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<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GKB3</accession>
    <name>A0A452GKB3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3 7">Carbonic anhydrase</fullName>
            <ecNumber evidence="3 7">4.2.1.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="7">Reversible hydration of carbon dioxide.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="11 12">
            <text>hydrogencarbonate + H(+) = CO2 + H2O</text>
            <dbReference type="Rhea" id="RHEA:10748"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:17544"/>
            <dbReference type="EC" id="4.2.1.1"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 7">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <comment type="similarity">
        <text evidence="2 7">Belongs to the alpha-carbonic anhydrase family.</text>
    </comment>
    <dbReference type="EC" id="4.2.1.1" evidence="3 7"/>
    <dbReference type="AlphaFoldDB" id="A0A452GKB3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002351.1">
        <property type="protein sequence ID" value="ENSGAGP00000002065.1"/>
        <property type="gene ID" value="ENSGAGG00000001445.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004089">
        <property type="term" value="F:carbonate dehydratase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.200.10:FF:000001">
        <property type="entry name" value="Carbonic anhydrase 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.200.10">
        <property type="entry name" value="Alpha carbonic anhydrase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001148">
        <property type="entry name" value="CA_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036398">
        <property type="entry name" value="CA_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023561">
        <property type="entry name" value="Carbonic_anhydrase_a-class"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018338">
        <property type="entry name" value="Carbonic_anhydrase_a-class_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18952">
        <property type="entry name" value="CARBONIC ANHYDRASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18952:SF25">
        <property type="entry name" value="CARBONIC ANHYDRASE 5B, MITOCHONDRIAL-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00194">
        <property type="entry name" value="Carb_anhydrase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01057">
        <property type="entry name" value="Carb_anhydrase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51069">
        <property type="entry name" value="Carbonic anhydrase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00162">
        <property type="entry name" value="ALPHA_CA_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51144">
        <property type="entry name" value="ALPHA_CA_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6 7" id="KW-0456">Lyase</keyword>
    <keyword evidence="4 7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 7" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="Alpha-carbonic anhydrase" evidence="8">
        <location>
            <begin position="8"/>
            <end position="266"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010718"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012925"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU367011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51144"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002065.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="287" mass="32571" checksum="7E9F97C0703243ED" modified="2019-05-08" version="1">AVRPGLCSQAGAAEGGAVHPLWQSELSVSGGTRQSPINIQWRDSVYDPHLKPLKIHYDPTTCLYIWNNGYSFQVEFDDSADKSVIMGGPLENHYRLKQFHFHWGAINEWGSEHMVDSKVYPAELHLVHWNPVTSTSFEEAIMENNGLVVIGVFLKLGAHHAGLQKLVDALPAVKHKDTIVEFDVFDPSCLMPSCPDYWTYDGSLTTPPLTESVTWIIKKTPIEVDQNQLEEFRMLLFTSAGEEEKRMVDNFRPLQPLMNRTVRSSFRNHHLLNIETRSSELAHQHSP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GKB4</accession>
    <name>A0A452GKB4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">CCHC-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GKB4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002484.1">
        <property type="protein sequence ID" value="ENSGAGP00000002177.1"/>
        <property type="gene ID" value="ENSGAGG00000001754.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003676">
        <property type="term" value="F:nucleic acid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035091">
        <property type="term" value="F:phosphatidylinositol binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd09558">
        <property type="entry name" value="SAM_ZCCH14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1520.10">
        <property type="entry name" value="Phox-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.50">
        <property type="entry name" value="Transcription Factor, Ets-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.60.10">
        <property type="entry name" value="Zinc finger, CCHC-type"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058599">
        <property type="entry name" value="PHAT_Smg/ZCCHC2-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036871">
        <property type="entry name" value="PX_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013761">
        <property type="entry name" value="SAM/pointed_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057327">
        <property type="entry name" value="Vts1_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037632">
        <property type="entry name" value="ZCCH14_SAM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042344">
        <property type="entry name" value="ZCCHC14"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001878">
        <property type="entry name" value="Znf_CCHC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036875">
        <property type="entry name" value="Znf_CCHC_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16195">
        <property type="entry name" value="ZINC FINGER CCHC DOMAIN CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16195:SF16">
        <property type="entry name" value="ZINC FINGER CCHC DOMAIN-CONTAINING PROTEIN 14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26034">
        <property type="entry name" value="PHAT_SMAUG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25479">
        <property type="entry name" value="Vts1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00098">
        <property type="entry name" value="zf-CCHC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00343">
        <property type="entry name" value="ZnF_C2HC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF64268">
        <property type="entry name" value="PX domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57756">
        <property type="entry name" value="Retrovirus zinc finger-like domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47769">
        <property type="entry name" value="SAM/Pointed domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50158">
        <property type="entry name" value="ZF_CCHC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0862">Zinc</keyword>
    <keyword evidence="1" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="CCHC-type" evidence="3">
        <location>
            <begin position="1029"/>
            <end position="1044"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="399"/>
            <end position="428"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="485"/>
            <end position="521"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="536"/>
            <end position="585"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="697"/>
            <end position="721"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="876"/>
            <end position="900"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="407"/>
            <end position="426"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="496"/>
            <end position="505"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="546"/>
            <end position="555"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="561"/>
            <end position="585"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="876"/>
            <end position="891"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00047"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS50158"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002177.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1070" mass="115863" checksum="E7966BF89F5E7035" modified="2019-05-08" version="1">MVEKRCPRLQRDSVYRWFSELPSPQRVEFLCGLLDLCIPLELRFLGACLEDLARKDYHSLRDSEIKANNPADLGSLTNLTDEVVRSKLLVSLALLGSAHREAAGVLFRTLTHIDSVINNYGLQLNEGRTGDEFLLLFTMASNHPAFSFHQKQVLRQELTQIQNIMASTSKNPSIPALGTMAAYPGCHKVAPRSETPINSMSNSLENALHTSAHSIEESLRKRSSGKHSKVSVEKVEVKGLAHKKNERNVDYSFEVLWSDSSVTLVTKSSTEVTEFISKLSQLYPEENLEKFIPCLAGPDSFYLERNHMDLESDIRYLASLPSHVLKNDHVKKFFSTSSSSQQLQSSSPGNPSLYKVVTTLGASGRPVCGVAGIQSSQNHVQHSAVSSVALPHCSHSGSTGSSLAYRSQMDTSSPMLSSSSQTPQTQEQNGILDWLRKLRLHKYYPVFKQLTMEKFLSLTEEDLNKFESLTMGAKKKLKTQLELEKEKSEKRCLNPGTPSSVTSSGVARVPPTTHGGPIQSIRGNHAAELRVDVDQPAHPLPREGSSSEYSSSSSSPMGIQTREESSDSAEENERRLEIHLDSSEKEKPVMLLNHFTSSSARPTAQVLPVQNDAGSSPSGHHILPMQMMTAASTHISPIRMLNAMHKSERNTDMKLLSSSVHSLLSLEERNKASPGPRSSIKMDKSFGNTVVDVMPTSSPHQPLQVLSGVPENSSSSSVNYGPRTKVVHASTLDRVIKTTQQTSLIAEASTAAPVTSSTVFHMARPPIKLLVSSSVPTDSATAGQTSCSSNMQISVSPAIINPRTALYTANTKVAFSAMSSMPVAPMPGSFCANSNTASSSSHPSTSFANMANLPSCPAPSSSPTISSVAENNFYSSSSSGGSAGSIPIPNQNHHHHHHHQQQPPGCMVCSSCGCSGNCGSGGMTVSYANYFQHPFSGSSMFTFPFLPFNPLCSNGYINTQQYNGSTTFPMVHSPYNSGLTPDSVMSGQTTFAVPPMQNFMAGTAGVYQAQGMMGNSNGSSHKKNGNLSCYNCGASGHRAQDCKQPPMDFNRQGTFRLKYAPPAESLDSTD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GKB5</accession>
    <name>A0A452GKB5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Succinate--CoA ligase [ADP-forming] subunit beta, mitochondrial</fullName>
            <ecNumber evidence="10">6.2.1.5</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="10">ATP-specific succinyl-CoA synthetase subunit beta</fullName>
            <shortName evidence="10">A-SCS</shortName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="10">Succinyl-CoA synthetase beta-A chain</fullName>
            <shortName evidence="10">SCS-betaA</shortName>
        </alternativeName>
    </protein>
    <gene>
        <name evidence="10" type="primary">SUCLA2</name>
    </gene>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">ATP-specific succinyl-CoA synthetase functions in the citric acid cycle (TCA), coupling the hydrolysis of succinyl-CoA to the synthesis of ATP and thus represents the only step of substrate-level phosphorylation in the TCA. The beta subunit provides nucleotide specificity of the enzyme and binds the substrate succinate, while the binding sites for coenzyme A and phosphate are found in the alpha subunit.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="16 10">
            <text>succinate + ATP + CoA = succinyl-CoA + ADP + phosphate</text>
            <dbReference type="Rhea" id="RHEA:17661"/>
            <dbReference type="ChEBI" id="CHEBI:30031"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57292"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="6.2.1.5"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="10">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
        <text evidence="10">Binds 1 Mg(2+) ion per subunit.</text>
    </comment>
    <comment type="pathway">
        <text evidence="1 10">Carbohydrate metabolism; tricarboxylic acid cycle; succinate from succinyl-CoA (ligase route): step 1/1.</text>
    </comment>
    <comment type="subunit">
        <text evidence="10">Heterodimer of an alpha and a beta subunit. The beta subunit determines specificity for ATP.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="10">Mitochondrion</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="10">Belongs to the succinate/malate CoA ligase beta subunit family. ATP-specific subunit beta subfamily.</text>
    </comment>
    <dbReference type="EC" id="6.2.1.5" evidence="10"/>
    <dbReference type="AlphaFoldDB" id="A0A452GKB5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002330.1">
        <property type="protein sequence ID" value="ENSGAGP00000002047.1"/>
        <property type="gene ID" value="ENSGAGG00000000169.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00223">
        <property type="reaction ID" value="UER00999"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
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        <property type="project" value="UniProtKB-SubCell"/>
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        <property type="term" value="C:succinate-CoA ligase complex"/>
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        <property type="term" value="F:ATP binding"/>
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    <dbReference type="GO" id="GO:0000287">
        <property type="term" value="F:magnesium ion binding"/>
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        <property type="term" value="F:succinate-CoA ligase (ADP-forming) activity"/>
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        <property type="term" value="P:tricarboxylic acid cycle"/>
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    <dbReference type="InterPro" id="IPR034723">
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        <property type="entry name" value="Succ_CoA_ligase-like_bsu"/>
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        <property type="entry name" value="Succinyl-CoA_synth-like"/>
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        <property type="entry name" value="PRK00696.1"/>
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    </dbReference>
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    </dbReference>
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        <property type="entry name" value="SUCCINATE--COA LIGASE [ADP-FORMING] SUBUNIT BETA, MITOCHONDRIAL"/>
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        <property type="entry name" value="SUCCINYL-COA SYNTHETASE BETA CHAIN"/>
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        <property type="entry name" value="ATP-grasp_2"/>
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    </dbReference>
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        <property type="entry name" value="Ligase_CoA"/>
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    </dbReference>
    <dbReference type="PIRSF" id="PIRSF001554">
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    </dbReference>
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        <property type="entry name" value="Glutathione synthetase ATP-binding domain-like"/>
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    </dbReference>
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        <property type="entry name" value="Succinyl-CoA synthetase domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50975">
        <property type="entry name" value="ATP_GRASP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01217">
        <property type="entry name" value="SUCCINYL_COA_LIG_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6 10 11" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="3 10 12" id="KW-0436">Ligase</keyword>
    <keyword evidence="7 10" id="KW-0460">Magnesium</keyword>
    <keyword evidence="4 10" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9 10" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="5 10 11" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0809">Transit peptide</keyword>
    <keyword evidence="2 10" id="KW-0816">Tricarboxylic acid cycle</keyword>
    <feature type="domain" description="ATP-grasp" evidence="13">
        <location>
            <begin position="3"/>
            <end position="228"/>
        </location>
    </feature>
    <feature type="binding site" evidence="10">
        <location>
            <position position="40"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="10">
        <location>
            <begin position="47"/>
            <end position="49"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
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        <location>
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        <ligand>
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        </ligand>
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        <location>
            <position position="214"/>
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        <ligand>
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        </ligand>
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    <feature type="binding site" evidence="10">
        <location>
            <position position="265"/>
        </location>
        <ligand>
            <name>substrate</name>
            <note>ligand shared with subunit alpha</note>
        </ligand>
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    <feature type="binding site" evidence="10">
        <location>
            <begin position="322"/>
            <end position="324"/>
        </location>
        <ligand>
            <name>substrate</name>
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    </feature>
    <feature type="site" description="Important for substrate specificity" evidence="10">
        <location>
            <position position="36"/>
        </location>
    </feature>
    <feature type="site" description="Important for substrate specificity" evidence="10">
        <location>
            <position position="104"/>
        </location>
    </feature>
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        <source>
            <dbReference type="ARBA" id="ARBA00005064"/>
        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
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    </evidence>
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    </evidence>
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        <source>
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        </source>
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        </source>
    </evidence>
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    </evidence>
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        <source>
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    </evidence>
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        <source>
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        </source>
    </evidence>
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        <source>
            <dbReference type="PROSITE" id="PS50975"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002047.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="405" mass="44102" checksum="E59FE83AD5352CFA" modified="2025-10-08" version="2">MSMELLQEAGISVPHGLVAKTPEEAYKIAKEIGSKDLVVKAQVLAGGRGKGTFEGGLKGGVKIVFSPEEAKEISSKMIGKKLFTKQTGEKGRICNQVFICERRYPRREYYFAITMERSFQGPVLIGSSHGGVNIEDVAAETPDAIIKEPIDIVEGIKKEQAVRLAQKMGFPPNLVDEAADNMIKIYNLFMKYDATMVEINPMVEDASGVVMCMDAKINFDSNAFYRQQKIFDLQDWTQEDERDKEAAKADLNYIGLDGNIGCLVNGAGLAMATMDIIKLHGGTPANFLDVGGGATVHQVTEAFKLITSDKKVLAILVNIFGGIMRCDVIAQGIIMAVKDLDLKIPIVVRLQGTRVDDAKALITASGLKILACDDLDEAAKMVVKLSEIVTLAKQAQVDVKFQLPI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GKC1</accession>
    <name>A0A452GKC1_9SAUR</name>
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        <submittedName>
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        </submittedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
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                <consortium name="Ensembl"/>
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        <scope>IDENTIFICATION</scope>
    </reference>
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        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
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        </citation>
        <scope>IDENTIFICATION</scope>
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    </dbReference>
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    <dbReference type="PANTHER" id="PTHR45920:SF2">
        <property type="entry name" value="FH1_FH2 DOMAIN-CONTAINING PROTEIN 1"/>
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    </dbReference>
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        <property type="entry name" value="FORMIN HOMOLOGY 2 DOMAIN CONTAINING, ISOFORM I"/>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
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        <property type="entry name" value="GBD_FH3"/>
        <property type="match status" value="1"/>
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    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
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        <location>
            <begin position="57"/>
            <end position="452"/>
        </location>
    </feature>
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        <location>
            <begin position="726"/>
            <end position="1120"/>
        </location>
    </feature>
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        <location>
            <begin position="328"/>
            <end position="405"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
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            <end position="530"/>
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    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="557"/>
            <end position="624"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="673"/>
            <end position="732"/>
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    </feature>
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            <end position="1122"/>
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            <end position="1146"/>
        </location>
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        <location>
            <begin position="1153"/>
            <end position="1243"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="362"/>
            <end position="372"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="488"/>
            <end position="499"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="575"/>
            <end position="587"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="600"/>
            <end position="624"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="2">
        <location>
            <begin position="699"/>
            <end position="718"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="2">
        <location>
            <begin position="1105"/>
            <end position="1119"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="1177"/>
            <end position="1198"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="2">
        <location>
            <begin position="1227"/>
            <end position="1242"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS51232"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS51444"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002154.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1264" mass="139456" checksum="5DDB084B1D78D33F" modified="2019-05-08" version="1">MEAGPGAALCRLQYLDDSDPFSCASFPEPRRAPVHPLLEALPLGAQLPALHRLLGAPLPLADCTLQVSPSGQYLDLDLSLLEQKDELEGFYEEISKGRRPTLILRTQLSVRVHAILEKLYNSQGPELRRSLFSLKQLFQEDKDLVPEFVNLEGLTCLIKVGAEADQNYQNYILRALSQVMLFVDGMLGVIAHNETVQWLYTLCGSLYRLVVKTALKLLLVFVEYTESNAQLLIQAVNTVEQARGAYPWSNLVAILEERNGADTELLVFAMTLINKTLAALPDQDSFYDVTDCLEQQGMEHIVQRHMHNKGLDPDVKQQFAIYESSLKHEDGVEELPPGARKERRKTDENRRGRRSLGGNVDAGAGASSPSPAVEDNKLPEPSSPPREPTAKTVMESSSAPPCFESVYNSTSNVRLSASASPEEREHPSLGERSVFKARFLENLASSQKERLSAMSKERLDILAEAIPGGGQLSPEPPPAGRLVLNEEKEQNETVNEQKPRLPSVGIALGRDENGDGTSGRCVPPQESATPCRSISLDKKLMHDMLFAKSQVEPERDLLFPEASASAQHHQVAGDQLSQAELQGSQDLENTRPKGQAPDRQVSSAQTRLPRTQSSVEAEANTQNVERALLTPFKKDLEFMWDRLEPSPMLLKIKDLDFSDLGEEEDSDALEQGAVGNDSFHVPPPLPPGTEALTVGGLLAPPPPPPPPXPVPGCPPPPALLGAPAPDGRSPAKKKKTVKLFWKELKHPSSQAVAGRFGRGTVWASLERVEIDAAKLEHLFESRAKEVLSSKKAADGKKCTVVLDPKRSNAINIGLTVLPPAHIIKTAILSFDEFAINKEGIEKILTMVPTDEEKQKIQEAQLANPDLPLGSAEQFLLTLSSISELTSRLQLWAFKLDYESMEQEIAEPLFDLKLGMEQLAKNQTFKCILATLLAMGNVLNGSQSRGFELSYLEKVSEVKDTVHRQSLLHHLCHAVVERFPQTSDLYSEIAAITRSAKVDFDELAESLSQLERRCRASWDNLKAIAKHETKPGLKSKLTDFLKESSQKIHILKVVHRRILHRFRSFLLFLGYPPAAARDMKVTGFCRLLREFALEYRTCRERVLQQQRKRATHRERNKTRGRMITEMERFSGMANSPEASPSPAVGLTFPEQQADAGHETMKSVLSSAVEPQSRRSRASRGVGRSSPSGSSVPQDDSSGSPADVSDEIMDRLVQSVTHSTGPRSGAAKERKRSRVSRKSLRRTLKSGLSDDLVQALGLAQAPGVQV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GKC5</accession>
    <name>A0A452GKC5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">Acyl-CoA dehydrogenase 6</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="9">
            <text>decanoyl-CoA + oxidized [electron-transfer flavoprotein] + H(+) = (2E)-decenoyl-CoA + reduced [electron-transfer flavoprotein]</text>
            <dbReference type="Rhea" id="RHEA:48176"/>
            <dbReference type="Rhea" id="RHEA-COMP:10685"/>
            <dbReference type="Rhea" id="RHEA-COMP:10686"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
            <dbReference type="ChEBI" id="CHEBI:58307"/>
            <dbReference type="ChEBI" id="CHEBI:61406"/>
            <dbReference type="ChEBI" id="CHEBI:61430"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="9">
            <dbReference type="Rhea" id="RHEA:48177"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="10">
            <text>dodecanoyl-CoA + oxidized [electron-transfer flavoprotein] + H(+) = (2E)-dodecenoyl-CoA + reduced [electron-transfer flavoprotein]</text>
            <dbReference type="Rhea" id="RHEA:47296"/>
            <dbReference type="Rhea" id="RHEA-COMP:10685"/>
            <dbReference type="Rhea" id="RHEA-COMP:10686"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57330"/>
            <dbReference type="ChEBI" id="CHEBI:57375"/>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
            <dbReference type="ChEBI" id="CHEBI:58307"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="10">
            <dbReference type="Rhea" id="RHEA:47297"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="14">
            <text>hexanoyl-CoA + oxidized [electron-transfer flavoprotein] + H(+) = (2E)-hexenoyl-CoA + reduced [electron-transfer flavoprotein]</text>
            <dbReference type="Rhea" id="RHEA:43464"/>
            <dbReference type="Rhea" id="RHEA-COMP:10685"/>
            <dbReference type="Rhea" id="RHEA-COMP:10686"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
            <dbReference type="ChEBI" id="CHEBI:58307"/>
            <dbReference type="ChEBI" id="CHEBI:62077"/>
            <dbReference type="ChEBI" id="CHEBI:62620"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="14">
            <dbReference type="Rhea" id="RHEA:43465"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="12">
            <text>octanoyl-CoA + oxidized [electron-transfer flavoprotein] + H(+) = (2E)-octenoyl-CoA + reduced [electron-transfer flavoprotein]</text>
            <dbReference type="Rhea" id="RHEA:48180"/>
            <dbReference type="Rhea" id="RHEA-COMP:10685"/>
            <dbReference type="Rhea" id="RHEA-COMP:10686"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57386"/>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
            <dbReference type="ChEBI" id="CHEBI:58307"/>
            <dbReference type="ChEBI" id="CHEBI:62242"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="12">
            <dbReference type="Rhea" id="RHEA:48181"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>oxidized [electron-transfer flavoprotein] + hexadecanoyl-CoA + H(+) = (2E)-hexadecenoyl-CoA + reduced [electron-transfer flavoprotein]</text>
            <dbReference type="Rhea" id="RHEA:43448"/>
            <dbReference type="Rhea" id="RHEA-COMP:10685"/>
            <dbReference type="Rhea" id="RHEA-COMP:10686"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57379"/>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
            <dbReference type="ChEBI" id="CHEBI:58307"/>
            <dbReference type="ChEBI" id="CHEBI:61526"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="11">
            <dbReference type="Rhea" id="RHEA:43449"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>tetradecanoyl-CoA + oxidized [electron-transfer flavoprotein] + H(+) = (2E)-tetradecenoyl-CoA + reduced [electron-transfer flavoprotein]</text>
            <dbReference type="Rhea" id="RHEA:47316"/>
            <dbReference type="Rhea" id="RHEA-COMP:10685"/>
            <dbReference type="Rhea" id="RHEA-COMP:10686"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57385"/>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
            <dbReference type="ChEBI" id="CHEBI:58307"/>
            <dbReference type="ChEBI" id="CHEBI:61405"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="13">
            <dbReference type="Rhea" id="RHEA:47317"/>
        </physiologicalReaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 15">
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </cofactor>
    </comment>
    <comment type="pathway">
        <text evidence="2">Lipid metabolism; mitochondrial fatty acid beta-oxidation.</text>
    </comment>
    <comment type="similarity">
        <text evidence="3 15">Belongs to the acyl-CoA dehydrogenase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GKC5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002499.1">
        <property type="protein sequence ID" value="ENSGAGP00000002190.1"/>
        <property type="gene ID" value="ENSGAGG00000001768.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003995">
        <property type="term" value="F:acyl-CoA dehydrogenase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050660">
        <property type="term" value="F:flavin adenine dinucleotide binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033539">
        <property type="term" value="P:fatty acid beta-oxidation using acyl-CoA dehydrogenase"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.110.10:FF:000002">
        <property type="entry name" value="Acyl-CoA dehydrogenase fadE12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.540.10">
        <property type="entry name" value="Acyl-CoA dehydrogenase/oxidase, N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.110.10">
        <property type="entry name" value="Butyryl-CoA Dehydrogenase, subunit A, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.140.10">
        <property type="entry name" value="Butyryl-CoA Dehydrogenase, subunit A, domain 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050741">
        <property type="entry name" value="Acyl-CoA_dehydrogenase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006089">
        <property type="entry name" value="Acyl-CoA_DH_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006091">
        <property type="entry name" value="Acyl-CoA_Oxase/DH_mid-dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046373">
        <property type="entry name" value="Acyl-CoA_Oxase/DH_mid-dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036250">
        <property type="entry name" value="AcylCo_DH-like_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009075">
        <property type="entry name" value="AcylCo_DH/oxidase_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013786">
        <property type="entry name" value="AcylCoA_DH/ox_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037069">
        <property type="entry name" value="AcylCoA_DH/ox_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009100">
        <property type="entry name" value="AcylCoA_DH/oxidase_NM_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48083:SF6">
        <property type="entry name" value="ACYL-COA DEHYDROGENASE 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48083">
        <property type="entry name" value="MEDIUM-CHAIN SPECIFIC ACYL-COA DEHYDROGENASE, MITOCHONDRIAL-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00441">
        <property type="entry name" value="Acyl-CoA_dh_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02770">
        <property type="entry name" value="Acyl-CoA_dh_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02771">
        <property type="entry name" value="Acyl-CoA_dh_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47203">
        <property type="entry name" value="Acyl-CoA dehydrogenase C-terminal domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56645">
        <property type="entry name" value="Acyl-CoA dehydrogenase NM domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00072">
        <property type="entry name" value="ACYL_COA_DH_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 15" id="KW-0274">FAD</keyword>
    <keyword evidence="6" id="KW-0276">Fatty acid metabolism</keyword>
    <keyword evidence="4 15" id="KW-0285">Flavoprotein</keyword>
    <keyword evidence="6" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="8 15" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0809">Transit peptide</keyword>
    <feature type="domain" description="Acyl-CoA dehydrogenase/oxidase N-terminal" evidence="18">
        <location>
            <begin position="52"/>
            <end position="164"/>
        </location>
    </feature>
    <feature type="domain" description="Acyl-CoA oxidase/dehydrogenase middle" evidence="17">
        <location>
            <begin position="168"/>
            <end position="263"/>
        </location>
    </feature>
    <feature type="domain" description="Acyl-CoA dehydrogenase/oxidase C-terminal" evidence="16">
        <location>
            <begin position="275"/>
            <end position="403"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001974"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005198"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00009347"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022630"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022827"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022832"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00047546"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00047893"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00047916"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00048877"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00049038"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00049192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="RuleBase" id="RU362125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF00441"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF02770"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF02771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002190.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="21">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="408" mass="44992" checksum="CD3AD6D40EB1015C" modified="2019-05-08" version="1">MGRCWPGLLTRGERGWWSSGLPGPARSQLRAASSSRAPGATGEGANLLYSGEHNALRASLRKIIDKEINPFVDKWEEEGQFPAHKVFKTLGQAGFLGVNKPIEYGGMGLDFSYSIAVAEELGNIHCGGVPMAIGVQTDMATPALTRFGSDELKRQFLVPTIAGEVVACLGVSEPGAGSDVASIKTTAVRKGDDYIINGGKMWITSGCQADWMCLLANSSEGPRYQNKSLICLPMNLPGVHIAKKIDKMGMRSSDTAQIFFEDVRVPSKYLIGEEGMGFTYQMLQFQEERLWAVANILAPMENVIQETIDYTRQRKIYNQPVLHNQTVHFRLAELATEVELVRSLFYQAVALYVRGNDVTKFASMAKLKEGRLARELSDNCLQFWGGMGFTSEVLVNRLYRTTASERQA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GKC7</accession>
    <name>A0A452GKC7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Cas scaffolding protein family member 4</fullName>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">Docking protein that plays a role in tyrosine kinase-based signaling related to cell adhesion and cell spreading. Regulates PTK2/FAK1 activity, focal adhesion integrity, and cell spreading.</text>
    </comment>
    <comment type="subunit">
        <text evidence="11">Interacts (via SH3 domain) with PTK2/FAK1 (via C-terminus).</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell junction</location>
            <location evidence="2">Focal adhesion</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the CAS family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GKC7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002466.1">
        <property type="protein sequence ID" value="ENSGAGP00000002160.1"/>
        <property type="gene ID" value="ENSGAGG00000001321.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005925">
        <property type="term" value="C:focal adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007169">
        <property type="term" value="P:cell surface receptor protein tyrosine kinase signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd12000">
        <property type="entry name" value="SH3_CASS4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.830:FF:000001">
        <property type="entry name" value="BCAR1 scaffold protein, Cas family member"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.40:FF:000147">
        <property type="entry name" value="Cas scaffold protein family member 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.230">
        <property type="entry name" value="Alpha-catenin/vinculin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.830">
        <property type="entry name" value="Serine-rich domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021901">
        <property type="entry name" value="CAS_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037362">
        <property type="entry name" value="CAS_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035744">
        <property type="entry name" value="CASS4_SH3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014928">
        <property type="entry name" value="Serine_rich_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038319">
        <property type="entry name" value="Serine_rich_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10654">
        <property type="entry name" value="CAS SCAFFOLDING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10654:SF19">
        <property type="entry name" value="CAS SCAFFOLDING PROTEIN FAMILY MEMBER 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12026">
        <property type="entry name" value="CAS_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08824">
        <property type="entry name" value="Serine_rich"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14604">
        <property type="entry name" value="SH3_9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00452">
        <property type="entry name" value="SH3DOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00326">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="8" id="KW-0965">Cell junction</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 13" id="KW-0728">SH3 domain</keyword>
    <keyword evidence="15" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="15">
        <location>
            <begin position="1"/>
            <end position="18"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019353470" description="Cas scaffolding protein family member 4" evidence="15">
        <location>
            <begin position="19"/>
            <end position="785"/>
        </location>
    </feature>
    <feature type="domain" description="SH3" evidence="16">
        <location>
            <begin position="28"/>
            <end position="90"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="270"/>
            <end position="294"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="358"/>
            <end position="386"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="400"/>
            <end position="420"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="270"/>
            <end position="286"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="365"/>
            <end position="376"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="402"/>
            <end position="418"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007848"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00055892"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00064074"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00071492"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002160.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="785" mass="88036" checksum="9B7017729092058E" modified="2019-05-08" version="1">VAVLAVLCKCGHILSCSAVTPSPALQLSSNTLAKALYDNKAECSDELAFRKGDILTVLEQNVLESEGWWKCSLHGRQGLAPANRLQLLAGSQAGMLPPSSEHDSQELHICQQNIYQVPSAPKSSTLSPTYERMDSWIKPVRSPVSSPPTQEIYQVPALAAQLFNSAIVLSDCTLLLHLELPPHHPCKTAKYYLVYDVPPTTARDHDISLQNLPQIKCLGHYNTLPNPRKSEWIYDIPVSPEKMGLKKDPSDSSLEKQVLYDIPPARYGTSLQNVPPTNIKSKSMNPQMYDVPPTQRKLTFPDISRYNVPSSRDMLLLQQNGNYHVPPSFLTPRVEQQNSEPNVYDIPKVLPTALQHRKGTEKNKSSFGNHTYNIPPQLSKDAKLEQDRLSVSSVDSRINTLSTSSSTSAESSFTSSSEEPAKEINLELDLAIETLTKLQHSVSSSVASLMIFVSSRWRFQEHLETNLEEIHRAIDHIKVSLGEFLDFAQVIKVNASCVTDNNLQTRIKTQLKILKDSFQILVETREALNNCKWSLEVLVIKKPQNNPDDLDRFVMVARAIPDDIKRFVSIIIANGKLLFKKNSKEQEIKSPKIGTDHKMAKPIPIPRRRELDSLQRNTLDKPNKNKVYSEKSRENVTEDCDYVQLQVSVLPPSKKHNIQSNLVLAKKITLSENCRLYFGALHKAISVFTSSLSNNQPPEIFIAQSKLIIMVGQKLVDSLCQETQEKDIRNDILCSSSHFCSLLKNLAVATKNAAVQYANAGAMQELQNQAKELSKYTQQFRAMME</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GKC8</accession>
    <name>A0A452GKC8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">EGF-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cell projection</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the tenascin family. Teneurin subfamily.</text>
    </comment>
    <comment type="caution">
        <text evidence="13">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000002350.1">
        <property type="protein sequence ID" value="ENSGAGP00000002064.1"/>
        <property type="gene ID" value="ENSGAGG00000000665.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043005">
        <property type="term" value="C:neuron projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050839">
        <property type="term" value="F:cell adhesion molecule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046982">
        <property type="term" value="F:protein heterodimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042803">
        <property type="term" value="F:protein homodimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007157">
        <property type="term" value="P:heterophilic cell-cell adhesion via plasma membrane cell adhesion molecules"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048666">
        <property type="term" value="P:neuron development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00054">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000021">
        <property type="entry name" value="Teneurin transmembrane protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.120.10.30:FF:000005">
        <property type="entry name" value="Teneurin transmembrane protein 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.120.10.30:FF:000006">
        <property type="entry name" value="Teneurin transmembrane protein 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.180.10.10:FF:000001">
        <property type="entry name" value="Teneurin transmembrane protein 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.180.10.10:FF:000005">
        <property type="entry name" value="Teneurin transmembrane protein 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.180.10.10">
        <property type="entry name" value="RHS repeat-associated core"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.120.10.30">
        <property type="entry name" value="TolB, C-terminal domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011042">
        <property type="entry name" value="6-blade_b-propeller_TolB-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008969">
        <property type="entry name" value="CarboxyPept-like_regulatory"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057627">
        <property type="entry name" value="FN-plug_TEN1-4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022385">
        <property type="entry name" value="Rhs_assc_core"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056823">
        <property type="entry name" value="TEN-like_YD-shell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056822">
        <property type="entry name" value="TEN_NHL"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056820">
        <property type="entry name" value="TEN_TTR-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051216">
        <property type="entry name" value="Teneurin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028916">
        <property type="entry name" value="Tox-GHH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006530">
        <property type="entry name" value="YD"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR03696">
        <property type="entry name" value="Rhs_assc_core"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01643">
        <property type="entry name" value="YD_repeat_2x"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11219">
        <property type="entry name" value="TENEURIN AND N-ACETYLGLUCOSAMINE-1-PHOSPHODIESTER ALPHA-N-ACETYLGLUCOSAMINIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11219:SF9">
        <property type="entry name" value="TENEURIN-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25024">
        <property type="entry name" value="EGF_TEN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24329">
        <property type="entry name" value="FN-plug_TEN1-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25021">
        <property type="entry name" value="TEN_NHL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25023">
        <property type="entry name" value="TEN_YD-shell"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23538">
        <property type="entry name" value="Teneurin_ABD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15636">
        <property type="entry name" value="Tox-GHH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25020">
        <property type="entry name" value="TTR_TEN1-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF63829">
        <property type="entry name" value="Calcium-dependent phosphotriesterase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49464">
        <property type="entry name" value="Carboxypeptidase regulatory domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01186">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="12" id="KW-0966">Cell projection</keyword>
    <keyword evidence="11 13" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6 13" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="10" id="KW-0472">Membrane</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9" id="KW-1133">Transmembrane helix</keyword>
    <feature type="domain" description="EGF-like" evidence="14">
        <location>
            <begin position="136"/>
            <end position="171"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="13">
        <location>
            <begin position="140"/>
            <end position="150"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="13">
        <location>
            <begin position="161"/>
            <end position="170"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004167"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004316"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00009385"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002064.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2100" mass="235237" checksum="61D38B8FF3B59E52" modified="2019-05-08" version="1">PIEVWSQCSGGGAVHGAPCRPPSNHHHHHLGAGPAGHFPGAAQCQERLLTARSAVLTGAARVPFQPGVLVENQAPEICAADCGGHGICVGGTCRCEEGWMGAACDQRACHPRCNEHGTCRDGKCECSPGWNGEHCTIEGCPGLCNGNGRCTLDMNGWHCVCQLGWRGAGCDTSMETACGDGKDNDGDGLVDCMDPDCCLQPLCHVNPLCLGSPDPLDIIQETQTPTSQQNLHSFYDRIKFLIGKDSTHVIPGENPFDGGHACVIRGQVITSDGTPLVGVNISFVNNPLFGYTISRQDGSFDLVTNGGISIILHFERAPFITQEHTLWLPWDRFFVMETIVMRHEENEIPSCDLSNFARPNPVVSPSPLTAFASSCSEKGPIVPEIQALQEEINIAGSKIKLSYLSSRTAGYKSVLRISMTHPTIPFNLMKVHLMVAVEGRLFRKWFAAAPDLSYYFIWDKTDVYSQKVYGLSEAFVSVGYEYESCPDLILWEKRTAVLQGYEIDASKLGGWSLDQHHALNIQSGILHKGNGENQFVSQQPPVIGSIMGNGRRRSISCPSCNGLADGNKLLAPVALTCGSDGSLYVGDFNYIRRIFPSGNVTNILELRNKDFRHSHSPAHKYYLTSDPITGSIYLSDTNSRRIYKIKSTSVVKDVVKNSEVVAGTGDQCLPFDETRCGDGGKGTEATLTNPRGKTQSLLTSILKWASRKVPNQQFWNTTEQVYKLPQGNPLMCLRVRLEWPTDLAVSPMDNSLYVLDNNVVLQISENHQVRIVAGRPMHCQVPGIDHFLLSKVAIHATLESATALAVSHNGVLYIAETDEKKINRIRQVTTNGEISLVAGAPSSCDCKNDANCDCFSGDDGYAKDARLNAPSSLAVCADGELYVADLGNIRIRFIRKNKPYLNTQNLYELSSPIDQELYLFDTSGKHLYTQSLPTGDYLYNFTYTGDGDITLITDNNGNLVNVRRDSTGMPLWLVVPDGQVYWVTIGTNSALKSVTAQGHEVGMMTYHGNSGLLATKTNENGWTTFYEYDSFGRLTNVTFPTGQVSSFRSDTDRSVHVQVETSSKDDVTITTNLSASGAFYTLLQDQVRNSYYIGADGSLRLMLANGMEVALQTEPHLLAGTVNPTVGKRNVTLPIDNGLNLVEWRQRKEQARGQVTVFGRRLRVHNRNLLSLDFDRVTRTEKIYDDHRKFTLRILYDQAGRPSLWSPSSRLNGVNVTYSPGGHIAGIQRGTMSERMEYDQAGRIISRIFADGKSWSYTYLEKSMVLLLHSQRQYIFEFDKNDRLSSVTMPNVARQTLETIRSIGYYRNIYRPPEGNASVIQDFTEEGQLLHTFYLGTGRRVIYKYGKLSKLAEMLYDTTKIGFTYDETAGMLKTINLQNEGFTCTIRYRQIGPLIDRQIFRFTEEGMVNARFDYNYDNSFRVTSMQAVINETPLPIDLYRYDDVSGKTEQFGKFGVIYYDINQIITTAVMTHTKHFDAYGRMKEVQYEIFRSLMYWMTVQYDNMGRVVKKELKVGPYANTTRYSYEYDADGQLQTVSINDKPLWRYSYDLNGNLHLLSPGNSARLTPLRYDLRDRITRLGDVQYKMDEDGFLRQRGNDIFEYNSAGLLIKAYNKVSGWSVKYRYDGLGRRVSSKTAHSHHLQFFYADLTNPTKVTHLYNHSSSEITSLYYDLQGHLFAMELSSGDEFYIACDNIGTPLAVFSGTGLMIKQILYTAYGEIYMDTNPNFQVIIGYHGGLYDPLTKLIHMGRRDYDVLAGRWTSPDHDMWKHLSSNNIMPFNLYMFKNNNPISNAQDIKCYMTDVNSWLLTFGFQLHNVIPGYPKPDTDAMEPSYELIHTQMKTQEWDNSKSILGVQCEVQKQLKAFVTLERFEQIYSSSIAGCRTVKENKNFASGGSVFGKGVKFAMKDGRVATDIISLANEDGRRIAAILNNAHYLENLHFTIEGVDTHYFVKQGPSESDLSILGLSGGRRTLENGVNVTVSQINTVLSGRTRRYTDIQLQYGALCLNTRYGTTLDEEKARVLELARQRAIAQAWAREQQRLRDGEEGIRSWTEGEKQQVLNTGRVQGYDGYFVISVEQYPELSDSANNIHFMRQSEMGRR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GKE4</accession>
    <name>A0A452GKE4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4 7">Methanethiol oxidase</fullName>
            <shortName evidence="7">MTO</shortName>
            <ecNumber evidence="3 7">1.8.3.4</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="7">Selenium-binding protein 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="7">Catalyzes the oxidation of methanethiol, an organosulfur compound known to be produced in substantial amounts by gut bacteria. Selenium-binding protein which may be involved in the sensing of reactive xenobiotics in the cytoplasm. May be involved in intra-Golgi protein transport.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="6 7">
            <text>methanethiol + O2 + H2O = hydrogen sulfide + formaldehyde + H2O2 + H(+)</text>
            <dbReference type="Rhea" id="RHEA:11812"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:15379"/>
            <dbReference type="ChEBI" id="CHEBI:16007"/>
            <dbReference type="ChEBI" id="CHEBI:16240"/>
            <dbReference type="ChEBI" id="CHEBI:16842"/>
            <dbReference type="ChEBI" id="CHEBI:29919"/>
            <dbReference type="EC" id="1.8.3.4"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="1 7">Organosulfur degradation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="7">Nucleus</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="7">Cytoplasm</location>
            <location evidence="7">Cytosol</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="7">Membrane</location>
            <topology evidence="7">Peripheral membrane protein</topology>
        </subcellularLocation>
        <text evidence="7">May associate with Golgi membrane. May associate with the membrane of autophagosomes.</text>
    </comment>
    <comment type="similarity">
        <text evidence="2 7">Belongs to the selenium-binding protein family.</text>
    </comment>
    <dbReference type="EC" id="1.8.3.4" evidence="3 7"/>
    <dbReference type="AlphaFoldDB" id="A0A452GKE4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002532.1">
        <property type="protein sequence ID" value="ENSGAGP00000002215.1"/>
        <property type="gene ID" value="ENSGAGG00000001587.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0018549">
        <property type="term" value="F:methanethiol oxidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008430">
        <property type="term" value="F:selenium binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008826">
        <property type="entry name" value="Se-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23300">
        <property type="entry name" value="METHANETHIOL OXIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23300:SF0">
        <property type="entry name" value="METHANETHIOL OXIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05694">
        <property type="entry name" value="SBP56"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF75011">
        <property type="entry name" value="3-carboxy-cis,cis-mucoante lactonizing enzyme"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0007">Acetylation</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="7" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="7" id="KW-0653">Protein transport</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 7" id="KW-0711">Selenium</keyword>
    <keyword evidence="7" id="KW-0813">Transport</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00005177"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005606"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012510"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00015601"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023266"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00047539"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU369071"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002215.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="466" mass="52257" checksum="440964EE32B504A3" modified="2019-05-08" version="1">SSAKCGACGPGYKTPLDAMKGPREEIVYLPCIYRNTGIEKPDYLATVDVDPKSSHYCQVIHRLPMPNLRDELHHSGWNTCSSSFGDVTKKRNRLLLPSLISSRIYVVDVGTDMRAPRIHKMIEPVDVFWKCGLANPHTTHCLGNGEIMISSMGDPSGNGKGGFILLDGETFEVKGNWEKGSNLPALGYDFWYQPRHNVLMSTEWGIPKVRIGNPKRSYGRHINVWDWTTHVFTQSIDLGEDSIPLEIRFLHNPDAAEGYVGCALSSAIHRFYKTGNGTWAAEKVVQVPSKKVNGWLFPDMPGFITDILISLDDRFLYFSNWLHGDIRQYDISNTRKPRLVGQVFLGGKIPRDGPIVVVEDQELKCQPEPFVIKGKRVQGGPQMIQLSLDGKRLYVSTSLYSAWDKQFYPDLVKEGSVMLQIDVDTEKGGLKVNKNFLVDFGKEPDGPVLAHEIRYPGGDCSSDIWI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GKE6</accession>
    <name>A0A452GKE6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Centrosomal protein of 164 kDa</fullName>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Plays a role in microtubule organization and/or maintenance for the formation of primary cilia (PC), a microtubule-based structure that protrudes from the surface of epithelial cells. Plays a critical role in G2/M checkpoint and nuclear divisions. A key player in the DNA damage-activated ATR/ATM signaling cascade since it is required for the proper phosphorylation of H2AX, RPA, CHEK2 and CHEK1. Plays a critical role in chromosome segregation, acting as a mediator required for the maintenance of genomic stability through modulation of MDC1, RPA and CHEK1.</text>
    </comment>
    <comment type="subunit">
        <text evidence="15">Interacts (via N-terminus) with ATRIP. Interacts with ATM, ATR and MDC1. Interacts with XPA (via N-terminus) upon UV irradiation. Interacts with CEP83, CCDC92, TTBK2, DVL3, NPHP3 and weakly with NPHP4. Interacts with DZIP1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Microtubule organizing center</location>
            <location evidence="1">Centrosome</location>
            <location evidence="1">Centriole</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000002386.1">
        <property type="protein sequence ID" value="ENSGAGP00000002098.1"/>
        <property type="gene ID" value="ENSGAGG00000001670.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005814">
        <property type="term" value="C:centriole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097539">
        <property type="term" value="C:ciliary transition fiber"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060271">
        <property type="term" value="P:cilium assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006281">
        <property type="term" value="P:DNA repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00201">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1470.10:FF:000001">
        <property type="entry name" value="Centrosomal protein of 164 kDa"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1470.10">
        <property type="entry name" value="Photosystem I PsaD, reaction center subunit II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051841">
        <property type="entry name" value="MT-Golgi_org_protein"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001202">
        <property type="entry name" value="WW_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036020">
        <property type="entry name" value="WW_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18902:SF27">
        <property type="entry name" value="CENTROSOMAL PROTEIN OF 164 KDA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18902">
        <property type="entry name" value="NUCLEAR MITOTIC APPARATUS PROTEIN 1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00397">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00456">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51045">
        <property type="entry name" value="WW domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01159">
        <property type="entry name" value="WW_DOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50020">
        <property type="entry name" value="WW_DOMAIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="13" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="5" id="KW-0132">Cell division</keyword>
    <keyword evidence="8" id="KW-0970">Cilium biogenesis/degradation</keyword>
    <keyword evidence="9 17" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="6" id="KW-0227">DNA damage</keyword>
    <keyword evidence="10" id="KW-0234">DNA repair</keyword>
    <keyword evidence="7" id="KW-0498">Mitosis</keyword>
    <keyword evidence="12" id="KW-0539">Nucleus</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="WW" evidence="19">
        <location>
            <begin position="58"/>
            <end position="91"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="105"/>
            <end position="151"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="163"/>
            <end position="215"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="237"/>
            <end position="263"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="315"/>
            <end position="372"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="434"/>
            <end position="467"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="505"/>
            <end position="537"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="572"/>
            <end position="604"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="630"/>
            <end position="660"/>
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    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="845"/>
            <end position="896"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="972"/>
            <end position="992"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1041"/>
            <end position="1077"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1093"/>
            <end position="1125"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1682"/>
            <end position="1702"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="17">
        <location>
            <begin position="1139"/>
            <end position="1627"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="18">
        <location>
            <begin position="110"/>
            <end position="126"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="127"/>
            <end position="138"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
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            <begin position="169"/>
            <end position="194"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="241"/>
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    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
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            <begin position="315"/>
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        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="337"/>
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    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="441"/>
            <end position="450"/>
        </location>
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        <location>
            <begin position="580"/>
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        </location>
    </feature>
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        <location>
            <begin position="633"/>
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        </location>
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    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="1044"/>
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        </location>
    </feature>
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        <location>
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            <begin position="1102"/>
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        <location>
            <begin position="1687"/>
            <end position="1702"/>
        </location>
    </feature>
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            <dbReference type="ARBA" id="ARBA00004114"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
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            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022776"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022794"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00056906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00061715"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00067900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50020"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002098.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1991" mass="224165" checksum="A6B365431E687BB8" modified="2019-05-08" version="1">MDMAGPAIRIGDQLILEEDYDETYIPSEQEIQEFAREIGIDPGNEPELMWLAREGIVAPLPPEWKPCQDITGDIYYFNFSNGQSTWDHPCDEHYRELVIQEREKLSAHGSIKKKDKKKKKDKKEKKEKKEREPVKHAAEMQPEPGILPSSSFYRMSSPILSSECASPDLDQHSQMRIEGFLKKGKRKPSERPPETSDSNKQFAGPSPNKLQPLFTAKSCRTHQILADVEKILGRMPSFSRSDVDHQQGQDRSPKIGDKPCLGFSDSESEGLDVVRTKPISYVARKSCSQDLENGRSVLDSCEVLEDKGMLFDEEKLSDNWKQEEMTGGSVCQPDDGLSVRERSTGPGGDKERGPGFSQADENEAFLTPSAPDHTFYSRKCEMILFDASPLEDLSLMEEGDVHCPQRELRAHSRAVESRRKRKLQDQALSQATKFCKSSEVVGKHERESSKQAEFVESEGAGLQAAPQKDLLDSHLAVPEVTADTLPYTRGVEAYAHAGADISASEPVSLSAAQHKDKEKANGLGPEPDNSLTSNSADHFASQILGEIDNFSWDLQSSRESDRPTDLLAAAAKGSLPTHPFTGTLQTPPRSSANGKSQSDCNSEDEKFYQHVQENDQVPSQEHSCVSFNMKLNRSRDAPEKNNEGRQEETGEGEQKTEPVPRGLEVDVFTEGLQVSPGWLYQTAECQESSRLCQQVEGELTQSGKVTEDYCKLKLCEGAGSNESPGSLLAPIQVPPGSLAPLRGVMDGPASAFHGLLSTNVEISGVPSPLLREGTLPLQTLKSTGCTKSLLGSMHEEKASLNLLALDEEEEEESENQSPRGTARLLKNLHMDISALGGGFEYEESLKMSHAEESQTFPLDSDGVRPPAPDKLLNRDADSSLCGLNKEESSGKPLGNELLEKEDTDVEDTVSVAYEQSGTLGERTAEGTHVVSSQGHPVAASPDVRAAETDQMESVAVTVDAVSVSEKIVNGMREEAADLETDSKLDAGKESEASEHVKELQVSDCSDHELLHLTDFGFQSRISEQVLDMDILSAALDSPKVEAQGLREEEKDQSEASLEEEQSKRPKAAESERNQRPFSACKMAEEKCLSLENASQEETAALEQKEGSLDSPKEEEPAAPHQAQLGEEIKQVVSLNWPSAESLNEIARKQEQELEQEKERLLQAKEERVRRFQEELRQEEEEEAQVLHQQKEKSLWSLKEELAKASEEEELRMREEEVEKLSKLRSQISSETKTEKEKIRAEQEAALQKLREEWESLQKVERERLEERKKRVLEMLKVEVEEAQQREAAELEQEKQRVLNELKERLERKRKEAAEVLEKQFAAELQELKSTAEEKHQKVVSSLRKQIAEAQRSEEAQLREELERAEQKVQQKMYQVAEFERELSELMKEKRQEVERDHDRKMERMKEEHKEVLARIQEQYEEEERKQRTKKLEELKGELERLRQLHDGELKALQKQLDEQLSDLRQSHKVKERKLQELEMELEIQATDAKARCAQLNHQEEAMRKKRQQLLDEEKQTELQRNEAALAAQLRLEESRKEHADLVESIRQLRRTLEELQDQKAELESQVDLLQTRSQRLQRRISELEAAVQSKQETLKELVAEDSMASPRREAELHLEDLRETLQAHSSREPASLTSHSNEDSDLQLDNVRHFISAEGVSIKNAKEFLVRQTRSMRKRHTALKAAKQQWRHNMQQAQDAVQDPDSSQLLEDVRRNLEEESAQLDKMKSAMQKGQVLLKKKEEKLSQLESSLLEELSDEDTLKGAACKKVVTFDLSDSEETHSLSSTDVPQHKFDLKPDVQFPQLDKIQHLTSTLQHITAELNGVLSTLGSLNQRQPPLFAPVQVPMTALPSDGVPLSMYTSLARAQAAGPIMPPAGIPLVNQWAWSTGLNSSPSSTAGQSVDDILAEKWHKYFPGGGSPLWSPTMLATLMTALFFMVRALGQKGGSSRHDSIYSRKGNVVPDSSSKCLCDPTERAAVTIQTHYRKYQQHKQEKK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GKF3</accession>
    <name>A0A452GKF3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">ubiquitinyl hydrolase 1</fullName>
            <ecNumber evidence="3">3.4.19.12</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>Thiol-dependent hydrolysis of ester, thioester, amide, peptide and isopeptide bonds formed by the C-terminal Gly of ubiquitin (a 76-residue protein attached to proteins as an intracellular targeting signal).</text>
            <dbReference type="EC" id="3.4.19.12"/>
        </reaction>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the peptidase C19 family.</text>
    </comment>
    <dbReference type="EC" id="3.4.19.12" evidence="3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002502.1">
        <property type="protein sequence ID" value="ENSGAGP00000002193.1"/>
        <property type="gene ID" value="ENSGAGG00000001383.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004843">
        <property type="term" value="F:cysteine-type deubiquitinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016579">
        <property type="term" value="P:protein deubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd02659">
        <property type="entry name" value="peptidase_C19C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.70.10">
        <property type="entry name" value="Cysteine proteinases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056850">
        <property type="entry name" value="ARM_UBP34_24_USP9X_Y"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021905">
        <property type="entry name" value="DUF3517"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038765">
        <property type="entry name" value="Papain-like_cys_pep_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050164">
        <property type="entry name" value="Peptidase_C19"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001394">
        <property type="entry name" value="Peptidase_C19_UCH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055176">
        <property type="entry name" value="UBP24/USP9X/USP9Y_UBL"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028889">
        <property type="entry name" value="USP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018200">
        <property type="entry name" value="USP_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24006">
        <property type="entry name" value="UBIQUITIN CARBOXYL-TERMINAL HYDROLASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24006:SF925">
        <property type="entry name" value="UBIQUITINYL HYDROLASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25010">
        <property type="entry name" value="ARM_UBP24_USP9X-Y"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12030">
        <property type="entry name" value="DUF3517"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00443">
        <property type="entry name" value="UCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22900">
        <property type="entry name" value="UCH_UBL1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54001">
        <property type="entry name" value="Cysteine proteinases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00972">
        <property type="entry name" value="USP_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00973">
        <property type="entry name" value="USP_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50235">
        <property type="entry name" value="USP_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="5" id="KW-0645">Protease</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0788">Thiol protease</keyword>
    <keyword evidence="6" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="USP" evidence="10">
        <location>
            <begin position="1381"/>
            <end position="1779"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="790"/>
            <end position="823"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="1419"/>
            <end position="1455"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="2268"/>
            <end position="2346"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="793"/>
            <end position="813"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="1425"/>
            <end position="1455"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="9">
        <location>
            <begin position="2268"/>
            <end position="2277"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="2295"/>
            <end position="2306"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="2320"/>
            <end position="2330"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000707"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009085"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012759"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022807"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50235"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002193.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2346" mass="267327" checksum="C27FD892D19C9F1F" modified="2019-05-08" version="1">MYLEGEVSVLEGAMHISLSHGVCVCLPCCLPSLHSCCLIECAGWLVDLINKFGTLNGFQILHDRFMSGSALNVQIIAALIKPFGQCYEFLTLHTVKKYFLPIIEMVPQFLENLTDDELKKEAKNEAKNDALSMIIKSLKNLASRVPGQEETVKNLEIFRLKMILRLLQISSFNGKMNALNEVNKVISSVSYYTHRHGNPEEEEWLTAERMAEWIQQNNILSIVLRDSLHQPQYVEKLEKILRFVIKEKALTLQDLDNIWAAQAGKHEAIVKNVHDLLAKLAWDFSPEQLDHLFDCFKASWTNASKKQREKLLELIRRLAEDDKDGVMAHKVLNLLWNLAHSDDVPVDIMDQALSAHIKILDYSCSQDRDTQKIQWIDRFIEELRTNDKWVIPALKQIREICSLFGEAPQNLSQTQRSPHVFYRHDLINQLQHNHALVTLVAENLSAYMESMRQYAKEHGEYDPQTVRPGSRYSHVQEVQERLNFLRFLLKDGQLWLCAPQAKQIWKCLAENAVYLCDREACFKWYSKLMGDEPDLDPDINKDFFENNVLQLDPSLLTENGMKCFERFFKAVNCREGKLVAKRRAYMMDDLELIGLDYLWRVVIQGNDDIASRAIDLLKEIYTNLGPRLQVNQVVIHEDFIQSCFDRLKASYDTLCVLDGDKDSINCARQEAIRMVRVLTVLREYINECDSDYHEERTILPMSRAFRGKHITLVVRFPNQGRQVEDLDIWSHTNDTIGSVRRCILNRIKANSAHTKVELFIGGELVDPADDRKLIGQLNLKDKTLITAKLTQISSNMPSSPDSSSDSSTGSPGNHGNHYSDGPNPEVESCLPGVIMSLHPRYISFLWQVADLGSSLNMPPLRDGARVLMKLMPPDNTTVEKLRAICLDHAKLGESSLSPSLDSLFFGPSASQVLYLTEVVYALLMPASAPLGEDASDFQYNFLKSGGLPLVLSMLTRNNFLPNADMETRRGAYLNALKIAKLLLTAIGFGHVRAVAEACQPVVEGTSPISPINQATHDQAVVLQTALQNIPNPTSECMLRNVAIRLAQQISDEASKYIPDICVIRAVQKIVWASGCGSVQLVFSSNEEISKIYEKTNAGNEPDMEDEQVCCEALEVMTLCFALIPTALDALSKEKAWQTFIIDLLLHCHSKTVRQMAQEQFFLMATRCCMGQRPLLFFITLLFTVLGSTARERAKHSGDYFTLLRHLLNYAYNSNINVPNAEVLLNNEIDWLKRIRDEVKRTGETGVEETILEGHLGVTKELLAFQTPEKKYHIGCEKGGANLIKELIDDFIFPASNVYLQYMRNGELPAEQAIPVCSSPATINAGFELLVALAVGCVRNLKQIVDTLTEMYYIGTAITTCEALTEWEYLPPVGPRPSKGFVGLKNAGATCYMNSVIQQLYMIPAIRNGILAIEGTGSDVDDDMSGDEKQDNESNVDPRDDVFGYPHQYEDKPTLSKTEDRKEYNIGVLRHLQVIFGHLAASRLQYYVPRGFWKQFRLWGEPVNLREQHDALEFFNSLVDSLDEALKALGHPAMLSKVLGGSFADQKICQGCPHRYECEESFTTLNVDIRNHQNLLDSLEQYVKGDLLEGANAYHCEKCNKKVDTVKRLLIKKLPPVLAIQLKRFDYDWERECAIKFNDYFEFPRELDMEPYTVAGVAKLEGDDVNPENQLIQNEQPENEHSGSTKYRLVGVLVHSGQASGGHYYSYIIQRNGGDGEKNRWYKFDDGDVTECKMDDDEEMKNQCFGGEYMGEVFDHMMKRMSYRRQKRWWNAYILFYERMDTMDKDNELIKYISELAMTTKPHQIKMPSAIERSVRKQNVQFMHNRMQYSLEYFQFIKKLLTCNSIFLNPPPGYLCCQEIFFDHCLEFLSPSKSYKNVRFWFAHNVLFNVSNRFSEYLLECPSAEVRGAFAKLIVFIAHFSLQDGPCPSPFASPGPSSQAYDNLSLSDHLLRAVLNLLRREVSEHGRHLQQYFNLFVMYANLGVAEKTQLLKLSVPATFMLVALDEGPGPPIKYQYAELGKLYTVVSQLIRCCNVSSRMQSSINGNPPLSNPYGDPNLSQPIMALQQNVADILFVRTSYVKKIIEDCSNSEETIKLLRFCCWENPQFSSTVLSELLWQVAYSYTYELRPYLDLLLQILLIEDSWQTHRIHNALKGIPDDRDGLFDTIQRSKNHYQKRAYQCIKCMVALFSNCPVAYQILQSNGDLKRKWTWAVEWLGDELERRPYTGNPQYTYNNWSPPVQSNETSNGYFLERSHSARMTLAKACELCPEEEPDDPDAPDEHESSPPEDAPLYPHSPSSQYQQNNHVHGQPYTGPAAHHMNNPQRTGQRAQENYEGSEEVSPPQTKD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GKF8</accession>
    <name>A0A452GKF8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">ARID domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GKF8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002564.1">
        <property type="protein sequence ID" value="ENSGAGP00000002240.1"/>
        <property type="gene ID" value="ENSGAGG00000001821.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000976">
        <property type="term" value="F:transcription cis-regulatory region binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.490">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.60">
        <property type="entry name" value="ARID DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051232">
        <property type="entry name" value="ARID/SWI1_ChromRemod"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001606">
        <property type="entry name" value="ARID_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036431">
        <property type="entry name" value="ARID_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043151">
        <property type="entry name" value="BAH_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13964:SF37">
        <property type="entry name" value="AT-RICH INTERACTIVE DOMAIN-CONTAINING PROTEIN 5B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13964">
        <property type="entry name" value="RBP-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01014">
        <property type="entry name" value="ARID"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00501">
        <property type="entry name" value="BRIGHT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46774">
        <property type="entry name" value="ARID-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51011">
        <property type="entry name" value="ARID"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0539">Nucleus</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0804">Transcription</keyword>
    <keyword evidence="1" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="ARID" evidence="5">
        <location>
            <begin position="232"/>
            <end position="325"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="248"/>
            <end position="282"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="325"/>
            <end position="503"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="799"/>
            <end position="860"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="875"/>
            <end position="894"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="953"/>
            <end position="977"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="4">
        <location>
            <begin position="250"/>
            <end position="259"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="359"/>
            <end position="373"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="385"/>
            <end position="411"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="481"/>
            <end position="491"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="829"/>
            <end position="860"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="953"/>
            <end position="969"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS51011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002240.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1101" mass="123300" checksum="592A55922E2F80B8" modified="2019-05-08" version="1">MEPNSVQWVGSPCGLHGPYIFYKAFQFHLEGRPRILSLGDFFFVRCKPEDPICIAELQLLWEERTTRQLLSSSKLYFLPEDTPQGRNSDHGEDEVIAVSEKVTVKLEDLAKWAHSDFSKWKCGLRADPVIPMELGKNGQKEALMRYRQSTLNSGLNFKDILKEKADLGEDDEDSNLLILSYPQYCRYRSMLKRVQDKPSSILTDQFILALGGIAVISKNPQILYCRDTFDHPTLIENESVCDEFAPNLKGRPRKKKPCPQRRDSLNGVKDSNNNSESKAVAKITARRQWKHIYDELGGNPGSTSAATCTRRHYERLILPYERFIKGEEDKPLPPVKPRKQDNSSQESETKTKVSGTKRIKNENQKSKKEKDNAQKPQDASEVSSEQEKDQDSSDQKSLSEHPAVEETKKPIEGPQSLLPEATGLVSLEKIDLMESSSSSEKPKEEVQHSVSFSSASVPPEEDVILEPTVNKHLHTPTDALDDTKQEQRDHSFTGSLESEPQELPFNNFPAIQVQLPSQNDMEDDKLPEMADYIANCTVKLDQLGNEDIHNALKQTPKVLVVQNFDMFKEKELPGSINDDSSFSYTPLIYSKGNPGIMSPLAKKKLLSQVSGATLSCSYAYGSPPPLISKKKLNSRDELSASISQAPYAPNSDSVAVNRPSVIQHVQSFKTKNSEERKSINEAFKPDTLSKPDPKRCDFSKHHLNSLAESYVLKPDIQDCKDKMNEKRALQHSHVPSFLADFYSSPHLHSLYRHTEHHLNNEQTSKYLPRDMFRDSENISTFTQHKHQEKLNLNYHPSLHQQEKKTAVEASSDDQPTDLSLPKSVHKQTAKTPGSSLPHSSMVPQESKSISQFHTSGSQGVSIDCNPKACRVSPMTMSAPKKHGESLHRSGKQQNQRLENLRKMEGMVHPIISRKSSPQNIGAARPLKRNLDGLDKVISEKKIRAVSPLHLPKEVPAKEKVPDREGEGSKSVHGLHSGNTLESHKFPLSTPIFTGLYPGTLCAGLNNRIPAGYSHPLQYLKNQTVLSPLMQPLALHSFMVQRQFLTSPANSQQLYRHLAAATPVGSSYGDLLHNSIYPLAAINPQAAFPPSQLSSVHPSTKL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GKG4</accession>
    <name>A0A452GKG4_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="5">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GKG4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002410.1">
        <property type="protein sequence ID" value="ENSGAGP00000002120.1"/>
        <property type="gene ID" value="ENSGAGG00000001705.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051242">
        <property type="entry name" value="WD-EF-hand_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019775">
        <property type="entry name" value="WD40_repeat_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036322">
        <property type="entry name" value="WD40_repeat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001680">
        <property type="entry name" value="WD40_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR44324:SF8">
        <property type="entry name" value="WD REPEAT-CONTAINING PROTEIN 64"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR44324">
        <property type="entry name" value="WD40 REPEAT DOMAIN 95"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00400">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00320">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50978">
        <property type="entry name" value="WD40 repeat-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00678">
        <property type="entry name" value="WD_REPEATS_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50082">
        <property type="entry name" value="WD_REPEATS_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50294">
        <property type="entry name" value="WD_REPEATS_REGION"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <keyword evidence="1 3" id="KW-0853">WD repeat</keyword>
    <feature type="repeat" description="WD" evidence="3">
        <location>
            <begin position="1"/>
            <end position="15"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="3">
        <location>
            <begin position="147"/>
            <end position="181"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="223"/>
            <end position="251"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="240"/>
            <end position="249"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022574"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002120.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="322" mass="35971" checksum="D996595835F44360" modified="2019-05-08" version="1">MASSSYDGEILIWNLISGHVCNRLSAVSQAVMEDGTGYITFWNIFGGGKLFAYFIGSKDRSTLSDMAVSDDDCLLCAADHIGYSYIWNITKYALQGPEEEPPACEQHFLLPFQSQCPLTLSHVADSCITDLYLIFFSPSGCQVLLSWRAHSCSVTSVAWMSEHQLLLTSSVDCTVKLWSLEGEYIGTFGQKDLWNIKDRMSWQRGPNPDSYITPAAPARQVSAIKPSCATESEDEANSAEPESQVTSPLSLIKDDEIAEELKQRQKAKSEHRAHHSQLKCIKHETRALTTYHSLQLCELTCLPATFHKLNSRRCNDPDNLTF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GKG7</accession>
    <name>A0A452GKG7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">NADH dehydrogenase [ubiquinone] 1 beta subcomplex subunit 11, mitochondrial</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="15">Complex I-ESSS</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="14">NADH-ubiquinone oxidoreductase ESSS subunit</fullName>
        </alternativeName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">Accessory subunit of the mitochondrial membrane respiratory chain NADH dehydrogenase (Complex I), that is believed not to be involved in catalysis. Complex I functions in the transfer of electrons from NADH to the respiratory chain. The immediate electron acceptor for the enzyme is believed to be ubiquinone.</text>
    </comment>
    <comment type="subunit">
        <text evidence="16">Complex I is composed of 45 different subunits. Interacts with BCAP31.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Mitochondrion inner membrane</location>
            <topology evidence="2">Single-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the complex I NDUFB11 subunit family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GKG7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002574.1">
        <property type="protein sequence ID" value="ENSGAGP00000002250.1"/>
        <property type="gene ID" value="ENSGAGG00000001774.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005743">
        <property type="term" value="C:mitochondrial inner membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019329">
        <property type="entry name" value="NADH_UbQ_OxRdtase_ESSS_su"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13327:SF0">
        <property type="entry name" value="NADH DEHYDROGENASE [UBIQUINONE] 1 BETA SUBCOMPLEX SUBUNIT 11, MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13327">
        <property type="entry name" value="NADH-UBIQUINONE OXIDOREDUCTASE ESSS SUBUNIT, MITOCHONDRIAL PRECURSOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10183">
        <property type="entry name" value="ESSS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0249">Electron transport</keyword>
    <keyword evidence="13 17" id="KW-0472">Membrane</keyword>
    <keyword evidence="12" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="8" id="KW-0999">Mitochondrion inner membrane</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0679">Respiratory chain</keyword>
    <keyword evidence="9" id="KW-0809">Transit peptide</keyword>
    <keyword evidence="7 17" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 17" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="5" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="81"/>
            <end position="101"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00003195"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004434"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00008915"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00018632"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022660"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022792"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022982"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00030753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00031387"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00046528"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002250.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="147" mass="16671" checksum="9E0F63B6A31C85B8" modified="2019-05-08" version="1">MHRYMWLFSGCPAPEVAASQSWVSHPRAAVPQLPPQPSIPSPCSSLELSLCPSLSLLSSWLNPDYHGFDADPVMDVWNMRLAFFFGISLCIVVGATFLHYLPEHGMRKWARREAERMVKERERLGLPVLDSNYFDPSKLVLPAEDEE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GKG8</accession>
    <name>A0A452GKG8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">EGF-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cell projection</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the tenascin family. Teneurin subfamily.</text>
    </comment>
    <comment type="caution">
        <text evidence="13">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000002422.1">
        <property type="protein sequence ID" value="ENSGAGP00000002127.1"/>
        <property type="gene ID" value="ENSGAGG00000000665.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043005">
        <property type="term" value="C:neuron projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050839">
        <property type="term" value="F:cell adhesion molecule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046982">
        <property type="term" value="F:protein heterodimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042803">
        <property type="term" value="F:protein homodimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007157">
        <property type="term" value="P:heterophilic cell-cell adhesion via plasma membrane cell adhesion molecules"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048666">
        <property type="term" value="P:neuron development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00054">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.120.10.30:FF:000005">
        <property type="entry name" value="Teneurin transmembrane protein 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.120.10.30:FF:000006">
        <property type="entry name" value="Teneurin transmembrane protein 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.180.10.10:FF:000001">
        <property type="entry name" value="Teneurin transmembrane protein 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.180.10.10:FF:000005">
        <property type="entry name" value="Teneurin transmembrane protein 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.180.10.10">
        <property type="entry name" value="RHS repeat-associated core"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.120.10.30">
        <property type="entry name" value="TolB, C-terminal domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011042">
        <property type="entry name" value="6-blade_b-propeller_TolB-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008969">
        <property type="entry name" value="CarboxyPept-like_regulatory"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057627">
        <property type="entry name" value="FN-plug_TEN1-4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022385">
        <property type="entry name" value="Rhs_assc_core"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056823">
        <property type="entry name" value="TEN-like_YD-shell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056822">
        <property type="entry name" value="TEN_NHL"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056820">
        <property type="entry name" value="TEN_TTR-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051216">
        <property type="entry name" value="Teneurin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028916">
        <property type="entry name" value="Tox-GHH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006530">
        <property type="entry name" value="YD"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR03696">
        <property type="entry name" value="Rhs_assc_core"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01643">
        <property type="entry name" value="YD_repeat_2x"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11219">
        <property type="entry name" value="TENEURIN AND N-ACETYLGLUCOSAMINE-1-PHOSPHODIESTER ALPHA-N-ACETYLGLUCOSAMINIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11219:SF9">
        <property type="entry name" value="TENEURIN-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23106">
        <property type="entry name" value="EGF_Teneurin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24329">
        <property type="entry name" value="FN-plug_TEN1-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25021">
        <property type="entry name" value="TEN_NHL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25023">
        <property type="entry name" value="TEN_YD-shell"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23538">
        <property type="entry name" value="Teneurin_ABD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15636">
        <property type="entry name" value="Tox-GHH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25020">
        <property type="entry name" value="TTR_TEN1-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF63829">
        <property type="entry name" value="Calcium-dependent phosphotriesterase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49464">
        <property type="entry name" value="Carboxypeptidase regulatory domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01186">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="12" id="KW-0966">Cell projection</keyword>
    <keyword evidence="11 13" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6 13" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="10" id="KW-0472">Membrane</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9" id="KW-1133">Transmembrane helix</keyword>
    <feature type="domain" description="EGF-like" evidence="14">
        <location>
            <begin position="143"/>
            <end position="178"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="13">
        <location>
            <begin position="147"/>
            <end position="157"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="13">
        <location>
            <begin position="168"/>
            <end position="177"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004167"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004316"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00009385"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002127.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2107" mass="236865" checksum="A676455D84610E97" modified="2019-05-08" version="1">RCEEGWMGAACDQRACHPRCNEHGTCRDGKCECSPGWNGDVSYDTFLSLCLSLTHTQTHSHTHTLTHSLTVSLPPSPGIGPSPSAGKFLFCLSILRMEHVQLLWVDAAYGVRLIHSSCKGDGACLMRQNLWKIYALECVIVCFAEGCPGLCNGNGRCTLDMNGWHCVCQLGWRGAGCDTSMETACGDGKDNDGDGLVDCMDPDCCLQPLCHVNPLCLGSPDPLDIIQETQTPTSQQNLHSFYDRIKFLIGKDSTHVIPGENPFDGGHACVIRGQVITSDGTPLVGVNISFVNNPLFGYTISRQDGSFDLVTNGGISIILHFERAPFITQEHTLWLPWDRFFVMETIVMRHEENEIPSCDLSNFARPNPVVSPSPLTAFASSCSEKGPIVPEIQALQEEINIAGSKIKLSYLSSRTAGYKSVLRISMTHPTIPFNLMKVHLMVAVEGRLFRKWFAAAPDLSYYFIWDKTDVYSQKVYGLSEAFVSVGYEYESCPDLILWEKRTAVLQGYEIDASKLGGWSLDQHHALNIQSGILHKGNGENQFVSQQPPVIGSIMGNGRRRSISCPSCNGLADGNKLLAPVALTCGSDGSLYVGDFNYIRRIFPSGNVTNILELRNKDFRHSHSPAHKYYLTSDPITGSIYLSDTNSRRIYKIKSTSVVKDVVKNSEVVAGTGDQCLPFDETRCGDGGKGTEATLTNPRGKTQSLLTSILKWASRKVPNQQFWNTTEQVYKLPQGNPLMCLRVRLEWPTDLAVSPMDNSLYVLDNNVVLQISENHQVRIVAGRPMHCQVPGIDHFLLSKVAIHATLESATALAVSHNGVLYIAETDEKKINRIRQVTTNGEISLVAGAPSSCDCKNDANCDCFSGDDGYAKDARLNAPSSLAVCADGELYVADLGNIRIRFIRKNKPYLNTQNLYELSSPIDQELYLFDTSGKHLYTQSLPTGDYLYNFTYTGDGDITLITDNNGNLVNVRRDSTGMPLWLVVPDGQVYWVTIGTNSALKSVTAQGHEVGMMTYHGNSGLLATKTNENGWTTFYEYDSFGRLTNVTFPTGQVSSFRSDTDRSVHVQVETSSKDDVTITTNLSASGAFYTLLQDQVRNSYYIGADGSLRLMLANGMEVALQTEPHLLAGTVNPTVGKRNVTLPIDNGLNLVEWRQRKEQARGQVTVFGRRLRVHNRNLLSLDFDRVTRTEKIYDDHRKFTLRILYDQAGRPSLWSPSSRLNGVNVTYSPGGHIAGIQRGTMSERMEYDQAGRIISRIFADGKSWSYTYLEKSMVLLLHSQRQYIFEFDKNDRLSSVTMPNVARQTLETIRSIGYYRNIYRPPEGNASVIQDFTEEGQLLHTFYLGTGRRVIYKYGKLSKLAEMLYDTTKIGFTYDETAGMLKTINLQNEGFTCTIRYRQIGPLIDRQIFRFTEEGMVNARFDYNYDNSFRVTSMQAVINETPLPIDLYRYDDVSGKTEQFGKFGVIYYDINQIITTAVMTHTKHFDAYGRMKEVQYEIFRSLMYWMTVQYDNMGRVVKKELKVGPYANTTRYSYEYDADGQLQTVSINDKPLWRYSYDLNGNLHLLSPGNSARLTPLRYDLRDRITRLGDVQYKMDEDGFLRQRGNDIFEYNSAGLLIKAYNKVSGWSVKYRYDGLGRRVSSKTAHSHHLQFFYADLTNPTKVTHLYNHSSSEITSLYYDLQGHLFAMELSSGDEFYIACDNIGTPLAVFSGTGLMIKQILYTAYGEIYMDTNPNFQVIIGYHGGLYDPLTKLIHMGRRDYDVLAGRWTSPDHDMWKHLSSNNIMPFNLYMFKNNNPISNAQDIKCYMTDVNSWLLTFGFQLHNVIPGYPKPDTDAMEPSYELIHTQMKTQEWDNSKSILGVQCEVQKQLKAFVTLERFEQIYSSSIAGCRTVKENKNFASGGSVFGKGVKFAMKDGRVATDIISLANEDGRRIAAILNNAHYLENLHFTIEGVDTHYFVKQGPSESDLSILGLSGGRRTLENGVNVTVSQINTVLSGRTRRYTDIQLQYGALCLNTRYGTTLDEEKARVLELARQRAIAQAWAREQQRLRDGEEGIRSWTEGEKQQVLNTGRVQGYDGYFVISVEQYPELSDSANNIHFMRQSEMGRR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GKG9</accession>
    <name>A0A452GKG9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Kinesin-like protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="5 6">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Kinesin family.</text>
    </comment>
    <comment type="caution">
        <text evidence="5">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GKG9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002537.1">
        <property type="protein sequence ID" value="ENSGAGP00000002219.1"/>
        <property type="gene ID" value="ENSGAGG00000000085.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005874">
        <property type="term" value="C:microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008017">
        <property type="term" value="F:microtubule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003777">
        <property type="term" value="F:microtubule motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035091">
        <property type="term" value="F:phosphatidylinositol binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007018">
        <property type="term" value="P:microtubule-based movement"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd22732">
        <property type="entry name" value="FHA_KIF16B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd06874">
        <property type="entry name" value="PX_KIF16B_SNX23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.200.20:FF:000005">
        <property type="entry name" value="Kinesin family member 16B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1520.10:FF:000022">
        <property type="entry name" value="Kinesin family member 16B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000167">
        <property type="entry name" value="Uncharacterized protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.200.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1520.10">
        <property type="entry name" value="Phox-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000253">
        <property type="entry name" value="FHA_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019821">
        <property type="entry name" value="Kinesin_motor_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001752">
        <property type="entry name" value="Kinesin_motor_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001683">
        <property type="entry name" value="PX_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036871">
        <property type="entry name" value="PX_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008984">
        <property type="entry name" value="SMAD_FHA_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47117:SF8">
        <property type="entry name" value="KINESIN FAMILY MEMBER 16B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47117">
        <property type="entry name" value="STAR-RELATED LIPID TRANSFER PROTEIN 9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00498">
        <property type="entry name" value="FHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00225">
        <property type="entry name" value="Kinesin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00787">
        <property type="entry name" value="PX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00380">
        <property type="entry name" value="KINESINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00129">
        <property type="entry name" value="KISc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00312">
        <property type="entry name" value="PX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF64268">
        <property type="entry name" value="PX domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49879">
        <property type="entry name" value="SMAD/FHA domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00411">
        <property type="entry name" value="KINESIN_MOTOR_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50067">
        <property type="entry name" value="KINESIN_MOTOR_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50195">
        <property type="entry name" value="PX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 6" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="3 7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="6" id="KW-0493">Microtubule</keyword>
    <keyword evidence="4 6" id="KW-0505">Motor protein</keyword>
    <keyword evidence="1 6" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="8">
        <location>
            <begin position="1"/>
            <end position="17"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5018978517" description="Kinesin-like protein" evidence="8">
        <location>
            <begin position="18"/>
            <end position="994"/>
        </location>
    </feature>
    <feature type="domain" description="Kinesin motor" evidence="9">
        <location>
            <begin position="1"/>
            <end position="166"/>
        </location>
    </feature>
    <feature type="domain" description="PX" evidence="10">
        <location>
            <begin position="859"/>
            <end position="994"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="384"/>
            <end position="621"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="703"/>
            <end position="790"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00283"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="RuleBase" id="RU000394"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50067"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50195"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002219.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="994" mass="115798" checksum="6FED16C1D0BC1643" modified="2019-05-08" version="1">ITFISILVLIYNHFCFCSDLSKHLVQNYSDVEELMEAGNINRTTAATGMNDVSSRSHAIFTINFTQAKFDAEMPCETVSKIHLVDLAGSERADATGATGVRLKEGGNINKSLVTLGNVISALADLSQDATNPLAKKKQVFVPYRDSVLTWLLKDSLGGNSKTIMIAIRLTRYVFYNTNVIHRELFIIALLDSPTALSMEEKLQQNEARVQELTKEWTNKWNETQNILKEQTLALRKEGIGVVLDSELPHLIGIDDDLLSTGIILYHLKEGQTYVGREDAATEQDIVLHGLDLESEHCVFENLSGTVNLIPLNGAQCSVNGIQITEATQLNQGAVILLGRTNMFRFNHPKEAAKLREKRKSGLLSSFSLSMTDLSKSLYSKQKLIEEMEEKQKSDKAELERMQQEVESQRKETEIVQLEIRKQEESLKRRSFHIESRLKDLLAEKEKFEEERLREHQEIEFQKTKQQEEIFARVKEELQRLQELNNNEKVEKMQIFRELEKLKKEKDEQCIKLELEKKRLEEQEREQVMQVAHLEGQLREKQVMIQLLKRGDVQRVEEEKRELEEIRESLLRVKEARSEGDEDSEELEKAQHNFVDFKRKQLEQLTILEKDLFQQRDHLEKEVADEHETLDRLKYAHKERLNLLKKDDENVVDATLMVEEFDKIKPAEYRLQYKERQLQYLMHNHLPALLEEKQRATEVLDRGLLGLDNTLYQVEKEMEEKEEQLVQYRVSTNQLQQLQETFEFTANIARQEEKVRKKEKEILESREKQQREALEQAVAKLERRHSALQRSSTIDVEIEEQKQKLAILNSSCSEQAGLQASLEAEQKDNEMLHNGTVQRKLKYERMICRSLGANSDDLKDPIKISIPRYVLCGQGKDEHYEFEIKITVLDETWTVFRRYSRFREMHKTLKLKYPELATLEFPPKKLFGNKDERVIAERRSHLEKYLRNFFCMMLKSPSSPLHIDKVGPTLSKHTICEFSPFFKKGVFDYSSHGTS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GKH0</accession>
    <name>A0A452GKH0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">NADH dehydrogenase [ubiquinone] 1 beta subcomplex subunit 11, mitochondrial</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="15">Complex I-ESSS</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="14">NADH-ubiquinone oxidoreductase ESSS subunit</fullName>
        </alternativeName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">Accessory subunit of the mitochondrial membrane respiratory chain NADH dehydrogenase (Complex I), that is believed not to be involved in catalysis. Complex I functions in the transfer of electrons from NADH to the respiratory chain. The immediate electron acceptor for the enzyme is believed to be ubiquinone.</text>
    </comment>
    <comment type="subunit">
        <text evidence="16">Complex I is composed of 45 different subunits. Interacts with BCAP31.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Mitochondrion inner membrane</location>
            <topology evidence="2">Single-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the complex I NDUFB11 subunit family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000002580.1">
        <property type="protein sequence ID" value="ENSGAGP00000002256.1"/>
        <property type="gene ID" value="ENSGAGG00000001774.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005743">
        <property type="term" value="C:mitochondrial inner membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019329">
        <property type="entry name" value="NADH_UbQ_OxRdtase_ESSS_su"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13327:SF0">
        <property type="entry name" value="NADH DEHYDROGENASE [UBIQUINONE] 1 BETA SUBCOMPLEX SUBUNIT 11, MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13327">
        <property type="entry name" value="NADH-UBIQUINONE OXIDOREDUCTASE ESSS SUBUNIT, MITOCHONDRIAL PRECURSOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10183">
        <property type="entry name" value="ESSS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0249">Electron transport</keyword>
    <keyword evidence="13 17" id="KW-0472">Membrane</keyword>
    <keyword evidence="12" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="8" id="KW-0999">Mitochondrion inner membrane</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0679">Respiratory chain</keyword>
    <keyword evidence="9" id="KW-0809">Transit peptide</keyword>
    <keyword evidence="7 17" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 17" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="5" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="85"/>
            <end position="105"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00003195"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004434"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00008915"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00018632"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022660"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022792"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022982"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00030753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00031387"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00046528"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002256.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="151" mass="17115" checksum="93C9ECB16E187FDB" modified="2019-05-08" version="1">MVPGRLLWECHRQTVERQIGRRLVTGRLHWGESAGXGCQVRGQLGEGRAIPLESCERSGYCGGGGNPDYHGFDADPVMDVWNMRLAFFFGISLCIVVGATFLHYLPEHGMRKWARREAERMVKERERLGLPVLDSNYFDPSKLVLPAEDEE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GKH3</accession>
    <name>A0A452GKH3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Centrosomal protein of 164 kDa</fullName>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Plays a role in microtubule organization and/or maintenance for the formation of primary cilia (PC), a microtubule-based structure that protrudes from the surface of epithelial cells. Plays a critical role in G2/M checkpoint and nuclear divisions. A key player in the DNA damage-activated ATR/ATM signaling cascade since it is required for the proper phosphorylation of H2AX, RPA, CHEK2 and CHEK1. Plays a critical role in chromosome segregation, acting as a mediator required for the maintenance of genomic stability through modulation of MDC1, RPA and CHEK1.</text>
    </comment>
    <comment type="subunit">
        <text evidence="15">Interacts (via N-terminus) with ATRIP. Interacts with ATM, ATR and MDC1. Interacts with XPA (via N-terminus) upon UV irradiation. Interacts with CEP83, CCDC92, TTBK2, DVL3, NPHP3 and weakly with NPHP4. Interacts with DZIP1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Microtubule organizing center</location>
            <location evidence="1">Centrosome</location>
            <location evidence="1">Centriole</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000002430.1">
        <property type="protein sequence ID" value="ENSGAGP00000002133.1"/>
        <property type="gene ID" value="ENSGAGG00000001670.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005814">
        <property type="term" value="C:centriole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097539">
        <property type="term" value="C:ciliary transition fiber"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060271">
        <property type="term" value="P:cilium assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006281">
        <property type="term" value="P:DNA repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00201">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1470.10:FF:000001">
        <property type="entry name" value="Centrosomal protein of 164 kDa"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1470.10">
        <property type="entry name" value="Photosystem I PsaD, reaction center subunit II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051841">
        <property type="entry name" value="MT-Golgi_org_protein"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001202">
        <property type="entry name" value="WW_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036020">
        <property type="entry name" value="WW_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18902:SF27">
        <property type="entry name" value="CENTROSOMAL PROTEIN OF 164 KDA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18902">
        <property type="entry name" value="NUCLEAR MITOTIC APPARATUS PROTEIN 1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00397">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00456">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51045">
        <property type="entry name" value="WW domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01159">
        <property type="entry name" value="WW_DOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50020">
        <property type="entry name" value="WW_DOMAIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="13" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="5" id="KW-0132">Cell division</keyword>
    <keyword evidence="8" id="KW-0970">Cilium biogenesis/degradation</keyword>
    <keyword evidence="9 17" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="6" id="KW-0227">DNA damage</keyword>
    <keyword evidence="10" id="KW-0234">DNA repair</keyword>
    <keyword evidence="7" id="KW-0498">Mitosis</keyword>
    <keyword evidence="12" id="KW-0539">Nucleus</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="WW" evidence="19">
        <location>
            <begin position="58"/>
            <end position="91"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="105"/>
            <end position="151"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="163"/>
            <end position="215"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="237"/>
            <end position="263"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="315"/>
            <end position="372"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="434"/>
            <end position="467"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="505"/>
            <end position="537"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="572"/>
            <end position="604"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="630"/>
            <end position="660"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="845"/>
            <end position="896"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1041"/>
            <end position="1077"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1093"/>
            <end position="1125"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1683"/>
            <end position="1702"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="17">
        <location>
            <begin position="1139"/>
            <end position="1627"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="18">
        <location>
            <begin position="110"/>
            <end position="126"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="127"/>
            <end position="138"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="169"/>
            <end position="194"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="241"/>
            <end position="257"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="315"/>
            <end position="324"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="337"/>
            <end position="353"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="441"/>
            <end position="450"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="580"/>
            <end position="600"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="633"/>
            <end position="658"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="1044"/>
            <end position="1053"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="1060"/>
            <end position="1074"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="1102"/>
            <end position="1117"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="1687"/>
            <end position="1702"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004114"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022776"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022794"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00056906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00061715"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00067900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50020"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002133.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2007" mass="225712" checksum="A111157F2A3F8380" modified="2019-05-08" version="1">MDMAGPAIRIGDQLILEEDYDETYIPSEQEIQEFAREIGIDPGNEPELMWLAREGIVAPLPPEWKPCQDITGDIYYFNFSNGQSTWDHPCDEHYRELVIQEREKLSAHGSIKKKDKKKKKDKKEKKEKKEREPVKHAAEMQPEPGILPSSSFYRMSSPILSSECASPDLDQHSQMRIEGFLKKGKRKPSERPPETSDSNKQFAGPSPNKLQPLFTAKSCRTHQILADVEKILGRMPSFSRSDVDHQQGQDRSPKIGDKPCLGFSDSESEGLDVVRTKPISYVARKSCSQDLENGRSVLDSCEVLEDKGMLFDEEKLSDNWKQEEMTGGSVCQPDDGLSVRERSTGPGGDKERGPGFSQADENEAFLTPSAPDHTFYSRKCEMILFDASPLEDLSLMEEGDVHCPQRELRAHSRAVESRRKRKLQDQALSQATKFCKSSEVVGKHERESSKQAEFVESEGAGLQAAPQKDLLDSHLAVPEVTADTLPYTRGVEAYAHAGADISASEPVSLSAAQHKDKEKANGLGPEPDNSLTSNSADHFASQILGEIDNFSWDLQSSRESDRPTDLLAAAAKGSLPTHPFTGTLQTPPRSSANGKSQSDCNSEDEKFYQHVQENDQVPSQEHSCVSFNMKLNRSRDAPEKNNEGRQEETGEGEQKTEPVPRGLEVDVFTEGLQVSPGWLYQTAECQESSRLCQQVEGELTQSGKVTEDYCKLKLCEGAGSNESPGSLLAPIQVPPGSLAPLRGVMDGPASAFHGLLSTNVEISGVPSPLLREGTLPLQTLKSTGCTKSLLGSMHEEKASLNLLALDEEEEEESENQSPRGTARLLKNLHMDISALGGGFEYEESLKMSHAEESQTFPLDSDGVRPPAPDKLLNRDADSSLCGLNKEESSGKPLGNELLEKEDTDVEDTVSVAYEQSGTLGERTAEGTHVVSSQGHPVAASPDVRAAETDQMESVAVTVDAVSVSEKIVNGMREEAADLETDSKLDAGKESEASEHVKELQVSDCSDHELLHLTDFGFQSRISEQVLDMDILSAALDSPKVEAQGLREEEKDQSEASLEEEQSKRPKAAESERNQRPFSACKMAEEKCLSLENASQEETAALEQKEGSLDSPKEEEPAAPHQAQLGEEIKQVVSLNWPSAESLNEIARKQEQELEQEKERLLQAKEERVRRFQEELRQEEEEEAQVLHQQKEKSLWSLKEELAKASEEEELRMREEEVEKLSKLRSQISSETKTEKEKIRAEQEAALQKLREEWESLQKVERERLEERKKRVLEMLKVEVEEAQQREAAELEQEKQRVLNELKERLERKRKEAAEVLEKQFAAELQELKSTAEEKHQKVVSSLRKQIAEAQRSEEAQLREELERAEQKVQQKMYQVAEFERELSELMKEKRQEVERDHDRKMERMKEEHKEVLARIQEQYEEEERKQRTKKLEELKGELERLRQLHDGELKALQKQLDEQLSDLRQSHKVKERKLQELEMELEIQATDAKARCAQLNHQEEAMRKKRQQLLDEEKQTELQRNEAALAAQLRLEESRKEHADLVESIRQLRRTLEELQDQKAELESQVDLLQTRSQRLQRRISELEAAVQSKQETLKELVAEDSMASPRREAELHLEDLRETLQAHSSREPASLTSHSNEDSDLQLDNVRHFISAEGVSIKNAKEFLVRQTRSMRKRHTALKAAKQQWRHNMQQAQDAVQDPDSSQLLEDVRRNLEEESAQLDKMKSAMQKGQVLLKKKEEKLSQLESSLLEELSDEDTLKGAACKKVVTFDLSDSEETHSLSSTDVPQHKFDLKPDVQFPQLDKIQHLTSTLQHITAELNGVLSTLGSLNQRQPPLFAPVQVPMTALPSDGVPLSMYTSLARAQAAGPIMPPAGIPLVNQWAWSTGLNSSPSSTAGQSVDDILAEKWHKYFPGGFPSLSGSPAPLDSNLGYLPAGEQIGLFQRSQLQGPETDSTSVQGMIDANKKWLENFKKDSKVPLFPRTQKPFASSPGLVQLGLDENNQIKVYHC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GKH5</accession>
    <name>A0A452GKH5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">PLAC domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GKH5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002555.1">
        <property type="protein sequence ID" value="ENSGAGP00000002232.1"/>
        <property type="gene ID" value="ENSGAGG00000001788.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031012">
        <property type="term" value="C:extracellular matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004222">
        <property type="term" value="F:metalloendopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030198">
        <property type="term" value="P:extracellular matrix organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.830:FF:000001">
        <property type="entry name" value="A disintegrin and metalloproteinase with thrombospondin motifs 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000005">
        <property type="entry name" value="ADAM metallopeptidase with thrombospondin type 1 motif 9"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.830">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.20.100.10">
        <property type="entry name" value="Thrombospondin type-1 (TSP1) repeat"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050439">
        <property type="entry name" value="ADAMTS_ADAMTS-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045371">
        <property type="entry name" value="ADAMTS_CR_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010294">
        <property type="entry name" value="ADAMTS_spacer1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010909">
        <property type="entry name" value="PLAC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000884">
        <property type="entry name" value="TSP1_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036383">
        <property type="entry name" value="TSP1_rpt_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13723">
        <property type="entry name" value="ADAMTS A DISINTEGRIN AND METALLOPROTEASE WITH THROMBOSPONDIN MOTIFS PROTEASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13723:SF144">
        <property type="entry name" value="ADAMTS-LIKE PROTEIN 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19236">
        <property type="entry name" value="ADAMTS_CR_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05986">
        <property type="entry name" value="ADAMTS_spacer1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08686">
        <property type="entry name" value="PLAC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19030">
        <property type="entry name" value="TSP1_ADAMTS"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00090">
        <property type="entry name" value="TSP_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00209">
        <property type="entry name" value="TSP1"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF82895">
        <property type="entry name" value="TSP-1 type 1 repeat"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50900">
        <property type="entry name" value="PLAC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50092">
        <property type="entry name" value="TSP1"/>
        <property type="match status" value="6"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="3 6" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="6">
        <location>
            <begin position="1"/>
            <end position="29"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019311771" description="PLAC domain-containing protein" evidence="6">
        <location>
            <begin position="30"/>
            <end position="1089"/>
        </location>
    </feature>
    <feature type="domain" description="PLAC" evidence="7">
        <location>
            <begin position="1043"/>
            <end position="1080"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="136"/>
            <end position="199"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="225"/>
            <end position="278"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="299"/>
            <end position="376"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="5">
        <location>
            <begin position="353"/>
            <end position="363"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002232.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1089" mass="119841" checksum="FBCA03E3B7399AA3" modified="2019-05-08" version="1">MLTGQVCAPQPSWLPFSPVFWLLFPLLWSLPRYQLSCCRIQEALFAQIFPSVPAPETWAEAAPWAPNATPEPWLLDQALRCVPQRTSRQVPEEDEAGNQVAGAWSSWGPWSSCSQPCGVGVLERVRTCLPPYQRAPWGGPHGPHTYAQPLYPEERPNPYPPRPSYPLHTDGDSRHGALGSKWVVSPTQESRAARRSRIRESIKPGKYGYGKVPFALPLHKEVAEGGTPRFKRHPGPSSDQPPPPTKTPKRKAKEPSRPGIPVGERAPSSGRTPRPRGELRLTHEGLKNHSSALEAQLVPKGRGAGGGHKEAPELSQPHSHSHRRATGAPKPRPEATHLTQPEGAGPVDPPARSHPHQLARPRAQRQNEYRPGGYGTRASQSLFLEQPSRHRLAEPDIWLLHRGSPVQFQARGQRPRLGLHPGPEVPQWNLYHPGTETFHCEGESKQFKACRQEPCPADQPDARAVQCAAFNTQEFMGRLYQWEPFTDGKGLGARTGMRCSHVRGSQRCELNCRPVGFRFYVRHTEKVQDGTPCEAASLDICVAGECLSPGCDGVLGSNRTLDSCGVCGGDHSTCKLVVGNFSDSNVPIGYHKILQIPQGATRINVTEMTQSPNYLALRSRSGKSLINGNWAVDPPGRYEAGGTVFIYTRPGHEEPRGGGESLTAEGPTTEPIDVYMIFQQDNPGVAYPVSMTPVLYGTLFPSPPRVSAGLWQPVFRCAARHSQEEVEEEQCADAPKPPALDEACNTQPCPAYWDMGEWSACSKSCGPGTQHRQVLCRQTYANRSTMVHPQHCSQLDKPSPTQPCQLRVCSHWEVRTNWSCSVLCGLGHRTRHIRCISNHGDLLSDGECNGSHRPPANEACDMGPCVRSWFHSDWSNTCSVECGTGIQRRSVVCLSSYANGQVEETCAGTKPADMRACNSGPCHRTVKWYTGPWSECSADCGTGTQRRDVICVSKLGSEFNVTEASECAPLEKPPSLQPCTGSACDARWFSTSWSACSKSCVGGVQVREVQCLTQNKTFSNLCQPDLKPIKKRSCNTQPCTPDLGENCRDKYHNCPVVVQARLCVYSYYKLVCCASCTRAVERSKDMQSR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GKI0</accession>
    <name>A0A452GKI0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">NF-kappa-B inhibitor zeta</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="11">I-kappa-B-zeta</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="12">Molecule possessing ankyrin repeats induced by lipopolysaccharide</fullName>
        </alternativeName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="8">Involved in regulation of NF-kappa-B transcription factor complexes. Inhibits NF-kappa-B activity without affecting its nuclear translocation upon stimulation. Inhibits DNA-binding of RELA and NFKB1/p50, and of the NF-kappa-B p65-p50 heterodimer and the NF-kappa-B p50-p50 homodimer. Also seems to activate NF-kappa-B-mediated transcription. In vitro, upon association with NFKB1/p50 has transcriptional activation activity and, together with NFKB1/p50 and RELA, is recruited to LCN2 promoters. Promotes transcription of LCN2 and DEFB4. Is recruited to IL-6 promoters and activates IL-6 but decreases TNF-alpha production in response to LPS. Seems to be involved in the induction of inflammatory genes activated through TLR/IL-1 receptor signaling. Involved in the induction of T helper 17 cells (Th17) differentiation upon recognition of antigen by T cell antigen receptor (TCR).</text>
    </comment>
    <comment type="subunit">
        <text evidence="9">Interacts with NFKB1/p50. Interacts with RELA. Interacts with AKIRIN2.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GKI0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002561.1">
        <property type="protein sequence ID" value="ENSGAGP00000002237.1"/>
        <property type="gene ID" value="ENSGAGG00000001827.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070974">
        <property type="term" value="F:POU domain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031349">
        <property type="term" value="P:positive regulation of defense response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010468">
        <property type="term" value="P:regulation of gene expression"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000179">
        <property type="entry name" value="NF-kappa-B inhibitor zeta isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000188">
        <property type="entry name" value="NF-kappa-B inhibitor zeta isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047571">
        <property type="entry name" value="OCA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24124">
        <property type="entry name" value="ANKYRIN REPEAT FAMILY A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24124:SF5">
        <property type="entry name" value="NF-KAPPA-B INHIBITOR ZETA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01415">
        <property type="entry name" value="ANKYRIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS52003">
        <property type="entry name" value="OCA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0010">Activator</keyword>
    <keyword evidence="4 13" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0804">Transcription</keyword>
    <keyword evidence="3" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="OCA" evidence="15">
        <location>
            <begin position="8"/>
            <end position="30"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="13">
        <location>
            <begin position="375"/>
            <end position="407"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="13">
        <location>
            <begin position="508"/>
            <end position="530"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="13">
        <location>
            <begin position="544"/>
            <end position="580"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="162"/>
            <end position="197"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="172"/>
            <end position="197"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
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        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00057515"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00064472"/>
        </source>
    </evidence>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00075776"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00078782"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="PROSITE" id="PS52003"/>
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    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002237.1"/>
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    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="614" mass="67858" checksum="829817860CBE226A" modified="2019-05-08" version="1">MGGGKQHRGPFQGVRVKNSVKELLLHIRSNKKMSSGHVADEFKEQAGLMNYEQFTELKNILSHSGKRKAHESLCDGPSYKRPATFHSHLLTPPQTPTSMDNMEEAHKNEPKHNCSDMLQNIINIKNESNPVSLNTVQVSWLHSISNHNSPGERYQDIQGAQAFSPSEKYQAFQANSPQQMLDPPQNYQFSSSQTQDLPQKYTQDSLLEYRPFSGNDQSPAYQQNVFESHELPYCPGQSFASLLNESEHSENILQPLATAHQQSDVNAPAQNFGLVQNNNCSTLDGHNSSLAALNASLQHQGIVRNTTQLGKSFFQWQVEQEENKLANISQEQILAKDSDGDTFLHIAVAQGRRALSYVLARKMAALHMLDIKEHNGQSAFQVAVAANQHLIVQDLVSLGAQVNTTDCWGRTPLHVCAEKGHSQVLQAIQKGAIGSSQYVALEATNYDGFTALHCAVLAHNAVVHELRNSQPPHSPEVQELLLKNKSLVDTIKALIQMGASVEAKDGKSGRSALHLAAEEANLELIRFFLDLPNCLSFVNAKAYNGNTALHVAASLQYRLTQLDAVRLLMRKGADPSARNLENEQPVHLVPDGPVGEQIRRILKGKAVQQRASPY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GKI7</accession>
    <name>A0A452GKI7_9SAUR</name>
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        <recommendedName>
            <fullName evidence="4">Mitotic spindle assembly checkpoint protein MAD2B</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="5">Mitotic arrest deficient 2-like protein 2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GKI7"/>
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        <property type="gene ID" value="ENSGAGG00000001763.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006974">
        <property type="term" value="P:DNA damage response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.900.10:FF:000003">
        <property type="entry name" value="Mitotic spindle assembly checkpoint protein MAD2B"/>
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    </dbReference>
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        <property type="entry name" value="HORMA domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003511">
        <property type="entry name" value="HORMA_dom"/>
    </dbReference>
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        <property type="entry name" value="HORMA_dom_sf"/>
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    <dbReference type="InterPro" id="IPR045091">
        <property type="entry name" value="Mad2-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11842">
        <property type="entry name" value="MITOTIC SPINDLE ASSEMBLY CHECKPOINT PROTEIN MAD2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11842:SF10">
        <property type="entry name" value="MITOTIC SPINDLE ASSEMBLY CHECKPOINT PROTEIN MAD2B"/>
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    </dbReference>
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    </dbReference>
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        <property type="entry name" value="The spindle assembly checkpoint protein mad2"/>
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    </dbReference>
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            <dbReference type="ARBA" id="ARBA00022763"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
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    </evidence>
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    </evidence>
    <evidence type="ECO:0000256" key="5">
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            <dbReference type="ARBA" id="ARBA00044264"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
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    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50815"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002274.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="184" mass="20848" checksum="99B2F4DE33485E87" modified="2019-05-08" version="1">MTTLTRQDLNFGQVVADVLSEFLEVAVHLILYVREVYPIGIFQKRKKYNVPVQMSCHPELNQYIQDTLHCVKPLLEKNDVEKVVVVILDKEHHPVERFVFEITQPPLLSISSDSLLSHVEQLLRAFILKISVCDAVLDNNPPGRIESLLFSRLHLHCPGSYTGGCHTKHGEDPGDKGFPMDPGR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNE3</accession>
    <name>A0A452HNE3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Protein capicua homolog-like C-terminal tri-helical domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000018836.1">
        <property type="protein sequence ID" value="ENSGAGP00000016504.1"/>
        <property type="gene ID" value="ENSGAGG00000012212.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000977">
        <property type="term" value="F:RNA polymerase II transcription regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052412">
        <property type="entry name" value="CC-Dev_Transcription_Reg"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058606">
        <property type="entry name" value="HTH_Cic_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13059">
        <property type="entry name" value="HMG-BOX TRANSCRIPTION FACTOR BBX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13059:SF13">
        <property type="entry name" value="PROTEIN CAPICUA HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25981">
        <property type="entry name" value="HTH_Cic_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="5" id="KW-0539">Nucleus</keyword>
    <keyword evidence="1" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0804">Transcription</keyword>
    <keyword evidence="2" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="Protein capicua homolog-like C-terminal tri-helical" evidence="7">
        <location>
            <begin position="916"/>
            <end position="970"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="1"/>
            <end position="157"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="400"/>
            <end position="433"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="677"/>
            <end position="801"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="813"/>
            <end position="883"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="970"/>
            <end position="1035"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="1"/>
            <end position="10"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="6">
        <location>
            <begin position="11"/>
            <end position="21"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="30"/>
            <end position="40"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="6">
        <location>
            <begin position="41"/>
            <end position="52"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="141"/>
            <end position="151"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="6">
        <location>
            <begin position="407"/>
            <end position="420"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="6">
        <location>
            <begin position="685"/>
            <end position="696"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="741"/>
            <end position="753"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="791"/>
            <end position="801"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="1015"/>
            <end position="1025"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF25981"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016504.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1035" mass="104448" checksum="849489AC15D2ECAA" modified="2019-05-08" version="1">MTSDEERMVICEEEGDDDVIADDGFNPPDIDLKCKERVTDSESDISSGEEPDGQGFGRKLFSPVIRSPPLPPSSGAFAPCRPPAPPELDLPHGPAQPYSPFPLALGPFKAQDSRGPRPPAPPPAKRPEPATPYRRKRADSAGGAEPGAAGPSVISSPGGVLQALVLPEPGRLSAGTVLVPAPSLSSCGGAPGLARTASTVVTNVVKPVSSTPVPIASKPFPRGAGEGGPLLSPPPLEPAPPPLAVLGGAKPEAVPVGRIGLLYADKKPPPAASQAPSPHVVVGPMSKVPAAPGNMVTNLLVGQAGGTASGPAPVTVLPPQPSVQFITQNPPGAGPNGPVPLGILQPQGLLSGAPGKAGGITQVQYILPTLPQQLQVAGGKGSGAGTASIHFTLPPTSSKVLATAPHAPAPPSPSSPPPQSVSPVQAPGPGGSAHLVQGKVLVPVAAPQVTVRSGGGAGPMAQVAPPFGVPLQNGSQPTSKIIQLTPLTPAPNSAALAPPLSPATALVGPHSQAQKVLLPSSTRITYVQSTGGHQLALGTSASPSQAAATPAATYVQAPVGPAPMALGFTAIGPSGPAIVQPLLSGQSPLLAPGQVGVSPLPSPQLPPSCSGGPVITAIYPGPPSAGPAPSPAHGLVYSVASPAPAPILPKGSSAGSQGTMGPGPMGMNGAGLGRAPFIPTTWPQNGGGAMEQGAGHGSTHMATEWGRGXSPLQTPRGPVSPDWRVPGPESRPEAPAPPAGPSSSSALSPAPAGSGDGSRGAGPTPSVPEGPERKEGAGGKKVKVRPPPLKKTFDSVDNRVLSEVDFEERFAELPEFKPEEVLPSPTLQSLATSPRAILGSYRKKRKNSTDLDSSTEDPISPKRKMRRRSSCSSEPNTPKSAKCEGDIFTFDKTGTEAEDVLGELEYEKVPYSSLRRTLDQRRALVMQLFQDHGFFPSAQATAAFQARYSDIFPTKVCLQLKIREVRQKIMQSATPTEQAPSTPEPGGPGSGSAAEGQPPEPVPPQDSDPGAEAGGSTSWDNSQMSPPGGAAGSSR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNE7</accession>
    <name>A0A452HNE7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Nicotinamide-nucleotide adenylyltransferase</fullName>
            <ecNumber evidence="20">2.7.7.1</ecNumber>
            <ecNumber evidence="20">2.7.7.18</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>beta-nicotinamide D-ribonucleotide + ATP + H(+) = diphosphate + NAD(+)</text>
            <dbReference type="Rhea" id="RHEA:21360"/>
            <dbReference type="ChEBI" id="CHEBI:14649"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:33019"/>
            <dbReference type="ChEBI" id="CHEBI:57540"/>
            <dbReference type="EC" id="2.7.7.1"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="18">
            <dbReference type="Rhea" id="RHEA:21361"/>
        </physiologicalReaction>
        <physiologicalReaction direction="right-to-left" evidence="18">
            <dbReference type="Rhea" id="RHEA:21362"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>nicotinate beta-D-ribonucleotide + ATP + H(+) = deamido-NAD(+) + diphosphate</text>
            <dbReference type="Rhea" id="RHEA:22860"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:33019"/>
            <dbReference type="ChEBI" id="CHEBI:57502"/>
            <dbReference type="ChEBI" id="CHEBI:58437"/>
            <dbReference type="EC" id="2.7.7.18"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="17">
            <dbReference type="Rhea" id="RHEA:22861"/>
        </physiologicalReaction>
        <physiologicalReaction direction="right-to-left" evidence="17">
            <dbReference type="Rhea" id="RHEA:22862"/>
        </physiologicalReaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="2">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <comment type="pathway">
        <text evidence="4 20">Cofactor biosynthesis; NAD(+) biosynthesis; NAD(+) from nicotinamide D-ribonucleotide: step 1/1.</text>
    </comment>
    <comment type="pathway">
        <text evidence="5">Cofactor biosynthesis; NAD(+) biosynthesis; deamido-NAD(+) from nicotinate D-ribonucleotide: step 1/1.</text>
    </comment>
    <comment type="subunit">
        <text evidence="19">Homohexamer. Interacts with ADPRT/PARP1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="6 20">Belongs to the eukaryotic NMN adenylyltransferase family.</text>
    </comment>
    <dbReference type="EC" id="2.7.7.1" evidence="20"/>
    <dbReference type="EC" id="2.7.7.18" evidence="20"/>
    <dbReference type="AlphaFoldDB" id="A0A452HNE7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018804.1">
        <property type="protein sequence ID" value="ENSGAGP00000016476.1"/>
        <property type="gene ID" value="ENSGAGG00000012328.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00253">
        <property type="reaction ID" value="UER00332"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000309">
        <property type="term" value="F:nicotinamide-nucleotide adenylyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004515">
        <property type="term" value="F:nicotinate-nucleotide adenylyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009435">
        <property type="term" value="P:NAD+ biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="CDD" id="cd09286">
        <property type="entry name" value="NMNAT_Eukarya"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.620:FF:000101">
        <property type="entry name" value="Nicotinamide-nucleotide adenylyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.620">
        <property type="entry name" value="HUPs"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004821">
        <property type="entry name" value="Cyt_trans-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051182">
        <property type="entry name" value="Euk_NMN_adenylyltrnsfrase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005248">
        <property type="entry name" value="NadD/NMNAT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045094">
        <property type="entry name" value="NMNAT_euk"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014729">
        <property type="entry name" value="Rossmann-like_a/b/a_fold"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00482">
        <property type="entry name" value="nicotinate (nicotinamide) nucleotide adenylyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12039">
        <property type="entry name" value="NICOTINAMIDE MONONUCLEOTIDE ADENYLYLTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12039:SF21">
        <property type="entry name" value="NICOTINAMIDE_NICOTINIC ACID MONONUCLEOTIDE ADENYLYLTRANSFERASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01467">
        <property type="entry name" value="CTP_transf_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52374">
        <property type="entry name" value="Nucleotidylyl transferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="13 20" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="14" id="KW-0460">Magnesium</keyword>
    <keyword evidence="15 20" id="KW-0520">NAD</keyword>
    <keyword evidence="11 20" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="10 20" id="KW-0548">Nucleotidyltransferase</keyword>
    <keyword evidence="16" id="KW-0539">Nucleus</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="8 20" id="KW-0662">Pyridine nucleotide biosynthesis</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9 20" id="KW-0808">Transferase</keyword>
    <keyword evidence="12" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="Cytidyltransferase-like" evidence="21">
        <location>
            <begin position="13"/>
            <end position="231"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004658"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00005019"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00007064"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022642"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022695"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023027"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00048514"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00048969"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00064648"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="RuleBase" id="RU362021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="Pfam" id="PF01467"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016476.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="286" mass="32593" checksum="8B55371C86232156" modified="2019-05-08" version="1">MENPDNKTEVVLLACGSFNPITNMHLRLFELAKDYLHETGKYNVIKGIISPVGDAYKKKSLISASHRVTMAKLATESSDWVQVDDWESSQNEWLETLKVLRYHHQKLASSTPANSLQNAVPLIKPGRKRKQEANRYIPAKKYQLSPETKGVPQLKLLCGADILESFGIPNLWKLEDITEIVANHGLVCITRAGNNAQKFIYESDMLWQHKNNIHLVEEWITNDISSTKIRRALRRGQSIRYLVPDAVRAYIEKHNLYSPESEDRNTGVILAPLQKHARGCKQEQTL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNE8</accession>
    <name>A0A452HNE8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Enoyl-[acyl-carrier-protein] reductase, mitochondrial</fullName>
            <ecNumber evidence="11">1.3.1.104</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="13">2-enoyl thioester reductase</fullName>
        </alternativeName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Mitochondrion</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the zinc-containing alcohol dehydrogenase family. Quinone oxidoreductase subfamily.</text>
    </comment>
    <dbReference type="EC" id="1.3.1.104" evidence="11"/>
    <dbReference type="AlphaFoldDB" id="A0A452HNE8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018859.1">
        <property type="protein sequence ID" value="ENSGAGP00000016524.1"/>
        <property type="gene ID" value="ENSGAGG00000012357.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0141148">
        <property type="term" value="F:enoyl-[acyl-carrier-protein] reductase (NADPH) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006633">
        <property type="term" value="P:fatty acid biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd08290">
        <property type="entry name" value="ETR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.720:FF:000112">
        <property type="entry name" value="Enoyl-[acyl-carrier-protein] reductase 1, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.180.10:FF:000010">
        <property type="entry name" value="Enoyl-[acyl-carrier-protein] reductase, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.180.10">
        <property type="entry name" value="Medium-chain alcohol dehydrogenases, catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.720">
        <property type="entry name" value="NAD(P)-binding Rossmann-like Domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013149">
        <property type="entry name" value="ADH-like_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013154">
        <property type="entry name" value="ADH-like_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020843">
        <property type="entry name" value="ER"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011032">
        <property type="entry name" value="GroES-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051034">
        <property type="entry name" value="Mito_Enoyl-ACP_Reductase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036291">
        <property type="entry name" value="NAD(P)-bd_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43981">
        <property type="entry name" value="ENOYL-[ACYL-CARRIER-PROTEIN] REDUCTASE, MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43981:SF9">
        <property type="entry name" value="ENOYL-[ACYL-CARRIER-PROTEIN] REDUCTASE, MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08240">
        <property type="entry name" value="ADH_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00107">
        <property type="entry name" value="ADH_zinc_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00829">
        <property type="entry name" value="PKS_ER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50129">
        <property type="entry name" value="GroES-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51735">
        <property type="entry name" value="NAD(P)-binding Rossmann-fold domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0275">Fatty acid biosynthesis</keyword>
    <keyword evidence="4" id="KW-0276">Fatty acid metabolism</keyword>
    <keyword evidence="3" id="KW-0444">Lipid biosynthesis</keyword>
    <keyword evidence="8" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="9" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="5" id="KW-0521">NADP</keyword>
    <keyword evidence="7" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0809">Transit peptide</keyword>
    <feature type="domain" description="Enoyl reductase (ER)" evidence="14">
        <location>
            <begin position="3"/>
            <end position="229"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004173"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010371"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022516"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022832"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022857"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023160"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00038963"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00041058"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00042123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="SMART" id="SM00829"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016524.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="276" mass="29907" checksum="5990EAD0F1F0AA1C" modified="2019-05-08" version="1">MLSILKDLERAELGHSGVHVKMLAAPINPADINMIQGTYALLPDLPAVGGNEGVGHVVEVGSHVTSLKPGDWVIPADAELGTWRTDAVFSEETLVKLPRDIPLLCAATLSVNPCTAYRMLADFETLGPGDSIIQNAANSGVGQAVIQIAAALGIKTINVIRDRPNLQEVVDRLTSLGADHVLTEEMLRKPEMKDLFKKVPKPRLALNCVGGKSATEMLRHIQQKGTMVTYGGMAKQPVTVPVSAFIFKDVKLRGFWMTQWKKDQTQSKLKVIPVGM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNF1</accession>
    <name>A0A452HNF1_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="5">GTP dependent ribosome recycling factor mitochondrial 2</fullName>
        </submittedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HNF1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018837.1">
        <property type="protein sequence ID" value="ENSGAGP00000016505.1"/>
        <property type="gene ID" value="ENSGAGG00000012347.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005525">
        <property type="term" value="F:GTP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003924">
        <property type="term" value="F:GTPase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032543">
        <property type="term" value="P:mitochondrial translation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032790">
        <property type="term" value="P:ribosome disassembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000795">
        <property type="entry name" value="T_Tr_GTP-bd_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43261:SF1">
        <property type="entry name" value="RIBOSOME-RELEASING FACTOR 2, MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43261">
        <property type="entry name" value="TRANSLATION ELONGATION FACTOR G-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00009">
        <property type="entry name" value="GTP_EFTU"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0342">GTP-binding</keyword>
    <keyword evidence="1" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="2" id="KW-0648">Protein biosynthesis</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Tr-type G" evidence="4">
        <location>
            <begin position="66"/>
            <end position="101"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022917"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="Pfam" id="PF00009"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016505.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="120" mass="13655" checksum="D0DEB497A95DD4ED" modified="2019-05-08" version="1">MLRVPRKFSVTVLKVSSLCSECIYCRWARTMVVKRLKSQECHQRNYSFPPGDVKSLRSIITPPISKIRNIGIMAHIDAGKTTTTERMLYYSGYTRALGECSSQTQMEVLARSSILLQHAK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNF3</accession>
    <name>A0A452HNF3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Oxysterol-binding protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="2">Belongs to the OSBP family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HNF3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018846.1">
        <property type="protein sequence ID" value="ENSGAGP00000016514.1"/>
        <property type="gene ID" value="ENSGAGG00000012290.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097038">
        <property type="term" value="C:perinuclear endoplasmic reticulum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015485">
        <property type="term" value="F:cholesterol binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006869">
        <property type="term" value="P:lipid transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.160.120">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037239">
        <property type="entry name" value="OSBP_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000648">
        <property type="entry name" value="Oxysterol-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018494">
        <property type="entry name" value="Oxysterol-bd_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10972">
        <property type="entry name" value="OXYSTEROL-BINDING PROTEIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10972:SF153">
        <property type="entry name" value="OXYSTEROL-BINDING PROTEIN-RELATED PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01237">
        <property type="entry name" value="Oxysterol_BP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF144000">
        <property type="entry name" value="Oxysterol-binding protein-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01013">
        <property type="entry name" value="OSBP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0445">Lipid transport</keyword>
    <keyword evidence="1" id="KW-0446">Lipid-binding</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00023121"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="RuleBase" id="RU003844"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="RuleBase" id="RU003845"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016514.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="232" mass="26400" checksum="9F6C6404DF4D54FC" modified="2019-05-08" version="1">MLSHKVLEKVTDLYFLYTGFDSDNSSGDFSETNHKVAEMLDLDPPQSNRTGKHNGARQLHENGIKKHRTSLPAPMFSRSDFSVWSILKKCIGLELSKITMPIVFNEPLSFLQRITEYMEHIYLIHKASSHANPLERMQAVAAFAVSAVASQWERTGKPFNPLLGETYELIREDLGFRFISEQVSHHPPVSAFYSEGLNKDFVFHGSIYPKLKFWGKSVEAEPRGTITLELLK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNF4</accession>
    <name>A0A452HNF4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">BAT2 N-terminal domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000018864.1">
        <property type="protein sequence ID" value="ENSGAGP00000016529.1"/>
        <property type="gene ID" value="ENSGAGG00000012361.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009738">
        <property type="entry name" value="BAT2_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033184">
        <property type="entry name" value="PRRC2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14038">
        <property type="entry name" value="BAT2 HLA-B-ASSOCIATED TRANSCRIPT 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14038:SF4">
        <property type="entry name" value="PROTEIN PRRC2B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07001">
        <property type="entry name" value="BAT2_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="1" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="BAT2 N-terminal" evidence="4">
        <location>
            <begin position="1"/>
            <end position="189"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="50"/>
            <end position="469"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="542"/>
            <end position="564"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="594"/>
            <end position="660"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="717"/>
            <end position="747"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="876"/>
            <end position="1057"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="1090"/>
            <end position="1247"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="1271"/>
            <end position="1527"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="1627"/>
            <end position="1673"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="1726"/>
            <end position="1755"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="1978"/>
            <end position="2103"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="2">
        <location>
            <begin position="496"/>
            <end position="523"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="88"/>
            <end position="137"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="179"/>
            <end position="190"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="233"/>
            <end position="247"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="3">
        <location>
            <begin position="303"/>
            <end position="318"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="375"/>
            <end position="388"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="423"/>
            <end position="434"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="441"/>
            <end position="456"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="544"/>
            <end position="564"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="612"/>
            <end position="621"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="3">
        <location>
            <begin position="629"/>
            <end position="639"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="717"/>
            <end position="735"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="876"/>
            <end position="885"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="928"/>
            <end position="939"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="956"/>
            <end position="990"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="3">
        <location>
            <begin position="1024"/>
            <end position="1033"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="1034"/>
            <end position="1048"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="1158"/>
            <end position="1192"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="1222"/>
            <end position="1235"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="1271"/>
            <end position="1283"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="1305"/>
            <end position="1322"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="1382"/>
            <end position="1392"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="1425"/>
            <end position="1434"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="1502"/>
            <end position="1522"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="1627"/>
            <end position="1642"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="1730"/>
            <end position="1740"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="1990"/>
            <end position="2003"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="2013"/>
            <end position="2022"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="3">
        <location>
            <begin position="2031"/>
            <end position="2044"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="Pfam" id="PF07001"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016529.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2103" mass="233114" checksum="CF543FB76EB8ED07" modified="2019-05-08" version="1">MSDRLGQITKGKDGKSKYSTLSLFDKYKGKSIEAIRTTVIPRHGLQSLGKVAAARRMPPPANLPSLKSENKGNDPNIIIVPKDGTGWANKQDQPDQKSSSATTAQPQESLPQQGLQKSVSNLQKPTQSISQESTNSVPGGPKSWAQLNGKPAGQEGGSRVSSRLLSFSPEEFPTLKAAGEQDKVGKEKGALDLSYGPGPSLRPQNVTSWREGGGRNIISATSLTASPPEPGIKTSSTGDGDPSSVSASDPKEPSLRPAQPIRKGASQFMGNAYHPPTYHDMLPAFMCPQQSSEVPGSLDRGSFPLPQLRLQPRVPFRQYQMNDQDGKENRPGVSRPTRPVRQQGERAPRPTIINAENLKELDELDTDADDGWAGIHDEVDYSEKLKFSEDEEEEEALKDGRQKWNTWDPRRQRQLSLSSADSADIKHTPEEGKNWGDQVGLSRSVSKVQGPQQPSRKLNGWTPASEYQKPPLGCILRQQSLEDKEEKVPLRQKFVHSEISEAVERARRRREEEERRAREERLAACAAKLKQLDQKCKLAQKTGETQKHVENEDLKPPSVEKKVAQENGHTFRRATSEFHAQDVSVGYLEEETTATTVAAQNSSEEEIREATSPAQEFNKYQKSLPPRFQRQQQQQQQEQLYKMQHWQQQQVYPPPSHSHPQRTFYPPHPQMLGFDPRWMMMPSYMDPRMAQGRTPVDFYPSALYPSGIMKPMIQQDSINGSGCRSEEQNCQPSLQQERKAASMDPAPVWGQESYTSLQNKGYPLPHQKQTENMSMEGLHTRSDSSYSASSGRSESINVQRNLFEDRGDEYLNTFDKKAQSDFDSSISSQRIGQDLLFQHQETVQDACVSGARHASLRCSPLESDFIQVDKKPEYNGWDINHHQKSSEPAAEADEEVPRDEQSFSADPWKKEGAVSKQPAEETDWAPENRNTNNQHQEQMGRTRRSGPIKKPVLKALKVEEKEKEQEKIKLEGEELSHPAKERAPVYKVENDSEDSSTLLNSTHYLLDEKGSSQANFTQDAEKSQEEEEEEEEEKAERAWENKLSRESNDLPPTKRNNWIFIDEEQAFGGRGQGRGRGRGFREFTFRGRGTAVSGRGVYNNQRSSRGRGLREFNQLEDFPRGKPRRRIASETHSEGSEYEELPKRRRQRGSENSNESSLLEREESDLKKGDFKDSWRSNKIYSDDHSSLDAKMRAPRAFGRSLPPRLSNSGYGRRGFMAKEPSQWQGRSGGTTWQEYSHCPPSDTFGIRRQSDRDYIQDSYKHVDSFTGRGFEENHIDNRRPLFQEDYTADQENIENRPFRRRRPPRQDKPPRFRRLRQERESVGQWNPEEGGTSLLPSQWRPKLTSVEKNGVSGRRSPELSYQNSSDHANEEWETASESSDFSERRERRDGVLESEAQLDGGLASGSLGEKRELAKRSFSSQRPLVDRQSRKAEPAGFGEQLVRAGGGAASRFESQQNGMPMKNKRSPEEGGGLGNAKSGSSHSVYSLERSLCTSSDSAEGPGKKQEKEPKSTAQRPSEKGETMSQFEVNYGNAIIDHRVSSTAEENEVGSMAGEGFIEVLTKKQRRLLEEERRKKEQAAQGPTKGRVLQSRIPPRFAKKQNSMCLEQGDVTVSGSSLGTEIWESNSPALSVQSPGSDSWSKPVNAFNGTESSSTEDSDFSLPPGSASGTAASPVTKLQDALASNAGLTQSIPILRRDHHLQRCIGLNPMSFPTADLTLKMESARKAWENSPSLPEQNSPGGAGSGIQPPSSVGASSGVSYSSFGGVSMPPMPVASVAPSASIPGNHIPPLYLDGHVFASQPRLVPQTIPQQQSYQQAAAAQQIPISLHTSLQAQAQLGLRGGLPVSQSQEIYSSIQPFRSPVYMHPSLSQPSTMVLTGGAALKPPYSAFPGMQPLEMVKTQSGSPYQPMSGSQTLVYEGQINQAASLGASQMMDSQLTQLTMPMPGSQLPLPRYGSGQQPLILPQSIQLPQGQNLSVGAPRRILPPGSQPSVLTTSRESSQMEMKGFHFTDGKQNMSSGGSVPSPHTYRPSSASPSGKPSGPAVNMGSVQGHYVQQAKQRVDENKTNLGAVKLQEPSSANQMKPVRTGAIKPQAVKVEESKA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNF7</accession>
    <name>A0A452HNF7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Kv channel-interacting protein 4</fullName>
        </recommendedName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Peripheral membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Peroxisome</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the recoverin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HNF7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018815.1">
        <property type="protein sequence ID" value="ENSGAGP00000016486.1"/>
        <property type="gene ID" value="ENSGAGG00000012335.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005777">
        <property type="term" value="C:peroxisome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008076">
        <property type="term" value="C:voltage-gated potassium channel complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005267">
        <property type="term" value="F:potassium channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015459">
        <property type="term" value="F:potassium channel regulator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1901379">
        <property type="term" value="P:regulation of potassium ion transmembrane transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00051">
        <property type="entry name" value="EFh"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.238.10:FF:000043">
        <property type="entry name" value="Kv channel-interacting protein 1 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.238.10">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011992">
        <property type="entry name" value="EF-hand-dom_pair"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018247">
        <property type="entry name" value="EF_Hand_1_Ca_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002048">
        <property type="entry name" value="EF_hand_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028846">
        <property type="entry name" value="Recoverin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23055">
        <property type="entry name" value="CALCIUM BINDING PROTEINS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23055:SF30">
        <property type="entry name" value="KV CHANNEL-INTERACTING PROTEIN 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13499">
        <property type="entry name" value="EF-hand_7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13833">
        <property type="entry name" value="EF-hand_8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00450">
        <property type="entry name" value="RECOVERIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00054">
        <property type="entry name" value="EFh"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47473">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00018">
        <property type="entry name" value="EF_HAND_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50222">
        <property type="entry name" value="EF_HAND_2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="13" id="KW-0106">Calcium</keyword>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="19" id="KW-0407">Ion channel</keyword>
    <keyword evidence="16" id="KW-0406">Ion transport</keyword>
    <keyword evidence="17" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="18" id="KW-0576">Peroxisome</keyword>
    <keyword evidence="9" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="15" id="KW-0630">Potassium</keyword>
    <keyword evidence="12" id="KW-0631">Potassium channel</keyword>
    <keyword evidence="8" id="KW-0633">Potassium transport</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="11" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0813">Transport</keyword>
    <keyword evidence="14" id="KW-0851">Voltage-gated channel</keyword>
    <feature type="domain" description="EF-hand" evidence="22">
        <location>
            <begin position="86"/>
            <end position="121"/>
        </location>
    </feature>
    <feature type="domain" description="EF-hand" evidence="22">
        <location>
            <begin position="122"/>
            <end position="157"/>
        </location>
    </feature>
    <feature type="domain" description="EF-hand" evidence="22">
        <location>
            <begin position="170"/>
            <end position="205"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="21">
        <location>
            <begin position="1"/>
            <end position="24"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="21">
        <location>
            <begin position="9"/>
            <end position="21"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004202"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004275"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00006049"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022538"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022826"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022882"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022958"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023140"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00040738"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS50222"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016486.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="216" mass="24960" checksum="59ACC8268E6094EE" modified="2019-05-08" version="1">MNARRVESISAQLEEASSTGDSVEDEMEIATVRHRPEALELLEAQSKFTKKELQILYRGFKNECPSGIVNEETFKEIYSQFFPQGDSTTYAHFLFNAFDTDHNGSVSFEDFIMGLSILLRGTVQEKLNWAFNLYDINKDGYITKEEMLDIMKAIYDMMGKCTYPVLKEDAPRQHVETFFQKMDKNKDGVVTIDEFIESCQKDENIMCSMQLFENVI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNF9</accession>
    <name>A0A452HNF9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="15">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cell projection</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018869.1">
        <property type="protein sequence ID" value="ENSGAGP00000016534.1"/>
        <property type="gene ID" value="ENSGAGG00000012280.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042995">
        <property type="term" value="C:cell projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="CDD" id="cd00603">
        <property type="entry name" value="IPT_PCSR"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.160.20.10">
        <property type="entry name" value="Single-stranded right-handed beta-helix, Pectin lyase-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055401">
        <property type="entry name" value="CEMIP_beta-hel_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052387">
        <property type="entry name" value="Fibrocystin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019316">
        <property type="entry name" value="G8_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014756">
        <property type="entry name" value="Ig_E-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002909">
        <property type="entry name" value="IPT_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037524">
        <property type="entry name" value="PA14/GLEYA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006626">
        <property type="entry name" value="PbH1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012334">
        <property type="entry name" value="Pectin_lyas_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011050">
        <property type="entry name" value="Pectin_lyase_fold/virulence"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46769:SF1">
        <property type="entry name" value="FIBROCYSTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46769">
        <property type="entry name" value="POLYCYSTIC KIDNEY AND HEPATIC DISEASE 1 (AUTOSOMAL RECESSIVE)-LIKE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24606">
        <property type="entry name" value="CEMIP_beta-hel"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10162">
        <property type="entry name" value="G8"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01833">
        <property type="entry name" value="TIG"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SMART" id="SM01225">
        <property type="entry name" value="G8"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00429">
        <property type="entry name" value="IPT"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00710">
        <property type="entry name" value="PbH1"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56988">
        <property type="entry name" value="Anthrax protective antigen"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81296">
        <property type="entry name" value="E set domains"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51126">
        <property type="entry name" value="Pectin lyase-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51484">
        <property type="entry name" value="G8"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51820">
        <property type="entry name" value="PA14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11" id="KW-0966">Cell projection</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9 12" id="KW-0472">Membrane</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="5 12" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 12" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="3605"/>
            <end position="3628"/>
        </location>
    </feature>
    <feature type="domain" description="PA14" evidence="14">
        <location>
            <begin position="141"/>
            <end position="300"/>
        </location>
    </feature>
    <feature type="domain" description="G8" evidence="13">
        <location>
            <begin position="1714"/>
            <end position="1835"/>
        </location>
    </feature>
    <feature type="domain" description="G8" evidence="13">
        <location>
            <begin position="2502"/>
            <end position="2628"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004167"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004316"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS51484"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS51820"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016534.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="3658" mass="399613" checksum="ECB823CE39E2D527" modified="2019-05-08" version="1">SAGGSCSICYPRITVFLCFILRCPIQLPWYYGEIYPQLKFHGVSQAFLSCCRSMVHKDAWLISAKQEVFLYQTFSDIASVFPVSGSLGGGSDLTITGDFFDQPVQVTVAGVPCQIKHCSPRKISCITGPIAEIRRLAAPQAGNRGLLFEVWDDAAGTDLTETAPGYRWQIVPNASSPERFLSQAQQSFSSRLRGFFVAPQTNNYTFWIQADGRASLHLSLSEDPGKKVKVAFIPAGASEWSEHWEKNWSKSWQPKSQKFELMGGSKYYLEALHHGKAPSSGMRVGVQLHNTWLNPDVVNTYRRERHEIWASALRLPEIQMLTLSGTGWFRISWDNVSSNMMATNATADQVQTVIEELLSVKCETEPLSAKFLLRNGFEEGNVCFAYKGYVSDTLYVSVSYINTFLNTVKKNLTCLWSLNETSPESWKFTCADLWRCCVNGSESLRDLEANSSVFVHQIDMLIPEREEETSSWFYLDEIIVSDRNIAVFQKDFKPARPGGNVIQEVSVVGSPPTYNVSFLVAGCGSNLPLLGALPSQGSEEDHLAAFMEGANVSMTVQRVQAASPPIGGTFRILLPNTVISGVPVHVSSHHLRKLLQSNSDNSTGQYFNASDFTVTKDVSTCYQDVWTLTWNSMTGDLPIVIDVSAENLTGLNPTIAARVVYDGGVFIGPIFGDMLATANNHTQVVVVVNDVPANCSGSCSFQYSQEATPLVSHVDYSLGNVYITGSGFAEDSQALQIEVNHTTCQVMTSNQTAVVCWMNMLPMGLYHVTLLVRPYGFARNASEGQGIYLRVKPRLEAIEPPTTSEIGGCSVILRGTGFDGISQVLFGSHPCPINTNASNSVRIECEVPSRGDEEYVVHVTLTAGSQSATFTEVFKYDPSLNPAIVSLSRNRSSTAGLQSLRIGISSFANCTGLDVQVKIEATVAQIQARTAHGVDVVLPALADGWYNLSVIINGIAIGSHGVDPFIQYSTEILSIEPCCGSVLGGTILTISGKGFSQNPSLVSVFIGTQTCDVTRSMEETIWCQTPPAADFSNIASQDIPVLVGVFIGNRSSPLLPSWSFPKRNITFTYQMALTPIVTSMEVKMMNDSLCLGIEGINITGSVAKLGDSECELEFQCANESTTLSQCSFPLSTLEPGIYPIRFLQKQLGYANVTAALQNFTLLPEVKAIIPSRGSACGGLLLTISGLGLKSRRNSVHVSLTGNYTCEIRSVDHNRISCALLPREPPLGDWWLPDASHALNVTVTVNGIQSVCLSDCALHLLKEWTPRVDAVVSEINGTLSSLLIRGQRLALATDELVIQVDNHDPCNVTFWNETSIQCQTGSLSPGEHSLSVSNRRSGHACFSRDSRIFTVIPCVLKFYPQNFSTNGGGLLAIQGAALKGRNSTSILVGNRPCLITDVSYVVLRCSVPLGNGIQDLTLEIDGISYHLGEISYSEEFTPIFLFLLPPVGLLLTMTVSRITGVESMHISIGDSPCTNITGNHTTLQCSCSAPWLPAGEYRVLGGDFLRGWASSNLTFTSELTVISTRNNWGKCGLGGGEVHVHGTGFAPGNTSVTICGAPCEMLGDITTTDVSCLAPRLDASLAVLCGLTHPSEDCKGAGTTYVQCDVHITAGAYSLAGARPYIYLCDDDSLLDSKTKVNSPQALFAGLFFSPKVERDEVLIYNSSCNITMETEAEMECEGANQPITAKITEIRKNWGQNTQRHLPLQFCGRWSKNSSWLDGHLPQDGANVTVERGQTLLLDTTTGILNLLHVKGGKLLLAGPGPIEIHAHYILVTDGGEFQVGSPDKPLRDKAIIHLYGSFHSATFFPYGAKFLAVRNGTVSMHGWVPQVVSTHLKSAACVNDSRIFLEEPVDWQPGDEVIVSGTGPGDTYQQEEIVTIEAVNNTELYLRSPLRFPHSSGEEWVHGQRFALKAVVALLSRRIVVRGNLTRERMSHLRACAEAGLRKGNENASACPYKRSERMLGSRDLGAVIIVQAFQGEDSHIHLEGVQFQNMGQAFQQHLSALTIAGDAHMTDSYIRGCSVWDSFGRGLSLSRTSELRVDNNVFYNIFGHGILLGESLEQGNRVRHNTVIGLSGTDGLSNIETLSPAGIYIRAPANQIEANTVCAAGYGYFFHLSPEGPSQTPLLTFSKNTAHSCTRYGLLVYPKYQPKCEDGLGPVLFQSFTAWRNQGGAQIFSSRNLELRNFQIYACKDFGIDIVESLGNTTVANSLLLGHFGQKDSSCMSSGLKTPQRHELLVSNTSFMNFDISTCTAISTCSGCYQGQGGFTVRAERLAFLNSPNQASFPFPHSAILEDLDGSLTEQEGSHLLASLDTLGASCVASANFSHAARGSVCGRDVIFHRMSIEAPDFPYNLTVTDSNNKTTTVYYVSDTLSNPYGWMALLLDQETYTLTFGNPFVSKQLQYSATFDNFIAGNYLLVEHENLPSCVEVMVFCGTRPGQPLQSLPSHDHDKGCDWFFNSKLRKLTYLGKCNASNIGNVVQMSISPPDDILKWSFPESWSGVKKGWGGYNHSIPAPGEDVIILPNRSILVDTTLPPLRSLYILGTLEFPVNSSNVLSVTCIVVAGGELKVGTFQHPLERGMKLLILLRASEGVYCDRLDGINVHPGTIGVYGKVQIHSAYPRKSWTHLGSDIAPGNERIWVEDEVDWSPGGNIVISSSSYDAHQAEVVTLKEINGHSIRFQERLFHRHIGRSHSIEDGRRVPLAAEVGLLTRNIQIKSDTACSGRLLVGRFRNTSGTEFAGDLRLSNIEILNFGPSQLPAIGFTNISLGSSIISSSIHHSCGGGIQAVTSSGISLRDNVVFNTVGPGIDLEGQTHSLIRNLIILSKQPKGLPDWVAGIKVNKIDDSYLLGNVMAGSERIAFHIRGQECFLAGEYFAENVAHSSLHGVHLYRGDGFQNCTKITGFLSYKNYDYGVMFQLESSVVMDNMMLVDNAVGVLPVVYSPSAKQHRLGKKYIELRNCVIIATSSTFDCIKDRITPLSANMTSRDRAPYSPQRGRTGILWPSFASEPSQWPNNPWHKIRNYAAVSGIMKLQEVTFTGFTKSCYSDDRDICIMSNPDSVGIMYPITAEMTRMSHVHEQNKFYFHTLETSINGSKDFEYLEMRCEGSRKALFKDLDGSALGLQPPVSVFPKSKSEWTESCLHAGIYREDTKCIYKPSTQGYFCKDTDHALVIIESLDTEADGRRLSPVVSVTGSFVEAFSSGVSHVSCCPSEHPRAFYSVLPSNKLTKVCFAGLTPPSLRLRLISGQNSTRLLIAIFYDEPRSLHVFMNGNYIPLASSSSSVFLTNAMAGTHSFSFENNLLYILLHGDDPIEIYTSFSLHTAFTIAETIGEEGQRRVVHRLADFLQIGRDHVRIVQSVSGSENTLKVISDNVGKRRRECPPMTSCTASCSRAGWEKTRTGAASVWILQSSHAATPLRVLIIEIGEPPGLFRNKFFPSLSSDRLKSLASTVINAQGYLGNCLGIGIGCFKIILNNMMAFFFPHCCLYVRPYNISVLVQPSDGEMEQELPVQPQIIFLDKQGRRVETLGLPSEPWVMTACLKGSSEAVLKGHTRVEVRDGRAIFTNLAISNSGSNWCLVFTVISPPGADFTVESQPFTIFPIPKGGKSITVLAAVLGSAASVLVMGLLVFCWSKKSKSFSKWESPGELSICRVSPISLVSKE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNH4</accession>
    <name>A0A452HNH4_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="15">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cell projection</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018867.1">
        <property type="protein sequence ID" value="ENSGAGP00000016532.1"/>
        <property type="gene ID" value="ENSGAGG00000012280.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042995">
        <property type="term" value="C:cell projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="CDD" id="cd00603">
        <property type="entry name" value="IPT_PCSR"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.160.20.10">
        <property type="entry name" value="Single-stranded right-handed beta-helix, Pectin lyase-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055401">
        <property type="entry name" value="CEMIP_beta-hel_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052387">
        <property type="entry name" value="Fibrocystin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019316">
        <property type="entry name" value="G8_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014756">
        <property type="entry name" value="Ig_E-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002909">
        <property type="entry name" value="IPT_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037524">
        <property type="entry name" value="PA14/GLEYA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006626">
        <property type="entry name" value="PbH1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012334">
        <property type="entry name" value="Pectin_lyas_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011050">
        <property type="entry name" value="Pectin_lyase_fold/virulence"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46769:SF1">
        <property type="entry name" value="FIBROCYSTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46769">
        <property type="entry name" value="POLYCYSTIC KIDNEY AND HEPATIC DISEASE 1 (AUTOSOMAL RECESSIVE)-LIKE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24606">
        <property type="entry name" value="CEMIP_beta-hel"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10162">
        <property type="entry name" value="G8"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01833">
        <property type="entry name" value="TIG"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SMART" id="SM01225">
        <property type="entry name" value="G8"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00429">
        <property type="entry name" value="IPT"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00710">
        <property type="entry name" value="PbH1"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56988">
        <property type="entry name" value="Anthrax protective antigen"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81296">
        <property type="entry name" value="E set domains"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51126">
        <property type="entry name" value="Pectin lyase-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51484">
        <property type="entry name" value="G8"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51820">
        <property type="entry name" value="PA14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11" id="KW-0966">Cell projection</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9 12" id="KW-0472">Membrane</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="5 12" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 12" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="3619"/>
            <end position="3642"/>
        </location>
    </feature>
    <feature type="domain" description="PA14" evidence="14">
        <location>
            <begin position="147"/>
            <end position="306"/>
        </location>
    </feature>
    <feature type="domain" description="G8" evidence="13">
        <location>
            <begin position="1720"/>
            <end position="1841"/>
        </location>
    </feature>
    <feature type="domain" description="G8" evidence="13">
        <location>
            <begin position="2516"/>
            <end position="2642"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004167"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004316"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS51484"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS51820"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016532.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="3672" mass="401539" checksum="EF32659D9A228132" modified="2019-05-08" version="1">EGDLQRSENRHYSICYPRITVFLCFILRCPIQLPWYYGEIYPQLKFHGVSQAFLSCCRSMVHKDAWLISAKQEVFLYQTFSDIASVFPVSGSLGGGSDLTITGDFFDQPVQVTVAGVPCQIKHCSPRKISCITGPIAEIRRLAAPQAGNRGLLFEVWDDAAGTDLTETAPGYRWQIVPNASSPERFLSQAQQSFSSRLRGFFVAPQTNNYTFWIQADGRASLHLSLSEDPGKKVKVAFIPAGASEWSEHWEKNWSKSWQPKSQKFELMGGSKYYLEALHHGKAPSSGMRVGVQLHNTWLNPDVVNTYRRERHEIWASALRLPEIQMLTLSGTGWFRISWDNVSSNMMATNATADQVQTVIEELLSVKCETEPLSAKFLLRNGFEEGNVCFAYKGYVSDTLYVSVSYINTFLNTVKKNLTCLWSLNETSPESWKFTCADLWRCCVNGSESLRDLEANSSVFVHQIDMLIPEREEETSSWFYLDEIIVSDRNIAVFQKDFKPARPGGNVIQEVSVVGSPPTYNVSFLVAGCGSNLPLLGALPSQGSEEDHLAAFMEGANVSMTVQRVQAASPPIGGTFRILLPNTVISGVPVHVSSHHLRKLLQSNSDNSTGQYFNASDFTVTKDVSTCYQDVWTLTWNSMTGDLPIVIDVSAENLTGLNPTIAARVVYDGGVFIGPIFGDMLATANNHTQVVVVVNDVPANCSGSCSFQYSQEATPLVSHVDYSLGNVYITGSGFAEDSQALQIEVNHTTCQVMTSNQTAVVCWMNMLPMGLYHVTLLVRPYGFARNASEGQGIYLRVKPRLEAIEPPTTSEIGGCSVILRGTGFDGISQVLFGSHPCPINTNASNSVRIECEVPSRGDEEYVVHVTLTAGSQSATFTEVFKYDPSLNPAIVSLSRNRSSTAGLQSLRIGISSFANCTGLDVQVKIEATVAQIQARTAHGVDVVLPALADGWYNLSVIINGIAIGSHGVDPFIQYSTEILSIEPCCGSVLGGTILTISGKGFSQNPSLVSVFIGTQTCDVTRSMEETIWCQTPPAADFSNIASQDIPVLVGVFIGNRSSPLLPSWSFPKRNITFTYQMALTPIVTSMEVKMMNDSLCLGIEGINITGSVAKLGDSECELEFQCANESTTLSQCSFPLSTLEPGIYPIRFLQKQLGYANVTAALQNFTLLPEVKAIIPSRGSACGGLLLTISGLGLKSRRNSVHVSLTGNYTCEIRSVDHNRISCALLPREPPLGDWWLPDASHALNVTVTVNGIQSVCLSDCALHLLKEWTPRVDAVVSEINGTLSSLLIRGQRLALATDELVIQVDNHDPCNVTFWNETSIQCQTGSLSPGEHSLSVSNRRSGHACFSRDSRIFTVIPCVLKFYPQNFSTNGGGLLAIQGAALKGRNSTSILVGNRPCLITDVSYVVLRCSVPLGNGIQDLTLEIDGISYHLGEISYSEEFTPIFLFLLPPVGLLLTMTVSRITGVESMHISIGDSPCTNITGNHTTLQCSCSAPWLPAGEYRVLGGDFLRGWASSNLTFTSELTVISTRNNWGKCGLGGGEVHVHGTGFAPGNTSVTICGAPCEMLGDITTTDVSCLAPRLDASLAVLCGLTHPSEDCKGAGTTYVQCDVHITAGAYSLAGARPYIYLCDDDSLLDSKTKVNSPQALFAGLFFSPKVERDEVLIYNSSCNITMETEAEMECEGANQPITAKITEIRKNWGQNTQRHLPLQFCGRWSKNSSWLDGHLPQDGANVTVERGQTLLLDTTTGILNLLHVKGGKLLLAGPGPIEIHAHYILVTDGGEFQVGSPDKPLRDKAIIHLYGSFHSATFFPYGAKFLAVRNGTVSMHGWVPQVVSTHLKSAACVNDSRIFLEEPVDWQPGDEVIVSGTGPGDTYQQEEIVTIEAVNNTELYLRSPLRFPHSSGEEWVHGQRFALKAVVALLSRRIVVRGNLTRERMSHLRACAEAGLRKGNENASACPYKRSERMLGSRDLGAVIIVQAFQGEDSHIHLEGVQFQNMGQAFQQHLSALTIAGDAHMTDSYIRGCSVWDSFGRGLSLSRTSELRVDNNVFYNIFGHGILLGKLVMSPFTKSLEQGNRVRHNTVIGLSGTDGLSNIETLSPAGIYIRAPANQIEANTVCAAGYGYFFHLSPEGPSQTPLLTFSKNTAHSCTRYGLLVYPKYQPKCEDGLGPVLFQSFTAWRNQGGAQIFSSRNLELRNFQIYACKDFGIDIVESLGNTTVANSLLLGHFGQKDSSCMSSGLKTPQRHELLVSNTSFMNFDISTCTAISTCSGCYQGQGGFTVRAERLAFLNSPNQASFPFPHSAILEDLDGSLTEQEGSHLLASLDTLGASCVASANFSHAARGSVCGRDVIFHRMSIEAPDFPYNLTVTDSNNKTTTVYYVSDTLSNPYGWMALLLDQETYTLTFGNPFVSKQLQYSATFDNFIAGNYLLVEHENLPSCVEVMVFCGTRPGQPLQSLPSHDHDKGCDWFFNSKLRKLTYLGKCNASNIGNVVQMSISPPDDILKWSFPESWSGVKKGWGGYNHSIPAPGEDVIILPNRSILVDTTLPPLRSLYILGTLEFPVNSSNVLSVTCIVVAGGELKVGTFQHPLERGMKLLILLRASEGVYCDRLDGINVHPGTIGVYGKVQIHSAYPRKSWTHLGSDIAPGNERIWVEDEVDWSPGGNIVISSSSYDAHQAEVVTLKEINGHSIRFQERLFHRHIGRSHSIEDGRRVPLAAEVGLLTRNIQIKSDTACSGRLLVGRFRNTSGTEFAGDLRLSNIEILNFGPSQLPAIGFTNISLGSSIISSSIHHSCGGGIQAVTSSGISLRDNVVFNTVGPGIDLEGQTHSLIRNLIILSKQPKGLPDWVAGIKVNKIDDSYLLGNVMAGSERIAFHIRGQECFLAGEYFAENVAHSSLHGVHLYRGDGFQNCTKITGFLSYKNYDYGVMFQLESSVVMDNMMLVDNAVGVLPVVYSPSAKQHRLGKKYIELRNCVIIATSSTFDCIKDRITPLSANMTSRDRAPYSPQRGRTGILWPSFASEPSQWPNNPWHKIRNYAAVSGIMKLQEVTFTGFTKSCYSDDRDICIMSNPDSVGIMYPITAEMTRMSHVHEQNKFYFHTLETSINGSKDFEYLEMRCEGSRKALFKDLDGSALGLQPPVSVFPKSKSEWTESCLHAGIYREDTKCIYKPSTQGYFCKDTDHALVIIESLDTEADGRRLSPVVSVTGSFVEAFSSGVSHVSCCPSEHPRAFYSVLPSNKLTKVCFAGLTPPSLRLRLISGQNSTRLLIAIFYDEPRSLHVFMNGNYIPLASSSSSVFLTNAMAGTHSFSFENNLLYILLHGDDPIEIYTSFSLHTAFTIAETIGEEGQRRVVHRLADFLQIGRDHVRIVQSVSGSENTLKVISDNVGKRRRECPPMTSCTASCSRAGWEKTRTGAASVWILQSSHAATPLRVLIIEIGEPPGLFRNKFFPSLSSDRLKSLASTVINAQGYLGNCLGIGIGCFKIILNNMMAFFFPHCCLYVRPYNISVLVQPSDGEMEQELPVQPQIIFLDKQGRRVETLGLPSEPWVMTACLKGSSEAVLKGHTRVEVRDGRAIFTNLAISNSGSNWCLVFTVISPPGADFTVESQPFTIFPIPKGGKSITVLAAVLGSAASVLVMGLLVFCWSKKSKSFSKWESPGELSICRVSPISLVSKE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNH7</accession>
    <name>A0A452HNH7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Attractin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="27 28">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="28" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="27" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="27" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="16">Involved in the initial immune cell clustering during inflammatory response and may regulate chemotactic activity of chemokines. May play a role in melanocortin signaling pathways that regulate energy homeostasis and hair color. Low-affinity receptor for agouti. Has a critical role in normal myelination in the central nervous system.</text>
    </comment>
    <comment type="subunit">
        <text evidence="17">Monomer and homotrimer.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="19">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018852.1">
        <property type="protein sequence ID" value="ENSGAGP00000016518.1"/>
        <property type="gene ID" value="ENSGAGG00000011749.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030246">
        <property type="term" value="F:carbohydrate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006954">
        <property type="term" value="P:inflammatory response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00041">
        <property type="entry name" value="CUB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00055">
        <property type="entry name" value="EGF_Lam"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.120.10.80:FF:000031">
        <property type="entry name" value="attractin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.120.10.80:FF:000034">
        <property type="entry name" value="Attractin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000079">
        <property type="entry name" value="Attractin like 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000164">
        <property type="entry name" value="Attractin like 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.290:FF:000008">
        <property type="entry name" value="Attractin like 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.120.10.80">
        <property type="entry name" value="Kelch-type beta propeller"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.100.10">
        <property type="entry name" value="Mannose-Binding Protein A, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.290">
        <property type="entry name" value="Spermadhesin, CUB domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056737">
        <property type="entry name" value="Beta-prop_ATRN-MKLN-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001304">
        <property type="entry name" value="C-type_lectin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016186">
        <property type="entry name" value="C-type_lectin-like/link_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016187">
        <property type="entry name" value="CTDL_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000859">
        <property type="entry name" value="CUB_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056732">
        <property type="entry name" value="GBD_ATRN"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015915">
        <property type="entry name" value="Kelch-typ_b-propeller"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002049">
        <property type="entry name" value="LE_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056863">
        <property type="entry name" value="LMN_ATRN_NET-like_EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051568">
        <property type="entry name" value="LZTR1/Attractin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002165">
        <property type="entry name" value="Plexin_repeat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016201">
        <property type="entry name" value="PSI"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035914">
        <property type="entry name" value="Sperma_CUB_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46376:SF3">
        <property type="entry name" value="ATTRACTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46376">
        <property type="entry name" value="LEUCINE-ZIPPER-LIKE TRANSCRIPTIONAL REGULATOR 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24981">
        <property type="entry name" value="Beta-prop_ATRN-LZTR1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00431">
        <property type="entry name" value="CUB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24973">
        <property type="entry name" value="EGF_LMN_ATRN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24972">
        <property type="entry name" value="GBD_ATRN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01437">
        <property type="entry name" value="PSI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00034">
        <property type="entry name" value="CLECT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00042">
        <property type="entry name" value="CUB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00180">
        <property type="entry name" value="EGF_Lam"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00423">
        <property type="entry name" value="PSI"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56436">
        <property type="entry name" value="C-type lectin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF117281">
        <property type="entry name" value="Kelch motif"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49854">
        <property type="entry name" value="Spermadhesin, CUB domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50041">
        <property type="entry name" value="C_TYPE_LECTIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01180">
        <property type="entry name" value="CUB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01248">
        <property type="entry name" value="EGF_LAM_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50027">
        <property type="entry name" value="EGF_LAM_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11 19" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="4 19" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="14" id="KW-0395">Inflammatory response</keyword>
    <keyword evidence="2" id="KW-0880">Kelch repeat</keyword>
    <keyword evidence="15 20" id="KW-0424">Laminin EGF-like domain</keyword>
    <keyword evidence="7" id="KW-0430">Lectin</keyword>
    <keyword evidence="10" id="KW-0472">Membrane</keyword>
    <keyword evidence="12" id="KW-0675">Receptor</keyword>
    <keyword evidence="28" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="6 22" id="KW-0732">Signal</keyword>
    <keyword evidence="5" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="22">
        <location>
            <begin position="1"/>
            <end position="48"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019210833" description="Attractin" evidence="22">
        <location>
            <begin position="49"/>
            <end position="1322"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="24">
        <location>
            <begin position="53"/>
            <end position="86"/>
        </location>
    </feature>
    <feature type="domain" description="CUB" evidence="23">
        <location>
            <begin position="88"/>
            <end position="204"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="24">
        <location>
            <begin position="202"/>
            <end position="239"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="26">
        <location>
            <begin position="755"/>
            <end position="875"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="25">
        <location>
            <begin position="1015"/>
            <end position="1060"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="21">
        <location>
            <begin position="1"/>
            <end position="24"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="19">
        <location>
            <begin position="57"/>
            <end position="67"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="19">
        <location>
            <begin position="76"/>
            <end position="85"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="19">
        <location>
            <begin position="206"/>
            <end position="216"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="19">
        <location>
            <begin position="210"/>
            <end position="227"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="19">
        <location>
            <begin position="229"/>
            <end position="238"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="1032"/>
            <end position="1041"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="1044"/>
            <end position="1058"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022441"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022734"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023198"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023292"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00058400"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00065783"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00074596"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00460"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS01180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS50027"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="26">
        <source>
            <dbReference type="PROSITE" id="PS50041"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016518.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="28">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1322" mass="147262" checksum="E5103AE10072A20A" modified="2019-05-08" version="1">MERSSRRQPAAGRLGSQGGRPQRFGGRPPAQLLLVLLMLLVPGPEAAAAAAVEGKECDKPCANGGRCQPGSGQCECQAGWVGDQCQHCGGRFRLTGSSGYVTDGPGHYKYKTKCTWLIEGKPNTILRLRFNHFATECSWDHLYVYDGDSVYAPLVAAFSGLIVPEKDGNETIPEVVATSGYVLLHFFSDAAYNLTGFNITYNFDMCPNNCSGRGECKHSNGSNTVKCECSEHWKGEACDISYCTDDCGYPARGQCNMNETKGCLCNSGWQGPGCAVPVPANQSFWTREEYSSPKLPRASHKAVVNGNVMWVVGGYMFNHSDSHMILAYDLVSEEWLPLNSTVNSVEIRYGHSLALHENKIYMYGGKIDAMGNVTNQLWVFHIHNESWVHVIPKTKEQYAVVGHSAHIVNLGDGSAVMLVIFGHCPLYGYISNVQEYNLATNTWNILQTSGALVQGGYGHSSVYDPNTQSIYIHGGYKAFSANKYRLADDLYKYEVDTRMWTILKDSRFFRYLHTAVIMSGTMLVFGGNTHNDTSMSHGAKCFSSDFMAYDIACDHWSVLPRPDLHHDVNRFGHSAVLSNSTVYIFGGFNSLLLSDILRYTPESCEAYDNETSCLRAGPGVRCVWNTTFSLCVPWERATVETLVRKCLVLVMQVFSLIVVDHEKCNQITDCYSCTANTNNCQWCTDQCILKNNNCTEDQVPITEYDKCPKDNPAYYCNKKTSCKSCAMDQNCQWEPRNQECIALPENICGANWHLVGNSCLKINSTKESYDSGKLACTTQNALLASLTTQKKVEFVLKELQRMQSLAKTLTPWVGLRKINVSYWCWEDMSPFTNTLVQWLPGEPSGAGFCGYLAEPYQRGLKAATCIDHVNGSICERLANHSAKQCRTPCALRTACGECTSGSSECMWCSNMKQCVDSNAYVASFPYGQCMEWYTMTANCSGYCTCAHCLEQPGCGWCTDPSNTGKGKCMEGSYRGPVKMPSPPATGKYYLEPILNVSMCPAENKYNWSFIQCPACQCNGHSKCVNESVCEKCENLTTGKHCETCISGYYGDPTNGGTCQSCKCNGHASVCNTNTGKCFCTTKGIKGDECQLCEVENRYQGNPLKGTCYYTLLIDYQFTFSLSQEDDRYYTAINFVATPEEQNRDLDMFINASKNFNLNITWSTSFAAGTQAGEEIPVVSKSNIKEYKDSFSNEKFDFRSNPNITFFVYVSNFTWPIKIQIAFSQHSNFMDLVQFFVTFFRGVLQVAFLTVLHFRYTCASQALDWRFPVSSICSTYACALYNFVPRPRAYRAARANHPQFLPSNLCLRWSSSMTALCVPQCVL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNI3</accession>
    <name>A0A452HNI3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Protein O-mannosyl-transferase 1</fullName>
            <ecNumber evidence="4">2.4.1.109</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>a di-trans,poly-cis-dolichyl beta-D-mannosyl phosphate + L-seryl-[protein] = 3-O-(alpha-D-mannosyl)-L-seryl-[protein] + a di-trans,poly-cis-dolichyl phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17377"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:13546"/>
            <dbReference type="Rhea" id="RHEA-COMP:19498"/>
            <dbReference type="Rhea" id="RHEA-COMP:19501"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:57683"/>
            <dbReference type="ChEBI" id="CHEBI:58211"/>
            <dbReference type="ChEBI" id="CHEBI:137321"/>
            <dbReference type="EC" id="2.4.1.109"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="12">
            <text>a di-trans,poly-cis-dolichyl beta-D-mannosyl phosphate + L-threonyl-[protein] = 3-O-(alpha-D-mannosyl)-L-threonyl-[protein] + a di-trans,poly-cis-dolichyl phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:53396"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:13547"/>
            <dbReference type="Rhea" id="RHEA-COMP:19498"/>
            <dbReference type="Rhea" id="RHEA-COMP:19501"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:57683"/>
            <dbReference type="ChEBI" id="CHEBI:58211"/>
            <dbReference type="ChEBI" id="CHEBI:137323"/>
            <dbReference type="EC" id="2.4.1.109"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="2">Protein modification; protein glycosylation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the glycosyltransferase 39 family.</text>
    </comment>
    <dbReference type="EC" id="2.4.1.109" evidence="4"/>
    <dbReference type="AlphaFoldDB" id="A0A452HNI3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018882.1">
        <property type="protein sequence ID" value="ENSGAGP00000016546.1"/>
        <property type="gene ID" value="ENSGAGG00000012372.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00378"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004169">
        <property type="term" value="F:dolichyl-phosphate-mannose-protein mannosyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="CDD" id="cd23281">
        <property type="entry name" value="beta-trefoil_MIR_POMT1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.80.10.50:FF:000012">
        <property type="entry name" value="Protein O-mannosyl-transferase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.80.10.50">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003342">
        <property type="entry name" value="ArnT-like_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036300">
        <property type="entry name" value="MIR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016093">
        <property type="entry name" value="MIR_motif"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027005">
        <property type="entry name" value="PMT-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032421">
        <property type="entry name" value="PMT_4TMC"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10050">
        <property type="entry name" value="DOLICHYL-PHOSPHATE-MANNOSE--PROTEIN MANNOSYLTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10050:SF51">
        <property type="entry name" value="PROTEIN O-MANNOSYL-TRANSFERASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02815">
        <property type="entry name" value="MIR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02366">
        <property type="entry name" value="PMT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16192">
        <property type="entry name" value="PMT_4TMC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00472">
        <property type="entry name" value="MIR"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF82109">
        <property type="entry name" value="MIR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50919">
        <property type="entry name" value="MIR"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="5" id="KW-0328">Glycosyltransferase</keyword>
    <keyword evidence="11 15" id="KW-0472">Membrane</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0808">Transferase</keyword>
    <keyword evidence="7 15" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="10 15" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="68"/>
            <end position="88"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="100"/>
            <end position="120"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="126"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="154"/>
            <end position="174"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="186"/>
            <end position="219"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="240"/>
            <end position="261"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="575"/>
            <end position="597"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="609"/>
            <end position="627"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="639"/>
            <end position="659"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="671"/>
            <end position="691"/>
        </location>
    </feature>
    <feature type="domain" description="MIR" evidence="16">
        <location>
            <begin position="296"/>
            <end position="359"/>
        </location>
    </feature>
    <feature type="domain" description="MIR" evidence="16">
        <location>
            <begin position="370"/>
            <end position="427"/>
        </location>
    </feature>
    <feature type="domain" description="MIR" evidence="16">
        <location>
            <begin position="431"/>
            <end position="491"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004477"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004922"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007222"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00012839"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022676"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00045085"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00045102"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00067952"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50919"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016546.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="725" mass="82798" checksum="FFF63E1733616EA1" modified="2019-05-08" version="1">MLGFLKQPVVVTAEINMNLVALTVMGLVNRLWALSYPRAVVFDEVYYGQFVSLYMKRIFFVDDSGPPFGHMLLAFGGYLGGFDGNFLWNRIGAEYSMNVPVWSLRLLPALTGALCVPLAYQILVELHFSHCAAFGAALLILFENSLITQSRLMLLESVLIFFILLAILSYLKFYNLQKCSPFTAGWWFWLLLTGVACSCAVGVKYMGLFTYMLLLAIAGVHSWHLIGDQTLSNVSVLCHLLARGLALVVVPVAMYLSFFYVHLTVLYRSGPHDQIMTSAFQASLEGGLARITQGQPLEVAYGSQITLRNVLGKPVPCWLHSHKNTYPIRYENGRGSSHQQQVTCYPFKDVNNWWIVKDPGMQQLVVSNPPRPVRHGHIVQLVHGITTRYLNTHDVAAPLSPHSQEISCYIDYNISMPAQNLWRVEIVNRESDTDVWKTILSEVRFVHVNTSAVLKLSGASLPEWGYRQLEIIGEKLSKGHHQSMLWNVEEHRYGKSQEQKEREVELHSPTQADISKNLSFMARFTELQWKILTLKNEDTEHKYSSSPLDWITMDTNIAYWFHPISGAQIHLIGNILTWASANVATLVYVCLFLWYLLRRRRKICDIPEDAWRLWVSAGGLCVGGWAVNYLPFFMMEKTLFLYHYLPALTFQILLIPVVLQHLTDHLCRSQLPKNMFSALMVAWFSSIYLVYRTFSPLTYGEPTLSAAELRSLRWKDSWDILIRKH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNI4</accession>
    <name>A0A452HNI4_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="14">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HNI4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018850.1">
        <property type="protein sequence ID" value="ENSGAGP00000016516.1"/>
        <property type="gene ID" value="ENSGAGG00000012353.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006355">
        <property type="term" value="P:regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd07765">
        <property type="entry name" value="KRAB_A-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001772">
        <property type="entry name" value="Uncharacterized protein"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002026">
        <property type="entry name" value="Zinc finger protein 160"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002343">
        <property type="entry name" value="Zinc finger protein 33A"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002090">
        <property type="entry name" value="Zinc finger protein 473"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001532">
        <property type="entry name" value="Zinc finger protein 483"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002254">
        <property type="entry name" value="Zinc finger protein 540"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001909">
        <property type="entry name" value="KRAB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036051">
        <property type="entry name" value="KRAB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050758">
        <property type="entry name" value="Znf_C2H2-type"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23234:SF8">
        <property type="entry name" value="C2H2-TYPE DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23234">
        <property type="entry name" value="ZNF44 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01352">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="SMART" id="SM00349">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109640">
        <property type="entry name" value="KRAB domain (Kruppel-associated box)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50805">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="13"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="9" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 11" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="KRAB" evidence="13">
        <location>
            <begin position="9"/>
            <end position="80"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="125"/>
            <end position="152"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="153"/>
            <end position="180"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="181"/>
            <end position="208"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="209"/>
            <end position="236"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="237"/>
            <end position="264"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="265"/>
            <end position="292"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="293"/>
            <end position="320"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="321"/>
            <end position="348"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="349"/>
            <end position="376"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="377"/>
            <end position="404"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="405"/>
            <end position="432"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="433"/>
            <end position="460"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="461"/>
            <end position="488"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016516.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="488" mass="56153" checksum="77B0AACE5A82B295" modified="2019-05-08" version="1">MHPMKWMPVTFEKVAVYFTQGQGALLDPTQRALYKDVMLENYELVTSLGFPIPKPDLIARLEQGEEPWVPDIQNEENQQEEGPGKVVSQEMVFRKAEGNFSQCLEQGNRSLLLAHQRLHTGERPYRCLECGKSFSLKSTLNRHRKTHTGEKPHKCLDCGKTFSQRSNLTKHRGIHTAERPYKCLDCGKTFGQSSYLIKHQRIHTGERPYKCLECGKNFIRHSNLTKHRRIHTGDKPYECNDCGKSFIERTKLTRHQAIHTGEKPHKCLDCGKTFSQKSQLLIHGRLHTGERPYKCLECGRSFSQSSNLITHQRTHTGERPNICLECGKRFKHSSSLINHRRIHTGERPYKCLECGKSFMESSSLTKHGRIHTGERPHKCLECGKGFIESSSLTKHGRIHTGERPYKCLECGKSFSLKSTLNSHQKTHAEEKPHKCLDCGNTFARRSSLIKHERIHMGERPYKCFDCGKSFSKSSELTKHQRIHKGERT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNI6</accession>
    <name>A0A452HNI6_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="15">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="9">Belongs to the CSMD family.</text>
    </comment>
    <comment type="caution">
        <text evidence="11">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018899.1">
        <property type="protein sequence ID" value="ENSGAGP00000016560.1"/>
        <property type="gene ID" value="ENSGAGG00000012256.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="CDD" id="cd00033">
        <property type="entry name" value="CCP"/>
        <property type="match status" value="25"/>
    </dbReference>
    <dbReference type="CDD" id="cd00041">
        <property type="entry name" value="CUB"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.70.10:FF:000011">
        <property type="entry name" value="CUB and sushi domain-containing protein 3 isoform A"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.290:FF:000001">
        <property type="entry name" value="CUB and sushi domain-containing protein 3 isoform X1"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.70.10:FF:000002">
        <property type="entry name" value="CUB and Sushi multiple domains 3"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.70.10">
        <property type="entry name" value="Complement Module, domain 1"/>
        <property type="match status" value="25"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.290">
        <property type="entry name" value="Spermadhesin, CUB domain"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000859">
        <property type="entry name" value="CUB_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051277">
        <property type="entry name" value="SEZ6_CSMD_C4BPB_Regulators"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035914">
        <property type="entry name" value="Sperma_CUB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035976">
        <property type="entry name" value="Sushi/SCR/CCP_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000436">
        <property type="entry name" value="Sushi_SCR_CCP_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45656:SF6">
        <property type="entry name" value="CUB AND SUSHI DOMAIN-CONTAINING PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45656">
        <property type="entry name" value="PROTEIN CBR-CLEC-78"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00431">
        <property type="entry name" value="CUB"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00084">
        <property type="entry name" value="Sushi"/>
        <property type="match status" value="25"/>
    </dbReference>
    <dbReference type="SMART" id="SM00032">
        <property type="entry name" value="CCP"/>
        <property type="match status" value="25"/>
    </dbReference>
    <dbReference type="SMART" id="SM00042">
        <property type="entry name" value="CUB"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57535">
        <property type="entry name" value="Complement control module/SCR domain"/>
        <property type="match status" value="25"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49854">
        <property type="entry name" value="Spermadhesin, CUB domain"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01180">
        <property type="entry name" value="CUB"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50923">
        <property type="entry name" value="SUSHI"/>
        <property type="match status" value="25"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7 11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="8" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6 12" id="KW-0472">Membrane</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="2 11" id="KW-0768">Sushi</keyword>
    <keyword evidence="3 12" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 12" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="2993"/>
            <end position="3014"/>
        </location>
    </feature>
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            <begin position="99"/>
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    <sequence length="3069" mass="335650" checksum="22D3DE8A6694CCD8" modified="2019-05-08" version="1">MGCRALQLPGHCFEISKVSRLRQRIESVVCRECHSDRVPLKDFGISLICDYFIFLIIGLGVQQGSSYKSIIQTLQQSFQLPEPFFPPLLFYFLSEIDQGSCGDPGIPAYGKREGSTFRHGDTLKFECQPAFELKGQKTITCQKSSQWSAQKPVCVFSCFFNFTTPSGIVLSPNYPEEYGSSLHCVWLIIAKPESRIHLAFNDFDVEPQFDFLAVKDGGTPESPVLGTFSGSQIPSSLTSSGHVARLEFQTDHSMEKRGFNITFTTFRHNECPDPGVPVNGKRFGDSLQLGSSVSFLCDEGFIRTHGSETITCILKEGNVVWNNAVLRCEAPCGGHLTSPSGMILSPGWPGFYKDSLNCVWVIEAQPGYPIKITFDRFKTEVNYDTLEVRDGRSYSSPLIGVYDGTQVPQFLISTSNYLYLLFSTDKSHSDIGFQIRYETVKLQSDHCLDPGIPVNGQRHGNDFYVGALVTFSCDSGYTLSDGEALECEPNFQWSRPLPSCEALCGGYIRSSSGTILSPGFPDFYPNNLNCTWVIETSHGKGVFFTFHTFHLESGHDYLLVTENGSFTQPLKQLTGSRLPPPISAGLFGNFSAQIRFISDFSMSYEGFNITFSGKTECGSSVTGTQGVLLSPNYPINYNNNHECIYSIQSQPGKGIQLKARTFELDSGDVLKVYDGNNNSARLLGSFSHTEMLGTSINSTSSSMWLEFISNSENTSKGFELQFSSFELIKCEDPGIPQFGYKVRDEGHFAGSSVSFNCDPGYTLRGSRALVCLTGERRAWDQPLPVCIAECGGTIRGESSGRILSPAYPTPYDHNLNCIWTIEAEAGCTIGLHFMVFHTEEFHDVLRIWDGPVENGIILKEMSGSSLPSDVHSTFNSVVLQFNTDFFTSKQGFAIQFSVSTATSCNDPGIPQNGSRSADSKEAGDSIVFQCDPGYTLQGEAKISCVQIENRFFWQPDPPSCIAPCGGILTGPAGVILSPQYPEPYPPGKECDWKLTVSPDYVIALVFNIFNLEPGYDFLHIYDGPSSLSPLIGSFYGSQLPDRIESSSNNLFLAFRSDASVSNAGFVIDYTENPRESCFDPGSVKNGTRVGTDLKLGSTITYYCDGGYEIEGVSTLTCIMGGDGKPTWNQPRPTCTAPCGGQYTGSDGVVLSPNYPRNYTSGQICLYFVIVPKDYVVFGQFAFFQTALNDVIEVHDGPNQHSRLLSSLSGSHTGESLPLATTNQILIKFSAKGQATAKGFHFVYQAVPRTSATQCSSVPEPRYGRRIGSDFAVGAIVRFECSAGYALQGSRSIECLTMPGALAQWNVSVPTCLVPCGGNLTERKGTILSPGFPEPYLNSLNCVWKITVPEGAGIQIQVISFVTEQNWDSLEVFDGGDNTVTMLGSFSGTTVPALLNSTSNQLYLHFYSDISVSAAGFHLEYKTVGLSSCPEPAVPGNGLKIGERYLVNDVVSFQCEPGYALQGHSHISCMPGTVRRWNYPPPLCIAQCGGTAEEMEGVILSPGFPGNYPSNMDCSWRISLPVGFGAHIQFLNFSTEPNHDFIEIRNGPYDTSNVIGRFSGTEIPGSLLSTSHETTVYFHSDHSQNKPGFKLEYQAYELQECPDPEPFANGIVRGAGYNVGQSVSFECLPGYQLIGHPVLTCQHGTNRNWDHSLPRCEVPCGGNITTQNGTVYSPGFPNQYPNSQDCTWLITMSAGYGIHLNFTLLQMEPYNDFITIWDGPQQTAPQLGMFTGNSVKKSAQSSSNQILMKFHSDATNGGIFAIYFYAYQLFKCHPPTIVPNAEIITENEEFNIGDIVRYQCLPGFTLSGTEILTCKLGTHLQFGGPPPTCEVHCLMNEVLMDSTGVILSQSYLGRYPHFQTCSWVIKVEPGYNITLTIEYFLSEKQYDEFEIFDGPAGQSPLLLSLSGNYSAPLTVTSTGNKVYLRWSSDHTYSRKGFKIRYSAPYCSLPLPPFHGSILSHTGLQPGSEILFGCDAGYRLIGHSGTTCSRHPEGYYHWSEAVPLCQAISCGVPRAPTNGIVFGKEYTVGTKAVYHCNQGFHLQATEESTTECLQTGQWSNSNIPPQCMAVTCPDINSITVEHGRWRLTYETQYQYDAQLMLICDPGYYYTGQRVIRCQANGTWSIGDPMPTCKIISCGDLPTPPNGNRIGTLTIYGATAIFSCNTGYTLVGSRVRECMANGLWSGSEVKCLAGHCGIPNPIVNGQITGENYNYRGSVVYQCNPGFRLIGMSVRICQQDHHWSGKTPVCVPITCGHPGNPANGLTQGSQFNLNDMAKFVCNVGYHLEGASQSQCLANGQWSNSLPTCHIVNCTDPGHLENSIRQVQPSGPHRFSFGTTVSYQCSHGYYLLGTHVLSCQGDGTWDRSLPQCLLVSCGHPGSPPHSQISGDRYTVGSVIRYSCLEKRSLIGNSTRMCQLDGHWSGSLPHCSGGSQGTCGDPGIPSHGIRLGSEFGVDSMVRFSCELGYTLRGSSERTCHANGSWSGMQPECEVISCGNPGTPSNARVVFNDGLVFSSSIIYQCREGYYSTGVLSRHCTVNGTWTGSTPECTVINCGDPGVPANGFRLGNDFNYNKTVVFQCIPGYMMESDRASSLICTKDRTWNGTKPVCKAITCKSPQVIPNGKVVGSDFSWGSSVSYACLEGYQLSLPAILTCEGNGTWTGELPQCFPVFCGDPGTPAQGRREDRGFTYRSSVSFSCLSPLVLVGSARRFCQSDGTWSGTQPSCIDPTLTTCADPGVPLFGMQNNSQGYQVGSIMFFRCQKGYLLQGSTTRTCLPNLTWSGVQPDCVPHHCKQPETPSHANVGALDLPSLGYTLIYSCQSGYYLTGGSEHRTCKADGSWTGKTPICLGKSPEPAGRERTDWLGLFPADVFAKNSLWKGSYEYMGKKQPAMLSVTSFDPTTSKVNATLIDHSGVELLVSGIYKREDMHLLLQVYQITGPVEIFVNKFKNDKWALDGHVSISQLGFKRIYFCCVSDISLIENDPESIGHHFASNSSSVAAAILVPFIALIIAGFVLYLYKHRRRPKVPFNGYAGHENTNVRATFENPMYDRNLQPTDIMANEADFTVSTVCTAV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNI8</accession>
    <name>A0A452HNI8_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="3">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the SAPAP family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HNI8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018887.1">
        <property type="protein sequence ID" value="ENSGAGP00000016551.1"/>
        <property type="gene ID" value="ENSGAGG00000012373.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031616">
        <property type="term" value="C:spindle pole centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008017">
        <property type="term" value="F:microtubule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051642">
        <property type="term" value="P:centrosome localization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007059">
        <property type="term" value="P:chromosome segregation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051382">
        <property type="term" value="P:kinetochore assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007052">
        <property type="term" value="P:mitotic spindle organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007346">
        <property type="term" value="P:regulation of mitotic cell cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0023052">
        <property type="term" value="P:signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005026">
        <property type="entry name" value="SAPAP"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12353:SF1">
        <property type="entry name" value="DISKS LARGE-ASSOCIATED PROTEIN 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12353">
        <property type="entry name" value="DISKS LARGE-ASSOCIATED PROTEIN DAP SAP90/PSD-95-ASSOCIATED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03359">
        <property type="entry name" value="GKAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="64"/>
            <end position="92"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="211"/>
            <end position="239"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="350"/>
            <end position="404"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="609"/>
            <end position="636"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="659"/>
            <end position="711"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="229"/>
            <end position="239"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="369"/>
            <end position="402"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="618"/>
            <end position="636"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="683"/>
            <end position="702"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00008839"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016551.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="851" mass="94897" checksum="5C04F23519AE8B18" modified="2019-05-08" version="1">MAATSQFANRYRKDLSTETLRTKVARRKSLIQKENRHKQFERSRQFGLADVNIQLPKERGLPKLDETNEVCSQEKTTAKRKQTTNAAINKSENERKEMLQRYKEEKELRKLKEQREKAKRGVFKVGLYRPNAPGFLPFVPQNPVIVKPKEKVAPTSARITRSKTKSQAEQTVVSARSQQSMIYADHATNGHSMRPTQTGCRQTCADKVTEKKKALGNQQQRKTASRGKQQKEVKAEDVEAHKHEVEGNTLMDQLTEVSVSNSKNPTSEVHQEDTLLEKENILVTHMPVPVRKRTRSFAPQNFVFQPLDGLTTYKVTPMTPSRANAFLTPCLSWSPMKIDANVSKEESRKECLLKSQASPAQEGTDKGSQEQQIGTNSLKVKNECASTKTQDAVQSSHETSPVSADISAIETKTGETDHDVPYFRNVLASETERLMSHCLQWDGKFELDIPEDAKDLIRTTVGQTRLLIAERFQQFEGLVDNCEFKRGDKETTCTDLDGFWDMVSFQVEDVNKKFDYLSKLQENGWQPVNIPSKRAIKKRAVPSGISKPKQGAAGRTAARNRIAAIRAAMKEKMKQGETAADSAVQEMPKEEEKVVFEAGFFRIESPLKSFPALPSKTPGRSSQRISKQLTTPRSASRTLLQSSLAILCSSEDTVAKQTTPAASGFHVPQNPNRLQAPTRKTLFGSTPEQSGSPLVEEQSSMTDGAAERTNETNVAVLEVCLRPDERSSSGPVNSQALKDPDTKIEIPDVTEEMELSATDLTGQDIVMCSPEKNAQSETDFIQAEKPNTPEYKSDLLHSDFISSDGTPFDVPSLNCSLPFTPVRNEAQKFAAAELYNDLIVFSPLSPPSVEK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNJ1</accession>
    <name>A0A452HNJ1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Serine/threonine protein phosphatase 2A regulatory subunit</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the phosphatase 2A regulatory subunit B56 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HNJ1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018870.1">
        <property type="protein sequence ID" value="ENSGAGP00000016535.1"/>
        <property type="gene ID" value="ENSGAGG00000011705.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000159">
        <property type="term" value="C:protein phosphatase type 2A complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072542">
        <property type="term" value="F:protein phosphatase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000003">
        <property type="entry name" value="Serine/threonine-protein phosphatase 2A 56 kDa regulatory subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002554">
        <property type="entry name" value="PP2A_B56"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10257">
        <property type="entry name" value="SERINE/THREONINE PROTEIN PHOSPHATASE 2A PP2A REGULATORY SUBUNIT B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10257:SF89">
        <property type="entry name" value="SERINE_THREONINE-PROTEIN PHOSPHATASE 2A 56 KDA REGULATORY SUBUNIT DELTA ISOFORM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01603">
        <property type="entry name" value="B56"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF028043">
        <property type="entry name" value="PP2A_B56"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0539">Nucleus</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="493"/>
            <end position="530"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="518"/>
            <end position="530"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009745"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PIRNR" id="PIRNR028043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016535.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="530" mass="62110" checksum="4793469B1B8FA44A" modified="2019-05-08" version="1">NKIKYSGGPQIVKKERRHSSSRFNLSKNRELQKLPALKDAPAHEREELFIQKLRQCCVLFDFISDPLSDLKFKEVKRAGLNEMVEYITHNRDVVTEAIYPEAVIMFSVNLFRTLPPSSNPTGAEFDPEEDEPTLEAAWPHLQLVYEFFLRFLESPDFQPNIAKKYIDQKFILSLLDLFDSEDPRERDFLKTILHRIYGKFLGLRAYVRRQINNIFYRFIYETEHHNGIAELLEILGSIINGFALPLKEEHKMFLIRVLLPLHKVKSLSVYHPQLAYCVVQFLEKDSSLTEPVIVGLLKFWPKTHSPKEVMFLNELEEILDVIEPSEFMKVMEPLFRQLAKCVSSPHFQVAERALYYWNNEYIMSLISDNAAKILPIMFPALYKNSKSHWNKTIHGLIYNALKLFMEMNQKLFDDCTQQYKAEKQKGRFKLREREEMWHKVEELARQNPQYPMYYAPPPLPPIYCMETETPTAEDIQLLKKTVETEAVQVSQGPAPWQAACHPTSEVHQKIPGSRHRPRGDCRGSRWPEHV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNJ3</accession>
    <name>A0A452HNJ3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Neurobeachin-like protein 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the WD repeat neurobeachin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018904.1">
        <property type="protein sequence ID" value="ENSGAGP00000016565.1"/>
        <property type="gene ID" value="ENSGAGG00000012367.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005783">
        <property type="term" value="C:endoplasmic reticulum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019901">
        <property type="term" value="F:protein kinase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008104">
        <property type="term" value="P:intracellular protein localization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030099">
        <property type="term" value="P:myeloid cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd06071">
        <property type="entry name" value="Beach"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd01201">
        <property type="entry name" value="PH_BEACH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.1540.10:FF:000001">
        <property type="entry name" value="neurobeachin isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:001375">
        <property type="entry name" value="Neurobeachin-like protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000301">
        <property type="entry name" value="Neurobeachin-like protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1540.10">
        <property type="entry name" value="BEACH domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000409">
        <property type="entry name" value="BEACH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036372">
        <property type="entry name" value="BEACH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050865">
        <property type="entry name" value="BEACH_Domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046851">
        <property type="entry name" value="NBCH_WD40"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031570">
        <property type="entry name" value="NBEA/BDCP_DUF4704"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046852">
        <property type="entry name" value="Neurobeachin_a-sol"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023362">
        <property type="entry name" value="PH-BEACH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036322">
        <property type="entry name" value="WD40_repeat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001680">
        <property type="entry name" value="WD40_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13743">
        <property type="entry name" value="BEIGE/BEACH-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13743:SF111">
        <property type="entry name" value="NEUROBEACHIN-LIKE PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02138">
        <property type="entry name" value="Beach"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15787">
        <property type="entry name" value="DUF4704"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16057">
        <property type="entry name" value="DUF4800"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20426">
        <property type="entry name" value="NBCH_WD40"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20425">
        <property type="entry name" value="Neurobeachin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14844">
        <property type="entry name" value="PH_BEACH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01026">
        <property type="entry name" value="Beach"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00320">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81837">
        <property type="entry name" value="BEACH domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50978">
        <property type="entry name" value="WD40 repeat-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50197">
        <property type="entry name" value="BEACH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51783">
        <property type="entry name" value="PH_BEACH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50082">
        <property type="entry name" value="WD_REPEATS_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="3 7" id="KW-0853">WD repeat</keyword>
    <feature type="domain" description="BEACH-type PH" evidence="10">
        <location>
            <begin position="1912"/>
            <end position="2012"/>
        </location>
    </feature>
    <feature type="domain" description="BEACH" evidence="9">
        <location>
            <begin position="2025"/>
            <end position="2317"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="7">
        <location>
            <begin position="2470"/>
            <end position="2503"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="7">
        <location>
            <begin position="2521"/>
            <end position="2562"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="718"/>
            <end position="743"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1342"/>
            <end position="1362"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1399"/>
            <end position="1435"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="718"/>
            <end position="729"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="1399"/>
            <end position="1408"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022574"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00068540"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50197"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS51783"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016565.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2727" mass="305954" checksum="AF7A0431DDC63862" modified="2019-05-08" version="1">MNFEKIITVQSLDPRRPEDSVSEIPLLPRDVLHVLSQQLAHCVAQIAREEAAEAGLGQALLHVKFFIIICRNLENVQADRTPVFIHEVLSLLVACTAKLKNTPEQCAPGRSQLETVTIYALHLYECLFDPYQTWRRQLSGEIVSMKEKSKYKFAPAALPLEFSTFFQGSFQDEGQLPEALQLRLIHLLGAIISGAKQNALQVITPASVEVLMLVLKRWCSPRNGDTKDPKLLQLTLKTLVGMIHILHASNPGQRKVEIRTILDSYFKVLNSDQPMASLEGSPAARWEEGMIALRVNMLDAIPEMLNCGDRPVLQAMFLSNNCFEHIIRLIQNSKLYVSGARKADETGSELAARLLTESDVQKVFDSSSDAITVHAISVLTSIMRNSPSAKEVFKERIGYSHLYEVLRCQSQPTARLLEELLNMAVEGDHSAFPVRLIRNEQPLLMLIEWLPELKSRDLQVFLSDWLKRVCDASLPSRVTCVKAGMVSCLLDTLGSEQELDRKCSENLIYLLQVLGSLSIRPGELRQLIRLLWTEPGKGPHPYTMLVIRALSGMARKDRSERALQYFDLTPSMAGIMVPAIQKWPGSGFAFHAWVCLNEEPEGLPDLPTRLKRKQLYSFFTASGTGFEAFFTANGMLVVAVCTKKEYMTVALPELTFNDSAWHCIDIVHVTGRRPFGQNLVNIYTDGQLRQVAQLRFPSLNESFTSCCIGSAGHRTTTTSTVYPSPSHSPELAFPSHPSLSRSQSFPASLATHSWTPGAAQPPREGQVSTMAAGSQDTEWGTPTSLEGHLGSVSIFCEALQQTQVKALFSSGPNVTSPLKPEGDFLELSSKLLLYYTPQDVVNCVGGMGVLLPLLEQVAAERKESEDSQETHDLVGPELTSSRNRQGMLIPLGKSSESRLEKNGVAAFLLLVKNFIRHHPVNQESLVQCHGPAIIGALLQKVSSLMMDMNVLMASQILMEQVAAESHSFLLHLLYQYLLFDFRIWSKSDFAVRLGHIQYLSNTIKDHKQRIRKKYGVQYILDSIRTYYGTHKENPVPTDVKTVQMSLFSLVKDFLCRSFSADEMQSTLSYLAAVHDEHQVCGVLEVIHSLLKDSASQEQLYAFLFESGNVEVLFSLLVQRKFSDEVRERVFKVLYKMLKYEKAHERSKYRLKLKDVGYQGLISNLNDIPVSMLLFRCLLEQVLGSDSPSYKDLMAVVYLSHRAELAVRLDTCRKLFHLIYSQHDMVRQLAKLAGWQDTLTKLYVKESYESRQHSLSGSANGGCLGLLKLSEHAGKESGTEAGKEQLDSASSALADLQDLDVFLPMGFESSDHELSEGFSDHSVSPTGGSKPFRSYNFKSFDSLDRTSHSSSSNTIDIPGQREASSTDALQFDSIYHPLSPFCMSPFDLGLDLASTCSTATVESGNQTPGSLPGTPSPLEYFKPFPGTQARKSSSLSNVLDESSYQETLASDDVSNTSNPQQTPEEELCNLLTNIIFSVTWRGVEGWDDAAWKERGQVFSVLTKLGTACELVRPPDEIKCSLLEMMLESAFTDIKESAPATLPSLTQNVLKLLRLLQDFLFSEGHNNQALWSEKIYEGVNSLLDKLGVWYHLANGTSDLKEMAQIGLRVLVGYSNLEEAQLHSLAYVKLHGLLQTASAPKEEEACYLLGKLESPLSRALHPKSEAFSWLLPIIRTLLDQCYETLQLQRRLPSLPPTNGSPTFYEDFQLFCTSPEWRAFIEKHVQPMMAQFEIDTFAKSHDHMSNFWNSCYDALMSSSLRREREKEESRKKFQELILEPVMKRCKYEHVRHVNVLKQINNHHSIVLKQWKALRRLLTSQRSAWADGNPPEIRWKLSSAETYSRMRLKLVPNYHFDPHVEASALRDNLGADHSHNLVEPLPLAVAKEAKVSELEDDQLAEEDLEFLDNQAEPKEQSQREKLVISEDCELITIVAVIQGRLEVTTQHVYFYDCSSEKEETEEGIGYDFKRPLSQLREVHLRRYNLRRSALEFFFIDQANYFVNFKKKVRNKVYSSILGLRPPNQIYFGSRSPQELLKASGLTQKWVYREISNFEYLMQLNTIAGRTYNDLSQYPVFPWILQDYVSETLDLSNPGVFRDLSKPIGVANERHAKDVKEKYESFEDSTGMIDKFHYGTHYSNAAGVMHYMIRTEPFTTLHIQLQSGRFDCSDRQFHSVPAAWQARMENPVDVKELIPEFFYFTDFLENQNGFDLGCLQMSNDQVNDVVLPKWARSPEDFIHKHRKALESEHMSAHLHEWIDLIFGYKQRGPAAVEALNVFYYCTYEGAVDLDAIADETQRKALEGIISNFGQTPCQLLKEPHPVRLSAENATRRLARLDTYSPNVFENLNQLKSFFVEGISDGVPLVQAVVPKNQAHSFITQGSPDVLVTVSANGLLGTHSWLPYDKNISNYFCFTKDPTISNAKAQRFLSGPFAPGVEVSSKTLTVSPDGKLLFSGGHWDNSLRVTSLSKGKVIGHITRHIDVVTCLALDLCGIYLISGSRDTTCMVWQVLQQGGFSSGLAPKPVQVLYGHDAEVTCVAISTELDMAVSGSKDGTIIVHTIRRGQFMKSIKPPCESSLPATVSNLAVGLEGQIIVQTTVAGRSSLKDKFALHLYSVNGKHLSSVPLEEQVTVLCVTEEFVVLGTMQCSLQILDLQSLQAAVLPMPMKVPVHSVSVTKEKSHILVGLEDGKLIVVGAGQPSEVRTGQFHRRLWRSTRRISQVSSGETEYNPPESK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNJ5</accession>
    <name>A0A452HNJ5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Neurobeachin-like protein 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the WD repeat neurobeachin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018888.1">
        <property type="protein sequence ID" value="ENSGAGP00000016552.1"/>
        <property type="gene ID" value="ENSGAGG00000012367.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005783">
        <property type="term" value="C:endoplasmic reticulum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019901">
        <property type="term" value="F:protein kinase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008104">
        <property type="term" value="P:intracellular protein localization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030099">
        <property type="term" value="P:myeloid cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd06071">
        <property type="entry name" value="Beach"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd01201">
        <property type="entry name" value="PH_BEACH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.1540.10:FF:000001">
        <property type="entry name" value="neurobeachin isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:001375">
        <property type="entry name" value="Neurobeachin-like protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000301">
        <property type="entry name" value="Neurobeachin-like protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1540.10">
        <property type="entry name" value="BEACH domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000409">
        <property type="entry name" value="BEACH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036372">
        <property type="entry name" value="BEACH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050865">
        <property type="entry name" value="BEACH_Domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046851">
        <property type="entry name" value="NBCH_WD40"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031570">
        <property type="entry name" value="NBEA/BDCP_DUF4704"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046852">
        <property type="entry name" value="Neurobeachin_a-sol"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023362">
        <property type="entry name" value="PH-BEACH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036322">
        <property type="entry name" value="WD40_repeat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001680">
        <property type="entry name" value="WD40_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13743">
        <property type="entry name" value="BEIGE/BEACH-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13743:SF111">
        <property type="entry name" value="NEUROBEACHIN-LIKE PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02138">
        <property type="entry name" value="Beach"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15787">
        <property type="entry name" value="DUF4704"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16057">
        <property type="entry name" value="DUF4800"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20426">
        <property type="entry name" value="NBCH_WD40"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20425">
        <property type="entry name" value="Neurobeachin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14844">
        <property type="entry name" value="PH_BEACH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01026">
        <property type="entry name" value="Beach"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00320">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81837">
        <property type="entry name" value="BEACH domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50978">
        <property type="entry name" value="WD40 repeat-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50197">
        <property type="entry name" value="BEACH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51783">
        <property type="entry name" value="PH_BEACH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50082">
        <property type="entry name" value="WD_REPEATS_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="3 7" id="KW-0853">WD repeat</keyword>
    <feature type="domain" description="BEACH-type PH" evidence="10">
        <location>
            <begin position="1945"/>
            <end position="2045"/>
        </location>
    </feature>
    <feature type="domain" description="BEACH" evidence="9">
        <location>
            <begin position="2058"/>
            <end position="2350"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="7">
        <location>
            <begin position="2503"/>
            <end position="2536"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="7">
        <location>
            <begin position="2554"/>
            <end position="2595"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="720"/>
            <end position="745"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1375"/>
            <end position="1395"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1432"/>
            <end position="1468"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="720"/>
            <end position="731"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="1432"/>
            <end position="1441"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022574"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00068540"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50197"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS51783"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016552.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2760" mass="309930" checksum="18F0042ECBF0E6F9" modified="2019-05-08" version="1">MELPSCQMGSPAIAFQKDLTYLQQWLEAFVMNFEKIITVQSLDPRRPEDSVSEIPLLPRDVLHVLSQQLAHCVAQIAREEAAEAGLGQALLHVKFFIIICRNLENVQADRTPVFIHEVLSLLVACTAKLKNTPEQCAPGRSQLETVTIYALHLYECLFDPYQTWRRQLSGEIVSMKEKSKYKFAPAALPLEFSTFFQGSFQDEGQLPEALQLRLIHLLGAIISGAKQNALQVITPASVEVLMLVLKRWCSPRNGDTKDPKLLQLTLKTLVGMIHILHASNPGQRKVEIRTILDSYFKVLNSDQPMASLEGSPAARWEEGMIALRVNMLDAIPEMLNCGDRPVLQAMFLSNNCFEHIIRLIQNSKVFDSSSDAITVHAISVLTSIMRNSPSAKEVFKERIGYSHLYEVLRCQSQPTARLLEELLNMAVEGDHSAFPVRLIRNEQPLLMLIEWLPELKSRDLQVFLSDWLKRVCDASLPSRVTCVKAGMVSCLLDTLGSEQELDRKCSENLIYLLQVLGSLSIRPGELRQLIRLLWTEPGKGPHPYTMLVIRALSGMARKDRSERALQYFDLTPSMAGIMVPAIQKWPGSGFAFHAWVCLNEEPEGLPDLPTRLKRKQLYSFFTASGTGFEAFFTANGMLVVAVCTKKEYMTVALPELTFNDSAWHCIDIVHVTGRRPFGQNLVNIYTDGQLRQVAQLRFPSLNESFTSCCIGSAGHRTTTTSTVYPSPSHSPELAFPSHPSLSRSQSFPASLATHSWTPGAAQPPREGQVSTMAAGSQDTEWGTPTSLEGHLGSVSIFCEALQQTQVKALFSSGPNVTSPLKPEGDFLELSSKLLLYYTPQACKNNICLDLSPNHFFDGRLTGHKVVNWDIKDVVNCVGGMGVLLPLLEQVAAERKESEDSQETHDLVGPELTSSRNRQGMLIPLGKSSESRLEKNGVAAFLLLVKNFIRHHPVNQESLVQCHGPAIIGALLQKVSSLMMDMNVLMASQILMEQVAAESHSFLLHLLYQYLLFDFRIWSKSDFAVRLGHIQYLSNTIKDHKQRIRKKYGVQYILDSIRTYYGTHKENPVPTDVKTVQMSLFSLVKDFLCRSFSADEMQSTLSYLAAVHDEHQVCGVLEVIHSLLKDSASQEQLYAFLFESGNVEVLFSLLVQRKFSDEVRERVFKVLYKMLKYEKAHERSKYRLKLKDVGYQGLISNLNDIPVSMLLFRCLLEQVLGSDSPSYKDLMAVVYLSHRAELAVRLDTCRKLFHLIYSQHDMVRQLAKLAGWQDTLTKLYVKESYESRQHSLSGSANGGCLGLLKLSEHAGKESGTEAGKEQLDSASSALADLQDLDVFLPMGFESSDHELSEGFSDHSVSPTGGSKPFRSYNFKSFDSLDRTSHSSSSNTIDIPGQREASSTDALQFDSIYHPLSPFCMSPFDLGLDLASTCSTATVESGNQTPGSLPGTPSPLEYFKPFPGTQARKSSSLSNVLDESSYQETLASDDVSNTSNPQQTPEEELCNLLTNIIFSVTWRGVEGWDDAAWKERGQVFSVLTKLGTACELVRPPDEIKCSLLEMMLESAFTDIKESAPATLPSLTQNVLKLLRLLQDFLFSEGHNNQALWSEKIYEGVNSLLDKLGVWYHLANGTSDLKEMAQIGLRVLVGYSNLEEAQLHSLAYVKLHGLLQTASAPKEEEACYLLGKLESPLSRALHPKSEAFSWLLPIIRTLLDQCYETLQLQRRLPSLPPTNGSPTFYEDFQLFCTSPEWRAFIEKHVQPMMAQFEIDTFAKSHDHMSNFWNSCYDALMSSSLRREREKEESRKKFQELILEPVMKRCKYEHVRHVNVLKQINNHHSIVLKQWKALRRLLTSQRSAWADGNPPEIRWKLSSAETYSRMRLKLVPNYHFDPHVEASALRDNLGADHSHNLVEPLPLAVAKEAKVSELEDDQLAEEDLEFLDNQAEPKEQSQREKLVISEDCELITIVAVIQGRLEVTTQHVYFYDCSSEKEETEEGIGYDFKRPLSQLREVHLRRYNLRRSALEFFFIDQANYFVNFKKKVRNKVYSSILGLRPPNQIYFGSRSPQELLKASGLTQKWVYREISNFEYLMQLNTIAGRTYNDLSQYPVFPWILQDYVSETLDLSNPGVFRDLSKPIGVANERHAKDVKEKYESFEDSTGMIDKFHYGTHYSNAAGVMHYMIRTEPFTTLHIQLQSGRFDCSDRQFHSVPAAWQARMENPVDVKELIPEFFYFTDFLENQNGFDLGCLQMSNDQVNDVVLPKWARSPEDFIHKHRKALESEHMSAHLHEWIDLIFGYKQRGPAAVEALNVFYYCTYEGAVDLDAIADETQRKALEGIISNFGQTPCQLLKEPHPVRLSAENATRRLARLDTYSPNVFENLNQLKSFFVEGISDGVPLVQAVVPKNQAHSFITQGSPDVLVTVSANGLLGTHSWLPYDKNISNYFCFTKDPTISNAKAQRFLSGPFAPGVEVSSKTLTVSPDGKLLFSGGHWDNSLRVTSLSKGKVIGHITRHIDVVTCLALDLCGIYLISGSRDTTCMVWQVLQQGGFSSGLAPKPVQVLYGHDAEVTCVAISTELDMAVSGSKDGTIIVHTIRRGQFMKSIKPPCESSLPATVSNLAVGLEGQIIVQTTVAGRSSLKDKFALHLYSVNGKHLSSVPLEEQVTVLCVTEEFVVLGTMQCSLQILDLQSLQAAVLPMPMKVPVHSVSVTKEKSHILVGLEDGKLIVVGAGQPSEVRTGQFHRRLWRSTRRISQVSSGETEYNPPESK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNJ9</accession>
    <name>A0A452HNJ9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">BAT2 N-terminal domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000018868.1">
        <property type="protein sequence ID" value="ENSGAGP00000016533.1"/>
        <property type="gene ID" value="ENSGAGG00000012361.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009738">
        <property type="entry name" value="BAT2_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033184">
        <property type="entry name" value="PRRC2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14038">
        <property type="entry name" value="BAT2 HLA-B-ASSOCIATED TRANSCRIPT 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14038:SF4">
        <property type="entry name" value="PROTEIN PRRC2B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07001">
        <property type="entry name" value="BAT2_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="1" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="BAT2 N-terminal" evidence="4">
        <location>
            <begin position="1"/>
            <end position="189"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="50"/>
            <end position="469"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="542"/>
            <end position="564"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="596"/>
            <end position="660"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="876"/>
            <end position="1057"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="1090"/>
            <end position="1247"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="1271"/>
            <end position="1527"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="1569"/>
            <end position="1604"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="1618"/>
            <end position="1762"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="1855"/>
            <end position="1884"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="2">
        <location>
            <begin position="496"/>
            <end position="523"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="88"/>
            <end position="137"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="179"/>
            <end position="190"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="233"/>
            <end position="247"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="3">
        <location>
            <begin position="303"/>
            <end position="318"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="375"/>
            <end position="388"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="423"/>
            <end position="434"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="441"/>
            <end position="456"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="544"/>
            <end position="564"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="612"/>
            <end position="621"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="3">
        <location>
            <begin position="629"/>
            <end position="639"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="876"/>
            <end position="885"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="928"/>
            <end position="939"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="956"/>
            <end position="990"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="3">
        <location>
            <begin position="1024"/>
            <end position="1033"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="1034"/>
            <end position="1048"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="1158"/>
            <end position="1192"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="1222"/>
            <end position="1235"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="1271"/>
            <end position="1283"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="1305"/>
            <end position="1322"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="1382"/>
            <end position="1392"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="1425"/>
            <end position="1434"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="1502"/>
            <end position="1522"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="1569"/>
            <end position="1578"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="1618"/>
            <end position="1657"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="3">
        <location>
            <begin position="1677"/>
            <end position="1686"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="1704"/>
            <end position="1734"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="1741"/>
            <end position="1756"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="1859"/>
            <end position="1869"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="Pfam" id="PF07001"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016533.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2185" mass="242391" checksum="00A248068A78A6D5" modified="2019-05-08" version="1">MSDRLGQITKGKDGKSKYSTLSLFDKYKGKSIEAIRTTVIPRHGLQSLGKVAAARRMPPPANLPSLKSENKGNDPNIIIVPKDGTGWANKQDQPDQKSSSATTAQPQESLPQQGLQKSVSNLQKPTQSISQESTNSVPGGPKSWAQLNGKPAGQEGGSRVSSRLLSFSPEEFPTLKAAGEQDKVGKEKGALDLSYGPGPSLRPQNVTSWREGGGRNIISATSLTASPPEPGIKTSSTGDGDPSSVSASDPKEPSLRPAQPIRKGASQFMGNAYHPPTYHDMLPAFMCPQQSSEVPGSLDRGSFPLPQLRLQPRVPFRQYQMNDQDGKENRPGVSRPTRPVRQQGERAPRPTIINAENLKELDELDTDADDGWAGIHDEVDYSEKLKFSEDEEEEEALKDGRQKWNTWDPRRQRQLSLSSADSADIKHTPEEGKNWGDQVGLSRSVSKVQGPQQPSRKLNGWTPASEYQKPPLGCILRQQSLEDKEEKVPLRQKFVHSEISEAVERARRRREEEERRAREERLAACAAKLKQLDQKCKLAQKTGETQKHVENEDLKPPSVEKKVAQENGHTFRRATSEFHAQDVSVGYLEEETTATTVAAQNSSEEEIREATSPAQEFNKYQKSLPPRFQRQQQQQQQEQLYKMQHWQQQQVYPPPSHSHPQRTFYPPHPQMLGFDPRWMMMPSYMDPRMAQGRTPVDFYPSALYPSGIMKPMIQQDSINGSGCRSEEQNCQPSLQQERKAASMDPAPVWGQESYTSLQNKGYPLPHQKQTENMSMEGLHTRSDSSYSASSGRSESINVQRNLFEDRGDEYLNTFDKKAQSDFDSSISSQRIGQDLLFQHQETVQDACVSGARHASLRCSPLESDFIQVDKKPEYNGWDINHHQKSSEPAAEADEEVPRDEQSFSADPWKKEGAVSKQPAEETDWAPENRNTNNQHQEQMGRTRRSGPIKKPVLKALKVEEKEKEQEKIKLEGEELSHPAKERAPVYKVENDSEDSSTLLNSTHYLLDEKGSSQANFTQDAEKSQEEEEEEEEEKAERAWENKLSRESNDLPPTKRNNWIFIDEEQAFGGRGQGRGRGRGFREFTFRGRGTAVSGRGVYNNQRSSRGRGLREFNQLEDFPRGKPRRRIASETHSEGSEYEELPKRRRQRGSENSNESSLLEREESDLKKGDFKDSWRSNKIYSDDHSSLDAKMRAPRAFGRSLPPRLSNSGYGRRGFMAKEPSQWQGRSGGTTWQEYSHCPPSDTFGIRRQSDRDYIQDSYKHVDSFTGRGFEENHIDNRRPLFQEDYTADQENIENRPFRRRRPPRQDKPPRFRRLRQERESVGQWNPEEGGTSLLPSQWRPKLTSVEKNGVSGRRSPELSYQNSSDHANEEWETASESSDFSERRERRDGVLESEAQLDGGLASGSLGEKRELAKRSFSSQRPLVDRQSRKAEPAGFGEQLVRAGGGAASRFESQQNGMPMKNKRSPEEGGGLGNAKSGSSHSVYSLERSLCTSSDSAEGPGKKQEKEPKSTAQRPSEKGETMSQFEVNYGNAIIDHRVSSTAEENEVGSMAGEGFIEVLTKKQRRLLEEERRKKEQAAQGPTKGRVLQSRIPPRFAKKQNSMCLEQGDVTVSGSSLGTEIWESNSPALSVQSPGSDSWSKPVNAFNGTESSSTEQGFKGSQGDSGIDLSAESRESSATSSQRSSPYGTLKPEEMNGAGLVDPKPDCQKEQVQKQSDKKDSDQGSGQNKEHKPGPIGNERSLKNRKGSEGTERLEGNIPPVNGVEIHVDSVLPVPPIEFGVNPKDSDFSLPPGSASGTAASPVTKLQDALASNAGLTQSIPILRRDHHLQRCIGLNPMSFPTADLTLKMESARKAWENSPSLPEQNSPGGAGSGIQPPSSVGASSGVSYSSFGGVSMPPMPVASVAPSASIPGNHIPPLYLDGHVFASQPRLVPQTIPQQQSYQQAAAAQQIPISLHTSLQAQAQLGLRGGLPVSQSQEIYSSIQPFRSPVYMHPSLSQPSTMVLTGGAALKPPYSAFPGMQPLEMVKTQSGSPYQPMSGSQTLVYEGQINQAASLGASQMMDSQLTQGLKPVMERKAEASVNLPDVPVPEITPAEEYSRFLDVDIELKTKADNAYAWVPASSPPVWFWPAATDFTTVHPASSGAESVCRSSPQDSSTWISAFCPHYQQGVLPNGNERVSLHRW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNK2</accession>
    <name>A0A452HNK2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">phospholipase A2</fullName>
            <ecNumber evidence="1">3.1.1.4</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="6">
            <text>a 1,2-diacyl-sn-glycero-3-phosphocholine + H2O = a 1-acyl-sn-glycero-3-phosphocholine + a fatty acid + H(+)</text>
            <dbReference type="Rhea" id="RHEA:15801"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:28868"/>
            <dbReference type="ChEBI" id="CHEBI:57643"/>
            <dbReference type="ChEBI" id="CHEBI:58168"/>
            <dbReference type="EC" id="3.1.1.4"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="6">
            <dbReference type="Rhea" id="RHEA:15802"/>
        </physiologicalReaction>
    </comment>
    <dbReference type="EC" id="3.1.1.4" evidence="1"/>
    <dbReference type="AlphaFoldDB" id="A0A452HNK2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018881.1">
        <property type="protein sequence ID" value="ENSGAGP00000016545.1"/>
        <property type="gene ID" value="ENSGAGG00000011814.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0047499">
        <property type="term" value="F:calcium-independent phospholipase A2 activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0052816">
        <property type="term" value="F:long-chain fatty acyl-CoA hydrolase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035965">
        <property type="term" value="P:cardiolipin acyl-chain remodeling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016042">
        <property type="term" value="P:lipid catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:2000304">
        <property type="term" value="P:positive regulation of ceramide biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07212">
        <property type="entry name" value="Pat_PNPLA9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.1090.10">
        <property type="entry name" value="Cytosolic phospholipase A2 catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016035">
        <property type="entry name" value="Acyl_Trfase/lysoPLipase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047148">
        <property type="entry name" value="PLPL9"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002641">
        <property type="entry name" value="PNPLA_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24139:SF34">
        <property type="entry name" value="85_88 KDA CALCIUM-INDEPENDENT PHOSPHOLIPASE A2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24139">
        <property type="entry name" value="CALCIUM-INDEPENDENT PHOSPHOLIPASE A2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01734">
        <property type="entry name" value="Patatin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01415">
        <property type="entry name" value="ANKYRIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52151">
        <property type="entry name" value="FabD/lysophospholipase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51635">
        <property type="entry name" value="PNPLA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4 7" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="3 8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="8" id="KW-0442">Lipid degradation</keyword>
    <keyword evidence="5 8" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <feature type="repeat" description="ANK" evidence="7">
        <location>
            <begin position="54"/>
            <end position="86"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="7">
        <location>
            <begin position="184"/>
            <end position="216"/>
        </location>
    </feature>
    <feature type="domain" description="PNPLA" evidence="10">
        <location>
            <begin position="320"/>
            <end position="504"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="249"/>
            <end position="280"/>
        </location>
    </feature>
    <feature type="short sequence motif" description="GXGXXG" evidence="8">
        <location>
            <begin position="324"/>
            <end position="329"/>
        </location>
    </feature>
    <feature type="short sequence motif" description="GXSXG" evidence="8">
        <location>
            <begin position="356"/>
            <end position="360"/>
        </location>
    </feature>
    <feature type="short sequence motif" description="DGA/G" evidence="8">
        <location>
            <begin position="491"/>
            <end position="493"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="263"/>
            <end position="280"/>
        </location>
    </feature>
    <feature type="active site" description="Nucleophile" evidence="8">
        <location>
            <position position="358"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="8">
        <location>
            <position position="491"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00013278"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023422"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01161"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS51635"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016545.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="645" mass="71373" checksum="ECFFB9B3B16811B7" modified="2019-05-08" version="1">MECLCELVECHARLDVTDKNGETVFHYAVRGSNPEIVKLLGKKPSVGLNHLSNQGQTPLHLACQLGKEDVVLALLKCHAKCNVMGTLGYPIHTAVKYSQKGCVQALLEVDVNQVQFKDPRYGAAVLHWAKNAEMTRMLIEYGCDVNSVSKTADMALHIMVKRGRFDSAMVLLTHGALTNAKGQDGNTPLHLAMKQDHLETIKALIVFGADVEIPNEFGETPGLVAARTSKGANRKVLLGLLQMVGTERCHPPIPESPHMASAESPSLSTSPERPQLPRNNGHSLGYESIIHISTALGHLLKTPDVVDSASDAKRTYDRLLCLDGGGIRGLVLIQLLIAVEKAAGRPIRELFDWIAGTSTGGILALAIVHGKPVDYMRCLYFRMKNEVFRGSRPYESEPLDEFLKREFGEHTKMTDVQKPKVIVTGTLCDRQPAELHLFRNYKAPATKNKPEVKTTDPFKPLTKPEDQLVWRAARCSGAAPTYFRPIGRFLDGGLLANNPTLDAMTEIHEYNKSLSQKGQSQQVRKLGVVVSLGTGRPPQVPVRSVDVFRPSNPWELAKTVFGAKELGKMVVDCCTDADGPAVDRARAWCEMTDTRYFRFSPQLQSDVMLDEVNDAILVNALWDTQLYICQQQEQFKQLVQELLTR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNK4</accession>
    <name>A0A452HNK4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">PA14 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000018873.1">
        <property type="protein sequence ID" value="ENSGAGP00000016538.1"/>
        <property type="gene ID" value="ENSGAGG00000012280.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="CDD" id="cd00603">
        <property type="entry name" value="IPT_PCSR"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052387">
        <property type="entry name" value="Fibrocystin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014756">
        <property type="entry name" value="Ig_E-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002909">
        <property type="entry name" value="IPT_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037524">
        <property type="entry name" value="PA14/GLEYA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46769:SF1">
        <property type="entry name" value="FIBROCYSTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46769">
        <property type="entry name" value="POLYCYSTIC KIDNEY AND HEPATIC DISEASE 1 (AUTOSOMAL RECESSIVE)-LIKE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01833">
        <property type="entry name" value="TIG"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SMART" id="SM00429">
        <property type="entry name" value="IPT"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56988">
        <property type="entry name" value="Anthrax protective antigen"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81296">
        <property type="entry name" value="E set domains"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51820">
        <property type="entry name" value="PA14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0732">Signal</keyword>
    <feature type="domain" description="PA14" evidence="2">
        <location>
            <begin position="88"/>
            <end position="247"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="PROSITE" id="PS51820"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016538.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1482" mass="161186" checksum="FE0F35A2B93C1039" modified="2019-05-08" version="1">MVHKDAWLISAKQEVFLYQTFSDIASVFPVSGSLGGGSDLTITGDFFDQPVQVTVAGVPCQIKHCSPRKISCITGPIAEIRRLAAPQAGNRGLLFEVWDDAAGTDLTETAPGYRWQIVPNASSPERFLSQAQQSFSSRLRGFFVAPQTNNYTFWIQADGRASLHLSLSEDPGKKVKVAFIPAGASEWSEHWEKNWSKSWQPKSQKFELMGGSKYYLEALHHGKAPSSGMRVGVQLHNTWLNPDVVNTYRRERHEIWASALRLPEIQMLTLSGTGWFRISWDNVSSNMMATNATADQVQTVIEELLSVKCETEPLSAKFLLRNGFEEGNRDPATEGHVASWTEPFCGRFSIHRPKHLIKMSPSILPRYNLTEYTHVCFAYKGYVSDTLYVSVSYINTFLNTVKKNLTCLWSLNETSPESWKFTCADLWRCCVNGSESLRDLEANSSVFVHQIDMLIPEREEETSSWFYLDEIIVSDRNIAVFQKDFKPARPGGNVIQEVSVVGSPPTYNVSFLVAGCGSNLPLLGALPSQGSEEDHLAAFMEGANVSMTVQRVQAASPPIGGTFRILLPNTVISGVPVHVSSHHLRKLLQSNSDNSTGQYFNASDFTVTKDVSTCYQDVWTLTWNSMTGDLPIVIDVSAENLTGLNPTIAARVVYDGGVFIGPIFGDMLATANNHTQVVVVVNDVPANCSGSCSFQYSQEATPLVSHVDYSLGNVYITGSGFAEDSQALQIEVNHTTCQVMTSNQTAVVCWMNMLPMGLYHVTLLVRPYGFARNASEGQGIYLRVKPRLEAIEPPTTSEIGGCSVILRGTGFDGISQVLFGSHPCPINTNASNSVRIECEVPSRGDEEYVVHVTLTAGSQSATFTEVFKYDPSLNPAIVSLSRNRSSTAGLQSLRIGISSFANCTGLDVQVKIEATVAQIQARTAHGVDVVLPALADGWYNLSVIINGIAIIPSRGSACGGLLLTISGLGLKSRRNSVHVSLTGNYTCEIRSVDHNRISCALLPREPPLGDWWLPDASHALNVTVTVNGIQSVCLSDCALHLLKEWTPRVDAVVSEINGTLSSLLIRGQRLALATDELVIQVDNHDPCNVTFWNETSIQCQTGSLSPGEHSLSVSNRRSGHACFSRDSRIFTVIPCVLKFYPQNFSTNGGGLLAIQGAALKGRNSTSILVGNRPCLITDVSYVVLRCSVPLGNGIQDLTLEIDGISYHLGEISYSEEFTPIFLFLLPPVGLLLTMTVSRITGVESMHISIGDSPCTNITGNHTTLQCSCSAPWLPAGEYRVLGGDFLRGWASSNLTFTSELTVISTRNNWGKCGLGGGEVHVHGTGFAPGNTSVTICGAPCEMLGDITTTDVSCLAPRLDASLAVLCGLTHPSEDCKGAGTTYVQCDVHITAGAYSLAGARPYIYLCDDDSLLDSKTKVNSPQALFAGLFFSPKVERDEVLIYNSSCNITMETEAEMECEGANQPITAKITEIRKNWGQNTQV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNK6</accession>
    <name>A0A452HNK6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Protein tyrosine phosphatase domain-containing protein 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="4">May play roles in cilia formation and/or maintenance.</text>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the protein-tyrosine phosphatase family. Non-receptor class PTPDC1 subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HNK6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018902.1">
        <property type="protein sequence ID" value="ENSGAGP00000016563.1"/>
        <property type="gene ID" value="ENSGAGG00000012384.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060271">
        <property type="term" value="P:cilium assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd14506">
        <property type="entry name" value="PTP_PTPDC1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000027">
        <property type="entry name" value="Protein tyrosine phosphatase domain containing 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000340">
        <property type="entry name" value="Dual-sp_phosphatase_cat-dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050561">
        <property type="entry name" value="PTP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049573">
        <property type="entry name" value="PTPDC1_PTP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016130">
        <property type="entry name" value="Tyr_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003595">
        <property type="entry name" value="Tyr_Pase_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020422">
        <property type="entry name" value="TYR_PHOSPHATASE_DUAL_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23339">
        <property type="entry name" value="TYROSINE SPECIFIC PROTEIN PHOSPHATASE AND DUAL SPECIFICITY PROTEIN PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00782">
        <property type="entry name" value="DSPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00195">
        <property type="entry name" value="DSPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00404">
        <property type="entry name" value="PTPc_motif"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00383">
        <property type="entry name" value="TYR_PHOSPHATASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50054">
        <property type="entry name" value="TYR_PHOSPHATASE_DUAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="1" id="KW-0970">Cilium biogenesis/degradation</keyword>
    <keyword evidence="2" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="3" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="7">
        <location>
            <begin position="83"/>
            <end position="254"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="8">
        <location>
            <begin position="169"/>
            <end position="236"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022794"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00056295"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00060867"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00072096"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016563.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="675" mass="77042" checksum="AA8016E53BD0C4B4" modified="2019-05-08" version="1">MAAGVLLRNELPYCSLVKSSAYLSTMSSGNLRRPTAKYTKVGERLRHVIPGHMQCSMACGGRACKYENPARWSDQEQAIKGLYSSWITDNILAMARPSTEIIEKYNIIEQFQRCDIKTVINLQRPGEHASCGNPLEQESGFTYLPEAFMEDGIYFYNFGWKDYGVASLTTILDMVKVMAFALQEGRVAVHCHAGLGRTGVLIACYLVFATRMTADQAILFVRAKRPNSIQTRGQLVCVREFTQFLIPLRNVFACCEPKAHAVTLSQYLIRQRHLLHGYESRHLKYVPKLIHLVCKLLLDLAENRQVIEEELLDIPDLSTEIEEAVSQLDSTQLDKELTRQNSDTSEPFSHSVVGNVTFETQDSVFSHERKYDPLWKRRNVECLQPLTHSKRRLSYSDSDLRRTAFLLEQGETPWTVPAQVPFFDQLKQQNTREHFLSSGIQSPQLDLNKEALVRSTFIFWSQGKFSLDGQNNFYTKKSPKEVQRSKTFSSGFERVHNDREARTSKCSSAKDIGHRKKRETNVYGRRVCASEDSDNSTLEEESSIGYESQDSKDLLEAVPCIVLQSELSLDARRILAAKALANLNEFMSEEEVKQKVEMWQEQYQTILCIFHCVVNLQTIPADVEEALLTRAIKAFTKMSLDSENGPQVYNTLRKVFKQILEAKRKEAKKETDNLS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNL0</accession>
    <name>A0A452HNL0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Zinc finger matrin-type protein 5</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="10">U11/U12 small nuclear ribonucleoprotein 20 kDa protein</fullName>
        </alternativeName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HNL0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018879.1">
        <property type="protein sequence ID" value="ENSGAGP00000016543.1"/>
        <property type="gene ID" value="ENSGAGG00000012370.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005689">
        <property type="term" value="C:U12-type spliceosomal complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006397">
        <property type="term" value="P:mRNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008380">
        <property type="term" value="P:RNA splicing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000741">
        <property type="entry name" value="Zinc finger matrin-type protein 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.1000.10">
        <property type="entry name" value="Zinc finger, CCCH-type"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013085">
        <property type="entry name" value="U1-CZ_Znf_C2H2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000571">
        <property type="entry name" value="Znf_CCCH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036855">
        <property type="entry name" value="Znf_CCCH_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16465">
        <property type="entry name" value="NUCLEASE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16465:SF0">
        <property type="entry name" value="ZINC FINGER MATRIN-TYPE PROTEIN 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00642">
        <property type="entry name" value="zf-CCCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06220">
        <property type="entry name" value="zf-U1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00356">
        <property type="entry name" value="ZnF_C3H1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90229">
        <property type="entry name" value="CCCH zinc finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50103">
        <property type="entry name" value="ZF_C3H1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="12" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3 11" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="2" id="KW-0507">mRNA processing</keyword>
    <keyword evidence="7" id="KW-0508">mRNA splicing</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0747">Spliceosome</keyword>
    <keyword evidence="6 11" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 11" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C3H1-type" evidence="14">
        <location>
            <begin position="51"/>
            <end position="79"/>
        </location>
    </feature>
    <feature type="zinc finger region" description="C3H1-type" evidence="11">
        <location>
            <begin position="51"/>
            <end position="79"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="145"/>
            <end position="167"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="12">
        <location>
            <begin position="75"/>
            <end position="102"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="13">
        <location>
            <begin position="145"/>
            <end position="157"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022664"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022728"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023187"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00067764"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00076547"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50103"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016543.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="167" mass="19391" checksum="A2E150AF78CB9CE8" modified="2019-05-08" version="1">MGKRYFCDYCDRSFQDNLHNRKKHLNGVQHLRAKKAWYDLFRDAAAVLLEEQSKKPCRKFLQTGQCDFGFNCRFSHMTEEDLEKLNAQVQEERRSKEEGAEVPVGTVEDWLEKREKRVSLAQSNSSLTEKMLVFQYPPGWPPIQDLPPSLQAPPPGGWPVLPNLQWG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNL3</accession>
    <name>A0A452HNL3_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="15">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="9">Belongs to the CSMD family.</text>
    </comment>
    <comment type="caution">
        <text evidence="11">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018929.1">
        <property type="protein sequence ID" value="ENSGAGP00000016586.1"/>
        <property type="gene ID" value="ENSGAGG00000012256.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="CDD" id="cd00033">
        <property type="entry name" value="CCP"/>
        <property type="match status" value="25"/>
    </dbReference>
    <dbReference type="CDD" id="cd00041">
        <property type="entry name" value="CUB"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.70.10:FF:000011">
        <property type="entry name" value="CUB and sushi domain-containing protein 3 isoform A"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.290:FF:000001">
        <property type="entry name" value="CUB and sushi domain-containing protein 3 isoform X1"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.70.10:FF:000002">
        <property type="entry name" value="CUB and Sushi multiple domains 3"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.70.10">
        <property type="entry name" value="Complement Module, domain 1"/>
        <property type="match status" value="25"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.290">
        <property type="entry name" value="Spermadhesin, CUB domain"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000859">
        <property type="entry name" value="CUB_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051277">
        <property type="entry name" value="SEZ6_CSMD_C4BPB_Regulators"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035914">
        <property type="entry name" value="Sperma_CUB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035976">
        <property type="entry name" value="Sushi/SCR/CCP_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000436">
        <property type="entry name" value="Sushi_SCR_CCP_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45656:SF6">
        <property type="entry name" value="CUB AND SUSHI DOMAIN-CONTAINING PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45656">
        <property type="entry name" value="PROTEIN CBR-CLEC-78"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00431">
        <property type="entry name" value="CUB"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00084">
        <property type="entry name" value="Sushi"/>
        <property type="match status" value="25"/>
    </dbReference>
    <dbReference type="SMART" id="SM00032">
        <property type="entry name" value="CCP"/>
        <property type="match status" value="25"/>
    </dbReference>
    <dbReference type="SMART" id="SM00042">
        <property type="entry name" value="CUB"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57535">
        <property type="entry name" value="Complement control module/SCR domain"/>
        <property type="match status" value="25"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49854">
        <property type="entry name" value="Spermadhesin, CUB domain"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01180">
        <property type="entry name" value="CUB"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50923">
        <property type="entry name" value="SUSHI"/>
        <property type="match status" value="25"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7 11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="8" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6 12" id="KW-0472">Membrane</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="2 11" id="KW-0768">Sushi</keyword>
    <keyword evidence="3 12" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 12" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="2904"/>
            <end position="2925"/>
        </location>
    </feature>
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        <location>
            <begin position="1"/>
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    <sequence length="2980" mass="325357" checksum="5CC8FDCB872F8FD0" modified="2019-05-08" version="1">RHGDTLKFECQPAFELKGQKTITCQKSSQWSAQKPVCVCKSLGEYKVKCGCVSCFFNFTTPSGIVLSPNYPEEYGSSLHCVWLIIAKPESRIHLAFNDFDVEPQFDFLAVKDGGTPESPVLGTFSGSQIPSSLTSSGHVARLEFQTDHSMEKRGFNITFTTFRHNECPDPGVPVNGKRFGDSLQLGSSVSFLCDEGFIRTHGSETITCILKEGNVVWNNAVLRCEAPCGGHLTSPSGMILSPGWPGFYKDSLNCVWVIEAQPGYPIKITFDRFKTEVNYDTLEVRDGRSYSSPLIGVYDGTQVPQFLISTSNYLYLLFSTDKSHSDIGFQIRYESKYHCLDPGIPVNGQRHGNDFYVGALVTFSCDSGYTLSDGEALECEPNFQWSRPLPSCEALCGGYIRSSSGTILSPGFPDFYPNNLNCTWVIETSHGKGVFFTFHTFHLESGHDYLLVTENGSFTQPLKQLTGSRLPPPISAGLFGNFSAQIRFISDFSMSYEGFNITFSGKRFTFVLGDTECGSSVTGTQGVLLSPNYPINYNNNHECIYSIQSQPGKGIQLKARTFELDSGDVLKVYDGNNNSARLLGSFSHTEMLGTSINSTSSSMWLEFISNSENTSKGFELQFSSFELIKCEDPGIPQFGYKVRDEGHFAGSSVSFNCDPGYTLRGSRALVCLTGERRAWDQPLPVCIAECGGTIRGESSGRILSPAYPTPYDHNLNCIWTIEAEAGCTIGLHFMVFHTEEFHDVLRIWDGPVENGIILKEMSGSSLPSDVHSTFNSVVLQFNTDFFTSKQGFAIQFSVSTATSCNDPGIPQNGSRSADSKEAGDSIVFQCDPGYTLQGEAKISCVQIENRFFWQPDPPSCIAPCGGILTGPAGVILSPQYPEPYPPGKECDWKLTVSPDYVIALVFNIFNLEPGYDFLHIYDGPSSLSPLIGSFYGSQLPDRIESSSNNLFLAFRSDASVSNAGFVIDYTENPRESCFDPGSVKNGTRVGTDLKLGSTITYYCDGGYEIEGVSTLTCIMGGDGKPTWNQPRPTCTAPCGGQYTGSDGVVLSPNYPRNYTSGQICLYFVIVPKDYVVFGQFAFFQTALNDVIEVHDGPNQHSRLLSSLSGSHTGESLPLATTNQILIKFSAKGQATAKGFHFVYQAVPRTSATQCSSVPEPRYGRRIGSDFAVGAIVRFECSAGYALQGSRSIECLTMPGALAQWNVSVPTCLVPCGGNLTERKGTILSPGFPEPYLNSLNCVWKITVPEGAGIQIQVISFVTEQNWDSLEVFDGGDNTVTMLGSFSGTTVPALLNSTSNQLYLHFYSDISVSAAGFHLEYKTVGLSSCPEPAVPGNGLKIGERYLVNDVVSFQCEPGYALQGHSHISCMPGTVRRWNYPPPLCIAQCGGTAEEMEGVILSPGFPGNYPSNMDCSWRISLPVGFGAHIQFLNFSTEPNHDFIEIRNGPYDTSNVIGRFSGTEIPGSLLSTSHETTVYFHSDHSQNKPGFKLEYQAYELQECPDPEPFANGIVRGAGYNVGQSVSFECLPGYQLIGHPVLTCQHGTNRNWDHSLPRCEVPCGGNITTQNGTVYSPGFPNQYPNSQDCTWLITMSAGYGIHLNFTLLQMEPYNDFITIWDGPQQTAPQLGMFTGNSVKKSAQSSSNQILMKFHSDATNGGIFAIYFYAYQLFKCHPPTIVPNAEIITENEEFNIGDIVRYQCLPGFTLSGTEILTCKLGTHLQFGGPPPTCEVHCLMNEVLMDSTGVILSQSYLGRYPHFQTCSWVIKVEPGYNITLTIEYFLSEKQYDEFEIFDGPAGQSPLLLSLSGNYSAPLTVTSTGNKVYLRWSSDHTYSRKGFKIRYSAPYCSLPLPPFHGSILSHTGLQPGSEILFGCDAGYRLIGHSGTTCSRHPEGYYHWSEAVPLCQAISCGVPRAPTNGIVFGKEYTVGTKAVYHCNQGFHLQATEESTTECLQTGQWSNSNIPPQCMAVTCPDINSITVEHGRWRLTYETQYQYDAQLMLICDPGYYYTGQRVIRCQANGTWSIGDPMPTCKIISCGDLPTPPNGNRIGTLTIYGATAIFSCNTGYTLVGSRVRECMANGLWSGSEVKCLAGHCGIPNPIVNGQITGENYNYRGSVVYQCNPGFRLIGMSVRICQQDHHWSGKTPVCVPITCGHPGNPANGLTQGSQFNLNDMAKFVCNVGYHLEGASQSQCLANGQWSNSLPTCHIVNCTDPGHLENSIRQVQPSGPHRFSFGTTVSYQCSHGYYLLGTHVLSCQGDGTWDRSLPQCLLVSCGHPGSPPHSQISGDRYTVGSVIRYSCLEKRSLIGNSTRMCQLDGHWSGSLPHCSGGSQGTCGDPGIPSHGIRLGSEFGVDSMVRFSCELGYTLRGSSERTCHANGSWSGMQPECEVISCGNPGTPSNARVVFNDGLVFSSSIIYQCREGYYSTGVLSRHCTVNGTWTGSTPECTVINCGDPGVPANGFRLGNDFNYNKTVVFQCIPGYMMESDRASSLICTKDRTWNGTKPVCKAITCKSPQVIPNGKVVGSDFSWGSSVSYACLEGYQLSLPAILTCEGNGTWTGELPQCFPVFCGDPGTPAQGRREDRGFTYRSSVSFSCLSPLVLVGSARRFCQSDGTWSGTQPSCIDPTLTTCADPGVPLFGMQNNSQGYQVGSIMFFRCQKGYLLQGSTTRTCLPNLTWSGVQPDCVPHHCKQPETPSHANVGALDLPSLGYTLIYSCQSGYYLTGGSEHRTCKADGSWTGKTPICLGKSPEPAGRERTDWLGLFPADVFAKNSLWKGSYEYMGKKQPAMLSVTSFDPTTSKVNATLIDHSGVELLVSGIYKREDMHLLLQVYQITGPVEIFVNKFKNDKWALDGHVSISAHVGTFVYQGFVGGKGFGQFGLQRLGNLIENDPESIGHHFASNSSSVAAAILVPFIALIIAGFVLYLYKHRRRPKVPFNGYAGHENTNVRATFENPMYDRNLQPTDIMANEADFTVSTVCTAV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNL8</accession>
    <name>A0A452HNL8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Protein tyrosine phosphatase domain-containing protein 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="4">May play roles in cilia formation and/or maintenance.</text>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the protein-tyrosine phosphatase family. Non-receptor class PTPDC1 subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HNL8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018900.1">
        <property type="protein sequence ID" value="ENSGAGP00000016561.1"/>
        <property type="gene ID" value="ENSGAGG00000012384.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060271">
        <property type="term" value="P:cilium assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd14506">
        <property type="entry name" value="PTP_PTPDC1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000027">
        <property type="entry name" value="Protein tyrosine phosphatase domain containing 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000340">
        <property type="entry name" value="Dual-sp_phosphatase_cat-dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050561">
        <property type="entry name" value="PTP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049573">
        <property type="entry name" value="PTPDC1_PTP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016130">
        <property type="entry name" value="Tyr_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003595">
        <property type="entry name" value="Tyr_Pase_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020422">
        <property type="entry name" value="TYR_PHOSPHATASE_DUAL_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23339">
        <property type="entry name" value="TYROSINE SPECIFIC PROTEIN PHOSPHATASE AND DUAL SPECIFICITY PROTEIN PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00782">
        <property type="entry name" value="DSPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00195">
        <property type="entry name" value="DSPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00404">
        <property type="entry name" value="PTPc_motif"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00383">
        <property type="entry name" value="TYR_PHOSPHATASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50054">
        <property type="entry name" value="TYR_PHOSPHATASE_DUAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="1" id="KW-0970">Cilium biogenesis/degradation</keyword>
    <keyword evidence="2" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="3" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="7">
        <location>
            <begin position="83"/>
            <end position="254"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="8">
        <location>
            <begin position="169"/>
            <end position="236"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022794"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00056295"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00060867"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00072096"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016561.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="737" mass="84249" checksum="E5CC2BBEC9502F69" modified="2019-05-08" version="1">MAAGVLLRNELPYCSLVKSSAYLSTMSSGNLRRPTAKYTKVGERLRHVIPGHMQCSMACGGRACKYENPARWSDQEQAIKGLYSSWITDNILAMARPSTEIIEKYNIIEQFQRCDIKTVINLQRPGEHASCGNPLEQESGFTYLPEAFMEDGIYFYNFGWKDYGVASLTTILDMVKVMAFALQEGRVAVHCHAGLGRTGVLIACYLVFATRMTADQAILFVRAKRPNSIQTRGQLVCVREFTQFLIPLRNVFACCEPKAHAVTLSQYLIRQRHLLHGYESRHLKYVPKLIHLVCKLLLDLAENRQVIEEELLDIPDLSTEIEEAVSQLDSTQLDKELTRQNSDTSEPFSHSVVGNVTFETQDSVFSHERKYDPLWKRRNVECLQPLTHSKRRLSYSDSDLRRTAFLLEQGETPWTVPAQVPFFDQLKQQNTREHFLSSGIQSPQLDLNKEALVRSTFIFWSQGKFSLDGQNNFYTKKSPKEVQRSKTFSSGFERVHNDREARTSKCSSAKDIGHRKKRETNVYGRRVCASEDSDNSTLEEESSIGYESQDSKDLLEAVPCIVLQSELSLDARRILAAKALANLNEFMSEEEVKQKVEMWQKELNSRDGAWDRICAERDPFVLCSLMWSWIEQLKEPVITKEDIDMLSKKCTESPETLNLLAKEQYQTILCIFHCVVNLQTIPADVEEALLTRAIKAFTKMSLDSENGPQVYNTLRKVFKQILEAKRKEAKKETDNLS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQH2</accession>
    <name>A0A452IQH2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="19">Voltage-dependent R-type calcium channel subunit alpha</fullName>
        </recommendedName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="19">Voltage-sensitive calcium channels (VSCC) mediate the entry of calcium ions into excitable cells and are also involved in a variety of calcium-dependent processes, including muscle contraction, hormone or neurotransmitter release, gene expression, cell motility, cell division and cell death. The isoform alpha-1E gives rise to R-type calcium currents.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>Ca(2+)(in) = Ca(2+)(out)</text>
            <dbReference type="Rhea" id="RHEA:29671"/>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 19">Membrane</location>
            <topology evidence="1 19">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="19">Belongs to the calcium channel alpha-1 subunit (TC 1.A.1.11) family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000034453.1">
        <property type="protein sequence ID" value="ENSGAGP00000030358.1"/>
        <property type="gene ID" value="ENSGAGG00000002084.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043025">
        <property type="term" value="C:neuronal cell body"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045202">
        <property type="term" value="C:synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="GOC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005891">
        <property type="term" value="C:voltage-gated calcium channel complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008331">
        <property type="term" value="F:high voltage-gated calcium channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098703">
        <property type="term" value="P:calcium ion import across plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007268">
        <property type="term" value="P:chemical synaptic transmission"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.350:FF:000001">
        <property type="entry name" value="Voltage-dependent L-type calcium channel subunit alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.238.10:FF:000063">
        <property type="entry name" value="Voltage-dependent N-type calcium channel subunit alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.350:FF:000011">
        <property type="entry name" value="Voltage-dependent N-type calcium channel subunit alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.350:FF:000013">
        <property type="entry name" value="Voltage-dependent N-type calcium channel subunit alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.350:FF:000015">
        <property type="entry name" value="Voltage-dependent N-type calcium channel subunit alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.70:FF:000023">
        <property type="entry name" value="Voltage-dependent R-type calcium channel subunit alpha"/>
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    </dbReference>
    <dbReference type="FunFam" id="1.10.287.70:FF:000025">
        <property type="entry name" value="Voltage-dependent R-type calcium channel subunit alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="4"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.2500">
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Voltage-gated potassium channels. Chain C"/>
        <property type="match status" value="4"/>
    </dbReference>
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        <property type="entry name" value="GPHH_dom"/>
    </dbReference>
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        <property type="entry name" value="Ion_trans_dom"/>
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    <dbReference type="InterPro" id="IPR014873">
        <property type="entry name" value="VDCC_a1su_IQ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050599">
        <property type="entry name" value="VDCC_alpha-1_subunit"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005449">
        <property type="entry name" value="VDCC_R_a1su"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002077">
        <property type="entry name" value="VDCCAlpha1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027359">
        <property type="entry name" value="Volt_channel_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45628">
        <property type="entry name" value="VOLTAGE-DEPENDENT CALCIUM CHANNEL TYPE A SUBUNIT ALPHA-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45628:SF5">
        <property type="entry name" value="VOLTAGE-DEPENDENT R-TYPE CALCIUM CHANNEL SUBUNIT ALPHA-1E"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08763">
        <property type="entry name" value="Ca_chan_IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16905">
        <property type="entry name" value="GPHH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00520">
        <property type="entry name" value="Ion_trans"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00167">
        <property type="entry name" value="CACHANNEL"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01633">
        <property type="entry name" value="RVDCCALPHA1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01062">
        <property type="entry name" value="Ca_chan_IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81324">
        <property type="entry name" value="Voltage-gated potassium channels"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9 18 19" id="KW-0106">Calcium</keyword>
    <keyword evidence="5 19" id="KW-0107">Calcium channel</keyword>
    <keyword evidence="4 19" id="KW-0109">Calcium transport</keyword>
    <keyword evidence="20" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="14" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="15" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="16" id="KW-0407">Ion channel</keyword>
    <keyword evidence="12" id="KW-0406">Ion transport</keyword>
    <keyword evidence="13 22" id="KW-0472">Membrane</keyword>
    <keyword evidence="7 18" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="6 22" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 22" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="2" id="KW-0813">Transport</keyword>
    <keyword evidence="10 19" id="KW-0851">Voltage-gated channel</keyword>
    <feature type="transmembrane region" description="Helical" evidence="22">
        <location>
            <begin position="141"/>
            <end position="162"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="22">
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    <feature type="transmembrane region" description="Helical" evidence="22">
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    <feature type="transmembrane region" description="Helical" evidence="22">
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    <feature type="transmembrane region" description="Helical" evidence="22">
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    <feature type="transmembrane region" description="Helical" evidence="22">
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    <feature type="transmembrane region" description="Helical" evidence="22">
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        <location>
            <position position="669"/>
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            <position position="1338"/>
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        <ligand>
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        <source>
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        <source>
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    </evidence>
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        <source>
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        <source>
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        <source>
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    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
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        <source>
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        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
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    </evidence>
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        <source>
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    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="PIRSR" id="PIRSR602077-1"/>
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        <source>
            <dbReference type="RuleBase" id="RU003808"/>
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    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="Coils"/>
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    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
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    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="SMART" id="SM01062"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030358.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2234" mass="252306" checksum="F7E512CC42A0DA5E" modified="2019-05-08" version="1">MARFGEAVAGRLGSGDASSEQNRIRQGPPAPSGATPGAFKQTKAQRARTMALYNPIPVRQNCFTVNRSLFLFGEDNMVRKYAKKLVPSLFCHFCSLTVPSTPFEYMILATIIANCIVLALEQHLPEDDKTPMSRRLEKTEPYFIGIFCFEAGIKIVALGFVFHKGSYLRNGWNVMDFIVVLSGILATAGTHFNTHVDLRTLRAVRVLRPLKLVSGIPSLQIVLKSIMKAMVPLLQIGLLLFFAILMFAIIGLEFYSGKLHRACYTNNSGELEELDPPHPCGVQGCPLGYECKEWIGPNDGITQFDNILFAVLTVFQCITMEGWTTVLYNTNDALGATWNWLYFIPLIIIGSFFVLNLVLGVLSGEFAKERERVENRRAFMKLRRQQQIERELNGYRAWIDKAEEVMLAEENKNAGTSALEVLRRATIKRNRTEAVNRDSSEEHCVDISSVGTPLARGSIRSAKVDGASYFRHKERLLRISVRHMVKSQVFYWTVLSLVALNTACVAIVHHNQPPWLTHLLYYAEFLFLGLFLLEMSLKMYGMGPRLYFHSSFNCFDCGVTVGSIFEVVWAIFRPGTSFGISVLRALRLLRIFKITKYWASLRNLVVSLMSSMKSIISLLFLLFLFIVVFALLGMQLFGGRFNFMDGTPSANFDTFPAAIMTVFQILTGEDWNEVMYNGIRSQGGVRSGMWSSIYFIILTLFGNYTLLNVFLAIAVDNLANAQELTKDEQEEEEAFNQKHALQKAKEVITLCIHHVSNCFCLRHHMSVWEQRTSQLRRHMQMSSQEALNKDAPPLINPHASIFRRKIPGENVTLEKGEEGHGGSTEHQQVECLDQPIAGTNSCPSTKEDQKSPSPRAKRDREQWHQKSCHGNCDLTEQDGAGRGGGGGGAGIEDRARLRQSQRRSRHRRVRTEGKEPSGALEGRSASQEAGLEEANPTEGEQARNEEGKEDTNQEEPLAGEELNRCSSKLSVSPCDAAFDGAMQEVSPLKVPHELNEEKNTSLTEQDCSGLDTSDQALLGSLQMEMSRTVSRSEPELSSVTANTEKATESTAIMIDVQDSTVVQISNKTDGEASPLKEAETKEDEGKEKKKRKKEKSSETGKAMVPHSSMFIFSTTNPVRRACHYIVNLRYFEMCILLVIAASSIALAAEDPVLTNSDRNKVIVFTACVFILSLQMIDQGLILQDGSYFRDLWNILDFIVVVGALVAFALATNKGRDIKTIKSLRVLRVLRPLKTIKRLPKLKAVFDCVVTSLKNVFNILIVYKLFMFIFAVIAVQLFKGKFFYCTDSSKDTEKDCIGNYVDHEKNKMEVKCREWKRHEFHYDNIIWALLTLFTVSTGEGWPQVLQHSVDVTEEDRGPSRSNRMEMSIFYVVYFVVFPFFFVNIFVALIIITFQEQGDKMMEECSLEKNERACIDFAISAKPLTRYMPQNRHTFQYRVWHFVVSPSFEYTIMAMIALNTVVLMMKYYSAPYTYELALKYLNIAFTMVFSLECVLKIIAFGFLNYFRDTWNIFDFITVIGSITEIILTDTKLVNTSSFNMSFLKLFRAARLIKLLRQGYTIRILLWTFVQSFKALPYVCLLIAMLFFIYAIIGMQVFGNIKLDEESHINRHNNFRSFLASLMLLFRSATGEAWQEIMLSCLQGKGCEPDTTATSGQHENEHCGTDLAYVYFVSFIFFCSFLMLNLFVAVIMDNFEYLTRDSSILGPHHLDEFVRIWAEYDRAACGRIPYKDMYKLVRVISPPLGLGKNCPYRVACKRLVLMNMPVAEDMTVHFTSTLMALIRTALDIKIAKGGADWQQLDSELQKEILAIWPHLSQKMLDVLVPLPKTSDLTVGKIYAAMMIMDYYKQSKAKKQKQQLEEQKNAPMFQRMEPSSLPPEILSNAKALPYLQQDTLSGLSSRSGYPSLSPLSPQEIFQLACMDSTHGQFQEQQSLVVTDTSSMRRSFSTIRDKRPNSSWLDEFSMERSSENTYKSRRRSYHSSLQLSARRLNADSGHRSETHRSGGRERGRSKERKHLLSPDISRCNSEERSPQADGESPVRRQSRSPSEGRSQTPNRQGTGSLSESSIPSISDTSTPRRGRRQLPPVPPKPRPLLSYASMMRHPGDTSPQPDESEAGSPLFSQALETNDTCLTESSSSPQGKQSRHSTPQRYISEPYLALHDDSHASDCGEEETLTFEAAVATSLGRSNTIGSAPPLRHGWQMPNGHYRRRRRGAAQGVMCGASIDVLSDTEEDDKC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQH4</accession>
    <name>A0A452IQH4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">DNA-directed RNA polymerases I, II, and III subunit RPABC2</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="8">DNA-directed RNA polymerase II subunit F</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="7">RPB6 homolog</fullName>
        </alternativeName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">DNA-dependent RNA polymerase catalyzes the transcription of DNA into RNA using the four ribonucleoside triphosphates as substrates. Common component of RNA polymerases I, II, and III which synthesize ribosomal RNA precursors, mRNA precursors and many functional non-coding RNAs, and small RNAs, such as 5S rRNA and tRNAs, respectively. Pol II is the central component of the basal RNA polymerase II transcription machinery. Pols are composed of mobile elements that move relative to each other. In Pol II, POLR2F/RPABC2 is part of the clamp element and together with parts of POLR2A/RPB1 and POLR2B/RPB2 forms a pocket to which the POLR2D/RPB4-POLR2G/RPB7 subcomplex binds.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the archaeal Rpo6/eukaryotic RPB6 RNA polymerase subunit family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQH4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034234.1">
        <property type="protein sequence ID" value="ENSGAGP00000030156.1"/>
        <property type="gene ID" value="ENSGAGG00000021757.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005736">
        <property type="term" value="C:RNA polymerase I complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005665">
        <property type="term" value="C:RNA polymerase II, core complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005666">
        <property type="term" value="C:RNA polymerase III complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003899">
        <property type="term" value="F:DNA-directed RNA polymerase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006360">
        <property type="term" value="P:transcription by RNA polymerase I"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006366">
        <property type="term" value="P:transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042797">
        <property type="term" value="P:tRNA transcription by RNA polymerase III"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.940.10:FF:000003">
        <property type="entry name" value="DNA-directed RNA polymerases I, II, and III subunit RPABC2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.940.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020708">
        <property type="entry name" value="DNA-dir_RNA_polK_14-18kDa_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006110">
        <property type="entry name" value="Pol_omega/Rpo6/RPB6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028363">
        <property type="entry name" value="RPB6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036161">
        <property type="entry name" value="RPB6/omega-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006111">
        <property type="entry name" value="Rpo6/Rpb6"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF002207">
        <property type="entry name" value="PRK01099.1-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF002208">
        <property type="entry name" value="PRK01099.1-3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47227">
        <property type="entry name" value="DNA-DIRECTED RNA POLYMERASE SUBUNIT K"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47227:SF5">
        <property type="entry name" value="DNA-DIRECTED RNA POLYMERASES I, II, AND III SUBUNIT RPABC2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01192">
        <property type="entry name" value="RNA_pol_Rpb6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF500154">
        <property type="entry name" value="RPB6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000778">
        <property type="entry name" value="RpoK/RPB6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01409">
        <property type="entry name" value="RNA_pol_Rpb6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF63562">
        <property type="entry name" value="RPB6/omega subunit-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01111">
        <property type="entry name" value="RNA_POL_K_14KD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0240">DNA-directed RNA polymerase</keyword>
    <keyword evidence="5" id="KW-0539">Nucleus</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0804">Transcription</keyword>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1"/>
            <end position="53"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="10">
        <location>
            <begin position="1"/>
            <end position="34"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="43"/>
            <end position="53"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00020808"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022478"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00025773"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00030456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00033085"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00044490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030156.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="127" mass="14509" checksum="2262B10D404ED38A" modified="2019-05-08" version="1">MSDNEDNFDGDDFDDVEEDEGLDDLENAEEEGQENVEILPSGERQQANQKRITTPYMTKYERARVLGTRALQIAMCAPVMVELEGETDPLLIAMKELKARKIPIIIRRYLPDGSYEDWGVDELIITD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQH7</accession>
    <name>A0A452IQH7_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="18">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQH7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034326.1">
        <property type="protein sequence ID" value="ENSGAGP00000030244.1"/>
        <property type="gene ID" value="ENSGAGG00000021690.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098794">
        <property type="term" value="C:postsynapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043083">
        <property type="term" value="C:synaptic cleft"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004930">
        <property type="term" value="F:G protein-coupled receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007189">
        <property type="term" value="P:adenylate cyclase-activating G protein-coupled receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007166">
        <property type="term" value="P:cell surface receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016525">
        <property type="term" value="P:negative regulation of angiogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016322">
        <property type="term" value="P:neuron remodeling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd15251">
        <property type="entry name" value="7tmB2_BAI_Adhesion_VII"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1070.10:FF:000016">
        <property type="entry name" value="Adhesion G protein-coupled receptor B2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000003">
        <property type="entry name" value="Adhesion G protein-coupled receptor B2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000004">
        <property type="entry name" value="Adhesion G protein-coupled receptor B2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000012">
        <property type="entry name" value="Adhesion G protein-coupled receptor B2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.1240.10:FF:000002">
        <property type="entry name" value="Adhesion G protein-coupled receptor B2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.220.50:FF:000004">
        <property type="entry name" value="Adhesion G protein-coupled receptor B3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.220.50">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.1240.10">
        <property type="entry name" value="GPCR, family 2, extracellular hormone receptor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1070.10">
        <property type="entry name" value="Rhodopsin 7-helix transmembrane proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.20.100.10">
        <property type="entry name" value="Thrombospondin type-1 (TSP1) repeat"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032471">
        <property type="entry name" value="AGRL2-4_GAIN_subdom_A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057244">
        <property type="entry name" value="GAIN_B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046338">
        <property type="entry name" value="GAIN_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017981">
        <property type="entry name" value="GPCR_2-like_7TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008077">
        <property type="entry name" value="GPCR_2_brain_angio_inhib"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036445">
        <property type="entry name" value="GPCR_2_extracell_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001879">
        <property type="entry name" value="GPCR_2_extracellular_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000832">
        <property type="entry name" value="GPCR_2_secretin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017983">
        <property type="entry name" value="GPCR_2_secretin-like_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000203">
        <property type="entry name" value="GPS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000884">
        <property type="entry name" value="TSP1_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036383">
        <property type="entry name" value="TSP1_rpt_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12011:SF40">
        <property type="entry name" value="ADHESION G PROTEIN-COUPLED RECEPTOR B3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12011">
        <property type="entry name" value="ADHESION G-PROTEIN COUPLED RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00002">
        <property type="entry name" value="7tm_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16489">
        <property type="entry name" value="GAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01825">
        <property type="entry name" value="GPS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02793">
        <property type="entry name" value="HRM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00090">
        <property type="entry name" value="TSP_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01694">
        <property type="entry name" value="BAIPRECURSOR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00249">
        <property type="entry name" value="GPCRSECRETIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00303">
        <property type="entry name" value="GPS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00008">
        <property type="entry name" value="HormR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00209">
        <property type="entry name" value="TSP1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF82895">
        <property type="entry name" value="TSP-1 type 1 repeat"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00650">
        <property type="entry name" value="G_PROTEIN_RECEP_F2_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50227">
        <property type="entry name" value="G_PROTEIN_RECEP_F2_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50261">
        <property type="entry name" value="G_PROTEIN_RECEP_F2_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50221">
        <property type="entry name" value="GAIN_B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50092">
        <property type="entry name" value="TSP1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="10" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="8" id="KW-0297">G-protein coupled receptor</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9 14" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="11" id="KW-0675">Receptor</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <keyword evidence="13" id="KW-0807">Transducer</keyword>
    <keyword evidence="4 14" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 14" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="555"/>
            <end position="577"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="589"/>
            <end position="607"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="619"/>
            <end position="637"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="658"/>
            <end position="677"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="697"/>
            <end position="720"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="741"/>
            <end position="763"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="769"/>
            <end position="793"/>
        </location>
    </feature>
    <feature type="domain" description="G-protein coupled receptors family 2 profile 1" evidence="16">
        <location>
            <begin position="208"/>
            <end position="278"/>
        </location>
    </feature>
    <feature type="domain" description="GAIN-B" evidence="15">
        <location>
            <begin position="387"/>
            <end position="545"/>
        </location>
    </feature>
    <feature type="domain" description="G-protein coupled receptors family 2 profile 2" evidence="17">
        <location>
            <begin position="553"/>
            <end position="794"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023040"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023224"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50227"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50261"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030244.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1060" mass="118413" checksum="F8039D11337B1AF3" modified="2019-05-08" version="1">MEVPYRQCSRENEDKTRKSVTADTQFHLGRIPFVFIFNLLCYRCPVCFVTVHGVWEEWSPWSLCSFTCGRGQRTRTRSCIPPQYGGRSCDGPETQHKPCNIALCPVDGQWQEWSSWSQCSVTCSNGTQQRSRQCTAAAHGGSECRGPWAESRECHNPECTANGQWNQWGHWSGCSKSCDGGWERRIRICQGAAVTGQQCEGTGEEIRRCSEQRCPAPYEICPEDYLLSMVWKRTPAGDMAFNRCPLNATGTASRRCSLSVHGVAFWEHPSFARCISNEYRHLQHSIKEHLAKGQRMLAGDGMSQVTKTLLDLTQRKNFYAGDLLISVEILRNVTDTFKRASYIPASDGVQNFFQIVSNLLDEENKEKWEDAQQVRFASVAIYIQKLPAASVLTDINFPMKGRKGMVDWARNSEDRVVIPKNIFTPMSSELDESTVFVLGAVLYKNLELILPTLRNFTVVNSKIIVVTIRPEPKTADSFLEIELAHLANGTLNPYCVLWDDSRMNDSLGTWSTQGCKTVLTDASHTKCLCDRLSTFAILAQFSVLSIMESSGAPSVTLIVGSGLSCLALITLAVVYAALWRYIRSERSIILINFCLSIISSNILILVGQTQTHNKGICTTTTAFLHFFFLASFCWVLTEAWQSYMAVTGKIRTQLIRKRFLCLGWGLPALVVAISIGFTKTKGYGTSHYCWLSLEGGLLYAFVGPAAAVVLVNMVIGILVFNKLVSRDGILDKKLKHRAGASLWSSCVVLPLLALTWMSAVLAMTDKRSILFQILFAVFDSLQGFVIVMVHCILRREVQDAFRCRLRNCQDPINADSSSSFPNGHAQIMTDFEKDVDIACRSVLHKDIGPCRAATITGTLSRISLNDDEEEKGANPEGMSYSTLPGSIISKVIIQQPAGLHMPMSMSELANQCLKKDNSELRRTVYLCTDDNLRGADMDIVNPQERMMESDYIVMPRGSVNTQQSLKDESKLNIGMDTLPHERLLHYKVNPEFNMNPSVMDQFNINLDQHLPTQEHMQNLPFEPRTAVKNFIASELDDNAGLARSETGSTISMSSLEVSDA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQH8</accession>
    <name>A0A452IQH8_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="8">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the BRMS1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQH8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034240.1">
        <property type="protein sequence ID" value="ENSGAGP00000030161.1"/>
        <property type="gene ID" value="ENSGAGG00000021762.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005654">
        <property type="term" value="C:nucleoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010468">
        <property type="term" value="P:regulation of gene expression"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.1500:FF:000002">
        <property type="entry name" value="breast cancer metastasis-suppressor 1-like protein-A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.1500">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013907">
        <property type="entry name" value="Sds3"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21964">
        <property type="entry name" value="BREAST CANCER METASTASIS-SUPPRESSOR 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08598">
        <property type="entry name" value="Sds3"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Sds3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0539">Nucleus</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0678">Repressor</keyword>
    <keyword evidence="4" id="KW-0804">Transcription</keyword>
    <keyword evidence="3" id="KW-0805">Transcription regulation</keyword>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1"/>
            <end position="54"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="1"/>
            <end position="15"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="7">
        <location>
            <begin position="16"/>
            <end position="51"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
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            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022491"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00038256"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030161.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="305" mass="35816" checksum="635D2A8CABC9841E" modified="2019-05-08" version="1">MPVHSREKKENNHDEMEVDYGENEGSSSEEEDTESSSVSEEGDSSEMDDEDCERRRMECLDEMSNLEKQFTDLKDQLYKERLSQVDAKLQEVIAGKAQEYLEPLAALQENMQIRTKVAGIYRELCLESVKNKYECEIQASRQHCESEKLLLYDTVQSELEEKIRRLEEDRHSIDITSELWNDELQSRKKRKDPFSPDKKKPVVISGPYIVYMLQDLDILEDWTTIRKAMATLGPHRVKPEPPIKLEKHLHSARSEEGRLYYDGEWYGRGQTICIDKKDECPTRTYINRHETNLWKLLVLSHFALH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQI9</accession>
    <name>A0A452IQI9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">Nucleolar 27S pre-rRNA processing Urb2/Npa2 C-terminal domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000034470.1">
        <property type="protein sequence ID" value="ENSGAGP00000030373.1"/>
        <property type="gene ID" value="ENSGAGG00000021879.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005730">
        <property type="term" value="C:nucleolus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042254">
        <property type="term" value="P:ribosome biogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052609">
        <property type="entry name" value="Ribosome_Biogenesis_Reg"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018849">
        <property type="entry name" value="Urb2/Npa2_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15682">
        <property type="entry name" value="UNHEALTHY RIBOSOME BIOGENESIS PROTEIN 2 HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15682:SF2">
        <property type="entry name" value="UNHEALTHY RIBOSOME BIOGENESIS PROTEIN 2 HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10441">
        <property type="entry name" value="Urb2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Nucleolar 27S pre-rRNA processing Urb2/Npa2 C-terminal" evidence="1">
        <location>
            <begin position="1349"/>
            <end position="1468"/>
        </location>
    </feature>
    <evidence type="ECO:0000259" key="1">
        <source>
            <dbReference type="Pfam" id="PF10441"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030373.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1494" mass="168579" checksum="1DC3BE330E1C5D96" modified="2019-05-08" version="1">MAAIYSGIHLKLKSAKTSWEDKIKLAHFAWISHQCFLPNKEQVLLDWVSHALVSYYNKKLELEDEIVEKLWTYLDNILHSRKLQNLLKDGKTINLSFLTAQVINERISETCVQKSHGNVGTVLSCCRGILSTPPLAIIYTAKCELMVNLLSKLCWLACQLLTLEDAVTSQLFDVLQLTFAQYLLIQRQQINPNRVFGQVTSHLFQACLLLRHLLMARVWTPADDGRVRQHLSREIRNQVEAVLQAGIFQPELLSSYREELLSERQPQEKKKRALKTILLPVNTVLSKLCDASFCEPTLQVIIVANSVPLLYKLFLDSYCKAENHMVCFHMFTRLFGCLRISHLQEDLWQETLSPVDWSTELLAVEQLLNLVLSNDIYNVAADRIRHKETQFQFYRKLAEMLVNHSQASIPAWFRCLKALISLNHLIVEPDLDDLVASAWIDAEVSEPRTKKPQETLISAMFQTYTKLRQFPKLFEEVLSVICRPAADELRLPVLSAGLTEKLGECLLELPPSQILDILFLILEKCQTRIIPDVKDDSDMALKLLSVSSVLHTFLFNMKSLDNVTPLPVVLRTQSLLDKMQREVIQPLLGLLKEHWAEENKPGLWLEKVSDSALLLVYTWVELDTLLSINCSKYVSRLAEAAGGVTECAAESWDFSALLPGVDRQCWKRVVRLSSSFCSASKYCLELLMLQKIKKILMQTSFRTEADLHTLQRAAAFILHSGRSSMSMEESEPWNGNISAINALTYPAAHWYLVVSNLTILIPHLSLKDVEYVADVLLKILSSTKALGVSADQDSFITAEKVSKALLHSTFLPEIKVLHCAFITSVIQQCASVLCSATQGVADQPIQQLSADNIPWHEEVLSSCKMVGLVEAQSKNNQSKDEAFHCWTTMEKIAQHILLLTRAGSPITLEEDQIEGFLHLLEIISALKLDSLFPSDHARCFLLLLSLVANTRANVSCSKLLSLKFLITCFHLLTCLQTGRSVNSIFKVFHASDALEAVMTALLAASKCFADVLTTPDWAEFLQVIQTFLECFLQMIVEKSQSLKLNLEKFTFFLASCKPYADADRGKPWNTAVSQLLLVPLTTLCHVLTLYLQQQSEKKQKLTEMLSVLLEPTVLQTGTAIQLCLRNLTKDQPLPLAFIPSVTTLLKADLSYLSMGGFMKMEGEQENPKDPGKYREYSHTELYQKFYIQILRELTSAGGHLQFLHSALQFLTVFCSVTDLYPGEETALTVFHAVKKLLAGPGTTAQVIQDLEMQLTELMTQLVESCTTEEFCVMMRLVLQGLEVSNIWKQNPKEVLSAVTLIKVLLSCPLTGEKERAFWFTSPQIITALVIQTKEVCRDQLLISTIVVPILETVAVLLRQGEENLSNPHHVTLAFSILLTVPLDHLKMEEYGSIFLGIHEVLFSILQCHPKVMLKAAPSFLNSFNRLVVSVMHEGRQKGDRGNTDEFGVVLKCAHLIERMYSHIAAKTEEFTVFSVFIVAQYVIELQKVIHSHPP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQK4</accession>
    <name>A0A452IQK4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Cytochrome c domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the TPPP family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQK4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034328.1">
        <property type="protein sequence ID" value="ENSGAGP00000030246.1"/>
        <property type="gene ID" value="ENSGAGG00000021800.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000034330.1">
        <property type="protein sequence ID" value="ENSGAGP00000030248.1"/>
        <property type="gene ID" value="ENSGAGG00000021800.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005874">
        <property type="term" value="C:microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009055">
        <property type="term" value="F:electron transfer activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0020037">
        <property type="term" value="F:heme binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015631">
        <property type="term" value="F:tubulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001578">
        <property type="term" value="P:microtubule bundle formation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046785">
        <property type="term" value="P:microtubule polymerization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032273">
        <property type="term" value="P:positive regulation of protein polymerization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.238.10:FF:000057">
        <property type="entry name" value="Tubulin polymerization-promoting protein family member 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.238.10">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009056">
        <property type="entry name" value="Cyt_c-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011992">
        <property type="entry name" value="EF-hand-dom_pair"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008907">
        <property type="entry name" value="TPP/p25"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12932">
        <property type="entry name" value="P25 ALPHA-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12932:SF18">
        <property type="entry name" value="TUBULIN POLYMERIZATION-PROMOTING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05517">
        <property type="entry name" value="p25-alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47473">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51007">
        <property type="entry name" value="CYTC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0349">Heme</keyword>
    <keyword evidence="5 6" id="KW-0408">Iron</keyword>
    <keyword evidence="4 6" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Cytochrome c" evidence="8">
        <location>
            <begin position="73"/>
            <end position="158"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1"/>
            <end position="51"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="175"/>
            <end position="229"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="14"/>
            <end position="26"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="33"/>
            <end position="43"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="185"/>
            <end position="205"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="219"/>
            <end position="229"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010994"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00433"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51007"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030248.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="229" mass="25102" checksum="654AA118D9DB46F1" modified="2019-05-08" version="1">MISGSALIAMMSDNKTKSTKPADKTPPKSPADPTRERATKRLSYESNSSHEGAMAAELTALEEAFRKFAIHGDTRATGKEMHGKNWSKLCKDCHVIDGKNVTITDVDIVFSKIKGKSSRTITFDQFKEALQELSKKRFKDKSNEEAIQEIHKLIEGKAPIISGVTKTISSPTVSRLTDTSKFTGSHKERFDPSGKGRGKAGREDLVDASGYVPGYKHAGTYDHKVQGSK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQK5</accession>
    <name>A0A452IQK5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">USP6 N-terminal-like protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="8">Acts as a GTPase-activating protein for RAB5A and RAB43. Involved in receptor trafficking. In complex with EPS8 inhibits internalization of EGFR. Involved in retrograde transport from the endocytic pathway to the Golgi apparatus. Involved in the transport of Shiga toxin from early and recycling endosomes to the trans-Golgi network. Required for structural integrity of the Golgi complex.</text>
    </comment>
    <comment type="subunit">
        <text evidence="9">Interacts with EPS8.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasmic vesicle</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Golgi apparatus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQK5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034267.1">
        <property type="protein sequence ID" value="ENSGAGP00000030187.1"/>
        <property type="gene ID" value="ENSGAGG00000021776.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031410">
        <property type="term" value="C:cytoplasmic vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031267">
        <property type="term" value="F:small GTPase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.8.270:FF:000010">
        <property type="entry name" value="Putative USP6 N-terminal-like protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.750:FF:000001">
        <property type="entry name" value="TBC1 domain family member 10A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.472.80:FF:000019">
        <property type="entry name" value="USP6 N-terminal like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.8.270">
        <property type="entry name" value="putative rabgap domain of human tbc1 domain family member 14 like domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.750">
        <property type="entry name" value="Ypt/Rab-GAP domain of gyp1p, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.472.80">
        <property type="entry name" value="Ypt/Rab-GAP domain of gyp1p, domain 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000195">
        <property type="entry name" value="Rab-GAP-TBC_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035969">
        <property type="entry name" value="Rab-GAP_TBC_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050302">
        <property type="entry name" value="Rab_GAP_TBC_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47219">
        <property type="entry name" value="RAB GTPASE-ACTIVATING PROTEIN 1-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47219:SF19">
        <property type="entry name" value="USP6 N-TERMINAL-LIKE PROTEIN ISOFORM X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00566">
        <property type="entry name" value="RabGAP-TBC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00164">
        <property type="entry name" value="TBC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47923">
        <property type="entry name" value="Ypt/Rab-GAP domain of gyp1p"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50086">
        <property type="entry name" value="TBC_RABGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0007">Acetylation</keyword>
    <keyword evidence="7" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="6" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="3" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Rab-GAP TBC" evidence="12">
        <location>
            <begin position="21"/>
            <end position="213"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="311"/>
            <end position="411"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="433"/>
            <end position="455"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="493"/>
            <end position="575"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="729"/>
            <end position="762"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="328"/>
            <end position="352"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="366"/>
            <end position="376"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="379"/>
            <end position="389"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="390"/>
            <end position="400"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="501"/>
            <end position="513"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="524"/>
            <end position="544"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004541"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004555"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00059926"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00064037"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00070172"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50086"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030187.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="762" mass="87586" checksum="8C62573DAB92039C" modified="2019-05-08" version="1">MLKSWEKYKNSEKFHRRIYKGIPLQLRGEVWCLLLDVPKMKEEMKDFYNKLKHRARGSSPDIRQIDLDVNRTYRDHIMFRDRYGVKQQSLFHVLAAYSIYNTEVGYCQGMSQITALLLMYMNEEDAFWALVKLLSGPKHAMHGFFVPGFPKLLRFQEHHEKILKKFLSKLKQHFDSQEISTSFYTMKWFFQCFLDRTPFTLTLRIWDIYILEGERVLTAMSYTILKLHRKHLLKLQMEELVEFLQETLAKDFFLEDDFVIEQLQNSMSELKRAKLDLPAAGKEDEFPKKPLGQIPLEPQPAMLNLTHMLNGQSNTSISRTQTRRRTERRLSPEKVQEDSPNNRRINSLEKKPSLKQQKPGPVVRSELSKNDVDIRKKPQSTVQDSSRQYNNAAANQNSNATSDTRREFVPKWNKPSDIKVIEKTTKLTVQVKGRSVNHAISGTPPSPGEPQPLNVKQNIWGLDADEGKRESNASQYDNIPGVDHENENPVEEMLERAHPQSPRNVTYSTSPRKQSAKIPGPLKISNNYAVSSQLRSPRYSSQSDGSDHRSNVKQPPPSYSNPPVYHGNTPKHASSSISVSFVPSQYNHGNRTNTSGRPYGSIFSTDISPDKPHMNSYPANRQNAISPPSSRIEVLPVDAYPGNSRSPSGFKFIIPPVDYLPDNRKRPDAAYMYRHEIYRQPWNQEANQSHLGNLQKYAACQPTPIQDHNLPTVSVDSPIRYRTSPTLEEHGSAQYQYSDPPAPAQHYRNQNDGMSMHESVLL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQK7</accession>
    <name>A0A452IQK7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Ig-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the MHC class II family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQK7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034208.1">
        <property type="protein sequence ID" value="ENSGAGP00000030130.1"/>
        <property type="gene ID" value="ENSGAGG00000021737.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042613">
        <property type="term" value="C:MHC class II protein complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019882">
        <property type="term" value="P:antigen processing and presentation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006955">
        <property type="term" value="P:immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.320.10">
        <property type="entry name" value="Class II Histocompatibility Antigen, M Beta Chain, Chain B, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003597">
        <property type="entry name" value="Ig_C1-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050160">
        <property type="entry name" value="MHC/Immunoglobulin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011162">
        <property type="entry name" value="MHC_I/II-like_Ag-recog"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014745">
        <property type="entry name" value="MHC_II_a/b_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001003">
        <property type="entry name" value="MHC_II_a_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19944:SF86">
        <property type="entry name" value="HLA CLASS II HISTOCOMPATIBILITY ANTIGEN, DR ALPHA CHAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19944">
        <property type="entry name" value="MHC CLASS II-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07654">
        <property type="entry name" value="C1-set"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00993">
        <property type="entry name" value="MHC_II_alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00407">
        <property type="entry name" value="IGc1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00920">
        <property type="entry name" value="MHC_II_alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54452">
        <property type="entry name" value="MHC antigen-recognition domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="7" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="5" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 8" id="KW-0732">Signal</keyword>
    <keyword evidence="3" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="8">
        <location>
            <begin position="1"/>
            <end position="18"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5045591903" description="Ig-like domain-containing protein" evidence="8">
        <location>
            <begin position="19"/>
            <end position="234"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="9">
        <location>
            <begin position="117"/>
            <end position="163"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007394"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030130.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="234" mass="26143" checksum="F1FFF35296FBDF3C" modified="2025-10-08" version="2">MPCHRLCVTLGQLLTSLDLPPLLGPEDNGPVAFYQLMDPTQQEAGEFMFEFDQDEIFHMNLERKTIWRLPNFGKFTSFEAQRALGNITVLKKNMEIMIRRSNRRQAQNSTVWEGGAPEMTVFPEDHVELGEPNILICFMDNFSPPVLSLTWLKNGTTPSASSPTCSSSRQSNFYDYQVEHGGLPEPFTKHWAQVPIPISETTEALGPAMGIIGIIVGTILIMKMNATGNLQGLL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQK8</accession>
    <name>A0A452IQK8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Monocarboxylate transporter 10</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="14">Solute carrier family 16 member 10</fullName>
        </alternativeName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="12">Sodium- and proton-independent thyroid hormones and aromatic acids transporter. Mediates both uptake and efflux of 3,5,3'-triiodothyronine (T3) and 3,5,3',5'-tetraiodothyronine (T4) with high affinity, suggesting a role in the homeostasis of thyroid hormone levels. Responsible for low affinity bidirectional transport of the aromatic amino acids, such as phenylalanine, tyrosine, tryptophan and L-3,4-dihydroxyphenylalanine (L-dopa). Plays an important role in homeostasis of aromatic amino acids.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="8">
            <text>3,3',5-triiodo-L-thyronine(out) = 3,3',5-triiodo-L-thyronine(in)</text>
            <dbReference type="Rhea" id="RHEA:71811"/>
            <dbReference type="ChEBI" id="CHEBI:533015"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="8">
            <dbReference type="Rhea" id="RHEA:71812"/>
        </physiologicalReaction>
        <physiologicalReaction direction="right-to-left" evidence="8">
            <dbReference type="Rhea" id="RHEA:71813"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>L-phenylalanine(in) = L-phenylalanine(out)</text>
            <dbReference type="Rhea" id="RHEA:27950"/>
            <dbReference type="ChEBI" id="CHEBI:58095"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="11">
            <dbReference type="Rhea" id="RHEA:27951"/>
        </physiologicalReaction>
        <physiologicalReaction direction="right-to-left" evidence="11">
            <dbReference type="Rhea" id="RHEA:27952"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="10">
            <text>L-thyroxine(out) = L-thyroxine(in)</text>
            <dbReference type="Rhea" id="RHEA:71819"/>
            <dbReference type="ChEBI" id="CHEBI:58448"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="10">
            <dbReference type="Rhea" id="RHEA:71820"/>
        </physiologicalReaction>
        <physiologicalReaction direction="right-to-left" evidence="10">
            <dbReference type="Rhea" id="RHEA:71821"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="7">
            <text>L-tryptophan(in) = L-tryptophan(out)</text>
            <dbReference type="Rhea" id="RHEA:70947"/>
            <dbReference type="ChEBI" id="CHEBI:57912"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="7">
            <dbReference type="Rhea" id="RHEA:70948"/>
        </physiologicalReaction>
        <physiologicalReaction direction="right-to-left" evidence="7">
            <dbReference type="Rhea" id="RHEA:70949"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="9">
            <text>L-tyrosine(in) = L-tyrosine(out)</text>
            <dbReference type="Rhea" id="RHEA:68572"/>
            <dbReference type="ChEBI" id="CHEBI:58315"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="9">
            <dbReference type="Rhea" id="RHEA:68573"/>
        </physiologicalReaction>
        <physiologicalReaction direction="right-to-left" evidence="9">
            <dbReference type="Rhea" id="RHEA:68574"/>
        </physiologicalReaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Basolateral cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the major facilitator superfamily. Monocarboxylate porter (TC 2.A.1.13) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQK8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034488.1">
        <property type="protein sequence ID" value="ENSGAGP00000030390.1"/>
        <property type="gene ID" value="ENSGAGG00000021892.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016323">
        <property type="term" value="C:basolateral plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015192">
        <property type="term" value="F:L-phenylalanine transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015196">
        <property type="term" value="F:L-tryptophan transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005302">
        <property type="term" value="F:L-tyrosine transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015349">
        <property type="term" value="F:thyroid hormone transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1250.20:FF:001016">
        <property type="entry name" value="Solute carrier family 16 member 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1250.20">
        <property type="entry name" value="MFS general substrate transporter like domains"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011701">
        <property type="entry name" value="MFS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036259">
        <property type="entry name" value="MFS_trans_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050327">
        <property type="entry name" value="Proton-linked_MCT"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11360">
        <property type="entry name" value="MONOCARBOXYLATE TRANSPORTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11360:SF123">
        <property type="entry name" value="MONOCARBOXYLATE TRANSPORTER 8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07690">
        <property type="entry name" value="MFS_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103473">
        <property type="entry name" value="MFS general substrate transporter"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="6 16" id="KW-0472">Membrane</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 16" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 16" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="44"/>
            <end position="71"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="91"/>
            <end position="108"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="145"/>
            <end position="168"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="175"/>
            <end position="197"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="209"/>
            <end position="231"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="272"/>
            <end position="288"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="308"/>
            <end position="326"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="338"/>
            <end position="357"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="363"/>
            <end position="387"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="399"/>
            <end position="421"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="427"/>
            <end position="451"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="1"/>
            <end position="26"/>
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    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004554"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00006727"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
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    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00050480"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00050518"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00051690"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00052346"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00058829"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00073865"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00078724"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030390.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="486" mass="53164" checksum="9C6E77A0E1A213AE" modified="2019-05-08" version="1">SWEEDPPPPPRENQGPGTESDPPGGAGLRCRGLDPAGLVYRGGFGWIVVFAATWCNGSIFGIQNSFGILYVMLQSDLGGGEKDPTLEFKTALVGALAMGMIFFCSPIVSIFTDRIGCRTTAASGAAIAFIGLLSSSFTKSLEVRYFTYGILFGCGSSFAFQPSLVILGHYFKHRLGLANGIVAGGSCLFSMSLPFFLKTIGSTIGLAHTFQVLSAFMFIQILLSLTFRPILPPSCDSHHNRQDKLASRNMKQQCGSQMRKYFNMRVFRRQTYRIWAFGIATAVLGYFVPYVHLIKYVEEEFKEKEKEWILLVCLGATSGLGRLVSGRIGDCIPGLKKIFLQVASFMLLGLMCMMIPQCHAFEGVIVVCLFLGLCDGFFITIMAPIAFELVGPMQASQAIGYLLGLMAIPMTAGPPIAGYLHDYFGNYYLAFYLAGVPPIIGGLVLSVVPLIHQRMLKKQQMDSGKDKMLVSETVMNVHLDLGEKPQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQL0</accession>
    <name>A0A452IQL0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Palladin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="19">Interacts with EPS8. Interacts with LASP1. Interacts with VASP. Interacts with ACTN. Interacts with SORBS2. Interacts with PFN1. Interacts with LPP. Interacts with SPIN90. Interacts with SRC. Interacts with EZR. Interacts with RAI14.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="4">Cell junction</location>
            <location evidence="4">Focal adhesion</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="6">Cell projection</location>
            <location evidence="6">Axon</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="8">Cell projection</location>
            <location evidence="8">Growth cone</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="7">Cell projection</location>
            <location evidence="7">Lamellipodium</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cell projection</location>
            <location evidence="1">Podosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="5">Cell projection</location>
            <location evidence="5">Ruffle</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Myofibril</location>
            <location evidence="2">Sarcomere</location>
            <location evidence="2">Z line</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="18">Belongs to the myotilin/palladin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQL0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034273.1">
        <property type="protein sequence ID" value="ENSGAGP00000030192.1"/>
        <property type="gene ID" value="ENSGAGG00000021781.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005925">
        <property type="term" value="C:focal adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
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    <dbReference type="GO" id="GO:0030426">
        <property type="term" value="C:growth cone"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
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    <dbReference type="GO" id="GO:0030027">
        <property type="term" value="C:lamellipodium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002102">
        <property type="term" value="C:podosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001726">
        <property type="term" value="C:ruffle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
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    <dbReference type="GO" id="GO:0001725">
        <property type="term" value="C:stress fiber"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
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    <dbReference type="GO" id="GO:0030018">
        <property type="term" value="C:Z disc"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
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    <dbReference type="GO" id="GO:0003779">
        <property type="term" value="F:actin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
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    <dbReference type="GO" id="GO:0004672">
        <property type="term" value="F:protein kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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        <property type="entry name" value="IgI_Myotilin_C"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="palladin isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="palladin isoform X2"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="palladin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="3"/>
    </dbReference>
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        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
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        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013098">
        <property type="entry name" value="Ig_I-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47633">
        <property type="entry name" value="IMMUNOGLOBULIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47633:SF10">
        <property type="entry name" value="PALLADIN, CYTOSKELETAL ASSOCIATED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07679">
        <property type="entry name" value="I-set"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="14" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="12" id="KW-0965">Cell junction</keyword>
    <keyword evidence="16" id="KW-0966">Cell projection</keyword>
    <keyword evidence="9" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="15" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="13" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="17" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="10" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="11" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="Ig-like" evidence="22">
        <location>
            <begin position="369"/>
            <end position="453"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="22">
        <location>
            <begin position="503"/>
            <end position="588"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="22">
        <location>
            <begin position="601"/>
            <end position="692"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="21">
        <location>
            <begin position="30"/>
            <end position="93"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="21">
        <location>
            <begin position="110"/>
            <end position="233"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="21">
        <location>
            <begin position="343"/>
            <end position="365"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="21">
        <location>
            <begin position="464"/>
            <end position="493"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="21">
        <location>
            <begin position="49"/>
            <end position="61"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="21">
        <location>
            <begin position="64"/>
            <end position="93"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="21">
        <location>
            <begin position="116"/>
            <end position="135"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="21">
        <location>
            <begin position="164"/>
            <end position="174"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="21">
        <location>
            <begin position="195"/>
            <end position="206"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="21">
        <location>
            <begin position="208"/>
            <end position="217"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="21">
        <location>
            <begin position="343"/>
            <end position="361"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004188"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004216"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004466"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00004489"/>
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    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00004510"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
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            <dbReference type="ARBA" id="ARBA00004624"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
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    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00061540"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00065150"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00072991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030192.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="751" mass="83041" checksum="8D3C7207107BE8D9" modified="2019-05-08" version="1">SATNKVCTVGGVTLISRPKDPLKLQQLQNQIRLEQEGGQRYHHHQQHQNPPPSPPFPPPPSFQELLSSQSVTSPVQMSILNSHTSPTMQSSSTFSYARPKQFIAAQNISPASGYVTPSSGSSTSSLPSPMSPTASQKQFGRAPVPPFSQPFAPEGEYLWSPSSSSPPPPPPPVFSPTATFSVSDSFPPPPPPLPSSVSSPSRSLSPTARFSNSSQSPAAFLSSMLPSQPPPISVNALGLPKGVMPPGFPKKTGRTARIASDEEIQGTKDAVIQDLERKLRFKEELLNNGQPRLTYEEKMARRLLGADNAATVFNIQEPDEEPTTQEYKVSSFEQRLISEIEYRLERSPVEESDDEVQHGDEPTDNSVAPYFEMKLKHYKIFEGTSVTFTCRVAGNPKPKIYWFKDGKQISKRSDHYRIQREPDGTCSLHTTASTLDDDGNYTIMAANPQGRISCTGRLMVQAVNQRGRSPRTPPSQPHIRRPRSRSRDSGDENEPIQERFFRPHFLQAPGDLIVQEGKLCRMDCKVSGLPTPDLSWQLNGRPIRPDSSHKMLVRENGVHSLIIEPVTARDSGIYTCIATNRAGQNTFSLELIVAAKEEHKPPVFIEKLQNTGVADGYPVRLECRVSGVPYPQIFWKKENESLTYNTERVSMHQDNHGYICLLIQGATKEDAGWYTVSAKNEAGIVSCTARLDVYTQWQQQPQSTKPKKVRPSASRYAALSDQGLDIKAAFQPEANPVHLTLQCGLVESDDL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQL6</accession>
    <name>A0A452IQL6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Cadherin domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">Cadherins are calcium-dependent cell adhesion proteins.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 14">Cell membrane</location>
            <topology evidence="1 14">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQL6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034278.1">
        <property type="protein sequence ID" value="ENSGAGP00000030197.1"/>
        <property type="gene ID" value="ENSGAGG00000021782.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005912">
        <property type="term" value="C:adherens junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016342">
        <property type="term" value="C:catenin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008013">
        <property type="term" value="F:beta-catenin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045296">
        <property type="term" value="F:cadherin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034332">
        <property type="term" value="P:adherens junction organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016339">
        <property type="term" value="P:calcium-dependent cell-cell adhesion via plasma membrane cell adhesion molecules"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000902">
        <property type="term" value="P:cell morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0044331">
        <property type="term" value="P:cell-cell adhesion mediated by cadherin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007043">
        <property type="term" value="P:cell-cell junction assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007156">
        <property type="term" value="P:homophilic cell adhesion via plasma membrane adhesion molecules"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd11304">
        <property type="entry name" value="Cadherin_repeat"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.900.10:FF:000001">
        <property type="entry name" value="Cadherin 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.60:FF:000009">
        <property type="entry name" value="Cadherin 24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.60:FF:000012">
        <property type="entry name" value="Cadherin 24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.60:FF:000017">
        <property type="entry name" value="Cadherin 24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.60:FF:000014">
        <property type="entry name" value="Cadherin 8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.60">
        <property type="entry name" value="Cadherins"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.900.10">
        <property type="entry name" value="TCF3-CBD (Catenin binding domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039808">
        <property type="entry name" value="Cadherin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002126">
        <property type="entry name" value="Cadherin-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015919">
        <property type="entry name" value="Cadherin-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020894">
        <property type="entry name" value="Cadherin_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000233">
        <property type="entry name" value="Cadherin_Y-type_LIR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027397">
        <property type="entry name" value="Catenin-bd_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24027:SF323">
        <property type="entry name" value="CADHERIN-19"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24027">
        <property type="entry name" value="CADHERIN-23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01049">
        <property type="entry name" value="CADH_Y-type_LIR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00028">
        <property type="entry name" value="Cadherin"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00205">
        <property type="entry name" value="CADHERIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00112">
        <property type="entry name" value="CA"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49313">
        <property type="entry name" value="Cadherin-like"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00232">
        <property type="entry name" value="CADHERIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50268">
        <property type="entry name" value="CADHERIN_2"/>
        <property type="match status" value="5"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8 13 14" id="KW-0106">Calcium</keyword>
    <keyword evidence="9 14" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="3" id="KW-0165">Cleavage on pair of basic residues</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11 16" id="KW-0472">Membrane</keyword>
    <keyword evidence="5" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="6 17" id="KW-0732">Signal</keyword>
    <keyword evidence="4 14 16" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="10 16" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="17">
        <location>
            <begin position="1"/>
            <end position="21"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019286044" description="Cadherin domain-containing protein" evidence="17">
        <location>
            <begin position="22"/>
            <end position="710"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="548"/>
            <end position="569"/>
        </location>
    </feature>
    <feature type="domain" description="Cadherin" evidence="18">
        <location>
            <begin position="69"/>
            <end position="149"/>
        </location>
    </feature>
    <feature type="domain" description="Cadherin" evidence="18">
        <location>
            <begin position="150"/>
            <end position="204"/>
        </location>
    </feature>
    <feature type="domain" description="Cadherin" evidence="18">
        <location>
            <begin position="203"/>
            <end position="316"/>
        </location>
    </feature>
    <feature type="domain" description="Cadherin" evidence="18">
        <location>
            <begin position="317"/>
            <end position="421"/>
        </location>
    </feature>
    <feature type="domain" description="Cadherin" evidence="18">
        <location>
            <begin position="421"/>
            <end position="535"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022685"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="RuleBase" id="RU003318"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="RuleBase" id="RU004357"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50268"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030197.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="710" mass="79833" checksum="454D1755F47EAB7B" modified="2019-05-08" version="1">MNCYAWLSLLLTLSHLWPCLPSMQNHNTHKMNHPFATLQRVKRGWVWSQFFVLEEQILKEPLCVGQLKSDSDKHDGTFKYILTGDGAGSIFTIDEYTGTIHVTQKLDREEKPFYTLRAQAINRNTQLPVEPESEFIIKVQDINDQEPKFLDGPYEATVPEMSPEGTSVIQVTATDGDDPAYGNSARLLYSILQGQPYFSVEPKTEFYYMNISEAAPVGSTIGKIMAEDSDIGENAAMNYVIEGEESLVFDIITNNETQEGIVILKERVDFESKRRYSIRVKAINRQMDDHFMKEGPFEDTTIIKISVEDADEPPVFTSQNYVMEIPEGATNGSFVGAITARDPDNANSPIRYSIVHSTYLKRLFSINAHSGTIIINKLLDRETAAWHNITVTATEMRNPEQVSEVTVYIQVLDVNDHAPEFPKHYEIYVCENARSGQLIQTISAVDKDDSVEGHHFYFTLAREATNNSNFTVRDNQDNTAGIFTAKNGFSRQEQFAFFLPVLIVDNGTPSLTSTNTLTVSVCDCDTEASAQSCRFGAFMFSMGLSIEALVGVLACILVILVFFLAMLVLRQRRKQSLFPDKPEEFRENIVRYDDEGGGEEDTEAFDISTLRTRTVMREHKSRRNITTEIQSLYRQSLQVGPDSAVFREFILEKLKEANADPTAPPYDSLQTYAFEGTGSLAGSLSSLGSSISHVDENYDYLTDGGWTLNN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQL7</accession>
    <name>A0A452IQL7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">CMP/dCMP-type deaminase domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the cytidine and deoxycytidylate deaminase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQL7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034218.1">
        <property type="protein sequence ID" value="ENSGAGP00000030140.1"/>
        <property type="gene ID" value="ENSGAGG00000021744.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004126">
        <property type="term" value="F:cytidine deaminase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016554">
        <property type="term" value="P:cytidine to uridine editing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.140.10">
        <property type="entry name" value="Cytidine Deaminase, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016192">
        <property type="entry name" value="APOBEC/CMP_deaminase_Zn-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050610">
        <property type="entry name" value="APOBEC_Cyt_Deaminase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002125">
        <property type="entry name" value="CMP_dCMP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016193">
        <property type="entry name" value="Cytidine_deaminase-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13857:SF26">
        <property type="entry name" value="C-U-EDITING ENZYME APOBEC-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13857">
        <property type="entry name" value="MRNA EDITING ENZYME"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18750">
        <property type="entry name" value="SNAD4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53927">
        <property type="entry name" value="Cytidine deaminase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00903">
        <property type="entry name" value="CYT_DCMP_DEAMINASES_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51747">
        <property type="entry name" value="CYT_DCMP_DEAMINASES_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="CMP/dCMP-type deaminase" evidence="6">
        <location>
            <begin position="63"/>
            <end position="166"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006576"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS51747"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030140.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="255" mass="30405" checksum="638C850960445CD1" modified="2019-05-08" version="1">MFLLSGIETEREKHTPVKQDTAWEAVVKAMATGWNRDHESRGITKGGKISQETFIDTYDPSVLRREQYMLYEIKWSNNKRPRQSCCHSTQTEHAEIYFLEDVFQKQRSDPSAHCFITWFVSWSPCGDCCKQIRDFLKEQPNVKLFIYVARIYWHKREINRQGLRNLMNIGVSIRVMYLPVYRYCWRTFVDKEDKYEDYWPRHFAPWIMLYSLELQSILQNIPSCLKISSFKNQSPVFSLCVDEKQKRALALANPY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQL9</accession>
    <name>A0A452IQL9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="7">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IQL9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034352.1">
        <property type="protein sequence ID" value="ENSGAGP00000030265.1"/>
        <property type="gene ID" value="ENSGAGG00000021815.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006897">
        <property type="term" value="P:endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016197">
        <property type="term" value="P:endosomal transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00052">
        <property type="entry name" value="EH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.238.10">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011992">
        <property type="entry name" value="EF-hand-dom_pair"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018247">
        <property type="entry name" value="EF_Hand_1_Ca_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002048">
        <property type="entry name" value="EF_hand_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000261">
        <property type="entry name" value="EH_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11216">
        <property type="entry name" value="EH DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11216:SF64">
        <property type="entry name" value="RALBP1-ASSOCIATED EPS DOMAIN-CONTAINING PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12763">
        <property type="entry name" value="EH"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00027">
        <property type="entry name" value="EH"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47473">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00018">
        <property type="entry name" value="EF_HAND_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50222">
        <property type="entry name" value="EF_HAND_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50031">
        <property type="entry name" value="EH"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0106">Calcium</keyword>
    <keyword evidence="3" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="1" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="EH" evidence="5">
        <location>
            <begin position="23"/>
            <end position="114"/>
        </location>
    </feature>
    <feature type="domain" description="EH" evidence="5">
        <location>
            <begin position="261"/>
            <end position="345"/>
        </location>
    </feature>
    <feature type="domain" description="EF-hand" evidence="6">
        <location>
            <begin position="294"/>
            <end position="329"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="149"/>
            <end position="225"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="456"/>
            <end position="502"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="518"/>
            <end position="544"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="617"/>
            <end position="647"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="3">
        <location>
            <begin position="650"/>
            <end position="688"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="149"/>
            <end position="164"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="4">
        <location>
            <begin position="168"/>
            <end position="178"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="214"/>
            <end position="225"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="4">
        <location>
            <begin position="522"/>
            <end position="532"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="534"/>
            <end position="544"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="619"/>
            <end position="630"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50031"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50222"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030265.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="698" mass="77212" checksum="AE0227C940422FC1" modified="2019-05-08" version="1">MEQGPGGAAAGGAGLLLPLSDTEQQRYSELFSLCCSPAPEGGSAPAPAAGGSKVGDLFRASQLPADTLHQITELCGAKRVGYFGPAQFYVALKLIAAAQSGLPVRIESIKSELPLPRFMALKNDIEMRYGNAAEILGVKFQIPHPILEKNSFRRSDEGDKKQETKSPPMSPMCSPPASPSNYQRIPLSYGYGKSRSKLEQPHGVSYEARHSTHQQDGPATGNYGSKSALVHSSLSRFLSVESEPQDSSNYPDDPWRITEEQREYYINQFKSLQPDLNSLISGSVAKNFFTKSKLPIPELSHVWELSDVDCDGALTLPEFCAAFHLIVARKNGYPLPDMLPETLLPDYLQPASLKPKRDCALLDSYSESLSVSQQTRDFNRTELETVCEDPTNSEPLILFDVATSIVEPTSVEEMPDKSALTQDANSDDNQALNISTTTKTIPKDFKPSQHLSVKTVVQESNTLKARPRSRSYSSTSIEEAMKKGEEPPMPPPRPQKSHSRASSLDLNKIFQQNTQAVKSGWLPPPPALPPRPFVSQQGPHHVTNTESVAQAKSSYVEFRLKEQTEQVSEIELHPQMNRAPSQTEESSPMRKEILLTQPPSKPIRRKFRSENQMLENQELTPTSSVTSSLPAVKPHLPAQNRQSSKQKRAIQTAIRKNKEANAVLARLNSELQQQLKEVHKERIALETQLEQLRPVTVL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQM0</accession>
    <name>A0A452IQM0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Glycine N-acyltransferase-like protein</fullName>
            <ecNumber evidence="3">2.3.1.-</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="3">Belongs to the glycine N-acyltransferase family.</text>
    </comment>
    <dbReference type="EC" id="2.3.1.-" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452IQM0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034503.1">
        <property type="protein sequence ID" value="ENSGAGP00000030405.1"/>
        <property type="gene ID" value="ENSGAGG00000021897.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0047961">
        <property type="term" value="F:glycine N-acyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.630.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016181">
        <property type="entry name" value="Acyl_CoA_acyltransferase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010313">
        <property type="entry name" value="Glycine_N-acyltransferase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013652">
        <property type="entry name" value="Glycine_N-acyltransferase_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015938">
        <property type="entry name" value="Glycine_N-acyltransferase_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000182">
        <property type="entry name" value="GNAT_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15298:SF1">
        <property type="entry name" value="GLYCINE N-ACYLTRANSFERASE-LIKE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15298">
        <property type="entry name" value="L-COA N-ACYLTRANSFERASE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08444">
        <property type="entry name" value="Gly_acyl_tr_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06021">
        <property type="entry name" value="Gly_acyl_tr_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55729">
        <property type="entry name" value="Acyl-CoA N-acyltransferases (Nat)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51186">
        <property type="entry name" value="GNAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 3" id="KW-0012">Acyltransferase</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1 3" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="N-acetyltransferase" evidence="4">
        <location>
            <begin position="157"/>
            <end position="286"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023315"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="RuleBase" id="RU368002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS51186"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030405.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="303" mass="33685" checksum="280C9BA2122A9444" modified="2019-05-08" version="1">RKLMGLSHKGIPSLTPTPPCPWQVYGTVMTINQGNPAAQEVLVDSWPEFKVVLTRPRREVALDDSDLYTNTYTVYYRDIGAYRALLASAGAVNWGQSFRILGNLFQLSRETSNHSVIVLTNLMVLGPCHDSEVPLSSLTFGALLCCFLPPPRLDPAMRLSSLDVSHADLLNETWSFGGNEKSRKYLATLIRHFPSICLLDAAGHPVSWIASYPFGAMGPAYTLHEHRQRGYMAMLGNLLAKRLHALGYPSYGYVDVENYPMQRLQESQGFQCLPNLWYFILHNPVPVKAPTLTPRPAPGTAPT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQM2</accession>
    <name>A0A452IQM2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">E3 ubiquitin-protein ligase Arkadia</fullName>
            <ecNumber evidence="6">2.3.2.27</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="20">RING finger protein 111</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="19">RING-type E3 ubiquitin transferase Arkadia</fullName>
        </alternativeName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>S-ubiquitinyl-[E2 ubiquitin-conjugating enzyme]-L-cysteine + [acceptor protein]-L-lysine = [E2 ubiquitin-conjugating enzyme]-L-cysteine + N(6)-ubiquitinyl-[acceptor protein]-L-lysine.</text>
            <dbReference type="EC" id="2.3.2.27"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="4">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
            <location evidence="2">PML body</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the Arkadia family.</text>
    </comment>
    <dbReference type="EC" id="2.3.2.27" evidence="6"/>
    <dbReference type="AlphaFoldDB" id="A0A452IQM2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034283.1">
        <property type="protein sequence ID" value="ENSGAGP00000030202.1"/>
        <property type="gene ID" value="ENSGAGG00000021784.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016605">
        <property type="term" value="C:PML body"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061630">
        <property type="term" value="F:ubiquitin protein ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006281">
        <property type="term" value="P:DNA repair"/>
        <property type="evidence" value="ECO:0007669"/>
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</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
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            <taxon>Vertebrata</taxon>
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            <taxon>Durocryptodira</taxon>
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        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
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        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
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        <citation type="submission" date="2025-08" db="UniProtKB">
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        <scope>IDENTIFICATION</scope>
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        <citation type="submission" date="2025-09" db="UniProtKB">
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        </source>
    </evidence>
    <sequence length="677" mass="74576" checksum="39DE4059C16C1086" modified="2019-05-08" version="1">MEQGPGGAAAGGAGLLLPLSDTEQQRYSELFSLCCSPAPEGGSAPAPAAGGSKVGDLFRASQLPADTLHQITELCGAKRVGYFGPAQFYVALKLIAAAQSGLPVRIESIKSELPLPRFMALKNDIEMRYGNAAEILGVKFQIPHPILEKNSFRRSDEGDKKETKSPPMSPMCSPPASPSNYQRIPLSYGYGKSRSKLEQPHGVSYEARHSTHQQDGPATGNYGSKSALVHSSLSRFLSVESEPQDSSNYPDDPWRITEEQREYYINQFKSLQPDLNSLISGSVAKNFFTKSKLPIPELSHVWELSDVDCDGALTLPEFCAAFHLIVARKNGYPLPDMLPETLLPDYLQPASLKPKRDCALLDSYSESLSVSQQTRDFNRTELETVCEDPTNSEPLILFDVATSIVEPTSVEEMPDKSALTQDANSDDNQALNISTTTKTIPKDFKPSQHLSVKTVVQESNTLKARPRSRSYSSTSIEEAMKKGEEPPMPPPRPQKSHSRASSLDLNKIFQQNTQAVKSGWLPPPPALPPRPFVSQQGPHHVTNTESVAQAKSSYVEFRLKEQEILLTQPPSKPIRRKFRSENQMLENQELTPTSSVTSSLPAVKPHLPAQNRQSSKQKRAIQTAIRKNKEANAVLARLNSELQQQLKVKNALNITITRLDAIGCFSPLAEDLVPLYR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQM6</accession>
    <name>A0A452IQM6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Peptidase S1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the peptidase S1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQM6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034288.1">
        <property type="protein sequence ID" value="ENSGAGP00000030207.1"/>
        <property type="gene ID" value="ENSGAGG00000021723.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030141">
        <property type="term" value="C:secretory granule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00190">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.10:FF:000077">
        <property type="entry name" value="Predicted protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.10.10">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009003">
        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043504">
        <property type="entry name" value="Peptidase_S1_PA_chymotrypsin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001314">
        <property type="entry name" value="Peptidase_S1A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001254">
        <property type="entry name" value="Trypsin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018114">
        <property type="entry name" value="TRYPSIN_HIS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033116">
        <property type="entry name" value="TRYPSIN_SER"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271:SF48">
        <property type="entry name" value="KALLIKREIN-14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271">
        <property type="entry name" value="KALLIKREIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00089">
        <property type="entry name" value="Trypsin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00722">
        <property type="entry name" value="CHYMOTRYPSIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00020">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50494">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50240">
        <property type="entry name" value="TRYPSIN_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00134">
        <property type="entry name" value="TRYPSIN_HIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00135">
        <property type="entry name" value="TRYPSIN_SER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3 6" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="2 6" id="KW-0645">Protease</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 6" id="KW-0720">Serine protease</keyword>
    <feature type="domain" description="Peptidase S1" evidence="7">
        <location>
            <begin position="17"/>
            <end position="240"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007664"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="RuleBase" id="RU363034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030207.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="242" mass="26664" checksum="73AFE2AD7A5B00D2" modified="2019-05-08" version="1">MENSVFYPILSSQNNRIVGGAPCIPHSMPWQAALFMGSQLNCGGTLIDRNWVVTAAHCHVQCSISVRLGEHNIRHLDWTEQLRLSEKTILHPGYNSASKDNDIMLIKLLTPANLNKNVQPLELPTSCPSPKEECVVSGWGTTKSPQLYFPPVLHCGKVSIFSQEQCRSIYPRYYTENMLCAGVLEGGIDSCQGDSGGPLLCNGKLQGIVSWGTQTCALPNKPGVYVAVCKYVDWIRETINNN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQM9</accession>
    <name>A0A452IQM9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IQM9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034233.1">
        <property type="protein sequence ID" value="ENSGAGP00000030155.1"/>
        <property type="gene ID" value="ENSGAGG00000021756.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007914">
        <property type="entry name" value="UPF0193"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR28348">
        <property type="entry name" value="UPF0193 PROTEIN EVG1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR28348:SF1">
        <property type="entry name" value="UPF0193 PROTEIN EVG1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05250">
        <property type="entry name" value="UPF0193"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1"/>
            <end position="27"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="65"/>
            <end position="86"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="65"/>
            <end position="74"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030155.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="233" mass="26608" checksum="AD68015BF46AC9CE" modified="2019-05-08" version="1">MSSQDGRRAAPAVPKGTGFWHSPGTTHYGQETKELLKVMMEESKLTNFQQRHLMDCMKRGVSLPTHCNPTSSKEPQLAQPAPSPPKPCLSVTLLARPHLRPAEICRAADAYTREKFKPRATRDLEREKQRLQNILATGKDMVEQKPQQKPVQTEEEIAEPDRFEELVNEIQERKEFLAEMEALGQSKQYRGTVLTEISQKLREMEIIDKRRSKDMRKAMAKIFPVGNKSDPNS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQN1</accession>
    <name>A0A452IQN1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Peptidase S1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the peptidase S1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQN1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034293.1">
        <property type="protein sequence ID" value="ENSGAGP00000030212.1"/>
        <property type="gene ID" value="ENSGAGG00000021723.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030141">
        <property type="term" value="C:secretory granule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00190">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.10:FF:000077">
        <property type="entry name" value="Predicted protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.10.10">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009003">
        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043504">
        <property type="entry name" value="Peptidase_S1_PA_chymotrypsin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001314">
        <property type="entry name" value="Peptidase_S1A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001254">
        <property type="entry name" value="Trypsin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018114">
        <property type="entry name" value="TRYPSIN_HIS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033116">
        <property type="entry name" value="TRYPSIN_SER"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271:SF48">
        <property type="entry name" value="KALLIKREIN-14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271">
        <property type="entry name" value="KALLIKREIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00089">
        <property type="entry name" value="Trypsin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00722">
        <property type="entry name" value="CHYMOTRYPSIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00020">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50494">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50240">
        <property type="entry name" value="TRYPSIN_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00134">
        <property type="entry name" value="TRYPSIN_HIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00135">
        <property type="entry name" value="TRYPSIN_SER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3 6" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="2 6" id="KW-0645">Protease</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 6" id="KW-0720">Serine protease</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="7">
        <location>
            <begin position="1"/>
            <end position="21"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019367677" description="Peptidase S1 domain-containing protein" evidence="7">
        <location>
            <begin position="22"/>
            <end position="251"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase S1" evidence="8">
        <location>
            <begin position="26"/>
            <end position="249"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007664"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="RuleBase" id="RU363034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030212.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="251" mass="27454" checksum="0243FA12811AD76A" modified="2019-05-08" version="1">ISPILFVGFIVNLSACLPVSSQNNRIVGGAPCIPHSMPWQAALFMGSQLNCGGTLIDRNWVVTAAHCHVQCSISVRLGEHNIRHLDWTEQLRLSEKTILHPGYNSASKDNDIMLIKLLTPANLNKNVQPLELPTSCPSPKEECVVSGWGTTKSPQLYFPPVLHCGKVSIFSQEQCRSIYPRYYTENMLCAGVLEGGIDSCQGDSGGPLLCNGKLQGIVSWGTQTCALPNKPGVYVAVCKYVDWIRETINNN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQN7</accession>
    <name>A0A452IQN7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Glycine amidinotransferase</fullName>
            <ecNumber evidence="10">2.1.4.1</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="10">L-arginine:glycine amidinotransferase</fullName>
        </alternativeName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">Catalyzes the biosynthesis of guanidinoacetate, the immediate precursor of creatine. Creatine plays a vital role in energy metabolism in muscle tissues. May play a role in embryonic and central nervous system development.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="10">
            <text>L-arginine + glycine = guanidinoacetate + L-ornithine</text>
            <dbReference type="Rhea" id="RHEA:13201"/>
            <dbReference type="ChEBI" id="CHEBI:32682"/>
            <dbReference type="ChEBI" id="CHEBI:46911"/>
            <dbReference type="ChEBI" id="CHEBI:57305"/>
            <dbReference type="ChEBI" id="CHEBI:57742"/>
            <dbReference type="EC" id="2.1.4.1"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="2 10">Amine and polyamine biosynthesis; creatine biosynthesis; creatine from L-arginine and glycine: step 1/2.</text>
    </comment>
    <comment type="subunit">
        <text evidence="10">Homodimer.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 10">Mitochondrion inner membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 10">Belongs to the amidinotransferase family.</text>
    </comment>
    <dbReference type="EC" id="2.1.4.1" evidence="10"/>
    <dbReference type="AlphaFoldDB" id="A0A452IQN7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034519.1">
        <property type="protein sequence ID" value="ENSGAGP00000030420.1"/>
        <property type="gene ID" value="ENSGAGG00000021905.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00104">
        <property type="reaction ID" value="UER00579"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005743">
        <property type="term" value="C:mitochondrial inner membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005758">
        <property type="term" value="C:mitochondrial intermembrane space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015068">
        <property type="term" value="F:glycine amidinotransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006601">
        <property type="term" value="P:creatine biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="CDD" id="cd21136">
        <property type="entry name" value="amidinotransferase_AGAT-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.75.10.10:FF:000005">
        <property type="entry name" value="Glycine amidinotransferase, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.75.10.10">
        <property type="entry name" value="L-arginine/glycine Amidinotransferase, Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033195">
        <property type="entry name" value="AmidinoTrfase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10488">
        <property type="entry name" value="GLYCINE AMIDINOTRANSFERASE, MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10488:SF1">
        <property type="entry name" value="GLYCINE AMIDINOTRANSFERASE, MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55909">
        <property type="entry name" value="Pentein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 10" id="KW-0472">Membrane</keyword>
    <keyword evidence="7 10" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="5 10" id="KW-0999">Mitochondrion inner membrane</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 10" id="KW-0808">Transferase</keyword>
    <keyword evidence="6" id="KW-0809">Transit peptide</keyword>
    <feature type="active site" evidence="9">
        <location>
            <position position="254"/>
        </location>
    </feature>
    <feature type="active site" evidence="9">
        <location>
            <position position="303"/>
        </location>
    </feature>
    <feature type="active site" description="Amidino-cysteine intermediate" evidence="9">
        <location>
            <position position="407"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004858"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00006943"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022792"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PIRSR" id="PIRSR633195-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="RuleBase" id="RU367092"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030420.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="418" mass="47732" checksum="8CDA79C4525FF17A" modified="2019-05-08" version="1">MLRVRCLRGGSRGAEAAHYIGSQLGRSFTGWVQRTFQSTQAAAASQNSCAVDDNKVTAPVPEDCPVCSYNEWDPLEEVIVGRAENACVPPFSVEVKANTYEKNWEFYQKFGGQSFPRDHLKKAIAEIEEMCNILKMEGVIVKRPEPIDWSVKYKTPDFESTGMYAAMPRDILLVVGNEIIEAPMAWRARFFEYRAYRPVIKDYFHRGAKWTTAPKPTMADELYYQDYPIHSVQDRHKLAAQGKFVTTEFEPCFDAADFIRAGRDIFVQRSQVTNNSGIEWMRRHLAPEYRVHTLSFKDPNPMHIDATLNIIGPGLVLSNPDRPCHQIEFFKKAGWTVVSPPLPLIPDDHPLWMSSKWLSMNVLMLDEKRVMVDANEIPIQKMFENLGISTIKVNIRHANSLGGGFHCWTCDIRRRGIL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IR00</accession>
    <name>A0A452IR00_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">Hepatocyte nuclear factor 4 alpha</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 11">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the nuclear hormone receptor family. NR2 subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IR00"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034370.1">
        <property type="protein sequence ID" value="ENSGAGP00000030283.1"/>
        <property type="gene ID" value="ENSGAGG00000021816.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003700">
        <property type="term" value="F:DNA-binding transcription factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045893">
        <property type="term" value="P:positive regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd06960">
        <property type="entry name" value="NR_DBD_HNF4A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd06931">
        <property type="entry name" value="NR_LBD_HNF4_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.565.10:FF:000007">
        <property type="entry name" value="Hepatocyte nuclear factor 4 alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.50.10:FF:000012">
        <property type="entry name" value="Hepatocyte nuclear factor 4, alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.50.10">
        <property type="entry name" value="Erythroid Transcription Factor GATA-1, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.565.10">
        <property type="entry name" value="Retinoid X Receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049636">
        <property type="entry name" value="HNF4-like_DBD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049635">
        <property type="entry name" value="HNF4_LBD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035500">
        <property type="entry name" value="NHR-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000536">
        <property type="entry name" value="Nucl_hrmn_rcpt_lig-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050274">
        <property type="entry name" value="Nuclear_hormone_rcpt_NR2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001723">
        <property type="entry name" value="Nuclear_hrmn_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001628">
        <property type="entry name" value="Znf_hrmn_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013088">
        <property type="entry name" value="Znf_NHR/GATA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24083">
        <property type="entry name" value="NUCLEAR HORMONE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00104">
        <property type="entry name" value="Hormone_recep"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00105">
        <property type="entry name" value="zf-C4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01282">
        <property type="entry name" value="COUPTNFACTOR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00398">
        <property type="entry name" value="STRDHORMONER"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00047">
        <property type="entry name" value="STROIDFINGER"/>
    </dbReference>
    <dbReference type="SMART" id="SM00430">
        <property type="entry name" value="HOLI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00399">
        <property type="entry name" value="ZnF_C4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57716">
        <property type="entry name" value="Glucocorticoid receptor-like (DNA-binding domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48508">
        <property type="entry name" value="Nuclear receptor ligand-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51843">
        <property type="entry name" value="NR_LBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00031">
        <property type="entry name" value="NUCLEAR_REC_DBD_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51030">
        <property type="entry name" value="NUCLEAR_REC_DBD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7 11" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3 11" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10 11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="9 11" id="KW-0675">Receptor</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8 11" id="KW-0804">Transcription</keyword>
    <keyword evidence="6 11" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="5 11" id="KW-0862">Zinc</keyword>
    <keyword evidence="4 11" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="Nuclear receptor" evidence="13">
        <location>
            <begin position="30"/>
            <end position="105"/>
        </location>
    </feature>
    <feature type="domain" description="NR LBD" evidence="14">
        <location>
            <begin position="120"/>
            <end position="350"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1"/>
            <end position="21"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006421"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="RuleBase" id="RU004334"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS51030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS51843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030283.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="17">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="442" mass="49114" checksum="3DBD46C42C755846" modified="2019-05-08" version="1">TPDPLNPLGGPDTSPSDATNLSTSNSIGVSALCAICGDRATGKHYGASSCDGCKGFFRRSVRKNHMYSCRFNRQCVVDKDKRNQCRYCRLKKCFRAGMKKEAVQNERDRISTRRSSYEDSSLPSINILLQAEVLAQQISSPISVINGDIRGKKIANITDVCESMKQQLLVLVEWAKYIPAFCELPLDDQVALLRAHAGEHLLLGAAKRSMVFKDILLLGNDHIIPRNCPELVEVIRVAIRILDELVLPFQELQIDDNEYACLKAIIFFDPDAKGLSDPTKIKRMRYQVQVSLEDYINDRQYDSRGRFGELLLLLPTLQSITWQMIEQIQFVKLFGMAKIDNLLQEMLLGGSSSETPHAHHPLHPHLIQENLGTNVIVANTMPAQMHNGQMLFLSAATPETPQPSPPAGSGSEQYKLLPGAIATVAKQPASIPQPTITKQEVI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IR01</accession>
    <name>A0A452IR01_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="5">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the RPAP1 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000034501.1">
        <property type="protein sequence ID" value="ENSGAGP00000030403.1"/>
        <property type="gene ID" value="ENSGAGG00000021786.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006366">
        <property type="term" value="P:transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039913">
        <property type="entry name" value="RPAP1/Rba50"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013929">
        <property type="entry name" value="RPAP1_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013930">
        <property type="entry name" value="RPAP1_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21483">
        <property type="entry name" value="RNA POLYMERASE II-ASSOCIATED PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21483:SF18">
        <property type="entry name" value="RNA POLYMERASE II-ASSOCIATED PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08620">
        <property type="entry name" value="RPAP1_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08621">
        <property type="entry name" value="RPAP1_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="RPAP1 N-terminal" evidence="4">
        <location>
            <begin position="238"/>
            <end position="280"/>
        </location>
    </feature>
    <feature type="domain" description="RPAP1 C-terminal" evidence="3">
        <location>
            <begin position="373"/>
            <end position="438"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="31"/>
            <end position="56"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="190"/>
            <end position="213"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="275"/>
            <end position="303"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="513"/>
            <end position="541"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="47"/>
            <end position="56"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="199"/>
            <end position="211"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="277"/>
            <end position="291"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="2">
        <location>
            <begin position="515"/>
            <end position="528"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00009953"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="Pfam" id="PF08620"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="Pfam" id="PF08621"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030403.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="944" mass="106164" checksum="A62C7D46E8F505F6" modified="2019-05-08" version="1">MLSRPKPGESEADLLRFQNQFLAARASPAVKVVKKADKRRGAGGSGDEERPPLQDSRDVVLLDDFPDVLPALIPAPPKKSKFKNACVRFDDEDPEERLECHDRHITAVFSKIIERDTSMAAVTMPVPTGDPFPRAFHRSEVKNEMKVESERKSIFAQKIAAKRAAEEAVTAPPAVETRYTHRRLAVPGAICSPGSAEEEPLHTTKDSKPDDDVSFLRPHIITGEGLGNLESEREALAIHEENLQKLHSMTQEEILQEQRRLLVQLDPSLVAFLKSRRSVDEGQERQEKPGHPEPGTESQHDGEETVKDFAVGGSGMEESVRPERDTRMEITEVDLPIKPQKEWIHMNNVEFEKLEWIKDLPRPRQRRTKKGMQARFSLKGELIPPDADLPTHLGLHHHGEEAERAGYSLQELFHLCRSQIIQQRTLALQVLGHVVQKAKAGEFASSLKGSVLRLLLDAGFLFLLRFSLDDTVDNVIAASLRALHALLVSPEDEEYLDQTFSWYQGMAVFPFTPNSEEEEEEEEEEELTLMEKTQGKKSKVENKSDPDVARYDVIKGLLKTQFLQRLRYILEVVRPVPAIVLDILDILIRVARHSSEACSQLLACPRLIETIVREFLPVQWDPRVAEPGHLLTSLHGVACATAMKLIRVLASGGRNATARLLNKFELKSRLSRFIAEDPQDLPLRREEAIRLSTEAFRLWAVAASCGQSCDLYSSGPGDGEQIPPPPVTWNQVSGLRPFLETNLRRSVQEMSRLETWQALQPLTTTYIIYLGVYYSVCSQQPSLNPIDCLEEVERLTYEVLLPLFSQPAMHRMWDLLRPCSALCNPLSCSPAPESVPSIVSLSCTGGKPQMSLVGSKSPFPFLTALLFLINNITHIHKGLVSKYTSVLDLKGLKDYLLQSWPTGPLSITPSSAWILRHEYHLQYFVLALAQXPPPQHFNHSYPDN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IR02</accession>
    <name>A0A452IR02_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="5">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the RPAP1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IR02"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034518.1">
        <property type="protein sequence ID" value="ENSGAGP00000030419.1"/>
        <property type="gene ID" value="ENSGAGG00000021786.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006366">
        <property type="term" value="P:transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039913">
        <property type="entry name" value="RPAP1/Rba50"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013929">
        <property type="entry name" value="RPAP1_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013930">
        <property type="entry name" value="RPAP1_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21483">
        <property type="entry name" value="RNA POLYMERASE II-ASSOCIATED PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21483:SF18">
        <property type="entry name" value="RNA POLYMERASE II-ASSOCIATED PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08620">
        <property type="entry name" value="RPAP1_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08621">
        <property type="entry name" value="RPAP1_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="RPAP1 N-terminal" evidence="4">
        <location>
            <begin position="191"/>
            <end position="233"/>
        </location>
    </feature>
    <feature type="domain" description="RPAP1 C-terminal" evidence="3">
        <location>
            <begin position="334"/>
            <end position="399"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="32"/>
            <end position="57"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="229"/>
            <end position="256"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="48"/>
            <end position="57"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="230"/>
            <end position="244"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00009953"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="Pfam" id="PF08620"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="Pfam" id="PF08621"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030419.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="831" mass="94107" checksum="D2F27379327D5BDE" modified="2019-05-08" version="1">IMLSRPKPGESEADLLRFQNQFLAARASPAVKVVKKADKRRGAGGSGDEERPPLQDSRDVVLLDDFPDVLPALIPAPPKKSKFKNACVRFDDEDPEERLECHDRHITAVFSKIIERDTSMAAVTMPVPTGDPFPRAFHRSEVKNEVRNAASPESSLTFFFNYQNFPLFLRPHIITGEGLGNLESEREALAIHEENLQKLHSMTQEEILQEQRRLLVQLDPSLVAFLKSRRSVDEGQERQEKPGHPEPGTESQHDGEETVKDFAVGGSGMEESVRPERDTRMEITGNKNLYLVAFDLPIKPQKEWIHMNNVEFEKLEWIKDLPRPRQRRTKKGMQARFSLKGELIPPDADLPTHLGLHHHGEEAERAGYSLQELFHLCRSQIIQQRTLALQVLGHVVQKAKAGEFASSLKGSVLRLLLDAGFLFLLRFSLDDTVDNVIAASLRALHALLVSPEDEEYLDQTFSWYQGMAVFPFTPNSEEEEEEEEEEELTLMEKTQGKKSKVENKSDPDVARYDVIKGLLKTQFLQRLRYILEVVRPVPAIVLDILDILIRVARHSSEACSQLLACPRLIETIVREFLPVQWDPRVAEPGHLLTSLHGVACATAMKLIRVLASGGRNATARLLNKFELKSRLSRFIAEDPQDLPLRREEAIRLSTEAFRLWAVAASCGQSCDLYRDLYPVLVKILQSLPHLLSDCQENPTFPLSIQRATSVITLLTHVTQTAGCTAELSGPGDGEQIPPPPVTWNQVSGLRPFLETNLRRSVQEMSRLETWQALQPLTTTYIIYLGVYYSVCSQQPSLNPIDCLEEVERLTYEVLLPLFSQPAMHRMWDLLR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IR03</accession>
    <name>A0A452IR03_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="13">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">Component of the transition zone in primary cilia. Required for ciliogenesis.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell projection</location>
            <location evidence="1">Cilium</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Membrane</location>
            <topology evidence="2">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the TMEM237 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IR03"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034434.1">
        <property type="protein sequence ID" value="ENSGAGP00000030340.1"/>
        <property type="gene ID" value="ENSGAGG00000021865.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000034435.1">
        <property type="protein sequence ID" value="ENSGAGP00000030341.1"/>
        <property type="gene ID" value="ENSGAGG00000021865.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035869">
        <property type="term" value="C:ciliary transition zone"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060271">
        <property type="term" value="P:cilium assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029409">
        <property type="entry name" value="TMEM237"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR28388">
        <property type="entry name" value="TRANSMEMBRANE PROTEIN 237"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR28388:SF1">
        <property type="entry name" value="TRANSMEMBRANE PROTEIN 237"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15383">
        <property type="entry name" value="TMEM237"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0966">Cell projection</keyword>
    <keyword evidence="7" id="KW-0969">Cilium</keyword>
    <keyword evidence="5" id="KW-0970">Cilium biogenesis/degradation</keyword>
    <keyword evidence="8 12" id="KW-0472">Membrane</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 12" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 12" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="218"/>
            <end position="241"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="261"/>
            <end position="282"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="294"/>
            <end position="315"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="346"/>
            <end position="367"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1"/>
            <end position="129"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="11">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="83"/>
            <end position="102"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="11">
        <location>
            <begin position="115"/>
            <end position="124"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004138"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00008783"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022794"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023069"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00025631"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030340.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="399" mass="44735" checksum="5ED4826369FBA4F8" modified="2019-05-08" version="1">METPPQGPRQCPPRALPPVPSGSQDEIPLSRPKRKKPKGKNALEGIVHAAVRRPSESSEPLTPEVQDAPPQRKRKKKRMPPESETSFIQQNLHNTSFFQTENGMDPQPSEETVIRKHRKKTKKTRPAEMQCSNELGVEEEDIIEDEQTKSPDQQPVFAAPAGASQPVSKVFVEKNRRFQAADRTELVKTTENIDVFMDVKALWTTRDVALSVHRGFRVIGLFTHGFLAGYAVWNIIVIYILAGNQLSTLSNLLEQYKALAYPAQSFLYLLLAISTVSAFDRIDLAKASVAMRGFLTLDPAAIASFLYFVALILSLSQQMTSDRINLYTPPSENGSLWTAGAEEQILQPWIVMNLVVALLVGLSWLFLSYRPELDHSEELMFHAEVEEFPQLDKGLNVQS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IR05</accession>
    <name>A0A452IR05_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="9">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the MAGUK family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IR05"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034647.1">
        <property type="protein sequence ID" value="ENSGAGP00000030531.1"/>
        <property type="gene ID" value="ENSGAGG00000021869.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="CDD" id="cd00071">
        <property type="entry name" value="GMPK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd06799">
        <property type="entry name" value="PDZ_MPP3-MPP4-MPP7-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12034">
        <property type="entry name" value="SH3_MPP4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.63.10:FF:000002">
        <property type="entry name" value="Guanylate kinase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.42.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008145">
        <property type="entry name" value="GK/Ca_channel_bsu"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008144">
        <property type="entry name" value="Guanylate_kin-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020590">
        <property type="entry name" value="Guanylate_kinase_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014775">
        <property type="entry name" value="L27_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004172">
        <property type="entry name" value="L27_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050716">
        <property type="entry name" value="MAGUK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035600">
        <property type="entry name" value="MPP4_SH3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001478">
        <property type="entry name" value="PDZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036034">
        <property type="entry name" value="PDZ_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23122">
        <property type="entry name" value="MEMBRANE-ASSOCIATED GUANYLATE KINASE MAGUK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00625">
        <property type="entry name" value="Guanylate_kin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02828">
        <property type="entry name" value="L27"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00595">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00072">
        <property type="entry name" value="GuKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00569">
        <property type="entry name" value="L27"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00228">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00326">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50156">
        <property type="entry name" value="PDZ domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00856">
        <property type="entry name" value="GUANYLATE_KINASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50052">
        <property type="entry name" value="GUANYLATE_KINASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51022">
        <property type="entry name" value="L27"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50106">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51257">
        <property type="entry name" value="PROKAR_LIPOPROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 3" id="KW-0728">SH3 domain</keyword>
    <feature type="domain" description="L27" evidence="8">
        <location>
            <begin position="25"/>
            <end position="81"/>
        </location>
    </feature>
    <feature type="domain" description="PDZ" evidence="7">
        <location>
            <begin position="128"/>
            <end position="209"/>
        </location>
    </feature>
    <feature type="domain" description="SH3" evidence="5">
        <location>
            <begin position="216"/>
            <end position="286"/>
        </location>
    </feature>
    <feature type="domain" description="Guanylate kinase-like" evidence="6">
        <location>
            <begin position="366"/>
            <end position="555"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="4">
        <location>
            <begin position="502"/>
            <end position="560"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007014"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50052"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50106"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51022"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030531.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="575" mass="65247" checksum="65C9795C17778ED5" modified="2019-05-08" version="1">MIHSEKRGESPRPFVLKCTALSCFVGSCLSHVLTLVLQELSLFCKRDVNGVGVLYDLLRSPWLQALLKVYECLHQSIRKKPVPVTPQARALSREALLSAHDTVAQKDFEPVLPPLPDNIPENEEAMRIVCLVKNKQPLGATIKRHEITGDIVVARVIHGGLADRSGLLYAGDKLVEVNGVPVEGLEPEQVIHILSLSQGTIMFKLIPVSDRPVSNQTTLYVRAMADYWPLQDPAIPCADAGLPFKKGEILQIVDQNDVLWWQARQVSDLSACAGLIPSNHLLKRKQREFWWSQPFQPHVCLKSTICDGSGRRVFIAGFRRSMRLCRRKSQNSQQPCYARCPSSCYSTAAAPYEEVVRYQRHPTDRNRLIVLVGPSGVGVNELRRRLIGMNPHHFQSAVPHTTRAQKSYEENGREYHYVSKETFENMVYSHRMLEHGEYKGHLYGTSVDAVRTVLDEGKICIVDLEPQGIQMARTHELKPYIIFIKPPSMSCMKQSRKNARIITDYYVNMKFKEEDLQEMEDRAKKMEVQFGQFFDQVIVNDNLQEACVQLLSAVHQAQDEPQWVPATWICSATQP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IR09</accession>
    <name>A0A452IR09_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="16">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 12">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 12">Belongs to the nuclear hormone receptor family. NR3 subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IR09"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034653.1">
        <property type="protein sequence ID" value="ENSGAGP00000030536.1"/>
        <property type="gene ID" value="ENSGAGG00000021989.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003707">
        <property type="term" value="F:nuclear steroid receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043565">
        <property type="term" value="F:sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005496">
        <property type="term" value="F:steroid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd06946">
        <property type="entry name" value="NR_LBD_ERR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.565.10:FF:000009">
        <property type="entry name" value="estrogen-related receptor gamma isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.50.10:FF:000092">
        <property type="entry name" value="Steroid hormone receptor ERR1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.50.10">
        <property type="entry name" value="Erythroid Transcription Factor GATA-1, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.565.10">
        <property type="entry name" value="Retinoid X Receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024178">
        <property type="entry name" value="Est_rcpt/est-rel_rcp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035500">
        <property type="entry name" value="NHR-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000536">
        <property type="entry name" value="Nucl_hrmn_rcpt_lig-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050200">
        <property type="entry name" value="Nuclear_hormone_rcpt_NR3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001723">
        <property type="entry name" value="Nuclear_hrmn_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027289">
        <property type="entry name" value="Oest-rel_rcp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001628">
        <property type="entry name" value="Znf_hrmn_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013088">
        <property type="entry name" value="Znf_NHR/GATA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48092">
        <property type="entry name" value="KNIRPS-RELATED PROTEIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00104">
        <property type="entry name" value="Hormone_recep"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00105">
        <property type="entry name" value="zf-C4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF002527">
        <property type="entry name" value="ER-like_NR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF500939">
        <property type="entry name" value="ERR1-2-3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00398">
        <property type="entry name" value="STRDHORMONER"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00047">
        <property type="entry name" value="STROIDFINGER"/>
    </dbReference>
    <dbReference type="SMART" id="SM00430">
        <property type="entry name" value="HOLI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00399">
        <property type="entry name" value="ZnF_C4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57716">
        <property type="entry name" value="Glucocorticoid receptor-like (DNA-binding domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48508">
        <property type="entry name" value="Nuclear receptor ligand-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51843">
        <property type="entry name" value="NR_LBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00031">
        <property type="entry name" value="NUCLEAR_REC_DBD_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51030">
        <property type="entry name" value="NUCLEAR_REC_DBD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0007">Acetylation</keyword>
    <keyword evidence="8" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11 12" id="KW-0539">Nucleus</keyword>
    <keyword evidence="10 12" id="KW-0675">Receptor</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9 12" id="KW-0804">Transcription</keyword>
    <keyword evidence="7 12" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <keyword evidence="4" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="Nuclear receptor" evidence="14">
        <location>
            <begin position="125"/>
            <end position="156"/>
        </location>
    </feature>
    <feature type="domain" description="NR LBD" evidence="15">
        <location>
            <begin position="194"/>
            <end position="418"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="42"/>
            <end position="92"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="13">
        <location>
            <begin position="42"/>
            <end position="52"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="13">
        <location>
            <begin position="57"/>
            <end position="70"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005413"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PIRNR" id="PIRNR002527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS51030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS51843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030536.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="419" mass="46764" checksum="BCF162430D2933BC" modified="2019-05-08" version="1">MDSVELCLPESFSLHYEEELLCRMSSKDRHIDSSCSSYIKTEPSSPASLTDSINHHSPGGSSDASGSYSSTMNGHQNGLDSPPLYPSAAGLGGNGPVRKRYDDCSSTIAEDSQTKCEYMLNSMPKRLCLVCGDIASGYHYGVASCEACKAFFKRTIQGVRLDRVRGGRQKYKRRIDAENSPYLNPQLVQPAKKPYNKIVSHLLVAEPEKIYAMPDPTVPDSDIKALTTLCDLADRELVVIIGWAKHIPGFSTLSLADQMSLLQSAWMEILILGVVYRSLSFEDELVYAEDYIMDEDQSKLAGLLDLNNAILQLVKKYKSMKLEKEEFVTLKAIALANSDSMHIEDVEAVQKLQDVLHEALQDYEAGQHMEDPRRAGKMLMTLPLLRQTSTKAVQHFYNIKLEGKVPMHKLFLEMLEAKV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2H1</accession>
    <name>A0A452J2H1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Sodium/hydrogen exchanger</fullName>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Recycling endosome membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 13">Belongs to the monovalent cation:proton antiporter 1 (CPA1) transporter (TC 2.A.36) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2H1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039173.1">
        <property type="protein sequence ID" value="ENSGAGP00000034599.1"/>
        <property type="gene ID" value="ENSGAGG00000024543.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055038">
        <property type="term" value="C:recycling endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015386">
        <property type="term" value="F:potassium:proton antiporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015385">
        <property type="term" value="F:sodium:proton antiporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051453">
        <property type="term" value="P:regulation of intracellular pH"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098719">
        <property type="term" value="P:sodium ion import across plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.1330">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018422">
        <property type="entry name" value="Cation/H_exchanger_CPA1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006153">
        <property type="entry name" value="Cation/H_exchanger_TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004709">
        <property type="entry name" value="NaH_exchanger"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002090">
        <property type="entry name" value="NHE-6/7/9"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00840">
        <property type="entry name" value="b_cpa1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10110">
        <property type="entry name" value="SODIUM/HYDROGEN EXCHANGER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10110:SF94">
        <property type="entry name" value="SODIUM_HYDROGEN EXCHANGER 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00999">
        <property type="entry name" value="Na_H_Exchanger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01084">
        <property type="entry name" value="NAHEXCHNGR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01088">
        <property type="entry name" value="NAHEXCHNGR6"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="13" id="KW-0050">Antiport</keyword>
    <keyword evidence="5" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="7" id="KW-0967">Endosome</keyword>
    <keyword evidence="10 13" id="KW-0406">Ion transport</keyword>
    <keyword evidence="11 15" id="KW-0472">Membrane</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0915">Sodium</keyword>
    <keyword evidence="12 13" id="KW-0739">Sodium transport</keyword>
    <keyword evidence="6 13 15" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 15" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="4 13" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="70"/>
            <end position="87"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="99"/>
            <end position="116"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="178"/>
            <end position="195"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="207"/>
            <end position="233"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="322"/>
            <end position="349"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="361"/>
            <end position="390"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="411"/>
            <end position="432"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="444"/>
            <end position="466"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="478"/>
            <end position="497"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="503"/>
            <end position="524"/>
        </location>
    </feature>
    <feature type="domain" description="Cation/H+ exchanger transmembrane" evidence="16">
        <location>
            <begin position="80"/>
            <end position="532"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="680"/>
            <end position="700"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004195"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007367"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023053"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023201"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="RuleBase" id="RU003722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF00999"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034599.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="700" mass="77204" checksum="64B4F7745EA683FB" modified="2019-05-08" version="1">GAPRCTRLAPLRTRVLRCPGSLWPPSAMCIRDRAGAAARGEAAARPVALEEEIVSEKAAEESHRQDSANLLIFILLLTLTILTIWLFKHRRARFLHETGLAMIYGVLMGVILRYGIHVPSDVNNVTLSCQVQTSPATLLVNVSGKYYEYTLKGEISPHELNNVQDNEMLRKVTFDPEVFFNILLPPIIFYAGYSLKRRHFFRNLGSILAYAFLGTAISCFVIGSIMYGCVALMKVTGQLGGDFYFTDCLLFGAIVSATDPVTVLAIFHELQVDVELYALLFGESVLNDAVAIVLSSSIVAYQPAGDNSHTFDVTAMFKSIGIFLGIFSGSFAMGAATGVVTALISFSLFTKLREFPLLETGLFFLMSWSTFLLAEACGFTGVVAVLFCGITQAHYTYNNLSTESQHRTKQLFELLNFLAENFIFSYMGLTLFTFQNHDFNPTFIVGAFFAIFLGRAANIYPLSFLLNLGRRNKIGTNLQHMMMFAGLRGAMAFALAIRDTATYARQMMFSTTLLIVFFTVWVFGGGTTAMLSCLHIRVGVDSDQEHLGVPESERRSTKAESAWLFRLWYNFDHNYLKPLLTHSGPPLTTTLPSCCGPVARCLTSPQAYENQEQLKDDDSDLILNDGDISLTYGDSTVNADSITSSGTSRRIVGNSSEDALDRELAFGDHELVIRGTRLVLPMDDSEPPSSLVDSARHGPA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2H4</accession>
    <name>A0A452J2H4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Nuclear protein MDM1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Microtubule-binding protein that negatively regulates centriole duplication. Binds to and stabilizes microtubules.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Microtubule organizing center</location>
            <location evidence="1">Centrosome</location>
            <location evidence="1">Centriole</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the MDM1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2H4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039200.1">
        <property type="protein sequence ID" value="ENSGAGP00000034625.1"/>
        <property type="gene ID" value="ENSGAGG00000024610.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005814">
        <property type="term" value="C:centriole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005874">
        <property type="term" value="C:microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008017">
        <property type="term" value="F:microtubule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046600">
        <property type="term" value="P:negative regulation of centriole replication"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060041">
        <property type="term" value="P:retina development in camera-type eye"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029136">
        <property type="entry name" value="MDM1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR32078">
        <property type="entry name" value="NUCLEAR PROTEIN MDM1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR32078:SF1">
        <property type="entry name" value="NUCLEAR PROTEIN MDM1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15501">
        <property type="entry name" value="MDM1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="6" id="KW-0493">Microtubule</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="80"/>
            <end position="127"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="278"/>
            <end position="299"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="434"/>
            <end position="477"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="552"/>
            <end position="581"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="10">
        <location>
            <begin position="290"/>
            <end position="299"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="457"/>
            <end position="477"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="555"/>
            <end position="581"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004114"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00010494"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00013508"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00045771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034625.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="622" mass="70603" checksum="9FD2FA50839644CF" modified="2019-05-08" version="1">MPVRFKGLSEYKRNFKWKKSHLSEFCNPSQEQKSSRAGLRSDQLGITREPNFISKRRVPYFNPQISESFEWRGDSDLEGYLDDAPEIEGAGPAESHTNNHSNDRNLENIETPEAPRLPKKIRSHSADSRIESAITLAENSRKKLSPPTPVNQKAVFVPPKKHLEKMDNGFQRVLQKKSGMNISPSDSFPRNSEYQRQFLWKSPPKNSPILAADQVIHNTSKSIPPFKSSSIIPETEYERSFKVSPSAVGPKLRCDLEEREFPECAPVNISPERKVYTSLNGKSETEQKPPKQKNKQHITQKKVNTEYRSKFLSPAQYLYKDGAWSRIRRNIPDQVSTNKCTLQRVQGTHFSRYHLNQILSDNNRLWDVSSNSSSEETISNTIRALDLAGAPEKQMSASQNVLLQPDSIEQSQQNNTEKLGMSDASTVPVRRRLVWGEQDSTEQVENQPPGLEEDEEKENKQESAKAQKLEENDKNPTVDNKIKGYILKFHCLICFLRQFPSEAKGKMEEEIDPELLQGFGSANDFRETEAVSLLASPAAGLKTLDPLPLRKDSWPTVNASDERASPTSAHPTTAPVQKSVKSTSPSYWSPCCRIQGTLRDPEFQHNGCFFLNRNFPSTEFVW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2H7</accession>
    <name>A0A452J2H7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">VWFA domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J2H7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039180.1">
        <property type="protein sequence ID" value="ENSGAGP00000034606.1"/>
        <property type="gene ID" value="ENSGAGG00000024596.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032039">
        <property type="term" value="C:integrator complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034472">
        <property type="term" value="P:snRNA 3'-end processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00198">
        <property type="entry name" value="vWFA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.410:FF:000010">
        <property type="entry name" value="Integrator complex subunit 6 like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.410">
        <property type="entry name" value="von Willebrand factor, type A domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057413">
        <property type="entry name" value="Beta-barrel_INTS6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029307">
        <property type="entry name" value="INT_SG_DDX_CT_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051113">
        <property type="entry name" value="Integrator_subunit6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002035">
        <property type="entry name" value="VWF_A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036465">
        <property type="entry name" value="vWFA_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12957">
        <property type="entry name" value="DEAD/H BOX POLYPEPTIDE 26/DICE1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12957:SF22">
        <property type="entry name" value="INTEGRATOR COMPLEX SUBUNIT 6-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25462">
        <property type="entry name" value="Beta-barrel_INTS6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15300">
        <property type="entry name" value="INT_SG_DDX_CT_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13519">
        <property type="entry name" value="VWA_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53300">
        <property type="entry name" value="vWA-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50234">
        <property type="entry name" value="VWFA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="VWFA" evidence="2">
        <location>
            <begin position="3"/>
            <end position="227"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="601"/>
            <end position="683"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="1">
        <location>
            <begin position="648"/>
            <end position="658"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="PROSITE" id="PS50234"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034606.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="862" mass="96446" checksum="FC3FDEBCBDF8D630" modified="2019-05-08" version="1">MPILLFLIDTSASMNQRAHLGTSYLDIAKGAVELFMKLRARDPASRGDRYMLVTFDEPPYCIKAGWKENHATFMNELKNLQASGLTTLGQALRSSFDLLNLNRLVSGIDNYGQGRNPFFLEPSILITITDGNKLTHTAGVQEELHLPLNSPLPGSELTKEPFRWDQRLFALVLRLPGASSVEPEQLGSVPTDESAITQMCEVTGGRSYCVRTQRMLNQCLESLVQKVQSGVVINFEKSGPDPAPIGEDGLVDSSRPINSFASQPWHSCHKLIYVRPNPKTGVPVGHWPIPESFWPDQNSPTLPPRTAHPVVRFSCVDCEPMVIDKLPFDKYELEPSPLTQYILERKSPHACWQVFVSNSGKYSELGHPFGYLKASTTLTCVNLFVMPYNYPALLPLLDDLFKVHKLKPNLKWRQAFDSYLKTMPPYYVLTKIESERIIGSVGKKVPQETGIKVKNHSSGLSLVHSRDFKQLLQGITGDSPLRLAEMNVKEFAGFHIGLLNKDLKPQAFRNAYDIPRRSLLDQLTRMRTNLLKTHKLILGQDEDCLHSVPVAQMGNYQEYLKMMPSPLREIDPDQPKRLHTFGNPFKQDKKGMMIDEADEFVAGPQNKIKRPGDPNTPASLKRRRSMSPLLRRPQSPPVITNHVGGKGPPSASGSPSYPNLKAIPGHKDANSSVVQDANGGKKAENCQSLDKQIDGLSIQMVPSVPAAMVDDDLLPNDLDSLPDEFNCLSEDGLDHKSATNALVRGPLNYSMPVDDQKMVLESTSGTVPDSLQITPTMLEGNNADIKIKVMKEVRKPGRNYEKIFALLEEVQGPMEIQKYFIEFAIKEAARFKKRVLIQHLERILEEIEFSSLPNRVNHVNSR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2I0</accession>
    <name>A0A452J2I0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">Glutamate receptor</fullName>
        </recommendedName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="21">Receptor for glutamate that functions as a ligand-gated ion channel in the central nervous system and plays an important role in excitatory synaptic transmission. L-glutamate acts as an excitatory neurotransmitter at many synapses in the central nervous system.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>Ca(2+)(in) = Ca(2+)(out)</text>
            <dbReference type="Rhea" id="RHEA:29671"/>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="16 21">Postsynaptic cell membrane</location>
            <topology evidence="16 21">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="21">Belongs to the glutamate-gated ion channel (TC 1.A.10.1) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2I0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039205.1">
        <property type="protein sequence ID" value="ENSGAGP00000034630.1"/>
        <property type="gene ID" value="ENSGAGG00000024584.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015276">
        <property type="term" value="F:ligand-gated monoatomic ion channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0038023">
        <property type="term" value="F:signaling receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd06382">
        <property type="entry name" value="PBP1_iGluR_Kainate"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.2300:FF:000010">
        <property type="entry name" value="Glutamate ionotropic receptor kainate type subunit 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.190.10:FF:000210">
        <property type="entry name" value="Glutamate receptor ionotropic, kainate 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.190.10:FF:000240">
        <property type="entry name" value="Glutamate receptor ionotropic, kainate 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.70:FF:000010">
        <property type="entry name" value="Putative glutamate receptor ionotropic kainate 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.70">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.2300">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.190.10">
        <property type="entry name" value="Periplasmic binding protein-like II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001828">
        <property type="entry name" value="ANF_lig-bd_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019594">
        <property type="entry name" value="Glu/Gly-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001508">
        <property type="entry name" value="Iono_Glu_rcpt_met"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015683">
        <property type="entry name" value="Ionotropic_Glu_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001320">
        <property type="entry name" value="Iontro_rcpt_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028082">
        <property type="entry name" value="Peripla_BP_I"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18966">
        <property type="entry name" value="IONOTROPIC GLUTAMATE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01094">
        <property type="entry name" value="ANF_receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00060">
        <property type="entry name" value="Lig_chan"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10613">
        <property type="entry name" value="Lig_chan-Glu_bd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00177">
        <property type="entry name" value="NMDARECEPTOR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00918">
        <property type="entry name" value="Lig_chan-Glu_bd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00079">
        <property type="entry name" value="PBPe"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53822">
        <property type="entry name" value="Periplasmic binding protein-like I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53850">
        <property type="entry name" value="Periplasmic binding protein-like II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 21" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="10 20" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="15 21" id="KW-0407">Ion channel</keyword>
    <keyword evidence="8 21" id="KW-0406">Ion transport</keyword>
    <keyword evidence="14 21" id="KW-1071">Ligand-gated ion channel</keyword>
    <keyword evidence="9 21" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="13 21" id="KW-0628">Postsynaptic cell membrane</keyword>
    <keyword evidence="11 21" id="KW-0675">Receptor</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <keyword evidence="7 21" id="KW-0770">Synapse</keyword>
    <keyword evidence="4 21" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 21" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="1 21" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="540"/>
            <end position="559"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="616"/>
            <end position="638"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="797"/>
            <end position="821"/>
        </location>
    </feature>
    <feature type="domain" description="Ionotropic glutamate receptor C-terminal" evidence="22">
        <location>
            <begin position="409"/>
            <end position="778"/>
        </location>
    </feature>
    <feature type="domain" description="Ionotropic glutamate receptor L-glutamate and glycine-binding" evidence="23">
        <location>
            <begin position="419"/>
            <end position="484"/>
        </location>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="493"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="500"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="667"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="715"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="site" description="Crucial to convey clamshell closure to channel opening" evidence="19">
        <location>
            <position position="645"/>
        </location>
    </feature>
    <feature type="site" description="Interaction with the cone snail toxin Con-ikot-ikot" evidence="19">
        <location>
            <position position="672"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="73"/>
            <end position="324"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="727"/>
            <end position="781"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023257"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00034104"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00036634"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="RuleBase" id="RU367118"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="SMART" id="SM00079"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="SMART" id="SM00918"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034630.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="885" mass="99760" checksum="117B9D6D5A98681C" modified="2019-05-08" version="1">QTNKKPVMQNVPDLKGGIFECVESGPMGAEELAFRFAVNTINRNRTLLPNTTLTYDTQKINLYDSFEASKKACDQLSLGVAAIFGPSHSSSANAVQSICNALGVPHIQTRWKHQVSDNKDSFYVSLYPDFSSLSRAILDLVQFFKWKTVTVVYDDSTGLIRLQELIKAPSRYNLRLKIRQLPADTKDAKPLLKEMKRGKEFHVIFDCSHEMAAGILKQALAMGMMTEYYHYIFTTLDLFALDVEPYRYSGVNMTGFRILNTENSQVASIIEKWSMERLQAPPKPDSGLLDGFMTTDAALMYDAVHVVSVAVQQFPQMTVSSLQCNRHKPWRFGTRFMSLIKEAHWEGLTGRITFNKTNGLRTDFDLDVISLKEEGLEKIGTWDPASGLNMTENQKGKPANITDSLSNRSLIVTTILEEPYVMFKKSDKPLYGNDRFEGYCIDLLRELATILGFTYEIRLVEDGKYGAQDDASGQWNGMVRELIDHKADLAVAPLAITYVREKVIDFSKPFMTLGISILYRKPNGTNPGVFSFLNPLSPDIWMYILLAYLGVSCVLFVIARFSPYEWYNPHPCNPDSDVVENNFTLLNSFWFGVGALMQQGSELMPKALSTRIVGGIWWFFTLIIISSYTANLAAFLTVERMESPIDSADDLAKQTKIEYGAVEDGATMTFFKKSKISTYDKMWAFMSSRRQSVLVKSNEEGIQRVLTSDYAFLMESTTIEFVTQRNCNLTQIGGLIDSKGYGVGTPMGSPYRDKITIAILQLQEEGKLHMMKEKWWRGNGCPEEESKEASALGVQNIGGIFIVLAAGLVLSVFVAVGEFLYKSKKNAQLEKRSFCSAMVEELRMSLKCQRRLKHKPQAPVIVKTEEVINMHTFNDRRLPGKETMA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2I3</accession>
    <name>A0A452J2I3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Ig-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J2I3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039078.1">
        <property type="protein sequence ID" value="ENSGAGP00000034515.1"/>
        <property type="gene ID" value="ENSGAGG00000024508.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002764">
        <property type="term" value="P:immune response-regulating signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000049">
        <property type="entry name" value="Leukocyte immunoglobulin-like receptor subfamily B member 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050412">
        <property type="entry name" value="Ig-like_Receptors_ImmuneReg"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11738">
        <property type="entry name" value="MHC CLASS I NK CELL RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11738:SF186">
        <property type="entry name" value="OSTEOCLAST-ASSOCIATED IMMUNOGLOBULIN-LIKE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13895">
        <property type="entry name" value="Ig_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0732">Signal</keyword>
    <feature type="domain" description="Ig-like" evidence="4">
        <location>
            <begin position="28"/>
            <end position="96"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034515.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="197" mass="20855" checksum="96725BD7E9F8DCA7" modified="2019-05-08" version="1">SEPTEQVLLSLGLASIPSPRAGHEFLKPTIWVSPSRVVALGRNVTIRCEGQDLGMEFVLCKAGHPNQQVQSVVPEGKMAEFPIASVGQEDGGSYTCYYRSITDQSHWSFPSNPVEIIVGGEGGAWLSGPAPSPTPRQTHRGSLQGVRFVLNKEGRHFPPTSSHPSAQASPAPSHSLVLLWGTKVGVLGLLAACSPGV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2I6</accession>
    <name>A0A452J2I6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Rho-GAP domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J2I6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039197.1">
        <property type="protein sequence ID" value="ENSGAGP00000034622.1"/>
        <property type="gene ID" value="ENSGAGG00000024607.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:1902533">
        <property type="term" value="P:positive regulation of intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.555.10:FF:000017">
        <property type="entry name" value="Rho GTPase activating protein 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.555.10">
        <property type="entry name" value="Rho GTPase activation protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037863">
        <property type="entry name" value="RHOGAP6/36"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000198">
        <property type="entry name" value="RhoGAP_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12635:SF7">
        <property type="entry name" value="RHO GTPASE ACTIVATING PROTEIN 6-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12635">
        <property type="entry name" value="RHO-GTPASE-ACTIVATING PROTEIN 6 FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00620">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00324">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50238">
        <property type="entry name" value="RHOGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Rho-GAP" evidence="2">
        <location>
            <begin position="187"/>
            <end position="387"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="PROSITE" id="PS50238"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034622.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="395" mass="44623" checksum="CF9A775CF52A9C8E" modified="2019-05-08" version="1">MQPVPLHSLSELERACLQEIAFYQLQKKNIISGIGLSKAGPKRNKFIRRKLENSSKEKKDCCPQAFGIPLSQVIANDWTHKQMQEAVKGSRRDCLDVEETVAKFRAQIQKKSPFVGGCVLGLSGEFADEPLSPTFLHTVYRSHRRGGVSVDSITDLGDNTSRLLEALQLSHPNELGLRRTLERNKMLSLNPIYRQVPRVVDRCCQHIEAYGLQTVGIFRVGSSKKRVQQLREEFDQGFDVFLDDSYCVHDVAALLKEFFRDMPDSLIPKELYSAFVSTAFMERMAQLSMLQLLVYLLPPCNSDTLYQLLQFLDKVAQHAQDSTGPERQKIPGNKMTASNLATIFGPNLLQREKLLERDCSAHAVEIENSASVILVLQSLIENYEALFMEDVSITT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2I7</accession>
    <name>A0A452J2I7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">Glutamate receptor</fullName>
        </recommendedName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="21">Receptor for glutamate that functions as a ligand-gated ion channel in the central nervous system and plays an important role in excitatory synaptic transmission. L-glutamate acts as an excitatory neurotransmitter at many synapses in the central nervous system.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>Ca(2+)(in) = Ca(2+)(out)</text>
            <dbReference type="Rhea" id="RHEA:29671"/>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="16 21">Postsynaptic cell membrane</location>
            <topology evidence="16 21">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="21">Belongs to the glutamate-gated ion channel (TC 1.A.10.1) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2I7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039210.1">
        <property type="protein sequence ID" value="ENSGAGP00000034635.1"/>
        <property type="gene ID" value="ENSGAGG00000024584.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
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        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
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    <dbReference type="GO" id="GO:0015276">
        <property type="term" value="F:ligand-gated monoatomic ion channel activity"/>
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        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0038023">
        <property type="term" value="F:signaling receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd06382">
        <property type="entry name" value="PBP1_iGluR_Kainate"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.2300:FF:000010">
        <property type="entry name" value="Glutamate ionotropic receptor kainate type subunit 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.190.10:FF:000210">
        <property type="entry name" value="Glutamate receptor ionotropic, kainate 1"/>
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    </dbReference>
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        <property type="entry name" value="Glutamate receptor ionotropic, kainate 2"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Putative glutamate receptor ionotropic kainate 1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.2300">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.190.10">
        <property type="entry name" value="Periplasmic binding protein-like II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001828">
        <property type="entry name" value="ANF_lig-bd_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019594">
        <property type="entry name" value="Glu/Gly-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001508">
        <property type="entry name" value="Iono_Glu_rcpt_met"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015683">
        <property type="entry name" value="Ionotropic_Glu_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001320">
        <property type="entry name" value="Iontro_rcpt_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028082">
        <property type="entry name" value="Peripla_BP_I"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18966">
        <property type="entry name" value="IONOTROPIC GLUTAMATE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01094">
        <property type="entry name" value="ANF_receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00060">
        <property type="entry name" value="Lig_chan"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10613">
        <property type="entry name" value="Lig_chan-Glu_bd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00177">
        <property type="entry name" value="NMDARECEPTOR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00918">
        <property type="entry name" value="Lig_chan-Glu_bd"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="PBPe"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Periplasmic binding protein-like I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53850">
        <property type="entry name" value="Periplasmic binding protein-like II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 21" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="10 20" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="15 21" id="KW-0407">Ion channel</keyword>
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    <keyword evidence="14 21" id="KW-1071">Ligand-gated ion channel</keyword>
    <keyword evidence="9 21" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="13 21" id="KW-0628">Postsynaptic cell membrane</keyword>
    <keyword evidence="11 21" id="KW-0675">Receptor</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <keyword evidence="7 21" id="KW-0770">Synapse</keyword>
    <keyword evidence="4 21" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 21" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="1 21" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="541"/>
            <end position="560"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="617"/>
            <end position="639"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="798"/>
            <end position="822"/>
        </location>
    </feature>
    <feature type="domain" description="Ionotropic glutamate receptor C-terminal" evidence="22">
        <location>
            <begin position="410"/>
            <end position="779"/>
        </location>
    </feature>
    <feature type="domain" description="Ionotropic glutamate receptor L-glutamate and glycine-binding" evidence="23">
        <location>
            <begin position="420"/>
            <end position="485"/>
        </location>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="494"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="501"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="668"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="716"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="site" description="Crucial to convey clamshell closure to channel opening" evidence="19">
        <location>
            <position position="646"/>
        </location>
    </feature>
    <feature type="site" description="Interaction with the cone snail toxin Con-ikot-ikot" evidence="19">
        <location>
            <position position="673"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="74"/>
            <end position="325"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="728"/>
            <end position="782"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023257"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00034104"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00036634"/>
        </source>
    </evidence>
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    </evidence>
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        </source>
    </evidence>
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            <dbReference type="RuleBase" id="RU367118"/>
        </source>
    </evidence>
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        </source>
    </evidence>
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            <dbReference type="SMART" id="SM00918"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034635.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="845" mass="95121" checksum="628F0016B3A9DCEE" modified="2019-05-08" version="1">GMLLEKGRGWGRETVFRGIFECVESGPMGAEELAFRFAVNTINRNRTLLPNTTLTYDTQKINLYDSFEASKKACDQLSLGVAAIFGPSHSSSANAVQSICNALGVPHIQTRWKHQVSDNKDSFYVSLYPDFSSLSRAILDLVQFFKWKTVTVVYDDSTGLIRLQELIKAPSRYNLRLKIRQLPADTKDAKPLLKEMKRGKEFHVIFDCSHEMAAGILKQALAMGMMTEYYHYIFTTLDLFALDVEPYRYSGVNMTGFRILNTENSQVASIIEKWSMERLQAPPKPDSGLLDGFMTTDAALMYDAVHVVSVAVQQFPQMTVSSLQCNRHKPWRFGTRFMSLIKEAHWEGLTGRITFNKTNGLRTDFDLDVISLKEEGLEKIGTWDPASGLNMTENQKGKPANITDSLSNRSLIVTTILEEPYVMFKKSDKPLYGNDRFEGYCIDLLRELATILGFTYEIRLVEDGKYGAQDDASGQWNGMVRELIDHKADLAVAPLAITYVREKVIDFSKPFMTLGISILYRKPNGTNPGVFSFLNPLSPDIWMYILLAYLGVSCVLFVIARFSPYEWYNPHPCNPDSDVVENNFTLLNSFWFGVGALMQQGSELMPKALSTRIVGGIWWFFTLIIISSYTANLAAFLTVERMESPIDSADDLAKQTKIEYGAVEDGATMTFFKKSKISTYDKMWAFMSSRRQSVLVKSNEEGIQRVLTSDYAFLMESTTIEFVTQRNCNLTQIGGLIDSKGYGVGTPMGSPYRDKITIAILQLQEEGKLHMMKEKWWRGNGCPEEESKEASALGVQNIGGIFIVLAAGLVLSVFVAVGEFLYKSKKNAQCEANSMTYPWILHQEG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2K2</accession>
    <name>A0A452J2K2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Galactocerebrosidase</fullName>
            <ecNumber evidence="3">3.2.1.46</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="15">Galactosylceramidase</fullName>
        </alternativeName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="14">
            <text>a D-galactosylceramide + H2O = an N-acyl-sphingoid base + D-galactose</text>
            <dbReference type="Rhea" id="RHEA:43412"/>
            <dbReference type="ChEBI" id="CHEBI:4139"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:36498"/>
            <dbReference type="ChEBI" id="CHEBI:83273"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="14">
            <dbReference type="Rhea" id="RHEA:43413"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="16">
            <text>beta-D-galactosyl-(1&lt;-&gt;1)-sphing-4-enine + H2O = sphing-4-enine + D-galactose</text>
            <dbReference type="Rhea" id="RHEA:43908"/>
            <dbReference type="ChEBI" id="CHEBI:4139"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:57756"/>
            <dbReference type="ChEBI" id="CHEBI:57934"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="16">
            <dbReference type="Rhea" id="RHEA:43909"/>
        </physiologicalReaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Lysosome</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the glycosyl hydrolase 59 family.</text>
    </comment>
    <dbReference type="EC" id="3.2.1.46" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452J2K2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039212.1">
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        <property type="gene ID" value="ENSGAGG00000024615.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="term" value="C:lysosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="GOC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004336">
        <property type="term" value="F:galactosylceramidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006683">
        <property type="term" value="P:galactosylceramide catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
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        <property type="entry name" value="galactocerebrosidase precursor"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="galactocerebrosidase precursor"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="galactocerebrosidase precursor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.20.20.70">
        <property type="entry name" value="Aldolase class I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.560">
        <property type="entry name" value="Exo-inulinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.20.20.80">
        <property type="entry name" value="Glycosidases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013785">
        <property type="entry name" value="Aldolase_TIM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017853">
        <property type="entry name" value="GH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049162">
        <property type="entry name" value="GH59_C"/>
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        <property type="entry name" value="Glyco_hydro_59"/>
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        <property type="entry name" value="Glyco_hydro_59_dom"/>
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        <property type="entry name" value="GALACTOCEREBROSIDASE"/>
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        <property type="entry name" value="Glyco_hydro_59M"/>
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        <property type="entry name" value="GLHYDRLASE59"/>
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    <dbReference type="SUPFAM" id="SSF51445">
        <property type="entry name" value="(Trans)glycosidases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13" id="KW-0326">Glycosidase</keyword>
    <keyword evidence="6" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="8" id="KW-0442">Lipid degradation</keyword>
    <keyword evidence="9" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="12" id="KW-0458">Lysosome</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <keyword evidence="7" id="KW-0746">Sphingolipid metabolism</keyword>
    <feature type="domain" description="Glycosyl hydrolase family 59 catalytic" evidence="18">
        <location>
            <begin position="2"/>
            <end position="262"/>
        </location>
    </feature>
    <feature type="domain" description="Glycosyl hydrolase family 59 central" evidence="19">
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            <begin position="270"/>
            <end position="384"/>
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    <feature type="domain" description="Glycosyl hydrolase family 59 C-terminal lectin" evidence="20">
        <location>
            <begin position="421"/>
            <end position="595"/>
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    </feature>
    <feature type="active site" description="Proton donor/acceptor" evidence="17">
        <location>
            <position position="111"/>
        </location>
    </feature>
    <feature type="active site" description="Nucleophile" evidence="17">
        <location>
            <position position="187"/>
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    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004371"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00005637"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00012657"/>
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            <dbReference type="ARBA" id="ARBA00019657"/>
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            <dbReference type="ARBA" id="ARBA00022729"/>
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            <dbReference type="ARBA" id="ARBA00023295"/>
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        <source>
            <dbReference type="PIRSR" id="PIRSR601286-50"/>
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        <source>
            <dbReference type="Pfam" id="PF02057"/>
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        <source>
            <dbReference type="Pfam" id="PF17387"/>
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        <source>
            <dbReference type="Pfam" id="PF21708"/>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034637.1"/>
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    </evidence>
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        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
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    <sequence length="596" mass="67505" checksum="8AF5AF5925D279F8" modified="2019-05-08" version="1">MPNFGASLHILKVEIGGDGQSTDGTEPSHMHYENDENYFRGYEWWLMKEAKKRNPNIKLIGLPWAFPGWIGRGQNWPYNHPDVTAYYIVSWILGAKQYHNLDIDYIGIWNERSFSSKYIKLLRYTLDKQGLQRVRIIASDNMWEPISFFMMFDSELHQVVDIIGAHYPGTRTVPNALATQKKLWASEDYSTFNDEVGAGCWARILNQNYVNGNMTSTIAWNLVASYYEDLPFGRDGLMTAQEPWSGCYAVEGPIWITAHTTQFTHPGWHYLQVDGRLEHGGSYVALTDGLGNLTIIIETMTYRHSQCIRPPLLPFIVLPQKATFYLKGSFHKVTILKMWYTKLGFKSANSTLFHPLPPVKVHEGMFSLNLDVDEVYTLTTLTTGRKGTYPDPPQSKPFPSNYKDDFNIRNPPFSEAPHFADQTGVFEYFVNTSDPGDHIFTLRQVVVQRPITWASDADQAISIIGDFKWVNVSITCDVYIEHLGNGGVFIAGRVNNGGIYVRHSKGLFFWVFADGTYQVTGDLSGKEVLMKGMSGVRARVWHTLTLNLKGSSASGSLNGYPLWENVTLSETSHGWAAIGTRSFEFAQFDNFHIEAS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2K3</accession>
    <name>A0A452J2K3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">Glutamate receptor</fullName>
        </recommendedName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="21">Receptor for glutamate that functions as a ligand-gated ion channel in the central nervous system and plays an important role in excitatory synaptic transmission. L-glutamate acts as an excitatory neurotransmitter at many synapses in the central nervous system.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>Ca(2+)(in) = Ca(2+)(out)</text>
            <dbReference type="Rhea" id="RHEA:29671"/>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="16 21">Postsynaptic cell membrane</location>
            <topology evidence="16 21">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="21">Belongs to the glutamate-gated ion channel (TC 1.A.10.1) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2K3"/>
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        <property type="protein sequence ID" value="ENSGAGP00000034629.1"/>
        <property type="gene ID" value="ENSGAGG00000024584.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015276">
        <property type="term" value="F:ligand-gated monoatomic ion channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0038023">
        <property type="term" value="F:signaling receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd06382">
        <property type="entry name" value="PBP1_iGluR_Kainate"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.2300:FF:000010">
        <property type="entry name" value="Glutamate ionotropic receptor kainate type subunit 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.190.10:FF:000210">
        <property type="entry name" value="Glutamate receptor ionotropic, kainate 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.190.10:FF:000240">
        <property type="entry name" value="Glutamate receptor ionotropic, kainate 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.70:FF:000010">
        <property type="entry name" value="Putative glutamate receptor ionotropic kainate 1"/>
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    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="2"/>
    </dbReference>
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        <property type="entry name" value="Periplasmic binding protein-like II"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="ANF_lig-bd_rcpt"/>
    </dbReference>
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        <property type="entry name" value="Glu/Gly-bd"/>
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        <property type="entry name" value="Iono_Glu_rcpt_met"/>
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        <property type="entry name" value="Ionotropic_Glu_rcpt"/>
    </dbReference>
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        <property type="entry name" value="Iontro_rcpt_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028082">
        <property type="entry name" value="Peripla_BP_I"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18966">
        <property type="entry name" value="IONOTROPIC GLUTAMATE RECEPTOR"/>
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    </dbReference>
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        <property type="entry name" value="ANF_receptor"/>
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    </dbReference>
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        <property type="entry name" value="Lig_chan"/>
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    </dbReference>
    <dbReference type="Pfam" id="PF10613">
        <property type="entry name" value="Lig_chan-Glu_bd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00177">
        <property type="entry name" value="NMDARECEPTOR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00918">
        <property type="entry name" value="Lig_chan-Glu_bd"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="PBPe"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Periplasmic binding protein-like I"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Periplasmic binding protein-like II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 21" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="10 20" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="15 21" id="KW-0407">Ion channel</keyword>
    <keyword evidence="8 21" id="KW-0406">Ion transport</keyword>
    <keyword evidence="14 21" id="KW-1071">Ligand-gated ion channel</keyword>
    <keyword evidence="9 21" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="13 21" id="KW-0628">Postsynaptic cell membrane</keyword>
    <keyword evidence="11 21" id="KW-0675">Receptor</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <keyword evidence="7 21" id="KW-0770">Synapse</keyword>
    <keyword evidence="4 21" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 21" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="1 21" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="541"/>
            <end position="560"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="617"/>
            <end position="639"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="798"/>
            <end position="822"/>
        </location>
    </feature>
    <feature type="domain" description="Ionotropic glutamate receptor C-terminal" evidence="22">
        <location>
            <begin position="410"/>
            <end position="779"/>
        </location>
    </feature>
    <feature type="domain" description="Ionotropic glutamate receptor L-glutamate and glycine-binding" evidence="23">
        <location>
            <begin position="420"/>
            <end position="485"/>
        </location>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="494"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="501"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="668"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="716"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="site" description="Crucial to convey clamshell closure to channel opening" evidence="19">
        <location>
            <position position="646"/>
        </location>
    </feature>
    <feature type="site" description="Interaction with the cone snail toxin Con-ikot-ikot" evidence="19">
        <location>
            <position position="673"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="74"/>
            <end position="325"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="728"/>
            <end position="782"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023257"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00034104"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00036634"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-1"/>
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    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="RuleBase" id="RU367118"/>
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    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="SMART" id="SM00079"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="SMART" id="SM00918"/>
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    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034629.1"/>
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    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="886" mass="99955" checksum="87B48FA58F43AE8F" modified="2019-05-08" version="1">GMLLEKGRGWGRETVFRGIFECVESGPMGAEELAFRFAVNTINRNRTLLPNTTLTYDTQKINLYDSFEASKKACDQLSLGVAAIFGPSHSSSANAVQSICNALGVPHIQTRWKHQVSDNKDSFYVSLYPDFSSLSRAILDLVQFFKWKTVTVVYDDSTGLIRLQELIKAPSRYNLRLKIRQLPADTKDAKPLLKEMKRGKEFHVIFDCSHEMAAGILKQALAMGMMTEYYHYIFTTLDLFALDVEPYRYSGVNMTGFRILNTENSQVASIIEKWSMERLQAPPKPDSGLLDGFMTTDAALMYDAVHVVSVAVQQFPQMTVSSLQCNRHKPWRFGTRFMSLIKEAHWEGLTGRITFNKTNGLRTDFDLDVISLKEEGLEKIGTWDPASGLNMTENQKGKPANITDSLSNRSLIVTTILEEPYVMFKKSDKPLYGNDRFEGYCIDLLRELATILGFTYEIRLVEDGKYGAQDDASGQWNGMVRELIDHKADLAVAPLAITYVREKVIDFSKPFMTLGISILYRKPNGTNPGVFSFLNPLSPDIWMYILLAYLGVSCVLFVIARFSPYEWYNPHPCNPDSDVVENNFTLLNSFWFGVGALMQQGSELMPKALSTRIVGGIWWFFTLIIISSYTANLAAFLTVERMESPIDSADDLAKQTKIEYGAVEDGATMTFFKKSKISTYDKMWAFMSSRRQSVLVKSNEEGIQRVLTSDYAFLMESTTIEFVTQRNCNLTQIGGLIDSKGYGVGTPMGSPYRDKITIAILQLQEEGKLHMMKEKWWRGNGCPEEESKEASALGVQNIGGIFIVLAAGLVLSVFVAVGEFLYKSKKNAQLEKRSFCSAMVEELRMSLKCQRRLKHKPQAPVIVKTEEVINMHTFNDRRLPGKETMA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2K6</accession>
    <name>A0A452J2K6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">Glutamate receptor</fullName>
        </recommendedName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="21">Receptor for glutamate that functions as a ligand-gated ion channel in the central nervous system and plays an important role in excitatory synaptic transmission. L-glutamate acts as an excitatory neurotransmitter at many synapses in the central nervous system.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>Ca(2+)(in) = Ca(2+)(out)</text>
            <dbReference type="Rhea" id="RHEA:29671"/>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="16 21">Postsynaptic cell membrane</location>
            <topology evidence="16 21">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="21">Belongs to the glutamate-gated ion channel (TC 1.A.10.1) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2K6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039211.1">
        <property type="protein sequence ID" value="ENSGAGP00000034636.1"/>
        <property type="gene ID" value="ENSGAGG00000024584.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015276">
        <property type="term" value="F:ligand-gated monoatomic ion channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0038023">
        <property type="term" value="F:signaling receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd06382">
        <property type="entry name" value="PBP1_iGluR_Kainate"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.2300:FF:000010">
        <property type="entry name" value="Glutamate ionotropic receptor kainate type subunit 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.190.10:FF:000210">
        <property type="entry name" value="Glutamate receptor ionotropic, kainate 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.190.10:FF:000240">
        <property type="entry name" value="Glutamate receptor ionotropic, kainate 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.70:FF:000010">
        <property type="entry name" value="Putative glutamate receptor ionotropic kainate 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.70">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.2300">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.190.10">
        <property type="entry name" value="Periplasmic binding protein-like II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001828">
        <property type="entry name" value="ANF_lig-bd_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019594">
        <property type="entry name" value="Glu/Gly-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001508">
        <property type="entry name" value="Iono_Glu_rcpt_met"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015683">
        <property type="entry name" value="Ionotropic_Glu_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001320">
        <property type="entry name" value="Iontro_rcpt_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028082">
        <property type="entry name" value="Peripla_BP_I"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18966">
        <property type="entry name" value="IONOTROPIC GLUTAMATE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01094">
        <property type="entry name" value="ANF_receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00060">
        <property type="entry name" value="Lig_chan"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10613">
        <property type="entry name" value="Lig_chan-Glu_bd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00177">
        <property type="entry name" value="NMDARECEPTOR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00918">
        <property type="entry name" value="Lig_chan-Glu_bd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00079">
        <property type="entry name" value="PBPe"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53822">
        <property type="entry name" value="Periplasmic binding protein-like I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53850">
        <property type="entry name" value="Periplasmic binding protein-like II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 21" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="10 20" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="15 21" id="KW-0407">Ion channel</keyword>
    <keyword evidence="8 21" id="KW-0406">Ion transport</keyword>
    <keyword evidence="14 21" id="KW-1071">Ligand-gated ion channel</keyword>
    <keyword evidence="9 21" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="13 21" id="KW-0628">Postsynaptic cell membrane</keyword>
    <keyword evidence="11 21" id="KW-0675">Receptor</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <keyword evidence="7 21" id="KW-0770">Synapse</keyword>
    <keyword evidence="4 21" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 21" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="1 21" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="531"/>
            <end position="550"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="607"/>
            <end position="629"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="788"/>
            <end position="812"/>
        </location>
    </feature>
    <feature type="domain" description="Ionotropic glutamate receptor C-terminal" evidence="22">
        <location>
            <begin position="400"/>
            <end position="769"/>
        </location>
    </feature>
    <feature type="domain" description="Ionotropic glutamate receptor L-glutamate and glycine-binding" evidence="23">
        <location>
            <begin position="410"/>
            <end position="475"/>
        </location>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="484"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="491"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="658"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="706"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="site" description="Crucial to convey clamshell closure to channel opening" evidence="19">
        <location>
            <position position="636"/>
        </location>
    </feature>
    <feature type="site" description="Interaction with the cone snail toxin Con-ikot-ikot" evidence="19">
        <location>
            <position position="663"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="64"/>
            <end position="315"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="718"/>
            <end position="772"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023257"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00034104"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00036634"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="RuleBase" id="RU367118"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="SMART" id="SM00079"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="SMART" id="SM00918"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034636.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="859" mass="96565" checksum="FCB51A376E64CBCC" modified="2019-05-08" version="1">HILVLTRGIFECVESGPMGAEELAFRFAVNTINRNRTLLPNTTLTYDTQKINLYDSFEASKKACDQLSLGVAAIFGPSHSSSANAVQSICNALGVPHIQTRWKHQVSDNKDSFYVSLYPDFSSLSRAILDLVQFFKWKTVTVVYDDSTGLIRLQELIKAPSRYNLRLKIRQLPADTKDAKPLLKEMKRGKEFHVIFDCSHEMAAGILKQALAMGMMTEYYHYIFTTLDLFALDVEPYRYSGVNMTGFRILNTENSQVASIIEKWSMERLQAPPKPDSGLLDGFMTTDAALMYDAVHVVSVAVQQFPQMTVSSLQCNRHKPWRFGTRFMSLIKEAHWEGLTGRITFNKTNGLRTDFDLDVISLKEEGLEKIGTWDPASGLNMTENQKGKPANITDSLSNRSLIVTTILEEPYVMFKKSDKPLYGNDRFEGYCIDLLRELATILGFTYEIRLVEDGKYGAQDDASGQWNGMVRELIDHKADLAVAPLAITYVREKVIDFSKPFMTLGISILYRKPNGTNPGVFSFLNPLSPDIWMYILLAYLGVSCVLFVIARFSPYEWYNPHPCNPDSDVVENNFTLLNSFWFGVGALMQQGSELMPKALSTRIVGGIWWFFTLIIISSYTANLAAFLTVERMESPIDSADDLAKQTKIEYGAVEDGATMTFFKKSKISTYDKMWAFMSSRRQSVLVKSNEEGIQRVLTSDYAFLMESTTIEFVTQRNCNLTQIGGLIDSKGYGVGTPMGSPYRDKITIAILQLQEEGKLHMMKEKWWRGNGCPEEESKEASALGVQNIGGIFIVLAAGLVLSVFVAVGEFLYKSKKNINIIFCGFTTDYDLDSCNLNSSSFETANVNLSCTARLHKYGN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2K9</accession>
    <name>A0A452J2K9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="8">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J2K9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039231.1">
        <property type="protein sequence ID" value="ENSGAGP00000034655.1"/>
        <property type="gene ID" value="ENSGAGG00000024614.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070182">
        <property type="term" value="F:DNA polymerase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006281">
        <property type="term" value="P:DNA repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd11720">
        <property type="entry name" value="FANCI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026171">
        <property type="entry name" value="FANCI"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029310">
        <property type="entry name" value="FANCI_HD1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029312">
        <property type="entry name" value="FANCI_HD2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029308">
        <property type="entry name" value="FANCI_S1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029305">
        <property type="entry name" value="FANCI_S1-cap"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029315">
        <property type="entry name" value="FANCI_S2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029313">
        <property type="entry name" value="FANCI_S3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029314">
        <property type="entry name" value="FANCI_S4"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21818">
        <property type="entry name" value="BC025462 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21818:SF0">
        <property type="entry name" value="FANCONI ANEMIA GROUP I PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14679">
        <property type="entry name" value="FANCI_HD1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14680">
        <property type="entry name" value="FANCI_HD2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14675">
        <property type="entry name" value="FANCI_S1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14674">
        <property type="entry name" value="FANCI_S1-cap"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14676">
        <property type="entry name" value="FANCI_S2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14677">
        <property type="entry name" value="FANCI_S3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14678">
        <property type="entry name" value="FANCI_S4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="FANCI solenoid 1 cap" evidence="1">
        <location>
            <begin position="33"/>
            <end position="85"/>
        </location>
    </feature>
    <feature type="domain" description="FANCI solenoid 1" evidence="2">
        <location>
            <begin position="96"/>
            <end position="313"/>
        </location>
    </feature>
    <feature type="domain" description="FANCI helical" evidence="6">
        <location>
            <begin position="318"/>
            <end position="404"/>
        </location>
    </feature>
    <feature type="domain" description="FANCI solenoid 2" evidence="3">
        <location>
            <begin position="412"/>
            <end position="574"/>
        </location>
    </feature>
    <feature type="domain" description="FANCI helical" evidence="7">
        <location>
            <begin position="589"/>
            <end position="822"/>
        </location>
    </feature>
    <feature type="domain" description="FANCI solenoid 3" evidence="4">
        <location>
            <begin position="840"/>
            <end position="1066"/>
        </location>
    </feature>
    <feature type="domain" description="FANCI solenoid 4" evidence="5">
        <location>
            <begin position="1080"/>
            <end position="1127"/>
        </location>
    </feature>
    <evidence type="ECO:0000259" key="1">
        <source>
            <dbReference type="Pfam" id="PF14674"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF14675"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="Pfam" id="PF14676"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="Pfam" id="PF14677"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="Pfam" id="PF14678"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="Pfam" id="PF14679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF14680"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034655.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1151" mass="128362" checksum="3A339BB7380705E1" modified="2019-05-08" version="1">MMPSPADPTMTCKDGEHIRCVTVLQSLMTRATMDQKILSLAAEDALDKLEECLQNLKESELDDLITRQAVKGKETAVLLRGIFKGSPCSQQNGVLRRLQVYKHCIQLVESGDLPLDIASEIIGLLMLEARQLPGAALAELATLFVDVIKGGSLSNGKSLELFSSVLTALATSKESLVYGKGELSGEEFKKQLINTLCSSRWNHQCVIHLTNMFRDIPLSAEELQFVVEKVLRMFSKLDLQEIPPLVYQLLLLSAKGSKKTVLEGIIAFFNQLDKRQKEEQRSPESVDLEAATVPLDQLRHVEGTIILHIISVINLDQDLGRELIKLLKAEQQRDLGKTLCPFSVALVLSVAIKHRFEEQVFDFLKMSIVRSCKDLQFLQGSKFLQDLFPQQYDVASIILEVVENSAFGWDHVTQGLVNLGFTLMESYEPKKVLGKAAETSYGLCKTPAQQACRLGANILLETFKVHEPIRSAILEQVLNRVITKAASPINHFIDLLSNIVVSAPLVLQNSSSKVTEAFDNLSFLPLDMVQGLLKAVQPLLKVSMSVRDSLILVLRKAIFSRQLDARKAAVAGFLLLLKNFKVLGSLAFSQCSQAIGASQVQADIHACYNSAANEAFCLEILGSLRRCLSQQADIRLMLYEGFYDVLRRNSQLSNSIMETLLSQIKRYYEPEPDLLPPLKLEGCIVAQGDQIFLQEPLAHLLSCIQHCLAWYKSTMQLHHEAEEEEDEDDDYVGFQQDLEEMLESITRRMIKSELEDFELDKSADFSLSSGVGVKNNIYAILVMGVCEVLIEYNFNIGNFSKNKSEDVLGLFKCYSKLSEILKEKAGKNKSSLANKTARSFLSMSFVSTLLTALFRDSAQSHEESLAVLRSSSEFMCYTINVALQKVQQLEETGQTDGPDGQNPEKMFQALCKITRVLLWRYTSIPTAVEESSGKKKGKSISLLCLEGLLKIFNTVQQLYKPKIPQFLRALDIANDDAEEEEDINVTEKAAFQIRQFQRSLVNQLSSAGDDFNSKEALSLVTVISTLSKLLEPSSPQFIQLFSWTVKICKENSLEDASCCKGLLILLFNLHVLYKSPVGLLREFAQDIHAHLGDIDQDMEVENQSHFAIVNTKTAAPTACVSWNSICKTQSLRYYCAAGAKTHLGTCGLQNF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2L1</accession>
    <name>A0A452J2L1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">Glutamate receptor</fullName>
        </recommendedName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="21">Receptor for glutamate that functions as a ligand-gated ion channel in the central nervous system and plays an important role in excitatory synaptic transmission. L-glutamate acts as an excitatory neurotransmitter at many synapses in the central nervous system.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>Ca(2+)(in) = Ca(2+)(out)</text>
            <dbReference type="Rhea" id="RHEA:29671"/>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="16 21">Postsynaptic cell membrane</location>
            <topology evidence="16 21">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="21">Belongs to the glutamate-gated ion channel (TC 1.A.10.1) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2L1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039214.1">
        <property type="protein sequence ID" value="ENSGAGP00000034639.1"/>
        <property type="gene ID" value="ENSGAGG00000024584.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015276">
        <property type="term" value="F:ligand-gated monoatomic ion channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0038023">
        <property type="term" value="F:signaling receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd06382">
        <property type="entry name" value="PBP1_iGluR_Kainate"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.2300:FF:000010">
        <property type="entry name" value="Glutamate ionotropic receptor kainate type subunit 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.190.10:FF:000210">
        <property type="entry name" value="Glutamate receptor ionotropic, kainate 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.190.10:FF:000240">
        <property type="entry name" value="Glutamate receptor ionotropic, kainate 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.70:FF:000010">
        <property type="entry name" value="Putative glutamate receptor ionotropic kainate 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.70">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.2300">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.190.10">
        <property type="entry name" value="Periplasmic binding protein-like II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001828">
        <property type="entry name" value="ANF_lig-bd_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019594">
        <property type="entry name" value="Glu/Gly-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001508">
        <property type="entry name" value="Iono_Glu_rcpt_met"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015683">
        <property type="entry name" value="Ionotropic_Glu_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001320">
        <property type="entry name" value="Iontro_rcpt_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028082">
        <property type="entry name" value="Peripla_BP_I"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18966">
        <property type="entry name" value="IONOTROPIC GLUTAMATE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01094">
        <property type="entry name" value="ANF_receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00060">
        <property type="entry name" value="Lig_chan"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10613">
        <property type="entry name" value="Lig_chan-Glu_bd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00177">
        <property type="entry name" value="NMDARECEPTOR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00918">
        <property type="entry name" value="Lig_chan-Glu_bd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00079">
        <property type="entry name" value="PBPe"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53822">
        <property type="entry name" value="Periplasmic binding protein-like I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53850">
        <property type="entry name" value="Periplasmic binding protein-like II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 21" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="10 20" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="15 21" id="KW-0407">Ion channel</keyword>
    <keyword evidence="8 21" id="KW-0406">Ion transport</keyword>
    <keyword evidence="14 21" id="KW-1071">Ligand-gated ion channel</keyword>
    <keyword evidence="9 21" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="13 21" id="KW-0628">Postsynaptic cell membrane</keyword>
    <keyword evidence="11 21" id="KW-0675">Receptor</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <keyword evidence="7 21" id="KW-0770">Synapse</keyword>
    <keyword evidence="4 21" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 21" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="1 21" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="534"/>
            <end position="553"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="610"/>
            <end position="632"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="791"/>
            <end position="815"/>
        </location>
    </feature>
    <feature type="domain" description="Ionotropic glutamate receptor C-terminal" evidence="22">
        <location>
            <begin position="403"/>
            <end position="772"/>
        </location>
    </feature>
    <feature type="domain" description="Ionotropic glutamate receptor L-glutamate and glycine-binding" evidence="23">
        <location>
            <begin position="413"/>
            <end position="478"/>
        </location>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="487"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="494"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="661"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="709"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="site" description="Crucial to convey clamshell closure to channel opening" evidence="19">
        <location>
            <position position="639"/>
        </location>
    </feature>
    <feature type="site" description="Interaction with the cone snail toxin Con-ikot-ikot" evidence="19">
        <location>
            <position position="666"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="67"/>
            <end position="318"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="721"/>
            <end position="775"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023257"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00034104"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00036634"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="RuleBase" id="RU367118"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="SMART" id="SM00079"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="SMART" id="SM00918"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034639.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="834" mass="94052" checksum="5E5C3C62AEBBF784" modified="2019-05-08" version="1">MGSKVPENAKGVLFDWESELLIFKELAFRFAVNTINRNRTLLPNTTLTYDTQKINLYDSFEASKKACDQLSLGVAAIFGPSHSSSANAVQSICNALGVPHIQTRWKHQVSDNKDSFYVSLYPDFSSLSRAILDLVQFFKWKTVTVVYDDSTGLIRLQELIKAPSRYNLRLKIRQLPADTKDAKPLLKEMKRGKEFHVIFDCSHEMAAGILKQALAMGMMTEYYHYIFTTLDLFALDVEPYRYSGVNMTGFRILNTENSQVASIIEKWSMERLQAPPKPDSGLLDGFMTTDAALMYDAVHVVSVAVQQFPQMTVSSLQCNRHKPWRFGTRFMSLIKEAHWEGLTGRITFNKTNGLRTDFDLDVISLKEEGLEKIGTWDPASGLNMTENQKGKPANITDSLSNRSLIVTTILEEPYVMFKKSDKPLYGNDRFEGYCIDLLRELATILGFTYEIRLVEDGKYGAQDDASGQWNGMVRELIDHKADLAVAPLAITYVREKVIDFSKPFMTLGISILYRKPNGTNPGVFSFLNPLSPDIWMYILLAYLGVSCVLFVIARFSPYEWYNPHPCNPDSDVVENNFTLLNSFWFGVGALMQQGSELMPKALSTRIVGGIWWFFTLIIISSYTANLAAFLTVERMESPIDSADDLAKQTKIEYGAVEDGATMTFFKKSKISTYDKMWAFMSSRRQSVLVKSNEEGIQRVLTSDYAFLMESTTIEFVTQRNCNLTQIGGLIDSKGYGVGTPMGSPYRDKITIAILQLQEEGKLHMMKEKWWRGNGCPEEESKEASALGVQNIGGIFIVLAAGLVLSVFVAVGEFLYKSKKNAQLEKVNIVFFHLI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2L3</accession>
    <name>A0A452J2L3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="24">Glutamate receptor</fullName>
        </recommendedName>
    </protein>
    <organism evidence="27 28">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="28" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="27" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="27" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="18">Independent of its ionotropic glutamate receptor activity, acts as a thermoreceptor conferring sensitivity to cold temperatures. Functions in dorsal root ganglion neurons.</text>
    </comment>
    <comment type="function">
        <text evidence="19">Ionotropic glutamate receptor that functions as a cation-permeable ligand-gated ion channel, gated by L-glutamate and the glutamatergic agonist kainic acid. L-glutamate acts as an excitatory neurotransmitter at many synapses in the central nervous system. Binding of the excitatory neurotransmitter L-glutamate induces a conformation change, leading to the opening of the cation channel, and thereby converts the chemical signal to an electrical impulse. The receptor then desensitizes rapidly and enters a transient inactive state, characterized by the presence of bound agonist.</text>
    </comment>
    <comment type="function">
        <text evidence="24">Receptor for glutamate that functions as a ligand-gated ion channel in the central nervous system and plays an important role in excitatory synaptic transmission. L-glutamate acts as an excitatory neurotransmitter at many synapses in the central nervous system.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="16">
            <text>Ca(2+)(in) = Ca(2+)(out)</text>
            <dbReference type="Rhea" id="RHEA:29671"/>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>Na(+)(in) = Na(+)(out)</text>
            <dbReference type="Rhea" id="RHEA:34963"/>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
        </reaction>
    </comment>
    <comment type="subunit">
        <text evidence="20">Homotetramer and heterotetramer with GRIK5. Tetramers may be formed by the dimerization of dimers.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="14 24">Postsynaptic cell membrane</location>
            <topology evidence="14 24">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="17">Belongs to the glutamate-gated ion channel (TC 1.A.10.1) family. GRIK2 subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2L3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039216.1">
        <property type="protein sequence ID" value="ENSGAGP00000034641.1"/>
        <property type="gene ID" value="ENSGAGG00000024584.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015276">
        <property type="term" value="F:ligand-gated monoatomic ion channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0038023">
        <property type="term" value="F:signaling receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd06382">
        <property type="entry name" value="PBP1_iGluR_Kainate"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.2300:FF:000010">
        <property type="entry name" value="Glutamate ionotropic receptor kainate type subunit 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.190.10:FF:000240">
        <property type="entry name" value="Glutamate receptor ionotropic, kainate 2"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="glutamate receptor ionotropic, kainate 4"/>
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    </dbReference>
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        <property type="entry name" value="Putative glutamate receptor ionotropic kainate 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.70">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.2300">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.190.10">
        <property type="entry name" value="Periplasmic binding protein-like II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001828">
        <property type="entry name" value="ANF_lig-bd_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019594">
        <property type="entry name" value="Glu/Gly-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001508">
        <property type="entry name" value="Iono_Glu_rcpt_met"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015683">
        <property type="entry name" value="Ionotropic_Glu_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001320">
        <property type="entry name" value="Iontro_rcpt_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028082">
        <property type="entry name" value="Peripla_BP_I"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18966">
        <property type="entry name" value="IONOTROPIC GLUTAMATE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01094">
        <property type="entry name" value="ANF_receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00060">
        <property type="entry name" value="Lig_chan"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10613">
        <property type="entry name" value="Lig_chan-Glu_bd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00177">
        <property type="entry name" value="NMDARECEPTOR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00918">
        <property type="entry name" value="Lig_chan-Glu_bd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00079">
        <property type="entry name" value="PBPe"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53822">
        <property type="entry name" value="Periplasmic binding protein-like I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53850">
        <property type="entry name" value="Periplasmic binding protein-like II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 24" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="8 23" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13 24" id="KW-0407">Ion channel</keyword>
    <keyword evidence="6 24" id="KW-0406">Ion transport</keyword>
    <keyword evidence="12 24" id="KW-1071">Ligand-gated ion channel</keyword>
    <keyword evidence="7 24" id="KW-0472">Membrane</keyword>
    <keyword evidence="11 24" id="KW-0628">Postsynaptic cell membrane</keyword>
    <keyword evidence="9 24" id="KW-0675">Receptor</keyword>
    <keyword evidence="28" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 24" id="KW-0770">Synapse</keyword>
    <keyword evidence="3 24" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 24" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="1 24" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="24">
        <location>
            <begin position="476"/>
            <end position="495"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="24">
        <location>
            <begin position="552"/>
            <end position="574"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="24">
        <location>
            <begin position="733"/>
            <end position="757"/>
        </location>
    </feature>
    <feature type="domain" description="Ionotropic glutamate receptor C-terminal" evidence="25">
        <location>
            <begin position="339"/>
            <end position="714"/>
        </location>
    </feature>
    <feature type="domain" description="Ionotropic glutamate receptor L-glutamate and glycine-binding" evidence="26">
        <location>
            <begin position="355"/>
            <end position="420"/>
        </location>
    </feature>
    <feature type="binding site" evidence="21">
        <location>
            <position position="429"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="21">
        <location>
            <position position="436"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="21">
        <location>
            <position position="603"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="21">
        <location>
            <position position="651"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="site" description="Crucial to convey clamshell closure to channel opening" evidence="22">
        <location>
            <position position="581"/>
        </location>
    </feature>
    <feature type="site" description="Interaction with the cone snail toxin Con-ikot-ikot" evidence="22">
        <location>
            <position position="608"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="23">
        <location>
            <begin position="47"/>
            <end position="298"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="23">
        <location>
            <begin position="663"/>
            <end position="717"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023257"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023286"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00034104"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00036239"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00036634"/>
        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00045840"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00059880"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="RuleBase" id="RU367118"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="SMART" id="SM00079"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="26">
        <source>
            <dbReference type="SMART" id="SM00918"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034641.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="28">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="776" mass="87681" checksum="E73F21E50B147642" modified="2019-05-08" version="1">MGAEELAFRFAVNTINRNRTLLPNTTLTYDTQKINLYDSFEASKKACDQLSLGVAAIFGPSHSSSANAVQSICNALGVPHIQTRWKHQVSDNKDSFYVSLYPDFSSLSRAILDLVQFFKWKTVTVVYDDSTGLIRLQELIKAPSRYNLRLKIRQLPADTKDAKPLLKEMKRGKEFHVIFDCSHEMAAGILKQALAMGMMTEYYHYIFTTLDLFALDVEPYRYSGVNMTGFRILNTENSQVASIIEKWSMERLQAPPKPDSGLLDGFMTTDAALMYDAVHVVSVAVQQFPQMTVSSLQCNRHKPWRFGTRFMSLIKEAHWEGLTGRITFNKTNGLRTDFDLDVISLKEEGLEKEEPYVMFKKSDKPLYGNDRFEGYCIDLLRELATILGFTYEIRLVEDGKYGAQDDASGQWNGMVRELIDHKADLAVAPLAITYVREKVIDFSKPFMTLGISILYRKPNGTNPGVFSFLNPLSPDIWMYILLAYLGVSCVLFVIARFSPYEWYNPHPCNPDSDVVENNFTLLNSFWFGVGALMQQGSELMPKALSTRIVGGIWWFFTLIIISSYTANLAAFLTVERMESPIDSADDLAKQTKIEYGAVEDGATMTFFKKSKISTYDKMWAFMSSRRQSVLVKSNEEGIQRVLTSDYAFLMESTTIEFVTQRNCNLTQIGGLIDSKGYGVGTPMGSPYRDKITIAILQLQEEGKLHMMKEKWWRGNGCPEEESKEASALGVQNIGGIFIVLAAGLVLSVFVAVGEFLYKSKKNAQLEKVNIVFFHLI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2L8</accession>
    <name>A0A452J2L8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Torsin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Endomembrane system</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3 11">Endoplasmic reticulum lumen</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4 11">Belongs to the ClpA/ClpB family. Torsin subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2L8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039221.1">
        <property type="protein sequence ID" value="ENSGAGP00000034645.1"/>
        <property type="gene ID" value="ENSGAGG00000024621.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000039222.1">
        <property type="protein sequence ID" value="ENSGAGP00000034646.1"/>
        <property type="gene ID" value="ENSGAGG00000024621.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005788">
        <property type="term" value="C:endoplasmic reticulum lumen"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005635">
        <property type="term" value="C:nuclear envelope"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019894">
        <property type="term" value="F:kinesin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071763">
        <property type="term" value="P:nuclear membrane organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034504">
        <property type="term" value="P:protein localization to nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000743">
        <property type="entry name" value="Torsin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049337">
        <property type="entry name" value="TOR1A_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010448">
        <property type="entry name" value="Torsin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017378">
        <property type="entry name" value="Torsin_1/2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10760">
        <property type="entry name" value="TORSIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10760:SF14">
        <property type="entry name" value="TORSIN-1B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21376">
        <property type="entry name" value="TOR1A_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06309">
        <property type="entry name" value="Torsin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF038079">
        <property type="entry name" value="Torsin_2A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="7 11" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="9" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="8" id="KW-0472">Membrane</keyword>
    <keyword evidence="12" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 13" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="13">
        <location>
            <begin position="1"/>
            <end position="17"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5044604615" description="Torsin" evidence="13">
        <location>
            <begin position="18"/>
            <end position="334"/>
        </location>
    </feature>
    <feature type="domain" description="Torsin-1A C-terminal" evidence="14">
        <location>
            <begin position="273"/>
            <end position="330"/>
        </location>
    </feature>
    <feature type="binding site" evidence="12">
        <location>
            <begin position="103"/>
            <end position="110"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004308"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00006235"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PIRNR" id="PIRNR038079"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PIRSR" id="PIRSR038079-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="Pfam" id="PF21376"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034646.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="334" mass="37742" checksum="BE78B125007176D6" modified="2019-05-08" version="1">MQLLAGLLWLLVPGSAALEPISLSIVIGAASVLTGYLSYPSFYCGFVECCPGEADRPNTTALREDLDNKLFGQHLAKDVILKAVTGFRNNPNPKKPLTLSLHGWGGTGKNFVSQFIAKSIHKAGQKSKFVHLFIATLHFPHEHQITLYKDQLQTWIQGNVSACARSVFIFDEMDKLHPGLIDAIKPFLDYYEQIDGVSYRRAIFIFLSNAGGDLINKVALDFWRSGKDRKQIQLKDLEPVLSVAVFNNKNSGLWHSSLIDKNLIDYFVPFLPLEYKHLKMCIRAEIEAGGKKINEAIVDEVAKEMTFFPKDEKIYSDKGCKLVQQKLLYYWERL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2M2</accession>
    <name>A0A452J2M2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Centrosomal protein of 120 kDa</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="8">Coiled-coil domain-containing protein 100</fullName>
        </alternativeName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Microtubule organizing center</location>
            <location evidence="1">Centrosome</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the CEP120 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2M2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039227.1">
        <property type="protein sequence ID" value="ENSGAGP00000034651.1"/>
        <property type="gene ID" value="ENSGAGG00000024622.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0022027">
        <property type="term" value="P:interkinetic nuclear migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903724">
        <property type="term" value="P:positive regulation of centriole elongation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032880">
        <property type="term" value="P:regulation of protein localization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000112">
        <property type="entry name" value="centrosomal protein of 120 kDa isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000008">
        <property type="entry name" value="C2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039893">
        <property type="entry name" value="CEP120-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022136">
        <property type="entry name" value="DUF3668"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21574">
        <property type="entry name" value="CENTROSOMAL PROTEIN OF 120 KDA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21574:SF0">
        <property type="entry name" value="CENTROSOMAL PROTEIN OF 120 KDA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00168">
        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12416">
        <property type="entry name" value="DUF3668"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49562">
        <property type="entry name" value="C2 domain (Calcium/lipid-binding domain, CaLB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50004">
        <property type="entry name" value="C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="5" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="C2" evidence="11">
        <location>
            <begin position="1"/>
            <end position="112"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="350"/>
            <end position="409"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="619"/>
            <end position="645"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="669"/>
            <end position="922"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="368"/>
            <end position="379"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="380"/>
            <end position="392"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="636"/>
            <end position="645"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00060869"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00071268"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00076226"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034651.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="986" mass="113083" checksum="542449EEB6AAE5A1" modified="2019-05-08" version="1">MVSKSDQLLIVVSILEGRHFPRRAKHMLVVEAKFDGEQLATDPVDHADQPEFATELAWELDRKALHQHRLQRTPIKLQCFALDPRSSTKENIGYIVLDLRAAQEKKQAPKWYPLLSNKYIKFKSEIQISIMLETDTKTPVDGFKAKEAPPREGKVPALLSGLDPKNIVAVLNEEEGYHQIGPAEYCREYFVLSVTIAFATQLEQLIPSTMKLPERQPEFFFYYSLLGNDVTNEPFTDLINPNFEPERASVRIRSTVEVLQVYFSAQPKIQIHLCCGDQSLGSTEIPLTGLLKKGNVEIDQQPVAVEGAFNLVPPNRAKQKLPPLALDMAPTVGVSVTLQKEGINTQSLLHLDLQTEPEHSQKRVLSATRDKTKSPEQRSHSAPSAPLQNGHSPPTKDEATESEVESLQYDRGTKLNKKALNALAPVSQSHPGAASGAESTSGQKIAVPAASHHFCFSIDLRSIHSLEVGFPVNCILRYSYPFFGSAAPIMTNPPIEVRKNMEVFLPQSYCAFDFATMSHQLQDTFFRLPLLVELWHKDKMTKDILLGVARLQLSNVLGLEKTRFLGTNGEQCWRQTCSETVGVMAAQGANNRIAELSYTVTLEDYGLVKMHEVLVSDSSQCSGAGQQQVPPPQPHHVPETEPEPRETLEYKAALELEMWKEMQEDIFENQLKKKELAHMQALAEEWKKRDKEREALVKKKVAEYTNLEEQLQKTLTDLEKRERQLVSAESELQRVKRELQAQHERNFQELQDSVRRVKEECAHQVELERSKIKQLEEDKVRLQQHFYEAENKYKILEKEFQQYKEQQCSKPEIRLQSEINLLTLEKVELERKLESATKSKLHYKQQWARALKELARLKQREQENAMARLKKQQQELEHMRLRYLAAEEKEMGKTERKELEDIRNELNRLKQQEDKKQSQDSRDISVEQVGSLHTRQLNESLDDYLTRLIEERDTLLRTGVYNHEDHIVSELDRQIREALAKRSSSK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2M5</accession>
    <name>A0A452J2M5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4 9">Signal recognition particle subunit SRP72</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Component of the signal recognition particle (SRP) complex, a ribonucleoprotein complex that mediates the cotranslational targeting of secretory and membrane proteins to the endoplasmic reticulum (ER).</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2 9">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 9">Belongs to the SRP72 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2M5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039246.1">
        <property type="protein sequence ID" value="ENSGAGP00000034670.1"/>
        <property type="gene ID" value="ENSGAGG00000024611.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005783">
        <property type="term" value="C:endoplasmic reticulum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005786">
        <property type="term" value="C:signal recognition particle, endoplasmic reticulum targeting"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008312">
        <property type="term" value="F:7S RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043022">
        <property type="term" value="F:ribosome binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006614">
        <property type="term" value="P:SRP-dependent cotranslational protein targeting to membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000133">
        <property type="entry name" value="Signal recognition particle subunit SRP72"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000191">
        <property type="entry name" value="Signal recognition particle subunit SRP72"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000369">
        <property type="entry name" value="Signal recognition particle subunit SRP72"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.10">
        <property type="entry name" value="Tetratricopeptide repeat domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013699">
        <property type="entry name" value="Signal_recog_part_SRP72_RNA-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026270">
        <property type="entry name" value="SRP72"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031545">
        <property type="entry name" value="SRP72_TPR-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011990">
        <property type="entry name" value="TPR-like_helical_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019734">
        <property type="entry name" value="TPR_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14094">
        <property type="entry name" value="SIGNAL RECOGNITION PARTICLE 72"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14094:SF9">
        <property type="entry name" value="SIGNAL RECOGNITION PARTICLE SUBUNIT SRP72"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08492">
        <property type="entry name" value="SRP72"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17004">
        <property type="entry name" value="SRP_TPR_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13181">
        <property type="entry name" value="TPR_8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF038922">
        <property type="entry name" value="SRP72"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00028">
        <property type="entry name" value="TPR"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48452">
        <property type="entry name" value="TPR-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50005">
        <property type="entry name" value="TPR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 9" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8 9" id="KW-0687">Ribonucleoprotein</keyword>
    <keyword evidence="7 9" id="KW-0733">Signal recognition particle</keyword>
    <keyword evidence="10" id="KW-0802">TPR repeat</keyword>
    <feature type="repeat" description="TPR" evidence="10">
        <location>
            <begin position="173"/>
            <end position="206"/>
        </location>
    </feature>
    <feature type="domain" description="Signal recognition particle SRP72 subunit RNA-binding" evidence="12">
        <location>
            <begin position="480"/>
            <end position="533"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="473"/>
            <end position="616"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="11">
        <location>
            <begin position="497"/>
            <end position="507"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="509"/>
            <end position="530"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="544"/>
            <end position="566"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="574"/>
            <end position="584"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="11">
        <location>
            <begin position="600"/>
            <end position="616"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007676"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00018350"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023135"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023274"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PIRNR" id="PIRNR038922"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00339"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF08492"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034670.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="616" mass="68741" checksum="BCBF906B1D2FBC14" modified="2019-05-08" version="1">ARVSGAGQDSPVGWMWLAGCSLPTDSITFEKAYCEYRLNRIENALKTIQSASQQTDKLKELHGQVLYRLERYDDCLAVYRDLIRNSQDDYEEERKTNLSAVVAAQSTWEKVEPEDLGLQEATYELCYNAACALIGQGRLNEAMENLQKAEDLCRQSLSEDSDVTEEDIEAELAIIHGQMAYIMQLQGRTEDALQLYNQIIKLKPTDVGLIAVIANNIITINKDQNVFDSKKKVKLTNADGVEHKLSKKQLQAIEFNKALLAMYTNQADQCRKLSASLQSQNPEHLLPVLIQAAQLCREKQHAKAIELLQDFAEQSPADSAQIKLTMAQLKIAQGSVTKACIILRSIEELKHKPGMVSALVTMYSHEEDIDSAIEVFTQAIQWYQEYQPKSPVHLLLIREAANFKLKYGRKKEAISDLEELWKQNPKDIHTLAQLISAYSLVDPEKAKVLSKHLPSSDTMSLKVDVDALENSPGATYVRKKGGKLTGENQQKEQGDVKKKKRKKKGKLPKNYDPKVTPDPERWLPMRERSYYRGRKKGKKKDQIGKGTQGATTAGASEDASKTASSPPTSPRLGSSTAMSSATTSNMIPPRHQKPAGAAATKKKQQQKKKKGGKGGW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2M6</accession>
    <name>A0A452J2M6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Centrosomal protein of 120 kDa</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="8">Coiled-coil domain-containing protein 100</fullName>
        </alternativeName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Microtubule organizing center</location>
            <location evidence="1">Centrosome</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the CEP120 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2M6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039230.1">
        <property type="protein sequence ID" value="ENSGAGP00000034654.1"/>
        <property type="gene ID" value="ENSGAGG00000024622.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0022027">
        <property type="term" value="P:interkinetic nuclear migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903724">
        <property type="term" value="P:positive regulation of centriole elongation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032880">
        <property type="term" value="P:regulation of protein localization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000112">
        <property type="entry name" value="centrosomal protein of 120 kDa isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000008">
        <property type="entry name" value="C2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039893">
        <property type="entry name" value="CEP120-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022136">
        <property type="entry name" value="DUF3668"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21574">
        <property type="entry name" value="CENTROSOMAL PROTEIN OF 120 KDA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21574:SF0">
        <property type="entry name" value="CENTROSOMAL PROTEIN OF 120 KDA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00168">
        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12416">
        <property type="entry name" value="DUF3668"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49562">
        <property type="entry name" value="C2 domain (Calcium/lipid-binding domain, CaLB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50004">
        <property type="entry name" value="C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="5" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="C2" evidence="11">
        <location>
            <begin position="1"/>
            <end position="112"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="284"/>
            <end position="343"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="553"/>
            <end position="579"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="603"/>
            <end position="856"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="302"/>
            <end position="313"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="314"/>
            <end position="326"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="570"/>
            <end position="579"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00060869"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00071268"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00076226"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034654.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="920" mass="105378" checksum="C923FCC47367FE3D" modified="2019-05-08" version="1">MVSKSDQLLIVVSILEGRHFPRRAKHMLVVEAKFDGEQLATDPVDHADQPEFATELAWELDRKALHQHRLQRTPIKLQCFALDPRSSTKENIGYIVLDLRAAQEKKQAPKWYPLLSNKYIKFKSEIQISIMLETDTKTPVDGFKAKEAPPREGKVPALLSGLDPKNIVAVLNEEEGYHQIGPAEYCREYFVLSVTIAFATQLEQIHLCCGDQSLGSTEIPLTGLLKKGNVEIDQQPVAVEGAFNLVPPNRAKQKLPPLALDMAPTVGVSVTLQKEGINTQSLLHLDLQTEPEHSQKRVLSATRDKTKSPEQRSHSAPSAPLQNGHSPPTKDEATESEVESLQYDRGTKLNKKALNALAPVSQSHPGAASGAESTSGQKIAVPAASHHFCFSIDLRSIHSLEVGFPVNCILRYSYPFFGSAAPIMTNPPIEVRKNMEVFLPQSYCAFDFATMSHQLQDTFFRLPLLVELWHKDKMTKDILLGVARLQLSNVLGLEKTRFLGTNGEQCWRQTCSETVGVMAAQGANNRIAELSYTVTLEDYGLVKMHEVLVSDSSQCSGAGQQQVPPPQPHHVPETEPEPRETLEYKAALELEMWKEMQEDIFENQLKKKELAHMQALAEEWKKRDKEREALVKKKVAEYTNLEEQLQKTLTDLEKRERQLVSAESELQRVKRELQAQHERNFQELQDSVRRVKEECAHQVELERSKIKQLEEDKVRLQQHFYEAENKYKILEKEFQQYKEQQCSKPEIRLQSEINLLTLEKVELERKLESATKSKLHYKQQWARALKELARLKQREQENAMARLKKQQQELEHMRLRYLAAEEKEMGKTERKELEDIRNELNRLKQQEDKKQSQDSRDISVEQVGSLHTRQLNESLDDYLTRLIEERDTLLRTGVYNHEDHIVSELDRQIREALAKRSSSK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2M8</accession>
    <name>A0A452J2M8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Putative histone-lysine N-methyltransferase PRDM6</fullName>
            <ecNumber evidence="2">2.1.1.361</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="19">PR domain zinc finger protein 6</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="20">PR domain-containing protein 6</fullName>
        </alternativeName>
    </protein>
    <organism evidence="25 26">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="26" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="25" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="25" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="16">Putative histone methyltransferase that acts as a transcriptional repressor of smooth muscle gene expression. Promotes the transition from differentiated to proliferative smooth muscle by suppressing differentiation and maintaining the proliferative potential of vascular smooth muscle cells. Also plays a role in endothelial cells by inhibiting endothelial cell proliferation, survival and differentiation. It is unclear whether it has histone methyltransferase activity in vivo. According to some authors, it does not act as a histone methyltransferase by itself and represses transcription by recruiting EHMT2/G9a. According to others, it possesses histone methyltransferase activity when associated with other proteins and specifically methylates 'Lys-20' of histone H4 in vitro. 'Lys-20' methylation represents a specific tag for epigenetic transcriptional repression.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>L-lysyl(20)-[histone H4] + S-adenosyl-L-methionine = N(6)-methyl-L-lysyl(20)-[histone H4] + S-adenosyl-L-homocysteine + H(+)</text>
            <dbReference type="Rhea" id="RHEA:60344"/>
            <dbReference type="Rhea" id="RHEA-COMP:15554"/>
            <dbReference type="Rhea" id="RHEA-COMP:15555"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29969"/>
            <dbReference type="ChEBI" id="CHEBI:57856"/>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
            <dbReference type="ChEBI" id="CHEBI:61929"/>
            <dbReference type="EC" id="2.1.1.361"/>
        </reaction>
    </comment>
    <comment type="subunit">
        <text evidence="17">Interacts with HDAC1, HDAC2, HDAC3, CBX1 and EP300.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="2.1.1.361" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452J2M8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039238.1">
        <property type="protein sequence ID" value="ENSGAGP00000034662.1"/>
        <property type="gene ID" value="ENSGAGG00000024630.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140944">
        <property type="term" value="F:histone H4K20 monomethyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042802">
        <property type="term" value="F:identical protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032259">
        <property type="term" value="P:methylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
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    <dbReference type="GO" id="GO:0045892">
        <property type="term" value="P:negative regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd19191">
        <property type="entry name" value="PR-SET_PRDM6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000654">
        <property type="entry name" value="PR/SET domain 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000778">
        <property type="entry name" value="PR/SET domain 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.170.270.10:FF:000023">
        <property type="entry name" value="putative histone-lysine N-methyltransferase PRDM6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000868">
        <property type="entry name" value="putative histone-lysine N-methyltransferase PRDM6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.170.270.10">
        <property type="entry name" value="SET domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044416">
        <property type="entry name" value="PRDM6_PR-SET"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001214">
        <property type="entry name" value="SET_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046341">
        <property type="entry name" value="SET_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050331">
        <property type="entry name" value="Zinc_finger"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16515:SF22">
        <property type="entry name" value="HISTONE-LYSINE N-METHYLTRANSFERASE PRDM6-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16515">
        <property type="entry name" value="PR DOMAIN ZINC FINGER PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21549">
        <property type="entry name" value="PRDM2_PR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00317">
        <property type="entry name" value="SET"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF82199">
        <property type="entry name" value="SET domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50280">
        <property type="entry name" value="SET"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="11" id="KW-0156">Chromatin regulator</keyword>
    <keyword evidence="7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="4" id="KW-0489">Methyltransferase</keyword>
    <keyword evidence="14" id="KW-0539">Nucleus</keyword>
    <keyword evidence="26" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0678">Repressor</keyword>
    <keyword evidence="6" id="KW-0949">S-adenosyl-L-methionine</keyword>
    <keyword evidence="13" id="KW-0804">Transcription</keyword>
    <keyword evidence="12" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="5" id="KW-0808">Transferase</keyword>
    <keyword evidence="10" id="KW-0862">Zinc</keyword>
    <keyword evidence="9 21" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="SET" evidence="24">
        <location>
            <begin position="218"/>
            <end position="337"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="445"/>
            <end position="472"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="473"/>
            <end position="500"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="501"/>
            <end position="528"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="529"/>
            <end position="556"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="31"/>
            <end position="79"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="22">
        <location>
            <begin position="60"/>
            <end position="79"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00012187"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022491"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022603"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022691"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00047784"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00054067"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00063348"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00074801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00076320"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00082168"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS50280"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034662.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="567" mass="62458" checksum="61D448261D46AD4E" modified="2019-05-08" version="1">MLKPGDPGGSAFLKVDPAYLQHWQQLFPQSSQLKSSGALTQLPPPERADPSADSLRQRPASLSSASSSSSSSTPSSSTSSCAASAASLAGLTSISVAQIPVFGPLPSSESPSGAPASLQSKDLCGGGAKCSERAAPRFRCTAEELDYYLYGQQRMEIIPLNQHTSDPNNRCDMCADNRNGECPMHGPLHSLRRLVGTSSAAAAAPPPEIPEWLRDLPREVCLCTSTVPGLAYGICAAQRIQQGTWIGPFQGVVLLPEKVQTGAIRNTQHLWEIYDQDGTLQHFIDGGEPSKSSWMRYIRCARHCGEQNLTVVQYRSNIFYRACIDIPRGTELLVWYNDSYTSFFGIPLQCIAQDENLNVPSTVMEAMSRPDALQPFNKSSKLPPATQQRSIVFPQTPCSRNFSLLDKSGPIESGFNQINVKNQRVLASPTSTSQLNSEFSDWHLWKCGQCFKTFTQRILLQMHVCTQNPDRPYQCGHCSQSFSQPSELRNHVVTHSSDRPFKCGYCGRAFAGATTLNNHIRTHTGEKPFKCERCERSFTQATQLSRHQRMPNECKPITESTESIEVD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2N0</accession>
    <name>A0A452J2N0_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="8">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J2N0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039235.1">
        <property type="protein sequence ID" value="ENSGAGP00000034659.1"/>
        <property type="gene ID" value="ENSGAGG00000024614.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070182">
        <property type="term" value="F:DNA polymerase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006281">
        <property type="term" value="P:DNA repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd11720">
        <property type="entry name" value="FANCI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026171">
        <property type="entry name" value="FANCI"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029310">
        <property type="entry name" value="FANCI_HD1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029312">
        <property type="entry name" value="FANCI_HD2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029308">
        <property type="entry name" value="FANCI_S1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029305">
        <property type="entry name" value="FANCI_S1-cap"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029315">
        <property type="entry name" value="FANCI_S2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029313">
        <property type="entry name" value="FANCI_S3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029314">
        <property type="entry name" value="FANCI_S4"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21818">
        <property type="entry name" value="BC025462 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21818:SF0">
        <property type="entry name" value="FANCONI ANEMIA GROUP I PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14679">
        <property type="entry name" value="FANCI_HD1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14680">
        <property type="entry name" value="FANCI_HD2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14675">
        <property type="entry name" value="FANCI_S1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14674">
        <property type="entry name" value="FANCI_S1-cap"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14676">
        <property type="entry name" value="FANCI_S2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14677">
        <property type="entry name" value="FANCI_S3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14678">
        <property type="entry name" value="FANCI_S4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="FANCI solenoid 1 cap" evidence="1">
        <location>
            <begin position="6"/>
            <end position="58"/>
        </location>
    </feature>
    <feature type="domain" description="FANCI solenoid 1" evidence="2">
        <location>
            <begin position="69"/>
            <end position="286"/>
        </location>
    </feature>
    <feature type="domain" description="FANCI helical" evidence="6">
        <location>
            <begin position="291"/>
            <end position="377"/>
        </location>
    </feature>
    <feature type="domain" description="FANCI solenoid 2" evidence="3">
        <location>
            <begin position="385"/>
            <end position="547"/>
        </location>
    </feature>
    <feature type="domain" description="FANCI helical" evidence="7">
        <location>
            <begin position="562"/>
            <end position="795"/>
        </location>
    </feature>
    <feature type="domain" description="FANCI solenoid 3" evidence="4">
        <location>
            <begin position="813"/>
            <end position="1039"/>
        </location>
    </feature>
    <feature type="domain" description="FANCI solenoid 4" evidence="5">
        <location>
            <begin position="1053"/>
            <end position="1100"/>
        </location>
    </feature>
    <evidence type="ECO:0000259" key="1">
        <source>
            <dbReference type="Pfam" id="PF14674"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF14675"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="Pfam" id="PF14676"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="Pfam" id="PF14677"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="Pfam" id="PF14678"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="Pfam" id="PF14679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF14680"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034659.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1124" mass="125419" checksum="59A347CCA57D08CD" modified="2019-05-08" version="1">MTRATMDQKILSLAAEDALDKLEECLQNLKESELDDLITRQAVKGKETAVLLRGIFKGSPCSQQNGVLRRLQVYKHCIQLVESGDLPLDIASEIIGLLMLEARQLPGAALAELATLFVDVIKGGSLSNGKSLELFSSVLTALATSKESLVYGKGELSGEEFKKQLINTLCSSRWNHQCVIHLTNMFRDIPLSAEELQFVVEKVLRMFSKLDLQEIPPLVYQLLLLSAKGSKKTVLEGIIAFFNQLDKRQKEEQRSPESVDLEAATVPLDQLRHVEGTIILHIISVINLDQDLGRELIKLLKAEQQRDLGKTLCPFSVALVLSVAIKHRFEEQVFDFLKMSIVRSCKDLQFLQGSKFLQDLFPQQYDVASIILEVVENSAFGWDHVTQGLVNLGFTLMESYEPKKVLGKAAETSYGLCKTPAQQACRLGANILLETFKVHEPIRSAILEQVLNRVITKAASPINHFIDLLSNIVVSAPLVLQNSSSKVTEAFDNLSFLPLDMVQGLLKAVQPLLKVSMSVRDSLILVLRKAIFSRQLDARKAAVAGFLLLLKNFKVLGSLAFSQCSQAIGASQVQADIHACYNSAANEAFCLEILGSLRRCLSQQADIRLMLYEGFYDVLRRNSQLSNSIMETLLSQIKRYYEPEPDLLPPLKLEGCIVAQGDQIFLQEPLAHLLSCIQHCLAWYKSTMQLHHEAEEEEDEDDDYVGFQQDLEEMLESITRRMIKSELEDFELDKSADFSLSSGVGVKNNIYAILVMGVCEVLIEYNFNIGNFSKNKSEDVLGLFKCYSKLSEILKEKAGKNKSSLANKTARSFLSMSFVSTLLTALFRDSAQSHEESLAVLRSSSEFMCYTINVALQKVQQLEETGQTDGPDGQNPEKMFQALCKITRVLLWRYTSIPTAVEESSGKKKGKSISLLCLEGLLKIFNTVQQLYKPKIPQFLRALDIANDDAEEEEDINVTEKAAFQIRQFQRSLVNQLSSAGDDFNSKEALSLVTVISTLSKLLEPSSPQFIQLFSWTVKICKENSLEDASCCKGLLILLFNLHVLYKSPVGLLREFAQDIHAHLGDIDQDMEVENQSHFAIVNTKTAAPTACVSWNSICKTQSLRYYCAAGAKTHLGTCGLQNF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2N1</accession>
    <name>A0A452J2N1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4 9">Signal recognition particle subunit SRP72</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Component of the signal recognition particle (SRP) complex, a ribonucleoprotein complex that mediates the cotranslational targeting of secretory and membrane proteins to the endoplasmic reticulum (ER).</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2 9">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 9">Belongs to the SRP72 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2N1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039252.1">
        <property type="protein sequence ID" value="ENSGAGP00000034675.1"/>
        <property type="gene ID" value="ENSGAGG00000024611.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005783">
        <property type="term" value="C:endoplasmic reticulum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005786">
        <property type="term" value="C:signal recognition particle, endoplasmic reticulum targeting"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008312">
        <property type="term" value="F:7S RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043022">
        <property type="term" value="F:ribosome binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006614">
        <property type="term" value="P:SRP-dependent cotranslational protein targeting to membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000133">
        <property type="entry name" value="Signal recognition particle subunit SRP72"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000191">
        <property type="entry name" value="Signal recognition particle subunit SRP72"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000369">
        <property type="entry name" value="Signal recognition particle subunit SRP72"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.10">
        <property type="entry name" value="Tetratricopeptide repeat domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013699">
        <property type="entry name" value="Signal_recog_part_SRP72_RNA-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026270">
        <property type="entry name" value="SRP72"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011990">
        <property type="entry name" value="TPR-like_helical_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019734">
        <property type="entry name" value="TPR_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14094">
        <property type="entry name" value="SIGNAL RECOGNITION PARTICLE 72"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14094:SF9">
        <property type="entry name" value="SIGNAL RECOGNITION PARTICLE SUBUNIT SRP72"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08492">
        <property type="entry name" value="SRP72"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13181">
        <property type="entry name" value="TPR_8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF038922">
        <property type="entry name" value="SRP72"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00028">
        <property type="entry name" value="TPR"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48452">
        <property type="entry name" value="TPR-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50005">
        <property type="entry name" value="TPR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 9" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8 9" id="KW-0687">Ribonucleoprotein</keyword>
    <keyword evidence="7 9" id="KW-0733">Signal recognition particle</keyword>
    <keyword evidence="10" id="KW-0802">TPR repeat</keyword>
    <feature type="repeat" description="TPR" evidence="10">
        <location>
            <begin position="160"/>
            <end position="193"/>
        </location>
    </feature>
    <feature type="domain" description="Signal recognition particle SRP72 subunit RNA-binding" evidence="12">
        <location>
            <begin position="466"/>
            <end position="522"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="460"/>
            <end position="606"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="11">
        <location>
            <begin position="486"/>
            <end position="496"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="498"/>
            <end position="519"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="533"/>
            <end position="556"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="564"/>
            <end position="574"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="11">
        <location>
            <begin position="590"/>
            <end position="606"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007676"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00018350"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023135"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023274"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PIRNR" id="PIRNR038922"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00339"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF08492"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034675.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="606" mass="67840" checksum="CDA4D98A57360854" modified="2019-05-08" version="1">IYSLMRISQNWVTGASNKAYCEYRLNRIENALKTIQSASQQTDKLKELHGQVLYRLERYDDCLAVYRDLIRNSQDDYEEERKTNLSAVVAAQSTWEKVEPEDLGLQEATYELCYNAACALIGQGRLNEAMENLQKAEDLCRQSLSEDSDVTEEDIEAELAIIHGQMAYIMQLQGRTEDALQLYNQIIKLKPTDVGLIAVIANNIITINKDQNVFDSKKKVKLTNADGVEHKLSKKQLQAIEFNKALLAMYTNQADQCRKLSASLQSQNPEHLLPVLIQAAQLCREKQHAKAIELLQDFAEQSPADSAQIKLTMAQLKIAQGSVTKACIILRSIEELKHKPGMVSALVTMYSHEEDIDSAIEVFTQAIQWYQEYQPKSPVHLLLIREAANFKLKYGRKKEAISDLEELWKQNPKDIHTLAQLISAYSLVDPEKAKVLSKHLPSSDTMSLKVDVDALENSPGATYVRKKGGKLTGENQQKEQGQGDVKKKKRKKKGKLPKNYDPKVTPDPERWLPMRERSYYRGRKKGKKKDQIGKGTQGATTAGASELDASKTASSPPTSPRLGSSTAMSSATTSNMIPPRHQKPAGAAATKKKQQQKKKKGGKGGW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2N4</accession>
    <name>A0A452J2N4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">small monomeric GTPase</fullName>
            <ecNumber evidence="3">3.6.5.2</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>GTP + H2O = GDP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:19669"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:37565"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:58189"/>
            <dbReference type="EC" id="3.6.5.2"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="13">
            <dbReference type="Rhea" id="RHEA:19670"/>
        </physiologicalReaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the small GTPase superfamily. Rab family.</text>
    </comment>
    <dbReference type="EC" id="3.6.5.2" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452J2N4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039240.1">
        <property type="protein sequence ID" value="ENSGAGP00000034664.1"/>
        <property type="gene ID" value="ENSGAGG00000024632.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003925">
        <property type="term" value="F:G protein activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005525">
        <property type="term" value="F:GTP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd04112">
        <property type="entry name" value="Rab26"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000459">
        <property type="entry name" value="ras-related protein Rab-37 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005225">
        <property type="entry name" value="Small_GTP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001806">
        <property type="entry name" value="Small_GTPase"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00231">
        <property type="entry name" value="small_GTP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47978">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00071">
        <property type="entry name" value="Ras"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00449">
        <property type="entry name" value="RASTRNSFRMNG"/>
    </dbReference>
    <dbReference type="SMART" id="SM00177">
        <property type="entry name" value="ARF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00175">
        <property type="entry name" value="RAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00176">
        <property type="entry name" value="RAN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00173">
        <property type="entry name" value="RAS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00174">
        <property type="entry name" value="RHO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51419">
        <property type="entry name" value="RAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51421">
        <property type="entry name" value="RAS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51420">
        <property type="entry name" value="RHO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0342">GTP-binding</keyword>
    <keyword evidence="7" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="11" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="8" id="KW-0460">Magnesium</keyword>
    <keyword evidence="5" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="6" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="12" id="KW-0636">Prenylation</keyword>
    <keyword evidence="9" id="KW-0653">Protein transport</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0813">Transport</keyword>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006270"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00011984"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023289"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00047660"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034664.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="227" mass="25075" checksum="4F2EBBECCEFB835B" modified="2019-05-08" version="1">MSGRAGSAAPAQNGGPAAAPERSPSLTQGYDLACKVMLLGDSGVGKTCFLLQFKDGAFLSGTFIATVGIDFRNKVVTVDDVKVKLQIWDTAGQERFRSVTHAYYRDAQALLLFYDITSKMSFDNIRAWLTEIHEYAQKDVVIMLLGNKADVSSGRVIKMEDGESLAREYGVPFMETSAKTGMNVELAFLAIAKELKQRAVQQPDEPRFQIQEYIESQKKKSSCCSFM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2N5</accession>
    <name>A0A452J2N5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Myosin IG</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="6">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000039149.1">
        <property type="protein sequence ID" value="ENSGAGP00000034577.1"/>
        <property type="gene ID" value="ENSGAGG00000024461.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005902">
        <property type="term" value="C:microvillus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016459">
        <property type="term" value="C:myosin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000146">
        <property type="term" value="F:microfilament motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007015">
        <property type="term" value="P:actin filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030048">
        <property type="term" value="P:actin filament-based movement"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006897">
        <property type="term" value="P:endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.4820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010926">
        <property type="entry name" value="Myosin_TH1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13140">
        <property type="entry name" value="MYOSIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13140:SF381">
        <property type="entry name" value="UNCONVENTIONAL MYOSIN-IG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06017">
        <property type="entry name" value="Myosin_TH1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51757">
        <property type="entry name" value="TH1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 6" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="2 6" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="4 6" id="KW-0505">Motor protein</keyword>
    <keyword evidence="3 6" id="KW-0518">Myosin</keyword>
    <keyword evidence="1 6" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="7">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5045551665" description="Myosin IG" evidence="7">
        <location>
            <begin position="23"/>
            <end position="959"/>
        </location>
    </feature>
    <feature type="domain" description="Myosin motor" evidence="8">
        <location>
            <begin position="1"/>
            <end position="654"/>
        </location>
    </feature>
    <feature type="domain" description="TH1" evidence="9">
        <location>
            <begin position="771"/>
            <end position="958"/>
        </location>
    </feature>
    <feature type="region of interest" description="Actin-binding" evidence="6">
        <location>
            <begin position="531"/>
            <end position="553"/>
        </location>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <begin position="72"/>
            <end position="79"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51757"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034577.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="12">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="959" mass="110272" checksum="63B385A7E971B49B" modified="2025-10-08" version="2">MXXXXXXXXXLVLTILSSLVSSADFATSLFTPFSRSFMNKLNRIGPRTDPWGTTKPRPSCCPLPILSVCLAGESGAGKTEASKYIMQYIAAITNPSQREEVERVKNVLLKSNCVLEAFGNAKTNRNDNSSRFGKYMDINFDFKGDPTGGHINNYLLEKSRILHQQPGERNFHSFYQVSWELLISLLLFPPPHPISQLLASDQSNFKDVADAMKVIGFSPEEIDSVYKILATILHLGNVSFVAEGDAVAIENRELVSQLAELTSTQPESLLRTLLYRAVATGGGEVIEKGHSAKEANYARDACAKAIYERLFCWIVGRINTVIEVKNYDARLHGKNTVIAVLDIYGFEIFDSNSFEQFCINYCNEKLQQLFIELILRQEQEEYQREGIEWQHIEYFNNQIIVDLVEQPHKGIISILDEACLTGGQVTDAMFLESMNTKLGKHPHYTSRKLCATDKSMEFGRDFRIKHYAGNVTYSVEGFIDKNKDTLFQDFKRLMYNSTDPVLRDMWPEGEVSITEVTKRPLTAATLFKNSMVALVENLASKEPYYVRCIKPNDQKSPLLFDAERCRHQVEYLGLLENVRVRRAGFAYRQPYDRFLLRYKMTCEYTWPNHLMPTDQEATRALIDQHGFQDDVAYGHTKLFIRMPQILFCLEQERAQLIPIIVLLLQKAWRGALARRRCRYLRAIYTIMGHYRRHKVKTYLLELVRRFQGVRSMADYGKGVEWPDPPAVLAQFQENSKLLFCRWRARQIVKNIPPSDMGQIQAKVAAMEVLQGLRKDWGCQRNWGRDYLSSVSVRPGELPGALRDKVHFSSVLFSCHVRKINRFNKSRDRAILVTDQHLYKLEPRKQYKVMRTLPLSMVTGVSVTSCQDQLVVFHTKNHDDLVICFHKMQPEEDNRVGELVGVLVHHFKTTKRELQVRVSDCIQLSLRGRKRLVAVEMRHELAFPDFGKSRDGFVLYCPRR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2N6</accession>
    <name>A0A452J2N6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Signal recognition particle subunit SRP72</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the SRP72 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2N6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039257.1">
        <property type="protein sequence ID" value="ENSGAGP00000034680.1"/>
        <property type="gene ID" value="ENSGAGG00000024611.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005783">
        <property type="term" value="C:endoplasmic reticulum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005786">
        <property type="term" value="C:signal recognition particle, endoplasmic reticulum targeting"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008312">
        <property type="term" value="F:7S RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043022">
        <property type="term" value="F:ribosome binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006614">
        <property type="term" value="P:SRP-dependent cotranslational protein targeting to membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000133">
        <property type="entry name" value="Signal recognition particle subunit SRP72"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000191">
        <property type="entry name" value="Signal recognition particle subunit SRP72"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000369">
        <property type="entry name" value="Signal recognition particle subunit SRP72"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.10">
        <property type="entry name" value="Tetratricopeptide repeat domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013699">
        <property type="entry name" value="Signal_recog_part_SRP72_RNA-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026270">
        <property type="entry name" value="SRP72"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011990">
        <property type="entry name" value="TPR-like_helical_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019734">
        <property type="entry name" value="TPR_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14094">
        <property type="entry name" value="SIGNAL RECOGNITION PARTICLE 72"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14094:SF9">
        <property type="entry name" value="SIGNAL RECOGNITION PARTICLE SUBUNIT SRP72"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08492">
        <property type="entry name" value="SRP72"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13181">
        <property type="entry name" value="TPR_8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF038922">
        <property type="entry name" value="SRP72"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00028">
        <property type="entry name" value="TPR"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48452">
        <property type="entry name" value="TPR-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50005">
        <property type="entry name" value="TPR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0687">Ribonucleoprotein</keyword>
    <keyword evidence="7" id="KW-0733">Signal recognition particle</keyword>
    <keyword evidence="9" id="KW-0802">TPR repeat</keyword>
    <feature type="repeat" description="TPR" evidence="9">
        <location>
            <begin position="160"/>
            <end position="193"/>
        </location>
    </feature>
    <feature type="domain" description="Signal recognition particle SRP72 subunit RNA-binding" evidence="11">
        <location>
            <begin position="465"/>
            <end position="496"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="464"/>
            <end position="580"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="472"/>
            <end position="493"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="507"/>
            <end position="530"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="538"/>
            <end position="548"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="10">
        <location>
            <begin position="564"/>
            <end position="580"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007676"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00018350"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023135"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023274"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00339"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF08492"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034680.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="580" mass="64932" checksum="3FBA78F95BD3C4EE" modified="2019-05-08" version="1">IYSLMRISQNWVTGASNKAYCEYRLNRIENALKTIQSASQQTDKLKELHGQVLYRLERYDDCLAVYRDLIRNSQDDYEEERKTNLSAVVAAQSTWEKVEPEDLGLQEATYELCYNAACALIGQGRLNEAMENLQKAEDLCRQSLSEDSDVTEEDIEAELAIIHGQMAYIMQLQGRTEDALQLYNQIIKLKPTDVGLIAVIANNIITINKDQNVFDSKKKVKLTNADGVEHKLSKKQLQAIEFNKALLAMYTNQADQCRKLSASLQSQNPEHLLPVLIQAAQLCREKQHAKAIELLQDFAEQSPADSAQIKLTMAQLKIAQGSVTKACIILRSIEELKHKPGMVSALVTMYSHEEDIDSAIEVFTQAIQWYQEYQPKSPVHLLLIREAANFKLKYGRKKEAISDLEELWKQNPKDIHTLAQLISAYSLVDPEKAKVLSKHLPSSDTMSLKVDVDALENSPGATYVRKKGKLPKNYDPKVTPDPERWLPMRERSYYRGRKKGKKKDQIGKGTQGATTAGASELDASKTASSPPTSPRLGSSTAMSSATTSNMIPPRHQKPAGAAATKKKQQQKKKKGGKGGW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2N7</accession>
    <name>A0A452J2N7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Sorting nexin-2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">Involved in several stages of intracellular trafficking. Interacts with membranes containing phosphatidylinositol 3-phosphate (PtdIns(3P)) or phosphatidylinositol 3,5-bisphosphate (PtdIns(3,5)P2). Acts in part as component of the retromer membrane-deforming SNX-BAR subcomplex. The SNX-BAR retromer mediates retrograde transport of cargo proteins from endosomes to the trans-Golgi network (TGN) and is involved in endosome-to-plasma membrane transport for cargo protein recycling. The SNX-BAR subcomplex functions to deform the donor membrane into a tubular profile called endosome-to-TGN transport carrier (ETC). Can sense membrane curvature and has in vitro vesicle-to-membrane remodeling activity. Required for retrograde endosome-to-TGN transport of TGN38. Promotes KALRN- and RHOG-dependent but retromer-independent membrane remodeling such as lamellipodium formation; the function is dependent on GEF activity of KALRN.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell projection</location>
            <location evidence="2">Lamellipodium</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Early endosome membrane</location>
            <topology evidence="1">Peripheral membrane protein</topology>
            <orientation evidence="1">Cytoplasmic side</orientation>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the sorting nexin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2N7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039248.1">
        <property type="protein sequence ID" value="ENSGAGP00000034672.1"/>
        <property type="gene ID" value="ENSGAGG00000024638.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="GOC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031901">
        <property type="term" value="C:early endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030027">
        <property type="term" value="C:lamellipodium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030904">
        <property type="term" value="C:retromer complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035091">
        <property type="term" value="F:phosphatidylinositol binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034498">
        <property type="term" value="P:early endosome to Golgi transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006886">
        <property type="term" value="P:intracellular protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd07664">
        <property type="entry name" value="BAR_SNX2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd07282">
        <property type="entry name" value="PX_SNX2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1270.60:FF:000012">
        <property type="entry name" value="Sorting nexin 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1520.10:FF:000016">
        <property type="entry name" value="Sorting nexin 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.60">
        <property type="entry name" value="Arfaptin homology (AH) domain/BAR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1520.10">
        <property type="entry name" value="Phox-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027267">
        <property type="entry name" value="AH/BAR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001683">
        <property type="entry name" value="PX_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036871">
        <property type="entry name" value="PX_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037918">
        <property type="entry name" value="SNX2_PX"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005329">
        <property type="entry name" value="Sorting_nexin_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015404">
        <property type="entry name" value="Vps5_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10555">
        <property type="entry name" value="SORTING NEXIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10555:SF31">
        <property type="entry name" value="SORTING NEXIN-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00787">
        <property type="entry name" value="PX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03700">
        <property type="entry name" value="Sorting_nexin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09325">
        <property type="entry name" value="Vps5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00312">
        <property type="entry name" value="PX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103657">
        <property type="entry name" value="BAR/IMD domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF64268">
        <property type="entry name" value="PX domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50195">
        <property type="entry name" value="PX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0007">Acetylation</keyword>
    <keyword evidence="12" id="KW-0966">Cell projection</keyword>
    <keyword evidence="14" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-0967">Endosome</keyword>
    <keyword evidence="10" id="KW-0446">Lipid-binding</keyword>
    <keyword evidence="11" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="8" id="KW-0653">Protein transport</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0813">Transport</keyword>
    <feature type="domain" description="PX" evidence="16">
        <location>
            <begin position="140"/>
            <end position="269"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="1"/>
            <end position="63"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="79"/>
            <end position="102"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="14">
        <location>
            <begin position="436"/>
            <end position="470"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="15">
        <location>
            <begin position="27"/>
            <end position="50"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004469"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004510"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00010883"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00020435"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023121"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00045620"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50195"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034672.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="519" mass="58676" checksum="08A875533C49591F" modified="2019-05-08" version="1">MAAEREPPPLGEARQPELEELEDGEDLFTSTVSTLESSPLSPEPASLPTEDISINSNGPKPAEIILDDDREDLFAEATEEVSLDSPEREPILSPDPSPAITPVTPTTLVTPRMESKSVTAPVIFDRSREEIEEEANGDVFDIEINVSDPEKVGDGMNAYMAYRVTTKTSLSMFHKNEFSVKRRFSDFLGLHSKLATKYLHVGYIVPPAPEKSIVGMTKVKVGKEDSSSTEFVEKRRAALERYLQRTVKHPTLLQDPDLRQFLESSELPRAVNTQALSGAGILRMVNKAADAVNKMTIKMNESDAWFEEKQQQFENLDQQLRKLHASVEALVCHRKELSANTAAFAKSAAMLGNSEDHTALSRALSQLAEVEEKIDQLHQEQAFADFYVFSELLGDYIRLIAAVKGVLDHRMKCWQKWQDAQVTLQKKREAEAKLQLANKPDKLQQAKDEIKEWETKVQQGERDFEQISKTIRKEVGRFEKERVKDFKTVIIKYLESLVQTQQQLIKYWEAFLPEAKAIA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J300</accession>
    <name>A0A452J300_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">UBA domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the VPS13 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000039377.1">
        <property type="protein sequence ID" value="ENSGAGP00000034784.1"/>
        <property type="gene ID" value="ENSGAGG00000024709.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006869">
        <property type="term" value="P:lipid transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007005">
        <property type="term" value="P:mitochondrion organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045053">
        <property type="term" value="P:protein retention in Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006623">
        <property type="term" value="P:protein targeting to vacuole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd23453">
        <property type="entry name" value="beta-trefoil_Ricin_VPS13D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd14306">
        <property type="entry name" value="UBA_VP13D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.8.10:FF:000107">
        <property type="entry name" value="Vacuolar protein sorting 13 homolog D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.8.10">
        <property type="entry name" value="DNA helicase RuvA subunit, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015940">
        <property type="entry name" value="UBA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009060">
        <property type="entry name" value="UBA-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041969">
        <property type="entry name" value="VP13D_UBA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026847">
        <property type="entry name" value="VPS13"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056747">
        <property type="entry name" value="VPS13-like_M"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026854">
        <property type="entry name" value="VPS13_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009543">
        <property type="entry name" value="VPS13_VAB"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16166:SF141">
        <property type="entry name" value="INTERMEMBRANE LIPID TRANSFER PROTEIN VPS13D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16166">
        <property type="entry name" value="VACUOLAR PROTEIN SORTING-ASSOCIATED PROTEIN VPS13"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25033">
        <property type="entry name" value="VPS13_M"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12624">
        <property type="entry name" value="VPS13_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25036">
        <property type="entry name" value="VPS13_VAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00165">
        <property type="entry name" value="UBA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46934">
        <property type="entry name" value="UBA-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50030">
        <property type="entry name" value="UBA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0445">Lipid transport</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0813">Transport</keyword>
    <feature type="domain" description="UBA" evidence="5">
        <location>
            <begin position="2276"/>
            <end position="2316"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="375"/>
            <end position="421"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="1382"/>
            <end position="1407"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="1704"/>
            <end position="1730"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="1759"/>
            <end position="1780"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="2246"/>
            <end position="2268"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="407"/>
            <end position="421"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="1759"/>
            <end position="1779"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="2251"/>
            <end position="2264"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006545"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034784.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="4003" mass="446561" checksum="0D09590396C63C5C" modified="2019-05-08" version="1">MHRIEEEQTFEELKILREIVYDKFHDVAESVKEPLSESPGVSPTADSECSSSRGMLQYLQSWFPGWGGWYGYAQQTYEGEPFEGLLPETQEQWNPEEILGADEFFDPAADASSMNTFTKRDHVFAKLNLQLHGGTVTLLHKEKRAPRINESAFMQLEFSGVKILAESLPRRNSSLLTVQLGGLFLRDLATEGTIFPVLVFPNPQKEVGCVSSFGLQTSSAETTNQSTDADPNAPVFEMLYERNPAHSNFERRLEVSTRPLNVIYNPQAIKKVADFFYKGRVHTSGFGYQSELELRVAEAARRQYNKLKMQTKAEIRQTIDRLLVGDFIENSKRWTVCLDISAPQVIFPDDFKFVDPVLVVLDLGRILLTNSQDESKKKSAGGLSSEVNELSDNEYKTPLATPPNTPPSEATTSSGIKTSELTGVEISEEQLQAHLMSKKMYERYTLSFMDLQIMVGRVKDNWKHVQDSDVGPTHVVEKFNVHLQLERRLIYTSDPKYPGAVLSGNLPDLKIHINEDKISALKNCFARLSTSETKSPDALHIGKEKIFTDVDQRGSLHDSVMNLTQSVMMVEQHTREVLVESQLLLAEFKVNCMQLGVESSGRYISVLKVFGTNAHFVKRPYDAEVSLTVHGLLLVDTMQTYGSDFDLLMASHKNLSFDVPTGRLQDSRAQSPIYGPNEAYPIDVASLTEKCAAASNISSGNYAKDQESLIKLEYQFVSAECPSMNLDSTLQVISVQVNNLDIILNPETIVELIGFLQKSFPKEKEDSSPQPLMTDLERNFREQETFQSTHARNIEVAVDIHWLNILLLRTVGMTNGEKYGRKIATASIGGTKVNVSMCNKLDVNGSLGCLHLMDLTQDNVKNQYVVSIGNTVRYENLINDIGYFESLFVRLHEESSLPDALSFNFVEQSKEECSLNLKMASLHYNHSAKFLKELTLSMDELEENFCSMLKTAATKVTSVLATKTAEYSEMVSLFDTTPRSRDPFNLEDNEEDGFGPSSLKTDTIKLILNVNVESPIVSIPRKPGSPELLVGHLGQICIQNFVPGEDESRSDRLQVEIKDIKMYSLNSNQFMVRKESASEMTQGLHPPSSTPSVTSQEEANFTRHDFFESLHKGHAFHILNNTTIQFKLEKIPINRDLDLMFPLSNEDIGISGIMRIEGKFVNPLQVVLAKHVYEQVLQTLDNLIYNEDLNKFASSSTSSACPSSPSVSLRSVGSEFSNEQKENGLFSHSSFSAAPNKSVPVRETKSFTQIQANFCISELQVQLCGDLTLGAQGLVSLKFQDFDVEFAKDHPHTLSVQITLRSLLMEDLLEKCPDSIHKNLMVSRGAPKPSSLMHKEYLSQSCPSISHVEYPDMPRSLPSHMEEAPNVFQLYQRPTCASRKAHKEDADSEYPLTPPPSPTLNKSKVPSGKNNFDDSLVHINIFLVDKKHCEFSSRYNKINRSVDVDFNCLDVLITLQTWVVILDFFGIGSTADNHAMKLQPEDIQQMVKSEASILSTSEVQDPVNTKLNLKVHSLSLVLNQTTSELAKANVSKLVTYLEMIDGDLALQGSIGSLSLSDLTSHGELYRERFTTSGEEALIFQTYRYGRPDPLLQRECDIRVSLRMASVQYVHTQRFQAEVVSFIQHFTQLQDVLGRQRAAIEGQTVRDQAQRTSRVFLDIEAGAPVLLIPESSKSNNLIVANLGKLKVKNRFLFAGSPGTFSLKDKQGSVPFSSPLGTPKHSTKKTTSADESKGVISLKEGLFMKSSSGTNITNLKSESELSTLTKQQGQPPKSPVAQTSDSPEDHICLLDCIVVDLQDMDIFAAERYSREYSKAMEDTSADLTFPSYFVRQTGGSLLTEPFGLKLQVERNLDKEMSHAVPDISIHGNLSSVHCSLDLSKYKLIRGLLENNLGEPIEEFMRPYDLQDPRIHTVLSGEVYTCMSFLIDMVNVSLELVEPKGKDGLSSLARFDFKKSKLLFESFSNQTKSINLVSHSMMAFDTRYAGQKTTLGVPNVFNCIFQSSKNSSNKGAIQIELHFRSTKDSSCFTVVLNNLRVFLIFDWLLLVHDFLCTPSDVKKQDNLTSSRQRSSSCETTVVPKTVKSGVVTKRSSLQVTTEKQLEVKINVTGTEFVVVEDMSCFDTNAIILKGTTVLTYKPKLVDRPFSGSLFGIEVFSCRLGNEQDTALSIIDPVHIQMELVGNSSYPSGSGLLDAFNSEDFPPILEIQLQALDIRLSYNDVQLFLAIAKSIPGQTSAAVPESCVLDTDTAAKNLSNSSSENESSSSTSENRDISKHVLDPVLEVQLARLQELGFGMDDCRKALLICQGQLKKAASWLFKNAEPLKSLSLASGRRDSQGSVPIQLISGVEIKAESMCICFIDDCMDCDVPLAELTFSRLNFLQHVRASPEGYAHFTLSGDYYNRELSGWEPFIEPWPCSVSWQQQAASRLHPSRLKLEAKAKHRLDINITSVLTEQYTSTKQSWMADYCKQEKVDTVKSEDWMGSSVDPPSFGQTEMKTPKRRQPFVPFALRNHTGCTLWFATLTTTPTRAALSHSGSPGVVPEENGTFLDDAHNVSEWREVLTSGEIPFEFEARGKLRHRHTHDLRIHQLQVRVNGWEQVSPVSVDKVGTFFRYAAPDKNTSSSTVGSPISRTNIIHPHVYFSSLPPVRVVFEVTMEGSARKVITVRSALIVKNRLETPMELRLDSPSAPDKPVLLPAVMPGDSFAIPLHLTSWRLQARPKGMGVFFCKTPIHWTNVQKAAEVCSSKRECHSMESENSRFFRFCVAIKKENYPDYMPSNIFSDSAKQIFRQPGHTIYLLPTVVICNLLPCELEFYVKGMPINGTLKPGKEAALHTADTSQNIELGVSLENFPLCKELLIPPGTQNYMVRMRLYDVNKRQLNLTIRIVCRAEGSLKIFISAPYWLINKTGLPLIFRQDNAKTDAAGQFEEHELARSLSPLLFCYADKEQANFCTMRIGKGIHPRGMPGWCQGFSLDGGSGVRALKVIQQGNRPGLIYNIGIDVKKGRGRYIDTCMVTFAPRYLLDNKSSHKLAFAQREFARGQGTSNPDGYISTLPGSSVVFHWPRNDYDQLLCVRLMEVPHCVWSGGFEVNKNSSFHINMRDTLGKCYFLRVEITLRGATYRISFGDTDQLPPPFRIDNFSKVPVVFTQHGVAEPRLCTEVKPMTSLDYAWDEPTLPPFIMLTVKGAGSSEITCSMNDFQDSKQLYYENFIYIAATYTFSGLQEGTGRPVASNKDVACAELVLDVSPKTQRVILKKKEPGKRSQLWRMTGTGMLCHEGSSVPHNPNKPSSPRSVESSMILDIAGLAAVTDNRYEPLMLRKPDRRRSTTQTWSFRDGKLTCGMHGLVVQAKGGISGLFDGAEVVLGPDMSLELLGPIPPEQQFINQKMRPGSGVLAVRVVPDGPTRVLQITDFNQRRNDRLSCEGGELPVTEQDLRKLKNPDTDQELEVLVKLEGGIGLSLVNKVPEELVFASLTGINVHYTQLSTSQVLELSIQDVQVDNQLIGTTQPFMLFLTPQINENEAIETGPAVQLNAMKFPSKNALTDIYKHLMVTARRFTVQIEEKLLLKLLSFFGYDQSESEVEKYDENLHEKTVDQNGAQKRYYFENLKINVPQIKLSVFTSNKLPLDLKALKSTLGFPLIRFEDAVINLDPFTRVHPYETKEFIINDILKHFQEELLSQAARILGSVDFLGNPMGLLNDVSEGVTGLIKYGNVGGLIRNVTHGVSNSAAKFAGTLSDGLGKTMDNRHQTEREYIRYHGATSGEHLIAGIHGLAHGIIGGLTSVITSTVEGVKTEGGVSGFISGLGKGLVGTVTKPMAGALDFASETAQAVRDTATLSGPRTQAERVRKPRCCRGPQGLLPRYSESQAEGQEQLFKLTDNIQDEFFIAVENTDSYCVLISSKAVYFLKSGDYVDREAIFLEVKYDHLYHCLISKDHGKVYVQLTKKAGNTSSSVAIPGPSQQKPMVHVKSEVLAVKLSQEINYAKSLYYEQQLMLRLSENQEELELDS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J301</accession>
    <name>A0A452J301_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Myosin light chain kinase, smooth muscle</fullName>
            <ecNumber evidence="5">2.7.11.18</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="21">Telokin</fullName>
        </alternativeName>
    </protein>
    <organism evidence="27 28">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="28" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="27" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="27" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="2">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Myofibril</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the protein kinase superfamily. CAMK Ser/Thr protein kinase family.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.18" evidence="5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039376.1">
        <property type="protein sequence ID" value="ENSGAGP00000034783.1"/>
        <property type="gene ID" value="ENSGAGG00000024612.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030016">
        <property type="term" value="C:myofibril"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004687">
        <property type="term" value="F:myosin light chain kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055013">
        <property type="term" value="P:cardiac muscle cell development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003007">
        <property type="term" value="P:heart morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060298">
        <property type="term" value="P:positive regulation of sarcomere organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045989">
        <property type="term" value="P:positive regulation of striated muscle contraction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00096">
        <property type="entry name" value="Ig"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05762">
        <property type="entry name" value="IgI_8_hMLCK_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd20973">
        <property type="entry name" value="IgI_telokin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000147">
        <property type="entry name" value="Myosin light chain kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000425">
        <property type="entry name" value="Myosin light chain kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000175">
        <property type="entry name" value="Myosin light chain kinase, smooth muscle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000080">
        <property type="entry name" value="Myosin light chain kinase, smooth muscle"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000372">
        <property type="entry name" value="Myosin light chain kinase, smooth muscle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000580">
        <property type="entry name" value="Myosin light chain kinase, smooth muscle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000198">
        <property type="entry name" value="Myosin light chain kinase, smooth muscle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000107">
        <property type="entry name" value="Myosin, light chain kinase a"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:001127">
        <property type="entry name" value="Myosin, light chain kinase a"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000032">
        <property type="entry name" value="palladin isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013098">
        <property type="entry name" value="Ig_I-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47633">
        <property type="entry name" value="IMMUNOGLOBULIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47633:SF1">
        <property type="entry name" value="MYOSIN LIGHT CHAIN KINASE, SMOOTH MUSCLE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16620">
        <property type="entry name" value="23ISL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07679">
        <property type="entry name" value="I-set"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="16 22" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="15" id="KW-0106">Calcium</keyword>
    <keyword evidence="18" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="19" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="20" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="14" id="KW-0418">Kinase</keyword>
    <keyword evidence="17" id="KW-0460">Magnesium</keyword>
    <keyword evidence="11" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13 22" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="9" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="28" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="12" id="KW-0677">Repeat</keyword>
    <keyword evidence="8" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="10" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Ig-like" evidence="25">
        <location>
            <begin position="1"/>
            <end position="72"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="25">
        <location>
            <begin position="102"/>
            <end position="190"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="25">
        <location>
            <begin position="313"/>
            <end position="413"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="25">
        <location>
            <begin position="437"/>
            <end position="525"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="25">
        <location>
            <begin position="535"/>
            <end position="630"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="25">
        <location>
            <begin position="870"/>
            <end position="946"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="25">
        <location>
            <begin position="996"/>
            <end position="1084"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="26">
        <location>
            <begin position="1092"/>
            <end position="1185"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="24">
        <location>
            <begin position="1222"/>
            <end position="1477"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="25">
        <location>
            <begin position="1567"/>
            <end position="1656"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="23">
        <location>
            <begin position="216"/>
            <end position="337"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="23">
        <location>
            <begin position="629"/>
            <end position="652"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="23">
        <location>
            <begin position="772"/>
            <end position="819"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="23">
        <location>
            <begin position="826"/>
            <end position="845"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="23">
        <location>
            <begin position="964"/>
            <end position="983"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="23">
        <location>
            <begin position="1178"/>
            <end position="1205"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="23">
        <location>
            <begin position="225"/>
            <end position="236"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="23">
        <location>
            <begin position="243"/>
            <end position="268"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="23">
        <location>
            <begin position="272"/>
            <end position="283"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="23">
        <location>
            <begin position="312"/>
            <end position="323"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="23">
        <location>
            <begin position="633"/>
            <end position="650"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="23">
        <location>
            <begin position="805"/>
            <end position="819"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="23">
        <location>
            <begin position="831"/>
            <end position="844"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="23">
        <location>
            <begin position="1178"/>
            <end position="1188"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="23">
        <location>
            <begin position="1189"/>
            <end position="1203"/>
        </location>
    </feature>
    <feature type="binding site" evidence="22">
        <location>
            <position position="1251"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001913"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004657"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00006692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012430"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00021842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00030959"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="26">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034783.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="28">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1671" mass="185949" checksum="E6BD46A7DA512128" modified="2019-05-08" version="1">MLCLPPRGYPEPQITWYRNGLPITEGEYYVVDRSIRGIFSLVIKNVQEEDGGKYTCEAANDGGVRQVTVELTVEGKRNIIILGRFSVPSVENRPSIWGESPPKFATKPNRVIVREGQTGKFSCKITGRPQPQITWFKGDIQLQQKDRFNMFEKSGIQFLEIQNVRLADAGTYTCTVTNSSGKASVSAELMVQGTTTRGGRDKFKLATSNGIAKELKSSSTGLTAETKERTAVKKESSSISSSKEAKDNRQRQLREPNTEPAQEPKGEARGSQGLQKSSSTITLQAIKVQPEPKAESTATSAGQAEDRKGATQPLTTTRQTASLIRQAGPRSREAENRVGTXCVSHQTVSGKPKPDVEWFKEGAPVKGKDGIHIYEEDGIHCLCLMKTHLGDSGSYSCTAFNPRGRTSTSWVLSVNKKKSSKKVESAQSVQTNKTFAPVFLKGLTDLKVMDGSQVIMTVEVSANPPPEIIWLHNGKEIQETEDFHFEKRGNEYSLYIQEVFPEDTGKYTCEAWNDLGETQTQATLTVQEPQDGIQPWFISKPRSVTAAPGQNVLISCAIAGDPFPTVHWFKDGQEIMPGAGCEILQNEDIFTLIMRNVQRRQAGQYEIQLRNQVGECSCQVSLMLRESSVSEGEMNREEPACSGEQRDGGDHGALTFGRISGFRKSGGETRPAEEEQEDVRGVLKRRVETREHTEERLRQQEAEQLDFRDILGKKVNTKSFSEEELKEIPAEQMDFRANLQRQVKPKTLSEEERKVHSPQQVDFRAVLAKKGTAKVSLPEKVPPPKPATPDFRSVLGSKKKPPAENGSTNTQAQDTSTNSVALNASTNSEAQHAKPEVKEEEKNDTNCAHGCSMVDGGIIEKKVDRKGMAPSFTEKLQDTHAVDGEKLVLQCRISSDPPAAVTWTLDGKAIKSSKFIVISQEGSLCSLTIDKAFPEDVGQYKCIAENAAGRAECACKVLVDGPSPTKTPKSAEKKTKKPKSSLPPVLATESKWFLSPQIIQFPEDQKVRAGESVELFCKVVGTPPLTCTWMKFRKQIQENEHIKIENAENSSKLTISSTKQEHCGCYTLVVENKMGCRQAQVNLTVVDKPDPPAGKPCASDIQSSSLTLSWYGSSYDGGSAVQSYTVEIWNSVDNKWTDLTTCRSTSYNVKSLLADREYKFRVRAANVYGISEPSQESELVKVGEKQEEPKDEAELSDDEEKETEVDYRTVTINTQQKVAEFYDIEERLGSGKFGQVFKLVEKKTGKVWAGKFFKAYSAKDKENIRQEIGIMNCLHHPKLVQCVDAFEEKANIVMVLEIVSGGELFERIIDEDFELTERECIKYMRQISEGVQYIHKQGIVHLDLKPENIMCVNKTGARIKLIDFGLARRLESTGSLKVLFGTPEFVAPEVINYEPIGYATDMWSIGVICYILVSGLSPFMGDNDNETLANVTSATWDFDDEAFDEISDDAKDFISNLLKKDMKSRLDCTQCLQHPWLQKDTKNMEAKKLSKDRMKKYMARRKWQKTGHAVRAIGRLSSMAMISGLSGRKSSTSSPTSPINTDKVDLNDDAARAFLEAVAEEKPHVKPYFTKTILDIEVVEGSAARFDCKIEGYPDPEVIWYKDDQSIKESRHFQIDYDEDGNCSLTISEVCGDDDAKYTCKAVNSLGEATCTAELIVETMGKEEEEEEEEE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J302</accession>
    <name>A0A452J302_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="23">Polycystin 1, transient receptor potential channel interacting</fullName>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cell projection</location>
            <location evidence="1">Cilium</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the polycystin family.</text>
    </comment>
    <comment type="caution">
        <text evidence="13">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000039276.1">
        <property type="protein sequence ID" value="ENSGAGP00000034695.1"/>
        <property type="gene ID" value="ENSGAGG00000024640.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005929">
        <property type="term" value="C:cilium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005261">
        <property type="term" value="F:monoatomic cation channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006816">
        <property type="term" value="P:calcium ion transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00037">
        <property type="entry name" value="CLECT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00146">
        <property type="entry name" value="PKD"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="CDD" id="cd01752">
        <property type="entry name" value="PLAT_polycystin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.60.20:FF:000012">
        <property type="entry name" value="polycystin-1 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.100.10">
        <property type="entry name" value="Mannose-Binding Protein A, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.60.20">
        <property type="entry name" value="PLAT/LH2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001304">
        <property type="entry name" value="C-type_lectin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016186">
        <property type="entry name" value="C-type_lectin-like/link_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018378">
        <property type="entry name" value="C-type_lectin_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016187">
        <property type="entry name" value="CTDL_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000203">
        <property type="entry name" value="GPS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000434">
        <property type="entry name" value="PC1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022409">
        <property type="entry name" value="PKD/Chitinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002859">
        <property type="entry name" value="PKD/REJ-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013122">
        <property type="entry name" value="PKD1_2_channel"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000601">
        <property type="entry name" value="PKD_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035986">
        <property type="entry name" value="PKD_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001024">
        <property type="entry name" value="PLAT/LH2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036392">
        <property type="entry name" value="PLAT/LH2_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042060">
        <property type="entry name" value="PLAT_polycystin1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046791">
        <property type="entry name" value="Polycystin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014010">
        <property type="entry name" value="REJ_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46730">
        <property type="entry name" value="POLYCYSTIN-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46730:SF2">
        <property type="entry name" value="POLYCYSTIN-1 ISOFORM X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00059">
        <property type="entry name" value="Lectin_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00801">
        <property type="entry name" value="PKD"/>
        <property type="match status" value="12"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08016">
        <property type="entry name" value="PKD_channel"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01477">
        <property type="entry name" value="PLAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20519">
        <property type="entry name" value="Polycystin_dom"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02010">
        <property type="entry name" value="REJ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00500">
        <property type="entry name" value="POLYCYSTIN1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00034">
        <property type="entry name" value="CLECT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00303">
        <property type="entry name" value="GPS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00308">
        <property type="entry name" value="LH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00089">
        <property type="entry name" value="PKD"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56436">
        <property type="entry name" value="C-type lectin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49723">
        <property type="entry name" value="Lipase/lipooxygenase domain (PLAT/LH2 domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49299">
        <property type="entry name" value="PKD domain"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00615">
        <property type="entry name" value="C_TYPE_LECTIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50041">
        <property type="entry name" value="C_TYPE_LECTIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50093">
        <property type="entry name" value="PKD"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50095">
        <property type="entry name" value="PLAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51111">
        <property type="entry name" value="REJ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="12" id="KW-0966">Cell projection</keyword>
    <keyword evidence="8" id="KW-0969">Cilium</keyword>
    <keyword evidence="10" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9 15" id="KW-0472">Membrane</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="16" id="KW-0732">Signal</keyword>
    <keyword evidence="5 15" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 15" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="16">
        <location>
            <begin position="1"/>
            <end position="39"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019171829" description="Polycystin 1, transient receptor potential channel interacting" evidence="16">
        <location>
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    <sequence length="4021" mass="440991" checksum="28DBF5B5D7F50F07" modified="2019-05-08" version="1">MDPSLGDRDTLRSCMAGVMLSQGLQLGWLLLLLLARGTADESSCPQGGRIHFQNLNCYWLSHAKETWQNARRLCREIPGGDLAVVRTQEVQSFLQETFSGAGMVWIGLSSLGSLHWVDGSLVDSFQNWTPFGRPKGKESCVQMHLLDPEGVWSSRPCSAKSSFLCERRAGVTLPSPDVFLTGMPVFSGVYDVRNVSTELLPPLLGPDSIELMLFPGLWFSHAGILVSVDFGVQGTKKPVLVQFQILRPHCNPNQHLVPPGCKFLHTPFACCHPRPLCNTTGWCPSRQQWCPLSEGCLNISSPCSSYAFENVTSSILPIRNPPRYSGTPPFYSPVADFPLLLAPSSANRHVQVLLPREEISVYPDDIIGIQHNAGVSSLLRCLQHSHSPWRQSYISLVKAGWWEGSISSFTNPTWVDHVVCDLRVAFVGKTQSFAATPPLARHSERGAYTYTATVRNAVSSRQMNCTVEVQTKVGGLQIIHPTLVSGKLNVATKKETLIVIKILSGCHAVAHWMAPVGKAGVLFEATCPGSIIALVPACRRDTADTWFSSTLLRMEEPKMETLNVLVSNQISTQKLSVKIQSYNAIEGLRVVPPGPRRMLVDVSQVFSAELTQGSSVSYSWVIDNMDMFAYNGQTYSVKFKRPATYQLKLRAENPVSSEVVEVALTVEAMNPLANPEIVGLPGVVAVDTLQTLVFRVEVDTAMEVTVRWGFGDSSPDVESLLSPPYDSQLSRPDPQGERVQVLARVTHTYTQPGNYSVTAEAINKYDRVEQVANVRAVCPIASVTILVTPASPLVNEVTKFKALPRPSPYGIVYSWDFNDGSPTQEGVNSLVHHRFEKSGVYNVMLWANNNLSEVPSDVAVMVSEGVAGLRVVASGPTELGAATVVNAALASGTDVLWTFNMGDGFVYSNLNLSFISHTFAKEGNYTVMVKAHNAATSTHVSTIAEVYRLQITKILAPVCLSSGEQTPFQAVVTGPSKGVQFCWDFQDGNPPSVGVGDPTVFHSYPVAGSYELNLTVRGTVSTTSRRLAICVEDRIRSVKLMAPTLALALGEPTCFQAEVVPVPDPQHQYRYHWDFGVGEDPVTSGSPEITFTYLEGGSYRVTVTVWNMVSRHNASADILAQRVIGTISILHSGETESFLALGTSYLFMTEVPWDSAATFLWNFGDHSPPQVGQSVSHTYHTVGDITVTVTGENQVSRRNASLDVTVLAPVQLLALSTEWPICEVRQEVAFHASLAAGDQVRYLWAIGQPRSFHQGAATFSHAFPAAGVFVVFVTAENAVSTEKANITIEVQERAQGVQIHSGNVVQGRYLASAEPFMLTAKVTQGSNITFRWVISQGLRRVFTATGQSLMYCYNASGELLVEVRAGNVLGEVSAGLALQVVERACGVKVQAASDSVAVGKPVNLSVSVASGTDLLYSWHIEEYAQPLLTNSSSLSYAYSTLGSKLVFVTVSNVLGSSNGSTELRVEEPVSRVSFAVEGSSPPFFLMSDTSRQLWGSVAAGSDVAWEWQLQGKEKRQLFHGQNISCKFSAAGVYQVDLRAWNDVSEDTVQHPMVVQDPVAGLAVEVDRQDVCTDDRVVFTLHTQRGTNVSFALTIPSLGLSLDVRGGTHHFSFPTAGSHQVLASASNNVSMQLASVVVQVLEKVRGLHIPNCCPAIMESNKELALTAQVQAGENVTFFWIFHLPGQPAYEAAGQQVGYVPPGAGNLTIHVEASNPFCATSLTVTVALQVPIEAASLSSNGTGAFLNQTVAFDVALTGGSDLHSEWKFGDSAATFAGSGVERMFHHYGRAGDFVVEVKIYNEVSFVLVQTTVTVRELGCDSPMVRLVDPPSSVPRSLTSYFEASIDLKGCTAYKALYLWEVFSSPSCEQLAEANQVPLQHADLFTPSLTLPKLSLDIGTHCLRFTASLQHTPLARTLSLLITVIPSKLVPVIRGGSWRSCSAQLDLVLDGSGSYDPDVGTAADHSLEYHWDCENVSSPWCAPPRLQGGASVTIPRTVLCRGTGYRFILTVKKPGKDPVSMAQMVWIKPGQILPVTIECRSCSALSSYEISRSVHVTLAGQCQSCDNRTMYKWTAQSSDGQNLTLNHVTTSTGDSNRDLVIRQGVLRDGVNYSFTLSASQPGGELQGGSSITLTPNNPPRGGICTLTPEHNLYLLETLVSYQCTGWVDEDSSPVQLIYTLIAEQCSPNSRHCQHFCLYRGIKSSFSTFLPAATGSSRTTVNILVELEDSQGAKTLALNRTLTLVMPVLPAGFPTVTSWLKNKSQSELWGVVQQGNPQEVIPYSLALITVLNQNSGSQGNEEELRDRISIRSNVTAALASLNISSVKEVAQLSAALQQCVAFPGELSLESQARTLAAAQRMIEVISSETEEGHETPTSAGNSILGILGSTLSALDTAPHPLLFGTPGLSTALTAFNLTRALMKSLMRSRVLNEETLLLSVSEIRTQGKRANSPNLLCLAPSDRCLFSVPSTVARQLAGSQEVVQVLMDISLNPFPFNYYANSSISTRLALLEFTTPGGVPIPVSNLSGERAIGLRLPTGQQEQRSSPPTLILLPPGESVNFTVRALAGSRAEGVHIHVGITMQEGFDPSQEREPSIHFFGHYTPVPNEFHYTWKEEMIFDSLQGGSSREVTILLSSPHSFNGTLRDYHVNITNRFSQAPVSAAVTVFASLCQYFHFPSMQWSTEGLRPTAATSRKEIVCLTEHLTVFGASLFVPPHSILFLPPSKRARQAPLVVITCCVLFAIYLGMVLIAHKLDDIDITRVGIVPRCGQPGRYKYWVTVKTGWKRGSGTTAHVGISLYGLNKSGSRHLDKGWAFQRNSQDIFQVETDANLGEIWKIRIWHDNTGLDPSWYLQHVIVWDRQTDHMYFFLVDDWLSVENEQNEGLVEKEVLAACPQELRCFSRVFPAQLRLGFSDWHVWLSIWGRSPHSRFTRVQRITCCVLAVYLFLAICVLWYGAIGVEGYSVPLGSQVSVTAESVAVGMVVAVMVFPIQLLFTFIFRKTRSKVVVEDPDPPVQDSQTVEMDVCLEHADLSSSSFLSILGGMESIMDGSSASGESPGSRRPGSTQENLTKVGNTRVVKRRPSCDSIFDVPDLLTSDPLVSQSRSLKRKKALLKLGIESTSNSDGDPLSFSLEDSEDSWRDSRGHRLTRSEEDLLNSIVAETQRSSSSDCVTSDSGRFSPQAEADLVSDVMESPSSAWSDGIPEQGRRGWGFLRKSFSYISALTSPGSVLLPDPEPPSAPVSSFSTRIGVSRRPLEWLFPGWVLPMTYAFIFLLLAGCFTITILYGSSLHDHTALMWLISCIFSFITSFVLLEPLKSCLTHLLFLLVVLTVNYQSCFHHRNIHLLHAAVRQELTAPNDRGLNFSSMRSYTDTWEWIDSVLLSYLYSNSRLTLVGVPRLRQESSAGNIPLDLAPRPEISRSGQVDRDACCDRSLQNSSHNSIGFWIWGHSGVYHTNTGCWRELGNSSAKARRTLADLRASNWINERTRAVFLEFTQYNVDVNLYVVVMLLLELPSVRLAVPSIIILPYQMLRLGTGLDLPLALVISLLIFSVAFLLPELSLLSQEGASCLGQSRCWLQLLLALLSIAVGALHFAHIWLADVQLEHYWGHRQAFTSFYEVALLAQMEVCLSALLLTITMLKIVRQLRFVRRWSAFGKTFQHALRDLVAVMLIFLLLLLVYAQCGYLAFSATVEEFRTFQCAFYALLSALHGKVAFLSVIQEFPTLGTAYVLSFMVSLLCVGSRFLCAVILHSYRTVQAEVYRPAMEPQDYEMIEFFVKRFKLWMGLSKTKEFRHKVKFEGMDSLISRSSGNSKRSRLLSAGTIFRCASSTISSSSEELALLESPVPEQYNVQFYLDRLPSAVNNLLDQFDRVIRVMEDVCQLESGLEQAQKRINDNKKGQKSLQGEKVTLPKQLELPRTYSTFSESALLRLRAHRAKVSNCNATDGHKLVQQPPADGIPHPVAGCTSWLVGPPISTDLCQRCSPAPGVSCKWRPKNEEEQGCLCHEVPQRQIPLKRRAWQTEGAGNI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J303</accession>
    <name>A0A452J303_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">protein-tyrosine-phosphatase</fullName>
            <ecNumber evidence="1">3.1.3.48</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="EC" id="3.1.3.48" evidence="1"/>
    <dbReference type="AlphaFoldDB" id="A0A452J303"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039396.1">
        <property type="protein sequence ID" value="ENSGAGP00000034802.1"/>
        <property type="gene ID" value="ENSGAGG00000024713.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0099560">
        <property type="term" value="P:synaptic membrane adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd14629">
        <property type="entry name" value="R-PTP-F-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000002">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000001">
        <property type="entry name" value="Receptor-type tyrosine-protein phosphatase F isoform A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050348">
        <property type="entry name" value="Protein-Tyr_Phosphatase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000242">
        <property type="entry name" value="PTP_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016130">
        <property type="entry name" value="Tyr_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003595">
        <property type="entry name" value="Tyr_Pase_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19134">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19134:SF203">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE F"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00102">
        <property type="entry name" value="Y_phosphatase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00700">
        <property type="entry name" value="PRTYPHPHTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00194">
        <property type="entry name" value="PTPc"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00404">
        <property type="entry name" value="PTPc_motif"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00383">
        <property type="entry name" value="TYR_PHOSPHATASE_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50055">
        <property type="entry name" value="TYR_PHOSPHATASE_PTP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="2" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="3">
        <location>
            <begin position="73"/>
            <end position="328"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="4">
        <location>
            <begin position="248"/>
            <end position="319"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="3">
        <location>
            <begin position="360"/>
            <end position="619"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="4">
        <location>
            <begin position="537"/>
            <end position="610"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00013064"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS50055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034802.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="628" mass="71568" checksum="AA16F2330FFF6053" modified="2019-05-08" version="1">MFGTGLHILFCFACEKCDYKDAAFGEYGGQAALWWMGHVGYAVYLGGRPGRGNPATPRRSASPADGPACHNSLSSLLQSIDPGQQFTWENSNLEVNKPKNRYANVIAYDHSRVILTSVDGVPGSDYINANYIDGYRKQNAYIATQGPLPETLSDFWRMVWEQRTATIVMMTRLEEKSRVKCDQYWPGRGTETYGLIQVTLLDTVELATYTVRTFALYKNGSSEKRELRQFQFMAWPDHGVPEYPTPILAFLRRVKACNPLDAGPMVVHCSAGVGRTGCFIVIDAMLERMKHEKTVDIYGHVTCMRSQRNYMVQTEDQYIFIHESLLEAATCGNTEVPARNLFAHLQKLSQVPPGESVTAMELEFKLLANSKAHTSRFISANLPCNKFKNRLVNIMPYELTRVCLQPIRGVEGSDYINASFIDGYRQQKAYIATQGPLAETTEDFWRMLWEHNSTIVVMLTKLREMGREKCHQYWPAERSARYQYFVVDPMAEYNMPQYILREFKVTDARDGQSRTIRQFQFTDWPEQGVPKTGEGFIDFIGQVHKTKEQFGQDGPITVHCSAGVGRTGVFITLSIVLERMRYEGVVDMFQTVKTLRTQRPAMVQTEDQYQLCYRAALEYLGSFDHYAT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J304</accession>
    <name>A0A452J304_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Myosin light chain kinase, smooth muscle</fullName>
            <ecNumber evidence="5">2.7.11.18</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="20">Telokin</fullName>
        </alternativeName>
    </protein>
    <organism evidence="26 27">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="27" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="26" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="26" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="2">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the protein kinase superfamily. CAMK Ser/Thr protein kinase family.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.18" evidence="5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039379.1">
        <property type="protein sequence ID" value="ENSGAGP00000034786.1"/>
        <property type="gene ID" value="ENSGAGG00000024612.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004687">
        <property type="term" value="F:myosin light chain kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055013">
        <property type="term" value="P:cardiac muscle cell development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003007">
        <property type="term" value="P:heart morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060298">
        <property type="term" value="P:positive regulation of sarcomere organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045989">
        <property type="term" value="P:positive regulation of striated muscle contraction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00096">
        <property type="entry name" value="Ig"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05762">
        <property type="entry name" value="IgI_8_hMLCK_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd20973">
        <property type="entry name" value="IgI_telokin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000425">
        <property type="entry name" value="Myosin light chain kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000175">
        <property type="entry name" value="Myosin light chain kinase, smooth muscle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000080">
        <property type="entry name" value="Myosin light chain kinase, smooth muscle"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000372">
        <property type="entry name" value="Myosin light chain kinase, smooth muscle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000580">
        <property type="entry name" value="Myosin light chain kinase, smooth muscle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000198">
        <property type="entry name" value="Myosin light chain kinase, smooth muscle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000107">
        <property type="entry name" value="Myosin, light chain kinase a"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:001127">
        <property type="entry name" value="Myosin, light chain kinase a"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013098">
        <property type="entry name" value="Ig_I-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47633">
        <property type="entry name" value="IMMUNOGLOBULIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47633:SF1">
        <property type="entry name" value="MYOSIN LIGHT CHAIN KINASE, SMOOTH MUSCLE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16620">
        <property type="entry name" value="23ISL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07679">
        <property type="entry name" value="I-set"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="16 21" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="15" id="KW-0106">Calcium</keyword>
    <keyword evidence="18" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="19" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="14" id="KW-0418">Kinase</keyword>
    <keyword evidence="17" id="KW-0460">Magnesium</keyword>
    <keyword evidence="11" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13 21" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="9" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="27" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="12" id="KW-0677">Repeat</keyword>
    <keyword evidence="8" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="10" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Ig-like" evidence="24">
        <location>
            <begin position="7"/>
            <end position="81"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="24">
        <location>
            <begin position="281"/>
            <end position="355"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="24">
        <location>
            <begin position="379"/>
            <end position="467"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="24">
        <location>
            <begin position="477"/>
            <end position="572"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="24">
        <location>
            <begin position="812"/>
            <end position="888"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="24">
        <location>
            <begin position="938"/>
            <end position="1026"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="25">
        <location>
            <begin position="1034"/>
            <end position="1127"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="23">
        <location>
            <begin position="1164"/>
            <end position="1419"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="24">
        <location>
            <begin position="1509"/>
            <end position="1598"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="107"/>
            <end position="228"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="571"/>
            <end position="594"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="714"/>
            <end position="761"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="768"/>
            <end position="787"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="906"/>
            <end position="925"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="1120"/>
            <end position="1147"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="22">
        <location>
            <begin position="116"/>
            <end position="127"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="22">
        <location>
            <begin position="134"/>
            <end position="159"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="22">
        <location>
            <begin position="163"/>
            <end position="174"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="22">
        <location>
            <begin position="203"/>
            <end position="214"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="22">
        <location>
            <begin position="575"/>
            <end position="592"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="22">
        <location>
            <begin position="747"/>
            <end position="761"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="22">
        <location>
            <begin position="773"/>
            <end position="786"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="22">
        <location>
            <begin position="1120"/>
            <end position="1130"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="22">
        <location>
            <begin position="1131"/>
            <end position="1145"/>
        </location>
    </feature>
    <feature type="binding site" evidence="21">
        <location>
            <position position="1193"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001913"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00006692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012430"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00021842"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
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    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
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    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
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    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00030959"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034786.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1613" mass="179276" checksum="CB878E684DFAAB86" modified="2019-05-08" version="1">HGRFSLCLLDQRILHYQKSSFHVDTSRPGDIQLQQKDRFNMFEKSGIQFLEIQNVRLADAGTYTCTVTNSSGKASVSAELMVQGTTTRGGRDKFKLATSNGIAKELKSSSTGLTAETKERTAVKKESSSISSSKEAKDNRQRQLREPNTEPAQEPKGEARGSQGLQKSSSTITLQAIKVQPEPKAESTATSAGQAEDRKGATQPLTTTRQTASLIRQAGPRSREAENRVGTXCVSHQTVSGKPKPDVEWFKEGAPVKGKDGIHIYEEDGIHCLCLMKTHLGDSGSYSCTAFNPRGRTSTSWVLSVNNEPIQYAHSIFEDGLAKLTVQDALPEDDGVYTCLAENSTGRASCSAQVTVKEKKSSKKVESAQSVQTNKTFAPVFLKGLTDLKVMDGSQVIMTVEVSANPPPEIIWLHNGKEIQETEDFHFEKRGNEYSLYIQEVFPEDTGKYTCEAWNDLGETQTQATLTVQEPQDGIQPWFISKPRSVTAAPGQNVLISCAIAGDPFPTVHWFKDGQEIMPGAGCEILQNEDIFTLIMRNVQRRQAGQYEIQLRNQVGECSCQVSLMLRESSVSEGEMNREEPACSGEQRDGGDHGALTFGRISGFRKSGGETRPAEEEQEDVRGVLKRRVETREHTEERLRQQEAEQLDFRDILGKKVNTKSFSEEELKEIPAEQMDFRANLQRQVKPKTLSEEERKVHSPQQVDFRAVLAKKGTAKVSLPEKVPPPKPATPDFRSVLGSKKKPPAENGSTNTQAQDTSTNSVALNASTNSEAQHAKPEVKEEEKNDTNCAHGCSMVDGGIIEKKVDRKGMAPSFTEKLQDTHAVDGEKLVLQCRISSDPPAAVTWTLDGKAIKSSKFIVISQEGSLCSLTIDKAFPEDVGQYKCIAENAAGRAECACKVLVDGPSPTKTPKSAEKKTKKPKSSLPPVLATESKWFLSPQIIQFPEDQKVRAGESVELFCKVVGTPPLTCTWMKFRKQIQENEHIKIENAENSSKLTISSTKQEHCGCYTLVVENKMGCRQAQVNLTVVDKPDPPAGKPCASDIQSSSLTLSWYGSSYDGGSAVQSYTVEIWNSVDNKWTDLTTCRSTSYNVKSLLADREYKFRVRAANVYGISEPSQESELVKVGEKQEEPKDEAELSDDEEKETEVDYRTVTINTQQKVAEFYDIEERLGSGKFGQVFKLVEKKTGKVWAGKFFKAYSAKDKENIRQEIGIMNCLHHPKLVQCVDAFEEKANIVMVLEIVSGGELFERIIDEDFELTERECIKYMRQISEGVQYIHKQGIVHLDLKPENIMCVNKTGARIKLIDFGLARRLESTGSLKVLFGTPEFVAPEVINYEPIGYATDMWSIGVICYILVSGLSPFMGDNDNETLANVTSATWDFDDEAFDEISDDAKDFISNLLKKDMKSRLDCTQCLQHPWLQKDTKNMEAKKLSKDRMKKYMARRKWQKTGHAVRAIGRLSSMAMISGLSGRKSSTSSPTSPINTDKVDLNDDAARAFLEAVAEEKPHVKPYFTKTILDIEVVEGSAARFDCKIEGYPDPEVIWYKDDQSIKESRHFQIDYDEDGNCSLTISEVCGDDDAKYTCKAVNSLGEATCTAELIVETMGKEEEEEEEEE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J306</accession>
    <name>A0A452J306_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Perilipin 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Lipid droplet</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the perilipin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J306"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039382.1">
        <property type="protein sequence ID" value="ENSGAGP00000034789.1"/>
        <property type="gene ID" value="ENSGAGG00000024712.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005811">
        <property type="term" value="C:lipid droplet"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006629">
        <property type="term" value="P:lipid metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004279">
        <property type="entry name" value="Perilipin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042998">
        <property type="entry name" value="PLIN1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47138">
        <property type="entry name" value="PERILIPIN-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47138:SF1">
        <property type="entry name" value="PERILIPIN-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03036">
        <property type="entry name" value="Perilipin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109775">
        <property type="entry name" value="Mannose-6-phosphate receptor binding protein 1 (Tip47), C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0551">Lipid droplet</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="194"/>
            <end position="218"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="278"/>
            <end position="311"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="393"/>
            <end position="422"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="202"/>
            <end position="212"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="280"/>
            <end position="303"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004502"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006311"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022677"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034789.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="7">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="517" mass="56969" checksum="974DBCC7799A0813" modified="2019-05-08" version="1">MMAANKSQKMQNGSTEENVLQRVLQLPVVNATCESLQRTYTSTKEVHPLVASVCEAYERGVKGASSLAVWSMEPVVRRLEPQFTAANMLACRGLDHLEEKIPALQYPVEKIASELKGSISTPIQSARSTIGNSIAGTLDRVLGLTAEGYELTKSTVRSTAEYTWSNRVSQMAAAGVDGALGQLEKLVELLLPEEEHEPGNPPSSIISGCSPSPTAPTSRQIGALASTISHRTYQQTARTIQRTRARGQELTTWIPGLVSAPAQQVLYDVQNAAAGWLSKEQSKEPEQEQEKEELKKEETKASEVQRGTPGLLGSMAQSLQTAYLSTISNVKRVPSAAWGTAGELLQLTPHKAAFIAREKLGTLGDALRSVTGSMVETLSHYVPLPRLLAKKEEPAHEKEPPAESQQSQEGSRPVAPPYPEKSQLRGHWRANRAHHPLSFLGLEDPFFLQPSSIQRQAPQRSPTFEPDYPMSRKSAFFPYREGRSTRRVSKNSYRYSPEPVYTRANYSNLYTMAFQKN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJX6</accession>
    <name>A0A452GJX6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">RING-type E3 ubiquitin transferase</fullName>
            <ecNumber evidence="5">2.3.2.27</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1 22">
            <text>S-ubiquitinyl-[E2 ubiquitin-conjugating enzyme]-L-cysteine + [acceptor protein]-L-lysine = [E2 ubiquitin-conjugating enzyme]-L-cysteine + N(6)-ubiquitinyl-[acceptor protein]-L-lysine.</text>
            <dbReference type="EC" id="2.3.2.27"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="3">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the TRIM/RBCC family.</text>
    </comment>
    <dbReference type="EC" id="2.3.2.27" evidence="5"/>
    <dbReference type="AlphaFoldDB" id="A0A452GJX6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002284.1">
        <property type="protein sequence ID" value="ENSGAGP00000002007.1"/>
        <property type="gene ID" value="ENSGAGG00000001264.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061630">
        <property type="term" value="F:ubiquitin protein ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd16598">
        <property type="entry name" value="RING-HC_TRIM26_C-IV"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12888">
        <property type="entry name" value="SPRY_PRY_TRIM7_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.920:FF:000004">
        <property type="entry name" value="Butyrophilin subfamily 1 member A1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.920">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001870">
        <property type="entry name" value="B30.2/SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043136">
        <property type="entry name" value="B30.2/SPRY_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003879">
        <property type="entry name" value="Butyrophylin_SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006574">
        <property type="entry name" value="PRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003877">
        <property type="entry name" value="SPRY_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050143">
        <property type="entry name" value="TRIM/RBCC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000315">
        <property type="entry name" value="Znf_B-box"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020457">
        <property type="entry name" value="Znf_B-box_chordata"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001510">
        <property type="entry name" value="Znf_PARP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001841">
        <property type="entry name" value="Znf_RING"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017907">
        <property type="entry name" value="Znf_RING_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24103">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE TRIM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13765">
        <property type="entry name" value="PRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00622">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00643">
        <property type="entry name" value="zf-B_box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15227">
        <property type="entry name" value="zf-C3HC4_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01406">
        <property type="entry name" value="BBOXZNFINGER"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01407">
        <property type="entry name" value="BUTYPHLNCDUF"/>
    </dbReference>
    <dbReference type="SMART" id="SM00336">
        <property type="entry name" value="BBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00589">
        <property type="entry name" value="PRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00184">
        <property type="entry name" value="RING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00449">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57845">
        <property type="entry name" value="B-box zinc-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57850">
        <property type="entry name" value="RING/U-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50188">
        <property type="entry name" value="B302_SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50119">
        <property type="entry name" value="ZF_BBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50064">
        <property type="entry name" value="ZF_PARP_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00518">
        <property type="entry name" value="ZF_RING_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50089">
        <property type="entry name" value="ZF_RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12 14" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0808">Transferase</keyword>
    <keyword evidence="10" id="KW-0833">Ubl conjugation pathway</keyword>
    <keyword evidence="11" id="KW-0862">Zinc</keyword>
    <keyword evidence="9 13" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="RING-type" evidence="17">
        <location>
            <begin position="16"/>
            <end position="59"/>
        </location>
    </feature>
    <feature type="domain" description="PARP-type" evidence="16">
        <location>
            <begin position="43"/>
            <end position="68"/>
        </location>
    </feature>
    <feature type="domain" description="B box-type" evidence="18">
        <location>
            <begin position="81"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="domain" description="B30.2/SPRY" evidence="19">
        <location>
            <begin position="251"/>
            <end position="440"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="251"/>
            <end position="272"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="14">
        <location>
            <begin position="176"/>
            <end position="203"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00008518"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
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        <source>
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        <source>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002007.1"/>
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    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="440" mass="50659" checksum="6D11E6470959B00C" modified="2019-05-08" version="1">MASSLSGLDMREEATCPICLEYLTDPVSVDCGHNFCRGCVTQYCVEWAGRGSLTCPICRSQILKGNLRPNWQLGNLVEKMKHVAVCTRHKKTLDLFCEEDEEAVCVVCERASQHRSHTVLLIEEAAQKYKEKIQAHLKTLKEEREKQLGLKATGEGKRQEYLVGRQKIVSEFQQLRQFLEEQERLLLAQLEKLDKEILRIQKENVTKLSGQISHLSELIHELEGKCQKPASEFLQVRLSLKHSRSMCRDRRAPESSLRSVRNGPSLPRVHMTLDPDTAHPQLILSEDRKTVRYGRLWNKLPNTTKRFDYERCVLGSEGFTMGRHCWEVEVGERGGWAVGVARESLGRKGEISFKPKGGIWAVEYWEGQYWALTDPLTPLPLSWVPRRIWIYLDYEQGQVTFFNADNESLIFTFPPASFAGERIRPLFVVWETESQLRLCP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJX7</accession>
    <name>A0A452GJX7_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="20">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="15">Homodimer; mediates homophilic interactions to promote cell adhesion.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="14">Synapse</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000002283.1">
        <property type="protein sequence ID" value="ENSGAGP00000002006.1"/>
        <property type="gene ID" value="ENSGAGG00000001306.1"/>
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        <property type="component" value="Unassembled WGS sequence"/>
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        <property type="project" value="UniProtKB-SubCell"/>
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    </dbReference>
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        <property type="term" value="F:cell-cell adhesion mediator activity"/>
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        <property type="project" value="TreeGrafter"/>
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        <property type="term" value="P:camera-type eye photoreceptor cell differentiation"/>
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        <property type="project" value="UniProtKB-ARBA"/>
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    <dbReference type="GO" id="GO:0070593">
        <property type="term" value="P:dendrite self-avoidance"/>
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        <property type="term" value="P:retina layer formation"/>
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        <property type="project" value="UniProtKB-ARBA"/>
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        <property type="term" value="P:synapse assembly"/>
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    </dbReference>
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        <property type="entry name" value="BASIGIN RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="CELL ADHESION MOLECULE DSCAML1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
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        <property type="match status" value="3"/>
    </dbReference>
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    </dbReference>
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    </dbReference>
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    </dbReference>
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        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="8"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="10 17" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="9" id="KW-0770">Synapse</keyword>
    <keyword evidence="3 17" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 17" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="1477"/>
            <end position="1497"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="18">
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            <begin position="980"/>
            <end position="1076"/>
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            <begin position="1268"/>
            <end position="1362"/>
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        <location>
            <begin position="1363"/>
            <end position="1463"/>
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        <location>
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            <begin position="1797"/>
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        <source>
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        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
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        <source>
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    </evidence>
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        <source>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00034103"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00065695"/>
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        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
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        <source>
            <dbReference type="SAM" id="Phobius"/>
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        <source>
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        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1808" mass="198966" checksum="892DBB2BDE32A876" modified="2019-05-08" version="1">TAHFHGLCSGAMWQHCAASEQLPCPATEVAALQWWAQSFCTLWGNNQTFSSLLLVSPENRFFITFYGGLYISDVQKEDALSTYRCITKHKYSGETRQSNGARLSVSDPAESIPTILDGFQSRELKAGQLVELPCIASGYPNPAIRWIKDGRPLPADSRWTKRITGLTISDLRGEDSGTYICEVTNTFGSAEVTGTLTVIDPLHVSLTPKKLKTGIGSTVILSCALSGSPEYALRWYRNTDLVVLDNYISIRGISNETLLITAAQKSHSGAYQCFATRKAQTAQDFSIIALEDGTPRIVSSFSEKVVNPGEQFSLMCAAKGAPPPTVTWALDDEPIQRDNGHRTNQYTMSDGTTVSHMNVTSPQIKDGGVYRCTARNSVGSAEYQARINVRGPPSIRAMKNITAVAGRDTFINCRVIGYPYYSIKWYKDALLLPDNHRQVVFENGTLKLMDVQKGMDEGEYLCSVLIQPQLSISQSVHVTVKVPPLIQPFEFPPASIGQLLYIPCVVSSGDMPIRITWRKDGQVIISGSGVTIESKEFMSSLQISSVSLKHNGNYTCIASNAAATVSRERQLIVRVPPRFVVQPNNQDGIYGKAGVLNCSVDGYPPPKVMWKHAKGSGNPQQYHPVPLTGRIQILPNSSLLIRHVLEEDIGYYLCQASNGVGTDISKSMFLTVKIPAMITSHPNTTIAIKGQMKELNCTARGERPIIIRWEKGDTVIDPDRNMRYAISTKDNGDEVISTLKLKPADRGDSVFFSCHAINSYGEDRGLIQLTVQEPPDPPELEIREVKARSMNLRWTQRFDGNSIITGFDIEYKNKSDSWDFKQSTRNISPTINQANIVDLHPASVYSIRMYSFNKIGRSEPSKELTISTEEAAPDGPPMDVTLQPITSQSIQVTWKAPKKELQNGVIRGYQIGYRENSPGSNGQYSIVEMKATGDSEVYTLDNLKKFAQYGVVVQAFNRAGTGPSSSEINATTLEDVPSQPPENVRALSITSDVAVISWSEPPRSTLNGVLKGYRVIFWSLYMDGEWGEMQNITTTRERVELRGMEKFTNYSVQVLAYTQAGDGVRSSVLYIQTKEDIPGPPARIKAVPSSASSVVVSWLPPTKPNGIIRKYTIFCSSPGSGQPAPSEYETSPEQLFYRIAHLNRGQQYMLWVAAVTSAGRGNISEKVTIEPAGKAPAKIISFGGTVTTPWMKDVRLPCNSVGEPAPAIKWTKDREAGWRLVDGHRLIQANGTLVLRSVKAEDSGYYTCTATNTWGFDTIIINLLVQVPPDQPRLTVSKTSASSITLAWIPGDNGGSSIRGFVLQYSVDNSEEWKDVFISSSERAFQLDSLKCGTWYKVKLAAKNSVGAGRISEIIEAKTHGREPSFSKDQHLFTHINSTHARLNLQGWSSGGCPIMAIVLEYRPKGNWVWQSLKTNNSNEVFLTELREATWYELRMKACNSAGCGNETTQFATLDYDGSTIPPIKSAQGEGDDVKKLFTIACPVILATLGVALLFIIRKKRKEKRLKRLRGELSELSPQTVLLAGDDKATIPVTDTEFSQAVNPQSFCTGVSLHHPALIQNTGPLIDMSDIRPGTNPVSRKSVKSAHSTRNRYSSQWTLTKCQASTPARTLTSDWRTVGSQHGIMVTESDSYSASLSQDTDKGRNSMVSTESASSTYEELARAYEHAKLEEQLQHAKFEITECFISDSSSDQMTTGTNENADSMTSMSTPSEPGICRFTASPPKPQDSDRGKNVAVPIPHRELGKLLGWGEPVQEKWAPPKPKQNQTAGTKHLKGCRAVFKLGDGGKPTAAEEHVDQTWTSLRGESND</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJX8</accession>
    <name>A0A452GJX8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">RING-type E3 ubiquitin transferase</fullName>
            <ecNumber evidence="5">2.3.2.27</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1 21">
            <text>S-ubiquitinyl-[E2 ubiquitin-conjugating enzyme]-L-cysteine + [acceptor protein]-L-lysine = [E2 ubiquitin-conjugating enzyme]-L-cysteine + N(6)-ubiquitinyl-[acceptor protein]-L-lysine.</text>
            <dbReference type="EC" id="2.3.2.27"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="3">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the TRIM/RBCC family.</text>
    </comment>
    <dbReference type="EC" id="2.3.2.27" evidence="5"/>
    <dbReference type="AlphaFoldDB" id="A0A452GJX8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002263.1">
        <property type="protein sequence ID" value="ENSGAGP00000001988.1"/>
        <property type="gene ID" value="ENSGAGG00000001264.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061630">
        <property type="term" value="F:ubiquitin protein ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd16598">
        <property type="entry name" value="RING-HC_TRIM26_C-IV"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12888">
        <property type="entry name" value="SPRY_PRY_TRIM7_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.920:FF:000004">
        <property type="entry name" value="Butyrophilin subfamily 1 member A1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.920">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001870">
        <property type="entry name" value="B30.2/SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043136">
        <property type="entry name" value="B30.2/SPRY_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003879">
        <property type="entry name" value="Butyrophylin_SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006574">
        <property type="entry name" value="PRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003877">
        <property type="entry name" value="SPRY_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050143">
        <property type="entry name" value="TRIM/RBCC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000315">
        <property type="entry name" value="Znf_B-box"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020457">
        <property type="entry name" value="Znf_B-box_chordata"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001510">
        <property type="entry name" value="Znf_PARP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001841">
        <property type="entry name" value="Znf_RING"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017907">
        <property type="entry name" value="Znf_RING_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24103">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE TRIM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13765">
        <property type="entry name" value="PRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00622">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00643">
        <property type="entry name" value="zf-B_box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15227">
        <property type="entry name" value="zf-C3HC4_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01406">
        <property type="entry name" value="BBOXZNFINGER"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01407">
        <property type="entry name" value="BUTYPHLNCDUF"/>
    </dbReference>
    <dbReference type="SMART" id="SM00336">
        <property type="entry name" value="BBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00589">
        <property type="entry name" value="PRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00184">
        <property type="entry name" value="RING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00449">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57845">
        <property type="entry name" value="B-box zinc-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57850">
        <property type="entry name" value="RING/U-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50188">
        <property type="entry name" value="B302_SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50119">
        <property type="entry name" value="ZF_BBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50064">
        <property type="entry name" value="ZF_PARP_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00518">
        <property type="entry name" value="ZF_RING_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50089">
        <property type="entry name" value="ZF_RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12 14" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0808">Transferase</keyword>
    <keyword evidence="10" id="KW-0833">Ubl conjugation pathway</keyword>
    <keyword evidence="11" id="KW-0862">Zinc</keyword>
    <keyword evidence="9 13" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="RING-type" evidence="16">
        <location>
            <begin position="16"/>
            <end position="59"/>
        </location>
    </feature>
    <feature type="domain" description="PARP-type" evidence="15">
        <location>
            <begin position="43"/>
            <end position="68"/>
        </location>
    </feature>
    <feature type="domain" description="B box-type" evidence="17">
        <location>
            <begin position="81"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="domain" description="B30.2/SPRY" evidence="18">
        <location>
            <begin position="259"/>
            <end position="448"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="14">
        <location>
            <begin position="123"/>
            <end position="196"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00008518"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00024"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50064"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50119"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50188"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001988.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="448" mass="51762" checksum="D3F3FD3B3CBF2C28" modified="2019-05-08" version="1">MASSLSGLDMREEATCPICLEYLTDPVSVDCGHNFCRGCVTQYCVEWAGRGSLTCPICRSQILKGNLRPNWQLGNLVEKMKHVAVCTRHKKTLDLFCEEDEEAVCVVCERASQHRSHTVLLIEEAAQKYKEKIQAHLKTLKEEREKQLGLKKQTQTERQKIVSEFQQLRQFLEEQERLLLAQLEKLDKEILRIQKENVTKLSGQISHLSELIHELEGKCQKPASEFLQVRLSLKHSRSMCRDRRAPESCPLESSSQRSQCNCLCALGRKQLRSYRNLHMTLDPDTAHPQLILSEDRKTVRYGRLWNKLPNTTKRFDYERCVLGSEGFTMGRHCWEVEVGERGGWAVGVARESLGRKGEISFKPKGGIWAVEYWEGQYWALTDPLTPLPLSWVPRRIWIYLDYEQGQVTFFNADNESLIFTFPPASFAGERIRPLFVVWETESQLRLCP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJY6</accession>
    <name>A0A452GJY6_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="20">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="15">Homodimer; mediates homophilic interactions to promote cell adhesion.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="14">Synapse</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000002296.1">
        <property type="protein sequence ID" value="ENSGAGP00000002018.1"/>
        <property type="gene ID" value="ENSGAGG00000001306.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030424">
        <property type="term" value="C:axon"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045202">
        <property type="term" value="C:synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098632">
        <property type="term" value="F:cell-cell adhesion mediator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007411">
        <property type="term" value="P:axon guidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060219">
        <property type="term" value="P:camera-type eye photoreceptor cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070593">
        <property type="term" value="P:dendrite self-avoidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007156">
        <property type="term" value="P:homophilic cell adhesion via plasma membrane adhesion molecules"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010842">
        <property type="term" value="P:retina layer formation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007416">
        <property type="term" value="P:synapse assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="CDD" id="cd00096">
        <property type="entry name" value="Ig"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="CDD" id="cd05734">
        <property type="entry name" value="Ig_DSCAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05735">
        <property type="entry name" value="Ig_DSCAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd20958">
        <property type="entry name" value="IgI_5_Dscam"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000333">
        <property type="entry name" value="Down syndrome cell adhesion molecule"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Down syndrome cell adhesion molecule"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Down syndrome cell adhesion molecule"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Down syndrome cell adhesion molecule a"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Down syndrome cell adhesion molecule a"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Down syndrome cell adhesion molecule a"/>
        <property type="match status" value="1"/>
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    <dbReference type="FunFam" id="2.60.40.10:FF:000017">
        <property type="entry name" value="Down syndrome cell adhesion molecule b"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000104">
        <property type="entry name" value="Down syndrome cell adhesion molecule b"/>
        <property type="match status" value="1"/>
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    <dbReference type="FunFam" id="2.60.40.10:FF:000167">
        <property type="entry name" value="Down syndrome cell adhesion molecule b"/>
        <property type="match status" value="1"/>
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    <dbReference type="FunFam" id="2.60.40.10:FF:000172">
        <property type="entry name" value="Down syndrome cell adhesion molecule b"/>
        <property type="match status" value="1"/>
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    <dbReference type="FunFam" id="2.60.40.10:FF:000219">
        <property type="entry name" value="Down syndrome cell adhesion molecule homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Down syndrome cell adhesion molecule homolog"/>
        <property type="match status" value="1"/>
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    <dbReference type="FunFam" id="2.60.40.10:FF:000120">
        <property type="entry name" value="Down syndrome cell adhesion molecule like 1"/>
        <property type="match status" value="1"/>
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    <dbReference type="FunFam" id="2.60.40.10:FF:000477">
        <property type="entry name" value="DS cell adhesion molecule like 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056754">
        <property type="entry name" value="DSCAM/DSCAML_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013098">
        <property type="entry name" value="Ig_I-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013106">
        <property type="entry name" value="Ig_V-set"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10075">
        <property type="entry name" value="BASIGIN RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10075:SF72">
        <property type="entry name" value="CELL ADHESION MOLECULE DSCAML1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25059">
        <property type="entry name" value="FN3_DSCAM-DSCAML_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07679">
        <property type="entry name" value="I-set"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13927">
        <property type="entry name" value="Ig_3"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SMART" id="SM00406">
        <property type="entry name" value="IGv"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="8"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="10 17" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="9" id="KW-0770">Synapse</keyword>
    <keyword evidence="3 17" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 17" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="1439"/>
            <end position="1459"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="18">
        <location>
            <begin position="75"/>
            <end position="159"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="18">
        <location>
            <begin position="163"/>
            <end position="248"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="18">
        <location>
            <begin position="257"/>
            <end position="350"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="18">
        <location>
            <begin position="355"/>
            <end position="435"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="18">
        <location>
            <begin position="445"/>
            <end position="534"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="18">
        <location>
            <begin position="539"/>
            <end position="633"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="18">
        <location>
            <begin position="637"/>
            <end position="734"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="19">
        <location>
            <begin position="736"/>
            <end position="833"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="19">
        <location>
            <begin position="838"/>
            <end position="937"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="19">
        <location>
            <begin position="942"/>
            <end position="1038"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="19">
        <location>
            <begin position="1042"/>
            <end position="1137"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="18">
        <location>
            <begin position="1127"/>
            <end position="1214"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="19">
        <location>
            <begin position="1230"/>
            <end position="1324"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="19">
        <location>
            <begin position="1325"/>
            <end position="1425"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1526"/>
            <end position="1552"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1592"/>
            <end position="1613"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1651"/>
            <end position="1673"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="16">
        <location>
            <begin position="1543"/>
            <end position="1552"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00034103"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00065695"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002018.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1832" mass="201791" checksum="AEE0ED95A6C5FE4C" modified="2019-05-08" version="1">MLLVTCCPLESSPWCSSISLRSFFITFYGGLYISDVQKEDALSTYRCITKHKYSGETRQSNGARLSVSDPAESIPTILDGFQSRELKAGQLVELPCIASGYPNPAIRWIKDGRPLPADSRWTKRITGLTISDLRGEDSGTYICEVTNTFGSAEVTGTLTVIDPLHVSLTPKKLKTGIGSTVILSCALSGSPEYALRWYRNTDLVVLDNYISIRGISNETLLITAAQKSHSGAYQCFATRKAQTAQDFSIIALEDGTPRIVSSFSEKVVNPGEQFSLMCAAKGAPPPTVTWALDDEPIQRDNGHRTNQYTMSDGTTVSHMNVTSPQIKDGGVYRCTARNSVGSAEYQARINVRGPPSIRAMKNITAVAGRDTFINCRVIGYPYYSIKWYKDALLLPDNHRQVVFENGTLKLMDVQKGMDEGEYLCSVLIQPQLSISQSVHVTVKVPPLIQPFEFPPASIGQLLYIPCVVSSGDMPIRITWRKDGQVIISGSGVTIESKEFMSSLQISSVSLKHNGNYTCIASNAAATVSRERQLIVRVPPRFVVQPNNQDGIYGKAGVLNCSVDGYPPPKVMWKHAKGSGNPQQYHPVPLTGRIQILPNSSLLIRHVLEEDIGYYLCQASNGVGTDISKSMFLTVKIPAMITSHPNTTIAIKGQMKELNCTARGERPIIIRWEKGDTVIDPDRNMRYAISTKDNGDEVISTLKLKPADRGDSVFFSCHAINSYGEDRGLIQLTVQEPPDPPELEIREVKARSMNLRWTQRFDGNSIITGFDIEYKNKSDSWDFKQSTRNISPTINQANIVDLHPASVYSIRMYSFNKIGRSEPSKELTISTEEAAPDGPPMDVTLQPITSQSIQVTWKAPKKELQNGVIRGYQIGYRENSPGSNGQYSIVEMKATGDSEVYTLDNLKKFAQYGVVVQAFNRAGTGPSSSEINATTLEDVPSQPPENVRALSITSDVAVISWSEPPRSTLNGVLKGYRVIFWSLYMDGEWGEMQNITTTRERVELRGMEKFTNYSVQVLAYTQAGDGVRSSVLYIQTKEDIPGPPARIKAVPSSASSVVVSWLPPTKPNGIIRKYTIFCSSPGSGQPAPSEYETSPEQLFYRIAHLNRGQQYMLWVAAVTSAGRGNISEKVTIEPAGKAPAKIISFGGTVTTPWMKDVRLPCNSVGEPAPAIKWTKDREAGWRLVDGHRLIQANGTLVLRSVKAEDSGYYTCTATNTWGFDTIIINLLVQVPPDQPRLTVSKTSASSITLAWIPGDNGGSSIRGFVLQYSVDNSEEWKDVFISSSERAFQLDSLKCGTWYKVKLAAKNSVGAGRISEIIEAKTHGREPSFSKDQHLFTHINSTHARLNLQGWSSGGCPIMAIVLEYRPKGNWVWQSLKTNNSNEVFLTELREATWYELRMKACNSAGCGNETTQFATLDYDGSTIPPIKSAQGEGDDVKKLFTIACPVILATLGVALLFIIRKKRKEKRLKRLRGELSELSPQTVLLAGDDKATIPVTDTEFSQAVNPQSFCTGVSLHHPALIQNTGPLIDMSDIRPGTNPVSRKSVKSAHSTRNRYSSQWTLTKCQASTPARTLTSDWRTVGSQHGIMVTESDSYSASLSQDTDKGRNSMVSTESASSTYEELARAYEHAKLEEQLQHAKFEITECFISDSSSDQMTTGTNENADSMTSMSTPSEPGICRFTASPPKPQDSDRGKNVAVPIPHRANKRTDLWSLWGLLLPCPLHAVLSQLGLRPFQVAERQGPNSHRGQGWDYLITITPAFAFTRPESAAIGRRAARIALLRDLQCQEGSKTERDPYCPRPLFCLCSDVPWGMDFLLPTPPLTAAGLGEARMK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJY9</accession>
    <name>A0A452GJY9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">C2H2-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJY9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002176.1">
        <property type="protein sequence ID" value="ENSGAGP00000001916.1"/>
        <property type="gene ID" value="ENSGAGG00000001510.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001997">
        <property type="entry name" value="Uncharacterized protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000056">
        <property type="entry name" value="Zinc finger and SCAN domain-containing 20"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:003000">
        <property type="entry name" value="Zinc finger and SCAN domain-containing 20"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000355">
        <property type="entry name" value="zinc finger and SCAN domain-containing protein 20 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Zinc finger protein 473"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="10"/>
    </dbReference>
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        <property type="entry name" value="Snail/Krueppel_Znf"/>
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        <property type="entry name" value="Znf_C2H2_sf"/>
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        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24388:SF96">
        <property type="entry name" value="GENE, 32687-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24388">
        <property type="entry name" value="ZINC FINGER PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
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        <property type="match status" value="10"/>
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        <property type="match status" value="5"/>
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        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
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    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
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    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="12" id="KW-0732">Signal</keyword>
    <keyword evidence="9" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 11" id="KW-0863">Zinc-finger</keyword>
    <feature type="signal peptide" evidence="12">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019107420" description="C2H2-type domain-containing protein" evidence="12">
        <location>
            <begin position="24"/>
            <end position="447"/>
        </location>
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            <end position="192"/>
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        <location>
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            <end position="248"/>
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            <end position="304"/>
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    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="305"/>
            <end position="332"/>
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        <location>
            <begin position="333"/>
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        <location>
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        <location>
            <begin position="417"/>
            <end position="444"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
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        <source>
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        <source>
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        <source>
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        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
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        <source>
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        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001916.1"/>
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    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="447" mass="50306" checksum="40715A7A3D8EC2BC" modified="2019-05-08" version="1">MRGRCTPCIVFSLFAIPLSFLSTSPGNDSCLDFLSSPADDEIMSENEDGNPQQEGPGQVELQGRFLRRAKGNFSQCWEEREAWSHRHGSERKLGNSQRNKERDPINVLSVGKVSVYSQPLLYIREPTQERNLMNVWTVGKASIGAQILLPIKECTQERDHINRPYKCLDCGKSFNQSSNLTTHQRVHIGERPYKCLECGKSFSVLSTLIVHQRTHTGEKPYECLDCGKSFYRSSNLTTHQRVHTGEGPYKCLECGKSFSAPSVLLIHQRTHTGEKPYKCLECGKSFSRRSGLISHERIHTGKRPYKCLDCGKSFSRRSDLGNHGRIHTGERPYKCLECGKSFSQSSGLINHSRIHTGERPYTCLDCGKSFYGSSNLTAHQRVHIGEGPYKCLECGKSFSAPSVLHIHQRTHTGEKPYKCLKCGKSFSQSSGLISHERIHTGERPAPQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJZ6</accession>
    <name>A0A452GJZ6_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="20">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="15">Homodimer; mediates homophilic interactions to promote cell adhesion.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="14">Synapse</location>
        </subcellularLocation>
    </comment>
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        <property type="gene ID" value="ENSGAGG00000001306.1"/>
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        <property type="term" value="P:camera-type eye photoreceptor cell differentiation"/>
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        <property type="project" value="UniProtKB-ARBA"/>
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        <property type="project" value="UniProtKB-ARBA"/>
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        <property type="project" value="UniProtKB-ARBA"/>
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        <property type="project" value="UniProtKB-ARBA"/>
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        <property type="entry name" value="Ig_V-set"/>
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    </dbReference>
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        <property type="entry name" value="CELL ADHESION MOLECULE DSCAML1"/>
        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
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        <property type="match status" value="6"/>
    </dbReference>
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    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="10 17" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="9" id="KW-0770">Synapse</keyword>
    <keyword evidence="3 17" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 17" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="1402"/>
            <end position="1422"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="18">
        <location>
            <begin position="38"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="18">
        <location>
            <begin position="126"/>
            <end position="211"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="18">
        <location>
            <begin position="220"/>
            <end position="313"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="18">
        <location>
            <begin position="318"/>
            <end position="398"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="18">
        <location>
            <begin position="408"/>
            <end position="497"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="18">
        <location>
            <begin position="502"/>
            <end position="596"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="18">
        <location>
            <begin position="600"/>
            <end position="697"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="19">
        <location>
            <begin position="699"/>
            <end position="796"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="19">
        <location>
            <begin position="801"/>
            <end position="900"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="19">
        <location>
            <begin position="905"/>
            <end position="1001"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="19">
        <location>
            <begin position="1005"/>
            <end position="1100"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="18">
        <location>
            <begin position="1090"/>
            <end position="1177"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="19">
        <location>
            <begin position="1193"/>
            <end position="1287"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="19">
        <location>
            <begin position="1288"/>
            <end position="1388"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1489"/>
            <end position="1515"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1555"/>
            <end position="1577"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1613"/>
            <end position="1708"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="16">
        <location>
            <begin position="1506"/>
            <end position="1515"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1613"/>
            <end position="1636"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1671"/>
            <end position="1681"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1689"/>
            <end position="1708"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00034103"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00065695"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002030.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1735" mass="190464" checksum="6C61EDB4822B8255" modified="2019-05-08" version="1">KEDALSTYRCITKHKYSGETRQSNGARLSVSDPAESIPTILDGFQSRELKAGQLVELPCIASGYPNPAIRWIKDGRPLPADSRWTKRITGLTISDLRGEDSGTYICEVTNTFGSAEVTGTLTVIDPLHVSLTPKKLKTGIGSTVILSCALSGSPEYALRWYRNTDLVVLDNYISIRGISNETLLITAAQKSHSGAYQCFATRKAQTAQDFSIIALEDGTPRIVSSFSEKVVNPGEQFSLMCAAKGAPPPTVTWALDDEPIQRDNGHRTNQYTMSDGTTVSHMNVTSPQIKDGGVYRCTARNSVGSAEYQARINVRGPPSIRAMKNITAVAGRDTFINCRVIGYPYYSIKWYKDALLLPDNHRQVVFENGTLKLMDVQKGMDEGEYLCSVLIQPQLSISQSVHVTVKVPPLIQPFEFPPASIGQLLYIPCVVSSGDMPIRITWRKDGQVIISGSGVTIESKEFMSSLQISSVSLKHNGNYTCIASNAAATVSRERQLIVRVPPRFVVQPNNQDGIYGKAGVLNCSVDGYPPPKVMWKHAKGSGNPQQYHPVPLTGRIQILPNSSLLIRHVLEEDIGYYLCQASNGVGTDISKSMFLTVKIPAMITSHPNTTIAIKGQMKELNCTARGERPIIIRWEKGDTVIDPDRNMRYAISTKDNGDEVISTLKLKPADRGDSVFFSCHAINSYGEDRGLIQLTVQEPPDPPELEIREVKARSMNLRWTQRFDGNSIITGFDIEYKNKSDSWDFKQSTRNISPTINQANIVDLHPASVYSIRMYSFNKIGRSEPSKELTISTEEAAPDGPPMDVTLQPITSQSIQVTWKAPKKELQNGVIRGYQIGYRENSPGSNGQYSIVEMKATGDSEVYTLDNLKKFAQYGVVVQAFNRAGTGPSSSEINATTLEDVPSQPPENVRALSITSDVAVISWSEPPRSTLNGVLKGYRVIFWSLYMDGEWGEMQNITTTRERVELRGMEKFTNYSVQVLAYTQAGDGVRSSVLYIQTKEDIPGPPARIKAVPSSASSVVVSWLPPTKPNGIIRKYTIFCSSPGSGQPAPSEYETSPEQLFYRIAHLNRGQQYMLWVAAVTSAGRGNISEKVTIEPAGKAPAKIISFGGTVTTPWMKDVRLPCNSVGEPAPAIKWTKDREAGWRLVDGHRLIQANGTLVLRSVKAEDSGYYTCTATNTWGFDTIIINLLVQVPPDQPRLTVSKTSASSITLAWIPGDNGGSSIRGFVLQYSVDNSEEWKDVFISSSERAFQLDSLKCGTWYKVKLAAKNSVGAGRISEIIEAKTHGREPSFSKDQHLFTHINSTHARLNLQGWSSGGCPIMAIVLEYRPKGNWVWQSLKTNNSNEVFLTELREATWYELRMKACNSAGCGNETTQFATLDYDGSTIPPIKSAQGEGDDVKKLFTIACPVILATLGVALLFIIRKKRKEKRLKRLRGELSELSPQTVLLAGDDKATIPVTDTEFSQAVNPQSFCTGVSLHHPALIQNTGPLIDMSDIRPGTNPVSRKSVKSAHSTRNRYSSQWTLTKCQASTPARTLTSDWRTVGSQHGIMVTESDSYSASLSQDTDKGRNSMVSTESASSTYEELARAYEHAKLEEQLQHAKFEITECFISDSSSDQMTTGTNENADSMTSMSTPSEPGICRFTASPPKPQDSDRGKNVAVPIPHRANKSKYPSNGPSPQHTHGPAPNGFSSSPAVSTNWSSPWVSGSTAGVAPAGMQVLTYICSYYLFHDSFGS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GK71</accession>
    <name>A0A452GK71_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="11">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Mitochondrion outer membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the Tom40 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GK71"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002408.1">
        <property type="protein sequence ID" value="ENSGAGP00000002118.1"/>
        <property type="gene ID" value="ENSGAGG00000001704.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005741">
        <property type="term" value="C:mitochondrial outer membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008320">
        <property type="term" value="F:protein transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030150">
        <property type="term" value="P:protein import into mitochondrial matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd07305">
        <property type="entry name" value="Porin3_Tom40"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.160.10:FF:000005">
        <property type="entry name" value="mitochondrial import receptor subunit TOM40 homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.160.10">
        <property type="entry name" value="Porin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023614">
        <property type="entry name" value="Porin_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027246">
        <property type="entry name" value="Porin_Euk/Tom40"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037930">
        <property type="entry name" value="Tom40"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10802">
        <property type="entry name" value="MITOCHONDRIAL IMPORT RECEPTOR SUBUNIT TOM40"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01459">
        <property type="entry name" value="Porin_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="6" id="KW-1000">Mitochondrion outer membrane</keyword>
    <keyword evidence="7" id="KW-0653">Protein transport</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4" id="KW-1134">Transmembrane beta strand</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1"/>
            <end position="59"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="18"/>
            <end position="29"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004374"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010510"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022452"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022787"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002118.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="337" mass="36200" checksum="E21B6DFF89E6E2EE" modified="2019-05-08" version="1">MGNVLAASSPTPSPTMPSPGSGLVSVPPGFTMPPVSGLGPPAQERPEEKGDRLPNPGTFEECHRKCKELFPIQMEGVKLTLNKGLSNHFQVNHSVALSTVGDSSYHFGVTYVGTKQLSPTEAFPVLVGDMDNSGSLNAQIIHQLSTRLRSKVAFQTQQSKFVNWQVDGEYRGEDFTAAVTLGNPDILVGSGILVAHYLQSITPCLALGGELVYHRRPGEEGTVMSLAGKYTAPNWIGTLTVGQAGAHATYYHKANDQLQVGVEFEASTRMQDTSVSFGYQLDLPKANLLFKGSVDSNWIVGAVLEKKLLPLPLTLAMGAFLNHRKSKFQCGFGLTIG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GK96</accession>
    <name>A0A452GK96_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">threonine--tRNA ligase</fullName>
            <ecNumber evidence="3">6.1.1.3</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="10">Threonyl-tRNA synthetase</fullName>
        </alternativeName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="12">Catalyzes the attachment of threonine to tRNA(Thr) in a two-step reaction: threonine is first activated by ATP to form Thr-AMP and then transferred to the acceptor end of tRNA(Thr). Also edits incorrectly charged tRNA(Thr) via its editing domain, at the post-transfer stage.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>tRNA(Thr) + L-threonine + ATP = L-threonyl-tRNA(Thr) + AMP + diphosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:24624"/>
            <dbReference type="Rhea" id="RHEA-COMP:9670"/>
            <dbReference type="Rhea" id="RHEA-COMP:9704"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:33019"/>
            <dbReference type="ChEBI" id="CHEBI:57926"/>
            <dbReference type="ChEBI" id="CHEBI:78442"/>
            <dbReference type="ChEBI" id="CHEBI:78534"/>
            <dbReference type="ChEBI" id="CHEBI:456215"/>
            <dbReference type="EC" id="6.1.1.3"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the class-II aminoacyl-tRNA synthetase family.</text>
    </comment>
    <dbReference type="EC" id="6.1.1.3" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452GK96"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002461.1">
        <property type="protein sequence ID" value="ENSGAGP00000002155.1"/>
        <property type="gene ID" value="ENSGAGG00000001743.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004829">
        <property type="term" value="F:threonine-tRNA ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006435">
        <property type="term" value="P:threonyl-tRNA aminoacylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd01667">
        <property type="entry name" value="TGS_ThrRS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00860">
        <property type="entry name" value="ThrRS_anticodon"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00771">
        <property type="entry name" value="ThrRS_core"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.930.10:FF:000009">
        <property type="entry name" value="Threonine--tRNA ligase 2, cytoplasmic"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.800:FF:000003">
        <property type="entry name" value="Threonine--tRNA ligase 2, cytoplasmic"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.20.30:FF:000006">
        <property type="entry name" value="Threonine--tRNA ligase, cytoplasmic"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.980.10:FF:000003">
        <property type="entry name" value="Threonine--tRNA ligase, cytoplasmic"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.20.30">
        <property type="match status" value="1"/>
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        <property type="entry name" value="Anticodon-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.930.10">
        <property type="entry name" value="Bira Bifunctional Protein, Domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.980.10">
        <property type="entry name" value="Threonyl-trna Synthetase, Chain A, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="HAMAP" id="MF_00184">
        <property type="entry name" value="Thr_tRNA_synth"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002314">
        <property type="entry name" value="aa-tRNA-synt_IIb"/>
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    <dbReference type="InterPro" id="IPR006195">
        <property type="entry name" value="aa-tRNA-synth_II"/>
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    <dbReference type="InterPro" id="IPR045864">
        <property type="entry name" value="aa-tRNA-synth_II/BPL/LPL"/>
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    <dbReference type="InterPro" id="IPR004154">
        <property type="entry name" value="Anticodon-bd"/>
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    <dbReference type="InterPro" id="IPR036621">
        <property type="entry name" value="Anticodon-bd_dom_sf"/>
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        <property type="entry name" value="Beta-grasp_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004095">
        <property type="entry name" value="TGS"/>
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    <dbReference type="InterPro" id="IPR012676">
        <property type="entry name" value="TGS-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002320">
        <property type="entry name" value="Thr-tRNA-ligase_IIa"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018163">
        <property type="entry name" value="Thr/Ala-tRNA-synth_IIc_edit"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047246">
        <property type="entry name" value="ThrRS_anticodon"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033728">
        <property type="entry name" value="ThrRS_core"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012947">
        <property type="entry name" value="tRNA_SAD"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00418">
        <property type="entry name" value="thrS"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="THREONINE--TRNA LIGASE, MITOCHONDRIAL"/>
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    </dbReference>
    <dbReference type="PANTHER" id="PTHR11451">
        <property type="entry name" value="THREONINE-TRNA LIGASE"/>
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    </dbReference>
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        <property type="entry name" value="HGTP_anticodon"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="TGS"/>
        <property type="match status" value="1"/>
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    <dbReference type="Pfam" id="PF00587">
        <property type="entry name" value="tRNA-synt_2b"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07973">
        <property type="entry name" value="tRNA_SAD"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="TRNASYNTHTHR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00863">
        <property type="entry name" value="tRNA_SAD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52954">
        <property type="entry name" value="Class II aaRS ABD-related"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55681">
        <property type="entry name" value="Class II aaRS and biotin synthetases"/>
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    </dbReference>
    <dbReference type="SUPFAM" id="SSF81271">
        <property type="entry name" value="TGS-like"/>
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        <property type="entry name" value="ThrRS/AlaRS common domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50862">
        <property type="entry name" value="AA_TRNA_LIGASE_II"/>
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    </dbReference>
    <dbReference type="PROSITE" id="PS51880">
        <property type="entry name" value="TGS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0030">Aminoacyl-tRNA synthetase</keyword>
    <keyword evidence="7" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="5" id="KW-0436">Ligase</keyword>
    <keyword evidence="6" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="8" id="KW-0648">Protein biosynthesis</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="TGS" evidence="14">
        <location>
            <begin position="59"/>
            <end position="121"/>
        </location>
    </feature>
    <feature type="domain" description="Aminoacyl-transfer RNA synthetases class-II family profile" evidence="13">
        <location>
            <begin position="327"/>
            <end position="591"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008226"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00013163"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022598"/>
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    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
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    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
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    </evidence>
    <evidence type="ECO:0000256" key="8">
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            <dbReference type="ARBA" id="ARBA00022917"/>
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    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023146"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00031900"/>
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    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00049515"/>
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    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00058080"/>
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    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50862"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS51880"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002155.1"/>
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    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="701" mass="79569" checksum="6D33B5D0D8FD1598" modified="2019-05-08" version="1">MRLLAPWGKVLAGRGPAWSRRWYQAGALLPHLAERLCLYEKLKAAYDEQCVDRAQKAGGPIRISLPDGQQVEGESLRTTPYHVASQIGQGLAEGAVAARVNGALYDLDRPLESSATLEFLGFDSPEGQAVFWHSSAHILGVAAERFYGALLCHGPSTESGFFYDMYLAGRTVLGSELPALEEACKSIVREKHPFERLEVSREDLLALFKYNKFKLQVIEEKVKSPTATVYRCGGLIDLCRGPHVRHTGKIQALKILKSSSAFWKGDPSLESLQRVYGISFPSPVRLEEWEQLQEAAASRDHRRIGKEQELFFFHELSPGSCFFLPRGAHIYNTLIDFIKSEYRKRGFSEVVTPNIYNAKLWELSGHWQHYSDHMFCFPVENETFALKPMNCPGHCLMFAHRPRSWRELPLRLADFGVLHRNESSGTLTGLTRVRRFQQDDAHIFCTLGQLEGEIGGCLDFLQAVYSVFGFSFRFYLSTRPAGFLGDAHVWEQAEQQLEKSLNDFGQPWELSPGDGAFYGPKIDIQLKDALGRYHQCATIQLDFQMPVRFDLTYISKDGSTSERPVMIHRAVLGSVERILAVLAENYGRKWPLWLSPFQVMVIPVGPDVEAYAHEVRETFHQAGFMADIDADWSATLNRKIRKAQLAQYNFQLVVGRKELSSRTVNVRTRDARQHGEHGLQEVLLKLRELRDARLRNAEELF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GK98</accession>
    <name>A0A452GK98_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Centrosomal protein of 164 kDa</fullName>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Plays a role in microtubule organization and/or maintenance for the formation of primary cilia (PC), a microtubule-based structure that protrudes from the surface of epithelial cells. Plays a critical role in G2/M checkpoint and nuclear divisions. A key player in the DNA damage-activated ATR/ATM signaling cascade since it is required for the proper phosphorylation of H2AX, RPA, CHEK2 and CHEK1. Plays a critical role in chromosome segregation, acting as a mediator required for the maintenance of genomic stability through modulation of MDC1, RPA and CHEK1.</text>
    </comment>
    <comment type="subunit">
        <text evidence="15">Interacts (via N-terminus) with ATRIP. Interacts with ATM, ATR and MDC1. Interacts with XPA (via N-terminus) upon UV irradiation. Interacts with CEP83, CCDC92, TTBK2, DVL3, NPHP3 and weakly with NPHP4. Interacts with DZIP1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Microtubule organizing center</location>
            <location evidence="1">Centrosome</location>
            <location evidence="1">Centriole</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000002414.1">
        <property type="protein sequence ID" value="ENSGAGP00000002122.1"/>
        <property type="gene ID" value="ENSGAGG00000001670.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005814">
        <property type="term" value="C:centriole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097539">
        <property type="term" value="C:ciliary transition fiber"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060271">
        <property type="term" value="P:cilium assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006281">
        <property type="term" value="P:DNA repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00201">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Centrosomal protein of 164 kDa"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1470.10">
        <property type="entry name" value="Photosystem I PsaD, reaction center subunit II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051841">
        <property type="entry name" value="MT-Golgi_org_protein"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001202">
        <property type="entry name" value="WW_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036020">
        <property type="entry name" value="WW_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18902:SF27">
        <property type="entry name" value="CENTROSOMAL PROTEIN OF 164 KDA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18902">
        <property type="entry name" value="NUCLEAR MITOTIC APPARATUS PROTEIN 1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00397">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00456">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51045">
        <property type="entry name" value="WW domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01159">
        <property type="entry name" value="WW_DOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50020">
        <property type="entry name" value="WW_DOMAIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="13" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="5" id="KW-0132">Cell division</keyword>
    <keyword evidence="8" id="KW-0970">Cilium biogenesis/degradation</keyword>
    <keyword evidence="9 17" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="6" id="KW-0227">DNA damage</keyword>
    <keyword evidence="10" id="KW-0234">DNA repair</keyword>
    <keyword evidence="7" id="KW-0498">Mitosis</keyword>
    <keyword evidence="12" id="KW-0539">Nucleus</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="WW" evidence="19">
        <location>
            <begin position="58"/>
            <end position="91"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="105"/>
            <end position="151"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="163"/>
            <end position="215"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="237"/>
            <end position="263"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="315"/>
            <end position="372"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="434"/>
            <end position="467"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="505"/>
            <end position="537"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="572"/>
            <end position="604"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="630"/>
            <end position="660"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="845"/>
            <end position="896"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1041"/>
            <end position="1077"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1093"/>
            <end position="1125"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1681"/>
            <end position="1702"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="17">
        <location>
            <begin position="1139"/>
            <end position="1627"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="18">
        <location>
            <begin position="110"/>
            <end position="126"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="127"/>
            <end position="138"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="169"/>
            <end position="194"/>
        </location>
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    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="241"/>
            <end position="257"/>
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    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="315"/>
            <end position="324"/>
        </location>
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    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="337"/>
            <end position="353"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="441"/>
            <end position="450"/>
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    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="580"/>
            <end position="600"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="633"/>
            <end position="658"/>
        </location>
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    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="1044"/>
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        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="1060"/>
            <end position="1074"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="1102"/>
            <end position="1117"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="1687"/>
            <end position="1702"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004114"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022776"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022794"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00056906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00061715"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00067900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50020"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002122.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1996" mass="224704" checksum="00382851C2101EBF" modified="2019-05-08" version="1">MDMAGPAIRIGDQLILEEDYDETYIPSEQEIQEFAREIGIDPGNEPELMWLAREGIVAPLPPEWKPCQDITGDIYYFNFSNGQSTWDHPCDEHYRELVIQEREKLSAHGSIKKKDKKKKKDKKEKKEKKEREPVKHAAEMQPEPGILPSSSFYRMSSPILSSECASPDLDQHSQMRIEGFLKKGKRKPSERPPETSDSNKQFAGPSPNKLQPLFTAKSCRTHQILADVEKILGRMPSFSRSDVDHQQGQDRSPKIGDKPCLGFSDSESEGLDVVRTKPISYVARKSCSQDLENGRSVLDSCEVLEDKGMLFDEEKLSDNWKQEEMTGGSVCQPDDGLSVRERSTGPGGDKERGPGFSQADENEAFLTPSAPDHTFYSRKCEMILFDASPLEDLSLMEEGDVHCPQRELRAHSRAVESRRKRKLQDQALSQATKFCKSSEVVGKHERESSKQAEFVESEGAGLQAAPQKDLLDSHLAVPEVTADTLPYTRGVEAYAHAGADISASEPVSLSAAQHKDKEKANGLGPEPDNSLTSNSADHFASQILGEIDNFSWDLQSSRESDRPTDLLAAAAKGSLPTHPFTGTLQTPPRSSANGKSQSDCNSEDEKFYQHVQENDQVPSQEHSCVSFNMKLNRSRDAPEKNNEGRQEETGEGEQKTEPVPRGLEVDVFTEGLQVSPGWLYQTAECQESSRLCQQVEGELTQSGKVTEDYCKLKLCEGAGSNESPGSLLAPIQVPPGSLAPLRGVMDGPASAFHGLLSTNVEISGVPSPLLREGTLPLQTLKSTGCTKSLLGSMHEEKASLNLLALDEEEEEESENQSPRGTARLLKNLHMDISALGGGFEYEESLKMSHAEESQTFPLDSDGVRPPAPDKLLNRDADSSLCGLNKEESSGKPLGNELLEKEDTDVEDTVSVAYEQSGTLGERTAEGTHVVSSQGHPVAASPDVRAAETDQMESVAVTVDAVSVSEKIVNGMREEAADLETDSKLDAGKESEASEHVKELQVSDCSDHELLHLTDFGFQSRISEQVLDMDILSAALDSPKVEAQGLREEEKDQSEASLEEEQSKRPKAAESERNQRPFSACKMAEEKCLSLENASQEETAALEQKEGSLDSPKEEEPAAPHQAQLGEEIKQVVSLNWPSAESLNEIARKQEQELEQEKERLLQAKEERVRRFQEELRQEEEEEAQVLHQQKEKSLWSLKEELAKASEEEELRMREEEVEKLSKLRSQISSETKTEKEKIRAEQEAALQKLREEWESLQKVERERLEERKKRVLEMLKVEVEEAQQREAAELEQEKQRVLNELKERLERKRKEAAEVLEKQFAAELQELKSTAEEKHQKVVSSLRKQIAEAQRSEEAQLREELERAEQKVQQKMYQVAEFERELSELMKEKRQEVERDHDRKMERMKEEHKEVLARIQEQYEEEERKQRTKKLEELKGELERLRQLHDGELKALQKQLDEQLSDLRQSHKVKERKLQELEMELEIQATDAKARCAQLNHQEEAMRKKRQQLLDEEKQTELQRNEAALAAQLRLEESRKEHADLVESIRQLRRTLEELQDQKAELESQVDLLQTRSQRLQRRISELEAAVQSKQETLKELVAEDSMASPRREAELHLEDLRETLQAHSSREPASLTSHSNEDSDLQLDNVRHFISAEGVSIKNAKEFLVRQTRSMRKRHTALKAAKQQWRHNMQQAQDAVQDPDSSQLLEDVRRNLEEESAQLDKMKSAMQKGQVLLKKKEEKLSQLESSLLEELSDEDTLKGAACKKVVTFDLSDSEETHSLSSTDVPQHKFDLKPDVQFPQLDKIQHLTSTLQHITAELNGVLSTLGSLNQRQPPLFAPVQVPMTALPSDGVPLSMYTSLARAQAAGPIMPPAGIPLVNQWAWSTGLNSSPSSTAGQSVDDILAEKWHKYFPGGGSPLWSPTMLATLMTALFFMVRALGQLRTSLFHEEPLPYLVYQPLPWQVQYVTQSRRASRDCGVRVTLLVSFCSHGEGPTAPS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GKA1</accession>
    <name>A0A452GKA1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4 7">Methanethiol oxidase</fullName>
            <shortName evidence="7">MTO</shortName>
            <ecNumber evidence="3 7">1.8.3.4</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="7">Selenium-binding protein 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="7">Catalyzes the oxidation of methanethiol, an organosulfur compound known to be produced in substantial amounts by gut bacteria. Selenium-binding protein which may be involved in the sensing of reactive xenobiotics in the cytoplasm. May be involved in intra-Golgi protein transport.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="6 7">
            <text>methanethiol + O2 + H2O = hydrogen sulfide + formaldehyde + H2O2 + H(+)</text>
            <dbReference type="Rhea" id="RHEA:11812"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:15379"/>
            <dbReference type="ChEBI" id="CHEBI:16007"/>
            <dbReference type="ChEBI" id="CHEBI:16240"/>
            <dbReference type="ChEBI" id="CHEBI:16842"/>
            <dbReference type="ChEBI" id="CHEBI:29919"/>
            <dbReference type="EC" id="1.8.3.4"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="1 7">Organosulfur degradation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="7">Nucleus</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="7">Cytoplasm</location>
            <location evidence="7">Cytosol</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="7">Membrane</location>
            <topology evidence="7">Peripheral membrane protein</topology>
        </subcellularLocation>
        <text evidence="7">May associate with Golgi membrane. May associate with the membrane of autophagosomes.</text>
    </comment>
    <comment type="similarity">
        <text evidence="2 7">Belongs to the selenium-binding protein family.</text>
    </comment>
    <dbReference type="EC" id="1.8.3.4" evidence="3 7"/>
    <dbReference type="AlphaFoldDB" id="A0A452GKA1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002467.1">
        <property type="protein sequence ID" value="ENSGAGP00000002161.1"/>
        <property type="gene ID" value="ENSGAGG00000001587.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0018549">
        <property type="term" value="F:methanethiol oxidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008430">
        <property type="term" value="F:selenium binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008826">
        <property type="entry name" value="Se-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23300">
        <property type="entry name" value="METHANETHIOL OXIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23300:SF0">
        <property type="entry name" value="METHANETHIOL OXIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05694">
        <property type="entry name" value="SBP56"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF75011">
        <property type="entry name" value="3-carboxy-cis,cis-mucoante lactonizing enzyme"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0007">Acetylation</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="7" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="7" id="KW-0653">Protein transport</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 7" id="KW-0711">Selenium</keyword>
    <keyword evidence="7" id="KW-0813">Transport</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00005177"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005606"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012510"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00015601"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023266"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00047539"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU369071"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002161.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="467" mass="52281" checksum="C12CCB5C4512EC0E" modified="2019-05-08" version="1">SSAKCGACGPGYKTPLDAMKGPREEIVYLPCIYRNTGIEKPDYLATVDVDPKSSHYCQVIHRLPMPNLRDELHHSGWNTCSSSFGDVTKKRNRLLLPSLISSRIYVVDVGTDMRAPRIHKMIEPVDVFWKCGLANPHTTHCLGNGEIMISSMGDPSGNGKGGFILLDGETFEVKGNWEKGSNLPALGYDFWYQPRHNVLMSTEWGIPKFDPADLKTGSYGRHINVWDWTTHVFTQSIDLGEDSIPLEIRFLHNPDAAEGYVGCALSSAIHRFYKTGNGTWAAEKVVQVPSKKVNGWLFPDMPGFITDILISLDDRFLYFSNWLHGDIRQYDISNTRKPRLVGQVFLGGKIPRDGPIVVVEDQELKCQPEPFVIKGKRVQGGPQMIQLSLDGKRLYVSTSLYSAWDKQFYPDLVKEGSVMLQIDVDTEKGGLKVNKNFLVDFGKEPDGPVLAHEIRYPGGDCSSDIWI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GKA3</accession>
    <name>A0A452GKA3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Carbonic anhydrase</fullName>
            <ecNumber evidence="4">4.2.1.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="4">Reversible hydration of carbon dioxide.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="4">
            <text>hydrogencarbonate + H(+) = CO2 + H2O</text>
            <dbReference type="Rhea" id="RHEA:10748"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:17544"/>
            <dbReference type="EC" id="4.2.1.1"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="4">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <comment type="similarity">
        <text evidence="1 4">Belongs to the alpha-carbonic anhydrase family.</text>
    </comment>
    <dbReference type="EC" id="4.2.1.1" evidence="4"/>
    <dbReference type="AlphaFoldDB" id="A0A452GKA3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002423.1">
        <property type="protein sequence ID" value="ENSGAGP00000002128.1"/>
        <property type="gene ID" value="ENSGAGG00000001445.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004089">
        <property type="term" value="F:carbonate dehydratase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.200.10">
        <property type="entry name" value="Alpha carbonic anhydrase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001148">
        <property type="entry name" value="CA_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036398">
        <property type="entry name" value="CA_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023561">
        <property type="entry name" value="Carbonic_anhydrase_a-class"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018338">
        <property type="entry name" value="Carbonic_anhydrase_a-class_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18952">
        <property type="entry name" value="CARBONIC ANHYDRASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18952:SF25">
        <property type="entry name" value="CARBONIC ANHYDRASE 5B, MITOCHONDRIAL-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00194">
        <property type="entry name" value="Carb_anhydrase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01057">
        <property type="entry name" value="Carb_anhydrase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51069">
        <property type="entry name" value="Carbonic anhydrase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00162">
        <property type="entry name" value="ALPHA_CA_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51144">
        <property type="entry name" value="ALPHA_CA_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0456">Lyase</keyword>
    <keyword evidence="2 4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 4" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="Alpha-carbonic anhydrase" evidence="5">
        <location>
            <begin position="1"/>
            <end position="231"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00010718"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="RuleBase" id="RU367011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS51144"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002128.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="252" mass="29124" checksum="007048C6F6081E7F" modified="2019-05-08" version="1">PINIQWRDSVYDPHLKPLKIHYDPTTCLYIWNNGYSFQVEFDDSADKSVIMGGPLENHYRLKQFHFHWGAINEWGSEHMVDSKVYPAELHLVHWNPVTSTSFEEAIMENNGLVVIGVFLKLGAHHAGLQKLVDALPAVKHKDTIVEFDVFDPSCLMPSCPDYWTYDGSLTTPPLTESVTWIIKKTPIEVDQNQLEEFRMLLFTSAGEEEKRMVDNFRPLQPLMNRTVRSSFRNHHLLNIETRSSELAHQHSP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GKD4</accession>
    <name>A0A452GKD4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">EGF-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cell projection</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the tenascin family. Teneurin subfamily.</text>
    </comment>
    <comment type="caution">
        <text evidence="13">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000002356.1">
        <property type="protein sequence ID" value="ENSGAGP00000002069.1"/>
        <property type="gene ID" value="ENSGAGG00000000665.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043005">
        <property type="term" value="C:neuron projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050839">
        <property type="term" value="F:cell adhesion molecule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046982">
        <property type="term" value="F:protein heterodimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042803">
        <property type="term" value="F:protein homodimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007157">
        <property type="term" value="P:heterophilic cell-cell adhesion via plasma membrane cell adhesion molecules"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048666">
        <property type="term" value="P:neuron development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00054">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.180.10.10">
        <property type="entry name" value="RHS repeat-associated core"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.120.10.30">
        <property type="entry name" value="TolB, C-terminal domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011042">
        <property type="entry name" value="6-blade_b-propeller_TolB-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008969">
        <property type="entry name" value="CarboxyPept-like_regulatory"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057627">
        <property type="entry name" value="FN-plug_TEN1-4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022385">
        <property type="entry name" value="Rhs_assc_core"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056823">
        <property type="entry name" value="TEN-like_YD-shell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056822">
        <property type="entry name" value="TEN_NHL"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056820">
        <property type="entry name" value="TEN_TTR-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051216">
        <property type="entry name" value="Teneurin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028916">
        <property type="entry name" value="Tox-GHH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006530">
        <property type="entry name" value="YD"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR03696">
        <property type="entry name" value="Rhs_assc_core"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01643">
        <property type="entry name" value="YD_repeat_2x"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11219">
        <property type="entry name" value="TENEURIN AND N-ACETYLGLUCOSAMINE-1-PHOSPHODIESTER ALPHA-N-ACETYLGLUCOSAMINIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11219:SF9">
        <property type="entry name" value="TENEURIN-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25024">
        <property type="entry name" value="EGF_TEN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24329">
        <property type="entry name" value="FN-plug_TEN1-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25021">
        <property type="entry name" value="TEN_NHL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25023">
        <property type="entry name" value="TEN_YD-shell"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23538">
        <property type="entry name" value="Teneurin_ABD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15636">
        <property type="entry name" value="Tox-GHH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25020">
        <property type="entry name" value="TTR_TEN1-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF63829">
        <property type="entry name" value="Calcium-dependent phosphotriesterase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49464">
        <property type="entry name" value="Carboxypeptidase regulatory domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01186">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="12" id="KW-0966">Cell projection</keyword>
    <keyword evidence="11 13" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6 13" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="10" id="KW-0472">Membrane</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="14" id="KW-0732">Signal</keyword>
    <keyword evidence="7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="14">
        <location>
            <begin position="1"/>
            <end position="18"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5046375804" description="EGF-like domain-containing protein" evidence="14">
        <location>
            <begin position="19"/>
            <end position="2029"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="15">
        <location>
            <begin position="68"/>
            <end position="103"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="13">
        <location>
            <begin position="72"/>
            <end position="82"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="13">
        <location>
            <begin position="93"/>
            <end position="102"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004167"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004316"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00009385"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002069.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2029" mass="227918" checksum="58CE33C9F413FB60" modified="2025-10-08" version="2">MTLLCSAVICAADCGGHGICVGGTCRCEEGWMGAACDQRACHPRCNEHGTCRDGKCECSPGWNGEHCTIEGCPGLCNGNGRCTLDMNGWHCVCQLGWRGAGCDTSMETACGDGKDNDGDGLVDCMDPDCCLQPLCHVNPLCLGSPDPLDIIQETQTPTSQQNLHSFYDRIKFLIGKDSTHVIPGENPFDGGHACVIRGQVITSDGTPLVGVNISFVNNPLFGYTISRQDGSFDLVTNGGISIILHFERAPFITQEHTLWLPWDRFFVMETIVMRHEENEIPSCDLSNFARPNPVVSPSPLTAFASSCSEKGPIVPEIQALQEEINIAGSKIKLSYLSSRTAGYKSVLRISMTHPTIPFNLMKVHLMVAVEGRLFRKWFAAAPDLSYYFIWDKTDVYSQKVYGLSEAFVSVGYEYESCPDLILWEKRTAVLQGYEIDASKLGGWSLDQHHALNIQSGILHKGNGENQFVSQQPPVIGSIMGNGRRRSISCPSCNGLADGNKLLAPVALTCGSDGSLYVGDFNYIRRIFPSGNVTNILELRYTRHHSPAHKYYLTSDPITGSIYLSDTNSRRIYKIKSTSVVKDVVKNSEVVAGTGDQCLPFDETRCGDGGKGTEATLTNPRGKTQSLLTSILKWASRKVPNQQFWNTTEQVYKLPQGNPLMCLRVRLEWPTDLAVSPMDNSLYVLDNNVVLQISENHQVRIVAGRPMHCQVPGIDHFLLSKVAIHATLESATALAVSHNGVLYIAETDEKKINRIRQVTTNGEISLVAGAPSSCDCKNDANCDCFSGDDGYAKDARLNAPSSLAVCADGELYVADLGNIRIRFIRKNKPYLNTQNLYELSSPIDQELYLFDTSGKHLYTQSLPTGDYLYNFTYTGDGDITLITDNNGNLVNVRRDSTGMPLWLVVPDGQVYWVTIGTNSALKSVTAQGHEVGMMTYHGNSGLLATKTNENGWTTFYEYDSFGRLTNVTFPTGQVSSFRSDTDRSVHVQVETSSKDDVTITTNLSASGAFYTLLQDQVRNSYYIGADGSLRLMLANGMEVALQTEPHLLAGTVNPTVGKRNVTLPIDNGLNLVEWRQRKEQARGQVTVFGRRLRVHNRNLLSLDFDRVTRTEKIYDDHRKFTLRILYDQAGRPSLWSPSSRLNGVNVTYSPGGHIAGIQRGTMSERMEYDQAGRIISRIFADGKSWSYTYLEKSMVLLLHSQRQYIFEFDKNDRLSSVTMPNVARQTLETIRSIGYYRNIYRPPEGNASVIQDFTEEGQLLHTFYLGTGRRVIYKYGKLSKLAEMLYDTTKIGFTYDETAGMLKTINLQNEGFTCTIRYRQIGPLIDRQIFRFTEEGMVNARFDYNYDNSFRVTSMQAVINETPLPIDLYRYDDVSGKTEQFGKFGVIYYDINQIITTAVMTHTKHFDAYGRMKEVQYEIFRSLMYWMTVQYDNMGRVVKKELKVGPYANTTRYSYEYDADGQLQTVSINDKPLWRYSYDLNGNLHLLSPGNSARLTPLRYDLRDRITRLGDVQYKMDEDGFLRQRGNDIFEYNSAGLLIKAYNKVSGWSVKYRYDGLGRRVSSKTAHSHHLQFFYADLTNPTKVTHLYNHSSSEITSLYYDLQGHLFAMELSSGDEFYIACDNIGTPLAVFSGTGLMIKQILYTAYGEIYMDTNPNFQVIIGYHGGLYDPLTKLIHMGRRDYDVLAGRWTSPDHDMWKHLSSNNIMPFNLYMFKNNNPISNAQDIKCYMTDVNSWLLTFGFQLHNVIPGYPKPDTDAMEPSYELIHTQMKTQEWDNSKSILGVQCEVQKQLKAFVTLERFEQIYSSSIAGCRTVKENKNFASGGSVFGKGVKFAMKDGRVATDIISLANEDGRRIAAILNNAHYLENLHFTIEGVDTHYFVKQGPSESDLSILGLSGGRRTLENGVNVTVSQINTVLSGRTRRYTDIQLQYGALCLNTRYGTTLDEEKARVLELARQRAIAQAWAREQQRLRDGEEGIRSWTEGEKQQVLNTGRVQGYDGYFVISVEQYPELSDSANNIHFMRQSEMGRR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GKD6</accession>
    <name>A0A452GKD6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Serine/threonine-protein kinase TOR</fullName>
            <ecNumber evidence="10">2.7.11.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="9 16">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="8 16">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="similarity">
        <text evidence="1 10">Belongs to the PI3/PI4-kinase family.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="10"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002374.1">
        <property type="protein sequence ID" value="ENSGAGP00000002087.1"/>
        <property type="gene ID" value="ENSGAGG00000000429.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031931">
        <property type="term" value="C:TORC1 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031932">
        <property type="term" value="C:TORC2 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0106310">
        <property type="term" value="F:protein serine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="RHEA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0044877">
        <property type="term" value="F:protein-containing complex binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016242">
        <property type="term" value="P:negative regulation of macroautophagy"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009891">
        <property type="term" value="P:positive regulation of biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051094">
        <property type="term" value="P:positive regulation of developmental process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051240">
        <property type="term" value="P:positive regulation of multicellular organismal process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008361">
        <property type="term" value="P:regulation of cell size"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051896">
        <property type="term" value="P:regulation of phosphatidylinositol 3-kinase/protein kinase B signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0080090">
        <property type="term" value="P:regulation of primary metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0038202">
        <property type="term" value="P:TORC1 signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05169">
        <property type="entry name" value="PIKKc_TOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.1070.11:FF:000074">
        <property type="entry name" value="DJ576K7.1 (FK506 binding protein 12-rapamycin associated protein 1)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000060">
        <property type="entry name" value="Serine/threonine-protein kinase mTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000094">
        <property type="entry name" value="Serine/threonine-protein kinase mTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000532">
        <property type="entry name" value="Serine/threonine-protein kinase mTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.150:FF:000001">
        <property type="entry name" value="Serine/threonine-protein kinase TOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000083">
        <property type="entry name" value="Serine/threonine-protein kinase TOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1010.10:FF:000004">
        <property type="entry name" value="Serine/threonine-protein kinase TOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.150">
        <property type="entry name" value="FKBP12-rapamycin binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1070.11">
        <property type="entry name" value="Phosphatidylinositol 3-/4-kinase, catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1010.10">
        <property type="entry name" value="Phosphatidylinositol 3-kinase Catalytic Subunit, Chain A, domain 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.10">
        <property type="entry name" value="Tetratricopeptide repeat domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050517">
        <property type="entry name" value="DDR_Repair_Kinase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009076">
        <property type="entry name" value="FRB_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036738">
        <property type="entry name" value="FRB_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057564">
        <property type="entry name" value="HEAT_ATR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024585">
        <property type="entry name" value="mTOR_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000403">
        <property type="entry name" value="PI3/4_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036940">
        <property type="entry name" value="PI3/4_kinase_cat_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018936">
        <property type="entry name" value="PI3/4_kinase_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003151">
        <property type="entry name" value="PIK-rel_kinase_FAT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014009">
        <property type="entry name" value="PIK_FAT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026683">
        <property type="entry name" value="TOR_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011990">
        <property type="entry name" value="TPR-like_helical_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11139">
        <property type="entry name" value="ATAXIA TELANGIECTASIA MUTATED ATM -RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11139:SF9">
        <property type="entry name" value="SERINE_THREONINE-PROTEIN KINASE MTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02259">
        <property type="entry name" value="FAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08771">
        <property type="entry name" value="FRB_dom"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23593">
        <property type="entry name" value="HEAT_ATR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11865">
        <property type="entry name" value="mTOR_dom"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00454">
        <property type="entry name" value="PI3_PI4_kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01346">
        <property type="entry name" value="DUF3385"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00146">
        <property type="entry name" value="PI3Kc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01345">
        <property type="entry name" value="Rapamycin_bind"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47212">
        <property type="entry name" value="FKBP12-rapamycin-binding domain of FKBP-rapamycin-associated protein (FRAP)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51189">
        <property type="entry name" value="FAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00915">
        <property type="entry name" value="PI3_4_KINASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00916">
        <property type="entry name" value="PI3_4_KINASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50290">
        <property type="entry name" value="PI3_4_KINASE_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7 10" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="5 10" id="KW-0418">Kinase</keyword>
    <keyword evidence="4 10" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="10" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="6" id="KW-0802">TPR repeat</keyword>
    <keyword evidence="2 10" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="FAT" evidence="13">
        <location>
            <begin position="1359"/>
            <end position="1945"/>
        </location>
    </feature>
    <feature type="domain" description="PI3K/PI4K catalytic" evidence="12">
        <location>
            <begin position="2119"/>
            <end position="2372"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1789"/>
            <end position="1829"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="1800"/>
            <end position="1823"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00011031"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022803"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00047899"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00048679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="RuleBase" id="RU364109"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50290"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS51189"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002087.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2372" mass="269926" checksum="D566B9BD73EE8C88" modified="2019-05-08" version="1">MSSTVSILQQFASGLKSRNEEIRAKAAKDLQHYVTMELREMSQEESTRFYDQLNHHIFELVSSSDANERKGGILAIASLIGVEGGNATRIGRFANYLRNLLPSNDPVVMEMASKAIGRLALAGDTFTAEYVEFEVKRALEWLGADRNEGRRHAAVLVLRELAISVPTFFFQQVQPFFDNIFVAVWDPKQAIREGAVSALRACLILTTQREPKEMQKPQWYRHTYEEAEKGFDETLAKEKGMNRDDRIHGALLILNELVRISSMEGERLREEMEEITQQQMVHDKYCKDLLGFGTKPRHITPFTSFQSVQPPQSNALVGLLGYSPHQGIMGFGASPIPAKSALVESRCCRDLMEEKFDQVCLWVLKCRTSKNSLIQMTILNLLPRLAAFRPSAFTADQYLPDTMNHILSCVKKEKERTAAFQALGLLSVAVRSEFQAYLPKVLEIIKAALPPKDFAHKRQKSMQVDATVFTCISMLARAMGPCIQQDIKELLEPMLAVGLSPALTAVLYDLSRQIPQLKKDIQDGLLKMLSLVLMHKPLRHPGMPKGLAHQLASPSLTNIPEASDVGSITLALRTLGSFEFEGHSLTQFVRHCADHFLNSEHKEIRMEAARTCSRLLTPSIHLISGHAHVVSQTAVQVVADVLSKLLVVGITDPDPDIRYCVLASLDERFDAHLAQAENLQALFVALNDQVFEIRELAICTVGRLSSMNPAFVMPFLRKMLIQILTELEHSGVGRIKEQSARMLGHLVSNAPRLIRPYMEPILKALIVKLKDPDPDPNPGVINNVLATIGELAQVSGLEMRKWVDELFIIIMDMLQDSSLLAKRQVALWTLGQLVASTGYVVEPYRKYPTLLDVLLNFLKTEQNQGTRREAIRVLGLLGALDPYKHKVNIGMIDQSRDASAVSLSESKSSQDSCKQRETGERMCTLHFSMVALMRIFRDQSLSHHHTMVVQAITFIFKSLGLKCVQFLPQVMPTFLNVIRVCDGAIREFLFQQLGMLVSFVRSHIRPYMDEIVTLMRDFWVMNNSIQSTIILLIEQIVVALGGEFKLYLPQLIPHMLRVFMHDSSPGRNVSVKLLNAIQLFGANLDDYLHLLLPPIVKLFDAPDVPLVARKAALETVDRLTESLDFTDYASRIIHPIVRTLDLSPELRPTAMDTLSSLVFQLGKKYQIFIPMVNKVLMRHRINHQRYDVLICRIVKGYTLADEEEDPLIYQHRMLRSNQGEMLTSGPVETGPMKKLHVSTINLQKAWGAARRVSKDDWLEWLRRLSLELLKDSSSPSLRSCWALAQAYNPMARDLFNAAFVSCWSELNEDQQDELIRSIELALTSQDIAEVTQTLLNLAEFMEHSDKGPLPLRDDNGIVLLGERAAKCRAYAKALHYKELEFQKGPTPAILESLISINNKLQQPEAAAGVLEYAMKHFGELEIQATWYEKLHEWEDALVAYDKKMDTNKDDPELMLGRMRCLEALGEWGQLHQQCCEKWTLVNDETQAKMARMAAAAAWGLGQWDSMEEYTCMIPRDTHDGAFYRAVLALHQDLFSLAQQCIDKARDLLDAELTAMAGESYSRAYGAMVSCQMLSELEEVIQYKLVPERREIIRQIWWERLQGCQRIVEDWQRILMVRSLVVNPHEDMRTWLKYASLCGKSGRLALAHKTLVLLLGVDPSRQLDHPLPTVHPQVTYAYMKHMWKSARKIDAFQHMQHFVQTMQEQAQHAIATEDQQHKQELHKLMARCFLKLGEWQLNLQGINESTIPKVLQYYSASTEHDGNWYKAWHAWAVMNFEAVLHYKHQNQARDEKKKLRHASGASITSANTEGSNSDSEAESTENSPIPSPVQKKVTEDLSRTLLLYTVPAVQGFFRSISLSRGNNLQDTLRVLTLWFDYGHWPDVNEALVEGVKAIQIDTWLQVIPQLIARIDTPRPLVGRLIHQLLTDIGRYHPQALIYPLTVASKSTTTARHNAANKILKNMCEHSNTLVQQAMMVSEELIRVAILWHEMWHEGLEEASRLYFGERNVKGMFEVLEPLHAMMERGPQTLKETSFNQAYGRDLMEAQEWCRKYMKSGNVKDLTQAWDLYYHVFRRISKQLPQLTSLELQYVSPKLLMCRDLELAVPGTYDPNQPIIRIQSIAPSLQVITSKQRPRKLTLMGSNGHEFVFLLKGHEDLRQDERVMQLFGLVNTLLANDPTSLRKNLSIQRYAVIPLSTNSGLIGWVPHCDTLHALIRDYREKKKILLNIEHRIMLRMAPDYDHLTLMQKVEVFEHAVNNTAGDDLAKLLWLKSPSSEVWFDRRTNYTRSLAVMSMVGYILGLGDRHPSNLMLDRLSGKILHIDFGDCFEVAMTREKFPEKIPFRLTRMLTNAMEQIQKAINDHEQGQTPIQLVSQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GKD7</accession>
    <name>A0A452GKD7_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="15">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GKD7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002477.1">
        <property type="protein sequence ID" value="ENSGAGP00000002170.1"/>
        <property type="gene ID" value="ENSGAGG00000001747.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000398">
        <property type="term" value="P:mRNA splicing, via spliceosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd16167">
        <property type="entry name" value="OCRE_RBM10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12753">
        <property type="entry name" value="RRM1_RBM10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12754">
        <property type="entry name" value="RRM2_RBM10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.330:FF:000110">
        <property type="entry name" value="RNA-binding protein 10 isoform X1"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="RNA-binding protein 10 isoform X1"/>
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        <property type="entry name" value="RNA-binding protein 10 isoform X2"/>
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        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.1060.10">
        <property type="entry name" value="Zinc finger, RanBP2-type"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000467">
        <property type="entry name" value="G_patch_dom"/>
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    <dbReference type="InterPro" id="IPR012677">
        <property type="entry name" value="Nucleotide-bd_a/b_plait_sf"/>
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    <dbReference type="InterPro" id="IPR041591">
        <property type="entry name" value="OCRE"/>
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        <property type="entry name" value="RBD_domain_sf"/>
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    <dbReference type="InterPro" id="IPR035618">
        <property type="entry name" value="RBM10_OCRE"/>
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    <dbReference type="InterPro" id="IPR034992">
        <property type="entry name" value="RBM10_RRM2"/>
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        <property type="entry name" value="RRM_dom"/>
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    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
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        <property type="entry name" value="Znf_RanBP2"/>
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        <property type="entry name" value="Znf_RanBP2_sf"/>
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    <dbReference type="PANTHER" id="PTHR13948">
        <property type="entry name" value="RNA-BINDING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13948:SF4">
        <property type="entry name" value="RNA-BINDING PROTEIN 10"/>
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        <property type="entry name" value="OCRE"/>
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        <property type="entry name" value="RRM_1"/>
        <property type="match status" value="1"/>
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    <dbReference type="Pfam" id="PF00641">
        <property type="entry name" value="Zn_ribbon_RanBP"/>
        <property type="match status" value="1"/>
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        <property type="match status" value="2"/>
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        <property type="entry name" value="ZnF_RBZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90209">
        <property type="entry name" value="Ran binding protein zinc finger-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54928">
        <property type="entry name" value="RNA-binding domain, RBD"/>
        <property type="match status" value="2"/>
    </dbReference>
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        <property type="entry name" value="G_PATCH"/>
        <property type="match status" value="1"/>
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        <property type="match status" value="2"/>
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    <dbReference type="PROSITE" id="PS01358">
        <property type="entry name" value="ZF_RANBP2_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50199">
        <property type="entry name" value="ZF_RANBP2_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="6 8" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <keyword evidence="4 9" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="RRM" evidence="11">
        <location>
            <begin position="114"/>
            <end position="194"/>
        </location>
    </feature>
    <feature type="domain" description="RanBP2-type" evidence="14">
        <location>
            <begin position="197"/>
            <end position="227"/>
        </location>
    </feature>
    <feature type="domain" description="RRM" evidence="11">
        <location>
            <begin position="284"/>
            <end position="368"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="754"/>
            <end position="784"/>
        </location>
    </feature>
    <feature type="domain" description="G-patch" evidence="13">
        <location>
            <begin position="852"/>
            <end position="898"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1"/>
            <end position="111"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="523"/>
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    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="612"/>
            <end position="642"/>
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    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="711"/>
            <end position="746"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="811"/>
            <end position="830"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="1"/>
            <end position="14"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="21"/>
            <end position="55"/>
        </location>
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    <feature type="compositionally biased region" description="Acidic residues" evidence="10">
        <location>
            <begin position="59"/>
            <end position="70"/>
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    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="95"/>
            <end position="111"/>
        </location>
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    <feature type="compositionally biased region" description="Pro residues" evidence="10">
        <location>
            <begin position="527"/>
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    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="618"/>
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    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="811"/>
            <end position="820"/>
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    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
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            <dbReference type="ARBA" id="ARBA00023242"/>
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    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
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    </evidence>
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        <source>
            <dbReference type="PROSITE" id="PS50102"/>
        </source>
    </evidence>
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        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
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        <source>
            <dbReference type="PROSITE" id="PS50199"/>
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    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002170.1"/>
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    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="924" mass="102071" checksum="ADA49F4891BB67E8" modified="2019-05-08" version="1">MEYERRGGRGDRTGRYGAADRSQDDGVDGRTQRDHDYRDMDYRSYPRDFSSHETANEYDSSEEQSAEDSYEASSGSETQRKRDSPTEPLGYPGDGDYRDQDYRPEPEEEQKASSIIMLRMLPQSASENDIRAQLQAHGIQPREVRLMRNKASGQSRGFAFVEFNHLQDATRWMEANQHSLSILGQKVSMHYSDPKPKINEDWLCSKCGVQNFKRREKCFKCGVPKSEAEQKLPPGSRLDQLVALSGRELSQGLLPLPQPYQSSVLSATQPLAQASEPCAENANDTIILRNLNPHSTMESILAALAPYAVLSASNVRVIKDKQTQLNRGFAFIQLATIVEAAQLLQILQALHPPLHIDGKTINVEFAKGSKRDMGSSDSNRISAASVASTAIAAAQWAISQASQGSEGAWATSEEQPTDYAYYQQEEAYGAEPALYSHAYLKGSPAQPGANGEAAATVAEPSLEPGVPGVDSLPVLQSFPHAAQTTAVPVVYQPPGGAETAGTAQGTAATAQSQPYTIVSPAVLKPETPSPAQPSSAAPPAPSTGLPLPTPIPQEPYGQYPVPDVSTYQYDETSGYYYDPLTGLYYDPNSQYYYNAQSQQYLYWDGERRTYVPASDQGADAHKDGSGGSGKEGKEKKEKHKTKTAQQIAKDMERWARSLNKQKENFKNSFQPISSLREDERRESATADAGYAILEKKGALAERQHTPMDLSKLASDDRLSPPRGLVAAYSGESDSEEEPERASDREEKLTDWHKLACLLCRRQFPSKEALIRHQQLSGLHKQNLEIHRRAHLSEQELEALEKNDMEMKYRDRAAERREKYGVPEPPEPKRRKYSAVMPATVDFEQPTRDGLGSDNIGSRMLQAMGWKEGSGLGRKKQGIVTPIEAQTRVRGSGLGARGSSYGVSAAESYRETLHKTMLTRFNEAE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GKE0</accession>
    <name>A0A452GKE0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Centrosomal protein of 164 kDa</fullName>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Plays a role in microtubule organization and/or maintenance for the formation of primary cilia (PC), a microtubule-based structure that protrudes from the surface of epithelial cells. Plays a critical role in G2/M checkpoint and nuclear divisions. A key player in the DNA damage-activated ATR/ATM signaling cascade since it is required for the proper phosphorylation of H2AX, RPA, CHEK2 and CHEK1. Plays a critical role in chromosome segregation, acting as a mediator required for the maintenance of genomic stability through modulation of MDC1, RPA and CHEK1.</text>
    </comment>
    <comment type="subunit">
        <text evidence="15">Interacts (via N-terminus) with ATRIP. Interacts with ATM, ATR and MDC1. Interacts with XPA (via N-terminus) upon UV irradiation. Interacts with CEP83, CCDC92, TTBK2, DVL3, NPHP3 and weakly with NPHP4. Interacts with DZIP1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Microtubule organizing center</location>
            <location evidence="1">Centrosome</location>
            <location evidence="1">Centriole</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000002361.1">
        <property type="protein sequence ID" value="ENSGAGP00000002074.1"/>
        <property type="gene ID" value="ENSGAGG00000001670.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005814">
        <property type="term" value="C:centriole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097539">
        <property type="term" value="C:ciliary transition fiber"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060271">
        <property type="term" value="P:cilium assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006281">
        <property type="term" value="P:DNA repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
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        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Centrosomal protein of 164 kDa"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1470.10">
        <property type="entry name" value="Photosystem I PsaD, reaction center subunit II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051841">
        <property type="entry name" value="MT-Golgi_org_protein"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001202">
        <property type="entry name" value="WW_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036020">
        <property type="entry name" value="WW_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18902:SF27">
        <property type="entry name" value="CENTROSOMAL PROTEIN OF 164 KDA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18902">
        <property type="entry name" value="NUCLEAR MITOTIC APPARATUS PROTEIN 1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00397">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="WW domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01159">
        <property type="entry name" value="WW_DOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50020">
        <property type="entry name" value="WW_DOMAIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="13" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="5" id="KW-0132">Cell division</keyword>
    <keyword evidence="8" id="KW-0970">Cilium biogenesis/degradation</keyword>
    <keyword evidence="9 17" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="6" id="KW-0227">DNA damage</keyword>
    <keyword evidence="10" id="KW-0234">DNA repair</keyword>
    <keyword evidence="7" id="KW-0498">Mitosis</keyword>
    <keyword evidence="12" id="KW-0539">Nucleus</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="WW" evidence="19">
        <location>
            <begin position="58"/>
            <end position="91"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="105"/>
            <end position="151"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="163"/>
            <end position="215"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="237"/>
            <end position="263"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="315"/>
            <end position="372"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="434"/>
            <end position="467"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="505"/>
            <end position="537"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="572"/>
            <end position="604"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="630"/>
            <end position="660"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="845"/>
            <end position="896"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="972"/>
            <end position="992"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1046"/>
            <end position="1070"/>
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    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1629"/>
            <end position="1650"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="17">
        <location>
            <begin position="1087"/>
            <end position="1575"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="18">
        <location>
            <begin position="110"/>
            <end position="126"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="127"/>
            <end position="138"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="169"/>
            <end position="194"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="241"/>
            <end position="257"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="315"/>
            <end position="324"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="337"/>
            <end position="353"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="441"/>
            <end position="450"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="580"/>
            <end position="600"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="633"/>
            <end position="658"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="1060"/>
            <end position="1070"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="1635"/>
            <end position="1650"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004114"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022776"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022794"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00056906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00061715"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00067900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50020"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002074.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2020" mass="226988" checksum="AD2CD43086C7A633" modified="2019-05-08" version="1">MDMAGPAIRIGDQLILEEDYDETYIPSEQEIQEFAREIGIDPGNEPELMWLAREGIVAPLPPEWKPCQDITGDIYYFNFSNGQSTWDHPCDEHYRELVIQEREKLSAHGSIKKKDKKKKKDKKEKKEKKEREPVKHAAEMQPEPGILPSSSFYRMSSPILSSECASPDLDQHSQMRIEGFLKKGKRKPSERPPETSDSNKQFAGPSPNKLQPLFTAKSCRTHQILADVEKILGRMPSFSRSDVDHQQGQDRSPKIGDKPCLGFSDSESEGLDVVRTKPISYVARKSCSQDLENGRSVLDSCEVLEDKGMLFDEEKLSDNWKQEEMTGGSVCQPDDGLSVRERSTGPGGDKERGPGFSQADENEAFLTPSAPDHTFYSRKCEMILFDASPLEDLSLMEEGDVHCPQRELRAHSRAVESRRKRKLQDQALSQATKFCKSSEVVGKHERESSKQAEFVESEGAGLQAAPQKDLLDSHLAVPEVTADTLPYTRGVEAYAHAGADISASEPVSLSAAQHKDKEKANGLGPEPDNSLTSNSADHFASQILGEIDNFSWDLQSSRESDRPTDLLAAAAKGSLPTHPFTGTLQTPPRSSANGKSQSDCNSEDEKFYQHVQENDQVPSQEHSCVSFNMKLNRSRDAPEKNNEGRQEETGEGEQKTEPVPRGLEVDVFTEGLQVSPGWLYQTAECQESSRLCQQVEGELTQSGKVTEDYCKLKLCEGAGSNESPGSLLAPIQVPPGSLAPLRGVMDGPASAFHGLLSTNVEISGVPSPLLREGTLPLQTLKSTGCTKSLLGSMHEEKASLNLLALDEEEEEESENQSPRGTARLLKNLHMDISALGGGFEYEESLKMSHAEESQTFPLDSDGVRPPAPDKLLNRDADSSLCGLNKEESSGKPLGNELLEKEDTDVEDTVSVAYEQSGTLGERTAEGTHVVSSQGHPVAASPDVRAAETDQMESVAVTVDAVSVSEKIVNGMREEAADLETDSKLDAGKESEASEHVKELQVSDCSDHELLHLTDFGFQSRISEQVLDMDILSAALDSPKVEAQGLREEEKDQSEASLEEEQSKRPKAAERQLGEEIKQVVSLNWPSAESLNEIARKQEQELEQEKERLLQAKEERVRRFQEELRQEEEEEAQVLHQQKEKSLWSLKEELAKASEEEELRMREEEVEKLSKLRSQISSETKTEKEKIRAEQEAALQKLREEWESLQKVERERLEERKKRVLEMLKVEVEEAQQREAAELEQEKQRVLNELKERLERKRKEAAEVLEKQFAAELQELKSTAEEKHQKVVSSLRKQIAEAQRSEEAQLREELERAEQKVQQKMYQVAEFERELSELMKEKRQEVERDHDRKMERMKEEHKEVLARIQEQYEEEERKQRTKKLEELKGELERLRQLHDGELKALQKQLDEQLSDLRQSHKVKERKLQELEMELEIQATDAKARCAQLNHQEEAMRKKRQQLLDEEKQTELQRNEAALAAQLRLEESRKEHADLVESIRQLRRTLEELQDQKAELESQVDLLQTRSQRLQRRISELEAAVQSKQETLKELVAEDSMASPRREAELHLEDLRETLQAHSSREPASLTSHSNEDSDLQLDNVRHFISAEGVSIKNAKEFLVRQTRSMRKRHTALKAAKQQWRHNMQQAQDAVQDPDSSQLLEDVRRNLEEESAQLDKMKSAMQKGQVLLKKKEEKLSQLESSLLEELSDEDTLKGAACKKVVTFDLSDSEETHSLSSTDVPQHKFDLKPDVQFPQLDKIQHLTSTLQHITAELNGVLSTLGSLNQRQPPLFAPVQVPMTALPSDGVPLSMYTSLARAQAAGPIMPPAGIPLVNQWAWSTGLNSSPSSTAGQSVDDILAEKWHKYFPGGFPSLSGSPAPLDSNLGYLPAGKVGALGTTAFTRGKVTWFLIQAASACVTRPSGLPSPSRLTTGNISSTSRRRNRYGRDPFLMCRCVGFPCEQEAFRSLLWWLPGSTLACSLLSSAVWVQRFRDMSMFVRVSVLALSYKPIPTYHSPSRGSEQGTGRASV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMY4</accession>
    <name>A0A452HMY4_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="3">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMY4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018651.1">
        <property type="protein sequence ID" value="ENSGAGP00000016343.1"/>
        <property type="gene ID" value="ENSGAGG00000012116.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008028">
        <property type="term" value="F:monocarboxylic acid transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1250.20">
        <property type="entry name" value="MFS general substrate transporter like domains"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011701">
        <property type="entry name" value="MFS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036259">
        <property type="entry name" value="MFS_trans_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050327">
        <property type="entry name" value="Proton-linked_MCT"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11360">
        <property type="entry name" value="MONOCARBOXYLATE TRANSPORTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11360:SF14">
        <property type="entry name" value="MONOCARBOXYLATE TRANSPORTER 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07690">
        <property type="entry name" value="MFS_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103473">
        <property type="entry name" value="MFS general substrate transporter"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="2" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="20"/>
            <end position="47"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="110"/>
            <end position="137"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="149"/>
            <end position="169"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="224"/>
            <end position="247"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="259"/>
            <end position="279"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="285"/>
            <end position="307"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016343.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="327" mass="35567" checksum="B54F9774B3A3BAED" modified="2019-05-08" version="1">LTAMEEEEQQPIPPPDGGWGWMIVLHFFLVNVLVMGMLKTFGIFFVVFQEEVGGSSEQISWIGSIMSSLRFSAGPLVTIACGKLGDQLTSIIGAGLVSSGCLISTQATSISFLCVSMGLLLGLGFACLYQAAAVMTARYFKKRLALSNAIARSGMGLTFILAPFTQLLIDFYGWQGITETVSQLLSGWIADQNWIKKYHYHKTYLILCGITNLLGPLATSFPLLMVYTVSFAIFCGGYLALLLPVLVDFVGVPRLHSSLGFASFFAGIAAIAGPPIAGWLSDYTQTYACSFLFAGTCYLIAPISFFFEPSAHKWKLQKEKELNEVTS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNA1</accession>
    <name>A0A452HNA1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">PH domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HNA1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018777.1">
        <property type="protein sequence ID" value="ENSGAGP00000016451.1"/>
        <property type="gene ID" value="ENSGAGG00000012309.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="CDD" id="cd13359">
        <property type="entry name" value="PH_ELMO1_CED-12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000053">
        <property type="entry name" value="Engulfment and cell motility protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.810">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16457">
        <property type="entry name" value="PH_12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="PH" evidence="1">
        <location>
            <begin position="23"/>
            <end position="149"/>
        </location>
    </feature>
    <evidence type="ECO:0000259" key="1">
        <source>
            <dbReference type="Pfam" id="PF16457"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016451.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="202" mass="23342" checksum="DF3962C3E5BF8D85" modified="2019-05-08" version="1">ENQETYSSTNLKEKIQPEILELIKQQRLNRLVEGTCFRKLNSRRRQDKFWYCRLSPNHKVLHYGDLEESPQGEVPHDSLQDKLPVADIKAVVTGKDCPHMKEKGALKQNKEVLELAFSILYDSNGQLNFIAPDKHEYCIWTDGLNALLGKDMMSDLTRNDLDTLLSMEIKLRLLDLENIQIPDAPPPIPKEPSNYDFVYDCN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNA2</accession>
    <name>A0A452HNA2_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="17">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">May be involved in transcriptional regulation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HNA2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018764.1">
        <property type="protein sequence ID" value="ENSGAGP00000016439.1"/>
        <property type="gene ID" value="ENSGAGG00000012304.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000785">
        <property type="term" value="C:chromatin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031519">
        <property type="term" value="C:PcG protein complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005667">
        <property type="term" value="C:transcription regulator complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd07765">
        <property type="entry name" value="KRAB_A-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002063">
        <property type="entry name" value="RB associated KRAB zinc finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000703">
        <property type="entry name" value="Uncharacterized protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000506">
        <property type="entry name" value="Zinc finger protein 23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000247">
        <property type="entry name" value="Zinc finger protein 236"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000358">
        <property type="entry name" value="zinc finger protein 24"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002343">
        <property type="entry name" value="Zinc finger protein 33A"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001119">
        <property type="entry name" value="zinc finger protein 408"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001343">
        <property type="entry name" value="Zinc finger protein 568"/>
        <property type="match status" value="2"/>
    </dbReference>
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        <property type="entry name" value="zinc finger protein 583 isoform X1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002134">
        <property type="entry name" value="Zinc finger protein 616"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="17"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001909">
        <property type="entry name" value="KRAB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036051">
        <property type="entry name" value="KRAB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14003">
        <property type="entry name" value="TRANSCRIPTIONAL REPRESSOR PROTEIN YY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14003:SF23">
        <property type="entry name" value="ZINC FINGER PROTEIN 143"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01352">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="SMART" id="SM00349">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109640">
        <property type="entry name" value="KRAB domain (Kruppel-associated box)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50805">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="16"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="11" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="4" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="5" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13" id="KW-0539">Nucleus</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="12" id="KW-0804">Transcription</keyword>
    <keyword evidence="10" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="9" id="KW-0832">Ubl conjugation</keyword>
    <keyword evidence="8" id="KW-0862">Zinc</keyword>
    <keyword evidence="7 14" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="KRAB" evidence="16">
        <location>
            <begin position="11"/>
            <end position="83"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="105"/>
            <end position="132"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="133"/>
            <end position="160"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="161"/>
            <end position="188"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="189"/>
            <end position="216"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="220"/>
            <end position="247"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="248"/>
            <end position="275"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="276"/>
            <end position="303"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="304"/>
            <end position="331"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="332"/>
            <end position="359"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="360"/>
            <end position="387"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="388"/>
            <end position="415"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="416"/>
            <end position="443"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="444"/>
            <end position="471"/>
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        <location>
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        <location>
            <begin position="500"/>
            <end position="527"/>
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    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="528"/>
            <end position="555"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00003767"/>
        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
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        <source>
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    </evidence>
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    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
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    </evidence>
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        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016439.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="584" mass="67434" checksum="36AFF0E053C37CCD" modified="2019-05-08" version="1">MTLTLLFQMPVTFEEVAVYFTQGQGALLDPAQRALYRDVMQENYETRLLRNYSREQMDEFIICGEGYRDPTAQQTTPKEDKHHEWFGYGNEFILRPQFVSPQKSYQCFKSRNSLNWKSALTGHQISHTGEKPHKCLDCGKSFIHRSALVNHQAIHTGERPHKCLDCGKSFTHRSNLKHHQWIHTGERPHKCLHCGKGFVHRSDLVKHQAIHTGERPYKPHKCLNCGKSFIQRSDLVKHQAIHTGQRPHKCLDCGKSFIRRSNLVKHQAIHTGERPYKCLDCGKSFIQRSNLVKHQAIHTGERSHKCLDCGKSFVWKSGLANHQAIHTGERSHKCLDCGKSFIQRSYLLKHQAIHTGERPHMCLDCGKSFIQRSNLITHQKIHTGERSHKCLDCGKNFIWKSALVKHQAVHTGERSHKCLDCGKSFIHKSDLVKHQAIHTGERPHTCLDCRKSFIQGSDLVKHQAIHTGERPHRCLDCGKSFTQRSHLKTHQRIHTGERPHKCLDCGKSFIQRSDLVKHQIIHTGERPHKCLDCGKSFIRRSTLLNHQTIHTGERPHKCLDCGKNMLKKFVNDIWGHSKLYRLEL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNA6</accession>
    <name>A0A452HNA6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Microtubule-associated protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 7">Cytoplasm</location>
            <location evidence="1 7">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018750.1">
        <property type="protein sequence ID" value="ENSGAGP00000016432.1"/>
        <property type="gene ID" value="ENSGAGG00000012294.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="term" value="C:microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043005">
        <property type="term" value="C:neuron projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008017">
        <property type="term" value="F:microtubule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000226">
        <property type="term" value="P:microtubule cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031175">
        <property type="term" value="P:neuron projection development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027324">
        <property type="entry name" value="MAP2/MAP4/Tau"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013588">
        <property type="entry name" value="MAP2_projctn"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001084">
        <property type="entry name" value="MAP_tubulin-bd_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11501">
        <property type="entry name" value="MICROTUBULE-ASSOCIATED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11501:SF15">
        <property type="entry name" value="MICROTUBULE-ASSOCIATED PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08377">
        <property type="entry name" value="MAP2_projctn"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00418">
        <property type="entry name" value="Tubulin-binding"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00229">
        <property type="entry name" value="TAU_MAP_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51491">
        <property type="entry name" value="TAU_MAP_2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2 7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6 7" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="4 7" id="KW-0493">Microtubule</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="MAP2/Tau projection" evidence="9">
        <location>
            <begin position="366"/>
            <end position="1538"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1"/>
            <end position="90"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="108"/>
            <end position="283"/>
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    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
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            <end position="412"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="433"/>
            <end position="584"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="599"/>
            <end position="646"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="916"/>
            <end position="980"/>
        </location>
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    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1076"/>
            <end position="1203"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1215"/>
            <end position="1278"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1332"/>
            <end position="1391"/>
        </location>
    </feature>
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        <location>
            <begin position="1426"/>
            <end position="1522"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1537"/>
            <end position="1673"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1813"/>
            <end position="1837"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="1"/>
            <end position="11"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="14"/>
            <end position="26"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="81"/>
            <end position="90"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="108"/>
            <end position="119"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="170"/>
            <end position="191"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="200"/>
            <end position="215"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="251"/>
            <end position="270"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="328"/>
            <end position="363"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="441"/>
            <end position="452"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="465"/>
            <end position="555"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="601"/>
            <end position="611"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="628"/>
            <end position="645"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="916"/>
            <end position="925"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="938"/>
            <end position="948"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="955"/>
            <end position="980"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="1084"/>
            <end position="1094"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="1111"/>
            <end position="1120"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="1129"/>
            <end position="1168"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="1263"/>
            <end position="1278"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="8">
        <location>
            <begin position="1358"/>
            <end position="1370"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="1440"/>
            <end position="1458"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="8">
        <location>
            <begin position="1459"/>
            <end position="1468"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="1473"/>
            <end position="1492"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="1499"/>
            <end position="1509"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="1555"/>
            <end position="1572"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="1574"/>
            <end position="1585"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="1587"/>
            <end position="1596"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="1597"/>
            <end position="1609"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="1622"/>
            <end position="1639"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="1645"/>
            <end position="1664"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="1818"/>
            <end position="1837"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU000686"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF08377"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016432.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1863" mass="205861" checksum="E1979AF5C7ECA463" modified="2019-05-08" version="1">MADDRKDEAKAPHWTSSQLTDASSHPHSPEIKEQAGAGAELVRSANGFPYREDEEAGYGERGQQGTYSRTKENGINGELSTGDRETAEEVSARIVQVVTAEAVAVLKGEQEKEAQHKDQPGPLPLVEESANLPPSPPPSPASEQTGTLAEASKMEIRDQEGTGPFTAEPLDTKERESEKDSKTDEQSKHDALVPQPAKTETLDKKESWGKEEKMHPTVSGQTLDADLQKKGEVSFADPVADSTAEPLVSQEHMDLSADLLKESKTDERSIDTQIPSNQAIPLKYKDDFTETQKSNLALLPSESRIDLSSAARSEPLSETPMTVSQKSQLEDSCKTKQEFSDKEIAKDLLKDEKIHKDQTEKPEAGQVSSMAADAPKPTTEERPIQKVSTWGEEKDMTRDESDEEERYGFLDWQPSKVLDDTKLYAEQEDKIVLPDKAVVSEAKKSSDGLKAEEETDRTGVSTVVDAKKEEQQSTETHTGKLSHESVPEKAKEPIDTDHLSKPTEKAPDAPRLTSEKGPTHELHQDRTIEKDAKEDRFHISDSASHEIQVEEDRSGMSKYFETSALKEEATKSDTQQQGSDYYELSDTKESAYELYHADPLLSKKEDKKLQPETDQPSTVPAHEIGYSTLAQSFPSDKSEEPSSPQERMFTIDPKVYGDKRQIHSKNKDDLTLSRSLGLGGRSAIEQRSMSINLPMSCLDSIALGFSFGRAQDLSPLASDILTNTSGSMDEGDDYLPATTPALERAPCFPIDSTEEDENVEEEKAMEEENIQVESLCESPFLAKDYYKNGAVMAPDLPEMLDLAGTRSRLASVSADTEVARRKSVPSDAAVEDSGASLPPVTDENHITLKAESQLEDLGYCVFNKYTVPLPSPVQDSESLTGEICPFYEGRDEKMRRGLAADLSLIEMKLAAAEKSKEEFSSEKDLSPNISSDTTVLGREIEQEKKGSDKLATVLEKSEEQVDSKEIHSLEDTKHEKKRPDAILEMKEEGTVDKLHIAGPESDIANDKALEKEVTKEKDLVSLLTQKEEKSLNIIPKVAEIETPHRPDYDAIKHDMDVAARKADQEYQTQLHTEINEFAAAGSKQDTDSVKKAEPEAQEAQQQQKDLASLPKEPKEADVLSRTEPSQMKDSTKLSESEIKEKVPKPDLVHQEAVDKEESYESSGEHDQIQEDLNGESLKPDYGKIKPQKPPVSEEEITSKLPVEEPSIELAISKADSVQEETADIQMESIPQPKEEIKETTPDIAVRPAETTEQLLSDLTPESVEGKEHKEEEDDDKHQLMSDVHQEIELEAPAGEEALTEAGPEEMPELKGVIESVVTIEDDFITVVQMTVEESESASHSVRFAAAQQEEMERRDFLPEEEVEMEMEEPADIQAEPKEGSLEAPASPEKEEILLTDYKTEIYDDFKDETTIDDSVMDTDSLWADTQDDDRSIMTEQLETVPKEEKTERELRRSSLDKHKKEKPFKTGRGRISTPERKIAKKEPSTLSRDEVRRKKAVYKKAELAKKSEVQAHSPSRKIILKPAIKYTRPTHLSCVKRKQTAAGGETNQAPGVFKQAKEKLSDGITKSPEKRSSLPRPSSILPTRRGVSGDRDRDENSFSLNSSISSSVRRTTRSEPIRSRTGKSGTSTPTTPGSTAITPGTPPSYSSRTPGTPGTPSYSRTPHTPGTPKSAILVPTEKKVAIIRTPPKSPATPKQLRLINQPLPDLKNVRSKIGSTDNIRYQPKGGQIQIVTKKIDLSHVTSKCGSLKNIRHRPGGGRVKIESVKLDFKEKAHAKVGSLENAHHTPGGGNIKIDSQKLNFREHAKARVDHGAEIITQSPGRSSVPSPRRLSNVSSSGSINLLESPQLATLAEDVTAALAKQGL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNB0</accession>
    <name>A0A452HNB0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Erbin</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="13">Densin-180-like protein</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="15">Erbb2-interacting protein</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="14">Protein LAP2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Basolateral cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="11">Cell junction</location>
            <location evidence="11">Hemidesmosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the LAP (LRR and PDZ) protein family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018797.1">
        <property type="protein sequence ID" value="ENSGAGP00000016469.1"/>
        <property type="gene ID" value="ENSGAGG00000006183.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005912">
        <property type="term" value="C:adherens junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016323">
        <property type="term" value="C:basolateral plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030056">
        <property type="term" value="C:hemidesmosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031965">
        <property type="term" value="C:nuclear membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0014069">
        <property type="term" value="C:postsynaptic density"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019901">
        <property type="term" value="F:protein kinase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098609">
        <property type="term" value="P:cell-cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045197">
        <property type="term" value="P:establishment or maintenance of epithelial cell apical/basal polarity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070433">
        <property type="term" value="P:negative regulation of nucleotide-binding oligomerization domain containing 2 signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098968">
        <property type="term" value="P:neurotransmitter receptor transport postsynaptic membrane to endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098887">
        <property type="term" value="P:neurotransmitter receptor transport, endosome to postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043113">
        <property type="term" value="P:receptor clustering"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010468">
        <property type="term" value="P:regulation of gene expression"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd06749">
        <property type="entry name" value="PDZ_densin_erbin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000036">
        <property type="entry name" value="Erbin isoform 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000013">
        <property type="entry name" value="Erbin isoform 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000020">
        <property type="entry name" value="Erbin isoform 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000022">
        <property type="entry name" value="Erbin isoform 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.42.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001611">
        <property type="entry name" value="Leu-rich_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003591">
        <property type="entry name" value="Leu-rich_rpt_typical-subtyp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001478">
        <property type="entry name" value="PDZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036034">
        <property type="entry name" value="PDZ_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050614">
        <property type="entry name" value="Synaptic_Scaffolding_LAP-MAGUK"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23119">
        <property type="entry name" value="DISCS LARGE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23119:SF46">
        <property type="entry name" value="ERBB2 INTERACTING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13855">
        <property type="entry name" value="LRR_8"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00595">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00364">
        <property type="entry name" value="LRR_BAC"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SMART" id="SM00365">
        <property type="entry name" value="LRR_SD22"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00369">
        <property type="entry name" value="LRR_TYP"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="SMART" id="SM00228">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52058">
        <property type="entry name" value="L domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50156">
        <property type="entry name" value="PDZ domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51450">
        <property type="entry name" value="LRR"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50106">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0965">Cell junction</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="6" id="KW-0433">Leucine-rich repeat</keyword>
    <keyword evidence="9" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="PDZ" evidence="17">
        <location>
            <begin position="1191"/>
            <end position="1280"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="449"/>
            <end position="475"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="492"/>
            <end position="590"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="617"/>
            <end position="673"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="981"/>
            <end position="1000"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1055"/>
            <end position="1079"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1114"/>
            <end position="1173"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="16">
        <location>
            <begin position="456"/>
            <end position="466"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="16">
        <location>
            <begin position="492"/>
            <end position="520"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="553"/>
            <end position="568"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="16">
        <location>
            <begin position="578"/>
            <end position="588"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="16">
        <location>
            <begin position="632"/>
            <end position="644"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="645"/>
            <end position="663"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1055"/>
            <end position="1068"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1137"/>
            <end position="1157"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004126"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004187"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007772"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022614"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00060371"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00074479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00076296"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00083415"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00084015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50106"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016469.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1282" mass="143508" checksum="24C5DB1827FDCCEF" modified="2019-05-08" version="1">MTTKRNLFVRLVPCRCLRGEEETVTALDYSHCSLEQVPKEIFTFEKTLEELYLDANQIEELPKQLFNCQSLHKLSLPDNDLTTLPASIANLINLRELDVSKNGIQEFPENIKNCKVLAIVEASVNPISKLPDGFSQLLNLTQLYLNDAFLEFLPANFGRLTKLQILELRENQLKMLPKTMNRLTQLERLDLGSNEFTEVPEVLEQLSGLKEFWMDGNRLTLIPGLIGSLKQLTYLDVSKNNIEIVEEGVSGCESLQDLLLSSNSLQQLPESIGSLKKVTTLKIDENQLMYLPDSIGGLIAVEELDCSFNEIEALPSSIGQLTNVRTFASDHNFLTQLPPEIGNWKNVTVLFLHSNKLEFLPEEMGDMQKLKVINLSDNRDFNYFFFSQSKPLIPLQKEVDPETQKTVLTNYMFPQQPRSEDVLFISDNESFNPSLWEEQRKQRAQVAFECDEDKDEREAPPREGNLKRYPTPYPDELKNMVKTVQTIVHRLKDEETTEDGGRGSKKEGQQVSVKDVDVKTSEMASAKSKTEDGENYMMENCVQKPTEPEAEHSTGNSQMTSHVKTLENTKIIVNHEDTLEESEELSSDEEMKMAEMRPPLIETSINQPKVVALSNNKKDKDADSLSDEATHNSNHNNSNCSSPSRMSDSVSLNTDSSNDTSVCSPDKETHSAAISKIRQEDENFNSLLQNGSELNITAEENISAQEKVTNLPEYELRIEERLALIEKGVDLNTTTEESHKLDQINMNINRLTSEDTIQPNVMERLKTQEIVSVKGCSNYNMKEETEHLENRNQYSNLCSINKVNGHSEREPQSLRKAELKRSITVDTDSSVDICISKSTEDLSPQRSGPVVKSHSITNIDNGALKIFDIVNDNGSQEPNSVVKSASAGPDGKNIVRSKSATLLYDHPLQVFPGSSSSSDLVSCTKTLFKFDSNHNPEGANVVRGPVASGAQMFSVPQYNIQYSSSTTAKDTLWLPKQNAQIEQGSLPPQRLSRSDSTESSNYVKHSANMNFSNHNNVRANAAYNMHQRMGGRHIEMWAIPPNDRLVPGATRNTLQRQSSISSTASINVGDTGPPRRAQVAEGDCLTYRDLHSMGRAPPIMSGQPRPLSARTYSIDGPNVPRPQSARPSVNEIPERTMSVSDFNYSRTSPTKRPNPRVNSEHSLLDPPGKSKVPHDWREQVLRHIEAKKLEKIRVRIEKDPELGFSISGGVGGRGNPFRPDDDGIFVTRVQPEGPASKLLQPGDKIIQANGYSFINIDHGQAVSLLKTFQNTVELIILREVSS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNB4</accession>
    <name>A0A452HNB4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Erbin</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="13">Densin-180-like protein</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="15">Erbb2-interacting protein</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="14">Protein LAP2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Basolateral cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="11">Cell junction</location>
            <location evidence="11">Hemidesmosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the LAP (LRR and PDZ) protein family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018782.1">
        <property type="protein sequence ID" value="ENSGAGP00000016456.1"/>
        <property type="gene ID" value="ENSGAGG00000006183.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005912">
        <property type="term" value="C:adherens junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016323">
        <property type="term" value="C:basolateral plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030056">
        <property type="term" value="C:hemidesmosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031965">
        <property type="term" value="C:nuclear membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0014069">
        <property type="term" value="C:postsynaptic density"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019901">
        <property type="term" value="F:protein kinase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098609">
        <property type="term" value="P:cell-cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045197">
        <property type="term" value="P:establishment or maintenance of epithelial cell apical/basal polarity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070433">
        <property type="term" value="P:negative regulation of nucleotide-binding oligomerization domain containing 2 signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098968">
        <property type="term" value="P:neurotransmitter receptor transport postsynaptic membrane to endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098887">
        <property type="term" value="P:neurotransmitter receptor transport, endosome to postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043113">
        <property type="term" value="P:receptor clustering"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010468">
        <property type="term" value="P:regulation of gene expression"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd06749">
        <property type="entry name" value="PDZ_densin_erbin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000036">
        <property type="entry name" value="Erbin isoform 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000013">
        <property type="entry name" value="Erbin isoform 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000020">
        <property type="entry name" value="Erbin isoform 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000022">
        <property type="entry name" value="Erbin isoform 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.42.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001611">
        <property type="entry name" value="Leu-rich_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003591">
        <property type="entry name" value="Leu-rich_rpt_typical-subtyp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001478">
        <property type="entry name" value="PDZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036034">
        <property type="entry name" value="PDZ_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050614">
        <property type="entry name" value="Synaptic_Scaffolding_LAP-MAGUK"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23119">
        <property type="entry name" value="DISCS LARGE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23119:SF46">
        <property type="entry name" value="ERBB2 INTERACTING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13855">
        <property type="entry name" value="LRR_8"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00595">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00364">
        <property type="entry name" value="LRR_BAC"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SMART" id="SM00365">
        <property type="entry name" value="LRR_SD22"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00369">
        <property type="entry name" value="LRR_TYP"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="SMART" id="SM00228">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52058">
        <property type="entry name" value="L domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50156">
        <property type="entry name" value="PDZ domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51450">
        <property type="entry name" value="LRR"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50106">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0965">Cell junction</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="6" id="KW-0433">Leucine-rich repeat</keyword>
    <keyword evidence="9" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="PDZ" evidence="17">
        <location>
            <begin position="1252"/>
            <end position="1341"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="474"/>
            <end position="500"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="517"/>
            <end position="555"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="650"/>
            <end position="688"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1002"/>
            <end position="1021"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1076"/>
            <end position="1100"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1134"/>
            <end position="1194"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="16">
        <location>
            <begin position="481"/>
            <end position="491"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="16">
        <location>
            <begin position="517"/>
            <end position="545"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="16">
        <location>
            <begin position="653"/>
            <end position="665"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="666"/>
            <end position="684"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1076"/>
            <end position="1089"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1158"/>
            <end position="1178"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004126"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004187"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007772"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022614"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00060371"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00074479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00076296"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00083415"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00084015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50106"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016456.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1343" mass="150239" checksum="A602393D2B1FB11A" modified="2019-05-08" version="1">MSSDGDSTTSFGSPIQYLTILICLKMTTKRNLFVRLVPCRCLRGEEETVTALDYSHCSLEQVPKEIFTFEKTLEELYLDANQIEELPKQLFNCQSLHKLSLPDNDLTTLPASIANLINLRELDVSKNGIQEFPENIKNCKVLAIVEASVNPISKLPDGFSQLLNLTQLYLNDAFLEFLPANFGRLTKLQILELRENQLKMLPKTMNRLTQLERLDLGSNEFTEVPEVLEQLSGLKEFWMDGNRLTLIPGLIGSLKQLTYLDVSKNNIEIVEEGVSGCESLQDLLLSSNSLQQLPESIGSLKKVTTLKIDENQLMYLPDSIGGLIAVEELDCSFNEIEALPSSIGQLTNVRTFASDHNFLTQLPPEIGNWKNVTVLFLHSNKLEFLPEEMGDMQKLKVINLSDNRDFNYFFFSQSKPLIPLQKEVDPETQKTVLTNYMFPQQPRSEDVLFISDNESFNPSLWEEQRKQRAQVAFECDEDKDEREAPPREGNLKRYPTPYPDELKNMVKTVQTIVHRLKDEETTEDGGRGSKKEGQQVSVKDVDVKTSEMASAKSKTEDGENYMMENCVQKPTEPEAEHSTGNSQMTSHVKTLENTKIIVNHEDTLEESEELSSDEEMKMAEMRPPLIETSINQPKVITKHTKSRVLLYDEATHNSNHNNSNCSSPSRMSDSVSLNTDSSNDTSVCSPDKETHSAAISKIRQEDENFNSLLQNGSELNITAEENISAQEKVTNLPEYELRIEERLALIEKGVDLNTTTEESHKLDQINMNINRLTSEDTIQPNVMERLKTQEIVSVKGCSNYNMKEETEHLENRNQYSNLCSINKVNGHSEREPQSLRKAELKRSITVDTDSSVDICISKSTEDLSPQRSGPVVKSHSITNIDNGALKIFDIVNDNGSQEPNSVVKSASAGPDGKNIVRSKSATLLYDHPLQVFPGSSSSSDLVSCTKTLFKFDSNHNPEGANVVRGPVASGAQMFSVPQYNIQYSSSTTAKDTLWLPKQNAQIEQGSLPPQRLSRSDSTESSNYVKHSANMNFSNHNNVRANAAYNMHQRMGGRHIEMWAIPPNDRLVPGATRNTLQRQSSISSTASINVGDTGPPRRAQVAEGDCLTYRDLHSMGRAPPIMSGQPRPLSARTYSIDGPNVPRPQSARPSVNEIPERTMSVSDFNYSRTSPTKRPNPRVNSEHSLLDPPGKSKVPHDWREQVLRHIEAKKLEKHPQTSTAGEPCQEDVFISGQQNYSSGTLNLKDVPPDSMKKIRVRIEKDPELGFSISGGVGGRGNPFRPDDDGIFVTRVQPEGPASKLLQPGDKIIQANGYSFINIDHGQAVSLLKTFQNTVELIILREVSS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNC1</accession>
    <name>A0A452HNC1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">PH domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HNC1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018787.1">
        <property type="protein sequence ID" value="ENSGAGP00000016461.1"/>
        <property type="gene ID" value="ENSGAGG00000012309.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.810">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16457">
        <property type="entry name" value="PH_12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="PH" evidence="1">
        <location>
            <begin position="68"/>
            <end position="133"/>
        </location>
    </feature>
    <evidence type="ECO:0000259" key="1">
        <source>
            <dbReference type="Pfam" id="PF16457"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016461.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="135" mass="16195" checksum="BA63FB96CD3D71BA" modified="2019-05-08" version="1">MQVVKEQIMRALTTKPSSLDQFKSKLQNLSYTEILKIRQSERMNQEDFQSRPILELKEKIQPEILELIKQQRLNRLVEGTCFRKLNSRRRQDKFWYCRLSPNHKVLHYGDLEESPQGEVPHDSLQDKCNIWEINP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNC6</accession>
    <name>A0A452HNC6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">IGFBP N-terminal domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HNC6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018829.1">
        <property type="protein sequence ID" value="ENSGAGP00000016497.1"/>
        <property type="gene ID" value="ENSGAGG00000012345.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031012">
        <property type="term" value="C:extracellular matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045597">
        <property type="term" value="P:positive regulation of cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.40.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050941">
        <property type="entry name" value="CCN"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009030">
        <property type="entry name" value="Growth_fac_rcpt_cys_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000867">
        <property type="entry name" value="IGFBP-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017891">
        <property type="entry name" value="Insulin_GF-bd_Cys-rich_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11348:SF3">
        <property type="entry name" value="CELLULAR COMMUNICATION NETWORK FACTOR 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11348">
        <property type="entry name" value="CONNECTIVE TISSUE GROWTH FACTOR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00219">
        <property type="entry name" value="IGFBP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00121">
        <property type="entry name" value="IB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57603">
        <property type="entry name" value="FnI-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57184">
        <property type="entry name" value="Growth factor receptor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00222">
        <property type="entry name" value="IGFBP_N_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51323">
        <property type="entry name" value="IGFBP_N_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1 3" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="3">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019530136" description="IGFBP N-terminal domain-containing protein" evidence="3">
        <location>
            <begin position="23"/>
            <end position="254"/>
        </location>
    </feature>
    <feature type="domain" description="IGFBP N-terminal" evidence="4">
        <location>
            <begin position="45"/>
            <end position="117"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS51323"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016497.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="254" mass="28168" checksum="DB952412545D8536" modified="2019-05-08" version="1">MQWLLILTLFTIIYIQQFFCRAQGNGQLEVKPGEKSAEASEVHQRKEFCHWPCKCPHVPTCTPGVSLVKDGCECCKICAKQAGETCNEAETCDPYKGLYCDYSADRPRYETGVCAYMIAVGCELNGVYYVNGQAFKPTPLYKCLCVSGAIGCTPVFTPKAAASQCVMVKGRKKSGHSNCGPGQQKQLQSTGYRSMPDLCQAFNLLPALRSLTCCLPSRLQKSITCFEGEVLSTGNFLDTLFQNLWHRYIQQSDQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNC7</accession>
    <name>A0A452HNC7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Erbin</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="13">Densin-180-like protein</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="15">Erbb2-interacting protein</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="14">Protein LAP2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Basolateral cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="11">Cell junction</location>
            <location evidence="11">Hemidesmosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the LAP (LRR and PDZ) protein family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018778.1">
        <property type="protein sequence ID" value="ENSGAGP00000016452.1"/>
        <property type="gene ID" value="ENSGAGG00000006183.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005912">
        <property type="term" value="C:adherens junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016323">
        <property type="term" value="C:basolateral plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030056">
        <property type="term" value="C:hemidesmosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031965">
        <property type="term" value="C:nuclear membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0014069">
        <property type="term" value="C:postsynaptic density"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019901">
        <property type="term" value="F:protein kinase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098609">
        <property type="term" value="P:cell-cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045197">
        <property type="term" value="P:establishment or maintenance of epithelial cell apical/basal polarity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070433">
        <property type="term" value="P:negative regulation of nucleotide-binding oligomerization domain containing 2 signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098968">
        <property type="term" value="P:neurotransmitter receptor transport postsynaptic membrane to endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098887">
        <property type="term" value="P:neurotransmitter receptor transport, endosome to postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043113">
        <property type="term" value="P:receptor clustering"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010468">
        <property type="term" value="P:regulation of gene expression"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd06749">
        <property type="entry name" value="PDZ_densin_erbin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000036">
        <property type="entry name" value="Erbin isoform 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000013">
        <property type="entry name" value="Erbin isoform 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000020">
        <property type="entry name" value="Erbin isoform 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000022">
        <property type="entry name" value="Erbin isoform 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.42.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001611">
        <property type="entry name" value="Leu-rich_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003591">
        <property type="entry name" value="Leu-rich_rpt_typical-subtyp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001478">
        <property type="entry name" value="PDZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036034">
        <property type="entry name" value="PDZ_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050614">
        <property type="entry name" value="Synaptic_Scaffolding_LAP-MAGUK"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23119">
        <property type="entry name" value="DISCS LARGE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23119:SF46">
        <property type="entry name" value="ERBB2 INTERACTING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13855">
        <property type="entry name" value="LRR_8"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00595">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00364">
        <property type="entry name" value="LRR_BAC"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SMART" id="SM00365">
        <property type="entry name" value="LRR_SD22"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00369">
        <property type="entry name" value="LRR_TYP"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="SMART" id="SM00228">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52058">
        <property type="entry name" value="L domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50156">
        <property type="entry name" value="PDZ domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51450">
        <property type="entry name" value="LRR"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50106">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0965">Cell junction</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="6" id="KW-0433">Leucine-rich repeat</keyword>
    <keyword evidence="9" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="PDZ" evidence="17">
        <location>
            <begin position="1282"/>
            <end position="1371"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="474"/>
            <end position="500"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="517"/>
            <end position="555"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="650"/>
            <end position="688"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1002"/>
            <end position="1021"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1076"/>
            <end position="1100"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1135"/>
            <end position="1194"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="16">
        <location>
            <begin position="481"/>
            <end position="491"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="16">
        <location>
            <begin position="517"/>
            <end position="545"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="16">
        <location>
            <begin position="653"/>
            <end position="665"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="666"/>
            <end position="684"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1076"/>
            <end position="1089"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1158"/>
            <end position="1178"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004126"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004187"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007772"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022614"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00060371"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00074479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00076296"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00083415"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00084015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50106"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016452.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1373" mass="153847" checksum="361E78A2626508C2" modified="2019-05-08" version="1">MSSDGDSTTSFGSPIQYLTILICLKMTTKRNLFVRLVPCRCLRGEEETVTALDYSHCSLEQVPKEIFTFEKTLEELYLDANQIEELPKQLFNCQSLHKLSLPDNDLTTLPASIANLINLRELDVSKNGIQEFPENIKNCKVLAIVEASVNPISKLPDGFSQLLNLTQLYLNDAFLEFLPANFGRLTKLQILELRENQLKMLPKTMNRLTQLERLDLGSNEFTEVPEVLEQLSGLKEFWMDGNRLTLIPGLIGSLKQLTYLDVSKNNIEIVEEGVSGCESLQDLLLSSNSLQQLPESIGSLKKVTTLKIDENQLMYLPDSIGGLIAVEELDCSFNEIEALPSSIGQLTNVRTFASDHNFLTQLPPEIGNWKNVTVLFLHSNKLEFLPEEMGDMQKLKVINLSDNRDFNYFFFSQSKPLIPLQKEVDPETQKTVLTNYMFPQQPRSEDVLFISDNESFNPSLWEEQRKQRAQVAFECDEDKDEREAPPREGNLKRYPTPYPDELKNMVKTVQTIVHRLKDEETTEDGGRGSKKEGQQVSVKDVDVKTSEMASAKSKTEDGENYMMENCVQKPTEPEAEHSTGNSQMTSHVKTLENTKIIVNHEDTLEESEELSSDEEMKMAEMRPPLIETSINQPKVITKHTKSRVLLYDEATHNSNHNNSNCSSPSRMSDSVSLNTDSSNDTSVCSPDKETHSAAISKIRQEDENFNSLLQNGSELNITAEENISAQEKVTNLPEYELRIEERLALIEKGVDLNTTTEESHKLDQINMNINRLTSEDTIQPNVMERLKTQEIVSVKGCSNYNMKEETEHLENRNQYSNLCSINKVNGHSEREPQSLRKAELKRSITVDTDSSVDICISKSTEDLSPQRSGPVVKSHSITNIDNGALKIFDIVNDNGSQEPNSVVKSASAGPDGKNIVRSKSATLLYDHPLQVFPGSSSSSDLVSCTKTLFKFDSNHNPEGANVVRGPVASGAQMFSVPQYNIQYSSSTTAKDTLWLPKQNAQIEQGSLPPQRLSRSDSTESSNYVKHSANMNFSNHNNVRANAAYNMHQRMGGRHIEMWAIPPNDRLVPGATRNTLQRQSSISSTASINVGDTGPPRRAQVAEGDCLTYRDLHSMGRAPPIMSGQPRPLSARTYSIDGPNVPRPQSARPSVNEIPERTMSVSDFNYSRTSPTKRPNPRVNSEHSLLDPPGKSKVPHDWREQVLRHIEAKKLEKIPLTNGQMCQPHRPQTNYSQVHHPASQASVARHPSREQLIDYLMLKVAHQPPFTQPQCSPKQGNEMAKQEIRVRIEKDPELGFSISGGVGGRGNPFRPDDDGIFVTRVQPEGPASKLLQPGDKIIQANGYSFINIDHGQAVSLLKTFQNTVELIILREVSS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNC8</accession>
    <name>A0A452HNC8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">DUF4550 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000018814.1">
        <property type="protein sequence ID" value="ENSGAGP00000016485.1"/>
        <property type="gene ID" value="ENSGAGG00000012332.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027876">
        <property type="entry name" value="DUF4550"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR33667:SF7">
        <property type="entry name" value="RIKEN CDNA 1810020O05 GENE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR33667">
        <property type="entry name" value="SI:DKEY-57N24.6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15084">
        <property type="entry name" value="DUF4550"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="DUF4550" evidence="2">
        <location>
            <begin position="132"/>
            <end position="225"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1"/>
            <end position="70"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="360"/>
            <end position="399"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="1"/>
            <end position="10"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="1">
        <location>
            <begin position="28"/>
            <end position="39"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="360"/>
            <end position="376"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF15084"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016485.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1298" mass="147530" checksum="59BE710A2EFED48B" modified="2019-05-08" version="1">MALQKQKEDVSGELTEPFAQDSKTKMVEEEEEGLVEVELEALGSESSFETETPDLLGDGGHPTPEDEDGRPLFFDKNLEQLDTSHTVMCTFSISLAVPMAAGPKHRTFNLTDRQRKKSNIGKDAAAIPKMRRYYHIEYFLLPDDVEPRKLDLALFGVVAKLFVECESKVVKPWFENDQIWVSWNHSINIHVTNEFLIKLRDHKVTLTVWDTKDKVSAKARFSKPRVPFSQVEDSDAIGGVKYTVLLQRKLFEDNQPELSCVKMKGVKVPSAPEKPAMGFAAGESKPLPEEDELCETPASLSVFQSLLPPSRTDDFPSVSRFSVSGSIVSNRDKILNEGNYLKQNTIAGFSDYVIPKKPEKTKFSLENPADHPDKKDPKGKRKSSTPKRGPALKERRQSKVSNAETLIAALTKKCGIASFQLDLMPLLSGERLVISRLQEKSSKILDAYMTFTTEEPLMSERQISELNPLIIKILSATCLPMTPVPIEVLQATCVPAYCRYQFHKLPPHQTQGQTHGTHIYFKDVNVILTGTISPGELREYFRGPPLEIEVHDRDRKMAKDIKKPSLFGEEQDDGKLSNVGFVSYKHTAYNPFTEKDNLWHPYGVAKVSLTDLLLGEKYLNISVPIHSCSVPDPGYQKDSKTEKIMGVLGSVAGPQSSPLPMGHYLESDSVLKVRVEIAVPLGLQAETADTEVANCPYGCIIYIFDYNNAPCLHDLLQEITEINAEALQLDSYPLQVIQMALATFKLKTTTKEISDLDIITGFHILDGTIHLLVLEGLKDKAIKRLWEQQFDKTHRTENGRLDILYNSQLSFHQRLYTDLEAILYHIHLCKPLSAITRQPLLYVRDMVRQACFQALSRLDYLCHSKKLRDVVQRGLLPSAEMITVLSQEFGIPLTKEDLFIQRPPLPSVSSFALQKSRKVPTKRGAMHSLLDNHNEKYIQCKKEMEDKMQHYKDHIQTNIDAVYLLSRKVKKPKFRAIRSSPADGKSVYNYSSQHLNSAERAKMQLRKELAKKPERRFAYCHEYLSAMFDPVDLDSVRKESIAKSKNLWVSPRGFVCPGFKSYIESNLHPRMPDETRLKELTEKWQENILFANILEPVLSRDRWSWDQRHIDFDLYKKPPVYLPASAPATHRAVDTWLHEDSSQLQKTVFDDTKLKVYRCPTAAELTTRGPKASAQLDKLQGLLKDEPEKFSLKRPAMILRPIPALSVLQHIPACGSYPDLSRKRSTTNGFVPGLEDQHSLKWNGNIVPCRDMEHRKFEKLKGADFNLLCYEHSFLYKRKRPEKKGEDTTLLLQQDSTQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HND2</accession>
    <name>A0A452HND2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Calsyntenin-1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="26 27">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="27" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="26" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="26" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell projection</location>
            <location evidence="2">Neuron projection</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Golgi apparatus membrane</location>
            <topology evidence="3">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="18">Postsynaptic cell membrane</location>
            <topology evidence="18">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="19">Belongs to the calsyntenin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HND2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018785.1">
        <property type="protein sequence ID" value="ENSGAGP00000016459.1"/>
        <property type="gene ID" value="ENSGAGG00000012316.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009986">
        <property type="term" value="C:cell surface"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000139">
        <property type="term" value="C:Golgi membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043005">
        <property type="term" value="C:neuron projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007156">
        <property type="term" value="P:homophilic cell adhesion via plasma membrane adhesion molecules"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051965">
        <property type="term" value="P:positive regulation of synapse assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050806">
        <property type="term" value="P:positive regulation of synaptic transmission"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd11304">
        <property type="entry name" value="Cadherin_repeat"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.200:FF:000036">
        <property type="entry name" value="Calsyntenin 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.60:FF:000025">
        <property type="entry name" value="Calsyntenin 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.60:FF:000062">
        <property type="entry name" value="Calsyntenin 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.200">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.60">
        <property type="entry name" value="Cadherins"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002126">
        <property type="entry name" value="Cadherin-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015919">
        <property type="entry name" value="Cadherin-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045588">
        <property type="entry name" value="CLSTN_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14139">
        <property type="entry name" value="CALSYNTENIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14139:SF4">
        <property type="entry name" value="CALSYNTENIN-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19699">
        <property type="entry name" value="CLSTN_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13385">
        <property type="entry name" value="Laminin_G_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00205">
        <property type="entry name" value="CADHERIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00112">
        <property type="entry name" value="CA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49313">
        <property type="entry name" value="Cadherin-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50268">
        <property type="entry name" value="CADHERIN_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9 21" id="KW-0106">Calcium</keyword>
    <keyword evidence="10" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="17" id="KW-0966">Cell projection</keyword>
    <keyword evidence="8" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="15" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="14 23" id="KW-0472">Membrane</keyword>
    <keyword evidence="16" id="KW-0628">Postsynaptic cell membrane</keyword>
    <keyword evidence="27" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="6 24" id="KW-0732">Signal</keyword>
    <keyword evidence="12" id="KW-0770">Synapse</keyword>
    <keyword evidence="5 23" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 23" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="24">
        <location>
            <begin position="1"/>
            <end position="28"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019182138" description="Calsyntenin-1" evidence="24">
        <location>
            <begin position="29"/>
            <end position="960"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="23">
        <location>
            <begin position="841"/>
            <end position="861"/>
        </location>
    </feature>
    <feature type="domain" description="Cadherin" evidence="25">
        <location>
            <begin position="38"/>
            <end position="164"/>
        </location>
    </feature>
    <feature type="domain" description="Cadherin" evidence="25">
        <location>
            <begin position="165"/>
            <end position="265"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="897"/>
            <end position="960"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="22">
        <location>
            <begin position="906"/>
            <end position="942"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="22">
        <location>
            <begin position="944"/>
            <end position="960"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004115"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004487"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004614"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023257"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00035006"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00035015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00069678"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS50268"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016459.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="960" mass="108046" checksum="2F6D3519B0111AD8" modified="2019-05-08" version="1">MLRPGMLCPLLTPVWLLTAGLLCGGLWAARVNKHKPWIETTYHGIVTENDNTVLLDPPLIALDKDAPLRFAESFEVTVTKEGEICGFKIHGQNIPFHAVVVDKSTGEGIIRSKEKLDCELQKDYTFTIQAYDCGKGPDGTNAKKSHKAIVHIQVNDVNEYAPVFKEKSYKATVIEGKRYDNILKVEAVDADCSPQFSQICTYEIVTPDVPFAIDKDGYIKNTEKLIYGKEHQYKLTVTAYDCGKKRAAEDVLVKISIKPTCKPAWQGWSKRIEYEPGTGSLALFPNIHLETCDEPITSIQATVELETTHIGKGCDRDTYSEKSLHRLCGASSGTAELLPPPSSTANWTVGLPTDNGHDSDQVFEFNGTQAVKIPDGVVTVSLKEPFMISVWMRHGPGTKEKETILCNSDKTDMNRHHYSLYVHNCRLVFLFRQDPSEGKTYKPAEFHWKLNQVCDKEWHHYVLSTEFPTVTLYVDGVSYDPFPVTEDYPLHPSKIETQLVVGACWQGGELHMAQFFRGNLAGLMIRSGKLENKKVIDCLYTCKEGLDLQIADGIGKGMKINVNPSQSTLTLEGDDIDRFDKAMQHVSYLNSRQFPTPGIRRLKITSNVKCFNEEACIAIPFVDGYIMVLQPEEPKISLSGINHFARSASEFESPEGVSLFPELRIISTITREVEPEGDGDEDPTVQESLVSEEIMHNLDTCEVTVLGEELNQEQESLEVDMTQLQQKGIEMSSSNLGMTITGVDTMASYEEVLHLIRYRNWHTVSLFDRKFKLVCSELNGRYVSNEFEVEVNVIHTANPVEHANHIAMQPQFVHPVHHTFVDLSGHNLANPHPFSVVPSTATIVIVVCVSFLVFMIILGVFRIRAAHQRTMHDQDTGKENEMDWDDSALTITVNPMETYEDQHSSEEEEEEEESEDGEEEDDITSAESESSEEEEGEQEEDQQNVNRQQQLEWDDSTLSY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HND4</accession>
    <name>A0A452HND4_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="21">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="14">
            <text>eicosanoyl-CoA + oxidized [electron-transfer flavoprotein] + H(+) = (2E)-eicosenoyl-CoA + reduced [electron-transfer flavoprotein]</text>
            <dbReference type="Rhea" id="RHEA:47236"/>
            <dbReference type="Rhea" id="RHEA-COMP:10685"/>
            <dbReference type="Rhea" id="RHEA-COMP:10686"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57380"/>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
            <dbReference type="ChEBI" id="CHEBI:58307"/>
            <dbReference type="ChEBI" id="CHEBI:74691"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="14">
            <dbReference type="Rhea" id="RHEA:47237"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>octadecanoyl-CoA + oxidized [electron-transfer flavoprotein] + H(+) = (2E)-octadecenoyl-CoA + reduced [electron-transfer flavoprotein]</text>
            <dbReference type="Rhea" id="RHEA:47240"/>
            <dbReference type="Rhea" id="RHEA-COMP:10685"/>
            <dbReference type="Rhea" id="RHEA-COMP:10686"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57394"/>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
            <dbReference type="ChEBI" id="CHEBI:58307"/>
            <dbReference type="ChEBI" id="CHEBI:71412"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="15">
            <dbReference type="Rhea" id="RHEA:47241"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="12">
            <text>oxidized [electron-transfer flavoprotein] + hexadecanoyl-CoA + H(+) = (2E)-hexadecenoyl-CoA + reduced [electron-transfer flavoprotein]</text>
            <dbReference type="Rhea" id="RHEA:43448"/>
            <dbReference type="Rhea" id="RHEA-COMP:10685"/>
            <dbReference type="Rhea" id="RHEA-COMP:10686"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57379"/>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
            <dbReference type="ChEBI" id="CHEBI:58307"/>
            <dbReference type="ChEBI" id="CHEBI:61526"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="12">
            <dbReference type="Rhea" id="RHEA:43449"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>tetradecanoyl-CoA + oxidized [electron-transfer flavoprotein] + H(+) = (2E)-tetradecenoyl-CoA + reduced [electron-transfer flavoprotein]</text>
            <dbReference type="Rhea" id="RHEA:47316"/>
            <dbReference type="Rhea" id="RHEA-COMP:10685"/>
            <dbReference type="Rhea" id="RHEA-COMP:10686"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57385"/>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
            <dbReference type="ChEBI" id="CHEBI:58307"/>
            <dbReference type="ChEBI" id="CHEBI:61405"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="13">
            <dbReference type="Rhea" id="RHEA:47317"/>
        </physiologicalReaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 16">
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Mitochondrion inner membrane</location>
            <topology evidence="2">Peripheral membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 16">Belongs to the acyl-CoA dehydrogenase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HND4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018820.1">
        <property type="protein sequence ID" value="ENSGAGP00000016490.1"/>
        <property type="gene ID" value="ENSGAGG00000012337.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005743">
        <property type="term" value="C:mitochondrial inner membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003995">
        <property type="term" value="F:acyl-CoA dehydrogenase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050660">
        <property type="term" value="F:flavin adenine dinucleotide binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006631">
        <property type="term" value="P:fatty acid metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.140.10:FF:000023">
        <property type="entry name" value="Acyl-CoA dehydrogenase family member 9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.110.10:FF:000006">
        <property type="entry name" value="very long-chain specific acyl-CoA dehydrogenase, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.540.10:FF:000001">
        <property type="entry name" value="Very long-chain-specific acyl-CoA dehydrogenase, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.540.10">
        <property type="entry name" value="Acyl-CoA dehydrogenase/oxidase, N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.110.10">
        <property type="entry name" value="Butyryl-CoA Dehydrogenase, subunit A, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.140.10">
        <property type="entry name" value="Butyryl-CoA Dehydrogenase, subunit A, domain 3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049448">
        <property type="entry name" value="ACAD9/ACADV-like_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006089">
        <property type="entry name" value="Acyl-CoA_DH_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006091">
        <property type="entry name" value="Acyl-CoA_Oxase/DH_mid-dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046373">
        <property type="entry name" value="Acyl-CoA_Oxase/DH_mid-dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036250">
        <property type="entry name" value="AcylCo_DH-like_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009075">
        <property type="entry name" value="AcylCo_DH/oxidase_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013786">
        <property type="entry name" value="AcylCoA_DH/ox_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037069">
        <property type="entry name" value="AcylCoA_DH/ox_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009100">
        <property type="entry name" value="AcylCoA_DH/oxidase_NM_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43884">
        <property type="entry name" value="ACYL-COA DEHYDROGENASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43884:SF9">
        <property type="entry name" value="COMPLEX I ASSEMBLY FACTOR ACAD9, MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21343">
        <property type="entry name" value="ACAD9-ACADV_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00441">
        <property type="entry name" value="Acyl-CoA_dh_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02770">
        <property type="entry name" value="Acyl-CoA_dh_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02771">
        <property type="entry name" value="Acyl-CoA_dh_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47203">
        <property type="entry name" value="Acyl-CoA dehydrogenase C-terminal domain-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56645">
        <property type="entry name" value="Acyl-CoA dehydrogenase NM domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00072">
        <property type="entry name" value="ACYL_COA_DH_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7 16" id="KW-0274">FAD</keyword>
    <keyword evidence="5 16" id="KW-0285">Flavoprotein</keyword>
    <keyword evidence="11" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="6" id="KW-0999">Mitochondrion inner membrane</keyword>
    <keyword evidence="9 16" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0809">Transit peptide</keyword>
    <feature type="domain" description="Acyl-CoA dehydrogenase/oxidase N-terminal" evidence="19">
        <location>
            <begin position="57"/>
            <end position="165"/>
        </location>
    </feature>
    <feature type="domain" description="Acyl-CoA oxidase/dehydrogenase middle" evidence="18">
        <location>
            <begin position="169"/>
            <end position="269"/>
        </location>
    </feature>
    <feature type="domain" description="Acyl-CoA dehydrogenase/oxidase C-terminal" evidence="17">
        <location>
            <begin position="282"/>
            <end position="395"/>
        </location>
    </feature>
    <feature type="domain" description="ACAD9/ACADV-like C-terminal" evidence="20">
        <location>
            <begin position="441"/>
            <end position="559"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001974"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004637"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00009347"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022630"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022792"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022827"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00047916"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00049038"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00049140"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00049224"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="RuleBase" id="RU362125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF00441"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF02770"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="Pfam" id="PF02771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="Pfam" id="PF21343"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016490.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="570" mass="62298" checksum="8EE1E4AEEF54D44E" modified="2019-05-08" version="1">MSGLSLLLRAARAAFPAGARQLRTGGFRSAYAKELFLGKMWKEEVFPYPEISNEELEEINQFVGPVEKFFSEEVDSKKIDQDAKIPPETLKGLKNLGLFGMQIPEEYGGLGLSNTMYARLGEIISLDGSVAVTLAAHQAIGLKGILIAGTDEQKAKYLPKLASGEHIAAFCLTEPGSGSDAASIQTRATMSEDGKSFLLNGSKIWISNGGLADVFTVFARTEIVDNDGAKKDKITALIVERAFGGVTNGKPEDKLGIRGSNTCEVHFENTKVPVENVIGEVGGGFKVAMNILNSGRFSMGSASAGMIKKLIEMTAEYACTRKQFNKKLSDFGLIQEKFTLMAVKAYVMESMAYLTAGMMDRPGVPDCSVEAAMVKGTNEILRMYIALTGMQYAGKILTGKIKEMKKGNVGVALEIFVNKFRDSMGRKVDLGLVGKDGVAHPSLEESSKKLEENIYYFGTTVEGLLRRFGKTIVDEQLVLKRVADVSVNLYAMTAAISRASRSISIGLRNHDHEVLLTNIFCTEAYFKNNYAMAELEKYAAENLDDSIKKVAQQVLEKRAYICSHPLERTF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HND5</accession>
    <name>A0A452HND5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Protein capicua homolog-like C-terminal tri-helical domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000018825.1">
        <property type="protein sequence ID" value="ENSGAGP00000016494.1"/>
        <property type="gene ID" value="ENSGAGG00000012212.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000977">
        <property type="term" value="F:RNA polymerase II transcription regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052412">
        <property type="entry name" value="CC-Dev_Transcription_Reg"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058606">
        <property type="entry name" value="HTH_Cic_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13059">
        <property type="entry name" value="HMG-BOX TRANSCRIPTION FACTOR BBX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13059:SF13">
        <property type="entry name" value="PROTEIN CAPICUA HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25981">
        <property type="entry name" value="HTH_Cic_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="5" id="KW-0539">Nucleus</keyword>
    <keyword evidence="1" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0804">Transcription</keyword>
    <keyword evidence="2" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="Protein capicua homolog-like C-terminal tri-helical" evidence="7">
        <location>
            <begin position="929"/>
            <end position="983"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="1"/>
            <end position="157"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="718"/>
            <end position="814"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="826"/>
            <end position="897"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="983"/>
            <end position="1048"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="1"/>
            <end position="10"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="6">
        <location>
            <begin position="11"/>
            <end position="21"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="30"/>
            <end position="40"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="6">
        <location>
            <begin position="41"/>
            <end position="52"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="141"/>
            <end position="151"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="754"/>
            <end position="766"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="804"/>
            <end position="814"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="1028"/>
            <end position="1038"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF25981"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016494.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1048" mass="105960" checksum="EBF7CB848AEE74A3" modified="2019-05-08" version="1">MTSDEERMVICEEEGDDDVIADDGFNPPDIDLKCKERVTDSESDISSGEEPDGQGFGRKLFSPVIRSPPLPPSSGAFAPCRPPAPPELDLPHGPAQPYSPFPLALGPFKAQDSRGPRPPAPPPAKRPEPATPYRRKRADSAGGAEPGAAGPSVISSPGGVLQALVLPEPGRLSAGTVLVPAPSLSSCGGAPGLARTASTVVTNVVKPVSSTPVPIASKPFPRGAGEGGPLLSPPPLEPAPPPLAVLGGAKPEAVPVGRIGLLYADKKPPPAASQAPSPHVVVGPMSKVPAAPGNMVTNLLVGQAGGTASGPAPVTVLPPQPSVQFITQNPPGAGPNGPVPLGILQPQGLLSGAPGKAGGITQVQYILPTLPQQLQVAGGKGSGAGTASIHFTLPPTSSKVLATAPHGLPILQPGPAQSVSPVQAPGPGGSAHLVQGKVLVPVAAPQVTVRSGGGAGPMAQVAPPFGVPLQNGSQPTSKIIQLTPLTPAPNSAALAPPLSPATALVGPHSQAQKVLLPSSTRITYVQSTGGHQLALGTSASPSQAAATPAATYVQAPVGPAPMALGFTAIGPSGPAIVQPLLSGQSPLLAPGQVGVSPLPSPQLPPSCSGGPVITAIYPGPPSAGPAPSPAHGLVYSVASPAPAPILPKGSSAGSQGTMGPGPMGMNGAGLGRAPFIPTTWPQNGGGAMEQGAGHGSTHMATEWGRGYGTGGWAWLHPHETGXSPLQTPRGPVSPDWRVPGPESRPEAPAPPAGPSSSSALSPAPAGSGDGSRGAGPTPSVPEGPERKEGAGGKKVKVRPPPLKKTFDSVDNRVLSEVDFEERFAELPEFKPEEVLPSPTLQSLATSPRAILGSYRKKRKNSTDLDSSTEDPISPKRKMRRRSSCSSEPNTPKSAKCEGDIFTFDKTGTEAEDVLGELEYEKVPYSSLRRTLDQRRALVMQLFQDHGFFPSAQATAAFQARYSDIFPTKVCLQLKIREVRQKIMQSATPTEQAPSTPEPGGPGSGSAAEGQPPEPVPPQDSDPGAEAGGSTSWDNSQMSPPGGAAGSSR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HND6</accession>
    <name>A0A452HND6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Oxysterol-binding protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1 5">Belongs to the OSBP family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HND6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018806.1">
        <property type="protein sequence ID" value="ENSGAGP00000016478.1"/>
        <property type="gene ID" value="ENSGAGG00000012290.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097038">
        <property type="term" value="C:perinuclear endoplasmic reticulum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015485">
        <property type="term" value="F:cholesterol binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006869">
        <property type="term" value="P:lipid transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.160.120:FF:000005">
        <property type="entry name" value="Oxysterol-binding protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.3490:FF:000003">
        <property type="entry name" value="Oxysterol-binding protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.160.120">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.3490">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037239">
        <property type="entry name" value="OSBP_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000648">
        <property type="entry name" value="Oxysterol-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018494">
        <property type="entry name" value="Oxysterol-bd_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10972">
        <property type="entry name" value="OXYSTEROL-BINDING PROTEIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10972:SF153">
        <property type="entry name" value="OXYSTEROL-BINDING PROTEIN-RELATED PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01237">
        <property type="entry name" value="Oxysterol_BP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF144000">
        <property type="entry name" value="Oxysterol-binding protein-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01013">
        <property type="entry name" value="OSBP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3 6" id="KW-0445">Lipid transport</keyword>
    <keyword evidence="4" id="KW-0446">Lipid-binding</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 6" id="KW-0813">Transport</keyword>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1"/>
            <end position="48"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="284"/>
            <end position="311"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="372"/>
            <end position="406"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="284"/>
            <end position="300"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="8">
        <location>
            <begin position="302"/>
            <end position="311"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00008842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023121"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="RuleBase" id="RU003844"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="RuleBase" id="RU003845"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016478.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="443" mass="50782" checksum="22E187560019D344" modified="2019-05-08" version="1">LNSEQVKSFDSDNSSGDFSETNHKVAEMLDLDPPQSNRTGKHNGARQLHENGIKKHRTSLPAPMFSRSDFSVWSILKKCIGLELSKITMPIVFNEPLSFLQRITEYMEHIYLIHKASSHANPLERMQAVAAFAVSAVASQWERTGKPFNPLLGETYELIREDLGFRFISEQVSHHPPVSAFYSEGLNKDFVFHGSIYPKLKFWGKSVEAEPRGTITLELLKHNEAYTWTNPTCCVHNIIIGKLWIEQYGTIEIVNHSRKKLSVIYGKWTECLWCVNPATYDTSKKNEKRGGDQKKMKPSEEAGNDEADDMPEIQETVQVIQGSKLLWRINSRPPNSSQMYNFTSFAVSLNELETGMEAVLAPTDCRLRPDIRGMENGELDLASKEKERLEEKQRAARKERAKDEVEWRTRWFHQGNNPYTGTSDWLYSGGYFDRNFSDCPDIY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HND7</accession>
    <name>A0A452HND7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Calsyntenin-1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="26 27">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="27" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="26" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="26" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell projection</location>
            <location evidence="2">Neuron projection</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Golgi apparatus membrane</location>
            <topology evidence="3">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="18">Postsynaptic cell membrane</location>
            <topology evidence="18">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="19">Belongs to the calsyntenin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HND7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018793.1">
        <property type="protein sequence ID" value="ENSGAGP00000016466.1"/>
        <property type="gene ID" value="ENSGAGG00000012316.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009986">
        <property type="term" value="C:cell surface"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000139">
        <property type="term" value="C:Golgi membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043005">
        <property type="term" value="C:neuron projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007156">
        <property type="term" value="P:homophilic cell adhesion via plasma membrane adhesion molecules"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051965">
        <property type="term" value="P:positive regulation of synapse assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050806">
        <property type="term" value="P:positive regulation of synaptic transmission"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd11304">
        <property type="entry name" value="Cadherin_repeat"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.200:FF:000036">
        <property type="entry name" value="Calsyntenin 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.60:FF:000025">
        <property type="entry name" value="Calsyntenin 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.60:FF:000062">
        <property type="entry name" value="Calsyntenin 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.200">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.60">
        <property type="entry name" value="Cadherins"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002126">
        <property type="entry name" value="Cadherin-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015919">
        <property type="entry name" value="Cadherin-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045588">
        <property type="entry name" value="CLSTN_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14139">
        <property type="entry name" value="CALSYNTENIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14139:SF4">
        <property type="entry name" value="CALSYNTENIN-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19699">
        <property type="entry name" value="CLSTN_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00205">
        <property type="entry name" value="CADHERIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00112">
        <property type="entry name" value="CA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49313">
        <property type="entry name" value="Cadherin-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50268">
        <property type="entry name" value="CADHERIN_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9 21" id="KW-0106">Calcium</keyword>
    <keyword evidence="10" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="17" id="KW-0966">Cell projection</keyword>
    <keyword evidence="8" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="15" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="14 23" id="KW-0472">Membrane</keyword>
    <keyword evidence="16" id="KW-0628">Postsynaptic cell membrane</keyword>
    <keyword evidence="27" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="6 24" id="KW-0732">Signal</keyword>
    <keyword evidence="12" id="KW-0770">Synapse</keyword>
    <keyword evidence="5 23" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 23" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="24">
        <location>
            <begin position="1"/>
            <end position="28"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019411429" description="Calsyntenin-1" evidence="24">
        <location>
            <begin position="29"/>
            <end position="967"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="23">
        <location>
            <begin position="848"/>
            <end position="868"/>
        </location>
    </feature>
    <feature type="domain" description="Cadherin" evidence="25">
        <location>
            <begin position="38"/>
            <end position="154"/>
        </location>
    </feature>
    <feature type="domain" description="Cadherin" evidence="25">
        <location>
            <begin position="155"/>
            <end position="255"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="904"/>
            <end position="967"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="22">
        <location>
            <begin position="913"/>
            <end position="949"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="22">
        <location>
            <begin position="951"/>
            <end position="967"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004115"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004487"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004614"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023257"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00035006"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00035015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00069678"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS50268"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016466.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="967" mass="108747" checksum="3A1C94D1E642DF8D" modified="2019-05-08" version="1">MLRPGMLCPLLTPVWLLTAGLLCGGLWAARVNKHKPWIETTYHGIVTENDNTVLLDPPLIALDKDAPLRFAGEICGFKIHGQNIPFHAVVVDKSTGEGIIRSKEKLDCELQKDYTFTIQAYDCGKGPDGTNAKKSHKAIVHIQVNDVNEYAPVFKEKSYKATVIEGKRYDNILKVEAVDADCSPQFSQICTYEIVTPDVPFAIDKDGYIKNTEKLIYGKEHQYKLTVTAYDCGKKRAAEDVLVKISIKPTCKPAWQGWSKRIEYEPGTGSLALFPNIHLETCDEPITSIQATVELETTHIGKGCDRDTYSEKSLHRLCGASSGTAELLPPPSSTANWTVGLPTDNGHDSDQVFEFNGTQAVKIPDGVVTVSLKEPFMISVWMRHGPGTKEKETILCNSDKTDMNRHHYSLYVHNCRLVFLFRQDPSEGKTYKPAEFHWKLNQVCDKEWHHYVLSTEFPTVTLYVDGVSYDPFPVTEDYPLHPSKIETQLVVGACWQEYTGNENDNETVPETSAGGELHMAQFFRGNLAGLMIRSGKLENKKVIDCLYTCKEGLDLQIADGIGKGMKINVNPSQSTLTLEGDDIDRFDKAMQHVSYLNSRQFPTPGIRRLKITSNVKCFNEEACIAIPFVDGYIMVLQPEEPKISLSGINHFARSASEFESPEGVSLFPELRIISTITREVEPEGDGDEDPTVQESLVSEEIMHNLDTCEVTVLGEELNQEQESLEVDMTQLQQKGIEMSSSNLGMTITGVDTMASYEEVLHLIRYRNWHTVSLFDRKFKLVCSELNGRYVSNEFEVEVNVIHTANPVEHANHIAMQPQFVHPVHHTFVDLSGHNLANPHPFSVVPSTATIVIVVCVSFLVFMIILGVFRIRAAHQRTMHDQDTGKENEMDWDDSALTITVNPMETYEDQHSSEEEEEEEESEDGEEEDDITSAESESSEEEEGEQEEDQQNVNRQQQLEWDDSTLSY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HND8</accession>
    <name>A0A452HND8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Nuclear pore complex protein Nup214</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="13">214 kDa nucleoporin</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="12">Nucleoporin Nup214</fullName>
        </alternativeName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Part of the nuclear pore complex. Has a critical role in nucleocytoplasmic transport. May serve as a docking site in the receptor-mediated import of substrates across the nuclear pore complex.</text>
    </comment>
    <comment type="subunit">
        <text evidence="10">Homodimer. Part of the nuclear pore complex (NPC). Interacts with NUP88. Interacts with ZFP36; this interaction increases upon lipopolysaccharide (LPS) stimulation. Interacts with DDX19. Interacts with XPO1. Interacts with XPO5.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
            <location evidence="1">Nuclear pore complex</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018831.1">
        <property type="protein sequence ID" value="ENSGAGP00000016499.1"/>
        <property type="gene ID" value="ENSGAGG00000012340.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005643">
        <property type="term" value="C:nuclear pore"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008139">
        <property type="term" value="F:nuclear localization sequence binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017056">
        <property type="term" value="F:structural constituent of nuclear pore"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006406">
        <property type="term" value="P:mRNA export from nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006606">
        <property type="term" value="P:protein import into nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:000142">
        <property type="entry name" value="Nuclear pore complex protein Nup214"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026054">
        <property type="entry name" value="Nucleoporin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039462">
        <property type="entry name" value="Nup159/Nup146_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041553">
        <property type="entry name" value="Nup214_FG"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001680">
        <property type="entry name" value="WD40_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23193">
        <property type="entry name" value="NUCLEAR PORE COMPLEX PROTEIN NUP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23193:SF21">
        <property type="entry name" value="NUCLEAR PORE COMPLEX PROTEIN NUP214"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16755">
        <property type="entry name" value="Beta-prop_NUP159_NUP214"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18617">
        <property type="entry name" value="Nup214_FG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00320">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF117289">
        <property type="entry name" value="Nucleoporin domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0509">mRNA transport</keyword>
    <keyword evidence="7" id="KW-0906">Nuclear pore complex</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="5" id="KW-0653">Protein transport</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0811">Translocation</keyword>
    <keyword evidence="2" id="KW-0813">Transport</keyword>
    <feature type="domain" description="Nucleoporin Nup159/Nup146 N-terminal" evidence="15">
        <location>
            <begin position="34"/>
            <end position="376"/>
        </location>
    </feature>
    <feature type="domain" description="Nuclear pore complex protein Nup214 phenylalanine-glycine (FG)" evidence="16">
        <location>
            <begin position="1878"/>
            <end position="1938"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="545"/>
            <end position="606"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1257"/>
            <end position="1280"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1445"/>
            <end position="1524"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1599"/>
            <end position="1655"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1718"/>
            <end position="1753"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1846"/>
            <end position="1865"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="545"/>
            <end position="566"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="574"/>
            <end position="584"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="1479"/>
            <end position="1495"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="1599"/>
            <end position="1622"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="1639"/>
            <end position="1651"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="1720"/>
            <end position="1731"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="1737"/>
            <end position="1748"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004567"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022816"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023010"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023132"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00055090"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00063892"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00068360"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00077390"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00083901"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="Pfam" id="PF16755"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF18617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016499.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2052" mass="210417" checksum="293F4FC8D1AF2A02" modified="2019-05-08" version="1">MGDDMDVVPEREMKDFQFRALKKVRIFDAPEELPKERSSLLAISNKYGLVFVGGATGLQIFHTKDLLMQNQAGEDRNKIVENVQSILVPMKFPVHHLALSCDNLTLSVCMTSSEFGSFIGFFDVRTFSNQAKQQKRPFAYHKLAKDSGSMAMVCDLKWNPASPSMVAVCLSDGSISVLQVTESVKVCATLPSSVAVTSVCWSPKGKQLAVGKQNGTVVQYLPSLQEKKVIPCPPFYDSDNPVKVLDVLWVGTYVFTIVYAAADGTLETSPEVVMAVLPKKEEKRPEKFMNFMEPCYGSCTERQHHYFLNYVEEWALVLAASAASTEVSILSRQADQNNWESWVLEDSSRAELPVTDKNDDTLPVGVAIDYTSNMPILIRSTATTPTTPTAPSKFETAPVVTPATASPLAPASSASGLLSAGGGAATFSFTSSAVKTSGSMSGDPPPYPFASDVSKQTLGSSLTGAAAFSFSSVSSKASMTPSPAAGPMAATANSFSFGATNFKPATNSAPVPPLSTTPAGLKPCITPTASTVKVNLSEKFTAIDTSASAGSSSHNSSTMFSFSTTSKPIPGGPLSQSTPLSASPATPVKASTPVSATMARPAQNVPATSNLLKTARITPPATKSISIQGKSALSSTGSEKQVHQWKDSDPVIAGIKEEIAHFQKEMEELKARTSNACFQVGTAEEMKQLRTDSHDLQTFLLEIKETTESLHGDISILKTTLLEGFAGVEDAREQNERNHNSGYLHLLYKKPLDPKSEAQLQEIRRLHQYVKFAVQDVNDVLDLEWDRYLEQKKTQKRLIVPERETLFNTLANNREIINQQRKRLNHLVDSLQQLRLYNQTSQWSMPSDTSSLSSSQSFDSDLESLRSALVKTTLESYAKSLPKAPAKLSPVKQAQLRNFLAKRKTPPVRSTAPANLSRSAFLSLRYYEDLDEVSSTSSISQSLENDDSQIVEQEEEAAPVQVPRHAPVVRTPSIQPGLMAQSPPFGKPHLTMGTLVPTSSSRIVPQGADSTMLATKTVKHGAPAPTVTVPAPQAAAAAALRRQMASQTPAVSTSLTESTLKNVPQVVNVRELKNNASAPAVSAVIGSSVPQSAAQAVHQVLATVATNQAKQGSLTSSLKAQSSPISASIPCTITTTAGQIPAGSKISGQGTAKAVAAVTSPTMTSTPVAAAQFNQTFSFSSLGTGFNFGAATSTASTPPSLSASQVVTAPAKESSQSDAFSFDGGTKFGTVPESSFTFGSIKPANVSGVSMTSGSSVDATQMNSKPAPAATSVPSTTSGKLEVLPSKPGDNLFQSNMAGETLGSFSGLRVGQADDNTKVNVKAPSTNIAGGQPTKTSTIPSGFTFNSPLQLGKPVEATSTLVMATNTASTSVSTNTTGSIFGNVQLTSTGSSGVFSLGGSSGGGKPAFSFGLPQANSITTSTSVPATLPTSASLSFGSLLSSAPTAVSPATSSNGAGDVKLPPLSEKPPEGEILTTVSAPLASQPQTTQPQLQLSDSVKDEPALSQSIAGDADVTGPSTSLTPSTDTISTAICATESEAEAVPPVSTSAPLDQNISVTTAAADVVIAPSAEITSTLATTKDIDTSVQSPAAGTSVFGTVSSSSSVFSQPPNSSSSSSAFSHPVGDTTTTPSTPVFGPQTAATTASSPFGQHTGSTISTTAATTQVASSGFGASAFGTAAAGGFGQPAFGQAPAFGQPASSSSSGFTFTQTGFGTMPVFGQPSSSTATSSGGNVFGAPSSTNSPSSFSFGQPSASTGGGLFGQSSAPVFGQSAGFGQTGSVFGNNATATTTTSSSGFSFCQASGFGSSNTGSVFGQAANTGGAVFGQQSSTSSGGLFGSGSGGRSGGFFSGLGGKPSQDAANKNPFSSATGGFGSAASQNTSSLFGNNGAKTFRFGNSSYGEQKPTGTFSSGGGTVAAQGFGFSSPTKAGGFGAAPVFGSPPTFGGSPGFGGVPAFGSAPAFTSPLGSTGGKVFGEGTAAANAGGFGFGGTSNNAASFGTLANQNAPTFGLLSQQTTGFGTQSSGFSGFGSSGGGFGFGSSNSSTSGFSGWRS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNG0</accession>
    <name>A0A452HNG0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Nuclear pore complex protein Nup214</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="13">214 kDa nucleoporin</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="12">Nucleoporin Nup214</fullName>
        </alternativeName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Part of the nuclear pore complex. Has a critical role in nucleocytoplasmic transport. May serve as a docking site in the receptor-mediated import of substrates across the nuclear pore complex.</text>
    </comment>
    <comment type="subunit">
        <text evidence="10">Homodimer. Part of the nuclear pore complex (NPC). Interacts with NUP88. Interacts with ZFP36; this interaction increases upon lipopolysaccharide (LPS) stimulation. Interacts with DDX19. Interacts with XPO1. Interacts with XPO5.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
            <location evidence="1">Nuclear pore complex</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018835.1">
        <property type="protein sequence ID" value="ENSGAGP00000016503.1"/>
        <property type="gene ID" value="ENSGAGG00000012340.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005643">
        <property type="term" value="C:nuclear pore"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008139">
        <property type="term" value="F:nuclear localization sequence binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017056">
        <property type="term" value="F:structural constituent of nuclear pore"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006406">
        <property type="term" value="P:mRNA export from nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006606">
        <property type="term" value="P:protein import into nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:000142">
        <property type="entry name" value="Nuclear pore complex protein Nup214"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026054">
        <property type="entry name" value="Nucleoporin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039462">
        <property type="entry name" value="Nup159/Nup146_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041553">
        <property type="entry name" value="Nup214_FG"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23193">
        <property type="entry name" value="NUCLEAR PORE COMPLEX PROTEIN NUP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23193:SF21">
        <property type="entry name" value="NUCLEAR PORE COMPLEX PROTEIN NUP214"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16755">
        <property type="entry name" value="Beta-prop_NUP159_NUP214"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18617">
        <property type="entry name" value="Nup214_FG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF117289">
        <property type="entry name" value="Nucleoporin domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0509">mRNA transport</keyword>
    <keyword evidence="7" id="KW-0906">Nuclear pore complex</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="5" id="KW-0653">Protein transport</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0811">Translocation</keyword>
    <keyword evidence="2" id="KW-0813">Transport</keyword>
    <feature type="domain" description="Nucleoporin Nup159/Nup146 N-terminal" evidence="15">
        <location>
            <begin position="79"/>
            <end position="280"/>
        </location>
    </feature>
    <feature type="domain" description="Nuclear pore complex protein Nup214 phenylalanine-glycine (FG)" evidence="16">
        <location>
            <begin position="1813"/>
            <end position="1873"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="305"/>
            <end position="336"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="480"/>
            <end position="524"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1192"/>
            <end position="1215"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1380"/>
            <end position="1460"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1534"/>
            <end position="1590"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1653"/>
            <end position="1688"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1781"/>
            <end position="1800"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="312"/>
            <end position="336"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="480"/>
            <end position="501"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="509"/>
            <end position="519"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="1414"/>
            <end position="1430"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="1451"/>
            <end position="1460"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="1534"/>
            <end position="1557"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="1574"/>
            <end position="1586"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="1655"/>
            <end position="1666"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="1672"/>
            <end position="1683"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004567"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022816"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023010"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023132"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00055090"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00063892"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00068360"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00077390"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00083901"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="Pfam" id="PF16755"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF18617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016503.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1987" mass="203233" checksum="636D644627F5F2FA" modified="2019-05-08" version="1">MGDDMDVVPEREMKDFQFRALKKVRIFDAPEELPKERSSLLAISNKYGLVFVGGATGLQIFHTKDLLMQNQAGEDRNKIVCWSPKGKQLAVGKQNGTVVQYLPSLQEKKVIPCPPFYDSDNPVKVLDVLWVGTYVFTIVYAAADGTLETSPEVVMAVLPKKEEKRPEKFMNFMEPCYGSCTERQHHYFLNYVEEWALVLAASAASTEVSILSRQADQNNWESWVLEDSSRAELPVTDKNDDTLPVGVAIDYTSNMPILISDEKTLPPAPVLILLSTDGVLCPFYMINQNPGVKSLIVTPEPLSIEGERAPKSAGSTATTPTTPTAPSKFETAPVVTPATASPLAPASSASGLLSAGGGAATFSFTSSAVKTSGSMSGDPPPYPFASDVSKQTLGSSLTGAAAFSFSSVSSKASMTPSPAAGPMAATANSFSFGATNFKPATNSAPVPPLSTTPAGLKPCITPTASTVKVNLSEKFTAIDTSASAGSSSHNSSTMFSFSTTSKPIPGGPLSQSTPLSASPATPVKASTPVSATMARPAQNVPATSNLLKTARITPPATKSISIQGKSALSSTGSEKQVHQWKDSDPVIAGIKEEIAHFQKEMEELKARTSNACFQVGTAEEMKQLRTDSHDLQTFLLEIKETTESLHGDISILKTTLLEGFAGVEDAREQNERNHNSGYLHLLYKKPLDPKSEAQLQEIRRLHQYVKFAVQDVNDVLDLEWDRYLEQKKTQKRLIVPERETLFNTLANNREIINQQRKRLNHLVDSLQQLRLYNQTSQWSMPSDTSSLSSSQSFDSDLESLRSALVKTTLESYAKSLPKAPAKLSPVKQAQLRNFLAKRKTPPVRSTAPANLSRSAFLSLRYYEDLDEVSSTSSISQSLENDDSQIVEQEEEAAPVQVPRHAPVVRTPSIQPGLMAQSPPFGKPHLTMGTLVPTSSSRIVPQGADSTMLATKTVKHGAPAPTVTVPAPQAAAAAALRRQMASQTPAVSTSLTESTLKNVPQVVNVRELKNNASAPAVSAVIGSSVPQSAAQAVHQVLATVATNQAKQGSLTSSLKAQSSPISASIPCTITTTAGQIPAGSKISGQGTAKAVAAVTSPTMTSTPVAAAQFNQTFSFSSLGTGFNFGAATSTASTPPSLSASQVVTAPAKESSQSDAFSFDGGTKFGTVPESSFTFGSIKPANVSGVSMTSGSSVDATQMNSKPAPAATSVPSTTSGKLEVLPSKPGDNLFQSNMAGETLGSFSGLRVGQADDNTKVNVKAPSTNIAGGQPTKTSTIPSGFTFNSPLQLGKPVEATSTLVMATNTASTSVSTNTTGSIFGNVQLTSTGSSGVFSLGGSSGGGKPAFSFGLPQANSITTSTSVPATLPTSASLSFGSLLSSAPTAVSPATSSNGAGDVKLPPLSEKPPEGEILTTVSAPLASQPQTTQPQLQLSDSVKDEPALSQSIAGDADVTGPSTSLTPSTDTISTAICATESEAEAVPPVSTSAPLDQNISVTTAAADVVIAPSAEITSTLATTKDIDTSVQSPAAGTSVFGTVSSSSSVFSQPPNSSSSSSAFSHPVGDTTTTPSTPVFGPQTAATTASSPFGQHTGSTISTTAATTQVASSGFGASAFGTAAAGGFGQPAFGQAPAFGQPASSSSSGFTFTQTGFGTMPVFGQPSSSTATSSGGNVFGAPSSTNSPSSFSFGQPSASTGGGLFGQSSAPVFGQSAGFGQTGSVFGNNATATTTTSSSGFSFCQASGFGSSNTGSVFGQAANTGGAVFGQQSSTSSGGLFGSGSGGRSGGFFSGLGGKPSQDAANKNPFSSATGGFGSAASQNTSSLFGNNGAKTFRFGNSSYGEQKPTGTFSSGGGTVAAQGFGFSSPTKAGGFGAAPVFGSPPTFGGSPGFGGVPAFGSAPAFTSPLGSTGGKVFGEGTAAANAGGFGFGGTSNNAASFGTLANQNAPTFGLLSQQTTGFGTQSSGFSGFGSSGGGFGFGSSNSSTSGFSGWRS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNG3</accession>
    <name>A0A452HNG3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Attractin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="27 28">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="28" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="27" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="27" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="16">Involved in the initial immune cell clustering during inflammatory response and may regulate chemotactic activity of chemokines. May play a role in melanocortin signaling pathways that regulate energy homeostasis and hair color. Low-affinity receptor for agouti. Has a critical role in normal myelination in the central nervous system.</text>
    </comment>
    <comment type="subunit">
        <text evidence="17">Monomer and homotrimer.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="19">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018861.1">
        <property type="protein sequence ID" value="ENSGAGP00000016526.1"/>
        <property type="gene ID" value="ENSGAGG00000011749.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030246">
        <property type="term" value="F:carbohydrate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006954">
        <property type="term" value="P:inflammatory response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00041">
        <property type="entry name" value="CUB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00055">
        <property type="entry name" value="EGF_Lam"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.120.10.80:FF:000031">
        <property type="entry name" value="attractin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.120.10.80:FF:000034">
        <property type="entry name" value="Attractin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000079">
        <property type="entry name" value="Attractin like 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000164">
        <property type="entry name" value="Attractin like 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.290:FF:000008">
        <property type="entry name" value="Attractin like 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.120.10.80">
        <property type="entry name" value="Kelch-type beta propeller"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.100.10">
        <property type="entry name" value="Mannose-Binding Protein A, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.290">
        <property type="entry name" value="Spermadhesin, CUB domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056737">
        <property type="entry name" value="Beta-prop_ATRN-MKLN-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001304">
        <property type="entry name" value="C-type_lectin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016186">
        <property type="entry name" value="C-type_lectin-like/link_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016187">
        <property type="entry name" value="CTDL_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000859">
        <property type="entry name" value="CUB_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056732">
        <property type="entry name" value="GBD_ATRN"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015915">
        <property type="entry name" value="Kelch-typ_b-propeller"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002049">
        <property type="entry name" value="LE_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056863">
        <property type="entry name" value="LMN_ATRN_NET-like_EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051568">
        <property type="entry name" value="LZTR1/Attractin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002165">
        <property type="entry name" value="Plexin_repeat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016201">
        <property type="entry name" value="PSI"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035914">
        <property type="entry name" value="Sperma_CUB_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46376:SF3">
        <property type="entry name" value="ATTRACTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46376">
        <property type="entry name" value="LEUCINE-ZIPPER-LIKE TRANSCRIPTIONAL REGULATOR 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24981">
        <property type="entry name" value="Beta-prop_ATRN-LZTR1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00431">
        <property type="entry name" value="CUB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24973">
        <property type="entry name" value="EGF_LMN_ATRN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24972">
        <property type="entry name" value="GBD_ATRN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01437">
        <property type="entry name" value="PSI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00034">
        <property type="entry name" value="CLECT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00042">
        <property type="entry name" value="CUB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00180">
        <property type="entry name" value="EGF_Lam"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00423">
        <property type="entry name" value="PSI"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56436">
        <property type="entry name" value="C-type lectin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF117281">
        <property type="entry name" value="Kelch motif"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49854">
        <property type="entry name" value="Spermadhesin, CUB domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50041">
        <property type="entry name" value="C_TYPE_LECTIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01180">
        <property type="entry name" value="CUB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01248">
        <property type="entry name" value="EGF_LAM_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50027">
        <property type="entry name" value="EGF_LAM_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11 19" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="4 19" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="14" id="KW-0395">Inflammatory response</keyword>
    <keyword evidence="2" id="KW-0880">Kelch repeat</keyword>
    <keyword evidence="15 20" id="KW-0424">Laminin EGF-like domain</keyword>
    <keyword evidence="7" id="KW-0430">Lectin</keyword>
    <keyword evidence="10" id="KW-0472">Membrane</keyword>
    <keyword evidence="12" id="KW-0675">Receptor</keyword>
    <keyword evidence="28" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="6 22" id="KW-0732">Signal</keyword>
    <keyword evidence="5" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="22">
        <location>
            <begin position="1"/>
            <end position="48"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019585376" description="Attractin" evidence="22">
        <location>
            <begin position="49"/>
            <end position="1296"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="24">
        <location>
            <begin position="53"/>
            <end position="86"/>
        </location>
    </feature>
    <feature type="domain" description="CUB" evidence="23">
        <location>
            <begin position="88"/>
            <end position="204"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="24">
        <location>
            <begin position="202"/>
            <end position="239"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="26">
        <location>
            <begin position="732"/>
            <end position="852"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="25">
        <location>
            <begin position="992"/>
            <end position="1037"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="21">
        <location>
            <begin position="1"/>
            <end position="24"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="19">
        <location>
            <begin position="57"/>
            <end position="67"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="19">
        <location>
            <begin position="76"/>
            <end position="85"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="19">
        <location>
            <begin position="206"/>
            <end position="216"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="19">
        <location>
            <begin position="210"/>
            <end position="227"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="19">
        <location>
            <begin position="229"/>
            <end position="238"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="1009"/>
            <end position="1018"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="1021"/>
            <end position="1035"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022441"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022734"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023198"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023292"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00058400"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00065783"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00074596"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00460"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS01180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS50027"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="26">
        <source>
            <dbReference type="PROSITE" id="PS50041"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016526.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="28">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1296" mass="144301" checksum="DAE146EB9D346385" modified="2019-05-08" version="1">MERSSRRQPAAGRLGSQGGRPQRFGGRPPAQLLLVLLMLLVPGPEAAAAAAVEGKECDKPCANGGRCQPGSGQCECQAGWVGDQCQHCGGRFRLTGSSGYVTDGPGHYKYKTKCTWLIEGKPNTILRLRFNHFATECSWDHLYVYDGDSVYAPLVAAFSGLIVPEKDGNETIPEVVATSGYVLLHFFSDAAYNLTGFNITYNFDMCPNNCSGRGECKHSNGSNTVKCECSEHWKGEACDISYCTDDCGYPARGQCNMNETKGCLCNSGWQGPGCAVPVPANQSFWTREEYSSPKLPRASHKAVVNGNVMWVVGGYMFNHSDSHMILAYDLVSEEWLPLNSTVNSVEIRYGHSLALHENKIYMYGGKIDAMGNVTNQLWVFHIHNESWVHVIPKTKEQYAVVGHSAHIVNLGDGSAVMLVIFGHCPLYGYISNVQEYNLATNTWNILQTSGALVQGGYGHSSVYDPNTQSIYIHGGYKAFSANKYRLADDLYKYEVDTRMWTILKDSRFFRYLHTAVIMSGTMLVFGGNTHNDTSMSHGAKCFSSDFMAYDIACDHWSVLPRPDLHHDVNRFGHSAVLSNSTVYIFGGFNSLLLSDILRYTPESCEAYDNETSCLRAGPGVRCVWNTTFSLCVPWERATEASSDNCYCYSCTANTNNCQWCTDQCILKNNNCTEDQVPITEYDKCPKDNPAYYCNKKTSCKSCAMDQNCQWEPRNQECIALPENICGANWHLVGNSCLKINSTKESYDSGKLACTTQNALLASLTTQKKVEFVLKELQRMQSLAKTLTPWVGLRKINVSYWCWEDMSPFTNTLVQWLPGEPSGAGFCGYLAEPYQRGLKAATCIDHVNGSICERLANHSAKQCRTPCALRTACGECTSGSSECMWCSNMKQCVDSNAYVASFPYGQCMEWYTMTANCSGYCTCAHCLEQPGCGWCTDPSNTGKGKCMEGSYRGPVKMPSPPATGKYYLEPILNVSMCPAENKYNWSFIQCPACQCNGHSKCVNESVCEKCENLTTGKHCETCISGYYGDPTNGGTCQSCKCNGHASVCNTNTGKCFCTTKGIKGDECQLCEVENRYQGNPLKGTCYYTLLIDYQFTFSLSQEDDRYYTAINFVATPEEQNRDLDMFINASKNFNLNITWSTTGTQAGEEIPVVSKSNIKEYKDSFSNEKFDFRSNPNITFFVYVSNFTWPIKIQIAFSQHSNFMDLVQFFVTFFRGVLQVAFLTVLHFRYTCASQALDWRFPVSSICSTYACALYNFVPRPRAYRAARANHPQFLPSNLCLRWSSSMTALCVPQCVL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNG9</accession>
    <name>A0A452HNG9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="9">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the TBP family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HNG9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018862.1">
        <property type="protein sequence ID" value="ENSGAGP00000016527.1"/>
        <property type="gene ID" value="ENSGAGG00000012359.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032991">
        <property type="term" value="C:protein-containing complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006352">
        <property type="term" value="P:DNA-templated transcription initiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd04516">
        <property type="entry name" value="TBP_eukaryotes"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.310.10:FF:000005">
        <property type="entry name" value="TATA box-binding protein-like 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.310.10:FF:000002">
        <property type="entry name" value="TATA-box-binding protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.310.10">
        <property type="entry name" value="TATA-Binding Protein"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000814">
        <property type="entry name" value="TBP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR030491">
        <property type="entry name" value="TBP_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012295">
        <property type="entry name" value="TBP_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033710">
        <property type="entry name" value="TBP_eukaryotic"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10126">
        <property type="entry name" value="TATA-BOX BINDING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00352">
        <property type="entry name" value="TBP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00686">
        <property type="entry name" value="TIFACTORIID"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55945">
        <property type="entry name" value="TATA-box binding protein-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00351">
        <property type="entry name" value="TFIID"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0804">Transcription</keyword>
    <keyword evidence="4" id="KW-0805">Transcription regulation</keyword>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1"/>
            <end position="33"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005560"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016527.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="278" mass="30638" checksum="960660A245B0ECCF" modified="2019-05-08" version="1">RGLAPSSPPTTLGPATREHRAALSPRSSLPIFLPPPLPALLRPEATPVPSLSSYGHEVQREQYGSQLTEQDSGFLNDSNLSQLSYADATQPSPETSGTCPPLTPMTPITPMTPVSESSGIVPQLQRAKHQNGKRRFAAVIMRIREPRTTALIFSSGKMVCTGAKSEEQSRLAARKYARVVQKLGFPAKFLDFKIQNMVGSCDVRFPIRLEGLVLTHQQFSSYEPELFPGLIYRMVKPRIVLLIFVSGKVVLTGAKERSEIYEAFENIYPILKGFKKAS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HNH0</accession>
    <name>A0A452HNH0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Beta-hexosaminidase</fullName>
            <ecNumber evidence="20">3.2.1.52</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="26 27">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="27" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="26" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="26" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="16">Hydrolyzes the non-reducing end N-acetyl-D-hexosamine and/or sulfated N-acetyl-D-hexosamine of glycoconjugates, such as the oligosaccharide moieties from proteins and neutral glycolipids, or from certain mucopolysaccharides. The isozyme B does not hydrolyze each of these substrates, however hydrolyzes efficiently neutral oligosaccharide. Only the isozyme A is responsible for the degradation of GM2 gangliosides in the presence of GM2A. During fertilization is responsible, at least in part, for the zona block to polyspermy. Present in the cortical granules of non-activated oocytes, is exocytosed during the cortical reaction in response to oocyte activation and inactivates the sperm galactosyltransferase-binding site, accounting for the block in sperm binding to the zona pellucida.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>Hydrolysis of terminal non-reducing N-acetyl-D-hexosamine residues in N-acetyl-beta-D-hexosaminides.</text>
            <dbReference type="EC" id="3.2.1.52"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>N-acetyl-beta-D-6-sulfogalactosaminyl-(1-&gt;4)-alpha-L-iduronyl-(1-&gt;3)-N-acetyl-D-6-sulfogalactosamine + H2O = alpha-L-iduronyl-(1-&gt;3)-N-acetyl-D-6-sulfogalactosamine + N-acetyl-D-6-sulfogalactosamine</text>
            <dbReference type="Rhea" id="RHEA:64384"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:152567"/>
            <dbReference type="ChEBI" id="CHEBI:152568"/>
            <dbReference type="ChEBI" id="CHEBI:153064"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="19">
            <dbReference type="Rhea" id="RHEA:64385"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>N-acetyl-beta-D-galactosaminyl-(1-&gt;4)-beta-D-3-sulfogalactosyl-(1-&gt;4)-beta-D-glucosyl-(1&lt;-&gt;1')-ceramide + H2O = a beta-D-3-sulfogalactosyl-(1-&gt;4)-beta-D-glucosyl-(1&lt;-&gt;1')-ceramide + N-acetyl-beta-D-galactosamine</text>
            <dbReference type="Rhea" id="RHEA:48276"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:28497"/>
            <dbReference type="ChEBI" id="CHEBI:90163"/>
            <dbReference type="ChEBI" id="CHEBI:90164"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="18">
            <dbReference type="Rhea" id="RHEA:48277"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="14">
            <text>a ganglioside GM2 (d18:1(4E)) + H2O = a ganglioside GM3 (d18:1(4E)) + N-acetyl-beta-D-galactosamine</text>
            <dbReference type="Rhea" id="RHEA:47940"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:28497"/>
            <dbReference type="ChEBI" id="CHEBI:60065"/>
            <dbReference type="ChEBI" id="CHEBI:71502"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="14">
            <dbReference type="Rhea" id="RHEA:47941"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>a ganglioside GM2 + H2O = a ganglioside GM3 + N-acetyl-beta-D-galactosamine</text>
            <dbReference type="Rhea" id="RHEA:47968"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:28497"/>
            <dbReference type="ChEBI" id="CHEBI:79210"/>
            <dbReference type="ChEBI" id="CHEBI:79218"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="15">
            <dbReference type="Rhea" id="RHEA:47969"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="12">
            <text>beta-D-GalNAc-(1-&gt;4)-alpha-L-IdoA-(1-&gt;3)-beta-D-GalNAc-4-sulfate-(1-&gt;4)-alpha-L-IdoA-(1-&gt;3)-D-GalNAc-4-sulfate + H2O = alpha-L-IdoA-(1-&gt;3)-beta-D-GalNAc-4-sulfate-(1-&gt;4)-alpha-L-IdoA-(1-&gt;3)-D-GalNAc-4-sulfate + N-acetyl-D-galactosamine</text>
            <dbReference type="Rhea" id="RHEA:64372"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:28037"/>
            <dbReference type="ChEBI" id="CHEBI:152565"/>
            <dbReference type="ChEBI" id="CHEBI:152566"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="12">
            <dbReference type="Rhea" id="RHEA:64373"/>
        </physiologicalReaction>
    </comment>
    <comment type="subunit">
        <text evidence="17">There are 3 forms of beta-hexosaminidase: hexosaminidase A is a heterodimer composed of one subunit alpha and one subunit beta (chain A and B); hexosaminidase B is a homodimer of two beta subunits (two chains A and B); hexosaminidase S is a homodimer of two alpha subunits. The composition of the dimer (isozyme A versus isozyme S) has a significant effect on the substrate specificity of the alpha subunit active site.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="13">Cytoplasmic vesicle</location>
            <location evidence="13">Secretory vesicle</location>
            <location evidence="13">Cortical granule</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Lysosome</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 20">Belongs to the glycosyl hydrolase 20 family.</text>
    </comment>
    <dbReference type="EC" id="3.2.1.52" evidence="20"/>
    <dbReference type="AlphaFoldDB" id="A0A452HNH0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018833.1">
        <property type="protein sequence ID" value="ENSGAGP00000016501.1"/>
        <property type="gene ID" value="ENSGAGG00000012344.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060473">
        <property type="term" value="C:cortical granule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005764">
        <property type="term" value="C:lysosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004563">
        <property type="term" value="F:beta-N-acetylhexosaminidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005975">
        <property type="term" value="P:carbohydrate metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006689">
        <property type="term" value="P:ganglioside catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030203">
        <property type="term" value="P:glycosaminoglycan metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd06562">
        <property type="entry name" value="GH20_HexA_HexB-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.20.20.80:FF:000063">
        <property type="entry name" value="Beta-hexosaminidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.379.10">
        <property type="entry name" value="Chitobiase/beta-hexosaminidase domain 2-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.20.20.80">
        <property type="entry name" value="Glycosidases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025705">
        <property type="entry name" value="Beta_hexosaminidase_sua/sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017853">
        <property type="entry name" value="GH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015883">
        <property type="entry name" value="Glyco_hydro_20_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029018">
        <property type="entry name" value="Hex-like_dom2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029019">
        <property type="entry name" value="HEX_eukaryotic_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22600">
        <property type="entry name" value="BETA-HEXOSAMINIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22600:SF38">
        <property type="entry name" value="BETA-HEXOSAMINIDASE SUBUNIT BETA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00728">
        <property type="entry name" value="Glyco_hydro_20"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14845">
        <property type="entry name" value="Glycohydro_20b2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF001093">
        <property type="entry name" value="B-hxosamndse_ab_euk"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00738">
        <property type="entry name" value="GLHYDRLASE20"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51445">
        <property type="entry name" value="(Trans)glycosidases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55545">
        <property type="entry name" value="beta-N-acetylhexosaminidase-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="11" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="7 22" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="8" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10 20" id="KW-0326">Glycosidase</keyword>
    <keyword evidence="5 20" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="6" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="9" id="KW-0458">Lysosome</keyword>
    <keyword evidence="27" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 23" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="23">
        <location>
            <begin position="1"/>
            <end position="18"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019314410" description="Beta-hexosaminidase" evidence="23">
        <location>
            <begin position="19"/>
            <end position="534"/>
        </location>
    </feature>
    <feature type="domain" description="Beta-hexosaminidase eukaryotic type N-terminal" evidence="25">
        <location>
            <begin position="48"/>
            <end position="172"/>
        </location>
    </feature>
    <feature type="domain" description="Glycoside hydrolase family 20 catalytic" evidence="24">
        <location>
            <begin position="194"/>
            <end position="492"/>
        </location>
    </feature>
    <feature type="active site" description="Proton donor" evidence="21">
        <location>
            <position position="331"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="22">
        <location>
            <begin position="83"/>
            <end position="131"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="22">
        <location>
            <begin position="285"/>
            <end position="336"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="22">
        <location>
            <begin position="510"/>
            <end position="527"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001231"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004371"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00006285"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023228"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023295"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023505"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00037865"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00043767"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00043827"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00045511"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00046959"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00047301"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00049464"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="PIRNR" id="PIRNR001093"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="PIRSR" id="PIRSR001093-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="PIRSR" id="PIRSR001093-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="Pfam" id="PF00728"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="Pfam" id="PF14845"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016501.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="534" mass="60514" checksum="761D4BACD737E470" modified="2019-05-08" version="1">MEPARLFLGLLCASAALAASSLHQRPPQPEFLLEELELELGESPSGSLWPLPRSLRILPRQLQLAPARFQLVHGPGSTAGPGCALLQDAFRRYYEYIFGYSKWQNQGQRPLPAGTELLQLQVIIVSKDSECDNYPNITSNEAYQLSVTEPVAVLRAEKVWGALRGLETFSQLVYEDDYGTFLINESEITDYPRFAHRGILLDTSRHFLPLKSILANLVSWHFVVTSISYQLITGSYSYDHVYTPTDVRLVIDYARLRGIRVIPEFDSPGHTKSWGKGQKDLLTPCFNGEHRSGSYGPVNPILNTTYHFMTDLFKEISSVFPDAYIHLGGDEVDFTCWKSNPDVTEFMKQQGFGIDYAKLESYYIQKILDIVSSANKGYIVWQEVFDKGVKIKPDTIVEVWIGSLYTAELSRVTGAGFTTILAAPWYLDYISYGQDWKKYYSVEPLNFPGCEKQKKLVIGGEACLWGEFVDATNLTPRLWPRASAVGERLWSSENVTNIQDAYNRLVKHRCRMLRRGIAAEPLFVGYCDPEYRGR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQA0</accession>
    <name>A0A452IQA0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">UBC core domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="4">Belongs to the ubiquitin-conjugating enzyme family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQA0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034147.1">
        <property type="protein sequence ID" value="ENSGAGP00000030075.1"/>
        <property type="gene ID" value="ENSGAGG00000021696.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016740">
        <property type="term" value="F:transferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.110.10:FF:000074">
        <property type="entry name" value="ubiquitin-conjugating enzyme E2 D1 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.110.10">
        <property type="entry name" value="Ubiquitin Conjugating Enzyme"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000608">
        <property type="entry name" value="UBC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023313">
        <property type="entry name" value="UBQ-conjugating_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016135">
        <property type="entry name" value="UBQ-conjugating_enzyme/RWD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24068">
        <property type="entry name" value="UBIQUITIN-CONJUGATING ENZYME E2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00179">
        <property type="entry name" value="UQ_con"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00212">
        <property type="entry name" value="UBCc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54495">
        <property type="entry name" value="UBC-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00183">
        <property type="entry name" value="UBC_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50127">
        <property type="entry name" value="UBC_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="4" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0808">Transferase</keyword>
    <keyword evidence="2 4" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="UBC core" evidence="5">
        <location>
            <begin position="1"/>
            <end position="123"/>
        </location>
    </feature>
    <feature type="active site" description="Glycyl thioester intermediate" evidence="3">
        <location>
            <position position="61"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10133"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="RuleBase" id="RU362109"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50127"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030075.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="123" mass="13943" checksum="AE27D829AA0C58E3" modified="2019-05-08" version="1">MSGRRRTCLLLLGAPKPDSAYQGGVFFLTVHFPTDYPFKPPKIAFTTKIYHPNINSNGSICLDILRSQWSPALTVSKVLLSICSLLCDPNPDDPLVPDIAQIYKSDKEKYNRHAREWTQKYAM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQA1</accession>
    <name>A0A452IQA1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Zinc finger protein 367</fullName>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQA1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034093.1">
        <property type="protein sequence ID" value="ENSGAGP00000030023.1"/>
        <property type="gene ID" value="ENSGAGG00000021660.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000785">
        <property type="term" value="C:chromatin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031519">
        <property type="term" value="C:PcG protein complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005667">
        <property type="term" value="C:transcription regulator complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000474">
        <property type="entry name" value="zinc finger protein 367"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000535">
        <property type="entry name" value="Zinc finger protein 367"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14003">
        <property type="entry name" value="TRANSCRIPTIONAL REPRESSOR PROTEIN YY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14003:SF26">
        <property type="entry name" value="ZINC FINGER PROTEIN 367"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13912">
        <property type="entry name" value="zf-C2H2_6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0010">Activator</keyword>
    <keyword evidence="8" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="9" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="12" id="KW-0539">Nucleus</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="11" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 14" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="16">
        <location>
            <begin position="139"/>
            <end position="166"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="16">
        <location>
            <begin position="167"/>
            <end position="196"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="72"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="219"/>
            <end position="238"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="257"/>
            <end position="304"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="15">
        <location>
            <begin position="72"/>
            <end position="85"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="15">
        <location>
            <begin position="110"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="15">
        <location>
            <begin position="258"/>
            <end position="268"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00068072"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030023.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="327" mass="36481" checksum="7355AF81613AB8F2" modified="2019-05-08" version="1">PPLQAPGLAMAERPPPVIFCQDSPKRVLVSVIKTTPIKPSRGGEEPAPAAPGFSDFMVYPWRWGENAHNVTLSPSSATAPSSVQQPRVGVVRTSSGEDEEAGSPDSGGSHLKDGIRRGRPRADTVRDLISEGEHSSSRIRCNICNRVFPREKSLQAHKRTHTGERPYLCDYPDCGKAFVQSGQLKTHQRLHTGEKPFVCSENGCLSRFTHANRHCPKHPYARLKREEPTGRLSKKQTADNKAVADWLAKYWETREQRTPTLKSKTAQKADQEQQDPMEYLQSDEEDEEEKNSAPSAARRRLQEQRERLHGALALIELANLAGAPMRQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQA6</accession>
    <name>A0A452IQA6_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="12">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 7">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 7">Belongs to the ETS family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQA6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034099.1">
        <property type="protein sequence ID" value="ENSGAGP00000030029.1"/>
        <property type="gene ID" value="ENSGAGG00000021670.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043565">
        <property type="term" value="F:sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd08540">
        <property type="entry name" value="SAM_PNT-ERG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.50:FF:000010">
        <property type="entry name" value="Fli-1 proto-oncogene, ETS transcription factor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.10:FF:000039">
        <property type="entry name" value="Friend leukemia integration 1 transcription factor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.50">
        <property type="entry name" value="Transcription Factor, Ets-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.10">
        <property type="entry name" value="Winged helix-like DNA-binding domain superfamily/Winged helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Ets_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046328">
        <property type="entry name" value="ETS_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003118">
        <property type="entry name" value="Pointed_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001660">
        <property type="entry name" value="SAM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013761">
        <property type="entry name" value="SAM/pointed_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036388">
        <property type="entry name" value="WH-like_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036390">
        <property type="entry name" value="WH_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11849">
        <property type="entry name" value="ETS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11849:SF216">
        <property type="entry name" value="TRANSCRIPTIONAL REGULATOR ERG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00178">
        <property type="entry name" value="Ets"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02198">
        <property type="entry name" value="SAM_PNT"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="ETSDOMAIN"/>
    </dbReference>
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        <property type="entry name" value="ETS"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47769">
        <property type="entry name" value="SAM/Pointed domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46785">
        <property type="entry name" value="Winged helix' DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00345">
        <property type="entry name" value="ETS_DOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00346">
        <property type="entry name" value="ETS_DOMAIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="ETS_DOMAIN_3"/>
        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
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        <property type="entry name" value="SAM_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 7" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="6 7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0804">Transcription</keyword>
    <keyword evidence="3" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="PNT" evidence="11">
        <location>
            <begin position="113"/>
            <end position="199"/>
        </location>
    </feature>
    <feature type="domain" description="SAM" evidence="10">
        <location>
            <begin position="134"/>
            <end position="179"/>
        </location>
    </feature>
    <feature type="domain" description="ETS" evidence="9">
        <location>
            <begin position="311"/>
            <end position="391"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="34"/>
            <end position="56"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="244"/>
            <end position="304"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="34"/>
            <end position="47"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="270"/>
            <end position="281"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005562"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU004019"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50061"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50105"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="PROSITE" id="PS51433"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030029.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="479" mass="54088" checksum="F859B8E8EC8DC15B" modified="2019-05-08" version="1">MASTIKEALSVVSEDQSLFECAYGTQHLTKTEMTASSSSEYGQTSKMSPRVPQQDWLSQPPARVTIKMECNPNQINGSRNSPDDCSVAKGGKMVSGSDNVGMNYGSYMEEKHVPPPNMTTNERRVIVPADPTLWSTDHVRQWLEWAVKEYGLPDVDILLFQNIDGKELCKMTKDDFQRLTPSYNADILLSHLHYLRETPLPHLTSDDVDKALQNSPRLMHARNTGGAAFIFPNTSVYPEATQRITTRPDLSYEQARRSAWTSHNHPAPQSKATQPSSSTVPKTEDQRPQLDPYQILGPTSSRLANPGSGQIQLWQFLLELLSDSSNSNCITWEGTNGEFKMTDPDEVARRWGERKSKPNMNYDKLSRALRYYYDKNIMTKVHGKRYAYKFDFHGIAQALQPHPPESSMYKYPTDLPYMSPYHAHPQKMNFVAPHPPALPVTSSSFFAAPTTYWNSPTGSIYPNTRLPATHMPSHLGTYY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQA9</accession>
    <name>A0A452IQA9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="4">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the small GTPase superfamily. Rab family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQA9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034157.1">
        <property type="protein sequence ID" value="ENSGAGP00000030085.1"/>
        <property type="gene ID" value="ENSGAGG00000021701.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005525">
        <property type="term" value="F:GTP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003924">
        <property type="term" value="F:GTPase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd01861">
        <property type="entry name" value="Rab6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000139">
        <property type="entry name" value="ras-related protein Rab-6A isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050227">
        <property type="entry name" value="Rab"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005225">
        <property type="entry name" value="Small_GTP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001806">
        <property type="entry name" value="Small_GTPase"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00231">
        <property type="entry name" value="small_GTP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47977">
        <property type="entry name" value="RAS-RELATED PROTEIN RAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00071">
        <property type="entry name" value="Ras"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00449">
        <property type="entry name" value="RASTRNSFRMNG"/>
    </dbReference>
    <dbReference type="SMART" id="SM00175">
        <property type="entry name" value="RAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00176">
        <property type="entry name" value="RAN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00173">
        <property type="entry name" value="RAS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00174">
        <property type="entry name" value="RHO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51419">
        <property type="entry name" value="RAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51421">
        <property type="entry name" value="RAS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51420">
        <property type="entry name" value="RHO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0342">GTP-binding</keyword>
    <keyword evidence="2" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006270"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030085.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="208" mass="23521" checksum="DF53D3DC9600F98B" modified="2019-05-08" version="1">MSTGGDFGNPLRKFKLVFLGEQSVGKTSLITRFMYDSFDNTYQATIGIDFLSKTMYLEDRTIRLQLWDTAGQERFRSLIPSYIRDSAAAVVVYDITNVNSFQQTTKWIDDVRTERGSDVIIMLVGNKTDLADKRQVSIEEGERKAKELNVMFIETSAKAGYNVKQLFRRVAAALPGMESTQDKSREDMIDIKLEKPQEQPVSEGGCSC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQB2</accession>
    <name>A0A452IQB2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">SAFB-like transcription modulator</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQB2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034258.1">
        <property type="protein sequence ID" value="ENSGAGP00000030178.1"/>
        <property type="gene ID" value="ENSGAGG00000021279.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043565">
        <property type="term" value="F:sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050684">
        <property type="term" value="P:regulation of mRNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd12678">
        <property type="entry name" value="RRM_SLTM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.720.30:FF:000010">
        <property type="entry name" value="SAFB-like transcription modulator isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.330:FF:000238">
        <property type="entry name" value="SAFB-like transcription modulator isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.330">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.720.30">
        <property type="entry name" value="SAP domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012677">
        <property type="entry name" value="Nucleotide-bd_a/b_plait_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035979">
        <property type="entry name" value="RBD_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000504">
        <property type="entry name" value="RRM_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051738">
        <property type="entry name" value="SAF_Modulators"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003034">
        <property type="entry name" value="SAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036361">
        <property type="entry name" value="SAP_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15683:SF5">
        <property type="entry name" value="SAFB-LIKE TRANSCRIPTION MODULATOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15683">
        <property type="entry name" value="SCAFFOLD ATTACHMENT FACTOR B-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00076">
        <property type="entry name" value="RRM_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02037">
        <property type="entry name" value="SAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00360">
        <property type="entry name" value="RRM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00513">
        <property type="entry name" value="SAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54928">
        <property type="entry name" value="RNA-binding domain, RBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF68906">
        <property type="entry name" value="SAP domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50102">
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="SAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0678">Repressor</keyword>
    <keyword evidence="3 9" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="6" id="KW-0804">Transcription</keyword>
    <keyword evidence="4" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="SAP" evidence="12">
        <location>
            <begin position="26"/>
            <end position="60"/>
        </location>
    </feature>
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            <end position="464"/>
        </location>
    </feature>
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            <begin position="1"/>
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        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
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            <begin position="60"/>
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        </location>
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    <feature type="region of interest" description="Disordered" evidence="10">
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            <end position="1036"/>
        </location>
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            <begin position="1"/>
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        </location>
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            <begin position="94"/>
            <end position="105"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="10">
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            <begin position="113"/>
            <end position="124"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
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            <begin position="139"/>
            <end position="171"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="10">
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        </location>
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        </location>
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        </location>
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        </location>
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            <begin position="1024"/>
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        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022491"/>
        </source>
    </evidence>
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        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
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            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        <source>
            <dbReference type="PROSITE" id="PS50800"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030178.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1036" mass="117638" checksum="90F683B895B830EA" modified="2019-05-08" version="1">MAAAAAVSSPAAAPSVGPAQMESKKITDLRVIDLKSELKRRNLDITGVKTVLISRLKQAIEEEGGDPDNIEITVSADTPIKKPTKGKGKKQETDELSGDASVEDDSFVKESELEVQDASDQDGNDELKDFEEAGENEEENSHSKELRPSEKKKYSDSKQRKTAEDAEKDFESQENDLQDTEDDTFPTVHNGEEEENEKDKAGSGDGIQEVSKPLPSEESLAEADHSAHEEMEANTSMKEAEDDNISVTIQAEDAITLDFDGDDLLETGKNVKITDSEASKAKDGQDTIAQSLEKERKDYEMTENHKDGKKEDCVKGDPVKKEAREGSKKAESGDKEKDTLKKGPSSTGASGQAKSSAKESKESKTTSKDDKGSASSVSGSSGSSTRNLWVSGLSSNTKAADLKNLFGKYGKVLSAKVVTNARSPGAKCYGIVTMSSSTEVARCIAHLHRTELHGQQISVEKVKGDPSKKELKKEIDEKSGSGRNMGDKKTASIDKTSKTQSAKKEEKKSDKSEKKESKEAKKTEGKDEKNDNGSSGPNQESTKKTEEKKRISGKSPGQMVVLDQTKGDQGHTRTVRRGRFDKPLVLRNKERFIQDKMKFREYRGRKDILPFEKMKEQRMREHMVRLERIRRAVELRRRRELAERERRERERIRIIREREERERLQRERERLEIERQKLERERMERERLERERIRIEQERRKEAERIAREREELRRQQQQLRYEQEKRNSLKRPRDVDHRRDDPYWNENKKMALDTDARFSHGSDYSRQQNRFNDFDHRERGRYPEGSSVPSSSFERRERFVNQSEGKKTRPTARREEPGFERYPKNFSESRRNEPPQPRNELRDTDRREVRGDRDERRTVIIHDRPEISHGRHPRETGSNPPRQTNWKNEGGISTDKRDARGERPERSGREVSGHVRSAAPGSRSSTSGYGSREGERGVIGERGSGGQHYNEDRHVVERHSRETGPRKEWHGPSSQGSGYHDARRMGDGRGGGGMMSPHTSNSSPINRVVQITGSSMQRGSGSGFKPFKSGPPRRF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQB4</accession>
    <name>A0A452IQB4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2 11">Arginase</fullName>
            <ecNumber evidence="2 11">3.5.3.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="8 11">
            <text>L-arginine + H2O = urea + L-ornithine</text>
            <dbReference type="Rhea" id="RHEA:20569"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:16199"/>
            <dbReference type="ChEBI" id="CHEBI:32682"/>
            <dbReference type="ChEBI" id="CHEBI:46911"/>
            <dbReference type="EC" id="3.5.3.1"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="11">
            <name>Mn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29035"/>
        </cofactor>
        <text evidence="11">Binds 2 manganese ions per subunit.</text>
    </comment>
    <comment type="pathway">
        <text evidence="1 11">Nitrogen metabolism; urea cycle; L-ornithine and urea from L-arginine: step 1/1.</text>
    </comment>
    <comment type="similarity">
        <text evidence="9 10">Belongs to the arginase family.</text>
    </comment>
    <dbReference type="EC" id="3.5.3.1" evidence="2 11"/>
    <dbReference type="AlphaFoldDB" id="A0A452IQB4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034162.1">
        <property type="protein sequence ID" value="ENSGAGP00000030090.1"/>
        <property type="gene ID" value="ENSGAGG00000021614.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00158">
        <property type="reaction ID" value="UER00270"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004053">
        <property type="term" value="F:arginase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030145">
        <property type="term" value="F:manganese ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006525">
        <property type="term" value="P:arginine metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000050">
        <property type="term" value="P:urea cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="CDD" id="cd09989">
        <property type="entry name" value="Arginase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.800.10:FF:000012">
        <property type="entry name" value="Arginase"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Ureohydrolase domain"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Arginase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006035">
        <property type="entry name" value="Ureohydrolase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023696">
        <property type="entry name" value="Ureohydrolase_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020855">
        <property type="entry name" value="Ureohydrolase_Mn_BS"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01229">
        <property type="entry name" value="rocF_arginase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43782">
        <property type="entry name" value="ARGINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43782:SF2">
        <property type="entry name" value="ARGINASE-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00491">
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00116">
        <property type="entry name" value="ARGINASE"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52768">
        <property type="entry name" value="Arginase/deacetylase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01053">
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51409">
        <property type="entry name" value="ARGINASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 11" id="KW-0056">Arginine metabolism</keyword>
    <keyword evidence="6 10" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="7 11" id="KW-0464">Manganese</keyword>
    <keyword evidence="5 11" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 11" id="KW-0835">Urea cycle</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00005098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00012168"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022436"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022503"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023211"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00047391"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00742"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="RuleBase" id="RU003684"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="RuleBase" id="RU361159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030090.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="266" mass="28737" checksum="1EDE1086C7F7DE19" modified="2019-05-08" version="1">MYFLLKIDIIPNDLPHHNAKNPRTVGCANEKLASAVEEVKKDGRTCLVVGGDHSLAIGSISGHAKTEPDLGVIWFDAHADINTPLTSLSGNIHGQPVAFLLKELKVSGVPGFKWLKPCLSVENIVYIGLRDVDKAEHYFLKSLGIKYYSMTEIDQLGIGKVMEEALDYLLANKKRPIHLSFDVDGLDPSLTPATGTPVPGGVTLREGIYVAEQLYKTGLLSAVDIVEINPRLGKTQEGVDSTVNTAISIILSCFGKAREGSHSCSC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQC2</accession>
    <name>A0A452IQC2_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="20">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Cell cortex</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the FNBP1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQC2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034115.1">
        <property type="protein sequence ID" value="ENSGAGP00000030044.1"/>
        <property type="gene ID" value="ENSGAGG00000021677.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005938">
        <property type="term" value="C:cell cortex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008289">
        <property type="term" value="F:lipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006897">
        <property type="term" value="P:endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd07676">
        <property type="entry name" value="F-BAR_FBP17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd11629">
        <property type="entry name" value="HR1_FBP17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12071">
        <property type="entry name" value="SH3_FBP17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1270.60:FF:000002">
        <property type="entry name" value="Formin-binding protein 1-like isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.40:FF:000017">
        <property type="entry name" value="Formin-binding protein 1-like isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.470">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.60">
        <property type="entry name" value="Arfaptin homology (AH) domain/BAR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027267">
        <property type="entry name" value="AH/BAR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031160">
        <property type="entry name" value="F_BAR_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001060">
        <property type="entry name" value="FCH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037449">
        <property type="entry name" value="FNBP1_F-BAR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035492">
        <property type="entry name" value="FNBP1_SH3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011072">
        <property type="entry name" value="HR1_rho-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057870">
        <property type="entry name" value="HR1_TOCA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15735">
        <property type="entry name" value="FCH AND DOUBLE SH3 DOMAINS PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15735:SF13">
        <property type="entry name" value="FORMIN-BINDING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00611">
        <property type="entry name" value="FCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25610">
        <property type="entry name" value="HR1_TOCA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00018">
        <property type="entry name" value="SH3_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00055">
        <property type="entry name" value="FCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00326">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103657">
        <property type="entry name" value="BAR/IMD domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51741">
        <property type="entry name" value="F_BAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51860">
        <property type="entry name" value="REM_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="9 14 15" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="12" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="8" id="KW-0254">Endocytosis</keyword>
    <keyword evidence="10" id="KW-0446">Lipid-binding</keyword>
    <keyword evidence="11" id="KW-0472">Membrane</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 13" id="KW-0728">SH3 domain</keyword>
    <feature type="domain" description="F-BAR" evidence="18">
        <location>
            <begin position="1"/>
            <end position="264"/>
        </location>
    </feature>
    <feature type="domain" description="REM-1" evidence="19">
        <location>
            <begin position="416"/>
            <end position="493"/>
        </location>
    </feature>
    <feature type="domain" description="SH3" evidence="17">
        <location>
            <begin position="562"/>
            <end position="623"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="293"/>
            <end position="320"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="333"/>
            <end position="366"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="501"/>
            <end position="545"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="15">
        <location>
            <begin position="423"/>
            <end position="483"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="16">
        <location>
            <begin position="338"/>
            <end position="347"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="507"/>
            <end position="524"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004544"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00009426"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022583"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023121"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01077"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS51741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS51860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030044.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="633" mass="73233" checksum="F7747674B42645C6" modified="2019-05-08" version="1">MSWGTELWDQFDNLEKHTQWGIDVLEKYIKFVKERTEIEISYAKQLRNLSKKYQPKKNSKEEEEYKYTSCKAFLATLNEMNDYAGQHEVISENMTSQITAELARFVQELKQERKSHFHDGRKAQQYIETFWKQLEASKRRFERDCKEADRAQQYFEKMDADINVTKADVEKARQQAQIRHQMAEDSKAEYSSILQKFNHDQHEHYHTHIPNIFQKIQEMEERRIVRIGECMKTYAEVDRQVIPIIGKCLDGITKAAESIDQKNDSLLVIEAFKSGFEPPGDIDFEDFTQPMKRTVSESSLSNSRWDGKSEPKPGSKSKGKLWPFIKKNKLMSLLTSPHQPPPPPPASASPSAVPNGPQSPKQQKEPLSHRFNEFMTSKPKIHCFRSLKRGLSLKLITLNTQVWNLIEGGGPEDFSNLPPEQRRKKLQQKVDELNKEIQKEMDQRDALTKMKDVYIKNPQMGDAASVDHKLAELGQNMEKLRLEAQKFEGWLAEVEGRLPVRTEQSRRQSGMYDTQNTSGVNNCAQDRESPDGSYTEEPSQESEMKVPVTEFDDEFDDEEPLPTIGTCKALYTFEGQNEGTISVMEGEMLYVIEEDKGDGWTRIRRNEDEEGYVPTSYVEVYLDKNAKGAMTYI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQC5</accession>
    <name>A0A452IQC5_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="14">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endomembrane system</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="9">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000034404.1">
        <property type="protein sequence ID" value="ENSGAGP00000030312.1"/>
        <property type="gene ID" value="ENSGAGG00000021780.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005770">
        <property type="term" value="C:late endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005802">
        <property type="term" value="C:trans-Golgi network"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005537">
        <property type="term" value="F:D-mannose binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005520">
        <property type="term" value="F:insulin-like growth factor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0038023">
        <property type="term" value="F:signaling receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007041">
        <property type="term" value="P:lysosomal transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00062">
        <property type="entry name" value="FN2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.70.130.10:FF:000004">
        <property type="entry name" value="Insulin-like growth factor 2 receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.70.130.10:FF:000005">
        <property type="entry name" value="Insulin-like growth factor 2 receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.70.130.10:FF:000006">
        <property type="entry name" value="Insulin-like growth factor 2 receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.70.130.10:FF:000007">
        <property type="entry name" value="Insulin-like growth factor 2 receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.70.130.10:FF:000009">
        <property type="entry name" value="Insulin-like growth factor 2 receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.70.130.10:FF:000011">
        <property type="entry name" value="Insulin-like growth factor 2 receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.70.130.10:FF:000012">
        <property type="entry name" value="Insulin-like growth factor 2 receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.70.130.10:FF:000013">
        <property type="entry name" value="Insulin-like growth factor 2 receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.70.130.10:FF:000016">
        <property type="entry name" value="Insulin-like growth factor 2 receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.70.130.10:FF:000017">
        <property type="entry name" value="Insulin-like growth factor 2 receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.70.130.10:FF:000028">
        <property type="entry name" value="Mannose-6-phosphate/insulin-like growth factor II receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.10.10">
        <property type="entry name" value="Fibronectin, type II, collagen-binding"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.70.130.10">
        <property type="entry name" value="Mannose-6-phosphate receptor binding domain"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000479">
        <property type="entry name" value="CIMR_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000562">
        <property type="entry name" value="FN_type2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036943">
        <property type="entry name" value="FN_type2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013806">
        <property type="entry name" value="Kringle-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009011">
        <property type="entry name" value="Man6P_isomerase_rcpt-bd_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044865">
        <property type="entry name" value="MRH_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15071:SF17">
        <property type="entry name" value="CATION-INDEPENDENT MANNOSE-6-PHOSPHATE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15071">
        <property type="entry name" value="MANNOSE-6-PHOSPHATE RECEPTOR FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00878">
        <property type="entry name" value="CIMR"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00040">
        <property type="entry name" value="fn2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00013">
        <property type="entry name" value="FNTYPEII"/>
    </dbReference>
    <dbReference type="SMART" id="SM01404">
        <property type="entry name" value="CIMR"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="SMART" id="SM00059">
        <property type="entry name" value="FN2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57440">
        <property type="entry name" value="Kringle-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50911">
        <property type="entry name" value="Mannose 6-phosphate receptor domain"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51092">
        <property type="entry name" value="FN2_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51914">
        <property type="entry name" value="MRH"/>
        <property type="match status" value="14"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="7 11" id="KW-0472">Membrane</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="3 11" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 11" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="2" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="2234"/>
            <end position="2256"/>
        </location>
    </feature>
    <feature type="domain" description="MRH" evidence="13">
        <location>
            <begin position="17"/>
            <end position="134"/>
        </location>
    </feature>
    <feature type="domain" description="MRH" evidence="13">
        <location>
            <begin position="143"/>
            <end position="289"/>
        </location>
    </feature>
    <feature type="domain" description="MRH" evidence="13">
        <location>
            <begin position="295"/>
            <end position="442"/>
        </location>
    </feature>
    <feature type="domain" description="MRH" evidence="13">
        <location>
            <begin position="447"/>
            <end position="591"/>
        </location>
    </feature>
    <feature type="domain" description="MRH" evidence="13">
        <location>
            <begin position="597"/>
            <end position="729"/>
        </location>
    </feature>
    <feature type="domain" description="MRH" evidence="13">
        <location>
            <begin position="732"/>
            <end position="889"/>
        </location>
    </feature>
    <feature type="domain" description="MRH" evidence="13">
        <location>
            <begin position="1024"/>
            <end position="1160"/>
        </location>
    </feature>
    <feature type="domain" description="MRH" evidence="13">
        <location>
            <begin position="1166"/>
            <end position="1302"/>
        </location>
    </feature>
    <feature type="domain" description="MRH" evidence="13">
        <location>
            <begin position="1306"/>
            <end position="1447"/>
        </location>
    </feature>
    <feature type="domain" description="MRH" evidence="13">
        <location>
            <begin position="1453"/>
            <end position="1591"/>
        </location>
    </feature>
    <feature type="domain" description="MRH" evidence="13">
        <location>
            <begin position="1593"/>
            <end position="1735"/>
        </location>
    </feature>
    <feature type="domain" description="MRH" evidence="13">
        <location>
            <begin position="1740"/>
            <end position="1925"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-II" evidence="12">
        <location>
            <begin position="1834"/>
            <end position="1880"/>
        </location>
    </feature>
    <feature type="domain" description="MRH" evidence="13">
        <location>
            <begin position="1928"/>
            <end position="2063"/>
        </location>
    </feature>
    <feature type="domain" description="MRH" evidence="13">
        <location>
            <begin position="2087"/>
            <end position="2215"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="2349"/>
            <end position="2422"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="2396"/>
            <end position="2410"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004308"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS51092"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS51914"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030312.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2422" mass="269555" checksum="F63DC8ED6E79C191" modified="2019-05-08" version="1">MALSIPVVRYVLTYRLPFCFSFSYTWEAFDVDKHVRYKINPCGGIEDCDGSSAICAYDLNKKVYLSVGDSSLKRSSKSLLEFNTTKPCLQQGAQHVVQSNINFLCGKTLGTPELVTATECVHYFEWRTFIACKKGSFKAKKEVPCYVFDDLKKHDLNPLIKTSGGYLVDDSDEDSLYINVCRDIGSSSSETRSCPVGSAACLLSEGHAFDVGSPQEQLKLRDKDRLVLNYEKTYSEEEKPNFCQGHNPAVTITFVCPSKRREERTGPKLTAKLKCRYEIEWVTEYACHRDYLESRNCSLTSEQHDVSIDLTPLTQQPSYVTPYVAKDEKEEYYYYLNICGKTSAGNCKDDTESVSSCQVKTRGQQEKVAGKFKNQTLRYSDGDLTLIYPDGDECSSGFQRMTVINFECNETAGNEGKGAPVFTGEVDCTYFFTWDTKYACVQEKEDLLCRVADKKKRYDLSPLIRSSESAQNWEAVNSSPVEADKHFFINVCHKVLKKGAAVVCSDDAAVCSVDKNNKPKNLGTVMSALKVGERIQLTYSNGDSCGNNNMIKTVITLVCKPGDLESAPVFNLENDECLYVFEWYTAAACVLSKTEGDNCRVSDAQAGFSFDLSPLTKRSGSYVINTTDYDFYINICSSVPEEHCLLKSAAACQVAKNLGEPSSKLSYYDGMIQLTYKNGTTYNNEEKTERSTLITFLCDYEAGIGQPEYQVEDNYTYNFKWYTKYACPEMPLECVVTDPNTMEQYDLSSLSKSEDRGENWYAMDNSEPNERRKYYINVCRPLVPVAGCDRYASGCQMKFEKDNDSFSETTSISNLGIASKGLVIEGRGRVLLTYTNGSVCISADGKRTSYTTIIHFTCSKGTLNSSPRFLDNQECVVTFLWETEAACPVTTTKDETQSCSVRDPNSGFLFNMQPLASEKGYITSGIGKTFMLNICGPAPDCGDGSDGKPAGGCEVEGLKSVRIVGLDKTLSLSSEGFLTLTYRGNFEVKSASDTFTVTFICNDSYPGELKFVDEEINSAAGIHNTFFEFHTALYDLSDLSKEGEPWIAVDTSKDAKTRTFYLNVCKPLPYVPGCPGGAIGSCVKSADKGQNLGIIQISPQAATDGSLSIIYLNGDTCKDKQRYSTRILFECDQTLGSPMFQHEDGCEFVFIWRTLAACPVHRAEGDNCQVKDPRYGFVYDLRPLSEKEKAVSTDEYDYYFRVCERITEACPQVNRLSNVSSCQVKKQDHSVKVAGLYTEKLTFEHGLIKINYTSGERCHKIYERSTAIFFYCDHNTLEPVFLKETPDCTYLFEWHTQYACPPFRSIECSYKDNKGNSFDLSPLTRHRENWEAIPMSGSTQKYYINVCQSLVLHAGAGSCPSDAAACLVDGSKYINLGEVADGPRWENGISVLKYVNGEPCPDKIRKRTTIIRLKCDESRVESKPELVTTIEECEYTFLWFTAAACPLKRNVQNDCRVTNPATGHLFDLNSLKRESGYRLYDSKKRRTLQLNVCNEAKSPCGSGVGACIIGGQKPINAGRLSKMLIYEDQLLKLVYSGGDPCPANAELKHTSFFSFVCKSDAGAGSQPVLLSFDEQTCTHYFSWHTTLACEEEVKCSVLNGSSVIDLSPLIHRTGYYEAFVDELTDSTPDFYINICEPLNPVADVNCPPGAAVCMDPVNGPPIDIGRITGPPKINPAVNEVYITFNSTTACEADKTFSSLIVFHCKRGTSLGNPKMIRKSDCSFVFEWATPLVCPDKVNMLGCSVTDELLNYTFNLSSLSGSSYKVTLTVYHIGVCAEVADVPQGKCIDGAVCLVSENNAASFGIIKGMKMDYRHQDETVILQYTGGDRCPPVTEKGELCVFPFKYKGKRYEDCIVDEEKRRWCATTNDYDKDGKRGFCGNATGNREATIIFKCDDNAGNGNPQLLSETLGCAVTFEWKTQFVCPPKKMECKFIRKHKTYDLRVLSSLTGSWLFNYHGIAYYMNLCQRINNGPTNCPERAAICSKSRNGAVRVLGVVHTQKLNVTDDKVLVSYSGGHECRKNKKATTIIELKCAKTIGEPRLHRIDEENCTYYITWGTRAACAVKPQEVEIVNGMVINPTTGKNFSLGDIYYKLYQASGDIRTNGDTYVYEIQLSAITDSAHPQCSGAKVCQIKTTSTYIRKVGSSDKAKYYVQDDDLDVVFSSDSACGKDKSKFVSSTILFHCNLQAKEGIPEFLHETADCQYLFTWYTSAVCPLGQSDEGAQIFEGLSGRSQAVGAVLSVLLVVLTACLLILLFYKKERRETVIHKITNCCRRSSNVSYKYTKINTEEEANENETEWLMEEIAASNQRPGKGGQENGHITTTSVKSEALTSLHVDDLDSEDEVLTVPDVKIHSTRGVQSRGSDQNRHQQKKPFSLGSDAKTNLLNGGKDRKTKPNPGQQKTQNSTNTLSFHDDSDEDLLNV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQC6</accession>
    <name>A0A452IQC6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Palladin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="19">Interacts with EPS8. Interacts with LASP1. Interacts with VASP. Interacts with ACTN. Interacts with SORBS2. Interacts with PFN1. Interacts with LPP. Interacts with SPIN90. Interacts with SRC. Interacts with EZR. Interacts with RAI14.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="4">Cell junction</location>
            <location evidence="4">Focal adhesion</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="6">Cell projection</location>
            <location evidence="6">Axon</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="8">Cell projection</location>
            <location evidence="8">Growth cone</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="7">Cell projection</location>
            <location evidence="7">Lamellipodium</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cell projection</location>
            <location evidence="1">Podosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="5">Cell projection</location>
            <location evidence="5">Ruffle</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Myofibril</location>
            <location evidence="2">Sarcomere</location>
            <location evidence="2">Z line</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="18">Belongs to the myotilin/palladin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQC6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034275.1">
        <property type="protein sequence ID" value="ENSGAGP00000030194.1"/>
        <property type="gene ID" value="ENSGAGG00000021781.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005925">
        <property type="term" value="C:focal adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030426">
        <property type="term" value="C:growth cone"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030027">
        <property type="term" value="C:lamellipodium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002102">
        <property type="term" value="C:podosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001726">
        <property type="term" value="C:ruffle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001725">
        <property type="term" value="C:stress fiber"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030018">
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        <property type="term" value="F:protein kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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        <property type="match status" value="1"/>
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        <property type="entry name" value="palladin isoform X1"/>
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        <property type="match status" value="3"/>
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        <property type="entry name" value="IMMUNOGLOBULIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47633:SF10">
        <property type="entry name" value="PALLADIN, CYTOSKELETAL ASSOCIATED PROTEIN"/>
        <property type="match status" value="1"/>
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        <property type="match status" value="3"/>
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        <property type="match status" value="3"/>
    </dbReference>
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        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="14" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="12" id="KW-0965">Cell junction</keyword>
    <keyword evidence="16" id="KW-0966">Cell projection</keyword>
    <keyword evidence="9" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="15" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="13" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="17" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="10" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="11" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="Ig-like" evidence="22">
        <location>
            <begin position="350"/>
            <end position="434"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="22">
        <location>
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            <end position="569"/>
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            <end position="57"/>
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            <end position="99"/>
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            <end position="342"/>
        </location>
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        <source>
            <dbReference type="ARBA" id="ARBA00004188"/>
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    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
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    <evidence type="ECO:0000256" key="21">
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        <source>
            <dbReference type="PROSITE" id="PS50835"/>
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    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="732" mass="80829" checksum="BB27E32EA0A3073B" modified="2019-05-08" version="1">GGQRYHHHQQHQNPPPSPPFPPPPSFQELLSSQSVTSPVQMSILNSHTSPTMQSSSTFSYARPKQFIAAQNISPASGYVTPSSGSSTSSLPSPMSPTASQKQFGRAPVPPFSQPFAPEGEYLWSPSSSSPPPPPPPVFSPTATFSVSDSFPPPPPPLPSSVSSPSRSLSPTARFSNSSQSPAAFLSSMLPSQPPPISVNALGLPKGVMPPGFPKKTGRTARIASDEEIQGTKDAVIQDLERKLRFKEELLNNGQPRLTYEEKMARRLLGADNAATVFNIQEPDEEPTTQEACSLDASVMSPVDNQKEYKVSSFEQRLISEIEYRLERSPVEESDDEVQHGDEPTDNSVAPYFEMKLKHYKIFEGTSVTFTCRVAGNPKPKIYWFKDGKQISKRSDHYRIQREPDGTCSLHTTASTLDDDGNYTIMAANPQGRISCTGRLMVQAVNQRGRSPRTPPSQPHIRRPRSRSRDSGDENEPIQERFFRPHFLQAPGDLIVQEGKLCRMDCKVSGLPTPDLSWQLNGRPIRPDSSHKMLVRENGVHSLIIEPVTARDSGIYTCIATNRAGQNTFSLELIVAAKEEHKPPVFIEKLQNTGVADGYPVRLECRVSGVPYPQIFWKKENESLTYNTERVSMHQDNHGYICLLIQGATKEDAGWYTVSAKNEAGIVSCTARLDVYTQWQQQPQSTKPKKVRPSASRYAALSDQGLDIKAAFQPEANPVHLTLQCGLVESDDL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQC7</accession>
    <name>A0A452IQC7_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="19">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Cell cortex</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the FNBP1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQC7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034120.1">
        <property type="protein sequence ID" value="ENSGAGP00000030049.1"/>
        <property type="gene ID" value="ENSGAGG00000021677.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005938">
        <property type="term" value="C:cell cortex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008289">
        <property type="term" value="F:lipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006897">
        <property type="term" value="P:endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd07676">
        <property type="entry name" value="F-BAR_FBP17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd11629">
        <property type="entry name" value="HR1_FBP17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12071">
        <property type="entry name" value="SH3_FBP17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1270.60:FF:000002">
        <property type="entry name" value="Formin-binding protein 1-like isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.40:FF:000017">
        <property type="entry name" value="Formin-binding protein 1-like isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.60">
        <property type="entry name" value="Arfaptin homology (AH) domain/BAR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027267">
        <property type="entry name" value="AH/BAR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031160">
        <property type="entry name" value="F_BAR_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001060">
        <property type="entry name" value="FCH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037449">
        <property type="entry name" value="FNBP1_F-BAR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035492">
        <property type="entry name" value="FNBP1_SH3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011072">
        <property type="entry name" value="HR1_rho-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057870">
        <property type="entry name" value="HR1_TOCA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15735">
        <property type="entry name" value="FCH AND DOUBLE SH3 DOMAINS PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15735:SF13">
        <property type="entry name" value="FORMIN-BINDING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00611">
        <property type="entry name" value="FCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25610">
        <property type="entry name" value="HR1_TOCA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00018">
        <property type="entry name" value="SH3_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00055">
        <property type="entry name" value="FCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00326">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103657">
        <property type="entry name" value="BAR/IMD domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51741">
        <property type="entry name" value="F_BAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51860">
        <property type="entry name" value="REM_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="9 14" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="12" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="8" id="KW-0254">Endocytosis</keyword>
    <keyword evidence="10" id="KW-0446">Lipid-binding</keyword>
    <keyword evidence="11" id="KW-0472">Membrane</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 13" id="KW-0728">SH3 domain</keyword>
    <feature type="domain" description="F-BAR" evidence="17">
        <location>
            <begin position="1"/>
            <end position="264"/>
        </location>
    </feature>
    <feature type="domain" description="REM-1" evidence="18">
        <location>
            <begin position="399"/>
            <end position="476"/>
        </location>
    </feature>
    <feature type="domain" description="SH3" evidence="16">
        <location>
            <begin position="545"/>
            <end position="606"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="293"/>
            <end position="321"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="333"/>
            <end position="366"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="384"/>
            <end position="411"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="484"/>
            <end position="528"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="15">
        <location>
            <begin position="338"/>
            <end position="347"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="15">
        <location>
            <begin position="490"/>
            <end position="507"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004544"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00009426"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022583"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023121"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01077"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS51741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS51860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030049.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="616" mass="71253" checksum="B62083D1D3001182" modified="2019-05-08" version="1">MSWGTELWDQFDNLEKHTQWGIDVLEKYIKFVKERTEIEISYAKQLRNLSKKYQPKKNSKEEEEYKYTSCKAFLATLNEMNDYAGQHEVISENMTSQITAELARFVQELKQERKSHFHDGRKAQQYIETFWKQLEASKRRFERDCKEADRAQQYFEKMDADINVTKADVEKARQQAQIRHQMAEDSKAEYSSILQKFNHDQHEHYHTHIPNIFQKIQEMEERRIVRIGECMKTYAEVDRQVIPIIGKCLDGITKAAESIDQKNDSLLVIEAFKSGFEPPGDIDFEDFTQPMKRTVSESSLSNSRWDGKSEPKPGSKSKGKLWPFIKKNKLMSLLTSPHQPPPPPPASASPSAVPNGPQSPKQQKEPLSHRFNEFMTSKPKIHCFRSLKRGGGGPEDFSNLPPEQRRKKLQQKVDELNKEIQKEMDQRDALTKMKDVYIKNPQMGDAASVDHKLAELGQNMEKLRLEAQKFEGWLAEVEGRLPVRTEQSRRQSGMYDTQNTSGVNNCAQDRESPDGSYTEEPSQESEMKVPVTEFDDEFDDEEPLPTIGTCKALYTFEGQNEGTISVMEGEMLYVIEEDKGDGWTRIRRNEDEEGYVPTSYVEVYLDKNAKGAMTYI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQC9</accession>
    <name>A0A452IQC9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Amine oxidase</fullName>
            <ecNumber evidence="16">1.4.3.-</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="14">
            <text>an L-alpha-amino acid + O2 + H2O = a 2-oxocarboxylate + H2O2 + NH4(+)</text>
            <dbReference type="Rhea" id="RHEA:13781"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15379"/>
            <dbReference type="ChEBI" id="CHEBI:16240"/>
            <dbReference type="ChEBI" id="CHEBI:28938"/>
            <dbReference type="ChEBI" id="CHEBI:35179"/>
            <dbReference type="ChEBI" id="CHEBI:59869"/>
            <dbReference type="EC" id="1.4.3.2"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 16">
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the flavin monoamine oxidase family. FIG1 subfamily.</text>
    </comment>
    <dbReference type="EC" id="1.4.3.-" evidence="16"/>
    <dbReference type="AlphaFoldDB" id="A0A452IQC9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034180.1">
        <property type="protein sequence ID" value="ENSGAGP00000030105.1"/>
        <property type="gene ID" value="ENSGAGG00000021719.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001716">
        <property type="term" value="F:L-amino-acid oxidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090729">
        <property type="term" value="F:toxin activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009063">
        <property type="term" value="P:amino acid catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042742">
        <property type="term" value="P:defense response to bacterium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.405.10:FF:000004">
        <property type="entry name" value="Amine oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.660.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.50.50.60">
        <property type="entry name" value="FAD/NAD(P)-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.405.10">
        <property type="entry name" value="Guanine Nucleotide Dissociation Inhibitor, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002937">
        <property type="entry name" value="Amino_oxidase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036188">
        <property type="entry name" value="FAD/NAD-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001613">
        <property type="entry name" value="Flavin_amine_oxidase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050281">
        <property type="entry name" value="Flavin_monoamine_oxidase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10742:SF355">
        <property type="entry name" value="AMINE OXIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10742">
        <property type="entry name" value="FLAVIN MONOAMINE OXIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01593">
        <property type="entry name" value="Amino_oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00757">
        <property type="entry name" value="AMINEOXDASEF"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54373">
        <property type="entry name" value="FAD-linked reductases, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51905">
        <property type="entry name" value="FAD/NAD(P)-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0044">Antibiotic</keyword>
    <keyword evidence="5" id="KW-0929">Antimicrobial</keyword>
    <keyword evidence="11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="8 16" id="KW-0274">FAD</keyword>
    <keyword evidence="6 16" id="KW-0285">Flavoprotein</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13" id="KW-1199">Hemostasis impairing toxin</keyword>
    <keyword evidence="9 16" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="7" id="KW-0800">Toxin</keyword>
    <feature type="domain" description="Amine oxidase" evidence="17">
        <location>
            <begin position="66"/>
            <end position="497"/>
        </location>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="67"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <begin position="86"/>
            <end position="87"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <begin position="108"/>
            <end position="111"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="111"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="276"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="474"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001974"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005465"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022529"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022630"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022656"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022827"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023022"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00047637"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PIRSR" id="PIRSR601613-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="RuleBase" id="RU362067"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF01593"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030105.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="517" mass="59080" checksum="AF497077AC5F1214" modified="2019-05-08" version="1">NLNGTDAKSTLHGFSWWGWLMRDSQCMTRWEKCFIDPDYEELVAIAKNGLETGCRSKKVVIVGAGISGLTAAKLLKDAGCKVFILEASNHLGGRILTHREQDWYVELGAMRLPGHHRLVREFIKKLKLKLNPFYNSNDKGWYFVNNVRARAGEVERNPDILQYPVLPKERGKSASQLYNQTLDKVTTNCSVLKEKYDSFSTKEYLIKEGNLSRGAIDMIGDVMNEDGLFHMSFLYSVMIFTTFSEKTYDEITGGFDQLPQAFYQDIHRSVIFNSPVVKILQDNDQVKVFYRRPHSPFPLSVMADYVLVTATAKATRLIKFSPPLSAKKTHALRSIHYARATKIFLICSEKFWQKDGICNGRSITDHPSRVIYYPSHNFSSGAAVILASYTWVDDAEFFIPLDDEKCVDVVLEDLAEIHRVPKNYIQKICNKHVIKKWGLDKYTMGGYVSFTPYQFVDYSKALFQNEGRVHFSGEHTAQPHAWIETSMKSAVRAARNIHGAINALPWQQEKKLKENEL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQD1</accession>
    <name>A0A452IQD1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">SAFB-like transcription modulator</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQD1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034280.1">
        <property type="protein sequence ID" value="ENSGAGP00000030199.1"/>
        <property type="gene ID" value="ENSGAGG00000021279.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043565">
        <property type="term" value="F:sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050684">
        <property type="term" value="P:regulation of mRNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd12678">
        <property type="entry name" value="RRM_SLTM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.720.30:FF:000010">
        <property type="entry name" value="SAFB-like transcription modulator isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.330:FF:000238">
        <property type="entry name" value="SAFB-like transcription modulator isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.330">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.720.30">
        <property type="entry name" value="SAP domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012677">
        <property type="entry name" value="Nucleotide-bd_a/b_plait_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035979">
        <property type="entry name" value="RBD_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000504">
        <property type="entry name" value="RRM_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051738">
        <property type="entry name" value="SAF_Modulators"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003034">
        <property type="entry name" value="SAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036361">
        <property type="entry name" value="SAP_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15683:SF5">
        <property type="entry name" value="SAFB-LIKE TRANSCRIPTION MODULATOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15683">
        <property type="entry name" value="SCAFFOLD ATTACHMENT FACTOR B-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00076">
        <property type="entry name" value="RRM_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02037">
        <property type="entry name" value="SAP"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="SAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54928">
        <property type="entry name" value="RNA-binding domain, RBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF68906">
        <property type="entry name" value="SAP domain"/>
        <property type="match status" value="1"/>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0678">Repressor</keyword>
    <keyword evidence="3 9" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="6" id="KW-0804">Transcription</keyword>
    <keyword evidence="4" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="SAP" evidence="12">
        <location>
            <begin position="26"/>
            <end position="60"/>
        </location>
    </feature>
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        <location>
            <begin position="288"/>
            <end position="366"/>
        </location>
    </feature>
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            <begin position="1"/>
            <end position="25"/>
        </location>
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        </location>
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            <begin position="359"/>
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        </location>
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        <location>
            <begin position="619"/>
            <end position="938"/>
        </location>
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            <begin position="1"/>
            <end position="19"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="10">
        <location>
            <begin position="94"/>
            <end position="105"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="119"/>
            <end position="133"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
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            <begin position="174"/>
            <end position="187"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="194"/>
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        </location>
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            <begin position="247"/>
            <end position="257"/>
        </location>
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        </location>
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            <begin position="275"/>
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    </feature>
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            <begin position="362"/>
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    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
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    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="624"/>
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        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="665"/>
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        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="675"/>
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    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
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        <location>
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            <begin position="824"/>
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            <begin position="900"/>
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    </feature>
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        <location>
            <begin position="926"/>
            <end position="938"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022491"/>
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    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00071318"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00176"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50102"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50800"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030199.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="938" mass="106589" checksum="0FAFC05890BD4220" modified="2019-05-08" version="1">MAAAAAVSSPAAAPSVGPAQMESKKITDLRVIDLKSELKRRNLDITGVKTVLISRLKQAIEEEGGDPDNIEITVSADTPIKKPTKGKGKKQETDELSGDASVEDDSFVKNGEEEENEKEESLAEADHSAHEEMEANTSMKEAEDDNISVTIQAEDAITLDFDGDDLLETGKNVKITDSEASKAKDGQDTIAQSLEKERKDYEMTENHKDGKKEDCVKGDPVKKEAREGSKKAESGDKEKDTLKKGPSSTGASGQAKSSAKESKESKTTSKDDKGSASSVSGSSGSSTRNLWVSGLSSNTKAADLKNLFGKYGKVLSAKVVTNARSPGAKCYGIVTMSSSTEVARCIAHLHRTELHGQQISVEKVKGDPSKKELKKEIDEKSGSGRNMGDKKTASIDKTSKTQSAKKEEKKSDKSEKKESKEAKKTEGKDEKNDNGSSGPNQESTKKTEEKKRISGKSPGQMVVLDQTKGDQGHTRTVRRGRFDKVSICKNKERFIQDKMKFREYRGRKDILPFEKMKEQRMREHMVRLERIRRAVELRRRRELAERERRERERIRIIREREERERLQRERERLEIERQKLERERMERERLERERIRIEQERRKEAERIAREREELRRQQQQLRYEQEKRNSLKRPRDVDHRRDDPYWNENKKMALDTDARFSHGSDYSRQQNRFNDFDHRERGRYPEGSSVPSSSFERRERFVNQSEGKKTRPTARREEPGFERYPKNFSESRRNEPPQPRNELRDTDRREVRGDRDERRTVIIHDRPEISHGRHPRETGSNPPRQTNWKNEGGISTDKRDARGERPERSGREVSGHVRSAAPGSRSSTSGYGSREGERGVIGERGSGGQHYNEDRHVVERHSRETGPRKEWHGPSSQGSGYHDARRMGDGRGGGGMMSPHTSNSSPINRVVQITGSSMQRGSGSGFKPFKSGPPRRF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQD3</accession>
    <name>A0A452IQD3_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="4">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IQD3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034252.1">
        <property type="protein sequence ID" value="ENSGAGP00000030172.1"/>
        <property type="gene ID" value="ENSGAGG00000021771.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000034253.1">
        <property type="protein sequence ID" value="ENSGAGP00000030173.1"/>
        <property type="gene ID" value="ENSGAGG00000021771.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005109">
        <property type="term" value="F:frizzled binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060070">
        <property type="term" value="P:canonical Wnt signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035556">
        <property type="term" value="P:intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd04438">
        <property type="entry name" value="DEP_dishevelled"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd06717">
        <property type="entry name" value="PDZ_Dishevelled-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.10:FF:000040">
        <property type="entry name" value="segment polarity protein dishevelled homolog DVL-3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000014">
        <property type="entry name" value="Segment polarity protein dishevelled homolog DVL-3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.42.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.10">
        <property type="entry name" value="Winged helix-like DNA-binding domain superfamily/Winged helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="DEP_dom"/>
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        <property type="entry name" value="Dishevelled_C-dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008339">
        <property type="entry name" value="Dishevelled_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003351">
        <property type="entry name" value="Dishevelled_protein_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015506">
        <property type="entry name" value="Dsh/Dvl-rel"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001478">
        <property type="entry name" value="PDZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036034">
        <property type="entry name" value="PDZ_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036388">
        <property type="entry name" value="WH-like_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036390">
        <property type="entry name" value="WH_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10878">
        <property type="entry name" value="SEGMENT POLARITY PROTEIN DISHEVELLED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10878:SF5">
        <property type="entry name" value="SEGMENT POLARITY PROTEIN DISHEVELLED HOMOLOG DVL-1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="DEP"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Dsh_C"/>
        <property type="match status" value="1"/>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01760">
        <property type="entry name" value="DISHEVELLED"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01761">
        <property type="entry name" value="DISHEVELLED1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50156">
        <property type="entry name" value="PDZ domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46785">
        <property type="entry name" value="Winged helix' DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="DEP"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="PDZ" evidence="2">
        <location>
            <begin position="152"/>
            <end position="224"/>
        </location>
    </feature>
    <feature type="domain" description="DEP" evidence="3">
        <location>
            <begin position="301"/>
            <end position="375"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="50"/>
            <end position="138"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="429"/>
            <end position="571"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="64"/>
            <end position="74"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="1">
        <location>
            <begin position="77"/>
            <end position="93"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="1">
        <location>
            <begin position="102"/>
            <end position="115"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="1">
        <location>
            <begin position="116"/>
            <end position="129"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="1">
        <location>
            <begin position="437"/>
            <end position="460"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="461"/>
            <end position="474"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="PROSITE" id="PS50106"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS50186"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030172.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="586" mass="64454" checksum="8A2CEB97FBA4E1F3" modified="2019-05-08" version="1">MTMQSYRASMEEWCPGWSWLKAPTPTRAPSALKATRTSLLPSREQVALETHGLPRFIPRLNGHPKLDRHREHPPGYDSSSTMMSSELESSSFIDSDDDDNTSRLSSSTEQSTSSRLIRKHKRRRRKQRMRQIDRSSSFSSITDSTMSLNIITVTLNMEKYNFLGISIVGQSNDRGDGGIYIGSIMKGGAVAADGRIEPGDMLLQVNEVNFENMSNDDAVRVLREIVSKPGPISLTVAKCWDPTPRSYFTIPRAEPVRPIDPAAWISHTTAMTGAYPRYELEESPLTVKSDMATIVKVMQLPDSGLEIRDRMWLKITISNAVIGADVVDWLYTHVEGFKDRREARKYASSMLKHGYLRHTVNKITFSEQCYYVFGDLCSNMAALNLNNGSSGASDQDTLAPLPHPAAPWPLGQGYPYQYPLPPPCFPPAYQDPGFSYGSGSAGSQQSEGSKSSGSNRSNSENSRRALGRDKDRKSAGSGSESDHTACSGGGASVVRRERPVSQLSHRSHVSATHSERSHHSHHSYGPPGVPPLYNLPKLGSKVYGTSGPPGGPPVRELSSVPPELTGSRQSFQKAMGNPCEFFVDIM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQD4</accession>
    <name>A0A452IQD4_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="20">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Cell cortex</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the FNBP1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQD4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034125.1">
        <property type="protein sequence ID" value="ENSGAGP00000030054.1"/>
        <property type="gene ID" value="ENSGAGG00000021677.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005938">
        <property type="term" value="C:cell cortex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008289">
        <property type="term" value="F:lipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006897">
        <property type="term" value="P:endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd07676">
        <property type="entry name" value="F-BAR_FBP17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd11629">
        <property type="entry name" value="HR1_FBP17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1270.60:FF:000002">
        <property type="entry name" value="Formin-binding protein 1-like isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.40:FF:000017">
        <property type="entry name" value="Formin-binding protein 1-like isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.470">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.60">
        <property type="entry name" value="Arfaptin homology (AH) domain/BAR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027267">
        <property type="entry name" value="AH/BAR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031160">
        <property type="entry name" value="F_BAR_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001060">
        <property type="entry name" value="FCH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037449">
        <property type="entry name" value="FNBP1_F-BAR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011072">
        <property type="entry name" value="HR1_rho-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057870">
        <property type="entry name" value="HR1_TOCA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15735">
        <property type="entry name" value="FCH AND DOUBLE SH3 DOMAINS PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15735:SF13">
        <property type="entry name" value="FORMIN-BINDING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00611">
        <property type="entry name" value="FCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25610">
        <property type="entry name" value="HR1_TOCA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00018">
        <property type="entry name" value="SH3_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00055">
        <property type="entry name" value="FCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00326">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103657">
        <property type="entry name" value="BAR/IMD domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51741">
        <property type="entry name" value="F_BAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51860">
        <property type="entry name" value="REM_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="9 14 15" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="12" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="8" id="KW-0254">Endocytosis</keyword>
    <keyword evidence="10" id="KW-0446">Lipid-binding</keyword>
    <keyword evidence="11" id="KW-0472">Membrane</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 13" id="KW-0728">SH3 domain</keyword>
    <feature type="domain" description="F-BAR" evidence="18">
        <location>
            <begin position="1"/>
            <end position="264"/>
        </location>
    </feature>
    <feature type="domain" description="REM-1" evidence="19">
        <location>
            <begin position="338"/>
            <end position="415"/>
        </location>
    </feature>
    <feature type="domain" description="SH3" evidence="17">
        <location>
            <begin position="484"/>
            <end position="545"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="293"/>
            <end position="345"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="423"/>
            <end position="467"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="15">
        <location>
            <begin position="345"/>
            <end position="405"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="429"/>
            <end position="446"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004544"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00009426"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022583"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023121"/>
        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01077"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS51741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS51860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030054.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="555" mass="64572" checksum="AA06E7143C730609" modified="2019-05-08" version="1">MSWGTELWDQFDNLEKHTQWGIDVLEKYIKFVKERTEIEISYAKQLRNLSKKYQPKKNSKEEEEYKYTSCKAFLATLNEMNDYAGQHEVISENMTSQITAELARFVQELKQERKSHFHDGRKAQQYIETFWKQLEASKRRFERDCKEADRAQQYFEKMDADINVTKADVEKARQQAQIRHQMAEDSKAEYSSILQKFNHDQHEHYHTHIPNIFQKIQEMEERRIVRIGECMKTYAEVDRQVIPIIGKCLDGITKAAESIDQKNDSLLVIEAFKSGFEPPGDIDFEDFTQPMKRTVSESSLSNSRWDGKSEPKPGSKSKGKLWPFIKKNKGGGPEDFSNLPPEQRRKKLQQKVDELNKEIQKEMDQRDALTKMKDVYIKNPQMGDAASVDHKLAELGQNMEKLRLEAQKFEGWLAEVEGRLPVRTEQSRRQSGMYDTQNTSGVNNCAQDRESPDGSYTEEPSQESEMKVPVTEFDDEFDDEEPLPTIGTCKALYTFEGQNEGTISVMEGEMLYVIEEDKGDGWTRIRRNEDEEGYVPTSYVEVYLDKNAKGAMTYI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQD6</accession>
    <name>A0A452IQD6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">SH2 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell junction</location>
            <location evidence="1">Focal adhesion</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the PTEN phosphatase protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQD6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034415.1">
        <property type="protein sequence ID" value="ENSGAGP00000030322.1"/>
        <property type="gene ID" value="ENSGAGG00000021831.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005925">
        <property type="term" value="C:focal adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="CDD" id="cd01213">
        <property type="entry name" value="PTB_tensin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.505.10">
        <property type="entry name" value="SH2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013625">
        <property type="entry name" value="PTB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006020">
        <property type="entry name" value="PTB/PI_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000980">
        <property type="entry name" value="SH2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036860">
        <property type="entry name" value="SH2_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033929">
        <property type="entry name" value="Tensin_PTB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051484">
        <property type="entry name" value="Tensin_PTEN_phosphatase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45734">
        <property type="entry name" value="TENSIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45734:SF6">
        <property type="entry name" value="TENSIN-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08416">
        <property type="entry name" value="PTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00017">
        <property type="entry name" value="SH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00462">
        <property type="entry name" value="PTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00252">
        <property type="entry name" value="SH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55550">
        <property type="entry name" value="SH2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50001">
        <property type="entry name" value="SH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 4" id="KW-0727">SH2 domain</keyword>
    <feature type="domain" description="SH2" evidence="6">
        <location>
            <begin position="442"/>
            <end position="548"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="61"/>
            <end position="91"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="116"/>
            <end position="143"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="261"/>
            <end position="308"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="331"/>
            <end position="354"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="384"/>
            <end position="430"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="261"/>
            <end position="288"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="5">
        <location>
            <begin position="331"/>
            <end position="345"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="5">
        <location>
            <begin position="396"/>
            <end position="409"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="410"/>
            <end position="423"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007881"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022999"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00191"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50001"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030322.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="689" mass="73982" checksum="34DC28F4320916B8" modified="2019-05-08" version="1">MSQIIPSHVLRVGQTIRVASQEESRSLQPAGSRCCSSLTTKCSYYSAEGWLSQPMMAQTKQVASMERASPTHQAEGEEETEGNSKASPVSPTLDVSIESLNQLILEIDPTFQPLLLKPDSSKKHPAQSMSPGNAAVSEKPDPDSLEIKYIEMTPTRAKSHESMQGSASPCITPLSMSPQGSCFLLARDNCTPNGSLVFSTPQNQPPTGSSPQTSYRASECVSIPQRGQKDCISYGPNTLLATSPGLESLRKALYIGRAQQRTSRVSMLSTSPGSDTSYVLGSTQSLLNDDSDSYQPAHRSAESPLSVGSFGSPCLTSCSFQVDRFGDAFSPGSNSGKANNSSSPPLQKGHASSCPPSIVNSLADIPILLVNGCLEHGETSPRMGKDFHGSVKQTVPLSSSPSSGINSLNKTSSETSLSFSPDGSSKDGEPTMKFVMDTSKYWFKPSITRDQAIQLLKEAEPGAFLMRDSTSYRGSFGLAMKVLSSPSELNSPGESYTDLVRHFLIESSAKGVHLKGASEEPYFGSLSAFVYQHAMTALALPCKLAIPTRGGRRPSVALTLVLCCSLAACNVLYLHSVSVETLTGASAVQKAVSSTFELETPPTPTIVHFKVTKQGITLTDVQRKVFFRRHYPLAALSFCCMDPENRKIFGFVAKSQTDAENLCHLFAEYDTVQPASLVIDFISKLLPEQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQD7</accession>
    <name>A0A452IQD7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Homeobox domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 9 10">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQD7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034192.1">
        <property type="protein sequence ID" value="ENSGAGP00000030115.1"/>
        <property type="gene ID" value="ENSGAGG00000021725.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00086">
        <property type="entry name" value="homeodomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.60:FF:000142">
        <property type="entry name" value="homeobox protein OTX2 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.60">
        <property type="entry name" value="Homeodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001356">
        <property type="entry name" value="HD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017970">
        <property type="entry name" value="Homeobox_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009057">
        <property type="entry name" value="Homeodomain-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003025">
        <property type="entry name" value="Otx_TF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013851">
        <property type="entry name" value="Otx_TF_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45793:SF22">
        <property type="entry name" value="CONE-ROD HOMEOBOX PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45793">
        <property type="entry name" value="HOMEOBOX PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00046">
        <property type="entry name" value="Homeodomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03529">
        <property type="entry name" value="TF_Otx"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01255">
        <property type="entry name" value="OTXHOMEOBOX"/>
    </dbReference>
    <dbReference type="SMART" id="SM00389">
        <property type="entry name" value="HOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46689">
        <property type="entry name" value="Homeodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00027">
        <property type="entry name" value="HOMEOBOX_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50071">
        <property type="entry name" value="HOMEOBOX_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="3" id="KW-0221">Differentiation</keyword>
    <keyword evidence="5 9 10" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="6 9 10" id="KW-0371">Homeobox</keyword>
    <keyword evidence="8 9 10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0804">Transcription</keyword>
    <keyword evidence="4" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="Homeobox" evidence="12">
        <location>
            <begin position="14"/>
            <end position="74"/>
        </location>
    </feature>
    <feature type="DNA-binding region" description="Homeobox" evidence="9">
        <location>
            <begin position="16"/>
            <end position="75"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="71"/>
            <end position="123"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="73"/>
            <end position="84"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="113"/>
            <end position="123"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023155"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00108"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="RuleBase" id="RU000682"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50071"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030115.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="268" mass="28850" checksum="7140E04DC9F46F9F" modified="2019-05-08" version="1">PCLLHPNAGSATPRKQRRERTTFTRAQLDILEALFAKTRYPDIFMREEVALKINLPESRVQVWFKNRRAKCRQQQQQSSGQAKARPAKKKSSPARETSSETSTNGQYSPPPAGTSGTPSSTASATVSIWSPASISPIPDPLASSATPCMQRSAAYPMTYTQAAGYPQSYAGSSSYFTGLDCGSYLSPMHPQLSAPGAALSPIATPAMGSHLSQSPASLSSQGFGTAGLSFGSVDCLDYKDQTGSWKLNFNATDCLDYKDQSSWKFQVL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQD8</accession>
    <name>A0A452IQD8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="19">F-box DNA helicase 1</fullName>
            <ecNumber evidence="17">5.6.2.4</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="21">DNA 3'-5' helicase 1</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="20">F-box only protein 18</fullName>
        </alternativeName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>ATP + H2O = ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:13065"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="5.6.2.4"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="16">
            <text>Couples ATP hydrolysis with the unwinding of duplex DNA by translocating in the 3'-5' direction.</text>
            <dbReference type="EC" id="5.6.2.4"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="3">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Chromosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the helicase family. UvrD subfamily.</text>
    </comment>
    <dbReference type="EC" id="5.6.2.4" evidence="17"/>
    <dbReference type="AlphaFoldDB" id="A0A452IQD8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034257.1">
        <property type="protein sequence ID" value="ENSGAGP00000030177.1"/>
        <property type="gene ID" value="ENSGAGG00000021773.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005694">
        <property type="term" value="C:chromosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043138">
        <property type="term" value="F:3'-5' DNA helicase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016787">
        <property type="term" value="F:hydrolase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000724">
        <property type="term" value="P:double-strand break repair via homologous recombination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031297">
        <property type="term" value="P:replication fork processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd22095">
        <property type="entry name" value="F-box_FBXO18"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18786">
        <property type="entry name" value="SF1_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1280.50:FF:000011">
        <property type="entry name" value="F-box DNA helicase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1280.50">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014017">
        <property type="entry name" value="DNA_helicase_UvrD-like_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000212">
        <property type="entry name" value="DNA_helicase_UvrD/REP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036047">
        <property type="entry name" value="F-box-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001810">
        <property type="entry name" value="F-box_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014016">
        <property type="entry name" value="UvrD-like_ATP-bd"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11070:SF30">
        <property type="entry name" value="F-BOX DNA HELICASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11070">
        <property type="entry name" value="UVRD / RECB / PCRA DNA HELICASE FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12937">
        <property type="entry name" value="F-box-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00580">
        <property type="entry name" value="UvrD-helicase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13361">
        <property type="entry name" value="UvrD_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00256">
        <property type="entry name" value="FBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81383">
        <property type="entry name" value="F-box domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50181">
        <property type="entry name" value="FBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="11" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="5" id="KW-0158">Chromosome</keyword>
    <keyword evidence="7" id="KW-0227">DNA damage</keyword>
    <keyword evidence="13" id="KW-0234">DNA repair</keyword>
    <keyword evidence="12" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="10" id="KW-0347">Helicase</keyword>
    <keyword evidence="9" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="14" id="KW-0413">Isomerase</keyword>
    <keyword evidence="6" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="15" id="KW-0539">Nucleus</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="F-box" evidence="22">
        <location>
            <begin position="99"/>
            <end position="148"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00009922"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022454"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022806"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023235"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00034617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00034808"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00048988"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00071173"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00075040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00079567"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS50181"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030177.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="943" mass="106342" checksum="CD66CECCA65B9397" modified="2019-05-08" version="1">MDEGLEDHCALLGCDGDAKTAPWGPVASRKRPLCSEGAGSNYTEYSKCGGTDKKPMIDSSLKYSPGHTEIRQDPDVIEIDPIPDSYFGLLGTSCGKEPQGHIDQLPNEVLSIIFAFLPVTDLCLNLNLVCHRWREIISDPLFILWKKLYRQYLMKEAQALLTVEWILDHYGITQEQDQCMLGFIRYMATSNAARRMDPDAVLRCLKKHHLFTKAEICIIQRLPDLEGTAGAVNVWAVIAAIVLFSDGVHDIQKLMACLRKPGSTLSLLDITEMLYCMATLLNAMREKGINISNRFHYSIFYSLYLMENACPVTKTTMLETPSSNCGRGAWPSNKLDIKLTHEQQRILNHTILPGQVVKIMAFAGTGKTSTLVKYAEKFPHLSFLYVAFNKAIVENGKSVFPPNVMCKTFHSLAYTSVGKHYNEKGKLNFSNLSPYTISFILKHRMGQSLFIRAKTVTQTLESFFASSDEFISVEHTPIWCKNTHGMRVLVSEEEKQIIVEEARRIWDNMKMLNGNTELNCRMTSDGYLKLWQLRKPRLFAYDAIFVDEAQDCTPAIMDIVLSQSCGIIFVGDPHQQIYTFRGAVNALFAVPHTHIFYLTKSFRFGPEIAYVGSTILDVCKKIRNKMLVGGNQEGDVRGSMNGKTAFLSRGNLNVFENAVTITGGERPTKIHLLGGPKNFGLDKIHDIWKLLQPEEARVRLKLQIDDHFIKKWVEKDGFQGLKNYAAKTEDRELEVKITIVEKYNVRIPELVARIKNSHVSNIAFADYVIGTVYKAKGLEFDTVLIADDFMKVSCARHNLQRLPRFIPDMSAEDEWNLLYVAVTRAKKHLLMTKSLEYLLTLAGEYFLRPELTSEALKEGAIHCSVSQCHNAVPADSILTMKKVPLTYSNGSKDQGGYLCHACVQQRLGPLTQLTISPELVQTMDVVDEVLEIPRHLAVLYGLF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQD9</accession>
    <name>A0A452IQD9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Mitogen-activated protein kinase kinase kinase kinase</fullName>
            <ecNumber evidence="9">2.7.11.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Serine/threonine kinase that plays a role in the response to environmental stress. Appears to act upstream of the JUN N-terminal pathway.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="18 19">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="20 19">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 9">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="similarity">
        <text evidence="2 9">Belongs to the protein kinase superfamily. STE Ser/Thr protein kinase family. STE20 subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="9"/>
    <dbReference type="AlphaFoldDB" id="A0A452IQD9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034131.1">
        <property type="protein sequence ID" value="ENSGAGP00000030059.1"/>
        <property type="gene ID" value="ENSGAGG00000021689.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008349">
        <property type="term" value="F:MAP kinase kinase kinase kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0106310">
        <property type="term" value="F:protein serine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="RHEA"/>
    </dbReference>
    <dbReference type="CDD" id="cd06613">
        <property type="entry name" value="STKc_MAP4K3_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000031">
        <property type="entry name" value="Mitogen-activated protein kinase kinase kinase kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001180">
        <property type="entry name" value="CNH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021160">
        <property type="entry name" value="MAPKKKK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050629">
        <property type="entry name" value="STE20/SPS1-PAK"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48012:SF6">
        <property type="entry name" value="MITOGEN-ACTIVATED PROTEIN KINASE KINASE KINASE KINASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48012">
        <property type="entry name" value="STERILE20-LIKE KINASE, ISOFORM B-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00780">
        <property type="entry name" value="CNH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF038172">
        <property type="entry name" value="MAPKKKK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00036">
        <property type="entry name" value="CNH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50219">
        <property type="entry name" value="CNH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 9 11" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="7 9" id="KW-0418">Kinase</keyword>
    <keyword evidence="6 9 11" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 9" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="5 9" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Protein kinase" evidence="14">
        <location>
            <begin position="16"/>
            <end position="273"/>
        </location>
    </feature>
    <feature type="domain" description="CNH" evidence="15">
        <location>
            <begin position="506"/>
            <end position="819"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="425"/>
            <end position="463"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="10">
        <location>
            <position position="136"/>
        </location>
    </feature>
    <feature type="binding site" evidence="11">
        <location>
            <begin position="22"/>
            <end position="30"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="11 12">
        <location>
            <position position="45"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008874"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PIRNR" id="PIRNR038172"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PIRSR" id="PIRSR038172-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PIRSR" id="PIRSR038172-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50219"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030059.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="846" mass="94671" checksum="1521D4FB88C40531" modified="2019-05-08" version="1">MSLLRDVSLLDPRLRYELIQRIGAGTYGDVYKARDTQTAELAAVKIIKLDPGDDISSIQQEITTLRDCGHPNVVAYFGSYLRNDRLWICMEFCGGGSLQEIYHTTGPLSEKQIAYVCRETLQGLSHLHAKGKMHRDIKGANILLTLFGDVKLADFGVSAELTASVAKRKSFIGTPYWMAPEVAAVEKKGGYNQLCDIWAIGITAIELAELQPPLFDLHPMRALMLMSKSSFQPPKLKDKAHWSPEFHQFLKLALTKNPKKRPTAEKLLQHLFTSQPLARTLAIELLDKVNSPDLAAPSLDDGDFEVYDIFPDKIHSCGAHGQAERTLSEIQFNQVKFGPPLRKETEPYCGLREEEDDWTLLGAEESLLQSVEEALEERSLTIRPASGRVPTPEDQPVDSHCIRENETGTIKRGSLWDLMNWELGGMEPGGGSETMRRQAPAGSPESGDIQPPLHTGSPEPGASLLSTEWATMKRREETERSSCHGLPPTPKVHMGACFSKVFNGCPLKINSAVTWIHSETRDQYLLVGAEEGIYTLNLHELHEDTMEKLLPHRCSWLYCMNNMLLSLAGKSSQISSHNLLGLFEQRRQLQKRQVPLSIATNRLTERIIPRKFALSSKIADTKGCLKCRVVRNPYSGNTFLCAALPSSLVLLQWYEPLQKFMLLKHFPATLPMPLSLFELLVVETEEYPQVCVRVVDRDRPGQELEFNVLSLSTSSSLSTEPSADVASSVACHMTQMDRDTVLVCFEHYVRTVNLRGVPKRALAPELTFNFPIETIVCLQDSVLAFWKHGMQGRSLEGNEVTQEITDKSRVFRVLGANRDIILESTPTENPLAHSNLYILTGHESSY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQE1</accession>
    <name>A0A452IQE1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Peptidase S1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IQE1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034290.1">
        <property type="protein sequence ID" value="ENSGAGP00000030209.1"/>
        <property type="gene ID" value="ENSGAGG00000021723.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030141">
        <property type="term" value="C:secretory granule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00190">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.10:FF:000120">
        <property type="entry name" value="Putative serine protease"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.10.10">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009003">
        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043504">
        <property type="entry name" value="Peptidase_S1_PA_chymotrypsin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001314">
        <property type="entry name" value="Peptidase_S1A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001254">
        <property type="entry name" value="Trypsin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018114">
        <property type="entry name" value="TRYPSIN_HIS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033116">
        <property type="entry name" value="TRYPSIN_SER"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271:SF48">
        <property type="entry name" value="KALLIKREIN-14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271">
        <property type="entry name" value="KALLIKREIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00089">
        <property type="entry name" value="Trypsin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00722">
        <property type="entry name" value="CHYMOTRYPSIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00020">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50494">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50240">
        <property type="entry name" value="TRYPSIN_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00134">
        <property type="entry name" value="TRYPSIN_HIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00135">
        <property type="entry name" value="TRYPSIN_SER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3 6" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="1 6" id="KW-0645">Protease</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 6" id="KW-0720">Serine protease</keyword>
    <keyword evidence="2" id="KW-0732">Signal</keyword>
    <feature type="domain" description="Peptidase S1" evidence="7">
        <location>
            <begin position="28"/>
            <end position="261"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="RuleBase" id="RU363034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030209.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="263" mass="28622" checksum="8409E7EAC3ECE6AD" modified="2019-05-08" version="1">SPALLCLQFTPVVNLSACLPVSSQNNRIVGGAPCIPHSMPWQAALFMGSQLNCGGTLIDRNWVVTAAHCHCGCACCEGRGCVIVRVSISVRLGEHNIRHLDWTEQLRLSEKTILHPGYNSASKDNDIMLIKLLTPANLNKNVQPLELPTSCPSPKEECVVSGWGTTKMYFPPVLHCGKVSIFSQEQCRSIYPRYYTENMLCAGVLEGGIDSCQGDSGGPLLCNGKLQGIVSWGTQTCALPNKPGVYVAVCKYVDWIRETINNN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQE5</accession>
    <name>A0A452IQE5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">SH2 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell junction</location>
            <location evidence="1">Focal adhesion</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the PTEN phosphatase protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQE5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034427.1">
        <property type="protein sequence ID" value="ENSGAGP00000030333.1"/>
        <property type="gene ID" value="ENSGAGG00000021831.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005925">
        <property type="term" value="C:focal adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="CDD" id="cd01213">
        <property type="entry name" value="PTB_tensin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.505.10">
        <property type="entry name" value="SH2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013625">
        <property type="entry name" value="PTB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006020">
        <property type="entry name" value="PTB/PI_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000980">
        <property type="entry name" value="SH2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036860">
        <property type="entry name" value="SH2_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033929">
        <property type="entry name" value="Tensin_PTB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051484">
        <property type="entry name" value="Tensin_PTEN_phosphatase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45734">
        <property type="entry name" value="TENSIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45734:SF6">
        <property type="entry name" value="TENSIN-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08416">
        <property type="entry name" value="PTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00017">
        <property type="entry name" value="SH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00462">
        <property type="entry name" value="PTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00252">
        <property type="entry name" value="SH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55550">
        <property type="entry name" value="SH2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50001">
        <property type="entry name" value="SH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 4" id="KW-0727">SH2 domain</keyword>
    <feature type="domain" description="SH2" evidence="6">
        <location>
            <begin position="434"/>
            <end position="535"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="66"/>
            <end position="85"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="112"/>
            <end position="138"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="190"/>
            <end position="224"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="265"/>
            <end position="313"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="336"/>
            <end position="358"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="72"/>
            <end position="85"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="190"/>
            <end position="221"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="265"/>
            <end position="293"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="5">
        <location>
            <begin position="336"/>
            <end position="350"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007881"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022999"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00191"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50001"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030333.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="680" mass="72945" checksum="86253AA513546D82" modified="2019-05-08" version="1">MSQIIPSHVLRVGQTIRVASQEESRSLQPAGSRCCSSLTTKCSYYSAEGWLSQPMMAQTKAHLLPNGGLHGANSNAGNSKASPVSPTLDVSIESLNQLILEIDPTFQPLLLKPDSSKKHPAQSMSPGNAAVSEKPDPDSLEIKYIEMTPTRAKSHESMQGSASPCITPLSMSPQGSCFLLARDNCTPNGSLVFSSPSRPDSNRPAPQNQPPTGSSPQTSYRASECVSIPQRGQKDCISYGPNTLLATSPGLESLRKALYIGRAQQRTSRVSMLSTSPGSDTSYVLGSTQSLLNDDSDSYQPAHRSAESPLSVGSFGSPCLTSCSFQVDRFGDAFSPGSNSGKANNSSSPPLQKGHASSCPPSIVNSLADIPILLVNGCLEHGETSPRMGKDFHGSVKQTVRSCVMHEGMPLSLAGSSKDGEPTMKFVMDTSKYWFKPSITRDQAIQLLKEAEPGAFLMRDSTSYRGSFGLAMKVLSSPSGDDSSDLVRHFLIESSAKGVHLKGASEEPYFGSLSAFVYQHAMTALALPCKLAIPTRGTGPTWRPTPLESAISAMKKTAACNVLYLHSVSVETLTGASAVQKAVSSTFELETPPTPTIVHFKVTKQGITLTDVQRKVFFRRHYPLAALSFCCMDPENRKIFGFVAKSQTDAENLCHLFAEYDTVQPASLVIDFISKLLPEQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQE7</accession>
    <name>A0A452IQE7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Peptidase S1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IQE7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034295.1">
        <property type="protein sequence ID" value="ENSGAGP00000030214.1"/>
        <property type="gene ID" value="ENSGAGG00000021723.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030141">
        <property type="term" value="C:secretory granule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00190">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.10.10">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009003">
        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043504">
        <property type="entry name" value="Peptidase_S1_PA_chymotrypsin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001314">
        <property type="entry name" value="Peptidase_S1A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001254">
        <property type="entry name" value="Trypsin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018114">
        <property type="entry name" value="TRYPSIN_HIS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033116">
        <property type="entry name" value="TRYPSIN_SER"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271:SF48">
        <property type="entry name" value="KALLIKREIN-14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271">
        <property type="entry name" value="KALLIKREIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00089">
        <property type="entry name" value="Trypsin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00722">
        <property type="entry name" value="CHYMOTRYPSIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00020">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50494">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50240">
        <property type="entry name" value="TRYPSIN_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00134">
        <property type="entry name" value="TRYPSIN_HIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00135">
        <property type="entry name" value="TRYPSIN_SER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="2 5" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="1 5" id="KW-0645">Protease</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 5" id="KW-0720">Serine protease</keyword>
    <feature type="domain" description="Peptidase S1" evidence="7">
        <location>
            <begin position="41"/>
            <end position="264"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="1"/>
            <end position="21"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="RuleBase" id="RU363034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030214.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="266" mass="28939" checksum="441AC63D22214564" modified="2025-10-08" version="2">MQRVRWSRGFGGPPAGVPASPPSPVVNLSACLPVSSQNNRIVGGAPCIPHSMPWQAALFMGSQLNCGGTLIDRNWVVTAAHCHVQCSISVRLGEHNIRHLDWTEQLRLSEKTILHPGYNSASKDNDIMLIKLLTPANLNKNVQPLELPTSCPSPKEECVVSGWGTTKSPQLYFPPVLHCGKVSIFSQEQCRSIYPRYYTENMLCAGVLEGGIDSCQGDSGGPLLCNGKLQGIVSWGTQTCALPNKPGVYVAVCKYVDWIRETINNN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQF1</accession>
    <name>A0A452IQF1_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="19">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the G-protein coupled receptor 2 family. Adhesion G-protein coupled receptor (ADGR) subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQF1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034300.1">
        <property type="protein sequence ID" value="ENSGAGP00000030219.1"/>
        <property type="gene ID" value="ENSGAGG00000021690.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098794">
        <property type="term" value="C:postsynapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043083">
        <property type="term" value="C:synaptic cleft"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004930">
        <property type="term" value="F:G protein-coupled receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007189">
        <property type="term" value="P:adenylate cyclase-activating G protein-coupled receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007166">
        <property type="term" value="P:cell surface receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016525">
        <property type="term" value="P:negative regulation of angiogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016322">
        <property type="term" value="P:neuron remodeling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="entry name" value="Adhesion G protein-coupled receptor B2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000003">
        <property type="entry name" value="Adhesion G protein-coupled receptor B2"/>
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    </dbReference>
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        <property type="entry name" value="Adhesion G protein-coupled receptor B2"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Adhesion G protein-coupled receptor B2"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Adhesion G protein-coupled receptor B2"/>
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        <property type="entry name" value="Adhesion G protein-coupled receptor B3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.610">
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.1240.10">
        <property type="entry name" value="GPCR, family 2, extracellular hormone receptor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1070.10">
        <property type="entry name" value="Rhodopsin 7-helix transmembrane proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.20.100.10">
        <property type="entry name" value="Thrombospondin type-1 (TSP1) repeat"/>
        <property type="match status" value="3"/>
    </dbReference>
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        <property type="entry name" value="AGRL2-4_GAIN_subdom_A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057244">
        <property type="entry name" value="GAIN_B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046338">
        <property type="entry name" value="GAIN_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017981">
        <property type="entry name" value="GPCR_2-like_7TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008077">
        <property type="entry name" value="GPCR_2_brain_angio_inhib"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036445">
        <property type="entry name" value="GPCR_2_extracell_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001879">
        <property type="entry name" value="GPCR_2_extracellular_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000832">
        <property type="entry name" value="GPCR_2_secretin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017983">
        <property type="entry name" value="GPCR_2_secretin-like_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000203">
        <property type="entry name" value="GPS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000884">
        <property type="entry name" value="TSP1_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036383">
        <property type="entry name" value="TSP1_rpt_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12011:SF40">
        <property type="entry name" value="ADHESION G PROTEIN-COUPLED RECEPTOR B3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12011">
        <property type="entry name" value="ADHESION G-PROTEIN COUPLED RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00002">
        <property type="entry name" value="7tm_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16489">
        <property type="entry name" value="GAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01825">
        <property type="entry name" value="GPS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02793">
        <property type="entry name" value="HRM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00090">
        <property type="entry name" value="TSP_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01694">
        <property type="entry name" value="BAIPRECURSOR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00249">
        <property type="entry name" value="GPCRSECRETIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00303">
        <property type="entry name" value="GPS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00008">
        <property type="entry name" value="HormR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00209">
        <property type="entry name" value="TSP1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF82895">
        <property type="entry name" value="TSP-1 type 1 repeat"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00650">
        <property type="entry name" value="G_PROTEIN_RECEP_F2_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50227">
        <property type="entry name" value="G_PROTEIN_RECEP_F2_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50261">
        <property type="entry name" value="G_PROTEIN_RECEP_F2_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50221">
        <property type="entry name" value="GAIN_B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50092">
        <property type="entry name" value="TSP1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="9" id="KW-0297">G-protein coupled receptor</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10 15" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="12" id="KW-0675">Receptor</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="14" id="KW-0807">Transducer</keyword>
    <keyword evidence="5 15" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 15" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="16"/>
            <end position="35"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="531"/>
            <end position="553"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="565"/>
            <end position="583"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="595"/>
            <end position="613"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="634"/>
            <end position="653"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="673"/>
            <end position="696"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="751"/>
            <end position="772"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="778"/>
            <end position="799"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="806"/>
            <end position="825"/>
        </location>
    </feature>
    <feature type="domain" description="G-protein coupled receptors family 2 profile 1" evidence="17">
        <location>
            <begin position="203"/>
            <end position="273"/>
        </location>
    </feature>
    <feature type="domain" description="GAIN-B" evidence="16">
        <location>
            <begin position="395"/>
            <end position="521"/>
        </location>
    </feature>
    <feature type="domain" description="G-protein coupled receptors family 2 profile 2" evidence="18">
        <location>
            <begin position="529"/>
            <end position="803"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007343"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023224"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50227"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50261"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030219.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1111" mass="124905" checksum="6BFEBDF005543170" modified="2019-05-08" version="1">MNFTTATVLQKRLTRLWAKSPLGFFALFVFIFNLLCYRCPVCFVTVHGVWEEWSPWSLCSFTCGRGQRTRTRSCIPPQYGGRSCDGPETQHKPCNIALCPVDGQWQEWSSWSQCSVTCSNGTQQRSRQCTAAAHGGSECRGPWAESRECHNPECTANGQWNQWGHWSGCSKSCDGGWERRIRICQGAAVTGQQCEGTGEEIRRCSEQRCPAPYEICPEDYLLSMVWKRTPAGDMAFNRCPLNATGTASRRCSLSVHGVAFWEHPSFARCISNEYRHLQHSIKEHLAKGQRMLAGDGMSQVTKTLLDLTQRKNFYAGDLLISVEILRNVTDTFKRASYIPASDGVQNFFQIVSNLLDEENKEKWEDAQQIYPGSVELMQVIEDFIHIVGMGMMDFQNSYLMTGNVVASIQKLPAASVLTDINFPMKGRKGMVDWARNSEDRVVIPKNIFTPMSSELDESTVFVLGAVLYKNLELILPTLRNDSLGTWSTQGCKTVLTDASHTKCLCDRLSTFAILAQQPREIIMESSGAPSVTLIVGSGLSCLALITLAVVYAALWRYIRSERSIILINFCLSIISSNILILVGQTQTHNKGICTTTTAFLHFFFLASFCWVLTEAWQSYMAVTGKIRTQLIRKRFLCLGWGLPALVVAISIGFTKTKGYGTSHYCWLSLEGGLLYAFVGPAAAVVLVNMVIGILVFNKLVSRDGILDKKLKHRAGQMSEPHSGLTLKCAKCGVVSTTALSATTASNAMASLWSSCVVLPLLALTWMSAVLAMTDKRSILFQILFAVFDSLQGFVIVMVHCILRREVLWFGFNLIMAVLFHCFLLQTDFEKDVDIACRSVLHKDIGPCRAATITGTLSRISLNDDEEEKGANPEGMSYSTLPGSIISKVIIQQPAGLHMPMNNSELRRTVYLCTDDNLRGADMDIVNPQERMMESDYIVMPRGSVNTQQSLKDESKLNIGMDTLPHERLLHYKVNPEFNMNPSVMDQFNINLDQHLPTQEHMQNLPFEPRTAVKNFIASERKSRYSDLDFEKVMHTRKRHMELFQELNQKFQTLDRFRDVPKSGSMVSILLSSYEFKIYWFAKDALELRPAEWEKCLNLPLDVQEGDFQTEV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQF2</accession>
    <name>A0A452IQF2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">PDZ domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQF2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034141.1">
        <property type="protein sequence ID" value="ENSGAGP00000030069.1"/>
        <property type="gene ID" value="ENSGAGG00000021685.1"/>
    </dbReference>
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    <sequence length="817" mass="89108" checksum="4B771DA03E0171B0" modified="2019-05-08" version="1">MHFIRLSCLFPNCLEGTIKPGDRLLSVDGIRLLGTSHAEAMSILKQCGQEATLLIEYDVSVMDSVATASGPLLVEVAKTPGAALGVALTTSMCCNKQVIVIDKIKSASFADRCGALHVGDHILSIDGTSMEYCTLAEATQFLANTADNVKLEILPHHQTRLALKGAEHAALVSSSFSPTSMSAYSLSSLNMGTLPRSLYSTSPRGTMMRRRMKKKDFKSSLSLASSTVGLAGQVVHTETTEVVLTADPIVGFGIQLQGSVFATETLSSPPLISYIECDSPAERCGVLQIGDRLMAINGIPTEDSTFEEANQLLRDSSITNKVTLEIEFDVAESVIPSSGTFHVKLPKKHNVELGITISSPSSRKPGDPLVISDIKKGSVAHRTGTLELGDKLLAIDNIRLDNCSMEDAVQILQHCEDLVKLKIRKDEDNSDGEFEAIIMFIFKPYLAASPKKFSVSGHLSELSDAEDETSSTQKPGKLSDIYSTTVPSVDSAVESWDGSGIDTSFGSQGPTYQASGYNFSAYEWRSPKQRDNLSPPSRPQNHPFHDTGLSDDEWDRPTASGFSGAHDNSETDQEENFWSQALEDLETCGQSGILRELEDKADRRLYLRNMTLLATIMSGSTMSLNHENPQPRSQLGRQASFQERSNVRPHYSQTARSNTLPSDVGRKSVVLRKIKQEMKEIMSPTPVELHKVTLYKESDIEDFGFSVSDGLLEKGVYVKNIRPAGPGDLGGLKPYDRLLQVNHVRTRDFDCCLVVPLIAESGNKLELVISRNPLASQKVTSDQQTLAGGEWNDQNNAFLQQTGHTASVETRDPTNTI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQF3</accession>
    <name>A0A452IQF3_9SAUR</name>
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        <recommendedName>
            <fullName evidence="8">SAFB-like transcription modulator</fullName>
        </recommendedName>
    </protein>
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        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQF3"/>
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        <property type="protein sequence ID" value="ENSGAGP00000030193.1"/>
        <property type="gene ID" value="ENSGAGG00000021279.1"/>
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        <property type="project" value="UniProtKB-SubCell"/>
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        <property type="entry name" value="RRM_dom"/>
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        <property type="entry name" value="SCAFFOLD ATTACHMENT FACTOR B-RELATED"/>
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    <keyword evidence="6" id="KW-0804">Transcription</keyword>
    <keyword evidence="4" id="KW-0805">Transcription regulation</keyword>
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            <begin position="1"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00022491"/>
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            <dbReference type="ARBA" id="ARBA00022884"/>
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            <dbReference type="PROSITE-ProRule" id="PRU00176"/>
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            <dbReference type="PROSITE" id="PS50800"/>
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            <dbReference type="Ensembl" id="ENSGAGP00000030193.1"/>
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        </source>
    </evidence>
    <sequence length="988" mass="112065" checksum="2705B228588C88BB" modified="2019-05-08" version="1">MAAAAAVSSPAAAPSVGPAQMESKKITDLRVIDLKSELKRRNLDITGVKTVLISRLKQAIEEEGGDPDNIEITVSADTPIKKPTKGKGNASNASDQDGNDELKDFEEAGENEEENSHSKELRPSEKKKYSDSKQRKTAEDAEKDFESQVPYKAGSGDGIQEVSKPLPSEESLAEADHSAHEEMEANTSMKEAEDDNISVTIQAEDAITLDFDGDDLLETGKNVKITDSEASKAKDGQDTIAQSLEKERKDYEMTENHKDGKKEDCVKGDPVKKEAREGSKKAESGDKEKDTLKKGPSSTGASGQAKSSAKESKESKTTSKDDKGSASSVSGSSGSSTRNLWVSGLSSNTKAADLKNLFGKYGKVLSAKVVTNARSPGAKCYGIVTMSSSTEVARCIAHLHRTELHGQQISVEKVKGDPSKKELKKEIDEKSGSGRNMGDKKTASIDKTSKTQSAKKEEKKSDKSEKKESKEAKKTEGKDEKNDNGSSGPNQESTKKTEEKKRISGKSPGQMVVLDQTKGDQGHTRTVRRGRFDKVSICKNKERFIQDKMKFREYRGRKDILPFEKMKEQRMREHMVRLERIRRAVELRRRRELAERERRERERIRIIREREERERLQRERERLEIERQKLERERMERERLERERIRIEQERRKEAERIAREREELRRQQQQLRYEQEKRNSLKRPRDVDHRRDDPYWNENKKMALDTDARFSHGSDYSRQQNRFNDFDHRERGRYPEGSSVPSSSFERRERFVNQSEGKKTRPTARREEPGFERYPKNFSESRRNEPPQPRNELRDTDRREVRGDRDERRTVIIHDRPEISHGRHPRETGSNPPRQTNWKNEGGISTDKRDARGERPERSGREVSGHVRSAAPGSRSSTSGYGSREGERGVIGERGSGGQHYNEDRHVVERHSRETGPRKEWHGPSSQGSGYHDARRMGDGRGGGGMMSPHTSNSSPINRVVQITGSSMQRGSGSGFKPFKSGPPRRF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQF5</accession>
    <name>A0A452IQF5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">SWI/SNF-related matrix-associated actin-dependent regulator of chromatin subfamily E member 1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="11">BRG1-associated factor 57</fullName>
        </alternativeName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IQF5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034437.1">
        <property type="protein sequence ID" value="ENSGAGP00000030343.1"/>
        <property type="gene ID" value="ENSGAGG00000021867.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016514">
        <property type="term" value="C:SWI/SNF complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016922">
        <property type="term" value="F:nuclear receptor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031492">
        <property type="term" value="F:nucleosomal DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006325">
        <property type="term" value="P:chromatin organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045892">
        <property type="term" value="P:negative regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd21983">
        <property type="entry name" value="HMG-box_SMARCE1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.30.10:FF:000011">
        <property type="entry name" value="Putative SWI/SNF-related matrix-associated actin-dependent regulator of chromatin subfamily E member 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.30.10">
        <property type="entry name" value="High mobility group box domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009071">
        <property type="entry name" value="HMG_box_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036910">
        <property type="entry name" value="HMG_box_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46232">
        <property type="entry name" value="SMARCE1 REGULATOR OF CHROMATIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46232:SF1">
        <property type="entry name" value="SWI_SNF-RELATED MATRIX-ASSOCIATED ACTIN-DEPENDENT REGULATOR OF CHROMATIN SUBFAMILY E MEMBER 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00505">
        <property type="entry name" value="HMG_box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00398">
        <property type="entry name" value="HMG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47095">
        <property type="entry name" value="HMG-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50118">
        <property type="entry name" value="HMG_BOX_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0156">Chromatin regulator</keyword>
    <keyword evidence="7 13" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="8 12" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="2" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="1" id="KW-0488">Methylation</keyword>
    <keyword evidence="6" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="9 12" id="KW-0539">Nucleus</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="HMG box" evidence="15">
        <location>
            <begin position="31"/>
            <end position="99"/>
        </location>
    </feature>
    <feature type="DNA-binding region" description="HMG box" evidence="12">
        <location>
            <begin position="31"/>
            <end position="99"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1"/>
            <end position="36"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="118"/>
            <end position="140"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="259"/>
            <end position="377"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="13">
        <location>
            <begin position="70"/>
            <end position="101"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="118"/>
            <end position="133"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="259"/>
            <end position="273"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="348"/>
            <end position="360"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00067740"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00079499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00267"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50118"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030343.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="377" mass="42745" checksum="56CC5456E3068205" modified="2019-05-08" version="1">MSKRPSYAPPPTPAPTTASSGITIPKPPKPPDKPLMPYMRYSRKVWDQVKASNPDLKLWEIGKIIGGMWRDLTDEEKQEYLNEYEAEKIEYNESMKAYHNSPAYLAYINAKSRAEAALEEESRQRQSRMEKGEPYMSIQPAEDPDDYDDGFSMKHTATARFQRNHRLISEILSESVVPDVRSVVTTARMQVLKRQVQSLMVHQRKLEAELLQIEERHQEKKRKFLESTESFNNELKRLCSLKVEVDMEKIAAEIAQAEEQARKRQEEREKEAAEQAECSQTNAASEEEQAVNKTEEKKEEESAPMETEEAHPEETTENQQNGGEGTSTPEDKESGQEGVDSTAEEGTSDSNTGSESNSATVEELPADPAPDDSEKKE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQF8</accession>
    <name>A0A452IQF8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Lethal giant larvae homologue 2 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the WD repeat L(2)GL family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQF8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034305.1">
        <property type="protein sequence ID" value="ENSGAGP00000030224.1"/>
        <property type="gene ID" value="ENSGAGG00000021760.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031201">
        <property type="term" value="C:SNARE complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045159">
        <property type="term" value="F:myosin II binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019905">
        <property type="term" value="F:syntaxin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006887">
        <property type="term" value="P:exocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006893">
        <property type="term" value="P:Golgi to plasma membrane transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:000521">
        <property type="entry name" value="syntaxin-binding protein 5-like isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000664">
        <property type="entry name" value="Lethal2_giant"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013577">
        <property type="entry name" value="LLGL2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036322">
        <property type="entry name" value="WD40_repeat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001680">
        <property type="entry name" value="WD40_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10241">
        <property type="entry name" value="LETHAL 2 GIANT LARVAE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10241:SF22">
        <property type="entry name" value="SYNTAXIN-BINDING PROTEIN 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08366">
        <property type="entry name" value="LLGL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00400">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00962">
        <property type="entry name" value="LETHAL2GIANT"/>
    </dbReference>
    <dbReference type="SMART" id="SM00320">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50978">
        <property type="entry name" value="WD40 repeat-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50082">
        <property type="entry name" value="WD_REPEATS_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50294">
        <property type="entry name" value="WD_REPEATS_REGION"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0268">Exocytosis</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="3 5" id="KW-0853">WD repeat</keyword>
    <feature type="repeat" description="WD" evidence="5">
        <location>
            <begin position="234"/>
            <end position="269"/>
        </location>
    </feature>
    <feature type="domain" description="Lethal giant larvae homologue 2" evidence="7">
        <location>
            <begin position="278"/>
            <end position="385"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="17"/>
            <end position="38"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="556"/>
            <end position="596"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="675"/>
            <end position="694"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="17"/>
            <end position="31"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="570"/>
            <end position="580"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="582"/>
            <end position="591"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00008070"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022574"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF08366"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030224.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="733" mass="81864" checksum="6A54BB54E062305A" modified="2019-05-08" version="1">MRKFNIRKVLDGLTAVSSSSSASSSSQQQQSGNRESDVPETLQSEHFQLCKTVRHGFPYQPSALAFDPVQRILAIGTQTGALRLFGRPGVECYCQHDAETAVIQLQFLINEGALVSALADDTLHLWNLRQKRPAIFHSLKFNRERITFCHLPFQSKWLYVGTERGNIHIVNVESFTLSGYVIMWNKAIELSSKSHPGPVVHISDNPMDEGKLLIGYESGIVVLWDLKSKKADYRYTHDEAIHSVAWHHEGKQFICSHSDGTLTIWNIKSPAKPLQTITPHGKQLKDGKKPEPCKPILKVEYKTARAGEPFIILSGGLSYDTVGRRPCLTVMHGKSTAVLEMDYSIVDFLTLCETPYPNDFQEPYAVVVLLEKDLVLIDLAQNGYPIFENPYPLNIHESPVTCCEYFADCPVDLIPALYSVGARQKRQGYSKKEWPISGGNWGLVTQSYPEIIITGHADGSIKFWDASAITLQVLYKLKTAKVFEKSRNKDDKPNTDIVDEDPYAIQIISWCPESRMLCIAGVSAHVIIYRFSKQEVTTEVIPMLEVRLLYEINDVESPDGEQAPPLPTPGTGSNPQSIAPQSHPSTSSSSSDGLRDNVPCLKIKNSPLKQSAGYQIELVIQLVWVSGEPPQQITSLAVNSAYGLVVFGNCNGIAMVDYLQKTVLLNLGTTELYGSNDPYRREPRSPRKSRQPSGGKNCWMLHFNNVRIGQTVPIAYLPSLRSLWDSKFFLMSH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQG1</accession>
    <name>A0A452IQG1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">E3 ubiquitin-protein ligase Arkadia</fullName>
            <ecNumber evidence="6">2.3.2.27</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="20">RING finger protein 111</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="19">RING-type E3 ubiquitin transferase Arkadia</fullName>
        </alternativeName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1 26">
            <text>S-ubiquitinyl-[E2 ubiquitin-conjugating enzyme]-L-cysteine + [acceptor protein]-L-lysine = [E2 ubiquitin-conjugating enzyme]-L-cysteine + N(6)-ubiquitinyl-[acceptor protein]-L-lysine.</text>
            <dbReference type="EC" id="2.3.2.27"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="4">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
            <location evidence="2">PML body</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the Arkadia family.</text>
    </comment>
    <dbReference type="EC" id="2.3.2.27" evidence="6"/>
    <dbReference type="AlphaFoldDB" id="A0A452IQG1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034284.1">
        <property type="protein sequence ID" value="ENSGAGP00000030203.1"/>
        <property type="gene ID" value="ENSGAGG00000021784.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016605">
        <property type="term" value="C:PML body"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061630">
        <property type="term" value="F:ubiquitin protein ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006281">
        <property type="term" value="P:DNA repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016567">
        <property type="term" value="P:protein ubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd16681">
        <property type="entry name" value="RING-H2_RNF111"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.40.10:FF:000165">
        <property type="entry name" value="E3 ubiquitin-protein ligase Arkadia isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029306">
        <property type="entry name" value="RNF111_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001841">
        <property type="entry name" value="Znf_RING"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051073">
        <property type="entry name" value="ZNRF3_Arkadia_E3_ligases"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16200">
        <property type="entry name" value="RING ZINC FINGER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15303">
        <property type="entry name" value="RNF111_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13639">
        <property type="entry name" value="zf-RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00184">
        <property type="entry name" value="RING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57850">
        <property type="entry name" value="RING/U-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50089">
        <property type="entry name" value="ZF_RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="12" id="KW-0227">DNA damage</keyword>
    <keyword evidence="16" id="KW-0234">DNA repair</keyword>
    <keyword evidence="9" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="11" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="17" id="KW-0539">Nucleus</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0808">Transferase</keyword>
    <keyword evidence="14" id="KW-0833">Ubl conjugation pathway</keyword>
    <keyword evidence="15" id="KW-0862">Zinc</keyword>
    <keyword evidence="13 21" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="RING-type" evidence="23">
        <location>
            <begin position="909"/>
            <end position="950"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="68"/>
            <end position="107"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="122"/>
            <end position="233"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="245"/>
            <end position="293"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="334"/>
            <end position="365"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="387"/>
            <end position="473"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="505"/>
            <end position="558"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="652"/>
            <end position="741"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="22">
        <location>
            <begin position="130"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="22">
        <location>
            <begin position="249"/>
            <end position="268"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="22">
        <location>
            <begin position="280"/>
            <end position="290"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="22">
        <location>
            <begin position="334"/>
            <end position="354"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="22">
        <location>
            <begin position="392"/>
            <end position="404"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="22">
        <location>
            <begin position="405"/>
            <end position="417"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="22">
        <location>
            <begin position="426"/>
            <end position="450"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="22">
        <location>
            <begin position="506"/>
            <end position="520"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="22">
        <location>
            <begin position="521"/>
            <end position="534"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="22">
        <location>
            <begin position="549"/>
            <end position="558"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="22">
        <location>
            <begin position="665"/>
            <end position="675"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="22">
        <location>
            <begin position="687"/>
            <end position="698"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="22">
        <location>
            <begin position="699"/>
            <end position="709"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="22">
        <location>
            <begin position="710"/>
            <end position="719"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004322"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00007622"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00012483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00074642"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00076609"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00080426"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS50089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030203.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="961" mass="105050" checksum="3E4772A468941A8A" modified="2019-05-08" version="1">MSQWTPECNIVYTIKADMKSEVPSDAPKRQESLKGILLNPEPIGAAKSFPAEVEMIANKVGNEFSHLCGDSQKQDMNGNHPEQDKGIVVRKKRKSQQAGPSYAQNCGVKENQGILGLRQHLETQSEDNDSSFSDCVSSPSSSLHFGDSDTITSDEEKETPVRHPQAVLSKNRTHSARSQKWPRTETEAVPGLLIKRPCFHSNSPRRPPYRKRFVKTSSSQRTQNQKERILIQRKKREVLARRKYALLPSSSSSSENDLSSESSSSSSTEGEEDLFVSPGDNLQNSTTVPSGSIDEDVVVIEASSTPQVAANEEINVTSTDSEVEIVTVGESYRSRSTIGHSRSHWNQNSSSHAARSQEQRNRSRISTVIQPLRQNAAEVVDLTVDEDEPTVVPTTSARVETQVVSSTSSNNTNTSTSEQASDAAISVSSSQISTAPDTLTTLPNSSTAGTSVGDDVRRTASNTTLETGPPAMPRLPSCCPQHSPCGGSSQNHHVLGHPHSSCFQQHGHHFQHHHHHHHNPHSTVPLSPSFSDSSCPVERPPPVSAPCGASSSTGTSYHDQQALPVDLSNNGIRSHGNGAFHGASAFDPCCPGSSSRTAIYGHQAGAGPSQPIAIDGYGTSMVAQPQPQPPAQASLSSCRHYMHSPYALTRPLHQSSACPHSHGNAPPPQPQPPPQVDYVIPHPVHPFHPSISSHASSHPVPPPPPPPTHPLASAAAPIPQHLPPTHQPITHHIPATAPSAQRLHPHEVIQRMEVQRRRMMQHPTRAHERPPPHPHRMHPNYGHGHHIHVPQTMSSHPRQAPERSAWELGIEAGVAAATYPPGPLHPHLAHYHAPPRLHHLQIGALPLMELIHLEERLGNVNRGASQGTIERCTYPHKYKKVTTDWFSQRKLHCKQDGEEGTEEDTEEKCTICLSILEEGEDVRRLPCMHLFHQVCVDQWLITNKKCPICRVDIEAQLPSES</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQG2</accession>
    <name>A0A452IQG2_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="14">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endomembrane system</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="9">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000034442.1">
        <property type="protein sequence ID" value="ENSGAGP00000030348.1"/>
        <property type="gene ID" value="ENSGAGG00000021780.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005770">
        <property type="term" value="C:late endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005802">
        <property type="term" value="C:trans-Golgi network"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005537">
        <property type="term" value="F:D-mannose binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005520">
        <property type="term" value="F:insulin-like growth factor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0038023">
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        <property type="entry name" value="CATION-INDEPENDENT MANNOSE-6-PHOSPHATE RECEPTOR"/>
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    <keyword evidence="6 11" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="2" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="2257"/>
            <end position="2279"/>
        </location>
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        <location>
            <begin position="37"/>
            <end position="154"/>
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        <location>
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        <location>
            <begin position="752"/>
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        <location>
            <begin position="919"/>
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        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
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        <source>
            <dbReference type="SAM" id="Phobius"/>
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    <sequence length="2445" mass="271913" checksum="05B36C241025D926" modified="2019-05-08" version="1">PSRQPAQLCVSLPSPAGRQLRGPPPPHQQYVLTYRLPFCFSFSYTWEAFDVDKHVRYKINPCGGIEDCDGSSAICAYDLNKKVYLSVGDSSLKRSSKSLLEFNTTKPCLQQGAQHVVQSNINFLCGKTLGTPELVTATECVHYFEWRTFIACKKGSFKAKKEVPCYVFDDLKKHDLNPLIKTSGGYLVDDSDEDSLYINVCRDIGSSSSETRSCPVGSAACLLSEGHAFDVGSPQEQLKLRDKDRLVLNYEKTYSEEEKPNFCQGHNPAVTITFVCPSKRREERTGPKLTAKLKCRYEIEWVTEYACHRDYLESRNCSLTSEQHDVSIDLTPLTQQPSYVTPYVAKDEKEEYYYYLNICGKTSAGNCKDDTESVSSCQVKTRGQQEKVAGKFKNQTLRYSDGDLTLIYPDGDECSSGFQRMTVINFECNETAGNEGKGAPVFTGEVDCTYFFTWDTKYACVQEKEDLLCRVADKKKRYDLSPLIRSSESAQNWEAVNSSPVEADKHFFINVCHKVLKKGAAVVCSDDAAVCSVDKNNKPKNLGTVMSALKVGERIQLTYSNGDSCGNNNMIKTVITLVCKPGDLESAPVFNLENDECLYVFEWYTAAACVLSKTEGDNCRVSDAQAGFSFDLSPLTKRSGSYVINTTDYDFYINICSSVPEEHCLLKSAAACQVAKNLGEPSSKLSYYDGMIQLTYKNGTTYNNEEKTERSTLITFLCDYEAGIGQPEYQVEDNYTYNFKWYTKYACPEMPLECVVTDPNTMEQYDLSSLSKSEDRGENWYAMDNSEPNERRKYYINVCRPLVPVAGCDRYASGCQMKFEKDNDSFSETTSISNLGIASKGLVIEGRGRVLLTYTNGSVCISADGKRTSYTTIIHFTCSKGTLNSSPRFLDNQECVVTFLWETEAACPVTTTKDETQVSSCSVRDPNSGFLFNMQPLASEKGYITSGIGKTFMLNICGPAPDCGDGSDGKPAGGCEVEGLKSVRIVGLDKTLSLSSEGFLTLTYRGNFEVKSGASDTFTVTFICNDSYPGELKFVDEEINSAAGIHNTFFEFHTALYDLSDLSKEGEPWIAVDTSKDAKTRTFYLNVCKPLPYVPGCPGGAIGSCVKSADKGQNLGIIQISPQAATDGSLSIIYLNGDTCKDKQRYSTRILFECDQTLGSPMFQHEDGCEFVFIWRTLAACPVHRAEGDNCQVKDPRYGFVYDLRPLSEKEKAVSTDEYDYYFRVCERITEACPQVNRLSNVSSCQVKKQDHSVKVAGLYTEKLTFEHGLIKINYTSGERCHKIYERSTAIFFYCDHNTLEPVFLKETPDCTYLFEWHTQYACPPFRSIECSYKDNKGNSFDLSPLTRHRENWEAIPMSGSTQKYYINVCQSLVLHAGAGSCPSDAAACLVDGSKYINLGEVADGPRWENGISVLKYVNGEPCPDKIRKRTTIIRLKCDESRVESKPELVTTIEECEYTFLWFTAAACPLKRNVQNDCRVTNPATGHLFDLNSLKRESGYRLYDSKKRRTLQLNVCNEAKSPCGSGVGACIIGGQKPINAGRLSKMLIYEDQLLKLVYSGGDPCPANAELKHTSFFSFVCKSDAGAGSQPVLLSFDEQTCTHYFSWHTTLACEEEVKCSVLNGSSVIDLSPLIHRTGYYEAFVDELTDSTPDFYINICEPLNPVADVNCPPGAAVCMDPVNGPPIDIGRITGPPKINPAVNEVYITFNSTTACEADKTFSSLIVFHCKRGTSLGNPKMIRKSDCSFVFEWATPLVCPDKVNMLGCSVTDELLNYTFNLSSLSGSSYKVTLTVYHIGVCAEVADVPQGKCIDGAVCLVSENNAASFGIIKGMKMDYRHQDETVILQYTGGDRCPPVTEKGELCVFPFKYKGKRYEDCIVDEEKRRWCATTNDYDKDGKRGFCGNATGNREATIIFKCDDNAGNGNPQLLSETLGCAVTFEWKTQFVCPPKKMECKFIRKHKTYDLRVLSSLTGSWLFNYHGIAYYMNLCQRINNGPTNCPERAAICSKSRNGAVRVLGVVHTQKLNVTDDKVLVSYSGGHECRKNKKATTIIELKCAKTIGEPRLHRIDEENCTYYITWGTRAACAVKPQEVEIVNGMVINPTTGKNFSLGDIYYKLYQASGDIRTNGDTYVYEIQLSAITDSAHPQCSGAKVCQIKTTSTYIRKVGSSDKAKYYVQDDDLDVVFSSDSACGKDKSKFVSSTILFHCNLQAKEGIPEFLHETADCQYLFTWYTSAVCPLGQSDEGAQIFEGLSGRSQAVGAVLSVLLVVLTACLLILLFYKKERRETVIHKITNCCRRSSNVSYKYTKINTEEEANENETEWLMEEIAASNQRPGKGGQENGHITTTSVKSEALTSLHVDDLDSEDEVLTVPDVKIHSTRGVQSRGSDQNRHQQKKPFSLGSDAKTNLLNGGKDRKTKPNPGQQKTQNSTNTLSFHDDSDEDLLNV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQG5</accession>
    <name>A0A452IQG5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Ig-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the MHC class II family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQG5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034156.1">
        <property type="protein sequence ID" value="ENSGAGP00000030084.1"/>
        <property type="gene ID" value="ENSGAGG00000021674.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042613">
        <property type="term" value="C:MHC class II protein complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002250">
        <property type="term" value="P:adaptive immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002504">
        <property type="term" value="P:antigen processing and presentation of peptide or polysaccharide antigen via MHC class II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd05767">
        <property type="entry name" value="IgC1_MHC_II_alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.320.10">
        <property type="entry name" value="Class II Histocompatibility Antigen, M Beta Chain, Chain B, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003006">
        <property type="entry name" value="Ig/MHC_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003597">
        <property type="entry name" value="Ig_C1-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050160">
        <property type="entry name" value="MHC/Immunoglobulin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011162">
        <property type="entry name" value="MHC_I/II-like_Ag-recog"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014745">
        <property type="entry name" value="MHC_II_a/b_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001003">
        <property type="entry name" value="MHC_II_a_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19944:SF86">
        <property type="entry name" value="HLA CLASS II HISTOCOMPATIBILITY ANTIGEN, DR ALPHA CHAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19944">
        <property type="entry name" value="MHC CLASS II-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07654">
        <property type="entry name" value="C1-set"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00993">
        <property type="entry name" value="MHC_II_alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00407">
        <property type="entry name" value="IGc1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00920">
        <property type="entry name" value="MHC_II_alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54452">
        <property type="entry name" value="MHC antigen-recognition domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00290">
        <property type="entry name" value="IG_MHC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-1064">Adaptive immunity</keyword>
    <keyword evidence="9" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="5" id="KW-0391">Immunity</keyword>
    <keyword evidence="8 12" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-0491">MHC II</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="3 12" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 12" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="212"/>
            <end position="229"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="13">
        <location>
            <begin position="71"/>
            <end position="190"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007394"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022859"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023130"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023182"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030084.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="244" mass="27419" checksum="579F4974468AA917" modified="2019-05-08" version="1">SALNKPPVLGAGVGADGDACFPTVDHVTSQVVFYQHMDGSQQGSEKFTQEFDQDEILYVDLERKKVVWRLPEFSKFTSFEVQHALGNMAVIKLYQEKLIERSSHTRAQNVPPELGEPNVLICFMDKFFPPVLSLTWLKNGQKVTEGVYETDFYPHQDNSFRKFSYLPFFPSHGDFYDCQVEHRGLPKPFTKHWEAQLPTSVAETTETLVCTLGLAVGIIGIIAGTILIIKGMKMNAAHNPRGPL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQG6</accession>
    <name>A0A452IQG6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Peptidase S1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQG6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034289.1">
        <property type="protein sequence ID" value="ENSGAGP00000030208.1"/>
        <property type="gene ID" value="ENSGAGG00000021723.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030141">
        <property type="term" value="C:secretory granule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051604">
        <property type="term" value="P:protein maturation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00190">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.10:FF:000047">
        <property type="entry name" value="Trypsin eta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.10.10">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009003">
        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043504">
        <property type="entry name" value="Peptidase_S1_PA_chymotrypsin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001314">
        <property type="entry name" value="Peptidase_S1A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001254">
        <property type="entry name" value="Trypsin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018114">
        <property type="entry name" value="TRYPSIN_HIS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033116">
        <property type="entry name" value="TRYPSIN_SER"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271:SF48">
        <property type="entry name" value="KALLIKREIN-14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271">
        <property type="entry name" value="KALLIKREIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00089">
        <property type="entry name" value="Trypsin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00722">
        <property type="entry name" value="CHYMOTRYPSIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00020">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50494">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50240">
        <property type="entry name" value="TRYPSIN_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00134">
        <property type="entry name" value="TRYPSIN_HIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00135">
        <property type="entry name" value="TRYPSIN_SER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="4 7" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="3 7" id="KW-0645">Protease</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="5 7" id="KW-0720">Serine protease</keyword>
    <feature type="domain" description="Peptidase S1" evidence="9">
        <location>
            <begin position="31"/>
            <end position="251"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU363034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030208.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="253" mass="27451" checksum="51FB9B4AF2293018" modified="2019-05-08" version="1">VNCSGQGAGSTPTTPAHFPSPPPALTPNNRIVGGAPCIPHSMPWQAALFMGSQLNCGGTLIDRNWVVTAAHCHVQCVRLGEHNIRHLDWTEQLRLSEKTILHPGYNSASKDNDIMLIKLLTPANLNKNVQPLELPTSCPSPKEECVVSGWGTTKSPQLYFPPVLHCGKVSIFSQEQCRSIYPRYYTENMLCAGVLEGGIDSCQGDSGGPLLCNGKLQGIVSWGTQTCALPNKPGVYVAVCKYVDWIRETINNN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQG7</accession>
    <name>A0A452IQG7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Conserved oligomeric Golgi complex subunit 2</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="8">Component of oligomeric Golgi complex 2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Golgi apparatus membrane</location>
            <topology evidence="1">Peripheral membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the COG2 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQG7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034447.1">
        <property type="protein sequence ID" value="ENSGAGP00000030353.1"/>
        <property type="gene ID" value="ENSGAGG00000021858.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000139">
        <property type="term" value="C:Golgi membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017119">
        <property type="term" value="C:Golgi transport complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007030">
        <property type="term" value="P:Golgi organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006891">
        <property type="term" value="P:intra-Golgi vesicle-mediated transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009316">
        <property type="entry name" value="COG2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024603">
        <property type="entry name" value="COG_complex_COG2_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024602">
        <property type="entry name" value="COG_su2_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12961">
        <property type="entry name" value="CONSERVED OLIGOMERIC GOLGI COMPLEX COMPONENT 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12961:SF0">
        <property type="entry name" value="CONSERVED OLIGOMERIC GOLGI COMPLEX SUBUNIT 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12022">
        <property type="entry name" value="COG2_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06148">
        <property type="entry name" value="COG2_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="7" id="KW-0472">Membrane</keyword>
    <keyword evidence="5" id="KW-0653">Protein transport</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0813">Transport</keyword>
    <feature type="domain" description="Conserved oligomeric Golgi complex subunit 2 N-terminal" evidence="10">
        <location>
            <begin position="16"/>
            <end position="80"/>
        </location>
    </feature>
    <feature type="domain" description="COG complex component COG2 C-terminal" evidence="11">
        <location>
            <begin position="370"/>
            <end position="592"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="464"/>
            <end position="497"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="469"/>
            <end position="486"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004395"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007603"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00020977"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00031344"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="Pfam" id="PF06148"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF12022"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030353.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="609" mass="69347" checksum="27F17D36A3286B31" modified="2019-05-08" version="1">MFFIYIPVSFCLLHFQPDFDVDHFVSDCRKRVQLEELRDDLELYYKLLKTAMVELINKDYADFVNLSTNLVGMDKALNQLSVPLGQLREEVMSLKSCVSEGIRTVDERMSKQEDIRRKKMCVLRLIQVIQSVEKIEKILHSQSSKELSLLEANSPLLTGQILERIATEFNQLQFHAVQSKGMPLLDKVRPRIAGITAMLQQSLEGLLLEGLQTSNVDIVRHCLRTYATIDKTRDAEALVGQVLVKPYVNEVIVEQYVQSHPNGLQAIYNKLLEFVPHHCRLLREVTGGAMSSEKANTVPGYDFLVNSVWPEIVRGLEEKLPSLFNPGNPDVFHEKYTISMDFIQRFERQCGSQASVKRLRLHPAYHSFNNKWNLPVYFQIRFREVAGALEAALSDSLEEAPAGSSYCLLATHLVWVSLLKCWSDQMFLPLLAHRLWKFTLQILARFSVFINELLLRPISNESTKDNKRSVATGSKESSVNPNPTEDQGNDAFPETQHLPSISNTQLVYVAADLDRLKEQLPELLEIIKPKLEMIGFKNISCITGALEDSQASLSASMPTLNNKIIQDLSESCFSYLKSALEVPRHYRRTNKVSADLMMLMFRDSFPSCK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQG8</accession>
    <name>A0A452IQG8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Protein Jade-1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="19">Jade family PHD finger protein 1</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="20">PHD finger protein 17</fullName>
        </alternativeName>
    </protein>
    <organism evidence="25 26">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="26" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="25" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="25" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Chromosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Cilium basal body</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="17">Belongs to the JADE family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQG8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034315.1">
        <property type="protein sequence ID" value="ENSGAGP00000030234.1"/>
        <property type="gene ID" value="ENSGAGG00000021796.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000123">
        <property type="term" value="C:histone acetyltransferase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006915">
        <property type="term" value="P:apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd15671">
        <property type="entry name" value="ePHD_JADE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd15679">
        <property type="entry name" value="PHD_JADE1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.40.10:FF:000004">
        <property type="entry name" value="Jade family PHD finger 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.40.10:FF:000030">
        <property type="entry name" value="Protein Jade-1 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019542">
        <property type="entry name" value="Enhancer_polycomb-like_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034732">
        <property type="entry name" value="EPHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050701">
        <property type="entry name" value="Histone_Mod_Regulator"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039546">
        <property type="entry name" value="Jade-1_PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019786">
        <property type="entry name" value="Zinc_finger_PHD-type_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011011">
        <property type="entry name" value="Znf_FYVE_PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001965">
        <property type="entry name" value="Znf_PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019787">
        <property type="entry name" value="Znf_PHD-finger"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13793">
        <property type="entry name" value="PHD FINGER PROTEINS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13793:SF79">
        <property type="entry name" value="PROTEIN JADE-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10513">
        <property type="entry name" value="EPL1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13831">
        <property type="entry name" value="PHD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13832">
        <property type="entry name" value="zf-HC5HC2H_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00249">
        <property type="entry name" value="PHD"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57903">
        <property type="entry name" value="FYVE/PHD zinc finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51805">
        <property type="entry name" value="EPHD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01359">
        <property type="entry name" value="ZF_PHD_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50016">
        <property type="entry name" value="ZF_PHD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12" id="KW-0010">Activator</keyword>
    <keyword evidence="6" id="KW-0053">Apoptosis</keyword>
    <keyword evidence="16" id="KW-0966">Cell projection</keyword>
    <keyword evidence="4" id="KW-0158">Chromosome</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="14" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="15" id="KW-0539">Nucleus</keyword>
    <keyword evidence="26" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="13" id="KW-0804">Transcription</keyword>
    <keyword evidence="11" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="10" id="KW-0862">Zinc</keyword>
    <keyword evidence="9 21" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="PHD-type" evidence="23">
        <location>
            <begin position="202"/>
            <end position="252"/>
        </location>
    </feature>
    <feature type="domain" description="PHD-type" evidence="24">
        <location>
            <begin position="254"/>
            <end position="368"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="1"/>
            <end position="42"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="367"/>
            <end position="400"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="588"/>
            <end position="623"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="669"/>
            <end position="714"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="785"/>
            <end position="827"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="22">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="22">
        <location>
            <begin position="26"/>
            <end position="35"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="22">
        <location>
            <begin position="379"/>
            <end position="394"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="22">
        <location>
            <begin position="597"/>
            <end position="620"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="22">
        <location>
            <begin position="681"/>
            <end position="691"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="22">
        <location>
            <begin position="785"/>
            <end position="797"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004120"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022454"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022703"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00038371"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00040212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00041998"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00042970"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00146"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS50016"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS51805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030234.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="842" mass="96229" checksum="E589D4075EA3A523" modified="2019-05-08" version="1">MKRSRLPSSSEDSDDNGSLSTWSQHSRSRHRRNSCSRHEDRKPSEVFRTDLITAMKLPDSYQLNPDEYYVLADPWRQEWEKGVQVPVSPGAIPEPVARVVSETKAVTFTRPRKYILSSGSEPPELGYVDIRTLADSVCRYDLNDVDATWLELANEELKEMGMPELDEYTMERVMEEFEQRCYDSMNHAIETEEGLGIEYDEDVVCDVCQSPDGEDGNEMVFCDKCNICVHQACYGILKVPEGSWLCRTCALGVQPKCLLCPKKGGAMKPTRSGTKWVHVSCALWIPEVSIGSPEKMEPITKVSHIPSSRWALVCSLCNEKIGASIQCSVKNCRTAFHVTCAFDRGLEMKTILAENDEVKFKSYCPKHSSTKKLDDENLCENTSQENENGTQDSSPCADIEPFHSLDQNQEEAHRVSVRKQKLQQLEDEFYTFVNVLEVTKALRLPEELVEFLYQYWKLKRKGNFNKPLITPKKDEEDNLAKREQDVLFRRLQLFTHLRQDLERVRNLTYMVTRREKIKRSVCKVQEQIFNLYTKLMDQERVPGVPSSSSSAENMVLFNSPPLGPDAPKIEDLKWHSAFFRKQMGTSLTHTLKKSHKGDRVRNSSGSDRKSLLKQPSRREGVVTPGSFLNFEKAFAEARIVSAQQRNGIVMPDHRKRRDNRSSYEVIKAELKEKPSKHSHKPLTSTEVSQRQSENKKSANHPGGRSVPGTRRDIVPKCNGSLIKINYNRPAVKVPTTPTSPVKNWGGFRIPKKGERQHQGEIQEETCHQNSNYPYLSMDRVLPKDRAKSKLKTDRENDGYAPDAEMSDSETEVAEKKCRQQRLSPNSTINRRTDIIRRSILAS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQJ1</accession>
    <name>A0A452IQJ1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Amine oxidase</fullName>
            <ecNumber evidence="16">1.4.3.-</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="14">
            <text>an L-alpha-amino acid + O2 + H2O = a 2-oxocarboxylate + H2O2 + NH4(+)</text>
            <dbReference type="Rhea" id="RHEA:13781"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15379"/>
            <dbReference type="ChEBI" id="CHEBI:16240"/>
            <dbReference type="ChEBI" id="CHEBI:28938"/>
            <dbReference type="ChEBI" id="CHEBI:35179"/>
            <dbReference type="ChEBI" id="CHEBI:59869"/>
            <dbReference type="EC" id="1.4.3.2"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 16">
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the flavin monoamine oxidase family. FIG1 subfamily.</text>
    </comment>
    <dbReference type="EC" id="1.4.3.-" evidence="16"/>
    <dbReference type="AlphaFoldDB" id="A0A452IQJ1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034184.1">
        <property type="protein sequence ID" value="ENSGAGP00000030109.1"/>
        <property type="gene ID" value="ENSGAGG00000021720.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001716">
        <property type="term" value="F:L-amino-acid oxidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090729">
        <property type="term" value="F:toxin activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009063">
        <property type="term" value="P:amino acid catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042742">
        <property type="term" value="P:defense response to bacterium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.405.10:FF:000004">
        <property type="entry name" value="Amine oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.660.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.50.50.60">
        <property type="entry name" value="FAD/NAD(P)-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.405.10">
        <property type="entry name" value="Guanine Nucleotide Dissociation Inhibitor, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002937">
        <property type="entry name" value="Amino_oxidase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036188">
        <property type="entry name" value="FAD/NAD-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001613">
        <property type="entry name" value="Flavin_amine_oxidase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050281">
        <property type="entry name" value="Flavin_monoamine_oxidase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10742:SF355">
        <property type="entry name" value="AMINE OXIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10742">
        <property type="entry name" value="FLAVIN MONOAMINE OXIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01593">
        <property type="entry name" value="Amino_oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00757">
        <property type="entry name" value="AMINEOXDASEF"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54373">
        <property type="entry name" value="FAD-linked reductases, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51905">
        <property type="entry name" value="FAD/NAD(P)-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0044">Antibiotic</keyword>
    <keyword evidence="5" id="KW-0929">Antimicrobial</keyword>
    <keyword evidence="11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="8 16" id="KW-0274">FAD</keyword>
    <keyword evidence="6 16" id="KW-0285">Flavoprotein</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13" id="KW-1199">Hemostasis impairing toxin</keyword>
    <keyword evidence="9 16" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="7" id="KW-0800">Toxin</keyword>
    <feature type="domain" description="Amine oxidase" evidence="17">
        <location>
            <begin position="51"/>
            <end position="482"/>
        </location>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="52"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <begin position="71"/>
            <end position="72"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <begin position="93"/>
            <end position="96"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="96"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="261"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="459"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001974"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005465"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022529"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022630"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022656"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022827"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023022"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00047637"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PIRSR" id="PIRSR601613-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="RuleBase" id="RU362067"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF01593"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030109.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="502" mass="57286" checksum="8208481C23008657" modified="2019-05-08" version="1">QRLLLESLPGKLKKWEQCFDDPDYGELVDIAKNGLERKCNSKKVVIVGAGISGLTAAKLLRDAGCEVLILEASGRLGGRIETYRERDWYMDLGPMRLPKHHRIVLEFIKQFKLKLNTFYNTDANTWYLVSNVRARYKDVEQNPNILRYPVHPSEKGKSATQLYEQTLDKVTTDCNALKEKYDSFSTKEYLIKEGHLSRGAIDMIGDLLNEDGRFFMSFLYSVMAHVTFSNSSFYEITGGFDQLPNAFYQYVHGSIQFRSAVVKILTKGDQVIVSYRKANTSDLLSVTADYVLVTTTAKAARLIKFSPPLSPLKTHALRSIHYTKDVKIALACTKKFWEKDGIHGGRSITDRPSRFIYYPSHNFPSGLGVILASYTWGDDAEFFIPLSDEKCVDVVLEDLAKIHQLPKDYIQRVCNKYVIKKWGLDRYSMGSYVSFTQYQYVDYSKALFQNEGRIHFSGELTAQPHAWIDTAMKSGLRAARNIHAVINASPRQQQHELKENEL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQJ6</accession>
    <name>A0A452IQJ6_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="9">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the RPAP1 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000034478.1">
        <property type="protein sequence ID" value="ENSGAGP00000030380.1"/>
        <property type="gene ID" value="ENSGAGG00000021786.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006366">
        <property type="term" value="P:transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039913">
        <property type="entry name" value="RPAP1/Rba50"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013929">
        <property type="entry name" value="RPAP1_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013930">
        <property type="entry name" value="RPAP1_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057989">
        <property type="entry name" value="TPR_RPAP1/MINIYO-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21483">
        <property type="entry name" value="RNA POLYMERASE II-ASSOCIATED PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21483:SF18">
        <property type="entry name" value="RNA POLYMERASE II-ASSOCIATED PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08620">
        <property type="entry name" value="RPAP1_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08621">
        <property type="entry name" value="RPAP1_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25766">
        <property type="entry name" value="TPR_RPAP1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0539">Nucleus</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0804">Transcription</keyword>
    <feature type="domain" description="RPAP1 N-terminal" evidence="7">
        <location>
            <begin position="190"/>
            <end position="232"/>
        </location>
    </feature>
    <feature type="domain" description="RPAP1 C-terminal" evidence="6">
        <location>
            <begin position="325"/>
            <end position="390"/>
        </location>
    </feature>
    <feature type="domain" description="RPAP1/MINIYO-like TPR repeats" evidence="8">
        <location>
            <begin position="1069"/>
            <end position="1294"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="31"/>
            <end position="56"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="228"/>
            <end position="255"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="465"/>
            <end position="493"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="47"/>
            <end position="56"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="229"/>
            <end position="243"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="5">
        <location>
            <begin position="467"/>
            <end position="480"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009953"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="Pfam" id="PF08620"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF08621"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF25766"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030380.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1366" mass="153574" checksum="E2784C60CF024AC8" modified="2019-05-08" version="1">MLSRPKPGESEADLLRFQNQFLAARASPAVKVVKKADKRRGAGGSGDEERPPLQDSRDVVLLDDFPDVLPALIPAPPKKSKFKNACVRFDDEDPEERLECHDRHITAVFSKIIERDTSMAAVTMPVPTGDPFPRAFHRSEVKNEVRNAASPESSLTFFFNYQNFPLFLRPHIITGEGLGNLESEREALAIHEENLQKLHSMTQEEILQEQRRLLVQLDPSLVAFLKSRRSVDEGQERQEKPGHPEPGTESQHDGEETVKDFAVGGSGMEESVRPERDTRMEITEVDLPIKPQKEWIHMNNVEFEKLEWIKDLPRPRQRRTKKGMQARFSLKGELIPPDADLPTHLGLHHHGEEAERAGYSLQELFHLCRSQIIQQRTLALQVLGHVVQKAKAGEFASSLKGSVLRLLLDAGFLFLLRFSLDDTVDNVIAASLRALHALLVSPEDEEYLDQTFSWYQGMAVFPFTPNSEEEEEEEEEEELTLMEKTQGKKSKVENKSDPDVARYDVIKGLLKTQFLQRLRYILEVVRPVPAIVLDILDILIRVARHSSEACSQLLACPRLIETIVREFLPVQWDPRVAEPGHLLTSLHGVACATAMKLIRVLASGGRNATARLLNKFELKSRLSRFIAEDPQDLPLRREEAIRLSTEAFRLWAVAASCGQSCDLYRDLYPVLVKILQSLPHLLSDCQENPTFPLSIQRATSVITLLTHVTQTAGCTAELSGPGDGEQIPPPPVTWNQVSGLRPFLETNLRRSVQEMSRLETWQALQPLTTTYIIYLGVYYSVCSQQPSLNPIDCLEEVERLTYEVLLPLFSQPAMHRMWDLLRPCSALCNPLSCSPAPESVPSIVSLSCTGGKPQMSLVGSKSPFPFLTALLFLINNITHIHKGLVSKYTSVLDLKGLKDYLLQSWPTGPLSITPSSAWILRHEYHLQYFVLALAGICPDYTQHASLHHCVAMALLSRLLPGSEHLAHELLLGLAFNPEFIPEGKTGGPEAADFSDILHLGTSFPKPHRGVLLGEAYQHLPFIRACYLSQFVHLEPALTRSQVAYQGRTYLVQSLLLPEVKGPILPSDWPFLPLISLYNKVTGAETGGAVLNTLPPDLVNTVTRNLQWVLLLETWRARVLQNIPAAAKLARLMCVFLTGGDLFLEGPVHCYTASLLALYCQPKVLDSLELDAPLPGLASFHDLYISVLEQFEGVSFGDPLFGAFVLLPLQRRFNVQLRLAVFGEHVSTLRALGVPLKQFPVPLERYTLPPEDNLNLLRQYFRTLVTGALRHAWCPVLYAVAVAHVNSFIFSQNSTTEADAARKSMLRKTWRLVDEVLKKHLLYYKVHNPESPMGVDLYEQLPPIRLKYLQMVTKKEPKENMQMSDPE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQJ9</accession>
    <name>A0A452IQJ9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="19">F-box DNA helicase 1</fullName>
            <ecNumber evidence="17">5.6.2.4</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="21">DNA 3'-5' helicase 1</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="20">F-box only protein 18</fullName>
        </alternativeName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>ATP + H2O = ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:13065"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="5.6.2.4"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="16">
            <text>Couples ATP hydrolysis with the unwinding of duplex DNA by translocating in the 3'-5' direction.</text>
            <dbReference type="EC" id="5.6.2.4"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="3">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Chromosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the helicase family. UvrD subfamily.</text>
    </comment>
    <dbReference type="EC" id="5.6.2.4" evidence="17"/>
    <dbReference type="AlphaFoldDB" id="A0A452IQJ9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034261.1">
        <property type="protein sequence ID" value="ENSGAGP00000030181.1"/>
        <property type="gene ID" value="ENSGAGG00000021773.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005694">
        <property type="term" value="C:chromosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043138">
        <property type="term" value="F:3'-5' DNA helicase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016787">
        <property type="term" value="F:hydrolase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000724">
        <property type="term" value="P:double-strand break repair via homologous recombination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031297">
        <property type="term" value="P:replication fork processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd22095">
        <property type="entry name" value="F-box_FBXO18"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="F-box DNA helicase 1"/>
        <property type="match status" value="1"/>
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    <dbReference type="Gene3D" id="1.20.1280.50">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000212">
        <property type="entry name" value="DNA_helicase_UvrD/REP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036047">
        <property type="entry name" value="F-box-like_dom_sf"/>
    </dbReference>
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        <property type="entry name" value="F-box_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014016">
        <property type="entry name" value="UvrD-like_ATP-bd"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11070:SF30">
        <property type="entry name" value="F-BOX DNA HELICASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11070">
        <property type="entry name" value="UVRD / RECB / PCRA DNA HELICASE FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12937">
        <property type="entry name" value="F-box-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00580">
        <property type="entry name" value="UvrD-helicase"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="FBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81383">
        <property type="entry name" value="F-box domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50181">
        <property type="entry name" value="FBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="11" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="5" id="KW-0158">Chromosome</keyword>
    <keyword evidence="7" id="KW-0227">DNA damage</keyword>
    <keyword evidence="13" id="KW-0234">DNA repair</keyword>
    <keyword evidence="12" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="10" id="KW-0347">Helicase</keyword>
    <keyword evidence="9" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="14" id="KW-0413">Isomerase</keyword>
    <keyword evidence="6" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="15" id="KW-0539">Nucleus</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="F-box" evidence="23">
        <location>
            <begin position="197"/>
            <end position="246"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="22"/>
            <end position="48"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="22">
        <location>
            <begin position="22"/>
            <end position="39"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00009922"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022454"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022806"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023204"/>
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    </evidence>
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        <source>
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        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00034617"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00034808"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00048988"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00071173"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00075040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00079567"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS50181"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030181.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="999" mass="112615" checksum="A8942253E2D4868E" modified="2019-05-08" version="1">MLVPVRAFKRKHLTASDCEAFSQSREGSSSLTQPFSQRRASGDLNRGLYPTYRTKRQQEGQGHQRSITDYFKVIQTPPTGNQVAKSSEVKEESLDSFFMDEGLEDHCALLGCDGDAKTAPWGPVASRKRPLCSEGAGSNYTEYSKCGGTDKKPMIDSSLKYSPGHTEIRQDPDVIEIDPIPDSYFGLLGTSCGKEPQGHIDQLPNEVLSIIFAFLPVTDLCLNLNLVCHRWREIISDPLFILWKKLYRQYLMKEAQALLTVEWILDHYGITQEQDQCMLGFIRYMATSNAARRMDPDAVLRCLKKHHLFTKAEICIIQRLPDLEGTAGAVNVWAVIAAIVLFSDGVHDIQKLMACLRKPGSTLSLLDITEMLYCMATLLNAMREKGINISNRFHYSIFYSLYLMENACPVTKTTMLETPSSNCGRGAWPSNKLDIKLTHEQQRILNHTILPGQVVKIMAFAGTGKTSTLVKYAEKFPHLSFLYVAFNKAIVENGKSVFPPNVMCKTFHSLAYTSVGKHYNEKGKLNFSNLSPYTISFILKHRMGQSLFIRAKTVTQTLESFFASSDEFISVEHTPIWCKNTHGMRVLVSEEEKQIIVEEARRIWDNMKMLNGNTELNCRMTSDGYLKLWQLRKPRLFAYDAIFVDEAQDCTPAIMDIVLSQSCGIIFVGDPHQQIYTFRGAVNALFAVPHTHIFYLTKSFRFGPEIAYVGSTILDVCKKIRNKMLVGGNQEGDVRGSMNGKTAFLSRGNLNVFENAVTITGGERPTKIHLLGGPKNFGLDKIHDIWKLLQPEEARVRLKLQIDDHFIKKWVEKDGFQGLKNYAAKTEDRELEVKITIVEKYNVRIPELVARIKNSHVSNIAFADMSAEDEWNLLYVAVTRAKKHLLMTKSLEYLLTLAGEYFLRPELTSEALKEGAIHCSVSQCHNAVPADSILTMKKVPLTYSNGSKDQGGYLCHACVQQRLGPLTQLTISPELVQTMDVVDEVLEIPRHLAVLYGLF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2A1</accession>
    <name>A0A452J2A1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Ig-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J2A1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039095.1">
        <property type="protein sequence ID" value="ENSGAGP00000034526.1"/>
        <property type="gene ID" value="ENSGAGG00000024508.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002764">
        <property type="term" value="P:immune response-regulating signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000033">
        <property type="entry name" value="Killer cell immunoglobulin-like receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000049">
        <property type="entry name" value="Leukocyte immunoglobulin-like receptor subfamily B member 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050412">
        <property type="entry name" value="Ig-like_Receptors_ImmuneReg"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11738">
        <property type="entry name" value="MHC CLASS I NK CELL RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11738:SF186">
        <property type="entry name" value="OSTEOCLAST-ASSOCIATED IMMUNOGLOBULIN-LIKE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13895">
        <property type="entry name" value="Ig_2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="4" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="5" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <keyword evidence="1" id="KW-0732">Signal</keyword>
    <feature type="domain" description="Ig-like" evidence="7">
        <location>
            <begin position="34"/>
            <end position="102"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="371"/>
            <end position="393"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034526.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="393" mass="42954" checksum="791BB340E4FA3216" modified="2019-05-08" version="1">MTDGLNWGERRRGAVSPLFNSCLLLKAGHEFLKPTIWVSPSRVVALGRNVTIRCEGQDLGMEFVLCKAGHPNQQVQSVVPEGKMAEFPIASVGQEDGGSYTCYYRSITDQSHWSFPSNPVEIIVGGEGGAWLSGPAPRVSVGEAMAIWCRRQRQGVRFVLNKEGRHFPPVDSDGFGAVFPISNVSREDGGNYSCSYHSRSEPFAVSDPSDPMELVVRGEGYPKPNISGIPRRGVSVGEAMAIWCRRQRQGVRFVLNKEGRHFPPVDSDGFGAVFPISNVSREDGGNYSCSYHSRSEPFAVSDPSDPMELVCFCGGPRSGCWGFLQPARPASDPPLVVKGGGRSLSWCFSPSLLPDRRWFYSCRGNRPDPAWNRAGSHPPGQHGARYRAGGGIY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2A4</accession>
    <name>A0A452J2A4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">PID domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="3">Plays a role in the process of neurogenesis.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2A4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039110.1">
        <property type="protein sequence ID" value="ENSGAGP00000034540.1"/>
        <property type="gene ID" value="ENSGAGG00000024554.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016323">
        <property type="term" value="C:basolateral plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031410">
        <property type="term" value="C:cytoplasmic vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050769">
        <property type="term" value="P:positive regulation of neurogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="CDD" id="cd01268">
        <property type="entry name" value="PTB_Numb"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000031">
        <property type="entry name" value="protein numb isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016698">
        <property type="entry name" value="Numb/numb-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010449">
        <property type="entry name" value="Numb_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006020">
        <property type="entry name" value="PTB/PI_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47368">
        <property type="entry name" value="NUMB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47368:SF5">
        <property type="entry name" value="PROTEIN NUMB HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06311">
        <property type="entry name" value="NumbF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00640">
        <property type="entry name" value="PID"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF017607">
        <property type="entry name" value="Numb/numb-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00462">
        <property type="entry name" value="PTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01179">
        <property type="entry name" value="PID"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1 3" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="3" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="PID" evidence="5">
        <location>
            <begin position="37"/>
            <end position="173"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="217"/>
            <end position="248"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="403"/>
            <end position="428"/>
        </location>
    </feature>
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        <location>
            <begin position="493"/>
            <end position="512"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="518"/>
            <end position="601"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="223"/>
            <end position="247"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="4">
        <location>
            <begin position="410"/>
            <end position="421"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="580"/>
            <end position="594"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="PIRNR" id="PIRNR017607"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS01179"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034540.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="601" mass="65925" checksum="CC5609C4A37B5238" modified="2019-05-08" version="1">MNKLRQSFRRKKDVYVPEASRPHQWQTDEEGVRTGKCSFPVKYLGHVEVDESRGMHVCEDAVKRLKAERKFFKGFFGKTGRKAVKAVLWVSADGLRVVDEKTKDLIVDQTIEKVSFCAPDRNFDRAFSYICRDGTTRRWICHCFMAVKDTGERLSHAVGCAFAACLERKQKREKECGVTATFDASRTTFTREGSFRVTTATEQAEREEIMRQIQDAKAETEVKTTAAPSSTPSNTAPSPGSPASPTADLTASLDKEMNHPHAIPRRHAPIEQLARQGSFRGFPALSQKMSPFKRQLSLRINELPSTVQRKTDFPMKNSVPEMEGEVDSISSLCSQITNAFSTPPEDPFSSAPMTKPVTVVAPQSPAFQGTEWSTSSGAASPSLFQMTHRRTPSEADRWLEEVSKTVRAQQPPPAAPQPLAQPPLAASKPATPFPVNTFVAPQPVPVGVVPPMQPAFMPAQPYSVANGMTFSAPSVPVVGITPSQMVANVFGTASHTPASHPHQSPSLVKQPTFPQYETSNATASPFFKSPAQHNGSAAFNGVDSGKWATEDKQLQPPASAHQVDPFEAQWAALESKSKQRTNPSPTNPFSSDLQKTFEIEL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2A8</accession>
    <name>A0A452J2A8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Ig-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
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            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J2A8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039102.1">
        <property type="protein sequence ID" value="ENSGAGP00000034533.1"/>
        <property type="gene ID" value="ENSGAGG00000024508.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="2"/>
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    </dbReference>
    <dbReference type="InterPro" id="IPR050412">
        <property type="entry name" value="Ig-like_Receptors_ImmuneReg"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11738">
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    </dbReference>
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    </dbReference>
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    </dbReference>
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        <property type="entry name" value="Immunoglobulin"/>
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    </dbReference>
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    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034533.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="4">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="253" mass="27533" checksum="EDE11613E15BF8B0" modified="2025-10-08" version="2">MGLLPVSCCSPGLRGNSGPCEKASRSPGVSVGEAMAIWCRRQRQGVRFVLNKEGRHFPPVDSDGFGAVFPISNVSREDGGNYSCSYHSRSEPFAVSDPSDPMELVVRGEGYPKPNISGIPRRGVSVGEAMAIWCRRQRQGVRFVLNKEGRHFPPVDSDGFGAVFPISNVSREDGGNYSCSYHSRSEPFAVSDPSDPMELVNSWSHPTHHPPRKAVTPPPCSSQLFLCSSLCLSLSSSSPAWLLHEAGWGTLWE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2A9</accession>
    <name>A0A452J2A9_9SAUR</name>
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        <recommendedName>
            <fullName evidence="4">Ig-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J2A9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039107.1">
        <property type="protein sequence ID" value="ENSGAGP00000034537.1"/>
        <property type="gene ID" value="ENSGAGG00000024508.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="2"/>
    </dbReference>
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        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
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        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050412">
        <property type="entry name" value="Ig-like_Receptors_ImmuneReg"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11738">
        <property type="entry name" value="MHC CLASS I NK CELL RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11738:SF186">
        <property type="entry name" value="OSTEOCLAST-ASSOCIATED IMMUNOGLOBULIN-LIKE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13895">
        <property type="entry name" value="Ig_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034537.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="4">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="249" mass="27094" checksum="A6B44A7643BA29BF" modified="2025-10-08" version="2">MGLLPVSCCSPGLRGNSGPCEKASRSPGVSVGEAMAIWCRRQRQGVRFVLNKEGRHFPPVDSDGFGAVFPISNVSREDGGNYSCSYHSRSEPFAVSDPSDPMELVVRGEGYPKPNISGIPRRGVSVGEAMAIWCRRQRQGVRFVLNKEGRHFPPVDSDGFGAVFPISNVSREDGGNYSCSYHSRSEPFAVSDPSDPMELLTGLWIRFVHAMAGFSNSGSGPLREWQGFSMVGSRAVSLHSKTLLCIQHL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2B1</accession>
    <name>A0A452J2B1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">PID domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="3">Plays a role in the process of neurogenesis.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2B1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039109.1">
        <property type="protein sequence ID" value="ENSGAGP00000034539.1"/>
        <property type="gene ID" value="ENSGAGG00000024554.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016323">
        <property type="term" value="C:basolateral plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031410">
        <property type="term" value="C:cytoplasmic vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050769">
        <property type="term" value="P:positive regulation of neurogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="CDD" id="cd01268">
        <property type="entry name" value="PTB_Numb"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000031">
        <property type="entry name" value="protein numb isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016698">
        <property type="entry name" value="Numb/numb-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010449">
        <property type="entry name" value="Numb_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006020">
        <property type="entry name" value="PTB/PI_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47368">
        <property type="entry name" value="NUMB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47368:SF5">
        <property type="entry name" value="PROTEIN NUMB HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06311">
        <property type="entry name" value="NumbF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00640">
        <property type="entry name" value="PID"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF017607">
        <property type="entry name" value="Numb/numb-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00462">
        <property type="entry name" value="PTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01179">
        <property type="entry name" value="PID"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1 3" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="3" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="PID" evidence="5">
        <location>
            <begin position="37"/>
            <end position="162"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="206"/>
            <end position="237"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="405"/>
            <end position="465"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="530"/>
            <end position="549"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="555"/>
            <end position="638"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="212"/>
            <end position="236"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="405"/>
            <end position="422"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="428"/>
            <end position="441"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="4">
        <location>
            <begin position="447"/>
            <end position="458"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="617"/>
            <end position="631"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PIRNR" id="PIRNR017607"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS01179"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034539.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="638" mass="69259" checksum="76DF7D2F1373A7FE" modified="2019-05-08" version="1">MNKLRQSFRRKKDVYVPEASRPHQWQTDEEGVRTGKCSFPVKYLGHVEVDESRGMHVCEDAVKRLKATGRKAVKAVLWVSADGLRVVDEKTKDLIVDQTIEKVSFCAPDRNFDRAFSYICRDGTTRRWICHCFMAVKDTGERLSHAVGCAFAACLERKQKREKECGVTATFDASRTTFTREGSFRVTTATEQAEREEIMRQIQDAKAETEVKTTAAPSSTPSNTAPSPGSPASPTADLTASLDKEMNHPHAIPRRHAPIEQLARQGSFRGFPALSQKMSPFKRQLSLRINELPSTVQRKTDFPMKNSVPEMEGEVDSISSLCSQITNAFSTPPEDPFSSAPMTKPVTVVAPQSPAFQVNGTASAFCVLAVKPSQAAVAPTAMPVRETNPWAHAPAANTGAAATFSGTEWSTSSGAASPSLFQMTHRRTPSEADRWLEEVSKTVRAQQPPPAAPQPLAQPPLAASKPATPFPVNTFVAPQPVPVGVVPPMQPAFMPAQPYSVANGMTFSAPSVPVVGITPSQMVANVFGTASHTPASHPHQSPSLVKQPTFPQYETSNATASPFFKSPAQHNGSAAFNGVDSGKWATEDKQLQPPASAHQVDPFEAQWAALESKSKQRTNPSPTNPFSSDLQKTFEIEL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2B3</accession>
    <name>A0A452J2B3_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J2B3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039115.1">
        <property type="protein sequence ID" value="ENSGAGP00000034545.1"/>
        <property type="gene ID" value="ENSGAGG00000024550.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027886">
        <property type="entry name" value="SPMIP4"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31393">
        <property type="entry name" value="C5ORF31"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31393:SF2">
        <property type="entry name" value="CHROMOSOME 7 OPEN READING FRAME 31"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15093">
        <property type="entry name" value="SPMIP4-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="371"/>
            <end position="405"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034545.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="475" mass="54134" checksum="8093BEEC8B6DA8B8" modified="2019-05-08" version="1">MSVLNTDDQIPPKCNSSDLQLSLPFPVSHPYQTHISRYAMFPNFKSPEDQDTGIQASSHRPFHPNVPTKAYDVIVLRKTKGNLYRHEVINFPSDSKKEALHWPGQHAYFHFPKFVEGSNQIYYPKPPKTVAPNTTCNPLEPNLSPRTTNMLRNVERAQWITTYNHDFTGIGPMNPLILDDYDMKLIGRVTGELGEDVELKETFLPSLSQVRPLEGRIARLLQSRRPHESILQEQDSSDIHVPPALYTVHAVIPSCSGVILSNRKTANLDQQCKHTENEAKKRDEDSLQTLHLPQACYQRGESQDNRPIVNKFQKITDTLKTEALYRRQLAVRPESEPSPTPSWNSLYYEDLKPSHLDQYIVWHNPVSLSKPGVPKNQTGGEGGNFQPECSDQEETQYALSLDGNPPNMALPEWIPNSGVPRPQTSLLELQDSFSKTGAHKRFHNSITGEAKDLRDNIHEGRRHTFYGFNSFYFYN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2B4</accession>
    <name>A0A452J2B4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">WD repeat-containing protein 75 second beta-propeller domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
            <location evidence="1">Nucleolus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2B4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038994.1">
        <property type="protein sequence ID" value="ENSGAGP00000034439.1"/>
        <property type="gene ID" value="ENSGAGG00000024445.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032040">
        <property type="term" value="C:small-subunit processome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:2000234">
        <property type="term" value="P:positive regulation of rRNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045943">
        <property type="term" value="P:positive regulation of transcription by RNA polymerase I"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006364">
        <property type="term" value="P:rRNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057644">
        <property type="entry name" value="Beta-prop_WDR75_2nd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036322">
        <property type="entry name" value="WD40_repeat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001680">
        <property type="entry name" value="WD40_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR053826">
        <property type="entry name" value="WDR75"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR44215">
        <property type="entry name" value="WD REPEAT-CONTAINING PROTEIN 75"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR44215:SF1">
        <property type="entry name" value="WD REPEAT-CONTAINING PROTEIN 75"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23869">
        <property type="entry name" value="Beta-prop_WDR75_1st"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23769">
        <property type="entry name" value="Beta-prop_WDR75_2nd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00320">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50978">
        <property type="entry name" value="WD40 repeat-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0690">Ribosome biogenesis</keyword>
    <keyword evidence="3" id="KW-0698">rRNA processing</keyword>
    <keyword evidence="6" id="KW-0804">Transcription</keyword>
    <keyword evidence="4" id="KW-0853">WD repeat</keyword>
    <feature type="domain" description="WD repeat-containing protein 75 second beta-propeller" evidence="8">
        <location>
            <begin position="117"/>
            <end position="442"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004604"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022517"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022552"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022574"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF23769"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034439.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="610" mass="68832" checksum="8433D9EB57B18FBB" modified="2019-05-08" version="1">TFVVGYKLFAFYMLAGSEDSVFIIIPKNDEDFLDWFQLVSVKLPKTPDQEVEAKELSLILDDIGQWPKCTAFGREPLPCLANLYVFVHLEIIIIHSSLSVSAVIQGLVKGSDVKTGLVVDPRTKALVLNGKPGHLQFYCLQTDKQLYNLDIVQQEYIHQIGLNQVDLVKAAFSAQGNWLATVEQREEKEVELELQMKLWNYDEQTQSFTLNTRINMPHEDHVTALCFRDTDELEDGTPTLVTAGNDGRFKVWVLVNDSDSEKPSMGWSCDFVGSYHNYPATNCCFSEDGSLLAVSFEETVTVWNSDTWDLICTFCHPPGRIQNLCFGRLSCSKYLLGTTDCGFLCCWNLLSCALEWSAQLNVMVLQPDPLSENIAAVSRLLEGSDLFVFKPNEPRPFCIQRNVCRERVQCAVFIPRDVPDSIGSEKHQWLSRSQLYFLTETQDLLTFSTKSAEERLTPSSRQLAVEESLPMTPFSLLLGKHRHQQQSKENADLDTAVVQSHLAQGSPAVKELLHTPAHVLPSASFLCSIFINSVLISKENKSAGEVVDVVEMDSEKAEDESDEDLELTEMEQDINPTEFLGEITEIKLSKSQEKQLRKIRRMDYSWVAAL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2B6</accession>
    <name>A0A452J2B6_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="18">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 13">Membrane</location>
            <topology evidence="1 13">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 13">Belongs to the integrin alpha chain family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2B6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039135.1">
        <property type="protein sequence ID" value="ENSGAGP00000034563.1"/>
        <property type="gene ID" value="ENSGAGG00000024571.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009897">
        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008305">
        <property type="term" value="C:integrin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033627">
        <property type="term" value="P:cell adhesion mediated by integrin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098609">
        <property type="term" value="P:cell-cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007160">
        <property type="term" value="P:cell-matrix adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007229">
        <property type="term" value="P:integrin-mediated signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050900">
        <property type="term" value="P:leukocyte migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1530:FF:000001">
        <property type="entry name" value="Integrin subunit alpha 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.930">
        <property type="entry name" value="Bicelle-embedded integrin alpha(iib) transmembrane segment"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.130">
        <property type="entry name" value="Integrin alpha, N-terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1460">
        <property type="entry name" value="Integrin domains. Chain A, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1510">
        <property type="entry name" value="ntegrin, alpha v. Chain A, domain 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1530">
        <property type="entry name" value="ntegrin, alpha v. Chain A, domain 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013517">
        <property type="entry name" value="FG-GAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013519">
        <property type="entry name" value="Int_alpha_beta-p"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000413">
        <property type="entry name" value="Integrin_alpha"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018184">
        <property type="entry name" value="Integrin_alpha_C_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013649">
        <property type="entry name" value="Integrin_alpha_Ig-like_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048285">
        <property type="entry name" value="Integrin_alpha_Ig-like_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048286">
        <property type="entry name" value="Integrin_alpha_Ig-like_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028994">
        <property type="entry name" value="Integrin_alpha_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032695">
        <property type="entry name" value="Integrin_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23220">
        <property type="entry name" value="INTEGRIN ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23220:SF90">
        <property type="entry name" value="INTEGRIN ALPHA-7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01839">
        <property type="entry name" value="FG-GAP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08441">
        <property type="entry name" value="Integrin_A_Ig_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20805">
        <property type="entry name" value="Integrin_A_Ig_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20806">
        <property type="entry name" value="Integrin_A_Ig_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01185">
        <property type="entry name" value="INTEGRINA"/>
    </dbReference>
    <dbReference type="SMART" id="SM00191">
        <property type="entry name" value="Int_alpha"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69318">
        <property type="entry name" value="Integrin alpha N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69179">
        <property type="entry name" value="Integrin domains"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51470">
        <property type="entry name" value="FG_GAP"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00242">
        <property type="entry name" value="INTEGRIN_ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6 13" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="8 13" id="KW-0401">Integrin</keyword>
    <keyword evidence="9 13" id="KW-0472">Membrane</keyword>
    <keyword evidence="10 13" id="KW-0675">Receptor</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="4 13" id="KW-0732">Signal</keyword>
    <keyword evidence="3 13" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 13" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="13">
        <location>
            <begin position="1"/>
            <end position="21"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5018820331" evidence="13">
        <location>
            <begin position="22"/>
            <end position="1125"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="13">
        <location>
            <begin position="1066"/>
            <end position="1088"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="12">
        <location>
            <begin position="338"/>
            <end position="399"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="12">
        <location>
            <begin position="400"/>
            <end position="455"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="12">
        <location>
            <begin position="459"/>
            <end position="518"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha first immunoglubulin-like" evidence="15">
        <location>
            <begin position="503"/>
            <end position="667"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha second immunoglobulin-like" evidence="16">
        <location>
            <begin position="669"/>
            <end position="837"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha third immunoglobulin-like" evidence="17">
        <location>
            <begin position="843"/>
            <end position="1053"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="942"/>
            <end position="964"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="943"/>
            <end position="956"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023037"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00803"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="RuleBase" id="RU003762"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="Pfam" id="PF08441"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF20805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF20806"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034563.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1125" mass="122570" checksum="B86AC274EBDEF095" modified="2019-05-08" version="1">MRGASGAWLCCLGFQLLAGAAFNLDGTDPLLKDGAGGSLFGFSVALHRQLRPEPASWMLVGAPQALALPGQKANRTGGLYACPLTREIDDCWRVPIDEGVDLHKESKENQWLGVSVKSQGAGGKIVTCAHLYESRNRVNQPLETRDVIGRCYVLSQDLTVSEELDGGEWKFCEGRPQGHDRFGFCQQGVSAGFTADKHYILFGAPGTYNWKGDLRVELVNQSSLELVTYDDGPYEAGGEKDQDPSLIPVPANSYFGLLFMTNVHSADPDQLVYKTAEPSERLPGTVGDVAQNSNLGFSVDSGKGLTRQDQLSFVTGAPRANHTGAVVILRRDNVNRLVPEVVLPGEQLTSSFGYAVAVVDLNSDGWMDLVVGAPHFFDRQEEIGGAAYVYINQAGRWADIRPIRLNGTSHSMFGIGLTALGDLNQDGFQDIAVGAPFDGAGKVFIYHGSSLGIVTKPAQVLDGEGVGVRTFGYSLSGGLDVDGNLYPDLLVGSLSDSVVLYRARPVIHVSKNVSLSPQNIDLESRNCPHHLGICVEVRACFSYTASPASYSPRVTLTYTLDADVDRQKLGQAPRVAFLDRKPTDPEHQFSDVVELPHQRAPSCVKATFQLQESIRDKLRPIVVMLTYSIEHAQSKRQVQAAALPPLMPVLNAQLPNSQRAEVNFLKQGCGEDKICQSNLQLRYQFCSRVGDADFLPLPRDADGTAVFAMSDQKDVALEIQVTNLPSDPAAPQRDGDDAHEALLTATFPEALPYSGVRAHDMWAGPGLEKQPCLPNLNGSQVQCELGNPMKRGAQGRFYLIVSTSGITLETRELELELELSTISEQPGLRPVTARARVVIELPLSVTGVAIPQRLFFGGRVQGESAIRTEGQVGSAVRYEVTVSNRGQSLNTLGSAFLNLMWPHEIANGKWLLYPLRLELAARPGQRVACSPAANPLRLALESHGQDRSRREAERPEVPSTGPWWHRAHSERKNVTLDCAQGTARCLVFRCPLHSFDRSAVLTAWGRLWNSTFLEEYPSVTSVELLVHANITVKSTIQNLVLKDAATQIPVTIYLDPSAVVARGVPWWIILIAVLAGVLVLGLLVSILWKCGFFQRSSRAAPYATNYYRARLAVQPSEVEKQAGEA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2B7</accession>
    <name>A0A452J2B7_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J2B7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039114.1">
        <property type="protein sequence ID" value="ENSGAGP00000034544.1"/>
        <property type="gene ID" value="ENSGAGG00000024550.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027886">
        <property type="entry name" value="SPMIP4"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31393">
        <property type="entry name" value="C5ORF31"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31393:SF2">
        <property type="entry name" value="CHROMOSOME 7 OPEN READING FRAME 31"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15093">
        <property type="entry name" value="SPMIP4-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="478"/>
            <end position="512"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034544.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="582" mass="66775" checksum="081747D578F4F08C" modified="2019-05-08" version="1">MELLHSLPYYYREREGTDILSKTLESNEALTPLQIPSRPTVSQDRYEELRDSFQWCRLPWGTEREYGGIAPISLPEDHRPKNEPPCLFAKGHQHYGYGGVSWPRGLPIEQYYDVTQMKKSDVRMNDDLQLSLPFPVSHPYQTHISRYAMFPNFKSPEDQDTGIQASSHRPFHPNVPTKAYDVIVLRKTKGKIPHVLKSPHLDAARFLHFTYFILFLQFPKFVEGSNQIYYPKPPKTVAPNTTCNPLEPNLSPRTTNMLRNVERAQWITTYNHDFTGIGPMNPLILDDYDMKLIGRVTGELGEDVELKETFLPSLSQVRPLEGRIARLLQSRRPHESILQEQDSSDIHVPPALYTVHAVIPSCSGVILSNRKTANLDQQCKHTENEAKKRDEDSLQTLHLPQACYQRGESQDNRPIVNKFQKITDTLKTEALYRRQLAVRPESEPSPTPSWNSLYYEDLKPSHLDQYIVWHNPVSLSKPGVPKNQTGGEGGNFQPECSDQEETQYALSLDGNPPNMALPEWIPNSGVPRPQTSLLELQDSFSKTGAHKRFHNSITGEAKDLRDNIHEGRRHTFYGFNSFYFYN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2C3</accession>
    <name>A0A452J2C3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">HECT-type E3 ubiquitin transferase</fullName>
            <ecNumber evidence="4">2.3.2.26</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>S-ubiquitinyl-[E2 ubiquitin-conjugating enzyme]-L-cysteine + [acceptor protein]-L-lysine = [E2 ubiquitin-conjugating enzyme]-L-cysteine + N(6)-ubiquitinyl-[acceptor protein]-L-lysine.</text>
            <dbReference type="EC" id="2.3.2.26"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="3">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="2.3.2.26" evidence="4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039127.1">
        <property type="protein sequence ID" value="ENSGAGP00000034555.1"/>
        <property type="gene ID" value="ENSGAGG00000024483.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061630">
        <property type="term" value="F:ubiquitin protein ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="CDD" id="cd00078">
        <property type="entry name" value="HECTc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12881">
        <property type="entry name" value="SPRY_HERC1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="UBA_HERC1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.30:FF:000001">
        <property type="entry name" value="LOW QUALITY PROTEIN: probable E3 ubiquitin-protein ligase HERC1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.920:FF:000015">
        <property type="entry name" value="LOW QUALITY PROTEIN: probable E3 ubiquitin-protein ligase HERC1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.2410.10:FF:000006">
        <property type="entry name" value="probable E3 ubiquitin-protein ligase HERC1 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.920">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.2160.10">
        <property type="entry name" value="Hect, E3 ligase catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.2410.10">
        <property type="entry name" value="Hect, E3 ligase catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1750.10">
        <property type="entry name" value="Hect, E3 ligase catalytic domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.30">
        <property type="entry name" value="Regulator of chromosome condensation 1/beta-lactamase-inhibitor protein II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001870">
        <property type="entry name" value="B30.2/SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043136">
        <property type="entry name" value="B30.2/SPRY_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000569">
        <property type="entry name" value="HECT_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035983">
        <property type="entry name" value="Hect_E3_ubiquitin_ligase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058923">
        <property type="entry name" value="RCC1-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009091">
        <property type="entry name" value="RCC1/BLIP-II"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000408">
        <property type="entry name" value="Reg_chr_condens"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051625">
        <property type="entry name" value="Signaling_Regulatory_Domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003877">
        <property type="entry name" value="SPRY_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035768">
        <property type="entry name" value="SPRY_HERC1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019775">
        <property type="entry name" value="WD40_repeat_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036322">
        <property type="entry name" value="WD40_repeat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001680">
        <property type="entry name" value="WD40_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22872">
        <property type="entry name" value="BTK-BINDING PROTEIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22872:SF6">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE HERC1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00632">
        <property type="entry name" value="HECT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00622">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00400">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25390">
        <property type="entry name" value="WD40_RLD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00633">
        <property type="entry name" value="RCCNDNSATION"/>
    </dbReference>
    <dbReference type="SMART" id="SM00119">
        <property type="entry name" value="HECTc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00449">
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00320">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56204">
        <property type="entry name" value="Hect, E3 ligase catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50985">
        <property type="entry name" value="RCC1/BLIP-II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50978">
        <property type="entry name" value="WD40 repeat-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50188">
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50237">
        <property type="entry name" value="HECT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00626">
        <property type="entry name" value="RCC1_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50012">
        <property type="entry name" value="RCC1_3"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00678">
        <property type="entry name" value="WD_REPEATS_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50082">
        <property type="entry name" value="WD_REPEATS_2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50294">
        <property type="entry name" value="WD_REPEATS_REGION"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0677">Repeat</keyword>
    <keyword evidence="8" id="KW-0808">Transferase</keyword>
    <keyword evidence="10 11" id="KW-0833">Ubl conjugation pathway</keyword>
    <keyword evidence="7 12" id="KW-0853">WD repeat</keyword>
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        <location>
            <begin position="777"/>
            <end position="968"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="12">
        <location>
            <begin position="2199"/>
            <end position="2240"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="12">
        <location>
            <begin position="2397"/>
            <end position="2428"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="12">
        <location>
            <begin position="2507"/>
            <end position="2548"/>
        </location>
    </feature>
    <feature type="repeat" description="RCC1" evidence="13">
        <location>
            <begin position="2761"/>
            <end position="2809"/>
        </location>
    </feature>
    <feature type="repeat" description="RCC1" evidence="13">
        <location>
            <begin position="2810"/>
            <end position="2864"/>
        </location>
    </feature>
    <feature type="repeat" description="RCC1" evidence="13">
        <location>
            <begin position="2865"/>
            <end position="2916"/>
        </location>
    </feature>
    <feature type="repeat" description="RCC1" evidence="13">
        <location>
            <begin position="2917"/>
            <end position="2968"/>
        </location>
    </feature>
    <feature type="repeat" description="RCC1" evidence="13">
        <location>
            <begin position="2970"/>
            <end position="3021"/>
        </location>
    </feature>
    <feature type="repeat" description="RCC1" evidence="13">
        <location>
            <begin position="3022"/>
            <end position="3073"/>
        </location>
    </feature>
    <feature type="repeat" description="RCC1" evidence="13">
        <location>
            <begin position="3074"/>
            <end position="3125"/>
        </location>
    </feature>
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        <location>
            <begin position="3265"/>
            <end position="3612"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="94"/>
            <end position="152"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="170"/>
            <end position="203"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="260"/>
            <end position="304"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="638"/>
            <end position="660"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1027"/>
            <end position="1061"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1181"/>
            <end position="1219"/>
        </location>
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        <location>
            <begin position="1391"/>
            <end position="1450"/>
        </location>
    </feature>
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            <begin position="1467"/>
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        </location>
    </feature>
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        <location>
            <begin position="1573"/>
            <end position="1651"/>
        </location>
    </feature>
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        <location>
            <begin position="2009"/>
            <end position="2028"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="111"/>
            <end position="121"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="129"/>
            <end position="140"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="260"/>
            <end position="276"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="14">
        <location>
            <begin position="639"/>
            <end position="650"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
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        </location>
    </feature>
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        <location>
            <begin position="1037"/>
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        </location>
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            <begin position="1185"/>
            <end position="1194"/>
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    </feature>
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            <begin position="1195"/>
            <end position="1209"/>
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            <begin position="1476"/>
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            <begin position="2011"/>
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            <position position="3575"/>
        </location>
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    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000885"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00004906"/>
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    </evidence>
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        <source>
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        </source>
    </evidence>
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    </evidence>
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        </source>
    </evidence>
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    </evidence>
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        <source>
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        </source>
    </evidence>
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        </source>
    </evidence>
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    </evidence>
    <evidence type="ECO:0000256" key="11">
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            <dbReference type="PROSITE-ProRule" id="PRU00104"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
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            <dbReference type="PROSITE-ProRule" id="PRU00235"/>
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    </evidence>
    <evidence type="ECO:0000256" key="14">
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            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
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        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50237"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034555.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="3625" mass="399199" checksum="E45B0183F44F2792" modified="2019-05-08" version="1">QALINFLEIPVHFNGSFLFSDWDSATLSNESLLDTVSRFVLSALLKHTNLLSQACGESRYQPGKALAEVYRCVYKVRSRLLACKNMELIQTRSSSRDRWMSENQESADIDPQEHSFTRTIDEEAEMEEQAERDREEGHPEQEDEEEEREHEVMTAGKIFQCFLSAREVARSRDRDRMNTGNGTRVDDQPPQSQQERRVSTDLPEGQDVYTAACNSVIHRCALLILGVSPVIEELQKRREEGQLQQPSTSTAEGIGLMTRSESLTAESRSTHASSNYRLIKSRSESDLSQPESDEEGYSLSGRRNVDLDLASSHRKRGAMHSQVEPLSDSWARLKHSRDWLYSSSCTFVESDFDLSKSLGVHTLIENVVSFVSGDVGNSPGFKEPEESMSTSPQASIIAMEQQQLRAELRLEALHQILVLLSGMEEKGSAPLMGSRQGPAFQSSSLLTSVRLQFLAGCFGLSTVGHAGAKGESVRLHHYQDGIRAAKRNIQIEIQVAVHKIYQQLSVTLERALQANKHHIEAQQRLLLVTVFALSVHYQPVDVSLAISTGLLNVLSQLCGTDTMLGQPLQLLPKTGISQLSTALKVASTRLLQILAITTGTYADKLSPKVVQSLLDLLCSQLKNLLSQAGVMLMASFGEGEGEENEEEEEEEKKLDSNGDNEKKDFRAALRKQHAAELHLGDFLVFLRRVVSSKAIQSKMASPKWTEVLLNIASQKCCSGVPLVGNLRTRLLALHVLEAVLPACESGVEDDQMAQVVERLFSLLSDCMWETPIAQAKYAIQIKEREQEMKLQKQGESEEEDENLPIQEVSFDPEKAQCCIVESGQILTHGSGGKGYGLASAGVTSGCYQWKFYIVKENRGNEGTCVGVSRWPVHDFNHRTTSDMWLYRAYSGNLYHNGEQTLTLSSFTQGDFITCVLDMEARTISFGKNGEEPKLAFEDVDAAELYPCVMFYSSNPGEKVKICDMQMRGTPRDLLPGDPICSPVAAVLAEATIQLIRILHRADRWTHCINKKMMERLHKIKMCLKESGQKLKKSRSVQSRDENDMREEKENKEEEKNKHSRHGLADLSELQLKILCVEVWPVLAVIGGVDAGLRVGGRCVHKQTGRHATLLGVVKEGSTSAKVQWDEAEITISFPTFWSPSDTPLYNLEPCEPLPFDVGRFRGLTATVLLDLTCLTGIHEDMGKQSAKRHEKKHRHETEDKGDIEQKTEGDTGSDTRSLLSADDIRGPITTCSKSENEIASFSLESTPLCMESQHQPIEGKKKNHEHIPRNHDVVQSEIRAVQLSYLYLGAMKSLSALLSCSKYAELLLIPKVLAENGHNSDCASSPVVHEDVEMRAALQFLMRHMVKRAVMRSPIKRALGLADLERAQAMIYKLVVHGLLEDQFGGKIKQEVDQQAEESEQAQQAQTPITTSPSASSTTSFMSSSLEDTTTATTPVTDTETVPASESPGVMPLSLLRQMFSSYPTTTVLPTRRAQTPPVSSLPTSPSDEVGRRQSLTSPDSQSARPANRTALSDPSSRLSTSPPPPAIAVPLLEMGFSLRQIAKAMEATGARGEADAQNITVLAMWMIEHPGNEEDEEPQSEGTADSHHGTVVSGSSGKSSEQCYLQSPGDIPSADATEMEEGFSESPDNMDHTENAASGSGPPSRGRSAVTRRHKFDLAARTLLARAAGLYRSVQAHRNQSRREGISLQQDPGALYDFNLDEELEIDLDDEAMEAMFGQDLASDNDILGMWIPEVLDWPTWHVCESEDREEVVVCELCESSVVSFNQHMKRNHPGCGRSANRQGYRSNGSYVDGWFGGECGSGNPYYLLCGSCREKYLALKSKSKTSTSERYKGQAPDLIGKQDSVYEEDWDMLDVDEDEKLTGEEEFELLAGPLGLNDRRIVPEAVQFPDSDPLGASVAMVTATNSMEETLMQIGCHGSIEKSSSGRITLGEQAAALVNPHDRVMALRRVTAAAQVLLARTMVMRALSLLSVSGSSCSLAAGLESLGLTDIRTLVRLMCLAAAGRAGLSTSPSTMSSSTERSRGVHSKTSKPISCLAYLSTAVGCLASNTPSAAKLLVQLCTQNLISAATGVNLTTVDDPIQRKFLPSFLRGIAEENKLVTSPNFVVTQALVALLADKGAKLRPNYDKAEIEKKGPLELANALAACCLSSRLSSQHRQWAAQQLVRTLAAHDRDNQISPQTLADMGGDLRKCSFIKLEAHQNRVMTCVWCNKKGLLATSGNDGTIRVWNVTKKQYSLQQTCVFNKLEDDSEESLGSPSDPSLSLICWSISGKYLAGALEKMVNIWQVNGGKGLLDLQPHWVSALAWPEEGPSATWTGDTSEVLLVGRMDGSLGLIEVVDVSTMHRLELEHCYRKDVSVTSLAWFSEDRPFAVGYSDGKLLIGTKEPLEKGGIVLVDAHKDTIVSVKWDPTGKILMTCAKEETVKLWRYVGGCWRRLHSLSHPAIVNSITWCSLPGKGPKLQLLLATYKIYTLVAKKSKSETLFYSQFSMYDGYRKAVDAKCIYQLRGHITPVRTVAFSSDGLVLVSGGLGGLMNIWSLRDGSVLQSVVIGSGAIQTAVWIPDVGVAACSNRSKDVLVVNCTSEWISSNHVLATCRTALKQQGVFGLNMAPCMRAFLERLPIMLQEQYAYEKPHVVCGDQLVHSPYMQCLASLAVGLHLDQLLCNPPIPPHHQNCLLDPSAWNPTEWAWLECFSTTIKAAEALTKGAQFPESFTVPDLEPVPEDELTLLMDNSKWINGMDEQIMSWATSRPEDWHLGGKCDVYLWGAGRHGQLAEAGRNVMIPVAAPSFSQAQQVICGQNCTFVIQANGTVLACGEGSYGRLGQGNSDDLHVLTVISALQGFVVTQLVTSCGSDGHSMALTESGEVFSWGDGDYGKLGHGNSDRQRRPRQIEALQGEEVAQMSCGFKHSAVVTADGKLFTFGNGDYGRLGLGNTSNKKLPERVTALEGYQIGQVACGLNHTIVMSTDGSMVWAFGDGDYGKLGLGNSTAKSSPQKVDILCGIGIKKVACGTQFSVALTKDGHVYTFGQDRLIGLPEGRARNHNRPQQVPVLSGIFIEDIAVGAEHTLALSSSGDVYAWGSNSEGQLGLGHTNHVREPTLITVLQGKNIRQISAGRCHSAAWTAPPVPPRAPGVSVPLQLGLPDAIPPQYGALKEVSIHTVRARLRLLYHFSDLMYSSWRLLNLSPNNQNSTSHYNAGTWGIVQGQLRPLLAPRVYTLPMVRSIGKTMVQGKNYGPQITVKRISTRGRKCKPIFVQIARQVVKLNASDLRLPSRAWKVKLVGEGADDAGGVFDDTITEMCQELETGVVDLLIPSPNATAEVGYNRDRFLFNSSACLDEHLMQFKFLGILMGVAIRTKKPLDLHLAPMVWKQLCCIPLTLEDLEEVDLLYVQTLNSILHIEDSGITEENFHEMIPLDSFVGQSADGKMVPIIPGGNSIPLTFSNRKEYVERAIEYRLHEIDRQVAAVREGMSWIVPVPLLSLLTAKQLEQMVCGMPEISVEVLKKVVRYREVDEQHQLVQWFWHTLEEFSNEERVLFMRFVSGRSRLPANTADISQRFQIMKVDRPYDSLPTSQTCFFQLRLPPYSSQLIMAERLRYAINNCRSIDMDNYMLSRNVDNAEGSDTDY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2C8</accession>
    <name>A0A452J2C8_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="18">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 13">Membrane</location>
            <topology evidence="1 13">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 13">Belongs to the integrin alpha chain family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2C8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039146.1">
        <property type="protein sequence ID" value="ENSGAGP00000034574.1"/>
        <property type="gene ID" value="ENSGAGG00000024571.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009897">
        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008305">
        <property type="term" value="C:integrin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033627">
        <property type="term" value="P:cell adhesion mediated by integrin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098609">
        <property type="term" value="P:cell-cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007160">
        <property type="term" value="P:cell-matrix adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007229">
        <property type="term" value="P:integrin-mediated signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050900">
        <property type="term" value="P:leukocyte migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1530:FF:000001">
        <property type="entry name" value="Integrin subunit alpha 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.930">
        <property type="entry name" value="Bicelle-embedded integrin alpha(iib) transmembrane segment"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.130">
        <property type="entry name" value="Integrin alpha, N-terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1460">
        <property type="entry name" value="Integrin domains. Chain A, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1510">
        <property type="entry name" value="ntegrin, alpha v. Chain A, domain 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1530">
        <property type="entry name" value="ntegrin, alpha v. Chain A, domain 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013517">
        <property type="entry name" value="FG-GAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013519">
        <property type="entry name" value="Int_alpha_beta-p"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000413">
        <property type="entry name" value="Integrin_alpha"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018184">
        <property type="entry name" value="Integrin_alpha_C_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013649">
        <property type="entry name" value="Integrin_alpha_Ig-like_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048285">
        <property type="entry name" value="Integrin_alpha_Ig-like_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048286">
        <property type="entry name" value="Integrin_alpha_Ig-like_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028994">
        <property type="entry name" value="Integrin_alpha_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032695">
        <property type="entry name" value="Integrin_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23220">
        <property type="entry name" value="INTEGRIN ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23220:SF90">
        <property type="entry name" value="INTEGRIN ALPHA-7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01839">
        <property type="entry name" value="FG-GAP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08441">
        <property type="entry name" value="Integrin_A_Ig_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20805">
        <property type="entry name" value="Integrin_A_Ig_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20806">
        <property type="entry name" value="Integrin_A_Ig_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01185">
        <property type="entry name" value="INTEGRINA"/>
    </dbReference>
    <dbReference type="SMART" id="SM00191">
        <property type="entry name" value="Int_alpha"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69318">
        <property type="entry name" value="Integrin alpha N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69179">
        <property type="entry name" value="Integrin domains"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51470">
        <property type="entry name" value="FG_GAP"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00242">
        <property type="entry name" value="INTEGRIN_ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6 13" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="8 13" id="KW-0401">Integrin</keyword>
    <keyword evidence="9 13" id="KW-0472">Membrane</keyword>
    <keyword evidence="10 13" id="KW-0675">Receptor</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="4 13" id="KW-0732">Signal</keyword>
    <keyword evidence="3 13" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 13" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="13">
        <location>
            <begin position="1"/>
            <end position="21"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5018813402" evidence="13">
        <location>
            <begin position="22"/>
            <end position="1108"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="13">
        <location>
            <begin position="1026"/>
            <end position="1048"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="12">
        <location>
            <begin position="242"/>
            <end position="297"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="12">
        <location>
            <begin position="298"/>
            <end position="359"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="12">
        <location>
            <begin position="360"/>
            <end position="415"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="12">
        <location>
            <begin position="419"/>
            <end position="478"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha first immunoglubulin-like" evidence="15">
        <location>
            <begin position="463"/>
            <end position="627"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha second immunoglobulin-like" evidence="16">
        <location>
            <begin position="629"/>
            <end position="797"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha third immunoglobulin-like" evidence="17">
        <location>
            <begin position="803"/>
            <end position="1013"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="902"/>
            <end position="924"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="903"/>
            <end position="916"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023037"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00803"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="RuleBase" id="RU003762"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="Pfam" id="PF08441"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF20805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF20806"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034574.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1108" mass="121167" checksum="0AEE2F33ECF93E78" modified="2019-05-08" version="1">MRGASGAWLCCLGFQLLAGAAFNLDGTDPLLKDGAGGSLFGFSVALHRQLRPEPASWMLVGAPQALALPGQKANRTGGLYACPLTREIDDCWRVPIDEGVDLHKESKENQWLGVSVKSQGAGGKIVTCAHLYESRNRVNQPLETRDVIGRCYVLSQDLTVSEELDGGEWKFCEGRPQGHDRFGFCQQGVSAGFTADKHYILFGAPGTYNWKGDLRVELVNQSSLELVTYDDGPYEAGGEKDQDPSLIPVPANSYFGFSVDSGKGLTRQDQLSFVTGAPRANHTGAVVILRRDNVNRLVPEVVLPGEQLTSSFGYAVAVVDLNSDGWMDLVVGAPHFFDRQEEIGGAAYVYINQAGRWADIRPIRLNGTSHSMFGIGLTALGDLNQDGFQDIAVGAPFDGAGKVFIYHGSSLGIVTKPAQVLDGEGVGVRTFGYSLSGGLDVDGNLYPDLLVGSLSDSVVLYRARPVIHVSKNVSLSPQNIDLESRNCPHHLGICVEVRACFSYTASPASYSPRVTLTYTLDADVDRQKLGQAPRVAFLDRKPTDPEHQFSDVVELPHQRAPSCVKATFQLQESIRDKLRPIVVMLTYSIEHAQSKRQVQAAALPPLMPVLNAQLPNSQRAEVNFLKQGCGEDKICQSNLQLRYQFCSRVGDADFLPLPRDADGTAVFAMSDQKDVALEIQVTNLPSDPAAPQRDGDDAHEALLTATFPEALPYSGVRAHDMWAGPGLEKQPCLPNLNGSQVQCELGNPMKRGAQGRFYLIVSTSGITLETRELELELELSTISEQPGLRPVTARARVVIELPLSVTGVAIPQRLFFGGRVQGESAIRTEGQVGSAVRYEVTVSNRGQSLNTLGSAFLNLMWPHEIANGKWLLYPLRLELAARPGQRVACSPAANPLRLALESHGQDRSRREAERPEVPSTGPWWHRAHSERKNVTLDCAQGTARCLVFRCPLHSFDRSAVLTAWGRLWNSTFLEEYPSVTSVELLVHANITVKSTIQNLVLKDAATQIPVTIYLDPSAVVARGVPWWIILIAVLAGVLVLGLLVSILWKVGFFKRPRHQQAAVPRYHAIKIPREERQLFRQEKTGAMQKKEWVTNWSEGSDGHVPVSG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2C9</accession>
    <name>A0A452J2C9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">RNA helicase</fullName>
            <ecNumber evidence="5">3.6.4.13</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="28">DEAH box protein 9</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="29">Nuclear DNA helicase II</fullName>
        </alternativeName>
    </protein>
    <organism evidence="35 36">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="36" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="35" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="35" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="37">
            <text>ATP + H2O = ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:13065"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="3.6.4.13"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Microtubule organizing center</location>
            <location evidence="1">Centrosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
            <location evidence="2">Nucleolus</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Nucleus</location>
            <location evidence="3">Nucleoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the DEAD box helicase family. DEAH subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.6.4.13" evidence="5"/>
    <dbReference type="AlphaFoldDB" id="A0A452J2C9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039012.1">
        <property type="protein sequence ID" value="ENSGAGP00000034456.1"/>
        <property type="gene ID" value="ENSGAGG00000024443.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005730">
        <property type="term" value="C:nucleolus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005654">
        <property type="term" value="C:nucleoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:1990904">
        <property type="term" value="C:ribonucleoprotein complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043138">
        <property type="term" value="F:3'-5' DNA helicase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033679">
        <property type="term" value="F:3'-5' DNA/RNA helicase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003725">
        <property type="term" value="F:double-stranded RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003724">
        <property type="term" value="F:RNA helicase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006353">
        <property type="term" value="P:DNA-templated transcription termination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006397">
        <property type="term" value="P:mRNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051028">
        <property type="term" value="P:mRNA transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045944">
        <property type="term" value="P:positive regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050684">
        <property type="term" value="P:regulation of mRNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006417">
        <property type="term" value="P:regulation of translation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031047">
        <property type="term" value="P:regulatory ncRNA-mediated gene silencing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008380">
        <property type="term" value="P:RNA splicing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd17972">
        <property type="entry name" value="DEXHc_DHX9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd19854">
        <property type="entry name" value="DSRM_DHX9_rpt1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd19855">
        <property type="entry name" value="DSRM_DHX9_rpt2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18791">
        <property type="entry name" value="SF2_C_RHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.20:FF:000026">
        <property type="entry name" value="ATP-dependent RNA helicase A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.20:FF:000028">
        <property type="entry name" value="ATP-dependent RNA helicase A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000677">
        <property type="entry name" value="ATP-dependent RNA helicase A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.1080:FF:000006">
        <property type="entry name" value="ATP-dependent RNA helicase A protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000284">
        <property type="entry name" value="probable ATP-dependent RNA helicase YTHDC2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.1080">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.20">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011709">
        <property type="entry name" value="DEAD-box_helicase_OB_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011545">
        <property type="entry name" value="DEAD/DEAH_box_helicase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044447">
        <property type="entry name" value="DHX9_DEXHc"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044445">
        <property type="entry name" value="DHX9_DSRM_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044446">
        <property type="entry name" value="DHX9_DSRM_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002464">
        <property type="entry name" value="DNA/RNA_helicase_DEAH_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014720">
        <property type="entry name" value="dsRBD_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048333">
        <property type="entry name" value="HA2_WH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007502">
        <property type="entry name" value="Helicase-assoc_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014001">
        <property type="entry name" value="Helicase_ATP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001650">
        <property type="entry name" value="Helicase_C-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18934">
        <property type="entry name" value="ATP-DEPENDENT RNA HELICASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18934:SF119">
        <property type="entry name" value="ATP-DEPENDENT RNA HELICASE A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00270">
        <property type="entry name" value="DEAD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00035">
        <property type="entry name" value="dsrm"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21010">
        <property type="entry name" value="HA2_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00271">
        <property type="entry name" value="Helicase_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07717">
        <property type="entry name" value="OB_NTP_bind"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04408">
        <property type="entry name" value="WHD_HA2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00487">
        <property type="entry name" value="DEXDc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00358">
        <property type="entry name" value="DSRM"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00847">
        <property type="entry name" value="HA2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00490">
        <property type="entry name" value="HELICc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54768">
        <property type="entry name" value="dsRNA-binding domain-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00690">
        <property type="entry name" value="DEAH_ATP_HELICASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50137">
        <property type="entry name" value="DS_RBD"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51192">
        <property type="entry name" value="HELICASE_ATP_BIND_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51194">
        <property type="entry name" value="HELICASE_CTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="22" id="KW-0010">Activator</keyword>
    <keyword evidence="16" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="8" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="26" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="20" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="14" id="KW-0347">Helicase</keyword>
    <keyword evidence="13" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="25" id="KW-0464">Manganese</keyword>
    <keyword evidence="10" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9" id="KW-0507">mRNA processing</keyword>
    <keyword evidence="24" id="KW-0508">mRNA splicing</keyword>
    <keyword evidence="15" id="KW-0509">mRNA transport</keyword>
    <keyword evidence="12" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="27" id="KW-0539">Nucleus</keyword>
    <keyword evidence="36" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="11" id="KW-0677">Repeat</keyword>
    <keyword evidence="18 31" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="21" id="KW-0943">RNA-mediated gene silencing</keyword>
    <keyword evidence="23" id="KW-0804">Transcription</keyword>
    <keyword evidence="19" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="7" id="KW-0806">Transcription termination</keyword>
    <keyword evidence="17" id="KW-0810">Translation regulation</keyword>
    <keyword evidence="6" id="KW-0813">Transport</keyword>
    <feature type="domain" description="DRBM" evidence="32">
        <location>
            <begin position="3"/>
            <end position="71"/>
        </location>
    </feature>
    <feature type="domain" description="DRBM" evidence="32">
        <location>
            <begin position="166"/>
            <end position="238"/>
        </location>
    </feature>
    <feature type="domain" description="Helicase ATP-binding" evidence="33">
        <location>
            <begin position="384"/>
            <end position="550"/>
        </location>
    </feature>
    <feature type="domain" description="Helicase C-terminal" evidence="34">
        <location>
            <begin position="622"/>
            <end position="795"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004604"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004642"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00008792"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012552"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022472"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022664"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022806"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022816"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022845"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00023158"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00023187"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00023211"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="ARBA" id="ARBA00031576"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="ARBA" id="ARBA00033216"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="ARBA" id="ARBA00047984"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="31">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00266"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="32">
        <source>
            <dbReference type="PROSITE" id="PS50137"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="33">
        <source>
            <dbReference type="PROSITE" id="PS51192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="34">
        <source>
            <dbReference type="PROSITE" id="PS51194"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="35">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034456.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="36">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1193" mass="134711" checksum="46CAD313D77D1CD4" modified="2019-05-08" version="1">MGDVKNFLYAWCGKRKVTPTYEIRAVGNKNRQKFMCEVRVEGFNYTGMGNSTNKKDAQSNAARDFLNYLVRLNEIKSEEIPALGVTPDFSACISQLLGEGLTFFLPLGGGPAPGVGSSGYGGAQWDRGANLKDYYSRKEEQEAQATLESEEVDLNAGLHGNWTLENAKARLNQFFQKEKVQGEYKYTQVGPDHNRSFIAEMTIYVKQLGRRIFAREHGSNKKLAAQSCALSLVRQLYHLGVIEPYTGQTKKKEGETVEPYEVVISPDLENQLNSIIQELSLEIMPQPEDPSYPVSLNIGKLAHFEPSVRQNQMGVVPWSPPQSNWNPWTSCNIDEGPLAYATPEQISMDLKNDLMYRLEQDQELQRILQERETLPVKKFEIEILDAIHHSPVVVIRGATGCGKTTQVPQYILDEYIKNGKAAECNIVVTQPRRISAVSVAERVSYERGEEPGQSCGYSVRFESVLPRPHASVMFCTVGVLLRKLEAGIRGISHVIVDEIHERDINTDFLMVVLRDVVEAYPEIRVILMSATIDTSMFCEYFFNCPIIEVYGRTFPVEEYFLEDCIQMTQFVPPLKDKKKKDKDEEGGEDDDANCNLICSDEYGPETKRCMAQLNEKETPFELIEALLKYIETLNVPGAVLVFLPGWNLIYTMQKHLEMNPHFGSHRYRILPLHSQIPRDEQRRVFDPVPSGVTKVILSTNIAETSITINDVVYVIDSCKQKVKLFTAHNNMTNYATVWASKTNLEQRKGRAGRVRAGFCFHLCSRARFERLETHMTPEMFRTPLHEIALSIKLLRLGGIGQFLAKAIEPPPLDAVIEAEHTLRELDALDSNDELTPLGRILAKLPIEPRLGKMMIMGCIFYVGDAVCTISAATCFPEPFISEGKRLGYVHRNFAGNRFSDHVALLSVFQAWDDARMGGEDAEIRFCEHKRLNMATLRMTWEAKVQLKDILINSGFPEECLMTQIFNNTGPDNNLDVVISLLAFGVYPNVCYHKEKRKILTTEGRNALIHKSSVNCPFSSQDMKYPSPFFVFGEKIRTRAISAKGMTLVSPLQLLLFASKKVLSDGEIILVDDWIKLRMSHDAAACVTALRAAMEALVVEVTKDPEIISQLDPVNERMLNVIRQISRPSAAGINLMAGNTRLVPICSRIQLDPWSRKTFLVTWKAETCQRPKCVSELPCQQELLLWDLVWKQAA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2D2</accession>
    <name>A0A452J2D2_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="18">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 13">Membrane</location>
            <topology evidence="1 13">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 13">Belongs to the integrin alpha chain family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2D2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039151.1">
        <property type="protein sequence ID" value="ENSGAGP00000034579.1"/>
        <property type="gene ID" value="ENSGAGG00000024571.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009897">
        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008305">
        <property type="term" value="C:integrin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033627">
        <property type="term" value="P:cell adhesion mediated by integrin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098609">
        <property type="term" value="P:cell-cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007160">
        <property type="term" value="P:cell-matrix adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007229">
        <property type="term" value="P:integrin-mediated signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050900">
        <property type="term" value="P:leukocyte migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1530:FF:000001">
        <property type="entry name" value="Integrin subunit alpha 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.930">
        <property type="entry name" value="Bicelle-embedded integrin alpha(iib) transmembrane segment"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.130">
        <property type="entry name" value="Integrin alpha, N-terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1460">
        <property type="entry name" value="Integrin domains. Chain A, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1510">
        <property type="entry name" value="ntegrin, alpha v. Chain A, domain 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1530">
        <property type="entry name" value="ntegrin, alpha v. Chain A, domain 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013517">
        <property type="entry name" value="FG-GAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013519">
        <property type="entry name" value="Int_alpha_beta-p"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000413">
        <property type="entry name" value="Integrin_alpha"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018184">
        <property type="entry name" value="Integrin_alpha_C_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013649">
        <property type="entry name" value="Integrin_alpha_Ig-like_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048285">
        <property type="entry name" value="Integrin_alpha_Ig-like_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048286">
        <property type="entry name" value="Integrin_alpha_Ig-like_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028994">
        <property type="entry name" value="Integrin_alpha_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032695">
        <property type="entry name" value="Integrin_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23220">
        <property type="entry name" value="INTEGRIN ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23220:SF90">
        <property type="entry name" value="INTEGRIN ALPHA-7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01839">
        <property type="entry name" value="FG-GAP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08441">
        <property type="entry name" value="Integrin_A_Ig_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20805">
        <property type="entry name" value="Integrin_A_Ig_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20806">
        <property type="entry name" value="Integrin_A_Ig_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01185">
        <property type="entry name" value="INTEGRINA"/>
    </dbReference>
    <dbReference type="SMART" id="SM00191">
        <property type="entry name" value="Int_alpha"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69318">
        <property type="entry name" value="Integrin alpha N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69179">
        <property type="entry name" value="Integrin domains"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51470">
        <property type="entry name" value="FG_GAP"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00242">
        <property type="entry name" value="INTEGRIN_ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6 13" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="8 13" id="KW-0401">Integrin</keyword>
    <keyword evidence="9 13" id="KW-0472">Membrane</keyword>
    <keyword evidence="10 13" id="KW-0675">Receptor</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="4 13" id="KW-0732">Signal</keyword>
    <keyword evidence="3 13" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 13" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="13">
        <location>
            <begin position="1"/>
            <end position="21"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5018811078" evidence="13">
        <location>
            <begin position="22"/>
            <end position="1104"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="13">
        <location>
            <begin position="1022"/>
            <end position="1044"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="12">
        <location>
            <begin position="294"/>
            <end position="355"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="12">
        <location>
            <begin position="356"/>
            <end position="411"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="12">
        <location>
            <begin position="415"/>
            <end position="474"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha first immunoglubulin-like" evidence="15">
        <location>
            <begin position="459"/>
            <end position="623"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha second immunoglobulin-like" evidence="16">
        <location>
            <begin position="625"/>
            <end position="793"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha third immunoglobulin-like" evidence="17">
        <location>
            <begin position="799"/>
            <end position="1009"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="898"/>
            <end position="920"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="899"/>
            <end position="912"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023037"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00803"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="RuleBase" id="RU003762"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="Pfam" id="PF08441"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF20805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF20806"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034579.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1104" mass="120653" checksum="FBD5A7E2E06B131D" modified="2019-05-08" version="1">MRGASGAWLCCLGFQLLAGAAFNLDGTDPLLKDGAGGSLFGFSVALHRQLRPEPASWMLVGAPQALALPGQKANRTGGLYACPLTREIDDCWRVPIDEGVDLHKESKENQWLGVSVKSQGAGGKIVTCAHLYESRNRVNQPLETRDVIGRCYVLSQDLTVSEELDGGEWKFCEGRPQGHDRFGFCQQGVSAGFTADKHYILFGAPGTYNWKGLLFMTNVHSADPDQLVYKTAEPSERLPGTVGDVAQNSNLGFSVDSGKGLTRQDQLSFVTGAPRANHTGAVVILRRDNVNRLVPEVVLPGEQLTSSFGYAVAVVDLNSDGWMDLVVGAPHFFDRQEEIGGAAYVYINQAGRWADIRPIRLNGTSHSMFGIGLTALGDLNQDGFQDIAVGAPFDGAGKVFIYHGSSLGIVTKPAQVLDGEGVGVRTFGYSLSGGLDVDGNLYPDLLVGSLSDSVVLYRARPVIHVSKNVSLSPQNIDLESRNCPHHLGICVEVRACFSYTASPASYSPRVTLTYTLDADVDRQKLGQAPRVAFLDRKPTDPEHQFSDVVELPHQRAPSCVKATFQLQESIRDKLRPIVVMLTYSIEHAQSKRQVQAAALPPLMPVLNAQLPNSQRAEVNFLKQGCGEDKICQSNLQLRYQFCSRVGDADFLPLPRDADGTAVFAMSDQKDVALEIQVTNLPSDPAAPQRDGDDAHEALLTATFPEALPYSGVRAHDMWAGPGLEKQPCLPNLNGSQVQCELGNPMKRGAQGRFYLIVSTSGITLETRELELELELSTISEQPGLRPVTARARVVIELPLSVTGVAIPQRLFFGGRVQGESAIRTEGQVGSAVRYEVTVSNRGQSLNTLGSAFLNLMWPHEIANGKWLLYPLRLELAARPGQRVACSPAANPLRLALESHGQDRSRREAERPEVPSTGPWWHRAHSERKNVTLDCAQGTARCLVFRCPLHSFDRSAVLTAWGRLWNSTFLEEYPSVTSVELLVHANITVKSTIQNLVLKDAATQIPVTIYLDPSAVVARGVPWWIILIAVLAGVLVLGLLVSILWKVGFFKRPRHQQAAVPRYHAIKIPREERQLFRQEKTGAMQKKEWVTNWSEGSDGHVPVSG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2D4</accession>
    <name>A0A452J2D4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Aggrecan core protein</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="13">Cartilage-specific proteoglycan core protein</fullName>
        </alternativeName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the aggrecan/versican proteoglycan family.</text>
    </comment>
    <comment type="caution">
        <text evidence="14">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000039138.1">
        <property type="protein sequence ID" value="ENSGAGP00000034566.1"/>
        <property type="gene ID" value="ENSGAGG00000024567.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072534">
        <property type="term" value="C:perineuronal net"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045202">
        <property type="term" value="C:synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005540">
        <property type="term" value="F:hyaluronic acid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007417">
        <property type="term" value="P:central nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010001">
        <property type="term" value="P:glial cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002052">
        <property type="term" value="P:positive regulation of neuroblast proliferation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001501">
        <property type="term" value="P:skeletal system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00033">
        <property type="entry name" value="CCP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd03588">
        <property type="entry name" value="CLECT_CSPGs"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00054">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="CDD" id="cd03517">
        <property type="entry name" value="Link_domain_CSPGs_modules_1_3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="CDD" id="cd03520">
        <property type="entry name" value="Link_domain_CSPGs_modules_2_4"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000451">
        <property type="entry name" value="aggrecan core protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000009">
        <property type="entry name" value="Aggrecan core protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000011">
        <property type="entry name" value="Aggrecan core protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000002">
        <property type="entry name" value="Hyaluronan proteoglycan link protein 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000472">
        <property type="entry name" value="Uncharacterized protein, isoform A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.70.10:FF:000003">
        <property type="entry name" value="Versican core protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000003">
        <property type="entry name" value="Versican core protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.70.10">
        <property type="entry name" value="Complement Module, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.100.10">
        <property type="entry name" value="Mannose-Binding Protein A, subunit A"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001304">
        <property type="entry name" value="C-type_lectin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016186">
        <property type="entry name" value="C-type_lectin-like/link_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018378">
        <property type="entry name" value="C-type_lectin_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033987">
        <property type="entry name" value="CSPG_CTLD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016187">
        <property type="entry name" value="CTDL_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001881">
        <property type="entry name" value="EGF-like_Ca-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000152">
        <property type="entry name" value="EGF-type_Asp/Asn_hydroxyl_site"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018097">
        <property type="entry name" value="EGF_Ca-bd_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050691">
        <property type="entry name" value="Hyaluronan_bind_Proteoglycan"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003006">
        <property type="entry name" value="Ig/MHC_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013106">
        <property type="entry name" value="Ig_V-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000538">
        <property type="entry name" value="Link_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035976">
        <property type="entry name" value="Sushi/SCR/CCP_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000436">
        <property type="entry name" value="Sushi_SCR_CCP_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22804:SF42">
        <property type="entry name" value="AGGRECAN CORE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22804">
        <property type="entry name" value="AGGRECAN/VERSICAN PROTEOGLYCAN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00008">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00059">
        <property type="entry name" value="Lectin_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00084">
        <property type="entry name" value="Sushi"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07686">
        <property type="entry name" value="V-set"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00193">
        <property type="entry name" value="Xlink"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01265">
        <property type="entry name" value="LINKMODULE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00032">
        <property type="entry name" value="CCP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00034">
        <property type="entry name" value="CLECT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00179">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00406">
        <property type="entry name" value="IGv"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00445">
        <property type="entry name" value="LINK"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56436">
        <property type="entry name" value="C-type lectin-like"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57535">
        <property type="entry name" value="Complement control module/SCR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00010">
        <property type="entry name" value="ASX_HYDROXYL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00615">
        <property type="entry name" value="C_TYPE_LECTIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50041">
        <property type="entry name" value="C_TYPE_LECTIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01186">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01187">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00290">
        <property type="entry name" value="IG_MHC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01241">
        <property type="entry name" value="LINK_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50963">
        <property type="entry name" value="LINK_2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50923">
        <property type="entry name" value="SUSHI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9 14" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="14" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="4" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="5" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8" id="KW-0654">Proteoglycan</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0964">Secreted</keyword>
    <keyword evidence="6 18" id="KW-0732">Signal</keyword>
    <keyword evidence="15" id="KW-0768">Sushi</keyword>
    <feature type="signal peptide" evidence="18">
        <location>
            <begin position="1"/>
            <end position="16"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019569100" description="Aggrecan core protein" evidence="18">
        <location>
            <begin position="17"/>
            <end position="2190"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="21">
        <location>
            <begin position="34"/>
            <end position="149"/>
        </location>
    </feature>
    <feature type="domain" description="Link" evidence="23">
        <location>
            <begin position="153"/>
            <end position="248"/>
        </location>
    </feature>
    <feature type="domain" description="Link" evidence="23">
        <location>
            <begin position="254"/>
            <end position="350"/>
        </location>
    </feature>
    <feature type="domain" description="Link" evidence="23">
        <location>
            <begin position="506"/>
            <end position="601"/>
        </location>
    </feature>
    <feature type="domain" description="Link" evidence="23">
        <location>
            <begin position="607"/>
            <end position="703"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="19">
        <location>
            <begin position="1897"/>
            <end position="1932"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="19">
        <location>
            <begin position="1934"/>
            <end position="1970"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="20">
        <location>
            <begin position="1983"/>
            <end position="2097"/>
        </location>
    </feature>
    <feature type="domain" description="Sushi" evidence="22">
        <location>
            <begin position="2101"/>
            <end position="2161"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="776"/>
            <end position="873"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="898"/>
            <end position="926"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1022"/>
            <end position="1044"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="2170"/>
            <end position="2190"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="17">
        <location>
            <begin position="822"/>
            <end position="837"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="17">
        <location>
            <begin position="2170"/>
            <end position="2182"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="199"/>
            <end position="220"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="297"/>
            <end position="318"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="552"/>
            <end position="573"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="650"/>
            <end position="671"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="14">
        <location>
            <begin position="1901"/>
            <end position="1911"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="14">
        <location>
            <begin position="1922"/>
            <end position="1931"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="14">
        <location>
            <begin position="1960"/>
            <end position="1969"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="2103"/>
            <end position="2146"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="2132"/>
            <end position="2159"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006838"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022974"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00039399"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00042947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00302"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00323"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50041"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS50923"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS50963"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034566.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2190" mass="232219" checksum="90C6FB178BF7C0DE" modified="2019-05-08" version="1">MTTLLLVFVCLRVITAAISVEVSDHGNTLSVSIPERSPLRVILGNSLTIPCYFINSLDHVTTAPYTPPLTPRIKWSRLSNGKEVVLLVATEGQVRINTEYKEVISLPNYPAIPTDATLEIRALRSNDTGIYRCEVMHGIEDSQDTIEVIVKGIVFHYRAISTXYTLDFEKAKQACIQNSAVIATPEQLQAAYDDGFDQCDAGWLADQTVRYPMHHPREGCYGDKDEFPGVRTYGIRETDETYDVYCYAEEMQGEVFYATSPEKFTFQEAAEKCLSLGTRLATTGQLYLAWKGGMDVCSAGWLADGSVRYPISKARPNCGGNLVGVRTVYLHVNQTGYPHPHSRYNAICYTDEDFLVPENFTIQTVTQTEVEIPLPYNVTEEEARGSIATLEPIDFTPTASELEEESTVTPDLFATGITAETAFPEAENVTEEGVTAVWMTPAEVTTSLLDTLFTSLAPEVSSAKEDLEVTATPELESASVEEMIVQVTAAPASGLAPEQPISPTGVVFHYRPATSRYSLTFTKAQQACLDNNAVIATPEQLQAAYQTGFHQCDAGWLRDQTVRYPIVTPRTQCVGDKDSSPGVRTYGMRPASEMYDVYCYIDRLKGEVFFATQADQFTLLEAQEYCESQNATLASVGQLHAAWKQGLDRCRAGWLADGSIRYPIVTPRPACGGDKPGVRTIYLHPNQTGFPDPLSRHHAFCFRALPSMDKDEFVEEQIVTQVIPGLEEVPSGEEPTVETEFATEPENQTAWGTEVFPTDISLLSVSPSAFPPVAAIPDEPSAETSLSGVPSGEISASGVLETSGQPSASGWIPGELASGTFEPSGEPSGIEESGVPSVDLDFSGFPSGLPSGDESGLPSGIIEGSGLPSGEEDLLVSASGIPDLSGVPSGLQEISGIPEISGLPSGEVSGVELTSGLPSGEYDESGLPSGFPTVSLVDTTLVEVVTTVAESQLEGKGTIGVSGEGGLSGLPSAEWDISGKVSGLPSGVDISGAPSGIPEISGAPSGIPDISGLPSGIDFSGAPSGIPDISGAPSGIPDISGFPSGIDFSGEPSGIPDISGLVDLSGLVSVVDRSGEPSSITFLNISLVEVTTPSVTEAEAKKFLEISGLPSGDEDISGTVSGILDVSGQPSGQIDFSGGVSGAPEMSGFPSGVIDISGYTSGVEVVSGLPSGVYDYSGLPSGFPTISLVDTTLVEVVTQTSVAQEVGEGPSGILEISGLPSGERETSGEVSGVMDINGLPSGTIDISGETSGISYFSGEISGLTDLSGESLAVTGVSGEASGLPEVTLVTSDLVEVVTKPTVSQELGGETAVTLPHIFESSGDGEIGREASAFPEINVETSGFHEISIETSTAPETNGETSALPEITIEISTVHETSGETSAISEISIETSTVHKTSDKISALPEISIETSTVYKTSSETSVLPEVNLETSGIVVVSGETSGAPEERREDFSGTLVTPLPFVSXDEDISGTVSGILDVSGQPSGQIDFSGGVSGAPEMSGFPSGVIDISGYTSGVEVVSGLPSGVYDYSGLPSGFPTISLVDTTLVEVVTQTSVAQEVGEGPSGILEISGLPSGERETSGEVSGVMDINGLPSGTIDISGETSGISYFSGEISGLTDLSGESLAVTGVSGEASGLPEVTLVTSDLVEVVTKPTVSQELGGETAVTLPHIFESSGDGEIGREASAFPEINVETSGFHEISIETSTAPETNGETSALPEITIEISTVHETSGETSAISEISIETSTVHKTSDKISALPEISIETSTVYKTSSETSVLPEVNLETSGIVVVSGETSGAPEERREDFSGTLVTPLPFVSGETSAPEVVISTSMPDIELTQEPMGPEEAWLEIQSSAPMVSEQETKIAIVQETLEISPTATAILPQVSPEATNVPEPTAEVIPKTCDEKPCGAGTCQERDGNITCVCLPGYMGDHCDIDIDECHSSPCLNGATCADGIDSFKCLCLPSYGGDLCEIDLEDCDEGWIKFHGHCYKHFEQRETWVEAETRCREHQSHLSSIITPEEQEFVNNYARDYQWLGLSDRAIEHDFRWSDGNSLQYENWRPNQPDNFFAAEEDCVVMIWHEQGEWNDVPCNYHLPFTCKKGTVACGDPPVVENARTFGRKKDRYEINSMLRYQCNQGYIQRHVPTIRCQPNGHWEEPRIACIDPSIYKRRLHKRSPRNRSRTNRRAAQTPTH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2D5</accession>
    <name>A0A452J2D5_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="13">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell projection</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 10">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2D5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039137.1">
        <property type="protein sequence ID" value="ENSGAGP00000034565.1"/>
        <property type="gene ID" value="ENSGAGG00000024461.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005902">
        <property type="term" value="C:microvillus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016459">
        <property type="term" value="C:myosin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000146">
        <property type="term" value="F:microfilament motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007015">
        <property type="term" value="P:actin filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030048">
        <property type="term" value="P:actin filament-based movement"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006897">
        <property type="term" value="P:endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.4820:FF:000003">
        <property type="entry name" value="Unconventional myosin ID"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.530:FF:000004">
        <property type="entry name" value="Unconventional myosin ID"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.720:FF:000009">
        <property type="entry name" value="Unconventional myosin-Id"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.4820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010926">
        <property type="entry name" value="Myosin_TH1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13140">
        <property type="entry name" value="MYOSIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13140:SF381">
        <property type="entry name" value="UNCONVENTIONAL MYOSIN-IG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06017">
        <property type="entry name" value="Myosin_TH1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51757">
        <property type="entry name" value="TH1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 10" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="4 10" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="5" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="9" id="KW-0966">Cell projection</keyword>
    <keyword evidence="7 10" id="KW-0505">Motor protein</keyword>
    <keyword evidence="6 10" id="KW-0518">Myosin</keyword>
    <keyword evidence="3 10" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Myosin motor" evidence="11">
        <location>
            <begin position="1"/>
            <end position="638"/>
        </location>
    </feature>
    <feature type="domain" description="TH1" evidence="12">
        <location>
            <begin position="749"/>
            <end position="892"/>
        </location>
    </feature>
    <feature type="region of interest" description="Actin-binding" evidence="10">
        <location>
            <begin position="515"/>
            <end position="537"/>
        </location>
    </feature>
    <feature type="binding site" evidence="10">
        <location>
            <begin position="56"/>
            <end position="63"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004316"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS51757"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034565.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="892" mass="102438" checksum="0EAEA6298A5B61AF" modified="2019-05-08" version="1">MILYTSIISPLSLLFSKLKRPSLFNLPSYGTLSKPLIILPRPSCCPLPILSVCLAGESGAGKTEASKYIMQYIAAITNPSQREEVERVKNVLLKSNCVLEAFGNAKTNRNDNSSRFGKYMDINFDFKGDPTGGHINNYLLEKSRILHQQPGERNFHSFYQVSWELLISLLLFPPPHPISQLLASDQSNFKDVADAMKVIGFSPEEIDSVYKILATILHLGNVSFVAEGDAVAIENRELVSQLAELTSTQPESLLRTLLYRAVATGGGEVIEKGHSAKEANYARDACAKAIYERLFCWIVGRINTVIEVKNYDARLHGKNTVIAVLDIYGFEIFDSNSFEQFCINYCNEKLQQLFIELILRQEQEEYQREGIEWQHIEYFNNQIIVDLVEQPHKGIISILDEACLTGGQVTDAMFLESMNTKLGKHPHYTSRKLCATDKSMEFGRDFRIKHYAGNVTYSVEGFIDKNKDTLFQDFKRLMYNSTDPVLRDMWPEGEVSITEVTKRPLTAATLFKNSMVALVENLASKEPYYVRCIKPNDQKSPLLFDAERCRHQVEYLGLLENVRVRRAGFAYRQPYDRFLLRYKMTCEYTWPNHLMPTDQEATRALIDQHGFQDDVAYGHTKLFIRMPQILFCLEQERAQLIPIIVLLLQKAWRGALARRRCRYLRAIYTIMGHYRRHKVKTYLLELVRRFQGVRSMADYGKGVEWPDPPAVLAQFQENSKLGHPIVKNIPPSDMGQIQAKVAAMEVLQGLRKDWGCQRNWGRDYLSSVAENPVMATQFTRRVQALRDKVHFSSVLFSCHVRKINRFNKSRDRAILVTDQHLYKLEPRKQYKVMRTLPLSMVTGVSVTSCQDQLVVFHTKNHDDLVICFHKMQPEEDNRVGELVGVLVHHFKT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2D7</accession>
    <name>A0A452J2D7_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="18">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 13">Membrane</location>
            <topology evidence="1 13">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 13">Belongs to the integrin alpha chain family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2D7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039143.1">
        <property type="protein sequence ID" value="ENSGAGP00000034571.1"/>
        <property type="gene ID" value="ENSGAGG00000024571.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009897">
        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008305">
        <property type="term" value="C:integrin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033627">
        <property type="term" value="P:cell adhesion mediated by integrin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098609">
        <property type="term" value="P:cell-cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007160">
        <property type="term" value="P:cell-matrix adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007229">
        <property type="term" value="P:integrin-mediated signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050900">
        <property type="term" value="P:leukocyte migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.930">
        <property type="entry name" value="Bicelle-embedded integrin alpha(iib) transmembrane segment"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.130">
        <property type="entry name" value="Integrin alpha, N-terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1460">
        <property type="entry name" value="Integrin domains. Chain A, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1510">
        <property type="entry name" value="ntegrin, alpha v. Chain A, domain 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1530">
        <property type="entry name" value="ntegrin, alpha v. Chain A, domain 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013517">
        <property type="entry name" value="FG-GAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013519">
        <property type="entry name" value="Int_alpha_beta-p"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000413">
        <property type="entry name" value="Integrin_alpha"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018184">
        <property type="entry name" value="Integrin_alpha_C_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013649">
        <property type="entry name" value="Integrin_alpha_Ig-like_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048285">
        <property type="entry name" value="Integrin_alpha_Ig-like_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048286">
        <property type="entry name" value="Integrin_alpha_Ig-like_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028994">
        <property type="entry name" value="Integrin_alpha_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032695">
        <property type="entry name" value="Integrin_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23220">
        <property type="entry name" value="INTEGRIN ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23220:SF90">
        <property type="entry name" value="INTEGRIN ALPHA-7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01839">
        <property type="entry name" value="FG-GAP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08441">
        <property type="entry name" value="Integrin_A_Ig_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20805">
        <property type="entry name" value="Integrin_A_Ig_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20806">
        <property type="entry name" value="Integrin_A_Ig_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01185">
        <property type="entry name" value="INTEGRINA"/>
    </dbReference>
    <dbReference type="SMART" id="SM00191">
        <property type="entry name" value="Int_alpha"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69318">
        <property type="entry name" value="Integrin alpha N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69179">
        <property type="entry name" value="Integrin domains"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51470">
        <property type="entry name" value="FG_GAP"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00242">
        <property type="entry name" value="INTEGRIN_ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6 13" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="8 13" id="KW-0401">Integrin</keyword>
    <keyword evidence="9 13" id="KW-0472">Membrane</keyword>
    <keyword evidence="10 13" id="KW-0675">Receptor</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="4 13" id="KW-0732">Signal</keyword>
    <keyword evidence="3 13" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 13" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="13">
        <location>
            <begin position="1"/>
            <end position="21"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5018813839" evidence="13">
        <location>
            <begin position="22"/>
            <end position="1110"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="13">
        <location>
            <begin position="1028"/>
            <end position="1050"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="12">
        <location>
            <begin position="338"/>
            <end position="399"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="12">
        <location>
            <begin position="400"/>
            <end position="455"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="12">
        <location>
            <begin position="459"/>
            <end position="518"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha first immunoglubulin-like" evidence="15">
        <location>
            <begin position="503"/>
            <end position="667"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha second immunoglobulin-like" evidence="16">
        <location>
            <begin position="669"/>
            <end position="837"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha third immunoglobulin-like" evidence="17">
        <location>
            <begin position="843"/>
            <end position="959"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="942"/>
            <end position="964"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="943"/>
            <end position="956"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023037"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00803"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="RuleBase" id="RU003762"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="Pfam" id="PF08441"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF20805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF20806"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034571.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1110" mass="121123" checksum="AAAB9257A525F07D" modified="2019-05-08" version="1">MRGASGAWLCCLGFQLLAGAAFNLDGTDPLLKDGAGGSLFGFSVALHRQLRPEPASWMLVGAPQALALPGQKANRTGGLYACPLTREIDDCWRVPIDEGVDLHKESKENQWLGVSVKSQGAGGKIVTCAHLYESRNRVNQPLETRDVIGRCYVLSQDLTVSEELDGGEWKFCEGRPQGHDRFGFCQQGVSAGFTADKHYILFGAPGTYNWKGDLRVELVNQSSLELVTYDDGPYEAGGEKDQDPSLIPVPANSYFGLLFMTNVHSADPDQLVYKTAEPSERLPGTVGDVAQNSNLGFSVDSGKGLTRQDQLSFVTGAPRANHTGAVVILRRDNVNRLVPEVVLPGEQLTSSFGYAVAVVDLNSDGWMDLVVGAPHFFDRQEEIGGAAYVYINQAGRWADIRPIRLNGTSHSMFGIGLTALGDLNQDGFQDIAVGAPFDGAGKVFIYHGSSLGIVTKPAQVLDGEGVGVRTFGYSLSGGLDVDGNLYPDLLVGSLSDSVVLYRARPVIHVSKNVSLSPQNIDLESRNCPHHLGICVEVRACFSYTASPASYSPRVTLTYTLDADVDRQKLGQAPRVAFLDRKPTDPEHQFSDVVELPHQRAPSCVKATFQLQESIRDKLRPIVVMLTYSIEHAQSKRQVQAAALPPLMPVLNAQLPNSQRAEVNFLKQGCGEDKICQSNLQLRYQFCSRVGDADFLPLPRDADGTAVFAMSDQKDVALEIQVTNLPSDPAAPQRDGDDAHEALLTATFPEALPYSGVRAHDMWAGPGLEKQPCLPNLNGSQVQCELGNPMKRGAQGRFYLIVSTSGITLETRELELELELSTISEQPGLRPVTARARVVIELPLSVTGVAIPQRLFFGGRVQGESAIRTEGQVGSAVRYEVTVSNRGQSLNTLGSAFLNLMWPHEIANGKWLLYPLRLELAARPGQRVACSPAANPLRLALESHGQDRSRREAERPEVPSTGPWWHRAHSERKNVTLEYPSVTSVELLVHANITVKSTIQNLVLKDAATQIPVTIYLDPSAVVARGVPWWIILIAVLAGVLVLGLLVSILWKVGFFKRPRHQQAAVPRYHAIKIPREERQLFRQEKTGAMQKKEWVTNWSEGSDGHVPVSG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2D8</accession>
    <name>A0A452J2D8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Major facilitator superfamily (MFS) profile domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the major facilitator superfamily. Sugar transporter (TC 2.A.1.1) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2D8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039023.1">
        <property type="protein sequence ID" value="ENSGAGP00000034467.1"/>
        <property type="gene ID" value="ENSGAGG00000024490.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005903">
        <property type="term" value="C:brush border"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055056">
        <property type="term" value="F:D-glucose transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046323">
        <property type="term" value="P:D-glucose import"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070837">
        <property type="term" value="P:dehydroascorbic acid transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd17431">
        <property type="entry name" value="MFS_GLUT_Class1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1250.20:FF:000029">
        <property type="entry name" value="solute carrier family 2, facilitated glucose transporter member 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1250.20">
        <property type="entry name" value="MFS general substrate transporter like domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045263">
        <property type="entry name" value="GLUT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020846">
        <property type="entry name" value="MFS_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005828">
        <property type="entry name" value="MFS_sugar_transport-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036259">
        <property type="entry name" value="MFS_trans_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003663">
        <property type="entry name" value="Sugar/inositol_transpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005829">
        <property type="entry name" value="Sugar_transporter_CS"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00879">
        <property type="entry name" value="SP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23503">
        <property type="entry name" value="SOLUTE CARRIER FAMILY 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23503:SF27">
        <property type="entry name" value="SOLUTE CARRIER FAMILY 2, FACILITATED GLUCOSE TRANSPORTER MEMBER 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00083">
        <property type="entry name" value="Sugar_tr"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00171">
        <property type="entry name" value="SUGRTRNSPORT"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103473">
        <property type="entry name" value="MFS general substrate transporter"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50850">
        <property type="entry name" value="MFS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00217">
        <property type="entry name" value="SUGAR_TRANSPORT_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 6" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 6" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="3 6" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="5" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="31"/>
            <end position="51"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="63"/>
            <end position="81"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="87"/>
            <end position="112"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="124"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="154"/>
            <end position="175"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="240"/>
            <end position="263"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="275"/>
            <end position="296"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="303"/>
            <end position="325"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="331"/>
            <end position="349"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="370"/>
            <end position="394"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="400"/>
            <end position="418"/>
        </location>
    </feature>
    <feature type="domain" description="Major facilitator superfamily (MFS) profile" evidence="7">
        <location>
            <begin position="1"/>
            <end position="424"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="RuleBase" id="RU003346"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50850"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034467.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="461" mass="50249" checksum="2A3E012CD7AD3247" modified="2019-05-08" version="1">FFQYGYSLGVINAPQRANLEELDVGKSTLTLYWSLSVSIFAIGGMISSFTVGWVGDKLGRVKAMLAVNVLSIIGNLLMGLAKFGPSHILIIAGRAITGLYCGLSSGLVPMYVGEISPTALRGALGTLHQLAIVTGILLSQVLGLDFLLGNDDMWPLLLGLSGAAALLQFFLLLLCPESPRYLYIKLGKVEEAKKSLKRLRGDCDPTKEITEMEKEKQEAASEKKVSIRKLFSSSSYRQPIIVALMVQISQQFSGINAIFYYSTNIFVRAGVDQPVYATIGVGVVNTIFTIISVFLVERTGRRSLFIAGLIGMLASAVAMTVGLVLLGTFAWMSYVSMVAIFLFVIFFEVGPGPIPWFIVAELFSQGPRPAAIAIAGFCNWTCNFIIGMCFQYIADLCGPYVFTIFAVLLLGFILFAHFKVPETKGKSFEEISAAFRRKRRSAIKGHKAATELEDLRGSEEA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2D9</accession>
    <name>A0A452J2D9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Aggrecan core protein</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="13">Cartilage-specific proteoglycan core protein</fullName>
        </alternativeName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the aggrecan/versican proteoglycan family.</text>
    </comment>
    <comment type="caution">
        <text evidence="14">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000039134.1">
        <property type="protein sequence ID" value="ENSGAGP00000034562.1"/>
        <property type="gene ID" value="ENSGAGG00000024567.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072534">
        <property type="term" value="C:perineuronal net"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045202">
        <property type="term" value="C:synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005540">
        <property type="term" value="F:hyaluronic acid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007417">
        <property type="term" value="P:central nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010001">
        <property type="term" value="P:glial cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002052">
        <property type="term" value="P:positive regulation of neuroblast proliferation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001501">
        <property type="term" value="P:skeletal system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00033">
        <property type="entry name" value="CCP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd03588">
        <property type="entry name" value="CLECT_CSPGs"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00054">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd03517">
        <property type="entry name" value="Link_domain_CSPGs_modules_1_3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="CDD" id="cd03520">
        <property type="entry name" value="Link_domain_CSPGs_modules_2_4"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000451">
        <property type="entry name" value="aggrecan core protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000009">
        <property type="entry name" value="Aggrecan core protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000011">
        <property type="entry name" value="Aggrecan core protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000002">
        <property type="entry name" value="Hyaluronan proteoglycan link protein 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.70.10:FF:000003">
        <property type="entry name" value="Versican core protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000003">
        <property type="entry name" value="Versican core protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.70.10">
        <property type="entry name" value="Complement Module, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.100.10">
        <property type="entry name" value="Mannose-Binding Protein A, subunit A"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001304">
        <property type="entry name" value="C-type_lectin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016186">
        <property type="entry name" value="C-type_lectin-like/link_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018378">
        <property type="entry name" value="C-type_lectin_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033987">
        <property type="entry name" value="CSPG_CTLD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016187">
        <property type="entry name" value="CTDL_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050691">
        <property type="entry name" value="Hyaluronan_bind_Proteoglycan"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003006">
        <property type="entry name" value="Ig/MHC_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013106">
        <property type="entry name" value="Ig_V-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000538">
        <property type="entry name" value="Link_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035976">
        <property type="entry name" value="Sushi/SCR/CCP_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000436">
        <property type="entry name" value="Sushi_SCR_CCP_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22804:SF54">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22804">
        <property type="entry name" value="AGGRECAN/VERSICAN PROTEOGLYCAN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00059">
        <property type="entry name" value="Lectin_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00084">
        <property type="entry name" value="Sushi"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07686">
        <property type="entry name" value="V-set"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00193">
        <property type="entry name" value="Xlink"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01265">
        <property type="entry name" value="LINKMODULE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00032">
        <property type="entry name" value="CCP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00034">
        <property type="entry name" value="CLECT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00406">
        <property type="entry name" value="IGv"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00445">
        <property type="entry name" value="LINK"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56436">
        <property type="entry name" value="C-type lectin-like"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57535">
        <property type="entry name" value="Complement control module/SCR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00615">
        <property type="entry name" value="C_TYPE_LECTIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50041">
        <property type="entry name" value="C_TYPE_LECTIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01186">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00290">
        <property type="entry name" value="IG_MHC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01241">
        <property type="entry name" value="LINK_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50963">
        <property type="entry name" value="LINK_2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50923">
        <property type="entry name" value="SUSHI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9 14" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="14" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="4" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="5" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8" id="KW-0654">Proteoglycan</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0964">Secreted</keyword>
    <keyword evidence="6 18" id="KW-0732">Signal</keyword>
    <keyword evidence="15" id="KW-0768">Sushi</keyword>
    <feature type="signal peptide" evidence="18">
        <location>
            <begin position="1"/>
            <end position="16"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019350400" description="Aggrecan core protein" evidence="18">
        <location>
            <begin position="17"/>
            <end position="2152"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="21">
        <location>
            <begin position="34"/>
            <end position="149"/>
        </location>
    </feature>
    <feature type="domain" description="Link" evidence="23">
        <location>
            <begin position="153"/>
            <end position="248"/>
        </location>
    </feature>
    <feature type="domain" description="Link" evidence="23">
        <location>
            <begin position="254"/>
            <end position="350"/>
        </location>
    </feature>
    <feature type="domain" description="Link" evidence="23">
        <location>
            <begin position="506"/>
            <end position="601"/>
        </location>
    </feature>
    <feature type="domain" description="Link" evidence="23">
        <location>
            <begin position="607"/>
            <end position="703"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="19">
        <location>
            <begin position="1897"/>
            <end position="1932"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="20">
        <location>
            <begin position="1945"/>
            <end position="2059"/>
        </location>
    </feature>
    <feature type="domain" description="Sushi" evidence="22">
        <location>
            <begin position="2063"/>
            <end position="2123"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="776"/>
            <end position="873"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="898"/>
            <end position="926"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1022"/>
            <end position="1044"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="2132"/>
            <end position="2152"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="17">
        <location>
            <begin position="822"/>
            <end position="837"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="17">
        <location>
            <begin position="2132"/>
            <end position="2144"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="199"/>
            <end position="220"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="297"/>
            <end position="318"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="552"/>
            <end position="573"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="650"/>
            <end position="671"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="14">
        <location>
            <begin position="1901"/>
            <end position="1911"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="14">
        <location>
            <begin position="1922"/>
            <end position="1931"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="2065"/>
            <end position="2108"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="2094"/>
            <end position="2121"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006838"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022974"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00039399"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00042947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00302"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00323"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50041"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS50923"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS50963"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034562.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2152" mass="228257" checksum="06F69A219BF869E8" modified="2019-05-08" version="1">MTTLLLVFVCLRVITAAVSVEVSDHGNTLSVSIPERSPLRVILGNSLTIPCYFINSLDHVTTAPYTPPLTPRIKWSRLSNGKEVVLLVATEGQVRINTEYKEVISLPNYPAIPTDATLEIRALRSNDTGIYRCEVMHGIEDSQDTIEVIVKGIVFHYRAISTXYTLDFEKAKQACIQNSAVIATPEQLQAAYDDGFDQCDAGWLADQTVRYPMHHPREGCYGDKDEFPGVRTYGIRETDETYDVYCYAEEMQGEVFYATSPEKFTFQEAAEKCLSLGTRLATTGQLYLAWKGGMDVCSAGWLADGSVRYPISKARPNCGGNLVGVRTVYLHVNQTGYPHPHSRYNAICYTDEDFLVPENFTIQTVTQTEVEIPLPYNVTEEEARGSIATLEPIDFTPTASELEEESTVTPDLFATGITAETAFPEAENVTEEGVTAVWMTPAEVTTSLLDTLFTSLAPEVSSAKEDLEVTATPELESASVEEMIVQVTAAPASGLAPEQPISPTGVVFHYRPATSRYSLTFTKAQQACLDNNAVIATPEQLQAAYQTGFHQCDAGWLRDQTVRYPIVTPRTQCVGDKDSSPGVRTYGMRPASEMYDVYCYIDRLKGEVFFATQADQFTLLEAQEYCESQNATLASVGQLHAAWKQGLDRCRAGWLADGSIRYPIVTPRPACGGDKPGVRTIYLHPNQTGFPDPLSRHHAFCFRALPSMDKDEFVEEQIVTQVIPGLEEVPSGEEPTVETEFATEPENQTAWGTEVFPTDISLLSVSPSAFPPVAAIPDEPSAETSLSGVPSGEISASGVLETSGQPSASGWIPGELASGTFEPSGEPSGIEESGVPSVDLDFSGFPSGLPSGDESGLPSGIIEGSGLPSGEEDLLVSASGIPDLSGVPSGLQEISGIPEISGLPSGEVSGVELTSGLPSGEYDESGLPSGFPTVSLVDTTLVEVVTTVAESQLEGKGTIGVSGEGGLSGLPSAEWDISGKVSGLPSGVDISGAPSGIPEISGAPSGIPDISGLPSGIDFSGAPSGIPDISGAPSGIPDISGFPSGIDFSGEPSGIPDISGLVDLSGLVSVVDRSGEPSSITFLNISLVEVTTPSVTEAEAKKFLEISGLPSGDEDISGTVSGILDVSGQPSGQIDFSGGVSGAPEMSGFPSGVIDISGYTSGVEVVSGLPSGVYDYSGLPSGFPTISLVDTTLVEVVTQTSVAQEVGEGPSGILEISGLPSGERETSGEVSGVMDINGLPSGTIDISGETSGISYFSGEISGLTDLSGESLAVTGVSGEASGLPEVTLVTSDLVEVVTKPTVSQELGGETAVTLPHIFESSGDGEIGREASAFPEINVETSGFHEISIETSTAPETNGETSALPEITIEISTVHETSGETSAISEISIETSTVHKTSDKISALPEISIETSTVYKTSSETSVLPEVNLETSGIVVVSGETSGAPEERREDFSGTLVTPLPFVSXDEDISGTVSGILDVSGQPSGQIDFSGGVSGAPEMSGFPSGVIDISGYTSGVEVVSGLPSGVYDYSGLPSGFPTISLVDTTLVEVVTQTSVAQEVGEGPSGILEISGLPSGERETSGEVSGVMDINGLPSGTIDISGETSGISYFSGEISGLTDLSGESLAVTGVSGEASGLPEVTLVTSDLVEVVTKPTVSQELGGETAVTLPHIFESSGDGEIGREASAFPEINVETSGFHEISIETSTAPETNGETSALPEITIEISTVHETSGETSAISEISIETSTVHKTSDKISALPEISIETSTVYKTSSETSVLPEVNLETSGIVVVSGETSGAPEERREDFSGTLVTPLPFVSGETSAPEVVISTSMPDIELTQEPMGPEEAWLEIQSSAPMVSEQETKIAIVQETLEISPTATAILPQVSPEATNVPEPTAEVIPKTCDEKPCGAGTCQERDGNITCVCLPGYMGDHCDIDLEDCDEGWIKFHGHCYKHFEQRETWVEAETRCREHQSHLSSIITPEEQEFVNNYARDYQWLGLSDRAIEHDFRWSDGNSLQYENWRPNQPDNFFAAEEDCVVMIWHEQGEWNDVPCNYHLPFTCKKGTVACGDPPVVENARTFGRKKDRYEINSMLRYQCNQGYIQRHVPTIRCQPNGHWEEPRIACIDPSIYKRRLHKRSPRNRSRTNRRAAQTPTH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2E1</accession>
    <name>A0A452J2E1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Sodium/hydrogen exchanger</fullName>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Recycling endosome membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 13">Belongs to the monovalent cation:proton antiporter 1 (CPA1) transporter (TC 2.A.36) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2E1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039161.1">
        <property type="protein sequence ID" value="ENSGAGP00000034589.1"/>
        <property type="gene ID" value="ENSGAGG00000024543.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055038">
        <property type="term" value="C:recycling endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015386">
        <property type="term" value="F:potassium:proton antiporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015385">
        <property type="term" value="F:sodium:proton antiporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051453">
        <property type="term" value="P:regulation of intracellular pH"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098719">
        <property type="term" value="P:sodium ion import across plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.1330">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018422">
        <property type="entry name" value="Cation/H_exchanger_CPA1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006153">
        <property type="entry name" value="Cation/H_exchanger_TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004709">
        <property type="entry name" value="NaH_exchanger"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002090">
        <property type="entry name" value="NHE-6/7/9"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00840">
        <property type="entry name" value="b_cpa1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10110">
        <property type="entry name" value="SODIUM/HYDROGEN EXCHANGER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10110:SF94">
        <property type="entry name" value="SODIUM_HYDROGEN EXCHANGER 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00999">
        <property type="entry name" value="Na_H_Exchanger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01084">
        <property type="entry name" value="NAHEXCHNGR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01088">
        <property type="entry name" value="NAHEXCHNGR6"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="13" id="KW-0050">Antiport</keyword>
    <keyword evidence="5" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="7" id="KW-0967">Endosome</keyword>
    <keyword evidence="10 13" id="KW-0406">Ion transport</keyword>
    <keyword evidence="11 15" id="KW-0472">Membrane</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0915">Sodium</keyword>
    <keyword evidence="12 13" id="KW-0739">Sodium transport</keyword>
    <keyword evidence="6 13 15" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 15" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="4 13" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="77"/>
            <end position="94"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="106"/>
            <end position="131"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="143"/>
            <end position="166"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="178"/>
            <end position="200"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="221"/>
            <end position="243"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="263"/>
            <end position="287"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="308"/>
            <end position="329"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="341"/>
            <end position="363"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="375"/>
            <end position="394"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="400"/>
            <end position="421"/>
        </location>
    </feature>
    <feature type="domain" description="Cation/H+ exchanger transmembrane" evidence="16">
        <location>
            <begin position="65"/>
            <end position="429"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="577"/>
            <end position="597"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004195"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007367"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023053"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023201"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="RuleBase" id="RU003722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF00999"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034589.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="597" mass="65806" checksum="8D34B1F4802247CA" modified="2019-05-08" version="1">MIYGVLMGVILRYGIHVPSDVNNVTLSCQVQTSPATLLVNVSGKYYEYTLKGEISPHELNNVQDNEMLRKVTFDPEVFFNILLPPIIFYAGYSLKRRHFFRNLGSILAYAFLGTAISCFVIGSIMYGCVALMKVTGQLGGDFYFTDCLLFGAIVSATDPVTVLAIFHELQVDVELYALLFGESVLNDAVAIVLSSSIVAYQPAGDNSHTFDVTAMFKSIGIFLGIFSGSFAMGAATGVVTALVTKFTKLREFPLLETGLFFLMSWSTFLLAEACGFTGVVAVLFCGITQAHYTYNNLSTESQHRTKQLFELLNFLAENFIFSYMGLTLFTFQNHDFNPTFIVGAFFAIFLGRAANIYPLSFLLNLGRRNKIGTNLQHMMMFAGLRGAMAFALAIRDTATYARQMMFSTTLLIVFFTVWVFGGGTTAMLSCLHIRVGVDSDQEHLGVPESERRSTKAESAWLFRLWYNFDHNYLKPLLTHSGPPLTTTLPSCCGPVARCLTSPQAYENQEQLKDDDSDLILNDGDISLTYGDSTVNADSITSSGTSRRIVGNSSEDALDRELAFGDHELVIRGTRLVLPMDDSEPPSSLVDSARHGPA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2E3</accession>
    <name>A0A452J2E3_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="14">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the requiem/DPF family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2E3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039029.1">
        <property type="protein sequence ID" value="ENSGAGP00000034472.1"/>
        <property type="gene ID" value="ENSGAGG00000024499.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071565">
        <property type="term" value="C:nBAF complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd15692">
        <property type="entry name" value="PHD1_DPF3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd15530">
        <property type="entry name" value="PHD2_d4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.40.10:FF:000005">
        <property type="entry name" value="zinc finger protein isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
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    </dbReference>
    <dbReference type="InterPro" id="IPR025750">
        <property type="entry name" value="DPF1-3_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011011">
        <property type="entry name" value="Znf_FYVE_PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001965">
        <property type="entry name" value="Znf_PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019787">
        <property type="entry name" value="Znf_PHD-finger"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45888">
        <property type="entry name" value="HL01030P-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45888:SF10">
        <property type="entry name" value="ZINC FINGER PROTEIN DPF3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14051">
        <property type="entry name" value="DPF1-3_N"/>
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    </dbReference>
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        <property type="entry name" value="PHD"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00249">
        <property type="entry name" value="PHD"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57903">
        <property type="entry name" value="FYVE/PHD zinc finger"/>
        <property type="match status" value="2"/>
    </dbReference>
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        <property type="entry name" value="ZF_PHD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="8" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 10" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="177"/>
            <end position="205"/>
        </location>
    </feature>
    <feature type="domain" description="PHD-type" evidence="12">
        <location>
            <begin position="238"/>
            <end position="298"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="204"/>
            <end position="233"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010539"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50016"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034472.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="358" mass="40454" checksum="550B331A345D4F0C" modified="2019-05-08" version="1">MQSLTLCSSPPGRLGDQFYKEAIEHCRSYNSRLCAERSVRLPFLDSQTGVAQNNCYIWMEKRHRGPGLAPGQLYTYPARCWRKKRRLHPPEDSRLKLLEIKPEVDLPLKKDGFTSEGTTLEALLRGEGIEKKVDTKEEDTIQEIQVKKCIYIGHARGSGGGRRRNDAASQDDHDKPFVCDICGKRYKNRPGLSYHYAHTHLASEEGDEAQEQETRSPPVHRNENHKPQKGPDGAIIPNNYCDFCLGGSSMNKKSGCPEELVSCSDCGRSGHPTCLQFTTNMTEAVKTYKWQCIECKSCSLCGTSENDDQLLFCDDCDRGYHMYCLNPPVSEPPEGNMFWKSTLRATLIKQMDTGNNFI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2E5</accession>
    <name>A0A452J2E5_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="6">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="caution">
        <text evidence="2">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2E5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039147.1">
        <property type="protein sequence ID" value="ENSGAGP00000034575.1"/>
        <property type="gene ID" value="ENSGAGG00000024575.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0038023">
        <property type="term" value="F:signaling receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00054">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="CDD" id="cd00057">
        <property type="entry name" value="FA58C"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.260:FF:000002">
        <property type="entry name" value="Coagulation factor VIII"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.260">
        <property type="entry name" value="Galactose-binding domain-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000421">
        <property type="entry name" value="FA58C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008979">
        <property type="entry name" value="Galactose-bd-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050633">
        <property type="entry name" value="Neuropilin_MCO_CoagFactor"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46806">
        <property type="entry name" value="F5/8 TYPE C DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46806:SF11">
        <property type="entry name" value="MILK FAT GLOBULE EGF AND FACTOR V_VIII DOMAIN CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00008">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00754">
        <property type="entry name" value="F5_F8_type_C"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00231">
        <property type="entry name" value="FA58C"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49785">
        <property type="entry name" value="Galactose-binding domain-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01186">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01285">
        <property type="entry name" value="FA58C_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01286">
        <property type="entry name" value="FA58C_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50022">
        <property type="entry name" value="FA58C_3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1 2" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="2" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="3">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019280094" evidence="3">
        <location>
            <begin position="23"/>
            <end position="430"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="5">
        <location>
            <begin position="24"/>
            <end position="62"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="5">
        <location>
            <begin position="65"/>
            <end position="107"/>
        </location>
    </feature>
    <feature type="domain" description="F5/8 type C" evidence="4">
        <location>
            <begin position="110"/>
            <end position="266"/>
        </location>
    </feature>
    <feature type="domain" description="F5/8 type C" evidence="4">
        <location>
            <begin position="271"/>
            <end position="428"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="2">
        <location>
            <begin position="33"/>
            <end position="50"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="2">
        <location>
            <begin position="52"/>
            <end position="61"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="2">
        <location>
            <begin position="97"/>
            <end position="106"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50022"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034575.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="430" mass="48277" checksum="D973A60E6157A163" modified="2019-05-08" version="1">MKESSPLRTLFGAILAVSVVLAVTGDFCDVNHCENGGTCLTGSNETPFFCICPEGFTGIDCNDTEKGPCHSNPCHNNGECQLLPNRGDVFTEYVCKCPVGYAGTHCQKSCAVLLGMEGRAIADAQLSASSVYYGFLGLQRWGPELARLNNKGIVNAWTSSNYDKNPWIQVNLLRKMRLIGIITQGARRAGWAEYVHTFKVAHSLDGRVFVFYRDENQDRDKVFPGNMNNDGMKSNTFNPPITAQYIRIYPVMCHRACTLRFELVGCELNGCSEPLGMKSRLISDQQITASSVYKTWGIDGFTWYPHYARLDKTGKTNAWTALNNDQSEWLQIDLLTQKKVTGVVTQGARDFGHIQYVAAYKVAYSDDGTSWTVYKDNRTNSSKIFQGNYDNTSHKKNVFDVPFYTRFVRILPVAWHNRITLRMELLGCDE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2E7</accession>
    <name>A0A452J2E7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Sodium/hydrogen exchanger</fullName>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Recycling endosome membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 13">Belongs to the monovalent cation:proton antiporter 1 (CPA1) transporter (TC 2.A.36) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2E7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039153.1">
        <property type="protein sequence ID" value="ENSGAGP00000034581.1"/>
        <property type="gene ID" value="ENSGAGG00000024543.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055038">
        <property type="term" value="C:recycling endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015386">
        <property type="term" value="F:potassium:proton antiporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015385">
        <property type="term" value="F:sodium:proton antiporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051453">
        <property type="term" value="P:regulation of intracellular pH"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098719">
        <property type="term" value="P:sodium ion import across plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.1330">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018422">
        <property type="entry name" value="Cation/H_exchanger_CPA1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006153">
        <property type="entry name" value="Cation/H_exchanger_TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004709">
        <property type="entry name" value="NaH_exchanger"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002090">
        <property type="entry name" value="NHE-6/7/9"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00840">
        <property type="entry name" value="b_cpa1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10110">
        <property type="entry name" value="SODIUM/HYDROGEN EXCHANGER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10110:SF94">
        <property type="entry name" value="SODIUM_HYDROGEN EXCHANGER 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00999">
        <property type="entry name" value="Na_H_Exchanger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01084">
        <property type="entry name" value="NAHEXCHNGR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01088">
        <property type="entry name" value="NAHEXCHNGR6"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="13" id="KW-0050">Antiport</keyword>
    <keyword evidence="5" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="7" id="KW-0967">Endosome</keyword>
    <keyword evidence="10 13" id="KW-0406">Ion transport</keyword>
    <keyword evidence="11 15" id="KW-0472">Membrane</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0915">Sodium</keyword>
    <keyword evidence="12 13" id="KW-0739">Sodium transport</keyword>
    <keyword evidence="6 13 15" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 15" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="4 13" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="71"/>
            <end position="88"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="100"/>
            <end position="117"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="179"/>
            <end position="196"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="208"/>
            <end position="234"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="323"/>
            <end position="350"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="362"/>
            <end position="391"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="412"/>
            <end position="433"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="445"/>
            <end position="467"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="479"/>
            <end position="498"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="504"/>
            <end position="525"/>
        </location>
    </feature>
    <feature type="domain" description="Cation/H+ exchanger transmembrane" evidence="16">
        <location>
            <begin position="81"/>
            <end position="533"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1"/>
            <end position="25"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004195"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007367"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023053"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023201"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="RuleBase" id="RU003722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF00999"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034581.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="664" mass="73141" checksum="5D5627322BC52042" modified="2019-05-08" version="1">ASGRGWAIAPAVTPPPRAKQALRRAAPRCAAGDRAGAAARGEAAARPVALEEEIVSEKAAEESHRQDSANLLIFILLLTLTILTIWLFKHRRARFLHETGLAMIYGVLMGVILRYGIHVPSDVNNVTLSCQVQTSPATLLVNVSGKYYEYTLKGEISPHELNNVQDNEMLRKVTFDPEVFFNILLPPIIFYAGYSLKRRHFFRNLGSILAYAFLGTAISCFVIGSIMYGCVALMKVTGQLGGDFYFTDCLLFGAIVSATDPVTVLAIFHELQVDVELYALLFGESVLNDAVAIVLSSSIVAYQPAGDNSHTFDVTAMFKSIGIFLGIFSGSFAMGAATGVVTALISFSLFTKLREFPLLETGLFFLMSWSTFLLAEACGFTGVVAVLFCGITQAHYTYNNLSTESQHRTKQLFELLNFLAENFIFSYMGLTLFTFQNHDFNPTFIVGAFFAIFLGRAANIYPLSFLLNLGRRNKIGTNLQHMMMFAGLRGAMAFALAIRDTATYARQMMFSTTLLIVFFTVWVFGGGTTAMLSCLHIRVGVDSDQEHLGVPESERRSTKAESAWLFRLWYNFDHNYLKPLLTHSGPPLTTTLPSCCGPVARCLTSPQAYENQEQLKDDDSDLILNDGDISLTYGDSTVNADSITSSGTSSLELNCSFHSHKYTH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2F2</accession>
    <name>A0A452J2F2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Myosin IG</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="6">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000039158.1">
        <property type="protein sequence ID" value="ENSGAGP00000034586.1"/>
        <property type="gene ID" value="ENSGAGG00000024461.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005902">
        <property type="term" value="C:microvillus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016459">
        <property type="term" value="C:myosin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000146">
        <property type="term" value="F:microfilament motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007015">
        <property type="term" value="P:actin filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030048">
        <property type="term" value="P:actin filament-based movement"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006897">
        <property type="term" value="P:endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.4820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010926">
        <property type="entry name" value="Myosin_TH1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13140">
        <property type="entry name" value="MYOSIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13140:SF381">
        <property type="entry name" value="UNCONVENTIONAL MYOSIN-IG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06017">
        <property type="entry name" value="Myosin_TH1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51757">
        <property type="entry name" value="TH1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 6" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="2 6" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="4 6" id="KW-0505">Motor protein</keyword>
    <keyword evidence="3 6" id="KW-0518">Myosin</keyword>
    <keyword evidence="1 6" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="7">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5046961090" description="Myosin IG" evidence="7">
        <location>
            <begin position="23"/>
            <end position="937"/>
        </location>
    </feature>
    <feature type="domain" description="Myosin motor" evidence="8">
        <location>
            <begin position="1"/>
            <end position="654"/>
        </location>
    </feature>
    <feature type="domain" description="TH1" evidence="9">
        <location>
            <begin position="771"/>
            <end position="937"/>
        </location>
    </feature>
    <feature type="region of interest" description="Actin-binding" evidence="6">
        <location>
            <begin position="531"/>
            <end position="553"/>
        </location>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <begin position="72"/>
            <end position="79"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51757"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034586.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="12">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="937" mass="107555" checksum="6BC085DEE7B191A3" modified="2025-10-08" version="2">MXXXXXXXXXLVLTILSSLVSSADFATSLFTPFSRSFMNKLNRIGPRTDPWGTTKPRPSCCPLPILSVCLAGESGAGKTEASKYIMQYIAAITNPSQREEVERVKNVLLKSNCVLEAFGNAKTNRNDNSSRFGKYMDINFDFKGDPTGGHINNYLLEKSRILHQQPGERNFHSFYQVSWELLISLLLFPPPHPISQLLASDQSNFKDVADAMKVIGFSPEEIDSVYKILATILHLGNVSFVAEGDAVAIENRELVSQLAELTSTQPESLLRTLLYRAVATGGGEVIEKGHSAKEANYARDACAKAIYERLFCWIVGRINTVIEVKNYDARLHGKNTVIAVLDIYGFEIFDSNSFEQFCINYCNEKLQQLFIELILRQEQEEYQREGIEWQHIEYFNNQIIVDLVEQPHKGIISILDEACLTGGQVTDAMFLESMNTKLGKHPHYTSRKLCATDKSMEFGRDFRIKHYAGNVTYSVEGFIDKNKDTLFQDFKRLMYNSTDPVLRDMWPEGEVSITEVTKRPLTAATLFKNSMVALVENLASKEPYYVRCIKPNDQKSPLLFDAERCRHQVEYLGLLENVRVRRAGFAYRQPYDRFLLRYKMTCEYTWPNHLMPTDQEATRALIDQHGFQDDVAYGHTKLFIRMPQILFCLEQERAQLIPIIVLLLQKAWRGALARRRCRYLRAIYTIMGHYRRHKVKTYLLELVRRFQGVRSMADYGKGVEWPDPPAVLAQFQENSKLLFCRWRARQIVKNIPPSDMGQIQAKVAAMEVLQGLRKDWGCQRNWGRDYLSSVSVRPGELPGALRDKVHFSSVLFSCHVRKINRFNKSRDRAILVTDQHLYKLEPRKQYKVMRTLPLSMVTGVSVTSCQDQLVVFHTKNHDDLVICFHKMQPEEDNRVGELVGVLVHHFKTRAAGARWMSVYRSPGRIDRGRRVERGGWG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2F5</accession>
    <name>A0A452J2F5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Myosin IG</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="6">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000039157.1">
        <property type="protein sequence ID" value="ENSGAGP00000034585.1"/>
        <property type="gene ID" value="ENSGAGG00000024461.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005902">
        <property type="term" value="C:microvillus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016459">
        <property type="term" value="C:myosin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000146">
        <property type="term" value="F:microfilament motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007015">
        <property type="term" value="P:actin filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030048">
        <property type="term" value="P:actin filament-based movement"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006897">
        <property type="term" value="P:endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.4820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010926">
        <property type="entry name" value="Myosin_TH1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13140">
        <property type="entry name" value="MYOSIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13140:SF381">
        <property type="entry name" value="UNCONVENTIONAL MYOSIN-IG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06017">
        <property type="entry name" value="Myosin_TH1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51757">
        <property type="entry name" value="TH1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 6" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="2 6" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="4 6" id="KW-0505">Motor protein</keyword>
    <keyword evidence="3 6" id="KW-0518">Myosin</keyword>
    <keyword evidence="1 6" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="7">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5047395846" description="Myosin IG" evidence="7">
        <location>
            <begin position="23"/>
            <end position="918"/>
        </location>
    </feature>
    <feature type="domain" description="Myosin motor" evidence="8">
        <location>
            <begin position="1"/>
            <end position="654"/>
        </location>
    </feature>
    <feature type="domain" description="TH1" evidence="9">
        <location>
            <begin position="771"/>
            <end position="918"/>
        </location>
    </feature>
    <feature type="region of interest" description="Actin-binding" evidence="6">
        <location>
            <begin position="531"/>
            <end position="553"/>
        </location>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <begin position="72"/>
            <end position="79"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51757"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034585.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="12">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="918" mass="105036" checksum="9B90FA6AB8006846" modified="2025-10-08" version="2">MXXXXXXXXXLVLTILSSLVSSADFATSLFTPFSRSFMNKLNRIGPRTDPWGTTKPRPSCCPLPILSVCLAGESGAGKTEASKYIMQYIAAITNPSQREEVERVKNVLLKSNCVLEAFGNAKTNRNDNSSRFGKYMDINFDFKGDPTGGHINNYLLEKSRILHQQPGERNFHSFYQVSWELLISLLLFPPPHPISQLLASDQSNFKDVADAMKVIGFSPEEIDSVYKILATILHLGNVSFVAEGDAVAIENRELVSQLAELTSTQPESLLRTLLYRAVATGGGEVIEKGHSAKEANYARDACAKAIYERLFCWIVGRINTVIEVKNYDARLHGKNTVIAVLDIYGFEIFDSNSFEQFCINYCNEKLQQLFIELILRQEQEEYQREGIEWQHIEYFNNQIIVDLVEQPHKGIISILDEACLTGGQVTDAMFLESMNTKLGKHPHYTSRKLCATDKSMEFGRDFRIKHYAGNVTYSVEGFIDKNKDTLFQDFKRLMYNSTDPVLRDMWPEGEVSITEVTKRPLTAATLFKNSMVALVENLASKEPYYVRCIKPNDQKSPLLFDAERCRHQVEYLGLLENVRVRRAGFAYRQPYDRFLLRYKMTCEYTWPNHLMPTDQEATRALIDQHGFQDDVAYGHTKLFIRMPQILFCLEQERAQLIPIIVLLLQKAWRGALARRRCRYLRAIYTIMGHYRRHKVKTYLLELVRRFQGVRSMADYGKGVEWPDPPAVLAQFQENTPSPHCRWRARQIVKNIPPSDMGQIQAKVAAMEVLQGLRKDWGCQRNWGRDYLSSVSVRPGELPGALRDKVHFSSVLFSCHVRKINRFNKSRDRAILVTDQHLYKLEPRKQYKVMRTLPLSMVSALLHCLLGPTHAHIYTHMPTRARAHTHTAHWQFPGSQAVIGWVKRSAGDILEELQGSGEG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2G2</accession>
    <name>A0A452J2G2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Sodium/hydrogen exchanger</fullName>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Recycling endosome membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 13">Belongs to the monovalent cation:proton antiporter 1 (CPA1) transporter (TC 2.A.36) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2G2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039171.1">
        <property type="protein sequence ID" value="ENSGAGP00000034597.1"/>
        <property type="gene ID" value="ENSGAGG00000024543.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
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        <property type="term" value="C:recycling endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015386">
        <property type="term" value="F:potassium:proton antiporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015385">
        <property type="term" value="F:sodium:proton antiporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051453">
        <property type="term" value="P:regulation of intracellular pH"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098719">
        <property type="term" value="P:sodium ion import across plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.1330">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018422">
        <property type="entry name" value="Cation/H_exchanger_CPA1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006153">
        <property type="entry name" value="Cation/H_exchanger_TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004709">
        <property type="entry name" value="NaH_exchanger"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002090">
        <property type="entry name" value="NHE-6/7/9"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00840">
        <property type="entry name" value="b_cpa1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10110">
        <property type="entry name" value="SODIUM/HYDROGEN EXCHANGER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10110:SF94">
        <property type="entry name" value="SODIUM_HYDROGEN EXCHANGER 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00999">
        <property type="entry name" value="Na_H_Exchanger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01084">
        <property type="entry name" value="NAHEXCHNGR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01088">
        <property type="entry name" value="NAHEXCHNGR6"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="13" id="KW-0050">Antiport</keyword>
    <keyword evidence="5" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="7" id="KW-0967">Endosome</keyword>
    <keyword evidence="10 13" id="KW-0406">Ion transport</keyword>
    <keyword evidence="11 15" id="KW-0472">Membrane</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0915">Sodium</keyword>
    <keyword evidence="12 13" id="KW-0739">Sodium transport</keyword>
    <keyword evidence="6 13 15" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 15" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="4 13" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="38"/>
            <end position="55"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="67"/>
            <end position="84"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="146"/>
            <end position="163"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="175"/>
            <end position="201"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="290"/>
            <end position="317"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="329"/>
            <end position="358"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="379"/>
            <end position="400"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="412"/>
            <end position="434"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="446"/>
            <end position="465"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="471"/>
            <end position="492"/>
        </location>
    </feature>
    <feature type="domain" description="Cation/H+ exchanger transmembrane" evidence="16">
        <location>
            <begin position="48"/>
            <end position="500"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1"/>
            <end position="28"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="648"/>
            <end position="668"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="14">
        <location>
            <begin position="1"/>
            <end position="15"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="16"/>
            <end position="28"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004195"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00007367"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023053"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023201"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="RuleBase" id="RU003722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF00999"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034597.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="668" mass="73573" checksum="0BF02F928782A889" modified="2019-05-08" version="1">RSRGGAQGAPGGGGARRAEEIVSEKAAEESHRQDSANLLIFILLLTLTILTIWLFKHRRARFLHETGLAMIYGVLMGVILRYGIHVPSDVNNVTLSCQVQTSPATLLVNVSGKYYEYTLKGEISPHELNNVQDNEMLRKVTFDPEVFFNILLPPIIFYAGYSLKRRHFFRNLGSILAYAFLGTAISCFVIGSIMYGCVALMKVTGQLGGDFYFTDCLLFGAIVSATDPVTVLAIFHELQVDVELYALLFGESVLNDAVAIVLSSSIVAYQPAGDNSHTFDVTAMFKSIGIFLGIFSGSFAMGAATGVVTALISFSLFTKLREFPLLETGLFFLMSWSTFLLAEACGFTGVVAVLFCGITQAHYTYNNLSTESQHRTKQLFELLNFLAENFIFSYMGLTLFTFQNHDFNPTFIVGAFFAIFLGRAANIYPLSFLLNLGRRNKIGTNLQHMMMFAGLRGAMAFALAIRDTATYARQMMFSTTLLIVFFTVWVFGGGTTAMLSCLHIRVGVDSDQEHLGVPESERRSTKAESAWLFRLWYNFDHNYLKPLLTHSGPPLTTTLPSCCGPVARCLTSPQAYENQEQLKDDDSDLILNDGDISLTYGDSTVNADSITSSGTSRRIVGNSSEDALDRELAFGDHELVIRGTRLVLPMDDSEPPSSLVDSARHGPA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2J1</accession>
    <name>A0A452J2J1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Nuclear protein MDM1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Microtubule-binding protein that negatively regulates centriole duplication. Binds to and stabilizes microtubules.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Microtubule organizing center</location>
            <location evidence="1">Centrosome</location>
            <location evidence="1">Centriole</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the MDM1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2J1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039202.1">
        <property type="protein sequence ID" value="ENSGAGP00000034627.1"/>
        <property type="gene ID" value="ENSGAGG00000024610.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005814">
        <property type="term" value="C:centriole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
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        <property type="term" value="C:microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
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        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
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        <property type="term" value="F:microtubule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
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        <property type="term" value="P:negative regulation of centriole replication"/>
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        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060041">
        <property type="term" value="P:retina development in camera-type eye"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="entry name" value="MDM1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR32078">
        <property type="entry name" value="NUCLEAR PROTEIN MDM1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR32078:SF1">
        <property type="entry name" value="NUCLEAR PROTEIN MDM1"/>
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    </dbReference>
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    </dbReference>
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    <keyword evidence="6" id="KW-0493">Microtubule</keyword>
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    <keyword evidence="11" id="KW-0812">Transmembrane</keyword>
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            <begin position="176"/>
            <end position="199"/>
        </location>
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        </location>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00010494"/>
        </source>
    </evidence>
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        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00045771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034627.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="205" mass="23541" checksum="6CB12C49233777AE" modified="2019-05-08" version="1">MPVRFKGLSEYKRNFKWKKSHLSEFCNPSQEQKSSRAGLRSDQLGITREPNFISKRRVPYFNPQISESFEWRGDSDLEGYLDDAPEIEGAGPAESHTNNHSNDRNLENIETPEAPRLPKKIRSHSADSRIESAITLAENSRKKLSPPTPVNQKAVFVPPKKHLEKMDNGVGSFENLFPFLIVYGFIIITFDIAVHWEWLCSKKRV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2J2</accession>
    <name>A0A452J2J2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">BHLH domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2J2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039099.1">
        <property type="protein sequence ID" value="ENSGAGP00000034530.1"/>
        <property type="gene ID" value="ENSGAGG00000024549.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046983">
        <property type="term" value="F:protein dimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd18930">
        <property type="entry name" value="bHLHzip_MXI1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.280.10:FF:000014">
        <property type="entry name" value="Max dimerization protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.280.10">
        <property type="entry name" value="Helix-loop-helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011598">
        <property type="entry name" value="bHLH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036638">
        <property type="entry name" value="HLH_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11969">
        <property type="entry name" value="MAX DIMERIZATION, MAD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11969:SF13">
        <property type="entry name" value="MAX-INTERACTING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00010">
        <property type="entry name" value="HLH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00353">
        <property type="entry name" value="HLH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47459">
        <property type="entry name" value="HLH, helix-loop-helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50888">
        <property type="entry name" value="BHLH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="6" id="KW-0539">Nucleus</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0678">Repressor</keyword>
    <keyword evidence="5" id="KW-0804">Transcription</keyword>
    <keyword evidence="3" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="BHLH" evidence="9">
        <location>
            <begin position="101"/>
            <end position="153"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="70"/>
            <end position="110"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="193"/>
            <end position="262"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="143"/>
            <end position="184"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="8">
        <location>
            <begin position="82"/>
            <end position="92"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="93"/>
            <end position="104"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="8">
        <location>
            <begin position="207"/>
            <end position="217"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="242"/>
            <end position="262"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022491"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50888"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034530.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="262" mass="29476" checksum="25116C5A088E8957" modified="2019-05-08" version="1">MCSGQQQEMPAADMSRCPFSDIFSSSGDSMEKINTFLQNVQILLEAASYLEKIEKENKKCEHGYASTFPLLPSPGLQDPKPMRRLSRVRKHSSGGSSTSTANRSTHNELEKNRRAHLRLCLERLKVLIPLGPDCTRHTTLGLLNKAKAHIKKLEEAERRSQHQLENLEREQRFLKRRLEQLQGPQEIERIRMDSIGSTISSDRSDSEREEIEVDVESTEFSHGEVDDISTTSISDIDDHSSLQSVGSDEGYSSASVKLSFAS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2J5</accession>
    <name>A0A452J2J5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Nuclear protein MDM1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Microtubule-binding protein that negatively regulates centriole duplication. Binds to and stabilizes microtubules.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Microtubule organizing center</location>
            <location evidence="1">Centrosome</location>
            <location evidence="1">Centriole</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the MDM1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2J5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039201.1">
        <property type="protein sequence ID" value="ENSGAGP00000034626.1"/>
        <property type="gene ID" value="ENSGAGG00000024610.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005814">
        <property type="term" value="C:centriole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005874">
        <property type="term" value="C:microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008017">
        <property type="term" value="F:microtubule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046600">
        <property type="term" value="P:negative regulation of centriole replication"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060041">
        <property type="term" value="P:retina development in camera-type eye"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029136">
        <property type="entry name" value="MDM1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR32078">
        <property type="entry name" value="NUCLEAR PROTEIN MDM1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR32078:SF1">
        <property type="entry name" value="NUCLEAR PROTEIN MDM1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15501">
        <property type="entry name" value="MDM1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="11" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-0493">Microtubule</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="11" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="176"/>
            <end position="197"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="80"/>
            <end position="125"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004114"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00010494"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00013508"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00045771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034626.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="223" mass="25603" checksum="B4DADCD0BE935697" modified="2019-05-08" version="1">MPVRFKGLSEYKRNFKWKKSHLSEFCNPSQEQKSSRAGLRSDQLGITREPNFISKRRVPYFNPQISESFEWRGDSDLEGYLDDAPEIEGAGPAESHTNNHSNDRNLENIETPEAPRLPKKIRSHSADSRIESAITLAENSRKKLSPPTPVNQKAVFVPPKKHLEKMDNGVGSFENLFPFLIVYGYMMWLSECCLVLFNPLYIGYMMWLSECCSRNLNIGISRL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2J7</accession>
    <name>A0A452J2J7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">Glutamate receptor</fullName>
        </recommendedName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="21">Receptor for glutamate that functions as a ligand-gated ion channel in the central nervous system and plays an important role in excitatory synaptic transmission. L-glutamate acts as an excitatory neurotransmitter at many synapses in the central nervous system.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>Ca(2+)(in) = Ca(2+)(out)</text>
            <dbReference type="Rhea" id="RHEA:29671"/>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="16 21">Postsynaptic cell membrane</location>
            <topology evidence="16 21">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="21">Belongs to the glutamate-gated ion channel (TC 1.A.10.1) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2J7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039207.1">
        <property type="protein sequence ID" value="ENSGAGP00000034632.1"/>
        <property type="gene ID" value="ENSGAGG00000024584.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015276">
        <property type="term" value="F:ligand-gated monoatomic ion channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0038023">
        <property type="term" value="F:signaling receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd06382">
        <property type="entry name" value="PBP1_iGluR_Kainate"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.2300:FF:000010">
        <property type="entry name" value="Glutamate ionotropic receptor kainate type subunit 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.190.10:FF:000210">
        <property type="entry name" value="Glutamate receptor ionotropic, kainate 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.190.10:FF:000240">
        <property type="entry name" value="Glutamate receptor ionotropic, kainate 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.70:FF:000010">
        <property type="entry name" value="Putative glutamate receptor ionotropic kainate 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.70">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.2300">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.190.10">
        <property type="entry name" value="Periplasmic binding protein-like II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001828">
        <property type="entry name" value="ANF_lig-bd_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019594">
        <property type="entry name" value="Glu/Gly-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001508">
        <property type="entry name" value="Iono_Glu_rcpt_met"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015683">
        <property type="entry name" value="Ionotropic_Glu_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001320">
        <property type="entry name" value="Iontro_rcpt_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028082">
        <property type="entry name" value="Peripla_BP_I"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18966">
        <property type="entry name" value="IONOTROPIC GLUTAMATE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01094">
        <property type="entry name" value="ANF_receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00060">
        <property type="entry name" value="Lig_chan"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Lig_chan-Glu_bd"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="NMDARECEPTOR"/>
    </dbReference>
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    </dbReference>
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        <property type="entry name" value="PBPe"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53822">
        <property type="entry name" value="Periplasmic binding protein-like I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53850">
        <property type="entry name" value="Periplasmic binding protein-like II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 21" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="10 20" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="15 21" id="KW-0407">Ion channel</keyword>
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    <keyword evidence="14 21" id="KW-1071">Ligand-gated ion channel</keyword>
    <keyword evidence="9 21" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="13 21" id="KW-0628">Postsynaptic cell membrane</keyword>
    <keyword evidence="11 21" id="KW-0675">Receptor</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <keyword evidence="7 21" id="KW-0770">Synapse</keyword>
    <keyword evidence="4 21" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 21" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="1 21" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="544"/>
            <end position="563"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="620"/>
            <end position="642"/>
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        <location>
            <begin position="801"/>
            <end position="825"/>
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            <begin position="423"/>
            <end position="488"/>
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    <feature type="binding site" evidence="18">
        <location>
            <position position="497"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="504"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="671"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="719"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="site" description="Crucial to convey clamshell closure to channel opening" evidence="19">
        <location>
            <position position="649"/>
        </location>
    </feature>
    <feature type="site" description="Interaction with the cone snail toxin Con-ikot-ikot" evidence="19">
        <location>
            <position position="676"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="77"/>
            <end position="328"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="731"/>
            <end position="785"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023257"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023286"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00034104"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00036634"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-1"/>
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    </evidence>
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        <source>
            <dbReference type="PIRSR" id="PIRSR601508-2"/>
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    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-3"/>
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    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="RuleBase" id="RU367118"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="SMART" id="SM00079"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="SMART" id="SM00918"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034632.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="889" mass="100223" checksum="ED25AB6E550222CE" modified="2019-05-08" version="1">IKIESSYTSQSDFQCVVAVRGIFECVESGPMGAEELAFRFAVNTINRNRTLLPNTTLTYDTQKINLYDSFEASKKACDQLSLGVAAIFGPSHSSSANAVQSICNALGVPHIQTRWKHQVSDNKDSFYVSLYPDFSSLSRAILDLVQFFKWKTVTVVYDDSTGLIRLQELIKAPSRYNLRLKIRQLPADTKDAKPLLKEMKRGKEFHVIFDCSHEMAAGILKQALAMGMMTEYYHYIFTTLDLFALDVEPYRYSGVNMTGFRILNTENSQVASIIEKWSMERLQAPPKPDSGLLDGFMTTDAALMYDAVHVVSVAVQQFPQMTVSSLQCNRHKPWRFGTRFMSLIKEAHWEGLTGRITFNKTNGLRTDFDLDVISLKEEGLEKIGTWDPASGLNMTENQKGKPANITDSLSNRSLIVTTILEEPYVMFKKSDKPLYGNDRFEGYCIDLLRELATILGFTYEIRLVEDGKYGAQDDASGQWNGMVRELIDHKADLAVAPLAITYVREKVIDFSKPFMTLGISILYRKPNGTNPGVFSFLNPLSPDIWMYILLAYLGVSCVLFVIARFSPYEWYNPHPCNPDSDVVENNFTLLNSFWFGVGALMQQGSELMPKALSTRIVGGIWWFFTLIIISSYTANLAAFLTVERMESPIDSADDLAKQTKIEYGAVEDGATMTFFKKSKISTYDKMWAFMSSRRQSVLVKSNEEGIQRVLTSDYAFLMESTTIEFVTQRNCNLTQIGGLIDSKGYGVGTPMGSPYRDKITIAILQLQEEGKLHMMKEKWWRGNGCPEEESKEASALGVQNIGGIFIVLAAGLVLSVFVAVGEFLYKSKKNAQLEKRSFCSAMVEELRMSLKCQRRLKHKPQAPVIVKTEEVINMHTFNDRRLPGKETMA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J2K1</accession>
    <name>A0A452J2K1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">Glutamate receptor</fullName>
        </recommendedName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="21">Receptor for glutamate that functions as a ligand-gated ion channel in the central nervous system and plays an important role in excitatory synaptic transmission. L-glutamate acts as an excitatory neurotransmitter at many synapses in the central nervous system.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>Ca(2+)(in) = Ca(2+)(out)</text>
            <dbReference type="Rhea" id="RHEA:29671"/>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="16 21">Postsynaptic cell membrane</location>
            <topology evidence="16 21">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="21">Belongs to the glutamate-gated ion channel (TC 1.A.10.1) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J2K1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039206.1">
        <property type="protein sequence ID" value="ENSGAGP00000034631.1"/>
        <property type="gene ID" value="ENSGAGG00000024584.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015276">
        <property type="term" value="F:ligand-gated monoatomic ion channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0038023">
        <property type="term" value="F:signaling receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd06382">
        <property type="entry name" value="PBP1_iGluR_Kainate"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.2300:FF:000010">
        <property type="entry name" value="Glutamate ionotropic receptor kainate type subunit 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.190.10:FF:000210">
        <property type="entry name" value="Glutamate receptor ionotropic, kainate 1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Glutamate receptor ionotropic, kainate 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.70:FF:000010">
        <property type="entry name" value="Putative glutamate receptor ionotropic kainate 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.70">
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="2"/>
    </dbReference>
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        <property type="entry name" value="Periplasmic binding protein-like II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001828">
        <property type="entry name" value="ANF_lig-bd_rcpt"/>
    </dbReference>
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        <property type="entry name" value="Glu/Gly-bd"/>
    </dbReference>
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        <property type="entry name" value="Iono_Glu_rcpt_met"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015683">
        <property type="entry name" value="Ionotropic_Glu_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001320">
        <property type="entry name" value="Iontro_rcpt_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028082">
        <property type="entry name" value="Peripla_BP_I"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18966">
        <property type="entry name" value="IONOTROPIC GLUTAMATE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01094">
        <property type="entry name" value="ANF_receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Lig_chan"/>
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    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="NMDARECEPTOR"/>
    </dbReference>
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        <property type="entry name" value="Lig_chan-Glu_bd"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Periplasmic binding protein-like I"/>
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    </dbReference>
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        <property type="entry name" value="Periplasmic binding protein-like II"/>
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    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 21" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="10 20" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="15 21" id="KW-0407">Ion channel</keyword>
    <keyword evidence="8 21" id="KW-0406">Ion transport</keyword>
    <keyword evidence="14 21" id="KW-1071">Ligand-gated ion channel</keyword>
    <keyword evidence="9 21" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="13 21" id="KW-0628">Postsynaptic cell membrane</keyword>
    <keyword evidence="11 21" id="KW-0675">Receptor</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <keyword evidence="7 21" id="KW-0770">Synapse</keyword>
    <keyword evidence="4 21" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 21" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="1 21" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="520"/>
            <end position="539"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="596"/>
            <end position="618"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="777"/>
            <end position="801"/>
        </location>
    </feature>
    <feature type="domain" description="Ionotropic glutamate receptor C-terminal" evidence="22">
        <location>
            <begin position="389"/>
            <end position="758"/>
        </location>
    </feature>
    <feature type="domain" description="Ionotropic glutamate receptor L-glutamate and glycine-binding" evidence="23">
        <location>
            <begin position="399"/>
            <end position="464"/>
        </location>
    </feature>
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        <location>
            <position position="473"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
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        <location>
            <position position="480"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="647"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
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        <location>
            <position position="695"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="site" description="Crucial to convey clamshell closure to channel opening" evidence="19">
        <location>
            <position position="625"/>
        </location>
    </feature>
    <feature type="site" description="Interaction with the cone snail toxin Con-ikot-ikot" evidence="19">
        <location>
            <position position="652"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="53"/>
            <end position="304"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="707"/>
            <end position="761"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023257"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00034104"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00036634"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="RuleBase" id="RU367118"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="SMART" id="SM00079"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="SMART" id="SM00918"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034631.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="865" mass="97710" checksum="183A7A51EC6A3987" modified="2019-05-08" version="1">MLSVAKVFFIELAFRFAVNTINRNRTLLPNTTLTYDTQKINLYDSFEASKKACDQLSLGVAAIFGPSHSSSANAVQSICNALGVPHIQTRWKHQVSDNKDSFYVSLYPDFSSLSRAILDLVQFFKWKTVTVVYDDSTGLIRLQELIKAPSRYNLRLKIRQLPADTKDAKPLLKEMKRGKEFHVIFDCSHEMAAGILKQALAMGMMTEYYHYIFTTLDLFALDVEPYRYSGVNMTGFRILNTENSQVASIIEKWSMERLQAPPKPDSGLLDGFMTTDAALMYDAVHVVSVAVQQFPQMTVSSLQCNRHKPWRFGTRFMSLIKEAHWEGLTGRITFNKTNGLRTDFDLDVISLKEEGLEKIGTWDPASGLNMTENQKGKPANITDSLSNRSLIVTTILEEPYVMFKKSDKPLYGNDRFEGYCIDLLRELATILGFTYEIRLVEDGKYGAQDDASGQWNGMVRELIDHKADLAVAPLAITYVREKVIDFSKPFMTLGISILYRKPNGTNPGVFSFLNPLSPDIWMYILLAYLGVSCVLFVIARFSPYEWYNPHPCNPDSDVVENNFTLLNSFWFGVGALMQQGSELMPKALSTRIVGGIWWFFTLIIISSYTANLAAFLTVERMESPIDSADDLAKQTKIEYGAVEDGATMTFFKKSKISTYDKMWAFMSSRRQSVLVKSNEEGIQRVLTSDYAFLMESTTIEFVTQRNCNLTQIGGLIDSKGYGVGTPMGSPYRDKITIAILQLQEEGKLHMMKEKWWRGNGCPEEESKEASALGVQNIGGIFIVLAAGLVLSVFVAVGEFLYKSKKNAQLEKRSFCSAMVEELRMSLKCQRRLKHKPQAPVIVKTEEVINMHTFNDRRLPGKETMA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJP3</accession>
    <name>A0A452GJP3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">TLC domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>sphinganine + octadecanoyl-CoA = N-(octadecanoyl)-sphinganine + CoA + H(+)</text>
            <dbReference type="Rhea" id="RHEA:36547"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57394"/>
            <dbReference type="ChEBI" id="CHEBI:57817"/>
            <dbReference type="ChEBI" id="CHEBI:67033"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="11">
            <dbReference type="Rhea" id="RHEA:36548"/>
        </physiologicalReaction>
    </comment>
    <comment type="pathway">
        <text evidence="2">Lipid metabolism; sphingolipid metabolism.</text>
    </comment>
    <comment type="pathway">
        <text evidence="3">Sphingolipid metabolism.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="12 14">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJP3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002154.1">
        <property type="protein sequence ID" value="ENSGAGP00000001895.1"/>
        <property type="gene ID" value="ENSGAGG00000001512.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00222"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050291">
        <property type="term" value="F:sphingosine N-acyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046513">
        <property type="term" value="P:ceramide biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00086">
        <property type="entry name" value="homeodomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.60:FF:000020">
        <property type="entry name" value="Ceramide synthase 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.60">
        <property type="entry name" value="Homeodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001356">
        <property type="entry name" value="HD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009057">
        <property type="entry name" value="Homeodomain-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016439">
        <property type="entry name" value="Lag1/Lac1-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006634">
        <property type="entry name" value="TLC-dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12560:SF6">
        <property type="entry name" value="CERAMIDE SYNTHASE 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12560">
        <property type="entry name" value="LONGEVITY ASSURANCE FACTOR 1 LAG1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00046">
        <property type="entry name" value="Homeodomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03798">
        <property type="entry name" value="TRAM_LAG1_CLN8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00724">
        <property type="entry name" value="TLC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46689">
        <property type="entry name" value="Homeodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50071">
        <property type="entry name" value="HOMEOBOX_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50922">
        <property type="entry name" value="TLC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="12 14" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="7" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="12 14" id="KW-0371">Homeobox</keyword>
    <keyword evidence="4" id="KW-0444">Lipid biosynthesis</keyword>
    <keyword evidence="9" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="10 13 15" id="KW-0472">Membrane</keyword>
    <keyword evidence="12 14" id="KW-0539">Nucleus</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0808">Transferase</keyword>
    <keyword evidence="6 13 15" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 15" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="98"/>
            <end position="116"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="152"/>
            <end position="172"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="179"/>
            <end position="197"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="217"/>
            <end position="245"/>
        </location>
    </feature>
    <feature type="domain" description="TLC" evidence="17">
        <location>
            <begin position="1"/>
            <end position="249"/>
        </location>
    </feature>
    <feature type="domain" description="Homeobox" evidence="16">
        <location>
            <begin position="13"/>
            <end position="74"/>
        </location>
    </feature>
    <feature type="DNA-binding region" description="Homeobox" evidence="12">
        <location>
            <begin position="15"/>
            <end position="75"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004477"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004760"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022516"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00049036"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00108"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00205"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="RuleBase" id="RU000682"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50071"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50922"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001895.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="20">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="252" mass="29822" checksum="7EAFFD35A475CE1D" modified="2019-05-08" version="1">LRLKFIALPLGRKLGVRDKVRLNAPPNPVLETFYATHSKKPKEGELICLSKQCDLPARKVETWFRYRRNQDKPSLTTKFCSVSLFVLLLQPLQRSVYWYYMMEFSFALLLTFTMAFDVRRKDFKEQMVHHAATIILISYSYCANYLRIGSLVMLLHVSSTFLLELTKLLHYLNWRRASHLLFLIFSSIFLVTRLIVFPCRVLYTSFYGSMEFYQPYFGYYLMNALLMVLQLLHVFWASLIIHMLYKFVNGTV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJQ0</accession>
    <name>A0A452GJQ0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Signal peptide peptidase-like 2B</fullName>
        </recommendedName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="4">Cell membrane</location>
            <topology evidence="4">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Endosome membrane</location>
            <topology evidence="2">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="5">Golgi apparatus membrane</location>
            <topology evidence="5">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Lysosome membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Membrane</location>
            <topology evidence="3">Multi-pass membrane protein</topology>
            <orientation evidence="3">Lumenal side</orientation>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the peptidase A22B family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJQ0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002178.1">
        <property type="protein sequence ID" value="ENSGAGP00000001918.1"/>
        <property type="gene ID" value="ENSGAGG00000001535.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098554">
        <property type="term" value="C:cytoplasmic side of endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010008">
        <property type="term" value="C:endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000139">
        <property type="term" value="C:Golgi membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030660">
        <property type="term" value="C:Golgi-associated vesicle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098553">
        <property type="term" value="C:lumenal side of endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005765">
        <property type="term" value="C:lysosomal membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042500">
        <property type="term" value="F:aspartic endopeptidase activity, intramembrane cleaving"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033619">
        <property type="term" value="P:membrane protein proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.50.30.30:FF:000006">
        <property type="entry name" value="Putative signal peptide peptidase-like 2B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.50.30.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003137">
        <property type="entry name" value="PA_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007369">
        <property type="entry name" value="Peptidase_A22B_SPP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006639">
        <property type="entry name" value="Preselin/SPP"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12174">
        <property type="entry name" value="SIGNAL PEPTIDE PEPTIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12174:SF39">
        <property type="entry name" value="SIGNAL PEPTIDE PEPTIDASE-LIKE 2B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02225">
        <property type="entry name" value="PA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04258">
        <property type="entry name" value="Peptidase_A22B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00730">
        <property type="entry name" value="PSN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11" id="KW-0967">Endosome</keyword>
    <keyword evidence="16" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="14" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="12" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="17" id="KW-0458">Lysosome</keyword>
    <keyword evidence="15 20" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-0645">Protease</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0732">Signal</keyword>
    <keyword evidence="9 20" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="13 20" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="117"/>
            <end position="138"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="167"/>
            <end position="188"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="194"/>
            <end position="215"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="241"/>
            <end position="258"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="264"/>
            <end position="281"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="293"/>
            <end position="313"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="358"/>
            <end position="376"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="388"/>
            <end position="411"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="417"/>
            <end position="435"/>
        </location>
    </feature>
    <feature type="domain" description="PA" evidence="21">
        <location>
            <begin position="20"/>
            <end position="94"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="461"/>
            <end position="553"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="473"/>
            <end position="488"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="506"/>
            <end position="516"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="19">
        <location>
            <begin position="520"/>
            <end position="535"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="543"/>
            <end position="553"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004155"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004337"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004366"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004653"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00006859"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023228"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00069829"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="Pfam" id="PF02225"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001918.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="553" mass="61912" checksum="A87370C119CE8B25" modified="2019-05-08" version="1">MFLSLQSLLQLQDQTASVLCSPSDVPDGGFHNRIPMVMRGNCTFYEKVRLAQINGARGLLIVSREKLAPPKGNWSQYEEIDIPVALLSYTDMLDIGKSFGRSVKVAMYAPNEPVLDYNMVIIFVMAVGTVAIGGYWAGSRDVKKRYMKHKRDDGAEKHDDETVDVTPIMICVFVVMCCSMLVLLYFFYDHLVYVIIGIFCLAASIGLYSCLSPFVRRFPLGKCRIPDNNLPYFHKRPQVRMLLLAVFCISISVVWGVFRNEDQWAWVLQDALGIAFCLYMLKTIRLPTFKGCTLLLMVLFVYDVFFVFITPFLTKTGQSIMVEVAAGPTDSATHEKLPMVLKVPRLNFSPLALCDRPFSLLGFGDILVPGLLVAYCHRFDIQVQSSRVYFVACTIAYGIGLLVTFVALALMQMGQPALLYLVPCTLITSFAVALWRKELAMFWTGSGFAKDLPHPPLVLAPVSCPNPRKDTNAPASQQEAEQQMTNPTLHADDRNNPSQCAEELPNLNTKELTNPSIRAEQPQISIPQIEQQTSQNKNDPESKSLTVEKNQLD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJQ2</accession>
    <name>A0A452GJQ2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Sushi domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="caution">
        <text evidence="6">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJQ2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002079.1">
        <property type="protein sequence ID" value="ENSGAGP00000001827.1"/>
        <property type="gene ID" value="ENSGAGG00000001465.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="CDD" id="cd00033">
        <property type="entry name" value="CCP"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.70.10:FF:000055">
        <property type="entry name" value="Complement decay-accelerating factor, GPI-anchored"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.70.10:FF:000014">
        <property type="entry name" value="Membrane cofactor protein"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.70.10">
        <property type="entry name" value="Complement Module, domain 1"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051277">
        <property type="entry name" value="SEZ6_CSMD_C4BPB_Regulators"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035976">
        <property type="entry name" value="Sushi/SCR/CCP_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000436">
        <property type="entry name" value="Sushi_SCR_CCP_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45656:SF3">
        <property type="entry name" value="CUB AND SUSHI DOMAIN-CONTAINING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45656">
        <property type="entry name" value="PROTEIN CBR-CLEC-78"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00084">
        <property type="entry name" value="Sushi"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SMART" id="SM00032">
        <property type="entry name" value="CCP"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57535">
        <property type="entry name" value="Complement control module/SCR domain"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50923">
        <property type="entry name" value="SUSHI"/>
        <property type="match status" value="6"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4 6" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0732">Signal</keyword>
    <keyword evidence="1 6" id="KW-0768">Sushi</keyword>
    <keyword evidence="7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="402"/>
            <end position="422"/>
        </location>
    </feature>
    <feature type="domain" description="Sushi" evidence="8">
        <location>
            <begin position="9"/>
            <end position="72"/>
        </location>
    </feature>
    <feature type="domain" description="Sushi" evidence="8">
        <location>
            <begin position="73"/>
            <end position="132"/>
        </location>
    </feature>
    <feature type="domain" description="Sushi" evidence="8">
        <location>
            <begin position="133"/>
            <end position="194"/>
        </location>
    </feature>
    <feature type="domain" description="Sushi" evidence="8">
        <location>
            <begin position="195"/>
            <end position="254"/>
        </location>
    </feature>
    <feature type="domain" description="Sushi" evidence="8">
        <location>
            <begin position="255"/>
            <end position="317"/>
        </location>
    </feature>
    <feature type="domain" description="Sushi" evidence="8">
        <location>
            <begin position="318"/>
            <end position="377"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="6">
        <location>
            <begin position="11"/>
            <end position="54"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="6">
        <location>
            <begin position="103"/>
            <end position="130"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="6">
        <location>
            <begin position="348"/>
            <end position="375"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022659"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00302"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50923"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001827.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="424" mass="45312" checksum="A9F4EB01BACB56B1" modified="2019-05-08" version="1">MLTFFASAIPCLPPPDITHGSHTGQGEEEFSFGSAVIYKCDQGFSLIGNASIHCTTKDNNGEWSGPAPECKVVRCPEPEVKNGKKQSGFGLDYSYGNVVIFECDSGYTLTGSSSVKCEANNSWVPSLPTCLRTCNSPPWLNFAELPGAVNDPYLVGTKLKYSCRPGYILGSGKSPFVTCLANSTWSVDPEFCVGRPCKPLELENGRVDLTDLRFGATATFSCNEGYRLIGTTSAKCVVAGNGVDWDKELPLCQAIPCLPPPDITHGSHTGQGEEFFFGSAVIYKCDQGFSLIGNASIHCTTKDNNGEWSGPAPECKVVKCPEPEVKNGKKQSGFGPNYSYGNIVIFECDSGYTLNSSSSVKCEANNSWVPSLPTCLRVTYTTTTTTSSPGGTTDTSGSNTGIMIAIGKMPVILAVLITNLFIRQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJQ3</accession>
    <name>A0A452GJQ3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">RING-type E3 ubiquitin transferase</fullName>
            <ecNumber evidence="5">2.3.2.27</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>S-ubiquitinyl-[E2 ubiquitin-conjugating enzyme]-L-cysteine + [acceptor protein]-L-lysine = [E2 ubiquitin-conjugating enzyme]-L-cysteine + N(6)-ubiquitinyl-[acceptor protein]-L-lysine.</text>
            <dbReference type="EC" id="2.3.2.27"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="3">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the TRIM/RBCC family.</text>
    </comment>
    <dbReference type="EC" id="2.3.2.27" evidence="5"/>
    <dbReference type="AlphaFoldDB" id="A0A452GJQ3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002188.1">
        <property type="protein sequence ID" value="ENSGAGP00000001925.1"/>
        <property type="gene ID" value="ENSGAGG00000001264.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061630">
        <property type="term" value="F:ubiquitin protein ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd12888">
        <property type="entry name" value="SPRY_PRY_TRIM7_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.920:FF:000004">
        <property type="entry name" value="Butyrophilin subfamily 1 member A1"/>
        <property type="match status" value="1"/>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="B30.2/SPRY"/>
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        <property type="entry name" value="B30.2/SPRY_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003879">
        <property type="entry name" value="Butyrophylin_SPRY"/>
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    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
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        <property type="entry name" value="PRY"/>
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    <dbReference type="InterPro" id="IPR003877">
        <property type="entry name" value="SPRY_dom"/>
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    <dbReference type="InterPro" id="IPR050143">
        <property type="entry name" value="TRIM/RBCC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000315">
        <property type="entry name" value="Znf_B-box"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020457">
        <property type="entry name" value="Znf_B-box_chordata"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001510">
        <property type="entry name" value="Znf_PARP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001841">
        <property type="entry name" value="Znf_RING"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017907">
        <property type="entry name" value="Znf_RING_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24103">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE TRIM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13765">
        <property type="entry name" value="PRY"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00643">
        <property type="entry name" value="zf-B_box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15227">
        <property type="entry name" value="zf-C3HC4_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01406">
        <property type="entry name" value="BBOXZNFINGER"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01407">
        <property type="entry name" value="BUTYPHLNCDUF"/>
    </dbReference>
    <dbReference type="SMART" id="SM00336">
        <property type="entry name" value="BBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="RING"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="B-box zinc-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57850">
        <property type="entry name" value="RING/U-box"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="B302_SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="ZF_BBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50064">
        <property type="entry name" value="ZF_PARP_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00518">
        <property type="entry name" value="ZF_RING_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50089">
        <property type="entry name" value="ZF_RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12 14" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0808">Transferase</keyword>
    <keyword evidence="10" id="KW-0833">Ubl conjugation pathway</keyword>
    <keyword evidence="11" id="KW-0862">Zinc</keyword>
    <keyword evidence="9 13" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="RING-type" evidence="16">
        <location>
            <begin position="18"/>
            <end position="61"/>
        </location>
    </feature>
    <feature type="domain" description="PARP-type" evidence="15">
        <location>
            <begin position="45"/>
            <end position="70"/>
        </location>
    </feature>
    <feature type="domain" description="B box-type" evidence="17">
        <location>
            <begin position="83"/>
            <end position="124"/>
        </location>
    </feature>
    <feature type="domain" description="B30.2/SPRY" evidence="18">
        <location>
            <begin position="282"/>
            <end position="471"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="14">
        <location>
            <begin position="182"/>
            <end position="209"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00008518"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00024"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50064"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50119"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50188"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001925.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="471" mass="54172" checksum="4F0D163DB0C2BEFD" modified="2019-05-08" version="1">GRSTGPRSGDLLLKEATCPICLEYLTDPVSVDCGHNFCRGCVTQYCVEWAGRGSLTCPICRSQILKGNLRPNWQLGNLVEKMKHVAVCTRHKKTLDLFCEEDEEAVCVVCERASQHRSHTVLLIEEAAQKYKEKIQAHLKTLKEEREKQLGLKATGEGKRQEYLKQTQTERQKIVSEFQQLRQFLEEQERLLLAQLEKLDKEILRIQKENVTKLSGQISHLSELIHELEGKCQKPASEFLQVRLSLKHSRSMCRDRRAPESCPLESSSQRSQCNCLCALCGKGKFQQPVEISLELEEKLMHMTLDPDTAHPQLILSEDRKTVRYGRLWNKLPNTTKRFDYERCVLGSEGFTMGRHCWEVEVGERGGWAVGVARESLGRKGEISFKPKGGIWAVEYWEGQYWALTDPLTPLPLSWVPRRIWIYLDYEQGQVTFFNADNESLIFTFPPASFAGERIRPLFVVWETESQLRLCP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJQ6</accession>
    <name>A0A452GJQ6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Tight junction protein 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cell junction</location>
            <location evidence="3">Tight junction</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Peripheral membrane protein</topology>
            <orientation evidence="2">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the MAGUK family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJQ6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002072.1">
        <property type="protein sequence ID" value="ENSGAGP00000001821.1"/>
        <property type="gene ID" value="ENSGAGG00000001448.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005923">
        <property type="term" value="C:bicellular tight junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050839">
        <property type="term" value="F:cell adhesion molecule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098609">
        <property type="term" value="P:cell-cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045216">
        <property type="term" value="P:cell-cell junction organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090557">
        <property type="term" value="P:establishment of endothelial intestinal barrier"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1905605">
        <property type="term" value="P:positive regulation of blood-brain barrier permeability"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0150105">
        <property type="term" value="P:protein localization to cell-cell junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd06727">
        <property type="entry name" value="PDZ1_ZO1-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd06728">
        <property type="entry name" value="PDZ2_ZO1-like_ds"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd06729">
        <property type="entry name" value="PDZ3_ZO1-like_domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12027">
        <property type="entry name" value="SH3_ZO-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000009">
        <property type="entry name" value="Putative tight junction protein ZO-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000013">
        <property type="entry name" value="Putative tight junction protein ZO-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000110">
        <property type="entry name" value="tight junction protein ZO-1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000075">
        <property type="entry name" value="Tight junction protein ZO-2 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.42.10">
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008145">
        <property type="entry name" value="GK/Ca_channel_bsu"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008144">
        <property type="entry name" value="Guanylate_kin-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001478">
        <property type="entry name" value="PDZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036034">
        <property type="entry name" value="PDZ_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005417">
        <property type="entry name" value="ZO"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005419">
        <property type="entry name" value="ZO-2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035598">
        <property type="entry name" value="ZO-2_SH3"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13865">
        <property type="entry name" value="TIGHT JUNCTION PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13865:SF26">
        <property type="entry name" value="TIGHT JUNCTION PROTEIN ZO-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00625">
        <property type="entry name" value="Guanylate_kin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00595">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07653">
        <property type="entry name" value="SH3_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01597">
        <property type="entry name" value="ZONOCCLUDNS"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01599">
        <property type="entry name" value="ZONOCCLUDNS2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00072">
        <property type="entry name" value="GuKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00228">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00326">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50156">
        <property type="entry name" value="PDZ domain-like"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50052">
        <property type="entry name" value="GUANYLATE_KINASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50106">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0965">Cell junction</keyword>
    <keyword evidence="7" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11" id="KW-0472">Membrane</keyword>
    <keyword evidence="12" id="KW-0539">Nucleus</keyword>
    <keyword evidence="8" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0677">Repeat</keyword>
    <keyword evidence="6 13" id="KW-0728">SH3 domain</keyword>
    <keyword evidence="5" id="KW-0796">Tight junction</keyword>
    <feature type="domain" description="PDZ" evidence="17">
        <location>
            <begin position="34"/>
            <end position="121"/>
        </location>
    </feature>
    <feature type="domain" description="PDZ" evidence="17">
        <location>
            <begin position="320"/>
            <end position="398"/>
        </location>
    </feature>
    <feature type="domain" description="PDZ" evidence="17">
        <location>
            <begin position="524"/>
            <end position="605"/>
        </location>
    </feature>
    <feature type="domain" description="SH3" evidence="15">
        <location>
            <begin position="619"/>
            <end position="684"/>
        </location>
    </feature>
    <feature type="domain" description="Guanylate kinase-like" evidence="16">
        <location>
            <begin position="789"/>
            <end position="891"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="148"/>
            <end position="317"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="419"/>
            <end position="473"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="493"/>
            <end position="518"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="937"/>
            <end position="1097"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1133"/>
            <end position="1206"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="171"/>
            <end position="180"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="181"/>
            <end position="225"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="233"/>
            <end position="263"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="272"/>
            <end position="295"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="431"/>
            <end position="460"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="958"/>
            <end position="970"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="1009"/>
            <end position="1029"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="1030"/>
            <end position="1045"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="1047"/>
            <end position="1059"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="14">
        <location>
            <begin position="1075"/>
            <end position="1086"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="14">
        <location>
            <begin position="1153"/>
            <end position="1164"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="1181"/>
            <end position="1191"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004413"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004435"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00007014"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022427"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50052"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50106"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001821.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="20">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1206" mass="135901" checksum="A209F7E3DD1CB718" modified="2019-05-08" version="1">MLARGGDPLLYPGRELSAWLGAPGMEELIWEQYTVTLQKDSKRGFGIAVSGGRDNPHFENGETSIVISDVLPGGPADGLLQENDRVVMVNGTPMENVPHSFAVQQLRKSGKVAAIVVKRPRKVQLTALRRSPSLDHDDRAFDILDDPAEFDGRSARSGYSDRSWHSGHGGRSQSWGNSPDRSYRRDQDRGRNYSRDHSRERSYSRDGSRGRSIDRERSLDRERRRDRSRGRSINRERSLDREHRRDRSRGRSIDRDDGYERGAGDYSPPRDYSYRHQPDPRYDKEARSHSRDRLNSHSPLLEPRRQHGYPGQQYQDRPISVLLTKSKPNEEYGLRLGSQIFIKEMTSTGLATKDGNLYEGDIILKINGTVTENMSLADARKLIEKSRGKLQLVVLRDRKQTLLNIPSLHDSDSEIEDISEIESNRSSSPQDDQKLHHSDFDSHSSNEKLKEKPNTKEDPPNRLSRMGAMPTPFKSTSDIAAAVIVTDTNKEPKYKDDLPVSQPKVAPRTFLRPSPEDEAIYGPNTKMVKFKKGDSVGLRLAGGNDVGIFVAGIQEGTSADQEGLQEGDQILKVNTQDFRGIVREDAVLYLLEIPKGETVTILAQSKYEVYRDIMACGRGDSFFIRSHFECEKESPQSLAFTRGEIFRVVDTLYDGKLGNWLAVRIGNELEKGIIPNKSRAEQMASVQNAQRDGSSDRADFWRMRGQRSGMKKNLKKSREDLTAIASVSTKFPAYERVLLREAGFKRPVVIFGPIADLAMEKLSNDLPDLYQTAKTEPKDAGSEKSTGVVRLNTVRQIIEQNKHALLDVTPKAVDLLNYTQWFPIVVFFNPDNKQGVKTMRQRLSPTSNKSSRKLYDQANKLKKTCSYLFTATINLNSANDSWYGSLKDTIQQQQVEAVWVSEGKMEGMDDDMEDRMSYLTAMGADYLSCDSRLISDLEDTDGEGGAYTDNELDEPSDEPSVSSISRSSEPVQLEESIRKPSPEPKAQMRRVGSREILRDPSPPPAFKPEPPKGKLQNKEDVYDFPKNYESKPNSSSTVSNETSAVLSKAAAPPVSVKPAFGRPILKTSQPAVPAAEEEEEEDDVDGGESAPKSVLGKVKIFEKMDHKARLQRIQELQEAQNARLEIAQKHPDIYAVPIKTQKPDQNWPQPMSSRPPEPQKPPTRPYLENRGSYGSDAEEEDYRRQLSDNSKRGYYGQPSRYRDTEL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJQ9</accession>
    <name>A0A452GJQ9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">E2 ubiquitin-conjugating enzyme</fullName>
            <ecNumber evidence="3">2.3.2.23</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>S-ubiquitinyl-[E1 ubiquitin-activating enzyme]-L-cysteine + [E2 ubiquitin-conjugating enzyme]-L-cysteine = [E1 ubiquitin-activating enzyme]-L-cysteine + S-ubiquitinyl-[E2 ubiquitin-conjugating enzyme]-L-cysteine.</text>
            <dbReference type="EC" id="2.3.2.23"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="2">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="similarity">
        <text evidence="9">Belongs to the ubiquitin-conjugating enzyme family.</text>
    </comment>
    <dbReference type="EC" id="2.3.2.23" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452GJQ9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002196.1">
        <property type="protein sequence ID" value="ENSGAGP00000001930.1"/>
        <property type="gene ID" value="ENSGAGG00000001519.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061631">
        <property type="term" value="F:ubiquitin conjugating enzyme activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.110.10:FF:000101">
        <property type="entry name" value="Ubiquitin-conjugating enzyme E2 D2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.110.10">
        <property type="entry name" value="Ubiquitin Conjugating Enzyme"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000608">
        <property type="entry name" value="UBC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023313">
        <property type="entry name" value="UBQ-conjugating_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016135">
        <property type="entry name" value="UBQ-conjugating_enzyme/RWD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24068">
        <property type="entry name" value="UBIQUITIN-CONJUGATING ENZYME E2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00179">
        <property type="entry name" value="UQ_con"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00212">
        <property type="entry name" value="UBCc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54495">
        <property type="entry name" value="UBC-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00183">
        <property type="entry name" value="UBC_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50127">
        <property type="entry name" value="UBC_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7 9" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="5 9" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0808">Transferase</keyword>
    <keyword evidence="6 9" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="UBC core" evidence="10">
        <location>
            <begin position="1"/>
            <end position="118"/>
        </location>
    </feature>
    <feature type="active site" description="Glycyl thioester intermediate" evidence="8">
        <location>
            <position position="56"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000485"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012486"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10133"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="RuleBase" id="RU362109"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50127"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001930.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="118" mass="13530" checksum="E71E2057CC06B30E" modified="2019-05-08" version="1">MLEPVTNDQYQDSIFYSGGVFFLDITFTPEYPFKPPKVTFRTRIYHCNINSQGVICLDILKDNWSPALTISKVLLSICSLLTDCNPADPLVGSIATQYMTNRAEHDRMARQWTKRYAT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJT3</accession>
    <name>A0A452GJT3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Focal adhesion kinase 1</fullName>
            <ecNumber evidence="4">2.7.10.2</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="19">Protein-tyrosine kinase 2</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="20">pp125FAK</fullName>
        </alternativeName>
    </protein>
    <organism evidence="26 27">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="27" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="26" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="26" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="21 28 29">
            <text>L-tyrosyl-[protein] + ATP = O-phospho-L-tyrosyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10596"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.10.2"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell junction</location>
            <location evidence="2">Focal adhesion</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cell membrane</location>
            <topology evidence="3">Peripheral membrane protein</topology>
            <orientation evidence="3">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Cilium basal body</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="2.7.10.2" evidence="4"/>
    <dbReference type="AlphaFoldDB" id="A0A452GJT3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002229.1">
        <property type="protein sequence ID" value="ENSGAGP00000001958.1"/>
        <property type="gene ID" value="ENSGAGG00000001572.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005925">
        <property type="term" value="C:focal adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004715">
        <property type="term" value="F:non-membrane spanning protein tyrosine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008284">
        <property type="term" value="P:positive regulation of cell population proliferation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007172">
        <property type="term" value="P:signal complex assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd14473">
        <property type="entry name" value="FERM_B-lobe"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd13190">
        <property type="entry name" value="FERM_C_FAK1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05056">
        <property type="entry name" value="PTKc_FAK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.330:FF:000001">
        <property type="entry name" value="focal adhesion kinase 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000058">
        <property type="entry name" value="focal adhesion kinase 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.20.90:FF:000021">
        <property type="entry name" value="focal adhesion kinase 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000047">
        <property type="entry name" value="focal adhesion kinase 1 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000039">
        <property type="entry name" value="Focal adhesion kinase, isoform D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.80.10:FF:000004">
        <property type="entry name" value="Protein-tyrosine kinase 2-beta isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.80.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.330">
        <property type="entry name" value="Nucleotidyltransferases domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.20.90">
        <property type="entry name" value="Phosphatidylinositol 3-kinase Catalytic Subunit, Chain A, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.540">
        <property type="entry name" value="Single helix bin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019749">
        <property type="entry name" value="Band_41_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041390">
        <property type="entry name" value="FADK_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049385">
        <property type="entry name" value="FAK1-like_FERM_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041784">
        <property type="entry name" value="FAK1/PYK2_FERM_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014352">
        <property type="entry name" value="FERM/acyl-CoA-bd_prot_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035963">
        <property type="entry name" value="FERM_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019748">
        <property type="entry name" value="FERM_central"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000299">
        <property type="entry name" value="FERM_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036137">
        <property type="entry name" value="Focal_adhe_kin_target_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005189">
        <property type="entry name" value="Focal_adhesion_kin_target_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001245">
        <property type="entry name" value="Ser-Thr/Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008266">
        <property type="entry name" value="Tyr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020635">
        <property type="entry name" value="Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029071">
        <property type="entry name" value="Ubiquitin-like_domsf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46221">
        <property type="entry name" value="FERM AND PDZ DOMAIN-CONTAINING PROTEIN FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46221:SF12">
        <property type="entry name" value="NON-SPECIFIC PROTEIN-TYROSINE KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21477">
        <property type="entry name" value="FERM_C_FAK1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00373">
        <property type="entry name" value="FERM_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18038">
        <property type="entry name" value="FERM_N_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03623">
        <property type="entry name" value="Focal_AT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07714">
        <property type="entry name" value="PK_Tyr_Ser-Thr"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00109">
        <property type="entry name" value="TYRKINASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00295">
        <property type="entry name" value="B41"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00219">
        <property type="entry name" value="TyrKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF68993">
        <property type="entry name" value="FAT domain of focal adhesion kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47031">
        <property type="entry name" value="Second domain of FERM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54236">
        <property type="entry name" value="Ubiquitin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50057">
        <property type="entry name" value="FERM_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00109">
        <property type="entry name" value="PROTEIN_KINASE_TYR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="12 22" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="13" id="KW-0965">Cell junction</keyword>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="17" id="KW-0966">Cell projection</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="16" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="5" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="11" id="KW-0418">Kinase</keyword>
    <keyword evidence="14" id="KW-0472">Membrane</keyword>
    <keyword evidence="10 22" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="8" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="27" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0808">Transferase</keyword>
    <keyword evidence="15" id="KW-0829">Tyrosine-protein kinase</keyword>
    <feature type="domain" description="FERM" evidence="25">
        <location>
            <begin position="35"/>
            <end position="355"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="24">
        <location>
            <begin position="456"/>
            <end position="714"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="23">
        <location>
            <begin position="1"/>
            <end position="27"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="23">
        <location>
            <begin position="720"/>
            <end position="774"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="23">
        <location>
            <begin position="871"/>
            <end position="954"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="23">
        <location>
            <begin position="10"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="23">
        <location>
            <begin position="720"/>
            <end position="731"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="23">
        <location>
            <begin position="871"/>
            <end position="883"/>
        </location>
    </feature>
    <feature type="binding site" evidence="22">
        <location>
            <position position="488"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004120"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004413"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00011903"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023137"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00039644"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00042078"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00043012"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00051245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS50057"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001958.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1086" mass="123052" checksum="E8E475CAB891B0F2" modified="2019-05-08" version="1">MAAAYLDPNLNHTPSSSAKTHLSTGVERSPGAMERVLKVFHYFESSSEPTTWASIIRHGDATDVRGIIQKIVDSHKVKNVACYGLRLSHLQSEEVHWLHLDMGVSNVRERFELAHPPEEWKYELRIRYLPKGFLNQFTEDKPTLNFFYQQVKNDYMLEIADQVDQEIALKLGCLEIRRSYWEMRGNALEKKSNYEVLEKDVGLKRFFPKSLLDSVKAKTLRKLIQQTFRQFAVLNREESIQKFFEILSPVYRFDKECFKCALGSSWIISVELAIGPEEGISYLTDKGSNPTHLADFNQVQTIQYSNSEDKDRKGMLQLKIAGAPEPLTVTAPSLTIAENMADLIDGYCRLVNGATQSFIIRPQKEGERALPSIPKLANNEKQGVRSHTVSVSGVSHCQHKVKKARRFLPFVFCSHEPPPKDEISGDETDDYAEIIDEEDTYTMPSTRDYEIQRERIELGRCIGEGQFGDVHQGVYMSPENPAMAVAIKTCKNCTSDSVREKFLQEALTMRQFDHPHIVKLIGVITENPVWIIMELCTLGELRSFLQVRKYSLDLASLILYAYQLSTALAYLESKRFVHRDIAARNVLVSSTDCVKLGDFGLSRYMEDSTYYKASKGKLPIKWMAPESINFRRFTSASDVWMFGVCMWEILMHGVKPFQGVKNNDVIGRIENGERLPMPPNCPPTLYSLMTKCWAYDPSRRPRFTELKAQLSTILEEEKLQQEERMRMESRRQVTVSWDSGGSDEAPPKPSRPGYPSPRSSEGFYPSPQHMVPPNHYQVSGYPGSHGIPAMAGSIYPGQASLLDQADSWNHRPQEISMWPPTMEDSGTLDMRGMGQVLPTHLMEERLIRQQQEMEEDQRWLEKEERFLKPDVRLSRGSIDREDGSLQGPAGNQHIYQPVGKPDHAAPPKKPPRPGAPSHLGSLASLNSPVDSYNEGVKLQPQEISPPPTANLDRSNDKVYENVTGLVKAVIEMSSKIQPAPPEEYVPMVKEVGLALRTLLATVDETIPVLPASTHREIEMAQKLLNSDLAELINKMKLAQQYVMTSLQQEYKKQMLTAAHALAVDAKNLLDVIDQARLKMIGQSRPH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJT8</accession>
    <name>A0A452GJT8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">KRR1 small subunit processome component</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="10">KRR-R motif-containing protein 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="8">Part of the small subunit (SSU) processome, first precursor of the small eukaryotic ribosomal subunit. During the assembly of the SSU processome in the nucleolus, many ribosome biogenesis factors, an RNA chaperone and ribosomal proteins associate with the nascent pre-rRNA and work in concert to generate RNA folding, modifications, rearrangements and cleavage as well as targeted degradation of pre-ribosomal RNA by the RNA exosome.</text>
    </comment>
    <comment type="function">
        <text evidence="10">Required for 40S ribosome biogenesis. Involved in nucleolar processing of pre-18S ribosomal RNA and ribosome assembly.</text>
    </comment>
    <comment type="subunit">
        <text evidence="10">Component of the ribosomal small subunit (SSU) processome.</text>
    </comment>
    <comment type="subunit">
        <text evidence="9">Part of the small subunit (SSU) processome, composed of more than 70 proteins and the RNA chaperone small nucleolar RNA (snoRNA) U3.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 10">Nucleus</location>
            <location evidence="1 10">Nucleolus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 10">Belongs to the KRR1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJT8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002106.1">
        <property type="protein sequence ID" value="ENSGAGP00000001852.1"/>
        <property type="gene ID" value="ENSGAGG00000001474.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032040">
        <property type="term" value="C:small-subunit processome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006364">
        <property type="term" value="P:rRNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd22393">
        <property type="entry name" value="KH-I_KRR1_rpt1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd22394">
        <property type="entry name" value="KH-I_KRR1_rpt2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1370.10:FF:000011">
        <property type="entry name" value="KRR1 small subunit processome component"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1370.10:FF:000014">
        <property type="entry name" value="KRR1 small subunit processome component"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1370.10">
        <property type="entry name" value="K Homology domain, type 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004087">
        <property type="entry name" value="KH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036612">
        <property type="entry name" value="KH_dom_type_1_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041174">
        <property type="entry name" value="KRR1-like_KH1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048550">
        <property type="entry name" value="KRR1-like_KH1_euk"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048548">
        <property type="entry name" value="KRR1-like_KH2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048549">
        <property type="entry name" value="KRR1-like_KH2_euk"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024166">
        <property type="entry name" value="rRNA_assembly_KRR1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12581">
        <property type="entry name" value="HIV-1 REV BINDING PROTEIN 2, 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12581:SF0">
        <property type="entry name" value="KRR1 SMALL SUBUNIT PROCESSOME COMPONENT HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17903">
        <property type="entry name" value="KH_KRR1_1st"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21800">
        <property type="entry name" value="KH_KRR1_2nd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF006515">
        <property type="entry name" value="KRR1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00322">
        <property type="entry name" value="KH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54791">
        <property type="entry name" value="Eukaryotic type KH-domain (KH-domain type I)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="11" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="13" id="KW-0472">Membrane</keyword>
    <keyword evidence="6 10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7 10" id="KW-0687">Ribonucleoprotein</keyword>
    <keyword evidence="3 10" id="KW-0690">Ribosome biogenesis</keyword>
    <keyword evidence="5 10" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="4 10" id="KW-0698">rRNA processing</keyword>
    <keyword evidence="13" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="13" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="13">
        <location>
            <begin position="336"/>
            <end position="357"/>
        </location>
    </feature>
    <feature type="domain" description="K Homology" evidence="14">
        <location>
            <begin position="136"/>
            <end position="208"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1"/>
            <end position="29"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="247"/>
            <end position="275"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="11">
        <location>
            <begin position="286"/>
            <end position="316"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="12">
        <location>
            <begin position="247"/>
            <end position="259"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004604"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009344"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022517"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022552"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023274"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00035000"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00035020"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PIRNR" id="PIRNR006515"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="SMART" id="SM00322"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001852.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="365" mass="42336" checksum="FE21A429D1ABB861" modified="2019-05-08" version="1">MAAPKKAESGQVAPASKGRSKKKAVNESELLTVPDGWKEPAFTREDNPKGLLEESSFATLFPKYREAYLKECWPLVQKALNEHHVNATLDLIEGSITVTTTKKTFDPYMIVRARDLIKLLARSVPFEQAVRVLQDDTACDIIKIGSLVSNRERFIKRRQRLIGPKGSTLKALELLTNCYIMVQGNTVSALGPFNGLKEVRKVVLDTMKNIHPIYNIKTLMIKRELSKDPELRTQSWERFLPQFKHKNLNKRKEPKKKSVKKEYTPFPPPQPENQIDKELASGEYFLKESQKKRKRMEELKAKQAESFRRRQEERNKAFIPPKEKLVVKPKKGNVCFLSLESFLLFYLKLLSFGWWLFSKPRYMEH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJT9</accession>
    <name>A0A452GJT9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">BHLH domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the MiT/TFE family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJT9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002235.1">
        <property type="protein sequence ID" value="ENSGAGP00000001964.1"/>
        <property type="gene ID" value="ENSGAGG00000001583.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046983">
        <property type="term" value="F:protein dimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030318">
        <property type="term" value="P:melanocyte differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd18926">
        <property type="entry name" value="bHLHzip_MITF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.280.10:FF:000003">
        <property type="entry name" value="microphthalmia-associated transcription factor isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.280.10">
        <property type="entry name" value="Helix-loop-helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011598">
        <property type="entry name" value="bHLH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036638">
        <property type="entry name" value="HLH_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021802">
        <property type="entry name" value="MiT/TFE_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031867">
        <property type="entry name" value="MiT/TFE_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45776:SF4">
        <property type="entry name" value="MICROPHTHALMIA-ASSOCIATED TRANSCRIPTION FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45776">
        <property type="entry name" value="MIP04163P"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11851">
        <property type="entry name" value="DUF3371"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00010">
        <property type="entry name" value="HLH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15951">
        <property type="entry name" value="MITF_TFEB_C_3_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00353">
        <property type="entry name" value="HLH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47459">
        <property type="entry name" value="HLH, helix-loop-helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50888">
        <property type="entry name" value="BHLH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0010">Activator</keyword>
    <keyword evidence="9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0804">Transcription</keyword>
    <keyword evidence="4" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="BHLH" evidence="11">
        <location>
            <begin position="285"/>
            <end position="338"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="9"/>
            <end position="28"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="40"/>
            <end position="62"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="462"/>
            <end position="493"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="331"/>
            <end position="372"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="41"/>
            <end position="51"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="52"/>
            <end position="62"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="466"/>
            <end position="477"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00008289"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50888"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001964.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="493" mass="55253" checksum="3D2E61026C6CB02E" modified="2019-05-08" version="1">MQPYFCCIDDSSEHPGASKPPLSSSTMTSRILLRQQLMREQMQEQERREQQQKQQAAQFMQQRVPLSQTPAINVSVPSNLPPPTQVPMEVLKVQTHLENPTKYHIQQAQRQQVKQYLSTTLANKHTNQALSLPCPNQPGDHVMPPGTGSSAPNSPMAMLTLNSNCEKEGFYKFEEQSRVESECPALNTHTRASCMQMDDVIDDIISLESSYNEEILGLMDPALQMANTLPVSGNLIDLYGNQGMPPSGLNISNSCPANLPNIKRELTACIFPTESEARALAKERQKKDNHNLIERRRRFNINDRIKELGTLIPKSNDPDMRWNKGTILKASVDYIRKLQREQQRTKELENRQKKLEHANRHLLLRIQELEMQARAHGLSLIPSTGLCSPDMVNRVIKQEPVIDNCNQDVLQRHSDLSCTTTLDLTDGTITFSDNLGTYSVPTKVGSKLEDILMDDTLSPVGVTDPLLSSVSPGASKTSSRRSSMSMEDTDHAC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJU2</accession>
    <name>A0A452GJU2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">C2H2-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJU2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002219.1">
        <property type="protein sequence ID" value="ENSGAGP00000001951.1"/>
        <property type="gene ID" value="ENSGAGG00000001569.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000785">
        <property type="term" value="C:chromatin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031519">
        <property type="term" value="C:PcG protein complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005667">
        <property type="term" value="C:transcription regulator complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002063">
        <property type="entry name" value="RB associated KRAB zinc finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002343">
        <property type="entry name" value="Zinc finger protein 33A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000382">
        <property type="entry name" value="zinc finger protein 35 isoform X4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000990">
        <property type="entry name" value="zinc finger protein 629 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14003">
        <property type="entry name" value="TRANSCRIPTIONAL REPRESSOR PROTEIN YY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14003:SF23">
        <property type="entry name" value="ZINC FINGER PROTEIN 143"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="9" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 11" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="110"/>
            <end position="137"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="138"/>
            <end position="165"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="166"/>
            <end position="193"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="194"/>
            <end position="231"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1"/>
            <end position="52"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001951.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="246" mass="27874" checksum="CCE1FD5D16563116" modified="2019-05-08" version="1">WGKGEPRTGTAGGAVCLDSHSVPAGDGNMNENEEEDPQQEGPKPVEAPGTLSDFFIREAFPSSDQDKACERQEICSPEKRQEKPIRCERGFRKRKDTIVHQKTLLGEKPYVCAKCGRNFYQNSGLIRHANFHTGEKPYKCPECGKCFSDSSNLIAHQRLHTGEKPYKCTDCDKCFSESSKLIIHRRTHTGEKPYMCPECGKSFSQRSHLVHQLMDSSQQLLMLKTNSKENPGIKNLVSEKHWHFPH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJU4</accession>
    <name>A0A452GJU4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">BHLH domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the MiT/TFE family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJU4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002240.1">
        <property type="protein sequence ID" value="ENSGAGP00000001969.1"/>
        <property type="gene ID" value="ENSGAGG00000001583.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046983">
        <property type="term" value="F:protein dimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030318">
        <property type="term" value="P:melanocyte differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd18926">
        <property type="entry name" value="bHLHzip_MITF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.280.10:FF:000003">
        <property type="entry name" value="microphthalmia-associated transcription factor isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.280.10">
        <property type="entry name" value="Helix-loop-helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011598">
        <property type="entry name" value="bHLH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036638">
        <property type="entry name" value="HLH_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021802">
        <property type="entry name" value="MiT/TFE_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031867">
        <property type="entry name" value="MiT/TFE_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45776:SF4">
        <property type="entry name" value="MICROPHTHALMIA-ASSOCIATED TRANSCRIPTION FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45776">
        <property type="entry name" value="MIP04163P"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11851">
        <property type="entry name" value="DUF3371"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00010">
        <property type="entry name" value="HLH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15951">
        <property type="entry name" value="MITF_TFEB_C_3_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00353">
        <property type="entry name" value="HLH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47459">
        <property type="entry name" value="HLH, helix-loop-helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50888">
        <property type="entry name" value="BHLH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0010">Activator</keyword>
    <keyword evidence="9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0804">Transcription</keyword>
    <keyword evidence="4" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="BHLH" evidence="11">
        <location>
            <begin position="266"/>
            <end position="319"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="50"/>
            <end position="72"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="443"/>
            <end position="474"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="312"/>
            <end position="353"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="50"/>
            <end position="60"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="61"/>
            <end position="72"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="447"/>
            <end position="458"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00008289"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50888"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001969.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="474" mass="53102" checksum="3A5FF7360237E142" modified="2019-05-08" version="1">MDALRVQMFMPCSFESFYLSSEHPGASKPPLSSSTMTSRILLRQQLMREQMQEQERREQQQKQQAAQFMQQRVPLSQTPAINVSVPSNLPPPTQVPMEVLKVQTHLENPTKYHIQQAQRQQVKQYLSTTLANKHTNQALSLPCPNQPGDHVMPPGTGSSAPNSPMAMLTLNSNCEKEMDDVIDDIISLESSYNEEILGLMDPALQMANTLPVSGNLIDLYGNQGMPPSGLNISNSCPANLPNIKRELTACIFPTESEARALAKERQKKDNHNLIERRRRFNINDRIKELGTLIPKSNDPDMRWNKGTILKASVDYIRKLQREQQRTKELENRQKKLEHANRHLLLRIQELEMQARAHGLSLIPSTGLCSPDMVNRVIKQEPVIDNCNQDVLQRHSDLSCTTTLDLTDGTITFSDNLGTYSVPTKVGSKLEDILMDDTLSPVGVTDPLLSSVSPGASKTSSRRSSMSMEDTDHAC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJU5</accession>
    <name>A0A452GJU5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">ubiquitinyl hydrolase 1</fullName>
            <ecNumber evidence="3">3.4.19.12</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>Thiol-dependent hydrolysis of ester, thioester, amide, peptide and isopeptide bonds formed by the C-terminal Gly of ubiquitin (a 76-residue protein attached to proteins as an intracellular targeting signal).</text>
            <dbReference type="EC" id="3.4.19.12"/>
        </reaction>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the peptidase C19 family.</text>
    </comment>
    <dbReference type="EC" id="3.4.19.12" evidence="3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002242.1">
        <property type="protein sequence ID" value="ENSGAGP00000001970.1"/>
        <property type="gene ID" value="ENSGAGG00000001383.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004843">
        <property type="term" value="F:cysteine-type deubiquitinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016579">
        <property type="term" value="P:protein deubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd02659">
        <property type="entry name" value="peptidase_C19C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.70.10">
        <property type="entry name" value="Cysteine proteinases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056850">
        <property type="entry name" value="ARM_UBP34_24_USP9X_Y"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021905">
        <property type="entry name" value="DUF3517"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038765">
        <property type="entry name" value="Papain-like_cys_pep_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050164">
        <property type="entry name" value="Peptidase_C19"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001394">
        <property type="entry name" value="Peptidase_C19_UCH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055176">
        <property type="entry name" value="UBP24/USP9X/USP9Y_UBL"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028889">
        <property type="entry name" value="USP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018200">
        <property type="entry name" value="USP_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24006">
        <property type="entry name" value="UBIQUITIN CARBOXYL-TERMINAL HYDROLASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24006:SF925">
        <property type="entry name" value="UBIQUITINYL HYDROLASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25010">
        <property type="entry name" value="ARM_UBP24_USP9X-Y"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12030">
        <property type="entry name" value="DUF3517"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00443">
        <property type="entry name" value="UCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22900">
        <property type="entry name" value="UCH_UBL1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54001">
        <property type="entry name" value="Cysteine proteinases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00972">
        <property type="entry name" value="USP_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00973">
        <property type="entry name" value="USP_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50235">
        <property type="entry name" value="USP_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="5" id="KW-0645">Protease</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0788">Thiol protease</keyword>
    <keyword evidence="6" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="USP" evidence="10">
        <location>
            <begin position="1381"/>
            <end position="1779"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="790"/>
            <end position="823"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="1419"/>
            <end position="1455"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="2298"/>
            <end position="2376"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="793"/>
            <end position="813"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="1425"/>
            <end position="1455"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="9">
        <location>
            <begin position="2298"/>
            <end position="2307"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="2325"/>
            <end position="2336"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="2350"/>
            <end position="2360"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000707"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009085"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012759"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022807"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50235"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001970.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2376" mass="270609" checksum="1CAC928CC7DE7100" modified="2019-05-08" version="1">MYLEGEVSVLEGAMHISLSHGVCVCLPCCLPSLHSCCLIECAGWLVDLINKFGTLNGFQILHDRFMSGSALNVQIIAALIKPFGQCYEFLTLHTVKKYFLPIIEMVPQFLENLTDDELKKEAKNEAKNDALSMIIKSLKNLASRVPGQEETVKNLEIFRLKMILRLLQISSFNGKMNALNEVNKVISSVSYYTHRHGNPEEEEWLTAERMAEWIQQNNILSIVLRDSLHQPQYVEKLEKILRFVIKEKALTLQDLDNIWAAQAGKHEAIVKNVHDLLAKLAWDFSPEQLDHLFDCFKASWTNASKKQREKLLELIRRLAEDDKDGVMAHKVLNLLWNLAHSDDVPVDIMDQALSAHIKILDYSCSQDRDTQKIQWIDRFIEELRTNDKWVIPALKQIREICSLFGEAPQNLSQTQRSPHVFYRHDLINQLQHNHALVTLVAENLSAYMESMRQYAKEHGEYDPQTVRPGSRYSHVQEVQERLNFLRFLLKDGQLWLCAPQAKQIWKCLAENAVYLCDREACFKWYSKLMGDEPDLDPDINKDFFENNVLQLDPSLLTENGMKCFERFFKAVNCREGKLVAKRRAYMMDDLELIGLDYLWRVVIQGNDDIASRAIDLLKEIYTNLGPRLQVNQVVIHEDFIQSCFDRLKASYDTLCVLDGDKDSINCARQEAIRMVRVLTVLREYINECDSDYHEERTILPMSRAFRGKHITLVVRFPNQGRQVEDLDIWSHTNDTIGSVRRCILNRIKANSAHTKVELFIGGELVDPADDRKLIGQLNLKDKTLITAKLTQISSNMPSSPDSSSDSSTGSPGNHGNHYSDGPNPEVESCLPGVIMSLHPRYISFLWQVADLGSSLNMPPLRDGARVLMKLMPPDNTTVEKLRAICLDHAKLGESSLSPSLDSLFFGPSASQVLYLTEVVYALLMPASAPLGEDASDFQYNFLKSGGLPLVLSMLTRNNFLPNADMETRRGAYLNALKIAKLLLTAIGFGHVRAVAEACQPVVEGTSPISPINQATHDQAVVLQTALQNIPNPTSECMLRNVAIRLAQQISDEASKYIPDICVIRAVQKIVWASGCGSVQLVFSSNEEISKIYEKTNAGNEPDMEDEQVCCEALEVMTLCFALIPTALDALSKEKAWQTFIIDLLLHCHSKTVRQMAQEQFFLMATRCCMGQRPLLFFITLLFTVLGSTARERAKHSGDYFTLLRHLLNYAYNSNINVPNAEVLLNNEIDWLKRIRDEVKRTGETGVEETILEGHLGVTKELLAFQTPEKKYHIGCEKGGANLIKELIDDFIFPASNVYLQYMRNGELPAEQAIPVCSSPATINAGFELLVALAVGCVRNLKQIVDTLTEMYYIGTAITTCEALTEWEYLPPVGPRPSKGFVGLKNAGATCYMNSVIQQLYMIPAIRNGILAIEGTGSDVDDDMSGDEKQDNESNVDPRDDVFGYPHQYEDKPTLSKTEDRKEYNIGVLRHLQVIFGHLAASRLQYYVPRGFWKQFRLWGEPVNLREQHDALEFFNSLVDSLDEALKALGHPAMLSKVLGGSFADQKICQGCPHRYECEESFTTLNVDIRNHQNLLDSLEQYVKGDLLEGANAYHCEKCNKKVDTVKRLLIKKLPPVLAIQLKRFDYDWERECAIKFNDYFEFPRELDMEPYTVAGVAKLEGDDVNPENQLIQNEQPENEHSGSTKYRLVGVLVHSGQASGGHYYSYIIQRNGGDGEKNRWYKFDDGDVTECKMDDDEEMKNQCFGGEYMGEVFDHMMKRMSYRRQKRWWNAYILFYERMDTMDKDNELIKYISELAMTTKPHQIKMPSAIERSVRKQNVQFMHNRMQYSLEYFQFIKKLLTCNSIFLNPPPGQDHLLPEAEEITMISIQLAARFLFTTGFHTKKIVRGPASDWYDALCILLRHSKNVRFWFAHNVLFNVSNRFSEYLLECPSAEVRGAFAKLIVFIAHFSLQDGPCPSPFASPGPSSQAYDNLSLSDHLLRAVLNLLRREVSEHGRHLQQYFNLFVMYANLGVAEKTQLLKLSVPATFMLVALDEGPGPPIKYQYAELGKLYTVVSQLIRCCNVSSRMQSSINGNPPLSNPYGDPNLSQPIMALQQNVADILFVRTSYVKKIIEDCSNSEETIKLLRFCCWENPQFSSTVLSELLWQVAYSYTYELRPYLDLLLQILLIEDSWQTHRIHNALKGIPDDRDGLFDTIQRSKNHYQKRAYQCIKCMVALFSNCPVAYQILQSNGDLKRKWTWAVEWLGDELERRPYTGNPQYTYNNWSPPVQSNETSNGYFLERSHSARMTLAKACELCPEEEPDDPDAPDEHESSPPEDAPLYPHSPSSQYQQNNHVHGQPYTGPAAHHMNNPQRTGQRAQENYEGSEEVSPPQTKD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJV3</accession>
    <name>A0A452GJV3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">RING-type E3 ubiquitin transferase</fullName>
            <ecNumber evidence="5">2.3.2.27</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>S-ubiquitinyl-[E2 ubiquitin-conjugating enzyme]-L-cysteine + [acceptor protein]-L-lysine = [E2 ubiquitin-conjugating enzyme]-L-cysteine + N(6)-ubiquitinyl-[acceptor protein]-L-lysine.</text>
            <dbReference type="EC" id="2.3.2.27"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="3">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the TRIM/RBCC family.</text>
    </comment>
    <dbReference type="EC" id="2.3.2.27" evidence="5"/>
    <dbReference type="AlphaFoldDB" id="A0A452GJV3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002253.1">
        <property type="protein sequence ID" value="ENSGAGP00000001980.1"/>
        <property type="gene ID" value="ENSGAGG00000001264.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061630">
        <property type="term" value="F:ubiquitin protein ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd16598">
        <property type="entry name" value="RING-HC_TRIM26_C-IV"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12888">
        <property type="entry name" value="SPRY_PRY_TRIM7_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.920:FF:000004">
        <property type="entry name" value="Butyrophilin subfamily 1 member A1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.920">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001870">
        <property type="entry name" value="B30.2/SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043136">
        <property type="entry name" value="B30.2/SPRY_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003879">
        <property type="entry name" value="Butyrophylin_SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006574">
        <property type="entry name" value="PRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003877">
        <property type="entry name" value="SPRY_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050143">
        <property type="entry name" value="TRIM/RBCC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000315">
        <property type="entry name" value="Znf_B-box"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020457">
        <property type="entry name" value="Znf_B-box_chordata"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001510">
        <property type="entry name" value="Znf_PARP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001841">
        <property type="entry name" value="Znf_RING"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017907">
        <property type="entry name" value="Znf_RING_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24103">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE TRIM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13765">
        <property type="entry name" value="PRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00622">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00643">
        <property type="entry name" value="zf-B_box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15227">
        <property type="entry name" value="zf-C3HC4_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01406">
        <property type="entry name" value="BBOXZNFINGER"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01407">
        <property type="entry name" value="BUTYPHLNCDUF"/>
    </dbReference>
    <dbReference type="SMART" id="SM00336">
        <property type="entry name" value="BBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00589">
        <property type="entry name" value="PRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00184">
        <property type="entry name" value="RING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00449">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57845">
        <property type="entry name" value="B-box zinc-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57850">
        <property type="entry name" value="RING/U-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50188">
        <property type="entry name" value="B302_SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50119">
        <property type="entry name" value="ZF_BBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50064">
        <property type="entry name" value="ZF_PARP_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00518">
        <property type="entry name" value="ZF_RING_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50089">
        <property type="entry name" value="ZF_RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12 14" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0808">Transferase</keyword>
    <keyword evidence="10" id="KW-0833">Ubl conjugation pathway</keyword>
    <keyword evidence="11" id="KW-0862">Zinc</keyword>
    <keyword evidence="9 13" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="RING-type" evidence="16">
        <location>
            <begin position="16"/>
            <end position="59"/>
        </location>
    </feature>
    <feature type="domain" description="PARP-type" evidence="15">
        <location>
            <begin position="43"/>
            <end position="68"/>
        </location>
    </feature>
    <feature type="domain" description="B box-type" evidence="17">
        <location>
            <begin position="78"/>
            <end position="107"/>
        </location>
    </feature>
    <feature type="domain" description="B30.2/SPRY" evidence="18">
        <location>
            <begin position="245"/>
            <end position="434"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="14">
        <location>
            <begin position="165"/>
            <end position="192"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00008518"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00024"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50064"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50119"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50188"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001980.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="434" mass="50024" checksum="8F7C64AEE1F6FD75" modified="2019-05-08" version="1">TTGPVLCLDMREEATCPICLEYLTDPVSVDCGHNFCRGCVTQYCVEWAGRGSLTCPICRSQILKGNLRPNWQLGNLTLDLFCEEDEEAVCVVCERASQHRSHTVLLIEEAAQKYKEKIQAHLKTLKEEREKQLGLKATGEGKRQEYLKQTQTERQKIVSEFQQLRQFLEEQERLLLAQLEKLDKEILRIQKENVTKLSGQISHLSELIHELEGKCQKPASEFLQVRLSLKHSRSMCRDRRAPESCPLESTLRSVRNGPSLPRVHMTLDPDTAHPQLILSEDRKTVRYGRLWNKLPNTTKRFDYERCVLGSEGFTMGRHCWEVEVGERGGWAVGVARESLGRKGEISFKPKGGIWAVEYWEGQYWALTDPLTPLPLSWVPRRIWIYLDYEQGQVTFFNADNESLIFTFPPASFAGERIRPLFVVWETESQLRLCP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJV6</accession>
    <name>A0A452GJV6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">RING-type E3 ubiquitin transferase</fullName>
            <ecNumber evidence="5">2.3.2.27</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>S-ubiquitinyl-[E2 ubiquitin-conjugating enzyme]-L-cysteine + [acceptor protein]-L-lysine = [E2 ubiquitin-conjugating enzyme]-L-cysteine + N(6)-ubiquitinyl-[acceptor protein]-L-lysine.</text>
            <dbReference type="EC" id="2.3.2.27"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="3">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the TRIM/RBCC family.</text>
    </comment>
    <dbReference type="EC" id="2.3.2.27" evidence="5"/>
    <dbReference type="AlphaFoldDB" id="A0A452GJV6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002260.1">
        <property type="protein sequence ID" value="ENSGAGP00000001985.1"/>
        <property type="gene ID" value="ENSGAGG00000001264.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061630">
        <property type="term" value="F:ubiquitin protein ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd16598">
        <property type="entry name" value="RING-HC_TRIM26_C-IV"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12888">
        <property type="entry name" value="SPRY_PRY_TRIM7_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.920:FF:000004">
        <property type="entry name" value="Butyrophilin subfamily 1 member A1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.920">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001870">
        <property type="entry name" value="B30.2/SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043136">
        <property type="entry name" value="B30.2/SPRY_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003879">
        <property type="entry name" value="Butyrophylin_SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006574">
        <property type="entry name" value="PRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003877">
        <property type="entry name" value="SPRY_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050143">
        <property type="entry name" value="TRIM/RBCC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000315">
        <property type="entry name" value="Znf_B-box"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020457">
        <property type="entry name" value="Znf_B-box_chordata"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001510">
        <property type="entry name" value="Znf_PARP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001841">
        <property type="entry name" value="Znf_RING"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017907">
        <property type="entry name" value="Znf_RING_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24103">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE TRIM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13765">
        <property type="entry name" value="PRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00622">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00643">
        <property type="entry name" value="zf-B_box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15227">
        <property type="entry name" value="zf-C3HC4_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01406">
        <property type="entry name" value="BBOXZNFINGER"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01407">
        <property type="entry name" value="BUTYPHLNCDUF"/>
    </dbReference>
    <dbReference type="SMART" id="SM00336">
        <property type="entry name" value="BBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00589">
        <property type="entry name" value="PRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00184">
        <property type="entry name" value="RING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00449">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57845">
        <property type="entry name" value="B-box zinc-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57850">
        <property type="entry name" value="RING/U-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50188">
        <property type="entry name" value="B302_SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50119">
        <property type="entry name" value="ZF_BBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50064">
        <property type="entry name" value="ZF_PARP_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00518">
        <property type="entry name" value="ZF_RING_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50089">
        <property type="entry name" value="ZF_RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12 14" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0808">Transferase</keyword>
    <keyword evidence="10" id="KW-0833">Ubl conjugation pathway</keyword>
    <keyword evidence="11" id="KW-0862">Zinc</keyword>
    <keyword evidence="9 13" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="RING-type" evidence="16">
        <location>
            <begin position="16"/>
            <end position="59"/>
        </location>
    </feature>
    <feature type="domain" description="PARP-type" evidence="15">
        <location>
            <begin position="43"/>
            <end position="68"/>
        </location>
    </feature>
    <feature type="domain" description="B box-type" evidence="17">
        <location>
            <begin position="81"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="domain" description="B30.2/SPRY" evidence="18">
        <location>
            <begin position="277"/>
            <end position="467"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="14">
        <location>
            <begin position="180"/>
            <end position="207"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00008518"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00024"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50064"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50119"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50188"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001985.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="467" mass="53998" checksum="0064F324C214D67E" modified="2019-05-08" version="1">MASSLSGLDMREEATCPICLEYLTDPVSVDCGHNFCRGCVTQYCVEWAGRGSLTCPICRSQILKGNLRPNWQLGNLVEKMKHVAVCTRHKKTLDLFCEEDEEAVCVVCERASQHRSHTVLLIEEAAQKYKEKIQAHLKTLKEEREKQLGLKATGEGKRQEYLKQTQTERQKIVSEFQQLRQFLEEQERLLLAQLEKLDKEILRIQKENVTKLSGQISHLSELIHELEGKCQKPASEFLQVRLSLKHSRSMCRDRRAPESCPLESSSQRSQCNCLCALLEWELLFFELQTHKYLGLLHMTLDPDTAHPQLILSEDRKTVRYGRLWNKLPNTTKRFDYERCVLGSEGFTMGRHCWEVEVGERGGWAVGVARESLGRKGEISFKPKGGIWAVEYWEGQYWALTDPLTPLPLSWVPRRIWIYLDYEQGQVTFFNADNESLIFTFPPASFAGERIRPLFVVWETESQLRLCP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJV9</accession>
    <name>A0A452GJV9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">RING-type E3 ubiquitin transferase</fullName>
            <ecNumber evidence="5">2.3.2.27</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>S-ubiquitinyl-[E2 ubiquitin-conjugating enzyme]-L-cysteine + [acceptor protein]-L-lysine = [E2 ubiquitin-conjugating enzyme]-L-cysteine + N(6)-ubiquitinyl-[acceptor protein]-L-lysine.</text>
            <dbReference type="EC" id="2.3.2.27"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="3">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the TRIM/RBCC family.</text>
    </comment>
    <dbReference type="EC" id="2.3.2.27" evidence="5"/>
    <dbReference type="AlphaFoldDB" id="A0A452GJV9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002259.1">
        <property type="protein sequence ID" value="ENSGAGP00000001984.1"/>
        <property type="gene ID" value="ENSGAGG00000001264.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061630">
        <property type="term" value="F:ubiquitin protein ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd12888">
        <property type="entry name" value="SPRY_PRY_TRIM7_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.920:FF:000004">
        <property type="entry name" value="Butyrophilin subfamily 1 member A1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.920">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001870">
        <property type="entry name" value="B30.2/SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043136">
        <property type="entry name" value="B30.2/SPRY_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003879">
        <property type="entry name" value="Butyrophylin_SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006574">
        <property type="entry name" value="PRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003877">
        <property type="entry name" value="SPRY_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050143">
        <property type="entry name" value="TRIM/RBCC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000315">
        <property type="entry name" value="Znf_B-box"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020457">
        <property type="entry name" value="Znf_B-box_chordata"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001510">
        <property type="entry name" value="Znf_PARP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001841">
        <property type="entry name" value="Znf_RING"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017907">
        <property type="entry name" value="Znf_RING_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24103">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE TRIM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13765">
        <property type="entry name" value="PRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00622">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00643">
        <property type="entry name" value="zf-B_box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15227">
        <property type="entry name" value="zf-C3HC4_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01406">
        <property type="entry name" value="BBOXZNFINGER"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01407">
        <property type="entry name" value="BUTYPHLNCDUF"/>
    </dbReference>
    <dbReference type="SMART" id="SM00336">
        <property type="entry name" value="BBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00589">
        <property type="entry name" value="PRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00184">
        <property type="entry name" value="RING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00449">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57845">
        <property type="entry name" value="B-box zinc-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57850">
        <property type="entry name" value="RING/U-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50188">
        <property type="entry name" value="B302_SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50119">
        <property type="entry name" value="ZF_BBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50064">
        <property type="entry name" value="ZF_PARP_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00518">
        <property type="entry name" value="ZF_RING_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50089">
        <property type="entry name" value="ZF_RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12 14" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0808">Transferase</keyword>
    <keyword evidence="10" id="KW-0833">Ubl conjugation pathway</keyword>
    <keyword evidence="11" id="KW-0862">Zinc</keyword>
    <keyword evidence="9 13" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="RING-type" evidence="16">
        <location>
            <begin position="13"/>
            <end position="56"/>
        </location>
    </feature>
    <feature type="domain" description="PARP-type" evidence="15">
        <location>
            <begin position="40"/>
            <end position="65"/>
        </location>
    </feature>
    <feature type="domain" description="B box-type" evidence="17">
        <location>
            <begin position="78"/>
            <end position="119"/>
        </location>
    </feature>
    <feature type="domain" description="B30.2/SPRY" evidence="18">
        <location>
            <begin position="252"/>
            <end position="442"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="14">
        <location>
            <begin position="155"/>
            <end position="182"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00008518"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00024"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50064"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50119"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50188"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001984.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="442" mass="51189" checksum="6F98C8B1FE001EE6" modified="2019-05-08" version="1">MKNFSVKREEATCPICLEYLTDPVSVDCGHNFCRGCVTQYCVEWAGRGSLTCPICRSQILKGNLRPNWQLGNLVEKMKHVAVCTRHKKTLDLFCEEDEEAVCVVCERASQHRSHTVLLIEEAAQKYKVGNTWILVNSKQTQTERQKIVSEFQQLRQFLEEQERLLLAQLEKLDKEILRIQKENVTKLSGQISHLSELIHELEGKCQKPASEFLQVRLSLKHSRSMCRDRRAPESCPLESSSQRSQCNCLCALLEWELLFFELQTHKYLGLLHMTLDPDTAHPQLILSEDRKTVRYGRLWNKLPNTTKRFDYERCVLGSEGFTMGRHCWEVEVGERGGWAVGVARESLGRKGEISFKPKGGIWAVEYWEGQYWALTDPLTPLPLSWVPRRIWIYLDYEQGQVTFFNADNESLIFTFPPASFAGERIRPLFVVWETESQLRLCP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJW0</accession>
    <name>A0A452GJW0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Focal adhesion kinase 1</fullName>
            <ecNumber evidence="4">2.7.10.2</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="19">Protein-tyrosine kinase 2</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="20">pp125FAK</fullName>
        </alternativeName>
    </protein>
    <organism evidence="26 27">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="27" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="26" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="26" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="21 28 29">
            <text>L-tyrosyl-[protein] + ATP = O-phospho-L-tyrosyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10596"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.10.2"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell junction</location>
            <location evidence="2">Focal adhesion</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cell membrane</location>
            <topology evidence="3">Peripheral membrane protein</topology>
            <orientation evidence="3">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Cilium basal body</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="2.7.10.2" evidence="4"/>
    <dbReference type="AlphaFoldDB" id="A0A452GJW0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002239.1">
        <property type="protein sequence ID" value="ENSGAGP00000001968.1"/>
        <property type="gene ID" value="ENSGAGG00000001572.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005925">
        <property type="term" value="C:focal adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004715">
        <property type="term" value="F:non-membrane spanning protein tyrosine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008284">
        <property type="term" value="P:positive regulation of cell population proliferation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007172">
        <property type="term" value="P:signal complex assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd14473">
        <property type="entry name" value="FERM_B-lobe"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd13190">
        <property type="entry name" value="FERM_C_FAK1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05056">
        <property type="entry name" value="PTKc_FAK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.330:FF:000001">
        <property type="entry name" value="focal adhesion kinase 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000058">
        <property type="entry name" value="focal adhesion kinase 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.20.90:FF:000021">
        <property type="entry name" value="focal adhesion kinase 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000047">
        <property type="entry name" value="focal adhesion kinase 1 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000039">
        <property type="entry name" value="Focal adhesion kinase, isoform D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.80.10">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.330">
        <property type="entry name" value="Nucleotidyltransferases domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.20.90">
        <property type="entry name" value="Phosphatidylinositol 3-kinase Catalytic Subunit, Chain A, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.540">
        <property type="entry name" value="Single helix bin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019749">
        <property type="entry name" value="Band_41_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041390">
        <property type="entry name" value="FADK_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049385">
        <property type="entry name" value="FAK1-like_FERM_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041784">
        <property type="entry name" value="FAK1/PYK2_FERM_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014352">
        <property type="entry name" value="FERM/acyl-CoA-bd_prot_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035963">
        <property type="entry name" value="FERM_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019748">
        <property type="entry name" value="FERM_central"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000299">
        <property type="entry name" value="FERM_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036137">
        <property type="entry name" value="Focal_adhe_kin_target_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005189">
        <property type="entry name" value="Focal_adhesion_kin_target_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001245">
        <property type="entry name" value="Ser-Thr/Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008266">
        <property type="entry name" value="Tyr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020635">
        <property type="entry name" value="Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029071">
        <property type="entry name" value="Ubiquitin-like_domsf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46221">
        <property type="entry name" value="FERM AND PDZ DOMAIN-CONTAINING PROTEIN FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46221:SF12">
        <property type="entry name" value="NON-SPECIFIC PROTEIN-TYROSINE KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21477">
        <property type="entry name" value="FERM_C_FAK1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00373">
        <property type="entry name" value="FERM_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18038">
        <property type="entry name" value="FERM_N_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03623">
        <property type="entry name" value="Focal_AT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07714">
        <property type="entry name" value="PK_Tyr_Ser-Thr"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00109">
        <property type="entry name" value="TYRKINASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00295">
        <property type="entry name" value="B41"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00219">
        <property type="entry name" value="TyrKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF68993">
        <property type="entry name" value="FAT domain of focal adhesion kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47031">
        <property type="entry name" value="Second domain of FERM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54236">
        <property type="entry name" value="Ubiquitin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50057">
        <property type="entry name" value="FERM_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00109">
        <property type="entry name" value="PROTEIN_KINASE_TYR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="12 22" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="13" id="KW-0965">Cell junction</keyword>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="17" id="KW-0966">Cell projection</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="16" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="5" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="11" id="KW-0418">Kinase</keyword>
    <keyword evidence="14" id="KW-0472">Membrane</keyword>
    <keyword evidence="10 22" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="8" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="27" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0808">Transferase</keyword>
    <keyword evidence="15" id="KW-0829">Tyrosine-protein kinase</keyword>
    <feature type="domain" description="FERM" evidence="25">
        <location>
            <begin position="35"/>
            <end position="334"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="24">
        <location>
            <begin position="442"/>
            <end position="700"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="23">
        <location>
            <begin position="1"/>
            <end position="27"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="23">
        <location>
            <begin position="706"/>
            <end position="760"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="23">
        <location>
            <begin position="857"/>
            <end position="940"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="23">
        <location>
            <begin position="10"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="23">
        <location>
            <begin position="706"/>
            <end position="717"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="23">
        <location>
            <begin position="857"/>
            <end position="869"/>
        </location>
    </feature>
    <feature type="binding site" evidence="22">
        <location>
            <position position="474"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004120"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004413"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00011903"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023137"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00039644"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00042078"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00043012"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00051245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS50057"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001968.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1072" mass="121230" checksum="77A3E38F64583B86" modified="2019-05-08" version="1">MAAAYLDPNLNHTPSSSAKTHLSTGVERSPGAMERVLKVFHYFESSSEPTTWASIIRHGDATDVRGIIQKIVDSHKVKNVACYGLRLSHLQSEEVHWLHLDMGVSNVRERFELAHPPEEWKYELRIRYLPKGFLNQFTEDKPTLNFFYQQVKNDYMLEIADQVDQEIALKLGCLEIRKDVGLKRFFPKSLLDSVKAKTLRKLIQQTFRQFAVLNREESIQKFFEILSPVYRFDKECFKCALGSSWIISVELAIGPEEGISYLTDKGSNPTHLADFNQVQTIQYSNSEDKDRKGMLQLKIAGAPEPLTVTAPSLTIAENMADLIDGYCRLVNGATQSFIIRPQKEGERALPSIPKLANNEKQGVRSHTVSVSGVSHCQHKVKKARRFLPFVFCSHEPPPKDEISGDETDDYAEIIDEEDTYTMPSKSYGIDEARDYEIQRERIELGRCIGEGQFGDVHQGVYMSPENPAMAVAIKTCKNCTSDSVREKFLQEALTMRQFDHPHIVKLIGVITENPVWIIMELCTLGELRSFLQVRKYSLDLASLILYAYQLSTALAYLESKRFVHRDIAARNVLVSSTDCVKLGDFGLSRYMEDSTYYKASKGKLPIKWMAPESINFRRFTSASDVWMFGVCMWEILMHGVKPFQGVKNNDVIGRIENGERLPMPPNCPPTLYSLMTKCWAYDPSRRPRFTELKAQLSTILEEEKLQQEERMRMESRRQVTVSWDSGGSDEAPPKPSRPGYPSPRSSEGFYPSPQHMVPPNHYQVSGYPGSHGIPAMAGSIYPGQASLLDQADSWNHRPQEISMWPPTMEDSGTLDMRGMGQVLPTHLMEERLIRQQQEMEEDQRWLEKEERFLKPDVRLSRGSIDREDGSLQGPAGNQHIYQPVGKPDHAAPPKKPPRPGAPSHLGSLASLNSPVDSYNEGVKLQPQEISPPPTANLDRSNDKVYENVTGLVKAVIEMSSKIQPAPPEEYVPMVKEVGLALRTLLATVDETIPVLPASTHREIEMAQKLLNSDLAELINKMKLAQQYVMTSLQQEYKKQMLTAAHALAVDAKNLLDVIDQARLKMIGQSRPH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJW8</accession>
    <name>A0A452GJW8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="26">Phospholipid-transporting ATPase</fullName>
            <ecNumber evidence="26">7.6.2.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="30 31">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="31" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="30" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="30" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="21">Catalytic component of P4-ATPase flippase complex, which catalyzes the hydrolysis of ATP coupled to the transport of phosphatidylcholine (PC) from the outer to the inner leaflet of the plasma membrane. Initiates inward plasma membrane bending and recruitment of Bin/amphiphysin/Rvs (BAR) domain-containing proteins involved in membrane tubulation and cell trafficking. Facilitates ITGB1/beta1 integrin endocytosis, delaying cell adhesion and cell spreading on extracellular matrix. Has low flippase activity toward glucosylceramide (GlcCer).</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="18 26">
            <text>ATP + H2O + phospholipidSide 1 = ADP + phosphate + phospholipidSide 2.</text>
            <dbReference type="EC" id="7.6.2.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="20">
            <text>a 1,2-diacyl-sn-glycero-3-phosphocholine(out) + ATP + H2O = a 1,2-diacyl-sn-glycero-3-phosphocholine(in) + ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:38583"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:57643"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="20">
            <dbReference type="Rhea" id="RHEA:38584"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>a beta-D-glucosyl-(1&lt;-&gt;1')-N-acylsphing-4-enine(out) + ATP + H2O = a beta-D-glucosyl-(1&lt;-&gt;1')-N-acylsphing-4-enine(in) + ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:66036"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:22801"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="19">
            <dbReference type="Rhea" id="RHEA:66037"/>
        </physiologicalReaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 25">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subunit">
        <text evidence="22">Component of a P4-ATPase flippase complex which consists of a catalytic alpha subunit ATP10A and an accessory beta subunit TMEM30A.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cell membrane</location>
            <topology evidence="3">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Endoplasmic reticulum membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="26">Membrane</location>
            <topology evidence="26">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4 26">Belongs to the cation transport ATPase (P-type) (TC 3.A.3) family. Type IV subfamily.</text>
    </comment>
    <dbReference type="EC" id="7.6.2.1" evidence="26"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002272.1">
        <property type="protein sequence ID" value="ENSGAGP00000001997.1"/>
        <property type="gene ID" value="ENSGAGG00000000613.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:1990531">
        <property type="term" value="C:phospholipid-translocating ATPase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140351">
        <property type="term" value="F:glycosylceramide flippase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000287">
        <property type="term" value="F:magnesium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090554">
        <property type="term" value="F:phosphatidylcholine floppase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="RHEA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045332">
        <property type="term" value="P:phospholipid translocation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd02073">
        <property type="entry name" value="P-type_ATPase_APLT_Dnf-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.70.150.10:FF:000022">
        <property type="entry name" value="Phospholipid-transporting ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.1110.10:FF:000009">
        <property type="entry name" value="Phospholipid-transporting ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.1110.10:FF:000109">
        <property type="entry name" value="Phospholipid-transporting ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1000:FF:000110">
        <property type="entry name" value="Phospholipid-transporting ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1000:FF:000001">
        <property type="entry name" value="Phospholipid-transporting ATPase IC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.1110.10">
        <property type="entry name" value="Calcium-transporting ATPase, cytoplasmic domain N"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.70.150.10">
        <property type="entry name" value="Calcium-transporting ATPase, cytoplasmic transduction domain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1110.10">
        <property type="entry name" value="Calcium-transporting ATPase, transmembrane domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1000">
        <property type="entry name" value="HAD superfamily/HAD-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023299">
        <property type="entry name" value="ATPase_P-typ_cyto_dom_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018303">
        <property type="entry name" value="ATPase_P-typ_P_site"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023298">
        <property type="entry name" value="ATPase_P-typ_TM_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008250">
        <property type="entry name" value="ATPase_P-typ_transduc_dom_A_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036412">
        <property type="entry name" value="HAD-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023214">
        <property type="entry name" value="HAD_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006539">
        <property type="entry name" value="P-type_ATPase_IV"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032631">
        <property type="entry name" value="P-type_ATPase_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001757">
        <property type="entry name" value="P_typ_ATPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032630">
        <property type="entry name" value="P_typ_ATPase_c"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044492">
        <property type="entry name" value="P_typ_ATPase_HD_dom"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01652">
        <property type="entry name" value="ATPase-Plipid"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01494">
        <property type="entry name" value="ATPase_P-type"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24092:SF81">
        <property type="entry name" value="PHOSPHOLIPID-TRANSPORTING ATPASE VA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24092">
        <property type="entry name" value="PROBABLE PHOSPHOLIPID-TRANSPORTING ATPASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13246">
        <property type="entry name" value="Cation_ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16212">
        <property type="entry name" value="PhoLip_ATPase_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16209">
        <property type="entry name" value="PhoLip_ATPase_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00119">
        <property type="entry name" value="CATATPASE"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDG00002">
        <property type="entry name" value="C1.7:_P-type_atpase_like"/>
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        <location>
            <begin position="53"/>
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    <feature type="transmembrane region" description="Helical" evidence="26">
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            <begin position="308"/>
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            <begin position="1042"/>
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            <begin position="1074"/>
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            <begin position="1124"/>
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            <begin position="1153"/>
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            <begin position="1181"/>
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            <begin position="1221"/>
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        <location>
            <begin position="25"/>
            <end position="76"/>
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            <begin position="1010"/>
            <end position="1252"/>
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            <begin position="421"/>
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            <begin position="1321"/>
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            <begin position="434"/>
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</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GK28</accession>
    <name>A0A452GK28_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3 7">Carbonic anhydrase</fullName>
            <ecNumber evidence="3 7">4.2.1.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="7">Reversible hydration of carbon dioxide.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="7">
            <text>hydrogencarbonate + H(+) = CO2 + H2O</text>
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        </reaction>
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            <name>Zn(2+)</name>
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        </cofactor>
    </comment>
    <comment type="similarity">
        <text evidence="2 7">Belongs to the alpha-carbonic anhydrase family.</text>
    </comment>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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        <property type="term" value="F:carbonate dehydratase activity"/>
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        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
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        <property type="entry name" value="Carbonic anhydrase 2"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Alpha carbonic anhydrase"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="CA_dom"/>
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    <dbReference type="InterPro" id="IPR036398">
        <property type="entry name" value="CA_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023561">
        <property type="entry name" value="Carbonic_anhydrase_a-class"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018338">
        <property type="entry name" value="Carbonic_anhydrase_a-class_CS"/>
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    <dbReference type="PANTHER" id="PTHR18952">
        <property type="entry name" value="CARBONIC ANHYDRASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18952:SF25">
        <property type="entry name" value="CARBONIC ANHYDRASE 5B, MITOCHONDRIAL-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Carb_anhydrase"/>
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        <property type="entry name" value="Carb_anhydrase"/>
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        <property type="entry name" value="ALPHA_CA_1"/>
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    </dbReference>
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        <property type="entry name" value="ALPHA_CA_2"/>
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    </dbReference>
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    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 7" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="Alpha-carbonic anhydrase" evidence="8">
        <location>
            <begin position="4"/>
            <end position="266"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010718"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012925"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU367011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51144"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002072.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="287" mass="33135" checksum="4C512E4167EC9FB2" modified="2019-05-08" version="1">FMLRGWRENCLVTCHVNVHPLWQSELSVSGGTRQSPINIQWRDSVYDPHLKPLKIHYDPTTCLYIWNNGYSFQVEFDDSADKSVIMGGPLENHYRLKQFHFHWGAINEWGSEHMVDSKVYPAELHLVHWNPVTSTSFEEAIMENNGLVVIGVFLKLGAHHAGLQKLVDALPAVKHKDTIVEFDVFDPSCLMPSCPDYWTYDGSLTTPPLTESVTWIIKKTPIEVDQNQLEEFRMLLFTSAGEEEKRMVDNFRPLQPLMNRTVRSSFRNHHLLNIETRSSELAHQHSP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GK29</accession>
    <name>A0A452GK29_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="16">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GK29"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002217.1">
        <property type="protein sequence ID" value="ENSGAGP00000001949.1"/>
        <property type="gene ID" value="ENSGAGG00000001571.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071011">
        <property type="term" value="C:precatalytic spliceosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003725">
        <property type="term" value="F:double-stranded RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003727">
        <property type="term" value="F:single-stranded RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051607">
        <property type="term" value="P:defense response to virus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd19910">
        <property type="entry name" value="DSRM_ILF3_rpt1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd19912">
        <property type="entry name" value="DSRM_ILF3_rpt2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.1410.40:FF:000001">
        <property type="entry name" value="interleukin enhancer-binding factor 3 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.20:FF:000006">
        <property type="entry name" value="interleukin enhancer-binding factor 3 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.20:FF:000008">
        <property type="entry name" value="interleukin enhancer-binding factor 3 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1410.40">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.20">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.460.10">
        <property type="entry name" value="Beta Polymerase, domain 2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014720">
        <property type="entry name" value="dsRBD_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033099">
        <property type="entry name" value="DSRM1_ILF3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006561">
        <property type="entry name" value="DZF_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049402">
        <property type="entry name" value="DZF_dom_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049401">
        <property type="entry name" value="DZF_dom_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043519">
        <property type="entry name" value="NT_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45762:SF4">
        <property type="entry name" value="INTERLEUKIN ENHANCER-BINDING FACTOR 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45762">
        <property type="entry name" value="ZINC FINGER RNA-BINDING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00035">
        <property type="entry name" value="dsrm"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20965">
        <property type="entry name" value="DZF_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07528">
        <property type="entry name" value="DZF_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00358">
        <property type="entry name" value="DSRM"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00572">
        <property type="entry name" value="DZF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54768">
        <property type="entry name" value="dsRNA-binding domain-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50137">
        <property type="entry name" value="DS_RBD"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51703">
        <property type="entry name" value="DZF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8" id="KW-0051">Antiviral defense</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="6 12" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="10" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="DZF" evidence="15">
        <location>
            <begin position="2"/>
            <end position="375"/>
        </location>
    </feature>
    <feature type="domain" description="DRBM" evidence="14">
        <location>
            <begin position="395"/>
            <end position="464"/>
        </location>
    </feature>
    <feature type="domain" description="DRBM" evidence="14">
        <location>
            <begin position="526"/>
            <end position="592"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="48"/>
            <end position="77"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="360"/>
            <end position="398"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="464"/>
            <end position="527"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="635"/>
            <end position="663"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="721"/>
            <end position="901"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="13">
        <location>
            <begin position="57"/>
            <end position="70"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="13">
        <location>
            <begin position="369"/>
            <end position="380"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="13">
        <location>
            <begin position="469"/>
            <end position="478"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="13">
        <location>
            <begin position="496"/>
            <end position="514"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="13">
        <location>
            <begin position="646"/>
            <end position="663"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="13">
        <location>
            <begin position="742"/>
            <end position="798"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="13">
        <location>
            <begin position="813"/>
            <end position="832"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="13">
        <location>
            <begin position="840"/>
            <end position="858"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="13">
        <location>
            <begin position="864"/>
            <end position="886"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023118"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00266"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50137"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS51703"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001949.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="901" mass="96223" checksum="BFB6E018E8ACC04E" modified="2019-05-08" version="1">MRIFVNDDRHVMAKHSAVYPTQEELEAVQNMVSHTERALKAVSDWIDEQEKVSGEQPEPEAMETAAEEENKEGGDQKAAEHLTRTLRGVMRVGLVAKGLLLKGDLDLELVLLCKEKPTTGLLDKVAENLGVQLATITEDKYEIIQSVSEAAIIIKNTQEPPLTLTIHLTSPVVREEMEKLLAGETLSVNDTPDVLDRQKCLAALASLRHAKWFQARANGLKSCVIVIRVLRDLCTRVPTWGPLRGWPLELLCEKSIGTANRPMGAGEALRRVLECLASGIVMPDGSGIYDPCEKEATDAIGHLDRQQREDITQSAQHALRLAAFGQLHKVLGMDPLPSKMPKKPKNENPVDYTVQIPPSTTYAITPMKRPMEEDGEEKSPSKKKKKIQKKEEKTEPPQAMNALMKLNQLKPGLQYKLVSQTGPVHAPIFTMSVEVDGSTFEASGPSKKTAKLHVAVKVLQDMGLPTGVEGKESGKGDESAEETEQKPVVVAPPPVVETISTPSAASPPSEQTSENVKQQGPILTKHGKNPVMELNEKRRGLKYELISETGGSHDKRFVMEVEVDGQKFQGAGSNKKVAKAYAALAALEKLFPDTPVAIEPNKKKRAPVPARGGPKFAVKQHNPGFGMGGPMHNEVPPPPNLRGRGRGGNIRGRGRGRGGFGGGNHGGYMNAGAGYGSYGYGGNSATAGYSQFYSNGGHSGNTGSGGGGGGGGGSSGYGSYYQGGDNYSSPVPPKHAGKKQQHGGQQKPSYGSGYQSHQGQQQQSYNQNQYSNYGPPQGKQKGYNHGQGNYSYSNSYNSSGGGSDYNYESKFNYGGGSGRGGGGNNYGSGGGSSYNTGSHSGYGGGSGGGGSSYQGKQGGYSSQTNYSSPGSGQNYSGPPSSYQASQGGYGRNDHSMNYQYR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GK54</accession>
    <name>A0A452GK54_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Focal adhesion kinase 1</fullName>
            <ecNumber evidence="4">2.7.10.2</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="19">Protein-tyrosine kinase 2</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="20">pp125FAK</fullName>
        </alternativeName>
    </protein>
    <organism evidence="26 27">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="27" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="26" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="26" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="21 28 29">
            <text>L-tyrosyl-[protein] + ATP = O-phospho-L-tyrosyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10596"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.10.2"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell junction</location>
            <location evidence="2">Focal adhesion</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cell membrane</location>
            <topology evidence="3">Peripheral membrane protein</topology>
            <orientation evidence="3">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Cilium basal body</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="2.7.10.2" evidence="4"/>
    <dbReference type="AlphaFoldDB" id="A0A452GK54"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002249.1">
        <property type="protein sequence ID" value="ENSGAGP00000001976.1"/>
        <property type="gene ID" value="ENSGAGG00000001572.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005925">
        <property type="term" value="C:focal adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004715">
        <property type="term" value="F:non-membrane spanning protein tyrosine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008284">
        <property type="term" value="P:positive regulation of cell population proliferation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007172">
        <property type="term" value="P:signal complex assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd14473">
        <property type="entry name" value="FERM_B-lobe"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd13190">
        <property type="entry name" value="FERM_C_FAK1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05056">
        <property type="entry name" value="PTKc_FAK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.330:FF:000001">
        <property type="entry name" value="focal adhesion kinase 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000058">
        <property type="entry name" value="focal adhesion kinase 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.20.90:FF:000021">
        <property type="entry name" value="focal adhesion kinase 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000047">
        <property type="entry name" value="focal adhesion kinase 1 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000039">
        <property type="entry name" value="Focal adhesion kinase, isoform D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.80.10:FF:000004">
        <property type="entry name" value="Protein-tyrosine kinase 2-beta isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.80.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.330">
        <property type="entry name" value="Nucleotidyltransferases domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.20.90">
        <property type="entry name" value="Phosphatidylinositol 3-kinase Catalytic Subunit, Chain A, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.540">
        <property type="entry name" value="Single helix bin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019749">
        <property type="entry name" value="Band_41_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041390">
        <property type="entry name" value="FADK_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049385">
        <property type="entry name" value="FAK1-like_FERM_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041784">
        <property type="entry name" value="FAK1/PYK2_FERM_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014352">
        <property type="entry name" value="FERM/acyl-CoA-bd_prot_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035963">
        <property type="entry name" value="FERM_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019748">
        <property type="entry name" value="FERM_central"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000299">
        <property type="entry name" value="FERM_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036137">
        <property type="entry name" value="Focal_adhe_kin_target_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005189">
        <property type="entry name" value="Focal_adhesion_kin_target_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001245">
        <property type="entry name" value="Ser-Thr/Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008266">
        <property type="entry name" value="Tyr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020635">
        <property type="entry name" value="Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029071">
        <property type="entry name" value="Ubiquitin-like_domsf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46221">
        <property type="entry name" value="FERM AND PDZ DOMAIN-CONTAINING PROTEIN FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46221:SF12">
        <property type="entry name" value="NON-SPECIFIC PROTEIN-TYROSINE KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21477">
        <property type="entry name" value="FERM_C_FAK1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00373">
        <property type="entry name" value="FERM_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18038">
        <property type="entry name" value="FERM_N_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03623">
        <property type="entry name" value="Focal_AT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07714">
        <property type="entry name" value="PK_Tyr_Ser-Thr"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00109">
        <property type="entry name" value="TYRKINASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00295">
        <property type="entry name" value="B41"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00219">
        <property type="entry name" value="TyrKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF68993">
        <property type="entry name" value="FAT domain of focal adhesion kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47031">
        <property type="entry name" value="Second domain of FERM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54236">
        <property type="entry name" value="Ubiquitin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50057">
        <property type="entry name" value="FERM_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00109">
        <property type="entry name" value="PROTEIN_KINASE_TYR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="12 22" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="13" id="KW-0965">Cell junction</keyword>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="17" id="KW-0966">Cell projection</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="16" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="5" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="11" id="KW-0418">Kinase</keyword>
    <keyword evidence="14" id="KW-0472">Membrane</keyword>
    <keyword evidence="10 22" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="8" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="27" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0808">Transferase</keyword>
    <keyword evidence="15" id="KW-0829">Tyrosine-protein kinase</keyword>
    <feature type="domain" description="FERM" evidence="25">
        <location>
            <begin position="35"/>
            <end position="355"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="24">
        <location>
            <begin position="435"/>
            <end position="693"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="23">
        <location>
            <begin position="1"/>
            <end position="27"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="23">
        <location>
            <begin position="699"/>
            <end position="754"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="23">
        <location>
            <begin position="850"/>
            <end position="933"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="23">
        <location>
            <begin position="10"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="23">
        <location>
            <begin position="699"/>
            <end position="710"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="23">
        <location>
            <begin position="850"/>
            <end position="862"/>
        </location>
    </feature>
    <feature type="binding site" evidence="22">
        <location>
            <position position="467"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004120"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004413"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00011903"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023137"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00039644"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00042078"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00043012"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00051245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS50057"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001976.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1065" mass="120572" checksum="B944853BA624EE10" modified="2019-05-08" version="1">MAAAYLDPNLNHTPSSSAKTHLSTGVERSPGAMERVLKVFHYFESSSEPTTWASIIRHGDATDVRGIIQKIVDSHKVKNVACYGLRLSHLQSEEVHWLHLDMGVSNVRERFELAHPPEEWKYELRIRYLPKGFLNQFTEDKPTLNFFYQQVKNDYMLEIADQVDQEIALKLGCLEIRRSYWEMRGNALEKKSNYEVLEKDVGLKRFFPKSLLDSVKAKTLRKLIQQTFRQFAVLNREESIQKFFEILSPVYRFDKECFKCALGSSWIISVELAIGPEEGISYLTDKGSNPTHLADFNQVQTIQYSNSEDKDRKGMLQLKIAGAPEPLTVTAPSLTIAENMADLIDGYCRLVNGATQSFIIRPQKEGERALPSIPKLANNEKQGVRSHTVSVSDEISGDETDDYAEIIDEEDTYTMPSKSYGIDEARDYEIQRERIELGRCIGEGQFGDVHQGVYMSPENPAMAVAIKTCKNCTSDSVREKFLQEALTMRQFDHPHIVKLIGVITENPVWIIMELCTLGELRSFLQVRKYSLDLASLILYAYQLSTALAYLESKRFVHRDIAARNVLVSSTDCVKLGDFGLSRYMEDSTYYKASKGKLPIKWMAPESINFRRFTSASDVWMFGVCMWEILMHGVKPFQGVKNNDVIGRIENGERLPMPPNCPPTLYSLMTKCWAYDPSRRPRFTELKAQLSTILEEEKLQQEERMRMESRRQVTVSWDSGGSDEAPPKPSRPGYPSPRSSEGFYPSPQHMVPPNHYQVSGYPGSHGIPAMAGSIYPGQASLLDQADSWNHRPQEISMWPPTMEDSGTLDMRGMGQVLPTHLMEERLIRQQQEMEEDQRWLEKEERFLKPDVRLSRGSIDREDGSLQGPAGNQHIYQPVGKPDHAAPPKKPPRPGAPSHLGSLASLNSPVDSYNEGVKLQPQEISPPPTANLDRSNDKVYENVTGLVKAVIEMSSKIQPAPPEEYVPMVKEVGLALRTLLATVDETIPVLPASTHREIEMAQKLLNSDLAELINKMKLAQQYVMTSLQQEYKKQMLTAAHALAVDAKNLLDVIDQARLKMIGQSRPH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GK56</accession>
    <name>A0A452GK56_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">EGF-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cell projection</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the tenascin family. Teneurin subfamily.</text>
    </comment>
    <comment type="caution">
        <text evidence="13">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000002365.1">
        <property type="protein sequence ID" value="ENSGAGP00000002078.1"/>
        <property type="gene ID" value="ENSGAGG00000000665.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043005">
        <property type="term" value="C:neuron projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050839">
        <property type="term" value="F:cell adhesion molecule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046982">
        <property type="term" value="F:protein heterodimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042803">
        <property type="term" value="F:protein homodimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007157">
        <property type="term" value="P:heterophilic cell-cell adhesion via plasma membrane cell adhesion molecules"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048666">
        <property type="term" value="P:neuron development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00054">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.180.10.10">
        <property type="entry name" value="RHS repeat-associated core"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.120.10.30">
        <property type="entry name" value="TolB, C-terminal domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011042">
        <property type="entry name" value="6-blade_b-propeller_TolB-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008969">
        <property type="entry name" value="CarboxyPept-like_regulatory"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057627">
        <property type="entry name" value="FN-plug_TEN1-4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022385">
        <property type="entry name" value="Rhs_assc_core"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056823">
        <property type="entry name" value="TEN-like_YD-shell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056822">
        <property type="entry name" value="TEN_NHL"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056820">
        <property type="entry name" value="TEN_TTR-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051216">
        <property type="entry name" value="Teneurin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028916">
        <property type="entry name" value="Tox-GHH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006530">
        <property type="entry name" value="YD"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR03696">
        <property type="entry name" value="Rhs_assc_core"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01643">
        <property type="entry name" value="YD_repeat_2x"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11219">
        <property type="entry name" value="TENEURIN AND N-ACETYLGLUCOSAMINE-1-PHOSPHODIESTER ALPHA-N-ACETYLGLUCOSAMINIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11219:SF9">
        <property type="entry name" value="TENEURIN-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25024">
        <property type="entry name" value="EGF_TEN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24329">
        <property type="entry name" value="FN-plug_TEN1-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25021">
        <property type="entry name" value="TEN_NHL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25023">
        <property type="entry name" value="TEN_YD-shell"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23538">
        <property type="entry name" value="Teneurin_ABD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15636">
        <property type="entry name" value="Tox-GHH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25020">
        <property type="entry name" value="TTR_TEN1-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF63829">
        <property type="entry name" value="Calcium-dependent phosphotriesterase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49464">
        <property type="entry name" value="Carboxypeptidase regulatory domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01186">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="12" id="KW-0966">Cell projection</keyword>
    <keyword evidence="11 13" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6 13" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="10" id="KW-0472">Membrane</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="14" id="KW-0732">Signal</keyword>
    <keyword evidence="7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="14">
        <location>
            <begin position="1"/>
            <end position="18"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5045512604" description="EGF-like domain-containing protein" evidence="14">
        <location>
            <begin position="19"/>
            <end position="2003"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="15">
        <location>
            <begin position="68"/>
            <end position="103"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="13">
        <location>
            <begin position="72"/>
            <end position="82"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="13">
        <location>
            <begin position="93"/>
            <end position="102"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004167"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004316"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00009385"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002078.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2003" mass="224822" checksum="4F4E468CC5B7E65F" modified="2025-10-08" version="2">MTLLCSAVICAADCGGHGICVGGTCRCEEGWMGAACDQRACHPRCNEHGTCRDGKCECSPGWNGEHCTIEGCPGLCNGNGRCTLDMNGWHCVCQLGWRGAGCDTSMETACGDGKDNDGDGLVDCMDPDCCLQPLCHVNPLCLGSPDPLDIIQETQTPTSQQNLHSFYDRIKFLIGKDSTHVIPGENPFDGGHACVIRGQVITSDGTPLVGVNISFVNNPLFGYTISRQDGSFDLVTNGGISIILHFERAPFITQEHTLWLPWDRFFVMETIVMRHEENEIPSCDLSNFARPNPVVSPSPLTAFASSCSEKGPIVPEIQALQEEINIAGSKIKLSYLSSRTAGYKSVLRISMTHPTIPFNLMKVHLMVAVEGRLFRKWFAAAPDLSYYFIWDKTDVYSQKVYGLSEAFVSVGYEYESCPDLILWEKRTAVLQGYEIDASKLGGWSLDQHHALNIQSGILHKGNGENQFVSQQPPVIGSIMGNGRRRSISCPSCNGLADGNKLLAPVALTCGSDGSLYVGDFNYIRRIFPSGNVTNILELSHSPAHKYYLTSDPITGSIYLSDTNSRRIYKIKSTSVVKDVVKNSEVVAGTGDQCLPFDETRCGDGGKGTEATLTNPRGKTQSLLTSILKWASRKVPNQQFWNTTEQVYKLPQGNPLMCLRVRLEWPTDLAVSPMDNSLYVLDNNVVLQISENHQVRIVAGRPMHCQVPGIDHFLLSKVAIHATLESATALAVSHNGVLYIAETDEKKINRIRQVTTNGEISLVAGAPSSCDCKNDANCDCFSGDDGYAKDARLNAPSSLAVCADGELYVADLGNIRIRFIRKNKPYLNTQNLYELSSPIDQELYLFDTSGKHLYTQSLPTGDYLYNFTYTGDGDITLITDNNGNLVNVRRDSTGMPLWLVVPDGQVYWVTIGTNSALKSVTAQGHEVGMMTYHGNSGLLATKTNENGWTTFYEYDSFGRLTNVTFPTGQVSSFRSDTDRSVHVQVETSSKDDVTITTNLSASGAFYTLLQDQVRNSYYIGADGSLRLMLANGMEVALQTEPHLLAGTVNPTVGKRNVTLPIDNGLNLVEWRQRKEQARGQVTVFGRRLRVHNRNLLSLDFDRVTRTEKIYDDHRKFTLRILYDQAGRPSLWSPSRGHIAGIQRGTMSERMEYDQAGRIISRIFADGKSWSYTYLEKSMVLLLHSQRQYIFEFDKNDRLSSVTMPNVARQTLETIRSIEGNASVIQDFTEEGQLLHTFYLGTGRRVIYKYGKLSKLAEMLYDTTKIGFTYDETAGMLKTINLQNEGFTCTIRYRQIGPLIDRQIFRFTEEGMVNARFDYNYDNSFRVTSMQAVINETPLPIDLYRYDDVSGKTEQFGKFGVIYYDINQIITTAVMTHTKHFDAYGRMKEVQYEIFRSLMYWMTVQYDNMGRVVKKELKVGPYANTTRYSYEYDADGQLQTVSINDKPLWRYSYDLNGNLHLLSPGNSARLTPLRYDLRDRITRLGDVQYKMDEDGFLRQRGNDIFEYNSAGLLIKAYNKVSGWSVKYRYDGLGRRVSSKTAHSHHLQFFYADLTNPTKVTHLYNHSSSEITSLYYDLQGHLFAMELSSGDEFYIACDNIGTPLAVFSGTGLMIKQILYTAYGEIYMDTNPNFQVIIGYHGGLYDPLTKLIHMGRRDYDVLAGRWTSPDHDMWKHLSSNNIMPFNLYMFKNNNPISNAQDIKCYMTDVNSWLLTFGFQLHNVIPGYPKPDTDAMEPSYELIHTQMKTQEWDNSKSILGVQCEVQKQLKAFVTLERFEQIYSSSIAGCRTVKENKNFASGGSVFGKGVKFAMKDGRVATDIISLANEDGRRIAAILNNAHYLENLHFTIEGVDTHYFVKQGPSESDLSILGLSGGRRTLENGVNVTVSQINTVLSGRTRRYTDIQLQYGALCLNTRYGTTLDEEKARVLELARQRAIAQAWAREQQRLRDGEEGIRSWTEGEKQQVLNTGRVQGYDGYFVISVEQYPELSDSANNIHFMRQSEMGRR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GK59</accession>
    <name>A0A452GK59_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="26">Phospholipid-transporting ATPase</fullName>
            <ecNumber evidence="26">7.6.2.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="30 31">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="31" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="30" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="30" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="21">Catalytic component of P4-ATPase flippase complex, which catalyzes the hydrolysis of ATP coupled to the transport of phosphatidylcholine (PC) from the outer to the inner leaflet of the plasma membrane. Initiates inward plasma membrane bending and recruitment of Bin/amphiphysin/Rvs (BAR) domain-containing proteins involved in membrane tubulation and cell trafficking. Facilitates ITGB1/beta1 integrin endocytosis, delaying cell adhesion and cell spreading on extracellular matrix. Has low flippase activity toward glucosylceramide (GlcCer).</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="18 26">
            <text>ATP + H2O + phospholipidSide 1 = ADP + phosphate + phospholipidSide 2.</text>
            <dbReference type="EC" id="7.6.2.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="20">
            <text>a 1,2-diacyl-sn-glycero-3-phosphocholine(out) + ATP + H2O = a 1,2-diacyl-sn-glycero-3-phosphocholine(in) + ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:38583"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:57643"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="20">
            <dbReference type="Rhea" id="RHEA:38584"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>a beta-D-glucosyl-(1&lt;-&gt;1')-N-acylsphing-4-enine(out) + ATP + H2O = a beta-D-glucosyl-(1&lt;-&gt;1')-N-acylsphing-4-enine(in) + ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:66036"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:22801"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="19">
            <dbReference type="Rhea" id="RHEA:66037"/>
        </physiologicalReaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 25">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subunit">
        <text evidence="22">Component of a P4-ATPase flippase complex which consists of a catalytic alpha subunit ATP10A and an accessory beta subunit TMEM30A.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cell membrane</location>
            <topology evidence="3">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Endoplasmic reticulum membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="26">Membrane</location>
            <topology evidence="26">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4 26">Belongs to the cation transport ATPase (P-type) (TC 3.A.3) family. Type IV subfamily.</text>
    </comment>
    <dbReference type="EC" id="7.6.2.1" evidence="26"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002398.1">
        <property type="protein sequence ID" value="ENSGAGP00000002108.1"/>
        <property type="gene ID" value="ENSGAGG00000000613.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:1990531">
        <property type="term" value="C:phospholipid-translocating ATPase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140351">
        <property type="term" value="F:glycosylceramide flippase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000287">
        <property type="term" value="F:magnesium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090554">
        <property type="term" value="F:phosphatidylcholine floppase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="RHEA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045332">
        <property type="term" value="P:phospholipid translocation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd02073">
        <property type="entry name" value="P-type_ATPase_APLT_Dnf-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.70.150.10:FF:000022">
        <property type="entry name" value="Phospholipid-transporting ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.1110.10:FF:000009">
        <property type="entry name" value="Phospholipid-transporting ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.1110.10:FF:000109">
        <property type="entry name" value="Phospholipid-transporting ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1000:FF:000110">
        <property type="entry name" value="Phospholipid-transporting ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1000:FF:000001">
        <property type="entry name" value="Phospholipid-transporting ATPase IC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.1110.10">
        <property type="entry name" value="Calcium-transporting ATPase, cytoplasmic domain N"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.70.150.10">
        <property type="entry name" value="Calcium-transporting ATPase, cytoplasmic transduction domain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1110.10">
        <property type="entry name" value="Calcium-transporting ATPase, transmembrane domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1000">
        <property type="entry name" value="HAD superfamily/HAD-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023299">
        <property type="entry name" value="ATPase_P-typ_cyto_dom_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018303">
        <property type="entry name" value="ATPase_P-typ_P_site"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023298">
        <property type="entry name" value="ATPase_P-typ_TM_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008250">
        <property type="entry name" value="ATPase_P-typ_transduc_dom_A_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036412">
        <property type="entry name" value="HAD-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023214">
        <property type="entry name" value="HAD_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006539">
        <property type="entry name" value="P-type_ATPase_IV"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032631">
        <property type="entry name" value="P-type_ATPase_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001757">
        <property type="entry name" value="P_typ_ATPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032630">
        <property type="entry name" value="P_typ_ATPase_c"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044492">
        <property type="entry name" value="P_typ_ATPase_HD_dom"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01652">
        <property type="entry name" value="ATPase-Plipid"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01494">
        <property type="entry name" value="ATPase_P-type"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24092:SF81">
        <property type="entry name" value="PHOSPHOLIPID-TRANSPORTING ATPASE VA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24092">
        <property type="entry name" value="PROBABLE PHOSPHOLIPID-TRANSPORTING ATPASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13246">
        <property type="entry name" value="Cation_ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16212">
        <property type="entry name" value="PhoLip_ATPase_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16209">
        <property type="entry name" value="PhoLip_ATPase_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00119">
        <property type="entry name" value="CATATPASE"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDG00002">
        <property type="entry name" value="C1.7:_P-type_atpase_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDS00003">
        <property type="entry name" value="Haloacid_Dehalogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDF00027">
        <property type="entry name" value="p-type_atpase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81653">
        <property type="entry name" value="Calcium ATPase, transduction domain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81665">
        <property type="entry name" value="Calcium ATPase, transmembrane domain M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56784">
        <property type="entry name" value="HAD-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81660">
        <property type="entry name" value="Metal cation-transporting ATPase, ATP-binding domain N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00154">
        <property type="entry name" value="ATPASE_E1_E2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12 24 26" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="16" id="KW-0445">Lipid transport</keyword>
    <keyword evidence="13 25 26" id="KW-0460">Magnesium</keyword>
    <keyword evidence="17 26" id="KW-0472">Membrane</keyword>
    <keyword evidence="9 25" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10 24 26" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="31" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="14 26" id="KW-1278">Translocase</keyword>
    <keyword evidence="8 26" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="15 26" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="5" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="26">
        <location>
            <begin position="72"/>
            <end position="90"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="26">
        <location>
            <begin position="96"/>
            <end position="114"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="26">
        <location>
            <begin position="294"/>
            <end position="317"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="26">
        <location>
            <begin position="337"/>
            <end position="359"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="26">
        <location>
            <begin position="1071"/>
            <end position="1091"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="26">
        <location>
            <begin position="1103"/>
            <end position="1123"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="26">
        <location>
            <begin position="1153"/>
            <end position="1176"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="26">
        <location>
            <begin position="1182"/>
            <end position="1203"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="26">
        <location>
            <begin position="1210"/>
            <end position="1230"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="26">
        <location>
            <begin position="1250"/>
            <end position="1269"/>
        </location>
    </feature>
    <feature type="domain" description="P-type ATPase N-terminal" evidence="28">
        <location>
            <begin position="42"/>
            <end position="95"/>
        </location>
    </feature>
    <feature type="domain" description="P-type ATPase C-terminal" evidence="29">
        <location>
            <begin position="1039"/>
            <end position="1281"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="27">
        <location>
            <begin position="1"/>
            <end position="48"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="27">
        <location>
            <begin position="449"/>
            <end position="498"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="27">
        <location>
            <begin position="603"/>
            <end position="623"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="27">
        <location>
            <begin position="1350"/>
            <end position="1382"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="27">
        <location>
            <begin position="10"/>
            <end position="25"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="27">
        <location>
            <begin position="463"/>
            <end position="475"/>
        </location>
    </feature>
    <feature type="active site" description="4-aspartylphosphate intermediate" evidence="23">
        <location>
            <position position="413"/>
        </location>
    </feature>
    <feature type="binding site" evidence="24">
        <location>
            <position position="413"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="25">
        <location>
            <position position="413"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="24">
        <location>
            <position position="414"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="24">
        <location>
            <position position="415"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="25">
        <location>
            <position position="415"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="24">
        <location>
            <position position="684"/>
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        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
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        <location>
            <position position="726"/>
        </location>
        <ligand>
            <name>ATP</name>
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    <feature type="binding site" evidence="24">
        <location>
            <position position="750"/>
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        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
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        <location>
            <position position="793"/>
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        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
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        <location>
            <position position="873"/>
        </location>
        <ligand>
            <name>ATP</name>
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        </ligand>
    </feature>
    <feature type="binding site" evidence="24">
        <location>
            <position position="874"/>
        </location>
        <ligand>
            <name>ATP</name>
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        </ligand>
    </feature>
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        <location>
            <position position="875"/>
        </location>
        <ligand>
            <name>ATP</name>
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        <location>
            <position position="987"/>
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        <ligand>
            <name>ATP</name>
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        <location>
            <position position="993"/>
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        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
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        <location>
            <position position="1013"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
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        </ligand>
    </feature>
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        <location>
            <position position="1016"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
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        <location>
            <position position="1017"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
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        <location>
            <position position="1017"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004586"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
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    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00008109"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
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    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
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    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
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    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
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    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
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    </evidence>
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            <dbReference type="ARBA" id="ARBA00022824"/>
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    </evidence>
    <evidence type="ECO:0000256" key="12">
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            <dbReference type="ARBA" id="ARBA00022840"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
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    </evidence>
    <evidence type="ECO:0000256" key="14">
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            <dbReference type="ARBA" id="ARBA00022967"/>
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    </evidence>
    <evidence type="ECO:0000256" key="15">
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023055"/>
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    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
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    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00034036"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00050913"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00052223"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00056521"/>
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            <dbReference type="ARBA" id="ARBA00065679"/>
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    </evidence>
    <evidence type="ECO:0000256" key="23">
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            <dbReference type="PIRSR" id="PIRSR606539-1"/>
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    <evidence type="ECO:0000256" key="24">
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            <dbReference type="PIRSR" id="PIRSR606539-2"/>
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            <dbReference type="PIRSR" id="PIRSR606539-3"/>
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    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="RuleBase" id="RU362033"/>
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    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="28">
        <source>
            <dbReference type="Pfam" id="PF16209"/>
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    <evidence type="ECO:0000259" key="29">
        <source>
            <dbReference type="Pfam" id="PF16212"/>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002108.1"/>
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    </evidence>
    <evidence type="ECO:0000313" key="31">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1490" mass="168566" checksum="86696D6C9FC82C5F" modified="2019-05-08" version="1">RVRESEGKKKLSPPKHRRRRKRTRTVRSNLLLPGPEAEKPQRSPMANNRLKTTKYTALSFLPKNLFEQFHRLANVYFVFIALLNFVPAVNAFQPELALAPVLFILAVTAIKDLWEDYSRYRSDNEINHTECLVYCRNEKKYMNRYWKEVEVGDFVQLRCNEIIPADILLLSSSDPDGLCHIETANLDGETNLKQRQVVRRFLELDSEFDPLKFTSVIECEKPNNDLSRFRGYIVHTSGKRDGLYKENLLLRGCTVRNTEEVAGIVIYAGHETKALLNNSGPRYKRSKLERQMNIDVLWCVLILIVMCLFSTIGHGLWVWQYGEKKPVFDVPGLDGNYLSPVLAAVYIFLTMIIVFQVLIPISLYVSIEIVKACQVFFIHQDKDLYDEETDSQLQCRALNITEDLGQMQYIFSDKTGTLTENKMVFRRCTVSGIEYSHDANARRLAMYQEADSEEEETAAKGGTLSQRNSVSSHQSIKVVHRSLSMKSHRRTGSRAEAKRASILSKHTAFSSPMEKDITPDPNLLEKVNECARSLEVTRSHEQPLSKLTPELSDIFDFFIALTICNTVVVTSPDQPRQKVRVRFELKSPVKTIEDFIRRFTPSRLTSGSNSSSSSSLTTSKSMPRLGSSILSSVSSESTLLKLEEKLAHSPQMNNNDYRSQQDRRAIGCRPEEGELRYEAESPDEAALVYAARAYNCALVGRLSDQVSVELPHLGRLTFELLHTLGFDSIRKRMSVVVRHPLTDEISVYTKGADSVIMDLLLPCSSDDPRGNHQKKIQSKTQNYLNLYAVDGLRTLCIAKRVLSKEEYASWLKSHLEAESSIENREELLFQSALRIETNLHLLGATGVEDRLQDGVPETIANLRKAGLQIWILTGDKQETAINIAYACKLLGHDEEIITLNAESLETSATLLDHCLQYIESKFSNNTIDKTAGNTTVGFCPLSPPSSSVHSSLGLVIDGKTLAYALDKTLEDKFLSLARRCRSVLCCRSTPLQKSMVVKLVRDKLKAMTLAIGDGANDVSMIQVADVGVGISGPEGMQAVMASDFAVPRFRHLEKLLLVHGHWCYSRLANMVLYFFYKNAMFVALLFWYQFYCGFSGSSMIDQWYLIFFNVLFSSLPQLVTGVLDKDIPAEVLIGVPHLYKSGQKMEEYQPHMFWMNMIDALYQSLVCFYIPFFTYYDSEVDLFSWGTPITVIALFTIILHLAIETKTWTLFHWSSCIFSILLFFIVALVYNVSCPICYPPSNPYWTMEKLMGEPVFYLICIISPVVALLPRYSILFLYRTLQGTIFPTQLQLGRQLNRLPPETHSQILNRLNIKKETIPQKPPCAEHSPQNPLSNDSDCFKACNHTPSLQSSQYGNDPAFPSNTKAEHTTGLSVTASPPESLLCGGNKQMESMSVEETFPWNTTVDQSGFSLLNWITSTPLFSRFGRVLQLSPHRLQNEAQDNMCVLGSISSLHRDFKGLTGKNSNDFQEQLKGTPTNCCKSETLETTFL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GK61</accession>
    <name>A0A452GK61_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Focal adhesion kinase 1</fullName>
            <ecNumber evidence="4">2.7.10.2</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="19">Protein-tyrosine kinase 2</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="20">pp125FAK</fullName>
        </alternativeName>
    </protein>
    <organism evidence="26 27">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="27" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="26" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="26" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="21 28 29">
            <text>L-tyrosyl-[protein] + ATP = O-phospho-L-tyrosyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10596"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.10.2"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell junction</location>
            <location evidence="2">Focal adhesion</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cell membrane</location>
            <topology evidence="3">Peripheral membrane protein</topology>
            <orientation evidence="3">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Cilium basal body</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="2.7.10.2" evidence="4"/>
    <dbReference type="AlphaFoldDB" id="A0A452GK61"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002256.1">
        <property type="protein sequence ID" value="ENSGAGP00000001981.1"/>
        <property type="gene ID" value="ENSGAGG00000001572.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005925">
        <property type="term" value="C:focal adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004715">
        <property type="term" value="F:non-membrane spanning protein tyrosine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008284">
        <property type="term" value="P:positive regulation of cell population proliferation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007172">
        <property type="term" value="P:signal complex assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd14473">
        <property type="entry name" value="FERM_B-lobe"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd13190">
        <property type="entry name" value="FERM_C_FAK1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05056">
        <property type="entry name" value="PTKc_FAK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.330:FF:000001">
        <property type="entry name" value="focal adhesion kinase 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000058">
        <property type="entry name" value="focal adhesion kinase 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.20.90:FF:000021">
        <property type="entry name" value="focal adhesion kinase 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000047">
        <property type="entry name" value="focal adhesion kinase 1 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000039">
        <property type="entry name" value="Focal adhesion kinase, isoform D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.80.10:FF:000004">
        <property type="entry name" value="Protein-tyrosine kinase 2-beta isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.80.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.330">
        <property type="entry name" value="Nucleotidyltransferases domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.20.90">
        <property type="entry name" value="Phosphatidylinositol 3-kinase Catalytic Subunit, Chain A, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.540">
        <property type="entry name" value="Single helix bin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019749">
        <property type="entry name" value="Band_41_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041390">
        <property type="entry name" value="FADK_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049385">
        <property type="entry name" value="FAK1-like_FERM_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041784">
        <property type="entry name" value="FAK1/PYK2_FERM_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014352">
        <property type="entry name" value="FERM/acyl-CoA-bd_prot_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035963">
        <property type="entry name" value="FERM_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019748">
        <property type="entry name" value="FERM_central"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000299">
        <property type="entry name" value="FERM_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036137">
        <property type="entry name" value="Focal_adhe_kin_target_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005189">
        <property type="entry name" value="Focal_adhesion_kin_target_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001245">
        <property type="entry name" value="Ser-Thr/Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008266">
        <property type="entry name" value="Tyr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020635">
        <property type="entry name" value="Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029071">
        <property type="entry name" value="Ubiquitin-like_domsf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46221">
        <property type="entry name" value="FERM AND PDZ DOMAIN-CONTAINING PROTEIN FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46221:SF12">
        <property type="entry name" value="NON-SPECIFIC PROTEIN-TYROSINE KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21477">
        <property type="entry name" value="FERM_C_FAK1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00373">
        <property type="entry name" value="FERM_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18038">
        <property type="entry name" value="FERM_N_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03623">
        <property type="entry name" value="Focal_AT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07714">
        <property type="entry name" value="PK_Tyr_Ser-Thr"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00109">
        <property type="entry name" value="TYRKINASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00295">
        <property type="entry name" value="B41"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00219">
        <property type="entry name" value="TyrKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF68993">
        <property type="entry name" value="FAT domain of focal adhesion kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47031">
        <property type="entry name" value="Second domain of FERM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54236">
        <property type="entry name" value="Ubiquitin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50057">
        <property type="entry name" value="FERM_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00109">
        <property type="entry name" value="PROTEIN_KINASE_TYR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="12 22" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="13" id="KW-0965">Cell junction</keyword>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="17" id="KW-0966">Cell projection</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="16" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="5" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="11" id="KW-0418">Kinase</keyword>
    <keyword evidence="14" id="KW-0472">Membrane</keyword>
    <keyword evidence="10 22" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="8" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="27" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0808">Transferase</keyword>
    <keyword evidence="15" id="KW-0829">Tyrosine-protein kinase</keyword>
    <feature type="domain" description="FERM" evidence="25">
        <location>
            <begin position="35"/>
            <end position="355"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="24">
        <location>
            <begin position="429"/>
            <end position="687"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="23">
        <location>
            <begin position="1"/>
            <end position="27"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="23">
        <location>
            <begin position="693"/>
            <end position="748"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="23">
        <location>
            <begin position="844"/>
            <end position="927"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="23">
        <location>
            <begin position="10"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="23">
        <location>
            <begin position="693"/>
            <end position="704"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="23">
        <location>
            <begin position="844"/>
            <end position="856"/>
        </location>
    </feature>
    <feature type="binding site" evidence="22">
        <location>
            <position position="461"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004120"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004413"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00011903"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023137"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00039644"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00042078"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00043012"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00051245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS50057"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001981.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1059" mass="119956" checksum="08E8F589CF612A22" modified="2019-05-08" version="1">MAAAYLDPNLNHTPSSSAKTHLSTGVERSPGAMERVLKVFHYFESSSEPTTWASIIRHGDATDVRGIIQKIVDSHKVKNVACYGLRLSHLQSEEVHWLHLDMGVSNVRERFELAHPPEEWKYELRIRYLPKGFLNQFTEDKPTLNFFYQQVKNDYMLEIADQVDQEIALKLGCLEIRRSYWEMRGNALEKKSNYEVLEKDVGLKRFFPKSLLDSVKAKTLRKLIQQTFRQFAVLNREESIQKFFEILSPVYRFDKECFKCALGSSWIISVELAIGPEEGISYLTDKGSNPTHLADFNQVQTIQYSNSEDKDRKGMLQLKIAGAPEPLTVTAPSLTIAENMADLIDGYCRLVNGATQSFIIRPQKEGERALPSIPKLANNEKQGVRSHTVSVSETDDYAEIIDEEDTYTMPSKSYGIDEARDYEIQRERIELGRCIGEGQFGDVHQGVYMSPENPAMAVAIKTCKNCTSDSVREKFLQEALTMRQFDHPHIVKLIGVITENPVWIIMELCTLGELRSFLQVRKYSLDLASLILYAYQLSTALAYLESKRFVHRDIAARNVLVSSTDCVKLGDFGLSRYMEDSTYYKASKGKLPIKWMAPESINFRRFTSASDVWMFGVCMWEILMHGVKPFQGVKNNDVIGRIENGERLPMPPNCPPTLYSLMTKCWAYDPSRRPRFTELKAQLSTILEEEKLQQEERMRMESRRQVTVSWDSGGSDEAPPKPSRPGYPSPRSSEGFYPSPQHMVPPNHYQVSGYPGSHGIPAMAGSIYPGQASLLDQADSWNHRPQEISMWPPTMEDSGTLDMRGMGQVLPTHLMEERLIRQQQEMEEDQRWLEKEERFLKPDVRLSRGSIDREDGSLQGPAGNQHIYQPVGKPDHAAPPKKPPRPGAPSHLGSLASLNSPVDSYNEGVKLQPQEISPPPTANLDRSNDKVYENVTGLVKAVIEMSSKIQPAPPEEYVPMVKEVGLALRTLLATVDETIPVLPASTHREIEMAQKLLNSDLAELINKMKLAQQYVMTSLQQEYKKQMLTAAHALAVDAKNLLDVIDQARLKMIGQSRPH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GK63</accession>
    <name>A0A452GK63_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Centrosomal protein of 164 kDa</fullName>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Plays a role in microtubule organization and/or maintenance for the formation of primary cilia (PC), a microtubule-based structure that protrudes from the surface of epithelial cells. Plays a critical role in G2/M checkpoint and nuclear divisions. A key player in the DNA damage-activated ATR/ATM signaling cascade since it is required for the proper phosphorylation of H2AX, RPA, CHEK2 and CHEK1. Plays a critical role in chromosome segregation, acting as a mediator required for the maintenance of genomic stability through modulation of MDC1, RPA and CHEK1.</text>
    </comment>
    <comment type="subunit">
        <text evidence="15">Interacts (via N-terminus) with ATRIP. Interacts with ATM, ATR and MDC1. Interacts with XPA (via N-terminus) upon UV irradiation. Interacts with CEP83, CCDC92, TTBK2, DVL3, NPHP3 and weakly with NPHP4. Interacts with DZIP1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Microtubule organizing center</location>
            <location evidence="1">Centrosome</location>
            <location evidence="1">Centriole</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000002370.1">
        <property type="protein sequence ID" value="ENSGAGP00000002083.1"/>
        <property type="gene ID" value="ENSGAGG00000001670.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005814">
        <property type="term" value="C:centriole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097539">
        <property type="term" value="C:ciliary transition fiber"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060271">
        <property type="term" value="P:cilium assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006281">
        <property type="term" value="P:DNA repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00201">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1470.10:FF:000001">
        <property type="entry name" value="Centrosomal protein of 164 kDa"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1470.10">
        <property type="entry name" value="Photosystem I PsaD, reaction center subunit II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051841">
        <property type="entry name" value="MT-Golgi_org_protein"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001202">
        <property type="entry name" value="WW_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036020">
        <property type="entry name" value="WW_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18902:SF27">
        <property type="entry name" value="CENTROSOMAL PROTEIN OF 164 KDA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18902">
        <property type="entry name" value="NUCLEAR MITOTIC APPARATUS PROTEIN 1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00397">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00456">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51045">
        <property type="entry name" value="WW domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01159">
        <property type="entry name" value="WW_DOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50020">
        <property type="entry name" value="WW_DOMAIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="13" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="5" id="KW-0132">Cell division</keyword>
    <keyword evidence="8" id="KW-0970">Cilium biogenesis/degradation</keyword>
    <keyword evidence="9 17" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="6" id="KW-0227">DNA damage</keyword>
    <keyword evidence="10" id="KW-0234">DNA repair</keyword>
    <keyword evidence="7" id="KW-0498">Mitosis</keyword>
    <keyword evidence="12" id="KW-0539">Nucleus</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="WW" evidence="19">
        <location>
            <begin position="58"/>
            <end position="91"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="105"/>
            <end position="151"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="163"/>
            <end position="215"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="237"/>
            <end position="263"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="315"/>
            <end position="372"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="434"/>
            <end position="467"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="505"/>
            <end position="537"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="572"/>
            <end position="604"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="630"/>
            <end position="660"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="895"/>
            <end position="931"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="948"/>
            <end position="979"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1536"/>
            <end position="1556"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="17">
        <location>
            <begin position="993"/>
            <end position="1481"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="18">
        <location>
            <begin position="110"/>
            <end position="126"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="127"/>
            <end position="138"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="169"/>
            <end position="194"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="241"/>
            <end position="257"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="315"/>
            <end position="324"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="337"/>
            <end position="353"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="441"/>
            <end position="450"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="580"/>
            <end position="600"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="633"/>
            <end position="658"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="898"/>
            <end position="907"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="914"/>
            <end position="928"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="956"/>
            <end position="971"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="1541"/>
            <end position="1556"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004114"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022776"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022794"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00056906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00061715"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00067900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50020"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002083.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1926" mass="217222" checksum="1CE3D168AD019B5C" modified="2019-05-08" version="1">MDMAGPAIRIGDQLILEEDYDETYIPSEQEIQEFAREIGIDPGNEPELMWLAREGIVAPLPPEWKPCQDITGDIYYFNFSNGQSTWDHPCDEHYRELVIQEREKLSAHGSIKKKDKKKKKDKKEKKEKKEREPVKHAAEMQPEPGILPSSSFYRMSSPILSSECASPDLDQHSQMRIEGFLKKGKRKPSERPPETSDSNKQFAGPSPNKLQPLFTAKSCRTHQILADVEKILGRMPSFSRSDVDHQQGQDRSPKIGDKPCLGFSDSESEGLDVVRTKPISYVARKSCSQDLENGRSVLDSCEVLEDKGMLFDEEKLSDNWKQEEMTGGSVCQPDDGLSVRERSTGPGGDKERGPGFSQADENEAFLTPSAPDHTFYSRKCEMILFDASPLEDLSLMEEGDVHCPQRELRAHSRAVESRRKRKLQDQALSQATKFCKSSEVVGKHERESSKQAEFVESEGAGLQAAPQKDLLDSHLAVPEVTADTLPYTRGVEAYAHAGADISASEPVSLSAAQHKDKEKANGLGPEPDNSLTSNSADHFASQILGEIDNFSWDLQSSRESDRPTDLLAAAAKGSLPTHPFTGTLQTPPRSSANGKSQSDCNSEDEKFYQHVQENDQVPSQEHSCVSFNMKLNRSRDAPEKNNEGRQEETGEGEQKTEPVPRGLEVDVFTEGLQVSPGWLYQTAECQESSRLCQQVEGELTQSGKVTEDYCKLKLCEGAGSNESPGSLLAPIQVPPGSLAPLRGVMDGPASAFHGLLSTNVEISGVPSPLLREGTLPLQTLKSTGCTKSLLGSMHEEKASLNLLALDEEEEEESENQSPRGTARLLKNLHMDISALGGGFEYEESEASEHVKELQVSDCSDHELLHLTDFGFQSRISEQVLDMDILSAALDSPKVEAQGLREEEKDQSEASLEEEQSKRPKAAESERNQRPFSACKMAEEKCLSLENASQEETAALEQKEGSLDSPKEEEPAAPHQAQLGEEIKQVVSLNWPSAESLNEIARKQEQELEQEKERLLQAKEERVRRFQEELRQEEEEEAQVLHQQKEKSLWSLKEELAKASEEEELRMREEEVEKLSKLRSQISSETKTEKEKIRAEQEAALQKLREEWESLQKVERERLEERKKRVLEMLKVEVEEAQQREAAELEQEKQRVLNELKERLERKRKEAAEVLEKQFAAELQELKSTAEEKHQKVVSSLRKQIAEAQRSEEAQLREELERAEQKVQQKMYQVAEFERELSELMKEKRQEVERDHDRKMERMKEEHKEVLARIQEQYEEEERKQRTKKLEELKGELERLRQLHDGELKALQKQLDEQLSDLRQSHKVKERKLQELEMELEIQATDAKARCAQLNHQEEAMRKKRQQLLDEEKQTELQRNEAALAAQLRLEESRKEHADLVESIRQLRRTLEELQDQKAELESQVDLLQTRSQRLQRRISELEAAVQSKQETLKELVAEDSMASPRREAELHLEDLRETLQAHSSREPASLTSHSNEDSDLQLDNVRHFISAEGVSIKNAKEFLVRQTRSMRKRHTALKAAKQQWRHNMQQAQDAVQDPDSSQLLEDVRRNLEEESAQLDKMKSAMQKGQVLLKKKEEKLSQLESSLLEELSDEDTLKGAACKKVVTFDLSDSEETHSLSSTDVPQHKFDLKPDVQFPQLDKIQHLTSTLQHITAELNGVLSTLGSLNQRQPPLFAPVQVPMTALPSDGVPLSMYTSLARAQAAGPIMPPAGIPLVNQWAWSTGLNSSPSSTAGQSVDDILAEKWHKYFPGGFPSLSGSPAPLDSNLGYLPAGKVGALGTTAFTRGKVTWFLIQAASACVTRPSGLPSPSRLTTGNISSTSRRRNRYGRDPFLMCRCVGFPCEQEAFRSLLWWLPGSTLACSLLSSAVWVQRFRDMSMFVRVSVLALSYKPIPTYHSPSRGSEQGTGRASV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GK65</accession>
    <name>A0A452GK65_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="6">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
            <location evidence="1">Nucleolus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the eukaryotic RPA49/POLR1E RNA polymerase subunit family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GK65"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002403.1">
        <property type="protein sequence ID" value="ENSGAGP00000002113.1"/>
        <property type="gene ID" value="ENSGAGG00000001659.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000428">
        <property type="term" value="C:DNA-directed RNA polymerase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005730">
        <property type="term" value="C:nucleolus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006351">
        <property type="term" value="P:DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009668">
        <property type="entry name" value="RNA_pol-assoc_fac_A49-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14440">
        <property type="entry name" value="DNA-DIRECTED RNA POLYMERASE I SUBUNIT RPA49"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06870">
        <property type="entry name" value="RNA_pol_I_A49"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0240">DNA-directed RNA polymerase</keyword>
    <keyword evidence="5" id="KW-0539">Nucleus</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0804">Transcription</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004604"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009430"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022478"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002113.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="358" mass="40274" checksum="54BDC334EB3DE6C1" modified="2019-05-08" version="1">MLSLCKFIPVQFSNGKLRNSENMDFTLYKSKDTTNPRKKHQRILTTETDRLSYVGKNFGSGILKCNSLCRYFVGVLDKDAGQMEVYNAELFNMQPLLSAFGTNKQKRALNSRRMHQVGSETLNVAVTKAAENIIETKGVAASTERAKCSLLFLKTSSVLSPAEYEALQTPAAAFVSITPEEILKKTEEKSHCSFVLEELKSMPPSEDRADHKARCLWFLDALIKFSFQKVIKKKYALGPDCPHVINSKLMKNFTALTYNNGSIQNLISASMKAKITAYVIALALHINNFQTDLTVLQRDMKLREKRILEIAKAMRLKISKSKGPSGLTDNEDHKLATLSLPLPVYKPAGRQRKRKKMT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GK66</accession>
    <name>A0A452GK66_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Focal adhesion kinase 1</fullName>
            <ecNumber evidence="4">2.7.10.2</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="19">Protein-tyrosine kinase 2</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="20">pp125FAK</fullName>
        </alternativeName>
    </protein>
    <organism evidence="26 27">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="27" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="26" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="26" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="21 28 29">
            <text>L-tyrosyl-[protein] + ATP = O-phospho-L-tyrosyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10596"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.10.2"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell junction</location>
            <location evidence="2">Focal adhesion</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cell membrane</location>
            <topology evidence="3">Peripheral membrane protein</topology>
            <orientation evidence="3">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Cilium basal body</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="2.7.10.2" evidence="4"/>
    <dbReference type="AlphaFoldDB" id="A0A452GK66"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002261.1">
        <property type="protein sequence ID" value="ENSGAGP00000001986.1"/>
        <property type="gene ID" value="ENSGAGG00000001572.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005925">
        <property type="term" value="C:focal adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004715">
        <property type="term" value="F:non-membrane spanning protein tyrosine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008284">
        <property type="term" value="P:positive regulation of cell population proliferation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007172">
        <property type="term" value="P:signal complex assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd14473">
        <property type="entry name" value="FERM_B-lobe"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd13190">
        <property type="entry name" value="FERM_C_FAK1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05056">
        <property type="entry name" value="PTKc_FAK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.330:FF:000001">
        <property type="entry name" value="focal adhesion kinase 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000058">
        <property type="entry name" value="focal adhesion kinase 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.20.90:FF:000021">
        <property type="entry name" value="focal adhesion kinase 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000047">
        <property type="entry name" value="focal adhesion kinase 1 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000039">
        <property type="entry name" value="Focal adhesion kinase, isoform D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.80.10:FF:000004">
        <property type="entry name" value="Protein-tyrosine kinase 2-beta isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.80.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.330">
        <property type="entry name" value="Nucleotidyltransferases domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.20.90">
        <property type="entry name" value="Phosphatidylinositol 3-kinase Catalytic Subunit, Chain A, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.540">
        <property type="entry name" value="Single helix bin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019749">
        <property type="entry name" value="Band_41_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041390">
        <property type="entry name" value="FADK_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049385">
        <property type="entry name" value="FAK1-like_FERM_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041784">
        <property type="entry name" value="FAK1/PYK2_FERM_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014352">
        <property type="entry name" value="FERM/acyl-CoA-bd_prot_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035963">
        <property type="entry name" value="FERM_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019748">
        <property type="entry name" value="FERM_central"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000299">
        <property type="entry name" value="FERM_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036137">
        <property type="entry name" value="Focal_adhe_kin_target_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005189">
        <property type="entry name" value="Focal_adhesion_kin_target_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001245">
        <property type="entry name" value="Ser-Thr/Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008266">
        <property type="entry name" value="Tyr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020635">
        <property type="entry name" value="Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029071">
        <property type="entry name" value="Ubiquitin-like_domsf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46221">
        <property type="entry name" value="FERM AND PDZ DOMAIN-CONTAINING PROTEIN FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46221:SF12">
        <property type="entry name" value="NON-SPECIFIC PROTEIN-TYROSINE KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21477">
        <property type="entry name" value="FERM_C_FAK1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00373">
        <property type="entry name" value="FERM_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18038">
        <property type="entry name" value="FERM_N_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03623">
        <property type="entry name" value="Focal_AT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07714">
        <property type="entry name" value="PK_Tyr_Ser-Thr"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00109">
        <property type="entry name" value="TYRKINASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00295">
        <property type="entry name" value="B41"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00219">
        <property type="entry name" value="TyrKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF68993">
        <property type="entry name" value="FAT domain of focal adhesion kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47031">
        <property type="entry name" value="Second domain of FERM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54236">
        <property type="entry name" value="Ubiquitin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50057">
        <property type="entry name" value="FERM_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00109">
        <property type="entry name" value="PROTEIN_KINASE_TYR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="12 22" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="13" id="KW-0965">Cell junction</keyword>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="17" id="KW-0966">Cell projection</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="16" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="5" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="11" id="KW-0418">Kinase</keyword>
    <keyword evidence="14" id="KW-0472">Membrane</keyword>
    <keyword evidence="10 22" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="8" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="27" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0808">Transferase</keyword>
    <keyword evidence="15" id="KW-0829">Tyrosine-protein kinase</keyword>
    <feature type="domain" description="FERM" evidence="25">
        <location>
            <begin position="35"/>
            <end position="355"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="24">
        <location>
            <begin position="463"/>
            <end position="721"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="23">
        <location>
            <begin position="1"/>
            <end position="27"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="23">
        <location>
            <begin position="727"/>
            <end position="785"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="23">
        <location>
            <begin position="840"/>
            <end position="896"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="23">
        <location>
            <begin position="10"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="23">
        <location>
            <begin position="727"/>
            <end position="738"/>
        </location>
    </feature>
    <feature type="binding site" evidence="22">
        <location>
            <position position="495"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004120"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004413"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00011903"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00022553"/>
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    </evidence>
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            <dbReference type="ARBA" id="ARBA00022679"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
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            <dbReference type="ARBA" id="ARBA00039644"/>
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    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00042078"/>
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            <dbReference type="ARBA" id="ARBA00051245"/>
        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS50057"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001986.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1049" mass="118443" checksum="188BA6F6AE502FDA" modified="2019-05-08" version="1">MAAAYLDPNLNHTPSSSAKTHLSTGVERSPGAMERVLKVFHYFESSSEPTTWASIIRHGDATDVRGIIQKIVDSHKVKNVACYGLRLSHLQSEEVHWLHLDMGVSNVRERFELAHPPEEWKYELRIRYLPKGFLNQFTEDKPTLNFFYQQVKNDYMLEIADQVDQEIALKLGCLEIRRSYWEMRGNALEKKSNYEVLEKDVGLKRFFPKSLLDSVKAKTLRKLIQQTFRQFAVLNREESIQKFFEILSPVYRFDKECFKCALGSSWIISVELAIGPEEGISYLTDKGSNPTHLADFNQVQTIQYSNSEDKDRKGMLQLKIAGAPEPLTVTAPSLTIAENMADLIDGYCRLVNGATQSFIIRPQKEGERALPSIPKLANNEKQGVRSHTVSVSGVSHCQHKVKKARRFLPFVFCSHEPPPKDEISGDETDDYAEIIDEEDTYTMPSKSYGIDEARDYEIQRERIELGRCIGEGQFGDVHQGVYMSPENPAMAVAIKTCKNCTSDSVREKFLQEALTMRQFDHPHIVKLIGVITENPVWIIMELCTLGELRSFLQVRKYSLDLASLILYAYQLSTALAYLESKRFVHRDIAARNVLVSSTDCVKLGDFGLSRYMEDSTYYKASKGKLPIKWMAPESINFRRFTSASDVWMFGVCMWEILMHGVKPFQGVKNNDVIGRIENGERLPMPPNCPPTLYSLMTKCWAYDPSRRPRFTELKAQLSTILEEEKLQQEERMRMESRRQVTVSWDSGGSDEAPPKPSRPGYPSPRSSEGFYPSPQHMVPPNHYQVSGYPGSHGIPAMAGSIYPGQASLLDQADSWNHRPQEISMWPPTMEKPDVRLSRGSIDREDGSLQGPAGNQHIYQPVGKPDHAAPPKKPPRPGAPSHLGSLASLNSPVDSYNEGVKLQPQEISPPPTANLDRSNDKVYENVTGLVKAVIEMSSKIQPAPPEEYVPMVKEVGLALRTLLATVDETIPVLPASTHREIEMAQKLLNSDLAELINKMKLAQQYVMTSLQQEYKKQMLTAAHALAVDAKNLLDVIDQARLKMIGQSRPH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GK77</accession>
    <name>A0A452GK77_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3 7">Carbonic anhydrase</fullName>
            <ecNumber evidence="3 7">4.2.1.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="7">Reversible hydration of carbon dioxide.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="7">
            <text>hydrogencarbonate + H(+) = CO2 + H2O</text>
            <dbReference type="Rhea" id="RHEA:10748"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:17544"/>
            <dbReference type="EC" id="4.2.1.1"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 7">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <comment type="similarity">
        <text evidence="2 7">Belongs to the alpha-carbonic anhydrase family.</text>
    </comment>
    <dbReference type="EC" id="4.2.1.1" evidence="3 7"/>
    <dbReference type="AlphaFoldDB" id="A0A452GK77"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002388.1">
        <property type="protein sequence ID" value="ENSGAGP00000002100.1"/>
        <property type="gene ID" value="ENSGAGG00000001445.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004089">
        <property type="term" value="F:carbonate dehydratase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.200.10:FF:000001">
        <property type="entry name" value="Carbonic anhydrase 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.200.10">
        <property type="entry name" value="Alpha carbonic anhydrase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001148">
        <property type="entry name" value="CA_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036398">
        <property type="entry name" value="CA_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023561">
        <property type="entry name" value="Carbonic_anhydrase_a-class"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018338">
        <property type="entry name" value="Carbonic_anhydrase_a-class_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18952">
        <property type="entry name" value="CARBONIC ANHYDRASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18952:SF25">
        <property type="entry name" value="CARBONIC ANHYDRASE 5B, MITOCHONDRIAL-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00194">
        <property type="entry name" value="Carb_anhydrase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01057">
        <property type="entry name" value="Carb_anhydrase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51069">
        <property type="entry name" value="Carbonic anhydrase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00162">
        <property type="entry name" value="ALPHA_CA_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51144">
        <property type="entry name" value="ALPHA_CA_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6 7" id="KW-0456">Lyase</keyword>
    <keyword evidence="4 7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 7" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="Alpha-carbonic anhydrase" evidence="8">
        <location>
            <begin position="7"/>
            <end position="261"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010718"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012925"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU367011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51144"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002100.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="282" mass="32418" checksum="EA3605EAAE5C16E6" modified="2019-05-08" version="1">MATGLVTQDPYRHKMWGMSELSVSGGTRQSPINIQWRDSVYDPHLKPLKIHYDPTTCLYIWNNGYSFQVEFDDSADKSVIMGGPLENHYRLKQFHFHWGAINEWGSEHMVDSKVYPAELHLVHWNPVTSTSFEEAIMENNGLVVIGVFLKLGAHHAGLQKLVDALPAVKHKDTIVEFDVFDPSCLMPSCPDYWTYDGSLTTPPLTESVTWIIKKTPIEVDQNQLEEFRMLLFTSAGEEEKRMVDNFRPLQPLMNRTVRSSFRNHHLLNIETRSSELAHQHSP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GK79</accession>
    <name>A0A452GK79_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Carbohydrate-binding/sugar hydrolysis domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="pathway">
        <text evidence="1">Protein modification; protein ubiquitination.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GK79"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002426.1">
        <property type="protein sequence ID" value="ENSGAGP00000002130.1"/>
        <property type="gene ID" value="ENSGAGG00000001710.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042981">
        <property type="term" value="P:regulation of apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006511">
        <property type="term" value="P:ubiquitin-dependent protein catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.160.20.10:FF:000017">
        <property type="entry name" value="F-box only protein 10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.160.20.10:FF:000022">
        <property type="entry name" value="F-box only protein 10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.160.20.10">
        <property type="entry name" value="Single-stranded right-handed beta-helix, Pectin lyase-like"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039448">
        <property type="entry name" value="Beta_helix"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006633">
        <property type="entry name" value="Carb-bd_sugar_hydrolysis-dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022441">
        <property type="entry name" value="Para_beta_helix_rpt-2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006626">
        <property type="entry name" value="PbH1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012334">
        <property type="entry name" value="Pectin_lyas_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011050">
        <property type="entry name" value="Pectin_lyase_fold/virulence"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051550">
        <property type="entry name" value="SCF-Subunits/Alg-Epimerases"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR03804">
        <property type="entry name" value="para_beta_helix"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22990">
        <property type="entry name" value="F-BOX ONLY PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22990:SF15">
        <property type="entry name" value="F-BOX ONLY PROTEIN 10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13229">
        <property type="entry name" value="Beta_helix"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00722">
        <property type="entry name" value="CASH"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00710">
        <property type="entry name" value="PbH1"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51126">
        <property type="entry name" value="Pectin lyase-like"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="Carbohydrate-binding/sugar hydrolysis" evidence="5">
        <location>
            <begin position="36"/>
            <end position="167"/>
        </location>
    </feature>
    <feature type="domain" description="Carbohydrate-binding/sugar hydrolysis" evidence="5">
        <location>
            <begin position="272"/>
            <end position="412"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="446"/>
            <end position="467"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="457"/>
            <end position="467"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="SMART" id="SM00722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002130.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="495" mass="53080" checksum="460D9091C655F554" modified="2019-05-08" version="1">FPGWCKGSISIIMLKNDIHHCKASGIFLRLAAGGLIADNNIHSNCEAGVDIRKGANPLILCNKIHSGLRSGIVVLGNGKGIIRSNQIYGNKEAGIYILYNGNPIVSGNHIFQGLAAGIAVNENGRGQITENVIRENQWGGADIRRGGDPVLRSNLICCGYSDGVVVGERGKGLIEGNTIYGNKGCGVWIMSSSLPHITNNRIGHNSIYGVAVFCRKDDANDYLANQGGNENFNDEGEAASWENDLDSEDECLTSRRPISVALVESNSINHNGAAGLYVKSSEALNIIANAIHANHDSGVAVLQSSQLTRIANNSISCNSLGGVLIESEARVELRGNGIYDNSSHGITSKGDGVIVENDIVGNRGCGLQLLQTADMKVTKNRIQSFWDYGIVMLDQTKGLVQENLIFQGRSKKTILQQVSNAEDCIVQNNKLLTFKSKSDIAWTLENPPARPHIEGSTRGTSMATNSQKVTTMSTRIAARMDGGCRNNGSIFCTIL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GK81</accession>
    <name>A0A452GK81_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3 7">Carbonic anhydrase</fullName>
            <ecNumber evidence="3 7">4.2.1.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="7">Reversible hydration of carbon dioxide.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="7">
            <text>hydrogencarbonate + H(+) = CO2 + H2O</text>
            <dbReference type="Rhea" id="RHEA:10748"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:17544"/>
            <dbReference type="EC" id="4.2.1.1"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 7">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <comment type="similarity">
        <text evidence="2 7">Belongs to the alpha-carbonic anhydrase family.</text>
    </comment>
    <dbReference type="EC" id="4.2.1.1" evidence="3 7"/>
    <dbReference type="AlphaFoldDB" id="A0A452GK81"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002394.1">
        <property type="protein sequence ID" value="ENSGAGP00000002105.1"/>
        <property type="gene ID" value="ENSGAGG00000001445.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004089">
        <property type="term" value="F:carbonate dehydratase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.200.10:FF:000001">
        <property type="entry name" value="Carbonic anhydrase 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.200.10">
        <property type="entry name" value="Alpha carbonic anhydrase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001148">
        <property type="entry name" value="CA_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036398">
        <property type="entry name" value="CA_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023561">
        <property type="entry name" value="Carbonic_anhydrase_a-class"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018338">
        <property type="entry name" value="Carbonic_anhydrase_a-class_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18952">
        <property type="entry name" value="CARBONIC ANHYDRASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18952:SF25">
        <property type="entry name" value="CARBONIC ANHYDRASE 5B, MITOCHONDRIAL-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00194">
        <property type="entry name" value="Carb_anhydrase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01057">
        <property type="entry name" value="Carb_anhydrase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51069">
        <property type="entry name" value="Carbonic anhydrase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00162">
        <property type="entry name" value="ALPHA_CA_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51144">
        <property type="entry name" value="ALPHA_CA_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6 7" id="KW-0456">Lyase</keyword>
    <keyword evidence="4 7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 7" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="Alpha-carbonic anhydrase" evidence="8">
        <location>
            <begin position="14"/>
            <end position="269"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010718"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012925"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU367011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51144"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002105.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="290" mass="33194" checksum="1225D42B73079097" modified="2019-05-08" version="1">TCSLTPVTAGLQRAPCRYADLHPLWQSELSVSGGTRQSPINIQWRDSVYDPHLKPLKIHYDPTTCLYIWNNGYSFQVEFDDSADKSVIMGGPLENHYRLKQFHFHWGAINEWGSEHMVDSKVYPAELHLVHWNPVTSTSFEEAIMENNGLVVIGVFLKLGAHHAGLQKLVDALPAVKHKDTIVEFDVFDPSCLMPSCPDYWTYDGSLTTPPLTESVTWIIKKTPIEVDQNQLEEFRMLLFTSAGEEEKRMVDNFRPLQPLMNRTVRSSFRNHHLLNIETRSSELAHQHSP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GK82</accession>
    <name>A0A452GK82_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3 7">Carbonic anhydrase</fullName>
            <ecNumber evidence="3 7">4.2.1.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="7">Reversible hydration of carbon dioxide.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="7">
            <text>hydrogencarbonate + H(+) = CO2 + H2O</text>
            <dbReference type="Rhea" id="RHEA:10748"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:17544"/>
            <dbReference type="EC" id="4.2.1.1"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 7">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <comment type="similarity">
        <text evidence="2 7">Belongs to the alpha-carbonic anhydrase family.</text>
    </comment>
    <dbReference type="EC" id="4.2.1.1" evidence="3 7"/>
    <dbReference type="AlphaFoldDB" id="A0A452GK82"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002424.1">
        <property type="protein sequence ID" value="ENSGAGP00000002129.1"/>
        <property type="gene ID" value="ENSGAGG00000001445.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004089">
        <property type="term" value="F:carbonate dehydratase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.200.10:FF:000001">
        <property type="entry name" value="Carbonic anhydrase 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.200.10">
        <property type="entry name" value="Alpha carbonic anhydrase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001148">
        <property type="entry name" value="CA_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036398">
        <property type="entry name" value="CA_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023561">
        <property type="entry name" value="Carbonic_anhydrase_a-class"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018338">
        <property type="entry name" value="Carbonic_anhydrase_a-class_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18952">
        <property type="entry name" value="CARBONIC ANHYDRASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18952:SF25">
        <property type="entry name" value="CARBONIC ANHYDRASE 5B, MITOCHONDRIAL-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00194">
        <property type="entry name" value="Carb_anhydrase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01057">
        <property type="entry name" value="Carb_anhydrase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51069">
        <property type="entry name" value="Carbonic anhydrase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00162">
        <property type="entry name" value="ALPHA_CA_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51144">
        <property type="entry name" value="ALPHA_CA_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6 7" id="KW-0456">Lyase</keyword>
    <keyword evidence="4 7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 7" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="Alpha-carbonic anhydrase" evidence="8">
        <location>
            <begin position="1"/>
            <end position="247"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010718"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012925"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU367011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51144"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002129.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="268" mass="30798" checksum="D30E162825F9EFC7" modified="2019-05-08" version="1">KKNNSELSVSGGTRQSPINIQWRDSVYDPHLKPLKIHYDPTTCLYIWNNGYSFQVEFDDSADKSVIMGGPLENHYRLKQFHFHWGAINEWGSEHMVDSKVYPAELHLVHWNPVTSTSFEEAIMENNGLVVIGVFLKLGAHHAGLQKLVDALPAVKHKDTIVEFDVFDPSCLMPSCPDYWTYDGSLTTPPLTESVTWIIKKTPIEVDQNQLEEFRMLLFTSAGEEEKRMVDNFRPLQPLMNRTVRSSFRNHHLLNIETRSSELAHQHSP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GK83</accession>
    <name>A0A452GK83_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="20">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="15">Homodimer; mediates homophilic interactions to promote cell adhesion.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="14">Synapse</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000002298.1">
        <property type="protein sequence ID" value="ENSGAGP00000002020.1"/>
        <property type="gene ID" value="ENSGAGG00000001306.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030424">
        <property type="term" value="C:axon"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045202">
        <property type="term" value="C:synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098632">
        <property type="term" value="F:cell-cell adhesion mediator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007411">
        <property type="term" value="P:axon guidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060219">
        <property type="term" value="P:camera-type eye photoreceptor cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070593">
        <property type="term" value="P:dendrite self-avoidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007156">
        <property type="term" value="P:homophilic cell adhesion via plasma membrane adhesion molecules"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010842">
        <property type="term" value="P:retina layer formation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007416">
        <property type="term" value="P:synapse assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="CDD" id="cd00096">
        <property type="entry name" value="Ig"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="CDD" id="cd05734">
        <property type="entry name" value="Ig_DSCAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05735">
        <property type="entry name" value="Ig_DSCAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd20958">
        <property type="entry name" value="IgI_5_Dscam"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000333">
        <property type="entry name" value="Down syndrome cell adhesion molecule"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000729">
        <property type="entry name" value="Down syndrome cell adhesion molecule"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:001868">
        <property type="entry name" value="Down syndrome cell adhesion molecule"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000141">
        <property type="entry name" value="Down syndrome cell adhesion molecule a"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000176">
        <property type="entry name" value="Down syndrome cell adhesion molecule a"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000215">
        <property type="entry name" value="Down syndrome cell adhesion molecule a"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000017">
        <property type="entry name" value="Down syndrome cell adhesion molecule b"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000104">
        <property type="entry name" value="Down syndrome cell adhesion molecule b"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000167">
        <property type="entry name" value="Down syndrome cell adhesion molecule b"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000172">
        <property type="entry name" value="Down syndrome cell adhesion molecule b"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000219">
        <property type="entry name" value="Down syndrome cell adhesion molecule homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000229">
        <property type="entry name" value="Down syndrome cell adhesion molecule homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000120">
        <property type="entry name" value="Down syndrome cell adhesion molecule like 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000477">
        <property type="entry name" value="DS cell adhesion molecule like 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056754">
        <property type="entry name" value="DSCAM/DSCAML_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013098">
        <property type="entry name" value="Ig_I-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013106">
        <property type="entry name" value="Ig_V-set"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10075">
        <property type="entry name" value="BASIGIN RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10075:SF72">
        <property type="entry name" value="CELL ADHESION MOLECULE DSCAML1"/>
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        <property type="entry name" value="fn3"/>
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        <property type="entry name" value="I-set"/>
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    </dbReference>
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        <property type="entry name" value="IG"/>
        <property type="match status" value="9"/>
    </dbReference>
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        <property type="entry name" value="IGc2"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SMART" id="SM00406">
        <property type="entry name" value="IGv"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="8"/>
    </dbReference>
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        <property type="match status" value="6"/>
    </dbReference>
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        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="8"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="10 17" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="9" id="KW-0770">Synapse</keyword>
    <keyword evidence="3 17" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 17" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="1453"/>
            <end position="1473"/>
        </location>
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    <feature type="domain" description="Ig-like" evidence="18">
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            <begin position="87"/>
            <end position="171"/>
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    <feature type="domain" description="Ig-like" evidence="18">
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            <end position="260"/>
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    <feature type="domain" description="Ig-like" evidence="18">
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            <end position="362"/>
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    <feature type="domain" description="Ig-like" evidence="18">
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            <end position="447"/>
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    <feature type="domain" description="Ig-like" evidence="18">
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    <feature type="domain" description="Ig-like" evidence="18">
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            <begin position="551"/>
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        <location>
            <begin position="649"/>
            <end position="746"/>
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        <location>
            <begin position="748"/>
            <end position="845"/>
        </location>
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    <feature type="domain" description="Fibronectin type-III" evidence="19">
        <location>
            <begin position="850"/>
            <end position="949"/>
        </location>
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    <feature type="domain" description="Fibronectin type-III" evidence="19">
        <location>
            <begin position="954"/>
            <end position="1050"/>
        </location>
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    <feature type="domain" description="Fibronectin type-III" evidence="19">
        <location>
            <begin position="1054"/>
            <end position="1149"/>
        </location>
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        <location>
            <begin position="1139"/>
            <end position="1228"/>
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        <location>
            <begin position="1244"/>
            <end position="1338"/>
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    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="19">
        <location>
            <begin position="1339"/>
            <end position="1439"/>
        </location>
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        <location>
            <begin position="1544"/>
            <end position="1567"/>
        </location>
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    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1609"/>
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            <begin position="1850"/>
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        <location>
            <begin position="1802"/>
            <end position="1826"/>
        </location>
    </feature>
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        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
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    </evidence>
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    </evidence>
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            <dbReference type="ARBA" id="ARBA00022889"/>
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    </evidence>
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            <dbReference type="ARBA" id="ARBA00022902"/>
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            <dbReference type="ARBA" id="ARBA00023136"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
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        <source>
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        <source>
            <dbReference type="ARBA" id="ARBA00065695"/>
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    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
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    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002020.1"/>
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    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1869" mass="205254" checksum="15CD7F32F3990296" modified="2019-05-08" version="1">TIPMECPCPGTGFDQTKSNNQTFSSLLLVSPENRFFITFYGGLYISDVQKEDALSTYRCITKHKYSGETRQSNGARLSVSDPAESIPTILDGFQSRELKAGQLVELPCIASGYPNPAIRWIKDGRPLPADSRWTKRITGLTISDLRGEDSGTYICEVTNTFGSAEVTGTLTVIDPLHVSLTPKKLKTGIGSTVILSCALSGSPEYALRWYRNTDLVVLDNYISIRGISNETLLITAAQKSHSGAYQCFATRKAQTAQDFSIIALEDGTPRIVSSFSEKVVNPGEQFSLMCAAKGAPPPTVTWALDDEPIQRDNGHRTNQYTMSDGTTVSHMNVTSPQIKDGGVYRCTARNSVGSAEYQARINVRGPPSIRAMKNITAVAGRDTFINCRVIGYPYYSIKWYKDALLLPDNHRQVVFENGTLKLMDVQKGMDEGEYLCSVLIQPQLSISQSVHVTVKVPPLIQPFEFPPASIGQLLYIPCVVSSGDMPIRITWRKDGQVIISGSGVTIESKEFMSSLQISSVSLKHNGNYTCIASNAAATVSRERQLIVRVPPRFVVQPNNQDGIYGKAGVLNCSVDGYPPPKVMWKHAKGSGNPQQYHPVPLTGRIQILPNSSLLIRHVLEEDIGYYLCQASNGVGTDISKSMFLTVKIPAMITSHPNTTIAIKGQMKELNCTARGERPIIIRWEKGDTVIDPDRNMRYAISTKDNGDEVISTLKLKPADRGDSVFFSCHAINSYGEDRGLIQLTVQEPPDPPELEIREVKARSMNLRWTQRFDGNSIITGFDIEYKNKSDSWDFKQSTRNISPTINQANIVDLHPASVYSIRMYSFNKIGRSEPSKELTISTEEAAPDGPPMDVTLQPITSQSIQVTWKAPKKELQNGVIRGYQIGYRENSPGSNGQYSIVEMKATGDSEVYTLDNLKKFAQYGVVVQAFNRAGTGPSSSEINATTLEDVPSQPPENVRALSITSDVAVISWSEPPRSTLNGVLKGYRVIFWSLYMDGEWGEMQNITTTRERVELRGMEKFTNYSVQVLAYTQAGDGVRSSVLYIQTKEDIPGPPARIKAVPSSASSVVVSWLPPTKPNGIIRKYTIFCSSPGSGQPAPSEYETSPEQLFYRIAHLNRGQQYMLWVAAVTSAGRGNISEKVTIEPAGKAPAKIISFGGTVTTPWMKDVRLPCNSVGEPAPAIKWTKDSEDSAIPVAVDGHRLIQANGTLVLRSVKAEDSGYYTCTATNTWGFDTIIINLLVQVPPDQPRLTVSKTSASSITLAWIPGDNGGSSIRGFVLQYSVDNSEEWKDVFISSSERAFQLDSLKCGTWYKVKLAAKNSVGAGRISEIIEAKTHGREPSFSKDQHLFTHINSTHARLNLQGWSSGGCPIMAIVLEYRPKGNWVWQSLKTNNSNEVFLTELREATWYELRMKACNSAGCGNETTQFATLDYDGSTIPPIKSAQGEGDDVKKLFTIACPVILATLGVALLFIIRKKRKEKRLKRLRDAKSLAEMLISKNNRSFDTPVKGPPQGPRLHIDIPRVQLLIEDKEGIKQLGEWANPPSANPFLLKTPDPVSRKSVKSAHSTRNRYSSQWTLTKCQASTPARTLTSDWRTVGSQHGIMVTESDSYSASLSQDTDKGRNSMVSTESASSTYEELARAYEHAKLEEQLQHAKFEITECFISDSSSDQMTTGTNENADSMTSMSTPSEPGICRFTASPPKPQDSDRGKNVAVPIPHRANKSDYCNLPLYMKSEAFFRKPDIHDPCPVVPPREASIRNLARAYHTQARHMTLDPGSKPLGLPHPASSTTTLPQRTLPMPVSANSASATTASASAPATDPTSATTAEPRMPPHTKMGGSRDSLLEMSTSGIGRSQKQGAGAYSKSYTLV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GK87</accession>
    <name>A0A452GK87_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Centrosomal protein of 164 kDa</fullName>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Plays a role in microtubule organization and/or maintenance for the formation of primary cilia (PC), a microtubule-based structure that protrudes from the surface of epithelial cells. Plays a critical role in G2/M checkpoint and nuclear divisions. A key player in the DNA damage-activated ATR/ATM signaling cascade since it is required for the proper phosphorylation of H2AX, RPA, CHEK2 and CHEK1. Plays a critical role in chromosome segregation, acting as a mediator required for the maintenance of genomic stability through modulation of MDC1, RPA and CHEK1.</text>
    </comment>
    <comment type="subunit">
        <text evidence="15">Interacts (via N-terminus) with ATRIP. Interacts with ATM, ATR and MDC1. Interacts with XPA (via N-terminus) upon UV irradiation. Interacts with CEP83, CCDC92, TTBK2, DVL3, NPHP3 and weakly with NPHP4. Interacts with DZIP1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Microtubule organizing center</location>
            <location evidence="1">Centrosome</location>
            <location evidence="1">Centriole</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000002401.1">
        <property type="protein sequence ID" value="ENSGAGP00000002111.1"/>
        <property type="gene ID" value="ENSGAGG00000001670.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005814">
        <property type="term" value="C:centriole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097539">
        <property type="term" value="C:ciliary transition fiber"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060271">
        <property type="term" value="P:cilium assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006281">
        <property type="term" value="P:DNA repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00201">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Centrosomal protein of 164 kDa"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1470.10">
        <property type="entry name" value="Photosystem I PsaD, reaction center subunit II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051841">
        <property type="entry name" value="MT-Golgi_org_protein"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001202">
        <property type="entry name" value="WW_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036020">
        <property type="entry name" value="WW_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18902:SF27">
        <property type="entry name" value="CENTROSOMAL PROTEIN OF 164 KDA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18902">
        <property type="entry name" value="NUCLEAR MITOTIC APPARATUS PROTEIN 1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00397">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00456">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51045">
        <property type="entry name" value="WW domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01159">
        <property type="entry name" value="WW_DOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50020">
        <property type="entry name" value="WW_DOMAIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="13" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="5" id="KW-0132">Cell division</keyword>
    <keyword evidence="8" id="KW-0970">Cilium biogenesis/degradation</keyword>
    <keyword evidence="9 17" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="6" id="KW-0227">DNA damage</keyword>
    <keyword evidence="10" id="KW-0234">DNA repair</keyword>
    <keyword evidence="7" id="KW-0498">Mitosis</keyword>
    <keyword evidence="12" id="KW-0539">Nucleus</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="WW" evidence="19">
        <location>
            <begin position="58"/>
            <end position="91"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="107"/>
            <end position="147"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="312"/>
            <end position="366"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="440"/>
            <end position="460"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="509"/>
            <end position="545"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="562"/>
            <end position="593"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1150"/>
            <end position="1170"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="17">
        <location>
            <begin position="607"/>
            <end position="1095"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="18">
        <location>
            <begin position="110"/>
            <end position="126"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="512"/>
            <end position="521"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="528"/>
            <end position="542"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="570"/>
            <end position="585"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="1155"/>
            <end position="1170"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004114"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022776"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022794"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00056906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00061715"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00067900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50020"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002111.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1459" mass="165482" checksum="E7194F8731672FC6" modified="2019-05-08" version="1">MDMAGPAIRIGDQLILEEDYDETYIPSEQEIQEFAREIGIDPGNEPELMWLAREGIVAPLPPEWKPCQDITGDIYYFNFSNGQSTWDHPCDEHYRELVIQEREKLSAHGSIKKKDKKKKKDKKEKKEKKEREPVKHAASPGSLLAPIQVPPGSLAPLRGVMDGPASAFHGLLSTNVEISGVPSPLLREGTLPLQTLKSTGCTKSLLGSMHEEKASLNLLALDEEEEEESENQSPRGTARLLKNLHMDISALGGGFEYEFRGVAASVWSSAHQGSCRAEEDNCGTRLASVTEPGLPHLRNSEDEAGGLKEKESLKMSHAEESQTFPLDSDGVRPPAPDKLLNRDADSSLCGLNKEESSGKPLGNELLEKEDTDVEDTVSVAYEQSGTLGERTAEGTHVVSSQGHPVAASPDVRAAETDQMESVAVTVDAVSVSEKIVNGMREEAADLETDSKLDAGKESEASEHVKELQVSDCSDHELLHLTDFGFQSRISEQVLDMDILSAALDSPKVEAQGLREEEKDQSEASLEEEQSKRPKAAESERNQRPFSACKMAEEKCLSLENASQEETAALEQKEGSLDSPKEEEPAAPHQAQLGEEIKQVVSLNWPSAESLNEIARKQEQELEQEKERLLQAKEERVRRFQEELRQEEEEEAQVLHQQKEKSLWSLKEELAKASEEEELRMREEEVEKLSKLRSQISSETKTEKEKIRAEQEAALQKLREEWESLQKVERERLEERKKRVLEMLKVEVEEAQQREAAELEQEKQRVLNELKERLERKRKEAAEVLEKQFAAELQELKSTAEEKHQKVVSSLRKQIAEAQRSEEAQLREELERAEQKVQQKMYQVAEFERELSELMKEKRQEVERDHDRKMERMKEEHKEVLARIQEQYEEEERKQRTKKLEELKGELERLRQLHDGELKALQKQLDEQLSDLRQSHKVKERKLQELEMELEIQATDAKARCAQLNHQEEAMRKKRQQLLDEEKQTELQRNEAALAAQLRLEESRKEHADLVESIRQLRRTLEELQDQKAELESQVDLLQTRSQRLQRRISELEAAVQSKQETLKELVAEDSMASPRREAELHLEDLRETLQAHSSREPASLTSHSNEDSDLQLDNVRHFISAEGVSIKNAKEFLVRQTRSMRKRHTALKAAKQQWRHNMQQAQDAVQDPDSSQLLEDVRRNLEEESAQLDKMKSAMQKGQVLLKKKEEKLSQLESSLLEELSDEDTLKGAACKKVVTFDLSDSEETHSLSSTDVPQHKFDLKPDVQFPQLDKIQHLTSTLQHITAELNGVLSTLGSLNQRQPPLFAPVQVPMTALPSDGVPLSMYTSLARAQAAGPIMPPAGIPLVNQWAWSTGLNSSPSSTAGQSVDDILAEKWHKYFPGGGSPLWSPTMLATLMTALFFMVRALGQKGGSSRHDSIYSRKGNVVPDSSSKCLCDPTERAAVTIQTHYRKYQQHKQEKK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GK92</accession>
    <name>A0A452GK92_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="7">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000002451.1">
        <property type="protein sequence ID" value="ENSGAGP00000002148.1"/>
        <property type="gene ID" value="ENSGAGG00000001670.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005814">
        <property type="term" value="C:centriole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097539">
        <property type="term" value="C:ciliary transition fiber"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060271">
        <property type="term" value="P:cilium assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051841">
        <property type="entry name" value="MT-Golgi_org_protein"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18902:SF27">
        <property type="entry name" value="CENTROSOMAL PROTEIN OF 164 KDA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18902">
        <property type="entry name" value="NUCLEAR MITOTIC APPARATUS PROTEIN 1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4 5" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="24"/>
            <end position="75"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="98"/>
            <end position="124"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="176"/>
            <end position="233"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="295"/>
            <end position="328"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="366"/>
            <end position="398"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="433"/>
            <end position="465"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="491"/>
            <end position="522"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="757"/>
            <end position="810"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="885"/>
            <end position="904"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="954"/>
            <end position="990"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="1007"/>
            <end position="1038"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="1596"/>
            <end position="1615"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="5">
        <location>
            <begin position="1052"/>
            <end position="1540"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="30"/>
            <end position="55"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="102"/>
            <end position="118"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="176"/>
            <end position="185"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="198"/>
            <end position="214"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="302"/>
            <end position="311"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="441"/>
            <end position="461"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="494"/>
            <end position="519"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="957"/>
            <end position="966"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="973"/>
            <end position="987"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="1015"/>
            <end position="1030"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="1600"/>
            <end position="1615"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002148.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1920" mass="214984" checksum="3B090453A68822CE" modified="2019-05-08" version="1">MQPEPGILPSSSFYRMSSPILSSECASPDLDQHSQMRIEGFLKKGKRKPSERPPETSDSNKQFAGPSPNKLQPLFTAKSCRTHQILADVEKILGRMPSFSRSDVDHQQGQDRSPKIGDKPCLGFSDSESEGLDVVRTKPISYVARKSCSQDLENGRSVLDSCEVLEDKGMLFDEEKLSDNWKQEEMTGGSVCQPDDGLSVRERSTGPGGDKERGPGFSQADENEAFLTPSAPDHTFYSRKCEMILFDASPLEDLSLMEEGDVHCPQRELRAHSRAVESRRKRKLQDQALSQATKFCKSSEVVGKHERESSKQAEFVESEGAGLQAAPQKDLLDSHLAVPEVTADTLPYTRGVEAYAHAGADISASEPVSLSAAQHKDKEKANGLGPEPDNSLTSNSADHFASQILGEIDNFSWDLQSSRESDRPTDLLAAAAKGSLPTHPFTGTLQTPPRSSANGKSQSDCNSEDEKFYQHVQENDQVPSQEHSCVSFNMKLNRSRDAPEKNNEGRQEETGEGEQKTEPVPRGLEVDVFTEGLQVSPGWLYQTAECQESSRLCQQVEGELTQSGKVTEDYCKLKLCEGAGSNESPGSLLAPIQVPPGSLAPLRGVMDGPASAFHGLLSTNVEISGVPSPLLREGTLPLQTLKSTGCTKSLLGSMHEEKASLNLLALDEEEEEESENQSPRGTARLLKNLHMDISALGGGFEYEFRGVAASVWSSAHQGSCRAEEDNCGTRLASVTEPGLPHLRNSEDEAGGLKEKESLKMSHAEESQTFPLDSDGVRPPAPDKLLNRDADSSLCGLNKEESSGKPLGNELLEKEDTDVEDTVSVAYEQSGTLGERTAEGTHVVSSQGHPVAASPDVRAAETDQMESVAVTVDAVSVSEKIVNGMREEAADLETDSKLDAGKESEASEHVKELQVSDCSDHELLHLTDFGFQSRISEQVLDMDILSAALDSPKVEAQGLREEEKDQSEASLEEEQSKRPKAAESERNQRPFSACKMAEEKCLSLENASQEETAALEQKEGSLDSPKEEEPAAPHQAQLGEEIKQVVSLNWPSAESLNEIARKQEQELEQEKERLLQAKEERVRRFQEELRQEEEEEAQVLHQQKEKSLWSLKEELAKASEEEELRMREEEVEKLSKLRSQISSETKTEKEKIRAEQEAALQKLREEWESLQKVERERLEERKKRVLEMLKVEVEEAQQREAAELEQEKQRVLNELKERLERKRKEAAEVLEKQFAAELQELKSTAEEKHQKVVSSLRKQIAEAQRSEEAQLREELERAEQKVQQKMYQVAEFERELSELMKEKRQEVERDHDRKMERMKEEHKEVLARIQEQYEEEERKQRTKKLEELKGELERLRQLHDGELKALQKQLDEQLSDLRQSHKVKERKLQELEMELEIQATDAKARCAQLNHQEEAMRKKRQQLLDEEKQTELQRNEAALAAQLRLEESRKEHADLVESIRQLRRTLEELQDQKAELESQVDLLQTRSQRLQRRISELEAAVQSKQETLKELVAEDSMASPRREAELHLEDLRETLQAHSSREPASLTSHSNEDSDLQLDNVRHFISAEGVSIKNAKEFLVRQTRSMRKRHTALKAAKQQWRHNMQQAQDAVQDPDSSQLLEDVRRNLEEESAQLDKMKSAMQKGQVLLKKKEEKLSQLESSLLEELSDEDTLKGAACKKVVTFDLSDSEETHSLSSTDVPQHKFDLKPDVQFPQLDKIQHLTSTLQHITAELNGVLSTLGSLNQRQPPLFAPVQVPMTALPSDGVPLSMYTSLARAQAAGPIMPPAGIPLVNQWAWSTGLNSSPSSTAGQSVDDILAEKWHKYFPGGFPSLSGSPAPLDSNLGYLPAGEQIGLFQRSQLQGPETDSTSVQGMIDANKKWLENFKKDSKVPLFPRTQKPFASSPGLVQLGLDENNQIKVYHC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GK93</accession>
    <name>A0A452GK93_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">KRAB domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GK93"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002406.1">
        <property type="protein sequence ID" value="ENSGAGP00000002116.1"/>
        <property type="gene ID" value="ENSGAGG00000001694.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006355">
        <property type="term" value="P:regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd07765">
        <property type="entry name" value="KRAB_A-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001909">
        <property type="entry name" value="KRAB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036051">
        <property type="entry name" value="KRAB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050169">
        <property type="entry name" value="Krueppel_C2H2_ZnF"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23232">
        <property type="entry name" value="KRAB DOMAIN C2H2 ZINC FINGER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23232:SF133">
        <property type="entry name" value="RIKEN CDNA 1700020N01 GENE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01352">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00349">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109640">
        <property type="entry name" value="KRAB domain (Kruppel-associated box)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50805">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="KRAB" evidence="2">
        <location>
            <begin position="31"/>
            <end position="88"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="1"/>
            <end position="10"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="PROSITE" id="PS50805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002116.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="88" mass="9915" checksum="2E100FA6A164DE3F" modified="2019-05-08" version="1">MEKGGWERKGNTGKNQQGPTGRGLLLFQGPVTFEEVAVYFTREEWALLDPRQRALCRDVMQENYENVTLVKVHATPTMPPLGWGVVAL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GK95</accession>
    <name>A0A452GK95_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">C2H2-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GK95"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002320.1">
        <property type="protein sequence ID" value="ENSGAGP00000002042.1"/>
        <property type="gene ID" value="ENSGAGG00000001645.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000012">
        <property type="entry name" value="RB-associated KRAB zinc finger protein-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002343">
        <property type="entry name" value="Zinc finger protein 33A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001437">
        <property type="entry name" value="Zinc finger protein 594"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23226">
        <property type="entry name" value="ZINC FINGER AND SCAN DOMAIN-CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23226:SF366">
        <property type="entry name" value="ZINC FINGER PROTEIN ZFP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="13" id="KW-0732">Signal</keyword>
    <keyword evidence="9" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 11" id="KW-0863">Zinc-finger</keyword>
    <feature type="signal peptide" evidence="13">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019341140" description="C2H2-type domain-containing protein" evidence="13">
        <location>
            <begin position="21"/>
            <end position="268"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="14">
        <location>
            <begin position="109"/>
            <end position="136"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="14">
        <location>
            <begin position="137"/>
            <end position="164"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="14">
        <location>
            <begin position="165"/>
            <end position="192"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="40"/>
            <end position="75"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="221"/>
            <end position="268"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="51"/>
            <end position="75"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="247"/>
            <end position="260"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002042.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="268" mass="30801" checksum="98751BACA14A8274" modified="2019-05-08" version="1">MIKWAHWLFLFHPFLQKPQSLFPACSLELHISEAHGELLRRSKGNVSSPREQGKAGERYHRPERQQGNHSEERVDEPINCQRTYRDLKETTSQEKILRKKRANTCSEPYECCECGKSFNRMSVLIRHQRIHKGDKPYECCECGKSFTQRYALIAHQRIHTEERPYECCKCRKSFTQRSALISHQTTHTGERPYECCECGKKATPPHPLWACSNWRQSIKGSSSAQPGWTKEENKPMLLAPPRKLLDPEAAASSHAEAPADTQSTAKIG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMS3</accession>
    <name>A0A452HMS3_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="13">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Mitochondrion membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 10">Belongs to the mitochondrial carrier (TC 2.A.29) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMS3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018563.1">
        <property type="protein sequence ID" value="ENSGAGP00000016261.1"/>
        <property type="gene ID" value="ENSGAGG00000012073.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031966">
        <property type="term" value="C:mitochondrial membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015227">
        <property type="term" value="F:O-acyl-L-carnitine transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1902603">
        <property type="term" value="P:carnitine transmembrane transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006839">
        <property type="term" value="P:mitochondrial transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.50.40.10:FF:000051">
        <property type="entry name" value="Mitochondrial carnitine/acylcarnitine carrier protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.50.40.10">
        <property type="entry name" value="Mitochondrial carrier domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023395">
        <property type="entry name" value="MCP_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018108">
        <property type="entry name" value="MCP_transmembrane"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050567">
        <property type="entry name" value="Mitochondrial_Carrier"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45624">
        <property type="entry name" value="MITOCHONDRIAL BASIC AMINO ACIDS TRANSPORTER-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45624:SF56">
        <property type="entry name" value="MITOCHONDRIAL CARNITINE_ACYLCARNITINE CARRIER PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00153">
        <property type="entry name" value="Mito_carr"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103506">
        <property type="entry name" value="Mitochondrial carrier"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50920">
        <property type="entry name" value="SOLCAR"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 9 12" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="4 9 10" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 12" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="3 10" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="47"/>
            <end position="67"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="79"/>
            <end position="101"/>
        </location>
    </feature>
    <feature type="repeat" description="Solcar" evidence="9">
        <location>
            <begin position="1"/>
            <end position="71"/>
        </location>
    </feature>
    <feature type="repeat" description="Solcar" evidence="9">
        <location>
            <begin position="80"/>
            <end position="168"/>
        </location>
    </feature>
    <feature type="repeat" description="Solcar" evidence="9">
        <location>
            <begin position="179"/>
            <end position="265"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1"/>
            <end position="21"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004225"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006375"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00282"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="RuleBase" id="RU000488"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016261.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="273" mass="29871" checksum="3A673D562E3A211B" modified="2019-05-08" version="1">TAPGLHRVRLQTQPKPQAGQPPLYSGTFDCFRQTLVREGVRGLYKGMAAPIVGVTPMFAVCFFGFGLGKKLQQKNPDDILTYPQLFAAGMLSGVFTTAIMAPGERIKCLLQIQAASGEIKYGGPMDCAKQLYREAGIRGIYKGTLLTLMRDVPASGMYFMTYEWLKNILTPEGKSVSDLSVPRILFAGGMAGIFNWAVAIPPDVLKSRFQTAASGKYPNGFRDVLRELIREEGVSSLYKGFTAVMIRAFPANAACFLGFEVAMKFLNWIAPSL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMS4</accession>
    <name>A0A452HMS4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">calcium/calmodulin-dependent protein kinase</fullName>
            <ecNumber evidence="2">2.7.11.17</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="9">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.17"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="8">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.17"/>
        </reaction>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the protein kinase superfamily. CAMK Ser/Thr protein kinase family. CaMK subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.17" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452HMS4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018566.1">
        <property type="protein sequence ID" value="ENSGAGP00000016264.1"/>
        <property type="gene ID" value="ENSGAGG00000006503.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004683">
        <property type="term" value="F:calcium/calmodulin-dependent protein kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd14086">
        <property type="entry name" value="STKc_CaMKII"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.450.50">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.620">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013543">
        <property type="entry name" value="Ca/CaM-dep_prot_kinase-assoc"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032710">
        <property type="entry name" value="NTF2-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24347">
        <property type="entry name" value="SERINE/THREONINE-PROTEIN KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08332">
        <property type="entry name" value="CaMKII_AD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54427">
        <property type="entry name" value="NTF2-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="6" id="KW-0418">Kinase</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="5" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Protein kinase" evidence="10">
        <location>
            <begin position="1"/>
            <end position="240"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00005354"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00012434"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00047307"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00047430"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016264.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="431" mass="48944" checksum="36EDC46ECAF1EAB9" modified="2025-10-08" version="2">MLICCGSSSSADPKVYSTVGYYHQKLEREARICRLLKHPNIVRLHDSISEEGFHYLVFDLVTGGELFEDIVAREYYSEADASHCIQQILEAVLHCHQMGVVHRDLKPENLLLASKSKGAAVKLADFGLAIEVQGEQQAWFGFAGTPGYLSPEVLRKDPYGKPVDMWACGVILYILLVGYPPFWDEDQHRLYQQIKAGAYDFPSPEWDTVTPEAKDLINKMLTINPAKRITASEALKHPWICQRSTVASMMHRQETVDCLKKFNARRKLKVRKKIFFRGILVESTESSNTTIEDEDVKARKQEIIKVTEQLIEAINNGDFEAYTKICDPGLTSFEPEALGNLVEGMDFHRFYFENALSKGNKPIHTIILNPHVHLVGDDAACIAYIRLTQYMDGSGMPKTMQSEETRVWHRRDGKWQNVHFHCSGSPTVPIK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMS5</accession>
    <name>A0A452HMS5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Pleckstrin and Sec7 domain containing 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell projection</location>
            <location evidence="1">Ruffle membrane</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMS5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018605.1">
        <property type="protein sequence ID" value="ENSGAGP00000016299.1"/>
        <property type="gene ID" value="ENSGAGG00000012132.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032587">
        <property type="term" value="C:ruffle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005543">
        <property type="term" value="F:phospholipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032012">
        <property type="term" value="P:regulation of ARF protein signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd13295">
        <property type="entry name" value="PH_EFA6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00171">
        <property type="entry name" value="Sec7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.1000.11:FF:000004">
        <property type="entry name" value="PH and SEC7 domain-containing protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000054">
        <property type="entry name" value="PH and SEC7 domain-containing protein 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1000.11">
        <property type="entry name" value="Arf Nucleotide-binding Site Opener,domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041681">
        <property type="entry name" value="PH_9"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001605">
        <property type="entry name" value="PH_dom-spectrin-type"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023394">
        <property type="entry name" value="Sec7_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000904">
        <property type="entry name" value="Sec7_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035999">
        <property type="entry name" value="Sec7_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10663">
        <property type="entry name" value="GUANYL-NUCLEOTIDE EXCHANGE FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10663:SF329">
        <property type="entry name" value="PH AND SEC7 DOMAIN-CONTAINING PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15410">
        <property type="entry name" value="PH_9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01369">
        <property type="entry name" value="Sec7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00683">
        <property type="entry name" value="SPECTRINPH"/>
    </dbReference>
    <dbReference type="SMART" id="SM00233">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00222">
        <property type="entry name" value="Sec7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48425">
        <property type="entry name" value="Sec7 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50190">
        <property type="entry name" value="SEC7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="6" id="KW-0966">Cell projection</keyword>
    <keyword evidence="4 7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="SEC7" evidence="9">
        <location>
            <begin position="40"/>
            <end position="203"/>
        </location>
    </feature>
    <feature type="domain" description="PH" evidence="8">
        <location>
            <begin position="253"/>
            <end position="366"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="394"/>
            <end position="421"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004632"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50190"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016299.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="12">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="512" mass="58169" checksum="C4A55AD631C31359" modified="2019-05-08" version="1">MVHNRGGFMNASGMNNCCCHNTLSDSDSDMGSTEQLDHGSTDTLANGCKADSDAAKRLAKRLYHLEGFKRCDVARQLGKNNEFSKLVAEEYLSFFDFTGLTLDKALRTFLKAFPLMGETQERERVLIHFSRRYCQCNPEESTSEDGIHTLTCALMLLNTDLHGHNIGKKMSCQQFITNLDGLNDGKDFAKDLLKTLYNSIKNEKLEWAIDEDELRKSLSELVDDKLGASAKKVTRIVDGSNPFLDIPQALNAITYKHGVLTRKTHADTDGKRTPRGRRGWKKFYAVLKGTILYLQKDEYKPDKDLSEVDLKNAIRVHHALATKASDYSKKSNVLKLKTADWRVFLFQAPSKEEMLSWILRINLVAAIFSAPAFPAAICSMKKFCRPLLPSSMTKLCQEEQLRSHENKMRQIADELTEHKLHPVEKSLKSKEAEEYRLKEHYLIFEKSRYETYINLLCMKIKVGTDDLEKIEASLFMVEADDISLRKTHSSPSLSQGHLSVHSKVEKDVIDEA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMS8</accession>
    <name>A0A452HMS8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">glutamine--fructose-6-phosphate transaminase (isomerizing)</fullName>
            <ecNumber evidence="3">2.6.1.16</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>D-fructose 6-phosphate + L-glutamine = D-glucosamine 6-phosphate + L-glutamate</text>
            <dbReference type="Rhea" id="RHEA:13237"/>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
            <dbReference type="ChEBI" id="CHEBI:58359"/>
            <dbReference type="ChEBI" id="CHEBI:58725"/>
            <dbReference type="ChEBI" id="CHEBI:61527"/>
            <dbReference type="EC" id="2.6.1.16"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="2">Nucleotide-sugar biosynthesis; UDP-N-acetyl-alpha-D-glucosamine biosynthesis; alpha-D-glucosamine 6-phosphate from D-fructose 6-phosphate: step 1/1.</text>
    </comment>
    <dbReference type="EC" id="2.6.1.16" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452HMS8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018554.1">
        <property type="protein sequence ID" value="ENSGAGP00000016253.1"/>
        <property type="gene ID" value="ENSGAGG00000012193.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00113">
        <property type="reaction ID" value="UER00528"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097367">
        <property type="term" value="F:carbohydrate derivative binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004360">
        <property type="term" value="F:glutamine-fructose-6-phosphate transaminase (isomerizing) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006002">
        <property type="term" value="P:fructose 6-phosphate metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006487">
        <property type="term" value="P:protein N-linked glycosylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006048">
        <property type="term" value="P:UDP-N-acetylglucosamine biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="CDD" id="cd00714">
        <property type="entry name" value="GFAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05008">
        <property type="entry name" value="SIS_GlmS_GlmD_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05009">
        <property type="entry name" value="SIS_GlmS_GlmD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10490:FF:000001">
        <property type="entry name" value="Glutamine--fructose-6-phosphate aminotransferase [isomerizing]"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10490:FF:000126">
        <property type="entry name" value="Glutamine--fructose-6-phosphate aminotransferase [isomerizing] 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.10490">
        <property type="entry name" value="Glucose-6-phosphate isomerase like protein, domain 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.60.20.10">
        <property type="entry name" value="Glutamine Phosphoribosylpyrophosphate, subunit 1, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017932">
        <property type="entry name" value="GATase_2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005855">
        <property type="entry name" value="GFAT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047084">
        <property type="entry name" value="GFAT_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035466">
        <property type="entry name" value="GlmS/AgaS_SIS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035490">
        <property type="entry name" value="GlmS/FrlB_SIS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029055">
        <property type="entry name" value="Ntn_hydrolases_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001347">
        <property type="entry name" value="SIS_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046348">
        <property type="entry name" value="SIS_dom_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01135">
        <property type="entry name" value="glmS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF001484">
        <property type="entry name" value="PRK00331.1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10937">
        <property type="entry name" value="GLUCOSAMINE--FRUCTOSE-6-PHOSPHATE AMINOTRANSFERASE, ISOMERIZING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10937:SF10">
        <property type="entry name" value="GLUTAMINE--FRUCTOSE-6-PHOSPHATE AMINOTRANSFERASE [ISOMERIZING] 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13522">
        <property type="entry name" value="GATase_6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01380">
        <property type="entry name" value="SIS"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56235">
        <property type="entry name" value="N-terminal nucleophile aminohydrolases (Ntn hydrolases)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53697">
        <property type="entry name" value="SIS domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51278">
        <property type="entry name" value="GATASE_TYPE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51464">
        <property type="entry name" value="SIS"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0315">Glutamine amidotransferase</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Glutamine amidotransferase type-2" evidence="8">
        <location>
            <begin position="52"/>
            <end position="338"/>
        </location>
    </feature>
    <feature type="domain" description="SIS" evidence="9">
        <location>
            <begin position="410"/>
            <end position="549"/>
        </location>
    </feature>
    <feature type="domain" description="SIS" evidence="9">
        <location>
            <begin position="581"/>
            <end position="722"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001031"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004775"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012916"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022962"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51278"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51464"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016253.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="732" mass="83166" checksum="0E118E0D1DF9BDB6" modified="2019-05-08" version="1">MLKQGRKLCLQGAAQFRRQQNHGNTEDDGWMQCDRTARFLRWRQDLQMNVFSGIFAYLNYRVPRTRKEIFETLIKGLQRLEYRGYDSAGVAIDGNNNEDKERYIRLIKKRGKVKALDEELYKQDGMDLKADFETHFGIAHTRWATHGVPNAVNSHPQRSDKRNEFVVIHNGIITNYKDLRKFLETKGYEFESETDTETIPKLVKYMYDNRETEDISFSALVERVIQQLEGAFALVFKSIHYPGEAVATRRGSPLLIGVRSKYKLSTEQIPILYRTCNIENVKNICKARMKRLDSSTCLHAIGDKAVEFFFASDASAIIEHTNRVIFLEDDDIAAVADGKLSLHRLKRSASDDPSRAIQTLQMELQQIMKGNFSAFMQKEIFEQPESVVNTMRGRVNFETKTVLLGGLKDHLKEIRRCRRLIIIGCGTSYHAAIATRQVLEELTELPVMVELASDFLDRNTPVFRDDVCFFISQSGETADTLMALRYCKDHGALTVGITNTVGSSISRETDCGVHINAGPEIGVASTKAYTSQFISLVMFGLMMSEDRISLQNRRQEIISGLQMLPEMIKEVLSLDEKIHDLALELYKQRSLLVMGRGYNYATCLEGALKIKEITYMHSEGILAGELKHGPLALIDKHMPVIMVIMKDPCFTKCQNALQQVIARQGRPIILSSREDTESSKFAYKTIELPHTVDCLQGVLSVVPLQLLSFHLAVLRGYDVDFPRNLAKSVTVE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMS9</accession>
    <name>A0A452HMS9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HMS9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018572.1">
        <property type="protein sequence ID" value="ENSGAGP00000016270.1"/>
        <property type="gene ID" value="ENSGAGG00000012200.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1"/>
            <end position="69"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="91"/>
            <end position="134"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="8"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="1">
        <location>
            <begin position="24"/>
            <end position="34"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="40"/>
            <end position="69"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="1">
        <location>
            <begin position="95"/>
            <end position="134"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016270.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="134" mass="16533" checksum="1B3FBA14FB33BFEC" modified="2019-05-08" version="1">MRHSKRSHCSEWDDRKNWDQGKHSAPHKRKKRSHSSVQESKPDKSHHLEVSDSHYLDDRSINERDHHDRRYVEEYRNDFCEGYDHRHYHRDYEKSHHHHYSKSSGRSRKSSHKRKHKRHHCSSHHSHSMKLLPR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMT6</accession>
    <name>A0A452HMT6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">SAM domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the BicC family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMT6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018561.1">
        <property type="protein sequence ID" value="ENSGAGP00000016260.1"/>
        <property type="gene ID" value="ENSGAGG00000012189.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="CDD" id="cd22421">
        <property type="entry name" value="KH-I_BICC1_rpt2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd22422">
        <property type="entry name" value="KH-I_BICC1_rpt3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd09520">
        <property type="entry name" value="SAM_BICC1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.50:FF:000025">
        <property type="entry name" value="Ankyrin repeat and sterile alpha motif domain-containing 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.310.270:FF:000001">
        <property type="entry name" value="BicC family RNA binding protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.310.270:FF:000002">
        <property type="entry name" value="BicC family RNA binding protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.310.270">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1370.10">
        <property type="entry name" value="K Homology domain, type 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.50">
        <property type="entry name" value="Transcription Factor, Ets-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR054727">
        <property type="entry name" value="BICC1_KH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047549">
        <property type="entry name" value="BICC1_KH-I_rpt1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047554">
        <property type="entry name" value="BICC1_KH-I_rpt2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047553">
        <property type="entry name" value="BICC1_KH-I_rpt3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037974">
        <property type="entry name" value="BICC1_SAM_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004087">
        <property type="entry name" value="KH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004088">
        <property type="entry name" value="KH_dom_type_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036612">
        <property type="entry name" value="KH_dom_type_1_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001660">
        <property type="entry name" value="SAM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013761">
        <property type="entry name" value="SAM/pointed_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10627:SF78">
        <property type="entry name" value="PROTEIN BICAUDAL C HOMOLOG 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10627">
        <property type="entry name" value="SCP160"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00013">
        <property type="entry name" value="KH_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22985">
        <property type="entry name" value="KH_BICC1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24234">
        <property type="entry name" value="KH_BICC1_1st"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00536">
        <property type="entry name" value="SAM_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00322">
        <property type="entry name" value="KH"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00454">
        <property type="entry name" value="SAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54791">
        <property type="entry name" value="Eukaryotic type KH-domain (KH-domain type I)"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47769">
        <property type="entry name" value="SAM/Pointed domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50084">
        <property type="entry name" value="KH_TYPE_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50105">
        <property type="entry name" value="SAM_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="4 5" id="KW-0694">RNA-binding</keyword>
    <feature type="domain" description="SAM" evidence="7">
        <location>
            <begin position="795"/>
            <end position="858"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="512"/>
            <end position="573"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="601"/>
            <end position="620"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="702"/>
            <end position="767"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="512"/>
            <end position="541"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="556"/>
            <end position="573"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="714"/>
            <end position="735"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007662"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00117"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50105"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016260.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="877" mass="94909" checksum="913018D2E8EDA3BA" modified="2019-05-08" version="1">MEETNTQIAWPSKLKIGAKSKKDPHIKVSGKKENVKEAKEKIMSVLDTKSNRVTLKMDVSHTEHSHVIGKGGNNIKKVMEETGCHIHFPDSNRNNQAEKSNQVSIAGQPAGVESARVRIRELLPLVLMFELPIAGILQPIPDPNSPTILHISQTYNISVSFKQRSRMYGATVIVRGSQNNTSAVKEGTAMLLEHLAGSLASAIPVSTQLDIAAQHHLFMMGRNGSNIKHIMQRTGAQIHFPDPSNPQKKSTVYLQGTIESVCLARQYLMGCLPLVLMFDMKEEIEIEPQIITQLMEQLDVFISIKPKPKQPSKSVIVKSVERNALNMYETRKRLLGLESNGVTIAANPSTVSCHVGLSCPGLDILPSAGLGLTGLGLLGPTALSVNTSTAPNSLLNALNSSVSPLQSPSSGTPSPTLWATSLANTSATGFSAIPHLMIPSTAQAALTSILLSGVPNYSQNTPSPPPGLTPVDIHVNGMQSENKKGSASLNGHVKTSNIKYAALSASSLGENVLSTNHSDSSRQTNSHSTSEQVTTKSNTAEGCNDAFVEVGMPRSPSHSANASDLKQMMSSSKQSCTKRQTVELLQGTKNSHLHTTERLLSDTELSASENPVADKKAPGSERAAERAAAAQQNSERVRLAPQSSYVNMQAFDYEQKKLLATKAMLKKPVVTEVRTPTNTWSGLGFSKSMPAETIKELRRANHVSYKPSMTTTYEGSSMSLSPSSSREHLGSGSESDNWRDRNGIGPSAHSEFVSSVGSPKRKQNKSTEHYLSSSNYMDCISSLTGSNGCNLNSLFKGSDLPELFSKLGLGKYTDVFQQQEIDLQTFLTLTDQDLKELGITTFGARRKMLLAISDVLLWIDEGFSYNEGNHKVPLFLQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMW3</accession>
    <name>A0A452HMW3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">phospholipase A2</fullName>
            <ecNumber evidence="1">3.1.1.4</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="6">
            <text>a 1,2-diacyl-sn-glycero-3-phosphocholine + H2O = a 1-acyl-sn-glycero-3-phosphocholine + a fatty acid + H(+)</text>
            <dbReference type="Rhea" id="RHEA:15801"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:28868"/>
            <dbReference type="ChEBI" id="CHEBI:57643"/>
            <dbReference type="ChEBI" id="CHEBI:58168"/>
            <dbReference type="EC" id="3.1.1.4"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="6">
            <dbReference type="Rhea" id="RHEA:15802"/>
        </physiologicalReaction>
    </comment>
    <dbReference type="EC" id="3.1.1.4" evidence="1"/>
    <dbReference type="AlphaFoldDB" id="A0A452HMW3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018623.1">
        <property type="protein sequence ID" value="ENSGAGP00000016316.1"/>
        <property type="gene ID" value="ENSGAGG00000011814.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0047499">
        <property type="term" value="F:calcium-independent phospholipase A2 activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0052816">
        <property type="term" value="F:long-chain fatty acyl-CoA hydrolase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035965">
        <property type="term" value="P:cardiolipin acyl-chain remodeling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016042">
        <property type="term" value="P:lipid catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:2000304">
        <property type="term" value="P:positive regulation of ceramide biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07212">
        <property type="entry name" value="Pat_PNPLA9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000338">
        <property type="entry name" value="85/88 kDa calcium-independent phospholipase A2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.1090.10">
        <property type="entry name" value="Cytosolic phospholipase A2 catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016035">
        <property type="entry name" value="Acyl_Trfase/lysoPLipase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047148">
        <property type="entry name" value="PLPL9"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002641">
        <property type="entry name" value="PNPLA_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24139:SF34">
        <property type="entry name" value="85_88 KDA CALCIUM-INDEPENDENT PHOSPHOLIPASE A2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24139">
        <property type="entry name" value="CALCIUM-INDEPENDENT PHOSPHOLIPASE A2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01734">
        <property type="entry name" value="Patatin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01415">
        <property type="entry name" value="ANKYRIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52151">
        <property type="entry name" value="FabD/lysophospholipase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51635">
        <property type="entry name" value="PNPLA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4 7" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="3 8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="8" id="KW-0442">Lipid degradation</keyword>
    <keyword evidence="5 8" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <feature type="repeat" description="ANK" evidence="7">
        <location>
            <begin position="162"/>
            <end position="194"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="7">
        <location>
            <begin position="229"/>
            <end position="261"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="7">
        <location>
            <begin position="359"/>
            <end position="391"/>
        </location>
    </feature>
    <feature type="domain" description="PNPLA" evidence="9">
        <location>
            <begin position="432"/>
            <end position="616"/>
        </location>
    </feature>
    <feature type="short sequence motif" description="GXGXXG" evidence="8">
        <location>
            <begin position="436"/>
            <end position="441"/>
        </location>
    </feature>
    <feature type="short sequence motif" description="GXSXG" evidence="8">
        <location>
            <begin position="468"/>
            <end position="472"/>
        </location>
    </feature>
    <feature type="short sequence motif" description="DGA/G" evidence="8">
        <location>
            <begin position="603"/>
            <end position="605"/>
        </location>
    </feature>
    <feature type="active site" description="Nucleophile" evidence="8">
        <location>
            <position position="470"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="8">
        <location>
            <position position="603"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00013278"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023422"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01161"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51635"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016316.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="757" mass="85065" checksum="B9C410DD50D18F6F" modified="2019-05-08" version="1">MGSGRSCFPQKMQFFGRLVNTINSYSNLFSNPYRVKEVPLAEYVAHTRLRENDRVILYKSKNARSWDCLLVNPQSHLVAFRLFQLDNETDAIERFEQYARRLRPFYESSPQPLQLEVLQQLSDCFRNHPYWSLAHVAVETELPGSFQHNRILSCVNSTECEDGCTPLQLACRKGNMECLCELVECHARLDVTDKNGETVFHYAVRGSNPEIVKLLGKKPSVGLNHLSNQGQTPLHLACQLGKEDVVLALLKCHAKCNVMGTLGYPIHTAVKYSQKGCVQALLEVDVNQVQFKDPRYGAAVLHWAKNAEMTRMLIEYGCDVNSVSKTADMALHIMVKRGRFDSAMVLLTHGALTNAKGQDGNTPLHLAMKQDHLETIKALIVFGADVEIPNEFGETPGLVAARTSKGDIIHISTALAAIFLFSWCLSRYDRLLCLDGGGIRGLVLIQLLIAVEKAAGRPIRELFDWIAGTSTGGILALAIVHGKPVDYMRCLYFRMKNEVFRGSRPYESEPLDEFLKREFGEHTKMTDVQKPKVIVTGTLCDRQPAELHLFRNYKAPATKNKPEVKTTDPFKPLTKPEDQLVWRAARCSGAAPTYFRPIGRFLDGGLLANNPTLDAMTEIHEYNKSLSQKGQSQQVRKLGVVVSLGTGRPPQVPVRSVDVFRPSNPWELAKTVFGAKELGKMVVDCCTDADGPAVDRARAWCEMTDTRYFRFSPQLQSDVMLDEVNDAILVNALWDTQLYICQQQEQFKQLVQELLTR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMW5</accession>
    <name>A0A452HMW5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">FAM69 protein-kinase domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HMW5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018614.1">
        <property type="protein sequence ID" value="ENSGAGP00000016308.1"/>
        <property type="gene ID" value="ENSGAGG00000012224.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004715">
        <property type="term" value="F:non-membrane spanning protein tyrosine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001501">
        <property type="term" value="P:skeletal system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022049">
        <property type="entry name" value="FAM69_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042983">
        <property type="entry name" value="PKDCC"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46448:SF3">
        <property type="entry name" value="EXTRACELLULAR TYROSINE-PROTEIN KINASE PKDCC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46448">
        <property type="entry name" value="PROTEIN KINASE DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12260">
        <property type="entry name" value="PIP49_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="FAM69 protein-kinase" evidence="1">
        <location>
            <begin position="30"/>
            <end position="158"/>
        </location>
    </feature>
    <evidence type="ECO:0000259" key="1">
        <source>
            <dbReference type="Pfam" id="PF12260"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016308.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="282" mass="32238" checksum="13094F0EFD971947" modified="2019-05-08" version="1">MALYGYCYQDSNDIPDTLTAITELGSPLEMIQLLQTSWEDRFRICLSLVRLLHYLAHSPLGSVTLLDFRPRQFVIVDGELKVTDLDDASIEESSCTSNRDCFIEFPARNFTLPCSVEGRCQNMNEKRNLYNAYRFFFTYLLPHSAPSSLRPLLDMIVNATGELHWGIDETVAQLEKVLHLYKSGTYLQNTTRMPKAEYQQIPEAAIPDENYRCWPSYHHKGCLLSVFDVGEAIEVCENHSQCKAFVLTNQTTWTGRQLVFFKMGSNHIVPDPDKITYVKATG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMW7</accession>
    <name>A0A452HMW7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Pleckstrin and Sec7 domain containing 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell projection</location>
            <location evidence="1">Ruffle membrane</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMW7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018608.1">
        <property type="protein sequence ID" value="ENSGAGP00000016302.1"/>
        <property type="gene ID" value="ENSGAGG00000012132.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032587">
        <property type="term" value="C:ruffle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005543">
        <property type="term" value="F:phospholipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032012">
        <property type="term" value="P:regulation of ARF protein signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd13295">
        <property type="entry name" value="PH_EFA6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00171">
        <property type="entry name" value="Sec7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.1000.11:FF:000004">
        <property type="entry name" value="PH and SEC7 domain-containing protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000054">
        <property type="entry name" value="PH and SEC7 domain-containing protein 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1000.11">
        <property type="entry name" value="Arf Nucleotide-binding Site Opener,domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041681">
        <property type="entry name" value="PH_9"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001605">
        <property type="entry name" value="PH_dom-spectrin-type"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023394">
        <property type="entry name" value="Sec7_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000904">
        <property type="entry name" value="Sec7_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035999">
        <property type="entry name" value="Sec7_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10663">
        <property type="entry name" value="GUANYL-NUCLEOTIDE EXCHANGE FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10663:SF329">
        <property type="entry name" value="PH AND SEC7 DOMAIN-CONTAINING PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15410">
        <property type="entry name" value="PH_9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01369">
        <property type="entry name" value="Sec7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00683">
        <property type="entry name" value="SPECTRINPH"/>
    </dbReference>
    <dbReference type="SMART" id="SM00233">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00222">
        <property type="entry name" value="Sec7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48425">
        <property type="entry name" value="Sec7 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50190">
        <property type="entry name" value="SEC7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="6" id="KW-0966">Cell projection</keyword>
    <keyword evidence="4 7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="SEC7" evidence="10">
        <location>
            <begin position="244"/>
            <end position="407"/>
        </location>
    </feature>
    <feature type="domain" description="PH" evidence="9">
        <location>
            <begin position="457"/>
            <end position="570"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="161"/>
            <end position="180"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="226"/>
            <end position="246"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="692"/>
            <end position="717"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="598"/>
            <end position="625"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="706"/>
            <end position="717"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004632"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50190"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016302.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="13">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="717" mass="81267" checksum="2F6DF000E41C71CE" modified="2019-05-08" version="1">MQELREFRYLQREVPEQGLRHLTPSEPSDLPAVTQGVWDRAGSCTWYFSCLPLFSRFAVGEAKQSCMEDDKEELQLKDIRDGFSSTFEKIVESDLMKGTYYSSLDSLDVLSLTDETDSCVSFESPLTPLIQQRVKESPELLEQKLVAQQKGALQHMVAEKQEQAAAKPVEEDLGSPLRHSITSSRSENVLSRLSLKNIPNGFHIDGAEEDALKLINSISDASLKDTLSDSDSDMGSTEQLDHGSTDTLANGCKADSDAAKRLAKRLYHLEGFKRCDVARQLGKNNEFSKLVAEEYLSFFDFTGLTLDKALRTFLKAFPLMGETQERERVLIHFSRRYCQCNPEESTSEDGIHTLTCALMLLNTDLHGHNIGKKMSCQQFITNLDGLNDGKDFAKDLLKTLYNSIKNEKLEWAIDEDELRKSLSELVDDKLGASAKKVTRIVDGSNPFLDIPQALNAITYKHGVLTRKTHADTDGKRTPRGRRGWKKFYAVLKGTILYLQKDEYKPDKDLSEVDLKNAIRVHHALATKASDYSKKSNVLKLKTADWRVFLFQAPSKEEMLSWILRINLVAAIFSAPAFPAAICSMKKFCRPLLPSSMTKLCQEEQLRSHENKMRQIADELTEHKLHPVEKSLKSKEAEEYRLKEHYLIFEKSRYETYINLLCMKIKVGTDDLEKIEASLFMVEADDISLRKTHSSPSLSQGHLSVHSKVEKDVIEQNT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMX1</accession>
    <name>A0A452HMX1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">HOOK N-terminal domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the CCDC88 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018626.1">
        <property type="protein sequence ID" value="ENSGAGP00000016319.1"/>
        <property type="gene ID" value="ENSGAGG00000012232.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051959">
        <property type="term" value="F:dynein light intermediate chain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008017">
        <property type="term" value="F:microtubule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031122">
        <property type="term" value="P:cytoplasmic microtubule organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030705">
        <property type="term" value="P:cytoskeleton-dependent intracellular transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.418.10:FF:000035">
        <property type="entry name" value="girdin isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.418.10">
        <property type="entry name" value="Calponin-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036872">
        <property type="entry name" value="CH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043936">
        <property type="entry name" value="HOOK_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18947:SF30">
        <property type="entry name" value="GIRDIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18947">
        <property type="entry name" value="HOOK PROTEINS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19047">
        <property type="entry name" value="HOOK_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF116907">
        <property type="entry name" value="Hook domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 6" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="3" id="KW-0344">Guanine-nucleotide releasing factor</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="HOOK N-terminal" evidence="8">
        <location>
            <begin position="8"/>
            <end position="111"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="771"/>
            <end position="797"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="957"/>
            <end position="986"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1367"/>
            <end position="1411"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1512"/>
            <end position="1556"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1695"/>
            <end position="1724"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1736"/>
            <end position="1764"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1789"/>
            <end position="1835"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="194"/>
            <end position="373"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="1077"/>
            <end position="1335"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="965"/>
            <end position="980"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="1370"/>
            <end position="1382"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="1397"/>
            <end position="1411"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="1512"/>
            <end position="1531"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="1546"/>
            <end position="1556"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="1743"/>
            <end position="1758"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022658"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00061299"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF19047"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016319.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1835" mass="211985" checksum="E363A082C7929E67" modified="2019-05-08" version="1">MRQINPKSTTQRVNKKVNNDASLRIQNLSILVRQIKSYYQETLQQLIMMSLPNVLIIGRNPFSEPGTEEIKKLLLLLLGCAVQCQKKEEFIERIQGLDFDTRAAVAAHIQEVTHNQENVFDLQWMDVSTLSQEYIEPLLKNMALHLKRLIDERDEHSETIVELSEERDCLHFLPHASAAQSPCGSPGIKRTESRQHLSVELADAKAKIRRLRQELEEKTEQLLDCKQELEQMEAELKRLQQENMNLLSDARSARVYRDELDVLREKAIRVDKLESEVSRYKERLHDIVFYKARVEELKEDNQVLLETKTMLEDQLEGTRARSDKLHELEKENLQLKAKLHDMEMERDMDRKKIEELMEENMTLEMAQKQSMDESLHLGWELEQMNRTTELSEVPRKSLGHEVNELTSSRLLKLEMENQSLLKTVEELQNVAGSVEGSNSKILKMEKENQRLSKKLEGLENEIKQEKQSLQNSQNLSKDLMKEKAQLEKTIETLRENSERQIKILEQENEHLNQTVDALRQRSQISAEARVKDIEKENKIIHESIKEASSKLNKIEFEKKQMRKELEHYKEKGERAEELENELHHLEKENELLQKKITNLKITCEKTEGLEQENSDLEVENRKLKKTLDSFKNLTFQLESLEKENSQLDEENLELRRTVESLKCTSIKMAQLQLENKELESEKKQLKKGLELMKASCKKTERLEVSYQGLDTENQRLQKALENSTKKIQQLESELQDLESENQTLQKNLEELKISSKRLEQLEKENKLLEQETSQLEKDKKQLEKENKRLRQQAEIKDSTLEENNTKISNLEKENKSLCKEISIFKESCVRLKELEKENKELVKRATIDKKTLITLREDLVNEKLKTQQMNNDLEKLTHELEKIGLNKERLLHDEQSTDDSRYKLLESKLESTLKKSLEIKEEKIAALEARLEESTNLNQQLRQELKTVKKNYEALKQRQEEERMVQNSPPRTGEETQSVNKWEKENQETTRELLKVKDRLIEVERNNATLQAEKQALKTQLKQLETQNNNLQAQILSLQRQTVSLQEQNTTLQTQNAKLQVENSTLNSQSTSLMNQNAQLLIQQSALENENESILREREDLKSLYDSLVKDHEKLEHLHERQASEYESLIAKHGSLKSAHKNLEVEHKDLEDRYNQLLKQKVQLEELEKVLKVEQEKMLLQSKKHKTVAAEYKKLCDENDRLNHTYNQLLRENEVLQTDHKNLKSLLNNSKLEQTRLEAEFSKLKEQYQQLDITSTKLNNQCELLSQLKGNLEEENRHLLDQIQTLMLQNRTLLEQNMESKDLFHVEQRQYIDKLNELRRQKEKLEEKIMDQYKFYEPSPPRRRGNWITLKMRKLIKSKKDVNRERLKSLTLTPTRSESSEGFLQLPHQDSQDSSSVGSNSLEDGQTLGTKKSSMIALKRLPFLRNRPKEKDKMKAIYRRSMSMNDLVQSMVLAGGQWTGSSEHLEGSDDVSTGKRKKELGAMAFSTTAINFATVNSSSGFRSKQLLNNKDTTSFEDVSPQGISDDSSTGSRVHASRPASLDSGRTSTSNSNNNASLHEVKAGGINNQSRPQSHSSGEFSLLHEHEAWSSSSSSPIQYLKSHTKLSPMIQEKMPETVDNQGKQKIKTGSPGSEVVTLQQFLEESNKVTSTEIKCASQENLLDEVMKSLSESAELAGKEKLRKQSSVSSGIVKSLSGKNTVDFSDGRSAKPEQVVRPSPRKTEDTYFTSSPIKLMSGTQGKVKSIKEKVQTDPSRRQSKDCSPYATLPRASNVISTAEGTTRRTSIHDFLSKDSRQPVSIDPAPSTADRSIPSASNVEAIPESRNSKSRSREQQSS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMX2</accession>
    <name>A0A452HMX2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">PAS domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMX2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018613.1">
        <property type="protein sequence ID" value="ENSGAGP00000016307.1"/>
        <property type="gene ID" value="ENSGAGG00000011530.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000976">
        <property type="term" value="F:transcription cis-regulatory region binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001222">
        <property type="term" value="F:transcription corepressor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032922">
        <property type="term" value="P:circadian regulation of gene expression"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043153">
        <property type="term" value="P:entrainment of circadian clock by photoperiod"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000122">
        <property type="term" value="P:negative regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00130">
        <property type="entry name" value="PAS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.450.20:FF:000013">
        <property type="entry name" value="Period circadian protein homolog 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.450.20:FF:000004">
        <property type="entry name" value="Period circadian protein homolog 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.450.20">
        <property type="entry name" value="PAS domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000014">
        <property type="entry name" value="PAS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035965">
        <property type="entry name" value="PAS-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013655">
        <property type="entry name" value="PAS_fold_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057310">
        <property type="entry name" value="PER1-3_bHLH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048814">
        <property type="entry name" value="Per1-3_PAS-A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022728">
        <property type="entry name" value="Period_circadian-like_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050760">
        <property type="entry name" value="Period_circadian_regulator"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11269">
        <property type="entry name" value="PERIOD CIRCADIAN PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11269:SF8">
        <property type="entry name" value="PERIOD CIRCADIAN PROTEIN HOMOLOG 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23170">
        <property type="entry name" value="bHLH_PER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08447">
        <property type="entry name" value="PAS_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21353">
        <property type="entry name" value="Per3-like_PAS-A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12114">
        <property type="entry name" value="Period_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00091">
        <property type="entry name" value="PAS"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55785">
        <property type="entry name" value="PYP-like sensor domain (PAS domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50112">
        <property type="entry name" value="PAS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0090">Biological rhythms</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="7" id="KW-0804">Transcription</keyword>
    <keyword evidence="5" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="PAS" evidence="10">
        <location>
            <begin position="357"/>
            <end position="400"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="1"/>
            <end position="119"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="552"/>
            <end position="593"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="618"/>
            <end position="660"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="699"/>
            <end position="721"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="759"/>
            <end position="817"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="854"/>
            <end position="940"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="954"/>
            <end position="1011"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="36"/>
            <end position="45"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="46"/>
            <end position="56"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="65"/>
            <end position="99"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="100"/>
            <end position="116"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="9">
        <location>
            <begin position="776"/>
            <end position="795"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="9">
        <location>
            <begin position="805"/>
            <end position="817"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="879"/>
            <end position="888"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="893"/>
            <end position="902"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="915"/>
            <end position="925"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="956"/>
            <end position="996"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023108"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50112"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016307.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1047" mass="112471" checksum="4801058E251B6A43" modified="2019-05-08" version="1">MSAGCEPGQAPFEEGSVGGAGSGAPRRARATDDMDATSNGSSGTESHGRDSLHSDRSSSGNGKDSALLETTESSKSTNSQSPSPPSSSIAYSLLSTSSEQDNPSTSGCSSEQSARVKTQKELMKALKEMKIRLPSEKRGKGQSGTLATLQYALSCVKQVQASHDYYQQWAIDESQPCYLDVATYTVAELEGMTSEYTLKNPDTFAVAVSFVTGKILYISDQAAFVLHCKRDVFKGATFAEFLAPQDVSIFYGSTAPYHLPSWSACTSATAAAVDYTQEKSVFCRISGGRGRGRELCYTPFRLTPYLAKVRPAAGAEGQPCCLLIAERIHSGYEAPRIPPDKRVFTTRHTPSCLFQDVDERAAPLLGYLPQDLIGTPVLFYLHPEDRPLMLAIHKKILQCGGQPFDHSPLRFCTRHGEYVTIDTSWSSFVNPWSRKISFVMGRHKVRTGPLNEDVFMAPKALKMRPLEPDIQELSEQIHRLLLQPVHSNGSVGLCSMASNTSHEQFLSAASSSDSNGLLAEETPAQRPMTFQQFCKDVHMVKNQGQQMFIESHSKPLPAKPPTAGERPGPLTAQRGLGQVVSSEESGRKEPSSYSYQQINCLDGIIRYLESYNIPSRVKRKCGSSSYTGSSNSDDDKQRAGREAAGSAKGSVLPAPGEIQPEPAAAVGPALAPLALPSKAESVVSLTSQCSFSSTIVHVGDKKAPESDTAMLEEGPAPPDREQYRRVGLTKEVLSVHTQQEEQAFLTRFQELSRLHVFDAPAGRSRQEGPGAEPGHRGPRRGAGRRGKSKAKRVKQHQTSDSTGSLPPPGPCQPPAEPAPHFPAPLVAPVVALMLPNYVLPPPYFPAPFGEPFGPRLPGPPEASRSTTPERPDSPPFDSRCSSPLQLNLLQMEEAPKAAERQEAPGGQAGGTGAGSQDSWVLSRTPTPAEGQDSSNNDALSSSSDLLDLLLQEDSHSGTGSAASGSGSVRSGSNGCSTSASGTTSSKSSHTSKYFGSIDSSEAERARQPPGRVGAGEQFIKYVLQDPLWLLMANTDETVMMTYQLPDR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMX4</accession>
    <name>A0A452HMX4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Olfactory receptor</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
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                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
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            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
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        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
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        <scope>IDENTIFICATION</scope>
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        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
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        <subcellularLocation>
            <location evidence="1 12">Cell membrane</location>
            <topology evidence="1 12">Multi-pass membrane protein</topology>
        </subcellularLocation>
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    <comment type="similarity">
        <text evidence="11">Belongs to the G-protein coupled receptor 1 family.</text>
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    <dbReference type="AlphaFoldDB" id="A0A452HMX4"/>
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        <property type="gene ID" value="ENSGAGG00000012231.1"/>
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        <property type="project" value="UniProtKB-SubCell"/>
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        <property type="match status" value="1"/>
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    </evidence>
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        </source>
    </evidence>
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        <source>
            <dbReference type="PROSITE" id="PS50262"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016318.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="287" mass="32466" checksum="3BB8027DE2BBC92C" modified="2019-05-08" version="1">INHTRVTEFIMQGLFDHPHHQGLFFGLFLCLYIAVVMGNSLIIVAIVIHPPLHTPMYFFIANLALVDILCTSSVVPKILENLMQEKKTISFGGCIIQLFVLTSSLGTELLLLTVMSYDRYVAICHPLHYVKLMNKELCICLAVSVWAVGTINSFHFFCEFPTILALSWSSTYVNEIMIFMADIFLAMGNFLLTIVSYCFIIITILKIQSSKGKQRAFSTCSSHLLVVTLYYSTIIYTYIQPTSSNSLDKNKMVAIMYTLVTPTLNPVIYSLRNKEVKVHLRIDFSAC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMX5</accession>
    <name>A0A452HMX5_9SAUR</name>
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        <recommendedName>
            <fullName evidence="8">Calponin-homology (CH) domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
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        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
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        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
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        </subcellularLocation>
    </comment>
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        <property type="gene ID" value="ENSGAGG00000012232.1"/>
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        <property type="project" value="TreeGrafter"/>
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    </dbReference>
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        <property type="entry name" value="girdin isoform X1"/>
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        <property type="entry name" value="Calponin-like domain"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="CH_dom_sf"/>
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        <property type="entry name" value="HOOK_N"/>
    </dbReference>
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        <property type="entry name" value="GIRDIN"/>
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    </dbReference>
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        <property type="entry name" value="HOOK PROTEINS"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
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    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="3" id="KW-0344">Guanine-nucleotide releasing factor</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
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        </location>
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    </feature>
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        <location>
            <begin position="1840"/>
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        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="245"/>
            <end position="424"/>
        </location>
    </feature>
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            <end position="1386"/>
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        <location>
            <begin position="1016"/>
            <end position="1031"/>
        </location>
    </feature>
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            <begin position="1421"/>
            <end position="1433"/>
        </location>
    </feature>
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        <location>
            <begin position="1448"/>
            <end position="1462"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
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            <end position="1582"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="1597"/>
            <end position="1607"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="1794"/>
            <end position="1809"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022658"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00061299"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="PROSITE" id="PS50021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016328.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1886" mass="217586" checksum="C150BF0E08358F34" modified="2019-05-08" version="1">MENEIFTPLMEQFISSPLVTWVKTFGPLAGGNGSNLEEYVALVDGIFLNEVMRQINPKSTTQRVNKKVNNDASLRIQNLSILVRQIKSYYQETLQQLIMMSLPNVLIIGRNPFSEPGTEEIKKLLLLLLGCAVQCQKKEEFIERIQGLDFDTRAAVAAHIQEVTHNQENVFDLQWMDVSTLSQEYIEPLLKNMALHLKRLIDERDEHSETIVELSEERDCLHFLPHASAAQSPCGSPGIKRTESRQHLSVELADAKAKIRRLRQELEEKTEQLLDCKQELEQMEAELKRLQQENMNLLSDARSARVYRDELDVLREKAIRVDKLESEVSRYKERLHDIVFYKARVEELKEDNQVLLETKTMLEDQLEGTRARSDKLHELEKENLQLKAKLHDMEMERDMDRKKIEELMEENMTLEMAQKQSMDESLHLGWELEQMNRTTELSEVPRKSLGHEVNELTSSRLLKLEMENQSLLKTVEELQNVAGSVEGSNSKILKMEKENQRLSKKLEGLENEIKQEKQSLQNSQNLSKDLMKEKAQLEKTIETLRENSERQIKILEQENEHLNQTVDALRQRSQISAEARVKDIEKENKIIHESIKEASSKLNKIEFEKKQMRKELEHYKEKGERAEELENELHHLEKENELLQKKITNLKITCEKTEGLEQENSDLEVENRKLKKTLDSFKNLTFQLESLEKENSQLDEENLELRRTVESLKCTSIKMAQLQLENKELESEKKQLKKGLELMKASCKKTERLEVSYQGLDTENQRLQKALENSTKKIQQLESELQDLESENQTLQKNLEELKISSKRLEQLEKENKLLEQETSQLEKDKKQLEKENKRLRQQAEIKDSTLEENNTKISNLEKENKSLCKEISIFKESCVRLKELEKENKELVKRATIDKKTLITLREDLVNEKLKTQQMNNDLEKLTHELEKIGLNKERLLHDEQSTDDSRYKLLESKLESTLKKSLEIKEEKIAALEARLEESTNLNQQLRQELKTVKKNYEALKQRQEEERMVQNSPPRTGEETQSVNKWEKENQETTRELLKVKDRLIEVERNNATLQAEKQALKTQLKQLETQNNNLQAQILSLQRQTVSLQEQNTTLQTQNAKLQVENSTLNSQSTSLMNQNAQLLIQQSALENENESILREREDLKSLYDSLVKDHEKLEHLHERQASEYESLIAKHGSLKSAHKNLEVEHKDLEDRYNQLLKQKVQLEELEKVLKVEQEKMLLQSKKHKTVAAEYKKLCDENDRLNHTYNQLLRENEVLQTDHKNLKSLLNNSKLEQTRLEAEFSKLKEQYQQLDITSTKLNNQCELLSQLKGNLEEENRHLLDQIQTLMLQNRTLLEQNMESKDLFHVEQRQYIDKLNELRRQKEKLEEKIMDQYKFYEPSPPRRRGNWITLKMRKLIKSKKDVNRERLKSLTLTPTRSESSEGFLQLPHQDSQDSSSVGSNSLEDGQTLGTKKSSMIALKRLPFLRNRPKEKDKMKAIYRRSMSMNDLVQSMVLAGGQWTGSSEHLEGSDDVSTGKRKKELGAMAFSTTAINFATVNSSSGFRSKQLLNNKDTTSFEDVSPQGISDDSSTGSRVHASRPASLDSGRTSTSNSNNNASLHEVKAGGINNQSRPQSHSSGEFSLLHEHEAWSSSSSSPIQYLKSHTKLSPMIQEKMPETVDNQGKQKIKTGSPGSEVVTLQQFLEESNKVTSTEIKCASQENLLDEVMKSLSESAELAGKEKLRKQSSVSSGIVKSLSGKNTVDFSDGRSAKPEQVVRPSPRKTEDTYFTSSPIKLMSGTQGKVKSIKEKVQTDPSRRQSKDCSPYATLPRASNVISTAEGTTRRTSIHDFLSKDSRQPVSIDPAPSTADRSIPSASNVEAIPESRNSKSRSREQQSS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMY1</accession>
    <name>A0A452HMY1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">PAS domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMY1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018624.1">
        <property type="protein sequence ID" value="ENSGAGP00000016317.1"/>
        <property type="gene ID" value="ENSGAGG00000011530.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000976">
        <property type="term" value="F:transcription cis-regulatory region binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001222">
        <property type="term" value="F:transcription corepressor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032922">
        <property type="term" value="P:circadian regulation of gene expression"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043153">
        <property type="term" value="P:entrainment of circadian clock by photoperiod"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000122">
        <property type="term" value="P:negative regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00130">
        <property type="entry name" value="PAS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.450.20:FF:000013">
        <property type="entry name" value="Period circadian protein homolog 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.450.20:FF:000004">
        <property type="entry name" value="Period circadian protein homolog 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.450.20">
        <property type="entry name" value="PAS domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000014">
        <property type="entry name" value="PAS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035965">
        <property type="entry name" value="PAS-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013655">
        <property type="entry name" value="PAS_fold_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057310">
        <property type="entry name" value="PER1-3_bHLH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048814">
        <property type="entry name" value="Per1-3_PAS-A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050760">
        <property type="entry name" value="Period_circadian_regulator"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11269">
        <property type="entry name" value="PERIOD CIRCADIAN PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11269:SF8">
        <property type="entry name" value="PERIOD CIRCADIAN PROTEIN HOMOLOG 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23170">
        <property type="entry name" value="bHLH_PER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08447">
        <property type="entry name" value="PAS_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21353">
        <property type="entry name" value="Per3-like_PAS-A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00091">
        <property type="entry name" value="PAS"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55785">
        <property type="entry name" value="PYP-like sensor domain (PAS domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50112">
        <property type="entry name" value="PAS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0090">Biological rhythms</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="7" id="KW-0804">Transcription</keyword>
    <keyword evidence="5" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="PAS" evidence="10">
        <location>
            <begin position="357"/>
            <end position="400"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="1"/>
            <end position="119"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="552"/>
            <end position="574"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="622"/>
            <end position="669"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="703"/>
            <end position="739"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="36"/>
            <end position="45"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="46"/>
            <end position="56"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="65"/>
            <end position="99"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="100"/>
            <end position="116"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="9">
        <location>
            <begin position="723"/>
            <end position="736"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023108"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50112"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016317.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="920" mass="98960" checksum="FD0DF38F414474A4" modified="2019-05-08" version="1">MSAGCEPGQAPFEEGSVGGAGSGAPRRARATDDMDATSNGSSGTESHGRDSLHSDRSSSGNGKDSALLETTESSKSTNSQSPSPPSSSIAYSLLSTSSEQDNPSTSGCSSEQSARVKTQKELMKALKEMKIRLPSEKRGKGQSGTLATLQYALSCVKQVQASHDYYQQWAIDESQPCYLDVATYTVAELEGMTSEYTLKNPDTFAVAVSFVTGKILYISDQAAFVLHCKRDVFKGATFAEFLAPQDVSIFYGSTAPYHLPSWSACTSATAAAVDYTQEKSVFCRISGGRGRGRELCYTPFRLTPYLAKVRPAAGAEGQPCCLLIAERIHSGYEAPRIPPDKRVFTTRHTPSCLFQDVDERAAPLLGYLPQDLIGTPVLFYLHPEDRPLMLAIHKKILQCGGQPFDHSPLRFCTRHGEYVTIDTSWSSFVNPWSRKISFVMGRHKVRTGPLNEDVFMAPKALKMRPLEPDIQELSEQIHRLLLQPVHSNGSVGLCSMASNTSHEQFLSAASSSDSNGLLAEETPAQRPMTFQQFCKDVHMVKNQGQQMFIESHSKPLPAKPPTAGERPGPLTAPCEAQRGLGQVVSSEESGRKEPSSYSYQQINCLDGIIRYLESYNIPSRVKRKCGSSSYTGSSNSDDDKQRAGREAAGSAKGSVLPAPREEEEEPAAAVGPALAPLALPSKAESVVSLTSQCSFSSTIVHVGDKKAPESGTRGEAPLASAPPRTPSPLPSPGPAPPDREQYRRVGLTKEVLSVHTQQEEQAFLTRFQELSRLHVFDAPAGRSRQEGPGAEPGGVFVRGGGFVPGPCPPFANPPPSAPVSDRLGVDWLKPWWGRSQDSWVLSPALGGEWGLVGQSREGWEPGLLGSLLGVGRAVGASGSEQGGLGARTPGFSPRCWEGSGDWWLEQGGLGARTPGFSPRH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMY5</accession>
    <name>A0A452HMY5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Protein transport protein SEC23</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="12">Component of the coat protein complex II (COPII) which promotes the formation of transport vesicles from the endoplasmic reticulum (ER). The coat has two main functions, the physical deformation of the endoplasmic reticulum membrane into vesicles and the selection of cargo molecules for their transport to the Golgi complex.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="13">Cytoplasmic vesicle</location>
            <location evidence="13">COPII-coated vesicle membrane</location>
            <topology evidence="13">Peripheral membrane protein</topology>
            <orientation evidence="13">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1 13">Endoplasmic reticulum membrane</location>
            <topology evidence="1 13">Peripheral membrane protein</topology>
            <orientation evidence="1 13">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="13">Cytoplasm</location>
            <location evidence="13">Cytosol</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 13">Belongs to the SEC23/SEC24 family. SEC23 subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMY5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018637.1">
        <property type="protein sequence ID" value="ENSGAGP00000016330.1"/>
        <property type="gene ID" value="ENSGAGG00000006201.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030127">
        <property type="term" value="C:COPII vesicle coat"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070971">
        <property type="term" value="C:endoplasmic reticulum exit site"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090110">
        <property type="term" value="P:COPII-coated vesicle cargo loading"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006886">
        <property type="term" value="P:intracellular protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd01478">
        <property type="entry name" value="Sec23-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd11287">
        <property type="entry name" value="Sec23_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.380:FF:000001">
        <property type="entry name" value="Protein transport protein SEC23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1670:FF:000006">
        <property type="entry name" value="Protein transport protein SEC23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.20.10:FF:000003">
        <property type="entry name" value="Protein transport protein SEC23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.410:FF:000011">
        <property type="entry name" value="Protein transport protein SEC23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1670">
        <property type="entry name" value="beta-sandwich domain of Sec23/24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.730">
        <property type="entry name" value="Sec23/Sec24 helical domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.20.10">
        <property type="entry name" value="Severin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.410">
        <property type="entry name" value="von Willebrand factor, type A domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.380">
        <property type="entry name" value="Zn-finger domain of Sec23/24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029006">
        <property type="entry name" value="ADF-H/Gelsolin-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012990">
        <property type="entry name" value="Beta-sandwich_Sec23_24"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007123">
        <property type="entry name" value="Gelsolin-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036180">
        <property type="entry name" value="Gelsolin-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037364">
        <property type="entry name" value="Sec23"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006900">
        <property type="entry name" value="Sec23/24_helical_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036175">
        <property type="entry name" value="Sec23/24_helical_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006896">
        <property type="entry name" value="Sec23/24_trunk_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037550">
        <property type="entry name" value="Sec23_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036465">
        <property type="entry name" value="vWFA_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006895">
        <property type="entry name" value="Znf_Sec23_Sec24"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036174">
        <property type="entry name" value="Znf_Sec23_Sec24_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11141">
        <property type="entry name" value="PROTEIN TRANSPORT PROTEIN SEC23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11141:SF10">
        <property type="entry name" value="PROTEIN TRANSPORT PROTEIN SEC23B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00626">
        <property type="entry name" value="Gelsolin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08033">
        <property type="entry name" value="Sec23_BS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04815">
        <property type="entry name" value="Sec23_helical"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04811">
        <property type="entry name" value="Sec23_trunk"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04810">
        <property type="entry name" value="zf-Sec23_Sec24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81995">
        <property type="entry name" value="beta-sandwich domain of Sec23/24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF82754">
        <property type="entry name" value="C-terminal, gelsolin-like domain of Sec23/24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81811">
        <property type="entry name" value="Helical domain of Sec23/24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53300">
        <property type="entry name" value="vWA-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF82919">
        <property type="entry name" value="Zn-finger domain of Sec23/24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 13" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11 13" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="6 13" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="8 13" id="KW-0931">ER-Golgi transport</keyword>
    <keyword evidence="10 13" id="KW-0472">Membrane</keyword>
    <keyword evidence="5 13" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9 13" id="KW-0653">Protein transport</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 13" id="KW-0813">Transport</keyword>
    <keyword evidence="7 13" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="Zinc finger Sec23/Sec24-type" evidence="15">
        <location>
            <begin position="58"/>
            <end position="98"/>
        </location>
    </feature>
    <feature type="domain" description="Sec23/Sec24 trunk" evidence="16">
        <location>
            <begin position="127"/>
            <end position="380"/>
        </location>
    </feature>
    <feature type="domain" description="Sec23/Sec24 beta-sandwich" evidence="18">
        <location>
            <begin position="391"/>
            <end position="494"/>
        </location>
    </feature>
    <feature type="domain" description="Sec23/Sec24 helical" evidence="17">
        <location>
            <begin position="543"/>
            <end position="580"/>
        </location>
    </feature>
    <feature type="domain" description="Gelsolin-like" evidence="14">
        <location>
            <begin position="596"/>
            <end position="682"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004397"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009210"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022892"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00037370"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="RuleBase" id="RU365030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="Pfam" id="PF00626"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="Pfam" id="PF04810"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF04811"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF04815"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF08033"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016330.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="729" mass="82119" checksum="36BDB93706FD3221" modified="2019-05-08" version="1">MATYLEFIQQNEERDGVRFSWNVWPSSRLEATRMVVPLACLLTPLKERLDLPPVQYEPVLCSRQTCKAVLNPLCQVDYRAKLWACNFCFQRNQFPPAYAGISEINQPAELMPQFSTIEYIVQVNASWLKEDLQALKESLQMSLSLLPPEALVGLITFGRMVQVHELSCEGISKSYVFRGTKDLTAKQIQDMLGLTRPVVPVQQGRPLQPQEQPPISSRFLQPVHKIDMNLTDLLGELQRDPWPVTQGKRPLRSTGVALSIAVGLLEGTFPNTGARIMLFTGGPPTQGPGMVVGDELKTPIRSWHDIEKDNARFMKKATKHYETLANRTAANGHCIDIYACALDQTGLLEMKCCANLTGGHMMMGDSFNTSLFKQTFQRVFSKDFNGEFRMAFGATLEVKTSRELKISGAIGPCVSLNAKGPCVSENELGIGGTCQWKICSLDPSSTLAIYFEVVNQHNAPIPQGGRGAVQFVTQYQHSSTQRRIRVTTIARNWADAQSQLQHIEAAFDQEVAAVLMARLGVYRAESEEGPDVLRWLDRQLIRLFMFHLRRSPFLQVFNNSPDESSFYRHHFSRQDLTQSLIMIQPILYSYSFSGPPEPVLLDSSSILADRILLMDTFFQIVIYHGETIAQWQKAGYQDMPEYENFKHLLQAPLDDAQEILQTRFPMPRYINTEHGSSQARFLLSKVNPSQTHNNLYAWGQESGSPILTDDVSLQVFMDHLKKLAVSSAS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMY7</accession>
    <name>A0A452HMY7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5 12">Dolichyl-diphosphooligosaccharide--protein glycosyltransferase subunit 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1 12">Subunit of the oligosaccharyl transferase (OST) complex that catalyzes the initial transfer of a defined glycan (Glc(3)Man(9)GlcNAc(2) in eukaryotes) from the lipid carrier dolichol-pyrophosphate to an asparagine residue within an Asn-X-Ser/Thr consensus motif in nascent polypeptide chains, the first step in protein N-glycosylation. N-glycosylation occurs cotranslationally and the complex associates with the Sec61 complex at the channel-forming translocon complex that mediates protein translocation across the endoplasmic reticulum (ER). All subunits are required for a maximal enzyme activity.</text>
    </comment>
    <comment type="pathway">
        <text evidence="3 12">Protein modification; protein glycosylation.</text>
    </comment>
    <comment type="subunit">
        <text evidence="11">Component of the oligosaccharyltransferase (OST) complex. OST exists in two different complex forms which contain common core subunits RPN1, RPN2, OST48, OST4, DAD1 and TMEM258, either STT3A or STT3B as catalytic subunits, and form-specific accessory subunits. STT3A complex assembly occurs through the formation of 3 subcomplexes. Subcomplex 1 contains RPN1 and TMEM258, subcomplex 2 contains the STT3A-specific subunits STT3A, DC2/OSTC, and KCP2 as well as the core subunit OST4, and subcomplex 3 contains RPN2, DAD1, and OST48. The STT3A complex can form stable complexes with the Sec61 complex or with both the Sec61 and TRAP complexes. Interacts with TMEM35A/NACHO.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2 12">Endoplasmic reticulum membrane</location>
            <topology evidence="2 12">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4 12">Belongs to the OST1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMY7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018638.1">
        <property type="protein sequence ID" value="ENSGAGP00000016331.1"/>
        <property type="gene ID" value="ENSGAGG00000012238.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00378"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008250">
        <property type="term" value="C:oligosaccharyltransferase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0018279">
        <property type="term" value="P:protein N-linked glycosylation via asparagine"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007676">
        <property type="entry name" value="Ribophorin_I"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21049:SF0">
        <property type="entry name" value="DOLICHYL-DIPHOSPHOOLIGOSACCHARIDE--PROTEIN GLYCOSYLTRANSFERASE SUBUNIT 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21049">
        <property type="entry name" value="RIBOPHORIN I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04597">
        <property type="entry name" value="Ribophorin_I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 12" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="10 12" id="KW-0472">Membrane</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7 12" id="KW-0732">Signal</keyword>
    <keyword evidence="6 12" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9 12" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="12">
        <location>
            <begin position="1"/>
            <end position="17"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5018808884" description="Dolichyl-diphosphooligosaccharide--protein glycosyltransferase subunit 1" evidence="12">
        <location>
            <begin position="18"/>
            <end position="546"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="428"/>
            <end position="446"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00002791"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004115"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004922"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00008905"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00017611"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00046898"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="RuleBase" id="RU361143"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016331.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="546" mass="61624" checksum="EB5E83B4A9D59035" modified="2019-05-08" version="1">MEGLACRLLLCLGCALAAPELLNEEVKRTVDLSTHLAKVTAELSLANPAGGSGGASSFLLALDPGLESRLAYLGVQVKGEEEEENTLEVRETKVKGKSGKFFTVKLPSPLAPGAKIRLSIETVFTHVLQPYPTHITQAEKQFVVFEGNHYFYSPYPTKTQTTRVKLASRNIENYTKLGNPTRSEDMIEYGPFKDIPPYSQDILKVHYENNNPFLTITSMIRVIEVSHWGNIAVEETVDLKHTGAVLKGPFSRYDYQRQPDSGISSVKSFKTILPAAAQDVYYRDEIGNISTSHLLILDDSVEMEIRPRFPLFGGWKTHYIIGYNLPSYEYLYNLGDQYALKMRFIDHVFDEQVADSLTVKIILPEGAKNIHVDSPYEINRATDELHYTYLDTFGRPVIVAHKNNLVEQHIQDIVVHYTFNKILMLQEPLLVVGAFYILFFTVIFYVRLDFSITKDPAAEARMKVACITEQVLTLVNKRLGLYRHFDEAVNKYKQSRDISTLNSGKKTLETEHKALTNEVASLQSKLKTEGSDMCDKQTQLLAAPRP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMZ2</accession>
    <name>A0A452HMZ2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Calponin-homology (CH) domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the CCDC88 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018643.1">
        <property type="protein sequence ID" value="ENSGAGP00000016336.1"/>
        <property type="gene ID" value="ENSGAGG00000012232.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051959">
        <property type="term" value="F:dynein light intermediate chain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008017">
        <property type="term" value="F:microtubule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031122">
        <property type="term" value="P:cytoplasmic microtubule organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030705">
        <property type="term" value="P:cytoskeleton-dependent intracellular transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd22229">
        <property type="entry name" value="HkD_Girdin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.418.10:FF:000035">
        <property type="entry name" value="girdin isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.418.10">
        <property type="entry name" value="Calponin-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001715">
        <property type="entry name" value="CH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036872">
        <property type="entry name" value="CH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043936">
        <property type="entry name" value="HOOK_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18947:SF30">
        <property type="entry name" value="GIRDIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18947">
        <property type="entry name" value="HOOK PROTEINS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19047">
        <property type="entry name" value="HOOK_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF116907">
        <property type="entry name" value="Hook domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50021">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 6" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="3" id="KW-0344">Guanine-nucleotide releasing factor</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Calponin-homology (CH)" evidence="8">
        <location>
            <begin position="12"/>
            <end position="132"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="776"/>
            <end position="802"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="962"/>
            <end position="991"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1372"/>
            <end position="1416"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1517"/>
            <end position="1561"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1700"/>
            <end position="1729"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1741"/>
            <end position="1769"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1794"/>
            <end position="1840"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="245"/>
            <end position="424"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="1082"/>
            <end position="1340"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="970"/>
            <end position="985"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="1375"/>
            <end position="1387"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="1402"/>
            <end position="1416"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="1517"/>
            <end position="1536"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="1551"/>
            <end position="1561"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="1748"/>
            <end position="1763"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022658"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00061299"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016336.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1840" mass="212173" checksum="4B16350AA4CF09A0" modified="2019-05-08" version="1">MENEIFTPLMEQFISSPLVTWVKTFGPLAGGNGSNLEEYVALVDGIFLNEVMRQINPKSTTQRVNKKVNNDASLRIQNLSILVRQIKSYYQETLQQLIMMSLPNVLIIGRNPFSEPGTEEIKKLLLLLLGCAVQCQKKEEFIERIQGLDFDTRAAVAAHIQEVTHNQENVFDLQWMDVSTLSQEYIEPLLKNMALHLKRLIDERDEHSETIVELSEERDCLHFLPHASAAQSPCGSPGIKRTESRQHLSVELADAKAKIRRLRQELEEKTEQLLDCKQELEQMEAELKRLQQENMNLLSDARSARVYRDELDVLREKAIRVDKLESEVSRYKERLHDIVFYKARVEELKEDNQVLLETKTMLEDQLEGTRARSDKLHELEKENLQLKAKLHDMEMERDMDRKKIEELMEENMTLEMAQKQSMDESLHLGWELEQMNRTTELSEVPRKSLGHEVNELTSSRLLKLEMENQSLLKTVEELQNVAGSVEGSNSKILKMEKENQRLSKKIKILEQENEHLNQTVDALRQRSQISAEARVKDIEKENKIIHESIKEASSKLNKIEFEKKQMRKELEHYKEKGERAEELENELHHLEKENELLQKKITNLKITCEKTEGLEQENSDLEVENRKLKKTLDSFKNLTFQLESLEKENSQLDEENLELRRTVESLKCTSIKMAQLQLENKELESEKKQLKKGLELMKASCKKTERLEVSYQGLDTENQRLQKALENSTKKIQQLESELQDLESENQTLQKNLEELKISSKRLEQLEKENKLLEQETSQLEKDKKQLEKENKRLRQQAEIKDSTLEENNTKISNLEKENKSLCKEISIFKESCVRLKELEKENKELVKRATIDKKTLITLREDLVNEKLKTQQMNNDLEKLTHELEKIGLNKERLLHDEQSTDDSRYKLLESKLESTLKKSLEIKEEKIAALEARLEESTNLNQQLRQELKTVKKNYEALKQRQEEERMVQNSPPRTGEETQSVNKWEKENQETTRELLKVKDRLIEVERNNATLQAEKQALKTQLKQLETQNNNLQAQILSLQRQTVSLQEQNTTLQTQNAKLQVENSTLNSQSTSLMNQNAQLLIQQSALENENESILREREDLKSLYDSLVKDHEKLEHLHERQASEYESLIAKHGSLKSAHKNLEVEHKDLEDRYNQLLKQKVQLEELEKVLKVEQEKMLLQSKKHKTVAAEYKKLCDENDRLNHTYNQLLRENEVLQTDHKNLKSLLNNSKLEQTRLEAEFSKLKEQYQQLDITSTKLNNQCELLSQLKGNLEEENRHLLDQIQTLMLQNRTLLEQNMESKDLFHVEQRQYIDKLNELRRQKEKLEEKIMDQYKFYEPSPPRRRGNWITLKMRKLIKSKKDVNRERLKSLTLTPTRSESSEGFLQLPHQDSQDSSSVGSNSLEDGQTLGTKKSSMIALKRLPFLRNRPKEKDKMKAIYRRSMSMNDLVQSMVLAGGQWTGSSEHLEGSDDVSTGKRKKELGAMAFSTTAINFATVNSSSGFRSKQLLNNKDTTSFEDVSPQGISDDSSTGSRVHASRPASLDSGRTSTSNSNNNASLHEVKAGGINNQSRPQSHSSGEFSLLHEHEAWSSSSSSPIQYLKSHTKLSPMIQEKMPETVDNQGKQKIKTGSPGSEVVTLQQFLEESNKVTSTEIKCASQENLLDEVMKSLSESAELAGKEKLRKQSSVSSGIVKSLSGKNTVDFSDGRSAKPEQVVRPSPRKTEDTYFTSSPIKLMSGTQGKVKSIKEKVQTDPSRRQSKDCSPYATLPRASNVISTAEGTTRRTSIHDFLSKDSRQPVSIDPAPSTADRSIPSASNVEAIPESRNSKSRSREQQSS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMZ3</accession>
    <name>A0A452HMZ3_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="4">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMZ3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018664.1">
        <property type="protein sequence ID" value="ENSGAGP00000016354.1"/>
        <property type="gene ID" value="ENSGAGG00000012116.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008028">
        <property type="term" value="F:monocarboxylic acid transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1250.20">
        <property type="entry name" value="MFS general substrate transporter like domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036259">
        <property type="entry name" value="MFS_trans_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050327">
        <property type="entry name" value="Proton-linked_MCT"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11360">
        <property type="entry name" value="MONOCARBOXYLATE TRANSPORTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11360:SF14">
        <property type="entry name" value="MONOCARBOXYLATE TRANSPORTER 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103473">
        <property type="entry name" value="MFS general substrate transporter"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0472">Membrane</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="3" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="3">
        <location>
            <begin position="229"/>
            <end position="247"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="3">
        <location>
            <begin position="299"/>
            <end position="315"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="3">
        <location>
            <begin position="321"/>
            <end position="344"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="3">
        <location>
            <begin position="356"/>
            <end position="374"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="166"/>
            <end position="196"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="176"/>
            <end position="195"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016354.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="386" mass="41754" checksum="47B65B8AEBC8243F" modified="2019-05-08" version="1">MGEILIPWTSLGLVFNPGAAPGASSRHQLKTQVLGAANTKGQHSGRYRGGMSGSLAAGPSVPLGAKDLPLTCHPREERCTLLIFGGITLNLVPSSMLLRPVIKKLQDFSRNVESGSLTQSKHPATLTSHPDKISTTETQFNIRLDSPMQDSAVTNGFQSAETLGSLVPGKSAGLEGSERETSNRYTPGRDSDKDCSYNNFPLRKEWNSNSRPVPSRQPLLDFSLLKDPFFCIFVWSFLFSQLAYFIPTFHLAARAKTLGIDSMDASYLISVAGITETVSQLLSGWIADQNWIKKYHYHKTYLILCGITNLLGPLATSFPLLMVYTVSFAIFCGGYLALLLPVLVSKGSYQEVIRSLGFASFFAGIAAIAGPPIAGNISNISPLSSQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMZ5</accession>
    <name>A0A452HMZ5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Major facilitator superfamily (MFS) profile domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMZ5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018647.1">
        <property type="protein sequence ID" value="ENSGAGP00000016340.1"/>
        <property type="gene ID" value="ENSGAGG00000012116.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008028">
        <property type="term" value="F:monocarboxylic acid transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1250.20:FF:000141">
        <property type="entry name" value="Solute carrier family 16, member 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1250.20">
        <property type="entry name" value="MFS general substrate transporter like domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011701">
        <property type="entry name" value="MFS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020846">
        <property type="entry name" value="MFS_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036259">
        <property type="entry name" value="MFS_trans_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050327">
        <property type="entry name" value="Proton-linked_MCT"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11360">
        <property type="entry name" value="MONOCARBOXYLATE TRANSPORTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11360:SF14">
        <property type="entry name" value="MONOCARBOXYLATE TRANSPORTER 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07690">
        <property type="entry name" value="MFS_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103473">
        <property type="entry name" value="MFS general substrate transporter"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50850">
        <property type="entry name" value="MFS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="3" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="3">
        <location>
            <begin position="21"/>
            <end position="48"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="3">
        <location>
            <begin position="68"/>
            <end position="89"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="3">
        <location>
            <begin position="101"/>
            <end position="121"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="3">
        <location>
            <begin position="127"/>
            <end position="148"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="3">
        <location>
            <begin position="275"/>
            <end position="293"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="3">
        <location>
            <begin position="313"/>
            <end position="333"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="3">
        <location>
            <begin position="345"/>
            <end position="361"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="3">
        <location>
            <begin position="367"/>
            <end position="390"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="3">
        <location>
            <begin position="402"/>
            <end position="422"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="3">
        <location>
            <begin position="428"/>
            <end position="450"/>
        </location>
    </feature>
    <feature type="domain" description="Major facilitator superfamily (MFS) profile" evidence="4">
        <location>
            <begin position="274"/>
            <end position="470"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="212"/>
            <end position="242"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="222"/>
            <end position="241"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50850"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016340.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="470" mass="51996" checksum="589CC1F97B1BC89D" modified="2019-05-08" version="1">LTAMEEEEQQPIPPPDGGWGWMIVLHFFLVNVLVMGMLKTFGIFFVVFQEEVGGSSEQISWIGSIMSSLRFSAGLGFACLYQAAAVMTARYFKKRLALSNAIARSGMGLTFILAPFTQLLIDFYGTLLIFGGITLNLVPSSMLLRPVIKKLQDFSRNVESGSLTQSKHPATLTSHPDKISTTETQFNIRLDSPMQDSAVTNGFQSAETLGSLVPGKSAGLEGSERETSNRYTPGRDSDKDCSYNNFPLRKEWNSNSRPVPSRQPLLDFSLLKDPFFCIFVWSFLFSQLAYFIPTFHLAARAKTLGIDSMDASYLISVAGITETVSQLLSGWIADQNWIKKYHYHKTYLILCGITNLLGPLATSFPLLMVYTVSFAIFCGGYLALLLPVLVSKGSYQEVIRSLGFASFFAGIAAIAGPPIAGWLSDYTQTYACSFLFAGTCYLIAPISFFFEPSAHKWKLQKEKELNEVTS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN43</accession>
    <name>A0A452HN43_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Laminin subunit alpha-5</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="14">Laminin-10 subunit alpha</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="16">Laminin-11 subunit alpha</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="15">Laminin-15 subunit alpha</fullName>
        </alternativeName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="12">Laminin is a complex glycoprotein, consisting of three different polypeptide chains (alpha, beta, gamma), which are bound to each other by disulfide bonds into a cross-shaped molecule comprising one long and three short arms with globules at each end. Alpha-5 is a subunit of laminin-10 (laminin-511), laminin-11 (laminin-521) and laminin-15 (laminin-523).</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
            <location evidence="1">Basement membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="17">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018724.1">
        <property type="protein sequence ID" value="ENSGAGP00000016408.1"/>
        <property type="gene ID" value="ENSGAGG00000012074.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043259">
        <property type="term" value="C:laminin-10 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045202">
        <property type="term" value="C:synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009887">
        <property type="term" value="P:animal organ morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007411">
        <property type="term" value="P:axon guidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070831">
        <property type="term" value="P:basement membrane assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048732">
        <property type="term" value="P:gland development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002009">
        <property type="term" value="P:morphogenesis of an epithelium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030155">
        <property type="term" value="P:regulation of cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030334">
        <property type="term" value="P:regulation of cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045995">
        <property type="term" value="P:regulation of embryonic development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060541">
        <property type="term" value="P:respiratory system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034446">
        <property type="term" value="P:substrate adhesion-dependent cell spreading"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035239">
        <property type="term" value="P:tube morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd00055">
        <property type="entry name" value="EGF_Lam"/>
        <property type="match status" value="17"/>
    </dbReference>
    <dbReference type="CDD" id="cd00110">
        <property type="entry name" value="LamG"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000074">
        <property type="entry name" value="Laminin subunit alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000083">
        <property type="entry name" value="Laminin subunit alpha"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000090">
        <property type="entry name" value="laminin subunit alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000069">
        <property type="entry name" value="Laminin subunit alpha 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000454">
        <property type="entry name" value="Laminin subunit alpha 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000033">
        <property type="entry name" value="Laminin subunit alpha 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000034">
        <property type="entry name" value="Laminin subunit alpha 3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000084">
        <property type="entry name" value="Laminin subunit alpha 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000051">
        <property type="entry name" value="Laminin subunit alpha 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000209">
        <property type="entry name" value="Laminin subunit alpha 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000405">
        <property type="entry name" value="Laminin subunit alpha 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000430">
        <property type="entry name" value="Laminin subunit alpha 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000437">
        <property type="entry name" value="Laminin subunit alpha 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000467">
        <property type="entry name" value="Laminin subunit alpha 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.200:FF:000151">
        <property type="entry name" value="Laminin subunit alpha 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.260:FF:000444">
        <property type="entry name" value="Laminin subunit alpha 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.200">
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.260">
        <property type="entry name" value="Galactose-binding domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="18"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050440">
        <property type="entry name" value="Laminin/Netrin_ECM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009254">
        <property type="entry name" value="Laminin_aI"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010307">
        <property type="entry name" value="Laminin_dom_II"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001791">
        <property type="entry name" value="Laminin_G"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000034">
        <property type="entry name" value="Laminin_IV"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008211">
        <property type="entry name" value="Laminin_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002049">
        <property type="entry name" value="LE_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056863">
        <property type="entry name" value="LMN_ATRN_NET-like_EGF"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10574:SF406">
        <property type="entry name" value="LAMININ SUBUNIT ALPHA 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10574">
        <property type="entry name" value="NETRIN/LAMININ-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00053">
        <property type="entry name" value="EGF_laminin"/>
        <property type="match status" value="16"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24973">
        <property type="entry name" value="EGF_LMN_ATRN"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00052">
        <property type="entry name" value="Laminin_B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00054">
        <property type="entry name" value="Laminin_G_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02210">
        <property type="entry name" value="Laminin_G_2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06008">
        <property type="entry name" value="Laminin_I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06009">
        <property type="entry name" value="Laminin_II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00055">
        <property type="entry name" value="Laminin_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00011">
        <property type="entry name" value="EGFLAMININ"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="SMART" id="SM00180">
        <property type="entry name" value="EGF_Lam"/>
        <property type="match status" value="20"/>
    </dbReference>
    <dbReference type="SMART" id="SM00281">
        <property type="entry name" value="LamB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00282">
        <property type="entry name" value="LamG"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00136">
        <property type="entry name" value="LamNT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="17"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01248">
        <property type="entry name" value="EGF_LAM_1"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50027">
        <property type="entry name" value="EGF_LAM_2"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50025">
        <property type="entry name" value="LAM_G_DOMAIN"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51115">
        <property type="entry name" value="LAMININ_IVA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51117">
        <property type="entry name" value="LAMININ_NTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0084">Basement membrane</keyword>
    <keyword evidence="7" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="8 18" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="9 17" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11 17" id="KW-0424">Laminin EGF-like domain</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <feature type="domain" description="Laminin N-terminal" evidence="23">
        <location>
            <begin position="1"/>
            <end position="135"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="21">
        <location>
            <begin position="265"/>
            <end position="309"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="21">
        <location>
            <begin position="331"/>
            <end position="377"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="21">
        <location>
            <begin position="378"/>
            <end position="423"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="21">
        <location>
            <begin position="424"/>
            <end position="473"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="21">
        <location>
            <begin position="514"/>
            <end position="559"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="21">
        <location>
            <begin position="611"/>
            <end position="663"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="21">
        <location>
            <begin position="1247"/>
            <end position="1292"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="21">
        <location>
            <begin position="1337"/>
            <end position="1385"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="21">
        <location>
            <begin position="1386"/>
            <end position="1436"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin IV type A" evidence="22">
        <location>
            <begin position="1457"/>
            <end position="1635"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="21">
        <location>
            <begin position="1669"/>
            <end position="1718"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="21">
        <location>
            <begin position="1719"/>
            <end position="1774"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="21">
        <location>
            <begin position="1775"/>
            <end position="1828"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="21">
        <location>
            <begin position="1829"/>
            <end position="1875"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="21">
        <location>
            <begin position="1876"/>
            <end position="1922"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin G" evidence="20">
        <location>
            <begin position="2637"/>
            <end position="2805"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin G" evidence="20">
        <location>
            <begin position="2812"/>
            <end position="2985"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin G" evidence="20">
        <location>
            <begin position="2997"/>
            <end position="3163"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin G" evidence="20">
        <location>
            <begin position="3170"/>
            <end position="3306"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="2132"/>
            <end position="2152"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="18">
        <location>
            <begin position="2332"/>
            <end position="2392"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="285"/>
            <end position="294"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="331"/>
            <end position="343"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="354"/>
            <end position="363"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="378"/>
            <end position="390"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="380"/>
            <end position="397"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="399"/>
            <end position="408"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="424"/>
            <end position="436"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="444"/>
            <end position="453"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="514"/>
            <end position="526"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="516"/>
            <end position="533"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="535"/>
            <end position="544"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="634"/>
            <end position="643"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="1247"/>
            <end position="1259"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="1249"/>
            <end position="1266"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="1268"/>
            <end position="1277"/>
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    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="1361"/>
            <end position="1370"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="1386"/>
            <end position="1398"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="1388"/>
            <end position="1405"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="1407"/>
            <end position="1416"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="1688"/>
            <end position="1697"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="1745"/>
            <end position="1754"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="1800"/>
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    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="1812"/>
            <end position="1826"/>
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    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="1848"/>
            <end position="1857"/>
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    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="1896"/>
            <end position="1905"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004302"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022869"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023292"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00063580"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00072596"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00077092"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00078366"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00081617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00460"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50025"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50027"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS51115"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS51117"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016408.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="3309" mass="363278" checksum="98330F20F937BCB8" modified="2019-05-08" version="1">PRPDLWVLERSTDFGLTYEPWQYFASSRRDCIEKFGPNTVKRITKDDDTICTTEYSRIVPLENGEIVVSLVNGRPGAMNFSYSPLLRNFTKATNIRLRFLQTNTLLGHLMGKALRDPTVTRRYYYSIKDISIGGRCVCHGHADVCDAKDPTDPYRLQCDCQHNTCGGSCDRCCPGFNQLPWKPATTDSANECQPCNCNGHAYDCYYDPEVDRHKASKNYDNQYKGGGVCIECQHHTTGINCERCIPGYYKSPDHSIDSPYICYRCNCDSEFTDGTCEDLTGRCYCKPNYTGELCNACAEGYTDFPHCYPVPAFSHNDTGEQVLPAGQIINCDCNAAGTEGNACRKDSQRGMCVCKPNFQGAQCDQCAPGHYGPGCQPCQCSGPGQYDGTCDSETGQCLCRSGFEGYSCNQCAPGYFSYPLCQLCGCSAVGTLPGGCDSGGRCFCRPEFDGPHCDQCSMGYHSYPRCQACSCDPRGSVDNNCSPAGQCRCHSNYAGHTCSQCAPGFYGYPSCTSCQCSTEGSLHGTCDQETGQCSCRPKVAGLRCDTCMPGAYGFPHCEVGSCNPAGLATADPSLSPGSCVCRAYVEGPACDRCKPLYWNLTPDNPYGCTSCQCDTKGTISGVAECQQGNGQCFCKPSICGQLCSTCKDGYFNLDHANYFGCQGCRCDVGGSIGPACEARTGACHCSCHHRAGTTLDGRAVRFGYNPLEFENFSWRGYAQMSPIQPKVVVTLNVTSPDLFRIVFRYVNRGPANVEGRVSVLEEGKFNVCGNCTEQTKQIIFAPSTEPAFVTVPQNSFGEPFVLNPSVWSLIVEAEGILLDYLVLLPSSYYEAPILQLKVTEACTYSPAPEQASQNCLLYKYLSVDSFPSASGGDMVCRLDNSLPRPCPMVPITPSHPQMALCSGSDVEVQLRLVVPQPGRYVLLVEYANKDAMQRASVAISSLQLATQQGSLTFHTCDYRYEARSEQEAERRGSWSEYGNVPLLHCTVRGQAFLFSLYLIPYGQFTMEFVEPKVHCISLHGTFSPSSSSCIPSRFMKLSQSIVLEAGQALSIPPDFPLTHVFTVAPTGIPTEPLPRPPTAVDSSAELILLQPSQAAAVFNSRIQTLGRYAFIMHYYQPNHPTFPVEVLINGGRIWQGQAKAAFCPHGYGCRSVVVSENQVILDVTDNELTVIVRVPAGRQLWLDYILVVPEDSYSSHYLLEEPLDKSYEFISSCGVNSFYINPTTSSKFCRDSAISLSLFYNNGAQSCRCHEAGAIGGQCELFGGQCACKSNIIGRECSRCATGYWGFPNCRPCDCGTRLCDEVTGQCICPPHTIKPECVVCEPQTFGCHALIGCEDCNCSRTGVQELAEPGCDMDSGQCRCKPNIIGRRCDVCAPGYYHYPHCRRCYCHQAGTEVSVCDPVTGQCHCKENVEGVRCDQCRLGTFSLDANNPKGCTRCFCFGATDRCHSAEKYRVEFINMNGWVLLSGDRQEVATSLRLEEQLIHADLKNVPEAYQELYWHAPSSYLGDRVSSYGGYLRYELRSDTRRGDVFIPMEPRPDVILKGNQMSIMFLEGTSPSPGEVHEGRLQLMEGNFRHTQTHNPVSREELMMVLANLEQLQIRALFSQLASSVYLRQVVLETVTEISGGIRASNVELCMCPANYRGDSCQECAPGYYRDTKGLFLGKCVPCHCNGHSDQCLPGSGICINCQHNTEGDRCERCKDGYVGNFSLDGPLQCMGCPCPLSVASNNFAVGCVHKGSTMQCLCKPGYAGAMCERCAPGYYGNPLVIGSSCQPCDCSENTDPNMLFSDCDSLTGACTGCMYNTAGPRCEMCAPGYHGDAVVAKNCTRCNCLPCGTELCDPRTGRCLCKPGVTGQHCDCCQDGYYGYHDCSGCRKCDCDIGSIGTSCHAETGQCSCQPGVTGLRCQQCAPGYWGLSKHGCTKCRCKGGSCDPRSGECTCSNGLTGKQCDTCIEKYEIPVANGPDSMRCEVCDSCVLLLLDDLQKIKASFPIIKNQLINLNASSIAWARLHGLNSSVKNVTVCGGLNVFSPLMDKLGTADLSTASSEEFWQKMAEVQRMLKEMKERDFLSQKGVAEHEHEKAQKLLHRVRSELYSKWESNQDLVKSIKDMLSRYSSELMDLRDALNEAVNKTRQTEDLNSLNRNNLEESQRKSKDCRSEELARLVNAPCGCVLLQEYEKLAARLDGARYPLTERVKKFSPASSKIPVVEEAEEYARLLDELARNLSSSIQGANQDGFIQRAINASNAYASIIKAVKNAERAAKDADEAAGEALTVCNFPSGPHQPLQNPGSSELARGPLLPCQKLGALPSLLLFAEFKILVSFPLTDNTTAVIQNAKAAAAAANDTVAMVEDTLSNMKKNLDEWKDQYGGLKNEELNQAVQDARKSVSNLESTIPLLLEKLSGLENRRNKNSTVSENILRIRQLITQARNAASKVKVPVKFNGTSGIQVRTPSDLQDLSAYTSLKFYIQNPEPKPRQRRQDGADATGDYMGIVLKDRKVQWIYKLGDEEPTYLTVDEEIGEQFAAVSINRILQYGHMSVIVEKQTLHETKGDSVAKGEQGLLNLEPHSVVFYVGGYPTGFTPPVTLRYPSYRGCIEMDTLNEEVVSLYNFQRTFHLDTAADKPCARSKSTGDPWLTDGSYFDGTGYAEIKFESQIGSTKRFEQEIRVVSYNGILFFLEHQGQFLCLAVQDGKLVLYYDFKQVLEMAPPSSDPTSLTIQIFLLTMSSKQRILVRLERQTIFMVEHENSLENAVSYYLGGVPMPLLPKSLKAIFPKGGSIRGCMKGLKALGKYVDLKRMNTVGVSYGCTSDLLVARSVKFHGHGYLTLPLKNVPSLQDFYTGFSFRTSQSHGLLYSHTTTVGLQGNCQVSLKNGRVAVTVLKTELTTKHAYADDASHYVAIYSDATGVRLYIDDQLQGPKQGADRYKRHRRQEEQMLFRLGGLPEAGALSNLTGCIGNVFMPWSCVRGQTVPSQAALKDRRHYQDRTCQLPRHPKATKGAFQFGGASTSHVEFDDIPASFRERSHFSLDVRLNSPSGLLFYVASELQGSFMALFVSSGRFVFLADLNGKKLRVRSKDKYHDGRWHTVSRGTAGLVLGRPIYWSHAFPAHVLCQQVMDQACECLPTLALSLSLQDTSVPSFSGCLKNLKLDGKALDSPSRVFGVTPCYEGMLESGIFFAAEGGYVTLGIRLLAWPWLRLDVRPRSASGLIFHMGTQRKHYLTLYADDTQVPLLAANAGAGEFSTSVARPSLCDGQWHSIAGIATLEAASPSLLTPLPHYVGCIRNLVINQNPVDLLQAGTIRGSVGLTGCPVI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN46</accession>
    <name>A0A452HN46_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">C2H2-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">May be involved in transcriptional regulation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HN46"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018710.1">
        <property type="protein sequence ID" value="ENSGAGP00000016395.1"/>
        <property type="gene ID" value="ENSGAGG00000012269.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001228">
        <property type="term" value="F:DNA-binding transcription activator activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002063">
        <property type="entry name" value="RB associated KRAB zinc finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001155">
        <property type="entry name" value="Zinc finger 30C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000688">
        <property type="entry name" value="zinc finger protein 197 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002343">
        <property type="entry name" value="Zinc finger protein 33A"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002274">
        <property type="entry name" value="Zinc finger protein 432"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000060">
        <property type="entry name" value="zinc finger protein 436"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000340">
        <property type="entry name" value="zinc finger protein 473 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000145">
        <property type="entry name" value="Zinc finger protein 574"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000624">
        <property type="entry name" value="zinc finger protein 697"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="12"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056438">
        <property type="entry name" value="Znf-C2H2_CTCF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24393:SF158">
        <property type="entry name" value="C2H2-TYPE DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24393">
        <property type="entry name" value="ZINC FINGER PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23611">
        <property type="entry name" value="zf-C2H2_16"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="10"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="10" id="KW-0804">Transcription</keyword>
    <keyword evidence="8" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="7" id="KW-0862">Zinc</keyword>
    <keyword evidence="6 12" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="151"/>
            <end position="178"/>
        </location>
    </feature>
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        <location>
            <begin position="179"/>
            <end position="206"/>
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        <location>
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            <end position="252"/>
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            <end position="280"/>
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        <location>
            <begin position="281"/>
            <end position="308"/>
        </location>
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        <location>
            <begin position="310"/>
            <end position="337"/>
        </location>
    </feature>
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        <location>
            <begin position="338"/>
            <end position="365"/>
        </location>
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            <begin position="366"/>
            <end position="393"/>
        </location>
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        <location>
            <begin position="394"/>
            <end position="421"/>
        </location>
    </feature>
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        <location>
            <begin position="422"/>
            <end position="449"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00003767"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
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    </evidence>
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        <source>
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    </evidence>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
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    </evidence>
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        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
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    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016395.1"/>
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    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="530" mass="59750" checksum="0B3CFB3A9189FFAF" modified="2019-05-08" version="1">MRNIHLLPLHTQFPPYSPDAFSPLQNVQGQTPLCLSHGRLFLMLRWLRLGETLAGFTLQPYRVLFRGSEEGAVLSPELQGTALRSCGSQRPESELGATQGCDWCSSNSPNMRSGPETGEGPSPCLVCAPRANGTAPPGVQESSVPKKEKPHKCPECGKAFRRKWDLTKHQRTHSGERPYPCTDCSKSFSHVSNLIKHQRIHTGERPFACNLCGRSFTLKQVLRPFACTDCDKSFKDKQKLTIHQRIHTGERPYVCADCGKSFRQSQRLKTHRRVHTGEKPYLCSDCGRAFSQVSNLYTHWRTHTGEHRFYSCLDCRRSFSQMAGLTRHQKTHLGDRPFPCMDCGKSFCLRQDLMSHQRTHTGERPFLCSQCEKGFSQLAHLAAHQRTHAAERPYPCGTCEKRFKQSQDLRKHQRTHTGERPYPCTECGKCFSHVSNLNAHRKVHMGARPHTCKDCGKGFRPSAGTVLHWGYSALQPHCTGLSKGPQGTPTSSLCTARPKASPCPYCPCWPETRAQPLGNENRDTEKWGRD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN47</accession>
    <name>A0A452HN47_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Collagen alpha-1(XII) chain</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">Type XII collagen interacts with type I collagen-containing fibrils, the COL1 domain could be associated with the surface of the fibrils, and the COL2 and NC3 domains may be localized in the perifibrillar matrix.</text>
    </comment>
    <comment type="subunit">
        <text evidence="14">Trimer of identical chains each containing 190 kDa of non-triple-helical sequences.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="12">Belongs to the fibril-associated collagens with interrupted helices (FACIT) family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018691.1">
        <property type="protein sequence ID" value="ENSGAGP00000016377.1"/>
        <property type="gene ID" value="ENSGAGG00000011657.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005581">
        <property type="term" value="C:collagen trimer"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005614">
        <property type="term" value="C:interstitial matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035987">
        <property type="term" value="P:endodermal cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="entry name" value="FN3"/>
        <property type="match status" value="18"/>
    </dbReference>
    <dbReference type="CDD" id="cd01482">
        <property type="entry name" value="vWA_collagen_alphaI-XII-like"/>
        <property type="match status" value="4"/>
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    <dbReference type="FunFam" id="2.60.40.10:FF:000018">
        <property type="entry name" value="collagen alpha-1(XII) chain isoform X1"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000489">
        <property type="entry name" value="collagen alpha-1(XII) chain isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000554">
        <property type="entry name" value="collagen alpha-1(XII) chain isoform X1"/>
        <property type="match status" value="1"/>
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    <dbReference type="FunFam" id="2.60.40.10:FF:000639">
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        <property type="entry name" value="collagen alpha-1(XII) chain isoform X1"/>
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        <property type="entry name" value="ECM_Assembly_Org"/>
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    </dbReference>
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        <property type="entry name" value="FN3_sf"/>
    </dbReference>
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        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
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        <property type="entry name" value="TSPN-like_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002035">
        <property type="entry name" value="VWF_A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036465">
        <property type="entry name" value="vWFA_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24020">
        <property type="entry name" value="COLLAGEN ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24020:SF17">
        <property type="entry name" value="COLLAGEN ALPHA-1(XII) CHAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01391">
        <property type="entry name" value="Collagen"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="18"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00092">
        <property type="entry name" value="VWA"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00453">
        <property type="entry name" value="VWFADOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="18"/>
    </dbReference>
    <dbReference type="SMART" id="SM00210">
        <property type="entry name" value="TSPN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00327">
        <property type="entry name" value="VWA"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="12"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53300">
        <property type="entry name" value="vWA-like"/>
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    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
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        <property type="match status" value="17"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50234">
        <property type="entry name" value="VWFA"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="8" id="KW-0176">Collagen</keyword>
    <keyword evidence="9" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11" id="KW-0379">Hydroxylation</keyword>
    <keyword evidence="7" id="KW-0654">Proteoglycan</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="16"/>
            <end position="106"/>
        </location>
    </feature>
    <feature type="domain" description="VWFA" evidence="17">
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            <begin position="129"/>
            <end position="305"/>
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    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
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            <begin position="429"/>
            <end position="605"/>
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    <feature type="domain" description="Fibronectin type-III" evidence="18">
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            <begin position="623"/>
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    </feature>
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    </feature>
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    </feature>
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            <begin position="1374"/>
            <end position="1463"/>
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    </feature>
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            <begin position="1464"/>
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            <begin position="1556"/>
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    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
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            <begin position="1646"/>
            <end position="1743"/>
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    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
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            <begin position="1749"/>
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    </feature>
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            <begin position="1931"/>
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        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
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            <begin position="2022"/>
            <end position="2112"/>
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        <location>
            <begin position="2113"/>
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    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="2203"/>
            <end position="2291"/>
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            <begin position="2319"/>
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    </feature>
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    <feature type="region of interest" description="Disordered" evidence="16">
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            <begin position="2006"/>
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    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
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    <feature type="region of interest" description="Disordered" evidence="16">
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            <begin position="2876"/>
            <end position="2992"/>
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    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="2008"/>
            <end position="2020"/>
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    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="16">
        <location>
            <begin position="2769"/>
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    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="2798"/>
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    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="16">
        <location>
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    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="16">
        <location>
            <begin position="2885"/>
            <end position="2894"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="16">
        <location>
            <begin position="2901"/>
            <end position="2910"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="16">
        <location>
            <begin position="2951"/>
            <end position="2964"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004498"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
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    </evidence>
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        <source>
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    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022974"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023119"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023278"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00049648"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00053577"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00064391"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00067989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50234"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016377.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="3010" mass="329339" checksum="987DF2736736796B" modified="2019-05-08" version="1">MPLAFILYWVSAIFNPPSDLNFTIIDEHTVRMSWTRPPDAIEGYKITVVPTTDGPTKEFTLAPSVTDTILSDLTPDTEYVVTITSYDDREESLPVFGQLTIQTGVAVGPEDKKPDQSHIKKCSVSALTDLVFLVDGSWSVGRSNFRYILDFMATLVSAFDIGEEKTRIGVVQYSTDTRTEFNLNQYFRRNDLLNAIKGIPYKGGNTMTGEAIDYLIRNAFTESAGARKGIPKVAIIITDGKSQDEVEIPARELRNVGVEVFSLGIKAADAKELKLMASQPSLKHVFNVANFDGIVDIQNEIISQVCSGVDEQLGELVSGEEAVEPPSNLVATQISSKSIRVTWDPSASQITGYRVQFFPMIAGAKQHVLSVGPQTTALNVRDLSPETEYQINVYAMRGMAASEPITIMEKTQPVKVQVECSRGVDVKADIVLLVDGSYSIGIANFVKVRAFLEVLVKSFEISPQKVQISLVQYSRDPYTEFYLNRYNKIEDIIQAIHTFPYRGGSTNTGKAMTYVREKVFVINRGSRPNVPKVMILITDGKSSDAFKEPAIKLRESDVEIFAVGVKDAVRTELEAIASPPAETHVYTVEDFDAFQRISFELTQSVCLRIEQELAAIRKKSYLPARNLAFSEVTSDSFKVNWSPAGSEVLSYLIKYKVAAGGEEFLVSVPASSTSSVLTHLLPETLYSVSVIAEYEDGDGPSLDGEETTLEVKGAPRNLMITDETTDSFKVGWTPAPGNVLRYRVEYRPVTGRERKHVTVPANERSTTLLNLTPDTRYEVYVIPEYQSGPGNSLNGYAKTEEVLGTPRNLRVSDATTSTMKLSWSAAPGKVQRYLVTYTPVAGGETKEVTIRGDTTTTVLRDLEHGTRYALSVTALYASGAGDALSGQGETPEDRGSPRDLLIRDITDTTMGLSWTAAPGAVRRYRIVWRSLYDGQTGESTVPGNVINTVLENLQPETKYKISVIASYSSGDGAPLEGEATTEVSPNARRVRVDEETENSMRVTWQPAPGKVVSYRVVYRPVSGGRQVVAKVPATSTATVLKRLQPLTKYNISVIPMYKSGEGKHRQGEGTTASPYKPPRNLKTSDSTMSSFRVTWEPAPGEVRGYKITFHPTGEDRLGELIVGPYDNTVVLEELRAGTTYKVNVFGMFEGGESAPLVGQEMTTLSDVTVMPFLARGLECTTRAEADIVLLVDGSWSIGRPNFKTVRNFIARIVEVFDIGPDKVQIALAQYSGDPRTEWHLNTYGSKQSLLDAVANLPYKGGNTLTGMALSFILKDNFKPEVGLRPKARKIGVLITDGKSQDDIVAPSRRLRDEGVELYAIGIKNADENELKQIATDPDDTHVFNVADFSFLTNIVDDLTINLCNSVKGPGGLSSPSNLVTSEVTPRSFRVTWTAPVESVDRYRVEYYPVSGGPPQQFYVNRVETTTVLKNLKPETEYVVKVFSVVEDESSEPLSGTETTLPIPTIRNLNAYDIGSTSMRVRWEPLRGATGYRLLYEPLNATIPAVEKEMRVGSTVNDVQLVDLIPNAAYTLTIHAEFGDLSSDPLTTQEVTLPLAGAKSLRVRDITHSSMNVIWDPAPGKVRKYLLRYKTTLGDEPKEVEVDKSKTSTVLPSLLSQTLYDLQLVAVYDEGESIPVAAKATTLPVPAPVNLRLTEVTKQSFRGTWDHGAPDVALYRITWGPYGGTEKQETILNGDENSLVFENLNPDTLYDVSVTAIYPDESESDDLIGSEFSYILISVTTQNTKSPKSGPRNLQVYNATSNSLTVKWDPASGRVQRYRITYQRTTGDGAEQSTMVGGRQNSVVLQKLQPDTPYTITVSSMYADGEGGRMTGRGKTKPLNTVKNLRVYDPTTSTLSVRWDHAEGSPRQYKVYYVPTTGGAEEMVIILVGNTNYIILRSLQPDTPYKITVVPTYPEGDGGRVSDTGRTLVRGAARNIQVYNPTPNTLNVRWEPAPGPVQQYRVVYAPLVGPRPSESIVVPANTRDVVLERLTPDTPYSVNVVALYADGEGNQSSGQGRTTPRSGPRNLKVFDPTTNSLSVQWDHADGPVQQYRIIYSPTVGDPIDEYTTVPGRRNNVILQPLHSDTPYKITVVAVYDDGDGGQLTGNGRTVGLLPPQNIHISDEWYTRFRVSWDPSPSPVLGYKLAYKPVGSAEPPMEVFVGEVTSYTLHNLSPSTIYDVNVYAQYDSGLSAPLIDQGTTLYLNVTDLTSYKVGWDTFCIRWSPHRSASSYRLKLNPSDGSRGQEITVRGTETSHCFTGLSPDTEYNATVFVQTPNLEGPPISLKERTFIKPTEAPTQPPPPPPPPTIPPARDVCKGAKADIVFLTDASWSIGDDNFNKVVKFVFNTVGAFDLINPAGIQVSFVQYSDDAKSEFKLNTYDDKAQALGALQNIRYKGGNTRTGKALTFINEKVLTWESGMRRGVPKVLVVVTDGRSQDEVKKAAAIIQHSGFSIFVVGVADVDYHELSKIASKPSERHVFIVDDFDAFEKIEENLLTFICETATSTCPLIYLEGYTSPGFKMLESYNLTEKHFASVQGVSLESGSFPSHVAYRLHRNAFISQPASEIHPDGLPHAYTIILLFRLLSETPNEPFAIWQITDRDFKPQVGVILDPSTKLLSFFNKDTRGEVQTVTFDDNEVKKIFYGSFHKVHIVVTSTSVKIYIDCSEVMEKPIKESGNITTDGFEILGKLVKGDRKSATFEIQSFDIVCNAVWTSRDRCCDLPSMRDEAKCPALPNACTCTQDSVGPPGPPGPAKPWFNIFLILCFKGPPGLPGEAGPPGPQGPPGPQGINGLSIPGEPVSNHSVSVSETVWRQRTDTSWERSPKISDHHGPPGMKGEKGDRGDIASQNMMRAVARQVCEQLINGQMNRFNHMLNQIPNDYYANRNQPGPPGPPGPPGSAGARGEPGPGGRPGFPGTPGDQGPPGERGLPGEKGERGIGSQGPRGLPGPPGPPGESRTGPPGSTGSRGPPGPPGRPGNGGIRGPPGPPGYCDSSQCSSIAYNGQGFPGRLLTT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN51</accession>
    <name>A0A452HN51_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Collagen alpha-1(XII) chain</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">Type XII collagen interacts with type I collagen-containing fibrils, the COL1 domain could be associated with the surface of the fibrils, and the COL2 and NC3 domains may be localized in the perifibrillar matrix.</text>
    </comment>
    <comment type="subunit">
        <text evidence="14">Trimer of identical chains each containing 190 kDa of non-triple-helical sequences.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="12">Belongs to the fibril-associated collagens with interrupted helices (FACIT) family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018709.1">
        <property type="protein sequence ID" value="ENSGAGP00000016394.1"/>
        <property type="gene ID" value="ENSGAGG00000011657.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000018719.1">
        <property type="protein sequence ID" value="ENSGAGP00000016404.1"/>
        <property type="gene ID" value="ENSGAGG00000011657.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005581">
        <property type="term" value="C:collagen trimer"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005614">
        <property type="term" value="C:interstitial matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009888">
        <property type="term" value="P:tissue development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="17"/>
    </dbReference>
    <dbReference type="CDD" id="cd01482">
        <property type="entry name" value="vWA_collagen_alphaI-XII-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000018">
        <property type="entry name" value="collagen alpha-1(XII) chain isoform X1"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000489">
        <property type="entry name" value="collagen alpha-1(XII) chain isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000639">
        <property type="entry name" value="collagen alpha-1(XII) chain isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000816">
        <property type="entry name" value="collagen alpha-1(XII) chain isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.200:FF:000008">
        <property type="entry name" value="Collagen type XII alpha 1 chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000121">
        <property type="entry name" value="Collagen type XII alpha 1 chain"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000234">
        <property type="entry name" value="Collagen, type XII, alpha 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000480">
        <property type="entry name" value="Collagen, type XII, alpha 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.410:FF:000001">
        <property type="entry name" value="Collagen, type XII, alpha 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.200">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="17"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.410">
        <property type="entry name" value="von Willebrand factor, type A domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008160">
        <property type="entry name" value="Collagen"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050991">
        <property type="entry name" value="ECM_Regulatory_Proteins"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048287">
        <property type="entry name" value="TSPN-like_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002035">
        <property type="entry name" value="VWF_A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036465">
        <property type="entry name" value="vWFA_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46708:SF7">
        <property type="entry name" value="FIBRONECTIN TYPE-III DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46708">
        <property type="entry name" value="TENASCIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01391">
        <property type="entry name" value="Collagen"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="17"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00092">
        <property type="entry name" value="VWA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00453">
        <property type="entry name" value="VWFADOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="17"/>
    </dbReference>
    <dbReference type="SMART" id="SM00210">
        <property type="entry name" value="TSPN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00327">
        <property type="entry name" value="VWA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53300">
        <property type="entry name" value="vWA-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="16"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50234">
        <property type="entry name" value="VWFA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="8" id="KW-0176">Collagen</keyword>
    <keyword evidence="9" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11" id="KW-0379">Hydroxylation</keyword>
    <keyword evidence="7" id="KW-0654">Proteoglycan</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="1"/>
            <end position="84"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="87"/>
            <end position="181"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="182"/>
            <end position="267"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="269"/>
            <end position="358"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="360"/>
            <end position="450"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="451"/>
            <end position="539"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="541"/>
            <end position="630"/>
        </location>
    </feature>
    <feature type="domain" description="VWFA" evidence="17">
        <location>
            <begin position="650"/>
            <end position="822"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="838"/>
            <end position="927"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="928"/>
            <end position="1019"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="1020"/>
            <end position="1109"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="1110"/>
            <end position="1200"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="1210"/>
            <end position="1304"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="1391"/>
            <end position="1481"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="1482"/>
            <end position="1572"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="1573"/>
            <end position="1662"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="1663"/>
            <end position="1751"/>
        </location>
    </feature>
    <feature type="domain" description="VWFA" evidence="17">
        <location>
            <begin position="1779"/>
            <end position="1952"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="163"/>
            <end position="182"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="524"/>
            <end position="551"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1466"/>
            <end position="1485"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="2203"/>
            <end position="2352"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="2388"/>
            <end position="2559"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1468"/>
            <end position="1480"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="16">
        <location>
            <begin position="2232"/>
            <end position="2249"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="16">
        <location>
            <begin position="2273"/>
            <end position="2284"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="16">
        <location>
            <begin position="2312"/>
            <end position="2330"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="16">
        <location>
            <begin position="2397"/>
            <end position="2406"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="16">
        <location>
            <begin position="2413"/>
            <end position="2422"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="16">
        <location>
            <begin position="2463"/>
            <end position="2476"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="2501"/>
            <end position="2510"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022974"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023119"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023278"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00049648"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00053577"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00064391"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00067989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50234"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016394.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2559" mass="276820" checksum="C9FC669896944FDF" modified="2019-05-08" version="1">MITDETTDSFKVGWTPAPGNVLRYRVEYRPVTGRERKHVTVPANERSTTLLNLTPDTRYEVYVIPEYQSGPGNSLNGYAKTEEVLGTPRNLRVSDATTSTMKLSWSAAPGKVQRYLVTYTPVAGGETKEVTIRGDTTTTVLRDLEHGTRYALSVTALYASGAGDALSGQGETPEARGSPRNLRVSDATTSTMKLSWSAAPGKVERYRITYTPVAGGDTKEVTTRGDTTTTVLRDLEHGTRYALSVTALYASGAGDALPGQGETLEARGSPRNLRVSDATTSTMKLSWSAAPGKVQRYRITYTPVAGGEAKEVTIRGDTTTTVLRDLEQGTRYTLSVTALYASGAGDALPGQGETLEDRGSPRDLLIRDITDTTMGLSWTAAPGAVRRYRIVWRSLYDGQTGESTVPGNVINTVLENLQPETKYKISVIASYSSGDGAPLEGEATTEVSPNARRVRVDEETENSMRVTWQPAPGKVVSYRVVYRPVSGGRQVVAKVPATSTATVLKRLQPLTKYNISVIPMYKSGEGKHRQGEGTTASPYKPPRNLKTSDSTMSSFRVTWEPAPGEVRGYKITFHPTGEDRLGELIVGPYDNTVVLEELRAGTTYKVNVFGMFEGGESAPLVGQEMTTLSDVTVMPFLARGLECTTRAEADIVLLVDGSWSIGRPNFKTVRNFIARIVEVFDIGPDKVQIALAQYSGDPRTEWHLNTYGSKQSLLDAVANLPYKGGNTLTGMALSFILKDNFKPEVGLRPKARKIGVLITDGKSQDDIVAPSRRLRDEGVELYAIGIKNADENELKQIATDPDDTHVFNVADFSFLTNIVDDLTINLCNSVKGPGGLSSPSNLVTSEVTPRSFRVTWTAPVESVDRYRVEYYPVSGGPPQQFYVNRVETTTVLKNLKPETEYVVKVFSVVEDESSEPLSGTETTLPIPTIRNLNAYDIGSTSMRVRWEPLRGATGYRLLYEPLNATIPAVEKEMRVGSTVNDVQLVDLIPNAAYTLTIHAEFGDLSSDPLTTQEVTLPLAGAKSLRVRDITHSSMNVIWDPAPGKVRKYLLRYKTTLGDEPKEVEVDKSKTSTVLPSLLSQTLYDLQLVAVYDEGESIPVAAKATTLPVPAPVNLRLTEVTKQSFRGTWDHGAPDVALYRITWGPYGGTEKQETILNGDENSLVFENLNPDTLYDVSVTAIYPDESESDDLIGSERTLPAIRPTTQAPKSGPRNLQVYNATSNSLTVKWDPASGRVQRYRITYQRTTGDGAEQSTMVGGRQNSVVLQKLQPDTPYTITVSSMYADGEGGRMTGRGKTKPLNTVKNLRVYDPTTSTLSVRWDHAEGSPRQYKVYYVPTTGGAEEMTTVPGNTNYIILRSLQPDTPYKITVVPTYPEGDGGRVSDTGRTLVRGAARNIQVYNPTPNTLNVRWEPAPGPVQQYRVVYAPLVGPRPSESIVVPANTRDVVLERLTPDTPYSVNVVALYADGEGNQSSGQGRTTPRSGPRNLKVFDPTTNSLSVQWDHADGPVQQYRIIYSPTVGDPIDEYTTVPGRRNNVILQPLHSDTPYKITVVAVYDDGDGGQLTGNGRTVGLLPPQNIHISDEWYTRFRVSWDPSPSPVLGYKLAYKPVGSAEPPMEVFVGEVTSYTLHNLSPSTIYDVNVYAQYDSGLSAPLIDQGTTLYLNVTDLTSYKVGWDTFCIRWSPHRSASSYRLKLNPSDGSRGQEITVRGTETSHCFTGLSPDTEYNATVFVQTPNLEGPPISLKERTFIKPTEAPTQPPPPPPPPTIPPARDVCKGAKADIVFLTDASWSIGDDNFNKVVKFVFNTVGAFDLINPAGIQVSFVQYSDDAKSEFKLNTYDDKAQALGALQNIRYKGGNTRTGKALTFINEKVLTWESGMRRGVPKVLVVVTDGRSQDEVKKAAAIIQHSGFSIFVVGVADVDYHELSKIASKPSERHVFIVDDFDAFEKIEENLLTFICETATSTCPLIYLEGYTSPGFKMLESYNLTEKHFASVQGVSLESGSFPSHVAYRLHRNAFISQPASEIHPDGLPHAYTIILLFRLLSETPNEPFAIWQITDRDFKPQVGVILDPSTKLLSFFNKDTRGEVQTVTFDDNEVKKIFYGSFHKVHIVVTSTSVKIYIDCSEVMEKPIKESGNITTDGFEILGKLVKGDRKSATFEIQSFDIVCNAVWTSRDRCCDLPSMRDEAKCPALPNACTCTQDSVGPPGPPGPAGGPGTKGPRGERGLTGTAGPPGLPGEAGPPGPQGPPGPQGINGLSIPGEPGRQGMKGDAGEPGLPGRSGTPGLSGPPGPVGPPGDRGFTGKDGATGPRGPPGSAGTPGVPGVAGPSGKPGKPGDRGPQGPPGMKGEKGDRGDIASQNMMRAVARQVCEQLINGQMNRFNHMLNQIPNDYYANRNQPGPPGPPGPPGSAGARGEPGPGGRPGFPGTPGDQGPPGERGLPGEKGERGIGSQGPRGLPGPPGPPGESRTGPPGSTGSRGPPGPPGRPGNGGIRGPPGPPGYCDSSQCSSIAYNGQGFPESYVPESGPYQPESEPYIVPMESERREDEYEDYEVEMHSPGLP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN52</accession>
    <name>A0A452HN52_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Collagen alpha-1(XII) chain</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">Type XII collagen interacts with type I collagen-containing fibrils, the COL1 domain could be associated with the surface of the fibrils, and the COL2 and NC3 domains may be localized in the perifibrillar matrix.</text>
    </comment>
    <comment type="subunit">
        <text evidence="14">Trimer of identical chains each containing 190 kDa of non-triple-helical sequences.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="12">Belongs to the fibril-associated collagens with interrupted helices (FACIT) family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018696.1">
        <property type="protein sequence ID" value="ENSGAGP00000016382.1"/>
        <property type="gene ID" value="ENSGAGG00000011657.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005581">
        <property type="term" value="C:collagen trimer"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005614">
        <property type="term" value="C:interstitial matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035987">
        <property type="term" value="P:endodermal cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="18"/>
    </dbReference>
    <dbReference type="CDD" id="cd01482">
        <property type="entry name" value="vWA_collagen_alphaI-XII-like"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000018">
        <property type="entry name" value="collagen alpha-1(XII) chain isoform X1"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000489">
        <property type="entry name" value="collagen alpha-1(XII) chain isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000554">
        <property type="entry name" value="collagen alpha-1(XII) chain isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000639">
        <property type="entry name" value="collagen alpha-1(XII) chain isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000704">
        <property type="entry name" value="collagen alpha-1(XII) chain isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000816">
        <property type="entry name" value="collagen alpha-1(XII) chain isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.200:FF:000008">
        <property type="entry name" value="Collagen type XII alpha 1 chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000121">
        <property type="entry name" value="Collagen type XII alpha 1 chain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000234">
        <property type="entry name" value="Collagen, type XII, alpha 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000480">
        <property type="entry name" value="Collagen, type XII, alpha 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.410:FF:000001">
        <property type="entry name" value="Collagen, type XII, alpha 1"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.200">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.320">
        <property type="entry name" value="6-Phosphogluconate Dehydrogenase, domain 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="18"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.410">
        <property type="entry name" value="von Willebrand factor, type A domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050525">
        <property type="entry name" value="ECM_Assembly_Org"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048287">
        <property type="entry name" value="TSPN-like_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002035">
        <property type="entry name" value="VWF_A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036465">
        <property type="entry name" value="vWFA_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24020">
        <property type="entry name" value="COLLAGEN ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24020:SF17">
        <property type="entry name" value="COLLAGEN ALPHA-1(XII) CHAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="18"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00092">
        <property type="entry name" value="VWA"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00453">
        <property type="entry name" value="VWFADOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="18"/>
    </dbReference>
    <dbReference type="SMART" id="SM00210">
        <property type="entry name" value="TSPN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00327">
        <property type="entry name" value="VWA"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="12"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53300">
        <property type="entry name" value="vWA-like"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="17"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50234">
        <property type="entry name" value="VWFA"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="8" id="KW-0176">Collagen</keyword>
    <keyword evidence="9" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11" id="KW-0379">Hydroxylation</keyword>
    <keyword evidence="7" id="KW-0654">Proteoglycan</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="26"/>
            <end position="116"/>
        </location>
    </feature>
    <feature type="domain" description="VWFA" evidence="17">
        <location>
            <begin position="139"/>
            <end position="315"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="335"/>
            <end position="424"/>
        </location>
    </feature>
    <feature type="domain" description="VWFA" evidence="17">
        <location>
            <begin position="439"/>
            <end position="615"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="633"/>
            <end position="722"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="724"/>
            <end position="818"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="819"/>
            <end position="903"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="906"/>
            <end position="996"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="997"/>
            <end position="1085"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="1087"/>
            <end position="1176"/>
        </location>
    </feature>
    <feature type="domain" description="VWFA" evidence="17">
        <location>
            <begin position="1196"/>
            <end position="1368"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="1384"/>
            <end position="1473"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="1474"/>
            <end position="1565"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="1566"/>
            <end position="1655"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="1656"/>
            <end position="1753"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="1759"/>
            <end position="1853"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="1941"/>
            <end position="2031"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="2032"/>
            <end position="2122"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="2123"/>
            <end position="2212"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="2213"/>
            <end position="2301"/>
        </location>
    </feature>
    <feature type="domain" description="VWFA" evidence="17">
        <location>
            <begin position="2329"/>
            <end position="2502"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1070"/>
            <end position="1097"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="2016"/>
            <end position="2035"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="2779"/>
            <end position="2850"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="2886"/>
            <end position="2977"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="2018"/>
            <end position="2030"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="16">
        <location>
            <begin position="2779"/>
            <end position="2794"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="2808"/>
            <end position="2818"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="16">
        <location>
            <begin position="2819"/>
            <end position="2850"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="16">
        <location>
            <begin position="2895"/>
            <end position="2904"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="16">
        <location>
            <begin position="2911"/>
            <end position="2920"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="16">
        <location>
            <begin position="2937"/>
            <end position="2951"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022974"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023119"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023278"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00049648"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00053577"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00064391"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00067989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50234"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016382.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2997" mass="328099" checksum="AF64148CD42DA55F" modified="2019-05-08" version="1">MLYAYIGLCRIRALGAQFLGPSIVNPPSDLNFTIIDEHTVRMSWTRPPDAIEGYKITVVPTTDGPTKEFTLAPSVTDTILSDLTPDTEYVVTITSYDDREESLPVFGQLTIQTGVAVGPEDKKPDQSHIKKCSVSALTDLVFLVDGSWSVGRSNFRYILDFMATLVSAFDIGEEKTRIGVVQYSTDTRTEFNLNQYFRRNDLLNAIKGIPYKGGNTMTGEAIDYLIRNAFTESAGARKGIPKVAIIITDGKSQDEVEIPARELRNVGVEVFSLGIKAADAKELKLMASQPSLKHVFNVANFDGIVDIQNEIISQVCSGVDEQLGELVSGEEAVEPPSNLVATQISSKSIRVTWDPSASQITGYRVQFFPMIAGAKQHVLSVGPQTTALNVRDLSPETEYQINVYAMRGMAASEPITIMEKTQPVKVQVECSRGVDVKADIVLLVDGSYSIGIANFVKVRAFLEVLVKSFEISPQKVQISLVQYSRDPYTEFYLNRYNKIEDIIQAIHTFPYRGGSTNTGKAMTYVREKVFVINRGSRPNVPKVMILITDGKSSDAFKEPAIKLRESDVEIFAVGVKDAVRTELEAIASPPAETHVYTVEDFDAFQRISFELTQSVCLRIEQELAAIRKKSYLPARNLAFSEVTSDSFKVNWSPAGSEVLSYLIKYKVAAGGEEFLVSVPASSTSSVLTHLLPETLYSVSVIAEYEDGDGPSLDGEETTLEVKGAPRNLMITDETTDSFKVGWTPAPGNVLRYRVEYRPVTGRERKHVTVPANERSTTLLNLTPDTRYEVYVIPEYQSGPGNSLNGYAKTEEVLGTPRNLRVSDATTSTMKLSWSAAPGKVQRYLVTYTPVAGGETKEVTIRGDTTTTVLRDLEHGTRYALSVTALYASGAGDALSGQGETPEDRGSPRDLLIRDITDTTMGLSWTAAPGAVRRYRIVWRSLYDGQTGESTVPGNVINTVLENLQPETKYKISVIASYSSGDGAPLEGEATTEVSPNARRVRVDEETENSMRVTWQPAPGKVVSYRVVYRPVSGGRQVVAKVPATSTATVLKRLQPLTKYNISVIPMYKSGEGKHRQGEGTTASPYKPPRNLKTSDSTMSSFRVTWEPAPGEVRGYKITFHPTGEDRLGELIVGPYDNTVVLEELRAGTTYKVNVFGMFEGGESAPLVGQEMTTLSDVTVMPFLARGLECTTRAEADIVLLVDGSWSIGRPNFKTVRNFIARIVEVFDIGPDKVQIALAQYSGDPRTEWHLNTYGSKQSLLDAVANLPYKGGNTLTGMALSFILKDNFKPEVGLRPKARKIGVLITDGKSQDDIVAPSRRLRDEGVELYAIGIKNADENELKQIATDPDDTHVFNVADFSFLTNIVDDLTINLCNSVKGPGGLSSPSNLVTSEVTPRSFRVTWTAPVESVDRYRVEYYPVSGGPPQQFYVNRVETTTVLKNLKPETEYVVKVFSVVEDESSEPLSGTETTLPIPTIRNLNAYDIGSTSMRVRWEPLRGATGYRLLYEPLNATIPAVEKEMRVGSTVNDVQLVDLIPNAAYTLTIHAEFGDLSSDPLTTQEVTLPLAGAKSLRVRDITHSSMNVIWDPAPGKVRKYLLRYKTTLGDEPKEVEVDKSKTSTVLPSLLSQTLYDLQLVAVYDEGESIPVAAKATTLPVPAPVNLRLTEVTKQSFRGTWDHGAPDVALYRITWGPYGGTEKQETILNGDENSLVFENLNPDTLYDVSVTAIYPDESESDDLIGSEFSYILISVTTQNTKSPKSGPRNLQVYNATSNSLTVKWDPASGRVQRYRITYQRTTGDGAEQSTMVGGRQNSVVLQKLQPDTPYTITVSSMYADGEGGRMTGRGKTKPLNTVKNLRVYDPTTSTLSVRWDHAEGSPRQYKVYYVPTTGGAEEMVIILVGNTNYIILRSLQPDTPYKITVVPTYPEGDGGRVSDTGRTLVRGAARNIQVYNPTPNTLNVRWEPAPGPVQQYRVVYAPLVGPRPSESIVVPANTRDVVLERLTPDTPYSVNVVALYADGEGNQSSGQGRTTPRSGPRNLKVFDPTTNSLSVQWDHADGPVQQYRIIYSPTVGDPIDEYTTVPGRRNNVILQPLHSDTPYKITVVAVYDDGDGGQLTGNGRTVGLLPPQNIHISDEWYTRFRVSWDPSPSPVLGYKLAYKPVGSAEPPMEVFVGEVTSYTLHNLSPSTIYDVNVYAQYDSGLSAPLIDQGTTLYLNVTDLTSYKVGWDTFCIRWSPHRSASSYRLKLNPSDGSRGQEITVRGTETSHCFTGLSPDTEYNATVFVQTPNLEGPPISLKERTFIKPTEAPTQPPPPPPPPTIPPARDVCKGAKADIVFLTDASWSIGDDNFNKVVKFVFNTVGAFDLINPAGIQVSFVQYSDDAKSEFKLNTYDDKAQALGALQNIRYKGGNTRTGKALTFINEKVLTWESGMRRGVPKVLVVVTDGRSQDEVKKAAAIIQHSGFSIFVVGVADVDYHELSKIASKPSERHVFIVDDFDAFEKIEENLLTFICETATSTCPLIYLEGYTSPGFKMLESYNLTEKHFASVQGVSLESGSFPSHVAYRLHRNAFISQPASEIHPDGLPHAYTIILLFRLLSETPNEPFAIWQITDRDFKPQVGVILDPSTKLLSFFNKDTRGEVQTVTFDDNEVKKIFYGSFHKVHIVVTSTSVKIYIDCSEVMEKPIKESGNITTDGFEILGKLVKGDRKSATFEIQSFDIVCNAVWTSRDRCCDLPSMRDEAKCPALPNACTCTQDSVGPPGPPGPAKPWFNIFLILCFKGPPGLPGEAGPPGPQGPPGPQGINGLSIPGEPVSNHSVSVSETVWRQRTDTSWERSPKISDHHGPPGMKGEKGDRGDIASQNMMRAVARQVCEQLINGQMNRFNHMLNQIPNDYYANRNQPGPPGPPGPPGSAGARGEPGPGGRPGFPGTPGDQGPPGERGPPGESRTGPPGSTGSRGPPGPPGRPGNGGIRGPPGPPGYCDSSQCSSIAYNGQGFPGRLLTT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN53</accession>
    <name>A0A452HN53_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">phospholipase A2</fullName>
            <ecNumber evidence="1">3.1.1.4</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="6">
            <text>a 1,2-diacyl-sn-glycero-3-phosphocholine + H2O = a 1-acyl-sn-glycero-3-phosphocholine + a fatty acid + H(+)</text>
            <dbReference type="Rhea" id="RHEA:15801"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:28868"/>
            <dbReference type="ChEBI" id="CHEBI:57643"/>
            <dbReference type="ChEBI" id="CHEBI:58168"/>
            <dbReference type="EC" id="3.1.1.4"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="6">
            <dbReference type="Rhea" id="RHEA:15802"/>
        </physiologicalReaction>
    </comment>
    <dbReference type="EC" id="3.1.1.4" evidence="1"/>
    <dbReference type="AlphaFoldDB" id="A0A452HN53"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018734.1">
        <property type="protein sequence ID" value="ENSGAGP00000016418.1"/>
        <property type="gene ID" value="ENSGAGG00000011814.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0047499">
        <property type="term" value="F:calcium-independent phospholipase A2 activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0052816">
        <property type="term" value="F:long-chain fatty acyl-CoA hydrolase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035965">
        <property type="term" value="P:cardiolipin acyl-chain remodeling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016042">
        <property type="term" value="P:lipid catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:2000304">
        <property type="term" value="P:positive regulation of ceramide biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07212">
        <property type="entry name" value="Pat_PNPLA9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000338">
        <property type="entry name" value="85/88 kDa calcium-independent phospholipase A2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.1090.10">
        <property type="entry name" value="Cytosolic phospholipase A2 catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016035">
        <property type="entry name" value="Acyl_Trfase/lysoPLipase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047148">
        <property type="entry name" value="PLPL9"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002641">
        <property type="entry name" value="PNPLA_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24139:SF34">
        <property type="entry name" value="85_88 KDA CALCIUM-INDEPENDENT PHOSPHOLIPASE A2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24139">
        <property type="entry name" value="CALCIUM-INDEPENDENT PHOSPHOLIPASE A2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01734">
        <property type="entry name" value="Patatin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01415">
        <property type="entry name" value="ANKYRIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52151">
        <property type="entry name" value="FabD/lysophospholipase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51635">
        <property type="entry name" value="PNPLA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4 7" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="3 8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="8" id="KW-0442">Lipid degradation</keyword>
    <keyword evidence="5 8" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <feature type="repeat" description="ANK" evidence="7">
        <location>
            <begin position="151"/>
            <end position="183"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="7">
        <location>
            <begin position="218"/>
            <end position="250"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="7">
        <location>
            <begin position="348"/>
            <end position="380"/>
        </location>
    </feature>
    <feature type="domain" description="PNPLA" evidence="10">
        <location>
            <begin position="476"/>
            <end position="660"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="415"/>
            <end position="444"/>
        </location>
    </feature>
    <feature type="short sequence motif" description="GXGXXG" evidence="8">
        <location>
            <begin position="480"/>
            <end position="485"/>
        </location>
    </feature>
    <feature type="short sequence motif" description="GXSXG" evidence="8">
        <location>
            <begin position="512"/>
            <end position="516"/>
        </location>
    </feature>
    <feature type="short sequence motif" description="DGA/G" evidence="8">
        <location>
            <begin position="647"/>
            <end position="649"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="427"/>
            <end position="444"/>
        </location>
    </feature>
    <feature type="active site" description="Nucleophile" evidence="8">
        <location>
            <position position="514"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="8">
        <location>
            <position position="647"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00013278"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023422"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01161"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS51635"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016418.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="801" mass="89937" checksum="73C544EC441F5FA8" modified="2019-05-08" version="1">MQFFGRLVNTINSYSNLFSNPYRVKEVPLAEYVAHTRLRENDRVILYKSKNARSWDCLLVNPQSHLVAFRLFQLDNETDAIERFEQYARRLRPFYESSPQPLQLEVLQQLSDCFRNHPYWSLAHVAVETELPGSFQHNRILSCVNSTECEDGCTPLQLACRKGNMECLCELVECHARLDVTDKNGETVFHYAVRGSNPEIVKLLGKKPSVGLNHLSNQGQTPLHLACQLGKEDVVLALLKCHAKCNVMGTLGYPIHTAVKYSQKGCVQALLEVDVNQVQFKDPRYGAAVLHWAKNAEMTRMLIEYGCDVNSVSKTADMALHIMVKRGRFDSAMVLLTHGALTNAKGQDGNTPLHLAMKQDHLETIKALIVFGADVEIPNEFGETPGLVAARTSKGANRKVLLGLLQMVGTERCHPPIPESPHMASAESPSLSTSPERPQLPRNNGHSLGKHTYVLMHNLAAIFLFSWCLSRYDRLLCLDGGGIRGLVLIQLLIAVEKAAGRPIRELFDWIAGTSTGGILALAIVHGKPVDYMRCLYFRMKNEVFRGSRPYESEPLDEFLKREFGEHTKMTDVQKPKVIVTGTLCDRQPAELHLFRNYKAPATKNKPEVKTTDPFKPLTKPEDQLVWRAARCSGAAPTYFRPIGRFLDGGLLANNPTLDAMTEIHEYNKSLSQKGQSQQVRKLGVVVSLGTGRPPQVPVRSVDVFRPSNPWELAKTVFGAKELGKMVVDCCTDADGPAVDRARAWCEMTDTRYFRFSPQLQSDVMLDEVNDAILVNALWDTQLYICQQQEQFKQLVQELLTR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN54</accession>
    <name>A0A452HN54_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Laminin subunit alpha-5</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="14">Laminin-10 subunit alpha</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="16">Laminin-11 subunit alpha</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="15">Laminin-15 subunit alpha</fullName>
        </alternativeName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="12">Laminin is a complex glycoprotein, consisting of three different polypeptide chains (alpha, beta, gamma), which are bound to each other by disulfide bonds into a cross-shaped molecule comprising one long and three short arms with globules at each end. Alpha-5 is a subunit of laminin-10 (laminin-511), laminin-11 (laminin-521) and laminin-15 (laminin-523).</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
            <location evidence="1">Basement membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="17">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018721.1">
        <property type="protein sequence ID" value="ENSGAGP00000016405.1"/>
        <property type="gene ID" value="ENSGAGG00000012074.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043259">
        <property type="term" value="C:laminin-10 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045202">
        <property type="term" value="C:synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005201">
        <property type="term" value="F:extracellular matrix structural constituent"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009887">
        <property type="term" value="P:animal organ morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007411">
        <property type="term" value="P:axon guidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048732">
        <property type="term" value="P:gland development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002009">
        <property type="term" value="P:morphogenesis of an epithelium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030155">
        <property type="term" value="P:regulation of cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030334">
        <property type="term" value="P:regulation of cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045995">
        <property type="term" value="P:regulation of embryonic development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060541">
        <property type="term" value="P:respiratory system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035239">
        <property type="term" value="P:tube morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd00055">
        <property type="entry name" value="EGF_Lam"/>
        <property type="match status" value="18"/>
    </dbReference>
    <dbReference type="CDD" id="cd00110">
        <property type="entry name" value="LamG"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000074">
        <property type="entry name" value="Laminin subunit alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000083">
        <property type="entry name" value="Laminin subunit alpha"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000090">
        <property type="entry name" value="laminin subunit alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000069">
        <property type="entry name" value="Laminin subunit alpha 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000454">
        <property type="entry name" value="Laminin subunit alpha 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000033">
        <property type="entry name" value="Laminin subunit alpha 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000034">
        <property type="entry name" value="Laminin subunit alpha 3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000084">
        <property type="entry name" value="Laminin subunit alpha 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000051">
        <property type="entry name" value="Laminin subunit alpha 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000209">
        <property type="entry name" value="Laminin subunit alpha 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000405">
        <property type="entry name" value="Laminin subunit alpha 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000430">
        <property type="entry name" value="Laminin subunit alpha 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000437">
        <property type="entry name" value="Laminin subunit alpha 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000467">
        <property type="entry name" value="Laminin subunit alpha 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.200:FF:000151">
        <property type="entry name" value="Laminin subunit alpha 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.260:FF:000444">
        <property type="entry name" value="Laminin subunit alpha 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.200">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.260">
        <property type="entry name" value="Galactose-binding domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="18"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050440">
        <property type="entry name" value="Laminin/Netrin_ECM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009254">
        <property type="entry name" value="Laminin_aI"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010307">
        <property type="entry name" value="Laminin_dom_II"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001791">
        <property type="entry name" value="Laminin_G"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000034">
        <property type="entry name" value="Laminin_IV"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008211">
        <property type="entry name" value="Laminin_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002049">
        <property type="entry name" value="LE_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056863">
        <property type="entry name" value="LMN_ATRN_NET-like_EGF"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10574:SF406">
        <property type="entry name" value="LAMININ SUBUNIT ALPHA 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10574">
        <property type="entry name" value="NETRIN/LAMININ-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00053">
        <property type="entry name" value="EGF_laminin"/>
        <property type="match status" value="16"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24973">
        <property type="entry name" value="EGF_LMN_ATRN"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00052">
        <property type="entry name" value="Laminin_B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00054">
        <property type="entry name" value="Laminin_G_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02210">
        <property type="entry name" value="Laminin_G_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06008">
        <property type="entry name" value="Laminin_I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06009">
        <property type="entry name" value="Laminin_II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00055">
        <property type="entry name" value="Laminin_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00011">
        <property type="entry name" value="EGFLAMININ"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="SMART" id="SM00180">
        <property type="entry name" value="EGF_Lam"/>
        <property type="match status" value="20"/>
    </dbReference>
    <dbReference type="SMART" id="SM00281">
        <property type="entry name" value="LamB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00282">
        <property type="entry name" value="LamG"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00136">
        <property type="entry name" value="LamNT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="18"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01248">
        <property type="entry name" value="EGF_LAM_1"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50027">
        <property type="entry name" value="EGF_LAM_2"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50025">
        <property type="entry name" value="LAM_G_DOMAIN"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51115">
        <property type="entry name" value="LAMININ_IVA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51117">
        <property type="entry name" value="LAMININ_NTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0084">Basement membrane</keyword>
    <keyword evidence="7" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="8 18" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="9 17" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11 17" id="KW-0424">Laminin EGF-like domain</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <feature type="domain" description="Laminin N-terminal" evidence="22">
        <location>
            <begin position="1"/>
            <end position="135"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="20">
        <location>
            <begin position="265"/>
            <end position="309"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="20">
        <location>
            <begin position="331"/>
            <end position="377"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="20">
        <location>
            <begin position="378"/>
            <end position="423"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="20">
        <location>
            <begin position="424"/>
            <end position="473"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="20">
        <location>
            <begin position="514"/>
            <end position="559"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="20">
        <location>
            <begin position="611"/>
            <end position="663"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="20">
        <location>
            <begin position="1244"/>
            <end position="1289"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="20">
        <location>
            <begin position="1334"/>
            <end position="1382"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="20">
        <location>
            <begin position="1383"/>
            <end position="1433"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin IV type A" evidence="21">
        <location>
            <begin position="1454"/>
            <end position="1632"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="20">
        <location>
            <begin position="1666"/>
            <end position="1715"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="20">
        <location>
            <begin position="1716"/>
            <end position="1771"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="20">
        <location>
            <begin position="1772"/>
            <end position="1825"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="20">
        <location>
            <begin position="1826"/>
            <end position="1872"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="20">
        <location>
            <begin position="1873"/>
            <end position="1919"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin G" evidence="19">
        <location>
            <begin position="2452"/>
            <end position="2650"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin G" evidence="19">
        <location>
            <begin position="2664"/>
            <end position="2838"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin G" evidence="19">
        <location>
            <begin position="2845"/>
            <end position="3003"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin G" evidence="19">
        <location>
            <begin position="3159"/>
            <end position="3304"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="18">
        <location>
            <begin position="2160"/>
            <end position="2187"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="18">
        <location>
            <begin position="2347"/>
            <end position="2407"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="285"/>
            <end position="294"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="331"/>
            <end position="343"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="354"/>
            <end position="363"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="378"/>
            <end position="390"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="380"/>
            <end position="397"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="399"/>
            <end position="408"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="424"/>
            <end position="436"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="444"/>
            <end position="453"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="514"/>
            <end position="526"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="516"/>
            <end position="533"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="535"/>
            <end position="544"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="634"/>
            <end position="643"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="1244"/>
            <end position="1256"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="1246"/>
            <end position="1263"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="1265"/>
            <end position="1274"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="1358"/>
            <end position="1367"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="1383"/>
            <end position="1395"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="1385"/>
            <end position="1402"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="1404"/>
            <end position="1413"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="1685"/>
            <end position="1694"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="1742"/>
            <end position="1751"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="1797"/>
            <end position="1806"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="1809"/>
            <end position="1823"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="1845"/>
            <end position="1854"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="1893"/>
            <end position="1902"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004302"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022869"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023292"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00063580"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00072596"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00077092"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00078366"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00081617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00460"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50025"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50027"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS51115"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS51117"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016405.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="3327" mass="364731" checksum="E468A06DD1321A7F" modified="2019-05-08" version="1">PRPDLWVLERSTDFGLTYEPWQYFASSRRDCIEKFGPNTVKRITKDDDTICTTEYSRIVPLENGEIVVSLVNGRPGAMNFSYSPLLRNFTKATNIRLRFLQTNTLLGHLMGKALRDPTVTRRYYYSIKDISIGGRCVCHGHADVCDAKDPTDPYRLQCDCQHNTCGGSCDRCCPGFNQLPWKPATTDSANECQPCNCNGHAYDCYYDPEVDRHKASKNYDNQYKGGGVCIECQHHTTGINCERCIPGYYKSPDHSIDSPYICYRCNCDSEFTDGTCEDLTGRCYCKPNYTGELCNACAEGYTDFPHCYPVPAFSHNDTGEQVLPAGQIINCDCNAAGTEGNACRKDSQRGMCVCKPNFQGAQCDQCAPGHYGPGCQPCQCSGPGQYDGTCDSETGQCLCRSGFEGYSCNQCAPGYFSYPLCQLCGCSAVGTLPGGCDSGGRCFCRPEFDGPHCDQCSMGYHSYPRCQACSCDPRGSVDNNCSPAGQCRCHSNYAGHTCSQCAPGFYGYPSCTSCQCSTEGSLHGTCDQETGQCSCRPKVAGLRCDTCMPGAYGFPHCEVGSCNPAGLATADPSLSPGSCVCRAYVEGPACDRCKPLYWNLTPDNPYGCTSCQCDTKGTISGVAECQQGNGQCFCKPSICGQLCSTCKDGYFNLDHANYFGCQGCRCDVGGSIGPACEARTGACHCSCHHRAGTTLDGRAVRFGYNPLEFENFSWRGYAQMSPIQPKVVVTLNVTSPDLFRIVFRYVNRGPANVEGRVSVLEEGKFNVCGNCTEQTKQIIFAPSTEPAFVTVPQNSFGEPFVLNPSVWSLIVEAEGILLDYLVLLPSSYYEAPILQLKVTEACTYSPAPDCLLYKYLSVDSFPSASGGDMVCRLDNSLPRPCPMVPITPSHPQMALCSGSDVEVQLRLVVPQPGRYVLLVEYANKDAMQRASVAISSLQLATQQGSLTFHTCDYSFLCRGIVLDSQNRLAAFELTTEGTIRFAADQAHFFLYKVYLIPYGQFTMEFVEPKVHCISLHGTFSPSSSSCIPSRFMKLSQSIVLEAGQALSIPPDFPLTHVFTVAPTGIPTEPLPRPPTAVDSSAELILLQPSQAAAVFNSRIQTLGRYAFIMHYYQPNHPTFPVEVLINGGRIWQGQAKAAFCPHGYGCRSVVVSENQVILDVTDNELTVIVRVPAGRQLWLDYILVVPEDSYSSHYLLEEPLDKSYEFISSCGVNSFYINPTTSSKFCRDSAISLSLFYNNGAQSCRCHEAGAIGGQCELFGGQCACKSNIIGRECSRCATGYWGFPNCRPCDCGTRLCDEVTGQCICPPHTIKPECVVCEPQTFGCHALIGCEDCNCSRTGVQELAEPGCDMDSGQCRCKPNIIGRRCDVCAPGYYHYPHCRRCYCHQAGTEVSVCDPVTGQCHCKENVEGVRCDQCRLGTFSLDANNPKGCTRCFCFGATDRCHSAEKYRVEFINMNGWVLLSGDRQEVATSLRLEEQLIHADLKNVPEAYQELYWHAPSSYLGDRVSSYGGYLRYELRSDTRRGDVFIPMEPRPDVILKGNQMSIMFLEGTSPSPGEVHEGRLQLMEGNFRHTQTHNPVSREELMMVLANLEQLQIRALFSQLASSVYLRQVVLETVTEISGGIRASNVELCMCPANYRGDSCQECAPGYYRDTKGLFLGKCVPCHCNGHSDQCLPGSGICINCQHNTEGDRCERCKDGYVGNFSLDGPLQCMGCPCPLSVASNNFAVGCVHKGSTMQCLCKPGYAGAMCERCAPGYYGNPLVIGSSCQPCDCSENTDPNMLFSDCDSLTGACTGCMYNTAGPRCEMCAPGYHGDAVVAKNCTRCNCLPCGTELCDPRTGRCLCKPGVTGQHCDCCQDGYYGYHDCSGCRKCDCDIGSIGTSCHAETGQCSCQPGVTGLRCQQCAPGYWGLSKHGCTKCRCKGGSCDPRSGECTCSNGLTGKQCDTCIEKYEIPVANGPDSMRCEVCDSCVLLLLDDLQKIKASFPIIKNQLINLNASSIAWARLHGLNSSVKNVTVCGGLNVFSPLLGTADLSTASSEEFWQKMAEVQRMLKEMKERDFLSQKGVAEHEHEKAQKLLHRVRSELYSKWESNQDLVKSIKDMLSRYSSELMDLRDALNEAVNKTRQTEDLNSLNRNNLEESQKQYGVVQDTLQMAEDSLVQVSDLLQIIENAKEEYEKLAARLDGARYPLTERVKKFSPASSKIPVVEEAEEYARLLDELARNLSSSIQGANQDGFIQRAINASNAYASIIKAVKNAERAAKDADEAAGEALTVCNFPSGPHQPLQNPGSSDVEPPTHVRPITAKLMQKRSADCSQKLGALPSLLLFAEFKILVSFPLTDNTTAVIQNAKAAAAAANDTVAMVEDTLSNMKKNLDEWKDQYGGLKNEELNQAVQDARKSVSNLESTIPLLLEKLSGLENRRNKNSTVSENILRIRQLITQARNAASKVKVPVKFNGTSGIQVRTPSDLQDLSAYTSLKFYIQNPEPKPRQRRQDGADAGQFILYLGNKDATGDYMGIVLKDRKVQWIYKLGDEEPTYLTVDEEIGEQFAAVSINRILQYGHMSVIVEKQTLHETKGDSVAKGEQGLLNLEPHSVVFYVGGYPTGFTPPVTLRYPSYRGCIEMDTLNEEVVSLYNFQRTFHLDTAADKPCARSKSTGDPWLTDGSYFDGTGYAEIKFESQIGSTKRFEQEIRVVSYNGILFFLEHQFLCLAVQDGKLVLYYDFKQVLEMAPPSSDPTSLTVSSTTNKAIQIFLLTMSSKQRILVRLERQTIFMVEHENSLENAVSYYLGGVPMPLLPKSLKAIFPKGGSIRGCMKGLKALGKYVDLKRMNTVGVSYGCTSDLLVARSVKFHGHGYLTLPLKNVPSLQDFYTGFSFRTSQSHGLLYSHTTTEGNCQVSLKNGRVAVTVLKTELTTKHAYADDASHYVAIYSDATGVRLYIDDQLQGPKQGADRYKRHRRQEEQMLFRLGGLPEAGALSNLTGCIGNVFVNWGENCDTERPLLFCRSHFSLDVRLNSPSGLLFYVASELQGSFMALFVSSGRFVFLADLNGKKLRVRSKDKYHDGRWHTVSRGTAGLVLGRAKSLQPIYWSHAFPAHVLCQQVMDQACECLPTLALSLSLQDTSVPSFSGCLKNLKLDGKALDSPSRVFGVTPCYEGMLESGIFFAAEGGYVTLANSVAMGRDLELRLDVRPRSASGLIFHMGTQRKHYLTLYADDTQVPLLAANAGAGEFSTSVARPSLCDGQWHSIAAPLCLSAGIATLEAASPSLLTPLPHYVGCIRNLVINQNPVDLLQAGTIRGSVGLTGCPLHYPPGALLGLDTQPAISCSLD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN57</accession>
    <name>A0A452HN57_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HN57"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018701.1">
        <property type="protein sequence ID" value="ENSGAGP00000016387.1"/>
        <property type="gene ID" value="ENSGAGG00000012269.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="103"/>
            <end position="122"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016387.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="303" mass="32662" checksum="B4C9DBC6A0AA8280" modified="2019-05-08" version="1">MDSSLPARWPPPPAGERVGAPGLSSSAVSLYGQLRCPILSPWTGAPSWECCRRLLWGKIWAVSSQVGVTGRCHLPSISGTHEGLARRWPRKQASWGIISAGPPPAHPHGRAALPAQDAPARPHGREALPLQRLRPGLQPGLQSVHALADPHRRAPLPLRLLWQELQLEARPHQAPAGAHGRASLPLHGLQQELQPEAGPHQAPAGAHGRAPLPLRPVWQELQPPKAPAMPLALGLRRAVLHWGYSALQPHCTGLSKGPQGTPTSSLCTARPKASPCPYCPCWPETRAQPLGNENRDTEKWGRD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN59</accession>
    <name>A0A452HN59_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">A-kinase anchor protein 7-like phosphoesterase domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="5">Binds cAMP-dependent protein kinase (PKA). Interacts with PRKCA; only the cytoplasmic form is capable of interacting with PRKCA.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HN59"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018726.1">
        <property type="protein sequence ID" value="ENSGAGP00000016410.1"/>
        <property type="gene ID" value="ENSGAGG00000012287.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034237">
        <property type="term" value="F:protein kinase A regulatory subunit binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010738">
        <property type="term" value="P:regulation of protein kinase A signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1140.10:FF:000004">
        <property type="entry name" value="A-kinase anchoring protein 7 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1140.10">
        <property type="entry name" value="Cyclic phosphodiesterase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019510">
        <property type="entry name" value="AKAP7-like_phosphoesterase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052641">
        <property type="entry name" value="AKAP7_isoform_gamma"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009097">
        <property type="entry name" value="Cyclic_Pdiesterase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15934:SF6">
        <property type="entry name" value="A-KINASE ANCHOR PROTEIN 7 ISOFORM GAMMA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15934">
        <property type="entry name" value="RNA 2',3'-CYCLIC PHOSPHODIESTERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10469">
        <property type="entry name" value="AKAP7_NLS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55144">
        <property type="entry name" value="LigT-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0539">Nucleus</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="A-kinase anchor protein 7-like phosphoesterase" evidence="7">
        <location>
            <begin position="81"/>
            <end position="276"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="1"/>
            <end position="27"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="61"/>
            <end position="81"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="6">
        <location>
            <begin position="1"/>
            <end position="18"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00038702"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF10469"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016410.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="335" mass="38581" checksum="EB4FB6E9E28F4888" modified="2019-05-08" version="1">MQEKKIIRKKKKRTKIRKGSQTPPDSMDYLMAEIPFASTDTGDEFGTTSTSEVCFKKRRKRTIRSKTEDDDQTKKKKQDQPNYFISLPITNPKITRGIQAVQDIVVQKDHRLSQAMVRCGSLHITLLVMHLSSEEEVGTAVGALLESKDAVEELLQGKHLDLSFQGIDDYRNEVIFVKFAASDHTAALTEISETMKKIFQEKGILAGENRAFKPHLTFMKLSKSPRLRKQVRKIDPHLYENFKSHYFGDETLYRLDLCSMLKKKQPNGYYHCESSIMVGKSHESDLIKKALYRESLALFSKVNKIKELLSNPETRMKINRELFQDRQTAVIKKSC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN84</accession>
    <name>A0A452HN84_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="5">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000018740.1">
        <property type="protein sequence ID" value="ENSGAGP00000016424.1"/>
        <property type="gene ID" value="ENSGAGG00000012286.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030424">
        <property type="term" value="C:axon"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034703">
        <property type="term" value="C:cation channel complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005261">
        <property type="term" value="F:monoatomic cation channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055080">
        <property type="term" value="P:monoatomic cation homeostasis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046460">
        <property type="entry name" value="UNC80_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045852">
        <property type="entry name" value="UNC80_central"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031542">
        <property type="entry name" value="UNC80_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31781:SF1">
        <property type="entry name" value="PROTEIN UNC-80 HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31781">
        <property type="entry name" value="UNC80"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19424">
        <property type="entry name" value="UNC80"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20262">
        <property type="entry name" value="UNC80_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15778">
        <property type="entry name" value="UNC80_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Cation channel complex component UNC80 N-terminal" evidence="2">
        <location>
            <begin position="17"/>
            <end position="236"/>
        </location>
    </feature>
    <feature type="domain" description="Protein UNC80 central region" evidence="3">
        <location>
            <begin position="1436"/>
            <end position="1784"/>
        </location>
    </feature>
    <feature type="domain" description="Protein UNC80 C-terminal" evidence="4">
        <location>
            <begin position="1829"/>
            <end position="2460"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="152"/>
            <end position="173"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="289"/>
            <end position="312"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="450"/>
            <end position="471"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="699"/>
            <end position="790"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="971"/>
            <end position="1026"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1044"/>
            <end position="1081"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1412"/>
            <end position="1452"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1824"/>
            <end position="1843"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="296"/>
            <end position="307"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="450"/>
            <end position="460"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="701"/>
            <end position="715"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="1">
        <location>
            <begin position="741"/>
            <end position="773"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="775"/>
            <end position="786"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="971"/>
            <end position="981"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="1">
        <location>
            <begin position="1044"/>
            <end position="1060"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="1425"/>
            <end position="1437"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF15778"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="Pfam" id="PF19424"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="Pfam" id="PF20262"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016424.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2494" mass="279061" checksum="364FDDD697251C02" modified="2019-05-08" version="1">MVKRKSSQGPEQEGGRGVPLAIQTFLWRQTRCELGRGCGGPWRGPLSSLSFERVLVENKLHGLSPALSEAIQSISRWELVQAALPHVLHCTATLLSNRNKLGHQDKLGVAETKLLHTLHWMLLEAPQDCSNDRFGGERGSSWGGSSSAFIHQVENQGSPGHPRRGSTNEEEENNRRKFFQNSMATVELFVFLFAPLVHRIKESDLTFRLASGLVIWQPMWEHRQPEVSAFTALVKPIRNIITAKRSSPTSNQSLACESPNLDTGHTEGLQVVCETTQPDAVPPRATVSGCHRGNSFDGSVSSQTSQDRGPPLSRVSLVIPPCQKSRYATYFDVAVLRCLLQTHWSEEGTQWSLMYYLQRLRHMLEEKPEKPPEPETMHLPRPRSSSMVAAAPSLVNTHKTQDLTMKCNEEEKSLSTEAFAKVSLTNLRRLAVPDLSTDLGMNIFKKFKSRKEDRERERKGSIPFHHTGKKRQRRMGVPFLLHEDHLDVSPTRSTFSFGSFSGIGEDRRSIERGGWQTAILGKFTRRGSSDTATEMESLSARHSHSHHTLVTDLPDHSNSHAENTVKEVRSQISTITVATFNTTLASFNVGYADFFSEHMRKLCNQVPIPEMPHEPLACANLPRSLTDSCINYSCLEDTDHIDGTNNFVHKNGMLDLSVVLKAVYLVLNHDISSRICDVALNIVECLLQLGVVPCVEKVRRKSENKENESAEKRPSEGSFQLKGATSGGPACGFGAPPVSGTGDGGGEEGGGGGSGGGGGGGADGGGGGGGGGPYEKNDKNQEKDESTPISTHRLALTMLIKIVKSLGCAYGCGEGHRGLSGDRLRHQVFQENAQNCLTKLYKLDKMQFRQTMRDYVNRDSLNNVVDFLHALLGFCMEPVTDNKAGFGNNFTTVDNKSTAQSVEGIIVGAMFKSLITRCASTTHELHSPENLGLYCDIRQLVQFIKEAHGNVFRRVALSALLDSAEKLIPGKKTDETTEQDLKPLGSKKSEVGSVVIDKGQTSSAPDECRSYISSRPQQTPEHDEQMQGAVLGRKDFWRKMFKSQSAASDTSSQSEQDTSECTTAHSGTTTDRRSRSRSRRISLRKKLKLPIGKWNWLKRSSLSGLADGVEDLLDISSVDRLSFIRQSSKVKFTSAVKLSEGGPGGGLENGRDEEENFFKRLGCHSFDDHLTSNQEGGKSKNVVNLGAIRQGMKRFQFLLNCCEPGTIPDASILAAALDLEAPVVARAALFLECARFVHRCNRGNWPEWMKGHHVNITKKGLSRGRSPIVGNKRNQKLQWSAAKLFYQWGDAIGIRLNELCHSESESPANLLGLIYDEEAKRRLRKEDEEEDFLDDSTVNPTKCCCPFALKMAACQLLLEITTFLRETFPYMPRPRTEPLVDLESCRLRLDPELDRHRYERKISFAGILDENEDSKDSLHSSSHTLKSDTGCEEKKEGSPWSASEPSIEPEVLSNTGTEENYHRNMSWLHVMILLCNQQSFICTHIDYCHPHCYLHHSRSCARLVRAIKLLYGDTVDSLRENNVLNNVARGKKQKECSDKSCLRTPSLKKRVMDINLEGKKDSGMLKYIRQQVMSLSPAPLSLLIKAAPILTEEMYGDIQPAAWELLLSVDEHMAGAAAAMFLLCAVKVPEAVSDMLMSEFHHPETVQRLNAVLKFHTLWRFRYQVWPRMEEGAQQIFKIPPPSINFTLPSPVLGMPSVPMFDPPWVPQCSGSVQDPINEDQSKSFSARAVSRSHQRAEHILKNLQQEEEKKRLGREASLITAIPITQEACYEPTCTPNSEQEEEVEEVTNLASRRLSVSPSCTSSTSHRNYSFRRGSVWSVRSAVSAEDEEHTTEHTPNHHVPQPPQAVFPACICAAVLPIVHLMEDGEVREDGVAVSAVAQQVLWNCLIEDPSTVLRHFLEKLTISNRQDELMYMLRKLLLNIGDFPAQTSHILFNYLVGLIMYFVRTPCEWGMDAISATLTFLWEVVGYVEGLFFKDLKQTMKKEQCEVKLLVTASMPGTKTLVVHGQNECDIPTQLPVHEDTQFEALLKECLEFFNIPETQSAHYFLMDKRWNLIHYNKTYVRDIYPFRRSVSPQLNLVHMHPEKGQELIQKQVFTRKLEEVGRVLFLISLTQKIPVAHKQSHVSMLQEDLLRLPSFPRSAIDAEFSLFSDPQAGKELFGLDTLQKSLWIKLLEEMFLGMPSEFPWGDEIMLFLNVFNGALILHPEDSALLRQYTATVINTAVHFNHLFSLSGYQWILPTMLQVYADYESNPQLRQAIEFACRQFYILHRKPFVLQLFASVAPLLEFPDATNNGPSKGVSAQCLFDLLQSLEGDTPDILDILELVKAEKPLKSLDFCYGNEDLTFSISEAIKLCVTVVAYAPESFRSLQMLMVLEALVPCYLQKLKRQTSQVETVPAAREEIAATAALATSLQALLYSVEVLTRPMTAPQMSRCDQGHKGATTANHTMSAGVNTRSQYFCSKRFCSEFYFPCGICPHGLHINKENHFLKTI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN86</accession>
    <name>A0A452HN86_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Erbin</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="13">Densin-180-like protein</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="15">Erbb2-interacting protein</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="14">Protein LAP2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Basolateral cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="11">Cell junction</location>
            <location evidence="11">Hemidesmosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the LAP (LRR and PDZ) protein family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018780.1">
        <property type="protein sequence ID" value="ENSGAGP00000016454.1"/>
        <property type="gene ID" value="ENSGAGG00000006183.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005912">
        <property type="term" value="C:adherens junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016323">
        <property type="term" value="C:basolateral plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030056">
        <property type="term" value="C:hemidesmosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031965">
        <property type="term" value="C:nuclear membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0014069">
        <property type="term" value="C:postsynaptic density"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019901">
        <property type="term" value="F:protein kinase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098609">
        <property type="term" value="P:cell-cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045197">
        <property type="term" value="P:establishment or maintenance of epithelial cell apical/basal polarity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070433">
        <property type="term" value="P:negative regulation of nucleotide-binding oligomerization domain containing 2 signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098968">
        <property type="term" value="P:neurotransmitter receptor transport postsynaptic membrane to endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098887">
        <property type="term" value="P:neurotransmitter receptor transport, endosome to postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043113">
        <property type="term" value="P:receptor clustering"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010468">
        <property type="term" value="P:regulation of gene expression"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd06749">
        <property type="entry name" value="PDZ_densin_erbin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000036">
        <property type="entry name" value="Erbin isoform 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000013">
        <property type="entry name" value="Erbin isoform 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000020">
        <property type="entry name" value="Erbin isoform 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000022">
        <property type="entry name" value="Erbin isoform 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.42.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001611">
        <property type="entry name" value="Leu-rich_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003591">
        <property type="entry name" value="Leu-rich_rpt_typical-subtyp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001478">
        <property type="entry name" value="PDZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036034">
        <property type="entry name" value="PDZ_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050614">
        <property type="entry name" value="Synaptic_Scaffolding_LAP-MAGUK"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23119">
        <property type="entry name" value="DISCS LARGE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23119:SF46">
        <property type="entry name" value="ERBB2 INTERACTING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13855">
        <property type="entry name" value="LRR_8"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00595">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00364">
        <property type="entry name" value="LRR_BAC"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SMART" id="SM00365">
        <property type="entry name" value="LRR_SD22"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00369">
        <property type="entry name" value="LRR_TYP"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="SMART" id="SM00228">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52058">
        <property type="entry name" value="L domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50156">
        <property type="entry name" value="PDZ domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51450">
        <property type="entry name" value="LRR"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50106">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0965">Cell junction</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="6" id="KW-0433">Leucine-rich repeat</keyword>
    <keyword evidence="9" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="PDZ" evidence="17">
        <location>
            <begin position="1260"/>
            <end position="1349"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="474"/>
            <end position="500"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="517"/>
            <end position="555"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="650"/>
            <end position="688"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1002"/>
            <end position="1021"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1076"/>
            <end position="1099"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1135"/>
            <end position="1194"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="16">
        <location>
            <begin position="481"/>
            <end position="491"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="16">
        <location>
            <begin position="517"/>
            <end position="545"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="16">
        <location>
            <begin position="653"/>
            <end position="665"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="666"/>
            <end position="684"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1076"/>
            <end position="1089"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1158"/>
            <end position="1178"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004126"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004187"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007772"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022614"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00060371"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00074479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00076296"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00083415"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00084015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50106"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016454.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1351" mass="151123" checksum="EAF0B8A7070F1260" modified="2019-05-08" version="1">MSSDGDSTTSFGSPIQYLTILICLKMTTKRNLFVRLVPCRCLRGEEETVTALDYSHCSLEQVPKEIFTFEKTLEELYLDANQIEELPKQLFNCQSLHKLSLPDNDLTTLPASIANLINLRELDVSKNGIQEFPENIKNCKVLAIVEASVNPISKLPDGFSQLLNLTQLYLNDAFLEFLPANFGRLTKLQILELRENQLKMLPKTMNRLTQLERLDLGSNEFTEVPEVLEQLSGLKEFWMDGNRLTLIPGLIGSLKQLTYLDVSKNNIEIVEEGVSGCESLQDLLLSSNSLQQLPESIGSLKKVTTLKIDENQLMYLPDSIGGLIAVEELDCSFNEIEALPSSIGQLTNVRTFASDHNFLTQLPPEIGNWKNVTVLFLHSNKLEFLPEEMGDMQKLKVINLSDNRDFNYFFFSQSKPLIPLQKEVDPETQKTVLTNYMFPQQPRSEDVLFISDNESFNPSLWEEQRKQRAQVAFECDEDKDEREAPPREGNLKRYPTPYPDELKNMVKTVQTIVHRLKDEETTEDGGRGSKKEGQQVSVKDVDVKTSEMASAKSKTEDGENYMMENCVQKPTEPEAEHSTGNSQMTSHVKTLENTKIIVNHEDTLEESEELSSDEEMKMAEMRPPLIETSINQPKVITKHTKSRVLLYDEATHNSNHNNSNCSSPSRMSDSVSLNTDSSNDTSVCSPDKETHSAAISKIRQEDENFNSLLQNGSELNITAEENISAQEKVTNLPEYELRIEERLALIEKGVDLNTTTEESHKLDQINMNINRLTSEDTIQPNVMERLKTQEIVSVKGCSNYNMKEETEHLENRNQYSNLCSINKVNGHSEREPQSLRKAELKRSITVDTDSSVDICISKSTEDLSPQRSGPVVKSHSITNIDNGALKIFDIVNDNGSQEPNSVVKSASAGPDGKNIVRSKSATLLYDHPLQVFPGSSSSSDLVSCTKTLFKFDSNHNPEGANVVRGPVASGAQMFSVPQYNIQYSSSTTAKDTLWLPKQNAQIEQGSLPPQRLSRSDSTESSNYVKHSANMNFSNHNNVRANAAYNMHQRMGGRHIEMWAIPPNDRLVPGATRNTLQRQSSISSTASINVGDTGPPRRAQVAEGDCLTYRDLHSMGRAPPIMSGQPRPLSARTYSIDGPNVPRPQSARPSVNEIPERTMSVSDFNYSRTSPTKRPNPRVNSEHSLLDPPGKSKVPHDWREQVLRHIEAKKLEKSMLSRSFNSNYAAISNFQYGSSRDLHGSQSSVALSVADRRSCGVHIVRIRVRIEKDPELGFSISGGVGGRGNPFRPDDDGIFVTRVQPEGPASKLLQPGDKIIQANGYSFINIDHGQAVSLLKTFQNTVELIILREVSS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN88</accession>
    <name>A0A452HN88_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Collagen alpha-1(XII) chain</fullName>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">Type XII collagen interacts with type I collagen-containing fibrils, the COL1 domain could be associated with the surface of the fibrils, and the COL2 and NC3 domains may be localized in the perifibrillar matrix.</text>
    </comment>
    <comment type="subunit">
        <text evidence="14">Trimer of identical chains each containing 190 kDa of non-triple-helical sequences.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="12">Belongs to the fibril-associated collagens with interrupted helices (FACIT) family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018733.1">
        <property type="protein sequence ID" value="ENSGAGP00000016417.1"/>
        <property type="gene ID" value="ENSGAGG00000011657.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005581">
        <property type="term" value="C:collagen trimer"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005614">
        <property type="term" value="C:interstitial matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035987">
        <property type="term" value="P:endodermal cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="CDD" id="cd01482">
        <property type="entry name" value="vWA_collagen_alphaI-XII-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000018">
        <property type="entry name" value="collagen alpha-1(XII) chain isoform X1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000489">
        <property type="entry name" value="collagen alpha-1(XII) chain isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000639">
        <property type="entry name" value="collagen alpha-1(XII) chain isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000816">
        <property type="entry name" value="collagen alpha-1(XII) chain isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.200:FF:000008">
        <property type="entry name" value="Collagen type XII alpha 1 chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000121">
        <property type="entry name" value="Collagen type XII alpha 1 chain"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000234">
        <property type="entry name" value="Collagen, type XII, alpha 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000480">
        <property type="entry name" value="Collagen, type XII, alpha 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.410:FF:000001">
        <property type="entry name" value="Collagen, type XII, alpha 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.200">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.320">
        <property type="entry name" value="6-Phosphogluconate Dehydrogenase, domain 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.410">
        <property type="entry name" value="von Willebrand factor, type A domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008160">
        <property type="entry name" value="Collagen"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050525">
        <property type="entry name" value="ECM_Assembly_Org"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048287">
        <property type="entry name" value="TSPN-like_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002035">
        <property type="entry name" value="VWF_A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036465">
        <property type="entry name" value="vWFA_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24020">
        <property type="entry name" value="COLLAGEN ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24020:SF17">
        <property type="entry name" value="COLLAGEN ALPHA-1(XII) CHAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01391">
        <property type="entry name" value="Collagen"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00092">
        <property type="entry name" value="VWA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00453">
        <property type="entry name" value="VWFADOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="SMART" id="SM00210">
        <property type="entry name" value="TSPN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00327">
        <property type="entry name" value="VWA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53300">
        <property type="entry name" value="vWA-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50234">
        <property type="entry name" value="VWFA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="8" id="KW-0176">Collagen</keyword>
    <keyword evidence="9" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11" id="KW-0379">Hydroxylation</keyword>
    <keyword evidence="7" id="KW-0654">Proteoglycan</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="4 17" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="17">
        <location>
            <begin position="1"/>
            <end position="35"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019037798" description="Collagen alpha-1(XII) chain" evidence="17">
        <location>
            <begin position="36"/>
            <end position="1772"/>
        </location>
    </feature>
    <feature type="domain" description="VWFA" evidence="18">
        <location>
            <begin position="36"/>
            <end position="208"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="19">
        <location>
            <begin position="224"/>
            <end position="313"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="19">
        <location>
            <begin position="314"/>
            <end position="405"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="19">
        <location>
            <begin position="406"/>
            <end position="495"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="19">
        <location>
            <begin position="496"/>
            <end position="586"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="19">
        <location>
            <begin position="587"/>
            <end position="681"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="19">
        <location>
            <begin position="769"/>
            <end position="859"/>
        </location>
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</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN89</accession>
    <name>A0A452HN89_9SAUR</name>
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            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
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            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
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            <taxon>Gopherus</taxon>
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        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
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                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
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        </source>
    </evidence>
    <sequence length="1938" mass="213626" checksum="CFE3DC50B2C2BED8" modified="2019-05-08" version="1">MADDRKDEAKAPHWTSSQLTDASSHPHSPEIKEQAGAGAELVRSANGFPYREDEEAGYGERGQQGTYSRTKENGINGELSTGDRETAVEESANLPPSPPPSPASEQTGTLAEEEETVESPNAEEAKPSALPGKELGKDSGSAEPSERAEGQLDSGSEAKVYHEDSVLSVAPSGKSITVVEEVPLKANIPSLAEEASKMEIRDQEGTGPFTAEPLDTKERESEKDSKTDEQSKHDALVPQPAKTETLDKKESWGKEEKMHPTVSGQTLDADLQKKGEVSFADPVADSTAEPLVSQEHMDLSADLLKESKTDERSIDTQIPSNQAIPLKYKDDFTETQKSNLALLPSESRIDLSSAARSEPLSETPMTVSQKSQLEDSCKTKQEFSDKEIAKDLLKDEKIHKDQTEKPEAGQVSSMAADAPKPTTEERPIQKVSTWGEEKDMTRDESDEEERYGFLDWQPSKVLDDTKLYAEQEDKIVLPDKAVVSEAKKSSDGLKAEEETDRTGVSTVVDAKKEEQQSTETHTGKLSHESVPEKAKEPIDTDHLSKPTEKAPDAPRLTSEKGPTHELHQDRTIEKDAKEDRFHISDSASHEIQVEEDRSGMSKYFETSALKEEATKSDTQQQGSDYYELSDTKESAYELYHADPLLSKKEDKKLQPETDQPSTVPAHEIGYSTLAQSFPSDKSEEPSSPQERMFTIDPKVYGDKRQIHSKNKDDLTLSRSLGLGGRSAIEQRSMSINLPMSCLDSIALGFSFGRAQDLSPLASDILTNTSGSMDEGDDYLPATTPALERAPCFPIDSTEEDENVEEEKAMEEENIQVESLCESPFLAKDYYKNGAVMAPDLPEMLDLAGTRSRLASVSADTEVARRKSVPSDAAVEDSGASLPPVTDENHITLKAESQLEDLGYCVFNKYTVPLPSPVQDSESLTGEICPFYEGRDEKMRRGLAADLSLIEMKLAAAEKSKEEFSSEKDLSPNISSDTTVLGREIEQEKKGSDKLATVLEKSEEQVDSKEIHSLEDTKHEKKRPDAILEMKEEGTVDKLHIAGPESDIANDKALEKEVTKEKDLVSLLTQKEEKSLNIIPKVAEIETPHRPDYDAIKHDMDVAARKADQEYQTQLHTEINEFAAAGSKQDTDSVKKAEPEAQEAQQQQKDLASLPKEPKEADVLSRTEPSQMKDSTKLSESEIKEKVPKPDLVHQEAVDKEESYESSGEHDQIQEDLNGESLKPDYGKIKPQKPPVSEEEITSKLPVEEPSIELAISKADSVQEETADIQMESIPQPKEEIKETTPDIAVRPAETTEQLLSDLTPESVEGKEHKEEEDDDKHQLMSDVHQEIELEAPAGEEALTEAGPEEMPELKGVIESVVTIEDDFITVVQMTVEESESASHSVRFAAAQQEEMERRDFLPEEEVEMEMEEPADIQAEPKEGSLEAPASPEKEEILLTDYKTEIYDDFKDETTIDDSVMDTDSLWADTQDDDRSIMTEQLETVPKEEKTERELRRSSLDKHKKEKPFKTGRGRISTPERKIAKKEPSTLSRDEVRRKKAVYKKAELAKKSEVQAHSPSRKIILKPAIKYTRPTHLSCVKRKQTAAGGETNQAPGVFKQAKEKLSDGITKSPEKRSSLPRPSSILPTRRGVSGDRDRDENSFSLNSSISSSVRRTTRSEPIRSRTGKSGTSTPTTPGSTAITPGTPPSYSSRTPGTPGTPSYSRTPHTPGTPKSAILVPTEKKVAIIRTPPKSPATPKQLRLINQPLPDLKNVRSKIGSTDNIRYQPKGGQVRILNKKIDFSDIQSRCGSKDNIKHSAGGGNIQIVTKKIDLSHVTSKCGSLKNIRHRPGGGRVKIESVKLDFKEKAHAKVGSLENAHHTPGGGNIKIDSQKLNFREHAKARVDHGAEIITQSPGRSSVPSPRRLSNVSSSGSINLLESPQLATLAEDVTAALAKQGL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN91</accession>
    <name>A0A452HN91_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Protein transport protein SEC23</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="5">Component of the coat protein complex II (COPII) which promotes the formation of transport vesicles from the endoplasmic reticulum (ER). The coat has two main functions, the physical deformation of the endoplasmic reticulum membrane into vesicles and the selection of cargo molecules.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="5">Cytoplasmic vesicle</location>
            <location evidence="5">COPII-coated vesicle membrane</location>
            <topology evidence="5">Peripheral membrane protein</topology>
            <orientation evidence="5">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1 5">Endoplasmic reticulum membrane</location>
            <topology evidence="1 5">Peripheral membrane protein</topology>
            <orientation evidence="1 5">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="5">Cytoplasm</location>
            <location evidence="5">Cytosol</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 5">Belongs to the SEC23/SEC24 family. SEC23 subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HN91"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018771.1">
        <property type="protein sequence ID" value="ENSGAGP00000016445.1"/>
        <property type="gene ID" value="ENSGAGG00000006201.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030127">
        <property type="term" value="C:COPII vesicle coat"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070971">
        <property type="term" value="C:endoplasmic reticulum exit site"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090110">
        <property type="term" value="P:COPII-coated vesicle cargo loading"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006886">
        <property type="term" value="P:intracellular protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd01478">
        <property type="entry name" value="Sec23-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd11287">
        <property type="entry name" value="Sec23_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.730:FF:000003">
        <property type="entry name" value="Protein transport protein SEC23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1670:FF:000006">
        <property type="entry name" value="Protein transport protein SEC23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.20.10:FF:000003">
        <property type="entry name" value="Protein transport protein SEC23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.410:FF:000011">
        <property type="entry name" value="Protein transport protein SEC23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1670">
        <property type="entry name" value="beta-sandwich domain of Sec23/24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.730">
        <property type="entry name" value="Sec23/Sec24 helical domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.20.10">
        <property type="entry name" value="Severin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.410">
        <property type="entry name" value="von Willebrand factor, type A domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029006">
        <property type="entry name" value="ADF-H/Gelsolin-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012990">
        <property type="entry name" value="Beta-sandwich_Sec23_24"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007123">
        <property type="entry name" value="Gelsolin-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036180">
        <property type="entry name" value="Gelsolin-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037364">
        <property type="entry name" value="Sec23"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006900">
        <property type="entry name" value="Sec23/24_helical_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036175">
        <property type="entry name" value="Sec23/24_helical_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006896">
        <property type="entry name" value="Sec23/24_trunk_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037550">
        <property type="entry name" value="Sec23_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036465">
        <property type="entry name" value="vWFA_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036174">
        <property type="entry name" value="Znf_Sec23_Sec24_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11141">
        <property type="entry name" value="PROTEIN TRANSPORT PROTEIN SEC23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11141:SF10">
        <property type="entry name" value="PROTEIN TRANSPORT PROTEIN SEC23B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00626">
        <property type="entry name" value="Gelsolin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08033">
        <property type="entry name" value="Sec23_BS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04815">
        <property type="entry name" value="Sec23_helical"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04811">
        <property type="entry name" value="Sec23_trunk"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81995">
        <property type="entry name" value="beta-sandwich domain of Sec23/24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF82754">
        <property type="entry name" value="C-terminal, gelsolin-like domain of Sec23/24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81811">
        <property type="entry name" value="Helical domain of Sec23/24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53300">
        <property type="entry name" value="vWA-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF82919">
        <property type="entry name" value="Zn-finger domain of Sec23/24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4 5" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="5" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="5" id="KW-0931">ER-Golgi transport</keyword>
    <keyword evidence="5" id="KW-0472">Membrane</keyword>
    <keyword evidence="5" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="5" id="KW-0653">Protein transport</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0813">Transport</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="Sec23/Sec24 trunk" evidence="7">
        <location>
            <begin position="96"/>
            <end position="349"/>
        </location>
    </feature>
    <feature type="domain" description="Sec23/Sec24 beta-sandwich" evidence="9">
        <location>
            <begin position="360"/>
            <end position="463"/>
        </location>
    </feature>
    <feature type="domain" description="Sec23/Sec24 helical" evidence="8">
        <location>
            <begin position="477"/>
            <end position="575"/>
        </location>
    </feature>
    <feature type="domain" description="Gelsolin-like" evidence="6">
        <location>
            <begin position="591"/>
            <end position="677"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004397"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009210"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="RuleBase" id="RU365030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="Pfam" id="PF00626"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF04811"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF04815"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF08033"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016445.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="724" mass="81610" checksum="EF16B65261D2149A" modified="2019-05-08" version="1">MATYLEFIQQNEERDGVRFSWNVWPSSRLEATRMVVPLACLLTPLKERLDLPPVQYEPVLCSRQTCKAVLNPLCQVDYRAKLWACNFCFQRNQVGSESEDLQALKESLQMSLSLLPPEALVGLITFGRMVQVHELSCEGISKSYVFRGTKDLTAKQIQDMLGLTRPVVPVQQGRPLQPQEQPPISSRFLQPVHKIDMNLTDLLGELQRDPWPVTQGKRPLRSTGVALSIAVGLLEGTFPNTGARIMLFTGGPPTQGPGMVVGDELKTPIRSWHDIEKDNARFMKKATKHYETLANRTAANGHCIDIYACALDQTGLLEMKCCANLTGGHMMMGDSFNTSLFKQTFQRVFSKDFNGEFRMAFGATLEVKTSRELKISGAIGPCVSLNAKGPCVSENELGIGGTCQWKICSLDPSSTLAIYFEVVNQHNAPIPQGGRGAVQFVTQYQHSSTQRRIRVTTIARNWADAQSQLQHIEAAFDQEVAAVLMARLGVYRAESEEGPDVLRWLDRQLIRLCQKFGQYNKDDPNSFRLSDSFSLYPQFMFHLRRSPFLQVFNNSPDESSFYRHHFSRQDLTQSLIMIQPILYSYSFSGPPEPVLLDSSSILADRILLMDTFFQIVIYHGETIAQWQKAGYQDMPEYENFKHLLQAPLDDAQEILQTRFPMPRYINTEHGSSQARFLLSKVNPSQTHNNLYAWGQESGSPILTDDVSLQVFMDHLKKLAVSSAS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN92</accession>
    <name>A0A452HN92_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">PRELI/MSF1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HN92"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018786.1">
        <property type="protein sequence ID" value="ENSGAGP00000016460.1"/>
        <property type="gene ID" value="ENSGAGG00000012317.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005758">
        <property type="term" value="C:mitochondrial intermembrane space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006797">
        <property type="entry name" value="PRELI/MSF1_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037365">
        <property type="entry name" value="Slowmo/Ups"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11158">
        <property type="entry name" value="MSF1/PX19 RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04707">
        <property type="entry name" value="PRELI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50904">
        <property type="entry name" value="PRELI_MSF1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="PRELI/MSF1" evidence="1">
        <location>
            <begin position="1"/>
            <end position="161"/>
        </location>
    </feature>
    <evidence type="ECO:0000259" key="1">
        <source>
            <dbReference type="PROSITE" id="PS50904"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016460.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="161" mass="17919" checksum="5870B89827BE9BC5" modified="2019-05-08" version="1">GNWEPAQHTLFPARCVPPPARVTAPPPLSRLFLCPERALGRSPLGLARRGQRRSYEDLELGARVRILGTSRTLTYIKEHSVVDPVEKKMELCSTNITLTNLVSVDERLVYMPHPENPEKTVLTQEAIITVKGVSLSSYLESLMANTISSNARKVCFFDSNS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN94</accession>
    <name>A0A452HN94_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Erbin</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="13">Densin-180-like protein</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="15">Erbb2-interacting protein</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="14">Protein LAP2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Basolateral cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="11">Cell junction</location>
            <location evidence="11">Hemidesmosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the LAP (LRR and PDZ) protein family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018776.1">
        <property type="protein sequence ID" value="ENSGAGP00000016450.1"/>
        <property type="gene ID" value="ENSGAGG00000006183.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005912">
        <property type="term" value="C:adherens junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016323">
        <property type="term" value="C:basolateral plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030056">
        <property type="term" value="C:hemidesmosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031965">
        <property type="term" value="C:nuclear membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0014069">
        <property type="term" value="C:postsynaptic density"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019901">
        <property type="term" value="F:protein kinase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098609">
        <property type="term" value="P:cell-cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045197">
        <property type="term" value="P:establishment or maintenance of epithelial cell apical/basal polarity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070433">
        <property type="term" value="P:negative regulation of nucleotide-binding oligomerization domain containing 2 signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098968">
        <property type="term" value="P:neurotransmitter receptor transport postsynaptic membrane to endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098887">
        <property type="term" value="P:neurotransmitter receptor transport, endosome to postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043113">
        <property type="term" value="P:receptor clustering"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010468">
        <property type="term" value="P:regulation of gene expression"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd06749">
        <property type="entry name" value="PDZ_densin_erbin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000036">
        <property type="entry name" value="Erbin isoform 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000013">
        <property type="entry name" value="Erbin isoform 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000020">
        <property type="entry name" value="Erbin isoform 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000022">
        <property type="entry name" value="Erbin isoform 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.42.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001611">
        <property type="entry name" value="Leu-rich_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003591">
        <property type="entry name" value="Leu-rich_rpt_typical-subtyp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001478">
        <property type="entry name" value="PDZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036034">
        <property type="entry name" value="PDZ_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050614">
        <property type="entry name" value="Synaptic_Scaffolding_LAP-MAGUK"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23119">
        <property type="entry name" value="DISCS LARGE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23119:SF46">
        <property type="entry name" value="ERBB2 INTERACTING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13855">
        <property type="entry name" value="LRR_8"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00595">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00364">
        <property type="entry name" value="LRR_BAC"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SMART" id="SM00365">
        <property type="entry name" value="LRR_SD22"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00369">
        <property type="entry name" value="LRR_TYP"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="SMART" id="SM00228">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52058">
        <property type="entry name" value="L domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50156">
        <property type="entry name" value="PDZ domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51450">
        <property type="entry name" value="LRR"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50106">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0965">Cell junction</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="6" id="KW-0433">Leucine-rich repeat</keyword>
    <keyword evidence="9" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="PDZ" evidence="17">
        <location>
            <begin position="1330"/>
            <end position="1419"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="474"/>
            <end position="500"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="517"/>
            <end position="555"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="650"/>
            <end position="688"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1002"/>
            <end position="1021"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1076"/>
            <end position="1099"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1135"/>
            <end position="1194"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="16">
        <location>
            <begin position="481"/>
            <end position="491"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="16">
        <location>
            <begin position="517"/>
            <end position="545"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="16">
        <location>
            <begin position="653"/>
            <end position="665"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="666"/>
            <end position="684"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1076"/>
            <end position="1089"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1158"/>
            <end position="1178"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004126"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004187"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007772"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022614"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00060371"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00074479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00076296"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00083415"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00084015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50106"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016450.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1421" mass="159048" checksum="89321A062C1C70FA" modified="2019-05-08" version="1">MSSDGDSTTSFGSPIQYLTILICLKMTTKRNLFVRLVPCRCLRGEEETVTALDYSHCSLEQVPKEIFTFEKTLEELYLDANQIEELPKQLFNCQSLHKLSLPDNDLTTLPASIANLINLRELDVSKNGIQEFPENIKNCKVLAIVEASVNPISKLPDGFSQLLNLTQLYLNDAFLEFLPANFGRLTKLQILELRENQLKMLPKTMNRLTQLERLDLGSNEFTEVPEVLEQLSGLKEFWMDGNRLTLIPGLIGSLKQLTYLDVSKNNIEIVEEGVSGCESLQDLLLSSNSLQQLPESIGSLKKVTTLKIDENQLMYLPDSIGGLIAVEELDCSFNEIEALPSSIGQLTNVRTFASDHNFLTQLPPEIGNWKNVTVLFLHSNKLEFLPEEMGDMQKLKVINLSDNRDFNYFFFSQSKPLIPLQKEVDPETQKTVLTNYMFPQQPRSEDVLFISDNESFNPSLWEEQRKQRAQVAFECDEDKDEREAPPREGNLKRYPTPYPDELKNMVKTVQTIVHRLKDEETTEDGGRGSKKEGQQVSVKDVDVKTSEMASAKSKTEDGENYMMENCVQKPTEPEAEHSTGNSQMTSHVKTLENTKIIVNHEDTLEESEELSSDEEMKMAEMRPPLIETSINQPKVITKHTKSRVLLYDEATHNSNHNNSNCSSPSRMSDSVSLNTDSSNDTSVCSPDKETHSAAISKIRQEDENFNSLLQNGSELNITAEENISAQEKVTNLPEYELRIEERLALIEKGVDLNTTTEESHKLDQINMNINRLTSEDTIQPNVMERLKTQEIVSVKGCSNYNMKEETEHLENRNQYSNLCSINKVNGHSEREPQSLRKAELKRSITVDTDSSVDICISKSTEDLSPQRSGPVVKSHSITNIDNGALKIFDIVNDNGSQEPNSVVKSASAGPDGKNIVRSKSATLLYDHPLQVFPGSSSSSDLVSCTKTLFKFDSNHNPEGANVVRGPVASGAQMFSVPQYNIQYSSSTTAKDTLWLPKQNAQIEQGSLPPQRLSRSDSTESSNYVKHSANMNFSNHNNVRANAAYNMHQRMGGRHIEMWAIPPNDRLVPGATRNTLQRQSSISSTASINVGDTGPPRRAQVAEGDCLTYRDLHSMGRAPPIMSGQPRPLSARTYSIDGPNVPRPQSARPSVNEIPERTMSVSDFNYSRTSPTKRPNPRVNSEHSLLDPPGKSKVPHDWREQVLRHIEAKKLEKSMLSRSFNSNYAAISNFQYGSSRDLHGSQSSVALSVADRRSCGVHIVRIPLTNGQMCQPHRPQTNYSQVHHPASQASVARHPSREQLIDYLMLKVAHQPPFTQPQCSPKQGNEMAKQEIRVRIEKDPELGFSISGGVGGRGNPFRPDDDGIFVTRVQPEGPASKLLQPGDKIIQANGYSFINIDHGQAVSLLKTFQNTVELIILREVSS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN95</accession>
    <name>A0A452HN95_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Anillin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="3">Required for cytokinesis. Essential for the structural integrity of the cleavage furrow and for completion of cleavage furrow ingression. Plays a role in bleb assembly during metaphase and anaphase of mitosis. May play a significant role in podocyte cell migration.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell projection</location>
            <location evidence="2">Bleb</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HN95"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018738.1">
        <property type="protein sequence ID" value="ENSGAGP00000016422.1"/>
        <property type="gene ID" value="ENSGAGG00000012291.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005826">
        <property type="term" value="C:actomyosin contractile ring"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032059">
        <property type="term" value="C:bleb"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000915">
        <property type="term" value="P:actomyosin contractile ring assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000281">
        <property type="term" value="P:mitotic cytokinesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031106">
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            <taxon>Durocryptodira</taxon>
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                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
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                <person name="Murphy R.W."/>
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    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF08377"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016427.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1945" mass="214362" checksum="A65F3D80BFA9238E" modified="2019-05-08" version="1">MADDRKDEAKAPHWTSSQLTDASSHPHSPEIKEQAGAGAELVRSANGFPYREDEEAGYGERGQQGTYSRTKENGINGELSTGDRETAEEVSARIVQVVTAEAVAVLKGEQEKEAQHKDQPGPLPLVEESANLPPSPPPSPASEQTGTLAEEEETVESPNAEEAKPSALPGKELGKDSGSAEPSERAEGQLDSGSEAKVYHEDSVLSVAPSGKSITVVEEVPLKANIPSLAEEASKMEIRDQEGTGPFTAEPLDTKERESEKDSKTDEQSKHDALVPQPAKTETLDKKESWGKEEKMHPTVSGQTLDADLQKKGEVSFADPVADSTAEPLVSQEHMDLSADLLKESKTDERSIDTQIPSNQAIPLKYKDDFTETQKSNLALLPSESRIDLSSAARSEPLSETPMTVSQKSQLEDSCKTKQEFSDKEIAKDLLKDEKIHKDQTEKPEAGQVSSMAADAPKPTTEERPIQKVSTWGEEKDMTRDESDEEERYGFLDWQPSKVLDDTKLYAEQEDKIVLPDKAVVSEAKKSSDGLKAEEETDRTGVSTVVDAKKEEQQSTETHTGKLSHESVPEKAKEPIDTDHLSKPTEKAPDAPRLTSEKGPTHELHQDRTIEKDAKEDRFHISDSASHEIQVEEDRSGMSKYFETSALKEEATKSDTQQQGSDYYELSDTKESAYELYHADPLLSKKEDKKLQPETDQPSTVPAHEIGYSTLAQSFPSDKSEEPSSPQERMFTIDPKVYGDKRQIHSKNKDDLTLSRSLGLGGRSAIEQRSMSINLPMSCLDSIALGFSFGRAQDLSPLASDILTNTSGSMDEGDDYLPATTPALERAPCFPIDSTEEDENVEEEKAMEEENIQVESLCESPFLAKDYYKNGAVMAPDLPEMLDLAGTRSRLASVSADTEVARRKSVPSDAAVEDSGASLPPVTDENHITLKAESQLEDLGYCVFNKYTVPLPSPVQDSESLTGEICPFYEGRDEKMRRGLAADLSLIEMKLAAAEKSKEEFSSEKDLSPNISSDTTVLGREIEQEKKGSDKLATVLEKSEEQVDSKEIHSLEDTKHEKKRPDAILEMKEEGTVDKLHIAGPESDIANDKALEKEVTKEKDLVSLLTQKEEKSLNIIPKVAEIETPHRPDYDAIKHDMDVAARKADQEYQTQLHTEINEFAAAGSKQDTDSVKKAEPEAQEAQQQQKDLASLPKEPKEADVLSRTEPSQMKDSTKLSESEIKEKVPKPDLVHQEAVDKEESYESSGEHDQIQEDLNGESLKPDYGKIKPQKPPVSEEEITSKLPVEEPSIELAISKADSVQEETADIQMESIPQPKEEIKETTPDIAVRPAETTEQLLSDLTPESVEGKEHKEEEDDDKHQLMSDVHQEIELEAPAGEEALTEAGPEEMPELKGVIESVVTIEDDFITVVQMTVEESESASHSVRFAAAQQEEMERRDFLPEEEVEMEMEEPADIQAEPKEGSLEAPASPEKEEILLTDYKTEIYDDFKDETTIDDSVMDTDSLWADTQDDDRSIMTEQLETVPKEEKTERELRRSSLDKHKKEKPFKTGRGRISTPERKIAKKEPSTLSRDEVRRKKAVYKKAELAKKSEVQAHSPSRKIILKPAIKYTRPTHLSCVKRKQTAAGGETNQAPGVFKQAKEKLSDGITKSPEKRSSLPRPSSILPTRRGVSGDRDRDENSFSLNSSISSSVRRTTRSEPIRSRTGKSGTSTPTTPGSTAITPGTPPSYSSRTPGTPGTPSYSRTPHTPGTPKSAILVPTEKKVAIIRTPPKSPATPKQLRLINQPLPDLKNVRSKIGSTDNIRYQPKGGQIQIVTKKIDLSHVTSKCGSLKNIRHRPGGGRVKIESVKLDFKEKAHAKVGSLENAHHTPGGGNIKIDSQKLNFREHAKARVDHGAEIITQSPGRSSVPSPRRLSNVSSSGSINLLESPQLATLAEDVTAALAKQGL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN99</accession>
    <name>A0A452HN99_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Roundabout guidance receptor 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018800.1">
        <property type="protein sequence ID" value="ENSGAGP00000016472.1"/>
        <property type="gene ID" value="ENSGAGG00000012321.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008046">
        <property type="term" value="F:axon guidance receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016199">
        <property type="term" value="P:axon midline choice point recognition"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035385">
        <property type="term" value="P:Roundabout signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="CDD" id="cd05725">
        <property type="entry name" value="IgI_3_Robo"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05726">
        <property type="entry name" value="IgI_4_Robo"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd20952">
        <property type="entry name" value="IgI_5_Robo"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000053">
        <property type="entry name" value="Roundabout guidance receptor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000055">
        <property type="entry name" value="roundabout homolog 1 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000065">
        <property type="entry name" value="roundabout homolog 1 isoform X3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000008">
        <property type="entry name" value="roundabout homolog 2 isoform X2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000043">
        <property type="entry name" value="roundabout homolog 2 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000058">
        <property type="entry name" value="roundabout homolog 2 isoform X3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013098">
        <property type="entry name" value="Ig_I-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013106">
        <property type="entry name" value="Ig_V-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051170">
        <property type="entry name" value="Neural/epithelial_adhesion"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032986">
        <property type="entry name" value="Robo1_Ig-like3"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12231">
        <property type="entry name" value="CTX-RELATED TYPE I TRANSMEMBRANE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12231:SF243">
        <property type="entry name" value="ROUNDABOUT HOMOLOG 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07679">
        <property type="entry name" value="I-set"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13927">
        <property type="entry name" value="Ig_3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00406">
        <property type="entry name" value="IGv"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="8" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="6" id="KW-0472">Membrane</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0732">Signal</keyword>
    <keyword evidence="2" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5" id="KW-1133">Transmembrane helix</keyword>
    <feature type="domain" description="Ig-like" evidence="10">
        <location>
            <begin position="1"/>
            <end position="78"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="10">
        <location>
            <begin position="83"/>
            <end position="167"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="10">
        <location>
            <begin position="172"/>
            <end position="267"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="10">
        <location>
            <begin position="276"/>
            <end position="362"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="11">
        <location>
            <begin position="384"/>
            <end position="478"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="11">
        <location>
            <begin position="497"/>
            <end position="594"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="11">
        <location>
            <begin position="599"/>
            <end position="695"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="1"/>
            <end position="32"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="896"/>
            <end position="918"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="939"/>
            <end position="1148"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="1168"/>
            <end position="1210"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="1233"/>
            <end position="1441"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="941"/>
            <end position="958"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="959"/>
            <end position="975"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="9">
        <location>
            <begin position="998"/>
            <end position="1008"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="9">
        <location>
            <begin position="1040"/>
            <end position="1049"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="1068"/>
            <end position="1082"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="1094"/>
            <end position="1106"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="9">
        <location>
            <begin position="1109"/>
            <end position="1120"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="9">
        <location>
            <begin position="1135"/>
            <end position="1148"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="1197"/>
            <end position="1210"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="1256"/>
            <end position="1265"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="9">
        <location>
            <begin position="1273"/>
            <end position="1284"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="9">
        <location>
            <begin position="1294"/>
            <end position="1304"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="1330"/>
            <end position="1387"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="1406"/>
            <end position="1415"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004167"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016472.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="14">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1465" mass="160246" checksum="525AF0131C3E8D92" modified="2019-05-08" version="1">MVAVGEPAVMECQPPRGHPEPTISWKKDGSPLDDKDERITIRGGKLMITYTKKNDAGKYVCVGTNMVGERESEVAELTVLERPSFVKRPSNLAVTVEDSAEFKCEARGDPVPTVRWRKDDGELPKARYEIRDDHTLKIRKVMAGDMGSYTCVAENMVGKAEASATLTVQEPPHFVVKPRDQVAALGRTVTFQCEATGNPQPAIFWRREGSQNLLFSYQPPQSSSRFSVSQTGDLTITSVQRSDVGYYICQTLNVAGSITTKAYLEVTDVIADRPPPVIRQGPVNQTVAVDGTLVLSCVATGTLVPTILWRKDGVLISTQDSRIKQLETGALQIRYAKLGDTGRYTCIASTPSGEATWSAYIEVQEFGVPVQPPRPTDPNLIPSAPSKPEVTDVSRNTVTLSWQPNLNSGATPTSYIIEAFSHASGSSWQTVAENVKTESFAVKGLKPNAIYLFLVRAANAYGLSDPSQISDPVKTQDVPPTSQGVDHKQVQRELGDVVLHLHNPTILSSSSIEVHWTVDQQSQYIQGYKILYRPATAPRGESEWLIFEVRTPSKNSVIIPELKKGVNYEIKARPFFNEFQGADSEVKFAKTLEEAPSAPPQSVSVTKNDGNGTAVVVTWQPPPEDTQNGMVQEYKVWCLGNESRYHINKTVDGSTFSVVIPSLVPGIRYSVEVAASTGAGPGVKSEPQFIQLDSHGNPVSPEDQVSLAQQISDVVKQPAFIAGIGAACWIILMVFSIWLYRHQKNRNGLTSTYAGIRKVTYQRGGEAVSSGGRPGLLNISEPAPQPWLADTWPNTGNNHNDCSINCCTSANGNSDSNLTTYSRPADCIANYNNQLESKQTNLMLPESTVYGDVDLSNKINEMKTFNSPNLKDGRFVNQPGQPTPYATTQLIQSSISNNVNSSSGDTNEKHWKPSVQQKQEVAPIQYNIMEQNKLNKDYRVNDNNAPTIPYNQSYDQNTGGSYNSSDRGSSTSGSQGHKKGARTPKVPKQAGMNWADLLPPPPAHPPPHSNSEEYSLSVDESYDQEIPCPVPPARMYLQQDELEEEDDDERGPTPPVRGAASSPAAVSYSHQSTATLTPSPQEELQPMLQDCQEEIGHMQHQPDRRRQPVSPPPPPRPISPPHTYGYISGPLVSDMDTDAPEEEEDEAEIEVAKMQNRRLLLRGLEQTPASSVGDLESSVTGSMINGWGSASEEDNISSGRSSVSSSDGSFFTDADFAQAVAAAAEYAGLKVARRQVQDAAGGRRHFHASHCPRPTSPVSTDSNMSAVVIQKIRPAKKQKHQPGHLRREVYTDDLPPPPVPPPAIKSPTAQSKSQLEVRPVMLPKLASIEPRTDRSTERKGASYKGREGADGRQHSDVRTNSGERRELQEQQNDGKLRGNKAAKREGTPAKTHLLQEDILPYCRPTFPTSNNPRDPSSSSSMSSRGSGGRQRADQTNIGRRNAAEMQILGAYEQGEEEEELEETES</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPV9</accession>
    <name>A0A452IPV9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">EDR1/CTR1/ARMC3-like peptidase-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IPV9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034003.1">
        <property type="protein sequence ID" value="ENSGAGP00000029944.1"/>
        <property type="gene ID" value="ENSGAGG00000021609.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000225">
        <property type="entry name" value="Armadillo"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052441">
        <property type="entry name" value="Armadillo-Ser/Thr_Kinase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055164">
        <property type="entry name" value="EDR1/CTR1/ARMC3-like_pept-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46618">
        <property type="entry name" value="ARMADILLO REPEAT-CONTAINING PROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46618:SF1">
        <property type="entry name" value="ARMADILLO REPEAT-CONTAINING PROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00514">
        <property type="entry name" value="Arm"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14381">
        <property type="entry name" value="EDR1_CTR1_ARMC3_pept"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13646">
        <property type="entry name" value="HEAT_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00185">
        <property type="entry name" value="ARM"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50176">
        <property type="entry name" value="ARM_REPEAT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0677">Repeat</keyword>
    <feature type="repeat" description="ARM" evidence="2">
        <location>
            <begin position="150"/>
            <end position="192"/>
        </location>
    </feature>
    <feature type="repeat" description="ARM" evidence="2">
        <location>
            <begin position="356"/>
            <end position="398"/>
        </location>
    </feature>
    <feature type="domain" description="EDR1/CTR1/ARMC3-like peptidase-like" evidence="4">
        <location>
            <begin position="712"/>
            <end position="850"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="622"/>
            <end position="692"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="3">
        <location>
            <begin position="654"/>
            <end position="664"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="665"/>
            <end position="685"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00259"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="Pfam" id="PF14381"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029944.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="863" mass="94895" checksum="11BF95969B7D2812" modified="2019-05-08" version="1">MGKKVKKEVEPPPKDVFDPLSIESKKAVTVVLMLNSPEEEVLAKACDAIYKFASKGDENKVTLLSLGAVEPLFKLVSHEDPLVRRNAIMVFGIMASNNDVRKLLRELDVTNSVIARLAPEEDIVVHEFASLCLAHMAVEYTSKVQIFEQGGLEPLIRLLSSPDPDVKKNSVECIYHLVQDFQSRAAVRELNAIPALLELLKSEYPVVQLLALKTFGVISNDRETRIILRENQGLDHLFKILETKEFNDLHVEALAVVANCLEDVDTMQLLQQTGGLKKLLSFAEMSTIPDIQKNAAKAITKAAYDPENRKILHEEEIEKCLITLLGTDSDGAKAAASQAISAMCENLASKDAFRIQGIPQLVQLLSSENEEVKEAAAIALANLTTASPNNASAVAEAEGIEPLINVLSAKRDGTVANAATVLTNMATQEPLRLSIHGVMNAIVEPLQSTNSLVQSKATLTVAAFGCDADARTELRNASGLEPLVKLLHSKNDEVRRNACWAVMVCASDELTAVELCRLGALDILEEINLSTGRKNNFSEGALDKLLDNNLSQKYSQMGYLSSSNIITDGFYDYGQVKPGVKLLSLEELSIQELNDHRAIILVNAKLPEASLSAEDKSQDLYYGRTTSSSSIRKSSKERASSAVASPVEEKQESSGRSPSSLSKSSTREKGSRKGKGKKEEDKPKEEEELQTMPKVQQEVILEKLLWLPPFDSILLDYIIDASKIILPLTTTKEQVIALALFVADKMGGPIERDKLHEFSWELHISEIEFELKCNVVPIGKIKKGTFYHRALLFKVIADRIGIGCSLVRGEYGRAWNEVKLVDDSPQGITGLLLPPQTYIVDLMYQPGCLMKQGSAEADHYLRI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPW2</accession>
    <name>A0A452IPW2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Phosphodiesterase</fullName>
            <ecNumber evidence="10">3.1.4.-</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="6">
            <text>3',5'-cyclic AMP + H2O = AMP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:25277"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:58165"/>
            <dbReference type="ChEBI" id="CHEBI:456215"/>
            <dbReference type="EC" id="3.1.4.53"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="6">
            <dbReference type="Rhea" id="RHEA:25278"/>
        </physiologicalReaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="10">
            <name>a divalent metal cation</name>
            <dbReference type="ChEBI" id="CHEBI:60240"/>
        </cofactor>
        <text evidence="10">Binds 2 divalent metal cations per subunit. Site 1 may preferentially bind zinc ions, while site 2 has a preference for magnesium and/or manganese ions.</text>
    </comment>
    <comment type="pathway">
        <text evidence="1">Purine metabolism; 3',5'-cyclic AMP degradation; AMP from 3',5'-cyclic AMP: step 1/1.</text>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the cyclic nucleotide phosphodiesterase family. PDE4 subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.1.4.-" evidence="10"/>
    <dbReference type="AlphaFoldDB" id="A0A452IPW2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033935.1">
        <property type="protein sequence ID" value="ENSGAGP00000029883.1"/>
        <property type="gene ID" value="ENSGAGG00000021546.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00762">
        <property type="reaction ID" value="UER00747"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004115">
        <property type="term" value="F:3',5'-cyclic-AMP phosphodiesterase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006198">
        <property type="term" value="P:cAMP catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00077">
        <property type="entry name" value="HDc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.1300.10:FF:000001">
        <property type="entry name" value="Phosphodiesterase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1300.10">
        <property type="entry name" value="3'5'-cyclic nucleotide phosphodiesterase, catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003607">
        <property type="entry name" value="HD/PDEase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040844">
        <property type="entry name" value="PDE4_UCR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023088">
        <property type="entry name" value="PDEase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002073">
        <property type="entry name" value="PDEase_catalytic_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036971">
        <property type="entry name" value="PDEase_catalytic_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023174">
        <property type="entry name" value="PDEase_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11347">
        <property type="entry name" value="CYCLIC NUCLEOTIDE PHOSPHODIESTERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18100">
        <property type="entry name" value="PDE4_UCR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00233">
        <property type="entry name" value="PDEase_I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00387">
        <property type="entry name" value="PDIESTERASE1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00471">
        <property type="entry name" value="HDc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109604">
        <property type="entry name" value="HD-domain/PDEase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00126">
        <property type="entry name" value="PDEASE_I_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51845">
        <property type="entry name" value="PDEASE_I_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0114">cAMP</keyword>
    <keyword evidence="4 10" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="3 9 10" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="PDEase" evidence="12">
        <location>
            <begin position="265"/>
            <end position="567"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="36"/>
            <end position="58"/>
        </location>
    </feature>
    <feature type="active site" description="Proton donor" evidence="7">
        <location>
            <position position="341"/>
        </location>
    </feature>
    <feature type="binding site" evidence="8">
        <location>
            <begin position="341"/>
            <end position="345"/>
        </location>
        <ligand>
            <name>AMP</name>
            <dbReference type="ChEBI" id="CHEBI:456215"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="345"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="381"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="8">
        <location>
            <position position="382"/>
        </location>
        <ligand>
            <name>AMP</name>
            <dbReference type="ChEBI" id="CHEBI:456215"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="382"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="382"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <label>2</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="8">
        <location>
            <position position="499"/>
        </location>
        <ligand>
            <name>AMP</name>
            <dbReference type="ChEBI" id="CHEBI:456215"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="499"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="8">
        <location>
            <position position="550"/>
        </location>
        <ligand>
            <name>AMP</name>
            <dbReference type="ChEBI" id="CHEBI:456215"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004703"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009517"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023149"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00033681"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PIRSR" id="PIRSR623088-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PIRSR" id="PIRSR623088-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PIRSR" id="PIRSR623088-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="RuleBase" id="RU363067"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS51845"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029883.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="567" mass="64513" checksum="C70E9850011D34D8" modified="2019-05-08" version="1">MKFRFKQMSRSESGLPKSLRMLGSIVFIHFDVENGPSPGRSPLDPQASSSSGLVLHTTFPGHSQRRESFLYRSDSDYDLSPKTMSRNSSLPSEQHGDDLIVTPFAQVLASLRSVRNNFTLLTNLHGTSNKRSPAASQPPVSRVSLQEESYQKLAMETLEELDWCLDQLETIQTYRSVSEMASNKFKRMLNRELTHLSEMSRSGNQVSEYISNTFLDKQNDVEIPSPTQKDREKKKKQQLMTQISGVKKLMHSSSLNNTSISRFGVKTEKEDHLAKELEDLNKWGLNIFNVARYSHSRPLTCIMYAIFQERDLLKTFKISSDTFVTYMMTLEDHYHSDVAYHNSLHAADVAQSTHVLLSTPALDAVFTDLEILAAIFAAAIHDVDHPGVSNQFLINTNSELALMYNDESVLENHHLAVGFKLLQEEHCDIFQNLSKKQRQTLRKMVIDMVLATDMSKHMSLLADLKTMVETKKVTSSGVLLLDNYTDRIQVLRNMVHCADLSNPTKSLELYRQWTDRIMEEFFQQGDKERERGMEISPMCDKHTASVEKSQVLYIGLAPLKTGIFLTL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPW5</accession>
    <name>A0A452IPW5_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="12">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 7">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 7">Belongs to the ETS family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPW5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034097.1">
        <property type="protein sequence ID" value="ENSGAGP00000030027.1"/>
        <property type="gene ID" value="ENSGAGG00000021670.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043565">
        <property type="term" value="F:sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd08540">
        <property type="entry name" value="SAM_PNT-ERG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.50:FF:000010">
        <property type="entry name" value="Fli-1 proto-oncogene, ETS transcription factor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.10:FF:000039">
        <property type="entry name" value="Friend leukemia integration 1 transcription factor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.50">
        <property type="entry name" value="Transcription Factor, Ets-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.10">
        <property type="entry name" value="Winged helix-like DNA-binding domain superfamily/Winged helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000418">
        <property type="entry name" value="Ets_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046328">
        <property type="entry name" value="ETS_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003118">
        <property type="entry name" value="Pointed_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001660">
        <property type="entry name" value="SAM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013761">
        <property type="entry name" value="SAM/pointed_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036388">
        <property type="entry name" value="WH-like_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036390">
        <property type="entry name" value="WH_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11849">
        <property type="entry name" value="ETS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11849:SF216">
        <property type="entry name" value="TRANSCRIPTIONAL REGULATOR ERG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00178">
        <property type="entry name" value="Ets"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02198">
        <property type="entry name" value="SAM_PNT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00454">
        <property type="entry name" value="ETSDOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00413">
        <property type="entry name" value="ETS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00251">
        <property type="entry name" value="SAM_PNT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47769">
        <property type="entry name" value="SAM/Pointed domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46785">
        <property type="entry name" value="Winged helix' DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00345">
        <property type="entry name" value="ETS_DOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00346">
        <property type="entry name" value="ETS_DOMAIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50061">
        <property type="entry name" value="ETS_DOMAIN_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51433">
        <property type="entry name" value="PNT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50105">
        <property type="entry name" value="SAM_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 7" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="6 7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0804">Transcription</keyword>
    <keyword evidence="3" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="PNT" evidence="11">
        <location>
            <begin position="120"/>
            <end position="206"/>
        </location>
    </feature>
    <feature type="domain" description="SAM" evidence="10">
        <location>
            <begin position="141"/>
            <end position="186"/>
        </location>
    </feature>
    <feature type="domain" description="ETS" evidence="9">
        <location>
            <begin position="291"/>
            <end position="371"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="41"/>
            <end position="63"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="235"/>
            <end position="283"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="41"/>
            <end position="54"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="250"/>
            <end position="261"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005562"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU004019"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50061"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50105"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS51433"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030027.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="459" mass="51947" checksum="4BC8999420178186" modified="2019-05-08" version="1">MIQTVPDPAAHIKEALSVVSEDQSLFECAYGTQHLTKTEMTASSSSEYGQTSKMSPRVPQQDWLSQPPARVTIKMECNPNQINGSRNSPDDCSVAKGGKMVSGSDNVGMNYGSYMEEKHVPPPNMTTNERRVIVPADPTLWSTDHVRQWLEWAVKEYGLPDVDILLFQNIDGKELCKMTKDDFQRLTPSYNADILLSHLHYLRERGAAFIFPNTSVYPEATQRITTRPDLSYEQARRSAWTSHNHPAPQSKATQPSSSTVPKTEDQRPQLDPYQILGPTSSRLANPGSGQIQLWQFLLELLSDSSNSNCITWEGTNGEFKMTDPDEVARRWGERKSKPNMNYDKLSRALRYYYDKNIMTKVHGKRYAYKFDFHGIAQALQPHPPESSMYKYPTDLPYMSPYHAHPQKMNFVAPHPPALPVTSSSFFAAPTTYWNSPTGSIYPNTRLPATHMPSHLGTYY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPW7</accession>
    <name>A0A452IPW7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Fringe-like glycosyltransferase domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="9">Endomembrane system</location>
            <topology evidence="9">Single-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type II membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the glycosyltransferase 31 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPW7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033940.1">
        <property type="protein sequence ID" value="ENSGAGP00000029888.1"/>
        <property type="gene ID" value="ENSGAGG00000021571.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012505">
        <property type="term" value="C:endomembrane system"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016757">
        <property type="term" value="F:glycosyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.550.50:FF:000021">
        <property type="entry name" value="Beta 3-glucosyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.550.50">
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003378">
        <property type="entry name" value="Fringe-like_glycosylTrfase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10811">
        <property type="entry name" value="FRINGE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02434">
        <property type="entry name" value="Fringe"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0328">Glycosyltransferase</keyword>
    <keyword evidence="8" id="KW-0472">Membrane</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0732">Signal</keyword>
    <keyword evidence="6" id="KW-0735">Signal-anchor</keyword>
    <keyword evidence="4" id="KW-0808">Transferase</keyword>
    <keyword evidence="5" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="10">
        <location>
            <begin position="1"/>
            <end position="28"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019412158" description="Fringe-like glycosyltransferase domain-containing protein" evidence="10">
        <location>
            <begin position="29"/>
            <end position="468"/>
        </location>
    </feature>
    <feature type="domain" description="Fringe-like glycosyltransferase" evidence="11">
        <location>
            <begin position="124"/>
            <end position="215"/>
        </location>
    </feature>
    <feature type="domain" description="Fringe-like glycosyltransferase" evidence="11">
        <location>
            <begin position="273"/>
            <end position="364"/>
        </location>
    </feature>
    <feature type="domain" description="Fringe-like glycosyltransferase" evidence="11">
        <location>
            <begin position="366"/>
            <end position="443"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004606"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008661"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022676"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00037847"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF02434"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029888.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="468" mass="53577" checksum="0A7A9C579A6D80AA" modified="2019-05-08" version="1">MPWERGGGGWWLLPALLLLPPLCAPALAISVAAEDAEEQIEQLVSQESGKSGKSNQINIELQEIVFVIQSQSNSFHSKRAEDLERNILKQAEELGKELPRILLIHQMDRHDGAWTILPLMQDFSASYGRNSSWIFFCEEDTRIQIVKLLETLRRFDKSKEWFLGKALYDEESTIIHHYAFAENPTVFKFPDFAAGWALSIPLVNKLANRLKSEPLKSDFTIDLKHEIALYIWQKGNGPHLTSVPEFCTDDLNSYKAVHCATMFSNFLPVCGDPVKKEDIFVAVKTCRKFHRDRIPIVKQTWEGEAALVEYYSDYAEISIPTIDLGVPNTERGHCGKTFAILERFLNHSSSKTPWLVIVDDDTLIRMVFSREAIQKLLASKCQCYSNDAPDDMVLGMCFSGLGIPITHCPLFHQARPVDYPKDYLAHQIPISFHKHWNIDPVKVYFTWLAPKVEDSFTEKKQRHIREDL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPW8</accession>
    <name>A0A452IPW8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">Glycosyltransferase 2-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000034064.1">
        <property type="protein sequence ID" value="ENSGAGP00000029995.1"/>
        <property type="gene ID" value="ENSGAGG00000021650.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008376">
        <property type="term" value="F:acetylgalactosaminyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019276">
        <property type="term" value="P:UDP-N-acetylgalactosamine metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006047">
        <property type="term" value="P:UDP-N-acetylglucosamine metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00761">
        <property type="entry name" value="Glyco_tranf_GTA_type"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.550.10">
        <property type="entry name" value="Spore Coat Polysaccharide Biosynthesis Protein SpsA, Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001173">
        <property type="entry name" value="Glyco_trans_2-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029044">
        <property type="entry name" value="Nucleotide-diphossugar_trans"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15046:SF2">
        <property type="entry name" value="BETA-1,4 N-ACETYLGALACTOSAMINYLTRANSFERASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15046">
        <property type="entry name" value="GLYCO_TRANS_2-LIKE DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00535">
        <property type="entry name" value="Glycos_transf_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53448">
        <property type="entry name" value="Nucleotide-diphospho-sugar transferases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Glycosyltransferase 2-like" evidence="1">
        <location>
            <begin position="181"/>
            <end position="292"/>
        </location>
    </feature>
    <evidence type="ECO:0000259" key="1">
        <source>
            <dbReference type="Pfam" id="PF00535"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029995.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="421" mass="47783" checksum="51CC3216AD4475B7" modified="2019-05-08" version="1">GLLPPSSEDRAESSSYWPGIQLRRQAPLPAAAQKYPSMTQDIVIAKPNVPLSYPIQGVEVMPLHTIMIPGIRGPSVPGTIQIHRRTGQEXTLANVSDDVVRGRGKKELLILTSDVELLNFILEHVTYTSVVYQLSAVDMMSFKSRDHVVQFPVIIRQPSLPKLIDPGADRKISSLVTITTKTFLRYHKLRFLIKSIRQYYPDIKIIVADDSEVPEKIIEENIEHYIMPFGKGWFAGRNLAISQVTTKYYLWVDDDYLFTENTKIEKLLDVLERTNLDMVGGSVKGDTYSFQLLYEQSDDGYCLHLRYGSFHKLDGFPNCVVTSGVVNFYLAHTDKSRYVSYDPLLQRVAHSEFFVDGLGILLVGSCSDVSVGHQDHNPASDPNLAKIEDQYKTFRDNTDAQVQFKLALHYFKNRLKCYSTR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPZ7</accession>
    <name>A0A452IPZ7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Arrestin domain-containing protein 3</fullName>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">Adapter protein that plays a role in regulating cell-surface expression of adrenergic receptors and probably also other G protein-coupled receptors. Plays a role in NEDD4-mediated ubiquitination and endocytosis af activated ADRB2 and subsequent ADRB2 degradation. May recruit NEDD4 to ADRB2. Alternatively, may function as adapter protein that does not play a major role in recruiting NEDD4 to ADRB2, but rather plays a role in a targeting ADRB2 to endosomes.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cell membrane</location>
            <topology evidence="3">Peripheral membrane protein</topology>
            <orientation evidence="3">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Early endosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Lysosome</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the arrestin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPZ7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033979.1">
        <property type="protein sequence ID" value="ENSGAGP00000029921.1"/>
        <property type="gene ID" value="ENSGAGG00000021451.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005769">
        <property type="term" value="C:early endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005764">
        <property type="term" value="C:lysosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.640:FF:000005">
        <property type="entry name" value="Arrestin domain-containing protein 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.640:FF:000007">
        <property type="entry name" value="Arrestin domain-containing protein 3 mRNA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.640">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014752">
        <property type="entry name" value="Arrestin-like_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011021">
        <property type="entry name" value="Arrestin-like_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011022">
        <property type="entry name" value="Arrestin_C-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050357">
        <property type="entry name" value="Arrestin_domain-protein"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014756">
        <property type="entry name" value="Ig_E-set"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11188">
        <property type="entry name" value="ARRESTIN DOMAIN CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11188:SF49">
        <property type="entry name" value="ARRESTIN DOMAIN-CONTAINING PROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02752">
        <property type="entry name" value="Arrestin_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00339">
        <property type="entry name" value="Arrestin_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01017">
        <property type="entry name" value="Arrestin_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81296">
        <property type="entry name" value="E set domains"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9" id="KW-0967">Endosome</keyword>
    <keyword evidence="11" id="KW-0458">Lysosome</keyword>
    <keyword evidence="10" id="KW-0472">Membrane</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="Arrestin C-terminal-like" evidence="15">
        <location>
            <begin position="144"/>
            <end position="271"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="350"/>
            <end position="371"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="362"/>
            <end position="371"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004371"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004412"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004413"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00005298"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023228"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00041132"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00045942"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="SMART" id="SM01017"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029921.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="371" mass="41701" checksum="12A95DDC37802C3A" modified="2019-05-08" version="1">MVLGKVKMKSLKIHARGHAKVRWTESRNAGSNTAYTQNYTEEVEYFNHKDILIGHERGLNTIHSGRHEYAFSFELPQTPLATSFEGRHGSVRYWVKAELHRPWLLPVKLKKEFTVFEHIDINTPSLLSPQAGTKEKTLCCWFCTSGPISLSAKIERKGFTPGESIQIFAEIENCSSRMVVPKAAIYQTQAFYAKGKMKEVKQLVANLRGESLSSGKTETWNGKQLKIPPVSPSILDCSIIRVEYSLMVYVDIPGAMDLFLNLPLVIGTIPLHPFGSRTSSVSSQCSMNMNWLGLPLPERPEAPPSYAEVVTEEQRQSSLVPIAACDDFERALQGPLFAYIQEFRFLPPPLYSEIDPNPDQPTDDRPSCPSR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPZ9</accession>
    <name>A0A452IPZ9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Sugar phosphate transporter domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPZ9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034096.1">
        <property type="protein sequence ID" value="ENSGAGP00000030026.1"/>
        <property type="gene ID" value="ENSGAGG00000021671.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004853">
        <property type="entry name" value="Sugar_P_trans_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050186">
        <property type="entry name" value="TPT_transporter"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11132">
        <property type="entry name" value="SOLUTE CARRIER FAMILY 35"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03151">
        <property type="entry name" value="TPT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4 5" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 5" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="3 5" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="20"/>
            <end position="37"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="49"/>
            <end position="67"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="109"/>
            <end position="130"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="137"/>
            <end position="156"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="194"/>
            <end position="214"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="226"/>
            <end position="247"/>
        </location>
    </feature>
    <feature type="domain" description="Sugar phosphate transporter" evidence="6">
        <location>
            <begin position="28"/>
            <end position="296"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="Pfam" id="PF03151"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030026.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="308" mass="34545" checksum="7DA2B487BD24F563" modified="2019-05-08" version="1">MSARQAEEEPAAAHPPLPRLLSAFFYGTCSFLIVLVNKTVLTTYRFPSPMVLGIGQMATTILILYVSKLNKIIHFPDFDKSIPVKLFPLPLLYVGNHISGLTSTSKLSLPMFTVLRKFTIPLTLLLEIIILGKRYSLGIIVSVFTIILGAFIAAGSDLSFNLEGYTFVLLNDIFTAANGVYTKQKMDPKELGKYGILFYNAYFMVIPTMIISFSTGDLQQVIDFKYWTNFFFLFQFLLSCFMGFLLMYSTVLCSHYNSALTTSVVGAIKNISIAYIGMLVGGDYIFSILNFIGLNIWYCITIFCQPYI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ74</accession>
    <name>A0A452IQ74_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="21">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cell projection</location>
            <location evidence="1">Cilium</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the polycystin family.</text>
    </comment>
    <comment type="caution">
        <text evidence="13">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000034132.1">
        <property type="protein sequence ID" value="ENSGAGP00000030060.1"/>
        <property type="gene ID" value="ENSGAGG00000021465.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005929">
        <property type="term" value="C:cilium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005261">
        <property type="term" value="F:monoatomic cation channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006816">
        <property type="term" value="P:calcium ion transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00037">
        <property type="entry name" value="CLECT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00146">
        <property type="entry name" value="PKD"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="CDD" id="cd01752">
        <property type="entry name" value="PLAT_polycystin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000825">
        <property type="entry name" value="Polycystin 1, transient receptor potential channel interacting"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.60.20:FF:000012">
        <property type="entry name" value="polycystin-1 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.70">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.100.10">
        <property type="entry name" value="Mannose-Binding Protein A, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.60.20">
        <property type="entry name" value="PLAT/LH2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001304">
        <property type="entry name" value="C-type_lectin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016186">
        <property type="entry name" value="C-type_lectin-like/link_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016187">
        <property type="entry name" value="CTDL_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057244">
        <property type="entry name" value="GAIN_B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000203">
        <property type="entry name" value="GPS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000434">
        <property type="entry name" value="PC1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022409">
        <property type="entry name" value="PKD/Chitinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002859">
        <property type="entry name" value="PKD/REJ-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013122">
        <property type="entry name" value="PKD1_2_channel"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000601">
        <property type="entry name" value="PKD_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035986">
        <property type="entry name" value="PKD_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001024">
        <property type="entry name" value="PLAT/LH2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036392">
        <property type="entry name" value="PLAT/LH2_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042060">
        <property type="entry name" value="PLAT_polycystin1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006228">
        <property type="entry name" value="Polycystin_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046791">
        <property type="entry name" value="Polycystin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014010">
        <property type="entry name" value="REJ_dom"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00864">
        <property type="entry name" value="PCC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46730">
        <property type="entry name" value="POLYCYSTIN-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46730:SF3">
        <property type="entry name" value="POLYCYSTIN-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00059">
        <property type="entry name" value="Lectin_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00801">
        <property type="entry name" value="PKD"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08016">
        <property type="entry name" value="PKD_channel"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01477">
        <property type="entry name" value="PLAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20519">
        <property type="entry name" value="Polycystin_dom"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02010">
        <property type="entry name" value="REJ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00500">
        <property type="entry name" value="POLYCYSTIN1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00034">
        <property type="entry name" value="CLECT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00303">
        <property type="entry name" value="GPS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00308">
        <property type="entry name" value="LH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00089">
        <property type="entry name" value="PKD"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56436">
        <property type="entry name" value="C-type lectin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49723">
        <property type="entry name" value="Lipase/lipooxygenase domain (PLAT/LH2 domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49299">
        <property type="entry name" value="PKD domain"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50041">
        <property type="entry name" value="C_TYPE_LECTIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50221">
        <property type="entry name" value="GAIN_B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50093">
        <property type="entry name" value="PKD"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50095">
        <property type="entry name" value="PLAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51111">
        <property type="entry name" value="REJ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="12" id="KW-0966">Cell projection</keyword>
    <keyword evidence="8" id="KW-0969">Cilium</keyword>
    <keyword evidence="10" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9 15" id="KW-0472">Membrane</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5 15" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 15" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="2665"/>
            <end position="2687"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="2873"/>
            <end position="2894"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="2914"/>
            <end position="2936"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3155"/>
            <end position="3177"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3183"/>
            <end position="3202"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3270"/>
            <end position="3289"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3512"/>
            <end position="3531"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3543"/>
            <end position="3570"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3590"/>
            <end position="3614"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3640"/>
            <end position="3662"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3668"/>
            <end position="3687"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3699"/>
            <end position="3724"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="16">
        <location>
            <begin position="11"/>
            <end position="125"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="17">
        <location>
            <begin position="559"/>
            <end position="621"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="17">
        <location>
            <begin position="651"/>
            <end position="723"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="17">
        <location>
            <begin position="741"/>
            <end position="803"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="17">
        <location>
            <begin position="912"/>
            <end position="985"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="17">
        <location>
            <begin position="1014"/>
            <end position="1069"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="17">
        <location>
            <begin position="1087"/>
            <end position="1151"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="17">
        <location>
            <begin position="1323"/>
            <end position="1409"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="17">
        <location>
            <begin position="1426"/>
            <end position="1486"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="17">
        <location>
            <begin position="1662"/>
            <end position="1745"/>
        </location>
    </feature>
    <feature type="domain" description="REJ" evidence="20">
        <location>
            <begin position="1748"/>
            <end position="2441"/>
        </location>
    </feature>
    <feature type="domain" description="GAIN-B" evidence="19">
        <location>
            <begin position="2462"/>
            <end position="2655"/>
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    </feature>
    <feature type="domain" description="PLAT" evidence="18">
        <location>
            <begin position="2710"/>
            <end position="2825"/>
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    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="3777"/>
            <end position="3806"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="3902"/>
            <end position="3925"/>
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    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="3955"/>
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        </location>
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    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="3794"/>
            <end position="3806"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="14">
        <location>
            <begin position="3960"/>
            <end position="3977"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
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    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
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    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007200"/>
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    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
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    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
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    <evidence type="ECO:0000256" key="6">
        <source>
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        <source>
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        <source>
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        <source>
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        <source>
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    </evidence>
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        <source>
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        <source>
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        <source>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030060.1"/>
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    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="3977" mass="441876" checksum="9143C3F61DDC98DC" modified="2019-05-08" version="1">MCPSDGLVFPGNNHCYQLVVEKAQWLEAQQHCQVHGDGELAFVSSPEMKSFLVSHVIRGLDVWIGFNDFASAGAQQRGEGFNLESCQNWLPGEPHPSNADHCVRMGPTGQCNTDLCMAKHSYVCEYKPKGILLNAENFYVGDPAFDTPRSVRYVTREASLPAPLDTVETMVFPSLEFQQEAFLTALEFVTQELQRSIHVRFQVQRPTRREGYKEGKNATYPFHTSQDIGNWTLLECPAGFQWCHLTNTCLSLSSCCTPVECANDSITNSTNSLSWHNESQLSYELLKEFLFTIPANTSAQYLVMFKGEDTFVRPKDVFSIQHDAAPGLFLQCQPSTLSPYRMNYLSTNSSDWVPSLSANLSGAIWKNDTVCSLRVLYATEQITPLISQPNPGLEHHGLYTVKATVDNGVFSANLSCSFWVTSLVSGLRVIHPSPHSGKVYVPTNHSSLVVKISSGVNATASWLGENRTFPFEGACPPAIVPLTADCTRDTNDTWFSVVELSGGGESVSTRLLMVENSVSSQNITVKVKVEEPIRGLRATPDPETRVLLNTRVSYIPVMEAGSDVTFRWTVDDKPSFTFYNVVFNVIYQSPAVYKLSLTASNHVSNVTVDYNVTVEKMNRMKDLTVSEIPLILPQNTTVELTARVQIDSAVDATFLWDFGDGGHEMYQFKPPYNESFLIPDPSVHQVVIEHNVSYTYQVPGEYILVVLVSNSFENLTHQIPVHVHNYLTDLTVEADADVLVVGQPVTFKANPLPSPYGVWYTWDFGDGSLLLLESQPRVTHSYQTKGVYNVSLTANNTISSVETHQCYRVFEEITGLRVSSDEVAEQGISISINASVETGDNITWIFDMGDGTILMSTVPAVEHRYEKDVNCCTINVTAVNPVNSVSQTVPVRVFVLEVLKIEPSSCILEHPNVQFTAYVSGNSENYIFDWTFGDGSSNVTVYGDPVVTHNFTHSGIFSLSLILSSSVNKAHYFTIVCVEPEIVNVTLLPSKQFVSLGKESNFQVSAMPLYQYRYHWDFGTNDSARSGGTEVSYTYKSTGIYLVTVTVSNNVSFNNDTAFVEVQEPVGVAKIEYNGTRVLELNQIYLFSASGNGTKVSYYWDFGDGCNQLGPIITHSYNNTGNYTISLTGWNNVSNCETRLNLTVKRRLQGLTINASRTVVPLNGSVSFVATLLAGNASRYSWILCDRCTPIQGSYTISYTFRSVGTFNVIVTAENEISSLQDSIFVYVLQQIEGLQIASSDLVEDTYFPTNRTLHLQAVVRDGTNISYSWMALKDSTPVQTFSGKIFSLSILEAGNYTIYLKATNMLGSTTVNRTVEFIESIGLLKPSAFPNPAAINASVNISTSVTSGTCLIYIWHLEDGFSPVTIEPFIIHSFQRPGATAVTVTAENKIGSTNASVYIFIQEPIVGLRIGTAELDSYIPSGSVVTFSGELEKGTNVTWLWELPKGLQTGQTVAVTFPKAGFFSVRLNVSNDISWAVASKNITVQDRIEGLELFVSKKVVEPGEQISFVISMSSGSSVSYSVNISGGYSVVLNGSRFTHEFTESGDYLVMVMVQNEVSRECTQVLISVLEAIHDLRLVNCCEKGIPTGTKKSFTAQVGSGSRVSYSWQFSLQKEHSRALVSLAGRSVSYAPAAPGLLEIHLSAFNDLGCLNITRVIQVQDPIVQVSLKPALSFVNRTVLFEASVQPSAWQVVFCWNFGDGSPAQITEVAAANYSYLKAGDYQVEVNATNLVSFMTAHLTVTVRVLECEEPEVELALPTQVVMKRSQRNYLEAQIDLRGCVKYQTEYLWEIYRAPSCLHFGNADRVHLLSVDVSRPQLVIPKLALAVGNYCFVFFVSFGDTPLSKSIFANVTVTPSKLVPIIDGGSYRVWSNTRDLILDGEKSYDPNLNDGEQTPLLYRWSCTSCSKSSPAGCSLHFSATGGVITISKAILEADVEYTFDLSVWKMGLNPESTNQTVLIKKGRVPIVSLECVSCKAQSVYEVSKSSYVYLEGTCRNCQNDSKLGRWAAHSFKNKSLVLDKTTTSTGNTGMNLVLRPGALKDGEGYTFTLHITDLTTGEEGYASIDLLPNRPPIGGLCQLSPNGEVKALMAKVHFECAGWRDTEDEEAPLVYILLATRCKAGQCEEFGMYKGSRSEHSAFLPPGFSANSFLVYVSVLVQDQLGATVMALNRSMTITLPTSAAGVHSLSHWLYDQTDTVLQGLVKQGDPQHVIEYSLALITILNEHEHLMGSEAEAGNELDLRTWTRRNITETLISLNVNTVDDIQQISAALAQCLVASKELLCRSCLTGTLNKLESMMAILQGETTQGIMTPTAIADNILNIMGDLIHLVNTVPQESKLQELCNDPHRLQVASQAYRLSSSLMRILMKSRVLNEEPLELVGGEIVAKGKRSDPFNLLCYENTLDCQFSIPQAFNTTFSDLTDVIQVMFRMDSNPFPFGYISNYTVSTEVASMEFQTNNGTQIPIGSLDAEKAITVMVLNNGMTNISAGSAIIQGKTSVNLVVMTENNNREAGLHFQLTYRVLNDRYISSELEPFIAAYLSPSPAPNENNCSAAKKIGLEAVERRDHKRYTFFISPITDDTTKDYYFNITNHFSWSPVEVTLSLYTSLCQYFCEQERRWKTEGIIPLEETSPDQAVCLTQHLTAFGASLFVPPNSVQFIFPAPGPGLNYIVLLTCAVCFVTYSVAMLIVHKLDLIDINRVGVIPFCGKNGMYKYEILVKTGWGRGSGTTAHVGITLYGVDNKSGHRHLDGENAFHRNSLDVFQIATEQSLGSVWKIRIWHDNKGLSPSWYLQHIIVQDLQTSKSYFFLVNDWLSVESEENDGMVEKEVYAASETELRSFPRIFVAELQRGFFEKHVWLSMWDRPPRSRFTRVQRATCCSALIFLFLCANAVWYGVVGDVHLSHVAVSNLIPLSVDTVAVGLVSSIVVYPLYLVILFLFRMARSKASISHTLTHSDQQSLEIDNYLDSSLLESSFLTFPGIRAEAFSEQTKTDLFLDDSKSLIRWHSNEALLNWPDLLSDPSIMGNTIQKLKRGRASRHLGLEVPLPAEEDSLSLGLHHGQTKYFSASDEDLIRQILADGASGISQDTGQYARAETDLISGLSSVFGEKTETIMLQRLNEKGQNIAAPPREVSRSAKSTWTVVDHTFRKRLLPHWCSYLAHTVSLLFFATSIGVSVWIGVGFTSSVALMWLISGIFSFLASFLVWEPLKVLLEALYFALVAKRLHPEEDDTLVEYPFVEHVSEKINKVRPPQGFALFQAKEEARKVKLLHKMLKSFLIYMLFLLVTLLTNYGDASRNGRAFLLQSSLKQQLGGEEFLRIKRSDEFWAWMSQVLLPSLYNNQSGRESCSTMLGVPRLRQLRLQEAECLDTIHNFLSGTDASVAAKKCTYESNSFNTGDYAVGWKRVSVNSTQTWSYSPPDLAGVWYWGYVSVYDSGGYIQELGATLEESRVHVDYLQQNNWINNMSRAVFVELTQYNPSVDLHAAITLQLEFPTAGRAVAAITVGPFPLLRLSGGVTLQLVMMVFLMAFVVYFVVSESLFIKKEGRAYFAWFGSYSQWLLILLTTCTVVVHLSQATLADQQWAKYLKNRKGFTSFYQVAFLNSVFNSLAASLLFLLTLKAAQQLRFVRQWSVFGKTLRKSMKELAAAGLAFALLVLAYTQLGFLLFSSCSESFHSFSSSLLLLVAAVHGSVNLRLCLPEPSGLYYLFCASYIVLEMWIVLRLFAAVLVHSYREMNFELYRPAFEPQDYEMVELFVRRLKLWMGFSKVKEFRHKVRFEGMEPLPSRSSSDKSFRGPTPSAASDSSRASTSSSQLDGLSLVLSTRESSEVDADVQRLLSLFEMLLAQFDRVNQVTEDVYRMEQQLEDSQSRRTRMKYIQMTEDLLSQYRLGCTDQGPQGLYPDNFSNTDLHPPRDPLLPSPPTSTLGALQSSTGCVPSRIVRASRGLSVASLLSPHKTYATTAPAVRKKKPLRAKNRVHPSIK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ75</accession>
    <name>A0A452IQ75_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="7">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IQ75"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034354.1">
        <property type="protein sequence ID" value="ENSGAGP00000030267.1"/>
        <property type="gene ID" value="ENSGAGG00000021815.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006897">
        <property type="term" value="P:endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016197">
        <property type="term" value="P:endosomal transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00052">
        <property type="entry name" value="EH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.238.10">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011992">
        <property type="entry name" value="EF-hand-dom_pair"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018247">
        <property type="entry name" value="EF_Hand_1_Ca_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002048">
        <property type="entry name" value="EF_hand_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000261">
        <property type="entry name" value="EH_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11216">
        <property type="entry name" value="EH DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11216:SF64">
        <property type="entry name" value="RALBP1-ASSOCIATED EPS DOMAIN-CONTAINING PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12763">
        <property type="entry name" value="EH"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00027">
        <property type="entry name" value="EH"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47473">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00018">
        <property type="entry name" value="EF_HAND_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50222">
        <property type="entry name" value="EF_HAND_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50031">
        <property type="entry name" value="EH"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0106">Calcium</keyword>
    <keyword evidence="3" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="1" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="EH" evidence="5">
        <location>
            <begin position="23"/>
            <end position="114"/>
        </location>
    </feature>
    <feature type="domain" description="EH" evidence="5">
        <location>
            <begin position="260"/>
            <end position="344"/>
        </location>
    </feature>
    <feature type="domain" description="EF-hand" evidence="6">
        <location>
            <begin position="293"/>
            <end position="328"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="149"/>
            <end position="224"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="371"/>
            <end position="402"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="425"/>
            <end position="475"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="491"/>
            <end position="517"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="590"/>
            <end position="620"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="3">
        <location>
            <begin position="623"/>
            <end position="657"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="149"/>
            <end position="164"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="4">
        <location>
            <begin position="167"/>
            <end position="177"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="213"/>
            <end position="224"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="392"/>
            <end position="402"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="425"/>
            <end position="436"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="4">
        <location>
            <begin position="495"/>
            <end position="505"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="507"/>
            <end position="517"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="592"/>
            <end position="603"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50031"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50222"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030267.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="679" mass="74985" checksum="65EDE45746D549AE" modified="2019-05-08" version="1">MEQGPGGAAAGGAGLLLPLSDTEQQRYSELFSLCCSPAPEGGSAPAPAAGGSKVGDLFRASQLPADTLHQITELCGAKRVGYFGPAQFYVALKLIAAAQSGLPVRIESIKSELPLPRFMALKNDIEMRYGNAAEILGVKFQIPHPILEKNSFRRSDEGDKKETKSPPMSPMCSPPASPSNYQRIPLSYGYGKSRSKLEQPHGVSYEARHSTHQQDGPATGNYGSKSALVHSSLSRFLSVESEPQDSSNYPDDPWRITEEQREYYINQFKSLQPDLNSLISGSVAKNFFTKSKLPIPELSHVWELSDVDCDGALTLPEFCAAFHLIVARKNGYPLPDMLPETLLPDYLQPASLKPKRDCALLDSYSESLSVSQQTRDFNRTETSVEEMPDKSALTQDANSDDNQALNISTTTKTIPKDFKPSQHLSVKTVVQESNTLKARPRSRSYSSTSIEEAMKKGEEPPMPPPRPQKSHSRASSLDLNKIFQQNTQAVKSGWLPPPPALPPRPFVSQQGPHHVTNTESVAQAKSSYVEFRLKEQTEQVSEIELHPQMNRAPSQTEESSPMRKEILLTQPPSKPIRRKFRSENQMLENQELTPTSSVTSSLPAVKPHLPAQNRQSSKQKRAIQTAIRKNKEANAVLARLNSELQQQLKVKNALNITITRLDAIGCFSPLAEDLVPLYR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ78</accession>
    <name>A0A452IQ78_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Amine oxidase</fullName>
            <ecNumber evidence="16">1.4.3.-</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="14">
            <text>an L-alpha-amino acid + O2 + H2O = a 2-oxocarboxylate + H2O2 + NH4(+)</text>
            <dbReference type="Rhea" id="RHEA:13781"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15379"/>
            <dbReference type="ChEBI" id="CHEBI:16240"/>
            <dbReference type="ChEBI" id="CHEBI:28938"/>
            <dbReference type="ChEBI" id="CHEBI:35179"/>
            <dbReference type="ChEBI" id="CHEBI:59869"/>
            <dbReference type="EC" id="1.4.3.2"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 16">
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the flavin monoamine oxidase family. FIG1 subfamily.</text>
    </comment>
    <dbReference type="EC" id="1.4.3.-" evidence="16"/>
    <dbReference type="AlphaFoldDB" id="A0A452IQ78"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034186.1">
        <property type="protein sequence ID" value="ENSGAGP00000030111.1"/>
        <property type="gene ID" value="ENSGAGG00000021720.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001716">
        <property type="term" value="F:L-amino-acid oxidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090729">
        <property type="term" value="F:toxin activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009063">
        <property type="term" value="P:amino acid catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042742">
        <property type="term" value="P:defense response to bacterium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
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        <property type="entry name" value="Amine oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.660.10">
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="FAD/NAD(P)-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.405.10">
        <property type="entry name" value="Guanine Nucleotide Dissociation Inhibitor, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002937">
        <property type="entry name" value="Amino_oxidase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036188">
        <property type="entry name" value="FAD/NAD-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001613">
        <property type="entry name" value="Flavin_amine_oxidase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050281">
        <property type="entry name" value="Flavin_monoamine_oxidase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10742:SF355">
        <property type="entry name" value="AMINE OXIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10742">
        <property type="entry name" value="FLAVIN MONOAMINE OXIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01593">
        <property type="entry name" value="Amino_oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00757">
        <property type="entry name" value="AMINEOXDASEF"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54373">
        <property type="entry name" value="FAD-linked reductases, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51905">
        <property type="entry name" value="FAD/NAD(P)-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0044">Antibiotic</keyword>
    <keyword evidence="5" id="KW-0929">Antimicrobial</keyword>
    <keyword evidence="11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="8 16" id="KW-0274">FAD</keyword>
    <keyword evidence="6 16" id="KW-0285">Flavoprotein</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13" id="KW-1199">Hemostasis impairing toxin</keyword>
    <keyword evidence="9 16" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="7" id="KW-0800">Toxin</keyword>
    <feature type="domain" description="Amine oxidase" evidence="17">
        <location>
            <begin position="39"/>
            <end position="470"/>
        </location>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="40"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <begin position="59"/>
            <end position="60"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <begin position="81"/>
            <end position="84"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="84"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="249"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="447"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001974"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005465"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022529"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022630"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022656"/>
        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00022827"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
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        </source>
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            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023240"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00047637"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PIRSR" id="PIRSR601613-1"/>
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    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="RuleBase" id="RU362067"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF01593"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030111.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="490" mass="55938" checksum="E9AA7E94E6D82FB9" modified="2019-05-08" version="1">KEWEQCFDDPDYGELVDIAKNGLERKCNSKKVVIVGAGISGLTAAKLLRDAGCEVLILEASGRLGGRIETYRERDWYMDLGPMRLPKHHRIVLEFIKQFKLKLNTFYNTDANTWYLVSNVRARYKDVEQNPNILRYPVHPSEKGKSATQLYEQTLDKVTTDCNALKEKYDSFSTKEYLIKEGHLSRGAIDMIGDLLNEDGRFFMSFLYSVMAHVTFSNSSFYEITGGFDQLPNAFYQYVHGSIQFRSAVVKILTKGDQVIVSYRKANTSDLLSVTADYVLVTTTAKAARLIKFSPPLSPLKTHALRSIHYTKDVKIALACTKKFWEKDGIHGGRSITDRPSRFIYYPSHNFPSGLGVILASYTWGDDAEFFIPLSDEKCVDVVLEDLAKIHQLPKDYIQRVCNKYVIKKWGLDRYSMGSYVSFTQYQYVDYSKALFQNEGRIHFSGELTAQPHAWIDTAMKSGLRAARNIHAVINASPRQQQHELKENEL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ80</accession>
    <name>A0A452IQ80_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Lipocalin/cytosolic fatty-acid binding domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the calycin superfamily. Fatty-acid binding protein (FABP) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQ80"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034193.1">
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        <property type="gene ID" value="ENSGAGG00000021726.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
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        <property type="entry name" value="Fatty acid-binding protein, adipocyte"/>
        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
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        <property type="entry name" value="Calycin"/>
    </dbReference>
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        <property type="entry name" value="Fatty_acid-bd"/>
    </dbReference>
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        <property type="entry name" value="ILBP"/>
    </dbReference>
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        <property type="entry name" value="Lipocln_cytosolic_FA-bd_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11955">
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00061">
        <property type="entry name" value="Lipocalin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00178">
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    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Lipocalin/cytosolic fatty-acid binding" evidence="2">
        <location>
            <begin position="7"/>
            <end position="113"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00008390"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF00061"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030116.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="120" mass="13508" checksum="8E1BCF8C757F3EA3" modified="2019-05-08" version="1">MCDQFLGTWKLVSSEKFEDYMKELGVGFAMRKLGSLAKPTVIISADEDRLTIKTESPVKNTAISFKLGQEFEETTTDNRKTKSVVTLDNGSMIHVQKWDDKEITIKREVVDGKMVVVSGM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ83</accession>
    <name>A0A452IQ83_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Cytosolic fatty-acid binding proteins domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1 2">Belongs to the calycin superfamily. Fatty-acid binding protein (FABP) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQ83"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034137.1">
        <property type="protein sequence ID" value="ENSGAGP00000030065.1"/>
        <property type="gene ID" value="ENSGAGG00000021676.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008289">
        <property type="term" value="F:lipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd19443">
        <property type="entry name" value="FABP3-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.128.20:FF:000001">
        <property type="entry name" value="Fatty acid-binding protein, adipocyte"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.128.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012674">
        <property type="entry name" value="Calycin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000463">
        <property type="entry name" value="Fatty_acid-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031259">
        <property type="entry name" value="ILBP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000566">
        <property type="entry name" value="Lipocln_cytosolic_FA-bd_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11955">
        <property type="entry name" value="FATTY ACID BINDING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00061">
        <property type="entry name" value="Lipocalin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00178">
        <property type="entry name" value="FATTYACIDBP"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50814">
        <property type="entry name" value="Lipocalins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00214">
        <property type="entry name" value="FABP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0813">Transport</keyword>
    <feature type="domain" description="Cytosolic fatty-acid binding proteins" evidence="3">
        <location>
            <begin position="7"/>
            <end position="24"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00008390"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="RuleBase" id="RU003696"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS00214"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030065.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="126" mass="13877" checksum="CFD7834F2E9BE6A9" modified="2019-05-08" version="1">MCELFLGSWKLISSENFESYMKELGVGFATRKLGSLAKPSVVISTNDDIITIKTESTFKNTEISFKLGEEFEETTADDRKTKSIITLDNGALIHVQKWDGKETTIKRELVDSKGGGVTGLPPSLLC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ86</accession>
    <name>A0A452IQ86_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">THO complex subunit 1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="21">Nuclear matrix protein p84</fullName>
        </alternativeName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus matrix</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Nucleus</location>
            <location evidence="3">Nucleoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="19">Belongs to the THOC1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQ86"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034077.1">
        <property type="protein sequence ID" value="ENSGAGP00000030008.1"/>
        <property type="gene ID" value="ENSGAGG00000021230.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016363">
        <property type="term" value="C:nuclear matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005654">
        <property type="term" value="C:nucleoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000445">
        <property type="term" value="C:THO complex part of transcription export complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
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        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
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        <property type="term" value="P:apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006406">
        <property type="term" value="P:mRNA export from nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="P:mRNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
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        <property type="term" value="P:RNA splicing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
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        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
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        <property type="entry name" value="Death_NMPP84"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="THO complex subunit 1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Death Domain, Fas"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="DEATH-like_dom_sf"/>
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        <property type="entry name" value="Death_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021861">
        <property type="entry name" value="THO_THOC1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13265">
        <property type="entry name" value="THO COMPLEX SUBUNIT 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13265:SF2">
        <property type="entry name" value="THO COMPLEX SUBUNIT 1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Death"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11957">
        <property type="entry name" value="efThoc1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00005">
        <property type="entry name" value="DEATH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47986">
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50017">
        <property type="entry name" value="DEATH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="13" id="KW-0007">Acetylation</keyword>
    <keyword evidence="9" id="KW-0053">Apoptosis</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="15" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="6" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="8" id="KW-0507">mRNA processing</keyword>
    <keyword evidence="17" id="KW-0508">mRNA splicing</keyword>
    <keyword evidence="10" id="KW-0509">mRNA transport</keyword>
    <keyword evidence="18" id="KW-0539">Nucleus</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="12" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="16" id="KW-0804">Transcription</keyword>
    <keyword evidence="14" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="4" id="KW-0813">Transport</keyword>
    <keyword evidence="11" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="Death" evidence="23">
        <location>
            <begin position="564"/>
            <end position="640"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="22">
        <location>
            <begin position="1"/>
            <end position="10"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004109"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004642"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022664"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022703"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022816"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00023187"/>
        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00023242"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00061272"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
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            <dbReference type="ARBA" id="ARBA00069888"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
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            <dbReference type="ARBA" id="ARBA00080564"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="PROSITE" id="PS50017"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030008.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="644" mass="74186" checksum="95AF36F6866A5B87" modified="2019-05-08" version="1">AAERWRHKRGPQSAEPTRETMSPPLFSLPEARLRFTVSSAGAAPRHGLPRRLATKTCLFFSENEKKCTLDQSFRVVLEEEIINQASCENVLAIISLAINAVTDGICTASTPFVLLGDVLDCLPLDQCDKIFTFVEKNVTTWKSNTFYSAGKNYLLRMCNDLLRRLSKSQNTVFCGRIQLFLARLFPLSEKSGLNLQSQFNLENVTVFNTNEQESTLGQKHIEERDEGMEVEEGEMGDDEAPTSCSIPIDYNLYRKFWSLQDYFRNPVQCYEKISWKTFLKYSEEVLAVFKSYKLDDTQASRKKLEELKTGGEHVYFAKFLTSEKLMDLQLSDSNFRRHILLQYLILFQYLKGQVKFKSSNYVLTDEQSLWIEDTTKLVYQLLSENPPDGERFSKMVEHILNTEENWNTWKNEGCPSFVKESEELTRLWNLCPDNMEACKSETREYMPTLEEFFEEAIEQADPENMVENEYKAVNNSNYGWRALRLLARRSPHFFQPTNQQFKSLPEYLENMVIKLAKELPPPSEEIKTGEDEDEEDNDALLKENNESPDVQRDKPVTGGQIESFANKLGEQWKTLAPYLEMKDSDIRQIESDSEDVKMRAKQLLVAWQDQEGIHATPENLITAMNKAGLSDLAESLSNDTENSS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ87</accession>
    <name>A0A452IQ87_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Peptidase S1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the peptidase S1 family. Snake venom subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQ87"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034226.1">
        <property type="protein sequence ID" value="ENSGAGP00000030148.1"/>
        <property type="gene ID" value="ENSGAGG00000021723.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030141">
        <property type="term" value="C:secretory granule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00190">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Kallikrein related peptidase 11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.10.10">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009003">
        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043504">
        <property type="entry name" value="Peptidase_S1_PA_chymotrypsin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001314">
        <property type="entry name" value="Peptidase_S1A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001254">
        <property type="entry name" value="Trypsin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018114">
        <property type="entry name" value="TRYPSIN_HIS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033116">
        <property type="entry name" value="TRYPSIN_SER"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271:SF48">
        <property type="entry name" value="KALLIKREIN-14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271">
        <property type="entry name" value="KALLIKREIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00089">
        <property type="entry name" value="Trypsin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00722">
        <property type="entry name" value="CHYMOTRYPSIN"/>
    </dbReference>
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        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50494">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="TRYPSIN_DOM"/>
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    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00135">
        <property type="entry name" value="TRYPSIN_SER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3 6" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="2 6" id="KW-0645">Protease</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 6" id="KW-0720">Serine protease</keyword>
    <feature type="domain" description="Peptidase S1" evidence="7">
        <location>
            <begin position="22"/>
            <end position="256"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00009228"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="RuleBase" id="RU363034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030148.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="258" mass="27609" checksum="2AF7576ACC62779F" modified="2019-05-08" version="1">SAPISFVNLSACLPVSSQNNRIVGGAPCIPHSMPWQAALFMGSQLNCGGTLIDRNWVVTAAHCHCGCACCEGRGCVIVRVSISVRLGEHNIRHLDWTEQLRLSEKTILHPGYNSASKDNDIMLIKLLTPANLNKNVQPLELPTSCPSPKEECVVSGWGTTKSPQCTVTPMLGGGLSFKGEGASSIYPRYYTENMLCAGVLEGGIDSCQGDSGGPLLCNGKLQGIVSWGTQTCALPNKPGVYVAVCKYVDWIRETINNN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1V6</accession>
    <name>A0A452J1V6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Beta-2-glycoprotein 1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="11">Apolipoprotein H</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="12">Beta-2-glycoprotein I</fullName>
        </alternativeName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">Binds to various kinds of negatively charged substances such as heparin, phospholipids, and dextran sulfate. May prevent activation of the intrinsic blood coagulation cascade by binding to phospholipids on the surface of damaged cells.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="13">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1V6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038915.1">
        <property type="protein sequence ID" value="ENSGAGP00000034372.1"/>
        <property type="gene ID" value="ENSGAGG00000024413.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
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    <dbReference type="GO" id="GO:0005576">
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    </dbReference>
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        <property type="entry name" value="Complement Module, domain 1"/>
        <property type="match status" value="4"/>
    </dbReference>
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        <property type="entry name" value="Sushi/SCR/CCP_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015104">
        <property type="entry name" value="Sushi_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000436">
        <property type="entry name" value="Sushi_SCR_CCP_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46393:SF7">
        <property type="entry name" value="COMPLEMENT C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46393">
        <property type="entry name" value="SUSHI DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00084">
        <property type="entry name" value="Sushi"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09014">
        <property type="entry name" value="Sushi_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00032">
        <property type="entry name" value="CCP"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57535">
        <property type="entry name" value="Complement control module/SCR domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50923">
        <property type="entry name" value="SUSHI"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="9 13" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6" id="KW-0358">Heparin-binding</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="7 14" id="KW-0732">Signal</keyword>
    <keyword evidence="5 13" id="KW-0768">Sushi</keyword>
    <feature type="signal peptide" evidence="14">
        <location>
            <begin position="1"/>
            <end position="19"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019116946" description="Beta-2-glycoprotein 1" evidence="14">
        <location>
            <begin position="20"/>
            <end position="290"/>
        </location>
    </feature>
    <feature type="domain" description="Sushi" evidence="15">
        <location>
            <begin position="21"/>
            <end position="81"/>
        </location>
    </feature>
    <feature type="domain" description="Sushi" evidence="15">
        <location>
            <begin position="82"/>
            <end position="144"/>
        </location>
    </feature>
    <feature type="domain" description="Sushi" evidence="15">
        <location>
            <begin position="145"/>
            <end position="204"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="13">
        <location>
            <begin position="23"/>
            <end position="66"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="13">
        <location>
            <begin position="147"/>
            <end position="190"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00003651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00020104"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022659"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022674"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00029855"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00033414"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00302"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50923"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034372.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="290" mass="32582" checksum="8DDBD048FC069EA6" modified="2019-05-08" version="1">AMPTVLFLWIVTLAHCALAGHVCHKPPEVPFATVDVDKRVYEVGEEITYTCDPGYSHQSGSRKYTCPLSGKWPINRMRCIPVTCPPPSLPEFGVISYHKLTPGNESVFQDIIRFECLPSFALFGNETATCMANGNWSDIPECRYVKCPHPTGIENGFINFEVRRTYHYMDSVDFGCQPHYVLDGPRESRCEKTGSWSIKPTCKAPCKIPVKKATVLYNGRKIKVQKDLKEGIQHAETIFFFCKNEIKKCGYTIPTQCINGQLTVPSCFKEIGIFAAIWKTDPADLTPCDP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1V8</accession>
    <name>A0A452J1V8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Parathyroid hormone 2 receptor</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the G-protein coupled receptor 2 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1V8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038917.1">
        <property type="protein sequence ID" value="ENSGAGP00000034374.1"/>
        <property type="gene ID" value="ENSGAGG00000024421.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
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        <property type="term" value="F:G protein-coupled peptide receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017046">
        <property type="term" value="F:peptide hormone binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="P:adenylate cyclase-modulating G protein-coupled receptor signaling pathway"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="P:cell surface receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.1240.10">
        <property type="entry name" value="GPCR, family 2, extracellular hormone receptor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1070.10">
        <property type="entry name" value="Rhodopsin 7-helix transmembrane proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050332">
        <property type="entry name" value="GPCR_2"/>
    </dbReference>
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        <property type="entry name" value="GPCR_2-like_7TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036445">
        <property type="entry name" value="GPCR_2_extracell_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001879">
        <property type="entry name" value="GPCR_2_extracellular_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000832">
        <property type="entry name" value="GPCR_2_secretin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017983">
        <property type="entry name" value="GPCR_2_secretin-like_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45620:SF18">
        <property type="entry name" value="PARATHYROID HORMONE_PARATHYROID HORMONE-RELATED PEPTIDE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45620">
        <property type="entry name" value="PDF RECEPTOR-LIKE PROTEIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00002">
        <property type="entry name" value="7tm_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02793">
        <property type="entry name" value="HRM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00249">
        <property type="entry name" value="GPCRSECRETIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00008">
        <property type="entry name" value="HormR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF111418">
        <property type="entry name" value="Hormone receptor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00649">
        <property type="entry name" value="G_PROTEIN_RECEP_F2_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50227">
        <property type="entry name" value="G_PROTEIN_RECEP_F2_3"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="G_PROTEIN_RECEP_F2_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="6" id="KW-0297">G-protein coupled receptor</keyword>
    <keyword evidence="9" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7 11" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-0675">Receptor</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0807">Transducer</keyword>
    <keyword evidence="4 11" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 11" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="150"/>
            <end position="173"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="185"/>
            <end position="206"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="240"/>
            <end position="258"/>
        </location>
    </feature>
    <feature type="domain" description="G-protein coupled receptors family 2 profile 1" evidence="12">
        <location>
            <begin position="35"/>
            <end position="133"/>
        </location>
    </feature>
    <feature type="domain" description="G-protein coupled receptors family 2 profile 2" evidence="13">
        <location>
            <begin position="148"/>
            <end position="281"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023224"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50227"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50261"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034374.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="16">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="281" mass="32087" checksum="C3D2582F00056A35" modified="2019-05-08" version="1">MTAAPVSLFLARQVDTDDVLTKEEQIYLLGEAKEKCQTNLKAQLEKIKGTISVLLSRVFAYWLDASCFPEWDGIICWPEGFPSQEVSVPCPEYIYDFNHKGHAYRHCDAYGNWELALSINKTWANYTECTDFLTPESRSREKEVFDRLHMMYTIGYSISLASLIVAVCILCYFKRLHCTRNYIHIHLFASFICRAASIFVKDVVLYSGSLIMALISQELGDVGCKVVVTLFLYFLATNHYWILVEGLYLHSLIFMAFLSNKNYLWALTIIGWDAGTSALGI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1W1</accession>
    <name>A0A452J1W1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Beta-2-glycoprotein 1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="11">Apolipoprotein H</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="12">Beta-2-glycoprotein I</fullName>
        </alternativeName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">Binds to various kinds of negatively charged substances such as heparin, phospholipids, and dextran sulfate. May prevent activation of the intrinsic blood coagulation cascade by binding to phospholipids on the surface of damaged cells.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="13">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
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        <property type="protein sequence ID" value="ENSGAGP00000034375.1"/>
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        <property type="project" value="UniProtKB-SubCell"/>
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        <property type="entry name" value="Complement Module, domain 1"/>
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    </dbReference>
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        <property type="entry name" value="Compl-Cell_Adhes-Reg"/>
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        <property type="entry name" value="Sushi_SCR_CCP_dom"/>
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        <property type="entry name" value="BETA-2-GLYCOPROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19325">
        <property type="entry name" value="COMPLEMENT COMPONENT-RELATED SUSHI DOMAIN-CONTAINING"/>
        <property type="match status" value="1"/>
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        <property type="match status" value="4"/>
    </dbReference>
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    </dbReference>
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    </dbReference>
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    <proteinExistence type="predicted"/>
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    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6" id="KW-0358">Heparin-binding</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
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    <keyword evidence="5 13" id="KW-0768">Sushi</keyword>
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        <location>
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        </location>
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        </location>
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    </feature>
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        <location>
            <begin position="171"/>
            <end position="214"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00003651"/>
        </source>
    </evidence>
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        <source>
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    </evidence>
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        </source>
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            <dbReference type="PROSITE" id="PS50923"/>
        </source>
    </evidence>
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        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="314" mass="35306" checksum="A143E66879BF0B0D" modified="2019-05-08" version="1">PLSPILSLSRDLVVSFLPVCHKPPEVPFATVDVDKRVYEVGEEITYTCDPGYSHQSGSRKYTCPLSGKWPINRMRCIRKYILHGTDTSQCMADGQWSGKLPECQPVTCPPPSLPEFGVISYHKLTPGNESVFQDIIRFECLPSFALFGNETATCMANGNWSDIPECRYVKCPHPTGIENGFINFEVRRTYHYMDSVDFGCQPHYVLDGPRESRCEKTGSWSIKPTCKAPCKIPVKKATVLYNGRKIKVQKDLKEGIQHAETIFFFCKNEIKKCGYTIPTQCINGQLTVPSCFKEIGIFAAIWKTDPADLTPCDP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1W4</accession>
    <name>A0A452J1W4_9SAUR</name>
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        <recommendedName>
            <fullName evidence="4">USP domain-containing protein</fullName>
        </recommendedName>
    </protein>
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        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
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            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
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        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
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        <property type="entry name" value="UBIQUITIN SPECIFIC PEPTIDASE 12-LIKE"/>
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        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034284.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="345" mass="39955" checksum="2C1C511004493F2B" modified="2019-05-08" version="1">MELLMTVRRLASICTMGANASALEKEIGPEQFPVNEHYFGLVNFGNTCYCNSVLQALYFCRPFREKVLAYKVQPRKKESLLTCLSDLFNSIATQKKKVGVIPPKKFISRLRKENELFDNYMQQDAHEFLNYLLNTIADLLQEEKKQERQNGKLQNGSIESEEGDKTDLTWVHEIFQGTLTNETRCLNCEAVSSKDEDFLDLSVDVEQNTSITHCLRGFSNTETLCSEYKYYCEQCRSKQEAQKRMRVKKLPMILALHLKRFKYMDQLHRYTKLSYRVVFPLELRLFNTSGDATNPDRMYDLVAVVVHCGSGPNRGHYITIVKSHGFWLLFDDDIVEVGIQIAFML</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1X1</accession>
    <name>A0A452J1X1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">DUF4485 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
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        <lineage>
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            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J1X1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038928.1">
        <property type="protein sequence ID" value="ENSGAGP00000034385.1"/>
        <property type="gene ID" value="ENSGAGG00000024427.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055310">
        <property type="entry name" value="CEP112"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027831">
        <property type="entry name" value="DUF4485"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18871">
        <property type="entry name" value="CENTROSOMAL PROTEIN OF 112 KDA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18871:SF2">
        <property type="entry name" value="CENTROSOMAL PROTEIN OF 112 KDA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14846">
        <property type="entry name" value="DUF4485"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="DUF4485" evidence="3">
        <location>
            <begin position="11"/>
            <end position="96"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="165"/>
            <end position="194"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="284"/>
            <end position="329"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="353"/>
            <end position="530"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="554"/>
            <end position="815"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="840"/>
            <end position="867"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="174"/>
            <end position="194"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="Pfam" id="PF14846"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034385.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="875" mass="103391" checksum="E3779D4765F96A4A" modified="2019-05-08" version="1">MSSEEETWEKLDAEFDHYVVDMKPFVLKLPHKSERQRCALWIKKLCEPSGAGAGIMGRKNRNLYAKLLLHMLKRGVLEGPFTHKPEPGILKTLPSYMSIYFDEPNSTRAQSGSPEALPDWVMGELGTSESKLEESWKFSSREESALVASPLVHRERYKYNGKLALRSHSMSPPHRSDEDNRAMQTPEDQHKKPISLDDSDLEARLNSWNLGLENPRYLREKPIPLSLMTPKMGLGKISTFRDEQSLFRIHEKELEMKTKIMEAKFHEEKLKLQQKHDADVQKILDRKNNETEVLKTLYKSKQNEAEETIRKLEKKVQTLVRESQVVRETKEKQIVELKKLCEQSTDSLNNDWEKRLHNAVAEMEKEKFDLQKKHTENIQELLEDTNARLSKMETEYLAQTKATNQMVKELEARVQQLTVEAESSNLQRQKLSQEKTDLEQCYQAVCTELQEMKSRCNCLQKEKDRIIQDYEQNIQQQQNKYDIDINFMKQEHALSAAKASDVIEELEQSVSELKQQLQESEHQRQQQLRDQEHKFQQEKLHLERIHEKKIHGFQNDLDKERGDAQKKIRKLEEALKEKEEQLTRVTEVQRLQAQQADAALEEFKLQVEQNSEKVYAEMKQQMEKVESDLSRSKSLREKQSKEFSWQLEELKQRYEQQIVELKLEHEQEKTHLFQQHNTEKDCLVRDHEREIEKLEKQLRAAMTEHENKTQECRKRDAQANSRLKHIEKEYSQKLAKSSQMVAELKTTISSLTEESSRQQLAAERRLQDIVQKFEDEKQQMIRDNDRAIKALQDEMENYCNQVRSTEKKLQHKELETQEQVTYIRQEYETKLKGLMPVSLRKELEDTIASLKSQVNFLQKRASVLQEELNVYHTRR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1Z6</accession>
    <name>A0A452J1Z6_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="13">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1Z6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038855.1">
        <property type="protein sequence ID" value="ENSGAGP00000034314.1"/>
        <property type="gene ID" value="ENSGAGG00000024379.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005604">
        <property type="term" value="C:basement membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009887">
        <property type="term" value="P:animal organ morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016358">
        <property type="term" value="P:dendrite development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008045">
        <property type="term" value="P:motor neuron axon guidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009888">
        <property type="term" value="P:tissue development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00055">
        <property type="entry name" value="EGF_Lam"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="CDD" id="cd03579">
        <property type="entry name" value="NTR_netrin-1_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000081">
        <property type="entry name" value="Netrin 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.50.120:FF:000001">
        <property type="entry name" value="Netrin 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000048">
        <property type="entry name" value="Netrin 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.50.120">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.260">
        <property type="entry name" value="Galactose-binding domain-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050440">
        <property type="entry name" value="Laminin/Netrin_ECM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008211">
        <property type="entry name" value="Laminin_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002049">
        <property type="entry name" value="LE_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056863">
        <property type="entry name" value="LMN_ATRN_NET-like_EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001134">
        <property type="entry name" value="Netrin_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018933">
        <property type="entry name" value="Netrin_module_non-TIMP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008993">
        <property type="entry name" value="TIMP-like_OB-fold"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10574:SF443">
        <property type="entry name" value="NETRIN 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10574">
        <property type="entry name" value="NETRIN/LAMININ-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00053">
        <property type="entry name" value="EGF_laminin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24973">
        <property type="entry name" value="EGF_LMN_ATRN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00055">
        <property type="entry name" value="Laminin_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01759">
        <property type="entry name" value="NTR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00643">
        <property type="entry name" value="C345C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00180">
        <property type="entry name" value="EGF_Lam"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00136">
        <property type="entry name" value="LamNT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50242">
        <property type="entry name" value="TIMP-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01248">
        <property type="entry name" value="EGF_LAM_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50027">
        <property type="entry name" value="EGF_LAM_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51117">
        <property type="entry name" value="LAMININ_NTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50189">
        <property type="entry name" value="NTR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5 8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7 8" id="KW-0424">Laminin EGF-like domain</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="3 9" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="9">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019153074" evidence="9">
        <location>
            <begin position="23"/>
            <end position="474"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin N-terminal" evidence="12">
        <location>
            <begin position="1"/>
            <end position="229"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="10">
        <location>
            <begin position="271"/>
            <end position="320"/>
        </location>
    </feature>
    <feature type="domain" description="NTR" evidence="11">
        <location>
            <begin position="342"/>
            <end position="471"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="8">
        <location>
            <begin position="271"/>
            <end position="283"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="8">
        <location>
            <begin position="273"/>
            <end position="290"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="8">
        <location>
            <begin position="292"/>
            <end position="301"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="8">
        <location>
            <begin position="304"/>
            <end position="318"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023292"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00460"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50027"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50189"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS51117"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034314.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="474" mass="52781" checksum="F84E59D946521F12" modified="2019-05-08" version="1">MPGLWQGLGVLLSLQLALLGLASDPCYDAARRPRYCLPEPTNLAAGRLVQASATCGRPPQRLCAFRHPGNTSLTLALGGRFELLYVALRFCSPRPHSLALYKSTDHGHSWMPFQFFSARCPQAYGLPLASVISKDWVTATDLRVLQVGGRCRCNGHAARCGAAGPSGAPQCQCRHNTAGPECETCRAFYCDRPWQRATPSDAHECVACECNRHSHRCRFNMELYKLSGRKSGGVCLNCRHHTAGRHCHYCQPGFKRDLARPITSRRACKACQCHPIGAVSTMCNQTTGQCQCKMGVTGLTCNRCAQGFQQSRTAHIPCIPGDELTPLYPCAPIYALPLGTECQTHCSPSHGRVHMNLRKYCKKDYVLHAQLLAMVESGEWWQFTTSVLVVYRQRQVPIRRGEQPLWVPRQDLACGCLRLQVGKAYLVIGNDAESPDPARLVLDRNSLALPWRDVWAHKLRRFQQQDRQGNCRSP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1Z9</accession>
    <name>A0A452J1Z9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Ig-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Lipid-anchor</topology>
            <topology evidence="1">GPI-anchor</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="12">Belongs to the immunoglobulin superfamily. IgLON family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1Z9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038962.1">
        <property type="protein sequence ID" value="ENSGAGP00000034413.1"/>
        <property type="gene ID" value="ENSGAGG00000024455.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098552">
        <property type="term" value="C:side of membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000013">
        <property type="entry name" value="cell adhesion molecule 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000500">
        <property type="entry name" value="limbic system-associated membrane protein isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000113">
        <property type="entry name" value="Opioid-binding protein/cell adhesion molecule"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013098">
        <property type="entry name" value="Ig_I-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050876">
        <property type="entry name" value="IgLON_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR42757">
        <property type="entry name" value="IGLON FAMILY OF IMMUNOGLOBULIN SUPERFAMILY-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR42757:SF22">
        <property type="entry name" value="LIMBIC SYSTEM-ASSOCIATED MEMBRANE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07679">
        <property type="entry name" value="I-set"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13927">
        <property type="entry name" value="Ig_3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="9" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="3" id="KW-0336">GPI-anchor</keyword>
    <keyword evidence="11" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="10" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="7" id="KW-0472">Membrane</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="4 13" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="13">
        <location>
            <begin position="1"/>
            <end position="28"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019048610" description="Ig-like domain-containing protein" evidence="13">
        <location>
            <begin position="29"/>
            <end position="309"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="14">
        <location>
            <begin position="24"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="14">
        <location>
            <begin position="132"/>
            <end position="214"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="14">
        <location>
            <begin position="219"/>
            <end position="306"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004609"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022622"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00037995"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034413.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="309" mass="34527" checksum="014E2A01FE343B7C" modified="2019-05-08" version="1">MGGRAQPDRKQLPLVLLRLLCLLPTGLPVRSLDFTRGTDNITVRQGDTAILRCYVEDRSSKVAWLNRSGIIFAGQDKWSLDPRVELEKRTPLEYSLRIQKVDVYDEGSYTCSVQTQHHPRTSQVYLIVQVPPKISNISSDVTVNEGSNVTLVCMANGRPEPVITWRHLTPTGREFEGEEEYLEILGITREQSGKYECKAANEVSSADVKQVRVTVNYPPTITESKSNEAATGRQALLKCEASAVPTPDFEWYRDDTRINSANGLEIKSMGSQSLLMVANVTEEHYGNYTCVAANKLGVTNASLYLYSKY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J275</accession>
    <name>A0A452J275_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="12">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Golgi apparatus membrane</location>
            <topology evidence="1">Peripheral membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the DOP1 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000039066.1">
        <property type="protein sequence ID" value="ENSGAGP00000034503.1"/>
        <property type="gene ID" value="ENSGAGG00000024506.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="GOC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005768">
        <property type="term" value="C:endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000139">
        <property type="term" value="C:Golgi membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005802">
        <property type="term" value="C:trans-Golgi network"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006895">
        <property type="term" value="P:Golgi to endosome transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040314">
        <property type="entry name" value="DOP1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056457">
        <property type="entry name" value="DOP1_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007249">
        <property type="entry name" value="DOP1_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056459">
        <property type="entry name" value="TPR_DOP1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056458">
        <property type="entry name" value="TPR_DOP1_M"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14042">
        <property type="entry name" value="DOPEY-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14042:SF22">
        <property type="entry name" value="PROTEIN DOPEY-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24598">
        <property type="entry name" value="DOP1_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04118">
        <property type="entry name" value="Dopey_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24601">
        <property type="entry name" value="TPR_DOP1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24597">
        <property type="entry name" value="TPR_DOP1_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="5" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0653">Protein transport</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0813">Transport</keyword>
    <feature type="domain" description="DOP1 N-terminal" evidence="8">
        <location>
            <begin position="11"/>
            <end position="294"/>
        </location>
    </feature>
    <feature type="domain" description="DOP1-like middle TPR" evidence="9">
        <location>
            <begin position="318"/>
            <end position="471"/>
        </location>
    </feature>
    <feature type="domain" description="DOP1-like TPR" evidence="11">
        <location>
            <begin position="1361"/>
            <end position="1735"/>
        </location>
    </feature>
    <feature type="domain" description="DOP1-like C-terminal" evidence="10">
        <location>
            <begin position="1902"/>
            <end position="2402"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="578"/>
            <end position="599"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="622"/>
            <end position="648"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="715"/>
            <end position="735"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1173"/>
            <end position="1217"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1286"/>
            <end position="1311"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="633"/>
            <end position="648"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="720"/>
            <end position="731"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="1173"/>
            <end position="1202"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="1203"/>
            <end position="1217"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004395"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00046326"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF04118"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF24597"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="Pfam" id="PF24598"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF24601"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034503.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2473" mass="278543" checksum="5B7669E3F1AFB115" modified="2019-05-08" version="1">MNTEEVELLSDSKYRNYVAAVDKALKNFEYSSEWADLISALGKLNKVLQNNAKYQVVPKKLTIGKRLAQCLHPALPGGVHRKALETYEIIFKIIGPKRLAKDLFLYSSGLFPLLANAAMSVKPALLSLYEIYYLPLGKTLKPGLQGLLTGILPGLEEGSEYYERTNTLLEKVAGVDQSAFYSALWGSLLTSPAVRLPGITYVLSHLNRKLSMEDQLYIIGSDIELMVEAVSTSVQDSSVLVQRSTLDLILFCFPFHMSQATRPDMIRILSAALHVVLRRDMSLNRRLYAWLLGFDNNGAIVGPRSTRQNNPEEHATYYFNTFSKEMLVQAMVGILQVNGHGEESTMMQDLKPFRILISLLDKPELGPVILEDVLIEVFRTLYTQCKAELDLQMEPSFSKDHTQLSSKLRENKKTAELIKTANLLFNSFEPYYMWDYIARWFEECCRRTLHARLQTGPVGGNESSELPLTNFCLLVDFLLDIVSLPTRSMRVLCQETYIEIQTEHLPQLLLRMISALTSHLHTLHLSELTDSLRLCSKILSKVQPPLLSAGTDGTLQLPSGHNSSIKEWEDKKVPSFSIENPHDVFEDGENPPSSRSSESGFTEFVQYQADTADDIDRALSEGHGAPGIPIIGSTSSETETASTVGSEETIVHPPSIITQGSATQSGKMIQKTAMQCCLEYVQQFLTRFINLYIIQSNSLSQPLATELQVDLTREQGQPAKWDRESQGDAKGKKTNKTPKEYLSAFIAACQLYLECSSFPVYIAEGNHTSELHPEKSDFDCEQVQPPLWLQTLMNACRQASDFSIQGIAISLIMDLVGLTQSVAVVTGENLNSVETAQPLSPNQGRVAVVIRPPLTQGNLRYMVEKTDFFKHVALTLWDQLGDGTPQHHQKSVELFYQLHNLVPSSSICEDVISQQLTHRDKKTRMETHAKFAVLWHLTRDLHINKSSSFGRSFDRSLFIMLDSLNSLDGSTWSVGQAWLNQVLQRHDIARVLEPLLLLLLHPKTQRVSVQRIQAECYWNKSPHYPEEETEKHFMQKFSCGDAFSHMPVSQGQLITPKENNEKQLIMDEMENFSLTVNPLSDRLSLLSTSSETIPMVVSDFELPDHQIEILQSSDSGCSQSSAGDNLSYEVEAESLSAQNSSQTLREDSPDEIVQQVVIDLICKVVSGLGEETESVKHDLHPDDASSKFSTLDHSEEVSKSEDQNIQSSENSLFGNDSSQLLSASTEIGLERLTDSPPCIEVSSQAPCDLTFSSTDGKSGQRSHSSIQFSFKGKLPEKVSEKETIVKEAGKQPGAKPKVKIARRKDEDKKKAQTEKLKQTSIFFSDGLDLENWYSCGEGEISEIESDVGSPGIRKSPSFNVHPLYQHVLLYLQLYDSSRTLYAFSAIKAILKTNPSAFVNAISTTSVNNVYTPQLSLLQNLLARHRISVMGKDFYSHIPVDSNHNFRSSMYIEIIISLCLYYMRSHYPTHVKVTPQDLIGNRNMQMMSIEILTLLFSELAKVIESSAKGFPSFISDMLSKCKVQKVILHCLLSSIFSAQKWHSEKVAGKNIVVIEEGFSEDSLINFSEDEFDSGSTLQSQLLKVLQRLIVLEHRVMTVPEENETGFEFAITDLEQINPQQPMTSLQYLHSQPITSQGMFLCAVIRALHQHCACKMHPQWIGLITSTLPYMGKVLQRVVVSVTLQLCRNLDNLIQQYKYETGLSNSRPLWMASITPPDMILTLLEGITTIIHYCLLDPSTQYHQLLVNVDQKHLLEARSGILSILHMIMSSVTLLWSILHQADSSEKTTAAAAASVTTINLGSTKNLRQQILELLGPISMNHGVHFMAAVAFVWNERRQNKNTSRTKVIPTASDEQLLLVELVRSISVMRTETVIQTVKEVLKQPPAIAKDKKHLSLEVCMLQFFYAYIQRIPVSSLVDSWATLLLLLKDSIQLSLPAPGQFLILGVLNEFIMKNPSLENKKDQRDLQDVTHKIVDAIGAIAGSSLEQTTWLRRNLEVKPSLKIMVDGNNLESDVEDMLYPALETSNITPSVYSVHALTLLSEVLAHLLDMVFYSDEKERVIPLLVNIMHYVVPYLRNHSAHNAPSYRACVQLLSSLSGYQYTRKAWKKEAFDLFMDSSFFQMDASCVNHWRAIMDNLMTHDKTTFRDLMTRVAVAQSSSLNLFANRDVELEQRAMLLKRLAFAIFSSEVDQYQKYLPDIQERLVESLRLPQVPTLHSQVFLFFRVLLLRMSPQHLTSLWPTMITELVQVFLLMEQELTADEDISRTSGPSVAGLETTYTGGNGFSTSYNSQRWLNLYLSACKFLDLALALPSENLPQFQMYRWAFIPEVSDDSGLEVRRQGTHQREFKPYVVRLAKLLRKRAKKNPEEDCSGKTLSWEPGHLLLTIYTVRSIEQLLPFFNVLSQVFNSKVTSRCVGHSGSPVLYPNSFPSKDIKMENQKTFSSKARQKIEEMVEKDFLEGVIKTHLQPRGTRKA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J279</accession>
    <name>A0A452J279_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">HMG box domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the TCF/LEF family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J279"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039069.1">
        <property type="protein sequence ID" value="ENSGAGP00000034506.1"/>
        <property type="gene ID" value="ENSGAGG00000024527.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:1990907">
        <property type="term" value="C:beta-catenin-TCF complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000785">
        <property type="term" value="C:chromatin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060070">
        <property type="term" value="P:canonical Wnt signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd21996">
        <property type="entry name" value="HMG-box_TCF7-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.30.10:FF:000001">
        <property type="entry name" value="transcription factor 7 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.900.10:FF:000016">
        <property type="entry name" value="Uncharacterized protein"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="High mobility group box domain"/>
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    </dbReference>
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    </dbReference>
    <dbReference type="Pfam" id="PF08347">
        <property type="entry name" value="CTNNB1_binding"/>
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    </dbReference>
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    </dbReference>
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    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="HMG_BOX_2"/>
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    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0010">Activator</keyword>
    <keyword evidence="6 10" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="9 10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0678">Repressor</keyword>
    <keyword evidence="8" id="KW-0804">Transcription</keyword>
    <keyword evidence="5" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="4" id="KW-0879">Wnt signaling pathway</keyword>
    <feature type="domain" description="HMG box" evidence="12">
        <location>
            <begin position="302"/>
            <end position="370"/>
        </location>
    </feature>
    <feature type="DNA-binding region" description="HMG box" evidence="10">
        <location>
            <begin position="302"/>
            <end position="370"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1"/>
            <end position="81"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="180"/>
            <end position="218"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="274"/>
            <end position="300"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="365"/>
            <end position="400"/>
        </location>
    </feature>
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        <location>
            <begin position="412"/>
            <end position="458"/>
        </location>
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            <begin position="28"/>
            <end position="56"/>
        </location>
    </feature>
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            <begin position="274"/>
            <end position="289"/>
        </location>
    </feature>
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        <location>
            <begin position="429"/>
            <end position="446"/>
        </location>
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        <location>
            <begin position="447"/>
            <end position="458"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00006569"/>
        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
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    </evidence>
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        <source>
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        </source>
    </evidence>
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        <source>
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        </source>
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        <source>
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        </source>
    </evidence>
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        <source>
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        <source>
            <dbReference type="PROSITE" id="PS50118"/>
        </source>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034506.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="540" mass="58286" checksum="BF2AA70097BD7043" modified="2019-05-08" version="1">MPQLPPAGGDDLGAPDELIPFQDEGDEQDKGAGRGSAHGDLDELKSSLVHESENRRASASGSDSEAERPPPPQPRESFQKPRDYLAEVVRRQQDGGFFKGPPYPGYPFLMIPELGSTYLPNGALSPGGARTYLQMKWPLLDVPAGATLKDSRSPSPAHLSNKVPVVQHAHHMHPLTPLITYSSDHFSPGSPPSHLSPEIDPKTGIPRPPHPSELSPYYPLSPGAVGQIPHALGWLVPHLMSSRFSPHMVSPAAHGLHPSGIPHPTIVSPIVKQEPTQASMSPGGNSKSPGSVKKEEEKKPHIKKPLNAFMLYMKEMRAKVVAECTLKESAAINQILGRRWHSLSREEQAKYYELARKERQLHSQLYPTWSARDNYGKKKKRKREKQPQQQIQDTESSLASKSKKPYVQYLPIEKPCDSPASSHGSMLDSPATPSTALASPAAPAATHSEQAQPLSLTTKPEARAQLSLHSAAFLSSKAASSSSSLSSPSPLLSRPIPFATMTHTALLTSPPSFPAAIPSPQAALAVLQAQPLSLVTKPAD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J282</accession>
    <name>A0A452J282_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Caspase family p20 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
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            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the peptidase C14A family.</text>
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    </dbReference>
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        <property type="project" value="InterPro"/>
    </dbReference>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="P:positive regulation of neuron apoptotic process"/>
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    </dbReference>
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    </dbReference>
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        <property type="entry name" value="Caspase-like_dom_sf"/>
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    </dbReference>
    <dbReference type="InterPro" id="IPR015917">
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    </dbReference>
    <dbReference type="PANTHER" id="PTHR10454:SF199">
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    </dbReference>
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    </dbReference>
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    </dbReference>
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    </dbReference>
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    </dbReference>
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    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
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            <begin position="6"/>
            <end position="124"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00010134"/>
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    </evidence>
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        <source>
            <dbReference type="PROSITE" id="PS50208"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034514.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="140" mass="16296" checksum="A04D095B8936E01E" modified="2019-05-08" version="1">MSMQQRKNRAVIIVNYAFCSRTEPLQPRKGAKREADKLFKALSKLNYAVKLYYDQAAKDIEEIYRQESREEHGDCFVSILSSHGEEGAIYDCWGELVKLTRIFQILSPQRCPVLAGKPKIFFIQNLCRYLERLCIFLLAI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J283</accession>
    <name>A0A452J283_9SAUR</name>
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        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
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            <taxon>Archelosauria</taxon>
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            <taxon>Durocryptodira</taxon>
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            <taxon>Gopherus</taxon>
        </lineage>
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    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J283"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039096.1">
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        <property type="gene ID" value="ENSGAGG00000024542.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="38"/>
            <end position="65"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="38"/>
            <end position="50"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034527.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="112" mass="12051" checksum="3A97E44DC87D975D" modified="2019-05-08" version="1">MSAQVKDKAAFQRLSFLYQAAHCVLAQILRTRSWHGSTVTCSAPSVTEPSANPAREDAATGTGSYQKPVLQGSVLAAAIPDFLPTSPQKTFDGKEPPDSLIAREIPFSFQKD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J287</accession>
    <name>A0A452J287_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Ig-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J287"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039074.1">
        <property type="protein sequence ID" value="ENSGAGP00000034511.1"/>
        <property type="gene ID" value="ENSGAGG00000024508.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002764">
        <property type="term" value="P:immune response-regulating signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000033">
        <property type="entry name" value="Killer cell immunoglobulin-like receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000049">
        <property type="entry name" value="Leukocyte immunoglobulin-like receptor subfamily B member 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050412">
        <property type="entry name" value="Ig-like_Receptors_ImmuneReg"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11738">
        <property type="entry name" value="MHC CLASS I NK CELL RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11738:SF186">
        <property type="entry name" value="OSTEOCLAST-ASSOCIATED IMMUNOGLOBULIN-LIKE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13895">
        <property type="entry name" value="Ig_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="4" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="5" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <keyword evidence="1" id="KW-0732">Signal</keyword>
    <feature type="domain" description="Ig-like" evidence="7">
        <location>
            <begin position="28"/>
            <end position="96"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="328"/>
            <end position="356"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="345"/>
            <end position="356"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034511.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="356" mass="38569" checksum="F3ECDC6EE19B1E60" modified="2019-05-08" version="1">SEPTEQVLLSLGLASIPSPRAGHEFLKPTIWVSPSRVVALGRNVTIRCEGQDLGMEFVLCKAGHPNQQVQSVVPEGKMAEFPIASVGQEDGGSYTCYYRSITDQSHWSFPSNPVEIIVGGEGGAWLSGPAPSPTPRQTHRGSLQGVRFVLNKEGRHFPPVDSDGFGAVFPISNVSREDGGNYSCSYHSRSEPFAVSDPSDPMELPSYPKPNISGIPRRGVSVGEAMAIWCRRQRQGVRFVLNKEGRHFPPVDSDGFGAVFPISNVSREDGGNYSCSYHSRSEPFAVSDPSDPMELVVLQAKRGAKPPAPEPAQPSMYAAINVNWGARGKRPPQSWGTGPQGATDFKARRVHPDHLV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J289</accession>
    <name>A0A452J289_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Presenilin</fullName>
            <ecNumber evidence="20">3.4.23.-</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="20">Probable subunit of the gamma-secretase complex, an endoprotease complex that catalyzes the intramembrane cleavage of integral membrane proteins such as Notch receptors.</text>
    </comment>
    <comment type="subunit">
        <text evidence="20">Homodimer.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cell projection</location>
            <location evidence="3">Axon</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasmic granule</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="20">Endoplasmic reticulum membrane</location>
            <topology evidence="20">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="20">Golgi apparatus membrane</location>
            <topology evidence="20">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="19">Synapse</location>
        </subcellularLocation>
    </comment>
    <comment type="domain">
        <text evidence="20">The PAL motif is required for normal active site conformation.</text>
    </comment>
    <comment type="similarity">
        <text evidence="4 20">Belongs to the peptidase A22A family.</text>
    </comment>
    <dbReference type="EC" id="3.4.23.-" evidence="20"/>
    <dbReference type="AlphaFoldDB" id="A0A452J289"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039079.1">
        <property type="protein sequence ID" value="ENSGAGP00000034516.1"/>
        <property type="gene ID" value="ENSGAGG00000024531.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000039084.1">
        <property type="protein sequence ID" value="ENSGAGP00000034520.1"/>
        <property type="gene ID" value="ENSGAGG00000024531.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030424">
        <property type="term" value="C:axon"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070765">
        <property type="term" value="C:gamma-secretase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000139">
        <property type="term" value="C:Golgi membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045202">
        <property type="term" value="C:synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042500">
        <property type="term" value="F:aspartic endopeptidase activity, intramembrane cleaving"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034205">
        <property type="term" value="P:amyloid-beta formation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006915">
        <property type="term" value="P:apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055074">
        <property type="term" value="P:calcium ion homeostasis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035556">
        <property type="term" value="P:intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006509">
        <property type="term" value="P:membrane protein ectodomain proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007219">
        <property type="term" value="P:Notch signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016485">
        <property type="term" value="P:protein processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.472.100:FF:000001">
        <property type="entry name" value="Presenilin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.472.100">
        <property type="entry name" value="Presenilin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002031">
        <property type="entry name" value="Pept_A22A_PS1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001108">
        <property type="entry name" value="Peptidase_A22A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006639">
        <property type="entry name" value="Preselin/SPP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042524">
        <property type="entry name" value="Presenilin_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10202">
        <property type="entry name" value="PRESENILIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10202:SF18">
        <property type="entry name" value="PRESENILIN-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01080">
        <property type="entry name" value="Presenilin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01072">
        <property type="entry name" value="PRESENILIN"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01073">
        <property type="entry name" value="PRESENILIN1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00730">
        <property type="entry name" value="PSN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0053">Apoptosis</keyword>
    <keyword evidence="12" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="5" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="18" id="KW-0966">Cell projection</keyword>
    <keyword evidence="11 20" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="16 20" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="10 20" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="17 20" id="KW-0472">Membrane</keyword>
    <keyword evidence="13 20" id="KW-0914">Notch signaling pathway</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="7 20" id="KW-0645">Protease</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="15" id="KW-0770">Synapse</keyword>
    <keyword evidence="8 20" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="14 20" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="82"/>
            <end position="100"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="133"/>
            <end position="154"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="161"/>
            <end position="183"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="195"/>
            <end position="213"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="225"/>
            <end position="241"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="247"/>
            <end position="263"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="406"/>
            <end position="426"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="432"/>
            <end position="451"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="21">
        <location>
            <begin position="1"/>
            <end position="68"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="21">
        <location>
            <begin position="326"/>
            <end position="351"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="21">
        <location>
            <begin position="12"/>
            <end position="30"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="21">
        <location>
            <begin position="31"/>
            <end position="47"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="21">
        <location>
            <begin position="50"/>
            <end position="60"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004463"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004489"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00008604"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022703"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022976"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00034103"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="RuleBase" id="RU361148"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034516.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="465" mass="52637" checksum="5447F16E11B825C6" modified="2019-05-08" version="1">MTELSAHLPQFQHGQLTENFPDNHLCNTNDNSERRRHDNSERRRNDNPESETNGQPQNRTQQVVEQDEEDEELTLKYGAKHVIMLFAPVTLCMVVVVATIKSVSFYTRKDGQLIYTPFTEDTDTVGQRALNSILNAAIMISVIIVMTILLVVLYKYRCYKVIHGWLIISSLLLLFFFSFIYLGEVFKTYNVAMDYITLALMIWNFGVVGMICIHWKGPLRLQQAYLIMISALMALVFIKYLPEWTAWLILAVISVYDLVAVLCPKGPLRMLVETAQERNETLFPALIYSSTMMWLVNMAEEDPEAQRKASKSSTYDKQANQIQNEDADLDDGGFNPEWEQQRDSRIGPLESTPESRAAVQALPSNSQASEDPEERGVKLGLGDFIFYSVLVGKASATASGDWNTTLACFVAILIGLCLTLLLLAIFKKALPALPISITFGLVFYFATDNLVQPFMDQLAFHQFYI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJL4</accession>
    <name>A0A452GJL4_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="9">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endosome</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJL4"/>
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        <property type="protein sequence ID" value="ENSGAGP00000001791.1"/>
        <property type="gene ID" value="ENSGAGG00000001421.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="term" value="C:endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="CDD" id="cd21255">
        <property type="entry name" value="CH_EHBP1L1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="EH domain-binding protein 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.418.10">
        <property type="entry name" value="Calponin-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022735">
        <property type="entry name" value="bMERB_dom"/>
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        <property type="entry name" value="CH_dom"/>
    </dbReference>
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        <property type="entry name" value="CH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050540">
        <property type="entry name" value="F-actin_Monoox_Mical"/>
    </dbReference>
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        <property type="entry name" value="NT-C2"/>
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    <dbReference type="PANTHER" id="PTHR23167">
        <property type="entry name" value="CALPONIN HOMOLOGY DOMAIN-CONTAINING PROTEIN DDB_G0272472-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23167:SF42">
        <property type="entry name" value="EH DOMAIN-BINDING PROTEIN 1-LIKE PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="bMERB_dom"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="CH"/>
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    </dbReference>
    <dbReference type="SMART" id="SM00033">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01203">
        <property type="entry name" value="DUF3585"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47576">
        <property type="entry name" value="Calponin-homology domain, CH-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51848">
        <property type="entry name" value="BMERB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51840">
        <property type="entry name" value="C2_NT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50021">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0967">Endosome</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="C2 NT-type" evidence="7">
        <location>
            <begin position="8"/>
            <end position="175"/>
        </location>
    </feature>
    <feature type="domain" description="Calponin-homology (CH)" evidence="6">
        <location>
            <begin position="291"/>
            <end position="396"/>
        </location>
    </feature>
    <feature type="domain" description="BMERB" evidence="8">
        <location>
            <begin position="735"/>
            <end position="877"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="161"/>
            <end position="289"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="439"/>
            <end position="468"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="510"/>
            <end position="744"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="5">
        <location>
            <begin position="169"/>
            <end position="178"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="179"/>
            <end position="188"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="230"/>
            <end position="243"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="5">
        <location>
            <begin position="269"/>
            <end position="286"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="550"/>
            <end position="559"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="678"/>
            <end position="693"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="705"/>
            <end position="723"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004177"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS51840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51848"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001791.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="877" mass="96222" checksum="F753BB8AF25C624D" modified="2019-05-08" version="1">MSSAWKRLQRVGKRASKFHFVACYQELMVECTKKWQPDKLIVVWTRRNRRVCSKAHSWQPGIKNPYRGMVVWMVPENVDIMVTLYRESKGQRKVLASADVNLQRFARPTPAQAELKLKLKPRSVKVVAATLQLTLSCVFLREGKATDEDMQSLASLMSVKPSDVGNLDDFADSDEDDEAARRVQHEDGSSQGTAPTDTSRELNTLAEEEENPSAPGTSRAAAGRATPHGRPREAGSAKEEPVRVADTPDAPLRDTPHPALELVLQGAAPSPPPELMGPPPEEPPATSPGLVSCSQSLLEWCQNVTAGYRGVRVTNFTTSWRNGLAFCAILHHFHPEKINYETLDPLAIKENNKLAYDGFASLGISRVLEPADMVFLTVPDKLIVMTYLCQIRAFFTGQELNVVQIANHSSQSTYKVGKFDSDPAFSIDPARFYAERFQSATRGPEEPGSTAPAARDDGKAAANKTVTGGTELAKKLEAELTAPKDAGTDAKIGDAEDAGKVTVNGTISQTTVQSEVEPSVTKDAGKVLINDVTTKPAISPEAEPPTPKDGGSKAAKDAGKSLVNGAMSPSAKKPEAEPLAPEDAGKVPANGDEKLVPPPRLKRQSVRNGEAPDQAERSLQRSLSTGSQGPVAPPRSHGSKSGFSHVRDADLLKKRRSRLKSESLSMDEGEAGGPPGETSRRRSDWLGTDDREGMMTPKAGPPASARTQQPPEIAPSSSTTTQEPSRDPSKETSTTEEEVPRFQDTSQYVVAELQALENEQKQIDARAAVVEKELRFLMESGTNQAQEEELIQEWFTLVNKKNALIRRQDQLQLLMEEQDLERRFELLSRELRAMLATEDCLKTEAQQRREQLLLEELVSLVNQRDELVRDLDIKERM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2025-10-08" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJL9</accession>
    <name>A0A452GJL9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Nuclear pore complex protein Nup98-Nup96</fullName>
        </recommendedName>
    </protein>
    <organism evidence="25 26">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="26" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="25" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="22">Plays a role in the nuclear pore complex (NPC) assembly and/or maintenance. NUP98 and NUP96 are involved in the bidirectional transport across the NPC. May anchor NUP153 and TPR to the NPC. In cooperation with DHX9, plays a role in transcription and alternative splicing activation of a subset of genes. Involved in the localization of DHX9 in discrete intranuclear foci (GLFG-body).</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Nucleus membrane</location>
            <topology evidence="2">Peripheral membrane protein</topology>
            <orientation evidence="2">Nucleoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
            <location evidence="1">Nuclear pore complex</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Nucleus</location>
            <location evidence="3">Nucleoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the nucleoporin GLFG family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000002028.1">
        <property type="protein sequence ID" value="ENSGAGP00000001783.1"/>
        <property type="gene ID" value="ENSGAGG00000001410.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000002046.1">
        <property type="protein sequence ID" value="ENSGAGP00000001796.1"/>
        <property type="gene ID" value="ENSGAGG00000001410.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031965">
        <property type="term" value="C:nuclear membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0044614">
        <property type="term" value="C:nuclear pore cytoplasmic filaments"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005654">
        <property type="term" value="C:nucleoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008139">
        <property type="term" value="F:nuclear localization sequence binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008236">
        <property type="term" value="F:serine-type peptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017056">
        <property type="term" value="F:structural constituent of nuclear pore"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051028">
        <property type="term" value="P:mRNA transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000973">
        <property type="term" value="P:post-transcriptional tethering of RNA polymerase II gene DNA at nuclear periphery"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006606">
        <property type="term" value="P:protein import into nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006405">
        <property type="term" value="P:RNA export from nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034398">
        <property type="term" value="P:telomere tethering at nuclear periphery"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.2360:FF:000001">
        <property type="entry name" value="Nuclear pore complex protein Nup98-Nup96"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.690:FF:000001">
        <property type="entry name" value="Nuclear pore complex protein Nup98-Nup96"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1610.10:FF:000001">
        <property type="entry name" value="Nuclear pore complex protein Nup98-Nup96"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.690">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1610.10">
        <property type="entry name" value="Peptidase S59, nucleoporin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037665">
        <property type="entry name" value="Nucleoporin_S59-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007230">
        <property type="entry name" value="Nup98_auto-Pept-S59_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036903">
        <property type="entry name" value="Nup98_auto-Pept-S59_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021967">
        <property type="entry name" value="Nup98_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23198:SF6">
        <property type="entry name" value="NUCLEAR PORE COMPLEX PROTEIN NUP98-NUP96"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23198">
        <property type="entry name" value="NUCLEOPORIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04096">
        <property type="entry name" value="Nucleoporin2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12110">
        <property type="entry name" value="Nup96"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21240">
        <property type="entry name" value="Nup98_GLEBS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF82215">
        <property type="entry name" value="C-terminal autoproteolytic domain of nucleoporin nup98"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51434">
        <property type="entry name" value="NUP_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="17" id="KW-0007">Acetylation</keyword>
    <keyword evidence="12" id="KW-0068">Autocatalytic cleavage</keyword>
    <keyword evidence="11" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="7" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="20" id="KW-0472">Membrane</keyword>
    <keyword evidence="13" id="KW-0509">mRNA transport</keyword>
    <keyword evidence="19" id="KW-0906">Nuclear pore complex</keyword>
    <keyword evidence="21" id="KW-0539">Nucleus</keyword>
    <keyword evidence="8" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="9" id="KW-0645">Protease</keyword>
    <keyword evidence="16" id="KW-0653">Protein transport</keyword>
    <keyword evidence="26" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0677">Repeat</keyword>
    <keyword evidence="14" id="KW-0720">Serine protease</keyword>
    <keyword evidence="18" id="KW-0811">Translocation</keyword>
    <keyword evidence="6" id="KW-0813">Transport</keyword>
    <keyword evidence="15" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="Peptidase S59" evidence="24">
        <location>
            <begin position="726"/>
            <end position="868"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="23">
        <location>
            <begin position="501"/>
            <end position="524"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="23">
        <location>
            <begin position="510"/>
            <end position="519"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004567"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004620"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004642"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00008926"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00013472"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022813"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022816"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023010"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023132"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00059309"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS51434"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001796.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1696" mass="183783" checksum="58BBBA575FB05281" modified="2019-05-08" version="1">MFNKSFGTPFGGSAGGFGTTSTFGQNAGFGTASGGAFGTSAFGSNNNTGSLFGNTQTKPGGLFGSSTFSQPATSSTSTGFGFGTSTGTSNSLFGTASTGGGLFSSQNNAFAQNKPAGFGNFGTSTSSGGLFGTTNTASNPFGGTSGSLFGPTSFTAAPTGTTIKFNPPTGTDTMVKSGVSTNISTKHQCITAMKEYESKSLEELRLEDYQANRKGPSNPAVAAAGLFGSSTATSSAATGLFGSSTTNAGFSYGQNKTAFGTSTTGFGTGTAGSLFGQQPQQTTSLFSKPFGQATTTQNTGFSFGNTSTLGQPNTNTMGLFGATQASPSGSLFGTAANTNAATGFGAGTGTGLFGQANTGFGVGGSTLFGKPAGFGTTTTSAPSFGTTTGSGLFGFGASTAGNSIFGNKPAAGTLGAGLGTGFGTALSAGQTSLFGNTQPKLGGTLGTGAFGAPGFNTSTATLGFGAPQPAVALTDPNVSAAQQAALQQHFNSLTYSPFGDSPLFRNPMSDPKKKEERLKPTNPAAQKALTTPTHYKLTPRPATRVRPKALQTAGSSKSHLFDGLDDDEPSLSNGAFMPKKSIKKLVLKNLNSSNLFSPVSREAEDLDPPSEYPENGERFNFLSRSVDENHRQEGEREEEDDHHEVTRFYTNPIAKPIPQTPENAVHKHQNNVDDTIVALNMRVALRNGLEGSSEDASFHDESLQDDREEENENVYSLHPAGIVLTKAGYYTIPSMEDLAKLTTDRGECIVTDFTIGRKGYGSIYFEGEVNLTNLNLDEIVHIRRKEVIVYPDDDQKPPIGEGLNRRAEVTLDGVWPTDKTSRCLIKSPERLADINYEGRLEAVSRKQGAQFKEYRPETGSWVFKVAHFSKYGLQDSDEEEEEHPSKMDAKKLKTAQVPPQGQLIPQQMALNGKPTSPSQSPEVEQLGRVVELDSDMADITQEPAQDSILEDSITEEPELVPASTHIASSLGINPHVLQIMKASLLADEEDVDLILDHHFGKQPTKMDTSQEICSPRLPISALQGQKRHSSVGGLLQSKFASVSFLQSASLPDFHGPRASSFGNPPSAASWTGLSPLPSAFSMPSAAPEVQLKTVGACRQQGLIPLEKSVTYGKGRLLMDMALFMGRSFRVGWGPNWTLVNSGDKLSGSSEPKGLRNDSMEYGFLPTPVAPKSLSESPFKVHIEKLTLEQRKTDGDQQLYLIPLEIKLKHSTVHVDERCPLLAANPGVSAIHDYADWVRDFCGDSVEIEPVVKYWCLTWTLCEAIWGHLKELEASLEEPSEYIVTLERRKAFSRWLSQTAAARIEEEVSLSQHENHIEAVFSYLTGKRISEACRVAQQSGDHRLALLLSQLVGSPQVRELLTMQLVDWHGLQADCFIQEERLRIFALLAGKPVWQLSERKSINVCSQLDWKRSIAIHLWYLLPPTASISKALNMYEAAFQNTPECEKYACCPLPPYLEDSGCVLEEDDSTERPLRDVCFHLLKLYSDRCYDLNQLLDPRSITSDPLDYRLSWHLWEVLRALNYTHLSEQCQGVLNASYAAQLESEGLWEWAIFVLLHTSNTYTREKAVREFLNRHCVLSETPESWTKETFLTQKLCIPPEWIQEAKAVRARMEGDKHKEALYLFKAGHWNQCHKLVIRHLASDAIINENYNYLKGFLEDLSPPERSTLIQDWDTAGLVYLDYIRVIEMLDRIQQVTV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJM0</accession>
    <name>A0A452GJM0_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="5">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the small GTPase superfamily. Ras family. KappaB-Ras subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJM0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002144.1">
        <property type="protein sequence ID" value="ENSGAGP00000001885.1"/>
        <property type="gene ID" value="ENSGAGG00000001505.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005525">
        <property type="term" value="F:GTP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032794">
        <property type="term" value="F:GTPase activating protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003924">
        <property type="term" value="F:GTPase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043124">
        <property type="term" value="P:negative regulation of canonical NF-kappaB signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032484">
        <property type="term" value="P:Ral protein signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042227">
        <property type="entry name" value="KBRS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005225">
        <property type="entry name" value="Small_GTP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001806">
        <property type="entry name" value="Small_GTPase"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00231">
        <property type="entry name" value="small_GTP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46152">
        <property type="entry name" value="NF-KAPPA-B INHIBITOR-INTERACTING RAS-LIKE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46152:SF1">
        <property type="entry name" value="NF-KAPPA-B INHIBITOR-INTERACTING RAS-LIKE PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00071">
        <property type="entry name" value="Ras"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00449">
        <property type="entry name" value="RASTRNSFRMNG"/>
    </dbReference>
    <dbReference type="SMART" id="SM00175">
        <property type="entry name" value="RAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00173">
        <property type="entry name" value="RAS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51419">
        <property type="entry name" value="RAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51421">
        <property type="entry name" value="RAS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0342">GTP-binding</keyword>
    <keyword evidence="2" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="172"/>
            <end position="191"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00008094"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001885.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="191" mass="21289" checksum="BFE28A2B5E6230A2" modified="2019-05-08" version="1">MGKGYKVVICGMASVGKTAILEQLLYGKHTVGLEEGATVEDVYLASVETDRGVKEQLRLYDTTGLQEGVEFPKHYFSVADGFILVYAVNSLESFQRAEVIKKEIDTVRDKKEVTVVVLGNKTDLTEQRQVETEIAQQWAKAEKVRLWEVTVTDRKTLIEPFTSLASKLSQSQNKSAFPLPGRKSKGNNCDN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJM1</accession>
    <name>A0A452GJM1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">C2H2-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="11">Involved in histone 3'-end pre-mRNA processing by associating with U7 snRNP and interacting with SLBP/pre-mRNA complex. Increases histone 3'-end pre-mRNA processing but has no effect on U7 snRNP levels, when overexpressed. Required for cell cycle progression from G1 to S phases.</text>
    </comment>
    <comment type="subunit">
        <text evidence="12">Interacts with the SLBP/pre-mRNA complex but not with SLBP alone. Interacts with LSM11 in a U7 snRNP-dependent manner.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000002145.1">
        <property type="protein sequence ID" value="ENSGAGP00000001886.1"/>
        <property type="gene ID" value="ENSGAGG00000001510.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0022603">
        <property type="term" value="P:regulation of anatomical structure morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001621">
        <property type="entry name" value="Uncharacterized protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:003000">
        <property type="entry name" value="Zinc finger and SCAN domain-containing 20"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000355">
        <property type="entry name" value="zinc finger and SCAN domain-containing protein 20 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000358">
        <property type="entry name" value="zinc finger protein 24"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002343">
        <property type="entry name" value="Zinc finger protein 33A"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000690">
        <property type="entry name" value="Zinc finger protein 354C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002090">
        <property type="entry name" value="Zinc finger protein 473"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000340">
        <property type="entry name" value="zinc finger protein 473 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001270">
        <property type="entry name" value="zinc finger protein 583 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="17"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23235:SF178">
        <property type="entry name" value="C2H2-TYPE DOMAIN-CONTAINING PROTEIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23235">
        <property type="entry name" value="KRUEPPEL-LIKE TRANSCRIPTION FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="17"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="17"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="16"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="17"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="9" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 13" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="24"/>
            <end position="64"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="65"/>
            <end position="92"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="93"/>
            <end position="120"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="121"/>
            <end position="148"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="159"/>
            <end position="186"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="187"/>
            <end position="214"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="215"/>
            <end position="242"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="243"/>
            <end position="270"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="271"/>
            <end position="298"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="299"/>
            <end position="326"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="327"/>
            <end position="354"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="355"/>
            <end position="382"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="383"/>
            <end position="410"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="411"/>
            <end position="438"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="439"/>
            <end position="466"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="467"/>
            <end position="494"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="495"/>
            <end position="522"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="519"/>
            <end position="543"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="519"/>
            <end position="536"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00054787"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00062613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001886.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="543" mass="61520" checksum="216DC57435DF3031" modified="2019-05-08" version="1">SVYSQPLLYIREPTQERNLMNRPYKCLDCGKSFSRRSDLERPYTCLDCGKILHIHQRTHTGEKPYKCLKCGKSFSQSSGLISHERIHTGERPYKCLECGKSFSHRSGLSNHGRIHTGERPYKCLECGKSFSRHSDLSNHGRIHTGEKPYKCXHSGERPYKCLDCGKSFNQSSNLTTHQRVHIGERPYKCLECGKSFSVLSTLIVHQRTHTGEKPYECLDCGKSFYRSSNLTTHQRVHTGEGPYKCLECGKSFSAPSVLLIHQRTHTGEKPYKCLECGKSFSRRSGLISHERIHTGKRPYKCLDCGKSFSRRSDLGNHGRIHTGERPYKCLECGKSFSQSSGLINHSRIHTGERPYTCLDCGKSFYGSSNLTAHQRVHIGEGPYKCLECGKSFSAPSVLHIHQRTHTGEKPYKCLKCGKSFSQSSGLISHERIHTGERPYKCLECGKSFSHRSGLSNHGRIHTGERPYKCLECGKSFSRHSDLSNHGRIHTGEKPYKCLICEKSFNRSSNLHTHWRVHTGESGKSFRESSKHQDRNHVSASAVG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJM6</accession>
    <name>A0A452GJM6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">C2H2-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000002150.1">
        <property type="protein sequence ID" value="ENSGAGP00000001891.1"/>
        <property type="gene ID" value="ENSGAGG00000001510.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001997">
        <property type="entry name" value="Uncharacterized protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:003000">
        <property type="entry name" value="Zinc finger and SCAN domain-containing 20"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000446">
        <property type="entry name" value="Zinc finger protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000706">
        <property type="entry name" value="Zinc finger protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000824">
        <property type="entry name" value="Zinc finger protein 184"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000358">
        <property type="entry name" value="zinc finger protein 24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002343">
        <property type="entry name" value="Zinc finger protein 33A"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000902">
        <property type="entry name" value="Zinc finger protein 445"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002090">
        <property type="entry name" value="Zinc finger protein 473"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001270">
        <property type="entry name" value="zinc finger protein 583 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000642">
        <property type="entry name" value="Zinc finger with KRAB and SCAN domains 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23235:SF178">
        <property type="entry name" value="C2H2-TYPE DOMAIN-CONTAINING PROTEIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23235">
        <property type="entry name" value="KRUEPPEL-LIKE TRANSCRIPTION FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="14"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="12" id="KW-0539">Nucleus</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="11" id="KW-0804">Transcription</keyword>
    <keyword evidence="9" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="8" id="KW-0832">Ubl conjugation</keyword>
    <keyword evidence="7" id="KW-0862">Zinc</keyword>
    <keyword evidence="6 13" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="30"/>
            <end position="55"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="56"/>
            <end position="83"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="84"/>
            <end position="111"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="112"/>
            <end position="139"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="150"/>
            <end position="177"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="178"/>
            <end position="205"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="206"/>
            <end position="233"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="234"/>
            <end position="261"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="262"/>
            <end position="289"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="290"/>
            <end position="317"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="318"/>
            <end position="345"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="346"/>
            <end position="373"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="374"/>
            <end position="401"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="402"/>
            <end position="429"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1"/>
            <end position="24"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001891.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="452" mass="50668" checksum="65DE05F3E4A02345" modified="2019-05-08" version="1">LPLPQGLGDHVPICSDPTTPHRRSDLERPYTCLDCGKSFYGILHIHQRTHTGEKPYKCLKCGKSFSQSSGLISHERIHTGERPYKCLECGKSFSHRSGLSNHGRIHTGERPYKCLECGKSFSRHSDLSNHGRIHTGEKPYKCXHSGERPYKCLDCGKSFNQSSNLTTHQRVHIGERPYKCLECGKSFSVLSTLIVHQRTHTGEKPYECLDCGKSFYRSSNLTTHQRVHTGEGPYKCLECGKSFSAPSVLLIHQRTHTGEKPYKCLECGKSFSRRSGLISHERIHTGKRPYKCLDCGKSFSRRSDLGNHGRIHTGERPYKCLECGKSFSQSSGLINHSRIHTGERPYTCLDCGKSFYGSSNLTAHQRVHIGEGPYKCLECGKSFSAPSVLHIHQRTHTGEKPYKCLKCGKSFSQSSGLISHERIHTGERPYKCLECGKSFSHPFKVVSGGKSG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJM7</accession>
    <name>A0A452GJM7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Nuclear pore complex protein Nup98-Nup96</fullName>
        </recommendedName>
    </protein>
    <organism evidence="25 26">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="26" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="25" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="25" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="22">Plays a role in the nuclear pore complex (NPC) assembly and/or maintenance. NUP98 and NUP96 are involved in the bidirectional transport across the NPC. May anchor NUP153 and TPR to the NPC. In cooperation with DHX9, plays a role in transcription and alternative splicing activation of a subset of genes. Involved in the localization of DHX9 in discrete intranuclear foci (GLFG-body).</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Nucleus membrane</location>
            <topology evidence="2">Peripheral membrane protein</topology>
            <orientation evidence="2">Nucleoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
            <location evidence="1">Nuclear pore complex</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Nucleus</location>
            <location evidence="3">Nucleoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the nucleoporin GLFG family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000002035.1">
        <property type="protein sequence ID" value="ENSGAGP00000001790.1"/>
        <property type="gene ID" value="ENSGAGG00000001410.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031965">
        <property type="term" value="C:nuclear membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0044614">
        <property type="term" value="C:nuclear pore cytoplasmic filaments"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005654">
        <property type="term" value="C:nucleoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008139">
        <property type="term" value="F:nuclear localization sequence binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008236">
        <property type="term" value="F:serine-type peptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017056">
        <property type="term" value="F:structural constituent of nuclear pore"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051028">
        <property type="term" value="P:mRNA transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000973">
        <property type="term" value="P:post-transcriptional tethering of RNA polymerase II gene DNA at nuclear periphery"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006606">
        <property type="term" value="P:protein import into nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006405">
        <property type="term" value="P:RNA export from nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034398">
        <property type="term" value="P:telomere tethering at nuclear periphery"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.2360:FF:000001">
        <property type="entry name" value="Nuclear pore complex protein Nup98-Nup96"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.690:FF:000001">
        <property type="entry name" value="Nuclear pore complex protein Nup98-Nup96"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1610.10:FF:000001">
        <property type="entry name" value="Nuclear pore complex protein Nup98-Nup96"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.2360">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.690">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1610.10">
        <property type="entry name" value="Peptidase S59, nucleoporin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037665">
        <property type="entry name" value="Nucleoporin_S59-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007230">
        <property type="entry name" value="Nup98_auto-Pept-S59_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036903">
        <property type="entry name" value="Nup98_auto-Pept-S59_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021967">
        <property type="entry name" value="Nup98_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23198:SF6">
        <property type="entry name" value="NUCLEAR PORE COMPLEX PROTEIN NUP98-NUP96"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23198">
        <property type="entry name" value="NUCLEOPORIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04096">
        <property type="entry name" value="Nucleoporin2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12110">
        <property type="entry name" value="Nup96"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21240">
        <property type="entry name" value="Nup98_GLEBS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF82215">
        <property type="entry name" value="C-terminal autoproteolytic domain of nucleoporin nup98"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51434">
        <property type="entry name" value="NUP_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="17" id="KW-0007">Acetylation</keyword>
    <keyword evidence="12" id="KW-0068">Autocatalytic cleavage</keyword>
    <keyword evidence="11" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="7" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="20" id="KW-0472">Membrane</keyword>
    <keyword evidence="13" id="KW-0509">mRNA transport</keyword>
    <keyword evidence="19" id="KW-0906">Nuclear pore complex</keyword>
    <keyword evidence="21" id="KW-0539">Nucleus</keyword>
    <keyword evidence="8" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="9" id="KW-0645">Protease</keyword>
    <keyword evidence="16" id="KW-0653">Protein transport</keyword>
    <keyword evidence="26" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0677">Repeat</keyword>
    <keyword evidence="14" id="KW-0720">Serine protease</keyword>
    <keyword evidence="18" id="KW-0811">Translocation</keyword>
    <keyword evidence="6" id="KW-0813">Transport</keyword>
    <keyword evidence="15" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="Peptidase S59" evidence="24">
        <location>
            <begin position="726"/>
            <end position="868"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="23">
        <location>
            <begin position="501"/>
            <end position="524"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="23">
        <location>
            <begin position="510"/>
            <end position="519"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004567"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004620"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004642"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00008926"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00013472"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022813"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022816"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023010"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023132"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00059309"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS51434"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001790.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1747" mass="189704" checksum="FD3CB04640F2D157" modified="2019-05-08" version="1">MFNKSFGTPFGGSAGGFGTTSTFGQNAGFGTASGGAFGTSAFGSNNNTGSLFGNTQTKPGGLFGSSTFSQPATSSTSTGFGFGTSTGTSNSLFGTASTGGGLFSSQNNAFAQNKPAGFGNFGTSTSSGGLFGTTNTASNPFGGTSGSLFGPTSFTAAPTGTTIKFNPPTGTDTMVKSGVSTNISTKHQCITAMKEYESKSLEELRLEDYQANRKGPSNPAVAAAGLFGSSTATSSAATGLFGSSTTNAGFSYGQNKTAFGTSTTGFGTGTAGSLFGQQPQQTTSLFSKPFGQATTTQNTGFSFGNTSTLGQPNTNTMGLFGATQASPSGSLFGTAANTNAATGFGAGTGTGLFGQANTGFGVGGSTLFGKPAGFGTTTTSAPSFGTTTGSGLFGFGASTAGNSIFGNKPAAGTLGAGLGTGFGTALSAGQTSLFGNTQPKLGGTLGTGAFGAPGFNTSTATLGFGAPQPAVALTDPNVSAAQQAALQQHFNSLTYSPFGDSPLFRNPMSDPKKKEERLKPTNPAAQKALTTPTHYKLTPRPATRVRPKALQTAGSSKSHLFDGLDDDEPSLSNGAFMPKKSIKKLVLKNLNSSNLFSPVSREAEDLDPPSEYPENGERFNFLSRSVDENHRQEGEREEEDDHHEVTRFYTNPIAKPIPQTPENAVHKHQNNVDDTIVALNMRVALRNGLEGSSEDASFHDESLQDDREEENENVYSLHPAGIVLTKAGYYTIPSMEDLAKLTTDRGECIVTDFTIGRKGYGSIYFEGEVNLTNLNLDEIVHIRRKEVIVYPDDDQKPPIGEGLNRRAEVTLDGVWPTDKTSRCLIKSPERLADINYEGRLEAVSRKQGAQFKEYRPETGSWVFKVAHFSKYGLQDSDEEEEEHPSKMDAKKLKTAQVPPQGQLIPQQMALNGKPTSPSQSPEVEQLGRVVELDSDMADITQEPAQDSILEDSITEEPELVPASTHIASSLGINPHVLQIMKASLLADEEDVDLILDHHFGKQPTKMDTSQEICSPRLPISALQGQKRHSSVGGLLQSKFASVSFLQSASLPDFHGPRASSFGNPPSAASWTGLSPLPSAFSMPSAAPEVQLKTVGACRQQGLIPLEKSVTYGKGRLLMDMALFMGRSFRVGWGPNWTLVNSGDKLSGSSEPKGLRNDSMEYGFLPTPVAPKSLSESPFKVHIEKLTLEQRKTDGDQQLYLIPLEIKLKHSTVHVDERCPLLAANPGVSAIHDYADWVRDFCGDSVEIEPVVKYWCLTWTLCEAIWGHLKELEASLEEPSEYIVTLERRKAFSRWLSQTAAARIEEEVSLSQHENHIEAVFSYLTGKRISEACRVAQQSGDHRLALLLSQLVGSPQVRELLTMQLVDWHGLQADCFIQEERLRIFALLAGKPVWQLSERKSINVCSQLDWKRSIAIHLWYLLPPTASISKALNMYEAAFQNTPECEKYACCPLPPYLEDSGCVLEEDDSTERPLRDVCFHLLKLYSDRCYDLNQLLDPRSITSDPLDYRLSWHLWEVLRALNYTHLSEQCQGVLNASYAAQLESEGLWEWAIFVLLHTSNTYTREKAVREFLNRHCVLSETPESWTKETFLTQKLCIPPEWIQEAKAVRARMEGDKHKEALYLFKAGHWNQCHKLVIRHLASDAIINENYNYLKGFLEDLSPPERSTLIQDWDTAGLVYLDYIRVIEMLDRIQQLDCSAYELERLHTKVTSLCNRIEQIQCHNAKDRLAQSGKALFVQCHRCAWQYTG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJM9</accession>
    <name>A0A452GJM9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Peptidase A1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1 8">Belongs to the peptidase A1 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000002155.1">
        <property type="protein sequence ID" value="ENSGAGP00000001896.1"/>
        <property type="gene ID" value="ENSGAGG00000001520.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004190">
        <property type="term" value="F:aspartic-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007586">
        <property type="term" value="P:digestion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.70.10:FF:000004">
        <property type="entry name" value="Pepsin A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.70.10">
        <property type="entry name" value="Acid Proteases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001461">
        <property type="entry name" value="Aspartic_peptidase_A1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001969">
        <property type="entry name" value="Aspartic_peptidase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033121">
        <property type="entry name" value="PEPTIDASE_A1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021109">
        <property type="entry name" value="Peptidase_aspartic_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47966">
        <property type="entry name" value="BETA-SITE APP-CLEAVING ENZYME, ISOFORM A-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47966:SF22">
        <property type="entry name" value="PEPSIN A-3-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00026">
        <property type="entry name" value="Asp"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00792">
        <property type="entry name" value="PEPSIN"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50630">
        <property type="entry name" value="Acid proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00141">
        <property type="entry name" value="ASP_PROTEASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51767">
        <property type="entry name" value="PEPTIDASE_A1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 8" id="KW-0064">Aspartyl protease</keyword>
    <keyword evidence="4" id="KW-0222">Digestion</keyword>
    <keyword evidence="6 7" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5 8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="2 8" id="KW-0645">Protease</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Peptidase A1" evidence="9">
        <location>
            <begin position="33"/>
            <end position="375"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="7">
        <location>
            <begin position="64"/>
            <end position="69"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007447"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022750"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022757"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PIRSR" id="PIRSR601461-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="RuleBase" id="RU000454"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51767"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001896.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="375" mass="41245" checksum="9A096186B9389819" modified="2019-05-08" version="1">SLHIHSKGKSLRQSLKEHEAASEPLTNYMDVDYYGTISIGTPAQDFTVVFDTGSSNLWAPSVYCSSAACTNHNKFNPSDSITYEATSQSLSIQYGTGSMTGILAYDTVQVGGIADTKQIFGLSETEPGSTFYYAPFDGILGLAFPSIASSGATPIFDNMIKEGLVSQDLFSVYLSSDEQSGSVVMFSVIDSSYYSGSLNWIPLSAETYWEITMDRYHPPLGPSPSSGHLEETGKRRCMQELNLQSLDKMRDKGVGITPSLELLERDSPREVILFSRTGNGEGGVPLFRLTEFTSVYLSRNTWRGGEEKRFISLCCETQGIWVLGSPGKVXVFCQTIALPLGGKYQHYIHANKVVFLVYLTLKRLAKGRKGKRHCC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJR4</accession>
    <name>A0A452GJR4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Sushi domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="caution">
        <text evidence="6">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJR4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002083.1">
        <property type="protein sequence ID" value="ENSGAGP00000001831.1"/>
        <property type="gene ID" value="ENSGAGG00000001465.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="CDD" id="cd00033">
        <property type="entry name" value="CCP"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.70.10:FF:000055">
        <property type="entry name" value="Complement decay-accelerating factor, GPI-anchored"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.70.10:FF:000014">
        <property type="entry name" value="Membrane cofactor protein"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.70.10">
        <property type="entry name" value="Complement Module, domain 1"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050350">
        <property type="entry name" value="Compl-Cell_Adhes-Reg"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035976">
        <property type="entry name" value="Sushi/SCR/CCP_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000436">
        <property type="entry name" value="Sushi_SCR_CCP_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19325:SF570">
        <property type="entry name" value="COMPLEMENT COMPONENT 4 BINDING PROTEIN, MEMBRANE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19325">
        <property type="entry name" value="COMPLEMENT COMPONENT-RELATED SUSHI DOMAIN-CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00084">
        <property type="entry name" value="Sushi"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00032">
        <property type="entry name" value="CCP"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57535">
        <property type="entry name" value="Complement control module/SCR domain"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50923">
        <property type="entry name" value="SUSHI"/>
        <property type="match status" value="5"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4 6" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0732">Signal</keyword>
    <keyword evidence="1 6" id="KW-0768">Sushi</keyword>
    <keyword evidence="7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="342"/>
            <end position="362"/>
        </location>
    </feature>
    <feature type="domain" description="Sushi" evidence="8">
        <location>
            <begin position="9"/>
            <end position="72"/>
        </location>
    </feature>
    <feature type="domain" description="Sushi" evidence="8">
        <location>
            <begin position="73"/>
            <end position="132"/>
        </location>
    </feature>
    <feature type="domain" description="Sushi" evidence="8">
        <location>
            <begin position="133"/>
            <end position="194"/>
        </location>
    </feature>
    <feature type="domain" description="Sushi" evidence="8">
        <location>
            <begin position="195"/>
            <end position="254"/>
        </location>
    </feature>
    <feature type="domain" description="Sushi" evidence="8">
        <location>
            <begin position="255"/>
            <end position="317"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="6">
        <location>
            <begin position="11"/>
            <end position="54"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="6">
        <location>
            <begin position="103"/>
            <end position="130"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022659"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00302"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50923"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001831.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="364" mass="38757" checksum="A3760F54FBA78354" modified="2019-05-08" version="1">MLTFFASAIPCLPPPDITHGSHTGQGEEEFSFGSAVIYKCDQGFSLIGNASIHCTTKDNNGEWSGPAPECKVVRCPEPEVKNGKKQSGFGLDYSYGNVVIFECDSGYTLTGSSSVKCEANNSWVPSLPTCLRTCNSPPWLNFAELPGAVNDPYLVGTKLKYSCRPGYILGSGKSPFVTCLANSTWSVDPEFCVGRPCKPLELENGRVDLTDLRFGATATFSCNEGYRLIGTTSAKCVVAGNGVDWDKELPLCQAIPCLPPPDITHGSHTGQGEEFFFGSAVIYKCDQGFSLIGNASIHCTTKDNNGEWSGPAPECKGVTYTTTTTTSSPGGTTDTSGSNTGIMIAIGKMPVILAVLITNLFIRQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJS2</accession>
    <name>A0A452GJS2_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="8">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the ENTREP family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJS2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002088.1">
        <property type="protein sequence ID" value="ENSGAGP00000001836.1"/>
        <property type="gene ID" value="ENSGAGG00000001468.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007237">
        <property type="entry name" value="CD20-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR030431">
        <property type="entry name" value="ENTREP1-3"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR17615:SF8">
        <property type="entry name" value="ENDOSOMAL TRANSMEMBRANE EPSIN INTERACTOR 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR17615">
        <property type="entry name" value="PROTEIN FAM189A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04103">
        <property type="entry name" value="CD20"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 7" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="3 7" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="66"/>
            <end position="88"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="100"/>
            <end position="119"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="125"/>
            <end position="147"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="209"/>
            <end position="232"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="1"/>
            <end position="34"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="345"/>
            <end position="385"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="1"/>
            <end position="19"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="6">
        <location>
            <begin position="20"/>
            <end position="30"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="367"/>
            <end position="379"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00034309"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001836.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="573" mass="61934" checksum="21FA3A184E439B7E" modified="2019-05-08" version="1">MAGLSAALQAAAPGPAASPAQPPPPPPPPLRRFQHYRPPPAAFSPLSARRAAAAAGAQGLLPARPLLSLGLLQLILGCCMVALSFGALSLSGSPQVKNSCPFWAGSSVILSGIIGLTTWKRPVVLLVNLFVLLSVVCVLLNLAGFILGCQGAQFVSSVPRCDLVDLGESKICFCCEEFHPAKCTEKENALKLYPVQSCSAVHLLLKKVLFALCALNALTTTVCLVAAALRYLQIFATRRSCIDESQVSVEEVEEQGRISDPEDFVPPVPPPSYFATFYSCTPRMNRRMLGSDVIPLPHIYGARIKGVEVFCPLDPPPPYEAVVSQISCEQGGAFQIPDVAEAVIGPPEPNDTQVSQVGGDIPDSFSRESLSPSNPSSAPAEQKPACKRAFNPLRKRSKSDPVLHGLPPRGPVLSCEAATQTELKPKLATVILRKGLRARALRARPRSLVDYKSYIDTKLLVARFLEQSSCSMTPDVHELVENIKSVLKSDEEHMEEAITSASFLEQVMTPAQPSTSRTHTLPFRRHPGMLHLESCGDLSTFTTSENKVAERRTQRLEHERPHSLIGVVRETVL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJS5</accession>
    <name>A0A452GJS5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">SUEL-type lectin domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the EVA1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJS5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002215.1">
        <property type="protein sequence ID" value="ENSGAGP00000001947.1"/>
        <property type="gene ID" value="ENSGAGG00000001568.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030246">
        <property type="term" value="F:carbohydrate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd22828">
        <property type="entry name" value="Gal_Rha_Lectin_EVA1_EVA1C_rpt1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd22829">
        <property type="entry name" value="Gal_Rha_Lectin_EVA1_EVA1C_rpt2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.740:FF:000003">
        <property type="entry name" value="Protein eva-1 homolog C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.740">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039500">
        <property type="entry name" value="EVA1_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000922">
        <property type="entry name" value="Lectin_gal-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043159">
        <property type="entry name" value="Lectin_gal-bd_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46780">
        <property type="entry name" value="PROTEIN EVA-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14851">
        <property type="entry name" value="FAM176"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02140">
        <property type="entry name" value="SUEL_Lectin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50228">
        <property type="entry name" value="SUEL_LECTIN"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0430">Lectin</keyword>
    <keyword evidence="7 9" id="KW-0472">Membrane</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="3 9" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 9" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="56"/>
            <end position="76"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="320"/>
            <end position="340"/>
        </location>
    </feature>
    <feature type="domain" description="SUEL-type lectin" evidence="10">
        <location>
            <begin position="97"/>
            <end position="190"/>
        </location>
    </feature>
    <feature type="domain" description="SUEL-type lectin" evidence="10">
        <location>
            <begin position="199"/>
            <end position="291"/>
        </location>
    </feature>
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        <location>
            <begin position="367"/>
            <end position="389"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="369"/>
            <end position="381"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004167"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022734"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50228"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001947.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="443" mass="49165" checksum="94CFA0067BB4ABF7" modified="2019-05-08" version="1">MQRTNPLLMDNRSLRATAPAAMKDEIPPRLESRWLWDSGNRESGTSYMMTVYQRLLGARVSCLGAVLGFLCFSTVLKAAPEFSGYLQKVLKNHTAHACDGEQLGVICPHKTSISILSAFYGRRVPSQNLCPSTGNISGESTTCVSATARQKLLDECQDQRWCQFSVHSPVFGPDPCPGTHKYLIVSYKCRPANHRIKTVCENDNLRLQCRAKSVLAIYSASYGRSMRGKPECDSATRTGLNAECLAPDALRRVSRKCHRKENCTVLANKATFGDPCFPGVKKQLRVSYTCVPRQLFEEVGQDSPDPFSLSDYTHGVPEKVGLYFLCGVSGGLVFLLCIFSPKMAFIQDMKEVLKASKLGHSSELSQAKLRNDHDEDNHNDDSSSDSSFRRLTRTYRASNNIFSPELTAALEGAAEHRAQDGEEIWMLQDSSPYAIHKLKSATK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJS6</accession>
    <name>A0A452GJS6_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="7">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GJS6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002107.1">
        <property type="protein sequence ID" value="ENSGAGP00000001853.1"/>
        <property type="gene ID" value="ENSGAGG00000001477.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="C:dendritic spine"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="F:amyloid-beta binding"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="P:chemical synaptic transmission"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="entry name" value="Mint1_CID"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="PDZ1_APBA1_3-like"/>
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    </dbReference>
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    </dbReference>
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        <property type="entry name" value="PTB_X11"/>
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        <property type="entry name" value="amyloid beta A4 precursor protein-binding family A member 1"/>
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        <property type="entry name" value="Amyloid beta A4 protein-binding family A member"/>
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        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
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        <property type="entry name" value="PDZ"/>
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        <property type="entry name" value="PDZ_sf"/>
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    </dbReference>
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    </dbReference>
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    </dbReference>
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    </dbReference>
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        <property type="entry name" value="PDZ domain-like"/>
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    </dbReference>
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    </dbReference>
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    </dbReference>
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    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="1" id="KW-0813">Transport</keyword>
    <feature type="domain" description="PID" evidence="5">
        <location>
            <begin position="444"/>
            <end position="619"/>
        </location>
    </feature>
    <feature type="domain" description="PDZ" evidence="6">
        <location>
            <begin position="632"/>
            <end position="717"/>
        </location>
    </feature>
    <feature type="domain" description="PDZ" evidence="6">
        <location>
            <begin position="723"/>
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        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="1"/>
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        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
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        </location>
    </feature>
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        <location>
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        </location>
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        <location>
            <begin position="343"/>
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        </location>
    </feature>
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        <location>
            <begin position="1"/>
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        </location>
    </feature>
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            <begin position="23"/>
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        </location>
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        <location>
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        </location>
    </feature>
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        </location>
    </feature>
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        <location>
            <begin position="384"/>
            <end position="400"/>
        </location>
    </feature>
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        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001853.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="813" mass="90651" checksum="AB873763355E2CEC" modified="2019-05-08" version="1">MNHLEGPSEGKVNDEPANGEVNESVEADLDHSEVEEEQQQYAAHYPRHANGSQEGQSGQEEEGMLARSASTESGFHNHTDTAEGDVIATAEDGYYTDRVQENEDESAYAVQYRPESEEYTENAEAEHAEAIQTRTLPNHLHFHSIEHEEAMNAAYSGYVYTHGLFHRGEDEQYAEPYADYGLQEHVYEEIGDTPDLDAREGLRLYEQEREEADNYRKEALGARLHHYDERSDGESDSPEKEAEFAPYPRMDSYEQEEDIDQIVAEVKQSMSSQSLDKAAEDMPEAEQNLECDHTLSSGTLESNGQLPAGSRPTVNSGESVQRYSKEKRDAISLAIKDIKEAIEEVKTRTIRSPYTPDEPKEPIWVMRQDISPTRDCDDQRPLDGDSPSPDSSSPRGAESSSRQHHQDVCGTAEPSTNKESRKSLASFPTYVEVPGPCDPEDLIDGIIFAANYLGSTQLLSDKTPSKNVRMMQAQEAVSRIKAPEGESQPMTEVDLFISTQRIKVLNADTQETMMDHPLRTISYIADIGNIVVLMARRRMPRSNSQDNVEASHPSQDGKRQYKMICHVFESEDAQLIAQSIGQAFSVAYQEFLRANGINPEDLSQKEYSDLLNTQDMYNDDLIHFSKSENCKDVYIEKQKGEILGVVIVESGWGSILPTVIIANMMHGGPAEKSGKLNIGDQIMSINGTSLVGLPLSTCQSIIKGLKNQARIKLNIVRCPPVTTVLIRRPDLRYQLGFSVQNGIICSLMRGGIAERGGVRVGHRIIEINGQSVVATPHEKIVHILSNAVGEIHMKTMPAAMYRLLTAQEQPVYI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GK10</accession>
    <name>A0A452GK10_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="8">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GK10"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002191.1">
        <property type="protein sequence ID" value="ENSGAGP00000001926.1"/>
        <property type="gene ID" value="ENSGAGG00000000203.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="term" value="C:neuromuscular junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098831">
        <property type="term" value="C:presynaptic active zone cytoplasmic component"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042734">
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030672">
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043195">
        <property type="term" value="C:terminal bouton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019992">
        <property type="term" value="F:diacylglycerol binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005543">
        <property type="term" value="F:phospholipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017075">
        <property type="term" value="F:syntaxin-1 binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061789">
        <property type="term" value="P:dense core granule priming"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0099525">
        <property type="term" value="P:presynaptic dense core vesicle exocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035249">
        <property type="term" value="P:synaptic transmission, glutamatergic"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016081">
        <property type="term" value="P:synaptic vesicle docking"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016082">
        <property type="term" value="P:synaptic vesicle priming"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd04027">
        <property type="entry name" value="C2B_Munc13"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.357.50:FF:000001">
        <property type="entry name" value="Protein unc-13 homolog B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000002">
        <property type="entry name" value="Protein unc-13 homolog B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.357.50">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.1100">
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000008">
        <property type="entry name" value="C2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010439">
        <property type="entry name" value="MUN_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014770">
        <property type="entry name" value="Munc13_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014772">
        <property type="entry name" value="Munc13_dom-2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027080">
        <property type="entry name" value="Unc-13"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037302">
        <property type="entry name" value="Unc-13_C2B"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10480">
        <property type="entry name" value="PROTEIN UNC-13 HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10480:SF2">
        <property type="entry name" value="PROTEIN UNC-13 HOMOLOG C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00168">
        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06292">
        <property type="entry name" value="MUN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00360">
        <property type="entry name" value="C2DOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00239">
        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM01145">
        <property type="entry name" value="DUF1041"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49562">
        <property type="entry name" value="C2 domain (Calcium/lipid-binding domain, CaLB)"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50004">
        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51258">
        <property type="entry name" value="MHD1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51259">
        <property type="entry name" value="MHD2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0106">Calcium</keyword>
    <keyword evidence="1" id="KW-0268">Exocytosis</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0862">Zinc</keyword>
    <keyword evidence="3" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2" evidence="5">
        <location>
            <begin position="36"/>
            <end position="160"/>
        </location>
    </feature>
    <feature type="domain" description="MHD1" evidence="6">
        <location>
            <begin position="464"/>
            <end position="607"/>
        </location>
    </feature>
    <feature type="domain" description="MHD2" evidence="7">
        <location>
            <begin position="706"/>
            <end position="849"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="5">
        <location>
            <begin position="838"/>
            <end position="965"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS51258"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS51259"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001926.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1010" mass="116831" checksum="3D6F81697E4DCA33" modified="2019-05-08" version="1">IIAMKERMKIRERNRPEVFEVIQEMFQISKEDFVQYTKAAKQCVLDGTSKWSAKITITVVCAQGLQAKDKTGSSDPYVTVQVGKTKRRTKTIFGNLNPVWDEKFYFECHNSTDRIKVRVWDEDDDIKSRVKQHFKKESDDFLGQTIIEVRTLSGEMDVWYNLEKRTDKSAVSGAIRLKINVEIKGEEKVAPYHVQYTYLHENLFHYLTEVKSNGSVKIPEVRGDEAWKVYFDAAAQEIVDEFAMRYGIEYIYQAMTHFSCLSSKYMCSGVPAVMSTLLANINAFYAHTIATTNVSASDRFAATNFGREKFIKLLDQLHNSLRIDLSKYRDNFPASNSERLQDLKSTVDLLTSITFFRMKVLELQSPPRASMVVKDCVRACLDSTYKYIFDNCYELYSQLMDQEVPREEQGPTTKNLDFWAQLITLMVTIIDEDKTAYTPVLNQFPQELNMGKVSAEIMWTLFAQDMKYALEEHEKQRLCKSTDYMNLHFKVKWFYNEYVRELPALQDAVPEYSLWFEPFVIQWLDENEDVSMEFLHGALERDKKDGFQQTSDHALFSCSVVDVFTQLNQSFEIIKKLECPNPEALSHLMRRFAKTINKVLLQYAAVISNDFNSYCDKEMLPCILMNNIQQLRVQLEKMFESMGGKELMEQNLNSHLLSYNSSTLFFLFSFQLVVEECVRQMSFELNQMRGNGNISANKNNAAIDAEIVLRPLMDFLDKILSLSAKICEKTVLKRVLKELWKSVLNKIEKQIVLPPLTDQTGPQMIFSAAKDLGQLSKLKDHVTREEARSLTLRQCAIMEVALSTIKVQLKYMMEVFACTMVWHKCTRVIKILVLLSQCAWGAQTSCKVVLDSPRWAEHKAMVPRSLPLPRRWQTAAMFRPFVEVCILGPNLSDKKRKHGTKTKSNTWSPKYNETFHILSNEEGLRAYELHLSVKDYCFAREDRIVGMTVIQLQNTAEKGSCASWYPLLKNIFMDDTGLTILRILSQRTNDEVAKEFVRLKSETRSTEETV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GK31</accession>
    <name>A0A452GK31_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3 7">Carbonic anhydrase</fullName>
            <ecNumber evidence="3 7">4.2.1.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="7">Reversible hydration of carbon dioxide.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="7">
            <text>hydrogencarbonate + H(+) = CO2 + H2O</text>
            <dbReference type="Rhea" id="RHEA:10748"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:17544"/>
            <dbReference type="EC" id="4.2.1.1"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 7">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <comment type="similarity">
        <text evidence="2 7">Belongs to the alpha-carbonic anhydrase family.</text>
    </comment>
    <dbReference type="EC" id="4.2.1.1" evidence="3 7"/>
    <dbReference type="AlphaFoldDB" id="A0A452GK31"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002360.1">
        <property type="protein sequence ID" value="ENSGAGP00000002073.1"/>
        <property type="gene ID" value="ENSGAGG00000001445.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004089">
        <property type="term" value="F:carbonate dehydratase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.200.10:FF:000001">
        <property type="entry name" value="Carbonic anhydrase 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.200.10">
        <property type="entry name" value="Alpha carbonic anhydrase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001148">
        <property type="entry name" value="CA_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036398">
        <property type="entry name" value="CA_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023561">
        <property type="entry name" value="Carbonic_anhydrase_a-class"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018338">
        <property type="entry name" value="Carbonic_anhydrase_a-class_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18952">
        <property type="entry name" value="CARBONIC ANHYDRASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18952:SF25">
        <property type="entry name" value="CARBONIC ANHYDRASE 5B, MITOCHONDRIAL-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00194">
        <property type="entry name" value="Carb_anhydrase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01057">
        <property type="entry name" value="Carb_anhydrase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51069">
        <property type="entry name" value="Carbonic anhydrase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00162">
        <property type="entry name" value="ALPHA_CA_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51144">
        <property type="entry name" value="ALPHA_CA_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6 7" id="KW-0456">Lyase</keyword>
    <keyword evidence="4 7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 7" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="Alpha-carbonic anhydrase" evidence="8">
        <location>
            <begin position="29"/>
            <end position="284"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010718"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012925"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU367011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51144"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002073.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="305" mass="34970" checksum="7CC7E63C78CEA2A2" modified="2019-05-08" version="1">TLRCKVLGRLQKSWPGRAETLGRTVRKMTSHGCEGVHPLWQSELSVSGGTRQSPINIQWRDSVYDPHLKPLKIHYDPTTCLYIWNNGYSFQVEFDDSADKSVIMGGPLENHYRLKQFHFHWGAINEWGSEHMVDSKVYPAELHLVHWNPVTSTSFEEAIMENNGLVVIGVFLKLGAHHAGLQKLVDALPAVKHKDTIVEFDVFDPSCLMPSCPDYWTYDGSLTTPPLTESVTWIIKKTPIEVDQNQLEEFRMLLFTSAGEEEKRMVDNFRPLQPLMNRTVRSSFRNHHLLNIETRSSELAHQHSP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GK33</accession>
    <name>A0A452GK33_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">DS cell adhesion molecule like 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="12">Synapse</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000002238.1">
        <property type="protein sequence ID" value="ENSGAGP00000001967.1"/>
        <property type="gene ID" value="ENSGAGG00000001306.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030424">
        <property type="term" value="C:axon"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045202">
        <property type="term" value="C:synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098632">
        <property type="term" value="F:cell-cell adhesion mediator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007411">
        <property type="term" value="P:axon guidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060219">
        <property type="term" value="P:camera-type eye photoreceptor cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070593">
        <property type="term" value="P:dendrite self-avoidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007156">
        <property type="term" value="P:homophilic cell adhesion via plasma membrane adhesion molecules"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010842">
        <property type="term" value="P:retina layer formation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007416">
        <property type="term" value="P:synapse assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="CDD" id="cd00096">
        <property type="entry name" value="Ig"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="CDD" id="cd05734">
        <property type="entry name" value="Ig_DSCAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05735">
        <property type="entry name" value="Ig_DSCAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd20958">
        <property type="entry name" value="IgI_5_Dscam"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056754">
        <property type="entry name" value="DSCAM/DSCAML_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
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        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013098">
        <property type="entry name" value="Ig_I-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013106">
        <property type="entry name" value="Ig_V-set"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10075">
        <property type="entry name" value="BASIGIN RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10075:SF72">
        <property type="entry name" value="CELL ADHESION MOLECULE DSCAML1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25059">
        <property type="entry name" value="FN3_DSCAM-DSCAML_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07679">
        <property type="entry name" value="I-set"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13927">
        <property type="entry name" value="Ig_3"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SMART" id="SM00406">
        <property type="entry name" value="IGv"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="8"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="10" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="11" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="9 14" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="3 15" id="KW-0732">Signal</keyword>
    <keyword evidence="8" id="KW-0770">Synapse</keyword>
    <keyword evidence="2 14" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 14" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="15">
        <location>
            <begin position="1"/>
            <end position="19"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5045309832" description="DS cell adhesion molecule like 1" evidence="15">
        <location>
            <begin position="20"/>
            <end position="1750"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="1456"/>
            <end position="1476"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="16">
        <location>
            <begin position="92"/>
            <end position="176"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="16">
        <location>
            <begin position="180"/>
            <end position="265"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="16">
        <location>
            <begin position="274"/>
            <end position="367"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="16">
        <location>
            <begin position="372"/>
            <end position="452"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="16">
        <location>
            <begin position="462"/>
            <end position="551"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="16">
        <location>
            <begin position="556"/>
            <end position="650"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="16">
        <location>
            <begin position="654"/>
            <end position="751"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="17">
        <location>
            <begin position="753"/>
            <end position="850"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="17">
        <location>
            <begin position="855"/>
            <end position="954"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="17">
        <location>
            <begin position="959"/>
            <end position="1055"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="17">
        <location>
            <begin position="1059"/>
            <end position="1154"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="16">
        <location>
            <begin position="1144"/>
            <end position="1231"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="17">
        <location>
            <begin position="1247"/>
            <end position="1341"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="17">
        <location>
            <begin position="1342"/>
            <end position="1442"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="1543"/>
            <end position="1569"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="1609"/>
            <end position="1630"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="1668"/>
            <end position="1690"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="13">
        <location>
            <begin position="1560"/>
            <end position="1569"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004167"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00034103"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001967.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="20">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1750" mass="192243" checksum="10D2EE006A0F73CA" modified="2025-10-08" version="2">MAVVFVCTLLECFWASSGATPLSNNQTFSSLLLVSPENRFFITFYGGLYISDVQKEDALSTYRCITKHKYSGETRQSNGARLSVSDPAESIPTILDGFQSRELKAGQLVELPCIASGYPNPAIRWIKDGRPLPADSRWTKRITGLTISDLRGEDSGTYICEVTNTFGSAEVTGTLTVIDPLHVSLTPKKLKTGIGSTVILSCALSGSPEYALRWYRNTDLVVLDNYISIRGISNETLLITAAQKSHSGAYQCFATRKAQTAQDFSIIALEDGTPRIVSSFSEKVVNPGEQFSLMCAAKGAPPPTVTWALDDEPIQRDNGHRTNQYTMSDGTTVSHMNVTSPQIKDGGVYRCTARNSVGSAEYQARINVRGPPSIRAMKNITAVAGRDTFINCRVIGYPYYSIKWYKDALLLPDNHRQVVFENGTLKLMDVQKGMDEGEYLCSVLIQPQLSISQSVHVTVKVPPLIQPFEFPPASIGQLLYIPCVVSSGDMPIRITWRKDGQVIISGSGVTIESKEFMSSLQISSVSLKHNGNYTCIASNAAATVSRERQLIVRVPPRFVVQPNNQDGIYGKAGVLNCSVDGYPPPKVMWKHAKGSGNPQQYHPVPLTGRIQILPNSSLLIRHVLEEDIGYYLCQASNGVGTDISKSMFLTVKIPAMITSHPNTTIAIKGQMKELNCTARGERPIIIRWEKGDTVIDPDRNMRYAISTKDNGDEVISTLKLKPADRGDSVFFSCHAINSYGEDRGLIQLTVQEPPDPPELEIREVKARSMNLRWTQRFDGNSIITGFDIEYKNKSDSWDFKQSTRNISPTINQANIVDLHPASVYSIRMYSFNKIGRSEPSKELTISTEEAAPDGPPMDVTLQPITSQSIQVTWKAPKKELQNGVIRGYQIGYRENSPGSNGQYSIVEMKATGDSEVYTLDNLKKFAQYGVVVQAFNRAGTGPSSSEINATTLEDVPSQPPENVRALSITSDVAVISWSEPPRSTLNGVLKGYRVIFWSLYMDGEWGEMQNITTTRERVELRGMEKFTNYSVQVLAYTQAGDGVRSSVLYIQTKEDIPGPPARIKAVPSSASSVVVSWLPPTKPNGIIRKYTIFCSSPGSGQPAPSEYETSPEQLFYRIAHLNRGQQYMLWVAAVTSAGRGNISEKVTIEPAGKAPAKIISFGGTVTTPWMKDVRLPCNSVGEPAPAIKWTKDREAGWRLVDGHRLIQANGTLVLRSVKAEDSGYYTCTATNTWGFDTIIINLLVQVPPDQPRLTVSKTSASSITLAWIPGDNGGSSIRGFVLQYSVDNSEEWKDVFISSSERAFQLDSLKCGTWYKVKLAAKNSVGAGRISEIIEAKTHGREPSFSKDQHLFTHINSTHARLNLQGWSSGGCPIMAIVLEYRPKGNWVWQSLKTNNSNEVFLTELREATWYELRMKACNSAGCGNETTQFATLDYDGSTIPPIKSAQGEGDDVKKLFTIACPVILATLGVALLFIIRKKRKEKRLKRLRGELSELSPQTVLLAGDDKATIPVTDTEFSQAVNPQSFCTGVSLHHPALIQNTGPLIDMSDIRPGTNPVSRKSVKSAHSTRNRYSSQWTLTKCQASTPARTLTSDWRTVGSQHGIMVTESDSYSASLSQDTDKGRNSMVSTESASSTYEELARAYEHAKLEEQLQHAKFEITECFISDSSSDQMTTGTNENADSMTSMSTPSEPGICRFTASPPKPQDSDRGKNVAVPIPHRANKKKRGLRAIGTGETFGMGGACTGEMGST</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GK34</accession>
    <name>A0A452GK34_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="10">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000002222.1">
        <property type="protein sequence ID" value="ENSGAGP00000001954.1"/>
        <property type="gene ID" value="ENSGAGG00000001570.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046983">
        <property type="term" value="F:protein dimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000977">
        <property type="term" value="F:RNA polymerase II transcription regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd19731">
        <property type="entry name" value="bHLH-PAS_NPAS1_PASD5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00130">
        <property type="entry name" value="PAS"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.450.20:FF:000025">
        <property type="entry name" value="Neuronal PAS domain protein 3 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.450.20:FF:000021">
        <property type="entry name" value="Neuronal PAS domain-containing protein 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.280.10:FF:000007">
        <property type="entry name" value="single-minded homolog 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.280.10">
        <property type="entry name" value="Helix-loop-helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.450.20">
        <property type="entry name" value="PAS domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011598">
        <property type="entry name" value="bHLH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036638">
        <property type="entry name" value="HLH_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000014">
        <property type="entry name" value="PAS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035965">
        <property type="entry name" value="PAS-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013655">
        <property type="entry name" value="PAS_fold_3"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23043">
        <property type="entry name" value="HYPOXIA-INDUCIBLE FACTOR 1 ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23043:SF25">
        <property type="entry name" value="NEURONAL PAS DOMAIN-CONTAINING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23171">
        <property type="entry name" value="bHLH_HIF1A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08447">
        <property type="entry name" value="PAS_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00353">
        <property type="entry name" value="HLH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00091">
        <property type="entry name" value="PAS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47459">
        <property type="entry name" value="HLH, helix-loop-helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55785">
        <property type="entry name" value="PYP-like sensor domain (PAS domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50888">
        <property type="entry name" value="BHLH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50112">
        <property type="entry name" value="PAS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="6" id="KW-0539">Nucleus</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0804">Transcription</keyword>
    <keyword evidence="3" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="BHLH" evidence="9">
        <location>
            <begin position="35"/>
            <end position="88"/>
        </location>
    </feature>
    <feature type="domain" description="PAS" evidence="8">
        <location>
            <begin position="292"/>
            <end position="347"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="96"/>
            <end position="117"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="165"/>
            <end position="212"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="414"/>
            <end position="540"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="653"/>
            <end position="692"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="182"/>
            <end position="199"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="424"/>
            <end position="437"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="7">
        <location>
            <begin position="523"/>
            <end position="534"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50112"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50888"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001954.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="864" mass="93593" checksum="2BDF35D962BF9485" modified="2019-05-08" version="1">MAAPFGNEVKCVSVEWDFLQGLLVKNQPVPCLQALRKEKSRNAARSRRGKENFEFYELAKMLPLPGAITSQLDKASIVRLTISYLKMRDFASHGEPPWGLRAEGSPPPGPASKGPGRRNMAALLTKLFEQHLGGHILQVELTGSSVFDYIHPGDHPEVAEQLGLKPPFDAVGSPLRSQKPPGLSGASSSSSCSSSLNEASEPDPAGSHARPRHDALERSFFVRMKSTLTKRGLHVKTSGYKVIHVTGRLRPRLPSFSHAHSVLSRVLGLVALAHTLPPSTLSEVRIECHMFVFRVNMDLQIVYCESRISDYMDLCPSELVGKSCYQFIHGEDVEGIRQSHLDLLKKGQVVTRYYRWLQKNGGFVWIQSCATISVNIKNPSEKNMVWVNYILSKPECKNMALDIFQLPGIPARQTDASDVSDLEPDFKESDHGLEHKQGNQFLPKAAGVLAKGYSRSSPDDGTRCSMAENDASSPETSYKDEEVSEGAGASDAESRRTGPRSKRIKLEFQPDSPPQEASLVSTEESDSGSEEEPDLPPQPVALERRVNGYNMAAGRNAGPSRPCTSEFTSVIQTQRSYPPKASAVLSIKTEQSQAPKYDKAALWTYPSARDALHEGPPYTVSKALNLEKPILRQATSHLYVSILDSVLTPPEMDSSGGLAKAPFSTSPRAMLPAPSAETLSPPLSGSPCEERATSTPFAPMVYPGEMEALQRFHAGNVVLPLVHQLGXFHAGNVVLPLVHQLGGHHGIASTTGSQSLYTTSTIRYAPADVTLAVPGSVLPASHAINLMDIGSAESKTSMELMYHHIQRLNMVAPFGNSANATGLAQISGATGGVFTAAESLFSTLPFSVASSSIHSLSAAERKED</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GK35</accession>
    <name>A0A452GK35_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Haptoglobin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">As a result of hemolysis, hemoglobin is found to accumulate in the kidney and is secreted in the urine. Haptoglobin captures, and combines with free plasma hemoglobin to allow hepatic recycling of heme iron and to prevent kidney damage. Haptoglobin also acts as an antioxidant, has antibacterial activity and plays a role in modulating many aspects of the acute phase response. Hemoglobin/haptoglobin complexes are rapidly cleared by the macrophage CD163 scavenger receptor expressed on the surface of liver Kupfer cells through an endocytic lysosomal degradation pathway.</text>
    </comment>
    <comment type="subunit">
        <text evidence="15">Tetramer of two alpha and two beta chains; disulfide-linked. The hemoglobin/haptoglobin complex is composed of a haptoglobin dimer bound to two hemoglobin alpha-beta dimers. Interacts with CD163. Interacts with ERGIC3.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="16">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GK35"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002339.1">
        <property type="protein sequence ID" value="ENSGAGP00000002055.1"/>
        <property type="gene ID" value="ENSGAGG00000001627.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072562">
        <property type="term" value="C:blood microparticle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016209">
        <property type="term" value="F:antioxidant activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030492">
        <property type="term" value="F:hemoglobin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006953">
        <property type="term" value="P:acute-phase response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042742">
        <property type="term" value="P:defense response to bacterium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031638">
        <property type="term" value="P:zymogen activation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00033">
        <property type="entry name" value="CCP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00190">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.10:FF:000054">
        <property type="entry name" value="Complement C1r subcomponent"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.70.10">
        <property type="entry name" value="Complement Module, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.10.10">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009003">
        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043504">
        <property type="entry name" value="Peptidase_S1_PA_chymotrypsin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035976">
        <property type="entry name" value="Sushi/SCR/CCP_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000436">
        <property type="entry name" value="Sushi_SCR_CCP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001254">
        <property type="entry name" value="Trypsin_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24255">
        <property type="entry name" value="COMPLEMENT COMPONENT 1, S SUBCOMPONENT-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24255:SF27">
        <property type="entry name" value="HAPTOGLOBIN-RELATED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00089">
        <property type="entry name" value="Trypsin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00020">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57535">
        <property type="entry name" value="Complement control module/SCR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50494">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50923">
        <property type="entry name" value="SUSHI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50240">
        <property type="entry name" value="TRYPSIN_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0011">Acute phase</keyword>
    <keyword evidence="10" id="KW-0044">Antibiotic</keyword>
    <keyword evidence="5" id="KW-0929">Antimicrobial</keyword>
    <keyword evidence="9" id="KW-0049">Antioxidant</keyword>
    <keyword evidence="11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13" id="KW-0351">Hemoglobin-binding</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="6" id="KW-0721">Serine protease homolog</keyword>
    <keyword evidence="8 17" id="KW-0732">Signal</keyword>
    <keyword evidence="7 16" id="KW-0768">Sushi</keyword>
    <feature type="signal peptide" evidence="17">
        <location>
            <begin position="1"/>
            <end position="17"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019568212" description="Haptoglobin" evidence="17">
        <location>
            <begin position="18"/>
            <end position="329"/>
        </location>
    </feature>
    <feature type="domain" description="Sushi" evidence="19">
        <location>
            <begin position="23"/>
            <end position="80"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase S1" evidence="18">
        <location>
            <begin position="95"/>
            <end position="327"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00014396"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022486"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022529"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022542"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022659"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022862"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023022"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023227"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00025619"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00046627"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00302"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50923"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002055.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="329" mass="36428" checksum="F99C11685D525855" modified="2019-05-08" version="1">CRTPMLLVASLLWGALASPDTSQSCPVPRWVEHGQVEHLARYRCDPYYQLRSRGDGLYRCSQQHVWVNEAAGEVLPICEPVCGKPKNPARQVQRIIGGMMAAKNSFPWQGQLLSRHNHTAGATLISDQWLLTTGRNLYLGHSENSTLEEIAPTLRLFLGRETPAGAVERIVLHPEFPGAVDLALLKLKHKVPIGEAVMPICLAQQDYAKVGRVGFVSGWGWTVLLEHPKQLKYVLLPVADSGSCQAYYQAHSLQPLQNNHTFCVGMSELRESTCLGDAGSAFAIHDPEDDTWYAAGILSFDRSCLAARYGVYVRVLSVLDWIKETMAAH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GK37</accession>
    <name>A0A452GK37_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">C-type lectin domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GK37"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002366.1">
        <property type="protein sequence ID" value="ENSGAGP00000002079.1"/>
        <property type="gene ID" value="ENSGAGG00000001663.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030246">
        <property type="term" value="F:carbohydrate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000015">
        <property type="entry name" value="C-type lectin Cal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.100.10">
        <property type="entry name" value="Mannose-Binding Protein A, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001304">
        <property type="entry name" value="C-type_lectin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016186">
        <property type="entry name" value="C-type_lectin-like/link_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050111">
        <property type="entry name" value="C-type_lectin/snaclec_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016187">
        <property type="entry name" value="CTDL_fold"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22803">
        <property type="entry name" value="MANNOSE, PHOSPHOLIPASE, LECTIN RECEPTOR RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00059">
        <property type="entry name" value="Lectin_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01504">
        <property type="entry name" value="PNCREATITSAP"/>
    </dbReference>
    <dbReference type="SMART" id="SM00034">
        <property type="entry name" value="CLECT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56436">
        <property type="entry name" value="C-type lectin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50041">
        <property type="entry name" value="C_TYPE_LECTIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0430">Lectin</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <feature type="domain" description="C-type lectin" evidence="5">
        <location>
            <begin position="27"/>
            <end position="138"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022734"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50041"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002079.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="180" mass="20124" checksum="9D8197C3F0FBE82E" modified="2019-05-08" version="1">APARGERLLPAPPGVEATSCPNDWLYYHDHCYGFFPEKTTWSEAEVECHYQHKGGHLASILSEAEGDLVARYISQSGSKDDVWIGLHNPRHNKRWRWTDGSLYHYSAWNPGEPNNLGNIEYCTELLNFRGKQMLKRTCTSNGAKTGPWPHPPPLSTDMASCPGTLPLSLSSHCHSAEIIP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GK38</accession>
    <name>A0A452GK38_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Leucine-rich repeat-containing protein 51</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GK38"/>
    <dbReference type="STRING" id="38772.ENSGAGP00000001972"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002245.1">
        <property type="protein sequence ID" value="ENSGAGP00000001972.1"/>
        <property type="gene ID" value="ENSGAGG00000001592.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000002250.1">
        <property type="protein sequence ID" value="ENSGAGP00000001977.1"/>
        <property type="gene ID" value="ENSGAGG00000001592.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001611">
        <property type="entry name" value="Leu-rich_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46545">
        <property type="entry name" value="LEUCINE-RICH REPEAT-CONTAINING PROTEIN 51"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46545:SF1">
        <property type="entry name" value="LEUCINE-RICH REPEAT-CONTAINING PROTEIN 51"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14580">
        <property type="entry name" value="LRR_9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52058">
        <property type="entry name" value="L domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51450">
        <property type="entry name" value="LRR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0433">Leucine-rich repeat</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00014223"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022614"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001972.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="193" mass="22097" checksum="A6A8F0428946DE04" modified="2019-05-08" version="1">MNVRWDCCKISLQAPPLDYSFRGISFIQDLLTEQPRAGLKVIKHSAGGKLLTQAVRLNNNTINELTDFASTMEQLLEYPDELSWVDLSFNDLPTIDPVLTMYSNLRALNLHGNSIQSLSEVDKLAVLPHLRTLTLHGNPIEEEKGYRSYVLSVLPQLKSFDFSGVTKQDRSTATFWRRMNVKPKKVKKRRDDY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GK42</accession>
    <name>A0A452GK42_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="16">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GK42"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002230.1">
        <property type="protein sequence ID" value="ENSGAGP00000001959.1"/>
        <property type="gene ID" value="ENSGAGG00000001571.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071011">
        <property type="term" value="C:precatalytic spliceosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003725">
        <property type="term" value="F:double-stranded RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003727">
        <property type="term" value="F:single-stranded RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051607">
        <property type="term" value="P:defense response to virus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd19910">
        <property type="entry name" value="DSRM_ILF3_rpt1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd19912">
        <property type="entry name" value="DSRM_ILF3_rpt2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.1410.40:FF:000001">
        <property type="entry name" value="interleukin enhancer-binding factor 3 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.20:FF:000006">
        <property type="entry name" value="interleukin enhancer-binding factor 3 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.20:FF:000008">
        <property type="entry name" value="interleukin enhancer-binding factor 3 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1410.40">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.20">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.460.10">
        <property type="entry name" value="Beta Polymerase, domain 2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014720">
        <property type="entry name" value="dsRBD_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033099">
        <property type="entry name" value="DSRM1_ILF3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006561">
        <property type="entry name" value="DZF_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049402">
        <property type="entry name" value="DZF_dom_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049401">
        <property type="entry name" value="DZF_dom_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043519">
        <property type="entry name" value="NT_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45762:SF4">
        <property type="entry name" value="INTERLEUKIN ENHANCER-BINDING FACTOR 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45762">
        <property type="entry name" value="ZINC FINGER RNA-BINDING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00035">
        <property type="entry name" value="dsrm"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20965">
        <property type="entry name" value="DZF_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07528">
        <property type="entry name" value="DZF_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00358">
        <property type="entry name" value="DSRM"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00572">
        <property type="entry name" value="DZF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54768">
        <property type="entry name" value="dsRNA-binding domain-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50137">
        <property type="entry name" value="DS_RBD"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51703">
        <property type="entry name" value="DZF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8" id="KW-0051">Antiviral defense</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="6 12" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="10" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="DZF" evidence="15">
        <location>
            <begin position="5"/>
            <end position="378"/>
        </location>
    </feature>
    <feature type="domain" description="DRBM" evidence="14">
        <location>
            <begin position="398"/>
            <end position="467"/>
        </location>
    </feature>
    <feature type="domain" description="DRBM" evidence="14">
        <location>
            <begin position="529"/>
            <end position="595"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="51"/>
            <end position="80"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="363"/>
            <end position="401"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="467"/>
            <end position="530"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="641"/>
            <end position="665"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="13">
        <location>
            <begin position="60"/>
            <end position="73"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="13">
        <location>
            <begin position="372"/>
            <end position="383"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="13">
        <location>
            <begin position="472"/>
            <end position="481"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="13">
        <location>
            <begin position="499"/>
            <end position="517"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="13">
        <location>
            <begin position="649"/>
            <end position="665"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023118"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00266"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50137"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS51703"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001959.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="708" mass="76890" checksum="1CFD0B7153C24384" modified="2019-05-08" version="1">MRPMRIFVNDDRHVMAKHSAVYPTQEELEAVQNMVSHTERALKAVSDWIDEQEKVSGEQPEPEAMETAAEEENKEGGDQKAAEHLTRTLRGVMRVGLVAKGLLLKGDLDLELVLLCKEKPTTGLLDKVAENLGVQLATITEDKYEIIQSVSEAAIIIKNTQEPPLTLTIHLTSPVVREEMEKLLAGETLSVNDTPDVLDRQKCLAALASLRHAKWFQARANGLKSCVIVIRVLRDLCTRVPTWGPLRGWPLELLCEKSIGTANRPMGAGEALRRVLECLASGIVMPDGSGIYDPCEKEATDAIGHLDRQQREDITQSAQHALRLAAFGQLHKVLGMDPLPSKMPKKPKNENPVDYTVQIPPSTTYAITPMKRPMEEDGEEKSPSKKKKKIQKKEEKTEPPQAMNALMKLNQLKPGLQYKLVSQTGPVHAPIFTMSVEVDGSTFEASGPSKKTAKLHVAVKVLQDMGLPTGVEGKESGKGDESAEETEQKPVVVAPPPVVETISTPSAASPPSEQTSENVKQQGPILTKHGKNPVMELNEKRRGLKYELISETGGSHDKRFVMEVEVDGQKFQGAGSNKKVAKAYAALAALEKLFPDTPVAIEPNKKKRAPVPARGGPKFAVKQHNPGFGMGGPMHNEVPPPPNLRGRGRGGNIRGRGRGRGGFGGGNHGGYMNAGAGYGSYGYGGNSATAGYSDFFTDCYGYHDFGSS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GK44</accession>
    <name>A0A452GK44_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">Serine/threonine-protein kinase TOR</fullName>
            <ecNumber evidence="1">2.7.11.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="5">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the PI3/PI4-kinase family.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002377.1">
        <property type="protein sequence ID" value="ENSGAGP00000002089.1"/>
        <property type="gene ID" value="ENSGAGG00000000429.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031931">
        <property type="term" value="C:TORC1 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031932">
        <property type="term" value="C:TORC2 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016242">
        <property type="term" value="P:negative regulation of macroautophagy"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0038202">
        <property type="term" value="P:TORC1 signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000060">
        <property type="entry name" value="Serine/threonine-protein kinase mTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000488">
        <property type="entry name" value="Serine/threonine-protein kinase mTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000532">
        <property type="entry name" value="Serine/threonine-protein kinase mTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050517">
        <property type="entry name" value="DDR_Repair_Kinase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024585">
        <property type="entry name" value="mTOR_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11139">
        <property type="entry name" value="ATAXIA TELANGIECTASIA MUTATED ATM -RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11139:SF9">
        <property type="entry name" value="SERINE_THREONINE-PROTEIN KINASE MTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11865">
        <property type="entry name" value="mTOR_dom"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01346">
        <property type="entry name" value="DUF3385"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="1" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="1" id="KW-0418">Kinase</keyword>
    <keyword evidence="1" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="1" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Serine/threonine-protein kinase mTOR" evidence="2">
        <location>
            <begin position="852"/>
            <end position="1020"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="RuleBase" id="RU364109"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="SMART" id="SM01346"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002089.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1029" mass="114886" checksum="DE89BFE42CCD8077" modified="2019-05-08" version="1">GLGSRPXQLAAIMSSTVSILQQFASGLKSRNEEIRAKAAKDLQHYVTMELREMSQEESTRFYDQLNHHIFELVSSSDANERKGGILAIASLIGVEGGNATRIGRFANYLRNLLPSNDPVVMEMASKAIGRLALAGDTFTAEYVEFEVKRALEWLGADRNEGRRHAAVLVLRELAISVPTFFFQQVQPFFDNIFVAVWDPKQAIREGAVSALRACLILTTQREPKEMQKPQWYRHTYEEAEKGFDETLAKEKGMNRDDRIHGALLILNELVRISSMEGERLREEMEEITQQQMVHDKYCKDLLGFGTKPRHITPFTSFQSVQPPQSNALVGLLGYSPHQGIMGFGASPIPAKSALVESRCCRDLMEEKFDQVCLWVLKCRTSKNSLIQMTILNLLPRLAAFRPSAFTADQYLPDTMNHILSCVKKEKERTAAFQALGLLSVAVRSEFQAYLPKVLEIIKAALPPKDFAHKRQKSMQVDATVFTCISMLARAMGPCIQQDIKELLEPMLAVGLSPALTAVLYDLSRQIPQLKKDIQDGLLKMLSLVLMHKPLRHPGMPKGLAHQLASPSLTNIPEASDVGSITLALRTLGSFEFEGHSLTQFVRHCADHFLNSEHKEIRMEAARTCSRLLTPSIHLISGHAHVVSQTAVQVVADVLSKLLVVGITDPDPDIRYCVLASLDERFDAHLAQAENLQALFVALNDQVFEIRELAICTVGRLSSMNPAFVMPFLRKMLIQILTELEHSGVGRIKEQSARMLGHLVSNAPRLIRPYMEPILKALIVKLKDPDPDPNPGVINNVLATIGELAQVSGLEMRKWVDELFIIIMDMLQDSSLLAKRQVALWTLGQLVASTGYVVEPYRKYPTLLDVLLNFLKTEQNQGTRREAIRVLGLLGALDPYKHKVNIGMIDQSRDASAVSLSESKSSQDSSDYSTSEMLVNMGNLPLDEFYPAVSMVALMRIFRDQSLSHHHTMVVQAITFIFKSLGLKCVQFLPQVMPTFLNVIRVCDGAIREVSVVCAILSLLELNLGFANYC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMP3</accession>
    <name>A0A452HMP3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Slit homolog 2 protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="15">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018502.1">
        <property type="protein sequence ID" value="ENSGAGP00000016206.1"/>
        <property type="gene ID" value="ENSGAGG00000011388.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048495">
        <property type="term" value="F:Roundabout binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072359">
        <property type="term" value="P:circulatory system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060560">
        <property type="term" value="P:developmental growth involved in morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060562">
        <property type="term" value="P:epithelial tube morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001822">
        <property type="term" value="P:kidney development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050919">
        <property type="term" value="P:negative chemotaxis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030308">
        <property type="term" value="P:negative regulation of cell growth"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010605">
        <property type="term" value="P:negative regulation of macromolecule metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010977">
        <property type="term" value="P:negative regulation of neuron projection development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0021772">
        <property type="term" value="P:olfactory bulb development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050770">
        <property type="term" value="P:regulation of axonogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032535">
        <property type="term" value="P:regulation of cellular component size"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050767">
        <property type="term" value="P:regulation of neurogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031290">
        <property type="term" value="P:retinal ganglion cell axon guidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0022029">
        <property type="term" value="P:telencephalon cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd00054">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="CDD" id="cd00110">
        <property type="entry name" value="LamG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000045">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000053">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000054">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000062">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000063">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000099">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000289">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.200:FF:000013">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000002">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000004">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000039">
        <property type="entry name" value="slit homolog 2 protein isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.200">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000483">
        <property type="entry name" value="Cys-rich_flank_reg_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006207">
        <property type="entry name" value="Cys_knot_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001881">
        <property type="entry name" value="EGF-like_Ca-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000152">
        <property type="entry name" value="EGF-type_Asp/Asn_hydroxyl_site"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018097">
        <property type="entry name" value="EGF_Ca-bd_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003645">
        <property type="entry name" value="Fol_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001791">
        <property type="entry name" value="Laminin_G"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001611">
        <property type="entry name" value="Leu-rich_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003591">
        <property type="entry name" value="Leu-rich_rpt_typical-subtyp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000372">
        <property type="entry name" value="LRRNT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051355">
        <property type="entry name" value="Notch/Slit_guidance"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45836">
        <property type="entry name" value="SLIT HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45836:SF2">
        <property type="entry name" value="SLIT HOMOLOG 2 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00008">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00054">
        <property type="entry name" value="Laminin_G_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00560">
        <property type="entry name" value="LRR_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13855">
        <property type="entry name" value="LRR_8"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01463">
        <property type="entry name" value="LRRCT"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01462">
        <property type="entry name" value="LRRNT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00041">
        <property type="entry name" value="CT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="SMART" id="SM00179">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SMART" id="SM00274">
        <property type="entry name" value="FOLN"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00282">
        <property type="entry name" value="LamG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00368">
        <property type="entry name" value="LRR_RI"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SMART" id="SM00365">
        <property type="entry name" value="LRR_SD22"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SMART" id="SM00369">
        <property type="entry name" value="LRR_TYP"/>
        <property type="match status" value="12"/>
    </dbReference>
    <dbReference type="SMART" id="SM00082">
        <property type="entry name" value="LRRCT"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00013">
        <property type="entry name" value="LRRNT"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52058">
        <property type="entry name" value="L domain-like"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00010">
        <property type="entry name" value="ASX_HYDROXYL"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01185">
        <property type="entry name" value="CTCK_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01225">
        <property type="entry name" value="CTCK_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01186">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01187">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50025">
        <property type="entry name" value="LAM_G_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51450">
        <property type="entry name" value="LRR"/>
        <property type="match status" value="5"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0145">Chemotaxis</keyword>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="10" id="KW-0221">Differentiation</keyword>
    <keyword evidence="12 15" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5 15" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7" id="KW-0358">Heparin-binding</keyword>
    <keyword evidence="6" id="KW-0433">Leucine-rich repeat</keyword>
    <keyword evidence="11" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="8" id="KW-0732">Signal</keyword>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="743"/>
            <end position="778"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="780"/>
            <end position="819"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="821"/>
            <end position="857"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="859"/>
            <end position="897"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="899"/>
            <end position="935"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="944"/>
            <end position="980"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin G" evidence="17">
        <location>
            <begin position="983"/>
            <end position="1156"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="1160"/>
            <end position="1191"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="1194"/>
            <end position="1230"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="1235"/>
            <end position="1271"/>
        </location>
    </feature>
    <feature type="domain" description="CTCK" evidence="16">
        <location>
            <begin position="1276"/>
            <end position="1351"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="768"/>
            <end position="777"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="809"/>
            <end position="818"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="847"/>
            <end position="856"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="887"/>
            <end position="896"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="925"/>
            <end position="934"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="970"/>
            <end position="979"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="1181"/>
            <end position="1190"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="1198"/>
            <end position="1208"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="1220"/>
            <end position="1229"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="1239"/>
            <end position="1249"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="1261"/>
            <end position="1270"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022500"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022614"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022674"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00074955"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS01225"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50025"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016206.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1352" mass="149735" checksum="E1305F2A8B13661F" modified="2019-05-08" version="1">AVDIKNLQLDYNQISCIEDGAFRALRDLEVLTLNNNNITRLSVASFNHMPKLRTFRLHSNNLYCDCHLAWLSDWLRQRPRVGLYTQCMGPSHLRGHNVAEVQKREFVCSGHQSFMAPSCSVLHCPTACTCSNNIVDCRGKGLTEIPTNLPETITEIRLEQNSIKVIPPGAFSPYKKLRRIDLSNNQISETAPDAFQGLRKISNIFGIQLLNANKINCLRVDAFQDLHNLNLLSLYDNKLQTIAKGTFSPLRSIQTLHLAQNPFICDCHLKWLADYLHANPIETSGARCTSPRRLANKRIGQIKSKKFRCSGTEDYRSKLSGDCFADLACPEKCRCEGTTVDCSNQKLNKIPEHVPQYTAELRLNNNEFTVLEATGIFKKLPQLRKINLSNNKITDIEEGAFEGASGVNELLLTSNRLETIRHKMFKGLESLKTLMLRSNRISCIGNDSFTGLSSVRLLSLYDNQITTVAPGAFDTLHSLSTLNLLANPFNCNCHLAWLGDWLRKKRIVTGNPRCQKPYFLKEIPIQDVAIQDFTCDDGNDDNSCSPLSRCPAECTCLDTVVRCSNKGLKVLPKGIPKDVTELYLDGNQFTLVPKELSNYKHLTLIDLSNNRISTLSNQSFSNMTQLLTLILSYNRLRCIPARTFDGLKSLRLLSLHGNDISVVPEGAFNDLSALSHLAIGANPLYCDCNMQWLSDWVKSEYKEPGIARCAGPGEMADKLLLTTPSKKFTCQGPVDVNILAKCSPCLSNPCKNDGTCNNDPVDFYTCTCPYGFKGQDCDIPIHACISNPCQHGGTCHLKEGEKDGFWCTCADGFEGENCEVNVDDCEDNDCENNSTCVDGINNYTCLCPPEYTGELCEEKLDFCAQDLNPCQHDSKCILTLKGYKCDCTPGYVGEHCDMDYDDCQDNKCKNGAQCTDAVNGYTCICPEGYSGLFCEFSPPVVLPRTSPCDNYECQNGAQCIVRVNEPICQCLSGYQGDKCEKLVSVNFVSKESYLQIPSSKVRPQTNITLQIATDEDSGILLYKGDRDHIAMELYRGRVRVSYDTGSYPASAIYSVETINDGNFHIVELLAMDQILSLSVDGGSPKIITNLSKQSTLNVDSPLYVGGMPVKNNVAALRQSPGQNGSSFHGCIRNLYINSELQDFRNVPLQVGILPGCEPCHKKVCVHGTCHAMSQSGFTCECEGGWTGPLCDQQTNDPCLGNKCVHGTCLPINAFSYSCKCLQGHGGVLCDEEEELFNPCQSIKCKHGKCRLSGLGKPYCECSSGYTGDSCDKEISCRGERIRDYYQKQQGYAACQTTKKVSRLECKGGCSTGQCCGPLRSKRRKYSFECTDGSSFVDEVEKVVKCGCTKCPS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMP4</accession>
    <name>A0A452HMP4_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="12">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Mitochondrion membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 10">Belongs to the mitochondrial carrier (TC 2.A.29) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMP4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018569.1">
        <property type="protein sequence ID" value="ENSGAGP00000016267.1"/>
        <property type="gene ID" value="ENSGAGG00000012073.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031966">
        <property type="term" value="C:mitochondrial membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015227">
        <property type="term" value="F:O-acyl-L-carnitine transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1902603">
        <property type="term" value="P:carnitine transmembrane transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006839">
        <property type="term" value="P:mitochondrial transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.50.40.10:FF:000051">
        <property type="entry name" value="Mitochondrial carnitine/acylcarnitine carrier protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.50.40.10">
        <property type="entry name" value="Mitochondrial carrier domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023395">
        <property type="entry name" value="MCP_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018108">
        <property type="entry name" value="MCP_transmembrane"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050567">
        <property type="entry name" value="Mitochondrial_Carrier"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45624">
        <property type="entry name" value="MITOCHONDRIAL BASIC AMINO ACIDS TRANSPORTER-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45624:SF56">
        <property type="entry name" value="MITOCHONDRIAL CARNITINE_ACYLCARNITINE CARRIER PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00153">
        <property type="entry name" value="Mito_carr"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103506">
        <property type="entry name" value="Mitochondrial carrier"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50920">
        <property type="entry name" value="SOLCAR"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 9 11" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="4 9 10" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 11" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="3 10" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="53"/>
            <end position="73"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="85"/>
            <end position="107"/>
        </location>
    </feature>
    <feature type="repeat" description="Solcar" evidence="9">
        <location>
            <begin position="1"/>
            <end position="77"/>
        </location>
    </feature>
    <feature type="repeat" description="Solcar" evidence="9">
        <location>
            <begin position="86"/>
            <end position="174"/>
        </location>
    </feature>
    <feature type="repeat" description="Solcar" evidence="9">
        <location>
            <begin position="185"/>
            <end position="271"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004225"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006375"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00282"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="RuleBase" id="RU000488"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016267.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="279" mass="30524" checksum="0BEE0CB7FEB606F7" modified="2019-05-08" version="1">MLAYWGGGVGVFFVRLQTQPKPQAGQPPLYSGTFDCFRQTLVREGVRGLYKGMAAPIVGVTPMFAVCFFGFGLGKKLQQKNPDDILTYPQLFAAGMLSGVFTTAIMAPGERIKCLLQIQAASGEIKYGGPMDCAKQLYREAGIRGIYKGTLLTLMRDVPASGMYFMTYEWLKNILTPEGKSVSDLSVPRILFAGGMAGIFNWAVAIPPDVLKSRFQTAASGKYPNGFRDVLRELIREEGVSSLYKGFTAVMIRAFPANAACFLGFEVAMKFLNWIAPSL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMP9</accession>
    <name>A0A452HMP9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">tRNA (guanosine(18)-2'-O)-methyltransferase TARBP1</fullName>
            <ecNumber evidence="10">2.1.1.34</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="12">TAR RNA-binding protein 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">S-adenosyl-L-methionine-dependent 2'-O-ribose methyltransferase that catalyzes the formation of 2'-O-methylguanosine at position 18 (Gm18) in a subset of tRNA. Selectively mediates Gm18 methylation of tRNAGln-TTG/CTG and tRNASer-TGA/GCT. Gm18 modification can enhance the stability of modified tRNAs.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="8">
            <text>guanosine(18) in tRNA + S-adenosyl-L-methionine = 2'-O-methylguanosine(18) in tRNA + S-adenosyl-L-homocysteine + H(+)</text>
            <dbReference type="Rhea" id="RHEA:20077"/>
            <dbReference type="Rhea" id="RHEA-COMP:10190"/>
            <dbReference type="Rhea" id="RHEA-COMP:10192"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57856"/>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
            <dbReference type="ChEBI" id="CHEBI:74269"/>
            <dbReference type="ChEBI" id="CHEBI:74445"/>
            <dbReference type="EC" id="2.1.1.34"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="8">
            <dbReference type="Rhea" id="RHEA:20078"/>
        </physiologicalReaction>
    </comment>
    <comment type="subunit">
        <text evidence="2">Monomer and homodimer.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the class IV-like SAM-binding methyltransferase superfamily. RNA methyltransferase TrmH family.</text>
    </comment>
    <dbReference type="EC" id="2.1.1.34" evidence="10"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018577.1">
        <property type="protein sequence ID" value="ENSGAGP00000016273.1"/>
        <property type="gene ID" value="ENSGAGG00000012102.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0141100">
        <property type="term" value="F:tRNA (guanine(18)-2'-O)-methyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030488">
        <property type="term" value="P:tRNA methylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd18091">
        <property type="entry name" value="SpoU-like_TRM3-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.1280.10:FF:000010">
        <property type="entry name" value="probable methyltransferase TARBP1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.1280.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029028">
        <property type="entry name" value="Alpha/beta_knot_MTases"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001537">
        <property type="entry name" value="SpoU_MeTrfase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025806">
        <property type="entry name" value="TARBP1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056921">
        <property type="entry name" value="TARBP1_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045330">
        <property type="entry name" value="TRM3/TARBP1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044748">
        <property type="entry name" value="Trm3/TARBP1_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029026">
        <property type="entry name" value="tRNA_m1G_MTases_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12029:SF11">
        <property type="entry name" value="METHYLTRANSFERASE TARBP1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12029">
        <property type="entry name" value="RNA METHYLTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00588">
        <property type="entry name" value="SpoU_methylase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25050">
        <property type="entry name" value="TARBP1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF75217">
        <property type="entry name" value="alpha/beta knot"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51624">
        <property type="entry name" value="SAM_MT_TRMH_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0007">Acetylation</keyword>
    <keyword evidence="3" id="KW-0489">Methyltransferase</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="5" id="KW-0949">S-adenosyl-L-methionine</keyword>
    <keyword evidence="4" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="TARBP1" evidence="14">
        <location>
            <begin position="198"/>
            <end position="302"/>
        </location>
    </feature>
    <feature type="domain" description="tRNA/rRNA methyltransferase SpoU type" evidence="13">
        <location>
            <begin position="1369"/>
            <end position="1510"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007228"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00011407"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022603"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022691"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00093266"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00093361"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00093594"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00093636"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00093656"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="Pfam" id="PF00588"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="Pfam" id="PF25050"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016273.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1525" mass="171836" checksum="697E297D94E86CFD" modified="2019-05-08" version="1">MELVLADALLGLAQSPDPCALLRAVCGGDGAAPLSVERVETLRFLLQWLEAGGAGGEALRRAAWEVALGRCWPLLRGGGGAAGPERRRLLRAACGALRLCARLCGAQLAARLAGEALAALRAEPPGAGLGAEAAVEALVAVAPELXXVWAGLLPGRGEERRVSRTLLALTALADLLFPGLEGDGGGEGPALDARLCGGFWRVLQAGLTDGDGLCRKRARYLLKRAVEVSDQLRTDCTCPPDDGDEPSLFWWSVEKNDKLLQFWENYILIMETLEGNQIHVIKPVLPKLNSLYEYALSEKGCWLFHPSWHMCIYKRMFESENKTLTREGVAQFLEIYETKRLPNSPGFSEFIIGPFMDALSESSLYGRSPGQLIGACPPLGMKLQKFLSTYLTLLSEEERRSFLLKFIQKMTSRHWCAVPILFLSMALACIPSCKALGTEGLHSLRDVLQCTMITHQILLRGAAQCFLLQTAMHLTDVEKVSLPEISSFLMSLRPEESLSRGTILWMEVRIETVTGFVLIKMFCNTNRENCFMPDWFEAKLIATMILLAADVEEMKNKHRNTERVELKSFLHPLLDVLLKLGTNAYIPTLKTDKSLQLLLKLLQTCLLKQYSPQDDVVLSFLSNSVMTAAESILEFILRRLTASELNTVSDLDRCNLYLALLSELVNLCSNAGWKKGPSIWNFISSLINASVKNLQETGNEQEPKLGEQIQKAVSMASLTVVCEIIDQKPELQLKSLHAVETLKGFISSVQLNQVLKKPSYIEEKNRYVHYFLCYSLFEEATSSQGWGKIVASYVHDQWVCLHFILNKYHKCIPLSNDETLELSLLAVQRPIRTLQSALEALTLLPSDQVLPVFHCMKVLVPKLLCSSESLCIDAFDLAWKVISSLSNTQLIFWSNLKAFVQFVFDTEVLAIAASSKGQAYSKIKEIIVKLVEMSTARTGVFNTVISYCCQSWLFPTADDTGALENPFSNAGNYSELITEACVFGMVFRRDQRLIQDVHAFVENLGDECAANVVTERDDHYVRVCAIKFLCWLDGSNTLHKMFMEDLVIKLLDKDELVSKSRTRYYVNSLQHRMKNRLWQTLLVIFPKLDQNFLTGTLDKIFQAGFGNNQASIKYLIEWIIILILQRYPQCLEKFWNCFSYGEEKLKTSICTFLSVLSHFDIILENTSEKKPILKKALLVVLQWCFNHNFSVRLYALVALKKIWGMCKALYVEEFEALTPVIESSLHQVENMHGAGNARKNWQRIQDHFFFATFHPLKDYSLETIFYTLPRLSELVEDEWIPLCKFTRYTDISLPAAFHWLHSRTELLNLKPSDWSQQDMGSDLVEMDNQTEWTDVQKKIIPWKNSSPDLDLELLFQDRAAKLGKSNSRLIVVASLIDKPTNLGGLCRTSEIFGASALVVGSLHYINDKQFQHLSVSAEQWLPLVEVKPSQLVNYLQQKKTEGYTIIGVEQTAKSSDLTEYCFPEKSLLLLGNEHEGIPANLIQHLDVCVEIPQQGIIRSLNVHVSGALLIWEYTRQQLIKQKQLK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMS0</accession>
    <name>A0A452HMS0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Slit homolog 2 protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="15">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018544.1">
        <property type="protein sequence ID" value="ENSGAGP00000016244.1"/>
        <property type="gene ID" value="ENSGAGG00000011388.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048495">
        <property type="term" value="F:Roundabout binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072359">
        <property type="term" value="P:circulatory system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060560">
        <property type="term" value="P:developmental growth involved in morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060562">
        <property type="term" value="P:epithelial tube morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001822">
        <property type="term" value="P:kidney development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050919">
        <property type="term" value="P:negative chemotaxis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030308">
        <property type="term" value="P:negative regulation of cell growth"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010605">
        <property type="term" value="P:negative regulation of macromolecule metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010977">
        <property type="term" value="P:negative regulation of neuron projection development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0021772">
        <property type="term" value="P:olfactory bulb development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050770">
        <property type="term" value="P:regulation of axonogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032535">
        <property type="term" value="P:regulation of cellular component size"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050767">
        <property type="term" value="P:regulation of neurogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031290">
        <property type="term" value="P:retinal ganglion cell axon guidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0022029">
        <property type="term" value="P:telencephalon cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd00054">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="CDD" id="cd00110">
        <property type="entry name" value="LamG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000045">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000053">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000054">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000062">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000063">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000099">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000289">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.200:FF:000013">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000002">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000039">
        <property type="entry name" value="slit homolog 2 protein isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.200">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000483">
        <property type="entry name" value="Cys-rich_flank_reg_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006207">
        <property type="entry name" value="Cys_knot_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001881">
        <property type="entry name" value="EGF-like_Ca-bd_dom"/>
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        <property type="entry name" value="EGF-type_Asp/Asn_hydroxyl_site"/>
    </dbReference>
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        <property type="entry name" value="EGF_Ca-bd_CS"/>
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        <property type="entry name" value="Fol_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001791">
        <property type="entry name" value="Laminin_G"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001611">
        <property type="entry name" value="Leu-rich_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003591">
        <property type="entry name" value="Leu-rich_rpt_typical-subtyp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000372">
        <property type="entry name" value="LRRNT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051355">
        <property type="entry name" value="Notch/Slit_guidance"/>
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    <dbReference type="PANTHER" id="PTHR45836">
        <property type="entry name" value="SLIT HOMOLOG"/>
        <property type="match status" value="1"/>
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    <dbReference type="PANTHER" id="PTHR45836:SF2">
        <property type="entry name" value="SLIT HOMOLOG 2 PROTEIN"/>
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    </dbReference>
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        <property type="entry name" value="EGF"/>
        <property type="match status" value="6"/>
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        <property type="entry name" value="Laminin_G_1"/>
        <property type="match status" value="1"/>
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    <dbReference type="Pfam" id="PF13855">
        <property type="entry name" value="LRR_8"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01463">
        <property type="entry name" value="LRRCT"/>
        <property type="match status" value="4"/>
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        <property type="entry name" value="LRRNT"/>
        <property type="match status" value="2"/>
    </dbReference>
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        <property type="entry name" value="CT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="SMART" id="SM00179">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="7"/>
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        <property type="entry name" value="FOLN"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00282">
        <property type="entry name" value="LamG"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="LRR_RI"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00365">
        <property type="entry name" value="LRR_SD22"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SMART" id="SM00369">
        <property type="entry name" value="LRR_TYP"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="SMART" id="SM00082">
        <property type="entry name" value="LRRCT"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00013">
        <property type="entry name" value="LRRNT"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52058">
        <property type="entry name" value="L domain-like"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00010">
        <property type="entry name" value="ASX_HYDROXYL"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01185">
        <property type="entry name" value="CTCK_1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="CTCK_2"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="9"/>
    </dbReference>
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        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="7"/>
    </dbReference>
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        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="9"/>
    </dbReference>
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        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50025">
        <property type="entry name" value="LAM_G_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51450">
        <property type="entry name" value="LRR"/>
        <property type="match status" value="5"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0145">Chemotaxis</keyword>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="10" id="KW-0221">Differentiation</keyword>
    <keyword evidence="12 15" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5 15" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7" id="KW-0358">Heparin-binding</keyword>
    <keyword evidence="6" id="KW-0433">Leucine-rich repeat</keyword>
    <keyword evidence="11" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="8" id="KW-0732">Signal</keyword>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="717"/>
            <end position="752"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="754"/>
            <end position="793"/>
        </location>
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    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="795"/>
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        </location>
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    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="833"/>
            <end position="871"/>
        </location>
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    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="873"/>
            <end position="909"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="918"/>
            <end position="954"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin G" evidence="17">
        <location>
            <begin position="957"/>
            <end position="1130"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="1134"/>
            <end position="1165"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="1168"/>
            <end position="1204"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="1209"/>
            <end position="1245"/>
        </location>
    </feature>
    <feature type="domain" description="CTCK" evidence="16">
        <location>
            <begin position="1250"/>
            <end position="1325"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="742"/>
            <end position="751"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="783"/>
            <end position="792"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="821"/>
            <end position="830"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="861"/>
            <end position="870"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="899"/>
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        </location>
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    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="944"/>
            <end position="953"/>
        </location>
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    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="1155"/>
            <end position="1164"/>
        </location>
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        </location>
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        </location>
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        </location>
    </feature>
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        <location>
            <begin position="1235"/>
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        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022500"/>
        </source>
    </evidence>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00074955"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
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    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS01225"/>
        </source>
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        <source>
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        </source>
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        <source>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016244.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
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        </source>
    </evidence>
    <sequence length="1326" mass="147303" checksum="C707BBE87C2E0692" modified="2019-05-08" version="1">FNRQLDYNQISCIEDGAFRALRDLEVLTLNNNNITRLSVASFNHMPKLRTFRLHSNNLYCDCHLAWLSDWLRQRPRVGLYTQCMGPSHLRGHNVAEVQKREFVCSGHQSFMAPSCSVLHCPTACTCSNNIVDCRGKGLTEIPTNLPETITEIRLEQNSIKVIPPGAFSPYKKLRRIDLSNNQISETAPDAFQGLRKISNIFGIQFLKILLNANKINCLRVDAFQDLHNLNLLSLYDNKLQTIAKGTFSPLRSIQTLHLAQNPFICDCHLKWLADYLHANPIETSGARCTSPRRLANKRIGQIKSKKFRCSAKEQYFIPGTEDYRSKLSGDCFADLACPEKCRCEGTTVDCSNQKLNKIPEHVPQYTAELRLNNNEFTVLEATGIFKKLPQLRKINLSNNKITDIEEGAFEGASGVNELLLTSNRLETIRHKMFKGLESLKTLMLRSNRISCIGNDSFTGLSSVRLLSLYDNQITTVAPGAFDTLHSLSTLNLLANPFNCNCHLAWLGDWLRKKRIVTGNPRCQKPYFLKEIPIQDVAIQDFTCDDGNDDNSCSPLSRCPAECTCLDTVVRCSNKGLKVLPKGIPKDVTELMMLPNLDLSNNRISTLSNQSFSNMTQLLTFAFSFQEFSADHRFVFVIQSKHIKLLCFFPPSAIGANPLYCDCNMQWLSDWVKSEYKEPGIARCAGPGEMADKLLLTTPSKKFTCQGPVDVNILAKCSPCLSNPCKNDGTCNNDPVDFYTCTCPYGFKGQDCDIPIHACISNPCQHGGTCHLKEGEKDGFWCTCADGFEGENCEVNVDDCEDNDCENNSTCVDGINNYTCLCPPEYTGELCEEKLDFCAQDLNPCQHDSKCILTLKGYKCDCTPGYVGEHCDMDYDDCQDNKCKNGAQCTDAVNGYTCICPEGYSGLFCEFSPPVVLPRTSPCDNYECQNGAQCIVRVNEPICQCLSGYQGDKCEKLVSVNFVSKESYLQIPSSKVRPQTNITLQIATDEDSGILLYKGDRDHIAMELYRGRVRVSYDTGSYPASAIYSVETINDGNFHIVELLAMDQILSLSVDGGSPKIITNLSKQSTLNVDSPLYVGGMPVKNNVAALRQSPGQNGSSFHGCIRNLYINSELQDFRNVPLQVGILPGCEPCHKKVCVHGTCHAMSQSGFTCECEGGWTGPLCDQQTNDPCLGNKCVHGTCLPINAFSYSCKCLQGHGGVLCDEEEELFNPCQSIKCKHGKCRLSGLGKPYCECSSGYTGDSCDKEISCRGERIRDYYQKQQGYAACQTTKKVSRLECKGGCSTGQCCGPLRSKRRKYSFECTDGSSFVDEVEKVVKCGCTKCPS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMU1</accession>
    <name>A0A452HMU1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">SAM domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the BicC family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMU1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018567.1">
        <property type="protein sequence ID" value="ENSGAGP00000016265.1"/>
        <property type="gene ID" value="ENSGAGG00000012189.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="CDD" id="cd22421">
        <property type="entry name" value="KH-I_BICC1_rpt2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd22422">
        <property type="entry name" value="KH-I_BICC1_rpt3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd09520">
        <property type="entry name" value="SAM_BICC1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.50:FF:000025">
        <property type="entry name" value="Ankyrin repeat and sterile alpha motif domain-containing 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.310.270:FF:000001">
        <property type="entry name" value="BicC family RNA binding protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.310.270:FF:000002">
        <property type="entry name" value="BicC family RNA binding protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1370.10">
        <property type="entry name" value="K Homology domain, type 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.50">
        <property type="entry name" value="Transcription Factor, Ets-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR054727">
        <property type="entry name" value="BICC1_KH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047549">
        <property type="entry name" value="BICC1_KH-I_rpt1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047554">
        <property type="entry name" value="BICC1_KH-I_rpt2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047553">
        <property type="entry name" value="BICC1_KH-I_rpt3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037974">
        <property type="entry name" value="BICC1_SAM_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004087">
        <property type="entry name" value="KH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004088">
        <property type="entry name" value="KH_dom_type_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036612">
        <property type="entry name" value="KH_dom_type_1_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001660">
        <property type="entry name" value="SAM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013761">
        <property type="entry name" value="SAM/pointed_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10627:SF78">
        <property type="entry name" value="PROTEIN BICAUDAL C HOMOLOG 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10627">
        <property type="entry name" value="SCP160"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00013">
        <property type="entry name" value="KH_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22985">
        <property type="entry name" value="KH_BICC1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24234">
        <property type="entry name" value="KH_BICC1_1st"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00536">
        <property type="entry name" value="SAM_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00322">
        <property type="entry name" value="KH"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00454">
        <property type="entry name" value="SAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54791">
        <property type="entry name" value="Eukaryotic type KH-domain (KH-domain type I)"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47769">
        <property type="entry name" value="SAM/Pointed domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50084">
        <property type="entry name" value="KH_TYPE_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50105">
        <property type="entry name" value="SAM_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="4 5" id="KW-0694">RNA-binding</keyword>
    <feature type="domain" description="SAM" evidence="7">
        <location>
            <begin position="742"/>
            <end position="805"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="459"/>
            <end position="519"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="548"/>
            <end position="568"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="649"/>
            <end position="714"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="459"/>
            <end position="488"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="503"/>
            <end position="519"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="559"/>
            <end position="568"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="661"/>
            <end position="682"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007662"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00117"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50105"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016265.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="824" mass="89773" checksum="2E8556DE74E9F85C" modified="2019-05-08" version="1">MEETNTQIAWPSKLKIGAKSKKDPHIKVSGKKENVKEAKEKIMSVLDTKSNRVTLKMDVSHTEHSHVIGKGGNNIKKVMEETGCHIHFPDSNRNNQAEKSNQVSIAGQPAGVESARVRIRELLPLVLMFELPIAGILQPIPDPNSPTILHISQTYNISVSFKQRSRMYGATVIVRGSQNNTSAVKEGTAMLLEHLAGSLASAIPVSTQLDIAAQHHLFMMGRNGSNIKHIMQRTGAQIHFPDPSNPQKKSTVYLQGTIESVCLARQYLMGCLPLVLMFDMKEEIEIEPQIITQLMEQLDVFISIKPKPKQPSKSVIVKSVERNALNMYETRKRLLGLESNGVTIAANPSTVSCHVGLSCPGLDILPSAGLGLTGLGFSAIPHLMIPSTAQAALTSILLSGVPNYSQNTPSPPPGLTPVDIHVNGMQSENKKGSASLNGHVKTSNIKYAALSASSLGENVLSTNHSDSSRQTNSHSTSEQVTTKSNTAEGCNDAFVEVGMPRSPSHSANASDLKQMMSSSKQSCTKRQTVELLQGTKNSHLHTTERLLSDTELSASENPVADKKAPGSERAAERAAAAQQNSERVRLAPQSSYVNMQAFDYEQKKLLATKAMLKKPVVTEVRTPTNTWSGLGFSKSMPAETIKELRRANHVSYKPSMTTTYEGSSMSLSPSSSREHLGSGSESDNWRDRNGIGPSAHSEFVSSVGSPKRKQNKSTEHYLSSSNYMDCISSLTGSNGCNLNSLFKGSDLPELFSKLGLGKYTDVFQQQEIDLQTFLTLTDQDLKELGITTFGARRKMLLAISDVLLWIDEGFSYNEGNHKVPLFLQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMU3</accession>
    <name>A0A452HMU3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">SAM domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the BicC family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMU3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018587.1">
        <property type="protein sequence ID" value="ENSGAGP00000016282.1"/>
        <property type="gene ID" value="ENSGAGG00000012189.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="CDD" id="cd22421">
        <property type="entry name" value="KH-I_BICC1_rpt2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd22422">
        <property type="entry name" value="KH-I_BICC1_rpt3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd09520">
        <property type="entry name" value="SAM_BICC1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.50:FF:000025">
        <property type="entry name" value="Ankyrin repeat and sterile alpha motif domain-containing 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.310.270:FF:000001">
        <property type="entry name" value="BicC family RNA binding protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.310.270:FF:000002">
        <property type="entry name" value="BicC family RNA binding protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.310.270">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1370.10">
        <property type="entry name" value="K Homology domain, type 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.50">
        <property type="entry name" value="Transcription Factor, Ets-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR054727">
        <property type="entry name" value="BICC1_KH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047549">
        <property type="entry name" value="BICC1_KH-I_rpt1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047554">
        <property type="entry name" value="BICC1_KH-I_rpt2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047553">
        <property type="entry name" value="BICC1_KH-I_rpt3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037974">
        <property type="entry name" value="BICC1_SAM_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004087">
        <property type="entry name" value="KH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004088">
        <property type="entry name" value="KH_dom_type_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036612">
        <property type="entry name" value="KH_dom_type_1_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001660">
        <property type="entry name" value="SAM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013761">
        <property type="entry name" value="SAM/pointed_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10627:SF78">
        <property type="entry name" value="PROTEIN BICAUDAL C HOMOLOG 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10627">
        <property type="entry name" value="SCP160"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00013">
        <property type="entry name" value="KH_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22985">
        <property type="entry name" value="KH_BICC1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24234">
        <property type="entry name" value="KH_BICC1_1st"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00536">
        <property type="entry name" value="SAM_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00322">
        <property type="entry name" value="KH"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00454">
        <property type="entry name" value="SAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54791">
        <property type="entry name" value="Eukaryotic type KH-domain (KH-domain type I)"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47769">
        <property type="entry name" value="SAM/Pointed domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50084">
        <property type="entry name" value="KH_TYPE_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50105">
        <property type="entry name" value="SAM_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="4 5" id="KW-0694">RNA-binding</keyword>
    <feature type="domain" description="SAM" evidence="7">
        <location>
            <begin position="803"/>
            <end position="866"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="520"/>
            <end position="580"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="609"/>
            <end position="628"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="710"/>
            <end position="776"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="520"/>
            <end position="549"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="564"/>
            <end position="580"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="722"/>
            <end position="743"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007662"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00117"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50105"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016282.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="904" mass="97819" checksum="4CD8157CE3D5DBC5" modified="2019-05-08" version="1">LDFKLPCAPIMEETNTQIAWPSKLKIGAKSKKDPHIKVSGKKENVKEAKEKIMSVLDTKSNRVTLKMDVSHTEHSHVIGKGGNNIKKVMEETGCHIHFPDSNRNNQAEKSNQVSIAGQPAGVESARVRIRELLPLVLMFELPIAGILQPIPDPNSPTILHISQTYNISVSFKQRSRMYGATVIVRGSQNNTSAVKEGTAMLLEHLAGSLASAIPVSTQLDIAAQHHLFMMGRNGSNIKHIMQRTGAQIHFPDPSNPQKKSTVYLQGTIESVCLARQYLMGCLPLVLMFDMKEEIEIEPQIITQLMEQLDVFISIKPKPKQPSKSVIVKSVERNALNMYETRKRLLGLESNGVTIAANPSTVSCHVGLSCPGLDILPSAGLGLTGLGLLGPTALSVNTSTAPNSLLNALNSSVSPLQSPSSGTPSPTLWATSLANTSATGFSAIPHLMIPSTAQAALTSILLSGVPNYSQNTPSPPPGLTPVDIHVNGMQSENKKGSASLNGHTSNIKYAALSASSLGENVLSTNHSDSSRQTNSHSTSEQVTTKSNTAEGCNDAFVEVGMPRSPSHSANASDLKQMMSSSKQSCTKRQTVELLQGTKNSHLHTTERLLSDTELSASENPVADKKAPGSERAAERAAAAQQNSERVRLAPQSSYVNMQAFDYEQKKLLATKAMLKKPVVTEVRTPTNTWSGLGFSKSMPAETIKELRRANHVSYKPSMTTTYEGSSMSLSPSSSREHLGSGSESDNWRDRNGIGPSAHSEFVSSVGSPKRKQNKSTEHYLSSSNYMDCISSLTGSNGCNLNSLFKGSDLPELFSKLGLGKYTDVFQQQEIDLQTFLTLTDQDLKELGITTFGARRKMLLAISELNKNRRKLFDPSNVRASFLEGGASGRLPRQYHSDIASVSGRW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMU6</accession>
    <name>A0A452HMU6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">tRNA (guanosine(18)-2'-O)-methyltransferase TARBP1</fullName>
            <ecNumber evidence="10">2.1.1.34</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="12">TAR RNA-binding protein 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">S-adenosyl-L-methionine-dependent 2'-O-ribose methyltransferase that catalyzes the formation of 2'-O-methylguanosine at position 18 (Gm18) in a subset of tRNA. Selectively mediates Gm18 methylation of tRNAGln-TTG/CTG and tRNASer-TGA/GCT. Gm18 modification can enhance the stability of modified tRNAs.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="8">
            <text>guanosine(18) in tRNA + S-adenosyl-L-methionine = 2'-O-methylguanosine(18) in tRNA + S-adenosyl-L-homocysteine + H(+)</text>
            <dbReference type="Rhea" id="RHEA:20077"/>
            <dbReference type="Rhea" id="RHEA-COMP:10190"/>
            <dbReference type="Rhea" id="RHEA-COMP:10192"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57856"/>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
            <dbReference type="ChEBI" id="CHEBI:74269"/>
            <dbReference type="ChEBI" id="CHEBI:74445"/>
            <dbReference type="EC" id="2.1.1.34"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="8">
            <dbReference type="Rhea" id="RHEA:20078"/>
        </physiologicalReaction>
    </comment>
    <comment type="subunit">
        <text evidence="2">Monomer and homodimer.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the class IV-like SAM-binding methyltransferase superfamily. RNA methyltransferase TrmH family.</text>
    </comment>
    <dbReference type="EC" id="2.1.1.34" evidence="10"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018580.1">
        <property type="protein sequence ID" value="ENSGAGP00000016276.1"/>
        <property type="gene ID" value="ENSGAGG00000012102.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0141100">
        <property type="term" value="F:tRNA (guanine(18)-2'-O)-methyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030488">
        <property type="term" value="P:tRNA methylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd18091">
        <property type="entry name" value="SpoU-like_TRM3-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.1280.10:FF:000010">
        <property type="entry name" value="probable methyltransferase TARBP1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.1280.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029028">
        <property type="entry name" value="Alpha/beta_knot_MTases"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001537">
        <property type="entry name" value="SpoU_MeTrfase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025806">
        <property type="entry name" value="TARBP1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056921">
        <property type="entry name" value="TARBP1_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045330">
        <property type="entry name" value="TRM3/TARBP1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044748">
        <property type="entry name" value="Trm3/TARBP1_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029026">
        <property type="entry name" value="tRNA_m1G_MTases_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12029:SF11">
        <property type="entry name" value="METHYLTRANSFERASE TARBP1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12029">
        <property type="entry name" value="RNA METHYLTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00588">
        <property type="entry name" value="SpoU_methylase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25050">
        <property type="entry name" value="TARBP1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF75217">
        <property type="entry name" value="alpha/beta knot"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51624">
        <property type="entry name" value="SAM_MT_TRMH_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0007">Acetylation</keyword>
    <keyword evidence="3" id="KW-0489">Methyltransferase</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="5" id="KW-0949">S-adenosyl-L-methionine</keyword>
    <keyword evidence="4" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="TARBP1" evidence="14">
        <location>
            <begin position="198"/>
            <end position="302"/>
        </location>
    </feature>
    <feature type="domain" description="tRNA/rRNA methyltransferase SpoU type" evidence="13">
        <location>
            <begin position="1324"/>
            <end position="1465"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007228"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00011407"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022603"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022691"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00093266"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00093361"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00093594"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00093636"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00093656"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="Pfam" id="PF00588"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="Pfam" id="PF25050"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016276.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1480" mass="167138" checksum="78968FF193C4D7CC" modified="2019-05-08" version="1">MELVLADALLGLAQSPDPCALLRAVCGGDGAAPLSVERVETLRFLLQWLEAGGAGGEALRRAAWEVALGRCWPLLRGGGGAAGPERRRLLRAACGALRLCARLCGAQLAARLAGEALAALRAEPPGAGLGAEAAVEALVAVAPELXXVWAGLLPGRGEERRVSRTLLALTALADLLFPGLEGDGGGEGPALDARLCGGFWRVLQAGLTDGDGLCRKRARYLLKRAVEVSDQLRTDCTCPPDDGDEPSLFWWSVEKNDKLLQFWENYILIMETLEGNQIHVIKPVLPKLNSLYEYALSEKGCWLFHPSWHMCIYKRMFESENKTLTREGVAQFLEIYETKRLPNSPGFSEFIIGPFMDALSESSLYGRSPGQLIGACPPLGMKLQKFLSTYLTLLSEEERRSFLLKFIQKMTSRHWCAVPILFLSMALACIPSCKALGTEGLHSLRDVLQCTMITHQILLRGAAQCFLLQTAMHLTDVEKVSLPEISSFLMSLRPEESLSRGTILWMELCDWLYTHDRCFRKSINRHVEHQERSSLSVYVHTLVEDYLKAPASTRENCFMPDWFEAKLIATMILLAADVEEMKNKHSKKRNTERVELKSFLHPLLDVLLKLGTNAYIPTLKTDKSLQLLLKLLQTCLLKQYSPQDDLDRCNLYLALLSELVNLCSNAGWKKGPSIWNFISSLINASVKNLQETGNEQEPKLGEQIQKAVSMASLTVVCEIIDQKPELQLKSLHAVETLKGFISSVQLNQVLKKPSYIEEKNSLFEEATSSQGWGKIVASYVHDQWVCLHFILNKYHKCIPLSNDETLELSLLAVQRPIRTLQSALEALTLLPSDQVLPVFHCMKVLVPKLLCSSESLCIDAFDLAWKVISSLSNTQLIFWSNLKAFVQFVFDTEVLAIAASSKGQAYSKIKEIIVKLVEMSTARTGVFNTVISYCCQSWLFPTADDTGSTYLISELVDSHFSDNKMFYSTNRDDHYVRVCAIKFLCWLDGSNTLHKMFMEDLVIKLLDKDELVSKSRTRYYVNSLQHRMKNRLWQTLLVIFPKLDQNFLTGTLDKIFQAGFGNNQASIKYLIEWIIILILQRYPQCLEKFWNCFSYGEEKLKTSICTFLSVLSHFDIILENTSEKKPILKKALLVVLQWCFNHNFSVRLYALVALKKIWGMCKALYVEEFEALTPVIESSLHQVENMHGAGNARKNWQRIQDHFFFATFHPLKDYSLETIFYTLPRLSELVEDEWIPLCKFTRYTDISLPAAFHWLHSRTELLNLKPSDWSQQDMGSDLVEMDNQTEWTDVQKKIIPWKNSSPDLDLELLFQDRAAKLGKSNSRLIVVASLIDKPTNLGGLCRTSEIFGASALVVGSLHYINDKQFQHLSVSAEQWLPLVEVKPSQLVNYLQQKKTEGYTIIGVEQTAKSSDLTEYCFPEKSLLLLGNEHEGIPANLIQHLDVCVEIPQQGIIRSLNVHVSGALLIWEYTRQQLIKQKQLK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMU7</accession>
    <name>A0A452HMU7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Pleckstrin and Sec7 domain containing 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell projection</location>
            <location evidence="1">Ruffle membrane</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMU7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018602.1">
        <property type="protein sequence ID" value="ENSGAGP00000016296.1"/>
        <property type="gene ID" value="ENSGAGG00000012132.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032587">
        <property type="term" value="C:ruffle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005543">
        <property type="term" value="F:phospholipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032012">
        <property type="term" value="P:regulation of ARF protein signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd13295">
        <property type="entry name" value="PH_EFA6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00171">
        <property type="entry name" value="Sec7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.1000.11:FF:000004">
        <property type="entry name" value="PH and SEC7 domain-containing protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000054">
        <property type="entry name" value="PH and SEC7 domain-containing protein 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1000.11">
        <property type="entry name" value="Arf Nucleotide-binding Site Opener,domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041681">
        <property type="entry name" value="PH_9"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001605">
        <property type="entry name" value="PH_dom-spectrin-type"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023394">
        <property type="entry name" value="Sec7_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000904">
        <property type="entry name" value="Sec7_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035999">
        <property type="entry name" value="Sec7_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10663">
        <property type="entry name" value="GUANYL-NUCLEOTIDE EXCHANGE FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10663:SF329">
        <property type="entry name" value="PH AND SEC7 DOMAIN-CONTAINING PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15410">
        <property type="entry name" value="PH_9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01369">
        <property type="entry name" value="Sec7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00683">
        <property type="entry name" value="SPECTRINPH"/>
    </dbReference>
    <dbReference type="SMART" id="SM00233">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00222">
        <property type="entry name" value="Sec7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48425">
        <property type="entry name" value="Sec7 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50190">
        <property type="entry name" value="SEC7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="6" id="KW-0966">Cell projection</keyword>
    <keyword evidence="4 7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="SEC7" evidence="10">
        <location>
            <begin position="53"/>
            <end position="216"/>
        </location>
    </feature>
    <feature type="domain" description="PH" evidence="9">
        <location>
            <begin position="266"/>
            <end position="379"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="34"/>
            <end position="55"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="407"/>
            <end position="434"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004632"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50190"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016296.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="13">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="538" mass="60835" checksum="F87D08D056D76E27" modified="2019-05-08" version="1">RTAPAALTLGFPENTTFVSLIVFCSPRSDASLKDTLSDSDSDMGSTEQLDHGSTDTLANGCKADSDAAKRLAKRLYHLEGFKRCDVARQLGKNNEFSKLVAEEYLSFFDFTGLTLDKALRTFLKAFPLMGETQERERVLIHFSRRYCQCNPEESTSEDGIHTLTCALMLLNTDLHGHNIGKKMSCQQFITNLDGLNDGKDFAKDLLKTLYNSIKNEKLEWAIDEDELRKSLSELVDDKLGASAKKVTRIVDGSNPFLDIPQALNAITYKHGVLTRKTHADTDGKRTPRGRRGWKKFYAVLKGTILYLQKDEYKPDKDLSEVDLKNAIRVHHALATKASDYSKKSNVLKLKTADWRVFLFQAPSKEEMLSWILRINLVAAIFSAPAFPAAICSMKKFCRPLLPSSMTKLCQEEQLRSHENKMRQIADELTEHKLHPVEKSLKSKEAEEYRLKEHYLIFEKSRYETYINLLCMKIKVGTDDLEKIEASLFMVEADDISLRKTHSSPSLNCLTCPVSQEICGDPGTEARSSDSQCNASQEW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMU8</accession>
    <name>A0A452HMU8_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="14">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the IRF2BP family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMU8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018593.1">
        <property type="protein sequence ID" value="ENSGAGP00000016288.1"/>
        <property type="gene ID" value="ENSGAGG00000012209.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
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        <property type="project" value="UniProtKB-SubCell"/>
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        <property type="term" value="F:zinc ion binding"/>
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        <property type="project" value="UniProtKB-KW"/>
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        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
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        <property type="entry name" value="vRING-HC_IRF2BP2"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="interferon regulatory factor 2-binding protein 2"/>
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        <property type="entry name" value="Interferon regulatory factor 2-binding protein"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="I2BP1/2_C3HC4-RING_sf"/>
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    <dbReference type="InterPro" id="IPR058682">
        <property type="entry name" value="IRF-2BP1/2-like_M"/>
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        <property type="entry name" value="IRF-2BP1_2-like_Znf"/>
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    <dbReference type="InterPro" id="IPR057414">
        <property type="entry name" value="Zf-C3HC4_IRF-2BP1_2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10816:SF18">
        <property type="entry name" value="INTERFERON REGULATORY FACTOR 2-BINDING PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="MYELIN TRANSCRIPTION FACTOR 1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11261">
        <property type="entry name" value="IRF-2BP1_2"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="IRF-2BP1_2_M"/>
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        <property type="entry name" value="zf-C3HC4_IRF-2BP1_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57850">
        <property type="entry name" value="RING/U-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0678">Repressor</keyword>
    <keyword evidence="8" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="Interferon regulatory factor 2-binding protein 1/2-like zinc finger" evidence="11">
        <location>
            <begin position="11"/>
            <end position="62"/>
        </location>
    </feature>
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        <location>
            <begin position="207"/>
            <end position="280"/>
        </location>
    </feature>
    <feature type="domain" description="Interferon regulatory factor 2-binding protein 1/2-like C3HC4 zinc finger" evidence="12">
        <location>
            <begin position="442"/>
            <end position="513"/>
        </location>
    </feature>
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            <begin position="61"/>
            <end position="80"/>
        </location>
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    <feature type="region of interest" description="Disordered" evidence="10">
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            <begin position="93"/>
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        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="176"/>
            <end position="240"/>
        </location>
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            <begin position="283"/>
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    </feature>
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            <begin position="372"/>
            <end position="430"/>
        </location>
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        <location>
            <begin position="193"/>
            <end position="202"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="206"/>
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        </location>
    </feature>
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            <begin position="312"/>
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            <begin position="343"/>
            <end position="352"/>
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            <begin position="374"/>
            <end position="385"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00010802"/>
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            <dbReference type="ARBA" id="ARBA00022491"/>
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            <dbReference type="Pfam" id="PF11261"/>
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        <source>
            <dbReference type="Pfam" id="PF25454"/>
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            <dbReference type="Pfam" id="PF25457"/>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016288.1"/>
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    </evidence>
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        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="525" mass="55579" checksum="333F5566CBDBF179" modified="2019-05-08" version="1">MSSAAVAAASRRQSCYLCDLPRMPWAMIWDFTEPVCRGCVNYEGADRVEFVIDTARQLKRAHSFQEGRAPPPPHAKQPPLSAKEILVQQQQQQLNHTAVRQPPRLGAEYGGSRLVNGNLVPNGFPKPEEPPELNRQSPNPRRTHTVPPTLVPLMNGASLPAPISISSRATAASLAAISGAPPPPGELGHKRPGSVSSSSSSSGGAGEHESKEKQPRGSAEELGVEGKGRGGAEQEWLGKPKTVRDTLLALHQHGGHAVPFESKFKKEPGMAGSRLLGYESNGAVAKAGARAARKRKPSPEPEGEAGPPKINGESQPWLPTSSEGMKMSMTATSFMSPPPPTASPHSSRTTPPEAAQNGQSPMAALILVADNAGGNHSSKDANQVHSTTRRNSNSPPSPSSMNQRRLGPSREVPSQGTNSAGGLEPVHPASLPDSSLAASVPLCCTLCHERLEDTHFVQCPSVPSHKFCFPCSRQSIKQQGASGEVYCPSGEKCPLVGSNVPWAFMQGEIATILAGDVKVKKERDS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMV2</accession>
    <name>A0A452HMV2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">PAS domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018586.1">
        <property type="protein sequence ID" value="ENSGAGP00000016281.1"/>
        <property type="gene ID" value="ENSGAGG00000011530.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
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        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
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        <property type="term" value="F:transcription cis-regulatory region binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001222">
        <property type="term" value="F:transcription corepressor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032922">
        <property type="term" value="P:circadian regulation of gene expression"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043153">
        <property type="term" value="P:entrainment of circadian clock by photoperiod"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000122">
        <property type="term" value="P:negative regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00130">
        <property type="entry name" value="PAS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.450.20:FF:000013">
        <property type="entry name" value="Period circadian protein homolog 2"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Period circadian protein homolog 3"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="PAS domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000014">
        <property type="entry name" value="PAS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035965">
        <property type="entry name" value="PAS-like_dom_sf"/>
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        <property type="entry name" value="PER1-3_bHLH"/>
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        <property type="entry name" value="Per1-3_PAS-A"/>
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    <dbReference type="InterPro" id="IPR022728">
        <property type="entry name" value="Period_circadian-like_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050760">
        <property type="entry name" value="Period_circadian_regulator"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11269">
        <property type="entry name" value="PERIOD CIRCADIAN PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11269:SF8">
        <property type="entry name" value="PERIOD CIRCADIAN PROTEIN HOMOLOG 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23170">
        <property type="entry name" value="bHLH_PER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08447">
        <property type="entry name" value="PAS_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21353">
        <property type="entry name" value="Per3-like_PAS-A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12114">
        <property type="entry name" value="Period_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00091">
        <property type="entry name" value="PAS"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55785">
        <property type="entry name" value="PYP-like sensor domain (PAS domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50112">
        <property type="entry name" value="PAS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0090">Biological rhythms</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="7" id="KW-0804">Transcription</keyword>
    <keyword evidence="5" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="PAS" evidence="10">
        <location>
            <begin position="357"/>
            <end position="400"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="1"/>
            <end position="119"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="552"/>
            <end position="585"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="610"/>
            <end position="657"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="694"/>
            <end position="713"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="750"/>
            <end position="803"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="901"/>
            <end position="929"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="942"/>
            <end position="985"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="1030"/>
            <end position="1063"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="36"/>
            <end position="45"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="46"/>
            <end position="56"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="65"/>
            <end position="99"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="100"/>
            <end position="116"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="573"/>
            <end position="582"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="9">
        <location>
            <begin position="754"/>
            <end position="776"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="1036"/>
            <end position="1051"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023108"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50112"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016281.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1099" mass="118414" checksum="D3F11012BE7CE57B" modified="2019-05-08" version="1">MSAGCEPGQAPFEEGSVGGAGSGAPRRARATDDMDATSNGSSGTESHGRDSLHSDRSSSGNGKDSALLETTESSKSTNSQSPSPPSSSIAYSLLSTSSEQDNPSTSGCSSEQSARVKTQKELMKALKEMKIRLPSEKRGKGQSGTLATLQYALSCVKQVQASHDYYQQWAIDESQPCYLDVATYTVAELEGMTSEYTLKNPDTFAVAVSFVTGKILYISDQAAFVLHCKRDVFKGATFAEFLAPQDVSIFYGSTAPYHLPSWSACTSATAAAVDYTQEKSVFCRISGGRGRGRELCYTPFRLTPYLAKVRPAAGAEGQPCCLLIAERIHSGYEAPRIPPDKRVFTTRHTPSCLFQDVDERAAPLLGYLPQDLIGTPVLFYLHPEDRPLMLAIHKKILQCGGQPFDHSPLRFCTRHGEYVTIDTSWSSFVNPWSRKISFVMGRHKVRTGPLNEDVFMAPKALKMRPLEPDIQELSEQIHRLLLQPVHSNGSVGLCSMASNTSHEQFLSAASSSDSNGLLAEETPAQRPMTFQQFCKDVHMVKNQGQQMFIESHSKPLPAKPPTAGERPGPLTARGEESGRKEPSSYSYQQINCLDGIIRYLESYNIPSRVKRKCGSSSYTGSSNSDDDKQRAGREAAGSAKGSVLPAPREEEEEPAAAVGPALAPLALPSKAESVVSLTSQCSFSSTIVHVGDKKAPESDTAMLEEGPAPPDREQYRRVGLTKEVLSVHTQQEEQAFLTRFQELSRLHVFDAPAGRHRGPRRGAGRRGKSKAKRVKQHQTSDSTGSLPPPGPWPPSXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXTTSLPFPAVLPVFPPSQPPAEPAPHFPAPLVAPVVALMLPNYVLPPPYFPAPFGEPFGPRLPGPPEASRSTTPERPDSPPFDSRCSSPLQLNLLQMEEAPKAAERQEAPGGQAGGTPGAVELGPQKETGLAEGQDSSNNDALSSSSDLLDLLLQEDSHSGTGSAASGSGSPGLLGSVSASGADLPLSPLPRAASSKSSHTSKYFGSIDSSEAERARQPPGRVGAGEQFIKYVLQDPLWLLMANTDETVMMTYQLPDR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMV5</accession>
    <name>A0A452HMV5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Pleckstrin and Sec7 domain containing 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell projection</location>
            <location evidence="1">Ruffle membrane</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMV5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018603.1">
        <property type="protein sequence ID" value="ENSGAGP00000016297.1"/>
        <property type="gene ID" value="ENSGAGG00000012132.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000018604.1">
        <property type="protein sequence ID" value="ENSGAGP00000016298.1"/>
        <property type="gene ID" value="ENSGAGG00000012132.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032587">
        <property type="term" value="C:ruffle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005543">
        <property type="term" value="F:phospholipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032012">
        <property type="term" value="P:regulation of ARF protein signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd13295">
        <property type="entry name" value="PH_EFA6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00171">
        <property type="entry name" value="Sec7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.1000.11:FF:000004">
        <property type="entry name" value="PH and SEC7 domain-containing protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000054">
        <property type="entry name" value="PH and SEC7 domain-containing protein 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1000.11">
        <property type="entry name" value="Arf Nucleotide-binding Site Opener,domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041681">
        <property type="entry name" value="PH_9"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001605">
        <property type="entry name" value="PH_dom-spectrin-type"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023394">
        <property type="entry name" value="Sec7_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000904">
        <property type="entry name" value="Sec7_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035999">
        <property type="entry name" value="Sec7_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10663">
        <property type="entry name" value="GUANYL-NUCLEOTIDE EXCHANGE FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10663:SF329">
        <property type="entry name" value="PH AND SEC7 DOMAIN-CONTAINING PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15410">
        <property type="entry name" value="PH_9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01369">
        <property type="entry name" value="Sec7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00683">
        <property type="entry name" value="SPECTRINPH"/>
    </dbReference>
    <dbReference type="SMART" id="SM00233">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00222">
        <property type="entry name" value="Sec7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48425">
        <property type="entry name" value="Sec7 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50190">
        <property type="entry name" value="SEC7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="6" id="KW-0966">Cell projection</keyword>
    <keyword evidence="4 7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="SEC7" evidence="10">
        <location>
            <begin position="178"/>
            <end position="342"/>
        </location>
    </feature>
    <feature type="domain" description="PH" evidence="9">
        <location>
            <begin position="392"/>
            <end position="505"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="95"/>
            <end position="114"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="160"/>
            <end position="180"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="627"/>
            <end position="652"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="533"/>
            <end position="560"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="641"/>
            <end position="652"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004632"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50190"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016298.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="13">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="652" mass="73778" checksum="1CE006CABFA81292" modified="2019-05-08" version="1">MEDDKEELQLKDIRDGFSSTFEKIVESDLMKGTYYSSLDSLDVLSLTDETDSCVSFESPLTPLIQQRVKESPELLEQKLVAQQKGALQHMVAEKQEQAAAKPVEEDLGSPLRHSITSSRSENVLSRLSLKNIPNGFHIDGAEEDALKLINSISDASLKDTLSDSDSDMGSTEQLDHGSTDTLANGCKADSDAAKRLAKRLYHLEGFKRCDVARQLGKNNEFSKLVAEEYLSFFDFTGLTLDKALRTFLKAFPLMGETQERERVLIHFSRRYCQCNPEESTSEDGIHTLTCALMLLNTDLHGHVNIGKKMSCQQFITNLDGLNDGKDFAKDLLKTLYNSIKNEKLEWAIDEDELRKSLSELVDDKLGASAKKVTRIVDGSNPFLDIPQALNAITYKHGVLTRKTHADTDGKRTPRGRRGWKKFYAVLKGTILYLQKDEYKPDKDLSEVDLKNAIRVHHALATKASDYSKKSNVLKLKTADWRVFLFQAPSKEEMLSWILRINLVAAIFSAPAFPAAICSMKKFCRPLLPSSMTKLCQEEQLRSHENKMRQIADELTEHKLHPVEKSLKSKEAEEYRLKEHYLIFEKSRYETYINLLCMKIKVGTDDLEKIEASLFMVEADDISLRKTHSSPSLSQGHLSVHSKVEKDVIEQNT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMV8</accession>
    <name>A0A452HMV8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">calcium/calmodulin-dependent protein kinase</fullName>
            <ecNumber evidence="2">2.7.11.17</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="9">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.17"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="8">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.17"/>
        </reaction>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the protein kinase superfamily. CAMK Ser/Thr protein kinase family. CaMK subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.17" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452HMV8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018609.1">
        <property type="protein sequence ID" value="ENSGAGP00000016303.1"/>
        <property type="gene ID" value="ENSGAGG00000006503.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004683">
        <property type="term" value="F:calcium/calmodulin-dependent protein kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd14086">
        <property type="entry name" value="STKc_CaMKII"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000001">
        <property type="entry name" value="Calcium/calmodulin-dependent protein kinase type II subunit delta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.450.50:FF:000001">
        <property type="entry name" value="calcium/calmodulin-dependent protein kinase type II subunit gamma isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.450.50">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.620">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013543">
        <property type="entry name" value="Ca/CaM-dep_prot_kinase-assoc"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032710">
        <property type="entry name" value="NTF2-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24347">
        <property type="entry name" value="SERINE/THREONINE-PROTEIN KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08332">
        <property type="entry name" value="CaMKII_AD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54427">
        <property type="entry name" value="NTF2-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="6" id="KW-0418">Kinase</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="5" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Protein kinase" evidence="10">
        <location>
            <begin position="1"/>
            <end position="220"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00005354"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00012434"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00047307"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00047430"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016303.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="411" mass="46780" checksum="0EA08CC3A2C9F8FE" modified="2019-05-08" version="1">FINHSNVEEARICRLLKHPNIVRLHDSISEEGFHYLVFDLVTGGELFEDIVAREYYSEADASHCIQQILEAVLHCHQMGVVHRDLKPENLLLASKSKGAAVKLADFGLAIEVQGEQQAWFGFAGTPGYLSPEVLRKDPYGKPVDMWACGVILYILLVGYPPFWDEDQHRLYQQIKAGAYDFPSPEWDTVTPEAKDLINKMLTINPAKRITASEALKHPWICQRSTVASMMHRQETVDCLKKFNARRKLKVRKKIFFRGILVESTESSNTTIEDEDVKARKQEIIKVTEQLIEAINNGDFEAYTKICDPGLTSFEPEALGNLVEGMDFHRFYFENALSKGNKPIHTIILNPHVHLVGDDAACIAYIRLTQYMDGSGMPKTMQSEETRVWHRRDGKWQNVHFHCSGSPTVPIK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMW0</accession>
    <name>A0A452HMW0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Attractin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="25 26">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="26" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="25" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="25" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="16">Involved in the initial immune cell clustering during inflammatory response and may regulate chemotactic activity of chemokines. May play a role in melanocortin signaling pathways that regulate energy homeostasis and hair color. Low-affinity receptor for agouti. Has a critical role in normal myelination in the central nervous system.</text>
    </comment>
    <comment type="subunit">
        <text evidence="17">Monomer and homotrimer.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="19">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018617.1">
        <property type="protein sequence ID" value="ENSGAGP00000016311.1"/>
        <property type="gene ID" value="ENSGAGG00000011749.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030246">
        <property type="term" value="F:carbohydrate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006954">
        <property type="term" value="P:inflammatory response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00041">
        <property type="entry name" value="CUB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00055">
        <property type="entry name" value="EGF_Lam"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.120.10.80:FF:000031">
        <property type="entry name" value="attractin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.120.10.80:FF:000034">
        <property type="entry name" value="Attractin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000079">
        <property type="entry name" value="Attractin like 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000164">
        <property type="entry name" value="Attractin like 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.290:FF:000008">
        <property type="entry name" value="Attractin like 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.120.10.80">
        <property type="entry name" value="Kelch-type beta propeller"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.100.10">
        <property type="entry name" value="Mannose-Binding Protein A, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.290">
        <property type="entry name" value="Spermadhesin, CUB domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056737">
        <property type="entry name" value="Beta-prop_ATRN-MKLN-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001304">
        <property type="entry name" value="C-type_lectin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016186">
        <property type="entry name" value="C-type_lectin-like/link_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016187">
        <property type="entry name" value="CTDL_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000859">
        <property type="entry name" value="CUB_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056732">
        <property type="entry name" value="GBD_ATRN"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015915">
        <property type="entry name" value="Kelch-typ_b-propeller"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002049">
        <property type="entry name" value="LE_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056863">
        <property type="entry name" value="LMN_ATRN_NET-like_EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051568">
        <property type="entry name" value="LZTR1/Attractin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002165">
        <property type="entry name" value="Plexin_repeat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016201">
        <property type="entry name" value="PSI"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035914">
        <property type="entry name" value="Sperma_CUB_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46376:SF3">
        <property type="entry name" value="ATTRACTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46376">
        <property type="entry name" value="LEUCINE-ZIPPER-LIKE TRANSCRIPTIONAL REGULATOR 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24981">
        <property type="entry name" value="Beta-prop_ATRN-LZTR1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24973">
        <property type="entry name" value="EGF_LMN_ATRN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24972">
        <property type="entry name" value="GBD_ATRN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01437">
        <property type="entry name" value="PSI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00034">
        <property type="entry name" value="CLECT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00042">
        <property type="entry name" value="CUB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00180">
        <property type="entry name" value="EGF_Lam"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00423">
        <property type="entry name" value="PSI"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56436">
        <property type="entry name" value="C-type lectin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF117281">
        <property type="entry name" value="Kelch motif"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49854">
        <property type="entry name" value="Spermadhesin, CUB domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50041">
        <property type="entry name" value="C_TYPE_LECTIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01180">
        <property type="entry name" value="CUB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01248">
        <property type="entry name" value="EGF_LAM_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50027">
        <property type="entry name" value="EGF_LAM_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11 19" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="4 19" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="14" id="KW-0395">Inflammatory response</keyword>
    <keyword evidence="2" id="KW-0880">Kelch repeat</keyword>
    <keyword evidence="15 20" id="KW-0424">Laminin EGF-like domain</keyword>
    <keyword evidence="7" id="KW-0430">Lectin</keyword>
    <keyword evidence="10" id="KW-0472">Membrane</keyword>
    <keyword evidence="12" id="KW-0675">Receptor</keyword>
    <keyword evidence="26" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="5" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9" id="KW-1133">Transmembrane helix</keyword>
    <feature type="domain" description="EGF-like" evidence="22">
        <location>
            <begin position="48"/>
            <end position="81"/>
        </location>
    </feature>
    <feature type="domain" description="CUB" evidence="21">
        <location>
            <begin position="83"/>
            <end position="193"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="22">
        <location>
            <begin position="191"/>
            <end position="228"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="24">
        <location>
            <begin position="737"/>
            <end position="857"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="23">
        <location>
            <begin position="1001"/>
            <end position="1046"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="19">
        <location>
            <begin position="52"/>
            <end position="62"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="19">
        <location>
            <begin position="71"/>
            <end position="80"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="19">
        <location>
            <begin position="195"/>
            <end position="205"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="19">
        <location>
            <begin position="199"/>
            <end position="216"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="19">
        <location>
            <begin position="218"/>
            <end position="227"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="1018"/>
            <end position="1027"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="1030"/>
            <end position="1044"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022441"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022734"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023198"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023292"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00058400"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00065783"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00074596"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00460"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS01180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS50027"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS50041"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016311.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1294" mass="144077" checksum="043CB0F1D8E9FADB" modified="2019-05-08" version="1">MSPCCEPVREWPDSKCETPGQSVGGNRRLYKIKPGQLVTLAAAAAAVEGKECDKPCANGGRCQPGSGQCECQAGWVGDQCQHCGGRFRLTGSSGYVTDGPGHYKYKTKSNTLIFRELRFNHFATECSWDHLYVYDGDSVYAPLVAAFSGLIVPEKDGNETIPEVVATSGYVLLHFFSDAAYNLTGFNITYNFDMCPNNCSGRGECKHSNGSNTVKCECSEHWKGEACDISYCTDDCGYPARGQCNMNETKGCLCNSGWQGPGCAVPVPANQSFWTREEYSSPKLPRASHKAVVNGNVMWVVGGYMFNHSDSHMILAYDLVSEEWLPLNSTVNSVEIRYGHSLALHENKIYMYGGKIDAMGNVTNQLWVFHIHNESWVHVIPKTKEQYAVVGHSAHIVNLGDGSAVMLVIFGHCPLYGYISNVQEYNLATNTWNILQTSGALVQGGYGHSSVYDPNTQSIYIHGGYKAFSANKYRLADDLYKYEVDTRMTILKDSRFFRYLHTAVIMSGTMLVFGGNTHNDTSMSHGAKCFSSDFMAYDIACDHWSVLPRPDLHHDVNRFGHSAVLSNSTVYIFGGFNSLLLSDILRYTPESCEAYDNETSCLRAGPGVRCVWNTTFSLCVPWERATVEQQQKVFEECPSKPDHEKCNQITDCYSCTANTNNCQWCTDQCILKNNNCTEDQVPITEYDKCPKDNPAYYCNKKTSCKSCAMDQNCQWEPRNQECIALPENICGANWHLVGNSCLKINSTKESYDSGKLACTTQNALLASLTTQKKVEFVLKELQRMQSLAKTLTPWVGLRKINVSYWCWEDMSPFTNTLVQWLPGEPSGAGFCGYLAEPYQRGLKAATCIDHVNGSICERLANHSAKQCRTPCALRTACGECTSGSSECMWCSNMKQCVDSNAYVASFPYGQCMEWYTMSSCPPANCSGYCTCAHCLEQPGCGWCTDPSNTGKGKCMEGSYRGPVKMPSPPATGKYYLEPILNVSMCPAENKYNWSFIQCPACQCNGHSKCVNESVCEKCENLTTGKHCETCISGYYGDPTNGGTCQSCKCNGHASVCNTNTGKCFCTTKGIKGDECQLCEVENRYQGNPLKGTCYYTLLIDYQFTFSLSQEDDRYYTAINFVATPEEQNRDLDMFINASKNFNLNITWSTSFAAGTQAGEEIPVVSKSNIKEYKDSFSNEKFDFRSNPNITFFVYVSNFTWPIKIQIAFSQHSNFMDLVQFFVTFFSTKITVSRLPVLPATAELSIFLYLPPPPLPLWRTLRRIKETFNQPPYLCGVPLCIITETHILTKSPSAT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN05</accession>
    <name>A0A452HN05_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Protein transport protein SEC23</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="12">Component of the coat protein complex II (COPII) which promotes the formation of transport vesicles from the endoplasmic reticulum (ER). The coat has two main functions, the physical deformation of the endoplasmic reticulum membrane into vesicles and the selection of cargo molecules for their transport to the Golgi complex.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="13">Cytoplasmic vesicle</location>
            <location evidence="13">COPII-coated vesicle membrane</location>
            <topology evidence="13">Peripheral membrane protein</topology>
            <orientation evidence="13">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1 13">Endoplasmic reticulum membrane</location>
            <topology evidence="1 13">Peripheral membrane protein</topology>
            <orientation evidence="1 13">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="13">Cytoplasm</location>
            <location evidence="13">Cytosol</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 13">Belongs to the SEC23/SEC24 family. SEC23 subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HN05"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018639.1">
        <property type="protein sequence ID" value="ENSGAGP00000016332.1"/>
        <property type="gene ID" value="ENSGAGG00000006201.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030127">
        <property type="term" value="C:COPII vesicle coat"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070971">
        <property type="term" value="C:endoplasmic reticulum exit site"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090110">
        <property type="term" value="P:COPII-coated vesicle cargo loading"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006886">
        <property type="term" value="P:intracellular protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd01478">
        <property type="entry name" value="Sec23-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd11287">
        <property type="entry name" value="Sec23_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.730:FF:000003">
        <property type="entry name" value="Protein transport protein SEC23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.380:FF:000001">
        <property type="entry name" value="Protein transport protein SEC23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.20.10:FF:000003">
        <property type="entry name" value="Protein transport protein SEC23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.410:FF:000011">
        <property type="entry name" value="Protein transport protein SEC23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1670">
        <property type="entry name" value="beta-sandwich domain of Sec23/24"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.730">
        <property type="entry name" value="Sec23/Sec24 helical domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.20.10">
        <property type="entry name" value="Severin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.410">
        <property type="entry name" value="von Willebrand factor, type A domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.380">
        <property type="entry name" value="Zn-finger domain of Sec23/24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029006">
        <property type="entry name" value="ADF-H/Gelsolin-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012990">
        <property type="entry name" value="Beta-sandwich_Sec23_24"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007123">
        <property type="entry name" value="Gelsolin-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036180">
        <property type="entry name" value="Gelsolin-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037364">
        <property type="entry name" value="Sec23"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006900">
        <property type="entry name" value="Sec23/24_helical_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036175">
        <property type="entry name" value="Sec23/24_helical_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006896">
        <property type="entry name" value="Sec23/24_trunk_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037550">
        <property type="entry name" value="Sec23_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036465">
        <property type="entry name" value="vWFA_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006895">
        <property type="entry name" value="Znf_Sec23_Sec24"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036174">
        <property type="entry name" value="Znf_Sec23_Sec24_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11141">
        <property type="entry name" value="PROTEIN TRANSPORT PROTEIN SEC23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11141:SF10">
        <property type="entry name" value="PROTEIN TRANSPORT PROTEIN SEC23B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00626">
        <property type="entry name" value="Gelsolin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08033">
        <property type="entry name" value="Sec23_BS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04815">
        <property type="entry name" value="Sec23_helical"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04811">
        <property type="entry name" value="Sec23_trunk"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04810">
        <property type="entry name" value="zf-Sec23_Sec24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81995">
        <property type="entry name" value="beta-sandwich domain of Sec23/24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF82754">
        <property type="entry name" value="C-terminal, gelsolin-like domain of Sec23/24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81811">
        <property type="entry name" value="Helical domain of Sec23/24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53300">
        <property type="entry name" value="vWA-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF82919">
        <property type="entry name" value="Zn-finger domain of Sec23/24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 13" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11 13" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="6 13" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="8 13" id="KW-0931">ER-Golgi transport</keyword>
    <keyword evidence="10 13" id="KW-0472">Membrane</keyword>
    <keyword evidence="5 13" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9 13" id="KW-0653">Protein transport</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 13" id="KW-0813">Transport</keyword>
    <keyword evidence="7 13" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="Zinc finger Sec23/Sec24-type" evidence="15">
        <location>
            <begin position="58"/>
            <end position="98"/>
        </location>
    </feature>
    <feature type="domain" description="Sec23/Sec24 trunk" evidence="16">
        <location>
            <begin position="127"/>
            <end position="380"/>
        </location>
    </feature>
    <feature type="domain" description="Sec23/Sec24 beta-sandwich" evidence="18">
        <location>
            <begin position="386"/>
            <end position="467"/>
        </location>
    </feature>
    <feature type="domain" description="Sec23/Sec24 helical" evidence="17">
        <location>
            <begin position="481"/>
            <end position="579"/>
        </location>
    </feature>
    <feature type="domain" description="Gelsolin-like" evidence="14">
        <location>
            <begin position="595"/>
            <end position="681"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004397"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009210"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022892"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00037370"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="RuleBase" id="RU365030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="Pfam" id="PF00626"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="Pfam" id="PF04810"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF04811"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF04815"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF08033"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016332.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="728" mass="82473" checksum="BBF2F1F1A3988551" modified="2019-05-08" version="1">MATYLEFIQQNEERDGVRFSWNVWPSSRLEATRMVVPLACLLTPLKERLDLPPVQYEPVLCSRQTCKAVLNPLCQVDYRAKLWACNFCFQRNQFPPAYAGISEINQPAELMPQFSTIEYIVQVNASWLKEDLQALKESLQMSLSLLPPEALVGLITFGRMVQVHELSCEGISKSYVFRGTKDLTAKQIQDMLGLTRPVVPVQQGRPLQPQEQPPISSRFLQPVHKIDMNLTDLLGELQRDPWPVTQGKRPLRSTGVALSIAVGLLEGTFPNTGARIMLFTGGPPTQGPGMVVGDELKTPIRSWHDIEKDNARFMKKATKHYETLANRTAANGHCIDIYACALDQTGLLEMKCCANLTGGHMMMGDSFNTSLFKQTFQRVFSKDFNGEFRMAFGATLEVKELGIGGTCQWKICSLDPSSTLAIYFEVVNQHNAPIPQGGRGAVQFVTQYQHSSTQRRIRVTTIARNWADAQSQLQHIEAAFDQEVAAVLMARLGVYRAESEEGPDVLRWLDRQLIRLCQKFGQYNKDDPNSFRLSDSFSLYPQFMFHLRRSPFLQVFNNSPDESSFYRHHFSRQDLTQSLIMIQPILYSYSFSGPPEPVLLDSSSILADRILLMDTFFQIVIYHGETIAQWQKAGYQDMPEYENFKHLLQAPLDDAQEILQTRFPMPRYINTEHGSSQARFLLSKVNPSQTHNNLYAWGQESGSPILTDDVSLQVFMDHLKKLAVSSAS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN09</accession>
    <name>A0A452HN09_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Saccharopine dehydrogenase-like oxidoreductase</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the saccharopine dehydrogenase family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018652.1">
        <property type="protein sequence ID" value="ENSGAGP00000016344.1"/>
        <property type="gene ID" value="ENSGAGG00000012242.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005811">
        <property type="term" value="C:lipid droplet"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009247">
        <property type="term" value="P:glycolipid biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.720">
        <property type="entry name" value="NAD(P)-binding Rossmann-like Domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036291">
        <property type="entry name" value="NAD(P)-bd_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051276">
        <property type="entry name" value="Saccharopine_DH-like_oxidrdct"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005097">
        <property type="entry name" value="Sacchrp_dh_NADP-bd"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12286">
        <property type="entry name" value="SACCHAROPINE DEHYDROGENASE-LIKE OXIDOREDUCTASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12286:SF5">
        <property type="entry name" value="SACCHAROPINE DEHYDROGENASE-LIKE OXIDOREDUCTASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03435">
        <property type="entry name" value="Sacchrp_dh_NADP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51735">
        <property type="entry name" value="NAD(P)-binding Rossmann-fold domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="3" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="3">
        <location>
            <begin position="252"/>
            <end position="278"/>
        </location>
    </feature>
    <feature type="domain" description="Saccharopine dehydrogenase NADP binding" evidence="4">
        <location>
            <begin position="16"/>
            <end position="114"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00038048"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00039852"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="Pfam" id="PF03435"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016344.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="401" mass="44121" checksum="59F04AB6199A97B0" modified="2019-05-08" version="1">MATRSGAAAGRPYELVVFGASGFTGHFVAEELARVAAGAELRGSLRWAVAGRSRDKLQGALDRAAERLGKPELKSEVGIILCDVSDPSSLADMAKQAAVVLNCVGPFLEGMYLKYNDKAAEKGVYVVGSSGFDSIPADMGVLYTRDNLKGTLTAVESFLTVKSGPEGYSIHDGTWKSAIHGLADQDNLRKLRKQIGHKPLPIIGAKLKRRGLVFYSQEFKEYSIPFMGSDVSVVKRTQRYLHTDLQETPVQYAAYATVGGIASVIKLMFAGLLFFLLVKFSFGRNLLTKYPELFSCGRFTKKGPTQKQIDGSSFTMTFFGEGYCEGQDPQEGKPNVKICTQVKGPGKEARMFYYGHLKNSRSAVCHVPLFYXKKPECFIMVILKIQGQLFVMYPSSTDHYF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN10</accession>
    <name>A0A452HN10_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Major facilitator superfamily (MFS) profile domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HN10"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018644.1">
        <property type="protein sequence ID" value="ENSGAGP00000016337.1"/>
        <property type="gene ID" value="ENSGAGG00000012116.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008028">
        <property type="term" value="F:monocarboxylic acid transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd17421">
        <property type="entry name" value="MFS_MCT5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1250.20:FF:000277">
        <property type="entry name" value="Solute carrier family 16 member 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1250.20:FF:000141">
        <property type="entry name" value="Solute carrier family 16, member 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1250.20">
        <property type="entry name" value="MFS general substrate transporter like domains"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011701">
        <property type="entry name" value="MFS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020846">
        <property type="entry name" value="MFS_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036259">
        <property type="entry name" value="MFS_trans_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050327">
        <property type="entry name" value="Proton-linked_MCT"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11360">
        <property type="entry name" value="MONOCARBOXYLATE TRANSPORTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11360:SF14">
        <property type="entry name" value="MONOCARBOXYLATE TRANSPORTER 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07690">
        <property type="entry name" value="MFS_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103473">
        <property type="entry name" value="MFS general substrate transporter"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50850">
        <property type="entry name" value="MFS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0472">Membrane</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="2" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="20"/>
            <end position="47"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="110"/>
            <end position="132"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="144"/>
            <end position="165"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="171"/>
            <end position="195"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="288"/>
            <end position="306"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="326"/>
            <end position="346"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="358"/>
            <end position="374"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="380"/>
            <end position="403"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="415"/>
            <end position="435"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="441"/>
            <end position="463"/>
        </location>
    </feature>
    <feature type="domain" description="Major facilitator superfamily (MFS) profile" evidence="3">
        <location>
            <begin position="287"/>
            <end position="483"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS50850"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016337.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="483" mass="52806" checksum="BC97B300912288B0" modified="2019-05-08" version="1">LTAMEEEEQQPIPPPDGGWGWMIVLHFFLVNVLVMGMLKTFGIFFVVFQEEVGGSSEQISWIGSIMSSLRFSAGPLVTIACGKLGDQLTSIIGAGLVSSGCLISTQATSISFLCVSMGLLLGLGFACLYQAAAVMTARYFKKRLALSNAIARSGMGLTFILAPFTQLLIDFYGWQGTLLIFGGITLNLVPSSMLLRPVIKKLQDFSRNVESGSLTQSKHPATLTSHPDKISTTETQFNIRLDSPMQDSAVTNGFQSAETLGSLEWNSNSRPVPSRQPLLDFSLLKDPFFCIFVWSFLFSQLAYFIPTFHLAARAKTLGIDSMDASYLISVAGITETVSQLLSGWIADQNWIKKYHYHKTYLILCGITNLLGPLATSFPLLMVYTVSFAIFCGGYLALLLPVLVDFVGVPRLHSSLGFASFFAGIAAIAGPPIAGWLSDYTQTYACSFLFAGTCYLIAPISFFFEPSAHKWKLQKEKELNEVTS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN13</accession>
    <name>A0A452HN13_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="5">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HN13"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018672.1">
        <property type="protein sequence ID" value="ENSGAGP00000016362.1"/>
        <property type="gene ID" value="ENSGAGG00000012258.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045202">
        <property type="term" value="C:synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="GOC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030550">
        <property type="term" value="F:acetylcholine receptor inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0095500">
        <property type="term" value="P:acetylcholine receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10036">
        <property type="entry name" value="CD59 GLYCOPROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10036:SF7">
        <property type="entry name" value="LY6_PLAUR DOMAIN-CONTAINING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1 4" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="4">
        <location>
            <begin position="1"/>
            <end position="32"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019295564" evidence="4">
        <location>
            <begin position="33"/>
            <end position="149"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="125"/>
            <end position="149"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016362.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="149" mass="16501" checksum="57C3416B55BD04DC" modified="2019-05-08" version="1">MRRGGCRAARGPGSTRMWLLFGAATFWGLVLAPGFGLQIQCYQCEEFQLNNDCSAPEFIVNCTVNVQDMCQKEVMEKSSGESPWSSCCRFSHPTHSRMQQREEVLPVQAGGTRSLLSVNSCSGGGFSKRNAQDREMKTQQSILNPKAEN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN14</accession>
    <name>A0A452HN14_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Calponin-homology (CH) domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the CCDC88 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018650.1">
        <property type="protein sequence ID" value="ENSGAGP00000016342.1"/>
        <property type="gene ID" value="ENSGAGG00000012232.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051959">
        <property type="term" value="F:dynein light intermediate chain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008017">
        <property type="term" value="F:microtubule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031122">
        <property type="term" value="P:cytoplasmic microtubule organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030705">
        <property type="term" value="P:cytoskeleton-dependent intracellular transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd22229">
        <property type="entry name" value="HkD_Girdin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.418.10:FF:000035">
        <property type="entry name" value="girdin isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.418.10">
        <property type="entry name" value="Calponin-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001715">
        <property type="entry name" value="CH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036872">
        <property type="entry name" value="CH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043936">
        <property type="entry name" value="HOOK_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18947:SF30">
        <property type="entry name" value="GIRDIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18947">
        <property type="entry name" value="HOOK PROTEINS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19047">
        <property type="entry name" value="HOOK_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF116907">
        <property type="entry name" value="Hook domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50021">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 6" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="3" id="KW-0344">Guanine-nucleotide releasing factor</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Calponin-homology (CH)" evidence="8">
        <location>
            <begin position="12"/>
            <end position="132"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="820"/>
            <end position="848"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1008"/>
            <end position="1037"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1418"/>
            <end position="1462"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1563"/>
            <end position="1607"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="245"/>
            <end position="424"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="1128"/>
            <end position="1386"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="1016"/>
            <end position="1031"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="1421"/>
            <end position="1433"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="1448"/>
            <end position="1462"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="1563"/>
            <end position="1582"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="1597"/>
            <end position="1607"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022658"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00061299"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016342.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1811" mass="210060" checksum="0310E12F5546319C" modified="2019-05-08" version="1">MENEIFTPLMEQFISSPLVTWVKTFGPLAGGNGSNLEEYVALVDGIFLNEVMRQINPKSTTQRVNKKVNNDASLRIQNLSILVRQIKSYYQETLQQLIMMSLPNVLIIGRNPFSEPGTEEIKKLLLLLLGCAVQCQKKEEFIERIQGLDFDTRAAVAAHIQEVTHNQENVFDLQWMDVSTLSQEYIEPLLKNMALHLKRLIDERDEHSETIVELSEERDCLHFLPHASAAQSPCGSPGIKRTESRQHLSVELADAKAKIRRLRQELEEKTEQLLDCKQELEQMEAELKRLQQENMNLLSDARSARVYRDELDVLREKAIRVDKLESEVSRYKERLHDIVFYKARVEELKEDNQVLLETKTMLEDQLEGTRARSDKLHELEKENLQLKAKLHDMEMERDMDRKKIEELMEENMTLEMAQKQSMDESLHLGWELEQMNRTTELSEVPRKSLGHEVNELTSSRLLKLEMENQSLLKTVEELQNVAGSVEGSNSKILKMEKENQRLSKKLEGLENEIKQEKQSLQNSQNLSKDLMKEKAQLEKTIETLRENSERQIKILEQENEHLNQTVDALRQRSQISAEARVKDIEKENKIIHESIKEASSKLNKIEFEKKQMRKELEHYKEKGERAEELENELHHLEKENELLQKKITNLKITCEKTEGLEQENSDLEVENRKLKKTLDSFKNLTFQLESLEKENSQLDEENLELRRTVESLKCTSIKMAQLQLENKELESEKKQLKKGLELMKASCKKTERLEVSYQGLDTENQRLQKALENSTKKIQQLESELQDLESENQTLQKNLEELKISSKRLEQLEKENKLLEQETSQLEKDKKQLEKENKRLRQQAEIKDSTLEENNTKISNLEKENKSLCKEISIFKESCVRLKELEKENKELVKRATIDKKTLITLREDLVNEKLKTQQMNNDLEKLTHELEKIGLNKERLLHDEQSTDDSRYKLLESKLESTLKKSLEIKEEKIAALEARLEESTNLNQQLRQELKTVKKNYEALKQRQEEERMVQNSPPRTGEETQSVNKWEKENQETTRELLKVKDRLIEVERNNATLQAEKQALKTQLKQLETQNNNLQAQILSLQRQTVSLQEQNTTLQTQNAKLQVENSTLNSQSTSLMNQNAQLLIQQSALENENESILREREDLKSLYDSLVKDHEKLEHLHERQASEYESLIAKHGSLKSAHKNLEVEHKDLEDRYNQLLKQKVQLEELEKVLKVEQEKMLLQSKKHKTVAAEYKKLCDENDRLNHTYNQLLRENEVLQTDHKNLKSLLNNSKLEQTRLEAEFSKLKEQYQQLDITSTKLNNQCELLSQLKGNLEEENRHLLDQIQTLMLQNRTLLEQNMESKDLFHVEQRQYIDKLNELRRQKEKLEEKIMDQYKFYEPSPPRRRGNWITLKMRKLIKSKKDVNRERLKSLTLTPTRSESSEGFLQLPHQDSQDSSSVGSNSLEDGQTLGTKKSSMIALKRLPFLRNRPKEKDKMKAIYRRSMSMNDLVQSMVLAGGQWTGSSEHLEGSDDVSTGKRKKELGAMAFSTTAINFATVNSSSGFRSKQLLNNKDTTSFEDVSPQGISDDSSTGSRVHASRPASLDSGRTSTSNSNNNASLHEVKAGGINNQSRPQSHSSGEFSLLHEHEAWSSSSSSPIQYLKSHTKLSPMIQEKMPETVDNQGKQKIKTGSPGSEVVTLQQFLEESNKVTSTEMWKLYQKAEIQKAGLGSNKAPNAITHYMRCGPSGEFPAQNVNEINFIYNKLTALPIRISAHCINRFKYYLLNWLSILLISKARPDSCSWSILPVWAGNEKSLNTFFLVK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN15</accession>
    <name>A0A452HN15_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Calponin-homology (CH) domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the CCDC88 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018660.1">
        <property type="protein sequence ID" value="ENSGAGP00000016350.1"/>
        <property type="gene ID" value="ENSGAGG00000012232.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051959">
        <property type="term" value="F:dynein light intermediate chain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008017">
        <property type="term" value="F:microtubule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031122">
        <property type="term" value="P:cytoplasmic microtubule organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030705">
        <property type="term" value="P:cytoskeleton-dependent intracellular transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd22229">
        <property type="entry name" value="HkD_Girdin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.418.10:FF:000035">
        <property type="entry name" value="girdin isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.418.10">
        <property type="entry name" value="Calponin-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001715">
        <property type="entry name" value="CH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036872">
        <property type="entry name" value="CH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043936">
        <property type="entry name" value="HOOK_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18947:SF30">
        <property type="entry name" value="GIRDIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18947">
        <property type="entry name" value="HOOK PROTEINS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19047">
        <property type="entry name" value="HOOK_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF116907">
        <property type="entry name" value="Hook domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50021">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 6" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="3" id="KW-0344">Guanine-nucleotide releasing factor</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Calponin-homology (CH)" evidence="8">
        <location>
            <begin position="12"/>
            <end position="132"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="820"/>
            <end position="848"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1008"/>
            <end position="1037"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1418"/>
            <end position="1464"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1535"/>
            <end position="1579"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="245"/>
            <end position="424"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="1128"/>
            <end position="1386"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="1016"/>
            <end position="1031"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="1421"/>
            <end position="1433"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="1448"/>
            <end position="1464"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="1535"/>
            <end position="1554"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="1569"/>
            <end position="1579"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022658"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00061299"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016350.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1783" mass="206601" checksum="73B4A85BF26F4924" modified="2019-05-08" version="1">MENEIFTPLMEQFISSPLVTWVKTFGPLAGGNGSNLEEYVALVDGIFLNEVMRQINPKSTTQRVNKKVNNDASLRIQNLSILVRQIKSYYQETLQQLIMMSLPNVLIIGRNPFSEPGTEEIKKLLLLLLGCAVQCQKKEEFIERIQGLDFDTRAAVAAHIQEVTHNQENVFDLQWMDVSTLSQEYIEPLLKNMALHLKRLIDERDEHSETIVELSEERDCLHFLPHASAAQSPCGSPGIKRTESRQHLSVELADAKAKIRRLRQELEEKTEQLLDCKQELEQMEAELKRLQQENMNLLSDARSARVYRDELDVLREKAIRVDKLESEVSRYKERLHDIVFYKARVEELKEDNQVLLETKTMLEDQLEGTRARSDKLHELEKENLQLKAKLHDMEMERDMDRKKIEELMEENMTLEMAQKQSMDESLHLGWELEQMNRTTELSEVPRKSLGHEVNELTSSRLLKLEMENQSLLKTVEELQNVAGSVEGSNSKILKMEKENQRLSKKLEGLENEIKQEKQSLQNSQNLSKDLMKEKAQLEKTIETLRENSERQIKILEQENEHLNQTVDALRQRSQISAEARVKDIEKENKIIHESIKEASSKLNKIEFEKKQMRKELEHYKEKGERAEELENELHHLEKENELLQKKITNLKITCEKTEGLEQENSDLEVENRKLKKTLDSFKNLTFQLESLEKENSQLDEENLELRRTVESLKCTSIKMAQLQLENKELESEKKQLKKGLELMKASCKKTERLEVSYQGLDTENQRLQKALENSTKKIQQLESELQDLESENQTLQKNLEELKISSKRLEQLEKENKLLEQETSQLEKDKKQLEKENKRLRQQAEIKDSTLEENNTKISNLEKENKSLCKEISIFKESCVRLKELEKENKELVKRATIDKKTLITLREDLVNEKLKTQQMNNDLEKLTHELEKIGLNKERLLHDEQSTDDSRYKLLESKLESTLKKSLEIKEEKIAALEARLEESTNLNQQLRQELKTVKKNYEALKQRQEEERMVQNSPPRTGEETQSVNKWEKENQETTRELLKVKDRLIEVERNNATLQAEKQALKTQLKQLETQNNNLQAQILSLQRQTVSLQEQNTTLQTQNAKLQVENSTLNSQSTSLMNQNAQLLIQQSALENENESILREREDLKSLYDSLVKDHEKLEHLHERQASEYESLIAKHGSLKSAHKNLEVEHKDLEDRYNQLLKQKVQLEELEKVLKVEQEKMLLQSKKHKTVAAEYKKLCDENDRLNHTYNQLLRENEVLQTDHKNLKSLLNNSKLEQTRLEAEFSKLKEQYQQLDITSTKLNNQCELLSQLKGNLEEENRHLLDQIQTLMLQNRTLLEQNMESKDLFHVEQRQYIDKLNELRRQKEKLEEKIMDQYKFYEPSPPRRRGNWITLKMRKLIKSKKDVNRERLKSLTLTPTRSESSEGFLQLPHQDSQDSSSVGSNSLEDGQTLGTKKSSTMNDLVQSMVLAGGQWTGSSEHLEGSDDVSTGKRKKELGAMAFSTTAINFATVNSSSGFRSKQLLNNKDTTSFEDVSPQGISDDSSTGSRVHASRPASLDSGRTSTSNSNNNASLHEVKAGGINNQSRPQSHSSGEFSLLHEHEAWSSSSSSPIQYLKSHTKLSPMIQEKMPETVDNQGKQKIKTGSPGSEVVTLQQFLEESNKVTSTEMWKLYQKAEIQKAGLGSNKAPNAITHYMRCGPSGEFPAQNVNEINFIYNKLTALPIRISAHCINRFKYYLLNWLSILLISKARPDSCSWSILPVWAGNEKSLNTFFLVK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN17</accession>
    <name>A0A452HN17_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">CWH43-like N-terminal domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endomembrane system</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the DRAM/TMEM150 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HN17"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018697.1">
        <property type="protein sequence ID" value="ENSGAGP00000016383.1"/>
        <property type="gene ID" value="ENSGAGG00000012271.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012505">
        <property type="term" value="C:endomembrane system"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019402">
        <property type="entry name" value="CWH43_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050911">
        <property type="entry name" value="DRAM/TMEM150_Autophagy_Mod"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21324">
        <property type="entry name" value="FASTING-INDUCIBLE INTEGRAL MEMBRANE PROTEIN TM6P1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21324:SF9">
        <property type="entry name" value="TRANSMEMBRANE PROTEIN 150A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10277">
        <property type="entry name" value="Frag1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 7" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 7" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="48"/>
            <end position="66"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="78"/>
            <end position="99"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="111"/>
            <end position="133"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="145"/>
            <end position="166"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="172"/>
            <end position="192"/>
        </location>
    </feature>
    <feature type="domain" description="CWH43-like N-terminal" evidence="8">
        <location>
            <begin position="32"/>
            <end position="201"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="219"/>
            <end position="242"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="222"/>
            <end position="231"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004127"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006565"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF10277"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016383.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="242" mass="26563" checksum="B3602EA263DE51D1" modified="2019-05-08" version="1">MWGWATRWTHLPRIYNQSCGLDGPISCCTLDHIPLVSKCGTFPPESCFFSLICSLGSFMVMLVGLLRYAHVIECCGPSLLNTLGLAAGWICAAGLTMVGNFQVDYAKVLHYIGAGVAFPTSMLFVFLQSVLTYRMAKTRGHYWTGHLRSILSALAFITLVFSGVFFVQESFVLQHVAALCEWIFIINVLVFYGTFTFEFGAISTDTFLVLLKASQAPKSYKSDSSTASTPHGHSHSEGLAMV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN18</accession>
    <name>A0A452HN18_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">C2H2-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HN18"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018657.1">
        <property type="protein sequence ID" value="ENSGAGP00000016347.1"/>
        <property type="gene ID" value="ENSGAGG00000012247.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd21440">
        <property type="entry name" value="KLF8_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000021">
        <property type="entry name" value="Basic krueppel-like factor 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000018">
        <property type="entry name" value="Krueppel-like factor 15"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000563">
        <property type="entry name" value="Krueppel-like factor 8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23235:SF46">
        <property type="entry name" value="KRUEPPEL-LIKE FACTOR 8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23235">
        <property type="entry name" value="KRUEPPEL-LIKE TRANSCRIPTION FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0678">Repressor</keyword>
    <keyword evidence="9" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 11" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="250"/>
            <end position="279"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="280"/>
            <end position="309"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="310"/>
            <end position="335"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1"/>
            <end position="54"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="202"/>
            <end position="245"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="1"/>
            <end position="10"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022491"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016347.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="335" mass="36580" checksum="62EB8FFBFDB515F0" modified="2019-05-08" version="1">MHSEEQKEKAAGSVQSRVLPKMSSSSLLEDNSIEPPVLLTDIKTEPPEELLASDCNQPQAEPVDLSLNKPKVSLQSSTLLPSSIRSSPMLVAPPIQAPVVSMSPTGLGSSSAIPAVLSPGSILASTQGSGGQQILHVIHTIPSVNLPSKMGNLQTIPVVVQSLPVVYTTMPTDGVTAAITVPLIGGDGKNAGSVKIDPSSMYPLEMNSDSEDSVTESSSAPFQDLQRESRSPVARMQRAESPDLKKRRIHQCDFEGCNKVYTKSSHLKAHRRIHTGEKPYKCTWEGCTWKFARSDELTRHFRKHTGIKPFRCSDCDRSFSRSDHLALHRRRHIMM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN63</accession>
    <name>A0A452HN63_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">A-kinase anchor protein 7 RI-RII subunit-binding domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="1">Binds cAMP-dependent protein kinase (PKA). Interacts with PRKCA; only the cytoplasmic form is capable of interacting with PRKCA.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HN63"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018731.1">
        <property type="protein sequence ID" value="ENSGAGP00000016415.1"/>
        <property type="gene ID" value="ENSGAGG00000012287.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034237">
        <property type="term" value="F:protein kinase A regulatory subunit binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010738">
        <property type="term" value="P:regulation of protein kinase A signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052641">
        <property type="entry name" value="AKAP7_isoform_gamma"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019511">
        <property type="entry name" value="AKAP7_RI-RII-bd_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15934:SF6">
        <property type="entry name" value="A-KINASE ANCHOR PROTEIN 7 ISOFORM GAMMA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15934">
        <property type="entry name" value="RNA 2',3'-CYCLIC PHOSPHODIESTERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10470">
        <property type="entry name" value="AKAP7_RIRII_bdg"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="A-kinase anchor protein 7 RI-RII subunit-binding" evidence="3">
        <location>
            <begin position="22"/>
            <end position="75"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="51"/>
            <end position="77"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00038702"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="Pfam" id="PF10470"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016415.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="77" mass="8599" checksum="D5095E59B05F4452" modified="2019-05-08" version="1">MGQLCCFPFSRNEEKISEKNGAEPDDAELVSLSKKLVENAVLKAVQQYLEETQNKNRQTDGSPVKTEEAANGNRNEK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN66</accession>
    <name>A0A452HN66_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Collagen alpha-1(XII) chain</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">Type XII collagen interacts with type I collagen-containing fibrils, the COL1 domain could be associated with the surface of the fibrils, and the COL2 and NC3 domains may be localized in the perifibrillar matrix.</text>
    </comment>
    <comment type="subunit">
        <text evidence="14">Trimer of identical chains each containing 190 kDa of non-triple-helical sequences.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="12">Belongs to the fibril-associated collagens with interrupted helices (FACIT) family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018725.1">
        <property type="protein sequence ID" value="ENSGAGP00000016409.1"/>
        <property type="gene ID" value="ENSGAGG00000011657.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005581">
        <property type="term" value="C:collagen trimer"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005614">
        <property type="term" value="C:interstitial matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005201">
        <property type="term" value="F:extracellular matrix structural constituent"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043394">
        <property type="term" value="F:proteoglycan binding"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007160">
        <property type="term" value="P:cell-matrix adhesion"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007044">
        <property type="term" value="P:cell-substrate junction assembly"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007507">
        <property type="term" value="P:heart development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009888">
        <property type="term" value="P:tissue development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
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        <property type="match status" value="17"/>
    </dbReference>
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        <property type="entry name" value="vWA_collagen_alphaI-XII-like"/>
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        <property type="entry name" value="collagen alpha-1(XII) chain isoform X1"/>
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        <property type="entry name" value="collagen alpha-1(XII) chain isoform X1"/>
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        <property type="entry name" value="ECM_Regulatory_Proteins"/>
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        <property type="entry name" value="Ig-like_fold"/>
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    <dbReference type="InterPro" id="IPR048287">
        <property type="entry name" value="TSPN-like_N"/>
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    <dbReference type="InterPro" id="IPR002035">
        <property type="entry name" value="VWF_A"/>
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        <property type="entry name" value="vWFA_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46708:SF7">
        <property type="entry name" value="FIBRONECTIN TYPE-III DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="TENASCIN"/>
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        <property type="match status" value="2"/>
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        <property type="entry name" value="VWFADOMAIN"/>
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    </dbReference>
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        <property type="match status" value="2"/>
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    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="10"/>
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    <dbReference type="PROSITE" id="PS50234">
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    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="8" id="KW-0176">Collagen</keyword>
    <keyword evidence="9" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11" id="KW-0379">Hydroxylation</keyword>
    <keyword evidence="7" id="KW-0654">Proteoglycan</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="1"/>
            <end position="84"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="87"/>
            <end position="181"/>
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        <location>
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            <end position="539"/>
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        <location>
            <begin position="541"/>
            <end position="630"/>
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    </feature>
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        <location>
            <begin position="650"/>
            <end position="822"/>
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    </feature>
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            <begin position="838"/>
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            <end position="1304"/>
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            <begin position="2413"/>
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        </location>
    </feature>
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        <location>
            <begin position="2463"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
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        <source>
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        <source>
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        <source>
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        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
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        <source>
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        <source>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016409.1"/>
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        <source>
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        </source>
    </evidence>
    <sequence length="2519" mass="272075" checksum="554C34BE6101E954" modified="2019-05-08" version="1">MITDETTDSFKVGWTPAPGNVLRYRVEYRPVTGRERKHVTVPANERSTTLLNLTPDTRYEVYVIPEYQSGPGNSLNGYAKTEEVLGTPRNLRVSDATTSTMKLSWSAAPGKVQRYLVTYTPVAGGETKEVTIRGDTTTTVLRDLEHGTRYALSVTALYASGAGDALSGQGETPEARGSPRNLRVSDATTSTMKLSWSAAPGKVERYRITYTPVAGGDTKEVTTRGDTTTTVLRDLEHGTRYALSVTALYASGAGDALPGQGETLEARGSPRNLRVSDATTSTMKLSWSAAPGKVQRYRITYTPVAGGEAKEVTIRGDTTTTVLRDLEQGTRYTLSVTALYASGAGDALPGQGETLEDRGSPRDLLIRDITDTTMGLSWTAAPGAVRRYRIVWRSLYDGQTGESTVPGNVINTVLENLQPETKYKISVIASYSSGDGAPLEGEATTEVSPNARRVRVDEETENSMRVTWQPAPGKVVSYRVVYRPVSGGRQVVAKVPATSTATVLKRLQPLTKYNISVIPMYKSGEGKHRQGEGTTASPYKPPRNLKTSDSTMSSFRVTWEPAPGEVRGYKITFHPTGEDRLGELIVGPYDNTVVLEELRAGTTYKVNVFGMFEGGESAPLVGQEMTTLSDVTVMPFLARGLECTTRAEADIVLLVDGSWSIGRPNFKTVRNFIARIVEVFDIGPDKVQIALAQYSGDPRTEWHLNTYGSKQSLLDAVANLPYKGGNTLTGMALSFILKDNFKPEVGLRPKARKIGVLITDGKSQDDIVAPSRRLRDEGVELYAIGIKNADENELKQIATDPDDTHVFNVADFSFLTNIVDDLTINLCNSVKGPGGLSSPSNLVTSEVTPRSFRVTWTAPVESVDRYRVEYYPVSGGPPQQFYVNRVETTTVLKNLKPETEYVVKVFSVVEDESSEPLSGTETTLPIPTIRNLNAYDIGSTSMRVRWEPLRGATGYRLLYEPLNATIPAVEKEMRVGSTVNDVQLVDLIPNAAYTLTIHAEFGDLSSDPLTTQEVTLPLAGAKSLRVRDITHSSMNVIWDPAPGKVRKYLLRYKTTLGDEPKEVEVDKSKTSTVLPSLLSQTLYDLQLVAVYDEGESIPVAAKATTLPVPAPVNLRLTEVTKQSFRGTWDHGAPDVALYRITWGPYGGTEKQETILNGDENSLVFENLNPDTLYDVSVTAIYPDESESDDLIGSERTLPAIRPTTQAPKSGPRNLQVYNATSNSLTVKWDPASGRVQRYRITYQRTTGDGAEQSTMVGGRQNSVVLQKLQPDTPYTITVSSMYADGEGGRMTGRGKTKPLNTVKNLRVYDPTTSTLSVRWDHAEGSPRQYKVYYVPTTGGAEEMTTVPGNTNYIILRSLQPDTPYKITVVPTYPEGDGGRVSDTGRTLVRGAARNIQVYNPTPNTLNVRWEPAPGPVQQYRVVYAPLVGPRPSESIVVPANTRDVVLERLTPDTPYSVNVVALYADGEGNQSSGQGRTTPRSGPRNLKVFDPTTNSLSVQWDHADGPVQQYRIIYSPTVGDPIDEYTTVPGRRNNVILQPLHSDTPYKITVVAVYDDGDGGQLTGNGRTVGLLPPQNIHISDEWYTRFRVSWDPSPSPVLGYKLAYKPVGSAEPPMEVFVGEVTSYTLHNLSPSTIYDVNVYAQYDSGLSAPLIDQGTTLYLNVTDLTSYKVGWDTFCIRWSPHRSASSYRLKLNPSDGSRGQEITVRGTETSHCFTGLSPDTEYNATVFVQTPNLEGPPISLKERTFIKPTEAPTQPPPPPPPPTIPPARDVCKGAKADIVFLTDASWSIGDDNFNKVVKFVFNTVGAFDLINPAGIQVSFVQYSDDAKSEFKLNTYDDKAQALGALQNIRYKGGNTRTGKALTFINEKVLTWESGMRRGVPKVLVVVTDGRSQDEVKKAAAIIQHSGFSIFVVGVADVDYHELSKIASKPSERHVFIVDDFDAFEKIEENLLTFICETATSTCPLIYLEGYTSPGFKMLESYNLTEKHFASVQGVSLESGSFPSHVAYRLHRNAFISQPASEIHPDGLPHAYTIILLFRLLSETPNEPFAIWQITDRDFKPQVGVILDPSTKLLSFFNKDTRGEVQTVTFDDNEVKKIFYGSFHKVHIVVTSTSVKIYIDCSEVMEKPIKESGNITTDGFEILGKLVKGDRKSATFEIQSFDIVCNAVWTSRDRCCDLPSMRDEAKCPALPNACTCTQDSVGPPGPPGPAGGPGTKGPRGERGLTGTAGPPGLPGEAGPPGPQGPPGPQGINGLSIPGEPGRQGMKGDAGEPGLPGRSGTPGLSGPPGPVGPPGDRGFTGKDGATGPRGPPGSAGTPGVPGVAGPSGKPGKPGDRGPQGPPGMKGEKGDRGDIASQNMMRAVARQVCEQLINGQMNRFNHMLNQIPNDYYANRNQPGPPGPPGPPGSAGARGEPGPGGRPGFPGTPGDQGPPGERGLPGEKGERGIGSQGPRGLPGPPGPPGESRTGPPGSTGSRGPPGPPGRPGNGGIRGPPGPPGYCDSSQCSSIAYNGQGFPEQG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN67</accession>
    <name>A0A452HN67_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">C2H2-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HN67"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018712.1">
        <property type="protein sequence ID" value="ENSGAGP00000016397.1"/>
        <property type="gene ID" value="ENSGAGG00000012269.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005654">
        <property type="term" value="C:nucleoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001227">
        <property type="term" value="F:DNA-binding transcription repressor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042802">
        <property type="term" value="F:identical protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001817">
        <property type="term" value="P:regulation of cytokine production"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002682">
        <property type="term" value="P:regulation of immune system process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000508">
        <property type="entry name" value="Myeloid zinc finger 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001155">
        <property type="entry name" value="Zinc finger 30C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000464">
        <property type="entry name" value="Zinc finger and SCAN domain containing 25"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000478">
        <property type="entry name" value="Zinc finger protein 133"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002716">
        <property type="entry name" value="Zinc finger protein 212"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002343">
        <property type="entry name" value="Zinc finger protein 33A"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001343">
        <property type="entry name" value="Zinc finger protein 568"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000145">
        <property type="entry name" value="Zinc finger protein 574"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000624">
        <property type="entry name" value="zinc finger protein 697"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="12"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24399:SF54">
        <property type="entry name" value="GASTRULA ZINC FINGER PROTEIN XLCGF26.1-LIKE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24399">
        <property type="entry name" value="ZINC FINGER AND BTB DOMAIN-CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="12"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="12"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="9" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 11" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="25"/>
            <end position="56"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="57"/>
            <end position="84"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="85"/>
            <end position="112"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="113"/>
            <end position="140"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="218"/>
            <end position="245"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="246"/>
            <end position="273"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="274"/>
            <end position="301"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="302"/>
            <end position="329"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="330"/>
            <end position="357"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="358"/>
            <end position="385"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="386"/>
            <end position="413"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="414"/>
            <end position="441"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="134"/>
            <end position="171"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="144"/>
            <end position="165"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016397.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="441" mass="50377" checksum="151E73C370CD49FF" modified="2019-05-08" version="1">MNLAHCLFPVEIAGPSPLPAGARPYVCADCGKSFRQSQRQVSNLYTHWRTHTGERPYPCAYCGKSFSLKHDLTKHQRVHTGERPYPCTDCNKSFSQKQDLTKHQRVHTGERPYPCAQCGKSFSNVSNLLAHQRTHLGQRLPRIPAEKEPDSPRESEKCSRKHAESSSEVGAKAHGKVPSCSACEQSLKGKIFFQSLGSFLRKKDLARHQGVHTGDRFYSCLDCRRSFSQMAGLTRHQKTHLGDRPFPCMDCGKSFCLRQDLMSHQRTHTGERPFLCSQCEKGFSQLAHLAAHQRTHAAERPYPCGTCEKRFKQSQDLRKHQRTHTGERPYPCTECGKCFSHVSNLNAHRKVHMGARPHTCKDCGKGFRKRQDLLRHERIHTGESPFSCSHCEKSFRQKKDLNRHRSVHTGERPYPCLECGKSFRLKHTLVRHQATHPRLSP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN71</accession>
    <name>A0A452HN71_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="15">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
            <location evidence="1">Basement membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="12">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018717.1">
        <property type="protein sequence ID" value="ENSGAGP00000016402.1"/>
        <property type="gene ID" value="ENSGAGG00000012074.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005604">
        <property type="term" value="C:basement membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009887">
        <property type="term" value="P:animal organ morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009888">
        <property type="term" value="P:tissue development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00055">
        <property type="entry name" value="EGF_Lam"/>
        <property type="match status" value="12"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000083">
        <property type="entry name" value="Laminin subunit alpha"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000090">
        <property type="entry name" value="laminin subunit alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000388">
        <property type="entry name" value="Laminin subunit alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000069">
        <property type="entry name" value="Laminin subunit alpha 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000034">
        <property type="entry name" value="Laminin subunit alpha 3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000084">
        <property type="entry name" value="Laminin subunit alpha 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000405">
        <property type="entry name" value="Laminin subunit alpha 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000430">
        <property type="entry name" value="Laminin subunit alpha 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.260:FF:000444">
        <property type="entry name" value="Laminin subunit alpha 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.260">
        <property type="entry name" value="Galactose-binding domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.170.300.10">
        <property type="entry name" value="Tie2 ligand-binding domain superfamily"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050440">
        <property type="entry name" value="Laminin/Netrin_ECM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008211">
        <property type="entry name" value="Laminin_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002049">
        <property type="entry name" value="LE_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056863">
        <property type="entry name" value="LMN_ATRN_NET-like_EGF"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10574:SF406">
        <property type="entry name" value="LAMININ SUBUNIT ALPHA 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10574">
        <property type="entry name" value="NETRIN/LAMININ-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00053">
        <property type="entry name" value="EGF_laminin"/>
        <property type="match status" value="12"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24973">
        <property type="entry name" value="EGF_LMN_ATRN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00055">
        <property type="entry name" value="Laminin_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00011">
        <property type="entry name" value="EGFLAMININ"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SMART" id="SM00180">
        <property type="entry name" value="EGF_Lam"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="SMART" id="SM00136">
        <property type="entry name" value="LamNT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="12"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01248">
        <property type="entry name" value="EGF_LAM_1"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50027">
        <property type="entry name" value="EGF_LAM_2"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51117">
        <property type="entry name" value="LAMININ_NTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0084">Basement membrane</keyword>
    <keyword evidence="7" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="8" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="9 12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11 12" id="KW-0424">Laminin EGF-like domain</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <feature type="domain" description="Laminin N-terminal" evidence="14">
        <location>
            <begin position="1"/>
            <end position="200"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="13">
        <location>
            <begin position="330"/>
            <end position="374"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="13">
        <location>
            <begin position="396"/>
            <end position="442"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="13">
        <location>
            <begin position="443"/>
            <end position="488"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="13">
        <location>
            <begin position="489"/>
            <end position="538"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="13">
        <location>
            <begin position="552"/>
            <end position="616"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="13">
        <location>
            <begin position="617"/>
            <end position="669"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="13">
        <location>
            <begin position="1279"/>
            <end position="1324"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="13">
        <location>
            <begin position="1369"/>
            <end position="1417"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="13">
        <location>
            <begin position="1418"/>
            <end position="1468"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="350"/>
            <end position="359"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="396"/>
            <end position="408"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="419"/>
            <end position="428"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="443"/>
            <end position="455"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="445"/>
            <end position="462"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="464"/>
            <end position="473"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="489"/>
            <end position="501"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="509"/>
            <end position="518"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="587"/>
            <end position="596"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="640"/>
            <end position="649"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="1279"/>
            <end position="1291"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="1281"/>
            <end position="1298"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="1300"/>
            <end position="1309"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="1393"/>
            <end position="1402"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="1418"/>
            <end position="1430"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="1420"/>
            <end position="1437"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="1439"/>
            <end position="1448"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004302"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022869"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023292"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00460"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50027"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS51117"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016402.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1560" mass="171770" checksum="120CCE602382853F" modified="2019-05-08" version="1">PPTHYPHFSSVLATAQSPSSPRSLGRLSVMSGLYCKGVRICCLCLPMSCPILFHVAYVLIKFANSPRPDLWVLERSTDFGLTYEPWQYFASSRRDCIEKFGPNTVKRITKDDDTICTTEYSRIVPLENGEIVVSLVNGRPGAMNFSYSPLLRNFTKATNIRLRFLQTNTLLGHLMGKALRDPTVTRRYYYSIKDISIGGRCVCHGHADVCDAKDPTDPYRLQCDCQHNTCGGSCDRCCPGFNQLPWKPATTDSANECQPCNCNGHAYDCYYDPEVDRHKASKNYDNQYKGGGVCIECQHHTTGINCERCIPGYYKSPDHSIDSPYICYRCNCDSEFTDGTCEDLTGRCYCKPNYTGELCNACAEGYTDFPHCYPVPAFSHNDTGEQVLPAGQIINCDCNAAGTEGNACRKDSQRGMCVCKPNFQGAQCDQCAPGHYGPGCQPCQCSGPGQYDGTCDSETGQCLCRSGFEGYSCNQCAPGYFSYPLCQLCGCSAVGTLPGGCDSGGRCFCRPEFDGPHCDQCSMGYHSYPRCQACSCDPRGSVDNNCSPAGQCRCHSNYAGHTCSQCAPGFYGYPSCTSPGILGSCVCRAYVEGPACDRCKPLYWNLTPDNPYGCTSCQCDTKGTISGVAECQQGNGQCFCKPSICGQLCSTCKDGYFNLDHANYFGCQGCRCDVGGSIGPACEARTGACHCRQNTQGPTCTLPTRDHYFPDLHHLKFELEEGTTLDGRAVRFGYNPLEFENFSWRGYAQMSPIQPKVVVTLNVTSPDLFRIVFRYVNRGPANVEGRVSVLEEGKFNVCGNCTEQTKQIIFAPSTEPAFVTVPQNSFGEPFVLNPSVWSLIVEAEGILLDYLVLLPSSYYEAPILQLKVTEACTYSPAPEQASQNCLLYKYLSVDSFPSASGGDMVCRLDNSLPRPCPMVPITPSHPQMALCSGSDVEVQLRLVVPQPGRYVLLVEYANKDAMQRASVAISSLQLATQQGSLTFHTCDYSFLCRGIVLDSQNRLAAFELTTEGTIRFAADQAHFFLYKVYLIPYGQFTMEFVEPKVHCISLHGTFSPSSSSCIPSRFMKLSQSIVLEAGQALSIPPDFPLTHVFTVAPTGIPTEPLPRPPTAVDSSAELILLQPSQVMKTGAGRIQTLGRYAFIMHYYQPNHPTFPVEVLINGGRIWQGQAKAAFCPHGYGCRSVVVSENQVILDVTDNELTVIVRVPAGRQLWLDYILVVPEDSYSSHYLLEEPLDKSYEFISSCGVNSFYINPTTSSKFCRDSAISLSLFYNNGAQSCRCHEAGAIGGQCELFGGQCACKSNIIGRECSRCATGYWGFPNCRPCDCGTRLCDEVTGQCICPPHTIKPECVVCEPQTFGCHALIGCEDCNCSRTGVQELAEPGCDMDSGQCRCKPNIIGRRCDVCAPGYYHYPHCRRCYCHQAGTEVSVCDPVTGQCHCKENVEGVRCDQCRLGTFSLDANNPKGCTRCFCFGATDRCHSAEKYRVEFINMNGWVLLSGDRQEVATSLRLEEQLIHADLKNVPEAYQELYWHAPSSYLGDRRSSPSSPHQCICVRWSWRL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN72</accession>
    <name>A0A452HN72_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">A-kinase anchor protein 7-like phosphoesterase domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="5">Binds cAMP-dependent protein kinase (PKA). Interacts with PRKCA; only the cytoplasmic form is capable of interacting with PRKCA.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HN72"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018730.1">
        <property type="protein sequence ID" value="ENSGAGP00000016414.1"/>
        <property type="gene ID" value="ENSGAGG00000012287.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034237">
        <property type="term" value="F:protein kinase A regulatory subunit binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010738">
        <property type="term" value="P:regulation of protein kinase A signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1140.10:FF:000004">
        <property type="entry name" value="A-kinase anchoring protein 7 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1140.10">
        <property type="entry name" value="Cyclic phosphodiesterase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019510">
        <property type="entry name" value="AKAP7-like_phosphoesterase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052641">
        <property type="entry name" value="AKAP7_isoform_gamma"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009097">
        <property type="entry name" value="Cyclic_Pdiesterase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15934:SF6">
        <property type="entry name" value="A-KINASE ANCHOR PROTEIN 7 ISOFORM GAMMA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15934">
        <property type="entry name" value="RNA 2',3'-CYCLIC PHOSPHODIESTERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10469">
        <property type="entry name" value="AKAP7_NLS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55144">
        <property type="entry name" value="LigT-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0539">Nucleus</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="A-kinase anchor protein 7-like phosphoesterase" evidence="7">
        <location>
            <begin position="100"/>
            <end position="295"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="80"/>
            <end position="101"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00038702"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF10469"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016414.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="354" mass="40751" checksum="6B31A5BB48E143B0" modified="2019-05-08" version="1">MLICMLHSLLPLFEDGLENMQEKKIIRKKKKRTKIRKGSQTPPDSMDYLMAEIPFASTDTGDEFGTTSTSEVCFKKRRKRTIRSKTEDDDQTKKKKQDQPNYFISLPITNPKITRGIQAVQDIVVQKDHRLSQAMVRCGSLHITLLVMHLSSEEEVGTAVGALLESKDAVEELLQGKHLDLSFQGIDDYRNEVIFVKFAASDHTAALTEISETMKKIFQEKGILAGENRAFKPHLTFMKLSKSPRLRKQVRKIDPHLYENFKSHYFGDETLYRLDLCSMLKKKQPNGYYHCESSIMVGKSHESDLIKKALYRESLALFSKVNKIKELLSNPETRMKINRELFQDRQTAVIKKSC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN73</accession>
    <name>A0A452HN73_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Anillin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="3">Required for cytokinesis. Essential for the structural integrity of the cleavage furrow and for completion of cleavage furrow ingression. Plays a role in bleb assembly during metaphase and anaphase of mitosis. May play a significant role in podocyte cell migration.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell projection</location>
            <location evidence="2">Bleb</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HN73"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018741.1">
        <property type="protein sequence ID" value="ENSGAGP00000016425.1"/>
        <property type="gene ID" value="ENSGAGG00000012291.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005826">
        <property type="term" value="C:actomyosin contractile ring"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032059">
        <property type="term" value="C:bleb"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000915">
        <property type="term" value="P:actomyosin contractile ring assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000281">
        <property type="term" value="P:mitotic cytokinesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031106">
        <property type="term" value="P:septin ring organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd01263">
        <property type="entry name" value="PH_anillin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000111">
        <property type="entry name" value="anillin isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012966">
        <property type="entry name" value="AHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031970">
        <property type="entry name" value="Anillin_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051364">
        <property type="entry name" value="Cytokinesis/Rho-signaling"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037840">
        <property type="entry name" value="PH_Anillin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21538:SF27">
        <property type="entry name" value="ANILLIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21538">
        <property type="entry name" value="ANILLIN/RHOTEKIN RTKN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08174">
        <property type="entry name" value="Anillin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16018">
        <property type="entry name" value="Anillin_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00169">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00233">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="PH" evidence="6">
        <location>
            <begin position="923"/>
            <end position="1047"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="1"/>
            <end position="93"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="122"/>
            <end position="210"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="240"/>
            <end position="272"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="293"/>
            <end position="327"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="405"/>
            <end position="436"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="548"/>
            <end position="581"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="700"/>
            <end position="723"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="57"/>
            <end position="70"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="82"/>
            <end position="91"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="192"/>
            <end position="210"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="299"/>
            <end position="317"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="407"/>
            <end position="421"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="423"/>
            <end position="436"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="5">
        <location>
            <begin position="548"/>
            <end position="557"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00043945"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00057106"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00071355"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016425.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1064" mass="118182" checksum="8A0DA25ECB287EE8" modified="2019-05-08" version="1">MAERPTAGMRPMMQTKRTREPLSETSNQSPLPSEEVKSSTKPSPSKRLCPDSMEVPVSSSENRQPVNSMSMKHLPTEMAEQPQATTKNRAPSVQPVPLLVEKEEQNLKSEVPVALTVKTRMQKLAQQRRCWDDNDPSESSPLPPLLSKEQAVSPPKQPSLASDTPVGRKGRLANLAATIGSWENDLSHPSAKRNNTQEQPGTTCLSKLSTTSEASARINSGSVKQEAASCSQRAVEVSVNKSATSKILGPNYLAKSSRVTSPSPKEPEVPQLLKKLCSPKKIEELKPTVKPILAQPVQSKEEPIKRTHVQLEHKDKPTTPGGSGIKPFLERFGERLQEHNTQSPTTTTTGYRTPIITPNTKTIQERLFKQNETSSTASLAQQLKQEREKELACIRGRFDKGSLWSAERGENSKSKHPEIKMESQAQASSKQLTSSDTTSLVAAEKTTASEIPVKSLSSEFSDAVCEIEISVNDKMNSSKVINDIFDEVLQEDGLDIEKLKKEMRINLEAESDDEQEESLNISSMSLLTPLAESVGVVSPEVFVSPSRSVAESSSISDESPKSGKFQRTRVPGAESGDSIGSLSEDHNLLYSIDAYRSQRFKETDRPSIKQVIVRKEDVASKLAVKRNGTSDQVNIKQKMQELNNEISLQQTVIHQASQALNCCIDEEHGKGSQEEAEAERLLLLATEKRKALLEELNKLKNEGPQGKKNKAASTPREFAPSRGSVTLSEMRLPLKADFVCSTVQKPEATNYYYIILLRSGAENMVATPVASTANSLNGDALTFTTTFTLQDVSNDFEINVEVYHLVQRKDGTGTTDKRKKTNKSKVITPKRLLTSITTKSTLLTPVMASPSGPNAVRTSNFVLVGSHKISLSSIGNSKFTLDKINYGGKERELLDFMFQDKVPFLSPLEGHIHLKLKCQVDSFVEEKGFLTMFEDVSGFGAWHRRWCVLSGNCISYWTYPDDEKRKHPMGRINLANCTNRQIEPANREFCARPNTFELITVRPQREDDKETLVSQCKDTLCVTKNWLSADTKEERNLWMQKLNQVLVDLRMWQPDACYKPIGKP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN76</accession>
    <name>A0A452HN76_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">phospholipase A2</fullName>
            <ecNumber evidence="1">3.1.1.4</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="6">
            <text>a 1,2-diacyl-sn-glycero-3-phosphocholine + H2O = a 1-acyl-sn-glycero-3-phosphocholine + a fatty acid + H(+)</text>
            <dbReference type="Rhea" id="RHEA:15801"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:28868"/>
            <dbReference type="ChEBI" id="CHEBI:57643"/>
            <dbReference type="ChEBI" id="CHEBI:58168"/>
            <dbReference type="EC" id="3.1.1.4"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="6">
            <dbReference type="Rhea" id="RHEA:15802"/>
        </physiologicalReaction>
    </comment>
    <dbReference type="EC" id="3.1.1.4" evidence="1"/>
    <dbReference type="AlphaFoldDB" id="A0A452HN76"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018735.1">
        <property type="protein sequence ID" value="ENSGAGP00000016419.1"/>
        <property type="gene ID" value="ENSGAGG00000011814.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0047499">
        <property type="term" value="F:calcium-independent phospholipase A2 activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0052816">
        <property type="term" value="F:long-chain fatty acyl-CoA hydrolase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035965">
        <property type="term" value="P:cardiolipin acyl-chain remodeling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016042">
        <property type="term" value="P:lipid catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:2000304">
        <property type="term" value="P:positive regulation of ceramide biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07212">
        <property type="entry name" value="Pat_PNPLA9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000338">
        <property type="entry name" value="85/88 kDa calcium-independent phospholipase A2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.1090.10">
        <property type="entry name" value="Cytosolic phospholipase A2 catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016035">
        <property type="entry name" value="Acyl_Trfase/lysoPLipase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047148">
        <property type="entry name" value="PLPL9"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002641">
        <property type="entry name" value="PNPLA_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24139:SF34">
        <property type="entry name" value="85_88 KDA CALCIUM-INDEPENDENT PHOSPHOLIPASE A2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24139">
        <property type="entry name" value="CALCIUM-INDEPENDENT PHOSPHOLIPASE A2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01734">
        <property type="entry name" value="Patatin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01415">
        <property type="entry name" value="ANKYRIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52151">
        <property type="entry name" value="FabD/lysophospholipase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51635">
        <property type="entry name" value="PNPLA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4 7" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="3 8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="8" id="KW-0442">Lipid degradation</keyword>
    <keyword evidence="5 8" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <feature type="repeat" description="ANK" evidence="7">
        <location>
            <begin position="151"/>
            <end position="183"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="7">
        <location>
            <begin position="218"/>
            <end position="250"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="7">
        <location>
            <begin position="348"/>
            <end position="380"/>
        </location>
    </feature>
    <feature type="domain" description="PNPLA" evidence="9">
        <location>
            <begin position="418"/>
            <end position="602"/>
        </location>
    </feature>
    <feature type="short sequence motif" description="GXGXXG" evidence="8">
        <location>
            <begin position="422"/>
            <end position="427"/>
        </location>
    </feature>
    <feature type="short sequence motif" description="GXSXG" evidence="8">
        <location>
            <begin position="454"/>
            <end position="458"/>
        </location>
    </feature>
    <feature type="short sequence motif" description="DGA/G" evidence="8">
        <location>
            <begin position="589"/>
            <end position="591"/>
        </location>
    </feature>
    <feature type="active site" description="Nucleophile" evidence="8">
        <location>
            <position position="456"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="8">
        <location>
            <position position="589"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00013278"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023422"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01161"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51635"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016419.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="743" mass="83533" checksum="16A3E3000AFBA0ED" modified="2019-05-08" version="1">MQFFGRLVNTINSYSNLFSNPYRVKEVPLAEYVAHTRLRENDRVILYKSKNARSWDCLLVNPQSHLVAFRLFQLDNETDAIERFEQYARRLRPFYESSPQPLQLEVLQQLSDCFRNHPYWSLAHVAVETELPGSFQHNRILSCVNSTECEDGCTPLQLACRKGNMECLCELVECHARLDVTDKNGETVFHYAVRGSNPEIVKLLGKKPSVGLNHLSNQGQTPLHLACQLGKEDVVLALLKCHAKCNVMGTLGYPIHTAVKYSQKGCVQALLEVDVNQVQFKDPRYGAAVLHWAKNAEMTRMLIEYGCDVNSVSKTADMALHIMVKRGRFDSAMVLLTHGALTNAKGQDGNTPLHLAMKQDHLETIKALIVFGADVEIPNEFGETPGLVAARTSKGDYTACPVLSEAKERSRSLYDRLLCLDGGGIRGLVLIQLLIAVEKAAGRPIRELFDWIAGTSTGGILALAIVHGKPVDYMRCLYFRMKNEVFRGSRPYESEPLDEFLKREFGEHTKMTDVQKPKVIVTGTLCDRQPAELHLFRNYKAPATKNKPEVKTTDPFKPLTKPEDQLVWRAARCSGAAPTYFRPIGRFLDGGLLANNPTLDAMTEIHEYNKSLSQKGQSQQVRKLGVVVSLGTGRPPQVPVRSVDVFRPSNPWELAKTVFGAKELGKMVVDCCTDADGPAVDRARAWCEMTDTRYFRFSPQLQSDVMLDEVNDAILVNALWDTQLYICQQQEQFKQLVQELLTR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN78</accession>
    <name>A0A452HN78_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">BZIP domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum membrane</location>
            <topology evidence="1">Single-pass type II membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the bZIP family. ATF subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HN78"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018748.1">
        <property type="protein sequence ID" value="ENSGAGP00000016430.1"/>
        <property type="gene ID" value="ENSGAGG00000012285.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd14689">
        <property type="entry name" value="bZIP_CREB3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.170:FF:000042">
        <property type="entry name" value="Cyclic AMP-responsive element-binding protein 3-like protein 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.170">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004827">
        <property type="entry name" value="bZIP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046347">
        <property type="entry name" value="bZIP_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051381">
        <property type="entry name" value="CREB_ATF_subfamily"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45996">
        <property type="entry name" value="AGAP001464-PB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45996:SF1">
        <property type="entry name" value="CYCLIC AMP-RESPONSIVE ELEMENT-BINDING PROTEIN 3-LIKE PROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00170">
        <property type="entry name" value="bZIP_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00338">
        <property type="entry name" value="BRLZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57959">
        <property type="entry name" value="Leucine zipper domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50217">
        <property type="entry name" value="BZIP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00036">
        <property type="entry name" value="BZIP_BASIC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0010">Activator</keyword>
    <keyword evidence="14" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="8" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="4" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9" id="KW-0472">Membrane</keyword>
    <keyword evidence="13" id="KW-0539">Nucleus</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0735">Signal-anchor</keyword>
    <keyword evidence="11" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="3" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6" id="KW-1133">Transmembrane helix</keyword>
    <feature type="domain" description="BZIP" evidence="16">
        <location>
            <begin position="218"/>
            <end position="281"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="56"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="388"/>
            <end position="443"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="14">
        <location>
            <begin position="243"/>
            <end position="284"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="15">
        <location>
            <begin position="56"/>
            <end position="72"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004648"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009050"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50217"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016430.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="443" mass="48283" checksum="E625007088F07B44" modified="2019-05-08" version="1">WWGDYSPGEAGCQMGSIDSLELLDLLFDRQDGILRNVEFGGHTLRGAQENEDFLSSILGSGDSGSDSPIWSPAASDSGISEDPNSDQLDSPQHYMPTGSPGPSEPAYPYQEASQAQPAPSLPGADLREAEVSIDLGEFSHDLPAPALIAASCTLTVKDLLLSSSCEMQQQQLMTPALLRQSHGHCQELVLTEDEKKLLAKEGVTLPTQLPLTKYEERVLKKIRRKIRNKQSAQESRKKKKEYIDGLESRMSACTAQNQELQRKVLHLEKQNSSLLEQLRKLQTLVVQSTNKAAQTGTCIAVLLLSFALIVFPSISPFTPNQAEPEGDFGPVRVFSRSLHNDASSRVVYALRKDAAESPQEPENPVWTEYAGEAQSVLRKFLSSHVLTSRPQAEAPQANGTGALPQDGHSDLEGLSHDGPIAASLAWTEPSQPSQVQLEPAEEL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN81</accession>
    <name>A0A452HN81_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">A-kinase anchor protein 7-like phosphoesterase domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="5">Binds cAMP-dependent protein kinase (PKA). Interacts with PRKCA; only the cytoplasmic form is capable of interacting with PRKCA.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HN81"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018728.1">
        <property type="protein sequence ID" value="ENSGAGP00000016412.1"/>
        <property type="gene ID" value="ENSGAGG00000012287.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034237">
        <property type="term" value="F:protein kinase A regulatory subunit binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010738">
        <property type="term" value="P:regulation of protein kinase A signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1140.10:FF:000004">
        <property type="entry name" value="A-kinase anchoring protein 7 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1140.10">
        <property type="entry name" value="Cyclic phosphodiesterase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019510">
        <property type="entry name" value="AKAP7-like_phosphoesterase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052641">
        <property type="entry name" value="AKAP7_isoform_gamma"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009097">
        <property type="entry name" value="Cyclic_Pdiesterase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15934:SF6">
        <property type="entry name" value="A-KINASE ANCHOR PROTEIN 7 ISOFORM GAMMA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15934">
        <property type="entry name" value="RNA 2',3'-CYCLIC PHOSPHODIESTERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10469">
        <property type="entry name" value="AKAP7_NLS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55144">
        <property type="entry name" value="LigT-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0539">Nucleus</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="A-kinase anchor protein 7-like phosphoesterase" evidence="7">
        <location>
            <begin position="174"/>
            <end position="369"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="53"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="154"/>
            <end position="175"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="66"/>
            <end position="76"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="85"/>
            <end position="98"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="6">
        <location>
            <begin position="99"/>
            <end position="111"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00038702"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF10469"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016412.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="428" mass="47944" checksum="2DD11C0A3AC6458D" modified="2019-05-08" version="1">MRRLLSTLRRHHARLLAAAAELPAPGHRAPPPGWGERRAAAAAWALGSGRSMAQCGGVPAPPPAPEEAQGSGAASGDPPPGAGESLEHDGLENMQEKKIIRKKKKRTKIRKGSQTPPDSMDYLMAEIPFASTDTGDEFGTTSTSEVCFKKRRKRTIRSKTEDDDQTKKKKQDQPNYFISLPITNPKITRGIQAVQDIVVQKDHRLSQAMVRCGSLHITLLVMHLSSEEEVGTAVGALLESKDAVEELLQGKHLDLSFQGIDDYRNEVIFVKFAASDHTAALTEISETMKKIFQEKGILAGENRAFKPHLTFMKLSKSPRLRKQVRKIDPHLYENFKSHYFGDETLYRLDLCSMLKKKQPNGYYHCESSIMVGKSHESDLIKKALYRESLALFSKVNKIKELLSNPETRMKINRELFQDRQTAVIKKSC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPR7</accession>
    <name>A0A452IPR7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Olfactomedin-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="3">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPR7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034165.1">
        <property type="protein sequence ID" value="ENSGAGP00000030093.1"/>
        <property type="gene ID" value="ENSGAGG00000021706.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003112">
        <property type="entry name" value="Olfac-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050605">
        <property type="entry name" value="Olfactomedin-like_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23192:SF7">
        <property type="entry name" value="OLFACTOMEDIN-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23192">
        <property type="entry name" value="OLFACTOMEDIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02191">
        <property type="entry name" value="OLF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00284">
        <property type="entry name" value="OLF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51132">
        <property type="entry name" value="OLF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <feature type="domain" description="Olfactomedin-like" evidence="5">
        <location>
            <begin position="193"/>
            <end position="457"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="4">
        <location>
            <begin position="70"/>
            <end position="154"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00446"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS51132"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030093.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="460" mass="53128" checksum="87D26250D236FBB7" modified="2019-05-08" version="1">NPPSLHVDNLLSETCFLQPFANVTGSLNDHRVCQCSVYLPDPVFPVQKVEMLEITAQVLSEKFDRELTKVSQYTKAIEMYEQKIQNLTIKVEHMESTSVSYMELDFQLLKLEISEMERLVTQLKSTLVGSNVIVEQLYIEIKNLTLMVNDLESLDKNNILAIHREIVALQNRLKECEKHRNQTTKPSYFPPGSCGHGGIVNISQPYVVQLNWKGFSFKYGSWGRDYSLRTPQKDLYWIAPLNTDGRLLEYYRLHNSYDDLLLLKNARELKIPYGYGEGSGTTVYNNFMYYNIYNSREIGKLDLNTNTLTVRKTLPNAAYGNRFSYAGVGWQDMDFAVDESGLWVIYSTEDNTGNIMISKLNETTLDVLHTWNTRQYKPSISNAFIICGVLYATRPVNTRKEEIFYTYDTNTEQEGKISIIMDKMLETIQSINYNPSDQVLYVYNDGYLVKYNLIFQPVLH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPR9</accession>
    <name>A0A452IPR9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="9">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Lipid-anchor</topology>
            <topology evidence="1">GPI-anchor</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the neuritin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPR9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033883.1">
        <property type="protein sequence ID" value="ENSGAGP00000029842.1"/>
        <property type="gene ID" value="ENSGAGG00000021529.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098552">
        <property type="term" value="C:side of membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:1990138">
        <property type="term" value="P:neuron projection extension"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026144">
        <property type="entry name" value="Neuritin_fam"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15902:SF2">
        <property type="entry name" value="NEURITIN-LIKE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15902">
        <property type="entry name" value="NEURITIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15056">
        <property type="entry name" value="NRN1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="7" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="4" id="KW-0336">GPI-anchor</keyword>
    <keyword evidence="8" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="6" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004609"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008377"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022622"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029842.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="155" mass="16969" checksum="E4E6C09D9B67857E" modified="2019-05-08" version="1">MVRWPPSPPASAGQCCARPCPLDLVSTAGRRCDTIYKGFAECLISLGDSMDGQELDTICRSWDDFHTCANRVLSRCPDEAAAIWESLRQESRKIQFQGNLHDLCSARARPPGTVRGSAADETNKETLRGSARLPWPSLLATLPLLVLTSLLVPFV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPS2</accession>
    <name>A0A452IPS2_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="5">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IPS2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033963.1">
        <property type="protein sequence ID" value="ENSGAGP00000029907.1"/>
        <property type="gene ID" value="ENSGAGG00000021582.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003676">
        <property type="term" value="F:nucleic acid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="CDD" id="cd02642">
        <property type="entry name" value="R3H_encore_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1370.50:FF:000001">
        <property type="entry name" value="R3H domain-containing protein 2 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1370.50">
        <property type="entry name" value="R3H-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001374">
        <property type="entry name" value="R3H_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036867">
        <property type="entry name" value="R3H_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051937">
        <property type="entry name" value="R3H_domain_containing"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024771">
        <property type="entry name" value="SUZ"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15672:SF14">
        <property type="entry name" value="CAMP-REGULATED PHOSPHOPROTEIN 21"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15672">
        <property type="entry name" value="CAMP-REGULATED PHOSPHOPROTEIN 21 RELATED R3H DOMAIN CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01424">
        <property type="entry name" value="R3H"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00393">
        <property type="entry name" value="R3H"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF82708">
        <property type="entry name" value="R3H domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51061">
        <property type="entry name" value="R3H"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51673">
        <property type="entry name" value="SUZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="R3H" evidence="3">
        <location>
            <begin position="179"/>
            <end position="242"/>
        </location>
    </feature>
    <feature type="domain" description="SUZ" evidence="4">
        <location>
            <begin position="243"/>
            <end position="324"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1"/>
            <end position="81"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="110"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="312"/>
            <end position="435"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="465"/>
            <end position="529"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="566"/>
            <end position="644"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="1"/>
            <end position="10"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="44"/>
            <end position="57"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="2">
        <location>
            <begin position="65"/>
            <end position="77"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="115"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="320"/>
            <end position="331"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="332"/>
            <end position="341"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="342"/>
            <end position="354"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="382"/>
            <end position="404"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="405"/>
            <end position="414"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="421"/>
            <end position="435"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="491"/>
            <end position="515"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="516"/>
            <end position="529"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="566"/>
            <end position="582"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="593"/>
            <end position="618"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="2">
        <location>
            <begin position="629"/>
            <end position="644"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS51061"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS51673"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029907.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="744" mass="82080" checksum="A1E65762C68A243A" modified="2019-05-08" version="1">MSEQAKLNQETVEETAKEQESAISDATPENCIINKTESLEAEEKEEKLSDAKTDLQKRGVNQNQQKKRSKSGAKGKLVRSLAVCEESSPRLGAEGLHDNQESIQLQLLSFPSLQEEDKSSKDDSEKEKEKDKNKEKDKNCEKSKIRMLSKDCSQEYTDSTGIDLHEFLVNTLKNNPRDRMILLKMEQEIIDFISDNNNHYKKFPQMSSYQRMLVHRVAAYFGMDHNVDQTGKSVIINKTSNTRIPEQRFSEHLKDEKGEESQKRFILKRDNSSIDKEDNQSVCSQENLFVENSRLLEDNSICNDTHKKRQLFRGNRDSTGKASGSRQSSTENEMKWTDHQRPWSSTDSDSSNRNLKPAITKTASFGGITVLTRGDSSSSNRSTGKLSKTGSESSSSAGSSGSLSRTHQPLQSASIAPGVTASPSGNVSYTENGMNGQVAPSNTSYIILPLEAAGIPPGSILLNPHTGQPFVNPDGTPAIYNPPSGQQPIRSQLVGQSQQQPSSQQQQQVQQPQQQMASHLVTQPVQALQPSSQSVQYPAVSYPPQHLLPVSPTQQFPVRDDMTTQFSQMSLSRQSSGDNPESPSGPVYPSPIIQPSGQQTGYVMASPSQQLPPGSFTGSGPPVSQQVLQPPPPPQGYVQQPPPAQMPVYYYPSGQYPTSATQQYRPIASVQYNAQRSQQISQTAQQAGYQPVLTNQQQGFQGLMGMQQPPQSQNLMNNQQGNQVQGLMVQYPAMSSYQVLTSCT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPS3</accession>
    <name>A0A452IPS3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Rap guanine nucleotide exchange factor 6</fullName>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the RAPGEF2 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000034054.1">
        <property type="protein sequence ID" value="ENSGAGP00000029986.1"/>
        <property type="gene ID" value="ENSGAGG00000021645.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007264">
        <property type="term" value="P:small GTPase-mediated signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00038">
        <property type="entry name" value="CAP_ED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd06755">
        <property type="entry name" value="PDZ_RapGEF2_RapGEF6-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd01785">
        <property type="entry name" value="RA_PDZ-GEF1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00155">
        <property type="entry name" value="RasGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd06224">
        <property type="entry name" value="REM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.840.10:FF:000001">
        <property type="entry name" value="Rap guanine nucleotide exchange factor (GEF) 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000024">
        <property type="entry name" value="rap guanine nucleotide exchange factor 2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.870.10:FF:000001">
        <property type="entry name" value="rap guanine nucleotide exchange factor 2 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.10:FF:000019">
        <property type="entry name" value="rap guanine nucleotide exchange factor 6 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.42.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.10">
        <property type="entry name" value="Jelly Rolls"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.840.10">
        <property type="entry name" value="Ras guanine-nucleotide exchange factors catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.870.10">
        <property type="entry name" value="Son of sevenless (SoS) protein Chain: S domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000595">
        <property type="entry name" value="cNMP-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018490">
        <property type="entry name" value="cNMP-bd_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001478">
        <property type="entry name" value="PDZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036034">
        <property type="entry name" value="PDZ_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000159">
        <property type="entry name" value="RA_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000651">
        <property type="entry name" value="Ras-like_Gua-exchang_fac_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023578">
        <property type="entry name" value="Ras_GEF_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001895">
        <property type="entry name" value="RASGEF_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036964">
        <property type="entry name" value="RASGEF_cat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014710">
        <property type="entry name" value="RmlC-like_jellyroll"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029071">
        <property type="entry name" value="Ubiquitin-like_domsf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45161">
        <property type="entry name" value="CYTOSKELETON-ASSOCIATED PROTEIN 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45161:SF4">
        <property type="entry name" value="RAP GUANINE NUCLEOTIDE EXCHANGE FACTOR 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00027">
        <property type="entry name" value="cNMP_binding"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00595">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00788">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00617">
        <property type="entry name" value="RasGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00618">
        <property type="entry name" value="RasGEF_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00100">
        <property type="entry name" value="cNMP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00228">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00314">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00147">
        <property type="entry name" value="RasGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00229">
        <property type="entry name" value="RasGEFN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51206">
        <property type="entry name" value="cAMP-binding domain-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50156">
        <property type="entry name" value="PDZ domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48366">
        <property type="entry name" value="Ras GEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54236">
        <property type="entry name" value="Ubiquitin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50042">
        <property type="entry name" value="CNMP_BINDING_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50106">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50200">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50009">
        <property type="entry name" value="RASGEF_CAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50212">
        <property type="entry name" value="RASGEF_NTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7 9" id="KW-0344">Guanine-nucleotide releasing factor</keyword>
    <keyword evidence="8" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Cyclic nucleotide-binding" evidence="12">
        <location>
            <begin position="259"/>
            <end position="359"/>
        </location>
    </feature>
    <feature type="domain" description="N-terminal Ras-GEF" evidence="15">
        <location>
            <begin position="391"/>
            <end position="504"/>
        </location>
    </feature>
    <feature type="domain" description="PDZ" evidence="13">
        <location>
            <begin position="509"/>
            <end position="579"/>
        </location>
    </feature>
    <feature type="domain" description="Ras-associating" evidence="14">
        <location>
            <begin position="734"/>
            <end position="820"/>
        </location>
    </feature>
    <feature type="domain" description="Ras-GEF" evidence="11">
        <location>
            <begin position="845"/>
            <end position="1072"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="151"/>
            <end position="223"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1059"/>
            <end position="1085"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1190"/>
            <end position="1273"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1456"/>
            <end position="1477"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1490"/>
            <end position="1533"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="166"/>
            <end position="183"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="10">
        <location>
            <begin position="206"/>
            <end position="218"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="1060"/>
            <end position="1072"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="1219"/>
            <end position="1230"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="1244"/>
            <end position="1253"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="1254"/>
            <end position="1271"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="1467"/>
            <end position="1477"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="1512"/>
            <end position="1521"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00010829"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022658"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00168"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50009"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50106"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50200"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029986.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="18">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1533" mass="172305" checksum="5AC9731CF283DE19" modified="2019-05-08" version="1">MDLNTIYSYLHGMDILSHLREHQLRIMSSGARYERYKGNQVLFCSDTIARCWYILLSGSVLMKDSMFLPPCSFGKQSGGKRGCECIILEPSEMIVVENSKDNEENILQREVPLRQSRRRFRKINQRGERQTITDNVDASSYLSLPADLTKMHLTDNPHPQVTHVPSSQSGCSIASDSGSSSLSDIYQATESDMGDVDLTGLPEAPVDSEDEEEEDEDIDRTSDPLLGRDVVRECLEKEPADKTDDDIEQLLEFMHQLPAFANMTMSVRRELCSVMIFEVVEQAGAIILEDGQELDSWYVILNGTVEISHPDGKTESLCMGNSFGINPSLDKQYMNGVVRTKVDDCQFVCIAQQDYWRILNHVEKNTHKVEEEGEIVMVKEHRELDRSGTRKGHIVIKATPERLIMHLIEEHSIVDPTYIEDFLLTYRTFLSSPLEVGDKLLEWFKVDSLRDKVTRVVLLWVNNHFNDFEGDPAMTRFLEEFEKNLEDTKMKGHLRLLNIACAAKAKWRQITLQKASRDSPLFFCLVGGSDKGFGIFVETVETGSKAAEAGLKRGDQIMEVNGQNFENITFVKAVEILRNNTHLSITVKTNIFVFKELLSRIGQEKNGVPHIPKIAEKKSNRYSIPDMPGDMEQMFPCEKGNKKIKANTVSGGRNRIRKILDKTRFSILPPKLFSDGSVSQSQDDSIVGTRQCRHSLAIMPIPGSLSSSSPDLLQPTTSMLDFTNPSAVGFYYIPDQVIRVFKADQQSCYIIISKDTTAKEVVSHAVHEFNLIGAPETYSLCEVSVSPEGVIKQRRLPDQFSKLADRIQLNGRYYLKNNMETETLCSDEDAQELLRESQISLLQLSTIEVATQLSMRDFELFRNIEPTEYIDDLYKMDSKTGNAHLKQFEDVINQETFWVATEILTEPNQLKRMKIVKHFIKIALHCRECKNFNSMFAVISGLNLAPVARLRGTWEKLPSKYEKHLRDLQDLFDPSRNMAKYRNILSSQSMQPPIIPLFPVVKKDITFLHEGNDSKVDGLVNFEKLRMIAKEIRQVVRMTSANMDPAVMFRQRKKRWRSLGSLSQGSTNSNMLDVQGGAHKKRARRSSLLNAKKLYEDAQMARKVKQYLSNLNVETDEEKFQIMSLQCEPAYSTLTKNLNERKGAKSSEMSPIPMRSIGQTTKAHLHQQNRMSQVLQVPAVNLYPIRKKGPAKDHVTFSSPQKSLSLSEEVSSKKQAEDSISMASSLHSSPPASPQGSPHKVGNIQRSDNCGQMNLSGSSSSLASETSNKNSGQRSYGIGYALIPSTKSDNFSDSSHSEISSRSSIVSNCSADSMSAALQDERCLPQSLIVMDSAGGVERKDVPQPSAEYSQPTTGWSLTRPAIIRGLAVTSSMSNEEISHEHITVEAADSGRGSWTSCSSSSHDNFQNIQNQKSWDLLSAYRHTHLDGPIAEVEPTNCCSEETYLYSEAFSKNNRQSKASTEHDQSRQSWASSSSLSDTYETNYGTIKRRGLEHSTAEQTEVLDSKPATDANYKTVTSSTEKGLIEDEQVSAV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPS4</accession>
    <name>A0A452IPS4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="19">UDP-glucose glycoprotein glucosyltransferase 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Recognizes glycoproteins with minor folding defects. Reglucosylates single N-glycans near the misfolded part of the protein, thus providing quality control for protein folding in the endoplasmic reticulum. Reglucosylated proteins are recognized by calreticulin for recycling to the endoplasmic reticulum and refolding or degradation.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="10">
            <text>N(4)-(alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;6)]-alpha-D-Man-(1-&gt;6)]-beta-D-Man-(1-&gt;4)-beta-D-GlcNAc-(1-&gt;4)-beta-D-GlcNAc)-L-asparaginyl-[protein] (N-glucan mannose isomer 9A1,2,3B1,2,3) + UDP-alpha-D-glucose = N(4)-(alpha-D-Glc-(1-&gt;3)-alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;6)]-alpha-D-Man-(1-&gt;6)]-beta-D-Man-(1-&gt;4)-beta-D-GlcNAc-(1-&gt;4)-beta-D-GlcNAc)-L-asparaginyl-[protein] + UDP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:61304"/>
            <dbReference type="Rhea" id="RHEA-COMP:14356"/>
            <dbReference type="Rhea" id="RHEA-COMP:14357"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:58223"/>
            <dbReference type="ChEBI" id="CHEBI:58885"/>
            <dbReference type="ChEBI" id="CHEBI:59080"/>
            <dbReference type="ChEBI" id="CHEBI:139493"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </cofactor>
    </comment>
    <comment type="pathway">
        <text evidence="3">Protein modification; protein glycosylation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Endoplasmic reticulum lumen</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the glycosyltransferase 8 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000033890.1">
        <property type="protein sequence ID" value="ENSGAGP00000029847.1"/>
        <property type="gene ID" value="ENSGAGG00000021377.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00378"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005788">
        <property type="term" value="C:endoplasmic reticulum lumen"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003980">
        <property type="term" value="F:UDP-glucose:glycoprotein glucosyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051082">
        <property type="term" value="F:unfolded protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036503">
        <property type="term" value="P:ERAD pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0018279">
        <property type="term" value="P:protein N-linked glycosylation via asparagine"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.550.10">
        <property type="entry name" value="Spore Coat Polysaccharide Biosynthesis Protein SpsA, Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040497">
        <property type="entry name" value="Glyco_transf_24"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029044">
        <property type="entry name" value="Nucleotide-diphossugar_trans"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009448">
        <property type="entry name" value="UDP-g_GGtrans"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040693">
        <property type="entry name" value="UGGT_TRXL_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040694">
        <property type="entry name" value="UGGT_TRXL_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040692">
        <property type="entry name" value="UGGT_TRXL_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040525">
        <property type="entry name" value="UGGT_TRXL_4"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11226">
        <property type="entry name" value="UDP-GLUCOSE GLYCOPROTEIN:GLUCOSYLTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11226:SF3">
        <property type="entry name" value="UDP-GLUCOSE:GLYCOPROTEIN GLUCOSYLTRANSFERASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18404">
        <property type="entry name" value="Glyco_transf_24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18400">
        <property type="entry name" value="Thioredoxin_12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18401">
        <property type="entry name" value="Thioredoxin_13"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18402">
        <property type="entry name" value="Thioredoxin_14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18403">
        <property type="entry name" value="Thioredoxin_15"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06427">
        <property type="entry name" value="UDP-g_GGTase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53448">
        <property type="entry name" value="Nucleotide-diphospho-sugar transferases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="8" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="5" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="UGGT thioredoxin-like" evidence="12">
        <location>
            <begin position="43"/>
            <end position="224"/>
        </location>
    </feature>
    <feature type="domain" description="UGGT thioredoxin-like" evidence="13">
        <location>
            <begin position="298"/>
            <end position="429"/>
        </location>
    </feature>
    <feature type="domain" description="UGGT thioredoxin-like" evidence="14">
        <location>
            <begin position="437"/>
            <end position="685"/>
        </location>
    </feature>
    <feature type="domain" description="UDP-glucose:glycoprotein glucosyltransferase thioredoxin-like" evidence="15">
        <location>
            <begin position="715"/>
            <end position="941"/>
        </location>
    </feature>
    <feature type="domain" description="Glucosyltransferase 24 catalytic" evidence="16">
        <location>
            <begin position="1236"/>
            <end position="1343"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1372"/>
            <end position="1393"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001913"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004922"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00006351"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00045874"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00048456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF18400"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="Pfam" id="PF18401"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="Pfam" id="PF18402"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="Pfam" id="PF18403"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF18404"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029847.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="19">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1393" mass="158277" checksum="A57EA8EF22056C21" modified="2019-05-08" version="1">LNWRGFKEEPQERLKIGKHTLCSSLAKADSKAVTTSLTTKWLSTPLLLEASEFLAEEGQEKFWTFVEASQDIRSSDHGGTNYSSYHAMLKAACQNLSPLQQNLLKFSLSLRSYSATIQAFQQIAADEPPPVGCNSFFAVHGKKTCEFDTLEILLQTASERLKPFLFKGDHRFPLSNPESPVVILYAEIGTEEFSKFHNLLVSKASAGDITYILRHYVANPSKEKVYLSGYGVELAIKSTEYKAKDDTQVKGMEVNATMIGENDPIDEVQGFLFGKLRQLHPDLKEELKELRKHLIESTNEMAPLKVWQLQDLSFQTAARILAAPTVDALMVMKDLSQNFPTKARAITKTVVSSELRTEIEENQKYFKGTLGLQSGDSAFFINGLHIDLDTQDIFSLFDVLRNEARVMEGLHSLGIEGLSLHNVLKLNIQPSDSDYAVDIRSPAISWINNLEVDSRYNSWPSSVQELLRPTFPGVIRQIRKNFHNFVLIVDPSHESTAELLNVAEMFFSNHIPLRIGLVFVVNDSEDVDGLQDAGVALLRAFNYVAQEVDNNYAFQTLISIYNKVKTGDQLQVEHVVSVLEKQYPYVEMNSILGIDSVYDQNRKEGRGYYEQTGVGPLPIVLFNGMPFQKEQLDPDDLETITMQKILETTSLFQRAVYLGELSSDQDVVEYIMNQPNVVPRINSRILMSDREYLDLTATNNFFVDDYARFTFLNSKDKTAAVANSMTYLTKKGMSSKEIYDDSFVRPVTFWIVGDFDKPSGRQLLYDAIKHQKSSNHVRISMINNPSEDPTSKNTVITRAIWAALQTQTSNNAKNFITKMAKEETAKALEAGADITEFAVGGMDVNTFKEAFESPKVDFILSHAMYCRDVLKLQKGQRAVISNGRIIGPLEDNELFNQDDFHLLENIILKTSGQKIKSHIQQLGVEEDLASDLVMKVDALLSAQPKGEARIEYQFFEDRYSAVKLRPKEGETYFDVVAIVDPATRDAQRLAPLLMVLNQLINMNLRVFMNCQSKLSDMPLKSFYRYVLEPEISFTADDNFAPGPIAKFLDMPQTPLFTLNLNTPESWMVESVRTPYDLDNIYLEEVDNRVAAEYELEYLLLEGHCYDITTGQPPRGLQFTLGTSANPVIVDTIVMANLGYFQLKANPGAWVLRLRKGRSEDIYRIYSHDGTDSPPDANEVTVVLNNFKSKIIKVKVSLKTVLQEYKYSNEDYIFPDFRGFTGGQKNDEMKQDKDDVINIFSVASGHLYERFLRIMMLSVLKHTKTPVKFWFLKNYLSPMFKEFIPYMAEKYNFQYELVQYKWPRWLHQQTEKQRIIWGYKILFLDVLFPLVVDKFLFVDADQVSYLMALGSLAGITRSAYIALQQMWQGSPCPSLPTDWRGTEPSRQGASGRAR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPS5</accession>
    <name>A0A452IPS5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Nck-associated protein 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="6">Cell projection</location>
            <location evidence="6">Lamellipodium membrane</location>
            <topology evidence="6">Single-pass membrane protein</topology>
            <orientation evidence="6">Cytoplasmic side</orientation>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="7">Belongs to the HEM-1/HEM-2 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPS5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034009.1">
        <property type="protein sequence ID" value="ENSGAGP00000029950.1"/>
        <property type="gene ID" value="ENSGAGG00000019065.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031258">
        <property type="term" value="C:lamellipodium membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031209">
        <property type="term" value="C:SCAR complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030031">
        <property type="term" value="P:cell projection assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030866">
        <property type="term" value="P:cortical actin cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048812">
        <property type="term" value="P:neuron projection morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019137">
        <property type="entry name" value="Nck-associated_protein-1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12093">
        <property type="entry name" value="NCK-ASSOCIATED PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12093:SF11">
        <property type="entry name" value="NCK-ASSOCIATED PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09735">
        <property type="entry name" value="Nckap1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="1" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="5" id="KW-0966">Cell projection</keyword>
    <keyword evidence="4" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="3" id="KW-1133">Transmembrane helix</keyword>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="519"/>
            <end position="544"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="530"/>
            <end position="544"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00037839"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00037947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00039689"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029950.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1007" mass="114836" checksum="36E24F1AFA94BBC8" modified="2019-05-08" version="1">QSTVNFDLTKNYLDLIITYTTLMILLSRIEERKAIIGLYNYAHEMTHGASDREYPRLGQMIVDYENPLKKMMEEFVPHSKSLSDALISLQMVYPRRNLSADQWRNAQLLSLISAPSTMLNPAQSDTMPCEYLSLDAMEKWIIFGFILCHGILNSDATALNLWKLALQSSSCLSLFRDEVFHIHKAAEDLFVNIRGYNKRINDIRECKEAAVSHAGSMHRERRKFLRSALKELATVLSDQPGLLGPKALFVFMALSFARDEIIWLLRHADNMPKKSADDFIDKHIAELIFYMEELRAHVRKYGPVMQRYYVQYLSGFDAVVLNELVQNLSVCPEDESIIMSSFVNTMTSLSVKQVEDGEVFDFRGMRLDWFRLQAYTSVSKASLSLADHRELGKMMNTIIFHTKMVDSLVEMLVETSDLSIFCFYSRAFEKMFQQCLELPSQSRYSIAFPLLCTHFMSCTHELCPEERHHIGDRSLSLCNMFLDEMAKQARNLITDICTEQCTLSDQLLPKHCAKTISQAVNKKSKKQTGKKGEPEREKPGVESMRKNRLVVTNLDKLHTALSELCFSINYVPNMVVWEHTFTPREYLTSHLEIRFTKSIVGMTMYNQATQEIAKPSELLTSVRAYMTVLQSIENYVQIDITRVFNNVLLQQTQHLDSHGEPTITSLYTNWYLETLLRQVSNGHIAYFPAMKAFVNLPTENELTFNAEEYSDISEMRALSELLGPYGMKFLSESLMWHISSQVAELKKLVVENVEVLTQMRTSFDKPDQMAALFKRLSSVDSVLKRMTIIGVILSFRSLAQEALRDVLSYHIPFLVSSIEDFKDHIPRETDMKVAMNVYELSSAAGLPCEIDPALVVALSSQKSENISPEEEYKIACLLMVFVAVSLPTLASNVMSQYSPAIEGHCNNIHCLAKAINQIAAALFTIHKGSIEDRLKEFLALASSSLLKIGQETDKTTTRNRESVYLLLDMIVQESPFLTMDLLESCFPYVLLRNAYHAVYKQSVTSSA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPS8</accession>
    <name>A0A452IPS8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Rap guanine nucleotide exchange factor 6</fullName>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the RAPGEF2 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000034060.1">
        <property type="protein sequence ID" value="ENSGAGP00000029991.1"/>
        <property type="gene ID" value="ENSGAGG00000021645.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007264">
        <property type="term" value="P:small GTPase-mediated signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00038">
        <property type="entry name" value="CAP_ED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd06755">
        <property type="entry name" value="PDZ_RapGEF2_RapGEF6-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd01785">
        <property type="entry name" value="RA_PDZ-GEF1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00155">
        <property type="entry name" value="RasGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd06224">
        <property type="entry name" value="REM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.840.10:FF:000001">
        <property type="entry name" value="Rap guanine nucleotide exchange factor (GEF) 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000024">
        <property type="entry name" value="rap guanine nucleotide exchange factor 2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.870.10:FF:000001">
        <property type="entry name" value="rap guanine nucleotide exchange factor 2 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.10:FF:000019">
        <property type="entry name" value="rap guanine nucleotide exchange factor 6 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.42.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.10">
        <property type="entry name" value="Jelly Rolls"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.840.10">
        <property type="entry name" value="Ras guanine-nucleotide exchange factors catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.870.10">
        <property type="entry name" value="Son of sevenless (SoS) protein Chain: S domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000595">
        <property type="entry name" value="cNMP-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018490">
        <property type="entry name" value="cNMP-bd_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001478">
        <property type="entry name" value="PDZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036034">
        <property type="entry name" value="PDZ_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000159">
        <property type="entry name" value="RA_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000651">
        <property type="entry name" value="Ras-like_Gua-exchang_fac_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023578">
        <property type="entry name" value="Ras_GEF_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001895">
        <property type="entry name" value="RASGEF_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036964">
        <property type="entry name" value="RASGEF_cat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014710">
        <property type="entry name" value="RmlC-like_jellyroll"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029071">
        <property type="entry name" value="Ubiquitin-like_domsf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45161">
        <property type="entry name" value="CYTOSKELETON-ASSOCIATED PROTEIN 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45161:SF4">
        <property type="entry name" value="RAP GUANINE NUCLEOTIDE EXCHANGE FACTOR 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00027">
        <property type="entry name" value="cNMP_binding"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00595">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00788">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00617">
        <property type="entry name" value="RasGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00618">
        <property type="entry name" value="RasGEF_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00100">
        <property type="entry name" value="cNMP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00228">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00314">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00147">
        <property type="entry name" value="RasGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00229">
        <property type="entry name" value="RasGEFN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51206">
        <property type="entry name" value="cAMP-binding domain-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50156">
        <property type="entry name" value="PDZ domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48366">
        <property type="entry name" value="Ras GEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54236">
        <property type="entry name" value="Ubiquitin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50042">
        <property type="entry name" value="CNMP_BINDING_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50106">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50200">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50009">
        <property type="entry name" value="RASGEF_CAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50212">
        <property type="entry name" value="RASGEF_NTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7 9" id="KW-0344">Guanine-nucleotide releasing factor</keyword>
    <keyword evidence="8" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Cyclic nucleotide-binding" evidence="12">
        <location>
            <begin position="259"/>
            <end position="359"/>
        </location>
    </feature>
    <feature type="domain" description="N-terminal Ras-GEF" evidence="15">
        <location>
            <begin position="391"/>
            <end position="504"/>
        </location>
    </feature>
    <feature type="domain" description="PDZ" evidence="13">
        <location>
            <begin position="509"/>
            <end position="579"/>
        </location>
    </feature>
    <feature type="domain" description="Ras-associating" evidence="14">
        <location>
            <begin position="729"/>
            <end position="815"/>
        </location>
    </feature>
    <feature type="domain" description="Ras-GEF" evidence="11">
        <location>
            <begin position="840"/>
            <end position="1068"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="151"/>
            <end position="223"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1177"/>
            <end position="1260"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1443"/>
            <end position="1464"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1477"/>
            <end position="1520"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="166"/>
            <end position="183"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="10">
        <location>
            <begin position="206"/>
            <end position="218"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="1206"/>
            <end position="1217"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="1231"/>
            <end position="1240"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="1241"/>
            <end position="1258"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="1454"/>
            <end position="1464"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="1499"/>
            <end position="1508"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00010829"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022658"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00168"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50009"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50106"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50200"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029991.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="18">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1520" mass="170708" checksum="C0485745A226F806" modified="2019-05-08" version="1">MDLNTIYSYLHGMDILSHLREHQLRIMSSGARYERYKGNQVLFCSDTIARCWYILLSGSVLMKDSMFLPPCSFGKQSGGKRGCECIILEPSEMIVVENSKDNEENILQREVPLRQSRRRFRKINQRGERQTITDNVDASSYLSLPADLTKMHLTDNPHPQVTHVPSSQSGCSIASDSGSSSLSDIYQATESDMGDVDLTGLPEAPVDSEDEEEEDEDIDRTSDPLLGRDVVRECLEKEPADKTDDDIEQLLEFMHQLPAFANMTMSVRRELCSVMIFEVVEQAGAIILEDGQELDSWYVILNGTVEISHPDGKTESLCMGNSFGINPSLDKQYMNGVVRTKVDDCQFVCIAQQDYWRILNHVEKNTHKVEEEGEIVMVKEHRELDRSGTRKGHIVIKATPERLIMHLIEEHSIVDPTYIEDFLLTYRTFLSSPLEVGDKLLEWFKVDSLRDKVTRVVLLWVNNHFNDFEGDPAMTRFLEEFEKNLEDTKMKGHLRLLNIACAAKAKWRQITLQKASRDSPLFFCLVGGSDKGFGIFVETVETGSKAAEAGLKRGDQIMEVNGQNFENITFVKAVEILRNNTHLSITVKTNIFVFKELLSRIGQEKNGVPHIPKIAEKKSNRYSIPDMPGDMEQMFPCEKGNKKIKANTVSGGRNRIRKILDKTRFSILPPKLFSDGSVSQSQDDSIVGTRQCRHSLAIMPIPGSLSSSSPDLLQPTTSMLDFTNPSDIPDQVIRVFKADQQSCYIIISKDTTAKEVVSHAVHEFNLIGAPETYSLCEVSVSPEGVIKQRRLPDQFSKLADRIQLNGRYYLKNNMETETLCSDEDAQELLRESQISLLQLSTIEVATQLSMRDFELFRNIEPTEYIDDLYKMDSKTGNAHLKQFEDVINQETFWVATEILTEPNQLKRMKIVKHFIKIALHCRECKNFNSMFAVISGLNLAPVARLRGTWEKLPSKYEKHLRDLQDLFDPSRNMAKYRNILSSQSMQPPIIPLFPVVKKDITFLHEGNDSKVDGLVNFEKLRMIAKEIRQVVRMTSANMDPAVMFRQRSLSQGSTNSNMLDVQGGAHKKRARRSSLLNAKKLYEDAQMARKVKQYLSNLNVETDEEKFQIMSLQCEPAYSTLTKNLNERKGAKSSEMSPIPMRSIGQTTKAHLHQQNRMSQVLQVPAVNLYPIRKKGPAKDHVTFSSPQKSLSLSEEVSSKKQAEDSISMASSLHSSPPASPQGSPHKVGNIQRSDNCGQMNLSGSSSSLASETSNKNSGQRSYGIGYALIPSTKSDNFSDSSHSEISSRSSIVSNCSADSMSAALQDERCLPQSLIVMDSAGGVERKDVPQPSAEYSQPTTGWSLTRPAIIRGLAVTSSMSNEEISHEHITVEAADSGRGSWTSCSSSSHDNFQNIQNQKSWDLLSAYRHTHLDGPIAEVEPTNCCSEETYLYSEAFSKNNRQSKASTEHDQSRQSWASSSSLSDTYETNYGTIKRRGLEHSTAEQTEVLDSKPATDANYKTVTSSTEKGLIEDEQVSAV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPS9</accession>
    <name>A0A452IPS9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">UDP-glucose glycoprotein glucosyltransferase 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Recognizes glycoproteins with minor folding defects. Reglucosylates single N-glycans near the misfolded part of the protein, thus providing quality control for protein folding in the endoplasmic reticulum. Reglucosylated proteins are recognized by calreticulin for recycling to the endoplasmic reticulum and refolding or degradation.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="10">
            <text>N(4)-(alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;6)]-alpha-D-Man-(1-&gt;6)]-beta-D-Man-(1-&gt;4)-beta-D-GlcNAc-(1-&gt;4)-beta-D-GlcNAc)-L-asparaginyl-[protein] (N-glucan mannose isomer 9A1,2,3B1,2,3) + UDP-alpha-D-glucose = N(4)-(alpha-D-Glc-(1-&gt;3)-alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;6)]-alpha-D-Man-(1-&gt;6)]-beta-D-Man-(1-&gt;4)-beta-D-GlcNAc-(1-&gt;4)-beta-D-GlcNAc)-L-asparaginyl-[protein] + UDP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:61304"/>
            <dbReference type="Rhea" id="RHEA-COMP:14356"/>
            <dbReference type="Rhea" id="RHEA-COMP:14357"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:58223"/>
            <dbReference type="ChEBI" id="CHEBI:58885"/>
            <dbReference type="ChEBI" id="CHEBI:59080"/>
            <dbReference type="ChEBI" id="CHEBI:139493"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </cofactor>
    </comment>
    <comment type="pathway">
        <text evidence="3">Protein modification; protein glycosylation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Endoplasmic reticulum lumen</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the glycosyltransferase 8 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000033895.1">
        <property type="protein sequence ID" value="ENSGAGP00000029852.1"/>
        <property type="gene ID" value="ENSGAGG00000021377.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00378"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005788">
        <property type="term" value="C:endoplasmic reticulum lumen"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003980">
        <property type="term" value="F:UDP-glucose:glycoprotein glucosyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051082">
        <property type="term" value="F:unfolded protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036503">
        <property type="term" value="P:ERAD pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0018279">
        <property type="term" value="P:protein N-linked glycosylation via asparagine"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.550.10">
        <property type="entry name" value="Spore Coat Polysaccharide Biosynthesis Protein SpsA, Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040497">
        <property type="entry name" value="Glyco_transf_24"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029044">
        <property type="entry name" value="Nucleotide-diphossugar_trans"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009448">
        <property type="entry name" value="UDP-g_GGtrans"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040693">
        <property type="entry name" value="UGGT_TRXL_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040694">
        <property type="entry name" value="UGGT_TRXL_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040692">
        <property type="entry name" value="UGGT_TRXL_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040525">
        <property type="entry name" value="UGGT_TRXL_4"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11226">
        <property type="entry name" value="UDP-GLUCOSE GLYCOPROTEIN:GLUCOSYLTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11226:SF3">
        <property type="entry name" value="UDP-GLUCOSE:GLYCOPROTEIN GLUCOSYLTRANSFERASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18404">
        <property type="entry name" value="Glyco_transf_24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18400">
        <property type="entry name" value="Thioredoxin_12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18401">
        <property type="entry name" value="Thioredoxin_13"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18402">
        <property type="entry name" value="Thioredoxin_14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18403">
        <property type="entry name" value="Thioredoxin_15"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06427">
        <property type="entry name" value="UDP-g_GGTase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53448">
        <property type="entry name" value="Nucleotide-diphospho-sugar transferases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="8" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="12" id="KW-0472">Membrane</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="5" id="KW-0808">Transferase</keyword>
    <keyword evidence="12" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="12" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="12"/>
            <end position="32"/>
        </location>
    </feature>
    <feature type="domain" description="UGGT thioredoxin-like" evidence="13">
        <location>
            <begin position="49"/>
            <end position="230"/>
        </location>
    </feature>
    <feature type="domain" description="UGGT thioredoxin-like" evidence="14">
        <location>
            <begin position="304"/>
            <end position="435"/>
        </location>
    </feature>
    <feature type="domain" description="UGGT thioredoxin-like" evidence="15">
        <location>
            <begin position="443"/>
            <end position="691"/>
        </location>
    </feature>
    <feature type="domain" description="UDP-glucose:glycoprotein glucosyltransferase thioredoxin-like" evidence="16">
        <location>
            <begin position="721"/>
            <end position="947"/>
        </location>
    </feature>
    <feature type="domain" description="Glucosyltransferase 24 catalytic" evidence="17">
        <location>
            <begin position="1246"/>
            <end position="1353"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1381"/>
            <end position="1408"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001913"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004922"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00006351"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00045874"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00048456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="Pfam" id="PF18400"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="Pfam" id="PF18401"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="Pfam" id="PF18402"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF18403"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF18404"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029852.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="20">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1408" mass="159768" checksum="78734CE6E7761266" modified="2019-05-08" version="1">MQVVRLGDPSGIFYKMGIFIALLGIWFSSSLAKADSKAVTTSLTTKWLSTPLLLEASEFLAEEGQEKFWTFVEASQDIRSSDHGGTNYSSYHAMLKAACQNLSPLQQNLLKFSLSLRSYSATIQAFQQIAADEPPPVGCNSFFAVHGKKTCEFDTLEILLQTASERLKPFLFKGDHRFPLSNPESPVVILYAEIGTEEFSKFHNLLVSKASAGDITYILRHYVANPSKEKVYLSGYGVELAIKSTEYKAKDDTQVKGMEVNATMIGENDPIDEVQGFLFGKLRQLHPDLKEELKELRKHLIESTNEMAPLKVWQLQDLSFQTAARILAAPTVDALMVMKDLSQNFPTKARAITKTVVSSELRTEIEENQKYFKGTLGLQSGDSAFFINGLHIDLDTQDIFSLFDVLRNEARVMEGLHSLGIEGLSLHNVLKLNIQPSDSDYAVDIRSPAISWINNLEVDSRYNSWPSSVQELLRPTFPGVIRQIRKNFHNFVLIVDPSHESTAELLNVAEMFFSNHIPLRIGLVFVVNDSEDVDGLQDAGVALLRAFNYVAQEVDNNYAFQTLISIYNKVKTGDQLQVEHVVSVLEKQYPYVEMNSILGIDSVYDQNRKEGRGYYEQTGVGPLPIVLFNGMPFQKEQLDPDDLETITMQKILETTSLFQRAVYLGELSSDQDVVEYIMNQPNVVPRINSRILMSDREYLDLTATNNFFVDDYARFTFLNSKDKTAAVANSMTYLTKKGMSSKEIYDDSFVRPVTFWIVGDFDKPSGRQLLYDAIKHQKSSNHVRISMINNPSEDPTSKNTVITRAIWAALQTQTSNNAKNFITKMAKEETAKALEAGADITEFAVGGMDVNTFKEAFESPKVDFILSHAMYCRDVLKLQKGQRAVISNGRIIGPLEDNELFNQDDFHLLENIILKTSGQKIKSHIQQLGVEEDLASDLVMKVDALLSAQPKGEARIEYQFFEDRYSAVKLRPKEGETYFDVVAIVDPATRDAQRLAPLLMVLNQLINMNLRVFMNCQSKLSDMPLKSFYRYVLEPEISFTADDNFAPGPIAKFLDMPQTPLFTLNLNTPESWMVESVRTPYDLDNIYLEEVDNRVAAEYELEYLLLEGHCYDITTGQPPRGLQFTLGTSANPVIVDTIVMANLGYFQLKANPGAWVLRLRKGRSEDIYRIYSHDGTDSPPDANEVTVVLNNFKSKIIKVKVSLKTVLQELLWKYKYSNEDYIFPDFRGFTGGQKNDEMKQDKDDVINIFSVASGHLYERFLRIMMLSVLKHTKTPVKFWFLKNYLSPMFKEFIPYMAEKYNFQYELVQYKWPRWLHQQTEKQRIIWGYKILFLDVLFPLVVDKFLFVDADQVSYLMALGSLAGITRSAYIAEEGSALQQMWQGSPCPSLPTDWRGTEPSRQGASGRAR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPV1</accession>
    <name>A0A452IPV1_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="5">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the FAM76 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPV1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033923.1">
        <property type="protein sequence ID" value="ENSGAGP00000029873.1"/>
        <property type="gene ID" value="ENSGAGG00000021565.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016607">
        <property type="term" value="C:nuclear speck"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032017">
        <property type="entry name" value="FAM76"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46176">
        <property type="entry name" value="LD21662P"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46176:SF2">
        <property type="entry name" value="PROTEIN FAM76A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16046">
        <property type="entry name" value="FAM76"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 3" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="120"/>
            <end position="161"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="3">
        <location>
            <begin position="197"/>
            <end position="274"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="140"/>
            <end position="161"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00009097"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029873.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="286" mass="32652" checksum="5BFFA7C2881257A5" modified="2019-05-08" version="1">LCFLLQECRIAHPIVKCTYCRTEFQQESKTNTICKKCAQNVKLYGTPKPCQYCNVIAAFIGNKCQRCTNSEKKYGPPLSCEQCKQQCAFDRKDDRKKVDGKLLCWLCTLSYKRVLQKTKEQRKHLSSSSRTSLQEKEQFSRLSSGSHYNSQKTLSTSSIQNEIPKKKPKFEAISANGDSFSPDLALDSPGTDHFVIIAQLKEEVATLKKMLHQKDQMILEKEKKITELKADLQYQESQMRAKMNQMEKTHKEVMEQLQAKNRELLKQAAALSKGKKPEKSGAITSP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPV2</accession>
    <name>A0A452IPV2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">G-protein coupled receptors family 1 profile domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPV2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034212.1">
        <property type="protein sequence ID" value="ENSGAGP00000030134.1"/>
        <property type="gene ID" value="ENSGAGG00000021739.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004930">
        <property type="term" value="F:G protein-coupled receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00637">
        <property type="entry name" value="7tm_classA_rhodopsin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1070.10">
        <property type="entry name" value="Rhodopsin 7-helix transmembrane proteins"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000276">
        <property type="entry name" value="GPCR_Rhodpsn"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017452">
        <property type="entry name" value="GPCR_Rhodpsn_7TM"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45695:SF37">
        <property type="entry name" value="FREE FATTY ACID RECEPTOR 4-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45695">
        <property type="entry name" value="LEUCOKININ RECEPTOR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00001">
        <property type="entry name" value="7tm_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00237">
        <property type="entry name" value="GPCRRHODOPSN"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81321">
        <property type="entry name" value="Family A G protein-coupled receptor-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50262">
        <property type="entry name" value="G_PROTEIN_RECEP_F1_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0297">G-protein coupled receptor</keyword>
    <keyword evidence="5 9" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-0675">Receptor</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0807">Transducer</keyword>
    <keyword evidence="2 9" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="3 9" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="96"/>
            <end position="117"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="129"/>
            <end position="153"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="165"/>
            <end position="184"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="205"/>
            <end position="230"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="272"/>
            <end position="295"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="302"/>
            <end position="322"/>
        </location>
    </feature>
    <feature type="domain" description="G-protein coupled receptors family 1 profile" evidence="10">
        <location>
            <begin position="108"/>
            <end position="327"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="353"/>
            <end position="373"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="357"/>
            <end position="366"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023224"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50262"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030134.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="373" mass="41690" checksum="9ED28CB0A76D3A33" modified="2019-05-08" version="1">MSEEEEEMRLLSARSPCKHPPLAPATQSCRPVPPLGTWVPGLGAAAASLALVACDALGASQMPGACKTPQGNCTRFPFFSDFKGQNRVALSALETAILSCIFLVSLLVNICAISLLARTKKKLRTANYLVLNLFCADLLFISAIPVILVVRWTESWVLGDVVCHMLFYVISLSGSVTILSLAAVSLERVVCIVRLRHTARFSCKLLAGTLLLIWGVSALATLPLCLFFSVEPLLRHGEITKASRRRLNVSLAYSEIHQIRVSQQDYKLFRTLFVLMISFFIMWSPIVITILLILVQNFKHDLNILPSFFFWIMAFTFANSAVNPVLYNVAHFRHEWRQFLLCCAALPGRRMTNTETTGRRNDHRESNLSVIYK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPX8</accession>
    <name>A0A452IPX8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Cohesin subunit SA</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="2">SCC3 homolog</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="2">Stromal antigen</fullName>
        </alternativeName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="2">Component of cohesin complex, a complex required for the cohesion of sister chromatids after DNA replication. The cohesin complex apparently forms a large proteinaceous ring within which sister chromatids can be trapped. At anaphase, the complex is cleaved and dissociates from chromatin, allowing sister chromatids to segregate.</text>
    </comment>
    <comment type="subunit">
        <text evidence="2">Part of the cohesin complex which is composed of a heterodimer between a SMC1 protein (SMC1A or SMC1B) and SMC3, which are attached via their hinge domain, and RAD21 which link them at their heads, and one STAG protein.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Chromosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Chromosome</location>
            <location evidence="2">Centromere</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="1 2">Belongs to the SCC3 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPX8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034013.1">
        <property type="protein sequence ID" value="ENSGAGP00000029954.1"/>
        <property type="gene ID" value="ENSGAGG00000021612.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000034025.1">
        <property type="protein sequence ID" value="ENSGAGP00000029964.1"/>
        <property type="gene ID" value="ENSGAGG00000021612.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000785">
        <property type="term" value="C:chromatin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000775">
        <property type="term" value="C:chromosome, centromeric region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008278">
        <property type="term" value="C:cohesin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003682">
        <property type="term" value="F:chromatin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007059">
        <property type="term" value="P:chromosome segregation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007062">
        <property type="term" value="P:sister chromatid cohesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039662">
        <property type="entry name" value="Cohesin_Scc3/SA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056396">
        <property type="entry name" value="HEAT_SCC3-SA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020839">
        <property type="entry name" value="SCD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013721">
        <property type="entry name" value="STAG"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11199:SF2">
        <property type="entry name" value="COHESIN SUBUNIT SA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11199">
        <property type="entry name" value="STROMAL ANTIGEN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24571">
        <property type="entry name" value="HEAT_SCC3-SA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21581">
        <property type="entry name" value="SCD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08514">
        <property type="entry name" value="STAG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51425">
        <property type="entry name" value="SCD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="2" id="KW-0132">Cell division</keyword>
    <keyword evidence="2" id="KW-0158">Chromosome</keyword>
    <keyword evidence="2" id="KW-0159">Chromosome partition</keyword>
    <keyword evidence="3" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="2" id="KW-0539">Nucleus</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="SCD" evidence="5">
        <location>
            <begin position="316"/>
            <end position="401"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="1"/>
            <end position="74"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="1079"/>
            <end position="1119"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="3">
        <location>
            <begin position="273"/>
            <end position="307"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="1"/>
            <end position="25"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="27"/>
            <end position="44"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="4">
        <location>
            <begin position="57"/>
            <end position="69"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="1101"/>
            <end position="1116"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00005486"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="RuleBase" id="RU369063"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS51425"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029964.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1193" mass="136762" checksum="D799206A0080C1ED" modified="2019-05-08" version="1">MLRLEGEKNRRLGHLGRENAIHDETVPESTVQSGSCLRNGNLQTPRGKRKVQEQCARVRRNSSRSRRRPSQPQNGEVVEAVTLFEVVSMGKRAMQSVVDDWVEAYIQDRDVALLDLINFYIQCSGCQGMVTAEMFQSLCNSDIMQKMTETFDEETGLQYKKFMAYPWILTVTWPVDMDNGDYPLIKPGPYWKKFKANFCEFTAVLIQQCQYSILYDSYLMDTVISLLTGLADSRVRAFRHTSTLAAVKLLTALVRVNLNLDISKCNAQRLYQVEKNRISVKRTNQRLDQLERKRKEYEQKPLEIENMMNAIFKGTFLQRYRDVIPEIRAICIEEIGSWMKMYPDAFLNDSYLKYIGWMLYDKQPEVRLKCLLGLQGIYDRKELVSTMDLFTSRFKDRIVSMTLDKDHEVAVQAMKLLMLMSQNCEDALSSEDCETLFQFVYTTYRPLAVAAGEFLYKRLLSYQGTEEEVPPKRRGKFGANTCYIRKLITFFLESELHKHVTYLVDSLWDWAGSLLKDWECMTTLLLETAEEEALSDAQESALIEIILVTIREAAEGHPPVGRGAAKKMLSAKEKKIQLEDCTKITEHFIVVLPQLLAKYSADAEKVANLLQIPQYYDLDVYSTGRLEKHVDALLSVVKGIVAKHSDMDVLEACSRMYYVLCNEELAIHRTVDLARTQLIDELVDKLNELLDDSWHEGEGLCTDEEGVCRIYSTLRRIAAFHNAHNLTKWNLYGKTSKLLVFEMEYGSLPVQMILPALQCTYFALLWQLASVVERSSSKETLLALNRELRFFCQICACCLSHKDKDLSEKAFMILCDLLMIVSHQSSRDDEAIGVLPSASLQSKMLLFIQDHVFTEEKEEIKDLTEEEEREKESYKLNDLHRRRSLLAVYCKLIVFTVVEMSAAAEIYKHYMKTYNDFGDIIKETLSRTRHNNKIQSARTLILCLQQLFQKHTETHGSTNCMDSSFTNIKELARRFSLTFGWDQVKSRESVAMIHKEGIEFAFQGAAGVEGKCLPPNLNFLLIISEFSNKLLKPDKRLVYSYLQRYITEPASCRGDEWHPLVWYRNSLLANEEEENSVISSLREWSPTSRSKTSSFKRKLSNGSLPETSHTETTNKTPDFLCTPELTAAAGKSRKKLKSQDTTLQECLPVFCPGGLSRRHSEDVIKAEDFSADGGAEDVGEEVDVVGADQVMVI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPX9</accession>
    <name>A0A452IPX9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">APC membrane recruitment protein 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Peripheral membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the Amer family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPX9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034244.1">
        <property type="protein sequence ID" value="ENSGAGP00000030164.1"/>
        <property type="gene ID" value="ENSGAGG00000021765.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008013">
        <property type="term" value="F:beta-catenin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005546">
        <property type="term" value="F:phosphatidylinositol-4,5-bisphosphate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060828">
        <property type="term" value="P:regulation of canonical Wnt signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016055">
        <property type="term" value="P:Wnt signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019003">
        <property type="entry name" value="AMER"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22237:SF0">
        <property type="entry name" value="APC MEMBRANE RECRUITMENT PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22237">
        <property type="entry name" value="APC MEMBRANE RECRUITMENT PROTEIN 2-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09422">
        <property type="entry name" value="AMER"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="5" id="KW-0446">Lipid-binding</keyword>
    <keyword evidence="6" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0879">Wnt signaling pathway</keyword>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="37"/>
            <end position="219"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="332"/>
            <end position="365"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="41"/>
            <end position="50"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="159"/>
            <end position="168"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="337"/>
            <end position="346"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004202"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007750"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022687"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023121"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030164.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="10">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="446" mass="48537" checksum="24252F7BD5656F61" modified="2025-10-08" version="2">MSSHSKSTAELEIAPPGKLKKTSFKLFGGKRSICTLPSFFGGKNKSQGKGASKKGLSKSKTHDGISDVTYEDGTGRLRLESPLNGGMDSHSRPLPSSQSAHLATDASVKFEFGRQESSHLGSAEGFEKKPNGDKSLSFPRPKKGLKGLFNSIRRHRKNKTAESEKAEWAADCGDGEQAKKTRGAGAETQRASEEESPRGTPLPGPYNDPPSVYSGDHFDSLTGCGDIIAEPEIDSIAESTISVERSRDAAKRSSCLVTYQGGGEEMATPEEIEEYLQQLWEGTTEVDSRYETKLPQPINSPKLQGVNSKLETCGLHEEAAHLYAGGVMDDVELLTPPSDQQESAPNSDEGYYDSTTPGPEDELGEIKKDRLPRDSYSGDALYEFYEPDDTLMSPSLGDEPLFESKASHPEIFSLSFPDSGINPYLARAQVPLTSMGHKYMFKYCPK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPY1</accession>
    <name>A0A452IPY1_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="20">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Cell cortex</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the FNBP1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPY1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034113.1">
        <property type="protein sequence ID" value="ENSGAGP00000030042.1"/>
        <property type="gene ID" value="ENSGAGG00000021677.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005938">
        <property type="term" value="C:cell cortex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008289">
        <property type="term" value="F:lipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006897">
        <property type="term" value="P:endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd07676">
        <property type="entry name" value="F-BAR_FBP17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd11629">
        <property type="entry name" value="HR1_FBP17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12071">
        <property type="entry name" value="SH3_FBP17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1270.60:FF:000002">
        <property type="entry name" value="Formin-binding protein 1-like isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.40:FF:000017">
        <property type="entry name" value="Formin-binding protein 1-like isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.470">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.60">
        <property type="entry name" value="Arfaptin homology (AH) domain/BAR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027267">
        <property type="entry name" value="AH/BAR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031160">
        <property type="entry name" value="F_BAR_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001060">
        <property type="entry name" value="FCH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037449">
        <property type="entry name" value="FNBP1_F-BAR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035492">
        <property type="entry name" value="FNBP1_SH3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011072">
        <property type="entry name" value="HR1_rho-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057870">
        <property type="entry name" value="HR1_TOCA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15735">
        <property type="entry name" value="FCH AND DOUBLE SH3 DOMAINS PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15735:SF13">
        <property type="entry name" value="FORMIN-BINDING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00611">
        <property type="entry name" value="FCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25610">
        <property type="entry name" value="HR1_TOCA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00018">
        <property type="entry name" value="SH3_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00055">
        <property type="entry name" value="FCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00326">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103657">
        <property type="entry name" value="BAR/IMD domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51741">
        <property type="entry name" value="F_BAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51860">
        <property type="entry name" value="REM_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="9 14 15" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="12" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="8" id="KW-0254">Endocytosis</keyword>
    <keyword evidence="10" id="KW-0446">Lipid-binding</keyword>
    <keyword evidence="11" id="KW-0472">Membrane</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 13" id="KW-0728">SH3 domain</keyword>
    <feature type="domain" description="F-BAR" evidence="18">
        <location>
            <begin position="1"/>
            <end position="264"/>
        </location>
    </feature>
    <feature type="domain" description="REM-1" evidence="19">
        <location>
            <begin position="432"/>
            <end position="509"/>
        </location>
    </feature>
    <feature type="domain" description="SH3" evidence="17">
        <location>
            <begin position="578"/>
            <end position="639"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="293"/>
            <end position="320"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="333"/>
            <end position="366"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="517"/>
            <end position="561"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="15">
        <location>
            <begin position="439"/>
            <end position="499"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="16">
        <location>
            <begin position="338"/>
            <end position="347"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="523"/>
            <end position="540"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004544"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00009426"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022583"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023121"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01077"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS51741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS51860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030042.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="649" mass="75097" checksum="6E2ACA903789C7D2" modified="2019-05-08" version="1">MSWGTELWDQFDNLEKHTQWGIDVLEKYIKFVKERTEIEISYAKQLRNLSKKYQPKKNSKEEEEYKYTSCKAFLATLNEMNDYAGQHEVISENMTSQITAELARFVQELKQERKSHFHDGRKAQQYIETFWKQLEASKRRFERDCKEADRAQQYFEKMDADINVTKADVEKARQQAQIRHQMAEDSKAEYSSILQKFNHDQHEHYHTHIPNIFQKIQEMEERRIVRIGECMKTYAEVDRQVIPIIGKCLDGITKAAESIDQKNDSLLVIEAFKSGFEPPGDIDFEDFTQPMKRTVSESSLSNSRWDGKSEPKPGSKSKGKLWPFIKKNKLMSLLTSPHQPPPPPPASASPSAVPNGPQSPKQQKEPLSHRFNEFMTSKPKIHCFRSLKRGNHSLYFHKELMKRTLGKPTCAVEARDYVLSLKLGGGPEDFSNLPPEQRRKKLQQKVDELNKEIQKEMDQRDALTKMKDVYIKNPQMGDAASVDHKLAELGQNMEKLRLEAQKFEGWLAEVEGRLPVRTEQSRRQSGMYDTQNTSGVNNCAQDRESPDGSYTEEPSQESEMKVPVTEFDDEFDDEEPLPTIGTCKALYTFEGQNEGTISVMEGEMLYVIEEDKGDGWTRIRRNEDEEGYVPTSYVEVYLDKNAKGAMTYI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPY2</accession>
    <name>A0A452IPY2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">N-acetylglucosaminyl-proteoglycan 4-beta-glucuronosyltransferase</fullName>
            <ecNumber evidence="4">2.4.1.225</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="pathway">
        <text evidence="2">Protein modification; protein glycosylation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum membrane</location>
            <topology evidence="1">Single-pass type II membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the glycosyltransferase 47 family.</text>
    </comment>
    <dbReference type="EC" id="2.4.1.225" evidence="4"/>
    <dbReference type="AlphaFoldDB" id="A0A452IPY2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033960.1">
        <property type="protein sequence ID" value="ENSGAGP00000029905.1"/>
        <property type="gene ID" value="ENSGAGG00000021583.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00378"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050508">
        <property type="term" value="F:glucuronosyl-N-acetylglucosaminyl-proteoglycan 4-alpha-N-acetylglucosaminyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050509">
        <property type="term" value="F:N-acetylglucosaminyl-proteoglycan 4-beta-glucuronosyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015012">
        <property type="term" value="P:heparan sulfate proteoglycan biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001503">
        <property type="term" value="P:ossification"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.550.10:FF:000034">
        <property type="entry name" value="Exostosin 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.550.10">
        <property type="entry name" value="Spore Coat Polysaccharide Biosynthesis Protein SpsA, Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004263">
        <property type="entry name" value="Exostosin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040911">
        <property type="entry name" value="Exostosin_GT47"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015338">
        <property type="entry name" value="GT64_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029044">
        <property type="entry name" value="Nucleotide-diphossugar_trans"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48261">
        <property type="entry name" value="ACETYLGLUCOSAMINYLTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48261:SF3">
        <property type="entry name" value="EXOSTOSIN GLYCOSYLTRANSFERASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03016">
        <property type="entry name" value="Exostosin_GT47"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09258">
        <property type="entry name" value="Glyco_transf_64"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53448">
        <property type="entry name" value="Nucleotide-diphospho-sugar transferases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="13" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="9" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="14" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="5" id="KW-0328">Glycosyltransferase</keyword>
    <keyword evidence="15" id="KW-0464">Manganese</keyword>
    <keyword evidence="12" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0735">Signal-anchor</keyword>
    <keyword evidence="6" id="KW-0808">Transferase</keyword>
    <keyword evidence="7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11" id="KW-1133">Transmembrane helix</keyword>
    <feature type="domain" description="Exostosin GT47" evidence="16">
        <location>
            <begin position="102"/>
            <end position="385"/>
        </location>
    </feature>
    <feature type="domain" description="Glycosyl transferase 64" evidence="17">
        <location>
            <begin position="481"/>
            <end position="720"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004648"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004922"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00010271"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00012192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022676"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023211"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF03016"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF09258"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029905.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="737" mass="85522" checksum="BFDFCBEE1AFC82D2" modified="2019-05-08" version="1">MQTRKKYIFITFFASWLLLFFFGGDQIRRLAFLSRKKATVLKNWPRWTDRSLLKSFADPEELQSDGGHHKQSAKARRESRINIYKHSRCRMETCFDFSRCEKHGFKVFVYPMEREDPVSESYLKIITSIEESRYYTSNPEEACLFILNIDTLDRDHLSIQYVHNINDKIQGFPLWNDGRNHLIFNLYSGTWPNYTEDLGFDIGQAILAKASFYVENFRPGFDISIPLFSKEHPQKGGERGWLYQNSVPPKKKYMLAFKGKRYLTGIGSDTRNALHHIHNGKEIISLTTCKHGKDWEKHKDTRCDKDNLDYEKFDYQELLHNSTFCIIPRGRRLGSFRFLEALQAACIPVLLSNGWELPFSEVIDWSKSAIVGDERLLLQIPSTVRCIHSDKILAFQQQTQFLWEAYFSSVEKIVQTTLEIIKDRLFLRESRPKFLWNTLPGGLLALPEYSTHLGDFPFYHLQHGSSPSGKFTAFIRAVSPVVSLSQPILKLIQAVSRSQHCAQILVLWSCEKPPPPRGRWPQTAVPLTVIHGRRKVSDRFFPYSAIGTDAVLSLDEHTSLSTSEVDFAFVVWRSFPDRIVGFPMRSHFWDSSQSQWGYTSKWTNEFSMVLTAAAFYHRYYHSLFTNYLPASLRSLIDQLSNCEDILMNFLVSAVTKLPPIKVTQKKQHQEAAPQQMSRSAAVSSGNLHFSQRQDCLNRFAAWFGYMPLVHSQLRLDPVLFKDQVSVLRKKYRHLEKP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPY4</accession>
    <name>A0A452IPY4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">HECT-type E3 ubiquitin transferase</fullName>
            <ecNumber evidence="2">2.3.2.26</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>S-ubiquitinyl-[E2 ubiquitin-conjugating enzyme]-L-cysteine + [acceptor protein]-L-lysine = [E2 ubiquitin-conjugating enzyme]-L-cysteine + N(6)-ubiquitinyl-[acceptor protein]-L-lysine.</text>
            <dbReference type="EC" id="2.3.2.26"/>
        </reaction>
    </comment>
    <comment type="caution">
        <text evidence="5">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="EC" id="2.3.2.26" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452IPY4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034249.1">
        <property type="protein sequence ID" value="ENSGAGP00000030169.1"/>
        <property type="gene ID" value="ENSGAGG00000021768.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061630">
        <property type="term" value="F:ubiquitin protein ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000209">
        <property type="term" value="P:protein polyubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1750.10:FF:000023">
        <property type="entry name" value="probable E3 ubiquitin-protein ligase HECTD2 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.2160.10:FF:000004">
        <property type="entry name" value="probable E3 ubiquitin-protein ligase HERC4 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.2160.10">
        <property type="entry name" value="Hect, E3 ligase catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1750.10">
        <property type="entry name" value="Hect, E3 ligase catalytic domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044611">
        <property type="entry name" value="E3A/B/C-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000569">
        <property type="entry name" value="HECT_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035983">
        <property type="entry name" value="Hect_E3_ubiquitin_ligase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45700:SF9">
        <property type="entry name" value="HECT-TYPE E3 UBIQUITIN TRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45700">
        <property type="entry name" value="UBIQUITIN-PROTEIN LIGASE E3C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00632">
        <property type="entry name" value="HECT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00119">
        <property type="entry name" value="HECTc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56204">
        <property type="entry name" value="Hect, E3 ligase catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50237">
        <property type="entry name" value="HECT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0808">Transferase</keyword>
    <keyword evidence="4 5" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="HECT" evidence="7">
        <location>
            <begin position="456"/>
            <end position="717"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="29"/>
            <end position="68"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000885"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00012485"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00104"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50237"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030169.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="726" mass="82921" checksum="2E4E81E31EB38C8C" modified="2019-05-08" version="1">MSLQTPQAATAGSDLPAVTEKGSLLSYKMGEDSFVPPSVEEEDGEEEKEKEKLPPILTASNTAGGGLERGTKSQFSTFSSFISTINQKKEAAGSSSSPTQLVIPNIKNVRDLPPICLDVRQKQRISIDTLPPDLKAPVPPEASLPIRTKTMKDFQEDMEKTKLSGDWKAVHDFYLTTFDSFLELNAAFKKDVSLPFSTIEDSGINAKFVNAVYDALLNTPQDIQKSVLKGIINGLLQEWKGPRTKDDLRAYCILLQNPQFSNTSTYVIYAHLLRQIATLAEADHHFLAHWFKKISQRRFKQLVDRLLQFISLRLFPAKPEEFPPMIKCTWWIPSATKVLSLFNAANSLANPAIIPYTDFYNSTLDHIDLMEEYHNWQCYGNSHRFSFCQYPFIISIAAKKFIIQRDSEQQMISMARQSLVDKVSRRQRPDMNMLFLNLKVRRMNLVSDSLDELTRKRADLKKKLKVTFVGEAGLDMGGLTKEWFLLLIRQIFHPDYGMFTYHKDSHCHWFSSYKCDDYSEFRLVGALMGLAVYNSITLDIRFPPCCYKKLLSPPIVPCDQSTPVGISGVSLDDLCHIMPELAHGLSELLSYEDNVEEDFYSTFQVFQEEFGVIKSYNLKPGGDKIPVTAQNRKEYVQLYVDFILNKSIYKQFAAFYYGFHSVCDSYALMLLRPEEVEILVCGSPELHKYIKEVNTREGEELVKLNENVDTRTNDYKLSMNELRVRN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPY6</accession>
    <name>A0A452IPY6_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="20">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Cell cortex</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the FNBP1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPY6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034118.1">
        <property type="protein sequence ID" value="ENSGAGP00000030047.1"/>
        <property type="gene ID" value="ENSGAGG00000021677.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005938">
        <property type="term" value="C:cell cortex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008289">
        <property type="term" value="F:lipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
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        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
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        <property type="match status" value="1"/>
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        <property type="entry name" value="Arfaptin homology (AH) domain/BAR domain"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="SH3 Domains"/>
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        <property type="entry name" value="FNBP1_F-BAR"/>
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        <property type="entry name" value="FNBP1_SH3"/>
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        <property type="entry name" value="HR1_rho-bd"/>
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        <property type="entry name" value="HR1_TOCA"/>
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        <property type="entry name" value="FCH AND DOUBLE SH3 DOMAINS PROTEIN"/>
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        <property type="match status" value="1"/>
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    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="9 14 15" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="12" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="8" id="KW-0254">Endocytosis</keyword>
    <keyword evidence="10" id="KW-0446">Lipid-binding</keyword>
    <keyword evidence="11" id="KW-0472">Membrane</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 13" id="KW-0728">SH3 domain</keyword>
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        <location>
            <begin position="1"/>
            <end position="264"/>
        </location>
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            <begin position="401"/>
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            <begin position="293"/>
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            <begin position="486"/>
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            <begin position="408"/>
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            <begin position="492"/>
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        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
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        <source>
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        <source>
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        <source>
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        <source>
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    </evidence>
    <sequence length="618" mass="71570" checksum="EA532A493F9392CB" modified="2019-05-08" version="1">MSWGTELWDQFDNLEKHTQWGIDVLEKYIKFVKERTEIEISYAKQLRNLSKKYQPKKNSKEEEEYKYTSCKAFLATLNEMNDYAGQHEVISENMTSQITAELARFVQELKQERKSHFHDGRKAQQYIETFWKQLEASKRRFERDCKEADRAQQYFEKMDADINVTKADVEKARQQAQIRHQMAEDSKAEYSSILQKFNHDQHEHYHTHIPNIFQKIQEMEERRIVRIGECMKTYAEVDRQVIPIIGKCLDGITKAAESIDQKNDSLLVIEAFKSGFEPPGDIDFEDFTQPMKRTVSESSLSNSRWDGKSEPKPGSKSKGKLWPFIKKNKPPPPPPASASPSAVPNGPQSPKQQKEPLSHRFNEFMTSKPKIHCFRSLKRGITLNTQVWNLIEGGGPEDFSNLPPEQRRKKLQQKVDELNKEIQKEMDQRDALTKMKDVYIKNPQMGDAASVDHKLAELGQNMEKLRLEAQKFEGWLAEVEGRLPVRTEQSRRQSGMYDTQNTSGVNNCAQDRESPDGSYTEEPSQESEMKVPVTEFDDEFDDEEPLPTIGTCKALYTFEGQNEGTISVMEGEMLYVIEEDKGDGWTRIRRNEDEEGYVPTSYVEVYLDKNAKGAMTYI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPY7</accession>
    <name>A0A452IPY7_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="5">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IPY7"/>
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        <property type="gene ID" value="ENSGAGG00000021582.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000033969.1">
        <property type="protein sequence ID" value="ENSGAGP00000029911.1"/>
        <property type="gene ID" value="ENSGAGG00000021582.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="F:nucleic acid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
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    </dbReference>
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        <property type="entry name" value="R3H_dom"/>
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        <property type="entry name" value="R3H_domain_containing"/>
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        <property type="entry name" value="SUZ"/>
    </dbReference>
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        <property type="entry name" value="CAMP-REGULATED PHOSPHOPROTEIN 21"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15672">
        <property type="entry name" value="CAMP-REGULATED PHOSPHOPROTEIN 21 RELATED R3H DOMAIN CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01424">
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        <property type="match status" value="1"/>
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        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
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        <location>
            <begin position="179"/>
            <end position="242"/>
        </location>
    </feature>
    <feature type="domain" description="SUZ" evidence="4">
        <location>
            <begin position="243"/>
            <end position="315"/>
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    </feature>
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        <location>
            <begin position="1"/>
            <end position="81"/>
        </location>
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            <begin position="110"/>
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            <begin position="1"/>
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    </feature>
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        <location>
            <begin position="44"/>
            <end position="57"/>
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    </feature>
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        <location>
            <begin position="65"/>
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        <location>
            <begin position="115"/>
            <end position="142"/>
        </location>
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        <location>
            <begin position="354"/>
            <end position="365"/>
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        <location>
            <begin position="366"/>
            <end position="375"/>
        </location>
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        <location>
            <begin position="376"/>
            <end position="388"/>
        </location>
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        <location>
            <begin position="416"/>
            <end position="438"/>
        </location>
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        <location>
            <begin position="439"/>
            <end position="448"/>
        </location>
    </feature>
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        <location>
            <begin position="455"/>
            <end position="469"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="525"/>
            <end position="549"/>
        </location>
    </feature>
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        <location>
            <begin position="550"/>
            <end position="563"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="600"/>
            <end position="616"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="627"/>
            <end position="652"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="2">
        <location>
            <begin position="663"/>
            <end position="678"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS51061"/>
        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029911.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="857" mass="94951" checksum="E7D087FB055BDE38" modified="2019-05-08" version="1">MSEQAKLNQETVEETAKEQESAISDATPENCIINKTESLEAEEKEEKLSDAKTDLQKRGVNQNQQKKRSKSGAKGKLVRSLAVCEESSPRLGAEGLHDNQESIQLQLLSFPSLQEEDKSSKDDSEKEKEKDKNKEKDKNCEKSKIRMLSKDCSQEYTDSTGIDLHEFLVNTLKNNPRDRMILLKMEQEIIDFISDNNNHYKKFPQMSSYQRMLVHRVAAYFGMDHNVDQTGKSVIINKTSNTRIPEQRFSEHLKDEKGEESQKRFILKRDNSSIDKEDNQQNRIHPFRDDRRSKSIEEREEEYQRVRERIFAQDSVCSQENLFVENSRLLEDNSICNDTHKKRQLFRGNRDSTGKASGSRQSSTENEMKWTDHQRPWSSTDSDSSNRNLKPAITKTASFGGITVLTRGDSSSSNRSTGKLSKTGSESSSSAGSSGSLSRTHQPLQSASIAPGVTASPSGNVSYTENGMNGQVAPSNTSYIILPLEAAGIPPGSILLNPHTGQPFVNPDGTPAIYNPPSGQQPIRSQLVGQSQQQPSSQQQQQVQQPQQQMASHLVTQPVQALQPSSQSVQYPAVSYPPQHLLPVSPTQQFPVRDDMTTQFSQMSLSRQSSGDNPESPSGPVYPSPIIQPSGQQTGYVMASPSQQLPPGSFTGSGPPVSQQVLQPPPPPQGYVQQPPPAQMPVYYYPSGQYPTSATQQYRPIASVQYNAQRSQQISQTAQQAGYQPVLTNQQQGFQGLMGMQQPPQSQNLMNNQQGNQVQGLMVQYPAMSSYQVPMTQGSQGLPQQSYQQPIMLPNQSGQGTITATGMPVYCNVIPPTPQNNLRLIAPHCPSNNVPVIAASCRTNCASINNTGWQIKY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPY8</accession>
    <name>A0A452IPY8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Structural maintenance of chromosomes protein 5</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="11">Forms a heterodimer with smc6. Component of the SMC5-SMC6 complex which consists at least of smc5, smc6, nsmce2, nsmce1 and nsmce4a.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Chromosome</location>
            <location evidence="2">Telomere</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the SMC family. SMC5 subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPY8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034041.1">
        <property type="protein sequence ID" value="ENSGAGP00000029974.1"/>
        <property type="gene ID" value="ENSGAGG00000021632.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000781">
        <property type="term" value="C:chromosome, telomeric region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030915">
        <property type="term" value="C:Smc5-Smc6 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003697">
        <property type="term" value="F:single-stranded DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000724">
        <property type="term" value="P:double-strand break repair via homologous recombination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:001301">
        <property type="entry name" value="Structural maintenance of chromosomes 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000793">
        <property type="entry name" value="Structural maintenance of chromosomes protein 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.1490">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038729">
        <property type="entry name" value="Rad50/SbcC_AAA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45916">
        <property type="entry name" value="STRUCTURAL MAINTENANCE OF CHROMOSOMES PROTEIN 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45916:SF1">
        <property type="entry name" value="STRUCTURAL MAINTENANCE OF CHROMOSOMES PROTEIN 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13476">
        <property type="entry name" value="AAA_23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="5" id="KW-0158">Chromosome</keyword>
    <keyword evidence="9 12" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="6" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0779">Telomere</keyword>
    <feature type="domain" description="Rad50/SbcC-type AAA" evidence="14">
        <location>
            <begin position="49"/>
            <end position="316"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="1"/>
            <end position="40"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="12">
        <location>
            <begin position="216"/>
            <end position="359"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="12">
        <location>
            <begin position="387"/>
            <end position="442"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="12">
        <location>
            <begin position="639"/>
            <end position="771"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="12">
        <location>
            <begin position="831"/>
            <end position="868"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="13">
        <location>
            <begin position="26"/>
            <end position="35"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004574"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00010171"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00018687"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022454"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022895"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00063886"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="Pfam" id="PF13476"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029974.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1041" mass="121697" checksum="47BDF317C82676DC" modified="2019-05-08" version="1">MAVSGKRKLGAATSDSGAGWSKRGSGAAGPEGSGRGQPQSRFVEGSIVKVTMENFLTYDNCEVCPGPHLNMIVGANGTGKSSIVCAICLGLAGKPSFIGRADKVGLYVKRGCSKGLVEIELFKTPKNLIITREIHVANNASVWFVNRKPSTLKIVEEQIAALNIQVGNLCQFLPQDKVGEFAKLSKTELLEATEKSVGPPEMYQFHCELKNFRERERELENSLQEKTNFLEKMKQRNERYKQDVERYYERKRHLDLIEMLERKRPWVEYENVRQEHEEVKQNRDHAKEELKKLQQTQSPMTQQIQEVEKQWRNLDIKIKEKTAEIKDTSQRCKQKQDAMEKKDKQIEEIQQALRIKNDEEMDRQKRIHNARKMIEDWQNELNSPGNSENLQPQIDSINSELRQLQEEKANIDGEMSDLRRERENLEREKKRVTDRIVQFNNMMNLKEDKLRGRYRDTHNALLWLRENKDRFKRSVCEPMMLAVRDHQKLKVNAVCAPTVSCAENRPSRSVEELHQYGFFSYLRELFDAPHPVMSYLCSQYRVHDVPVGTEKTRSMIERVIQETKFKQIYTAEERYSVRVSAYTNQTISSNTSLRPAQFLTVTVDADERRQLENQLKDINRLFQLRDSQLHTLYERQKGLERKDNELRQQKKERLERKNKRKQLESKISMKHDSLKQMEQDAINLEEESQQTNARIKEINVQKAKLVTELMQLIKNCIRLNILKVDLVLQSTTVNSKKNRLESDYKAATVQLRALEQQVSDLGGKKRVLLEKCKGLMRKARHACNLSPDQEIPKDFQTAFQDLPSTLDEIDALLNEEKSRASCFTGLNASVVEEYNRRAQEIQQVTEELEEKKNELESYRQNISQVKERWLTPLKQLVEQINEKFSNFFSSMQCAGEVDLHTENEEEYDKYGIRIRVKFRSNTQLHELTPHHQSGGERSVSTMLYLMALQELNRCPFRVVDEINQGMDPVNERRVFEMVVKTACKESTSQYFFITPKLLQNLTYAEKMTVLFVYNGPFMLEPTKWNLKAFRRRRRQVEQMDQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPY9</accession>
    <name>A0A452IPY9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">THO complex subunit 1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="21">Nuclear matrix protein p84</fullName>
        </alternativeName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus matrix</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Nucleus</location>
            <location evidence="3">Nucleoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="19">Belongs to the THOC1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPY9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034084.1">
        <property type="protein sequence ID" value="ENSGAGP00000030015.1"/>
        <property type="gene ID" value="ENSGAGG00000021230.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016363">
        <property type="term" value="C:nuclear matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005654">
        <property type="term" value="C:nucleoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000445">
        <property type="term" value="C:THO complex part of transcription export complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006915">
        <property type="term" value="P:apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006406">
        <property type="term" value="P:mRNA export from nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006397">
        <property type="term" value="P:mRNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008380">
        <property type="term" value="P:RNA splicing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd08318">
        <property type="entry name" value="Death_NMPP84"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.533.10:FF:000025">
        <property type="entry name" value="THO complex subunit 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.533.10">
        <property type="entry name" value="Death Domain, Fas"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011029">
        <property type="entry name" value="DEATH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000488">
        <property type="entry name" value="Death_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021861">
        <property type="entry name" value="THO_THOC1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13265">
        <property type="entry name" value="THO COMPLEX SUBUNIT 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13265:SF2">
        <property type="entry name" value="THO COMPLEX SUBUNIT 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00531">
        <property type="entry name" value="Death"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11957">
        <property type="entry name" value="efThoc1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00005">
        <property type="entry name" value="DEATH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47986">
        <property type="entry name" value="DEATH domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50017">
        <property type="entry name" value="DEATH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="13" id="KW-0007">Acetylation</keyword>
    <keyword evidence="9" id="KW-0053">Apoptosis</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="15" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="6" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="8" id="KW-0507">mRNA processing</keyword>
    <keyword evidence="17" id="KW-0508">mRNA splicing</keyword>
    <keyword evidence="10" id="KW-0509">mRNA transport</keyword>
    <keyword evidence="18" id="KW-0539">Nucleus</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="12" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="16" id="KW-0804">Transcription</keyword>
    <keyword evidence="14" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="4" id="KW-0813">Transport</keyword>
    <keyword evidence="11" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="Death" evidence="22">
        <location>
            <begin position="591"/>
            <end position="667"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004109"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004642"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022664"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022703"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022816"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023187"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00061272"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00069888"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00080564"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS50017"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030015.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="671" mass="77242" checksum="106938F141DFB5E4" modified="2019-05-08" version="1">RHKRGPQSAEPTRETMSPPLFSLPEARLRFTNSTRDALTNKTIKPLLNAFNQIPGSENEKKCTLDQSFRVVLEEEIINQASCENVLAIISLAINAVTDGICTASTPFVLLGDVLDCLPLDQCDKIFTFVEKNVTTWKSNTFYSAGKNYLLRMCNDLLRRLSKSQNTVFCGRIQLFLARLFPLSEKSGLNLQSQFNLENVTVFNTNEQESTLGQKHIEERDEGMEVEEGEMGDDEAPTSCSIPIDYNLYRKFWSLQDYFRNPVQCYEKISWKTFLKYSEEVLAVFKSYKLDDTQASRKKLEELKTGGEHVYFAKFLTSEKLMDLQLSDSNFRRHILLQYLILFQYLKGQVKFKSSNYVLTDEQSLWIEDTTKLVYQLLSENPPDGERFSKMVEHILNTEENWNTWKNEGCPSFVKERPSDSKPTRLVRKRPAPEDFLGKGPNKKILMGNEELTRLWNLCPDNMEACKSETREYMPTLEEFFEEAIEQADPENMVENEYKAVNNSNYGWRALRLLARRSPHFFQPTNQQFKSLPEYLENMVIKLAKELPPPSEEIKTGEDEDEEDNDALLKENNESPDVQRDKPVTGGQIESFANKLGEQWKTLAPYLEMKDSDIRQIESDSEDVKMRAKQLLVAWQDQEGIHATPENLITAMNKAGLSDLAESLSNDTENSS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPZ1</accession>
    <name>A0A452IPZ1_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="20">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Cell cortex</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the FNBP1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPZ1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034123.1">
        <property type="protein sequence ID" value="ENSGAGP00000030052.1"/>
        <property type="gene ID" value="ENSGAGG00000021677.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005938">
        <property type="term" value="C:cell cortex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008289">
        <property type="term" value="F:lipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006897">
        <property type="term" value="P:endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd07676">
        <property type="entry name" value="F-BAR_FBP17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd11629">
        <property type="entry name" value="HR1_FBP17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1270.60:FF:000002">
        <property type="entry name" value="Formin-binding protein 1-like isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.40:FF:000017">
        <property type="entry name" value="Formin-binding protein 1-like isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.470">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.60">
        <property type="entry name" value="Arfaptin homology (AH) domain/BAR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027267">
        <property type="entry name" value="AH/BAR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031160">
        <property type="entry name" value="F_BAR_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001060">
        <property type="entry name" value="FCH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037449">
        <property type="entry name" value="FNBP1_F-BAR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011072">
        <property type="entry name" value="HR1_rho-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057870">
        <property type="entry name" value="HR1_TOCA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15735">
        <property type="entry name" value="FCH AND DOUBLE SH3 DOMAINS PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15735:SF13">
        <property type="entry name" value="FORMIN-BINDING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00611">
        <property type="entry name" value="FCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25610">
        <property type="entry name" value="HR1_TOCA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00018">
        <property type="entry name" value="SH3_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00055">
        <property type="entry name" value="FCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00326">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103657">
        <property type="entry name" value="BAR/IMD domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51741">
        <property type="entry name" value="F_BAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51860">
        <property type="entry name" value="REM_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="9 14 15" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="12" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="8" id="KW-0254">Endocytosis</keyword>
    <keyword evidence="10" id="KW-0446">Lipid-binding</keyword>
    <keyword evidence="11" id="KW-0472">Membrane</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 13" id="KW-0728">SH3 domain</keyword>
    <feature type="domain" description="F-BAR" evidence="18">
        <location>
            <begin position="1"/>
            <end position="264"/>
        </location>
    </feature>
    <feature type="domain" description="REM-1" evidence="19">
        <location>
            <begin position="355"/>
            <end position="432"/>
        </location>
    </feature>
    <feature type="domain" description="SH3" evidence="17">
        <location>
            <begin position="501"/>
            <end position="562"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="293"/>
            <end position="319"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="440"/>
            <end position="484"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="15">
        <location>
            <begin position="362"/>
            <end position="422"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="446"/>
            <end position="463"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004544"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00009426"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022583"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023121"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01077"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS51741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS51860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030052.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="572" mass="66553" checksum="C1312536B760B8AB" modified="2019-05-08" version="1">MSWGTELWDQFDNLEKHTQWGIDVLEKYIKFVKERTEIEISYAKQLRNLSKKYQPKKNSKEEEEYKYTSCKAFLATLNEMNDYAGQHEVISENMTSQITAELARFVQELKQERKSHFHDGRKAQQYIETFWKQLEASKRRFERDCKEADRAQQYFEKMDADINVTKADVEKARQQAQIRHQMAEDSKAEYSSILQKFNHDQHEHYHTHIPNIFQKIQEMEERRIVRIGECMKTYAEVDRQVIPIIGKCLDGITKAAESIDQKNDSLLVIEAFKSGFEPPGDIDFEDFTQPMKRTVSESSLSNSRWDGKSEPKPGSKSKGKLWPFIKKNKLSLKLITLNTQVWNLIEGGGPEDFSNLPPEQRRKKLQQKVDELNKEIQKEMDQRDALTKMKDVYIKNPQMGDAASVDHKLAELGQNMEKLRLEAQKFEGWLAEVEGRLPVRTEQSRRQSGMYDTQNTSGVNNCAQDRESPDGSYTEEPSQESEMKVPVTEFDDEFDDEEPLPTIGTCKALYTFEGQNEGTISVMEGEMLYVIEEDKGDGWTRIRRNEDEEGYVPTSYVEVYLDKNAKGAMTYI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ36</accession>
    <name>A0A452IQ36_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Lethal giant larvae homologue 2 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the WD repeat L(2)GL family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQ36"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034306.1">
        <property type="protein sequence ID" value="ENSGAGP00000030225.1"/>
        <property type="gene ID" value="ENSGAGG00000021760.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031201">
        <property type="term" value="C:SNARE complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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    <dbReference type="GO" id="GO:0045159">
        <property type="term" value="F:myosin II binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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    <dbReference type="GO" id="GO:0019905">
        <property type="term" value="F:syntaxin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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    <dbReference type="GO" id="GO:0006887">
        <property type="term" value="P:exocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006893">
        <property type="term" value="P:Golgi to plasma membrane transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:000521">
        <property type="entry name" value="syntaxin-binding protein 5-like isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000664">
        <property type="entry name" value="Lethal2_giant"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013577">
        <property type="entry name" value="LLGL2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036322">
        <property type="entry name" value="WD40_repeat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001680">
        <property type="entry name" value="WD40_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10241">
        <property type="entry name" value="LETHAL 2 GIANT LARVAE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10241:SF22">
        <property type="entry name" value="SYNTAXIN-BINDING PROTEIN 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08366">
        <property type="entry name" value="LLGL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00400">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00962">
        <property type="entry name" value="LETHAL2GIANT"/>
    </dbReference>
    <dbReference type="SMART" id="SM00320">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50978">
        <property type="entry name" value="WD40 repeat-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50082">
        <property type="entry name" value="WD_REPEATS_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50294">
        <property type="entry name" value="WD_REPEATS_REGION"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0268">Exocytosis</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="3 5" id="KW-0853">WD repeat</keyword>
    <feature type="repeat" description="WD" evidence="5">
        <location>
            <begin position="234"/>
            <end position="269"/>
        </location>
    </feature>
    <feature type="domain" description="Lethal giant larvae homologue 2" evidence="7">
        <location>
            <begin position="278"/>
            <end position="385"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="17"/>
            <end position="38"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="556"/>
            <end position="596"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="675"/>
            <end position="694"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="17"/>
            <end position="31"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="570"/>
            <end position="580"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="582"/>
            <end position="591"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00008070"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022574"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF08366"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030225.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="733" mass="81811" checksum="9FAEB9109F60D76A" modified="2019-05-08" version="1">MRKFNIRKVLDGLTAVSSSSSASSSSQQQQSGNRESDVPETLQSEHFQLCKTVRHGFPYQPSALAFDPVQRILAIGTQTGALRLFGRPGVECYCQHDAETAVIQLQFLINEGALVSALADDTLHLWNLRQKRPAIFHSLKFNRERITFCHLPFQSKWLYVGTERGNIHIVNVESFTLSGYVIMWNKAIELSSKSHPGPVVHISDNPMDEGKLLIGYESGIVVLWDLKSKKADYRYTHDEAIHSVAWHHEGKQFICSHSDGTLTIWNIKSPAKPLQTITPHGKQLKDGKKPEPCKPILKVEYKTARAGEPFIILSGGLSYDTVGRRPCLTVMHGKSTAVLEMDYSIVDFLTLCETPYPNDFQEPYAVVVLLEKDLVLIDLAQNGYPIFENPYPLNIHESPVTCCEYFADCPVDLIPALYSVGARQKRQGYSKKEWPISGGNWGLVTQSYPEIIITGHADGSIKFWDASAITLQVLYKLKTAKVFEKSRNKDDKPNTDIVDEDPYAIQIISWCPESRMLCIAGVSAHVIIYRFSKQEVTTEVIPMLEVRLLYEINDVESPDGEQAPPLPTPGTGSNPQSIAPQSHPSTSSSSSDGLRDNVPCLKIKNSPLKQSAGYQIELVIQLVWVSGEPPQQITSLAVNSAYGLVVFGNCNGIAMVDYLQKTVLLNLGTTELYGSNDPYRREPRSPRKSRQPSGGKNCWMLHFNNVTTLNIKTLATQNVYFCMTITCLIFENN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ37</accession>
    <name>A0A452IQ37_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="22">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cell projection</location>
            <location evidence="1">Cilium</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the polycystin family.</text>
    </comment>
    <comment type="caution">
        <text evidence="13">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000034138.1">
        <property type="protein sequence ID" value="ENSGAGP00000030066.1"/>
        <property type="gene ID" value="ENSGAGG00000021465.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005929">
        <property type="term" value="C:cilium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005261">
        <property type="term" value="F:monoatomic cation channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006816">
        <property type="term" value="P:calcium ion transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00037">
        <property type="entry name" value="CLECT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00146">
        <property type="entry name" value="PKD"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="CDD" id="cd01752">
        <property type="entry name" value="PLAT_polycystin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000825">
        <property type="entry name" value="Polycystin 1, transient receptor potential channel interacting"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.60.20:FF:000012">
        <property type="entry name" value="polycystin-1 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.70">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.100.10">
        <property type="entry name" value="Mannose-Binding Protein A, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.60.20">
        <property type="entry name" value="PLAT/LH2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001304">
        <property type="entry name" value="C-type_lectin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016186">
        <property type="entry name" value="C-type_lectin-like/link_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016187">
        <property type="entry name" value="CTDL_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057244">
        <property type="entry name" value="GAIN_B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000203">
        <property type="entry name" value="GPS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000434">
        <property type="entry name" value="PC1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022409">
        <property type="entry name" value="PKD/Chitinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002859">
        <property type="entry name" value="PKD/REJ-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013122">
        <property type="entry name" value="PKD1_2_channel"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000601">
        <property type="entry name" value="PKD_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035986">
        <property type="entry name" value="PKD_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001024">
        <property type="entry name" value="PLAT/LH2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036392">
        <property type="entry name" value="PLAT/LH2_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042060">
        <property type="entry name" value="PLAT_polycystin1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006228">
        <property type="entry name" value="Polycystin_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046791">
        <property type="entry name" value="Polycystin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014010">
        <property type="entry name" value="REJ_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002889">
        <property type="entry name" value="WSC_carb-bd"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00864">
        <property type="entry name" value="PCC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46730">
        <property type="entry name" value="POLYCYSTIN-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46730:SF3">
        <property type="entry name" value="POLYCYSTIN-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00059">
        <property type="entry name" value="Lectin_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00801">
        <property type="entry name" value="PKD"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08016">
        <property type="entry name" value="PKD_channel"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01477">
        <property type="entry name" value="PLAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20519">
        <property type="entry name" value="Polycystin_dom"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02010">
        <property type="entry name" value="REJ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00500">
        <property type="entry name" value="POLYCYSTIN1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00034">
        <property type="entry name" value="CLECT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00303">
        <property type="entry name" value="GPS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00308">
        <property type="entry name" value="LH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00089">
        <property type="entry name" value="PKD"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="SMART" id="SM00321">
        <property type="entry name" value="WSC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56436">
        <property type="entry name" value="C-type lectin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49723">
        <property type="entry name" value="Lipase/lipooxygenase domain (PLAT/LH2 domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49299">
        <property type="entry name" value="PKD domain"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50041">
        <property type="entry name" value="C_TYPE_LECTIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50221">
        <property type="entry name" value="GAIN_B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50093">
        <property type="entry name" value="PKD"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50095">
        <property type="entry name" value="PLAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51111">
        <property type="entry name" value="REJ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51212">
        <property type="entry name" value="WSC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="12" id="KW-0966">Cell projection</keyword>
    <keyword evidence="8" id="KW-0969">Cilium</keyword>
    <keyword evidence="10" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9 15" id="KW-0472">Membrane</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5 15" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 15" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="2944"/>
            <end position="2966"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3152"/>
            <end position="3173"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3193"/>
            <end position="3215"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3415"/>
            <end position="3437"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3443"/>
            <end position="3462"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3530"/>
            <end position="3549"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3759"/>
            <end position="3778"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3790"/>
            <end position="3817"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3837"/>
            <end position="3861"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3887"/>
            <end position="3909"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3946"/>
            <end position="3971"/>
        </location>
    </feature>
    <feature type="domain" description="WSC" evidence="21">
        <location>
            <begin position="60"/>
            <end position="153"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="17">
        <location>
            <begin position="179"/>
            <end position="241"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="16">
        <location>
            <begin position="297"/>
            <end position="411"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="17">
        <location>
            <begin position="832"/>
            <end position="900"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="17">
        <location>
            <begin position="930"/>
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        <location>
            <begin position="1020"/>
            <end position="1082"/>
        </location>
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        <location>
            <begin position="1191"/>
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            <begin position="2027"/>
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        <location>
            <begin position="2989"/>
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        <location>
            <begin position="4024"/>
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        <location>
            <begin position="4041"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00004138"/>
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    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007200"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
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            <dbReference type="ARBA" id="ARBA00022989"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00023069"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
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    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
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        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00152"/>
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    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
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        <source>
            <dbReference type="SAM" id="Phobius"/>
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    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50041"/>
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    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50093"/>
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    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50095"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50221"/>
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    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS51111"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS51212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030066.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="4152" mass="459400" checksum="D875898B7C471096" modified="2019-05-08" version="1">[encoded data, 4152 characters]</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ42</accession>
    <name>A0A452IQ42_9SAUR</name>
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        <submittedName>
            <fullName evidence="12">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
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            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
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        </lineage>
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        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
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                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
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        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
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    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
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        <scope>IDENTIFICATION</scope>
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        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
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        <scope>IDENTIFICATION</scope>
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        <subcellularLocation>
            <location evidence="1 7">Nucleus</location>
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        </location>
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            <dbReference type="Proteomes" id="UP000291020"/>
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</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ43</accession>
    <name>A0A452IQ43_9SAUR</name>
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        <recommendedName>
            <fullName evidence="2">Lipocalin/cytosolic fatty-acid binding domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
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        <lineage>
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            <taxon>Metazoa</taxon>
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            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
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        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the calycin superfamily. Fatty-acid binding protein (FABP) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQ43"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034035.1">
        <property type="protein sequence ID" value="ENSGAGP00000029968.1"/>
        <property type="gene ID" value="ENSGAGG00000021627.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Fatty acid-binding protein, adipocyte"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.128.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012674">
        <property type="entry name" value="Calycin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000463">
        <property type="entry name" value="Fatty_acid-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031259">
        <property type="entry name" value="ILBP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000566">
        <property type="entry name" value="Lipocln_cytosolic_FA-bd_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11955">
        <property type="entry name" value="FATTY ACID BINDING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00061">
        <property type="entry name" value="Lipocalin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00178">
        <property type="entry name" value="FATTYACIDBP"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50814">
        <property type="entry name" value="Lipocalins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Lipocalin/cytosolic fatty-acid binding" evidence="2">
        <location>
            <begin position="7"/>
            <end position="131"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00008390"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF00061"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029968.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="132" mass="14698" checksum="D412723BBCF32540" modified="2019-05-08" version="1">MVDHLLGTWKSTSCENFDAYMKELGVGLATRKLGKLAKPSVTISADGDVVTIQTKSAFKNNEISFKLGEEFDETTPDDRKTKSIVTLENGSLNQVQNWDGKETTIKRQVVDGKMVVECTMNDITCTRIYEKA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ44</accession>
    <name>A0A452IQ44_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">G-protein coupled receptors family 1 profile domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQ44"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034177.1">
        <property type="protein sequence ID" value="ENSGAGP00000030102.1"/>
        <property type="gene ID" value="ENSGAGG00000021717.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004930">
        <property type="term" value="F:G protein-coupled receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004984">
        <property type="term" value="F:olfactory receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1070.10">
        <property type="entry name" value="Rhodopsin 7-helix transmembrane proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000276">
        <property type="entry name" value="GPCR_Rhodpsn"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017452">
        <property type="entry name" value="GPCR_Rhodpsn_7TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000725">
        <property type="entry name" value="Olfact_rcpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR26453">
        <property type="entry name" value="OLFACTORY RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13853">
        <property type="entry name" value="7tm_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00237">
        <property type="entry name" value="GPCRRHODOPSN"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00245">
        <property type="entry name" value="OLFACTORYR"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81321">
        <property type="entry name" value="Family A G protein-coupled receptor-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50262">
        <property type="entry name" value="G_PROTEIN_RECEP_F1_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="7 9" id="KW-0472">Membrane</keyword>
    <keyword evidence="5" id="KW-0552">Olfaction</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0716">Sensory transduction</keyword>
    <keyword evidence="8" id="KW-0807">Transducer</keyword>
    <keyword evidence="4 9" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 9" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="164"/>
            <end position="185"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="206"/>
            <end position="225"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="237"/>
            <end position="256"/>
        </location>
    </feature>
    <feature type="domain" description="G-protein coupled receptors family 1 profile" evidence="10">
        <location>
            <begin position="38"/>
            <end position="254"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022606"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022725"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023224"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50262"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030102.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="322" mass="35752" checksum="BD375CE778A4903B" modified="2025-10-08" version="2">MNWLSLGEFILLGVSDQPQLELLLFVLILISQRTSLLGNTITILVSWLDPHLHMPMYFFLSNLSFLDLCYTTSFGPQMLLYITLSLGCTKCLLLAVMAYDRYIAVCQPLRYTAITSHHLCLQLAATDWLLPQCGQNQINNLFCEVPALLKLACVETSANEAKLLVSAVIFLLVPLGFILISYGYISLTVLKIHSAKGRLKAFNTCASHLVVVSLFYGTGISMYLQPPSSYTQHLGKMVPLFYTMVTLMLNPLIYTLRNKDVHREALQKVLRRSPTTQGIQAGAVVGHEIICTSFGFCSIFPTAPGCQLSLKKFLSVSLLMVG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ45</accession>
    <name>A0A452IQ45_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="11">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQ45"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034316.1">
        <property type="protein sequence ID" value="ENSGAGP00000030235.1"/>
        <property type="gene ID" value="ENSGAGG00000021690.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098794">
        <property type="term" value="C:postsynapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043083">
        <property type="term" value="C:synaptic cleft"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004930">
        <property type="term" value="F:G protein-coupled receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007189">
        <property type="term" value="P:adenylate cyclase-activating G protein-coupled receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007166">
        <property type="term" value="P:cell surface receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016525">
        <property type="term" value="P:negative regulation of angiogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016322">
        <property type="term" value="P:neuron remodeling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.220.50:FF:000027">
        <property type="entry name" value="Adhesion G protein-coupled receptor B3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.220.50">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1070.10">
        <property type="entry name" value="Rhodopsin 7-helix transmembrane proteins"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057244">
        <property type="entry name" value="GAIN_B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046338">
        <property type="entry name" value="GAIN_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017981">
        <property type="entry name" value="GPCR_2-like_7TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008077">
        <property type="entry name" value="GPCR_2_brain_angio_inhib"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000832">
        <property type="entry name" value="GPCR_2_secretin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017983">
        <property type="entry name" value="GPCR_2_secretin-like_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000203">
        <property type="entry name" value="GPS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12011:SF40">
        <property type="entry name" value="ADHESION G PROTEIN-COUPLED RECEPTOR B3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12011">
        <property type="entry name" value="ADHESION G-PROTEIN COUPLED RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00002">
        <property type="entry name" value="7tm_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01825">
        <property type="entry name" value="GPS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01694">
        <property type="entry name" value="BAIPRECURSOR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00249">
        <property type="entry name" value="GPCRSECRETIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00303">
        <property type="entry name" value="GPS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00650">
        <property type="entry name" value="G_PROTEIN_RECEP_F2_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50261">
        <property type="entry name" value="G_PROTEIN_RECEP_F2_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50221">
        <property type="entry name" value="GAIN_B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6 8" id="KW-0472">Membrane</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0732">Signal</keyword>
    <keyword evidence="2 8" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 8" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="87"/>
            <end position="108"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="120"/>
            <end position="140"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="152"/>
            <end position="169"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="189"/>
            <end position="209"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="273"/>
            <end position="295"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="302"/>
            <end position="322"/>
        </location>
    </feature>
    <feature type="domain" description="GAIN-B" evidence="9">
        <location>
            <begin position="1"/>
            <end position="75"/>
        </location>
    </feature>
    <feature type="domain" description="G-protein coupled receptors family 2 profile 2" evidence="10">
        <location>
            <begin position="83"/>
            <end position="327"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50261"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030235.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="698" mass="78033" checksum="AC446E0D2B19B526" modified="2019-05-08" version="1">MIQVLPLGTEKASGVRCQCIIIPISLSLSLLCRNDSLGTWSTQGCKTVLTDASHTKCLCDRLSTFAILAQQPREIIMESSGAPSVTLIVGSGLSCLALITLAVVYAALWRYIRSERSIILINFCLSIISSNILILVGQTQTHNKGICTTTTAFLHFFFLASFCWVLTEAWQSYMAVTGKIRTQLIRKRFLCLGWGLPALVVAISIETIFCNNACRHFLSLSLKGQLDGILDKKLKHRAGQMSEPHSGLTLKCAKCGVVSTTALSATTASNAMASLWSSCVVLPLLALTWMSAVLAMTDKRSILFQILFAVFDSLQGFVIVMVHCILRREVQDAFRCRLRNCQDPINADSSSSFPNGHAQIMTDFEKDVDIACRSVLHKDIGPCRAATITGTLSRISLNDDEEEKGANPEGMSYSTLPGSIISKVIIQQPAGLHMPMSMSELANQCLKKDNSELRRTVYLCTDDNLRGADMDIVNPQERMMESDYIVMPRGSVNTQQSLKDESKLNIGMDTLPHERLLHYKVNPEFNMNPSVMDQFNINLDQHLPTQEHMQNLPFEPRTAVKNFIASELDDNAGLARSETGSTISMSSLERRKSRYSDLDFEKVMHTRKRHMELFQELNQKFQTLDRFRDVPKSGSMENPTPNKAPWDSFKNTNEYQHYTTINVLDTEAKDALELRPAEWEKCLNLPLDVQEGDFQTEV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ46</accession>
    <name>A0A452IQ46_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">PDZ domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQ46"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034143.1">
        <property type="protein sequence ID" value="ENSGAGP00000030071.1"/>
        <property type="gene ID" value="ENSGAGG00000021685.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098887">
        <property type="term" value="P:neurotransmitter receptor transport, endosome to postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd06686">
        <property type="entry name" value="PDZ4_GRIP1-2-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd06682">
        <property type="entry name" value="PDZ5_GRIP1-2-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd06683">
        <property type="entry name" value="PDZ6_GRIP1-2-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd06685">
        <property type="entry name" value="PDZ7_GRIP1-2-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000022">
        <property type="entry name" value="Glutamate receptor interacting protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000025">
        <property type="entry name" value="Glutamate receptor interacting protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000034">
        <property type="entry name" value="Glutamate receptor interacting protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000035">
        <property type="entry name" value="Glutamate receptor interacting protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.42.10">
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043545">
        <property type="entry name" value="GRIP1/2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001478">
        <property type="entry name" value="PDZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041489">
        <property type="entry name" value="PDZ_6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036034">
        <property type="entry name" value="PDZ_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46227:SF3">
        <property type="entry name" value="GLUTAMATE RECEPTOR-INTERACTING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46227">
        <property type="entry name" value="GLUTAMATE RECEPTOR-INTERACTING PROTEIN GRIP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00595">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17820">
        <property type="entry name" value="PDZ_6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00228">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50156">
        <property type="entry name" value="PDZ domain-like"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50106">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="5">
        <location>
            <begin position="1"/>
            <end position="21"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019177497" description="PDZ domain-containing protein" evidence="5">
        <location>
            <begin position="22"/>
            <end position="609"/>
        </location>
    </feature>
    <feature type="domain" description="PDZ" evidence="6">
        <location>
            <begin position="31"/>
            <end position="106"/>
        </location>
    </feature>
    <feature type="domain" description="PDZ" evidence="6">
        <location>
            <begin position="132"/>
            <end position="217"/>
        </location>
    </feature>
    <feature type="domain" description="PDZ" evidence="6">
        <location>
            <begin position="232"/>
            <end position="314"/>
        </location>
    </feature>
    <feature type="domain" description="PDZ" evidence="6">
        <location>
            <begin position="483"/>
            <end position="565"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="334"/>
            <end position="382"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="413"/>
            <end position="453"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="413"/>
            <end position="436"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="443"/>
            <end position="453"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50106"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030071.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="609" mass="66544" checksum="1A374C39DD10E32C" modified="2019-05-08" version="1">MFFPLVCSVLVSLASSTVGLAGQVVHTETTEVVLTADPIVGFGIQLQGSVFATETLSSPPLISYIECDSPAERCGVLQIGDRLMAINGIPTEDSTFEEANQLLRDSSITNKVTLEIEFDVAESVIPSSGTFHVKLPKKHNVELGITISSPSSRKPGDPLVISDIKKGSVAHRTGTLELGDKLLAIDNIRLDNCSMEDAVQILQHCEDLVKLKIRKDEDNSDEQESSGAIIYTVELKRYGGPLGITISGTEEPFDPIIISSLTKGGLAERTGAIHIGDRILAINSSSLKGKPLSEAIHLLQMAGETVTLKIKKQTDGPTYQASGYNFSAYEWRSPKQRDNLSPPSRPQNHPFHDTGLSDDEWDRPTASGFSGAHDNSETDQEENFWSQALEDLETCGQSGILRELEATIMSGSTMSLNHENPQPRSQLGRQASFQERSNVRPHYSQTARSNTLPSDVGRKSVVLRKIKQEMKEIMSPTPVELHKVTLYKESDIEDFGFSVSDGLLEKGVYVKNIRPAGPGDLGGLKPYDRLLQVNHVRTRDFDCCLVVPLIAESGNKLELVISRNPLASQKVTSDQQTLAGGEWNDQNNAFLQQTGHTASVETRDPTNTI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ47</accession>
    <name>A0A452IQ47_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Ig-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the MHC class II family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQ47"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034182.1">
        <property type="protein sequence ID" value="ENSGAGP00000030107.1"/>
        <property type="gene ID" value="ENSGAGG00000021674.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042613">
        <property type="term" value="C:MHC class II protein complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002250">
        <property type="term" value="P:adaptive immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002504">
        <property type="term" value="P:antigen processing and presentation of peptide or polysaccharide antigen via MHC class II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd05767">
        <property type="entry name" value="IgC1_MHC_II_alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.320.10">
        <property type="entry name" value="Class II Histocompatibility Antigen, M Beta Chain, Chain B, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003006">
        <property type="entry name" value="Ig/MHC_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003597">
        <property type="entry name" value="Ig_C1-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050160">
        <property type="entry name" value="MHC/Immunoglobulin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011162">
        <property type="entry name" value="MHC_I/II-like_Ag-recog"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014745">
        <property type="entry name" value="MHC_II_a/b_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001003">
        <property type="entry name" value="MHC_II_a_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19944:SF86">
        <property type="entry name" value="HLA CLASS II HISTOCOMPATIBILITY ANTIGEN, DR ALPHA CHAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19944">
        <property type="entry name" value="MHC CLASS II-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07654">
        <property type="entry name" value="C1-set"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00993">
        <property type="entry name" value="MHC_II_alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00407">
        <property type="entry name" value="IGc1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00920">
        <property type="entry name" value="MHC_II_alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54452">
        <property type="entry name" value="MHC antigen-recognition domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00290">
        <property type="entry name" value="IG_MHC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-1064">Adaptive immunity</keyword>
    <keyword evidence="9" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="5" id="KW-0391">Immunity</keyword>
    <keyword evidence="8 12" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-0491">MHC II</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="3 12" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 12" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="214"/>
            <end position="231"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="13">
        <location>
            <begin position="70"/>
            <end position="192"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007394"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022859"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023130"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023182"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030107.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="246" mass="27708" checksum="30903D5BC131C4E8" modified="2019-05-08" version="1">GPLRKAVLLKGAPPTDCMGPGVVDHVTSQVVFYQHMDGSQQGSEKFTQEFDQDEILYVDLERKKVVWRLPEFSKFTSFEVQHALGNMAVIKLYQEKLIERSSHTRAQNGGTFPCQLGEPNVLICFMDKFFPPVLSLTWLKNGQKVTEGVYETDFYPHQDNSFRKFSYLPFFPSHGDFYDCQVEHRGLPKPFTKHWEAQLPTSVAETTETLVCTLGLAVGIIGIIAGTILIIKGMKMNAAHNPRGPL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ71</accession>
    <name>A0A452IQ71_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="22">E3 ubiquitin-protein ligase TRIM9</fullName>
            <ecNumber evidence="6">2.3.2.27</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="24">RING-type E3 ubiquitin transferase TRIM9</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="23">Tripartite motif-containing protein 9</fullName>
        </alternativeName>
    </protein>
    <organism evidence="31 32">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="32" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
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            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="31" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="31" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
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            <text>S-ubiquitinyl-[E2 ubiquitin-conjugating enzyme]-L-cysteine + [acceptor protein]-L-lysine = [E2 ubiquitin-conjugating enzyme]-L-cysteine + N(6)-ubiquitinyl-[acceptor protein]-L-lysine.</text>
            <dbReference type="EC" id="2.3.2.27"/>
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        <text evidence="5">Protein modification; protein ubiquitination.</text>
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        <text evidence="21">Interacts with SNAP25.</text>
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        <subcellularLocation>
            <location evidence="4">Cell projection</location>
            <location evidence="4">Dendrite</location>
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        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
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            <location evidence="2">Cytoplasmic vesicle</location>
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        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE TRIM36-RELATED"/>
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    <keyword evidence="17" id="KW-0175">Coiled coil</keyword>
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    <keyword evidence="8" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="32" id="KW-1185">Reference proteome</keyword>
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    <keyword evidence="16" id="KW-0770">Synapse</keyword>
    <keyword evidence="9" id="KW-0808">Transferase</keyword>
    <keyword evidence="15" id="KW-0832">Ubl conjugation</keyword>
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    <keyword evidence="12 25" id="KW-0863">Zinc-finger</keyword>
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        <location>
            <begin position="10"/>
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        </location>
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            <end position="430"/>
        </location>
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            <begin position="438"/>
            <end position="533"/>
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    <sequence length="708" mass="79246" checksum="26246FEEC8D0C99B" modified="2019-05-08" version="1">MDEMEEELKCPVCGSFYREPLILPCSHNLCQACARNILVQTPEAESPQSRRASGVSDYDYLDLDKMSLYSEADSGYGSYGGFASAPTTPCQKSPNGVRVFPPAAPPPPHLPPAALAPVPRNACLTCPQCHRSLILDERGLRGFPKNRVLEGVIDRYQQSKAAALRCQLCEKAPKEATVMCEQCDVFYCEPCRLRCHPPRGPLAKHRLVPPAQGRVSRRLSPRKISTCTDHELENHSMYCVQCKMPVCYQCLEEGKHSSHEVKALGAMWKLHKSQLSQALNGLSDRAKEAKEFLVQLRNMVQQIQENSVEFEACLVAQCDALIDALNRRKAQLLSRVNKEHEHKLKVVRDQISHCTVKLRQTTGLMEYCLEVIKENDPSGFLQISDALIRRVHLTEDQWGKGTLTPRMTTDFDLNLDSAPLLQSIHQLDFVQMKASSPVPAPPILQLEECCTHNNSATLSWKQPPLSTVQVEGYILELDDGNGGQFREVYVGKETMCTVDGLHFNSTYSARVKAFNKTGVSQYSKTLVLQTSEVAWFSFDPGSAHSDILFSNDNLTVTCNSYDDRVVLGKTGFSKGLHYWELSIDRYDNHPDPAFGVARVDVLKDVMLGKDDKAWAMYVDNNRSWFMHNNSHTNRTEGGITKGATVGVLLDLTRRTLTFSINEDQQGPVAFENMEGLFFPAVSLNRNVQVTLHTGLPVPEFYNSRASMQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ93</accession>
    <name>A0A452IQ93_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="22">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cell projection</location>
            <location evidence="1">Cilium</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the polycystin family.</text>
    </comment>
    <comment type="caution">
        <text evidence="13">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000034142.1">
        <property type="protein sequence ID" value="ENSGAGP00000030070.1"/>
        <property type="gene ID" value="ENSGAGG00000021465.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005929">
        <property type="term" value="C:cilium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005261">
        <property type="term" value="F:monoatomic cation channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006816">
        <property type="term" value="P:calcium ion transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00037">
        <property type="entry name" value="CLECT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00146">
        <property type="entry name" value="PKD"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="CDD" id="cd01752">
        <property type="entry name" value="PLAT_polycystin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000825">
        <property type="entry name" value="Polycystin 1, transient receptor potential channel interacting"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.60.20:FF:000012">
        <property type="entry name" value="polycystin-1 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.100.10">
        <property type="entry name" value="Mannose-Binding Protein A, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.60.20">
        <property type="entry name" value="PLAT/LH2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001304">
        <property type="entry name" value="C-type_lectin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016186">
        <property type="entry name" value="C-type_lectin-like/link_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016187">
        <property type="entry name" value="CTDL_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057244">
        <property type="entry name" value="GAIN_B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000203">
        <property type="entry name" value="GPS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000434">
        <property type="entry name" value="PC1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022409">
        <property type="entry name" value="PKD/Chitinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002859">
        <property type="entry name" value="PKD/REJ-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013122">
        <property type="entry name" value="PKD1_2_channel"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000601">
        <property type="entry name" value="PKD_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035986">
        <property type="entry name" value="PKD_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001024">
        <property type="entry name" value="PLAT/LH2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036392">
        <property type="entry name" value="PLAT/LH2_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042060">
        <property type="entry name" value="PLAT_polycystin1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006228">
        <property type="entry name" value="Polycystin_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046791">
        <property type="entry name" value="Polycystin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014010">
        <property type="entry name" value="REJ_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002889">
        <property type="entry name" value="WSC_carb-bd"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00864">
        <property type="entry name" value="PCC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46730">
        <property type="entry name" value="POLYCYSTIN-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46730:SF3">
        <property type="entry name" value="POLYCYSTIN-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00059">
        <property type="entry name" value="Lectin_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00801">
        <property type="entry name" value="PKD"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08016">
        <property type="entry name" value="PKD_channel"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01477">
        <property type="entry name" value="PLAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20519">
        <property type="entry name" value="Polycystin_dom"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02010">
        <property type="entry name" value="REJ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00500">
        <property type="entry name" value="POLYCYSTIN1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00034">
        <property type="entry name" value="CLECT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00303">
        <property type="entry name" value="GPS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00308">
        <property type="entry name" value="LH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00089">
        <property type="entry name" value="PKD"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="SMART" id="SM00321">
        <property type="entry name" value="WSC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56436">
        <property type="entry name" value="C-type lectin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49723">
        <property type="entry name" value="Lipase/lipooxygenase domain (PLAT/LH2 domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49299">
        <property type="entry name" value="PKD domain"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50041">
        <property type="entry name" value="C_TYPE_LECTIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50221">
        <property type="entry name" value="GAIN_B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50093">
        <property type="entry name" value="PKD"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50095">
        <property type="entry name" value="PLAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51111">
        <property type="entry name" value="REJ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51212">
        <property type="entry name" value="WSC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="12" id="KW-0966">Cell projection</keyword>
    <keyword evidence="8" id="KW-0969">Cilium</keyword>
    <keyword evidence="10" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9 15" id="KW-0472">Membrane</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5 15" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 15" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="2910"/>
            <end position="2932"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3118"/>
            <end position="3139"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3159"/>
            <end position="3181"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3340"/>
            <end position="3362"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3368"/>
            <end position="3387"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3455"/>
            <end position="3474"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3677"/>
            <end position="3698"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3710"/>
            <end position="3727"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3747"/>
            <end position="3770"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3796"/>
            <end position="3815"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3847"/>
            <end position="3871"/>
        </location>
    </feature>
    <feature type="domain" description="WSC" evidence="21">
        <location>
            <begin position="37"/>
            <end position="130"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="17">
        <location>
            <begin position="156"/>
            <end position="218"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="16">
        <location>
            <begin position="274"/>
            <end position="388"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="17">
        <location>
            <begin position="804"/>
            <end position="866"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="17">
        <location>
            <begin position="896"/>
            <end position="968"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="17">
        <location>
            <begin position="986"/>
            <end position="1048"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="17">
        <location>
            <begin position="1157"/>
            <end position="1230"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="17">
        <location>
            <begin position="1259"/>
            <end position="1314"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="17">
        <location>
            <begin position="1332"/>
            <end position="1396"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="17">
        <location>
            <begin position="1568"/>
            <end position="1654"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="17">
        <location>
            <begin position="1671"/>
            <end position="1731"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="17">
        <location>
            <begin position="1907"/>
            <end position="1990"/>
        </location>
    </feature>
    <feature type="domain" description="REJ" evidence="20">
        <location>
            <begin position="1993"/>
            <end position="2686"/>
        </location>
    </feature>
    <feature type="domain" description="GAIN-B" evidence="19">
        <location>
            <begin position="2707"/>
            <end position="2900"/>
        </location>
    </feature>
    <feature type="domain" description="PLAT" evidence="18">
        <location>
            <begin position="2955"/>
            <end position="3070"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="3924"/>
            <end position="3953"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="4050"/>
            <end position="4072"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="4102"/>
            <end position="4124"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="3941"/>
            <end position="3953"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="14">
        <location>
            <begin position="4107"/>
            <end position="4124"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004138"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007200"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023069"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00152"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50041"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50093"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50095"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS51111"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS51212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030070.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="4124" mass="456019" checksum="53C7B7BB61686CE5" modified="2019-05-08" version="1">[encoded data, 4124 characters]</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ97</accession>
    <name>A0A452IQ97_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">SAM domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the liprin family. Liprin-alpha subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQ97"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034375.1">
        <property type="protein sequence ID" value="ENSGAGP00000030287.1"/>
        <property type="gene ID" value="ENSGAGG00000020741.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048786">
        <property type="term" value="C:presynaptic active zone"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050808">
        <property type="term" value="P:synapse organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd09562">
        <property type="entry name" value="SAM_liprin-alpha1_2_3_4_repeat1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd09565">
        <property type="entry name" value="SAM_liprin-alpha1_2_3_4_repeat2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd09568">
        <property type="entry name" value="SAM_liprin-alpha1_2_3_4_repeat3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.50:FF:000003">
        <property type="entry name" value="liprin-alpha-2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.50:FF:000002">
        <property type="entry name" value="PTPRF interacting protein alpha 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.50:FF:000004">
        <property type="entry name" value="PTPRF interacting protein alpha 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.50">
        <property type="entry name" value="Transcription Factor, Ets-1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057892">
        <property type="entry name" value="LIP-1_CC2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037620">
        <property type="entry name" value="LIP-1_SAM_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037621">
        <property type="entry name" value="LIP-1_SAM_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037622">
        <property type="entry name" value="LIP-1_SAM_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029515">
        <property type="entry name" value="Liprin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001660">
        <property type="entry name" value="SAM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013761">
        <property type="entry name" value="SAM/pointed_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12587">
        <property type="entry name" value="LAR INTERACTING PROTEIN LIP -RELATED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12587:SF15">
        <property type="entry name" value="LIPRIN-ALPHA-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25526">
        <property type="entry name" value="LIP-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00536">
        <property type="entry name" value="SAM_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07647">
        <property type="entry name" value="SAM_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00454">
        <property type="entry name" value="SAM"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47769">
        <property type="entry name" value="SAM/Pointed domain"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50105">
        <property type="entry name" value="SAM_DOMAIN"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6 7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="SAM" evidence="9">
        <location>
            <begin position="919"/>
            <end position="985"/>
        </location>
    </feature>
    <feature type="domain" description="SAM" evidence="9">
        <location>
            <begin position="1040"/>
            <end position="1097"/>
        </location>
    </feature>
    <feature type="domain" description="SAM" evidence="9">
        <location>
            <begin position="1121"/>
            <end position="1190"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1"/>
            <end position="31"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="223"/>
            <end position="246"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="451"/>
            <end position="471"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="682"/>
            <end position="703"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="716"/>
            <end position="791"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="803"/>
            <end position="884"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1237"/>
            <end position="1272"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="97"/>
            <end position="131"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="266"/>
            <end position="388"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="494"/>
            <end position="542"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="237"/>
            <end position="246"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="682"/>
            <end position="692"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="722"/>
            <end position="731"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="768"/>
            <end position="777"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="803"/>
            <end position="812"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="813"/>
            <end position="822"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="1238"/>
            <end position="1249"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="1254"/>
            <end position="1272"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50105"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030287.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1272" mass="143826" checksum="B79B13DBC2FEC275" modified="2019-05-08" version="1">MMCEVMPTISEAEIPSGGNGGHGSGSPLQSDADSHFEQLMVSMLEERDRLLDTLRETQETLVLTQGKLHEVSHERDSLQRQLNTALPQEFAALTKELNVCREQLLEREEEIAELKAERNNTRLLLEHLECLVSRHERSLRMTVVKRQAQSPAGVSSEVEVLKALKSLFEHHKALDEKVRERLRVALERCSLLEEELGTTHKELMILKEQSNQKRTLTDGLLDINHDQENTPSTNGKRSSDGSSSHDEDLAKVIELQEIIDKQTNEQGQMKERLTALSSRVTELEEDLDTARKDLIKSEEMNTKLQRDVREAMAQKEDMEERITTLEKRYLAAQREATSVHDLNDKLENEIANKDSMHRQSEDKNRQLQERLELAEQKLQQTLRKAETLPEVEAELAQRVAALSKSDPLSSGNSAAKDAKLLELASKFRKAEERHGNIEERLRQMEAQLEEKNQELQRARQREKMNEEHNKRLSDTVDKLLSESNERLQLHLKERMAALEDKNSLLREVENAKKQVEDLQHEKDQLVLNIEAMRAENDQMRIRATSLHHSRPHLGSVPDFRYPMTPLSVADSHTDPYSTSSVLRRPQKGRLAALRDEPSKVQTLNEQDWERAQQASVLANVAQAFESDVDVSDGDDDGETIFSSVDLLSPSGQADAQTLAMMLQEQLDAINKEIRLIQEEKENTEQRAEEIESRVGSGSLDNNGRFRSMNSIPLPFIGGTLAGSSPPGSGRSTPRRILHSPAREVDRLGIMTLPSDLRKHRRKSPVSGEEVRDDKTTIKCETSPPSSPRSLYLDRVHKGALHTLSHEDIRDIRNSTGSQDGQVSNPSSSNSSQDSLHKAPKKKGIKSSIGRLFGKKEKGRPGQMSKESLGQVGLAETDNSSQDALGLSKLGGQAEKNRKLQKKHELLEEARRQGLPFAQWDGPTVVVWLELWVGMPAWYVAACRANVKSGAIMSALSDTEIQREIGISNPLHRLKLRLAIQEIMSLTSPSAPPTSRTTTGNVWVTHEEMENLTATPQTVSYSTATSTLAYGDMNHEWIGNEWLPSLGLPQYRSYFMECLVDARMLDHLTKKDLRGQLKMVDSFHRNSFQCGIMCLRKLNYDRKELERRREESQNEIKDVLVWSNDRMIRWVLSIGLKEYANNLIESGVHGALVALEETFDFSALALLLQIPTQNTQARAVLEREFNNLLVMGTDRRFDEDDDKSFRRAPSWRKKFRPKDIRGLAAGSAETLPANFRVTSSMSSPSMQPKKMQIDGNVSGTQRLDSTTVRTYSC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ99</accession>
    <name>A0A452IQ99_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">APC membrane recruitment protein 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Peripheral membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the Amer family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQ99"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034243.1">
        <property type="protein sequence ID" value="ENSGAGP00000030163.1"/>
        <property type="gene ID" value="ENSGAGG00000021765.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008013">
        <property type="term" value="F:beta-catenin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005546">
        <property type="term" value="F:phosphatidylinositol-4,5-bisphosphate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060828">
        <property type="term" value="P:regulation of canonical Wnt signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016055">
        <property type="term" value="P:Wnt signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019003">
        <property type="entry name" value="AMER"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22237:SF0">
        <property type="entry name" value="APC MEMBRANE RECRUITMENT PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22237">
        <property type="entry name" value="APC MEMBRANE RECRUITMENT PROTEIN 2-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09422">
        <property type="entry name" value="AMER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="5" id="KW-0446">Lipid-binding</keyword>
    <keyword evidence="6" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0879">Wnt signaling pathway</keyword>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="36"/>
            <end position="210"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="419"/>
            <end position="452"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="41"/>
            <end position="50"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="159"/>
            <end position="168"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="424"/>
            <end position="433"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004202"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007750"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022687"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023121"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030163.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="10">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="533" mass="57461" checksum="8AC69777C9BE4B19" modified="2025-10-08" version="2">MSSHSKSTAELEIAPPGKLKKTSFKLFGGKRSICTLPSFFGGKNKSQGKGASKKGLSKSKTHDGISDVTYEDGTGRLRLESPLNGGMDSHSRPLPSSQSAHLATDASVKFEFGRQESSHLGSAEGFEKKPNGDKSLSFPRPKKGLKGLFNSIRRHRKNKTAESEKAEWAADCGDGEQAKKTRGAGAETQRASEEESPRGTPLPGPCTGNLDGNCSLSTTTNLGENADWLMAAQSSSEGDAAVVPVGKDDILDVKLDAETIVCASSPYGGLPDAHHADYPDNDPPSVYSGDQLSLIFEDVTSLKSFDSLTGCGDIIAEPEIDSIAESTISVERSRDAAKRSSCLVTYQGGGEEMATPEEIEEYLQQLWEGTTEVDSRYETKLPQPINSPKLQGVNSKLETCGLHEEAAHLYAGGVMDDVELLTPPSDQQESAPNSDEGYYDSTTPGPEDELGEIKKDRLPRDSYSGDALYEFYEPDDTLMSPSLGDEPLFESKASHPEIFSLSFPDSGINPYLARAQVPLTSMGHKYMFKYCPK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1R2</accession>
    <name>A0A452J1R2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="23">Glutamate receptor</fullName>
        </recommendedName>
    </protein>
    <organism evidence="26 27">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="27" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="26" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="26" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="23">Receptor for glutamate that functions as a ligand-gated ion channel in the central nervous system and plays an important role in excitatory synaptic transmission. L-glutamate acts as an excitatory neurotransmitter at many synapses in the central nervous system.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>Ca(2+)(in) = Ca(2+)(out)</text>
            <dbReference type="Rhea" id="RHEA:29671"/>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="18 23">Postsynaptic cell membrane</location>
            <topology evidence="18 23">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="23">Belongs to the glutamate-gated ion channel (TC 1.A.10.1) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1R2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038866.1">
        <property type="protein sequence ID" value="ENSGAGP00000034324.1"/>
        <property type="gene ID" value="ENSGAGG00000024385.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0022824">
        <property type="term" value="F:transmitter-gated monoatomic ion channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007166">
        <property type="term" value="P:cell surface receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd06387">
        <property type="entry name" value="PBP1_iGluR_AMPA_GluR3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd13715">
        <property type="entry name" value="PBP2_iGluR_AMPA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.70:FF:000067">
        <property type="entry name" value="glutamate receptor 2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.70:FF:000099">
        <property type="entry name" value="glutamate receptor 2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.190.10:FF:000001">
        <property type="entry name" value="Glutamate receptor ionotropic, kainate 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.2300:FF:000004">
        <property type="entry name" value="Glutamate receptor, ionotropic, AMPA 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.190.10:FF:000666">
        <property type="entry name" value="Glutamate receptor, ionotropic, AMPA 2a"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.70">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.2300">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.190.10">
        <property type="entry name" value="Periplasmic binding protein-like II"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001828">
        <property type="entry name" value="ANF_lig-bd_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019594">
        <property type="entry name" value="Glu/Gly-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001508">
        <property type="entry name" value="Iono_Glu_rcpt_met"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015683">
        <property type="entry name" value="Ionotropic_Glu_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001320">
        <property type="entry name" value="Iontro_rcpt_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028082">
        <property type="entry name" value="Peripla_BP_I"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18966">
        <property type="entry name" value="IONOTROPIC GLUTAMATE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01094">
        <property type="entry name" value="ANF_receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00060">
        <property type="entry name" value="Lig_chan"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10613">
        <property type="entry name" value="Lig_chan-Glu_bd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00177">
        <property type="entry name" value="NMDARECEPTOR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00918">
        <property type="entry name" value="Lig_chan-Glu_bd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00079">
        <property type="entry name" value="PBPe"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53822">
        <property type="entry name" value="Periplasmic binding protein-like I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53850">
        <property type="entry name" value="Periplasmic binding protein-like II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81324">
        <property type="entry name" value="Voltage-gated potassium channels"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 23" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11 22" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="17 23" id="KW-0407">Ion channel</keyword>
    <keyword evidence="8 23" id="KW-0406">Ion transport</keyword>
    <keyword evidence="15 23" id="KW-1071">Ligand-gated ion channel</keyword>
    <keyword evidence="16" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="9 23" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0564">Palmitate</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="14 23" id="KW-0628">Postsynaptic cell membrane</keyword>
    <keyword evidence="12 23" id="KW-0675">Receptor</keyword>
    <keyword evidence="27" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 23" id="KW-0732">Signal</keyword>
    <keyword evidence="7 23" id="KW-0770">Synapse</keyword>
    <keyword evidence="4 23" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 23" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="1 23" id="KW-0813">Transport</keyword>
    <feature type="signal peptide" evidence="23">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5027164192" description="Glutamate receptor" evidence="23">
        <location>
            <begin position="23"/>
            <end position="888"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="23">
        <location>
            <begin position="545"/>
            <end position="567"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="23">
        <location>
            <begin position="630"/>
            <end position="652"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="23">
        <location>
            <begin position="820"/>
            <end position="842"/>
        </location>
    </feature>
    <feature type="domain" description="Ionotropic glutamate receptor C-terminal" evidence="24">
        <location>
            <begin position="418"/>
            <end position="795"/>
        </location>
    </feature>
    <feature type="domain" description="Ionotropic glutamate receptor L-glutamate and glycine-binding" evidence="25">
        <location>
            <begin position="428"/>
            <end position="493"/>
        </location>
    </feature>
    <feature type="binding site" evidence="20">
        <location>
            <position position="502"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="20">
        <location>
            <position position="504"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="20">
        <location>
            <position position="509"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="20">
        <location>
            <position position="680"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="20">
        <location>
            <position position="681"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="20">
        <location>
            <position position="731"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="site" description="Interaction with the cone snail toxin Con-ikot-ikot" evidence="21">
        <location>
            <position position="477"/>
        </location>
    </feature>
    <feature type="site" description="Crucial to convey clamshell closure to channel opening" evidence="21">
        <location>
            <position position="659"/>
        </location>
    </feature>
    <feature type="site" description="Interaction with the cone snail toxin Con-ikot-ikot" evidence="21">
        <location>
            <position position="686"/>
        </location>
    </feature>
    <feature type="site" description="Interaction with the cone snail toxin Con-ikot-ikot" evidence="21">
        <location>
            <position position="778"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="22">
        <location>
            <begin position="85"/>
            <end position="334"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="22">
        <location>
            <begin position="744"/>
            <end position="799"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023139"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023257"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00034104"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00036634"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="RuleBase" id="RU367118"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="SMART" id="SM00079"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="SMART" id="SM00918"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034324.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="888" mass="100146" checksum="21B62B834EA0BC50" modified="2019-05-08" version="1">MGQPVPQTLLLLVAGLLGESLGGFPNTISIGGLFMRNTVQEHSAFRFAVQLYNTNQNITEKPFHLNYHVDHLDSSNSFSVTNAFCSQFSRGVYAIFGFYDQMSMNTLTSFCGALHTSFVTPSFPTDADVQFVIQMRPALKGAILSLLTHYKWEKFVYLYDTERGFSILQAIMEAAVQNNWQVTARSVGSIKDVQEFRRIIEEMDRRQEKRYLIDCEVERINTILEQVVILGKHSRGYHYMLANLGFTDIILQRVMHGGANVTGFQIVNSENPMVQQFLQRWIRLDEREFPEAKNSPLKYTSALTHDAILVIAEAFRYLRRQRVDVSRRGSAGDCLANPAVPWSQGIDIERALKMVQVQGMTGNIQFDTYGRRTNYTIDVYETKATGSRKAGYWNEYERFVPALDQQVSNDTSSVENRTIVVTTILESPYVMYKKNHEQLEGNERYEGYCVDLAYEIAKHVGIKYKLSIVGDGKYGARDPESKIWNGMVGELVYGRADIAVAPLTITLVREEVIDFSKPFMSLGISIMIKKPQKSKPGVFSFLDPLAYEIWMCIVFAYIGVSVVLFLVSRFSPYEWHLEDNNEEPRDPQNPPDPPNEFGIFNSLWFSLGAFMQQGCDISPRSLSGRIVGGVWWFFTLIIISSYTANLAAFLTVERMVSPIESAEDLAKQTEIAYGTLDSGSTKEFFRRSKIAVYEKMWSYMKSAEPSVFTKTTADGVARVRKSKGKFAFLLESTMNEYIEQRKPCDTMKVGGNLDSKGYGVATPKGSALRNAVNLAVLKLNEQGLLDKLKNKWWYDKGECGSGGGDSKDKTSALSLSNVAGVFYILVGGLGLAMMVALIEFCYKSRAESKRMKLTKNTQNFKPAPATNTQNYATYREGYNVYGTESVKI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1R5</accession>
    <name>A0A452J1R5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Eyes absent homolog</fullName>
            <ecNumber evidence="15">3.1.3.48</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="12 15">
            <text>O-phospho-L-tyrosyl-[protein] + H2O = L-tyrosyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:10684"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="EC" id="3.1.3.48"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="14 15">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
        <text evidence="14 15">Binds 1 Mg(2+) ion per subunit.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 15">Belongs to the HAD-like hydrolase superfamily. EYA family.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.48" evidence="15"/>
    <dbReference type="AlphaFoldDB" id="A0A452J1R5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038884.1">
        <property type="protein sequence ID" value="ENSGAGP00000034342.1"/>
        <property type="gene ID" value="ENSGAGG00000024390.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:2001240">
        <property type="term" value="P:negative regulation of extrinsic apoptotic signaling pathway in absence of ligand"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045739">
        <property type="term" value="P:positive regulation of DNA repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd02601">
        <property type="entry name" value="HAD_Eya"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.12350:FF:000001">
        <property type="entry name" value="Eyes absent homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.12350">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028472">
        <property type="entry name" value="EYA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006545">
        <property type="entry name" value="EYA_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042577">
        <property type="entry name" value="EYA_dom_metazoan"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038102">
        <property type="entry name" value="EYA_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036412">
        <property type="entry name" value="HAD-like_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01658">
        <property type="entry name" value="EYA-cons_domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10190">
        <property type="entry name" value="EYES ABSENT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10190:SF7">
        <property type="entry name" value="EYES ABSENT HOMOLOG 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00702">
        <property type="entry name" value="Hydrolase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDG01129">
        <property type="entry name" value="C1.5:_HAD__Beta-PGM__Phosphata"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDS00003">
        <property type="entry name" value="Haloacid_Dehalogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56784">
        <property type="entry name" value="HAD-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0010">Activator</keyword>
    <keyword evidence="3" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="5 15" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="6 14 15" id="KW-0460">Magnesium</keyword>
    <keyword evidence="4 14 15" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="7 15" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0804">Transcription</keyword>
    <keyword evidence="8 15" id="KW-0805">Transcription regulation</keyword>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="186"/>
            <end position="237"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="186"/>
            <end position="198"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="16">
        <location>
            <begin position="208"/>
            <end position="223"/>
        </location>
    </feature>
    <feature type="active site" description="Nucleophile" evidence="13">
        <location>
            <position position="248"/>
        </location>
    </feature>
    <feature type="active site" description="Proton donor" evidence="13">
        <location>
            <position position="250"/>
        </location>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="248"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="250"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="476"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010501"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00051722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PIRSR" id="PIRSR628472-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PIRSR" id="PIRSR628472-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="RuleBase" id="RU362036"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034342.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="512" mass="56793" checksum="622678A9D098CE28" modified="2019-05-08" version="1">MIPASCVTWQLACPGRCCRSPHCGLWGLQRRHHPHIVTVPSFPTMATYGQTQYSAGIQQAATYAAYPPPGQPYGIPSYSIKTEDSLSHSPGQSGFLSYGSTYSTPTTGQAPYTYQMHGTSGIYQGANGLTNSAGFSTVHQEYSSYPSFPQSQYSQYYTSSYNSPYVSANSISPSAIPTSTYSLQESSHNITSHSTESLSGEYATTPVKDTETDRQHRGSDGKLRGRSKRSNDPSPTVDNEIERVFVWDLDETIIIFHSLLTGTFASRYGKDTTTSVRIGLMMEEMIFNLADTHLFFNDLEDCDQIHIDDVSSDDNGQDLSTYNFSADGFHSSTAGANLCLGSGVHGGVDWMRKLAFRYRRVKEMYNTYKNNVGGLIGAPKRETWLQLRAELEALTDLWLTHALKALNLIHSRPNCVNVLVTTTQLIPALAKVLLYGLGTVFPIENIYSATKTGKESCFERIMQRFGRKAVYVVIGDGVEEEQGAKKHNMPFWRISCHADLEALRHALELEYL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1S4</accession>
    <name>A0A452J1S4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Ankyrin and armadillo repeat containing</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000038876.1">
        <property type="protein sequence ID" value="ENSGAGP00000034334.1"/>
        <property type="gene ID" value="ENSGAGG00000024398.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043379">
        <property type="entry name" value="ANKAR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000225">
        <property type="entry name" value="Armadillo"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46464">
        <property type="entry name" value="ANK_REP_REGION DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46464:SF1">
        <property type="entry name" value="ANKYRIN AND ARMADILLO REPEAT-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13637">
        <property type="entry name" value="Ank_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00185">
        <property type="entry name" value="ARM"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50176">
        <property type="entry name" value="ARM_REPEAT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="repeat" description="ANK" evidence="1">
        <location>
            <begin position="550"/>
            <end position="582"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="1">
        <location>
            <begin position="619"/>
            <end position="651"/>
        </location>
    </feature>
    <feature type="repeat" description="ARM" evidence="2">
        <location>
            <begin position="723"/>
            <end position="765"/>
        </location>
    </feature>
    <feature type="repeat" description="ARM" evidence="2">
        <location>
            <begin position="764"/>
            <end position="806"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="1"/>
            <end position="24"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="49"/>
            <end position="68"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="514"/>
            <end position="546"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="524"/>
            <end position="545"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00259"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034334.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="6">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1414" mass="158040" checksum="8ACFBFD98FE14CB0" modified="2019-05-08" version="1">MTSASCTAPPAGVAKGKRRRPALLRFPPGQVPRASAGWAAGLPPFASTTGQSGAWPPMETPTSPPACLDSNYDHSTSCQLPPAYIDTRIGQILISVDYMLKALWHGVYMPKEKRVRFSELWRSSMDIDMDGNPQTEKNIFSEFCSAGLVDLSNEPDFEGIYDENMNEDPTYDPNSAQEKNLFMQYADNILFKLTFGTIQVQQHESVCKFDAAYVLSNVIRLTEDHLDTVTYQRLQQRLILQQKLVKKHLEKKAEIRKNIEYLKIVSFLIPFLLGLKRKMKIPNLNQLLQPYSDDKVKTERELPPFVFGPDFKCQHFHYIKNEYFHVHGGIEFDVGTPSLEDVSEKIKAVFDDIQICASNHVAQLLDPDIPYREHYSIPTMEFDGKSYYVISIELETFYQQLYKTPWWGAINEIISTLKPKRLPLTDIQSLEQFKKRFGYKKAIKCKSLPYGMKSAAERGLSAIFHTFCRKTSTSSLSVVDEYGYALLHHASIHNRVPIICQLIKAHLNINQRRNVHFNPGRRSQKSETKGDRAGKPDLQEKETSGERTALGPTALHLAAQCGSIEALTCLLAFKADYKLTDDRGWMAIHFAAFYDNISCITVLYRKDPDLLEAETTAEYHSTPLLLAATSGALDTLQYLFSLGANWKKTDSEGNNIIHLAVLYFHTEILKHLIQLDNPDLPVWNTLVEMLQCEDSMRPEMAVRSLEVLCLAKDFYWKCILDSGAIPSLINLLKGHQIKLACLTSGVLSNISPHITISKALVEAGGISVLIELLGSGEPELQSRCAVILYDIAQFDNNQNVISELGGIPPLINLLKIRLQDLLVNVINCIRVLCIQNKQNQITVKEHQGIPALVEFLTSQSDVLQAVSSAALAELARGNEKMQDAIADADAIGPLVELLRGRKITVQVKGAMAIEALSDNNTHIQRLFLEKSATKYLLKLLKAFHLTVKEQGATTLWALAGQTLKQQKLMAEQIGYNFIIDMLLSPSDKMQYVGGEAVIALSKDSKLHQNQICEGNGIGPLVRLLRSTKIAEGTLLSVIRALGTICIGVAHTNNPVSQENIVDEQALPILVHLLKNNNSLQIKVEVAYSLACIVLRNSNLQTVLQEEEGFNYKDVLELLNTPDKDICLRAGYALALFAYNNPVQQFLILETGAIMMSTFEPFLRSEIETHKAMAAFQIVVLARVIVDVDQVTLSARGVTILVELLNSEKTATLVLTGKLLASLAHTRAGVPEAITELGTIKRLCYHLYSDKEEVRIASAGALGYLTFNRTAYRHLLVECRNKPKQFTRLMNNLSRDAKISQAFVKEFQRQRQVGLPSLSLVINGGPPAVPTYTKERSRSSSSRRQFKAKFHPKDSLLLMPISAHLMGKLRTTDKPHSRTTGLHSFAHATSDITKVSRPRIFNVSKIRLNSPGKKA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1S5</accession>
    <name>A0A452J1S5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">PHD-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000038894.1">
        <property type="protein sequence ID" value="ENSGAGP00000034352.1"/>
        <property type="gene ID" value="ENSGAGG00000024403.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032922">
        <property type="term" value="P:circadian regulation of gene expression"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034732">
        <property type="entry name" value="EPHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052440">
        <property type="entry name" value="Trans_Reg/Chrom_Remod"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14955">
        <property type="entry name" value="RETINOIC ACID INDUCED 1/TRANSCRIPTION FACTOR 20"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14955:SF6">
        <property type="entry name" value="RETINOIC ACID-INDUCED PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13771">
        <property type="entry name" value="zf-HC5HC2H"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51805">
        <property type="entry name" value="EPHD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="1" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0862">Zinc</keyword>
    <keyword evidence="3" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="PHD-type" evidence="6">
        <location>
            <begin position="1793"/>
            <end position="1907"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="1"/>
            <end position="39"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="177"/>
            <end position="237"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="261"/>
            <end position="282"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="318"/>
            <end position="397"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="442"/>
            <end position="492"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="516"/>
            <end position="550"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="638"/>
            <end position="661"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="789"/>
            <end position="811"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="871"/>
            <end position="915"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="990"/>
            <end position="1009"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="1189"/>
            <end position="1239"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="1283"/>
            <end position="1302"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="1504"/>
            <end position="1597"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="1732"/>
            <end position="1786"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="1809"/>
            <end position="1829"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="12"/>
            <end position="24"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="177"/>
            <end position="213"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="5">
        <location>
            <begin position="263"/>
            <end position="277"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="5">
        <location>
            <begin position="322"/>
            <end position="336"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="351"/>
            <end position="364"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="381"/>
            <end position="397"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="463"/>
            <end position="473"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="516"/>
            <end position="542"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="999"/>
            <end position="1008"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="1513"/>
            <end position="1535"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="5">
        <location>
            <begin position="1560"/>
            <end position="1571"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="5">
        <location>
            <begin position="1578"/>
            <end position="1587"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="5">
        <location>
            <begin position="1739"/>
            <end position="1748"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="1753"/>
            <end position="1764"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="1777"/>
            <end position="1786"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="1809"/>
            <end position="1819"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS51805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034352.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1907" mass="209070" checksum="1C07571DADFB2EA6" modified="2019-05-08" version="1">MQSFRERCGFHGNQQNYQQTSQDTSRLENYRHQSQAGPNCERQRLVAKEYYSQQQLPYPGYENSAVEKYHRGNKQLPGQQLQGRPAFSNYAVQENSPYPARYSGEESLQPWGAQQQSLAGGVAKYEENLMKKSPAPAGSRQYHEQTAQLPFRTHSLHLPQQPPQPPALTYPKLQRQKIQNDVSSPMSFSQTGHFTQHSQSQFPASSTYSSVPGGSQAAHSYKSCTAPSSQQHERPLGGAANLASGQRVQNLHGYQANRISYDQQQQPQQQPQQQPLQGRHHAQETLHYQNLAKYQHYNQPGQTYCQTDAPVRTPEQYYQTFSPSSSHSPARSVGRSPSYSSTPSPLMPNLENFQYSQQSLNTGSFPAGITDHSHFMPLLNPSPTDGTSPDAQSGNCKNLQKEKIPENLLSDLSLQSLTALTSQVENISNTVQQLLLSKSAVPQKKGIKNPPRTPEQLKGQHCSPESSTYSAEQVGTPLSDPLSTPQSVHAEAQDADYLSGSEDQLERNFLYCGQSRSPARVNSNSKAKPESVSTCSVTSPDDMSTKSDDSFQSIHASLPLETFTKYMTNERECPRLLLSTLSQEELASEIIVLQDAINEKADKAWANSPALNKDSAKSPFHVENHRTCLDSMVKSTWSNQGNSGTHTEPLKLDKASGGNNSKDFSEEVYENAQVEFTTTETKNALKVTGSLAYNSKPNIPAATSSSGAASYSCYSNTTANSVDSENAMESFDWPDESISESCLRWKELGPCLQSSDLSKGLFHSKLAGSCEEKKNACCMDICDGEQPAKNEQAEDFSQQQMGEEEEDTLTYDEATKADSERWLEDTRHCCSGGDFSEIPMISSPDLKESDLEPEEYSSLCELASSEQKSVIYDTSPPKPPENAAVLSSKDVPVSAEETVSAVEKENSGPSSRLSGQSVILLGPAVGTETKVKSWFESSLPHIKPEEDTTVGESVLQEKTETETTLLVKVKKQMLPENLLVKSESISRGKTLRSKRVHSRLSEGEESRKPVPSLFKDIQAAGMVANTCVGPDGPMETTSKNAHGQTPRFPNEGLPARMCTRSFTALSEPRMPAPLEGLKVLAHQEKLGKKPVCGMKQRVAFKARKRTGRPAPKVVQNPSDPLSLLVPNLVQNEDFIGQNAKGLEPHDAEVKDQRSMILRSRTKTQEVYTKRRREKRTAEVRIKNCKTPKKLLSNNHLPPTFKITPQGGPHKEGKVGKRMKLPKSGPGMSSKMSERPLHSLKRKSTFVSPIPTKKRNLVLRSSSSSTKEEKAEASPNLFKRMPSTKKARAKLPTKSSCEAALMPPPAKEPPNVCIKITSRAAFQGAMKTKVLPPRKGRGLKLEAIVQKITSPNLKKFACKTAAVTATAHSNPLSPAGAEKEQALKNASVTPAVGEAGPLNQGMAQKAPAASVAEQLCRNSNNRSLKGKLMNSKKLSSDCFKGEAYSSPETVQYNSSSMAAKSISVLPKKRNRKGKAAAFSVAKNPLDRRAHLAPALLLTARERAGAGDALGNGDDGQRDGKKPKTEDKDFAKKEPPEGRSSQTQTRAQKQRANHSNYNGYSKRQRKRHTHSKAKNVASRCKSRAKRRRQQQAPLLNPTEPEIRLKYISCKRLRTDSRAPPFSPYVRVEKHNEFTTTCTVINSAGEEARLRKEQQPSSSAQALLPGGASLQPKAALPLSSTMHLGPVVSKALNAACLVCCLCRNPANYKDLGDLCGPYYPEHCLPKKKSRLKEKIKVEGPSEEAPVPSPAERTLKTTDNNCTTSTISGKPPRLDSAADSAKQSALRSSSRGMFRKLQSCYCCDERTEGEEAAEKPRRHECNKAESPSQESVGDTQEHWVHEACAVWTAGVYLVAGKLYGLQEAIKVAAEVRCSSCQQAGATVGCCHKGCVQTYHYACAIDTGSKMPIGVL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1S8</accession>
    <name>A0A452J1S8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Sodium/calcium exchanger membrane region domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>Ca(2+)(out) + K(+)(out) + 4 Na(+)(in) = Ca(2+)(in) + K(+)(in) + 4 Na(+)(out)</text>
            <dbReference type="Rhea" id="RHEA:69967"/>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
            <dbReference type="ChEBI" id="CHEBI:29103"/>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the Ca(2+):cation antiporter (CaCA) (TC 2.A.19) family. SLC24A subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1S8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038778.1">
        <property type="protein sequence ID" value="ENSGAGP00000034249.1"/>
        <property type="gene ID" value="ENSGAGG00000024329.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005262">
        <property type="term" value="F:calcium channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008273">
        <property type="term" value="F:calcium, potassium:sodium antiporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015293">
        <property type="term" value="F:symporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098703">
        <property type="term" value="P:calcium ion import across plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006874">
        <property type="term" value="P:intracellular calcium ion homeostasis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060292">
        <property type="term" value="P:long-term synaptic depression"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060291">
        <property type="term" value="P:long-term synaptic potentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1420.30:FF:000002">
        <property type="entry name" value="Sodium/potassium/calcium exchanger 2 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1420.30:FF:000004">
        <property type="entry name" value="Sodium/potassium/calcium exchanger 2 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1420.30">
        <property type="entry name" value="NCX, central ion-binding region"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004481">
        <property type="entry name" value="K/Na/Ca-exchanger"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004837">
        <property type="entry name" value="NaCa_Exmemb"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044880">
        <property type="entry name" value="NCX_ion-bd_dom_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00367">
        <property type="entry name" value="calcium/sodium antiporter"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10846">
        <property type="entry name" value="SODIUM/POTASSIUM/CALCIUM EXCHANGER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10846:SF41">
        <property type="entry name" value="SODIUM_POTASSIUM_CALCIUM EXCHANGER 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01699">
        <property type="entry name" value="Na_Ca_ex"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0050">Antiport</keyword>
    <keyword evidence="9" id="KW-0106">Calcium</keyword>
    <keyword evidence="6" id="KW-0109">Calcium transport</keyword>
    <keyword evidence="15" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13" id="KW-0406">Ion transport</keyword>
    <keyword evidence="14 19" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-0630">Potassium</keyword>
    <keyword evidence="5" id="KW-0633">Potassium transport</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="16" id="KW-0915">Sodium</keyword>
    <keyword evidence="16" id="KW-0739">Sodium transport</keyword>
    <keyword evidence="10" id="KW-0769">Symport</keyword>
    <keyword evidence="7 19" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="12 19" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="36"/>
            <end position="60"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="136"/>
            <end position="155"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="176"/>
            <end position="200"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="206"/>
            <end position="228"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="240"/>
            <end position="259"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="457"/>
            <end position="473"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="485"/>
            <end position="508"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="520"/>
            <end position="542"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="587"/>
            <end position="610"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="617"/>
            <end position="636"/>
        </location>
    </feature>
    <feature type="domain" description="Sodium/calcium exchanger membrane region" evidence="20">
        <location>
            <begin position="141"/>
            <end position="282"/>
        </location>
    </feature>
    <feature type="domain" description="Sodium/calcium exchanger membrane region" evidence="20">
        <location>
            <begin position="486"/>
            <end position="634"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="414"/>
            <end position="444"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="18">
        <location>
            <begin position="434"/>
            <end position="443"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005364"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022449"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022538"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022568"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022847"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022958"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023201"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00033627"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="Pfam" id="PF01699"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034249.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="647" mass="72647" checksum="92797F1D2B95B230" modified="2019-05-08" version="1">MALSKNIAASILEEWCLEEPLFRRRRHQNVRKKLRLIRIIGLFMSIMAISTFSLSISAFFKMEPHSTEMTSNLGSQKSMHGHQRTLLDVKEQNQNGTPDSPISMKYETEHNNATEEHSKGEYPKDLFSLAERRRGAVILHIMGMIYMFIALAIVCDEFFVPSLTVITEKLAISDDVAGATFMAAGGSAPELFTSLIGVFISHSNVGIGTIVGSAVFNILFVIGMCALFSREILNLTWWPLFRDVSFYIMDLILLIIFFLDNLIMWWESLTLLIAYFCYVTFMKFNVQVEECVKKILNRNKVEKVTTADSEQKTKPRLQRGGSSASLHNSLMRNSIFQLMIHTLDPLAEELGSYGKLKYYDSMTEEGKFKEKASILHKIAKKKCQVEDSERWNGAANHAEKIELPNTTKVEVEVTPPNDGSGPVQNGNIAHGIEENSEDNDEEQPLSLAWPDTLRKQVMYLAVFPIVFPLWITLPDVRKPSSRKFFPVTFFGAITWIAVFSYLMVWWAHQVGETIGISEEIMGLTILAAGTSIPDLITSVIVARKGLGDMAVSSSVGSNIFDITVGLPLPWLLYAAINGFAPVTVSSNGLFCAIVLLFIMLLFVILSIALCKWRMNKFLGFTMFGLYFIFLIVSVLLEDKVILCPVSI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1V0</accession>
    <name>A0A452J1V0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">EF-hand domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="1">Myosin is a hexamer of 2 heavy chains and 4 light chains.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1V0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038921.1">
        <property type="protein sequence ID" value="ENSGAGP00000034378.1"/>
        <property type="gene ID" value="ENSGAGG00000024425.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031672">
        <property type="term" value="C:A band"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016460">
        <property type="term" value="C:myosin II complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003785">
        <property type="term" value="F:actin monomer binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060048">
        <property type="term" value="P:cardiac muscle contraction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00051">
        <property type="entry name" value="EFh"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.238.10:FF:000019">
        <property type="entry name" value="Myosin light chain 1 skeletal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.238.10:FF:000056">
        <property type="entry name" value="Myosin light chain 1 skeletal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.238.10">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050230">
        <property type="entry name" value="CALM/Myosin/TropC-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011992">
        <property type="entry name" value="EF-hand-dom_pair"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002048">
        <property type="entry name" value="EF_hand_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23048">
        <property type="entry name" value="MYOSIN LIGHT CHAIN 1, 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23048:SF1">
        <property type="entry name" value="MYOSIN LIGHT CHAIN 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47473">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50222">
        <property type="entry name" value="EF_HAND_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0505">Motor protein</keyword>
    <keyword evidence="4" id="KW-0514">Muscle protein</keyword>
    <keyword evidence="2" id="KW-0518">Myosin</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="EF-hand" evidence="5">
        <location>
            <begin position="118"/>
            <end position="153"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00011445"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023179"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50222"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034378.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="185" mass="20935" checksum="EC8671347C5038C0" modified="2019-05-08" version="1">MLKSHKVLRLLDFFYCNRELPALARIAPQACYQIDFSADQIEEFKEAFMLFDRTPTGEMKITYGQCGDVLRALGQNPTNAEVLKVLGKPKPEEMNTKMLDFETFLPMLQHISRSKDQGTFEDFVEGLRVFDKEGNGTVMGAELRHVLATLGEKLTESEVEQLMTGQEDANGCINYEAFVKHIMSG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1X4</accession>
    <name>A0A452J1X4_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J1X4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038933.1">
        <property type="protein sequence ID" value="ENSGAGP00000034390.1"/>
        <property type="gene ID" value="ENSGAGG00000024438.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019005">
        <property type="term" value="C:SCF ubiquitin ligase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031146">
        <property type="term" value="P:SCF-dependent proteasomal ubiquitin-dependent protein catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd22129">
        <property type="entry name" value="F-box_FBXL22"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036047">
        <property type="entry name" value="F-box-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006553">
        <property type="entry name" value="Leu-rich_rpt_Cys-con_subtyp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13318">
        <property type="entry name" value="PARTNER OF PAIRED, ISOFORM B-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00367">
        <property type="entry name" value="LRR_CC"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81383">
        <property type="entry name" value="F-box domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52047">
        <property type="entry name" value="RNI-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0833">Ubl conjugation pathway</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034390.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="226" mass="26062" checksum="95383C9C95F2CB86" modified="2019-05-08" version="1">MHITQLNRECLLHLFSFLDKNSRKSLAQTCHKLLEVFQEPFLWPVLNFHSPVELKKDNFLLGPALRYLSICWHSSRVKVCNIEDWMKNTFQKDICRKHENAVNHFLIQVCKRCPNLLSLTLSGCGHVTDDYLLLLLKSCPNLKILKLENCVRITDQTLEAVTLYGGSLQTLHVDFCRNITQAGLETVQEKCPSVMLKAERSAHMIPDNEPGEKLMLERAVRKLVLH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1X7</accession>
    <name>A0A452J1X7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Ig-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Lipid-anchor</topology>
            <topology evidence="1">GPI-anchor</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="12">Belongs to the immunoglobulin superfamily. IgLON family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1X7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038958.1">
        <property type="protein sequence ID" value="ENSGAGP00000034409.1"/>
        <property type="gene ID" value="ENSGAGG00000024455.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000038960.1">
        <property type="protein sequence ID" value="ENSGAGP00000034411.1"/>
        <property type="gene ID" value="ENSGAGG00000024455.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098552">
        <property type="term" value="C:side of membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000013">
        <property type="entry name" value="cell adhesion molecule 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000500">
        <property type="entry name" value="limbic system-associated membrane protein isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000113">
        <property type="entry name" value="Opioid-binding protein/cell adhesion molecule"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013098">
        <property type="entry name" value="Ig_I-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050876">
        <property type="entry name" value="IgLON_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR42757">
        <property type="entry name" value="IGLON FAMILY OF IMMUNOGLOBULIN SUPERFAMILY-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR42757:SF22">
        <property type="entry name" value="LIMBIC SYSTEM-ASSOCIATED MEMBRANE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07679">
        <property type="entry name" value="I-set"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13927">
        <property type="entry name" value="Ig_3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="9" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="3" id="KW-0336">GPI-anchor</keyword>
    <keyword evidence="11" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="10" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="7" id="KW-0472">Membrane</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="4 13" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="13">
        <location>
            <begin position="1"/>
            <end position="28"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5044604602" description="Ig-like domain-containing protein" evidence="13">
        <location>
            <begin position="29"/>
            <end position="338"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="14">
        <location>
            <begin position="24"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="14">
        <location>
            <begin position="132"/>
            <end position="214"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="14">
        <location>
            <begin position="219"/>
            <end position="306"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004609"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022622"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00037995"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034409.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="338" mass="37470" checksum="B471ECAAC77883FA" modified="2019-05-08" version="1">MGGRAQPDRKQLPLVLLRLLCLLPTGLPVRSLDFTRGTDNITVRQGDTAILRCYVEDRSSKVAWLNRSGIIFAGQDKWSLDPRVELEKRTPLEYSLRIQKVDVYDEGSYTCSVQTQHHPRTSQVYLIVQVPPKISNISSDVTVNEGSNVTLVCMANGRPEPVITWRHLTPTGREFEGEEEYLEILGITREQSGKYECKAANEVSSADVKQVRVTVNYPPTITESKSNEAATGRQALLKCEASAVPTPDFEWYRDDTRINSANGLEIKSMGSQSLLMVANVTEEHYGNYTCVAANKLGVTNASLYLYRPGTGRVDNGSVSLAVPLWLLAVSLLCLLSKC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="22" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1Y8</accession>
    <name>A0A452J1Y8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">UNC93-like protein MFSD11</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="8">Major facilitator superfamily domain-containing protein 11</fullName>
        </alternativeName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the unc-93 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1Y8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038953.1">
        <property type="protein sequence ID" value="ENSGAGP00000034404.1"/>
        <property type="gene ID" value="ENSGAGG00000024454.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000038955.1">
        <property type="protein sequence ID" value="ENSGAGP00000034406.1"/>
        <property type="gene ID" value="ENSGAGG00000024454.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="CDD" id="cd17407">
        <property type="entry name" value="MFS_MFSD11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1250.20">
        <property type="entry name" value="MFS general substrate transporter like domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010291">
        <property type="entry name" value="Ion_channel_UNC-93"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036259">
        <property type="entry name" value="MFS_trans_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051617">
        <property type="entry name" value="UNC-93-like_regulator"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23294">
        <property type="entry name" value="ET TRANSLATION PRODUCT-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23294:SF0">
        <property type="entry name" value="UNC93-LIKE PROTEIN MFSD11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05978">
        <property type="entry name" value="UNC-93"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103473">
        <property type="entry name" value="MFS general substrate transporter"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="5 9" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 9" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 9" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="7"/>
            <end position="26"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="46"/>
            <end position="65"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="77"/>
            <end position="94"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="100"/>
            <end position="118"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="138"/>
            <end position="158"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="170"/>
            <end position="190"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="239"/>
            <end position="257"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="277"/>
            <end position="298"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="310"/>
            <end position="329"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="387"/>
            <end position="405"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="411"/>
            <end position="432"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009172"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00040302"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00041910"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034406.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="449" mass="49434" checksum="DE00623E1E8AEC09" modified="2019-05-08" version="1">MSPKSKKLFNIIMLGVAFMFMFTAFQTCGNVAQTVITNLNNTNFHGSGYTSMAIIYGVFSASNLISPSVVAVVGPQLSMFVSGLFYSIYIAVFIQPFTWSFYTASVFIGIAAAVIWTAQGNCLTINSDENTIGRNSGIFWALLQSSLFFGNLYIYFAWRGKTHISESDRRTVFIALTVLSLVGTFLFFLIRKPEESNTLEEEVSSTDESIVDNSSVQSKATRAVDAFKRSIKLYFTKEILLLSVTTAYTGLELTFFSGVYGTCIGAVNRFGTEEKSLIGLSGIFIGIGEILGGGFFGLVNKDNRFGRNPVVMLGIVIHFVAFYLIFYNIPNDAPIASFEGTDNSAYMNPSKELAIFCSFLLGLGDSCFNTQLLSILGFLYSEDSAPAFAVFKFVQSICAAVAFFYSNYLLLQWQLLIMAVVGFFGTISFFTVEWEAAAFVARGSDYRSI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J234</accession>
    <name>A0A452J234_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">RNA helicase</fullName>
            <ecNumber evidence="5">3.6.4.13</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="28">DEAH box protein 9</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="29">Nuclear DNA helicase II</fullName>
        </alternativeName>
    </protein>
    <organism evidence="36 37">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="37" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="36" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="36" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="30">
            <text>ATP + H2O = ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:13065"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="3.6.4.13"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Microtubule organizing center</location>
            <location evidence="1">Centrosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
            <location evidence="2">Nucleolus</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Nucleus</location>
            <location evidence="3">Nucleoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the DEAD box helicase family. DEAH subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.6.4.13" evidence="5"/>
    <dbReference type="AlphaFoldDB" id="A0A452J234"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039007.1">
        <property type="protein sequence ID" value="ENSGAGP00000034451.1"/>
        <property type="gene ID" value="ENSGAGG00000024443.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005730">
        <property type="term" value="C:nucleolus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005654">
        <property type="term" value="C:nucleoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:1990904">
        <property type="term" value="C:ribonucleoprotein complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043138">
        <property type="term" value="F:3'-5' DNA helicase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033679">
        <property type="term" value="F:3'-5' DNA/RNA helicase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003725">
        <property type="term" value="F:double-stranded RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003724">
        <property type="term" value="F:RNA helicase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006353">
        <property type="term" value="P:DNA-templated transcription termination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006397">
        <property type="term" value="P:mRNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051028">
        <property type="term" value="P:mRNA transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045944">
        <property type="term" value="P:positive regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050684">
        <property type="term" value="P:regulation of mRNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006417">
        <property type="term" value="P:regulation of translation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031047">
        <property type="term" value="P:regulatory ncRNA-mediated gene silencing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008380">
        <property type="term" value="P:RNA splicing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd17972">
        <property type="entry name" value="DEXHc_DHX9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd19854">
        <property type="entry name" value="DSRM_DHX9_rpt1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd19855">
        <property type="entry name" value="DSRM_DHX9_rpt2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18791">
        <property type="entry name" value="SF2_C_RHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.20:FF:000026">
        <property type="entry name" value="ATP-dependent RNA helicase A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.20:FF:000028">
        <property type="entry name" value="ATP-dependent RNA helicase A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000677">
        <property type="entry name" value="ATP-dependent RNA helicase A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.1080:FF:000006">
        <property type="entry name" value="ATP-dependent RNA helicase A protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000284">
        <property type="entry name" value="probable ATP-dependent RNA helicase YTHDC2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.1080">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.20">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011709">
        <property type="entry name" value="DEAD-box_helicase_OB_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011545">
        <property type="entry name" value="DEAD/DEAH_box_helicase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044447">
        <property type="entry name" value="DHX9_DEXHc"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044445">
        <property type="entry name" value="DHX9_DSRM_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044446">
        <property type="entry name" value="DHX9_DSRM_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002464">
        <property type="entry name" value="DNA/RNA_helicase_DEAH_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014720">
        <property type="entry name" value="dsRBD_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048333">
        <property type="entry name" value="HA2_WH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007502">
        <property type="entry name" value="Helicase-assoc_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014001">
        <property type="entry name" value="Helicase_ATP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001650">
        <property type="entry name" value="Helicase_C-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18934">
        <property type="entry name" value="ATP-DEPENDENT RNA HELICASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18934:SF119">
        <property type="entry name" value="ATP-DEPENDENT RNA HELICASE A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00270">
        <property type="entry name" value="DEAD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00035">
        <property type="entry name" value="dsrm"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21010">
        <property type="entry name" value="HA2_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00271">
        <property type="entry name" value="Helicase_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07717">
        <property type="entry name" value="OB_NTP_bind"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04408">
        <property type="entry name" value="WHD_HA2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00487">
        <property type="entry name" value="DEXDc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00358">
        <property type="entry name" value="DSRM"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00847">
        <property type="entry name" value="HA2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00490">
        <property type="entry name" value="HELICc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54768">
        <property type="entry name" value="dsRNA-binding domain-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00690">
        <property type="entry name" value="DEAH_ATP_HELICASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50137">
        <property type="entry name" value="DS_RBD"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51192">
        <property type="entry name" value="HELICASE_ATP_BIND_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51194">
        <property type="entry name" value="HELICASE_CTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="22" id="KW-0010">Activator</keyword>
    <keyword evidence="16" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="8" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="26" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="20" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="14" id="KW-0347">Helicase</keyword>
    <keyword evidence="13" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="25" id="KW-0464">Manganese</keyword>
    <keyword evidence="10" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9" id="KW-0507">mRNA processing</keyword>
    <keyword evidence="24" id="KW-0508">mRNA splicing</keyword>
    <keyword evidence="15" id="KW-0509">mRNA transport</keyword>
    <keyword evidence="12" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="27" id="KW-0539">Nucleus</keyword>
    <keyword evidence="37" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="11" id="KW-0677">Repeat</keyword>
    <keyword evidence="18 31" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="21" id="KW-0943">RNA-mediated gene silencing</keyword>
    <keyword evidence="23" id="KW-0804">Transcription</keyword>
    <keyword evidence="19" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="7" id="KW-0806">Transcription termination</keyword>
    <keyword evidence="17" id="KW-0810">Translation regulation</keyword>
    <keyword evidence="6" id="KW-0813">Transport</keyword>
    <feature type="domain" description="DRBM" evidence="33">
        <location>
            <begin position="3"/>
            <end position="71"/>
        </location>
    </feature>
    <feature type="domain" description="DRBM" evidence="33">
        <location>
            <begin position="176"/>
            <end position="248"/>
        </location>
    </feature>
    <feature type="domain" description="Helicase ATP-binding" evidence="34">
        <location>
            <begin position="367"/>
            <end position="533"/>
        </location>
    </feature>
    <feature type="domain" description="Helicase C-terminal" evidence="35">
        <location>
            <begin position="605"/>
            <end position="778"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="32">
        <location>
            <begin position="106"/>
            <end position="131"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="32">
        <location>
            <begin position="1119"/>
            <end position="1154"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="32">
        <location>
            <begin position="1210"/>
            <end position="1230"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="32">
        <location>
            <begin position="117"/>
            <end position="131"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004604"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004642"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00008792"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012552"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022472"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022664"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022806"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022816"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022845"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00023158"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00023187"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00023211"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="ARBA" id="ARBA00031576"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="ARBA" id="ARBA00033216"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="ARBA" id="ARBA00047984"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="31">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00266"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="32">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="33">
        <source>
            <dbReference type="PROSITE" id="PS50137"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="34">
        <source>
            <dbReference type="PROSITE" id="PS51192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="35">
        <source>
            <dbReference type="PROSITE" id="PS51194"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="36">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034451.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="37">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1230" mass="136760" checksum="68F4D064A7C8F864" modified="2019-05-08" version="1">MGDVKNFLYAWCGKRKVTPTYEIRAVGNKNRQKFMCEVRVEGFNYTGMGNSTNKKDAQSNAARDFLNYLVRLNEIKSEEIPALGVNGIVPEARGDAGDRVSTLGGPLPPHLALKSETGGGPAPGVGSSGYGGAQWDRGANLKDYYSRKEEQEAQATLESEEVDLNAGLHGNWTLENAKARLNQFFQKEKVQGEYKYTQVGPDHNRSFIAEMTIYVKQLGRRIFAREHGSNKKLAAQSCALSLVRQLYHLGVIEPYTGQTKKKEGETVSLLQKEHYPVSLNIGKLAHFEPSVRQNQMGVVPWSPPQSNWNPWTSCNIDEGPLAYATPEQISMDLKNDLMYRLEQDQELQRILQERETLPVKKFEIEILDAIHHSPVVVIRGATGCGKTTQVPQYILDEYIKNGKAAECNIVVTQPRRISAVSVAERVSYERGEEPGQSCGYSVRFESVLPRPHASVMFCTVGVLLRKLEAGIRGISHVIVDEIHERDINTDFLMVVLRDVVEAYPEIRVILMSATIDTSMFCEYFFNCPIIEVYGRTFPVEEYFLEDCIQMTQFVPPLKDKKKKDKDEEGGEDDDANCNLICSDEYGPETKRCMAQLNEKETPFELIEALLKYIETLNVPGAVLVFLPGWNLIYTMQKHLEMNPHFGSHRYRILPLHSQIPRDEQRRVFDPVPSGVTKVILSTNIAETSITINDVVYVIDSCKQKVKLFTAHNNMTNYATVWASKTNLEQRKGRAGRVRAGFCFHLCSRARFERLETHMTPEMFRTPLHEIALSIKLLRLGGIGQFLAKAIEPPPLDAVIEAEHTLRELDALDSNDELTPLGRILAKLPIEPRLGKMMIMGCIFYVGDAVCTISAATCFPEPFISEGKRLGYVHRNFAGNRFSDHVALLSVFQAWDDARMGGEDAEIRFCEHKRLNMATLRMTWEAKVQLKDILINSGFPEECLMTQIFNNTGPDNNLDVVISLLAFGVYPNVCYHKEKRKILTTEGRNALIHKSSVNCPFSSQDMKYPSPFFVFGEKIRTRAISAKGMTLVSPLQLLLFASKKVLSDGEIILVDDWIKLRMSHDAAACVTALRAAMEALVVEVTKDPEIISQLDPVNERMLNVIRQISRPSAAGINLMAGNTRFGDGPRPPKMARYDNGGGFRGRGVNQRGSGYRGGGYGGGGYNSYRGGSGGRGYQGGSGGGYRGGGGYRGSGGGYRGSGGGYRGYQGGGNRGNYGGGYGGQGRGRGGY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J235</accession>
    <name>A0A452J235_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Malic enzyme</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="10">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
        <cofactor evidence="10">
            <name>Mn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29035"/>
        </cofactor>
        <text evidence="10">Divalent metal cations. Prefers magnesium or manganese.</text>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29035"/>
        </cofactor>
    </comment>
    <comment type="subunit">
        <text evidence="4">Homotetramer.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 11">Belongs to the malic enzymes family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J235"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039009.1">
        <property type="protein sequence ID" value="ENSGAGP00000034453.1"/>
        <property type="gene ID" value="ENSGAGG00000024492.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004473">
        <property type="term" value="F:malate dehydrogenase (decarboxylating) (NADP+) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051287">
        <property type="term" value="F:NAD binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006108">
        <property type="term" value="P:malate metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05312">
        <property type="entry name" value="NAD_bind_1_malic_enz"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.720:FF:000060">
        <property type="entry name" value="Malic enzyme"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10380:FF:000028">
        <property type="entry name" value="NADP-dependent malic enzyme 4 chloroplastic"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.10380">
        <property type="entry name" value="Malic enzyme, N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.720">
        <property type="entry name" value="NAD(P)-binding Rossmann-like Domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046346">
        <property type="entry name" value="Aminoacid_DH-like_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015884">
        <property type="entry name" value="Malic_enzyme_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012301">
        <property type="entry name" value="Malic_N_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037062">
        <property type="entry name" value="Malic_N_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012302">
        <property type="entry name" value="Malic_NAD-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001891">
        <property type="entry name" value="Malic_OxRdtase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036291">
        <property type="entry name" value="NAD(P)-bd_dom_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF010052">
        <property type="entry name" value="PRK13529.1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23406">
        <property type="entry name" value="MALIC ENZYME-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23406:SF17">
        <property type="entry name" value="NADP-DEPENDENT MALIC ENZYME"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00390">
        <property type="entry name" value="malic"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03949">
        <property type="entry name" value="Malic_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000106">
        <property type="entry name" value="ME"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00072">
        <property type="entry name" value="MALOXRDTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM01274">
        <property type="entry name" value="malic"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00919">
        <property type="entry name" value="Malic_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53223">
        <property type="entry name" value="Aminoacid dehydrogenase-like, N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51735">
        <property type="entry name" value="NAD(P)-binding Rossmann-fold domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00331">
        <property type="entry name" value="MALIC_ENZYMES"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6 10 11" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7" id="KW-0521">NADP</keyword>
    <keyword evidence="8 11" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Malic enzyme N-terminal" evidence="13">
        <location>
            <begin position="1"/>
            <end position="161"/>
        </location>
    </feature>
    <feature type="domain" description="Malic enzyme NAD-binding" evidence="12">
        <location>
            <begin position="171"/>
            <end position="423"/>
        </location>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="56"/>
        </location>
        <ligand>
            <name>(S)-malate</name>
            <dbReference type="ChEBI" id="CHEBI:15589"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="10">
        <location>
            <position position="146"/>
        </location>
        <ligand>
            <name>a divalent metal cation</name>
            <dbReference type="ChEBI" id="CHEBI:60240"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="10">
        <location>
            <position position="147"/>
        </location>
        <ligand>
            <name>a divalent metal cation</name>
            <dbReference type="ChEBI" id="CHEBI:60240"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="10">
        <location>
            <position position="170"/>
        </location>
        <ligand>
            <name>a divalent metal cation</name>
            <dbReference type="ChEBI" id="CHEBI:60240"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="309"/>
        </location>
        <ligand>
            <name>(S)-malate</name>
            <dbReference type="ChEBI" id="CHEBI:15589"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="354"/>
        </location>
        <ligand>
            <name>(S)-malate</name>
            <dbReference type="ChEBI" id="CHEBI:15589"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001936"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00008785"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00011881"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022857"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PIRSR" id="PIRSR000106-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PIRSR" id="PIRSR000106-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="RuleBase" id="RU003426"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="SMART" id="SM00919"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="SMART" id="SM01274"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034453.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="472" mass="52249" checksum="1C51F9BE6139E91C" modified="2019-05-08" version="1">AHTELPKHQPCVLPNIYLFACRGLFITIHDWGHIATMLKSWPEDAIRAIVVTDGERILGLGDLGCYGMGIPVGKLALYTACGGMKPQECLPVILDVGTDNEELLKDPLYIGLRHKRVRGQAYNDLLDEFMQAVTFRYGMNCLIQFEDFANANAFRLLNRYRNKYCTFNDDIQGTAAVAVAGLLAALRITENKLSDHTILFQGAGEAALGIANLVIMAMQKEGTPEHEAIKKIWMVDSKGLIVKERSSLTTEKERFAHKHDEMKNLEDIVRAIKPTVLIGVAAIGGAFTKQILKDMASFNKQPIIFALSNPTSKAECSAEQCYQMTEGRGIFASGSPFDPVTLPGGRTFYPGQGNNAYVFPGVALGVVSCGLRYISEDIFLTTAEVIAQQVSKKHLEEGRLYPPLVTIRDVSLKIAVKIAEQAYSNKTASFYPEPEDKEAFIRSQVYSTDYDNFVVDSYVWPEEAMKIQTGNP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J239</accession>
    <name>A0A452J239_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Malic enzyme</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="10">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
        <cofactor evidence="10">
            <name>Mn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29035"/>
        </cofactor>
        <text evidence="10">Divalent metal cations. Prefers magnesium or manganese.</text>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29035"/>
        </cofactor>
    </comment>
    <comment type="subunit">
        <text evidence="4">Homotetramer.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 11">Belongs to the malic enzymes family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J239"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039013.1">
        <property type="protein sequence ID" value="ENSGAGP00000034457.1"/>
        <property type="gene ID" value="ENSGAGG00000024492.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004473">
        <property type="term" value="F:malate dehydrogenase (decarboxylating) (NADP+) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051287">
        <property type="term" value="F:NAD binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006108">
        <property type="term" value="P:malate metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05312">
        <property type="entry name" value="NAD_bind_1_malic_enz"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.720:FF:000060">
        <property type="entry name" value="Malic enzyme"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10380:FF:000028">
        <property type="entry name" value="NADP-dependent malic enzyme 4 chloroplastic"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.10380">
        <property type="entry name" value="Malic enzyme, N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.720">
        <property type="entry name" value="NAD(P)-binding Rossmann-like Domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046346">
        <property type="entry name" value="Aminoacid_DH-like_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015884">
        <property type="entry name" value="Malic_enzyme_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012301">
        <property type="entry name" value="Malic_N_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037062">
        <property type="entry name" value="Malic_N_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012302">
        <property type="entry name" value="Malic_NAD-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001891">
        <property type="entry name" value="Malic_OxRdtase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036291">
        <property type="entry name" value="NAD(P)-bd_dom_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF010052">
        <property type="entry name" value="PRK13529.1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23406">
        <property type="entry name" value="MALIC ENZYME-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23406:SF17">
        <property type="entry name" value="NADP-DEPENDENT MALIC ENZYME"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00390">
        <property type="entry name" value="malic"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03949">
        <property type="entry name" value="Malic_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000106">
        <property type="entry name" value="ME"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00072">
        <property type="entry name" value="MALOXRDTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM01274">
        <property type="entry name" value="malic"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00919">
        <property type="entry name" value="Malic_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53223">
        <property type="entry name" value="Aminoacid dehydrogenase-like, N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51735">
        <property type="entry name" value="NAD(P)-binding Rossmann-fold domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00331">
        <property type="entry name" value="MALIC_ENZYMES"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6 10 11" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7" id="KW-0521">NADP</keyword>
    <keyword evidence="8 11" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Malic enzyme N-terminal" evidence="13">
        <location>
            <begin position="1"/>
            <end position="149"/>
        </location>
    </feature>
    <feature type="domain" description="Malic enzyme NAD-binding" evidence="12">
        <location>
            <begin position="159"/>
            <end position="411"/>
        </location>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="44"/>
        </location>
        <ligand>
            <name>(S)-malate</name>
            <dbReference type="ChEBI" id="CHEBI:15589"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="10">
        <location>
            <position position="134"/>
        </location>
        <ligand>
            <name>a divalent metal cation</name>
            <dbReference type="ChEBI" id="CHEBI:60240"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="10">
        <location>
            <position position="135"/>
        </location>
        <ligand>
            <name>a divalent metal cation</name>
            <dbReference type="ChEBI" id="CHEBI:60240"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="10">
        <location>
            <position position="158"/>
        </location>
        <ligand>
            <name>a divalent metal cation</name>
            <dbReference type="ChEBI" id="CHEBI:60240"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="297"/>
        </location>
        <ligand>
            <name>(S)-malate</name>
            <dbReference type="ChEBI" id="CHEBI:15589"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="342"/>
        </location>
        <ligand>
            <name>(S)-malate</name>
            <dbReference type="ChEBI" id="CHEBI:15589"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001936"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00008785"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00011881"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022857"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PIRSR" id="PIRSR000106-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PIRSR" id="PIRSR000106-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="RuleBase" id="RU003426"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="SMART" id="SM00919"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="SMART" id="SM01274"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034457.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="460" mass="50930" checksum="827A127BC5DFF6F8" modified="2019-05-08" version="1">MQSIYLFACRGLFITIHDWGHIATMLKSWPEDAIRAIVVTDGERILGLGDLGCYGMGIPVGKLALYTACGGMKPQECLPVILDVGTDNEELLKDPLYIGLRHKRVRGQAYNDLLDEFMQAVTFRYGMNCLIQFEDFANANAFRLLNRYRNKYCTFNDDIQGTAAVAVAGLLAALRITENKLSDHTILFQGAGEAALGIANLVIMAMQKEGTPEHEAIKKIWMVDSKGLIVKERSSLTTEKERFAHKHDEMKNLEDIVRAIKPTVLIGVAAIGGAFTKQILKDMASFNKQPIIFALSNPTSKAECSAEQCYQMTEGRGIFASGSPFDPVTLPGGRTFYPGQGNNAYVFPGVALGVVSCGLRYISEDIFLTTAEVIAQQVSKKHLEEGRLYPPLVTIRDVSLKIAVKIAEQAYSNKTASFYPEPEDKEAFIRSQVYSTDYDNFVVDSYVWPEEAMKIQTGNP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J246</accession>
    <name>A0A452J246_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="7">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the cornichon family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J246"/>
    <dbReference type="SMR" id="A0A452J246"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038908.1">
        <property type="protein sequence ID" value="ENSGAGP00000034365.1"/>
        <property type="gene ID" value="ENSGAGG00000024418.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016192">
        <property type="term" value="P:vesicle-mediated transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003377">
        <property type="entry name" value="Cornichon"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033466">
        <property type="entry name" value="Cornichon_conserved"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12290">
        <property type="entry name" value="CORNICHON-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03311">
        <property type="entry name" value="Cornichon"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01398">
        <property type="entry name" value="Cornichon"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01340">
        <property type="entry name" value="CORNICHON"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 6" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 6" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 6" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="7"/>
            <end position="29"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="56"/>
            <end position="82"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="123"/>
            <end position="142"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010095"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034365.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="144" mass="16715" checksum="F85D114D24C455DB" modified="2019-05-08" version="1">MAFTFAAFCYMLALLLTAALIFFAIWHIIAFDELKTDYKNPIDQCNTLNPLVLPEYLIHAFFCVMFLCAAEWLTLGLNMPLLAYHIWRYMSRPVMSGPGLYDPTTIMNADILAYCQKEGWCKLAFYLLSFFYYLYGMIYVLVSS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J247</accession>
    <name>A0A452J247_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="6">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J247"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039027.1">
        <property type="protein sequence ID" value="ENSGAGP00000034471.1"/>
        <property type="gene ID" value="ENSGAGG00000024490.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005903">
        <property type="term" value="C:brush border"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055056">
        <property type="term" value="F:D-glucose transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046323">
        <property type="term" value="P:D-glucose import"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070837">
        <property type="term" value="P:dehydroascorbic acid transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1250.20">
        <property type="entry name" value="MFS general substrate transporter like domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045263">
        <property type="entry name" value="GLUT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005828">
        <property type="entry name" value="MFS_sugar_transport-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036259">
        <property type="entry name" value="MFS_trans_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23503">
        <property type="entry name" value="SOLUTE CARRIER FAMILY 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23503:SF27">
        <property type="entry name" value="SOLUTE CARRIER FAMILY 2, FACILITATED GLUCOSE TRANSPORTER MEMBER 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00083">
        <property type="entry name" value="Sugar_tr"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4 5" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 5" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="3 5" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="12"/>
            <end position="30"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="86"/>
            <end position="109"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034471.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="130" mass="14248" checksum="708F19A490084E60" modified="2019-05-08" version="1">FTEEILPPHLTGTLVLSVFTAVLGFFQYGYSLGVINAPQRVIEAHYGRVLGIAPIDRSAIDFTDTNGSMIFPDLEELDVGKSTLTLYWSLSVSIFAIGGMISSFTVGWVGDKEVLFASNFLPSSYLKVQY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J248</accession>
    <name>A0A452J248_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Oxysterol-binding protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="5">Belongs to the OSBP family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J248"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039051.1">
        <property type="protein sequence ID" value="ENSGAGP00000034490.1"/>
        <property type="gene ID" value="ENSGAGG00000024516.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000039053.1">
        <property type="protein sequence ID" value="ENSGAGP00000034492.1"/>
        <property type="gene ID" value="ENSGAGG00000024516.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015485">
        <property type="term" value="F:cholesterol binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006869">
        <property type="term" value="P:lipid transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.2720:FF:000001">
        <property type="entry name" value="Oxysterol-binding OBPalpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.160.120:FF:000002">
        <property type="entry name" value="Oxysterol-binding protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.3490:FF:000001">
        <property type="entry name" value="Oxysterol-binding protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.2720">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.160.120">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.3490">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037239">
        <property type="entry name" value="OSBP_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000648">
        <property type="entry name" value="Oxysterol-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018494">
        <property type="entry name" value="Oxysterol-bd_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10972">
        <property type="entry name" value="OXYSTEROL-BINDING PROTEIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10972:SF47">
        <property type="entry name" value="OXYSTEROL-BINDING PROTEIN-RELATED PROTEIN 10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01237">
        <property type="entry name" value="Oxysterol_BP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF144000">
        <property type="entry name" value="Oxysterol-binding protein-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01013">
        <property type="entry name" value="OSBP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 6" id="KW-0445">Lipid transport</keyword>
    <keyword evidence="4" id="KW-0446">Lipid-binding</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1 6" id="KW-0813">Transport</keyword>
    <feature type="domain" description="PH" evidence="8">
        <location>
            <begin position="1"/>
            <end position="34"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="36"/>
            <end position="78"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="45"/>
            <end position="67"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="68"/>
            <end position="78"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023121"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="RuleBase" id="RU003844"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="RuleBase" id="RU003845"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034490.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="626" mass="70210" checksum="A7221FE1C57FBFC5" modified="2019-05-08" version="1">MLVIYFANGEMYKLRAADAKEKQYWVTQLRSCAKYHTEGNSKNTPSSRNRNLSLLPSGMSNSASPSSQKHQGQSGPTVVTVTHHKLPAAARRAKSQYTGQLHDVREIMSHVEGQQKNLVQAIESLPGSGPFSALDQDLLLLKATSAATLSCLGECLNLLQQSMHQAGQHNNRTLPSENILGWPGPKSHSTELKNGAFSSLSSASANITWAFVPAATQDGQTLQPNTEHSAPELILVEDEMTDTEDNEEEDLGVMEDQRSIILHLISQLKLGMDLTRVVLPTFILEKRSLLEMYANFMAHPDLFLSIAAGASPEERIICFVEYYLTAFHEGRKGAVAKKPYNPIIGESFHCSWDVPKDKVKSFRTNSASIVSTDQTKEFGQDQSECYKLRFVAEQVSHHPPVSCFYCECKEKRMCVNAHVWTKSKFLGMSIGVSMVGEGVLYLMEHEEEYVFTLPSAYARSILTIPWVELGGKISINCAKTGYSATVTFHTKPFYGGKVHRVTAEVKHNPTNTIVCKAQGEWNGILEFTYSNGETKVIDTNKLPVIRKKIRPIAKQGPFESRHLWQHVTNSLKEGNIDAATEHKHRLEERQRAEERQRVAINTPWKPKYFTKEGDGWIYFNPLWKSH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J270</accession>
    <name>A0A452J270_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="12">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Golgi apparatus membrane</location>
            <topology evidence="1">Peripheral membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the DOP1 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000039061.1">
        <property type="protein sequence ID" value="ENSGAGP00000034498.1"/>
        <property type="gene ID" value="ENSGAGG00000024506.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="GOC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005768">
        <property type="term" value="C:endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000139">
        <property type="term" value="C:Golgi membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005802">
        <property type="term" value="C:trans-Golgi network"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006895">
        <property type="term" value="P:Golgi to endosome transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040314">
        <property type="entry name" value="DOP1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056457">
        <property type="entry name" value="DOP1_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007249">
        <property type="entry name" value="DOP1_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056459">
        <property type="entry name" value="TPR_DOP1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056458">
        <property type="entry name" value="TPR_DOP1_M"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14042">
        <property type="entry name" value="DOPEY-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14042:SF22">
        <property type="entry name" value="PROTEIN DOPEY-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24598">
        <property type="entry name" value="DOP1_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04118">
        <property type="entry name" value="Dopey_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24601">
        <property type="entry name" value="TPR_DOP1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24597">
        <property type="entry name" value="TPR_DOP1_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="5" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0653">Protein transport</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0813">Transport</keyword>
    <feature type="domain" description="DOP1 N-terminal" evidence="8">
        <location>
            <begin position="11"/>
            <end position="294"/>
        </location>
    </feature>
    <feature type="domain" description="DOP1-like middle TPR" evidence="9">
        <location>
            <begin position="318"/>
            <end position="534"/>
        </location>
    </feature>
    <feature type="domain" description="DOP1-like TPR" evidence="11">
        <location>
            <begin position="1351"/>
            <end position="1725"/>
        </location>
    </feature>
    <feature type="domain" description="DOP1-like C-terminal" evidence="10">
        <location>
            <begin position="1892"/>
            <end position="2412"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="568"/>
            <end position="589"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="613"/>
            <end position="638"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="705"/>
            <end position="725"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1163"/>
            <end position="1207"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1276"/>
            <end position="1301"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="623"/>
            <end position="638"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="710"/>
            <end position="721"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="1163"/>
            <end position="1192"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="1193"/>
            <end position="1207"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004395"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00046326"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF04118"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF24597"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="Pfam" id="PF24598"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF24601"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034498.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2483" mass="279760" checksum="4FD1824E32C2E18F" modified="2019-05-08" version="1">MNTEEVELLSDSKYRNYVAAVDKALKNFEYSSEWADLISALGKLNKVLQNNAKYQVVPKKLTIGKRLAQCLHPALPGGVHRKALETYEIIFKIIGPKRLAKDLFLYSSGLFPLLANAAMSVKPALLSLYEIYYLPLGKTLKPGLQGLLTGILPGLEEGSEYYERTNTLLEKVAGVDQSAFYSALWGSLLTSPAVRLPGITYVLSHLNRKLSMEDQLYIIGSDIELMVEAVSTSVQDSSVLVQRSTLDLILFCFPFHMSQATRPDMIRILSAALHVVLRRDMSLNRRLYAWLLGFDNNGAIVGPRSTRQNNPEEHATYYFNTFSKEMLVQAMVGILQVNGHGEESTMMQDLKPFRILISLLDKPELGPVILEDVLIEVFRTLYTQCKAELDLQMEPSFSKDHTQLSSKLRENKKTAELIKTANLLFNSFEPYYMWDYIARWFEECCRRTLHARLQTGPVGGNESSELPLTNFCLLVDFLLDIVSLETYIEIQTEHLPQLLLRMISALTSHLHTLHLSELTDSLRLCSKILSKVQPPLLSAGTDGTLQLPSGHNSSIKEWEDKKVPSFSIENPHDVFEDGENPPSSRSSESGFTEFVQYQADTADDIDRALSEGHGAPGIPIIGSTSSETETASTVGSEETIVHPPSIITQGSATQSGKMIQKTAMQCCLEYVQQFLTRFINLYIIQSNSLSQPLATELQVDLTREQGQPAKWDRESQGDAKGKKTNKTPKEYLSAFIAACQLYLECSSFPVYIAEGNHTSELHPEKSDFDCEQVQPPLWLQTLMNACRQASDFSIQGIAISLIMDLVGLTQSVAVVTGENLNSVETAQPLSPNQGRVAVVIRPPLTQGNLRYMVEKTDFFKHVALTLWDQLGDGTPQHHQKSVELFYQLHNLVPSSSICEDVISQQLTHRDKKTRMETHAKFAVLWHLTRDLHINKSSSFGRSFDRSLFIMLDSLNSLDGSTWSVGQAWLNQVLQRHDIARVLEPLLLLLLHPKTQRVSVQRIQAECYWNKSPHYPEEETEKHFMQKFSCGDAFSHMPVSQGQLITPKENNEKQLIMDEMENFSLTVNPLSDRLSLLSTSSETIPMVVSDFELPDHQIEILQSSDSGCSQSSAGDNLSYEVEAESLSAQNSSQTLREDSPDEIVQQVVIDLICKVVSGLGEETESVKHDLHPDDASSKFSTLDHSEEVSKSEDQNIQSSENSLFGNDSSQLLSASTEIGLERLTDSPPCIEVSSQAPCDLTFSSTDGKSGQRSHSSIQFSFKGKLPEKVSEKETIVKEAGKQPGAKPKVKIARRKDEDKKKAQTEKLKQTSIFFSDGLDLENWYSCGEGEISEIESDVGSPGIRKSPSFNVHPLYQHVLLYLQLYDSSRTLYAFSAIKAILKTNPSAFVNAISTTSVNNVYTPQLSLLQNLLARHRISVMGKDFYSHIPVDSNHNFRSSMYIEIIISLCLYYMRSHYPTHVKVTPQDLIGNRNMQMMSIEILTLLFSELAKVIESSAKGFPSFISDMLSKCKVQKVILHCLLSSIFSAQKWHSEKVAGKNIVVIEEGFSEDSLINFSEDEFDSGSTLQSQLLKVLQRLIVLEHRVMTVPEENETGFEFAITDLEQINPQQPMTSLQYLHSQPITSQGMFLCAVIRALHQHCACKMHPQWIGLITSTLPYMGKVLQRVVVSVTLQLCRNLDNLIQQYKYETGLSNSRPLWMASITPPDMILTLLEGITTIIHYCLLDPSTQYHQLLVNVDQKHLLEARSGILSILHMIMSSVTLLWSILHQADSSEKTTAAAAASVTTINLGSTKNLRQQILELLGPISMNHGVHFMAAVAFVWNERRQNKNTSRTKVIPTASDEQLLLVELVRSISVMRTETVIQTVKEVLKQPPAIAKDKKHLSLEVCMLQFFYAYIQRIPVSSLVDSWATLLLLLKDSIQLSLPAPGQFLILGVLNEFIMKNPSLENKKDQRDLQDVTHKIVDAIGAIAGSSLEQTTWLRRNLEVKPSLKIMVDGNNLESDVEDMLYPALETSNITPSVYSVHALTLLSEVLAHLLDMVFYSDEKERVIPLLVNIMHYVVPYLRNHSAHNAPSYRACVQLLSSLSGYQYTRKAWKKEAFDLFMDSSFFQMDASCVNHWRAIMDNLMTHDKTTFRDLMTRVAVAQSSSLNLFANRDVELEQRAMLLKRLAFAIFSSEVDQYQKYLPDIQERLVESLRLPQVPTLHSQVFLFFRVLLLRMSPQHLTSLWPTMITELVQVFLLMEQELTADEDISRTSGPSVAGLETTYTGGNGFSTSYNSQRWLNLYLSACKFLDLALALPSENLPQFQMYRWAFIPEVSDDSGLEVRRQGTHQREFKPYVVRLAKLLRKRAKDKQEDFKMMILEGVEMAKHQKNPEEDCSGKTLSWEPGHLLLTIYTVRSIEQLLPFFNVLSQVFNSKVTSRCVGHSGSPVLYPNSFPSKDIKMENQKTFSSKARQKIEEMVEKDFLEGVIKTHLQPRGTRKA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J292</accession>
    <name>A0A452J292_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">T-box domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 6">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="6">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J292"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038961.1">
        <property type="protein sequence ID" value="ENSGAGP00000034412.1"/>
        <property type="gene ID" value="ENSGAGG00000024450.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000785">
        <property type="term" value="C:chromatin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002302">
        <property type="term" value="P:CD8-positive, alpha-beta T cell differentiation involved in immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001714">
        <property type="term" value="P:endodermal cell fate specification"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001707">
        <property type="term" value="P:mesoderm formation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045893">
        <property type="term" value="P:positive regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd20205">
        <property type="entry name" value="T-box_TBR2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.820:FF:000004">
        <property type="entry name" value="T-box, brain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.820">
        <property type="entry name" value="Transcription factor, T-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008967">
        <property type="entry name" value="p53-like_TF_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032385">
        <property type="entry name" value="T-box_assoc"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046360">
        <property type="entry name" value="T-box_DNA-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036960">
        <property type="entry name" value="T-box_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001699">
        <property type="entry name" value="TF_T-box"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018186">
        <property type="entry name" value="TF_T-box_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11267:SF13">
        <property type="entry name" value="EOMESODERMIN HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11267">
        <property type="entry name" value="T-BOX PROTEIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00907">
        <property type="entry name" value="T-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16176">
        <property type="entry name" value="T-box_assoc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00937">
        <property type="entry name" value="TBOX"/>
    </dbReference>
    <dbReference type="SMART" id="SM00425">
        <property type="entry name" value="TBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49417">
        <property type="entry name" value="p53-like transcription factors"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01283">
        <property type="entry name" value="TBOX_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01264">
        <property type="entry name" value="TBOX_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50252">
        <property type="entry name" value="TBOX_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3 6" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="5 6" id="KW-0539">Nucleus</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0804">Transcription</keyword>
    <keyword evidence="2" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="T-box" evidence="8">
        <location>
            <begin position="192"/>
            <end position="377"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1"/>
            <end position="85"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="528"/>
            <end position="548"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="553"/>
            <end position="572"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="579"/>
            <end position="601"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="22"/>
            <end position="39"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="586"/>
            <end position="597"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00201"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50252"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034412.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="625" mass="68282" checksum="5BAD3018114066E7" modified="2019-05-08" version="1">MQLGEQLLGSAASLAGPPFYPPESSGGQRGQPSSGSPQRLDLDKGPKKCGGAGPGMLSEAEAGEPFPGAGSKQGAPDGRKAAPCGEAELPPAAARYPLDSLSPERYYLQSPGPQGAELGAPCSLFPYPAAQHGSVYQAPGGARYPYGSMLAPGGFSAAVCSPGGRAQFGGGGGGYQYAVPGSGPGLRAQVFLCNRPLWLKFHRHQTEMIITKQGRRMFPFLSFNITGLNPTAHYNVFVEVVLADPNHWRFQGGKWVTCGKADNNMQGNKVYVHPESPNTGSHWMRQEISFGKLKLTNNKGANNNNTQMIVLQSLHKYQPRLHIVEVTEDGVEDLNDSSKTQTFIFPETQFIAVTAYQNTDITQLKIDHNPFAKGFRDNYDSMYTASENDRLTPSPTDSPRSHQIVPGARYSVQPFFQEQFVNNLPPARFYNGERTVPQTNGLLSPQQNEEVANPPQRWFVTPVQQPGTNKLDMNSYETEYSPSTLLPYGIKSLPLQTSHALGYYPDPTFSSMAGWGSRGSYQRKMATGLPWTSRTSPPAFSEDHLPKEKVKEEISSSWIETPPSIKSLDSNDSGIYTSACKRRRLSPSTSSNENSPTMKCEDMNAEDYSKDTSKGMGYYAFYTTT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J299</accession>
    <name>A0A452J299_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">BHLH domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J299"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039103.1">
        <property type="protein sequence ID" value="ENSGAGP00000034534.1"/>
        <property type="gene ID" value="ENSGAGG00000024549.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046983">
        <property type="term" value="F:protein dimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd18930">
        <property type="entry name" value="bHLHzip_MXI1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.280.10:FF:000014">
        <property type="entry name" value="Max dimerization protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.280.10">
        <property type="entry name" value="Helix-loop-helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011598">
        <property type="entry name" value="bHLH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036638">
        <property type="entry name" value="HLH_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11969">
        <property type="entry name" value="MAX DIMERIZATION, MAD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11969:SF13">
        <property type="entry name" value="MAX-INTERACTING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00010">
        <property type="entry name" value="HLH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00353">
        <property type="entry name" value="HLH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47459">
        <property type="entry name" value="HLH, helix-loop-helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50888">
        <property type="entry name" value="BHLH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="6" id="KW-0539">Nucleus</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0678">Repressor</keyword>
    <keyword evidence="5" id="KW-0804">Transcription</keyword>
    <keyword evidence="3" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="BHLH" evidence="9">
        <location>
            <begin position="67"/>
            <end position="119"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="35"/>
            <end position="76"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="159"/>
            <end position="228"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="109"/>
            <end position="150"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="8">
        <location>
            <begin position="48"/>
            <end position="58"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="59"/>
            <end position="70"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="8">
        <location>
            <begin position="173"/>
            <end position="183"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="208"/>
            <end position="228"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022491"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50888"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034534.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="228" mass="26004" checksum="C214F2017ED920CF" modified="2019-05-08" version="1">MERVQMINIQRLLEAAEYLERRERECEHGYASTFPLLPSPGLQDPKPMRRLSRVRKHSSGGSSTSTANRSTHNELEKNRRAHLRLCLERLKVLIPLGPDCTRHTTLGLLNKAKAHIKKLEEAERRSQHQLENLEREQRFLKRRLEQLQGPQEIERIRMDSIGSTISSDRSDSEREEIEVDVESTEFSHGEVDDISTTSISDIDDHSSLQSVGSDEGYSSASVKLSFAS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJH0</accession>
    <name>A0A452GJH0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">RRM domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJH0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001975.1">
        <property type="protein sequence ID" value="ENSGAGP00000001739.1"/>
        <property type="gene ID" value="ENSGAGG00000001387.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003729">
        <property type="term" value="F:mRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.330:FF:000058">
        <property type="entry name" value="RNA-binding motif protein 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.330:FF:000085">
        <property type="entry name" value="RNA-binding protein 4 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.330">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050502">
        <property type="entry name" value="Euk_RNA-bind_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012677">
        <property type="entry name" value="Nucleotide-bd_a/b_plait_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035979">
        <property type="entry name" value="RBD_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000504">
        <property type="entry name" value="RRM_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48025:SF26">
        <property type="entry name" value="HETEROGENEOUS NUCLEAR RIBONUCLEOPROTEIN M-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48025">
        <property type="entry name" value="OS02G0815200 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00076">
        <property type="entry name" value="RRM_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00360">
        <property type="entry name" value="RRM"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54928">
        <property type="entry name" value="RNA-binding domain, RBD"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50102">
        <property type="entry name" value="RRM"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="6 8" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <keyword evidence="4" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="RRM" evidence="9">
        <location>
            <begin position="2"/>
            <end position="72"/>
        </location>
    </feature>
    <feature type="domain" description="RRM" evidence="9">
        <location>
            <begin position="78"/>
            <end position="138"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00176"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50102"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001739.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="188" mass="21396" checksum="4F9197E310049666" modified="2019-05-08" version="1">MVKLFIGNLPREATEQEIRSLFEQYGKVLECDIIKNYGFVHIEDKTAAEDAIRNLHHHKLHGVCINVEASKNKSKASTKLHVGNISTTCTNLELRAKFEEYGPVIECDIVKDYAFVHMERAEDAVEAIRGLDNTEFQGSIFSGGPSGSHYTRKFVLLPPPSHCLRRHCGWRIPSLLFIYLLLLSWELI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJH1</accession>
    <name>A0A452GJH1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">ACHA9 protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="14">Synaptic cell membrane</location>
            <topology evidence="14">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="15">Belongs to the ligand-gated ion channel (TC 1.A.9) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJH1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002082.1">
        <property type="protein sequence ID" value="ENSGAGP00000001829.1"/>
        <property type="gene ID" value="ENSGAGG00000001456.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0022848">
        <property type="term" value="F:acetylcholine-gated monoatomic cation-selective channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004888">
        <property type="term" value="F:transmembrane signaling receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd19051">
        <property type="entry name" value="LGIC_TM_cation"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.390:FF:000009">
        <property type="entry name" value="Cholinergic receptor nicotinic alpha 9 subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.390:FF:000053">
        <property type="entry name" value="Neuronal acetylcholine receptor subunit alpha-9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.70.170.10:FF:000010">
        <property type="entry name" value="neuronal acetylcholine receptor subunit alpha-9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.70.170.10">
        <property type="entry name" value="Neurotransmitter-gated ion-channel ligand-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.390">
        <property type="entry name" value="Neurotransmitter-gated ion-channel transmembrane domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006202">
        <property type="entry name" value="Neur_chan_lig-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036734">
        <property type="entry name" value="Neur_chan_lig-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006201">
        <property type="entry name" value="Neur_channel"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036719">
        <property type="entry name" value="Neuro-gated_channel_TM_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038050">
        <property type="entry name" value="Neuro_actylchol_rec"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006029">
        <property type="entry name" value="Neurotrans-gated_channel_TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018000">
        <property type="entry name" value="Neurotransmitter_ion_chnl_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002394">
        <property type="entry name" value="Nicotinic_acetylcholine_rcpt"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00860">
        <property type="entry name" value="LIC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18945">
        <property type="entry name" value="NEUROTRANSMITTER GATED ION CHANNEL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02931">
        <property type="entry name" value="Neur_chan_LBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02932">
        <property type="entry name" value="Neur_chan_memb"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00254">
        <property type="entry name" value="NICOTINICR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00252">
        <property type="entry name" value="NRIONCHANNEL"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90112">
        <property type="entry name" value="Neurotransmitter-gated ion-channel transmembrane pore"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF63712">
        <property type="entry name" value="Nicotinic receptor ligand binding domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00236">
        <property type="entry name" value="NEUROTR_ION_CHANNEL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="9" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13 15" id="KW-0407">Ion channel</keyword>
    <keyword evidence="7 15" id="KW-0406">Ion transport</keyword>
    <keyword evidence="12" id="KW-1071">Ligand-gated ion channel</keyword>
    <keyword evidence="8 15" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0675">Receptor</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="6" id="KW-0770">Synapse</keyword>
    <keyword evidence="3 15" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 15" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="1 15" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="225"/>
            <end position="249"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="255"/>
            <end position="276"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="288"/>
            <end position="310"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="419"/>
            <end position="437"/>
        </location>
    </feature>
    <feature type="domain" description="Neurotransmitter-gated ion-channel ligand-binding" evidence="16">
        <location>
            <begin position="18"/>
            <end position="222"/>
        </location>
    </feature>
    <feature type="domain" description="Neurotransmitter-gated ion-channel transmembrane" evidence="17">
        <location>
            <begin position="230"/>
            <end position="435"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00034099"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="RuleBase" id="RU000687"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF02931"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF02932"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001829.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="20">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="440" mass="50322" checksum="B19669976667154B" modified="2019-05-08" version="1">FASQFPSSFAVSQGKFAYKLLHDLFANYSNALRPVEDTDRALNVTLQITLSQIIDMDERNQILTAYLWIRQVWLDAYLAWDKDDYDGIDTIRIPSSYVWRPDIVLYNNADDQFTESMETNVVIRYDGQVMWDSPAITKSSCKVDVSYFPFDGQQCRLTFGSWTYNGNQIDLLNGLDTGDLTDFVENVEWEVLGMPAKRNVIIYGCCSEPYPDITYTLLLKRRASFYIFNLLLPCIMISFLAPLGFYLPADSGEKVSLGVTVLLALTVFQLLVAESMPPSENVPLIGKYYIATMTMITASTALTIFIMNIHHCGPGAKPVPKWAKKFILQYMARLFFVYEVGESCKSPKRQLGHLPTVQVVSECGWEEQQHQVCVKTPCLCHHNSLLRNVEYIANCFRDQKAAQKQSGEWKKVAKVMDRIFMWIFFIMVFFMSMLIMGKAI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJH2</accession>
    <name>A0A452GJH2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Rho GTPase activating protein 33</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the PX domain-containing GAP family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000002068.1">
        <property type="protein sequence ID" value="ENSGAGP00000001817.1"/>
        <property type="gene ID" value="ENSGAGG00000001385.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015629">
        <property type="term" value="C:actin cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005938">
        <property type="term" value="C:cell cortex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001650">
        <property type="term" value="C:fibrillar center"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005654">
        <property type="term" value="C:nucleoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035091">
        <property type="term" value="F:phosphatidylinositol binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007264">
        <property type="term" value="P:small GTPase-mediated signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd04384">
        <property type="entry name" value="RhoGAP_CdGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd11835">
        <property type="entry name" value="SH3_ARHGAP32_33"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.555.10:FF:000002">
        <property type="entry name" value="rho GTPase-activating protein 32 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.40:FF:000030">
        <property type="entry name" value="rho GTPase-activating protein 32 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.555.10">
        <property type="entry name" value="Rho GTPase activation protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051576">
        <property type="entry name" value="PX-Rho_GAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036871">
        <property type="entry name" value="PX_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000198">
        <property type="entry name" value="RhoGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15729">
        <property type="entry name" value="CDC42 GTPASE-ACTIVATING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15729:SF11">
        <property type="entry name" value="RHO GTPASE-ACTIVATING PROTEIN 33"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00620">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14604">
        <property type="entry name" value="SH3_9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00324">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00326">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF64268">
        <property type="entry name" value="PX domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50238">
        <property type="entry name" value="RHOGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 4" id="KW-0728">SH3 domain</keyword>
    <feature type="domain" description="SH3" evidence="6">
        <location>
            <begin position="172"/>
            <end position="234"/>
        </location>
    </feature>
    <feature type="domain" description="Rho-GAP" evidence="7">
        <location>
            <begin position="300"/>
            <end position="495"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="506"/>
            <end position="566"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="616"/>
            <end position="666"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="5">
        <location>
            <begin position="617"/>
            <end position="638"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00008795"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50238"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001817.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="10">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="666" mass="73887" checksum="D5AE2FD1822611C0" modified="2019-05-08" version="1">SGAVWGTLGFSGMSFPPRLSITRTHFPKLAECAHFHYEAVDFGHIQLLLASEQGEGLPSTLRDRELLFIIQVTCKGKTWPVHRSYEEFCSLDGHLHCCIFDRRFSQLPELPPPCPDSAPAPQMLAPVLRQYLEKLSAIVDSNINCGPVLTWMEIDNHGNRLLVNEEASINVPAIAAAHVIKRYAAQAPDELSFEVGDIVSVIDMPPKEEVSWWRGKHGFQVGFFPSECVELFPDRLAPELKAGDAWAPQLGCFSPTMSLCPAVCKKHGKLLGFLRTFMKSRPSKQRLKQRGILRERVFGCDLGEHLLNSGHDVPQVLRCCSEFIEKHGMVDGIYRLSGVASNIQKLRHEFDSERVPELGRATYLQDIHCVSSLCKLYFRELPNPLLTYQLYNKFAEAVAVPGEEERLVRVHDVIQQLPPPHYRTLEFLLRHLARVATHSPETSMHIKNLAIVWAPNLLRSLELEAVGLSGAEAFHEVRVQSVVVEFLLSHVQVLFSDKFTSIGKDSAGGHAGGHEGDEAVGARPRGSRRRSKMGWGVCRRNRETEGGGHPPPPGSDSWTQAPLYPXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXYPAPSLPPMGPPTPRYPDPSLPPTGPSTWAVAYHGSVTPPPALVQSAAPPDKH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJH3</accession>
    <name>A0A452GJH3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Ephexin-1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="16">Eph-interacting exchange protein</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="17">Neuronal guanine nucleotide exchange factor</fullName>
        </alternativeName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">Acts as a guanine nucleotide exchange factor (GEF) which differentially activates the GTPases RHOA, RAC1 and CDC42. Plays a role in axon guidance regulating ephrin-induced growth cone collapse and dendritic spine morphogenesis. Upon activation by ephrin through EPHA4, the GEF activity switches toward RHOA resulting in its activation. Activated RHOA promotes cone retraction at the expense of RAC1- and CDC42-stimulated growth cone extension.</text>
    </comment>
    <comment type="subunit">
        <text evidence="14">Interacts with CDK5R1 and EPHA4; activated by EPHA4 through the CDK5 kinase.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cell projection</location>
            <location evidence="3">Growth cone</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Membrane</location>
        </subcellularLocation>
    </comment>
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        <property type="protein sequence ID" value="ENSGAGP00000001835.1"/>
        <property type="gene ID" value="ENSGAGG00000001461.1"/>
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        <property type="component" value="Unassembled WGS sequence"/>
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        <property type="project" value="UniProtKB-SubCell"/>
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        <property type="term" value="C:growth cone"/>
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        <property type="project" value="UniProtKB-SubCell"/>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
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        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
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        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
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        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
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        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd01221">
        <property type="entry name" value="PH_ephexin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00160">
        <property type="entry name" value="RhoGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd11939">
        <property type="entry name" value="SH3_ephexin1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Ephexin-1 isoform X1"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Neuronal guanine nucleotide exchange factor"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="rho guanine nucleotide exchange factor 19"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.900.10">
        <property type="entry name" value="Dbl homology (DH) domain"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="DBL_dom_sf"/>
    </dbReference>
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        <property type="entry name" value="DH_dom"/>
    </dbReference>
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        <property type="entry name" value="Ephexin-1_SH3"/>
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    <dbReference type="InterPro" id="IPR047271">
        <property type="entry name" value="Ephexin-like"/>
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        <property type="entry name" value="PH-like_dom_sf"/>
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    </dbReference>
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        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055251">
        <property type="entry name" value="SOS1_NGEF_PH"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12845:SF8">
        <property type="entry name" value="EPHEXIN-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12845">
        <property type="entry name" value="GUANINE NUCLEOTIDE EXCHANGE FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
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    </dbReference>
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        <property type="entry name" value="DBL homology domain (DH-domain)"/>
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        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
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    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="12" id="KW-0966">Cell projection</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="5" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="9" id="KW-0221">Differentiation</keyword>
    <keyword evidence="8" id="KW-0344">Guanine-nucleotide releasing factor</keyword>
    <keyword evidence="11" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 18" id="KW-0728">SH3 domain</keyword>
    <feature type="domain" description="DH" evidence="22">
        <location>
            <begin position="178"/>
            <end position="362"/>
        </location>
    </feature>
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        <location>
            <begin position="394"/>
            <end position="506"/>
        </location>
    </feature>
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        <location>
            <begin position="517"/>
            <end position="578"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1"/>
            <end position="26"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="109"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="19">
        <location>
            <begin position="115"/>
            <end position="128"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="133"/>
            <end position="142"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
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        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00004496"/>
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    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
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    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        </source>
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    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
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    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
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            <dbReference type="SAM" id="MobiDB-lite"/>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
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            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="609" mass="70391" checksum="D2BC871580A081D0" modified="2019-05-08" version="1">MASLKNGDSRSKPLNLDVLSRKGPPVSVKSEVKDLEEQWIQRESPSTAGLVLIPDESPVVPECPAVLSETLQMIHPMTTDSWKTLIEQIGLLYQEYRDKSTLQEIETRRLQDAQTDPEECSASEEVPSEAEPTNTAESRTLPQISLLRNSTSRFNLWQDLPEIRSSGVLNILQPDEIKLQEAMFELVTSEASYYKSLNLLVSHFMENERLKKILHPSEAHILFSNVLDVMAVSERFLLDLEQRVEENIVISDVCDIVYQHTVNHFSVYVTYVSNQTYQERAYKQLLQDKTAFREVISQLEMDPKCKGLPFSSFLILPFQRITRLKLLVQNILKKVEEKSEKETTALEAHKELETVVKACNEGVRKMSRTEQMISIQKKLEFKIKSVPIISHSRWLLKQGELQQMNGPKTSRTLRTKKLFREIYLFLFNDLLVLCRQIPGDKYQVFDSAPRGLLRVEELEDQGQSLANVFILRLLENTDDREVSYMLKASSQSEMKRWMISLAPNRRTKFVSFTSRLLDCPQVQCVHPYVAQQPDELSLELADVLNILDKTDDGWIFGERLHDQERGWFPSSMGEEILNPKIRSQNLKECFRVHKSDDSQRRKLGSRNRQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJH7</accession>
    <name>A0A452GJH7_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="8">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
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        <lineage>
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            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GJH7"/>
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    </dbReference>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="C:presynaptic membrane"/>
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        <property type="project" value="TreeGrafter"/>
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        <property type="project" value="TreeGrafter"/>
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        <property type="term" value="C:terminal bouton"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019992">
        <property type="term" value="F:diacylglycerol binding"/>
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        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005543">
        <property type="term" value="F:phospholipid binding"/>
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        <property type="project" value="InterPro"/>
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    </dbReference>
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        <property type="term" value="F:zinc ion binding"/>
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        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="P:presynaptic dense core vesicle exocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035249">
        <property type="term" value="P:synaptic transmission, glutamatergic"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="P:synaptic vesicle docking"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016082">
        <property type="term" value="P:synaptic vesicle priming"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="entry name" value="C2B_Munc13"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Protein unc-13 homolog B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000002">
        <property type="entry name" value="Protein unc-13 homolog B"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000008">
        <property type="entry name" value="C2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010439">
        <property type="entry name" value="MUN_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014770">
        <property type="entry name" value="Munc13_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014772">
        <property type="entry name" value="Munc13_dom-2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027080">
        <property type="entry name" value="Unc-13"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037302">
        <property type="entry name" value="Unc-13_C2B"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10480">
        <property type="entry name" value="PROTEIN UNC-13 HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10480:SF2">
        <property type="entry name" value="PROTEIN UNC-13 HOMOLOG C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00168">
        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06292">
        <property type="entry name" value="MUN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00360">
        <property type="entry name" value="C2DOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00239">
        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM01145">
        <property type="entry name" value="DUF1041"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49562">
        <property type="entry name" value="C2 domain (Calcium/lipid-binding domain, CaLB)"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50004">
        <property type="entry name" value="C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51258">
        <property type="entry name" value="MHD1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51259">
        <property type="entry name" value="MHD2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0106">Calcium</keyword>
    <keyword evidence="1" id="KW-0268">Exocytosis</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0862">Zinc</keyword>
    <keyword evidence="3" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2" evidence="5">
        <location>
            <begin position="36"/>
            <end position="160"/>
        </location>
    </feature>
    <feature type="domain" description="MHD1" evidence="6">
        <location>
            <begin position="468"/>
            <end position="611"/>
        </location>
    </feature>
    <feature type="domain" description="MHD2" evidence="7">
        <location>
            <begin position="710"/>
            <end position="845"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS51258"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS51259"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001756.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1031" mass="118940" checksum="CB1FC629654610A1" modified="2019-05-08" version="1">IIAMKERMKIRERNRPEVFEVIQEMFQISKEDFVQYTKAAKQCVLDGTSKWSAKITITVVCAQGLQAKDKTGSSDPYVTVQVGKTKRRTKTIFGNLNPVWDEKFYFECHNSTDRIKVRVWDEDDDIKSRVKQHFKKESDDFLGQTIIEVRTLSGEMDVWYNLEKRTDKSAVSGAIRLKINVEIKGEEKVAPYHVQYTYLHENLFHYLTEVKSNGSVKIPEVRGDEAWKVYFDAAAQEIVDEFAMRYGIEYIYQAMTHFSCLSSKYMCSGVPAVMSTLLANINAFYAHTIATTNVSASDRFAATNFGREKFIKLLDQLHNSLRIDLSKYRDNFPASNSERLQDLKSTVDLLTSITFFRMKVLELQSPPRASMVVKDCVRACLDSTYKYIFDNCYELYSQLMDQGKKQEVPREEQGPTTKNLDFWAQLITLMVTIIDEDKTAYTPVLNQFPQELNMGKVSAEIMWTLFAQDMKYALEEHEKQRLCKSTDYMNLHFKVKWFYNEYVRELPALQDAVPEYSLWFEPFVIQWLDENEDVSMEFLHGALERDKKDGFQQTSDHALFSCSVVDVFTQLNQSFEIIKKLECPNPEALSHLMRRFAKTINKVLLQYAAVISNDFNSYCDKEMLPCILMNNIQQLRVQLEKMFESMGGKELMEQNLNSHLLSYNSSTLFFLFSFQLVVEECVRQMSFELNQMRGNGNISANKNNAAIDAEIVLRPLMDFLDKILSLSAKICEKTVLKRVLKELWKSVLNKIEKQIVLPPLTDQTVSVQMFFKKAREDHVTREEARSLTLRQCAIMEVALSTIKQYFHAGGNGLKKNFLEKSPDLQSLRYALSLYTQTTDALIKKFIATQTSQSSSPHGQKEEYFALSHSTNNTVGEISVQVDVFTHPGTGEHRVTVKVVAINNLNWQTTAMFRPFVEVCILGPNLSDKKRKHGTKTKIFFSFKGLRAYELHLSVKDYCFAREDRIVGMTVIQLQNTAEKGSCASWYPLLKNIFMDDTGLTILRILSQRTNDEVAKEFVRLKSETRSTEETV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJI2</accession>
    <name>A0A452GJI2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">cAMP-dependent protein kinase catalytic subunit alpha</fullName>
            <ecNumber evidence="6">2.7.11.11</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="26 27">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="27" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="26" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="26" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="22">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.11"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="21">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.11"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Mitochondrion</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the protein kinase superfamily. AGC Ser/Thr protein kinase family. cAMP subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.11" evidence="6"/>
    <dbReference type="AlphaFoldDB" id="A0A452GJI2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002003.1">
        <property type="protein sequence ID" value="ENSGAGP00000001761.1"/>
        <property type="gene ID" value="ENSGAGG00000001402.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005952">
        <property type="term" value="C:cAMP-dependent protein kinase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004691">
        <property type="term" value="F:cAMP-dependent protein kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034237">
        <property type="term" value="F:protein kinase A regulatory subunit binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd14209">
        <property type="entry name" value="STKc_PKA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000005">
        <property type="entry name" value="cAMP-dependent protein kinase catalytic subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000005">
        <property type="entry name" value="cAMP-dependent protein kinase catalytic subunit alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000961">
        <property type="entry name" value="AGC-kinase_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044109">
        <property type="entry name" value="STKc_PKA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24353:SF82">
        <property type="entry name" value="CAMP-DEPENDENT PROTEIN KINASE CATALYTIC SUBUNIT ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24353">
        <property type="entry name" value="CYCLIC NUCLEOTIDE-DEPENDENT PROTEIN KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00133">
        <property type="entry name" value="S_TK_X"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51285">
        <property type="entry name" value="AGC_KINASE_CTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="16 23" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="19" id="KW-0114">cAMP</keyword>
    <keyword evidence="8" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="9" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="15" id="KW-0418">Kinase</keyword>
    <keyword evidence="13" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="18" id="KW-0472">Membrane</keyword>
    <keyword evidence="17" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="13" id="KW-0519">Myristate</keyword>
    <keyword evidence="14 23" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="20" id="KW-0539">Nucleus</keyword>
    <keyword evidence="11" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="27" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="12" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Protein kinase" evidence="24">
        <location>
            <begin position="31"/>
            <end position="259"/>
        </location>
    </feature>
    <feature type="domain" description="AGC-kinase C-terminal" evidence="25">
        <location>
            <begin position="260"/>
            <end position="312"/>
        </location>
    </feature>
    <feature type="binding site" evidence="23">
        <location>
            <position position="60"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004173"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00007115"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00012444"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00020608"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022707"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023149"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00047292"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00047454"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS51285"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001761.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="312" mass="36056" checksum="BBF4C9DD63439675" modified="2019-05-08" version="1">SIIGSSHVSLYPQKQHDLFSPFQNTASLDQFERIKTLGTGSFGRVLLVKHKETGNYYAMKVLEKQKVVKLKQIEHTLNEKRILQAINFPLLIKLEYSFKDNSNLYMVMEYIPGGEMFSHLQRIGRFSEPHARFYAAQIQGYIEVMDFGFDKRVKGRTWTLCGTPEYLAPEIILSKGYNKAVDWWALGVLIYEMAAGYPPFFADQPIQIYEKIVSGKVRFPSHFSSDLKDLLRNLLQVDLTKRFGNLKNGVNDIKNHKWFATTDWIAIYQRKVEAPFVPKCKGPGDTSNFDDYEEEEIRVSTTEECAMEFAEF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJI4</accession>
    <name>A0A452GJI4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">RRM domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GJI4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002098.1">
        <property type="protein sequence ID" value="ENSGAGP00000001845.1"/>
        <property type="gene ID" value="ENSGAGG00000001433.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009897">
        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003954">
        <property type="term" value="F:NADH dehydrogenase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003756">
        <property type="term" value="F:protein disulfide isomerase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007624">
        <property type="term" value="P:ultradian rhythm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.330">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038876">
        <property type="entry name" value="ENOX"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056611">
        <property type="entry name" value="ENOX1/2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012677">
        <property type="entry name" value="Nucleotide-bd_a/b_plait_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035979">
        <property type="entry name" value="RBD_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000504">
        <property type="entry name" value="RRM_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16001">
        <property type="entry name" value="ECTO-NOX DISULFIDE-THIOL EXCHANGER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16001:SF6">
        <property type="entry name" value="ECTO-NOX DISULFIDE-THIOL EXCHANGER 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23267">
        <property type="entry name" value="ENOX1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00076">
        <property type="entry name" value="RRM_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00360">
        <property type="entry name" value="RRM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54928">
        <property type="entry name" value="RNA-binding domain, RBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50102">
        <property type="entry name" value="RRM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0694">RNA-binding</keyword>
    <feature type="domain" description="RRM" evidence="4">
        <location>
            <begin position="76"/>
            <end position="132"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="295"/>
            <end position="314"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="2">
        <location>
            <begin position="206"/>
            <end position="233"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="2">
        <location>
            <begin position="325"/>
            <end position="424"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="3">
        <location>
            <begin position="298"/>
            <end position="307"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00176"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50102"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001845.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="540" mass="61864" checksum="97E9C3B01FC17ADF" modified="2019-05-08" version="1">VFTDSICVPGFDPSLSMMTGITPINPMIPGMGLVPPPPPTDVPVVKEIIHCKSCTLFPPNPNLPPPSTRERPPGCKTVFVGGLPENATEEIIQEVFEQCGDITAIRKSKKNFCHIRFAEEFMVDKAIYLSGTLHSLIDFLCFSFVFYDANVFPCFCLSGREYMSCLQIEPLVSPALKFSEAITVLLTWIERGEVNRRSANQFYSMVQSANSHVRRLMNEKATHEQEMEQAKESFKNALTGILTQFEQIVAVFNASTRQKAWDHFSKAQRKNIDIWRKHSEELRNAHSEQLMGIRREEEMEMSDDDSSDNPSKKLRVDESALTAQAYALKEENDSLRWQLDAYRNEVELLKQEKGQLFRTEESLTKDQQLKFLQQTMQGMQQQLLSIQEELNNKKSELQQAKEEQTHTEARLKVLQEQLKSAKELAEINGQDESQEINMLTVALVNQEREKNSEKRSQGLKSEKEALLIGIISTFLHVHPFGANIEYLWSYMQQLDSRISANEIETLLMRLPRMFKQEFTGVGATLEKRWKFCAFEGIKTT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJI6</accession>
    <name>A0A452GJI6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Zinc finger and BTB domain-containing protein 44</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJI6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002008.1">
        <property type="protein sequence ID" value="ENSGAGP00000001766.1"/>
        <property type="gene ID" value="ENSGAGG00000001414.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd18228">
        <property type="entry name" value="BTB_POZ_ZBTB44"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000266">
        <property type="entry name" value="zinc finger and BTB domain-containing protein 44 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000278">
        <property type="entry name" value="zinc finger and BTB domain-containing protein 44 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.710.10">
        <property type="entry name" value="Potassium Channel Kv1.1, Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000210">
        <property type="entry name" value="BTB/POZ_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011333">
        <property type="entry name" value="SKP1/BTB/POZ_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050457">
        <property type="entry name" value="ZnFinger_BTB_dom_contain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46105">
        <property type="entry name" value="AGAP004733-PA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46105:SF5">
        <property type="entry name" value="ZINC FINGER AND BTB DOMAIN-CONTAINING PROTEIN 44 ISOFORM X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00651">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00225">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54695">
        <property type="entry name" value="POZ domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50097">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="2" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9" id="KW-0539">Nucleus</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="8" id="KW-0804">Transcription</keyword>
    <keyword evidence="6" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <keyword evidence="4 11" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="BTB" evidence="13">
        <location>
            <begin position="31"/>
            <end position="98"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="14">
        <location>
            <begin position="399"/>
            <end position="426"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="14">
        <location>
            <begin position="427"/>
            <end position="449"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="295"/>
            <end position="364"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="304"/>
            <end position="318"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00070979"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50097"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001766.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="453" mass="50234" checksum="CDB712B5BB4431E4" modified="2019-05-08" version="1">MGIKTFTHNSPAHSQEMLGKLNMLRNDGHFCDITIRVQDKIFRAHKVVLAACSDFFRSKLVGQAEDDGKSVLDLHHVTVTGFIPLLEYAYTATLSINTENIIDVLAAASYMQMFSVASTCSEFMKSSILWNAPNSQQDKVLDAGQENSANRNFTSRAGSISPVSSECSVVERAIPVCPESRRKRKSYIVMSPESPLKCNTQTNSPQVLNPSASYAESRNQPVDSSLAFPWTFSFGIDQRLQSEKVKQVENARTLELPGPSEAARRITDYVTCESTKASSPLVIEEDVRVKVERLSDEEVHEEVSQPVSASQSSLSDQQTVPGSEQVQEDLLISPQPSSIGSIDEGVTEGLPTLQSTSGTNARADDDDRLENVQYPYQLYIAPSTSSTERPSPNCPDRPFQCSTCGVRFTRIQNLKQHMLIHSGIKPFQCDRCGKKFTRAYSLKMHRLKHEVIS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJJ9</accession>
    <name>A0A452GJJ9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Autophagy-related protein 16 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
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            <authorList>
                <consortium name="Ensembl"/>
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        <scope>IDENTIFICATION</scope>
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        <text evidence="1">Belongs to the WD repeat ATG16 family.</text>
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        <property type="component" value="Unassembled WGS sequence"/>
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        <property type="entry name" value="Autophagy-related protein 16-1 isoform 1"/>
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    </dbReference>
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    </dbReference>
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        <property type="entry name" value="AUTOPHAGY-RELATED PROTEIN 16-1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="ATG16"/>
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        <property type="entry name" value="GPROTEINBRPT"/>
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    </dbReference>
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    </dbReference>
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            <begin position="202"/>
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    </feature>
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        <source>
            <dbReference type="ARBA" id="ARBA00009271"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022574"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
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    </evidence>
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        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
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        </source>
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        <source>
            <dbReference type="Pfam" id="PF08614"/>
        </source>
    </evidence>
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        <source>
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    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="588" mass="66531" checksum="AE745A221269F2D0" modified="2019-05-08" version="1">MTSGLKAATFPTWKRHIAEELRRRDRLQRQAFEEIILQYNKLLEKTDLHAVLADKLQAEKYDLQSRHEISPGHDGTRNEAQLQEMAQLRIKHQEELTELHKKRGELAQSVIDLNNQMQQKDREMQINEAKIAEYMQKISELETECQELRSKLQELERANQTLKDEYDALQITFTALEEKLRKTTEDNQELVTRWMAEKAQEANRLNAENEKDSRRRQAKLQKELAEAAKEPLPVEQDDDIEVLVDETSDPAEETSPVRAVSRTSRRRSLSSFPAPQENVDVHPGSSKEVRVPTTAVYVFDAHDGEVNAVQFSPASRLLATGGMDRRVKLWEVFGDRCELKSSLSGSNAGITSIEFDSAGSYLLAASNDFASRIWTVDDYRLRHTLTGHSGKVLSAKFLLDNARIVSGSHDRTLKLWDLRSKVCIKTVFAGSSCNDIVCTEQCVMSGHFDKKIRFWDIRTESIVQELELFGRITALDLNPERTELLTCSRDDLLKIIDLRVNAVKQTFSAQGFKCGSDWTRVVFSPDGSYVAGGSADGVLYIWNVLTGKVERTLAKHHSSSINAVAWSPAGAHVVSVDKGNKAVLWSEF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJK0</accession>
    <name>A0A452GJK0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Autophagy-related protein 16 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the WD repeat ATG16 family.</text>
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    <dbReference type="AlphaFoldDB" id="A0A452GJK0"/>
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        <property type="gene ID" value="ENSGAGG00000001408.1"/>
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    </dbReference>
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    </dbReference>
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    </dbReference>
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    </dbReference>
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    </dbReference>
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    <feature type="domain" description="Autophagy-related protein 16" evidence="6">
        <location>
            <begin position="16"/>
            <end position="206"/>
        </location>
    </feature>
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        <location>
            <begin position="315"/>
            <end position="348"/>
        </location>
    </feature>
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        <location>
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            <end position="400"/>
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        <location>
            <begin position="401"/>
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    </feature>
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        <location>
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    </feature>
    <feature type="repeat" description="WD" evidence="4">
        <location>
            <begin position="537"/>
            <end position="568"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="4">
        <location>
            <begin position="570"/>
            <end position="604"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="202"/>
            <end position="303"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="202"/>
            <end position="229"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="5">
        <location>
            <begin position="235"/>
            <end position="252"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="5">
        <location>
            <begin position="264"/>
            <end position="274"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00009271"/>
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    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022574"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="Pfam" id="PF08614"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001765.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="604" mass="68111" checksum="8169399F6369950C" modified="2019-05-08" version="1">MTSGLKAATFPTWKRHIAEELRRRDRLQRQAFEEIILQYNKLLEKTDLHAVLADKLQAEKYDLQSRHEISPGHDGTRNEAQLQEMAQLRIKHQEELTELHKKRGELAQSVIDLNNQMQQKDREMQINEAKIAEYMQKISELETECQELRSKLQELERANQTLKDEYDALQITFTALEEKLRKTTEDNQELVTRWMAEKAQEANRLNAENEKDSRRRQAKLQKELAEAAKEPLPVEQDDDIEVLVDETSDPAEETSPVRAVSRTSSLSESSLLGHHSSDAARRRSLSSFPAPQENVDVHPGSSKEVRVPTTAVYVFDAHDGEVNAVQFSPASRLLATGGMDRRVKLWEVFGDRCELKSSLSGSNAGITSIEFDSAGSYLLAASNDFASRIWTVDDYRLRHTLTGHSGKVLSAKFLLDNARIVSGSHDRTLKLWDLRSKVCIKTVFAGSSCNDIVCTEQCVMSGHFDKKIRFWDIRTESIVQELELFGRITALDLNPERTELLTCSRDDLLKIIDLRVNAVKQTFSAQGFKCGSDWTRVVFSPDGSYVAGGSADGVLYIWNVLTGKVERTLAKHHSSSINAVAWSPAGAHVVSVDKGNKAVLWSEF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJK1</accession>
    <name>A0A452GJK1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3 12">Cytochrome b-c1 complex subunit 7</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="12">Component of the ubiquinol-cytochrome c oxidoreductase, a multisubunit transmembrane complex that is part of the mitochondrial electron transport chain which drives oxidative phosphorylation.</text>
    </comment>
    <comment type="subunit">
        <text evidence="11">Component of the ubiquinol-cytochrome c oxidoreductase (cytochrome b-c1 complex, complex III, CIII), a multisubunit enzyme composed of 11 subunits. The complex is composed of 3 respiratory subunits cytochrome b, cytochrome c1 and Rieske protein UQCRFS1, 2 core protein subunits UQCRC1/QCR1 and UQCRC2/QCR2, and 6 low-molecular weight protein subunits UQCRH/QCR6, UQCRB/QCR7, UQCRQ/QCR8, UQCR10/QCR9, UQCR11/QCR10 and subunit 9, the cleavage product of Rieske protein UQCRFS1. The complex exists as an obligatory dimer and forms supercomplexes (SCs) in the inner mitochondrial membrane with NADH-ubiquinone oxidoreductase (complex I, CI) and cytochrome c oxidase (complex IV, CIV), resulting in different assemblies (supercomplex SCI(1)III(2)IV(1) and megacomplex MCI(2)III(2)IV(2)).</text>
    </comment>
    <comment type="subunit">
        <text evidence="10">Component of the ubiquinol-cytochrome c oxidoreductase (cytochrome b-c1 complex, complex III, CIII), a multisubunit enzyme composed of 3 respiratory subunits cytochrome b, cytochrome c1 and Rieske protein, 2 core protein subunits, and additional low-molecular weight protein subunits. The complex exists as an obligatory dimer and forms supercomplexes (SCs) in the inner mitochondrial membrane with cytochrome c oxidase (complex IV, CIV).</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Mitochondrion inner membrane</location>
            <topology evidence="1">Peripheral membrane protein</topology>
            <orientation evidence="1">Matrix side</orientation>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 12">Belongs to the UQCRB/QCR7 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJK1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002122.1">
        <property type="protein sequence ID" value="ENSGAGP00000001865.1"/>
        <property type="gene ID" value="ENSGAGG00000001495.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005743">
        <property type="term" value="C:mitochondrial inner membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045275">
        <property type="term" value="C:respiratory chain complex III"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006122">
        <property type="term" value="P:mitochondrial electron transport, ubiquinol to cytochrome c"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.1090.10:FF:000001">
        <property type="entry name" value="Cytochrome b-c1 complex subunit 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1090.10">
        <property type="entry name" value="Cytochrome b-c1 complex subunit 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003197">
        <property type="entry name" value="QCR7"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036544">
        <property type="entry name" value="QCR7_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12022:SF0">
        <property type="entry name" value="CYTOCHROME B-C1 COMPLEX SUBUNIT 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12022">
        <property type="entry name" value="UBIQUINOL-CYTOCHROME C REDUCTASE COMPLEX 14 KD PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02271">
        <property type="entry name" value="UCR_14kD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000022">
        <property type="entry name" value="Bc1_14K"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81524">
        <property type="entry name" value="14 kDa protein of cytochrome bc1 complex (Ubiquinol-cytochrome c reductase)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7 12" id="KW-0249">Electron transport</keyword>
    <keyword evidence="9 12" id="KW-0472">Membrane</keyword>
    <keyword evidence="8 12" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="6 12" id="KW-0999">Mitochondrion inner membrane</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 12" id="KW-0679">Respiratory chain</keyword>
    <keyword evidence="4 12" id="KW-0813">Transport</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008554"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00016323"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022660"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022792"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022982"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00038521"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00046393"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PIRNR" id="PIRNR000022"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001865.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="111" mass="13396" checksum="2F75F8C4D4FF098C" modified="2019-05-08" version="1">MAARAPASGSGRLLEGLRKWYYNAAGFNKLGLMRDDTIYENDDVKVALKRLPENLYNERLFRIKRALDLSMRQQILPEEQWVKYEEDKKYLEPYLKEVVRERLEREEWDKK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJK4</accession>
    <name>A0A452GJK4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="23">Chromodomain-helicase-DNA-binding protein 4</fullName>
        </recommendedName>
    </protein>
    <organism evidence="31 32">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="32" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="31" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="31" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="24">ATP-dependent chromatin-remodeling factor that binds and distorts nucleosomal DNA. Acts as a component of the histone deacetylase NuRD complex which participates in the remodeling of chromatin. Localizes to acetylated damaged chromatin in a ZMYND8-dependent manner, to promote transcriptional repression and double-strand break repair by homologous recombination. Involved in neurogenesis.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="22">
            <text>ATP + H2O = ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:13065"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
            <location evidence="2">Microtubule organizing center</location>
            <location evidence="2">Centrosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the SNF2/RAD54 helicase family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000002026.1">
        <property type="protein sequence ID" value="ENSGAGP00000001781.1"/>
        <property type="gene ID" value="ENSGAGG00000001416.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016581">
        <property type="term" value="C:NuRD complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140658">
        <property type="term" value="F:ATP-dependent chromatin remodeler activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003682">
        <property type="term" value="F:chromatin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140297">
        <property type="term" value="F:DNA-binding transcription factor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042393">
        <property type="term" value="F:histone binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd18667">
        <property type="entry name" value="CD1_tandem_CHD3-4_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18662">
        <property type="entry name" value="CD2_tandem_CHD3-4_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd15531">
        <property type="entry name" value="PHD1_CHD_II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd15532">
        <property type="entry name" value="PHD2_CHD_II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18793">
        <property type="entry name" value="SF2_C_SNF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.60:FF:000037">
        <property type="entry name" value="chromodomain-helicase-DNA-binding protein 3 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.50.40:FF:000003">
        <property type="entry name" value="chromodomain-helicase-DNA-binding protein 3 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.40.10:FF:000001">
        <property type="entry name" value="chromodomain-helicase-DNA-binding protein 3 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.40.10:FF:000011">
        <property type="entry name" value="chromodomain-helicase-DNA-binding protein 4 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.50.40:FF:000011">
        <property type="entry name" value="chromodomain-helicase-DNA-binding protein 4 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000015">
        <property type="entry name" value="chromodomain-helicase-DNA-binding protein 9 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.50.40">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.60">
        <property type="entry name" value="Homeodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.10810">
        <property type="entry name" value="Tandem AAA-ATPase domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012957">
        <property type="entry name" value="CHD_C2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009462">
        <property type="entry name" value="CHD_II_SANT-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012958">
        <property type="entry name" value="CHD_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016197">
        <property type="entry name" value="Chromo-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000953">
        <property type="entry name" value="Chromo/chromo_shadow_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023780">
        <property type="entry name" value="Chromo_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002464">
        <property type="entry name" value="DNA/RNA_helicase_DEAH_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009463">
        <property type="entry name" value="DUF1087"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014001">
        <property type="entry name" value="Helicase_ATP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001650">
        <property type="entry name" value="Helicase_C-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038718">
        <property type="entry name" value="SNF2-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049730">
        <property type="entry name" value="SNF2/RAD54-like_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000330">
        <property type="entry name" value="SNF2_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019786">
        <property type="entry name" value="Zinc_finger_PHD-type_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011011">
        <property type="entry name" value="Znf_FYVE_PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001965">
        <property type="entry name" value="Znf_PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019787">
        <property type="entry name" value="Znf_PHD-finger"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45623">
        <property type="entry name" value="CHROMODOMAIN-HELICASE-DNA-BINDING PROTEIN 3-RELATED-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45623:SF22">
        <property type="entry name" value="CHROMODOMAIN-HELICASE-DNA-BINDING PROTEIN 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08074">
        <property type="entry name" value="CHDCT2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06461">
        <property type="entry name" value="CHDII_SANT-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08073">
        <property type="entry name" value="CHDNT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00385">
        <property type="entry name" value="Chromo"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06465">
        <property type="entry name" value="DUF1087"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00271">
        <property type="entry name" value="Helicase_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00628">
        <property type="entry name" value="PHD"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00176">
        <property type="entry name" value="SNF2-rel_dom"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00298">
        <property type="entry name" value="CHROMO"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00487">
        <property type="entry name" value="DEXDc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01146">
        <property type="entry name" value="DUF1086"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01147">
        <property type="entry name" value="DUF1087"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00490">
        <property type="entry name" value="HELICc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00249">
        <property type="entry name" value="PHD"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54160">
        <property type="entry name" value="Chromo domain-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57903">
        <property type="entry name" value="FYVE/PHD zinc finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50013">
        <property type="entry name" value="CHROMO_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00690">
        <property type="entry name" value="DEAH_ATP_HELICASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51192">
        <property type="entry name" value="HELICASE_ATP_BIND_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51194">
        <property type="entry name" value="HELICASE_CTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01359">
        <property type="entry name" value="ZF_PHD_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50016">
        <property type="entry name" value="ZF_PHD_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="16" id="KW-0007">Acetylation</keyword>
    <keyword evidence="13" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="15" id="KW-0156">Chromatin regulator</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="20" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="18" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="11" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="5" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="21" id="KW-0539">Nucleus</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="32" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="19" id="KW-0804">Transcription</keyword>
    <keyword evidence="17" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="14" id="KW-0832">Ubl conjugation</keyword>
    <keyword evidence="12" id="KW-0862">Zinc</keyword>
    <keyword evidence="10 25" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="PHD-type" evidence="28">
        <location>
            <begin position="425"/>
            <end position="472"/>
        </location>
    </feature>
    <feature type="domain" description="PHD-type" evidence="28">
        <location>
            <begin position="504"/>
            <end position="551"/>
        </location>
    </feature>
    <feature type="domain" description="Chromo" evidence="27">
        <location>
            <begin position="584"/>
            <end position="641"/>
        </location>
    </feature>
    <feature type="domain" description="Chromo" evidence="27">
        <location>
            <begin position="677"/>
            <end position="712"/>
        </location>
    </feature>
    <feature type="domain" description="Helicase ATP-binding" evidence="29">
        <location>
            <begin position="824"/>
            <end position="910"/>
        </location>
    </feature>
    <feature type="domain" description="Helicase C-terminal" evidence="30">
        <location>
            <begin position="1042"/>
            <end position="1191"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="26">
        <location>
            <begin position="26"/>
            <end position="208"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="26">
        <location>
            <begin position="303"/>
            <end position="350"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="26">
        <location>
            <begin position="380"/>
            <end position="419"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="26">
        <location>
            <begin position="571"/>
            <end position="592"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="26">
        <location>
            <begin position="633"/>
            <end position="658"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="26">
        <location>
            <begin position="1331"/>
            <end position="1388"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="26">
        <location>
            <begin position="1516"/>
            <end position="1660"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="26">
        <location>
            <begin position="1886"/>
            <end position="1908"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="26">
        <location>
            <begin position="33"/>
            <end position="53"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="26">
        <location>
            <begin position="95"/>
            <end position="105"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="26">
        <location>
            <begin position="167"/>
            <end position="185"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="26">
        <location>
            <begin position="189"/>
            <end position="199"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="26">
        <location>
            <begin position="380"/>
            <end position="393"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="26">
        <location>
            <begin position="394"/>
            <end position="407"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="26">
        <location>
            <begin position="1522"/>
            <end position="1531"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="26">
        <location>
            <begin position="1549"/>
            <end position="1572"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="26">
        <location>
            <begin position="1606"/>
            <end position="1637"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="26">
        <location>
            <begin position="1649"/>
            <end position="1660"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007025"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00049360"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00074666"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00093442"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00146"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="27">
        <source>
            <dbReference type="PROSITE" id="PS50013"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="28">
        <source>
            <dbReference type="PROSITE" id="PS50016"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="29">
        <source>
            <dbReference type="PROSITE" id="PS51192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="30">
        <source>
            <dbReference type="PROSITE" id="PS51194"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="31">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001781.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="32">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1908" mass="217649" checksum="4E04FC3BBAED2CA6" modified="2019-05-08" version="1">MGWKRERGGAWPGFFPFKRWCRSRGHFPFSGGSERRGAGGRAGKETKLQKLSEPKYPNLEDMASGIGSPSPCSAGSDDDEMEILLNNSIPQHPEPEEEPEEELLSEAETPKIKKKKKPKKLKEPKVPKLSKRQKKELGDSSGEGNEFVEEEEEVLHSDSEGSDYTPGRKKKKKLGPKKEKKNKAKRKEEEEEEEEDDDSKEPKSSAQLLEDWGMEDIDHIFTEEDYRTLTNYKAFSQFVRPLIAAKNPKIAVSKMMMVLGAKWREFSTNNPFKGSSGASVAAAAAAAVAVVESMVANVDAVLPQPPADVPLRKAKTKEGKGPNARRKPKASPRVPDIKKPKTKKVAPLKIKLGGFSSKRKRSSSEDEDLDVESDFDDASINSYSVSDGSTSRSSRSRKKLKAGKRKKKGEDDSTVPVDGYETDHQDYCEVCQQGGEIILCDTCPRAYHMVCLDPDMEKAPEGKWSCPHCEKEGIQWEAKEDNSEGEEILEDVVGDPEEEDDHHMEFCRVCKDGGELLCCDACPSSYHIHCLNPPLPEIPNGEWLCPRCTCPSLKGKVQKILIWKWGQPPLPTPVPRPADADPNTPSPRPLEGRPERQFFVKWQGMSYWHCSWVSELQLELHCQVMFRNYQRKNDMDEPPSGDFGGEEEKSRKRKNKDPKFAEMEERFYRYGIKPEWMMIHRILNHSVDKKGNVHYLIKWRDLPYDQASWESEDVEVQDYDLYKQAYWNHRELMRGEEGRPGKKIKKVKMRKLERPPDTPTVDPTVKYDRQPEYLDVTGGTLHPYQLEGLNWLRFSWAQGTDTILADEMGLGKTVQTAVFLYSLYKEKEASVKFHVLLTSYELITIDMAILGSIDWACLIVDEAHRLKNNQSKFFRVLNGYSLQHKLLLTGTPLQNNLEELFHLLNFLTPERFHNLEGFLEEFADIAKEDQIKKLHDMLGPHMLRRLKADVFKNMPSKTELIVRVELSPMQKKYYKYILTRNFEALNARGGGNQVSLLNVVMDLKKCCNHPYLFPVAAMEAPKMPNGMYDGSALIRASGKLLLLQKMLKNLKEGGHRVLIFSQMTKMLDLLEDFLEHEGYKYERIDGGITGNMRQEAIDRFNAPGAQQFCFLLSTRAGGLGINLATADTVIIYDSDWNPHNDIQAFSRAHRIGQNKKVMIYRFVTRASVEERITQVAKKKMMLTHLVVRPGLGSKTGSMSKQELDDILKFGTEELFKDEATEGGDSKEGEDSSVIHYDDKAIERLLDRNQDETEDTELQGMNEYLSSFKVAQYVVREEEMGEEEEVEREIIKQEESVDPDYWEKLLRHHYEQQQEDLARNLGKGKRIRKQVNYNDGSQEDRDWQDDQSDNQSDYSVASEEGDEDFDERSEAARRPSRKGLRNDKDKPLPPLLARVGGNIEVLGFNARQRKAFLNAIMRYGMPPQDAFTTQWLVRDLRGKSEKEFKAYVSLFMRHLCEPGADGAETFADGVPREGLSRQHVLTRIGVMSLIRKKVQEFEHVNGRWSMPELAEIEENKKLLQPGSPSPKTPTPSTPGDTQPNTPAAVPLLEEGVKVEEGASTREQGELMESEKEPSLVSAEAEVPMEQCAQPVETPPQEAKSPMNPPEAEEKKPEEPEVKEEPMEMESKADVEKIEDRVSAEPSAEPPTINLDEKEEKKEDDKRDVVMLQNGEMLKDSLDERHKKAVKQRFMFNIADGGFTELHSLWQNEERAATVTKKTYEIWHRRHDYWLLAGIINHGYARWQDIQNDPRYAILNEPFKGEMNRGNFLEIKNKFLARRFKLLEQALVIEEQLRRAAYLNMSEDPSHPSMALNTRFAEVECLAESHQHLSKESMAGNKPANAVLHKVLKQLEELLSDMKADVTRLPATIARIPPVAVRLQMSERNILSRLANRSSEPPPPPPPQQVAQQQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJK5</accession>
    <name>A0A452GJK5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="23">NACHT domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Endoplasmic reticulum</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Golgi apparatus</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Inflammasome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Secreted</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJK5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002129.1">
        <property type="protein sequence ID" value="ENSGAGP00000001871.1"/>
        <property type="gene ID" value="ENSGAGG00000001488.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061702">
        <property type="term" value="C:canonical inflammasome complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012505">
        <property type="term" value="C:endomembrane system"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006954">
        <property type="term" value="P:inflammatory response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045087">
        <property type="term" value="P:innate immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd08321">
        <property type="entry name" value="Pyrin_ASC-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000442">
        <property type="entry name" value="NACHT, LRR and PYD domains-containing protein 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004020">
        <property type="entry name" value="DAPIN"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011029">
        <property type="entry name" value="DEATH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007111">
        <property type="entry name" value="NACHT_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041267">
        <property type="entry name" value="NLRP_HD2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050637">
        <property type="entry name" value="NLRP_innate_immun_reg"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041075">
        <property type="entry name" value="NOD1/2_WH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45690">
        <property type="entry name" value="NACHT, LRR AND PYD DOMAINS-CONTAINING PROTEIN 12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45690:SF19">
        <property type="entry name" value="NACHT, LRR AND PYD DOMAINS-CONTAINING PROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05729">
        <property type="entry name" value="NACHT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17776">
        <property type="entry name" value="NLRC4_HD2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02758">
        <property type="entry name" value="PYRIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17779">
        <property type="entry name" value="WHD_NOD2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00368">
        <property type="entry name" value="LRR_RI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01289">
        <property type="entry name" value="PYRIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47986">
        <property type="entry name" value="DEATH domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52047">
        <property type="entry name" value="RNI-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50824">
        <property type="entry name" value="DAPIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50837">
        <property type="entry name" value="NACHT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="12" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="15" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="10" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="18" id="KW-1271">Inflammasome</keyword>
    <keyword evidence="17" id="KW-0395">Inflammatory response</keyword>
    <keyword evidence="9" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0964">Secreted</keyword>
    <keyword evidence="16" id="KW-0804">Transcription</keyword>
    <keyword evidence="14" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="13" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="Pyrin" evidence="19">
        <location>
            <begin position="1"/>
            <end position="91"/>
        </location>
    </feature>
    <feature type="domain" description="NACHT" evidence="20">
        <location>
            <begin position="130"/>
            <end position="328"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004110"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004555"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023198"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023233"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001871.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="23">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="669" mass="76693" checksum="1DFDBFE9968FB0D7" modified="2019-05-08" version="1">MAGRISDLLVHALNNLTQGDFKRFKDKLSYSALKGRGNIPRDCLDNANRIDTKNLLMKFYGGEAAVDVTIDVFTQINLRDSAAKLREGKEKGKKSIVTLKPGKHEIMAMTWSVVTMDTLFTCDKQGQTPQIVVLLGAAGIGKTMTTRKIMCDWAAGELYKKKFDYVFYINCRETNLTEQGSVVDMILKNWPNTNAPVNEILMKPEKLLFIIDGFDELRFSFDQTENLCSDPWEKKPMEIILSSLFRKLILPQSYLLITSRPTALEKLLQCLECSRYAEILGFSEADRKEYFHKFFGNKKQARQALQFVKENDILFTMCFIPIMCWIICTVVKQQLEKGEDLAQTSKSVTGVYMLYISSLVKPLSSNLKQHMQANLRGLCSLAAYGIWKRKILFEEAELKKFGLNQADSFPLSSNKNIFQKDIDCVNAYSFIHLSFQEFFAALFYVLEEDEETMEDSETPKKDMKTLLENYGNSRNYLMLTVRFLFGLLNEEIMQIMEEKFSCKISSKIKPDLLKWVQTNQQKYLPLSNVYQETCRIYNLEEFHCMFEMYEENFVKSALDHVVALTLDSNKLTQMDQMAFSFCVKNCHRLESLCISYLSPDTTFPFCLSVSCRLVNCSLTAACCRDLSSVLNTKTTLTELDLSDKVLGNSGIKLLCEGLKHPNCKVQKLR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJK7</accession>
    <name>A0A452GJK7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Ribosomal protein L15</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="4">Component of the large ribosomal subunit. The ribosome is a large ribonucleoprotein complex responsible for the synthesis of proteins in the cell.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1 5">Belongs to the eukaryotic ribosomal protein eL15 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJK7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002128.1">
        <property type="protein sequence ID" value="ENSGAGP00000001870.1"/>
        <property type="gene ID" value="ENSGAGG00000001496.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0022625">
        <property type="term" value="C:cytosolic large ribosomal subunit"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003735">
        <property type="term" value="F:structural constituent of ribosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002181">
        <property type="term" value="P:cytoplasmic translation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.1120.10">
        <property type="entry name" value="Ribosomal protein l15e"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024794">
        <property type="entry name" value="Rbsml_eL15_core_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000439">
        <property type="entry name" value="Ribosomal_eL15"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020925">
        <property type="entry name" value="Ribosomal_eL15_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012678">
        <property type="entry name" value="Ribosomal_uL23/eL15/eS24_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11847:SF4">
        <property type="entry name" value="LARGE RIBOSOMAL SUBUNIT PROTEIN EL15"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11847">
        <property type="entry name" value="RIBOSOMAL PROTEIN L15"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00827">
        <property type="entry name" value="Ribosomal_L15e"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01384">
        <property type="entry name" value="Ribosomal_L15e"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54189">
        <property type="entry name" value="Ribosomal proteins S24e, L23 and L15e"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01194">
        <property type="entry name" value="RIBOSOMAL_L15E"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 5" id="KW-0687">Ribonucleoprotein</keyword>
    <keyword evidence="2 5" id="KW-0689">Ribosomal protein</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006857"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022980"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023274"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00034092"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="RuleBase" id="RU000663"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001870.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="194" mass="22568" checksum="2D4DEAD2D9D8729E" modified="2019-05-08" version="1">MGAYKYIQELWRKKQSDVMRFLLRVRCWQYRQLSALHRAPRPTRPDKARRLGYKAKQGYVIYRIRVRRGGRKRPVPKGATYGKPVNHGVNQLKFARSLQSVAEVPHPRINQVNKEEKQNEFKLVAARPEEGNTSGQLDIINVIGQQLFCLCDNNTTAWETLTKEGLKGIHLRFYPEPSCMLGQDFVRFRSTVLV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJK8</accession>
    <name>A0A452GJK8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">KRR1 small subunit processome component</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="8">KRR-R motif-containing protein 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="8">Required for 40S ribosome biogenesis. Involved in nucleolar processing of pre-18S ribosomal RNA and ribosome assembly.</text>
    </comment>
    <comment type="subunit">
        <text evidence="8">Component of the ribosomal small subunit (SSU) processome.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 8">Nucleus</location>
            <location evidence="1 8">Nucleolus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 8">Belongs to the KRR1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJK8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002113.1">
        <property type="protein sequence ID" value="ENSGAGP00000001859.1"/>
        <property type="gene ID" value="ENSGAGG00000001474.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032040">
        <property type="term" value="C:small-subunit processome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006364">
        <property type="term" value="P:rRNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd22393">
        <property type="entry name" value="KH-I_KRR1_rpt1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd22394">
        <property type="entry name" value="KH-I_KRR1_rpt2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1370.10:FF:000014">
        <property type="entry name" value="KRR1 small subunit processome component"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1370.10">
        <property type="entry name" value="K Homology domain, type 1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036612">
        <property type="entry name" value="KH_dom_type_1_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041174">
        <property type="entry name" value="KRR1-like_KH1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048550">
        <property type="entry name" value="KRR1-like_KH1_euk"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048548">
        <property type="entry name" value="KRR1-like_KH2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048549">
        <property type="entry name" value="KRR1-like_KH2_euk"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024166">
        <property type="entry name" value="rRNA_assembly_KRR1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12581">
        <property type="entry name" value="HIV-1 REV BINDING PROTEIN 2, 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12581:SF0">
        <property type="entry name" value="KRR1 SMALL SUBUNIT PROCESSOME COMPONENT HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17903">
        <property type="entry name" value="KH_KRR1_1st"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21800">
        <property type="entry name" value="KH_KRR1_2nd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF006515">
        <property type="entry name" value="KRR1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="11" id="KW-0472">Membrane</keyword>
    <keyword evidence="6 8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7 8" id="KW-0687">Ribonucleoprotein</keyword>
    <keyword evidence="3 8" id="KW-0690">Ribosome biogenesis</keyword>
    <keyword evidence="5 8" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="4 8" id="KW-0698">rRNA processing</keyword>
    <keyword evidence="11" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="306"/>
            <end position="329"/>
        </location>
    </feature>
    <feature type="domain" description="KRR1 small subunit processome component first KH" evidence="12">
        <location>
            <begin position="55"/>
            <end position="135"/>
        </location>
    </feature>
    <feature type="domain" description="KRR1 small subunit processome component second KH" evidence="13">
        <location>
            <begin position="170"/>
            <end position="199"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1"/>
            <end position="45"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="219"/>
            <end position="246"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="258"/>
            <end position="288"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="36"/>
            <end position="45"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="10">
        <location>
            <begin position="219"/>
            <end position="231"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004604"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009344"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022517"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022552"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023274"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PIRNR" id="PIRNR006515"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF17903"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="Pfam" id="PF21800"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001859.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="337" mass="39357" checksum="6A485DC3120C2658" modified="2019-05-08" version="1">MAAPKKAESGQVAPASKGRSKKKAVNESELLTVPDGWKEPAFTREDNPKGLLEESSFATLFPKYREAYLKECWPLVQKALNEHHVNATLDLIEGSITVTTTKKTFDPYMIVRARDLIKLLARSVPFEQAVRVLQDDTACDIIKIGSLVSNRERFIKRRQRLIGPKGSTLKVRKVVLDTMKNIHPIYNIKTLMIKRELSKDPELRTQSWERFLPQFKHKNLNKRKEPKKKSVKKEYTPFPPPQPENQIDKELASGEYFLKESQKKRKRMEELKAKQAESFRRRQEERNKAFIPPKEKLVVKPKKGNVCFLSLESFLLFYLKLLSFGWWLFSKPRYMEH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJN1</accession>
    <name>A0A452GJN1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">C2H2-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJN1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002156.1">
        <property type="protein sequence ID" value="ENSGAGP00000001897.1"/>
        <property type="gene ID" value="ENSGAGG00000001510.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000977">
        <property type="term" value="F:RNA polymerase II transcription regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0022603">
        <property type="term" value="P:regulation of anatomical structure morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002315">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002063">
        <property type="entry name" value="RB associated KRAB zinc finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:003000">
        <property type="entry name" value="Zinc finger and SCAN domain-containing 20"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002343">
        <property type="entry name" value="Zinc finger protein 33A"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000690">
        <property type="entry name" value="Zinc finger protein 354C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002090">
        <property type="entry name" value="Zinc finger protein 473"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050717">
        <property type="entry name" value="C2H2-ZF_Transcription_Reg"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14196">
        <property type="entry name" value="ODD-SKIPPED - RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14196:SF12">
        <property type="entry name" value="ZINC FINGER PROTEIN 208-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="9"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="9" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 11" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="64"/>
            <end position="98"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="99"/>
            <end position="126"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="127"/>
            <end position="154"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="155"/>
            <end position="182"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="183"/>
            <end position="210"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="211"/>
            <end position="238"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="239"/>
            <end position="266"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="267"/>
            <end position="294"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="295"/>
            <end position="322"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="319"/>
            <end position="343"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="319"/>
            <end position="336"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001897.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="343" mass="38933" checksum="89F8B6F1CCE59FF7" modified="2019-05-08" version="1">MCITLHFSTLNFICPGQVELQGRFLRRAKGNFSQCWEEREAWSHRHGSERKLGNSQRNKVNEVIECGGGGKDPQETTAHFNQSSNLTTHQRVHIGERPYKCLECGKSFSVLSTLIVHQRTHTGEKPYECLDCGKSFYRSSNLTTHQRVHTGEGPYKCLECGKSFSAPSVLLIHQRTHTGEKPYKCLECGKSFSRRSVLHIHQRTHTGEKPYKCLKCGKSFSQSSGLISHERIHTGERPYKCLECGKSFSHRSGLSNHGRIHTGERPYKCLECGKSFSRHSDLSNHGRIHTGEKPYKCLICEKSFNRSSNLHTHWRVHTGESGKSFRESSKHQDRNHVSASAVG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GK04</accession>
    <name>A0A452GK04_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">BHLH domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the MiT/TFE family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GK04"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002293.1">
        <property type="protein sequence ID" value="ENSGAGP00000002015.1"/>
        <property type="gene ID" value="ENSGAGG00000000269.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046983">
        <property type="term" value="F:protein dimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006959">
        <property type="term" value="P:humoral immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045893">
        <property type="term" value="P:positive regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045670">
        <property type="term" value="P:regulation of osteoclast differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd18928">
        <property type="entry name" value="bHLHzip_TFE3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.280.10:FF:000003">
        <property type="entry name" value="microphthalmia-associated transcription factor isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.280.10">
        <property type="entry name" value="Helix-loop-helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011598">
        <property type="entry name" value="bHLH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024100">
        <property type="entry name" value="bHLHzip_TFE3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036638">
        <property type="entry name" value="HLH_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021802">
        <property type="entry name" value="MiT/TFE_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031867">
        <property type="entry name" value="MiT/TFE_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45776">
        <property type="entry name" value="MIP04163P"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45776:SF3">
        <property type="entry name" value="TRANSCRIPTION FACTOR E3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11851">
        <property type="entry name" value="DUF3371"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00010">
        <property type="entry name" value="HLH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15951">
        <property type="entry name" value="MITF_TFEB_C_3_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00353">
        <property type="entry name" value="HLH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47459">
        <property type="entry name" value="HLH, helix-loop-helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50888">
        <property type="entry name" value="BHLH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0010">Activator</keyword>
    <keyword evidence="9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0804">Transcription</keyword>
    <keyword evidence="4" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="BHLH" evidence="11">
        <location>
            <begin position="232"/>
            <end position="285"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="12"/>
            <end position="93"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="338"/>
            <end position="361"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="396"/>
            <end position="441"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="285"/>
            <end position="319"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="10">
        <location>
            <begin position="20"/>
            <end position="41"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="405"/>
            <end position="441"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00008289"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50888"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000002015.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="441" mass="46818" checksum="2DB68EC1F0411425" modified="2019-05-08" version="1">MAGAGFCTSPLMSLTLSVSPSPPAAPLQPPAPPAPGDPPGPSMSSSRVLLRQQLMRAQTQEQERREQQQAAQFLPPNSMPPTPAISVSSPLPRPSAQVPVEVLKVRAWGTCGEWGLSPLGGTGSHLGARGATPLIPSGSTPWLSPGSIDDVIDEIISLESSYNDEMLNYMSVDGGLQLPSTLPVAGNLLEIYSSPGLTDPTVPVSNSCPADLPNIKTELSENEAKALMKERQKKDNHNLIERRRRFNINDRIKELGTLIPKSNDPEMRWNKGTILKASVDYIRKLQKEQQRSKEMELRQRKLEQANRSLQLRVQELEMQAQLHGLPLTSSAGLLPLGCSGEGAKPEGPGPPPFRGGPSCAPAPTSSAACLLDLAFPSEELGEAGLGFPLGLGVDGGLEDILMEDGGPLSPLGASDPLLSSLSPGASKGSSRRSSFSMEEDS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMK3</accession>
    <name>A0A452HMK3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4 8">S-formylglutathione hydrolase</fullName>
            <ecNumber evidence="3 8">3.1.2.12</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1 8">Serine hydrolase involved in the detoxification of formaldehyde.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="8">
            <text>S-formylglutathione + H2O = formate + glutathione + H(+)</text>
            <dbReference type="Rhea" id="RHEA:14961"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:15740"/>
            <dbReference type="ChEBI" id="CHEBI:57688"/>
            <dbReference type="ChEBI" id="CHEBI:57925"/>
            <dbReference type="EC" id="3.1.2.12"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="8">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 8">Belongs to the esterase D family.</text>
    </comment>
    <dbReference type="EC" id="3.1.2.12" evidence="3 8"/>
    <dbReference type="AlphaFoldDB" id="A0A452HMK3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018471.1">
        <property type="protein sequence ID" value="ENSGAGP00000016177.1"/>
        <property type="gene ID" value="ENSGAGG00000012141.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000018473.1">
        <property type="protein sequence ID" value="ENSGAGP00000016179.1"/>
        <property type="gene ID" value="ENSGAGG00000012141.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0052689">
        <property type="term" value="F:carboxylic ester hydrolase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0018738">
        <property type="term" value="F:S-formylglutathione hydrolase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046294">
        <property type="term" value="P:formaldehyde catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1820:FF:000002">
        <property type="entry name" value="S-formylglutathione hydrolase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1820">
        <property type="entry name" value="alpha/beta hydrolase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029058">
        <property type="entry name" value="AB_hydrolase_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000801">
        <property type="entry name" value="Esterase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014186">
        <property type="entry name" value="S-formylglutathione_hydrol"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR02821">
        <property type="entry name" value="fghA_ester_D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10061">
        <property type="entry name" value="S-FORMYLGLUTATHIONE HYDROLASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10061:SF0">
        <property type="entry name" value="S-FORMYLGLUTATHIONE HYDROLASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00756">
        <property type="entry name" value="Esterase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53474">
        <property type="entry name" value="alpha/beta-Hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6 8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 8" id="KW-0719">Serine esterase</keyword>
    <feature type="active site" description="Charge relay system" evidence="7">
        <location>
            <position position="149"/>
        </location>
    </feature>
    <feature type="active site" description="Charge relay system" evidence="7">
        <location>
            <position position="226"/>
        </location>
    </feature>
    <feature type="active site" description="Charge relay system" evidence="7">
        <location>
            <position position="260"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00002608"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005622"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00016774"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022487"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PIRSR" id="PIRSR614186-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="RuleBase" id="RU363068"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016179.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="282" mass="31673" checksum="810A2643BB058338" modified="2019-05-08" version="1">MALKQISSNKCFEGFQKVFEHDSIELKCKMRFGIYLPPKAEIGKCPVLYWLSGLTCTEQNFITKSGFHQAAAEHGLIVVAPDTSPRGCNIEGEEESWDFGTGAGFYVDATEEPWKTNYRMYSYIKDELPKLIKANFPADSERMSIFGHSMGGHGALILALKNPGKYKSVSAFAPICNPVQCPWGKKAFGGYLGSDLTKWEAYDATNLVKSYPDSHLDILIDQGKDDQFLSAGQLLPDNFIAACTERKIPVVFRLQQGYDHSYFFITTFINDHIRHHAKYLNA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMK4</accession>
    <name>A0A452HMK4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Calponin-homology (CH) domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HMK4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018464.1">
        <property type="protein sequence ID" value="ENSGAGP00000016171.1"/>
        <property type="gene ID" value="ENSGAGG00000012125.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd21270">
        <property type="entry name" value="CH_LRCH1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.418.10:FF:000021">
        <property type="entry name" value="Leucine-rich repeat and calponin homology domain-containing protein 1 isoform 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000007">
        <property type="entry name" value="Leucine-rich repeat and calponin homology domain-containing protein 1 isoform 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.418.10">
        <property type="entry name" value="Calponin-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001715">
        <property type="entry name" value="CH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036872">
        <property type="entry name" value="CH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001611">
        <property type="entry name" value="Leu-rich_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003591">
        <property type="entry name" value="Leu-rich_rpt_typical-subtyp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050216">
        <property type="entry name" value="LRR_domain-containing"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48051">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48051:SF38">
        <property type="entry name" value="LEUCINE RICH REPEATS AND CALPONIN HOMOLOGY DOMAIN CONTAINING 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00307">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13855">
        <property type="entry name" value="LRR_8"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00033">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00364">
        <property type="entry name" value="LRR_BAC"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00369">
        <property type="entry name" value="LRR_TYP"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47576">
        <property type="entry name" value="Calponin-homology domain, CH-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52058">
        <property type="entry name" value="L domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50021">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51450">
        <property type="entry name" value="LRR"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0433">Leucine-rich repeat</keyword>
    <keyword evidence="4" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="4">
        <location>
            <begin position="555"/>
            <end position="573"/>
        </location>
    </feature>
    <feature type="domain" description="Calponin-homology (CH)" evidence="5">
        <location>
            <begin position="428"/>
            <end position="544"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="257"/>
            <end position="289"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022614"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016171.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="580" mass="65344" checksum="E66C2826A5470624" modified="2019-05-08" version="1">PSGSLASWGEEDGYLGQGGGGCSRAQGERGCRHLGKGGNVLFLFLSSDLSKNRLTEVPMELCHFVSLETLNLYHNCIKIIPDAIVNLQMLTYLNLSRNQLSSLPACLCGLPLKVLIASNNRLGSLPEEIGQLKQLMELDVSCNEITALPQQIGQLKSLRELNIRRNYLKVLPQELIELPLVKFDFSCNKVLVIPICFRKMLQLQVLLLENNPLQSPPAQICIKGKVHIFKYLSIQACQIKTADSLYLNAIERPHLQQPVEESKKDSDSGVGSDNGDKRLSATEPSDEDTVCLNVPMSNIMEEDQIIKEDSGHHINLVQGLNVKKRTLTVTGSLPTILFVMLYECNSQKTSQPKFLLTTWLKPTPSVYHYPPLTLPPISFNKLTQAQTWDNYSYSIPSEGDSDNVFLRPQRNLESIDPQFTIRRKMEQMREERELVEQLRENIEMRLKICLPEDLGSALMDGVVLCHLVNHIRPRSVGSIHVPSPAVPKLSMAKCRRNVENFLDACRKLGIPEADLCSPYDILQSDIRHIRKTVDTLLALGEKTPQSTSTFRSWDLIGFCLFHILFIVLMYITYHWNALTA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMK8</accession>
    <name>A0A452HMK8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Slit homolog 2 protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
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    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
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        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
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    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
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        <scope>IDENTIFICATION</scope>
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    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
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                <consortium name="Ensembl"/>
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    <comment type="caution">
        <text evidence="15">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
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        </source>
    </evidence>
    <sequence length="1336" mass="147988" checksum="DB70599D870E97D5" modified="2019-05-08" version="1">MPKLRTFRLHSNNLYCDCHLAWLSDWLRQRPRVGLYTQCMGPSHLRGHNVAEVQKREFVCSDEEEGHQSFMAPSCSVLHCPTACTCSNNIVDCRGKGLTEIPTNLPETITEIRLEQNSIKVIPPGAFSPYKKLRRIDLSNNQISETAPDAFQGLRSLNSLVLYGNKITELPKGLFEGLFSLQLLLLNANKINCLRVDAFQDLHNLNLLSLYDNKLQTIAKGTFSPLRSIQTLHLAQNPFICDCHLKWLADYLHANPIETSGARCTSPRRLANKRIGQIKSKKFRCSAKEQYFIPGTEDYRSKLSGDCFADLACPEKCRCEGTTVDCSNQKLNKIPEHVPQYTAELRLNNNEFTVLEATGIFKKLPQLRKINLSNNKITDIEEGAFEGASGVNELLLTSNRLETIRHKMFKGLESLKTLMLRSNRISCIGNDSFTGLSSVRLLSLYDNQITTVAPGAFDTLHSLSTLNLLANPFNCNCHLAWLGDWLRKKRIVTGNPRCQKPYFLKEIPIQDVAIQDFTCDDGNDDNSCSPLSRCPAECTCLDTVVRCSNKGLKVLPKGIPKDVTELYLDGNQFTLVPKELSNYKHLTLIDLSNNRISTLSNQSFSNMTQLLTLILSYNRLRCIPARTFDGLKSLRLLSLHGNDISVVPEGAFNDLSALSHLAIGANPLYCDCNMQWLSDWVKSEYKEPGIARCAGPGEMADKLLLTTPSKKFTCQGPVDVNILAKCSPCLSNPCKNDGTCNNDPVDFYTCTCPYGFKGQDCDIPIHACISNPCQHGGTCHLKEGEKDGFWCTCADGFEGENCEVNVDDCEDNDCENNSTCVDGINNYTCLCPPEYTGELCEEKLDFCAQDLNPCQHDSKCILTLKGYKCDCTPGYVGEHCDMDYDDCQDNKCKNGAQCTDAVNGYTCICPEGYSGLFCEFSPPVVLPRTSPCDNYECQNGAQCIVRVNEPICQCLSGYQGDKCEKLVSVNFVSKESYLQIPSSKVRPQTNITLQIATDEDSGILLYKGDRDHIAMELYRGRVRVSYDTGSYPASAIYSVETINDGNFHIVELLAMDQILSLSVDGGSPKIITNLSKQSTLNVDSPLYVGGMPVKNNVAALRQSPGQNGSSFHGCIRNLYINSELQDFRNVPLQVGILPGCEPCHKKVCVHGTCHAMSQSGFTCECEGGWTGPLCDQQTNDPCLGNKCVHGTCLPINAFSYSCKCLQGHGGVLCDEEEELFNPCQSIKCKHGKCRLSGLGKPYCECSSGYTGDSCDKEISCRGERIRDYYQKQQGYAACQTTKKVSRLECKGGCSTGQCCGPLRSKRRKYSFECTDGSSFVDEVEKVVKCGCTKCPS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HML1</accession>
    <name>A0A452HML1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Slit homolog 2 protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="15">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018477.1">
        <property type="protein sequence ID" value="ENSGAGP00000016183.1"/>
        <property type="gene ID" value="ENSGAGG00000011388.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048495">
        <property type="term" value="F:Roundabout binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072359">
        <property type="term" value="P:circulatory system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060560">
        <property type="term" value="P:developmental growth involved in morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060562">
        <property type="term" value="P:epithelial tube morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001822">
        <property type="term" value="P:kidney development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050919">
        <property type="term" value="P:negative chemotaxis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030308">
        <property type="term" value="P:negative regulation of cell growth"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010605">
        <property type="term" value="P:negative regulation of macromolecule metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010977">
        <property type="term" value="P:negative regulation of neuron projection development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0021772">
        <property type="term" value="P:olfactory bulb development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050770">
        <property type="term" value="P:regulation of axonogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032535">
        <property type="term" value="P:regulation of cellular component size"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050767">
        <property type="term" value="P:regulation of neurogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031290">
        <property type="term" value="P:retinal ganglion cell axon guidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0022029">
        <property type="term" value="P:telencephalon cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd00054">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="CDD" id="cd00110">
        <property type="entry name" value="LamG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000045">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000053">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000054">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000062">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000063">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000099">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000289">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.200:FF:000013">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000002">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000004">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.200">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000483">
        <property type="entry name" value="Cys-rich_flank_reg_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006207">
        <property type="entry name" value="Cys_knot_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001881">
        <property type="entry name" value="EGF-like_Ca-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000152">
        <property type="entry name" value="EGF-type_Asp/Asn_hydroxyl_site"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018097">
        <property type="entry name" value="EGF_Ca-bd_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003645">
        <property type="entry name" value="Fol_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001791">
        <property type="entry name" value="Laminin_G"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001611">
        <property type="entry name" value="Leu-rich_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003591">
        <property type="entry name" value="Leu-rich_rpt_typical-subtyp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000372">
        <property type="entry name" value="LRRNT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051355">
        <property type="entry name" value="Notch/Slit_guidance"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45836">
        <property type="entry name" value="SLIT HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45836:SF2">
        <property type="entry name" value="SLIT HOMOLOG 2 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00008">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00054">
        <property type="entry name" value="Laminin_G_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00560">
        <property type="entry name" value="LRR_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13855">
        <property type="entry name" value="LRR_8"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01463">
        <property type="entry name" value="LRRCT"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01462">
        <property type="entry name" value="LRRNT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00041">
        <property type="entry name" value="CT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="SMART" id="SM00179">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SMART" id="SM00274">
        <property type="entry name" value="FOLN"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00282">
        <property type="entry name" value="LamG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00368">
        <property type="entry name" value="LRR_RI"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SMART" id="SM00365">
        <property type="entry name" value="LRR_SD22"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SMART" id="SM00369">
        <property type="entry name" value="LRR_TYP"/>
        <property type="match status" value="12"/>
    </dbReference>
    <dbReference type="SMART" id="SM00082">
        <property type="entry name" value="LRRCT"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00013">
        <property type="entry name" value="LRRNT"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52058">
        <property type="entry name" value="L domain-like"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00010">
        <property type="entry name" value="ASX_HYDROXYL"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01185">
        <property type="entry name" value="CTCK_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01225">
        <property type="entry name" value="CTCK_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01186">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01187">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50025">
        <property type="entry name" value="LAM_G_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51450">
        <property type="entry name" value="LRR"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0145">Chemotaxis</keyword>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="10" id="KW-0221">Differentiation</keyword>
    <keyword evidence="12 15" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5 15" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7" id="KW-0358">Heparin-binding</keyword>
    <keyword evidence="6" id="KW-0433">Leucine-rich repeat</keyword>
    <keyword evidence="11" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="8" id="KW-0732">Signal</keyword>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="719"/>
            <end position="754"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="756"/>
            <end position="795"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="797"/>
            <end position="833"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="844"/>
            <end position="882"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="884"/>
            <end position="920"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="929"/>
            <end position="965"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin G" evidence="17">
        <location>
            <begin position="968"/>
            <end position="1141"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="1145"/>
            <end position="1176"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="1179"/>
            <end position="1215"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="1220"/>
            <end position="1256"/>
        </location>
    </feature>
    <feature type="domain" description="CTCK" evidence="16">
        <location>
            <begin position="1261"/>
            <end position="1336"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="744"/>
            <end position="753"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="785"/>
            <end position="794"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="872"/>
            <end position="881"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="910"/>
            <end position="919"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="955"/>
            <end position="964"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="1166"/>
            <end position="1175"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="1183"/>
            <end position="1193"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="1205"/>
            <end position="1214"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="1224"/>
            <end position="1234"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="1246"/>
            <end position="1255"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022500"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022614"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022674"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00074955"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS01225"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50025"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016183.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1337" mass="147980" checksum="705CF3C450A923B2" modified="2019-05-08" version="1">MPKLRTFRLHSNNLYCDCHLAWLSDWLRQRPRVGLYTQCMGPSHLRGHNVAEVQKREFVCSDEEEGHQSFMAPSCSVLHCPTACTCSNNIVDCRGKGLTEIPTNLPETITEIRLEQNSIKVIPPGAFSPYKKLRRIDLSNNQISETAPDAFQGLRSLNSLVLYGNKITELPKGLFEGLFSLQLLLLNANKINCLRVDAFQDLHNLNLLSLYDNKLQTIAKGTFSPLRSIQTLHLAQNPFICDCHLKWLADYLHANPIETSGARCTSPRRLANKRIGQIKSKKFRCSGTEDYRSKLSGDCFADLACPEKCRCEGTTVDCSNQKLNKIPEHVPQYTAELRLNNNEFTVLEATGIFKKLPQLRKINLSNNKITDIEEGAFEGASGVNELLLTSNRLETIRHKMFKGLESLKTLMLRSNRISCIGNDSFTGLSSVRLLSLYDNQITTVAPGAFDTLHSLSTLNLLANPFNCNCHLAWLGDWLRKKRIVTGNPRCQKPYFLKEIPIQDVAIQDFTCDDGNDDNSCSPLSRCPAECTCLDTVVRCSNKGLKVLPKGIPKDVTELYLDGNQFTLVPKELSNYKHLTLIDLSNNRISTLSNQSFSNMTQLLTLILSYNRLRCIPARTFDGLKSLRLLSLHGNDISVVPEGAFNDLSALSHLAIGANPLYCDCNMQWLSDWVKSEYKEPGIARCAGPGEMADKLLLTTPSKKFTCQGPVDVNILAKCSPCLSNPCKNDGTCNNDPVDFYTCTCPYGFKGQDCDIPIHACISNPCQHGGTCHLKEGEKDGFWCTCADGFEGENCEVNVDDCEDNDCENNSTCVDGINNYTCLCPPEYTAANLNEVEKGELCEEKLDFCAQDLNPCQHDSKCILTLKGYKCDCTPGYVGEHCDMDYDDCQDNKCKNGAQCTDAVNGYTCICPEGYSGLFCEFSPPVVLPRTSPCDNYECQNGAQCIVRVNEPICQCLSGYQGDKCEKLVSVNFVSKESYLQIPSSKVRPQTNITLQIATDEDSGILLYKGDRDHIAMELYRGRVRVSYDTGSYPASAIYSVETINDGNFHIVELLAMDQILSLSVDGGSPKIITNLSKQSTLNVDSPLYVGGMPVKNNVAALRQSPGQNGSSFHGCIRNLYINSELQDFRNVPLQVGILPGCEPCHKKVCVHGTCHAMSQSGFTCECEGGWTGPLCDQQTNDPCLGNKCVHGTCLPINAFSYSCKCLQGHGGVLCDEEEELFNPCQSIKCKHGKCRLSGLGKPYCECSSGYTGDSCDKEISCRGERIRDYYQKQQGYAACQTTKKVSRLECKGGCSTGQCCGPLRSKRRKYSFECTDGSSFVDEVEKVVKCGCTKCPS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HML6</accession>
    <name>A0A452HML6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">BAR domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="8">Involved in cytokinesis and septation where it has a role in the localization of F-actin.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HML6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018482.1">
        <property type="protein sequence ID" value="ENSGAGP00000016188.1"/>
        <property type="gene ID" value="ENSGAGG00000012148.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015629">
        <property type="term" value="C:actin cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008289">
        <property type="term" value="F:lipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051666">
        <property type="term" value="P:actin cortical patch localization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048589">
        <property type="term" value="P:developmental growth"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006897">
        <property type="term" value="P:endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097320">
        <property type="term" value="P:plasma membrane tubulation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07590">
        <property type="entry name" value="BAR_Bin3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1270.60:FF:000028">
        <property type="entry name" value="Bridging integrator 3 homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.60">
        <property type="entry name" value="Arfaptin homology (AH) domain/BAR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027267">
        <property type="entry name" value="AH/BAR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004148">
        <property type="entry name" value="BAR_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046982">
        <property type="entry name" value="BIN3/RVS161-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037428">
        <property type="entry name" value="Bin3_BAR"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47174">
        <property type="entry name" value="BRIDGING INTEGRATOR 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47174:SF3">
        <property type="entry name" value="BRIDGING INTEGRATOR 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03114">
        <property type="entry name" value="BAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00721">
        <property type="entry name" value="BAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103657">
        <property type="entry name" value="BAR/IMD domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51021">
        <property type="entry name" value="BAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="3" id="KW-0132">Cell division</keyword>
    <keyword evidence="4 9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0717">Septation</keyword>
    <feature type="domain" description="BAR" evidence="10">
        <location>
            <begin position="9"/>
            <end position="232"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="24"/>
            <end position="51"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="121"/>
            <end position="148"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023210"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00059510"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS51021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016188.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="253" mass="29521" checksum="F6EDE9925F9D1835" modified="2019-05-08" version="1">MSWIPFKIGQPKKQIVPKTVERDFEREYGKLQQLEDQTKKLQKDMKKSTDADVAMSKSAVKISSDLLSNPLCEQDPNFLAMVTALDTAMKRMDSFNQEKVNQIQKTVIEPLKKFSSVFPSLNMAVKRREQTLQDYKRLQSKVEKYEEKEKTGPILAKLHQAREELRPVKEDFEAKNKQLLEEMPKFYSSRIDYFEPSFESLIRAQVVYYTEMHKIFGDLTKQIDEPGLTDEQREKENEAKLSELRALSIVADD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMM5</accession>
    <name>A0A452HMM5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Slit homolog 2 protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="15">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018485.1">
        <property type="protein sequence ID" value="ENSGAGP00000016191.1"/>
        <property type="gene ID" value="ENSGAGG00000011388.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048495">
        <property type="term" value="F:Roundabout binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072359">
        <property type="term" value="P:circulatory system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060560">
        <property type="term" value="P:developmental growth involved in morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060562">
        <property type="term" value="P:epithelial tube morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001822">
        <property type="term" value="P:kidney development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050919">
        <property type="term" value="P:negative chemotaxis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030308">
        <property type="term" value="P:negative regulation of cell growth"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010605">
        <property type="term" value="P:negative regulation of macromolecule metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010977">
        <property type="term" value="P:negative regulation of neuron projection development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0021772">
        <property type="term" value="P:olfactory bulb development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050770">
        <property type="term" value="P:regulation of axonogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032535">
        <property type="term" value="P:regulation of cellular component size"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050767">
        <property type="term" value="P:regulation of neurogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031290">
        <property type="term" value="P:retinal ganglion cell axon guidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0022029">
        <property type="term" value="P:telencephalon cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd00054">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="CDD" id="cd00110">
        <property type="entry name" value="LamG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000045">
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        <property type="entry name" value="Ribonuclease Inhibitor"/>
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        <property type="entry name" value="ConA-like_dom_sf"/>
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        <property type="entry name" value="Cys-rich_flank_reg_C"/>
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        <property type="entry name" value="Cys_knot_C"/>
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        <property type="entry name" value="LamG"/>
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        <property type="match status" value="5"/>
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    </dbReference>
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    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
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    </dbReference>
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        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52058">
        <property type="entry name" value="L domain-like"/>
        <property type="match status" value="3"/>
    </dbReference>
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        <property type="entry name" value="ASX_HYDROXYL"/>
        <property type="match status" value="2"/>
    </dbReference>
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    </dbReference>
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        <property type="entry name" value="LRR"/>
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    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0145">Chemotaxis</keyword>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="10" id="KW-0221">Differentiation</keyword>
    <keyword evidence="12 15" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5 15" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7" id="KW-0358">Heparin-binding</keyword>
    <keyword evidence="6" id="KW-0433">Leucine-rich repeat</keyword>
    <keyword evidence="11" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="8" id="KW-0732">Signal</keyword>
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        <location>
            <begin position="703"/>
            <end position="738"/>
        </location>
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            <begin position="952"/>
            <end position="1125"/>
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            <begin position="1204"/>
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            <begin position="1245"/>
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    <sequence length="1321" mass="146278" checksum="DEE33A568F061005" modified="2019-05-08" version="1">MPKLRTFRLHSNNLYCDCHLAWLSDWLRQRPRVGLYTQCMGPSHLRGHNVAEVQKREFVCSDEEEGHQSFMAPSCSVLHCPTACTCSNNIVDCRGKGLTEIPTNLPETITEIRLEQNSIKVIPPGAFSPYKKLRRIDLSNNQISETAPDAFQGLRSLNSLVLYGNKITELPKGLFEGLFSLQLLLLNANKINCLRVDAFQDLHNLNLLSLYDNKLQTIAKGTFSPLRSIQTLHLAQNPFICDCHLKWLADYLHANPIETSGARCTSPRRLANKRIGQIKSKKFRCSAKEQYFIPGTEDYRSKLSGDCFADLACPEKCRCEGTTVDCSNQKLNKIPEHVPQYTAELRLNNNEFTVLEATGIFKKLPQLRKINLSNNKITDIEEGAFEGASGVNELLLTSNRLETIRHKMFKGLESLKTLMLRSNRISCIGNDSFTGLSSVRLLSLYDNQITTVAPGAFDTLHSLSTLNLLANPFNCNCHLAWLGDWLRKKRIVTGNPRCQKPYFLKEIPIQDVAIQDFTCDDGNDDNSCSPLSRCPAECTCLDTVVRCSNKGLKVLPKGIPKDVTELYLDGNQFTLVPKELSNYKHLTLIILSYNRLRCIPARTFDGLKSLRLLSLHGNDISVVPEGAFNDLSALSHLAIGANPLYCDCNMQWLSDWVKSEYKEPGIARCAGPGEMADKLLLTTPSKKFTCQGPVDVNILAKCSPCLSNPCKNDGTCNNDPVDFYTCTCPYGFKGQDCDIPIHACISNPCQHGGTCHLKEGEKDGFWCTCADGFEGENCEVNVDDCEDNDCENNSTCVDGINNYTCLCPPEYTAANLNEVEKGELCEEKLDFCAQDLNPCQHDSKCILTLKGYKCDCTPGYVGEHCDMDYDDCQDNKCKNGAQCTDAVNGYTCICPEGYSGLFCEFSPPVVLPRTSPCDNYECQNGAQCIVRVNEPICQCLSGYQGDKCEKLVSVNFVSKESYLQIPSSKVRPQTNITLQIATDEDSGILLYKGDRDHIAMELYRGRVRVSYDTGSYPASAIYSVETINDGNFHIVELLAMDQILSLSVDGGSPKIITNLSKQSTLNVDSPLYVGGMPVKNNVAALRQSPGQNGSSFHGCIRNLYINSELQDFRNVPLQVGILPGCEPCHKKVCVHGTCHAMSQSGFTCECEGGWTGPLCDQQTNDPCLGNKCVHGTCLPINAFSYSCKCLQGHGGVLCDEEEELFNPCQSIKCKHGKCRLSGLGKPYCECSSGYTGDSCDKEISCRGERIRDYYQKQQGYAACQTTKKVSRLECKGGCSTGQCCGPLRSKRRKYSFECTDGSSFVDEVEKVVKCGCTKCPS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMM7</accession>
    <name>A0A452HMM7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Fibroblast growth factor</fullName>
            <shortName evidence="9">FGF</shortName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 9">Belongs to the heparin-binding growth factors family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMM7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018503.1">
        <property type="protein sequence ID" value="ENSGAGP00000016207.1"/>
        <property type="gene ID" value="ENSGAGG00000012157.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008083">
        <property type="term" value="F:growth factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051781">
        <property type="term" value="P:positive regulation of cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd23318">
        <property type="entry name" value="beta-trefoil_FGF6"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Fibroblast growth factor"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002209">
        <property type="entry name" value="Fibroblast_GF_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008996">
        <property type="entry name" value="IL1/FGF"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11486">
        <property type="entry name" value="FIBROBLAST GROWTH FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00167">
        <property type="entry name" value="FGF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00263">
        <property type="entry name" value="HBGFFGF"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00262">
        <property type="entry name" value="IL1HBGF"/>
    </dbReference>
    <dbReference type="SMART" id="SM00442">
        <property type="entry name" value="FGF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50353">
        <property type="entry name" value="Cytokine"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="6" id="KW-0221">Differentiation</keyword>
    <keyword evidence="7" id="KW-0339">Growth factor</keyword>
    <keyword evidence="8" id="KW-0497">Mitogen</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="5 9" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="9">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5018811308" description="Fibroblast growth factor" evidence="9">
        <location>
            <begin position="23"/>
            <end position="189"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007936"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="RuleBase" id="RU049442"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016207.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="189" mass="21041" checksum="FF677384CE0FBB7C" modified="2019-05-08" version="1">MLVHWTLPAFVLLGFLAGIVSSYPVVSRTNGTLLERGWESLLSRSIAGMSGEKSDVNWESDYLLGIKRQRRLYCNVGIGFHLQILPDGRISGVHNENQYSLLEISTVERGVVSLFGVKSALFVAMNNKGRLYGTVNECKFRETLLPNNYNAYESSAYQGAYIALSKHGRVKKGNKVSPAMTVTHFLPRI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMN0</accession>
    <name>A0A452HMN0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Rho-related BTB domain-containing protein 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="7">Regulator of cell proliferation and apoptosis. It likely functions as a substrate-adapter that recruits key substrates, e.g. MSI2, to CUL3-based ubiquitin ligase complexes for degradation. Required for MSI2 ubiquitination and degradation.</text>
    </comment>
    <comment type="subunit">
        <text evidence="8">Interacts with HSP90AA1 and HSP90AB1. Forms a complex with CUL3 and RBX1. Interacts (via BTB 1 domain) with CUL3. Interacts with MSI2.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMN0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018508.1">
        <property type="protein sequence ID" value="ENSGAGP00000016212.1"/>
        <property type="gene ID" value="ENSGAGG00000012159.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010008">
        <property type="term" value="C:endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005525">
        <property type="term" value="F:GTP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003924">
        <property type="term" value="F:GTPase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007264">
        <property type="term" value="P:small GTPase-mediated signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd18531">
        <property type="entry name" value="BACK_RHOBTB2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18356">
        <property type="entry name" value="BTB1_POZ_RHOBTB2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18359">
        <property type="entry name" value="BTB2_POZ_RHOBTB2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd01873">
        <property type="entry name" value="RhoBTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.710.10:FF:000050">
        <property type="entry name" value="Rho related BTB domain containing 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.710.10:FF:000179">
        <property type="entry name" value="Rho-related BTB domain-containing protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000177">
        <property type="entry name" value="Rho-related BTB domain-containing protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.710.10:FF:000014">
        <property type="entry name" value="Rho-related BTB domain-containing protein 2 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.710.10">
        <property type="entry name" value="Potassium Channel Kv1.1, Chain A"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000210">
        <property type="entry name" value="BTB/POZ_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011333">
        <property type="entry name" value="SKP1/BTB/POZ_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001806">
        <property type="entry name" value="Small_GTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003578">
        <property type="entry name" value="Small_GTPase_Rho"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24072">
        <property type="entry name" value="RHO FAMILY GTPASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00651">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00071">
        <property type="entry name" value="Ras"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00449">
        <property type="entry name" value="RASTRNSFRMNG"/>
    </dbReference>
    <dbReference type="SMART" id="SM00225">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00175">
        <property type="entry name" value="RAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00173">
        <property type="entry name" value="RAS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00174">
        <property type="entry name" value="RHO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54695">
        <property type="entry name" value="POZ domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50097">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51419">
        <property type="entry name" value="RAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51421">
        <property type="entry name" value="RAS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51420">
        <property type="entry name" value="RHO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0342">GTP-binding</keyword>
    <keyword evidence="2" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0832">Ubl conjugation</keyword>
    <keyword evidence="3" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="BTB" evidence="10">
        <location>
            <begin position="266"/>
            <end position="445"/>
        </location>
    </feature>
    <feature type="domain" description="BTB" evidence="10">
        <location>
            <begin position="503"/>
            <end position="570"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="307"/>
            <end position="332"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="354"/>
            <end position="376"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="709"/>
            <end position="739"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="354"/>
            <end position="368"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00074749"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00093283"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00093525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50097"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016212.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="739" mass="84162" checksum="BE5E7F7AD91196E6" modified="2019-05-08" version="1">MDSDMDYERPNVETIKCVVVGDNAVGKTRLICARACNATLTQYQLLATHVPTVWAIDQYRVCQEVLERSRDVVDDVSVSLRLWDTFGDHHKDRRFAYGRSDVVVLCFSIANPNSLHHVKTMWYPEIKHFCPRAPVILVGCQLDLRYADLEAVNRARRPLARPIKPNEILPPEKGREVAKELGIPYYETSVVAQFGIKDVFDNAIRAALISRRHLQFWKSHLRNVQRPLLQAPFLPPKPPPPIIMVPDPPSNNEEHPAHLLEDPLCADVILVLQEKIKIYAHKIYLSTSSSKFYDLFLMDLSEEDQQGAAGGSSRVAVSTTATERTLHQEERHHGRDFLLRAASFDICESTEEFSSSQRPPCLRPSTSDGILRGNSYEKGQRGLRRGRILSSWSRAFVSIQEEMADDPLTYKSRLMVVVKMDPSIQPGPFRAVLKYLYTGELDENERDLMHIAHIAELLEVFDLRMMVANILNNEAFMNQEITKAFHVRRTNRVKECLAKGTFSDVTFILDDGAISAHKPLLISSCDWMAAMFGGPFVESSTQEVVFPYTSKSCMRAVLEYLYTGQFSSSPDLDDMKLIILANRLCLPHLVALTEQYMVAGLMEASQMLVDIDGDVLVFLELAQFHCAYQLADWCLHHICTNYNNVCRKFPRDMKAMSAENQEYFEKHRWPPVWYLKEEDHYQRAKKEREKEDYLHQKRQPKRRWLFWNTSSSPSASPSSSAATASSSSSSSSSSSSAVV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMN2</accession>
    <name>A0A452HMN2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Rho-related BTB domain-containing protein 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="7">Regulator of cell proliferation and apoptosis. It likely functions as a substrate-adapter that recruits key substrates, e.g. MSI2, to CUL3-based ubiquitin ligase complexes for degradation. Required for MSI2 ubiquitination and degradation.</text>
    </comment>
    <comment type="subunit">
        <text evidence="8">Interacts with HSP90AA1 and HSP90AB1. Forms a complex with CUL3 and RBX1. Interacts (via BTB 1 domain) with CUL3. Interacts with MSI2.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMN2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018505.1">
        <property type="protein sequence ID" value="ENSGAGP00000016209.1"/>
        <property type="gene ID" value="ENSGAGG00000012159.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010008">
        <property type="term" value="C:endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005525">
        <property type="term" value="F:GTP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003924">
        <property type="term" value="F:GTPase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007264">
        <property type="term" value="P:small GTPase-mediated signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd18531">
        <property type="entry name" value="BACK_RHOBTB2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18356">
        <property type="entry name" value="BTB1_POZ_RHOBTB2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18359">
        <property type="entry name" value="BTB2_POZ_RHOBTB2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd01873">
        <property type="entry name" value="RhoBTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.710.10:FF:000050">
        <property type="entry name" value="Rho related BTB domain containing 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.710.10:FF:000179">
        <property type="entry name" value="Rho-related BTB domain-containing protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000177">
        <property type="entry name" value="Rho-related BTB domain-containing protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.710.10:FF:000014">
        <property type="entry name" value="Rho-related BTB domain-containing protein 2 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.710.10">
        <property type="entry name" value="Potassium Channel Kv1.1, Chain A"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000210">
        <property type="entry name" value="BTB/POZ_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011333">
        <property type="entry name" value="SKP1/BTB/POZ_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001806">
        <property type="entry name" value="Small_GTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003578">
        <property type="entry name" value="Small_GTPase_Rho"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24072">
        <property type="entry name" value="RHO FAMILY GTPASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00651">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00071">
        <property type="entry name" value="Ras"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00449">
        <property type="entry name" value="RASTRNSFRMNG"/>
    </dbReference>
    <dbReference type="SMART" id="SM00225">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00175">
        <property type="entry name" value="RAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00173">
        <property type="entry name" value="RAS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00174">
        <property type="entry name" value="RHO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54695">
        <property type="entry name" value="POZ domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50097">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51419">
        <property type="entry name" value="RAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51421">
        <property type="entry name" value="RAS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51420">
        <property type="entry name" value="RHO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0342">GTP-binding</keyword>
    <keyword evidence="2" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0832">Ubl conjugation</keyword>
    <keyword evidence="3" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="BTB" evidence="10">
        <location>
            <begin position="280"/>
            <end position="459"/>
        </location>
    </feature>
    <feature type="domain" description="BTB" evidence="10">
        <location>
            <begin position="517"/>
            <end position="584"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="321"/>
            <end position="346"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="368"/>
            <end position="390"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="723"/>
            <end position="753"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="368"/>
            <end position="382"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00074749"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00093283"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00093525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50097"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016209.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="753" mass="85870" checksum="DA3B162B359E322D" modified="2019-05-08" version="1">MEREIPVLRVRSQLMDSDMDYERPNVETIKCVVVGDNAVGKTRLICARACNATLTQYQLLATHVPTVWAIDQYRVCQEVLERSRDVVDDVSVSLRLWDTFGDHHKDRRFAYGRSDVVVLCFSIANPNSLHHVKTMWYPEIKHFCPRAPVILVGCQLDLRYADLEAVNRARRPLARPIKPNEILPPEKGREVAKELGIPYYETSVVAQFGIKDVFDNAIRAALISRRHLQFWKSHLRNVQRPLLQAPFLPPKPPPPIIMVPDPPSNNEEHPAHLLEDPLCADVILVLQEKIKIYAHKIYLSTSSSKFYDLFLMDLSEEDQQGAAGGSSRVAVSTTATERTLHQEERHHGRDFLLRAASFDICESTEEFSSSQRPPCLRPSTSDGILRGNSYEKGQRGLRRGRILSSWSRAFVSIQEEMADDPLTYKSRLMVVVKMDPSIQPGPFRAVLKYLYTGELDENERDLMHIAHIAELLEVFDLRMMVANILNNEAFMNQEITKAFHVRRTNRVKECLAKGTFSDVTFILDDGAISAHKPLLISSCDWMAAMFGGPFVESSTQEVVFPYTSKSCMRAVLEYLYTGQFSSSPDLDDMKLIILANRLCLPHLVALTEQYMVAGLMEASQMLVDIDGDVLVFLELAQFHCAYQLADWCLHHICTNYNNVCRKFPRDMKAMSAENQEYFEKHRWPPVWYLKEEDHYQRAKKEREKEDYLHQKRQPKRRWLFWNTSSSPSASPSSSAATASSSSSSSSSSSSAVV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMN3</accession>
    <name>A0A452HMN3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">TP53 induced glycolysis regulatory phosphatase</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HMN3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018491.1">
        <property type="protein sequence ID" value="ENSGAGP00000016196.1"/>
        <property type="gene ID" value="ENSGAGG00000012153.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004331">
        <property type="term" value="F:fructose-2,6-bisphosphate 2-phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045820">
        <property type="term" value="P:negative regulation of glycolytic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043456">
        <property type="term" value="P:regulation of pentose-phosphate shunt"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07067">
        <property type="entry name" value="HP_PGM_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1240">
        <property type="entry name" value="Phosphoglycerate mutase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013078">
        <property type="entry name" value="His_Pase_superF_clade-1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029033">
        <property type="entry name" value="His_PPase_superfam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051695">
        <property type="entry name" value="Phosphoglycerate_Mutase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46517">
        <property type="entry name" value="FRUCTOSE-2,6-BISPHOSPHATASE TIGAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46517:SF1">
        <property type="entry name" value="FRUCTOSE-2,6-BISPHOSPHATASE TIGAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00300">
        <property type="entry name" value="His_Phos_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53254">
        <property type="entry name" value="Phosphoglycerate mutase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="binding site" evidence="2">
        <location>
            <position position="42"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="2">
        <location>
            <begin position="70"/>
            <end position="73"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="PIRSR" id="PIRSR613078-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016196.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="5">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="256" mass="28085" checksum="65D80CF12DABAFAF" modified="2019-05-08" version="1">MPFSGQGVDEPLSATGFRQADAAGIFLSNVKFTHVFSSDLLRAKQTASTIIGKNKFCKDIVIQYDARLRERKYGIAEGRPLSDLKAMANAAGEQCPSFTPSGGETLDEVRTRAKNFFEFLCQLSIEEECQKEQAVLGMAGNGLETLEGKPVFPLRNHCCGLELNSDTDKDTKMLNPNILVVSHGAYMRNWFGYFVLDLKCTLPSALKKSQLSSVSPNTGVSHFIINLGNGDVIKPEISCICLNRDDHLADVNTENV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMN4</accession>
    <name>A0A452HMN4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Stress-activated protein kinase JNK</fullName>
            <ecNumber evidence="12">2.7.11.24</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="12">Responds to activation by environmental stress and pro-inflammatory cytokines by phosphorylating a number of transcription factors, and thus regulates transcriptional activity.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.24"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="10">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.24"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 12">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="12">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 12">Belongs to the protein kinase superfamily. CMGC Ser/Thr protein kinase family. MAP kinase subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.24" evidence="12"/>
    <dbReference type="AlphaFoldDB" id="A0A452HMN4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018556.1">
        <property type="protein sequence ID" value="ENSGAGP00000016255.1"/>
        <property type="gene ID" value="ENSGAGG00000012196.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004707">
        <property type="term" value="F:MAP kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0106310">
        <property type="term" value="F:protein serine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048511">
        <property type="term" value="P:rhythmic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd07850">
        <property type="entry name" value="STKc_JNK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000009">
        <property type="entry name" value="Mitogen-activated protein kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000210">
        <property type="entry name" value="Mitogen-activated protein kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003527">
        <property type="entry name" value="MAP_kinase_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050117">
        <property type="entry name" value="MAPK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008351">
        <property type="entry name" value="MAPK_JNK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24055">
        <property type="entry name" value="MITOGEN-ACTIVATED PROTEIN KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01772">
        <property type="entry name" value="JNKMAPKINASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01351">
        <property type="entry name" value="MAPK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 12" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="9" id="KW-0090">Biological rhythms</keyword>
    <keyword evidence="7 12" id="KW-0418">Kinase</keyword>
    <keyword evidence="12" id="KW-0460">Magnesium</keyword>
    <keyword evidence="6 12" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="4 12" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 12" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="5 12" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Protein kinase" evidence="13">
        <location>
            <begin position="26"/>
            <end position="321"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008832"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023108"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00047592"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00048312"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="RuleBase" id="RU368052"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016255.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="382" mass="43915" checksum="B8F65E2EC14584E9" modified="2019-05-08" version="1">MSDGKCDSQFYSVQVADSTFTVLKQYQQLKPIGSGAQGIVCAAFDTVLGINVAVKKLSRPFQNQTHAKRAYRELVLLKCVNHKNIISLLNVFTPQKSLEEFQDVYLVMELMDANLCQVIHMELDHERMSYLLYQMLCGIKHLHSAGIIHRDLKPSNIVVKSDCTLKILDFGLARTACTNFMMTPYVVTRYYRAPEVILGMGYKENVDIWSVGCIMGELVKGCVIFQGTDHIDQWNKVIEQLGTPSADFMKKLQPTVRNYVENRPKYPGIKFEELFPDWIFPSESDRDKLKTSQARDLLSKMLVVDPDKRISVDEALRDPYITVWYDPAEAEAPPPKIYDAQLEEREHAIEEWKELIYKEVMDWEERSKNGVVKDQPSAQMQQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMN7</accession>
    <name>A0A452HMN7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Rho-related BTB domain-containing protein 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="7">Regulator of cell proliferation and apoptosis. It likely functions as a substrate-adapter that recruits key substrates, e.g. MSI2, to CUL3-based ubiquitin ligase complexes for degradation. Required for MSI2 ubiquitination and degradation.</text>
    </comment>
    <comment type="subunit">
        <text evidence="8">Interacts with HSP90AA1 and HSP90AB1. Forms a complex with CUL3 and RBX1. Interacts (via BTB 1 domain) with CUL3. Interacts with MSI2.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMN7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018504.1">
        <property type="protein sequence ID" value="ENSGAGP00000016208.1"/>
        <property type="gene ID" value="ENSGAGG00000012159.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010008">
        <property type="term" value="C:endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005525">
        <property type="term" value="F:GTP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003924">
        <property type="term" value="F:GTPase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007264">
        <property type="term" value="P:small GTPase-mediated signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd18531">
        <property type="entry name" value="BACK_RHOBTB2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18356">
        <property type="entry name" value="BTB1_POZ_RHOBTB2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18359">
        <property type="entry name" value="BTB2_POZ_RHOBTB2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd01873">
        <property type="entry name" value="RhoBTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.710.10:FF:000050">
        <property type="entry name" value="Rho related BTB domain containing 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.710.10:FF:000179">
        <property type="entry name" value="Rho-related BTB domain-containing protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000177">
        <property type="entry name" value="Rho-related BTB domain-containing protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.710.10:FF:000014">
        <property type="entry name" value="Rho-related BTB domain-containing protein 2 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.710.10">
        <property type="entry name" value="Potassium Channel Kv1.1, Chain A"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000210">
        <property type="entry name" value="BTB/POZ_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011333">
        <property type="entry name" value="SKP1/BTB/POZ_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001806">
        <property type="entry name" value="Small_GTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003578">
        <property type="entry name" value="Small_GTPase_Rho"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24072">
        <property type="entry name" value="RHO FAMILY GTPASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00651">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00071">
        <property type="entry name" value="Ras"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00449">
        <property type="entry name" value="RASTRNSFRMNG"/>
    </dbReference>
    <dbReference type="SMART" id="SM00225">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00175">
        <property type="entry name" value="RAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00173">
        <property type="entry name" value="RAS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00174">
        <property type="entry name" value="RHO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54695">
        <property type="entry name" value="POZ domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50097">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51419">
        <property type="entry name" value="RAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51421">
        <property type="entry name" value="RAS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51420">
        <property type="entry name" value="RHO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0342">GTP-binding</keyword>
    <keyword evidence="2" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0832">Ubl conjugation</keyword>
    <keyword evidence="3" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="BTB" evidence="10">
        <location>
            <begin position="289"/>
            <end position="468"/>
        </location>
    </feature>
    <feature type="domain" description="BTB" evidence="10">
        <location>
            <begin position="526"/>
            <end position="593"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="330"/>
            <end position="355"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="377"/>
            <end position="399"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="732"/>
            <end position="762"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="377"/>
            <end position="391"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00074749"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00093283"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00093525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50097"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016208.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="762" mass="86784" checksum="06EE86FD7F1E02FC" modified="2019-05-08" version="1">MEARAKAHHAVKKTKMYHSSSQLMDSDMDYERPNVETIKCVVVGDNAVGKTRLICARACNATLTQYQLLATHVPTVWAIDQYRVCQEVLERSRDVVDDVSVSLRLWDTFGDHHKDRRFAYGRSDVVVLCFSIANPNSLHHVKTMWYPEIKHFCPRAPVILVGCQLDLRYADLEAVNRARRPLARPIKPNEILPPEKGREVAKELGIPYYETSVVAQFGIKDVFDNAIRAALISRRHLQFWKSHLRNVQRPLLQAPFLPPKPPPPIIMVPDPPSNNEEHPAHLLEDPLCADVILVLQEKIKIYAHKIYLSTSSSKFYDLFLMDLSEEDQQGAAGGSSRVAVSTTATERTLHQEERHHGRDFLLRAASFDICESTEEFSSSQRPPCLRPSTSDGILRGNSYEKGQRGLRRGRILSSWSRAFVSIQEEMADDPLTYKSRLMVVVKMDPSIQPGPFRAVLKYLYTGELDENERDLMHIAHIAELLEVFDLRMMVANILNNEAFMNQEITKAFHVRRTNRVKECLAKGTFSDVTFILDDGAISAHKPLLISSCDWMAAMFGGPFVESSTQEVVFPYTSKSCMRAVLEYLYTGQFSSSPDLDDMKLIILANRLCLPHLVALTEQYMVAGLMEASQMLVDIDGDVLVFLELAQFHCAYQLADWCLHHICTNYNNVCRKFPRDMKAMSAENQEYFEKHRWPPVWYLKEEDHYQRAKKEREKEDYLHQKRQPKRRWLFWNTSSSPSASPSSSAATASSSSSSSSSSSSAVV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMQ3</accession>
    <name>A0A452HMQ3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">tRNA (guanosine(18)-2'-O)-methyltransferase TARBP1</fullName>
            <ecNumber evidence="10">2.1.1.34</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="12">TAR RNA-binding protein 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">S-adenosyl-L-methionine-dependent 2'-O-ribose methyltransferase that catalyzes the formation of 2'-O-methylguanosine at position 18 (Gm18) in a subset of tRNA. Selectively mediates Gm18 methylation of tRNAGln-TTG/CTG and tRNASer-TGA/GCT. Gm18 modification can enhance the stability of modified tRNAs.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="8">
            <text>guanosine(18) in tRNA + S-adenosyl-L-methionine = 2'-O-methylguanosine(18) in tRNA + S-adenosyl-L-homocysteine + H(+)</text>
            <dbReference type="Rhea" id="RHEA:20077"/>
            <dbReference type="Rhea" id="RHEA-COMP:10190"/>
            <dbReference type="Rhea" id="RHEA-COMP:10192"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57856"/>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
            <dbReference type="ChEBI" id="CHEBI:74269"/>
            <dbReference type="ChEBI" id="CHEBI:74445"/>
            <dbReference type="EC" id="2.1.1.34"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="8">
            <dbReference type="Rhea" id="RHEA:20078"/>
        </physiologicalReaction>
    </comment>
    <comment type="subunit">
        <text evidence="2">Monomer and homodimer.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the class IV-like SAM-binding methyltransferase superfamily. RNA methyltransferase TrmH family.</text>
    </comment>
    <dbReference type="EC" id="2.1.1.34" evidence="10"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018583.1">
        <property type="protein sequence ID" value="ENSGAGP00000016278.1"/>
        <property type="gene ID" value="ENSGAGG00000012102.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0141100">
        <property type="term" value="F:tRNA (guanine(18)-2'-O)-methyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030488">
        <property type="term" value="P:tRNA methylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd18091">
        <property type="entry name" value="SpoU-like_TRM3-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.1280.10:FF:000010">
        <property type="entry name" value="probable methyltransferase TARBP1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.1280.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029028">
        <property type="entry name" value="Alpha/beta_knot_MTases"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001537">
        <property type="entry name" value="SpoU_MeTrfase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025806">
        <property type="entry name" value="TARBP1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056921">
        <property type="entry name" value="TARBP1_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045330">
        <property type="entry name" value="TRM3/TARBP1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044748">
        <property type="entry name" value="Trm3/TARBP1_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029026">
        <property type="entry name" value="tRNA_m1G_MTases_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12029:SF11">
        <property type="entry name" value="METHYLTRANSFERASE TARBP1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12029">
        <property type="entry name" value="RNA METHYLTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00588">
        <property type="entry name" value="SpoU_methylase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25050">
        <property type="entry name" value="TARBP1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF75217">
        <property type="entry name" value="alpha/beta knot"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51624">
        <property type="entry name" value="SAM_MT_TRMH_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0007">Acetylation</keyword>
    <keyword evidence="3" id="KW-0489">Methyltransferase</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="5" id="KW-0949">S-adenosyl-L-methionine</keyword>
    <keyword evidence="4" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="TARBP1" evidence="14">
        <location>
            <begin position="198"/>
            <end position="302"/>
        </location>
    </feature>
    <feature type="domain" description="tRNA/rRNA methyltransferase SpoU type" evidence="13">
        <location>
            <begin position="1253"/>
            <end position="1394"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007228"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00011407"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022603"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022691"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00093266"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00093361"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00093594"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00093636"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00093656"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="Pfam" id="PF00588"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="Pfam" id="PF25050"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016278.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1409" mass="159088" checksum="5CA7FE409432AD9E" modified="2019-05-08" version="1">MELVLADALLGLAQSPDPCALLRAVCGGDGAAPLSVERVETLRFLLQWLEAGGAGGEALRRAAWEVALGRCWPLLRGGGGAAGPERRRLLRAACGALRLCARLCGAQLAARLAGEALAALRAEPPGAGLGAEAAVEALVAVAPELXXVWAGLLPGRGEERRVSRTLLALTALADLLFPGLEGDGGGEGPALDARLCGGFWRVLQAGLTDGDGLCRKRARYLLKRAVEVSDQLRTDCTCPPDDGDEPSLFWWSVEKNDKLLQFWENYILIMETLEGNQIHVIKPVLPKLNSLYEYALSEKGCWLFHPSWHMCIYKRMFESENKTLTREGVAQFLEIYETKRLPNSPGFSEVRAFRQLIGACPPLGMKLQKFLSTYLTLLSEEERRIFLLKFIQKMTSRHWCAVPILFLSMALACIPSCKALGTEGLHSLRDVLQCTMITHQILLRGAAQCFLLQTAMHLTDVEKVSLPEISSFLMSLRPEESLRENCFMPDWFEAKLIATMILLAADVEEMKNKHRNTERVELKSFLHPLLDVLLKLGTNAYIPTLKTDKSLQLLLKLLQTCLLKQYSPQDDLDRCNLYLALLSELVNLCSNAGWKKGPSIWNFISSLINASVKNLQETGNEQVILQGINVEWAFRLLILICFLLEHYIMAYSLYVKKTEEATSSQGWGKIVASYVHDQWVCLHFILNKYHKCIPLSNDETLELSLLAVQRPIRTLQSALEALTLLPSDQVLPVFHCMKVLVPKLLCSSESLCIDAFDLAWKVISSLSNTQLIFWSNLKAFVQFVFDTEVLAIAASSKGQAYSKIKEIIVKLVEMSTARTGVFNTVISYCCQSWLFPTADDTGALENPFSNAGNYSELITEACVFGMVFRRDQRLIQDVHAFVENLGDECAANVVTESTNRDDHYVRVCAIKFLCWLDGSNTLHKMFMEDLVIKLLDKDELVSKSRTRYYVNSLQHRMKNRLWQTLLVIFPKLDQNFLTGTLDKIFQAGFGNNQASIKYLIEWIIILILQRYPQCLEKFWNCFSYGEEKLKTSICTFLSVLSHFDIILENTSEKKPILKKALLVVLQWCFNHNFSVRLYALVALKKIWGMCKALYVEEFEALTPVIESSLHQVENMHGAGNARKNWQRIQDHFFFATFHPLKDYSLETIFYTLPRLSELVEDEWIPLCKFTRYTDISLPAAFHWLHSRTELLNLKPSDWSQQDMGSDLVEMDNQTEWTDVQKKIIPWKNSSPDLDLELLFQDRAAKLGKSNSRLIVVASLIDKPTNLGGLCRTSEIFGASALVVGSLHYINDKQFQHLSVSAEQWLPLVEVKPSQLVNYLQQKKTEGYTIIGVEQTAKSSDLTEYCFPEKSLLLLGNEHEGIPANLIQHLDVCVEIPQQGIIRSLNVHVSGALLIWEYTRQQLIKQKQLK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMR0</accession>
    <name>A0A452HMR0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">Receptor-type tyrosine-protein phosphatase alpha</fullName>
            <ecNumber evidence="4">3.1.3.48</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="16">Tyrosine protein phosphatase which is involved in integrin-mediated focal adhesion formation. Following integrin engagement, specifically recruits BCAR3, BCAR1 and CRK to focal adhesions thereby promoting SRC-mediated phosphorylation of BRAC1 and the subsequent activation of PAK and small GTPase RAC1 and CDC42.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell junction</location>
            <location evidence="1">Focal adhesion</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the protein-tyrosine phosphatase family. Receptor class 4 subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.48" evidence="4"/>
    <dbReference type="AlphaFoldDB" id="A0A452HMR0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018534.1">
        <property type="protein sequence ID" value="ENSGAGP00000016234.1"/>
        <property type="gene ID" value="ENSGAGG00000010212.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005925">
        <property type="term" value="C:focal adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="CDD" id="cd14623">
        <property type="entry name" value="R-PTPc-A-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000007">
        <property type="entry name" value="Receptor-type tyrosine-protein phosphatase alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000021">
        <property type="entry name" value="Receptor-type tyrosine-protein phosphatase alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050348">
        <property type="entry name" value="Protein-Tyr_Phosphatase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000242">
        <property type="entry name" value="PTP_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016130">
        <property type="entry name" value="Tyr_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003595">
        <property type="entry name" value="Tyr_Pase_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19134">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19134:SF433">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00102">
        <property type="entry name" value="Y_phosphatase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00700">
        <property type="entry name" value="PRTYPHPHTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00194">
        <property type="entry name" value="PTPc"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00404">
        <property type="entry name" value="PTPc_motif"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00383">
        <property type="entry name" value="TYR_PHOSPHATASE_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50055">
        <property type="entry name" value="TYR_PHOSPHATASE_PTP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12" id="KW-0965">Cell junction</keyword>
    <keyword evidence="5" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="15" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="14" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="11" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0677">Repeat</keyword>
    <keyword evidence="8" id="KW-0732">Signal</keyword>
    <keyword evidence="7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="13" id="KW-1133">Transmembrane helix</keyword>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="18">
        <location>
            <begin position="37"/>
            <end position="282"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="19">
        <location>
            <begin position="202"/>
            <end position="273"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="18">
        <location>
            <begin position="314"/>
            <end position="572"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="19">
        <location>
            <begin position="488"/>
            <end position="563"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007765"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00013064"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00054255"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00074216"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016234.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="583" mass="67253" checksum="50E7D6FA18D76CE8" modified="2019-05-08" version="1">MPLLARSPSTNRKYPPLPVDKLEEEINRRMADDNKLFREEFNALPACPIQATCEAASKEENKEKNRYVNILPYDHSRVHLTPIEGVPDSDYINASFINVSSCMGPKEETVNDFWRMIWEQNTATIVMVTNLKERKECKCAQYWPDQGCWTYGNIRVSVEDVTVLVDYTVRKFCIQQPQRLVTQFHFTSWPDFGVPFTPIGMLKFLKKVKTCNPQYAGAIVVHCSAGVGRTGTFIVIDAMLDMMHAERKVDVYGFVSRIRAQRCQMVQTDMQYVFIYQALLEHYLYGDTELEVTSLETHLQKIYNKVPGTSSNGLEEEFKKLTSIKIQNDKMRTGNLPANMKKNRVLQIIPYEFNRVIIPVKRGEENTDYVNASFIDGYRQKDSYIASQGPLQHTIEDFWRMIWEWKSCSIVMLTELEERGQEKCAQYWPSDGSVSYGDITMELKKEEECESYTVRDLLVTNTRENKSRQIRQFHFHGWPEVGIPSDGKGMINIIAAVQKQQQQSGNHPITVHCSAGAGRTGTFCALSTVLERVKAEGILDVFQTVKSLRLQRPHMVQTLEQYEFCYKVVQEYIDAFSDYANFK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMR1</accession>
    <name>A0A452HMR1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Charged multivesicular body protein 7</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="10">Chromatin-modifying protein 7</fullName>
        </alternativeName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus envelope</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the SNF7 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMR1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018521.1">
        <property type="protein sequence ID" value="ENSGAGP00000016221.1"/>
        <property type="gene ID" value="ENSGAGG00000012166.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009898">
        <property type="term" value="C:cytoplasmic side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000815">
        <property type="term" value="C:ESCRT III complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005771">
        <property type="term" value="C:multivesicular body"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005635">
        <property type="term" value="C:nuclear envelope"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032511">
        <property type="term" value="P:late endosome to vacuole transport via multivesicular body sorting pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006900">
        <property type="term" value="P:vesicle budding from membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.1230">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057471">
        <property type="entry name" value="CHMP7_WHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005024">
        <property type="entry name" value="Snf7_fam"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22761">
        <property type="entry name" value="CHARGED MULTIVESICULAR BODY PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22761:SF21">
        <property type="entry name" value="CHARGED MULTIVESICULAR BODY PROTEIN 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03357">
        <property type="entry name" value="Snf7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25239">
        <property type="entry name" value="WHD_CHMP7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25880">
        <property type="entry name" value="WHD_CHMP7_1st"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7 11" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="6" id="KW-0653">Protein transport</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0813">Transport</keyword>
    <feature type="domain" description="CHMP7 winged helix" evidence="13">
        <location>
            <begin position="154"/>
            <end position="222"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="385"/>
            <end position="429"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="11">
        <location>
            <begin position="246"/>
            <end position="306"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004259"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00006190"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00041077"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00041629"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="Pfam" id="PF25239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016221.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="429" mass="48553" checksum="A03AA57015D41994" modified="2019-05-08" version="1">MCSPAETLPGPAPEGDLPPEWESDEERMAFLFSAFKQSREVNSTEWDSKMGFWAELVLGRGRRRGAVRTCLRELQQAFQRRGSVPLGLGTVLRELLRRGKLQRESDFIASVDCSWISWGVGIFILKPLKWTLSSVLGDSKVPEEEEVLIFVELLQEKAEEVYRLYQNSALSSHPVVVLSELRSLCASICPDERTFYLLLLQLQKEKRVTVMEQNREKIVKFARGLHAKVSPVNDVDIGVYQLMQSEQLLSQKVESLSQEAERCKEDARSACRAGKKQLALRYLKSKRRTERRIEELLSKLDAVQGILDRIYASQTDQMVFNAYQAGVGALKLSMKDVTVEKTENLVDQIQELCDTQDEVVQALAGMSINGLEVDSEELEKELDSLLQDSTKEPLDLPPVPQKPLLPSISDTELEAELEKKDGGMYRQQS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMR2</accession>
    <name>A0A452HMR2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">SAM domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the BicC family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMR2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018588.1">
        <property type="protein sequence ID" value="ENSGAGP00000016283.1"/>
        <property type="gene ID" value="ENSGAGG00000012189.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="CDD" id="cd22421">
        <property type="entry name" value="KH-I_BICC1_rpt2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd22422">
        <property type="entry name" value="KH-I_BICC1_rpt3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd09520">
        <property type="entry name" value="SAM_BICC1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.50:FF:000025">
        <property type="entry name" value="Ankyrin repeat and sterile alpha motif domain-containing 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.310.270:FF:000001">
        <property type="entry name" value="BicC family RNA binding protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.310.270:FF:000002">
        <property type="entry name" value="BicC family RNA binding protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.310.270">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1370.10">
        <property type="entry name" value="K Homology domain, type 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.50">
        <property type="entry name" value="Transcription Factor, Ets-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR054727">
        <property type="entry name" value="BICC1_KH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047549">
        <property type="entry name" value="BICC1_KH-I_rpt1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047554">
        <property type="entry name" value="BICC1_KH-I_rpt2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047553">
        <property type="entry name" value="BICC1_KH-I_rpt3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037974">
        <property type="entry name" value="BICC1_SAM_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004087">
        <property type="entry name" value="KH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004088">
        <property type="entry name" value="KH_dom_type_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036612">
        <property type="entry name" value="KH_dom_type_1_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001660">
        <property type="entry name" value="SAM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013761">
        <property type="entry name" value="SAM/pointed_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10627:SF78">
        <property type="entry name" value="PROTEIN BICAUDAL C HOMOLOG 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10627">
        <property type="entry name" value="SCP160"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00013">
        <property type="entry name" value="KH_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22985">
        <property type="entry name" value="KH_BICC1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24234">
        <property type="entry name" value="KH_BICC1_1st"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00536">
        <property type="entry name" value="SAM_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00322">
        <property type="entry name" value="KH"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00454">
        <property type="entry name" value="SAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54791">
        <property type="entry name" value="Eukaryotic type KH-domain (KH-domain type I)"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47769">
        <property type="entry name" value="SAM/Pointed domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50084">
        <property type="entry name" value="KH_TYPE_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50105">
        <property type="entry name" value="SAM_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="4 5" id="KW-0694">RNA-binding</keyword>
    <feature type="domain" description="SAM" evidence="7">
        <location>
            <begin position="807"/>
            <end position="870"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="524"/>
            <end position="584"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="613"/>
            <end position="632"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="714"/>
            <end position="780"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="524"/>
            <end position="553"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="568"/>
            <end position="584"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="726"/>
            <end position="747"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007662"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00117"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50105"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016283.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="908" mass="98566" checksum="3E04D47E18724A39" modified="2019-05-08" version="1">RTLLPYWTDERWNIMEETNTQIAWPSKLKIGAKSKKDPHIKVSGKKENVKEAKEKIMSVLDTKSNRVTLKMDVSHTEHSHVIGKGGNNIKKVMEETGCHIHFPDSNRNNQAEKSNQVSIAGQPAGVESARVRIRELLPLVLMFELPIAGILQPIPDPNSPTILHISQTYNISVSFKQRSRMYGATVIVRGSQNNTSAVKEGTAMLLEHLAGSLASAIPVSTQLDIAAQHHLFMMGRNGSNIKHIMQRTGAQIHFPDPSNPQKKSTVYLQGTIESVCLARQYLMGCLPLVLMFDMKEEIEIEPQIITQLMEQLDVFISIKPKPKQPSKSVIVKSVERNALNMYETRKRLLGLESNGVTIAANPSTVSCHVGLSCPGLDILPSAGLGLTGLGLLGPTALSVNTSTAPNSLLNALNSSVSPLQSPSSGTPSPTLWATSLANTSATGFSAIPHLMIPSTAQAALTSILLSGVPNYSQNTPSPPPGLTPVDIHVNGMQSENKKGSASLNGHTSNIKYAALSASSLGENVLSTNHSDSSRQTNSHSTSEQVTTKSNTAEGCNDAFVEVGMPRSPSHSANASDLKQMMSSSKQSCTKRQTVELLQGTKNSHLHTTERLLSDTELSASENPVADKKAPGSERAAERAAAAQQNSERVRLAPQSSYVNMQAFDYEQKKLLATKAMLKKPVVTEVRTPTNTWSGLGFSKSMPAETIKELRRANHVSYKPSMTTTYEGSSMSLSPSSSREHLGSGSESDNWRDRNGIGPSAHSEFVSSVGSPKRKQNKSTEHYLSSSNYMDCISSLTGSNGCNLNSLFKGSDLPELFSKLGLGKYTDVFQQQEIDLQTFLTLTDQDLKELGITTFGARRKMLLAISELNKNRRKLFDPSNVRASFLEGGASGRLPRQYHSDIASVSGRW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMR3</accession>
    <name>A0A452HMR3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">glutamine--fructose-6-phosphate transaminase (isomerizing)</fullName>
            <ecNumber evidence="3">2.6.1.16</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>D-fructose 6-phosphate + L-glutamine = D-glucosamine 6-phosphate + L-glutamate</text>
            <dbReference type="Rhea" id="RHEA:13237"/>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
            <dbReference type="ChEBI" id="CHEBI:58359"/>
            <dbReference type="ChEBI" id="CHEBI:58725"/>
            <dbReference type="ChEBI" id="CHEBI:61527"/>
            <dbReference type="EC" id="2.6.1.16"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="2">Nucleotide-sugar biosynthesis; UDP-N-acetyl-alpha-D-glucosamine biosynthesis; alpha-D-glucosamine 6-phosphate from D-fructose 6-phosphate: step 1/1.</text>
    </comment>
    <dbReference type="EC" id="2.6.1.16" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452HMR3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018550.1">
        <property type="protein sequence ID" value="ENSGAGP00000016249.1"/>
        <property type="gene ID" value="ENSGAGG00000012193.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00113">
        <property type="reaction ID" value="UER00528"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097367">
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        <property type="project" value="InterPro"/>
    </dbReference>
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        <property type="term" value="F:glutamine-fructose-6-phosphate transaminase (isomerizing) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
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    <dbReference type="GO" id="GO:0006002">
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        <property type="project" value="TreeGrafter"/>
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    <dbReference type="GO" id="GO:0006487">
        <property type="term" value="P:protein N-linked glycosylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006048">
        <property type="term" value="P:UDP-N-acetylglucosamine biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
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        <property type="entry name" value="GFAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05008">
        <property type="entry name" value="SIS_GlmS_GlmD_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05009">
        <property type="entry name" value="SIS_GlmS_GlmD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10490:FF:000001">
        <property type="entry name" value="Glutamine--fructose-6-phosphate aminotransferase [isomerizing]"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10490:FF:000126">
        <property type="entry name" value="Glutamine--fructose-6-phosphate aminotransferase [isomerizing] 1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Glucose-6-phosphate isomerase like protein, domain 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.60.20.10">
        <property type="entry name" value="Glutamine Phosphoribosylpyrophosphate, subunit 1, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017932">
        <property type="entry name" value="GATase_2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005855">
        <property type="entry name" value="GFAT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047084">
        <property type="entry name" value="GFAT_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035466">
        <property type="entry name" value="GlmS/AgaS_SIS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035490">
        <property type="entry name" value="GlmS/FrlB_SIS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029055">
        <property type="entry name" value="Ntn_hydrolases_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001347">
        <property type="entry name" value="SIS_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046348">
        <property type="entry name" value="SIS_dom_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01135">
        <property type="entry name" value="glmS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF001484">
        <property type="entry name" value="PRK00331.1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10937">
        <property type="entry name" value="GLUCOSAMINE--FRUCTOSE-6-PHOSPHATE AMINOTRANSFERASE, ISOMERIZING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10937:SF10">
        <property type="entry name" value="GLUTAMINE--FRUCTOSE-6-PHOSPHATE AMINOTRANSFERASE [ISOMERIZING] 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13522">
        <property type="entry name" value="GATase_6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01380">
        <property type="entry name" value="SIS"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56235">
        <property type="entry name" value="N-terminal nucleophile aminohydrolases (Ntn hydrolases)"/>
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    </dbReference>
    <dbReference type="SUPFAM" id="SSF53697">
        <property type="entry name" value="SIS domain"/>
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    </dbReference>
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        <property type="entry name" value="GATASE_TYPE_2"/>
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    </dbReference>
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    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Glutamine amidotransferase type-2" evidence="8">
        <location>
            <begin position="22"/>
            <end position="308"/>
        </location>
    </feature>
    <feature type="domain" description="SIS" evidence="9">
        <location>
            <begin position="380"/>
            <end position="519"/>
        </location>
    </feature>
    <feature type="domain" description="SIS" evidence="9">
        <location>
            <begin position="551"/>
            <end position="692"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001031"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00004775"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00012916"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022962"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51278"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51464"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016249.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="702" mass="79697" checksum="88B28F51141E1C0A" modified="2019-05-08" version="1">MQCDRTARFLRWRQDLQMNVFSGIFAYLNYRVPRTRKEIFETLIKGLQRLEYRGYDSAGVAIDGNNNEDKERYIRLIKKRGKVKALDEELYKQDGMDLKADFETHFGIAHTRWATHGVPNAVNSHPQRSDKRNEFVVIHNGIITNYKDLRKFLETKGYEFESETDTETIPKLVKYMYDNRETEDISFSALVERVIQQLEGAFALVFKSIHYPGEAVATRRGSPLLIGVRSKYKLSTEQIPILYRTCNIENVKNICKARMKRLDSSTCLHAIGDKAVEFFFASDASAIIEHTNRVIFLEDDDIAAVADGKLSLHRLKRSASDDPSRAIQTLQMELQQIMKGNFSAFMQKEIFEQPESVVNTMRGRVNFETKTVLLGGLKDHLKEIRRCRRLIIIGCGTSYHAAIATRQVLEELTELPVMVELASDFLDRNTPVFRDDVCFFISQSGETADTLMALRYCKDHGALTVGITNTVGSSISRETDCGVHINAGPEIGVASTKAYTSQFISLVMFGLMMSEDRISLQNRRQEIISGLQMLPEMIKEVLSLDEKIHDLALELYKQRSLLVMGRGYNYATCLEGALKIKEITYMHSEGILAGELKHGPLALIDKHMPVIMVIMKDPCFTKCQNALQQVIARQGRPIILSSREDTESSKFAYKTIELPHTVDCLQGVLSVVPLQLLSFHLAVLRGYDVDFPRNLAKSVTVE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMR8</accession>
    <name>A0A452HMR8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">glutamine--fructose-6-phosphate transaminase (isomerizing)</fullName>
            <ecNumber evidence="3">2.6.1.16</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>D-fructose 6-phosphate + L-glutamine = D-glucosamine 6-phosphate + L-glutamate</text>
            <dbReference type="Rhea" id="RHEA:13237"/>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
            <dbReference type="ChEBI" id="CHEBI:58359"/>
            <dbReference type="ChEBI" id="CHEBI:58725"/>
            <dbReference type="ChEBI" id="CHEBI:61527"/>
            <dbReference type="EC" id="2.6.1.16"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="2">Nucleotide-sugar biosynthesis; UDP-N-acetyl-alpha-D-glucosamine biosynthesis; alpha-D-glucosamine 6-phosphate from D-fructose 6-phosphate: step 1/1.</text>
    </comment>
    <dbReference type="EC" id="2.6.1.16" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452HMR8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018555.1">
        <property type="protein sequence ID" value="ENSGAGP00000016254.1"/>
        <property type="gene ID" value="ENSGAGG00000012193.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00113">
        <property type="reaction ID" value="UER00528"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097367">
        <property type="term" value="F:carbohydrate derivative binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004360">
        <property type="term" value="F:glutamine-fructose-6-phosphate transaminase (isomerizing) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006002">
        <property type="term" value="P:fructose 6-phosphate metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006487">
        <property type="term" value="P:protein N-linked glycosylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006048">
        <property type="term" value="P:UDP-N-acetylglucosamine biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="CDD" id="cd00714">
        <property type="entry name" value="GFAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05008">
        <property type="entry name" value="SIS_GlmS_GlmD_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05009">
        <property type="entry name" value="SIS_GlmS_GlmD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10490:FF:000001">
        <property type="entry name" value="Glutamine--fructose-6-phosphate aminotransferase [isomerizing]"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10490:FF:000126">
        <property type="entry name" value="Glutamine--fructose-6-phosphate aminotransferase [isomerizing] 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.10490">
        <property type="entry name" value="Glucose-6-phosphate isomerase like protein, domain 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.60.20.10">
        <property type="entry name" value="Glutamine Phosphoribosylpyrophosphate, subunit 1, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017932">
        <property type="entry name" value="GATase_2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005855">
        <property type="entry name" value="GFAT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047084">
        <property type="entry name" value="GFAT_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035466">
        <property type="entry name" value="GlmS/AgaS_SIS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035490">
        <property type="entry name" value="GlmS/FrlB_SIS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029055">
        <property type="entry name" value="Ntn_hydrolases_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001347">
        <property type="entry name" value="SIS_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046348">
        <property type="entry name" value="SIS_dom_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01135">
        <property type="entry name" value="glmS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF001484">
        <property type="entry name" value="PRK00331.1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10937">
        <property type="entry name" value="GLUCOSAMINE--FRUCTOSE-6-PHOSPHATE AMINOTRANSFERASE, ISOMERIZING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10937:SF10">
        <property type="entry name" value="GLUTAMINE--FRUCTOSE-6-PHOSPHATE AMINOTRANSFERASE [ISOMERIZING] 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13522">
        <property type="entry name" value="GATase_6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01380">
        <property type="entry name" value="SIS"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56235">
        <property type="entry name" value="N-terminal nucleophile aminohydrolases (Ntn hydrolases)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53697">
        <property type="entry name" value="SIS domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51278">
        <property type="entry name" value="GATASE_TYPE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51464">
        <property type="entry name" value="SIS"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0315">Glutamine amidotransferase</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Glutamine amidotransferase type-2" evidence="8">
        <location>
            <begin position="2"/>
            <end position="288"/>
        </location>
    </feature>
    <feature type="domain" description="SIS" evidence="9">
        <location>
            <begin position="360"/>
            <end position="499"/>
        </location>
    </feature>
    <feature type="domain" description="SIS" evidence="9">
        <location>
            <begin position="531"/>
            <end position="672"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001031"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004775"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012916"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022962"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51278"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51464"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016254.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="682" mass="77147" checksum="2F93F1452D88E1D4" modified="2019-05-08" version="1">MCGIFAYLNYRVPRTRKEIFETLIKGLQRLEYRGYDSAGVAIDGNNNEDKERYIRLIKKRGKVKALDEELYKQDGMDLKADFETHFGIAHTRWATHGVPNAVNSHPQRSDKRNEFVVIHNGIITNYKDLRKFLETKGYEFESETDTETIPKLVKYMYDNRETEDISFSALVERVIQQLEGAFALVFKSIHYPGEAVATRRGSPLLIGVRSKYKLSTEQIPILYRTCNIENVKNICKARMKRLDSSTCLHAIGDKAVEFFFASDASAIIEHTNRVIFLEDDDIAAVADGKLSLHRLKRSASDDPSRAIQTLQMELQQIMKGNFSAFMQKEIFEQPESVVNTMRGRVNFETKTVLLGGLKDHLKEIRRCRRLIIIGCGTSYHAAIATRQVLEELTELPVMVELASDFLDRNTPVFRDDVCFFISQSGETADTLMALRYCKDHGALTVGITNTVGSSISRETDCGVHINAGPEIGVASTKAYTSQFISLVMFGLMMSEDRISLQNRRQEIISGLQMLPEMIKEVLSLDEKIHDLALELYKQRSLLVMGRGYNYATCLEGALKIKEITYMHSEGILAGELKHGPLALIDKHMPVIMVIMKDPCFTKCQNALQQVIARQGRPIILSSREDTESSKFAYKTIELPHTVDCLQGVLSVVPLQLLSFHLAVLRGYDVDFPRNLAKSVTVE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN01</accession>
    <name>A0A452HN01_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="3">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HN01"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018654.1">
        <property type="protein sequence ID" value="ENSGAGP00000016346.1"/>
        <property type="gene ID" value="ENSGAGG00000012116.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008028">
        <property type="term" value="F:monocarboxylic acid transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1250.20">
        <property type="entry name" value="MFS general substrate transporter like domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011701">
        <property type="entry name" value="MFS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036259">
        <property type="entry name" value="MFS_trans_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050327">
        <property type="entry name" value="Proton-linked_MCT"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11360">
        <property type="entry name" value="MONOCARBOXYLATE TRANSPORTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11360:SF14">
        <property type="entry name" value="MONOCARBOXYLATE TRANSPORTER 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07690">
        <property type="entry name" value="MFS_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103473">
        <property type="entry name" value="MFS general substrate transporter"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="2" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="20"/>
            <end position="47"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="110"/>
            <end position="137"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="149"/>
            <end position="169"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="204"/>
            <end position="224"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="231"/>
            <end position="250"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="270"/>
            <end position="294"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016346.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="314" mass="34383" checksum="70D5328713D23F21" modified="2019-05-08" version="1">LTAMEEEEQQPIPPPDGGWGWMIVLHFFLVNVLVMGMLKTFGIFFVVFQEEVGGSSEQISWIGSIMSSLRFSAGPLVTIACGKLGDQLTSIIGAGLVSSGCLISTQATSISFLCVSMGLLLGLGFACLYQAAAVMTARYFKKRLALSNAIARSGMGLTFILAPFTQLLIDFYGWQGITETVSQLLSGWIADQNWIKKYHYHKTYLILCGITNLLGPLATSFPLLMVYTVSFAIFCGGYLALLLPVLVTMWESPAALITKPSSIAGWLSDYTQTYACSFLFAGTCYLIAPISFFFEPSAHKWKLQKEKELNEVTS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN22</accession>
    <name>A0A452HN22_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Collagen alpha-1(XII) chain</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">Type XII collagen interacts with type I collagen-containing fibrils, the COL1 domain could be associated with the surface of the fibrils, and the COL2 and NC3 domains may be localized in the perifibrillar matrix.</text>
    </comment>
    <comment type="subunit">
        <text evidence="14">Trimer of identical chains each containing 190 kDa of non-triple-helical sequences.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="12">Belongs to the fibril-associated collagens with interrupted helices (FACIT) family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018702.1">
        <property type="protein sequence ID" value="ENSGAGP00000016388.1"/>
        <property type="gene ID" value="ENSGAGG00000011657.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005581">
        <property type="term" value="C:collagen trimer"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005614">
        <property type="term" value="C:interstitial matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005201">
        <property type="term" value="F:extracellular matrix structural constituent"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043394">
        <property type="term" value="F:proteoglycan binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007160">
        <property type="term" value="P:cell-matrix adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007044">
        <property type="term" value="P:cell-substrate junction assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007507">
        <property type="term" value="P:heart development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009888">
        <property type="term" value="P:tissue development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="17"/>
    </dbReference>
    <dbReference type="CDD" id="cd01482">
        <property type="entry name" value="vWA_collagen_alphaI-XII-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000018">
        <property type="entry name" value="collagen alpha-1(XII) chain isoform X1"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000489">
        <property type="entry name" value="collagen alpha-1(XII) chain isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000639">
        <property type="entry name" value="collagen alpha-1(XII) chain isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000816">
        <property type="entry name" value="collagen alpha-1(XII) chain isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.200:FF:000008">
        <property type="entry name" value="Collagen type XII alpha 1 chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000121">
        <property type="entry name" value="Collagen type XII alpha 1 chain"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000234">
        <property type="entry name" value="Collagen, type XII, alpha 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000480">
        <property type="entry name" value="Collagen, type XII, alpha 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.410:FF:000001">
        <property type="entry name" value="Collagen, type XII, alpha 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.200">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="17"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.410">
        <property type="entry name" value="von Willebrand factor, type A domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008160">
        <property type="entry name" value="Collagen"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050991">
        <property type="entry name" value="ECM_Regulatory_Proteins"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048287">
        <property type="entry name" value="TSPN-like_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002035">
        <property type="entry name" value="VWF_A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036465">
        <property type="entry name" value="vWFA_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46708:SF7">
        <property type="entry name" value="FIBRONECTIN TYPE-III DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46708">
        <property type="entry name" value="TENASCIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01391">
        <property type="entry name" value="Collagen"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="17"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00092">
        <property type="entry name" value="VWA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00453">
        <property type="entry name" value="VWFADOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="17"/>
    </dbReference>
    <dbReference type="SMART" id="SM00210">
        <property type="entry name" value="TSPN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00327">
        <property type="entry name" value="VWA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53300">
        <property type="entry name" value="vWA-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="16"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50234">
        <property type="entry name" value="VWFA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="8" id="KW-0176">Collagen</keyword>
    <keyword evidence="9" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11" id="KW-0379">Hydroxylation</keyword>
    <keyword evidence="7" id="KW-0654">Proteoglycan</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="1"/>
            <end position="84"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="87"/>
            <end position="181"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="182"/>
            <end position="267"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="269"/>
            <end position="358"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="360"/>
            <end position="450"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="451"/>
            <end position="539"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="541"/>
            <end position="630"/>
        </location>
    </feature>
    <feature type="domain" description="VWFA" evidence="17">
        <location>
            <begin position="650"/>
            <end position="822"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="838"/>
            <end position="927"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="928"/>
            <end position="1019"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="1020"/>
            <end position="1109"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="1110"/>
            <end position="1200"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="1210"/>
            <end position="1304"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="1391"/>
            <end position="1481"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="1482"/>
            <end position="1572"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="1573"/>
            <end position="1662"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="1663"/>
            <end position="1751"/>
        </location>
    </feature>
    <feature type="domain" description="VWFA" evidence="17">
        <location>
            <begin position="1779"/>
            <end position="1952"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="163"/>
            <end position="182"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="524"/>
            <end position="551"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1466"/>
            <end position="1485"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="2203"/>
            <end position="2352"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="2388"/>
            <end position="2503"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1468"/>
            <end position="1480"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="16">
        <location>
            <begin position="2232"/>
            <end position="2249"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="16">
        <location>
            <begin position="2273"/>
            <end position="2284"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="16">
        <location>
            <begin position="2312"/>
            <end position="2330"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="16">
        <location>
            <begin position="2397"/>
            <end position="2406"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="16">
        <location>
            <begin position="2413"/>
            <end position="2422"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="16">
        <location>
            <begin position="2463"/>
            <end position="2476"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022974"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023119"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023278"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00049648"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00053577"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00064391"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00067989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50234"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016388.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2521" mass="272336" checksum="F896C6772D6E6101" modified="2019-05-08" version="1">MITDETTDSFKVGWTPAPGNVLRYRVEYRPVTGRERKHVTVPANERSTTLLNLTPDTRYEVYVIPEYQSGPGNSLNGYAKTEEVLGTPRNLRVSDATTSTMKLSWSAAPGKVQRYLVTYTPVAGGETKEVTIRGDTTTTVLRDLEHGTRYALSVTALYASGAGDALSGQGETPEARGSPRNLRVSDATTSTMKLSWSAAPGKVERYRITYTPVAGGDTKEVTTRGDTTTTVLRDLEHGTRYALSVTALYASGAGDALPGQGETLEARGSPRNLRVSDATTSTMKLSWSAAPGKVQRYRITYTPVAGGEAKEVTIRGDTTTTVLRDLEQGTRYTLSVTALYASGAGDALPGQGETLEDRGSPRDLLIRDITDTTMGLSWTAAPGAVRRYRIVWRSLYDGQTGESTVPGNVINTVLENLQPETKYKISVIASYSSGDGAPLEGEATTEVSPNARRVRVDEETENSMRVTWQPAPGKVVSYRVVYRPVSGGRQVVAKVPATSTATVLKRLQPLTKYNISVIPMYKSGEGKHRQGEGTTASPYKPPRNLKTSDSTMSSFRVTWEPAPGEVRGYKITFHPTGEDRLGELIVGPYDNTVVLEELRAGTTYKVNVFGMFEGGESAPLVGQEMTTLSDVTVMPFLARGLECTTRAEADIVLLVDGSWSIGRPNFKTVRNFIARIVEVFDIGPDKVQIALAQYSGDPRTEWHLNTYGSKQSLLDAVANLPYKGGNTLTGMALSFILKDNFKPEVGLRPKARKIGVLITDGKSQDDIVAPSRRLRDEGVELYAIGIKNADENELKQIATDPDDTHVFNVADFSFLTNIVDDLTINLCNSVKGPGGLSSPSNLVTSEVTPRSFRVTWTAPVESVDRYRVEYYPVSGGPPQQFYVNRVETTTVLKNLKPETEYVVKVFSVVEDESSEPLSGTETTLPIPTIRNLNAYDIGSTSMRVRWEPLRGATGYRLLYEPLNATIPAVEKEMRVGSTVNDVQLVDLIPNAAYTLTIHAEFGDLSSDPLTTQEVTLPLAGAKSLRVRDITHSSMNVIWDPAPGKVRKYLLRYKTTLGDEPKEVEVDKSKTSTVLPSLLSQTLYDLQLVAVYDEGESIPVAAKATTLPVPAPVNLRLTEVTKQSFRGTWDHGAPDVALYRITWGPYGGTEKQETILNGDENSLVFENLNPDTLYDVSVTAIYPDESESDDLIGSERTLPAIRPTTQAPKSGPRNLQVYNATSNSLTVKWDPASGRVQRYRITYQRTTGDGAEQSTMVGGRQNSVVLQKLQPDTPYTITVSSMYADGEGGRMTGRGKTKPLNTVKNLRVYDPTTSTLSVRWDHAEGSPRQYKVYYVPTTGGAEEMTTVPGNTNYIILRSLQPDTPYKITVVPTYPEGDGGRVSDTGRTLVRGAARNIQVYNPTPNTLNVRWEPAPGPVQQYRVVYAPLVGPRPSESIVVPANTRDVVLERLTPDTPYSVNVVALYADGEGNQSSGQGRTTPRSGPRNLKVFDPTTNSLSVQWDHADGPVQQYRIIYSPTVGDPIDEYTTVPGRRNNVILQPLHSDTPYKITVVAVYDDGDGGQLTGNGRTVGLLPPQNIHISDEWYTRFRVSWDPSPSPVLGYKLAYKPVGSAEPPMEVFVGEVTSYTLHNLSPSTIYDVNVYAQYDSGLSAPLIDQGTTLYLNVTDLTSYKVGWDTFCIRWSPHRSASSYRLKLNPSDGSRGQEITVRGTETSHCFTGLSPDTEYNATVFVQTPNLEGPPISLKERTFIKPTEAPTQPPPPPPPPTIPPARDVCKGAKADIVFLTDASWSIGDDNFNKVVKFVFNTVGAFDLINPAGIQVSFVQYSDDAKSEFKLNTYDDKAQALGALQNIRYKGGNTRTGKALTFINEKVLTWESGMRRGVPKVLVVVTDGRSQDEVKKAAAIIQHSGFSIFVVGVADVDYHELSKIASKPSERHVFIVDDFDAFEKIEENLLTFICETATSTCPLIYLEGYTSPGFKMLESYNLTEKHFASVQGVSLESGSFPSHVAYRLHRNAFISQPASEIHPDGLPHAYTIILLFRLLSETPNEPFAIWQITDRDFKPQVGVILDPSTKLLSFFNKDTRGEVQTVTFDDNEVKKIFYGSFHKVHIVVTSTSVKIYIDCSEVMEKPIKESGNITTDGFEILGKLVKGDRKSATFEIQSFDIVCNAVWTSRDRCCDLPSMRDEAKCPALPNACTCTQDSVGPPGPPGPAGGPGTKGPRGERGLTGTAGPPGLPGEAGPPGPQGPPGPQGINGLSIPGEPGRQGMKGDAGEPGLPGRSGTPGLSGPPGPVGPPGDRGFTGKDGATGPRGPPGSAGTPGVPGVAGPSGKPGKPGDRGPQGPPGMKGEKGDRGDIASQNMMRAVARQVCEQLINGQMNRFNHMLNQIPNDYYANRNQPGPPGPPGPPGSAGARGEPGPGGRPGFPGTPGDQGPPGERGLPGEKGERGIGSQGPRGLPGPPGPPGESRTGPPGSTGSRGPPGPPGRPGNGGIRGPPGPPGYCDSSQCSSIAYNGQGFPVAYLQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN24</accession>
    <name>A0A452HN24_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Golgi-specific brefeldin A-resistance guanine nucleotide exchange factor 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="4">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Endoplasmic reticulum-Golgi intermediate compartment</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Golgi apparatus</location>
            <location evidence="2">cis-Golgi network</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="6">Golgi apparatus</location>
            <location evidence="6">trans-Golgi network</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="5">Lipid droplet</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Peripheral membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018683.1">
        <property type="protein sequence ID" value="ENSGAGP00000016373.1"/>
        <property type="gene ID" value="ENSGAGG00000012266.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005793">
        <property type="term" value="C:endoplasmic reticulum-Golgi intermediate compartment"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005811">
        <property type="term" value="C:lipid droplet"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010256">
        <property type="term" value="P:endomembrane system organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016197">
        <property type="term" value="P:endosomal transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032012">
        <property type="term" value="P:regulation of ARF protein signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00171">
        <property type="entry name" value="Sec7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.1000.11:FF:000007">
        <property type="entry name" value="Golgi-specific brefeldin A-resistance guanine nucleotide exchange factor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.220.20:FF:000004">
        <property type="entry name" value="Golgi-specific brefeldin A-resistance guanine nucleotide exchange factor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.220.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1000.11">
        <property type="entry name" value="Arf Nucleotide-binding Site Opener,domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056604">
        <property type="entry name" value="GBF1-like_TPR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032691">
        <property type="entry name" value="Mon2/Sec7/BIG1-like_HUS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023394">
        <property type="entry name" value="Sec7_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000904">
        <property type="entry name" value="Sec7_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035999">
        <property type="entry name" value="Sec7_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10663:SF388">
        <property type="entry name" value="GOLGI-SPECIFIC BREFELDIN A-RESISTANCE GUANINE NUCLEOTIDE EXCHANGE FACTOR 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10663">
        <property type="entry name" value="GUANYL-NUCLEOTIDE EXCHANGE FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01369">
        <property type="entry name" value="Sec7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12783">
        <property type="entry name" value="Sec7-like_HUS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23325">
        <property type="entry name" value="TPR_28"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00222">
        <property type="entry name" value="Sec7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48425">
        <property type="entry name" value="Sec7 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50190">
        <property type="entry name" value="SEC7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="13" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="10" id="KW-0344">Guanine-nucleotide releasing factor</keyword>
    <keyword evidence="11" id="KW-0551">Lipid droplet</keyword>
    <keyword evidence="14 17" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="12" id="KW-0653">Protein transport</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="17" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="17" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="7" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="1624"/>
            <end position="1644"/>
        </location>
    </feature>
    <feature type="domain" description="SEC7" evidence="18">
        <location>
            <begin position="693"/>
            <end position="883"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="216"/>
            <end position="257"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="273"/>
            <end position="308"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="324"/>
            <end position="348"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="610"/>
            <end position="637"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1436"/>
            <end position="1483"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1723"/>
            <end position="1778"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="16">
        <location>
            <begin position="227"/>
            <end position="241"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="244"/>
            <end position="257"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="16">
        <location>
            <begin position="273"/>
            <end position="288"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="16">
        <location>
            <begin position="329"/>
            <end position="347"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="614"/>
            <end position="637"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="16">
        <location>
            <begin position="1436"/>
            <end position="1449"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="16">
        <location>
            <begin position="1723"/>
            <end position="1734"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="16">
        <location>
            <begin position="1736"/>
            <end position="1749"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004222"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004399"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004502"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00004601"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022658"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022677"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00069794"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50190"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016373.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1852" mass="206591" checksum="52D2A4AA8DEB4B7B" modified="2019-05-08" version="1">MVDKNIYIVQGEINTVVGAIKRNARWNTHTHLDEERDPLLHSFSLLKEVLNTITELSEIEPNVFLRPFLEVIRSEDTTGPITGLALTSVNKFLSYALIDPSHEGTAEGMENMADAVTHARFVGTDPASDEVVLMKILQVLRTLLLTPVGAHLTNESVCEIMQSCFRICFEMRLSELLRKSAEHTLVDMVQLLFTRLPHFKEEPKSYVGANMKKLKMRAGGMSDSSKWKKQKRSPRPTRHATKVSPGTEQPATNGTVMPSNIIVDGVTFIDAPSPISSGSSENISSAVSPATDSGLELSSQTTSKEDLTDLEQVNSLGLSAAMPSIEPNATEHPHKHEPQNEGLRVEKTQSASVESIPEVLEECTSVAEHSDSASVHDMDYVNPRGVRFTQSSQKEGIALVPYGLPCIRELFRFLISLTNPHDRHNSEVMIHMGLQLLTVGLESAPVANCQSLLGLVKDELCRHLFQLLSVERLNLYAASLRVCFLLFESMREHLKFQLEMYIKKLMEIITVENPKMPYEMKEMALEAIVQLWRIPSFVTELYINYDCDYYCSNLFEDLTKLLSKNAFPVSGQLYTTHLLSLEALLTVIDSTEAHCQAKVLSNIHQQEKEVVKPSTETASSIKEPSNNNERVSEGNSPNVVTELSGACPPTSGCLMADQMKQGCQDLEGGSDAAEKNIPRKPTRFSCLLPRTQELMEIKSKKKLLITGTEQFNQKPRKGIQFLQEKNLLAIPMDNSEVAQWLRENPRLDKKMIGEFVSDRKNIDLLESFVGTFNFQGLRLDEALRLYLEAFRLPGEAPVIQRLLEAFTEHWRKSNGSPFANSDACFALAYAVIMLNTDQHNHNVRKQNVPMTLEEFRKNLKGVNGGKDFEQDILEDMYHAIKNDEIVMPEEQTGLVKENYIWNVLLHRGATDEGIFLHMPPGSYDHDLFTMTWGPTIAALSYVFDKSLDETIIQKAISGFRKCAMISAHYGLSDVFDNLIISLCKFTALSSESIENLPSVFGSNPKAHIAAKTVFHLAHRHGDILREGWKNIMEAMLQLFRAELLPKAMVEVEDFVDPNGKISLQREETPSNRGESTVLSFVSWLTLSGTEQSGMRGPSTENQEAKRMALECIKQCDPEKLITESKFLQLESLQELMKALISVTPDEETYDEEDAAFCLEMLLRIVLENRDRVGCVWQAVRDHLYHLCVHALEFCFLVERAVVGLLRLAIRLLRREEISAQVLLSLRILLMMKPSVLSRVSHQVAYGLHELLKTNAANIHSGDDWYTLFTLLECIGSGVKPPPALQVTARADNDTGAQSDSEISLSYHPSDVSLDRGYTSDSEVYADHSKQGKMHRTVTDMDVVNSGWLVVGKDDIDSPKPMAGLSRQGPSPLVNQYSLTVGLDLGPHDTKSLMKCVESLSFIVRDAAHVTPENFELCVKTIRIFVEASLNGGYKSQEKRVKSHKYDGKSNRFKKKPKENSLRRSRASSQHQSHSHSDEDDDESVPACYHTVSLQLLDLMHTLHTRAATIYSSWAEEQRHLEAAGKKIEADSQTLWSNCWCPLLQGIAWLCCDARRQVRMQALTYLQRALLVHDLQALDALEWESCFNKVLFPLLTKLLENVSPADVGGMEETRMRASTLLSKVFLQHLSPLLSLTTFAALWLTILDFMDKYMHAGSSDLLSEAIPESLKNMLLVMDTAGIFHSADSRTGYSDLWEITWERIDCFLPRLRDELFKQTVIQEPVPSVPAEPHPQKPVAPALSASAAHSAPLEKSSELGASDSERAPGPGPAIRGTSGSPLSAVASVKMSPVTDVPPHMAQSPLILQPLASPLQVGVPPMTLPIILNPALIEATSPVPLLATARPTDPMTTSEVN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN25</accession>
    <name>A0A452HN25_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Major facilitator superfamily (MFS) profile domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HN25"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018663.1">
        <property type="protein sequence ID" value="ENSGAGP00000016353.1"/>
        <property type="gene ID" value="ENSGAGG00000012116.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008028">
        <property type="term" value="F:monocarboxylic acid transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1250.20:FF:000141">
        <property type="entry name" value="Solute carrier family 16, member 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1250.20">
        <property type="entry name" value="MFS general substrate transporter like domains"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011701">
        <property type="entry name" value="MFS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020846">
        <property type="entry name" value="MFS_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036259">
        <property type="entry name" value="MFS_trans_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050327">
        <property type="entry name" value="Proton-linked_MCT"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11360">
        <property type="entry name" value="MONOCARBOXYLATE TRANSPORTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11360:SF14">
        <property type="entry name" value="MONOCARBOXYLATE TRANSPORTER 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07690">
        <property type="entry name" value="MFS_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103473">
        <property type="entry name" value="MFS general substrate transporter"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50850">
        <property type="entry name" value="MFS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="3" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="3">
        <location>
            <begin position="47"/>
            <end position="70"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="3">
        <location>
            <begin position="82"/>
            <end position="103"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="3">
        <location>
            <begin position="109"/>
            <end position="133"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="3">
        <location>
            <begin position="264"/>
            <end position="282"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="3">
        <location>
            <begin position="302"/>
            <end position="322"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="3">
        <location>
            <begin position="334"/>
            <end position="350"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="3">
        <location>
            <begin position="356"/>
            <end position="379"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="3">
        <location>
            <begin position="391"/>
            <end position="411"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="3">
        <location>
            <begin position="417"/>
            <end position="439"/>
        </location>
    </feature>
    <feature type="domain" description="Major facilitator superfamily (MFS) profile" evidence="4">
        <location>
            <begin position="263"/>
            <end position="459"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="202"/>
            <end position="231"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="211"/>
            <end position="230"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50850"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016353.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="459" mass="50006" checksum="4653435F96038FA7" modified="2019-05-08" version="1">MGMDDSASLLSGPLVTIACGKLGDQLTSIIGAGLVSSGCLISTQATSISFLCVSMGLLLGLGFACLYQAAAVMTARYFKKRLALSNAIARSGMGLTFILAPFTQLLIDFYGWQGTLLIFGGITLNLVPSSMLLRPVIKKLQDFSRNVESGSLTQSKHPATLTSHPDKISTTETQFNIRLDSPMQDSAVTNGFQSAETLGSLVPGKSAGLEGSERETSNRYTPGRDSDKDCSYNNFPLRKEWNSNSRPVPSRQPLLDFSLLKDPFFCIFVWSFLFSQLAYFIPTFHLAARAKTLGIDSMDASYLISVAGITETVSQLLSGWIADQNWIKKYHYHKTYLILCGITNLLGPLATSFPLLMVYTVSFAIFCGGYLALLLPVLVSKGSYQEVIRSLGFASFFAGIAAIAGPPIAGWLSDYTQTYACSFLFAGTCYLIAPISFFFEPSAHKWKLQKEKELNEVTS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN26</accession>
    <name>A0A452HN26_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">SEC7 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Golgi apparatus</location>
            <location evidence="1">trans-Golgi network</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018686.1">
        <property type="protein sequence ID" value="ENSGAGP00000016376.1"/>
        <property type="gene ID" value="ENSGAGG00000012266.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032012">
        <property type="term" value="P:regulation of ARF protein signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016192">
        <property type="term" value="P:vesicle-mediated transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd00171">
        <property type="entry name" value="Sec7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.1000.11:FF:000021">
        <property type="entry name" value="Golgi-specific brefeldin A-resistance factor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.220.20:FF:000004">
        <property type="entry name" value="Golgi-specific brefeldin A-resistance guanine nucleotide exchange factor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.220.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1000.11">
        <property type="entry name" value="Arf Nucleotide-binding Site Opener,domain 2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056604">
        <property type="entry name" value="GBF1-like_TPR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032691">
        <property type="entry name" value="Mon2/Sec7/BIG1-like_HUS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023394">
        <property type="entry name" value="Sec7_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000904">
        <property type="entry name" value="Sec7_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035999">
        <property type="entry name" value="Sec7_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10663:SF388">
        <property type="entry name" value="GOLGI-SPECIFIC BREFELDIN A-RESISTANCE GUANINE NUCLEOTIDE EXCHANGE FACTOR 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10663">
        <property type="entry name" value="GUANYL-NUCLEOTIDE EXCHANGE FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01369">
        <property type="entry name" value="Sec7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12783">
        <property type="entry name" value="Sec7-like_HUS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23325">
        <property type="entry name" value="TPR_28"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00222">
        <property type="entry name" value="Sec7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48425">
        <property type="entry name" value="Sec7 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50190">
        <property type="entry name" value="SEC7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="3" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="3">
        <location>
            <begin position="1582"/>
            <end position="1602"/>
        </location>
    </feature>
    <feature type="domain" description="SEC7" evidence="4">
        <location>
            <begin position="693"/>
            <end position="871"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="216"/>
            <end position="257"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="273"/>
            <end position="308"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="324"/>
            <end position="348"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="610"/>
            <end position="637"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1394"/>
            <end position="1441"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1681"/>
            <end position="1736"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="2">
        <location>
            <begin position="227"/>
            <end position="241"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="244"/>
            <end position="257"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="273"/>
            <end position="288"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="329"/>
            <end position="347"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="614"/>
            <end position="637"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="1394"/>
            <end position="1407"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="2">
        <location>
            <begin position="1681"/>
            <end position="1692"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="1694"/>
            <end position="1707"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004601"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50190"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016376.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1810" mass="201961" checksum="C3A6CE289C208B00" modified="2019-05-08" version="1">MVDKNIYIVQGEINTVVGAIKRNARWNTHTHLDEERDPLLHSFSLLKEVLNTITELSEIEPNVFLRPFLEVIRSEDTTGPITGLALTSVNKFLSYALIDPSHEGTAEGMENMADAVTHARFVGTDPASDEVVLMKILQVLRTLLLTPVGAHLTNESVCEIMQSCFRICFEMRLSELLRKSAEHTLVDMVQLLFTRLPHFKEEPKSYVGANMKKLKMRAGGMSDSSKWKKQKRSPRPTRHATKVSPGTEQPATNGTVMPSNIIVDGVTFIDAPSPISSGSSENISSAVSPATDSGLELSSQTTSKEDLTDLEQVNSLGLSAAMPSIEPNATEHPHKHEPQNEGLRVEKTQSASVESIPEVLEECTSVAEHSDSASVHDMDYVNPRGVRFTQSSQKEGIALVPYGLPCIRELFRFLISLTNPHDRHNSEVMIHMGLQLLTVGLESAPVANCQSLLGLVKDELCRHLFQLLSVERLNLYAASLRVCFLLFESMREHLKFQLEMYIKKLMEIITVENPKMPYEMKEMALEAIVQLWRIPSFVTELYINYDCDYYCSNLFEDLTKLLSKNAFPVSGQLYTTHLLSLEALLTVIDSTEAHCQAKVLSNIHQQEKEVVKPSTETASSIKEPSNNNERVSEGNSPNVVTELSGACPPTSGCLMADQMKQGCQDLEGGSDAAEKNIPRKPTRFSCLLPRTQELMEIKSKKKLLITGTEQFNQKPRKGIQFLQEKNLLAIPMDNSEVAQWLRENPRLDKKMIGEFVSDRKNIDLLESFVGTFNFQGLRLDEALRLYLEAFRLPGEAPVIQRLLEAFTEHWREFRKNLKGVNGGKDFEQDILEDMYHAIKNDEIVMPEEQTGLVKENYIWNVLLHRGATDEGIFLHMPPGSYDHDLFTMTWGPTIAALSYVFDKSLDETIIQKAISGFRKCAMISAHYGLSDVFDNLIISLCKFTALSSESIENLPSVFGSNPKAHIAAKTVFHLAHRHGDILREGWKNIMEAMLQLFRAELLPKAMVEVEDFVDPNGKISLQREETPSNRGESTVLSFVSWLTLSGTEQSGMRGPSTENQEAKRMALECIKQCDPEKLITESKFLQLESLQELMKALISVTPDEETYDEEDAAFCLEMLLRIVLENRDRVGCVWQAVRDHLYHLCVHALEFCFLVERAVVGLLRLAIRLLRREEISAQVLLSLRILLMMKPSVLSRVSHQVAYGLHELLKTNAANIHSGDDWYTLFTLLECIGSGVKPPPALQVTARADNDTGAQSDSEISLSYHPSDVSLDRGYTSDSEVYADHSKQGKMHRTVTDMDVVNSGWLVVGKDDIDSPKPMAGLSRQGPSPLVNQYSLTVGLDLGPHDTKSLMKCVESLSFIVRDAAHVTPENFELCVKTIRIFVEASLNGGYKSQEKRVKSHKYDGKSNRFKKKPKENSLRRSRASSQHQSHSHSDEDDDESVPACYHTVSLQLLDLMHTLHTRAATIYSSWAEEQRHLEAAGKKIEADSQTLWSNCWCPLLQGIAWLCCDARRQVRMQALTYLQRALLVHDLQALDALEWESCFNKVLFPLLTKLLENVSPADVGGMEETRMRASTLLSKVFLQHLSPLLSLTTFAALWLTILDFMDKYMHAGSSDLLSEAIPESLKNMLLVMDTAGIFHSADSRTGYSDLWEITWERIDCFLPRLRDELFKQTVIQEPVPSVPAEPHPQKPVAPALSASAAHSAPLEKSSELGASDSERAPGPGPAIRGTSGSPLSAVASVKMSPVTDVPPHMAQSPLILQPLASPLQVGVPPMTLPIILNPALIEATSPVPLLATARPTDPMTTSEVN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN27</accession>
    <name>A0A452HN27_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">C2H2-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">May be involved in transcriptional regulation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HN27"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018708.1">
        <property type="protein sequence ID" value="ENSGAGP00000016393.1"/>
        <property type="gene ID" value="ENSGAGG00000012269.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010468">
        <property type="term" value="P:regulation of gene expression"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001155">
        <property type="entry name" value="Zinc finger 30C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000688">
        <property type="entry name" value="zinc finger protein 197 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002716">
        <property type="entry name" value="Zinc finger protein 212"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002343">
        <property type="entry name" value="Zinc finger protein 33A"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000135">
        <property type="entry name" value="Zinc finger protein 358"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002274">
        <property type="entry name" value="Zinc finger protein 432"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000060">
        <property type="entry name" value="zinc finger protein 436"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000384">
        <property type="entry name" value="Zinc finger protein 550"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="12"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056438">
        <property type="entry name" value="Znf-C2H2_CTCF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050758">
        <property type="entry name" value="Znf_C2H2-type"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23234">
        <property type="entry name" value="ZNF44 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23611">
        <property type="entry name" value="zf-C2H2_16"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="10"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="10" id="KW-0804">Transcription</keyword>
    <keyword evidence="8" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="7" id="KW-0862">Zinc</keyword>
    <keyword evidence="6 12" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="156"/>
            <end position="183"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="184"/>
            <end position="211"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="230"/>
            <end position="257"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="258"/>
            <end position="285"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="286"/>
            <end position="313"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="314"/>
            <end position="341"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="342"/>
            <end position="369"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="370"/>
            <end position="397"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="415"/>
            <end position="442"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="443"/>
            <end position="470"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00003767"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016393.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="551" mass="61275" checksum="1940A87A0BED3947" modified="2019-05-08" version="1">LLGFHPQQCLCMGSYGVLFSAHGQGPLHGNAAAGCCGGRFGLSNGRSRSSSEEQHPDEGAETLQPYRVLFRGSEEGAVLSPELQGTALRSCGSQRPESELGATQGCDWCSSNSPNMRSGPETGEGPSPCLVCAPRANGTAPPGVQESSVPKKEKPHKCPECGKAFRRKWDLTKHQRTHSGERPYPCTDCSKSFSHVSNLIKHQRIHTGERPFACNLCGRSFTLKQVLRPFACTDCDKSFKDKQKLTIHQRIHTGERPYVCADCGKSFRQSQRLKTHRRVHTGEKPYLCSDCGRAFSQVSNLYTHWRTHTGERPYPCAYCGKSFSLKHDLTKHQRVHTGERPYPCTDCNKSFSQKQDLTKHQRVHTGERPYPCAQCGKSFSNVSNLLAHQRTHLGQRLPLAHLAAHQRTHAAERPYPCGTCEKRFKQSQDLRKHQRTHTGERPYPCTECGKCFSHVSNLNAHRKVHMGARPHTCKDCGKGFRPSAGTVLHWGYSALQPHCTGLSKGPQGTPTSSLCTARPKASPCPYCPCWPETRAQPLGNENRDTEKWGRD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HN29</accession>
    <name>A0A452HN29_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Collagen alpha-1(XII) chain</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">Type XII collagen interacts with type I collagen-containing fibrils, the COL1 domain could be associated with the surface of the fibrils, and the COL2 and NC3 domains may be localized in the perifibrillar matrix.</text>
    </comment>
    <comment type="subunit">
        <text evidence="14">Trimer of identical chains each containing 190 kDa of non-triple-helical sequences.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="12">Belongs to the fibril-associated collagens with interrupted helices (FACIT) family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018694.1">
        <property type="protein sequence ID" value="ENSGAGP00000016380.1"/>
        <property type="gene ID" value="ENSGAGG00000011657.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005581">
        <property type="term" value="C:collagen trimer"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005614">
        <property type="term" value="C:interstitial matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035987">
        <property type="term" value="P:endodermal cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="18"/>
    </dbReference>
    <dbReference type="CDD" id="cd01482">
        <property type="entry name" value="vWA_collagen_alphaI-XII-like"/>
        <property type="match status" value="4"/>
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        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
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    <keyword evidence="6" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="8" id="KW-0176">Collagen</keyword>
    <keyword evidence="9" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11" id="KW-0379">Hydroxylation</keyword>
    <keyword evidence="7" id="KW-0654">Proteoglycan</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
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            <begin position="26"/>
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        </location>
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            <begin position="906"/>
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            <begin position="997"/>
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            <begin position="1087"/>
            <end position="1176"/>
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            <begin position="1196"/>
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            <begin position="1384"/>
            <end position="1473"/>
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        <location>
            <begin position="1474"/>
            <end position="1565"/>
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    <feature type="domain" description="Fibronectin type-III" evidence="18">
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            <begin position="1566"/>
            <end position="1655"/>
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            <begin position="1656"/>
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            <begin position="1747"/>
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            <begin position="2201"/>
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            <begin position="2317"/>
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            <begin position="1070"/>
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            <begin position="1730"/>
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            <begin position="2874"/>
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            <begin position="2883"/>
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            <begin position="2899"/>
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            <begin position="2949"/>
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            <dbReference type="ARBA" id="ARBA00004498"/>
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        <source>
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        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
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    <sequence length="3008" mass="328955" checksum="20014630F3FA016C" modified="2019-05-08" version="1">MLYAYIGLCRIRALGAQFLGPSIVNPPSDLNFTIIDEHTVRMSWTRPPDAIEGYKITVVPTTDGPTKEFTLAPSVTDTILSDLTPDTEYVVTITSYDDREESLPVFGQLTIQTGVAVGPEDKKPDQSHIKKCSVSALTDLVFLVDGSWSVGRSNFRYILDFMATLVSAFDIGEEKTRIGVVQYSTDTRTEFNLNQYFRRNDLLNAIKGIPYKGGNTMTGEAIDYLIRNAFTESAGARKGIPKVAIIITDGKSQDEVEIPARELRNVGVEVFSLGIKAADAKELKLMASQPSLKHVFNVANFDGIVDIQNEIISQVCSGVDEQLGELVSGEEAVEPPSNLVATQISSKSIRVTWDPSASQITGYRVQFFPMIAGAKQHVLSVGPQTTALNVRDLSPETEYQINVYAMRGMAASEPITIMEKTQPVKVQVECSRGVDVKADIVLLVDGSYSIGIANFVKVRAFLEVLVKSFEISPQKVQISLVQYSRDPYTEFYLNRYNKIEDIIQAIHTFPYRGGSTNTGKAMTYVREKVFVINRGSRPNVPKVMILITDGKSSDAFKEPAIKLRESDVEIFAVGVKDAVRTELEAIASPPAETHVYTVEDFDAFQRISFELTQSVCLRIEQELAAIRKKSYLPARNLAFSEVTSDSFKVNWSPAGSEVLSYLIKYKVAAGGEEFLVSVPASSTSSVLTHLLPETLYSVSVIAEYEDGDGPSLDGEETTLEVKGAPRNLMITDETTDSFKVGWTPAPGNVLRYRVEYRPVTGRERKHVTVPANERSTTLLNLTPDTRYEVYVIPEYQSGPGNSLNGYAKTEEVLGTPRNLRVSDATTSTMKLSWSAAPGKVQRYLVTYTPVAGGETKEVTIRGDTTTTVLRDLEHGTRYALSVTALYASGAGDALSGQGETPEDRGSPRDLLIRDITDTTMGLSWTAAPGAVRRYRIVWRSLYDGQTGESTVPGNVINTVLENLQPETKYKISVIASYSSGDGAPLEGEATTEVSPNARRVRVDEETENSMRVTWQPAPGKVVSYRVVYRPVSGGRQVVAKVPATSTATVLKRLQPLTKYNISVIPMYKSGEGKHRQGEGTTASPYKPPRNLKTSDSTMSSFRVTWEPAPGEVRGYKITFHPTGEDRLGELIVGPYDNTVVLEELRAGTTYKVNVFGMFEGGESAPLVGQEMTTLSDVTVMPFLARGLECTTRAEADIVLLVDGSWSIGRPNFKTVRNFIARIVEVFDIGPDKVQIALAQYSGDPRTEWHLNTYGSKQSLLDAVANLPYKGGNTLTGMALSFILKDNFKPEVGLRPKARKIGVLITDGKSQDDIVAPSRRLRDEGVELYAIGIKNADENELKQIATDPDDTHVFNVADFSFLTNIVDDLTINLCNSVKGPGGLSSPSNLVTSEVTPRSFRVTWTAPVESVDRYRVEYYPVSGGPPQQFYVNRVETTTVLKNLKPETEYVVKVFSVVEDESSEPLSGTETTLPIPTIRNLNAYDIGSTSMRVRWEPLRGATGYRLLYEPLNATIPAVEKEMRVGSTVNDVQLVDLIPNAAYTLTIHAEFGDLSSDPLTTQEVTLPLAGAKSLRVRDITHSSMNVIWDPAPGKVRKYLLRYKTTLGDEPKEVEVDKSKTSTVLPSLLSQTLYDLQLVAVYDEGESIPVAAKATTLPVPAPVNLRLTEVTKQSFRGTWDHGAPDVALYRITWGPYGGTEKQETILNGDENSLVFENLNPDTLYDVSVTAIYPDESESDDLIGSERTSPKSGPRNLQVYNATSNSLTVKWDPASGRVQRYRITYQRTTGDGAEQSTMVGGRQNSVVLQKLQPDTPYTITVSSMYADGEGGRMTGRGKTKPLNTVKNLRVYDPTTSTLSVRWDHAEGSPRQYKVYYVPTTGGAEEMVIILVGNTNYIILRSLQPDTPYKITVVPTYPEGDGGRVSDTGRTLVRGAARNIQVYNPTPNTLNVRWEPAPGPVQQYRVVYAPLVGPRPSESIVVPANTRDVVLERLTPDTPYSVNVVALYADGEGNQSSGQGRTTPRSGPRNLKVFDPTTNSLSVQWDHADGPVQQYRIIYSPTVGDPIDEYTTVPGRRNNVILQPLHSDTPYKITVVAVYDDGDGGQLTGNGRTVGLLPPQNIHISDEWYTRFRVSWDPSPSPVLGYKLAYKPVGSAEPPMEVFVGEVTSYTLHNLSPSTIYDVNVYAQYDSGLSAPLIDQGTTLYLNVTDLTSYKVGWDTFCIRWSPHRSASSYRLKLNPSDGSRGQEITVRGTETSHCFTGLSPDTEYNATVFVQTPNLEGPPISLKERTFIKPTEAPTQPPPPPPPPTIPPARDVCKGAKADIVFLTDASWSIGDDNFNKVVKFVFNTVGAFDLINPAGIQVSFVQYSDDAKSEFKLNTYDDKAQALGALQNIRYKGGNTRTGKALTFINEKVLTWESGMRRGVPKVLVVVTDGRSQDEVKKAAAIIQHSGFSIFVVGVADVDYHELSKIASKPSERHVFIVDDFDAFEKIEENLLTFICETATSTCPLIYLEGYTSPGFKMLESYNLTEKHFASVQGVSLESGSFPSHVAYRLHRNAFISQPASEIHPDGLPHAYTIILLFRLLSETPNEPFAIWQITDRDFKPQVGVILDPSTKLLSFFNKDTRGEVQTVTFDDNEVKKIFYGSFHKVHIVVTSTSVKIYIDCSEVMEKPIKESGNITTDGFEILGKLVKGDRKSATFEIQSFDIVCNAVWTSRDRCCDLPSMRDEAKCPALPNACTCTQDSVGPPGPPGPAKPWFNIFLILCFKGPPGLPGEAGPPGPQGPPGPQGINGLSIPGEPVSNHSVSVSETVWRQRTDTSWERSPKISDHHGPPGMKGEKGDRGDIASQNMMRAVARQVCEQLINGQMNRFNHMLNQIPNDYYANRNQPGPPGPPGPPGSAGARGEPGPGGRPGFPGTPGDQGPPGERGLPGEKGERGIGSQGPRGLPGPPGPPGESRTGPPGSTGSRGPPGPPGRPGNGGIRGPPGPPGYCDSSQCSSIAYNGQGFPGRLLTT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPN0</accession>
    <name>A0A452IPN0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">VPS33A core subunit of CORVET and HOPS complexes</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the STXBP/unc-18/SEC1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPN0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033842.1">
        <property type="protein sequence ID" value="ENSGAGP00000029803.1"/>
        <property type="gene ID" value="ENSGAGG00000021422.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016192">
        <property type="term" value="P:vesicle-mediated transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.830.10:FF:000006">
        <property type="entry name" value="Vacuolar protein sorting-associated protein 33A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1910:FF:000005">
        <property type="entry name" value="vacuolar protein sorting-associated protein 33A isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1910:FF:000008">
        <property type="entry name" value="vacuolar protein sorting-associated protein 33A isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1910">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.2060">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043154">
        <property type="entry name" value="Sec-1-like_dom1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001619">
        <property type="entry name" value="Sec1-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027482">
        <property type="entry name" value="Sec1-like_dom2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036045">
        <property type="entry name" value="Sec1-like_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11679">
        <property type="entry name" value="VESICLE PROTEIN SORTING-ASSOCIATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00995">
        <property type="entry name" value="Sec1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56815">
        <property type="entry name" value="Sec1/munc18-like (SM) proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0653">Protein transport</keyword>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0813">Transport</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00009884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029803.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="5">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="493" mass="55969" checksum="D64C71238647FDD3" modified="2019-05-08" version="1">MFTLKAGRLPPADVKNIIFLVRPRLELMDIIAENVLSEDRVRSPLRDFHILFVPRRSLLCEQRLKDQGVLGSFIHREQYSLDLIPFDGDLLSMESESAFKECYLENDQTSLYQAAKGLMTLQALYGTIPQIFGKGDCARHVANMMIRMKREFPGGQNSIFPIFDTLLLLDRNVDLLSPLATQLTYEGLIDEIYGIQNTYVKLPPEKFAPKKQGEGGKDLPTEPKKLQLNSAEELYAEIRDKNFNAVGSVLSKKAKIISAAFEERHHAKTVGEIKQFVSQLPHMQAARSSLANHTSIAELIKDITTSEDFFDNLTVEQEFMSGIDTDKTYGYEHILTLNNLEKAGLLKLQTSSRNNYPTIRKTMRLWMDDVNEQNPNDISYVYSGYAPLSVRLAQLLARPGWRSIEEVLKMLPGPHFEERQQLPTGLQKKRQQGENRVTLVFFLGGVTYAEIAALRFLSQMEDGGTEYVIATTKLINGTNWIKSLMEKLEPPPF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPN1</accession>
    <name>A0A452IPN1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Arrestin domain-containing protein 3</fullName>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">Adapter protein that plays a role in regulating cell-surface expression of adrenergic receptors and probably also other G protein-coupled receptors. Plays a role in NEDD4-mediated ubiquitination and endocytosis af activated ADRB2 and subsequent ADRB2 degradation. May recruit NEDD4 to ADRB2. Alternatively, may function as adapter protein that does not play a major role in recruiting NEDD4 to ADRB2, but rather plays a role in a targeting ADRB2 to endosomes.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cell membrane</location>
            <topology evidence="3">Peripheral membrane protein</topology>
            <orientation evidence="3">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Early endosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Lysosome</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the arrestin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPN1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033909.1">
        <property type="protein sequence ID" value="ENSGAGP00000029864.1"/>
        <property type="gene ID" value="ENSGAGG00000021451.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005769">
        <property type="term" value="C:early endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005764">
        <property type="term" value="C:lysosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.640:FF:000005">
        <property type="entry name" value="Arrestin domain-containing protein 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.640:FF:000007">
        <property type="entry name" value="Arrestin domain-containing protein 3 mRNA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.640">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014752">
        <property type="entry name" value="Arrestin-like_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011021">
        <property type="entry name" value="Arrestin-like_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011022">
        <property type="entry name" value="Arrestin_C-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050357">
        <property type="entry name" value="Arrestin_domain-protein"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014756">
        <property type="entry name" value="Ig_E-set"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11188">
        <property type="entry name" value="ARRESTIN DOMAIN CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11188:SF49">
        <property type="entry name" value="ARRESTIN DOMAIN-CONTAINING PROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02752">
        <property type="entry name" value="Arrestin_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00339">
        <property type="entry name" value="Arrestin_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01017">
        <property type="entry name" value="Arrestin_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81296">
        <property type="entry name" value="E set domains"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9" id="KW-0967">Endosome</keyword>
    <keyword evidence="11" id="KW-0458">Lysosome</keyword>
    <keyword evidence="10" id="KW-0472">Membrane</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="Arrestin C-terminal-like" evidence="15">
        <location>
            <begin position="183"/>
            <end position="310"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="389"/>
            <end position="410"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="401"/>
            <end position="410"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004371"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004412"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004413"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00005298"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023228"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00041132"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00045942"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="SMART" id="SM01017"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029864.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="410" mass="45668" checksum="7D3EA305ED97E5C4" modified="2019-05-08" version="1">NGGETGGKVKSLTISFDCLNDSNVPVYSSGDTVSGRVNLEVTGEIRVKSLKIHARGHAKVRWTESRNAGSNTAYTQNYTEEVEYFNHKDILIGHERGLNTIHSGRHEYAFSFELPQTPLATSFEGRHGSVRYWVKAELHRPWLLPVKLKKEFTVFEHIDINTPSLLSPQAGTKEKTLCCWFCTSGPISLSAKIERKGFTPGESIQIFAEIENCSSRMVVPKAAIYQTQAFYAKGKMKEVKQLVANLRGESLSSGKTETWNGKQLKIPPVSPSILDCSIIRVEYSLMVYVDIPGAMDLFLNLPLVIGTIPLHPFGSRTSSVSSQCSMNMNWLGLPLPERPEAPPSYAEVVTEEQRQSSLVPIAACDDFERALQGPLFAYIQEFRFLPPPLYSEIDPNPDQPTDDRPSCPSR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPN3</accession>
    <name>A0A452IPN3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Mediator of RNA polymerase II transcription subunit 23</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="7">Mediator complex subunit 23</fullName>
        </alternativeName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the Mediator complex subunit 23 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPN3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033953.1">
        <property type="protein sequence ID" value="ENSGAGP00000029900.1"/>
        <property type="gene ID" value="ENSGAGG00000021033.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016592">
        <property type="term" value="C:mediator complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005667">
        <property type="term" value="C:transcription regulator complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010628">
        <property type="term" value="P:positive regulation of gene expression"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021629">
        <property type="entry name" value="Mediator_Med23"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12691">
        <property type="entry name" value="MEDIATOR OF RNA POLYMERASE II TRANSCRIPTION SUBUNIT 23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12691:SF10">
        <property type="entry name" value="MEDIATOR OF RNA POLYMERASE II TRANSCRIPTION SUBUNIT 23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11573">
        <property type="entry name" value="Med23"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0539">Nucleus</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0804">Transcription</keyword>
    <keyword evidence="4" id="KW-0805">Transcription regulation</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010222"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00019696"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00031961"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029900.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1227" mass="140808" checksum="7B04026F5BE75A1E" modified="2019-05-08" version="1">MRIKQSRRHTVFRGLNLFLLQKTEVIEEAFPGMFMDTPEDERTKLISCLGAFRQFWSSLSQESHEQCVQWIVRFIHSQHSPKRISFLYDCLAMAVETGLLPPRMVCESLINSETLEWERTQLWALTFKLVRKIIGGVDYKGVRDLLKVILEKILTIPNTVSSAVVQQLLAAREVVAYILERNACLLPAYFAVTEIRKLYPEGKLPHWLLGNLVSDFVDTFRPTARINSICGRCSLLPVVNNSGAMCNSWKLDPTTLRFPLKGLLPYDKDLFEPQTALLRYVLEQPYSRDMVCNMLGLNKQHKQRCPVLEDQLVDLVVYAMERSETEEKFDDGGTSQLLWQHLSSQLIFFVLFQFASFPHMVLSLHQKLAGRGLIKGRDHLMWVLLQFISGSIQKNALADFLPVMKLFDLLYPEKECIPVPDINKPQSTHAFAMTCIWIHLNRKAHSDNSKLQIPIPHSLKLHHEFLQQSLRNKSLQMNDYKIALLCNAYSTNSECFTLPMGVLVETIYGNGNMRIPLPGTNCMASGSITPLPMNLLDSLTVHAKMSLIHSIATRVIKLAHAKSSLALAPALVETYSRLLVYMEIESLGIKGFISQLLPTVFKSHAWGILHTLLEMFSYRMHHIQPHYRVQLLSHLHSLAAVPQTNQNQLHLCVESTALRLITALGSSEVQPQFTRFLSDPKTVLSAESEELNRALILTLARATHVTDFFTGSDSIQGTWCKDILQTIMSFTPHNWASHTLSCFPAPLQAFFKQNNVPQESRFNLKKNVEEEYRKWKSMTNENDIITHFSMQGSPPLFLCLLWKMLLETDHINQIGYRVLERIGARALVAHVRTFADFLVYEFSTSAGGQQLNKCIEILNDMVWKYNIVTLDRLILCLAMRSHEGNEAQVCYFIIQLLLLKPNDFRNRVSDFVKENSPEHWLQNDWHTKHMSYHKKYPEKLYFEGLAEQVNPPVQIQPQYLPIYFGNVCLRFLPVFDIVIHRFLELLPVSKSLETLLDHLGGLYKFHDRPVTYLYNTLHYYERHLRERTNLKRKLVHAIIGSLKDNRPQGWCLSDTYIKSAMNVREDNPWIPDDTYYCKLIGRLSPGPFPNCDWRFNEFPNPAAHALHVTCVELMALAVPGKDVGNALLNVVLKSQPLVPRENITAWMNAIGLIITALPVSLIGQHSLIPKFLTEVLIPIVKTEFQLLYVYHLVGPFLQRFQQERTRCMIEVKYSSCPGFYSLILSGL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPP8</accession>
    <name>A0A452IPP8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Phosphodiesterase</fullName>
            <ecNumber evidence="10">3.1.4.-</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="6">
            <text>3',5'-cyclic AMP + H2O = AMP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:25277"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:58165"/>
            <dbReference type="ChEBI" id="CHEBI:456215"/>
            <dbReference type="EC" id="3.1.4.53"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="6">
            <dbReference type="Rhea" id="RHEA:25278"/>
        </physiologicalReaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="10">
            <name>a divalent metal cation</name>
            <dbReference type="ChEBI" id="CHEBI:60240"/>
        </cofactor>
        <text evidence="10">Binds 2 divalent metal cations per subunit. Site 1 may preferentially bind zinc ions, while site 2 has a preference for magnesium and/or manganese ions.</text>
    </comment>
    <comment type="pathway">
        <text evidence="1">Purine metabolism; 3',5'-cyclic AMP degradation; AMP from 3',5'-cyclic AMP: step 1/1.</text>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the cyclic nucleotide phosphodiesterase family. PDE4 subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.1.4.-" evidence="10"/>
    <dbReference type="AlphaFoldDB" id="A0A452IPP8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033931.1">
        <property type="protein sequence ID" value="ENSGAGP00000029879.1"/>
        <property type="gene ID" value="ENSGAGG00000021546.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00762">
        <property type="reaction ID" value="UER00747"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004115">
        <property type="term" value="F:3',5'-cyclic-AMP phosphodiesterase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006198">
        <property type="term" value="P:cAMP catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00077">
        <property type="entry name" value="HDc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.1300.10:FF:000001">
        <property type="entry name" value="Phosphodiesterase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1300.10">
        <property type="entry name" value="3'5'-cyclic nucleotide phosphodiesterase, catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003607">
        <property type="entry name" value="HD/PDEase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040844">
        <property type="entry name" value="PDE4_UCR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023088">
        <property type="entry name" value="PDEase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002073">
        <property type="entry name" value="PDEase_catalytic_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036971">
        <property type="entry name" value="PDEase_catalytic_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023174">
        <property type="entry name" value="PDEase_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11347">
        <property type="entry name" value="CYCLIC NUCLEOTIDE PHOSPHODIESTERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18100">
        <property type="entry name" value="PDE4_UCR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00233">
        <property type="entry name" value="PDEase_I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00387">
        <property type="entry name" value="PDIESTERASE1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00471">
        <property type="entry name" value="HDc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109604">
        <property type="entry name" value="HD-domain/PDEase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00126">
        <property type="entry name" value="PDEASE_I_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51845">
        <property type="entry name" value="PDEASE_I_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0114">cAMP</keyword>
    <keyword evidence="4 10" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="3 9 10" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="PDEase" evidence="12">
        <location>
            <begin position="251"/>
            <end position="553"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="17"/>
            <end position="44"/>
        </location>
    </feature>
    <feature type="active site" description="Proton donor" evidence="7">
        <location>
            <position position="327"/>
        </location>
    </feature>
    <feature type="binding site" evidence="8">
        <location>
            <begin position="327"/>
            <end position="331"/>
        </location>
        <ligand>
            <name>AMP</name>
            <dbReference type="ChEBI" id="CHEBI:456215"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="331"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="367"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="8">
        <location>
            <position position="368"/>
        </location>
        <ligand>
            <name>AMP</name>
            <dbReference type="ChEBI" id="CHEBI:456215"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="368"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <label>2</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="368"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="8">
        <location>
            <position position="485"/>
        </location>
        <ligand>
            <name>AMP</name>
            <dbReference type="ChEBI" id="CHEBI:456215"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="485"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="8">
        <location>
            <position position="536"/>
        </location>
        <ligand>
            <name>AMP</name>
            <dbReference type="ChEBI" id="CHEBI:456215"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004703"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009517"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023149"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00033681"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PIRSR" id="PIRSR623088-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PIRSR" id="PIRSR623088-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PIRSR" id="PIRSR623088-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="RuleBase" id="RU363067"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS51845"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029879.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="553" mass="62873" checksum="0A12C2A383A481FC" modified="2019-05-08" version="1">SSFKELKDNMCFVHSFDVENGPSPGRSPLDPQASSSSGLVLHTTFPGHSQRRESFLYRSDSDYDLSPKTMSRNSSLPSEQHGDDLIVTPFAQVLASLRSVRNNFTLLTNLHGTSNKRSPAASQPPVSRVSLQEESYQKLAMETLEELDWCLDQLETIQTYRSVSEMASNKFKRMLNRELTHLSEMSRSGNQVSEYISNTFLDKQNDVEIPSPTQKDREKKKKQQLMTQISGVKKLMHSSSLNNTSISRFGVKTEKEDHLAKELEDLNKWGLNIFNVARYSHSRPLTCIMYAIFQERDLLKTFKISSDTFVTYMMTLEDHYHSDVAYHNSLHAADVAQSTHVLLSTPALDAVFTDLEILAAIFAAAIHDVDHPGVSNQFLINTNSELALMYNDESVLENHHLAVGFKLLQEEHCDIFQNLSKKQRQTLRKMVIDMVLATDMSKHMSLLADLKTMVETKKVTSSGVLLLDNYTDRIQVLRNMVHCADLSNPTKSLELYRQWTDRIMEEFFQQGDKERERGMEISPMCDKHTASVEKSQVLYIGLAPLKTGIFLTL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPP9</accession>
    <name>A0A452IPP9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="23">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 16">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000033984.1">
        <property type="protein sequence ID" value="ENSGAGP00000029925.1"/>
        <property type="gene ID" value="ENSGAGG00000021570.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005884">
        <property type="term" value="C:actin filament"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030027">
        <property type="term" value="C:lamellipodium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016459">
        <property type="term" value="C:myosin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001726">
        <property type="term" value="C:ruffle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000146">
        <property type="term" value="F:microfilament motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030048">
        <property type="term" value="P:actin filament-based movement"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072673">
        <property type="term" value="P:lamellipodium morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007266">
        <property type="term" value="P:Rho protein signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd20884">
        <property type="entry name" value="C1_Myosin-IXb"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd01385">
        <property type="entry name" value="MYSc_Myo9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd17217">
        <property type="entry name" value="RA_Myosin-IXb"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04407">
        <property type="entry name" value="RhoGAP_myosin_IXB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.720:FF:000003">
        <property type="entry name" value="Putative unconventional myosin-IXa"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000008">
        <property type="entry name" value="Putative unconventional myosin-IXa"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.820:FF:000003">
        <property type="entry name" value="unconventional myosin-IXa isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.555.10:FF:000009">
        <property type="entry name" value="unconventional myosin-IXa isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.530:FF:000005">
        <property type="entry name" value="unconventional myosin-IXa isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000013">
        <property type="entry name" value="unconventional myosin-IXa isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.190:FF:000013">
        <property type="entry name" value="unconventional myosin-IXa isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.530:FF:000009">
        <property type="entry name" value="unconventional myosin-IXb isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.190">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.60.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.20.240.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.555.10">
        <property type="entry name" value="Rho GTPase activation protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046349">
        <property type="entry name" value="C1-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000048">
        <property type="entry name" value="IQ_motif_EF-hand-BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046987">
        <property type="entry name" value="Myo9"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036023">
        <property type="entry name" value="MYSc_Myo9"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002219">
        <property type="entry name" value="PKC_DAG/PE"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000159">
        <property type="entry name" value="RA_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046989">
        <property type="entry name" value="RA_Myosin-IXb"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000198">
        <property type="entry name" value="RhoGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028557">
        <property type="entry name" value="RhoGAP_myosin_IXB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029071">
        <property type="entry name" value="Ubiquitin-like_domsf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46184:SF2">
        <property type="entry name" value="UNCONVENTIONAL MYOSIN-IXB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46184">
        <property type="entry name" value="UNCONVENTIONAL MYOSIN-IXB-LIKE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00612">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00788">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00620">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00109">
        <property type="entry name" value="C1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00015">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00314">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00324">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57889">
        <property type="entry name" value="Cysteine-rich domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54236">
        <property type="entry name" value="Ubiquitin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50200">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50238">
        <property type="entry name" value="RHOGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00479">
        <property type="entry name" value="ZF_DAG_PE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50081">
        <property type="entry name" value="ZF_DAG_PE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="15 16" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="11 16" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="12 17" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="3" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="6" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="14 16" id="KW-0505">Motor protein</keyword>
    <keyword evidence="13 16" id="KW-0518">Myosin</keyword>
    <keyword evidence="8 16" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="10" id="KW-0862">Zinc</keyword>
    <keyword evidence="9" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="Ras-associating" evidence="20">
        <location>
            <begin position="15"/>
            <end position="113"/>
        </location>
    </feature>
    <feature type="domain" description="Myosin motor" evidence="22">
        <location>
            <begin position="145"/>
            <end position="940"/>
        </location>
    </feature>
    <feature type="domain" description="Phorbol-ester/DAG-type" evidence="19">
        <location>
            <begin position="1645"/>
            <end position="1694"/>
        </location>
    </feature>
    <feature type="domain" description="Rho-GAP" evidence="21">
        <location>
            <begin position="1713"/>
            <end position="1901"/>
        </location>
    </feature>
    <feature type="region of interest" description="Actin-binding" evidence="16">
        <location>
            <begin position="822"/>
            <end position="844"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1087"/>
            <end position="1140"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1182"/>
            <end position="1221"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1238"/>
            <end position="1356"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1466"/>
            <end position="1494"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1618"/>
            <end position="1639"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1945"/>
            <end position="1976"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1993"/>
            <end position="2037"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="17">
        <location>
            <begin position="1028"/>
            <end position="1055"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="1110"/>
            <end position="1135"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="1182"/>
            <end position="1195"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="1238"/>
            <end position="1249"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="1265"/>
            <end position="1275"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="1304"/>
            <end position="1316"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="1342"/>
            <end position="1356"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="1479"/>
            <end position="1490"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="18">
        <location>
            <begin position="1626"/>
            <end position="1636"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="2024"/>
            <end position="2037"/>
        </location>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <begin position="238"/>
            <end position="245"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50081"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50200"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50238"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029925.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2037" mass="234604" checksum="BEEFF3648F52AFB1" modified="2019-05-08" version="1">MSLKDADSAVCQEKATYNLHIYPQLSTESTTCCRVTATKDSTTSDVIKDVIQILDLDVSKHYVLVEVKESGGEEWVLDLNDSPVHRVLLWPRRAQDEHPQNDGYYFLLQERDTDGTIKYVHMQLVSQEKDARRLVERGFLPWHQENFDDLCNLSNLTETTLLENLKRRFLKHKIYTYAGSILIAINPFKFLPIYNPKYVKMYENHQLGKLEPHIFAIADVAYHTMLKKHVNQCIVISGESGSGKTQSTNFLIHCFTALSQKGYASGVERTILGAGPVLEAFGNAKTAHNNNSSRFGKFIQVNYLENGIVRGAVVEKYLLEKSRLVSQEKDERNYHVFYYLLLGVSEEERKEFHLKQPEDYFYLNQHNLKIEDGEDLRHDFERLKQAMEMVGFLPATKKQIFSVLSAILYLGNVTYKKRATGRDEGLEVGPTEVLDTLSQLLKVKREVLVEVLTKRKTVTANDKLILPYSLSEAITARDSMAKSLYSALFDWIVLRINHALLNKKDMEESVSCLSIGVLDIFGFEDFETNSFEQFCINYANEQLQYYFNQHIFKLEQEEYQSEGIAWHNIDYTDNVTCIHLISKKPTGLFYLLDEESNFPHATNQTLLAKFKQQHEENKFFVGTPVKEPAFIIRHFAGKVKYQIKDFREKNMDYMRPDIVALLRSSDSAYVRELIGMDPVAVFRWAILRAAIRAMAVFAEAGRQRAQKTAGVVRHGPRVPLGELQRANTPIEKVYRCSVLDFSFDCSEDFDINAFEDIIAFYESKKKSRGTKQKQIIPKNLLDSKSLKLIVSMTLHDRTTKSLLHLHKKKKPPSISAQFQNSLNKLLETLGKAEPFFIRCIRSNAEKRELFFDEGLVLQQLRYTGMLETVRIRRSGYSAKYTFQDFIDQFQVLLPQNAKASKEEISAHLNKLKLDKNNYQIGKTKVFMKEAERQMLQDTLHKEVIRKIVLLQSWLRMVLERRHFLRMRQATITLQTCWRSYRIRRALERTNAAIYIQSVWRRYRERESYHQQRRRISLVQALVRGHLQRKRFQQMIVEKQQEMQQAQEGEDSVSQEDGNELVLEQLAVKPEMQLDQGIEQMAAGVDEAQNDGDEKLSQATEKAVPSQKSITDGHEKLTSSREKRESRRQRGLEHNKLQNKHVLFPLKEQSLVCQEEETSAKETPETVQEPNKFTAYKTVLPKGTEEERIPVEENKTLSDTQVPGEILDSDVSFPKQSNESKQEILDDKIIDKLKIQDNQKNQVKGSQSFTCPERPSDFSLHIKKNLTPTRSFQTGDENYGNKNKHRISKVDKDLDSPPSFQIQRYLHDPGELRDKREKWKGKRQSEAGQNDVLSESLDERNRTGQSPQTLLSKKGTTSSLDDIANLAQSVTMIQPSPETMAEEKSNKKQTLQRKSNEQLSLLDLAVSVQLPSQPMDAKSTFKSPLRKLLGKKPDKKYVRESPDVMEEGENLPLLSCILISESVESQKLSEAPSGHPRRPQAGERPVKETSKGKRNRTIKIKKISAVSEKWPASVIREITNANELKYLDEFLLNKINDLNSQKSATECLFFEATEKFRGNIKTMYSVPNGQIHVGYKDLIENYQLLVINLAKEREEKDVKLVLNLFQSLLDEFSRGYAKREESEQHKQSKTQKKKRKQERAIEEHNGHVFTSYQVSIRQSCEQCSSYIWPMEKACLCSTCKLICHKKCMSKTQSCCTSPCGKKNEQGVDSRHFGVCVSSLISDKNSVPVVMEKLLEYVEMHGLYTEGIYRKSGSANRMRELKQLLQTDPNSVKLENYPIHTITGILKQWLRELPDPLMTFAQYSDFLRAVELPEKQEQLIAIYSVLEQLPQANHNTLERLIFHLVKVALIEDVNRMSPNALAIVFAPCLLRCPDNSDPLTSMKDVSKTTMCVEMLIKEQMRKYKIKMDEINQLEAAESIAFRRLSLLRQNTLWPVKLGFSSPYEGVLSKSPQDKGNDDSFPELGSLQEEDEVSDVDNREKEILIERIQSIKEEKEDITYRLPELDQQGSDEENLDSETSVSTESLPEDRTGRLDTEGQY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPQ1</accession>
    <name>A0A452IPQ1_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="21">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="15">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
        <text evidence="15">Binds 1 zinc ion per subunit.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="17">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPQ1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033864.1">
        <property type="protein sequence ID" value="ENSGAGP00000029825.1"/>
        <property type="gene ID" value="ENSGAGG00000021496.1"/>
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    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
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        <property type="term" value="C:extracellular matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
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    <dbReference type="GO" id="GO:0004222">
        <property type="term" value="F:metalloendopeptidase activity"/>
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        <property type="project" value="InterPro"/>
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    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
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        <property type="entry name" value="ZnMc_ADAMTS_like"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="A disintegrin and metalloproteinase with thrombospondin motifs 1"/>
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        <property type="match status" value="1"/>
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        <property type="entry name" value="Collagenase (Catalytic Domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Thrombospondin type-1 (TSP1) repeat"/>
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        <property type="entry name" value="ADAMTS/ADAMTS-like"/>
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        <property type="entry name" value="ADAMTS_ADAMTS-like"/>
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        <property type="entry name" value="ADAMTS_CR_2"/>
    </dbReference>
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        <property type="entry name" value="ADAMTS_CR_3"/>
    </dbReference>
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        <property type="entry name" value="ADAMTS_spacer1"/>
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        <property type="entry name" value="MetalloPept_cat_dom_sf"/>
    </dbReference>
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        <property type="entry name" value="Peptidase_M12B"/>
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        <property type="entry name" value="Peptidase_M12B_N"/>
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        <property type="entry name" value="PLAC"/>
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        <property type="entry name" value="TSP1_rpt"/>
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        <property type="entry name" value="A DISINTEGRIN AND METALLOPROTEINASE WITH THROMBOSPONDIN MOTIFS 3"/>
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    </dbReference>
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        <property type="match status" value="1"/>
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        <property type="match status" value="1"/>
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        <property type="entry name" value="Pep_M12B_propep"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01421">
        <property type="entry name" value="Reprolysin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19030">
        <property type="entry name" value="TSP1_ADAMTS"/>
        <property type="match status" value="3"/>
    </dbReference>
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        <property type="entry name" value="TSP_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01857">
        <property type="entry name" value="ADAMTSFAMILY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00209">
        <property type="entry name" value="TSP1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55486">
        <property type="entry name" value="Metalloproteases ('zincins'), catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF82895">
        <property type="entry name" value="TSP-1 type 1 repeat"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50215">
        <property type="entry name" value="ADAM_MEPRO"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="PLAC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50092">
        <property type="entry name" value="TSP1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="15" id="KW-0106">Calcium</keyword>
    <keyword evidence="12 16" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="5 15 17" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0482">Metalloprotease</keyword>
    <keyword evidence="4" id="KW-0645">Protease</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="9 15 17" id="KW-0862">Zinc</keyword>
    <keyword evidence="11" id="KW-0865">Zymogen</keyword>
    <feature type="domain" description="Peptidase M12B" evidence="19">
        <location>
            <begin position="194"/>
            <end position="398"/>
        </location>
    </feature>
    <feature type="domain" description="PLAC" evidence="20">
        <location>
            <begin position="959"/>
            <end position="997"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1062"/>
            <end position="1103"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="1080"/>
            <end position="1103"/>
        </location>
    </feature>
    <feature type="active site" evidence="14 17">
        <location>
            <position position="337"/>
        </location>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="197"/>
        </location>
        <ligand>
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="197"/>
        </location>
        <ligand>
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
            <label>2</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="289"/>
        </location>
        <ligand>
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15 17">
        <location>
            <position position="336"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <note>catalytic</note>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15 17">
        <location>
            <position position="340"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <note>catalytic</note>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15 17">
        <location>
            <position position="346"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <note>catalytic</note>
        </ligand>
    </feature>
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        <location>
            <position position="393"/>
        </location>
        <ligand>
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="396"/>
        </location>
        <ligand>
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
            <label>2</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="396"/>
        </location>
        <ligand>
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="271"/>
            <end position="320"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="314"/>
            <end position="393"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="353"/>
            <end position="379"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="420"/>
            <end position="445"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="431"/>
            <end position="454"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="440"/>
            <end position="473"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="467"/>
            <end position="478"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="501"/>
            <end position="538"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="505"/>
            <end position="543"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="516"/>
            <end position="528"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
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        </source>
    </evidence>
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    </evidence>
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        <source>
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        </source>
    </evidence>
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    </evidence>
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        </source>
    </evidence>
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    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
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    </evidence>
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        <source>
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    </evidence>
    <evidence type="ECO:0000256" key="18">
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        <source>
            <dbReference type="PROSITE" id="PS50215"/>
        </source>
    </evidence>
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    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
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        </source>
    </evidence>
    <sequence length="1103" mass="124670" checksum="138487579D05DC9C" modified="2019-05-08" version="1">LLPTVLPRLYGLVTPISTSADGSFISNILSANHQRRLTRDVSQSPKQLYFNITAFGREFHLRLRPNTHLVAPEAVVEWYEDSVEKPLWTNCAYVGDITDIPGASVAISNCDGLAGMIRTDKDEYFIEPLERGKQMEEERGRIHMVYRRSAVTQDPGAPLVLDLGLDGLDSMYSNIEQQLNETLRRRRHAGDNDYNIEVLLGVDDSVVRFHGKEHVQNYLLTLMNIVNEIYHDESLGVHINVVLVRMIMLGYAKSVSLIERGNPSRSLENVCRWAYQQQKSDPNHSEHHDHAIFLTRQDFGPAGMQGYAPVTGMCHPVRSCTLNHEDGFSSAFVVAHETGHVLGMEHDGQGNRCGDETAMGSVMAPLVQAAFHRYHWSRCSGQELKRYIHSYDCLLDDPFEHDWPKLPELPGINYSMDEQCRFDFGVGYKMCTAFRTFDPCKQLWCSHPDNPYFCKTKKGPPLDGTECAPGKWCYKGHCMWKNANQLKQDGNWGSWTKFGSCSRTCGTGVRFRTRQCNNPMPVNGGQDCASVNFEYQLCSTEECPKHFEDFRAQQCQQRNSHFEYQKSKHHWLPYEHPDANKRCHLYCQSKEIGDIAYMKQLAHDGTRCSYKDPYSICVRGECVKVGCDKEIGSNKVEDKCGVCGGDNSHCRTVKGTFTRTPKKPGYLKMFDIPPGARHVFIQEDEASPHILAIKNQATGHYILNGKGEEAKSRSFIDLGVEWEYNTEDDIETLHTDGPLHDAVVVLVIPQENDTRSSLTYKYIIHEDSVPTINSNNVLQDEIDTFEWALKSWSQCSKPCAGGFQYTKYGCRRKSDNKMVHRSFCEASKKPKPIRRMLWMDEIWIADEWEYCTKTCGHSGYQIRTVRCIQPLHDGANRSVNIKYCSGDRPESRRPCNRTPCPAQWKAGPWAECSVTCGEGTESRQVTCRAGDQCDGEKPESVRVCKLAPCNGNQYCACFADEPCLGDKSIFCQMEVLARYCSIPGYNKLCCESCSKRSSTLPPPYLTEAAETKEEVMFHPRDLPRALAMPTSLVPQHAEVADENRSSVGRMPMEDDIDVLVSTSNSKSKGAELPQRRAYQAGSQTSRLIAMPYQSSTKIGNLSP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPQ3</accession>
    <name>A0A452IPQ3_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="5">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1 4">Belongs to the heat shock protein 70 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPQ3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033936.1">
        <property type="protein sequence ID" value="ENSGAGP00000029884.1"/>
        <property type="gene ID" value="ENSGAGG00000021572.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140662">
        <property type="term" value="F:ATP-dependent protein folding chaperone"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.30.30:FF:000001">
        <property type="entry name" value="heat shock 70 kDa protein-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.640.10:FF:000134">
        <property type="entry name" value="Heat shock cognate 71 kDa protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.420.40:FF:000026">
        <property type="entry name" value="Heat shock protein 70"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.30.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.420.40">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.640.10">
        <property type="entry name" value="Actin, Chain A, domain 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.34.10">
        <property type="entry name" value="Substrate Binding Domain Of DNAk, Chain A, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043129">
        <property type="entry name" value="ATPase_NBD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018181">
        <property type="entry name" value="Heat_shock_70_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029048">
        <property type="entry name" value="HSP70_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029047">
        <property type="entry name" value="HSP70_peptide-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013126">
        <property type="entry name" value="Hsp_70_fam"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19375">
        <property type="entry name" value="HEAT SHOCK PROTEIN 70KDA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00012">
        <property type="entry name" value="HSP70"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00301">
        <property type="entry name" value="HEATSHOCK70"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53067">
        <property type="entry name" value="Actin-like ATPase domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF100934">
        <property type="entry name" value="Heat shock protein 70kD (HSP70), C-terminal subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF100920">
        <property type="entry name" value="Heat shock protein 70kD (HSP70), peptide-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00297">
        <property type="entry name" value="HSP70_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00329">
        <property type="entry name" value="HSP70_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 4" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="2 4" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007381"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="RuleBase" id="RU003322"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029884.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="622" mass="68550" checksum="4D6EE9AC1DA577FF" modified="2019-05-08" version="1">GMPALKHPAIGIDLGTTKSCVGLFRDGKVEIIVSNQGIRTTPSCVAFTETGRLIGDEAKNQAALNSPNIIFSAKRLIGRTSEDPVVQADAEFWPFQIVKAEGKPQVQVSYKGKEKVFYPEEISAMVLGKLKEMAEDYLGHPITHAIITVPAHFNDSQRQATKDAGTIAGLNVLRILSETTAAALAYSLDNPSNGGHNVLIFDLGGGTFNVSVLTIDGGILEVKATSGDTHLGGDDFDNCLVSFFIDEFLKKHQENISKDRRAVWRLRMACEAAKCTLSSNIKASIDLYCLYKGIDFHADITRAQFEELCADLFRGTLKPVERALQDAKLSQAQIHDVVLVGASTRIPKIQELLQDFFGGRQLNKSLNLDEAVACGAAVQAAILIGERSEVTRGLLALDVVSMSLGIETDTGKMHTLIRRNTAMPAVVTRTFTTFDDHQPDLLIRAYEGERATAKENHLLGRFLLKGISPAPRGMVLVEATFYINADGILTVFALDKVASNANPITVDANKGRLSREEIQRLAAEAEKYRAEDRAQKWKLATKNYLQSCAFSMKRTSEDILNYFTLQAKKKVMEKCQLAFSWLSPTGWLKRRRSERVKQDKEMLCCPLLSGILTPKEDPSLKW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPQ6</accession>
    <name>A0A452IPQ6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Histone-arginine methyltransferase CARM1</fullName>
            <ecNumber evidence="3">2.1.1.319</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="14">Coactivator-associated arginine methyltransferase 1</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="13">Protein arginine N-methyltransferase 4</fullName>
        </alternativeName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>L-arginyl-[protein] + 2 S-adenosyl-L-methionine = N(omega),N(omega)-dimethyl-L-arginyl-[protein] + 2 S-adenosyl-L-homocysteine + 2 H(+)</text>
            <dbReference type="Rhea" id="RHEA:48096"/>
            <dbReference type="Rhea" id="RHEA-COMP:10532"/>
            <dbReference type="Rhea" id="RHEA-COMP:11991"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29965"/>
            <dbReference type="ChEBI" id="CHEBI:57856"/>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
            <dbReference type="ChEBI" id="CHEBI:61897"/>
            <dbReference type="EC" id="2.1.1.319"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="2.1.1.319" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452IPQ6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033870.1">
        <property type="protein sequence ID" value="ENSGAGP00000029830.1"/>
        <property type="gene ID" value="ENSGAGG00000021536.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070611">
        <property type="term" value="F:histone H3R2 methyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035242">
        <property type="term" value="F:protein-arginine omega-N asymmetric methyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032259">
        <property type="term" value="P:methylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd02440">
        <property type="entry name" value="AdoMet_MTases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.70.160.11:FF:000002">
        <property type="entry name" value="Probable histone-arginine methyltransferase CARM1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.150:FF:000031">
        <property type="entry name" value="Putative Histone-arginine methyltransferase CARM1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.70.160.11">
        <property type="entry name" value="Hnrnp arginine n-methyltransferase1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.150">
        <property type="entry name" value="Vaccinia Virus protein VP39"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025799">
        <property type="entry name" value="Arg_MeTrfase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055135">
        <property type="entry name" value="PRMT_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029063">
        <property type="entry name" value="SAM-dependent_MTases_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11006:SF49">
        <property type="entry name" value="HISTONE-ARGININE METHYLTRANSFERASE CARM1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11006">
        <property type="entry name" value="PROTEIN ARGININE N-METHYLTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06325">
        <property type="entry name" value="PrmA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22528">
        <property type="entry name" value="PRMT_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53335">
        <property type="entry name" value="S-adenosyl-L-methionine-dependent methyltransferases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51678">
        <property type="entry name" value="SAM_MT_PRMT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="9" id="KW-0156">Chromatin regulator</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6 16" id="KW-0489">Methyltransferase</keyword>
    <keyword evidence="12" id="KW-0539">Nucleus</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8 16" id="KW-0949">S-adenosyl-L-methionine</keyword>
    <keyword evidence="11" id="KW-0804">Transcription</keyword>
    <keyword evidence="10" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="7 16" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Protein arginine N-methyltransferase" evidence="17">
        <location>
            <begin position="285"/>
            <end position="443"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00011925"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00021804"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022603"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022691"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00030670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00033236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00049086"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF22528"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029830.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="468" mass="52997" checksum="60A21F2BA1088543" modified="2019-05-08" version="1">MPERAGRCLKCGTCLSVKPTRRLSLFLSSLARCAGPGKQQRPPELRGWAAAGIMAAPQQHQLLQYSVKLLLVSEEQLRGGLQLASPNQGALQLAVQPGEEDADVTVTDEYSTFSNALKRCWNQKKENSVFSQRTEEASAAQYFQFYGCLSQQQNMMQDFVRTATYHRAILQNHIDFKDKVVLDVGCGSGILSFFAVQAGARKVYAVEASSVAKYAEILVRSNNLSDKIMVLSGKIEEISLPENVDVVISEPMGYMLFNERMLESYLHSKKWLKSNGMMFPTFSDIHLAPFSDEQLYMEHYSRANFWYQQCFYGVNLSSLRSAAVDEYFRQPIVDTFDIRILMARTVKYTVNFTEAEEEDLHRVEIPFVFQMMQSGLIHGLAFWFDVAFVGSLVTVWLSTAPTEPLTHWYQVRCLLQTPLFAKEGETLSGKVLFVANKRQSYDIQIVAVVDQTGFKSGNILDLKNPFFR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPQ9</accession>
    <name>A0A452IPQ9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Nck-associated protein 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="6">Cell projection</location>
            <location evidence="6">Lamellipodium membrane</location>
            <topology evidence="6">Single-pass membrane protein</topology>
            <orientation evidence="6">Cytoplasmic side</orientation>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="7">Belongs to the HEM-1/HEM-2 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPQ9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033994.1">
        <property type="protein sequence ID" value="ENSGAGP00000029935.1"/>
        <property type="gene ID" value="ENSGAGG00000019065.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031258">
        <property type="term" value="C:lamellipodium membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031209">
        <property type="term" value="C:SCAR complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030031">
        <property type="term" value="P:cell projection assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030866">
        <property type="term" value="P:cortical actin cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048812">
        <property type="term" value="P:neuron projection morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019137">
        <property type="entry name" value="Nck-associated_protein-1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12093">
        <property type="entry name" value="NCK-ASSOCIATED PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12093:SF11">
        <property type="entry name" value="NCK-ASSOCIATED PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09735">
        <property type="entry name" value="Nckap1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="1" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="5" id="KW-0966">Cell projection</keyword>
    <keyword evidence="4" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="3" id="KW-1133">Transmembrane helix</keyword>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="507"/>
            <end position="532"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="518"/>
            <end position="532"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00037839"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00037947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00039689"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029935.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="995" mass="113438" checksum="5CE616F251C5AEBF" modified="2019-05-08" version="1">QNLIITYTTLMILLSRIEERKAIIGLYNYAHEMTHGASDREYPRLGQMIVDYENPLKKMMEEFVPHSKSLSDALISLQMVYPRRNLSADQWRNAQLLSLISAPSTMLNPAQSDTMPCEYLSLDAMEKWIIFGFILCHGILNSDATALNLWKLALQSSSCLSLFRDEVFHIHKAAEDLFVNIRGYNKRINDIRECKEAAVSHAGSMHRERRKFLRSALKELATVLSDQPGLLGPKALFVFMALSFARDEIIWLLRHADNMPKKSADDFIDKHIAELIFYMEELRAHVRKYGPVMQRYYVQYLSGFDAVVLNELVQNLSVCPEDESIIMSSFVNTMTSLSVKQVEDGEVFDFRGMRLDWFRLQAYTSVSKASLSLADHRELGKMMNTIIFHTKMVDSLVEMLVETSDLSIFCFYSRAFEKMFQQCLELPSQSRYSIAFPLLCTHFMSCTHELCPEERHHIGDRSLSLCNMFLDEMAKQARNLITDICTEQCTLSDQLLPKHCAKTISQAVNKKSKKQTGKKGEPEREKPGVESMRKNRLVVTNLDKLHTALSELCFSINYVPNMVVWEHTFTPREYLTSHLEIRFTKSIVGMTMYNQATQEIAKPSELLTSVRAYMTVLQSIENYVQIDITRVFNNVLLQQTQHLDSHGEPTITSLYTNWYLETLLRQVSNGHIAYFPAMKAFVNLPTENELTFNAEEYSDISEMRALSELLGPYGMKFLSESLMWHISSQVAELKKLVVENVEVLTQMRTSFDKPDQMAALFKRLSSVDSVLKRMTIIGVILSFRSLAQEALRDVLSYHIPFLVSSIEDFKDHIPRETDMKVAMNVYELSSAAGLPCEIDPALVVALSSQKSENISPEEEYKIACLLMVFVAVSLPTLASNVMSQYSPAIEGHCNNIHCLAKAINQIAAALFTIHKGSIEDRLKEFLALASSSLLKIGQETDKTTTRNRESVYLLLDMIVQESPFLTMDLLESCFPYVLLRNAYHAVYKQSVTSSA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPT1</accession>
    <name>A0A452IPT1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Cytosolic fatty-acid binding proteins domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1 2">Belongs to the calycin superfamily. Fatty-acid binding protein (FABP) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPT1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034183.1">
        <property type="protein sequence ID" value="ENSGAGP00000030108.1"/>
        <property type="gene ID" value="ENSGAGG00000021676.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008289">
        <property type="term" value="F:lipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd19443">
        <property type="entry name" value="FABP3-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.128.20:FF:000001">
        <property type="entry name" value="Fatty acid-binding protein, adipocyte"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.128.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012674">
        <property type="entry name" value="Calycin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000463">
        <property type="entry name" value="Fatty_acid-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031259">
        <property type="entry name" value="ILBP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000566">
        <property type="entry name" value="Lipocln_cytosolic_FA-bd_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11955">
        <property type="entry name" value="FATTY ACID BINDING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00061">
        <property type="entry name" value="Lipocalin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00178">
        <property type="entry name" value="FATTYACIDBP"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50814">
        <property type="entry name" value="Lipocalins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00214">
        <property type="entry name" value="FABP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0813">Transport</keyword>
    <feature type="domain" description="Cytosolic fatty-acid binding proteins" evidence="3">
        <location>
            <begin position="6"/>
            <end position="23"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00008390"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="RuleBase" id="RU003696"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS00214"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030108.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="131" mass="14815" checksum="C789E9A0BBFF538C" modified="2019-05-08" version="1">MALFLGTWKLVSSENFEEYMKELGVGFATRKLGSLAKPSVVISTNDDIITIKTESTFKNTEISFKLGEEFEETTADDRKTKSIITLDNGALIHVQKWDGKETTIKRELVDSKLVVECTMNKVTSTRVYERV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPT5</accession>
    <name>A0A452IPT5_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="3">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IPT5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033900.1">
        <property type="protein sequence ID" value="ENSGAGP00000029857.1"/>
        <property type="gene ID" value="ENSGAGG00000021552.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1"/>
            <end position="33"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="54"/>
            <end position="74"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="138"/>
            <end position="158"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="192"/>
            <end position="219"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="138"/>
            <end position="150"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029857.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="363" mass="40633" checksum="26A5FE9C42B363A6" modified="2019-05-08" version="1">MASKNRMRVTPGAEGAAEPDGSAGATGARPKGGRVVKARYLQYDRKPVGKSISANSSVVSGVKRSEKGETPTGRAGLFQKCKATTGTATSTLNRTVLGKDDLQSTLLEGHKIARPDLDLSAINDKSLLKKIPGSKSLFRAESRTRKKEQNSKSSDPDDVIGMLESQALLLNYASMKIEKNIAQLEEKGERNLSILCEEKEKQQKKLYELKRQLLLKKREQKLEEVLDKQVEVLTPLLTVCEQFKEEYKTFAAALDTTRHELPVKNIHIEGNRHTHLEDLKKHLAVTQSLLVEAAPGYLEENSKAFPVLKELKEVALKMDAELERSFKQVQDLSFEISKEVSLHNQKVCEDTHGIETLQHWYFN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPT8</accession>
    <name>A0A452IPT8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">PH domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IPT8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033908.1">
        <property type="protein sequence ID" value="ENSGAGP00000029863.1"/>
        <property type="gene ID" value="ENSGAGG00000021553.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="CDD" id="cd10573">
        <property type="entry name" value="PH_DAPP1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000229">
        <property type="entry name" value="Dual adapter for phosphotyrosine and 3-phosphotyrosine and 3-phosphoinositide"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051707">
        <property type="entry name" value="PI-Interact_SigTrans_Reg"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14336:SF15">
        <property type="entry name" value="DUAL ADAPTER FOR PHOSPHOTYROSINE AND 3-PHOSPHOTYROSINE AND 3-PHOSPHOINOSITIDE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14336">
        <property type="entry name" value="TANDEM PH DOMAIN CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00169">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00233">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="PH" evidence="2">
        <location>
            <begin position="116"/>
            <end position="211"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="53"/>
            <end position="112"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="1">
        <location>
            <begin position="53"/>
            <end position="81"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="89"/>
            <end position="101"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029863.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="226" mass="25371" checksum="FC699EA2A33FA5DD" modified="2019-05-08" version="1">RHNTLHTAQRALARASSSFPVRCREGPALTSSFGSLALQQQVAALGERLRRCCPGSGSRGGSPRSLPSLPARGGARAAGADGKSRAGRRREDEPRHEDPPRGARRGSARGAAEPRLGTKEGYLVKQGKIVKNWKMRWFTLHRNELKYFKDQMSTEPIRALDLTECSAVQFDYSQERVNCFCLVFPLRTYYLCAKTGIEADEWIKILRWKLSQIRKQQDQRDATSTP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPU0</accession>
    <name>A0A452IPU0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Lipocalin/cytosolic fatty-acid binding domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the calycin superfamily. Fatty-acid binding protein (FABP) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPU0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034070.1">
        <property type="protein sequence ID" value="ENSGAGP00000030001.1"/>
        <property type="gene ID" value="ENSGAGG00000021637.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008289">
        <property type="term" value="F:lipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd19443">
        <property type="entry name" value="FABP3-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.128.20:FF:000001">
        <property type="entry name" value="Fatty acid-binding protein, adipocyte"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.128.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012674">
        <property type="entry name" value="Calycin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000463">
        <property type="entry name" value="Fatty_acid-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031259">
        <property type="entry name" value="ILBP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000566">
        <property type="entry name" value="Lipocln_cytosolic_FA-bd_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11955">
        <property type="entry name" value="FATTY ACID BINDING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00061">
        <property type="entry name" value="Lipocalin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00178">
        <property type="entry name" value="FATTYACIDBP"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50814">
        <property type="entry name" value="Lipocalins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Lipocalin/cytosolic fatty-acid binding" evidence="2">
        <location>
            <begin position="8"/>
            <end position="132"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00008390"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF00061"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030001.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="133" mass="15124" checksum="CA6DC16F89E01A0C" modified="2019-05-08" version="1">RMCDLFMGIWKFVSSEKFEDYMKELGVGLATRTLGSLAKPTVIISMDGDVITIKTKSTFKNTEVSFKLGEEFEETTADDRKTKSIVTLEEGSLTQVQKWDGKETTIKRKLADGKMIVEYTMNNITCTRVYERA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPU7</accession>
    <name>A0A452IPU7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="19">Lymphocyte activation gene 3 protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="16">Lymphocyte activation gene 3 protein: Inhibitory receptor on antigen activated T-cells. Delivers inhibitory signals upon binding to ligands, such as FGL1. FGL1 constitutes a major ligand of LAG3 and is responsible for LAG3 T-cell inhibitory function. Following TCR engagement, LAG3 associates with CD3-TCR in the immunological synapse and directly inhibits T-cell activation. May inhibit antigen-specific T-cell activation in synergy with PDCD1/PD-1, possibly by acting as a coreceptor for PDCD1/PD-1. Negatively regulates the proliferation, activation, effector function and homeostasis of both CD8(+) and CD4(+) T-cells. Also mediates immune tolerance: constitutively expressed on a subset of regulatory T-cells (Tregs) and contributes to their suppressive function. Also acts as a negative regulator of plasmacytoid dendritic cell (pDCs) activation. Binds MHC class II (MHC-II); the precise role of MHC-II-binding is however unclear.</text>
    </comment>
    <comment type="function">
        <text evidence="15">May function as a ligand for MHC class II (MHC-II) on antigen-presenting cells (APC), promoting APC activation/maturation and driving Th1 immune response.</text>
    </comment>
    <comment type="subunit">
        <text evidence="18">Interacts with MHC class II (MHC-II); selectively recognizes stable complexes of peptide and MHC-II. Interacts with FGL1 (via the Fibrinogen C-terminal domain).</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="17">Belongs to the LAG3 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPU7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034207.1">
        <property type="protein sequence ID" value="ENSGAGP00000030129.1"/>
        <property type="gene ID" value="ENSGAGG00000021736.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009897">
        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045121">
        <property type="term" value="C:membrane raft"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042289">
        <property type="term" value="F:MHC class II protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1990782">
        <property type="term" value="F:protein tyrosine kinase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002250">
        <property type="term" value="P:adaptive immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035723">
        <property type="term" value="P:interleukin-15-mediated signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070374">
        <property type="term" value="P:positive regulation of ERK1 and ERK2 cascade"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042110">
        <property type="term" value="P:T cell activation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:002440">
        <property type="entry name" value="Lymphocyte activation gene 3 protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11422:SF12">
        <property type="entry name" value="MICROFIBRIL-ASSOCIATED GLYCOPROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11422">
        <property type="entry name" value="T-CELL SURFACE GLYCOPROTEIN CD4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13895">
        <property type="entry name" value="Ig_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-1064">Adaptive immunity</keyword>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="8" id="KW-0391">Immunity</keyword>
    <keyword evidence="14" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="11 20" id="KW-0472">Membrane</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="5 20" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9 20" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="32"/>
            <end position="55"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="21">
        <location>
            <begin position="134"/>
            <end position="215"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="21">
        <location>
            <begin position="236"/>
            <end position="314"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="21">
        <location>
            <begin position="316"/>
            <end position="400"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022859"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023130"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00057112"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00059221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00061264"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00065545"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00067553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030129.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="437" mass="47762" checksum="A22F6DDB0A57117F" modified="2019-05-08" version="1">MGKGNAGCEDGGAALFYSFPLSSPLPQSLIPIPSPLMCFFSLSMPFLLCLSLLLFPILSLPYSHKEPHMVLEVEYSGLLKRARSMMPRASVRETGFRQGNFSLWIEPLLSEDAGHYEALVRYGKETRRCQLELGMVTVTVNPPGLLVEAEPLLLNCNSSHPAKLVDMRWFHNGSLVPISSRFRSRHGALSISRPTVSDSGPWRCELTYSDDERVSASFNLQILGNSVSPVVYAAVGSAANLPCLLNRDPSASSILGVSAHWSHFAGGDLKIWDISRNGSNGSFTLHLSEVGPQDAGQYCCAVSIHGMTITRDVTLAVMTVTPSIKGPVAEGSHLLLICSLTHPRGHEHFQWRQLSSVQRYYPGSTLELIHVSQEDIGTWECSVHGSEGRLGAVEYELYITGTVSPVSHSAPLPPPSAPRFLLHLTSQHFPNLIHTLN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ01</accession>
    <name>A0A452IQ01_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Mediator of RNA polymerase II transcription subunit 23</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="7">Mediator complex subunit 23</fullName>
        </alternativeName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the Mediator complex subunit 23 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000033987.1">
        <property type="protein sequence ID" value="ENSGAGP00000029928.1"/>
        <property type="gene ID" value="ENSGAGG00000021033.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016592">
        <property type="term" value="C:mediator complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005667">
        <property type="term" value="C:transcription regulator complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010628">
        <property type="term" value="P:positive regulation of gene expression"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021629">
        <property type="entry name" value="Mediator_Med23"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12691">
        <property type="entry name" value="MEDIATOR OF RNA POLYMERASE II TRANSCRIPTION SUBUNIT 23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12691:SF10">
        <property type="entry name" value="MEDIATOR OF RNA POLYMERASE II TRANSCRIPTION SUBUNIT 23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11573">
        <property type="entry name" value="Med23"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0539">Nucleus</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0804">Transcription</keyword>
    <keyword evidence="4" id="KW-0805">Transcription regulation</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010222"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00019696"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00031961"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029928.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1307" mass="149957" checksum="46D9E61A461758AA" modified="2019-05-08" version="1">MRIKQSRRHTVFRGLNLFLLQKTEVIEEAFPGMFMDTPEDERTKLISCLGAFRQFWSSLSQESHEQCVQWIVRFIHSQHSPKRISFLYDCLAMAVETGLLPPRMVCESLINSETLEWERTQLWALTFKLVRKIIGGVDYKGVRDLLKVILEKILTIPNTVSSAVVQQLLAAREVVAYILERNACLLPAYFAVTEIRKLYPEGKLPHWLLGNLVSDFVDTFRPTARINSICGRCSLLPVVNNSGAMCNSWKLDPTTLRFPLKGLLPYDKDLFEPQTALLRYVLEQPYSRDMVCNMLGLNKQVLYNTHKQRCPVLEDQLVDLVVYAMERSETEEKFDDGGTSQLLWQHLSSQLIFFVLFQFASFPHMVLSLHQKLAGRGLIKGRDHLMWVLLQFISGSIQKNALADFLPVMKLFDLLYPEKECIPVPDINKPQSTHAFAMTCIWIHLNRKAHSDNSKLQIPIPHSLKLHHEFLQQSLRNKSLQMNDYKIALLCNAYSTNSECFTLPMGVLVETIYGNGNMRIPLPGTNCMASGSITPLPMNLLDSLTVHAKMSLIHSIATRVIKLAHAKSSLALAPALVETYSRLLVYMEIESLGIKGFISQLLPTVFKSHAWGILHTLLEMFSYRMHHIQPHYRVQLLSHLHSLAAVPQTNQNQLHLCVESTALRLITALGSSEVQPQFTRFLSDPKTVLSAESEELNRALILTLARATHVTDFFTGSDSIQGTWCKDILQTIMSFTPHNWASHTLSCFPAPLQAFFKQNNVPQESRFNLKKNVEEEYRKWKSMTNENDIITHFSMQGSPPLFLCLLWKMLLETDHINQIGYRVLERIGARALVAHVRTFADFLVYEFSTSAGGQQLNKCIEILNDMVWKYNIVTLDRLILCLAMRSHEGNEAQVCYFIIQLLLLKPNDFRNRVSDFVKENSPEHWLQNDWHTKHMSYHKKYPEKLYFEGLAEQVNPPVQIQPQYLPIYFGNVCLRFLPVFDIVIHRFLELLPVSKSLETLLDHLGGLYKFHDRPVTYLYNTLHYYERHLRERTNLKRKLVHAIIGSLKDNRPQGWCLSDTYIKSAMNVREDNPWIPDDTYYCKLIGRLSPGPFPNCDWRFNEFPNPAAHALHVTCVELMALAVPGKDVGNALLNVVLKSQPLVPRENITAWMNAIGLIITALPVSLYVPPSLLSLLPTVSYLKTSLCLCHKQIXVYHLVGPFLQRFQQERTRCMIEIGVAFYEMLLDVDQCSTHLNYMDPICDFLYHMKYMFTGDSVKDQVEKIICNLRPSLKLRLRFITHISKMEPAAIPQQSINSGSPAPQPITQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ03</accession>
    <name>A0A452IQ03_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Alpha-1,4 glucan phosphorylase</fullName>
            <ecNumber evidence="15">2.4.1.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="12 15">Allosteric enzyme that catalyzes the rate-limiting step in glycogen catabolism, the phosphorolytic cleavage of glycogen to produce glucose-1-phosphate, and plays a central role in maintaining cellular and organismal glucose homeostasis.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>[(1-&gt;4)-alpha-D-glucosyl](n) + phosphate = [(1-&gt;4)-alpha-D-glucosyl](n-1) + alpha-D-glucose 1-phosphate</text>
            <dbReference type="Rhea" id="RHEA:41732"/>
            <dbReference type="Rhea" id="RHEA-COMP:9584"/>
            <dbReference type="Rhea" id="RHEA-COMP:9586"/>
            <dbReference type="ChEBI" id="CHEBI:15444"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:58601"/>
            <dbReference type="EC" id="2.4.1.1"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="11">
            <dbReference type="Rhea" id="RHEA:41733"/>
        </physiologicalReaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 15">
            <name>pyridoxal 5'-phosphate</name>
            <dbReference type="ChEBI" id="CHEBI:597326"/>
        </cofactor>
    </comment>
    <comment type="subunit">
        <text evidence="13">Homodimer. Homotetramer; to form the enzymatically active phosphorylase A.</text>
    </comment>
    <comment type="similarity">
        <text evidence="2 15">Belongs to the glycogen phosphorylase family.</text>
    </comment>
    <dbReference type="EC" id="2.4.1.1" evidence="15"/>
    <dbReference type="AlphaFoldDB" id="A0A452IQ03"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034134.1">
        <property type="protein sequence ID" value="ENSGAGP00000030062.1"/>
        <property type="gene ID" value="ENSGAGG00000021692.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008184">
        <property type="term" value="F:glycogen phosphorylase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030170">
        <property type="term" value="F:pyridoxal phosphate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005980">
        <property type="term" value="P:glycogen catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd04300">
        <property type="entry name" value="GT35_Glycogen_Phosphorylase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.2000:FF:000005">
        <property type="entry name" value="Alpha-1,4 glucan phosphorylase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.2000:FF:000149">
        <property type="entry name" value="Glycogen phosphorylase, muscle form"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.2000">
        <property type="entry name" value="Glycogen Phosphorylase B"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011833">
        <property type="entry name" value="Glycg_phsphrylas"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000811">
        <property type="entry name" value="Glyco_trans_35"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035090">
        <property type="entry name" value="Pyridoxal_P_attach_site"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR02093">
        <property type="entry name" value="P_ylase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11468">
        <property type="entry name" value="GLYCOGEN PHOSPHORYLASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11468:SF32">
        <property type="entry name" value="GLYCOGEN PHOSPHORYLASE, MUSCLE FORM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00343">
        <property type="entry name" value="Phosphorylase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000460">
        <property type="entry name" value="Pprylas_GlgP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53756">
        <property type="entry name" value="UDP-Glycosyltransferase/glycogen phosphorylase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00102">
        <property type="entry name" value="PHOSPHORYLASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0007">Acetylation</keyword>
    <keyword evidence="3" id="KW-0021">Allosteric enzyme</keyword>
    <keyword evidence="10 15" id="KW-0119">Carbohydrate metabolism</keyword>
    <keyword evidence="5" id="KW-0321">Glycogen metabolism</keyword>
    <keyword evidence="6 15" id="KW-0328">Glycosyltransferase</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="8 14 15" id="KW-0663">Pyridoxal phosphate</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7 15" id="KW-0808">Transferase</keyword>
    <feature type="modified residue" description="N6-(pyridoxal phosphate)lysine" evidence="14">
        <location>
            <position position="681"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001933"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006047"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022533"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022600"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022676"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022898"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023277"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00036074"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00037413"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00046356"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PIRSR" id="PIRSR000460-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="RuleBase" id="RU000587"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030062.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="817" mass="93740" checksum="B640B491EE27A344" modified="2019-05-08" version="1">MSLPLSDYEKRKQISVRGLAGVENVSDLKKNFNRHLHFTLVKDRNVATPRDYYFALAHTVRDHLVGRWIRTQQYYYEKDPKRIYYLSLEFYMGRTLQNTMVNLGLQNACDEATYQLGLDMEELQEIEEDAGLGNGGLGRLAACFLDSMATLGLAAYGYGIRYEFGIFNQKISGGWQVEEADDWLRYGNPWEKARPEYMIPVHFYGHVDHGPDGTKWVDTQVILALPYDTPVPGYRNNTVNTMRLWSARAPNEFNLKDFNVGGYIQAVLDRNLAENISRVLYPNDNFFEGKELRLKQEYFTVAATLQDIIRRFKSSKFGSRDPVRTSFDAFPDKVAIQLNDTHPSLAIPELMRVLVDIEKLDWDKAWDITVRTCAYTNHTVLPEALERWPVHLLETLLPRHLEIIYEINQRFLDKVNAMFPGDFDRLRRMSLVEEGSIKRINMAHLCIVGSHAINGVARIHSDIIKKSVFKDFYEFEPHKFQNKTNGITPRRWLVLCNPGLAEVIAERIGDEYISNLDHLKKLLAYVDDEGFIRDVAKIKQENKLKFAAYLEKEYKVKINLNSLFDVQVKRIHEYKRQLLNCLHVITLYNRIKKEPNQHFVPRTVMIGGKAAPGYHMAKMIIKLVTAIGDVINNDPVIGDRLKMIFLENYRVSLAEKVIPAADLSEQISTAGTEASGTGNMKFMLNGALTIGTMDGANVEMAEEAGEENLFIFGMRVEDVEEMDRKGYCAKGYYDRIPELRQVIDQLSSGFFSPKQPQLFKDIVNMLMNNDRFKVFADYEAYIKCQENVSALYKVSVHPSGLPGVLHPALVREWGLVG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ04</accession>
    <name>A0A452IQ04_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Palladin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="19">Interacts with EPS8. Interacts with LASP1. Interacts with VASP. Interacts with ACTN. Interacts with SORBS2. Interacts with PFN1. Interacts with LPP. Interacts with SPIN90. Interacts with SRC. Interacts with EZR. Interacts with RAI14.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="4">Cell junction</location>
            <location evidence="4">Focal adhesion</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="6">Cell projection</location>
            <location evidence="6">Axon</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="8">Cell projection</location>
            <location evidence="8">Growth cone</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="7">Cell projection</location>
            <location evidence="7">Lamellipodium</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cell projection</location>
            <location evidence="1">Podosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="5">Cell projection</location>
            <location evidence="5">Ruffle</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Myofibril</location>
            <location evidence="2">Sarcomere</location>
            <location evidence="2">Z line</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="18">Belongs to the myotilin/palladin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQ04"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034271.1">
        <property type="protein sequence ID" value="ENSGAGP00000030190.1"/>
        <property type="gene ID" value="ENSGAGG00000021781.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005925">
        <property type="term" value="C:focal adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030426">
        <property type="term" value="C:growth cone"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030027">
        <property type="term" value="C:lamellipodium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002102">
        <property type="term" value="C:podosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001726">
        <property type="term" value="C:ruffle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001725">
        <property type="term" value="C:stress fiber"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030018">
        <property type="term" value="C:Z disc"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003779">
        <property type="term" value="F:actin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004672">
        <property type="term" value="F:protein kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
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        <property type="entry name" value="palladin isoform X1"/>
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        <property type="entry name" value="palladin isoform X2"/>
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        <property type="match status" value="3"/>
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        <property type="entry name" value="PALLADIN, CYTOSKELETAL ASSOCIATED PROTEIN"/>
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    </dbReference>
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    <keyword evidence="14" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="12" id="KW-0965">Cell junction</keyword>
    <keyword evidence="16" id="KW-0966">Cell projection</keyword>
    <keyword evidence="9" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="15" id="KW-0206">Cytoskeleton</keyword>
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    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
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        <location>
            <begin position="351"/>
            <end position="435"/>
        </location>
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            <begin position="485"/>
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            <end position="11"/>
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            <begin position="325"/>
            <end position="343"/>
        </location>
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        <source>
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        <source>
            <dbReference type="PROSITE" id="PS50835"/>
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        <source>
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        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
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    <sequence length="733" mass="81092" checksum="2AB229873179BF15" modified="2019-05-08" version="1">SQTPVPISFANQIRLEQEGGQRYHHHQQHQNPPPSPPFPPPPSFQELLSSQSVTSPVQMSILNSHTSPTMQSSSTFSYARPKQFIAAQNISPASGYVTPSSGSSTSSLPSPMSPTASQKQFGRAPVPPFSQPFAPEGEYLWSPSSSSPPPPPPPVFSPTATFSVSDSFPPPPPPLPSSVSSPSRSLSPTARFSNSSQSPAAFLSSMLPSQPPPISVNALGLPKGVMPPGFPKKTGRTARIASDEEIQGTKDAVIQDLERKLRFKEELLNNGQPRLTYEEKMARRLLGADNAATVFNIQEPDEEPTTQEYKVSSFEQRLISEIEYRLERSPVEESDDEVQHGDEPTDNSVAPYFEMKLKHYKIFEGTSVTFTCRVAGNPKPKIYWFKDGKQISKRSDHYRIQREPDGTCSLHTTASTLDDDGNYTIMAANPQGRISCTGRLMVQAVNQRGRSPRTPPSQPHIRRPRSRSRDSGDENEPIQERFFRPHFLQAPGDLIVQEGKLCRMDCKVSGLPTPDLSWQLNGRPIRPDSSHKMLVRENGVHSLIIEPVTARDSGIYTCIATNRAGQNTFSLELIVAAKEEHKPPVFIEKLQNTGVADGYPVRLECRVSGVPYPQIFWKKENESLTYNTERVSMHQDNHGYICLLIQGATKEDAGWYTVSAKNEAGIVSCTARLDVYTQWQQQPQSTKPKKVRPSASRYAALSDQGLDIKAAFQPEANPVHLTLQCGLVESDDL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ05</accession>
    <name>A0A452IQ05_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="12">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 7">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 7">Belongs to the ETS family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQ05"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034101.1">
        <property type="protein sequence ID" value="ENSGAGP00000030031.1"/>
        <property type="gene ID" value="ENSGAGG00000021670.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043565">
        <property type="term" value="F:sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd08540">
        <property type="entry name" value="SAM_PNT-ERG"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Fli-1 proto-oncogene, ETS transcription factor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.10:FF:000039">
        <property type="entry name" value="Friend leukemia integration 1 transcription factor"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Transcription Factor, Ets-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.10">
        <property type="entry name" value="Winged helix-like DNA-binding domain superfamily/Winged helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Ets_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046328">
        <property type="entry name" value="ETS_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003118">
        <property type="entry name" value="Pointed_dom"/>
    </dbReference>
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        <property type="entry name" value="SAM"/>
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        <property type="entry name" value="SAM/pointed_sf"/>
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        <property type="entry name" value="WH-like_DNA-bd_sf"/>
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        <property type="entry name" value="WH_DNA-bd_sf"/>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="TRANSCRIPTIONAL REGULATOR ERG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00178">
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="ETSDOMAIN"/>
    </dbReference>
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        <property type="entry name" value="ETS"/>
        <property type="match status" value="1"/>
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        <property type="match status" value="1"/>
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        <property type="entry name" value="SAM/Pointed domain"/>
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    </dbReference>
    <dbReference type="SUPFAM" id="SSF46785">
        <property type="entry name" value="Winged helix' DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00345">
        <property type="entry name" value="ETS_DOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00346">
        <property type="entry name" value="ETS_DOMAIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50061">
        <property type="entry name" value="ETS_DOMAIN_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51433">
        <property type="entry name" value="PNT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50105">
        <property type="entry name" value="SAM_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 7" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="6 7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0804">Transcription</keyword>
    <keyword evidence="3" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="PNT" evidence="11">
        <location>
            <begin position="110"/>
            <end position="196"/>
        </location>
    </feature>
    <feature type="domain" description="SAM" evidence="10">
        <location>
            <begin position="131"/>
            <end position="176"/>
        </location>
    </feature>
    <feature type="domain" description="ETS" evidence="9">
        <location>
            <begin position="308"/>
            <end position="388"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="31"/>
            <end position="53"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="241"/>
            <end position="301"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="31"/>
            <end position="44"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="267"/>
            <end position="278"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005562"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU004019"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50061"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50105"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS51433"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030031.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="476" mass="53845" checksum="945D941DDF9EB85B" modified="2019-05-08" version="1">MEEEALSVVSEDQSLFECAYGTQHLTKTEMTASSSSEYGQTSKMSPRVPQQDWLSQPPARVTIKMECNPNQINGSRNSPDDCSVAKGGKMVSGSDNVGMNYGSYMEEKHVPPPNMTTNERRVIVPADPTLWSTDHVRQWLEWAVKEYGLPDVDILLFQNIDGKELCKMTKDDFQRLTPSYNADILLSHLHYLRETPLPHLTSDDVDKALQNSPRLMHARNTGGAAFIFPNTSVYPEATQRITTRPDLSYEQARRSAWTSHNHPAPQSKATQPSSSTVPKTEDQRPQLDPYQILGPTSSRLANPGSGQIQLWQFLLELLSDSSNSNCITWEGTNGEFKMTDPDEVARRWGERKSKPNMNYDKLSRALRYYYDKNIMTKVHGKRYAYKFDFHGIAQALQPHPPESSMYKYPTDLPYMSPYHAHPQKMNFVAPHPPALPVTSSSFFAAPTTYWNSPTGSIYPNTRLPATHMPSHLGTYY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ07</accession>
    <name>A0A452IQ07_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Mediator of RNA polymerase II transcription subunit 23</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="7">Mediator complex subunit 23</fullName>
        </alternativeName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the Mediator complex subunit 23 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000033992.1">
        <property type="protein sequence ID" value="ENSGAGP00000029933.1"/>
        <property type="gene ID" value="ENSGAGG00000021033.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016592">
        <property type="term" value="C:mediator complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005667">
        <property type="term" value="C:transcription regulator complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010628">
        <property type="term" value="P:positive regulation of gene expression"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021629">
        <property type="entry name" value="Mediator_Med23"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12691">
        <property type="entry name" value="MEDIATOR OF RNA POLYMERASE II TRANSCRIPTION SUBUNIT 23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12691:SF10">
        <property type="entry name" value="MEDIATOR OF RNA POLYMERASE II TRANSCRIPTION SUBUNIT 23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11573">
        <property type="entry name" value="Med23"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0539">Nucleus</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0804">Transcription</keyword>
    <keyword evidence="4" id="KW-0805">Transcription regulation</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010222"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00019696"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00031961"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029933.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1317" mass="151073" checksum="7AF53175052499F1" modified="2019-05-08" version="1">MHERAEKNIVPAEASGACHFQETKTEVIEEAFPGMFMDTPEDERTKLISCLGAFRQFWSSLSQESHEQCVQWIVRFIHSQHSPKRISFLYDCLAMAVETGLLPPRMVCESLINSETLEWERTQLWALTFKLVRKIIGGVDYKGVRDLLKVILEKILTIPNTVSSAVVQQLLAAREVVAYILERNACLLPAYFAVTEIRKLYPEGKLPHWLLGNLVSDFVDTFRPTARINSICGRCSLLPVVNNSGAMCNSWKLDPTTLRFPLKGLLPYDKDLFEPQTALLRYVLEQPYSRDMVCNMLGLNKQVLYNTHKQRCPVLEDQLVDLVVYAMERSETEEKFDDGGTSQLLWQHLSSQLIFFVLFQFASFPHMVLSLHQKLAGRGLIKGRDHLMWVLLQFISGSIQKNALADFLPVMKLFDLLYPEKECIPVPDINKPQSTHAFAMTCIWIHLNRKAHSDNSKLQIPIPHSLKLHHEFLQQSLRNKSLQMNDYKIALLCNAYSTNSECFTLPMGVLVETIYGNGNMRIPLPGTNCMASGSITPLPMNLLDSLTVHAKMSLIHSIATRVIKLAHAKSSLALAPALVETYSRLLVYMEIESLGIKGFISQLLPTVFKSHAWGILHTLLEMFSYRMHHIQPHYRVQLLSHLHSLAAVPQTNQNQLHLCVESTALRLITALGSSEVQPQFTRFLSDPKTVLSAESEELNRALILTLARATHVTDFFTGSDSIQGTWCKDILQTIMSFTPHNWASHTLSCFPAPLQAFFKQNNVPQESRFNLKKNVEEEYRKWKSMTNENDIITHFSMQGSPPLFLCLLWKMLLETDHINQIGYRVLERIGARALVAHVRTFADFLVYEFSTSAGGQQLNKCIEILNDMVWKYNIVTLDRLILCLAMRSHEGNEAQVCYFIIQLLLLKPNDFRNRVSDFVKENSPEHWLQNDWHTKHMSYHKKYPEKLYFEGLAEQVNPPVQIQPQYLPIYFGNVCLRFLPVFDIVIHRFLELLPVSKSLETLLDHLGGLYKFHDRPVTYLYNTLHYYERHLRERTNLKRKLVHAIIGSLKDNRPQGWCLSDTYIKSAMNVREDNPWIPDDTYYCKLIGRLSPGPFPNCDWRFNEFPNPAAHALHVTCVELMALAVPGKDVGNALLNVVLKSQPLVPRENITAWMNAIGLIITALPVSLYFHASPSYITHPFETLHCLPISRCKKCKRLIFILQXVYHLVGPFLQRFQQERTRCMIEIGVAFYEMLLDVDQCSTHLNYMDPICDFLYHMKYMFTGDSVKDQVEKIICNLRPSLKLRLRFITHISKMEPAAIPQQSINSGSPAPQPITQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ21</accession>
    <name>A0A452IQ21_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Peptidase S1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the peptidase S1 family. Snake venom subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQ21"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034291.1">
        <property type="protein sequence ID" value="ENSGAGP00000030210.1"/>
        <property type="gene ID" value="ENSGAGG00000021723.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00190">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.10:FF:000010">
        <property type="entry name" value="Kallikrein related peptidase 11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.10:FF:000005">
        <property type="entry name" value="Serine protease 37"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.10.10">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009003">
        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043504">
        <property type="entry name" value="Peptidase_S1_PA_chymotrypsin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001314">
        <property type="entry name" value="Peptidase_S1A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050127">
        <property type="entry name" value="Serine_Proteases_S1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001254">
        <property type="entry name" value="Trypsin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018114">
        <property type="entry name" value="TRYPSIN_HIS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033116">
        <property type="entry name" value="TRYPSIN_SER"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24264:SF15">
        <property type="entry name" value="RIKEN CDNA 2210010C04 GENE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24264">
        <property type="entry name" value="TRYPSIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00089">
        <property type="entry name" value="Trypsin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00722">
        <property type="entry name" value="CHYMOTRYPSIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00020">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50494">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50240">
        <property type="entry name" value="TRYPSIN_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00134">
        <property type="entry name" value="TRYPSIN_HIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00135">
        <property type="entry name" value="TRYPSIN_SER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5 8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="4 8" id="KW-0645">Protease</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0964">Secreted</keyword>
    <keyword evidence="6 8" id="KW-0720">Serine protease</keyword>
    <feature type="domain" description="Peptidase S1" evidence="9">
        <location>
            <begin position="28"/>
            <end position="246"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009228"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="RuleBase" id="RU363034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030210.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="248" mass="27015" checksum="154EC8E7588CF9E0" modified="2019-05-08" version="1">SPALLCLQFTPVVNLSACLPVSSQNNRIVGGAPCIPHSMPWQAALFMGSQLNCGGTLIDRNWVVTAAHCHCGCACCEGRGCVIVRVSISVRLGEHNIRHLDWTEQLRLSEKTILHPGYNSASKDNDIMLIKLLTPANLNKNECVVSGWGTTKMYFPPVLHCGKVSIFSQEQCRSIYPRYYTENMLCAGVLEGGIDSCQGDSGGPLLCNGKLQGIVSWGTQTCALPNKPGVYVAVCKYVDWIRETINNN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ23</accession>
    <name>A0A452IQ23_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">Glycosyltransferase 2-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IQ23"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034078.1">
        <property type="protein sequence ID" value="ENSGAGP00000030009.1"/>
        <property type="gene ID" value="ENSGAGG00000021650.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008376">
        <property type="term" value="F:acetylgalactosaminyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019276">
        <property type="term" value="P:UDP-N-acetylgalactosamine metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006047">
        <property type="term" value="P:UDP-N-acetylglucosamine metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00761">
        <property type="entry name" value="Glyco_tranf_GTA_type"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.550.10">
        <property type="entry name" value="Spore Coat Polysaccharide Biosynthesis Protein SpsA, Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001173">
        <property type="entry name" value="Glyco_trans_2-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029044">
        <property type="entry name" value="Nucleotide-diphossugar_trans"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15046:SF2">
        <property type="entry name" value="BETA-1,4 N-ACETYLGALACTOSAMINYLTRANSFERASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15046">
        <property type="entry name" value="GLYCO_TRANS_2-LIKE DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00535">
        <property type="entry name" value="Glycos_transf_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53448">
        <property type="entry name" value="Nucleotide-diphospho-sugar transferases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Glycosyltransferase 2-like" evidence="1">
        <location>
            <begin position="174"/>
            <end position="285"/>
        </location>
    </feature>
    <evidence type="ECO:0000259" key="1">
        <source>
            <dbReference type="Pfam" id="PF00535"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030009.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="414" mass="47203" checksum="B3F5388E3E2B8FD7" modified="2019-05-08" version="1">PAKAGGDPIRVVALAKSRCSLPELNIPQCWEGSVQLRSEQCQYPSMTQDIVIAKPNVPLSYPIQGVEVMPLHTIMIPGIRVLCLFIACSWAQLPVESKTLSLVHTLYIGRGKKELLILTSDVELLNFILEHVTYTSVVYQLSAVDMSEYRLDNFFVPSLAVYRKISSLVTITTKTFLRYHKLRFLIKSIRQYYPDIKIIVADDSEVPEKIIEENIEHYIMPFGKGWFAGRNLAISQVTTKYYLWVDDDYLFTENTKIEKLLDVLERTNLDMVGGSVKGDTYSFQLLYEQSDDGYCLHLRYGSFHKLDGFPNCVVTSGVVNFYLAHTDKSRYVSYDPLLQRVAHSEFFVDGLGILLVGSCSDVSVGHQDHNPASDPNLAKIEDQYKTFRDNTDAQVQFKLALHYFKNRLKCYSTR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ26</accession>
    <name>A0A452IQ26_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Peptidase S1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the peptidase S1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQ26"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034296.1">
        <property type="protein sequence ID" value="ENSGAGP00000030215.1"/>
        <property type="gene ID" value="ENSGAGG00000021723.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030141">
        <property type="term" value="C:secretory granule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00190">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.10:FF:000077">
        <property type="entry name" value="Predicted protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.10.10">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009003">
        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043504">
        <property type="entry name" value="Peptidase_S1_PA_chymotrypsin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001314">
        <property type="entry name" value="Peptidase_S1A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001254">
        <property type="entry name" value="Trypsin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018114">
        <property type="entry name" value="TRYPSIN_HIS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033116">
        <property type="entry name" value="TRYPSIN_SER"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271:SF48">
        <property type="entry name" value="KALLIKREIN-14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271">
        <property type="entry name" value="KALLIKREIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00089">
        <property type="entry name" value="Trypsin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00722">
        <property type="entry name" value="CHYMOTRYPSIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00020">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50494">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50240">
        <property type="entry name" value="TRYPSIN_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00134">
        <property type="entry name" value="TRYPSIN_HIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00135">
        <property type="entry name" value="TRYPSIN_SER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3 6" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="2 6" id="KW-0645">Protease</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 6" id="KW-0720">Serine protease</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="7">
        <location>
            <begin position="1"/>
            <end position="25"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019588588" description="Peptidase S1 domain-containing protein" evidence="7">
        <location>
            <begin position="26"/>
            <end position="255"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase S1" evidence="8">
        <location>
            <begin position="30"/>
            <end position="253"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007664"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="RuleBase" id="RU363034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030215.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="255" mass="28071" checksum="F5A8B0B49DF80D92" modified="2019-05-08" version="1">TELANGEGMVWWSLLISTGCIWVSSQNNRIVGGAPCIPHSMPWQAALFMGSQLNCGGTLIDRNWVVTAAHCHCPPWTSVRLGEHNIRHLDWTEQLRLSEKTILHPGYNSASKDNDIMLIKLLTPANLNKNVQPLELPTSCPSPKEECVVSGWGTTKSPQLYFPPVLHCGKVSIFSQEQCRSIYPRYYTENMLCAGVLEGGIDSCQGDSGGPLLCNGKLQGIVSWGTQTCALPNKPGVYVAVCKYVDWIRETINNN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ27</accession>
    <name>A0A452IQ27_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Menin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQ27"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034128.1">
        <property type="protein sequence ID" value="ENSGAGP00000030056.1"/>
        <property type="gene ID" value="ENSGAGG00000021688.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000785">
        <property type="term" value="C:chromatin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035097">
        <property type="term" value="C:histone methyltransferase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003682">
        <property type="term" value="F:chromatin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000976">
        <property type="term" value="F:transcription cis-regulatory region binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000403">
        <property type="term" value="F:Y-form DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006325">
        <property type="term" value="P:chromatin organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045786">
        <property type="term" value="P:negative regulation of cell cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008285">
        <property type="term" value="P:negative regulation of cell population proliferation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd14456">
        <property type="entry name" value="Menin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007747">
        <property type="entry name" value="Menin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12693">
        <property type="entry name" value="MENIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12693:SF3">
        <property type="entry name" value="MENIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05053">
        <property type="entry name" value="Menin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0156">Chromatin regulator</keyword>
    <keyword evidence="7" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="9" id="KW-0539">Nucleus</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0678">Repressor</keyword>
    <keyword evidence="8" id="KW-0804">Transcription</keyword>
    <keyword evidence="6" id="KW-0805">Transcription regulation</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00021162"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022491"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030056.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="422" mass="47659" checksum="81023F8496A435CC" modified="2019-05-08" version="1">MGLKPWQKALFPLRSVEDVVRLFMAELQQEQPDLVLLSLVLGFVEHFLAVNRVIPTNVPGISFEPCPGPEPDSLTYFPVAELSIVAALYARFTAQIRGAVDLSLYPRPEGFSSRELVKKVSDVIWNSLSRSYFKDRAHIQSLFSFITGTKLDSSGVAFAVVGACQVLGLPDVHLALSEDHAWVVFGRDGEQTAEVTWHGKGNEDRRGQTVNAGVAERSWLYLKGSYLRCDRHMEVAFMVCAINPSIDLHTDSLELLQLQQRLLWVLYDMGHLERYPMALGNLADLEELEPTPGRPDPLSIYHKGISSARKYYNNEHIYPYMYLAGYHCRNKNVKEALEAWADTATVIQDYNYCREDEEIYKEFFDVANDVVPNLLKEVANLAEPLEEKGAGERGEVSGVLSWMGLWDLGGRKRDTRLEGLWS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ30</accession>
    <name>A0A452IQ30_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="19">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the G-protein coupled receptor 2 family. Adhesion G-protein coupled receptor (ADGR) subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQ30"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034301.1">
        <property type="protein sequence ID" value="ENSGAGP00000030220.1"/>
        <property type="gene ID" value="ENSGAGG00000021690.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098794">
        <property type="term" value="C:postsynapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043083">
        <property type="term" value="C:synaptic cleft"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004930">
        <property type="term" value="F:G protein-coupled receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007189">
        <property type="term" value="P:adenylate cyclase-activating G protein-coupled receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007166">
        <property type="term" value="P:cell surface receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016525">
        <property type="term" value="P:negative regulation of angiogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016322">
        <property type="term" value="P:neuron remodeling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd15251">
        <property type="entry name" value="7tmB2_BAI_Adhesion_VII"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1070.10:FF:000016">
        <property type="entry name" value="Adhesion G protein-coupled receptor B2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.610:FF:000002">
        <property type="entry name" value="Adhesion G protein-coupled receptor B2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000003">
        <property type="entry name" value="Adhesion G protein-coupled receptor B2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000004">
        <property type="entry name" value="Adhesion G protein-coupled receptor B2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000012">
        <property type="entry name" value="Adhesion G protein-coupled receptor B2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.1240.10:FF:000002">
        <property type="entry name" value="Adhesion G protein-coupled receptor B2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.220.50:FF:000027">
        <property type="entry name" value="Adhesion G protein-coupled receptor B3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.610">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.220.50">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.1240.10">
        <property type="entry name" value="GPCR, family 2, extracellular hormone receptor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1070.10">
        <property type="entry name" value="Rhodopsin 7-helix transmembrane proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.20.100.10">
        <property type="entry name" value="Thrombospondin type-1 (TSP1) repeat"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032471">
        <property type="entry name" value="AGRL2-4_GAIN_subdom_A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057244">
        <property type="entry name" value="GAIN_B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046338">
        <property type="entry name" value="GAIN_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017981">
        <property type="entry name" value="GPCR_2-like_7TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008077">
        <property type="entry name" value="GPCR_2_brain_angio_inhib"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036445">
        <property type="entry name" value="GPCR_2_extracell_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001879">
        <property type="entry name" value="GPCR_2_extracellular_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000832">
        <property type="entry name" value="GPCR_2_secretin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017983">
        <property type="entry name" value="GPCR_2_secretin-like_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000203">
        <property type="entry name" value="GPS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000884">
        <property type="entry name" value="TSP1_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036383">
        <property type="entry name" value="TSP1_rpt_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12011:SF40">
        <property type="entry name" value="ADHESION G PROTEIN-COUPLED RECEPTOR B3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12011">
        <property type="entry name" value="ADHESION G-PROTEIN COUPLED RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00002">
        <property type="entry name" value="7tm_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16489">
        <property type="entry name" value="GAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01825">
        <property type="entry name" value="GPS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02793">
        <property type="entry name" value="HRM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00090">
        <property type="entry name" value="TSP_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01694">
        <property type="entry name" value="BAIPRECURSOR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00249">
        <property type="entry name" value="GPCRSECRETIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00303">
        <property type="entry name" value="GPS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00008">
        <property type="entry name" value="HormR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00209">
        <property type="entry name" value="TSP1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF82895">
        <property type="entry name" value="TSP-1 type 1 repeat"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00650">
        <property type="entry name" value="G_PROTEIN_RECEP_F2_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50227">
        <property type="entry name" value="G_PROTEIN_RECEP_F2_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50261">
        <property type="entry name" value="G_PROTEIN_RECEP_F2_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50221">
        <property type="entry name" value="GAIN_B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50092">
        <property type="entry name" value="TSP1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="9" id="KW-0297">G-protein coupled receptor</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10 15" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="12" id="KW-0675">Receptor</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="14" id="KW-0807">Transducer</keyword>
    <keyword evidence="5 15" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 15" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="12"/>
            <end position="37"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="533"/>
            <end position="555"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="567"/>
            <end position="585"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="597"/>
            <end position="615"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="636"/>
            <end position="655"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="675"/>
            <end position="698"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="719"/>
            <end position="741"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="747"/>
            <end position="768"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="775"/>
            <end position="794"/>
        </location>
    </feature>
    <feature type="domain" description="G-protein coupled receptors family 2 profile 1" evidence="17">
        <location>
            <begin position="205"/>
            <end position="275"/>
        </location>
    </feature>
    <feature type="domain" description="GAIN-B" evidence="16">
        <location>
            <begin position="397"/>
            <end position="523"/>
        </location>
    </feature>
    <feature type="domain" description="G-protein coupled receptors family 2 profile 2" evidence="18">
        <location>
            <begin position="531"/>
            <end position="772"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007343"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023224"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50227"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50261"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030220.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1084" mass="122473" checksum="A03EF3899077FE08" modified="2019-05-08" version="1">PNPNVYPELNNILLVHIYLIPRTFPFNIFFVFIFNLLCYRCPVCFVTVHGVWEEWSPWSLCSFTCGRGQRTRTRSCIPPQYGGRSCDGPETQHKPCNIALCPVDGQWQEWSSWSQCSVTCSNGTQQRSRQCTAAAHGGSECRGPWAESRECHNPECTANGQWNQWGHWSGCSKSCDGGWERRIRICQGAAVTGQQCEGTGEEIRRCSEQRCPAPYEICPEDYLLSMVWKRTPAGDMAFNRCPLNATGTASRRCSLSVHGVAFWEHPSFARCISNEYRHLQHSIKEHLAKGQRMLAGDGMSQVTKTLLDLTQRKNFYAGDLLISVEILRNVTDTFKRASYIPASDGVQNFFQIVSNLLDEENKEKWEDAQQIYPGSVELMQVIEDFIHIVGMGMMDFQNSYLMTGNVVASIQKLPAASVLTDINFPMKGRKGMVDWARNSEDRVVIPKNIFTPMSSELDESTVFVLGAVLYKNLELILPTLRNDSLGTWSTQGCKTVLTDASHTKCLCDRLSTFAILAQQPREIIMESSGAPSVTLIVGSGLSCLALITLAVVYAALWRYIRSERSIILINFCLSIISSNILILVGQTQTHNKGICTTTTAFLHFFFLASFCWVLTEAWQSYMAVTGKIRTQLIRKRFLCLGWGLPALVVAISIGFTKTKGYGTSHYCWLSLEGGLLYAFVGPAAAVVLVNMVIGILVFNKLVSRDGILDKKLKHRAGASLWSSCVVLPLLALTWMSAVLAMTDKRSILFQILFAVFDSLQGFVIVMVHCILRREVLWFGFNLIMAVLFHCFLLQTDFEKDVDIACRSVLHKDIGPCRAATITGTLSRISLNDDEEEKGANPEGMSYSTLPGSIISKVIIQQPAGLHMPMNNSELRRTVYLCTDDNLRGADMDIVNPQERMMESDYIVMPRGSVNTQQSLKDESKLNIGMDTLPHERLLHYKVNPEFNMNPSVMDQFNINLDQHLPTQEHMQNLPFEPRTAVKNFIASERKSRYSDLDFEKVMHTRKRHMELFQELNQKFQTLDRFRDVPKSGSMVSILLSSYEFKIYWFVDFKAKDALELRPAEWEKCLNLPLDVQEGDFQTEV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ33</accession>
    <name>A0A452IQ33_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Secreted phosphoprotein 24</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="9">Secreted phosphoprotein 2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">Could coordinate an aspect of bone turnover.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the SPP2 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQ33"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034089.1">
        <property type="protein sequence ID" value="ENSGAGP00000030019.1"/>
        <property type="gene ID" value="ENSGAGG00000021666.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046849">
        <property type="term" value="P:bone remodeling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.450.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046350">
        <property type="entry name" value="Cystatin_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010892">
        <property type="entry name" value="Spp-24"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15444">
        <property type="entry name" value="SECRETED PHOSPHOPROTEIN 24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15444:SF4">
        <property type="entry name" value="SECRETED PHOSPHOPROTEIN 24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07448">
        <property type="entry name" value="Spp-24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54403">
        <property type="entry name" value="Cystatin/monellin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0964">Secreted</keyword>
    <keyword evidence="7 10" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="10">
        <location>
            <begin position="1"/>
            <end position="18"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019027647" description="Secreted phosphoprotein 24" evidence="10">
        <location>
            <begin position="19"/>
            <end position="143"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00002371"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00008576"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00020365"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00029627"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030019.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="143" mass="16219" checksum="AE5EC187B67FE3F5" modified="2019-05-08" version="1">MKTLFVFALVMSVWSCSGLPVYDYEPSITEEALRASIGRVNSRSQWPALFGVVRSYVRGVDLLDNNDYSIVLDFIVRETTCTKDSEKDPSLCDFRMGRYVQTAFCRSTVRVSEEQIQNLAVYCSQDGSSSESSSSEEVRRFHL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ50</accession>
    <name>A0A452IQ50_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="22">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cell projection</location>
            <location evidence="1">Cilium</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the polycystin family.</text>
    </comment>
    <comment type="caution">
        <text evidence="13">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000034148.1">
        <property type="protein sequence ID" value="ENSGAGP00000030076.1"/>
        <property type="gene ID" value="ENSGAGG00000021465.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005929">
        <property type="term" value="C:cilium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005261">
        <property type="term" value="F:monoatomic cation channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006816">
        <property type="term" value="P:calcium ion transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00037">
        <property type="entry name" value="CLECT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00146">
        <property type="entry name" value="PKD"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="CDD" id="cd01752">
        <property type="entry name" value="PLAT_polycystin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000825">
        <property type="entry name" value="Polycystin 1, transient receptor potential channel interacting"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.60.20:FF:000012">
        <property type="entry name" value="polycystin-1 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.70">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.100.10">
        <property type="entry name" value="Mannose-Binding Protein A, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.60.20">
        <property type="entry name" value="PLAT/LH2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001304">
        <property type="entry name" value="C-type_lectin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016186">
        <property type="entry name" value="C-type_lectin-like/link_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016187">
        <property type="entry name" value="CTDL_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057244">
        <property type="entry name" value="GAIN_B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000203">
        <property type="entry name" value="GPS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000434">
        <property type="entry name" value="PC1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022409">
        <property type="entry name" value="PKD/Chitinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002859">
        <property type="entry name" value="PKD/REJ-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013122">
        <property type="entry name" value="PKD1_2_channel"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000601">
        <property type="entry name" value="PKD_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035986">
        <property type="entry name" value="PKD_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001024">
        <property type="entry name" value="PLAT/LH2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036392">
        <property type="entry name" value="PLAT/LH2_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042060">
        <property type="entry name" value="PLAT_polycystin1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006228">
        <property type="entry name" value="Polycystin_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046791">
        <property type="entry name" value="Polycystin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014010">
        <property type="entry name" value="REJ_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002889">
        <property type="entry name" value="WSC_carb-bd"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00864">
        <property type="entry name" value="PCC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46730">
        <property type="entry name" value="POLYCYSTIN-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46730:SF3">
        <property type="entry name" value="POLYCYSTIN-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00059">
        <property type="entry name" value="Lectin_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00801">
        <property type="entry name" value="PKD"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08016">
        <property type="entry name" value="PKD_channel"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01477">
        <property type="entry name" value="PLAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20519">
        <property type="entry name" value="Polycystin_dom"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02010">
        <property type="entry name" value="REJ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00500">
        <property type="entry name" value="POLYCYSTIN1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00034">
        <property type="entry name" value="CLECT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00303">
        <property type="entry name" value="GPS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00308">
        <property type="entry name" value="LH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00089">
        <property type="entry name" value="PKD"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="SMART" id="SM00321">
        <property type="entry name" value="WSC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56436">
        <property type="entry name" value="C-type lectin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49723">
        <property type="entry name" value="Lipase/lipooxygenase domain (PLAT/LH2 domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49299">
        <property type="entry name" value="PKD domain"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50041">
        <property type="entry name" value="C_TYPE_LECTIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50221">
        <property type="entry name" value="GAIN_B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50093">
        <property type="entry name" value="PKD"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50095">
        <property type="entry name" value="PLAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51111">
        <property type="entry name" value="REJ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51212">
        <property type="entry name" value="WSC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="12" id="KW-0966">Cell projection</keyword>
    <keyword evidence="8" id="KW-0969">Cilium</keyword>
    <keyword evidence="10" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9 15" id="KW-0472">Membrane</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5 15" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 15" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="2899"/>
            <end position="2921"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3107"/>
            <end position="3128"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3148"/>
            <end position="3170"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3372"/>
            <end position="3394"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3400"/>
            <end position="3419"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3487"/>
            <end position="3506"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3750"/>
            <end position="3775"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3795"/>
            <end position="3819"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3845"/>
            <end position="3867"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="3904"/>
            <end position="3929"/>
        </location>
    </feature>
    <feature type="domain" description="WSC" evidence="21">
        <location>
            <begin position="18"/>
            <end position="111"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="17">
        <location>
            <begin position="137"/>
            <end position="199"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="16">
        <location>
            <begin position="258"/>
            <end position="372"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="17">
        <location>
            <begin position="795"/>
            <end position="857"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="17">
        <location>
            <begin position="887"/>
            <end position="959"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="17">
        <location>
            <begin position="977"/>
            <end position="1039"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="17">
        <location>
            <begin position="1148"/>
            <end position="1221"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="17">
        <location>
            <begin position="1250"/>
            <end position="1305"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="17">
        <location>
            <begin position="1323"/>
            <end position="1387"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="17">
        <location>
            <begin position="1559"/>
            <end position="1645"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="17">
        <location>
            <begin position="1662"/>
            <end position="1722"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="17">
        <location>
            <begin position="1898"/>
            <end position="1981"/>
        </location>
    </feature>
    <feature type="domain" description="REJ" evidence="20">
        <location>
            <begin position="1984"/>
            <end position="2675"/>
        </location>
    </feature>
    <feature type="domain" description="GAIN-B" evidence="19">
        <location>
            <begin position="2696"/>
            <end position="2889"/>
        </location>
    </feature>
    <feature type="domain" description="PLAT" evidence="18">
        <location>
            <begin position="2944"/>
            <end position="3059"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="3982"/>
            <end position="4011"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="4065"/>
            <end position="4085"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="3999"/>
            <end position="4011"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004138"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007200"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023069"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00152"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50041"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50093"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50095"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS51111"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS51212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030076.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="4116" mass="455672" checksum="1F7C735578AA70C6" modified="2019-05-08" version="1">[encoded data, 4116 characters]</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ51</accession>
    <name>A0A452IQ51_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">SH2 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
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        <property type="protein sequence ID" value="ENSGAGP00000030240.1"/>
        <property type="gene ID" value="ENSGAGG00000021795.1"/>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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        <property type="match status" value="1"/>
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        <property type="entry name" value="B-cell linker protein isoform 2"/>
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        <property type="match status" value="1"/>
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        <property type="entry name" value="Transcription Factor, Ets-1"/>
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        <property type="entry name" value="SAM/pointed_sf"/>
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        <property type="entry name" value="SH2"/>
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        <property type="entry name" value="SH2_dom_sf"/>
    </dbReference>
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        <property type="entry name" value="B-CELL LINKER PROTEIN"/>
        <property type="match status" value="1"/>
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        <property type="match status" value="1"/>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="SH2"/>
        <property type="match status" value="1"/>
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    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1 2" id="KW-0727">SH2 domain</keyword>
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        <location>
            <begin position="492"/>
            <end position="599"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="103"/>
            <end position="458"/>
        </location>
    </feature>
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        <location>
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            <end position="142"/>
        </location>
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            <begin position="358"/>
            <end position="372"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="410"/>
            <end position="431"/>
        </location>
    </feature>
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        <source>
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        </source>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
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        <source>
            <dbReference type="PROSITE" id="PS50001"/>
        </source>
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    <evidence type="ECO:0000313" key="5">
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        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="602" mass="67509" checksum="9FD970F20B9169F2" modified="2019-05-08" version="1">MSVKQSIYCHQKHIKSSSPDSKMATKLPLREECENWTTLQVADLLRQIRMPESAAVVEKLDINGAKFLNISDYELNKFNIMHQPQLQKMVHDIKKNETGIMSKFKKLKNKPPPSVPQRDYASEHAEDEDEQWSDDFDSDYENPDAHSDSEMYVVPSEENCDDSYEPPPTEQEKKKIPSSFSFSRGEYADNRTTQQEPLPSRPSQSLIKSKKPSKPSLPSSSVPLKPKVPPKTKEFNEDEDDYIVPVDDEDNYIEPTEGSTSQSTKPPINRSTKPTKPVLPSSAKSSPAPVSPFVQDVYEVPEEEEKPSPPASRITKPLPPKPVQSTGGHSHMLSSAKESSKPDTSRNILPLPRNHPQQKAEHDRNEEHDENHSSSGAQESKFPSVATPSPLPRALKKTPNPVKPPKPSLSHKENLSVTEEKPTAAERRRGPSQELPLPPLPTTAQKPLLQKPPALPKPLDTANRTMGTLPRSSISASLDTTDQDAGVHDKAWYAAYCDRKTAEDALYKSNKDGSFLIRKSSGQDSKQPYTLVVFYNKKVYNIPVRFIESTKRYALGREKTGEERFTSVAEIIENHQHTPLVLIDSQNNTKDSTKLKHIVRVS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ53</accession>
    <name>A0A452IQ53_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="20">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Cell cortex</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the FNBP1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQ53"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034111.1">
        <property type="protein sequence ID" value="ENSGAGP00000030040.1"/>
        <property type="gene ID" value="ENSGAGG00000021677.1"/>
    </dbReference>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
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        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Formin-binding protein 1-like isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
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        <property type="entry name" value="F_BAR_dom"/>
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        <property type="entry name" value="FCH_dom"/>
    </dbReference>
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        <property type="entry name" value="FNBP1_F-BAR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035492">
        <property type="entry name" value="FNBP1_SH3"/>
    </dbReference>
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        <property type="entry name" value="HR1_rho-bd"/>
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    <dbReference type="InterPro" id="IPR057870">
        <property type="entry name" value="HR1_TOCA"/>
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        <property type="entry name" value="SH3-like_dom_sf"/>
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        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15735">
        <property type="entry name" value="FCH AND DOUBLE SH3 DOMAINS PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15735:SF13">
        <property type="entry name" value="FORMIN-BINDING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00611">
        <property type="entry name" value="FCH"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="HR1_TOCA"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="REM_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
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        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="9 14 15" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="12" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="8" id="KW-0254">Endocytosis</keyword>
    <keyword evidence="10" id="KW-0446">Lipid-binding</keyword>
    <keyword evidence="11" id="KW-0472">Membrane</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 13" id="KW-0728">SH3 domain</keyword>
    <feature type="domain" description="F-BAR" evidence="18">
        <location>
            <begin position="1"/>
            <end position="264"/>
        </location>
    </feature>
    <feature type="domain" description="REM-1" evidence="19">
        <location>
            <begin position="434"/>
            <end position="511"/>
        </location>
    </feature>
    <feature type="domain" description="SH3" evidence="17">
        <location>
            <begin position="580"/>
            <end position="641"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="293"/>
            <end position="356"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="519"/>
            <end position="563"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="15">
        <location>
            <begin position="441"/>
            <end position="501"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="525"/>
            <end position="542"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004544"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00009426"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022583"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023121"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01077"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS51741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS51860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030040.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="651" mass="75415" checksum="2FD39D5658B23782" modified="2019-05-08" version="1">MSWGTELWDQFDNLEKHTQWGIDVLEKYIKFVKERTEIEISYAKQLRNLSKKYQPKKNSKEEEEYKYTSCKAFLATLNEMNDYAGQHEVISENMTSQITAELARFVQELKQERKSHFHDGRKAQQYIETFWKQLEASKRRFERDCKEADRAQQYFEKMDADINVTKADVEKARQQAQIRHQMAEDSKAEYSSILQKFNHDQHEHYHTHIPNIFQKIQEMEERRIVRIGECMKTYAEVDRQVIPIIGKCLDGITKAAESIDQKNDSLLVIEAFKSGFEPPGDIDFEDFTQPMKRTVSESSLSNSRWDGKSEPKPGSKSKGKLWPFIKKNKPPPPPPASASPSAVPNGPQSPKQQKEPLSHRFNEFMTSKPKIHCFRSLKRGNHSLYFHKELMKRTLGKPTCAVEARDYVLSLKLITLNTQVWNLIEGGGPEDFSNLPPEQRRKKLQQKVDELNKEIQKEMDQRDALTKMKDVYIKNPQMGDAASVDHKLAELGQNMEKLRLEAQKFEGWLAEVEGRLPVRTEQSRRQSGMYDTQNTSGVNNCAQDRESPDGSYTEEPSQESEMKVPVTEFDDEFDDEEPLPTIGTCKALYTFEGQNEGTISVMEGEMLYVIEEDKGDGWTRIRRNEDEEGYVPTSYVEVYLDKNAKGAMTYI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ55</accession>
    <name>A0A452IQ55_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2 11">Arginase</fullName>
            <ecNumber evidence="2 11">3.5.3.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="8 11">
            <text>L-arginine + H2O = urea + L-ornithine</text>
            <dbReference type="Rhea" id="RHEA:20569"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:16199"/>
            <dbReference type="ChEBI" id="CHEBI:32682"/>
            <dbReference type="ChEBI" id="CHEBI:46911"/>
            <dbReference type="EC" id="3.5.3.1"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="11">
            <name>Mn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29035"/>
        </cofactor>
        <text evidence="11">Binds 2 manganese ions per subunit.</text>
    </comment>
    <comment type="pathway">
        <text evidence="1 11">Nitrogen metabolism; urea cycle; L-ornithine and urea from L-arginine: step 1/1.</text>
    </comment>
    <comment type="similarity">
        <text evidence="9 10">Belongs to the arginase family.</text>
    </comment>
    <dbReference type="EC" id="3.5.3.1" evidence="2 11"/>
    <dbReference type="AlphaFoldDB" id="A0A452IQ55"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034153.1">
        <property type="protein sequence ID" value="ENSGAGP00000030081.1"/>
        <property type="gene ID" value="ENSGAGG00000021614.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00158">
        <property type="reaction ID" value="UER00270"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004053">
        <property type="term" value="F:arginase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030145">
        <property type="term" value="F:manganese ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010121">
        <property type="term" value="P:L-arginine catabolic process to proline via ornithine"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000050">
        <property type="term" value="P:urea cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="CDD" id="cd09989">
        <property type="entry name" value="Arginase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.800.10:FF:000005">
        <property type="entry name" value="Arginase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.800.10">
        <property type="entry name" value="Ureohydrolase domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014033">
        <property type="entry name" value="Arginase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006035">
        <property type="entry name" value="Ureohydrolase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023696">
        <property type="entry name" value="Ureohydrolase_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020855">
        <property type="entry name" value="Ureohydrolase_Mn_BS"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01229">
        <property type="entry name" value="rocF_arginase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43782">
        <property type="entry name" value="ARGINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43782:SF2">
        <property type="entry name" value="ARGINASE-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00491">
        <property type="entry name" value="Arginase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00116">
        <property type="entry name" value="ARGINASE"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52768">
        <property type="entry name" value="Arginase/deacetylase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01053">
        <property type="entry name" value="ARGINASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51409">
        <property type="entry name" value="ARGINASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 11" id="KW-0056">Arginine metabolism</keyword>
    <keyword evidence="6 10" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="7 11" id="KW-0464">Manganese</keyword>
    <keyword evidence="5 11" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 11" id="KW-0835">Urea cycle</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00005098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00012168"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022436"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022503"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023211"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00047391"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00742"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="RuleBase" id="RU003684"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="RuleBase" id="RU361159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030081.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="284" mass="30744" checksum="968313926530C839" modified="2019-05-08" version="1">MIWPEVVKKAGLIDKLTAQGCDVKDRELKIDIIPNDLPHHNAKNPRTVGCANEKLASAVEEVKKDGRTCLVVGGDHSLAIGSISGHAKTEPDLGVIWFDAHADINTPLTSLSGNIHGQPVAFLLKELKVSGVPGFKWLKPCLSVENIVYIGLRDVDKAEHYFLKSLGIKYYSMTEIDQLGIGKVMEEALDYLLANKKRPIHLSFDVDGLDPSLTPATGTPVPGGVTLREGIYVAEQLYKTGLLSAVDIVEINPRLGKTQEGVDSTVNTAISIILSCFGKPQKSI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ56</accession>
    <name>A0A452IQ56_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">SH2 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IQ56"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034327.1">
        <property type="protein sequence ID" value="ENSGAGP00000030245.1"/>
        <property type="gene ID" value="ENSGAGG00000021795.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007169">
        <property type="term" value="P:cell surface receptor protein tyrosine kinase signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035556">
        <property type="term" value="P:intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd09929">
        <property type="entry name" value="SH2_BLNK_SLP-76"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.505.10:FF:000016">
        <property type="entry name" value="B-cell linker protein isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.505.10">
        <property type="entry name" value="SH2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051751">
        <property type="entry name" value="Immunoreceptor_sig_adapters"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000980">
        <property type="entry name" value="SH2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036860">
        <property type="entry name" value="SH2_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14098:SF3">
        <property type="entry name" value="B-CELL LINKER PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14098">
        <property type="entry name" value="SH2 DOMAIN CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00017">
        <property type="entry name" value="SH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00252">
        <property type="entry name" value="SH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55550">
        <property type="entry name" value="SH2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50001">
        <property type="entry name" value="SH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1 2" id="KW-0727">SH2 domain</keyword>
    <feature type="domain" description="SH2" evidence="4">
        <location>
            <begin position="443"/>
            <end position="550"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="50"/>
            <end position="411"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="3">
        <location>
            <begin position="76"/>
            <end position="93"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="3">
        <location>
            <begin position="165"/>
            <end position="176"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="3">
        <location>
            <begin position="187"/>
            <end position="203"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="208"/>
            <end position="225"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="3">
        <location>
            <begin position="227"/>
            <end position="243"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="274"/>
            <end position="288"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="309"/>
            <end position="323"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="361"/>
            <end position="382"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022999"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00191"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50001"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030245.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="553" mass="61895" checksum="348207845D1ECD32" modified="2019-05-08" version="1">MDKLNKITEPAGQKLRQLQKMVHDIKKNETGIMSKFKKFQNEQVALICKTGKNTWDRLKNKPPPSVPQRDYASEHAEDEDEQWSDDFDSDYENPDAHSDSEMYVVPSEENCDDSYEPPPTEQEKKKIPSSFSFSRGEYADNRTTQQEPLPSRPSQSLIKSKKPSKPSLPSSSVPLKPKVPPKTKEFNEDEDDYIVPVDDEDNYIEPTEGSTSQSTKPPINRSTKPTKPVLPSSAKSSPAPVSPFVQDVYEVPEEEEKPSPPASRITKPLPPKPVQSTGGHSHMLSSAKESSKPDTSRNILPLPRNHPQQKAEHDRNEEHDENHSSSGAQESKFPSVATPSPLPRALKKTPNPVKPPKPSLSHKENLSVTEEKPTAAERRRGPSQELPLPPLPTTAQKPLLQKPPALPKPLDTANRTMGTLPRSSISASLDTTDQDAGVHDKAWYAAYCDRKTAEDALYKSNKDGSFLIRKSSGQDSKQPYTLVVFYNKKVYNIPVRFIESTKRYALGREKTGEERFTSVAEIIENHQHTPLVLIDSQNNTKDSTKLKHIVRVS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ57</accession>
    <name>A0A452IQ57_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="20">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Cell cortex</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the FNBP1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQ57"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034116.1">
        <property type="protein sequence ID" value="ENSGAGP00000030045.1"/>
        <property type="gene ID" value="ENSGAGG00000021677.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005938">
        <property type="term" value="C:cell cortex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008289">
        <property type="term" value="F:lipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006897">
        <property type="term" value="P:endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd11629">
        <property type="entry name" value="HR1_FBP17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12071">
        <property type="entry name" value="SH3_FBP17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1270.60:FF:000002">
        <property type="entry name" value="Formin-binding protein 1-like isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.40:FF:000017">
        <property type="entry name" value="Formin-binding protein 1-like isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.470">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.60">
        <property type="entry name" value="Arfaptin homology (AH) domain/BAR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027267">
        <property type="entry name" value="AH/BAR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031160">
        <property type="entry name" value="F_BAR_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001060">
        <property type="entry name" value="FCH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035492">
        <property type="entry name" value="FNBP1_SH3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011072">
        <property type="entry name" value="HR1_rho-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057870">
        <property type="entry name" value="HR1_TOCA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15735">
        <property type="entry name" value="FCH AND DOUBLE SH3 DOMAINS PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15735:SF13">
        <property type="entry name" value="FORMIN-BINDING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00611">
        <property type="entry name" value="FCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25610">
        <property type="entry name" value="HR1_TOCA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00018">
        <property type="entry name" value="SH3_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00055">
        <property type="entry name" value="FCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00326">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103657">
        <property type="entry name" value="BAR/IMD domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51741">
        <property type="entry name" value="F_BAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51860">
        <property type="entry name" value="REM_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="9 14 15" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="12" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="8" id="KW-0254">Endocytosis</keyword>
    <keyword evidence="10" id="KW-0446">Lipid-binding</keyword>
    <keyword evidence="11" id="KW-0472">Membrane</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 13" id="KW-0728">SH3 domain</keyword>
    <feature type="domain" description="F-BAR" evidence="18">
        <location>
            <begin position="1"/>
            <end position="245"/>
        </location>
    </feature>
    <feature type="domain" description="REM-1" evidence="19">
        <location>
            <begin position="413"/>
            <end position="490"/>
        </location>
    </feature>
    <feature type="domain" description="SH3" evidence="17">
        <location>
            <begin position="559"/>
            <end position="620"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="274"/>
            <end position="301"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="314"/>
            <end position="347"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="498"/>
            <end position="542"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="15">
        <location>
            <begin position="420"/>
            <end position="480"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="16">
        <location>
            <begin position="319"/>
            <end position="328"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="504"/>
            <end position="521"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004544"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00009426"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022583"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023121"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01077"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS51741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS51860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030045.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="630" mass="72745" checksum="AF17712224FF3238" modified="2019-05-08" version="1">MSWGTELWDQFDNLEKHTQWGIDVLEKYIKFVKERTEIEISYAKQLRYTSCKAFLATLNEMNDYAGQHEVISENMTSQITAELARFVQELKQERKSHFHDGRKAQQYIETFWKQLEASKRRFERDCKEADRAQQYFEKMDADINVTKADVEKARQQAQIRHQMAEDSKAEYSSILQKFNHDQHEHYHTHIPNIFQKIQEMEERRIVRIGECMKTYAEVDRQVIPIIGKCLDGITKAAESIDQKNDSLLVIEAFKSGFEPPGDIDFEDFTQPMKRTVSESSLSNSRWDGKSEPKPGSKSKGKLWPFIKKNKLMSLLTSPHQPPPPPPASASPSAVPNGPQSPKQQKEPLSHRFNEFMTSKPKIHCFRSLKRGNHSLYFHKELMKRTLGKPTCAVEARDYVLSLKLGGGPEDFSNLPPEQRRKKLQQKVDELNKEIQKEMDQRDALTKMKDVYIKNPQMGDAASVDHKLAELGQNMEKLRLEAQKFEGWLAEVEGRLPVRTEQSRRQSGMYDTQNTSGVNNCAQDRESPDGSYTEEPSQESEMKVPVTEFDDEFDDEEPLPTIGTCKALYTFEGQNEGTISVMEGEMLYVIEEDKGDGWTRIRRNEDEEGYVPTSYVEVYLDKNAKGAMTYI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ63</accession>
    <name>A0A452IQ63_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">SH3 and PX domain-containing protein 2A</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell projection</location>
            <location evidence="1">Podosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the SH3PXD2 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQ63"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034163.1">
        <property type="protein sequence ID" value="ENSGAGP00000030091.1"/>
        <property type="gene ID" value="ENSGAGG00000021705.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002102">
        <property type="term" value="C:podosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016176">
        <property type="term" value="F:superoxide-generating NADPH oxidase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042554">
        <property type="term" value="P:superoxide anion generation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd12074">
        <property type="entry name" value="SH3_Tks5_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12077">
        <property type="entry name" value="SH3_Tks5_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12019">
        <property type="entry name" value="SH3_Tks5_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12020">
        <property type="entry name" value="SH3_Tks5_5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.40:FF:000020">
        <property type="entry name" value="SH3 and PX domain-containing protein 2A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.40:FF:000031">
        <property type="entry name" value="SH3 and PX domain-containing protein 2A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.40:FF:000042">
        <property type="entry name" value="SH3 and PX domain-containing protein 2A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.40:FF:000059">
        <property type="entry name" value="SH3 and PX domain-containing protein 2A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.40:FF:000141">
        <property type="entry name" value="SH3 and PX domain-containing protein 2A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051228">
        <property type="entry name" value="NADPH_Oxidase/PX-Domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035450">
        <property type="entry name" value="SH3PXD2A_SH3_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035452">
        <property type="entry name" value="SH3PXD2A_SH3_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035453">
        <property type="entry name" value="SH3PXD2A_SH3_4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035454">
        <property type="entry name" value="SH3PXD2A_SH3_5"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15706:SF2">
        <property type="entry name" value="SH3 AND PX DOMAIN-CONTAINING PROTEIN 2A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15706">
        <property type="entry name" value="SH3 MULTIPLE DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00018">
        <property type="entry name" value="SH3_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07653">
        <property type="entry name" value="SH3_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00326">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="5"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0965">Cell junction</keyword>
    <keyword evidence="9" id="KW-0966">Cell projection</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="4 11" id="KW-0728">SH3 domain</keyword>
    <feature type="domain" description="SH3" evidence="13">
        <location>
            <begin position="35"/>
            <end position="94"/>
        </location>
    </feature>
    <feature type="domain" description="SH3" evidence="13">
        <location>
            <begin position="135"/>
            <end position="194"/>
        </location>
    </feature>
    <feature type="domain" description="SH3" evidence="13">
        <location>
            <begin position="316"/>
            <end position="375"/>
        </location>
    </feature>
    <feature type="domain" description="SH3" evidence="13">
        <location>
            <begin position="684"/>
            <end position="743"/>
        </location>
    </feature>
    <feature type="domain" description="SH3" evidence="13">
        <location>
            <begin position="913"/>
            <end position="974"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="284"/>
            <end position="314"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="371"/>
            <end position="658"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="822"/>
            <end position="894"/>
        </location>
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        <location>
            <begin position="409"/>
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        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="441"/>
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        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="475"/>
            <end position="488"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
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            <begin position="502"/>
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        </location>
    </feature>
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        <location>
            <begin position="557"/>
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        <location>
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    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="882"/>
            <end position="892"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004188"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00009628"/>
        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
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        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
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            <dbReference type="ARBA" id="ARBA00072115"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030091.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="974" mass="108513" checksum="E2F16BE847655DD7" modified="2019-05-08" version="1">MNWMDGGRSRGAMAGSWLTACFFPSPGADASSEPMILEQYVVVSSYEKQENSEISLQAGEVVDVIEKNESGWWFVSTAEEQGWVPATYLESQNGTRDDSEINTSKAGEVSKRRKAHLRRLDRRWTLGGIVNRQQSREEKYVTIQPYTSQGKDEIGFEKGVTVEVIQKNLEGWWYIRHLGKEGWAPASYLKKAKDDLPSRKKNLTGPVEIIGNIMEISNLLNKKPSSDKETQAENEMAESHITKKEISLPILCNDSNGNAMMTSDKQASKLAQGSPAVARIAPQRAQISSPNLRTRPPPRRESSLGFQLPKPPEPPSVEVEYYTIAEFQSCISDGISFRGGQKAEVIDKNSGGWWYVQIGEKEGWAPASYIDKRKKPNLNRRTSTLTRPKVPPPAPPSKPKDTEEGTAPGTVSSGEVQDSPQKLRYEEPEYDVPAFGFDSESEVHESHGDDSSLEKRLFQPLKPSPVSSLQRAKFRVGESSEDVAREEEIIYENEGFRPYVEDTSSCKESSGDSDSQKSSSVSLIRRNSPCSSSPKSSLLKLKSDKSIQPDLGKCHYSRNDEIKPRSASDVGLRSMPRTSAKKEPGQSPSTKAKPMVRPKPFLNKADSQSQEKMDISTLRRQLRPTGQLRGGLQGSRSEESESPPQIAPESQDDSLKRSSADLITSPSHAIFCSNHPAKPEQENDTKCFYVTSDSYQKVQDSELCFPAGVEVEVLEKQASGWWYMKYGDMEGWAPSHYLVLADNQKEETSETDISLARNRRNESKSNSLEKIEKRVQALNTINQSKRAMPPIPSKPPGGFSKTSGAVKMRNGVRQLAVRPQSVFVSPPPKDNNLSCSLRRNESLTTTDHLRAVRRNSSFSSARSQPGDSKAKQTDQSGIEGLETTSNLSTQRNGIPVSTVWPKPLEKSQFIHNNLKDIYVSIADYEGDEETAGFQEGVCMEVLERNPNGWWYCQIMDGGKPFKGWVPSNYLEKKN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1N2</accession>
    <name>A0A452J1N2_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J1N2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038736.1">
        <property type="protein sequence ID" value="ENSGAGP00000034209.1"/>
        <property type="gene ID" value="ENSGAGG00000024299.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="entry name" value="NHS-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23039">
        <property type="entry name" value="NANCE-HORAN SYNDROME PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23039:SF6">
        <property type="entry name" value="SIMILAR TO MKIAA1522 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15273">
        <property type="entry name" value="NHS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
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        <location>
            <begin position="43"/>
            <end position="243"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
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        </location>
    </feature>
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            <begin position="458"/>
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            <begin position="78"/>
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            <begin position="377"/>
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        <location>
            <begin position="492"/>
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    </feature>
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        <location>
            <begin position="1226"/>
            <end position="1240"/>
        </location>
    </feature>
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        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034209.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1270" mass="133489" checksum="2A48ADE07ED4B5B3" modified="2019-05-08" version="1">MGNMHHKRNPPNAKARSFWHFGHTGKLKTGSAKAENKRLNVQYKAGEERPDNVFFPSSRPPHLEELHNQAQAGLKSLQHQEKQKQTRSGWDHGDSNSIQSSHTSAADDEISFRSRTQSCTTENASEDALSIRSEMIQRKGSTFRPHDSFPKSSEKAGKRRKERRTTVLGIPQHVQKELGLRNSHELRKRPGSVSPQDGPIQAGSPHSLAPPTLVNGGEADGGMIHIPTINGNLQPPPAPKGGARVSLQALEEAESDAAIQRHINCVYYDDTLLGRRTAAKLSPLLRPKSLAVPGMTTNPSPPPELFGPVMSISPQATYMSKIIPNAVLPPMVDVIALSRSSVQTLSRCSLVSSSPASVRSLTRFSEPSARSREHSSSSDNWSHSQSTETIVSDSSTISSQGGSNSRKGEAGPPSEADTAGRSDTDQLSVYSAASCTNSCSQARPVYTAHGLLAVGPGGSSGRASPAYSTSSMADGSDTVSVRSDRSSTRSVSLRKMKKPPAPPRRTHSLHQKAQQHQQGVGEGGQKVLGLPPKPDPRPQREPWTPRLESQSPLGEDEVFSPCSLSETSSIRSDSLAYSADASSPDVSRCSPVPGEKLRREVEGVAVIFREQQATPRQSSPEGFKRTMSPSSGYSSQSGTPTLPTKGLGPHASSPGKRRPQPKKPERVCSLQSPALSISSSLTSLSSSTSDPAPAETVAPPAGPALSWQSRMDRFIIPPHPKVLAPISPPPIKPQQAAPSASPTVSSPAPSTASQKPSALPKPSSRSPPPSPPPPYHPPPPPVKKAGTSPEAPHSETIQEAPQDPSWPPPPPPALDEQDLSMADFPPPDEAGFSTLPELLPVTGTAGASSVAAQPEPVGSEQPARPKAAPRGDGPGAASLAISAKVSSRTQQQQGPLAGSALPAPAVELPMPTDMSPSARATPDLQPGQANPAVPAAPPHAPAAPAPPTALQPQASLKKTTNGSQLDPKKELVSRSKSSPTPKEDANLPIVTPSLLQMVRLRSVNMDGHAPAGPGKQSPGQNGQGADGVGKQPQQVPQKPIRKSLSLRQSPSSKDTVPLNQLQNAVRLKTSTLSSRDAPTSRLAGWTNGNKLTLPGHTASSSEHTTGDAKDGQVSPIHKSPASTASFIFAKSSKKLVIETPSSPEAQADLKRNLVAELMNFAGQRSTVPAMTQQGQVPPGKSQAQRKSSKIPPPIARKPLLGMGQPQSPVSPKPQGEEVLNCSLPDSKAKASSAEENRTKSELSVNAAPQADSSRAGHLAGSATPSQNPPA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1N9</accession>
    <name>A0A452J1N9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">HMG box domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
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                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the DNA mismatch repair MutL/HexB family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1N9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038837.1">
        <property type="protein sequence ID" value="ENSGAGP00000034298.1"/>
        <property type="gene ID" value="ENSGAGG00000024375.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032389">
        <property type="term" value="C:MutLalpha complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
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        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140664">
        <property type="term" value="F:ATP-dependent DNA damage sensor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030983">
        <property type="term" value="F:mismatched DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006298">
        <property type="term" value="P:mismatch repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd16926">
        <property type="entry name" value="HATPase_MutL-MLH-PMS-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd03485">
        <property type="entry name" value="MutL_Trans_hPMS_1_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.30.10:FF:000026">
        <property type="entry name" value="PMS1 homolog 1, mismatch repair system component"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.230.10:FF:000030">
        <property type="entry name" value="PMS1 homolog 1, mismatch repair system component"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.565.10:FF:000017">
        <property type="entry name" value="PMS1 homolog 1, mismatch repair system component"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.230.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.30.10">
        <property type="entry name" value="High mobility group box domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.565.10">
        <property type="entry name" value="Histidine kinase-like ATPase, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014762">
        <property type="entry name" value="DNA_mismatch_repair_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013507">
        <property type="entry name" value="DNA_mismatch_S5_2-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036890">
        <property type="entry name" value="HATPase_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009071">
        <property type="entry name" value="HMG_box_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036910">
        <property type="entry name" value="HMG_box_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002099">
        <property type="entry name" value="MutL/Mlh/PMS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038973">
        <property type="entry name" value="MutL/Mlh/Pms-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020568">
        <property type="entry name" value="Ribosomal_Su5_D2-typ_SF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014721">
        <property type="entry name" value="Ribsml_uS5_D2-typ_fold_subgr"/>
    </dbReference>
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        <property type="entry name" value="mutl"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10073">
        <property type="entry name" value="DNA MISMATCH REPAIR PROTEIN MLH, PMS, MUTL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10073:SF54">
        <property type="entry name" value="PMS1 PROTEIN HOMOLOG 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01119">
        <property type="entry name" value="DNA_mis_repair"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13589">
        <property type="entry name" value="HATPase_c_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00505">
        <property type="entry name" value="HMG_box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01340">
        <property type="entry name" value="DNA_mis_repair"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00398">
        <property type="entry name" value="HMG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55874">
        <property type="entry name" value="ATPase domain of HSP90 chaperone/DNA topoisomerase II/histidine kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47095">
        <property type="entry name" value="HMG-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54211">
        <property type="entry name" value="Ribosomal protein S5 domain 2-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00058">
        <property type="entry name" value="DNA_MISMATCH_REPAIR_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50118">
        <property type="entry name" value="HMG_BOX_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0227">DNA damage</keyword>
    <keyword evidence="3" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0539">Nucleus</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="HMG box" evidence="5">
        <location>
            <begin position="580"/>
            <end position="648"/>
        </location>
    </feature>
    <feature type="DNA-binding region" description="HMG box" evidence="3">
        <location>
            <begin position="580"/>
            <end position="648"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="376"/>
            <end position="400"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="461"/>
            <end position="482"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="535"/>
            <end position="560"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="641"/>
            <end position="668"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="384"/>
            <end position="400"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="468"/>
            <end position="478"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="550"/>
            <end position="560"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="641"/>
            <end position="662"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006082"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00267"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50118"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034298.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="902" mass="101927" checksum="85D825276D464117" modified="2019-05-08" version="1">MKQLSAETVRLLSSSQVITSVVSVVKELIENSLDANATSIDVKLENYGFDKIEVRDNGDGIKAADVPVMAVKHYTSKISCSEDLESLTTYGFRGEALGSICCISEVLITTKTAADDFSTQYILDSSGHVTSQKPSHLGQGTTVAALKLFKNLPVRKQFYSTVKKCKEELKKIQDLLMAYGIIKPELRIIFTHNKAVIWQKTRVSDHKMSFMSVLGTAVMGSMVPFQHRCEDPEVFLSGFLPKPESDNSLTSLSSSERSFIYINNRPVHQKEILKLVRWFYNLKSHKDSSRSYPVFFINITVPTSAVDVNLTPDKTQVLLHNKESVLSAVENVLTSLYGLPPDTVSCETNKTDVTSADVVVNETEQTDVLVNKMEPFGNSDPHAHTSSFSSSNDVQNGQAGKNTEVCLKHQRFSSDNTHGLLNRKEVSGTDVTVSDISQDSSINNLSCEDSQNGYSVTLTLDGSTHMDTQSKNRSEHPLRASYANEVEKNEANTVPEYVSEISADNWSKGNALKNATGENLEPVKILIPEVGGITDHKANENTDEQDRDQQSSNQNIKKTNVVNDKAGQIMAYDLISNQIIRKPMSAVALFTQEYRSRLLTDHPKANMEDLMLKIEGLWEKLSREEKKKYEEKATKDLERYNRQTKKAVEQRIQRSTKETEKRYKSKLKTSLSDQQKLDKLFQSQIEKKQRHNQDMKIVQVPFSMSSFKHKLQRLEKKNFDKNESCLICHLSFPDAWIIASEKKIMLLNPYRVEEALLYKKLLEQHKLPVEKLDKAIVLTDSLFSGSHYMDSLYNMTKDCHNLNGSIYLSDPRLIANGFRIKMTPGHSAIENQLEIEGMANCLPFYGISDLKEILNAVINKNAKEVYECRPLKVINYLEKDLIERKKREVVLVRVCVNVKLSR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1P8</accession>
    <name>A0A452J1P8_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="13">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="8">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1P8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038851.1">
        <property type="protein sequence ID" value="ENSGAGP00000034310.1"/>
        <property type="gene ID" value="ENSGAGG00000024379.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005604">
        <property type="term" value="C:basement membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009887">
        <property type="term" value="P:animal organ morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016358">
        <property type="term" value="P:dendrite development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008045">
        <property type="term" value="P:motor neuron axon guidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009888">
        <property type="term" value="P:tissue development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00055">
        <property type="entry name" value="EGF_Lam"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="CDD" id="cd03579">
        <property type="entry name" value="NTR_netrin-1_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000081">
        <property type="entry name" value="Netrin 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.50.120:FF:000001">
        <property type="entry name" value="Netrin 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000048">
        <property type="entry name" value="Netrin 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.50.120">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.260">
        <property type="entry name" value="Galactose-binding domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050440">
        <property type="entry name" value="Laminin/Netrin_ECM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008211">
        <property type="entry name" value="Laminin_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002049">
        <property type="entry name" value="LE_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056863">
        <property type="entry name" value="LMN_ATRN_NET-like_EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001134">
        <property type="entry name" value="Netrin_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018933">
        <property type="entry name" value="Netrin_module_non-TIMP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008993">
        <property type="entry name" value="TIMP-like_OB-fold"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10574:SF443">
        <property type="entry name" value="NETRIN 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10574">
        <property type="entry name" value="NETRIN/LAMININ-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00053">
        <property type="entry name" value="EGF_laminin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24973">
        <property type="entry name" value="EGF_LMN_ATRN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00055">
        <property type="entry name" value="Laminin_N"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01759">
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00643">
        <property type="entry name" value="C345C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00180">
        <property type="entry name" value="EGF_Lam"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00136">
        <property type="entry name" value="LamNT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50242">
        <property type="entry name" value="TIMP-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01248">
        <property type="entry name" value="EGF_LAM_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50027">
        <property type="entry name" value="EGF_LAM_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51117">
        <property type="entry name" value="LAMININ_NTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50189">
        <property type="entry name" value="NTR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5 8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7 8" id="KW-0424">Laminin EGF-like domain</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="3 9" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="9">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019578755" evidence="9">
        <location>
            <begin position="23"/>
            <end position="552"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin N-terminal" evidence="12">
        <location>
            <begin position="32"/>
            <end position="238"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="10">
        <location>
            <begin position="359"/>
            <end position="410"/>
        </location>
    </feature>
    <feature type="domain" description="NTR" evidence="11">
        <location>
            <begin position="420"/>
            <end position="549"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="8">
        <location>
            <begin position="359"/>
            <end position="371"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="8">
        <location>
            <begin position="361"/>
            <end position="378"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="8">
        <location>
            <begin position="380"/>
            <end position="389"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023292"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00460"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50027"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50189"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS51117"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034310.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="552" mass="61101" checksum="43FE57CA75BEA3F4" modified="2019-05-08" version="1">MPGLWQGLGVLLSLQLALLGLASDPCYDAARRPRYCLPEPTNLAAGRLVQASATCGRPPQRLCAFRHPGNTSSRRCRLCDGATCWQSGPVANASLTLALGGRFELLYVALRFCSPRPHSLALYKSTDHGHSWMPFQFFSARCPQAYGLPLASVISKASEHEAICTSAQTDPTPLVGGLVAFMPLAGRPLARAFEYSPVLQDWVTATDLRVALDRRHRGLGLRGREASYGVSELQVGGRCRCNGHAARCGAAGPSGAPQCQCRHNTAGPECETCRAFYCDRPWQRATPSDAHECVACECNRHSHRCRFNMELYKLSGRKSGGVCLNCRHHTAGRHCHYCQPGFKRDLARPITSRRACKACQCHPIGAVSTMCNQTTGQCQCKMGVTGLTCNRCAQGFQQSRTLTPLYPCAPIYALPLGTECQTHCSPSHGRVHMNLRKYCKKDYVLHAQLLAMVESGEWWQFTTSVLVVYRQRQVPIRRGEQPLWVPRQDLACGCLRLQVGKAYLVIGNDAESPDPARLVLDRNSLALPWRDVWAHKLRRFQQQDRQGNCRSP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1Q1</accession>
    <name>A0A452J1Q1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="23">Sterol regulatory element-binding protein 1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="24">Sterol regulatory element-binding transcription factor 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="31 32">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="32" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="31" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="31" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="26">Key transcription factor that regulates expression of genes involved in cholesterol biosynthesis and lipid homeostasis. Binds to the sterol regulatory element 1 (SRE-1) (5'-ATCACCCCAC-3'). Has dual sequence specificity binding to both an E-box motif (5'-ATCACGTGA-3') and to SRE-1 (5'-ATCACCCCAC-3'). Regulates the promoters of genes involved in cholesterol biosynthesis and the LDL receptor (LDLR) pathway of sterol regulation.</text>
    </comment>
    <comment type="function">
        <text evidence="25">Precursor of the transcription factor form (Processed sterol regulatory element-binding protein 1), which is embedded in the endoplasmic reticulum membrane. Low sterol concentrations promote processing of this form, releasing the transcription factor form that translocates into the nucleus and activates transcription of genes involved in cholesterol biosynthesis and lipid homeostasis.</text>
    </comment>
    <comment type="subunit">
        <text evidence="27">Efficient DNA binding of the soluble transcription factor fragment requires dimerization with another bHLH protein. Interacts with CEBPA, the interaction produces a transcriptional synergy. Interacts with LMNA.</text>
    </comment>
    <comment type="subunit">
        <text evidence="28">Forms a tight complex with SCAP, the SCAP-SREBP complex, in the endoplasmic reticulum membrane and the Golgi apparatus. Interacts with PAQR3; the interaction anchors the SCAP-SREBP complex to the Golgi apparatus in low cholesterol conditions.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cytoplasmic vesicle</location>
            <location evidence="3">COPII-coated vesicle membrane</location>
            <topology evidence="3">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Endoplasmic reticulum membrane</location>
            <topology evidence="2">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Golgi apparatus membrane</location>
            <topology evidence="4">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="22">Belongs to the SREBP family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1Q1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038854.1">
        <property type="protein sequence ID" value="ENSGAGP00000034313.1"/>
        <property type="gene ID" value="ENSGAGG00000024380.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012507">
        <property type="term" value="C:ER to Golgi transport vesicle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000139">
        <property type="term" value="C:Golgi membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046983">
        <property type="term" value="F:protein dimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008203">
        <property type="term" value="P:cholesterol metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd18921">
        <property type="entry name" value="bHLHzip_SREBP1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.280.10:FF:000016">
        <property type="entry name" value="Sterol regulatory element-binding transcription factor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.280.10">
        <property type="entry name" value="Helix-loop-helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011598">
        <property type="entry name" value="bHLH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036638">
        <property type="entry name" value="HLH_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46062">
        <property type="entry name" value="STEROL REGULATORY ELEMENT-BINDING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46062:SF2">
        <property type="entry name" value="STEROL REGULATORY ELEMENT-BINDING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00010">
        <property type="entry name" value="HLH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00353">
        <property type="entry name" value="HLH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47459">
        <property type="entry name" value="HLH, helix-loop-helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50888">
        <property type="entry name" value="BHLH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="16" id="KW-0010">Activator</keyword>
    <keyword evidence="5" id="KW-0153">Cholesterol metabolism</keyword>
    <keyword evidence="21" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="14" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="8" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="12" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="13" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="15" id="KW-0472">Membrane</keyword>
    <keyword evidence="20" id="KW-0539">Nucleus</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="32" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="19" id="KW-0753">Steroid metabolism</keyword>
    <keyword evidence="18" id="KW-1207">Sterol metabolism</keyword>
    <keyword evidence="17" id="KW-0804">Transcription</keyword>
    <keyword evidence="11" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="10" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="9" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="BHLH" evidence="30">
        <location>
            <begin position="292"/>
            <end position="342"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="29">
        <location>
            <begin position="23"/>
            <end position="120"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="29">
        <location>
            <begin position="389"/>
            <end position="437"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="29">
        <location>
            <begin position="469"/>
            <end position="493"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="29">
        <location>
            <begin position="999"/>
            <end position="1033"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="29">
        <location>
            <begin position="36"/>
            <end position="47"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="29">
        <location>
            <begin position="398"/>
            <end position="422"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="29">
        <location>
            <begin position="1022"/>
            <end position="1033"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004477"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004557"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004653"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022548"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023166"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00038460"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00039749"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00042215"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00045313"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00045371"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="ARBA" id="ARBA00047005"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="ARBA" id="ARBA00049702"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="30">
        <source>
            <dbReference type="PROSITE" id="PS50888"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="31">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034313.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="32">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1096" mass="118460" checksum="2A1EA51462FEFB38" modified="2019-05-08" version="1">MSRYHDMLQLINTQDNEFPGLFDTPFGAPAAPMPEASTSVGLPSTPGSLDGYLGPSQAPSSTPVSMYQVPPPLQSFSPQPQLRAGPGLTVSPASDIKEEPMAVSASQPRPLPQPQQGVMTAQSFVPASQGQFTPQPLVGYQGQNGFTAVQPGGAAQSLASLPHSSPQIQPVTLQAHVQNISPQQLLAATTTGTSQAISSQFQQVPVLLQPHFIKAESLFLTAVKTDVSGAKTSSHTSLATTAAVQSAPLQVPTLVSGGTILATVPLMMDADKLPINRLATSGKPVLVQNKGEKRTAHNAIEKRYRSSINDRILELKDLVVGSEAKMNKSAILRKAIDYIRFLRLSNQKLKQENVALKMAAQQNKSLKDLVATCGGKTDVPMEEIKPEMMDMLTPPPSDVGSPSHSSSLSLSSSSSDSEPDSPLFEETKGKKEQQLSTGSLGMLDRSRMALCTFVFLCLSFNPLASLLRGTSPEGTPESGASPGPGRSIMGQPDSIDEASGWPTLIFWLLNVLVVLGAFVRLFIYGEPVTRPHSEPSVLFWRHRKQADLDLARGDFAQASQHLWTALKALGRPLPTSNFDLTCSLMWNLIRHLLQRLWVGRWLAGRAGGFRRDSELKADVRKSARDAALVYHKLHQLHMTGKHVGGHLSGINMALSAVNLAECAGDTVSVATLAEIYVAAALRVKTSLHRRVHFLARPFLCSARQVSLSHSGTVPPGMQWLCQPVGHRFFVDGDWSLKGTPRDSIYSSTGNPVDPLAQVTQLFREHLLEKALCCVAMPEPSLPVAHGEGEFSDALEYLQLLNGCSDTAGTPSCTLSISAGMVAGTGSDPVAKWWASIITVAIHWLQGDDEAAERLYPLVETMPRALQVSEKPLPRAALHSFKAVRALLGKQDSSQASLGQCEKASGYLRDSLGLSLPATGTLDKAVQLLLCDLLLVTRTNLWQQQMNVSQQLSCVYQASALELRGFQQDLSSLRRLAQTFRPAMRRVFLHEATARLMARASPTRTHQLLDRSLRRRGPQSSKEASEPESHPTPREHAEALLLACCYLPPSFLSAPGQRVGMLAEAARTLEKLGDKRMLHDCQQMIIKLGSGTTVTTG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1Q6</accession>
    <name>A0A452J1Q6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="23">Glutamate receptor</fullName>
        </recommendedName>
    </protein>
    <organism evidence="26 27">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="27" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="26" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="26" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="23">Receptor for glutamate that functions as a ligand-gated ion channel in the central nervous system and plays an important role in excitatory synaptic transmission. L-glutamate acts as an excitatory neurotransmitter at many synapses in the central nervous system.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>Ca(2+)(in) = Ca(2+)(out)</text>
            <dbReference type="Rhea" id="RHEA:29671"/>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="18 23">Postsynaptic cell membrane</location>
            <topology evidence="18 23">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="23">Belongs to the glutamate-gated ion channel (TC 1.A.10.1) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1Q6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038874.1">
        <property type="protein sequence ID" value="ENSGAGP00000034332.1"/>
        <property type="gene ID" value="ENSGAGG00000024385.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0022824">
        <property type="term" value="F:transmitter-gated monoatomic ion channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007166">
        <property type="term" value="P:cell surface receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd06387">
        <property type="entry name" value="PBP1_iGluR_AMPA_GluR3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd13715">
        <property type="entry name" value="PBP2_iGluR_AMPA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.70:FF:000067">
        <property type="entry name" value="glutamate receptor 2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.190.10:FF:000001">
        <property type="entry name" value="Glutamate receptor ionotropic, kainate 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.2300:FF:000004">
        <property type="entry name" value="Glutamate receptor, ionotropic, AMPA 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.190.10:FF:000666">
        <property type="entry name" value="Glutamate receptor, ionotropic, AMPA 2a"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.70">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.2300">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.190.10">
        <property type="entry name" value="Periplasmic binding protein-like II"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001828">
        <property type="entry name" value="ANF_lig-bd_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019594">
        <property type="entry name" value="Glu/Gly-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001508">
        <property type="entry name" value="Iono_Glu_rcpt_met"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015683">
        <property type="entry name" value="Ionotropic_Glu_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001320">
        <property type="entry name" value="Iontro_rcpt_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028082">
        <property type="entry name" value="Peripla_BP_I"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18966">
        <property type="entry name" value="IONOTROPIC GLUTAMATE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01094">
        <property type="entry name" value="ANF_receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00060">
        <property type="entry name" value="Lig_chan"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10613">
        <property type="entry name" value="Lig_chan-Glu_bd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00177">
        <property type="entry name" value="NMDARECEPTOR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00918">
        <property type="entry name" value="Lig_chan-Glu_bd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00079">
        <property type="entry name" value="PBPe"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53822">
        <property type="entry name" value="Periplasmic binding protein-like I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53850">
        <property type="entry name" value="Periplasmic binding protein-like II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81324">
        <property type="entry name" value="Voltage-gated potassium channels"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 23" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11 22" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="17 23" id="KW-0407">Ion channel</keyword>
    <keyword evidence="8 23" id="KW-0406">Ion transport</keyword>
    <keyword evidence="15 23" id="KW-1071">Ligand-gated ion channel</keyword>
    <keyword evidence="16" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="9 23" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0564">Palmitate</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="14 23" id="KW-0628">Postsynaptic cell membrane</keyword>
    <keyword evidence="12 23" id="KW-0675">Receptor</keyword>
    <keyword evidence="27" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 23" id="KW-0732">Signal</keyword>
    <keyword evidence="7 23" id="KW-0770">Synapse</keyword>
    <keyword evidence="4 23" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 23" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="1 23" id="KW-0813">Transport</keyword>
    <feature type="signal peptide" evidence="23">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5027146921" description="Glutamate receptor" evidence="23">
        <location>
            <begin position="23"/>
            <end position="818"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="23">
        <location>
            <begin position="545"/>
            <end position="567"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="23">
        <location>
            <begin position="630"/>
            <end position="652"/>
        </location>
    </feature>
    <feature type="domain" description="Ionotropic glutamate receptor C-terminal" evidence="24">
        <location>
            <begin position="418"/>
            <end position="795"/>
        </location>
    </feature>
    <feature type="domain" description="Ionotropic glutamate receptor L-glutamate and glycine-binding" evidence="25">
        <location>
            <begin position="428"/>
            <end position="493"/>
        </location>
    </feature>
    <feature type="binding site" evidence="20">
        <location>
            <position position="502"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="20">
        <location>
            <position position="504"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="20">
        <location>
            <position position="509"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="20">
        <location>
            <position position="680"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="20">
        <location>
            <position position="681"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="20">
        <location>
            <position position="731"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="site" description="Interaction with the cone snail toxin Con-ikot-ikot" evidence="21">
        <location>
            <position position="477"/>
        </location>
    </feature>
    <feature type="site" description="Crucial to convey clamshell closure to channel opening" evidence="21">
        <location>
            <position position="659"/>
        </location>
    </feature>
    <feature type="site" description="Interaction with the cone snail toxin Con-ikot-ikot" evidence="21">
        <location>
            <position position="686"/>
        </location>
    </feature>
    <feature type="site" description="Interaction with the cone snail toxin Con-ikot-ikot" evidence="21">
        <location>
            <position position="778"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="22">
        <location>
            <begin position="85"/>
            <end position="334"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="22">
        <location>
            <begin position="744"/>
            <end position="799"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023139"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023257"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00034104"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00036634"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="RuleBase" id="RU367118"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="SMART" id="SM00079"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="SMART" id="SM00918"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034332.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="818" mass="92421" checksum="DF166AFF8AF20DB1" modified="2019-05-08" version="1">MGQPVPQTLLLLVAGLLGESLGGFPNTISIGGLFMRNTVQEHSAFRFAVQLYNTNQNITEKPFHLNYHVDHLDSSNSFSVTNAFCSQFSRGVYAIFGFYDQMSMNTLTSFCGALHTSFVTPSFPTDADVQFVIQMRPALKGAILSLLTHYKWEKFVYLYDTERGFSILQAIMEAAVQNNWQVTARSVGSIKDVQEFRRIIEEMDRRQEKRYLIDCEVERINTILEQVVILGKHSRGYHYMLANLGFTDIILQRVMHGGANVTGFQIVNSENPMVQQFLQRWIRLDEREFPEAKNSPLKYTSALTHDAILVIAEAFRYLRRQRVDVSRRGSAGDCLANPAVPWSQGIDIERALKMVQVQGMTGNIQFDTYGRRTNYTIDVYETKATGSRKAGYWNEYERFVPALDQQVSNDTSSVENRTIVVTTILESPYVMYKKNHEQLEGNERYEGYCVDLAYEIAKHVGIKYKLSIVGDGKYGARDPESKIWNGMVGELVYGRADIAVAPLTITLVREEVIDFSKPFMSLGISIMIKKPQKSKPGVFSFLDPLAYEIWMCIVFAYIGVSVVLFLVSRFSPYEWHLEDNNEEPRDPQNPPDPPNEFGIFNSLWFSLGAFMQQGCDISPRSLSGRIVGGVWWFFTLIIISSYTANLAAFLTVERMVSPIESAEDLAKQTEIAYGTLDSGSTKEFFRRSKIAVYEKMWSYMKSAEPSVFTKTTADGVARVRKSKGKFAFLLESTMNEYIEQRKPCDTMKVGGNLDSKGYGVATPKGSALRNAVNLAVLKLNEQGLLDKLKNKWWYDKGECGSGGGDSKNSCKPCSIETQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1Q9</accession>
    <name>A0A452J1Q9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="20">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 16">Membrane</location>
            <topology evidence="1 16">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 16">Belongs to the integrin alpha chain family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1Q9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038756.1">
        <property type="protein sequence ID" value="ENSGAGP00000034229.1"/>
        <property type="gene ID" value="ENSGAGG00000024233.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009897">
        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008305">
        <property type="term" value="C:integrin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033627">
        <property type="term" value="P:cell adhesion mediated by integrin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098609">
        <property type="term" value="P:cell-cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007160">
        <property type="term" value="P:cell-matrix adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007229">
        <property type="term" value="P:integrin-mediated signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.130:FF:000006">
        <property type="entry name" value="Integrin, alpha 9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.930">
        <property type="entry name" value="Bicelle-embedded integrin alpha(iib) transmembrane segment"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.130">
        <property type="entry name" value="Integrin alpha, N-terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1460">
        <property type="entry name" value="Integrin domains. Chain A, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1530">
        <property type="entry name" value="ntegrin, alpha v. Chain A, domain 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013517">
        <property type="entry name" value="FG-GAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013519">
        <property type="entry name" value="Int_alpha_beta-p"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000413">
        <property type="entry name" value="Integrin_alpha"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013649">
        <property type="entry name" value="Integrin_alpha_Ig-like_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048286">
        <property type="entry name" value="Integrin_alpha_Ig-like_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028994">
        <property type="entry name" value="Integrin_alpha_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032695">
        <property type="entry name" value="Integrin_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23220">
        <property type="entry name" value="INTEGRIN ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23220:SF78">
        <property type="entry name" value="INTEGRIN ALPHA-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01839">
        <property type="entry name" value="FG-GAP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08441">
        <property type="entry name" value="Integrin_A_Ig_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20806">
        <property type="entry name" value="Integrin_A_Ig_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01185">
        <property type="entry name" value="INTEGRINA"/>
    </dbReference>
    <dbReference type="SMART" id="SM00191">
        <property type="entry name" value="Int_alpha"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69318">
        <property type="entry name" value="Integrin alpha N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69179">
        <property type="entry name" value="Integrin domains"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51470">
        <property type="entry name" value="FG_GAP"/>
        <property type="match status" value="5"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0106">Calcium</keyword>
    <keyword evidence="8 16" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="14" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10 16" id="KW-0401">Integrin</keyword>
    <keyword evidence="11 16" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13 16" id="KW-0675">Receptor</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5 17" id="KW-0732">Signal</keyword>
    <keyword evidence="3 16" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9 16" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="17">
        <location>
            <begin position="1"/>
            <end position="37"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019369790" evidence="17">
        <location>
            <begin position="38"/>
            <end position="933"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="870"/>
            <end position="892"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="43"/>
            <end position="104"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="190"/>
            <end position="241"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="296"/>
            <end position="355"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="359"/>
            <end position="416"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="420"/>
            <end position="482"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha first immunoglubulin-like" evidence="18">
        <location>
            <begin position="467"/>
            <end position="572"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha third immunoglobulin-like" evidence="19">
        <location>
            <begin position="693"/>
            <end position="843"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023037"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00803"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="RuleBase" id="RU003762"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF08441"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="Pfam" id="PF20806"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034229.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="933" mass="103463" checksum="A4B5355081C356BB" modified="2019-05-08" version="1">MRSRARLPGGLASGGAPVWAALLLLLLLGRCMRAAHAYNLDTQRPVVFKGLSGTLFGYSVVLHSHGPSRWLVVGAPKANWLANSSVVNPGAIFRCRIGSNPSGTCEQLQMGNPAGDYCGKTCFEERDNQWLGVSLSRQPRENGTIVACGHRWKNIYYIKNEHKLPNGVCYEIPPDFRTQLSKRISPCYKDYVRKFGENHGSCQAGMSSFYFKDLIIMGAPGSYYWTGSVFVYNTTANTFQAYTDSRNQVKFGSYLGYAVGAGHFLTPASTEVIGGAPQQEQTGKAYIFSIGTRALTILSELKGKKLGSYFGAAVCAVDLNTDGLSDLLVGAPMQSKVREEGRVYVYINSGSGAKMLELEIELAGSDSYAARFGESIANLGDIDNDGFEDVAIGAPQENNLEGAIYIYNGRKDGISSSFSQRIQGHRISNTLSMFGQSISSGIDADNNGYPDVAVGAFLDDSAVLLRTRPVVIVEAFLKHPNAVNRTNLECVENGQPAVCVNLMLCFTYKGRDVPGYVVLLYDLSVDVSRKTNTPARFYFSSNGTSDMVSGSMEIYNSNVTCKSHLAFLKENIDRWKTLPHEKKTYLSVGSMKTLLLNISLSNAGDDAYETALHIQLPKGLYFVRVLELVRIEIGCCHNSLTILTLLGQTVHVSFVLQMDFSFLLDASSLTRAEDDLTITVNATCEENSPINCMKEKVNFTFHVISAGPSMAPAIGLEIMVPNSFAPRDEKLFNILDIQTADGECFYNNYTRDCTLPEKNGHMNILEDLVIFFSKPTKRRLYCMKEDHLCLRILCKFGNMENGKEAIVRMQLEATPSVLEMDEASSLKFEISATASPEKNPKVIELHKERQIAHVFLEGLHNQKPKHHVTMLIIGTGSVLGIFLFLLIVLLLWKVGFFFLYQYFSIIFIDSFKNRNNMLYCDLFLLKKNKSWYD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1R1</accession>
    <name>A0A452J1R1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Eyes absent homolog</fullName>
            <ecNumber evidence="15">3.1.3.48</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="12 15">
            <text>O-phospho-L-tyrosyl-[protein] + H2O = L-tyrosyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:10684"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="EC" id="3.1.3.48"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="14 15">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
        <text evidence="14 15">Binds 1 Mg(2+) ion per subunit.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 15">Belongs to the HAD-like hydrolase superfamily. EYA family.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.48" evidence="15"/>
    <dbReference type="AlphaFoldDB" id="A0A452J1R1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038868.1">
        <property type="protein sequence ID" value="ENSGAGP00000034326.1"/>
        <property type="gene ID" value="ENSGAGG00000024390.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:2001240">
        <property type="term" value="P:negative regulation of extrinsic apoptotic signaling pathway in absence of ligand"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045739">
        <property type="term" value="P:positive regulation of DNA repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd02601">
        <property type="entry name" value="HAD_Eya"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.12350">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028472">
        <property type="entry name" value="EYA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006545">
        <property type="entry name" value="EYA_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042577">
        <property type="entry name" value="EYA_dom_metazoan"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038102">
        <property type="entry name" value="EYA_dom_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01658">
        <property type="entry name" value="EYA-cons_domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10190">
        <property type="entry name" value="EYES ABSENT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10190:SF7">
        <property type="entry name" value="EYES ABSENT HOMOLOG 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00702">
        <property type="entry name" value="Hydrolase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0010">Activator</keyword>
    <keyword evidence="3" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="5 15" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="6 14 15" id="KW-0460">Magnesium</keyword>
    <keyword evidence="4 14 15" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="7 15" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0804">Transcription</keyword>
    <keyword evidence="8 15" id="KW-0805">Transcription regulation</keyword>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="208"/>
            <end position="259"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="208"/>
            <end position="220"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="16">
        <location>
            <begin position="230"/>
            <end position="245"/>
        </location>
    </feature>
    <feature type="active site" description="Nucleophile" evidence="13">
        <location>
            <position position="270"/>
        </location>
    </feature>
    <feature type="active site" description="Proton donor" evidence="13">
        <location>
            <position position="272"/>
        </location>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="270"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="272"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010501"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00051722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PIRSR" id="PIRSR628472-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PIRSR" id="PIRSR628472-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="RuleBase" id="RU362036"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034326.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="486" mass="53470" checksum="24110AB2C80D7567" modified="2019-05-08" version="1">MLDLVISPSLTVNRECPDRLKLNLSNIYATAPDDIEGNSKAASQCPLHLCSTEHHPHIVTVPSFPTMATYGQTQYSAGIQQAATYAAYPPPGQPYGIPSYSIKTEDSLSHSPGQSGFLSYGSTYSTPTTGQAPYTYQMHGTSGIYQGANGLTNSAGFSTVHQEYSSYPSFPQSQYSQYYTSSYNSPYVSANSISPSAIPTSTYSLQESSHNITSHSTESLSGEYATTPVKDTETDRQHRGSDGKLRGRSKRSNDPSPTVDNEIERVFVWDLDETIIIFHSLLTGTFASRYGKDTTTSVRIGLMMEEMIFNLADTHLFFNDLEDCDQIHIDDVSSDDNGQDLSTYNFSADGFHSSTAGANLCLGSGVHGGVDWMRKLAFRYRRVKEMYNTYKNNVGGLIGAPKRETWLQLRAELEALTDLWLTHALKALNLIHSRPNCVNVLVTTTQLIPALAKVLLYGLGTVFPIENIYSATKTAQHAFLENLLSR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1R3</accession>
    <name>A0A452J1R3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">CAP-Gly domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1R3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038870.1">
        <property type="protein sequence ID" value="ENSGAGP00000034328.1"/>
        <property type="gene ID" value="ENSGAGG00000024389.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005938">
        <property type="term" value="C:cell cortex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035371">
        <property type="term" value="C:microtubule plus-end"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051010">
        <property type="term" value="F:microtubule plus-end binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031122">
        <property type="term" value="P:cytoplasmic microtubule organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.190:FF:000002">
        <property type="entry name" value="CAP-Gly domain containing linker protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.190:FF:000001">
        <property type="entry name" value="Putative CAP-Gly domain-containing linker protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.190">
        <property type="entry name" value="CAP Gly-rich-like domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036859">
        <property type="entry name" value="CAP-Gly_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000938">
        <property type="entry name" value="CAP-Gly_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18916:SF10">
        <property type="entry name" value="CAP-GLY DOMAIN-CONTAINING LINKER PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18916">
        <property type="entry name" value="DYNACTIN 1-RELATED MICROTUBULE-BINDING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01302">
        <property type="entry name" value="CAP_GLY"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM01052">
        <property type="entry name" value="CAP_GLY"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF74924">
        <property type="entry name" value="Cap-Gly domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00845">
        <property type="entry name" value="CAP_GLY_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50245">
        <property type="entry name" value="CAP_GLY_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5 7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="3" id="KW-0493">Microtubule</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="CAP-Gly" evidence="9">
        <location>
            <begin position="90"/>
            <end position="132"/>
        </location>
    </feature>
    <feature type="domain" description="CAP-Gly" evidence="9">
        <location>
            <begin position="230"/>
            <end position="272"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1"/>
            <end position="59"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="135"/>
            <end position="168"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="186"/>
            <end position="206"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="284"/>
            <end position="330"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="975"/>
            <end position="999"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="345"/>
            <end position="446"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="505"/>
            <end position="571"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="638"/>
            <end position="853"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="895"/>
            <end position="922"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="23"/>
            <end position="35"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="153"/>
            <end position="168"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="186"/>
            <end position="201"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="305"/>
            <end position="329"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="975"/>
            <end position="993"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034328.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1026" mass="114565" checksum="217F5E765FB7C8B8" modified="2019-05-08" version="1">MLKPSGLKIPGRAGKHSSPVGRASGTAAAAATTGSKEGSPLHKQGPTSTASAEKPGSKVTEVGDDFLGDFVVGERVWVNGVKPGVIQYLGETQFAPGQWAGVVLDDPVGKNDGSVGGVRYFECQPLQGIFTRPSKLTRQPVAEGSGSDAHSVESLTTQNLSLHSGTATPPLSTRIIPLRESVLNSTMKTGNESGSNLSDSGSVKKGDKDLRLGDRVLVGGTKTGVVRYVGETDFAKGEWCGVELDEPLGKNDGAVAGTRYFQCPPKFGLFAPIHKVIRIGFPSTSPAKAKKSKRMAMGVSALTHSPSSSSISSVSSVASSVGGRPSRSGLLTETSSRYARKISGTTALQEALKEKQQHIEQLLAERDLERAEVAKATSHICEVEKEIAILKAQHEQYVTEAEANLQRARALVDGVQKEKLELLNQLEEEKRKVEDLQFRVEEESITKGDLELTTVAEKSRILQLEEELSVRRGEVEELRQCLRNSHQAETPEHNLGLQSEALRLRDQLLSASKEHQKESSQLKEKYEKTLKKHQQEIEKLKAVNEKYSQEIVDLKHKVQQATNENMGLMDNWKSKLDTLASDHQKSLEDLKATLNTGPDTQHKEIVELKAVVESIKMEHQLELENLKAKHDIETAVHIKEKESLKLKLQEASDEVEESNNNWKMQLETKTNQHLIELQDLKEKCRDAELRVHELEKLHSEYKDQAQAIAFLKEQISLAEKKMLDYETLQKTEAQSKQEIQRLQEKMLILENKLQSMEALHPSQHANMIETNDISEEKIKMKEAMEELQDKLSKRDKEVSSLLSQTETLRAQVSALENKCKAGEKKGDSLLKEKKRLEAELETLSRKTHDASGQLVLISQELLKKERSLNELRALLLEANRHSPGPERDLSREVHKAEWRLKEQKLKDDIKGLREKLIVLEKEKSVVDQRRYSLIDPSSESEVLRLQHRLVSTEDVLRNALEQGQQMEKLMEAMRSSSEKTQTLGNSGSANGIHQQDKSHKQEVGYTMLSYLNKKTPMQNFFRGVSL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1U5</accession>
    <name>A0A452J1U5_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="4">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000038896.1">
        <property type="protein sequence ID" value="ENSGAGP00000034354.1"/>
        <property type="gene ID" value="ENSGAGG00000024398.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043379">
        <property type="entry name" value="ANKAR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000225">
        <property type="entry name" value="Armadillo"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46464">
        <property type="entry name" value="ANK_REP_REGION DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46464:SF1">
        <property type="entry name" value="ANKYRIN AND ARMADILLO REPEAT-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13637">
        <property type="entry name" value="Ank_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00185">
        <property type="entry name" value="ARM"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50176">
        <property type="entry name" value="ARM_REPEAT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="repeat" description="ANK" evidence="1">
        <location>
            <begin position="550"/>
            <end position="582"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="1">
        <location>
            <begin position="619"/>
            <end position="651"/>
        </location>
    </feature>
    <feature type="repeat" description="ARM" evidence="2">
        <location>
            <begin position="723"/>
            <end position="765"/>
        </location>
    </feature>
    <feature type="repeat" description="ARM" evidence="2">
        <location>
            <begin position="764"/>
            <end position="806"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="1"/>
            <end position="24"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="49"/>
            <end position="68"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="514"/>
            <end position="546"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="524"/>
            <end position="545"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00259"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034354.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1343" mass="150288" checksum="FEA651FEE33612AE" modified="2019-05-08" version="1">MTSASCTAPPAGVAKGKRRRPALLRFPPGQVPRASAGWAAGLPPFASTTGQSGAWPPMETPTSPPACLDSNYDHSTSCQLPPAYIDTRIGQILISVDYMLKALWHGVYMPKEKRVRFSELWRSSMDIDMDGNPQTEKNIFSEFCSAGLVDLSNEPDFEGIYDENMNEDPTYDPNSAQEKNLFMQYADNILFKLTFGTIQVQQHESVCKFDAAYVLSNVIRLTEDHLDTVTYQRLQQRLILQQKLVKKHLEKKAEIRKNIEYLKIVSFLIPFLLGLKRKMKIPNLNQLLQPYSDDKVKTERELPPFVFGPDFKCQHFHYIKNEYFHVHGGIEFDVGTPSLEDVSEKIKAVFDDIQICASNHVAQLLDPDIPYREHYSIPTMEFDGKSYYVISIELETFYQQLYKTPWWGAINEIISTLKPKRLPLTDIQSLEQFKKRFGYKKAIKCKSLPYGMKSAAERGLSAIFHTFCRKTSTSSLSVVDEYGYALLHHASIHNRVPIICQLIKAHLNINQRRNVHFNPGRRSQKSETKGDRAGKPDLQEKETSGERTALGPTALHLAAQCGSIEALTCLLAFKADYKLTDDRGWMAIHFAAFYDNISCITVLYRKDPDLLEAETTAEYHSTPLLLAATSGALDTLQYLFSLGANWKKTDSEGNNIIHLAVLYFHTEILKHLIQLDNPDLPVWNTLVEMLQCEDSMRPEMAVRSLEVLCLAKDFYWKCILDSGAIPSLINLLKGHQIKLACLTSGVLSNISPHITISKALVEAGGISVLIELLGSGEPELQSRCAVILYDIAQFDNNQNVISELGGIPPLINLLKIRLQDLLVNVINCIRVLCIQNKQNQITVKEHQGIPALVEFLTSQSDVLQAVSSAALAELARGNEKMQDAIADADAIGPLVELLRGRKITVQVKGAMAIEALSDNNTHIQRLFLEKSATKYLLKLLKAFHLTVKEQGATTLWALAGQTLKQQKLMAEQIGYNFIIDMLLSPSDKMQYVGGEAVIALSKDSKLHQNQICEGNGIGPLVRLLRSTKIAEGTLLSVIRALGTICIGVAHTNNPVSQENIVDEQALPILVHLLKNNNSLQIKVEVAYSLACIVLRNSNLQTVLQEEEGFNYKDVLELLNTPDKDICLRAGYALALFAYNNPVQQFLILETGAIMMSTFEPFLRSEIETHKAMAAFQIVVLARVIVDVDQVTLSARGVTILVELLNSEKTATLVLTGKLLASLAHTRAGVPEAITELGTIKRLCYHLYSDKEEVRIASAGALGYLTFNRTAYRHLLVECRNKPKQFTRLMNNLSRDAKISQAFVKEFQRQRQVGLPSLRLTFSSDVRHLSATTKSSELESDMVW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J203</accession>
    <name>A0A452J203_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="15">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">May be involved in transcriptional regulation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J203"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038971.1">
        <property type="protein sequence ID" value="ENSGAGP00000034421.1"/>
        <property type="gene ID" value="ENSGAGG00000024460.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031981">
        <property type="term" value="C:nuclear lumen"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003700">
        <property type="term" value="F:DNA-binding transcription factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07765">
        <property type="entry name" value="KRAB_A-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:003000">
        <property type="entry name" value="Zinc finger and SCAN domain-containing 20"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000352">
        <property type="entry name" value="zinc finger protein 3 homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002343">
        <property type="entry name" value="Zinc finger protein 33A"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000016">
        <property type="entry name" value="zinc finger protein 37 homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002090">
        <property type="entry name" value="Zinc finger protein 473"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000990">
        <property type="entry name" value="zinc finger protein 629 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002190">
        <property type="entry name" value="Zinc finger protein 658"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000176">
        <property type="entry name" value="zinc finger protein 70"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050589">
        <property type="entry name" value="Ikaros_C2H2-ZF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001909">
        <property type="entry name" value="KRAB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036051">
        <property type="entry name" value="KRAB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24404">
        <property type="entry name" value="ZINC FINGER PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24404:SF100">
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        <source>
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    <sequence length="410" mass="46998" checksum="A9B876FBCA4B804F" modified="2019-05-08" version="1">LSPSPTPQSLLAVLSGGLMLFQMPVTFEEVAVYFTQGQGALLDPAQRALYRDVMQENYKMVTSLGKGFSSPWLLEAMGSVCLNLETTAPQTYHQEEKPYKRLENRKRFSVSRNLSSHLITHQRLHTGERPYKCLECGKGFNVRSNLIKHQRNHTGGKPHICLDCGKCFNHSSYLITHQRLHTGERPYKCLECGKSFNQSSSLITHLRTHTGNKPYQCLNCGKSFNVRSQLIIHQRIHTGERPHKCLDCGKSFSHSSHLTKHLRIHTGERPYKCLECGKTFNQSSHLIMHQRTHTGERHYKCLECGKSFVRRSALITHQRIHTGEKPHICLDCGKSFSQRSALIRHQRIHTGEKPHTCLNCGKRFIQRSHLITHQTIHKGEKTHKCLDCGKSFNNSSYLTKHRRIHKGETL</sequence>
</entry>
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    <name>A0A452J205_9SAUR</name>
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        <recommendedName>
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            <taxon>Archelosauria</taxon>
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            <taxon>Durocryptodira</taxon>
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                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
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        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
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        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
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            </authorList>
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        <scope>IDENTIFICATION</scope>
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        <citation type="submission" date="2025-09" db="UniProtKB">
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                <consortium name="Ensembl"/>
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        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
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        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
    <sequence length="400" mass="44816" checksum="90B49881E4B3685C" modified="2019-05-08" version="1">MVAPQPLRVVRCGGSKLSGARAAFSADAKYLLCASGDYVKVYSTATEKCVHILQGHTNLVTGIQLNPQNHMQLYSCSLDGTIKLWDFTDGILIKTFVVGYKLFAFYMLAGSEDSVFIIIPKNDEDFLDWFQLVSVKLPKTPDQEVEAKELSLILDDIGQWPKCTAFGREGEYVASVRGLHLLVYFFKKKKINRFSLSATSKKGANNYFTCVACHPKEDCIASGHKDGKIRLWRNFNHKREYTFSTLHWHHDVVMDLAFSVEGTSLLSGGVESVLVQWRDGSDYKKEFLPRLGSTIEHISASPDGTFYCTSHTENKIIIIHSSLSVSAVIQGLVKGSDVKTGLVVDPRTKALVLNGKPGHLQFYCLQTDKQLYNVSWIDWCCKLYTCVTELHLQYDVGKGE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J218</accession>
    <name>A0A452J218_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">WD repeat-containing protein 75 second beta-propeller domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
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            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
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        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
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                <person name="Dolby G.A."/>
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                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
            <location evidence="1">Nucleolus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J218"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038992.1">
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        <property type="gene ID" value="ENSGAGG00000024445.1"/>
    </dbReference>
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    </dbReference>
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        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
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    </dbReference>
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    </dbReference>
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        <property type="entry name" value="WD40_repeat_CS"/>
    </dbReference>
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        <property type="entry name" value="WD40_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR053826">
        <property type="entry name" value="WDR75"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR44215">
        <property type="entry name" value="WD REPEAT-CONTAINING PROTEIN 75"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR44215:SF1">
        <property type="entry name" value="WD REPEAT-CONTAINING PROTEIN 75"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23869">
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23769">
        <property type="entry name" value="Beta-prop_WDR75_2nd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00320">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50998">
        <property type="entry name" value="Quinoprotein alcohol dehydrogenase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50969">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00678">
        <property type="entry name" value="WD_REPEATS_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50082">
        <property type="entry name" value="WD_REPEATS_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50294">
        <property type="entry name" value="WD_REPEATS_REGION"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0690">Ribosome biogenesis</keyword>
    <keyword evidence="3" id="KW-0698">rRNA processing</keyword>
    <keyword evidence="6" id="KW-0804">Transcription</keyword>
    <keyword evidence="4 8" id="KW-0853">WD repeat</keyword>
    <feature type="repeat" description="WD" evidence="8">
        <location>
            <begin position="53"/>
            <end position="95"/>
        </location>
    </feature>
    <feature type="domain" description="WD repeat-containing protein 75 second beta-propeller" evidence="9">
        <location>
            <begin position="311"/>
            <end position="636"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004604"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022517"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022552"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022574"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF23769"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034438.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="804" mass="90594" checksum="38D5D04DFFFC42E2" modified="2019-05-08" version="1">MVAPQPLRVVRCGGSKLSGARAAFSADAKYLLCASGDYVKVYSTATEKCVHILQGHTNLVTGIQLNPQNHMQLYSCSLDGTIKLWDFTDGILIKVCKYFVLSQLFMDSRVVHYWFQLVSVKLPKTPDQEVEAKELSLILDDIGQWPKCTMFFFYIKCINHILFSLSATSKKGANNYFTCVACHPKEDCIASGHKDGKIRLWRNFNHKREYTFSTLHWHHDVVMDLAFSVEGTSLLSGGVESVLVQWRDGSDYKKEFLPRLGSTIEHISASPDGTFYCTSHTENKIIIIHSSLSVSAVIQGLVKGSDVKTGLVVDPRTKALVLNGKPGHLQFYCLQTDKQLYNLDIVQQEYIHQIGLNQVDLVKAAFSAQGNWLATVEQREEKEVELELQMKLWNYDEQTQSFTLNTRINMPHEDHVTALCFRDTDELEDGTPTLVTAGNDGRFKVWVLVNDSDSEKPSMGWSCDFVGSYHNYPATNCCFSEDGSLLAVSFEETVTVWNSDTWDLICTFCHPPGRIQNLCFGRLSCSKYLLGTTDCGFLCCWNLLSCALEWSAQLNVMVLQPDPLSENIAAVSRLLEGSDLFVFKPNEPRPFCIQRNVCRERVQCAVFIPRDVPDSIGSEKHQWLSRSQLYFLTETQDLLTFSTKSAEERLTPSSRQLAVEESLPMTPFSLLLGKHRHQQQSKENADLDTAVVQSHLAQGSPAVKELLHTPAHVLPSASFLCSIFINSVLISKENKSAGEVVDVVEMDSEKAEDESDEDLELTEMEQDINPTEFLGEITEIKLSKSQEKQLRKIRRMDYSWVAAL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J223</accession>
    <name>A0A452J223_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Major facilitator superfamily (MFS) profile domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the major facilitator superfamily. Sugar transporter (TC 2.A.1.1) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J223"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039015.1">
        <property type="protein sequence ID" value="ENSGAGP00000034459.1"/>
        <property type="gene ID" value="ENSGAGG00000024490.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005903">
        <property type="term" value="C:brush border"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055056">
        <property type="term" value="F:D-glucose transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046323">
        <property type="term" value="P:D-glucose import"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070837">
        <property type="term" value="P:dehydroascorbic acid transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd17431">
        <property type="entry name" value="MFS_GLUT_Class1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1250.20">
        <property type="entry name" value="MFS general substrate transporter like domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045263">
        <property type="entry name" value="GLUT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020846">
        <property type="entry name" value="MFS_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005828">
        <property type="entry name" value="MFS_sugar_transport-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036259">
        <property type="entry name" value="MFS_trans_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003663">
        <property type="entry name" value="Sugar/inositol_transpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005829">
        <property type="entry name" value="Sugar_transporter_CS"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00879">
        <property type="entry name" value="SP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23503">
        <property type="entry name" value="SOLUTE CARRIER FAMILY 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23503:SF27">
        <property type="entry name" value="SOLUTE CARRIER FAMILY 2, FACILITATED GLUCOSE TRANSPORTER MEMBER 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00083">
        <property type="entry name" value="Sugar_tr"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00171">
        <property type="entry name" value="SUGRTRNSPORT"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103473">
        <property type="entry name" value="MFS general substrate transporter"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50850">
        <property type="entry name" value="MFS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00217">
        <property type="entry name" value="SUGAR_TRANSPORT_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 6" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 6" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="3 6" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="5" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="50"/>
            <end position="72"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="79"/>
            <end position="99"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="105"/>
            <end position="130"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="142"/>
            <end position="160"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="172"/>
            <end position="193"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="258"/>
            <end position="281"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="293"/>
            <end position="314"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="321"/>
            <end position="343"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="349"/>
            <end position="367"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="388"/>
            <end position="412"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="418"/>
            <end position="436"/>
        </location>
    </feature>
    <feature type="domain" description="Major facilitator superfamily (MFS) profile" evidence="7">
        <location>
            <begin position="12"/>
            <end position="442"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="RuleBase" id="RU003346"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50850"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034459.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="479" mass="52382" checksum="F27273711FDBB109" modified="2025-10-08" version="2">MEGSSKRTLVLSVFTAVLGFFQYGYSLGVINAPQRAISVYRSGKSTLTLYWSLSVSIFAIGGMISSFTVGWVGDKLGRVKAMLAVNVLSIIGNLLMGLAKFGPSHILIIAGRAITGLYCGLSSGLVPMYVGEISPTALRGALGTLHQLAIVTGILLSQVLGLDFLLGNDDMWPLLLGLSGAAALLQFFLLLLCPESPRYLYIKLGKVEEAKKSLKRLRGDCDPTKEITEMEKEKQEAASEKKVSIRKLFSSSSYRQPIIVALMVQISQQFSGINAIFYYSTNIFVRAGVDQPVYATIGVGVVNTIFTIISVFLVERTGRRSLFIAGLIGMLASAVAMTVGLVLLGTFAWMSYVSMVAIFLFVIFFEVGPGPIPWFIVAELFSQGPRPAAIAIAGFCNWTCNFIIGMCFQYIADLCGPYVFTIFAVLLLGFILFAHFKVPETKGKSFEEISAAFRQKRRCAIIYSYGWFHPEKTKCYCSC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J228</accession>
    <name>A0A452J228_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Major facilitator superfamily (MFS) profile domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J228"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039020.1">
        <property type="protein sequence ID" value="ENSGAGP00000034464.1"/>
        <property type="gene ID" value="ENSGAGG00000024490.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005903">
        <property type="term" value="C:brush border"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055056">
        <property type="term" value="F:D-glucose transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046323">
        <property type="term" value="P:D-glucose import"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070837">
        <property type="term" value="P:dehydroascorbic acid transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1250.20">
        <property type="entry name" value="MFS general substrate transporter like domains"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045263">
        <property type="entry name" value="GLUT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020846">
        <property type="entry name" value="MFS_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005828">
        <property type="entry name" value="MFS_sugar_transport-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036259">
        <property type="entry name" value="MFS_trans_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003663">
        <property type="entry name" value="Sugar/inositol_transpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005829">
        <property type="entry name" value="Sugar_transporter_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23503">
        <property type="entry name" value="SOLUTE CARRIER FAMILY 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23503:SF27">
        <property type="entry name" value="SOLUTE CARRIER FAMILY 2, FACILITATED GLUCOSE TRANSPORTER MEMBER 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00083">
        <property type="entry name" value="Sugar_tr"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00171">
        <property type="entry name" value="SUGRTRNSPORT"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103473">
        <property type="entry name" value="MFS general substrate transporter"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50850">
        <property type="entry name" value="MFS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00217">
        <property type="entry name" value="SUGAR_TRANSPORT_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4 5" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 5" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="3 5" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="12"/>
            <end position="35"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="72"/>
            <end position="92"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="104"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="128"/>
            <end position="153"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="165"/>
            <end position="183"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="195"/>
            <end position="216"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="309"/>
            <end position="336"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="342"/>
            <end position="364"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="376"/>
            <end position="399"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="405"/>
            <end position="423"/>
        </location>
    </feature>
    <feature type="domain" description="Major facilitator superfamily (MFS) profile" evidence="6">
        <location>
            <begin position="22"/>
            <end position="429"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="PROSITE" id="PS50850"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034464.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="466" mass="50405" checksum="A15C16624A704FD6" modified="2019-05-08" version="1">IPRGFHLSTQSIQHLTGTLVLSVFTAVLGFFQYGYSLGVINAPQRATGLHSPSLVRGALGLVCPQGPSTLTLYWSLSVSIFAIGGMISSFTVGWVGDKLGRVKAMLAVNVLSIIGNLLMGLAKFGPSHILIIAGRAITGLYCGLSSGLVPMYVGEISPTALRGALGTLHQLAIVTGILLSQVLGLDFLLGNDDMWPLLLGLSGAAALLQFFLLLLCPESPRYLYIKLGKVEEAKKSLKRLRGDCDPTKEITEMEKEKQEAASEKKVSIRKLFSSSSYRQPIIVALMVQISQQFSGINAVILYITGRRSLFIAGLIGMLASAVAMTVGLVLLGTFAWMSYVSMVAIFLFVIFFEVGPGPIPWFIVAELFSQGPRPAAIAIAGFCNWTCNFIIGMCFQYIADLCGPYVFTIFAVLLLGFILFAHFKVPETKGKSFEEISAAFRRKRRSAIKGHKAATELEDLRGSEEA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J229</accession>
    <name>A0A452J229_9SAUR</name>
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        <recommendedName>
            <fullName evidence="5">Tumor necrosis factor alpha-induced protein 8</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the TNFAIP8 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J229"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039000.1">
        <property type="protein sequence ID" value="ENSGAGP00000034445.1"/>
        <property type="gene ID" value="ENSGAGG00000024487.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043027">
        <property type="term" value="F:cysteine-type endopeptidase inhibitor activity involved in apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006915">
        <property type="term" value="P:apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
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        <property type="term" value="P:regulation of apoptotic process"/>
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        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1440.160:FF:000001">
        <property type="entry name" value="Tumor necrosis factor alpha-induced protein 8-like 1"/>
        <property type="match status" value="1"/>
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    <dbReference type="Gene3D" id="1.20.1440.160">
        <property type="entry name" value="Tumor necrosis factor alpha-induced protein 8-like"/>
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        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
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    </dbReference>
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    <keyword evidence="3" id="KW-0053">Apoptosis</keyword>
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    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022703"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00038267"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00040051"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034445.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="188" mass="21802" checksum="D1A7E3AFF2522A87" modified="2019-05-08" version="1">MATDIFNSKNLAIQAQKKILGKMVSKSIATTLIDDTSSDVLDELYRVTREYTQNKKEAEKIIKNLIKTVIKLAILYRNNQFNQEEIALMEKFKKKVHQLAKTVVSFHQVDYTFDRIFLSKLLNDCREILHQIIQRHLTAKSHGRVNNVFDHFSDCEFLAALYNPFGPYKPHLQKLCDGVNKMLDEGNI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J232</accession>
    <name>A0A452J232_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Malic enzyme</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="10">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
        <cofactor evidence="10">
            <name>Mn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29035"/>
        </cofactor>
        <text evidence="10">Divalent metal cations. Prefers magnesium or manganese.</text>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29035"/>
        </cofactor>
    </comment>
    <comment type="subunit">
        <text evidence="4">Homotetramer.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 11">Belongs to the malic enzymes family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J232"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039011.1">
        <property type="protein sequence ID" value="ENSGAGP00000034455.1"/>
        <property type="gene ID" value="ENSGAGG00000024492.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004473">
        <property type="term" value="F:malate dehydrogenase (decarboxylating) (NADP+) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051287">
        <property type="term" value="F:NAD binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006108">
        <property type="term" value="P:malate metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05312">
        <property type="entry name" value="NAD_bind_1_malic_enz"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.720:FF:000060">
        <property type="entry name" value="Malic enzyme"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10380:FF:000028">
        <property type="entry name" value="NADP-dependent malic enzyme 4 chloroplastic"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.10380">
        <property type="entry name" value="Malic enzyme, N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.720">
        <property type="entry name" value="NAD(P)-binding Rossmann-like Domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046346">
        <property type="entry name" value="Aminoacid_DH-like_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015884">
        <property type="entry name" value="Malic_enzyme_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012301">
        <property type="entry name" value="Malic_N_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037062">
        <property type="entry name" value="Malic_N_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012302">
        <property type="entry name" value="Malic_NAD-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001891">
        <property type="entry name" value="Malic_OxRdtase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036291">
        <property type="entry name" value="NAD(P)-bd_dom_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF010052">
        <property type="entry name" value="PRK13529.1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23406">
        <property type="entry name" value="MALIC ENZYME-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23406:SF17">
        <property type="entry name" value="NADP-DEPENDENT MALIC ENZYME"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00390">
        <property type="entry name" value="malic"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03949">
        <property type="entry name" value="Malic_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000106">
        <property type="entry name" value="ME"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00072">
        <property type="entry name" value="MALOXRDTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM01274">
        <property type="entry name" value="malic"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00919">
        <property type="entry name" value="Malic_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53223">
        <property type="entry name" value="Aminoacid dehydrogenase-like, N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51735">
        <property type="entry name" value="NAD(P)-binding Rossmann-fold domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00331">
        <property type="entry name" value="MALIC_ENZYMES"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6 10 11" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7" id="KW-0521">NADP</keyword>
    <keyword evidence="8 11" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Malic enzyme N-terminal" evidence="13">
        <location>
            <begin position="2"/>
            <end position="156"/>
        </location>
    </feature>
    <feature type="domain" description="Malic enzyme NAD-binding" evidence="12">
        <location>
            <begin position="166"/>
            <end position="418"/>
        </location>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="51"/>
        </location>
        <ligand>
            <name>(S)-malate</name>
            <dbReference type="ChEBI" id="CHEBI:15589"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="10">
        <location>
            <position position="141"/>
        </location>
        <ligand>
            <name>a divalent metal cation</name>
            <dbReference type="ChEBI" id="CHEBI:60240"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="10">
        <location>
            <position position="142"/>
        </location>
        <ligand>
            <name>a divalent metal cation</name>
            <dbReference type="ChEBI" id="CHEBI:60240"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="10">
        <location>
            <position position="165"/>
        </location>
        <ligand>
            <name>a divalent metal cation</name>
            <dbReference type="ChEBI" id="CHEBI:60240"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="304"/>
        </location>
        <ligand>
            <name>(S)-malate</name>
            <dbReference type="ChEBI" id="CHEBI:15589"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="349"/>
        </location>
        <ligand>
            <name>(S)-malate</name>
            <dbReference type="ChEBI" id="CHEBI:15589"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001936"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00008785"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00011881"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022857"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PIRSR" id="PIRSR000106-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PIRSR" id="PIRSR000106-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="RuleBase" id="RU003426"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="SMART" id="SM00919"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="SMART" id="SM01274"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034455.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="467" mass="51782" checksum="AC337E55AC0BD284" modified="2019-05-08" version="1">SLFFQTKMQSIYLFACRGLFITIHDWGHIATMLKSWPEDAIRAIVVTDGERILGLGDLGCYGMGIPVGKLALYTACGGMKPQECLPVILDVGTDNEELLKDPLYIGLRHKRVRGQAYNDLLDEFMQAVTFRYGMNCLIQFEDFANANAFRLLNRYRNKYCTFNDDIQGTAAVAVAGLLAALRITENKLSDHTILFQGAGEAALGIANLVIMAMQKEGTPEHEAIKKIWMVDSKGLIVKERSSLTTEKERFAHKHDEMKNLEDIVRAIKPTVLIGVAAIGGAFTKQILKDMASFNKQPIIFALSNPTSKAECSAEQCYQMTEGRGIFASGSPFDPVTLPGGRTFYPGQGNNAYVFPGVALGVVSCGLRYISEDIFLTTAEVIAQQVSKKHLEEGRLYPPLVTIRDVSLKIAVKIAEQAYSNKTASFYPEPEDKEAFIRSQVYSTDYDNFVVDSYVWPEEAMKIQTGNP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J233</accession>
    <name>A0A452J233_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Major facilitator superfamily (MFS) profile domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the major facilitator superfamily. Sugar transporter (TC 2.A.1.1) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J233"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039025.1">
        <property type="protein sequence ID" value="ENSGAGP00000034469.1"/>
        <property type="gene ID" value="ENSGAGG00000024490.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005903">
        <property type="term" value="C:brush border"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055056">
        <property type="term" value="F:D-glucose transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046323">
        <property type="term" value="P:D-glucose import"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070837">
        <property type="term" value="P:dehydroascorbic acid transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1250.20:FF:000029">
        <property type="entry name" value="solute carrier family 2, facilitated glucose transporter member 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1250.20">
        <property type="entry name" value="MFS general substrate transporter like domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045263">
        <property type="entry name" value="GLUT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020846">
        <property type="entry name" value="MFS_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005828">
        <property type="entry name" value="MFS_sugar_transport-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036259">
        <property type="entry name" value="MFS_trans_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003663">
        <property type="entry name" value="Sugar/inositol_transpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005829">
        <property type="entry name" value="Sugar_transporter_CS"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00879">
        <property type="entry name" value="SP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23503">
        <property type="entry name" value="SOLUTE CARRIER FAMILY 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23503:SF27">
        <property type="entry name" value="SOLUTE CARRIER FAMILY 2, FACILITATED GLUCOSE TRANSPORTER MEMBER 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00083">
        <property type="entry name" value="Sugar_tr"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00171">
        <property type="entry name" value="SUGRTRNSPORT"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103473">
        <property type="entry name" value="MFS general substrate transporter"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50850">
        <property type="entry name" value="MFS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00217">
        <property type="entry name" value="SUGAR_TRANSPORT_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 6" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 6" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="3 6" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="5" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="33"/>
            <end position="53"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="65"/>
            <end position="83"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="89"/>
            <end position="114"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="126"/>
            <end position="144"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="156"/>
            <end position="177"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="244"/>
            <end position="267"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="279"/>
            <end position="300"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="307"/>
            <end position="331"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="337"/>
            <end position="358"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="370"/>
            <end position="393"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="399"/>
            <end position="417"/>
        </location>
    </feature>
    <feature type="domain" description="Major facilitator superfamily (MFS) profile" evidence="7">
        <location>
            <begin position="1"/>
            <end position="423"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="RuleBase" id="RU003346"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50850"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034469.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="460" mass="50087" checksum="C3B2102DBFDF3A31" modified="2019-05-08" version="1">IAINILNLHDFPFSLQVIEAHYGRVLGISTLTLYWSLSVSIFAIGGMISSFTVGWVGDKLGRVKAMLAVNVLSIIGNLLMGLAKFGPSHILIIAGRAITGLYCGLSSGLVPMYVGEISPTALRGALGTLHQLAIVTGILLSQVLGLDFLLGNDDMWPLLLGLSGAAALLQFFLLLLCPESPRYLYIKLGKVEEAKKSKKLILLRGDCDPTKEITEMEKEKQEAASEKKVSIRKLFSSSSYRQPIIVALMVQISQQFSGINAIFYYSTNIFVRAGVDQPVYATIGVGVVNTIFTIISVFLVERTGRRSLFIAGLIGMLASAVAMTVGLVLLVSYVSMVAIFLFVIFFEVGPGPIPWFIVAELFSQGPRPAAIAIAGFCNWTCNFIIGMCFQYIADLCGPYVFTIFAVLLLGFILFAHFKVPETKGKSFEEISAAFRRKRRSAIKGHKAATELEDLRGSEEA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J250</accession>
    <name>A0A452J250_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Alpha-type protein kinase domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J250"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039026.1">
        <property type="protein sequence ID" value="ENSGAGP00000034470.1"/>
        <property type="gene ID" value="ENSGAGG00000024500.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005929">
        <property type="term" value="C:cilium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048029">
        <property type="term" value="F:monosaccharide binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002753">
        <property type="term" value="P:cytoplasmic pattern recognition receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045087">
        <property type="term" value="P:innate immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd16969">
        <property type="entry name" value="Alpha_kinase_ALPK1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.20.200.10">
        <property type="entry name" value="MHCK/EF2 kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004166">
        <property type="entry name" value="a-kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043529">
        <property type="entry name" value="ALPK1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46747">
        <property type="entry name" value="ALPHA-PROTEIN KINASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46747:SF1">
        <property type="entry name" value="ALPHA-PROTEIN KINASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02816">
        <property type="entry name" value="Alpha_kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00811">
        <property type="entry name" value="Alpha_kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51158">
        <property type="entry name" value="ALPHA_KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0418">Kinase</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="2" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Alpha-type protein kinase" evidence="5">
        <location>
            <begin position="1003"/>
            <end position="1225"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="519"/>
            <end position="542"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="577"/>
            <end position="607"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="654"/>
            <end position="678"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="847"/>
            <end position="888"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="893"/>
            <end position="912"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="522"/>
            <end position="532"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="594"/>
            <end position="607"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="847"/>
            <end position="859"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS51158"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034470.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1231" mass="138841" checksum="BB751C40C4BEC409" modified="2019-05-08" version="1">MNNQKVVATLLQECKQALDVLSRKMSDASEEDKREYQQCKASLPDDLRTLIEEAKEMKWPFVPEKWQYKQAIGPEDKTNLQDMISARLHELLIYLKASIMVKDCATAAAVVFLIDRFLYWVDASSKLLRIAKGLHKLQPATPIAPQVVIRLARISVNSGKLLKAEYILSSLINDNGATGVWLYDKESDRILVQSVCIHIRGQILQKLGMWYEAAELIWASIVGYFKLPQPDKKSLLQEFDHRLLSAAEACKLAAAFSQYTPLFVLTAVNIRGTCLLSYSYSTECPPEKKKFYLSEAKESFQIGLLTKKENDPVTSRQELHSFIKAAFCLTTVHRRLYGESKKLCEVRQLCSEAMEKLYTYSTSPFTEDQDKETLAKEIMSLIMSVKEHLEVQSFPNSDARSYVPDNYKGGAEKPILRGEVNFEKLLEVHSQHHRSVSEVFESTCRSQETRHREAKVGACITALKTETKNVDTMCTTEEKLHCGKYSAKISDAQNSRRSRERLSGQRRWISSNVMNVSLEEEMERKTSDKVTSDEDGIFSSLQSRSQTTSSLKSWSKLSKSSSSACWEELDCNASNKMPSDWQPREEGSVDEQCCTASTESTEENVQAGSLHSLSFQTHHLSLQESQSGCSGPSRHSLKKNMLFANRLADASSASREGLLQGTEPERDASEQNAGDASSIDFRNNAISFMDIKGAHCSGPNRPRRETISSSLTELESFEMIESRTKDSNRDFSVIHSPAQHSIVEGNVIDAERYEKVTEPLSRNTYVWIDPEGDTVDNTEDLSVEPHVPIGKVATTTSINNKPKIASDSFVFNWINKSSAEMAGNESFEMPEVDTQADTEDEWEFAFSNSRSSVNNSPATNAPRPQLSHRVGIRVSPSGRKKPEADDFNDCATAEEDEGEKYRNIPNSSHSSSSSLKLWYKSAQFSSSFSELESPSFLNSSGSSFVFVPGRTKEQILKARFLEDADYEKLLSGVKHSWLVQRLQTTGIFKARKLHKAYSALLLKYSKKSGLWTGQETAVYIGDYLSVAKEGKQRNAFWIHFLHQEETLGRYVGKEYKEEKGLLNHFIDVERQMTAQYYVTEFNKRLYEQKIPTQIFYIPSAILLILEGKSIKGCVSVEPYILGEFVKLSNNTKVVKTEYKATEYGLAFGHFSYEFSHGTDVVVDLQGWVTGNGKGLIYLTDPQIHSLNGKDVSYSTNFGEKGIYYFFNNQHVECNEICHHLSLTRPSVAEPN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J251</accession>
    <name>A0A452J251_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Beta-2-glycoprotein 1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="11">Apolipoprotein H</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="12">Beta-2-glycoprotein I</fullName>
        </alternativeName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">Binds to various kinds of negatively charged substances such as heparin, phospholipids, and dextran sulfate. May prevent activation of the intrinsic blood coagulation cascade by binding to phospholipids on the surface of damaged cells.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="13">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J251"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038914.1">
        <property type="protein sequence ID" value="ENSGAGP00000034371.1"/>
        <property type="gene ID" value="ENSGAGG00000024413.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00033">
        <property type="entry name" value="CCP"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.70.10">
        <property type="entry name" value="Complement Module, domain 1"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050350">
        <property type="entry name" value="Compl-Cell_Adhes-Reg"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035976">
        <property type="entry name" value="Sushi/SCR/CCP_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015104">
        <property type="entry name" value="Sushi_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000436">
        <property type="entry name" value="Sushi_SCR_CCP_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19325:SF549">
        <property type="entry name" value="BETA-2-GLYCOPROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19325">
        <property type="entry name" value="COMPLEMENT COMPONENT-RELATED SUSHI DOMAIN-CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00084">
        <property type="entry name" value="Sushi"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09014">
        <property type="entry name" value="Sushi_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00032">
        <property type="entry name" value="CCP"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57535">
        <property type="entry name" value="Complement control module/SCR domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50923">
        <property type="entry name" value="SUSHI"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="9 13" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6" id="KW-0358">Heparin-binding</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="7 14" id="KW-0732">Signal</keyword>
    <keyword evidence="5 13" id="KW-0768">Sushi</keyword>
    <feature type="signal peptide" evidence="14">
        <location>
            <begin position="1"/>
            <end position="19"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019101085" description="Beta-2-glycoprotein 1" evidence="14">
        <location>
            <begin position="20"/>
            <end position="319"/>
        </location>
    </feature>
    <feature type="domain" description="Sushi" evidence="15">
        <location>
            <begin position="21"/>
            <end position="81"/>
        </location>
    </feature>
    <feature type="domain" description="Sushi" evidence="15">
        <location>
            <begin position="111"/>
            <end position="173"/>
        </location>
    </feature>
    <feature type="domain" description="Sushi" evidence="15">
        <location>
            <begin position="174"/>
            <end position="233"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="13">
        <location>
            <begin position="23"/>
            <end position="66"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="13">
        <location>
            <begin position="176"/>
            <end position="219"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00003651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00020104"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022659"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022674"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00029855"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00033414"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00302"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50923"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034371.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="319" mass="35895" checksum="A22A7FC12E8AAFE3" modified="2019-05-08" version="1">AMPTVLFLWIVTLAHCALAGHVCHKPPEVPFATVDVDKRVYEVGEEITYTCDPGYSHQSGSRKYTCPLSGKWPINRMRCIPKKCPYPEAMRNGIVHVIDLNYQGLIHFSSVTCPPPSLPEFGVISYHKLTPGNESVFQDIIRFECLPSFALFGNETATCMANGNWSDIPECRYVKCPHPTGIENGFINFEVRRTYHYMDSVDFGCQPHYVLDGPRESRCEKTGSWSIKPTCKAPCKIPVKKATVLYNGRKIKVQKDLKEGIQHAETIFFFCKNEIKKCGYTIPTQCINGQLTVPSCFKEIGIFAAIWKTDPADLTPCDP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J252</accession>
    <name>A0A452J252_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">C2H2-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J252"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039035.1">
        <property type="protein sequence ID" value="ENSGAGP00000034476.1"/>
        <property type="gene ID" value="ENSGAGG00000024499.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025750">
        <property type="entry name" value="DPF1-3_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051580">
        <property type="entry name" value="ZnF-Chromatin_assoc"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23057">
        <property type="entry name" value="JUXTAPOSED WITH ANOTHER ZINC FINGER PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23057:SF4">
        <property type="entry name" value="ZINC FINGER PROTEIN DPF3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14051">
        <property type="entry name" value="DPF1-3_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0862">Zinc</keyword>
    <keyword evidence="1" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="3">
        <location>
            <begin position="177"/>
            <end position="205"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="204"/>
            <end position="233"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="273"/>
            <end position="334"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="273"/>
            <end position="288"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="292"/>
            <end position="302"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034476.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="334" mass="37052" checksum="71589E055F99DF7C" modified="2019-05-08" version="1">MQSLTLCSSPPGRLGDQFYKEAIEHCRSYNSRLCAERSVRLPFLDSQTGVAQNNCYIWMEKRHRGPGLAPGQLYTYPARCWRKKRRLHPPEDSRLKLLEIKPEVDLPLKKDGFTSEGTTLEALLRGEGIEKKVDTKEEDTIQEIQVKKCIYIGHARGSGGGRRRNDAASQDDHDKPFVCDICGKRYKNRPGLSYHYAHTHLASEEGDEAQEQETRSPPVHRNENHKPQKGPDGAIIPNNYCDFCLGGSSMNKKSGCPEELVSCSDCGRSAHLGREGRKEEAAPARTTEDLFGSTSESDTSTFHGFDEDDAEEPLLSRGGGHGSSSPSTNKKGGC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J256</accession>
    <name>A0A452J256_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="12">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Golgi apparatus membrane</location>
            <topology evidence="1">Peripheral membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the DOP1 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000039045.1">
        <property type="protein sequence ID" value="ENSGAGP00000034485.1"/>
        <property type="gene ID" value="ENSGAGG00000024506.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="GOC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005768">
        <property type="term" value="C:endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000139">
        <property type="term" value="C:Golgi membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005802">
        <property type="term" value="C:trans-Golgi network"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006895">
        <property type="term" value="P:Golgi to endosome transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040314">
        <property type="entry name" value="DOP1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056457">
        <property type="entry name" value="DOP1_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007249">
        <property type="entry name" value="DOP1_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056459">
        <property type="entry name" value="TPR_DOP1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056458">
        <property type="entry name" value="TPR_DOP1_M"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14042">
        <property type="entry name" value="DOPEY-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14042:SF22">
        <property type="entry name" value="PROTEIN DOPEY-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24598">
        <property type="entry name" value="DOP1_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04118">
        <property type="entry name" value="Dopey_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24601">
        <property type="entry name" value="TPR_DOP1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24597">
        <property type="entry name" value="TPR_DOP1_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="5" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0653">Protein transport</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0813">Transport</keyword>
    <feature type="domain" description="DOP1 N-terminal" evidence="8">
        <location>
            <begin position="11"/>
            <end position="295"/>
        </location>
    </feature>
    <feature type="domain" description="DOP1-like middle TPR" evidence="9">
        <location>
            <begin position="309"/>
            <end position="462"/>
        </location>
    </feature>
    <feature type="domain" description="DOP1-like TPR" evidence="11">
        <location>
            <begin position="1352"/>
            <end position="1726"/>
        </location>
    </feature>
    <feature type="domain" description="DOP1-like C-terminal" evidence="10">
        <location>
            <begin position="1851"/>
            <end position="2371"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="569"/>
            <end position="590"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="613"/>
            <end position="639"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="706"/>
            <end position="726"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1164"/>
            <end position="1208"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1277"/>
            <end position="1302"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="624"/>
            <end position="639"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="711"/>
            <end position="722"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="1164"/>
            <end position="1193"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="1194"/>
            <end position="1208"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004395"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00046326"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF04118"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF24597"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="Pfam" id="PF24598"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF24601"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034485.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2442" mass="275156" checksum="FF09AA631D148D03" modified="2019-05-08" version="1">MNTEEVELLSDSKYRNYVAAVDKALKNFEYSSEWADLISALGKLNKVLQNNAKYQVVPKKLTIGKRLAQCLHPALPGGVHRKALETYEIIFKIIGPKRLAKDLFLYSSGLFPLLANAAMSVKPALLSLYEIYYLPLGKTLKPGLQGLLTGILPGLEEGSEYYERTNTLLEKVAGVDQSAFYSALWGSLLTSPAVRLPGITYVLSHLNRKLSMEDQLYIIGSDIELMVEAVSTSVQDSSVLVQRSTLDLILFCFPFHMSQATRPDMIRILSAALHVVLRRDMSLNRRLYAWLLGPRSTRQNNPEEHATYYFNTFSKEMLVQAMVGILQVNGHGEESTMMQDLKPFRILISLLDKPELGPVILEDVLIEVFRTLYTQCKAELDLQMEPSFSKDHTQLSSKLRENKKTAELIKTANLLFNSFEPYYMWDYIARWFEECCRRTLHARLQTGPVGGNESSELPLTNFCLLVDFLLDIVSLPTRSMRVLCQETYIEIQTEHLPQLLLRMISALTSHLHTLHLSELTDSLRLCSKILSKVQPPLLSAGTDGTLQLPSGHNSSIKEWEDKKVPSFSIENPHDVFEDGENPPSSRSSESGFTEFVQYQADTADDIDRALSEGHGAPGIPIIGSTSSETETASTVGSEETIVHPPSIITQGSATQSGKMIQKTAMQCCLEYVQQFLTRFINLYIIQSNSLSQPLATELQVDLTREQGQPAKWDRESQGDAKGKKTNKTPKEYLSAFIAACQLYLECSSFPVYIAEGNHTSELHPEKSDFDCEQVQPPLWLQTLMNACRQASDFSIQGIAISLIMDLVGLTQSVAVVTGENLNSVETAQPLSPNQGRVAVVIRPPLTQGNLRYMVEKTDFFKHVALTLWDQLGDGTPQHHQKSVELFYQLHNLVPSSSICEDVISQQLTHRDKKTRMETHAKFAVLWHLTRDLHINKSSSFGRSFDRSLFIMLDSLNSLDGSTWSVGQAWLNQVLQRHDIARVLEPLLLLLLHPKTQRVSVQRIQAECYWNKSPHYPEEETEKHFMQKFSCGDAFSHMPVSQGQLITPKENNEKQLIMDEMENFSLTVNPLSDRLSLLSTSSETIPMVVSDFELPDHQIEILQSSDSGCSQSSAGDNLSYEVEAESLSAQNSSQTLREDSPDEIVQQVVIDLICKVVSGLGEETESVKHDLHPDDASSKFSTLDHSEEVSKSEDQNIQSSENSLFGNDSSQLLSASTEIGLERLTDSPPCIEVSSQAPCDLTFSSTDGKSGQRSHSSIQFSFKGKLPEKVSEKETIVKEAGKQPGAKPKVKIARRKDEDKKKAQTEKLKQTSIFFSDGLDLENWYSCGEGEISEIESDVGSPGIRKSPSFNVHPLYQHVLLYLQLYDSSRTLYAFSAIKAILKTNPSAFVNAISTTSVNNVYTPQLSLLQNLLARHRISVMGKDFYSHIPVDSNHNFRSSMYIEIIISLCLYYMRSHYPTHVKVTPQDLIGNRNMQMMSIEILTLLFSELAKVIESSAKGFPSFISDMLSKCKVQKVILHCLLSSIFSAQKWHSEKVAGKNIVVIEEGFSEDSLINFSEDEFDSGSTLQSQLLKVLQRLIVLEHRVMTVPEENETGFEFAITDLEQINPQQPMTSLQYLHSQPITSQGMFLCAVIRALHQHCACKMHPQWIGLITSTLPYMGKVLQRVVVSVTLQLCRNLDNLIQQYKYETGLSNSRPLWMASITPPDMILTLLEGITTIIHYCLLDPSTQYHQLLVNVDQKHLLEARSGILSILHMIMSSVTLLWSILHQADSSEKTTAAAAASVTTINLGSTKVIPTASDEQLLLVELVRSISVMRTETVIQTVKEVLKQPPAIAKDKKHLSLEVCMLQFFYAYIQRIPVSSLVDSWATLLLLLKDSIQLSLPAPGQFLILGVLNEFIMKNPSLENKKDQRDLQDVTHKIVDAIGAIAGSSLEQTTWLRRNLEVKPSLKIMVDGNNLESDVEDMLYPALETSNITPSVYSVHALTLLSEVLAHLLDMVFYSDEKERVIPLLVNIMHYVVPYLRNHSAHNAPSYRACVQLLSSLSGYQYTRKAWKKEAFDLFMDSSFFQMDASCVNHWRAIMDNLMTHDKTTFRDLMTRVAVAQSSSLNLFANRDVELEQRAMLLKRLAFAIFSSEVDQYQKYLPDIQERLVESLRLPQVPTLHSQVFLFFRVLLLRMSPQHLTSLWPTMITELVQVFLLMEQELTADEDISRTSGPSVAGLETTYTGGNGFSTSYNSQRWLNLYLSACKFLDLALALPSENLPQFQMYRWAFIPEVSDDSGLEVRRQGTHQREFKPYVVRLAKLLRKRAKDKQEDFKMMILEGVEMAKHQKNPEEDCSGKTLSWEPGHLLLTIYTVRSIEQLLPFFNVLSQVFNSKVTSRCVGHSGSPVLYPNSFPSKDIKMENQKTFSSKARQKIEEMVEKDFLEGVIKTHLQPRGTRKA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J258</accession>
    <name>A0A452J258_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Ig-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J258"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039065.1">
        <property type="protein sequence ID" value="ENSGAGP00000034502.1"/>
        <property type="gene ID" value="ENSGAGG00000024508.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002764">
        <property type="term" value="P:immune response-regulating signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000049">
        <property type="entry name" value="Leukocyte immunoglobulin-like receptor subfamily B member 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050412">
        <property type="entry name" value="Ig-like_Receptors_ImmuneReg"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11738">
        <property type="entry name" value="MHC CLASS I NK CELL RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11738:SF186">
        <property type="entry name" value="OSTEOCLAST-ASSOCIATED IMMUNOGLOBULIN-LIKE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13895">
        <property type="entry name" value="Ig_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0732">Signal</keyword>
    <feature type="domain" description="Ig-like" evidence="4">
        <location>
            <begin position="37"/>
            <end position="105"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034502.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="324" mass="35199" checksum="65C0B3D925CAF5F7" modified="2019-05-08" version="1">MQICTRCKICVCVSRLPVVLPSGYCLNAATWHEFLKPTIWVSPSRVVALGRNVTIRCEGQDLGMEFVLCKAGHPNQQVQSVVPEGKMAEFPIASVGQEDGGSYTCYYRSITDQSHWSFPSNPVEIIVGEPSYPKPNISGIPRRGVSVGEAMAIWCRRQRQGVRFVLNKEGRHFPPVDSDGFGAVFPISNVSREDGGNYSCSYHSRSEPFAVSDPSDPMELVVRGEGPVASTGTRGSRSAHGRFWFVSLCRKRSHTETEQVGTWILPLLIDLPGSQGSWVLSPALGGVGGWGPGLLRDALGLLTLEGHDPRTAGVTPLTTHPGKQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J262</accession>
    <name>A0A452J262_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">DOP1 N-terminal domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="3">Belongs to the DOP1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J262"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039070.1">
        <property type="protein sequence ID" value="ENSGAGP00000034507.1"/>
        <property type="gene ID" value="ENSGAGG00000024506.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="GOC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005768">
        <property type="term" value="C:endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005802">
        <property type="term" value="C:trans-Golgi network"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006895">
        <property type="term" value="P:Golgi to endosome transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
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        <property type="entry name" value="DOP1"/>
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        <property type="entry name" value="DOP1_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14042">
        <property type="entry name" value="DOPEY-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14042:SF22">
        <property type="entry name" value="PROTEIN DOPEY-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04118">
        <property type="entry name" value="Dopey_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0653">Protein transport</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0813">Transport</keyword>
    <feature type="domain" description="DOP1 N-terminal" evidence="4">
        <location>
            <begin position="11"/>
            <end position="106"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00046326"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="Pfam" id="PF04118"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034507.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="133" mass="15210" checksum="C5E20E6AE42C7101" modified="2019-05-08" version="1">MNTEEVELLSDSKYRNYVAAVDKALKNFEYSSEWADLISALGKLNKVLQNNAKYQVVPKKLTIGKRLAQCLHPALPGGVHRKALETYEIIFKIIGPKRLAKDLFLYRFFADGQYILEQSEPLSSSILQYDKLQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J264</accession>
    <name>A0A452J264_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Ig-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J264"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039049.1">
        <property type="protein sequence ID" value="ENSGAGP00000034488.1"/>
        <property type="gene ID" value="ENSGAGG00000024508.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002764">
        <property type="term" value="P:immune response-regulating signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000033">
        <property type="entry name" value="Killer cell immunoglobulin-like receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000049">
        <property type="entry name" value="Leukocyte immunoglobulin-like receptor subfamily B member 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050412">
        <property type="entry name" value="Ig-like_Receptors_ImmuneReg"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11738">
        <property type="entry name" value="MHC CLASS I NK CELL RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11738:SF186">
        <property type="entry name" value="OSTEOCLAST-ASSOCIATED IMMUNOGLOBULIN-LIKE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13895">
        <property type="entry name" value="Ig_2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="3"/>
    </dbReference>
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        <property type="entry name" value="IGc2"/>
        <property type="match status" value="3"/>
    </dbReference>
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        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="4" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="5" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <keyword evidence="1" id="KW-0732">Signal</keyword>
    <feature type="domain" description="Ig-like" evidence="7">
        <location>
            <begin position="27"/>
            <end position="95"/>
        </location>
    </feature>
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        <location>
            <begin position="344"/>
            <end position="374"/>
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    </feature>
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        <location>
            <begin position="363"/>
            <end position="374"/>
        </location>
    </feature>
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        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034488.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="374" mass="41090" checksum="DC107B813ED34DC2" modified="2019-05-08" version="1">PSPALTAPLRGRMTDGLNWAGHEFLKPTIWVSPSRVVALGRNVTIRCEGQDLGMEFVLCKAGHPNQQVQSVVPEGKMAEFPIASVGQEDGGSYTCYYRSITDQSHWSFPSNPVEIIVGEPSYPKPNISGIPRRGVSVGEAMAIWCRRQRQGVRFVLNKEGRHFPPVDSDGFGAVFPISNVSREDGGNYSCSYHSRSEPFAVSDPSDPMELPSYPKPNISGIPRRGVSVGEAMAIWCRRQRQGVRFVLNKEGRHFPPVDSDGFGAVFPISNVSREDGGNYSCSYHSRSEPHFLPLHPQEPNPGADRLTYAELDNQVLQAKRGAKPPAPEPAQPSMYAAINVNWGARGKRPPQSWGTGPQGATDFKARRVHPDHLV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J267</accession>
    <name>A0A452J267_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">RNA-binding protein 25</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="13">RNA-binding motif protein 25</fullName>
        </alternativeName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="11">Interacts with LUC7L3 and SRRM1. Specifically associates with functional splicing complexes, including Sm proteins and U1, U2, U4, U5 and U6 snRNAs. Associates with exon junction complex (EJC) proteins, including APEX1, DDX39B, NCBP1, RBM8A and RNPS1. Interaction with NCBP1 is RNA-dependent.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J267"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039075.1">
        <property type="protein sequence ID" value="ENSGAGP00000034512.1"/>
        <property type="gene ID" value="ENSGAGG00000024524.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005681">
        <property type="term" value="C:spliceosomal complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003729">
        <property type="term" value="F:mRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006397">
        <property type="term" value="P:mRNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000381">
        <property type="term" value="P:regulation of alternative mRNA splicing, via spliceosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008380">
        <property type="term" value="P:RNA splicing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd12446">
        <property type="entry name" value="RRM_RBM25"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1390.10:FF:000004">
        <property type="entry name" value="RNA-binding motif protein 25"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.330:FF:000230">
        <property type="entry name" value="RNA-binding motif protein 25"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.330">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1390.10">
        <property type="entry name" value="PWI domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012677">
        <property type="entry name" value="Nucleotide-bd_a/b_plait_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002483">
        <property type="entry name" value="PWI_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036483">
        <property type="entry name" value="PWI_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035979">
        <property type="entry name" value="RBD_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052768">
        <property type="entry name" value="RBM25"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034268">
        <property type="entry name" value="RBM25_RRM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000504">
        <property type="entry name" value="RRM_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18806">
        <property type="entry name" value="RBM25 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18806:SF4">
        <property type="entry name" value="RNA-BINDING PROTEIN 25"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01480">
        <property type="entry name" value="PWI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00076">
        <property type="entry name" value="RRM_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00311">
        <property type="entry name" value="PWI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00360">
        <property type="entry name" value="RRM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF101233">
        <property type="entry name" value="PWI domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54928">
        <property type="entry name" value="RNA-binding domain, RBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51025">
        <property type="entry name" value="PWI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50102">
        <property type="entry name" value="RRM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8" id="KW-0007">Acetylation</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="3" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="5" id="KW-0507">mRNA processing</keyword>
    <keyword evidence="9" id="KW-0508">mRNA splicing</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7 14" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="6" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="RRM" evidence="16">
        <location>
            <begin position="50"/>
            <end position="127"/>
        </location>
    </feature>
    <feature type="domain" description="PWI" evidence="17">
        <location>
            <begin position="683"/>
            <end position="776"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="29"/>
            <end position="49"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="135"/>
            <end position="163"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="183"/>
            <end position="205"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="241"/>
            <end position="402"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="454"/>
            <end position="665"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="15">
        <location>
            <begin position="30"/>
            <end position="43"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="15">
        <location>
            <begin position="241"/>
            <end position="393"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="15">
        <location>
            <begin position="454"/>
            <end position="506"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="15">
        <location>
            <begin position="523"/>
            <end position="532"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="15">
        <location>
            <begin position="554"/>
            <end position="563"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022664"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023187"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00062484"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00067881"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00075704"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00176"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50102"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS51025"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034512.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="776" mass="92239" checksum="B0E62F1A2CC447B3" modified="2019-05-08" version="1">MIPVPMGIMAPAPTVLVPTAVSMVGKHLGARKEHPGLKNKENDENSGPTTTVFVGNISEKASDMLIRQLLAKCGLVLSWKRVQGASGKLQAFGFCEYKEPESTLRALRLLHDLQIGEKKLLVKVDAKTKAQLDEWKAKKRASNGNSRPESTNDDDEALDEETKRRDQLIKGAIEVLIREYSSELNAPSQESDSHPRKKKKEKKEDIFRRFPVAPLIPYPLITKEDINAIEMEEDKRDLISREISKFRDTHKKLEEEKGKKEKERQEIEKERRERERERERERERREREREREREREREKEKERERERERDRDRDRTKDRDRDRDRDRDRDRERSSDRNKDRSRSREKSRDREREREREREREREREREREREREREREREREREREKDKKRDREEDEEDAYERRKLERKLREKEAAYQEVIWEYYVSEKAFIRLFLTFKHKVCIFFRGSALQKRLRDREKEMEGDERDRKREKEELEEIRQRLLAEGHPDPDAELQRMEQEAERRRQPQIKQEPESEEEEEEKPEKEEKREEPEEEEEEPEQKPCLKPTLRPISSAPSVSSASGNATPNTPGDESPCGIIIPHENSPDQQQPEEHRPKIGLSLKLGACNSPSQPNAVKRKKLPVDSVFNKFEDEDSDDVPRKRKLVPLDYGEEDKGSSKGNVNTEEKRKHIKSLIEKIPTAKPELFAYPLDWSIVDSTLMERRIRPWINKKIIEYIGEEEATLVDFVCSKVMAHSSPQSILDDVAMVLDEEAEVFIVKMWRLLIYETEAKKIGLVK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J268</accession>
    <name>A0A452J268_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Ig-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J268"/>
    <dbReference type="Ensembl" id="ENSGAGT00000039062.1">
        <property type="protein sequence ID" value="ENSGAGP00000034499.1"/>
        <property type="gene ID" value="ENSGAGG00000024508.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002764">
        <property type="term" value="P:immune response-regulating signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000049">
        <property type="entry name" value="Leukocyte immunoglobulin-like receptor subfamily B member 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050412">
        <property type="entry name" value="Ig-like_Receptors_ImmuneReg"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11738">
        <property type="entry name" value="MHC CLASS I NK CELL RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11738:SF186">
        <property type="entry name" value="OSTEOCLAST-ASSOCIATED IMMUNOGLOBULIN-LIKE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13895">
        <property type="entry name" value="Ig_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0732">Signal</keyword>
    <feature type="domain" description="Ig-like" evidence="5">
        <location>
            <begin position="38"/>
            <end position="106"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="1"/>
            <end position="29"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="1"/>
            <end position="12"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034499.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="325" mass="35048" checksum="B810451FC2B411D0" modified="2019-05-08" version="1">MVSSASATVSASIEDLPPPRGSTQQNLPPPGWHEFLKPTIWVSPSRVVALGRNVTIRCEGQDLGMEFVLCKAGHPNQQVQSVVPEGKMAEFPIASVGQEDGGSYTCYYRSITDQSHWSFPSNPVEIIVGEPSYPKPNISGIPRRGVSVGEAMAIWCRRQRQGVRFVLNKEGRHFPPVDSDGFGAVFPISNVSREDGGNYSCSYHSRSEPFAVSDPSDPMELVVRGEGPVASTGTRGSRSAHGRFWFVSLCRKRSHTETEQVGTWILPLLIDLPGSQGSWVLSPALGGVGGWGPGLLRDALGLLTLEGHDPRTAGVTPLTTHPGKQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJA8</accession>
    <name>A0A452GJA8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Transmembrane protein 196</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJA8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001905.1">
        <property type="protein sequence ID" value="ENSGAGP00000001676.1"/>
        <property type="gene ID" value="ENSGAGG00000001339.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037661">
        <property type="entry name" value="TMEM196"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR28681">
        <property type="entry name" value="TRANSMEMBRANE PROTEIN 196"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR28681:SF1">
        <property type="entry name" value="TRANSMEMBRANE PROTEIN 196"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6 8" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 8" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 8" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="12"/>
            <end position="38"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="50"/>
            <end position="67"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="74"/>
            <end position="95"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="107"/>
            <end position="132"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00044525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001676.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="182" mass="19597" checksum="1B660F9B0F803EF1" modified="2019-05-08" version="1">MCTSSHTIGSLLVLSVLEIGLGVSSVAVGAVSFSLVLAEHKPQLGDSSPVWSGVCFLLCGICGILCAKKKSGLVMILFSACCICGLIGGILNFQFLRALTKKSSTLYSLHLTSMSLACIGIGGCTLSSWLTCRLASYEQRRMFSEREHSLHHSHEMAEKEMTDNMSNGGPQLIYNGRVYEKI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJD0</accession>
    <name>A0A452GJD0_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="10">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GJD0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002014.1">
        <property type="protein sequence ID" value="ENSGAGP00000001772.1"/>
        <property type="gene ID" value="ENSGAGG00000000203.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031594">
        <property type="term" value="C:neuromuscular junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098831">
        <property type="term" value="C:presynaptic active zone cytoplasmic component"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042734">
        <property type="term" value="C:presynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030672">
        <property type="term" value="C:synaptic vesicle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043195">
        <property type="term" value="C:terminal bouton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019992">
        <property type="term" value="F:diacylglycerol binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005543">
        <property type="term" value="F:phospholipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017075">
        <property type="term" value="F:syntaxin-1 binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061789">
        <property type="term" value="P:dense core granule priming"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0099525">
        <property type="term" value="P:presynaptic dense core vesicle exocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035249">
        <property type="term" value="P:synaptic transmission, glutamatergic"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016081">
        <property type="term" value="P:synaptic vesicle docking"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016082">
        <property type="term" value="P:synaptic vesicle priming"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd04027">
        <property type="entry name" value="C2B_Munc13"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd08395">
        <property type="entry name" value="C2C_Munc13"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.357.50:FF:000001">
        <property type="entry name" value="Protein unc-13 homolog B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000002">
        <property type="entry name" value="Protein unc-13 homolog B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000014">
        <property type="entry name" value="protein unc-13 homolog B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.357.50">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.1100">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000008">
        <property type="entry name" value="C2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010439">
        <property type="entry name" value="MUN_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014770">
        <property type="entry name" value="Munc13_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014772">
        <property type="entry name" value="Munc13_dom-2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027080">
        <property type="entry name" value="Unc-13"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037302">
        <property type="entry name" value="Unc-13_C2B"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10480">
        <property type="entry name" value="PROTEIN UNC-13 HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10480:SF2">
        <property type="entry name" value="PROTEIN UNC-13 HOMOLOG C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00168">
        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06292">
        <property type="entry name" value="MUN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00360">
        <property type="entry name" value="C2DOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00239">
        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM01145">
        <property type="entry name" value="DUF1041"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49562">
        <property type="entry name" value="C2 domain (Calcium/lipid-binding domain, CaLB)"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50004">
        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51258">
        <property type="entry name" value="MHD1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51259">
        <property type="entry name" value="MHD2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0106">Calcium</keyword>
    <keyword evidence="1" id="KW-0268">Exocytosis</keyword>
    <keyword evidence="2" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <keyword evidence="4" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2" evidence="7">
        <location>
            <begin position="36"/>
            <end position="160"/>
        </location>
    </feature>
    <feature type="domain" description="MHD1" evidence="8">
        <location>
            <begin position="466"/>
            <end position="609"/>
        </location>
    </feature>
    <feature type="domain" description="MHD2" evidence="9">
        <location>
            <begin position="715"/>
            <end position="815"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="7">
        <location>
            <begin position="847"/>
            <end position="973"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51258"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51259"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001772.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1018" mass="116915" checksum="CC91786046317228" modified="2019-05-08" version="1">IIAMKERMKIRERNRPEVFEVIQEMFQISKEDFVQYTKAAKQCVLDGTSKWSAKITITVVCAQGLQAKDKTGSSDPYVTVQVGKTKRRTKTIFGNLNPVWDEKFYFECHNSTDRIKVRVWDEDDDIKSRVKQHFKKESDDFLGQTIIEVRTLSGEMDVWYNLEKRTDKSAVSGAIRLKINVEIKGEEKVAPYHVQYTYLHENLFHYLTEVKSNGSVKIPEVRGDEAWKVYFDAAAQEIVDEFAMRYGIEYIYQAMTHFSCLSSKYMCSGVPAVMSTLLANINAFYAHTIATTNVSASDRFAATNFGREKFIKLLDQLHNSLRIDLSKYRDNFPASNSERLQDLKSTVDLLTSITFFRMKVLELQSPPRASMVVKDCVRACLDSTYKYIFDNCYELYSQLMDQKQEVPREEQGPTTKNLDFWAQLITLMVTIIDEDKTAYTPVLNQFPQELNMGKVSAEIMWTLFAQDMKYALEEHEKQRLCKSTDYMNLHFKVKWFYNEYVRELPALQDAVPEYSLWFEPFVIQWLDENEDVSMEFLHGALERDKKDGFQQTSDHALFSCSVVDVFTQLNQSFEIIKKLECPNPEALSHLMRRFAKTINKVLLQYAAVISNDFNSYCDKEMLPCILMNNIQQLRVQLEKMFESMGGKELDPDASVVLKELQVKLSNVLDELSIAYGASFQLVVEECVRQMSFELNQMRGNGNISANKNNAAIDAEIVLRPLMDFLDKILSLSAKICEKTVLKRVLKELWKSVLNKIEKQIVLPPLTDQTVSVQQYFHAGGNGLKKNFLEKSPDLQSLRYALSLYTQTTDALIKKFIVQLDDGVACDGGVGLTFSESYVPESHSTNNTVGEISVQVDVFTHPGTGEHRVTVKVVAINNLNWQTTAMFRPFVEVCILGPNLSDKKRKHGTKTKSNTWSPKYNETFHILSNEEGLRAYELHLSVKDYCFAREDRIVGMTVIQLQNTAEKGSCASWYPLLKNIFMDDTGLTILRILSQRTNDEVAKEFVRLKSETRSTEETV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJD2</accession>
    <name>A0A452GJD2_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="8">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJD2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001938.1">
        <property type="protein sequence ID" value="ENSGAGP00000001703.1"/>
        <property type="gene ID" value="ENSGAGG00000000765.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015643">
        <property type="term" value="F:toxic substance binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040416">
        <property type="entry name" value="TMEM181"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR054077">
        <property type="entry name" value="TMEM181_GOLD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047843">
        <property type="entry name" value="WLS-like_TM"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31918">
        <property type="entry name" value="TRANSMEMBRANE PROTEIN 181"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31918:SF1">
        <property type="entry name" value="TRANSMEMBRANE PROTEIN 181"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21885">
        <property type="entry name" value="TMEM181_GOLD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06664">
        <property type="entry name" value="WLS-like_TM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4 5" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 5" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="3 5" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="15"/>
            <end position="40"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="177"/>
            <end position="195"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="215"/>
            <end position="233"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="245"/>
            <end position="265"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="277"/>
            <end position="300"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="320"/>
            <end position="344"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="356"/>
            <end position="376"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="382"/>
            <end position="400"/>
        </location>
    </feature>
    <feature type="domain" description="TMEM181 GOLD" evidence="7">
        <location>
            <begin position="56"/>
            <end position="168"/>
        </location>
    </feature>
    <feature type="domain" description="Wntless-like transmembrane" evidence="6">
        <location>
            <begin position="169"/>
            <end position="401"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="Pfam" id="PF06664"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF21885"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001703.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="453" mass="52730" checksum="9DAC24AF75603804" modified="2019-05-08" version="1">MLAPMRLYTLSKRHFVLVFVVFFICFGLTVFIGIAGPSVIESTAVTATLSNLKIRPFKLNSKPLSTYNQQLWLTCMIQLDQPDESSFKTNVTITVKIRGVEENGSSSFIKKQVYNRTRMLNCAQKCAEIIVVHLGYLTYTQYEVIVGFEDLNKLMYPIKNVNFTWKTYNPSFSQVEIWFRFVFVVLTFMVTCLFAHSLRKFSMRDWGIEQKWMSILLPLLLLYNDPFFPLSFLVNSWFPGMLDDLFQSVFLCALLLFWLCVYHGIRVQGERKCLTFYFPKFFIVGLLWLACVTLGIWQTVNELHDPTYQFRVDTGNFQGMKIFFLVVATVYILYLLFLIVRACSELRNMPYVDLRLKFLTALTFVVLVISIVILYLPEFLSFYGLLNFYLYTLAFVYSPSKNALYESQLKDNPAFSMLNDSDDDVIYGSDYEEMPLQNGQAIRAKYKEESDSD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJD3</accession>
    <name>A0A452GJD3_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="9">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endosome</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJD3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002033.1">
        <property type="protein sequence ID" value="ENSGAGP00000001788.1"/>
        <property type="gene ID" value="ENSGAGG00000001421.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005768">
        <property type="term" value="C:endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="CDD" id="cd21255">
        <property type="entry name" value="CH_EHBP1L1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.418.10:FF:000023">
        <property type="entry name" value="EH domain-binding protein 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.418.10">
        <property type="entry name" value="Calponin-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022735">
        <property type="entry name" value="bMERB_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001715">
        <property type="entry name" value="CH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036872">
        <property type="entry name" value="CH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050540">
        <property type="entry name" value="F-actin_Monoox_Mical"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019448">
        <property type="entry name" value="NT-C2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23167">
        <property type="entry name" value="CALPONIN HOMOLOGY DOMAIN-CONTAINING PROTEIN DDB_G0272472-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23167:SF42">
        <property type="entry name" value="EH DOMAIN-BINDING PROTEIN 1-LIKE PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12130">
        <property type="entry name" value="bMERB_dom"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00307">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10358">
        <property type="entry name" value="NT-C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00033">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01203">
        <property type="entry name" value="DUF3585"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47576">
        <property type="entry name" value="Calponin-homology domain, CH-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51848">
        <property type="entry name" value="BMERB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51840">
        <property type="entry name" value="C2_NT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50021">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0967">Endosome</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="C2 NT-type" evidence="7">
        <location>
            <begin position="8"/>
            <end position="157"/>
        </location>
    </feature>
    <feature type="domain" description="Calponin-homology (CH)" evidence="6">
        <location>
            <begin position="309"/>
            <end position="414"/>
        </location>
    </feature>
    <feature type="domain" description="BMERB" evidence="8">
        <location>
            <begin position="753"/>
            <end position="895"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="179"/>
            <end position="307"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="457"/>
            <end position="486"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="528"/>
            <end position="762"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="5">
        <location>
            <begin position="187"/>
            <end position="196"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="197"/>
            <end position="206"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="248"/>
            <end position="261"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="5">
        <location>
            <begin position="287"/>
            <end position="304"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="568"/>
            <end position="577"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="696"/>
            <end position="711"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="723"/>
            <end position="741"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004177"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS51840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51848"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001788.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="895" mass="98516" checksum="E2768507D32E02AD" modified="2019-05-08" version="1">MSSAWKRLQRVGKRASKFHFVACYQELMVECTKKWQPDKLIVVWTRRNRRVCSKAHSWQPGIKNPYRGMVVWMVPENVDIMVTLYRDPHMEEYEDKEWTFVIENESKGQRKVLASADVNLQRFARPTPAQAELKLKLKPRSVKVVAATLQLTLSCVFLREGKATDEDMQSLASLMSVKPSDVGNLDDFADSDEDDEAARRVQHEDGSSQGTAPTDTSRELNTLAEEEENPSAPGTSRAAAGRATPHGRPREAGSAKEEPVRVADTPDAPLRDTPHPALELVLQGAAPSPPPELMGPPPEEPPATSPGLVSCSQSLLEWCQNVTAGYRGVRVTNFTTSWRNGLAFCAILHHFHPEKINYETLDPLAIKENNKLAYDGFASLGISRVLEPADMVFLTVPDKLIVMTYLCQIRAFFTGQELNVVQIANHSSQSTYKVGKFDSDPAFSIDPARFYAERFQSATRGPEEPGSTAPAARDDGKAAANKTVTGGTELAKKLEAELTAPKDAGTDAKIGDAEDAGKVTVNGTISQTTVQSEVEPSVTKDAGKVLINDVTTKPAISPEAEPPTPKDGGSKAAKDAGKSLVNGAMSPSAKKPEAEPLAPEDAGKVPANGDEKLVPPPRLKRQSVRNGEAPDQAERSLQRSLSTGSQGPVAPPRSHGSKSGFSHVRDADLLKKRRSRLKSESLSMDEGEAGGPPGETSRRRSDWLGTDDREGMMTPKAGPPASARTQQPPEIAPSSSTTTQEPSRDPSKETSTTEEEVPRFQDTSQYVVAELQALENEQKQIDARAAVVEKELRFLMESGTNQAQEEELIQEWFTLVNKKNALIRRQDQLQLLMEEQDLERRFELLSRELRAMLATEDCLKTEAQQRREQLLLEELVSLVNQRDELVRDLDIKERM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJD7</accession>
    <name>A0A452GJD7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Olfactory receptor</fullName>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 14">Cell membrane</location>
            <topology evidence="1 14">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 13">Belongs to the G-protein coupled receptor 1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJD7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001943.1">
        <property type="protein sequence ID" value="ENSGAGP00000001708.1"/>
        <property type="gene ID" value="ENSGAGG00000001364.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004930">
        <property type="term" value="F:G protein-coupled receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004984">
        <property type="term" value="F:olfactory receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd15911">
        <property type="entry name" value="7tmA_OR11A-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.1220.70:FF:000001">
        <property type="entry name" value="Olfactory receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1070.10:FF:000010">
        <property type="entry name" value="Olfactory receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1070.10">
        <property type="entry name" value="Rhodopsin 7-helix transmembrane proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000276">
        <property type="entry name" value="GPCR_Rhodpsn"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017452">
        <property type="entry name" value="GPCR_Rhodpsn_7TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000725">
        <property type="entry name" value="Olfact_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050516">
        <property type="entry name" value="Olfactory_GPCR"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR26452">
        <property type="entry name" value="OLFACTORY RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13853">
        <property type="entry name" value="7tm_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00237">
        <property type="entry name" value="GPCRRHODOPSN"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00245">
        <property type="entry name" value="OLFACTORYR"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81321">
        <property type="entry name" value="Family A G protein-coupled receptor-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00237">
        <property type="entry name" value="G_PROTEIN_RECEP_F1_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50262">
        <property type="entry name" value="G_PROTEIN_RECEP_F1_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 14" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="8 13" id="KW-0297">G-protein coupled receptor</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9 14" id="KW-0472">Membrane</keyword>
    <keyword evidence="6 14" id="KW-0552">Olfaction</keyword>
    <keyword evidence="10 13" id="KW-0675">Receptor</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 14" id="KW-0716">Sensory transduction</keyword>
    <keyword evidence="12 13" id="KW-0807">Transducer</keyword>
    <keyword evidence="5 13" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 14" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="27"/>
            <end position="53"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="60"/>
            <end position="80"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="100"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="142"/>
            <end position="166"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="196"/>
            <end position="228"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="240"/>
            <end position="263"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="275"/>
            <end position="294"/>
        </location>
    </feature>
    <feature type="domain" description="G-protein coupled receptors family 1 profile" evidence="15">
        <location>
            <begin position="43"/>
            <end position="292"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010663"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022606"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022725"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023224"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="RuleBase" id="RU000688"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="RuleBase" id="RU363047"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50262"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001708.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="311" mass="34858" checksum="BA66FBA8C0CBDEAE" modified="2019-05-08" version="1">MLLPLKVLSLSFKFILLGFGNLPELQILLFLVFLVIYIVTMSGNILIVALVVADQHLHTPMYFFLGNLSCLETCYTLTILPRLLASLLTGDRTISVSGCFVQFYCFGCLITTECYLLAVMSYDRYLAICKPLHYATLMNSRLCLQLAAGSWISGFLTCVITMWFMLQLIFCGPNEIDHFFCDFSPMLKLSCSDTSMITLVSFILASINTPCPFLLTVTSYVCIIATILRIPSTTGRQKAFSTCSSHLIVVTVFYGTLMTVYLLPKTSTLKALNKVFSVFYTVLTPMLNPLIYSLRNKEVKEALRKPRVCGI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJD9</accession>
    <name>A0A452GJD9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="9">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endosome</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000002027.1">
        <property type="protein sequence ID" value="ENSGAGP00000001782.1"/>
        <property type="gene ID" value="ENSGAGG00000001421.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005768">
        <property type="term" value="C:endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="CDD" id="cd21255">
        <property type="entry name" value="CH_EHBP1L1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.418.10:FF:000023">
        <property type="entry name" value="EH domain-binding protein 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.418.10">
        <property type="entry name" value="Calponin-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022735">
        <property type="entry name" value="bMERB_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001715">
        <property type="entry name" value="CH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036872">
        <property type="entry name" value="CH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050540">
        <property type="entry name" value="F-actin_Monoox_Mical"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019448">
        <property type="entry name" value="NT-C2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23167">
        <property type="entry name" value="CALPONIN HOMOLOGY DOMAIN-CONTAINING PROTEIN DDB_G0272472-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23167:SF42">
        <property type="entry name" value="EH DOMAIN-BINDING PROTEIN 1-LIKE PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12130">
        <property type="entry name" value="bMERB_dom"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00307">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10358">
        <property type="entry name" value="NT-C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00033">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01203">
        <property type="entry name" value="DUF3585"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47576">
        <property type="entry name" value="Calponin-homology domain, CH-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51848">
        <property type="entry name" value="BMERB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51840">
        <property type="entry name" value="C2_NT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50021">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0967">Endosome</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="C2 NT-type" evidence="7">
        <location>
            <begin position="8"/>
            <end position="157"/>
        </location>
    </feature>
    <feature type="domain" description="Calponin-homology (CH)" evidence="6">
        <location>
            <begin position="1058"/>
            <end position="1163"/>
        </location>
    </feature>
    <feature type="domain" description="BMERB" evidence="8">
        <location>
            <begin position="1502"/>
            <end position="1644"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="179"/>
            <end position="492"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="697"/>
            <end position="747"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="884"/>
            <end position="949"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="1000"/>
            <end position="1057"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="1206"/>
            <end position="1235"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="1277"/>
            <end position="1511"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="5">
        <location>
            <begin position="187"/>
            <end position="196"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="197"/>
            <end position="206"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="248"/>
            <end position="261"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="5">
        <location>
            <begin position="287"/>
            <end position="299"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="5">
        <location>
            <begin position="308"/>
            <end position="324"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="352"/>
            <end position="367"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="5">
        <location>
            <begin position="406"/>
            <end position="419"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="5">
        <location>
            <begin position="454"/>
            <end position="463"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="5">
        <location>
            <begin position="479"/>
            <end position="490"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="726"/>
            <end position="736"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="1014"/>
            <end position="1025"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="1317"/>
            <end position="1326"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="1445"/>
            <end position="1460"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="1472"/>
            <end position="1490"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004177"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS51840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51848"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001782.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1644" mass="175097" checksum="41F15F049B073392" modified="2019-05-08" version="1">MSSAWKRLQRVGKRASKFHFVACYQELMVECTKKWQPDKLIVVWTRRNRRVCSKAHSWQPGIKNPYRGMVVWMVPENVDIMVTLYRDPHMEEYEDKEWTFVIENESKGQRKVLASADVNLQRFARPTPAQAELKLKLKPRSVKVVAATLQLTLSCVFLREGKATDEDMQSLASLMSVKPSDVGNLDDFADSDEDDEAARRVQHEDGSSQGTAPTDTSRELNTLAEEEENPSAPGTSRAAAGRATPHGRPREAGSAKEEPVRVADTPDAPLRDTPHPALELVLQGAAPSPPPELMGPPPVSGAGQAGRAGQNMAAGGNPMGAAGAVEPGAKGEMGAQVPDQAHKPGTDPSQETGRHDGNPRAEEEEKTAALMKQPASGSDTKGRPSDIWMPRAESPVPAPRRKKPLLPEPTAPTAPPPEVPSIQRCQWEGAKLPPSQLKLETAKSEQGPRPATSLPVPVVPARVRLARPRSAELPVRPKPLPRPRRDRRARSASPGISVIMYLHKDAPGPAVVAGEGIGLARSGFGAGVPSAAALLAVSSVVGLPPVPWGRQEEKAAADANGWPSVAGAVAPVVVPLKEEQGRATGEGLRGASHLERLTADVAAGKDAGGAVVCVGGGDAASLVGPPKPWRYRSENVVGEGAKGASRVVMKAIPGQLEKGRPEEPTERENCRANRKACGGTAQSVAEVVPSLERHRTELVGQEDSGRGFWPSPAPDLSRAVTPPQLEQRRAAGERPRAASQPERPGAERAAWKDAAGAVVQGGAGDAPSLMAPPKPWRCRSEDVVGQGAGEVPSSVMKAEAGDGLWRVIPAWPEWGRLEGAGGEAAGEPGGGAVHSVTAPLPVEAPAWLERGRLEEAGREAAGRPCGGEAQLVVVDAPSLEASAQLEGAMESPGADGTVSGGATPSTPEPLLERGGPEEAVGSRECGGAVGSLPSKEDASGAGRPLEKQSGLCTDQLTTCIGPGRSSRQAAALALMPPQDAPILGPPSPFAEPYISQCRFTLSLPPTHPSRERSASPSSNEGTQASGREPSPPPCGAARDSVGQEAPQEEPPATSPGLVSCSQSLLEWCQNVTAGYRGVRVTNFTTSWRNGLAFCAILHHFHPEKINYETLDPLAIKENNKLAYDGFASLGISRVLEPADMVFLTVPDKLIVMTYLCQIRAFFTGQELNVVQIANHSSQSTYKVGKFDSDPAFSIDPARFYAERFQSATRGPEEPGSTAPAARDDGKAAANKTVTGGTELAKKLEAELTAPKDAGTDAKIGDAEDAGKVTVNGTISQTTVQSEVEPSVTKDAGKVLINDVTTKPAISPEAEPPTPKDGGSKAAKDAGKSLVNGAMSPSAKKPEAEPLAPEDAGKVPANGDEKLVPPPRLKRQSVRNGEAPDQAERSLQRSLSTGSQGPVAPPRSHGSKSGFSHVRDADLLKKRRSRLKSESLSMDEGEAGGPPGETSRRRSDWLGTDDREGMMTPKAGPPASARTQQPPEIAPSSSTTTQEPSRDPSKETSTTEEEVPRFQDTSQYVVAELQALENEQKQIDARAAVVEKELRFLMESGTNQAQEEELIQEWFTLVNKKNALIRRQDQLQLLMEEQDLERRFELLSRELRAMLATEDCLKTEAQQRREQLLLEELVSLVNQRDELVRDLDIKERM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJE0</accession>
    <name>A0A452GJE0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Adiponectin receptor 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the ADIPOR family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJE0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002044.1">
        <property type="protein sequence ID" value="ENSGAGP00000001794.1"/>
        <property type="gene ID" value="ENSGAGG00000001276.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0038023">
        <property type="term" value="F:signaling receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033211">
        <property type="term" value="P:adiponectin-activated signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004254">
        <property type="entry name" value="AdipoR/HlyIII-related"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR20855:SF35">
        <property type="entry name" value="ADIPONECTIN RECEPTOR 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR20855">
        <property type="entry name" value="ADIPOR/PROGESTIN RECEPTOR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03006">
        <property type="entry name" value="HlyIII"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 7" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 7" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="146"/>
            <end position="166"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="178"/>
            <end position="198"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="219"/>
            <end position="237"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="243"/>
            <end position="260"/>
        </location>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="199"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="316"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="320"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PIRSR" id="PIRSR604254-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001794.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="10">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="354" mass="38875" checksum="D6C5CA378A671A97" modified="2019-05-08" version="1">MQPPLGWSSRGTAALRQGTGLATPNPGAGWEPLPRLHLGPSLQQGAEVGAEPAGWGAAAQWGAEELDNADPVRVLTLPLQAQQAMGRMEEFVFKVWEGRWRVIPYDVLPDWLKDNDFLLHGHRPPMPSFRACFRSIFRLHTETGNIWTHLIGFLFFLILGIGYLFSPNVNYVAPVQERVVFGMFFLGATLCLCFSWLFHTVYCHSERVSRTFSKLDYSGITLLIVGSFVPWLYYSFYCSPQPQLIYLIIVCALGVTAITVSQSDHFSTPQYRAVRAGGGLGAQGGGGAWLAPARRGGRSQAHPSPFLPHSPQFNSHQIFHVLVVAGAGVHLHGVSQLQAFRSSLGGTCTESTVL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJE2</accession>
    <name>A0A452GJE2_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="5">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the THOC5 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJE2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002048.1">
        <property type="protein sequence ID" value="ENSGAGP00000001798.1"/>
        <property type="gene ID" value="ENSGAGG00000001442.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000445">
        <property type="term" value="C:THO complex part of transcription export complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003729">
        <property type="term" value="F:mRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006406">
        <property type="term" value="P:mRNA export from nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019163">
        <property type="entry name" value="THO_Thoc5"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13375">
        <property type="entry name" value="FMS INTERACTING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13375:SF3">
        <property type="entry name" value="THO COMPLEX SUBUNIT 5 HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09766">
        <property type="entry name" value="FmiP_Thoc5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0539">Nucleus</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="1"/>
            <end position="36"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="299"/>
            <end position="330"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="22"/>
            <end position="36"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="4">
        <location>
            <begin position="306"/>
            <end position="319"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008044"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001798.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="663" mass="76021" checksum="D9A7E5F8DB1844BC" modified="2019-05-08" version="1">MSSDSSKKRKPKVIRTDGAPQESKRSRADPDQDVKYYSEEAEVELRDPSKDYELYKQTCQELQRLMAEIQDLKSKGSKDSAVAIEERRIQSCMHFMTLKKLNRLAHIRLKKGRDQTHEAKQKVDAYHLQLQNLLYEVMHLQKEITKCLEFKSKHEEIDLVSVDEFYKEAPPEISKPDVTLNEPHQQTLARLDWELEQRKRLAEKYKECLANKEKILKEIEVKKEYLSSLQPRLNSIMQASLPVQEYLFMPFDQAHKQYETARHLPPPLYVLFVQANAYGQACDKKLAVAIEGSVEEAKALYKPPEDSQDDESDSDAEEEQTTKRRRPTLGVQLDDKRKEMLKRHPLSVTIDLKCKDDSVLHLIFYYLMNLNVMTVKTKVTTATEMATPISAGDLLSPGSLLNCLYPGDHGKRTPNPANQFQFDKVGILTLSDYVTDLGHPYVWVQKLGGLHFPKDQPQHTVTADNSLSASHMEMTMKLLRMRLQSRLALHKQFASLEHGIVPVSSECQHLFPSKVVSRLVKWAALPYEDYLELSYTKDVVEAGLAEDTHLYYTALVERGTAKLQAAVVLNPGYSSMPPIFNLCLNWKGEKTNSNDDNIRAMESEVNVCYKELCGPKPGYQLLTNQLQRLCVVLDVYLETESHDTSVEGPKEFPQEKMCLRLVR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJE5</accession>
    <name>A0A452GJE5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">ATP-binding cassette sub-family C member 10</fullName>
            <ecNumber evidence="3">7.6.2.2</ecNumber>
            <ecNumber evidence="15">7.6.2.3</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="22">Multidrug resistance-associated protein 7</fullName>
        </alternativeName>
    </protein>
    <organism evidence="27 28">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="28" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="27" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="27" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>17beta-estradiol 17-O-(beta-D-glucuronate)(in) + ATP + H2O = 17beta-estradiol 17-O-(beta-D-glucuronate)(out) + ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:60128"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:82961"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="19">
            <dbReference type="Rhea" id="RHEA:60129"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="16">
            <text>ATP + H2O + xenobioticSide 1 = ADP + phosphate + xenobioticSide 2.</text>
            <dbReference type="EC" id="7.6.2.2"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="20">
            <text>an S-substituted glutathione(in) + ATP + H2O = an S-substituted glutathione(out) + ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:19121"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:90779"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="7.6.2.3"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="20">
            <dbReference type="Rhea" id="RHEA:19122"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>leukotriene C4(in) + ATP + H2O = leukotriene C4(out) + ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:38963"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:57973"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="18">
            <dbReference type="Rhea" id="RHEA:38964"/>
        </physiologicalReaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="17">Basal cell membrane</location>
            <topology evidence="17">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Basolateral cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the ABC transporter superfamily. ABCC family. Conjugate transporter (TC 3.A.1.208) subfamily.</text>
    </comment>
    <dbReference type="EC" id="7.6.2.2" evidence="3"/>
    <dbReference type="EC" id="7.6.2.3" evidence="15"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002049.1">
        <property type="protein sequence ID" value="ENSGAGP00000001799.1"/>
        <property type="gene ID" value="ENSGAGG00000001439.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016323">
        <property type="term" value="C:basolateral plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015431">
        <property type="term" value="F:ABC-type glutathione S-conjugate transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008559">
        <property type="term" value="F:ABC-type xenobiotic transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006869">
        <property type="term" value="P:lipid transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd18605">
        <property type="entry name" value="ABC_6TM_MRP7_D2_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd03250">
        <property type="entry name" value="ABCC_MRP_domain1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd03244">
        <property type="entry name" value="ABCC_MRP_domain2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1560.10:FF:000037">
        <property type="entry name" value="ATP-binding cassette subfamily C member 10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:001090">
        <property type="entry name" value="ATP-binding cassette subfamily C member 10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000163">
        <property type="entry name" value="Multidrug resistance-associated protein member 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1560.10">
        <property type="entry name" value="ABC transporter type 1, transmembrane domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003593">
        <property type="entry name" value="AAA+_ATPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011527">
        <property type="entry name" value="ABC1_TM_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036640">
        <property type="entry name" value="ABC1_TM_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003439">
        <property type="entry name" value="ABC_transporter-like_ATP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017871">
        <property type="entry name" value="ABC_transporter-like_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050173">
        <property type="entry name" value="ABC_transporter_C-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24223">
        <property type="entry name" value="ATP-BINDING CASSETTE SUB-FAMILY C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24223:SF330">
        <property type="entry name" value="ATP-BINDING CASSETTE SUB-FAMILY C MEMBER 10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00664">
        <property type="entry name" value="ABC_membrane"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00005">
        <property type="entry name" value="ABC_tran"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00382">
        <property type="entry name" value="AAA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90123">
        <property type="entry name" value="ABC transporter transmembrane region"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50929">
        <property type="entry name" value="ABC_TM1F"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00211">
        <property type="entry name" value="ABC_TRANSPORTER_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50893">
        <property type="entry name" value="ABC_TRANSPORTER_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="5" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="13" id="KW-0445">Lipid transport</keyword>
    <keyword evidence="14 24" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="28" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="11" id="KW-1278">Translocase</keyword>
    <keyword evidence="7 24" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="12 24" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="4" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="24">
        <location>
            <begin position="70"/>
            <end position="90"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="24">
        <location>
            <begin position="96"/>
            <end position="121"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="24">
        <location>
            <begin position="133"/>
            <end position="153"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="24">
        <location>
            <begin position="168"/>
            <end position="189"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="24">
        <location>
            <begin position="341"/>
            <end position="363"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="24">
        <location>
            <begin position="375"/>
            <end position="391"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="24">
        <location>
            <begin position="445"/>
            <end position="469"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="24">
        <location>
            <begin position="475"/>
            <end position="494"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="24">
        <location>
            <begin position="515"/>
            <end position="536"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="24">
        <location>
            <begin position="862"/>
            <end position="882"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="24">
        <location>
            <begin position="920"/>
            <end position="938"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="24">
        <location>
            <begin position="950"/>
            <end position="972"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="24">
        <location>
            <begin position="1025"/>
            <end position="1044"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="24">
        <location>
            <begin position="1050"/>
            <end position="1069"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="24">
        <location>
            <begin position="1137"/>
            <end position="1156"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="24">
        <location>
            <begin position="1168"/>
            <end position="1186"/>
        </location>
    </feature>
    <feature type="domain" description="ABC transmembrane type-1" evidence="26">
        <location>
            <begin position="341"/>
            <end position="517"/>
        </location>
    </feature>
    <feature type="domain" description="ABC transporter" evidence="25">
        <location>
            <begin position="588"/>
            <end position="812"/>
        </location>
    </feature>
    <feature type="domain" description="ABC transmembrane type-1" evidence="26">
        <location>
            <begin position="875"/>
            <end position="1194"/>
        </location>
    </feature>
    <feature type="domain" description="ABC transporter" evidence="25">
        <location>
            <begin position="1230"/>
            <end position="1463"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="23">
        <location>
            <begin position="286"/>
            <end position="314"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="23">
        <location>
            <begin position="302"/>
            <end position="314"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004554"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009726"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012191"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022967"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00024220"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00034018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00034696"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00047523"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00047576"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00048007"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00067405"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00082787"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS50893"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="26">
        <source>
            <dbReference type="PROSITE" id="PS50929"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001799.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="28">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1467" mass="162348" checksum="B1CC1A55F1427901" modified="2019-05-08" version="1">MEEILADLCSTSPEDPLPVWVHGSVGHCFNQLILNVVPHAILALVSACYLGTPRSPVPGASHRPGWKCRIAASFILTGLSILDIISAAITQQEFGTLYLVVLTSGIALLAWLTHSLALLMLYRSTYGSSRGPAAMVFLALFPVPALIITVVWYCQNGVAWPHLHSAHIFRLSILCLQLASLLIYVIGYITPVASRQVFCPINDSWQREPLISEPGIPAPNWHGVAEDGESWLSRFSYAWMNPLMKRGYQWMLNQPQDVYLLPQRLQAARVHDHFYSCWQEKAALSQEEEDTMSSTSPIVTKSDGSGDASSSSGSPCDAQKAVQLLAVLHAAFGLRYYSLGLLRLAGSLLGFSGPLLLNLLVSFMESHQEPLSHGVLYALGLFAGSFLGAVLRNQFSYEVNKVMLTVRAAVISAIYRKALRVSGSSLTGFTMGEIVNFMSTDTDRLVNFCISFHEVWSLPFQFAITLYLLYQQVGVAFLGGLALALLLVPVNKVIANRIMENNKEMLKHKDTRVKVFTALALVGMLILPLNNFPWVLNGILDAKVSLDRIQHFLELSDQDLHAYYITDGPRDPSSALEMHHSTFSWSPPREESTRQHLSGGTLQIFIQDLAVAKGALVGVVGKVGCGKSSLLAAVTGELNRLGGQVYVWDLERGFGLATQEPWIQFTTVRENILFGRDYDARLYQEVLEACALSDDLNILPAGDQTEVGENGVTLSGGQKARIALARAVYQEKEIYLLDDPLAAVDADVANHLMQKCILGVLKHKTRILCTHRTEFLEKADTLLLMDNGRIVKIGTPSEILPLVEATLKVNEKNKRRQNKVAAEQGQEEDMEIEAEELGQATCLLQREEEKKEGAVAFQVYRAYWLAVGSCLALSILFSLLLMQASRNISVWWLSHWISNLPQTANISMRPMSASFPSPQLLLFSFVGLVSHVQVLGTAPVHPNSTLDVNFYLMVYGGIAGANSVFTILRAFLFAHGTIRAATVIHSRLLQRVMKATVTFFDITPMGRILNRFSSDLYCVDDTLPFILNIFLANVFGLLGMLVMITYGLPWIGLILLPLAVIYFSIQRYYRRTSRELKRLYSLTLSPIYTHFSETLTGLSIIRATRATDRFEAENQLQLEQNQRCLFASNTAMQWLDIRLQMIGVAVVTAIAGIAIIQHQRQLGNPGLVGLALSYALSVTNLLSGLISSFTQTETMMVSVERAEEYSTDIHIEPQEQLVQVAPDWPSRGHVEFQQVVLTYRPGLPNALDGVTFTIYPGEKVGIVGRTGSGKSTIFLALFRMVELRGGRILLDNVDSRSVGLESLRSKLAIIPQDPFLFSGTIRENLDPLGRHTDSDLYQVLEQCRLQAVITQMGGLDCELGERGKSLSVGQRQLVCLARALLTQAKVLCIDEATASVDQKTDQLLQQTIRQRFADKTVLTIAHRLNTILDSDRVLVMHAGKVAELDSPAVLSKREDSLFQRLLTSGQQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJE6</accession>
    <name>A0A452GJE6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">SRCR domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GJE6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002053.1">
        <property type="protein sequence ID" value="ENSGAGP00000001803.1"/>
        <property type="gene ID" value="ENSGAGG00000001441.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045217">
        <property type="term" value="P:cell-cell junction maintenance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000003">
        <property type="entry name" value="Deleted in malignant brain tumors 1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.250.10">
        <property type="entry name" value="SRCR-like domain"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR053243">
        <property type="entry name" value="SJ_maturation_regulator"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001190">
        <property type="entry name" value="SRCR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036772">
        <property type="entry name" value="SRCR-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47653:SF1">
        <property type="entry name" value="DELETED IN MALIGNANT BRAIN TUMORS 1 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47653">
        <property type="entry name" value="PROTEIN BARK BEETLE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00530">
        <property type="entry name" value="SRCR"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00258">
        <property type="entry name" value="SPERACTRCPTR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00202">
        <property type="entry name" value="SR"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56487">
        <property type="entry name" value="SRCR-like"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00420">
        <property type="entry name" value="SRCR_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50287">
        <property type="entry name" value="SRCR_2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3 5" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="4" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <keyword evidence="1 6" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="6">
        <location>
            <begin position="1"/>
            <end position="27"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019302625" description="SRCR domain-containing protein" evidence="6">
        <location>
            <begin position="28"/>
            <end position="346"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="7">
        <location>
            <begin position="22"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="7">
        <location>
            <begin position="126"/>
            <end position="226"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="7">
        <location>
            <begin position="230"/>
            <end position="330"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="47"/>
            <end position="111"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="60"/>
            <end position="121"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="91"/>
            <end position="101"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="151"/>
            <end position="215"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="164"/>
            <end position="225"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="195"/>
            <end position="205"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="255"/>
            <end position="319"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="268"/>
            <end position="329"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="299"/>
            <end position="309"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00196"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50287"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001803.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="346" mass="38373" checksum="0BAEC1F2AB1F51EB" modified="2019-05-08" version="1">LCLLSAFLVYTISAPLSSFISLRLVNGQNRCCGRVEVRYNGNWGTVCDDAWDIKDAEVVCRQLECGRAIAAPMNARFGEGFGRILLDDVQCRGDESSLLQCSHRGWGTHNCAHREDASVICSGPLVRVRNGQNGCSGRVEVRYKGSWGTVCDDAWDINDAEVVCRQLGCGRAIEAPTNARFGKGFGRILLDDVQCRGNEHSLWQCSHRGWGIHNCAHREDASVICSGPLVRVRNGQNGCSGRIEVRYKGSWGTVCDDAWDINDAEVVCRQLGCGRAIEAPTNARFGKGFGRILLDDVQCRGNEHSLWQCLHRGWGIHNCAHRKDASVICSEQRPIRHMVQDHLTYI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJF2</accession>
    <name>A0A452GJF2_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="15">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJF2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002059.1">
        <property type="protein sequence ID" value="ENSGAGP00000001809.1"/>
        <property type="gene ID" value="ENSGAGG00000001446.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd07765">
        <property type="entry name" value="KRAB_A-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000295">
        <property type="entry name" value="zinc finger protein 19"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000003">
        <property type="entry name" value="Zinc finger protein 3 homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001498">
        <property type="entry name" value="Zinc finger protein 404"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000281">
        <property type="entry name" value="Zinc finger protein 558 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000737">
        <property type="entry name" value="Zinc finger protein 565"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001437">
        <property type="entry name" value="Zinc finger protein 594"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001909">
        <property type="entry name" value="KRAB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036051">
        <property type="entry name" value="KRAB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23226">
        <property type="entry name" value="ZINC FINGER AND SCAN DOMAIN-CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23226:SF85">
        <property type="entry name" value="ZINC FINGER PROTEIN 397"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01352">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SMART" id="SM00349">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109640">
        <property type="entry name" value="KRAB domain (Kruppel-associated box)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50805">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="6"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="9" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 11" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="KRAB" evidence="14">
        <location>
            <begin position="11"/>
            <end position="83"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="157"/>
            <end position="184"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="185"/>
            <end position="212"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="213"/>
            <end position="240"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="241"/>
            <end position="268"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="269"/>
            <end position="296"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="297"/>
            <end position="324"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="334"/>
            <end position="363"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="339"/>
            <end position="349"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001809.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="393" mass="45322" checksum="B45DE06C23E82BD8" modified="2019-05-08" version="1">LVGLLLFQGPVTFEEVAVYFSREEGTLLDPTQRALYRDVMQGNYETMVLLAEFLVFQRDMISQLEQGEEPWVPELQGSEEREILRSPRTDEETLNQLRICKCLKETSRMPYKALGKSLSSLLSIAGKMDKLISCQGTQKGLKEARIQQEIGREKQKYTCAECGKNFTYRSRLSQHQRIHTGERPYECHECGKLFTQRSALIRHQRIHKGDRPYECRECGKTFNCSSNLITHQRIHTGDRPYECRECGKTFNCSSHLIRHRRIHIGDRPYECRECGKTFTHSSNLITHQRIHTGNRPYECRECGKTFNRSSHLIRHQKIHRRDRPYECREKHLPNLPVESSEGQSHSSLSPCQPGLSMLPTSHNAGTLPDLPSDLCSPPLLLGSLPHTIQMPCF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJF4</accession>
    <name>A0A452GJF4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">SRCR domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GJF4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002047.1">
        <property type="protein sequence ID" value="ENSGAGP00000001797.1"/>
        <property type="gene ID" value="ENSGAGG00000001441.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045217">
        <property type="term" value="P:cell-cell junction maintenance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000003">
        <property type="entry name" value="Deleted in malignant brain tumors 1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.250.10">
        <property type="entry name" value="SRCR-like domain"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR053243">
        <property type="entry name" value="SJ_maturation_regulator"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001190">
        <property type="entry name" value="SRCR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036772">
        <property type="entry name" value="SRCR-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47653:SF1">
        <property type="entry name" value="DELETED IN MALIGNANT BRAIN TUMORS 1 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47653">
        <property type="entry name" value="PROTEIN BARK BEETLE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00530">
        <property type="entry name" value="SRCR"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00258">
        <property type="entry name" value="SPERACTRCPTR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00202">
        <property type="entry name" value="SR"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56487">
        <property type="entry name" value="SRCR-like"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00420">
        <property type="entry name" value="SRCR_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50287">
        <property type="entry name" value="SRCR_2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3 5" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="4" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <keyword evidence="1 6" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="6">
        <location>
            <begin position="1"/>
            <end position="25"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019334426" description="SRCR domain-containing protein" evidence="6">
        <location>
            <begin position="26"/>
            <end position="336"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="7">
        <location>
            <begin position="20"/>
            <end position="120"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="7">
        <location>
            <begin position="124"/>
            <end position="224"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="7">
        <location>
            <begin position="228"/>
            <end position="328"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="45"/>
            <end position="109"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="58"/>
            <end position="119"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="89"/>
            <end position="99"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="149"/>
            <end position="213"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="162"/>
            <end position="223"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="193"/>
            <end position="203"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="253"/>
            <end position="317"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="266"/>
            <end position="327"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="297"/>
            <end position="307"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00196"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50287"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001797.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="336" mass="36956" checksum="E6F881CF4542F516" modified="2019-05-08" version="1">MALSFLVYTISAPLSSFISLRLVNGQNRCCGRVEVRYNGNWGTVCDDAWDIKDAEVVCRQLECGRAIAAPMNARFGEGFGRILLDDVQCRGDESSLLQCSHRGWGTHNCAHREDASVICSGPLVRVRNGQNGCSGRVEVRYKGSWGTVCDDAWDINDAEVVCRQLGCGRAIEAPTNARFGKGFGRILLDDVQCRGNEHSLWQCSHRGWGIHNCAHREDASVICSGPLVRVRNGQNGCSGRIEVRYKGSWGTVCDDAWDINDAEVVCRQLGCGRAIEAPTNARFGKGFGRILLDDVQCRGNEHSLWQCLHRGWGIHNCAHRKDASVICSVGQKDAVT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJF9</accession>
    <name>A0A452GJF9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Rap guanine nucleotide exchange factor 2</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="21">Cyclic nucleotide ras GEF</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="23">Neural RAP guanine nucleotide exchange protein</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="20">PDZ domain-containing guanine nucleotide exchange factor 1</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="19">RA-GEF-1</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="22">Ras/Rap1-associating GEF-1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="31 32">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="32" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="31" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="31" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell junction</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Perinuclear region</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Late endosome</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the RAPGEF2 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000001965.1">
        <property type="protein sequence ID" value="ENSGAGP00000001729.1"/>
        <property type="gene ID" value="ENSGAGG00000001344.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070161">
        <property type="term" value="C:anchoring junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005770">
        <property type="term" value="C:late endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048471">
        <property type="term" value="C:perinuclear region of cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007264">
        <property type="term" value="P:small GTPase-mediated signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00038">
        <property type="entry name" value="CAP_ED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd06755">
        <property type="entry name" value="PDZ_RapGEF2_RapGEF6-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd01785">
        <property type="entry name" value="RA_PDZ-GEF1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00155">
        <property type="entry name" value="RasGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd06224">
        <property type="entry name" value="REM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.10:FF:000008">
        <property type="entry name" value="Rap guanine nucleotide exchange factor (GEF) 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.840.10:FF:000001">
        <property type="entry name" value="Rap guanine nucleotide exchange factor (GEF) 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000024">
        <property type="entry name" value="rap guanine nucleotide exchange factor 2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.870.10:FF:000001">
        <property type="entry name" value="rap guanine nucleotide exchange factor 2 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.42.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.10">
        <property type="entry name" value="Jelly Rolls"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.840.10">
        <property type="entry name" value="Ras guanine-nucleotide exchange factors catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.870.10">
        <property type="entry name" value="Son of sevenless (SoS) protein Chain: S domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000595">
        <property type="entry name" value="cNMP-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018490">
        <property type="entry name" value="cNMP-bd_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001478">
        <property type="entry name" value="PDZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036034">
        <property type="entry name" value="PDZ_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000159">
        <property type="entry name" value="RA_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000651">
        <property type="entry name" value="Ras-like_Gua-exchang_fac_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023578">
        <property type="entry name" value="Ras_GEF_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001895">
        <property type="entry name" value="RASGEF_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036964">
        <property type="entry name" value="RASGEF_cat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014710">
        <property type="entry name" value="RmlC-like_jellyroll"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029071">
        <property type="entry name" value="Ubiquitin-like_domsf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45161">
        <property type="entry name" value="CYTOSKELETON-ASSOCIATED PROTEIN 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45161:SF2">
        <property type="entry name" value="RAP GUANINE NUCLEOTIDE EXCHANGE FACTOR 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00595">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00788">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00617">
        <property type="entry name" value="RasGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00618">
        <property type="entry name" value="RasGEF_N"/>
        <property type="match status" value="1"/>
    </dbReference>
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    <keyword evidence="9" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="10" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="14" id="KW-0221">Differentiation</keyword>
    <keyword evidence="13" id="KW-0967">Endosome</keyword>
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    <keyword evidence="32" id="KW-1185">Reference proteome</keyword>
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        <location>
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    <sequence length="1453" mass="162104" checksum="3A063A06D30255E3" modified="2019-05-08" version="1">MKPLAIPANHGVMGQQEKHSLPADFTKLHLTDSLHPQVTHVTSSHSGCSITSDSGSSSLSDIYQATESEAGDMDLSGLPETAVDSEDDDDEEDIERASDPLMSRDIVRDCLEKDPIDRTDDDIEQLLEFMHQLPAFANMTMSVRRELCAVMVFAVVERAGTIVLNDGEELDSWSVILNGSVEVTYPDGRTEILCMGNSFGVSPTMDKEYMKGVMRTKVDDCQFVCIAQQDYCRILNQVEKNMQKVEEEGEIVMVKEHRELDRTGTRKGHIVIKGTSERLTMHLVEEHSVVDPTFIEDFLLTYRTFLSSPMEVGKKLLEWFNDPSLRDKVTRVVLLWVNNHFNDFEGDPAMTRFLEEFENNLEREKMGGHLRLLNIACAAKAKRRLITLTKPSREAPLPFILLGGSEKGFGIFVDSVDLGSKATEAGLKRGDQILEVNGQNFENIQLSKAMEILRNNTHLSITVKTNLFVFKELLTRLSEEKRNGAPHLPKIGDIKKASRYSIPDLAVDVEQVIGLEKVNKKSKANTVGGRNKLKKILDKTRISILPQKPYNDIGIGQSQDDSIVGLRQTKHIPPALPVSGTLSSSNPDLLQSHHRILDFNSTPDLPDQVLRVFKADQQSRYIMISKDTTAKEVVIQAIREFAVTATPDAYSLCEVSVTPEGVIKQRRLPDQLSKLADRIQLSGRYYLKNNMETETLCSDEDAQELLRESQISLLQLSTVEMAAQLSMRNFELFRNIEPTEYIDDLFKLKSKTSCANLKKFEEVINQETFWVASEILRETNQLKRMKIIKHFIKIALHCRECKNFNSMFAIISGLNLAPVARLRTTWEKLPSKYEKLFQDLQDLFDPSRNMAKYRNVLNSQNLQPPIIPLFPVIKKDLTFLHEGNDSKVEGLVNFEKLRMIAKEIRHVGRMASVNMDPAVMFRTRKKKWRSLGSLSQSSTNAAVLDVAQTGGHKKRVRRSSFLNAKKLYEDAQMARKVKQYLSNLDLEMDEESLQTLSLQCEPATNTLPKNPGDKRSGKQEMSPVAPRAGIQQKVQQQHKINQALQVPAVSLYPSRKKVPVKDLPPFGINSPQALKKILSLSEEGSLDRHKKQAEDTISNASSQLSSPPTSPHTSPRKGYTLAPSGTVDNFSDSGRSEISSRSSIVSNSSFDSMPVSLHDERRQRHSVSIVETNLGVGRIDRRAMIEPDQYSLGSYAPLSENRGLYAGATVLSSPSTEELSQDQGDRTSLDAADSGRGSWTSCSSGSHDNIQTIQHQRSWETLPFGHVHFDSSGDGAGLWASGSHMDQMMFPDHGTKYGRQSQGREGLDQAQSRASWASSTGYWGEDSEGDTGTIKRRGGKDVSIEAETSSITSVTAEESKPAPMSSHITVSSSSAKGLIARKEGRYREPPPTPPGYVGIPIADFAEGVSHPTRKPPDYNVALQRSRMVARTCETHGTSTLQQQSQEDEQVSAV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJG0</accession>
    <name>A0A452GJG0_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="7">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
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        <property type="gene ID" value="ENSGAGG00000001447.1"/>
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        <property type="evidence" value="ECO:0007669"/>
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        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="Ig-like" evidence="5">
        <location>
            <begin position="46"/>
            <end position="141"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="5">
        <location>
            <begin position="177"/>
            <end position="255"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="6">
        <location>
            <begin position="269"/>
            <end position="364"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="6">
        <location>
            <begin position="397"/>
            <end position="495"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="6">
        <location>
            <begin position="498"/>
            <end position="591"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="6">
        <location>
            <begin position="597"/>
            <end position="692"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="6">
        <location>
            <begin position="699"/>
            <end position="800"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="5">
        <location>
            <begin position="1019"/>
            <end position="1104"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="5">
        <location>
            <begin position="1232"/>
            <end position="1321"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001820.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1335" mass="151124" checksum="27B15540B24B5EEE" modified="2019-05-08" version="1">MEKLEKGVIRNCRLLRVKADRKAFRKKALEKARREKEYIELLSSRPPLIWIPLRVHAVWERMGVKLTCTILASPPPQVTWYKNGVRIDPRLAPAGKYRIQNKFGMLTLDINRCSLEDSAEYSVEVKNPYGEAYSFATVLVQKYYGKESGFDSEIYKRSLIGQEADFAFSLKPLFSREKEPFTLSCFFSSDLLQHQRDITWFRDGEFLKDSDRQQIKCLDREASLTVLGAYKEDEGFYTVRVPTLDGYNEQSAYVFVRDAVAATAGAPGSPLNVKCHDINKDCLLLSWVAPSDSGGSPIFGYFIERCEVGTDEWVPCNKMPVKTCRCPVLCLTEGKTYQFRVKAVNHSGISNPSKASDPVTMKDPNEEKRVMTIPYDDGKKITISKDELEGKIKIPLPPTNVHASEIRENYVVLSWDEPDPRGKEPLSYYVEKSIAGSNSWQNVNLEMAVNSPRFALFDLVKGKSYCFRVRSANKYGVSEPSLPSEPITAGTTLAPLPPPSQVLAFRDSKTSVVVHWDKLKEGQEPLGYYIYCREVGTDEWQTVNNKPVTCNKFTVPGLKPGKEYVFCVKSVNEAGVGDSSPESDPIIVRPAISRPSAPRDFALLSCGRNDMTIGWKAPKHKGGGKILGYFLDQHDAAELGWHEVNIRPIAQRVYKVTNLNEGHFYEFRACATNMAGVGKVSEPSDLFKCEEWTMPQPGPPYDVRFTEVRDANLMLHXMLHWEAPLYTGTGPVTGYYIDVCEEGSEEWKQLNKQPTASTHMRVSDLETGKCYIFRVRALNKAGIGPPSIPSDPVVVETKPGTNEIELGVDKEGFVYMAFETPDMCDSSEFIWSKDYEGPPDPNRVKIEDKGNKSKIILKNASEKDLGTYSVEVTDVDDEISASHTLSQEELDKLRKISHEIRNPLIELISGWNIDVLEKGEVRLWLEVEKLSPEAELHLIFNGKELSSTPTHKINFDRANGLIELIIQDFCDDDKGTYTAQLQDGKAKNQFTLALVDDSFDKVEAEAAAKRRDWRRKQGPHFIENLQWKVTEDCEVLLTCKATNMKKDTNFKWFLDKKARLDGLYDPQTGAGTLRIKKLTKEDKGIYKAVVSDDRGEDITELDLSKEGFEDILKELCRISALSAKPLKVQPTAEGIRIYSEVKYYTDYMKTTWYHKEKRLESGERLKSGSTMDQIWLHILNPTDSDKGKYTLEIFDGKDSHRLSTDLSGKAFEEAMAEYLRLKEAVIVEKNRARVVQGLPDVATIMEDKTLCLTCCISGDPYPEITWFKNEKVVVFKDRYKMDVKGIVVTITIEKVCSDDTGKYSIYVKNKYGSETGQVTISVYKHGEEPIELREK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJG2</accession>
    <name>A0A452GJG2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Protein phosphatase 1 regulatory subunit 21</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="7">Coiled-coil domain-containing protein 128</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="8">Ferry endosomal RAB5 effector complex subunit 2</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="6">KLRAQ motif-containing protein 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Early endosome</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJG2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002070.1">
        <property type="protein sequence ID" value="ENSGAGP00000001819.1"/>
        <property type="gene ID" value="ENSGAGG00000001451.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005769">
        <property type="term" value="C:early endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040024">
        <property type="entry name" value="PPP1R21"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049372">
        <property type="entry name" value="PPP1R21_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019343">
        <property type="entry name" value="PPP1R21_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019348">
        <property type="entry name" value="PPP1R21_six_helix"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21448:SF0">
        <property type="entry name" value="PROTEIN PHOSPHATASE 1 REGULATORY SUBUNIT 21"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21448">
        <property type="entry name" value="SMOOTH MUSCLE MYOSIN HEAVY CHAIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10205">
        <property type="entry name" value="KLRAQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21636">
        <property type="entry name" value="PPP1R21_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10212">
        <property type="entry name" value="PPP1R21_helical"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01254">
        <property type="entry name" value="KLRAQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5 9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0967">Endosome</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0694">RNA-binding</keyword>
    <feature type="domain" description="Protein phosphatase 1 regulatory subunit 21 N-terminal" evidence="11">
        <location>
            <begin position="1"/>
            <end position="66"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="38"/>
            <end position="58"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="91"/>
            <end position="157"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="512"/>
            <end position="557"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="649"/>
            <end position="683"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004412"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00020102"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00031361"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00031617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00044824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="SMART" id="SM01254"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001819.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="734" mass="83798" checksum="82578DDCF8C53F1D" modified="2019-05-08" version="1">MKDQSLRKLQQEMDSLTFRNQQLAKRVELLQDELALSEAKGKKNKKSGDSSSQLSQEQKSVFDEDLQKKIEENERLHIQFFEADERHKRLEAELKNRLEVLETDAAQHQAVVDSLTRKYIDTIEKLQNDKTNLEIKSQTLEREAKECRFRAEECQQQLKNLHADLGRRLDDSLCIINEKVPFNDTRSIQYNALNVPLHNRRCQLKLRDIAGQALAFVQDLVTALLNFHTYMEQRVQIFPIDSATDTISPLNQKFSQYLHENASYVRPLEEGMLHLFESITEDTVTVLETTVKLKVFSEHLTSYTCFLRKILPYQLKSLEEECESSLCTAALRARNLELHNDMKKITAVFEKLHVYISLIVLPSIRPEGLLRTNYSSVFTHIAAGLHGFHNVMKEVSKHYSQKAALEHELPTATQKLITTNDCILSSVVALTNGAGKIASFFSNNLDHFMASLSYGPKGGTEFISPLSAECMLQYKKKAVAYMKSLKKPYSDSVPYEEALANRRVLLSSTESREGLAQQVQQSLEKIAKLEQEKEHWMLEAQLAKIKLEKENQKLKSSLSGQLIEPMPENSVLLNAAEQEKEVSIKKTSGEPVKKMSLIGMLTMTTGSKEAPEAKSREDLIKNHYMARIVELTSLLQLADSKSVHFHAECRALAKRLSLAEKSKESLVEEMKLASQSISRLQDELTTTKRSYEDQLSMMSDHLCSMNETLSKQREEIDTLKMSKVGYQVLYLLWA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJG3</accession>
    <name>A0A452GJG3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Tight junction protein 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cell junction</location>
            <location evidence="3">Tight junction</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Peripheral membrane protein</topology>
            <orientation evidence="2">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the MAGUK family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJG3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002057.1">
        <property type="protein sequence ID" value="ENSGAGP00000001807.1"/>
        <property type="gene ID" value="ENSGAGG00000001448.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005923">
        <property type="term" value="C:bicellular tight junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050839">
        <property type="term" value="F:cell adhesion molecule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098609">
        <property type="term" value="P:cell-cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045216">
        <property type="term" value="P:cell-cell junction organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090557">
        <property type="term" value="P:establishment of endothelial intestinal barrier"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1905605">
        <property type="term" value="P:positive regulation of blood-brain barrier permeability"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0150105">
        <property type="term" value="P:protein localization to cell-cell junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd06727">
        <property type="entry name" value="PDZ1_ZO1-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd06728">
        <property type="entry name" value="PDZ2_ZO1-like_ds"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd06729">
        <property type="entry name" value="PDZ3_ZO1-like_domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12027">
        <property type="entry name" value="SH3_ZO-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000009">
        <property type="entry name" value="Putative tight junction protein ZO-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000013">
        <property type="entry name" value="Putative tight junction protein ZO-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000110">
        <property type="entry name" value="tight junction protein ZO-1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000075">
        <property type="entry name" value="Tight junction protein ZO-2 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.42.10">
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008145">
        <property type="entry name" value="GK/Ca_channel_bsu"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008144">
        <property type="entry name" value="Guanylate_kin-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
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        <property type="entry name" value="ZO"/>
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        <property type="entry name" value="ZO-2"/>
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        <property type="entry name" value="ZO-2_SH3"/>
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        <property type="entry name" value="TIGHT JUNCTION PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="TIGHT JUNCTION PROTEIN ZO-2"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Guanylate_kin"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="ZONOCCLUDNS"/>
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        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
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    <keyword evidence="7" id="KW-1003">Cell membrane</keyword>
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    <keyword evidence="9" id="KW-0677">Repeat</keyword>
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        <location>
            <begin position="10"/>
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            <begin position="500"/>
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        <location>
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            <end position="660"/>
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            <begin position="765"/>
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            <begin position="913"/>
            <end position="1073"/>
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            <begin position="1109"/>
            <end position="1182"/>
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    <feature type="compositionally biased region" description="Polar residues" evidence="14">
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            <begin position="147"/>
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            <begin position="1051"/>
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        <source>
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    <sequence length="1182" mass="133422" checksum="D480DE59304CFE1A" modified="2019-05-08" version="1">MEELIWEQYTVTLQKDSKRGFGIAVSGGRDNPHFENGETSIVISDVLPGGPADGLLQENDRVVMVNGTPMENVPHSFAVQQLRKSGKVAAIVVKRPRKVQLTALRRSPSLDHDDRAFDILDDPAEFDGRSARSGYSDRSWHSGHGGRSQSWGNSPDRSYRRDQDRGRNYSRDHSRERSYSRDGSRGRSIDRERSLDRERRRDRSRGRSINRERSLDREHRRDRSRGRSIDRDDGYERGAGDYSPPRDYSYRHQPDPRYDKEARSHSRDRLNSHSPLLEPRRQHGYPGQQYQDRPISVLLTKSKPNEEYGLRLGSQIFIKEMTSTGLATKDGNLYEGDIILKINGTVTENMSLADARKLIEKSRGKLQLVVLRDRKQTLLNIPSLHDSDSEIEDISEIESNRSSSPQDDQKLHHSDFDSHSSNEKLKEKPNTKEDPPNRLSRMGAMPTPFKSTSDIAAAVIVTDTNKEPKYKDDLPVSQPKVAPRTFLRPSPEDEAIYGPNTKMVKFKKGDSVGLRLAGGNDVGIFVAGIQEGTSADQEGLQEGDQILKVNTQDFRGIVREDAVLYLLEIPKGETVTILAQSKYEVYRDIMACGRGDSFFIRSHFECEKESPQSLAFTRGEIFRVVDTLYDGKLGNWLAVRIGNELEKGIIPNKSRAEQMASVQNAQRDGSSDRADFWRMRGQRSGMKKNLKKSREDLTAIASVSTKFPAYERVLLREAGFKRPVVIFGPIADLAMEKLSNDLPDLYQTAKTEPKDAGSEKSTGVVRLNTVRQIIEQNKHALLDVTPKAVDLLNYTQWFPIVVFFNPDNKQGVKTMRQRLSPTSNKSSRKLYDQANKLKKTCSYLFTATINLNSANDSWYGSLKDTIQQQQVEAVWVSEGKMEGMDDDMEDRMSYLTAMGADYLSCDSRLISDLEDTDGEGGAYTDNELDEPSDEPSVSSISRSSEPVQLEESIRKPSPEPKAQMRRVGSREILRDPSPPPAFKPEPPKGKLQNKEDVYDFPKNYESKPNSSSTVSNETSAVLSKAAAPPVSVKPAFGRPILKTSQPAVPAAEEEEEEDDVDGGESAPKSVLGKVKIFEKMDHKARLQRIQELQEAQNARLEIAQKHPDIYAVPIKTQKPDQNWPQPMSSRPPEPQKPPTRPYLENRGSYGSDAEEEDYRRQLSDNSKRGYYGQPSRYRDTEL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJG4</accession>
    <name>A0A452GJG4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">RNA binding motif protein 4B</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
            <location evidence="1">Nucleolus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJG4"/>
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        <property type="protein sequence ID" value="ENSGAGP00000001734.1"/>
        <property type="gene ID" value="ENSGAGG00000001387.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
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        <property type="term" value="C:nucleolus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003729">
        <property type="term" value="F:mRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006397">
        <property type="term" value="P:mRNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008380">
        <property type="term" value="P:RNA splicing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
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        <property type="entry name" value="RRM1_RBM4"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="RRM2_RBM4"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="RNA-binding motif protein 4"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="RNA-binding protein 4 isoform X1"/>
        <property type="match status" value="1"/>
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        <property type="match status" value="2"/>
    </dbReference>
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        <property type="entry name" value="Zinc finger, CCHC-type"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Euk_RNA-bind_prot"/>
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        <property type="entry name" value="Nucleotide-bd_a/b_plait_sf"/>
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        <property type="entry name" value="RBD_domain_sf"/>
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        <property type="entry name" value="RBM4_RRM1"/>
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        <property type="entry name" value="RBM4_RRM2"/>
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        <property type="entry name" value="RRM_dom"/>
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        <property type="entry name" value="Znf_CCHC"/>
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        <property type="entry name" value="HETEROGENEOUS NUCLEAR RIBONUCLEOPROTEIN M-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="OS02G0815200 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="RRM_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00098">
        <property type="entry name" value="zf-CCHC"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="RRM"/>
        <property type="match status" value="2"/>
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        <property type="entry name" value="ZnF_C2HC"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="RNA-binding domain, RBD"/>
        <property type="match status" value="2"/>
    </dbReference>
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        <property type="entry name" value="RRM"/>
        <property type="match status" value="2"/>
    </dbReference>
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        <property type="entry name" value="ZF_CCHC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8" id="KW-0010">Activator</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="2" id="KW-0507">mRNA processing</keyword>
    <keyword evidence="9" id="KW-0508">mRNA splicing</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="7 12" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 11" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="RRM" evidence="13">
        <location>
            <begin position="2"/>
            <end position="72"/>
        </location>
    </feature>
    <feature type="domain" description="RRM" evidence="13">
        <location>
            <begin position="78"/>
            <end position="148"/>
        </location>
    </feature>
    <feature type="domain" description="CCHC-type" evidence="14">
        <location>
            <begin position="162"/>
            <end position="176"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004604"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022664"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023187"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00047"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00176"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50102"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50158"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001734.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="17">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="367" mass="40988" checksum="C2F8BAF7E31F1739" modified="2019-05-08" version="1">MVKLFIGNLPREATEQEIRSLFEQYGKVLECDIIKNYGFVHIEDKTAAEDAIRNLHHHKLHGVCINVEASKNKSKASTKLHVGNISTTCTNLELRAKFEEYGPVIECDIVKDYAFVHMERAEDAVEAIRGLDNTEFQGKRMRVQLSTSRLRTIPGMGDKSGCYRCGKEGHWSKECPVDRTGQVPDFTETYNEQYGAVRTPYPAGYGETMYYDDGYGAMVDYYKRYRVRPYATASAYDAYAEQTMAQYSQYAQYSQVQTTAMAATTAMANRLTATLDPYDRALLPTPGAAAAVAAAATAAAAAASSTYYTRDRSPLRRTATAATTIGEAYAYERCQLSPVSSVARASLYDVQRFERESYADRARYSAF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJG5</accession>
    <name>A0A452GJG5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Cholinergic receptor nicotinic alpha 10 subunit</fullName>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="14">Synaptic cell membrane</location>
            <topology evidence="14">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="15">Belongs to the ligand-gated ion channel (TC 1.A.9) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJG5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002077.1">
        <property type="protein sequence ID" value="ENSGAGP00000001825.1"/>
        <property type="gene ID" value="ENSGAGG00000001456.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0022848">
        <property type="term" value="F:acetylcholine-gated monoatomic cation-selective channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004888">
        <property type="term" value="F:transmembrane signaling receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd19051">
        <property type="entry name" value="LGIC_TM_cation"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.390:FF:000009">
        <property type="entry name" value="Cholinergic receptor nicotinic alpha 9 subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.390:FF:000053">
        <property type="entry name" value="Neuronal acetylcholine receptor subunit alpha-9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.70.170.10:FF:000010">
        <property type="entry name" value="neuronal acetylcholine receptor subunit alpha-9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.70.170.10">
        <property type="entry name" value="Neurotransmitter-gated ion-channel ligand-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.390">
        <property type="entry name" value="Neurotransmitter-gated ion-channel transmembrane domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006202">
        <property type="entry name" value="Neur_chan_lig-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036734">
        <property type="entry name" value="Neur_chan_lig-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006201">
        <property type="entry name" value="Neur_channel"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036719">
        <property type="entry name" value="Neuro-gated_channel_TM_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038050">
        <property type="entry name" value="Neuro_actylchol_rec"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006029">
        <property type="entry name" value="Neurotrans-gated_channel_TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018000">
        <property type="entry name" value="Neurotransmitter_ion_chnl_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002394">
        <property type="entry name" value="Nicotinic_acetylcholine_rcpt"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00860">
        <property type="entry name" value="LIC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18945">
        <property type="entry name" value="NEUROTRANSMITTER GATED ION CHANNEL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02931">
        <property type="entry name" value="Neur_chan_LBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02932">
        <property type="entry name" value="Neur_chan_memb"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00254">
        <property type="entry name" value="NICOTINICR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00252">
        <property type="entry name" value="NRIONCHANNEL"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90112">
        <property type="entry name" value="Neurotransmitter-gated ion-channel transmembrane pore"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF63712">
        <property type="entry name" value="Nicotinic receptor ligand binding domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00236">
        <property type="entry name" value="NEUROTR_ION_CHANNEL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="9" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13 15" id="KW-0407">Ion channel</keyword>
    <keyword evidence="7 15" id="KW-0406">Ion transport</keyword>
    <keyword evidence="12" id="KW-1071">Ligand-gated ion channel</keyword>
    <keyword evidence="8 15" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0675">Receptor</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 15" id="KW-0732">Signal</keyword>
    <keyword evidence="6" id="KW-0770">Synapse</keyword>
    <keyword evidence="3 15" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 15" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="1 15" id="KW-0813">Transport</keyword>
    <feature type="signal peptide" evidence="15">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5022253725" description="Cholinergic receptor nicotinic alpha 10 subunit" evidence="15">
        <location>
            <begin position="21"/>
            <end position="464"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="233"/>
            <end position="257"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="263"/>
            <end position="284"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="296"/>
            <end position="318"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="443"/>
            <end position="461"/>
        </location>
    </feature>
    <feature type="domain" description="Neurotransmitter-gated ion-channel ligand-binding" evidence="16">
        <location>
            <begin position="26"/>
            <end position="230"/>
        </location>
    </feature>
    <feature type="domain" description="Neurotransmitter-gated ion-channel transmembrane" evidence="17">
        <location>
            <begin position="238"/>
            <end position="459"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00034099"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="RuleBase" id="RU000687"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF02931"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF02932"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001825.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="20">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="464" mass="52924" checksum="B1CD7FE02FFBD33F" modified="2019-05-08" version="1">MGLVLCRVSFLSIYLGCSGAQGKFAYKLLHDLFANYSNALRPVEDTDRALNVTLQITLSQIIDMDERNQILTAYLWIRQVWLDAYLAWDKDDYDGIDTIRIPSSYVWRPDIVLYNNADDQFTESMETNVVIRYDGQVMWDSPAITKSSCKVDVSYFPFDGQQCRLTFGSWTYNGNQIDLLNGLDTGDLTDFVENVEWEVLGMPAKRNVIIYGCCSEPYPDITYTLLLKRRASFYIFNLLLPCIMISFLAPLGFYLPADSGEKVSLGVTVLLALTVFQLLVAESMPPSENVPLIGKYYIATMTMITASTALTIFIMNIHHCGPGAKPVPKWAKKFILQYMARLFFVYEVGESCKSPKRQLGHLPTVQVVSGRAAKEEQKTGTRMKLECGWEEQQHQVCVKTPCLCHHNSLLRNVEYIANCFRDQKAAQKQSGEWKKVAKVMDRIFMWIFFIMVFFMSMLIMGKAI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJG6</accession>
    <name>A0A452GJG6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Cholinergic receptor nicotinic alpha 10 subunit</fullName>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="14">Synaptic cell membrane</location>
            <topology evidence="14">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="15">Belongs to the ligand-gated ion channel (TC 1.A.9) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJG6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002075.1">
        <property type="protein sequence ID" value="ENSGAGP00000001824.1"/>
        <property type="gene ID" value="ENSGAGG00000001456.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0022848">
        <property type="term" value="F:acetylcholine-gated monoatomic cation-selective channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004888">
        <property type="term" value="F:transmembrane signaling receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd19051">
        <property type="entry name" value="LGIC_TM_cation"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.390:FF:000009">
        <property type="entry name" value="Cholinergic receptor nicotinic alpha 9 subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.390:FF:000053">
        <property type="entry name" value="Neuronal acetylcholine receptor subunit alpha-9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.70.170.10:FF:000010">
        <property type="entry name" value="neuronal acetylcholine receptor subunit alpha-9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.70.170.10">
        <property type="entry name" value="Neurotransmitter-gated ion-channel ligand-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.390">
        <property type="entry name" value="Neurotransmitter-gated ion-channel transmembrane domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006202">
        <property type="entry name" value="Neur_chan_lig-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036734">
        <property type="entry name" value="Neur_chan_lig-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006201">
        <property type="entry name" value="Neur_channel"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036719">
        <property type="entry name" value="Neuro-gated_channel_TM_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038050">
        <property type="entry name" value="Neuro_actylchol_rec"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006029">
        <property type="entry name" value="Neurotrans-gated_channel_TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018000">
        <property type="entry name" value="Neurotransmitter_ion_chnl_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002394">
        <property type="entry name" value="Nicotinic_acetylcholine_rcpt"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00860">
        <property type="entry name" value="LIC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18945">
        <property type="entry name" value="NEUROTRANSMITTER GATED ION CHANNEL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02931">
        <property type="entry name" value="Neur_chan_LBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02932">
        <property type="entry name" value="Neur_chan_memb"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00254">
        <property type="entry name" value="NICOTINICR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00252">
        <property type="entry name" value="NRIONCHANNEL"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90112">
        <property type="entry name" value="Neurotransmitter-gated ion-channel transmembrane pore"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF63712">
        <property type="entry name" value="Nicotinic receptor ligand binding domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00236">
        <property type="entry name" value="NEUROTR_ION_CHANNEL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="9" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13 15" id="KW-0407">Ion channel</keyword>
    <keyword evidence="7 15" id="KW-0406">Ion transport</keyword>
    <keyword evidence="12" id="KW-1071">Ligand-gated ion channel</keyword>
    <keyword evidence="8 15" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0675">Receptor</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="6" id="KW-0770">Synapse</keyword>
    <keyword evidence="3 15" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 15" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="1 15" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="243"/>
            <end position="267"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="273"/>
            <end position="294"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="306"/>
            <end position="328"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="428"/>
            <end position="446"/>
        </location>
    </feature>
    <feature type="domain" description="Neurotransmitter-gated ion-channel ligand-binding" evidence="16">
        <location>
            <begin position="36"/>
            <end position="240"/>
        </location>
    </feature>
    <feature type="domain" description="Neurotransmitter-gated ion-channel transmembrane" evidence="17">
        <location>
            <begin position="248"/>
            <end position="373"/>
        </location>
    </feature>
    <feature type="domain" description="Neurotransmitter-gated ion-channel transmembrane" evidence="17">
        <location>
            <begin position="378"/>
            <end position="444"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023286"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00034099"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="RuleBase" id="RU000687"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF02931"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF02932"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001824.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="20">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="449" mass="51176" checksum="7F1540CF568FC7B7" modified="2019-05-08" version="1">IDLRPHYPKLNPAKLILALLPAPGRRCSGAQGKFAYKLLHDLFANYSNALRPVEDTDRALNVTLQITLSQIIDMDERNQILTAYLWIRQVWLDAYLAWDKDDYDGIDTIRIPSSYVWRPDIVLYNNADDQFTESMETNVVIRYDGQVMWDSPAITKSSCKVDVSYFPFDGQQCRLTFGSWTYNGNQIDLLNGLDTGDLTDFVENVEWEVLGMPAKRNVIIYGCCSEPYPDITYTLLLKRRASFYIFNLLLPCIMISFLAPLGFYLPADSGEKVSLGVTVLLALTVFQLLVAESMPPSENVPLIGKYYIATMTMITASTALTIFIMNIHHCGPGAKPVPKWAKKFILQYMARLFFVYEVGESCKSPKRSASGQQDWKETGTLCVKTPCLCHHNSLLRNVEYIANCFRDQKAAQKQSGEWKKVAKVMDRIFMWIFFIMVFFMSMLIMGKAI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJJ7</accession>
    <name>A0A452GJJ7_9SAUR</name>
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        <recommendedName>
            <fullName evidence="3">Ecto-NOX disulfide-thiol exchanger 1/2 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
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            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GJJ7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000002102.1">
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        <property type="gene ID" value="ENSGAGG00000001433.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
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        <property type="term" value="F:NADH dehydrogenase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003756">
        <property type="term" value="F:protein disulfide isomerase activity"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007624">
        <property type="term" value="P:ultradian rhythm"/>
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        <property type="project" value="InterPro"/>
    </dbReference>
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    </dbReference>
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        <property type="entry name" value="ENOX1/2_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16001">
        <property type="entry name" value="ECTO-NOX DISULFIDE-THIOL EXCHANGER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16001:SF6">
        <property type="entry name" value="ECTO-NOX DISULFIDE-THIOL EXCHANGER 1"/>
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    </dbReference>
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    </dbReference>
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        <location>
            <begin position="174"/>
            <end position="270"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="145"/>
            <end position="164"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="56"/>
            <end position="83"/>
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    </feature>
    <feature type="coiled-coil region" evidence="1">
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            <begin position="175"/>
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        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="2">
        <location>
            <begin position="148"/>
            <end position="157"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001849.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="390" mass="45124" checksum="E69143D54F9712E3" modified="2019-05-08" version="1">APSPVEPETPVCSNKDPGSELLAPNLKFSEAITVLLTWIERGEVNRRSANQFYSMVQSANSHVRRLMNEKATHEQEMEQAKESFKNALTGILTQFEQIVAVFNASTRQKAWDHFSKAQRKNIDIWRKHSEELRNAHSEQLMGIRREEEMEMSDDDSSDNPSKKLRVDESALTAQAYALKEENDSLRWQLDAYRNEVELLKQEKGQLFRTEESLTKDQQLKFLQQTMQGMQQQLLSIQEELNNKKSELQQAKEEQTHTEARLKVLQEQLKSAKELAEINGQDESQEINMLTVALVNQEREKNSEKRSQGLKSEKEALLIGIISTFLHVHPFGANIEYLWSYMQQLDSRISANEIETLLMRLPRMFKQEFTGVGATLEKRWKFCAFEGIKTT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMC0</accession>
    <name>A0A452HMC0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Transmembrane protein 45B</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the TMEM45 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMC0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018378.1">
        <property type="protein sequence ID" value="ENSGAGP00000016092.1"/>
        <property type="gene ID" value="ENSGAGG00000012019.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006904">
        <property type="entry name" value="DUF716"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042127">
        <property type="entry name" value="TMEM45"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16007">
        <property type="entry name" value="EPIDIDYMAL MEMBRANE PROTEIN E9-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16007:SF21">
        <property type="entry name" value="TRANSMEMBRANE PROTEIN 45A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04819">
        <property type="entry name" value="DUF716"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 6" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 6" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 6" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="6"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="44"/>
            <end position="61"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="81"/>
            <end position="102"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="114"/>
            <end position="132"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="144"/>
            <end position="162"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="174"/>
            <end position="195"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="215"/>
            <end position="237"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006948"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016092.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="9">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="250" mass="28470" checksum="1DAE1689E11BABC4" modified="2019-05-08" version="1">MSSFKGNALRGTFFLIFGLGWSVKYPLKYLSRKLNADNQARKNMICICLYCLGMLAEQFVPDGPHMGLYSGGTYSWEKLTNWQYTTMYLFYGLSGIVDVLSYSPLKLSLGLDRLLLSVAMFTEGFLFCFHDYNNTALDQHLHSLLLIAIFGGALCALMEVFLRDHTILEFFRTSFFILQGSWFWQIGFVLSPPWGGPGWDQTDRGNFNFLTMCFFWHYAAALLVTAANSVFSYWYVVGQGNNFGALKNAF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMC1</accession>
    <name>A0A452HMC1_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="9">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HMC1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018381.1">
        <property type="protein sequence ID" value="ENSGAGP00000016095.1"/>
        <property type="gene ID" value="ENSGAGG00000012086.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004867">
        <property type="term" value="F:serine-type endopeptidase inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.22.10:FF:000002">
        <property type="entry name" value="Cysteine rich transmembrane BMP regulator 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.40.20:FF:000003">
        <property type="entry name" value="cysteine-rich motor neuron 1 protein isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
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    <dbReference type="Gene3D" id="6.20.200.20">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.22.10">
        <property type="entry name" value="Antistasin, domain 1"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.70.10">
        <property type="entry name" value="Complement Module, domain 1"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Antistasin-like"/>
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        <property type="entry name" value="CRIM1"/>
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    <dbReference type="InterPro" id="IPR045813">
        <property type="entry name" value="CRIM1_C"/>
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    <dbReference type="InterPro" id="IPR009030">
        <property type="entry name" value="Growth_fac_rcpt_cys_sf"/>
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    <dbReference type="InterPro" id="IPR011061">
        <property type="entry name" value="Hirudin/antistatin"/>
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    <dbReference type="InterPro" id="IPR000867">
        <property type="entry name" value="IGFBP-like"/>
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    <dbReference type="InterPro" id="IPR001007">
        <property type="entry name" value="VWF_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46439">
        <property type="entry name" value="CYSTEINE-RICH MOTOR NEURON 1 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46439:SF1">
        <property type="entry name" value="CYSTEINE-RICH MOTOR NEURON 1 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02822">
        <property type="entry name" value="Antistasin"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19442">
        <property type="entry name" value="CRIM1_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00219">
        <property type="entry name" value="IGFBP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00093">
        <property type="entry name" value="VWC"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23334">
        <property type="entry name" value="VWC2L_2nd"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00121">
        <property type="entry name" value="IB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00214">
        <property type="entry name" value="VWC"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SMART" id="SM00215">
        <property type="entry name" value="VWC_out"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57603">
        <property type="entry name" value="FnI-like domain"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57184">
        <property type="entry name" value="Growth factor receptor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57262">
        <property type="entry name" value="Leech antihemostatic proteins"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51252">
        <property type="entry name" value="ANTISTASIN"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51323">
        <property type="entry name" value="IGFBP_N_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01208">
        <property type="entry name" value="VWFC_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50184">
        <property type="entry name" value="VWFC_2"/>
        <property type="match status" value="6"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="4" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <keyword evidence="1 5" id="KW-0732">Signal</keyword>
    <keyword evidence="4" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="5">
        <location>
            <begin position="1"/>
            <end position="37"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019425068" evidence="5">
        <location>
            <begin position="38"/>
            <end position="956"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="4">
        <location>
            <begin position="861"/>
            <end position="881"/>
        </location>
    </feature>
    <feature type="domain" description="IGFBP N-terminal" evidence="8">
        <location>
            <begin position="36"/>
            <end position="115"/>
        </location>
    </feature>
    <feature type="domain" description="VWFC" evidence="6">
        <location>
            <begin position="255"/>
            <end position="312"/>
        </location>
    </feature>
    <feature type="domain" description="VWFC" evidence="6">
        <location>
            <begin position="322"/>
            <end position="378"/>
        </location>
    </feature>
    <feature type="domain" description="Antistasin-like" evidence="7">
        <location>
            <begin position="390"/>
            <end position="419"/>
        </location>
    </feature>
    <feature type="domain" description="Antistasin-like" evidence="7">
        <location>
            <begin position="426"/>
            <end position="453"/>
        </location>
    </feature>
    <feature type="domain" description="Antistasin-like" evidence="7">
        <location>
            <begin position="460"/>
            <end position="485"/>
        </location>
    </feature>
    <feature type="domain" description="Antistasin-like" evidence="7">
        <location>
            <begin position="488"/>
            <end position="513"/>
        </location>
    </feature>
    <feature type="domain" description="VWFC" evidence="6">
        <location>
            <begin position="527"/>
            <end position="584"/>
        </location>
    </feature>
    <feature type="domain" description="VWFC" evidence="6">
        <location>
            <begin position="598"/>
            <end position="656"/>
        </location>
    </feature>
    <feature type="domain" description="VWFC" evidence="6">
        <location>
            <begin position="672"/>
            <end position="730"/>
        </location>
    </feature>
    <feature type="domain" description="VWFC" evidence="6">
        <location>
            <begin position="738"/>
            <end position="795"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50184"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS51252"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51323"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016095.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="956" mass="105358" checksum="BF111F345BE64939" modified="2019-05-08" version="1">MYLAAVLAGRRRPGGSGWHLGAGWLLLVLLGESGTRALVCLPCDESKCEEPKSCPGSIVLGICGCCFMCARQRNESCGGVYGLHGACDRGLRCVIRPPLNGDSITEYEVGVCEDENWDDQLLGFEPCNENLVTGCNIINGKCECDTIRTCSNPFEFPSQDTCLSALKRIEVFSVDCSTVECPPVQQVVCPLDSYETQVRLTADGCCTLPTRCECLSGLCGFPMCEAGSIPQIVSHGDGTPGKCCDVFECVNEVKPACIFNSVEYYDGDMFRMDACRFCRCQGGVSICFSAQCGELHCDRYYVPEGECCPVCEDPIHPVNNPAGCYANGQIQAHGDRWREDDCTFCQCINGDPHCVATACGQSCLNPVKVPGECCPVCEEPTYITIGPSTCEILVNCTLTERDCIYSFRLDQNGCRICQCKTREELCTGLISGCSLDCSFGFQTDVHNCEICQCRSRPKKCKPIVCDKYCPFGYLKNKHGCDICRCKKCPELPCGKICPMGFQQNNHGCLLCKCREIPDSVMPPVKTGSCLSMDGRRHEKEESWHDGCRECYCHNGREMCALITCPVPNCGNPTIHPGQCCPSCPDESIVQKPELSNPSICHAPGGEYFVEGETWNIDSCTQCTCHSGRVLCETEVCPPLLCQNPIRTQDSCCPQCPDEPLQPSSSSNESMPSYCKNDEGDIFLTAESWKPNVCTSCICMDGVISCYSESCPPVSCERPVLRKGQCCPYCIEDTIPKKVVCHFSGKTYADEERWDIDSCTHCYCLQGQTLCSTVSCPPLPCVEPINVEGSCCPMCPEMYVPEPTNIPIEKTNHRDAELEAPIWPTPSENDIVHLHRDMSHLQGDFREGIGLHSSEDTSLNSVAWVTVPIMTALVLIIALLLVNQKKQWIPVSCYRAPTKPSCLNNQLVYVDCKKGTMVQVDSSQRMLRIADPDSRYSGFYSMQKQNNLQADNFYQTV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMC2</accession>
    <name>A0A452HMC2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">AT-rich interactive domain-containing protein 4A</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="15">Retinoblastoma-binding protein 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="12">DNA-binding protein which modulates activity of several transcription factors including RB1 (retinoblastoma-associated protein) and AR (androgen receptor). May function as part of an mSin3A repressor complex. Has no intrinsic transcriptional activity. Plays a role in the regulation of epigenetic modifications at the PWS/AS imprinting center near the SNRPN promoter, where it might function as part of a complex with RB1 and ARID4B. Involved in spermatogenesis, together with ARID4B, where it acts as a transcriptional coactivator for AR and enhances expression of genes required for sperm maturation. Regulates expression of the tight junction protein CLDN3 in the testis, which is important for integrity of the blood-testis barrier. Plays a role in myeloid homeostasis where it regulates the histone methylation state of bone marrow cells and expression of various genes involved in hematopoiesis. May function as a leukemia suppressor.</text>
    </comment>
    <comment type="subunit">
        <text evidence="13">Identified in mSin3A corepressor complexes together with SIN3A, SIN3B, RBBP4, RBBP7, SAP30, BRMS1, HDAC1 and HDAC2. Interacts with BRMS1. Interacts with RB1. Interacts with ARID4B. Interacts with AR.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMC2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018380.1">
        <property type="protein sequence ID" value="ENSGAGP00000016094.1"/>
        <property type="gene ID" value="ENSGAGG00000012085.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000976">
        <property type="term" value="F:transcription cis-regulatory region binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006325">
        <property type="term" value="P:chromatin organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007283">
        <property type="term" value="P:spermatogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd16882">
        <property type="entry name" value="ARID_ARID4A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18641">
        <property type="entry name" value="CBD_RBP1_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05162">
        <property type="entry name" value="PWWP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd20459">
        <property type="entry name" value="Tudor_ARID4A_rpt1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd20461">
        <property type="entry name" value="Tudor_ARID4A_rpt2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.140:FF:000050">
        <property type="entry name" value="AT-rich interactive domain 4A (RBP1-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.140:FF:000012">
        <property type="entry name" value="AT-rich interactive domain-containing protein 4A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.60:FF:000003">
        <property type="entry name" value="AT-rich interactive domain-containing protein 4B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.140:FF:000009">
        <property type="entry name" value="AT-rich interactive domain-containing protein 4B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.140">
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.60">
        <property type="entry name" value="ARID DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051232">
        <property type="entry name" value="ARID/SWI1_ChromRemod"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012603">
        <property type="entry name" value="ARID4A/B_PWWP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001606">
        <property type="entry name" value="ARID_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036431">
        <property type="entry name" value="ARID_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047473">
        <property type="entry name" value="CBD_RBP1-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016197">
        <property type="entry name" value="Chromo-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000953">
        <property type="entry name" value="Chromo/chromo_shadow_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002999">
        <property type="entry name" value="Tudor"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025995">
        <property type="entry name" value="Tudor-knot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047472">
        <property type="entry name" value="Tudor_ARID4A_rpt1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13964:SF26">
        <property type="entry name" value="AT-RICH INTERACTIVE DOMAIN-CONTAINING PROTEIN 4A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13964">
        <property type="entry name" value="RBP-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01388">
        <property type="entry name" value="ARID"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08169">
        <property type="entry name" value="RBB1NT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11717">
        <property type="entry name" value="Tudor-knot"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01014">
        <property type="entry name" value="ARID"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00501">
        <property type="entry name" value="BRIGHT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00298">
        <property type="entry name" value="CHROMO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00333">
        <property type="entry name" value="TUDOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46774">
        <property type="entry name" value="ARID-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54160">
        <property type="entry name" value="Chromo domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF63748">
        <property type="entry name" value="Tudor/PWWP/MBT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51011">
        <property type="entry name" value="ARID"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0156">Chromatin regulator</keyword>
    <keyword evidence="4" id="KW-0221">Differentiation</keyword>
    <keyword evidence="9" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="2" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0744">Spermatogenesis</keyword>
    <keyword evidence="10" id="KW-0804">Transcription</keyword>
    <keyword evidence="8" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="5" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="ARID" evidence="17">
        <location>
            <begin position="308"/>
            <end position="400"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="124"/>
            <end position="168"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
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</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMC3</accession>
    <name>A0A452HMC3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="22">Hydroxymethylglutaryl-CoA synthase</fullName>
            <shortName evidence="22">HMG-CoA synthase</shortName>
            <ecNumber evidence="22">2.3.3.10</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="22">3-hydroxy-3-methylglutaryl coenzyme A synthase</fullName>
        </alternativeName>
    </protein>
    <organism evidence="25 26">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="26" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="25" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="25" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="22">Catalyzes the condensation of acetyl-CoA with acetoacetyl-CoA to form HMG-CoA.</text>
    </comment>
    <comment type="function">
        <text evidence="18">Catalyzes the first irreversible step in ketogenesis, condensing acetyl-CoA to acetoacetyl-CoA to form HMG-CoA, which is converted by HMG-CoA reductase (HMGCR) into mevalonate.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>acetoacetyl-CoA + acetyl-CoA + H2O = (3S)-3-hydroxy-3-methylglutaryl-CoA + CoA + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10188"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:43074"/>
            <dbReference type="ChEBI" id="CHEBI:57286"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57288"/>
            <dbReference type="EC" id="2.3.3.10"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="19">
            <dbReference type="Rhea" id="RHEA:10189"/>
        </physiologicalReaction>
    </comment>
    <comment type="pathway">
        <text evidence="2 22">Metabolic intermediate biosynthesis; (R)-mevalonate biosynthesis; (R)-mevalonate from acetyl-CoA: step 2/3.</text>
    </comment>
    <comment type="subunit">
        <text evidence="4">Homodimer.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Mitochondrion</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 22">Belongs to the thiolase-like superfamily. HMG-CoA synthase family.</text>
    </comment>
    <dbReference type="EC" id="2.3.3.10" evidence="22"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018428.1">
        <property type="protein sequence ID" value="ENSGAGP00000016140.1"/>
        <property type="gene ID" value="ENSGAGG00000012117.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00058">
        <property type="reaction ID" value="UER00102"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004421">
        <property type="term" value="F:hydroxymethylglutaryl-CoA synthase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006084">
        <property type="term" value="P:acetyl-CoA metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006695">
        <property type="term" value="P:cholesterol biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010142">
        <property type="term" value="P:farnesyl diphosphate biosynthetic process, mevalonate pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00827">
        <property type="entry name" value="init_cond_enzymes"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.47.10:FF:000008">
        <property type="entry name" value="3-hydroxy-3-methylglutaryl coenzyme A synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.47.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000590">
        <property type="entry name" value="HMG_CoA_synt_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013746">
        <property type="entry name" value="HMG_CoA_synt_C_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013528">
        <property type="entry name" value="HMG_CoA_synth_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010122">
        <property type="entry name" value="HMG_CoA_synthase_euk"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016039">
        <property type="entry name" value="Thiolase-like"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01833">
        <property type="entry name" value="HMG-CoA-S_euk"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43323">
        <property type="entry name" value="3-HYDROXY-3-METHYLGLUTARYL COENZYME A SYNTHASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43323:SF1">
        <property type="entry name" value="HYDROXYMETHYLGLUTARYL-COA SYNTHASE, MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08540">
        <property type="entry name" value="HMG_CoA_synt_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01154">
        <property type="entry name" value="HMG_CoA_synt_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53901">
        <property type="entry name" value="Thiolase-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01226">
        <property type="entry name" value="HMG_COA_SYNTHASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12" id="KW-0007">Acetylation</keyword>
    <keyword evidence="9" id="KW-0152">Cholesterol biosynthesis</keyword>
    <keyword evidence="6" id="KW-0153">Cholesterol metabolism</keyword>
    <keyword evidence="5 22" id="KW-0444">Lipid biosynthesis</keyword>
    <keyword evidence="14 22" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="15" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="26" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="11 22" id="KW-0752">Steroid biosynthesis</keyword>
    <keyword evidence="17 22" id="KW-0753">Steroid metabolism</keyword>
    <keyword evidence="13 22" id="KW-0756">Sterol biosynthesis</keyword>
    <keyword evidence="16 22" id="KW-1207">Sterol metabolism</keyword>
    <keyword evidence="8 22" id="KW-0808">Transferase</keyword>
    <keyword evidence="10" id="KW-0809">Transit peptide</keyword>
    <feature type="domain" description="Hydroxymethylglutaryl-coenzyme A synthase N-terminal" evidence="23">
        <location>
            <begin position="18"/>
            <end position="191"/>
        </location>
    </feature>
    <feature type="domain" description="Hydroxymethylglutaryl-coenzyme A synthase C-terminal" evidence="24">
        <location>
            <begin position="192"/>
            <end position="475"/>
        </location>
    </feature>
    <feature type="active site" description="Proton donor/acceptor" evidence="20">
        <location>
            <position position="100"/>
        </location>
    </feature>
    <feature type="active site" description="Acyl-thioester intermediate" evidence="20">
        <location>
            <position position="134"/>
        </location>
    </feature>
    <feature type="active site" description="Proton donor/acceptor" evidence="20">
        <location>
            <position position="269"/>
        </location>
    </feature>
    <feature type="binding site" evidence="21">
        <location>
            <position position="172"/>
        </location>
        <ligand>
            <name>CoA</name>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="21">
        <location>
            <position position="226"/>
        </location>
        <ligand>
            <name>CoA</name>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="21">
        <location>
            <position position="274"/>
        </location>
        <ligand>
            <name>CoA</name>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="21">
        <location>
            <position position="278"/>
        </location>
        <ligand>
            <name>CoA</name>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004173"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
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            <dbReference type="ARBA" id="ARBA00005218"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00007061"/>
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    </evidence>
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            <dbReference type="ARBA" id="ARBA00011738"/>
        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00022516"/>
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    </evidence>
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            <dbReference type="ARBA" id="ARBA00022548"/>
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    </evidence>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
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            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
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            <dbReference type="ARBA" id="ARBA00022778"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022955"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023166"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00037714"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00049887"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="PIRSR" id="PIRSR610122-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="PIRSR" id="PIRSR610122-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="RuleBase" id="RU364071"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="Pfam" id="PF01154"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="Pfam" id="PF08540"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016140.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="494" mass="55378" checksum="89FFB7915A664DFC" modified="2019-05-08" version="1">IPHFSTVSVEHLSQLDSWPKDVGILALEVYFPSQYVEQVELEKYDGVEAGKYTLGLGQKQMGFCSAHEDINSLCLTVVQRLVERSRLSWDSIGRLEVGTETIIDKSKAVKTVLMQLFQDSGNTDVEGIDTTNACYGGTASLFNSVDWVESSSWDGRYAMVVCGDIAVYATGNARPTGGAGAVAMLVGPEAPLVLERGVRGTHMEHVYDFYKPDLASEYPMVDGKLSIQCYFRALDQCYSVYRKKIQSQWQKVGVDKPFTLEDFQFMIFHTPFCKLVQKSVGRMMLNDFLAAPNPDMQSGLYKGLQAFRGVKLEDTYSDKDVEKAFQKASLEMFSQKTKPSLLLSSRNGNMYTPSMYGCLASLLAQYSAQQLAGSRIGAFSYGSGLAASLFSMRVSQDAAPGSPLDKLVSSLSDLQTRLDARKRVPPEEFAEIMKQREESHHLVNHTPQGSKDDLFPGTWYLERVDDKHRRQYARRRLMPQIKHTYTXPLFISFF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMC5</accession>
    <name>A0A452HMC5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Fibronectin type-III domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the type I cytokine receptor family. Type 2 subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMC5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018383.1">
        <property type="protein sequence ID" value="ENSGAGP00000016097.1"/>
        <property type="gene ID" value="ENSGAGG00000011475.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009897">
        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004896">
        <property type="term" value="F:cytokine receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000494">
        <property type="entry name" value="Leptin receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000501">
        <property type="entry name" value="Leptin receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000568">
        <property type="entry name" value="Leptin receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000613">
        <property type="entry name" value="Leptin receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003529">
        <property type="entry name" value="Hematopoietin_rcpt_Gp130_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010457">
        <property type="entry name" value="IgC2-like_lig-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041182">
        <property type="entry name" value="LEP-R_IGD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23037">
        <property type="entry name" value="CYTOKINE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23037:SF44">
        <property type="entry name" value="LEPTIN RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06328">
        <property type="entry name" value="Lep_receptor_Ig"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18589">
        <property type="entry name" value="ObR_Ig"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01353">
        <property type="entry name" value="HEMATOPO_REC_L_F2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7 11" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-0675">Receptor</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="3 11" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 11" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="810"/>
            <end position="832"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="12">
        <location>
            <begin position="209"/>
            <end position="303"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="12">
        <location>
            <begin position="510"/>
            <end position="606"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="12">
        <location>
            <begin position="711"/>
            <end position="804"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008921"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016097.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="866" mass="98750" checksum="109BA6A56FEF118F" modified="2019-05-08" version="1">VAWSVVLTGATRVPFRLVAHCMVHQIPASSFMLPCMSPNETSVIPLTAGVVASSPGLHREHGTTETNSLMLVAEESFLCCLWSETNANCSLYTAGMETKKFTSSEVSSFAPQQTDLSWNIQCWTKGDLELFVLKYIGKLSLEDISTGSLKGNFMVTPCNGNGSDKYECHIPSLKLNYTYITWLNIINGVTLLQSPLMSVRPINIVKPESPLNLRLEMTDKGQLKICWSNPVLTPYPLQYEVKFSGNTTQNAWQVVEIVIETSLIIGNMLVGSSYLVQVRCKSLHGTGFWSDWSTPYNLNTEDVMYFPPKILTSVGSNVSFHCLYNDKNKMILSKKIVWWLNLAEEIPERQYTLVNDRVSRVTLFNLNATKPRGKFFYNALYCCNQNRECHHRYAELYVVDVNINITCETDGYLTKMTCRWSANTNTLLVGSSLQLKYYRSSIYCSDFPSIPPKSEAKECHLQRNHYYECTFQPIFLLSGYTMWIEIKHQLGTLESPPTCVIPADVVKPFPPSNVKAEITKNIGLLNVSWTNPAFPNSDLKFQIRYSVNREEILWEIFEVSNAPTRSVMIKVLDLCVVYIVQVRCSGLDGLGYWSDWSKPAYTIVQDIQAPLRGPEFWRVINEDPIRKQKNVTLVWKPLMKNYSLCSVCGYIVKQHTSENITWTEYVTSGTTYTYPWMEDAHTITILAVNSVGASSMNFNLTLSQQMSTVNIVQSLSAYPVNSSCVILTWTLSPQMYVITSFVIEWKNLNEEEQMKWIRVAPNISKYYIYDHFILIEKYQFSLYPIFPEGVGNPKTTDEFIKDRSEKQNDAGLYVILPIVISSFVLLLGTLLISQQRMKKLFWEDVPNPKNCSWAQGVNFQKRMDIL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMC9</accession>
    <name>A0A452HMC9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="7">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HMC9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018382.1">
        <property type="protein sequence ID" value="ENSGAGP00000016096.1"/>
        <property type="gene ID" value="ENSGAGG00000012086.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004867">
        <property type="term" value="F:serine-type endopeptidase inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.40.20:FF:000003">
        <property type="entry name" value="cysteine-rich motor neuron 1 protein isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.40.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.20.200.20">
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004094">
        <property type="entry name" value="Antistasin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052624">
        <property type="entry name" value="CRIM1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045813">
        <property type="entry name" value="CRIM1_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009030">
        <property type="entry name" value="Growth_fac_rcpt_cys_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000867">
        <property type="entry name" value="IGFBP-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001007">
        <property type="entry name" value="VWF_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46439">
        <property type="entry name" value="CYSTEINE-RICH MOTOR NEURON 1 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46439:SF1">
        <property type="entry name" value="CYSTEINE-RICH MOTOR NEURON 1 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02822">
        <property type="entry name" value="Antistasin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19442">
        <property type="entry name" value="CRIM1_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00219">
        <property type="entry name" value="IGFBP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00093">
        <property type="entry name" value="VWC"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23334">
        <property type="entry name" value="VWC2L_2nd"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00121">
        <property type="entry name" value="IB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00214">
        <property type="entry name" value="VWC"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SMART" id="SM00215">
        <property type="entry name" value="VWC_out"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57603">
        <property type="entry name" value="FnI-like domain"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57184">
        <property type="entry name" value="Growth factor receptor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51252">
        <property type="entry name" value="ANTISTASIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51323">
        <property type="entry name" value="IGFBP_N_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01208">
        <property type="entry name" value="VWFC_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50184">
        <property type="entry name" value="VWFC_2"/>
        <property type="match status" value="6"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="2" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0732">Signal</keyword>
    <keyword evidence="2" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="2" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="3">
        <location>
            <begin position="1"/>
            <end position="37"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019100208" evidence="3">
        <location>
            <begin position="38"/>
            <end position="944"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="849"/>
            <end position="869"/>
        </location>
    </feature>
    <feature type="domain" description="IGFBP N-terminal" evidence="6">
        <location>
            <begin position="36"/>
            <end position="115"/>
        </location>
    </feature>
    <feature type="domain" description="VWFC" evidence="4">
        <location>
            <begin position="336"/>
            <end position="393"/>
        </location>
    </feature>
    <feature type="domain" description="VWFC" evidence="4">
        <location>
            <begin position="403"/>
            <end position="459"/>
        </location>
    </feature>
    <feature type="domain" description="Antistasin-like" evidence="5">
        <location>
            <begin position="471"/>
            <end position="500"/>
        </location>
    </feature>
    <feature type="domain" description="VWFC" evidence="4">
        <location>
            <begin position="515"/>
            <end position="572"/>
        </location>
    </feature>
    <feature type="domain" description="VWFC" evidence="4">
        <location>
            <begin position="586"/>
            <end position="644"/>
        </location>
    </feature>
    <feature type="domain" description="VWFC" evidence="4">
        <location>
            <begin position="660"/>
            <end position="718"/>
        </location>
    </feature>
    <feature type="domain" description="VWFC" evidence="4">
        <location>
            <begin position="726"/>
            <end position="783"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50184"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS51252"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS51323"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016096.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="944" mass="103652" checksum="BA93E35DF943BE4B" modified="2019-05-08" version="1">MYLAAVLAGRRRPGGSGWHLGAGWLLLVLLGESGTRALVCLPCDESKCEEPKSCPGSIVLGICGCCFMCARQRNESCGGVYGLHGACDRGLRCVIRPPLNGDSITEYEVGVCEDENWDDQLLGFEPCNENLVTGCNIINGKCECDTIRTCSNPFEFPSQDTCLSALKRIEEEKPDCSKARCEVQFSPRCPEDSILIEGYAPPGECCPLPSRCVCNPAGCLRKVCQPGYLNILVSKASGKPGECCDLYECKPVFSVDCSTVECPPVQQVVCPLDSYETQVRLTADGCCTLPTRCECLSGLCGFPMCEAGSIPQIVSHGDGTPGKCCDVFECVNEVKPACIFNSVEYYDGDMFRMDACRFCRCQGGVSICFSAQCGELHCDRYYVPEGECCPVCEDPIHPVNNPAGCYANGQIQAHGDRWREDDCTFCQCINGDPHCVATACGQSCLNPVKVPGECCPVCEEPTYITIGPSTCEILVNCTLTERDCIYSFRLDQNGCRICQCKTKIPDSVMPPVKTGSCLSMDGRRHEKEESWHDGCRECYCHNGREMCALITCPVPNCGNPTIHPGQCCPSCPDESIVQKPELSNPSICHAPGGEYFVEGETWNIDSCTQCTCHSGRVLCETEVCPPLLCQNPIRTQDSCCPQCPDEPLQPSSSSNESMPSYCKNDEGDIFLTAESWKPNVCTSCICMDGVISCYSESCPPVSCERPVLRKGQCCPYCIEDTIPKKVVCHFSGKTYADEERWDIDSCTHCYCLQGQTLCSTVSCPPLPCVEPINVEGSCCPMCPEMYVPEPTNIPIEKTNHRDAELEAPIWPTPSENDIVHLHRDMSHLQGDFREGIGLHSSEDTSLNSVAWVTVPIMTALVLIIALLLVNQKKQWIPVSCYRAPTKPSCLNNQLVYVDCKKGTMVQVDSSQRMLRIADPDSRYSGFYSMQKQNNLQADNFYQTV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMD4</accession>
    <name>A0A452HMD4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Fibronectin type-III domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the type I cytokine receptor family. Type 2 subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMD4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018387.1">
        <property type="protein sequence ID" value="ENSGAGP00000016101.1"/>
        <property type="gene ID" value="ENSGAGG00000011475.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009897">
        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004896">
        <property type="term" value="F:cytokine receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000494">
        <property type="entry name" value="Leptin receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000501">
        <property type="entry name" value="Leptin receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000568">
        <property type="entry name" value="Leptin receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000613">
        <property type="entry name" value="Leptin receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003529">
        <property type="entry name" value="Hematopoietin_rcpt_Gp130_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010457">
        <property type="entry name" value="IgC2-like_lig-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041182">
        <property type="entry name" value="LEP-R_IGD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23037">
        <property type="entry name" value="CYTOKINE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23037:SF44">
        <property type="entry name" value="LEPTIN RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06328">
        <property type="entry name" value="Lep_receptor_Ig"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18589">
        <property type="entry name" value="ObR_Ig"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01353">
        <property type="entry name" value="HEMATOPO_REC_L_F2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7 11" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-0675">Receptor</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="3 11" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 11" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="809"/>
            <end position="831"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="861"/>
            <end position="883"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="12">
        <location>
            <begin position="211"/>
            <end position="302"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="12">
        <location>
            <begin position="509"/>
            <end position="605"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="12">
        <location>
            <begin position="710"/>
            <end position="803"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008921"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016101.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="885" mass="101282" checksum="379AFBC88CA29BF3" modified="2019-05-08" version="1">MQMAVAHCMVHQIPASSFMLPCMSPNETSVIPLTAGVVASSPGLHREHGTTETNSLMLVAEESFLCCLWSETNANCSLYTAGMETKKFTSSEVSSFAPQQTGSWNIQCWTKGDLELFVCILKLSNKKSYLNGEFKINLLYVVLMPFLESSLKGNFMVTPCNGNGSDKYECHIPSLKLNYTYITWLNIINGVTLLQSPLMSVRPINIVKPESPLNLRLEMTDKGQLKICWSNPVLTPYPLQYEVKFSGNTTQNAWQIVIETSLIIGNMLVGSSYLVQVRCKSLHGTGFWSDWSTPYNLNTEDVMYFPPKILTSVGSNVSFHCLYNDKNKMILSKKIVWWLNLAEEIPERQYTLVNDRVSRVTLFNLNATKPRGKFFYNALYCCNQNRECHHRYAELYVVDVNINITCETDGYLTKMTCRWSANTNTLLVGSSLQLKYYRSSIYCSDFPSIPPKSEAKECHLQRNHYYECTFQPIFLLSGYTMWIEIKHQLGTLESPPTCVIPADVVKPFPPSNVKAEITKNIGLLNVSWTNPAFPNSDLKFQIRYSVNREEILWEIFEVSNAPTRSVMIKVLDLCVVYIVQVRCSGLDGLGYWSDWSKPAYTIVQDIQAPLRGPEFWRVINEDPIRKQKNVTLVWKPLMKNYSLCSVCGYIVKQHTSENITWTEYVTSGTTYTYPWMEDAHTITILAVNSVGASSMNFNLTLSQQMSTVNIVQSLSAYPVNSSCVILTWTLSPQMYVITSFVIEWKNLNEEEQMKWIRVAPNISKYYIYDHFILIEKYQFSLYPIFPEGVGNPKTTDEFIKDRSEKQNDAGLYVILPIVISSFVLLLGTLLISQQRMKKLFWEDVPNPKNCSWAQGVNFQKVWVSIMIRFLVWVYVFHTLTFSLKC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMD5</accession>
    <name>A0A452HMD5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Transmembrane protein 45B</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the TMEM45 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMD5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018394.1">
        <property type="protein sequence ID" value="ENSGAGP00000016107.1"/>
        <property type="gene ID" value="ENSGAGG00000012019.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006904">
        <property type="entry name" value="DUF716"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042127">
        <property type="entry name" value="TMEM45"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16007">
        <property type="entry name" value="EPIDIDYMAL MEMBRANE PROTEIN E9-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16007:SF21">
        <property type="entry name" value="TRANSMEMBRANE PROTEIN 45A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04819">
        <property type="entry name" value="DUF716"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 6" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 6" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 6" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="12"/>
            <end position="31"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="52"/>
            <end position="70"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="90"/>
            <end position="111"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="123"/>
            <end position="144"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="156"/>
            <end position="174"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="186"/>
            <end position="207"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="227"/>
            <end position="248"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006948"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016107.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="9">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="272" mass="30325" checksum="E1707638E7C838C5" modified="2019-05-08" version="1">MSSFKGNALRGTFFLIFGLGWSVKYPLKYLSRKLNADNQASRWIEKLDILEGAAKAFLAVAGMLAEQFVPDGPHMGLYSGGTYSWEKLTNWQYTTMYLFYGLSGIVDVLSYSPLKLSLGLDRLILIAAACFGVGVGFLFCFHDYNNTALDQHLHSLLLIAIFGGALCALMEVFLRDHTILEFFRTSFFILQGSWFWQIGFVLSPPWGGPGWDQTDRGNFNFLTMCFFWHYAAALLVTAANSVFSYWYVVGQGSGIQICIYSAILSGSDEKGG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMD6</accession>
    <name>A0A452HMD6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Transmembrane protein 45B</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the TMEM45 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMD6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018397.1">
        <property type="protein sequence ID" value="ENSGAGP00000016110.1"/>
        <property type="gene ID" value="ENSGAGG00000012098.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006904">
        <property type="entry name" value="DUF716"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042127">
        <property type="entry name" value="TMEM45"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16007">
        <property type="entry name" value="EPIDIDYMAL MEMBRANE PROTEIN E9-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16007:SF59">
        <property type="entry name" value="TRANSMEMBRANE PROTEIN 45B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04819">
        <property type="entry name" value="DUF716"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4" id="KW-1133">Transmembrane helix</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006948"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00039264"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016110.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="241" mass="27900" checksum="F3043C30B4621477" modified="2019-05-08" version="1">MGIFGDHTLPDTLILMLEFWWSLNYPLKRLNENCCTNRCYQSGEFLEAAGKLFSFVGKLLKFLPAVSKQIHQHSTMYLSFLFPDRFVFSQVMFIEGLLFHFHVHYRSALQQHVHLLSTVALCNNAFSLCLEVFIPHHPHPTALQHPCSPHPGSWFWQISFVLYPPWGWLDWDQTDDGNIMFSTRCVSWHYTAALINLLSITQFLRETLPPSGLRVPTRAQDKSLFSSRKRGGLKQGVSYPR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMG0</accession>
    <name>A0A452HMG0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">SWIM-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HMG0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018425.1">
        <property type="protein sequence ID" value="ENSGAGP00000016137.1"/>
        <property type="gene ID" value="ENSGAGG00000012100.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031462">
        <property type="term" value="C:Cul2-RING ubiquitin ligase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007527">
        <property type="entry name" value="Znf_SWIM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048370">
        <property type="entry name" value="ZSWIM4-8_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22619">
        <property type="entry name" value="ZINC FINGER SWIM DOMAIN CONTAINING PROTEIN 4, 5, 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22619:SF3">
        <property type="entry name" value="ZINC FINGER SWIM DOMAIN-CONTAINING PROTEIN 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21055">
        <property type="entry name" value="ZSWIM4-8_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50966">
        <property type="entry name" value="ZF_SWIM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0862">Zinc</keyword>
    <keyword evidence="2 4" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="SWIM-type" evidence="5">
        <location>
            <begin position="9"/>
            <end position="46"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00325"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50966"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016137.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="978" mass="111160" checksum="6C190A63120A18F6" modified="2019-05-08" version="1">MQSEPETVCKVAISFDRCKITSVTCSCGNKDIFYCAHVVALSLYRIRKPDQVKLHLPISETLFQMNRDQLQKFVQYLITMHHTEVLPTAQKLADEILSQNSEINQVHGAPDPTAGASIDDENCWHLDEEQVQEQVKLFLSQGGYHGSGKQLNLLFAKVREMLKMRDSNGARMLTLITEQFMADPRLSLWRQQGTAMTDKYRQLWDELGALWMCIVLNPHCKLEQKASWLKQLKKWNCVDVCPWEDGNHGNELPNLTNALPQGANANQDSLNRPHRTVFTRAIEACDLHWQDSHLQHIISSDLYTNYCYHDDTENSLFDSHGWPLWHEHVPTACARVDALRSHGYPREALRLAIAIVNTLRRQQQKQLDMFRAQKKELLHKGVTSITNLEGWVGHPLDPVGTLFSSLMEACRVDDESFHGFSDFTENMGQCKSLEYQHLPSHKFLEEGESYLTLAVEVALIGLGQQRIMPDGLYAQEKVCRNEEQLISKLQEIELDDTLVKIFRKQAVFLLEAGPYSGLGEIIHRESVPMHTFAKYLFTSLLPHDAELAYKTALRAMRLLVLESTAPSGDMSRPHHIASVVPNRYPRWFTLSHIESQQCELASTMLTAAKGDVRRLETVLESIQKNIHSSSHIFKLAQDAFKIATLMDSLPDITLLKVSLELGLQVMRMTLSTLNWRRREMVRWLVTCATEVGVYALDSIMQNWFTLFTPTEATSIVATTVMSNSTIVRLHLDCLQQEKLASSARTLALQCAMKDPQNCALSALTLCEKDHIAFETAYQIVLDAATKGMSYTQLFTIARYMEHRGYPMRAYKLATLAMNHLNLSYNQDTHPAINDVLWACALSHSLGKNELAAIIPLVVKSVKCATVLSDILRRCTLTTPGMVGLHGRRNSGKLMSLDKAPLRQLLDATIGAYINTTHSRLTHISPRHYSEFIEFLSKARETFLMAHDGHIQFTQFIDNLKQIYKGKKKLMMLVRERFG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMG3</accession>
    <name>A0A452HMG3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Slit homolog 2 protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="15">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018474.1">
        <property type="protein sequence ID" value="ENSGAGP00000016180.1"/>
        <property type="gene ID" value="ENSGAGG00000011388.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048495">
        <property type="term" value="F:Roundabout binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072359">
        <property type="term" value="P:circulatory system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060560">
        <property type="term" value="P:developmental growth involved in morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060562">
        <property type="term" value="P:epithelial tube morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001822">
        <property type="term" value="P:kidney development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050919">
        <property type="term" value="P:negative chemotaxis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030308">
        <property type="term" value="P:negative regulation of cell growth"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010605">
        <property type="term" value="P:negative regulation of macromolecule metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010977">
        <property type="term" value="P:negative regulation of neuron projection development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0021772">
        <property type="term" value="P:olfactory bulb development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050770">
        <property type="term" value="P:regulation of axonogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032535">
        <property type="term" value="P:regulation of cellular component size"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050767">
        <property type="term" value="P:regulation of neurogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031290">
        <property type="term" value="P:retinal ganglion cell axon guidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0022029">
        <property type="term" value="P:telencephalon cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd00054">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="CDD" id="cd00110">
        <property type="entry name" value="LamG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000045">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000053">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000054">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000062">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000063">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000099">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000289">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.200:FF:000013">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000002">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000004">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.200">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000483">
        <property type="entry name" value="Cys-rich_flank_reg_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006207">
        <property type="entry name" value="Cys_knot_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001881">
        <property type="entry name" value="EGF-like_Ca-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000152">
        <property type="entry name" value="EGF-type_Asp/Asn_hydroxyl_site"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018097">
        <property type="entry name" value="EGF_Ca-bd_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003645">
        <property type="entry name" value="Fol_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001791">
        <property type="entry name" value="Laminin_G"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001611">
        <property type="entry name" value="Leu-rich_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003591">
        <property type="entry name" value="Leu-rich_rpt_typical-subtyp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000372">
        <property type="entry name" value="LRRNT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051355">
        <property type="entry name" value="Notch/Slit_guidance"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45836">
        <property type="entry name" value="SLIT HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45836:SF2">
        <property type="entry name" value="SLIT HOMOLOG 2 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00008">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00054">
        <property type="entry name" value="Laminin_G_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00560">
        <property type="entry name" value="LRR_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13855">
        <property type="entry name" value="LRR_8"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01463">
        <property type="entry name" value="LRRCT"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01462">
        <property type="entry name" value="LRRNT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00041">
        <property type="entry name" value="CT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="SMART" id="SM00179">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SMART" id="SM00274">
        <property type="entry name" value="FOLN"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00282">
        <property type="entry name" value="LamG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00368">
        <property type="entry name" value="LRR_RI"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SMART" id="SM00365">
        <property type="entry name" value="LRR_SD22"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SMART" id="SM00369">
        <property type="entry name" value="LRR_TYP"/>
        <property type="match status" value="12"/>
    </dbReference>
    <dbReference type="SMART" id="SM00082">
        <property type="entry name" value="LRRCT"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00013">
        <property type="entry name" value="LRRNT"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52058">
        <property type="entry name" value="L domain-like"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00010">
        <property type="entry name" value="ASX_HYDROXYL"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01185">
        <property type="entry name" value="CTCK_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01225">
        <property type="entry name" value="CTCK_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01186">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01187">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50025">
        <property type="entry name" value="LAM_G_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51450">
        <property type="entry name" value="LRR"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0145">Chemotaxis</keyword>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="10" id="KW-0221">Differentiation</keyword>
    <keyword evidence="12 15" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5 15" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7" id="KW-0358">Heparin-binding</keyword>
    <keyword evidence="6" id="KW-0433">Leucine-rich repeat</keyword>
    <keyword evidence="11" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="8" id="KW-0732">Signal</keyword>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="723"/>
            <end position="758"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="760"/>
            <end position="799"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="801"/>
            <end position="837"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="848"/>
            <end position="886"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="888"/>
            <end position="924"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="933"/>
            <end position="969"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin G" evidence="17">
        <location>
            <begin position="972"/>
            <end position="1145"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="1149"/>
            <end position="1180"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="1183"/>
            <end position="1219"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="1224"/>
            <end position="1260"/>
        </location>
    </feature>
    <feature type="domain" description="CTCK" evidence="16">
        <location>
            <begin position="1265"/>
            <end position="1340"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="748"/>
            <end position="757"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="789"/>
            <end position="798"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="876"/>
            <end position="885"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="914"/>
            <end position="923"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="959"/>
            <end position="968"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="1170"/>
            <end position="1179"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="1187"/>
            <end position="1197"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="1209"/>
            <end position="1218"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="1228"/>
            <end position="1238"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="1250"/>
            <end position="1259"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022500"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022614"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022674"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00074955"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS01225"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50025"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016180.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1341" mass="148455" checksum="1C196306B2D5F2D4" modified="2019-05-08" version="1">MPKLRTFRLHSNNLYCDCHLAWLSDWLRQRPRVGLYTQCMGPSHLRGHNVAEVQKREFVCSGHQSFMAPSCSVLHCPTACTCSNNIVDCRGKGLTEIPTNLPETITEIRLEQNSIKVIPPGAFSPYKKLRRIDLSNNQISETAPDAFQGLRSLNSLVLYGNKITELPKGLFEGLFSLQLLLLNANKINCLRVDAFQDLHNLNLLSLYDNKLQTIAKGTFSPLRSIQTLHLAQNPFICDCHLKWLADYLHANPIETSGARCTSPRRLANKRIGQIKSKKFRCSAKEQYFIPGTEDYRSKLSGDCFADLACPEKCRCEGTTVDCSNQKLNKIPEHVPQYTAELRLNNNEFTVLEATGIFKKLPQLRKINLSNNKITDIEEGAFEGASGVNELLLTSNRLETIRHKMFKGLESLKTLMLRSNRISCIGNDSFTGLSSVRLLSLYDNQITTVAPGAFDTLHSLSTLNLLANPFNCNCHLAWLGDWLRKKRIVTGNPRCQKPYFLKEIPIQDVAIQDFTCDDGNDDNSCSPLSRCPAECTCLDTVVRCSNKGLKVLPKGIPKDVTELYLDGNQFTLVPKELSNYKHLTLIDLSNNRISTLSNQSFSNMTQLLTLILSYNRLRCIPARTFDGLKSLRLLSLHGNDISVVPEGAFNDLSALSHLAIGANPLYCDCNMQWLSDWVKSEYKEPGIARCAGPGEMADKLLLTTPSKKFTCQGPVDVNILAKCSPCLSNPCKNDGTCNNDPVDFYTCTCPYGFKGQDCDIPIHACISNPCQHGGTCHLKEGEKDGFWCTCADGFEGENCEVNVDDCEDNDCENNSTCVDGINNYTCLCPPEYTAANLNEVEKGELCEEKLDFCAQDLNPCQHDSKCILTLKGYKCDCTPGYVGEHCDMDYDDCQDNKCKNGAQCTDAVNGYTCICPEGYSGLFCEFSPPVVLPRTSPCDNYECQNGAQCIVRVNEPICQCLSGYQGDKCEKLVSVNFVSKESYLQIPSSKVRPQTNITLQIATDEDSGILLYKGDRDHIAMELYRGRVRVSYDTGSYPASAIYSVETINDGNFHIVELLAMDQILSLSVDGGSPKIITNLSKQSTLNVDSPLYVGGMPVKNNVAALRQSPGQNGSSFHGCIRNLYINSELQDFRNVPLQVGILPGCEPCHKKVCVHGTCHAMSQSGFTCECEGGWTGPLCDQQTNDPCLGNKCVHGTCLPINAFSYSCKCLQGHGGVLCDEEEELFNPCQSIKCKHGKCRLSGLGKPYCECSSGYTGDSCDKEISCRGERIRDYYQKQQGYAACQTTKKVSRLECKGGCSTGQCCGPLRSKRRKYSFECTDGSSFVDEVEKVVKCGCTKCPS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMG6</accession>
    <name>A0A452HMG6_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the actin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMG6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018431.1">
        <property type="protein sequence ID" value="ENSGAGP00000016143.1"/>
        <property type="gene ID" value="ENSGAGG00000012115.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="CDD" id="cd10207">
        <property type="entry name" value="ASKHA_NBD_Arp10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.640.10:FF:000017">
        <property type="entry name" value="Actin-related protein 10 homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.420.40">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.640.10">
        <property type="entry name" value="Actin, Chain A, domain 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004000">
        <property type="entry name" value="Actin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043129">
        <property type="entry name" value="ATPase_NBD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11937">
        <property type="entry name" value="ACTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00022">
        <property type="entry name" value="Actin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00268">
        <property type="entry name" value="ACTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53067">
        <property type="entry name" value="Actin-like ATPase domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="RuleBase" id="RU000487"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016143.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="344" mass="38228" checksum="9965854F3A076ED0" modified="2019-05-08" version="1">MLYFRHLLVNPRDRRVVIVESVLCPSHFRETLTRVLFKYFEVPSVLFAPSHLMSLLTLGINSAMVLDCGYAESLVLPIYEGIPILSCWGALPLGGKAIHKELENHLLEQCSVDTGVAKGQRLPSVMGSIPEDIVEDIKVRTCFVSDLQRGLKIQAAKFNMDGSAERPSPPPDVDYPLDGEKILHVVGSIRDSVVEILFEQDNEEKSVATLILDALIECPIDTRKQLAENLVVIGGTAMLPGFLHRLMAEIRYLVDKPKYKEALATKTFRIHTPPAKPNCVAWLGGAIFGALQDILGSRSVSKEYYSQTGRIPDWCCLNNPPLEMMFDVGKAPPPLMKRAFSTEK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMG7</accession>
    <name>A0A452HMG7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">ZSWIM4-8 C-terminal domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HMG7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018427.1">
        <property type="protein sequence ID" value="ENSGAGP00000016139.1"/>
        <property type="gene ID" value="ENSGAGG00000012100.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031462">
        <property type="term" value="C:Cul2-RING ubiquitin ligase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048370">
        <property type="entry name" value="ZSWIM4-8_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22619">
        <property type="entry name" value="ZINC FINGER SWIM DOMAIN CONTAINING PROTEIN 4, 5, 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22619:SF3">
        <property type="entry name" value="ZINC FINGER SWIM DOMAIN-CONTAINING PROTEIN 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21055">
        <property type="entry name" value="ZSWIM4-8_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="3" id="KW-0862">Zinc</keyword>
    <keyword evidence="2" id="KW-0863">Zinc-finger</keyword>
    <feature type="signal peptide" evidence="4">
        <location>
            <begin position="1"/>
            <end position="17"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019145252" description="ZSWIM4-8 C-terminal domain-containing protein" evidence="4">
        <location>
            <begin position="18"/>
            <end position="784"/>
        </location>
    </feature>
    <feature type="domain" description="ZSWIM4-8 C-terminal" evidence="5">
        <location>
            <begin position="598"/>
            <end position="775"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="Pfam" id="PF21055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016139.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="784" mass="89093" checksum="9198D4E7CBD9E442" modified="2019-05-08" version="1">MWRYMCTVNFLKSGALWMCIVLNPHCKLEQKASWLKQLKKWNCVDVCPWEDGNHGNELPNLTNALPQGANANQDSLNRPHRTVFTRAIEACDLHWQDSHLQHIISSDLYTNYCYHDDTENSLFDSHGWPLWHEHVPTACARVDALRSHGYPREALRLAIAIVNTLRRQQQKQLDMFRAQKKELLHKGVTSITNLEGWVGHPLDPVGTLFSSLMEACRVDDESFHGFSDFTENMGQCKSLEYQHLPSHKFLEEGESYLTLAVEVALIGLGQQRIMPDGLYAQEKVCRNEEQLISKLQEIELDDTLVKIFRKQAVFLLEAGPYSGLGEIIHRESVPMHTFAKYLFTSLLPHDAELAYKTALRAMRLLVLESTAPSGDMSRPHHIASVVPNRYPRWFTLSHIESQQCELASTMLTAAKGDVRRLETVLESIQKNIHSSSHIFKLAQDAFKIATLMDSLPDITLLKVSLELGLQVMRMTLSTLNWRRREMVRWLVTCATEVGVYALDSIMQNWFTLFTPTEATSIVATTVMSNSTIVRLHLDCLQQEKLASSARTLALQCAMKDPQNCALSALTLCEKDHIAFETAYQIVLDAATKGMSYTQLFTIARYMEHRGYPMRAYKLATLAMNHLNLSYNQDTHPAINDVLWACALSHSLGKNELAAIIPLVVKSVKCATVLSDILRRCTLTTPGMVGLHGRRNSGKLMSLDKAPLRQLLDATIGAYINTTHSRLTHISPRHYSEFIEFLSKARETFLMAHDGHIQFTQFIDNLKQIYKGKKKLMMLVRERFG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMH2</accession>
    <name>A0A452HMH2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">C2H2-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMH2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018432.1">
        <property type="protein sequence ID" value="ENSGAGP00000016144.1"/>
        <property type="gene ID" value="ENSGAGG00000012119.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd21441">
        <property type="entry name" value="KLF12_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000021">
        <property type="entry name" value="Basic krueppel-like factor 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000018">
        <property type="entry name" value="Krueppel-like factor 15"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000563">
        <property type="entry name" value="Krueppel-like factor 8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23235:SF56">
        <property type="entry name" value="KRUEPPEL-LIKE FACTOR 12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23235">
        <property type="entry name" value="KRUEPPEL-LIKE TRANSCRIPTION FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0678">Repressor</keyword>
    <keyword evidence="9" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 11" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="290"/>
            <end position="319"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="320"/>
            <end position="349"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="350"/>
            <end position="372"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="86"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="218"/>
            <end position="246"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="268"/>
            <end position="287"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="87"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="272"/>
            <end position="284"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022491"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016144.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="375" mass="41247" checksum="9E5CABF2EEE9FC29" modified="2019-05-08" version="1">MKRKAIKDINAFENRMLMLDGMPAVRVKTELLESEQGSPNVHNYPDMEAVPLLLNNMKADPPEDSLSTDHFQTQTEPVDLSINKARSSPIAASSSPVSMTASASSPSSTSTSSSSRPASSPTVITSVSSAASVPSVLTPGPLVASASGVGGQQFLHIIHPVPPSSPMNLQSNKMSHVHRIPVVVQSVPVVYTAVRSPGNMNNTIVVPLLEDGRSHVKAQMDPRGISPRQIKSDSDDDDLPNVTLDSVNETGSTALSIARAVQDSISPFNIESTRRQRRSESPDSRKRRIHRCDFEGCNKVYTKSSHLKAHRRTHTGEKPYKCTWEGCTWKFARSDELTRHYRKHTGVKPFKCADCDRSFSRSDHLALHRRRHMLV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMH4</accession>
    <name>A0A452HMH4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Pro-neuregulin-2, membrane-bound isoform</fullName>
        </recommendedName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Direct ligand for ERBB3 and ERBB4 tyrosine kinase receptors. Concomitantly recruits ERBB1 and ERBB2 coreceptors, resulting in ligand-stimulated tyrosine phosphorylation and activation of the ERBB receptors. May also promote the heterodimerization with the EGF receptor.</text>
    </comment>
    <comment type="subunit">
        <text evidence="15">Interacts with ERBB3 and ERBB4.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the neuregulin family.</text>
    </comment>
    <comment type="caution">
        <text evidence="17">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMH4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018438.1">
        <property type="protein sequence ID" value="ENSGAGP00000016148.1"/>
        <property type="gene ID" value="ENSGAGG00000012118.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008083">
        <property type="term" value="F:growth factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048513">
        <property type="term" value="P:animal organ development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035556">
        <property type="term" value="P:intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
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            <end position="262"/>
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        <location>
            <begin position="212"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
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        <source>
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        <source>
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        <source>
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    </evidence>
    <sequence length="642" mass="72154" checksum="89A1499CBD8947AC" modified="2019-05-08" version="1">MVQPYPIFQMHSWFISLLRSIQIQPLSHYRVSAAADQMLLLEPQQRETRLYNFPCPVPLTEAGFHTGLISLFTAVPPKLKKLKSPNVHVGEKISLKCEATTGNPQPSYKWYKDGKELKKSKDIRIKYGNGKKISRLQFNKVKLEDAGEYSCEAENVLGKDTAKGSLNVRSVATTLSSWSGHARKCNETAKSYCVNGGVCYYIEGINQLSCKCPIEFTGDRCEHFAMVSFSKAEELYQKRVLTITGICVALLVVGIVCVVAYCKTKKQRKQMHNHLRQNMCPTHPNRSLANGPSHPQLDPEEIQMADYISKNVSATDHVIRRETETTFSGSHSCSPSHHCSTATPTSSQRQESHTWSLDRTESLTSDSQSGMMLSSVGTSKCNSPACVEARTRRGAAYCIEDSRRPTMQYRDSVDSLRDSPHSERYVSALTTPARLSPVDFHYSLAAQVPTFEITSPNSAHAVSLPPAAPVSFRVEEQQPLLRRYQVPFQDMQRYDSYYQRKSYLNESMGSLPSSPFCITEDDEYETTQEYIPALEQPKKTASSNRRWKRSKLNGQVSHRARTVRDSFSGSSASYSESDEELMAESTPFLSIENTEAMNTDSPPLYRPGDSRTYYSFERQSAQLESGQSRSTDTRPKQDSAPL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMH6</accession>
    <name>A0A452HMH6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Pro-neuregulin-2, membrane-bound isoform</fullName>
        </recommendedName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Direct ligand for ERBB3 and ERBB4 tyrosine kinase receptors. Concomitantly recruits ERBB1 and ERBB2 coreceptors, resulting in ligand-stimulated tyrosine phosphorylation and activation of the ERBB receptors. May also promote the heterodimerization with the EGF receptor.</text>
    </comment>
    <comment type="subunit">
        <text evidence="15">Interacts with ERBB3 and ERBB4.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the neuregulin family.</text>
    </comment>
    <comment type="caution">
        <text evidence="17">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMH6"/>
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        <property type="protein sequence ID" value="ENSGAGP00000016149.1"/>
        <property type="gene ID" value="ENSGAGG00000012118.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008083">
        <property type="term" value="F:growth factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048513">
        <property type="term" value="P:animal organ development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035556">
        <property type="term" value="P:intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
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        <property type="project" value="InterPro"/>
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        <property type="entry name" value="Neuregulin"/>
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        <property type="entry name" value="Neuregulin_C"/>
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    </dbReference>
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    </dbReference>
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    </dbReference>
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    </dbReference>
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    </dbReference>
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    </dbReference>
    <proteinExistence type="inferred from homology"/>
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    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9" id="KW-0339">Growth factor</keyword>
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    <keyword evidence="10 19" id="KW-0472">Membrane</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0964">Secreted</keyword>
    <keyword evidence="7 19" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 19" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="273"/>
            <end position="295"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="21">
        <location>
            <begin position="104"/>
            <end position="196"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="20">
        <location>
            <begin position="208"/>
            <end position="249"/>
        </location>
    </feature>
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            <end position="331"/>
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    </feature>
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        <location>
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    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="567"/>
            <end position="675"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="18">
        <location>
            <begin position="361"/>
            <end position="373"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="383"/>
            <end position="394"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="395"/>
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        </location>
    </feature>
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        <location>
            <begin position="599"/>
            <end position="608"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="620"/>
            <end position="634"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="650"/>
            <end position="663"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="664"/>
            <end position="675"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="220"/>
            <end position="237"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="239"/>
            <end position="248"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00008216"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00058865"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00065723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00068573"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016149.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="675" mass="75749" checksum="D12B85F176DE9EEB" modified="2019-05-08" version="1">SRCWEINSVSVSSSKGKRRCHSKKVSSSNGRIRKIHHELLPLRCVQPGLIQPLSHYRVSAAADQMLLLEPQQRETRLYNFPCPVPLTEAGFHTGLISLFTAVPPKLKKLKSPNVHVGEKISLKCEATTGNPQPSYKWYKDGKELKKSKDIRIKYGNGKKISRLQFNKVKLEDAGEYSCEAENVLGKDTAKGSLNVRSVATTLSSWSGHARKCNETAKSYCVNGGVCYYIEGINQLSCKCPNGFFGQRCLEKLPLRLYMPDPKQKAEELYQKRVLTITGICVALLVVGIVCVVAYCKTKKQRKQMHNHLRQNMCPTHPNRSLANGPSHPQLDPEEIQMADYISKNVSATDHVIRRETETTFSGSHSCSPSHHCSTATPTSSQRQESHTWSLDRTESLTSDSQSGMMLSSVGTSKCNSPACVEARTRRGAAYCIEDSRRPTMQYRDSVDSLRDSPHSERYVSALTTPARLSPVDFHYSLAAQVPTFEITSPNSAHAVSLPPAAPVSFRVEEQQPLLRRYQVPFQDMQRYDSYYQRKSYLNESMGSLPSSPFCITEDDEYETTQEYIPALEQPKKTASSNRRWKRSKLNGQVSHRARTVRDSFSGSSASYSESDEELMAESTPFLSIENTEAMNTDSPPLYRPGDSRTYYSFERQSAQLESGQSRSTDTRPKQDSAPL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMI0</accession>
    <name>A0A452HMI0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Serine/threonine-protein phosphatase</fullName>
            <ecNumber evidence="12">3.1.3.16</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="10">
            <text>O-phospho-L-seryl-[protein] + H2O = L-seryl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:20629"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="EC" id="3.1.3.16"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="11 12">
            <text>O-phospho-L-threonyl-[protein] + H2O = L-threonyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:47004"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="EC" id="3.1.3.16"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="2">
            <name>Fe(3+)</name>
            <dbReference type="ChEBI" id="CHEBI:29034"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the PPP phosphatase family. PP-2B subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.16" evidence="12"/>
    <dbReference type="AlphaFoldDB" id="A0A452HMI0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018449.1">
        <property type="protein sequence ID" value="ENSGAGP00000016157.1"/>
        <property type="gene ID" value="ENSGAGG00000012129.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033192">
        <property type="term" value="F:calmodulin-dependent protein phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097720">
        <property type="term" value="P:calcineurin-mediated signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd07416">
        <property type="entry name" value="MPP_PP2B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.60.21.10:FF:000002">
        <property type="entry name" value="Serine/threonine-protein phosphatase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.60.21.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004843">
        <property type="entry name" value="Calcineurin-like_PHP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029052">
        <property type="entry name" value="Metallo-depent_PP-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041751">
        <property type="entry name" value="MPP_PP2B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043360">
        <property type="entry name" value="PP2B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006186">
        <property type="entry name" value="Ser/Thr-sp_prot-phosphatase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45673">
        <property type="entry name" value="SERINE/THREONINE-PROTEIN PHOSPHATASE 2B CATALYTIC SUBUNIT 1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00149">
        <property type="entry name" value="Metallophos"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00114">
        <property type="entry name" value="STPHPHTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00156">
        <property type="entry name" value="PP2Ac"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56300">
        <property type="entry name" value="Metallo-dependent phosphatases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00125">
        <property type="entry name" value="SER_THR_PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="5 12" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="9" id="KW-0408">Iron</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="Serine/threonine specific protein phosphatases" evidence="14">
        <location>
            <begin position="144"/>
            <end position="149"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="464"/>
            <end position="501"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="13">
        <location>
            <begin position="464"/>
            <end position="478"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="13">
        <location>
            <begin position="485"/>
            <end position="494"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00001965"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00009905"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00047761"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00048336"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="RuleBase" id="RU004273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS00125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016157.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="501" mass="56886" checksum="EDF7876CF16E8B5F" modified="2019-05-08" version="1">MTGRGHNKLPTTERIVKAVPFPPTQRLTLKEVFEDGKPRLDVLKSHLVKEGRLEEDAALKIINDGAAILRHEKTMIEVEAPITVCGDIHGQFFDLMKLFEVGGSPNNTRYLFLGDYVDRGYFSIECVLYLWSLKINHPKTLFLLRGNHECRHLTEYFTFKQECRIKYSERVYDACMDTFDCLPLAALLNQQFLCVHGGLSPEITCLDDIRKLDRFKEPPAFGPMCDLLWSDPSEDYGNEKTLEHFTHNTVRGCSYFYSYPAVCEFLQNNSLLSVIRAHEAQDAGYRMYRKSQTTGFPSLITIFSAPNYLDVYNNKAAVLKYENNVMNIRQFNCSPHPYWLPNFMDVFTWSLPFVGEKVTEMLVNILNICSDDELISDGDETLEAGTTVRKEIIRNKIRAIGKMARVFSVLREESESVLTLKGLTPTGTLPLGVLSGGKQTLQTAIRGFSPQHKIRSFEEARGLDRINERMPPRKDSMHPDGPVNLVTSTNSAEKNGTGKKA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMI3</accession>
    <name>A0A452HMI3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Piwi-like protein 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="11">Belongs to the argonaute family. Piwi subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMI3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018442.1">
        <property type="protein sequence ID" value="ENSGAGP00000016151.1"/>
        <property type="gene ID" value="ENSGAGG00000012123.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016787">
        <property type="term" value="F:hydrolase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034584">
        <property type="term" value="F:piRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051321">
        <property type="term" value="P:meiotic cell cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006417">
        <property type="term" value="P:regulation of translation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031047">
        <property type="term" value="P:regulatory ncRNA-mediated gene silencing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd02845">
        <property type="entry name" value="PAZ_piwi_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04658">
        <property type="entry name" value="Piwi_piwi-like_Euk"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.2300:FF:000141">
        <property type="entry name" value="piwi-like protein 2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.420.10:FF:000014">
        <property type="entry name" value="Piwi-like RNA-mediated gene silencing 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.170.260.10:FF:000003">
        <property type="entry name" value="Piwi-like RNA-mediated gene silencing 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.2300">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.170.260.10">
        <property type="entry name" value="paz domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.420.10">
        <property type="entry name" value="Ribonuclease H-like superfamily/Ribonuclease H"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014811">
        <property type="entry name" value="ArgoL1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003100">
        <property type="entry name" value="PAZ_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036085">
        <property type="entry name" value="PAZ_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003165">
        <property type="entry name" value="Piwi"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012337">
        <property type="entry name" value="RNaseH-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036397">
        <property type="entry name" value="RNaseH_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22891">
        <property type="entry name" value="EUKARYOTIC TRANSLATION INITIATION FACTOR 2C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08699">
        <property type="entry name" value="ArgoL1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02170">
        <property type="entry name" value="PAZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02171">
        <property type="entry name" value="Piwi"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23278">
        <property type="entry name" value="Piwi_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01163">
        <property type="entry name" value="DUF1785"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00949">
        <property type="entry name" value="PAZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00950">
        <property type="entry name" value="Piwi"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF101690">
        <property type="entry name" value="PAZ domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53098">
        <property type="entry name" value="Ribonuclease H-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50821">
        <property type="entry name" value="PAZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50822">
        <property type="entry name" value="PIWI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="5" id="KW-0221">Differentiation</keyword>
    <keyword evidence="6" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="10" id="KW-0469">Meiosis</keyword>
    <keyword evidence="3" id="KW-0488">Methylation</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="9" id="KW-0943">RNA-mediated gene silencing</keyword>
    <keyword evidence="7" id="KW-0810">Translation regulation</keyword>
    <feature type="domain" description="PAZ" evidence="14">
        <location>
            <begin position="386"/>
            <end position="486"/>
        </location>
    </feature>
    <feature type="domain" description="Piwi" evidence="15">
        <location>
            <begin position="635"/>
            <end position="927"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022845"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023158"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023254"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00038291"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00039537"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50821"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50822"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016151.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="941" mass="107016" checksum="478D099016D6A3EC" modified="2019-05-08" version="1">MDSRRPPFQGTAPFCMPGLQPQLPQPEHLIRGRAVSLPCTTRMQRVPEEPRQPAGLIQQEPLPSVTTLHSMFQGLGFETIPRPALGRAVQPLGRGILGRGMGDMEAKPKAVMPVQPTFPSPNVMPVADSNVERMSLSPGRILGRGQMEAPGRGAMSLSRVLCKVPTDLTFLRTGVLQPQLSQPLLSPPRFAPQPESPPPTVVEQMERKEPLMKQGFKGTPFPLGLNLIKIHCKNEAVYQYHVTFSPNVECKSMRFGMMMEHRSVTGDVTAFDGSILYLPVKLPQTVDLTCQRKTDGAEISIKLQMTKILEPSSDLCIPFYNVVFRRVMRILDMKLVGRNFYDPTTPTMLQQYRLQIWPGYSASIRRTDGGLFLLADVSHKVIRNDSVLDIMHVIYQQRRETFQDECTKQLIGNIIITRYNNHTYRIDDIDWNKTPKDSFTMSDGKEITFIDYYSKNYGITVRELDQPLLIHRPNERRTPQGKDLTHQINLSPKQHHASLRNLLHRIEVNEAASKEMSRWGLCLDKDVHKTQGRVLPMEKINLRTSCFMTSEDLNWTKEITREAAISTIPMHYWALFYPKRVVDQARELVSMLEKVSGPIGMQLNQPAWVELKDDRIETYARTIKSILGTEGKMQMVVCIITGTRDDLYGAIKKLCCVQSPVPSQVINARTIMTQPNKLRSIAQKILLQINCKLGGELWGVDIPLKQLMVIGMDVYHDPSMGMRSVVGFIASTNHTLSKWYSRVVFQMPHQEIADSLKLCLVGALQKFYEVNHCLPEKIAIYRDGVSDTQLNTVVSYEIPQLQQCFKAFENYQPKMVVFVVQKKIGTNLYSASTEHFVAPTPGIVIDHTVTSRDWVDFYLLSHRVRQGCGIPTHYICVLNTANLSPDHMQRLTFKLCHMYWNWPGTIRVPAPCKYAHKLAFLAGQFLHHEPAIQLCERLFFL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMI7</accession>
    <name>A0A452HMI7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Fibroblast growth factor</fullName>
            <shortName evidence="5">FGF</shortName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1 5">Belongs to the heparin-binding growth factors family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMI7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018500.1">
        <property type="protein sequence ID" value="ENSGAGP00000016205.1"/>
        <property type="gene ID" value="ENSGAGG00000012157.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008083">
        <property type="term" value="F:growth factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051781">
        <property type="term" value="P:positive regulation of cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd23318">
        <property type="entry name" value="beta-trefoil_FGF6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.80.10.50">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002209">
        <property type="entry name" value="Fibroblast_GF_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008996">
        <property type="entry name" value="IL1/FGF"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11486">
        <property type="entry name" value="FIBROBLAST GROWTH FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00167">
        <property type="entry name" value="FGF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00263">
        <property type="entry name" value="HBGFFGF"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00262">
        <property type="entry name" value="IL1HBGF"/>
    </dbReference>
    <dbReference type="SMART" id="SM00442">
        <property type="entry name" value="FGF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50353">
        <property type="entry name" value="Cytokine"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00247">
        <property type="entry name" value="HBGF_FGF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="3" id="KW-0221">Differentiation</keyword>
    <keyword evidence="4" id="KW-0497">Mitogen</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007936"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="RuleBase" id="RU049442"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016205.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="201" mass="22260" checksum="0BDB7FA8FCD0CA12" modified="2025-10-08" version="2">MSSPNIALISSVHWTLPAFVLLGFLAGIVSSYPVVSRTNGTLLERGWESLLSRSIAGMSGEKSDVNWESDYLLGIKRQRRLYCNVGIGFHLQILPDGRISGVHNENQYSLLEISTVERGVVSLFGVKSALFVAMNNKGRLYGTAIFQDECKFRETLLPNNYNAYESSAYQGAYIALSKHGRVKKGNKVSPAMTVTHFLPRI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMI9</accession>
    <name>A0A452HMI9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Serine/threonine-protein phosphatase</fullName>
            <ecNumber evidence="12">3.1.3.16</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="10">
            <text>O-phospho-L-seryl-[protein] + H2O = L-seryl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:20629"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="EC" id="3.1.3.16"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="11 12">
            <text>O-phospho-L-threonyl-[protein] + H2O = L-threonyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:47004"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="EC" id="3.1.3.16"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="2">
            <name>Fe(3+)</name>
            <dbReference type="ChEBI" id="CHEBI:29034"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the PPP phosphatase family. PP-2B subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.16" evidence="12"/>
    <dbReference type="AlphaFoldDB" id="A0A452HMI9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018455.1">
        <property type="protein sequence ID" value="ENSGAGP00000016163.1"/>
        <property type="gene ID" value="ENSGAGG00000012129.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033192">
        <property type="term" value="F:calmodulin-dependent protein phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097720">
        <property type="term" value="P:calcineurin-mediated signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd07416">
        <property type="entry name" value="MPP_PP2B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.60.21.10:FF:000002">
        <property type="entry name" value="Serine/threonine-protein phosphatase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.60.21.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004843">
        <property type="entry name" value="Calcineurin-like_PHP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029052">
        <property type="entry name" value="Metallo-depent_PP-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041751">
        <property type="entry name" value="MPP_PP2B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043360">
        <property type="entry name" value="PP2B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006186">
        <property type="entry name" value="Ser/Thr-sp_prot-phosphatase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45673">
        <property type="entry name" value="SERINE/THREONINE-PROTEIN PHOSPHATASE 2B CATALYTIC SUBUNIT 1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00149">
        <property type="entry name" value="Metallophos"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00114">
        <property type="entry name" value="STPHPHTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00156">
        <property type="entry name" value="PP2Ac"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56300">
        <property type="entry name" value="Metallo-dependent phosphatases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00125">
        <property type="entry name" value="SER_THR_PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="5 12" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="9" id="KW-0408">Iron</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="Serine/threonine specific protein phosphatases" evidence="14">
        <location>
            <begin position="144"/>
            <end position="149"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="474"/>
            <end position="511"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="13">
        <location>
            <begin position="474"/>
            <end position="488"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="13">
        <location>
            <begin position="495"/>
            <end position="504"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00001965"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00009905"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00047761"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00048336"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="RuleBase" id="RU004273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS00125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016163.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="511" mass="57942" checksum="27388E45A9D83234" modified="2019-05-08" version="1">MTGRGHNKLPTTERIVKAVPFPPTQRLTLKEVFEDGKPRLDVLKSHLVKEGRLEEDAALKIINDGAAILRHEKTMIEVEAPITVCGDIHGQFFDLMKLFEVGGSPNNTRYLFLGDYVDRGYFSIECVLYLWSLKINHPKTLFLLRGNHECRHLTEYFTFKQECRIKYSERVYDACMDTFDCLPLAALLNQQFLCVHGGLSPEITCLDDIRKLDRFKEPPAFGPMCDLLWSDPSEDYGNEKTLEHFTHNTVRGCSYFYSYPAVCEFLQNNSLLSVIRAHEAQDAGYRMYRKSQTTGFPSLITIFSAPNYLDVYNNKAAVLKYENNVMNIRQFNCSPHPYWLPNFMDVFTWSLPFVGEKVTEMLVNILNICSDDELISDGDETLEAGTTVRKEIIRNKIRAIGKMARVFSVLREESESVLTLKGLTPTGTLPLGVLSGGKQTLQTATVEAVEAREAIRGFSPQHKIRSFEEARGLDRINERMPPRKDSMHPDGPVNLVTSTNSAEKNGTGKKA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMJ2</accession>
    <name>A0A452HMJ2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Slit homolog 2 protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="15">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018470.1">
        <property type="protein sequence ID" value="ENSGAGP00000016176.1"/>
        <property type="gene ID" value="ENSGAGG00000011388.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000018506.1">
        <property type="protein sequence ID" value="ENSGAGP00000016210.1"/>
        <property type="gene ID" value="ENSGAGG00000011388.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048495">
        <property type="term" value="F:Roundabout binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072359">
        <property type="term" value="P:circulatory system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060560">
        <property type="term" value="P:developmental growth involved in morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060562">
        <property type="term" value="P:epithelial tube morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001822">
        <property type="term" value="P:kidney development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050919">
        <property type="term" value="P:negative chemotaxis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030308">
        <property type="term" value="P:negative regulation of cell growth"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010605">
        <property type="term" value="P:negative regulation of macromolecule metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010977">
        <property type="term" value="P:negative regulation of neuron projection development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0021772">
        <property type="term" value="P:olfactory bulb development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050770">
        <property type="term" value="P:regulation of axonogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032535">
        <property type="term" value="P:regulation of cellular component size"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050767">
        <property type="term" value="P:regulation of neurogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031290">
        <property type="term" value="P:retinal ganglion cell axon guidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0022029">
        <property type="term" value="P:telencephalon cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd00054">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="CDD" id="cd00110">
        <property type="entry name" value="LamG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000045">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000053">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000054">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000062">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000063">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000099">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000289">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.200:FF:000013">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000002">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000004">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.200">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000483">
        <property type="entry name" value="Cys-rich_flank_reg_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006207">
        <property type="entry name" value="Cys_knot_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001881">
        <property type="entry name" value="EGF-like_Ca-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000152">
        <property type="entry name" value="EGF-type_Asp/Asn_hydroxyl_site"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018097">
        <property type="entry name" value="EGF_Ca-bd_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003645">
        <property type="entry name" value="Fol_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001791">
        <property type="entry name" value="Laminin_G"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001611">
        <property type="entry name" value="Leu-rich_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003591">
        <property type="entry name" value="Leu-rich_rpt_typical-subtyp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000372">
        <property type="entry name" value="LRRNT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051355">
        <property type="entry name" value="Notch/Slit_guidance"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45836">
        <property type="entry name" value="SLIT HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45836:SF2">
        <property type="entry name" value="SLIT HOMOLOG 2 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00008">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00054">
        <property type="entry name" value="Laminin_G_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00560">
        <property type="entry name" value="LRR_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13855">
        <property type="entry name" value="LRR_8"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01463">
        <property type="entry name" value="LRRCT"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01462">
        <property type="entry name" value="LRRNT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00041">
        <property type="entry name" value="CT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="SMART" id="SM00179">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SMART" id="SM00274">
        <property type="entry name" value="FOLN"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00282">
        <property type="entry name" value="LamG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00368">
        <property type="entry name" value="LRR_RI"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SMART" id="SM00365">
        <property type="entry name" value="LRR_SD22"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SMART" id="SM00369">
        <property type="entry name" value="LRR_TYP"/>
        <property type="match status" value="12"/>
    </dbReference>
    <dbReference type="SMART" id="SM00082">
        <property type="entry name" value="LRRCT"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00013">
        <property type="entry name" value="LRRNT"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52058">
        <property type="entry name" value="L domain-like"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00010">
        <property type="entry name" value="ASX_HYDROXYL"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01185">
        <property type="entry name" value="CTCK_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01225">
        <property type="entry name" value="CTCK_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01186">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01187">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50025">
        <property type="entry name" value="LAM_G_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51450">
        <property type="entry name" value="LRR"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0145">Chemotaxis</keyword>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="10" id="KW-0221">Differentiation</keyword>
    <keyword evidence="12 15" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5 15" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7" id="KW-0358">Heparin-binding</keyword>
    <keyword evidence="6" id="KW-0433">Leucine-rich repeat</keyword>
    <keyword evidence="11" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="8" id="KW-0732">Signal</keyword>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="727"/>
            <end position="762"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="764"/>
            <end position="803"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="805"/>
            <end position="841"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="852"/>
            <end position="890"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="892"/>
            <end position="928"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="937"/>
            <end position="973"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin G" evidence="17">
        <location>
            <begin position="976"/>
            <end position="1149"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="1153"/>
            <end position="1184"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="1187"/>
            <end position="1223"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="1228"/>
            <end position="1264"/>
        </location>
    </feature>
    <feature type="domain" description="CTCK" evidence="16">
        <location>
            <begin position="1269"/>
            <end position="1344"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="752"/>
            <end position="761"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="793"/>
            <end position="802"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="880"/>
            <end position="889"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="918"/>
            <end position="927"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="963"/>
            <end position="972"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="1174"/>
            <end position="1183"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="1191"/>
            <end position="1201"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="1213"/>
            <end position="1222"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="1232"/>
            <end position="1242"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="1254"/>
            <end position="1263"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022500"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022614"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022674"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00074955"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS01225"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50025"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016210.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1345" mass="148958" checksum="227DF2017CCC0775" modified="2019-05-08" version="1">MPKLRTFRLHSNNLYCDCHLAWLSDWLRQRPRVGLYTQCMGPSHLRGHNVAEVQKREFVCSDEEEGHQSFMAPSCSVLHCPTACTCSNNIVDCRGKGLTEIPTNLPETITEIRLEQNSIKVIPPGAFSPYKKLRRIDLSNNQISETAPDAFQGLRSLNSLVLYGNKITELPKGLFEGLFSLQLLLLNANKINCLRVDAFQDLHNLNLLSLYDNKLQTIAKGTFSPLRSIQTLHLAQNPFICDCHLKWLADYLHANPIETSGARCTSPRRLANKRIGQIKSKKFRCSAKEQYFIPGTEDYRSKLSGDCFADLACPEKCRCEGTTVDCSNQKLNKIPEHVPQYTAELRLNNNEFTVLEATGIFKKLPQLRKINLSNNKITDIEEGAFEGASGVNELLLTSNRLETIRHKMFKGLESLKTLMLRSNRISCIGNDSFTGLSSVRLLSLYDNQITTVAPGAFDTLHSLSTLNLLANPFNCNCHLAWLGDWLRKKRIVTGNPRCQKPYFLKEIPIQDVAIQDFTCDDGNDDNSCSPLSRCPAECTCLDTVVRCSNKGLKVLPKGIPKDVTELYLDGNQFTLVPKELSNYKHLTLIDLSNNRISTLSNQSFSNMTQLLTLILSYNRLRCIPARTFDGLKSLRLLSLHGNDISVVPEGAFNDLSALSHLAIGANPLYCDCNMQWLSDWVKSEYKEPGIARCAGPGEMADKLLLTTPSKKFTCQGPVDVNILAKCSPCLSNPCKNDGTCNNDPVDFYTCTCPYGFKGQDCDIPIHACISNPCQHGGTCHLKEGEKDGFWCTCADGFEGENCEVNVDDCEDNDCENNSTCVDGINNYTCLCPPEYTAANLNEVEKGELCEEKLDFCAQDLNPCQHDSKCILTLKGYKCDCTPGYVGEHCDMDYDDCQDNKCKNGAQCTDAVNGYTCICPEGYSGLFCEFSPPVVLPRTSPCDNYECQNGAQCIVRVNEPICQCLSGYQGDKCEKLVSVNFVSKESYLQIPSSKVRPQTNITLQIATDEDSGILLYKGDRDHIAMELYRGRVRVSYDTGSYPASAIYSVETINDGNFHIVELLAMDQILSLSVDGGSPKIITNLSKQSTLNVDSPLYVGGMPVKNNVAALRQSPGQNGSSFHGCIRNLYINSELQDFRNVPLQVGILPGCEPCHKKVCVHGTCHAMSQSGFTCECEGGWTGPLCDQQTNDPCLGNKCVHGTCLPINAFSYSCKCLQGHGGVLCDEEEELFNPCQSIKCKHGKCRLSGLGKPYCECSSGYTGDSCDKEISCRGERIRDYYQKQQGYAACQTTKKVSRLECKGGCSTGQCCGPLRSKRRKYSFECTDGSSFVDEVEKVVKCGCTKCPS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMJ4</accession>
    <name>A0A452HMJ4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Calponin-homology (CH) domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HMJ4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018462.1">
        <property type="protein sequence ID" value="ENSGAGP00000016169.1"/>
        <property type="gene ID" value="ENSGAGG00000012125.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd21271">
        <property type="entry name" value="CH_LRCH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.418.10:FF:000021">
        <property type="entry name" value="Leucine-rich repeat and calponin homology domain-containing protein 1 isoform 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000007">
        <property type="entry name" value="Leucine-rich repeat and calponin homology domain-containing protein 1 isoform 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.418.10">
        <property type="entry name" value="Calponin-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001715">
        <property type="entry name" value="CH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036872">
        <property type="entry name" value="CH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001611">
        <property type="entry name" value="Leu-rich_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003591">
        <property type="entry name" value="Leu-rich_rpt_typical-subtyp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050216">
        <property type="entry name" value="LRR_domain-containing"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48051">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48051:SF38">
        <property type="entry name" value="LEUCINE RICH REPEATS AND CALPONIN HOMOLOGY DOMAIN CONTAINING 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00307">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13855">
        <property type="entry name" value="LRR_8"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00033">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00364">
        <property type="entry name" value="LRR_BAC"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00369">
        <property type="entry name" value="LRR_TYP"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47576">
        <property type="entry name" value="Calponin-homology domain, CH-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52058">
        <property type="entry name" value="L domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50021">
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51450">
        <property type="entry name" value="LRR"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0433">Leucine-rich repeat</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="Calponin-homology (CH)" evidence="4">
        <location>
            <begin position="455"/>
            <end position="568"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="244"/>
            <end position="277"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022614"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016169.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="576" mass="65185" checksum="3879FB7AB013B8E1" modified="2019-05-08" version="1">GGWISWPGGWGLQPGSGGERVQASGNVLFLFLSSDLSKNRLTEVPMELCHFVSLETLNLYHNCIKIIPDAIVNLQMLTYLNLSRNQLSSLPACLCGLPLKVLIASNNRLGSLPEEIGQLKQLMELDVSCNEITALPQQIGQLKSLRELNIRRNYLKVLPQELIELPLVKFDFSCNKVLVIPICFRKMLQLQVLLLENNPLQSPPAQICIKGKVHIFKYLSIQACQIKTADSLYLNAIERPHLQQPVEESKKDSDSGVGSDNGDKRLSATEPSDEDTVCLNVPMSNIMEEDQIIKEDSGHHINLVQEFHQGPSLMIHIEKAREAGNQFAEGSQPLTTEFMSYIKATAECDELLRIEEDTHWQTEGITNTSNKQNIDIAMKDNLKEAVDLLQDPNRYVHIRGFSDVNNMLYRKFCEVFDGLFLLNISLFKITVFLRPQRNLESIDPQFTIRRKMEQMREERELVEQLRENIEMRLKICLPEDLGSALMDGVVLCHLVNHIRPRSVGSIHVPSPAVPKLSMAKCRRNVENFLDACRKLGIPEEKLCLPHHILEEKGLVKVSITVQALLDVTTMKQALFT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMJ5</accession>
    <name>A0A452HMJ5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Calponin-homology (CH) domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HMJ5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018465.1">
        <property type="protein sequence ID" value="ENSGAGP00000016172.1"/>
        <property type="gene ID" value="ENSGAGG00000012125.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd21271">
        <property type="entry name" value="CH_LRCH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.418.10:FF:000021">
        <property type="entry name" value="Leucine-rich repeat and calponin homology domain-containing protein 1 isoform 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000067">
        <property type="entry name" value="Leucine-rich repeat and calponin homology domain-containing protein 4 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.418.10">
        <property type="entry name" value="Calponin-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001715">
        <property type="entry name" value="CH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036872">
        <property type="entry name" value="CH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001611">
        <property type="entry name" value="Leu-rich_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003591">
        <property type="entry name" value="Leu-rich_rpt_typical-subtyp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050216">
        <property type="entry name" value="LRR_domain-containing"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48051">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48051:SF38">
        <property type="entry name" value="LEUCINE RICH REPEATS AND CALPONIN HOMOLOGY DOMAIN CONTAINING 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00307">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13855">
        <property type="entry name" value="LRR_8"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00033">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00364">
        <property type="entry name" value="LRR_BAC"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00369">
        <property type="entry name" value="LRR_TYP"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47576">
        <property type="entry name" value="Calponin-homology domain, CH-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52058">
        <property type="entry name" value="L domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50021">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51450">
        <property type="entry name" value="LRR"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0433">Leucine-rich repeat</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="Calponin-homology (CH)" evidence="4">
        <location>
            <begin position="360"/>
            <end position="473"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="199"/>
            <end position="231"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022614"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016172.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="481" mass="54681" checksum="1DE887466C39E7D8" modified="2019-05-08" version="1">MELCHFVSLETLNLYHNCIKIIPDAIVNLQMLTYLNLSRNQLSSLPACLCGLPLKVLIASNNRLGSLPEEIGQLKQLMELDVSCNEITALPQQIGQLKSLRELNIRRNYLKVLPQELIELPLVKFDFSCNKVLVIPICFRKMLQLQVLLLENNPLQSPPAQICIKGKVHIFKYLSIQACQIKTADSLYLNAIERPHLQQPVEESKKDSDSGVGSDNGDKRLSATEPSDEDTVCLNVPMSNIMEEDQIIKEDSGHHINLVQGLNVKKRTLTVTGSLPTILFVMLYECNSQKTSQPKFLLTTWLKPTPSVYHCFSWKIYEAFFDGLFLLNISLFKITVFLRPQRNLESIDPQFTIRRKMEQMREERELVEQLRENIEMRLKICLPEDLGSALMDGVVLCHLVNHIRPRSVGSIHVPSPAVPKLSMAKCRRNVENFLDACRKLGIPEEKLCLPHHILEEKGLVKVSITVQALLDVTTMKQALFT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMJ6</accession>
    <name>A0A452HMJ6_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="9">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell junction</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMJ6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018461.1">
        <property type="protein sequence ID" value="ENSGAGP00000016168.1"/>
        <property type="gene ID" value="ENSGAGG00000012137.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070161">
        <property type="term" value="C:anchoring junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="CDD" id="cd11921">
        <property type="entry name" value="SH3_Vinexin_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd11924">
        <property type="entry name" value="SH3_Vinexin_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd11918">
        <property type="entry name" value="SH3_Vinexin_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.40:FF:000001">
        <property type="entry name" value="Sorbin and SH3 domain-containing protein 1 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050384">
        <property type="entry name" value="Endophilin_SH3RF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003127">
        <property type="entry name" value="SoHo_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035609">
        <property type="entry name" value="Vinexin_SH3_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035608">
        <property type="entry name" value="Vinexin_SH3_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035607">
        <property type="entry name" value="Vinexin_SH3_3"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14167">
        <property type="entry name" value="SH3 DOMAIN-CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14167:SF54">
        <property type="entry name" value="VINEXIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00018">
        <property type="entry name" value="SH3_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14604">
        <property type="entry name" value="SH3_9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02208">
        <property type="entry name" value="Sorb"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00499">
        <property type="entry name" value="P67PHOX"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00452">
        <property type="entry name" value="SH3DOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00326">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00459">
        <property type="entry name" value="Sorb"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50831">
        <property type="entry name" value="SOHO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0965">Cell junction</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="2 5" id="KW-0728">SH3 domain</keyword>
    <feature type="domain" description="SoHo" evidence="8">
        <location>
            <begin position="119"/>
            <end position="183"/>
        </location>
    </feature>
    <feature type="domain" description="SH3" evidence="7">
        <location>
            <begin position="398"/>
            <end position="457"/>
        </location>
    </feature>
    <feature type="domain" description="SH3" evidence="7">
        <location>
            <begin position="472"/>
            <end position="533"/>
        </location>
    </feature>
    <feature type="domain" description="SH3" evidence="7">
        <location>
            <begin position="703"/>
            <end position="762"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="120"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="327"/>
            <end position="397"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="540"/>
            <end position="604"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="619"/>
            <end position="702"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="567"/>
            <end position="579"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="585"/>
            <end position="601"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="619"/>
            <end position="631"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="632"/>
            <end position="646"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="6">
        <location>
            <begin position="647"/>
            <end position="663"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="664"/>
            <end position="675"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004282"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50831"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016168.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="762" mass="82760" checksum="6F4B02C68994430D" modified="2019-05-08" version="1">MAFQAAGSGTLQPRSVQPLEPLLHGWGPEPAVQKGFLGNFSLPTPLEGTLRAAGNPHLRERHGAHSAGAGDSGCFPHSLSSPPPYAMAGGCLNPQGSQGLSLKDFVPSHLQKREFLPAARSPAAPMATSPQPSTWVPLRHRGPRSVDRPRDWYRSMFQQIHRKLPEPQLDWDFHHCTEPRPPAFAQKGRMPSGSSYLQNGLDWRGQEAADDAAMEPRSIFDYEPGKSSVLDHPNPVAEPQVGRAGSWHQFLKELEMGSLPKKPLAPFPVEPPPPSQPIEVLLERELQQLSEELDKDMKALETHRHPCKTSSASLCACTAARSLPSACRSHPALGQSPPASRRPPASPSMERAGLGVGTSDWSRSPPRKDPPGHSGKSPLADESESIEAVDGPVRREDKKMKAARVKFDFQAESPKELTMQKGDIVYIHKEVDRNWLEGEHYGRVGIFPSNYVEILSPTEVPKPIKPPTIQVLEYGEALAQYNFKGDLAVELSFRKGERICLVRKVDENWYEGRISGTNRQGIFPANYVQVLKEPRVKASEEFPASPNLTAPASPRLLAHTGSPAQLHSHGSPRSNPSSSFLAGSPTAQQRGTEGGALSSSPHHFGFTFPVSPKLQRAGVSSSLLQGPQNPQLAPSTTLLSPVSLSPSPQPLGPSRPQETPPTAAPSSLPGQSSTLEAPVPAGCHLGSAGHLPKAPPSDNGSSIQWTPFRAIYQYRPQNEDELELHEGDRVDVMQQCDDGWFVGVSRRTQKFGTFPGNYVAPV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMM1</accession>
    <name>A0A452HMM1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Early growth response protein 3</fullName>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="11">Probable transcription factor involved in muscle spindle development.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 14">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 14">Belongs to the EGR C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMM1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018487.1">
        <property type="protein sequence ID" value="ENSGAGP00000016193.1"/>
        <property type="gene ID" value="ENSGAGG00000012151.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005654">
        <property type="term" value="C:nucleoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048731">
        <property type="term" value="P:system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000392">
        <property type="entry name" value="early growth response protein 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000324">
        <property type="entry name" value="Early growth response protein 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000419">
        <property type="entry name" value="Early growth response protein 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021849">
        <property type="entry name" value="EGR_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23235:SF78">
        <property type="entry name" value="EARLY GROWTH RESPONSE PROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23235">
        <property type="entry name" value="KRUEPPEL-LIKE TRANSCRIPTION FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11928">
        <property type="entry name" value="DUF3446"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 14" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3 14" id="KW-0479">Metal-binding</keyword>
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    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="16" id="KW-0732">Signal</keyword>
    <keyword evidence="9 14" id="KW-0804">Transcription</keyword>
    <keyword evidence="7 14" id="KW-0805">Transcription regulation</keyword>
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            <begin position="1"/>
            <end position="17"/>
        </location>
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        </location>
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            <end position="315"/>
        </location>
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        </location>
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            <begin position="239"/>
            <end position="266"/>
        </location>
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        <location>
            <begin position="331"/>
            <end position="369"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="15">
        <location>
            <begin position="252"/>
            <end position="266"/>
        </location>
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        <location>
            <begin position="333"/>
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        </location>
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        <location>
            <begin position="350"/>
            <end position="369"/>
        </location>
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            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
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        <source>
            <dbReference type="ARBA" id="ARBA00005682"/>
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        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
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        </source>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016193.1"/>
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    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
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    </evidence>
    <sequence length="369" mass="41029" checksum="81477E32FB96837E" modified="2019-05-08" version="1">MCFYAFAVSAAAGFSFASRGCSGEWKLPDREVAENVMDIGLANEKTNQELSSYSGSFQPTPGNKTVTYLGKFAFDSPSNWCQDNIISLMSAGILGVPPSSSAITSTQTSTASMVQNQGEVDQMYPALPPYSTCSDLYPEPVSFHDPQSNPGLTYSPQDYQAAKPALDSNLFPMIPDYNLYHHPNEMGTLTEHKPYQSLDPIRVNPPPITPLETIKAFKDKQIHPSFGSLQQQPLTLKPIRPRKYPNRPSKTPLHERPHACPAEGCDRRFSRSDELTRHLRIHTGHKPFQCRICMRSFSRSDHLTTHIRTHTGEKPFACEFCGRKFARSDERKRHAKIHLKQKEKKADKGSSSSSSSPPVSLAPVVTTCA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMM2</accession>
    <name>A0A452HMM2_9SAUR</name>
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        <lineage>
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            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
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        </lineage>
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            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
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                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
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        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
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    </evidence>
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        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
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    </evidence>
    <sequence length="323" mass="36629" checksum="2FAC31665C261BC4" modified="2019-05-08" version="1">SEELLYIKGYSCTYHPGVPVFHDALKGWSCCKRRTTDFSDFLSIAGCTKGLHNSEKPPEPVKPEVKTTSERKELAELKPKFQEHIIRAPKPMETINRPSPDEPMTSLQLKVSASLKQALDKLKLASENEEEKKEEDSDEIKVGTSCKNGGCLKTFEGPQSTEEICIYHSGVPIFHEGMKYWSCCKRKTSDFNTFLSQEGCTTGTHVWTKKDAGKKVVPCRHDWHQTGGEVTISIYAKNSLPELSRVEANSTMLNIHIIFEGDKEFDRNVKLWGVIDVKRSYVNMTATKIELTMRKAEPMLWASLELPIPETQQKQNEDITDQE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMM6</accession>
    <name>A0A452HMM6_9SAUR</name>
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        <recommendedName>
            <fullName evidence="4">Cysteine and histidine-rich domain-containing protein 1</fullName>
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        </alternativeName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
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            <taxon>Eukaryota</taxon>
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            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HMM6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018493.1">
        <property type="protein sequence ID" value="ENSGAGP00000016198.1"/>
        <property type="gene ID" value="ENSGAGG00000012138.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
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    </dbReference>
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    </dbReference>
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        <property type="entry name" value="cysteine and histidine-rich domain-containing protein 1"/>
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    </dbReference>
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    </dbReference>
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    </dbReference>
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        <property type="entry name" value="CHORD_dom"/>
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        <property type="entry name" value="CHRD1"/>
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        <property type="entry name" value="CS_dom"/>
    </dbReference>
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        <property type="entry name" value="HSP20-like_chaperone"/>
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        <property type="entry name" value="CYSTEINE AND HISTIDINE-RICH DOMAIN-CONTAINING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46983:SF4">
        <property type="entry name" value="CYSTEINE AND HISTIDINE-RICH DOMAIN-CONTAINING PROTEIN 1"/>
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    </dbReference>
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    </dbReference>
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    </dbReference>
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    </dbReference>
    <proteinExistence type="predicted"/>
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    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="CHORD" evidence="8">
        <location>
            <begin position="1"/>
            <end position="56"/>
        </location>
    </feature>
    <feature type="domain" description="CHORD" evidence="8">
        <location>
            <begin position="150"/>
            <end position="209"/>
        </location>
    </feature>
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        <location>
            <begin position="220"/>
            <end position="309"/>
        </location>
    </feature>
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        <location>
            <begin position="116"/>
            <end position="143"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00034547"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00034558"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS51203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51401"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016198.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="327" mass="36949" checksum="F28DE852513075B9" modified="2019-05-08" version="1">SALLCADKASTEYYSCTYHPGVPVFHDALKGWSCCKRRTTDFSDFLSIAGCTKGLHNSEKPPEPVKPEVKTTSERKELAELKPKFQEHIIRAPKPMETINRPSPDEPMTSLQLKVSASLKQALDKLKLASENEEEKKEEDSDEIKVGTSCKNGGCLKTFEGPQSTEEICIYHSGVPIFHEGMKYWSCCKRKTSDFNTFLSQEGCTTGTHVWTKKDAGKKVVPCRHDWHQTGGEVTISIYAKNSLPELSRVEANSTMLNIHIIFEGDKEFDRNVKLWGVIDVKRSYVNMTATKIELTMRKAEPMLWASLELPIPETQQKQNEDITDQE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPF3</accession>
    <name>A0A452IPF3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Rap guanine nucleotide exchange factor 6</fullName>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the RAPGEF2 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000034050.1">
        <property type="protein sequence ID" value="ENSGAGP00000029982.1"/>
        <property type="gene ID" value="ENSGAGG00000021645.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007264">
        <property type="term" value="P:small GTPase-mediated signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00038">
        <property type="entry name" value="CAP_ED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd06755">
        <property type="entry name" value="PDZ_RapGEF2_RapGEF6-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd01785">
        <property type="entry name" value="RA_PDZ-GEF1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00155">
        <property type="entry name" value="RasGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd06224">
        <property type="entry name" value="REM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.840.10:FF:000001">
        <property type="entry name" value="Rap guanine nucleotide exchange factor (GEF) 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000024">
        <property type="entry name" value="rap guanine nucleotide exchange factor 2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.870.10:FF:000001">
        <property type="entry name" value="rap guanine nucleotide exchange factor 2 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.10:FF:000019">
        <property type="entry name" value="rap guanine nucleotide exchange factor 6 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.10:FF:000040">
        <property type="entry name" value="rap guanine nucleotide exchange factor 6 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.42.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.10">
        <property type="entry name" value="Jelly Rolls"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.840.10">
        <property type="entry name" value="Ras guanine-nucleotide exchange factors catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.870.10">
        <property type="entry name" value="Son of sevenless (SoS) protein Chain: S domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000595">
        <property type="entry name" value="cNMP-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018490">
        <property type="entry name" value="cNMP-bd_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001478">
        <property type="entry name" value="PDZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036034">
        <property type="entry name" value="PDZ_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000159">
        <property type="entry name" value="RA_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000651">
        <property type="entry name" value="Ras-like_Gua-exchang_fac_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023578">
        <property type="entry name" value="Ras_GEF_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001895">
        <property type="entry name" value="RASGEF_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036964">
        <property type="entry name" value="RASGEF_cat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014710">
        <property type="entry name" value="RmlC-like_jellyroll"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029071">
        <property type="entry name" value="Ubiquitin-like_domsf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45161">
        <property type="entry name" value="CYTOSKELETON-ASSOCIATED PROTEIN 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45161:SF4">
        <property type="entry name" value="RAP GUANINE NUCLEOTIDE EXCHANGE FACTOR 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00027">
        <property type="entry name" value="cNMP_binding"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00595">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00788">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00617">
        <property type="entry name" value="RasGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00618">
        <property type="entry name" value="RasGEF_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00100">
        <property type="entry name" value="cNMP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00228">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00314">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00147">
        <property type="entry name" value="RasGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00229">
        <property type="entry name" value="RasGEFN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51206">
        <property type="entry name" value="cAMP-binding domain-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50156">
        <property type="entry name" value="PDZ domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48366">
        <property type="entry name" value="Ras GEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54236">
        <property type="entry name" value="Ubiquitin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50042">
        <property type="entry name" value="CNMP_BINDING_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50106">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50200">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50009">
        <property type="entry name" value="RASGEF_CAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50212">
        <property type="entry name" value="RASGEF_NTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7 9" id="KW-0344">Guanine-nucleotide releasing factor</keyword>
    <keyword evidence="8" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Cyclic nucleotide-binding" evidence="12">
        <location>
            <begin position="281"/>
            <end position="381"/>
        </location>
    </feature>
    <feature type="domain" description="N-terminal Ras-GEF" evidence="15">
        <location>
            <begin position="413"/>
            <end position="526"/>
        </location>
    </feature>
    <feature type="domain" description="PDZ" evidence="13">
        <location>
            <begin position="531"/>
            <end position="601"/>
        </location>
    </feature>
    <feature type="domain" description="Ras-associating" evidence="14">
        <location>
            <begin position="756"/>
            <end position="842"/>
        </location>
    </feature>
    <feature type="domain" description="Ras-GEF" evidence="11">
        <location>
            <begin position="867"/>
            <end position="1094"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="173"/>
            <end position="245"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1081"/>
            <end position="1107"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1212"/>
            <end position="1295"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1478"/>
            <end position="1499"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1512"/>
            <end position="1555"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="188"/>
            <end position="205"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="10">
        <location>
            <begin position="228"/>
            <end position="240"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="1082"/>
            <end position="1094"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="1241"/>
            <end position="1252"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="1266"/>
            <end position="1275"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="1276"/>
            <end position="1293"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="1489"/>
            <end position="1499"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="1534"/>
            <end position="1543"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00010829"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022658"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00168"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50009"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50106"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50200"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029982.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="18">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1555" mass="174828" checksum="BC0FD0C8C6EB3532" modified="2019-05-08" version="1">MTSPVDPKIKQALRKKTSERTQEDLNTIYSYLHGMDILSHLREHQLRIMSSGARYERYKGNQVLFCSDTIARCWYILLSGSVLMKDSMFLPPCSFGKQSGGKRGCECIILEPSEMIVVENSKDNEENILQREVPLRQSRRRFRKINQRGERQTITDNVDASSYLSLPADLTKMHLTDNPHPQVTHVPSSQSGCSIASDSGSSSLSDIYQATESDMGDVDLTGLPEAPVDSEDEEEEDEDIDRTSDPLLGRDVVRECLEKEPADKTDDDIEQLLEFMHQLPAFANMTMSVRRELCSVMIFEVVEQAGAIILEDGQELDSWYVILNGTVEISHPDGKTESLCMGNSFGINPSLDKQYMNGVVRTKVDDCQFVCIAQQDYWRILNHVEKNTHKVEEEGEIVMVKEHRELDRSGTRKGHIVIKATPERLIMHLIEEHSIVDPTYIEDFLLTYRTFLSSPLEVGDKLLEWFKVDSLRDKVTRVVLLWVNNHFNDFEGDPAMTRFLEEFEKNLEDTKMKGHLRLLNIACAAKAKWRQITLQKASRDSPLFFCLVGGSDKGFGIFVETVETGSKAAEAGLKRGDQIMEVNGQNFENITFVKAVEILRNNTHLSITVKTNIFVFKELLSRIGQEKNGVPHIPKIAEKKSNRYSIPDMPGDMEQMFPCEKGNKKIKANTVSGGRNRIRKILDKTRFSILPPKLFSDGSVSQSQDDSIVGTRQCRHSLAIMPIPGSLSSSSPDLLQPTTSMLDFTNPSAVGFYYIPDQVIRVFKADQQSCYIIISKDTTAKEVVSHAVHEFNLIGAPETYSLCEVSVSPEGVIKQRRLPDQFSKLADRIQLNGRYYLKNNMETETLCSDEDAQELLRESQISLLQLSTIEVATQLSMRDFELFRNIEPTEYIDDLYKMDSKTGNAHLKQFEDVINQETFWVATEILTEPNQLKRMKIVKHFIKIALHCRECKNFNSMFAVISGLNLAPVARLRGTWEKLPSKYEKHLRDLQDLFDPSRNMAKYRNILSSQSMQPPIIPLFPVVKKDITFLHEGNDSKVDGLVNFEKLRMIAKEIRQVVRMTSANMDPAVMFRQRKKRWRSLGSLSQGSTNSNMLDVQGGAHKKRARRSSLLNAKKLYEDAQMARKVKQYLSNLNVETDEEKFQIMSLQCEPAYSTLTKNLNERKGAKSSEMSPIPMRSIGQTTKAHLHQQNRMSQVLQVPAVNLYPIRKKGPAKDHVTFSSPQKSLSLSEEVSSKKQAEDSISMASSLHSSPPASPQGSPHKVGNIQRSDNCGQMNLSGSSSSLASETSNKNSGQRSYGIGYALIPSTKSDNFSDSSHSEISSRSSIVSNCSADSMSAALQDERCLPQSLIVMDSAGGVERKDVPQPSAEYSQPTTGWSLTRPAIIRGLAVTSSMSNEEISHEHITVEAADSGRGSWTSCSSSSHDNFQNIQNQKSWDLLSAYRHTHLDGPIAEVEPTNCCSEETYLYSEAFSKNNRQSKASTEHDQSRQSWASSSSLSDTYETNYGTIKRRGLEHSTAEQTEVLDSKPATDANYKTVTSSTEKGLIEDEQVSAV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPF4</accession>
    <name>A0A452IPF4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">BTB domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IPF4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033814.1">
        <property type="protein sequence ID" value="ENSGAGP00000029777.1"/>
        <property type="gene ID" value="ENSGAGG00000021506.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031463">
        <property type="term" value="C:Cul3-RING ubiquitin ligase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097602">
        <property type="term" value="F:cullin family protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043161">
        <property type="term" value="P:proteasome-mediated ubiquitin-dependent protein catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051260">
        <property type="term" value="P:protein homooligomerization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd18390">
        <property type="entry name" value="BTB_POZ_KCTD5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.2000:FF:000001">
        <property type="entry name" value="Potassium channel tetramerization domain-containing 17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.710.10:FF:000005">
        <property type="entry name" value="Potassium channel tetramerization domain-containing 17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.2000">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.750">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.710.10">
        <property type="entry name" value="Potassium Channel Kv1.1, Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000210">
        <property type="entry name" value="BTB/POZ_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011333">
        <property type="entry name" value="SKP1/BTB/POZ_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003131">
        <property type="entry name" value="T1-type_BTB"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14958:SF12">
        <property type="entry name" value="BTB_POZ DOMAIN-CONTAINING PROTEIN KCTD5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14958">
        <property type="entry name" value="POTASSIUM CHANNEL TETRAMERISATION DOMAIN CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02214">
        <property type="entry name" value="BTB_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00225">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54695">
        <property type="entry name" value="POZ domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="BTB" evidence="2">
        <location>
            <begin position="41"/>
            <end position="143"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="210"/>
            <end position="231"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="217"/>
            <end position="231"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="SMART" id="SM00225"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029777.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="231" mass="25820" checksum="7DE3B7A7BACD09E1" modified="2019-05-08" version="1">MAAENHCEYPVPSMGVLGLPGTGGPCRRCSSGPSPSGVPGKWVRLNVGGTCFLTTRQTLCRDPKSFLFRLCQADPDLDSDKDETGAYLIDRDPTYFGPVLNYLRHGKLVINKDLAEEGVLEEAEFYNITSLIKLVKDKIRERDSKISQVPVKHVYRVLQCQEEELTQMVSTMSDGWKFEQLVSIGSSYNYGNEDQAEFLCVVSKELHNTPYGTTSEPSEKAKILQERGSRM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPF9</accession>
    <name>A0A452IPF9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="5">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the copine family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPF9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033819.1">
        <property type="protein sequence ID" value="ENSGAGP00000029782.1"/>
        <property type="gene ID" value="ENSGAGG00000021499.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005544">
        <property type="term" value="F:calcium-dependent phospholipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071277">
        <property type="term" value="P:cellular response to calcium ion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903265">
        <property type="term" value="P:positive regulation of tumor necrosis factor-mediated signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043122">
        <property type="term" value="P:regulation of canonical NF-kappaB signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd04048">
        <property type="entry name" value="C2A_Copine"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04047">
        <property type="entry name" value="C2B_Copine"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd01459">
        <property type="entry name" value="vWA_copine_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000099">
        <property type="entry name" value="Copine 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000008">
        <property type="entry name" value="C2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037768">
        <property type="entry name" value="C2B_Copine"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045052">
        <property type="entry name" value="Copine"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010734">
        <property type="entry name" value="Copine_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002035">
        <property type="entry name" value="VWF_A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036465">
        <property type="entry name" value="vWFA_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10857">
        <property type="entry name" value="COPINE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10857:SF2">
        <property type="entry name" value="COPINE-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00168">
        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07002">
        <property type="entry name" value="Copine"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00239">
        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00327">
        <property type="entry name" value="VWA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49562">
        <property type="entry name" value="C2 domain (Calcium/lipid-binding domain, CaLB)"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53300">
        <property type="entry name" value="vWA-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50004">
        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50234">
        <property type="entry name" value="VWFA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="C2" evidence="3">
        <location>
            <begin position="1"/>
            <end position="114"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="3">
        <location>
            <begin position="123"/>
            <end position="251"/>
        </location>
    </feature>
    <feature type="domain" description="VWFA" evidence="4">
        <location>
            <begin position="257"/>
            <end position="477"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00009048"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS50004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50234"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029782.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="501" mass="55992" checksum="353259BBE8F0F3D6" modified="2019-05-08" version="1">MAQCVTKVELSISCENLIDKDVGSKSDPLCVLLQNVGGDQWAELDRTEKIKNCQKPEFSKKLLIDYYFEKVQKLKFGVYDIDNKSYDLNDDDYLGGVECTLGQIVSSKTFTRPLELKKGKPAGKGTITISAEEIKDTRIVSLEIEAQNLDKKDFLGKSDPFLEFYKPSDGGKWQLIYRSEVIKNNLNPCWRKFSVPLQTFCGGDFNKPIKVEFECIHPGKKQKKKSYKNSGIVRIKSCKIETEYSFLDYIMGGCQINFTVGIDFTGSNGDPKSPDSLHYISPNGINEYLTAIWSVGNVVQDYDTDKLFPAFGFGAQVPPNWQVSHEFALNFNPNNPYCQGIQGIVDAYRQALPQVRLYGPTNFSPIINHVARFAAHSAQQGTAAQYFVLLIITDGEITDLDQTRQAIVNASKLPMSVIIVGVGNADFKAMEFLDGDNGVLKSLTGEPAVRDIVQFVPFRQFLNAPREALSQTVLAEVPKQLVSYYKMQGWAPVKLPGAKQE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPG0</accession>
    <name>A0A452IPG0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Inter-alpha-trypsin inhibitor heavy chain H3</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">May act as a carrier of hyaluronan in serum or as a binding protein between hyaluronan and other matrix protein, including those on cell surfaces in tissues to regulate the localization, synthesis and degradation of hyaluronan which are essential to cells undergoing biological processes.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the ITIH family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPG0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033921.1">
        <property type="protein sequence ID" value="ENSGAGP00000029871.1"/>
        <property type="gene ID" value="ENSGAGG00000021562.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004867">
        <property type="term" value="F:serine-type endopeptidase inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030212">
        <property type="term" value="P:hyaluronan metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.410:FF:000013">
        <property type="entry name" value="inter-alpha-trypsin inhibitor heavy chain H2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.410">
        <property type="entry name" value="von Willebrand factor, type A domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010600">
        <property type="entry name" value="ITI_HC_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050934">
        <property type="entry name" value="ITIH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013694">
        <property type="entry name" value="VIT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002035">
        <property type="entry name" value="VWF_A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036465">
        <property type="entry name" value="vWFA_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10338">
        <property type="entry name" value="INTER-ALPHA-TRYPSIN INHIBITOR HEAVY CHAIN FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10338:SF115">
        <property type="entry name" value="INTER-ALPHA-TRYPSIN INHIBITOR HEAVY CHAIN H3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06668">
        <property type="entry name" value="ITI_HC_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08487">
        <property type="entry name" value="VIT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00092">
        <property type="entry name" value="VWA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00609">
        <property type="entry name" value="VIT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00327">
        <property type="entry name" value="VWA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53300">
        <property type="entry name" value="vWA-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51468">
        <property type="entry name" value="VIT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50234">
        <property type="entry name" value="VWFA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="4" id="KW-0646">Protease inhibitor</keyword>
    <keyword evidence="7" id="KW-0654">Proteoglycan</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0964">Secreted</keyword>
    <keyword evidence="6" id="KW-0722">Serine protease inhibitor</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <feature type="domain" description="VIT" evidence="12">
        <location>
            <begin position="1"/>
            <end position="111"/>
        </location>
    </feature>
    <feature type="domain" description="VWFA" evidence="11">
        <location>
            <begin position="236"/>
            <end position="419"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010158"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022690"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022974"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00037051"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00039924"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50234"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS51468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029871.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="852" mass="95629" checksum="669084EDDDDF6574" modified="2019-05-08" version="1">MKIDSKVTSRFAHNVITSRAVNRANVSKEVVFDVDLPKTAFITNFSMTIDGVTYPGTIKEKEVAKKQYEKAVSQGQTAGLVRASGRKTEKFTVSVNIAAASKVTFELTYEELLKRNFGKYEMLIKVKPKQLINHFEIQANIFEPQGISELEAQGSFITNDLSQVIKKSFSGKKGHVSFKPTIDQQRTCANCSTSLLDGDFTIKYDVKRDRPGDLQIVNGYFVHFFAPTNLPNFPKNVVFVIDISGSMSGIGIQQTREALLKILEDVREDDYINFILFGSDVHVWKDTLLKATAENLKEAKDFVRAINTQGMTNLHGGLMKGIEMLTTAFETNSVPKRSASIIIMLTDGQPNEGVSDRQEIQMNVKKAIQGRYPLYNLAFGYGVDYNFLEKMALENKGLARRIYPQSDSAIQMEGFYEEVANPMLTEVELNYPENAISDLTQNIFKHYYDGSEIVVAGRLTDNDLDSLTADVKAQGAENDVMFSEQADVEETAKALQEQEYIFGDYIERLWAYLTIQQLLDKRTVAQGEEKENLTTEALELSLKYKFVTPLTSMVVTKPEDNEDKAAIADKPVEGTARAVPGTQSYPTHSYLYSRPPIPSLYTSVDGDPHFIISVPQKEDAICFNINEETGAVLNLINDPVTGIRVNGQLIGDKRVSSDAKIQNTYFGKLGIENKKLELKLEVTPEKITLQKGRKKTVFTWLDTVTLRQEGLTVIINRKKNLVLSMGDGASFVIVLHQVWKKHPLHQDFLGLYTLDSHKLSERTHGLLGQFFHHIDFNILEIHPGSDPKKPDATIVIKNNKLTVTRGWQKDYRKDPKHGIDIPCWFIHNNGAGLIDGVHTDYIVSSIFGTTVN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPG1</accession>
    <name>A0A452IPG1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Protein Wnt</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="11">Ligand for members of the frizzled family of seven transmembrane receptors.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 11">Secreted</location>
            <location evidence="1 11">Extracellular space</location>
            <location evidence="1 11">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 11">Belongs to the Wnt family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPG1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033755.1">
        <property type="protein sequence ID" value="ENSGAGP00000029728.1"/>
        <property type="gene ID" value="ENSGAGG00000021458.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005125">
        <property type="term" value="F:cytokine activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005109">
        <property type="term" value="F:frizzled binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060070">
        <property type="term" value="P:canonical Wnt signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045165">
        <property type="term" value="P:cell fate commitment"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030182">
        <property type="term" value="P:neuron differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd19336">
        <property type="entry name" value="Wnt_Wnt4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.2460.20:FF:000001">
        <property type="entry name" value="Wnt homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.2460.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005817">
        <property type="entry name" value="Wnt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009142">
        <property type="entry name" value="Wnt4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043158">
        <property type="entry name" value="Wnt_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018161">
        <property type="entry name" value="Wnt_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12027:SF104">
        <property type="entry name" value="PROTEIN WNT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12027">
        <property type="entry name" value="WNT RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00110">
        <property type="entry name" value="wnt"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01844">
        <property type="entry name" value="WNT4PROTEIN"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01349">
        <property type="entry name" value="WNTPROTEIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00097">
        <property type="entry name" value="WNT1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00246">
        <property type="entry name" value="WNT1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 11" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="9" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
    <keyword evidence="6 11" id="KW-0879">Wnt signaling pathway</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005683"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022687"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="RuleBase" id="RU003500"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029728.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="332" mass="36858" checksum="DF173F7B058D316E" modified="2019-05-08" version="1">MHSLFIPTCSCASTVWYLAKQPSLRGTLQDSSACEGLKGLAEEQVRICQRQVEAMDSVKHGAELAIEECQHQFHNRRWNCSTLQGLQVFGTRESAFIHAISSAGVAFAVTQACSRGELEQCGCDRKIRGVSPEGFQWSGCSDNLSYGIAFSQAFVDNPERSRGVSSSRALMNLHNNEAGRKVKLVECKCHGVSGSCEVRTCWKAMPPFRKVGNVLKEKFEGATEVHPKRVGSRKLLVPKSSRFKPYTAHDLVYLVASPDFCDRDPHSGVFGTSGRQCNRTSHAMDGCELLCCGRGFRTAQAERVERCSCKFHWCCSVKCKQCHHLVEVHTCR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPG3</accession>
    <name>A0A452IPG3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Pre-mRNA-splicing factor SYF1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the crooked-neck family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPG3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033859.1">
        <property type="protein sequence ID" value="ENSGAGP00000029820.1"/>
        <property type="gene ID" value="ENSGAGG00000021531.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071014">
        <property type="term" value="C:post-mRNA release spliceosomal complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000974">
        <property type="term" value="C:Prp19 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071007">
        <property type="term" value="C:U2-type catalytic step 2 spliceosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000349">
        <property type="term" value="P:generation of catalytic spliceosome for first transesterification step"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000023">
        <property type="entry name" value="Pre-mRNA-splicing factor SYF1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000137">
        <property type="entry name" value="Pre-mRNA-splicing factor syf1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000411">
        <property type="entry name" value="pre-mRNA-splicing factor SYF1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:001071">
        <property type="entry name" value="pre-mRNA-splicing factor SYF1-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.10">
        <property type="entry name" value="Tetratricopeptide repeat domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003107">
        <property type="entry name" value="HAT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056350">
        <property type="entry name" value="HAT_Syf1_central"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055430">
        <property type="entry name" value="HAT_Syf1_CNRKL1_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045075">
        <property type="entry name" value="Syf1-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011990">
        <property type="entry name" value="TPR-like_helical_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019734">
        <property type="entry name" value="TPR_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11246">
        <property type="entry name" value="PRE-MRNA SPLICING FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11246:SF5">
        <property type="entry name" value="PRE-MRNA-SPLICING FACTOR SYF1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23231">
        <property type="entry name" value="HAT_Syf1_CNRKL1_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23220">
        <property type="entry name" value="HAT_Syf1_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00386">
        <property type="entry name" value="HAT"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="SMART" id="SM00028">
        <property type="entry name" value="TPR"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48452">
        <property type="entry name" value="TPR-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0507">mRNA processing</keyword>
    <keyword evidence="6" id="KW-0508">mRNA splicing</keyword>
    <keyword evidence="7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0747">Spliceosome</keyword>
    <feature type="domain" description="Pre-mRNA-splicing factor SYF1 central HAT repeats" evidence="11">
        <location>
            <begin position="118"/>
            <end position="322"/>
        </location>
    </feature>
    <feature type="domain" description="Pre-mRNA-splicing factor Syf1/CRNKL1-like C-terminal HAT-repeats" evidence="12">
        <location>
            <begin position="324"/>
            <end position="711"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="742"/>
            <end position="792"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="703"/>
            <end position="730"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="10">
        <location>
            <begin position="753"/>
            <end position="775"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008644"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022664"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022728"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023187"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00039472"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF23220"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF23231"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029820.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="792" mass="92410" checksum="B3FDE9D45F60550F" modified="2019-05-08" version="1">MPALEETPVAAGEGKPDLFFEEEDLQYEEEILRNPFSVKCWIRYIEFKQNAPRHVLNLIYERALKELPGSYKLWYNYLKQRRKQVKNRCVTDPSYEEVNNCHERALVFMHKLSPENAEEYIEYLRSIDRLDEAALRLATVVNDERFISKEGKSNYQLWHELCDLISQNPDKVKSLNVGAIIRGGLTRFTDQLGKLWCSLADYYIRSGHFEKARDVYEEAIQTVMTVRDFTQVFDSYAQFEESMIAAKMETTSELGQEEEDDVDLELRLARFEQLITRRPLLLNSVLLRQNPHNVHEWHKRVKLFEGKPREIINTYTEAVQTVDPFKATGKTHTLWVSFAKFYEDNGQIEDARTIFEKATKVNFKQVDDLACVWCEYGEMELRHENYDQALRILRKATAIPAKKAEYFDASEPVQNRVYKSLKVWSMLADLEESLGTFKSTKAVYDRILDLRIATPQIIINYALFLEEHTYFEESFKAYERGISLFRWPNVYDIWNTYLTKFIDRYGGKKLERARDLFEQALDGCPQKYAKTIYLLYAKLEEEFGLARHAMAVYERATRAVQPSEQYEMYNIFIKRAAEIYGVTHTRSIYEKAIEVLSDEHAREMCQRFADMECKLGEIDRARAIYSYCSQICDPRTTGSFWQTWKDFEIRHGNEDTIREMLRIKRSVQATYNTQVNFMASQMLKVSSNATGTVSDLAPGQSGMDDMKLLEQRAEQLAAEAERDKPRAKEKILFVRSDASRSELAALSRQANPDEIDIEEEEEEGESEENEPDEVQLEQQSVPSAVFGGLKTD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPG5</accession>
    <name>A0A452IPG5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Peptidase S1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPG5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033929.1">
        <property type="protein sequence ID" value="ENSGAGP00000029876.1"/>
        <property type="gene ID" value="ENSGAGG00000021548.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00190">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.10:FF:000054">
        <property type="entry name" value="Complement C1r subcomponent"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.10.10">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009003">
        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043504">
        <property type="entry name" value="Peptidase_S1_PA_chymotrypsin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001314">
        <property type="entry name" value="Peptidase_S1A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001254">
        <property type="entry name" value="Trypsin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033116">
        <property type="entry name" value="TRYPSIN_SER"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24255">
        <property type="entry name" value="COMPLEMENT COMPONENT 1, S SUBCOMPONENT-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24255:SF13">
        <property type="entry name" value="MANNAN-BINDING LECTIN SERINE PROTEASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00089">
        <property type="entry name" value="Trypsin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00722">
        <property type="entry name" value="CHYMOTRYPSIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00020">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50494">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50240">
        <property type="entry name" value="TRYPSIN_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00135">
        <property type="entry name" value="TRYPSIN_SER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="3" id="KW-0768">Sushi</keyword>
    <feature type="domain" description="Peptidase S1" evidence="8">
        <location>
            <begin position="28"/>
            <end position="205"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="210"/>
            <end position="233"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022659"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029876.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="233" mass="25582" checksum="026EACA63F7EBDC7" modified="2019-05-08" version="1">PVRLTVLSFLSPVCGQPVRPLPGIIKRIIGGRNAELGFFPWQALIVVEDMSRVPNDKNKKDAVNRTVEEIILHVGFDIQNYNHDIALVKLKEKVTMDAYVMPVCLPQFEHELEGPHPNTLGLVAGWGISNPNITVDEIISNYSVTENMFCAGYYEGGKDTCLGDSGGAFVIQDPRTQRWVAQGLVSWGGPEECGMSCLPTVANARCLRGNEQNRESPSDPSPVTRSHLLANRG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPG8</accession>
    <name>A0A452IPG8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="22">Serine/threonine-protein kinase Nek4</fullName>
            <ecNumber evidence="5">2.7.11.1</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="24">Never in mitosis A-related kinase 4</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="23">Serine/threonine-protein kinase 2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="28 29">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="29" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="28" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="28" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="21">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="20">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29035"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell projection</location>
            <location evidence="2">Cilium</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the protein kinase superfamily. NEK Ser/Thr protein kinase family. NIMA subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="5"/>
    <dbReference type="AlphaFoldDB" id="A0A452IPG8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033933.1">
        <property type="protein sequence ID" value="ENSGAGP00000029881.1"/>
        <property type="gene ID" value="ENSGAGG00000021569.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005929">
        <property type="term" value="C:cilium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd08223">
        <property type="entry name" value="STKc_Nek4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000219">
        <property type="entry name" value="Putative serine/threonine-protein kinase Nek4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000247">
        <property type="entry name" value="serine/threonine-protein kinase Nek4 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051131">
        <property type="entry name" value="NEK_Ser/Thr_kinase_NIMA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR44899">
        <property type="entry name" value="CAMK FAMILY PROTEIN KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR44899:SF7">
        <property type="entry name" value="NIMA-RELATED KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="16 25" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="19" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="10" id="KW-0132">Cell division</keyword>
    <keyword evidence="18" id="KW-0966">Cell projection</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="15" id="KW-0418">Kinase</keyword>
    <keyword evidence="17" id="KW-0460">Magnesium</keyword>
    <keyword evidence="12" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="6" id="KW-0488">Methylation</keyword>
    <keyword evidence="14" id="KW-0498">Mitosis</keyword>
    <keyword evidence="13 25" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="9" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="29" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="11" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Protein kinase" evidence="27">
        <location>
            <begin position="6"/>
            <end position="261"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="26">
        <location>
            <begin position="336"/>
            <end position="355"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="26">
        <location>
            <begin position="400"/>
            <end position="426"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="26">
        <location>
            <begin position="576"/>
            <end position="623"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="26">
        <location>
            <begin position="341"/>
            <end position="354"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="26">
        <location>
            <begin position="576"/>
            <end position="585"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="26">
        <location>
            <begin position="607"/>
            <end position="617"/>
        </location>
    </feature>
    <feature type="binding site" evidence="25">
        <location>
            <position position="36"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001936"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004138"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00010886"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012513"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022776"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00047899"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00048679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00067731"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00080102"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00082679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="27">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="28">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029881.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="29">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="759" mass="87107" checksum="9722184306E2311F" modified="2019-05-08" version="1">MPLAAYFFLRAVGKGSYGEVSLVRHRQDNRQYVIKKLNLKHASSRERKAAEQEAQLLSQLKHPNIVTYRESWEGEDGLLYIVMGFCEGGDLYHKLKEQKGKPLPENQVVEWFVQIAMALQYLHEKHILHRDLKTQNVFLTRTNIIKVGDLGIARVLENQYDMASTLIGTPYYMSPELFSNKPYNYKSDVWALGCCVYEMATLKHAFNAKDMNSLVYRIIEGKLPPMPKDYSPQLAELVRTMLSKKPEERPNVKSILRQPYIKRQISLFLEATKAKTAKNHKKISDSKPRSAASVISVKTGYNNGTITHKKHHSEQARRYRANEEKHLIKDKTFEIYPSEKPAAEPEKNANKNEKNSTGVSIATVSKVDIDILLSERRNYSKSDHLLQDNKARCLSIPSEVESKITSNGPQTKEDRLQQNSEPGSRTEILDSKHFAVDVANEGDDTFKLLQPVSKDQKENDDQSLDSTEKFLASFVPVIIQDEASHGNPEDVQEKITFHLQPHSSFCEPSLSRQRWQKKRELDEVCSEKDRPLSARERRRLKQSQEKMFPFVPAARRASHSAVVEAKSLVENLIKVTQSSSDPSISQKKKLTHCLSEDELSSSTSSTDKSDGDSKEGKSNTNEMNDLVQLMTQTLKMDSKENCEHSSILIPASEFKLHRKYRDTLILHGKVSNEPEKFKFEEFPSDVLSGPEKIRRMVEILRSDVVQGLGVKLLEKVYDVMEEDDELKRELHLREHMREKYTDYSVKARHLKFLEENVKF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPJ0</accession>
    <name>A0A452IPJ0_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="21">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="15">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
        <text evidence="15">Binds 1 zinc ion per subunit.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="17">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPJ0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033858.1">
        <property type="protein sequence ID" value="ENSGAGP00000029819.1"/>
        <property type="gene ID" value="ENSGAGG00000021496.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031012">
        <property type="term" value="C:extracellular matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004222">
        <property type="term" value="F:metalloendopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030198">
        <property type="term" value="P:extracellular matrix organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd04273">
        <property type="entry name" value="ZnMc_ADAMTS_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000006">
        <property type="entry name" value="A disintegrin and metalloproteinase with thrombospondin motifs 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.830:FF:000001">
        <property type="entry name" value="A disintegrin and metalloproteinase with thrombospondin motifs 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000011">
        <property type="entry name" value="A disintegrin and metalloproteinase with thrombospondin motifs 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000022">
        <property type="entry name" value="A disintegrin and metalloproteinase with thrombospondin motifs 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.1620.60:FF:000001">
        <property type="entry name" value="A disintegrin and metalloproteinase with thrombospondin motifs 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.390.10:FF:000008">
        <property type="entry name" value="A disintegrin and metalloproteinase with thrombospondin motifs 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.830">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.1620.60">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.390.10">
        <property type="entry name" value="Collagenase (Catalytic Domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.20.100.10">
        <property type="entry name" value="Thrombospondin type-1 (TSP1) repeat"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013273">
        <property type="entry name" value="ADAMTS/ADAMTS-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050439">
        <property type="entry name" value="ADAMTS_ADAMTS-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041645">
        <property type="entry name" value="ADAMTS_CR_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045371">
        <property type="entry name" value="ADAMTS_CR_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010294">
        <property type="entry name" value="ADAMTS_spacer1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024079">
        <property type="entry name" value="MetalloPept_cat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001590">
        <property type="entry name" value="Peptidase_M12B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002870">
        <property type="entry name" value="Peptidase_M12B_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010909">
        <property type="entry name" value="PLAC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000884">
        <property type="entry name" value="TSP1_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036383">
        <property type="entry name" value="TSP1_rpt_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13723:SF158">
        <property type="entry name" value="A DISINTEGRIN AND METALLOPROTEINASE WITH THROMBOSPONDIN MOTIFS 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13723">
        <property type="entry name" value="ADAMTS A DISINTEGRIN AND METALLOPROTEASE WITH THROMBOSPONDIN MOTIFS PROTEASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17771">
        <property type="entry name" value="ADAMTS_CR_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19236">
        <property type="entry name" value="ADAMTS_CR_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05986">
        <property type="entry name" value="ADAMTS_spacer1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01562">
        <property type="entry name" value="Pep_M12B_propep"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01421">
        <property type="entry name" value="Reprolysin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19030">
        <property type="entry name" value="TSP1_ADAMTS"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00090">
        <property type="entry name" value="TSP_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01857">
        <property type="entry name" value="ADAMTSFAMILY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00209">
        <property type="entry name" value="TSP1"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55486">
        <property type="entry name" value="Metalloproteases ('zincins'), catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF82895">
        <property type="entry name" value="TSP-1 type 1 repeat"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50215">
        <property type="entry name" value="ADAM_MEPRO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50900">
        <property type="entry name" value="PLAC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50092">
        <property type="entry name" value="TSP1"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="15" id="KW-0106">Calcium</keyword>
    <keyword evidence="12 16" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="5 15 17" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0482">Metalloprotease</keyword>
    <keyword evidence="4" id="KW-0645">Protease</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="9 15 17" id="KW-0862">Zinc</keyword>
    <keyword evidence="11" id="KW-0865">Zymogen</keyword>
    <feature type="domain" description="Peptidase M12B" evidence="19">
        <location>
            <begin position="238"/>
            <end position="442"/>
        </location>
    </feature>
    <feature type="domain" description="PLAC" evidence="20">
        <location>
            <begin position="995"/>
            <end position="1038"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1104"/>
            <end position="1124"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1139"/>
            <end position="1174"/>
        </location>
    </feature>
    <feature type="active site" evidence="14 17">
        <location>
            <position position="381"/>
        </location>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="241"/>
        </location>
        <ligand>
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="241"/>
        </location>
        <ligand>
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
            <label>2</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="333"/>
        </location>
        <ligand>
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15 17">
        <location>
            <position position="380"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <note>catalytic</note>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15 17">
        <location>
            <position position="384"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <note>catalytic</note>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15 17">
        <location>
            <position position="390"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <note>catalytic</note>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="437"/>
        </location>
        <ligand>
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="440"/>
        </location>
        <ligand>
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
            <label>2</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="440"/>
        </location>
        <ligand>
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="315"/>
            <end position="364"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="358"/>
            <end position="437"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="397"/>
            <end position="423"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="464"/>
            <end position="489"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="475"/>
            <end position="498"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="484"/>
            <end position="517"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="511"/>
            <end position="522"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="545"/>
            <end position="582"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="549"/>
            <end position="587"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="560"/>
            <end position="572"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023049"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023145"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PIRSR" id="PIRSR613273-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PIRSR" id="PIRSR613273-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PIRSR" id="PIRSR613273-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00276"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50215"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029819.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1174" mass="132839" checksum="30BC1B3A342CE739" modified="2019-05-08" version="1">LYRNRDWLILFALADLPKIRPRLYGLVTPISTSADGSFISNILSANHQRRLTRDVSQSPKQLYFNITAFGREFHLRLRPNTHLVAPEAVVEWYEDSVESGNGTEAGNKGQTRSTKEHIWKKEPLWTNCAYVGDITDIPGASVAISNCDGLAGMIRTDKDEYFIEPLERGKQMEEERGRIHMVYRRSAVTQDPGDMLLDFHTEASLLSGLDGLDSMYSNIEQQLNETLRRRRHAGDNDYNIEVLLGVDDSVVRFHGKEHVQNYLLTLMNIVNEIYHDESLGVHINVVLVRMIMLGYAKSVSLIERGNPSRSLENVCRWAYQQQKSDPNHSEHHDHAIFLTRQDFGPAGMQGYAPVTGMCHPVRSCTLNHEDGFSSAFVVAHETGHVLGMEHDGQGNRCGDETAMGSVMAPLVQAAFHRYHWSRCSGQELKRYIHSYDCLLDDPFEHDWPKLPELPGINYSMDEQCRFDFGVGYKMCTAFRTFDPCKQLWCSHPDNPYFCKTKKGPPLDGTECAPGKWCYKGHCMWKNANQLKQDGNWGSWTKFGSCSRTCGTGVRFRTRQCNNPMPVNGGQDCASVNFEYQLCSTEECPKHFEDFRAQQCQQRNSHFEYQKSKHHWLPYEHPDGKSNKRCHLYCQSKEIGDIAYMKQLAHDGTRCSYKDPYSICVRGECVKVGCDKEIGSNKVEDKCGVCGGDNSHCRTVKGTFTRTPKKPGYLKMFDIPPGARHVFIQEDEASPHILAIKNQATGHYILNGKGEEAKSRSFIDLGVEWEYNTEDDIETLHTDGPLHDAVVVLVIPQENDTRSSLTYKYIIHEDSVPTINSNNVLQDEIDTFEWALKSWSQCSKPCAGGFQYTKYGCRRKSDNKMVHRSFCEASKKPKPIRRMCNLQECTQPLWIADEWEYCTKTCGHSGYQIRTVRCIQPLHDGANRSVNIKYCSGDRPESRRPCNRTPCPAQWKAGPWAECSVTCGEGTESRQVTCRAGDQCDGEKPESVRVCKLAPCNDEPCLGDKSIFCQMEVLARYCSIPGYNKLCCESCSKRSSTLPPPYLTEAAETKEEVMFHPRDLPRALAMPTSLVPQHAEVADENRSSVGRMPMEDDIDVLVSTSNSKSKGAELPQRRAYQAGSQTSRLIAMPYQSSTKIVNNTSAGALSPARTSRKNEKHVEKRHALRSSTVER</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPJ2</accession>
    <name>A0A452IPJ2_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="23">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 16">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000033967.1">
        <property type="protein sequence ID" value="ENSGAGP00000029909.1"/>
        <property type="gene ID" value="ENSGAGG00000021570.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005884">
        <property type="term" value="C:actin filament"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030027">
        <property type="term" value="C:lamellipodium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016459">
        <property type="term" value="C:myosin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001726">
        <property type="term" value="C:ruffle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000146">
        <property type="term" value="F:microfilament motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030048">
        <property type="term" value="P:actin filament-based movement"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072673">
        <property type="term" value="P:lamellipodium morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007266">
        <property type="term" value="P:Rho protein signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd01385">
        <property type="entry name" value="MYSc_Myo9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd17217">
        <property type="entry name" value="RA_Myosin-IXb"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04407">
        <property type="entry name" value="RhoGAP_myosin_IXB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.720:FF:000003">
        <property type="entry name" value="Putative unconventional myosin-IXa"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000008">
        <property type="entry name" value="Putative unconventional myosin-IXa"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.820:FF:000003">
        <property type="entry name" value="unconventional myosin-IXa isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.555.10:FF:000009">
        <property type="entry name" value="unconventional myosin-IXa isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.530:FF:000005">
        <property type="entry name" value="unconventional myosin-IXa isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000013">
        <property type="entry name" value="unconventional myosin-IXa isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.190:FF:000013">
        <property type="entry name" value="unconventional myosin-IXa isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.530:FF:000009">
        <property type="entry name" value="unconventional myosin-IXb isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.190">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.60.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.20.240.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.555.10">
        <property type="entry name" value="Rho GTPase activation protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046349">
        <property type="entry name" value="C1-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000048">
        <property type="entry name" value="IQ_motif_EF-hand-BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046987">
        <property type="entry name" value="Myo9"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036023">
        <property type="entry name" value="MYSc_Myo9"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002219">
        <property type="entry name" value="PKC_DAG/PE"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000159">
        <property type="entry name" value="RA_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046989">
        <property type="entry name" value="RA_Myosin-IXb"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000198">
        <property type="entry name" value="RhoGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028557">
        <property type="entry name" value="RhoGAP_myosin_IXB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029071">
        <property type="entry name" value="Ubiquitin-like_domsf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46184:SF2">
        <property type="entry name" value="UNCONVENTIONAL MYOSIN-IXB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46184">
        <property type="entry name" value="UNCONVENTIONAL MYOSIN-IXB-LIKE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00612">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00788">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00620">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00109">
        <property type="entry name" value="C1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00015">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00314">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00324">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57889">
        <property type="entry name" value="Cysteine-rich domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54236">
        <property type="entry name" value="Ubiquitin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50200">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50238">
        <property type="entry name" value="RHOGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00479">
        <property type="entry name" value="ZF_DAG_PE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50081">
        <property type="entry name" value="ZF_DAG_PE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="15 16" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="11 16" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="12 17" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="3" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="6" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="14 16" id="KW-0505">Motor protein</keyword>
    <keyword evidence="13 16" id="KW-0518">Myosin</keyword>
    <keyword evidence="8 16" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="10" id="KW-0862">Zinc</keyword>
    <keyword evidence="9" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="Ras-associating" evidence="20">
        <location>
            <begin position="15"/>
            <end position="113"/>
        </location>
    </feature>
    <feature type="domain" description="Myosin motor" evidence="22">
        <location>
            <begin position="145"/>
            <end position="948"/>
        </location>
    </feature>
    <feature type="domain" description="Phorbol-ester/DAG-type" evidence="19">
        <location>
            <begin position="1653"/>
            <end position="1702"/>
        </location>
    </feature>
    <feature type="domain" description="Rho-GAP" evidence="21">
        <location>
            <begin position="1721"/>
            <end position="1909"/>
        </location>
    </feature>
    <feature type="region of interest" description="Actin-binding" evidence="16">
        <location>
            <begin position="830"/>
            <end position="852"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1095"/>
            <end position="1148"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1190"/>
            <end position="1229"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1246"/>
            <end position="1364"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1474"/>
            <end position="1502"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1626"/>
            <end position="1647"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1953"/>
            <end position="1983"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="2004"/>
            <end position="2038"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="17">
        <location>
            <begin position="1036"/>
            <end position="1063"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="1118"/>
            <end position="1143"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="1190"/>
            <end position="1203"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="1246"/>
            <end position="1257"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="1273"/>
            <end position="1283"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="1312"/>
            <end position="1324"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="1350"/>
            <end position="1364"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="1487"/>
            <end position="1498"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="18">
        <location>
            <begin position="1634"/>
            <end position="1644"/>
        </location>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <begin position="238"/>
            <end position="245"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50081"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50200"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50238"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029909.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2187" mass="251315" checksum="39D986334776AA8A" modified="2019-05-08" version="1">MSLKDADSAVCQEKATYNLHIYPQLSTESTTCCRVTATKDSTTSDVIKDVIQILDLDVSKHYVLVEVKESGGEEWVLDLNDSPVHRVLLWPRRAQDEHPQNDGYYFLLQERDTDGTIKYVHMQLVSQEKDARRLVERGFLPWHQENFDDLCNLSNLTETTLLENLKRRFLKHKIYTYAGSILIAINPFKFLPIYNPKYVKMYENHQLGKLEPHIFAIADVAYHTMLKKHVNQCIVISGESGSGKTQSTNFLIHCFTALSQKGYASGVERTILGAGPVLEAFGNAKTAHNNNSSRFGKFIQVNYLENGIVRGAVVEKYLLEKSRLVSQEKDERNYHVFYYLLLGVSEEERKEFHLKQPEDYFYLNQHNLKIEDGEDLRHDFERLKQAMEMVGFLPATKKQIFSVLSAILYLGNVTYKKRATGRDEGLEVGPTEVLDTLSQLLKVKREVLVEVLTKRKTVTANDKLILPYSLSEAITARDSMAKSLYSALFDWIVLRINHALLNKKDMEESVSCLSIGVLDIFGFEDFETNSFEQFCINYANEQLQYYFNQHIFKLEQEEYQSEGIAWHNIDYTDNVTCIHLISKKPTGLFYLLDEESNFPHATNQTLLAKFKQQHEENKFFVGTPVKEPAFIIRHFAGKVKYQIKDFREKNMDYMRPDIVALLRSSDSAYVRELIGMDPVAVFRWAILRAAIRAMAVFAEAGRQRAQKTAGVVRHGPRVPLGELQRANTPIEKVYRDLHDQIIKSIKGLPWQGEDPCRLLRSLSRLQQPHVHFLKSRGTKQKQIIPKNLLDSKSLKLIVSMTLHDRTTKSLLHLHKKKKPPSISAQFQNSLNKLLETLGKAEPFFIRCIRSNAEKRELFFDEGLVLQQLRYTGMLETVRIRRSGYSAKYTFQDFIDQFQVLLPQNAKASKEEISAHLNKLKLDKNNYQIGKTKVFMKEAERQMLQDTLHKEVIRKIVLLQSWLRMVLERRHFLRMRQATITLQTCWRSYRIRRALERTNAAIYIQSVWRRYRERESYHQQRRRISLVQALVRGHLQRKRFQQMIVEKQQEMQQAQEGEDSVSQEDGNELVLEQLAVKPEMQLDQGIEQMAAGVDEAQNDGDEKLSQATEKAVPSQKSITDGHEKLTSSREKRESRRQRGLEHNKLQNKHVLFPLKEQSLVCQEEETSAKETPETVQEPNKFTAYKTVLPKGTEEERIPVEENKTLSDTQVPGEILDSDVSFPKQSNESKQEILDDKIIDKLKIQDNQKNQVKGSQSFTCPERPSDFSLHIKKNLTPTRSFQTGDENYGNKNKHRISKVDKDLDSPPSFQIQRYLHDPGELRDKREKWKGKRQSEAGQNDVLSESLDERNRTGQSPQTLLSKKGTTSSLDDIANLAQSVTMIQPSPETMAEEKSNKKQTLQRKSNEQLSLLDLAVSVQLPSQPMDAKSTFKSPLRKLLGKKPDKKYVRESPDVMEEGENLPLLSCILISESVESQKLSEAPSGHPRRPQAGERPVKETSKGKRNRTIKIKKISAVSEKWPASVIREITNANELKYLDEFLLNKINDLNSQKSATECLFFEATEKFRGNIKTMYSVPNGQIHVGYKDLIENYQLLVINLAKEREEKDVKLVLNLFQSLLDEFSRGYAKREESEQHKQSKTQKKKRKQERAIEEHNGHVFTSYQVSIRQSCEQCSSYIWPMEKACLCSTCKLICHKKCMSKTQSCCTSPCGKKNEQGVDSRHFGVCVSSLISDKNSVPVVMEKLLEYVEMHGLYTEGIYRKSGSANRMRELKQLLQTDPNSVKLENYPIHTITGILKQWLRELPDPLMTFAQYSDFLRAVELPEKQEQLIAIYSVLEQLPQANHNTLERLIFHLVKVALIEDVNRMSPNALAIVFAPCLLRCPDNSDPLTSMKDVSKTTMCVEMLIKEQMRKYKIKMDEINQLEAAESIAFRRLSLLRQNTLWPVKLGFSSPYEGVLSKSPQDKGNDDSFPELGSLQEEDEVSDVDNREKEILIERIQSIKEEKEDITYRLPELDQQGSDEENLDSETSVSTESLPEDRTGRLDTEGTMLSEFQCHGQCPSVPAKDTCGGPSISLSLASTFSSVTRRRPSSVLVKMKVPCRNPVIPTANIKLPYGIPKLTESQRRTSAEESQPVVRRRDRLARRTDKIHSVYIAQGSVVAQTQDPLDEYEPTAKVKRRFSDPYSHISCIDE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPJ3</accession>
    <name>A0A452IPJ3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Nck-associated protein 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="6">Cell projection</location>
            <location evidence="6">Lamellipodium membrane</location>
            <topology evidence="6">Single-pass membrane protein</topology>
            <orientation evidence="6">Cytoplasmic side</orientation>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="7">Belongs to the HEM-1/HEM-2 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPJ3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034087.1">
        <property type="protein sequence ID" value="ENSGAGP00000030017.1"/>
        <property type="gene ID" value="ENSGAGG00000019065.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031258">
        <property type="term" value="C:lamellipodium membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031209">
        <property type="term" value="C:SCAR complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030031">
        <property type="term" value="P:cell projection assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030866">
        <property type="term" value="P:cortical actin cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048812">
        <property type="term" value="P:neuron projection morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019137">
        <property type="entry name" value="Nck-associated_protein-1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12093">
        <property type="entry name" value="NCK-ASSOCIATED PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12093:SF11">
        <property type="entry name" value="NCK-ASSOCIATED PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09735">
        <property type="entry name" value="Nckap1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="1" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="5" id="KW-0966">Cell projection</keyword>
    <keyword evidence="4 10" id="KW-0472">Membrane</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 10" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="3 10" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="6"/>
            <end position="28"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="510"/>
            <end position="535"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="521"/>
            <end position="535"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00037839"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00037947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00039689"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030017.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="977" mass="111367" checksum="4DE5DF5C6F9B362E" modified="2019-05-08" version="1">MCVYILIFLLLKNYLDLIITYTTLMILLSRIEERKAIIGLYNYAHEMTHGASDREYPRLGQMIVDYENPLKKMMEEFVPHSKSLSDALISLQMVYPRRNLSADQWRNAQLLSLISAPSTMLNPAQSDTMPCEYLSLDAMEKWIIFGFILCHGILNSDATALNLWKLALQSSSCLSLFRDEVFHIHKAAEDLFVNIRGYGIFMNVFGSMHRERRKFLRSALKELATVLSDQPGLLGPKALFVFMALSFARDEIIWLLRHADNMPKKSADDFIDKHIAELIFYMEELRAHVRKYGPVMQRYYVQYLSGFDAVVLNELVQNLSVCPEDESIIMSSFVNTMTSLSVKQVEDGEVFDFRGMRLDWFRLQAYTSVSKASLSLADHRELGKMMNTIIFHTKMVDSLVEMLVETSDLSIFCFYSRAFEKMFQQCLELPSQSRYSIAFPLLCTHFMSCTHELCPEERHHIGDRSLSLCNMFLDEMAKQARNLITDICTEQCTLSDQLLPKHCAKTISQAVNKKSKKQTGKKGEPEREKPGVESMRKNRLVVTNLDKLHTALSELCFSINYVPNMVVWEHTFTPREYLTSHLEIRFTKSIVGMTMYNQATQEIAKPSELLTSVRAYMTVLQSIENYVQIDITRVFNNVLLQQTQHLDSHGEPTITSLYTNWYLETLLRQVSNGHIAYFPAMKAFVNLPTENELTFNAEEYSDISEMRALSELLGPYGMKFLSESLMWHISSQVAELKKLVVENVEVLTQMRTSFDKPDQMAALFKRLSSVDSVLKRMTIIGVILSFRSLAQEALRDCFSPCNNCYIVSQVAMNVYELSSAAGLPCEIDPALVVALSSQKSENISPEEEYKIACLLMVFVAVSLPTLASNVMSQYSPAIEGHCNNIHCLAKAINQIAAALFTIHKGSIEDQLTATKIRLKNQTKTLNICCNSDLPFRCTVIVQESPFLTMDLLESCFPYVLLRNAYHAVYKQSVTSSA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPJ4</accession>
    <name>A0A452IPJ4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">RAB GTPase activating protein 1 like</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IPJ4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033789.1">
        <property type="protein sequence ID" value="ENSGAGP00000029759.1"/>
        <property type="gene ID" value="ENSGAGG00000021483.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47728:SF1">
        <property type="entry name" value="RAB GTPASE ACTIVATING PROTEIN 1 LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47728">
        <property type="entry name" value="RAB GTPASE-ACTIVATING PROTEIN 1-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="229"/>
            <end position="254"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="9"/>
            <end position="114"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="174"/>
            <end position="222"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029759.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="5">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="254" mass="29079" checksum="A44249C1C3241208" modified="2019-05-08" version="1">MIESSSWSVTLQERENRRLQEASMRLEQENDDLAHELVTSKIALRNDLDQAEDKADVLNKELLLTKQKLVETEEEKRKQEEETAQLKEIFRKQLEKAESEIKKTTAIIAEYKQICSQLSTRLEKQQTASKEELEVVKGKVMACKHCSEIFSKEGALKLPAISRETQGTERDDEKDALKKQLREMELELAQTKLQLVEAKCKIQELEHQRGTLMNEIQAAKNSWFSKTLNSIKTATSTQPAQQPPPSQPPKDSST</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPJ6</accession>
    <name>A0A452IPJ6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Uronyl 2-sulfotransferase</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Golgi apparatus membrane</location>
            <topology evidence="1">Single-pass type II membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the sulfotransferase 3 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPJ6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033865.1">
        <property type="protein sequence ID" value="ENSGAGP00000029826.1"/>
        <property type="gene ID" value="ENSGAGG00000021514.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000139">
        <property type="term" value="C:Golgi membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008146">
        <property type="term" value="F:sulfotransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000580">
        <property type="entry name" value="uronyl 2-sulfotransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007734">
        <property type="entry name" value="Heparan_SO4_2-O-STrfase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005331">
        <property type="entry name" value="Sulfotransferase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12129">
        <property type="entry name" value="HEPARAN SULFATE 2-O-SULFOTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12129:SF15">
        <property type="entry name" value="URONYL 2-SULFOTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03567">
        <property type="entry name" value="Sulfotransfer_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="8" id="KW-0472">Membrane</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0735">Signal-anchor</keyword>
    <keyword evidence="3" id="KW-0808">Transferase</keyword>
    <keyword evidence="4" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6" id="KW-1133">Transmembrane helix</keyword>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="290"/>
            <end position="331"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="11">
        <location>
            <begin position="309"/>
            <end position="324"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004323"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010569"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00071728"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029826.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="331" mass="39226" checksum="948AE483C98BA918" modified="2019-05-08" version="1">SAGLRCLGGGGGARTRHRLPLPVLPFPSQVVYNRVGKCGSRTVVLLLRILSERHGFNLVTSDIHNKTRLTKNEQMELIKNISTAEQPYLFTRHVHFLNFSRFGGEQPVYINIIRDPANRFLSNYFFRRFGDWRGEQNHMIRTPSMRQEERYLDINICILENYPECSNPRLFYIIPYFCGQHPRCREPGEWALERAKLNVNENFLLVGILEELEDVLLLLERFLPHYFKDVLSIYKNPEHRKLGNLTVTVKKTVPSPEAIQILYQRMRYEYEFYYYVKEQFHLLKRKFGLKSHSSEPPPKPEFFIPSPLETEEPIDDEEQDDEKWLEDIYKR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPK0</accession>
    <name>A0A452IPK0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Peptidase S1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000033914.1">
        <property type="protein sequence ID" value="ENSGAGP00000029865.1"/>
        <property type="gene ID" value="ENSGAGG00000021548.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001867">
        <property type="term" value="P:complement activation, lectin pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00190">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.10:FF:000112">
        <property type="entry name" value="Mannan-binding lectin serine protease 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.10:FF:000068">
        <property type="entry name" value="transmembrane protease serine 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.10.10">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009003">
        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043504">
        <property type="entry name" value="Peptidase_S1_PA_chymotrypsin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001314">
        <property type="entry name" value="Peptidase_S1A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001254">
        <property type="entry name" value="Trypsin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033116">
        <property type="entry name" value="TRYPSIN_SER"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24255">
        <property type="entry name" value="COMPLEMENT COMPONENT 1, S SUBCOMPONENT-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24255:SF13">
        <property type="entry name" value="MANNAN-BINDING LECTIN SERINE PROTEASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00089">
        <property type="entry name" value="Trypsin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00722">
        <property type="entry name" value="CHYMOTRYPSIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00020">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50494">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50240">
        <property type="entry name" value="TRYPSIN_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00135">
        <property type="entry name" value="TRYPSIN_SER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 4" id="KW-0732">Signal</keyword>
    <keyword evidence="1" id="KW-0768">Sushi</keyword>
    <feature type="signal peptide" evidence="4">
        <location>
            <begin position="1"/>
            <end position="32"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019207613" description="Peptidase S1 domain-containing protein" evidence="4">
        <location>
            <begin position="33"/>
            <end position="291"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase S1" evidence="5">
        <location>
            <begin position="45"/>
            <end position="277"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022659"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029865.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="291" mass="32320" checksum="946DBB06F60617F9" modified="2019-05-08" version="1">MAHCLLTACPSSSVMLAPVRLTVLSFLSPVCGQPVRPLPGIIKRIIGGRNAELGFFPWQALIVVEDMSRVPNDKWFGSGALLSDSWVLTAAHVLRSQRRXAPSAGTTLSSRNKKDAVNRTVEEIILHVGFDIQNYNHDIALVKLKEKVTMDAYVMPVCLPQFEHELEGSGMRTLSDILQYVKLPVVLHDECKTSYESRSGNYSVTENMFCAGYYEGGKDTCLGDSGGAFVIQDPRTQRWVAQGLVSWGGPEECGSKQVYGVYTKVSNYVDWVEEKMGSSHKWTFLDPELNR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPK5</accession>
    <name>A0A452IPK5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Peptidase S1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000033920.1">
        <property type="protein sequence ID" value="ENSGAGP00000029870.1"/>
        <property type="gene ID" value="ENSGAGG00000021548.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001867">
        <property type="term" value="P:complement activation, lectin pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00190">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.10:FF:000062">
        <property type="entry name" value="mannan-binding lectin serine protease 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.10.10">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009003">
        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043504">
        <property type="entry name" value="Peptidase_S1_PA_chymotrypsin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001314">
        <property type="entry name" value="Peptidase_S1A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001254">
        <property type="entry name" value="Trypsin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033116">
        <property type="entry name" value="TRYPSIN_SER"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24255">
        <property type="entry name" value="COMPLEMENT COMPONENT 1, S SUBCOMPONENT-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24255:SF13">
        <property type="entry name" value="MANNAN-BINDING LECTIN SERINE PROTEASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00089">
        <property type="entry name" value="Trypsin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00722">
        <property type="entry name" value="CHYMOTRYPSIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00020">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50494">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50240">
        <property type="entry name" value="TRYPSIN_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00135">
        <property type="entry name" value="TRYPSIN_SER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 4" id="KW-0732">Signal</keyword>
    <keyword evidence="1" id="KW-0768">Sushi</keyword>
    <feature type="signal peptide" evidence="4">
        <location>
            <begin position="1"/>
            <end position="29"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019358788" description="Peptidase S1 domain-containing protein" evidence="4">
        <location>
            <begin position="30"/>
            <end position="255"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase S1" evidence="5">
        <location>
            <begin position="42"/>
            <end position="255"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022659"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029870.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="255" mass="28078" checksum="679FAB7BE16580B1" modified="2019-05-08" version="1">MLNRLSAVSCEVLAPVRLTVLSFLSPVCGQPVRPLPGIIKRIIGGRNAELGFFPWQALIVVEDMSRVPNDKWFGSGALLSDSWVLTAAHVLRSQRRXAPSEIILHVGFDIQNYNHDIALVKLKEKVTMDAYVMPVCLPQFEHELEGPHPNTLGLVAGWGISNPNITVDEIISSGMRTLSDILQYVKLPVVLHDECKTSYESRSGNYSVTENMFCAGYYEGGKDTCLGDSGGAFVIQDPRTQSYWANGTADPSWSL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPL0</accession>
    <name>A0A452IPL0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Inter-alpha-trypsin inhibitor heavy chain H3</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">May act as a carrier of hyaluronan in serum or as a binding protein between hyaluronan and other matrix protein, including those on cell surfaces in tissues to regulate the localization, synthesis and degradation of hyaluronan which are essential to cells undergoing biological processes.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the ITIH family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPL0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033925.1">
        <property type="protein sequence ID" value="ENSGAGP00000029875.1"/>
        <property type="gene ID" value="ENSGAGG00000021562.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004867">
        <property type="term" value="F:serine-type endopeptidase inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030212">
        <property type="term" value="P:hyaluronan metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.410:FF:000013">
        <property type="entry name" value="inter-alpha-trypsin inhibitor heavy chain H2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.410">
        <property type="entry name" value="von Willebrand factor, type A domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010600">
        <property type="entry name" value="ITI_HC_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050934">
        <property type="entry name" value="ITIH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013694">
        <property type="entry name" value="VIT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002035">
        <property type="entry name" value="VWF_A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036465">
        <property type="entry name" value="vWFA_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10338">
        <property type="entry name" value="INTER-ALPHA-TRYPSIN INHIBITOR HEAVY CHAIN FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10338:SF115">
        <property type="entry name" value="INTER-ALPHA-TRYPSIN INHIBITOR HEAVY CHAIN H3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06668">
        <property type="entry name" value="ITI_HC_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08487">
        <property type="entry name" value="VIT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00092">
        <property type="entry name" value="VWA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00609">
        <property type="entry name" value="VIT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00327">
        <property type="entry name" value="VWA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53300">
        <property type="entry name" value="vWA-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51468">
        <property type="entry name" value="VIT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50234">
        <property type="entry name" value="VWFA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="4" id="KW-0646">Protease inhibitor</keyword>
    <keyword evidence="7" id="KW-0654">Proteoglycan</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0964">Secreted</keyword>
    <keyword evidence="6" id="KW-0722">Serine protease inhibitor</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <feature type="domain" description="VIT" evidence="12">
        <location>
            <begin position="8"/>
            <end position="137"/>
        </location>
    </feature>
    <feature type="domain" description="VWFA" evidence="11">
        <location>
            <begin position="262"/>
            <end position="445"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010158"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022690"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022974"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00037051"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00039924"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50234"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS51468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029875.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="831" mass="93166" checksum="3CA1C17357B502EB" modified="2019-05-08" version="1">MEDGFLSHGGKRSADDLIVNGIEIYSMKIDSKVTSRFAHNVITSRAVNRANVSKEVVFDVDLPKTAFITNFSMTIDGVTYPGTIKEKEVAKKQYEKAVSQGQTAGLVRASGRKTEKFTVSVNIAAASKVTFELTYEELLKRNFGKYEMLIKVKPKQLINHFEIQANIFEPQGISELEAQGSFITNDLSQVIKKSFSGKKGHVSFKPTIDQQRTCANCSTSLLDGDFTIKYDVKRDRPGDLQIVNGYFVHFFAPTNLPNFPKNVVFVIDISGSMSGIGIQQTREALLKILEDVREDDYINFILFGSDVHVWKDTLLKATAENLKEAKDFVRAINTQGMTNLHGGLMKGIEMLTTAFETNSVPKRSASIIIMLTDGQPNEGVSDRQEIQMNVKKAIQGRYPLYNLAFGYGVDYNFLEKMALENKGLARRIYPQSDSAIQMEGFYEEVANPMLTEVELNYPENAISDLTQNIFKHYYDGSEIVVAGRLTDNDLDSLTADVKAQGAENDVMFSEQADVEETAKALQEQEYIFGDYIERLWAYLTIQQLLDKRTVAQGEEKENLTTEALELSLKYKFVTPLTSMVVTKPEDNEDKAAIADKPVEGTARAVPGTQSYPTHSYLYSRPPIPSLYTSVDGDPHFIISVPQKEDAICFNINEETGAVLNLINDPVTGIRVNGQLIGDKRVSSDAKIQNTYFGKLGIENKKLELKLEVTPEKITLQKGRKKTVFTWLDTVTLRQEGLTVIINRKKNLVLSMGDGASFVIVLHQVWKKHPLHQDFLGLYTLDSHKLSERTHGLLGQFFHHIDFNILEIHPGSDPKKPDATIVIKNNKLTVTR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPL2</accession>
    <name>A0A452IPL2_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="11">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="domain">
        <text evidence="6">The CKK domain binds microtubules.</text>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the CAMSAP1 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000033884.1">
        <property type="protein sequence ID" value="ENSGAGP00000029843.1"/>
        <property type="gene ID" value="ENSGAGG00000021541.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036449">
        <property type="term" value="C:microtubule minus-end"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051011">
        <property type="term" value="F:microtubule minus-end binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030507">
        <property type="term" value="F:spectrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031122">
        <property type="term" value="P:cytoplasmic microtubule organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007026">
        <property type="term" value="P:negative regulation of microtubule depolymerization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031175">
        <property type="term" value="P:neuron projection development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.20.360:FF:000001">
        <property type="entry name" value="Calmodulin-regulated spectrin-associated protein 3 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.20.360">
        <property type="entry name" value="CKK domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032940">
        <property type="entry name" value="CAMSAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031372">
        <property type="entry name" value="CAMSAP_CC1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022613">
        <property type="entry name" value="CH_CAMSAP_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001715">
        <property type="entry name" value="CH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036872">
        <property type="entry name" value="CH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014797">
        <property type="entry name" value="CKK_CAMSAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038209">
        <property type="entry name" value="CKK_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011033">
        <property type="entry name" value="PRC_barrel-like_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21595:SF2">
        <property type="entry name" value="CALMODULIN-REGULATED SPECTRIN-ASSOCIATED PROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21595">
        <property type="entry name" value="PATRONIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17095">
        <property type="entry name" value="CAMSAP_CC1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11971">
        <property type="entry name" value="CAMSAP_CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08683">
        <property type="entry name" value="CAMSAP_CKK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01051">
        <property type="entry name" value="CAMSAP_CKK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47576">
        <property type="entry name" value="Calponin-homology domain, CH-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50346">
        <property type="entry name" value="PRC-barrel domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50021">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51508">
        <property type="entry name" value="CKK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="5" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="3 6" id="KW-0493">Microtubule</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="8">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019486910" evidence="8">
        <location>
            <begin position="23"/>
            <end position="1255"/>
        </location>
    </feature>
    <feature type="domain" description="Calponin-homology (CH)" evidence="9">
        <location>
            <begin position="102"/>
            <end position="216"/>
        </location>
    </feature>
    <feature type="domain" description="CKK" evidence="10">
        <location>
            <begin position="1116"/>
            <end position="1250"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="54"/>
            <end position="80"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="426"/>
            <end position="574"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="598"/>
            <end position="654"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="698"/>
            <end position="993"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1062"/>
            <end position="1122"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="482"/>
            <end position="493"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="494"/>
            <end position="513"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="618"/>
            <end position="632"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="737"/>
            <end position="759"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="7">
        <location>
            <begin position="760"/>
            <end position="776"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="811"/>
            <end position="824"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="7">
        <location>
            <begin position="893"/>
            <end position="905"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="921"/>
            <end position="993"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00841"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS51508"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029843.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1255" mass="136685" checksum="5BD7979631B7D5A2" modified="2019-05-08" version="1">MGAHLAVIDSLMIAFAVESTHSLSAPPGTDLGAGSWEQKLLCWVDTVNRKLQESTEREGSQQLVPGTEGQSQSQSPASGPKCPTRWYWKLVPHAIAFCLKESGSKPPVIRYRKDKVLPKQTPCFTPVMGMKDLANGGAIAATIHYYCPDVVRLEDVCLKETMSVADSLYNLQLIQEFCIKYLGGCCPLALEDLLYVPPVLKINIGVFLAELFLCFEVLKPDFVRPKELRGLKDPPAVSDSLTPSSGNSNSGSLVFSFLHPLLPGGQPQSPLRGSLGSMHHSTSMSHVEGGFGKSWSKKQLSHPLSQAVSFSIPFGLDSDVDIVMGNPVGMLRSISSDSLAPVPTRLARSPGPLPEDAAQRLAKDEAPNGPVAPHRTVHKVALSAKGVVEREGPPVENGLADGYPDLPTIEEALQIIHSSERLHPEGAADGFYLHSPEPAKPTQAPEPTPALAVYRFHSGPPNGRAEAPTPPGNGSLDSDAEDPARLPGTKDDTSSGLSSLSSQPDSAGSSGAGVRMTSFAERKKKLAPPEGKTLQKSTGESSEAGAVPPEDSPGRSPALSSEMSQLGARLEEKRRAIEAQKKRIEAIFAKHRQRLGKSAFLQLKRRDGGEEEEEEEGEAKLSLEERLARLEAEEGGSLQVQAEEEGPGQGCPEKQVTFSSEIKGAPLDENLGDYNRAVAKLNTALSSLQLDMHRLSQQQQRLLQEKKPSQAWVIPAPKGPAPRASREFVPPRSVELSSSPSPSPSRXXXXXXXXXXPAAPRSPQPAPKKAAPTPPKSPKHARPVELKLPPLTRVLTPPHNVDTLPHLRKFSPSQVPMQTRSSIHFSEEVAGPEQPLESPELETQGNLRPVASVVPQGGVGSARVSGDGTSDVSSPGERRSSLIEIPLSSLQAEEGDGDDSLEESLTEVMDMEPRAGLGFFFKDDEKPEDEMAQKRASLLERQQRRNEEARQRKQWLEAEKEQKEEAARLQAEERPRPEEEASPRRGDFTRQEYQRRHQLKLMEDLDKVLRQKPTTVRALKKGRPKTVFCDDSALARSPVKGLLGSRLSKVYSQSTLSLSTVANEPGNSLTIKRPSRAASPSGLMSPSRLLGSQSRERDWENASTASSPASVPEYTGPKLYKEPSAKSNKHIIHNALSHCCLAGRVNEPHKNKILEEMEKSKAHHFLILFRDSSCQFRALYTPAGETEELTRLTGYGPRTISQAMIEGIYKYNSDRKRFTQIPAKTMSMNVDAITIQGHLWQTKKPGTPKKPGTPK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPL4</accession>
    <name>A0A452IPL4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Filensin</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="14">Beaded filament structural protein 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Peripheral membrane protein</topology>
            <orientation evidence="2">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Cell cortex</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPL4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033990.1">
        <property type="protein sequence ID" value="ENSGAGP00000029931.1"/>
        <property type="gene ID" value="ENSGAGG00000021598.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005938">
        <property type="term" value="C:cell cortex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005882">
        <property type="term" value="C:intermediate filament"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005212">
        <property type="term" value="F:structural constituent of eye lens"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070307">
        <property type="term" value="P:lens fiber cell development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.1160:FF:000009">
        <property type="entry name" value="filensin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.170">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.1160">
        <property type="entry name" value="Vasodilator-stimulated phosphoprotein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042358">
        <property type="entry name" value="BFSP1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039008">
        <property type="entry name" value="IF_rod_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14069">
        <property type="entry name" value="FILENSIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14069:SF0">
        <property type="entry name" value="FILENSIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00038">
        <property type="entry name" value="Filament"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01391">
        <property type="entry name" value="Filament"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF64593">
        <property type="entry name" value="Intermediate filament protein, coiled coil region"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51842">
        <property type="entry name" value="IF_ROD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11 15" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="13" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="8" id="KW-0273">Eye lens protein</keyword>
    <keyword evidence="10" id="KW-0403">Intermediate filament</keyword>
    <keyword evidence="12" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="IF rod" evidence="17">
        <location>
            <begin position="38"/>
            <end position="318"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="489"/>
            <end position="575"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="15">
        <location>
            <begin position="90"/>
            <end position="159"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="15">
        <location>
            <begin position="255"/>
            <end position="317"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="16">
        <location>
            <begin position="547"/>
            <end position="556"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004413"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004544"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00019025"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022754"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00031415"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS51842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029931.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="620" mass="71782" checksum="93F9E3C05F53E30B" modified="2019-05-08" version="1">MYRSSYLREVRKEKYERSDAYDELLGTSDFSSLAQAQGLENLQELNERFANYINRARMLEQRNTIFRKQLETFQRMDVLAGLEEAFAEQIELNRLRIRELSADRAKLEREDKDSQRMLDEYRNKYRNEREYQQKLKETLERLNKEADEALLCNLELQIESQFLQDDINATKDRYKKNIMEIQTYVSILQQIIQMTPSVTTITVGISEERLIAERRIPILQSQLEEYKSILCQLQTQKLKLQTETTMLEQAIKSTQESYDDEVQLYNEQIEALRKEIEDAERTLEKYTSDCHQLVIYQQSLENELERYKRIIENEDNRLNSAIIGTPVALFTQSYRPCQVTTWRRRDITLVMQEIASAKPRQKGLAKKISRRKEITSKDLDGFSPENSYERTLEGFHQDQLQFKHESTVICEPKQEGLELDEQEMVPEDVPDGARISKAFDELCNTVREKIRCYKRPEPTKGRYVLVTGEASYDHPCFYSSSTPAGGGIIVSTSNGKIPNGGEVEPIPELPEPSEPSANEKGEPSTQPDITEPEAAPPAGVVSPSEPDVFEETEYDKEDQREGQAFREPSLPSSMSYEKVEVVESIEKFSDDRIQTYEETAMIVETTIEKTSKKKLGDKGS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPL7</accession>
    <name>A0A452IPL7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">NACHT domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPL7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033823.1">
        <property type="protein sequence ID" value="ENSGAGP00000029786.1"/>
        <property type="gene ID" value="ENSGAGG00000021504.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006954">
        <property type="term" value="P:inflammatory response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045087">
        <property type="term" value="P:innate immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007111">
        <property type="entry name" value="NACHT_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041267">
        <property type="entry name" value="NLRP_HD2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050637">
        <property type="entry name" value="NLRP_innate_immun_reg"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041075">
        <property type="entry name" value="NOD1/2_WH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45690">
        <property type="entry name" value="NACHT, LRR AND PYD DOMAINS-CONTAINING PROTEIN 12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45690:SF19">
        <property type="entry name" value="NACHT, LRR AND PYD DOMAINS-CONTAINING PROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05729">
        <property type="entry name" value="NACHT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17776">
        <property type="entry name" value="NLRC4_HD2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17779">
        <property type="entry name" value="WHD_NOD2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50837">
        <property type="entry name" value="NACHT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-0395">Inflammatory response</keyword>
    <keyword evidence="4" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="NACHT" evidence="8">
        <location>
            <begin position="41"/>
            <end position="173"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023198"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029786.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="549" mass="63347" checksum="F86E43FE4D0FC55C" modified="2019-05-08" version="1">TFVYNGLSGTFVTLGNFHYRLVSVITMGTLFKPYKDGQTPQIVVLVGAAGIGKTITSKKIMLDWAAGELYKEMFDYVFYINCREMNLLTKEVSVADMILKHCPNNKTSINHILGQSDKLLFIIDELRFSFDQSEDNLCSERWEKKPVEIILSSLFRKTVLPESYLLITTRPAALEKLGRCERYAEILGFSEADRKEYFHKFFGNEKQARQAYRFVRENEILFTMCFVPIMCWIICTVLKQQLEKGEDLAQASKTATGMYVLYLSSLVNPPGSNSKPCKKASLRGLCSLAADGIWKQKILFEEEEIKMLGLDQADALPLFLNENIFQKDIDTECLYSFIHLSFQEFFAALFYVLEEDETMEDSGTPKKDVKTLLENYGNSRNYLMLTVRFLYGLLNEKSIKDMEEKFGCKISPKIKLDLIKWIQTYKQTVLSVPSVYDKIICQLEEFRCLYEIQDENFMKNALEPFTKIVICRNNLTPMDQIVLSFCVNNCHRLVSLEMYACTFRFEDRKKEVPQRHEELDQSLFPVSQYHVSCHSSSTNSSAGLQNNLN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPL9</accession>
    <name>A0A452IPL9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="5">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1 4">Belongs to the heat shock protein 70 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPL9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033937.1">
        <property type="protein sequence ID" value="ENSGAGP00000029885.1"/>
        <property type="gene ID" value="ENSGAGG00000021572.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140662">
        <property type="term" value="F:ATP-dependent protein folding chaperone"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.30.30:FF:000001">
        <property type="entry name" value="heat shock 70 kDa protein-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.640.10:FF:000134">
        <property type="entry name" value="Heat shock cognate 71 kDa protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.420.40:FF:000026">
        <property type="entry name" value="Heat shock protein 70"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.30.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.420.40">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.640.10">
        <property type="entry name" value="Actin, Chain A, domain 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.34.10">
        <property type="entry name" value="Substrate Binding Domain Of DNAk, Chain A, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043129">
        <property type="entry name" value="ATPase_NBD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018181">
        <property type="entry name" value="Heat_shock_70_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029048">
        <property type="entry name" value="HSP70_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029047">
        <property type="entry name" value="HSP70_peptide-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013126">
        <property type="entry name" value="Hsp_70_fam"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19375">
        <property type="entry name" value="HEAT SHOCK PROTEIN 70KDA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00012">
        <property type="entry name" value="HSP70"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00301">
        <property type="entry name" value="HEATSHOCK70"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53067">
        <property type="entry name" value="Actin-like ATPase domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF100934">
        <property type="entry name" value="Heat shock protein 70kD (HSP70), C-terminal subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF100920">
        <property type="entry name" value="Heat shock protein 70kD (HSP70), peptide-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00297">
        <property type="entry name" value="HSP70_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00329">
        <property type="entry name" value="HSP70_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 4" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="2 4" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007381"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="RuleBase" id="RU003322"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029885.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="621" mass="68493" checksum="ECE5840D13DF9CD4" modified="2019-05-08" version="1">MPALKHPAIGIDLGTTKSCVGLFRDGKVEIIVSNQGIRTTPSCVAFTETGRLIGDEAKNQAALNSPNIIFSAKRLIGRTSEDPVVQADAEFWPFQIVKAEGKPQVQVSYKGKEKVFYPEEISAMVLGKLKEMAEDYLGHPITHAIITVPAHFNDSQRQATKDAGTIAGLNVLRILSETTAAALAYSLDNPSNGGHNVLIFDLGGGTFNVSVLTIDGGILEVKATSGDTHLGGDDFDNCLVSFFIDEFLKKHQENISKDRRAVWRLRMACEAAKCTLSSNIKASIDLYCLYKGIDFHADITRAQFEELCADLFRGTLKPVERALQDAKLSQAQIHDVVLVGASTRIPKIQELLQDFFGGRQLNKSLNLDEAVACGAAVQAAILIGERSEVTRGLLALDVVSMSLGIETDTGKMHTLIRRNTAMPAVVTRTFTTFDDHQPDLLIRAYEGERATAKENHLLGRFLLKGISPAPRGMVLVEATFYINADGILTVFALDKVASNANPITVDANKGRLSREEIQRLAAEAEKYRAEDRAQKWKLATKNYLQSCAFSMKRTSEDILNYFTLQAKKKVMEKCQLAFSWLSPTGWLKRRRSERVKQDKEMLCCPLLSGILTPKEDPSLKW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPM0</accession>
    <name>A0A452IPM0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="19">Huntingtin-interacting protein 1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="20">Huntingtin-interacting protein I</fullName>
        </alternativeName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="17">Plays a role in clathrin-mediated endocytosis and trafficking. Involved in regulating AMPA receptor trafficking in the central nervous system in an NMDA-dependent manner. Regulates presynaptic nerve terminal activity. Enhances androgen receptor (AR)-mediated transcription. May act as a proapoptotic protein that induces cell death by acting through the intrinsic apoptosis pathway. Binds 3-phosphoinositides (via ENTH domain). May act through the ENTH domain to promote cell survival by stabilizing receptor tyrosine kinases following ligand-induced endocytosis. May play a functional role in the cell filament networks. May be required for differentiation, proliferation, and/or survival of somatic and germline progenitors.</text>
    </comment>
    <comment type="subunit">
        <text evidence="18">Homodimer. Binds actin. Binds HTT (via N-terminus). This interaction is restricted to the brain. Binds to IFT57. In normal conditions, it poorly interacts with IFT57, HIP1 being strongly associated with HTT. However, in mutant HTT proteins with a long poly-Gln region, interaction between HTT and HIP1 is inhibited, promoting the interaction between HIP1 and IFT57. Interacts with CLTB (via N-terminus). Interacts (via coiled coil domain) with AR. Interacts with AP2A1, AP2A2, CLTC and HIP1R. Interacts with GRIA1, GRIN2A and GRIN2B.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasmic vesicle membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the SLA2 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPM0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033995.1">
        <property type="protein sequence ID" value="ENSGAGP00000029936.1"/>
        <property type="gene ID" value="ENSGAGG00000021602.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030136">
        <property type="term" value="C:clathrin-coated vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030864">
        <property type="term" value="C:cortical actin cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030659">
        <property type="term" value="C:cytoplasmic vesicle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098793">
        <property type="term" value="C:presynapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035615">
        <property type="term" value="F:clathrin adaptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032051">
        <property type="term" value="F:clathrin light chain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043325">
        <property type="term" value="F:phosphatidylinositol-3,4-bisphosphate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0080025">
        <property type="term" value="F:phosphatidylinositol-3,5-bisphosphate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007015">
        <property type="term" value="P:actin filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006915">
        <property type="term" value="P:apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048268">
        <property type="term" value="P:clathrin coat assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006897">
        <property type="term" value="P:endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048260">
        <property type="term" value="P:positive regulation of receptor-mediated endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010646">
        <property type="term" value="P:regulation of cell communication"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0023051">
        <property type="term" value="P:regulation of signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd17013">
        <property type="entry name" value="ANTH_N_HIP1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1410.10:FF:000002">
        <property type="entry name" value="Huntingtin interacting protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.1700:FF:000002">
        <property type="entry name" value="Huntingtin interacting protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.90:FF:000022">
        <property type="entry name" value="huntingtin-interacting protein 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.1700">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.90">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1410.10">
        <property type="entry name" value="I/LWEQ domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011417">
        <property type="entry name" value="ANTH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013809">
        <property type="entry name" value="ENTH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008942">
        <property type="entry name" value="ENTH_VHS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032422">
        <property type="entry name" value="HIP1_clath-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035964">
        <property type="entry name" value="I/LWEQ_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002558">
        <property type="entry name" value="ILWEQ_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR030224">
        <property type="entry name" value="Sla2_fam"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10407">
        <property type="entry name" value="HUNTINGTIN INTERACTING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10407:SF14">
        <property type="entry name" value="HUNTINGTIN-INTERACTING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07651">
        <property type="entry name" value="ANTH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16515">
        <property type="entry name" value="HIP1_clath_bdg"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01608">
        <property type="entry name" value="I_LWEQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00273">
        <property type="entry name" value="ENTH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00307">
        <property type="entry name" value="ILWEQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48464">
        <property type="entry name" value="ENTH/VHS domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109885">
        <property type="entry name" value="I/LWEQ domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50942">
        <property type="entry name" value="ENTH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50945">
        <property type="entry name" value="I_LWEQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="14" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="12" id="KW-0010">Activator</keyword>
    <keyword evidence="7" id="KW-0053">Apoptosis</keyword>
    <keyword evidence="10 21" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="16" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="8" id="KW-0221">Differentiation</keyword>
    <keyword evidence="6" id="KW-0254">Endocytosis</keyword>
    <keyword evidence="11" id="KW-0472">Membrane</keyword>
    <keyword evidence="15" id="KW-0539">Nucleus</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="13" id="KW-0804">Transcription</keyword>
    <keyword evidence="9" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="ENTH" evidence="22">
        <location>
            <begin position="1"/>
            <end position="126"/>
        </location>
    </feature>
    <feature type="domain" description="I/LWEQ" evidence="23">
        <location>
            <begin position="717"/>
            <end position="958"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="21">
        <location>
            <begin position="334"/>
            <end position="480"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="21">
        <location>
            <begin position="509"/>
            <end position="561"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="21">
        <location>
            <begin position="924"/>
            <end position="953"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004156"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00010135"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022583"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022703"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00055592"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00062629"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00073538"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00080332"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS50942"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS50945"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029936.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="970" mass="108699" checksum="710176A2F61098F4" modified="2019-05-08" version="1">MEVGKMTVSINKAINTQEVAVKEKHARTCILGTHHEKGAQTFWSVVNRLPLSSNAVLCWKFCHVFHKLLRDGHPNVLKDSVRYKNELNDMSRMWSHLSEGYGQLCSIYLKLLRTKMEFHTKNPWFPGNLQMSDRQLDEAGENDVNNFFQLTVEMFDYLECELNLFQTVFSSLDMSRSVSVTAAGQCRLAPLIQVILDCSHLYDYTVKLLFKLHSCLPADTLQGHRDRFLEQFRKLKDLFYRSSNLQYFKRLIQIPQLPENPPNFLRASALSEHISPVVVIPAEASSPDSEPITDLVETDTASQQSLFDNKFDDIFGSSFSNDPFNFSGQNGMNKDDKDRLIEQLYQEISALKEELDNFKAESHRLGVQLRGRISELEAELAEQQHLKQQALDESDFLRTELEELKKQWEDTEKAQRSLTEIERKAQANEQRYTKLKEKYSELVQNHADLLRKTQEQAEVLETLKRELTASKQELQALQSTLQSSLQSEAERSTRTVGLEQERDTLVVAVTQRSQEVTALQAELQQLKDALDSEKESSSKSLEILQTRLGEKESREQALQQRLLDEQFALLQCTVQEAEQMVQDSLNRLDDPAHISCTSSADYLLCRTLAASDSIERLQGAHGKYLSNRSDVSSLLPCVALFAHLISDTILQGSATSHMVPMEPADLLSEMCKQCGSEALHFLGALKDEASLGGANCTAVRNCLSQISALGEELRPKGLDIEQEELGDLVDKEMAATSAAVETAVARIEDMLSKSRAGDTGVKLEVNERILGSCTDLMQAIQVLVLASKDLQREIVESGRGAASPKEFYARNSRWTEGLISASKAVGWGATVMVDAADLVVQGKGKFEELMVCSHEIAASTAQLVAASKVKADKDSANLARLRQASRGVNQATAGVVASTKSGKSQIEEKDSMDFSSMTLTQIKRQEMDSQVRVLELENQLQKERQKLGELRKKHYKLAGVAEGWDEDAAE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPM5</accession>
    <name>A0A452IPM5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Nck-associated protein 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="6">Cell projection</location>
            <location evidence="6">Lamellipodium membrane</location>
            <topology evidence="6">Single-pass membrane protein</topology>
            <orientation evidence="6">Cytoplasmic side</orientation>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="7">Belongs to the HEM-1/HEM-2 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPM5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034000.1">
        <property type="protein sequence ID" value="ENSGAGP00000029941.1"/>
        <property type="gene ID" value="ENSGAGG00000019065.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031258">
        <property type="term" value="C:lamellipodium membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031209">
        <property type="term" value="C:SCAR complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030031">
        <property type="term" value="P:cell projection assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030866">
        <property type="term" value="P:cortical actin cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048812">
        <property type="term" value="P:neuron projection morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019137">
        <property type="entry name" value="Nck-associated_protein-1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12093">
        <property type="entry name" value="NCK-ASSOCIATED PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12093:SF11">
        <property type="entry name" value="NCK-ASSOCIATED PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09735">
        <property type="entry name" value="Nckap1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="1" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="5" id="KW-0966">Cell projection</keyword>
    <keyword evidence="4" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="3" id="KW-1133">Transmembrane helix</keyword>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="530"/>
            <end position="555"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="541"/>
            <end position="555"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00037839"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00037947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00039689"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029941.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="979" mass="111961" checksum="80C8384A5AD4F617" modified="2025-10-08" version="2">MNGWETKREMIFSTVNFDLTKNYLDLIITYTTLMILLSRIEERKAIIGLYNYAHEMTHGASDREYPRLGQMIVDYENPLKKMMEEFVPHSKSLSDALISLQMVYPRRNLSADQWRNAQLLSLISAPSTMLNPAQSDTMPCEYLSLDAMEKWIIFGFILCHGILNSDATALNLWKLALQSSSCLSLFRDEVFHIHKAAEDLFVNIRGYNKRINDIRECKEAAVSHAGSMHRERRKFLRSALKELATVLSDQPGLLGPKALFVFMALSFARDEIIWLLRHADNMPKKSADDFIDKHIAELIFYMEELRAHVRKYGPVMQRYYVQYLSGFDAVVLNELVQNLSVCPEDESIIMSSFVNTMTSLSVKQVEDGEVFDFRGMRLDWFRLQAYTSVSKASLSLADHRELGKMMNTIIFHTKMVDSLVEMLVETSDLSIFCFYSRAFEKMFQQCLELPSQSRYSIAFPLLCTHFMSCTHELCPEERHHIGDRSLSLCNMFLDEMAKQARNLITDICTEQCTLSDQLLPKHCAKTISQAVNKKSKKQTGKKGEPEREKPGVESMRKNRLVVTNLDKLHTALSELCFSINYVPNMVVWEHTFTPREYLTSHLEIRFTKSIVGMTMYNQATQEIAKPSELLTSVRAYMTVLQSIENYVQIDITRVFNNVLLQQTQHLDSHGEPTITSLYTNWYLETLLRQVSNGHIAYFPAMKAFVNLPTENELTFNAEEYSDISEMRALSELLGPYGMKFLSESLMWHISSQVAELKKLVVENVEVLTQMRTSFDKPDQMAALFKRLSSVDSVLKRMTIIGVILSFRSLAQEALRDVLSYHIPFLVSSIEDFKDHIPRETDMKVAMNVYELSSAAGLPCEIDPALVVALSSQKSGHCNNIHCLAKAINQIAAALFTIHKGSIEDRLKEFLALASSSLLKIGQETDKTTTRNRESVYLLLDMIVQESPFLTMDLLESCFPYVLLRNAYHAVYKQSVTSSA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPP1</accession>
    <name>A0A452IPP1_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="21">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="15">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
        <text evidence="15">Binds 1 zinc ion per subunit.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="17">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPP1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033852.1">
        <property type="protein sequence ID" value="ENSGAGP00000029813.1"/>
        <property type="gene ID" value="ENSGAGG00000021496.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031012">
        <property type="term" value="C:extracellular matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004222">
        <property type="term" value="F:metalloendopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030198">
        <property type="term" value="P:extracellular matrix organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
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    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
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    <dbReference type="CDD" id="cd04273">
        <property type="entry name" value="ZnMc_ADAMTS_like"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="A disintegrin and metalloproteinase with thrombospondin motifs 1"/>
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        <property type="entry name" value="A disintegrin and metalloproteinase with thrombospondin motifs 1"/>
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        <property type="entry name" value="A disintegrin and metalloproteinase with thrombospondin motifs 3"/>
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        <property type="match status" value="1"/>
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        <property type="match status" value="1"/>
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    <dbReference type="Gene3D" id="3.40.390.10">
        <property type="entry name" value="Collagenase (Catalytic Domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.20.100.10">
        <property type="entry name" value="Thrombospondin type-1 (TSP1) repeat"/>
        <property type="match status" value="4"/>
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    <dbReference type="InterPro" id="IPR013273">
        <property type="entry name" value="ADAMTS/ADAMTS-like"/>
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    <dbReference type="InterPro" id="IPR050439">
        <property type="entry name" value="ADAMTS_ADAMTS-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041645">
        <property type="entry name" value="ADAMTS_CR_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045371">
        <property type="entry name" value="ADAMTS_CR_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010294">
        <property type="entry name" value="ADAMTS_spacer1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024079">
        <property type="entry name" value="MetalloPept_cat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001590">
        <property type="entry name" value="Peptidase_M12B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002870">
        <property type="entry name" value="Peptidase_M12B_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010909">
        <property type="entry name" value="PLAC"/>
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    <dbReference type="InterPro" id="IPR000884">
        <property type="entry name" value="TSP1_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036383">
        <property type="entry name" value="TSP1_rpt_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13723:SF158">
        <property type="entry name" value="A DISINTEGRIN AND METALLOPROTEINASE WITH THROMBOSPONDIN MOTIFS 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13723">
        <property type="entry name" value="ADAMTS A DISINTEGRIN AND METALLOPROTEASE WITH THROMBOSPONDIN MOTIFS PROTEASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17771">
        <property type="entry name" value="ADAMTS_CR_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19236">
        <property type="entry name" value="ADAMTS_CR_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05986">
        <property type="entry name" value="ADAMTS_spacer1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01562">
        <property type="entry name" value="Pep_M12B_propep"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01421">
        <property type="entry name" value="Reprolysin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19030">
        <property type="entry name" value="TSP1_ADAMTS"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00090">
        <property type="entry name" value="TSP_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01857">
        <property type="entry name" value="ADAMTSFAMILY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00209">
        <property type="entry name" value="TSP1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55486">
        <property type="entry name" value="Metalloproteases ('zincins'), catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF82895">
        <property type="entry name" value="TSP-1 type 1 repeat"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50215">
        <property type="entry name" value="ADAM_MEPRO"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="PLAC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50092">
        <property type="entry name" value="TSP1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="15" id="KW-0106">Calcium</keyword>
    <keyword evidence="12 16" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="5 15 17" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0482">Metalloprotease</keyword>
    <keyword evidence="4" id="KW-0645">Protease</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="9 15 17" id="KW-0862">Zinc</keyword>
    <keyword evidence="11" id="KW-0865">Zymogen</keyword>
    <feature type="domain" description="Peptidase M12B" evidence="19">
        <location>
            <begin position="247"/>
            <end position="451"/>
        </location>
    </feature>
    <feature type="domain" description="PLAC" evidence="20">
        <location>
            <begin position="1000"/>
            <end position="1043"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1109"/>
            <end position="1128"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1147"/>
            <end position="1180"/>
        </location>
    </feature>
    <feature type="active site" evidence="14 17">
        <location>
            <position position="390"/>
        </location>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="250"/>
        </location>
        <ligand>
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="250"/>
        </location>
        <ligand>
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
            <label>2</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="342"/>
        </location>
        <ligand>
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15 17">
        <location>
            <position position="389"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <note>catalytic</note>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15 17">
        <location>
            <position position="393"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <note>catalytic</note>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15 17">
        <location>
            <position position="399"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <note>catalytic</note>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="446"/>
        </location>
        <ligand>
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="449"/>
        </location>
        <ligand>
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
            <label>2</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="449"/>
        </location>
        <ligand>
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="324"/>
            <end position="373"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="367"/>
            <end position="446"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="406"/>
            <end position="432"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="473"/>
            <end position="498"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="484"/>
            <end position="507"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="493"/>
            <end position="526"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="520"/>
            <end position="531"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="554"/>
            <end position="591"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="558"/>
            <end position="596"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="569"/>
            <end position="581"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
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        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023049"/>
        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00023145"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="PIRSR" id="PIRSR613273-1"/>
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    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PIRSR" id="PIRSR613273-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PIRSR" id="PIRSR613273-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00276"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
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        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
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        </source>
    </evidence>
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        <source>
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    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1180" mass="133676" checksum="52005AF7FE18513F" modified="2019-05-08" version="1">MHLIKYVYLEAFLWHTSVSEPLNSKDVSSDLPKIRPRLYGLVTPISTSADGSFISNILSANHQRRLTRDVSQSPKQLYFNITAFGREFHLRLRPNTHLVAPEAVVEWYEDSVESGNGTETRSTKEHIWKKEPLWTNCAYVGDITDIPGASVAISNCDGLAGMIRTDKDEYFIEPLERGKQMEEERGRIHMVYRRSAVTQDPGDMLLDFHTEASLLSGLDGLDSMYSNIEQQLNETLRRRRHAGDNDYNIEVLLGVDDSVVRFHGKEHVQNYLLTLMNIVNEIYHDESLGVHINVVLVRMIMLGYAKSVSLIERGNPSRSLENVCRWAYQQQKSDPNHSEHHDHAIFLTRQDFGPAGMQGYAPVTGMCHPVRSCTLNHEDGFSSAFVVAHETGHVLGMEHDGQGNRCGDETAMGSVMAPLVQAAFHRYHWSRCSGQELKRYIHSYDCLLDDPFEHDWPKLPELPGINYSMDEQCRFDFGVGYKMCTAFRTFDPCKQLWCSHPDNPYFCKTKKGPPLDGTECAPGKWCYKGHCMWKNANQLKQDGNWGSWTKFGSCSRTCGTGVRFRTRQCNNPMPVNGGQDCASVNFEYQLCSTEECPKHFEDFRAQQCQQRNSHFEYQKSKHHWLPYEHPDANKRCHLYCQSKEIGDIAYMKQLAHDGTRCSYKDPYSICVRGECVKVGCDKEIGSNKVEDKCGVCGGDNSHCRTVKGTFTRTPKKPGYLKMFDIPPGARHVFIQEDEASPHILAIKNQATGHYILNGKGEEAKSRSFIDLGVEWEYNTEDDIETLHTDGPLHDAVVVLVIPQENDTRSSLTYKYIIHEDSVPTINSNNVLQDEIDTFEWALKSWSQCSKPCAGGFQYTKYGCRRKSDNKMVHRSFCEASKKPKPIRRMLWMDEMWASDENEWEYCTKTCGHSGYQIRTVRCIQPLHDGANRSVNIKYCSGDRPESRRPCNRTPCPAQWKAGPWAECSVTCGEGTESRQVTCRAGDQCDGEKPESVRVCKLAPCNDEPCLGDKSIFCQMEVLARYCSIPGYNKLCCESCSKRSSTLPPPYLTEAAETKEEVMFHPRDLPRALAMPTSLVPQHAEVADENRSSVGRMPMEDDIDVLVSTSNSKSKGAELPQRRAYQAGSQTSRLIAMPYQSSTKIAVNNTSAGALSPARTSRKNEKHVEKRHALRSSTVER</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPP4</accession>
    <name>A0A452IPP4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">PDZ domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPP4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034140.1">
        <property type="protein sequence ID" value="ENSGAGP00000030068.1"/>
        <property type="gene ID" value="ENSGAGG00000021685.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098887">
        <property type="term" value="P:neurotransmitter receptor transport, endosome to postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd06684">
        <property type="entry name" value="PDZ3_GRIP1-2-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd06686">
        <property type="entry name" value="PDZ4_GRIP1-2-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd06682">
        <property type="entry name" value="PDZ5_GRIP1-2-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd06685">
        <property type="entry name" value="PDZ7_GRIP1-2-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000022">
        <property type="entry name" value="Glutamate receptor interacting protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000025">
        <property type="entry name" value="Glutamate receptor interacting protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000031">
        <property type="entry name" value="Glutamate receptor interacting protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000035">
        <property type="entry name" value="Glutamate receptor interacting protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.42.10">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043545">
        <property type="entry name" value="GRIP1/2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001478">
        <property type="entry name" value="PDZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041489">
        <property type="entry name" value="PDZ_6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036034">
        <property type="entry name" value="PDZ_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46227:SF3">
        <property type="entry name" value="GLUTAMATE RECEPTOR-INTERACTING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46227">
        <property type="entry name" value="GLUTAMATE RECEPTOR-INTERACTING PROTEIN GRIP"/>
        <property type="match status" value="1"/>
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    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="PDZ" evidence="5">
        <location>
            <begin position="15"/>
            <end position="59"/>
        </location>
    </feature>
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            <begin position="73"/>
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            <begin position="728"/>
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            <begin position="515"/>
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            <begin position="658"/>
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            <begin position="658"/>
            <end position="681"/>
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        <location>
            <begin position="688"/>
            <end position="698"/>
        </location>
    </feature>
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        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50106"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030068.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="854" mass="93462" checksum="6D2E97ECC1562953" modified="2019-05-08" version="1">MHFIRLSCLFPNCLEGTIKPGDRLLSVDGIRLLGTSHAEAMSILKQCGQEATLLIEYDVSVMDSVATASGPLLVEVAKTPGAALGVALTTSMCCNKQVIVIDKIKSASFADRCGALHVGDHILSIDGTSMEYCTLAEATQFLANTADNVKLEILPHHQTRLALKGAEHVKVQRSNRQLTWDSCANNHSSLLTYHHYNTYHPDHCRMPALKFQKTPPQKNLPALVSSSFSPTSMSAYSLSSLNMGTLPRSLYSTSPRGTMMRRRMKKKDFKSSLSLASSTVGLAGQVVHTETTEVVLTADPIVGFGIQLQGSVFATETLSSPPLISYIECDSPAERCGVLQIGDRLMAINGIPTEDSTFEEANQLLRDSSITNKVTLEIEFDVAESVIPSSGTFHVKLPKKHNVELGITISSPSSRKPGDPLVISDIKKGSVAHRTGTLELGDKLLAIDNIRLDNCSMEDAVQILQHCEDLVKLKIRKDEDNSDGEFEAIIMFIFKPYLAASPKKFSVSGHLSELSDAEDETSSTQKPGKLSDIYSTTVPSVDSAVESWDGSGIDTSFGSQGPTYQASGYNFSAYEWRSPKQRDNLSPPSRPQNHPFHDTGLSDDEWDRPTASGFSGAHDNSETDQEENFWSQALEDLETCGQSGILRELEATIMSGSTMSLNHENPQPRSQLGRQASFQERSNVRPHYSQTARSNTLPSDVGRKSVVLRKIKQEMKEIMSPTPVELHKVTLYKESDIEDFGFSVSDGLLEKGVYVKNIRPAGPGDLGGLKPYDRLLQVNHVRTRDFDCCLVVPLIAESGNKLELVISRNPLASQKVTSDQQTLAGGEWNDQNNAFLQQTGHTASVETRDPTNTI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ10</accession>
    <name>A0A452IQ10_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="8">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQ10"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034276.1">
        <property type="protein sequence ID" value="ENSGAGP00000030195.1"/>
        <property type="gene ID" value="ENSGAGG00000021279.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
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        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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    <dbReference type="GO" id="GO:0050684">
        <property type="term" value="P:regulation of mRNA processing"/>
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    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd12678">
        <property type="entry name" value="RRM_SLTM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.330:FF:000238">
        <property type="entry name" value="SAFB-like transcription modulator isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.330">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.720.30">
        <property type="entry name" value="SAP domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012677">
        <property type="entry name" value="Nucleotide-bd_a/b_plait_sf"/>
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    <dbReference type="InterPro" id="IPR035979">
        <property type="entry name" value="RBD_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000504">
        <property type="entry name" value="RRM_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051738">
        <property type="entry name" value="SAF_Modulators"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003034">
        <property type="entry name" value="SAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036361">
        <property type="entry name" value="SAP_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15683:SF5">
        <property type="entry name" value="SAFB-LIKE TRANSCRIPTION MODULATOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15683">
        <property type="entry name" value="SCAFFOLD ATTACHMENT FACTOR B-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00076">
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    </dbReference>
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        <property type="match status" value="1"/>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="RNA-binding domain, RBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF68906">
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    </dbReference>
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    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 4" id="KW-0694">RNA-binding</keyword>
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        <location>
            <begin position="26"/>
            <end position="60"/>
        </location>
    </feature>
    <feature type="domain" description="RRM" evidence="6">
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            <begin position="316"/>
            <end position="394"/>
        </location>
    </feature>
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        </location>
    </feature>
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            <begin position="1"/>
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        </location>
    </feature>
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            <begin position="92"/>
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        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
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            <begin position="152"/>
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        </location>
    </feature>
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        <location>
            <begin position="202"/>
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        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="222"/>
            <end position="271"/>
        </location>
    </feature>
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            <begin position="275"/>
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        </location>
    </feature>
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            <end position="302"/>
        </location>
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        </location>
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        </location>
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        </location>
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        <location>
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        </location>
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        </location>
    </feature>
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        <location>
            <begin position="703"/>
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        </location>
    </feature>
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        <location>
            <begin position="724"/>
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        </location>
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        <location>
            <begin position="807"/>
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        </location>
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        <location>
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        <location>
            <begin position="928"/>
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        </location>
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        <location>
            <begin position="954"/>
            <end position="966"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00176"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50102"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50800"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030195.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="966" mass="109953" checksum="7CCEC54C843FCA18" modified="2019-05-08" version="1">MAAAAAVSSPAAAPSVGPAQMESKKITDLRVIDLKSELKRRNLDITGVKTVLISRLKQVRGRCRPRNASNASDQDGNDELKDFEEAGENEEENSHSKELRPSEKKKYSDSKQRKTAEDAEKDFESQVPYKAGSGDGIQEVSKPLPSEESLAEADHSAHEEMEANTSMKEAEDDNISVTIQAEDAITLDFDGDDLLETGKNVKITDSEASKAKDGQDTIAQSLEKERKDYEMTENHKDGKKEDCVKGDPVKKEAREGSKKAESGDKEKDTLKKGPSSTGASGQAKSSAKESKESKTTSKDDKGSASSVSGSSGSSTRNLWVSGLSSNTKAADLKNLFGKYGKVLSAKVVTNARSPGAKCYGIVTMSSSTEVARCIAHLHRTELHGQQISVEKVKGDPSKKELKKEIDEKSGSGRNMGDKKTASIDKTSKTQSAKKEEKKSDKSEKKESKEAKKTEGKDEKNDNGSSGPNQESTKKTEEKKRISGKSPGQMVVLDQTKGDQGHTRTVRRGRFDKVSICKNKERFIQDKMKFREYRGRKDILPFEKMKEQRMREHMVRLERIRRAVELRRRRELAERERRERERIRIIREREERERLQRERERLEIERQKLERERMERERLERERIRIEQERRKEAERIAREREELRRQQQQLRYEQEKRNSLKRPRDVDHRRDDPYWNENKKMALDTDARFSHGSDYSRQQNRFNDFDHRERGRYPEGSSVPSSSFERRERFVNQSEGKKTRPTARREEPGFERYPKNFSESRRNEPPQPRNELRDTDRREVRGDRDERRTVIIHDRPEISHGRHPRETGSNPPRQTNWKNEGGISTDKRDARGERPERSGREVSGHVRSAAPGSRSSTSGYGSREGERGVIGERGSGGQHYNEDRHVVERHSRETGPRKEWHGPSSQGSGYHDARRMGDGRGGGGMMSPHTSNSSPINRVVQITGSSMQRGSGSGFKPFKSGPPRRF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ14</accession>
    <name>A0A452IQ14_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Rho-GAP domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IQ14"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034281.1">
        <property type="protein sequence ID" value="ENSGAGP00000030200.1"/>
        <property type="gene ID" value="ENSGAGG00000021783.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:2001136">
        <property type="term" value="P:negative regulation of endocytic recycling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007264">
        <property type="term" value="P:small GTPase-mediated signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd04404">
        <property type="entry name" value="RhoGAP-p50rhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.555.10:FF:000024">
        <property type="entry name" value="Rho GTPase-activating protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.525.10">
        <property type="entry name" value="CRAL-TRIO lipid binding domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.555.10">
        <property type="entry name" value="Rho GTPase activation protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049592">
        <property type="entry name" value="ARHGAP1_RhoGAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001251">
        <property type="entry name" value="CRAL-TRIO_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036865">
        <property type="entry name" value="CRAL-TRIO_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000198">
        <property type="entry name" value="RhoGAP_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45808">
        <property type="entry name" value="RHO GTPASE-ACTIVATING PROTEIN 68F"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45808:SF4">
        <property type="entry name" value="RHO GTPASE-ACTIVATING PROTEIN 8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13716">
        <property type="entry name" value="CRAL_TRIO_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00620">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00324">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52087">
        <property type="entry name" value="CRAL/TRIO domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50238">
        <property type="entry name" value="RHOGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Rho-GAP" evidence="3">
        <location>
            <begin position="199"/>
            <end position="385"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="166"/>
            <end position="190"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="166"/>
            <end position="176"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="2">
        <location>
            <begin position="178"/>
            <end position="190"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS50238"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030200.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="396" mass="45721" checksum="54235C674496EEF3" modified="2019-05-08" version="1">MSEIDFEELAEIELQKDEAEYAIMEKRKTSFTPAACEDTELSINHPYYDVARHGIVQLAGDDNSGRRVITFSCCRMPPSHQLNHSRLLEYLRYTLDQYVENDYTLVYFHYGLNSRNKPSLNWLQSAYKEFDRNHKFGKKITYLNYLSDLREHLKYDQLNIPEDVIRHDQNIRDKQKGRPPPLAKLPPPRPPLPTQQFGVSLQYLKDKNKGQLIPPVMKQTVSYLKSKGLRVEGLFRRSASIQTIKEIHKLYNQGKPVNFDDYGDIHIPAVILKTFLRELPQPLLTFESYDQILGITSVESSLRVTSCKQVIQGLPEHNYVVLKYLICFLHMVSQESIYNKMTGSNLACVFGLNLIWPPKGASSLRALNPLNLFTELLIDFYSKIFSSRTVSGEILP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ15</accession>
    <name>A0A452IQ15_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Glycosyltransferase 2-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IQ15"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034073.1">
        <property type="protein sequence ID" value="ENSGAGP00000030004.1"/>
        <property type="gene ID" value="ENSGAGG00000021650.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008376">
        <property type="term" value="F:acetylgalactosaminyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019276">
        <property type="term" value="P:UDP-N-acetylgalactosamine metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006047">
        <property type="term" value="P:UDP-N-acetylglucosamine metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00761">
        <property type="entry name" value="Glyco_tranf_GTA_type"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.550.10">
        <property type="entry name" value="Spore Coat Polysaccharide Biosynthesis Protein SpsA, Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001173">
        <property type="entry name" value="Glyco_trans_2-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029044">
        <property type="entry name" value="Nucleotide-diphossugar_trans"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15046:SF2">
        <property type="entry name" value="BETA-1,4 N-ACETYLGALACTOSAMINYLTRANSFERASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15046">
        <property type="entry name" value="GLYCO_TRANS_2-LIKE DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00535">
        <property type="entry name" value="Glycos_transf_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53448">
        <property type="entry name" value="Nucleotide-diphospho-sugar transferases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="1">
        <location>
            <begin position="1"/>
            <end position="19"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019158978" description="Glycosyltransferase 2-like domain-containing protein" evidence="1">
        <location>
            <begin position="20"/>
            <end position="411"/>
        </location>
    </feature>
    <feature type="domain" description="Glycosyltransferase 2-like" evidence="2">
        <location>
            <begin position="171"/>
            <end position="282"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF00535"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030004.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="411" mass="46986" checksum="7DD28DC1F6A35ACA" modified="2019-05-08" version="1">RYFSTSILLWALTLSYIQPNFLTKEATFTCQGRRGSYSRPKPNVPLSYPIQGVEVMPLHTIMIPGIRVILRASLGTLNTLADVSDDVVRGRGRGKKELLILTSDVELLNFILEHVTYTSVVYQLSAVDMMSFKSRDHVVQFPVIIRQPSLPKLIDPGADRKISSLVTITTKTFLRYHKLRFLIKSIRQYYPDIKIIVADDSEVPEKIIEENIEHYIMPFGKGWFAGRNLAISQVTTKYYLWVDDDYLFTENTKIEKLLDVLERTNLDMVGGSVKGDTYSFQLLYEQSDDGYCLHLRYGSFHKLDGFPNCVVTSGVVNFYLAHTDKSRYVSYDPLLQRVAHSEFFVDGLGILLVGSCSDVSVGHQDHNPASDPNLAKIEDQYKTFRDNTDAQVQFKLALHYFKNRLKCYSTR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ16</accession>
    <name>A0A452IQ16_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Nck-associated protein 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="6">Cell projection</location>
            <location evidence="6">Lamellipodium membrane</location>
            <topology evidence="6">Single-pass membrane protein</topology>
            <orientation evidence="6">Cytoplasmic side</orientation>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="7">Belongs to the HEM-1/HEM-2 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IQ16"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034117.1">
        <property type="protein sequence ID" value="ENSGAGP00000030046.1"/>
        <property type="gene ID" value="ENSGAGG00000019065.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031258">
        <property type="term" value="C:lamellipodium membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031209">
        <property type="term" value="C:SCAR complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030031">
        <property type="term" value="P:cell projection assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030866">
        <property type="term" value="P:cortical actin cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048812">
        <property type="term" value="P:neuron projection morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019137">
        <property type="entry name" value="Nck-associated_protein-1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12093">
        <property type="entry name" value="NCK-ASSOCIATED PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12093:SF11">
        <property type="entry name" value="NCK-ASSOCIATED PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09735">
        <property type="entry name" value="Nckap1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="1" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="5" id="KW-0966">Cell projection</keyword>
    <keyword evidence="4" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="3" id="KW-1133">Transmembrane helix</keyword>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="460"/>
            <end position="485"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="471"/>
            <end position="485"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00037839"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00037947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00039689"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030046.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="919" mass="104690" checksum="B5E588D410429D86" modified="2019-05-08" version="1">MIVDYENPLKKMMEEFVPHSKSLSDALISLQMVYPRRNLSADQWRNAQLLSLISAPSTMLNPAQSDTMPCEYLSLDAMEKWIIFGFILCHGILNSDATALNLWKLALQSSSCLSLFRDEVFHIHKAAEDLFVNIRGYNKRINDIRECKEAAVSHAGSMHRERRKFLRSALKELATVLSDQPGLLGPKALFVFMALSFARDEIIWLLRHADNMPKKSADDFIDKHIAELIFYMEELRAHVRKYGPVMQRYYVQYLSGFDAVVLNELVQNLSVCPEDESIIMSSFVNTMTSLSVKQVEDGEVFDFRGMRLDWFRLQAYTSVSKASLSLADHRELGKMMNTIIFHTKMVDSLVEMLVETSDLSIFCFYSRAFEKMFQQCLELPSQSRYSIAFPLLCTHFMSCTHELCPEERHHIGDRSLSLCNMFLDEMAKQARNLITDICTEQCTLSDQLLPKHCAKTISQAVNKKSKKQTGKKGEPEREKPGVESMRKNRLVVTNLDKLHTALSELCFSINYVPNMVVWEHTFTPREYLTSHLEIRFTKSIVGMTMYNQATQEIAKPSELLTSVRAYMTVLQSIENYVQIDITRVFNNVLLQQTQHLDSHGEPTITSLYTNWYLETLLRQVSNGHIAYFPAMKAFVNLPTENELTFNAEEYSDISEMRALSELLGPYGMKFLSESLMWHISSQVAELKKLVVENVEVLTQMRTSFDKPDQMAALFKRLSSVDSVLKRMTIIGVILSFRSLAQEALRDVLSYHIPFLVSSIEDFKDHIPRETDMKVAMNVYELSSAAGLPCEIDPALVVALSSQKSENISPEEEYKIACLLMVFVAVSLPTLASNVMSQYSPAIEGHCNNIHCLAKAINQIAAALFTIHKGSIEDRLKEFLALASSSLLKIGQETDKTTTRNRESVYLLLDMVSSHAHCYF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2025-10-08" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IQ17</accession>
    <name>A0A452IQ17_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="10">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the prickle / espinas / testin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000034002.1">
        <property type="protein sequence ID" value="ENSGAGP00000029943.1"/>
        <property type="gene ID" value="ENSGAGG00000021608.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000034006.1">
        <property type="protein sequence ID" value="ENSGAGP00000029947.1"/>
        <property type="gene ID" value="ENSGAGG00000021608.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000034007.1">
        <property type="protein sequence ID" value="ENSGAGP00000029948.1"/>
        <property type="gene ID" value="ENSGAGG00000021608.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd09415">
        <property type="entry name" value="LIM1_Prickle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd09418">
        <property type="entry name" value="LIM2_Prickle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd09420">
        <property type="entry name" value="LIM3_Prickle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd09827">
        <property type="entry name" value="PET_Prickle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.110.10:FF:000022">
        <property type="entry name" value="prickle-like protein 2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.110.10:FF:000035">
        <property type="entry name" value="prickle-like protein 2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.110.10:FF:000005">
        <property type="entry name" value="Testin isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.110.10">
        <property type="entry name" value="Cysteine Rich Protein"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033725">
        <property type="entry name" value="LIM1_prickle"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033726">
        <property type="entry name" value="LIM2_prickle"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033727">
        <property type="entry name" value="LIM3_prickle"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010442">
        <property type="entry name" value="PET_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033723">
        <property type="entry name" value="PET_prickle"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047120">
        <property type="entry name" value="Pk/Esn/Tes"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001781">
        <property type="entry name" value="Znf_LIM"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24211">
        <property type="entry name" value="LIM DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24211:SF18">
        <property type="entry name" value="PRICKLE-LIKE PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00412">
        <property type="entry name" value="LIM"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06297">
        <property type="entry name" value="PET"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00132">
        <property type="entry name" value="LIM"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57716">
        <property type="entry name" value="Glucocorticoid receptor-like (DNA-binding domain)"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00478">
        <property type="entry name" value="LIM_DOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50023">
        <property type="entry name" value="LIM_DOMAIN_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51303">
        <property type="entry name" value="PET"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 6" id="KW-0440">LIM domain</keyword>
    <keyword evidence="2 6" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="4 6" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="PET" evidence="9">
        <location>
            <begin position="14"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="domain" description="LIM zinc-binding" evidence="8">
        <location>
            <begin position="124"/>
            <end position="188"/>
        </location>
    </feature>
    <feature type="domain" description="LIM zinc-binding" evidence="8">
        <location>
            <begin position="189"/>
            <end position="249"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="310"/>
            <end position="337"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="547"/>
            <end position="572"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="633"/>
            <end position="690"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="811"/>
            <end position="835"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="310"/>
            <end position="320"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="7">
        <location>
            <begin position="675"/>
            <end position="685"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00008268"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023038"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029943.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="835" mass="94982" checksum="3930D50E89EB4682" modified="2019-05-08" version="1">MPLEMEKTVTKLMFDFQRNSTSDDDSGCALEEYAWVPPGLKPEQVHQYYSCLPEDKVPYVNSPGEKLRIKQLLHQLPPHDNEVRYCNSLDEEEKRELKLFSNQRKRENLGRGNVRPFPVTMTGAICEQCGGQISGGDIAVFASRAGHGICWHPPCFICTVCNELLVDLIYFYQEGKIYCGRHHAECLKPRCAACDEIIFADECTEAEGRHWHMKHFCCFECETVLGGQRYIMKEGRPYCCNCFESLYAEYCDTCAXLKGIDQGQMTYDGQHWHATETCFCCAQCKKSLLGRPFLPKQGQIFCSRACSVGDDPNGSDSSDSAFQNARAKESRRSAKIGKNKGKVEENMLNQCSQLQVTSNRLSADVDPLSLQMDLLSLSSQTPSLNREHIWRSRDELYHYGNQMEQSQSQSPLQLLSQCNIRTSYNPGQVPGTQPDLWAKHFSNPKRSSSVAMKGHNGSFIQECREDYYSGRLRSQESYSDMSNQSFNETRGNIKVPKYDEGEEGGMSTKQCRTRHPINALKFTEDLTPTEQTPRGSMESLALSNATGLSADGGAKRQEHLSRFSMPDLSKDSGMNVSEKMSNMGTLNSSVQFRSAESVRSLLSVQQYQDMEPNLRDLANPANYREMPSHGRMHQSFEYDSGIPGNKIPGQDSARLPPMSERTRRRTNLRDDDRRYRPHRSRRSRRSRSDNALHLASEQCCRLKERPSLRAREDYDQFMHQRSCRETMEQGPCRDLYGQCPRTVSDLALQNHLGERWGPYFAEYDWCSTCSSSSESDNEGYFLGEPIPQPAHLRYVTSEELLHKYGSYGMPKSSTLGGRGQLHSRKRQKSKNCIIS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1F1</accession>
    <name>A0A452J1F1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Dynactin subunit 4</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="14">Subunit of dynactin, a multiprotein complex part of a tripartite complex with dynein and a adapter, such as BICDL1, BICD2 or HOOK3. The dynactin complex is built around ACTR1A/ACTB filament and consists of an actin-related filament composed of a shoulder domain, a pointed end and a barbed end. Its length is defined by its flexible shoulder domain. The soulder is composed of 2 DCTN1 subunits, 4 DCTN2 and 2 DCTN3. The 4 DCNT2 (via N-terminus) bind the ACTR1A filament and act as molecular rulers to determine the length. The pointed end is important for binding dynein-dynactin cargo adapters. Consists of 4 subunits: ACTR10, DCNT4, DCTN5 and DCTN6. The barbed end is composed of a CAPZA1:CAPZB heterodimers, which binds ACTR1A/ACTB filament and dynactin and stabilizes dynactin. Interacts with ATP7B, but not ATP7A, in a copper-dependent manner. Interacts with ANK2; this interaction is required for localization at costameres. Interacts with N4BP2L1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Cell cortex</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Microtubule organizing center</location>
            <location evidence="1">Centrosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
            <location evidence="2">Stress fiber</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cytoplasm</location>
            <location evidence="4">Myofibril</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="12">Belongs to the dynactin subunit 4 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1F1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038735.1">
        <property type="protein sequence ID" value="ENSGAGP00000034208.1"/>
        <property type="gene ID" value="ENSGAGG00000024291.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005938">
        <property type="term" value="C:cell cortex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005869">
        <property type="term" value="C:dynactin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030016">
        <property type="term" value="C:myofibril"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001725">
        <property type="term" value="C:stress fiber"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008603">
        <property type="entry name" value="DCTN4"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13034">
        <property type="entry name" value="DYNACTIN P62 SUBUNIT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13034:SF2">
        <property type="entry name" value="DYNACTIN SUBUNIT 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05502">
        <property type="entry name" value="Dynactin_p62"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0007">Acetylation</keyword>
    <keyword evidence="10" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="6" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0832">Ubl conjugation</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004529"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004544"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004657"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00034776"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00034864"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00093507"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034208.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="516" mass="58517" checksum="CDA63C6B73B297AA" modified="2019-05-08" version="1">MEKAHSSPYRPAWELPALNAHHVPLPGRKWLGDHVGAARPDHVRGRGGGCWAPAPKMASLLQSERVSYLVRGEKEIRAPLSQLYFCRYCSELRSLECVSHEVDSHYCPSCLENMPSAEAKLKKNRCANCFDCPCCMHTLSTRATSIPAPLPDDPAKTTMKKAYYLACGFCRWTSRDVGMADKSVASGGWQEPENPHTQRINKLVEYYQQLAQKEKIERDRKKLVRRRNYMPLAFSDKYGLGTRLQRQRPGMSISALSGLSLKEGEDLKEIKIEPADAVEEVEPLPEDYYTRPINLTEVTTLHQRLLQPDFQPICASQLYPRHKHLLIKRSLRCRKCEHNLSKPEFNPTSIKFKIQLVAFNYIPEVRIMSIPNLRYMKESQVLLTLTNPVENLTHVTLLECEEGDPDDINSTAKVVVPPKELILAGKDAAAEYDELAEPQDFQDDPDIVAFRKANKVGVFIKVTPQKEEGEVTVSFKMKHEFKNLAAPIRPSEEGDQSSEVIWLTHHVELSLGPLLP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1F2</accession>
    <name>A0A452J1F2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Ral GTPase-activating protein subunit beta</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="6">p170</fullName>
        </alternativeName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="3">Non-catalytic subunit of the heterodimeric RalGAP1 and RalGAP2 complexes which act as GTPase activators for the Ras-like small GTPases RALA and RALB.</text>
    </comment>
    <comment type="subunit">
        <text evidence="4">Component of the heterodimeric RalGAP1 complex with RALGAPA1 and of the heterodimeric RalGAP2 complex with RALGAPA2. Heterodimerization is required for activity.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038654.1">
        <property type="protein sequence ID" value="ENSGAGP00000034137.1"/>
        <property type="gene ID" value="ENSGAGG00000024251.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051056">
        <property type="term" value="P:regulation of small GTPase mediated signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.11210:FF:000005">
        <property type="entry name" value="Ral GTPase-activating protein, beta subunit (non-catalytic)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.11210">
        <property type="entry name" value="Rap/Ran-GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039930">
        <property type="entry name" value="RALGAPB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035974">
        <property type="entry name" value="Rap/Ran-GAP_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000331">
        <property type="entry name" value="Rap/Ran_GAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046859">
        <property type="entry name" value="RGPA/RALGAPB_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21344">
        <property type="entry name" value="RAL GTPASE-ACTIVATING PROTEIN SUBUNIT BETA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21344:SF1">
        <property type="entry name" value="RAL GTPASE-ACTIVATING PROTEIN SUBUNIT BETA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20412">
        <property type="entry name" value="RALGAPB_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF111347">
        <property type="entry name" value="Rap/Ran-GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50085">
        <property type="entry name" value="RAPGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Rap-GAP" evidence="8">
        <location>
            <begin position="1164"/>
            <end position="1408"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="355"/>
            <end position="386"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="399"/>
            <end position="438"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="721"/>
            <end position="753"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1089"/>
            <end position="1115"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1329"/>
            <end position="1351"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="369"/>
            <end position="381"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="399"/>
            <end position="411"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="412"/>
            <end position="428"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="734"/>
            <end position="750"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="1329"/>
            <end position="1339"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00053327"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00063020"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00072207"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00081608"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50085"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034137.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1510" mass="168369" checksum="9779A1CF609CF806" modified="2019-05-08" version="1">MYSEWRSLHLVVQNDQGHTSVLHSYPDNVGREVANAVVRPLGQALNTPSAAGSESLLKTDKEVKWTMEVVCYGLTLPLEGDTVKYCVDVYTDWIMALVLPKDSIPLPVIKEPNLYVQSILKHLQNLFVPRQDPGSSQNRLCLQVLKAVQKLARESTIMARETWEVLLLFLLQINDTLLAAPTVQGGIAENLAEKLIGVLFEVWLLACTRCFPTPPYWKTAKEMVANWRHHPAVVEQWSKVICALTSRLLRFIYGPSFPPFKIPDEDASLIPPEMDNECIAQTWFRFLHMLSNPVDLSNPAIISSTPKFQEQFLNVSGMTQELNQYPCLKHLPHIFFRAMRGISCLVDAFLGISRPRSDSAPPTPVNRLSMPQNTAINTTPPHNRRHRAVTVNKATMKTSTVSTVHASKVQHQPSSSTSPLSSPNQTSSEPRPLPAPHRPKVNSILNLFGSWLFDAGFVHCKLHNGINRDSSMTASFIQILLSYKSSIATQASMEFRRKGSQMSTDTMTSNSTFDSSDFPDNYEAGRAEACGTLCRIFCSKKTGEDILPAYLSRFYMLLIQGLQIADFICHPVLASIILNSPPLFCCDLKGIDVVVPYFISALETILPDRELSKFKTYVNPTDLRRSSINILLSLLPLPHHFGTIKSEVVLEGKFSNDDGSSSYDKPITFLSLKLRLVNILIGALQTETDPNNTQMILGAMLNIVQDSALLEAIGCQMETNGGENNLLKNHRRTNSSISTASGGSTEPTTPDSERPAQALLRDYALNTDTAAGLLIRSIHLVTQRLNSQWRQDMSISLAALELLSGLAKVKVIVDSADRKRAVSSVCSYIVYQCSRPAPLHSRDLHSMIVAAFQCLCVWLTEHPDMMDEKDCLKEVLEIVELGISGSKSKNSEQEVKYKGDKEPNPASMRVKDAAEATLTCIMQLLGAFPSPSGPASPCSLVNETTLIKYSRLPTINKHSFRYFVLDNSVILAMLEQPLGNEQNDSFPSVTVLIRGVSGRLAWAQQLCLLPRGAKANQKMFLPEPRPAPKNDVGLKYSVKHRPFPEEVDKIPFVKADLSIPDLHEIVNEELEERHEKLRNGMAQQITYETSVDQQSEEEWRKKSFPDPVTECKPPPPAQEFQTARLFLSHFGFLSLEALKETANSRLPPHLIALDSALPGFFDDLGYLDLLPCRPFDTVFIFYMKAGQKTSQEILKNVESSRNVQPHFLEFLLSLGWAVDVGKHPGWTGHVSTSWSINSCSDEEGSEQDEIVSSEDTGASIFNGQKKVLYYADALTEIAFVVPSPVESLTDSLESNVSDQESDSNMDLLPGILKQPSLTLELFPNHTDNLNSLQRLSPTSRSKKLPQGRTIPPMGPETKVSVVWVERYDDIENFPLSDLVSETSTGVETTSNSSTSLRSTTPEREVPVVFIHPLNTGLFRIKVQGATGKFNMVIPLVDGMIVSRRALGFLVRQTIINICRRKRLESDSYNPPHVRRKQKIADIVNKYRNKQLEPEFYTSLFQEVGLKNCNP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1F6</accession>
    <name>A0A452J1F6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Taste receptor type 1 member 2</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="17">Sweet taste receptor T1R2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">Putative taste receptor. TAS1R2/TAS1R3 recognizes diverse natural and synthetic sweeteners.</text>
    </comment>
    <comment type="subunit">
        <text evidence="15">Forms heterodimers with TAS1R3.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="14">Belongs to the G-protein coupled receptor 3 family. TAS1R subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1F6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038740.1">
        <property type="protein sequence ID" value="ENSGAGP00000034213.1"/>
        <property type="gene ID" value="ENSGAGG00000024281.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903767">
        <property type="term" value="C:sweet taste receptor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004930">
        <property type="term" value="F:G protein-coupled receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033041">
        <property type="term" value="F:sweet taste receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.50.30:FF:000004">
        <property type="entry name" value="Taste receptor type 1 member 3-like protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.2300">
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.50.30">
        <property type="entry name" value="GPCR, family 3, nine cysteines domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001828">
        <property type="entry name" value="ANF_lig-bd_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000337">
        <property type="entry name" value="GPCR_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011500">
        <property type="entry name" value="GPCR_3_9-Cys_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038550">
        <property type="entry name" value="GPCR_3_9-Cys_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017978">
        <property type="entry name" value="GPCR_3_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000068">
        <property type="entry name" value="GPCR_3_Ca_sens_rcpt-rel"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017979">
        <property type="entry name" value="GPCR_3_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028082">
        <property type="entry name" value="Peripla_BP_I"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24061">
        <property type="entry name" value="CALCIUM-SENSING RECEPTOR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24061:SF517">
        <property type="entry name" value="TASTE RECEPTOR TYPE 1 MEMBER 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00003">
        <property type="entry name" value="7tm_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01094">
        <property type="entry name" value="ANF_receptor"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07562">
        <property type="entry name" value="NCD3G"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00248">
        <property type="entry name" value="GPCRMGR"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53822">
        <property type="entry name" value="Periplasmic binding protein-like I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00980">
        <property type="entry name" value="G_PROTEIN_RECEP_F3_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50259">
        <property type="entry name" value="G_PROTEIN_RECEP_F3_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="8" id="KW-0297">G-protein coupled receptor</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9 18" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0675">Receptor</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0716">Sensory transduction</keyword>
    <keyword evidence="6 19" id="KW-0732">Signal</keyword>
    <keyword evidence="3" id="KW-0919">Taste</keyword>
    <keyword evidence="12" id="KW-0807">Transducer</keyword>
    <keyword evidence="5 18" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 18" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="19">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019074938" description="Taste receptor type 1 member 2" evidence="19">
        <location>
            <begin position="23"/>
            <end position="791"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="18">
        <location>
            <begin position="516"/>
            <end position="543"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="18">
        <location>
            <begin position="555"/>
            <end position="576"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="18">
        <location>
            <begin position="588"/>
            <end position="611"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="18">
        <location>
            <begin position="631"/>
            <end position="651"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="18">
        <location>
            <begin position="680"/>
            <end position="701"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="18">
        <location>
            <begin position="713"/>
            <end position="732"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="18">
        <location>
            <begin position="738"/>
            <end position="760"/>
        </location>
    </feature>
    <feature type="domain" description="G-protein coupled receptors family 3 profile" evidence="20">
        <location>
            <begin position="517"/>
            <end position="783"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022480"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022606"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023224"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00037659"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00038492"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00038699"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00040704"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00042616"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50259"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034213.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="791" mass="89714" checksum="9D31B49D2A5CBBCB" modified="2019-05-08" version="1">MRSTGGIAMCVSVLLTMAGSQTQYSDFHLAGDYLLGGLFSLHTETKSSPHLSYSAVPTCQAYKLKKASYSHLRAMRFAVEQINNSSTLLPGVSLGYEMVDVCYLTNFIHPILYFLANKHSVVQLHNNYTHYQPRVLAVIGPDSSQAVVIAAQVLSLFLVPQISYTATTNELSDRKRFPTAFQTISSMEQQIVAMQLLLKEFRWNWINILYSDDDYGRQNLKLLRARVVIVLSLELILPFFFQEAVRQNVTDLVWIATEAWATEPALYNITDLWRLGTILGVAVKEFSIPGLDDFRVSLAEGANEQASDQTTCNQECDQCLRAAQDAEHSMRAQGERTDFSVYSSVYIVAHALHRLLGCNTTGCHKRMVYPWQLVPEVKRVNFSLLNERINFRERGTIPNSFEIIQWKWDQPNNPFQKIASYSRNKKLLFTIQNISWHTPNNTVPISVCSQQCEYGQRRKPVTVSSCCFECIDCEEGTFLNNNDLYNCQPCPADQWSNARSQKCYPRLLKYLEWQEGIAVVLLLFSVLGLVATVVISVIFAIHLHTPVVKSAGGRMCFLMLASLIIGFINVPVYMGIPTEFKCMCRQIVYTLCFTVCISCLTVRSFQIISIFKMAARLPKAYSYWMKYNGQYIFITVILALKVTIVVTNVIIHPSIPVPLVLEDDPSTVELQCNKSFKLSLLMNNSLDTFLSILCFYFSYIGKELPKNYNEAKYITLCMTCYAVSEIAIFLAMSVYRGMLVTICDVIGVVINLLGISIGYFGPKCYLIFFHPERNTAAYFQSVIQSYTMRQD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1F8</accession>
    <name>A0A452J1F8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Calcitonin peptide-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the calcitonin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1F8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038742.1">
        <property type="protein sequence ID" value="ENSGAGP00000034215.1"/>
        <property type="gene ID" value="ENSGAGG00000024303.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031716">
        <property type="term" value="F:calcitonin receptor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005179">
        <property type="term" value="F:hormone activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007189">
        <property type="term" value="P:adenylate cyclase-activating G protein-coupled receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051480">
        <property type="term" value="P:regulation of cytosolic calcium ion concentration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021118">
        <property type="entry name" value="Calcitonin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021117">
        <property type="entry name" value="Calcitonin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021116">
        <property type="entry name" value="Calcitonin/adrenomedullin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018360">
        <property type="entry name" value="Calcitonin_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001693">
        <property type="entry name" value="Calcitonin_peptide-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10505:SF16">
        <property type="entry name" value="CALCITONIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10505">
        <property type="entry name" value="CALCITONIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00214">
        <property type="entry name" value="Calc_CGRP_IAPP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00270">
        <property type="entry name" value="CALCITONINA"/>
    </dbReference>
    <dbReference type="SMART" id="SM00113">
        <property type="entry name" value="CALCITONIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00258">
        <property type="entry name" value="CALCITONIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0027">Amidation</keyword>
    <keyword evidence="4" id="KW-0165">Cleavage on pair of basic residues</keyword>
    <keyword evidence="8 9" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5" id="KW-0372">Hormone</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0964">Secreted</keyword>
    <keyword evidence="6 10" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="10">
        <location>
            <begin position="1"/>
            <end position="25"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019317906" description="Calcitonin peptide-like domain-containing protein" evidence="10">
        <location>
            <begin position="26"/>
            <end position="138"/>
        </location>
    </feature>
    <feature type="domain" description="Calcitonin peptide-like" evidence="11">
        <location>
            <begin position="80"/>
            <end position="117"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="9">
        <location>
            <begin position="82"/>
            <end position="88"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009222"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022685"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022702"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022815"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PIRSR" id="PIRSR621116-50"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="SMART" id="SM00113"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034215.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="138" mass="15369" checksum="DFC54F50252B3072" modified="2019-05-08" version="1">MVMLKISPFLAVYALVMCQMDSSQATPLRPGLDSITDRVTLSDYEARRLLNALVKEFVQMRAEDVEQETEGNSLDKPISKRCASLSTCVLGKLSQELHKLQTYPRTDVGAGTPGKKRNVLNDLENERYANYGESLGNN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1F9</accession>
    <name>A0A452J1F9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">Anaphase-promoting complex subunit 4</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J1F9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038662.1">
        <property type="protein sequence ID" value="ENSGAGP00000034143.1"/>
        <property type="gene ID" value="ENSGAGG00000024170.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005680">
        <property type="term" value="C:anaphase-promoting complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034399">
        <property type="term" value="C:nuclear periphery"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031145">
        <property type="term" value="P:anaphase-promoting complex-dependent catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070979">
        <property type="term" value="P:protein K11-linked ubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024789">
        <property type="entry name" value="APC4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056358">
        <property type="entry name" value="APC4_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024790">
        <property type="entry name" value="APC4_long_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13260">
        <property type="entry name" value="ANAPHASE PROMOTING COMPLEX SUBUNIT 4 APC4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13260:SF0">
        <property type="entry name" value="ANAPHASE-PROMOTING COMPLEX SUBUNIT 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12896">
        <property type="entry name" value="ANAPC4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23405">
        <property type="entry name" value="WD40_APC4_C-half"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="2" id="KW-0132">Cell division</keyword>
    <keyword evidence="3" id="KW-0498">Mitosis</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="Anaphase-promoting complex subunit 4 long" evidence="6">
        <location>
            <begin position="123"/>
            <end position="321"/>
        </location>
    </feature>
    <feature type="domain" description="Anaphase-promoting complex subunit 4 C-terminal half WD40" evidence="7">
        <location>
            <begin position="465"/>
            <end position="624"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00016067"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022776"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="Pfam" id="PF12896"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF23405"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034143.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="654" mass="74362" checksum="03235FB907A47129" modified="2019-05-08" version="1">MHWMEVTEESSVLTSFYNAEDESNLLLPKLPALPKNYSTTAKIFSEEKSDEIMKLLGDVRLNALVLGGSSGFIEVYAYGMYKIATVTGVAGSCLGLCFSSDLKSLSVITEVQVSPDSAPEITYFQLDTSLLSVYLPEVTRMARKFTHISTLLQYIKLSLTCMCEAWEEILMQMDSRLTKFVQEKNTTTSVQDEFMQLLLWGKASLELQALLMNQLTVKGLKKLGQSIESSYSSIQKLVISHLQSGSEALLYHLSELKGMASWKQKYESLGLDAAGIEGAITAVGSFILKANELLQVIDSSMKNFKAFFRWLYVAMLRMSEDHVLPELNKMTQKDITFVADFLTEHFNEAPDLYNRKGKYFNVERVGQYLKDEDDDLVSPPNTEGNQWFNFLQNSTHLKESPLLFPYYPQKSLHFVKRRMEGIIDQCLQKPADVIGKSVHQAVCMCLYKGSKSEDSTPRLFKLPFMWNDKSSNLHYVLFTMLESSLSKIYILRRHTDVSRSANNGLLSVDFGNFLNNSINESSDTGFYSCVDAHFYDDETITVVLKENAEQEGKERILAQLPLSSVYVDEDQEREFSLNSNKRLDEQSDDIPTRTVFLESQWRVLENMKAQYVAVNGIRKVSCVILRKKEGDNLHLLSRVEKIIIIQLKYLLASV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1G1</accession>
    <name>A0A452J1G1_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="20">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 16">Membrane</location>
            <topology evidence="1 16">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 16">Belongs to the integrin alpha chain family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1G1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038758.1">
        <property type="protein sequence ID" value="ENSGAGP00000034231.1"/>
        <property type="gene ID" value="ENSGAGG00000024233.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009897">
        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008305">
        <property type="term" value="C:integrin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033627">
        <property type="term" value="P:cell adhesion mediated by integrin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098609">
        <property type="term" value="P:cell-cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007160">
        <property type="term" value="P:cell-matrix adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007229">
        <property type="term" value="P:integrin-mediated signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.130:FF:000006">
        <property type="entry name" value="Integrin, alpha 9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.930">
        <property type="entry name" value="Bicelle-embedded integrin alpha(iib) transmembrane segment"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.130">
        <property type="entry name" value="Integrin alpha, N-terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1460">
        <property type="entry name" value="Integrin domains. Chain A, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1510">
        <property type="entry name" value="ntegrin, alpha v. Chain A, domain 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013517">
        <property type="entry name" value="FG-GAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013519">
        <property type="entry name" value="Int_alpha_beta-p"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000413">
        <property type="entry name" value="Integrin_alpha"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013649">
        <property type="entry name" value="Integrin_alpha_Ig-like_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048285">
        <property type="entry name" value="Integrin_alpha_Ig-like_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028994">
        <property type="entry name" value="Integrin_alpha_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032695">
        <property type="entry name" value="Integrin_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23220">
        <property type="entry name" value="INTEGRIN ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23220:SF78">
        <property type="entry name" value="INTEGRIN ALPHA-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01839">
        <property type="entry name" value="FG-GAP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08441">
        <property type="entry name" value="Integrin_A_Ig_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20805">
        <property type="entry name" value="Integrin_A_Ig_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01185">
        <property type="entry name" value="INTEGRINA"/>
    </dbReference>
    <dbReference type="SMART" id="SM00191">
        <property type="entry name" value="Int_alpha"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69318">
        <property type="entry name" value="Integrin alpha N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69179">
        <property type="entry name" value="Integrin domains"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51470">
        <property type="entry name" value="FG_GAP"/>
        <property type="match status" value="5"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0106">Calcium</keyword>
    <keyword evidence="8 16" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="14" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10 16" id="KW-0401">Integrin</keyword>
    <keyword evidence="11 16" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13 16" id="KW-0675">Receptor</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5 17" id="KW-0732">Signal</keyword>
    <keyword evidence="3 16" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9 16" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="17">
        <location>
            <begin position="1"/>
            <end position="37"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019379327" evidence="17">
        <location>
            <begin position="38"/>
            <end position="911"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="846"/>
            <end position="866"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="43"/>
            <end position="104"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="190"/>
            <end position="241"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="296"/>
            <end position="355"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="359"/>
            <end position="416"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="420"/>
            <end position="482"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha first immunoglubulin-like" evidence="18">
        <location>
            <begin position="467"/>
            <end position="572"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha second immunoglobulin-like" evidence="19">
        <location>
            <begin position="591"/>
            <end position="711"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023037"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00803"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="RuleBase" id="RU003762"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF08441"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="Pfam" id="PF20805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034231.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="911" mass="101112" checksum="E182A6CD844E7D1C" modified="2019-05-08" version="1">MRSRARLPGGLASGGAPVWAALLLLLLLGRCMRAAHAYNLDTQRPVVFKGLSGTLFGYSVVLHSHGPSRWLVVGAPKANWLANSSVVNPGAIFRCRIGSNPSGTCEQLQMGNPAGDYCGKTCFEERDNQWLGVSLSRQPRENGTIVACGHRWKNIYYIKNEHKLPNGVCYEIPPDFRTQLSKRISPCYKDYVRKFGENHGSCQAGMSSFYFKDLIIMGAPGSYYWTGSVFVYNTTANTFQAYTDSRNQVKFGSYLGYAVGAGHFLTPASTEVIGGAPQQEQTGKAYIFSIGTRALTILSELKGKKLGSYFGAAVCAVDLNTDGLSDLLVGAPMQSKVREEGRVYVYINSGSGAKMLELEIELAGSDSYAARFGESIANLGDIDNDGFEDVAIGAPQENNLEGAIYIYNGRKDGISSSFSQRIQGHRISNTLSMFGQSISSGIDADNNGYPDVAVGAFLDDSAVLLRTRPVVIVEAFLKHPNAVNRTNLECVENGQPAVCVNLMLCFTYKGRDVPGYVVLLYDLSVDVSRKTNTPARFYFSSNGTSDMVSGSMEIYNSNVTCKSHLAFLKENIGIHIFGMIRAKDRWKTLPHEKKTYLSVGSMKTLLLNISLSNAGDDAYETALHIQLPKGLYFVRVLELEEKQIHCEISDKEVHAVRLDCSVGYLYVDHLSKMDFSFLLDASSLTRAEDDLTITVNATCEENSPINCMKEKVNFTFHVISAGPSMAPAIGLEIMVPNSFAPRDEKLFNILDIQVCLFYYINILHTALYVTKASTHQGLIHLLQYLCNLAFFFFPYPQQDEASSLKFEISATASPEKNPKVIELHKERQIAHVFLEGLHNQKPKHHVTMLIIGTGSVLGIFLFLLIVLLLWKVGFFFLYQYFSIIFIDSFKNRNNMLYCDLFLLKKNKSWYD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1G2</accession>
    <name>A0A452J1G2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Fork-head domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 6">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1G2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038754.1">
        <property type="protein sequence ID" value="ENSGAGP00000034227.1"/>
        <property type="gene ID" value="ENSGAGG00000024307.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009653">
        <property type="term" value="P:anatomical structure morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048513">
        <property type="term" value="P:animal organ development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048731">
        <property type="term" value="P:system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd20019">
        <property type="entry name" value="FH_FOXE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.10:FF:000201">
        <property type="entry name" value="Forkhead box E1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.10">
        <property type="entry name" value="Winged helix-like DNA-binding domain superfamily/Winged helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001766">
        <property type="entry name" value="Fork_head_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050211">
        <property type="entry name" value="FOX_domain-containing"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018122">
        <property type="entry name" value="TF_fork_head_CS_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR030456">
        <property type="entry name" value="TF_fork_head_CS_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036388">
        <property type="entry name" value="WH-like_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036390">
        <property type="entry name" value="WH_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11829">
        <property type="entry name" value="FORKHEAD BOX PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11829:SF156">
        <property type="entry name" value="FORKHEAD BOX PROTEIN E3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00250">
        <property type="entry name" value="Forkhead"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00053">
        <property type="entry name" value="FORKHEAD"/>
    </dbReference>
    <dbReference type="SMART" id="SM00339">
        <property type="entry name" value="FH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46785">
        <property type="entry name" value="Winged helix' DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00657">
        <property type="entry name" value="FORK_HEAD_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00658">
        <property type="entry name" value="FORK_HEAD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50039">
        <property type="entry name" value="FORK_HEAD_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3 6" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="5 6" id="KW-0539">Nucleus</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0804">Transcription</keyword>
    <keyword evidence="2" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="Fork-head" evidence="7">
        <location>
            <begin position="68"/>
            <end position="162"/>
        </location>
    </feature>
    <feature type="DNA-binding region" description="Fork-head" evidence="6">
        <location>
            <begin position="68"/>
            <end position="162"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50039"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034227.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="278" mass="30252" checksum="A1629CFCBCD56DE1" modified="2019-05-08" version="1">MFDNGSFLRRRKRFKRADLATYPAYAAFGPAAPYAPAGYSPPLPAGAAYPRGFTIDSLMGGRPVPRGKPPYSYIALIAMAIAHAAERRLTLGGIYRFITERFPFYRDNPKKWQNSIRHNLTLNDCFVKVPREPGHPGKGSYWALDPAAEDMFDNGSFLRRRKRFKRADLATYPAYAAFGPAAPYAPAGYSPPLPAGAAYPRGFTIDSLMGGAELGRAYPYSASPGAPLYGAAGRLGLPPVRSCAEHPEPLLALAGPQFGPNPAYMRQPGFASGLERYL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1G4</accession>
    <name>A0A452J1G4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Integrin alpha second immunoglobulin-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 16">Membrane</location>
            <topology evidence="1 16">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 16">Belongs to the integrin alpha chain family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1G4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038750.1">
        <property type="protein sequence ID" value="ENSGAGP00000034223.1"/>
        <property type="gene ID" value="ENSGAGG00000024233.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
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        <property type="term" value="C:external side of plasma membrane"/>
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        <property type="project" value="TreeGrafter"/>
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        <property type="project" value="InterPro"/>
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        <property type="term" value="F:integrin binding"/>
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        <property type="project" value="TreeGrafter"/>
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        <property type="term" value="F:metal ion binding"/>
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        <property type="project" value="UniProtKB-KW"/>
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        <property type="term" value="P:cell adhesion mediated by integrin"/>
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        <property type="project" value="TreeGrafter"/>
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        <property type="term" value="P:cell-cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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        <property type="term" value="P:integrin-mediated signaling pathway"/>
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        <property type="project" value="UniProtKB-KW"/>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Bicelle-embedded integrin alpha(iib) transmembrane segment"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Integrin alpha, N-terminal"/>
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    </dbReference>
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        <property type="entry name" value="Integrin domains. Chain A, domain 2"/>
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    </dbReference>
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        <property type="entry name" value="ntegrin, alpha v. Chain A, domain 3"/>
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        <property type="entry name" value="FG-GAP"/>
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        <property type="entry name" value="Int_alpha_beta-p"/>
    </dbReference>
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        <property type="entry name" value="Integrin_alpha"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048285">
        <property type="entry name" value="Integrin_alpha_Ig-like_2"/>
    </dbReference>
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        <property type="entry name" value="Integrin_alpha_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032695">
        <property type="entry name" value="Integrin_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23220">
        <property type="entry name" value="INTEGRIN ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23220:SF78">
        <property type="entry name" value="INTEGRIN ALPHA-4"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="FG-GAP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20805">
        <property type="entry name" value="Integrin_A_Ig_2"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="INTEGRINA"/>
    </dbReference>
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        <property type="entry name" value="Int_alpha"/>
        <property type="match status" value="5"/>
    </dbReference>
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        <property type="entry name" value="Integrin alpha N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Integrin domains"/>
        <property type="match status" value="3"/>
    </dbReference>
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        <property type="entry name" value="FG_GAP"/>
        <property type="match status" value="5"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0106">Calcium</keyword>
    <keyword evidence="8 16" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="12" id="KW-1015">Disulfide bond</keyword>
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    <keyword evidence="11 16" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13 16" id="KW-0675">Receptor</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
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    <keyword evidence="5 17" id="KW-0732">Signal</keyword>
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    <keyword evidence="9 16" id="KW-1133">Transmembrane helix</keyword>
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        <location>
            <begin position="1"/>
            <end position="37"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019513852" description="Integrin alpha second immunoglobulin-like domain-containing protein" evidence="17">
        <location>
            <begin position="38"/>
            <end position="941"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="878"/>
            <end position="900"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="43"/>
            <end position="104"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="190"/>
            <end position="241"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="296"/>
            <end position="355"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="359"/>
            <end position="416"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="420"/>
            <end position="482"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha second immunoglobulin-like" evidence="18">
        <location>
            <begin position="560"/>
            <end position="704"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
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            <dbReference type="ARBA" id="ARBA00008054"/>
        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
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    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
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            <dbReference type="Pfam" id="PF20805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034223.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="941" mass="104597" checksum="6F91A7B6DD50041F" modified="2019-05-08" version="1">MRSRARLPGGLASGGAPVWAALLLLLLLGRCMRAAHAYNLDTQRPVVFKGLSGTLFGYSVVLHSHGPSRWLVVGAPKANWLANSSVVNPGAIFRCRIGSNPSGTCEQLQMGNPAGDYCGKTCFEERDNQWLGVSLSRQPRENGTIVACGHRWKNIYYIKNEHKLPNGVCYEIPPDFRTQLSKRISPCYKDYVRKFGENHGSCQAGMSSFYFKDLIIMGAPGSYYWTGSVFVYNTTANTFQAYTDSRNQVKFGSYLGYAVGAGHFLTPASTEVIGGAPQQEQTGKAYIFSIGTRALTILSELKGKKLGSYFGAAVCAVDLNTDGLSDLLVGAPMQSKVREEGRVYVYINSGSGAKMLELEIELAGSDSYAARFGESIANLGDIDNDGFEDVAIGAPQENNLEGAIYIYNGRKDGISSSFSQRIQGHRISNTLSMFGQSISSGIDADNNGYPDVAVGAFLDDSAVLLRTRPVVIVEAFLKHPNAVNRTNLECVENGQPAVCVNLMLCFTYKGRDVPGYVVLLYDLSVDVSRKTNTPARFYFSSNGTSDMVSGSMEIYNSNVTCKSHLAFLKENIERAKDRWKTLPHEKKTYLSVGSMKTLLLNISLSNAGDDAYETALHIQLPKGLYFVRVLELEEKQIHCEISDKEVHAVRLDCSVGYLYVDHLSKMDFSFLLDASSLTRAEDDLTITVNATCEENSPINCMKEKVNFTFHVISAGPSMAPAIGLEIMVPNSFAPRDEKLFNILDIQTADGECFYNNYTRDCTLPEKNGHMNILEDLVIFFSKPTKRRLYCMKEDHLCLRILCKFGNMENGKEAIVRMQLEATPSVLEMDEASSLKFEISATASPEKNPKVIELHKERQIAHVFLEGLHNQKPKHHVTMLIIGTGSVLGIFLFLLIVLLLWKVGFFFLYQYFSIIFIDSFKNRNNMLYCDLFLLKKNKSWYD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1G5</accession>
    <name>A0A452J1G5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="19">Glutamate receptor</fullName>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
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                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="19">Receptor for glutamate that functions as a ligand-gated ion channel in the central nervous system and plays an important role in excitatory synaptic transmission. L-glutamate acts as an excitatory neurotransmitter at many synapses in the central nervous system.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>Ca(2+)(in) = Ca(2+)(out)</text>
            <dbReference type="Rhea" id="RHEA:29671"/>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="14 19">Postsynaptic cell membrane</location>
            <topology evidence="14 19">Multi-pass membrane protein</topology>
        </subcellularLocation>
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    <comment type="similarity">
        <text evidence="19">Belongs to the glutamate-gated ion channel (TC 1.A.10.1) family.</text>
    </comment>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015276">
        <property type="term" value="F:ligand-gated monoatomic ion channel activity"/>
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        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0038023">
        <property type="term" value="F:signaling receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd06382">
        <property type="entry name" value="PBP1_iGluR_Kainate"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.2300:FF:000010">
        <property type="entry name" value="Glutamate ionotropic receptor kainate type subunit 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.190.10:FF:000240">
        <property type="entry name" value="Glutamate receptor ionotropic, kainate 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.70:FF:000010">
        <property type="entry name" value="Putative glutamate receptor ionotropic kainate 1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.190.10">
        <property type="entry name" value="Periplasmic binding protein-like II"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="ANF_lig-bd_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019594">
        <property type="entry name" value="Glu/Gly-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001508">
        <property type="entry name" value="Iono_Glu_rcpt_met"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015683">
        <property type="entry name" value="Ionotropic_Glu_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001320">
        <property type="entry name" value="Iontro_rcpt_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028082">
        <property type="entry name" value="Peripla_BP_I"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18966">
        <property type="entry name" value="IONOTROPIC GLUTAMATE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01094">
        <property type="entry name" value="ANF_receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00060">
        <property type="entry name" value="Lig_chan"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10613">
        <property type="entry name" value="Lig_chan-Glu_bd"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="NMDARECEPTOR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00079">
        <property type="entry name" value="PBPe"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53822">
        <property type="entry name" value="Periplasmic binding protein-like I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53850">
        <property type="entry name" value="Periplasmic binding protein-like II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 19" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="8 18" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13 19" id="KW-0407">Ion channel</keyword>
    <keyword evidence="6 19" id="KW-0406">Ion transport</keyword>
    <keyword evidence="12 19" id="KW-1071">Ligand-gated ion channel</keyword>
    <keyword evidence="7 19" id="KW-0472">Membrane</keyword>
    <keyword evidence="11 19" id="KW-0628">Postsynaptic cell membrane</keyword>
    <keyword evidence="9 19" id="KW-0675">Receptor</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="19" id="KW-0732">Signal</keyword>
    <keyword evidence="5 19" id="KW-0770">Synapse</keyword>
    <keyword evidence="3 19" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 19" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="1 19" id="KW-0813">Transport</keyword>
    <feature type="signal peptide" evidence="19">
        <location>
            <begin position="1"/>
            <end position="19"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5041014849" description="Glutamate receptor" evidence="19">
        <location>
            <begin position="20"/>
            <end position="842"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="493"/>
            <end position="512"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="569"/>
            <end position="591"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="750"/>
            <end position="774"/>
        </location>
    </feature>
    <feature type="domain" description="Ionotropic glutamate receptor C-terminal" evidence="20">
        <location>
            <begin position="404"/>
            <end position="731"/>
        </location>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="446"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="448"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="453"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="619"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="620"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="668"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="site" description="Crucial to convey clamshell closure to channel opening" evidence="17">
        <location>
            <position position="598"/>
        </location>
    </feature>
    <feature type="site" description="Interaction with the cone snail toxin Con-ikot-ikot" evidence="17">
        <location>
            <position position="625"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="18">
        <location>
            <begin position="68"/>
            <end position="319"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="18">
        <location>
            <begin position="680"/>
            <end position="734"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023257"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00034104"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00036634"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="RuleBase" id="RU367118"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="SMART" id="SM00079"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034148.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="842" mass="95138" checksum="7ED27451CF78BD4B" modified="2019-05-08" version="1">MLLFYLLASTGGIFETVENEPVNVEELAFKFAVTNINRNKTLMPNTTLTYDIQRINLFDSFEASRRACDQLALGVAALFGPSHSSSVSAVQSICNALEVPHIQTRWKHPTVDNKDTFYINLYPDYAAISRAILDLVLYYNWKIVTVVYEDSTGLIRLQELIKAPSRYNIKIKIRQLPSGNKDARPLLKEMKKGKEFYVIFDCSHETAAEILKQILSMGMMTEYYHYFFTTLDLFALDLEPYRYSGVNMTGFRLLNIDNPQVTSVIEKWSMERLQAPPKPETGLLDGMMTTEAALMYDAVYMVAVASQRASQMTVSSLQCHRHKPWRFGPRFMNLIKEARWDGLTGHITFNKTDGLRKDFDLDIISLKEEGTEKIGVWNSNSGLNMTDSNKDRSTNITDSLANRTLIVTTILVSPAVEGIKICLLNAKKMRIFSNLCSQKADLAVAPLTITYVREKVIDFSKPFMTLGISILYRKPNGTNPGVFSFLNPLSPDIWMYVLLACLGVSCVLFVIARFTPYEWYNPHPCNPDSDVVENNFTLLNSFWFGVGALMQQGSELMPKALSTRIVGGIWWFFTLIIISSYTANLAAFLTVERMESPIDSADDLAKQTKIEYGAVRDGSTMTFFKKSKISTYEKMWAFMSSRQQTALVKNNDEGIQRVLTTDYALLMESTSIEYVTQRNCNLTQIGGLIDSKGYGVGTPIGSPYRDKITIAILQLQEEGKLHMMKEKWWRGNGCPEEDSKEASALGVENIGGIFIVLAAGLVLSVFVAIGEFIYKSRKNSDIEQVSFFQGKCLSFNAIMEELGISLRHQKKLKKRSRTKGRSSFTSILTCHQRRTLRKETMA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1I9</accession>
    <name>A0A452J1I9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">Alpha-N-acetylgalactosaminide alpha-2,6-sialyltransferase 5</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="19">GD1 alpha synthase</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="20">GalNAc alpha-2,6-sialyltransferase V</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="18">ST6GalNAc V</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="21">Sialyltransferase 7E</fullName>
        </alternativeName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="16">Predominantly catalyzes the biosynthesis of ganglioside GD1alpha from GM1b in the brain, by transferring the sialyl group (N-acetyl-alpha-neuraminyl or NeuAc) from CMP-NeuAc to the GalNAc residue on the NeuAc-alpha-2,3-Gal-beta-1,3-GalNAc sequence of GM1b. GD1alpha is a critical molecule in the communication and interaction between neuronal cells and their supportive cells, particularly in brain tissues, and functions as an adhesion molecule in the process of metastasis. Also shows activity towards sialyl Lc4Cer (N-acetyl-alpha-neuraminosyl-(2-&gt;3)-beta-D-galactosyl-(1-&gt;3)-N-acetyl-beta-D-glucosaminyl-(1-&gt;3)-beta-D-galactosyl-(1-&gt;4)-beta-D-glucosyl-(1&lt;-&gt;1')-N-acyl-sphing-4-enine) generating disialyl Lc4Cer, which can lead to the synthesis of disialyl Lewis a (Le(a)), suggested to be a cancer-associated antigen.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>N-acetyl-alpha-neuraminosyl-(2-&gt;3)-beta-D-galactosyl-(1-&gt;3)-N-acetyl-beta-D-glucosaminyl-(1-&gt;3)-beta-D-galactosyl-(1-&gt;4)-beta-D-glucosyl-(1&lt;-&gt;1')-N-acyl-sphing-4-enine + CMP-N-acetyl-beta-neuraminate = N-acetyl-alpha-neuraminosyl-(2-&gt;3)-beta-D-galactosyl-(1-&gt;3)-[N-acetyl-alpha-neuraminosyl-(2-&gt;6)]-N-acetyl-beta-D-glucosaminyl-(1-&gt;3)-beta-D-galactosyl-(1-&gt;4)-beta-D-glucosyl-(1&lt;-&gt;1')-N-acyl-sphing-4-enine + CMP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:47884"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57812"/>
            <dbReference type="ChEBI" id="CHEBI:60377"/>
            <dbReference type="ChEBI" id="CHEBI:88073"/>
            <dbReference type="ChEBI" id="CHEBI:88079"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="15">
            <dbReference type="Rhea" id="RHEA:47885"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="14">
            <text>a ganglioside GM1b (d18:1(4E)) + CMP-N-acetyl-beta-neuraminate = a ganglioside GD1alpha (d18:1(4E)) + CMP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:41968"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57812"/>
            <dbReference type="ChEBI" id="CHEBI:60377"/>
            <dbReference type="ChEBI" id="CHEBI:78568"/>
            <dbReference type="ChEBI" id="CHEBI:78569"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="14">
            <dbReference type="Rhea" id="RHEA:41969"/>
        </physiologicalReaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Golgi apparatus membrane</location>
            <topology evidence="1">Single-pass type II membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the glycosyltransferase 29 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1I9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038772.1">
        <property type="protein sequence ID" value="ENSGAGP00000034245.1"/>
        <property type="gene ID" value="ENSGAGG00000024326.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000139">
        <property type="term" value="C:Golgi membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001665">
        <property type="term" value="F:alpha-N-acetylgalactosaminide alpha-2,6-sialyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001574">
        <property type="term" value="P:ganglioside biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009311">
        <property type="term" value="P:oligosaccharide metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd23976">
        <property type="entry name" value="GT29_ST6GALNAC5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1480.20:FF:000011">
        <property type="entry name" value="ST6 N-acetylgalactosaminide alpha-2,6-sialyltransferase 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1480.20">
        <property type="entry name" value="Glycosyl transferase family 29"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001675">
        <property type="entry name" value="Glyco_trans_29"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038578">
        <property type="entry name" value="GT29-like_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45906">
        <property type="entry name" value="ALPHA-N-ACETYL-NEURAMINYL-2,3-BETA-GALACTOSYL-1, 3-N-ACETYL-GALACTOSAMINIDE ALPHA-2,6-SIALYLTRANSFERASE-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45906:SF5">
        <property type="entry name" value="ALPHA-N-ACETYLGALACTOSAMINIDE ALPHA-2,6-SIALYLTRANSFERASE 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00777">
        <property type="entry name" value="Glyco_transf_29"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="3" id="KW-0328">Glycosyltransferase</keyword>
    <keyword evidence="9" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="10" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="11" id="KW-0472">Membrane</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0730">Sialic acid</keyword>
    <keyword evidence="6" id="KW-0735">Signal-anchor</keyword>
    <keyword evidence="4" id="KW-0808">Transferase</keyword>
    <keyword evidence="5" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8" id="KW-1133">Transmembrane helix</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004323"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022676"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022981"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00043744"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00051061"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00056921"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00074917"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00076990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00077803"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00081108"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00082858"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034245.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="313" mass="35744" checksum="4B6B2FACED9512FF" modified="2019-05-08" version="1">MCTSLLLMYSSIGGGGQKEPSQQQQQQVAVAASGKPETKQLGSSQRLPVGAGLLEGYISVLDNKPLKMHCKSCALVTSSGHLLGSKQGNRIDQTECVIRMNDAPTRGYGKDVGNKTSLRVIAHSSIQRILRNRNELLNMSQGTVFIFWGPSSYMRRDGKGLVYNNLQLMNQILPQLKVYMVSRHKMLQFDDLFKWETGKDRKISNTWLSTGWFTMTIALELCDRINVYGMVPPDFCRDPNHSSVPYHYYEPLGPDECTMYISHERGRKGSHHRFITEKRVFENWARIFNIHFFQPDWKPESLTINHLEIKPVF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1J0</accession>
    <name>A0A452J1J0_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000038699.1">
        <property type="protein sequence ID" value="ENSGAGP00000034173.1"/>
        <property type="gene ID" value="ENSGAGG00000024257.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005777">
        <property type="term" value="C:peroxisome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033228">
        <property type="entry name" value="SZT2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14918">
        <property type="entry name" value="KICSTOR COMPLEX PROTEIN SZT2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14918:SF3">
        <property type="entry name" value="KICSTOR COMPLEX PROTEIN SZT2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="694"/>
            <end position="722"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1402"/>
            <end position="1426"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1696"/>
            <end position="1736"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1763"/>
            <end position="1804"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="2321"/>
            <end position="2342"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="2354"/>
            <end position="2399"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="2629"/>
            <end position="2649"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="2764"/>
            <end position="2787"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="706"/>
            <end position="715"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="1402"/>
            <end position="1412"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="1">
        <location>
            <begin position="1413"/>
            <end position="1424"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="1701"/>
            <end position="1724"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="1775"/>
            <end position="1787"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="2325"/>
            <end position="2340"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="2364"/>
            <end position="2377"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="2384"/>
            <end position="2393"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034173.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="3331" mass="373164" checksum="D051A792564489F5" modified="2019-05-08" version="1">MVTCSPLLQVEDAGQVFLLMKKDYRISRNVRLAWFLNHLHQAIWAVPESELVKSDNELDVLSILPNGWQPHESVRPRPYLLVPSTQVTFLARQYRFVIELDLSPSTGIVDDSTGEIIFDEVFHALSRCLVGLLKPFRIPGSEITYQPEIFVTIQAYSSIIGLQSHQVLFQGCQLDETKRETFLQQIYTQLCAFENKVAEMLQQQYEPKPQVRQAKELSGRKPGISMVTADIGLVSMIRQGILALQLLPSNSSAGIIIITDGVTSVPDVAVCETLLNQLRSSTVACSFVQVGGVYSYDCSFGHVPNVELMKFIAMATFGAYLSTCLELDPLSLDLNVYHKAFLLYSFLRTRESLNPDYYCVSQHRLFNEHLVSASSNPALAMRRKKHTEKEVHADLVSIVSVRLREGYSIREVNITKGGSQLEVKLVLLWKHNMRIEYLAVAAWPLDPSKRSTWVEVTMEGSYDILHDISCTMRKPITSLYRTTVIRRFWNTLQSINQTDQMLVHLQSFDTVPEHFTIPESTKNGVPLFYIPPGSTTPVLSLQHSGSDCSHSQFASYWKPILSMDANFWQRWLHMHRIVLMLEHDTPVPKHLHTPGNNGRYSTIQCRISHSALTSLLRDWSSFVLVEGYSYVKLMPSSEDPAPSSFYVVRIISKAPCMVLRLGFPIGTPAQARNQIVEELQDRILQLRFPHRVQSKEATPKVKRKTFGSSSPSKSQPVAGAPPALSDRPCLVVLHKPLEKLLIRYEKLPSDYRAPFILTLEHPPVSGPLTMVANRTASSTLASLSRYFYHQRWIWCIQSGLAPAVPIAAVAHLLSTLTEVRLSEGFHFASSGDGIINMVLELDDSSGREKHTCIVQYVLFPPHSTSTKDSFSTDDDNDTEVEAIEMDAELNLVTECWVEPQSGHIHSTAENWRYLQGLPYQKIPKALYPRDLACISTMLTFEYISQLCQNKEWQCLMFLALLADPDMGAGSNVHEIPFQFSLLKLLPKCQQIEMFFLMLTGGPLGPSYYNFFHGCLQHDLSDREIPLADSDHMAFMEQVLQRDRDGHLPPFTLPVIRGKGSSNPAPVCDDPPASLGCPGTGRSEGLALGQWRCYAKLVSPQHLFLTFLPATFPGDEAPWGSQEWSGWCLGLSMGSSPSGRLSSAEAGQDPGEPGTSLRKDIHISFNCQASQPELSEGHRLRCPIYIYSCSLELLREQMVSPRAHRPPRDIFFRSQFLERPPNATWLDHKHKELVSYCTLLQEHSHQCYVKGLFKSLQQSHSISSQDLLTAVDYCEELLQEIDITHFLLTLCSHVRAFRESHRHFHTHPRAASQEQEEELPSRERGMLISESSVEIEDYSEQDHRGGQACSGLGPEGVSLTFTISCLSRQEKFLEIGSLHFKPVLSNPHYFFYCPPCAKKEEEASRDQADRKDSEDMEGSEAELAAEEGTVSGCCIATESDPELEVEYREKLENEFPDLGSLSDSNTVNHDDDSFSVLGGDSLNEQELLEQEMPPLFVHLTCSVKLRSQHSSMPVRSLPTCLGEVLSCLESCQAMASIDLSDLCVTLDIFVLTLPLEIEGVNADLHHNRWVWGWAGYPSPTDRVPGWPGSPELTPPDKEGGIPPSLSNLPGVHRQAVEATMNEIRWLLDDEMVSALRHSQAVSASILERVATHVYSSQGRPSCHSEVVPLQFVFGPEHSLEKFREVRAAHRLHPLETPASAVPSLQQPTPGGSTGAAQSHVGQERSSGADAEGLQPNHPQVSCPILQSLGAWGLCAATSALLSAPSSPAGASKQRPSRVQSLVSSQGSMDSDHLGYDGGSSDSECEEALMSDHDPKCPLMPDFWLIVKIQQDRVEVAMAEAGKEPAGECLHLHQTVVRKIGEICRVVNQRLLLQDLHDSHVCNTLLVAESEEDIWKSETPYSSYRQRAQPTDDYSTEESYLAATMQFMPGHFACPVVWSTLIHVHSRLKMGPNMGVSRAIQALRSVLNAFCVVNRKNMFVYQERSTKSVFYLRLSETTHSGKAWEGESIQGCVCRSLALTRSQEPIYMEDQTGSRSSLDTASCRSADSARPVGQVDRHIQLMVHGVAPAGPEITDELVKVLRRRLDEATLDIITVMLVRNCKLTPADVEFIQPPGSLPTEVLQFTLPPASLPWLHAVACYLRQNLLIFLHTPKYTDSNVEHHFKHYFQQGGSPDLDLYLYNKPGGQGTGGKGIACIALSFVDEHGSPLSLAHWERSSSLKLPPSLPGTDLLQDADFESLTAVSRHHAEPGTLKPGESAYVPGMALRLDVWEKGNISPLQLSEKLRSAVRHALCDAVMEFRVLPAPLCMEAACPLEELGTPPPPLHFTSASSSSSGEPITPTNKMGRRSFWDMLSKADSSELGSPKTTDDIVLERPEESRTRRRHKTESVKQHLSQERASATAELEQAQRHQVCQLQEGEVGTMHPLFSQTCQQWMAFMSRLGCPSVQQCSVEIVSRFLLSSILEEVVSLVTALASDTTVKVFQQHCCPQGATFLPYNPHQSSSPRPAAIRHFILLGRNFHQWRYKTEQGKCGGSNSSPSLAPRQRFLLLEILDKKLMLYTYNWSPDLGSSLNSSLTRLLQWQNARSHIVHCLLSQKLGLFHHYCHLDTPWHEDSKQPNPFLNSTLEVDALIRSSSPPPSKEQGRLSSSARSLPSLHFPPDMMPFDEALRDVTTLKGVPLIPELGPRDPVSWHGAQFLEIKSLERRELEKQMKIENLFVTWQQRSAQSNMPISLADLETLKQSSRLVHYCATPLLFDPAFRRPSELFQPGNSCRHRSSDSTASGRDRSHSCDSAEVLPCKLKQEPWLQEMCNAFLQQYIQYLQSMGFILVQVRPPSPAVTLAFLGTEGRGSFSYKPKADGSPKSTSPPVTTYHLQRALPGGIVLMELGFQGCYFCVKQYALECSRIPMGQSVNSQLSMLFTEECDKVRDLMHVHSFSYDFHLRIVHQYLLGSHMALRQGYHLTSFLEDFIAHHPDIPKFGRNHVFQGMLAMPTNMITAHQLYNYVADHANTYHMKPLRMARLAVGSDGRKGTPGMEQNEYALVSIWNSSGSYKDSEGLRHHDDFDVSLLVCHSAAPFEEQSEAERHVLRLRFYVIMTSQRELFPRLTADMRRFKKQPRLELWQEVEDSSEFYAGGAQVKLGPGLFYTLPLFPLLASEVAVAQKQLSSMVQLAKCHCRRDNLWKRLFLLEPLASDKLKLGKLSLGELEELLNAVHGKSIADIDPQLGCFLTMTVPWYQSLIKVLLSRFPQSCRHFHNVETGTQYLVVLNQKFTDCFVLVFLDSHSGKTSLTVVFREPFPVQPQNSESPLPQLVSTYHHLESVINTACFNLWTGLL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1J1</accession>
    <name>A0A452J1J1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Peptidase S1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J1J1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038800.1">
        <property type="protein sequence ID" value="ENSGAGP00000034266.1"/>
        <property type="gene ID" value="ENSGAGG00000024324.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00190">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.10:FF:000014">
        <property type="entry name" value="Complement factor D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.10.10">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009003">
        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043504">
        <property type="entry name" value="Peptidase_S1_PA_chymotrypsin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001314">
        <property type="entry name" value="Peptidase_S1A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001254">
        <property type="entry name" value="Trypsin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018114">
        <property type="entry name" value="TRYPSIN_HIS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033116">
        <property type="entry name" value="TRYPSIN_SER"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271:SF81">
        <property type="entry name" value="GRANZYME B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271">
        <property type="entry name" value="KALLIKREIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00089">
        <property type="entry name" value="Trypsin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00722">
        <property type="entry name" value="CHYMOTRYPSIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00020">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50494">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50240">
        <property type="entry name" value="TRYPSIN_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00134">
        <property type="entry name" value="TRYPSIN_HIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00135">
        <property type="entry name" value="TRYPSIN_SER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3 7" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="1 7" id="KW-0645">Protease</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 7" id="KW-0720">Serine protease</keyword>
    <keyword evidence="2 8" id="KW-0732">Signal</keyword>
    <keyword evidence="5" id="KW-0865">Zymogen</keyword>
    <feature type="signal peptide" evidence="8">
        <location>
            <begin position="1"/>
            <end position="21"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019234469" description="Peptidase S1 domain-containing protein" evidence="8">
        <location>
            <begin position="22"/>
            <end position="212"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase S1" evidence="9">
        <location>
            <begin position="24"/>
            <end position="203"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023145"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU363034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034266.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="212" mass="23204" checksum="4165EC4436375820" modified="2019-05-08" version="1">MQLQILLLLPMAFLLSPGAQAEEIIGGRVAKPHSCPYMAYLKRKPCKGEETCGGLLVREDFVLTAAHCLAELAELTDAIYTIPLPQAGRTVDPGSVCSVAGWGRTSLNSMRTSNILHEVDLEVMSDKTCQEDGLLQHYYKPLKMMCVGDPAEDKASFSGDSGGPLVCDRTAQGIVSFGKRNGSPPRVFTRVSGYIPWIKKTMRAPQRYLPSW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1J3</accession>
    <name>A0A452J1J3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">ETS domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 6">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 6">Belongs to the ETS family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1J3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038779.1">
        <property type="protein sequence ID" value="ENSGAGP00000034250.1"/>
        <property type="gene ID" value="ENSGAGG00000024331.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
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        <property type="term" value="P:cell differentiation"/>
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    <dbReference type="GO" id="GO:0045893">
        <property type="term" value="P:positive regulation of DNA-templated transcription"/>
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        <property type="project" value="UniProtKB-ARBA"/>
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        <property type="entry name" value="ETS translocation variant 5"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Winged helix-like DNA-binding domain superfamily/Winged helix DNA-binding domain"/>
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        <property type="entry name" value="ETS_PEA3_N"/>
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        <property type="entry name" value="ETS"/>
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    </dbReference>
    <dbReference type="PANTHER" id="PTHR11849:SF181">
        <property type="entry name" value="ETS TRANSLOCATION VARIANT 4"/>
        <property type="match status" value="1"/>
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    </dbReference>
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        <property type="entry name" value="ETSDOMAIN"/>
    </dbReference>
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        <property type="entry name" value="Winged helix' DNA-binding domain"/>
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    </dbReference>
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        <property type="entry name" value="ETS_DOMAIN_1"/>
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    </dbReference>
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        <property type="entry name" value="ETS_DOMAIN_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 6" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="5 6" id="KW-0539">Nucleus</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="ETS" evidence="7">
        <location>
            <begin position="301"/>
            <end position="381"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00005562"/>
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            <dbReference type="ARBA" id="ARBA00022553"/>
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    </evidence>
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            <dbReference type="ARBA" id="ARBA00023242"/>
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            <dbReference type="PROSITE" id="PS50061"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034250.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="443" mass="50529" checksum="39413442349E4143" modified="2019-05-08" version="1">MGAKRPMDPGLAPLQDSEELFQDLSQFQETWLTEAQVPDSDEQFVPDFHSENLAFHSPPAKVKKEPQSPCSDPALSCSHKQPFQYRNGEQCLYASAYDQTRQAGLKSPNPGTPIQSPLQHFPRQDRSFLSPQGPSQPTSSRGYLMEHSSVFQPPTEMCRSFPSSRGMGREAPVAYQRQMSEPCLQYHPQGFKQEYHDPMYEQANQLGNSSDHQYPAGVVIKQEQTDYAYDSDVPGCPSMYMNAEGYQSHSNTEGTLGYGYNKQMRPFADDVCVVPEKFEGDIKQEVGGYREGPPYQRRGSLQLWQFLVALLDDPTNSHFIAWTGRGMEFKLIEPEEVARLWGIQKNRPAMNYDKLSRSLRYYYEKGIMQKVAGERYVYKFVCEPEALFSLAFPDNQRPTLKAEFDRQISEEDTVPLSHLDENTAYIQDLGSLPSQPYSKSYPY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1K1</accession>
    <name>A0A452J1K1_9SAUR</name>
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        <recommendedName>
            <fullName evidence="9">Peptidase S1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J1K1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038812.1">
        <property type="protein sequence ID" value="ENSGAGP00000034276.1"/>
        <property type="gene ID" value="ENSGAGG00000024324.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="project" value="InterPro"/>
    </dbReference>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Complement factor D"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Serine protease 37"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="2"/>
    </dbReference>
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        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
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    </dbReference>
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    </dbReference>
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    </dbReference>
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        <property type="entry name" value="TRYPSIN_SER"/>
    </dbReference>
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        <property type="entry name" value="GRANZYME B"/>
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    </dbReference>
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    </dbReference>
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    </dbReference>
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        <property type="entry name" value="CHYMOTRYPSIN"/>
    </dbReference>
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    </dbReference>
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    </dbReference>
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    </dbReference>
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    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-1015">Disulfide bond</keyword>
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    <feature type="signal peptide" evidence="8">
        <location>
            <begin position="1"/>
            <end position="21"/>
        </location>
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    <feature type="chain" id="PRO_5019298638" description="Peptidase S1 domain-containing protein" evidence="8">
        <location>
            <begin position="22"/>
            <end position="267"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase S1" evidence="9">
        <location>
            <begin position="24"/>
            <end position="251"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023145"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU363034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034276.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="267" mass="29221" checksum="334A06EAE9AE852A" modified="2019-05-08" version="1">MQLQILLLLPMAFLLSPGAQAEEIIGGRVAKPHSCPYMAYLKRKPCKGEETCGGLLVREDFVLTAAHCADGDITVLLGAHNVRRDEETQQQVSVRRKIPHPRYNETSFENDLMLLQLAELAELTDAIYTIPLPQAGRTVDPGSVCSVAGWGRTSLNSMRTSNILHEVDLEVMSDKTCQEDGLLQHYYKPLKMMCVGDPAEDKASFSGDSGGPLVCDRTAQGIVSFGKRNGSPPRVFTRVSGYIPWIKKTMRGLNSPGSHAESPASLH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1K3</accession>
    <name>A0A452J1K3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Peptidase S1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J1K3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038801.1">
        <property type="protein sequence ID" value="ENSGAGP00000034267.1"/>
        <property type="gene ID" value="ENSGAGG00000024324.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00190">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.10:FF:000014">
        <property type="entry name" value="Complement factor D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.10.10">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009003">
        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043504">
        <property type="entry name" value="Peptidase_S1_PA_chymotrypsin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001254">
        <property type="entry name" value="Trypsin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033116">
        <property type="entry name" value="TRYPSIN_SER"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271:SF81">
        <property type="entry name" value="GRANZYME B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271">
        <property type="entry name" value="KALLIKREIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00089">
        <property type="entry name" value="Trypsin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00020">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50494">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50240">
        <property type="entry name" value="TRYPSIN_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00135">
        <property type="entry name" value="TRYPSIN_SER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="1" id="KW-0645">Protease</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0720">Serine protease</keyword>
    <keyword evidence="2 6" id="KW-0732">Signal</keyword>
    <keyword evidence="4" id="KW-0865">Zymogen</keyword>
    <feature type="signal peptide" evidence="6">
        <location>
            <begin position="1"/>
            <end position="21"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019123630" description="Peptidase S1 domain-containing protein" evidence="6">
        <location>
            <begin position="22"/>
            <end position="164"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase S1" evidence="7">
        <location>
            <begin position="1"/>
            <end position="155"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023145"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034267.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="164" mass="17973" checksum="C96F503DDACD0C29" modified="2019-05-08" version="1">MQLQILLLLPMAFLLSPGAQLAELAELTDAIYTIPLPQAGRTVDPGSVCSVAGWGRTSLNSMRTSNILHEVDLEVMSDKTCQEDGLLQHYYKPLKMMCVGDPAEDKASFSGDSGGPLVCDRTAQGIVSFGKRNGSPPRVFTRVSGYIPWIKKTMRAPQRYLPSW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1K4</accession>
    <name>A0A452J1K4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Peptidase S1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J1K4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038791.1">
        <property type="protein sequence ID" value="ENSGAGP00000034258.1"/>
        <property type="gene ID" value="ENSGAGG00000024324.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00190">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.10:FF:000014">
        <property type="entry name" value="Complement factor D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.10:FF:000005">
        <property type="entry name" value="Serine protease 37"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.10.10">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009003">
        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043504">
        <property type="entry name" value="Peptidase_S1_PA_chymotrypsin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001314">
        <property type="entry name" value="Peptidase_S1A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001254">
        <property type="entry name" value="Trypsin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018114">
        <property type="entry name" value="TRYPSIN_HIS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033116">
        <property type="entry name" value="TRYPSIN_SER"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271:SF81">
        <property type="entry name" value="GRANZYME B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271">
        <property type="entry name" value="KALLIKREIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00089">
        <property type="entry name" value="Trypsin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00722">
        <property type="entry name" value="CHYMOTRYPSIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00020">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50494">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50240">
        <property type="entry name" value="TRYPSIN_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00134">
        <property type="entry name" value="TRYPSIN_HIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00135">
        <property type="entry name" value="TRYPSIN_SER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3 7" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="1 7" id="KW-0645">Protease</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 7" id="KW-0720">Serine protease</keyword>
    <keyword evidence="2" id="KW-0732">Signal</keyword>
    <keyword evidence="5" id="KW-0865">Zymogen</keyword>
    <feature type="domain" description="Peptidase S1" evidence="8">
        <location>
            <begin position="8"/>
            <end position="235"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023145"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU363034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034258.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="244" mass="27108" checksum="0132F9994EA14B7E" modified="2019-05-08" version="1">PQLFPKEIIGGRVAKPHSCPYMAYLKRKPCKGEETCGGLLVREDFVLTAAHCADGDITVLLGAHNVRRDEETQQQVSVRRKIPHPRYNETSFENDLMLLQLAELAELTDAIYTIPLPQAGRTVDPGSVCSVAGWGRTSLNSMRTSNILHEVDLEVMSDKTCQEDGLLQHYYKPLKMMCVGDPAEDKASFSGDSGGPLVCDRTAQGIVSFGKRNGSPPRVFTRVSGYIPWIKKTMRAPQRYLPSW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1K7</accession>
    <name>A0A452J1K7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">GTP-binding protein 8</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the TRAFAC class TrmE-Era-EngA-EngB-Septin-like GTPase superfamily. EngB GTPase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1K7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038714.1">
        <property type="protein sequence ID" value="ENSGAGP00000034188.1"/>
        <property type="gene ID" value="ENSGAGG00000024287.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005525">
        <property type="term" value="F:GTP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd01876">
        <property type="entry name" value="YihA_EngB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000857">
        <property type="entry name" value="GTP-binding protein 8 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052279">
        <property type="entry name" value="EngB_GTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR030393">
        <property type="entry name" value="G_ENGB_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006073">
        <property type="entry name" value="GTP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019987">
        <property type="entry name" value="GTP-bd_ribosome_bio_YsxC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR03598">
        <property type="entry name" value="GTPase_YsxC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46498">
        <property type="entry name" value="GTP-BINDING PROTEIN 8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46498:SF1">
        <property type="entry name" value="GTP-BINDING PROTEIN 8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01926">
        <property type="entry name" value="MMR_HSR1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51706">
        <property type="entry name" value="G_ENGB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0342">GTP-binding</keyword>
    <keyword evidence="6" id="KW-0460">Magnesium</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="5" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="EngB-type G" evidence="8">
        <location>
            <begin position="1"/>
            <end position="171"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009638"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00015370"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51706"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034188.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="177" mass="19919" checksum="A3F704ECF12F70D1" modified="2019-05-08" version="1">MVCFVGRSNVGKSSLIKALFSLAPDVEVRVSKTPGHTKKMNFFKVGKYFTLVDMPGYGYRAPQDFADMVEAYLQERQNLKRTFLLVDSVVGFQKADHTAIEMLEEFGIPYVFVLTKIDKASKGLLVKNVLEIQDFAEKHTQGCFPQLFPVSSIEYSGIHLLRCFIAHVTGNLPMMNS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1M5</accession>
    <name>A0A452J1M5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">Pikachurin</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="18">EGF-like, fibronectin type-III and laminin G-like domain-containing protein</fullName>
        </alternativeName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">Involved in both the retinal photoreceptor ribbon synapse formation and physiological functions of visual perception. Plays a key role in the synaptic organization of photoreceptors by mediating transsynaptic interaction between alpha-dystroglycan and GPR179 on the postsynaptic membrane. Necessary for proper bipolar dendritic tip apposition to the photoreceptor ribbon synapse. Promotes matrix assembly and cell adhesiveness.</text>
    </comment>
    <comment type="subunit">
        <text evidence="16">Interacts with DAG1 alpha-dystroglycan. Interacts with GPR158 and GPR179; transsynaptic interaction is required for synaptic organization of photoreceptor cells.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="13">Presynaptic active zone</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="14">Synaptic cleft</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="19">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1M5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038841.1">
        <property type="protein sequence ID" value="ENSGAGP00000034301.1"/>
        <property type="gene ID" value="ENSGAGG00000024373.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="project" value="UniProtKB-ARBA"/>
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        <property type="term" value="C:presynaptic active zone"/>
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        <property type="project" value="UniProtKB-SubCell"/>
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        <property type="term" value="C:synaptic cleft"/>
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        <property type="project" value="UniProtKB-SubCell"/>
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        <property type="term" value="F:calcium ion binding"/>
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        <property type="entry name" value="Immunoglobulins"/>
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        <property type="entry name" value="Ig-like_fold"/>
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        <property type="entry name" value="Neurexin-related_CASP"/>
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        <property type="entry name" value="PIKACHURIN"/>
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        <property type="entry name" value="PIKACHURIN-LIKE PROTEIN"/>
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        <property type="entry name" value="Laminin_G_1"/>
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        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="3"/>
    </dbReference>
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        <property type="entry name" value="Fibronectin type III"/>
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        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="3"/>
    </dbReference>
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    </dbReference>
    <dbReference type="PROSITE" id="PS50025">
        <property type="entry name" value="LAM_G_DOMAIN"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="12" id="KW-0966">Cell projection</keyword>
    <keyword evidence="9 19" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="4 19" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="3" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11" id="KW-0357">Heparan sulfate</keyword>
    <keyword evidence="7" id="KW-0654">Proteoglycan</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="5 20" id="KW-0732">Signal</keyword>
    <keyword evidence="8" id="KW-0770">Synapse</keyword>
    <feature type="signal peptide" evidence="20">
        <location>
            <begin position="1"/>
            <end position="24"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019431059" description="Pikachurin" evidence="20">
        <location>
            <begin position="25"/>
            <end position="986"/>
        </location>
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    <feature type="domain" description="Fibronectin type-III" evidence="23">
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            <begin position="38"/>
            <end position="146"/>
        </location>
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    <feature type="domain" description="Fibronectin type-III" evidence="23">
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            <begin position="154"/>
            <end position="249"/>
        </location>
    </feature>
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            <end position="582"/>
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        <location>
            <begin position="828"/>
            <end position="837"/>
        </location>
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        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00064047"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00068283"/>
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            <dbReference type="ARBA" id="ARBA00081917"/>
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        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
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        </source>
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    <evidence type="ECO:0000259" key="21">
        <source>
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        </source>
    </evidence>
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        <source>
            <dbReference type="PROSITE" id="PS50853"/>
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    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034301.1"/>
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    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="986" mass="108527" checksum="201DFED3AE096F04" modified="2019-05-08" version="1">MDLQSPARCFALILLASCIGHTAASRRNIPRRQDRLGPPLDILMDSLNCTAFSIQWRMPRQHASTITGYTVFYSEVEADKPVKQLSHDVPLSLDMLSMSQLDGQAIFDVVIGNLKPGTPHRISVGAYGWAGKGRPSMPRDITTLSQDKCMPPEPPYQPQVIVVSDSEVALSWKPGVNEGSSPIQYYTLEFLRPDLDSNWTVIKEQIQMESMVLKGLIPNTKYQFAIRAANSYGSSLPSTPSDIIRTFSSEELGSGSYGHRYITGTIVGDDDGFDIDDSDYDIYIEELKPLPAIKGGNRKFLVETKVGPGSSTRLLSRLITTTSDPITSPATTMLLIPTTTASTTTAKATRRNGQASAVRLFDLSCDETICSADSFCVNDYDRGGSRCHCNLGKGGETCTEDIIIQYPQFSGYSYITFEPLKNSYQTFQITLEFRAEAEDGLLLYCGENEHGRGDFMSLAIIRRNIQFRFNCGTGVAVIMSESKIKLGNWHSVTVFRDGLNGWLRLDNDTPVTGKSQGQYSKITFRTPFYLGGAPSAYWLVRAAGTNRGFQGCVQSLIVNGKIIDMRPWPLGKALSGADVGECSSGICDEASCINSGTCTASQADAYVCLCPLGFKGRHCEEAFILTIPQFNESLRSFATTPWPLEPQNYLSFMEFEMTFRPDSENGVLLYSYDTDSKDFLSLNMVASYVEFRFDCGSGTAIIRSEEPVILGQWHELRVSRTAKNAILQVDKQKPVEGMAEGAFTQIKCNTGIFIGGVPNYDNLKKNSGILKPFSGSIQKIILNDRTIDVKQDFASGVNVENAVHPCVNSPCANGGSCMPKKDSYECDCPLGFDGQHCQKECGNYCFNTVTEAIEIPQFIGRSYLTYDNPDILKRVSGSRSNVFMRFKTTLKDGLLMWRGGSPMRPNSDFISLGLQEGALVFSYNLGSGIASIMVNGSFNDGRWHRVKAVRDGQSGKVTVDDYGARTGKSPGMMRQLNINGDLYIGK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1M6</accession>
    <name>A0A452J1M6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Peptidase S1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J1M6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038819.1">
        <property type="protein sequence ID" value="ENSGAGP00000034283.1"/>
        <property type="gene ID" value="ENSGAGG00000024324.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00190">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.10:FF:000005">
        <property type="entry name" value="Serine protease 37"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.10.10">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009003">
        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043504">
        <property type="entry name" value="Peptidase_S1_PA_chymotrypsin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001314">
        <property type="entry name" value="Peptidase_S1A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001254">
        <property type="entry name" value="Trypsin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018114">
        <property type="entry name" value="TRYPSIN_HIS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033116">
        <property type="entry name" value="TRYPSIN_SER"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271:SF81">
        <property type="entry name" value="GRANZYME B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271">
        <property type="entry name" value="KALLIKREIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00089">
        <property type="entry name" value="Trypsin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00722">
        <property type="entry name" value="CHYMOTRYPSIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00020">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50494">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="TRYPSIN_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00134">
        <property type="entry name" value="TRYPSIN_HIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00135">
        <property type="entry name" value="TRYPSIN_SER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3 7" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="1 7" id="KW-0645">Protease</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 7" id="KW-0720">Serine protease</keyword>
    <keyword evidence="2 8" id="KW-0732">Signal</keyword>
    <keyword evidence="5" id="KW-0865">Zymogen</keyword>
    <feature type="signal peptide" evidence="8">
        <location>
            <begin position="1"/>
            <end position="21"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019124830" description="Peptidase S1 domain-containing protein" evidence="8">
        <location>
            <begin position="22"/>
            <end position="262"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase S1" evidence="9">
        <location>
            <begin position="33"/>
            <end position="246"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023145"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU363034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034283.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="262" mass="28712" checksum="E8390801E7484634" modified="2019-05-08" version="1">MQLQILLLLPMAFLLSPGAQAGEYGWQWGPLGLWGQMEAKPHSCPYMAYLKRKPCKGEETCGGLLVREDFVLTAAHCADGDITVLLGAHNVRRDEETQQQVSVRRKIPHPRYNETSFENDLMLLQLAELAELTDAIYTIPLPQAGRTVDPGSVCSVAGWGRTSLNSMRTSNILHEVDLEVMSDKTCQEDGWGSEGRTDQCCGDSGGPLVCDRTAQGIVSFGKRNGSPPRVFTRVSGYIPWIKKTMRGLNSPGSHAESPASLH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1P0</accession>
    <name>A0A452J1P0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Calcitonin peptide-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the calcitonin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1P0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038741.1">
        <property type="protein sequence ID" value="ENSGAGP00000034214.1"/>
        <property type="gene ID" value="ENSGAGG00000024303.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031716">
        <property type="term" value="F:calcitonin receptor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005179">
        <property type="term" value="F:hormone activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007189">
        <property type="term" value="P:adenylate cyclase-activating G protein-coupled receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051480">
        <property type="term" value="P:regulation of cytosolic calcium ion concentration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.2190">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021117">
        <property type="entry name" value="Calcitonin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021116">
        <property type="entry name" value="Calcitonin/adrenomedullin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018360">
        <property type="entry name" value="Calcitonin_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015476">
        <property type="entry name" value="Calcitonin_gene-rel_peptide"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001693">
        <property type="entry name" value="Calcitonin_peptide-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10505:SF16">
        <property type="entry name" value="CALCITONIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10505">
        <property type="entry name" value="CALCITONIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00214">
        <property type="entry name" value="Calc_CGRP_IAPP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00817">
        <property type="entry name" value="CALCITONINB"/>
    </dbReference>
    <dbReference type="SMART" id="SM00113">
        <property type="entry name" value="CALCITONIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00258">
        <property type="entry name" value="CALCITONIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0165">Cleavage on pair of basic residues</keyword>
    <keyword evidence="6 7" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0964">Secreted</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <feature type="domain" description="Calcitonin peptide-like" evidence="8">
        <location>
            <begin position="112"/>
            <end position="154"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="7">
        <location>
            <begin position="115"/>
            <end position="120"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009222"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022685"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PIRSR" id="PIRSR621116-50"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="SMART" id="SM00113"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034214.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="159" mass="17860" checksum="8D8D6ABD503BCD0E" modified="2019-05-08" version="1">MYLEKDCIFSRSVKIEEAGKWPLFDVTLCCREDIMVMLKISPFLAVYALVMCQMDSSQATPLRPGLDSITDRVTLSDYEARRLLNALVKEFVQMRAEDVEQETEGNSVTAQKRACNTATCVTHRLADFLSRSGGVGKNNFVPTNVGSKAFGRRRRNAQI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1P1</accession>
    <name>A0A452J1P1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">RING-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J1P1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038857.1">
        <property type="protein sequence ID" value="ENSGAGP00000034316.1"/>
        <property type="gene ID" value="ENSGAGG00000024374.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061630">
        <property type="term" value="F:ubiquitin protein ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016567">
        <property type="term" value="P:protein ubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd16679">
        <property type="entry name" value="RING-H2_RNF38"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.40.10:FF:000024">
        <property type="entry name" value="RING finger protein 44 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001841">
        <property type="entry name" value="Znf_RING"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46171:SF1">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE RNF38"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46171">
        <property type="entry name" value="GH10160P"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13639">
        <property type="entry name" value="zf-RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00184">
        <property type="entry name" value="RING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57850">
        <property type="entry name" value="RING/U-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50089">
        <property type="entry name" value="ZF_RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0862">Zinc</keyword>
    <keyword evidence="2 4" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="RING-type" evidence="6">
        <location>
            <begin position="380"/>
            <end position="421"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="1"/>
            <end position="63"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="1"/>
            <end position="21"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="5">
        <location>
            <begin position="32"/>
            <end position="51"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034316.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="432" mass="48594" checksum="379148DD9AF4650B" modified="2019-05-08" version="1">MRPWEMTSNRQPPSARPNQHHFSGERCNTPARNRRSPPVRRQRARRDRLSRHNSISQDENYHHLPYGQQQAIEEPRAFHPPNVSPRMLHPAAHPPQQNAVMVDIHEQIHQGTVPVSYTVTTVAPHGIPLCTGQHIPACNTQQVPGCSVVFSGQHLPVCSVPPPMLQACSVQHLPVPYAAFPPLISSDPFLLHPPHLSPHHPPHLPPPGQFVPFQTQQSRSPLQRIENEVELLGEHLPVGGFTYPPSAHPPTLTPSAPLQFLTHDPLHQEVSFGVPYPPFMPRRLTGRSRYRSQQPIPPPPYHPSLLPYVLSMLPVPPAVGPAFSFELDVEDGEVENYEALLNLAERLGEAKPRGLTKADIEQLPSYRFNPNNHQSEQTLCVVCMCDFESRQLLRVLPCNHEFHAKCVDKWLKANRTCPICRADASEVHRDSE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1P4</accession>
    <name>A0A452J1P4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">HMG box domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the DNA mismatch repair MutL/HexB family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1P4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038843.1">
        <property type="protein sequence ID" value="ENSGAGP00000034303.1"/>
        <property type="gene ID" value="ENSGAGG00000024375.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032389">
        <property type="term" value="C:MutLalpha complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140664">
        <property type="term" value="F:ATP-dependent DNA damage sensor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030983">
        <property type="term" value="F:mismatched DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006298">
        <property type="term" value="P:mismatch repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd16926">
        <property type="entry name" value="HATPase_MutL-MLH-PMS-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd03485">
        <property type="entry name" value="MutL_Trans_hPMS_1_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.30.10:FF:000026">
        <property type="entry name" value="PMS1 homolog 1, mismatch repair system component"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.230.10:FF:000030">
        <property type="entry name" value="PMS1 homolog 1, mismatch repair system component"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.565.10:FF:000017">
        <property type="entry name" value="PMS1 homolog 1, mismatch repair system component"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.30.10">
        <property type="entry name" value="High mobility group box domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.565.10">
        <property type="entry name" value="Histidine kinase-like ATPase, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014762">
        <property type="entry name" value="DNA_mismatch_repair_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013507">
        <property type="entry name" value="DNA_mismatch_S5_2-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036890">
        <property type="entry name" value="HATPase_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009071">
        <property type="entry name" value="HMG_box_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036910">
        <property type="entry name" value="HMG_box_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002099">
        <property type="entry name" value="MutL/Mlh/PMS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038973">
        <property type="entry name" value="MutL/Mlh/Pms-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020568">
        <property type="entry name" value="Ribosomal_Su5_D2-typ_SF"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00585">
        <property type="entry name" value="mutl"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10073">
        <property type="entry name" value="DNA MISMATCH REPAIR PROTEIN MLH, PMS, MUTL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10073:SF54">
        <property type="entry name" value="PMS1 PROTEIN HOMOLOG 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01119">
        <property type="entry name" value="DNA_mis_repair"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13589">
        <property type="entry name" value="HATPase_c_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00505">
        <property type="entry name" value="HMG_box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01340">
        <property type="entry name" value="DNA_mis_repair"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00398">
        <property type="entry name" value="HMG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55874">
        <property type="entry name" value="ATPase domain of HSP90 chaperone/DNA topoisomerase II/histidine kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47095">
        <property type="entry name" value="HMG-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54211">
        <property type="entry name" value="Ribosomal protein S5 domain 2-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00058">
        <property type="entry name" value="DNA_MISMATCH_REPAIR_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50118">
        <property type="entry name" value="HMG_BOX_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="2" id="KW-0539">Nucleus</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="HMG box" evidence="4">
        <location>
            <begin position="541"/>
            <end position="609"/>
        </location>
    </feature>
    <feature type="DNA-binding region" description="HMG box" evidence="2">
        <location>
            <begin position="541"/>
            <end position="609"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="337"/>
            <end position="361"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="422"/>
            <end position="443"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="496"/>
            <end position="521"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="602"/>
            <end position="629"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="345"/>
            <end position="361"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="429"/>
            <end position="439"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="511"/>
            <end position="521"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="602"/>
            <end position="623"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006082"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00267"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50118"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034303.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="905" mass="102292" checksum="BD9305876EC77D9D" modified="2019-05-08" version="1">MKQLSAETVRLLSSSQVITSVVSVVKELIENSLDANATSIDVKLENYGFDKIEVRDNGDGIKAADVPVMAVKHYTSKISCSEDLESLTTYGFRGEALGSICCISEVLITTKTAADDFSTQYILDSSGHVTSQKPSHLGQGTTVAALKLFKNLPVRKQFYSTVKKCKEELKKIQDLLMAYGIIKPELRIIFTHNKVFLSGFLPKPESDNSLTSLSSSERSFIYINNRPVHQKEILKLVRWFYNLKSHKDSSRSYPVFFINITVPTSAVDVNLTPDKTQVLLHNKESVLSAVENVLTSLYGLPPDTVSCETNKTDVTSADVVVNETEQTDVLVNKMEPFGNSDPHAHTSSFSSSNDVQNGQAGKNTEVCLKHQRFSSDNTHGLLNRKEVSGTDVTVSDISQDSSINNLSCEDSQNGYSVTLTLDGSTHMDTQSKNRSEHPLRASYANEVEKNEANTVPEYVSEISADNWSKGNALKNATGENLEPVKILIPEVGGITDHKANENTDEQDRDQQSSNQNIKKTNVVNDKAGQIMAYDLISNQIIRKPMSAVALFTQEYRSRLLTDHPKANMEDLMLKIEGLWEKLSREEKKKYEEKATKDLERYNRQTKKAVEQRIQRSTKETEKRYKSKLKTSLSDQQKLDKLFQSQIEKKQRHNQDMKIVQVPFSMSSFKHKLQRLEKKNFDKNESCLICHLSFPDAWIIASEKKIMLLNPYRVEEALLYKKLLEQHKLPVEKLDKAIVLTDSLFSGSHYMDSLYNMTKDCHNLNGSIYLSDPRLIANGFRIKMTPGHSAIENQLEIEGMANCLPFYGISDLKEILNAVINKNAKEVYECRPLKVINYLEGEAVRISRQLPLYMSKEDVEDTIYRMKQQLGNENKGCVHGRPFFHHLTDIPEFKLQAEIQGSYSVK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J210</accession>
    <name>A0A452J210_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7 26">Neuromodulin</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="22 26">Axonal membrane protein GAP-43</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="23 26">Growth-associated protein 43</fullName>
        </alternativeName>
    </protein>
    <organism evidence="30 31">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="31" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="30" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="30" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="21 26">This protein is associated with nerve growth. It is a major component of the motile 'growth cones' that form the tips of elongating axons. Plays a role in axonal and dendritic filopodia induction.</text>
    </comment>
    <comment type="subunit">
        <text evidence="25">Identified in a complex containing FGFR4, NCAM1, CDH2, PLCG1, FRS2, SRC, SHC1, GAP43 and CTTN. Interacts (via IQ domain) with calmodulin. Binds calmodulin with a greater affinity in the absence of Ca(2+) than in its presence.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="26">Cell membrane</location>
            <topology evidence="26">Peripheral membrane protein</topology>
            <orientation evidence="26">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="5 26">Cell projection</location>
            <location evidence="5 26">Growth cone membrane</location>
            <topology evidence="5 26">Peripheral membrane protein</topology>
            <orientation evidence="5 26">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="24 26">Synapse</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="26">Cell projection</location>
            <location evidence="26">Filopodium membrane</location>
            <topology evidence="26">Peripheral membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cell projection</location>
            <location evidence="3">Axon</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cell projection</location>
            <location evidence="1">Dendrite</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Perikaryon</location>
        </subcellularLocation>
    </comment>
    <comment type="PTM">
        <text evidence="26">Palmitoylated. Palmitoylation is essential for plasma membrane association.</text>
    </comment>
    <comment type="similarity">
        <text evidence="6 26">Belongs to the neuromodulin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J210"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038976.1">
        <property type="protein sequence ID" value="ENSGAGP00000034426.1"/>
        <property type="gene ID" value="ENSGAGG00000024465.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030425">
        <property type="term" value="C:dendrite"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031527">
        <property type="term" value="C:filopodium membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032584">
        <property type="term" value="C:growth cone membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043204">
        <property type="term" value="C:perikaryon"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0014069">
        <property type="term" value="C:postsynaptic density"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035727">
        <property type="term" value="F:lysophosphatidic acid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1901981">
        <property type="term" value="F:phosphatidylinositol phosphate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001786">
        <property type="term" value="F:phosphatidylserine binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016198">
        <property type="term" value="P:axon choice point recognition"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031103">
        <property type="term" value="P:axon regeneration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0040008">
        <property type="term" value="P:regulation of growth"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042246">
        <property type="term" value="P:tissue regeneration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd23767">
        <property type="entry name" value="IQCD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.190:FF:000075">
        <property type="entry name" value="Neuromodulin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.190">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000048">
        <property type="entry name" value="IQ_motif_EF-hand-BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001422">
        <property type="entry name" value="Neuromodulin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017454">
        <property type="entry name" value="Neuromodulin_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018947">
        <property type="entry name" value="Neuromodulin_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033137">
        <property type="entry name" value="Neuromodulin_P_site"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018243">
        <property type="entry name" value="Neuromodulin_palmitoyl_site"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10699">
        <property type="entry name" value="NEUROMODULIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10699:SF15">
        <property type="entry name" value="NEUROMODULIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00612">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06614">
        <property type="entry name" value="Neuromodulin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10580">
        <property type="entry name" value="Neuromodulin_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00215">
        <property type="entry name" value="NEUROMODULIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00015">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00412">
        <property type="entry name" value="NEUROMODULIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00413">
        <property type="entry name" value="NEUROMODULIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="14 26" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="9 26" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="19 26" id="KW-0966">Cell projection</keyword>
    <keyword evidence="10" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8 26" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="13 26" id="KW-0221">Differentiation</keyword>
    <keyword evidence="12 26" id="KW-0341">Growth regulation</keyword>
    <keyword evidence="20 26" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="17" id="KW-0472">Membrane</keyword>
    <keyword evidence="15 26" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="18 26" id="KW-0564">Palmitate</keyword>
    <keyword evidence="11 26" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="31" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="16 26" id="KW-0770">Synapse</keyword>
    <feature type="domain" description="Neuromodulin N-terminal" evidence="29">
        <location>
            <begin position="2"/>
            <end position="32"/>
        </location>
    </feature>
    <feature type="domain" description="Neuromodulin C-terminal" evidence="28">
        <location>
            <begin position="69"/>
            <end position="241"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="27">
        <location>
            <begin position="1"/>
            <end position="241"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="27">
        <location>
            <begin position="9"/>
            <end position="33"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="27">
        <location>
            <begin position="67"/>
            <end position="86"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="27">
        <location>
            <begin position="87"/>
            <end position="103"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="27">
        <location>
            <begin position="125"/>
            <end position="158"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="27">
        <location>
            <begin position="165"/>
            <end position="177"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="27">
        <location>
            <begin position="182"/>
            <end position="206"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="27">
        <location>
            <begin position="208"/>
            <end position="220"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="27">
        <location>
            <begin position="232"/>
            <end position="241"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004279"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004484"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004489"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004503"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00005890"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00021591"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022604"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023139"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00023288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00025215"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00030597"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00033250"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00034103"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00046855"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="RuleBase" id="RU368113"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="28">
        <source>
            <dbReference type="Pfam" id="PF06614"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="29">
        <source>
            <dbReference type="Pfam" id="PF10580"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="30">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034426.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="31">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="241" mass="25207" checksum="8966825D2D3C98C4" modified="2019-05-08" version="1">MLCCMRRTKQVEKNDDVDQKIEQDGIKPEDKAHKAATKIQASFRGHITRKKLKGEKKGDVQAADAEAAEKKDEAPSDGVAENKESDAPAATEAAPSDSAPPAEGSKDGSAPSEEKQGEGTADAGSEQPAPQAATPASSSEDKPAPTATETESATKASTDNSPSLKADEAQDKEEPKQADVPAAVTTTAAEDATAKAIAQTETAESSQTEEKTDAVEETKPTESAQQEEVKEDENKADQENA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J213</accession>
    <name>A0A452J213_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="4">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the eukaryotic ribosomal protein eL22 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J213"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038983.1">
        <property type="protein sequence ID" value="ENSGAGP00000034432.1"/>
        <property type="gene ID" value="ENSGAGG00000024474.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:1990904">
        <property type="term" value="C:ribonucleoprotein complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005840">
        <property type="term" value="C:ribosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003735">
        <property type="term" value="F:structural constituent of ribosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002181">
        <property type="term" value="P:cytoplasmic translation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1360.210:FF:000001">
        <property type="entry name" value="60S ribosomal protein L22 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1360.210">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002671">
        <property type="entry name" value="Ribosomal_eL22"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038526">
        <property type="entry name" value="Ribosomal_eL22_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10064">
        <property type="entry name" value="60S RIBOSOMAL PROTEIN L22"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10064:SF1">
        <property type="entry name" value="RIBOSOMAL PROTEIN EL22-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01776">
        <property type="entry name" value="Ribosomal_L22e"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0687">Ribonucleoprotein</keyword>
    <keyword evidence="2" id="KW-0689">Ribosomal protein</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007817"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022980"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023274"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034432.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="141" mass="16691" checksum="11881FFF3639335C" modified="2019-05-08" version="1">MFCCCTVCPFWELAKLTFNANLQRDRKSKKSAWKFSLDLTHPVEDGIFDSGNFEQFLKEKVKVNGKTGNLGNIVHIERFKNKITVVSEKQFSKRYLKYLTKKYLKKNNLRDWLRVVASDKETYELRYFQISQDEEGSESEE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIV7</accession>
    <name>A0A452GIV7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">DUF4704 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000001812.1">
        <property type="protein sequence ID" value="ENSGAGP00000001593.1"/>
        <property type="gene ID" value="ENSGAGG00000000400.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019901">
        <property type="term" value="F:protein kinase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008104">
        <property type="term" value="P:intracellular protein localization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050865">
        <property type="entry name" value="BEACH_Domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031570">
        <property type="entry name" value="NBEA/BDCP_DUF4704"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13743">
        <property type="entry name" value="BEIGE/BEACH-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13743:SF62">
        <property type="entry name" value="NEUROBEACHIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15787">
        <property type="entry name" value="DUF4704"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0853">WD repeat</keyword>
    <feature type="domain" description="DUF4704" evidence="3">
        <location>
            <begin position="23"/>
            <end position="438"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="462"/>
            <end position="485"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="730"/>
            <end position="766"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1330"/>
            <end position="1349"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="471"/>
            <end position="484"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="730"/>
            <end position="754"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="755"/>
            <end position="766"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="1334"/>
            <end position="1344"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022574"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="Pfam" id="PF15787"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001593.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1457" mass="161318" checksum="2148A1A5AAE4A7E5" modified="2025-10-08" version="2">MSVDWKRELEIGENGEKFCIAVHRILQSSVAMQEQMLGGKGFLVIGYLLEKSSRVHITRAVLEQFLSFAKYLDGLSHGAPLLKQLCDHILFNPAIWIHTPAKVQLSLYTYLSAEFIGTATIYTTIRRVGTVLQLMHTLKYYYWVVNPADSSGITPKGLEGPRPSQKEIISLRAFMLLFLKQLILKDRGVKEDELQSILNYLLTMHEDENIHDVLQLLVALMSEHPASMIPAFDQRNGIRVIYKLLASKSESIWVQALKVLGYFLKHLGHKRKVEIMHTHSLFTLLGERLMLHTNTVTITTYNTLYEILTEQVCTQVVHKPHPEPDSTVKIQNPMILKVVATLLKNSTPSAELMEVRRLFLSDMIKLFSNSRENRRCLLQCSVWQDWMFSLGYINPKNSEEQKITEMVYNIFRILLYHAIKYEWGGWRVWVDTLSIAHSKVTYEAHKEYLAKMYEEYQRQEEENIKKGKKGNVSTISGLSSQTTGAKGGLEIREIEDLSQSQSPESETDYPVSTDTRDLLMATKVSDEVLGSSDRPGGGVHVEVHDLLVDIKAEKVEATEVKLDDMDLSPETLVTGENGALVEVESLLDNVYCAAVEKLQNNVHGSVGIIKKSEEKDNGPLITLADEKDEPSTNNTSFLFDKIPSQEEKLLPELSSNHITIPNVQETQMHLGVSDDLGLLAHMTGSVDITCTSSIIEDKEFKIHTTSDGMSNLSERELGSSSKGLEYAEMTATTLEPESSGSKSLPNVDTGSIISDTERSDDGKEAGKEIRKIQTTTTTQAVQGRSVTQQDRDLRVDLGFRGMPMTEEQRRQFSPGPRTTMFRIPEFKWSPMHQRLLTDLLFALETDVHVWRSHSTKSVMDFVNSNENIIFVHNTIHLISQMVDNIIIACGGILPLLSAATSPTTELENIEVTQGMSAETAVTFLSRLMAMVDVLVFASSLNFSEIEAEKNMSSGGLMRQCLRLVCCVAVRNCLECRQRQRERVNKSSLISSKTQETLQGVTATALTKSPLENVPGNLSPIKDPDRLLQDVDINRLRAVVFRDVDDSKQAQFLALAVVYFISVLMVSKYRDILEPQRETARCGSQAGRNIRQEINSPTSTVVVIPSIPHPSLNHGFLAKLIPEQSFAHSFYKETPAVFPENIKEKETPTPVEDIQLESSIPHTDSGIGEEQMPSILNGTDLETSAGPDAMSELFSTLSSEVKKSQESLTESPSEILKPAPSISSISQSKGMNVKEILKSLVAAPIEIAECGPDPIPYPDPALKREAQSILPMQFHSFDRSVVVPVKKPPPGSLAVTTVGAATAGGVLPPGTTPNIFAATGATPKSMINTTGAVDSGSSSSSSSSSFVNGATSKNLPAVQTVAPMPEDSAENMSITAKLERALEKVAPLLREIFVDFAPFLSRTLLGSHGQELLIEGLVCMKSSTSVVELVMLLCSQEWQNSIQKNAGLAFIELINEGR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIV8</accession>
    <name>A0A452GIV8_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="12">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="pathway">
        <text evidence="2">Protein modification; protein glycosylation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Golgi apparatus membrane</location>
            <topology evidence="1">Single-pass type II membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="10">Belongs to the glycosyltransferase 14 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIV8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001829.1">
        <property type="protein sequence ID" value="ENSGAGP00000001609.1"/>
        <property type="gene ID" value="ENSGAGG00000001291.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000139">
        <property type="term" value="C:Golgi membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008375">
        <property type="term" value="F:acetylglucosaminyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003406">
        <property type="entry name" value="Glyco_trans_14"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19297:SF178">
        <property type="entry name" value="BETA-1,3-GALACTOSYL-O-GLYCOSYL-GLYCOPROTEIN BETA-1,6-N-ACETYLGLUCOSAMINYLTRANSFERASE 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19297">
        <property type="entry name" value="GLYCOSYLTRANSFERASE 14 FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02485">
        <property type="entry name" value="Branch"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="3" id="KW-0328">Glycosyltransferase</keyword>
    <keyword evidence="8 11" id="KW-0472">Membrane</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0735">Signal-anchor</keyword>
    <keyword evidence="4" id="KW-0808">Transferase</keyword>
    <keyword evidence="5 11" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 11" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="6"/>
            <end position="27"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004323"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004922"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022676"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00038150"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001609.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="400" mass="46608" checksum="AB0B1254BCAF668F" modified="2019-05-08" version="1">MNQLDATKSGFLVCIGLCIFILTFIYVRNTFSKVPSEQKSFPDTAECGVYPDELCSALFEGKSAAPQIGNWCQIIHQPEMPDCIRTPCNCSTVLKSLHFITRPLSEEEGNFSLAYIITIHKELEMFVKLLRAIYLPQNVYCIHIDEKSPQDYKTAVQNLVNCFENIFISSKRENVVYAGFSRLQADINCMKDLVNAKTQWNYVINLCGQDYPIKTNKEIIQYIKSKWNDKNITPGVVQPPHMKHRTHLSYKEYVHSGTSYVYPTKNMKDDPPHNLTIYFGSAYYVLTRKFVEFTLTDARAKDLLEWSRDTYSPDEHYWVTLNRLNGHYIRDICVYGLGDLQWIIESPNLFANKFEPTTYPLVMDCLERRYRLKVLQQAEVPLEAHWHFQESNYFNMKLNV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIW2</accession>
    <name>A0A452GIW2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Topoisomerase 6 subunit A/Spo11 TOPRIM domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>ATP-dependent breakage, passage and rejoining of double-stranded DNA.</text>
            <dbReference type="EC" id="5.6.2.2"/>
        </reaction>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the TOP6A family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIW2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001755.1">
        <property type="protein sequence ID" value="ENSGAGP00000001539.1"/>
        <property type="gene ID" value="ENSGAGG00000001218.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000228">
        <property type="term" value="C:nuclear chromosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003918">
        <property type="term" value="F:DNA topoisomerase type II (double strand cut, ATP-hydrolyzing) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042138">
        <property type="term" value="P:meiotic DNA double-strand break formation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000706">
        <property type="term" value="P:meiotic DNA double-strand break processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007131">
        <property type="term" value="P:reciprocal meiotic recombination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00223">
        <property type="entry name" value="TOPRIM_TopoIIB_SPO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.1360.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002815">
        <property type="entry name" value="Spo11/TopoVI_A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036078">
        <property type="entry name" value="Spo11/TopoVI_A_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034136">
        <property type="entry name" value="TOPRIM_Topo6A/Spo11"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10848">
        <property type="entry name" value="MEIOTIC RECOMBINATION PROTEIN SPO11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10848:SF0">
        <property type="entry name" value="MEIOTIC RECOMBINATION PROTEIN SPO11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21180">
        <property type="entry name" value="TOP6A-Spo11_Toprim"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01550">
        <property type="entry name" value="TOP6AFAMILY"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56726">
        <property type="entry name" value="DNA topoisomerase IV, alpha subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS52041">
        <property type="entry name" value="TOPO_IIB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="1" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="1" id="KW-0413">Isomerase</keyword>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0799">Topoisomerase</keyword>
    <feature type="domain" description="Topoisomerase 6 subunit A/Spo11 TOPRIM" evidence="2">
        <location>
            <begin position="85"/>
            <end position="227"/>
        </location>
    </feature>
    <feature type="active site" description="O-(5'-phospho-DNA)-tyrosine intermediate" evidence="1">
        <location>
            <position position="53"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01385"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF21180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001539.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="233" mass="26695" checksum="26757D6FCA0E7056" modified="2019-05-08" version="1">MLVPRAETSDQVAFQDVYGSCWIGRKALMLKILSVIYKMVQNNTYATKRDIYYSDTLLFGSQSVVDNIVNDISCMLKIPRRSLHIGKGVPDLNTRLLVRKLWDTCHIPIFTLMDADPHGIEIMCIYKYGSVSMSFEAHHLTVPAIRWLGLLPSDIERLNIRKDALIPLTKQDQNKLTSMQKRPYMACQPMWKKEMEIMAASKMKAEIQALTSLSSDYLSRVYLPSKLQFGGWI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIW4</accession>
    <name>A0A452GIW4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">TFIIS N-terminal domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="9">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="8">Belongs to the IWS1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIW4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001768.1">
        <property type="protein sequence ID" value="ENSGAGP00000001550.1"/>
        <property type="gene ID" value="ENSGAGG00000001253.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006397">
        <property type="term" value="P:mRNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016973">
        <property type="term" value="P:poly(A)+ mRNA export from nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008380">
        <property type="term" value="P:RNA splicing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.930.10:FF:000001">
        <property type="entry name" value="IWS1, SUPT6H interacting protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.930.10">
        <property type="entry name" value="Conserved domain common to transcription factors TFIIS, elongin A, CRSP70"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051037">
        <property type="entry name" value="RNAPII_TF_IWS1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035441">
        <property type="entry name" value="TFIIS/LEDGF_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017923">
        <property type="entry name" value="TFIIS_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46010">
        <property type="entry name" value="PROTEIN IWS1 HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46010:SF1">
        <property type="entry name" value="PROTEIN IWS1 HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08711">
        <property type="entry name" value="Med26"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47676">
        <property type="entry name" value="Conserved domain common to transcription factors TFIIS, elongin A, CRSP70"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51319">
        <property type="entry name" value="TFIIS_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0507">mRNA processing</keyword>
    <keyword evidence="6" id="KW-0508">mRNA splicing</keyword>
    <keyword evidence="3" id="KW-0509">mRNA transport</keyword>
    <keyword evidence="7 9" id="KW-0539">Nucleus</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0804">Transcription</keyword>
    <keyword evidence="4" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="1" id="KW-0813">Transport</keyword>
    <feature type="domain" description="TFIIS N-terminal" evidence="11">
        <location>
            <begin position="577"/>
            <end position="655"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1"/>
            <end position="481"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="659"/>
            <end position="693"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="34"/>
            <end position="51"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="66"/>
            <end position="82"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="99"/>
            <end position="134"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="151"/>
            <end position="211"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="243"/>
            <end position="292"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="321"/>
            <end position="330"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="408"/>
            <end position="421"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="10">
        <location>
            <begin position="442"/>
            <end position="463"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="662"/>
            <end position="673"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022664"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022816"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023187"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00037992"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00649"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS51319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001550.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="781" mass="87628" checksum="7E0A54FCA4B73BFA" modified="2019-05-08" version="1">MEAEYYGGDQSDDGGATPVQDERDSGSDVEDDVNDQHSGSDNESLGHHSENEQSDGEDGGQTRGHHMTDSENEDTPRQKDSDSENEDPPNHNASDSENEGAHGDKDSDSDIEDHPIHHISDSENEDALNHHASDSENEEPQKVPNSDSENEDHRKNLHSESENEEHQKGHASDSENEEPTKHVASDSENEEPLKHAASDSENEDLSKHAASDSENEEPPKQAASDSENEVPPKQVASDSENEEPLKHAASDSENEEPQKVHASDSENEEPQKHPASDSEIEDVPRRKQKIESDDSDGGNGKEAMQNDSRHSDSEQAGEGFHVSDSEEEGPKRRKITDSDEDEERDEDKAVKRKAAVFSDSEDDEKTQAAKKGRVVSDVEESDSDASEKSDKRKKNAVASDSEEEEESKDSVGKKKEEKDLFGSDSESGNEQENLIADIFGESGDEEEEEFTGFNQEDLEEEKTEAEMKETADDSDSDDNIKRGKHMDFMSDFEMMLQRKKSISGKRRRNRDGGTFISDADDVVSAMIVKMNEAAEEDRQLNTQKKPALKKLTLLPTVVMHLKKQDLKETFIDSGVMSAIKEWLSPLPDRSLPALKIREELLKILQELPSVSQETLKHSGIGRAVMYLYKHPKESRPNKDMAGKLINEWSRPIFGLTSNYKGMTREEREQRDLEQMPQRRRLSSCGGQTPRRDLEKVLTGEEKALRPGDPGFCARARVPMPSNKDYVVRPKWNVEMESSRSGSVKKGISRLEKHKRRFAEQKRLSKVHQAIKFSIEGNRMPL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIW6</accession>
    <name>A0A452GIW6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Zinc finger protein 423</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="19">Olf1/EBF-associated zinc finger protein</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="20">Smad- and Olf-interacting zinc finger protein</fullName>
        </alternativeName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="17">Transcription factor that can both act as an activator or a repressor depending on the context. Plays a central role in BMP signaling and olfactory neurogenesis. Associates with SMADs in response to BMP2 leading to activate transcription of BMP target genes. Acts as a transcriptional repressor via its interaction with EBF1, a transcription factor involved in terminal olfactory receptor neurons differentiation; this interaction preventing EBF1 to bind DNA and activate olfactory-specific genes. Involved in olfactory neurogenesis by participating in a developmental switch that regulates the transition from differentiation to maturation in olfactory receptor neurons. Controls proliferation and differentiation of neural precursors in cerebellar vermis formation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIW6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001834.1">
        <property type="protein sequence ID" value="ENSGAGP00000001614.1"/>
        <property type="gene ID" value="ENSGAGG00000001293.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043565">
        <property type="term" value="F:sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045892">
        <property type="term" value="P:negative regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000468">
        <property type="entry name" value="B-cell lymphoma 6 protein-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000244">
        <property type="entry name" value="zinc finger protein 423"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000469">
        <property type="entry name" value="Zinc finger protein 423"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000483">
        <property type="entry name" value="Zinc finger protein 423"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000548">
        <property type="entry name" value="Zinc finger protein 423"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000760">
        <property type="entry name" value="Zinc finger protein 423"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001417">
        <property type="entry name" value="zinc finger protein 423"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002384">
        <property type="entry name" value="zinc finger protein 423"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:004063">
        <property type="entry name" value="Zinc finger protein 423"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000143">
        <property type="entry name" value="Zinc finger protein 521"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000167">
        <property type="entry name" value="Zinc finger protein 521"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000261">
        <property type="entry name" value="Zinc finger protein 521"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050331">
        <property type="entry name" value="Zinc_finger"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16515:SF66">
        <property type="entry name" value="C2H2-TYPE DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16515">
        <property type="entry name" value="PR DOMAIN ZINC FINGER PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13912">
        <property type="entry name" value="zf-C2H2_6"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="30"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="26"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="24"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="14" id="KW-0010">Activator</keyword>
    <keyword evidence="3" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="9" id="KW-0221">Differentiation</keyword>
    <keyword evidence="13" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="6" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="16" id="KW-0539">Nucleus</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0678">Repressor</keyword>
    <keyword evidence="15" id="KW-0804">Transcription</keyword>
    <keyword evidence="12" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="10" id="KW-0862">Zinc</keyword>
    <keyword evidence="8 21" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="7"/>
            <end position="29"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="78"/>
            <end position="105"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="106"/>
            <end position="133"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="134"/>
            <end position="161"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="162"/>
            <end position="189"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="203"/>
            <end position="231"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="235"/>
            <end position="263"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="424"/>
            <end position="452"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="461"/>
            <end position="484"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="576"/>
            <end position="598"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="606"/>
            <end position="628"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="636"/>
            <end position="660"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="664"/>
            <end position="692"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="694"/>
            <end position="722"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="725"/>
            <end position="752"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="751"/>
            <end position="774"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="829"/>
            <end position="856"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="873"/>
            <end position="900"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="902"/>
            <end position="929"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="1063"/>
            <end position="1091"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="1111"/>
            <end position="1138"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="1141"/>
            <end position="1168"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="1172"/>
            <end position="1200"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="1202"/>
            <end position="1227"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="29"/>
            <end position="55"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="283"/>
            <end position="343"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="534"/>
            <end position="562"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="1085"/>
            <end position="1106"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="22">
        <location>
            <begin position="42"/>
            <end position="55"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="22">
        <location>
            <begin position="303"/>
            <end position="317"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="22">
        <location>
            <begin position="548"/>
            <end position="559"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="22">
        <location>
            <begin position="1091"/>
            <end position="1104"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022491"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00055444"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00067811"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00076623"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00080204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001614.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1227" mass="139219" checksum="A9C0D24F306DB8D7" modified="2019-05-08" version="1">MEDESIYTCDNCQQDFDSLADLTEHRAHHCPGDGDDDPQLSWVASSPSSKDVASPTQMIGDGCDLGIGEEEGGTGLPYPCQFCDKSFIRLSYLKRHEQIHSDKLPFKCTYCSRLFKHKRSRDRHIKLHTGDKKYHCHECEAAFSRSDHLKIHLKTHSSSKPFKCTVCKRGFSSTSSLQSHMQAHKKNKEHMAKTEKEVKKDDFMCDYCEETFSQTEDLEKHVMTRHPQLSEKADLQCIHCPEVFADENSLLSHIHQAHANKKHKCPMCPEQFSSVEEVYCHLDSHRQPDSSNHSVSPDPVLGSVASMSSATPDSSASVERGSTPDSTLKPLRGQKKIRSAEREEGQNWSKVTYSCPYCSKRDFNSLAVLEIHLKTIHADKPQQSHTCQICLDSMPTLYNLNEHVRKVHKNHAYPMMQFNNISAFHCNYCPEMFADINSLQEHIRITHCGPSATPQDGNNAFFCNQCSMGFLTEASLTEHIQQTHCNVGNSKLDSPVVQPTQSFMEVYSCPYCTNSPIFGSILKLTKHIKENHKNIPLAHNKKSKAEQSPVSSDVEVSSPKRQRLSASLNSVSNGEYPCNQCDLKFSNFESFQTHLKLHLELLLRKQSCPQCKEDFDSQESLLQHLTVHYMTTSTHYVCESCDKQFSSVDDLQKHLLDMHTFVLYHCTLCQEVFDSKVSIQVHLAVKHSNEKKMYRCTACNWDFRKEVDLQIHVKHSHLGNPTKSHKCIFCGETFSTEVELQCHITTHSKKYNCKFCSKAFHAIILLEKHLREKHCVFDAGAENGTANGMTPTNKKTEAADLQNMLMKNPDTSNSHEASEDDVDASEPMYGCDICGAAYTMEVLLQNHRLRDHNIRPGEDDCSRKKAEFIKGSHKCNICSRTFFSENGLREHMQTHRGPAKHYMCPICGERFPSLLTLTEHKVTHSKSLDTGTCRICKMPLQSEEEFIEHCQMHPDLRNSLTGFRCVVCMQTVTSTLELKIHGTFHMQKLAGNSAASSPNGQALQKLYKCALCLKEFRNKQDLVKLDVNGLPYGLCAGCMTRGTNGQSSGLTPQEVNERPCGSLRCPECSVKFESAEDLENHIQLDHRDLTPETSGQRKGTQTSPVPRKKTYQCIKCQMTFENEREIQIHVANHMIEEGINHECKLCNQMFDSPAKLLCHLIEHSFEGMGGTFKCPVCFTVFVQANKLQQHIFAVHGQEDKIYDCSQCPQKFFFQTELQNHTLSQHAQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIW8</accession>
    <name>A0A452GIW8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="22">Dynein axonemal heavy chain 11</fullName>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Cilium axoneme</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the dynein heavy chain family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000001760.1">
        <property type="protein sequence ID" value="ENSGAGP00000001544.1"/>
        <property type="gene ID" value="ENSGAGG00000000806.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005930">
        <property type="term" value="C:axoneme"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030286">
        <property type="term" value="C:dynein complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005874">
        <property type="term" value="C:microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031514">
        <property type="term" value="C:motile cilium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045505">
        <property type="term" value="F:dynein intermediate chain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051959">
        <property type="term" value="F:dynein light intermediate chain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007018">
        <property type="term" value="P:microtubule-based movement"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000667">
        <property type="entry name" value="Dynein axonemal heavy chain 11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.2620:FF:000004">
        <property type="entry name" value="Dynein axonemal heavy chain 17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.920.30:FF:000003">
        <property type="entry name" value="Dynein axonemal heavy chain 17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000219">
        <property type="entry name" value="Dynein axonemal heavy chain 17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.20.180.20:FF:000001">
        <property type="entry name" value="Dynein axonemal heavy chain 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.8.710:FF:000002">
        <property type="entry name" value="dynein heavy chain 17, axonemal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.140.100:FF:000001">
        <property type="entry name" value="dynein heavy chain 17, axonemal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.1120:FF:000002">
        <property type="entry name" value="Dynein heavy chain 9, axonemal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.472.130:FF:000001">
        <property type="entry name" value="Dynein, axonemal, heavy chain 9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.2620">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.472.130">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.8.710">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.1120">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.920.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.140.100">
        <property type="entry name" value="Dynein heavy chain, N-terminal domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.20.180.20">
        <property type="entry name" value="Dynein heavy chain, N-terminal domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035699">
        <property type="entry name" value="AAA_6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026983">
        <property type="entry name" value="DHC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041589">
        <property type="entry name" value="DNAH3_AAA_lid_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042222">
        <property type="entry name" value="Dynein_2_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043157">
        <property type="entry name" value="Dynein_AAA1S"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041466">
        <property type="entry name" value="Dynein_AAA5_ext"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013602">
        <property type="entry name" value="Dynein_heavy_linker"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013594">
        <property type="entry name" value="Dynein_heavy_tail"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042228">
        <property type="entry name" value="Dynein_linker_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45703:SF12">
        <property type="entry name" value="DYNEIN AXONEMAL HEAVY CHAIN 11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45703">
        <property type="entry name" value="DYNEIN HEAVY CHAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12774">
        <property type="entry name" value="AAA_6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12775">
        <property type="entry name" value="AAA_7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17857">
        <property type="entry name" value="AAA_lid_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08385">
        <property type="entry name" value="DHC_N1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08393">
        <property type="entry name" value="DHC_N2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17852">
        <property type="entry name" value="Dynein_AAA_lid"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="13" id="KW-0966">Cell projection</keyword>
    <keyword evidence="10" id="KW-0969">Cilium</keyword>
    <keyword evidence="9 14" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="12" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="8" id="KW-0243">Dynein</keyword>
    <keyword evidence="4" id="KW-0493">Microtubule</keyword>
    <keyword evidence="11" id="KW-0505">Motor protein</keyword>
    <keyword evidence="6" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="Dynein heavy chain tail" evidence="15">
        <location>
            <begin position="26"/>
            <end position="521"/>
        </location>
    </feature>
    <feature type="domain" description="Dynein heavy chain linker" evidence="16">
        <location>
            <begin position="1025"/>
            <end position="1430"/>
        </location>
    </feature>
    <feature type="domain" description="Dynein heavy chain hydrolytic ATP-binding dynein motor region" evidence="17">
        <location>
            <begin position="1564"/>
            <end position="1890"/>
        </location>
    </feature>
    <feature type="domain" description="Dynein heavy chain AAA 5 extension" evidence="18">
        <location>
            <begin position="2048"/>
            <end position="2166"/>
        </location>
    </feature>
    <feature type="domain" description="Dynein heavy chain 3 AAA+ lid" evidence="19">
        <location>
            <begin position="2383"/>
            <end position="2479"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="14">
        <location>
            <begin position="991"/>
            <end position="1018"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004430"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008887"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023017"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023069"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="Pfam" id="PF08385"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF08393"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF12774"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF17852"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="Pfam" id="PF17857"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001544.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="22">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="2592" mass="301466" checksum="3EC7BFC34AF2C3B8" modified="2019-05-08" version="1">VCSLKQGNVLQALGLTEGGCPSWLLRFLQFYHFLFTALMEAQDIELYLRPLRRHIQFLQETEFPKIHSLIPPLFHTICLIWSHSQFYSVPARIIVLLQEFCNLFIDQARSYLSPEDLLKGEIEETLEHVQIAVNTLRSFKNFFFSHREKLASYFTNSKEFKPWDFQSNMVFARFDLFLNRLVKIEDIFVVMLEFQKLEKLEFGGVKGKTLSEQIYRMNEEFIESCKVFKEKTYDPSDYNNVEFEEDYVEFKAKILDFDRRLGSILCVGFLDCSGLESAFKLLAIFGYFLEKPVIMEIFSPNYIILLQMFDEELDNCKHLYDEHIRQVEKGNEIIINKNMPFTSGNLKWSKALQERIQTFWSNFLYLILQDPETALVYQKYIETLTLIDQFEEQIYCKWKSQVDEVYQFNLDQPLIKRNPENGLLSVNFDSKLVALLREVTYLLKLNRSDIPDSALAIYEKRKIFAKYIGNLELIVQWYNKLKQTVLEVEYPLIETELEAIDGQLRETEEVFTWQKENCWKYIEQVKAAVYDLEQRVQQSKENIKTIQQIMNAWAEHTLFSRKDDRKDALLSLEDKEDRLAKKYKLLKEDGYKIHQLAEENLKLFKADSSSDSWKIYIEYIDDIIVDGFFNTLMHNLDFFLENTEKGLKPAPLFQAQMILSAPEIEFKPSLDKEAGDGFYDLVEELLSNVFKMSAQVKRVAAHLEVENYQSDMDNMLDLSEIRQEIMDRVADVIRKALDYRNSLDTYAYLWVDDRAEFMKQFLLYGHVLTSEEIDSHADEGAPEQPPTTEQFKEQIDIYENLYIQMSKFEDFRTFESWFKVDMKPFKMSLLNIIKKWSWMFKEHLLKFVIGSLTDLEEFIKVTDAGLQKEVNEGDYNGLVEIMGHLLAVRDRQASTDELFEPLKETITLLESYGQKMSDQVYVQLEELPEKWNTTKKIAASIKHEVAPLQTSEVTLIRKKFALFDGKQIEFRERFRVEAPFCFDAKYPYILLDKENRELEALEEEMLQMQKSANLFEVVVPDYKQLKQCRKEIRLLKEVWDINIYVTSSIGDWTKTQWRQINVEQMDAELRRFAKEVWSLDKEIRSWDVYIGLELTVKNLMTSLRAVTELQNPAIRERHWHQLMNATDVRLSITEGTTLADLLGLQLHKVEDEVRSIVDKAVKELGTEKILTEISQTWATVEFCYEEHYRTRTPLLKCDEQLFETLENNQVQLQTILQSKYVEYFIEQVTSWQRTLNVADSVIFIWMEIQRTWSHLESIFIGSDDIRTKLPKDAKRFDGVDADFKELMMETANTKNVLEATTKPNLYEKLESLQRRLSLCEKALAEYLETKCLAFPRFYFVSSADLLDILSKGTQPRQVTRHLTKLFDNIADLQFQDNKEKSTDIALGMYSKEKEYVPFHAECECIGQVETWLQQLEETMYKTVHRHIMEAIAAYEEKPREQWIFDYPAQVALTSSQIWWTTDVGIAFNRLEEGFETALKDYNKKQIAQLNALIALLLGELASGDRQKIMTICTIDVHARDVVAKLVAQKVTSSQAFAWLSQLRHRWDEAQKHCLANICDAQFQYFYEYLGNTSRLVITPLTDRCYITLTQSLHLTMSGAPAGPTGTGKTETTKDLGHALGMMVYVFNCSEQMDYKSIGNIYKGLVQTGAWGCFDEFNRISVEVLSVVAVQVKTIHDAIRNKRKKFLFLGENITLKTSVGIFITMNPGYAGRTELPENLKALFRPCAMVVPDIELICEIMLVAEGFTDARLLARKFITLYTLCRELLSKQDHYDWGLRAVKSVLVVAGSLKRGDKNRPEDQVLMRTLRDFNLPKIVTDDIPIFMGLISDLFPALDVPRRRDLQFEQMVKQSTLELHLQPEESFIFKVVQLEELLAVRHSVFVVGNAGTGKSKVLRTLNRTYVNMKQKLVWNDLNPKAVTTDELFGFIHHATREWKDGLFSSLLREQANITQDGPKWIVLDGDIDPMWIESLNTVMDDNKVLTLASNERVPLTPSMRLLFEIHHLKTATPATVSRAGILYVNPQDLGWNPYVASWIETRSHQSEKANLTILFDKYVPPCLDQLRTSFKTITPIPENSMVQTLCSLLDCLLTPENVPPDCPRELYEIYFVFACVWAFGGALFQDQLSNYQAEFSRWWLKEMKAVKFPSQGTIFDYYLDPKTRKFLPWTDKVPRFDMDPETPLQAVLVHTSETTRLRYFTDLLLERGKPVMLVGNAGVGKTVFVGDRLAAFPDDYMVSNVLFNYYTTSSVLQKILEKSLEKKAGRNYGPVGNKKLVYFIDDMNMPEMDVYGTVQPHTLIRQHIDYGHWYDRQKLTVKEIHNCHYVACMNPTAGNFTINPRLQRHFTVFALNFPSVDALNTIYSKILGFHFQRHVFNPSVLKNAPAMIQASIWLHQTMVQNFLPTAVKFHYIFNLRDLSNIFQGILFATPECLREPNDLVCLWLHESSRVYGDKLVEPKDCHFFRKKLIDTAHKYFEGVEDHLLRQHPLIYCHFANGVASPHYMPVKEWEVLRKILTETLESYNELNASMNLVLFEDAMQHGSPIQKLRTTELTPLEPLGMQGRGVSLGSVGKEVGGVGYCSSHVIPAPWLSWLYHC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIX0</accession>
    <name>A0A452GIX0_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="7">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the formin homology family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIX0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001835.1">
        <property type="protein sequence ID" value="ENSGAGP00000001615.1"/>
        <property type="gene ID" value="ENSGAGG00000001295.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031267">
        <property type="term" value="F:small GTPase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030866">
        <property type="term" value="P:cortical actin cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008360">
        <property type="term" value="P:regulation of cell shape"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="entry name" value="Formin-like 1, isoform CRA_c"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000036">
        <property type="entry name" value="Formin-like protein 3 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.2220">
        <property type="entry name" value="Formin, FH2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014767">
        <property type="entry name" value="DAD_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015425">
        <property type="entry name" value="FH2_Formin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042201">
        <property type="entry name" value="FH2_Formin_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010472">
        <property type="entry name" value="FH3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043592">
        <property type="entry name" value="FMNL_animal"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014768">
        <property type="entry name" value="GBD/FH3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010473">
        <property type="entry name" value="GTPase-bd"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45857">
        <property type="entry name" value="FORMIN-LIKE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45857:SF5">
        <property type="entry name" value="FORMIN-LIKE PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06367">
        <property type="entry name" value="Drf_FH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06371">
        <property type="entry name" value="Drf_GBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02181">
        <property type="entry name" value="FH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01139">
        <property type="entry name" value="Drf_FH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01140">
        <property type="entry name" value="Drf_GBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00498">
        <property type="entry name" value="FH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF101447">
        <property type="entry name" value="Formin homology 2 domain (FH2 domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51231">
        <property type="entry name" value="DAD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51444">
        <property type="entry name" value="FH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51232">
        <property type="entry name" value="GBD_FH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="GBD/FH3" evidence="5">
        <location>
            <begin position="1"/>
            <end position="409"/>
        </location>
    </feature>
    <feature type="domain" description="FH2" evidence="6">
        <location>
            <begin position="556"/>
            <end position="947"/>
        </location>
    </feature>
    <feature type="domain" description="DAD" evidence="4">
        <location>
            <begin position="979"/>
            <end position="1016"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="462"/>
            <end position="534"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="2">
        <location>
            <begin position="321"/>
            <end position="418"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="2">
        <location>
            <begin position="836"/>
            <end position="890"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="2">
        <location>
            <begin position="921"/>
            <end position="952"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="3">
        <location>
            <begin position="465"/>
            <end position="478"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="3">
        <location>
            <begin position="491"/>
            <end position="508"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="3">
        <location>
            <begin position="515"/>
            <end position="531"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00023449"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS51231"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS51232"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS51444"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001615.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1026" mass="116727" checksum="A3F3EF9DEE6DAA0A" modified="2019-05-08" version="1">MIMCSLQERFQVKNPPHTYIQKLKGYLDPAVTRKKFRRRVQESTQVLRELEISLRTNHIGWVREFLNEENKGLDVLVEYLSFAQYAVTFDFESLENNVENSMDKSKPWSRSIEDLHRGSNLPSPVGNSISRSGRHSTLRYNTLPSRRTLKNSRLVSKKDDVHVCIMCLRAIMNYQYGFNMVMSHPHAVNEIALSLNNKNPRTKALVLELLAAVCLVRGGHEIILSAFDNFKEVCGEKQRFEKLMEHFRNEDNNIDFMVACMQFINIVVHSVEDMNFRVHLQYEFTKLGLDEYLDKLKHTESDKLQVQIQAYLDNVFDVGALLEDAETKNAALERVEELEENISHLSEKLQDTENEAMAKIVELEKQLMQRNKELDVVREIYKDANTQVHTLRKMVKEKEEAIQRQSTLEKKIHELEKQGTIKIQKKGDGDISILPVAVASGTLPSGSEATAGTCVGPVTGAMPSVPLPPPPPPLPPLSPTSETVQNGPVTAPMPPPPPPPPPPPPLPGPAAETAPAPPLPPPPPPSAPPLPGTASPTVIFNSGLADGPIKLFSVKIKKPIKTKFRMPVFNWVALKPNQINGTVFNEIDDERILEDLNVDEFEEIFKTKAQGPAIDLTSSKQKITQKGSNKVTLLEANRAKNLAITLRKAGKTADEICKAIQVFDLKTLPVDFVECLMRFLPTENEVKVLRLYERERKPLENLTDEDRFMMQFSKIERLMQKMTIMAFIGNFTESIQMLTPQLHAVIAASVSIKSSQKLKKILEIILALGNYMNSSKRGAVYGFKLQSLDLLLDTKSTDRKQTLLHYISNVVKEKYHQVSLFYNELHYVEKAAAVSLENVLLDVKELQRGLDLTKREYTMHDHNTMLKEFIHNNEGKLKKLQDDAKIAQDAFDDAVKYFGENPKTTPPSVFFPVFVRFVKAYKQAEEENELRKKQEQALMEKLLEQEALMEQQDQKSPSHKTKRQQQELIAELRRRQVKDNRHVYEGKDGAIEDIITVLKTVPFTARTAKRGSRFFCEPVLTEEFHY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIX2</accession>
    <name>A0A452GIX2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Transmembrane protein 255A</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="7">Protein FAM70A</fullName>
        </alternativeName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the TMEM255 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIX2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001766.1">
        <property type="protein sequence ID" value="ENSGAGP00000001549.1"/>
        <property type="gene ID" value="ENSGAGG00000001248.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028014">
        <property type="entry name" value="TMEM255"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR33721:SF1">
        <property type="entry name" value="TRANSMEMBRANE PROTEIN 255A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR33721">
        <property type="entry name" value="TRANSMEMBRANE PROTEIN 255B-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14967">
        <property type="entry name" value="FAM70"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 9" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 9" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 9" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="30"/>
            <end position="50"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="62"/>
            <end position="82"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="89"/>
            <end position="115"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="174"/>
            <end position="199"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="258"/>
            <end position="279"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="264"/>
            <end position="279"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007903"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00039451"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00041817"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001549.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="302" mass="33055" checksum="87AA66419E0CB896" modified="2019-05-08" version="1">MRQSLTQQRSSAVPLPDSAGSFNRRKRNSLYVTVTLLIVSVLILTVGLAATTRTQNVTVGGYYPGVILGFGSFLGIIGSNLIENKRQMLVASIVFISFGVIAAFCCAIVDGVFAARHIGICNPQRRGPCTPKIKTNTCFCCDLYNCGNRVEISGGYYEYIDVSSCQDIIHLYHLLWSATILNIVGLFLGIITAAILGGFKDMTPSLPTLNCTVENAHPSVSYYSRPQVTSYNTYYHSTPHLPPYSAYDFQHSSVFPASTPSGLSDDPQSMSPSPSYMWSSNAPPRYSPPYFPPFEKPPPYTP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIX4</accession>
    <name>A0A452GIX4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Gastrin-releasing peptide</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasmic vesicle</location>
            <location evidence="1">Secretory vesicle lumen</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the bombesin/neuromedin-B/ranatensin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIX4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001828.1">
        <property type="protein sequence ID" value="ENSGAGP00000001608.1"/>
        <property type="gene ID" value="ENSGAGG00000001289.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031410">
        <property type="term" value="C:cytoplasmic vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005184">
        <property type="term" value="F:neuropeptide hormone activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007218">
        <property type="term" value="P:neuropeptide signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000874">
        <property type="entry name" value="Bombesin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16866">
        <property type="entry name" value="GASTRIN-RELEASING PEPTIDE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16866:SF2">
        <property type="entry name" value="GASTRIN-RELEASING PEPTIDE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02044">
        <property type="entry name" value="Bombesin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00257">
        <property type="entry name" value="BOMBESIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0027">Amidation</keyword>
    <keyword evidence="6" id="KW-0165">Cleavage on pair of basic residues</keyword>
    <keyword evidence="9" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0964">Secreted</keyword>
    <keyword evidence="7 10" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="10">
        <location>
            <begin position="1"/>
            <end position="28"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019272643" description="Gastrin-releasing peptide" evidence="10">
        <location>
            <begin position="29"/>
            <end position="153"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004263"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00010012"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00016270"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022685"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022815"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001608.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="153" mass="17431" checksum="51B4229E016D930F" modified="2019-05-08" version="1">MRGGEMPRSLLSLGLFALVLLEVHWGAAAPLQANGGASPLAKIYPRGSHWAVGHLMGKKSTGDFPYVYEEENKIPFSALPENVKQLGEYLQWEETLKNLLRLLEGNENRSTQILREELPWYTKNTWETDDDSSFKDMMDYLLQVMNRKESTPS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIX5</accession>
    <name>A0A452GIX5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Pumilio homolog 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">P-body</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasmic granule</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIX5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001840.1">
        <property type="protein sequence ID" value="ENSGAGP00000001620.1"/>
        <property type="gene ID" value="ENSGAGG00000001290.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000932">
        <property type="term" value="C:P-body"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003730">
        <property type="term" value="F:mRNA 3'-UTR binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035196">
        <property type="term" value="P:miRNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043488">
        <property type="term" value="P:regulation of mRNA stability"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006417">
        <property type="term" value="P:regulation of translation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd07920">
        <property type="entry name" value="Pumilio"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000004">
        <property type="entry name" value="Pumilio homolog 1 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033133">
        <property type="entry name" value="PUM-HD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033712">
        <property type="entry name" value="Pumilio_RNA-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001313">
        <property type="entry name" value="Pumilio_RNA-bd_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12537:SF1">
        <property type="entry name" value="PUMILIO HOMOLOG 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12537">
        <property type="entry name" value="RNA BINDING PROTEIN PUMILIO-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00806">
        <property type="entry name" value="PUF"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SMART" id="SM00025">
        <property type="entry name" value="Pumilio"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50302">
        <property type="entry name" value="PUM"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50303">
        <property type="entry name" value="PUM_HD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="5" id="KW-0810">Translation regulation</keyword>
    <feature type="domain" description="PUM-HD" evidence="10">
        <location>
            <begin position="730"/>
            <end position="1072"/>
        </location>
    </feature>
    <feature type="repeat" description="Pumilio" evidence="8">
        <location>
            <begin position="750"/>
            <end position="785"/>
        </location>
    </feature>
    <feature type="repeat" description="Pumilio" evidence="8">
        <location>
            <begin position="786"/>
            <end position="821"/>
        </location>
    </feature>
    <feature type="repeat" description="Pumilio" evidence="8">
        <location>
            <begin position="822"/>
            <end position="859"/>
        </location>
    </feature>
    <feature type="repeat" description="Pumilio" evidence="8">
        <location>
            <begin position="860"/>
            <end position="895"/>
        </location>
    </feature>
    <feature type="repeat" description="Pumilio" evidence="8">
        <location>
            <begin position="896"/>
            <end position="931"/>
        </location>
    </feature>
    <feature type="repeat" description="Pumilio" evidence="8">
        <location>
            <begin position="932"/>
            <end position="967"/>
        </location>
    </feature>
    <feature type="repeat" description="Pumilio" evidence="8">
        <location>
            <begin position="968"/>
            <end position="1003"/>
        </location>
    </feature>
    <feature type="repeat" description="Pumilio" evidence="8">
        <location>
            <begin position="1007"/>
            <end position="1046"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="22"/>
            <end position="73"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="233"/>
            <end position="286"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="490"/>
            <end position="525"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="644"/>
            <end position="677"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="45"/>
            <end position="58"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="250"/>
            <end position="277"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="490"/>
            <end position="503"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="512"/>
            <end position="525"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="665"/>
            <end position="677"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004201"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004463"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022845"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00040564"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00317"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001620.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1090" mass="117329" checksum="85B41B572A55D397" modified="2019-05-08" version="1">MSVACVLKRKAVLWQDSFSPHLKQHPQDTANPNMPVVLTPGTGSQAQQQPAANPALAAGTHSSPVPGSIGVAGRSQDDAMVDYFFQRQHGEQLGGGGSGGGGYNNSKHRWPTGDNIHAEHQVRSMDELNHDFQALALEGRAMGEQLLPGKKFWETDDSSKDGPKGIFLGDQWRDSAWGTSDHSVSQPIMVQRRPGQGFHVNSDVNSVLSPRSESGGLGVSMVEYVLSSSPGDSCLRKGGFGPRDAENDENDKSDKKNKGTFDGDKLGDLKEEGDVMDKTNGLPVQNGIDADVKDFSRTPGNCQNSANEVDLLGPNQNGSEGLAQLASTNGAKPVEDFSNMESQSVPLDPMEHVGMEPLQFDYSGTQVPVDSAAATVGLFDYNSQPQLFQRPNALAVQQLTAAQQQQYALAAAHQPHIAGLAPAAFVPNPYIISAAPPGTDPYAAGLAAAATLGPAVVPHQYYGVTPWGVYPASLFQQQAAAAAAASNSANQQTTQQTQQGQQQVLRGGASQRPLTPNQNQQGQQTDPLVAAAAVNSALAFGQGLAAGMPGYPMLAPAAYYDQTGALVVNAGARNGLGAPVRLVAPAPVIISSSAAQAVANSNTGSGSRRDSLTGSSDLYKRTSSSLTPIGHSFYNGLGFSSSPGPVGMPLPSQGPGHSQTPPPSLSSHGSSSSLNLGGLTNGSGRYISAAPGAEAKYRSASSASSLFSPSSTLFPSSRLRYGMSDVMPSGRSRLLEDFRNNRYPNLQLREIAGHIMEFSQDQHGSRFIQLKLERATPAERQLVFNEILQAAYQLMVDVFGNYVIQKFFEFGSLDQKLALAERIRGHVLSLALQMYGCRVIQKALEFIPPDQQVINEMVRELDGHVLKCVKDQNGNHVVQKCIECVQPQSLQFIIDAFKGQVFALSTHPYGCRVIQRILEHCLPEQTLPILEELHQHTEQLVQDQYGNYVIQHVLEHGRPEDKSKIVAEIRGNVLVLSQHKFASNVVEKCVTHASRTERAMLIDEVCTMNDGPHSALYTMMKDQYANYVVQKMIDVAEPAQRKIVMHKIRPHIATLRKYTYGKHILAKLEKYYMKNGVDLGPICGPPNGII</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIX6</accession>
    <name>A0A452GIX6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">RING-type E3 ubiquitin transferase</fullName>
            <ecNumber evidence="5">2.3.2.27</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>S-ubiquitinyl-[E2 ubiquitin-conjugating enzyme]-L-cysteine + [acceptor protein]-L-lysine = [E2 ubiquitin-conjugating enzyme]-L-cysteine + N(6)-ubiquitinyl-[acceptor protein]-L-lysine.</text>
            <dbReference type="EC" id="2.3.2.27"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="3">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the TRIM/RBCC family.</text>
    </comment>
    <dbReference type="EC" id="2.3.2.27" evidence="5"/>
    <dbReference type="AlphaFoldDB" id="A0A452GIX6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001833.1">
        <property type="protein sequence ID" value="ENSGAGP00000001613.1"/>
        <property type="gene ID" value="ENSGAGG00000001264.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061630">
        <property type="term" value="F:ubiquitin protein ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd16598">
        <property type="entry name" value="RING-HC_TRIM26_C-IV"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="SPRY_PRY_TRIM7_like"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Butyrophilin subfamily 1 member A1"/>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
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        <property type="entry name" value="B30.2/SPRY"/>
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        <property type="entry name" value="Butyrophylin_SPRY"/>
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        <property type="entry name" value="ConA-like_dom_sf"/>
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        <property type="entry name" value="PRY"/>
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        <property type="entry name" value="TRIM/RBCC"/>
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        <property type="entry name" value="Znf_B-box"/>
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        <property type="entry name" value="Znf_B-box_chordata"/>
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        <property type="entry name" value="Znf_PARP"/>
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        <property type="entry name" value="Znf_RING"/>
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        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
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        <property type="entry name" value="Znf_RING_CS"/>
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        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE TRIM"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="PRY"/>
        <property type="match status" value="1"/>
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        <property type="match status" value="1"/>
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        <property type="entry name" value="BBOXZNFINGER"/>
    </dbReference>
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        <property type="entry name" value="BUTYPHLNCDUF"/>
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        <property type="entry name" value="B-box zinc-binding domain"/>
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        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
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    <dbReference type="SUPFAM" id="SSF57850">
        <property type="entry name" value="RING/U-box"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="B302_SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="ZF_BBOX"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="ZF_PARP_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00518">
        <property type="entry name" value="ZF_RING_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50089">
        <property type="entry name" value="ZF_RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12 14" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0808">Transferase</keyword>
    <keyword evidence="10" id="KW-0833">Ubl conjugation pathway</keyword>
    <keyword evidence="11" id="KW-0862">Zinc</keyword>
    <keyword evidence="9 13" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="RING-type" evidence="16">
        <location>
            <begin position="16"/>
            <end position="59"/>
        </location>
    </feature>
    <feature type="domain" description="PARP-type" evidence="15">
        <location>
            <begin position="43"/>
            <end position="68"/>
        </location>
    </feature>
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        <location>
            <begin position="81"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="domain" description="B30.2/SPRY" evidence="18">
        <location>
            <begin position="266"/>
            <end position="453"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="14">
        <location>
            <begin position="123"/>
            <end position="196"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00008518"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00012483"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
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        <source>
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        <source>
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    </evidence>
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        <source>
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    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
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        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00024"/>
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        <source>
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    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50064"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="PROSITE" id="PS50089"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="PROSITE" id="PS50119"/>
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        <source>
            <dbReference type="PROSITE" id="PS50188"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001613.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="453" mass="52282" checksum="DAFC6285EE50144B" modified="2019-05-08" version="1">MASSLSGLDMREEATCPICLEYLTDPVSVDCGHNFCRGCVTQYCVEWAGRGSLTCPICRSQILKGNLRPNWQLGNLVEKMKHVAVCTRHKKTLDLFCEEDEEAVCVVCERASQHRSHTVLLIEEAAQKYKEKIQAHLKTLKEEREKQLGLKKQTQTERQKIVSEFQQLRQFLEEQERLLLAQLEKLDKEILRIQKENVTKLSGQISHLSELIHELEGKCQKPASEFLQVRLSLKHSRSMCRDRRAPESCPLESSSQRSQCNCLCALLEWELLFFELQTHKYLGLLHMTLDPDTAHPQLILSEDRKTVRYGRLWNKLPNTTKRFDYERCVLGSEGFTMGRHCWEVEVGERGGWAVGVARESLGRKGEISFKPKGGIWAVEYWEGQYWALTDPLTPLPLSWVPRRIWIYLDYEQGQVTFFNADNESLIFTFPPASFAGERIRPLFAITQENAAIF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIZ1</accession>
    <name>A0A452GIZ1_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="11">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIZ1"/>
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        <property type="protein sequence ID" value="ENSGAGP00000001571.1"/>
        <property type="gene ID" value="ENSGAGG00000001257.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd18331">
        <property type="entry name" value="BTB_POZ_ZBTB27_BCL6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000105">
        <property type="entry name" value="B-cell CLL/lymphoma 6, member B"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000457">
        <property type="entry name" value="B-cell lymphoma 6 protein-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.710.10:FF:000025">
        <property type="entry name" value="B-cell lymphoma 6 protein-like"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="BCL6A, transcription repressor a"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.710.10">
        <property type="entry name" value="Potassium Channel Kv1.1, Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000210">
        <property type="entry name" value="BTB/POZ_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011333">
        <property type="entry name" value="SKP1/BTB/POZ_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
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    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24394:SF36">
        <property type="entry name" value="B-CELL LYMPHOMA 6 PROTEIN ISOFORM X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24394">
        <property type="entry name" value="ZINC FINGER PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="BTB"/>
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        <property type="entry name" value="zf-C2H2"/>
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    </dbReference>
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        <property type="entry name" value="BTB"/>
        <property type="match status" value="1"/>
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    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="4"/>
    </dbReference>
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        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54695">
        <property type="entry name" value="POZ domain"/>
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        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="6" id="KW-0539">Nucleus</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <keyword evidence="4 7" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="BTB" evidence="9">
        <location>
            <begin position="32"/>
            <end position="99"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="10">
        <location>
            <begin position="521"/>
            <end position="548"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="10">
        <location>
            <begin position="549"/>
            <end position="576"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="10">
        <location>
            <begin position="577"/>
            <end position="604"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="10">
        <location>
            <begin position="605"/>
            <end position="628"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="304"/>
            <end position="373"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="395"/>
            <end position="469"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="304"/>
            <end position="318"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="334"/>
            <end position="371"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="403"/>
            <end position="455"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50097"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001571.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="653" mass="72903" checksum="8703D5C183DBFB35" modified="2019-05-08" version="1">MASPADSCIQFTRHASDVLLNLNRLRSRDILTDVVIIVNREQFRAHKTVLMACSGLFYSIFTDQLKCNLNIITLAPDINPEGFCILLDFMYTSRLNLRESNIMAVMATALYLQMEHVVDTCRRFVKSSEAEMVSAMKTPREEFLTGRMLSHPEVMAYRSREVSENNMPLRNTPLCDGRAFASTLYSGLSGSHLSYTGYNPIPVNGFLLDDELREARMPLPDVSRVNPFPKERTLPCDSSRTIPADYTRAVTDLSANTCYATIYSPKEAATEEARSDMHYSIAAGPKPVAPSVRNNPYFACDKVAKEEERTSSEDEISQHFEPTNTPMNRKGLISPQSPQKSDCQPNSPTESSSSKNARIAQSSGSLGTKSPTDPKACNWKKYKFIVLNNLNQSTKQDGADQNEAGTLSPRSYVSTSVCQQSMEPENLDIQSPTKLSLNGEDSTIPQASRLNNIVNRSLDGSPRSSEGQSPLYLHSSKCSSCGSQSPQHSEMCLHTPGSTFGEEMGETHSEYSDSSCGEKPYRCTICGAQFNRPANLKTHTRIHSGEKPYKCETCGARFVQVAHLRAHVLIHTGEKPYPCEICGTRFRHLQTLKSHLRIHTGEKPYHCEKCNLHFRHKSQLRLHLRQKHGAITNTKVQYRVSTTELPPELPKAC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIZ2</accession>
    <name>A0A452GIZ2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">DNA ligase</fullName>
            <ecNumber evidence="15">6.5.1.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">DNA ligase that seals nicks in double-stranded during DNA repair. Also involved in DNA replication and DNA recombination.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13 15">
            <text>ATP + (deoxyribonucleotide)n-3'-hydroxyl + 5'-phospho-(deoxyribonucleotide)m = (deoxyribonucleotide)n+m + AMP + diphosphate.</text>
            <dbReference type="EC" id="6.5.1.1"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 16">Belongs to the ATP-dependent DNA ligase family.</text>
    </comment>
    <dbReference type="EC" id="6.5.1.1" evidence="15"/>
    <dbReference type="AlphaFoldDB" id="A0A452GIZ2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001860.1">
        <property type="protein sequence ID" value="ENSGAGP00000001639.1"/>
        <property type="gene ID" value="ENSGAGG00000000751.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003910">
        <property type="term" value="F:DNA ligase (ATP) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071897">
        <property type="term" value="P:DNA biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006310">
        <property type="term" value="P:DNA recombination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006281">
        <property type="term" value="P:DNA repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903461">
        <property type="term" value="P:Okazaki fragment processing involved in mitotic DNA replication"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07900">
        <property type="entry name" value="Adenylation_DNA_ligase_I_Euk"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd07969">
        <property type="entry name" value="OBF_DNA_ligase_I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.3260.10:FF:000001">
        <property type="entry name" value="DNA ligase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.50.140:FF:000062">
        <property type="entry name" value="DNA ligase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.470.30:FF:000016">
        <property type="entry name" value="DNA ligase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1490.70">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.3260.10">
        <property type="entry name" value="DNA ligase, ATP-dependent, N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.470.30">
        <property type="entry name" value="DNA ligase/mRNA capping enzyme"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.50.140">
        <property type="entry name" value="Nucleic acid-binding proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050191">
        <property type="entry name" value="ATP-dep_DNA_ligase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000977">
        <property type="entry name" value="DNA_ligase_ATP-dep"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012309">
        <property type="entry name" value="DNA_ligase_ATP-dep_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012310">
        <property type="entry name" value="DNA_ligase_ATP-dep_cent"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016059">
        <property type="entry name" value="DNA_ligase_ATP-dep_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012308">
        <property type="entry name" value="DNA_ligase_ATP-dep_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036599">
        <property type="entry name" value="DNA_ligase_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012340">
        <property type="entry name" value="NA-bd_OB-fold"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00574">
        <property type="entry name" value="dnl1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45674:SF4">
        <property type="entry name" value="DNA LIGASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45674">
        <property type="entry name" value="DNA LIGASE 1/3 FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04679">
        <property type="entry name" value="DNA_ligase_A_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01068">
        <property type="entry name" value="DNA_ligase_A_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04675">
        <property type="entry name" value="DNA_ligase_A_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF117018">
        <property type="entry name" value="ATP-dependent DNA ligase DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56091">
        <property type="entry name" value="DNA ligase/mRNA capping enzyme, catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50249">
        <property type="entry name" value="Nucleic acid-binding proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00697">
        <property type="entry name" value="DNA_LIGASE_A1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00333">
        <property type="entry name" value="DNA_LIGASE_A2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50160">
        <property type="entry name" value="DNA_LIGASE_A3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 15" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="12" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="4" id="KW-0132">Cell division</keyword>
    <keyword evidence="7 15" id="KW-0227">DNA damage</keyword>
    <keyword evidence="9 15" id="KW-0233">DNA recombination</keyword>
    <keyword evidence="10 15" id="KW-0234">DNA repair</keyword>
    <keyword evidence="5" id="KW-0235">DNA replication</keyword>
    <keyword evidence="3 15" id="KW-0436">Ligase</keyword>
    <keyword evidence="6 15" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="ATP-dependent DNA ligase family profile" evidence="17">
        <location>
            <begin position="344"/>
            <end position="480"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007572"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022598"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022705"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023172"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00034003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00058910"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="RuleBase" id="RU000617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="RuleBase" id="RU004196"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50160"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001639.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="615" mass="69273" checksum="2E264BB9AE0F5F8F" modified="2019-05-08" version="1">MLKNIETLSNLFRSVVALSPSDILPCIYLCLNRLGPAYKGLELGIGETILMKAVAQAMGRQLDQIKAEAQEKGDLGLVAESSRSNQRTMFTPPKLSAAGVFSKLQEIARMTGSASMNKKIDIIKGLFVACRHSEARYITRSLGGKLRIGLAEQSVLSAIAHAVSLTPPGQGFPPEVLDAGQSKSAEARKAWLEEQTQILKQTFCELPSYDIIVPILLEHGVESLPQHCKITPGIPLKPMLAQPTKGIGEVLKRFEEAAFTCEYKYDGERAQIHILENGDVHIYSRNQEDNTSKYPDIISRIPKVKKDSVQSCILDAEAVAWDPEKRQIQPFQVLTTRKRKDVEAAEIKVQVCLYAFDILYLNGESLVKEPFSRRRQLLHESFVETEGEFLFATSMDTCSLDEISEFLDQSIKDSCEGLMVKTLEVDATYEIAKRSHKWLKLKKDYLEGVGDTLDLVVIGAYLGKGKRVGMYGGFLLACYDEESEEYQSICKIGTGFTEESLEKHYSFLKDHVLEKPRPYYRWDSSTEPNHWLAAVQVWEVKCADLSISPVHKAAIGLVDEEKGISLRFPRFLRVRDDKKPEEATTSSQVAELYRRQQQIQNQQLVEKAEEEEDFY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIZ3</accession>
    <name>A0A452GIZ3_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="6">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the formin homology family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIZ3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001848.1">
        <property type="protein sequence ID" value="ENSGAGP00000001628.1"/>
        <property type="gene ID" value="ENSGAGG00000001295.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031267">
        <property type="term" value="F:small GTPase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030866">
        <property type="term" value="P:cortical actin cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008360">
        <property type="term" value="P:regulation of cell shape"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.2220:FF:000001">
        <property type="entry name" value="Formin-like 1, isoform CRA_c"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000036">
        <property type="entry name" value="Formin-like protein 3 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.2220">
        <property type="entry name" value="Formin, FH2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015425">
        <property type="entry name" value="FH2_Formin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042201">
        <property type="entry name" value="FH2_Formin_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010472">
        <property type="entry name" value="FH3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043592">
        <property type="entry name" value="FMNL_animal"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014768">
        <property type="entry name" value="GBD/FH3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010473">
        <property type="entry name" value="GTPase-bd"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45857">
        <property type="entry name" value="FORMIN-LIKE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45857:SF5">
        <property type="entry name" value="FORMIN-LIKE PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06367">
        <property type="entry name" value="Drf_FH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06371">
        <property type="entry name" value="Drf_GBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02181">
        <property type="entry name" value="FH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01139">
        <property type="entry name" value="Drf_FH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01140">
        <property type="entry name" value="Drf_GBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00498">
        <property type="entry name" value="FH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF101447">
        <property type="entry name" value="Formin homology 2 domain (FH2 domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51444">
        <property type="entry name" value="FH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51232">
        <property type="entry name" value="GBD_FH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="GBD/FH3" evidence="4">
        <location>
            <begin position="1"/>
            <end position="409"/>
        </location>
    </feature>
    <feature type="domain" description="FH2" evidence="5">
        <location>
            <begin position="549"/>
            <end position="940"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="462"/>
            <end position="534"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="2">
        <location>
            <begin position="321"/>
            <end position="418"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="2">
        <location>
            <begin position="829"/>
            <end position="883"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="2">
        <location>
            <begin position="914"/>
            <end position="945"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="3">
        <location>
            <begin position="465"/>
            <end position="478"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="3">
        <location>
            <begin position="491"/>
            <end position="508"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="3">
        <location>
            <begin position="515"/>
            <end position="531"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00023449"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS51232"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS51444"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001628.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1020" mass="115895" checksum="5C7E5BD164CF6637" modified="2019-05-08" version="1">MIMCSLQERFQVKNPPHTYIQKLKGYLDPAVTRKKFRRRVQESTQVLRELEISLRTNHIGWVREFLNEENKGLDVLVEYLSFAQYAVTFDFESLENNVENSMDKSKPWSRSIEDLHRGSNLPSPVGNSISRSGRHSTLRYNTLPSRRTLKNSRLVSKKDDVHVCIMCLRAIMNYQYGFNMVMSHPHAVNEIALSLNNKNPRTKALVLELLAAVCLVRGGHEIILSAFDNFKEVCGEKQRFEKLMEHFRNEDNNIDFMVACMQFINIVVHSVEDMNFRVHLQYEFTKLGLDEYLDKLKHTESDKLQVQIQAYLDNVFDVGALLEDAETKNAALERVEELEENISHLSEKLQDTENEAMAKIVELEKQLMQRNKELDVVREIYKDANTQVHTLRKMVKEKEEAIQRQSTLEKKIHELEKQGTIKIQKKGDGDISILPVAVASGTLPSGSEATAGTCVGPVTGAMPSVPLPPPPPPLPPLSPTSETVQNGPVTAPMPPPPPPPPPPPPLPGPAAETAPAPPLPPPPPPSAPPLPGTASPTVIFNSGLAAVKIKKPIKTKFRMPVFNWVALKPNQINGTVFNEIDDERILEDLNVDEFEEIFKTKAQGPAIDLTSSKQKITQKGSNKVTLLEANRAKNLAITLRKAGKTADEICKAIQVFDLKTLPVDFVECLMRFLPTENEVKVLRLYERERKPLENLTDEDRFMMQFSKIERLMQKMTIMAFIGNFTESIQMLTPQLHAVIAASVSIKSSQKLKKILEIILALGNYMNSSKRGAVYGFKLQSLDLLLDTKSTDRKQTLLHYISNVVKEKYHQVSLFYNELHYVEKAAAVSLENVLLDVKELQRGLDLTKREYTMHDHNTMLKEFIHNNEGKLKKLQDDAKIAQDAFDDAVKYFGENPKTTPPSVFFPVFVRFVKAYKQAEEENELRKKQEQALMEKLLEQEALMEQQDQKSPSHKTKRQQQELIAELRRRQVKDNRHVYEGKDGAIEDIITALKKNNITKFPNVHSRVRISSSTPMVEDTQS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIZ8</accession>
    <name>A0A452GIZ8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">TGF-beta family profile domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 8">Belongs to the TGF-beta family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIZ8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001795.1">
        <property type="protein sequence ID" value="ENSGAGP00000001576.1"/>
        <property type="gene ID" value="ENSGAGG00000001267.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005125">
        <property type="term" value="F:cytokine activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008083">
        <property type="term" value="F:growth factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030509">
        <property type="term" value="P:BMP signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048468">
        <property type="term" value="P:cell development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051094">
        <property type="term" value="P:positive regulation of developmental process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042127">
        <property type="term" value="P:regulation of cell population proliferation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009888">
        <property type="term" value="P:tissue development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.970:FF:000008">
        <property type="entry name" value="Bone morphogenetic protein 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.970">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.90.10">
        <property type="entry name" value="Cystine-knot cytokines"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029034">
        <property type="entry name" value="Cystine-knot_cytokine"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001839">
        <property type="entry name" value="TGF-b_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001111">
        <property type="entry name" value="TGF-b_propeptide"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015615">
        <property type="entry name" value="TGF-beta-rel"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017948">
        <property type="entry name" value="TGFb_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11848:SF135">
        <property type="entry name" value="BONE MORPHOGENETIC PROTEIN 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11848">
        <property type="entry name" value="TGF-BETA FAMILY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00019">
        <property type="entry name" value="TGF_beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00688">
        <property type="entry name" value="TGFb_propeptide"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00669">
        <property type="entry name" value="INHIBINA"/>
    </dbReference>
    <dbReference type="SMART" id="SM00204">
        <property type="entry name" value="TGFB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57501">
        <property type="entry name" value="Cystine-knot cytokines"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00250">
        <property type="entry name" value="TGF_BETA_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51362">
        <property type="entry name" value="TGF_BETA_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="5" id="KW-0221">Differentiation</keyword>
    <keyword evidence="7" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6 8" id="KW-0339">Growth factor</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <feature type="domain" description="TGF-beta family profile" evidence="10">
        <location>
            <begin position="317"/>
            <end position="387"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="296"/>
            <end position="315"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006656"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="RuleBase" id="RU000354"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS51362"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001576.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="407" mass="46839" checksum="F161ECFE7E8CB4DD" modified="2019-05-08" version="1">MHVVWLRQPASWLLVLWAAAPFLWLRCILADFTLDNEVHSSFIHRKLRSQERREIQREILSILGLPHRPRPHLHGKQNSAPMFMLDLYNAMSVEEEAAGAAGEEGEGFSYPYKPIFSTQGPPLASLQDSNFLTEADMVMSFVNLVEHDREFHHQRYHHREFRFDLSRIPEGEAVTAAEFRIYKDYIRERFDNETFQISVYQVLQEHPGRDSDLFLLDTRIIWAAEEGWLVFDITATSNHWVVNPQHNLGLQLSVESIDGQSINPKLAGLIGRHGPQNKQPFTVAFFKATEVHLRSIRSTGGKQRSQNRSKTPKNQEAFRVSNIAENSSSDQRQACKKHELYVSFRDLGWQDWIIAPEGYAAYYCEGECAFPLNSYMNATNHAIVQTLPCTGRSFLSAARKNFYELQA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJ75</accession>
    <name>A0A452GJ75_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Rap guanine nucleotide exchange factor 2</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="21">Cyclic nucleotide ras GEF</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="23">Neural RAP guanine nucleotide exchange protein</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="20">PDZ domain-containing guanine nucleotide exchange factor 1</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="19">RA-GEF-1</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="22">Ras/Rap1-associating GEF-1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="31 32">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="32" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="31" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="31" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell junction</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Perinuclear region</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Late endosome</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the RAPGEF2 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000001950.1">
        <property type="protein sequence ID" value="ENSGAGP00000001715.1"/>
        <property type="gene ID" value="ENSGAGG00000001344.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070161">
        <property type="term" value="C:anchoring junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005770">
        <property type="term" value="C:late endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048471">
        <property type="term" value="C:perinuclear region of cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007264">
        <property type="term" value="P:small GTPase-mediated signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00038">
        <property type="entry name" value="CAP_ED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd06755">
        <property type="entry name" value="PDZ_RapGEF2_RapGEF6-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd01785">
        <property type="entry name" value="RA_PDZ-GEF1"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="RasGEF"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="REM"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Rap guanine nucleotide exchange factor (GEF) 2"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Rap guanine nucleotide exchange factor (GEF) 6"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="rap guanine nucleotide exchange factor 2 isoform X1"/>
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        <property type="entry name" value="rap guanine nucleotide exchange factor 2 isoform X2"/>
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        <property type="match status" value="1"/>
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    <dbReference type="Gene3D" id="2.60.120.10">
        <property type="entry name" value="Jelly Rolls"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.840.10">
        <property type="entry name" value="Ras guanine-nucleotide exchange factors catalytic domain"/>
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        <property type="entry name" value="Son of sevenless (SoS) protein Chain: S domain 1"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="cNMP-bd_dom"/>
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        <property type="entry name" value="cNMP-bd_dom_sf"/>
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        <property type="entry name" value="PDZ_sf"/>
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        <property type="entry name" value="RA_dom"/>
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        <property type="entry name" value="Ras-like_Gua-exchang_fac_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023578">
        <property type="entry name" value="Ras_GEF_dom_sf"/>
    </dbReference>
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        <property type="entry name" value="RASGEF_cat_dom"/>
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        <property type="entry name" value="RASGEF_cat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014710">
        <property type="entry name" value="RmlC-like_jellyroll"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029071">
        <property type="entry name" value="Ubiquitin-like_domsf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45161">
        <property type="entry name" value="CYTOSKELETON-ASSOCIATED PROTEIN 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45161:SF2">
        <property type="entry name" value="RAP GUANINE NUCLEOTIDE EXCHANGE FACTOR 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00595">
        <property type="entry name" value="PDZ"/>
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        <property type="entry name" value="RA"/>
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        <property type="entry name" value="RasGEF"/>
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    <dbReference type="Pfam" id="PF00618">
        <property type="entry name" value="RasGEF_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00100">
        <property type="entry name" value="cNMP"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="PDZ"/>
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        <property type="entry name" value="RasGEF"/>
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        <property type="entry name" value="RasGEFN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51206">
        <property type="entry name" value="cAMP-binding domain-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50156">
        <property type="entry name" value="PDZ domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48366">
        <property type="entry name" value="Ras GEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54236">
        <property type="entry name" value="Ubiquitin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50042">
        <property type="entry name" value="CNMP_BINDING_3"/>
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    </dbReference>
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        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="RA"/>
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    </dbReference>
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        <property type="entry name" value="RASGEF_CAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50212">
        <property type="entry name" value="RASGEF_NTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="17" id="KW-0965">Cell junction</keyword>
    <keyword evidence="9" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="10" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="14" id="KW-0221">Differentiation</keyword>
    <keyword evidence="13" id="KW-0967">Endosome</keyword>
    <keyword evidence="7" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="12 24" id="KW-0344">Guanine-nucleotide releasing factor</keyword>
    <keyword evidence="18" id="KW-0472">Membrane</keyword>
    <keyword evidence="16" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="11" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="32" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="15" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="Cyclic nucleotide-binding" evidence="27">
        <location>
            <begin position="248"/>
            <end position="348"/>
        </location>
    </feature>
    <feature type="domain" description="N-terminal Ras-GEF" evidence="30">
        <location>
            <begin position="380"/>
            <end position="493"/>
        </location>
    </feature>
    <feature type="domain" description="PDZ" evidence="28">
        <location>
            <begin position="498"/>
            <end position="568"/>
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    <feature type="domain" description="Ras-associating" evidence="29">
        <location>
            <begin position="719"/>
            <end position="805"/>
        </location>
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    <feature type="domain" description="Ras-GEF" evidence="26">
        <location>
            <begin position="830"/>
            <end position="1057"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="25">
        <location>
            <begin position="153"/>
            <end position="172"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="25">
        <location>
            <begin position="181"/>
            <end position="214"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="25">
        <location>
            <begin position="1115"/>
            <end position="1142"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="25">
        <location>
            <begin position="1275"/>
            <end position="1310"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="25">
        <location>
            <begin position="1352"/>
            <end position="1438"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="25">
        <location>
            <begin position="1494"/>
            <end position="1515"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="25">
        <location>
            <begin position="155"/>
            <end position="172"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="25">
        <location>
            <begin position="196"/>
            <end position="207"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="25">
        <location>
            <begin position="1275"/>
            <end position="1284"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="25">
        <location>
            <begin position="1299"/>
            <end position="1310"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="25">
        <location>
            <begin position="1360"/>
            <end position="1383"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="25">
        <location>
            <begin position="1408"/>
            <end position="1418"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="25">
        <location>
            <begin position="1428"/>
            <end position="1437"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="25">
        <location>
            <begin position="1496"/>
            <end position="1506"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
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            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00004282"/>
        </source>
    </evidence>
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        </source>
    </evidence>
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    </evidence>
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    </evidence>
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    </evidence>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022658"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00029925"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00030673"/>
        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00031545"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00031980"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00032021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00168"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="26">
        <source>
            <dbReference type="PROSITE" id="PS50009"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="27">
        <source>
            <dbReference type="PROSITE" id="PS50042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="28">
        <source>
            <dbReference type="PROSITE" id="PS50106"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="29">
        <source>
            <dbReference type="PROSITE" id="PS50200"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="30">
        <source>
            <dbReference type="PROSITE" id="PS50212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="31">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001715.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="32">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1515" mass="170159" checksum="693676391290A433" modified="2019-05-08" version="1">MCETVRYERHEANEVLYYPDDIGSCWYILLSGSVFIKESMFLPRSSFGKRSAGSFRRGCECIVLEPSEMIVVDYMDENEEYFQRQASHRQSRRRFRKINQKGERQTIIDTVDPYPVGKPPLPRGYHTECTKPQLPADFTKLHLTDSLHPQVTHVTSSHSGCSITSDSGSSSLSDIYQATESEAGDMDLSGLPETAVDSEDDDDEEDIERASDPLMSRDIVRDCLEKDPIDRTDDDIEQLLEFMHQLPAFANMTMSVRRELCAVMVFAVVERAGTIVLNDGEELDSWSVILNGSVEVTYPDGRTEILCMGNSFGVSPTMDKEYMKGVMRTKVDDCQFVCIAQQDYCRILNQVEKNMQKVEEEGEIVMVKEHRELDRTGTRKGHIVIKGTSERLTMHLVEEHSVVDPTFIEDFLLTYRTFLSSPMEVGKKLLEWFNDPSLRDKVTRVVLLWVNNHFNDFEGDPAMTRFLEEFENNLEREKMGGHLRLLNIACAAKAKRRLITLTKPSREAPLPFILLGGSEKGFGIFVDSVDLGSKATEAGLKRGDQILEVNGQNFENIQLSKAMEILRNNTHLSITVKTNLFVFKELLTRLSEEKRNGAPHLPKIGDIKKASRYSIPDLAVDVEQVIGLEKVNKKSKANTVGGRNKLKKILDKTRISILPQKPYNDIGIGQSQDDSIVGLRQTKHIPPALPVSGTLSSSNPDLLQSHHRILDFNSTPDLPDQVLRVFKADQQSRYIMISKDTTAKEVVIQAIREFAVTATPDAYSLCEVSVTPEGVIKQRRLPDQLSKLADRIQLSGRYYLKNNMETETLCSDEDAQELLRESQISLLQLSTVEMAAQLSMRNFELFRNIEPTEYIDDLFKLKSKTSCANLKKFEEVINQETFWVASEILRETNQLKRMKIIKHFIKIALHCRECKNFNSMFAIISGLNLAPVARLRTTWEKLPSKYEKLFQDLQDLFDPSRNMAKYRNVLNSQNLQPPIIPLFPVIKKDLTFLHEGNDSKVEGLVNFEKLRMIAKEIRHVGRMASVNMDPAVMFRTRKKKWRSLGSLSQSSTNAAVLDVAQTGGHKKRVRRSSFLNAKKLYEDAQMARKVKQYLSNLDLEMDEESLQTLSLQCEPATNTLPKNPGDKRSGKQEMSPVAPRAGIQQKVQQQHKINQALQVPAVSLYPSRKKVPVKDLPPFGYTLAPSGTVDNFSDSGRSEISSRSSIVSNSSFDSMPVSLHDERRQRHSVSIVETNLGVGRIDRRAMIEPDQYSLGSYAPLSENRGLYAGATVLSSPSTEELSQDQGDRTSLDAADSGRGSWTSCSSGSHDNIQTIQHQRSWETLPFGHVHFDSSGDGAGLWASGSHMDQMMFPDHGTKYGRQSQGREGLDQAQSRASWASSTGYWGEDSEGDTGTIKRRGGKDVSIEAETSSITSVTAEESKPAPMSSHITVSSSSAKGLIARKEGRYREPPPTPPGYVGIPIADFAEGVSHPTRKPPDYNVALQRSRMVARTCETHGTSTLQQQSQEDEQVSAV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJ78</accession>
    <name>A0A452GJ78_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="22">Dynein axonemal heavy chain 11</fullName>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Cilium axoneme</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the dynein heavy chain family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000001893.1">
        <property type="protein sequence ID" value="ENSGAGP00000001667.1"/>
        <property type="gene ID" value="ENSGAGG00000000806.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005930">
        <property type="term" value="C:axoneme"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030286">
        <property type="term" value="C:dynein complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005874">
        <property type="term" value="C:microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031514">
        <property type="term" value="C:motile cilium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045505">
        <property type="term" value="F:dynein intermediate chain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051959">
        <property type="term" value="F:dynein light intermediate chain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007018">
        <property type="term" value="P:microtubule-based movement"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.2620:FF:000001">
        <property type="entry name" value="Cytoplasmic dynein heavy chain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000667">
        <property type="entry name" value="Dynein axonemal heavy chain 11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.920.30:FF:000003">
        <property type="entry name" value="Dynein axonemal heavy chain 17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000219">
        <property type="entry name" value="Dynein axonemal heavy chain 17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.20.180.20:FF:000001">
        <property type="entry name" value="Dynein axonemal heavy chain 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.8.710:FF:000002">
        <property type="entry name" value="dynein heavy chain 17, axonemal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.140.100:FF:000001">
        <property type="entry name" value="dynein heavy chain 17, axonemal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.1120:FF:000002">
        <property type="entry name" value="Dynein heavy chain 9, axonemal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.472.130:FF:000001">
        <property type="entry name" value="Dynein, axonemal, heavy chain 9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.2620">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.472.130">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.8.710">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.1120">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.920.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.140.100">
        <property type="entry name" value="Dynein heavy chain, N-terminal domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.20.180.20">
        <property type="entry name" value="Dynein heavy chain, N-terminal domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035699">
        <property type="entry name" value="AAA_6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026983">
        <property type="entry name" value="DHC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041589">
        <property type="entry name" value="DNAH3_AAA_lid_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042222">
        <property type="entry name" value="Dynein_2_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043157">
        <property type="entry name" value="Dynein_AAA1S"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041466">
        <property type="entry name" value="Dynein_AAA5_ext"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013602">
        <property type="entry name" value="Dynein_heavy_linker"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013594">
        <property type="entry name" value="Dynein_heavy_tail"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042228">
        <property type="entry name" value="Dynein_linker_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45703:SF12">
        <property type="entry name" value="DYNEIN AXONEMAL HEAVY CHAIN 11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45703">
        <property type="entry name" value="DYNEIN HEAVY CHAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12774">
        <property type="entry name" value="AAA_6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12775">
        <property type="entry name" value="AAA_7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17857">
        <property type="entry name" value="AAA_lid_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08385">
        <property type="entry name" value="DHC_N1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08393">
        <property type="entry name" value="DHC_N2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17852">
        <property type="entry name" value="Dynein_AAA_lid"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="13" id="KW-0966">Cell projection</keyword>
    <keyword evidence="10" id="KW-0969">Cilium</keyword>
    <keyword evidence="9 14" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="12" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="8" id="KW-0243">Dynein</keyword>
    <keyword evidence="4" id="KW-0493">Microtubule</keyword>
    <keyword evidence="11" id="KW-0505">Motor protein</keyword>
    <keyword evidence="6" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="Dynein heavy chain tail" evidence="15">
        <location>
            <begin position="29"/>
            <end position="319"/>
        </location>
    </feature>
    <feature type="domain" description="Dynein heavy chain tail" evidence="15">
        <location>
            <begin position="338"/>
            <end position="484"/>
        </location>
    </feature>
    <feature type="domain" description="Dynein heavy chain linker" evidence="16">
        <location>
            <begin position="977"/>
            <end position="1380"/>
        </location>
    </feature>
    <feature type="domain" description="Dynein heavy chain hydrolytic ATP-binding dynein motor region" evidence="17">
        <location>
            <begin position="1511"/>
            <end position="1837"/>
        </location>
    </feature>
    <feature type="domain" description="Dynein heavy chain AAA 5 extension" evidence="18">
        <location>
            <begin position="1995"/>
            <end position="2113"/>
        </location>
    </feature>
    <feature type="domain" description="Dynein heavy chain 3 AAA+ lid" evidence="19">
        <location>
            <begin position="2330"/>
            <end position="2408"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="14">
        <location>
            <begin position="943"/>
            <end position="970"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004430"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008887"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023017"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023069"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="Pfam" id="PF08385"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF08393"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF12774"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF17852"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="Pfam" id="PF17857"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001667.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="22">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="2502" mass="291465" checksum="2E7E17616E30B99E" modified="2019-05-08" version="1">MVLEKLLSTNEGEQILNKKYTPDKITEFYHFLFTALMEAQDIELYLRPLRRHIQFLQETEFPKIHSLIPPLFHTICLIWSHSQFYSVPARIIVLLQEFCNLFIDQARSYLSPEDLLKGEIEETLEHVQIAVNTLRSFKNFFFSHREKLASYFTNSKEFKPWDFQSNMVFARFDLFLNRLDIFVVMLEFQKLEKLEFGGVKGKTLSEQIYRMNEEFIESCKVFKEKTYDPSDYNNEFEEDYVEFKAKILDFDRRLGSILCVGFLDCSGLESAFKLLAIFGYFLEKPVIMEIFSPNYIILLQMFDEELDNCKHLYDEHIRQVCSKAKIKAIFFLVFTALVYQKYIETLTLIDQFEEQIYCKWKSQVDEVYQFNLDQPLIKRNPENGLLSVNFDSKLVALLREVTYLLKLNRSDIPDSALEFYSMNVFFHQQYIGNLELIVQWYNKLKQTVLEVEYPLIETELEAIDGQLRETEEVFTWQKENCWKYIEQVKAAVYDLEQRVQQSKENIKTIQQIMNAWAEHTLFSRKDDRKDALLSLEDKEDRLAKKYKLLKEDGYKIHQLAEVISENLKLFKADSSSDSWKIYIEYIDDIIVDGFFNTLMHNLDFFLENTEKGLKPAPLFQAQMILSAPEIEFKPSLDKEAGDGFYDLVEELLSNVFKMSAQSDMDNMLDLSEIRQEIMDRVADVIRKALDYRNSLDTYAYLWVDDRAEFMKQFLLYGHVLTSEEIDSHADEGAPEQPPTTEQFKEQIDIYENLYIQMSKFEDFRTFESWFKVDMKPFKMSLLNIIKKWSWMFKEHLLKFVIGSLTDLEEFIKVTDAGLQKEVNEGDYNGLVEIMGHLLAVRDRQASTDELFEPLKETITLLESYGQKMSDQVYVQLEELPEKWNTTKKIAASIKHEVAPLQTSEVTLIRKKFALFDGKQIEFRERFRVEAPFCFDAKYPYILLDKENRELEALEEEMLQMQKSANLFEVVVPDYKQLKQCRKEIRLLKEVWDINIYVTSSIGDWTKTQWRQINVEQMDAELRRFAKAMWSLDKEIRSWDVYIGLELTVKNLMTSLRAVTELQNPAIRERHWHQLMNATDVRLSITEGTTLADLLGLQLHKVEDEVRSIVDKAVKELGTEKVEISQTWATVEFCYEEHYRTRTPLLKCDEQLFETLENNQVQLQTILQSKYVEYFIEQVTSWQRTLNVADSVIFIWMEIQRTWSHLESIFIGSDDIRTKLPKDAKRFDGVDADFKELMMETANTKNVLEATTKPNLYEKLESLQRRLSLCEKALAEYLETKCLAFPRFYFVSSADLLDILSKGTQPRQVTRHLTKLFDNIADLQFQDNKEKSTDIALGMYSKEKEYVPFHAECECIGQVETWLQQLEETMYKTVHRHIMEAIAAYEEKPREQWIFDYPAQVALTSSQIWWTTDVGIAFNRLEEGFETALKDYNKKQIAQLNALIALLLGELASGDRQKIMTICTIDVHARDVVAKLVVTSSQAFAWLSQLRHRWDEAQKHCLANICDAQFQYFYEYLGNTSRLVITPLTDRCYITLTQSLHLTMSGAPAGPTGTGKTETTKDLGHALGMMVYVFNCSEQMDYKSIGNIYKGLVQTGAWGCFDEFNRISVEVLSVVAVQVKTIHDAIRNKRKKFLFLGENITLKTSVGIFITMNPGYAGRTELPENLKALFRPCAMVVPDIELICEIMLVAEGFTDARLLARKFITLYTLCRELLSKQDHYDWGLRAVKSVLVVAGSLKRGDKNRPEDQVLMRTLRDFNLPKIVTDDIPIFMGLISDLFPALDVPRRRDLQFEQMVKQSTLELHLQPEESFIFKVVQLEELLAVRHSVFVVGNAGTGKSKVLRTLNRTYVNMKQKLVWNDLNPKAVTTDELFGFIHHATREWKDGLFSSLLREQANITQDGPKWIVLDGDIDPMWIESLNTVMDDNKVLTLASNERVPLTPSMRLLFEIHHLKTATPATVSRAGILYVNPQDLGWNPYVASWIETRSHQSEKANLTILFDKYVPPCLDQLRTSFKTITPIPENSMVQTLCSLLDCLLTPENVPPDCPRELYEIYFVFACVWAFGGALFQDQLSNYQAEFSRWWLKEMKAVKFPSQGTIFDYYLDPKTRKFLPWTDKVPRFDMDPETPLQAVLVHTSETTRLRYFTDLLLERGKPVMLVGNAGVGKTVFVGDRLAAFPDDYMVSNVLFNYYTTSSVLQKILEKSLEKKAGRNYGPVGNKKLVYFIDDMNMPEMDVYGTVQPHTLIRQHIDYGHWYDRQKLTVKEIHNCHYVACMNPTAGNFTINPRLQRHFTVFALNFPSVDALNTIYSKILGFHFQRHVFNPSVLKNAPAMIQASIWLHQTMVQNFLPTAVKFHYIFNLRDLSNIFQGILFATPECLREPNDLVCLWLHESSRVYGDKLVEPKDCHFFRKKLIDTGVEDHLLRQHPLIYCHFANGVASPHYMPVKEWEVLRKILTETLESYNELNASMNLVLFEDAMQHKTIQQCSGLTSSLPVKQGFNLFVFV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJ82</accession>
    <name>A0A452GJ82_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">RRM domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJ82"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001968.1">
        <property type="protein sequence ID" value="ENSGAGP00000001732.1"/>
        <property type="gene ID" value="ENSGAGG00000001387.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003729">
        <property type="term" value="F:mRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.330:FF:000058">
        <property type="entry name" value="RNA-binding motif protein 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.330:FF:000085">
        <property type="entry name" value="RNA-binding protein 4 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.330">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050502">
        <property type="entry name" value="Euk_RNA-bind_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012677">
        <property type="entry name" value="Nucleotide-bd_a/b_plait_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035979">
        <property type="entry name" value="RBD_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000504">
        <property type="entry name" value="RRM_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48025:SF26">
        <property type="entry name" value="HETEROGENEOUS NUCLEAR RIBONUCLEOPROTEIN M-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48025">
        <property type="entry name" value="OS02G0815200 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00076">
        <property type="entry name" value="RRM_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00360">
        <property type="entry name" value="RRM"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54928">
        <property type="entry name" value="RNA-binding domain, RBD"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50102">
        <property type="entry name" value="RRM"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="6 8" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <keyword evidence="4" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="RRM" evidence="9">
        <location>
            <begin position="2"/>
            <end position="72"/>
        </location>
    </feature>
    <feature type="domain" description="RRM" evidence="9">
        <location>
            <begin position="78"/>
            <end position="138"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00176"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50102"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001732.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="167" mass="18784" checksum="53DA349F76A0C8F9" modified="2019-05-08" version="1">MVKLFIGNLPREATEQEIRSLFEQYGKVLECDIIKNYGFVHIEDKTAAEDAIRNLHHHKLHGVCINVEASKNKSKASTKLHVGNISTTCTNLELRAKFEEYGPVIECDIVKDYAFVHMERAEDAVEAIRGLDNTEFQGSIFSGGPSGSHYTRKFVLLPPPSHCLRRG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJ84</accession>
    <name>A0A452GJ84_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Essential for reactive oxygen species protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum membrane</location>
            <topology evidence="1">Single-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the CYBC1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJ84"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001881.1">
        <property type="protein sequence ID" value="ENSGAGP00000001656.1"/>
        <property type="gene ID" value="ENSGAGG00000001318.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045087">
        <property type="term" value="P:innate immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027846">
        <property type="entry name" value="Cybc1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31837">
        <property type="entry name" value="CYTOCHROME B-245 CHAPERONE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31837:SF3">
        <property type="entry name" value="CYTOCHROME B-245 CHAPERONE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15169">
        <property type="entry name" value="Cybc1_Eros"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0143">Chaperone</keyword>
    <keyword evidence="5" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="6" id="KW-0391">Immunity</keyword>
    <keyword evidence="3" id="KW-0399">Innate immunity</keyword>
    <keyword evidence="8 11" id="KW-0472">Membrane</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 11" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 11" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="21"/>
            <end position="40"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="88"/>
            <end position="110"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004389"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009907"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022588"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022859"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023186"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00030424"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001656.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="188" mass="21162" checksum="497B098EFFB0B651" modified="2019-05-08" version="1">MYMLLENRTSSILHLKRSPGIRSWSLFVGISSIGLAAAYYSEDSLGWKLFYIAGCLFVAAQNLEEWEEAVFDKDKGKSLESQLYKSEMSMLGMAVTDVLPLLLLTVAASLNEIRDVNVEEEKVRYFGKGYLVVLRFVTGFSHPLTQSAVLGCRRYELFFSIMSVSIRGVGCFVTKPYLQICCDCCPLF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJ85</accession>
    <name>A0A452GJ85_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">S-arrestin</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="4">Retinal S-antigen</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="3">Rod photoreceptor arrestin</fullName>
        </alternativeName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the arrestin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJ85"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001961.1">
        <property type="protein sequence ID" value="ENSGAGP00000001726.1"/>
        <property type="gene ID" value="ENSGAGG00000001371.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001917">
        <property type="term" value="C:photoreceptor inner segment"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001750">
        <property type="term" value="C:photoreceptor outer segment"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001664">
        <property type="term" value="F:G protein-coupled receptor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002031">
        <property type="term" value="P:G protein-coupled receptor internalization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.840:FF:000002">
        <property type="entry name" value="Arrestin 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.640:FF:000011">
        <property type="entry name" value="S-arrestin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.640">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.840">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000698">
        <property type="entry name" value="Arrestin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014752">
        <property type="entry name" value="Arrestin-like_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011021">
        <property type="entry name" value="Arrestin-like_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011022">
        <property type="entry name" value="Arrestin_C-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017864">
        <property type="entry name" value="Arrestin_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014753">
        <property type="entry name" value="Arrestin_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014756">
        <property type="entry name" value="Ig_E-set"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11792">
        <property type="entry name" value="ARRESTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11792:SF15">
        <property type="entry name" value="S-ARRESTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02752">
        <property type="entry name" value="Arrestin_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00339">
        <property type="entry name" value="Arrestin_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00309">
        <property type="entry name" value="ARRESTIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM01017">
        <property type="entry name" value="Arrestin_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81296">
        <property type="entry name" value="E set domains"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00295">
        <property type="entry name" value="ARRESTINS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Arrestin C-terminal-like" evidence="5">
        <location>
            <begin position="212"/>
            <end position="373"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00005298"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00040206"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00041305"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00042071"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="SMART" id="SM01017"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001726.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="385" mass="43326" checksum="0144FE147BFF2BE4" modified="2019-05-08" version="1">MSCAESHKVATPSSICLKNTSPPQSHVVYKKMSRDKAVTIYLGKRDFIDHIDGVEPVDGVLLVDPEIVKGKKVYVTLTCAFRYGQEDIDVMGLTFRKDLFFSRIQLYPPVEKWESLSQLQECLMKKLGNNAYPFVLTFPDYLPCSVSLQPAPHDVGKCCGVDFEVKAFCTENVEERIPKRNFVRLLIRKVQYAPEESGPQPQAETTWQFFMSSKPLHLRARLSKEVFYHGEPIPVTVTVTNNTEKTVKKIRVLVEQVANVVLYSSDYYTKVVAMEEMQEKVQPNSIITKTLILLPLLANNREKRGIALDGKLKDEDTNLASSTIIKDGIDKTVLGILVAYKVKVKLIISGMLGDLTSSEVSTELPFRLMHPKPEEPNIRGLSSKG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJ87</accession>
    <name>A0A452GJ87_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">RRM domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJ87"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001973.1">
        <property type="protein sequence ID" value="ENSGAGP00000001737.1"/>
        <property type="gene ID" value="ENSGAGG00000001387.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003729">
        <property type="term" value="F:mRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.330:FF:000058">
        <property type="entry name" value="RNA-binding motif protein 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.330:FF:000085">
        <property type="entry name" value="RNA-binding protein 4 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.330">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050502">
        <property type="entry name" value="Euk_RNA-bind_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012677">
        <property type="entry name" value="Nucleotide-bd_a/b_plait_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035979">
        <property type="entry name" value="RBD_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000504">
        <property type="entry name" value="RRM_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48025:SF26">
        <property type="entry name" value="HETEROGENEOUS NUCLEAR RIBONUCLEOPROTEIN M-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48025">
        <property type="entry name" value="OS02G0815200 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00076">
        <property type="entry name" value="RRM_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00360">
        <property type="entry name" value="RRM"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54928">
        <property type="entry name" value="RNA-binding domain, RBD"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50102">
        <property type="entry name" value="RRM"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="6 8" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <keyword evidence="4" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="RRM" evidence="9">
        <location>
            <begin position="2"/>
            <end position="72"/>
        </location>
    </feature>
    <feature type="domain" description="RRM" evidence="9">
        <location>
            <begin position="78"/>
            <end position="138"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00176"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50102"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001737.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="186" mass="20708" checksum="1267872C35293368" modified="2019-05-08" version="1">MVKLFIGNLPREATEQEIRSLFEQYGKVLECDIIKNYGFVHIEDKTAAEDAIRNLHHHKLHGVCINVEASKNKSKASTKLHVGNISTTCTNLELRAKFEEYGPVIECDIVKDYAFVHMERAEDAVEAIRGLDNTEFQGSIFSGGPSGSHYTRKFVLLPPPSHCLRRVLAASSVFIFNGGSCVSVKL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJ89</accession>
    <name>A0A452GJ89_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Ig-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJ89"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001967.1">
        <property type="protein sequence ID" value="ENSGAGP00000001731.1"/>
        <property type="gene ID" value="ENSGAGG00000001370.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005769">
        <property type="term" value="C:early endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="C:late endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046790">
        <property type="term" value="F:virion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0075512">
        <property type="term" value="P:clathrin-dependent endocytosis of virus by host cell"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013162">
        <property type="entry name" value="CD80_C2-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013106">
        <property type="entry name" value="Ig_V-set"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47243">
        <property type="entry name" value="SIALOADHESIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47243:SF1">
        <property type="entry name" value="SIALOADHESIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08205">
        <property type="entry name" value="C2-set_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13895">
        <property type="entry name" value="Ig_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07686">
        <property type="entry name" value="V-set"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="4" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="2" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="3" id="KW-1133">Transmembrane helix</keyword>
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        <location>
            <begin position="1"/>
            <end position="25"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019230080" description="Ig-like domain-containing protein" evidence="6">
        <location>
            <begin position="26"/>
            <end position="344"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="7">
        <location>
            <begin position="244"/>
            <end position="328"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004167"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001731.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="344" mass="37800" checksum="EADB81A751EEBBF7" modified="2019-05-08" version="1">YPVGAVMPILFQLIFLASHALQAFGSWGVSYPLSLHSVKDSCILIPCTFSYPSTVSSTGSIVAIWYKDYDSQRIVVYHSATPSEVDGRFQGRTELLGDLAGHNCTLLLRGVRSEDSGKYNFRFEINQGDRWSDLRGVMVTVTDAPTSPAIAAPEDLRESMRVNFTCSSPYACPYDSITLQWRGYNPVLSTVTGTVQLDTTGVLSRQTLVSSLSWQDHSKILFCEVSVGSKQATGEIALRVKYSPKGTKVVINPSTKNIRVGDAVSLICSVNSTFPEVTTYKWYKGGTALGESQQILTLQSVAREDYGLYHCEAENSNDKGSEMSQAISLSVFCKLVLFFKHCPC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJ90</accession>
    <name>A0A452GJ90_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Essential for reactive oxygen species protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum membrane</location>
            <topology evidence="1">Single-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the CYBC1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJ90"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001886.1">
        <property type="protein sequence ID" value="ENSGAGP00000001661.1"/>
        <property type="gene ID" value="ENSGAGG00000001318.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045087">
        <property type="term" value="P:innate immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027846">
        <property type="entry name" value="Cybc1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31837">
        <property type="entry name" value="CYTOCHROME B-245 CHAPERONE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31837:SF3">
        <property type="entry name" value="CYTOCHROME B-245 CHAPERONE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15169">
        <property type="entry name" value="Cybc1_Eros"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0143">Chaperone</keyword>
    <keyword evidence="5" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="6" id="KW-0391">Immunity</keyword>
    <keyword evidence="3" id="KW-0399">Innate immunity</keyword>
    <keyword evidence="8" id="KW-0472">Membrane</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="12" id="KW-0732">Signal</keyword>
    <keyword evidence="4" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="12">
        <location>
            <begin position="1"/>
            <end position="17"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019564650" description="Essential for reactive oxygen species protein" evidence="12">
        <location>
            <begin position="18"/>
            <end position="104"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="84"/>
            <end position="104"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="11">
        <location>
            <begin position="93"/>
            <end position="104"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004389"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009907"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022588"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022859"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023186"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00030424"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001661.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="104" mass="11486" checksum="2C0BBE9AF87E5CB4" modified="2019-05-08" version="1">MAVTDVLPLLLLTVAASLNEIRDVNVEEEKVRYFGKGYLVVLRFVTGFSHPLTQSAVLGCRSDVEAVAKLITGFLELNKVENHQDLSQSSETEASDADEPEDEY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJB7</accession>
    <name>A0A452GJB7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">Diacylglycerol kinase</fullName>
            <shortName evidence="21">DAG kinase</shortName>
            <ecNumber evidence="21">2.7.1.107</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="26 27">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="27" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="26" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="26" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>1,2-di-(9Z-octadecenoyl)-sn-glycerol + ATP = 1,2-di-(9Z-octadecenoyl)-sn-glycero-3-phosphate + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:40327"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:52333"/>
            <dbReference type="ChEBI" id="CHEBI:74546"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="18">
            <dbReference type="Rhea" id="RHEA:40328"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>a 1,2-diacyl-sn-glycerol + ATP = a 1,2-diacyl-sn-glycero-3-phosphate + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10272"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:17815"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:58608"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.1.107"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="19">
            <dbReference type="Rhea" id="RHEA:10273"/>
        </physiologicalReaction>
    </comment>
    <comment type="pathway">
        <text evidence="20">Glycerolipid metabolism.</text>
    </comment>
    <comment type="pathway">
        <text evidence="3">Lipid metabolism; glycerolipid metabolism.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4 21">Belongs to the eukaryotic diacylglycerol kinase family.</text>
    </comment>
    <dbReference type="EC" id="2.7.1.107" evidence="21"/>
    <dbReference type="AlphaFoldDB" id="A0A452GJB7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001916.1">
        <property type="protein sequence ID" value="ENSGAGP00000001686.1"/>
        <property type="gene ID" value="ENSGAGG00000001299.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00230"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004143">
        <property type="term" value="F:ATP-dependent diacylglycerol kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046486">
        <property type="term" value="P:glycerolipid metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007200">
        <property type="term" value="P:phospholipase C-activating G protein-coupled receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd20847">
        <property type="entry name" value="C1_DGKdelta_rpt1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd20893">
        <property type="entry name" value="C1_DGKdelta_rpt2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd13274">
        <property type="entry name" value="PH_DGK_type2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000060">
        <property type="entry name" value="Diacylglycerol kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.200.40:FF:000001">
        <property type="entry name" value="Diacylglycerol kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.60.20:FF:000002">
        <property type="entry name" value="Diacylglycerol kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.60.20:FF:000029">
        <property type="entry name" value="Diacylglycerol kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10330:FF:000001">
        <property type="entry name" value="Diacylglycerol kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.200.40">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.60.20">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.10330">
        <property type="entry name" value="Probable inorganic polyphosphate/atp-NAD kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017438">
        <property type="entry name" value="ATP-NAD_kinase_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046349">
        <property type="entry name" value="C1-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047478">
        <property type="entry name" value="C1_DGKdelta_rpt1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047477">
        <property type="entry name" value="C1_DGKdelta_rpt2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037607">
        <property type="entry name" value="DGK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR054474">
        <property type="entry name" value="DGKD_4H"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000756">
        <property type="entry name" value="Diacylglycerol_kin_accessory"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001206">
        <property type="entry name" value="Diacylglycerol_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016064">
        <property type="entry name" value="NAD/diacylglycerol_kinase_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002219">
        <property type="entry name" value="PKC_DAG/PE"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11255">
        <property type="entry name" value="DIACYLGLYCEROL KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11255:SF30">
        <property type="entry name" value="DIACYLGLYCEROL KINASE DELTA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00130">
        <property type="entry name" value="C1_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00609">
        <property type="entry name" value="DAGK_acc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00781">
        <property type="entry name" value="DAGK_cat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22944">
        <property type="entry name" value="DGKD_4H"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00169">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00109">
        <property type="entry name" value="C1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00045">
        <property type="entry name" value="DAGKa"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00046">
        <property type="entry name" value="DAGKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00233">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57889">
        <property type="entry name" value="Cysteine-rich domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF111331">
        <property type="entry name" value="NAD kinase/diacylglycerol kinase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50146">
        <property type="entry name" value="DAGK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00479">
        <property type="entry name" value="ZF_DAG_PE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50081">
        <property type="entry name" value="ZF_DAG_PE_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="15 21" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="5" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="13 21" id="KW-0418">Kinase</keyword>
    <keyword evidence="16" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="17" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11 21" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="27" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0677">Repeat</keyword>
    <keyword evidence="8 21" id="KW-0808">Transferase</keyword>
    <keyword evidence="14" id="KW-0862">Zinc</keyword>
    <keyword evidence="12" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="PH" evidence="23">
        <location>
            <begin position="1"/>
            <end position="88"/>
        </location>
    </feature>
    <feature type="domain" description="Phorbol-ester/DAG-type" evidence="24">
        <location>
            <begin position="105"/>
            <end position="155"/>
        </location>
    </feature>
    <feature type="domain" description="Phorbol-ester/DAG-type" evidence="24">
        <location>
            <begin position="177"/>
            <end position="228"/>
        </location>
    </feature>
    <feature type="domain" description="DAGKc" evidence="25">
        <location>
            <begin position="259"/>
            <end position="394"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="497"/>
            <end position="543"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="600"/>
            <end position="626"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="1068"/>
            <end position="1112"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="22">
        <location>
            <begin position="497"/>
            <end position="506"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="22">
        <location>
            <begin position="523"/>
            <end position="537"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="22">
        <location>
            <begin position="1072"/>
            <end position="1081"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="22">
        <location>
            <begin position="1095"/>
            <end position="1106"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00009280"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023371"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023411"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00060536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="RuleBase" id="RU361128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS50081"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS50146"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001686.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1129" mass="125906" checksum="8A6B4EA0CEE29FE0" modified="2019-05-08" version="1">MLMKQTSSFQRWKRRYFKLRGRTLYYAKNAKSIIFDEVDLTDASVAESSTKNINNSFTVITPCRKLILCADNRKEMEDWIAALKTIQNREHFESTQYSMDHFSGMHNWYACSHARPTYCNVCREALSGVTSHGLSCEVCKFKAHKRCAVRATNNCKWTTLASIGKDIIEDEDGISMPHQWLEGNLPVSAKCMVCDKTCGSVLRLQDWRCLWCKAMVHTTCKDLLPTKCPLGLCKVSVIPPTALNSIDSDGFWKATCPPSCTSPLLVFVNSKSGDNQGVKFLRRFKQLLNPAQVFDLMNGGPHLGLRLFQKFDTFRILVCGGDGSVGWVLSEIDSLNLHKQCQLGVLPLGTGNDLARVLGWGSACDDDTQLPQILEKLERASTKMLDRWSIMVYETKLPRQSSTSTVTEDFSDDSEVQKILFYEDSVAAHLSKILTSDQHSVVISSAKVLCETVKDFVARVGKAYEKATESSEESEVMARKCSVLKEKLDSLLKTLNDESQASSSLSNPPPTIAEETEDGDGSGSACDSSSDHSMGSSCTARPQIFRPREQLMLRANSLKKAIRQIIEHAEKAVDEQNAQTQEQEGFILGLPTSEEGKELRNEDKTSLQPLHSGNYGVPKGRSHRKVSKSPCERLINKGSLSLGSSASLPPQTGNRDNLPMLNTKILYPNIRAGMSGSLPGSSVISRLLINADPFNSEPENLECYTEKCVMNNYFGIGLDAKISLDFNNKRDEHPEKCRSRTKNMMWYGVLGTKELLHRTYKNLEQKVLLECDGRPIPLPSLQGIAVLNIPSYAGGTNFWGGTKEDDTFTAPSFDDKILEVVAVFGSMQMAVSRVINLQHHRIAQCRTVKIAILGDEGVPVQVDGEAWIQPPGYIWIVHKNRAQTLTRDRAFESTLKSWEDKQKCELPRPSSFSLHPEIMSEEESTQINQFGQAAGALIHSIREIAQSYQDMEQELAHAVNASSKSMDKVYAKSKSTEGLNCSLVVEMVNNVKALHSETELLLAGKMALQLDPPQKEQLLAALMDMDLQLRKLADIPWLWQLMEPSDDENQMLDYSKRSRSGKFRLVTKFKKEKNNKNKETRSSMGLPGPGSDQSGSHEEDITRDQGAEPEYSCQRGLASVFTACVYHSP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJB8</accession>
    <name>A0A452GJB8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Nuclear pore complex protein Nup98-Nup96</fullName>
        </recommendedName>
    </protein>
    <organism evidence="25 26">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="26" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="25" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="25" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="22">Plays a role in the nuclear pore complex (NPC) assembly and/or maintenance. NUP98 and NUP96 are involved in the bidirectional transport across the NPC. May anchor NUP153 and TPR to the NPC. In cooperation with DHX9, plays a role in transcription and alternative splicing activation of a subset of genes. Involved in the localization of DHX9 in discrete intranuclear foci (GLFG-body).</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Nucleus membrane</location>
            <topology evidence="2">Peripheral membrane protein</topology>
            <orientation evidence="2">Nucleoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
            <location evidence="1">Nuclear pore complex</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Nucleus</location>
            <location evidence="3">Nucleoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the nucleoporin GLFG family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000002016.1">
        <property type="protein sequence ID" value="ENSGAGP00000001774.1"/>
        <property type="gene ID" value="ENSGAGG00000001410.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031965">
        <property type="term" value="C:nuclear membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0044614">
        <property type="term" value="C:nuclear pore cytoplasmic filaments"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005654">
        <property type="term" value="C:nucleoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008139">
        <property type="term" value="F:nuclear localization sequence binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008236">
        <property type="term" value="F:serine-type peptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017056">
        <property type="term" value="F:structural constituent of nuclear pore"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051028">
        <property type="term" value="P:mRNA transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000973">
        <property type="term" value="P:post-transcriptional tethering of RNA polymerase II gene DNA at nuclear periphery"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006606">
        <property type="term" value="P:protein import into nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006405">
        <property type="term" value="P:RNA export from nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034398">
        <property type="term" value="P:telomere tethering at nuclear periphery"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.2360:FF:000001">
        <property type="entry name" value="Nuclear pore complex protein Nup98-Nup96"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.690:FF:000001">
        <property type="entry name" value="Nuclear pore complex protein Nup98-Nup96"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1610.10:FF:000001">
        <property type="entry name" value="Nuclear pore complex protein Nup98-Nup96"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.2360">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.690">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1610.10">
        <property type="entry name" value="Peptidase S59, nucleoporin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037665">
        <property type="entry name" value="Nucleoporin_S59-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007230">
        <property type="entry name" value="Nup98_auto-Pept-S59_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036903">
        <property type="entry name" value="Nup98_auto-Pept-S59_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021967">
        <property type="entry name" value="Nup98_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23198:SF6">
        <property type="entry name" value="NUCLEAR PORE COMPLEX PROTEIN NUP98-NUP96"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23198">
        <property type="entry name" value="NUCLEOPORIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04096">
        <property type="entry name" value="Nucleoporin2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12110">
        <property type="entry name" value="Nup96"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21240">
        <property type="entry name" value="Nup98_GLEBS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF82215">
        <property type="entry name" value="C-terminal autoproteolytic domain of nucleoporin nup98"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51434">
        <property type="entry name" value="NUP_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="17" id="KW-0007">Acetylation</keyword>
    <keyword evidence="12" id="KW-0068">Autocatalytic cleavage</keyword>
    <keyword evidence="11" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="7" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="20" id="KW-0472">Membrane</keyword>
    <keyword evidence="13" id="KW-0509">mRNA transport</keyword>
    <keyword evidence="19" id="KW-0906">Nuclear pore complex</keyword>
    <keyword evidence="21" id="KW-0539">Nucleus</keyword>
    <keyword evidence="8" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="9" id="KW-0645">Protease</keyword>
    <keyword evidence="16" id="KW-0653">Protein transport</keyword>
    <keyword evidence="26" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0677">Repeat</keyword>
    <keyword evidence="14" id="KW-0720">Serine protease</keyword>
    <keyword evidence="18" id="KW-0811">Translocation</keyword>
    <keyword evidence="6" id="KW-0813">Transport</keyword>
    <keyword evidence="15" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="Peptidase S59" evidence="24">
        <location>
            <begin position="679"/>
            <end position="821"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="23">
        <location>
            <begin position="454"/>
            <end position="519"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="23">
        <location>
            <begin position="463"/>
            <end position="472"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004567"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004620"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004642"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00008926"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00013472"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022813"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022816"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023010"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023132"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00059309"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS51434"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001774.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1755" mass="191448" checksum="57CF136C26CC13B6" modified="2019-05-08" version="1">MFNKSFGTPFGGSAGGFGTTSTFGQNAGFGTASGGAFGTSAFGSNNNTGSLFGNTQTKPGGLFGSSTFSQPATSSTSTGFGFGTSTGTSNSLFGTASTGGGLFSSQNNAFAQNKPAGFGNFGTSTSSGGLFGTTNTASNPFGGTSGSLFGPTSFTAAPTGTTIKFNPPTGTDTMVKSGVSTNISTKHQCITAMKEYESKSLEELRLEDYQANRKGPSNPAVAAAGLFGSSTATSSAATGLFGSSTTNAGFSYGQNKTAFGTSTTGFGTGTAGSLFGQQPQQTTSLFSKPFGQATTTQNTGFSFGNTSTLGQPNTNTMGLFGATQASPSGSLFGTAANTNAATGFGAGTGTGLFGQANTGFGVGGSTLFGKPAGFGTTTTSAPSFGTTTGSGLFGFGASTAGNSIFGNKPAAGTLGAGLGTGFGTALTDPNVSAAQQAALQQHFNSLTYSPFGDSPLFRNPMSDPKKKEERLKPTNPAAQKALTTPTHYKLTPRPATRVRPKALQTAGSSKSHLFDGLDDDEPSLSNGAFMPKKSIKKLVLKNLNSSNLFSPVSREAEDLDPPSEYPENGERFNFLSRSVDENHRQEGEREEEDDHHEVTRFYTNPIAKPIPQTPENAVHKHQNNVDDTIVALNMRVALRNGLEGSSEDASFHDESLQDDREEENENVYSLHPAGIVLTKAGYYTIPSMEDLAKLTTDRGECIVTDFTIGRKGYGSIYFEGEVNLTNLNLDEIVHIRRKEVIVYPDDDQKPPIGEGLNRRAEVTLDGVWPTDKTSRCLIKSPERLADINYEGRLEAVSRKQGAQFKEYRPETGSWVFKVAHFSKYGLQDSDEEEEEHPSKMDAKKLKTAQVPPQGQLIPQQMALNGKPTSPSQSPEVEQLGRVVELDSDMADITQEPAQDSILEDSITEEPELVPASTHIASSLGINPHVLQIMKASLLADEEDVDLILDHHFGKQPTKMDTSQEICSPRLPISALQGQKRHSSVGGLLQSKFASVSFLQSASLPDFHGPRASSFGNPPSAASWTGLSPLPSAFSMPSAAPEVQLKTVGACRQQGLIPLEKSVTYGKGRLLMDMALFMGRSFRVGWGPNWTLVNSGDKLSGSSEPKGLRNDSMEYGFLPTPVAPKSLSESPFKVHIEKLTLEQRKTDGDQQLYLIPLEIKLKHSTVHVDERCPLLAANPGVSAIHDYADWVRDFCGDSVEIEPVVKYWCLTWTLCEAIWGHLKELEASLEEPSEYIVTLERRKAFSRWLSQTAAARIEEEVSLSQHENHIEAVFSYLTGKRISEACRVAQQSGDHRLALLLSQLVGSPQVRELLTMQLVDWHGLQADCFIQEERLRIFALLAGKPVWQLSERKSINVCSQLDWKRSIAIHLWYLLPPTASISKALNMYEAAFQNTPECEKYACCPLPPYLEDSGCVLEEDDSTERPLRDVCFHLLKLYSDRCYDLNQLLDPRSITSDPLDYRLSWHLWEVLRALNYTHLSEQCQGVLNASYAAQLESEGLWEWAIFVLLHTSNTYTREKAVREFLNRHCVLSETPESWTKETFLTQKLCIPPEWIQEAKAVRARMEGDKHKEALYLFKAGHWNQCHKLVIRHLASDAIINENYNYLKGFLEDLSPPERSTLIQDWDTAGLVYLDYIRVIEMLDRIQQLDCSAYELERLHTKVTSLCNRIEQIQCHNAKDRLAQSDMAKRVANLLRVVLSLQHSPEPASDSTPDVQRVPLRLLAPHIGRLPMPEDYALEELRSLTQSYLRELTVVTQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJB9</accession>
    <name>A0A452GJB9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">lysophospholipase</fullName>
            <ecNumber evidence="3">3.1.1.5</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>1-(9Z-octadecenoyl)-sn-glycero-3-phosphocholine + H2O = sn-glycerol 3-phosphocholine + (9Z)-octadecenoate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:40807"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16870"/>
            <dbReference type="ChEBI" id="CHEBI:28610"/>
            <dbReference type="ChEBI" id="CHEBI:30823"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="13">
            <dbReference type="Rhea" id="RHEA:40808"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="14">
            <text>1-hexadecanoyl-sn-glycero-3-phosphate + H2O = sn-glycerol 3-phosphate + hexadecanoate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:49092"/>
            <dbReference type="ChEBI" id="CHEBI:7896"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57518"/>
            <dbReference type="ChEBI" id="CHEBI:57597"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="14">
            <dbReference type="Rhea" id="RHEA:49093"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="16">
            <text>1-hexadecanoyl-sn-glycero-3-phosphocholine + H2O = sn-glycerol 3-phosphocholine + hexadecanoate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:40435"/>
            <dbReference type="ChEBI" id="CHEBI:7896"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16870"/>
            <dbReference type="ChEBI" id="CHEBI:72998"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="16">
            <dbReference type="Rhea" id="RHEA:40436"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>a 1-acyl-sn-glycero-3-phosphocholine + H2O = sn-glycerol 3-phosphocholine + a fatty acid + H(+)</text>
            <dbReference type="Rhea" id="RHEA:15177"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16870"/>
            <dbReference type="ChEBI" id="CHEBI:28868"/>
            <dbReference type="ChEBI" id="CHEBI:58168"/>
            <dbReference type="EC" id="3.1.1.5"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="15">
            <dbReference type="Rhea" id="RHEA:15178"/>
        </physiologicalReaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum membrane</location>
            <topology evidence="1">Single-pass type III membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the NTE family.</text>
    </comment>
    <dbReference type="EC" id="3.1.1.5" evidence="3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001944.1">
        <property type="protein sequence ID" value="ENSGAGP00000001709.1"/>
        <property type="gene ID" value="ENSGAGG00000001350.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004622">
        <property type="term" value="F:phosphatidylcholine lysophospholipase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016042">
        <property type="term" value="P:lipid catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046470">
        <property type="term" value="P:phosphatidylcholine metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00038">
        <property type="entry name" value="CAP_ED"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="CDD" id="cd07225">
        <property type="entry name" value="Pat_PNPLA6_PNPLA7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.10:FF:000010">
        <property type="entry name" value="neuropathy target esterase isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.10:FF:000012">
        <property type="entry name" value="neuropathy target esterase isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.1090.10:FF:000001">
        <property type="entry name" value="neuropathy target esterase isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.10:FF:000022">
        <property type="entry name" value="Patatin like phospholipase domain containing 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.1090.10">
        <property type="entry name" value="Cytosolic phospholipase A2 catalytic domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.10">
        <property type="entry name" value="Jelly Rolls"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016035">
        <property type="entry name" value="Acyl_Trfase/lysoPLipase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000595">
        <property type="entry name" value="cNMP-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018490">
        <property type="entry name" value="cNMP-bd_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001423">
        <property type="entry name" value="LysoPLipase_patatin_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050301">
        <property type="entry name" value="NTE"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056556">
        <property type="entry name" value="NTE1_P-loop_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002641">
        <property type="entry name" value="PNPLA_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014710">
        <property type="entry name" value="RmlC-like_jellyroll"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14226">
        <property type="entry name" value="NEUROPATHY TARGET ESTERASE/SWISS CHEESE D.MELANOGASTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14226:SF26">
        <property type="entry name" value="PATATIN-LIKE PHOSPHOLIPASE DOMAIN-CONTAINING PROTEIN 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00027">
        <property type="entry name" value="cNMP_binding"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24179">
        <property type="entry name" value="NTE_Ploop"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01734">
        <property type="entry name" value="Patatin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00100">
        <property type="entry name" value="cNMP"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51206">
        <property type="entry name" value="cAMP-binding domain-like"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52151">
        <property type="entry name" value="FabD/lysophospholipase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50042">
        <property type="entry name" value="CNMP_BINDING_3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51635">
        <property type="entry name" value="PNPLA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01237">
        <property type="entry name" value="UPF0028"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="7 17" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="9 17" id="KW-0442">Lipid degradation</keyword>
    <keyword evidence="11 17" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="12 19" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5 19" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="10 19" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="36"/>
            <end position="59"/>
        </location>
    </feature>
    <feature type="domain" description="Cyclic nucleotide-binding" evidence="20">
        <location>
            <begin position="174"/>
            <end position="301"/>
        </location>
    </feature>
    <feature type="domain" description="Cyclic nucleotide-binding" evidence="20">
        <location>
            <begin position="494"/>
            <end position="598"/>
        </location>
    </feature>
    <feature type="domain" description="Cyclic nucleotide-binding" evidence="20">
        <location>
            <begin position="626"/>
            <end position="731"/>
        </location>
    </feature>
    <feature type="domain" description="PNPLA" evidence="21">
        <location>
            <begin position="962"/>
            <end position="1128"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="336"/>
            <end position="372"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="427"/>
            <end position="452"/>
        </location>
    </feature>
    <feature type="short sequence motif" description="GXGXXG" evidence="17">
        <location>
            <begin position="966"/>
            <end position="971"/>
        </location>
    </feature>
    <feature type="short sequence motif" description="GXSXG" evidence="17">
        <location>
            <begin position="993"/>
            <end position="997"/>
        </location>
    </feature>
    <feature type="short sequence motif" description="DGA/G" evidence="17">
        <location>
            <begin position="1115"/>
            <end position="1117"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="427"/>
            <end position="441"/>
        </location>
    </feature>
    <feature type="active site" description="Nucleophile" evidence="17">
        <location>
            <position position="995"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="17">
        <location>
            <position position="1115"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004643"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006636"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00013274"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022963"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00047314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00048133"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00048454"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00048656"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01161"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS51635"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001709.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1344" mass="150898" checksum="B30ABDF7A3BE6B2A" modified="2019-05-08" version="1">MGQSESELQDEQEDVVLTAVLPNLKLFVEDQLQTSMVLGVMIGVGVSMLLIATLILVLVRRLRHKKVQTQETPKYRFRKRDKVLFYGRKIMRKVSQSTSSLVDTTISNTSRPRMKKKLKMLNIAKKILRIKKELPILQLKEPPPSVLEADLTEFDVANSHLPSEVLYMLKNVRVLGHFEKPLFLELCKHMIFQQFQQGEYVFRPGQPDSSIYVVQDGKLELFLTQQDGKETLVKEVFPGDSVHSLLSILDVITGHQRPYRTVSARAGEDSTVLRLPVEAFSAVFEKYPESLVRVVQIIMVRLQRVTFLALHNYLGLTNELFSHEMQLLRLFPQPNHASRTSPVRHNKRVISISSIDDHKESSGRQPDASAKDLADQLKLGSLEPSSVPLLSRCISMPVDISGLQKGPRSDFDMAYERGRISVSLQEESTGTMASFSQSVSQEPKERRSVTVEEQSPGTYHYNYCEDDSAGGACPFGPYQGRQTSAIFEAAKRELVKLMKIEDPSLLNNRVLLHHAKAGTVIARQGDQDVSLHFVLWGCLHVYQRMIDKAEDVCLFLTQPGEMVGQLAVLTGEPLIFTIKANRDCTFLRISKSDFYEIMREQPSVVLSVAHTVAARMSPFVRQMDFAIDWMAVEAGRALYRQGDKSDCTYIVLNGRLRSVIQKGNGKKELVGEYGRGDLIGVVEALTRQPRATTVHAVRDTELAKLPEGTLNNIKRRYPQVVTRLIHLLSQKILGNLQQLCGPFTGSSLGVASSSELTNPASNLSTVAVLPVCDDVPMAAFTLELEHALNAIGPTLLLTSDIIRARLGASALDSIQEYRLSGWLAQQEDIHRIVLYQTDCTLTPWTVRCIRQADCILIVGLGDQEPALGKLEQMLENTAVRALKQLILLHREDGPSPSRTVEWLNMRSWCSGHLHIKCPRRVFSRRSPNKLREMYAKVFEKDADRHSDFSRLARVLTGNSIALVLGGGGARGCSHIGVIKAMEEAGIPIDLIGGTSIGSFIGALYAEERSAVRTKQRAREWAKSMNSVFETVLDLTYPITSMFSGSAFNTSINKVFQDKQIEDLWLPYFNVTTDITASAMRVHKDGSLWRYVRASMTLSGYLPPLCDPKDGNLLMDGGYINNLPADIARNMGAKTVVAIDVGSQDETDLCNYGDSLSGWWLLWKRLNPWAQKVKVPDMAEIQSRLAYVSCVRQLEVVKSSSYCEYIRPPIDRFKTMDFGKFDEIYDVGYQHGKVVFSGWCRGDIIEKMVKDRRLADFYESKRMDVLTCPSAGFTDLAEIVSRIEPAKSYLSDGYADGEESDYLTEYEEEGLDPARGEEELLAPAEWIHAEDEEKSLRHRKSLPQE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJC2</accession>
    <name>A0A452GJC2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="22">Dynein axonemal heavy chain 11</fullName>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Cilium axoneme</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the dynein heavy chain family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000001925.1">
        <property type="protein sequence ID" value="ENSGAGP00000001692.1"/>
        <property type="gene ID" value="ENSGAGG00000000806.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005930">
        <property type="term" value="C:axoneme"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030286">
        <property type="term" value="C:dynein complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005874">
        <property type="term" value="C:microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031514">
        <property type="term" value="C:motile cilium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045505">
        <property type="term" value="F:dynein intermediate chain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051959">
        <property type="term" value="F:dynein light intermediate chain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007018">
        <property type="term" value="P:microtubule-based movement"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000667">
        <property type="entry name" value="Dynein axonemal heavy chain 11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.2620:FF:000004">
        <property type="entry name" value="Dynein axonemal heavy chain 17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.920.30:FF:000003">
        <property type="entry name" value="Dynein axonemal heavy chain 17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000219">
        <property type="entry name" value="Dynein axonemal heavy chain 17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.20.180.20:FF:000001">
        <property type="entry name" value="Dynein axonemal heavy chain 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.8.710:FF:000002">
        <property type="entry name" value="dynein heavy chain 17, axonemal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.140.100:FF:000001">
        <property type="entry name" value="dynein heavy chain 17, axonemal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.1120:FF:000002">
        <property type="entry name" value="Dynein heavy chain 9, axonemal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.472.130:FF:000001">
        <property type="entry name" value="Dynein, axonemal, heavy chain 9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.2620">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.472.130">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.8.710">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.1120">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.920.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.140.100">
        <property type="entry name" value="Dynein heavy chain, N-terminal domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.20.180.20">
        <property type="entry name" value="Dynein heavy chain, N-terminal domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035699">
        <property type="entry name" value="AAA_6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026983">
        <property type="entry name" value="DHC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041589">
        <property type="entry name" value="DNAH3_AAA_lid_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042222">
        <property type="entry name" value="Dynein_2_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043157">
        <property type="entry name" value="Dynein_AAA1S"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041466">
        <property type="entry name" value="Dynein_AAA5_ext"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013602">
        <property type="entry name" value="Dynein_heavy_linker"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013594">
        <property type="entry name" value="Dynein_heavy_tail"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042228">
        <property type="entry name" value="Dynein_linker_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45703:SF12">
        <property type="entry name" value="DYNEIN AXONEMAL HEAVY CHAIN 11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45703">
        <property type="entry name" value="DYNEIN HEAVY CHAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12774">
        <property type="entry name" value="AAA_6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12775">
        <property type="entry name" value="AAA_7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17857">
        <property type="entry name" value="AAA_lid_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08385">
        <property type="entry name" value="DHC_N1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08393">
        <property type="entry name" value="DHC_N2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17852">
        <property type="entry name" value="Dynein_AAA_lid"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="13" id="KW-0966">Cell projection</keyword>
    <keyword evidence="10" id="KW-0969">Cilium</keyword>
    <keyword evidence="9 14" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="12" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="8" id="KW-0243">Dynein</keyword>
    <keyword evidence="4" id="KW-0493">Microtubule</keyword>
    <keyword evidence="11" id="KW-0505">Motor protein</keyword>
    <keyword evidence="6" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="Dynein heavy chain tail" evidence="15">
        <location>
            <begin position="44"/>
            <end position="336"/>
        </location>
    </feature>
    <feature type="domain" description="Dynein heavy chain tail" evidence="15">
        <location>
            <begin position="354"/>
            <end position="509"/>
        </location>
    </feature>
    <feature type="domain" description="Dynein heavy chain linker" evidence="16">
        <location>
            <begin position="1004"/>
            <end position="1409"/>
        </location>
    </feature>
    <feature type="domain" description="Dynein heavy chain hydrolytic ATP-binding dynein motor region" evidence="17">
        <location>
            <begin position="1543"/>
            <end position="1869"/>
        </location>
    </feature>
    <feature type="domain" description="Dynein heavy chain AAA 5 extension" evidence="18">
        <location>
            <begin position="2027"/>
            <end position="2145"/>
        </location>
    </feature>
    <feature type="domain" description="Dynein heavy chain 3 AAA+ lid" evidence="19">
        <location>
            <begin position="2362"/>
            <end position="2441"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="14">
        <location>
            <begin position="970"/>
            <end position="997"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004430"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008887"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023017"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023069"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="Pfam" id="PF08385"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF08393"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF12774"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF17852"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="Pfam" id="PF17857"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001692.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="22">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="2524" mass="294360" checksum="360CD4949669C10B" modified="2019-05-08" version="1">GHGALRVVVEWGGKRRWVLRWGGGRAWDRARGRKQSYLRFYHFLFTALMEAQDIELYLRPLRRHIQFLQETEFPKIHSLIPPLFHTICLIWSHSQFYSVPARIIVLLQEFCNLFIDQARSYLSPEDLLKGEIEETLEHVQIAVNTLRSFKNFFFSHREKLASYFTNSKEFKPWDFQSNMVFARFDLFLNRLVKIEDIFVVMLEFQKLEKLEFGGVKGKTLSEQIYRMNEEFIESCKVFKEKTYDPSDYNNVEFEEDYVEFKAKILDFDRRLGSILCVGFLDCSGLESAFKLLAIFGYFLEKPVIMEIFSPNYIILLQMFDEELDNCKHLYDEHIRQVCSKAKIKAIFFLVFVRILQDPETALVYQKYIETLTLIDQFEEQIYCKWKSQVDEVYQFNLDQPLIKRNPENGLLSVNFDSKLVALLREVTYLLKLNRSDIPDSALEFYSMNVFFHQQYIGNLELIVQWYNKLKQTVLEVEYPLIETELEAIDGQLRETEEVFTWQKENCWKYIEQVKAAVYDLEQRVQQSKENIKTIQQIMNAWAEHTLFSRKDDRKDALLSLEDKEDRLAKKYKLLKEDGYKIHQLAEVISENLKLFKADSSSDSWKIYIEYIDDIIVDGFFNTLMHNLDFFLENTEKGLKPAPLFQAQMILSAPEIEFKPSLDKEAGDGFYDLVEELLRQHPLPGDLNYINSLKKEQEYLWHLRDQVRLADVIRKALDYRNSLDTYAYLWVDDRAEFMKQFLLYGHVLTSEEIDSHADEGAPEQPPTTEQFKEQIDIYENLYIQMSKFEDFRTFESWFKVDMKPFKMSLLNIIKKWSWMFKEHLLKFVIGSLTDLEEFIKVTDAGLQKEVNEGDYNGLVEIMGHLLAVRDRQASTDELFEPLKETITLLESYGQKMSDQVYVQLEELPEKWNTTKKIAASIKHEVAPLQTSEVTLIRKKFALFDGKQIEFRERFRVEAPFCFDAKYPYILLDKENRELEALEEEMLQMQKSANLFEVVVPDYKQLKQCRKEIRLLKEVWDINIYVTSSIGDWTKTQWRQINVEQMDAELRRFAKEVWSLDKEIRSWDVYIGLELTVKNLMTSLRAVTELQNPAIRERHWHQLMNATDVRLSITEGTTLADLLGLQLHKVEDEVRSIVDKAVKELGTEKILTEISQTWATVEFCYEEHYRTRTPLLKCDEQLFETLENNQVQLQTILQSKYVEYFIEQVTSWQRTLNVADSVIFIWMEIQRTWSHLESIFIGSDDIRTKLPKDAKRFDGVDADFKELMMETANTKNVLEATTKPNLYEKLESLQRRLSLCEKALAEYLETKCLAFPRFYFVSSADLLDILSKGTQPRQVTRHLTKLFDNIADLQFQDNKEKSTDIALGMYSKEKEYVPFHAECECIGQVETWLQQLEETMYKTVHRHIMEAIAAYEEKPREQWIFDYPAQVALTSSQIWWTTDVGIAFNRLEEGFETALKDYNKKQIAQLNALIALLLGELASGDRQKIMTICTIDVHARDVVAKLVAQKVTSSQAFAWLSQLRHRWDEAQKHCLANICDAQFQYFYEYLGNTSRLVITPLTDRCYITLTQSLHLTMSGAPAGPTGTGKTETTKDLGHALGMMVYVFNCSEQMDYKSIGNIYKGLVQTGAWGCFDEFNRISVEVLSVVAVQVKTIHDAIRNKRKKFLFLGENITLKTSVGIFITMNPGYAGRTELPENLKALFRPCAMVVPDIELICEIMLVAEGFTDARLLARKFITLYTLCRELLSKQDHYDWGLRAVKSVLVVAGSLKRGDKNRPEDQVLMRTLRDFNLPKIVTDDIPIFMGLISDLFPALDVPRRRDLQFEQMVKQSTLELHLQPEESFIFKVVQLEELLAVRHSVFVVGNAGTGKSKVLRTLNRTYVNMKQKLVWNDLNPKAVTTDELFGFIHHATREWKDGLFSSLLREQANITQDGPKWIVLDGDIDPMWIESLNTVMDDNKVLTLASNERVPLTPSMRLLFEIHHLKTATPATVSRAGILYVNPQDLGWNPYVASWIETRSHQSEKANLTILFDKYVPPCLDQLRTSFKTITPIPENSMVQTLCSLLDCLLTPENVPPDCPRELYEIYFVFACVWAFGGALFQDQLSNYQAEFSRWWLKEMKAVKFPSQGTIFDYYLDPKTRKFLPWTDKVPRFDMDPETPLQAVLVHTSETTRLRYFTDLLLERGKPVMLVGNAGVGKTVFVGDRLAAFPDDYMVSNVLFNYYTTSSVLQKILEKSLEKKAGRNYGPVGNKKLVYFIDDMNMPEMDVYGTVQPHTLIRQHIDYGHWYDRQKLTVKEIHNCHYVACMNPTAGNFTINPRLQRHFTVFALNFPSVDALNTIYSKILGFHFQRHVFNPSVLKNAPAMIQASIWLHQTMVQNFLPTAVKFHYIFNLRDLSNIFQGILFATPECLREPNDLVCLWLHESSRVYGDKLVEPKDCHFFHHLLRQHPLIYCHFANGVASPHYMPVKEWEVLRKILTETLESYNELNASMNLVLFEDAMQHKTIQQCSGLTSSLPVKQGFNLFVFV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZX0</accession>
    <name>A0A452GZX0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">Receptor-type tyrosine-protein phosphatase F</fullName>
            <ecNumber evidence="3">3.1.3.48</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>O-phospho-L-tyrosyl-[protein] + H2O = L-tyrosyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:10684"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="EC" id="3.1.3.48"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the protein-tyrosine phosphatase family. Receptor class 2A subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.48" evidence="3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008891.1">
        <property type="protein sequence ID" value="ENSGAGP00000007706.1"/>
        <property type="gene ID" value="ENSGAGG00000005093.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="CDD" id="cd05738">
        <property type="entry name" value="IgI_2_RPTP_IIa_LAR_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05739">
        <property type="entry name" value="IgI_3_RPTP_IIa_LAR_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000010">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000027">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000036">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000066">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000144">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000015">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000082">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000128">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000098">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase F isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000353">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase F isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013098">
        <property type="entry name" value="Ig_I-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050713">
        <property type="entry name" value="RTP_Phos/Ushers"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46957">
        <property type="entry name" value="CYTOKINE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46957:SF9">
        <property type="entry name" value="PROTEIN-TYROSINE-PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07679">
        <property type="entry name" value="I-set"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00014">
        <property type="entry name" value="FNTYPEIII"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="13" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="15" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="4" id="KW-0358">Heparin-binding</keyword>
    <keyword evidence="8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="16" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="12 20" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="14" id="KW-0675">Receptor</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="5 20" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 20" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="1161"/>
            <end position="1184"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="21">
        <location>
            <begin position="42"/>
            <end position="137"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="21">
        <location>
            <begin position="145"/>
            <end position="227"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="234"/>
            <end position="325"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="330"/>
            <end position="424"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="428"/>
            <end position="517"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="524"/>
            <end position="621"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="626"/>
            <end position="725"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="726"/>
            <end position="820"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="824"/>
            <end position="916"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="920"/>
            <end position="1004"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="606"/>
            <end position="627"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1270"/>
            <end position="1297"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004167"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010504"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00013064"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022674"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00044158"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00051722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007706.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1334" mass="147121" checksum="CAFF66B410B06F98" modified="2019-05-08" version="1">SSLTSCTMQATESHPPIPVSNLYLICCMLFPVAEDHLPAGFPTIDMGPQLKVVEKARTATMLCAASGNPDPEISWFKDFLPVDTATSNGRIKQLRSGGTPIRGALQIENSEESDQGKYECVATNSAGTRYSAPANLYVRVRRVAPRFSIPPSNHEVMPGGSVNLTCVAVGAPMPYVKWMAGVEELTKEDEMPVGRNVLELNNIVQSASYTCVAMSSLGMIEATAQITVKALPKPPTEPLVTETTATSVTLTWDSGNSDPISYYVIQYKPKSWEEGLFQEVDGVATTRYSIGGLSPFSEYEFRIIAVNNIGRGPPSELVEARTGEQAPSSPPLKVQARMLSASTMLVQWEQPEEPNGQIRGYRVYYTTDPHLPLSMWQKHNTDDSHLTTVGSLITGTTYSIRVLAFTSVGDGPPSDIIQVKTQQGVPAQPADFQAEAESDTRILLTWLPALQERITKYELLYWEGEDGSQRKVNFDPSSSYAVEGLKPDTLYKFQLGARSELGLGVYTPVLEARTAQSSKSPSAPPQEVECVSTSSTTIRVSWVPPPAQSRNGVITQYSIAYQAVDAEDGAKHVVDGIGRERSSWEIKDLEKWTEYKVWVRAHTDVGPGPESSPVHVRTDEDVPSAPPRKVEVESLNSTAIRVSWKLPLSNKQHGQIRGYQVTYVKLENNEPRGQPVIKDVMLAEAQETIIGGLLPETTYSITVAAYTTKGDGARSKPKVVTTTGAVPGKPSMLISTTAMNTALIQWHPPKEMVGELLGYRLQYKRTDEEKMNIMDFEKNVHHYTVTSLHKGATYIFKLSAKNRAGPGEEFEKEITTTEDVPSGFPQNLRVVGLTTSTTEVAWDPPVLAERNGRITNYTVMYRDINSQQELTNITRETSIVLTNLRPDTTYDIKVRACNSKGVGPLSPSIQSRTMPVEQVFAKNFRVNAVMKTSVLLSWEVPDSYKSAVPFKILYNSQSVEVDGHSMKKLISDLQPDTEYSFVLMNRGSSAGGLQHRVSIHTAPDVLQSKPISTNKYMQEGKFTLTLPKVQTSAPVRWYYIVVVPVDQSTSSRTARWRTPDEMELDQLLEAISQGRRQRRQADRLKPYIAALVDMLPETFTLGDEKTYKGFYNKPLSEDLSYRCFVLASLEDGDTKRYAASPYSDEIVMEVSSAKQQDEPEMLWVMGPVLAVILIIIIVIAILLFKRLKANDGLRFSQEYEVSCREGDRCAWHCPEPGVAGRRWAMAASPRGGSVDSFHAHSFSVHLPRGYPLSWLVPQLQPPAASLVTRRSCPQAPRPGLQRPGVPHXGSPYSCSADTPQSHLASPCPPTHTLALLIPLSPALPLPGLPLVLLC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZX1</accession>
    <name>A0A452GZX1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Alpha-2-macroglobulin like 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the protease inhibitor I39 (alpha-2-macroglobulin) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZX1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000009020.1">
        <property type="protein sequence ID" value="ENSGAGP00000007823.1"/>
        <property type="gene ID" value="ENSGAGG00000006228.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000009023.1">
        <property type="protein sequence ID" value="ENSGAGP00000007826.1"/>
        <property type="gene ID" value="ENSGAGG00000006228.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004867">
        <property type="term" value="F:serine-type endopeptidase inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd02897">
        <property type="entry name" value="A2M_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000312">
        <property type="entry name" value="Alpha-2-macroglobulin like 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.50.10.20:FF:000001">
        <property type="entry name" value="CD109 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.50.10.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.20.130.20">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.1540">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1930">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050473">
        <property type="entry name" value="A2M/Complement_sys"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011625">
        <property type="entry name" value="A2M_N_BRD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041813">
        <property type="entry name" value="A2M_TED"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047565">
        <property type="entry name" value="Alpha-macroglob_thiol-ester_cl"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011626">
        <property type="entry name" value="Alpha-macroglobulin_TED"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014756">
        <property type="entry name" value="Ig_E-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001599">
        <property type="entry name" value="Macroglobln_a2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019742">
        <property type="entry name" value="MacrogloblnA2_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040839">
        <property type="entry name" value="MG4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008930">
        <property type="entry name" value="Terpenoid_cyclase/PrenylTrfase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11412:SF185">
        <property type="entry name" value="ALPHA-2-MACROGLOBULIN-LIKE PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11412">
        <property type="entry name" value="MACROGLOBULIN / COMPLEMENT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00207">
        <property type="entry name" value="A2M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07703">
        <property type="entry name" value="A2M_BRD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17789">
        <property type="entry name" value="MG4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07678">
        <property type="entry name" value="TED_complement"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01360">
        <property type="entry name" value="A2M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01359">
        <property type="entry name" value="A2M_N_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01419">
        <property type="entry name" value="Thiol-ester_cl"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81296">
        <property type="entry name" value="E set domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48239">
        <property type="entry name" value="Terpenoid cyclases/Protein prenyltransferases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00477">
        <property type="entry name" value="ALPHA_2_MACROGLOBULIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="8" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="4" id="KW-0646">Protease inhibitor</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0964">Secreted</keyword>
    <keyword evidence="6" id="KW-0722">Serine protease inhibitor</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <feature type="domain" description="Alpha-2-macroglobulin bait region" evidence="9">
        <location>
            <begin position="143"/>
            <end position="293"/>
        </location>
    </feature>
    <feature type="domain" description="Alpha-2-macroglobulin" evidence="10">
        <location>
            <begin position="448"/>
            <end position="537"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010952"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022690"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="SMART" id="SM01359"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="SMART" id="SM01360"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007826.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="13">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1007" mass="109951" checksum="99FDA563768C8154" modified="2019-05-08" version="1">MSLQAKASLVEEGTGVELNATKSCDIVSEIATVTFEDADNTYKAGIPYTGKMLLKAADGSALKNEKLQLFVSYGDVRRDQTFLTDESGRASFKLDTSGWTGRVTLKGHFKQVDRSSVFGRVSPSYPDAHRNLEPFYSKSQSFLKIRWLGGVLPCDQAQQLQVDYIIARETLGNGSRSLDFIFLVVAKGAIARILHKGLDLREGAGLKGSFSVELPIGADLAPAAKVLGYTVLPDGEMAADSARLQLAKCLPNKVKLAFSQDRALPGSELQLRLQAAPGSLCAVRAVDQSVLLMKPEAELSVDTVYNLLPRFPQGDYPAAVREHESGDCPYFSEYPEYMGPWRRRHRKPQSVQFSDSDSYRLFKDAALKILTNANTKKPFICPDIPIFRYSSQPDFDLGLRGSPGDPFYRREILRENVLVARGGLGNALPEPEPSTEEAPAPRAYFPETWLWDLVPVGEGGSKDVPVSVPDTITEWKAGMFCTAEVGFGLSPTATFTAFKPFFVELALPYSVIRGEAFALKATVFNYLRQCIKVRVTLAESAQFQVQAGEDDAYTSCLCADEGKTFPWEVTASSLGEVNFTVSTEALHTEELCGNELAVVPAQGRVDTVIKPLLVQPGGILEEKAYSSLLCQEASEEISLQLPANVLAGSERAHVTVLGDIMGTALQNIDRLLAMPYGCGEQNMVRFAPNIYIQQYLEKSGQLSPEIRDKAKGFLLSGYQRELLYKHKDGSYSAFGERDATGNTWLTAFVLKSFGQARPYISIDEKHLEDALRWLQSHQLESGCFRSVGKLLNNALQGGVVDELSLSAYITAALLELGQPLMDPMVTRALQCLRESSTSDLYTQALLAYAFGLVGSTELRDALLQRLAQHSVSAGGQLYWQRKVKALPSPYGNQAPSAEVEMTAYVLLAYLSLPNVSAANMATAAQIVRWLSKQQNPYGGFASTQDTVVALQALAKYAALTYSASGAVSVTVSSQAGAQQQFHVDNANRLVLQQAALQEIPGQYTVQA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZX3</accession>
    <name>A0A452GZX3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">Receptor-type tyrosine-protein phosphatase F</fullName>
            <ecNumber evidence="3">3.1.3.48</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>O-phospho-L-tyrosyl-[protein] + H2O = L-tyrosyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:10684"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="EC" id="3.1.3.48"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the protein-tyrosine phosphatase family. Receptor class 2A subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.48" evidence="3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008931.1">
        <property type="protein sequence ID" value="ENSGAGP00000007743.1"/>
        <property type="gene ID" value="ENSGAGG00000005093.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="CDD" id="cd05738">
        <property type="entry name" value="IgI_2_RPTP_IIa_LAR_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05739">
        <property type="entry name" value="IgI_3_RPTP_IIa_LAR_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000010">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000027">
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        <property type="entry name" value="receptor-type tyrosine-protein phosphatase F isoform X1"/>
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        <property type="match status" value="9"/>
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        <property type="entry name" value="FN3_dom"/>
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        <property type="entry name" value="FN3_sf"/>
    </dbReference>
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        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
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        <property type="entry name" value="Ig-like_dom_sf"/>
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    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
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        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050713">
        <property type="entry name" value="RTP_Phos/Ushers"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46957">
        <property type="entry name" value="CYTOKINE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46957:SF9">
        <property type="entry name" value="PROTEIN-TYROSINE-PHOSPHATASE"/>
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        <property type="entry name" value="fn3"/>
        <property type="match status" value="6"/>
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        <property type="entry name" value="I-set"/>
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    <dbReference type="PRINTS" id="PR00014">
        <property type="entry name" value="FNTYPEIII"/>
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        <property type="match status" value="7"/>
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        <property type="entry name" value="IG"/>
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        <property type="entry name" value="IGc2"/>
        <property type="match status" value="2"/>
    </dbReference>
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        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="4"/>
    </dbReference>
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        <property type="match status" value="2"/>
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        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="13" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="15" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="4" id="KW-0358">Heparin-binding</keyword>
    <keyword evidence="8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="16" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="12 20" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="14" id="KW-0675">Receptor</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="5 20" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 20" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="1090"/>
            <end position="1113"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="21">
        <location>
            <begin position="81"/>
            <end position="170"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="21">
        <location>
            <begin position="178"/>
            <end position="260"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="267"/>
            <end position="358"/>
        </location>
    </feature>
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        <location>
            <begin position="363"/>
            <end position="457"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="461"/>
            <end position="550"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="555"/>
            <end position="654"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="655"/>
            <end position="749"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="753"/>
            <end position="845"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="849"/>
            <end position="933"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1261"/>
            <end position="1285"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004167"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010504"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00013064"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022674"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
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        </source>
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            <dbReference type="ARBA" id="ARBA00044158"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00051722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007743.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1285" mass="142893" checksum="895AE3CA343BE45E" modified="2019-05-08" version="1">FSVCVCFYLALVLAACLCSMQNTSLFLHKDKHLLLEFNNKRSSLHISENHGQAIDSNRVTFSSFCCMLFPVAEDHLPAGFPTIDMGPQLKVVEKARTATMLCAASGNPDPEISWFKDFLPVDTATSNGRIKQLRSGALQIENSEESDQGKYECVATNSAGTRYSAPANLYVRVRRVAPRFSIPPSNHEVMPGGSVNLTCVAVGAPMPYVKWMAGVEELTKEDEMPVGRNVLELNNIVQSASYTCVAMSSLGMIEATAQITVKALPKPPTEPLVTETTATSVTLTWDSGNSDPISYYVIQYKPKSWEEGLFQEVDGVATTRYSIGGLSPFSEYEFRIIAVNNIGRGPPSELVEARTGEQAPSSPPLKVQARMLSASTMLVQWEQPEEPNGQIRGYRVYYTTDPHLPLSMWQKHNTDDSHLTTVGSLITGTTYSIRVLAFTSVGDGPPSDIIQVKTQQGVPAQPADFQAEAESDTRILLTWLPALQERITKYELLYWEGEDGSQRKVNFDPSSSYAVEGLKPDTLYKFQLGARSELGLGVYTPVLEARTAQSMPSAPPRKVEVESLNSTAIRVSWKLPLSNKQHGQIRGYQVTYVKLENNEPRGQPVIKDVMLAEAQETIIGGLLPETTYSITVAAYTTKGDGARSKPKVVTTTGAVPGKPSMLISTTAMNTALIQWHPPKEMVGELLGYRLQYKRTDEEKMNIMDFEKNVHHYTVTSLHKGATYIFKLSAKNRAGPGEEFEKEITTTEDVPSGFPQNLRVVGLTTSTTEVAWDPPVLAERNGRITNYTVMYRDINSQQELTNITRETSIVLTNLRPDTTYDIKVRACNSKGVGPLSPSIQSRTMPVEQVFAKNFRVNAVMKTSVLLSWEVPDSYKSAVPFKILYNSQSVEVDGHSMKKLISDLQPDTEYSFVLMNRGSSAGGLQHRVSIHTAPDVLQSKPISTNKYMQEGKFTLTLPKVQTSAPVRWYYIVVVPVDQSTSSRTARWRTPDEMELDQLLEAISQGRRQRRQADRLKPYIAALVDMLPETFTLGDEKTYKGFYNKPLSEDLSYRCFVLASLEDGDTKRYAASPYSDEIVMEVSSAKQQDEPEMLWVMGPVLAVILIIIIVIAILLFKRKRANSPSSKDEHSIGLKDSLLAHSSDPVEMRRLNYQTPGMRDHPPIPVTDLADNIDRLKANDGLRFSQEYEVSCREGDRCAWHCPEPGVAGRRWAMAASPRGGSVELLPWVTELYRWGPTLGRVLGQQSACWPWWCQEPSLAQPGYRRRERESRGVPGSQNKRSDGCGAV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZX5</accession>
    <name>A0A452GZX5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">mannosyl-oligosaccharide 1,3-1,6-alpha-mannosidase</fullName>
            <ecNumber evidence="16">3.2.1.114</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="17">Mannosyl-oligosaccharide 1,3-1,6-alpha-mannosidase</fullName>
        </alternativeName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">Catalyzes the first committed step in the biosynthesis of complex N-glycans. It controls conversion of high mannose to complex N-glycans; the final hydrolytic step in the N-glycan maturation pathway.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>N(4)-{beta-D-GlcNAc-(1-&gt;2)-alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;6)]-alpha-D-Man-(1-&gt;6)]-beta-D-Man-(1-&gt;4)-beta-D-GlcNAc-(1-&gt;4)-beta-D-GlcNAc}-L-asparaginyl-[protein] + 2 H2O = 2 alpha-D-mannopyranose + an N(4)-{beta-D-GlcNAc-(1-&gt;2)-alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;6)]-beta-D-Man-(1-&gt;4)-beta-D-GlcNAc-(1-&gt;4)-beta-D-GlcNAc}-L-asparaginyl-[protein]</text>
            <dbReference type="Rhea" id="RHEA:56052"/>
            <dbReference type="Rhea" id="RHEA-COMP:14368"/>
            <dbReference type="Rhea" id="RHEA-COMP:14369"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:28729"/>
            <dbReference type="ChEBI" id="CHEBI:60615"/>
            <dbReference type="ChEBI" id="CHEBI:60625"/>
            <dbReference type="EC" id="3.2.1.114"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <comment type="pathway">
        <text evidence="3">Protein modification; protein glycosylation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Golgi apparatus membrane</location>
            <topology evidence="2">Single-pass type II membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the glycosyl hydrolase 38 family.</text>
    </comment>
    <dbReference type="EC" id="3.2.1.114" evidence="16"/>
    <dbReference type="AlphaFoldDB" id="A0A452GZX5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008831.1">
        <property type="protein sequence ID" value="ENSGAGP00000007649.1"/>
        <property type="gene ID" value="ENSGAGG00000002370.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000139">
        <property type="term" value="C:Golgi membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030246">
        <property type="term" value="F:carbohydrate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004572">
        <property type="term" value="F:mannosyl-oligosaccharide 1,3-1,6-alpha-mannosidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006013">
        <property type="term" value="P:mannose metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006491">
        <property type="term" value="P:N-glycan processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1270.50:FF:000001">
        <property type="entry name" value="Alpha-mannosidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.70.98.30:FF:000002">
        <property type="entry name" value="Alpha-mannosidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.20.110.10:FF:000003">
        <property type="entry name" value="Alpha-mannosidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.20.110.10">
        <property type="entry name" value="Glycoside hydrolase 38, N terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.50">
        <property type="entry name" value="Glycoside hydrolase family 38, central domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.70.98.30">
        <property type="entry name" value="Golgi alpha-mannosidase II, domain 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011013">
        <property type="entry name" value="Gal_mutarotase_sf_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011330">
        <property type="entry name" value="Glyco_hydro/deAcase_b/a-brl"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011682">
        <property type="entry name" value="Glyco_hydro_38_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015341">
        <property type="entry name" value="Glyco_hydro_38_cen"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037094">
        <property type="entry name" value="Glyco_hydro_38_cen_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000602">
        <property type="entry name" value="Glyco_hydro_38_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027291">
        <property type="entry name" value="Glyco_hydro_38_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028995">
        <property type="entry name" value="Glyco_hydro_57/38_cen_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050843">
        <property type="entry name" value="Glycosyl_Hydrlase_38"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11607">
        <property type="entry name" value="ALPHA-MANNOSIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11607:SF57">
        <property type="entry name" value="ALPHA-MANNOSIDASE 2X"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09261">
        <property type="entry name" value="Alpha-mann_mid"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07748">
        <property type="entry name" value="Glyco_hydro_38C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01074">
        <property type="entry name" value="Glyco_hydro_38N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00872">
        <property type="entry name" value="Alpha-mann_mid"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF88688">
        <property type="entry name" value="Families 57/38 glycoside transferase middle domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF74650">
        <property type="entry name" value="Galactose mutarotase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF88713">
        <property type="entry name" value="Glycoside hydrolase/deacetylase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="13" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="14" id="KW-0326">Glycosidase</keyword>
    <keyword evidence="11" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="7" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="12 19" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0735">Signal-anchor</keyword>
    <keyword evidence="5 19" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="10 19" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="8" id="KW-0862">Zinc</keyword>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="7"/>
            <end position="26"/>
        </location>
    </feature>
    <feature type="domain" description="Glycoside hydrolase family 38 central" evidence="20">
        <location>
            <begin position="505"/>
            <end position="591"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004323"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004922"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00009792"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023295"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00059516"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00066412"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00083602"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00093232"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="SMART" id="SM00872"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007649.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1070" mass="120732" checksum="FF4513D53D1936B3" modified="2019-05-08" version="1">MKLKKQVTVCGAAIFCVAVFSLYLMLDRAGGPLVPRRLLSPCVCNLQSQISVLQNRIEQLEQLLEENHEIISHIKESVLELTAHVDGQPVLLPYRTPNGSWVLPPESRPSFFSISPQDCQFALGSGGQKPDLQMLAVASVLPFDNQDGGVWKQGFDITYEPHDWDTEPLQVFVVPHSHNDPGWIKTFEKYYADQTQHILNSMVLKLQEDPRRRFIWSEISFFSKWWDNISTQKRAAVRRLVGGGQLEMVTGGWVMPDEANSHYFAMIDQLIEGHQWLEKNIGVTPRSGWAVDPFGHSATMPYLLRRANLTSMLIQRVHYTIKKHFAATRSLEFMWRQTWDPDSSTDIFCHMMPFYSYDVPHTCGPDPKICCQFDFKRLPGGRINCPWKVPPRAITEANVAERAHLLLDQYRKKSKLFRSKVLLVPLGDDFRYDKPQEWDAQFLNYQRLFDFLNAHPSLHVQAQFGTLSDYFDALSKRTGIIPGLKPPGFPVLSGDFFSYADREDHYWTGYYTSRPFYKSLDRVLEAHLRGAEILYSLALSHARHAGMDSRYPLSDYALLTEARRALGLFQHHDAITGTAKEAVVVDYALSLLVNSPRVRVLSEEGQPLAIQLSAQWSSATDMVPDVYQVREELAASPHPELGETYQTHSALGSSPHPPPCTLYTPISHIHPPLHTPHIGGRAGGSIVCLLPTHSLPAWPASLQSVRRAGEEHRLSSEFLIYGTRSAKDKSGAYLFLPDGDAKPYAPKEPPVVRVTEGPFFSEVASYYQHVQQVVRLYNVPGVEGLSLDVSCLVDIRDHVNKEMALRLSTTIASEDTFFTDLNGFQIQPRRFLKKLPLQANFYPMPAMAYIQDKQSRLTLHTAQALGVSSLHSGQLEVILDRRLMQDDNRGLGQGLKDNKRTCNRFRLLLERRTTANRVQDNRPISFPSLLSHMTSMHLNAEVLAMPVAREKPAPPALRSFRPLSATVPCDFHILNLRTLQAEVRDDLLPSAEIALILHRKGFDCGLEAKHLGFNCTTSQGKVGLLFHGLELGSLQPTSLTLMYPLGAASPNSTVIHMDPMEIATFRIHFE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZX8</accession>
    <name>A0A452GZX8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">BTB domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GZX8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008903.1">
        <property type="protein sequence ID" value="ENSGAGP00000007716.1"/>
        <property type="gene ID" value="ENSGAGG00000005859.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="CDD" id="cd18526">
        <property type="entry name" value="BACK_ABTB2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18350">
        <property type="entry name" value="BTB_POZ_ABTB2_BPOZ2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000045">
        <property type="entry name" value="Ankyrin repeat and BTB/POZ domain-containing protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.710.10:FF:000030">
        <property type="entry name" value="Ankyrin repeat and BTB/POZ domain-containing protein BTBD11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.710.10">
        <property type="entry name" value="Potassium Channel Kv1.1, Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048063">
        <property type="entry name" value="ABTB2_BTB_POZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052089">
        <property type="entry name" value="Ankyrin-BTB/POZ_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000210">
        <property type="entry name" value="BTB/POZ_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011333">
        <property type="entry name" value="SKP1/BTB/POZ_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46071">
        <property type="entry name" value="ANKYRIN REPEAT AND BTB/POZ DOMAIN-CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46071:SF3">
        <property type="entry name" value="ANKYRIN REPEAT AND BTB_POZ DOMAIN-CONTAINING PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00023">
        <property type="entry name" value="Ank"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00651">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00225">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54695">
        <property type="entry name" value="POZ domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50097">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2 3" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0677">Repeat</keyword>
    <feature type="repeat" description="ANK" evidence="3">
        <location>
            <begin position="306"/>
            <end position="338"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="3">
        <location>
            <begin position="352"/>
            <end position="384"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="3">
        <location>
            <begin position="391"/>
            <end position="417"/>
        </location>
    </feature>
    <feature type="domain" description="BTB" evidence="4">
        <location>
            <begin position="629"/>
            <end position="698"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50097"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007716.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="809" mass="90790" checksum="6B5F543598685A87" modified="2019-05-08" version="1">GEWDEALSSPFSEGWLLCYGALQKGEATRGKGDGSQDSVRDSVPFNGEKHTHLFPSIPEWWSIPLPLRFHLQQACSLLSPGVLSLPAYFSPYNDGSISSDGRADAYAQLELRTLEQSLLATCVGSISELSDLVSRAMHHMQCLTTLHPGMSPIRQTRQQQPITWSPDALHTLYYFLRCPQMESMENPNLDPPRMTLNNERPFMLLPPLMEWMRVAITYAEHRRSLTVDSDDIRQTARLLLPGLDCEPRQLKPECCFSSFRRLDAKAATERFQQDLGFRMLNCGRTDLINQAIEALGPDGVNTMDDQGMTPLMYACAAGDEAMVQMLIDAGANLDIPVPGSNPRYPSVHPDSRHWTALTFAVLHGHISVVQLLLDAGAHVEGSAVNSSEDNYAETPLQLASAAGNYELVSLLLSRGADPLLSMLDVNGMSSSLHEDMNCFSHSAAHGHRNVLRKLLTQPQQSKGDVLSLEEILAEGVESDTSSQDSSNEGQVRLCKTRMKALQEAMYYSAEHGYVDITMELRVLGIPWKLHVWIESLRTTFYQSRYSVVQNLLREFVTIKEEDYNEELVNEGLQLMFDILKTSKNDSINQLLAAIFTHCYGSSPIPNIPEIQKTLPARLDPHFLNNKEMSDVTFLVEGKLFYAHKVLLVTASNRFKTLMTNKTELDSQGSKTVEISDMKYNIFKMLMQYLYYGGTESMEIPTTDILELLSAASLFQLDGLQRHCEILCAQTISMESSVNIYKYAKIHSAPELASFCKGFFLKHMNSLLEQDSFRQLIYGRNSKVQGLDPLRDLQTTLASRLHSIYITSRV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZY6</accession>
    <name>A0A452GZY6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Zinc finger protein 148</fullName>
        </recommendedName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="16">Involved in transcriptional regulation. Represses the transcription of a number of genes including gastrin, stromelysin and enolase. Binds to the G-rich box in the enhancer region of these genes.</text>
    </comment>
    <comment type="subunit">
        <text evidence="17">Interacts with HNRNPDL. Interacts with the 5FMC complex; the interaction requires association with CHTOP. Interacts with CAVIN1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZY6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000009043.1">
        <property type="protein sequence ID" value="ENSGAGP00000007844.1"/>
        <property type="gene ID" value="ENSGAGG00000006250.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:1990837">
        <property type="term" value="F:sequence-specific double-stranded DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:004830">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000067">
        <property type="entry name" value="Vascular endothelial zinc finger 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000042">
        <property type="entry name" value="Zinc finger protein 148"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050636">
        <property type="entry name" value="C2H2-ZF_domain-containing"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47772:SF13">
        <property type="entry name" value="GASTRULA ZINC FINGER PROTEIN XLCGF49.1-LIKE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47772">
        <property type="entry name" value="ZINC FINGER PROTEIN 200"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="11" id="KW-0007">Acetylation</keyword>
    <keyword evidence="13" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="4" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="6" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="15" id="KW-0539">Nucleus</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0678">Repressor</keyword>
    <keyword evidence="14" id="KW-0804">Transcription</keyword>
    <keyword evidence="12" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="10" id="KW-0832">Ubl conjugation</keyword>
    <keyword evidence="9" id="KW-0862">Zinc</keyword>
    <keyword evidence="8 19" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="21">
        <location>
            <begin position="170"/>
            <end position="197"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="21">
        <location>
            <begin position="198"/>
            <end position="225"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="21">
        <location>
            <begin position="226"/>
            <end position="254"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="120"/>
            <end position="151"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="247"/>
            <end position="309"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="544"/>
            <end position="566"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="657"/>
            <end position="691"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="745"/>
            <end position="764"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="20">
        <location>
            <begin position="124"/>
            <end position="140"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="20">
        <location>
            <begin position="291"/>
            <end position="309"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="20">
        <location>
            <begin position="666"/>
            <end position="682"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022491"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00060069"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00064837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00067481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007844.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="764" mass="85160" checksum="2FB3989B28F7D616" modified="2019-05-08" version="1">MNIEDKLGGLFLKCGGIDQMQSSRAMVGMGAVSDQSSISGERQEAVLQDRTMAHQEILTTDEVLQESELRQQEMISHDELMVHEETVKNDDELEAQDRLPQGLQYAVNVPISVKQEITFTDASEQQKRDKKQIREPVDLQKKKKRKQRSPAKILTINEDGSLGLKTPKSHVCEHCNAAFRTNYHLQRHVFIHTGEKPFRCDECGMRFIQKYHMERHKRTHSGEKPYQCEYCLQYFSRTDRVLKHKRMCHESRDKKPNRSATKVGLLTSDEDPGFSTPMKDSSLPKKKRQKTEKAKSVDKEIGDKSEQKKDKNDYLPLYSASAKVKDEYMVAEYAVEMPHSSVSGSHLEDANSGEIHPPKLVLKKVTSKRSLKPPLEQSQTISPLSTYEENKVSKYAFDLVDKQGLLDADGNADIDQVDTLQEGPSKPVHSSTNYDDAMQFLKKKRYLQAASNNSREYALNVGTIASQPSVTQAAVASVIDESTTASILDSQALNVEIKSNHDKNVIPDEVLQTLLDHYSHKANGQHEISFNVTDTEVTSSISINSSEVSEVTQSETVGTSSQASSSDKASMLQEYSKFLQQALDRTSQNDAYLNNPSLNFVTDNQTLTSQPAFSSIDKQVYASIPVNSFRSGMNSPLRTTPDKSHFGLMVADSQHSFPFSGDEANHSSSSSTQDFLDQVTSQKKAEAQPVHQAYQMSSFEQPFRAQYHGARAGISPQFSTANGQVNLRGPGTSAEFPEFPLVNVNDNRAGMTSSPDATTGQTFG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLY6</accession>
    <name>A0A452HLY6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Tetraspanin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 7">Membrane</location>
            <topology evidence="1 7">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 7">Belongs to the tetraspanin (TM4SF) family.</text>
    </comment>
    <comment type="caution">
        <text evidence="7">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLY6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018233.1">
        <property type="protein sequence ID" value="ENSGAGP00000015958.1"/>
        <property type="gene ID" value="ENSGAGG00000012007.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1450.10">
        <property type="entry name" value="Tetraspanin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018499">
        <property type="entry name" value="Tetraspanin/Peripherin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000301">
        <property type="entry name" value="Tetraspanin_animals"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008952">
        <property type="entry name" value="Tetraspanin_EC2_sf"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00335">
        <property type="entry name" value="Tetraspanin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF002419">
        <property type="entry name" value="Tetraspanin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48652">
        <property type="entry name" value="Tetraspanin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5 7" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 7" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="12"/>
            <end position="34"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="60"/>
            <end position="84"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="174"/>
            <end position="200"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="6">
        <location>
            <begin position="130"/>
            <end position="160"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="6">
        <location>
            <begin position="131"/>
            <end position="148"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PIRSR" id="PIRSR002419-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU361218"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015958.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="217" mass="24807" checksum="933FA45983BC0777" modified="2019-05-08" version="1">MGRRHWVRTTKCQLLVTSLFVMLLGLAVTIMTVITRYGAHFKIISGVSYEANPYRIFHNVAFYSGICLSMILVLAALLSTAATVRESVCLMAMVEDAMMDVYDSVYEEVRRNLSSFRRQELATIHETFLCCGKQSPFGETGGIENETCPSEHMGTAREDCLQEIQDFLKKHMDFVSALLIITICFMVYGMILTSFLWFSIHFSNNLDRRGKYIFRET</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="22" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLY7</accession>
    <name>A0A452HLY7_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="1">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="1 2">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="2" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="1" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="1" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HLY7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018229.1">
        <property type="protein sequence ID" value="ENSGAGP00000015954.1"/>
        <property type="gene ID" value="ENSGAGG00000012004.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000313" key="1">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015954.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="220" mass="23514" checksum="942ECE40EFC1E505" modified="2019-05-08" version="1">MTFFRWDARARARLLVVIKDPVALVRRAQFSRQIPANCLLLWKFQLNAVCFVTSGPKLPCNLDVRLLNCCCVPPQQRLTIAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSTVSPGPSGSRWLLSCGLIAG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="22" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLZ1</accession>
    <name>A0A452HLZ1_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="1">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="1 2">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="2" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="1" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="1" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HLZ1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018234.1">
        <property type="protein sequence ID" value="ENSGAGP00000015959.1"/>
        <property type="gene ID" value="ENSGAGG00000012004.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000313" key="1">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015959.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="202" mass="21476" checksum="3AA4468690272B99" modified="2019-05-08" version="1">MTFFRWDARARARLLVVIKDPVALVRRAQFSRQIPANCLLLWKFQLNAVCFVTSGPKLPCNLDVRLLNCCCVPPQHAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRTVSPGPSGSRWLLSCGLIAG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="22" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLZ2</accession>
    <name>A0A452HLZ2_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="1">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="1 2">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="2" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="1" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="1" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HLZ2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018238.1">
        <property type="protein sequence ID" value="ENSGAGP00000015963.1"/>
        <property type="gene ID" value="ENSGAGG00000012004.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000313" key="1">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015963.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="208" mass="21853" checksum="B3907F02E866DC54" modified="2019-05-08" version="1">MSAWPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAPKLRAWCPPLPCSSAHRLPRARAAPDQSSRLCTSRWWQRRESQAPRAGSGCCQCSRPVGCATSI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLZ4</accession>
    <name>A0A452HLZ4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">non-specific serine/threonine protein kinase</fullName>
            <ecNumber evidence="1">2.7.11.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="EC" id="2.7.11.1" evidence="1"/>
    <dbReference type="AlphaFoldDB" id="A0A452HLZ4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018225.1">
        <property type="protein sequence ID" value="ENSGAGP00000015950.1"/>
        <property type="gene ID" value="ENSGAGG00000012006.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000596">
        <property type="entry name" value="CK1 family protein kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050235">
        <property type="entry name" value="CK1_Ser-Thr_kinase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11909">
        <property type="entry name" value="CASEIN KINASE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Protein kinase" evidence="3">
        <location>
            <begin position="1"/>
            <end position="220"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="244"/>
            <end position="353"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="244"/>
            <end position="258"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="324"/>
            <end position="346"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00012513"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015950.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="353" mass="40511" checksum="5873EDDB16C12DF1" modified="2019-05-08" version="1">MMQGGVGIPTIKWCGAEGDYNVMVMELLGPSLEDLFNFCSRKFSLKTVLLLADQMISRIEYIHSKNFIHRDVKPDNFLMGLGKKGNLVYIIDFGLAKKYRDARTHQHIPYRENKNLTGTARYASINTHLGIEQSRRDDLESLGYVLMYFNLGSLPWQGLKAATKRQKYERISEKKMSTPIEVLCKGYPSEFATYLNFCRSLRFDDKPDYSYLRQLFRNLFHRQGFSYDYVFDWNMLKFGASRAAEDAERERREREERLRHTRNSAVRGLPSTASGRLRGTQDVAPPPPPLTPTSHTANASPRPVSGMERERKVSMRLHRGAPVNISSSDLTGRQDTSRMSTSQNSIPFEHHGK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="22" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLZ6</accession>
    <name>A0A452HLZ6_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="1">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="1 2">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="2" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="1" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="1" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HLZ6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018236.1">
        <property type="protein sequence ID" value="ENSGAGP00000015961.1"/>
        <property type="gene ID" value="ENSGAGG00000012004.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000313" key="1">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015961.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="208" mass="21883" checksum="A89ADFB8F3DC7CE5" modified="2019-05-08" version="1">MSAWPSTPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAPKLRAWCPPLPCSSAHRLPRARAAPDQSSRLCTSRWWQRRESQAPRAGSGCCQCSRPVGCATSI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLZ8</accession>
    <name>A0A452HLZ8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Sema domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="caution">
        <text evidence="3">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLZ8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018287.1">
        <property type="protein sequence ID" value="ENSGAGP00000016007.1"/>
        <property type="gene ID" value="ENSGAGG00000011815.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045499">
        <property type="term" value="F:chemorepellent activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030215">
        <property type="term" value="F:semaphorin receptor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007411">
        <property type="term" value="P:axon guidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001755">
        <property type="term" value="P:neural crest cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030335">
        <property type="term" value="P:positive regulation of cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071526">
        <property type="term" value="P:semaphorin-plexin signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd11253">
        <property type="entry name" value="Sema_3E"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:000015">
        <property type="entry name" value="Semaphorin 3B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1680.10:FF:000001">
        <property type="entry name" value="Semaphorin 3F like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1680.10">
        <property type="entry name" value="ligand-binding face of the semaphorins, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016201">
        <property type="entry name" value="PSI"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001627">
        <property type="entry name" value="Semap_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036352">
        <property type="entry name" value="Semap_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027231">
        <property type="entry name" value="Semaphorin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015513">
        <property type="entry name" value="Semaphorin_3E_Sema"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11036">
        <property type="entry name" value="SEMAPHORIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11036:SF22">
        <property type="entry name" value="SEMAPHORIN-3E"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01403">
        <property type="entry name" value="Sema"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00423">
        <property type="entry name" value="PSI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00630">
        <property type="entry name" value="Sema"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103575">
        <property type="entry name" value="Plexin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF101912">
        <property type="entry name" value="Sema domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51004">
        <property type="entry name" value="SEMA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="2" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Sema" evidence="4">
        <location>
            <begin position="14"/>
            <end position="499"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00352"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS51004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016007.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="620" mass="71021" checksum="9B281D20C385D696" modified="2019-05-08" version="1">KIGTSGHPTGEFLFPPLPRDHKLWDLNRTSVFHSPSGYLGLHTMLLDEYQERLFVGGRDLVYSLSLDRVSENYREIHWPSTPQQAEECTIKGRDADECANYIRVLHRYNRTHLLACGTGAFDPLCTFIRVGHPSEDHLFQLDWHKFERGRGRCPFDPSSSFISTLIGGELFAGLYNDYWGRDAAVLRTMGRMPHIRTEPDDERLLKEPKFVGSYMIPDNEDRDDNKVYFFFTEKALEAETGAHAIYARVGRVCANDMGGQRMLVNKWSTFLKTRLICSVPGMNGINTHFDELEDVFLLQTRDNKNPVVFGLFNTTSNIFRGYAICVYHIASIRAAFNGPYAHKEGPEYHWSLYEGKVPYPRPGSCASKINGGLYGTTKDYPDEAVRFARSHPLMYQPIKPVHKRPILVKTDGKYNLKQIAVDRVEAEDGQYDVLFIGTDNGIVLKVITIYNQDTESMEEVILEEMQVFKVPVPIISMEISSKRQQLYVGSESAVAQVKFHQCDMYGTACADCCLARDPYCAWDGISCSRYYPTGMQAKRRFRRQDVRHGNAAQQCFGQQFIGEALEKTEERIVYGIEYNSTLLECTPRTLQAKVIWFVQRAHEARKEEFKEMSNILRHSM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMA0</accession>
    <name>A0A452HMA0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Putative malate dehydrogenase 1B</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the LDH/MDH superfamily. MDH type 2 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMA0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018349.1">
        <property type="protein sequence ID" value="ENSGAGP00000016065.1"/>
        <property type="gene ID" value="ENSGAGG00000012069.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016615">
        <property type="term" value="F:malate dehydrogenase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016616">
        <property type="term" value="F:oxidoreductase activity, acting on the CH-OH group of donors, NAD or NADP as acceptor"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006108">
        <property type="term" value="P:malate metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006099">
        <property type="term" value="P:tricarboxylic acid cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.720:FF:000144">
        <property type="entry name" value="Malate dehydrogenase [NADP]"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.110.10:FF:000006">
        <property type="entry name" value="putative malate dehydrogenase 1B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.110.10">
        <property type="entry name" value="Lactate dehydrogenase/glycoside hydrolase, family 4, C-terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.720">
        <property type="entry name" value="NAD(P)-binding Rossmann-like Domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022383">
        <property type="entry name" value="Lactate/malate_DH_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015955">
        <property type="entry name" value="Lactate_DH/Glyco_Ohase_4_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010945">
        <property type="entry name" value="Malate_DH_type2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036291">
        <property type="entry name" value="NAD(P)-bd_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23382">
        <property type="entry name" value="MALATE DEHYDROGENASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02866">
        <property type="entry name" value="Ldh_1_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56327">
        <property type="entry name" value="LDH C-terminal domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51735">
        <property type="entry name" value="NAD(P)-binding Rossmann-fold domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0520">NAD</keyword>
    <keyword evidence="3" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0816">Tricarboxylic acid cycle</keyword>
    <feature type="domain" description="Lactate/malate dehydrogenase C-terminal" evidence="7">
        <location>
            <begin position="332"/>
            <end position="412"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="462"/>
            <end position="523"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="503"/>
            <end position="523"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00009613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022532"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023027"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00039310"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF02866"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016065.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="523" mass="58567" checksum="E8EF224380B23F02" modified="2019-05-08" version="1">MAKFVLAGKADCPYFAKAELLADYLQINLPNFKIHKITQHPDKWEHYYGITSDMLSEQMLRIAKENLQTHIEIEKEEEYIKSLINPLQVWISSASAPACYNLIPLLANGEVFGMATEVSIHLLDSSHYKEILHGIVMEAEDLAFPLLRSVSMHTELDDAFVQADIVILLDDILFQHEIPSLEDCIRQVTEQCKVYGSLIEKNANSKVKVIVSGKTFVNLRALMIMTFAPSIDGQNIIAVAMFLENAAKAMLARKLNMNSAGVKDVIVWGNISGNSYVDLSKAKVYKYDSAIWGPPNFSRHLLDMIYDGSWVRSEFAFVLSSLSSWGHHCLGMSPAHVIATVLRYWYQDSPPGEIVSMGILSEGQFCIPEGIIFSMPVRFQDGSWEVITETEISEETQETLKRLAHDLIQEKQVALGEIPEMRPYKEPAFNGFNLLALLEEENVQTPPEDLDTLSNGLLQPETQEGEKVQATTDEPEAPPDGLLQSEMLEEDKVQVPIDDPETSSESLNQSETSLSNAENNENP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMA1</accession>
    <name>A0A452HMA1_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="3">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000018360.1">
        <property type="protein sequence ID" value="ENSGAGP00000016074.1"/>
        <property type="gene ID" value="ENSGAGG00000012070.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005814">
        <property type="term" value="C:centriole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036064">
        <property type="term" value="C:ciliary basal body"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007224">
        <property type="term" value="P:smoothened signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029246">
        <property type="entry name" value="TALPID3"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15721">
        <property type="entry name" value="KIAA0586 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15721:SF2">
        <property type="entry name" value="PROTEIN TALPID3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15324">
        <property type="entry name" value="TALPID3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="678"/>
            <end position="711"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1150"/>
            <end position="1176"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1207"/>
            <end position="1258"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1293"/>
            <end position="1319"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="249"/>
            <end position="293"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="500"/>
            <end position="527"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="572"/>
            <end position="599"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="698"/>
            <end position="711"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="1207"/>
            <end position="1219"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="1306"/>
            <end position="1319"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016074.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1622" mass="178115" checksum="67420F7544322845" modified="2019-05-08" version="1">MEVESNSSVSQDSMASFTAGDVLTRSTSLHTKEKPTGAWTAATDCTRKKVKISVKKLREVASPYQADDAALFRDKTSYIPPALSANKPLPNRSTVTTSSSVTQAVRSTLAERSSEHLLSVENDLEPTPLSQKDYSKEAAVKGVPLQVMEDGNKPEDGDIFISQYTTGEKEALRAVLKQRTQNPPVFKEVKVQLLENVSAEKKESTQDTRSAHSNIDSAATIAAATAAAIATTAPLLKVQSDLEAKVSSVSALLSKLQETDKQLQRVAEKQTNIKAQQQERSRCHDRVNELEKQMTVFMEQRIQHLEKLQQQQIHVQSHLISSAINTGGFQPVNIPSSNLVAKHSTNPELQQPLTNQVSSCHRNLFSTNAVPTQAYSSQFGNYGVCTQKSPLKTPAPRRYAPVPVSKNVKISQKISKKERPIAEKENVPKSAYGGVVDGGRLLEQILNSQETPLRQTESSEKSLLNNTKMAWHSERECRPPALQTDSFPAFEGPLQDYNSINKTVKKAEDVLQDLGQLKREMHGMLQEAKAWKSDMNDFVKPTNPVVAPSLAECHQVSKPSILQTVKAPKSVLKDAERILRGVQNNKKVLEENLEAIIRAKDGDAMYTFINALTTNRHVLEEIRIRKTVDEWIKAISVEIQAEMAKNDYEQTQYNQKDPKTPWIKRTQNTRDMKTNKEIKAKTKKPQGSLTKKYLSAAKPSQKQSENNFGRQSLRSYSVSETLLKKEGRAEGPLKASAVVQNEEYLSQVYGKPIYQGHRSTLKKAPYLRFNSPSPKSKPQRPKVIECVRGTKVKSARTQTNSCTQKVIISPKRQRPLSAPLKESHYLFSPSREMPDLSGPLEGHLIPMAIPLGQMQINGISPQPAGVVIGKPHPVTVTTSVLPTSPKPQPKLKKPNIAVIEMRSEKKDPPKLTVQVLPNVDIDSISNDSVTINQASTSSEAALLPVSSLIQGPEQTQYEEEELKFPGTNFVDVTDIIQDQEEEDRIPELSEPILEFNRQTEVVSPKYNGPPFPPTAPASQQPADILDEIIERRETLENRLISWVEQEVMARIISGMYPVQQERVPSVSASDSEESEAVTSDIVEATGGGGFQLFVSAGVPVNSEMISHFVNEALTETIAVMLGDREAQKAVPAANVLQSTTVLPESVVSTPLATPPHTPLPEKEPAPVKTPVSSPSVTEIGGDVGEQEQMTEKGFDIPAATTPICTPVFTPVDTPPRVTTPSPPVSEHGSDATRMGSPEPPNPWGDAELPLDEEKPSPLNEETLHPRAVVMSVAKDEEPEMLILPSPEPVRPFESLPCEPRVPSPLHTPSSGPSTHESSLTVTETETADKPISEGEVLFSYGQLVAARALAEGGLSLPNLSESLTSTLHDANEMDYDPPSEGQVMRRPHRGYHRDPMLSLFAKLNQAPVALHEVIYHSEDSDNSVGELSEGQRPRLTRAAESILMGHSVYTDQVAGLVTEQRPHRVPSPGECDRVTAELLGDAADVSHGPMSMAELESQPLLNPAPQTAQTNNRLTSLSEDLPEKQSQKVVQVEPVRSRVIHVRNKSEVTQQQDSQGDMDRTHVEPNMYLTLLLKGEETVPHLSSVTSPAKMSVMLPSMNIDDQTQSISTVHGDSDSSGTDTF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMA2</accession>
    <name>A0A452HMA2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Peptidase M14 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <comment type="similarity">
        <text evidence="2 10">Belongs to the peptidase M14 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMA2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018361.1">
        <property type="protein sequence ID" value="ENSGAGP00000016075.1"/>
        <property type="gene ID" value="ENSGAGG00000012077.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004181">
        <property type="term" value="F:metallocarboxypeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.630.10:FF:000084">
        <property type="entry name" value="Carboxypeptidase B2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.630.10">
        <property type="entry name" value="Zn peptidases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057246">
        <property type="entry name" value="CARBOXYPEPT_ZN_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000834">
        <property type="entry name" value="Peptidase_M14"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11705:SF19">
        <property type="entry name" value="CARBOXYPEPTIDASE O"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11705">
        <property type="entry name" value="PROTEASE FAMILY M14 CARBOXYPEPTIDASE A,B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00246">
        <property type="entry name" value="Peptidase_M14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00765">
        <property type="entry name" value="CRBOXYPTASEA"/>
    </dbReference>
    <dbReference type="SMART" id="SM00631">
        <property type="entry name" value="Zn_pept"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53187">
        <property type="entry name" value="Zn-dependent exopeptidases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00132">
        <property type="entry name" value="CARBOXYPEPT_ZN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS52035">
        <property type="entry name" value="PEPTIDASE_M14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0121">Carboxypeptidase</keyword>
    <keyword evidence="7" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="5" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9" id="KW-0482">Metalloprotease</keyword>
    <keyword evidence="4" id="KW-0645">Protease</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="8" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="Peptidase M14" evidence="11">
        <location>
            <begin position="20"/>
            <end position="300"/>
        </location>
    </feature>
    <feature type="active site" description="Proton donor/acceptor" evidence="10">
        <location>
            <position position="266"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005988"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022645"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023049"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01379"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS52035"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016075.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="330" mass="38211" checksum="720C57AA357934E7" modified="2019-05-08" version="1">FFRVLIRDVQKQLDSSMPRKGGSHWRIYHWMYQIKESSSELVTQHYLGMTYESRPMYYLKVSQPSNKAKKIIWMDCGIHAREWISPAFCQSFMKENYKTDPKIKSFLQNLDIYILPVRNIDGYIYSWTTERLWKKSRSALVNGTCFGTDLNCNFNGSWGSNSLTFCGTGPESEPETKAVARFIEKRKSDVLYFLTIHSYGQYILTLYGYKTDLASNHQDLKAEVTLKEKHGTNFTVGPTSLVLYKNSSSLRDWAHMIGIPFFYTFELRDKGAYGFILLPDQIQPTCDETTTAVMTIIEYVNQKYFPNSAVAVTSGSLWLWLIVLFTRTLS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMA8</accession>
    <name>A0A452HMA8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Fibronectin type-III domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the type I cytokine receptor family. Type 2 subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMA8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018368.1">
        <property type="protein sequence ID" value="ENSGAGP00000016082.1"/>
        <property type="gene ID" value="ENSGAGG00000011475.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009897">
        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004896">
        <property type="term" value="F:cytokine receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000494">
        <property type="entry name" value="Leptin receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000501">
        <property type="entry name" value="Leptin receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000568">
        <property type="entry name" value="Leptin receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000613">
        <property type="entry name" value="Leptin receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003529">
        <property type="entry name" value="Hematopoietin_rcpt_Gp130_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010457">
        <property type="entry name" value="IgC2-like_lig-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041182">
        <property type="entry name" value="LEP-R_IGD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23037">
        <property type="entry name" value="CYTOKINE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23037:SF44">
        <property type="entry name" value="LEPTIN RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06328">
        <property type="entry name" value="Lep_receptor_Ig"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18589">
        <property type="entry name" value="ObR_Ig"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01353">
        <property type="entry name" value="HEMATOPO_REC_L_F2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7 12" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-0675">Receptor</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="3 12" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 12" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="790"/>
            <end position="812"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="13">
        <location>
            <begin position="195"/>
            <end position="283"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="13">
        <location>
            <begin position="490"/>
            <end position="586"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="13">
        <location>
            <begin position="691"/>
            <end position="784"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="907"/>
            <end position="926"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="907"/>
            <end position="916"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="917"/>
            <end position="926"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008921"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016082.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1114" mass="126212" checksum="E384BCC067D812B8" modified="2019-05-08" version="1">PCCSLFPTLTHFHQAGAEVAHCMVHQIPASSFMLPCMSPNETSVIPLTAGVVASSPGLHREHGTTETNSLMLVAEESFLCCLWSETNANCSLYTAGMETKKFTSSEVSSFAPQQTGRFAKFLSELSLEDISTGSLKGNFMVTPCNGNGSDKYECHIPSLKLNYTYITWLNIINGVTLLQSPLMSVRPINIVKPESPLNLRLEMTDKGQLKICWSNPVLTPYPLQYEVKFSGNTTQNIVIETSLIIGNMLVGSSYLVQVRCKSLHGTGFWSDWSTPYNLNTEDVMYFPPKILTSVGSNVSFHCLYNDKNKMILSKKIVWWLNLAEEIPERQYTLVNDRVSRVTLFNLNATKPRGKFFYNALYCCNQNRECHHRYAELYVVDVNINITCETDGYLTKMTCRWSANTNTLLVGSSLQLKYYRSSIYCSDFPSIPPKSEAKECHLQRNHYYECTFQPIFLLSGYTMWIEIKHQLGTLESPPTCVIPADVVKPFPPSNVKAEITKNIGLLNVSWTNPAFPNSDLKFQIRYSVNREEILWEIFEVSNAPTRSVMIKVLDLCVVYIVQVRCSGLDGLGYWSDWSKPAYTIVQDIQAPLRGPEFWRVINEDPIRKQKNVTLVWKPLMKNYSLCSVCGYIVKQHTSENITWTEYVTSGTTYTYPWMEDAHTITILAVNSVGASSMNFNLTLSQQMSTVNIVQSLSAYPVNSSCVILTWTLSPQMYVITSFVIEWKNLNEEEQMKWIRVAPNISKYYIYDHFILIEKYQFSLYPIFPEGVGNPKTTDEFIKDRSEKQNDAGLYVILPIVISSFVLLLGTLLISQQRMKKLFWEDVPNPKNCSWAQGVNFQKPETFEHLFIKHPEAISFGPLLLEPEIVLEDISVAKALKSEDKQDLLAVDSMFTMIQDSEHDSACSSSHFSNSLSESSHDDKVSERTMRQSNIKYATIISNCKLSGLYEEQKNLSGSFSGCFLGEDSLVIDPFSRSWEVGNQAFLILPDQHPSQASKTISLSVVSSEGFSEPSDHDDTFSDGDSPERSLYYLGLTSVKKNENDIFLTENSRVMCHLHTNGLFKDMGFLQDTSSDLNPFIKNSLKYENSVKTFVPYMPQFQATTIKLQETTETKT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMB1</accession>
    <name>A0A452HMB1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">DUF4371 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000018371.1">
        <property type="protein sequence ID" value="ENSGAGP00000016085.1"/>
        <property type="gene ID" value="ENSGAGG00000012081.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046983">
        <property type="term" value="F:protein dimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025398">
        <property type="entry name" value="DUF4371"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008906">
        <property type="entry name" value="HATC_C_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012337">
        <property type="entry name" value="RNaseH-like_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45749">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45749:SF35">
        <property type="entry name" value="AC-LIKE TRANSPOSASE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05699">
        <property type="entry name" value="Dimer_Tnp_hAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14291">
        <property type="entry name" value="DUF4371"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53098">
        <property type="entry name" value="Ribonuclease H-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="DUF4371" evidence="2">
        <location>
            <begin position="45"/>
            <end position="225"/>
        </location>
    </feature>
    <feature type="domain" description="HAT C-terminal dimerisation" evidence="1">
        <location>
            <begin position="520"/>
            <end position="600"/>
        </location>
    </feature>
    <evidence type="ECO:0000259" key="1">
        <source>
            <dbReference type="Pfam" id="PF05699"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF14291"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016085.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="5">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="626" mass="72501" checksum="11A49E8969EAD9BC" modified="2019-05-08" version="1">MLGQKRKRLSGAAYQLQREKQQKELEMLSHTSKKLNSEILWWQAVLKRSLAIINFLAEQCLAFCRSSDILYENNNGNFLKLVELLAKFDNVMAEHVQRVKDGEIHTHYLGKNTQNEIIELISNGVRNEILSNLEHANYYSIIVDCTQDLSKTKQRSIVLRFLKTDENAEVKICEHFLEFIPVNKTTGKALTDVILQRLEHYRIPLKNMRGQGYDNGSNMRGRHAGVQQRILHLNNRAFFSPCHAHSLDLVVXARTTGKCTAKNIEFKQSNAVSYFTTVQAIYNFFSASTHRWAVLKKHLEQKLTLKPLSQTRWESRIDDLSYDPSFRHEAETLAQKMMQFQFSCCTVIWHNILNQINLTSKVMQNITINISKAKMILNKMLEYLRKYHSDEEFEETVKEASTIAEDLDFEPTFAPQQFIRPREKTRQFCYEAHDDPILNPNKRFKVECFYCILDVAITSIEERFCQLHGHCEIFEFLYDIGNIKNQFSKEDLMKQCQDLYLALSTDNAADIDAVELYDELIALAELISPKTSPLKVLEFIARNVFFTPNTAITLRILLTLPVSTASGQRSFSKLKLLKNYLRSIMSQNRMSGLALISIESTIAKKLDFTNLLKDYASIKTRKIQFI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMB4</accession>
    <name>A0A452HMB4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Fibronectin type-III domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the type I cytokine receptor family. Type 2 subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMB4"/>
    <dbReference type="STRING" id="38772.ENSGAGP00000016087"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018373.1">
        <property type="protein sequence ID" value="ENSGAGP00000016087.1"/>
        <property type="gene ID" value="ENSGAGG00000011475.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009897">
        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004896">
        <property type="term" value="F:cytokine receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000494">
        <property type="entry name" value="Leptin receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000501">
        <property type="entry name" value="Leptin receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000568">
        <property type="entry name" value="Leptin receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000613">
        <property type="entry name" value="Leptin receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003529">
        <property type="entry name" value="Hematopoietin_rcpt_Gp130_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010457">
        <property type="entry name" value="IgC2-like_lig-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041182">
        <property type="entry name" value="LEP-R_IGD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23037">
        <property type="entry name" value="CYTOKINE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23037:SF44">
        <property type="entry name" value="LEPTIN RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06328">
        <property type="entry name" value="Lep_receptor_Ig"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18589">
        <property type="entry name" value="ObR_Ig"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01353">
        <property type="entry name" value="HEMATOPO_REC_L_F2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7 12" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-0675">Receptor</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="3 12" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 12" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="830"/>
            <end position="852"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="13">
        <location>
            <begin position="229"/>
            <end position="323"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="13">
        <location>
            <begin position="530"/>
            <end position="626"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="13">
        <location>
            <begin position="731"/>
            <end position="824"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="947"/>
            <end position="966"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="947"/>
            <end position="956"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="957"/>
            <end position="966"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008921"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016087.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1154" mass="130964" checksum="B31505D117526EB4" modified="2019-05-08" version="1">VAWSVVLTGATRVPFRLVAHCMVHQIPASSFMLPCMSPNETSVIPLTAGVVASSPGLHREHGTTETNSLMLVAEESFLCCLWSETNANCSLYTAGMETKKFTSSEVSSFAPQQTDLSWNIQCWTKGDLELFVCILKLSNKKSYLNGEFKINLLYVLSELSLEDISTGSLKGNFMVTPCNGNGSDKYECHIPSLKLNYTYITWLNIINGVTLLQSPLMSVRPINIVKPESPLNLRLEMTDKGQLKICWSNPVLTPYPLQYEVKFSGNTTQNAWQVVEIVIETSLIIGNMLVGSSYLVQVRCKSLHGTGFWSDWSTPYNLNTEDVMYFPPKILTSVGSNVSFHCLYNDKNKMILSKKIVWWLNLAEEIPERQYTLVNDRVSRVTLFNLNATKPRGKFFYNALYCCNQNRECHHRYAELYVVDVNINITCETDGYLTKMTCRWSANTNTLLVGSSLQLKYYRSSIYCSDFPSIPPKSEAKECHLQRNHYYECTFQPIFLLSGYTMWIEIKHQLGTLESPPTCVIPADVVKPFPPSNVKAEITKNIGLLNVSWTNPAFPNSDLKFQIRYSVNREEILWEIFEVSNAPTRSVMIKVLDLCVVYIVQVRCSGLDGLGYWSDWSKPAYTIVQDIQAPLRGPEFWRVINEDPIRKQKNVTLVWKPLMKNYSLCSVCGYIVKQHTSENITWTEYVTSGTTYTYPWMEDAHTITILAVNSVGASSMNFNLTLSQQMSTVNIVQSLSAYPVNSSCVILTWTLSPQMYVITSFVIEWKNLNEEEQMKWIRVAPNISKYYIYDHFILIEKYQFSLYPIFPEGVGNPKTTDEFIKDRSEKQNDAGLYVILPIVISSFVLLLGTLLISQQRMKKLFWEDVPNPKNCSWAQGVNFQKPETFEHLFIKHPEAISFGPLLLEPEIVLEDISVAKALKSEDKQDLLAVDSMFTMIQDSEHDSACSSSHFSNSLSESSHDDKVSERTMRQSNIKYATIISNCKLSGLYEEQKNLSGSFSGCFLGEDSLVIDPFSRSWEVGNQAFLILPDQHPSQASKTISLSVVSSEGFSEPSDHDDTFSDGDSPERSLYYLGLTSVKKNENDIFLTENSRVMCHLHTNGLFKDMGFLQDTSSDLNPFIKNSLKYENSVKTFVPYMPQFQATTIKLQETTETKT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMB6</accession>
    <name>A0A452HMB6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Fibronectin type-III domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the type I cytokine receptor family. Type 2 subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMB6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018367.1">
        <property type="protein sequence ID" value="ENSGAGP00000016081.1"/>
        <property type="gene ID" value="ENSGAGG00000011475.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009897">
        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004896">
        <property type="term" value="F:cytokine receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000494">
        <property type="entry name" value="Leptin receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000501">
        <property type="entry name" value="Leptin receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000568">
        <property type="entry name" value="Leptin receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000613">
        <property type="entry name" value="Leptin receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003529">
        <property type="entry name" value="Hematopoietin_rcpt_Gp130_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010457">
        <property type="entry name" value="IgC2-like_lig-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041182">
        <property type="entry name" value="LEP-R_IGD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23037">
        <property type="entry name" value="CYTOKINE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23037:SF44">
        <property type="entry name" value="LEPTIN RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06328">
        <property type="entry name" value="Lep_receptor_Ig"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18589">
        <property type="entry name" value="ObR_Ig"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01353">
        <property type="entry name" value="HEMATOPO_REC_L_F2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7 11" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-0675">Receptor</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="3 11" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 11" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="831"/>
            <end position="853"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="883"/>
            <end position="905"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="12">
        <location>
            <begin position="233"/>
            <end position="324"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="12">
        <location>
            <begin position="531"/>
            <end position="627"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="12">
        <location>
            <begin position="732"/>
            <end position="825"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008921"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016081.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="907" mass="104131" checksum="E1A31C6CE219F2C8" modified="2019-05-08" version="1">YHYQKYYYVTHFLHYFFVFPDFMQMAVAHCMVHQIPASSFMLPCMSPNETSVIPLTAGVVASSPGLHREHGTTETNSLMLVAEESFLCCLWSETNANCSLYTAGMETKKFTSSEVSSFAPQQTGSWNIQCWTKGDLELFVCILKLSNKKSYLNGEFKINLLYLSLEDISTGSLKGNFMVTPCNGNGSDKYECHIPSLKLNYTYITWLNIINGVTLLQSPLMSVRPINIVKPESPLNLRLEMTDKGQLKICWSNPVLTPYPLQYEVKFSGNTTQNVVEIVIETSLIIGNMLVGSSYLVQVRCKSLHGTGFWSDWSTPYNLNTEDVMYFPPKILTSVGSNVSFHCLYNDKNKMILSKKIVWWLNLAEEIPERQYTLVNDRVSRVTLFNLNATKPRGKFFYNALYCCNQNRECHHRYAELYVVDVNINITCETDGYLTKMTCRWSANTNTLLVGSSLQLKYYRSSIYCSDFPSIPPKSEAKECHLQRNHYYECTFQPIFLLSGYTMWIEIKHQLGTLESPPTCVIPADVVKPFPPSNVKAEITKNIGLLNVSWTNPAFPNSDLKFQIRYSVNREEILWEIFEVSNAPTRSVMIKVLDLCVVYIVQVRCSGLDGLGYWSDWSKPAYTIVQDIQAPLRGPEFWRVINEDPIRKQKNVTLVWKPLMKNYSLCSVCGYIVKQHTSENITWTEYVTSGTTYTYPWMEDAHTITILAVNSVGASSMNFNLTLSQQMSTVNIVQSLSAYPVNSSCVILTWTLSPQMYVITSFVIEWKNLNEEEQMKWIRVAPNISKYYIYDHFILIEKYQFSLYPIFPEGVGNPKTTDEFIKDRSEKQNDAGLYVILPIVISSFVLLLGTLLISQQRMKKLFWEDVPNPKNCSWAQGVNFQKVWVSIMIRFLVWVYVFHTLTFSLKC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMB8</accession>
    <name>A0A452HMB8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">Cyclin dependent kinase 8</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="12">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.22"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.22"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>[DNA-directed RNA polymerase] + ATP = phospho-[DNA-directed RNA polymerase] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10216"/>
            <dbReference type="Rhea" id="RHEA-COMP:11321"/>
            <dbReference type="Rhea" id="RHEA-COMP:11322"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43176"/>
            <dbReference type="ChEBI" id="CHEBI:68546"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.23"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMB8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018423.1">
        <property type="protein sequence ID" value="ENSGAGP00000016135.1"/>
        <property type="gene ID" value="ENSGAGG00000012060.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004693">
        <property type="term" value="F:cyclin-dependent protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008353">
        <property type="term" value="F:RNA polymerase II CTD heptapeptide repeat kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050108">
        <property type="entry name" value="CDK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24056">
        <property type="entry name" value="CELL DIVISION PROTEIN KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24056:SF243">
        <property type="entry name" value="CYCLIN-DEPENDENT KINASE 8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="6" id="KW-0418">Kinase</keyword>
    <keyword evidence="5" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="9" id="KW-0804">Transcription</keyword>
    <keyword evidence="8" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="4" id="KW-0808">Transferase</keyword>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="146"/>
            <end position="249"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="158"/>
            <end position="176"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="194"/>
            <end position="211"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="221"/>
            <end position="249"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00047811"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00048367"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00049280"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016135.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="17">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="249" mass="28404" checksum="7B04985EC0191AD4" modified="2019-05-08" version="1">EPPNSSLEPLALLTSEPIFHCRQEDIKTSNPYHHDQLDRIFNVMGFPADKDWEDIKKMPEHSTLMKDFRRNTYTNCSLIKYMEKHKVKPDSKAFHLLQKLLTMDPIKRITSEQAMQDPYFLEDPLPTSDVFAGCQIPYPKREFLTEEEPDDKGDKKNQQQQQGNNHTNGTGHPGNQDNSHTQGPPLKKVRVVPPTTTSGGLIMTSDYQRSNPHAAYPNPGPSTSQPQSSMGYSTTSQQPPQYSHQTHRY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMB9</accession>
    <name>A0A452HMB9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Fibronectin type-III domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the type I cytokine receptor family. Type 2 subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMB9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018372.1">
        <property type="protein sequence ID" value="ENSGAGP00000016086.1"/>
        <property type="gene ID" value="ENSGAGG00000011475.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009897">
        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004896">
        <property type="term" value="F:cytokine receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000494">
        <property type="entry name" value="Leptin receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000501">
        <property type="entry name" value="Leptin receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000568">
        <property type="entry name" value="Leptin receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000613">
        <property type="entry name" value="Leptin receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003529">
        <property type="entry name" value="Hematopoietin_rcpt_Gp130_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010457">
        <property type="entry name" value="IgC2-like_lig-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041182">
        <property type="entry name" value="LEP-R_IGD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23037">
        <property type="entry name" value="CYTOKINE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23037:SF44">
        <property type="entry name" value="LEPTIN RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06328">
        <property type="entry name" value="Lep_receptor_Ig"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18589">
        <property type="entry name" value="ObR_Ig"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01353">
        <property type="entry name" value="HEMATOPO_REC_L_F2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7 12" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-0675">Receptor</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="3 12" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 12" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="830"/>
            <end position="852"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="13">
        <location>
            <begin position="229"/>
            <end position="323"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="13">
        <location>
            <begin position="530"/>
            <end position="626"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="13">
        <location>
            <begin position="731"/>
            <end position="824"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="947"/>
            <end position="966"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="947"/>
            <end position="956"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="957"/>
            <end position="966"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008921"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016086.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1150" mass="130585" checksum="C6D259E09D62246B" modified="2019-05-08" version="1">VAWSVVLTGATRVPFRLVAHCMVHQIPASSFMLPCMSPNETSVIPLTAGVVASSPGLHREHGTTETNSLMLVAEESFLCCLWSETNANCSLYTAGMETKKFTSSEVSSFAPQQTDLSWNIQCWTKGDLELFVCILKLSNKKSYLNGEFKINLLYVLSELSLEDISTGSLKGNFMVTPCNGNGSDKYECHIPSLKLNYTYITWLNIINGVTLLQSPLMSVRPINIVKPESPLNLRLEMTDKGQLKICWSNPVLTPYPLQYEVKFSGNTTQNAWQVVEIVIETSLIIGNMLVGSSYLVQVRCKSLHGTGFWSDWSTPYNLNTEDVMYFPPKILTSVGSNVSFHCLYNDKNKMILSKKIVWWLNLAEEIPERQYTLVNDRVSRVTLFNLNATKPRGKFFYNALYCCNQNRECHHRYAELYVVDVNINITCETDGYLTKMTCRWSANTNTLLVGSSLQLKYYRSSIYCSDFPSIPPKSEAKECHLQRNHYYECTFQPIFLLSGYTMWIEIKHQLGTLESPPTCVIPADVVKPFPPSNVKAEITKNIGLLNVSWTNPAFPNSDLKFQIRYSVNREEILWEIFEVSNAPTRSVMIKVLDLCVVYIVQVRCSGLDGLGYWSDWSKPAYTIVQDIQAPLRGPEFWRVINEDPIRKQKNVTLVWKPLMKNYSLCSVCGYIVKQHTSENITWTEYVTSGTTYTYPWMEDAHTITILAVNSVGASSMNFNLTLSQQMSTVNIVQSLSAYPVNSSCVILTWTLSPQMYVITSFVIEWKNLNEEEQMKWIRVAPNISKYYIYDHFILIEKYQFSLYPIFPEGVGNPKTTDEFIKDRSEKQNDAGLYVILPIVISSFVLLLGTLLISQQRMKKLFWEDVPNPKNCSWAQGVNFQKPETFEHLFIKHPEAISFGPLLLEPEIVLEDISVAKALKSEDKQDLLAVDSMFTMIQDSEHDSACSSSHFSNSLSESSHDDKVSERTMRQSNIKYATIISNCKLSGLYEEQKNLSGSFSGCFLGEDSLVIDPFSRSWEVGNQAFLILPDQHPSQASKTISLSVVSSEGFSEPSDHDDTFSDGDSPERSLYYLGLTSVKKNENDIFLTENSRVMCHLHTNGLFKDMGFLQDTSSDLNPFIKNSLKYENSVKTFVPYMPQFQIRNFSYVNSS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HME0</accession>
    <name>A0A452HME0_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="19">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="12">Belongs to the APP family.</text>
    </comment>
    <comment type="caution">
        <text evidence="12">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HME0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018402.1">
        <property type="protein sequence ID" value="ENSGAGP00000016115.1"/>
        <property type="gene ID" value="ENSGAGG00000012099.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009986">
        <property type="term" value="C:cell surface"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012505">
        <property type="term" value="C:endomembrane system"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043005">
        <property type="term" value="C:neuron projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0099503">
        <property type="term" value="C:secretory vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140677">
        <property type="term" value="F:molecular function activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004867">
        <property type="term" value="F:serine-type endopeptidase inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046914">
        <property type="term" value="F:transition metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007409">
        <property type="term" value="P:axonogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007417">
        <property type="term" value="P:central nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010604">
        <property type="term" value="P:positive regulation of macromolecule metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051246">
        <property type="term" value="P:regulation of protein metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd21709">
        <property type="entry name" value="JMTM_APLP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd22607">
        <property type="entry name" value="Kunitz_ABPP-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1490.140:FF:000001">
        <property type="entry name" value="Amyloid beta (A4) protein b"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.570.10:FF:000001">
        <property type="entry name" value="Amyloid beta A4 protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.770:FF:000001">
        <property type="entry name" value="Amyloid beta A4 protein-like isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.410.10:FF:000003">
        <property type="entry name" value="amyloid-like protein 2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.1670">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.770">
        <property type="entry name" value="Amyloid precursor protein, E2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1490.140">
        <property type="entry name" value="Amyloidogenic glycoprotein, copper-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.570.10">
        <property type="entry name" value="Amyloidogenic glycoprotein, heparin-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.410.10">
        <property type="entry name" value="Pancreatic trypsin inhibitor Kunitz domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036669">
        <property type="entry name" value="Amyloid_Cu-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008155">
        <property type="entry name" value="Amyloid_glyco"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011178">
        <property type="entry name" value="Amyloid_glyco_Cu-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024329">
        <property type="entry name" value="Amyloid_glyco_E2_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008154">
        <property type="entry name" value="Amyloid_glyco_extra"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015849">
        <property type="entry name" value="Amyloid_glyco_heparin-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036454">
        <property type="entry name" value="Amyloid_glyco_heparin-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019745">
        <property type="entry name" value="Amyloid_glyco_intracell_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019543">
        <property type="entry name" value="APP_amyloid_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019744">
        <property type="entry name" value="APP_CUBD_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036176">
        <property type="entry name" value="E2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002223">
        <property type="entry name" value="Kunitz_BPTI"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036880">
        <property type="entry name" value="Kunitz_BPTI_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020901">
        <property type="entry name" value="Prtase_inh_Kunz-CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23103">
        <property type="entry name" value="ALZHEIMER'S DISEASE BETA-AMYLOID RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23103:SF14">
        <property type="entry name" value="AMYLOID BETA PRECURSOR LIKE PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10515">
        <property type="entry name" value="APP_amyloid"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12924">
        <property type="entry name" value="APP_Cu_bd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12925">
        <property type="entry name" value="APP_E2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02177">
        <property type="entry name" value="APP_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00014">
        <property type="entry name" value="Kunitz_BPTI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00203">
        <property type="entry name" value="AMYLOIDA4"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00759">
        <property type="entry name" value="BASICPTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00006">
        <property type="entry name" value="A4_EXTRA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00131">
        <property type="entry name" value="KU"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56491">
        <property type="entry name" value="A heparin-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF89811">
        <property type="entry name" value="Amyloid beta a4 protein copper binding domain (domain 2)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57362">
        <property type="entry name" value="BPTI-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109843">
        <property type="entry name" value="CAPPD, an extracellular domain of amyloid beta A4 protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00319">
        <property type="entry name" value="APP_CUBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51869">
        <property type="entry name" value="APP_E1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51870">
        <property type="entry name" value="APP_E2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00320">
        <property type="entry name" value="APP_INTRA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00280">
        <property type="entry name" value="BPTI_KUNITZ_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50279">
        <property type="entry name" value="BPTI_KUNITZ_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0186">Copper</keyword>
    <keyword evidence="10 12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9 14" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="2" id="KW-0646">Protease inhibitor</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0722">Serine protease inhibitor</keyword>
    <keyword evidence="5 15" id="KW-0732">Signal</keyword>
    <keyword evidence="3 14" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 14" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="15">
        <location>
            <begin position="1"/>
            <end position="25"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019367676" evidence="15">
        <location>
            <begin position="26"/>
            <end position="736"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="667"/>
            <end position="689"/>
        </location>
    </feature>
    <feature type="domain" description="E1" evidence="17">
        <location>
            <begin position="40"/>
            <end position="201"/>
        </location>
    </feature>
    <feature type="domain" description="BPTI/Kunitz inhibitor" evidence="16">
        <location>
            <begin position="302"/>
            <end position="352"/>
        </location>
    </feature>
    <feature type="domain" description="E2" evidence="18">
        <location>
            <begin position="365"/>
            <end position="556"/>
        </location>
    </feature>
    <feature type="region of interest" description="GFLD subdomain" evidence="12">
        <location>
            <begin position="40"/>
            <end position="135"/>
        </location>
    </feature>
    <feature type="region of interest" description="CuBD subdomain" evidence="12">
        <location>
            <begin position="143"/>
            <end position="201"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="209"/>
            <end position="263"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="13">
        <location>
            <begin position="209"/>
            <end position="224"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="13">
        <location>
            <begin position="233"/>
            <end position="261"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="85"/>
            <end position="129"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="110"/>
            <end position="117"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="145"/>
            <end position="199"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="156"/>
            <end position="186"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="170"/>
            <end position="198"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022690"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023008"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01217"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50279"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS51869"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS51870"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016115.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="736" mass="84109" checksum="F806D2E73F56A5B9" modified="2019-05-08" version="1">MVAGGELALLLLPLLALTGPAATLAGYIEALAANAGTGFAVAEPQIAMFCGKLNMHVNIQTGKWEPDTSGTKSCFGTKEEILQYCEEIYPELQITNVVEANQPVSIDNWCKKGKKQCKGHTHIVVPYKCLVGEFVSDVLLVPEKCRFFHKERMDVCESHQHWHTVAKEACLTEGMILHSYGMLLPCGVDQFHGTEYVCCPQTKIVDEALSQEEEDEDEDEDDDYDLYKSEFPTEPDVEEFTGTAVEEDEEEEDEEEEEVVEDRDYYYDNYKVDDYNEETPTESSNDRSLSEKEMISDMKAVCSQEAMTGPCRAVMPRWYFDLYKRKCIRFIYGGCGGNRNNFESEEYCMAVCKKMIPPTPVPTDDVDVYFETPADDNEHARFQKAKEQLEVRHHNRMDRVKKEWEEAEYESKNLPKAERQTLIQHFQAMVKSLEKEAASEKQQLVETHLARVEAMLNDRRRIALENYLAALQADPPRPHRILQALKRYVRAENKDRLHTVRHYQHVLAVDPEKAAQMKSQVMTHLHVIEERMNQSLSLLYKVPYVAEEIQDEIDELLQEQRADMDQFTSSISESQVDVRVSSEESEEIPLQEGKPFRPFQVKTFPAVPGNEDPQPDLYHPMKKGSGMADLDGLIGAEEKVINSKNKVDENVDNESSLREDFSLSSSALIGLLVIAVAIATVIVISLVMLRKRQYGTISHGIVEVDPMLTPEERHLNKMQNHGYENPTYKYLEQMQI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HME3</accession>
    <name>A0A452HME3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Transmembrane protein 45A</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the TMEM45 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HME3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018398.1">
        <property type="protein sequence ID" value="ENSGAGP00000016111.1"/>
        <property type="gene ID" value="ENSGAGG00000012098.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006904">
        <property type="entry name" value="DUF716"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042127">
        <property type="entry name" value="TMEM45"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16007">
        <property type="entry name" value="EPIDIDYMAL MEMBRANE PROTEIN E9-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16007:SF15">
        <property type="entry name" value="TRANSMEMBRANE PROTEIN 45B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04819">
        <property type="entry name" value="DUF716"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4" id="KW-1133">Transmembrane helix</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006948"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016111.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="8">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="238" mass="27484" checksum="840A3F59DE64DE70" modified="2019-05-08" version="1">MGIFGDHTLPDTLILMLEFWWSLNYPLKRLNENCCTNRCYQSGEFLEAAGKLFSFVGKLLKFLPAVSKQIHQHSTMYLSFLFPDRFVFSQVMFIEGLLFHFHVHYRSALQQHVHLLSTVALCNNAFSLCLEVFIPHHPHPTALQHPCSPHPGSWFWQISFVLYPPWGWLDWDQTDDGNIMFSTRCVSWHYTAALINLLSITQLFSGMRWVKLTRPQDRKPKPCYAGLQPGAPSPTTQG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HME6</accession>
    <name>A0A452HME6_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="18">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="12">Belongs to the APP family.</text>
    </comment>
    <comment type="caution">
        <text evidence="12">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HME6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018404.1">
        <property type="protein sequence ID" value="ENSGAGP00000016117.1"/>
        <property type="gene ID" value="ENSGAGG00000012099.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009986">
        <property type="term" value="C:cell surface"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012505">
        <property type="term" value="C:endomembrane system"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043005">
        <property type="term" value="C:neuron projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0099503">
        <property type="term" value="C:secretory vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140677">
        <property type="term" value="F:molecular function activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004867">
        <property type="term" value="F:serine-type endopeptidase inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046914">
        <property type="term" value="F:transition metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007409">
        <property type="term" value="P:axonogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007417">
        <property type="term" value="P:central nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010604">
        <property type="term" value="P:positive regulation of macromolecule metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051246">
        <property type="term" value="P:regulation of protein metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd21709">
        <property type="entry name" value="JMTM_APLP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1490.140:FF:000001">
        <property type="entry name" value="Amyloid beta (A4) protein b"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.570.10:FF:000001">
        <property type="entry name" value="Amyloid beta A4 protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.770:FF:000001">
        <property type="entry name" value="Amyloid beta A4 protein-like isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.1670">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.770">
        <property type="entry name" value="Amyloid precursor protein, E2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1490.140">
        <property type="entry name" value="Amyloidogenic glycoprotein, copper-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.570.10">
        <property type="entry name" value="Amyloidogenic glycoprotein, heparin-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036669">
        <property type="entry name" value="Amyloid_Cu-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008155">
        <property type="entry name" value="Amyloid_glyco"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011178">
        <property type="entry name" value="Amyloid_glyco_Cu-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024329">
        <property type="entry name" value="Amyloid_glyco_E2_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008154">
        <property type="entry name" value="Amyloid_glyco_extra"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015849">
        <property type="entry name" value="Amyloid_glyco_heparin-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036454">
        <property type="entry name" value="Amyloid_glyco_heparin-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019745">
        <property type="entry name" value="Amyloid_glyco_intracell_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019543">
        <property type="entry name" value="APP_amyloid_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019744">
        <property type="entry name" value="APP_CUBD_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036176">
        <property type="entry name" value="E2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23103">
        <property type="entry name" value="ALZHEIMER'S DISEASE BETA-AMYLOID RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23103:SF14">
        <property type="entry name" value="AMYLOID BETA PRECURSOR LIKE PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10515">
        <property type="entry name" value="APP_amyloid"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12924">
        <property type="entry name" value="APP_Cu_bd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12925">
        <property type="entry name" value="APP_E2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02177">
        <property type="entry name" value="APP_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00203">
        <property type="entry name" value="AMYLOIDA4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00006">
        <property type="entry name" value="A4_EXTRA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56491">
        <property type="entry name" value="A heparin-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF89811">
        <property type="entry name" value="Amyloid beta a4 protein copper binding domain (domain 2)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109843">
        <property type="entry name" value="CAPPD, an extracellular domain of amyloid beta A4 protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00319">
        <property type="entry name" value="APP_CUBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51869">
        <property type="entry name" value="APP_E1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51870">
        <property type="entry name" value="APP_E2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00320">
        <property type="entry name" value="APP_INTRA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0186">Copper</keyword>
    <keyword evidence="10 12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9 14" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="2" id="KW-0646">Protease inhibitor</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0722">Serine protease inhibitor</keyword>
    <keyword evidence="5 15" id="KW-0732">Signal</keyword>
    <keyword evidence="3 14" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 14" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="15">
        <location>
            <begin position="1"/>
            <end position="25"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019329044" evidence="15">
        <location>
            <begin position="26"/>
            <end position="705"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="636"/>
            <end position="658"/>
        </location>
    </feature>
    <feature type="domain" description="E1" evidence="16">
        <location>
            <begin position="40"/>
            <end position="201"/>
        </location>
    </feature>
    <feature type="domain" description="E2" evidence="17">
        <location>
            <begin position="309"/>
            <end position="500"/>
        </location>
    </feature>
    <feature type="region of interest" description="GFLD subdomain" evidence="12">
        <location>
            <begin position="40"/>
            <end position="135"/>
        </location>
    </feature>
    <feature type="region of interest" description="CuBD subdomain" evidence="12">
        <location>
            <begin position="143"/>
            <end position="201"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="209"/>
            <end position="265"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="274"/>
            <end position="293"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="13">
        <location>
            <begin position="209"/>
            <end position="224"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="13">
        <location>
            <begin position="233"/>
            <end position="261"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="13">
        <location>
            <begin position="284"/>
            <end position="293"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="85"/>
            <end position="129"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="110"/>
            <end position="117"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="145"/>
            <end position="199"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="156"/>
            <end position="186"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="170"/>
            <end position="198"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022690"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023008"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01217"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS51869"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS51870"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016117.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="705" mass="80402" checksum="AA24D78F4779DCCF" modified="2019-05-08" version="1">MVAGGELALLLLPLLALTGPAATLAGYIEALAANAGTGFAVAEPQIAMFCGKLNMHVNIQTGKWEPDTSGTKSCFGTKEEILQYCEEIYPELQITNVVEANQPVSIDNWCKKGKKQCKGHTHIVVPYKCLVGEFVSDVLLVPEKCRFFHKERMDVCESHQHWHTVAKEACLTEGMILHSYGMLLPCGVDQFHGTEYVCCPQTKIVDEALSQEEEDEDEDEDDDYDLYKSEFPTEPDVEEFTGTAVEEDEEEEDEEEEEVVEDRDYYYDNYKVDDYNEETPTESSNDRSLSEKEMISDMKVPPTPVPTDDVDVYFETPADDNEHARFQKAKEQLEVRHHNRMDRVKKEWEEAEYESKNLPKAERQTLIQHFQAMVKSLEKEAASEKQQLVETHLARVEAMLNDRRRIALENYLAALQADPPRPHRILQALKRYVRAENKDRLHTVRHYQHVLAVDPEKAAQMKSQVMTHLHVIEERMNQSLSLLYKVPYVAEEIQDEIDELLQEQRADMDQFTSSISESQVDVRVSSEESEEIPLQEGKPFRPFQVKTFPAVPGNEDPQPDLYHPMKKGSGMADLDGLIGAEEKVINSKNKVDENVVIDETLDVKEMIFNAERVGGLMDEPDNESSLREDFSLSSSALIGLLVIAVAIATVIVISLVMLRKRQYGTISHGIVEVDPMLTPEERHLNKMQNHGYENPTYKYLEQMQI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HME7</accession>
    <name>A0A452HME7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Cysteine-rich protein 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="7">Interacts with TGFB1I1.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HME7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018403.1">
        <property type="protein sequence ID" value="ENSGAGP00000016116.1"/>
        <property type="gene ID" value="ENSGAGG00000012083.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd09401">
        <property type="entry name" value="LIM_TLP_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.110.10:FF:000025">
        <property type="entry name" value="Cysteine-rich protein 2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.110.10">
        <property type="entry name" value="Cysteine Rich Protein"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001781">
        <property type="entry name" value="Znf_LIM"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46074:SF2">
        <property type="entry name" value="CYSTEINE-RICH PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46074">
        <property type="entry name" value="CYSTEINE-RICH PROTEIN CRIP FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00412">
        <property type="entry name" value="LIM"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00132">
        <property type="entry name" value="LIM"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57716">
        <property type="entry name" value="Glucocorticoid receptor-like (DNA-binding domain)"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00478">
        <property type="entry name" value="LIM_DOMAIN_1"/>
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    </dbReference>
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        <property type="entry name" value="LIM_DOMAIN_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0007">Acetylation</keyword>
    <keyword evidence="6 9" id="KW-0440">LIM domain</keyword>
    <keyword evidence="2 9" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="1" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="4 9" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="LIM zinc-binding" evidence="10">
        <location>
            <begin position="3"/>
            <end position="64"/>
        </location>
    </feature>
    <feature type="domain" description="LIM zinc-binding" evidence="10">
        <location>
            <begin position="123"/>
            <end position="184"/>
        </location>
    </feature>
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        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023038"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00038732"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00039933"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016116.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="207" mass="22830" checksum="77C05D7D4DEF2EEA" modified="2019-05-08" version="1">PRSHCHLCRKTFAFPEKVTSLGKDWHKFCLKCERCNKTLTPGGHAEHDGKPFCHKPCYATLFGPKGVNIGGAGSYIYEKPQMEEQPAPGPIEHISKVEEKKVSGPPKGPSRASSVTTFTGEPNMCPRCGKKVYFAEKVTSLGKDWHRPCLRCERCGKTLTPGGHAEHDEQPYCHKPCYGILFGPKGVNTGAVGSYIYDKDPEAKDQP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HME8</accession>
    <name>A0A452HME8_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="18">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="12">Belongs to the APP family.</text>
    </comment>
    <comment type="caution">
        <text evidence="12">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HME8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018406.1">
        <property type="protein sequence ID" value="ENSGAGP00000016119.1"/>
        <property type="gene ID" value="ENSGAGG00000012099.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009986">
        <property type="term" value="C:cell surface"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012505">
        <property type="term" value="C:endomembrane system"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043005">
        <property type="term" value="C:neuron projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0099503">
        <property type="term" value="C:secretory vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140677">
        <property type="term" value="F:molecular function activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004867">
        <property type="term" value="F:serine-type endopeptidase inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046914">
        <property type="term" value="F:transition metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007409">
        <property type="term" value="P:axonogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007417">
        <property type="term" value="P:central nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010604">
        <property type="term" value="P:positive regulation of macromolecule metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051246">
        <property type="term" value="P:regulation of protein metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd21709">
        <property type="entry name" value="JMTM_APLP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd22607">
        <property type="entry name" value="Kunitz_ABPP-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1490.140:FF:000001">
        <property type="entry name" value="Amyloid beta (A4) protein b"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.570.10:FF:000001">
        <property type="entry name" value="Amyloid beta A4 protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.770:FF:000001">
        <property type="entry name" value="Amyloid beta A4 protein-like isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.410.10:FF:000003">
        <property type="entry name" value="amyloid-like protein 2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.1670">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.770">
        <property type="entry name" value="Amyloid precursor protein, E2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1490.140">
        <property type="entry name" value="Amyloidogenic glycoprotein, copper-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.570.10">
        <property type="entry name" value="Amyloidogenic glycoprotein, heparin-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.410.10">
        <property type="entry name" value="Pancreatic trypsin inhibitor Kunitz domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036669">
        <property type="entry name" value="Amyloid_Cu-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008155">
        <property type="entry name" value="Amyloid_glyco"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011178">
        <property type="entry name" value="Amyloid_glyco_Cu-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024329">
        <property type="entry name" value="Amyloid_glyco_E2_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008154">
        <property type="entry name" value="Amyloid_glyco_extra"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015849">
        <property type="entry name" value="Amyloid_glyco_heparin-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036454">
        <property type="entry name" value="Amyloid_glyco_heparin-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019745">
        <property type="entry name" value="Amyloid_glyco_intracell_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019543">
        <property type="entry name" value="APP_amyloid_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019744">
        <property type="entry name" value="APP_CUBD_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036176">
        <property type="entry name" value="E2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002223">
        <property type="entry name" value="Kunitz_BPTI"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036880">
        <property type="entry name" value="Kunitz_BPTI_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020901">
        <property type="entry name" value="Prtase_inh_Kunz-CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23103">
        <property type="entry name" value="ALZHEIMER'S DISEASE BETA-AMYLOID RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23103:SF14">
        <property type="entry name" value="AMYLOID BETA PRECURSOR LIKE PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10515">
        <property type="entry name" value="APP_amyloid"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12924">
        <property type="entry name" value="APP_Cu_bd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12925">
        <property type="entry name" value="APP_E2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02177">
        <property type="entry name" value="APP_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00014">
        <property type="entry name" value="Kunitz_BPTI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00203">
        <property type="entry name" value="AMYLOIDA4"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00759">
        <property type="entry name" value="BASICPTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00006">
        <property type="entry name" value="A4_EXTRA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00131">
        <property type="entry name" value="KU"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56491">
        <property type="entry name" value="A heparin-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF89811">
        <property type="entry name" value="Amyloid beta a4 protein copper binding domain (domain 2)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57362">
        <property type="entry name" value="BPTI-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109843">
        <property type="entry name" value="CAPPD, an extracellular domain of amyloid beta A4 protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00319">
        <property type="entry name" value="APP_CUBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51869">
        <property type="entry name" value="APP_E1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51870">
        <property type="entry name" value="APP_E2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00320">
        <property type="entry name" value="APP_INTRA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00280">
        <property type="entry name" value="BPTI_KUNITZ_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50279">
        <property type="entry name" value="BPTI_KUNITZ_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0186">Copper</keyword>
    <keyword evidence="10 12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9 14" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="2" id="KW-0646">Protease inhibitor</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0722">Serine protease inhibitor</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <keyword evidence="3 14" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 14" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="669"/>
            <end position="691"/>
        </location>
    </feature>
    <feature type="domain" description="E1" evidence="16">
        <location>
            <begin position="17"/>
            <end position="178"/>
        </location>
    </feature>
    <feature type="domain" description="BPTI/Kunitz inhibitor" evidence="15">
        <location>
            <begin position="279"/>
            <end position="329"/>
        </location>
    </feature>
    <feature type="domain" description="E2" evidence="17">
        <location>
            <begin position="342"/>
            <end position="533"/>
        </location>
    </feature>
    <feature type="region of interest" description="GFLD subdomain" evidence="12">
        <location>
            <begin position="17"/>
            <end position="112"/>
        </location>
    </feature>
    <feature type="region of interest" description="CuBD subdomain" evidence="12">
        <location>
            <begin position="120"/>
            <end position="178"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="186"/>
            <end position="240"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="13">
        <location>
            <begin position="186"/>
            <end position="201"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="13">
        <location>
            <begin position="210"/>
            <end position="238"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="62"/>
            <end position="106"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="87"/>
            <end position="94"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="122"/>
            <end position="176"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="133"/>
            <end position="163"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="147"/>
            <end position="175"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022690"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023008"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01217"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50279"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS51869"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS51870"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016119.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="738" mass="84740" checksum="674B256B6A4C03D5" modified="2019-05-08" version="1">MDGIFVALAANAGTGFAVAEPQIAMFCGKLNMHVNIQTGKWEPDTSGTKSCFGTKEEILQYCEEIYPELQITNVVEANQPVSIDNWCKKGKKQCKGHTHIVVPYKCLVGEFVSDVLLVPEKCRFFHKERMDVCESHQHWHTVAKEACLTEGMILHSYGMLLPCGVDQFHGTEYVCCPQTKIVDEALSQEEEDEDEDEDDDYDLYKSEFPTEPDVEEFTGTAVEEDEEEEDEEEEEVVEDRDYYYDNYKVDDYNEETPTESSNDRSLSEKEMISDMKAVCSQEAMTGPCRAVMPRWYFDLYKRKCIRFIYGGCGGNRNNFESEEYCMAVCKKMIPPTPVPTDDVDVYFETPADDNEHARFQKAKEQLEVRHHNRMDRVKKEWEEAEYESKNLPKAERQTLIQHFQAMVKSLEKEAASEKQQLVETHLARVEAMLNDRRRIALENYLAALQADPPRPHRILQALKRYVRAENKDRLHTVRHYQHVLAVDPEKAAQMKSQVMTHLHVIEERMNQSLSLLYKVPYVAEEIQDEIDELLQEQRADMDQFTSSISESQVDVRVSSEESEEIPLQEGKPFRPFQVKTFPAVPGNEDPQPDLYHPMKKGSGMADLDGLIGAEEKVINSKNKVDENVVIDETLDVKEMIFNAERVGGLMDEPDNESSLREDFSLSSSALIGLLVIAVAIATVIVISLVMLRKRQYGTISHGIVEVDPMLTPEERHLNKMQNHGYENPTYKYLEQMQI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMF0</accession>
    <name>A0A452HMF0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Proteasome subunit alpha type-3</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="9">The 26S proteasome consists of a 20S proteasome core and two 19S regulatory subunits. The 20S proteasome core is a barrel-shaped complex made of 28 subunits that are arranged in four stacked rings. The two outer rings are each formed by seven alpha subunits, and the two inner rings are formed by seven beta subunits. The proteolytic activity is exerted by three beta-subunits PSMB5, PSMB6 and PSMB7. Interacts with AURKB. Interacts with CDKN1A. Interacts with MDM2 and RB1. Interacts with the C-terminus of TBXA2R isoform 2. Interacts with DNAJB2.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="10">Belongs to the peptidase T1A family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMF0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018415.1">
        <property type="protein sequence ID" value="ENSGAGP00000016127.1"/>
        <property type="gene ID" value="ENSGAGG00000012088.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019773">
        <property type="term" value="C:proteasome core complex, alpha-subunit complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051603">
        <property type="term" value="P:proteolysis involved in protein catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd03751">
        <property type="entry name" value="proteasome_alpha_type_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.60.20.10:FF:000077">
        <property type="entry name" value="Proteasome subunit alpha type-3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.60.20.10">
        <property type="entry name" value="Glutamine Phosphoribosylpyrophosphate, subunit 1, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029055">
        <property type="entry name" value="Ntn_hydrolases_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050115">
        <property type="entry name" value="Proteasome_alpha"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023332">
        <property type="entry name" value="Proteasome_alpha-type"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001353">
        <property type="entry name" value="Proteasome_sua/b"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11599">
        <property type="entry name" value="PROTEASOME SUBUNIT ALPHA/BETA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00227">
        <property type="entry name" value="Proteasome"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56235">
        <property type="entry name" value="N-terminal nucleophile aminohydrolases (Ntn hydrolases)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51475">
        <property type="entry name" value="PROTEASOME_ALPHA_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0007">Acetylation</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="6 10" id="KW-0647">Proteasome</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00021338"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022942"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00064317"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00808"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016127.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="228" mass="25642" checksum="813A17F944A7F42B" modified="2019-05-08" version="1">MKAVENSSTAIGIRCKDGVVFGVEKLVLSKLYEEGSNKRLFNVDRHVGMAVAGLLADARSLADIAREEASNFRSNYGYNIPLKHLADRVAMYVHAYTLYSAVRPFGCSFMLGSYDEDDGAQLYMIDPSGVSYGYWGCAIGKARQAAKTEIEKLQMKEMTCRDVVKEVAKIIYIVHDEVKDKSFELELSWVGEITKGKHEIVPKDIREEAEKYAKESLKEEDESDEDNM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMF4</accession>
    <name>A0A452HMF4_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="6">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the PACS family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HMF4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018416.1">
        <property type="protein sequence ID" value="ENSGAGP00000016128.1"/>
        <property type="gene ID" value="ENSGAGG00000012108.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0044325">
        <property type="term" value="F:transmembrane transporter binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072659">
        <property type="term" value="P:protein localization to plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019381">
        <property type="entry name" value="PACS1/2_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057541">
        <property type="entry name" value="PACS1/2_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13280">
        <property type="entry name" value="PHOSPHOFURIN ACIDIC CLUSTER SORTING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13280:SF15">
        <property type="entry name" value="PHOSPHOFURIN ACIDIC CLUSTER SORTING PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25332">
        <property type="entry name" value="C2_PACS_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10254">
        <property type="entry name" value="Pacs-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Phosphofurin acidic cluster sorting protein 1/2 N-terminal C2" evidence="5">
        <location>
            <begin position="21"/>
            <end position="180"/>
        </location>
    </feature>
    <feature type="domain" description="Phosphofurin acidic cluster sorting protein 1/2 C-terminal" evidence="4">
        <location>
            <begin position="467"/>
            <end position="806"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="181"/>
            <end position="242"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="293"/>
            <end position="387"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="405"/>
            <end position="461"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="683"/>
            <end position="731"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="782"/>
            <end position="802"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="186"/>
            <end position="198"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="368"/>
            <end position="387"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="449"/>
            <end position="458"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="3">
        <location>
            <begin position="683"/>
            <end position="710"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="3">
        <location>
            <begin position="721"/>
            <end position="731"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="3">
        <location>
            <begin position="783"/>
            <end position="799"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00008590"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="Pfam" id="PF10254"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="Pfam" id="PF25332"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016128.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="832" mass="92989" checksum="55C51B0C35FFB339" modified="2019-05-08" version="1">MAAERGRLSFPGGAGVLSRPVPMNLFATWEIDGSSPSCVPRLCTLTLKKLVVLKELDKELISVVIAVKMQGSKRILRSHEIVLPPSGHVETELALTFSLQYPHFLKREGNKLQIMLQRRKRYKNRTILGYKTLAVGSINMAEVMQHPTEGGQVLSLFSNIKETTAKVAEIWIFSLSSQPIDHEDSTMQPSQKIKSTDNYSEEEYESFSSEQEASDDAVHGQDLDDDDYELGKPKKQRRSIVRTTSITRQQNFKQKVVALLRRFKVSDEVLDSEQDPAEHVPEVEEDLDLLYDTLDIDNPSDSGPEMEDDDSVLSTPKPKLKPYFEGLSHSSSQTEVGSIHSIRSQREPSSPVEVPEKTKSLGSKHPSDSISDTVSYEPSQQDSQEAELSTLDVFIEKLPPSGKITKTESLIIPSTSRPEGKQTGRRGRSTSLKERQPARPQNERANSLDNERSPDTRHHLQIPRKTVYDQLNHILISDDQLPENIILVNTSDWQGQFLSDVLQKHTLPVVCTCSSADVQAAFSTIVSRIQRYCNCNSQPPNPIKIAVAGAQNYLSAVLRLFVEQLSHKTPDWLSYMRFLIIPLGSHPVAKYLGSVDYRYNNFFQDFAWRDLFNKLEAQTSVQETPDIVSRITQYIAGANCAHQLPIAEAMLTYKQKSPDEESSQKFIPFVGVVKVGIVEQSSATSGDSDDAAPSSSSVLSSTPPSVSPAVKEASPTPPSSPSVTGSFSSSSQTLGAELMGLQVDYWIAAQPMEKKRDPEKKDPSTAKNTLKCTFRSLQVSRLPTSGETTSTPTMSMTVVTKEKNKKDKPLDFYYPFVGRKSWRRRRMRSWWS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HMF8</accession>
    <name>A0A452HMF8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">SWIM-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HMF8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018421.1">
        <property type="protein sequence ID" value="ENSGAGP00000016133.1"/>
        <property type="gene ID" value="ENSGAGG00000012100.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031462">
        <property type="term" value="C:Cul2-RING ubiquitin ligase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007527">
        <property type="entry name" value="Znf_SWIM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048370">
        <property type="entry name" value="ZSWIM4-8_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22619">
        <property type="entry name" value="ZINC FINGER SWIM DOMAIN CONTAINING PROTEIN 4, 5, 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22619:SF3">
        <property type="entry name" value="ZINC FINGER SWIM DOMAIN-CONTAINING PROTEIN 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21055">
        <property type="entry name" value="ZSWIM4-8_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50966">
        <property type="entry name" value="ZF_SWIM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0862">Zinc</keyword>
    <keyword evidence="2 4" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="SWIM-type" evidence="6">
        <location>
            <begin position="154"/>
            <end position="191"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="1"/>
            <end position="31"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="1"/>
            <end position="25"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00325"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50966"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016133.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1108" mass="125082" checksum="E83BC5ED5A1D22FB" modified="2019-05-08" version="1">PHSVPSGAQSNRGTGEVTRPTSSPRGSPESLLDIAARKVAEKWPFQRVEERFERIPEPVQRRIVYWSFPRSEREICMYSSFNTGAEDPAAPGGAAASSSPAGAADAAADENRLPFRRGIALLDGSCVDNVLQVGFHLSGTVTDPAMQSEPETVCKVAISFDRCKITSVTCSCGNKDIFYCAHVVALSLYRIRKPDQVKLHLPISETLFQMNRDQLQKFVQYLITMHHTEVLPTAQKLADEILSQNSEINQVHGAPDPTAGASIDDENCWHLDEEQVQEQVKLFLSQGGYHGSGKQLNLLFAKVREMLKMRDSNGARMLTLITEQFMADPRLSLWRQQGTAMTDKYRQLWDELGALWMCIVLNPHCKLEQKASWLKQLKKWNCVDVCPWEDGNHGNELPNLTNALPQDSLNRPHRTVFTRAIEACDLHWQDSHLQHIISSDLYTNYCYHDDTENSLFDSHGWPLWHEHVPTACARVDALRSHGYPREALRLAIAIVNTLRRQQQKQLDMFRAQKKELLHKGVTSITNLEGWVGHPLDPVGTLFSSLMEACRVDDEKNMGQCKSLEYQHLPSHKFLEEGESYLTLAVEVALIGLGQQRIMPDGLYAQEKVCRNEEQLISKLQEIELDDTLVKIFRKQAVFLLEAGPYSGLGEIIHRESVPMHTFAKYLFTSLLPHDAELAYKTALRAMRLLVLESTAPSGDMSRPHHIASVVPNRYPRWFTLSHIESQQCELASTMLTAAKGDVRRLETVLESIQKNIHSSSHIFKLAQDAFKIATLMDSLPDITLLKVSLELGLQVMRMTLSTLNWRRREMVRWLVTCATEVGVYALDSIMQNWFTLFTPTEATSIVATTVMSNSTIVRLHLDCLQQEKLASSARTLALQCAMKDPQNCALSALTLCEKDHIAFETAYQIVLDAATKGMSYTQLFTIARYMEHRGYPMRAYKLATLAMNHLNLSYNQDTHPAINDVLWACALSHSLGKNELAAIIPLVVKSVKCATVLSDILRRCTLTTPGMVGLHGRRNSGKLMSLDKAPLRQLLDATIGAYINTTHSRLTHISPRHYSEFIEFLSKARETFLMAHDGHIQFTQFIDNLKQIYKGKKKLMMLVRERFG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPA4</accession>
    <name>A0A452IPA4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">thioredoxin-disulfide reductase (NADPH)</fullName>
            <ecNumber evidence="2">1.8.1.9</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="11">
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </cofactor>
        <text evidence="11">Binds 1 FAD per subunit.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1 13">Belongs to the class-I pyridine nucleotide-disulfide oxidoreductase family.</text>
    </comment>
    <dbReference type="EC" id="1.8.1.9" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452IPA4"/>
    <dbReference type="STRING" id="38772.ENSGAGP00000029670"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033691.1">
        <property type="protein sequence ID" value="ENSGAGP00000029670.1"/>
        <property type="gene ID" value="ENSGAGG00000021420.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050660">
        <property type="term" value="F:flavin adenine dinucleotide binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004362">
        <property type="term" value="F:glutathione-disulfide reductase (NADPH) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004791">
        <property type="term" value="F:thioredoxin-disulfide reductase (NADPH) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045454">
        <property type="term" value="P:cell redox homeostasis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034599">
        <property type="term" value="P:cellular response to oxidative stress"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006749">
        <property type="term" value="P:glutathione metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.50.50.60:FF:000190">
        <property type="entry name" value="Thioredoxin reductase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.390.30:FF:000004">
        <property type="entry name" value="Thioredoxin reductase 1, cytoplasmic"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.390.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.50.50.60">
        <property type="entry name" value="FAD/NAD(P)-binding domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036188">
        <property type="entry name" value="FAD/NAD-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023753">
        <property type="entry name" value="FAD/NAD-binding_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016156">
        <property type="entry name" value="FAD/NAD-linked_Rdtase_dimer_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046952">
        <property type="entry name" value="GSHR/TRXR-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001100">
        <property type="entry name" value="Pyr_nuc-diS_OxRdtase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004099">
        <property type="entry name" value="Pyr_nucl-diS_OxRdtase_dimer"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012999">
        <property type="entry name" value="Pyr_OxRdtase_I_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006338">
        <property type="entry name" value="Thioredoxin/glutathione_Rdtase"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01438">
        <property type="entry name" value="TGR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR42737">
        <property type="entry name" value="GLUTATHIONE REDUCTASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR42737:SF7">
        <property type="entry name" value="THIOREDOXIN-DISULFIDE REDUCTASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07992">
        <property type="entry name" value="Pyr_redox_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02852">
        <property type="entry name" value="Pyr_redox_dim"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000350">
        <property type="entry name" value="Mercury_reductase_MerA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00368">
        <property type="entry name" value="FADPNR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00411">
        <property type="entry name" value="PNDRDTASEI"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51905">
        <property type="entry name" value="FAD/NAD(P)-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55424">
        <property type="entry name" value="FAD/NAD-linked reductases, dimerisation (C-terminal) domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00076">
        <property type="entry name" value="PYRIDINE_REDOX_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="4 11 13" id="KW-0274">FAD</keyword>
    <keyword evidence="3 13" id="KW-0285">Flavoprotein</keyword>
    <keyword evidence="11" id="KW-0520">NAD</keyword>
    <keyword evidence="5" id="KW-0521">NADP</keyword>
    <keyword evidence="11" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="7 13" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="9 13" id="KW-0676">Redox-active center</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0712">Selenocysteine</keyword>
    <feature type="domain" description="FAD/NAD(P)-binding" evidence="15">
        <location>
            <begin position="30"/>
            <end position="365"/>
        </location>
    </feature>
    <feature type="domain" description="Pyridine nucleotide-disulphide oxidoreductase dimerisation" evidence="14">
        <location>
            <begin position="385"/>
            <end position="496"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="10">
        <location>
            <position position="487"/>
        </location>
    </feature>
    <feature type="binding site" evidence="11">
        <location>
            <position position="85"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="11">
        <location>
            <position position="149"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="11">
        <location>
            <begin position="215"/>
            <end position="222"/>
        </location>
        <ligand>
            <name>NAD(+)</name>
            <dbReference type="ChEBI" id="CHEBI:57540"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="11">
        <location>
            <position position="307"/>
        </location>
        <ligand>
            <name>NAD(+)</name>
            <dbReference type="ChEBI" id="CHEBI:57540"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="11">
        <location>
            <position position="349"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="disulfide bond" description="Redox-active" evidence="12">
        <location>
            <begin position="76"/>
            <end position="81"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007532"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00012610"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022630"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022827"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022857"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022933"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023284"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PIRSR" id="PIRSR000350-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PIRSR" id="PIRSR000350-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PIRSR" id="PIRSR000350-4"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="RuleBase" id="RU003691"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="Pfam" id="PF02852"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="Pfam" id="PF07992"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029670.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="512" mass="55974" checksum="A41226BDF0C85329" modified="2019-05-08" version="1">MAALCRGRKRFQWEPVLCRTARVLSGKNEYDLLVIGGGSGGLACAKEAAQFGKKVAVLDYVEPSPKGTKWGLGGTCVNVGCIPKKLMHQASLLGSALKDAQHYGWNISHPVRHDWPVMAEAIQNYVKSLNWGHRVQLQDKKVKYFNLTGSLMDAHTVRGLTNAGKETIITAENIVIATGGRPKYPAHIAGALEYGITSDDIFWLKESPGKTLVVGASYVSLECAGFLTGIGQDTTVMVRSIPLRGFDQQMSFLVTDYMESCGTRFLKKCTPSKVEKLKSGKLQVTWKHADSGKEEMDDFDTVMWAVGRAPDTKTLNLETIGVKISSETGKIIVDASEATSVPHIYAIGDITEGRPELTPTAIAAGKLLAQRLFGQSSELMDYDNVPTTVFTPLEYGCVGLSEEKAVEHYGLDNIEVYHAYYKPLEFTVAERDATQCYIKMVCLRKKEQRILGLHFIGPNAGEVIQGFALGMKCGATYAQMMKTVGIHPTCAEEVTKLRITKRSGLDAKVTGC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPA8</accession>
    <name>A0A452IPA8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">NACHT domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPA8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033855.1">
        <property type="protein sequence ID" value="ENSGAGP00000029816.1"/>
        <property type="gene ID" value="ENSGAGG00000021527.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006954">
        <property type="term" value="P:inflammatory response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045087">
        <property type="term" value="P:innate immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007111">
        <property type="entry name" value="NACHT_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041267">
        <property type="entry name" value="NLRP_HD2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050637">
        <property type="entry name" value="NLRP_innate_immun_reg"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041075">
        <property type="entry name" value="NOD1/2_WH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45690">
        <property type="entry name" value="NACHT, LRR AND PYD DOMAINS-CONTAINING PROTEIN 12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45690:SF19">
        <property type="entry name" value="NACHT, LRR AND PYD DOMAINS-CONTAINING PROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05729">
        <property type="entry name" value="NACHT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17776">
        <property type="entry name" value="NLRC4_HD2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17779">
        <property type="entry name" value="WHD_NOD2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52047">
        <property type="entry name" value="RNI-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50837">
        <property type="entry name" value="NACHT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-0395">Inflammatory response</keyword>
    <keyword evidence="4" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="NACHT" evidence="8">
        <location>
            <begin position="14"/>
            <end position="332"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023198"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029816.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="510" mass="59435" checksum="B916CDBB81AB618B" modified="2019-05-08" version="1">CMLKSHLLYWDFNLTVVLLGAAGIGKTMTARKIMLDWAAGKLYQKKFDYVFYINCRKINLVTAQQSMADLLLNNFPDGVRPTKEIFEKPEKLLFIIDGFDELNSVLSMEEVSLWSDPFEKKPVKTVLCSLLRKEVLQKSFLLITSRPTALEKLKQNLKFPCYAEIRGFSGDDRKEYFHKFFGDEKKATQAFNFVKENEMLFTMCFVPIVCWIICTVMKQQMERKEDLAQTSKTTTSVYLLFLTSLLTDHFTDSVQQSKTTLWKLCSLAADGILERKILFEEGDLKKHNLSLSTIHTLFLNKSIFQKDTECESVYSFIHLSFQEFFAALFYVLEEDEKTMKKSLSSKKDVNIRFLFGLLNKERLNDMEKKLHCKTSSAIKLDLLKWALRNCLNQLELFHCLYESQEEEFAKSAMDNFQEIRLEHIRLSTMDEIALSFCVKNCCSKQSLYLCNCTLGIGEREEGWIAKLRKWLFPLEPHCWCTLGSLLHHRHCPKAGLIHKPRCQWLQFCSV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPA9</accession>
    <name>A0A452IPA9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Protein Wnt</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="11">Ligand for members of the frizzled family of seven transmembrane receptors.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 11">Secreted</location>
            <location evidence="1 11">Extracellular space</location>
            <location evidence="1 11">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 11">Belongs to the Wnt family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPA9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033696.1">
        <property type="protein sequence ID" value="ENSGAGP00000029675.1"/>
        <property type="gene ID" value="ENSGAGG00000021418.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005125">
        <property type="term" value="F:cytokine activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005109">
        <property type="term" value="F:frizzled binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060070">
        <property type="term" value="P:canonical Wnt signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045165">
        <property type="term" value="P:cell fate commitment"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030182">
        <property type="term" value="P:neuron differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd19354">
        <property type="entry name" value="Wnt_Wnt9b"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.2460.20:FF:000002">
        <property type="entry name" value="Protein Wnt"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.2460.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005817">
        <property type="entry name" value="Wnt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043158">
        <property type="entry name" value="Wnt_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018161">
        <property type="entry name" value="Wnt_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12027:SF84">
        <property type="entry name" value="PROTEIN WNT-9B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12027">
        <property type="entry name" value="WNT RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00110">
        <property type="entry name" value="wnt"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01349">
        <property type="entry name" value="WNTPROTEIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00097">
        <property type="entry name" value="WNT1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00246">
        <property type="entry name" value="WNT1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 11" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="9" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
    <keyword evidence="6 11" id="KW-0879">Wnt signaling pathway</keyword>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1"/>
            <end position="33"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="248"/>
            <end position="267"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="249"/>
            <end position="267"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005683"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022687"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="RuleBase" id="RU003500"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029675.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="346" mass="38542" checksum="55A6D1823F59C5F4" modified="2019-05-08" version="1">PDLLSHPTPSQSLGLTGKEGLTHLPSLGPSANPAQGKAHLKQCDLLKLSRKQKRLCQREPGLAETLRDAIRLSIMECQYQFRYERWNCSLDGRTSLLKRGFKETAFLYAVSSAALTHSLARACSAGRMERCTCDDSPDLENRKAWQWGVCGDNLKYSTKFLKNFLGQKKIGKDLRAKVDIHNTNVGIKAVKNGLKTTCKCHGVSGSCAVRTCWKQLSPFHETGKILKLRFDDAVKIFSATNDAVGHSELVSTQRRSHSPKSPATPRSTDLVYVEDSPSFCRPSKYSLGTAGRTCSREASCDSLCCGRGYNTQSRMVTFSCHCQVQWCCYVECQQCMQEEVIYSCKQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPB2</accession>
    <name>A0A452IPB2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Nck-associated protein 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="6">Cell projection</location>
            <location evidence="6">Lamellipodium membrane</location>
            <topology evidence="6">Single-pass membrane protein</topology>
            <orientation evidence="6">Cytoplasmic side</orientation>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="7">Belongs to the HEM-1/HEM-2 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPB2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034001.1">
        <property type="protein sequence ID" value="ENSGAGP00000029942.1"/>
        <property type="gene ID" value="ENSGAGG00000019065.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031258">
        <property type="term" value="C:lamellipodium membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031209">
        <property type="term" value="C:SCAR complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030031">
        <property type="term" value="P:cell projection assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030866">
        <property type="term" value="P:cortical actin cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048812">
        <property type="term" value="P:neuron projection morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019137">
        <property type="entry name" value="Nck-associated_protein-1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12093">
        <property type="entry name" value="NCK-ASSOCIATED PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12093:SF11">
        <property type="entry name" value="NCK-ASSOCIATED PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09735">
        <property type="entry name" value="Nckap1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="1" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="5" id="KW-0966">Cell projection</keyword>
    <keyword evidence="4" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="3" id="KW-1133">Transmembrane helix</keyword>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="509"/>
            <end position="534"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="520"/>
            <end position="534"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00037839"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00037947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00039689"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029942.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="997" mass="113668" checksum="49F293BDA24552A4" modified="2019-05-08" version="1">QNDDLIITYTTLMILLSRIEERKAIIGLYNYAHEMTHGASDREYPRLGQMIVDYENPLKKMMEEFVPHSKSLSDALISLQMVYPRRNLSADQWRNAQLLSLISAPSTMLNPAQSDTMPCEYLSLDAMEKWIIFGFILCHGILNSDATALNLWKLALQSSSCLSLFRDEVFHIHKAAEDLFVNIRGYNKRINDIRECKEAAVSHAGSMHRERRKFLRSALKELATVLSDQPGLLGPKALFVFMALSFARDEIIWLLRHADNMPKKSADDFIDKHIAELIFYMEELRAHVRKYGPVMQRYYVQYLSGFDAVVLNELVQNLSVCPEDESIIMSSFVNTMTSLSVKQVEDGEVFDFRGMRLDWFRLQAYTSVSKASLSLADHRELGKMMNTIIFHTKMVDSLVEMLVETSDLSIFCFYSRAFEKMFQQCLELPSQSRYSIAFPLLCTHFMSCTHELCPEERHHIGDRSLSLCNMFLDEMAKQARNLITDICTEQCTLSDQLLPKHCAKTISQAVNKKSKKQTGKKGEPEREKPGVESMRKNRLVVTNLDKLHTALSELCFSINYVPNMVVWEHTFTPREYLTSHLEIRFTKSIVGMTMYNQATQEIAKPSELLTSVRAYMTVLQSIENYVQIDITRVFNNVLLQQTQHLDSHGEPTITSLYTNWYLETLLRQVSNGHIAYFPAMKAFVNLPTENELTFNAEEYSDISEMRALSELLGPYGMKFLSESLMWHISSQVAELKKLVVENVEVLTQMRTSFDKPDQMAALFKRLSSVDSVLKRMTIIGVILSFRSLAQEALRDVLSYHIPFLVSSIEDFKDHIPRETDMKVAMNVYELSSAAGLPCEIDPALVVALSSQKSENISPEEEYKIACLLMVFVAVSLPTLASNVMSQYSPAIEGHCNNIHCLAKAINQIAAALFTIHKGSIEDRLKEFLALASSSLLKIGQETDKTTTRNRESVYLLLDMIVQESPFLTMDLLESCFPYVLLRNAYHAVYKQSVTSSA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPB7</accession>
    <name>A0A452IPB7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Uronyl 2-sulfotransferase</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Golgi apparatus membrane</location>
            <topology evidence="1">Single-pass type II membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the sulfotransferase 3 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPB7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033868.1">
        <property type="protein sequence ID" value="ENSGAGP00000029828.1"/>
        <property type="gene ID" value="ENSGAGG00000021514.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000139">
        <property type="term" value="C:Golgi membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008146">
        <property type="term" value="F:sulfotransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000580">
        <property type="entry name" value="uronyl 2-sulfotransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007734">
        <property type="entry name" value="Heparan_SO4_2-O-STrfase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005331">
        <property type="entry name" value="Sulfotransferase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12129">
        <property type="entry name" value="HEPARAN SULFATE 2-O-SULFOTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12129:SF15">
        <property type="entry name" value="URONYL 2-SULFOTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03567">
        <property type="entry name" value="Sulfotransfer_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="8" id="KW-0472">Membrane</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0735">Signal-anchor</keyword>
    <keyword evidence="3" id="KW-0808">Transferase</keyword>
    <keyword evidence="4" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6" id="KW-1133">Transmembrane helix</keyword>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1"/>
            <end position="21"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="306"/>
            <end position="347"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="1"/>
            <end position="15"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="11">
        <location>
            <begin position="325"/>
            <end position="340"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004323"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010569"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00071728"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029828.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="347" mass="41203" checksum="BE2CA64A500B8AED" modified="2019-05-08" version="1">PAGEREGREGGRREVASYGGSRQPNSIRVEYCTTLNEPVLPFPSQVVYNRVGKCGSRTVVLLLRILSERHGFNLVTSDIHNKTRLTKNEQMELIKNISTAEQPYLFTRHVHFLNFSRFGGEQPVYINIIRDPANRFLSNYFFRRFGDWRGEQNHMIRTPSMRQEERYLDINICILENYPECSNPRLFYIIPYFCGQHPRCREPGEWALERAKLNVNENFLLVGILEELEDVLLLLERFLPHYFKDVLSIYKNPEHRKLGNLTVTVKKTVPSPEAIQILYQRMRYEYEFYYYVKEQFHLLKRKFGLKSHSSEPPPKPEFFIPSPLETEEPIDDEEQDDEKWLEDIYKR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPC3</accession>
    <name>A0A452IPC3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Nck-associated protein 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="6">Cell projection</location>
            <location evidence="6">Lamellipodium membrane</location>
            <topology evidence="6">Single-pass membrane protein</topology>
            <orientation evidence="6">Cytoplasmic side</orientation>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="7">Belongs to the HEM-1/HEM-2 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPC3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034011.1">
        <property type="protein sequence ID" value="ENSGAGP00000029952.1"/>
        <property type="gene ID" value="ENSGAGG00000019065.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031258">
        <property type="term" value="C:lamellipodium membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031209">
        <property type="term" value="C:SCAR complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030031">
        <property type="term" value="P:cell projection assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030866">
        <property type="term" value="P:cortical actin cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048812">
        <property type="term" value="P:neuron projection morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019137">
        <property type="entry name" value="Nck-associated_protein-1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12093">
        <property type="entry name" value="NCK-ASSOCIATED PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12093:SF11">
        <property type="entry name" value="NCK-ASSOCIATED PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09735">
        <property type="entry name" value="Nckap1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="1" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="5" id="KW-0966">Cell projection</keyword>
    <keyword evidence="4" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="3" id="KW-1133">Transmembrane helix</keyword>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="520"/>
            <end position="545"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="531"/>
            <end position="545"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00037839"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00037947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00039689"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029952.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1008" mass="114960" checksum="C7D94F0FD4FBC4E5" modified="2019-05-08" version="1">LKKELEKDTHPYSGHLIITYTTLMILLSRIEERKAIIGLYNYAHEMTHGASDREYPRLGQMIVDYENPLKKMMEEFVPHSKSLSDALISLQMVYPRRNLSADQWRNAQLLSLISAPSTMLNPAQSDTMPCEYLSLDAMEKWIIFGFILCHGILNSDATALNLWKLALQSSSCLSLFRDEVFHIHKAAEDLFVNIRGYNKRINDIRECKEAAVSHAGSMHRERRKFLRSALKELATVLSDQPGLLGPKALFVFMALSFARDEIIWLLRHADNMPKKSADDFIDKHIAELIFYMEELRAHVRKYGPVMQRYYVQYLSGFDAVVLNELVQNLSVCPEDESIIMSSFVNTMTSLSVKQVEDGEVFDFRGMRLDWFRLQAYTSVSKASLSLADHRELGKMMNTIIFHTKMVDSLVEMLVETSDLSIFCFYSRAFEKMFQQCLELPSQSRYSIAFPLLCTHFMSCTHELCPEERHHIGDRSLSLCNMFLDEMAKQARNLITDICTEQCTLSDQLLPKHCAKTISQAVNKKSKKQTGKKGEPEREKPGVESMRKNRLVVTNLDKLHTALSELCFSINYVPNMVVWEHTFTPREYLTSHLEIRFTKSIVGMTMYNQATQEIAKPSELLTSVRAYMTVLQSIENYVQIDITRVFNNVLLQQTQHLDSHGEPTITSLYTNWYLETLLRQVSNGHIAYFPAMKAFVNLPTENELTFNAEEYSDISEMRALSELLGPYGMKFLSESLMWHISSQVAELKKLVVENVEVLTQMRTSFDKPDQMAALFKRLSSVDSVLKRMTIIGVILSFRSLAQEALRDVLSYHIPFLVSSIEDFKDHIPRETDMKVAMNVYELSSAAGLPCEIDPALVVALSSQKSENISPEEEYKIACLLMVFVAVSLPTLASNVMSQYSPAIEGHCNNIHCLAKAINQIAAALFTIHKGSIEDRLKEFLALASSSLLKIGQETDKTTTRNRESVYLLLDMIVQESPFLTMDLLESCFPYVLLRNAYHAVYKQSVTSSA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPC5</accession>
    <name>A0A452IPC5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">NACHT domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Inflammasome</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the NLRP family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPC5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033815.1">
        <property type="protein sequence ID" value="ENSGAGP00000029778.1"/>
        <property type="gene ID" value="ENSGAGG00000021504.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006954">
        <property type="term" value="P:inflammatory response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045087">
        <property type="term" value="P:innate immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001611">
        <property type="entry name" value="Leu-rich_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029495">
        <property type="entry name" value="NACHT-assoc"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007111">
        <property type="entry name" value="NACHT_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041267">
        <property type="entry name" value="NLRP_HD2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050637">
        <property type="entry name" value="NLRP_innate_immun_reg"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041075">
        <property type="entry name" value="NOD1/2_WH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45690">
        <property type="entry name" value="NACHT, LRR AND PYD DOMAINS-CONTAINING PROTEIN 12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45690:SF19">
        <property type="entry name" value="NACHT, LRR AND PYD DOMAINS-CONTAINING PROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14484">
        <property type="entry name" value="FISNA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13516">
        <property type="entry name" value="LRR_6"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05729">
        <property type="entry name" value="NACHT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17776">
        <property type="entry name" value="NLRC4_HD2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17779">
        <property type="entry name" value="WHD_NOD2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00368">
        <property type="entry name" value="LRR_RI"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52047">
        <property type="entry name" value="RNI-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50837">
        <property type="entry name" value="NACHT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9" id="KW-1271">Inflammasome</keyword>
    <keyword evidence="8" id="KW-0395">Inflammatory response</keyword>
    <keyword evidence="5" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="7" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="NACHT" evidence="10">
        <location>
            <begin position="112"/>
            <end position="244"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004110"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008665"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023198"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023233"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029778.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="877" mass="99528" checksum="5C7020242F9EF5EE" modified="2019-05-08" version="1">MNLVAACGRSAESTGLADPPQACCRMQHACCPHGNQQSAPCGLPNARLGENVKLNSRYTKLIIVDSHGYEKNREHEIIATGWKHTEAMTERASSVITMGTLFKPYKDGQTPQIVVLVGAAGIGKTITSKKIMLDWAAGELYKEMFDYVFYINCREMNLLTKEVSVADMILKHCPNNKTSINHILGQSDKLLFIIDELRFSFDQSEDNLCSERWEKKPVEIILSSLFRKTVLPESYLLITTRPAALEKLGRCERYAEILGFSEADRKEYFHKFFGNEKQARQAYRFVRENEILFTMCFVPIMCWIICTVLKQQLEKGEDLAQASKTATGMYVLYLSSLVNPPGSNSKPCKKASLRGLCSLAADGIWKQKILFEEEEIKMLGLDQADALPLFLNENIFQKDIDTECLYSFIHLSFQEFFAALFYVLEEDETMEDSGTPKKDVKTLLENYGNSRNYLMLTVRFLYGLLNEKSIKDMEEKFGCKISPKIKLDLIKWIQTYKQTVLSVPSVYDKIICQLEEFRCLYEIQDENFMKNALEPFTKIVICRNNLTPMDQIVLSFCVNNCHRLVSLEMYACTFRFEDRKKEEIELKSVSRDQDRTFLFSLSRDQHQDELKHSRVFLLCQALKDPNCKLKKLWKDRTDRTAHFLICASISCSLRSCKLTADCCGDLSSVLSTNQFLTDLDLTANEVRNTGVQLLCEGLKHQNCKLQRLSLGSCNISDDDCTDLAAALRTSQSLTELDLSFNHYLRDSGVQLLCEGLIQPTCKLQSLKLNNCNITHVSCKDLAAVLKTSRRLIELDVGTNELQEAGVQLLCEALKHPNCRLLGLCRLTAACCGDLSSALSTSQTLAELNLQDNRLEDSGVQQLCEALKHPNCKLQKIK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPD3</accession>
    <name>A0A452IPD3_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="11">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="domain">
        <text evidence="6">The CKK domain binds microtubules.</text>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the CAMSAP1 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000033888.1">
        <property type="protein sequence ID" value="ENSGAGP00000029845.1"/>
        <property type="gene ID" value="ENSGAGG00000021541.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036449">
        <property type="term" value="C:microtubule minus-end"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051011">
        <property type="term" value="F:microtubule minus-end binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030507">
        <property type="term" value="F:spectrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031122">
        <property type="term" value="P:cytoplasmic microtubule organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007026">
        <property type="term" value="P:negative regulation of microtubule depolymerization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031175">
        <property type="term" value="P:neuron projection development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.20.360:FF:000001">
        <property type="entry name" value="Calmodulin-regulated spectrin-associated protein 3 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.20.360">
        <property type="entry name" value="CKK domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032940">
        <property type="entry name" value="CAMSAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031372">
        <property type="entry name" value="CAMSAP_CC1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022613">
        <property type="entry name" value="CH_CAMSAP_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001715">
        <property type="entry name" value="CH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036872">
        <property type="entry name" value="CH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014797">
        <property type="entry name" value="CKK_CAMSAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038209">
        <property type="entry name" value="CKK_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011033">
        <property type="entry name" value="PRC_barrel-like_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21595:SF2">
        <property type="entry name" value="CALMODULIN-REGULATED SPECTRIN-ASSOCIATED PROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21595">
        <property type="entry name" value="PATRONIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17095">
        <property type="entry name" value="CAMSAP_CC1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11971">
        <property type="entry name" value="CAMSAP_CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08683">
        <property type="entry name" value="CAMSAP_CKK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01051">
        <property type="entry name" value="CAMSAP_CKK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47576">
        <property type="entry name" value="Calponin-homology domain, CH-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50346">
        <property type="entry name" value="PRC-barrel domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50021">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51508">
        <property type="entry name" value="CKK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="5" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="3 6" id="KW-0493">Microtubule</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="8">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019525654" evidence="8">
        <location>
            <begin position="23"/>
            <end position="1239"/>
        </location>
    </feature>
    <feature type="domain" description="Calponin-homology (CH)" evidence="9">
        <location>
            <begin position="77"/>
            <end position="200"/>
        </location>
    </feature>
    <feature type="domain" description="CKK" evidence="10">
        <location>
            <begin position="1100"/>
            <end position="1234"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="54"/>
            <end position="81"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="410"/>
            <end position="558"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="582"/>
            <end position="638"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="682"/>
            <end position="977"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1046"/>
            <end position="1106"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="466"/>
            <end position="477"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="478"/>
            <end position="497"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="602"/>
            <end position="616"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="721"/>
            <end position="743"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="7">
        <location>
            <begin position="744"/>
            <end position="760"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="795"/>
            <end position="808"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="7">
        <location>
            <begin position="877"/>
            <end position="889"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="905"/>
            <end position="977"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00841"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS51508"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029845.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1239" mass="135019" checksum="C42CBF4597CFF8AB" modified="2019-05-08" version="1">MGAHLAVIDSLMIAFAVESTHSLSAPPGTDLGAGSWEQKLLCWVDTVNRKLQESTEREGSQQLVPGTEGQSQSQSPASGPKCPTRWYWKLVPIRYRKDKVLPKQTPCFTPVMGMKDLANGGAIAATIHYYCPDVVRLEDVCLKETMSVADSLYNLQLIQEFCIKYLGGCCPLALEDLLYVPPVLKINIGVFLAELFLCFEVLKPDFVRPKELRGLKDPPAVSDSLTPSSGNSNSGSLVFSFLHPLLPGGQPQSPLRGSLGSMHHSTSMSHVEGGFGKSWSKKQLSHPLSQAVSFSIPFGLDSDVDIVMGNPVGMLRSISSDSLAPVPTRLARSPGPLPEDAAQRLAKDEAPNGPVAPHRTVHKVALSAKGVVEREGPPVENGLADGYPDLPTIEEALQIIHSSERLHPEGAADGFYLHSPEPAKPTQAPEPTPALAVYRFHSGPPNGRAEAPTPPGNGSLDSDAEDPARLPGTKDDTSSGLSSLSSQPDSAGSSGAGVRMTSFAERKKKLAPPEGKTLQKSTGESSEAGAVPPEDSPGRSPALSSEMSQLGARLEEKRRAIEAQKKRIEAIFAKHRQRLGKSAFLQLKRRDGGEEEEEEEGEAKLSLEERLARLEAEEGGSLQVQAEEEGPGQGCPEKQVTFSSEIKGAPLDENLGDYNRAVAKLNTALSSLQLDMHRLSQQQQRLLQEKKPSQAWVIPAPKGPAPRASREFVPPRSVELSSSPSPSPSRXXXXXXXXXXPAAPRSPQPAPKKAAPTPPKSPKHARPVELKLPPLTRVLTPPHNVDTLPHLRKFSPSQVPMQTRSSIHFSEEVAGPEQPLESPELETQGNLRPVASVVPQGGVGSARVSGDGTSDVSSPGERRSSLIEIPLSSLQAEEGDGDDSLEESLTEVMDMEPRAGLGFFFKDDEKPEDEMAQKRASLLERQQRRNEEARQRKQWLEAEKEQKEEAARLQAEERPRPEEEASPRRGDFTRQEYQRRHQLKLMEDLDKVLRQKPTTVRALKKGRPKTVFCDDSALARSPVKGLLGSRLSKVYSQSTLSLSTVANEPGNSLTIKRPSRAASPSGLMSPSRLLGSQSRERDWENASTASSPASVPEYTGPKLYKEPSAKSNKHIIHNALSHCCLAGRVNEPHKNKILEEMEKSKAHHFLILFRDSSCQFRALYTPAGETEELTRLTGYGPRTISQAMIEGIYKYNSDRKRFTQIPAKTMSMNVDAITIQGHLWQTKKPGTPKKPGTPK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPD4</accession>
    <name>A0A452IPD4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">UDP-glucose glycoprotein glucosyltransferase 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="pathway">
        <text evidence="2">Protein modification; protein glycosylation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum lumen</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the glycosyltransferase 8 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000033825.1">
        <property type="protein sequence ID" value="ENSGAGP00000029788.1"/>
        <property type="gene ID" value="ENSGAGG00000021377.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000033833.1">
        <property type="protein sequence ID" value="ENSGAGP00000029795.1"/>
        <property type="gene ID" value="ENSGAGG00000021377.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00378"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005788">
        <property type="term" value="C:endoplasmic reticulum lumen"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003980">
        <property type="term" value="F:UDP-glucose:glycoprotein glucosyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051082">
        <property type="term" value="F:unfolded protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036503">
        <property type="term" value="P:ERAD pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0018279">
        <property type="term" value="P:protein N-linked glycosylation via asparagine"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040497">
        <property type="entry name" value="Glyco_transf_24"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009448">
        <property type="entry name" value="UDP-g_GGtrans"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040693">
        <property type="entry name" value="UGGT_TRXL_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040694">
        <property type="entry name" value="UGGT_TRXL_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040692">
        <property type="entry name" value="UGGT_TRXL_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040525">
        <property type="entry name" value="UGGT_TRXL_4"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11226">
        <property type="entry name" value="UDP-GLUCOSE GLYCOPROTEIN:GLUCOSYLTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11226:SF3">
        <property type="entry name" value="UDP-GLUCOSE:GLYCOPROTEIN GLUCOSYLTRANSFERASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18404">
        <property type="entry name" value="Glyco_transf_24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18400">
        <property type="entry name" value="Thioredoxin_12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18401">
        <property type="entry name" value="Thioredoxin_13"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18402">
        <property type="entry name" value="Thioredoxin_14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18403">
        <property type="entry name" value="Thioredoxin_15"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06427">
        <property type="entry name" value="UDP-g_GGTase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="7" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 8" id="KW-0732">Signal</keyword>
    <keyword evidence="4" id="KW-0808">Transferase</keyword>
    <feature type="signal peptide" evidence="8">
        <location>
            <begin position="1"/>
            <end position="17"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5044604588" description="UDP-glucose glycoprotein glucosyltransferase 1" evidence="8">
        <location>
            <begin position="18"/>
            <end position="1281"/>
        </location>
    </feature>
    <feature type="domain" description="UGGT thioredoxin-like" evidence="9">
        <location>
            <begin position="34"/>
            <end position="215"/>
        </location>
    </feature>
    <feature type="domain" description="UGGT thioredoxin-like" evidence="10">
        <location>
            <begin position="289"/>
            <end position="420"/>
        </location>
    </feature>
    <feature type="domain" description="UGGT thioredoxin-like" evidence="11">
        <location>
            <begin position="428"/>
            <end position="676"/>
        </location>
    </feature>
    <feature type="domain" description="UDP-glucose:glycoprotein glucosyltransferase thioredoxin-like" evidence="12">
        <location>
            <begin position="706"/>
            <end position="932"/>
        </location>
    </feature>
    <feature type="domain" description="Glucosyltransferase 24 catalytic" evidence="13">
        <location>
            <begin position="1234"/>
            <end position="1278"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004922"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00006351"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF18400"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="Pfam" id="PF18401"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF18402"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF18403"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="Pfam" id="PF18404"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029788.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="16">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1281" mass="144856" checksum="0576B32C7FB07F3D" modified="2019-05-08" version="1">MGIFIALLGIWFSSSLAKADSKAVTTSLTTKWLSTPLLLEASEFLAEEGQEKFWTFVEASQDIRSSDHGGTNYSSYHAMLKAACQNLSPLQQNLLKFSLSLRSYSATIQAFQQIAADEPPPVGCNSFFAVHGKKTCEFDTLEILLQTASERLKPFLFKGDHRFPLSNPESPVVILYAEIGTEEFSKFHNLLVSKASAGDITYILRHYVANPSKEKVYLSGYGVELAIKSTEYKAKDDTQVKGMEVNATMIGENDPIDEVQGFLFGKLRQLHPDLKEELKELRKHLIESTNEMAPLKVWQLQDLSFQTAARILAAPTVDALMVMKDLSQNFPTKARAITKTVVSSELRTEIEENQKYFKGTLGLQSGDSAFFINGLHIDLDTQDIFSLFDVLRNEARVMEGLHSLGIEGLSLHNVLKLNIQPSDSDYAVDIRSPAISWINNLEVDSRYNSWPSSVQELLRPTFPGVIRQIRKNFHNFVLIVDPSHESTAELLNVAEMFFSNHIPLRIGLVFVVNDSEDVDGLQDAGVALLRAFNYVAQEVDNNYAFQTLISIYNKVKTGDQLQVEHVVSVLEKQYPYVEMNSILGIDSVYDQNRKEGRGYYEQTGVGPLPIVLFNGMPFQKEQLDPDDLETITMQKILETTSLFQRAVYLGELSSDQDVVEYIMNQPNVVPRINSRILMSDREYLDLTATNNFFVDDYARFTFLNSKDKTAAVANSMTYLTKKGMSSKEIYDDSFVRPVTFWIVGDFDKPSGRQLLYDAIKHQKSSNHVRISMINNPSEDPTSKNTVITRAIWAALQTQTSNNAKNFITKMAKEETAKALEAGADITEFAVGGMDVNTFKEAFESPKVDFILSHAMYCRDVLKLQKGQRAVISNGRIIGPLEDNELFNQDDFHLLENIILKTSGQKIKSHIQQLGVEEDLASDLVMKVDALLSAQPKGEARIEYQFFEDRYSAVKLRPKEGETYFDVVAIVDPATRDAQRLAPLLMVLNQLINMNLRVFMNCQSKLSDMPLKSFYRYVLEPEISFTADDNFAPGPIAKFLDMPQTPLFTLNLNTPESWMVESVRTPYDLDNIYLEEVDNRVAAEYELEYLLLEGHCYDITTGQPPRGLQFTLGTSANPVIVDTIVMANLGYFQLKANPGAWVLRLRKGRSEDIYRIYSHDGTDSPPDANEVTVVLNNFKSKIIKVKVQKKLDMMNEDLLSDGTSENESGFWESLKWGFTGGQKNDEMKQDKDDVINIFSVASGHLYERFLRIMMLSVLKHTKTPVKFWFLKNYLSPMFKVCY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPD5</accession>
    <name>A0A452IPD5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Large ribosomal subunit protein uL15</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="5">60S ribosomal protein L27a</fullName>
        </alternativeName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1 6">Belongs to the universal ribosomal protein uL15 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPD5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033724.1">
        <property type="protein sequence ID" value="ENSGAGP00000029703.1"/>
        <property type="gene ID" value="ENSGAGG00000021440.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0022625">
        <property type="term" value="C:cytosolic large ribosomal subunit"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003735">
        <property type="term" value="F:structural constituent of ribosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006412">
        <property type="term" value="P:translation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.100.10.10:FF:000024">
        <property type="entry name" value="RPL27A isoform 10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.100.10.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021131">
        <property type="entry name" value="Ribosomal_uL15/eL18"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036227">
        <property type="entry name" value="Ribosomal_uL15/eL18_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001196">
        <property type="entry name" value="Ribosomal_uL15_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11721">
        <property type="entry name" value="60S RIBOSOMAL PROTEIN L27A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11721:SF3">
        <property type="entry name" value="LARGE RIBOSOMAL SUBUNIT PROTEIN UL15"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00828">
        <property type="entry name" value="Ribosomal_L27A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52080">
        <property type="entry name" value="Ribosomal proteins L15p and L18e"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00475">
        <property type="entry name" value="RIBOSOMAL_L15"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 6" id="KW-0687">Ribonucleoprotein</keyword>
    <keyword evidence="2 6" id="KW-0689">Ribosomal protein</keyword>
    <feature type="domain" description="Large ribosomal subunit protein uL15/eL18" evidence="7">
        <location>
            <begin position="15"/>
            <end position="88"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007320"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022980"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023274"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00035200"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00035527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="RuleBase" id="RU003888"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF00828"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029703.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="91" mass="10279" checksum="0A1874DB99A84E4B" modified="2019-05-08" version="1">MRHYHLKKNQNFCPTVNLDKLWTLVSEQTRLHYAKNQAGLAPVIDVVRAGYYKVLGKGKLPKQPVIVKAKFFSRKAEEKIKEVGGACVLVA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPE1</accession>
    <name>A0A452IPE1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">pantothenate kinase</fullName>
            <ecNumber evidence="5">2.7.1.33</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>(R)-pantothenate + ATP = (R)-4'-phosphopantothenate + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:16373"/>
            <dbReference type="ChEBI" id="CHEBI:10986"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29032"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.1.33"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="3">Cofactor biosynthesis; coenzyme A biosynthesis; CoA from (R)-pantothenate: step 1/5.</text>
    </comment>
    <comment type="subunit">
        <text evidence="4">Homodimer.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="13">Belongs to the type II pantothenate kinase family.</text>
    </comment>
    <dbReference type="EC" id="2.7.1.33" evidence="5"/>
    <dbReference type="AlphaFoldDB" id="A0A452IPE1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033729.1">
        <property type="protein sequence ID" value="ENSGAGP00000029708.1"/>
        <property type="gene ID" value="ENSGAGG00000021410.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004594">
        <property type="term" value="F:pantothenate kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015937">
        <property type="term" value="P:coenzyme A biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd24135">
        <property type="entry name" value="ASKHA_NBD_PanK-II_Pank1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.420.40:FF:000244">
        <property type="entry name" value="pantothenate kinase 1-like isoform X3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.420.510:FF:000001">
        <property type="entry name" value="pantothenate kinase 2, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.420.40:FF:000392">
        <property type="entry name" value="pantothenate kinase 3-like isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.420.40">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.420.510">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043129">
        <property type="entry name" value="ATPase_NBD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004567">
        <property type="entry name" value="Type_II_PanK"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00555">
        <property type="entry name" value="panK_eukar"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12280">
        <property type="entry name" value="PANTOTHENATE KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12280:SF23">
        <property type="entry name" value="PANTOTHENATE KINASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03630">
        <property type="entry name" value="Fumble"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53067">
        <property type="entry name" value="Actin-like ATPase domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="11" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="12" id="KW-0173">Coenzyme A biosynthesis</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="10" id="KW-0418">Kinase</keyword>
    <keyword evidence="14" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0808">Transferase</keyword>
    <keyword evidence="14" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="14" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="12"/>
            <end position="32"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001206"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005225"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00011738"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012102"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022993"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00060870"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029708.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="391" mass="43638" checksum="9E46E50BAEB46EEF" modified="2019-05-08" version="1">RPWGSSPGNCSCRIFIAPLSFFFPPFLAFPWFGMDIGGTLVKLVYFEPKDITAEEEQEEVENLKSIRKYLTSNTAYGKTGIRDVHLELKNLTMCGRKGNLHFIRFPTSAMHRFIQMGSEKNFSSLHTTLCATGGGAYKFEEDFRTIADLQLHKLDELDCLIQGLLYVDSVGYNGQPECYYFENPTDPEQCQKKPYCLDNPYPMLLVNIGSGVSILAVYSKDNYKRVTGTSLGGGTFLGLCCLLTGCETFEEALEMAAKGDSTNVDKLVKDIYGGDYERFGLQGSAVASSFGHMMSKEKRDSISKEDLARATLVTITNNIGSIARMCALNENIDRVVFVGNFLRINMISMKVLAYAMDYWSKGQLKALFLEHEGYFGAVGALLELLKMPENQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPE6</accession>
    <name>A0A452IPE6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Acrosin-binding protein</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="6">Acrosin-binding protein, 60 kDa form</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="7">Proacrosin-binding protein sp32</fullName>
        </alternativeName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="8">Acrosomal protein that maintains proacrosin (pro-ACR) as an enzymatically inactive zymogen in the acrosome. Involved also in the acrosome formation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasmic vesicle</location>
            <location evidence="1">Secretory vesicle</location>
            <location evidence="1">Acrosome</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPE6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033736.1">
        <property type="protein sequence ID" value="ENSGAGP00000029713.1"/>
        <property type="gene ID" value="ENSGAGG00000021441.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001669">
        <property type="term" value="C:acrosomal vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009865">
        <property type="entry name" value="Proacrosin-bd"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21362">
        <property type="entry name" value="ACROSIN-BINDING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21362:SF1">
        <property type="entry name" value="ACROSIN-BINDING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07222">
        <property type="entry name" value="PBP_sp32"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004218"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00018940"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00032734"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00033453"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00045517"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029713.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="90" mass="10196" checksum="3A0CBAD63C90C40B" modified="2025-10-08" version="2">MPKPRLLLPMVQPVWSMSSLVPGSPLSDSEYQVFFSALMPSWKAGVACQIRRTQGCHDPKVIRLDQFENHGWIPEAALFPTSLTFLKWVP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPH2</accession>
    <name>A0A452IPH2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Protein Wnt</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="11">Ligand for members of the frizzled family of seven transmembrane receptors.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 11">Secreted</location>
            <location evidence="1 11">Extracellular space</location>
            <location evidence="1 11">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 11">Belongs to the Wnt family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPH2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033768.1">
        <property type="protein sequence ID" value="ENSGAGP00000029739.1"/>
        <property type="gene ID" value="ENSGAGG00000021458.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005125">
        <property type="term" value="F:cytokine activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005109">
        <property type="term" value="F:frizzled binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060070">
        <property type="term" value="P:canonical Wnt signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045165">
        <property type="term" value="P:cell fate commitment"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030182">
        <property type="term" value="P:neuron differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.2460.20:FF:000001">
        <property type="entry name" value="Wnt homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.2460.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005817">
        <property type="entry name" value="Wnt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009142">
        <property type="entry name" value="Wnt4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043158">
        <property type="entry name" value="Wnt_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12027:SF101">
        <property type="entry name" value="PROTEIN WNT-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12027">
        <property type="entry name" value="WNT RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00110">
        <property type="entry name" value="wnt"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01844">
        <property type="entry name" value="WNT4PROTEIN"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01349">
        <property type="entry name" value="WNTPROTEIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00097">
        <property type="entry name" value="WNT1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 11" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="9" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
    <keyword evidence="6 11" id="KW-0879">Wnt signaling pathway</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005683"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022687"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="RuleBase" id="RU003500"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029739.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="256" mass="28513" checksum="9EC2297F4987B481" modified="2019-05-08" version="1">KRARKQQPELCLFAVAMRVAGQKRGISSLEEYLAKQPSLRGTLQDSSACEGLKGLAEEQVRICQRQVEAMDSVKHGAELAIEECQHQFHNRRWNCSTLQGLQVFGTRESAFIHAISSAGVAFAVTQACSRGELEQCGCDRKIRGGATEVHPKRVGSRKLLVPKSSRFKPYTAHDLVYLVASPDFCDRDPHSGVFGTSGRQCNRTSHAMDGCELLCCGRGFRTAQAERVERCSCKFHWCCSVKCKQCHHLVEVHTCR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPH5</accession>
    <name>A0A452IPH5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Glycosyltransferase 2-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IPH5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000034071.1">
        <property type="protein sequence ID" value="ENSGAGP00000030002.1"/>
        <property type="gene ID" value="ENSGAGG00000021650.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008376">
        <property type="term" value="F:acetylgalactosaminyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019276">
        <property type="term" value="P:UDP-N-acetylgalactosamine metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006047">
        <property type="term" value="P:UDP-N-acetylglucosamine metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00761">
        <property type="entry name" value="Glyco_tranf_GTA_type"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.550.10">
        <property type="entry name" value="Spore Coat Polysaccharide Biosynthesis Protein SpsA, Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001173">
        <property type="entry name" value="Glyco_trans_2-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029044">
        <property type="entry name" value="Nucleotide-diphossugar_trans"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15046:SF2">
        <property type="entry name" value="BETA-1,4 N-ACETYLGALACTOSAMINYLTRANSFERASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15046">
        <property type="entry name" value="GLYCO_TRANS_2-LIKE DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00535">
        <property type="entry name" value="Glycos_transf_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53448">
        <property type="entry name" value="Nucleotide-diphospho-sugar transferases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="1">
        <location>
            <begin position="1"/>
            <end position="19"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5018993263" description="Glycosyltransferase 2-like domain-containing protein" evidence="1">
        <location>
            <begin position="20"/>
            <end position="415"/>
        </location>
    </feature>
    <feature type="domain" description="Glycosyltransferase 2-like" evidence="2">
        <location>
            <begin position="175"/>
            <end position="286"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF00535"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000030002.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="415" mass="47336" checksum="61DD7845829553C9" modified="2019-05-08" version="1">RYFSTSILLWALTLSYIQPNFLTKEATFTCQGRRGSYSRPKPNVPLSYPIQGVEVMPLHTIMIPGIHQGPSVILRASLGTLNTLADVSDDVVRGRGRGKKELLILTSDVELLNFILEHVTYTSVVYQLSAVDMMSFKSRDHVVQFPVIIRQPSLPKLIDPGADRKISSLVTITTKTFLRYHKLRFLIKSIRQYYPDIKIIVADDSEVPEKIIEENIEHYIMPFGKGWFAGRNLAISQVTTKYYLWVDDDYLFTENTKIEKLLDVLERTNLDMVGGSVKGDTYSFQLLYEQSDDGYCLHLRYGSFHKLDGFPNCVVTSGVVNFYLAHTDKSRYVSYDPLLQRVAHSEFFVDGLGILLVGSCSDVSVGHQDHNPASDPNLAKIEDQYKTFRDNTDAQVQFKLALHYFKNRLKCYSTR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPH9</accession>
    <name>A0A452IPH9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">3',5'-cyclic-AMP phosphodiesterase</fullName>
            <ecNumber evidence="3">3.1.4.53</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="7">
            <text>3',5'-cyclic AMP + H2O = AMP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:25277"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:58165"/>
            <dbReference type="ChEBI" id="CHEBI:456215"/>
            <dbReference type="EC" id="3.1.4.53"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="7">
            <dbReference type="Rhea" id="RHEA:25278"/>
        </physiologicalReaction>
    </comment>
    <comment type="pathway">
        <text evidence="1">Purine metabolism; 3',5'-cyclic AMP degradation; AMP from 3',5'-cyclic AMP: step 1/1.</text>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the cyclic nucleotide phosphodiesterase family. PDE4 subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.1.4.53" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452IPH9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033943.1">
        <property type="protein sequence ID" value="ENSGAGP00000029891.1"/>
        <property type="gene ID" value="ENSGAGG00000021546.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00762">
        <property type="reaction ID" value="UER00747"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004115">
        <property type="term" value="F:3',5'-cyclic-AMP phosphodiesterase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006198">
        <property type="term" value="P:cAMP catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00077">
        <property type="entry name" value="HDc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1300.10">
        <property type="entry name" value="3'5'-cyclic nucleotide phosphodiesterase, catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003607">
        <property type="entry name" value="HD/PDEase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040844">
        <property type="entry name" value="PDE4_UCR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023088">
        <property type="entry name" value="PDEase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002073">
        <property type="entry name" value="PDEase_catalytic_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036971">
        <property type="entry name" value="PDEase_catalytic_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023174">
        <property type="entry name" value="PDEase_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11347">
        <property type="entry name" value="CYCLIC NUCLEOTIDE PHOSPHODIESTERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18100">
        <property type="entry name" value="PDE4_UCR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00233">
        <property type="entry name" value="PDEase_I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00387">
        <property type="entry name" value="PDIESTERASE1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109604">
        <property type="entry name" value="HD-domain/PDEase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00126">
        <property type="entry name" value="PDEASE_I_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51845">
        <property type="entry name" value="PDEASE_I_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0114">cAMP</keyword>
    <keyword evidence="5" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="4 9" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="PDEase" evidence="10">
        <location>
            <begin position="97"/>
            <end position="328"/>
        </location>
    </feature>
    <feature type="active site" description="Proton donor" evidence="8">
        <location>
            <position position="173"/>
        </location>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="177"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="213"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="214"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="214"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <label>2</label>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004703"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009517"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012276"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023149"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00033681"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PIRSR" id="PIRSR623088-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PIRSR" id="PIRSR623088-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS51845"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029891.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="328" mass="37511" checksum="A23F9A00F1D483C4" modified="2019-05-08" version="1">MPEANYLLSVSWGYIKFKRMLNRELTHLSEMSRSGNQVSEYISNTFLDKQNDVEIPSPTQKDREKKKKQQLMTQISGVKKLMHSSSLNNTSISRFGVKTEKEDHLAKELEDLNKWGLNIFNVARYSHSRPLTCIMYAIFQERDLLKTFKISSDTFVTYMMTLEDHYHSDVAYHNSLHAADVAQSTHVLLSTPALDAVFTDLEILAAIFAAAIHDVDHPGVSNQFLINTNSELALMYNDESVLENHHLAVGFKLLQEEHCDIFQNLSKKQRQTLRKMVIDMVLATDMSKHMSLLADLKTMVETKKVTSSGVLLLDNYTDRIQVCDCNFP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPI4</accession>
    <name>A0A452IPI4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="19">UDP-glucose glycoprotein glucosyltransferase 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Recognizes glycoproteins with minor folding defects. Reglucosylates single N-glycans near the misfolded part of the protein, thus providing quality control for protein folding in the endoplasmic reticulum. Reglucosylated proteins are recognized by calreticulin for recycling to the endoplasmic reticulum and refolding or degradation.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="10">
            <text>N(4)-(alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;6)]-alpha-D-Man-(1-&gt;6)]-beta-D-Man-(1-&gt;4)-beta-D-GlcNAc-(1-&gt;4)-beta-D-GlcNAc)-L-asparaginyl-[protein] (N-glucan mannose isomer 9A1,2,3B1,2,3) + UDP-alpha-D-glucose = N(4)-(alpha-D-Glc-(1-&gt;3)-alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;6)]-alpha-D-Man-(1-&gt;6)]-beta-D-Man-(1-&gt;4)-beta-D-GlcNAc-(1-&gt;4)-beta-D-GlcNAc)-L-asparaginyl-[protein] + UDP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:61304"/>
            <dbReference type="Rhea" id="RHEA-COMP:14356"/>
            <dbReference type="Rhea" id="RHEA-COMP:14357"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:58223"/>
            <dbReference type="ChEBI" id="CHEBI:58885"/>
            <dbReference type="ChEBI" id="CHEBI:59080"/>
            <dbReference type="ChEBI" id="CHEBI:139493"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </cofactor>
    </comment>
    <comment type="pathway">
        <text evidence="3">Protein modification; protein glycosylation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Endoplasmic reticulum lumen</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the glycosyltransferase 8 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000033887.1">
        <property type="protein sequence ID" value="ENSGAGP00000029844.1"/>
        <property type="gene ID" value="ENSGAGG00000021377.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00378"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005788">
        <property type="term" value="C:endoplasmic reticulum lumen"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003980">
        <property type="term" value="F:UDP-glucose:glycoprotein glucosyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051082">
        <property type="term" value="F:unfolded protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036503">
        <property type="term" value="P:ERAD pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0018279">
        <property type="term" value="P:protein N-linked glycosylation via asparagine"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.550.10">
        <property type="entry name" value="Spore Coat Polysaccharide Biosynthesis Protein SpsA, Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040497">
        <property type="entry name" value="Glyco_transf_24"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029044">
        <property type="entry name" value="Nucleotide-diphossugar_trans"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009448">
        <property type="entry name" value="UDP-g_GGtrans"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040693">
        <property type="entry name" value="UGGT_TRXL_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040694">
        <property type="entry name" value="UGGT_TRXL_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040692">
        <property type="entry name" value="UGGT_TRXL_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040525">
        <property type="entry name" value="UGGT_TRXL_4"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11226">
        <property type="entry name" value="UDP-GLUCOSE GLYCOPROTEIN:GLUCOSYLTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11226:SF3">
        <property type="entry name" value="UDP-GLUCOSE:GLYCOPROTEIN GLUCOSYLTRANSFERASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18404">
        <property type="entry name" value="Glyco_transf_24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18400">
        <property type="entry name" value="Thioredoxin_12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18401">
        <property type="entry name" value="Thioredoxin_13"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18402">
        <property type="entry name" value="Thioredoxin_14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18403">
        <property type="entry name" value="Thioredoxin_15"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06427">
        <property type="entry name" value="UDP-g_GGTase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53448">
        <property type="entry name" value="Nucleotide-diphospho-sugar transferases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="8" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="5" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="UGGT thioredoxin-like" evidence="12">
        <location>
            <begin position="53"/>
            <end position="234"/>
        </location>
    </feature>
    <feature type="domain" description="UGGT thioredoxin-like" evidence="13">
        <location>
            <begin position="308"/>
            <end position="439"/>
        </location>
    </feature>
    <feature type="domain" description="UGGT thioredoxin-like" evidence="14">
        <location>
            <begin position="447"/>
            <end position="695"/>
        </location>
    </feature>
    <feature type="domain" description="UDP-glucose:glycoprotein glucosyltransferase thioredoxin-like" evidence="15">
        <location>
            <begin position="725"/>
            <end position="951"/>
        </location>
    </feature>
    <feature type="domain" description="Glucosyltransferase 24 catalytic" evidence="16">
        <location>
            <begin position="1246"/>
            <end position="1353"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1381"/>
            <end position="1408"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001913"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004922"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00006351"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00045874"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00048456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF18400"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="Pfam" id="PF18401"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="Pfam" id="PF18402"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="Pfam" id="PF18403"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF18404"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029844.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="19">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1408" mass="159738" checksum="92512F03824EB653" modified="2019-05-08" version="1">MVNKWGGKEDIMASCIFYKMGIFIALLGIWFSSSLAKADSKAVTTSLTTKWLSTPLLLEASEFLAEEGQEKFWTFVEASQDIRSSDHGGTNYSSYHAMLKAACQNLSPLQQNLLKFSLSLRSYSATIQAFQQIAADEPPPVGCNSFFAVHGKKTCEFDTLEILLQTASERLKPFLFKGDHRFPLSNPESPVVILYAEIGTEEFSKFHNLLVSKASAGDITYILRHYVANPSKEKVYLSGYGVELAIKSTEYKAKDDTQVKGMEVNATMIGENDPIDEVQGFLFGKLRQLHPDLKEELKELRKHLIESTNEMAPLKVWQLQDLSFQTAARILAAPTVDALMVMKDLSQNFPTKARAITKTVVSSELRTEIEENQKYFKGTLGLQSGDSAFFINGLHIDLDTQDIFSLFDVLRNEARVMEGLHSLGIEGLSLHNVLKLNIQPSDSDYAVDIRSPAISWINNLEVDSRYNSWPSSVQELLRPTFPGVIRQIRKNFHNFVLIVDPSHESTAELLNVAEMFFSNHIPLRIGLVFVVNDSEDVDGLQDAGVALLRAFNYVAQEVDNNYAFQTLISIYNKVKTGDQLQVEHVVSVLEKQYPYVEMNSILGIDSVYDQNRKEGRGYYEQTGVGPLPIVLFNGMPFQKEQLDPDDLETITMQKILETTSLFQRAVYLGELSSDQDVVEYIMNQPNVVPRINSRILMSDREYLDLTATNNFFVDDYARFTFLNSKDKTAAVANSMTYLTKKGMSSKEIYDDSFVRPVTFWIVGDFDKPSGRQLLYDAIKHQKSSNHVRISMINNPSEDPTSKNTVITRAIWAALQTQTSNNAKNFITKMAKEETAKALEAGADITEFAVGGMDVNTFKEAFESPKVDFILSHAMYCRDVLKLQKGQRAVISNGRIIGPLEDNELFNQDDFHLLENIILKTSGQKIKSHIQQLGVEEDLASDLVMKVDALLSAQPKGEARIEYQFFEDRYSAVKLRPKEGETYFDVVAIVDPATRDAQRLAPLLMVLNQLINMNLRVFMNCQSKLSDMPLKSFYRYVLEPEISFTADDNFAPGPIAKFLDMPQTPLFTLNLNTPESWMVESVRTPYDLDNIYLEEVDNRVAAEYELEYLLLEGHCYDITTGQPPRGLQFTLGTSANPVIVDTIVMANLGYFQLKANPGAWVLRLRKGRSEDIYRIYSHDGTDSPPDANEVTVVLNNFKSKIIKVKVSLKTVLQEYKYSNEDYIFPDFRGFTGGQKNDEMKQDKDDVINIFSVASGHLYERFLRIMMLSVLKHTKTPVKFWFLKNYLSPMFKEFIPYMAEKYNFQYELVQYKWPRWLHQQTEKQRIIWGYKILFLDVLFPLVVDKFLFVDADQVSYLMALGSLAGITRSAYIAEEGSALQQMWQGSPCPSLPTDWRGTEPSRQGASGRAR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPI7</accession>
    <name>A0A452IPI7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Pre-mRNA-splicing factor SYF1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the crooked-neck family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IPI7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033853.1">
        <property type="protein sequence ID" value="ENSGAGP00000029814.1"/>
        <property type="gene ID" value="ENSGAGG00000021531.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071014">
        <property type="term" value="C:post-mRNA release spliceosomal complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000974">
        <property type="term" value="C:Prp19 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071007">
        <property type="term" value="C:U2-type catalytic step 2 spliceosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000349">
        <property type="term" value="P:generation of catalytic spliceosome for first transesterification step"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000023">
        <property type="entry name" value="Pre-mRNA-splicing factor SYF1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000137">
        <property type="entry name" value="Pre-mRNA-splicing factor syf1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000220">
        <property type="entry name" value="Pre-mRNA-splicing factor SYF1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000411">
        <property type="entry name" value="pre-mRNA-splicing factor SYF1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.430:FF:000006">
        <property type="entry name" value="Pre-mRNA-splicing factor SYF1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:001071">
        <property type="entry name" value="pre-mRNA-splicing factor SYF1-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.10">
        <property type="entry name" value="Tetratricopeptide repeat domain"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003107">
        <property type="entry name" value="HAT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055433">
        <property type="entry name" value="HAT_Syf1-like_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056350">
        <property type="entry name" value="HAT_Syf1_central"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055430">
        <property type="entry name" value="HAT_Syf1_CNRKL1_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045075">
        <property type="entry name" value="Syf1-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011990">
        <property type="entry name" value="TPR-like_helical_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019734">
        <property type="entry name" value="TPR_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11246">
        <property type="entry name" value="PRE-MRNA SPLICING FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11246:SF5">
        <property type="entry name" value="PRE-MRNA-SPLICING FACTOR SYF1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23231">
        <property type="entry name" value="HAT_Syf1_CNRKL1_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23233">
        <property type="entry name" value="HAT_Syf1_CNRKL1_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23220">
        <property type="entry name" value="HAT_Syf1_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00386">
        <property type="entry name" value="HAT"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="SMART" id="SM00028">
        <property type="entry name" value="TPR"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48452">
        <property type="entry name" value="TPR-like"/>
        <property type="match status" value="5"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0507">mRNA processing</keyword>
    <keyword evidence="6" id="KW-0508">mRNA splicing</keyword>
    <keyword evidence="7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0747">Spliceosome</keyword>
    <feature type="domain" description="Pre-mRNA-splicing factor Syf1-like N-terminal HAT-repeats" evidence="13">
        <location>
            <begin position="23"/>
            <end position="181"/>
        </location>
    </feature>
    <feature type="domain" description="Pre-mRNA-splicing factor SYF1 central HAT repeats" evidence="11">
        <location>
            <begin position="184"/>
            <end position="388"/>
        </location>
    </feature>
    <feature type="domain" description="Pre-mRNA-splicing factor Syf1/CRNKL1-like C-terminal HAT-repeats" evidence="12">
        <location>
            <begin position="390"/>
            <end position="777"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="808"/>
            <end position="858"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="769"/>
            <end position="796"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="10">
        <location>
            <begin position="819"/>
            <end position="841"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008644"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022664"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022728"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023187"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00039472"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF23220"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF23231"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="Pfam" id="PF23233"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029814.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="858" mass="100745" checksum="90602E0ECD3C2368" modified="2019-05-08" version="1">MPALEETPVAAGEGKPDLFFEEEDLQYEEEILRNPFSVKCWIRYIEFKQNAPRHVLNLIYERALKELPGSYKLWYNYLKQRRKQVKNRCVTDPSYEEVNNCHERALVFMHKMPRIWLDYCQFLMDQCRITRTRRTFDRALRALPITQHHRIWPLYLKFVRLYPLPETAVRVYRRYLKLSPENAEEYIEYLRSIDRLDEAALRLATVVNDERFISKEGKSNYQLWHELCDLISQNPDKVKSLNVGAIIRGGLTRFTDQLGKLWCSLADYYIRSGHFEKARDVYEEAIQTVMTVRDFTQVFDSYAQFEESMIAAKMETTSELGQEEEDDVDLELRLARFEQLITRRPLLLNSVLLRQNPHNVHEWHKRVKLFEGKPREIINTYTEAVQTVDPFKATGKTHTLWVSFAKFYEDNGQIEDARTIFEKATKVNFKQVDDLACVWCEYGEMELRHENYDQALRILRKATAIPAKKAEYFDASEPVQNRVYKSLKVWSMLADLEESLGTFKSTKAVYDRILDLRIATPQIIINYALFLEEHTYFEESFKAYERGISLFRWPNVYDIWNTYLTKFIDRYGGKKLERARDLFEQALDGCPQKYAKTIYLLYAKLEEEFGLARHAMAVYERATRAVQPSEQYEMYNIFIKRAAEIYGVTHTRSIYEKAIEVLSDEHAREMCQRFADMECKLGEIDRARAIYSYCSQICDPRTTGSFWQTWKDFEIRHGNEDTIREMLRIKRSVQATYNTQVNFMASQMLKVSSNATGTVSDLAPGQSGMDDMKLLEQRAEQLAAEAERDKPRAKEKILFVRSDASRSELAALSRQANPDEIDIEEEEEEGESEENEPDEVQLEQQSVPSAVFGGLKTD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPI9</accession>
    <name>A0A452IPI9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="11">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="domain">
        <text evidence="6">The CKK domain binds microtubules.</text>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the CAMSAP1 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000033892.1">
        <property type="protein sequence ID" value="ENSGAGP00000029849.1"/>
        <property type="gene ID" value="ENSGAGG00000021541.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036449">
        <property type="term" value="C:microtubule minus-end"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051011">
        <property type="term" value="F:microtubule minus-end binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030507">
        <property type="term" value="F:spectrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031122">
        <property type="term" value="P:cytoplasmic microtubule organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007026">
        <property type="term" value="P:negative regulation of microtubule depolymerization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031175">
        <property type="term" value="P:neuron projection development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.20.360:FF:000001">
        <property type="entry name" value="Calmodulin-regulated spectrin-associated protein 3 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.20.360">
        <property type="entry name" value="CKK domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032940">
        <property type="entry name" value="CAMSAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031372">
        <property type="entry name" value="CAMSAP_CC1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022613">
        <property type="entry name" value="CH_CAMSAP_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001715">
        <property type="entry name" value="CH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036872">
        <property type="entry name" value="CH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014797">
        <property type="entry name" value="CKK_CAMSAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038209">
        <property type="entry name" value="CKK_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011033">
        <property type="entry name" value="PRC_barrel-like_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21595:SF2">
        <property type="entry name" value="CALMODULIN-REGULATED SPECTRIN-ASSOCIATED PROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21595">
        <property type="entry name" value="PATRONIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17095">
        <property type="entry name" value="CAMSAP_CC1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11971">
        <property type="entry name" value="CAMSAP_CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08683">
        <property type="entry name" value="CAMSAP_CKK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01051">
        <property type="entry name" value="CAMSAP_CKK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47576">
        <property type="entry name" value="Calponin-homology domain, CH-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50346">
        <property type="entry name" value="PRC-barrel domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50021">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51508">
        <property type="entry name" value="CKK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="5" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="3 6" id="KW-0493">Microtubule</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="8">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019022003" evidence="8">
        <location>
            <begin position="23"/>
            <end position="1228"/>
        </location>
    </feature>
    <feature type="domain" description="Calponin-homology (CH)" evidence="9">
        <location>
            <begin position="75"/>
            <end position="189"/>
        </location>
    </feature>
    <feature type="domain" description="CKK" evidence="10">
        <location>
            <begin position="1089"/>
            <end position="1223"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="54"/>
            <end position="81"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="399"/>
            <end position="547"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="571"/>
            <end position="627"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="671"/>
            <end position="966"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1035"/>
            <end position="1095"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="455"/>
            <end position="466"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="467"/>
            <end position="486"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="591"/>
            <end position="605"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="710"/>
            <end position="732"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="7">
        <location>
            <begin position="733"/>
            <end position="749"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="784"/>
            <end position="797"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="7">
        <location>
            <begin position="866"/>
            <end position="878"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="894"/>
            <end position="966"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00841"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS51508"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029849.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1228" mass="133588" checksum="A8E3064042B69D88" modified="2019-05-08" version="1">MGAHLAVIDSLMIAFAVESTHSLSAPPGTDLGAGSWEQKLLCWVDTVNRKLQESTEREGSQQLVPGTEGQSQSQSPASGPKIRYRKDKVLPKQTPCFTPVMGMKDLANGGAIAATIHYYCPDVVRLEDVCLKETMSVADSLYNLQLIQEFCIKYLGGCCPLALEDLLYVPPVLKINIGVFLAELFLCFEVLKPDFVRPKELRGLKDPPAVSDSLTPSSGNSNSGSLVFSFLHPLLPGGQPQSPLRGSLGSMHHSTSMSHVEGGFGKSWSKKQLSHPLSQAVSFSIPFGLDSDVDIVMGNPVGMLRSISSDSLAPVPTRLARSPGPLPEDAAQRLAKDEAPNGPVAPHRTVHKVALSAKGVVEREGPPVENGLADGYPDLPTIEEALQIIHSSERLHPEGAADGFYLHSPEPAKPTQAPEPTPALAVYRFHSGPPNGRAEAPTPPGNGSLDSDAEDPARLPGTKDDTSSGLSSLSSQPDSAGSSGAGVRMTSFAERKKKLAPPEGKTLQKSTGESSEAGAVPPEDSPGRSPALSSEMSQLGARLEEKRRAIEAQKKRIEAIFAKHRQRLGKSAFLQLKRRDGGEEEEEEEGEAKLSLEERLARLEAEEGGSLQVQAEEEGPGQGCPEKQVTFSSEIKGAPLDENLGDYNRAVAKLNTALSSLQLDMHRLSQQQQRLLQEKKPSQAWVIPAPKGPAPRASREFVPPRSVELSSSPSPSPSRXXXXXXXXXXPAAPRSPQPAPKKAAPTPPKSPKHARPVELKLPPLTRVLTPPHNVDTLPHLRKFSPSQVPMQTRSSIHFSEEVAGPEQPLESPELETQGNLRPVASVVPQGGVGSARVSGDGTSDVSSPGERRSSLIEIPLSSLQAEEGDGDDSLEESLTEVMDMEPRAGLGFFFKDDEKPEDEMAQKRASLLERQQRRNEEARQRKQWLEAEKEQKEEAARLQAEERPRPEEEASPRRGDFTRQEYQRRHQLKLMEDLDKVLRQKPTTVRALKKGRPKTVFCDDSALARSPVKGLLGSRLSKVYSQSTLSLSTVANEPGNSLTIKRPSRAASPSGLMSPSRLLGSQSRERDWENASTASSPASVPEYTGPKLYKEPSAKSNKHIIHNALSHCCLAGRVNEPHKNKILEEMEKSKAHHFLILFRDSSCQFRALYTPAGETEELTRLTGYGPRTISQAMIEGIYKYNSDRKRFTQIPAKTMSMNVDAITIQGHLWQTKKPGTPKKPGTPK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1A1</accession>
    <name>A0A452J1A1_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="14">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Golgi apparatus membrane</location>
            <topology evidence="1">Single-pass type II membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the glycosyltransferase 29 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1A1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038679.1">
        <property type="protein sequence ID" value="ENSGAGP00000034156.1"/>
        <property type="gene ID" value="ENSGAGG00000024270.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000139">
        <property type="term" value="C:Golgi membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003828">
        <property type="term" value="F:alpha-N-acetylneuraminate alpha-2,8-sialyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006491">
        <property type="term" value="P:N-glycan processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009311">
        <property type="term" value="P:oligosaccharide metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd23987">
        <property type="entry name" value="GT29_ST8SIA2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1480.20:FF:000001">
        <property type="entry name" value="ST8 alpha-N-acetyl-neuraminide alpha-2,8-sialyltransferase 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1480.20">
        <property type="entry name" value="Glycosyl transferase family 29"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001675">
        <property type="entry name" value="Glyco_trans_29"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050943">
        <property type="entry name" value="Glycosyltr_29_Sialyltrsf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038578">
        <property type="entry name" value="GT29-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012163">
        <property type="entry name" value="Sialyl_trans"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11987">
        <property type="entry name" value="ALPHA-2,8-SIALYLTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11987:SF30">
        <property type="entry name" value="ALPHA-2,8-SIALYLTRANSFERASE 8B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00777">
        <property type="entry name" value="Glyco_transf_29"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF005557">
        <property type="entry name" value="Sialyl_trans"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="3" id="KW-0328">Glycosyltransferase</keyword>
    <keyword evidence="8" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="9" id="KW-0472">Membrane</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="13" id="KW-0732">Signal</keyword>
    <keyword evidence="6" id="KW-0735">Signal-anchor</keyword>
    <keyword evidence="4" id="KW-0808">Transferase</keyword>
    <keyword evidence="5" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="13">
        <location>
            <begin position="1"/>
            <end position="28"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019269607" evidence="13">
        <location>
            <begin position="29"/>
            <end position="353"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="135"/>
            <end position="285"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004323"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022676"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PIRSR" id="PIRSR005557-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034156.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="353" mass="40201" checksum="CF2E56B953F03E58" modified="2019-05-08" version="1">MQLPCRSWALALLTLLVGFLVFVDISELEEETGAEVVINGSSSPAVVDRSNESIKHIKPASFRWRHNQTLSLKIRKQILKFLDAEKDISVLKGTLKPGDVIHYIFDRDSTMNVSQNLYELLPRTSPLKGKNFRTCGIVGNSGILLNSGCGREIDAHSFVIRCNLAPVQEYAWDVGMKTDLVTMNPSVIQRAFEDLVNETWREKLLQHLHSLNGSILWIPAFMAKGGKERVEWVNELILKHRINVRTAYPSLRLLHAVRGYWLTNKVHIKRPTTGLLMYTLATRFCNRIYLYGFWPFPRDQNQNPVKYHYYDSLKYGYTSQASPHTMPLEFKALKTLHQQGALKLTVGECGGAT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1A2</accession>
    <name>A0A452J1A2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">Anaphase-promoting complex subunit 4</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J1A2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038677.1">
        <property type="protein sequence ID" value="ENSGAGP00000034154.1"/>
        <property type="gene ID" value="ENSGAGG00000024170.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005680">
        <property type="term" value="C:anaphase-promoting complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034399">
        <property type="term" value="C:nuclear periphery"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031145">
        <property type="term" value="P:anaphase-promoting complex-dependent catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070979">
        <property type="term" value="P:protein K11-linked ubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024789">
        <property type="entry name" value="APC4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056358">
        <property type="entry name" value="APC4_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024790">
        <property type="entry name" value="APC4_long_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13260">
        <property type="entry name" value="ANAPHASE PROMOTING COMPLEX SUBUNIT 4 APC4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13260:SF0">
        <property type="entry name" value="ANAPHASE-PROMOTING COMPLEX SUBUNIT 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12896">
        <property type="entry name" value="ANAPC4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23405">
        <property type="entry name" value="WD40_APC4_C-half"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="2" id="KW-0132">Cell division</keyword>
    <keyword evidence="3" id="KW-0498">Mitosis</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="Anaphase-promoting complex subunit 4 long" evidence="7">
        <location>
            <begin position="99"/>
            <end position="297"/>
        </location>
    </feature>
    <feature type="domain" description="Anaphase-promoting complex subunit 4 C-terminal half WD40" evidence="8">
        <location>
            <begin position="441"/>
            <end position="611"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="614"/>
            <end position="669"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="6">
        <location>
            <begin position="614"/>
            <end position="629"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00016067"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022776"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF12896"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF23405"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034154.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="669" mass="75779" checksum="0B8A8743C735C477" modified="2019-05-08" version="1">MHWMEVTEESSVLTSFYNAEDESNLLLPKLPALPKKLNALVLGGSSGFIEVYAYGMYKIATVTGVAGSCLGLCFSSDLKSLSVITEVQVSPDSAPEITYFQLDTSLLSVYLPEVTRMARKFTHISTLLQYIKLSLTCMCEAWEEILMQMDSRLTKFVQEKNTTTSVQDEFMQLLLWGKASLELQALLMNQLTVKGLKKLGQSIESSYSSIQKLVISHLQSGSEALLYHLSELKGMASWKQKYESLGLDAAGIEGAITAVGSFILKANELLQVIDSSMKNFKAFFRWLYVAMLRMSEDHVLPELNKMTQKDITFVADFLTEHFNEAPDLYNRKGKYFNVERVGQYLKDEDDDLVSPPNTEGNQWFNFLQNSTHLKESPLLFPYYPQKSLHFVKRRMEGIIDQCLQKPADVIGKSVHQAVCMCLYKGSKSEDSTPRLFKLPFMWNDKSSNLHYVLFTMLESSLSKIYILRRHTDVSRSANNGLLSVDFGNFLNNSINESSDTGFYSCVDAHFYDDETITVVLKENAEQEGKERILAQLPLSSVYVDEDQEREFSLNSNKRLDEQSDDIPTRTVFLESQWRVLENMKAQYVAVNGIRKVSCVLSSNLRHVRVFEMDVEDDGEAEEEEEEEETTPIVGDEHEELTLSAVNHNSMCGGSAEADETEEHESSLDS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1A3</accession>
    <name>A0A452J1A3_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="12">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the GPSM family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1A3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038686.1">
        <property type="protein sequence ID" value="ENSGAGP00000034162.1"/>
        <property type="gene ID" value="ENSGAGG00000024269.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000038690.1">
        <property type="protein sequence ID" value="ENSGAGP00000034166.1"/>
        <property type="gene ID" value="ENSGAGG00000024269.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005938">
        <property type="term" value="C:cell cortex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001965">
        <property type="term" value="F:G-protein alpha-subunit binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005092">
        <property type="term" value="F:GDP-dissociation inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000132">
        <property type="term" value="P:establishment of mitotic spindle orientation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000043">
        <property type="entry name" value="G-protein-signaling modulator 2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.10">
        <property type="entry name" value="Tetratricopeptide repeat domain"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003109">
        <property type="entry name" value="GoLoco_motif"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052386">
        <property type="entry name" value="GPSM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011990">
        <property type="entry name" value="TPR-like_helical_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019734">
        <property type="entry name" value="TPR_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45954:SF2">
        <property type="entry name" value="G-PROTEIN-SIGNALING MODULATOR 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45954">
        <property type="entry name" value="LD33695P"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02188">
        <property type="entry name" value="GoLoco"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13374">
        <property type="entry name" value="TPR_10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13424">
        <property type="entry name" value="TPR_12"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13176">
        <property type="entry name" value="TPR_7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00390">
        <property type="entry name" value="GoLoco"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00028">
        <property type="entry name" value="TPR"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48452">
        <property type="entry name" value="TPR-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50877">
        <property type="entry name" value="GOLOCO"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50005">
        <property type="entry name" value="TPR"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="8 10" id="KW-0802">TPR repeat</keyword>
    <feature type="repeat" description="TPR" evidence="10">
        <location>
            <begin position="43"/>
            <end position="76"/>
        </location>
    </feature>
    <feature type="repeat" description="TPR" evidence="10">
        <location>
            <begin position="223"/>
            <end position="256"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="356"/>
            <end position="385"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="424"/>
            <end position="448"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="585"/>
            <end position="604"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="618"/>
            <end position="640"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="626"/>
            <end position="640"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00006600"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022803"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00339"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034162.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="640" mass="71486" checksum="EDC268DB7F5AA179" modified="2019-05-08" version="1">MEASCLELALEGERLCKAGDFKAGAAFFEAAVQVGTEDLKTLSAIYSQLGNAYFYLKEYSKALEYHKHDLTLARTIGDRIGEAKASGNLGNTLKILGQFDEAIVCCQRHLEISQEQGDKVGEARALYNIGNVYHAKGKQLSWSMAQDPGSLPPEVKETLQRASEYYERNLSLVKELGDRAAQGRAYGNLGNTQYLLGNFNEATAFHKERLAIAKEFGDKAAERRAYSNLGNAHIFLGRFDISAEYYKKTLQLSRQLKDQAVEAQACYSLGNTYTLLQDYERAIDYHLKHLLIAQELGDRVGEGRACWSLGNAYVSLGNHDQALRFARKHLEISQEIGDRNGELTARMNVAQLKSALGDEDGDATHHSSSYEAQGARPKRSQRNSMDSLDLLKFPSEKEQNGDSHHMGNLKITGKDFLSLPVRSRKYQEIQSSSERKSQDSNMSPLDASDVRLQVSQSKINRAPSDEECFFDLLSKFQSSRMDDQRCPLEECQAEAAEAAATSVPALEERISQSSLIASPQTEEFFDLIASSQSRRLDDQRASVGSLPGLRITHNNLGHLRMEGDAQEPGDEFFNMLMKCQSSRIDDQRCAPPDTVPRGPTMPDEDFFSLIQRVQAKRMDEQRVDLASAQEEQQQRRPSSS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1A4</accession>
    <name>A0A452J1A4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">THO complex subunit 2 N-terminal domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J1A4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038701.1">
        <property type="protein sequence ID" value="ENSGAGP00000034175.1"/>
        <property type="gene ID" value="ENSGAGG00000024263.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000445">
        <property type="term" value="C:THO complex part of transcription export complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003729">
        <property type="term" value="F:mRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006406">
        <property type="term" value="P:mRNA export from nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006397">
        <property type="term" value="P:mRNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040007">
        <property type="entry name" value="Tho2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032302">
        <property type="entry name" value="THOC2_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21597:SF0">
        <property type="entry name" value="THO COMPLEX SUBUNIT 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21597">
        <property type="entry name" value="THO2 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16134">
        <property type="entry name" value="THOC2_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="THO complex subunit 2 N-terminal" evidence="2">
        <location>
            <begin position="10"/>
            <end position="367"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="292"/>
            <end position="314"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="292"/>
            <end position="313"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF16134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034175.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="368" mass="42578" checksum="A3357AAF9AE26A6E" modified="2019-05-08" version="1">MLSENKSHENVTFRDIQQALYELANHVVKGNLKHEQASNVLSDVIEFRDDMSSILADVFCILDIETSCLEEKSKRDHFTQLVLACLYLVSDTVLKERLDPETLESLGLIKQSQQFNQKSVKIKTKLFYKQQKFNLLREENEGYAKLIAELGQDLSGNITSDLILENIKSLIGCFNLDPNRVLDIILEVYECRPEYDEFFVPLIESYMYMCEPQTLCHILGFKFKFYQEPNGETPSSLYRVAAVLLQHNLIDLEDLYVHLLPVDNAILEEHKREIVEAKQIVRKLTMVVLSSEKTEEKEKEKDKEEEKTEKPPDNQKLGLLEALLKVGDWQHAQSIMDQMPPFYATSHKPIAIALCQLVHVTVEPLYRR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1A5</accession>
    <name>A0A452J1A5_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="17">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
            <location evidence="1">Basement membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="12">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1A5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038607.1">
        <property type="protein sequence ID" value="ENSGAGP00000034095.1"/>
        <property type="gene ID" value="ENSGAGG00000024224.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005604">
        <property type="term" value="C:basement membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009887">
        <property type="term" value="P:animal organ morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007411">
        <property type="term" value="P:axon guidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009888">
        <property type="term" value="P:tissue development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00055">
        <property type="entry name" value="EGF_Lam"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000067">
        <property type="entry name" value="Laminin subunit gamma 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000441">
        <property type="entry name" value="Laminin subunit gamma 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000533">
        <property type="entry name" value="Laminin subunit gamma 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000174">
        <property type="entry name" value="Laminin subunit gamma-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050440">
        <property type="entry name" value="Laminin/Netrin_ECM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000034">
        <property type="entry name" value="Laminin_IV"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002049">
        <property type="entry name" value="LE_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10574:SF313">
        <property type="entry name" value="LAMININ SUBUNIT GAMMA-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10574">
        <property type="entry name" value="NETRIN/LAMININ-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00053">
        <property type="entry name" value="EGF_laminin"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00052">
        <property type="entry name" value="Laminin_B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00011">
        <property type="entry name" value="EGFLAMININ"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SMART" id="SM00180">
        <property type="entry name" value="EGF_Lam"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SMART" id="SM00281">
        <property type="entry name" value="LamB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01248">
        <property type="entry name" value="EGF_LAM_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50027">
        <property type="entry name" value="EGF_LAM_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51115">
        <property type="entry name" value="LAMININ_IVA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0084">Basement membrane</keyword>
    <keyword evidence="7" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="8 13" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="9 12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11 12" id="KW-0424">Laminin EGF-like domain</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="4 14" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="14">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5018969942" evidence="14">
        <location>
            <begin position="23"/>
            <end position="1149"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="15">
        <location>
            <begin position="80"/>
            <end position="138"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin IV type A" evidence="16">
        <location>
            <begin position="208"/>
            <end position="376"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="15">
        <location>
            <begin position="511"/>
            <end position="566"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="13">
        <location>
            <begin position="631"/>
            <end position="665"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="13">
        <location>
            <begin position="844"/>
            <end position="885"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="13">
        <location>
            <begin position="1012"/>
            <end position="1039"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="13">
        <location>
            <begin position="1077"/>
            <end position="1132"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="80"/>
            <end position="92"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="100"/>
            <end position="109"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="535"/>
            <end position="544"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004302"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
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        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023292"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00460"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
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        </source>
    </evidence>
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        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS51115"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034095.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1149" mass="126729" checksum="33C5B203A39BE778" modified="2019-05-08" version="1">QSTLGSLLRSLCLTVAIISVLSVCDCNGKSSQCIFDRDLLRETGNGYRCLNCIDNTDGIHCERCKEEFYRQRDGESCLPCNCSPRGSLGLQCDNYGRCSCKPGVMGDKCDRCQWGFHSFSEAGTLSFRLRSAQCDCDPAGSAGHCVSGRCVCKASVSGERCDRCKQGYYHLEAGNPEGCSQCFCYGHSAMCASAENYNIYKITSTFQQDSEGWRGVYVNGSPVQLQWSPRHRDVFVTARQSEPTYFLAPARFLGNQQVSYGQTLSFDYRMDRGGRRPSPQDVVLEGAGLRVTASLVPHGKMLPCGISKTYTFRLDEHPRSGWSPKLSYFEYRRLLGNLTALWIRATYGEYSTGYIDNVTLVSAQLTSGAPASWVEQCVCPPGYQGQFCERCAPGYKRDASNLGPFSTCVLCNCQGGGNCDPDTGDCYSGDENMDPNANCPFGFYRDPWNLQSCQLCPCRNGQGCSVAPGRKEVICNNCPLGVTGANCERCADGYFGNPLGDNGHVRPCQPCQCNNNVDPSAVGNCNRLTGECLKCLYNTAGFACDQCRDGFFGNPLALNPAEKCQACNCNSVGSDPLKCRSDGSCVCKPGFEGPNCEHTQCPACYSHVKTQVDQYLQQLQHLEVLVSRLQLGSGAEANEELEGKMQQAEETLREILRETLSLQASDKSLESRMSKMKGQESSYQSRLADIRVMVQRLQVLGHQYQAQVQDIRRLIEKAHLDLGQSKVTLSGLVSSPQDFSAQALQTSNIPGHCCRGQRNPLLLTSQGGSGLCVSWAVKHEVSVLAYDAFLFGRYDEAKLLASELETDTSRSALDADRAYQGSLVLLSSMSRFSKIDTGSFQKEANRLRQEADALMGLVETYMAEYRRLQSNTRTWEEEIKQLLQSGESKRLVSSQLLSRANLAKSRAQQALSAGNATFYEVELILKNLRGFHLQVDGRRREAEEAMRRLPLISNMVTNANDKTRRAEGALGSAATDAKTARSMAGEAKEITGGIQQEIRRLNLEANRTADGVLALEKGVITLLNEAREAERELTRKALEVDMDATTAQKAAVQETLHALEDFLHLMDQPGPVDEQALRLLELNLTAARTKNSQLKEQMAELEKTASLQRLRVQTLERSVNEILADIRNLEEIRNNLPPGCYNTNSIEKP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1A7</accession>
    <name>A0A452J1A7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="19">Glutamate receptor</fullName>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="19">Receptor for glutamate that functions as a ligand-gated ion channel in the central nervous system and plays an important role in excitatory synaptic transmission. L-glutamate acts as an excitatory neurotransmitter at many synapses in the central nervous system.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>Ca(2+)(in) = Ca(2+)(out)</text>
            <dbReference type="Rhea" id="RHEA:29671"/>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="14 19">Postsynaptic cell membrane</location>
            <topology evidence="14 19">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="19">Belongs to the glutamate-gated ion channel (TC 1.A.10.1) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1A7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038682.1">
        <property type="protein sequence ID" value="ENSGAGP00000034159.1"/>
        <property type="gene ID" value="ENSGAGG00000024221.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015276">
        <property type="term" value="F:ligand-gated monoatomic ion channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0038023">
        <property type="term" value="F:signaling receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd06382">
        <property type="entry name" value="PBP1_iGluR_Kainate"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.2300:FF:000010">
        <property type="entry name" value="Glutamate ionotropic receptor kainate type subunit 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.190.10:FF:000240">
        <property type="entry name" value="Glutamate receptor ionotropic, kainate 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.70:FF:000010">
        <property type="entry name" value="Putative glutamate receptor ionotropic kainate 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.70">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.2300">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.190.10">
        <property type="entry name" value="Periplasmic binding protein-like II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001828">
        <property type="entry name" value="ANF_lig-bd_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019594">
        <property type="entry name" value="Glu/Gly-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001508">
        <property type="entry name" value="Iono_Glu_rcpt_met"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015683">
        <property type="entry name" value="Ionotropic_Glu_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001320">
        <property type="entry name" value="Iontro_rcpt_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028082">
        <property type="entry name" value="Peripla_BP_I"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18966">
        <property type="entry name" value="IONOTROPIC GLUTAMATE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01094">
        <property type="entry name" value="ANF_receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00060">
        <property type="entry name" value="Lig_chan"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10613">
        <property type="entry name" value="Lig_chan-Glu_bd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00177">
        <property type="entry name" value="NMDARECEPTOR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00079">
        <property type="entry name" value="PBPe"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53822">
        <property type="entry name" value="Periplasmic binding protein-like I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53850">
        <property type="entry name" value="Periplasmic binding protein-like II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 19" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="8 18" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13 19" id="KW-0407">Ion channel</keyword>
    <keyword evidence="6 19" id="KW-0406">Ion transport</keyword>
    <keyword evidence="12 19" id="KW-1071">Ligand-gated ion channel</keyword>
    <keyword evidence="7 19" id="KW-0472">Membrane</keyword>
    <keyword evidence="11 19" id="KW-0628">Postsynaptic cell membrane</keyword>
    <keyword evidence="9 19" id="KW-0675">Receptor</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 19" id="KW-0770">Synapse</keyword>
    <keyword evidence="3 19" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 19" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="1 19" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="413"/>
            <end position="432"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="489"/>
            <end position="511"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="670"/>
            <end position="694"/>
        </location>
    </feature>
    <feature type="domain" description="Ionotropic glutamate receptor C-terminal" evidence="20">
        <location>
            <begin position="318"/>
            <end position="651"/>
        </location>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="366"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="368"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="373"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="539"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="540"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="588"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="site" description="Crucial to convey clamshell closure to channel opening" evidence="17">
        <location>
            <position position="518"/>
        </location>
    </feature>
    <feature type="site" description="Interaction with the cone snail toxin Con-ikot-ikot" evidence="17">
        <location>
            <position position="545"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="18">
        <location>
            <begin position="26"/>
            <end position="277"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="18">
        <location>
            <begin position="600"/>
            <end position="654"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023257"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00034104"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00036634"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="RuleBase" id="RU367118"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="SMART" id="SM00079"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034159.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="755" mass="85559" checksum="16A28192100E9D53" modified="2019-05-08" version="1">MPNTTLTYDIQRINLFDSFEASRRACDQLALGVAALFGPSHSSSVSAVQSICNALEVPHIQTRWKHPTVDNKDTFYINLYPDYAAISRAILDLVLYYNWKIVTVVYEDSTGLIRLQELIKAPSRYNIKIKIRQLPSGNKDARPLLKEMKKGKEFYVIFDCSHETAAEILKQILSMGMMTEYYHYFFTTLDLFALDLEPYRYSGVNMTGFRLLNIDNPQVTSVIEKWSMERLQAPPKPETGLLDGMMTTEAALMYDAVYMVAVASQRASQMTVSSLQCHRHKPWRFGPRFMNLIKEARWDGLTGHITFNKTDGLRKDFDLDIISLKEEGTEKVTNCVEGIKICLLNAKKMRIFSNLCSQKADLAVAPLTITYVREKVIDFSKPFMTLGISILYRKPNGTNPGVFSFLNPLSPDIWMYVLLACLGVSCVLFVIARFTPYEWYNPHPCNPDSDVVENNFTLLNSFWFGVGALMQQGSELMPKALSTRIVGGIWWFFTLIIISSYTANLAAFLTVERMESPIDSADDLAKQTKIEYGAVRDGSTMTFFKKSKISTYEKMWAFMSSRQQTALVKNNDEGIQRVLTTDYALLMESTSIEYVTQRNCNLTQIGGLIDSKGYGVGTPIGSPYRDKITIAILQLQEEGKLHMMKEKWWRGNGCPEEDSKEASALGVENIGGIFIVLAAGLVLSVFVAIGEFIYKSRKNSDIEQCLSFNAIMEELGISLRHQKKLKKRSRTKGRSSFTSILTCHQRRTLRKETMA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1B0</accession>
    <name>A0A452J1B0_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="20">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 16">Membrane</location>
            <topology evidence="1 16">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 16">Belongs to the integrin alpha chain family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1B0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038706.1">
        <property type="protein sequence ID" value="ENSGAGP00000034180.1"/>
        <property type="gene ID" value="ENSGAGG00000024233.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009897">
        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008305">
        <property type="term" value="C:integrin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033627">
        <property type="term" value="P:cell adhesion mediated by integrin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098609">
        <property type="term" value="P:cell-cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007160">
        <property type="term" value="P:cell-matrix adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007229">
        <property type="term" value="P:integrin-mediated signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.130:FF:000006">
        <property type="entry name" value="Integrin, alpha 9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.930">
        <property type="entry name" value="Bicelle-embedded integrin alpha(iib) transmembrane segment"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.130">
        <property type="entry name" value="Integrin alpha, N-terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1460">
        <property type="entry name" value="Integrin domains. Chain A, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1530">
        <property type="entry name" value="ntegrin, alpha v. Chain A, domain 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013517">
        <property type="entry name" value="FG-GAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013519">
        <property type="entry name" value="Int_alpha_beta-p"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000413">
        <property type="entry name" value="Integrin_alpha"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013649">
        <property type="entry name" value="Integrin_alpha_Ig-like_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048286">
        <property type="entry name" value="Integrin_alpha_Ig-like_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028994">
        <property type="entry name" value="Integrin_alpha_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032695">
        <property type="entry name" value="Integrin_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23220">
        <property type="entry name" value="INTEGRIN ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23220:SF78">
        <property type="entry name" value="INTEGRIN ALPHA-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01839">
        <property type="entry name" value="FG-GAP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08441">
        <property type="entry name" value="Integrin_A_Ig_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20806">
        <property type="entry name" value="Integrin_A_Ig_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01185">
        <property type="entry name" value="INTEGRINA"/>
    </dbReference>
    <dbReference type="SMART" id="SM00191">
        <property type="entry name" value="Int_alpha"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69318">
        <property type="entry name" value="Integrin alpha N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69179">
        <property type="entry name" value="Integrin domains"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51470">
        <property type="entry name" value="FG_GAP"/>
        <property type="match status" value="5"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0106">Calcium</keyword>
    <keyword evidence="8 16" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="14" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10 16" id="KW-0401">Integrin</keyword>
    <keyword evidence="11 16" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13 16" id="KW-0675">Receptor</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5 17" id="KW-0732">Signal</keyword>
    <keyword evidence="3 16" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9 16" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="17">
        <location>
            <begin position="1"/>
            <end position="48"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019329352" evidence="17">
        <location>
            <begin position="49"/>
            <end position="905"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="840"/>
            <end position="860"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="54"/>
            <end position="115"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="201"/>
            <end position="252"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="307"/>
            <end position="366"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="370"/>
            <end position="427"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="431"/>
            <end position="493"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha first immunoglubulin-like" evidence="18">
        <location>
            <begin position="478"/>
            <end position="583"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha third immunoglobulin-like" evidence="19">
        <location>
            <begin position="702"/>
            <end position="748"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023037"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00803"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="RuleBase" id="RU003762"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF08441"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="Pfam" id="PF20806"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034180.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="905" mass="100556" checksum="21123A0D81646166" modified="2019-05-08" version="1">MGSVGNAARAPRPGQCCFPRARGEHRAPVWAALLLLLLLGRCMRAAHAYNLDTQRPVVFKGLSGTLFGYSVVLHSHGPSRWLVVGAPKANWLANSSVVNPGAIFRCRIGSNPSGTCEQLQMGNPAGDYCGKTCFEERDNQWLGVSLSRQPRENGTIVACGHRWKNIYYIKNEHKLPNGVCYEIPPDFRTQLSKRISPCYKDYVRKFGENHGSCQAGMSSFYFKDLIIMGAPGSYYWTGSVFVYNTTANTFQAYTDSRNQVKFGSYLGYAVGAGHFLTPASTEVIGGAPQQEQTGKAYIFSIGTRALTILSELKGKKLGSYFGAAVCAVDLNTDGLSDLLVGAPMQSKVREEGRVYVYINSGSGAKMLELEIELAGSDSYAARFGESIANLGDIDNDGFEDVAIGAPQENNLEGAIYIYNGRKDGISSSFSQRIQGHRISNTLSMFGQSISSGIDADNNGYPDVAVGAFLDDSAVLLRTRPVVIVEAFLKHPNAVNRTNLECVENGQPAVCVNLMLCFTYKGRDVPGYVVLLYDLSVDVSRKTNTPARFYFSSNGTSDMVSGSMEIYNSNVTCKSHLAFLKENIDRWKTLPHEKKTYLSVGSMKTLLLNISLSNAGDDAYETALHIQLPKGLYFVRVLELFNDVTFYLKHLRHTQSFSTNSVSFVLQMDFSFLLDASSLTRAEDDLTITVNATCEENSPINCMKEKVNFTFHVISAGPSMAPAIGLEIMVPNSFAPRDEKLFNILDIQVCLFYYINILHTALYVTKASTHQGLIHLLQYLCNLAFFFFPYPQQDEASSLKFEISATASPEKNPKVIELHKERQIAHVFLEGLHNQKPKHHVTMLIIGTGSVLGIFLFLLIVLLLWKVGFFFLYQYFSIIFIDSFKNRNNMLYCDLFLLKKNKSWYD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1B5</accession>
    <name>A0A452J1B5_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="15">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="8">Ovomucin, the glycoprotein responsible for the gel properties of egg white, is composed for 2 subunits, alpha-ovomucin/MUC5B and beta-ovomucin/MUC6.</text>
    </comment>
    <comment type="subunit">
        <text evidence="9">Homomultimer; disulfide-linked. The N- and C-terminus mediate their assembly into higher order structures to form filaments. The CTCK domains of two polypeptides associate in the endoplasmic reticulum to generate intermolecularly disulfide-bonded dimers. These dimers progress to the Golgi apparatus, which is a more acidic environment than the endoplasmic reticulum. Under acidic conditions, the N-termini form non-covalent intermolecular interactions that juxtapose assemblies from different CTCK-linked dimers to produce long, disulfide-linked polymers that remain highly compact until secretion.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="10">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038711.1">
        <property type="protein sequence ID" value="ENSGAGP00000034185.1"/>
        <property type="gene ID" value="ENSGAGG00000024266.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046790">
        <property type="term" value="F:virion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042632">
        <property type="term" value="P:cholesterol homeostasis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030299">
        <property type="term" value="P:intestinal cholesterol absorption"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002281">
        <property type="term" value="P:macrophage activation involved in immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd19941">
        <property type="entry name" value="TIL"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000153">
        <property type="entry name" value="MUC5B isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000674">
        <property type="entry name" value="Mucin-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000414">
        <property type="entry name" value="von Willebrand factor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006207">
        <property type="entry name" value="Cys_knot_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050780">
        <property type="entry name" value="Mucin_vWF_Thrombospondin_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036084">
        <property type="entry name" value="Ser_inhib-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002919">
        <property type="entry name" value="TIL_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058753">
        <property type="entry name" value="TIL_OTOGL_Mucin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014853">
        <property type="entry name" value="VWF/SSPO/ZAN-like_Cys-rich_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001007">
        <property type="entry name" value="VWF_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001846">
        <property type="entry name" value="VWF_type-D"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025155">
        <property type="entry name" value="WxxW_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11339">
        <property type="entry name" value="EXTRACELLULAR MATRIX GLYCOPROTEIN RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11339:SF408">
        <property type="entry name" value="MUCIN-5B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08742">
        <property type="entry name" value="C8"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13330">
        <property type="entry name" value="Mucin2_WxxW"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01826">
        <property type="entry name" value="TIL"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25962">
        <property type="entry name" value="TIL_OTOGL_Mucin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00094">
        <property type="entry name" value="VWD"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00832">
        <property type="entry name" value="C8"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00041">
        <property type="entry name" value="CT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00214">
        <property type="entry name" value="VWC"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00215">
        <property type="entry name" value="VWC_out"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00216">
        <property type="entry name" value="VWD"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57567">
        <property type="entry name" value="Serine protease inhibitors"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01185">
        <property type="entry name" value="CTCK_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01225">
        <property type="entry name" value="CTCK_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01208">
        <property type="entry name" value="VWFC_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50184">
        <property type="entry name" value="VWFC_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51233">
        <property type="entry name" value="VWFD"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0186">Copper</keyword>
    <keyword evidence="6" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="7" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="3" id="KW-0732">Signal</keyword>
    <feature type="domain" description="VWFD" evidence="14">
        <location>
            <begin position="20"/>
            <end position="190"/>
        </location>
    </feature>
    <feature type="domain" description="VWFD" evidence="14">
        <location>
            <begin position="374"/>
            <end position="549"/>
        </location>
    </feature>
    <feature type="domain" description="VWFD" evidence="14">
        <location>
            <begin position="836"/>
            <end position="1007"/>
        </location>
    </feature>
    <feature type="domain" description="VWFD" evidence="14">
        <location>
            <begin position="2304"/>
            <end position="2488"/>
        </location>
    </feature>
    <feature type="domain" description="VWFC" evidence="13">
        <location>
            <begin position="2575"/>
            <end position="2645"/>
        </location>
    </feature>
    <feature type="domain" description="CTCK" evidence="12">
        <location>
            <begin position="2639"/>
            <end position="2706"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1421"/>
            <end position="1465"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1591"/>
            <end position="1611"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1764"/>
            <end position="1808"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1825"/>
            <end position="1870"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="2712"/>
            <end position="2746"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="2716"/>
            <end position="2727"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="2728"/>
            <end position="2746"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023008"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00054401"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00063950"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00039"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS01225"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50184"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS51233"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034185.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2746" mass="300326" checksum="76B91D9856C5DE2B" modified="2019-05-08" version="1">IQPLRTTLPIKPGNPAHNGRVCSTWGNFHFKTFDGDIFHFPGLCNYVFASHCKSAYEDFNIQIRRTVLESSPVISHITLKLDGMVIELTSDSIVVNGNPIQLPFSQSGILIERSSEYLKITAKMGMVFMWNQLDGLLVELDKKYANQTCGLCGDFNGIPIYNEFTSSSIELTDVQFGNMQKMDGPTEQCEDPTSSSLSNCSNEFSATCQTVLTGEAFVSCNALVDVQDYIDSCVQDLCRCDESMRDFCICNTFAEYSRQCAHAGGQPLNWRTKELCNKTCPYNMQHEECGSPCANTCTNPERSALCEDHCTDGCVCSPGTVFDDINSSGCIPVSECPCTYKGETYSPGASFASQCTSCNCTGGQWSCVSHSCPGTCSIEGGSHISTFDEKRYSFFGDCSYILTKLCDSDAFTVLGEVRKCGLTDTETCLKGVAISISGGQTIVVIKSSGSVFVNTIYTQLPFSAANVTIFRPSSFFIIVQTNFGLQIQMQLVPIMQLFVNVDPSHKGQTCGLCGNFNNIQTDDFKATSGIVEGTAAAFANTWKTQGDCPDVKNIFENPCTLSIENEKYAQHWCGLLTDTQGPFAECHSTVNPDNCLFDTCNCEKSEDCMCAALSTYVKACAAKGVLLNGWRTNVCTITSCPKSLTYSYVVNSCQPTCRSLSEPDVTCNIKFTPVDGCTCANGTYMDENGKCVPATSCPCYYKGSVLPSGEVVHDNGVICSCTHGKLNCVRVKPEPVCAAPMVYFDCNNITAGTIGSECQKSCQTLDMGCYSTQCVSGCVCPNDLVSDGKGGCVPVGECPCVHNEASYNPGETIKVNCNTCTCKNRMWQCTNESCLATCAVYGDGHYITFDGKRFSFSGGCEYTLVQDYCGKNSTSLGTFRVITENMPCGTTGTTCSKAIKVFLRNCELVFSDGKFEVIEKVRGGEVPYEVRYMGIYMVIDTDAGLTLMWDKKTSIFIKLSPDFKGQVCGLCGNYDGNGINDFTTRSQSVVEDVLEFGNSWKVSPTCPDAKSNKNPCRANPYRNSWAQKQCSIINSKVFSACHSQVEPNEYYEACVSDACACDTGGDCECFCTAVASYAQACNEAGICVAWRTPSICPLFCDYYNPQGECEWHYKPCGSPCMKTCRNPSGKCLHELQGLEGCYPKCPENKPYFDEDDMECVDHCGCYDTKGKYYKPGTPIRSEENCQSCECTMNGIKCEYDEYECHCTYDGKQYEYEDVIYNTTDGTGGCITATCGHNGTIDRKVYACSTPLTTTPFSFSSSTPISTTTVTPTTSVCVREVCQWSQWYDMSYPGSGINDGDFETFDNLRAKGYKVCKAPKAVECRSEKFPNTPLSELEQKVECSTTMGLICYNKDQHQMICDNFQIKILCCSFVPCEYTTPATTTPADTTISITKTTTSETTVTKRTPVTTTSTISTTLPTTTTVETTTSEATQSSTTTGSTTLPTTEIPTTSPKPTTPGTTTSSTTKCQPTCTWSKWFDVDFPASGPKEGDTETYNNIRAAGGIICAKPANIECRAENHPDISIEDVGQIVQCDVNFGLMCKNEDQIGKFKMCLNYEVRVLCCEENLNCPTPTRSIATTTIRTPPTTVTKTTIEVTTSTKPPSTTSLPTKEITTTSTISTTLPTTTSVEATTRTTTSSTTKCQPICTWSKWFDVDFPASGPKEGDIETYNNIRAAGGIICAKPANIECRAENYPDISIEEVGQIVQCDVNFGLMCKNEDQIGKFKMCLNYEVRVLCCEENLNCPTPTRSIATTTIRTPPTTVTKTTIEVTTSTKPPGKETPGTTTAETTSITTPKTTTRSTTGTTSLPTKQITTTSTISTTLPTTTSVEATTSEATQSTTTTGTTTLPTTEIPTTSPIPTTPGTTTSSATKCQPICTWSKWFDVDFPASGPKEGDIETYNNIRAAGGIICAKPANIECRAENHPDISIEEVDQIVQCDVNFGLMCKNEDQIGKFKMCLNYEVRVLCCEENLNCPTPTRSTATTTIRTPPTAVTKTTIETTTRSTTGTTSLPTKEITTTSTISTTLPTTTSVETTTTTKCQPICTWSKWFDVDFPASGPKEGDIETYNNIRAAGEIICAKPANIECRAENHPDISIEEVDQIVQCDVNFGLVCKNEDQIGKFKMCLNYEVRVLCCEANFNCPTPTRSTATTTVRTPPTTVTKSTIEVTSSTKPPEVTMRTTHCFCQIEDTVYVPGEMIYNETDKDGCNFYAICSKNCSVERFKGPCKSTTPATPTPIPTTSTSIPTMTPTATTGETWKISNCTTATCEGNNKVVIQKVKCPPVKKITCANRYPPVKNIWGFVFHGVCSGWGDPHYITFDGIYYTFLDNCTYTLVKQITPKYDNFRVDIDNYFCNAEDGLSCPQSIIVYYKYTVVVLTRELSNGVMANKIIFNKKVVNPGFQKDGISIYMLGINMVVEIPEIGATISFSGLIFSVKLPFSKFGNNTEGQCGTCTNTKTDDCRLPSGKIISSCTQMAPHWKVDDGKKQYCVGTPTATPEPSIPTPAPSCPPSALCKIILSEVFEECHKVIPPQDFYKGCVFDACHMTNTSMQCSSLEIYATLCATRGVCVDWRGETKGKCRKYCGINKYMDQWKSNCQDCVCDQYTFTVQCTKRPCKPLPPVSCDDPGFVPVQTLTQEDPCCGQTECRKNPLSSSIYSLEANVMQHKCKCCQEVKTSKRQVTLSCPDGSTTDYSYIHVEQCDCAGTECIPQTTPTTLAQEQQQQQQEQEQIPSKTGSLKTRKMNQNQQK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1B6</accession>
    <name>A0A452J1B6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Kinetochore protein NDC80</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">Acts as a component of the essential kinetochore-associated NDC80 complex, which is required for chromosome segregation and spindle checkpoint activity.</text>
    </comment>
    <comment type="subunit">
        <text evidence="10">Component of the NDC80 complex.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="10">Chromosome</location>
            <location evidence="10">Centromere</location>
            <location evidence="10">Kinetochore</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="10">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="1 10">Belongs to the NDC80/HEC1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1B6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038697.1">
        <property type="protein sequence ID" value="ENSGAGP00000034171.1"/>
        <property type="gene ID" value="ENSGAGG00000024278.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031262">
        <property type="term" value="C:Ndc80 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051315">
        <property type="term" value="P:attachment of mitotic spindle microtubules to kinetochore"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007051">
        <property type="term" value="P:spindle organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.418.30:FF:000002">
        <property type="entry name" value="NDC80, kinetochore complex component"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.1950">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.418.30">
        <property type="entry name" value="Ncd80 complex, Ncd80 subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040967">
        <property type="entry name" value="DUF5595"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005550">
        <property type="entry name" value="Kinetochore_Ndc80"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055260">
        <property type="entry name" value="Ndc80_CH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057091">
        <property type="entry name" value="NDC80_loop"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038273">
        <property type="entry name" value="Ndc80_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10643">
        <property type="entry name" value="KINETOCHORE PROTEIN NDC80"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10643:SF2">
        <property type="entry name" value="KINETOCHORE PROTEIN NDC80 HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18077">
        <property type="entry name" value="DUF5595"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03801">
        <property type="entry name" value="Ndc80_HEC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24487">
        <property type="entry name" value="NDC80_loop"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 10" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="3 10" id="KW-0132">Cell division</keyword>
    <keyword evidence="9 10" id="KW-0137">Centromere</keyword>
    <keyword evidence="2 10" id="KW-0158">Chromosome</keyword>
    <keyword evidence="6 11" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5 10" id="KW-0995">Kinetochore</keyword>
    <keyword evidence="4 10" id="KW-0498">Mitosis</keyword>
    <keyword evidence="7 10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Kinetochore protein Ndc80 CH" evidence="12">
        <location>
            <begin position="50"/>
            <end position="172"/>
        </location>
    </feature>
    <feature type="domain" description="DUF5595" evidence="13">
        <location>
            <begin position="190"/>
            <end position="261"/>
        </location>
    </feature>
    <feature type="domain" description="Kinetochore protein NDC80 loop region" evidence="14">
        <location>
            <begin position="353"/>
            <end position="584"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="11">
        <location>
            <begin position="234"/>
            <end position="376"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="11">
        <location>
            <begin position="441"/>
            <end position="546"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007050"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022454"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022776"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022838"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023328"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="RuleBase" id="RU368072"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF03801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="Pfam" id="PF18077"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="Pfam" id="PF24487"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034171.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="613" mass="70459" checksum="10D53627DBF660CA" modified="2019-05-08" version="1">MKQSSSSSGSNGRASLLPIRVQDTSRAGLYTPQATTSGSGVSRNTPWGFFTGTEKLKDPRPLHDKAYIQQCIRQLCEFLVENGYAQNVSVKSLQTPSTKDFFRIFAFIYTFLCPSYELPDSKIEEEIPKVFKELGYPFPLPKSSMYTVGVPHVWPQVAAALVWLIDCIKIYSVMRGNINVLVDAQCWSGEGEDEIVVNKLLMEYTEKCYDHFMNGGDAFEEMDAELQAKLNEVFKVDHSKLKTLEAENKRLNEEIARLERERESEPDRLEMLRKLKASLQADVEKYQAYMTNLESHSAVLNQKLHSLTEVCEESELEVEMMKQENARLQHTVDNQLYTAVDIERINHEKDELQQTINKLTKELETERQQLWAEELKYARGKESIEAQLDDYHKLARKLKLIPAGMENSRGHDFEIKFNPEDGPHSLAKYRRQINVPLMQIIDETEEAISKASNKKIILEDSLEQLNVMMAEKKSNIKLLKEEAQKLDDHHKQKLKVAEEEEEKSAAELQALEIHKNLIEAGVNEGLNEAMNELHEIQKKHQIAVRTTNEEKMQLLDKLYRVLEMVAFHVGSVEKYLAEHNAKADTVFEEFMSEDLLDNLKQILAKYKSKANAL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1C0</accession>
    <name>A0A452J1C0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Taste receptor type 1 member 2</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="17">Sweet taste receptor T1R2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">Putative taste receptor. TAS1R2/TAS1R3 recognizes diverse natural and synthetic sweeteners.</text>
    </comment>
    <comment type="subunit">
        <text evidence="15">Forms heterodimers with TAS1R3.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="14">Belongs to the G-protein coupled receptor 3 family. TAS1R subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1C0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038716.1">
        <property type="protein sequence ID" value="ENSGAGP00000034190.1"/>
        <property type="gene ID" value="ENSGAGG00000024281.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903767">
        <property type="term" value="C:sweet taste receptor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004930">
        <property type="term" value="F:G protein-coupled receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033041">
        <property type="term" value="F:sweet taste receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.2300:FF:000016">
        <property type="entry name" value="Taste 1 receptor member 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.50.30:FF:000004">
        <property type="entry name" value="Taste receptor type 1 member 3-like protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.2300">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.50.30">
        <property type="entry name" value="GPCR, family 3, nine cysteines domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001828">
        <property type="entry name" value="ANF_lig-bd_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000337">
        <property type="entry name" value="GPCR_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011500">
        <property type="entry name" value="GPCR_3_9-Cys_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038550">
        <property type="entry name" value="GPCR_3_9-Cys_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017978">
        <property type="entry name" value="GPCR_3_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000068">
        <property type="entry name" value="GPCR_3_Ca_sens_rcpt-rel"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017979">
        <property type="entry name" value="GPCR_3_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028082">
        <property type="entry name" value="Peripla_BP_I"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24061">
        <property type="entry name" value="CALCIUM-SENSING RECEPTOR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24061:SF517">
        <property type="entry name" value="TASTE RECEPTOR TYPE 1 MEMBER 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00003">
        <property type="entry name" value="7tm_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01094">
        <property type="entry name" value="ANF_receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07562">
        <property type="entry name" value="NCD3G"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00248">
        <property type="entry name" value="GPCRMGR"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53822">
        <property type="entry name" value="Periplasmic binding protein-like I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00980">
        <property type="entry name" value="G_PROTEIN_RECEP_F3_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50259">
        <property type="entry name" value="G_PROTEIN_RECEP_F3_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="8" id="KW-0297">G-protein coupled receptor</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9 18" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0675">Receptor</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0716">Sensory transduction</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="3" id="KW-0919">Taste</keyword>
    <keyword evidence="12" id="KW-0807">Transducer</keyword>
    <keyword evidence="5 18" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 18" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="18">
        <location>
            <begin position="557"/>
            <end position="584"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="18">
        <location>
            <begin position="596"/>
            <end position="617"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="18">
        <location>
            <begin position="629"/>
            <end position="652"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="18">
        <location>
            <begin position="672"/>
            <end position="692"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="18">
        <location>
            <begin position="721"/>
            <end position="742"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="18">
        <location>
            <begin position="754"/>
            <end position="773"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="18">
        <location>
            <begin position="779"/>
            <end position="801"/>
        </location>
    </feature>
    <feature type="domain" description="G-protein coupled receptors family 3 profile" evidence="19">
        <location>
            <begin position="558"/>
            <end position="824"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022480"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022606"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023224"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00037659"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00038492"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00038699"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00040704"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00042616"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50259"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034190.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="832" mass="94223" checksum="487FA23848DB6181" modified="2019-05-08" version="1">MGPLASCLSHYVLLTMAGSQTQYSDFHLAGDYLLGGLFSLHTETKSSPHLSYSAVPTCQAYKLKKASYSHLRAMRFAVEQINNSSTLLPGVSLGYEMVDVCYLTNFIHPILYFLANKHSVVQLHNNYTHYQPRVLAVIGPDSSQAVVIAAQVLSLFLVPQISYTATTNELSDRKRFPTAFQTISSMEQQIVAMQLLLKEFRWNWINILYSDDDYGRQNLKLLRARATEICVALQEVIPVPQSSQMLSLELQGKIKSVVRKVIHSTAKVVIVLSLELILPFFFQEAVRQNVTDLVWIATEAWATEPALYNITDLWRLGTILGVAVKEFSIPGLDDFRVSLAEGANEQASDQTTCNQECDQCLRAAQDAEHSMRAQGERTDFSVYSSVYIVAHALHRLLGCNTTGCHKRMVYPWQLVPEVKRVNFSLLNERINFRERGTIPNSFEIIQWKWDQPNNPFQKIASYSRNKKLLFTIQNISWHTPNNTVPISVCSQQCEYGQRRKPVTVSSCCFECIDCEEGTFLNNNDLYNCQPCPADQWSNARSQKCYPRLLKYLEWQEGIAVVLLLFSVLGLVATVVISVIFAIHLHTPVVKSAGGRMCFLMLASLIIGFINVPVYMGIPTEFKCMCRQIVYTLCFTVCISCLTVRSFQIISIFKMAARLPKAYSYWMKYNGQYIFITVILALKVTIVVTNVIIHPSIPVPLVLEDDPSTVELQCNKSFKLSLLMNNSLDTFLSILCFYFSYIGKELPKNYNEAKYITLCMTCYAVSEIAIFLAMSVYRGMLVTICDVIGVVINLLGISIGYFGPKCYLIFFHPERNTAAYFQSVIQSYTMRQD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1C2</accession>
    <name>A0A452J1C2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Kinetochore protein NDC80</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">Acts as a component of the essential kinetochore-associated NDC80 complex, which is required for chromosome segregation and spindle checkpoint activity.</text>
    </comment>
    <comment type="subunit">
        <text evidence="10">Component of the NDC80 complex.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="10">Chromosome</location>
            <location evidence="10">Centromere</location>
            <location evidence="10">Kinetochore</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="10">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="1 10">Belongs to the NDC80/HEC1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1C2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038700.1">
        <property type="protein sequence ID" value="ENSGAGP00000034174.1"/>
        <property type="gene ID" value="ENSGAGG00000024278.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031262">
        <property type="term" value="C:Ndc80 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051315">
        <property type="term" value="P:attachment of mitotic spindle microtubules to kinetochore"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007051">
        <property type="term" value="P:spindle organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.418.30:FF:000002">
        <property type="entry name" value="NDC80, kinetochore complex component"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.1950">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.418.30">
        <property type="entry name" value="Ncd80 complex, Ncd80 subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005550">
        <property type="entry name" value="Kinetochore_Ndc80"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055260">
        <property type="entry name" value="Ndc80_CH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057091">
        <property type="entry name" value="NDC80_loop"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038273">
        <property type="entry name" value="Ndc80_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10643">
        <property type="entry name" value="KINETOCHORE PROTEIN NDC80"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10643:SF2">
        <property type="entry name" value="KINETOCHORE PROTEIN NDC80 HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03801">
        <property type="entry name" value="Ndc80_HEC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24487">
        <property type="entry name" value="NDC80_loop"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 10" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="3 10" id="KW-0132">Cell division</keyword>
    <keyword evidence="9 10" id="KW-0137">Centromere</keyword>
    <keyword evidence="2 10" id="KW-0158">Chromosome</keyword>
    <keyword evidence="6 11" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5 10" id="KW-0995">Kinetochore</keyword>
    <keyword evidence="4 10" id="KW-0498">Mitosis</keyword>
    <keyword evidence="7 10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Kinetochore protein Ndc80 CH" evidence="12">
        <location>
            <begin position="66"/>
            <end position="196"/>
        </location>
    </feature>
    <feature type="domain" description="Kinetochore protein NDC80 loop region" evidence="13">
        <location>
            <begin position="347"/>
            <end position="578"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="11">
        <location>
            <begin position="228"/>
            <end position="370"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="11">
        <location>
            <begin position="435"/>
            <end position="540"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007050"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022454"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022776"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022838"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023328"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="RuleBase" id="RU368072"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF03801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="Pfam" id="PF24487"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034174.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="607" mass="69783" checksum="B3902A60CFFABA80" modified="2019-05-08" version="1">MKQSSSSSGSNGRASLLPIRVQDTSRAGLYTPQATERSKLSISKYMGGSEGKTSFSVKRTSGSGVSRNTPWGFFTGTEKLKDPRPLHDKAYIQQCIRQLCEFLVENGYAQNVSVKSLQTPSTKDFFRIFAFIYTFLCPSYELPDSKIEEEIPKVFKELGYPFPLPKSSMYTVGVPHVWPQVAAALVWLIDCIKLLMEYTEKCYDHFMNGGDAFEEMDAELQAKLNEVFKVDHSKLKTLEAENKRLNEEIARLERERESEPDRLEMLRKLKASLQADVEKYQAYMTNLESHSAVLNQKLHSLTEVCEESELEVEMMKQENARLQHTVDNQLYTAVDIERINHEKDELQQTINKLTKELETERQQLWAEELKYARGKESIEAQLDDYHKLARKLKLIPAGMENSRGHDFEIKFNPEDGPHSLAKYRRQINVPLMQIIDETEEAISKASNKKIILEDSLEQLNVMMAEKKSNIKLLKEEAQKLDDHHKQKLKVAEEEEEKSAAELQALEIHKNLIEAGVNEGLNEAMNELHEIQKKHQIAVRTTNEEKMQLLDKLYRVLEMVAFHVGSVEKYLAEHNAKADTVFEEFMSEDLLDNLKQILAKYKSKANAL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1C4</accession>
    <name>A0A452J1C4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="22">Cytosolic purine 5'-nucleotidase</fullName>
            <ecNumber evidence="21">2.7.1.77</ecNumber>
            <ecNumber evidence="4">3.1.3.5</ecNumber>
            <ecNumber evidence="5">3.1.3.99</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="23">Cytosolic nucleoside phosphotransferase 5'N</fullName>
        </alternativeName>
    </protein>
    <organism evidence="34 35">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="35" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="34" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="34" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="24">Broad specificity cytosolic 5'-nucleotidase that catalyzes the dephosphorylation of 6-hydroxypurine nucleoside 5'-monophosphates. In addition, possesses a phosphotransferase activity by which it can transfer a phosphate from a donor nucleoside monophosphate to an acceptor nucleoside, preferably inosine, deoxyinosine and guanosine. Has the highest activities for IMP and GMP followed by dIMP, dGMP and XMP. Could also catalyze the transfer of phosphates from pyrimidine monophosphates but with lower efficiency. Through these activities regulates the purine nucleoside/nucleotide pools within the cell.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>GMP + H2O = guanosine + phosphate</text>
            <dbReference type="Rhea" id="RHEA:27714"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:16750"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:58115"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="19">
            <dbReference type="Rhea" id="RHEA:27715"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="28">
            <text>IMP + H2O = inosine + phosphate</text>
            <dbReference type="Rhea" id="RHEA:27718"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:17596"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:58053"/>
            <dbReference type="EC" id="3.1.3.99"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="28">
            <dbReference type="Rhea" id="RHEA:27719"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="26">
            <text>XMP + H2O = xanthosine + phosphate</text>
            <dbReference type="Rhea" id="RHEA:28530"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:18107"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:57464"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="26">
            <dbReference type="Rhea" id="RHEA:28531"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="30">
            <text>a 2'-deoxyribonucleoside + a ribonucleoside 5'-phosphate = a ribonucleoside + a 2'-deoxyribonucleoside 5'-phosphate</text>
            <dbReference type="Rhea" id="RHEA:19961"/>
            <dbReference type="ChEBI" id="CHEBI:18254"/>
            <dbReference type="ChEBI" id="CHEBI:18274"/>
            <dbReference type="ChEBI" id="CHEBI:58043"/>
            <dbReference type="ChEBI" id="CHEBI:65317"/>
            <dbReference type="EC" id="2.7.1.77"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>a ribonucleoside 5'-phosphate + H2O = a ribonucleoside + phosphate</text>
            <dbReference type="Rhea" id="RHEA:12484"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:18254"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:58043"/>
            <dbReference type="EC" id="3.1.3.5"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="15">
            <dbReference type="Rhea" id="RHEA:12485"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="20">
            <text>dGMP + H2O = 2'-deoxyguanosine + phosphate</text>
            <dbReference type="Rhea" id="RHEA:29379"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:17172"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:57673"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="20">
            <dbReference type="Rhea" id="RHEA:29380"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>dGMP + inosine = 2'-deoxyguanosine + IMP</text>
            <dbReference type="Rhea" id="RHEA:69580"/>
            <dbReference type="ChEBI" id="CHEBI:17172"/>
            <dbReference type="ChEBI" id="CHEBI:17596"/>
            <dbReference type="ChEBI" id="CHEBI:57673"/>
            <dbReference type="ChEBI" id="CHEBI:58053"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="16">
            <text>dIMP + H2O = 2'-deoxyinosine + phosphate</text>
            <dbReference type="Rhea" id="RHEA:29383"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:28997"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:61194"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="16">
            <dbReference type="Rhea" id="RHEA:29384"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>dIMP + inosine = 2'-deoxyinosine + IMP</text>
            <dbReference type="Rhea" id="RHEA:69572"/>
            <dbReference type="ChEBI" id="CHEBI:17596"/>
            <dbReference type="ChEBI" id="CHEBI:28997"/>
            <dbReference type="ChEBI" id="CHEBI:58053"/>
            <dbReference type="ChEBI" id="CHEBI:61194"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="29">
            <text>inosine + AMP = IMP + adenosine</text>
            <dbReference type="Rhea" id="RHEA:69596"/>
            <dbReference type="ChEBI" id="CHEBI:16335"/>
            <dbReference type="ChEBI" id="CHEBI:17596"/>
            <dbReference type="ChEBI" id="CHEBI:58053"/>
            <dbReference type="ChEBI" id="CHEBI:456215"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="31">
            <text>inosine + CMP = cytidine + IMP</text>
            <dbReference type="Rhea" id="RHEA:69592"/>
            <dbReference type="ChEBI" id="CHEBI:17562"/>
            <dbReference type="ChEBI" id="CHEBI:17596"/>
            <dbReference type="ChEBI" id="CHEBI:58053"/>
            <dbReference type="ChEBI" id="CHEBI:60377"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="27">
            <text>inosine + GMP = guanosine + IMP</text>
            <dbReference type="Rhea" id="RHEA:69584"/>
            <dbReference type="ChEBI" id="CHEBI:16750"/>
            <dbReference type="ChEBI" id="CHEBI:17596"/>
            <dbReference type="ChEBI" id="CHEBI:58053"/>
            <dbReference type="ChEBI" id="CHEBI:58115"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="25">
            <text>inosine + UMP = uridine + IMP</text>
            <dbReference type="Rhea" id="RHEA:69588"/>
            <dbReference type="ChEBI" id="CHEBI:16704"/>
            <dbReference type="ChEBI" id="CHEBI:17596"/>
            <dbReference type="ChEBI" id="CHEBI:57865"/>
            <dbReference type="ChEBI" id="CHEBI:58053"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="33">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
        <text evidence="33">Binds 1 Mg(2+) ion per subunit.</text>
    </comment>
    <comment type="subunit">
        <text evidence="3">Homotetramer.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytosol</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the 5'(3')-deoxyribonucleotidase family.</text>
    </comment>
    <dbReference type="EC" id="2.7.1.77" evidence="21"/>
    <dbReference type="EC" id="3.1.3.5" evidence="4"/>
    <dbReference type="EC" id="3.1.3.99" evidence="5"/>
    <dbReference type="AlphaFoldDB" id="A0A452J1C4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038707.1">
        <property type="protein sequence ID" value="ENSGAGP00000034181.1"/>
        <property type="gene ID" value="ENSGAGG00000024283.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008253">
        <property type="term" value="F:5'-nucleotidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050146">
        <property type="term" value="F:nucleoside phosphotransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046085">
        <property type="term" value="P:adenosine metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046037">
        <property type="term" value="P:GMP metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046040">
        <property type="term" value="P:IMP metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07522">
        <property type="entry name" value="HAD_cN-II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1000:FF:000021">
        <property type="entry name" value="NT5C2 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1000">
        <property type="entry name" value="HAD superfamily/HAD-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036412">
        <property type="entry name" value="HAD-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008380">
        <property type="entry name" value="HAD-SF_hydro_IG_5-nucl"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023214">
        <property type="entry name" value="HAD_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016695">
        <property type="entry name" value="Pur_nucleotidase"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR02244">
        <property type="entry name" value="HAD-IG-Ncltidse"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12103">
        <property type="entry name" value="5'-NUCLEOTIDASE DOMAIN-CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12103:SF17">
        <property type="entry name" value="CYTOSOLIC PURINE 5'-NUCLEOTIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05761">
        <property type="entry name" value="5_nucleotid"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF017434">
        <property type="entry name" value="Purine_5'-nucleotidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56784">
        <property type="entry name" value="HAD-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0021">Allosteric enzyme</keyword>
    <keyword evidence="12" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="13 33" id="KW-0460">Magnesium</keyword>
    <keyword evidence="9 33" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="14" id="KW-0546">Nucleotide metabolism</keyword>
    <keyword evidence="10" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="35" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0808">Transferase</keyword>
    <feature type="active site" description="Nucleophile" evidence="32">
        <location>
            <position position="34"/>
        </location>
    </feature>
    <feature type="active site" description="Proton donor" evidence="32">
        <location>
            <position position="36"/>
        </location>
    </feature>
    <feature type="binding site" evidence="33">
        <location>
            <position position="34"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="33">
        <location>
            <position position="36"/>
        </location>
        <ligand>
            <name>GMP</name>
            <dbReference type="ChEBI" id="CHEBI:58115"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="33">
        <location>
            <position position="333"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004514"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009589"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00011881"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00012643"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012894"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022533"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023080"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00035871"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00036191"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00036258"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00036593"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00036695"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00036911"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00038866"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00040391"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00042328"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00046009"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00047396"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00047560"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="ARBA" id="ARBA00048038"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="ARBA" id="ARBA00048127"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="ARBA" id="ARBA00048155"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="ARBA" id="ARBA00048156"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="31">
        <source>
            <dbReference type="ARBA" id="ARBA00048783"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="32">
        <source>
            <dbReference type="PIRSR" id="PIRSR017434-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="33">
        <source>
            <dbReference type="PIRSR" id="PIRSR017434-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="34">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034181.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="35">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="519" mass="59876" checksum="8B41303562977C38" modified="2019-05-08" version="1">MIFSCCCSGTEDVYMLVFVNRSLAMEKIKCFGFDMDYTLAVYKSPEYESLGFDLTVERLVSIGYPQELLSFVYDPSFPTRGLVFDTLYGNLLKVDAYGNLLVCTHGFNFVRGPETREQYPNKFIQRDDTERFYILNTLFNLPETYLLACLVDFFTNCDRYSSCEKGFKDGDLFMSFRSMFQDVRDAVDWVHYQGTLKEKTLENLEKYVVKDGKMPLLLSRMNEVGKVFLATNSDYKYTDKIMTYLFDFSHGPKPGSSHQPWQSYFDLILVDARKPLFFGEGTVLRQVDTVTGKLKIGTYTGPLQHGIVYSGGSSDTICDLLGAKGKDILYIGDHIFGDILKSKKRQGWRTFLVIPELAQELHVWTDKSSLFEELQSLDIFLAELYKHLDSSSNERPDISSIQRRIKKVTHDMDMCYGMMGSLFRSGSRQTLFASQVMRYADLYAASFINLLYYPFSYLFRAAHVLMPHESTVEHIHVDINEKESPLATRNRTSVDFKDSDYKRHQLTRKGKESMKVSQR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1D0</accession>
    <name>A0A452J1D0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">5'-nucleotidase, cytosolic II</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J1D0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038710.1">
        <property type="protein sequence ID" value="ENSGAGP00000034184.1"/>
        <property type="gene ID" value="ENSGAGG00000024283.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016787">
        <property type="term" value="F:hydrolase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036412">
        <property type="entry name" value="HAD-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008380">
        <property type="entry name" value="HAD-SF_hydro_IG_5-nucl"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05761">
        <property type="entry name" value="5_nucleotid"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56784">
        <property type="entry name" value="HAD-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="3" id="KW-0460">Magnesium</keyword>
    <keyword evidence="1" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034184.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="6">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="69" mass="8038" checksum="A5F2182B6A86D986" modified="2019-05-08" version="1">MTTSWSDRLQNAADLPANMDRHALKKYRREAYHRVFVNRSLAMEKIKCFGFDMDYTLAGEPSVTPSRLE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1D1</accession>
    <name>A0A452J1D1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Integrin alpha second immunoglobulin-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 13">Membrane</location>
            <topology evidence="1 13">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 13">Belongs to the integrin alpha chain family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1D1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038726.1">
        <property type="protein sequence ID" value="ENSGAGP00000034200.1"/>
        <property type="gene ID" value="ENSGAGG00000024233.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009897">
        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008305">
        <property type="term" value="C:integrin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033627">
        <property type="term" value="P:cell adhesion mediated by integrin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098609">
        <property type="term" value="P:cell-cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007160">
        <property type="term" value="P:cell-matrix adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007229">
        <property type="term" value="P:integrin-mediated signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.930">
        <property type="entry name" value="Bicelle-embedded integrin alpha(iib) transmembrane segment"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.130">
        <property type="entry name" value="Integrin alpha, N-terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1460">
        <property type="entry name" value="Integrin domains. Chain A, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1510">
        <property type="entry name" value="ntegrin, alpha v. Chain A, domain 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013517">
        <property type="entry name" value="FG-GAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013519">
        <property type="entry name" value="Int_alpha_beta-p"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000413">
        <property type="entry name" value="Integrin_alpha"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048285">
        <property type="entry name" value="Integrin_alpha_Ig-like_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028994">
        <property type="entry name" value="Integrin_alpha_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032695">
        <property type="entry name" value="Integrin_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23220">
        <property type="entry name" value="INTEGRIN ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23220:SF78">
        <property type="entry name" value="INTEGRIN ALPHA-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01839">
        <property type="entry name" value="FG-GAP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20805">
        <property type="entry name" value="Integrin_A_Ig_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01185">
        <property type="entry name" value="INTEGRINA"/>
    </dbReference>
    <dbReference type="SMART" id="SM00191">
        <property type="entry name" value="Int_alpha"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69318">
        <property type="entry name" value="Integrin alpha N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69179">
        <property type="entry name" value="Integrin domains"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51470">
        <property type="entry name" value="FG_GAP"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6 13" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="8 13" id="KW-0401">Integrin</keyword>
    <keyword evidence="9 13" id="KW-0472">Membrane</keyword>
    <keyword evidence="10 13" id="KW-0675">Receptor</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="4 13" id="KW-0732">Signal</keyword>
    <keyword evidence="3 13" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 13" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="13">
        <location>
            <begin position="1"/>
            <end position="35"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5018814913" description="Integrin alpha second immunoglobulin-like domain-containing protein" evidence="13">
        <location>
            <begin position="36"/>
            <end position="928"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="13">
        <location>
            <begin position="865"/>
            <end position="887"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="12">
        <location>
            <begin position="41"/>
            <end position="102"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="12">
        <location>
            <begin position="188"/>
            <end position="239"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="12">
        <location>
            <begin position="294"/>
            <end position="353"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="12">
        <location>
            <begin position="357"/>
            <end position="414"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha second immunoglobulin-like" evidence="14">
        <location>
            <begin position="547"/>
            <end position="691"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023037"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00803"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="RuleBase" id="RU003762"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="Pfam" id="PF20805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034200.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="928" mass="103508" checksum="033FA9829AA6CEF5" modified="2019-05-08" version="1">AAHAARGGSGMRGAPVWAALLLLLLLGRCMRAAHAYNLDTQRPVVFKGLSGTLFGYSVVLHSHGPSRWLVVGAPKANWLANSSVVNPGAIFRCRIGSNPSGTCEQLQMGNPAGDYCGKTCFEERDNQWLGVSLSRQPRENGTIVACGHRWKNIYYIKNEHKLPNGVCYEIPPDFRTQLSKRISPCYKDYVRKFGENHGSCQAGMSSFYFKDLIIMGAPGSYYWTGSVFVYNTTANTFQAYTDSRNQVKFGSYLGYAVGAGHFLTPASTEVIGGAPQQEQTGKAYIFSIGTRALTILSELKGKKLGSYFGAAVCAVDLNTDGLSDLLVGAPMQSKVREEGRVYVYINSGSGAKMLELEIELAGSDSYAARFGESIANLGDIDNDGFEDVAIGAPQENNLEGAIYIYNGRKDGISSSFSQRIQGHRISNTLSMFGQSISNTIINIIFENNKFVQRTRPVVIVEAFLKHPNAVNRTNLECVENGQPAVCVNLMLCFTYKGRDVPGYVVLLYDLSVDVSRKTNTPARFYFSSNGTSDMVSGSMEIYNSNVTCKSHLAFLKENIERAKDRWKTLPHEKKTYLSVGSMKTLLLNISLSNAGDDAYETALHIQLPKGLYFVRVLELEEKQIHCEISDKEVHAVRLDCSVGYLYVDHLSKMDFSFLLDASSLTRAEDDLTITVNATCEENSPINCMKEKVNFTFHVISAGPSMAPAIGLEIMVPNSFAPRDEKLFNILDIQTADGECFYNNYTRDCTLPEKNGHMNILEDLVIFFSKPTKRRLYCMKEDHLCLRILCKFGNMENGKEAIVRMQLEATPSVLEMDEASSLKFEISATASPEKNPKVIELHKERQIAHVFLEGLHNQKPKHHVTMLIIGTGSVLGIFLFLLIVLLLWKVGFFFLYQYFSIIFIDSFKNRNNMLYCDLFLLKKNKSWYD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1D3</accession>
    <name>A0A452J1D3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">VWFD domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="7">Ovomucin, the glycoprotein responsible for the gel properties of egg white, is composed for 2 subunits, alpha-ovomucin/MUC5B and beta-ovomucin/MUC6.</text>
    </comment>
    <comment type="subunit">
        <text evidence="8">Homomultimer; disulfide-linked. The N- and C-terminus mediate their assembly into higher order structures to form filaments. The CTCK domains of two polypeptides associate in the endoplasmic reticulum to generate intermolecularly disulfide-bonded dimers. These dimers progress to the Golgi apparatus, which is a more acidic environment than the endoplasmic reticulum. Under acidic conditions, the N-termini form non-covalent intermolecular interactions that juxtapose assemblies from different CTCK-linked dimers to produce long, disulfide-linked polymers that remain highly compact until secretion.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1D3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038717.1">
        <property type="protein sequence ID" value="ENSGAGP00000034191.1"/>
        <property type="gene ID" value="ENSGAGG00000024266.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031012">
        <property type="term" value="C:extracellular matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046790">
        <property type="term" value="F:virion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042632">
        <property type="term" value="P:cholesterol homeostasis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030299">
        <property type="term" value="P:intestinal cholesterol absorption"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002281">
        <property type="term" value="P:macrophage activation involved in immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd19941">
        <property type="entry name" value="TIL"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000153">
        <property type="entry name" value="MUC5B isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000674">
        <property type="entry name" value="Mucin-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000414">
        <property type="entry name" value="von Willebrand factor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050780">
        <property type="entry name" value="Mucin_vWF_Thrombospondin_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036084">
        <property type="entry name" value="Ser_inhib-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002919">
        <property type="entry name" value="TIL_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058753">
        <property type="entry name" value="TIL_OTOGL_Mucin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014853">
        <property type="entry name" value="VWF/SSPO/ZAN-like_Cys-rich_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001007">
        <property type="entry name" value="VWF_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001846">
        <property type="entry name" value="VWF_type-D"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11339">
        <property type="entry name" value="EXTRACELLULAR MATRIX GLYCOPROTEIN RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11339:SF408">
        <property type="entry name" value="MUCIN-5B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08742">
        <property type="entry name" value="C8"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01826">
        <property type="entry name" value="TIL"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25962">
        <property type="entry name" value="TIL_OTOGL_Mucin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00094">
        <property type="entry name" value="VWD"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00832">
        <property type="entry name" value="C8"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00215">
        <property type="entry name" value="VWC_out"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00216">
        <property type="entry name" value="VWD"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57603">
        <property type="entry name" value="FnI-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57567">
        <property type="entry name" value="Serine protease inhibitors"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51233">
        <property type="entry name" value="VWFD"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="3" id="KW-0732">Signal</keyword>
    <feature type="domain" description="VWFD" evidence="9">
        <location>
            <begin position="40"/>
            <end position="203"/>
        </location>
    </feature>
    <feature type="domain" description="VWFD" evidence="9">
        <location>
            <begin position="387"/>
            <end position="562"/>
        </location>
    </feature>
    <feature type="domain" description="VWFD" evidence="9">
        <location>
            <begin position="833"/>
            <end position="1004"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00054401"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00063950"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51233"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034191.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1185" mass="129886" checksum="8F377BE127C09EC3" modified="2019-05-08" version="1">GPNPTPVEVKWCLCIDLKGNRIGPTTCFESTAGNPAHNGRVCSTWGNFHFKTFDGDIFHFPGLCNYVFASHCKSAYEDFNIQIRRTVLESSPVISHITLKLDGMVIELTSDSIVVNGNPQSGILIERSSEYLKITAKMGMVFMWNQLDGLLVELDKKYANQTCGLCGDFNGIPIYNEFTSSIELTDVQFGNMQKMDGPTEQCEDPTSSSLSNCSNEFSATCQTVLTGEAFVSCNALVDVQDYIDSCVQDLCRCDESMRDFCICNTFAEYSRQCAHAGGQPLNWRTKELCNKTCPYNMQHEECGSPCANTCTNPERSALCEDHCTDGCVCSPGTVFDDINSSGCIPVSECPCTYKGETYSPGASFASQCTSCNCTGGQWSCVSHSCPGTCSIEGGSHISTFDEKRYSFFGDCSYILTKLCDSDAFTVLGEVRKCGLTDTETCLKGVAISISGGQTVRSKNRYPTFRLNVFKTDSYWVVTNVTIFRPSSFFIIVQTNFGLQIQMQLVPIMQLFVNVDPSHKGQTCGLCGNFNNIQTDDFKATSGIVEGTAAAFANTWKTQGDCPDVKNIFENPCTLSIENEKYAQHWCGLLTDTQGPFAECHSTVNPDVYHTNCLFDTCNCEKSEDCMCAALSTYVKACAAKGVLLNGWRTNVCTITSCPKSLTYSYVVNSCQPTCRSLSEPDVTCNIKFTPVDGCTCANGTYMDENGKCVPATSCPCYYKGSVLPSGEVVHDNVCAAPMVYFDCNNITAGTIGSECQKSCQTLDMGCYSTQCVSGCVCPNDLVSDGKGGCVPVGECPCVHNEASYNPGETIKVNCNTCTCKNRMWQCTNESCLATCAVYGDGHYITFDGKRFSFSGGCEYTLVQDYCGKNSTSLGTFRVITENMPCGTTGTTCSKAIKVFLRNCELVFSDGKFEVIEKVRGGEVPYEVRYMGIYMVIDTDAGLTLMWDKKTSIFIKLSPDFKGQVCGLCGNYDGNGINDFTTRSQSVVEDVLEFGNSWKVSPTCPDAKSNKNPCRANPYRNSWAQKQCSIINSKVFSACHSQVEPNEYYEACVSDACACDTGGDCECFCTAVASYAQACNEAGICVAWRTPSICPLFCDYYNPQGECEWHYKPCGSPCMKTCRNPSGKCLHELQGLEGCYPKCPENKPYFDEDDMECVDHCGCYDTKGKYYKPGTPIRSEENCQSW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1D6</accession>
    <name>A0A452J1D6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Peptidase S1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J1D6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038732.1">
        <property type="protein sequence ID" value="ENSGAGP00000034206.1"/>
        <property type="gene ID" value="ENSGAGG00000024296.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00190">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.10.10">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009003">
        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043504">
        <property type="entry name" value="Peptidase_S1_PA_chymotrypsin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001254">
        <property type="entry name" value="Trypsin_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271:SF81">
        <property type="entry name" value="GRANZYME B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271">
        <property type="entry name" value="KALLIKREIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00089">
        <property type="entry name" value="Trypsin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00020">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50494">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50240">
        <property type="entry name" value="TRYPSIN_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0732">Signal</keyword>
    <keyword evidence="2" id="KW-0865">Zymogen</keyword>
    <feature type="domain" description="Peptidase S1" evidence="4">
        <location>
            <begin position="2"/>
            <end position="181"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023145"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034206.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="184" mass="21017" checksum="1E8356D714E87CFD" modified="2019-05-08" version="1">EIIRSREAKVHSRCYMAYLEIQHGKVTKKCGWFLVKPNLVLMDAHGKGEYITVTLGAHNISKPEWSQQKIPVRRQVPHLQYDKKTLNIVIMLLNTKTRVRPGSTCRVAGWDRTVIKNNMCELYPGYNRPMMLCGGSLGDSGDPLVCEGVVQGIASYVVNFYKPSVFTRLSKFIPWIQKILQNQH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1D8</accession>
    <name>A0A452J1D8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="19">Glutamate receptor</fullName>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="19">Receptor for glutamate that functions as a ligand-gated ion channel in the central nervous system and plays an important role in excitatory synaptic transmission. L-glutamate acts as an excitatory neurotransmitter at many synapses in the central nervous system.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>Ca(2+)(in) = Ca(2+)(out)</text>
            <dbReference type="Rhea" id="RHEA:29671"/>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="14 19">Postsynaptic cell membrane</location>
            <topology evidence="14 19">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="19">Belongs to the glutamate-gated ion channel (TC 1.A.10.1) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1D8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038644.1">
        <property type="protein sequence ID" value="ENSGAGP00000034127.1"/>
        <property type="gene ID" value="ENSGAGG00000024221.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015276">
        <property type="term" value="F:ligand-gated monoatomic ion channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0038023">
        <property type="term" value="F:signaling receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd06382">
        <property type="entry name" value="PBP1_iGluR_Kainate"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.2300:FF:000010">
        <property type="entry name" value="Glutamate ionotropic receptor kainate type subunit 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.190.10:FF:000240">
        <property type="entry name" value="Glutamate receptor ionotropic, kainate 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.70:FF:000010">
        <property type="entry name" value="Putative glutamate receptor ionotropic kainate 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.70">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.2300">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.190.10">
        <property type="entry name" value="Periplasmic binding protein-like II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001828">
        <property type="entry name" value="ANF_lig-bd_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019594">
        <property type="entry name" value="Glu/Gly-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001508">
        <property type="entry name" value="Iono_Glu_rcpt_met"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015683">
        <property type="entry name" value="Ionotropic_Glu_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001320">
        <property type="entry name" value="Iontro_rcpt_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028082">
        <property type="entry name" value="Peripla_BP_I"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18966">
        <property type="entry name" value="IONOTROPIC GLUTAMATE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01094">
        <property type="entry name" value="ANF_receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00060">
        <property type="entry name" value="Lig_chan"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10613">
        <property type="entry name" value="Lig_chan-Glu_bd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00177">
        <property type="entry name" value="NMDARECEPTOR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00079">
        <property type="entry name" value="PBPe"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53822">
        <property type="entry name" value="Periplasmic binding protein-like I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53850">
        <property type="entry name" value="Periplasmic binding protein-like II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 19" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="8 18" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13 19" id="KW-0407">Ion channel</keyword>
    <keyword evidence="6 19" id="KW-0406">Ion transport</keyword>
    <keyword evidence="12 19" id="KW-1071">Ligand-gated ion channel</keyword>
    <keyword evidence="7 19" id="KW-0472">Membrane</keyword>
    <keyword evidence="11 19" id="KW-0628">Postsynaptic cell membrane</keyword>
    <keyword evidence="9 19" id="KW-0675">Receptor</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 19" id="KW-0770">Synapse</keyword>
    <keyword evidence="3 19" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 19" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="1 19" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="497"/>
            <end position="516"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="573"/>
            <end position="595"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="754"/>
            <end position="778"/>
        </location>
    </feature>
    <feature type="domain" description="Ionotropic glutamate receptor C-terminal" evidence="20">
        <location>
            <begin position="385"/>
            <end position="735"/>
        </location>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="450"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="452"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="457"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="623"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="624"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="672"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="site" description="Crucial to convey clamshell closure to channel opening" evidence="17">
        <location>
            <position position="602"/>
        </location>
    </feature>
    <feature type="site" description="Interaction with the cone snail toxin Con-ikot-ikot" evidence="17">
        <location>
            <position position="629"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="18">
        <location>
            <begin position="80"/>
            <end position="331"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="18">
        <location>
            <begin position="684"/>
            <end position="738"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023257"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00034104"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00036634"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="RuleBase" id="RU367118"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="SMART" id="SM00079"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034127.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="839" mass="94863" checksum="A8ECAB3F43B49065" modified="2019-05-08" version="1">WTSFAPIKAPFLQVAPCGWTHYGGIFETVENEPVNVEELAFKFAVTNINRNKTLMPNTTLTYDIQRINLFDSFEASRRACDQLALGVAALFGPSHSSSVSAVQSICNALEVPHIQTRWKHPTVDNKDTFYINLYPDYAAISRAILDLVLYYNWKIVTVVYEDSTGLIRLQELIKAPSRYNIKIKIRQLPSGNKDARPLLKEMKKGKEFYVIFDCSHETAAEILKQILSMGMMTEYYHYFFTTLDLFALDLEPYRYSGVNMTGFRLLNIDNPQVTSVIEKWSMERLQAPPKPETGLLDGMMTTEAALMYDAVYMVAVASQRASQMTVSSLQCHRHKPWRFGPRFMNLIKEARWDGLTGHITFNKTDGLRKDFDLDIISLKEEGTEKIGVWNSNSGLNMTDSNKDRSTNITDSLANRTLIVTTILVRDREGAEVAGEWSGDREWGADLAVAPLTITYVREKVIDFSKPFMTLGISILYRKPNGTNPGVFSFLNPLSPDIWMYVLLACLGVSCVLFVIARFTPYEWYNPHPCNPDSDVVENNFTLLNSFWFGVGALMQQGSELMPKALSTRIVGGIWWFFTLIIISSYTANLAAFLTVERMESPIDSADDLAKQTKIEYGAVRDGSTMTFFKKSKISTYEKMWAFMSSRQQTALVKNNDEGIQRVLTTDYALLMESTSIEYVTQRNCNLTQIGGLIDSKGYGVGTPIGSPYRDKITIAILQLQEEGKLHMMKEKWWRGNGCPEEDSKEASALGVENIGGIFIVLAAGLVLSVFVAIGEFIYKSRKNSDIEQCLSFNAIMEELGISLRHQKKLKKRSRTKGRSSFTSILTCHQRRTLRKETMA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1D9</accession>
    <name>A0A452J1D9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Taste receptor type 1 member 2</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="17">Sweet taste receptor T1R2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">Putative taste receptor. TAS1R2/TAS1R3 recognizes diverse natural and synthetic sweeteners.</text>
    </comment>
    <comment type="subunit">
        <text evidence="15">Forms heterodimers with TAS1R3.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="14">Belongs to the G-protein coupled receptor 3 family. TAS1R subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1D9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038723.1">
        <property type="protein sequence ID" value="ENSGAGP00000034197.1"/>
        <property type="gene ID" value="ENSGAGG00000024281.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903767">
        <property type="term" value="C:sweet taste receptor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004930">
        <property type="term" value="F:G protein-coupled receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033041">
        <property type="term" value="F:sweet taste receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.2300:FF:000016">
        <property type="entry name" value="Taste 1 receptor member 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.50.30:FF:000004">
        <property type="entry name" value="Taste receptor type 1 member 3-like protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.2300">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.50.30">
        <property type="entry name" value="GPCR, family 3, nine cysteines domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001828">
        <property type="entry name" value="ANF_lig-bd_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000337">
        <property type="entry name" value="GPCR_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011500">
        <property type="entry name" value="GPCR_3_9-Cys_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038550">
        <property type="entry name" value="GPCR_3_9-Cys_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017978">
        <property type="entry name" value="GPCR_3_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000068">
        <property type="entry name" value="GPCR_3_Ca_sens_rcpt-rel"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017979">
        <property type="entry name" value="GPCR_3_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028082">
        <property type="entry name" value="Peripla_BP_I"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24061">
        <property type="entry name" value="CALCIUM-SENSING RECEPTOR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24061:SF517">
        <property type="entry name" value="TASTE RECEPTOR TYPE 1 MEMBER 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00003">
        <property type="entry name" value="7tm_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01094">
        <property type="entry name" value="ANF_receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07562">
        <property type="entry name" value="NCD3G"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00248">
        <property type="entry name" value="GPCRMGR"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53822">
        <property type="entry name" value="Periplasmic binding protein-like I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00980">
        <property type="entry name" value="G_PROTEIN_RECEP_F3_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50259">
        <property type="entry name" value="G_PROTEIN_RECEP_F3_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="8" id="KW-0297">G-protein coupled receptor</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9 18" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0675">Receptor</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0716">Sensory transduction</keyword>
    <keyword evidence="6 19" id="KW-0732">Signal</keyword>
    <keyword evidence="3" id="KW-0919">Taste</keyword>
    <keyword evidence="12" id="KW-0807">Transducer</keyword>
    <keyword evidence="5 18" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 18" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="19">
        <location>
            <begin position="1"/>
            <end position="26"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019588589" description="Taste receptor type 1 member 2" evidence="19">
        <location>
            <begin position="27"/>
            <end position="827"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="18">
        <location>
            <begin position="552"/>
            <end position="579"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="18">
        <location>
            <begin position="591"/>
            <end position="612"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="18">
        <location>
            <begin position="624"/>
            <end position="647"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="18">
        <location>
            <begin position="667"/>
            <end position="687"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="18">
        <location>
            <begin position="716"/>
            <end position="737"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="18">
        <location>
            <begin position="749"/>
            <end position="768"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="18">
        <location>
            <begin position="774"/>
            <end position="796"/>
        </location>
    </feature>
    <feature type="domain" description="G-protein coupled receptors family 3 profile" evidence="20">
        <location>
            <begin position="553"/>
            <end position="819"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022480"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022606"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023224"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00037659"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00038492"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00038699"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00040704"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00042616"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50259"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034197.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="827" mass="93417" checksum="C46D4FFF70A6580B" modified="2019-05-08" version="1">HHRTMRSTGGIAMCVSVLLTMAGSQTQYSDFHLAGDYLLGGLFSLHTETKSSPHLSYSAVHLRAMRFAVEQINNSSTLLPGVSLGYEMVDVCYLTNFIHPILYFLANKHSVVQLHNNYTHYQPRVLAVIGPDSSQAVVIAAQVLSLFLVPQISYTATTNELSDRKRFPTAFQTISSMEQQIVAMQLLLKEFRWNWINILYSDDDYGRQNLKLLRARATEICVALQEVIPVPQSSQMLSLELQGKIKSVVRKVIHSTAKVVIVLSLELILPFFFQEAVRQNVTDLVWIATEAWATEPALYNITDLWRLGTILGVAVKEFSIPGLDDFRVSLAEGANEQASDQTTCNQECDQCLRAAQDAEHSMRAQGERTDFSVYSSVYIVAHALHRLLGCNTTGCHKRMVYPWQVGHMLALRNPSHVLRALKKKQKQNNWGFVPLLSRGLDNIYDWLRFYYGTFRSSSSSLGSCYEIVIVFLQFPFSQVPISVCSQQCEYGQRRKPVTVSSCCFECIDCEEGTFLNNNDLYNCQPCPADQWSNARSQKCYPRLLKYLEWQEGIAVVLLLFSVLGLVATVVISVIFAIHLHTPVVKSAGGRMCFLMLASLIIGFINVPVYMGIPTEFKCMCRQIVYTLCFTVCISCLTVRSFQIISIFKMAARLPKAYSYWMKYNGQYIFITVILALKVTIVVTNVIIHPSIPVPLVLEDDPSTVELQCNKSFKLSLLMNNSLDTFLSILCFYFSYIGKELPKNYNEAKYITLCMTCYAVSEIAIFLAMSVYRGMLVTICDVIGVVINLLGISIGYFGPKCYLIFFHPERNTAAYFQSVIQSYTMRQD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1E0</accession>
    <name>A0A452J1E0_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J1E0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038738.1">
        <property type="protein sequence ID" value="ENSGAGP00000034211.1"/>
        <property type="gene ID" value="ENSGAGG00000024299.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024845">
        <property type="entry name" value="NHS-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23039">
        <property type="entry name" value="NANCE-HORAN SYNDROME PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23039:SF6">
        <property type="entry name" value="SIMILAR TO MKIAA1522 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15273">
        <property type="entry name" value="NHS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="65"/>
            <end position="233"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="349"/>
            <end position="413"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="448"/>
            <end position="1111"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1160"/>
            <end position="1260"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="68"/>
            <end position="84"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="85"/>
            <end position="94"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="103"/>
            <end position="113"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="134"/>
            <end position="146"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="164"/>
            <end position="175"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="1">
        <location>
            <begin position="349"/>
            <end position="358"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="1">
        <location>
            <begin position="367"/>
            <end position="395"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="1">
        <location>
            <begin position="482"/>
            <end position="500"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="552"/>
            <end position="561"/>
        </location>
    </feature>
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        <location>
            <begin position="562"/>
            <end position="575"/>
        </location>
    </feature>
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        <location>
            <begin position="601"/>
            <end position="610"/>
        </location>
    </feature>
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        <location>
            <begin position="618"/>
            <end position="631"/>
        </location>
    </feature>
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        <location>
            <begin position="659"/>
            <end position="679"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="1">
        <location>
            <begin position="723"/>
            <end position="754"/>
        </location>
    </feature>
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        <location>
            <begin position="755"/>
            <end position="772"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="1">
        <location>
            <begin position="794"/>
            <end position="803"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="874"/>
            <end position="883"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="1">
        <location>
            <begin position="884"/>
            <end position="895"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="1">
        <location>
            <begin position="924"/>
            <end position="939"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="1">
        <location>
            <begin position="1018"/>
            <end position="1028"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="1035"/>
            <end position="1067"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="1160"/>
            <end position="1175"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="1216"/>
            <end position="1230"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034211.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1260" mass="132307" checksum="34783B7D10105801" modified="2019-05-08" version="1">MVVFLGRNLPSLLTLFKKKGSAKAENKRLNVQYKAGEERPDNVFFPSSRPPHLEELHNQAQAGLKSLQHQEKQKQTRSGWDHGDSNSIQSSHTSAADDEISFRSRTQSCTTENASEDALSIRSEMIQRKGSTFRPHDSFPKSSEKAGKRRKERRTTVLGIPQHVQKELGLRNSHELRKRPGSVSPQDGPIQAGSPHSLAPPTLVNGGEADGGMIHIPTINGNLQPPPAPKGGARVSLQALEEAESDAAIQRHINCVYYDDTLLGRRTAAKLSPLLRPKSLAVPGMTTNPSPPPELFGPVMSISPQATYMSKIIPNAVLPPMVDVIALSRSSVQTLSRCSLVSSSPASVRSLTRFSEPSARSREHSSSSDNWSHSQSTETIVSDSSTISSQGGSNSRKGEAGPPSEADTAGRSDTDQLSVYSAASCTNSCSQARPVYTAHGLLAVGPGGSSGRASPAYSTSSMADGSDTVSVRSDRSSTRSVSLRKMKKPPAPPRRTHSLHQKAQQHQQGVGEGGQKVLGLPPKPDPRPQREPWTPRLESQSPLGEDEVFSPCSLSETSSIRSDSLAYSADASSPDVSRCSPVPGEKLRREVEGVAVIFREQQATPRQSSPEGFKRTMSPSSGYSSQSGTPTLPTKGLGPHASSPGKRRPQPKKPERVCSLQSPALSISSSLTSLSSSTSDPAPAETVAPPAGPALSWQSRMDRFIIPPHPKVLAPISPPPIKPQQAAPSASPTVSSPAPSTASQKPSALPKPSSRSPPPSPPPPYHPPPPPVKKAGTSPEAPHSETIQEAPQDPSWPPPPPPALDEQDLSMADFPPPDEAGFSTLPELLPVTGTAGASSVAAQPEPVGSEQPARPKAAPRGDGPGAASLAISAKVSSRTQQQQGPLAGSALPAPAVELPMPTDMSPSARATPDLQPGQANPAVPAAPPHAPAAPAPPTALQPQASLKKTTNGSQLDPKKELVSRSKSSPTPKEDANLPIVTPSLLQMVRLRSVNMDGHAPAGPGKQSPGQNGQGADGVGKQPQQVPQKPIRKSLSLRQSPSSKDTVPLNQLQNAVRLKTSTLSSRDAPTSRLAGWTNGNKLTLPGHTASSSEHTTGDAKDGQVSPIHKSPASTASFIFAKSSKKLVIETPSSPEAQADLKRNLVAELMNFAGQRSTVPAMTQQGQVPPGKSQAQRKSSKIPPPIARKPLLGMGQPQSPVSPKPQGEEVLNCSLPDSKAKASSAEENRTKSELSVNAAPQADSSRAGHLAGSATPSQNPPA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1E4</accession>
    <name>A0A452J1E4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Ral GTPase-activating protein subunit beta</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="6">p170</fullName>
        </alternativeName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="3">Non-catalytic subunit of the heterodimeric RalGAP1 and RalGAP2 complexes which act as GTPase activators for the Ras-like small GTPases RALA and RALB.</text>
    </comment>
    <comment type="subunit">
        <text evidence="4">Component of the heterodimeric RalGAP1 complex with RALGAPA1 and of the heterodimeric RalGAP2 complex with RALGAPA2. Heterodimerization is required for activity.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038649.1">
        <property type="protein sequence ID" value="ENSGAGP00000034132.1"/>
        <property type="gene ID" value="ENSGAGG00000024251.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051056">
        <property type="term" value="P:regulation of small GTPase mediated signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.11210:FF:000005">
        <property type="entry name" value="Ral GTPase-activating protein, beta subunit (non-catalytic)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.11210">
        <property type="entry name" value="Rap/Ran-GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039930">
        <property type="entry name" value="RALGAPB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035974">
        <property type="entry name" value="Rap/Ran-GAP_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000331">
        <property type="entry name" value="Rap/Ran_GAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046859">
        <property type="entry name" value="RGPA/RALGAPB_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21344">
        <property type="entry name" value="RAL GTPASE-ACTIVATING PROTEIN SUBUNIT BETA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21344:SF1">
        <property type="entry name" value="RAL GTPASE-ACTIVATING PROTEIN SUBUNIT BETA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20412">
        <property type="entry name" value="RALGAPB_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF111347">
        <property type="entry name" value="Rap/Ran-GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50085">
        <property type="entry name" value="RAPGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Rap-GAP" evidence="8">
        <location>
            <begin position="1168"/>
            <end position="1412"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="364"/>
            <end position="454"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="725"/>
            <end position="757"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1093"/>
            <end position="1119"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1333"/>
            <end position="1355"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="385"/>
            <end position="397"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="408"/>
            <end position="427"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="428"/>
            <end position="444"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="738"/>
            <end position="754"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="1333"/>
            <end position="1343"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00053327"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00063020"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00072207"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00081608"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50085"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034132.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1514" mass="168724" checksum="7F425C5DEE6F2084" modified="2019-05-08" version="1">MYSEWRSLHLVVQNDQGHTSVLHSYPDNVGREVANAVVRPLGQALNTPSAAGSESLLKTDKEVKWTMEVVCYGLTLPLEGDTVKYCVDVYTDWIMALVLPKDSIPLPVIKEPNLYVQSILKHLQNLFVPRQDPGSSQNRLCLQVLKAVQKLARESTIMARETWEVLLLFLLQINDTLLAAPTVQGGIAENLAEKLIGVLFEVWLLACTRCFPTPPYWKTAKEMVANWRHHPAVVEQWSKVICALTSRLLRFIYGPSFPPFKIPDEDASLIPPEMDNECIAQTWFRFLHMLSNPVDLSNPAIISSTPKFQEQFLNVSGMTQELNQYPCLKHLPHIFFRAMRGISCLVDAFLGVAAFTNSGCERLPRYGISRPRSDSAPPTPVNRLSMPQNTAINTTPPHNRRHRAVTVNKATMKTSTVSTVHASKVQHQPSSSTSPLSSPNQTSSEPRPLPAPHRPKVNSILNLFGSWLFDAGFVHCKLHNGINRDSSMTAIATQASMEFRRKGSQMSTDTMTSNSTFDSSDFPDNYEAGRAEACGTLCRIFCSKKTGEDILPAYLSRFYMLLIQGLQIADFICHPVLASIILNSPPLFCCDLKGIDVVVPYFISALETILPDRELSKFKTYVNPTDLRRSSINILLSLLPLPHHFGTIKSEVVLEGKFSNDDGSSSYDKPITFLSLKLRLVNILIGALQTETDPNNTQMILGAMLNIVQDSALLEAIGCQMETNGGENNLLKNHRRTNSSISTASGGSTEPTTPDSERPAQALLRDYALNTDTAAGLLIRSIHLVTQRLNSQWRQDMSISLAALELLSGLAKVKVIVDSADRKRAVSSVCSYIVYQCSRPAPLHSRDLHSMIVAAFQCLCVWLTEHPDMMDEKDCLKEVLEIVELGISGSKSKNSEQEVKYKGDKEPNPASMRVKDAAEATLTCIMQLLGAFPSPSGPASPCSLVNETTLIKYSRLPTINKHSFRYFVLDNSVILAMLEQPLGNEQNDSFPSVTVLIRGVSGRLAWAQQLCLLPRGAKANQKMFLPEPRPAPKNDVGLKYSVKHRPFPEEVDKIPFVKADLSIPDLHEIVNEELEERHEKLRNGMAQQITYETSVDQQSEEEWRKKSFPDPVTECKPPPPAQEFQTARLFLSHFGFLSLEALKETANSRLPPHLIALDSALPGFFDDLGYLDLLPCRPFDTVFIFYMKAGQKTSQEILKNVESSRNVQPHFLEFLLSLGWAVDVGKHPGWTGHVSTSWSINSCSDEEGSEQDEIVSSEDTGASIFNGQKKVLYYADALTEIAFVVPSPVESLTDSLESNVSDQESDSNMDLLPGILKQPSLTLELFPNHTDNLNSLQRLSPTSRSKKLPQGRTIPPMGPETKVSVVWVERYDDIENFPLSDLVSETSTGVETTSNSSTSLRSTTPEREVPVVFIHPLNTGLFRIKVQGATGKFNMVIPLVDGMIVSRRALGFLVRQTIINICRRKRLESDSYNPPHVRRKQKIADIVNKYRNKQLEPEFYTSLFQEVGLKNCNP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1E7</accession>
    <name>A0A452J1E7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Taste receptor type 1 member 2</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="17">Sweet taste receptor T1R2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">Putative taste receptor. TAS1R2/TAS1R3 recognizes diverse natural and synthetic sweeteners.</text>
    </comment>
    <comment type="subunit">
        <text evidence="15">Forms heterodimers with TAS1R3.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="14">Belongs to the G-protein coupled receptor 3 family. TAS1R subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1E7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038734.1">
        <property type="protein sequence ID" value="ENSGAGP00000034207.1"/>
        <property type="gene ID" value="ENSGAGG00000024281.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903767">
        <property type="term" value="C:sweet taste receptor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004930">
        <property type="term" value="F:G protein-coupled receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033041">
        <property type="term" value="F:sweet taste receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.2300:FF:000016">
        <property type="entry name" value="Taste 1 receptor member 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.50.30:FF:000004">
        <property type="entry name" value="Taste receptor type 1 member 3-like protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.2300">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.50.30">
        <property type="entry name" value="GPCR, family 3, nine cysteines domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001828">
        <property type="entry name" value="ANF_lig-bd_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000337">
        <property type="entry name" value="GPCR_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011500">
        <property type="entry name" value="GPCR_3_9-Cys_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038550">
        <property type="entry name" value="GPCR_3_9-Cys_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017978">
        <property type="entry name" value="GPCR_3_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000068">
        <property type="entry name" value="GPCR_3_Ca_sens_rcpt-rel"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017979">
        <property type="entry name" value="GPCR_3_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028082">
        <property type="entry name" value="Peripla_BP_I"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24061">
        <property type="entry name" value="CALCIUM-SENSING RECEPTOR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24061:SF517">
        <property type="entry name" value="TASTE RECEPTOR TYPE 1 MEMBER 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00003">
        <property type="entry name" value="7tm_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01094">
        <property type="entry name" value="ANF_receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07562">
        <property type="entry name" value="NCD3G"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00248">
        <property type="entry name" value="GPCRMGR"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53822">
        <property type="entry name" value="Periplasmic binding protein-like I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00980">
        <property type="entry name" value="G_PROTEIN_RECEP_F3_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50259">
        <property type="entry name" value="G_PROTEIN_RECEP_F3_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="8" id="KW-0297">G-protein coupled receptor</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9 18" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0675">Receptor</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0716">Sensory transduction</keyword>
    <keyword evidence="6 19" id="KW-0732">Signal</keyword>
    <keyword evidence="3" id="KW-0919">Taste</keyword>
    <keyword evidence="12" id="KW-0807">Transducer</keyword>
    <keyword evidence="5 18" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 18" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="19">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019370043" description="Taste receptor type 1 member 2" evidence="19">
        <location>
            <begin position="23"/>
            <end position="804"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="18">
        <location>
            <begin position="529"/>
            <end position="556"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="18">
        <location>
            <begin position="568"/>
            <end position="589"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="18">
        <location>
            <begin position="601"/>
            <end position="624"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="18">
        <location>
            <begin position="644"/>
            <end position="664"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="18">
        <location>
            <begin position="693"/>
            <end position="714"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="18">
        <location>
            <begin position="726"/>
            <end position="745"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="18">
        <location>
            <begin position="751"/>
            <end position="773"/>
        </location>
    </feature>
    <feature type="domain" description="G-protein coupled receptors family 3 profile" evidence="20">
        <location>
            <begin position="530"/>
            <end position="796"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022480"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022606"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023224"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00037659"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00038492"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00038699"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00040704"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00042616"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50259"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034207.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="804" mass="90806" checksum="CB444C50E56BC53E" modified="2019-05-08" version="1">MRSTGGIAMCVSVLLTMAGSQTQYSDFHLAGDYLLGGLFSLHTETKSSPHLRNGCRWASYSHLRAMRFAVEQINNSSTLLPGVSLGYEMVDVCYLTNFIHPILYFLANKHSVVQLHNNYTHYQPRVLAVIGPDSSQAVVIAAQVLSLFLVPQISYTATTNELSDRKRFPTAFQTISSMEQQIVAMQLLLKEFRWNWINILYSDDDYGRQNLKLLRARATEICVALQEVIPVPQSSQMLSLELQGKIKSVVRKVIHSTAKVVIVLSLELILPFFFQEAVRQNVTDLVWIATEAWATEPALYNITDLWRLGTILGVAVKEFSIPGLDDFRVSLAEGANEQASDQTTCNQECDQCLRAAQDAEHSMRAQGERTDFSVYSSVYIVAHALHRLLGCNTTGCHKRMVYPWQVGHMLALRNPSHVLRAWDYMAWGIGRMLVDRNKKLLFTIQNISWHTPNNTVPISVCSQQCEYGQRRKPVTVSSCCFECIDCEEGTFLNNNDLYNCQPCPADQWSNARSQKCYPRLLKYLEWQEGIAVVLLLFSVLGLVATVVISVIFAIHLHTPVVKSAGGRMCFLMLASLIIGFINVPVYMGIPTEFKCMCRQIVYTLCFTVCISCLTVRSFQIISIFKMAARLPKAYSYWMKYNGQYIFITVILALKVTIVVTNVIIHPSIPVPLVLEDDPSTVELQCNKSFKLSLLMNNSLDTFLSILCFYFSYIGKELPKNYNEAKYITLCMTCYAVSEIAIFLAMSVYRGMLVTICDVIGVVINLLGISIGYFGPKCYLIFFHPERNTAAYFQSVIQSYTMRQD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1H2</accession>
    <name>A0A452J1H2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">Integrin alpha first immunoglubulin-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 16">Membrane</location>
            <topology evidence="1 16">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 16">Belongs to the integrin alpha chain family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1H2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038773.1">
        <property type="protein sequence ID" value="ENSGAGP00000034246.1"/>
        <property type="gene ID" value="ENSGAGG00000024233.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009897">
        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008305">
        <property type="term" value="C:integrin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033627">
        <property type="term" value="P:cell adhesion mediated by integrin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098609">
        <property type="term" value="P:cell-cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007160">
        <property type="term" value="P:cell-matrix adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007229">
        <property type="term" value="P:integrin-mediated signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.130:FF:000006">
        <property type="entry name" value="Integrin, alpha 9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.130">
        <property type="entry name" value="Integrin alpha, N-terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1460">
        <property type="entry name" value="Integrin domains. Chain A, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013517">
        <property type="entry name" value="FG-GAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013519">
        <property type="entry name" value="Int_alpha_beta-p"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000413">
        <property type="entry name" value="Integrin_alpha"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013649">
        <property type="entry name" value="Integrin_alpha_Ig-like_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028994">
        <property type="entry name" value="Integrin_alpha_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032695">
        <property type="entry name" value="Integrin_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23220">
        <property type="entry name" value="INTEGRIN ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23220:SF78">
        <property type="entry name" value="INTEGRIN ALPHA-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01839">
        <property type="entry name" value="FG-GAP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08441">
        <property type="entry name" value="Integrin_A_Ig_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01185">
        <property type="entry name" value="INTEGRINA"/>
    </dbReference>
    <dbReference type="SMART" id="SM00191">
        <property type="entry name" value="Int_alpha"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69318">
        <property type="entry name" value="Integrin alpha N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69179">
        <property type="entry name" value="Integrin domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51470">
        <property type="entry name" value="FG_GAP"/>
        <property type="match status" value="5"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0106">Calcium</keyword>
    <keyword evidence="8 16" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="14" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10 16" id="KW-0401">Integrin</keyword>
    <keyword evidence="11" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13 16" id="KW-0675">Receptor</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5 16" id="KW-0732">Signal</keyword>
    <keyword evidence="3" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="16">
        <location>
            <begin position="1"/>
            <end position="31"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5018812272" description="Integrin alpha first immunoglubulin-like domain-containing protein" evidence="16">
        <location>
            <begin position="32"/>
            <end position="570"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="37"/>
            <end position="98"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="185"/>
            <end position="236"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="291"/>
            <end position="350"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="351"/>
            <end position="407"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="411"/>
            <end position="473"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha first immunoglubulin-like" evidence="17">
        <location>
            <begin position="458"/>
            <end position="504"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023037"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00803"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="RuleBase" id="RU003762"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF08441"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034246.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="570" mass="62781" checksum="2C0A521A5C0C8DE6" modified="2019-05-08" version="1">SPGPPAPRGAPVWAALLLLLLLGRCMRAAHAYNLDTQRPVVFKGLSGTLFGYSVVLHSHGPSRWLVVGAPKANWLANSSVVNPGAIFRCRIGSNPSGTCEQLQMGELNAGDYCGKTCFEERDNQWLGVSLSRQPRENGTIVACGHRWKNIYYIKNEHKLPNGVCYEIPPDFRTQLSKRISPCYKDYVRKFGENHGSCQAGMSSFYFKDLIIMGAPGSYYWTGSVFVYNTTANTFQAYTDSRNQVKFGSYLGYAVGAGHFLTPASTEVIGGAPQQEQTGKAYIFSIGTRALTILSELKGKKLGSYFGAAVCAVDLNTDGLSDLLVGAPMQSKVREEGRVYVYINSGSVKLEIELAGSDSYAARFGESIANLGDIDNDGFEDVAIGAPQENNLEGAIYIYNGRKDGISSSFSQRIQGHRISNTLSMFGQSISSGIDADNNGYPDVAVGAFLDDSAVLLRTRPVVIVEAFLKHPNAVNRTNLECVENGQPAVCVNLMLCFTYKGRDVPGYVGKLLDKYFVLNTWISFYAVKYFTIWTINKYCVCKVVHQPEHSRYHYEKSLKSYDSRIKENKQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1H3</accession>
    <name>A0A452J1H3_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="13">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Golgi apparatus membrane</location>
            <topology evidence="1">Single-pass type II membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the glycosyltransferase 29 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1H3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038681.1">
        <property type="protein sequence ID" value="ENSGAGP00000034158.1"/>
        <property type="gene ID" value="ENSGAGG00000024270.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000139">
        <property type="term" value="C:Golgi membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003828">
        <property type="term" value="F:alpha-N-acetylneuraminate alpha-2,8-sialyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006491">
        <property type="term" value="P:N-glycan processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009311">
        <property type="term" value="P:oligosaccharide metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd23987">
        <property type="entry name" value="GT29_ST8SIA2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1480.20:FF:000001">
        <property type="entry name" value="ST8 alpha-N-acetyl-neuraminide alpha-2,8-sialyltransferase 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1480.20">
        <property type="entry name" value="Glycosyl transferase family 29"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001675">
        <property type="entry name" value="Glyco_trans_29"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050943">
        <property type="entry name" value="Glycosyltr_29_Sialyltrsf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038578">
        <property type="entry name" value="GT29-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012163">
        <property type="entry name" value="Sialyl_trans"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11987">
        <property type="entry name" value="ALPHA-2,8-SIALYLTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11987:SF30">
        <property type="entry name" value="ALPHA-2,8-SIALYLTRANSFERASE 8B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00777">
        <property type="entry name" value="Glyco_transf_29"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF005557">
        <property type="entry name" value="Sialyl_trans"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="3" id="KW-0328">Glycosyltransferase</keyword>
    <keyword evidence="8" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="9" id="KW-0472">Membrane</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0735">Signal-anchor</keyword>
    <keyword evidence="4" id="KW-0808">Transferase</keyword>
    <keyword evidence="5" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7" id="KW-1133">Transmembrane helix</keyword>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="129"/>
            <end position="279"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004323"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022676"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PIRSR" id="PIRSR005557-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034158.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="347" mass="39476" checksum="86E708C03935BCE0" modified="2019-05-08" version="1">FFFAGWNSGGRGTIRSSVNSLHSKSNRAEVVINGSSSPAVVDRSNESIKHIKPASFRWRHNQTLSLKIRKQILKFLDAEKDISVLKGTLKPGDVIHYIFDRDSTMNVSQNLYELLPRTSPLKGKNFRTCGIVGNSGILLNSGCGREIDAHSFVIRCNLAPVQEYAWDVGMKTDLVTMNPSVIQRAFEDLVNETWREKLLQHLHSLNGSILWIPAFMAKGGKERVEWVNELILKHRINVRTAYPSLRLLHAVRGYWLTNKVHIKRPTTGLLMYTLATRFCNRIYLYGFWPFPRDQNQNPVKYHYYDSLKYGYTSQASPHTMPLEFKALKTLHQQGALKLTVGECGGAT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1H7</accession>
    <name>A0A452J1H7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Sodium/calcium exchanger membrane region domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>Ca(2+)(out) + K(+)(out) + 4 Na(+)(in) = Ca(2+)(in) + K(+)(in) + 4 Na(+)(out)</text>
            <dbReference type="Rhea" id="RHEA:69967"/>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
            <dbReference type="ChEBI" id="CHEBI:29103"/>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the Ca(2+):cation antiporter (CaCA) (TC 2.A.19) family. SLC24A subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J1H7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038784.1">
        <property type="protein sequence ID" value="ENSGAGP00000034251.1"/>
        <property type="gene ID" value="ENSGAGG00000024329.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005262">
        <property type="term" value="F:calcium channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008273">
        <property type="term" value="F:calcium, potassium:sodium antiporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015293">
        <property type="term" value="F:symporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098703">
        <property type="term" value="P:calcium ion import across plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006874">
        <property type="term" value="P:intracellular calcium ion homeostasis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060292">
        <property type="term" value="P:long-term synaptic depression"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060291">
        <property type="term" value="P:long-term synaptic potentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1420.30:FF:000002">
        <property type="entry name" value="Sodium/potassium/calcium exchanger 2 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1420.30:FF:000004">
        <property type="entry name" value="Sodium/potassium/calcium exchanger 2 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1420.30">
        <property type="entry name" value="NCX, central ion-binding region"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004481">
        <property type="entry name" value="K/Na/Ca-exchanger"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004837">
        <property type="entry name" value="NaCa_Exmemb"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044880">
        <property type="entry name" value="NCX_ion-bd_dom_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00367">
        <property type="entry name" value="calcium/sodium antiporter"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10846">
        <property type="entry name" value="SODIUM/POTASSIUM/CALCIUM EXCHANGER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10846:SF41">
        <property type="entry name" value="SODIUM_POTASSIUM_CALCIUM EXCHANGER 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01699">
        <property type="entry name" value="Na_Ca_ex"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0050">Antiport</keyword>
    <keyword evidence="9" id="KW-0106">Calcium</keyword>
    <keyword evidence="6" id="KW-0109">Calcium transport</keyword>
    <keyword evidence="15" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13" id="KW-0406">Ion transport</keyword>
    <keyword evidence="14 19" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-0630">Potassium</keyword>
    <keyword evidence="5" id="KW-0633">Potassium transport</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="16" id="KW-0915">Sodium</keyword>
    <keyword evidence="16" id="KW-0739">Sodium transport</keyword>
    <keyword evidence="10" id="KW-0769">Symport</keyword>
    <keyword evidence="7 19" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="12 19" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="36"/>
            <end position="60"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="136"/>
            <end position="155"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="176"/>
            <end position="200"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="206"/>
            <end position="228"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="240"/>
            <end position="259"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="453"/>
            <end position="469"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="481"/>
            <end position="504"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="516"/>
            <end position="538"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="583"/>
            <end position="606"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="613"/>
            <end position="632"/>
        </location>
    </feature>
    <feature type="domain" description="Sodium/calcium exchanger membrane region" evidence="20">
        <location>
            <begin position="141"/>
            <end position="282"/>
        </location>
    </feature>
    <feature type="domain" description="Sodium/calcium exchanger membrane region" evidence="20">
        <location>
            <begin position="482"/>
            <end position="630"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="307"/>
            <end position="336"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="410"/>
            <end position="440"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="323"/>
            <end position="336"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="18">
        <location>
            <begin position="430"/>
            <end position="439"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005364"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022449"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022538"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022568"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022847"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022958"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023201"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00033627"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="Pfam" id="PF01699"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034251.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="643" mass="72027" checksum="E836FA9F9893C2DC" modified="2019-05-08" version="1">MALSKNIAASILEEWCLEEPLFRRRRHQNVRKKLRLIRIIGLFMSIMAISTFSLSISAFFKMEPHSTEMTSNLGSQKSMHGHQRTLLDVKEQNQNGTPDSPISMKYETEHNNATEEHSKGEYPKDLFSLAERRRGAVILHIMGMIYMFIALAIVCDEFFVPSLTVITEKLAISDDVAGATFMAAGGSAPELFTSLIGVFISHSNVGIGTIVGSAVFNILFVIGMCALFSREILNLTWWPLFRDVSFYIMDLILLIIFFLDNLIMWWESLTLLIAYFCYVTFMKFNVQVEECVKKILNRNKVEKVTTADSEQKPANAGYKDDQTLTTKPRLQRGGSSASLHNSLMRNSIFQLMIHTLDPLAEGKFKEKASILHKIAKKKCQVEDSERWNGAANHAEKIELPNTTKVEVEVTPPNDGSGPVQNGNIAHGIEENSEDNDEEQPLSLAWPDTLRKQVMYLAVFPIVFPLWITLPDVRKPSSRKFFPVTFFGAITWIAVFSYLMVWWAHQVGETIGISEEIMGLTILAAGTSIPDLITSVIVARKGLGDMAVSSSVGSNIFDITVGLPLPWLLYAAINGFAPVTVSSNGLFCAIVLLFIMLLFVILSIALCKWRMNKFLGFTMFGLYFIFLIVSVLLEDKVILCPVSI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J1I8</accession>
    <name>A0A452J1I8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Peptidase S1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J1I8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038795.1">
        <property type="protein sequence ID" value="ENSGAGP00000034261.1"/>
        <property type="gene ID" value="ENSGAGG00000024324.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00190">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.10:FF:000014">
        <property type="entry name" value="Complement factor D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.10:FF:000005">
        <property type="entry name" value="Serine protease 37"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.10.10">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009003">
        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043504">
        <property type="entry name" value="Peptidase_S1_PA_chymotrypsin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001314">
        <property type="entry name" value="Peptidase_S1A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001254">
        <property type="entry name" value="Trypsin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018114">
        <property type="entry name" value="TRYPSIN_HIS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033116">
        <property type="entry name" value="TRYPSIN_SER"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271:SF81">
        <property type="entry name" value="GRANZYME B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271">
        <property type="entry name" value="KALLIKREIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00089">
        <property type="entry name" value="Trypsin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00722">
        <property type="entry name" value="CHYMOTRYPSIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00020">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50494">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50240">
        <property type="entry name" value="TRYPSIN_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00134">
        <property type="entry name" value="TRYPSIN_HIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00135">
        <property type="entry name" value="TRYPSIN_SER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3 7" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="1 7" id="KW-0645">Protease</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 7" id="KW-0720">Serine protease</keyword>
    <keyword evidence="2" id="KW-0732">Signal</keyword>
    <keyword evidence="5" id="KW-0865">Zymogen</keyword>
    <feature type="domain" description="Peptidase S1" evidence="8">
        <location>
            <begin position="21"/>
            <end position="248"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023145"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU363034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034261.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="267" mass="29396" checksum="04903308F009A4DD" modified="2019-05-08" version="1">IVPPALIPHLISSLPGVAEEIIGGRVAKPHSCPYMAYLKRKPCKGEETCGGLLVREDFVLTAAHCADGDITVLLGAHNVRRDEETQQQVSVRRKIPHPRYNETSFENDLMLLQLAELAELTDAIYTIPLPQAGRTVDPGSVCSVAGWGRTSLNSMRTSNILHEVDLEVMSDKTCQEDGLLQHYYKPLKMMCVGDPAEDKASFSGDSGGPLVCDRTAQGIVSFGKRNGSPPRVFTRVSGYIPWIKKTMRDLLIGRDRGPAPQRYLPSW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIR7</accession>
    <name>A0A452GIR7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">SCAN box domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000001707.1">
        <property type="protein sequence ID" value="ENSGAGP00000001497.1"/>
        <property type="gene ID" value="ENSGAGG00000000400.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019901">
        <property type="term" value="F:protein kinase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008104">
        <property type="term" value="P:intracellular protein localization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.200:FF:000010">
        <property type="entry name" value="neurobeachin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.200">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.4020.10">
        <property type="entry name" value="DNA breaking-rejoining enzymes"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050865">
        <property type="entry name" value="BEACH_Domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031570">
        <property type="entry name" value="NBEA/BDCP_DUF4704"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046852">
        <property type="entry name" value="Neurobeachin_a-sol"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003309">
        <property type="entry name" value="SCAN_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038269">
        <property type="entry name" value="SCAN_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13743">
        <property type="entry name" value="BEIGE/BEACH-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13743:SF62">
        <property type="entry name" value="NEUROBEACHIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15787">
        <property type="entry name" value="DUF4704"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13385">
        <property type="entry name" value="Laminin_G_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20425">
        <property type="entry name" value="Neurobeachin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02023">
        <property type="entry name" value="SCAN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00431">
        <property type="entry name" value="SCAN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47353">
        <property type="entry name" value="Retrovirus capsid dimerization domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50804">
        <property type="entry name" value="SCAN_BOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0853">WD repeat</keyword>
    <feature type="domain" description="SCAN box" evidence="3">
        <location>
            <begin position="1674"/>
            <end position="1749"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1"/>
            <end position="21"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="963"/>
            <end position="986"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="972"/>
            <end position="985"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022574"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS50804"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001497.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1758" mass="196662" checksum="72E01150CB16B372" modified="2019-05-08" version="1">MASEKPVSGPDPQPAGLIPVGAGGGGGGGSSAAVMGELRASGSGAVVLPAGMINPSVPIRNIRMKFAVLIGLIQVGEVSNRDIVETVLNLLVGGEFDLEMNFIIQDAESITCMSELLEHCDVTCQAEIWSMFTAILRKSVRNLQTSTEVGLIEQVLLKMSTVDDMIADLLVDMLGVLASYSITVKELKLLFSMLRGENGIWPRHAIKLLSVLNQMPQRHGPDTFFNFPGCSAAAIALPPIAKWPYQNGFTLNTWFRMDPLNNINVDKDKPYLYCFRTSKGVGYSAHFVGNCLIVTSLKSKGKGFQHCVKYDFQPRKWYMISIVHIYNRWRNSEIRCYVNGQLVSYGDMAWHVNTNDSYDKCFLGSSETADANRVFCGQLGAVYVFTEALNPAQIFAIHQLGPGYKSTFKFKSESDIHLAEHHKQVLYDGKLASSIAFTYNAKATDAQLCLESSPKENPSIFVHSPHALMLQDVKAIVTHSIHSAIHSIGGIQVLFPLFAQLDNRQLNDSQMETTVCATLLAFLVELLKSSVAMQEQMLGGKGFLVIGYLLEKSSRVHITRAVLEQFLSFAKYLDGLSHGAPLLKQLCDHILFNPAIWIHTPAKVQLSLYTYLSAEFIGTATIYTTIRRVGTVLQLMHTLKYYYWVVNPADSSGITPKGLEGPRPSQKEIISLRAFMLLFLKQLILKDRGVKEDELQSILNYLLTMHEDENIHDVLQLLVALMSEHPASMIPAFDQRNGIRVIYKLLASKSESIWVQALKVLGYFLKHLGHKRKVEIMHTHSLFTLLGERLMLHTNTVTITTYNTLYEILTEQVCTQVVHKPHPEPDSTVKIQNPMILKVVATLLKNSTPSAELMEVRRLFLSDMIKLFSNSRENRRCLLQCSVWQDWMFSLGYINPKNSEEQKITEMVYNIFRILLYHAIKYEWGGWRVWVDTLSIAHSKVTYEAHKEYLAKMYEEYQRQEEENIKKGKKGNVSTISGLSSQTTGAKGGLEIREIEDLSQSQSPESETDYPVSTDTRDLLMATKVSDEVLGSSDRPGGGVHVEVHDLLVDIKAEKVEATEAVQGRSVTQQDRDLRVDLGFRGMPMTEEQRRQFSPGPRTTMFRIPEFKWSPMHQRLLTDLLFALETDVHVWRSHSTKSVMDFVNSNENIIFVHNTIHLISQMVDNIIIACGGILPLLSAATSPTTELENIEVTQGMSAETAVTFLSRLMAMVDVLVFASSLNFSEIEAEKNMSSGGLMRQCLRLVCCVAVRNCLECRQRQRERVNKSSLISSKTQETLQGVTATSPLENVPGNLSPIKDPDRLLQDVDINRLRAVVFRDVDDSKQAQFLALAVVYFISVLMVSKYRDILEPQRETARCGSQLFKNKGSFKLNMETPAVFPENIKEKETPTPVEDIQLESSIPHTDSGIGEEQMPSILNGTDLETSAGPDAMSELFSTLSSEVKKSQESLTESPSEILKPAPSISSISQSKGMNVKEILKSLVAAPIEIAECGPDPIPYPDPALKREAQSILPMQFHSFDRYMAELIILYCCNAEIWSAVDSGSSSSSSSSSFVNGATSKNLPAVQTVAPMPEDSAENMSITAKLERALEKVAPLLREIFVDFAPFLSRTLLGSHGQELLIEGLVCMKSSTSVVELVMLLCSQEWQNSIQKNAGLAFIELINEGRSGLTTAVRAQRYHEWRYQEDKTPRSQLYDLIHLAQKWLQPESWSPEEILEVLVIDRYMRALPPDLCKWVGQNDPSTYDEMITLVERRMTARELT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIS0</accession>
    <name>A0A452GIS0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Platelet-derived growth factor (PDGF) family profile domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Growth factor active in angiogenesis, vasculogenesis and endothelial cell growth. Induces endothelial cell proliferation, promotes cell migration, inhibits apoptosis and induces permeabilization of blood vessels. Binds to the FLT1/VEGFR1 and KDR/VEGFR2 receptors, heparan sulfate and heparin.</text>
    </comment>
    <comment type="subunit">
        <text evidence="10">Homodimer; disulfide-linked. Also found as heterodimer with PGF.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1 11">Belongs to the PDGF/VEGF growth factor family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIS0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001789.1">
        <property type="protein sequence ID" value="ENSGAGP00000001570.1"/>
        <property type="gene ID" value="ENSGAGG00000001262.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042056">
        <property type="term" value="F:chemoattractant activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008083">
        <property type="term" value="F:growth factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005172">
        <property type="term" value="F:vascular endothelial growth factor receptor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050930">
        <property type="term" value="P:induction of positive chemotaxis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045766">
        <property type="term" value="P:positive regulation of angiogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051781">
        <property type="term" value="P:positive regulation of cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001938">
        <property type="term" value="P:positive regulation of endothelial cell proliferation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060754">
        <property type="term" value="P:positive regulation of mast cell chemotaxis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001666">
        <property type="term" value="P:response to hypoxia"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002040">
        <property type="term" value="P:sprouting angiogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048010">
        <property type="term" value="P:vascular endothelial growth factor receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0038084">
        <property type="term" value="P:vascular endothelial growth factor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00135">
        <property type="entry name" value="PDGF"/>
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        <property type="entry name" value="Cystine-knot cytokines"/>
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        <property type="entry name" value="PDGF/VEGF_dom"/>
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        <property type="entry name" value="PDGF/VEGF_growth_factor"/>
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        <property type="entry name" value="VEGF_C"/>
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        <property type="entry name" value="VEGF_C_sf"/>
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        <property type="entry name" value="VASCULAR ENDOTHELIAL GROWTH FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="VASCULAR ENDOTHELIAL GROWTH FACTOR A, LONG FORM"/>
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    </dbReference>
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        <property type="entry name" value="PDGF"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="VEGF_C"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="PDGF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57501">
        <property type="entry name" value="Cystine-knot cytokines"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57593">
        <property type="entry name" value="Heparin-binding domain from vascular endothelial growth factor"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="PDGF_1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="PDGF_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0037">Angiogenesis</keyword>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="4" id="KW-0221">Differentiation</keyword>
    <keyword evidence="6" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="7" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="5 11" id="KW-0339">Growth factor</keyword>
    <keyword evidence="8" id="KW-0497">Mitogen</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="12" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="12">
        <location>
            <begin position="1"/>
            <end position="26"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019529522" description="Platelet-derived growth factor (PDGF) family profile domain-containing protein" evidence="12">
        <location>
            <begin position="27"/>
            <end position="192"/>
        </location>
    </feature>
    <feature type="domain" description="Platelet-derived growth factor (PDGF) family profile" evidence="13">
        <location>
            <begin position="39"/>
            <end position="135"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006686"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022657"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00054340"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00064697"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="RuleBase" id="RU003818"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50278"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001570.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="192" mass="22474" checksum="ECC284CE0CAF4012" modified="2019-05-08" version="1">MNFLFTWIHWGLAALLYLHNAKLSQAAPTLGDGERKPNEVIKFLEVYERSVCRTIETLVDIFQEYPDEVEYIFKPSCVPLLRCAGCCGDEGLECVPMEVHNVTMEIMRIKPHQSQHIVHMSFLQHSRCDCRPKKDVKAKQENHCEPCSERRKHLFVQDPQTCKCSCKFTDSRCKSRQLELNERTCRCEKPRR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIS3</accession>
    <name>A0A452GIS3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="22">Dynein axonemal heavy chain 11</fullName>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Cilium axoneme</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the dynein heavy chain family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000001794.1">
        <property type="protein sequence ID" value="ENSGAGP00000001575.1"/>
        <property type="gene ID" value="ENSGAGG00000000806.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005930">
        <property type="term" value="C:axoneme"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030286">
        <property type="term" value="C:dynein complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005874">
        <property type="term" value="C:microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031514">
        <property type="term" value="C:motile cilium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045505">
        <property type="term" value="F:dynein intermediate chain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051959">
        <property type="term" value="F:dynein light intermediate chain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007018">
        <property type="term" value="P:microtubule-based movement"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000667">
        <property type="entry name" value="Dynein axonemal heavy chain 11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.2620:FF:000004">
        <property type="entry name" value="Dynein axonemal heavy chain 17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.920.30:FF:000003">
        <property type="entry name" value="Dynein axonemal heavy chain 17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000219">
        <property type="entry name" value="Dynein axonemal heavy chain 17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.20.180.20:FF:000001">
        <property type="entry name" value="Dynein axonemal heavy chain 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.8.710:FF:000002">
        <property type="entry name" value="dynein heavy chain 17, axonemal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.140.100:FF:000001">
        <property type="entry name" value="dynein heavy chain 17, axonemal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.1120:FF:000002">
        <property type="entry name" value="Dynein heavy chain 9, axonemal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.472.130:FF:000001">
        <property type="entry name" value="Dynein, axonemal, heavy chain 9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.2620">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.472.130">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.8.710">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.1120">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.920.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.140.100">
        <property type="entry name" value="Dynein heavy chain, N-terminal domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.20.180.20">
        <property type="entry name" value="Dynein heavy chain, N-terminal domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035699">
        <property type="entry name" value="AAA_6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026983">
        <property type="entry name" value="DHC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041589">
        <property type="entry name" value="DNAH3_AAA_lid_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042222">
        <property type="entry name" value="Dynein_2_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043157">
        <property type="entry name" value="Dynein_AAA1S"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041466">
        <property type="entry name" value="Dynein_AAA5_ext"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013602">
        <property type="entry name" value="Dynein_heavy_linker"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013594">
        <property type="entry name" value="Dynein_heavy_tail"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042228">
        <property type="entry name" value="Dynein_linker_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45703:SF12">
        <property type="entry name" value="DYNEIN AXONEMAL HEAVY CHAIN 11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45703">
        <property type="entry name" value="DYNEIN HEAVY CHAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12774">
        <property type="entry name" value="AAA_6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12775">
        <property type="entry name" value="AAA_7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17857">
        <property type="entry name" value="AAA_lid_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08385">
        <property type="entry name" value="DHC_N1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08393">
        <property type="entry name" value="DHC_N2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17852">
        <property type="entry name" value="Dynein_AAA_lid"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="13" id="KW-0966">Cell projection</keyword>
    <keyword evidence="10" id="KW-0969">Cilium</keyword>
    <keyword evidence="9 14" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="12" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="8" id="KW-0243">Dynein</keyword>
    <keyword evidence="4" id="KW-0493">Microtubule</keyword>
    <keyword evidence="11" id="KW-0505">Motor protein</keyword>
    <keyword evidence="6" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="Dynein heavy chain tail" evidence="15">
        <location>
            <begin position="53"/>
            <end position="540"/>
        </location>
    </feature>
    <feature type="domain" description="Dynein heavy chain linker" evidence="16">
        <location>
            <begin position="1044"/>
            <end position="1454"/>
        </location>
    </feature>
    <feature type="domain" description="Dynein heavy chain hydrolytic ATP-binding dynein motor region" evidence="17">
        <location>
            <begin position="1588"/>
            <end position="1914"/>
        </location>
    </feature>
    <feature type="domain" description="Dynein heavy chain AAA 5 extension" evidence="18">
        <location>
            <begin position="2072"/>
            <end position="2190"/>
        </location>
    </feature>
    <feature type="domain" description="Dynein heavy chain 3 AAA+ lid" evidence="19">
        <location>
            <begin position="2407"/>
            <end position="2503"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="14">
        <location>
            <begin position="1010"/>
            <end position="1037"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004430"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008887"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023017"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023069"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="Pfam" id="PF08385"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF08393"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF12774"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF17852"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="Pfam" id="PF17857"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001575.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="22">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="2599" mass="302767" checksum="C2819708B88A3310" modified="2019-05-08" version="1">GSGGNREEQGHGALRVVVEWGGKRRWVLRWGGGRAWDRARGRKQSYLRFYHFLFTALMEAQDIELYLRPLRRHIQFLQETEFPKIHSLIPPLFHTICLIWSHSQFYSVPARIIVLLQEFCNLFIDQARSYLSPEDLLKGEIEETLEHVQIAVNTLRSFKNFFFSHREKLASYFTNSKEFKPWDFQSNMVFARFDLFLNRLVKIEDIFVVMLEFQKLEKLEFGGVKGKTLSEQIYRMNEEFIESCKVFKEKTYDPSDYNNVEFEEDYVEFKAKILDFDRRLGSILCVGFLDCSGLESAFKLLAIFGYFLEKPVIMEIFSPNYIILLQMFDEELDNCKHLYDEHIRQVEKGNEIIINKNMPFTSGNLKWSKALQERIQTFWSNFLYLILQDPETALVYQKYIETLTLIDQFEEQIYCKWKSQVDEVYQFNLDQPLIKRNPENGLLSVNFDSKLVALLREVTYLLKLNRSDIPDSALAIYEKRKIFAKYIGNLELIVQWYNKLKQTVLEVEYPLIETELEAIDGQLRETEEVFTWQKENCWKYIEQVKAAVYDLEQRVQQSKENIKTIQQIMNAWAEHTLFSRKDDRKDALLSLEDKEDRLAKKYKLLKEDGYKIHQLAEENLKLFKADSSSDSWKIYIEYIDDIIVDGFFNTLMHNLDFFLENTEKGLKPAPLFQAQMILSAPEIEFKPSLDKEAGDGFYDLVEELLSNVFKMSAQVKRVAAHLEVENYQSDMDNMLDLSEIRQEIMDRVADVIRKALDYRNSLDTYAYLWVDDRAEFMKQFLLYGHVLTSEEIDSHADEGAPEQPPTTEQFKEQIDIYENLYIQMSKFEDFRTFESWFKVDMKPFKMSLLNIIKKWSWMFKEHLLKFVIGSLTDLEEFIKVTDAGLQKEVNEGDYNGLVEIMGHLLAVRDRQASTDELFEPLKETITLLESYGQKMSDQVYVQLEELPEKWNTTKKIAASIKHEVAPLQTSEVTLIRKKFALFDGKQIEFRERFRVEAPFCFDAKYPYILLDKENRELEALEEEMLQMQKSANLFEVVVPDYKQLKQCRKEIRLLKEVWDINIYVTSSIGDWTKTQWRQINVEQMDAELRRFAKASSIAEVWSLDKEIRSWDVYIGLELTVKNLMTSLRAVTELQNPAIRERHWHQLMNATDVRLSITEGTTLADLLGLQLHKVEDEVRSIVDKAVKELGTEKILTEISQTWATVEFCYEEHYRTRTPLLKCDEQLFETLENNQVQLQTILQSKYVEYFIEQVTSWQRTLNVADSVIFIWMEIQRTWSHLESIFIGSDDIRTKLPKDAKRFDGVDADFKELMMETANTKNVLEATTKPNLYEKLESLQRRLSLCEKALAEYLETKCLAFPRFYFVSSADLLDILSKGTQPRQVTRHLTKLFDNIADLQFQDNKEKSTDIALGMYSKEKEYVPFHAECECIGQVETWLQQLEETMYKTVHRHIMEAIAAYEEKPREQWIFDYPAQVALTSSQIWWTTDVGIAFNRLEEGFETALKDYNKKQIAQLNALIALLLGELASGDRQKIMTICTIDVHARDVVAKLVAQKVTSSQAFAWLSQLRHRWDEAQKHCLANICDAQFQYFYEYLGNTSRLVITPLTDRCYITLTQSLHLTMSGAPAGPTGTGKTETTKDLGHALGMMVYVFNCSEQMDYKSIGNIYKGLVQTGAWGCFDEFNRISVEVLSVVAVQVKTIHDAIRNKRKKFLFLGENITLKTSVGIFITMNPGYAGRTELPENLKALFRPCAMVVPDIELICEIMLVAEGFTDARLLARKFITLYTLCRELLSKQDHYDWGLRAVKSVLVVAGSLKRGDKNRPEDQVLMRTLRDFNLPKIVTDDIPIFMGLISDLFPALDVPRRRDLQFEQMVKQSTLELHLQPEESFIFKVVQLEELLAVRHSVFVVGNAGTGKSKVLRTLNRTYVNMKQKLVWNDLNPKAVTTDELFGFIHHATREWKDGLFSSLLREQANITQDGPKWIVLDGDIDPMWIESLNTVMDDNKVLTLASNERVPLTPSMRLLFEIHHLKTATPATVSRAGILYVNPQDLGWNPYVASWIETRSHQSEKANLTILFDKYVPPCLDQLRTSFKTITPIPENSMVQTLCSLLDCLLTPENVPPDCPRELYEIYFVFACVWAFGGALFQDQLSNYQAEFSRWWLKEMKAVKFPSQGTIFDYYLDPKTRKFLPWTDKVPRFDMDPETPLQAVLVHTSETTRLRYFTDLLLERGKPVMLVGNAGVGKTVFVGDRLAAFPDDYMVSNVLFNYYTTSSVLQKILEKSLEKKAGRNYGPVGNKKLVYFIDDMNMPEMDVYGTVQPHTLIRQHIDYGHWYDRQKLTVKEIHNCHYVACMNPTAGNFTINPRLQRHFTVFALNFPSVDALNTIYSKILGFHFQRHVFNPSVLKNAPAMIQASIWLHQTMVQNFLPTAVKFHYIFNLRDLSNIFQGILFATPECLREPNDLVCLWLHESSRVYGDKLVEPKDCHFFRKKLIDTAHKYFEGVEDHLLRQHPLIYCHFANGVASPHYMPVKEWEVLRKILTETLESYNELNASMNLVLFEDAMQHHWTXPLWGLEQIAPLGILKNRKNHGQCLLSFKTCCGFLN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIS9</accession>
    <name>A0A452GIS9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Platelet-derived growth factor (PDGF) family profile domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Growth factor active in angiogenesis, vasculogenesis and endothelial cell growth. Induces endothelial cell proliferation, promotes cell migration, inhibits apoptosis and induces permeabilization of blood vessels. Binds to the FLT1/VEGFR1 and KDR/VEGFR2 receptors, heparan sulfate and heparin.</text>
    </comment>
    <comment type="subunit">
        <text evidence="10">Homodimer; disulfide-linked. Also found as heterodimer with PGF.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1 11">Belongs to the PDGF/VEGF growth factor family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIS9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001781.1">
        <property type="protein sequence ID" value="ENSGAGP00000001562.1"/>
        <property type="gene ID" value="ENSGAGG00000001262.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042056">
        <property type="term" value="F:chemoattractant activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008083">
        <property type="term" value="F:growth factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005172">
        <property type="term" value="F:vascular endothelial growth factor receptor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050930">
        <property type="term" value="P:induction of positive chemotaxis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045766">
        <property type="term" value="P:positive regulation of angiogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051781">
        <property type="term" value="P:positive regulation of cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001938">
        <property type="term" value="P:positive regulation of endothelial cell proliferation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060754">
        <property type="term" value="P:positive regulation of mast cell chemotaxis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001666">
        <property type="term" value="P:response to hypoxia"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002040">
        <property type="term" value="P:sprouting angiogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048010">
        <property type="term" value="P:vascular endothelial growth factor receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0038084">
        <property type="term" value="P:vascular endothelial growth factor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00135">
        <property type="entry name" value="PDGF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.160.10:FF:000001">
        <property type="entry name" value="Vascular endothelial growth factor A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.90.10:FF:000009">
        <property type="entry name" value="Vascular endothelial growth factor A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.90.10">
        <property type="entry name" value="Cystine-knot cytokines"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.160.10">
        <property type="entry name" value="Vascular endothelial growth factor, heparin-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029034">
        <property type="entry name" value="Cystine-knot_cytokine"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023581">
        <property type="entry name" value="PD_growth_factor_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000072">
        <property type="entry name" value="PDGF/VEGF_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050507">
        <property type="entry name" value="PDGF/VEGF_growth_factor"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027928">
        <property type="entry name" value="VEGF_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036841">
        <property type="entry name" value="VEGF_C_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12025">
        <property type="entry name" value="VASCULAR ENDOTHELIAL GROWTH FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12025:SF5">
        <property type="entry name" value="VASCULAR ENDOTHELIAL GROWTH FACTOR A, LONG FORM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00341">
        <property type="entry name" value="PDGF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14554">
        <property type="entry name" value="VEGF_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00141">
        <property type="entry name" value="PDGF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57501">
        <property type="entry name" value="Cystine-knot cytokines"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57593">
        <property type="entry name" value="Heparin-binding domain from vascular endothelial growth factor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00249">
        <property type="entry name" value="PDGF_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50278">
        <property type="entry name" value="PDGF_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0037">Angiogenesis</keyword>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="4" id="KW-0221">Differentiation</keyword>
    <keyword evidence="6" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="7" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="5 11" id="KW-0339">Growth factor</keyword>
    <keyword evidence="8" id="KW-0497">Mitogen</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="13" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="13">
        <location>
            <begin position="1"/>
            <end position="26"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019474301" description="Platelet-derived growth factor (PDGF) family profile domain-containing protein" evidence="13">
        <location>
            <begin position="27"/>
            <end position="216"/>
        </location>
    </feature>
    <feature type="domain" description="Platelet-derived growth factor (PDGF) family profile" evidence="14">
        <location>
            <begin position="39"/>
            <end position="135"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="133"/>
            <end position="159"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="12">
        <location>
            <begin position="142"/>
            <end position="159"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006686"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022657"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00054340"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00064697"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="RuleBase" id="RU003818"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50278"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001562.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="216" mass="25364" checksum="1721883231F110DA" modified="2019-05-08" version="1">MNFLFTWIHWGLAALLYLHNAKLSQAAPTLGDGERKPNEVIKFLEVYERSVCRTIETLVDIFQEYPDEVEYIFKPSCVPLLRCAGCCGDEGLECVPMEVHNVTMEIMRIKPHQSQHIVHMSFLQHSRCDCRPKKDVKAKQEKKSKRGKGKGQKRKRKKSRYKLLSFHCEPCSERRKHLFVQDPQTCKCSCKFTDSRCKSRQLELNERTCRCEKPRR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIT4</accession>
    <name>A0A452GIT4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">KRAB domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GIT4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001787.1">
        <property type="protein sequence ID" value="ENSGAGP00000001568.1"/>
        <property type="gene ID" value="ENSGAGG00000001242.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006355">
        <property type="term" value="P:regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd07765">
        <property type="entry name" value="KRAB_A-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001909">
        <property type="entry name" value="KRAB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036051">
        <property type="entry name" value="KRAB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050169">
        <property type="entry name" value="Krueppel_C2H2_ZnF"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23232:SF142">
        <property type="entry name" value="GASTRULA ZINC FINGER PROTEIN XLCGF57.1-LIKE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23232">
        <property type="entry name" value="KRAB DOMAIN C2H2 ZINC FINGER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01352">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00349">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109640">
        <property type="entry name" value="KRAB domain (Kruppel-associated box)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50805">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="KRAB" evidence="1">
        <location>
            <begin position="18"/>
            <end position="89"/>
        </location>
    </feature>
    <evidence type="ECO:0000259" key="1">
        <source>
            <dbReference type="PROSITE" id="PS50805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001568.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="96" mass="11148" checksum="380BBCE39BC70512" modified="2019-05-08" version="1">LFKTYFQCVGLFFFQGPVTFEEVAVYFTREEWALLDPVQRALYRDVMQENYEMVVLLGFPISKPDVISQLERGEEPWVADLQDLKVSCPLIGPLVK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIT6</accession>
    <name>A0A452GIT6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Armadillo repeat-containing protein 8</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIT6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001726.1">
        <property type="protein sequence ID" value="ENSGAGP00000001512.1"/>
        <property type="gene ID" value="ENSGAGG00000001203.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034657">
        <property type="term" value="C:GID complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043161">
        <property type="term" value="P:proteasome-mediated ubiquitin-dependent protein catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000489">
        <property type="entry name" value="Armadillo repeat containing 8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000061">
        <property type="entry name" value="armadillo repeat-containing protein 8 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000225">
        <property type="entry name" value="Armadillo"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038739">
        <property type="entry name" value="ARMC8/Vid28"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15651">
        <property type="entry name" value="ARMADILLO REPEAT-CONTAINING PROTEIN 8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15651:SF7">
        <property type="entry name" value="ARMADILLO REPEAT-CONTAINING PROTEIN 8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00514">
        <property type="entry name" value="Arm"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00185">
        <property type="entry name" value="ARM"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50176">
        <property type="entry name" value="ARM_REPEAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0539">Nucleus</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <feature type="repeat" description="ARM" evidence="7">
        <location>
            <begin position="61"/>
            <end position="97"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00013746"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00259"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001512.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="646" mass="72415" checksum="2A08D6C8B8EB4A18" modified="2019-05-08" version="1">MACVLETPLRMSVLSEVTASSRHYVDRLFDPDPQKVLQGVIDMKNAVIGNNKQKANLIVLGAVPRLLYLLQQETSSTELRTECAVVLGSLAMGTENNVKSLLDCHIIPALLQGLLSPELKFIEACLRCLRTIFTSPVTPEELLYTDATVIPHLMALLSRSRYTQEYICQIFSHCCKVRMQALKCFSVLAFENPQVSMTLVNVLVDGELLPQIFVKMLQRDKPIEMQLTSAKCLTYMCRAGAIQTDDNCIVLKALPSLVRMCSKERLLEERVEGAETLAYLIETDVELQRIASITDHLIAMLSDYFKYPSSVSAITDIKRLDHDLKHAHELRQAAFKLYASLGANDEDIRKKIIETENMMDRIVNGLSESSIKVRLSAVRCLHSLSRSVQQLRTSFQDHAVWKPLMKVLQNAPDEVLVVASSTLCNLLLEFSPSKEPILESGAIELLCSLTQSENLALRVNGIWALMNMAFQAEQKIKADILQGLSTEQLFQLLSDSDVNVLMKTLGLLRNLLSTRPHIDQIMSTHGKQIMQAVTLILEGEHNIEVKEQTLCILANIADGTTAKELIMTNDDILQKIKYYMSHSNAKLQLAAMFCISNLIWNEEEGSQERQDKLRDIGVVDILHKLSQSSDPNLCDKAKTALQQYLA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIU0</accession>
    <name>A0A452GIU0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">DUF4704 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000001809.1">
        <property type="protein sequence ID" value="ENSGAGP00000001590.1"/>
        <property type="gene ID" value="ENSGAGG00000000400.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019901">
        <property type="term" value="F:protein kinase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008104">
        <property type="term" value="P:intracellular protein localization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.200:FF:000010">
        <property type="entry name" value="neurobeachin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.200">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050865">
        <property type="entry name" value="BEACH_Domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031570">
        <property type="entry name" value="NBEA/BDCP_DUF4704"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13743">
        <property type="entry name" value="BEIGE/BEACH-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13743:SF62">
        <property type="entry name" value="NEUROBEACHIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15787">
        <property type="entry name" value="DUF4704"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13385">
        <property type="entry name" value="Laminin_G_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0853">WD repeat</keyword>
    <feature type="domain" description="DUF4704" evidence="3">
        <location>
            <begin position="361"/>
            <end position="840"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="864"/>
            <end position="887"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1132"/>
            <end position="1168"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1710"/>
            <end position="1729"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="873"/>
            <end position="886"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="1132"/>
            <end position="1156"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="1157"/>
            <end position="1168"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="1714"/>
            <end position="1724"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022574"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="Pfam" id="PF15787"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001590.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1837" mass="203890" checksum="11D3A33B30D5A716" modified="2019-05-08" version="1">MNFIIQDAESITCMSELLEHCDVTCQAEIWSMFTAILRKSVRNLQTSTEVGLIEQVLLKMSTVDDMIADLLVDMLGVLASYSITVKELKLLFSMLRGENGIWPRHAIKLLSVLNQMPQRHGPDTFFNFPGCSAAAIALPPIAKWPYQNGFTLNTWFRMDPLNNINVDKDKPYLYCFRTSKGVGYSAHFVGNCLIVTSLKSKGKGFQHCVKYDFQPRKWYMISIVHIYNRWRNSEIRCYVNGQLVSYGDMAWHVNTNDSYDKCFLGSSETADANRVFCGQLGAVYVFTEALNPAQIFAIHQLGPGYKSTFKFKSESDIHLAEHHKQVLYDGKLASSIAFTYNAKATDAQLCLESSPKENPSIFVHSPHALMLQDVKAIVTHSIHSAIHSIGGIQVLFPLFAQLDNRQLNDSQMETTVCATLLAFLVELLKSSVAMQEQMLGGKGFLVIGYLLEKSSRVHITRAVLEQFLSFAKYLDGLSHGAPLLKQLCDHILFNPAIWIHTPAKVQLSLYTYLSAEFIGTATIYTTIRRVGTVLQLMHTLKYYYWVVNPADSSGITPKGLEGPRPSQKEIISLRAFMLLFLKQLILKDRGVKEDELQSILNYLLTMHEDENIHDVLQLLVALMSEHPASMIPAFDQRNGIRVIYKLLASKSESIWVQALKVLGYFLKHLGHKRKVEIMHTHSLFTLLGERLMLHTNTVTITTYNTLYEILTEQVCTQVVHKPHPEPDSTVKIQNPMILKVVATLLKNSTPSAELMEVRRLFLSDMIKLFSNSRENRRCLLQCSVWQDWMFSLGYINPKNSEEQKITEMVYNIFRILLYHAIKYEWGGWRVWVDTLSIAHSKVTYEAHKEYLAKMYEEYQRQEEENIKKGKKGNVSTISGLSSQTTGAKGGLEIREIEDLSQSQSPESETDYPVSTDTRDLLMATKVSDEVLGSSDRPGGGVHVEVHDLLVDIKAEKVEATEVKLDDMDLSPETLVTGENGALVEVESLLDNVYCAAVEKLQNNVHGSVGIIKKSEEKDNGPLITLADEKDEPSTNNTSFLFDKIPSQEEKLLPELSSNHITIPNVQETQMHLGVSDDLGLLAHMTGSVDITCTSSIIEDKEFKIHTTSDGMSNLSERELGSSSKGLEYAEMTATTLEPESSGSKSLPNVDTGSIISDTERSDDGKEAGKEIRKIQTTTTTQAVQGRSVTQQDRDLRVDLGFRGMPMTEEQRRQFSPGPRTTMFRIPEFKWSPMHQRLLTDLLFALETDVHVWRSHSTKSVMDFVNSNENIIFVHNTIHLISQMVDNIIIACGGILPLLSAATSPTVSTIFPSSGHTELENIEVTQGMSAETAVTFLSRLMAMVDVLVFASSLNFSEIEAEKNMSSGGLMRQCLRLVCCVAVRNCLECRQRQRERVNKSSLISSKTQETLQGVTATALTKSPLENVPGNLSPIKDPDRLLQDVDINRLRAVVFRDVDDSKQAQFLALAVVYFISVLMVSKYRDILEPQRETARCGSQAGRNIRQEINSPTSTETPAVFPENIKEKETPTPVEDIQLESSIPHTDSGIGEEQMPSILNGTDLETSAGPDAMSELFSTLSSEVKKSQESLTESPSEILKPAPSISSISQSKGMNVKEILKSLVAAPIEIAECGPDPIPYPDPALKREAQSILPMQFHSFDRSVVVPVKKPPPGSLAVTTVGAATAGGVLPPGTTPNIFAATGATPKSMINTTGAVDSGSSSSSSSSSFVNGATSKNLPAVQTVAPMPEDSAENMSITAKLERALEKVAPLLREIFVDFAPFLSRTLLGSHGQELLIEGLVCMKSSTSVVELVMLLCSQEWQNSIQKNAGLAFIELINEGR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIU1</accession>
    <name>A0A452GIU1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">DNA ligase</fullName>
            <ecNumber evidence="15">6.5.1.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">DNA ligase that seals nicks in double-stranded during DNA repair. Also involved in DNA replication and DNA recombination.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13 15">
            <text>ATP + (deoxyribonucleotide)n-3'-hydroxyl + 5'-phospho-(deoxyribonucleotide)m = (deoxyribonucleotide)n+m + AMP + diphosphate.</text>
            <dbReference type="EC" id="6.5.1.1"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 16">Belongs to the ATP-dependent DNA ligase family.</text>
    </comment>
    <dbReference type="EC" id="6.5.1.1" evidence="15"/>
    <dbReference type="AlphaFoldDB" id="A0A452GIU1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001803.1">
        <property type="protein sequence ID" value="ENSGAGP00000001584.1"/>
        <property type="gene ID" value="ENSGAGG00000000751.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003910">
        <property type="term" value="F:DNA ligase (ATP) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071897">
        <property type="term" value="P:DNA biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006310">
        <property type="term" value="P:DNA recombination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006281">
        <property type="term" value="P:DNA repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903461">
        <property type="term" value="P:Okazaki fragment processing involved in mitotic DNA replication"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07900">
        <property type="entry name" value="Adenylation_DNA_ligase_I_Euk"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd07969">
        <property type="entry name" value="OBF_DNA_ligase_I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.3260.10:FF:000001">
        <property type="entry name" value="DNA ligase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.50.140:FF:000062">
        <property type="entry name" value="DNA ligase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.470.30:FF:000002">
        <property type="entry name" value="DNA ligase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1490.70">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.3260.10">
        <property type="entry name" value="DNA ligase, ATP-dependent, N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.470.30">
        <property type="entry name" value="DNA ligase/mRNA capping enzyme"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.50.140">
        <property type="entry name" value="Nucleic acid-binding proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050191">
        <property type="entry name" value="ATP-dep_DNA_ligase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000977">
        <property type="entry name" value="DNA_ligase_ATP-dep"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012309">
        <property type="entry name" value="DNA_ligase_ATP-dep_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012310">
        <property type="entry name" value="DNA_ligase_ATP-dep_cent"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016059">
        <property type="entry name" value="DNA_ligase_ATP-dep_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012308">
        <property type="entry name" value="DNA_ligase_ATP-dep_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036599">
        <property type="entry name" value="DNA_ligase_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012340">
        <property type="entry name" value="NA-bd_OB-fold"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00574">
        <property type="entry name" value="dnl1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45674:SF4">
        <property type="entry name" value="DNA LIGASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45674">
        <property type="entry name" value="DNA LIGASE 1/3 FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04679">
        <property type="entry name" value="DNA_ligase_A_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01068">
        <property type="entry name" value="DNA_ligase_A_M"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04675">
        <property type="entry name" value="DNA_ligase_A_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF117018">
        <property type="entry name" value="ATP-dependent DNA ligase DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56091">
        <property type="entry name" value="DNA ligase/mRNA capping enzyme, catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50249">
        <property type="entry name" value="Nucleic acid-binding proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00697">
        <property type="entry name" value="DNA_LIGASE_A1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00333">
        <property type="entry name" value="DNA_LIGASE_A2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50160">
        <property type="entry name" value="DNA_LIGASE_A3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 15" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="12" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="4" id="KW-0132">Cell division</keyword>
    <keyword evidence="7 15" id="KW-0227">DNA damage</keyword>
    <keyword evidence="9 15" id="KW-0233">DNA recombination</keyword>
    <keyword evidence="10 15" id="KW-0234">DNA repair</keyword>
    <keyword evidence="5" id="KW-0235">DNA replication</keyword>
    <keyword evidence="3 15" id="KW-0436">Ligase</keyword>
    <keyword evidence="6 15" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="ATP-dependent DNA ligase family profile" evidence="18">
        <location>
            <begin position="527"/>
            <end position="663"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1"/>
            <end position="34"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="17">
        <location>
            <begin position="1"/>
            <end position="28"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007572"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022598"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022705"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023172"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00034003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00058910"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="RuleBase" id="RU000617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="RuleBase" id="RU004196"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50160"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001584.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="798" mass="88632" checksum="2CEBF23173520530" modified="2019-05-08" version="1">MQTPQSDISADSLSPSSSHSAGDSPALSDSPAISHAGIPRRRTASVYVPPHEVQFLQGQRIPANIRPGLHPGLCAHRVSCDSAASPMHALLQLGACLIHAIIVPVATCAGLPSLGVLVQANCWETLRNTSAVGPVYWTWWLPISLWLCDPSSFSKPVVPLDPMEYNPSKSSYHPVQDACWGHGQRVPYLAVARTFERIEEVSARLKNIETLSNLFRSVVALSPSDILPCIYLCLNRLGPAYKGLELGIGETILMKAVAQAMGRQLDQIKAEAQEKGDLGLVAESSRSNQRTMFTPPKLSAAGVFSKLQEIARMTGSASMNKKIDIIKGLFVACRHSEARYITRSLGGKLRIGLAEQSVLSAIAHAVSLTPPGQGFPPEVLDAGQSKSAEARKAWLEEQTQILKQTFCELPSYDIIVPILLEHGVESLPQHCKITPGIPLKPMLAQPTKGIGEVLKRFEEAAFTCEYKYDGERAQPSVLPCIDTPVQVKKDSVQSCILDAEAVAWDPEKRQIQPFQVLTTRKRKDVEAAEIKVQVCLYAFDILYLNGESLVKEPFSRRRQLLHESFVETEGEFLFATSMDTCSLDEISEFLDQSIKDSCEGLMVKTLEVDATYEIAKRSHKWLKLKKDYLEGVGDTLDLVVIGAYLGKGKRVGMYGGFLLACYDEESEEYQSICKIGTGFTEESLEKHYSFLKDHVLEKPRPYYRWDSSTEPNHWLAAVQVWEVKCADLSISPVHKAAIGLVDEEKGISLRFPRFLRVRDDKKPEEATTSSQVAELYRRQQQIQNQQLVEKAEEEEDFY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIU5</accession>
    <name>A0A452GIU5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Armadillo repeat-containing protein 8</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIU5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001744.1">
        <property type="protein sequence ID" value="ENSGAGP00000001529.1"/>
        <property type="gene ID" value="ENSGAGG00000001203.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034657">
        <property type="term" value="C:GID complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043161">
        <property type="term" value="P:proteasome-mediated ubiquitin-dependent protein catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000061">
        <property type="entry name" value="armadillo repeat-containing protein 8 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000070">
        <property type="entry name" value="armadillo repeat-containing protein 8 isoform X1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000225">
        <property type="entry name" value="Armadillo"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038739">
        <property type="entry name" value="ARMC8/Vid28"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15651">
        <property type="entry name" value="ARMADILLO REPEAT-CONTAINING PROTEIN 8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15651:SF7">
        <property type="entry name" value="ARMADILLO REPEAT-CONTAINING PROTEIN 8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00185">
        <property type="entry name" value="ARM"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0539">Nucleus</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00013746"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001529.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="638" mass="71851" checksum="B7F4281D6F17BB38" modified="2019-05-08" version="1">MENLYYWRTLLLYLLQQETSSTELRTECAVVLGSLAMGTENNVKSLLDCHIIPALLQGLLSPELKFIEACLRCLRTIFTSPVTPEELLYTDATVIPHLMALLSRSRYTQEYICQIFSHCCKLRHWHSCGFKSSPQPLLCANGPDHQTILFNHGAVQNIAHLLSSTSYKVRMQALKCFSVLAFENPQVSMTLVNVLVDGELLPQIFVKMLQRDKPIEMQLTSAKCLTYMCRAGAIQTDDNCIVLKALPSLVRMCSKERLLEERVEGAETLAYLIETDVELQRIASITDHLIAMLSDYFKYPSSVSAITDIKRLDHDLKHAHELRQAAFKLYASLGANDEDIRKKIIETENMMDRIVNGLSESSIKVRLSAVRCLHSLSRSVQQLRTSFQDHAVWKPLMKVLQNAPDEVLVVASSTLCNLLLEFSPSKEPILESGAIELLCSLTQSENLALRVNGIWALMNMAFQAEQKIKADILQGLSTEQLFQLLSDSDVNVLMKTLGLLRNLLSTRPHIDQIMSTHGKQIMQAVTLILEGEHNIEVKEQTLCILANIADGTTAKELIMTNDDILQKIKYYMSHSNAKLQLAAMFCISNLIWNEEEGSQERQDKLRDIGVVDILHKLSQSSDPNLCDKAKTALQQYLA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIU7</accession>
    <name>A0A452GIU7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Ig-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIU7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001737.1">
        <property type="protein sequence ID" value="ENSGAGP00000001523.1"/>
        <property type="gene ID" value="ENSGAGG00000001226.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009897">
        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042612">
        <property type="term" value="C:MHC class I protein complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002474">
        <property type="term" value="P:antigen processing and presentation of peptide antigen via MHC class I"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006955">
        <property type="term" value="P:immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.500.10">
        <property type="entry name" value="MHC class I-like antigen recognition-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003006">
        <property type="entry name" value="Ig/MHC_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003597">
        <property type="entry name" value="Ig_C1-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050208">
        <property type="entry name" value="MHC_class-I_related"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011161">
        <property type="entry name" value="MHC_I-like_Ag-recog"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037055">
        <property type="entry name" value="MHC_I-like_Ag-recog_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011162">
        <property type="entry name" value="MHC_I/II-like_Ag-recog"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16675:SF242">
        <property type="entry name" value="MAJOR HISTOCOMPATIBILITY COMPLEX CLASS I-RELATED GENE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16675">
        <property type="entry name" value="MHC CLASS I-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07654">
        <property type="entry name" value="C1-set"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00129">
        <property type="entry name" value="MHC_I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00407">
        <property type="entry name" value="IGc1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54452">
        <property type="entry name" value="MHC antigen-recognition domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00290">
        <property type="entry name" value="IG_MHC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="9" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="5" id="KW-0391">Immunity</keyword>
    <keyword evidence="7 11" id="KW-0472">Membrane</keyword>
    <keyword evidence="2" id="KW-0490">MHC I</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 12" id="KW-0732">Signal</keyword>
    <keyword evidence="3 11" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 11" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="12">
        <location>
            <begin position="1"/>
            <end position="21"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019458698" description="Ig-like domain-containing protein" evidence="12">
        <location>
            <begin position="22"/>
            <end position="277"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="206"/>
            <end position="228"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="13">
        <location>
            <begin position="104"/>
            <end position="194"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="237"/>
            <end position="277"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022451"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022859"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001523.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="277" mass="30801" checksum="D67C9A2C6737E754" modified="2019-05-08" version="1">APAISPPQAMLLLPHWQHSLAVLVTAVTHEEGTYHFTMTAQLDDVQIAYYSSDAREVRPSQEWVAQALDAEYLQEKTQQFWRLEEIFKGDIRRWMKLHNQTGVPPMLSVSRRDNPDGFTTLSCHARGFYLRPIHVSWVQDREDILAETDSSGILPNDIHSYYTQSSLEISPQQEDGHRYACQVEHSSLPEPTLVWGKGRGPLPPGVLAAIVLAVLVVAGAVGAGVVLWRRKSAGKRRSPWSRDVGMSGEWGSPVSPRGNGSSLSTPKERWGRAMGQG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIU9</accession>
    <name>A0A452GIU9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">TGF-beta family profile domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 8">Belongs to the TGF-beta family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIU9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001791.1">
        <property type="protein sequence ID" value="ENSGAGP00000001572.1"/>
        <property type="gene ID" value="ENSGAGG00000001267.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000001802.1">
        <property type="protein sequence ID" value="ENSGAGP00000001583.1"/>
        <property type="gene ID" value="ENSGAGG00000001267.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005125">
        <property type="term" value="F:cytokine activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008083">
        <property type="term" value="F:growth factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030509">
        <property type="term" value="P:BMP signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048468">
        <property type="term" value="P:cell development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051094">
        <property type="term" value="P:positive regulation of developmental process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042127">
        <property type="term" value="P:regulation of cell population proliferation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009888">
        <property type="term" value="P:tissue development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.970:FF:000008">
        <property type="entry name" value="Bone morphogenetic protein 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.970">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.90.10">
        <property type="entry name" value="Cystine-knot cytokines"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029034">
        <property type="entry name" value="Cystine-knot_cytokine"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001839">
        <property type="entry name" value="TGF-b_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001111">
        <property type="entry name" value="TGF-b_propeptide"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015615">
        <property type="entry name" value="TGF-beta-rel"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017948">
        <property type="entry name" value="TGFb_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11848:SF135">
        <property type="entry name" value="BONE MORPHOGENETIC PROTEIN 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11848">
        <property type="entry name" value="TGF-BETA FAMILY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00019">
        <property type="entry name" value="TGF_beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00688">
        <property type="entry name" value="TGFb_propeptide"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00669">
        <property type="entry name" value="INHIBINA"/>
    </dbReference>
    <dbReference type="SMART" id="SM00204">
        <property type="entry name" value="TGFB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57501">
        <property type="entry name" value="Cystine-knot cytokines"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00250">
        <property type="entry name" value="TGF_BETA_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51362">
        <property type="entry name" value="TGF_BETA_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="5" id="KW-0221">Differentiation</keyword>
    <keyword evidence="7" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6 8" id="KW-0339">Growth factor</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <feature type="domain" description="TGF-beta family profile" evidence="10">
        <location>
            <begin position="317"/>
            <end position="407"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="296"/>
            <end position="317"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006656"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="RuleBase" id="RU000354"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS51362"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001583.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="407" mass="46864" checksum="4D89AF0092E9CF6D" modified="2019-05-08" version="1">MHVVWLRQPASWLLVLWAAAPFLWLRCILADFTLDNEVHSSFIHRKLRSQERREIQREILSILGLPHRPRPHLHGKQNSAPMFMLDLYNAMSVEEEAAGAAGEEGEGFSYPYKPIFSTQGPPLASLQDSNFLTEADMVMSFVNLVEHDREFHHQRYHHREFRFDLSRIPEGEAVTAAEFRIYKDYIRERFDNETFQISVYQVLQEHPGRDSDLFLLDTRIIWAAEEGWLVFDITATSNHWVVNPQHNLGLQLSVESIDGQSINPKLAGLIGRHGPQNKQPFTVAFFKATEVHLRSIRSTGGKQRSQNRSKTPKNQEAFRVSNIAENSSSDQRQACKKHELYVSFRDLGWQDWIIAPEGYAAYYCEGECAFPLNSYMNATNHAIVQTLLCLLLEETLLDGRKPEILCL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIV2</accession>
    <name>A0A452GIV2_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="10">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Golgi apparatus membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the nucleotide-sugar transporter family. SLC35A subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIV2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001751.1">
        <property type="protein sequence ID" value="ENSGAGP00000001535.1"/>
        <property type="gene ID" value="ENSGAGG00000001243.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000139">
        <property type="term" value="C:Golgi membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015165">
        <property type="term" value="F:pyrimidine nucleotide-sugar transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.3730.20:FF:000037">
        <property type="entry name" value="Nucleotide Sugar TransPorter family"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037185">
        <property type="entry name" value="EmrE-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007271">
        <property type="entry name" value="Nuc_sug_transpt"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00803">
        <property type="entry name" value="nst"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10231">
        <property type="entry name" value="NUCLEOTIDE-SUGAR TRANSMEMBRANE TRANSPORTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04142">
        <property type="entry name" value="Nuc_sug_transp"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF005799">
        <property type="entry name" value="UDP-gal_transpt"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103481">
        <property type="entry name" value="Multidrug resistance efflux transporter EmrE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="8 9" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0762">Sugar transport</keyword>
    <keyword evidence="5 9" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 9" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="7"/>
            <end position="26"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="38"/>
            <end position="59"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="136"/>
            <end position="154"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="174"/>
            <end position="194"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="206"/>
            <end position="226"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="246"/>
            <end position="266"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="273"/>
            <end position="290"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004653"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009976"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022597"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001535.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="297" mass="33089" checksum="86E479AE862D94D4" modified="2019-05-08" version="1">MSTNLKYLSLGILVFQTTSLVLTMRYSRTLKEEGPRYLSSTAVVVAEFLKIMACILLVYKDSKCNLRALNRVLRDEILNKPMETLKLAIPSGIYTLQNNLLYVALSNLDAATYQVTYQLKILTTALFSVSMLSKKLGVYQWLSLVILMTGVAFVQWPSDSQATTTKELSAGSQFVGLIAVLIACFSSGFAGVYFEKILKETKQSVWIRNIQLGFFGSIFGLMGVYIYDGELLSKNGFFQGYNKLTWIVVVLQALGGLVIAAVIKYADNILKGFATSLSIILSTLISYFWLQDFVPTR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIY2</accession>
    <name>A0A452GIY2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">KRAB domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GIY2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001779.1">
        <property type="protein sequence ID" value="ENSGAGP00000001560.1"/>
        <property type="gene ID" value="ENSGAGG00000001242.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006355">
        <property type="term" value="P:regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd07765">
        <property type="entry name" value="KRAB_A-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001909">
        <property type="entry name" value="KRAB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036051">
        <property type="entry name" value="KRAB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050169">
        <property type="entry name" value="Krueppel_C2H2_ZnF"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23232:SF142">
        <property type="entry name" value="GASTRULA ZINC FINGER PROTEIN XLCGF57.1-LIKE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23232">
        <property type="entry name" value="KRAB DOMAIN C2H2 ZINC FINGER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01352">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00349">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109640">
        <property type="entry name" value="KRAB domain (Kruppel-associated box)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50805">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="KRAB" evidence="1">
        <location>
            <begin position="4"/>
            <end position="76"/>
        </location>
    </feature>
    <evidence type="ECO:0000259" key="1">
        <source>
            <dbReference type="PROSITE" id="PS50805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001560.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="89" mass="10258" checksum="5686B0E2AA34BEFE" modified="2019-05-08" version="1">MGPVTFEEVAVYFTREEWALLDPVQRALYRDVMQENYEMVVLLVGFPISKPDVISQLERGEEPWVADLQGKKYLCFTSTRSIYGIGYSS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJ34</accession>
    <name>A0A452GJ34_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Protein transport protein sec16</fullName>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="16">Plays a role in the organization of the endoplasmic reticulum exit sites (ERES), also known as transitional endoplasmic reticulum (tER). Required for secretory cargo traffic from the endoplasmic reticulum to the Golgi apparatus.</text>
    </comment>
    <comment type="subunit">
        <text evidence="16">SEC16A and SEC16B are each present in multiple copies in a heteromeric complex.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Cytosol</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="5">Cytoplasm</location>
            <location evidence="5">Perinuclear region</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Endoplasmic reticulum membrane</location>
            <topology evidence="2">Peripheral membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Golgi apparatus membrane</location>
            <topology evidence="1">Peripheral membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Microsome membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="6 16">Belongs to the SEC16 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000001831.1">
        <property type="protein sequence ID" value="ENSGAGP00000001611.1"/>
        <property type="gene ID" value="ENSGAGG00000001287.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070971">
        <property type="term" value="C:endoplasmic reticulum exit site"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012507">
        <property type="term" value="C:ER to Golgi transport vesicle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000139">
        <property type="term" value="C:Golgi membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048471">
        <property type="term" value="C:perinuclear region of cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007029">
        <property type="term" value="P:endoplasmic reticulum organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007030">
        <property type="term" value="P:Golgi organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051668">
        <property type="term" value="P:localization within membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070973">
        <property type="term" value="P:protein localization to endoplasmic reticulum exit site"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016192">
        <property type="term" value="P:vesicle-mediated transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd09233">
        <property type="entry name" value="ACE1-Sec16-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.1030:FF:000002">
        <property type="entry name" value="Protein transport protein sec16"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.1030">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024340">
        <property type="entry name" value="Sec16_CCD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024298">
        <property type="entry name" value="Sec16_Sec23-bd"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13402:SF13">
        <property type="entry name" value="PROTEIN TRANSPORT PROTEIN SEC16A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13402">
        <property type="entry name" value="RGPR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12932">
        <property type="entry name" value="Sec16"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12931">
        <property type="entry name" value="TPR_Sec16"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="10 16" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="12 16" id="KW-0931">ER-Golgi transport</keyword>
    <keyword evidence="14 16" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="15 16" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-0492">Microsome</keyword>
    <keyword evidence="9" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="13 16" id="KW-0653">Protein transport</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7 16" id="KW-0813">Transport</keyword>
    <feature type="domain" description="Sec16 central conserved" evidence="19">
        <location>
            <begin position="1544"/>
            <end position="1642"/>
        </location>
    </feature>
    <feature type="domain" description="Sec16 Sec23-binding" evidence="18">
        <location>
            <begin position="1714"/>
            <end position="1948"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1"/>
            <end position="113"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="184"/>
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            <dbReference type="Pfam" id="PF12931"/>
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            <dbReference type="Pfam" id="PF12932"/>
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            <dbReference type="Ensembl" id="ENSGAGP00000001611.1"/>
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    <sequence length="2467" mass="269707" checksum="2B26F78E96680802" modified="2019-05-08" version="1">MQPPPQTVPAGVGGPPPAGIARNTYWRNSPFSRRANAPVSGATAPVQPVTDPFAFGRQTPQGSLLGNPSKGNPLVMPGPAPSVFPRPAGVHSSQSYAGDNSHGLPTSLPTSVSQPGITSNNFSNVVTPSSPAHIINNTGEVHPNAEHGPHNSSAQSLYNTGAAHESSLSGHLAMAHVANRPLSRQDINRDPSNTSSVPTIAPFLPPPLQQTPPQWRPIRENLQPQVQNYWPYTEPPSQNAVYSVSHSSAVTSHLPPQADLHQGPVHQHIPQSTMQAPLPVGGEKNEISSFLVANHHVNSFQPENMFRQNIRVTNPWPSQPYQEQFYPQPLLPDSGFVNPITQENNSQKQSQNVSETPTGHTSTNADSGTLSMFFKGDEAENEEILSSEKKDLSSKANYDACQLNSGHIYHQPLPPQQAAASVLCQPQISTSSANEAVQKGMDVQYFSKTISSQHETQTSKNSMYISDDKANVSKARENVGSHCENVENLECIQNQEVLPSEPQTINRSSPSVGSDLYRYGSLPGQLLPKNAIVSHAERGPNLEAPDSLPHPVRSDSVSSNYSNISHKSVSSSTRPQELVGTFIQQESGKPDEESSASFFKQIDSSPLGGDSSEPNVSKNYHSNLSQPPTPSPPKPTGIFQTSANSSFEPVRSHGVGVKPAEVDQAKMVVELRENHPNQKNIKKSAAMPAASPGNLEQPPDNLETIFMPQVYPLPLTVTSEPGNILQPAGGPVVENIQSVPEKRPSTRAQGAAKKCESPATTLWAHNELPNFGGNVLLAPAAPAVYVPAKQTVEIIQPPEEGLSSQQPNKPECVPVQQSQHGNASSENLENPPKMGEEEALQSQASSGYASLLSSPPTDSLQNQPILIAQPNQSYNLAQPINFSLSLCNQLTRNENNLSLKDPGVEDKPVTGLQAPHAGGIPSGENVSLSVMQVGSSVNTPPSSNLSHSNLLQIPVNSSEITSNQSTNLLMQPPSQTPINLAPEGQKNANSEGCLPEFASKPGSNSAVSPGTNLPSGNASTLLVPPANTMVPNNNSTNDSHSHEEASGALDFTVTRTLDKSKTSNLIQMHNPLLSCGPVFPQQPVNHASQVGPNTHDKQHFYQQVTKDVQLQAPSDRVTQGALLSQQQKQTAQMLQTVPSGQSLVPSNYQMASGIKFTQALQQQENQVSTNRYYPAGPPEGSSTQPPTSYSQTSPDKQPFSGQSLSAPTSSASVTTSHPAMPTMQQEQNPPPPQTPQDACGPPQNPYYYYRHPYDAYQTPYPQPYPPLDPRTAAHLYYQDDVYGQYGPRYQHYDRSNAAYAEPGCYRYGEPERPSSRASQCSDRPPSRQGYTEDYYNPKSGWNDYYADYYANPYNYGDPNRWERYPSAYDSRYRDPRSYDQRYWYDTEQNPYQKREAYPYDNRQDRYEDHWRYDPRFAGSFDDETEPHRDPYGDEFDRRSVHSEHSAHSLRSSHSVHSRKSSFSSRSQQSQLYRSNHDLTANAYETTTQPVSLHTDYAYGGYPANFDGQQPFTDYGYSAESGWPSVEQVPSRPSTPEKFLVPHVCARFGPGGHLTKVLPNLPSEGQPALVEIHSMETMLQHMPEQEEMRAFPGPLAKDDTHKVDVINFAQNKVTQCFQNENLIDKESASLLWDFIVLLCRQNGTVVGTDIAELLLRDHKTVWLPGKSPNEANLIDFTNEALEQVEEESGEAQLSFLTNSLITTIDSLERETERFRELLLYGRKKDALESAMKHGLWGHALLLASKMDNRTHARVMTRFANSLPINDPLQTVYQLMSGRMPAASTCCGDEKWGDWRPHLAMVLSNLTNNMEVESRTITTMGDTLASKGLLDAAHFCYLMAQIGFGVYTKKTTKLVLIGSNHSLPFLKFATNEAIQRTEAYEYAQSLGTQPCCLPNFQVFKFIYACRLAEMGLAAQAFHYCEVISKTVLKNPYYYSPVLISQLIQISSQLRLFDPQIKEKPEQELFIEPSWLVRLRHLDGQIKDGTIAYNADRSTPQQYACSTPSSELDHISQCDGTGSGQEMGPATENPLLTSLLPNTMHPMQGVQLMPSAPQTILEGSAAMTPPTQQETVGGVPFYPVAPPSIGPGSGFAPPGFPNQYGAEQSSLYLGSTVPPGGPPPQAAEPQPEGHLNQETAMQRIPQESPIQKTFPEQREEDFYGKMANMGYGRRSRTTSESSTHSIGRERCNSAAKQPSPPPPSIPEGKETNKEIKKEPAARKSGANWFRWLMGKGKNEAHLPDDKNKSIVWDEKKQRWVNLDEPEEERKPPPPPPAGFPKALQTAPPGPGGPPGVNMFSRKAAGTRARYVDVLNPSGAKSSGAVPAPSDLFAPLAPMPIPTNLFLPNSVPEEQQPMEGSEAQEQTLSANQVSADAATEPQYLNSTMLPPGSELPGSNPDGSQSGELSRSSSMSSLSREVSQHFNQPLCSVPPSGGPPSGTVQLYNPSQFAQSPATTRSSRLGRIGQRKYPTLK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJ39</accession>
    <name>A0A452GJ39_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">Probable ATP-dependent RNA helicase DDX27</fullName>
            <ecNumber evidence="3">3.6.4.13</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="18">DEAD box protein 27</fullName>
        </alternativeName>
    </protein>
    <organism evidence="25 26">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="26" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="25" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="25" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">Probable ATP-dependent RNA helicase. Component of the nucleolar ribosomal RNA (rRNA) processing machinery that regulates 3' end formation of ribosomal 47S rRNA.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="14">
            <text>ATP + H2O = ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:13065"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="3.6.4.13"/>
        </reaction>
    </comment>
    <comment type="subunit">
        <text evidence="16">Associates with PeBoW complex, composed of BOP1, PES1 and WDR12. Interacts directly with BOP1 and PES1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Chromosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
            <location evidence="2">Nucleolus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="13">Belongs to the DEAD box helicase family. DDX27/DRS1 subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.6.4.13" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452GJ39"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001847.1">
        <property type="protein sequence ID" value="ENSGAGP00000001627.1"/>
        <property type="gene ID" value="ENSGAGG00000001296.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005694">
        <property type="term" value="C:chromosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005730">
        <property type="term" value="C:nucleolus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016787">
        <property type="term" value="F:hydrolase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003676">
        <property type="term" value="F:nucleic acid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003724">
        <property type="term" value="F:RNA helicase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006364">
        <property type="term" value="P:rRNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd17947">
        <property type="entry name" value="DEADc_DDX27"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18787">
        <property type="entry name" value="SF2_C_DEAD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000842">
        <property type="entry name" value="ATP-dependent RNA helicase DRS1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:001134">
        <property type="entry name" value="Probable ATP-dependent RNA helicase DDX27"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011545">
        <property type="entry name" value="DEAD/DEAH_box_helicase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050079">
        <property type="entry name" value="DEAD_box_RNA_helicase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014001">
        <property type="entry name" value="Helicase_ATP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001650">
        <property type="entry name" value="Helicase_C-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000629">
        <property type="entry name" value="RNA-helicase_DEAD-box_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014014">
        <property type="entry name" value="RNA_helicase_DEAD_Q_motif"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47959">
        <property type="entry name" value="ATP-DEPENDENT RNA HELICASE RHLE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47959:SF22">
        <property type="entry name" value="RNA HELICASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00270">
        <property type="entry name" value="DEAD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00271">
        <property type="entry name" value="Helicase_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00487">
        <property type="entry name" value="DEXDc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00490">
        <property type="entry name" value="HELICc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00039">
        <property type="entry name" value="DEAD_ATP_HELICASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51192">
        <property type="entry name" value="HELICASE_ATP_BIND_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51194">
        <property type="entry name" value="HELICASE_CTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51195">
        <property type="entry name" value="Q_MOTIF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="11 20" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="4" id="KW-0158">Chromosome</keyword>
    <keyword evidence="10 20" id="KW-0347">Helicase</keyword>
    <keyword evidence="9 20" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="8 20" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="12" id="KW-0539">Nucleus</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="26" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0690">Ribosome biogenesis</keyword>
    <keyword evidence="6" id="KW-0698">rRNA processing</keyword>
    <feature type="domain" description="DEAD-box RNA helicase Q" evidence="24">
        <location>
            <begin position="159"/>
            <end position="187"/>
        </location>
    </feature>
    <feature type="domain" description="Helicase ATP-binding" evidence="22">
        <location>
            <begin position="190"/>
            <end position="364"/>
        </location>
    </feature>
    <feature type="domain" description="Helicase C-terminal" evidence="23">
        <location>
            <begin position="398"/>
            <end position="544"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="21">
        <location>
            <begin position="1"/>
            <end position="123"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="21">
        <location>
            <begin position="580"/>
            <end position="599"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="21">
        <location>
            <begin position="620"/>
            <end position="639"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="21">
        <location>
            <begin position="655"/>
            <end position="736"/>
        </location>
    </feature>
    <feature type="short sequence motif" description="Q motif" evidence="19">
        <location>
            <begin position="159"/>
            <end position="187"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="21">
        <location>
            <begin position="16"/>
            <end position="31"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="21">
        <location>
            <begin position="57"/>
            <end position="67"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="21">
        <location>
            <begin position="72"/>
            <end position="101"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="21">
        <location>
            <begin position="102"/>
            <end position="112"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="21">
        <location>
            <begin position="625"/>
            <end position="639"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="21">
        <location>
            <begin position="657"/>
            <end position="666"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="21">
        <location>
            <begin position="716"/>
            <end position="736"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004604"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012552"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022454"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022517"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022552"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022806"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00043999"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00047984"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00053341"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00065947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00072908"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00075453"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00552"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="RuleBase" id="RU000492"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS51192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS51194"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS51195"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001627.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="736" mass="83374" checksum="B2303C96E5CAFDB0" modified="2019-05-08" version="1">MLQGLGLVATLGEGDAAPEEPESGDSADEEEPLGRGQQRRRGGDFSAGFVFAEKEAPGAASGAWAEALRQLRGKEREVKHVKMKGEDTKVEKDETKEKESEEGSEEEEEDVESQYSSTDENILTKADTLKVKERKRQKNVGEMETKIFFEDASQYDDNLSFQDMNLSRPLLKAITALSFKQPTPIQKACIPVGLLGKDICACAATGTGKTAAFTLPVLERLIYKPRQAPVTRVLVLVPTRELGIQVHSVTKQLAQFSSITTCLAVGGLDVKTQEAALRSGPDILIATPGRLIDHLHNCPSFHLSSIEVLILDEADRMLDEYFEEQMKEIIRLCSHHRQTMLFSATMTDEVKDLAAVSLKNPVRIFVNSNTDVAPFLRQEFIRVRPNREGDREAIVSALLTRTFTDHVMLFTQTKKQAHRMHILLGLMGLQVGELHGNLSQTQRLEALRRFKDEQIDVLVATDVAARGLDIEGVKTVINFTMPNTTKHYVHRVGRTARAGRAGRSVSLVGEEERKMLKEIVKTAKMPVKARILPQDVILKFREKIEKLEKDVYAVLRLEREEREMQQSEAQINMAKKRLEAGEQKVEEGPERSWFQSREERKKEKLAKALQEFDLALRGKKKRKKFMQDTQKKGQMTPEERSQFEILKSQMYAERVAKRNRRAKRARVVPEDEPPSTAAGLKRKKQSSVFDEELTNTSRKALKQYRAGPSFEERKRLGLARHRKGGNFKSKSRYKRK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJ41</accession>
    <name>A0A452GJ41_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">RING-type E3 ubiquitin transferase</fullName>
            <ecNumber evidence="5">2.3.2.27</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>S-ubiquitinyl-[E2 ubiquitin-conjugating enzyme]-L-cysteine + [acceptor protein]-L-lysine = [E2 ubiquitin-conjugating enzyme]-L-cysteine + N(6)-ubiquitinyl-[acceptor protein]-L-lysine.</text>
            <dbReference type="EC" id="2.3.2.27"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="3">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the TRIM/RBCC family.</text>
    </comment>
    <dbReference type="EC" id="2.3.2.27" evidence="5"/>
    <dbReference type="AlphaFoldDB" id="A0A452GJ41"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001836.1">
        <property type="protein sequence ID" value="ENSGAGP00000001616.1"/>
        <property type="gene ID" value="ENSGAGG00000001264.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061630">
        <property type="term" value="F:ubiquitin protein ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd16598">
        <property type="entry name" value="RING-HC_TRIM26_C-IV"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12888">
        <property type="entry name" value="SPRY_PRY_TRIM7_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.920:FF:000004">
        <property type="entry name" value="Butyrophilin subfamily 1 member A1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.920">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001870">
        <property type="entry name" value="B30.2/SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043136">
        <property type="entry name" value="B30.2/SPRY_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003879">
        <property type="entry name" value="Butyrophylin_SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006574">
        <property type="entry name" value="PRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003877">
        <property type="entry name" value="SPRY_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050143">
        <property type="entry name" value="TRIM/RBCC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000315">
        <property type="entry name" value="Znf_B-box"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020457">
        <property type="entry name" value="Znf_B-box_chordata"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001510">
        <property type="entry name" value="Znf_PARP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001841">
        <property type="entry name" value="Znf_RING"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017907">
        <property type="entry name" value="Znf_RING_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24103">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE TRIM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13765">
        <property type="entry name" value="PRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00622">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00643">
        <property type="entry name" value="zf-B_box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15227">
        <property type="entry name" value="zf-C3HC4_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01406">
        <property type="entry name" value="BBOXZNFINGER"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01407">
        <property type="entry name" value="BUTYPHLNCDUF"/>
    </dbReference>
    <dbReference type="SMART" id="SM00336">
        <property type="entry name" value="BBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00589">
        <property type="entry name" value="PRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00184">
        <property type="entry name" value="RING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00449">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57845">
        <property type="entry name" value="B-box zinc-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57850">
        <property type="entry name" value="RING/U-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50188">
        <property type="entry name" value="B302_SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50119">
        <property type="entry name" value="ZF_BBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50064">
        <property type="entry name" value="ZF_PARP_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00518">
        <property type="entry name" value="ZF_RING_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50089">
        <property type="entry name" value="ZF_RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12 14" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0808">Transferase</keyword>
    <keyword evidence="10" id="KW-0833">Ubl conjugation pathway</keyword>
    <keyword evidence="11" id="KW-0862">Zinc</keyword>
    <keyword evidence="9 13" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="RING-type" evidence="16">
        <location>
            <begin position="16"/>
            <end position="59"/>
        </location>
    </feature>
    <feature type="domain" description="PARP-type" evidence="15">
        <location>
            <begin position="43"/>
            <end position="68"/>
        </location>
    </feature>
    <feature type="domain" description="B box-type" evidence="17">
        <location>
            <begin position="81"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="domain" description="B30.2/SPRY" evidence="18">
        <location>
            <begin position="256"/>
            <end position="450"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="14">
        <location>
            <begin position="179"/>
            <end position="206"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00008518"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00024"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50064"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50119"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50188"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001616.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="457" mass="52351" checksum="C97DF20C538F599C" modified="2019-05-08" version="1">MASSLSGLDMREEATCPICLEYLTDPVSVDCGHNFCRGCVTQYCVEWAGRGSLTCPICRSQILKGNLRPNWQLGNLVEKMKHVAVCTRHKKTLDLFCEEDEEAVCVVCERASQHRSHTVLLIEEAAQKYKVGNTWILRHLDKGPRPSREMRFRIPVRNSGAKQTQTERQKIVSEFQQLRQFLEEQERLLLAQLEKLDKEILRIQKENVTKLSGQISHLSELIHELEGKCQKPASEFLQVRLSLKHSRSMCRDRRAPESCPLESSSQRSQCNCLLHMTLDPDTAHPQLILSEDRKTVRYGRLWNKLPNTTKRFDYERCVLGSEGFTMGRHCWEVEVGERGGWAVGVARESLGRKGEISFKPKGGIWAVEYWEGQYWALTDPLTPLPLSWVPRRIWIYLDYEQGQVTFFNADNESLIFTFPPASFAGERIRPLFVVWETDLGSSLAHLATLTPSSLGWG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJ46</accession>
    <name>A0A452GJ46_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Protein transport protein sec16</fullName>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="16">Plays a role in the organization of the endoplasmic reticulum exit sites (ERES), also known as transitional endoplasmic reticulum (tER). Required for secretory cargo traffic from the endoplasmic reticulum to the Golgi apparatus.</text>
    </comment>
    <comment type="subunit">
        <text evidence="16">SEC16A and SEC16B are each present in multiple copies in a heteromeric complex.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Cytosol</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="5">Cytoplasm</location>
            <location evidence="5">Perinuclear region</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Endoplasmic reticulum membrane</location>
            <topology evidence="2">Peripheral membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Golgi apparatus membrane</location>
            <topology evidence="1">Peripheral membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Microsome membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="6 16">Belongs to the SEC16 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000001841.1">
        <property type="protein sequence ID" value="ENSGAGP00000001621.1"/>
        <property type="gene ID" value="ENSGAGG00000001287.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070971">
        <property type="term" value="C:endoplasmic reticulum exit site"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012507">
        <property type="term" value="C:ER to Golgi transport vesicle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000139">
        <property type="term" value="C:Golgi membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048471">
        <property type="term" value="C:perinuclear region of cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007029">
        <property type="term" value="P:endoplasmic reticulum organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007030">
        <property type="term" value="P:Golgi organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051668">
        <property type="term" value="P:localization within membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070973">
        <property type="term" value="P:protein localization to endoplasmic reticulum exit site"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016192">
        <property type="term" value="P:vesicle-mediated transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd09233">
        <property type="entry name" value="ACE1-Sec16-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.1030:FF:000002">
        <property type="entry name" value="Protein transport protein sec16"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.1030">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024340">
        <property type="entry name" value="Sec16_CCD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024298">
        <property type="entry name" value="Sec16_Sec23-bd"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13402:SF13">
        <property type="entry name" value="PROTEIN TRANSPORT PROTEIN SEC16A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13402">
        <property type="entry name" value="RGPR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12932">
        <property type="entry name" value="Sec16"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12931">
        <property type="entry name" value="TPR_Sec16"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="10 16" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="12 16" id="KW-0931">ER-Golgi transport</keyword>
    <keyword evidence="14 16" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="15 16" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-0492">Microsome</keyword>
    <keyword evidence="9" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="13 16" id="KW-0653">Protein transport</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7 16" id="KW-0813">Transport</keyword>
    <feature type="domain" description="Sec16 central conserved" evidence="19">
        <location>
            <begin position="1544"/>
            <end position="1642"/>
        </location>
    </feature>
    <feature type="domain" description="Sec16 Sec23-binding" evidence="18">
        <location>
            <begin position="1714"/>
            <end position="1948"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1"/>
            <end position="113"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="184"/>
            <end position="207"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="249"/>
            <end position="276"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="335"/>
            <end position="370"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="539"/>
            <end position="659"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="800"/>
            <end position="857"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1161"/>
            <end position="1247"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1309"/>
            <end position="1334"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1417"/>
            <end position="1471"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="2104"/>
            <end position="2129"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="2160"/>
            <end position="2309"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="2353"/>
            <end position="2463"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="58"/>
            <end position="70"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="91"/>
            <end position="113"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="339"/>
            <end position="370"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="17">
        <location>
            <begin position="554"/>
            <end position="572"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="595"/>
            <end position="604"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="612"/>
            <end position="626"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="638"/>
            <end position="647"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="815"/>
            <end position="828"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="840"/>
            <end position="857"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="1161"/>
            <end position="1171"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="17">
        <location>
            <begin position="1181"/>
            <end position="1227"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="17">
        <location>
            <begin position="1235"/>
            <end position="1247"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="17">
        <location>
            <begin position="1425"/>
            <end position="1446"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="17">
        <location>
            <begin position="1460"/>
            <end position="1470"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="17">
        <location>
            <begin position="2216"/>
            <end position="2230"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="17">
        <location>
            <begin position="2245"/>
            <end position="2266"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="2371"/>
            <end position="2382"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="2430"/>
            <end position="2449"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004395"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004406"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004514"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004524"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004556"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00005927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022848"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022892"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="RuleBase" id="RU364101"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF12931"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="Pfam" id="PF12932"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001621.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2463" mass="269155" checksum="59C02E5AF3BA20AA" modified="2019-05-08" version="1">MQPPPQTVPAGVGGPPPAGIARNTYWRNSPFSRRANAPVSGATAPVQPVTDPFAFGRQTPQGSLLGNPSKGNPLVMPGPAPSVFPRPAGVHSSQSYAGDNSHGLPTSLPTSVSQPGITSNNFSNVVTPSSPAHIINNTGEVHPNAEHGPHNSSAQSLYNTGAAHESSLSGHLAMAHVANRPLSRQDINRDPSNTSSVPTIAPFLPPPLQQTPPQWRPIRENLQPQVQNYWPYTEPPSQNAVYSVSHSSAVTSHLPPQADLHQGPVHQHIPQSTMQAPLPVGGEKNEISSFLVANHHVNSFQPENMFRQNIRVTNPWPSQPYQEQFYPQPLLPDSGFVNPITQENNSQKQSQNVSETPTGHTSTNADSGTLSMFFKGDEAENEEILSSEKKDLSSKANYDACQLNSGHIYHQPLPPQQAAASVLCQPQISTSSANEAVQKGMDVQYFSKTISSQHETQTSKNSMYISDDKANVSKARENVGSHCENVENLECIQNQEVLPSEPQTINRSSPSVGSDLYRYGSLPGQLLPKNAIVSHAERGPNLEAPDSLPHPVRSDSVSSNYSNISHKSVSSSTRPQELVGTFIQQESGKPDEESSASFFKQIDSSPLGGDSSEPNVSKNYHSNLSQPPTPSPPKPTGIFQTSANSSFEPVRSHGVGVKPAEVDQAKMVVELRENHPNQKNIKKSAAMPAASPGNLEQPPDNLETIFMPQVYPLPLTVTSEPGNILQPAGGPVVENIQSVPEKRPSTRAQGAAKKCESPATTLWAHNELPNFGGNVLLAPAAPAVYVPAKQTVEIIQPPEEGLSSQQPNKPECVPVQQSQHGNASSENLENPPKMGEEEALQSQASSGYASLLSSPPTDSLQNQPILIAQPNQSYNLAQPINFSLSLCNQLTRNENNLSLKDPGVEDKPVTGLQAPHAGGIPSGENVSLSVMQVGSSVNTPPSSNLSHSNLLQIPVNSSEITSNQSTNLLMQPPSQTPINLAPEGQKNANSEGCLPEFASKPGSNSAVSPGTNLPSGNASTLLVPPANTMVPNNNSTNDSHSHEEASGALDFTVTRTLDKSKTSNLIQMHNPLLSCGPVFPQQPVNHASQVGPNTHDKQHFYQQVTKDVQLQAPSDRVTQGALLSQQQKQTAQMLQTVPSGQSLVPSNYQMASGIKFTQALQQQENQVSTNRYYPAGPPEGSSTQPPTSYSQTSPDKQPFSGQSLSAPTSSASVTTSHPAMPTMQQEQNPPPPQTPQDACGPPQNPYYYYRHPYDAYQTPYPQPYPPLDPRTAAHLYYQDDVYGQYGPRYQHYDRSNAAYAEPGCYRYGEPERPSSRASQCSDRPPSRQGYTEDYYNPKSGWNDYYADYYANPYNYGDPNRWERYPSAYDSRYRDPRSYDQRYWYDTEQNPYQKREAYPYDNRQDRYEDHWRYDPRFAGSFDDETEPHRDPYGDEFDRRSVHSEHSAHSLRSSHSVHSRKSSFSSRSQQSQLYRSNHDLTANAYETTTQPVSLHTDYAYGGYPANFDGQQPFTDYGYSAESGWPSVEQVPSRPSTPEKFLVPHVCARFGPGGHLTKVLPNLPSEGQPALVEIHSMETMLQHMPEQEEMRAFPGPLAKDDTHKVDVINFAQNKVTQCFQNENLIDKESASLLWDFIVLLCRQNGTVVGTDIAELLLRDHKTVWLPGKSPNEANLIDFTNEALEQVEEESGEAQLSFLTNSLITTIDSLERETERFRELLLYGRKKDALESAMKHGLWGHALLLASKMDNRTHARVMTRFANSLPINDPLQTVYQLMSGRMPAASTCCGDEKWGDWRPHLAMVLSNLTNNMEVESRTITTMGDTLASKGLLDAAHFCYLMAQIGFGVYTKKTTKLVLIGSNHSLPFLKFATNEAIQRTEAYEYAQSLGTQPCCLPNFQVFKFIYACRLAEMGLAAQAFHYCEVISKTVLKNPYYYSPVLISQLIQISSQLRLFDPQIKEKPEQELFIEPSWLVRLRHLDGQIKDGTIAYNADRSTPQQYACSTPSSELDHISQCDGTGSGQEMGPATENPLLTSLLPNTMHPMQGVQLMPSAPQTILEGSAAMTPPTQQETVGGVPFYPVAPPSIGPGSGFAPPGFPNQYGAEQSSLYLGSTVPPGGPPPQAAEPQPEGHLNQETAMQRIPQESPIQKTFPEQREEDFYGKMANMAPGRRSRSTSQSSAHMGYGRRSRTTSESSTHSIGRERCNSAAKQPSPPPPSIPEGKETNKEIKKEPAARKSGANWFRWLMGKGKNEAHLPDDKNKSIVWDEKKQRWVNLDEPEEERKPPPPPPAGFPKALQTAPPGPGGPPGVNMFSRKAAGTRARYVDVLNPSGAKSSGAVPAPSDLFAPLAPMPIPTNLFLPNSVPEEQQPMEGSEAQEQTLSANQVSADAATEPQYLNSTMLPPGSELPGSNPDGSQSGEPLCSVPPSGGPPSGTVQLYNPSQFAQSPATTRSSRLGRIGQRKYPTLK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJ47</accession>
    <name>A0A452GJ47_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Rap guanine nucleotide exchange factor 2</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="21">Cyclic nucleotide ras GEF</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="23">Neural RAP guanine nucleotide exchange protein</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="20">PDZ domain-containing guanine nucleotide exchange factor 1</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="19">RA-GEF-1</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="22">Ras/Rap1-associating GEF-1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="31 32">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="32" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
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                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
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        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
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        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
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        <citation type="submission" date="2025-09" db="UniProtKB">
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        </source>
    </evidence>
    <sequence length="1558" mass="174611" checksum="5094ADF7AFE8F0A4" modified="2019-05-08" version="1">MCETVRYERHEANEVLYYPDDIGSCWYILLSGSVFIKESMFLPRSSFGKRSAGSFRRGCECIVLEPSEMIVVDYMDENEEYFQRQASHRQSRRRFRKINQKGERQTIIDTVDPYPVGKPPLPRGYHTECTKPQLPADFTKLHLTDSLHPQVTHVTSSHSGCSITSDSGSSSLSDIYQATESEAGDMDLSGLPETAVDSEDDDDEEDIERASDPLMSRDIVRDCLEKDPIDRTDDDIEQLLEFMHQLPAFANMTMSVRRELCAVMVFAVVERAGTIVLNDGEELDSWSVILNGSVEVTYPDGRTEILCMGNSFGVSPTMDKEYMKGVMRTKVDDCQFVCIAQQDYCRILNQVEKNMQKVEEEGEIVMVKEHRELDRTGTRKGHIVIKGTSERLTMHLVEEHSVVDPTFIEDFLLTYRTFLSSPMEVGKKLLEWFNDPSLRDKVTRVVLLWVNNHFNDFEGDPAMTRFLEEFENNLEREKMGGHLRLLNIACAAKAKRRLITLTKPSREAPLPFILLGGSEKGFGIFVDSVDLGSKATEAGLKRGDQILEVNGQNFENIQLSKAMEILRNNTHLSITVKTNLFVFKELLTRLSEEKRNGAPHLPKIGDIKKASRYSIPDLAVDVEQVIGLEKVNKKSKANTVGGRNKLKKILDKTRISILPQKPYNDIGIGQSQDDSIVGLRQTKHIPPALPVSGTLSSSNPDLLQSHHRILDFNSTPDLPDQVLRVFKADQQSRYIMISKDTTAKEVVIQAIREFAVTATPDAYSLCEVSVTPEGVIKQRRLPDQLSKLADRIQLSGRYYLKNNMETETLCSDEDAQELLRESQISLLQLSTVEMAAQLSMRNFELFRNIEPTEYIDDLFKLKSKTSCANLKKFEEVINQETFWVASEILRETNQLKRMKIIKHFIKIALHCRECKNFNSMFAIISGLNLAPVARLRTTWEKLPSKYEKLFQDLQDLFDPSRNMAKYRNVLNSQNLQPPIIPLFPVIKKDLTFLHEGNDSKVEGLVNFEKLRMIAKEIRHVGRMASVNMDPAVMFRTRSLSQSSTNAAVLDVAQTGGHKKRVRRSSFLNAKKLYEDAQMARKVKQYLSNLDLEMDEESLQTLSLQCEPATNTLPKNPGDKRSGKQEMSPVAPRAGIQQKVQQQHKINQALQVPAVSLYPSRKKVPVKDLPPFGINSPQALKKILSLSEEGSLDRHKKQAEDTISNASSQLSSPPTSPHTSPRKGYTLAPSGTVDNFSDSGRSEISSRSSIVSNSSFDSMPVSLHDERRQRHSVSIVETNLGVGRIDRRAMIEPDQYSLGSYAPLSENRGLYAGATVLSSPSTEELSQDQGDRTSLDAADSGRGSWTSCSSGSHDNIQTIQHQRSWETLPFGHVHFDSSGDGAGLWASGSHMDQMMFPDHGTKYGRQSQGREGLDQAQSRASWASSTGYWGEDSEGDTGTIKRRGGKDVSIEAETSSITSVTAEESKPAPMSSHITVSSSSAKGLIARKEGRYREPPPTPPGYVGIPIADFAEGVSHPTRKPPDYNVALQRSRMVARTCETHGTSTLQQQSQEDEQVSAV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJ48</accession>
    <name>A0A452GJ48_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">lysophospholipase</fullName>
            <ecNumber evidence="3">3.1.1.5</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>1-(9Z-octadecenoyl)-sn-glycero-3-phosphocholine + H2O = sn-glycerol 3-phosphocholine + (9Z)-octadecenoate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:40807"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16870"/>
            <dbReference type="ChEBI" id="CHEBI:28610"/>
            <dbReference type="ChEBI" id="CHEBI:30823"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="13">
            <dbReference type="Rhea" id="RHEA:40808"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="14">
            <text>1-hexadecanoyl-sn-glycero-3-phosphate + H2O = sn-glycerol 3-phosphate + hexadecanoate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:49092"/>
            <dbReference type="ChEBI" id="CHEBI:7896"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57518"/>
            <dbReference type="ChEBI" id="CHEBI:57597"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="14">
            <dbReference type="Rhea" id="RHEA:49093"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="16">
            <text>1-hexadecanoyl-sn-glycero-3-phosphocholine + H2O = sn-glycerol 3-phosphocholine + hexadecanoate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:40435"/>
            <dbReference type="ChEBI" id="CHEBI:7896"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16870"/>
            <dbReference type="ChEBI" id="CHEBI:72998"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="16">
            <dbReference type="Rhea" id="RHEA:40436"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>a 1-acyl-sn-glycero-3-phosphocholine + H2O = sn-glycerol 3-phosphocholine + a fatty acid + H(+)</text>
            <dbReference type="Rhea" id="RHEA:15177"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16870"/>
            <dbReference type="ChEBI" id="CHEBI:28868"/>
            <dbReference type="ChEBI" id="CHEBI:58168"/>
            <dbReference type="EC" id="3.1.1.5"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="15">
            <dbReference type="Rhea" id="RHEA:15178"/>
        </physiologicalReaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum membrane</location>
            <topology evidence="1">Single-pass type III membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the NTE family.</text>
    </comment>
    <comment type="caution">
        <text evidence="17">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="EC" id="3.1.1.5" evidence="3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001931.1">
        <property type="protein sequence ID" value="ENSGAGP00000001696.1"/>
        <property type="gene ID" value="ENSGAGG00000001350.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004622">
        <property type="term" value="F:phosphatidylcholine lysophospholipase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016042">
        <property type="term" value="P:lipid catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046470">
        <property type="term" value="P:phosphatidylcholine metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00038">
        <property type="entry name" value="CAP_ED"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="CDD" id="cd07225">
        <property type="entry name" value="Pat_PNPLA6_PNPLA7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.10:FF:000010">
        <property type="entry name" value="neuropathy target esterase isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.10:FF:000012">
        <property type="entry name" value="neuropathy target esterase isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.1090.10:FF:000022">
        <property type="entry name" value="Neuropathy target esterase sws"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.1090.10:FF:000020">
        <property type="entry name" value="neuropathy target esterase-like isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.10:FF:000022">
        <property type="entry name" value="Patatin like phospholipase domain containing 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.1090.10">
        <property type="entry name" value="Cytosolic phospholipase A2 catalytic domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.10">
        <property type="entry name" value="Jelly Rolls"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016035">
        <property type="entry name" value="Acyl_Trfase/lysoPLipase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000595">
        <property type="entry name" value="cNMP-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018490">
        <property type="entry name" value="cNMP-bd_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001423">
        <property type="entry name" value="LysoPLipase_patatin_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050301">
        <property type="entry name" value="NTE"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056556">
        <property type="entry name" value="NTE1_P-loop_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002641">
        <property type="entry name" value="PNPLA_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014710">
        <property type="entry name" value="RmlC-like_jellyroll"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14226">
        <property type="entry name" value="NEUROPATHY TARGET ESTERASE/SWISS CHEESE D.MELANOGASTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14226:SF26">
        <property type="entry name" value="PATATIN-LIKE PHOSPHOLIPASE DOMAIN-CONTAINING PROTEIN 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00027">
        <property type="entry name" value="cNMP_binding"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24179">
        <property type="entry name" value="NTE_Ploop"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01734">
        <property type="entry name" value="Patatin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00100">
        <property type="entry name" value="cNMP"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51206">
        <property type="entry name" value="cAMP-binding domain-like"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52151">
        <property type="entry name" value="FabD/lysophospholipase-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50042">
        <property type="entry name" value="CNMP_BINDING_3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51635">
        <property type="entry name" value="PNPLA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01237">
        <property type="entry name" value="UPF0028"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="7 17" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="9 17" id="KW-0442">Lipid degradation</keyword>
    <keyword evidence="11 17" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="12 19" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5 19" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="10 19" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="14"/>
            <end position="36"/>
        </location>
    </feature>
    <feature type="domain" description="Cyclic nucleotide-binding" evidence="20">
        <location>
            <begin position="151"/>
            <end position="278"/>
        </location>
    </feature>
    <feature type="domain" description="Cyclic nucleotide-binding" evidence="20">
        <location>
            <begin position="471"/>
            <end position="575"/>
        </location>
    </feature>
    <feature type="domain" description="Cyclic nucleotide-binding" evidence="20">
        <location>
            <begin position="603"/>
            <end position="708"/>
        </location>
    </feature>
    <feature type="domain" description="PNPLA" evidence="21">
        <location>
            <begin position="939"/>
            <end position="1106"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="313"/>
            <end position="349"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="403"/>
            <end position="429"/>
        </location>
    </feature>
    <feature type="short sequence motif" description="GXGXXG" evidence="17">
        <location>
            <begin position="943"/>
            <end position="948"/>
        </location>
    </feature>
    <feature type="short sequence motif" description="GXSXG" evidence="17">
        <location>
            <begin position="970"/>
            <end position="974"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="403"/>
            <end position="418"/>
        </location>
    </feature>
    <feature type="active site" description="Nucleophile" evidence="17">
        <location>
            <position position="972"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="17">
        <location>
            <position position="1093"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004643"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006636"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00013274"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022963"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00047314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00048133"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00048454"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00048656"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01161"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS51635"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001696.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1361" mass="152759" checksum="43EC8F04BDF381A8" modified="2019-05-08" version="1">MGQSESELQDEQEVLGVMIGVGVSMLLIATLILVLVRRLRHKKVQTQETPKYRFRKRDKVLFYGRKIMRKVSQSTSSLVDTTISNTSRPRMKKKLKMLNIAKKILRIKKELPILQLKEPPPSVLEADLTEFDVANSHLPSEVLYMLKNVRVLGHFEKPLFLELCKHMIFQQFQQGEYVFRPGQPDSSIYVVQDGKLELFLTQQDGKETLVKEVFPGDSVHSLLSILDVITGHQRPYRTVSARAGEDSTVLRLPVEAFSAVFEKYPESLVRVVQIIMVRLQRVTFLALHNYLGLTNELFSHEMQLLRLFPQPNHASRTSPVRHNKRVISISSIDDHKESSGRQPDASAKDLADQLKLGSLEPSSVPLLSRCISMPVDISGLQKGPRSDFDMAYERGRISVSLQEESTGTMASFSQSVSQEPKERRSVTVEEQSPGTYHYNYCEDDSAGGACPFGPYQGRQTSAIFEAAKRELVKLMKIEDPSLLNNRVLLHHAKAGTVIARQGDQDVSLHFVLWGCLHVYQRMIDKAEDVCLFLTQPGEMVGQLAVLTGEPLIFTIKANRDCTFLRISKSDFYEIMREQPSVVLSVAHTVAARMSPFVRQMDFAIDWMAVEAGRALYRQGDKSDCTYIVLNGRLRSVIQKGNGKKELVGEYGRGDLIGVVEALTRQPRATTVHAVRDTELAKLPEGTLNNIKRRYPQVVTRLIHLLSQKILGNLQQLCGPFTGSSLGVASSSELTNPASNLSTVAVLPVCDDVPMAAFTLELEHALNAIGPTLLLTSDIIRARLGASALDSIQEYRLSGWLAQQEDIHRIVLYQTDCTLTPWTVRCIRQADCILIVGLGDQEPALGKLEQMLENTAVRALKQLILLHREDGPSPSRTVEWLNMRSWCSGHLHIKCPRRVFSRRSPNKLREMYAKVFEKDADRHSDFSRLARVLTGNSIALVLGGGGARGCSHIGVIKAMEEAGIPIDLIGGTSIGSFIGALYAEERSAVRTKQRAREWAKSMNSVFETVLDLTYPITSMFSGSAFNTSINKVFQDKQIEDLWLPYFNVTTDITASAMRVHKDAGSVWRYIRASASYPPYLPPLCDPKDSHLLVDGCYVNNVPGSLWRYVRASMTLSGYLPPLCDPKDGNLLMDGGYINNLPADIARNMGAKTVVAIDVGSQDETDLCNYGDSLSGWWLLWKRLNPWAQKVKVPDMAEIQSRLAYVSCVRQLEVVKSSSYCEYIRPPIDRFKTMDFGKFDEIYDVGYQHGKVVFSGWCRGDIIEKMVKDRRLADFYESKRMDVLTCPSAGFTDLAEIVSRIEPAKSYLSDGYADGEESDYLTEYEEEGLDPARGEEELLAPAEWIHAEDEEKSLRHRKSLPQE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJ50</accession>
    <name>A0A452GJ50_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Rap guanine nucleotide exchange factor 2</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="21">Cyclic nucleotide ras GEF</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="23">Neural RAP guanine nucleotide exchange protein</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="20">PDZ domain-containing guanine nucleotide exchange factor 1</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="19">RA-GEF-1</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="22">Ras/Rap1-associating GEF-1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="31 32">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="32" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="31" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="31" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell junction</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Perinuclear region</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Late endosome</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the RAPGEF2 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000001921.1">
        <property type="protein sequence ID" value="ENSGAGP00000001689.1"/>
        <property type="gene ID" value="ENSGAGG00000001344.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070161">
        <property type="term" value="C:anchoring junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005770">
        <property type="term" value="C:late endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048471">
        <property type="term" value="C:perinuclear region of cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007264">
        <property type="term" value="P:small GTPase-mediated signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00038">
        <property type="entry name" value="CAP_ED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd06755">
        <property type="entry name" value="PDZ_RapGEF2_RapGEF6-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd01785">
        <property type="entry name" value="RA_PDZ-GEF1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00155">
        <property type="entry name" value="RasGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd06224">
        <property type="entry name" value="REM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.10:FF:000008">
        <property type="entry name" value="Rap guanine nucleotide exchange factor (GEF) 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.840.10:FF:000001">
        <property type="entry name" value="Rap guanine nucleotide exchange factor (GEF) 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000024">
        <property type="entry name" value="rap guanine nucleotide exchange factor 2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.870.10:FF:000001">
        <property type="entry name" value="rap guanine nucleotide exchange factor 2 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.42.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.10">
        <property type="entry name" value="Jelly Rolls"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.840.10">
        <property type="entry name" value="Ras guanine-nucleotide exchange factors catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.870.10">
        <property type="entry name" value="Son of sevenless (SoS) protein Chain: S domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000595">
        <property type="entry name" value="cNMP-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018490">
        <property type="entry name" value="cNMP-bd_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001478">
        <property type="entry name" value="PDZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036034">
        <property type="entry name" value="PDZ_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000159">
        <property type="entry name" value="RA_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000651">
        <property type="entry name" value="Ras-like_Gua-exchang_fac_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023578">
        <property type="entry name" value="Ras_GEF_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001895">
        <property type="entry name" value="RASGEF_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036964">
        <property type="entry name" value="RASGEF_cat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014710">
        <property type="entry name" value="RmlC-like_jellyroll"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029071">
        <property type="entry name" value="Ubiquitin-like_domsf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45161">
        <property type="entry name" value="CYTOSKELETON-ASSOCIATED PROTEIN 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45161:SF2">
        <property type="entry name" value="RAP GUANINE NUCLEOTIDE EXCHANGE FACTOR 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00595">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00788">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00617">
        <property type="entry name" value="RasGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00618">
        <property type="entry name" value="RasGEF_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00100">
        <property type="entry name" value="cNMP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00228">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00314">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00147">
        <property type="entry name" value="RasGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00229">
        <property type="entry name" value="RasGEFN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51206">
        <property type="entry name" value="cAMP-binding domain-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50156">
        <property type="entry name" value="PDZ domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48366">
        <property type="entry name" value="Ras GEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54236">
        <property type="entry name" value="Ubiquitin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50042">
        <property type="entry name" value="CNMP_BINDING_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50106">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50200">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50009">
        <property type="entry name" value="RASGEF_CAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50212">
        <property type="entry name" value="RASGEF_NTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="17" id="KW-0965">Cell junction</keyword>
    <keyword evidence="9" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="10" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="14" id="KW-0221">Differentiation</keyword>
    <keyword evidence="13" id="KW-0967">Endosome</keyword>
    <keyword evidence="7" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="12 24" id="KW-0344">Guanine-nucleotide releasing factor</keyword>
    <keyword evidence="18" id="KW-0472">Membrane</keyword>
    <keyword evidence="16" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="11" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="32" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="15" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="Cyclic nucleotide-binding" evidence="27">
        <location>
            <begin position="242"/>
            <end position="342"/>
        </location>
    </feature>
    <feature type="domain" description="N-terminal Ras-GEF" evidence="30">
        <location>
            <begin position="374"/>
            <end position="487"/>
        </location>
    </feature>
    <feature type="domain" description="PDZ" evidence="28">
        <location>
            <begin position="492"/>
            <end position="562"/>
        </location>
    </feature>
    <feature type="domain" description="Ras-associating" evidence="29">
        <location>
            <begin position="713"/>
            <end position="799"/>
        </location>
    </feature>
    <feature type="domain" description="Ras-GEF" evidence="26">
        <location>
            <begin position="824"/>
            <end position="1051"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="25">
        <location>
            <begin position="175"/>
            <end position="206"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="25">
        <location>
            <begin position="1106"/>
            <end position="1146"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="25">
        <location>
            <begin position="1191"/>
            <end position="1271"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="25">
        <location>
            <begin position="1320"/>
            <end position="1355"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="25">
        <location>
            <begin position="1397"/>
            <end position="1483"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="25">
        <location>
            <begin position="1539"/>
            <end position="1560"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="25">
        <location>
            <begin position="190"/>
            <end position="201"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="25">
        <location>
            <begin position="1106"/>
            <end position="1115"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="25">
        <location>
            <begin position="1137"/>
            <end position="1146"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="25">
        <location>
            <begin position="1205"/>
            <end position="1220"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="25">
        <location>
            <begin position="1237"/>
            <end position="1259"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="25">
        <location>
            <begin position="1320"/>
            <end position="1329"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="25">
        <location>
            <begin position="1344"/>
            <end position="1355"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="25">
        <location>
            <begin position="1405"/>
            <end position="1428"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="25">
        <location>
            <begin position="1453"/>
            <end position="1463"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="25">
        <location>
            <begin position="1473"/>
            <end position="1482"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="25">
        <location>
            <begin position="1541"/>
            <end position="1551"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004282"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004556"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004603"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00010829"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00016709"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022658"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00029925"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00030673"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00031545"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00031980"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00032021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00168"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="26">
        <source>
            <dbReference type="PROSITE" id="PS50009"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="27">
        <source>
            <dbReference type="PROSITE" id="PS50042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="28">
        <source>
            <dbReference type="PROSITE" id="PS50106"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="29">
        <source>
            <dbReference type="PROSITE" id="PS50200"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="30">
        <source>
            <dbReference type="PROSITE" id="PS50212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="31">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001689.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="32">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1560" mass="174909" checksum="BA2972176A464AD2" modified="2019-05-08" version="1">MCETVRYERHEANEVLYYPDDIGSCWYILLSGSVFIKESMFLPRSSFGKRSAGSFRRGCECIVLEPSEMIVVDYMDENEEYFQRQASHRQSRRRFRKINQKGERQTIIDTVDPYPVGKPPLPRGYHTLPADFTKLHLTDSLHPQVTHVTSSHSGCSITSDSGSSSLSDIYQATESEAGDMDLSGLPETAVDSEDDDDEEDIERASDPLMSRDIVRDCLEKDPIDRTDDDIEQLLEFMHQLPAFANMTMSVRRELCAVMVFAVVERAGTIVLNDGEELDSWSVILNGSVEVTYPDGRTEILCMGNSFGVSPTMDKEYMKGVMRTKVDDCQFVCIAQQDYCRILNQVEKNMQKVEEEGEIVMVKEHRELDRTGTRKGHIVIKGTSERLTMHLVEEHSVVDPTFIEDFLLTYRTFLSSPMEVGKKLLEWFNDPSLRDKVTRVVLLWVNNHFNDFEGDPAMTRFLEEFENNLEREKMGGHLRLLNIACAAKAKRRLITLTKPSREAPLPFILLGGSEKGFGIFVDSVDLGSKATEAGLKRGDQILEVNGQNFENIQLSKAMEILRNNTHLSITVKTNLFVFKELLTRLSEEKRNGAPHLPKIGDIKKASRYSIPDLAVDVEQVIGLEKVNKKSKANTVGGRNKLKKILDKTRISILPQKPYNDIGIGQSQDDSIVGLRQTKHIPPALPVSGTLSSSNPDLLQSHHRILDFNSTPDLPDQVLRVFKADQQSRYIMISKDTTAKEVVIQAIREFAVTATPDAYSLCEVSVTPEGVIKQRRLPDQLSKLADRIQLSGRYYLKNNMETETLCSDEDAQELLRESQISLLQLSTVEMAAQLSMRNFELFRNIEPTEYIDDLFKLKSKTSCANLKKFEEVINQETFWVASEILRETNQLKRMKIIKHFIKIALHCRECKNFNSMFAIISGLNLAPVARLRTTWEKLPSKYEKLFQDLQDLFDPSRNMAKYRNVLNSQNLQPPIIPLFPVIKKDLTFLHEGNDSKVEGLVNFEKLRMIAKEIRHVGRMASVNMDPAVMFRTRKKKWRSLGSLSQSSTNAAVLDVAQTGGHKKRVRRSSFLNAKKLYEDAQMARKVKQYLSNLDLEMDEESLQTLSLQCEPATNTLPKNPGDKRSGKQEMSPVAPRAGIQQKVQQQHKINQALQVPAVSLYPSRKKVPVKDLPPFGINSPQALKKILSLSEEGSLDRHKKQAEDTISNASSQLSSPPTSPHTSPRKGYTLAPSGTVDNFSDSGRSEISSRSSIVSNSSFDSMPVSLHDERRQRHSVSIVETNLGVGRIDRRAMIEPDQYSLGSYAPLSENRGLYAGATVLSSPSTEELSQDQGDRTSLDAADSGRGSWTSCSSGSHDNIQTIQHQRSWETLPFGHVHFDSSGDGAGLWASGSHMDQMMFPDHGTKYGRQSQGREGLDQAQSRASWASSTGYWGEDSEGDTGTIKRRGGKDVSIEAETSSITSVTAEESKPAPMSSHITVSSSSAKGLIARKEGRYREPPPTPPGYVGIPIADFAEGVSHPTRKPPDYNVALQRSRMVARTCETHGTSTLQQQSQEDEQVSAV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJ51</accession>
    <name>A0A452GJ51_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Pumilio homolog 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">P-body</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasmic granule</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJ51"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001846.1">
        <property type="protein sequence ID" value="ENSGAGP00000001626.1"/>
        <property type="gene ID" value="ENSGAGG00000001290.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000932">
        <property type="term" value="C:P-body"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003730">
        <property type="term" value="F:mRNA 3'-UTR binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035196">
        <property type="term" value="P:miRNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043488">
        <property type="term" value="P:regulation of mRNA stability"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006417">
        <property type="term" value="P:regulation of translation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd07920">
        <property type="entry name" value="Pumilio"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000004">
        <property type="entry name" value="Pumilio homolog 1 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033133">
        <property type="entry name" value="PUM-HD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033712">
        <property type="entry name" value="Pumilio_RNA-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001313">
        <property type="entry name" value="Pumilio_RNA-bd_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12537:SF1">
        <property type="entry name" value="PUMILIO HOMOLOG 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12537">
        <property type="entry name" value="RNA BINDING PROTEIN PUMILIO-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00806">
        <property type="entry name" value="PUF"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SMART" id="SM00025">
        <property type="entry name" value="Pumilio"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50302">
        <property type="entry name" value="PUM"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50303">
        <property type="entry name" value="PUM_HD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="5" id="KW-0810">Translation regulation</keyword>
    <feature type="domain" description="PUM-HD" evidence="10">
        <location>
            <begin position="715"/>
            <end position="1057"/>
        </location>
    </feature>
    <feature type="repeat" description="Pumilio" evidence="8">
        <location>
            <begin position="735"/>
            <end position="770"/>
        </location>
    </feature>
    <feature type="repeat" description="Pumilio" evidence="8">
        <location>
            <begin position="771"/>
            <end position="806"/>
        </location>
    </feature>
    <feature type="repeat" description="Pumilio" evidence="8">
        <location>
            <begin position="807"/>
            <end position="844"/>
        </location>
    </feature>
    <feature type="repeat" description="Pumilio" evidence="8">
        <location>
            <begin position="845"/>
            <end position="880"/>
        </location>
    </feature>
    <feature type="repeat" description="Pumilio" evidence="8">
        <location>
            <begin position="881"/>
            <end position="916"/>
        </location>
    </feature>
    <feature type="repeat" description="Pumilio" evidence="8">
        <location>
            <begin position="917"/>
            <end position="952"/>
        </location>
    </feature>
    <feature type="repeat" description="Pumilio" evidence="8">
        <location>
            <begin position="953"/>
            <end position="988"/>
        </location>
    </feature>
    <feature type="repeat" description="Pumilio" evidence="8">
        <location>
            <begin position="992"/>
            <end position="1031"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="119"/>
            <end position="172"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="376"/>
            <end position="411"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="500"/>
            <end position="538"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="629"/>
            <end position="660"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="136"/>
            <end position="163"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="376"/>
            <end position="389"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="398"/>
            <end position="411"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="513"/>
            <end position="538"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="650"/>
            <end position="660"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004201"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004463"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022845"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00040564"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00317"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001626.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1075" mass="114810" checksum="55E69F3C72935C98" modified="2019-05-08" version="1">MVLLDHLVRSMDELNHDFQALALEGRAMGEQLLPGKKFWETDDSSKDGPKGIFLGDQWRDSAWGTSDHSVSQPIMVQRRPGQGFHVNSDVNSVLSPRSESGGLGVSMVEYVLSSSPGDSCLRKGGFGPRDAENDENDKSDKKNKGTFDGDKLGDLKEEGDVMDKTNGLPVQNGIDADVKDFSRTPGNCQNSANEVDLLGPNQNGSEGLAQLASTNGAKPVEDFSNMESQSVPLDPMEHVGMEPLQFDYSGTQVPVDSAAATVGLFDYNSQPQLFQRPNALAVQQLTAAQQQQYALAAAHQPHIAGLAPAAFVPNPYIISAAPPGTDPYAAGLAAAATLGPAVVPHQYYGVTPWGVYPASLFQQQAAAAAAASNSANQQTTQQTQQGQQQVLRGGASQRPLTPNQNQQGQQTDPLVAAAAVNSALAFGQGLAAGMPGYPMLAPAAYYDQTGALVVNAGARNGLGAPVRLVAPAPVIISSSAAQAAVAAAAASANGAAGGLAGTTNGPFRPLGTQQPQPQPQQQPTNNLASSSFYGNNSLSSNSQSSSLFSQGSAQAANTSLGFGSSSSLGATLGSALGGFGTAVANSNTGSGSRRDSLTGSSDLYKRTSSSLTPIGHSFYNGLGFSSSPGPVGMPLPSQGPGHSQTPPPSLSSHGSSSSLNLGGLTNGSGRYISAAPGAEAKYRSASSASSLFSPSSTLFPSSRLRYGMSDVMPSGRSRLLEDFRNNRYPNLQLREIAGHIMEFSQDQHGSRFIQLKLERATPAERQLVFNEILQAAYQLMVDVFGNYVIQKFFEFGSLDQKLALAERIRGHVLSLALQMYGCRVIQKALEFIPPDQQVINEMVRELDGHVLKCVKDQNGNHVVQKCIECVQPQSLQFIIDAFKGQVFALSTHPYGCRVIQRILEHCLPEQTLPILEELHQHTEQLVQDQYGNYVIQHVLEHGRPEDKSKIVAEIRGNVLVLSQHKFASNVVEKCVTHASRTERAMLIDEVCTMNDGPHSALYTMMKDQYANYVVQKMIDVAEPAQRKIVMHKIRPHIATLRKYTYGKHILAKLEKYYMKNGVDLGPICGPPNGII</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJ57</accession>
    <name>A0A452GJ57_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">DNA ligase</fullName>
            <ecNumber evidence="15">6.5.1.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">DNA ligase that seals nicks in double-stranded during DNA repair. Also involved in DNA replication and DNA recombination.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13 15">
            <text>ATP + (deoxyribonucleotide)n-3'-hydroxyl + 5'-phospho-(deoxyribonucleotide)m = (deoxyribonucleotide)n+m + AMP + diphosphate.</text>
            <dbReference type="EC" id="6.5.1.1"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 16">Belongs to the ATP-dependent DNA ligase family.</text>
    </comment>
    <dbReference type="EC" id="6.5.1.1" evidence="15"/>
    <dbReference type="AlphaFoldDB" id="A0A452GJ57"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001851.1">
        <property type="protein sequence ID" value="ENSGAGP00000001631.1"/>
        <property type="gene ID" value="ENSGAGG00000000751.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003910">
        <property type="term" value="F:DNA ligase (ATP) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071897">
        <property type="term" value="P:DNA biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006310">
        <property type="term" value="P:DNA recombination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006281">
        <property type="term" value="P:DNA repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903461">
        <property type="term" value="P:Okazaki fragment processing involved in mitotic DNA replication"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07900">
        <property type="entry name" value="Adenylation_DNA_ligase_I_Euk"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd07969">
        <property type="entry name" value="OBF_DNA_ligase_I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.3260.10:FF:000001">
        <property type="entry name" value="DNA ligase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.50.140:FF:000062">
        <property type="entry name" value="DNA ligase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.470.30:FF:000016">
        <property type="entry name" value="DNA ligase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1490.70">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.3260.10">
        <property type="entry name" value="DNA ligase, ATP-dependent, N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.470.30">
        <property type="entry name" value="DNA ligase/mRNA capping enzyme"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.50.140">
        <property type="entry name" value="Nucleic acid-binding proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050191">
        <property type="entry name" value="ATP-dep_DNA_ligase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000977">
        <property type="entry name" value="DNA_ligase_ATP-dep"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012309">
        <property type="entry name" value="DNA_ligase_ATP-dep_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012310">
        <property type="entry name" value="DNA_ligase_ATP-dep_cent"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016059">
        <property type="entry name" value="DNA_ligase_ATP-dep_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012308">
        <property type="entry name" value="DNA_ligase_ATP-dep_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036599">
        <property type="entry name" value="DNA_ligase_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012340">
        <property type="entry name" value="NA-bd_OB-fold"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00574">
        <property type="entry name" value="dnl1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45674:SF4">
        <property type="entry name" value="DNA LIGASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45674">
        <property type="entry name" value="DNA LIGASE 1/3 FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04679">
        <property type="entry name" value="DNA_ligase_A_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01068">
        <property type="entry name" value="DNA_ligase_A_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04675">
        <property type="entry name" value="DNA_ligase_A_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF117018">
        <property type="entry name" value="ATP-dependent DNA ligase DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56091">
        <property type="entry name" value="DNA ligase/mRNA capping enzyme, catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50249">
        <property type="entry name" value="Nucleic acid-binding proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00697">
        <property type="entry name" value="DNA_LIGASE_A1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00333">
        <property type="entry name" value="DNA_LIGASE_A2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50160">
        <property type="entry name" value="DNA_LIGASE_A3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 15" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="12" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="4" id="KW-0132">Cell division</keyword>
    <keyword evidence="7 15" id="KW-0227">DNA damage</keyword>
    <keyword evidence="9 15" id="KW-0233">DNA recombination</keyword>
    <keyword evidence="10 15" id="KW-0234">DNA repair</keyword>
    <keyword evidence="5" id="KW-0235">DNA replication</keyword>
    <keyword evidence="3 15" id="KW-0436">Ligase</keyword>
    <keyword evidence="6 15" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="ATP-dependent DNA ligase family profile" evidence="18">
        <location>
            <begin position="548"/>
            <end position="684"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="66"/>
            <end position="87"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="17">
        <location>
            <begin position="74"/>
            <end position="87"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007572"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022598"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022705"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023172"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00034003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00058910"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="RuleBase" id="RU000617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="RuleBase" id="RU004196"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50160"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001631.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="816" mass="91359" checksum="A07F08DC66E41A0E" modified="2019-05-08" version="1">MLSVTQCCSCFCTPALVHLWTPHPSFSHNLSWSKDSGLAPETPPSSLQEADSNLCSSARKQIPKRKIELSTSRENNRHEEPSAKRCREEQGKSLGACLIHAIIVPVATCAGLPSLGVLVQANCWETLRNTSAVGPVYWTWWLPISLWLCDPSSFSKPVVPLDPMEYNPSKSSYHPVQDACWGHGQRVPYLAVARTFERIEEVSARLKNIETLSNLFRSVVALSPSDILPCIYLCLNRLGPAYKGLELGIGETILMKAVAQAMGRQLDQIKAEAQEKGDLGLVAESSRSNQRTMFTPPKLSAAGVFSKLQEIARMTGSASMNKKIDIIKGLFVACRHSEARYITRSLGGKLRIGLAEQSVLSAIAHAVSLTPPGQGFPPEVLDAGQSKSAEARKAWLEEQTQILKQTFCELPSYDIIVPILLEHGVESLPQHCKITPGIPLKPMLAQPTKGIGEVLKRFEEAAFTCEYKYDGERAQIHILENGDVHIYSRNQEDNTSKYPDIISRIPKVKKDSVQSCILDAEAVAWDPEKRQIQPFQVLTTRKRKDVEAAEIKVQVCLYAFDILYLNGESLVKEPFSRRRQLLHESFVETEGEFLFATSMDTCSLDEISEFLDQSIKDSCEGLMVKTLEVDATYEIAKRSHKWLKLKKDYLEGVGDTLDLVVIGAYLGKGKRVGMYGGFLLACYDEESEEYQSICKVGSVGSLHRLDAHLQQTARPKPRPYYRWDSSTEPNHWLAAVQVWEVKCADLSISPVHKAAIGLVDEEKGISLRFPRFLRVRDDKKPEEATTSSQVAELYRRQQQIQNQQLVEKAEEEEDFY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJ58</accession>
    <name>A0A452GJ58_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Rap guanine nucleotide exchange factor 2</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="21">Cyclic nucleotide ras GEF</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="23">Neural RAP guanine nucleotide exchange protein</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="20">PDZ domain-containing guanine nucleotide exchange factor 1</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="19">RA-GEF-1</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="22">Ras/Rap1-associating GEF-1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="31 32">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="32" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="31" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="31" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell junction</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Perinuclear region</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Late endosome</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the RAPGEF2 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000001942.1">
        <property type="protein sequence ID" value="ENSGAGP00000001707.1"/>
        <property type="gene ID" value="ENSGAGG00000001344.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070161">
        <property type="term" value="C:anchoring junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005770">
        <property type="term" value="C:late endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048471">
        <property type="term" value="C:perinuclear region of cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007264">
        <property type="term" value="P:small GTPase-mediated signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00038">
        <property type="entry name" value="CAP_ED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd06755">
        <property type="entry name" value="PDZ_RapGEF2_RapGEF6-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd01785">
        <property type="entry name" value="RA_PDZ-GEF1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00155">
        <property type="entry name" value="RasGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd06224">
        <property type="entry name" value="REM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.10:FF:000008">
        <property type="entry name" value="Rap guanine nucleotide exchange factor (GEF) 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.840.10:FF:000001">
        <property type="entry name" value="Rap guanine nucleotide exchange factor (GEF) 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000024">
        <property type="entry name" value="rap guanine nucleotide exchange factor 2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.870.10:FF:000001">
        <property type="entry name" value="rap guanine nucleotide exchange factor 2 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.42.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.10">
        <property type="entry name" value="Jelly Rolls"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.840.10">
        <property type="entry name" value="Ras guanine-nucleotide exchange factors catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.870.10">
        <property type="entry name" value="Son of sevenless (SoS) protein Chain: S domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000595">
        <property type="entry name" value="cNMP-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018490">
        <property type="entry name" value="cNMP-bd_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001478">
        <property type="entry name" value="PDZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036034">
        <property type="entry name" value="PDZ_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000159">
        <property type="entry name" value="RA_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000651">
        <property type="entry name" value="Ras-like_Gua-exchang_fac_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023578">
        <property type="entry name" value="Ras_GEF_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001895">
        <property type="entry name" value="RASGEF_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036964">
        <property type="entry name" value="RASGEF_cat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014710">
        <property type="entry name" value="RmlC-like_jellyroll"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029071">
        <property type="entry name" value="Ubiquitin-like_domsf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45161">
        <property type="entry name" value="CYTOSKELETON-ASSOCIATED PROTEIN 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45161:SF2">
        <property type="entry name" value="RAP GUANINE NUCLEOTIDE EXCHANGE FACTOR 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00595">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00788">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00617">
        <property type="entry name" value="RasGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00618">
        <property type="entry name" value="RasGEF_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00100">
        <property type="entry name" value="cNMP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00228">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00314">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00147">
        <property type="entry name" value="RasGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00229">
        <property type="entry name" value="RasGEFN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51206">
        <property type="entry name" value="cAMP-binding domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50156">
        <property type="entry name" value="PDZ domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48366">
        <property type="entry name" value="Ras GEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54236">
        <property type="entry name" value="Ubiquitin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50042">
        <property type="entry name" value="CNMP_BINDING_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50106">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50200">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50009">
        <property type="entry name" value="RASGEF_CAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50212">
        <property type="entry name" value="RASGEF_NTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="17" id="KW-0965">Cell junction</keyword>
    <keyword evidence="9" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="10" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="14" id="KW-0221">Differentiation</keyword>
    <keyword evidence="13" id="KW-0967">Endosome</keyword>
    <keyword evidence="7" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="12 24" id="KW-0344">Guanine-nucleotide releasing factor</keyword>
    <keyword evidence="18" id="KW-0472">Membrane</keyword>
    <keyword evidence="16" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="11" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="32" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="15" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="Cyclic nucleotide-binding" evidence="27">
        <location>
            <begin position="220"/>
            <end position="320"/>
        </location>
    </feature>
    <feature type="domain" description="N-terminal Ras-GEF" evidence="30">
        <location>
            <begin position="352"/>
            <end position="465"/>
        </location>
    </feature>
    <feature type="domain" description="PDZ" evidence="28">
        <location>
            <begin position="470"/>
            <end position="540"/>
        </location>
    </feature>
    <feature type="domain" description="Ras-associating" evidence="29">
        <location>
            <begin position="691"/>
            <end position="777"/>
        </location>
    </feature>
    <feature type="domain" description="Ras-GEF" evidence="26">
        <location>
            <begin position="802"/>
            <end position="1029"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="25">
        <location>
            <begin position="125"/>
            <end position="144"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="25">
        <location>
            <begin position="153"/>
            <end position="186"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="25">
        <location>
            <begin position="1084"/>
            <end position="1124"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="25">
        <location>
            <begin position="1169"/>
            <end position="1249"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="25">
        <location>
            <begin position="1298"/>
            <end position="1333"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="25">
        <location>
            <begin position="1375"/>
            <end position="1461"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="25">
        <location>
            <begin position="1517"/>
            <end position="1538"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="25">
        <location>
            <begin position="127"/>
            <end position="144"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="25">
        <location>
            <begin position="168"/>
            <end position="179"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="25">
        <location>
            <begin position="1084"/>
            <end position="1093"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="25">
        <location>
            <begin position="1115"/>
            <end position="1124"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="25">
        <location>
            <begin position="1183"/>
            <end position="1198"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="25">
        <location>
            <begin position="1215"/>
            <end position="1237"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="25">
        <location>
            <begin position="1298"/>
            <end position="1307"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="25">
        <location>
            <begin position="1322"/>
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        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="25">
        <location>
            <begin position="1383"/>
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        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="25">
        <location>
            <begin position="1431"/>
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        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="25">
        <location>
            <begin position="1451"/>
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        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="25">
        <location>
            <begin position="1519"/>
            <end position="1529"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004282"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004556"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004603"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00010829"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00016709"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022658"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00029925"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00030673"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00031545"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00031980"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00032021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00168"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="26">
        <source>
            <dbReference type="PROSITE" id="PS50009"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="27">
        <source>
            <dbReference type="PROSITE" id="PS50042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="28">
        <source>
            <dbReference type="PROSITE" id="PS50106"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="29">
        <source>
            <dbReference type="PROSITE" id="PS50200"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="30">
        <source>
            <dbReference type="PROSITE" id="PS50212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="31">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001707.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="32">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1538" mass="172465" checksum="5204D07767914BE3" modified="2019-05-08" version="1">MCETVRYERHEANEVLYYFGKRSAGSFRRGCECIVLEPSEMIVVDYMDENEEYFQRQASHRQSRRRFRKINQKGERQTIIDTVDPYPVGKPPLPRGYHTECTKPQLPADFTKLHLTDSLHPQVTHVTSSHSGCSITSDSGSSSLSDIYQATESEAGDMDLSGLPETAVDSEDDDDEEDIERASDPLMSRDIVRDCLEKDPIDRTDDDIEQLLEFMHQLPAFANMTMSVRRELCAVMVFAVVERAGTIVLNDGEELDSWSVILNGSVEVTYPDGRTEILCMGNSFGVSPTMDKEYMKGVMRTKVDDCQFVCIAQQDYCRILNQVEKNMQKVEEEGEIVMVKEHRELDRTGTRKGHIVIKGTSERLTMHLVEEHSVVDPTFIEDFLLTYRTFLSSPMEVGKKLLEWFNDPSLRDKVTRVVLLWVNNHFNDFEGDPAMTRFLEEFENNLEREKMGGHLRLLNIACAAKAKRRLITLTKPSREAPLPFILLGGSEKGFGIFVDSVDLGSKATEAGLKRGDQILEVNGQNFENIQLSKAMEILRNNTHLSITVKTNLFVFKELLTRLSEEKRNGAPHLPKIGDIKKASRYSIPDLAVDVEQVIGLEKVNKKSKANTVGGRNKLKKILDKTRISILPQKPYNDIGIGQSQDDSIVGLRQTKHIPPALPVSGTLSSSNPDLLQSHHRILDFNSTPDLPDQVLRVFKADQQSRYIMISKDTTAKEVVIQAIREFAVTATPDAYSLCEVSVTPEGVIKQRRLPDQLSKLADRIQLSGRYYLKNNMETETLCSDEDAQELLRESQISLLQLSTVEMAAQLSMRNFELFRNIEPTEYIDDLFKLKSKTSCANLKKFEEVINQETFWVASEILRETNQLKRMKIIKHFIKIALHCRECKNFNSMFAIISGLNLAPVARLRTTWEKLPSKYEKLFQDLQDLFDPSRNMAKYRNVLNSQNLQPPIIPLFPVIKKDLTFLHEGNDSKVEGLVNFEKLRMIAKEIRHVGRMASVNMDPAVMFRTRKKKWRSLGSLSQSSTNAAVLDVAQTGGHKKRVRRSSFLNAKKLYEDAQMARKVKQYLSNLDLEMDEESLQTLSLQCEPATNTLPKNPGDKRSGKQEMSPVAPRAGIQQKVQQQHKINQALQVPAVSLYPSRKKVPVKDLPPFGINSPQALKKILSLSEEGSLDRHKKQAEDTISNASSQLSSPPTSPHTSPRKGYTLAPSGTVDNFSDSGRSEISSRSSIVSNSSFDSMPVSLHDERRQRHSVSIVETNLGVGRIDRRAMIEPDQYSLGSYAPLSENRGLYAGATVLSSPSTEELSQDQGDRTSLDAADSGRGSWTSCSSGSHDNIQTIQHQRSWETLPFGHVHFDSSGDGAGLWASGSHMDQMMFPDHGTKYGRQSQGREGLDQAQSRASWASSTGYWGEDSEGDTGTIKRRGGKDVSIEAETSSITSVTAEESKPAPMSSHITVSSSSAKGLIARKEGRYREPPPTPPGYVGIPIADFAEGVSHPTRKPPDYNVALQRSRMVARTCETHGTSTLQQQSQEDEQVSAV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJ60</accession>
    <name>A0A452GJ60_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="35">Serine/threonine-protein kinase TOR</fullName>
            <ecNumber evidence="35">2.7.11.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="39 40">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="40" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="39" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="39" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="33">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="32">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="34">
            <text>L-tyrosyl-[protein] + ATP = O-phospho-L-tyrosyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10596"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.10.2"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="34">
            <dbReference type="Rhea" id="RHEA:10597"/>
        </physiologicalReaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Peripheral membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasmic vesicle</location>
            <location evidence="3">Phagosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="5">Endoplasmic reticulum membrane</location>
            <topology evidence="5">Peripheral membrane protein</topology>
            <orientation evidence="5">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Golgi apparatus membrane</location>
            <topology evidence="2">Peripheral membrane protein</topology>
            <orientation evidence="2">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="8">Lysosome membrane</location>
            <topology evidence="8">Peripheral membrane protein</topology>
            <orientation evidence="8">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="6">Microsome membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="7">Mitochondrion outer membrane</location>
            <topology evidence="7">Peripheral membrane protein</topology>
            <orientation evidence="7">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Nucleus</location>
            <location evidence="4">PML body</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="9 35">Belongs to the PI3/PI4-kinase family.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="35"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001935.1">
        <property type="protein sequence ID" value="ENSGAGP00000001700.1"/>
        <property type="gene ID" value="ENSGAGG00000000429.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000139">
        <property type="term" value="C:Golgi membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005765">
        <property type="term" value="C:lysosomal membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005741">
        <property type="term" value="C:mitochondrial outer membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045335">
        <property type="term" value="C:phagocytic vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016605">
        <property type="term" value="C:PML body"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031931">
        <property type="term" value="C:TORC1 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031932">
        <property type="term" value="C:TORC2 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004715">
        <property type="term" value="F:non-membrane spanning protein tyrosine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0106310">
        <property type="term" value="F:protein serine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="RHEA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0044877">
        <property type="term" value="F:protein-containing complex binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016242">
        <property type="term" value="P:negative regulation of macroautophagy"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009891">
        <property type="term" value="P:positive regulation of biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045597">
        <property type="term" value="P:positive regulation of cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051240">
        <property type="term" value="P:positive regulation of multicellular organismal process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008361">
        <property type="term" value="P:regulation of cell size"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019216">
        <property type="term" value="P:regulation of lipid metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051896">
        <property type="term" value="P:regulation of phosphatidylinositol 3-kinase/protein kinase B signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
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    <dbReference type="GO" id="GO:0038202">
        <property type="term" value="P:TORC1 signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="entry name" value="PIKKc_TOR"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="DJ576K7.1 (FK506 binding protein 12-rapamycin associated protein 1)"/>
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        <property type="entry name" value="MTOR isoform 6"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Serine/threonine-protein kinase mTOR"/>
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        <property type="entry name" value="Serine/threonine-protein kinase mTOR"/>
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        <property type="entry name" value="Serine/threonine-protein kinase mTOR"/>
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        <property type="entry name" value="Serine/threonine-protein kinase TOR"/>
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    </dbReference>
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        <property type="entry name" value="Serine/threonine-protein kinase TOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1010.10:FF:000004">
        <property type="entry name" value="Serine/threonine-protein kinase TOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.150">
        <property type="entry name" value="FKBP12-rapamycin binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1070.11">
        <property type="entry name" value="Phosphatidylinositol 3-/4-kinase, catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1010.10">
        <property type="entry name" value="Phosphatidylinositol 3-kinase Catalytic Subunit, Chain A, domain 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.10">
        <property type="entry name" value="Tetratricopeptide repeat domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050517">
        <property type="entry name" value="DDR_Repair_Kinase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009076">
        <property type="entry name" value="FRB_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036738">
        <property type="entry name" value="FRB_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057564">
        <property type="entry name" value="HEAT_ATR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024585">
        <property type="entry name" value="mTOR_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000403">
        <property type="entry name" value="PI3/4_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036940">
        <property type="entry name" value="PI3/4_kinase_cat_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018936">
        <property type="entry name" value="PI3/4_kinase_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003151">
        <property type="entry name" value="PIK-rel_kinase_FAT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014009">
        <property type="entry name" value="PIK_FAT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026683">
        <property type="entry name" value="TOR_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011990">
        <property type="entry name" value="TPR-like_helical_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11139">
        <property type="entry name" value="ATAXIA TELANGIECTASIA MUTATED ATM -RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11139:SF9">
        <property type="entry name" value="SERINE_THREONINE-PROTEIN KINASE MTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02259">
        <property type="entry name" value="FAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08771">
        <property type="entry name" value="FRB_dom"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23593">
        <property type="entry name" value="HEAT_ATR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11865">
        <property type="entry name" value="mTOR_dom"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00454">
        <property type="entry name" value="PI3_PI4_kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01346">
        <property type="entry name" value="DUF3385"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00146">
        <property type="entry name" value="PI3Kc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01345">
        <property type="entry name" value="Rapamycin_bind"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47212">
        <property type="entry name" value="FKBP12-rapamycin-binding domain of FKBP-rapamycin-associated protein (FRAP)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51189">
        <property type="entry name" value="FAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00915">
        <property type="entry name" value="PI3_4_KINASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00916">
        <property type="entry name" value="PI3_4_KINASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50290">
        <property type="entry name" value="PI3_4_KINASE_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="25" id="KW-0007">Acetylation</keyword>
    <keyword evidence="22 35" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="10" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="31" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="21" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="26" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="12" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="18 35" id="KW-0418">Kinase</keyword>
    <keyword evidence="29" id="KW-0458">Lysosome</keyword>
    <keyword evidence="28" id="KW-0472">Membrane</keyword>
    <keyword evidence="24" id="KW-0492">Microsome</keyword>
    <keyword evidence="27" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="19" id="KW-1000">Mitochondrion outer membrane</keyword>
    <keyword evidence="17 35" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="30" id="KW-0539">Nucleus</keyword>
    <keyword evidence="14" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="40" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="16" id="KW-0677">Repeat</keyword>
    <keyword evidence="13 35" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="20" id="KW-0802">TPR repeat</keyword>
    <keyword evidence="15 35" id="KW-0808">Transferase</keyword>
    <keyword evidence="23" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="FAT" evidence="38">
        <location>
            <begin position="1371"/>
            <end position="1957"/>
        </location>
    </feature>
    <feature type="domain" description="PI3K/PI4K catalytic" evidence="37">
        <location>
            <begin position="2131"/>
            <end position="2444"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="36">
        <location>
            <begin position="1801"/>
            <end position="1841"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="36">
        <location>
            <begin position="1812"/>
            <end position="1835"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004202"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004255"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004262"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004322"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004397"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00004524"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00004570"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00004630"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00011031"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00022787"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00022803"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00022848"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="ARBA" id="ARBA00023228"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="31">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="32">
        <source>
            <dbReference type="ARBA" id="ARBA00047899"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="33">
        <source>
            <dbReference type="ARBA" id="ARBA00048679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="34">
        <source>
            <dbReference type="ARBA" id="ARBA00051942"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="35">
        <source>
            <dbReference type="RuleBase" id="RU364109"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="36">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="37">
        <source>
            <dbReference type="PROSITE" id="PS50290"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="38">
        <source>
            <dbReference type="PROSITE" id="PS51189"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="39">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001700.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="40">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2484" mass="282521" checksum="B5EFE909497EFE2E" modified="2019-05-08" version="1">GLGSRPXQLAAIMSSTVSILQQFASGLKSRNEEIRAKAAKDLQHYVTMELREMSQEESTRFYDQLNHHIFELVSSSDANERKGGILAIASLIGVEGGNATRIGRFANYLRNLLPSNDPVVMEMASKAIGRLALAGDTFTAEYVEFEVKRALEWLGADRNEGRRHAAVLVLRELAISVPTFFFQQVQPFFDNIFVAVWDPKQAIREGAVSALRACLILTTQREPKEMQKPQWYRHTYEEAEKGFDETLAKEKGMNRDDRIHGALLILNELVRISSMEGERLREEMEEITQQQMVHDKYCKDLLGFGTKPRHITPFTSFQSVQPPQSNALVGLLGYSPHQGIMGFGASPIPAKSALVESRCCRDLMEEKFDQVCLWVLKCRTSKNSLIQMTILNLLPRLAAFRPSAFTADQYLPDTMNHILSCVKKEKERTAAFQALGLLSVAVRSEFQAYLPKVLEIIKAALPPKDFAHKRQKSMQVDATVFTCISMLARAMGPCIQQDIKELLEPMLAVGLSPALTAVLYDLSRQIPQLKKDIQDGLLKMLSLVLMHKPLRHPGMPKGLAHQLASPSLTNIPEASDVGSITLALRTLGSFEFEGHSLTQFVRHCADHFLNSEHKEIRMEAARTCSRLLTPSIHLISGHAHVVSQTAVQVVADVLSKLLVVGITDPDPDIRYCVLASLDERFDAHLAQAENLQALFVALNDQVFEIRELAICTVGRLSSMNPAFVMPFLRKMLIQILTELEHSGVGRIKEQSARMLGHLVSNAPRLIRPYMEPILKALIVKLKDPDPDPNPGVINNVLATIGELAQVSGLEMRKWVDELFIIIMDMLQDSSLLAKRQVALWTLGQLVASTGYVVEPYRKYPTLLDVLLNFLKTEQNQGTRREAIRVLGLLGALDPYKHKVNIGMIDQSRDASAVSLSESKSSQDSCKQRETGERMCTLHFSMVALMRIFRDQSLSHHHTMVVQAITFIFKSLGLKCVQFLPQVMPTFLNVIRVCDGAIREFLFQQLGMLVSFVRSHIRPYMDEIVTLMRDFWVMNNSIQSTIILLIEQIVVALGGEFKLYLPQLIPHMLRVFMHDSSPGRNVSVKLLNAIQLFGANLDDYLHLLLPPIVKLFDAPDVPLVARKAALETVDRLTESLDFTDYASRIIHPIVRTLDLSPELRPTAMDTLSSLVFQLGKKYQIFIPMVNKVLMRHRINHQRYDVLICRIVKGYTLADEEEDPLIYQHRMLRSNQGEMLTSGPVETGPMKKLHVSTINLQKAWGAARRVSKDDWLEWLRRLSLELLKDSSSPSLRSCWALAQAYNPMARDLFNAAFVSCWSELNEDQQDELIRSIELALTSQDIAEVTQTLLNLAEFMEHSDKGPLPLRDDNGIVLLGERAAKCRAYAKALHYKELEFQKGPTPAILESLISINNKLQQPEAAAGVLEYAMKHFGELEIQATWYEKLHEWEDALVAYDKKMDTNKDDPELMLGRMRCLEALGEWGQLHQQCCEKWTLVNDETQAKMARMAAAAAWGLGQWDSMEEYTCMIPRDTHDGAFYRAVLALHQDLFSLAQQCIDKARDLLDAELTAMAGESYSRAYGAMVSCQMLSELEEVIQYKLVPERREIIRQIWWERLQGCQRIVEDWQRILMVRSLVVNPHEDMRTWLKYASLCGKSGRLALAHKTLVLLLGVDPSRQLDHPLPTVHPQVTYAYMKHMWKSARKIDAFQHMQHFVQTMQEQAQHAIATEDQQHKQELHKLMARCFLKLGEWQLNLQGINESTIPKVLQYYSASTEHDGNWYKAWHAWAVMNFEAVLHYKHQNQARDEKKKLRHASGASITSANTEGSNSDSEAESTENSPIPSPVQKKVTEDLSRTLLLYTVPAVQGFFRSISLSRGNNLQDTLRVLTLWFDYGHWPDVNEALVEGVKAIQIDTWLQVIPQLIARIDTPRPLVGRLIHQLLTDIGRYHPQALIYPLTVASKSTTTARHNAANKILKNMCEHSNTLVQQAMMVSEELIRVAILWHEMWHEGLEEASRLYFGERNVKGMFEVLEPLHAMMERGPQTLKETSFNQAYGRDLMEAQEWCRKYMKSGNVKDLTQAWDLYYHVFRRISKQLPQLTSLELQYVSPKLLMCRDLELAVPGTYDPNQPIIRIQSIAPSLQVITSKQRPRKLTLMGSNGHEFVFLLKGHEDLRQDERVMQLFGLVNTLLANDPTSLRKNLSIQRYAVIPLSTNSGLIGWVPHCDTLHALIRDYREKKKILLNIEHRIMLRMAPDYDHLTLMQKVEVFEHAVNNTAGDDLAKLLWLKSPSSEVWFDRRTNYTRSLAVMSMVGYILGLGDRHPSNLMLDRLSGKILHIDFGDCFEVAMTREKFPEKIPFRLTRMLTNAMEVTGLDGNYRITCHTVMEVLREHKDSVMAVLEAFVYDPLLNWRLMDTNTKGNKRSRTRTDSYSASQSVEMLDNVELGETAHKKTGTTVPESIHSFSKFLCKHNTFYIMATVGSLQRCVCPFW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJ64</accession>
    <name>A0A452GJ64_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">Diacylglycerol kinase</fullName>
            <shortName evidence="21">DAG kinase</shortName>
            <ecNumber evidence="21">2.7.1.107</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="27 28">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="28" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="27" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="27" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>1,2-di-(9Z-octadecenoyl)-sn-glycerol + ATP = 1,2-di-(9Z-octadecenoyl)-sn-glycero-3-phosphate + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:40327"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:52333"/>
            <dbReference type="ChEBI" id="CHEBI:74546"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="18">
            <dbReference type="Rhea" id="RHEA:40328"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>a 1,2-diacyl-sn-glycerol + ATP = a 1,2-diacyl-sn-glycero-3-phosphate + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10272"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:17815"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:58608"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.1.107"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="19">
            <dbReference type="Rhea" id="RHEA:10273"/>
        </physiologicalReaction>
    </comment>
    <comment type="pathway">
        <text evidence="20">Glycerolipid metabolism.</text>
    </comment>
    <comment type="pathway">
        <text evidence="3">Lipid metabolism; glycerolipid metabolism.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4 21">Belongs to the eukaryotic diacylglycerol kinase family.</text>
    </comment>
    <dbReference type="EC" id="2.7.1.107" evidence="21"/>
    <dbReference type="AlphaFoldDB" id="A0A452GJ64"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001877.1">
        <property type="protein sequence ID" value="ENSGAGP00000001652.1"/>
        <property type="gene ID" value="ENSGAGG00000001299.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00230"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004143">
        <property type="term" value="F:ATP-dependent diacylglycerol kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046982">
        <property type="term" value="F:protein heterodimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042803">
        <property type="term" value="F:protein homodimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046486">
        <property type="term" value="P:glycerolipid metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007200">
        <property type="term" value="P:phospholipase C-activating G protein-coupled receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd20847">
        <property type="entry name" value="C1_DGKdelta_rpt1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd20893">
        <property type="entry name" value="C1_DGKdelta_rpt2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd13274">
        <property type="entry name" value="PH_DGK_type2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd09575">
        <property type="entry name" value="SAM_DGK-delta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.50:FF:000021">
        <property type="entry name" value="Diacylglycerol kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000060">
        <property type="entry name" value="Diacylglycerol kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.200.40:FF:000001">
        <property type="entry name" value="Diacylglycerol kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.60.20:FF:000002">
        <property type="entry name" value="Diacylglycerol kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.60.20:FF:000029">
        <property type="entry name" value="Diacylglycerol kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10330:FF:000001">
        <property type="entry name" value="Diacylglycerol kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.200.40">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.60.20">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.10330">
        <property type="entry name" value="Probable inorganic polyphosphate/atp-NAD kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.50">
        <property type="entry name" value="Transcription Factor, Ets-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017438">
        <property type="entry name" value="ATP-NAD_kinase_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046349">
        <property type="entry name" value="C1-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047478">
        <property type="entry name" value="C1_DGKdelta_rpt1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047477">
        <property type="entry name" value="C1_DGKdelta_rpt2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037607">
        <property type="entry name" value="DGK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037606">
        <property type="entry name" value="DGK-delta_SAM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR054474">
        <property type="entry name" value="DGKD_4H"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000756">
        <property type="entry name" value="Diacylglycerol_kin_accessory"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001206">
        <property type="entry name" value="Diacylglycerol_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016064">
        <property type="entry name" value="NAD/diacylglycerol_kinase_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002219">
        <property type="entry name" value="PKC_DAG/PE"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001660">
        <property type="entry name" value="SAM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013761">
        <property type="entry name" value="SAM/pointed_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11255">
        <property type="entry name" value="DIACYLGLYCEROL KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11255:SF30">
        <property type="entry name" value="DIACYLGLYCEROL KINASE DELTA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00130">
        <property type="entry name" value="C1_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00609">
        <property type="entry name" value="DAGK_acc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00781">
        <property type="entry name" value="DAGK_cat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22944">
        <property type="entry name" value="DGKD_4H"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00169">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07647">
        <property type="entry name" value="SAM_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00109">
        <property type="entry name" value="C1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00045">
        <property type="entry name" value="DAGKa"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00046">
        <property type="entry name" value="DAGKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00233">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00454">
        <property type="entry name" value="SAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57889">
        <property type="entry name" value="Cysteine-rich domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF111331">
        <property type="entry name" value="NAD kinase/diacylglycerol kinase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47769">
        <property type="entry name" value="SAM/Pointed domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50146">
        <property type="entry name" value="DAGK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50105">
        <property type="entry name" value="SAM_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00479">
        <property type="entry name" value="ZF_DAG_PE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50081">
        <property type="entry name" value="ZF_DAG_PE_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="15 21" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="5" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="13 21" id="KW-0418">Kinase</keyword>
    <keyword evidence="16" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="17" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11 21" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="28" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0677">Repeat</keyword>
    <keyword evidence="8 21" id="KW-0808">Transferase</keyword>
    <keyword evidence="14" id="KW-0862">Zinc</keyword>
    <keyword evidence="12" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="PH" evidence="23">
        <location>
            <begin position="1"/>
            <end position="88"/>
        </location>
    </feature>
    <feature type="domain" description="Phorbol-ester/DAG-type" evidence="24">
        <location>
            <begin position="105"/>
            <end position="155"/>
        </location>
    </feature>
    <feature type="domain" description="Phorbol-ester/DAG-type" evidence="24">
        <location>
            <begin position="177"/>
            <end position="228"/>
        </location>
    </feature>
    <feature type="domain" description="DAGKc" evidence="26">
        <location>
            <begin position="259"/>
            <end position="394"/>
        </location>
    </feature>
    <feature type="domain" description="SAM" evidence="25">
        <location>
            <begin position="1060"/>
            <end position="1123"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="497"/>
            <end position="543"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="600"/>
            <end position="626"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="22">
        <location>
            <begin position="497"/>
            <end position="506"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="22">
        <location>
            <begin position="523"/>
            <end position="537"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00009280"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023371"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023411"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00060536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="RuleBase" id="RU361128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS50081"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS50105"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="26">
        <source>
            <dbReference type="PROSITE" id="PS50146"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001652.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="28">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1129" mass="126684" checksum="D497ED45911B5FED" modified="2019-05-08" version="1">MLMKQTSSFQRWKRRYFKLRGRTLYYAKNAKSIIFDEVDLTDASVAESSTKNINNSFTVITPCRKLILCADNRKEMEDWIAALKTIQNREHFESTQYSMDHFSGMHNWYACSHARPTYCNVCREALSGVTSHGLSCEVCKFKAHKRCAVRATNNCKWTTLASIGKDIIEDEDGISMPHQWLEGNLPVSAKCMVCDKTCGSVLRLQDWRCLWCKAMVHTTCKDLLPTKCPLGLCKVSVIPPTALNSIDSDGFWKATCPPSCTSPLLVFVNSKSGDNQGVKFLRRFKQLLNPAQVFDLMNGGPHLGLRLFQKFDTFRILVCGGDGSVGWVLSEIDSLNLHKQCQLGVLPLGTGNDLARVLGWGSACDDDTQLPQILEKLERASTKMLDRWSIMVYETKLPRQSSTSTVTEDFSDDSEVQKILFYEDSVAAHLSKILTSDQHSVVISSAKVLCETVKDFVARVGKAYEKATESSEESEVMARKCSVLKEKLDSLLKTLNDESQASSSLSNPPPTIAEETEDGDGSGSACDSSSDHSMGSSCTARPQIFRPREQLMLRANSLKKAIRQIIEHAEKAVDEQNAQTQEQEGFILGLPTSEEGKELRNEDKTSLQPLHSGNYGVPKGRSHRKVSKSPCERLINKGSLSLGSSASLPPQTGNRDNLPMLNTKILYPNIRAGMSGSLPGSSVISRLLINADPFNSEPENLECYTEKCVMNNYFGIGLDAKISLDFNNKRDEHPEKCRSRTKNMMWYGVLGTKELLHRTYKNLEQKVLLECDGRPIPLPSLQGIAVLNIPSYAGGTNFWGGTKEDDTFTAPSFDDKILEVVAVFGSMQMAVSRVINLQHHRIAQCRTVKIAILGDEGVPVQVDGEAWIQPPGYIWIVHKNRAQTLTRDRAFESTLKSWEDKQKCELPRPSSFSLHPEIMSEEESTQINQFGQAAGALIHSIREIAQSYQDMEQELAHAVNASSKSMDKVYAKSKSTEQLDPPQKEQLLAALMDMDLQLRKLADIPWLWQLMEPSDDENQMLDYSKRSRSGKFRLVTKFKKEKNNKNKETRSSMGLPVHLWGTEEVAAWLEHLSLCEYKDIFIRHDVRGSELLHLERRDLKDLGVTKVGHMKRILHGIKELSRNTLVSEV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJ66</accession>
    <name>A0A452GJ66_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">S-arrestin</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="4">Retinal S-antigen</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="3">Rod photoreceptor arrestin</fullName>
        </alternativeName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the arrestin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJ66"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001951.1">
        <property type="protein sequence ID" value="ENSGAGP00000001716.1"/>
        <property type="gene ID" value="ENSGAGG00000001371.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001917">
        <property type="term" value="C:photoreceptor inner segment"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001750">
        <property type="term" value="C:photoreceptor outer segment"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001664">
        <property type="term" value="F:G protein-coupled receptor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002031">
        <property type="term" value="P:G protein-coupled receptor internalization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.840:FF:000002">
        <property type="entry name" value="Arrestin 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.640:FF:000011">
        <property type="entry name" value="S-arrestin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.640">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.840">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000698">
        <property type="entry name" value="Arrestin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014752">
        <property type="entry name" value="Arrestin-like_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011021">
        <property type="entry name" value="Arrestin-like_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011022">
        <property type="entry name" value="Arrestin_C-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017864">
        <property type="entry name" value="Arrestin_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014753">
        <property type="entry name" value="Arrestin_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014756">
        <property type="entry name" value="Ig_E-set"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11792">
        <property type="entry name" value="ARRESTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11792:SF15">
        <property type="entry name" value="S-ARRESTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02752">
        <property type="entry name" value="Arrestin_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00339">
        <property type="entry name" value="Arrestin_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00309">
        <property type="entry name" value="ARRESTIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM01017">
        <property type="entry name" value="Arrestin_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81296">
        <property type="entry name" value="E set domains"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00295">
        <property type="entry name" value="ARRESTINS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Arrestin C-terminal-like" evidence="5">
        <location>
            <begin position="212"/>
            <end position="365"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00005298"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00040206"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00041305"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00042071"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="SMART" id="SM01017"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001716.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="426" mass="48741" checksum="8A590B6E5B6FD379" modified="2019-05-08" version="1">MSCAESHKVATPSSICLKNTSPPQSHVVYKKMSRDKAVTIYLGKRDFIDHIDGVEPVDGVLLVDPEIVKGKKVYVTLTCAFRYGQEDIDVMGLTFRKDLFFSRIQLYPPVEKWESLSQLQECLMKKLGNNAYPFVLTFPDYLPCSVSLQPAPHDVGKCCGVDFEVKAFCTENVEERIPKRNFVRLLIRKVQYAPEESGPQPQAETTWQFFMSSKPLHLRARLSKEVFYHGEPIPVTVTVTNNTEKTVKKIRVLVEQVANVVLYSSDYYTKVVAMEEMQEKVQPNSIITKTLILLPLLANNREKRGIALDGKLKDEDTNLASSTIIKDGIDKTVLGILVAYKVKVKLIISGEVSTELPFRLMHPKPEEPNIRVPRMKTWYLKSFLVNSCKIWTKMRIRLFLQMMSDLENRLLEDLWGLMVVTNTFRI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJ70</accession>
    <name>A0A452GJ70_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">Dynein axonemal heavy chain 11</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Cilium axoneme</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the dynein heavy chain family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000001945.1">
        <property type="protein sequence ID" value="ENSGAGP00000001710.1"/>
        <property type="gene ID" value="ENSGAGG00000000806.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005930">
        <property type="term" value="C:axoneme"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030286">
        <property type="term" value="C:dynein complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005874">
        <property type="term" value="C:microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031514">
        <property type="term" value="C:motile cilium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045505">
        <property type="term" value="F:dynein intermediate chain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051959">
        <property type="term" value="F:dynein light intermediate chain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007018">
        <property type="term" value="P:microtubule-based movement"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.2620:FF:000001">
        <property type="entry name" value="Cytoplasmic dynein heavy chain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000667">
        <property type="entry name" value="Dynein axonemal heavy chain 11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.920.30:FF:000003">
        <property type="entry name" value="Dynein axonemal heavy chain 17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000219">
        <property type="entry name" value="Dynein axonemal heavy chain 17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.20.180.20:FF:000001">
        <property type="entry name" value="Dynein axonemal heavy chain 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.8.710:FF:000002">
        <property type="entry name" value="dynein heavy chain 17, axonemal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.140.100:FF:000001">
        <property type="entry name" value="dynein heavy chain 17, axonemal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.1120:FF:000002">
        <property type="entry name" value="Dynein heavy chain 9, axonemal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.472.130:FF:000001">
        <property type="entry name" value="Dynein, axonemal, heavy chain 9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.2620">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.472.130">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.8.710">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.1120">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.920.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.140.100">
        <property type="entry name" value="Dynein heavy chain, N-terminal domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.20.180.20">
        <property type="entry name" value="Dynein heavy chain, N-terminal domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035699">
        <property type="entry name" value="AAA_6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026983">
        <property type="entry name" value="DHC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041589">
        <property type="entry name" value="DNAH3_AAA_lid_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042222">
        <property type="entry name" value="Dynein_2_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043157">
        <property type="entry name" value="Dynein_AAA1S"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041466">
        <property type="entry name" value="Dynein_AAA5_ext"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013602">
        <property type="entry name" value="Dynein_heavy_linker"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013594">
        <property type="entry name" value="Dynein_heavy_tail"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042228">
        <property type="entry name" value="Dynein_linker_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45703:SF12">
        <property type="entry name" value="DYNEIN AXONEMAL HEAVY CHAIN 11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45703">
        <property type="entry name" value="DYNEIN HEAVY CHAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12774">
        <property type="entry name" value="AAA_6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12775">
        <property type="entry name" value="AAA_7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17857">
        <property type="entry name" value="AAA_lid_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08385">
        <property type="entry name" value="DHC_N1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08393">
        <property type="entry name" value="DHC_N2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17852">
        <property type="entry name" value="Dynein_AAA_lid"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="13" id="KW-0966">Cell projection</keyword>
    <keyword evidence="10" id="KW-0969">Cilium</keyword>
    <keyword evidence="9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="12" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="8" id="KW-0243">Dynein</keyword>
    <keyword evidence="4" id="KW-0493">Microtubule</keyword>
    <keyword evidence="11" id="KW-0505">Motor protein</keyword>
    <keyword evidence="6" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="Dynein heavy chain tail" evidence="14">
        <location>
            <begin position="80"/>
            <end position="365"/>
        </location>
    </feature>
    <feature type="domain" description="Dynein heavy chain tail" evidence="14">
        <location>
            <begin position="383"/>
            <end position="535"/>
        </location>
    </feature>
    <feature type="domain" description="Dynein heavy chain linker" evidence="15">
        <location>
            <begin position="1033"/>
            <end position="1432"/>
        </location>
    </feature>
    <feature type="domain" description="Dynein heavy chain hydrolytic ATP-binding dynein motor region" evidence="16">
        <location>
            <begin position="1566"/>
            <end position="1892"/>
        </location>
    </feature>
    <feature type="domain" description="Dynein heavy chain AAA 5 extension" evidence="17">
        <location>
            <begin position="2050"/>
            <end position="2168"/>
        </location>
    </feature>
    <feature type="domain" description="Dynein heavy chain 3 AAA+ lid" evidence="18">
        <location>
            <begin position="2385"/>
            <end position="2463"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004430"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008887"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023017"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023069"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="Pfam" id="PF08385"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="Pfam" id="PF08393"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF12774"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF17852"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF17857"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001710.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="21">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="2533" mass="295061" checksum="3262EC45A294F069" modified="2019-05-08" version="1">MKSFDCRLELTEGNTESLKTAGDKKMKRDKKVARARNNTVTEDSRKTAVMVWIYIKKAPGEDTIHGDTYQSKQSYLRFYHFLFTALMEAQDIELYLRPLRRHIQFLQETEFPKIHSLIPPLFHTICLIWSHSQFYSVPARIIVLLQEFCNLFIDQARSYLSPEDLLKGEIEETLEHVQIAVNTLRSFKNFFFSHREKLASYFTNSKEFKPWDFQSNMVFARFDLFLNRLDIFVVMLEFQKLEKLEFGGVKGKTLSEQIYRMNEEFIESCKVFKEKTYDPSDYNNDYVEFKAKILDFDRRLGSILCVGFLDCSGLESAFKLLAIFGYFLEKPVIMEIFSPNYIILLQMFDEELDNCKHLYDEHIRQVCSKAKIKAIFFLVFVRILQDPETALVYQKYIETLTLIDQFEEQIYCKWKSQVDEVYQFNLDQPLIKRNPENGLLSVNFDSKLVALLREVTYLLKLNRSDIPDSALAIYEKRKIFYIGNLELIVQWYNKLKQTVLEVEYPLIETELEAIDGQLRETEEVFTWQKENCWKYIEQVKAAVYDLEQRVQQSKENIKTIQQIMNAWAEHTLFSRKDDRKDALLSLEDKEDRLAKKYKLLKEDGYKIHQLAEENLKLFKADSSSDSWKIYIEYIDDIIVDGFFNTLMHNLDFFLENTEKGLKPAPLFQAQMILSAPEIEFKPSLDKEAGDGFYDLVEELLSNVFKMSAQVKRVAAHLEVENYQSDMDNMLDLSEIRQEIMDRVADVIRKALDYRNSLDTYAYLWVDDRAEFMKQFLLYGHVLTSEEIDSHADEGAPEQPPTTEQFKEQIDIYENLYIQMSKFEDFRTFESWFKVDMKPFKMSLLNIIKKWSWMFKEHLLKFVIGSLTDLEEFIKVTDAGLQKEVNEGDYNGLVEIMGHLLAVRDRQASTDELFEPLKETITLLESYGQKMSDQVYVQLEELPEKWNTTKKIAASIKHEVAPLQTSEVTLIRKKFALFDGKQIEFRERFRVEAPFCFDAKYPYILLDKALEEEMLQMQKSANLFEVVVPDYKQLKQCRKEIRLLKEVWDINIYVTSSIGDWTKTQWRQINVEQMDAELRRFAKEVWSLDKEIRSWDVYIGLELTVKNLMTSLRAVTELQNPAIRERHWHQLMNATDVSTTLADLLGLQLHKVEDEVRSIVDKAVKELGTEKILTEISQTWATVEFCYEEHYRTRTPLLKCDEQLFETLENNQVQLQTILQSKYVEYFIEQVTSWQRTLNVADSVIFIWMEIQRTWSHLESIFIGSDDIRTKLPKDAKRFDGVDADFKELMMETANTKNVLEATTKPNLYEKLESLQRRLSLCEKALAEYLETKCLAFPRFYFVSSADLLDILSKGTQPRQVTRHLTKLFDNIADLQFQDNKEKSTDIALGMYSKEKEYVPFHAECECIGQVETWLQQLEETMYKTVHRHIMEAIAAYEEKPREQWIFDYPAQVALTSSQIWWTTDVGIAFNRLEEGFETALKDYNKKQIAQLNALIALLLGELASGDRQKIMTICTIDVHARDVVAKLVAQKVTSSQAFAWLSQLRHRWDEAQKHCLANICDAQFQYFYEYLGNTSRLVITPLTDRCYITLTQSLHLTMSGAPAGPTGTGKTETTKDLGHALGMMVYVFNCSEQMDYKSIGNIYKGLVQTGAWGCFDEFNRISVEVLSVVAVQVKTIHDAIRNKRKKFLFLGENITLKTSVGIFITMNPGYAGRTELPENLKALFRPCAMVVPDIELICEIMLVAEGFTDARLLARKFITLYTLCRELLSKQDHYDWGLRAVKSVLVVAGSLKRGDKNRPEDQVLMRTLRDFNLPKIVTDDIPIFMGLISDLFPALDVPRRRDLQFEQMVKQSTLELHLQPEESFIFKVVQLEELLAVRHSVFVVGNAGTGKSKVLRTLNRTYVNMKQKLVWNDLNPKAVTTDELFGFIHHATREWKDGLFSSLLREQANITQDGPKWIVLDGDIDPMWIESLNTVMDDNKVLTLASNERVPLTPSMRLLFEIHHLKTATPATVSRAGILYVNPQDLGWNPYVASWIETRSHQSEKANLTILFDKYVPPCLDQLRTSFKTITPIPENSMVQTLCSLLDCLLTPENVPPDCPRELYEIYFVFACVWAFGGALFQDQLSNYQAEFSRWWLKEMKAVKFPSQGTIFDYYLDPKTRKFLPWTDKVPRFDMDPETPLQAVLVHTSETTRLRYFTDLLLERGKPVMLVGNAGVGKTVFVGDRLAAFPDDYMVSNVLFNYYTTSSVLQKILEKSLEKKAGRNYGPVGNKKLVYFIDDMNMPEMDVYGTVQPHTLIRQHIDYGHWYDRQKLTVKEIHNCHYVACMNPTAGNFTINPRLQRHFTVFALNFPSVDALNTIYSKILGFHFQRHVFNPSVLKNAPAMIQASIWLHQTMVQNFLPTAVKFHYIFNLRDLSNIFQGILFATPECLREPNDLVCLWLHESSRVYGDKLVEPKDCHFFRKKLIDTGVEDHLLRQHPLIYCHFANGVASPHYMPVKEWEVLRKILTETLESYNELNASMNLVLFEDAMQHV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJ71</accession>
    <name>A0A452GJ71_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">Dynein axonemal heavy chain 11</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Cilium axoneme</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the dynein heavy chain family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000001956.1">
        <property type="protein sequence ID" value="ENSGAGP00000001721.1"/>
        <property type="gene ID" value="ENSGAGG00000000806.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005930">
        <property type="term" value="C:axoneme"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030286">
        <property type="term" value="C:dynein complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005874">
        <property type="term" value="C:microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031514">
        <property type="term" value="C:motile cilium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045505">
        <property type="term" value="F:dynein intermediate chain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051959">
        <property type="term" value="F:dynein light intermediate chain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007018">
        <property type="term" value="P:microtubule-based movement"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000667">
        <property type="entry name" value="Dynein axonemal heavy chain 11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.2620:FF:000004">
        <property type="entry name" value="Dynein axonemal heavy chain 17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.920.30:FF:000003">
        <property type="entry name" value="Dynein axonemal heavy chain 17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000219">
        <property type="entry name" value="Dynein axonemal heavy chain 17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.20.180.20:FF:000001">
        <property type="entry name" value="Dynein axonemal heavy chain 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.8.710:FF:000002">
        <property type="entry name" value="dynein heavy chain 17, axonemal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.140.100:FF:000001">
        <property type="entry name" value="dynein heavy chain 17, axonemal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.1120:FF:000002">
        <property type="entry name" value="Dynein heavy chain 9, axonemal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.472.130:FF:000001">
        <property type="entry name" value="Dynein, axonemal, heavy chain 9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.2620">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.472.130">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.8.710">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.1120">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.920.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.140.100">
        <property type="entry name" value="Dynein heavy chain, N-terminal domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.20.180.20">
        <property type="entry name" value="Dynein heavy chain, N-terminal domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035699">
        <property type="entry name" value="AAA_6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026983">
        <property type="entry name" value="DHC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041589">
        <property type="entry name" value="DNAH3_AAA_lid_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042222">
        <property type="entry name" value="Dynein_2_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043157">
        <property type="entry name" value="Dynein_AAA1S"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041466">
        <property type="entry name" value="Dynein_AAA5_ext"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013602">
        <property type="entry name" value="Dynein_heavy_linker"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013594">
        <property type="entry name" value="Dynein_heavy_tail"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042228">
        <property type="entry name" value="Dynein_linker_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45703:SF12">
        <property type="entry name" value="DYNEIN AXONEMAL HEAVY CHAIN 11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45703">
        <property type="entry name" value="DYNEIN HEAVY CHAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12774">
        <property type="entry name" value="AAA_6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12775">
        <property type="entry name" value="AAA_7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17857">
        <property type="entry name" value="AAA_lid_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08385">
        <property type="entry name" value="DHC_N1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08393">
        <property type="entry name" value="DHC_N2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17852">
        <property type="entry name" value="Dynein_AAA_lid"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="13" id="KW-0966">Cell projection</keyword>
    <keyword evidence="10" id="KW-0969">Cilium</keyword>
    <keyword evidence="9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="12" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="8" id="KW-0243">Dynein</keyword>
    <keyword evidence="4" id="KW-0493">Microtubule</keyword>
    <keyword evidence="11" id="KW-0505">Motor protein</keyword>
    <keyword evidence="6" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="Dynein heavy chain tail" evidence="14">
        <location>
            <begin position="66"/>
            <end position="311"/>
        </location>
    </feature>
    <feature type="domain" description="Dynein heavy chain tail" evidence="14">
        <location>
            <begin position="315"/>
            <end position="458"/>
        </location>
    </feature>
    <feature type="domain" description="Dynein heavy chain linker" evidence="15">
        <location>
            <begin position="957"/>
            <end position="1362"/>
        </location>
    </feature>
    <feature type="domain" description="Dynein heavy chain hydrolytic ATP-binding dynein motor region" evidence="16">
        <location>
            <begin position="1496"/>
            <end position="1822"/>
        </location>
    </feature>
    <feature type="domain" description="Dynein heavy chain AAA 5 extension" evidence="17">
        <location>
            <begin position="1980"/>
            <end position="2098"/>
        </location>
    </feature>
    <feature type="domain" description="Dynein heavy chain 3 AAA+ lid" evidence="18">
        <location>
            <begin position="2315"/>
            <end position="2411"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004430"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008887"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023017"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023069"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="Pfam" id="PF08385"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="Pfam" id="PF08393"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF12774"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF17852"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF17857"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001721.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="21">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="2469" mass="287394" checksum="79BE2B738CD79735" modified="2019-05-08" version="1">SNFCITPSWLAEAGEQTDRKERVVMHRSLTPSSRTDEQGFLGLHHNTVSCPRNLHIFLPGFYHFLFTALMEAQDIELYLRPLRRHIQFLQETEFPKIHSLIPPLFHTICLIWSHSQFYSVPARIIVLLQEFCNLFIDQARSYLSPEDLLKGEIEETLEHVQIAVNTLRSFKNFFFSHREKLASYFTNSKEFKPWDFQSNMVFARFDLFLNRLVKIEDIFVVMLEFQKLEKLEFGGVKGKTLSEQIYRMNEEFIESCKVFKEKTYDPSDYNNVEFEEDYVEFKAKILDFDRRLGSILCVGFLDCSGLESAFKTALVYQKYIETLTLIDQFEEQIYCKWKSQVDEVYQFNLDQPLIKRNPENGLLSVNFDSKLVALLREVTYLLKLNRSDIPDSALAIYEKRKIFYIGNLELIVQWYNKLKQTVLEVEYPLIETELEAIDGQLRETEEVFTWQKENCWKYIEQVKAAVYDLEQRVQQSKENIKTIQQIMNAWAEHTLFSRKDDRKDALLSLEDKEDRLAKKYKLLKEDGYKIHQLAEENLKLFKADSSSDSWKIYIEYIDDIIVDGFFNTLMHNLDFFLENTEKGLKPAPLFQAQMILSAPEIEFKPSLDKEAGDGFYDLVEELLSNVFKMSAQVKRVAAHLEVENYQSDMDNMLDLSEIRQEIMDRVADVIRKALDYRNSLDTYAYLWVDDRAEFMKQFLLYGHVLTSEEIDSHADEGAPEQPPTTEQFKEQIDIYENLYIQMSKFEDFRTFESWFKVDMKPFKMSLLNIIKKWSWMFKEHLLKFVIGSLTDLEEFIKVTDAGLQKEVNEGDYNGLVEIMGHLLAVRDRQASTDELFEPLKETITLLESYGQKMSDQVYVQLEELPEKWNTTKKIAASIKHEVAPLQTSEVTLIRKKFALFDGKQIEFRERFRVEAPFCFDAKYPYILLDKVHHIDTFIYSTKSANLFEVVVPDYKQLKQCRKEIRLLKEVWDINIYVTSSIGDWTKTQWRQINVEQMDAELRRFAKEVWSLDKEIRSWDVYIGLELTVKNLMTSLRAVTELQNPAIRERHWHQLMNATDVRLSITEGTTLADLLGLQLHKVEDEVRSIVDKAVKELGTEKILTEISQTWATVEFCYEEHYRTRTPLLKCDEQLFETLENNQVQLQTILQSKYVEYFIEQVTSWQRTLNVADSVIFIWMEIQRTWSHLESIFIGSDDIRTKLPKDAKRFDGVDADFKELMMETANTKNVLEATTKPNLYEKLESLQRRLSLCEKALAEYLETKCLAFPRFYFVSSADLLDILSKGTQPRQVTRHLTKLFDNIADLQFQDNKEKSTDIALGMYSKEKEYVPFHAECECIGQVETWLQQLEETMYKTVHRHIMEAIAAYEEKPREQWIFDYPAQVALTSSQIWWTTDVGIAFNRLEEGFETALKDYNKKQIAQLNALIALLLGELASGDRQKIMTICTIDVHARDVVAKLVAQKVTSSQAFAWLSQLRHRWDEAQKHCLANICDAQFQYFYEYLGNTSRLVITPLTDRCYITLTQSLHLTMSGAPAGPTGTGKTETTKDLGHALGMMVYVFNCSEQMDYKSIGNIYKGLVQTGAWGCFDEFNRISVEVLSVVAVQVKTIHDAIRNKRKKFLFLGENITLKTSVGIFITMNPGYAGRTELPENLKALFRPCAMVVPDIELICEIMLVAEGFTDARLLARKFITLYTLCRELLSKQDHYDWGLRAVKSVLVVAGSLKRGDKNRPEDQVLMRTLRDFNLPKIVTDDIPIFMGLISDLFPALDVPRRRDLQFEQMVKQSTLELHLQPEESFIFKVVQLEELLAVRHSVFVVGNAGTGKSKVLRTLNRTYVNMKQKLVWNDLNPKAVTTDELFGFIHHATREWKDGLFSSLLREQANITQDGPKWIVLDGDIDPMWIESLNTVMDDNKVLTLASNERVPLTPSMRLLFEIHHLKTATPATVSRAGILYVNPQDLGWNPYVASWIETRSHQSEKANLTILFDKYVPPCLDQLRTSFKTITPIPENSMVQTLCSLLDCLLTPENVPPDCPRELYEIYFVFACVWAFGGALFQDQLSNYQAEFSRWWLKEMKAVKFPSQGTIFDYYLDPKTRKFLPWTDKVPRFDMDPETPLQAVLVHTSETTRLRYFTDLLLERGKPVMLVGNAGVGKTVFVGDRLAAFPDDYMVSNVLFNYYTTSSVLQKILEKSLEKKAGRNYGPVGNKKLVYFIDDMNMPEMDVYGTVQPHTLIRQHIDYGHWYDRQKLTVKEIHNCHYVACMNPTAGNFTINPRLQRHFTVFALNFPSVDALNTIYSKILGFHFQRHVFNPSVLKNAPAMIQASIWLHQTMVQNFLPTAVKFHYIFNLRDLSNIFQGILFATPECLREPNDLVCLWLHESSRVYGDKLVEPKDCHFFRKKLIDTAHKYFEGVEDHLLRQHPLIYCHFANGVASPHYMPVKEWEVLRKILTETLESYNELNASMNLVLFEDAMQHV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJ74</accession>
    <name>A0A452GJ74_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">Diacylglycerol kinase</fullName>
            <shortName evidence="17">DAG kinase</shortName>
            <ecNumber evidence="17">2.7.1.107</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>1,2-di-(9Z-octadecenoyl)-sn-glycerol + ATP = 1,2-di-(9Z-octadecenoyl)-sn-glycero-3-phosphate + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:40327"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:52333"/>
            <dbReference type="ChEBI" id="CHEBI:74546"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="15">
            <dbReference type="Rhea" id="RHEA:40328"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>a 1,2-diacyl-sn-glycerol + ATP = a 1,2-diacyl-sn-glycero-3-phosphate + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10272"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:17815"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:58608"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.1.107"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="16">Glycerolipid metabolism.</text>
    </comment>
    <comment type="pathway">
        <text evidence="2">Lipid metabolism; glycerolipid metabolism.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 17">Belongs to the eukaryotic diacylglycerol kinase family.</text>
    </comment>
    <dbReference type="EC" id="2.7.1.107" evidence="17"/>
    <dbReference type="AlphaFoldDB" id="A0A452GJ74"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001870.1">
        <property type="protein sequence ID" value="ENSGAGP00000001646.1"/>
        <property type="gene ID" value="ENSGAGG00000001299.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00230"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004143">
        <property type="term" value="F:ATP-dependent diacylglycerol kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046982">
        <property type="term" value="F:protein heterodimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042803">
        <property type="term" value="F:protein homodimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046486">
        <property type="term" value="P:glycerolipid metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007200">
        <property type="term" value="P:phospholipase C-activating G protein-coupled receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd20847">
        <property type="entry name" value="C1_DGKdelta_rpt1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd20893">
        <property type="entry name" value="C1_DGKdelta_rpt2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd09575">
        <property type="entry name" value="SAM_DGK-delta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.50:FF:000021">
        <property type="entry name" value="Diacylglycerol kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.200.40:FF:000001">
        <property type="entry name" value="Diacylglycerol kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.60.20:FF:000002">
        <property type="entry name" value="Diacylglycerol kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.60.20:FF:000029">
        <property type="entry name" value="Diacylglycerol kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10330:FF:000001">
        <property type="entry name" value="Diacylglycerol kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.200.40">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.60.20">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.10330">
        <property type="entry name" value="Probable inorganic polyphosphate/atp-NAD kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.50">
        <property type="entry name" value="Transcription Factor, Ets-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017438">
        <property type="entry name" value="ATP-NAD_kinase_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046349">
        <property type="entry name" value="C1-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047478">
        <property type="entry name" value="C1_DGKdelta_rpt1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047477">
        <property type="entry name" value="C1_DGKdelta_rpt2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037607">
        <property type="entry name" value="DGK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037606">
        <property type="entry name" value="DGK-delta_SAM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR054474">
        <property type="entry name" value="DGKD_4H"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000756">
        <property type="entry name" value="Diacylglycerol_kin_accessory"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001206">
        <property type="entry name" value="Diacylglycerol_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016064">
        <property type="entry name" value="NAD/diacylglycerol_kinase_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002219">
        <property type="entry name" value="PKC_DAG/PE"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001660">
        <property type="entry name" value="SAM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013761">
        <property type="entry name" value="SAM/pointed_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11255">
        <property type="entry name" value="DIACYLGLYCEROL KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11255:SF30">
        <property type="entry name" value="DIACYLGLYCEROL KINASE DELTA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00130">
        <property type="entry name" value="C1_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00609">
        <property type="entry name" value="DAGK_acc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00781">
        <property type="entry name" value="DAGK_cat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22944">
        <property type="entry name" value="DGKD_4H"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07647">
        <property type="entry name" value="SAM_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00109">
        <property type="entry name" value="C1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00045">
        <property type="entry name" value="DAGKa"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00046">
        <property type="entry name" value="DAGKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00454">
        <property type="entry name" value="SAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57889">
        <property type="entry name" value="Cysteine-rich domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF111331">
        <property type="entry name" value="NAD kinase/diacylglycerol kinase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47769">
        <property type="entry name" value="SAM/Pointed domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50146">
        <property type="entry name" value="DAGK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50105">
        <property type="entry name" value="SAM_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00479">
        <property type="entry name" value="ZF_DAG_PE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50081">
        <property type="entry name" value="ZF_DAG_PE_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="13 17" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11 17" id="KW-0418">Kinase</keyword>
    <keyword evidence="14" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9 17" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="6 17" id="KW-0808">Transferase</keyword>
    <keyword evidence="12" id="KW-0862">Zinc</keyword>
    <keyword evidence="10" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="PH" evidence="19">
        <location>
            <begin position="1"/>
            <end position="61"/>
        </location>
    </feature>
    <feature type="domain" description="Phorbol-ester/DAG-type" evidence="20">
        <location>
            <begin position="78"/>
            <end position="128"/>
        </location>
    </feature>
    <feature type="domain" description="Phorbol-ester/DAG-type" evidence="20">
        <location>
            <begin position="150"/>
            <end position="201"/>
        </location>
    </feature>
    <feature type="domain" description="DAGKc" evidence="22">
        <location>
            <begin position="232"/>
            <end position="367"/>
        </location>
    </feature>
    <feature type="domain" description="SAM" evidence="21">
        <location>
            <begin position="1064"/>
            <end position="1127"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="470"/>
            <end position="516"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="573"/>
            <end position="599"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="470"/>
            <end position="479"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="18">
        <location>
            <begin position="496"/>
            <end position="510"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00009280"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023371"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00060536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="RuleBase" id="RU361128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50081"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50105"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS50146"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001646.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1133" mass="127063" checksum="6E355F3CC1E72DE5" modified="2019-05-08" version="1">MLMKQTSSFQRWKRRYFKLRGRTLYYAKNAKVITPCRKLILCADNRKEMEDWIAALKTIQNREHFESTQYSMDHFSGMHNWYACSHARPTYCNVCREALSGVTSHGLSCEVCKFKAHKRCAVRATNNCKWTTLASIGKDIIEDEDGISMPHQWLEGNLPVSAKCMVCDKTCGSVLRLQDWRCLWCKAMVHTTCKDLLPTKCPLGLCKVSVIPPTALNSIDSDGFWKATCPPSCTSPLLVFVNSKSGDNQGVKFLRRFKQLLNPAQVFDLMNGGPHLGLRLFQKFDTFRILVCGGDGSVGWVLSEIDSLNLHKQCQLGVLPLGTGNDLARVLGWGSACDDDTQLPQILEKLERASTKMLDRWSIMVYETKLPRQSSTSTVTEDFSDDSEVQKILFYEDSVAAHLSKILTSDQHSVVISSAKVLCETVKDFVARVGKAYEKATESSEESEVMARKCSVLKEKLDSLLKTLNDESQASSSLSNPPPTIAEETEDGDGSGSACDSSSDHSMGSSCTARPQIFRPREQLMLRANSLKKAIRQIIEHAEKAVDEQNAQTQEQEGFILGLPTSEEGKELRNEDKTSLQPLHSGNYGVPKGRSHRKVSKSPCERLINKGSLSLGSSASLPPQTGNRDNLPMLNTKILYPNIRAGMSGSLPGSSVISRLLINADPFNSEPENLECYTEKCVMNNYFGIGLDAKISLDFNNKRDEHPEKCRSRTKNMMWYGVLGTKELLHRTYKNLEQKVLLECDGRPIPLPSLQGIAVLNIPSYAGGTNFWGGTKEDDTFTAPSFDDKILEVVAVFGSMQMAVSRVINLQHHRIAQCRTVKIAILGDEGVPVQVDGEAWIQPPGYIWIVHKNRAQTLTRDRAFESTLKSWEDKQKCELPRPSSFSLHPEIMSEEESTQINQFGQAAGALIHSIREIAQSYQDMEQELAHAVNASSKSMDKVYAKSKSTEGLNCSLVVEMVNNVKALHSETELLLAGKMALQLDPPQKEQLLAALMDMDLQLRKLADIPWLWQLMEPSDDENQMLDYSKRSRSGKFRLVTKFKKEKNNKNKETRSSMGLPVHLWGTEEVAAWLEHLSLCEYKDIFIRHDVRGSELLHLERRDLKDLGVTKVGHMKRILHGIKELSRNTLVSEV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJ98</accession>
    <name>A0A452GJ98_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Autophagy-related protein 16 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the WD repeat ATG16 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJ98"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001996.1">
        <property type="protein sequence ID" value="ENSGAGP00000001754.1"/>
        <property type="gene ID" value="ENSGAGG00000001408.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034274">
        <property type="term" value="C:Atg12-Atg5-Atg16 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000421">
        <property type="term" value="C:autophagosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034045">
        <property type="term" value="C:phagophore assembly site membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043495">
        <property type="term" value="F:protein-membrane adaptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000045">
        <property type="term" value="P:autophagosome assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd22887">
        <property type="entry name" value="Atg16_CCD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00200">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.170:FF:000039">
        <property type="entry name" value="Autophagy-related protein 16-1 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:000364">
        <property type="entry name" value="Autophagy-related protein 16-1 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.170">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045160">
        <property type="entry name" value="ATG16"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013923">
        <property type="entry name" value="Autophagy-rel_prot_16_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020472">
        <property type="entry name" value="WD40_PAC1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019775">
        <property type="entry name" value="WD40_repeat_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036322">
        <property type="entry name" value="WD40_repeat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001680">
        <property type="entry name" value="WD40_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19878">
        <property type="entry name" value="AUTOPHAGY PROTEIN 16-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19878:SF6">
        <property type="entry name" value="AUTOPHAGY-RELATED PROTEIN 16-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08614">
        <property type="entry name" value="ATG16"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00400">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00320">
        <property type="entry name" value="GPROTEINBRPT"/>
    </dbReference>
    <dbReference type="SMART" id="SM00320">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50978">
        <property type="entry name" value="WD40 repeat-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00678">
        <property type="entry name" value="WD_REPEATS_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50082">
        <property type="entry name" value="WD_REPEATS_2"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50294">
        <property type="entry name" value="WD_REPEATS_REGION"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="2 4" id="KW-0853">WD repeat</keyword>
    <feature type="domain" description="Autophagy-related protein 16" evidence="6">
        <location>
            <begin position="16"/>
            <end position="206"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="4">
        <location>
            <begin position="318"/>
            <end position="351"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="4">
        <location>
            <begin position="362"/>
            <end position="403"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="4">
        <location>
            <begin position="404"/>
            <end position="445"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="4">
        <location>
            <begin position="462"/>
            <end position="484"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="4">
        <location>
            <begin position="540"/>
            <end position="571"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="4">
        <location>
            <begin position="573"/>
            <end position="607"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="202"/>
            <end position="231"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="246"/>
            <end position="272"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="202"/>
            <end position="229"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00009271"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022574"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="Pfam" id="PF08614"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001754.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="607" mass="68431" checksum="CDE4120362C316EB" modified="2019-05-08" version="1">MTSGLKAATFPTWKRHIAEELRRRDRLQRQAFEEIILQYNKLLEKTDLHAVLADKLQAEKYDLQSRHEISPGHDGTRNEAQLQEMAQLRIKHQEELTELHKKRGELAQSVIDLNNQMQQKDREMQINEAKIAEYMQKISELETECQELRSKLQELERANQTLKDEYDALQITFTALEEKLRKTTEDNQELVTRWMAEKAQEANRLNAENEKDSRRRQAKLQKELAEAAKEPLPVEQDDDIEVLVDETSDPAEETSPVRAVSRTSSKRLSQPAGGLLDSITNIFGRRSLSSFPAPQENVDVHPGSSKEVRVPTTAVYVFDAHDGEVNAVQFSPASRLLATGGMDRRVKLWEVFGDRCELKSSLSGSNAGITSIEFDSAGSYLLAASNDFASRIWTVDDYRLRHTLTGHSGKVLSAKFLLDNARIVSGSHDRTLKLWDLRSKVCIKTVFAGSSCNDIVCTEQCVMSGHFDKKIRFWDIRTESIVQELELFGRITALDLNPERTELLTCSRDDLLKIIDLRVNAVKQTFSAQGFKCGSDWTRVVFSPDGSYVAGGSADGVLYIWNVLTGKVERTLAKHHSSSINAVAWSPAGAHVVSVDKGNKAVLWSEF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJ99</accession>
    <name>A0A452GJ99_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Diaphanous related formin 3</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000001977.1">
        <property type="protein sequence ID" value="ENSGAGP00000001742.1"/>
        <property type="gene ID" value="ENSGAGG00000001377.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005884">
        <property type="term" value="C:actin filament"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030041">
        <property type="term" value="P:actin filament polymerization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.630:FF:000001">
        <property type="entry name" value="Diaphanous related formin 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.630">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.30.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.2220">
        <property type="entry name" value="Formin, FH2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014767">
        <property type="entry name" value="DAD_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010465">
        <property type="entry name" value="Drf_DAD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015425">
        <property type="entry name" value="FH2_Formin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042201">
        <property type="entry name" value="FH2_Formin_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051412">
        <property type="entry name" value="Formin_Homology_Diaphanous_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45691">
        <property type="entry name" value="PROTEIN DIAPHANOUS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45691:SF9">
        <property type="entry name" value="PROTEIN DIAPHANOUS HOMOLOG 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06345">
        <property type="entry name" value="Drf_DAD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02181">
        <property type="entry name" value="FH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00498">
        <property type="entry name" value="FH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF101447">
        <property type="entry name" value="Formin homology 2 domain (FH2 domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51231">
        <property type="entry name" value="DAD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51444">
        <property type="entry name" value="FH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="FH2" evidence="4">
        <location>
            <begin position="87"/>
            <end position="482"/>
        </location>
    </feature>
    <feature type="domain" description="DAD" evidence="3">
        <location>
            <begin position="505"/>
            <end position="535"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1"/>
            <end position="81"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="477"/>
            <end position="560"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="358"/>
            <end position="385"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="1"/>
            <end position="14"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="2">
        <location>
            <begin position="25"/>
            <end position="48"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="2">
        <location>
            <begin position="59"/>
            <end position="80"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="477"/>
            <end position="504"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS51231"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS51444"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001742.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="7">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="560" mass="64332" checksum="A7B046EAFAB85184" modified="2019-05-08" version="1">VHFLSSIQGGTSIPPSDAAAHEQLLPPPPLAPFNGAAPPPPPPPPPPFMNGSGMAPSFGAPPPPPPPGFPGVQWSPPPRTLPFGLKPKKEFKPEITMKRLNWSKIRPNEMNENCFWLKADEDKYESADLLCKLELTFCCQKKVKKEEDDFEEKKSIKKRIKELKVLDPKIAQNLSIFLGSFRVPYEEIKMMILEVDETQLAETMIQNLIKHLPEQEQLNTLSKFKSEYNNLSEPEQFGIVMSNVKRLQTRLSAILFKLQFEEQVNNIKPDIMAVSAACEEIKKSKSFSKLLELVLLIGNYMNAGSRNAQTFGFNLSSLCKLSHFQYIYILYFCVCVCVCVCXKYPDVLNFTEDLQHLDKASKVSAENLEKNLKQMERQLQQLEKDLKIFPPPDDIHDKFVTKIFAVYAREHYQILTTMHENMEKLYQNVMGYYAIDLKKVSVEEFLTDLNNFRTMFLQAAKENIRRKEAEEKQRRARIAKEKAEKEKLERQQKKKRLLEMKTEGDETGVMDSLLEALQSGAAFRDRRKRTPRPKDILQNLSPTNQRPVLKPCNHGNKPYS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZQ9</accession>
    <name>A0A452GZQ9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">U4/U6.U5 small nuclear ribonucleoprotein 27 kDa protein</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="9">U4/U6.U5 tri-snRNP-associated protein 3</fullName>
        </alternativeName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">May play a role in mRNA splicing.</text>
    </comment>
    <comment type="subunit">
        <text evidence="4">Part of a tri-snRNP complex.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the SNUT3 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZQ9"/>
    <dbReference type="STRING" id="38772.ENSGAGP00000007686"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008872.1">
        <property type="protein sequence ID" value="ENSGAGP00000007688.1"/>
        <property type="gene ID" value="ENSGAGG00000006150.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071011">
        <property type="term" value="C:precatalytic spliceosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006397">
        <property type="term" value="P:mRNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008380">
        <property type="term" value="P:RNA splicing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013957">
        <property type="entry name" value="SNRNP27"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31077">
        <property type="entry name" value="U4/U6.U5 SMALL NUCLEAR RIBONUCLEOPROTEIN 27 KDA PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31077:SF1">
        <property type="entry name" value="U4_U6.U5 SMALL NUCLEAR RIBONUCLEOPROTEIN 27 KDA PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08648">
        <property type="entry name" value="SNRNP27"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0507">mRNA processing</keyword>
    <keyword evidence="7" id="KW-0508">mRNA splicing</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="U4/U6.U5 small nuclear ribonucleoprotein 27kDa protein" evidence="11">
        <location>
            <begin position="90"/>
            <end position="145"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1"/>
            <end position="90"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="10">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="10">
        <location>
            <begin position="31"/>
            <end position="51"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="58"/>
            <end position="90"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00003632"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00008218"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00011825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00014357"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022664"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023187"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00031864"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF08648"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007688.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="147" mass="17836" checksum="6F9B11311167EAD7" modified="2019-05-08" version="1">MKPERRRSRSASRERERRRRERSRSRERDRRRSRSRSPHRRRSRSPRRHRSSSSSPSRQKERRDDEKKENKDTKSKEHQITEEDMEGKTEEEIEMMKTMGFASFDTTKGKKVDGSVNAYAINVSQKRKYRQYMNRKGGFNRPLDFIA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZT0</accession>
    <name>A0A452GZT0_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GZT0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008977.1">
        <property type="protein sequence ID" value="ENSGAGP00000007788.1"/>
        <property type="gene ID" value="ENSGAGG00000006197.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016491">
        <property type="term" value="F:oxidoreductase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.20.20.70">
        <property type="entry name" value="Aldolase class I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013785">
        <property type="entry name" value="Aldolase_TIM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050139">
        <property type="entry name" value="GMP_reductase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43170">
        <property type="entry name" value="GMP REDUCTASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43170:SF3">
        <property type="entry name" value="GMP REDUCTASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51412">
        <property type="entry name" value="Inosine monophosphate dehydrogenase (IMPDH)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007788.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="123" mass="14150" checksum="0D7272780ABACB0F" modified="2019-05-08" version="1">MPHVEADLKLDFHDVLLRPKRSSLKSRAEVDLIRTFTFRNSKQTYTGIPIMVANMDTVGTFEMAKVMTKVSADFHLSLQKQGNHLFGDRMSCSKSITYFIYVHWELGCKGNCCDKIKNRMVKT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZT1</accession>
    <name>A0A452GZT1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Fibrillar collagen NC1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000008894.1">
        <property type="protein sequence ID" value="ENSGAGP00000007709.1"/>
        <property type="gene ID" value="ENSGAGG00000005830.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005581">
        <property type="term" value="C:collagen trimer"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031012">
        <property type="term" value="C:extracellular matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030020">
        <property type="term" value="F:extracellular matrix structural constituent conferring tensile strength"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030198">
        <property type="term" value="P:extracellular matrix organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.1000:FF:000001">
        <property type="entry name" value="Collagen alpha-1 type I chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.1000">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008160">
        <property type="entry name" value="Collagen"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050149">
        <property type="entry name" value="Collagen_superfamily"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000885">
        <property type="entry name" value="Fib_collagen_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24023">
        <property type="entry name" value="COLLAGEN ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24023:SF1109">
        <property type="entry name" value="COLLAGEN ALPHA-4(IV) CHAIN-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01410">
        <property type="entry name" value="COLFI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01391">
        <property type="entry name" value="Collagen"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00038">
        <property type="entry name" value="COLFI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51461">
        <property type="entry name" value="NC1_FIB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0106">Calcium</keyword>
    <keyword evidence="7" id="KW-0176">Collagen</keyword>
    <keyword evidence="8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="9" id="KW-0379">Hydroxylation</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <feature type="domain" description="Fibrillar collagen NC1" evidence="11">
        <location>
            <begin position="1072"/>
            <end position="1305"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1"/>
            <end position="1075"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="18"/>
            <end position="33"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="10">
        <location>
            <begin position="42"/>
            <end position="54"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="233"/>
            <end position="249"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="422"/>
            <end position="432"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="494"/>
            <end position="504"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="10">
        <location>
            <begin position="505"/>
            <end position="514"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="516"/>
            <end position="527"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="10">
        <location>
            <begin position="538"/>
            <end position="547"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="548"/>
            <end position="558"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="582"/>
            <end position="593"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="632"/>
            <end position="647"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="680"/>
            <end position="689"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="10">
        <location>
            <begin position="700"/>
            <end position="709"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="869"/>
            <end position="878"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="899"/>
            <end position="920"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="933"/>
            <end position="947"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="10">
        <location>
            <begin position="974"/>
            <end position="987"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="10">
        <location>
            <begin position="1017"/>
            <end position="1032"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023119"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023278"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS51461"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007709.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1305" mass="124950" checksum="3A5DDCB15411C550" modified="2019-05-08" version="1">MAGLDEKSGISSQMGFMPGAVGPVGPRGPQGLQGQQGGVGPTGPPGEPGEPGPMGPAGARGPEGPPGKPGEDGEPGRSGQPGETGFPGSTGSRGFPGAPGLPGLKGHRGHKGLDGPKGEVGATGSKGEAGPTGPMGPTGPIGPRGMSGERGRIGPQGAPGARGSHGMPGKPGPVGPLGIIGSPGFPGPPGVKGEAGPTGARGPEGPQGQRGETGPPGPAGSQGLPGTVGTDGSPGAKGPTGSPGTSGPPGLAGPPGSPGPQGSTGPPGIRGQAGDPGVPGFKGEAGPKGEPGPHGPQGPIGSVGEEGKRGPRGDPGSVGPPGPVGERGAPGNRGFPGSDGLPGPKGAQGERGPAGSSGPKGSQGDPGRTGEPGLPGARGLTGNPGVQGPEGKIGPLGAPGEDGRPGPAGSIGIRGQPGSMGLPGPKGSSGDPGKPGEAGNAGVSGQRGAPGKDGEVGPSGPVGPPGLAGERGEQGPPGPTGFQGLPGPPGPPGEAGKPGDQGVPGDPGAGGPLGPRGERGNPGERGEPGAAGLQGEKGMAGGHGPDGPKGNPGPTGTPGDPGPPGLQGMPGERGIAGTPGPKGDRGGVGEKGSEGTAGNDGARGLPGPIGPMGPAGPTGEKGEPGPRGLVGPPGSRGNPGSRGENGPTGAVGFAGPPGPDGQPGVKGEPGEPGQKGDAGSPGPQGLSGSPGPPGPHGVPGLKGGRGTQGPPGATGFPGSAGRVGPPGPTGALGPAGPIGDPGKEGPPGLRGDPGAHGRVGDRGPAGPPGGPGDKGDSGEDGQPGPDGPPGPAGTTGQRGIVGMPGQRGERGMPGLPGPGGTPGKQGSTGPPGDKGPSGPIGQPGASGPVGEPGPEGPAGNDGTPGRDGAVGERGDRGEPGPAGLPGAQGGPGTPGPVGPTGEAGQRGEPGSRGPVGPPGRAGKRGLPGPQGPRGDKGDHGDRGDRGQKGHRGFTGLQGLPGPPGPIGEQGSPGIPGPFGPRGPPGPVGPSGKDGNPGPHGPLGPPGVRGSLGEAGPEGPPGDPGPPGPPGPPGHLTAAIGDIMGHFDESMADPLPEFTEDEAAPDDNNKTDPGVHATLKSLSSQIETMRSPDGSKKHPARTCDDLKLCHPSKKSGEYWIDPNQGCVEDALKVYCNMETGETCISANPSSIPRKMWWTSRSPDLKPVWYGLDMNRGSQFVYGDSESPNTAVTQMTFLRLLSKEASQNITYHCKNSIGYMDDQSKNLKKAVILKGANDLEIKAEGNNRFRYTVLQDSCSKRNGNIGKTVFEYRTQNVARLPIVDIAPVDIGNTDQEFGIEIGPVCFV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZT6</accession>
    <name>A0A452GZT6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Myosin VB</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="4">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <comment type="caution">
        <text evidence="4">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZT6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008901.1">
        <property type="protein sequence ID" value="ENSGAGP00000007714.1"/>
        <property type="gene ID" value="ENSGAGG00000005542.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016459">
        <property type="term" value="C:myosin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003779">
        <property type="term" value="F:actin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003774">
        <property type="term" value="F:cytoskeletal motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051020">
        <property type="term" value="F:GTPase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd23767">
        <property type="entry name" value="IQCD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd15477">
        <property type="entry name" value="Myo5b_CBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.190:FF:000006">
        <property type="entry name" value="Myosin VA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.190:FF:000001">
        <property type="entry name" value="unconventional myosin-Va"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.190">
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.1590">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002710">
        <property type="entry name" value="Dilute_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000048">
        <property type="entry name" value="IQ_motif_EF-hand-BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058662">
        <property type="entry name" value="Myo5a/b_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037990">
        <property type="entry name" value="Myo5b_CBD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052072">
        <property type="entry name" value="Vascular_dev_regulator"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16027:SF6">
        <property type="entry name" value="DILUTE DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16027">
        <property type="entry name" value="DILUTE DOMAIN-CONTAINING PROTEIN YPR089W"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01843">
        <property type="entry name" value="DIL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00612">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25966">
        <property type="entry name" value="Myo5a"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01132">
        <property type="entry name" value="DIL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00015">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51126">
        <property type="entry name" value="DILUTE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="2" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="3 5" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-0505">Motor protein</keyword>
    <keyword evidence="4" id="KW-0518">Myosin</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0677">Repeat</keyword>
    <keyword evidence="7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="12"/>
            <end position="33"/>
        </location>
    </feature>
    <feature type="domain" description="Myosin motor" evidence="9">
        <location>
            <begin position="51"/>
            <end position="114"/>
        </location>
    </feature>
    <feature type="domain" description="Dilute" evidence="8">
        <location>
            <begin position="878"/>
            <end position="1155"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="447"/>
            <end position="477"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="5">
        <location>
            <begin position="501"/>
            <end position="619"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="5">
        <location>
            <begin position="685"/>
            <end position="792"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="456"/>
            <end position="476"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51126"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007714.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="12">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1200" mass="140310" checksum="52E73751D0A2CAC0" modified="2019-05-08" version="1">MAETIMVYIVKVSIVQFAAYPCFCGVVMVYQVAVGGAHIPVFLKLGSLLVWSYHDFFNRYRVLMKKREISKNDKKLICKTLLENLIKDPDKFQFGRTKIFFRAGQVAYLEKLRADKFRAATIMIQKTVRGWLQRVKYRRMRNAAITIQRYTRGHLARRLADHLRKTRAAIIFQKQYRMLRIFHAFQKIRKATITIQAFTRGMFVRKIYHKILLEHKATIIQKHARAWLARKQFLHFKCAAVVIQCYFRRMKARQELKKLKIEARSAQHLKKLNIGMENKVVQLQRKIDDQNKEYMHLNEQLSALTSAYATDVEKLKKELEQYRQNQGDTNQLVNLKKEIESLRLELEKSHSERKIIEDTYDKEKDLLRKRVSDLEEENALLKQEKEELNNRILCQSEDEFAQNTVKENLLTKKELEEERSRYQNLVKEYSRLEQRYDNLRDEMTIIKQTPGHRRNPSNQSSLESDSNCPSVSTSEIGDTEDAIQQVEDVGLDKAAMDMTLFLKLQKRVRELEQERKKLQTQLEKKKSQVKDVIETKNEMDSDRDTDLAYNNLKRQELESENKKLKNELNELRKTIVDRVTQNNSSSDVQDSYTLLLNQLKSANEELEVRKEEVLILRAQIIKAAQQKDIGKNMDSNINTYTSWPNSDKHVDAEDAIEAYHGVCQTNRKTEDWGYLNEDGELGLAYQGLKQVARLLEAQLQDQRREHEEEVETLKTQMETLREEMEKQQQAFFQTLQLPPEAQVEFGIQQEIARLTNENLDLKELLEKLEKNEKKLKKQLKIYMKKVQDFEATQATAQTERRRHELTRQVTVQRKEKDFQGMLEYYKEDEPLLIRNLITDLKPQTVSATVPCLPAYILYMCIRHADYINDDQKVHSLLTSTINGIKKVLKKHNDDFQMTSFWLANTCRLLHCLKQYSGDTGFMTQNTAKQNEHCLKNFDLTEYRQVLSDLSIQIYQQLFKIAEGMLQPMIVSAMLESESIQGLSGVKPMGYRNRSSSMVDGDNSYSLDAVIRQLNTFHSIMCDQGLDPEIIQQVFKQLFYMINAVTLNNLLLRKDVCSWSTGMQLRYNISQLEEWLRGKNLQQSGAAETVEPLIQAAQLLQLKKKTSEDAEAICSLCTSLSTQQIVKILNLYTPVNEFEERVTVAFIRNIQANLQERNDPPQLLLDYKYMFPVLFPFSPSSITMDSIHIPASLNLEFLNEV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZT7</accession>
    <name>A0A452GZT7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Spondin-1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="11">F-spondin</fullName>
        </alternativeName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZT7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008775.1">
        <property type="protein sequence ID" value="ENSGAGP00000007602.1"/>
        <property type="gene ID" value="ENSGAGG00000005723.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031012">
        <property type="term" value="C:extracellular matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
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    <keyword evidence="8" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="9" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="4" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="5" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0964">Secreted</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
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        <source>
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        <source>
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    </evidence>
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        <source>
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    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="506" mass="57303" checksum="F23CED0CF04873C4" modified="2019-05-08" version="1">SAEFSVDRTRHLMSFLTMLGPSPDWNVGLSAEDLCTKDCGWVQKVIQDLIPWDAGTDSGVTYESPNKPTIPQEKIRPLTSLDHPQSPFYDPEGGSIAVVARVVIERIARKGEQCNIIPDNVDDIVADLVPEEKEEDDTPETCIYSNWSPWSACSSSTCDKGKRMRQRMLKAQLDLSVPCPDTQDFQPCMGPGCSDEDGSTCMMSEWITWSPCTVSCGMGMRSRERYVKQFPDDGSMCKVPTEETEKCIVNEECSPSSCLVTEWAEWNECSASCGIGMKKRHRMIKMTPADGSMCNTETTEVEKCMMPECHTIPCMLSPWSEWSDCSVTCGKGMRTRQRMLKSAAELGDCNEELEQVEKCMLPECPIDCELTEWSKWSECNKSCGKGHIIRTRMIKMEPQFGGAPCPETVQRKKCRIRKCLRGPSLEKRRWKEAREKRRSEHTKEDVEGEPYPGCKMRPWTAWSECTKLCGGGIQERFMTVKKRFKSTQFTSCKDKKEIRACNVHPC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZT9</accession>
    <name>A0A452GZT9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Voltage-dependent T-type calcium channel subunit alpha-1G</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="21">Voltage-gated calcium channel subunit alpha Cav3.1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="26 27">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="27" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="26" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="26" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>Ca(2+)(in) = Ca(2+)(out)</text>
            <dbReference type="Rhea" id="RHEA:29671"/>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="19">Belongs to the calcium channel alpha-1 subunit (TC 1.A.1.11) family. CACNA1G subfamily.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000008988.1">
        <property type="protein sequence ID" value="ENSGAGP00000007799.1"/>
        <property type="gene ID" value="ENSGAGG00000006199.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005891">
        <property type="term" value="C:voltage-gated calcium channel complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008331">
        <property type="term" value="F:high voltage-gated calcium channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008332">
        <property type="term" value="F:low voltage-gated calcium channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098703">
        <property type="term" value="P:calcium ion import across plasma membrane"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
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    </dbReference>
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    </dbReference>
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        <property type="entry name" value="Voltage-gated potassium channels. Chain C"/>
        <property type="match status" value="4"/>
    </dbReference>
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        <property type="entry name" value="Ion_trans_dom"/>
    </dbReference>
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        <property type="entry name" value="VDCC_alpha-1_subunit"/>
    </dbReference>
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        <property type="entry name" value="VDCC_T_a1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027359">
        <property type="entry name" value="Volt_channel_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45628">
        <property type="entry name" value="VOLTAGE-DEPENDENT CALCIUM CHANNEL TYPE A SUBUNIT ALPHA-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45628:SF33">
        <property type="entry name" value="VOLTAGE-DEPENDENT T-TYPE CALCIUM CHANNEL SUBUNIT ALPHA-1G"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00520">
        <property type="entry name" value="Ion_trans"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01629">
        <property type="entry name" value="TVDCCALPHA1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81324">
        <property type="entry name" value="Voltage-gated potassium channels"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="11" id="KW-0106">Calcium</keyword>
    <keyword evidence="8" id="KW-0107">Calcium channel</keyword>
    <keyword evidence="7" id="KW-0109">Calcium transport</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="22" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="16" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="17" id="KW-0407">Ion channel</keyword>
    <keyword evidence="14" id="KW-0406">Ion transport</keyword>
    <keyword evidence="15 24" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="27" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0677">Repeat</keyword>
    <keyword evidence="9 24" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="13 24" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <keyword evidence="12" id="KW-0851">Voltage-gated channel</keyword>
    <feature type="transmembrane region" description="Helical" evidence="24">
        <location>
            <begin position="36"/>
            <end position="53"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="24">
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    </feature>
    <feature type="transmembrane region" description="Helical" evidence="24">
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    <feature type="transmembrane region" description="Helical" evidence="24">
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    <feature type="transmembrane region" description="Helical" evidence="24">
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    <feature type="transmembrane region" description="Helical" evidence="24">
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            <begin position="711"/>
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    <feature type="transmembrane region" description="Helical" evidence="24">
        <location>
            <begin position="741"/>
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        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="24">
        <location>
            <begin position="832"/>
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        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="24">
        <location>
            <begin position="908"/>
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    <feature type="transmembrane region" description="Helical" evidence="24">
        <location>
            <begin position="1221"/>
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    </feature>
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            <begin position="1259"/>
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        <location>
            <begin position="1356"/>
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            <begin position="1772"/>
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            <begin position="31"/>
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        </location>
    </feature>
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    </feature>
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            <begin position="426"/>
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    </feature>
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            <begin position="469"/>
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        </location>
    </feature>
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    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="23">
        <location>
            <begin position="1029"/>
            <end position="1056"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="23">
        <location>
            <begin position="1131"/>
            <end position="1141"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="23">
        <location>
            <begin position="1163"/>
            <end position="1172"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="23">
        <location>
            <begin position="2005"/>
            <end position="2034"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022568"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022673"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022882"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00036634"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00061006"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00069348"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00078683"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="Pfam" id="PF00520"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007799.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2361" mass="264946" checksum="865BBD56904DB0CF" modified="2019-05-08" version="1">MGEHDVSSFSRPFPACVLKGLLISLTDLLCTWFERLSMLVILLNCVTLGMFHPCEDMACDSPRCRILQSFDDFIFAFFAVEMIVKMIALGIFGKKCYLGDTWNRLDFFIVIAGMLEYSLDLQNVSFSAVRTVRVLRPLRAINRVPSMRILVTLLLDTLPMLGNVLLLCFFVFFIFGIVGVQLWAGLLRNRCFLPENFSLPYTVELERYYQTENEDENPFICSQPRENGMRYCRSIPTRREEGLECTLDFYAYNDTTNTSCVNWNQYYTNCSAGEHNPFKGAINFDNIGYAWIAIFQVITLEGWVDIMYFVMDAHSFYNFIYFILLIIVGSFFMINLCLVVIATQFSETKQRESQLMKEQRIRYLSNASTLASFSEPGSCYDELLKYLVYVARKASKQLVKGYREAGFKMGLLSSPRNKSGAERQPCRHRQRKRSSVHHLIHHHHHHHHHYHLGNGNLRAPRASPEISDVETSSLHNGTNRLMLPPTTSNPHPTPNAAASNTESVHSIYHADCHFEPIHCRSSLPQPGLSLPSPEGLPKNVVGRKVYPTVHPSTSHEMLKEKNVADSTVNPGASMLTNLNIPPGPYSTMHKLLETQSTGFFSVHSTDTGAGFPGACKSSCKISNQCGKLDSGSCNPDSCPYCIKTLARDLELTDYETADSDSEGVYEFTQDAHYSDQRDPQRGKTKGRKTSQILAFWKVVCETFRKIVDSKYFGRGIMIAILINTLSMGIEYHEQPEELTNALEISNIVFTSLFALEMLLKVLVYGPFGYIKNPYNIFDGIIVVISVWEIVGQQGGGLSVLRTFRLMRVLKLVRFMPALQRQLVVLMKTMDNVATFCMLLMLFIFIFSILGMHLFGCKFASERDGDTLPDRKNFDSLLWAIVTVFQILTQEDWNKVLYNGMASTSSWAALYFIALMTFGNYVLFNLLVAILVEGFQTEGDASKSDSEGDLFPHSLEEEGGLKKNLSNPALMVLSDLPELKKSLTPPLIIHTAATPMPMPKSAMFGDAARGYDSRRASSVSMDPNAYELKSPSSIQSSPHSPWSAASSWNSRRSSWNSLGRAPSLKRRSQSGERKSLLSGNGKESSEEGESSDEERSSRAGSVNGSLPHRMASLETKGSFDLQDTLQVPSLYRTSSMHSSRTAASEHQDCNGKSSPGVLLQQLPLDDDDGDDEGNLSKSERVKAWIRAHLPSWCKERDSWSIYIFAPHSKFRLMCNKIITHKMFDHIVLVIIFLNCITIAMERPKIEPHSAERIFLTLSNYIFTVIFLAEMTVKVVALGLCFGEKAYLKSSWNVLDGVLVLISVIDILVSMVSDSGTKILGMLRVLRLLRTLRPLRVISRAQGLKLVVETLMSSLKPIGNIVVICCAFFIIFGILGVQLFKGKFFVCQGEDTRNITNKSDCAEASYKWVRHKYNFDNLGQALMSLFVLASKDGWVDIMYDGLDAVGVDQQPVMNYNPWMLLYFISFLLIVAFFVLNMFVGVVVENFHKCRQHQEEEEAKRREEKRLRRLEKKRRNLMLDDVIMESSASAVQEAQCKPYYSDYSRFRLLIHQMCTSHYLDLFITGVIGLNVITMAMEHYQQPKVLDEALKICNYIFTVIFVLESVFKLIAFGFRRFFQDRWNQLDLAIVLLSIMGITLEEIEVNASLPINPTIIRIMRVLRIARVLKLLKMAVGMRALLDTVMQALPQVGNLGLLFMLLFFIFAALGVELFGDLECDDTHHCEGLGRHATFRNFGMAFLTLFRVSTGDNWNGIMKDTLRDCDPESTCYNTVISPMYFVSFVLTAQFVLVNVVIAVLMKHLEESNKEAKEEAELEAELEMEMKTITPGPIPASDLFAWTGGNGGERPESPRGCTNPMQFKVDSQLSLFYPMERHLFDTISLLIQESLEGELKLMDNLSGSVCHHYALPAPEYYNSEKQIPLAEMEALSLTSDILSEKSWSLALTDDSFPDDTNTLLLNALESNTDHLSRDDTLTLSPSKDLLSVRKPSVGRTHSLPNDSYMFQPPNAGSCANSQAARNASHQKSQSGSTASVQSQPANTSSLLQIPKDHFHHIRPHGNLDWESKSKIPPTVHSPSAQRLLRRQVAIRNDSLDMYCSESKENLHAEVNELSDTNVPAGSTEDPSEASESACWGQSSVRTQHSHNQYNISKHTPALCACTDSYQETLRDSIDQDVSEINSSLEPFSSGTCTALSACSVVPPLTPKRNIGNSSNVTLKDLKKYYSVDTQGLLRKPSSWLDDQRRHSIEICSMQNSPQHHSTSSSSGFISQVLSEMEGLQGARQKKKLSPPCISIDPPDGQSLLPRGSHGISSPSADICLRRRAPSCDSKDSMDIGDSLLPDSMSTSPTPKTDLLTLPSFSFDQAEMDP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZU1</accession>
    <name>A0A452GZU1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Myosin VB</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="4">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <comment type="caution">
        <text evidence="4">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZU1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008906.1">
        <property type="protein sequence ID" value="ENSGAGP00000007719.1"/>
        <property type="gene ID" value="ENSGAGG00000005542.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016459">
        <property type="term" value="C:myosin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003779">
        <property type="term" value="F:actin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003774">
        <property type="term" value="F:cytoskeletal motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051020">
        <property type="term" value="F:GTPase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd23767">
        <property type="entry name" value="IQCD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd15477">
        <property type="entry name" value="Myo5b_CBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.190:FF:000006">
        <property type="entry name" value="Myosin VA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.190:FF:000001">
        <property type="entry name" value="unconventional myosin-Va"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.190">
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.1590">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002710">
        <property type="entry name" value="Dilute_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000048">
        <property type="entry name" value="IQ_motif_EF-hand-BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058662">
        <property type="entry name" value="Myo5a/b_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037990">
        <property type="entry name" value="Myo5b_CBD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052072">
        <property type="entry name" value="Vascular_dev_regulator"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16027:SF6">
        <property type="entry name" value="DILUTE DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16027">
        <property type="entry name" value="DILUTE DOMAIN-CONTAINING PROTEIN YPR089W"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01843">
        <property type="entry name" value="DIL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00612">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25966">
        <property type="entry name" value="Myo5a"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01132">
        <property type="entry name" value="DIL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00015">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51126">
        <property type="entry name" value="DILUTE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="2" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="3 5" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-0505">Motor protein</keyword>
    <keyword evidence="4" id="KW-0518">Myosin</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0677">Repeat</keyword>
    <keyword evidence="7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="12"/>
            <end position="33"/>
        </location>
    </feature>
    <feature type="domain" description="Myosin motor" evidence="9">
        <location>
            <begin position="51"/>
            <end position="114"/>
        </location>
    </feature>
    <feature type="domain" description="Dilute" evidence="8">
        <location>
            <begin position="829"/>
            <end position="1106"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="447"/>
            <end position="477"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="5">
        <location>
            <begin position="501"/>
            <end position="609"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="5">
        <location>
            <begin position="640"/>
            <end position="747"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="456"/>
            <end position="476"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51126"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007719.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="12">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1151" mass="134699" checksum="09C4F397A11BE8B1" modified="2019-05-08" version="1">MAETIMVYIVKVSIVQFAAYPCFCGVVMVYQVAVGGAHIPVFLKLGSLLVWSYHDFFNRYRVLMKKREISKNDKKLICKTLLENLIKDPDKFQFGRTKIFFRAGQVAYLEKLRADKFRAATIMIQKTVRGWLQRVKYRRMRNAAITIQRYTRGHLARRLADHLRKTRAAIIFQKQYRMLRIFHAFQKIRKATITIQAFTRGMFVRKIYHKILLEHKATIIQKHARAWLARKQFLHFKCAAVVIQCYFRRMKARQELKKLKIEARSAQHLKKLNIGMENKVVQLQRKIDDQNKEYMHLNEQLSALTSAYATDVEKLKKELEQYRQNQGDTNQLVNLKKEIESLRLELEKSHSERKIIEDTYDKEKDLLRKRVSDLEEENALLKQEKEELNNRILCQSEDEFAQNTVKENLLTKKELEEERSRYQNLVKEYSRLEQRYDNLRDEMTIIKQTPGHRRNPSNQSSLESDSNCPSVSTSEIGDTEDAIQQVEDVGLDKAAMDMTLFLKLQKRVRELEQERKKLQTQLEKKKSQRAPEEMAPVIICLFERQELESENKKLKNELNELRKTIVDRVTQNNSSSDVQDSYTLLLNQLKSANEELEVRKEEVLILRAQIIKAAQQKDIGKNMTEDWGYLNEDGELGLAYQGLKQVARLLEAQLQDQRREHEEEVETLKTQMETLREEMEKQQQAFFQTLQLPPEAQVEFGIQQEIARLTNENLDLKELLEKLEKNEKKLKKQLKIYMKKVQDFEGNSGFRRRHELTRQVTVQRKEKDFQGMLEYYKEDEPLLIRNLITDLKPQTVSATVPCLPAYILYMCIRHADYINDDQKVHSLLTSTINGIKKVLKKHNDDFQMTSFWLANTCRLLHCLKQYSGDTGFMTQNTAKQNEHCLKNFDLTEYRQVLSDLSIQIYQQLFKIAEGMLQPMIVSAMLESESIQGLSGVKPMGYRNRSSSMVDGDNSYSLDAVIRQLNTFHSIMCDQGLDPEIIQQVFKQLFYMINAVTLNNLLLRKDVCSWSTGMQLRYNISQLEEWLRGKNLQQSGAAETVEPLIQAAQLLQLKKKTSEDAEAICSLCTSLSTQQIVKILNLYTPVNEFEERVTVAFIRNIQANLQERNDPPQLLLDYKYMFPVLFPFSPSSITMDSIHIPASLNLEFLNEV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZU3</accession>
    <name>A0A452GZU3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">Receptor-type tyrosine-protein phosphatase F</fullName>
            <ecNumber evidence="3">3.1.3.48</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="25">
            <text>O-phospho-L-tyrosyl-[protein] + H2O = L-tyrosyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:10684"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="EC" id="3.1.3.48"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the protein-tyrosine phosphatase family. Receptor class 2A subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.48" evidence="3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008905.1">
        <property type="protein sequence ID" value="ENSGAGP00000007718.1"/>
        <property type="gene ID" value="ENSGAGG00000005093.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="CDD" id="cd05738">
        <property type="entry name" value="IgI_2_RPTP_IIa_LAR_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05739">
        <property type="entry name" value="IgI_3_RPTP_IIa_LAR_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000010">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000027">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000036">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000066">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000144">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000015">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000082">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000128">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000098">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase F isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000353">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase F isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013098">
        <property type="entry name" value="Ig_I-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050713">
        <property type="entry name" value="RTP_Phos/Ushers"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46957">
        <property type="entry name" value="CYTOKINE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46957:SF9">
        <property type="entry name" value="PROTEIN-TYROSINE-PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07679">
        <property type="entry name" value="I-set"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00014">
        <property type="entry name" value="FNTYPEIII"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="13" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="15" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="4" id="KW-0358">Heparin-binding</keyword>
    <keyword evidence="8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="16" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="12 20" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="14" id="KW-0675">Receptor</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="5 20" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 20" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="1146"/>
            <end position="1169"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="21">
        <location>
            <begin position="24"/>
            <end position="119"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="21">
        <location>
            <begin position="127"/>
            <end position="209"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="216"/>
            <end position="307"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="312"/>
            <end position="406"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="410"/>
            <end position="499"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="504"/>
            <end position="601"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="606"/>
            <end position="710"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="711"/>
            <end position="805"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="809"/>
            <end position="901"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="905"/>
            <end position="989"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="586"/>
            <end position="607"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1255"/>
            <end position="1282"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004167"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010504"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00013064"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022674"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
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            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
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            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
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        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00044158"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00051722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
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        </source>
    </evidence>
    <sequence length="1319" mass="145541" checksum="0236C087AD664450" modified="2019-05-08" version="1">MPSAAWICCMLFPVAEDHLPAGFPTIDMGPQLKVVEKARTATMLCAASGNPDPEISWFKDFLPVDTATSNGRIKQLRSGGTPIRGALQIENSEESDQGKYECVATNSAGTRYSAPANLYVRVRRVAPRFSIPPSNHEVMPGGSVNLTCVAVGAPMPYVKWMAGVEELTKEDEMPVGRNVLELNNIVQSASYTCVAMSSLGMIEATAQITVKALPKPPTEPLVTETTATSVTLTWDSGNSDPISYYVIQYKPKSWEEGLFQEVDGVATTRYSIGGLSPFSEYEFRIIAVNNIGRGPPSELVEARTGEQAPSSPPLKVQARMLSASTMLVQWEQPEEPNGQIRGYRVYYTTDPHLPLSMWQKHNTDDSHLTTVGSLITGTTYSIRVLAFTSVGDGPPSDIIQVKTQQGVPAQPADFQAEAESDTRILLTWLPALQERITKYELLYWEGEDGSQRKVNFDPSSSYAVEGLKPDTLYKFQLGARSELGLGVYTPVLEARTAQSTPSAPPQEVECVSTSSTTIRVSWVPPPAQSRNGVITQYSIAYQAVDAEDGAKHVVDGIGRERSSWEIKDLEKWTEYKVWVRAHTDVGPGPESSPVHVRTDEDVPSAPPRKVEVESLNSTAIRVSWKLPLSNKQHGQIRGYQVTYVKLENNEPRGQPVIKDVMLAEAQVHTTPETIIGGLLPETTYSITVAAYTTKGDGARSKPKVVTTTGAVPGKPSMLISTTAMNTALIQWHPPKEMVGELLGYRLQYKRTDEEKMNIMDFEKNVHHYTVTSLHKGATYIFKLSAKNRAGPGEEFEKEITTTEDVPSGFPQNLRVVGLTTSTTEVAWDPPVLAERNGRITNYTVMYRDINSQQELTNITRETSIVLTNLRPDTTYDIKVRACNSKGVGPLSPSIQSRTMPVEQVFAKNFRVNAVMKTSVLLSWEVPDSYKSAVPFKILYNSQSVEVDGHSMKKLISDLQPDTEYSFVLMNRGSSAGGLQHRVSIHTAPDVLQSKPISTNKYMQEGKFTLTLPKVQTSAPVRWYYIVVVPVDQSTSSRTARWRTPDEMELDQLLEAISQGRRQRRQADRLKPYIAALVDMLPETFTLGDEKTYKGFYNKPLSEDLSYRCFVLASLEDGDTKRYAASPYSDEIVMEVSSAKQQDEPEMLWVMGPVLAVILIIIIVIAILLFKRLKANDGLRFSQEYEVSCREGDRCAWHCPEPGVAGRRWAMAASPRGGSVDSFHAHSFSVHLPRGYPLSWLVPQLQPPAASLVTRRSCPQAPRPGLQRPGVPHXGSPYSCSADTPQSHLASPCPPTHTLALLIPLSPALPLPGLPLVLLC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZU6</accession>
    <name>A0A452GZU6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">RRM domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GZU6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008792.1">
        <property type="protein sequence ID" value="ENSGAGP00000007618.1"/>
        <property type="gene ID" value="ENSGAGG00000006087.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005654">
        <property type="term" value="C:nucleoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.330:FF:000084">
        <property type="entry name" value="Serine/arginine-rich splicing factor 11 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.330">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012677">
        <property type="entry name" value="Nucleotide-bd_a/b_plait_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035979">
        <property type="entry name" value="RBD_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000504">
        <property type="entry name" value="RRM_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR32343">
        <property type="entry name" value="SERINE/ARGININE-RICH SPLICING FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR32343:SF71">
        <property type="entry name" value="SPLICING REGULATORY GLUTAMIC ACID AND LYSINE RICH PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00076">
        <property type="entry name" value="RRM_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54928">
        <property type="entry name" value="RNA-binding domain, RBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="RRM" evidence="1">
        <location>
            <begin position="22"/>
            <end position="85"/>
        </location>
    </feature>
    <evidence type="ECO:0000259" key="1">
        <source>
            <dbReference type="Pfam" id="PF00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007618.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="131" mass="14179" checksum="D826A93E4DD5BA5D" modified="2019-05-08" version="1">MNSGGIGVPLGFPLGPTSVIQVTNLSSAVTSEQMRTLFGFLGEIEELRLYPPDNAPLAFSSKVCYIKFREASSVGVAQHLTNTVFIDRALIVVPCAEVMQPTRTVVMCNTTFDHRENTALGKRPCLISFGS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZU8</accession>
    <name>A0A452GZU8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">BTB domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GZU8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008910.1">
        <property type="protein sequence ID" value="ENSGAGP00000007723.1"/>
        <property type="gene ID" value="ENSGAGG00000005859.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="CDD" id="cd18526">
        <property type="entry name" value="BACK_ABTB2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18350">
        <property type="entry name" value="BTB_POZ_ABTB2_BPOZ2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000045">
        <property type="entry name" value="Ankyrin repeat and BTB/POZ domain-containing protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.710.10:FF:000030">
        <property type="entry name" value="Ankyrin repeat and BTB/POZ domain-containing protein BTBD11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.710.10">
        <property type="entry name" value="Potassium Channel Kv1.1, Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048063">
        <property type="entry name" value="ABTB2_BTB_POZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052089">
        <property type="entry name" value="Ankyrin-BTB/POZ_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000210">
        <property type="entry name" value="BTB/POZ_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011333">
        <property type="entry name" value="SKP1/BTB/POZ_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46071">
        <property type="entry name" value="ANKYRIN REPEAT AND BTB/POZ DOMAIN-CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46071:SF3">
        <property type="entry name" value="ANKYRIN REPEAT AND BTB_POZ DOMAIN-CONTAINING PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00023">
        <property type="entry name" value="Ank"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00651">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00225">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54695">
        <property type="entry name" value="POZ domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50097">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2 3" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0677">Repeat</keyword>
    <feature type="repeat" description="ANK" evidence="3">
        <location>
            <begin position="342"/>
            <end position="374"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="3">
        <location>
            <begin position="388"/>
            <end position="420"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="3">
        <location>
            <begin position="427"/>
            <end position="453"/>
        </location>
    </feature>
    <feature type="domain" description="BTB" evidence="4">
        <location>
            <begin position="665"/>
            <end position="734"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50097"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007723.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="845" mass="94820" checksum="0FE2C7D8A65B97B6" modified="2019-05-08" version="1">MGAAAHQDSRYGYTCSCTALPREHSHSSALKCEYGHGTTAGRELSQRCRYSTSPWGLACSGHCLHWRFAVLSMHCIPQSPLQKKSVLSVFPSIPEWWSIPLPLRFHLQQACSLLSPGVLSLPAYFSPYNDGSISSDGRADAYAQLELRTLEQSLLATCVGSISELSDLVSRAMHHMQCLTTLHPGMSPIRQTRQQQPITWSPDALHTLYYFLRCPQMESMENPNLDPPRMTLNNERPFMLLPPLMEWMRVAITYAEHRRSLTVDSDDIRQTARLLLPGLDCEPRQLKPECCFSSFRRLDAKAATERFQQDLGFRMLNCGRTDLINQAIEALGPDGVNTMDDQGMTPLMYACAAGDEAMVQMLIDAGANLDIPVPGSNPRYPSVHPDSRHWTALTFAVLHGHISVVQLLLDAGAHVEGSAVNSSEDNYAETPLQLASAAGNYELVSLLLSRGADPLLSMLDVNGMSSSLHEDMNCFSHSAAHGHRNVLRKLLTQPQQSKGDVLSLEEILAEGVESDTSSQDSSNEGQVRLCKTRMKALQEAMYYSAEHGYVDITMELRVLGIPWKLHVWIESLRTTFYQSRYSVVQNLLREFVTIKEEDYNEELVNEGLQLMFDILKTSKNDSINQLLAAIFTHCYGSSPIPNIPEIQKTLPARLDPHFLNNKEMSDVTFLVEGKLFYAHKVLLVTASNRFKTLMTNKTELDSQGSKTVEISDMKYNIFKMLMQYLYYGGTESMEIPTTDILELLSAASLFQLDGLQRHCEILCAQTISMESSVNIYKYAKIHSAPELASFCKGFFLKHMNSLLEQDSFRQLIYGRNSKVQGLDPLRDLQTTLASRLHSIYITSRV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZV4</accession>
    <name>A0A452GZV4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">ATP binding cassette subfamily C member 3</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>17beta-estradiol 17-O-(beta-D-glucuronate)(in) + ATP + H2O = 17beta-estradiol 17-O-(beta-D-glucuronate)(out) + ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:60128"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:82961"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="15">
            <dbReference type="Rhea" id="RHEA:60129"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>ATP + H2O + xenobioticSide 1 = ADP + phosphate + xenobioticSide 2.</text>
            <dbReference type="EC" id="7.6.2.2"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="14">
            <text>leukotriene C4(in) + ATP + H2O = leukotriene C4(out) + ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:38963"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:57973"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="14">
            <dbReference type="Rhea" id="RHEA:38964"/>
        </physiologicalReaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the ABC transporter superfamily. ABCC family. Conjugate transporter (TC 3.A.1.208) subfamily.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000009012.1">
        <property type="protein sequence ID" value="ENSGAGP00000007816.1"/>
        <property type="gene ID" value="ENSGAGG00000006224.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008559">
        <property type="term" value="F:ABC-type xenobiotic transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006869">
        <property type="term" value="P:lipid transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd18595">
        <property type="entry name" value="ABC_6TM_MRP1_2_3_6_D1_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18603">
        <property type="entry name" value="ABC_6TM_MRP1_2_3_6_D2_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd03250">
        <property type="entry name" value="ABCC_MRP_domain1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd03244">
        <property type="entry name" value="ABCC_MRP_domain2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000293">
        <property type="entry name" value="ATP binding cassette subfamily C member 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1560.10:FF:000001">
        <property type="entry name" value="ATP-binding cassette subfamily C member 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1560.10:FF:000007">
        <property type="entry name" value="ATP-binding cassette subfamily C member 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000074">
        <property type="entry name" value="Multidrug resistance-associated protein 5 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1560.10">
        <property type="entry name" value="ABC transporter type 1, transmembrane domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003593">
        <property type="entry name" value="AAA+_ATPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011527">
        <property type="entry name" value="ABC1_TM_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036640">
        <property type="entry name" value="ABC1_TM_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003439">
        <property type="entry name" value="ABC_transporter-like_ATP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017871">
        <property type="entry name" value="ABC_transporter-like_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050173">
        <property type="entry name" value="ABC_transporter_C-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005292">
        <property type="entry name" value="MRP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00957">
        <property type="entry name" value="MRP_assoc_pro"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24223">
        <property type="entry name" value="ATP-BINDING CASSETTE SUB-FAMILY C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24223:SF405">
        <property type="entry name" value="ATP-BINDING CASSETTE SUB-FAMILY C MEMBER 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00664">
        <property type="entry name" value="ABC_membrane"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00005">
        <property type="entry name" value="ABC_tran"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00382">
        <property type="entry name" value="AAA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90123">
        <property type="entry name" value="ABC transporter transmembrane region"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50929">
        <property type="entry name" value="ABC_TM1F"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00211">
        <property type="entry name" value="ABC_TRANSPORTER_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50893">
        <property type="entry name" value="ABC_TRANSPORTER_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11" id="KW-0445">Lipid transport</keyword>
    <keyword evidence="12 16" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="9" id="KW-1278">Translocase</keyword>
    <keyword evidence="5 16" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="10 16" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="20"/>
            <end position="37"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="58"/>
            <end position="75"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="241"/>
            <end position="261"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="317"/>
            <end position="339"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="345"/>
            <end position="365"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="423"/>
            <end position="450"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="856"/>
            <end position="883"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="903"/>
            <end position="925"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="973"/>
            <end position="995"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="1001"/>
            <end position="1021"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="1092"/>
            <end position="1108"/>
        </location>
    </feature>
    <feature type="domain" description="ABC transmembrane type-1" evidence="18">
        <location>
            <begin position="206"/>
            <end position="488"/>
        </location>
    </feature>
    <feature type="domain" description="ABC transporter" evidence="17">
        <location>
            <begin position="521"/>
            <end position="745"/>
        </location>
    </feature>
    <feature type="domain" description="ABC transmembrane type-1" evidence="18">
        <location>
            <begin position="863"/>
            <end position="1143"/>
        </location>
    </feature>
    <feature type="domain" description="ABC transporter" evidence="17">
        <location>
            <begin position="1180"/>
            <end position="1383"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009726"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022967"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00034018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00047523"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00047576"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50893"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50929"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007816.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="21">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1383" mass="155071" checksum="0F3520C194AAAF57" modified="2019-05-08" version="1">MVLATVLLQYERLRGVQSSGVLIIFWFLSVLCAVGPFRSKIIAATAQGQIKERFRFTTFFSYFALILIELILSCFKENPPFFSPVNSDPNPCPELNSGFLSKMTFWWFTRMAILGYKRPLEDKDLWSLNKDDTSSIIVQQLHKEWDKQKDEWKQKEDVTYVKTSNHVLNHVGDGPEEAEVLIQGKRQHKEPSFLKALLRTFGPYFLIGSFYKLIQDLLLFVNPQLLSILIGFTKDKDAPSWWGFLIAALMFFSAVLQTIILHQHFRYCFVTGMRLRTGITGMIYRKSLVITNSAKRSSTVGEIVNLMSVDGQRFMDLVIFLDMLWSAPLQICLALYFLWQNLGPSVLAGVAVMILLIPLNAAIAIKTRALQVEQMRYKDSRIKLMNEILSGIKVLKLYAWEPSFQEKVLAIRKNELHVLKKGAYLHCLSTFAWISAPFLVALTTFAVYVSVDENNILDAEKAFVSLSLFNILRFPLNMLPQVISNLAQTSVSLKRIQQFLSHEELDPNCVETKVVTPGYAITVTNGTFSWAKELKPSLKDINLLVPSGSLVAVVGHVGCGKSSLVSALLGEMEKLEGHVAVKGSVAYVPQQAWIQNATLKDNILFGQQHNEQKYQNVLEACALKTDLEVLPGGDHTEIGEKGINLSGGQRQRVSIARAVFSDADVYLLDDPLSAVDSHVAKHIFDNVIGPEGALKGKTRMLVTHGITFLPQVDHIVVLVDGKISEMGSYQELLKQNGAFAEFLRNYAPDEDVEEDELTMLEDEEVLLAEDTLSNHTDLADNEPVTNEVRKQFLRQISVISSEGECPKKMSARRRVCEKNSAEPPQPQKIPTEKLIQAETSETGTVKLTVFWQYMKAVGLVTSLVMCFLHGCQNAAAIGANVWLSDWTNEPVINGTQRNTSMRVGVYAALGLLQGLLVLISSFTLAMGGINAARKLHAALLENKFHTPQSFFDTTPTGRIINRFSKDIYVIDEVIPPTILMFLGTFFTSVSTMLVIISSTPLFAVVIIPLAIVYFFVQRFYVATSRQLKRLESVSRSPIYSHFSETITGASVIRAYRRENAFVCLSDLKVDENQKSYFPSIVSNRWLGVRIELIGNFVVLFAALFAVLGKNSLNAGLVGLSVSYALQVTLSLNWMVRMTSDLESNIVAVERVKEYSETETEAPWIIEDKRPPENWPAQGEVEFVDYSVRYRKGLDLVLKGLSLRVKGGEKIGIVGRTGAGKSSMTLCLFRILEAVKGEIKIDGVRISEIGLHDLRSKITIIPQDPVLFSGTLRMNLDPFNKYSEDEIWKALELSHLKKFVSSKPDKLEYECSEGGENLSVGQRQLVCLARALLRKTRILVLDEATAAIDLETDDLIQMTIRTQFEDCTVLTIAHRLNTIMDYTR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZV5</accession>
    <name>A0A452GZV5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">PSI domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZV5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008920.1">
        <property type="protein sequence ID" value="ENSGAGP00000007733.1"/>
        <property type="gene ID" value="ENSGAGG00000006159.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002116">
        <property type="term" value="C:semaphorin receptor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017154">
        <property type="term" value="F:semaphorin receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007162">
        <property type="term" value="P:negative regulation of cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050772">
        <property type="term" value="P:positive regulation of axonogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030334">
        <property type="term" value="P:regulation of cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008360">
        <property type="term" value="P:regulation of cell shape"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00102">
        <property type="entry name" value="IPT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="CDD" id="cd12789">
        <property type="entry name" value="RasGAP_plexin_C1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.506.10">
        <property type="entry name" value="GTPase Activation - p120gap, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.20.90">
        <property type="entry name" value="Phosphatidylinositol 3-kinase Catalytic Subunit, Chain A, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002909">
        <property type="entry name" value="IPT_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031148">
        <property type="entry name" value="Plexin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013548">
        <property type="entry name" value="Plexin_cytoplasmic_RasGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046800">
        <property type="entry name" value="Plexin_RBD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002165">
        <property type="entry name" value="Plexin_repeat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016201">
        <property type="entry name" value="PSI"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22625">
        <property type="entry name" value="PLEXIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22625:SF4">
        <property type="entry name" value="PLEXIN-C1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08337">
        <property type="entry name" value="Plexin_cytopl"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20170">
        <property type="entry name" value="Plexin_RBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01437">
        <property type="entry name" value="PSI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01833">
        <property type="entry name" value="TIG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00423">
        <property type="entry name" value="PSI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103575">
        <property type="entry name" value="Plexin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="2 4" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="4">
        <location>
            <begin position="469"/>
            <end position="491"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="4">
        <location>
            <begin position="598"/>
            <end position="619"/>
        </location>
    </feature>
    <feature type="domain" description="PSI" evidence="5">
        <location>
            <begin position="129"/>
            <end position="166"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004370"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="SMART" id="SM00423"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007733.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1066" mass="121826" checksum="8CFA5A5573BEF5A0" modified="2019-05-08" version="1">SMSTALRVDSGTPFGLMEFRNRCKRAQGSIHRVQTRHDISIPEQSILTRRCRGLVWTWAHKHPLINPKSPREKQLLAILNNNMEPNCPEILYEIEEESSVFPKLVLDPVDSDYIYLSFANEVRRIQVANCNKYGSCKECLSAKDPHCGWCYLSKSCSRCISTGVCAFCEIKHISPFVACKPDDSGQPICNISIEPLWISVLGKRDVIIKGANFTEASNITVVLTGTSSCKQDNIQVSKVLNDTHVRFSLPPRRKEAKSVCIKANGRKCSPPITVYYVSEPSCTKMEPNITWASGGRKITLFGRNFNVTDSVIISDDERLKFTVSLELLCNNKIYVSLSKGCKIVNVSLKVENVLIPCIKLQYCPDPIFIDYQLHTEMDPHLELKLYKKNDILDISENEIGVTLTHMMNDIRLEPIIFSVQNITKTPDHTTILCKVKRKKPGKIELSTVEVWVTLGNFSQEVQKKSSHKYLYVLTLLPILLLGVIVVAIIVTRYKSKQLTRKLSQQIELLECEIRKKIRDGFAELQMDQSDVVDSFGTVPFLDYKHFALRTFFPEVNGFCLIFDESLTMLHALICNKDFLVTLIHTLEKQKNFSVKDRCLFASFLTIALQSKLVYLTHILEVLTRDLMEQSSNTQPKLMLRRTESVVEKLLTNWMSVCLSGFLRETVGEPFYKLVTALNQRINKGPVDVTTCKALYTLNEDWLLWQVTEFKSVALNVFFEKIPEDECADTCLDIEANVLDCDTIEQAKEKIFQAFLNKNGSLYGLQLREIGLELQSGAQQKELLDIDGSSVILEDGITKLNTIVHALSPFSYCLDVEYSNKHCHLILPDSQEPQGAMHKGKQKFEVKEMYLTKLLSTKVAIHLVVENLFKSIWSLPNNKAPIAIKYFFDFLDAQAESKKITDPDVVHIWKTNSLPLRFWVNILKNPQFVFDIKKTPHIDGCLSVIAQAFMDAFSLTEQQLGKDAPTNKLLYAKDIPRYQEEVKAYYKAIRDLPLLPSEAEEFLTQESKKHENELNEDVALTEIYKYIAKYYDEIQSKLEREQGLEDAQKDLLHIKALFDEKKKCKWM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZW2</accession>
    <name>A0A452GZW2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Hyaluronidase</fullName>
            <ecNumber evidence="12">3.2.1.35</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1 12">
            <text>Random hydrolysis of (1-&gt;4)-linkages between N-acetyl-beta-D-glucosamine and D-glucuronate residues in hyaluronate.</text>
            <dbReference type="EC" id="3.2.1.35"/>
        </reaction>
    </comment>
    <comment type="subunit">
        <text evidence="4">Monomer.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 9 12">Belongs to the glycosyl hydrolase 56 family.</text>
    </comment>
    <dbReference type="EC" id="3.2.1.35" evidence="12"/>
    <dbReference type="AlphaFoldDB" id="A0A452GZW2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008927.1">
        <property type="protein sequence ID" value="ENSGAGP00000007739.1"/>
        <property type="gene ID" value="ENSGAGG00000006182.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031410">
        <property type="term" value="C:cytoplasmic vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004415">
        <property type="term" value="F:hyalurononglucosaminidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005975">
        <property type="term" value="P:carbohydrate metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030214">
        <property type="term" value="P:hyaluronan catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="entry name" value="Hyaluronidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.20.20.70">
        <property type="entry name" value="Aldolase class I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013785">
        <property type="entry name" value="Aldolase_TIM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017853">
        <property type="entry name" value="GH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018155">
        <property type="entry name" value="Hyaluronidase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11769">
        <property type="entry name" value="HYALURONIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11769:SF9">
        <property type="entry name" value="HYALURONIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01630">
        <property type="entry name" value="Glyco_hydro_56"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF038193">
        <property type="entry name" value="Hyaluronidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00846">
        <property type="entry name" value="GLHYDRLASE56"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51445">
        <property type="entry name" value="(Trans)glycosidases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7 11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="8 12" id="KW-0326">Glycosidase</keyword>
    <keyword evidence="6 12" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0964">Secreted</keyword>
    <feature type="active site" description="Proton donor" evidence="10">
        <location>
            <position position="136"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="11">
        <location>
            <begin position="212"/>
            <end position="228"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00008871"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00011245"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023295"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PIRNR" id="PIRNR038193"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PIRSR" id="PIRSR038193-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PIRSR" id="PIRSR038193-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="RuleBase" id="RU610713"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007739.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="349" mass="40895" checksum="18AA5A9E01EC5E75" modified="2019-05-08" version="1">ISNILGPIWLQQHLLLIMVDGQLLKQAKAPLLLHKPFIVVWNAPTEQCRLRYKVDLDLNVFDIASNTNETLSGSNVTIFYHTHLGHYPYYLDNGVPINGGVPQNESLIKHLNKAKSDIDHSIPLKKFQGLAVIDWENWRPQWDRNWGNKTIYRNKSLELVRRRHPQWSEDKIRKVAKEEFENAGKSFMNNTILLAEHMRPNGLWGYYLYPDCYNYDYKEHPKVYTGRCPDIESSRNDLLLWLWKESTALYPSIYLDYILKSSPNALKFVHYRVKEAIRIASVARKDYVLPVFVYSRPFYAYTLHVLTEVNYIFGSMSSMSGVWAGRIFIDRAPERAPMIRKTMFMALMF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZW6</accession>
    <name>A0A452GZW6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">AB hydrolase-1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GZW6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008885.1">
        <property type="protein sequence ID" value="ENSGAGP00000007700.1"/>
        <property type="gene ID" value="ENSGAGG00000006122.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005811">
        <property type="term" value="C:lipid droplet"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="F:lysophosphatidic acid acyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="F:phosphatidylcholine lysophospholipase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055088">
        <property type="term" value="P:lipid homeostasis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006654">
        <property type="term" value="P:phosphatidic acid biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="entry name" value="alpha/beta hydrolase"/>
        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
    <dbReference type="InterPro" id="IPR029058">
        <property type="entry name" value="AB_hydrolase_fold"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR42886:SF21">
        <property type="entry name" value="(LYSO)-N-ACYLPHOSPHATIDYLETHANOLAMINE LIPASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR42886">
        <property type="entry name" value="RE40534P-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Abhydrolase_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00111">
        <property type="entry name" value="ABHYDROLASE"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53474">
        <property type="entry name" value="alpha/beta-Hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="AB hydrolase-1" evidence="1">
        <location>
            <begin position="32"/>
            <end position="265"/>
        </location>
    </feature>
    <evidence type="ECO:0000259" key="1">
        <source>
            <dbReference type="Pfam" id="PF00561"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007700.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="282" mass="31547" checksum="B151A1724548DA2A" modified="2019-05-08" version="1">WSQFVGRYVTLPGRARIWTVSLCPEQGEGRTPLVLVHGFGGGVGLWVLNLDRLGARRPLHAFDLLGFGRSSRPPFPHDAQGAEAEFVRSIEDWRREMGITTMILLGHSLGGFLAASYSLTYPERVKHLILVDPWGFPVRPTDPSQIRPPPTWVKAMAAVLGRSNPLAVLRVAGPWGPGLVQRFRPDFKQKFAEFFEDDTISEYIYHCNAQAPRIHLLPRELPITLIYGAESWIDTSTGRRVRELRPHSYVRHLAIAGASHHVYADQPAAFNAAVQEICDSVD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZW8</accession>
    <name>A0A452GZW8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">Receptor-type tyrosine-protein phosphatase F</fullName>
            <ecNumber evidence="3">3.1.3.48</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="25">
            <text>O-phospho-L-tyrosyl-[protein] + H2O = L-tyrosyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:10684"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="EC" id="3.1.3.48"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the protein-tyrosine phosphatase family. Receptor class 2A subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.48" evidence="3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008932.1">
        <property type="protein sequence ID" value="ENSGAGP00000007744.1"/>
        <property type="gene ID" value="ENSGAGG00000005093.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="CDD" id="cd05738">
        <property type="entry name" value="IgI_2_RPTP_IIa_LAR_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05739">
        <property type="entry name" value="IgI_3_RPTP_IIa_LAR_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000010">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000027">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000036">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000066">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000144">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000015">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000082">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000128">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000098">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase F isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000353">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase F isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013098">
        <property type="entry name" value="Ig_I-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050713">
        <property type="entry name" value="RTP_Phos/Ushers"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46957">
        <property type="entry name" value="CYTOKINE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46957:SF9">
        <property type="entry name" value="PROTEIN-TYROSINE-PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07679">
        <property type="entry name" value="I-set"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00014">
        <property type="entry name" value="FNTYPEIII"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
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        <property type="entry name" value="IG"/>
        <property type="match status" value="2"/>
    </dbReference>
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        <property type="entry name" value="IGc2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="2"/>
    </dbReference>
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        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="13" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="15" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="4" id="KW-0358">Heparin-binding</keyword>
    <keyword evidence="8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="16" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="12 20" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="14" id="KW-0675">Receptor</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="5 20" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 20" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="1201"/>
            <end position="1224"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="21">
        <location>
            <begin position="88"/>
            <end position="177"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="21">
        <location>
            <begin position="185"/>
            <end position="267"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="274"/>
            <end position="365"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="370"/>
            <end position="464"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="468"/>
            <end position="557"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="564"/>
            <end position="661"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="666"/>
            <end position="765"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="766"/>
            <end position="860"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="864"/>
            <end position="956"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="960"/>
            <end position="1044"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="646"/>
            <end position="667"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1359"/>
            <end position="1387"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004167"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010504"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00013064"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022674"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
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    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
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    </evidence>
    <evidence type="ECO:0000256" key="9">
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            <dbReference type="ARBA" id="ARBA00022889"/>
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    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
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    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
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    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
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    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
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    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
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    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00044158"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00051722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007744.1"/>
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    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1424" mass="157021" checksum="D77E0A7561B3C8D7" modified="2019-05-08" version="1">MQNTSLFLHKDKHLLLEFNNKRSSLHISENHGQAIDSNRVTFSSCPCCCXVPRQLIQTELHLAAAPVAAMICCMLFPVAEDHLPAGFPTIDMGPQLKVVEKARTATMLCAASGNPDPEISWFKDFLPVDTATSNGRIKQLRSGALQIENSEESDQGKYECVATNSAGTRYSAPANLYVRVRRVAPRFSIPPSNHEVMPGGSVNLTCVAVGAPMPYVKWMAGVEELTKEDEMPVGRNVLELNNIVQSASYTCVAMSSLGMIEATAQITVKALPKPPTEPLVTETTATSVTLTWDSGNSDPISYYVIQYKPKSWEEGLFQEVDGVATTRYSIGGLSPFSEYEFRIIAVNNIGRGPPSELVEARTGEQAPSSPPLKVQARMLSASTMLVQWEQPEEPNGQIRGYRVYYTTDPHLPLSMWQKHNTDDSHLTTVGSLITGTTYSIRVLAFTSVGDGPPSDIIQVKTQQGVPAQPADFQAEAESDTRILLTWLPALQERITKYELLYWEGEDGSQRKVNFDPSSSYAVEGLKPDTLYKFQLGARSELGLGVYTPVLEARTAQSSKSPSAPPQEVECVSTSSTTIRVSWVPPPAQSRNGVITQYSIAYQAVDAEDGAKHVVDGIGRERSSWEIKDLEKWTEYKVWVRAHTDVGPGPESSPVHVRTDEDVPSAPPRKVEVESLNSTAIRVSWKLPLSNKQHGQIRGYQVTYVKLENNEPRGQPVIKDVMLAEAQETIIGGLLPETTYSITVAAYTTKGDGARSKPKVVTTTGAVPGKPSMLISTTAMNTALIQWHPPKEMVGELLGYRLQYKRTDEEKMNIMDFEKNVHHYTVTSLHKGATYIFKLSAKNRAGPGEEFEKEITTTEDVPSGFPQNLRVVGLTTSTTEVAWDPPVLAERNGRITNYTVMYRDINSQQELTNITRETSIVLTNLRPDTTYDIKVRACNSKGVGPLSPSIQSRTMPVEQVFAKNFRVNAVMKTSVLLSWEVPDSYKSAVPFKILYNSQSVEVDGHSMKKLISDLQPDTEYSFVLMNRGSSAGGLQHRVSIHTAPDVLQSKPISTNKYMQEGKFTLTLPKVQTSAPVRWYYIVVVPVDQSTSSRTARWRTPDEMELDQLLEAISQGRRQRRQADRLKPYIAALVDMLPETFTLGDEKTYKGFYNKPLSEDLSYRCFVLASLEDGDTKRYAASPYSDEIVMEVSSAKQQDEPEMLWVMGPVLAVILIIIIVIAILLFKSTKGPPSCHQTQRSLSGSGPTGMVEGMRSSIQGMRDHPPIPVTDLADNIDRLKANDGLRFSQEYEVSCREGDRCAWHCPEPGVAGRRWAMAASPRGGSVDSFHAHSFSVHLPRGYPLSWLVPQLQPPAASLVTRRSCPQAPRPGLQRPGVPHXGSPYSCSADTPQSHLASPCPPTHTLALLIPLSPALPLPGLPLVLLC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZW9</accession>
    <name>A0A452GZW9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">Receptor-type tyrosine-protein phosphatase F</fullName>
            <ecNumber evidence="3">3.1.3.48</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="25">
            <text>O-phospho-L-tyrosyl-[protein] + H2O = L-tyrosyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:10684"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="EC" id="3.1.3.48"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the protein-tyrosine phosphatase family. Receptor class 2A subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.48" evidence="3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008937.1">
        <property type="protein sequence ID" value="ENSGAGP00000007749.1"/>
        <property type="gene ID" value="ENSGAGG00000005093.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="CDD" id="cd05738">
        <property type="entry name" value="IgI_2_RPTP_IIa_LAR_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05739">
        <property type="entry name" value="IgI_3_RPTP_IIa_LAR_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000010">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000027">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000036">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000066">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000144">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000015">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000082">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000128">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000098">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase F isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000353">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase F isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013098">
        <property type="entry name" value="Ig_I-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050713">
        <property type="entry name" value="RTP_Phos/Ushers"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46957">
        <property type="entry name" value="CYTOKINE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46957:SF9">
        <property type="entry name" value="PROTEIN-TYROSINE-PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07679">
        <property type="entry name" value="I-set"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00014">
        <property type="entry name" value="FNTYPEIII"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="13" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="15" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="4" id="KW-0358">Heparin-binding</keyword>
    <keyword evidence="8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="16" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="12 20" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="14" id="KW-0675">Receptor</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="5 20" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 20" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="1147"/>
            <end position="1170"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="21">
        <location>
            <begin position="21"/>
            <end position="116"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="21">
        <location>
            <begin position="124"/>
            <end position="206"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="213"/>
            <end position="304"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="309"/>
            <end position="403"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="407"/>
            <end position="496"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="501"/>
            <end position="598"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="603"/>
            <end position="711"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="712"/>
            <end position="806"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="810"/>
            <end position="902"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="906"/>
            <end position="990"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="583"/>
            <end position="604"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1312"/>
            <end position="1340"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004167"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010504"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00013064"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022674"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00044158"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00051722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007749.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1377" mass="152021" checksum="D891CFAB2DC9D89E" modified="2019-05-08" version="1">SGGVCCMLFPVAEDHLPAGFPTIDMGPQLKVVEKARTATMLCAASGNPDPEISWFKDFLPVDTATSNGRIKQLRSGGTPIRGALQIENSEESDQGKYECVATNSAGTRYSAPANLYVRVRRVAPRFSIPPSNHEVMPGGSVNLTCVAVGAPMPYVKWMAGVEELTKEDEMPVGRNVLELNNIVQSASYTCVAMSSLGMIEATAQITVKALPKPPTEPLVTETTATSVTLTWDSGNSDPISYYVIQYKPKSWEEGLFQEVDGVATTRYSIGGLSPFSEYEFRIIAVNNIGRGPPSELVEARTGEQAPSSPPLKVQARMLSASTMLVQWEQPEEPNGQIRGYRVYYTTDPHLPLSMWQKHNTDDSHLTTVGSLITGTTYSIRVLAFTSVGDGPPSDIIQVKTQQGVPAQPADFQAEAESDTRILLTWLPALQERITKYELLYWEGEDGSQRKVNFDPSSSYAVEGLKPDTLYKFQLGARSELGLGVYTPVLEARTAQSTPSAPPQEVECVSTSSTTIRVSWVPPPAQSRNGVITQYSIAYQAVDAEDGAKHVVDGIGRERSSWEIKDLEKWTEYKVWVRAHTDVGPGPESSPVHVRTDEDVPSAPPRKVEVESLNSTAIRVSWKLPLSNKQHGQIRGYQVTYVKLENNEPRGQPVIKDVMLAEAQWRAEDSIDHETIIGGLLPETTYSITVAAYTTKGDGARSKPKVVTTTGAVPGKPSMLISTTAMNTALIQWHPPKEMVGELLGYRLQYKRTDEEKMNIMDFEKNVHHYTVTSLHKGATYIFKLSAKNRAGPGEEFEKEITTTEDVPSGFPQNLRVVGLTTSTTEVAWDPPVLAERNGRITNYTVMYRDINSQQELTNITRETSIVLTNLRPDTTYDIKVRACNSKGVGPLSPSIQSRTMPVEQVFAKNFRVNAVMKTSVLLSWEVPDSYKSAVPFKILYNSQSVEVDGHSMKKLISDLQPDTEYSFVLMNRGSSAGGLQHRVSIHTAPDVLQSKPISTNKYMQEGKFTLTLPKVQTSAPVRWYYIVVVPVDQSTSSRTARWRTPDEMELDQLLEAISQGRRQRRQADRLKPYIAALVDMLPETFTLGDEKTYKGFYNKPLSEDLSYRCFVLASLEDGDTKRYAASPYSDEIVMEVSSAKQQDEPEMLWVMGPVLAVILIIIIVIAILLFKRKRANSPSSKDEHSIGLKDSLLAHSSDPVEMRRLNYQTPGMRDHPPIPVTDLADNIDRLKANDGLRFSQEYEVSCREGDRCAWHCPEPGVAGRRWAMAASPRGGSVDSFHAHSFSVHLPRGYPLSWLVPQLQPPAASLVTRRSCPQAPRPGLQRPGVPHXGSPYSCSADTPQSHLASPCPPTHTLALLIPLSPALPLPGLPLVLLC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZZ0</accession>
    <name>A0A452GZZ0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Large ribosomal subunit protein mL38</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="9">39S ribosomal protein L38, mitochondrial</fullName>
        </alternativeName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Mitochondrion</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="7">Belongs to the phosphatidylethanolamine-binding protein family. Mitochondrion-specific ribosomal protein mL38 subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZZ0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008958.1">
        <property type="protein sequence ID" value="ENSGAGP00000007769.1"/>
        <property type="gene ID" value="ENSGAGG00000006174.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005743">
        <property type="term" value="C:mitochondrial inner membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005762">
        <property type="term" value="C:mitochondrial large ribosomal subunit"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00866">
        <property type="entry name" value="PEBP_euk"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.280.10:FF:000002">
        <property type="entry name" value="39S ribosomal protein L38, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.280.10">
        <property type="entry name" value="PEBP-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008914">
        <property type="entry name" value="PEBP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036610">
        <property type="entry name" value="PEBP-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035810">
        <property type="entry name" value="PEBP_euk"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11362:SF133">
        <property type="entry name" value="LARGE RIBOSOMAL SUBUNIT PROTEIN ML38"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11362">
        <property type="entry name" value="PHOSPHATIDYLETHANOLAMINE-BINDING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01161">
        <property type="entry name" value="PBP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49777">
        <property type="entry name" value="PEBP-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0687">Ribonucleoprotein</keyword>
    <keyword evidence="3" id="KW-0689">Ribosomal protein</keyword>
    <keyword evidence="2" id="KW-0809">Transit peptide</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004173"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022980"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023274"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00038016"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00039444"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00041206"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007769.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="369" mass="43497" checksum="82D6D004349E6106" modified="2019-05-08" version="1">MKPLNTWSSCCCAAALCKRAAPLGPMPNEDIDVSNLETLEKYRSFTRYFKVAEKESKKTPWWQTYRKHVTPEQEAKIDIGLPYYRPSRAKQLKERKEILKQNRQSMEMERASRLKTLVIPLDEVKAEWEKTNGLLHKQRVAEHYGIYRDMFDEATFVPRVVLRIEYNLDDEHVMPVYHGNFVTPTEASSSPEVIYEADEGSLWTLLLTNLDGHLRDADSEYIHWLVTNIPGNNVDSGTEICHYFPPFPAKGTGYHRFVFLLFKQDRPIDFTEDVRPTPCHSLKMRTFKTFDFYKKHQNDMTPAGLAFFQCQWDDSVTHVFHNHLNMKEPVFEFVRPPVYHPPQKKFPHLQPLRYLDRYRDSHEPTYGIY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZZ1</accession>
    <name>A0A452GZZ1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">G protein-coupled receptor 137B</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Lysosome membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZZ1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008959.1">
        <property type="protein sequence ID" value="ENSGAGP00000007770.1"/>
        <property type="gene ID" value="ENSGAGG00000006196.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005765">
        <property type="term" value="C:lysosomal membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045779">
        <property type="term" value="P:negative regulation of bone resorption"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045671">
        <property type="term" value="P:negative regulation of osteoclast differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1904263">
        <property type="term" value="P:positive regulation of TORC1 signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010506">
        <property type="term" value="P:regulation of autophagy"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029723">
        <property type="entry name" value="GPR137"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15146">
        <property type="entry name" value="INTEGRAL MEMBRANE PROTEIN GPR137"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15146:SF0">
        <property type="entry name" value="INTEGRAL MEMBRANE PROTEIN GPR137B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0458">Lysosome</keyword>
    <keyword evidence="4 6" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 6" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="3 6" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="33"/>
            <end position="53"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="65"/>
            <end position="85"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="97"/>
            <end position="121"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="141"/>
            <end position="165"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="223"/>
            <end position="248"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004155"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023228"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007770.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="9">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="335" mass="38925" checksum="069DA4794D9E7300" modified="2019-05-08" version="1">MEASEWDPVKNDTLPPTLTPAVPPYVKLGLTTVYTIFYSLLFVFIYVQLWLVLHYRHKRFSYQTVFLFLCLFWASLRTVLFSFYFKDFVTANSLRPFIFWLLYCFPVCLQFFTLTLMNLYFTQVIFKAKSKYSPELLKYRLPLYLASLFISLLFLLVNLTCAVLVRTENSERKVIVSVRVAINDTLFVLCAISLSICLYKISKMSLANIYLESKADLKCQLGDAGYVVFGVILFIWELLPTSLVVYFFRVRNPTKDLTNSGMVPSHGFSPRSYFFDNPRRYDSDDDLAWNISPQGIQGSFSPDYYDWGHQHNSFMAYIGSLQPESTMDTERPSPI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZZ9</accession>
    <name>A0A452GZZ9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">PSI domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZZ9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008968.1">
        <property type="protein sequence ID" value="ENSGAGP00000007779.1"/>
        <property type="gene ID" value="ENSGAGG00000006159.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002116">
        <property type="term" value="C:semaphorin receptor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017154">
        <property type="term" value="F:semaphorin receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007162">
        <property type="term" value="P:negative regulation of cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050772">
        <property type="term" value="P:positive regulation of axonogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030334">
        <property type="term" value="P:regulation of cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008360">
        <property type="term" value="P:regulation of cell shape"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00102">
        <property type="entry name" value="IPT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="CDD" id="cd12789">
        <property type="entry name" value="RasGAP_plexin_C1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.506.10">
        <property type="entry name" value="GTPase Activation - p120gap, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.20.90">
        <property type="entry name" value="Phosphatidylinositol 3-kinase Catalytic Subunit, Chain A, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002909">
        <property type="entry name" value="IPT_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031148">
        <property type="entry name" value="Plexin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013548">
        <property type="entry name" value="Plexin_cytoplasmic_RasGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046800">
        <property type="entry name" value="Plexin_RBD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002165">
        <property type="entry name" value="Plexin_repeat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016201">
        <property type="entry name" value="PSI"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22625">
        <property type="entry name" value="PLEXIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22625:SF4">
        <property type="entry name" value="PLEXIN-C1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08337">
        <property type="entry name" value="Plexin_cytopl"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20170">
        <property type="entry name" value="Plexin_RBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01437">
        <property type="entry name" value="PSI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01833">
        <property type="entry name" value="TIG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00423">
        <property type="entry name" value="PSI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103575">
        <property type="entry name" value="Plexin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="2 4" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="4">
        <location>
            <begin position="488"/>
            <end position="510"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="4">
        <location>
            <begin position="617"/>
            <end position="638"/>
        </location>
    </feature>
    <feature type="domain" description="PSI" evidence="5">
        <location>
            <begin position="131"/>
            <end position="180"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004370"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="SMART" id="SM00423"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007779.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1085" mass="123414" checksum="F5185217543CF5AC" modified="2019-05-08" version="1">MRLLLPAPTTAWPRLCSEPWPHSQLQPWPWPWPLAPSPAAALARALTPLSLSVPSFSQEPRLSPSHVSWAWTGKERPNAILNNNMEPNCPEILYEIEEESSVFPKLVLDPVDSDYIYLSFANEVRRIQVANCNKYGSCKECLSAKDPHCGWCYLSKRYCLIVVTAVGDVSAVSKGHFVCIVKDTKTNQVLCKKEEPQIMNCSCNLNNAPICNISIEPLWISVLGKRDVIIKGANFTEASNITVVLTGTSSCKQDNIQVSKVLNDTHVRFSLPPRRKEAKSVCIKANGRKCSPPITVYYVSEPSCTKMEPNITWASGGRKITLFGRNFNVTDSVIISDDERLKFTVSGCPGSTSPCSFLTPDVSLSKGCKIVNVSLKVENVLIPCIKLQYCPDPIFIDYQLHTEMDPHLELKLYKKNDILDISENEIGVTLTHMMNDIRLEPIIFSVQNITKTPDHTTILCKVKRKKPGKIELSTVEVWVIQKKSSHKYLYVLTLLPILLLGVIVVAIIVTRYKSKQLTRKLSQQIELLECEIRKKIRDGFAELQMDQSDVVDSFGTVPFLDYKHFALRTFFPEVNGFCLIFDESLTMLHALICNKDFLVTLIHTLEKQKNFSVKDRCLFASFLTIALQSKLVYLTHILEVLTRDLMEQSSNTQPKLMLRRTESVVEKLLTNWMSVCLSGFLRETVGEPFYKLVTALNQRINKGPVDVTTCKALYTLNEDWLLWQVTEFKSVALNVFFEKIPEDECADTCLDIEANVLDCDTIEQAKEKIFQAFLNKNGSLYGLQLREIGLELQSGAQQKELLDIDGSSVILEDGITKLNTIVHALSPFSYCLDVEYSNKHCHLILPDSQEPQGAMHKGKQKFEVKEMYLTKLLSTKVAIHLVVENLFKSIWSLPNNKAPIAIKYFFDFLDAQAESKKITDPDVVHIWKTNSLPLRFWVNILKNPQFVFDIKKTPHIDGCLSVIAQAFMDAFSLTEQQLGKDAPTNKLLYAKDIPRYQEEVKAYYKAIRDLPLLPSEAEEFLTQESKKHENELNEDVALTEIYKYIAKYYDEIQSKLEREQGLEDAQKDLLHIKALFDEKKKCKWM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLU3</accession>
    <name>A0A452HLU3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="30">Serine/threonine-protein kinase MARK2</fullName>
            <ecNumber evidence="8">2.7.11.1</ecNumber>
            <ecNumber evidence="7">2.7.11.26</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="31">ELKL motif kinase 1</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="32">MAP/microtubule affinity-regulating kinase 2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="38 39">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="39" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="38" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="38" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="28">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="27">
            <text>L-seryl-[tau protein] + ATP = O-phospho-L-seryl-[tau protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:12801"/>
            <dbReference type="Rhea" id="RHEA-COMP:13701"/>
            <dbReference type="Rhea" id="RHEA-COMP:13702"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.26"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="26">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="29">
            <text>L-threonyl-[tau protein] + ATP = O-phospho-L-threonyl-[tau protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:53904"/>
            <dbReference type="Rhea" id="RHEA-COMP:13703"/>
            <dbReference type="Rhea" id="RHEA-COMP:13704"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.26"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="5">Cell projection</location>
            <location evidence="5">Dendrite</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cytoplasm</location>
            <location evidence="4">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Endomembrane system</location>
            <topology evidence="3">Peripheral membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Lateral cell membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the protein kinase superfamily. CAMK Ser/Thr protein kinase family. SNF1 subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="8"/>
    <dbReference type="EC" id="2.7.11.26" evidence="7"/>
    <dbReference type="AlphaFoldDB" id="A0A452HLU3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018188.1">
        <property type="protein sequence ID" value="ENSGAGP00000015913.1"/>
        <property type="gene ID" value="ENSGAGG00000011970.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030425">
        <property type="term" value="C:dendrite"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012505">
        <property type="term" value="C:endomembrane system"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016328">
        <property type="term" value="C:lateral plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008289">
        <property type="term" value="F:lipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0106310">
        <property type="term" value="F:protein serine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="RHEA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050321">
        <property type="term" value="F:tau-protein kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035556">
        <property type="term" value="P:intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000226">
        <property type="term" value="P:microtubule cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016055">
        <property type="term" value="P:Wnt signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd14072">
        <property type="entry name" value="STKc_MARK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd14406">
        <property type="entry name" value="UBA_MARK2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:001032">
        <property type="entry name" value="KP78b, isoform A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.8.10:FF:000013">
        <property type="entry name" value="Non-specific serine/threonine protein kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000003">
        <property type="entry name" value="Non-specific serine/threonine protein kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.310.80:FF:000001">
        <property type="entry name" value="Non-specific serine/threonine protein kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.8.10">
        <property type="entry name" value="DNA helicase RuvA subunit, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.310.80">
        <property type="entry name" value="Kinase associated domain 1, KA1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028375">
        <property type="entry name" value="KA1/Ssp2_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001772">
        <property type="entry name" value="KA1_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049508">
        <property type="entry name" value="MARK1-4_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015940">
        <property type="entry name" value="UBA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24346">
        <property type="entry name" value="MAP/MICROTUBULE AFFINITY-REGULATING KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24346:SF56">
        <property type="entry name" value="SERINE_THREONINE-PROTEIN KINASE MARK2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02149">
        <property type="entry name" value="KA1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00627">
        <property type="entry name" value="UBA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00165">
        <property type="entry name" value="UBA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103243">
        <property type="entry name" value="KA1-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50032">
        <property type="entry name" value="KA1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50030">
        <property type="entry name" value="UBA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="20 33" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="10" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="25" id="KW-0966">Cell projection</keyword>
    <keyword evidence="11" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="24" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="9" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="19" id="KW-0221">Differentiation</keyword>
    <keyword evidence="18" id="KW-0418">Kinase</keyword>
    <keyword evidence="22" id="KW-0446">Lipid-binding</keyword>
    <keyword evidence="21" id="KW-0460">Magnesium</keyword>
    <keyword evidence="23" id="KW-0472">Membrane</keyword>
    <keyword evidence="16" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="17 33" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="13" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="39" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="12" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="14" id="KW-0808">Transferase</keyword>
    <keyword evidence="15" id="KW-0879">Wnt signaling pathway</keyword>
    <feature type="domain" description="Protein kinase" evidence="35">
        <location>
            <begin position="54"/>
            <end position="305"/>
        </location>
    </feature>
    <feature type="domain" description="UBA" evidence="36">
        <location>
            <begin position="324"/>
            <end position="363"/>
        </location>
    </feature>
    <feature type="domain" description="KA1" evidence="37">
        <location>
            <begin position="708"/>
            <end position="757"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="34">
        <location>
            <begin position="1"/>
            <end position="50"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="34">
        <location>
            <begin position="382"/>
            <end position="589"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="34">
        <location>
            <begin position="607"/>
            <end position="633"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="34">
        <location>
            <begin position="19"/>
            <end position="46"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="34">
        <location>
            <begin position="417"/>
            <end position="431"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="34">
        <location>
            <begin position="432"/>
            <end position="444"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="34">
        <location>
            <begin position="466"/>
            <end position="485"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="34">
        <location>
            <begin position="493"/>
            <end position="505"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="34">
        <location>
            <begin position="506"/>
            <end position="525"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="34">
        <location>
            <begin position="546"/>
            <end position="560"/>
        </location>
    </feature>
    <feature type="binding site" evidence="33">
        <location>
            <position position="83"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004124"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004184"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004279"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00006234"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00012407"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00012513"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022687"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00023121"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00047899"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="ARBA" id="ARBA00048291"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="ARBA" id="ARBA00048679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="ARBA" id="ARBA00048878"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="ARBA" id="ARBA00070360"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="31">
        <source>
            <dbReference type="ARBA" id="ARBA00079159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="32">
        <source>
            <dbReference type="ARBA" id="ARBA00083985"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="33">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="34">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="35">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="36">
        <source>
            <dbReference type="PROSITE" id="PS50030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="37">
        <source>
            <dbReference type="PROSITE" id="PS50032"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="38">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015913.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="39">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="757" mass="84609" checksum="7EBFBDDBAAA1A6D5" modified="2019-05-08" version="1">MSSSGRSPLPTVNERDTEQPTLGHTESKASSKSNMLRGRNSATSTDEQPHIGNYRLLKTIGKGNFAKVKLARHVLTGKEVAVKIIDKTQLNSSSLQKLFREVRIMKVLNHPNIVKLFEVIETEKTLYLVMEYASGGEVFDYLVAHGRMKEKEARAKFRQIVSAVQYCHQKFIVHRDLKAENLLLDADMNIKIADFGFSNEFTFGNKLDTFCGSPPYAAPELFQGKKYDGPEVDVWSLGVILYTLVSGSLPFDGQNLKELRERVLRGKYRIPFYMSTDCENLLKKFLILNPSKRGTLEQIMKDRWMNVGHEDDELKPYMEPVPDYKDPRRTELMISMGYTREEIQESLIGQKYNEVMATYLLLGYKNSELDNDNITLKPRPAAELANSSAPSPSHKVQRSVSANPKQRRFSDQVPAIPTSTSYSKKTQSNNAENKRPEEEREAGRKASSTVKVPASPLTGLERKKSTPTPSTNSVLSTSTNRSRNSPMLERTGLGQNSIQNGKDSLTTPGSRASTASASAAVGSARPRQHQKSMSASVHPNKPTLPPTENNCEAQRPSTAPQRVPGASPSAHNISSAIPERTNFPRGISSRSTFHAGQLRQVRDQQNLPYNVPPASPSGNSQGRRGASGSIFSKFTSKFVRRPHMVGGDKERDENRDAKPRSLRFTWSMKTTSSMEPNEMMKEIRKVLDANSCQCELQEKYMLLCMHGAPGHDSFVQWEMEVCKLPRLSLNGVRFKRITGTSMAFKNIASKVANELKL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="22" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLU7</accession>
    <name>A0A452HLU7_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="1">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="1 2">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="2" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="1" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="1" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HLU7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018232.1">
        <property type="protein sequence ID" value="ENSGAGP00000015957.1"/>
        <property type="gene ID" value="ENSGAGG00000012004.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000313" key="1">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015957.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="216" mass="23039" checksum="BC51CBF36C2ECFDB" modified="2019-05-08" version="1">MTFFRWDARARARLLVVIKDPVALVRRAQFSRQIPANCLLLWKFQLNAVCFVTSGPKLPCNLDVRLLNCCCVPPQHAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSTVSPGPSGSRWLLSCGLIAG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLU9</accession>
    <name>A0A452HLU9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="30">Serine/threonine-protein kinase MARK2</fullName>
            <ecNumber evidence="8">2.7.11.1</ecNumber>
            <ecNumber evidence="7">2.7.11.26</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="31">ELKL motif kinase 1</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="32">MAP/microtubule affinity-regulating kinase 2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="38 39">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="39" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="38" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="38" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="28">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="27">
            <text>L-seryl-[tau protein] + ATP = O-phospho-L-seryl-[tau protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:12801"/>
            <dbReference type="Rhea" id="RHEA-COMP:13701"/>
            <dbReference type="Rhea" id="RHEA-COMP:13702"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.26"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="26">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="29">
            <text>L-threonyl-[tau protein] + ATP = O-phospho-L-threonyl-[tau protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:53904"/>
            <dbReference type="Rhea" id="RHEA-COMP:13703"/>
            <dbReference type="Rhea" id="RHEA-COMP:13704"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.26"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="5">Cell projection</location>
            <location evidence="5">Dendrite</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cytoplasm</location>
            <location evidence="4">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Endomembrane system</location>
            <topology evidence="3">Peripheral membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Lateral cell membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the protein kinase superfamily. CAMK Ser/Thr protein kinase family. SNF1 subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="8"/>
    <dbReference type="EC" id="2.7.11.26" evidence="7"/>
    <dbReference type="AlphaFoldDB" id="A0A452HLU9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018183.1">
        <property type="protein sequence ID" value="ENSGAGP00000015910.1"/>
        <property type="gene ID" value="ENSGAGG00000011970.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030425">
        <property type="term" value="C:dendrite"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012505">
        <property type="term" value="C:endomembrane system"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016328">
        <property type="term" value="C:lateral plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008289">
        <property type="term" value="F:lipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0106310">
        <property type="term" value="F:protein serine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="RHEA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050321">
        <property type="term" value="F:tau-protein kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035556">
        <property type="term" value="P:intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000226">
        <property type="term" value="P:microtubule cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016055">
        <property type="term" value="P:Wnt signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd14072">
        <property type="entry name" value="STKc_MARK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd14406">
        <property type="entry name" value="UBA_MARK2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:001032">
        <property type="entry name" value="KP78b, isoform A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.8.10:FF:000013">
        <property type="entry name" value="Non-specific serine/threonine protein kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000003">
        <property type="entry name" value="Non-specific serine/threonine protein kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.310.80:FF:000001">
        <property type="entry name" value="Non-specific serine/threonine protein kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.8.10">
        <property type="entry name" value="DNA helicase RuvA subunit, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.310.80">
        <property type="entry name" value="Kinase associated domain 1, KA1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028375">
        <property type="entry name" value="KA1/Ssp2_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001772">
        <property type="entry name" value="KA1_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049508">
        <property type="entry name" value="MARK1-4_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015940">
        <property type="entry name" value="UBA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24346">
        <property type="entry name" value="MAP/MICROTUBULE AFFINITY-REGULATING KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24346:SF56">
        <property type="entry name" value="SERINE_THREONINE-PROTEIN KINASE MARK2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02149">
        <property type="entry name" value="KA1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00627">
        <property type="entry name" value="UBA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00165">
        <property type="entry name" value="UBA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103243">
        <property type="entry name" value="KA1-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50032">
        <property type="entry name" value="KA1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50030">
        <property type="entry name" value="UBA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="20 33" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="10" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="25" id="KW-0966">Cell projection</keyword>
    <keyword evidence="11" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="24" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="9" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="19" id="KW-0221">Differentiation</keyword>
    <keyword evidence="18" id="KW-0418">Kinase</keyword>
    <keyword evidence="22" id="KW-0446">Lipid-binding</keyword>
    <keyword evidence="21" id="KW-0460">Magnesium</keyword>
    <keyword evidence="23" id="KW-0472">Membrane</keyword>
    <keyword evidence="16" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="17 33" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="13" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="39" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="12" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="14" id="KW-0808">Transferase</keyword>
    <keyword evidence="15" id="KW-0879">Wnt signaling pathway</keyword>
    <feature type="domain" description="Protein kinase" evidence="35">
        <location>
            <begin position="54"/>
            <end position="305"/>
        </location>
    </feature>
    <feature type="domain" description="UBA" evidence="36">
        <location>
            <begin position="324"/>
            <end position="363"/>
        </location>
    </feature>
    <feature type="domain" description="KA1" evidence="37">
        <location>
            <begin position="717"/>
            <end position="766"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="34">
        <location>
            <begin position="1"/>
            <end position="50"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="34">
        <location>
            <begin position="382"/>
            <end position="589"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="34">
        <location>
            <begin position="607"/>
            <end position="628"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="34">
        <location>
            <begin position="19"/>
            <end position="46"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="34">
        <location>
            <begin position="417"/>
            <end position="431"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="34">
        <location>
            <begin position="432"/>
            <end position="444"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="34">
        <location>
            <begin position="466"/>
            <end position="485"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="34">
        <location>
            <begin position="493"/>
            <end position="505"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="34">
        <location>
            <begin position="506"/>
            <end position="525"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="34">
        <location>
            <begin position="546"/>
            <end position="560"/>
        </location>
    </feature>
    <feature type="binding site" evidence="33">
        <location>
            <position position="83"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004124"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004184"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004279"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00006234"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00012407"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00012513"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022687"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00023121"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00047899"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="ARBA" id="ARBA00048291"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="ARBA" id="ARBA00048679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="ARBA" id="ARBA00048878"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="ARBA" id="ARBA00070360"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="31">
        <source>
            <dbReference type="ARBA" id="ARBA00079159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="32">
        <source>
            <dbReference type="ARBA" id="ARBA00083985"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="33">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="34">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="35">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="36">
        <source>
            <dbReference type="PROSITE" id="PS50030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="37">
        <source>
            <dbReference type="PROSITE" id="PS50032"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="38">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015910.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="39">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="766" mass="85743" checksum="10BC86D2152456AC" modified="2019-05-08" version="1">MSSSGRSPLPTVNERDTEQPTLGHTESKASSKSNMLRGRNSATSTDEQPHIGNYRLLKTIGKGNFAKVKLARHVLTGKEVAVKIIDKTQLNSSSLQKLFREVRIMKVLNHPNIVKLFEVIETEKTLYLVMEYASGGEVFDYLVAHGRMKEKEARAKFRQIVSAVQYCHQKFIVHRDLKAENLLLDADMNIKIADFGFSNEFTFGNKLDTFCGSPPYAAPELFQGKKYDGPEVDVWSLGVILYTLVSGSLPFDGQNLKELRERVLRGKYRIPFYMSTDCENLLKKFLILNPSKRGTLEQIMKDRWMNVGHEDDELKPYMEPVPDYKDPRRTELMISMGYTREEIQESLIGQKYNEVMATYLLLGYKNSELDNDNITLKPRPAAELANSSAPSPSHKVQRSVSANPKQRRFSDQVPAIPTSTSYSKKTQSNNAENKRPEEEREAGRKASSTVKVPASPLTGLERKKSTPTPSTNSVLSTSTNRSRNSPMLERTGLGQNSIQNGKDSLTTPGSRASTASASAAVGSARPRQHQKSMSASVHPNKPTLPPTENNCEAQRPSTAPQRVPGASPSAHNISSAIPERTNFPRGISSRSTFHAGQLRQVRDQQNLPYNVPPASPSGNSQGRRGASGSIFSKFTSKFVRRNVSFRFARRPHMVGGDKERDENRDAKPRSLRFTWSMKTTSSMEPNEMMKEIRKVLDANSCQCELQEKYMLLCMHGAPGHDSFVQWEMEVCKLPRLSLNGVRFKRITGTSMAFKNIASKVANELKL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLV0</accession>
    <name>A0A452HLV0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="30">Serine/threonine-protein kinase MARK2</fullName>
            <ecNumber evidence="8">2.7.11.1</ecNumber>
            <ecNumber evidence="7">2.7.11.26</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="31">ELKL motif kinase 1</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="32">MAP/microtubule affinity-regulating kinase 2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="38 39">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="39" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="38" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="38" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="28">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="27">
            <text>L-seryl-[tau protein] + ATP = O-phospho-L-seryl-[tau protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:12801"/>
            <dbReference type="Rhea" id="RHEA-COMP:13701"/>
            <dbReference type="Rhea" id="RHEA-COMP:13702"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.26"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="26">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="29">
            <text>L-threonyl-[tau protein] + ATP = O-phospho-L-threonyl-[tau protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:53904"/>
            <dbReference type="Rhea" id="RHEA-COMP:13703"/>
            <dbReference type="Rhea" id="RHEA-COMP:13704"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.26"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="5">Cell projection</location>
            <location evidence="5">Dendrite</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cytoplasm</location>
            <location evidence="4">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Endomembrane system</location>
            <topology evidence="3">Peripheral membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Lateral cell membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the protein kinase superfamily. CAMK Ser/Thr protein kinase family. SNF1 subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="8"/>
    <dbReference type="EC" id="2.7.11.26" evidence="7"/>
    <dbReference type="AlphaFoldDB" id="A0A452HLV0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018193.1">
        <property type="protein sequence ID" value="ENSGAGP00000015918.1"/>
        <property type="gene ID" value="ENSGAGG00000011970.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030425">
        <property type="term" value="C:dendrite"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012505">
        <property type="term" value="C:endomembrane system"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016328">
        <property type="term" value="C:lateral plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008289">
        <property type="term" value="F:lipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0106310">
        <property type="term" value="F:protein serine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="RHEA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050321">
        <property type="term" value="F:tau-protein kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035556">
        <property type="term" value="P:intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000226">
        <property type="term" value="P:microtubule cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016055">
        <property type="term" value="P:Wnt signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd14072">
        <property type="entry name" value="STKc_MARK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd14406">
        <property type="entry name" value="UBA_MARK2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:001032">
        <property type="entry name" value="KP78b, isoform A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.8.10:FF:000013">
        <property type="entry name" value="Non-specific serine/threonine protein kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000003">
        <property type="entry name" value="Non-specific serine/threonine protein kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.310.80:FF:000001">
        <property type="entry name" value="Non-specific serine/threonine protein kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.8.10">
        <property type="entry name" value="DNA helicase RuvA subunit, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.310.80">
        <property type="entry name" value="Kinase associated domain 1, KA1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028375">
        <property type="entry name" value="KA1/Ssp2_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001772">
        <property type="entry name" value="KA1_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049508">
        <property type="entry name" value="MARK1-4_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015940">
        <property type="entry name" value="UBA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24346">
        <property type="entry name" value="MAP/MICROTUBULE AFFINITY-REGULATING KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24346:SF56">
        <property type="entry name" value="SERINE_THREONINE-PROTEIN KINASE MARK2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02149">
        <property type="entry name" value="KA1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00627">
        <property type="entry name" value="UBA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00165">
        <property type="entry name" value="UBA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103243">
        <property type="entry name" value="KA1-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50032">
        <property type="entry name" value="KA1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50030">
        <property type="entry name" value="UBA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="20 33" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="10" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="25" id="KW-0966">Cell projection</keyword>
    <keyword evidence="11" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="24" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="9" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="19" id="KW-0221">Differentiation</keyword>
    <keyword evidence="18" id="KW-0418">Kinase</keyword>
    <keyword evidence="22" id="KW-0446">Lipid-binding</keyword>
    <keyword evidence="21" id="KW-0460">Magnesium</keyword>
    <keyword evidence="23" id="KW-0472">Membrane</keyword>
    <keyword evidence="16" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="17 33" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="13" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="39" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="12" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="14" id="KW-0808">Transferase</keyword>
    <keyword evidence="15" id="KW-0879">Wnt signaling pathway</keyword>
    <feature type="domain" description="Protein kinase" evidence="35">
        <location>
            <begin position="47"/>
            <end position="298"/>
        </location>
    </feature>
    <feature type="domain" description="UBA" evidence="36">
        <location>
            <begin position="317"/>
            <end position="356"/>
        </location>
    </feature>
    <feature type="domain" description="KA1" evidence="37">
        <location>
            <begin position="725"/>
            <end position="774"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="34">
        <location>
            <begin position="16"/>
            <end position="41"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="34">
        <location>
            <begin position="375"/>
            <end position="582"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="34">
        <location>
            <begin position="600"/>
            <end position="622"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="34">
        <location>
            <begin position="642"/>
            <end position="674"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="34">
        <location>
            <begin position="21"/>
            <end position="39"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="34">
        <location>
            <begin position="410"/>
            <end position="424"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="34">
        <location>
            <begin position="425"/>
            <end position="437"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="34">
        <location>
            <begin position="459"/>
            <end position="478"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="34">
        <location>
            <begin position="486"/>
            <end position="498"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="34">
        <location>
            <begin position="499"/>
            <end position="518"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="34">
        <location>
            <begin position="539"/>
            <end position="553"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="34">
        <location>
            <begin position="644"/>
            <end position="674"/>
        </location>
    </feature>
    <feature type="binding site" evidence="33">
        <location>
            <position position="76"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004124"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004184"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004279"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00006234"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00012407"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00012513"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022687"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00023121"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00047899"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="ARBA" id="ARBA00048291"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="ARBA" id="ARBA00048679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="ARBA" id="ARBA00048878"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="ARBA" id="ARBA00070360"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="31">
        <source>
            <dbReference type="ARBA" id="ARBA00079159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="32">
        <source>
            <dbReference type="ARBA" id="ARBA00083985"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="33">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="34">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="35">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="36">
        <source>
            <dbReference type="PROSITE" id="PS50030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="37">
        <source>
            <dbReference type="PROSITE" id="PS50032"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="38">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015918.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="39">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="774" mass="86902" checksum="E1711577A2E98E6F" modified="2019-05-08" version="1">MTAWIWKKINGLPTLGHTESKASSKSNMLRGRNSATSTDEQPHIGNYRLLKTIGKGNFAKVKLARHVLTGKEVAVKIIDKTQLNSSSLQKLFREVRIMKVLNHPNIVKLFEVIETEKTLYLVMEYASGGEVFDYLVAHGRMKEKEARAKFRQIVSAVQYCHQKFIVHRDLKAENLLLDADMNIKIADFGFSNEFTFGNKLDTFCGSPPYAAPELFQGKKYDGPEVDVWSLGVILYTLVSGSLPFDGQNLKELRERVLRGKYRIPFYMSTDCENLLKKFLILNPSKRGTLEQIMKDRWMNVGHEDDELKPYMEPVPDYKDPRRTELMISMGYTREEIQESLIGQKYNEVMATYLLLGYKNSELDNDNITLKPRPAAELANSSAPSPSHKVQRSVSANPKQRRFSDQVPAIPTSTSYSKKTQSNNAENKRPEEEREAGRKASSTVKVPASPLTGLERKKSTPTPSTNSVLSTSTNRSRNSPMLERTGLGQNSIQNGKDSLTTPGSRASTASASAAVGSARPRQHQKSMSASVHPNKPTLPPTENNCEAQRPSTAPQRVPGASPSAHNISSAIPERTNFPRGISSRSTFHAGQLRQVRDQQNLPYNVPPASPSGNSQGRRGASGSIFSKFTSKFVRRNVSFRFARRNPHEPESKDRVETLRPHMVGGDKERDENRDAKPRSLRFTWSMKTTSSMEPNEMMKEIRKVLDANSCQCELQEKYMLLCMHGAPGHDSFVQWEMEVCKLPRLSLNGVRFKRITGTSMAFKNIASKVANELKL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLV1</accession>
    <name>A0A452HLV1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="30">Serine/threonine-protein kinase MARK2</fullName>
            <ecNumber evidence="8">2.7.11.1</ecNumber>
            <ecNumber evidence="7">2.7.11.26</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="31">ELKL motif kinase 1</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="32">MAP/microtubule affinity-regulating kinase 2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="38 39">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="39" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="38" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="38" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="28">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="27">
            <text>L-seryl-[tau protein] + ATP = O-phospho-L-seryl-[tau protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:12801"/>
            <dbReference type="Rhea" id="RHEA-COMP:13701"/>
            <dbReference type="Rhea" id="RHEA-COMP:13702"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.26"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="26">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="29">
            <text>L-threonyl-[tau protein] + ATP = O-phospho-L-threonyl-[tau protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:53904"/>
            <dbReference type="Rhea" id="RHEA-COMP:13703"/>
            <dbReference type="Rhea" id="RHEA-COMP:13704"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.26"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="5">Cell projection</location>
            <location evidence="5">Dendrite</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cytoplasm</location>
            <location evidence="4">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Endomembrane system</location>
            <topology evidence="3">Peripheral membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Lateral cell membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the protein kinase superfamily. CAMK Ser/Thr protein kinase family. SNF1 subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="8"/>
    <dbReference type="EC" id="2.7.11.26" evidence="7"/>
    <dbReference type="AlphaFoldDB" id="A0A452HLV1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018191.1">
        <property type="protein sequence ID" value="ENSGAGP00000015916.1"/>
        <property type="gene ID" value="ENSGAGG00000011970.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030425">
        <property type="term" value="C:dendrite"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012505">
        <property type="term" value="C:endomembrane system"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016328">
        <property type="term" value="C:lateral plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008289">
        <property type="term" value="F:lipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0106310">
        <property type="term" value="F:protein serine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="RHEA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050321">
        <property type="term" value="F:tau-protein kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035556">
        <property type="term" value="P:intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000226">
        <property type="term" value="P:microtubule cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016055">
        <property type="term" value="P:Wnt signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd12201">
        <property type="entry name" value="MARK2_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd14072">
        <property type="entry name" value="STKc_MARK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd14406">
        <property type="entry name" value="UBA_MARK2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:001032">
        <property type="entry name" value="KP78b, isoform A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.8.10:FF:000013">
        <property type="entry name" value="Non-specific serine/threonine protein kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000003">
        <property type="entry name" value="Non-specific serine/threonine protein kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.310.80:FF:000001">
        <property type="entry name" value="Non-specific serine/threonine protein kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.8.10">
        <property type="entry name" value="DNA helicase RuvA subunit, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.310.80">
        <property type="entry name" value="Kinase associated domain 1, KA1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028375">
        <property type="entry name" value="KA1/Ssp2_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001772">
        <property type="entry name" value="KA1_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049508">
        <property type="entry name" value="MARK1-4_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015940">
        <property type="entry name" value="UBA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24346">
        <property type="entry name" value="MAP/MICROTUBULE AFFINITY-REGULATING KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24346:SF56">
        <property type="entry name" value="SERINE_THREONINE-PROTEIN KINASE MARK2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02149">
        <property type="entry name" value="KA1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00627">
        <property type="entry name" value="UBA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00165">
        <property type="entry name" value="UBA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103243">
        <property type="entry name" value="KA1-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50032">
        <property type="entry name" value="KA1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50030">
        <property type="entry name" value="UBA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="20 33" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="10" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="25" id="KW-0966">Cell projection</keyword>
    <keyword evidence="11" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="24" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="9" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="19" id="KW-0221">Differentiation</keyword>
    <keyword evidence="18" id="KW-0418">Kinase</keyword>
    <keyword evidence="22" id="KW-0446">Lipid-binding</keyword>
    <keyword evidence="21" id="KW-0460">Magnesium</keyword>
    <keyword evidence="23" id="KW-0472">Membrane</keyword>
    <keyword evidence="16" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="17 33" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="13" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="39" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="12" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="14" id="KW-0808">Transferase</keyword>
    <keyword evidence="15" id="KW-0879">Wnt signaling pathway</keyword>
    <feature type="domain" description="Protein kinase" evidence="35">
        <location>
            <begin position="54"/>
            <end position="305"/>
        </location>
    </feature>
    <feature type="domain" description="UBA" evidence="36">
        <location>
            <begin position="324"/>
            <end position="363"/>
        </location>
    </feature>
    <feature type="domain" description="KA1" evidence="37">
        <location>
            <begin position="679"/>
            <end position="728"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="34">
        <location>
            <begin position="1"/>
            <end position="50"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="34">
        <location>
            <begin position="382"/>
            <end position="536"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="34">
        <location>
            <begin position="554"/>
            <end position="576"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="34">
        <location>
            <begin position="597"/>
            <end position="628"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="34">
        <location>
            <begin position="19"/>
            <end position="46"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="34">
        <location>
            <begin position="417"/>
            <end position="431"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="34">
        <location>
            <begin position="432"/>
            <end position="444"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="34">
        <location>
            <begin position="466"/>
            <end position="485"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="34">
        <location>
            <begin position="493"/>
            <end position="507"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="34">
        <location>
            <begin position="598"/>
            <end position="628"/>
        </location>
    </feature>
    <feature type="binding site" evidence="33">
        <location>
            <position position="83"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004124"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004184"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004279"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00006234"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00012407"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00012513"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022687"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00023121"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00047899"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="ARBA" id="ARBA00048291"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="ARBA" id="ARBA00048679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="ARBA" id="ARBA00048878"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="ARBA" id="ARBA00070360"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="31">
        <source>
            <dbReference type="ARBA" id="ARBA00079159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="32">
        <source>
            <dbReference type="ARBA" id="ARBA00083985"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="33">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="34">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="35">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="36">
        <source>
            <dbReference type="PROSITE" id="PS50030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="37">
        <source>
            <dbReference type="PROSITE" id="PS50032"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="38">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015916.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="39">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="728" mass="82081" checksum="AF173DF8FAE84424" modified="2019-05-08" version="1">MSSSGRSPLPTVNERDTEQPTLGHTESKASSKSNMLRGRNSATSTDEQPHIGNYRLLKTIGKGNFAKVKLARHVLTGKEVAVKIIDKTQLNSSSLQKLFREVRIMKVLNHPNIVKLFEVIETEKTLYLVMEYASGGEVFDYLVAHGRMKEKEARAKFRQIVSAVQYCHQKFIVHRDLKAENLLLDADMNIKIADFGFSNEFTFGNKLDTFCGSPPYAAPELFQGKKYDGPEVDVWSLGVILYTLVSGSLPFDGQNLKELRERVLRGKYRIPFYMSTDCENLLKKFLILNPSKRGTLEQIMKDRWMNVGHEDDELKPYMEPVPDYKDPRRTELMISMGYTREEIQESLIGQKYNEVMATYLLLGYKNSELDNDNITLKPRPAAELANSSAPSPSHKVQRSVSANPKQRRFSDQVPAIPTSTSYSKKTQSNNAENKRPEEEREAGRKASSTVKVPASPLTGLERKKSTPTPSTNSVLSTSTNRSRNSPMLERTGLGQNSIQNGKDSTAPQRVPGASPSAHNISSAIPERTNFPRGISSRSTFHAGQLRQVRDQQNLPYNVPPASPSGNSQGRRGASGSIFSKFTSKFVRRNVSFRFARRNPHEPESKDRVETLRPHMVGGDKERDENRDAKPRSLRFTWSMKTTSSMEPNEMMKEIRKVLDANSCQCELQEKYMLLCMHGAPGHDSFVQWEMEVCKLPRLSLNGVRFKRITGTSMAFKNIASKVANELKL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLV8</accession>
    <name>A0A452HLV8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="30">Serine/threonine-protein kinase MARK2</fullName>
            <ecNumber evidence="8">2.7.11.1</ecNumber>
            <ecNumber evidence="7">2.7.11.26</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="31">ELKL motif kinase 1</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="32">MAP/microtubule affinity-regulating kinase 2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="38 39">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="39" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="38" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="38" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="28">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="27">
            <text>L-seryl-[tau protein] + ATP = O-phospho-L-seryl-[tau protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:12801"/>
            <dbReference type="Rhea" id="RHEA-COMP:13701"/>
            <dbReference type="Rhea" id="RHEA-COMP:13702"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.26"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="26">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="29">
            <text>L-threonyl-[tau protein] + ATP = O-phospho-L-threonyl-[tau protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:53904"/>
            <dbReference type="Rhea" id="RHEA-COMP:13703"/>
            <dbReference type="Rhea" id="RHEA-COMP:13704"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.26"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="5">Cell projection</location>
            <location evidence="5">Dendrite</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cytoplasm</location>
            <location evidence="4">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Endomembrane system</location>
            <topology evidence="3">Peripheral membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Lateral cell membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the protein kinase superfamily. CAMK Ser/Thr protein kinase family. SNF1 subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="8"/>
    <dbReference type="EC" id="2.7.11.26" evidence="7"/>
    <dbReference type="AlphaFoldDB" id="A0A452HLV8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018199.1">
        <property type="protein sequence ID" value="ENSGAGP00000015924.1"/>
        <property type="gene ID" value="ENSGAGG00000011970.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030425">
        <property type="term" value="C:dendrite"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012505">
        <property type="term" value="C:endomembrane system"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016328">
        <property type="term" value="C:lateral plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008289">
        <property type="term" value="F:lipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0106310">
        <property type="term" value="F:protein serine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="RHEA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050321">
        <property type="term" value="F:tau-protein kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035556">
        <property type="term" value="P:intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000226">
        <property type="term" value="P:microtubule cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016055">
        <property type="term" value="P:Wnt signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd14072">
        <property type="entry name" value="STKc_MARK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd14406">
        <property type="entry name" value="UBA_MARK2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:001032">
        <property type="entry name" value="KP78b, isoform A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.8.10:FF:000013">
        <property type="entry name" value="Non-specific serine/threonine protein kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000003">
        <property type="entry name" value="Non-specific serine/threonine protein kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.310.80:FF:000001">
        <property type="entry name" value="Non-specific serine/threonine protein kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.8.10">
        <property type="entry name" value="DNA helicase RuvA subunit, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.310.80">
        <property type="entry name" value="Kinase associated domain 1, KA1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028375">
        <property type="entry name" value="KA1/Ssp2_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001772">
        <property type="entry name" value="KA1_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049508">
        <property type="entry name" value="MARK1-4_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015940">
        <property type="entry name" value="UBA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24346">
        <property type="entry name" value="MAP/MICROTUBULE AFFINITY-REGULATING KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24346:SF56">
        <property type="entry name" value="SERINE_THREONINE-PROTEIN KINASE MARK2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02149">
        <property type="entry name" value="KA1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00627">
        <property type="entry name" value="UBA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00165">
        <property type="entry name" value="UBA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103243">
        <property type="entry name" value="KA1-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50032">
        <property type="entry name" value="KA1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50030">
        <property type="entry name" value="UBA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="20 33" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="10" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="25" id="KW-0966">Cell projection</keyword>
    <keyword evidence="11" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="24" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="9" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="19" id="KW-0221">Differentiation</keyword>
    <keyword evidence="18" id="KW-0418">Kinase</keyword>
    <keyword evidence="22" id="KW-0446">Lipid-binding</keyword>
    <keyword evidence="21" id="KW-0460">Magnesium</keyword>
    <keyword evidence="23" id="KW-0472">Membrane</keyword>
    <keyword evidence="16" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="17 33" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="13" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="39" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="12" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="14" id="KW-0808">Transferase</keyword>
    <keyword evidence="15" id="KW-0879">Wnt signaling pathway</keyword>
    <feature type="domain" description="Protein kinase" evidence="35">
        <location>
            <begin position="48"/>
            <end position="299"/>
        </location>
    </feature>
    <feature type="domain" description="UBA" evidence="36">
        <location>
            <begin position="318"/>
            <end position="357"/>
        </location>
    </feature>
    <feature type="domain" description="KA1" evidence="37">
        <location>
            <begin position="726"/>
            <end position="775"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="34">
        <location>
            <begin position="1"/>
            <end position="42"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="34">
        <location>
            <begin position="376"/>
            <end position="583"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="34">
        <location>
            <begin position="601"/>
            <end position="623"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="34">
        <location>
            <begin position="643"/>
            <end position="675"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="34">
        <location>
            <begin position="22"/>
            <end position="40"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="34">
        <location>
            <begin position="411"/>
            <end position="425"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="34">
        <location>
            <begin position="426"/>
            <end position="438"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="34">
        <location>
            <begin position="460"/>
            <end position="479"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="34">
        <location>
            <begin position="487"/>
            <end position="499"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="34">
        <location>
            <begin position="500"/>
            <end position="519"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="34">
        <location>
            <begin position="540"/>
            <end position="554"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="34">
        <location>
            <begin position="645"/>
            <end position="675"/>
        </location>
    </feature>
    <feature type="binding site" evidence="33">
        <location>
            <position position="77"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004124"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004184"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004279"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00006234"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00012407"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00012513"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022687"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00023121"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00047899"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="ARBA" id="ARBA00048291"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="ARBA" id="ARBA00048679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="ARBA" id="ARBA00048878"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="ARBA" id="ARBA00070360"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00079159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="32">
        <source>
            <dbReference type="ARBA" id="ARBA00083985"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="33">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="34">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="35">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="36">
        <source>
            <dbReference type="PROSITE" id="PS50030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="37">
        <source>
            <dbReference type="PROSITE" id="PS50032"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="38">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015924.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="39">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="775" mass="86856" checksum="D2B1B20F09C5272A" modified="2019-05-08" version="1">MSRGSTYTRPMAGPTLGHTESKASSKSNMLRGRNSATSTDEQPHIGNYRLLKTIGKGNFAKVKLARHVLTGKEVAVKIIDKTQLNSSSLQKLFREVRIMKVLNHPNIVKLFEVIETEKTLYLVMEYASGGEVFDYLVAHGRMKEKEARAKFRQIVSAVQYCHQKFIVHRDLKAENLLLDADMNIKIADFGFSNEFTFGNKLDTFCGSPPYAAPELFQGKKYDGPEVDVWSLGVILYTLVSGSLPFDGQNLKELRERVLRGKYRIPFYMSTDCENLLKKFLILNPSKRGTLEQIMKDRWMNVGHEDDELKPYMEPVPDYKDPRRTELMISMGYTREEIQESLIGQKYNEVMATYLLLGYKNSELDNDNITLKPRPAAELANSSAPSPSHKVQRSVSANPKQRRFSDQVPAIPTSTSYSKKTQSNNAENKRPEEEREAGRKASSTVKVPASPLTGLERKKSTPTPSTNSVLSTSTNRSRNSPMLERTGLGQNSIQNGKDSLTTPGSRASTASASAAVGSARPRQHQKSMSASVHPNKPTLPPTENNCEAQRPSTAPQRVPGASPSAHNISSAIPERTNFPRGISSRSTFHAGQLRQVRDQQNLPYNVPPASPSGNSQGRRGASGSIFSKFTSKFVRRNVSFRFARRNPHEPESKDRVETLRPHMVGGDKERDENRDAKPRSLRFTWSMKTTSSMEPNEMMKEIRKVLDANSCQCELQEKYMLLCMHGAPGHDSFVQWEMEVCKLPRLSLNGVRFKRITGTSMAFKNIASKVANELKL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLV9</accession>
    <name>A0A452HLV9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">EF-hand domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="1">Myosin is a hexamer of 2 heavy chains and 4 light chains.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLV9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018203.1">
        <property type="protein sequence ID" value="ENSGAGP00000015928.1"/>
        <property type="gene ID" value="ENSGAGG00000011990.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000018206.1">
        <property type="protein sequence ID" value="ENSGAGP00000015931.1"/>
        <property type="gene ID" value="ENSGAGG00000011990.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00051">
        <property type="entry name" value="EFh"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.238.10:FF:000010">
        <property type="entry name" value="Myosin regulatory light chain 2, atrial isoform"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.238.10:FF:000007">
        <property type="entry name" value="Putative myosin regulatory light chain sqh"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.238.10">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011992">
        <property type="entry name" value="EF-hand-dom_pair"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018247">
        <property type="entry name" value="EF_Hand_1_Ca_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002048">
        <property type="entry name" value="EF_hand_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050403">
        <property type="entry name" value="Myosin_RLC"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23049">
        <property type="entry name" value="MYOSIN REGULATORY LIGHT CHAIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13499">
        <property type="entry name" value="EF-hand_7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00054">
        <property type="entry name" value="EFh"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47473">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00018">
        <property type="entry name" value="EF_HAND_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50222">
        <property type="entry name" value="EF_HAND_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0106">Calcium</keyword>
    <keyword evidence="2" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="EF-hand" evidence="5">
        <location>
            <begin position="29"/>
            <end position="64"/>
        </location>
    </feature>
    <feature type="domain" description="EF-hand" evidence="5">
        <location>
            <begin position="98"/>
            <end position="133"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00011445"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50222"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015928.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="172" mass="19776" checksum="ED2108C56216B209" modified="2019-05-08" version="1">MSSKRAKTKTAKKRPQRATSNVFAMFDQSQIQEFKEAFNMIDQNRDGFIDKEDLHDMLASLGKDPTDAYLDAMMNEAPGPINFTMFLTMFGEKLNGTDPEDVIRNAFACFDEEATGVIQEDYLRELLTTMGDRFTDEEVDELYREAPIDKKGNFNYIEFTRILKHGAKDKDD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLW1</accession>
    <name>A0A452HLW1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">PNPLA domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="caution">
        <text evidence="2">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLW1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018251.1">
        <property type="protein sequence ID" value="ENSGAGP00000015972.1"/>
        <property type="gene ID" value="ENSGAGG00000012001.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005811">
        <property type="term" value="C:lipid droplet"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004806">
        <property type="term" value="F:triacylglycerol lipase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055088">
        <property type="term" value="P:lipid homeostasis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019433">
        <property type="term" value="P:triglyceride catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.1090.10">
        <property type="entry name" value="Cytosolic phospholipase A2 catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016035">
        <property type="entry name" value="Acyl_Trfase/lysoPLipase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033562">
        <property type="entry name" value="PLPL"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002641">
        <property type="entry name" value="PNPLA_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12406">
        <property type="entry name" value="CALCIUM-INDEPENDENT PHOSPHOLIPASE A2 IPLA2 -RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12406:SF23">
        <property type="entry name" value="OMEGA-HYDROXYCERAMIDE TRANSACYLASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01734">
        <property type="entry name" value="Patatin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52151">
        <property type="entry name" value="FabD/lysophospholipase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51635">
        <property type="entry name" value="PNPLA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="2" id="KW-0442">Lipid degradation</keyword>
    <keyword evidence="1 2" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="PNPLA" evidence="3">
        <location>
            <begin position="5"/>
            <end position="153"/>
        </location>
    </feature>
    <feature type="short sequence motif" description="GXSXG" evidence="2">
        <location>
            <begin position="40"/>
            <end position="44"/>
        </location>
    </feature>
    <feature type="short sequence motif" description="DGA/G" evidence="2">
        <location>
            <begin position="140"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="active site" description="Nucleophile" evidence="2">
        <location>
            <position position="42"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="2">
        <location>
            <position position="140"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01161"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS51635"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015972.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="243" mass="27153" checksum="A192DAF72033E6A0" modified="2019-05-08" version="1">MMRVARTGSSGFLALYQIGVVQSLLELAPEILKSASKVYGASAGSLIAAAVMDFNAMKGSERNDLHNTITNNVIMLPENSHQVASGRLHISLTRVMDGQNVMVSEFSSKEDLIQALICSCFVPIYCGFIPPSFRGVRYVDGGFTNLQPCSDLEAVITVSPFTGELDICPRDCPAIFYSFQIFNSSIQISVENLCRISYALFPPSYLVRYQDAVLFLRRNSKWYSEPVKRQEAFSVWHCFPYRE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLW6</accession>
    <name>A0A452HLW6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Dysferlin</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="17">Dystrophy-associated fer-1-like protein</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="18">Fer-1-like protein 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">Key calcium ion sensor involved in the Ca(2+)-triggered synaptic vesicle-plasma membrane fusion. Plays a role in the sarcolemma repair mechanism of both skeletal muscle and cardiomyocytes that permits rapid resealing of membranes disrupted by mechanical stress.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <location evidence="1">Sarcolemma</location>
            <topology evidence="1">Single-pass type II membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasmic vesicle membrane</location>
            <topology evidence="2">Single-pass type II membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the ferlin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018257.1">
        <property type="protein sequence ID" value="ENSGAGP00000015977.1"/>
        <property type="gene ID" value="ENSGAGG00000011993.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030659">
        <property type="term" value="C:cytoplasmic vesicle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012505">
        <property type="term" value="C:endomembrane system"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015630">
        <property type="term" value="C:microtubule cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030315">
        <property type="term" value="C:T-tubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008289">
        <property type="term" value="F:lipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002281">
        <property type="term" value="P:macrophage activation involved in immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061025">
        <property type="term" value="P:membrane fusion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002280">
        <property type="term" value="P:monocyte activation involved in immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050765">
        <property type="term" value="P:negative regulation of phagocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007009">
        <property type="term" value="P:plasma membrane organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd08373">
        <property type="entry name" value="C2A_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04011">
        <property type="entry name" value="C2B_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04018">
        <property type="entry name" value="C2C_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04017">
        <property type="entry name" value="C2D_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04037">
        <property type="entry name" value="C2E_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd08374">
        <property type="entry name" value="C2F_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000009">
        <property type="entry name" value="dysferlin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000021">
        <property type="entry name" value="dysferlin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000026">
        <property type="entry name" value="dysferlin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000033">
        <property type="entry name" value="dysferlin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000037">
        <property type="entry name" value="dysferlin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000061">
        <property type="entry name" value="dysferlin isoform X8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000008">
        <property type="entry name" value="C2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037726">
        <property type="entry name" value="C2A_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037720">
        <property type="entry name" value="C2B_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037722">
        <property type="entry name" value="C2C_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037723">
        <property type="entry name" value="C2D_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037724">
        <property type="entry name" value="C2E_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037725">
        <property type="entry name" value="C2F_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012968">
        <property type="entry name" value="FerIin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037721">
        <property type="entry name" value="Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012560">
        <property type="entry name" value="Ferlin_A-domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012561">
        <property type="entry name" value="Ferlin_B-domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032362">
        <property type="entry name" value="Ferlin_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055072">
        <property type="entry name" value="Ferlin_DSRM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006614">
        <property type="entry name" value="Peroxin/Ferlin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12546:SF44">
        <property type="entry name" value="DYSFERLIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12546">
        <property type="entry name" value="FER-1-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00168">
        <property type="entry name" value="C2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22901">
        <property type="entry name" value="dsrm_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08165">
        <property type="entry name" value="FerA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08150">
        <property type="entry name" value="FerB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08151">
        <property type="entry name" value="FerI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16165">
        <property type="entry name" value="Ferlin_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00239">
        <property type="entry name" value="C2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SMART" id="SM00694">
        <property type="entry name" value="DysFC"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00693">
        <property type="entry name" value="DysFN"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM01200">
        <property type="entry name" value="FerA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01201">
        <property type="entry name" value="FerB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01202">
        <property type="entry name" value="FerI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49562">
        <property type="entry name" value="C2 domain (Calcium/lipid-binding domain, CaLB)"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50004">
        <property type="entry name" value="C2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0106">Calcium</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="14" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="12" id="KW-0446">Lipid-binding</keyword>
    <keyword evidence="13 20" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="10" id="KW-0735">Signal-anchor</keyword>
    <keyword evidence="6 20" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 20" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="2041"/>
            <end position="2063"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="1"/>
            <end position="101"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="197"/>
            <end position="315"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="354"/>
            <end position="490"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="1130"/>
            <end position="1256"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="1303"/>
            <end position="1431"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="1557"/>
            <end position="1675"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="1791"/>
            <end position="1939"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="127"/>
            <end position="208"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1992"/>
            <end position="2013"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="19">
        <location>
            <begin position="127"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="172"/>
            <end position="182"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="19">
        <location>
            <begin position="188"/>
            <end position="203"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="1994"/>
            <end position="2013"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004274"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007561"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023121"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00060070"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00068801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00078385"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00079079"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015977.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2076" mass="237309" checksum="0A4E1F1DA2E6ADE3" modified="2019-05-08" version="1">MLRVFILHAENVQTPDTDISDSYCSVVFEGVKKRTKVIKNNVNPVWNEGFDWDLKGIPLDQNAELTVVVKDHETVGRNRFLGEARVPLQDVLVNPTLTASFNAPLLDTKRQSTGAFLNLQVSYIPPPGAAPLFPPPTPPEPTPSTAELDTVTDTAGEEETEDQIVTGDETEPSTGPSGSEQPTLPRKPPSHHVHGIKRKKSAPKKLLSNKPQDFQIRVRVIEGRQLPGVNIKSVVKVTVAGQTKRTRIRKGNGPFFDEIFFFNVFESPAELFDEPIFITVVDSRSLRTDSVIGEFQLDVGTVYAEPKHALLRKWLLLSDSEDFSAGAKGYVKVSLFVLGPGDEAPLEKKEVAEEKEDIEGNLLRPTGVTLRGAHFCLKIYRAEDLPQMDDAIMDNVKQIFGFESNKKNLVDPFVEVSFAGRTISSKILEKNANPQWNQRITVPAMFPSMCEKMRIRVMDWDRVTHNDVIGTTHLRLSQISAPGGELEDEFPVLAKPLKAAAMDDGLGFLPTFGPCYVNLYGSPREFTGFPDPYEELNMGKGEGVAFRGRLLMELETKLVEHIEQKLEDIPPDDILRVEKHLRRRKYCLFAAFYSASMLQDVDAAIQFEVSIGNYGNKFDNTCLPLASTTQYSGAVFDGCQYYYLPWGNVKPVVVLSSYWEDISHRTDMQNLLQRTADRLEAGLELVHLALKARCSSNELDSLVSQLIDDIIADCSQPLVDVSNKTASTHLDQFVYKFRSTNLHQIIQAALNLKHEDSDLQTVLDQAEDWLLRLKAMAEEPQNSLPDIVIWMLQGDKRVAYARVPAHEVLFSRTGANCCGKNCGKLQTIFLKYPQEEVMGPRIPAQVRVQLWFGLAVDEKEFNKYAEGKLSVFAETYENQVKLALVGNWGTTGLTYPKYSDVTGKIKLPKDSFRPSVGWTWAGDWFISPEKTLLYDTDAGHMSFVEEVFENQVRLPGGQWIHMADAYTDVNGEKVLPKEEIECPPGWKWEDVEWDTDLNRAVDERGWEYGIAIPPDHKPKAWVPAEKMYHTSRRRRWVRLRRRDLKQMEALKKHKQEELEGEGWEYASLFGWKFHLNYRKTDTFRRRRWRRRMEPLERTGAAAVFALEGALGGVTDDKSDDGKSVSTLSFGVNRPTVSCIFDYGSRYHLRCYIYQARDLIAMDKDSFSDPYAIVSFLHQSQKTVVVKNTLNPLWDQTLIFYEIEIFGDPQNVADVPPNIVVEIYDHDTYGADEFMGRCICKPSLERSPRLSWHPVARGNRKAGELLAAFELIQREKPAVHHLPGFESELSASLDESADSDLPYPPPQREPNIYMVPQGIKPVLQRTAIEILAWGLRNLKSYQLASITSPSLQVECGGQTVQSCVIKNVRKNPNFDVCVLFMEVRLPKEALYCPPVIMKVIDNRQFGRKPVVGQCTIQSLEEFYCDPYKEETDAPEEQPDEVSLTPRDDVLIDIDDKEPLIPVQKIPWEEEFIDWWSKFYASTGEREKCGSYLEKGFDTLKVYETELENMEEFERLSDFCHTFKLYRGRTQDINEDPSVVGEFKGSFKMYPLPDDPRVPVPPRQFHQLPAKGPQECLIRIYIIRAFDLQPKDSNGKCDPYVKISVGKKSINDQENYIPCTLEPVFGKMFELTCTLPLEKDLKVTLYDYDLLSKDEKIGETVIDLENRFLSKFGARCGLPQTYCISGPNQWRDQLRPSQLLHLFSLQRNCKAPIYKPDRVVFREQEYFLSEFEDGKPPNPHLGAPEERLALYTLRKQGLVPEHVETRPLYSPLQPEIEQGKLQMWVDLFPKSLGLPGPPFNITPRRAKRFFLRCIIWNTKDVILDDLSITGEKMSDIYVKGWMTGHEENKQKTDVHYRSMGGEGNFNWRFVFPFDYLAAEQVCHIAKKEHFWSLDKTDNKVPPQLILQIWDNDKFSFDDYLGSVQMNLNRMPAPTKTAEKCSLNLLDETLSSDRFVSLFEQKAVKGWWPCVVEAEEKKTIAGKLEMTLEIVTEQEQEERPAGMGRDEPNMNPKLEDPKRPETSFLWFTSPYKTMKYILWRRFKWVFLIAIVVFILLLFIGIFIYAFPNYAAMKLVKPFS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLW8</accession>
    <name>A0A452HLW8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">SPIN-DOC-like zinc-finger domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HLW8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018213.1">
        <property type="protein sequence ID" value="ENSGAGP00000015938.1"/>
        <property type="gene ID" value="ENSGAGG00000011995.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045892">
        <property type="term" value="P:negative regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040647">
        <property type="entry name" value="SPIN-DOC_Znf-C2H2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052675">
        <property type="entry name" value="ZnF_transloc-Spindlin_int"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR34589">
        <property type="entry name" value="SIMILAR TO RIKEN CDNA 2700081O15"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR34589:SF1">
        <property type="entry name" value="SPINDLIN INTERACTOR AND REPRESSOR OF CHROMATIN-BINDING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18658">
        <property type="entry name" value="zf-C2H2_12"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="SPIN-DOC-like zinc-finger" evidence="2">
        <location>
            <begin position="67"/>
            <end position="129"/>
        </location>
    </feature>
    <feature type="domain" description="SPIN-DOC-like zinc-finger" evidence="2">
        <location>
            <begin position="339"/>
            <end position="399"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="128"/>
            <end position="203"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="221"/>
            <end position="287"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="1">
        <location>
            <begin position="181"/>
            <end position="198"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF18658"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015938.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="416" mass="44577" checksum="E9E83A2F07B8544B" modified="2019-05-08" version="1">MEVKAEVSRLEYIAVSLKCEAEEPVPIAACAQLCAPMGDLAPGVKLEEADKEPLDGGGAPAEAGPGRAWQQEYLVGDCPGRGGTVCMVCGTFLGTCGPSAAREHVLQHHAHSLGLSPEEKHNILEAWSQGGNLPEGAPPPCTPGQYHGAKEPAEIEVLIDPEEQTVTRKRGQPRGRPAPRPGTTSSSEEAAPSSAPPEIRLFTDGSFPAGFTLRLYCRPQSASGGREIRKNPSIQEQKSPGDRGKTQALGKRQKQLGKAGTVRRSAGNPVGNGPISALPSQDLDPGVPRLRRKGCSHHGITRRLRLSPSTVRCTLQSWQTQGKAPAAPSPPQARPFDPAWRADFLMDYDAGAGTMVCMVCEYSLLPIRLGTVMRHIRQCHAETLHLPRGVRRAIREVWETRGPVPTPPQHGALKGE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLX0</accession>
    <name>A0A452HLX0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">SPIN-DOC-like zinc-finger domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HLX0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018211.1">
        <property type="protein sequence ID" value="ENSGAGP00000015936.1"/>
        <property type="gene ID" value="ENSGAGG00000011995.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045892">
        <property type="term" value="P:negative regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040647">
        <property type="entry name" value="SPIN-DOC_Znf-C2H2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052675">
        <property type="entry name" value="ZnF_transloc-Spindlin_int"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR34589">
        <property type="entry name" value="SIMILAR TO RIKEN CDNA 2700081O15"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR34589:SF1">
        <property type="entry name" value="SPINDLIN INTERACTOR AND REPRESSOR OF CHROMATIN-BINDING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18658">
        <property type="entry name" value="zf-C2H2_12"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="SPIN-DOC-like zinc-finger" evidence="2">
        <location>
            <begin position="67"/>
            <end position="129"/>
        </location>
    </feature>
    <feature type="domain" description="SPIN-DOC-like zinc-finger" evidence="2">
        <location>
            <begin position="348"/>
            <end position="408"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="128"/>
            <end position="296"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="211"/>
            <end position="224"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF18658"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015936.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="425" mass="45868" checksum="D22DAF0AA9354C76" modified="2019-05-08" version="1">MEVKAEVSRLEYIAVSLKCEAEEPVPIAACAQLCAPMGDLAPGVKLEEADKEPLDGGGAPAEAGPGRAWQQEYLVGDCPGRGGTVCMVCGTFLGTCGPSAAREHVLQHHAHSLGLSPEEKHNILEAWSQGGNLPEGAPPPCTPGQYHGAKEPAEIEVLIDPEEQTVTRKRGQPRGRPAPRPDARALEPGQRPRGQRRRQSVRPVEGMALPARERRSKQLSEGQDRGTVSPASGGREIRKNPSIQEQKSPGDRGKTQALGKRQKQLGKAGTVRRSAGNPVGNGPISALPSQDLDPGVPRLRRKGCSHHGITRRLRLSPSTVRCTLQSWQTQGKAPAAPSPPQARPFDPAWRADFLMDYDAGAGTMVCMVCEYSLLPIRLGTVMRHIRQCHAETLHLPRGVRRAIREVWETRGPVPTPPQHGALKGE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLX4</accession>
    <name>A0A452HLX4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Zinc finger protein 135</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLX4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018262.1">
        <property type="protein sequence ID" value="ENSGAGP00000015982.1"/>
        <property type="gene ID" value="ENSGAGG00000012025.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd07765">
        <property type="entry name" value="KRAB_A-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001833">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002434">
        <property type="entry name" value="Zinc finger protein 1168"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000638">
        <property type="entry name" value="Zinc finger protein 184"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000295">
        <property type="entry name" value="zinc finger protein 19"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001823">
        <property type="entry name" value="Zinc finger protein 322"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002343">
        <property type="entry name" value="Zinc finger protein 33A"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002090">
        <property type="entry name" value="Zinc finger protein 473"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000737">
        <property type="entry name" value="Zinc finger protein 565"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003655">
        <property type="entry name" value="aKRAB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050752">
        <property type="entry name" value="C2H2-ZF_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001909">
        <property type="entry name" value="KRAB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036051">
        <property type="entry name" value="KRAB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24384">
        <property type="entry name" value="FINGER PUTATIVE TRANSCRIPTION FACTOR FAMILY-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24384:SF218">
        <property type="entry name" value="ZINC FINGER PROTEIN 502"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01352">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="SMART" id="SM00349">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109640">
        <property type="entry name" value="KRAB domain (Kruppel-associated box)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50805">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50806">
        <property type="entry name" value="KRAB_RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="11"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 9" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="KRAB-related" evidence="12">
        <location>
            <begin position="26"/>
            <end position="96"/>
        </location>
    </feature>
    <feature type="domain" description="KRAB" evidence="11">
        <location>
            <begin position="29"/>
            <end position="100"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="10">
        <location>
            <begin position="112"/>
            <end position="139"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="10">
        <location>
            <begin position="140"/>
            <end position="167"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="10">
        <location>
            <begin position="168"/>
            <end position="195"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="10">
        <location>
            <begin position="196"/>
            <end position="223"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="10">
        <location>
            <begin position="224"/>
            <end position="251"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="10">
        <location>
            <begin position="252"/>
            <end position="279"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="10">
        <location>
            <begin position="280"/>
            <end position="307"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="10">
        <location>
            <begin position="308"/>
            <end position="335"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="10">
        <location>
            <begin position="336"/>
            <end position="363"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="10">
        <location>
            <begin position="364"/>
            <end position="391"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="10">
        <location>
            <begin position="392"/>
            <end position="419"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50806"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015982.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="15">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="432" mass="49276" checksum="A939170864D106B8" modified="2019-05-08" version="1">NRPLGTLLTAIHPWLPFNGVGLLLFQGPVTFEEVAVYFTSEEWALLDPTQRALYWDVMQENYENVTSLGFPVSKPHIISQLEQGAEPWVPDLQGSEEILSKHQRIHTGKRPYKCSECGKSFITSSVLSQHQRIHTGERPYECLECGKSFITSSDLSQHQRIHTGERPYKCSECGKSFITSSDLSKHQRIHTGKRPYKCSECGKSFITSSVLSQHQRIHTGERPYECRECGKSFSSSSALSHHQRIHTGERPFECSDCGKTFAFRSVLSKHQRIHTGERPYKCSECGKSFTQSSQLTEHQRIHTGERPFECSDCGKTFTYRSVLSKHQRIHTGERPYKCHECGKSFTRSSHLTGHQRIHTGERPYKCHECGESFTRSSHLSGHQRIHTGERPYKCGKCGKTFIHRQELISHQRIHTGMRPNGSSECGKTLSCH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLX7</accession>
    <name>A0A452HLX7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Dysferlin</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="17">Dystrophy-associated fer-1-like protein</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="18">Fer-1-like protein 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">Key calcium ion sensor involved in the Ca(2+)-triggered synaptic vesicle-plasma membrane fusion. Plays a role in the sarcolemma repair mechanism of both skeletal muscle and cardiomyocytes that permits rapid resealing of membranes disrupted by mechanical stress.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <location evidence="1">Sarcolemma</location>
            <topology evidence="1">Single-pass type II membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasmic vesicle membrane</location>
            <topology evidence="2">Single-pass type II membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the ferlin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018210.1">
        <property type="protein sequence ID" value="ENSGAGP00000015935.1"/>
        <property type="gene ID" value="ENSGAGG00000011993.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030659">
        <property type="term" value="C:cytoplasmic vesicle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012505">
        <property type="term" value="C:endomembrane system"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015630">
        <property type="term" value="C:microtubule cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030315">
        <property type="term" value="C:T-tubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008289">
        <property type="term" value="F:lipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002281">
        <property type="term" value="P:macrophage activation involved in immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061025">
        <property type="term" value="P:membrane fusion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002280">
        <property type="term" value="P:monocyte activation involved in immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050765">
        <property type="term" value="P:negative regulation of phagocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007009">
        <property type="term" value="P:plasma membrane organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd08373">
        <property type="entry name" value="C2A_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04011">
        <property type="entry name" value="C2B_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04018">
        <property type="entry name" value="C2C_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04017">
        <property type="entry name" value="C2D_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04037">
        <property type="entry name" value="C2E_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd08374">
        <property type="entry name" value="C2F_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000009">
        <property type="entry name" value="dysferlin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000021">
        <property type="entry name" value="dysferlin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000026">
        <property type="entry name" value="dysferlin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000033">
        <property type="entry name" value="dysferlin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000037">
        <property type="entry name" value="dysferlin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000061">
        <property type="entry name" value="dysferlin isoform X8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000008">
        <property type="entry name" value="C2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037726">
        <property type="entry name" value="C2A_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037720">
        <property type="entry name" value="C2B_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037722">
        <property type="entry name" value="C2C_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037723">
        <property type="entry name" value="C2D_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037724">
        <property type="entry name" value="C2E_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037725">
        <property type="entry name" value="C2F_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012968">
        <property type="entry name" value="FerIin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037721">
        <property type="entry name" value="Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012560">
        <property type="entry name" value="Ferlin_A-domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012561">
        <property type="entry name" value="Ferlin_B-domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032362">
        <property type="entry name" value="Ferlin_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055072">
        <property type="entry name" value="Ferlin_DSRM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006614">
        <property type="entry name" value="Peroxin/Ferlin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12546:SF44">
        <property type="entry name" value="DYSFERLIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12546">
        <property type="entry name" value="FER-1-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00168">
        <property type="entry name" value="C2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22901">
        <property type="entry name" value="dsrm_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08165">
        <property type="entry name" value="FerA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08150">
        <property type="entry name" value="FerB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08151">
        <property type="entry name" value="FerI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16165">
        <property type="entry name" value="Ferlin_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00239">
        <property type="entry name" value="C2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SMART" id="SM00694">
        <property type="entry name" value="DysFC"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00693">
        <property type="entry name" value="DysFN"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM01200">
        <property type="entry name" value="FerA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01201">
        <property type="entry name" value="FerB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01202">
        <property type="entry name" value="FerI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49562">
        <property type="entry name" value="C2 domain (Calcium/lipid-binding domain, CaLB)"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50004">
        <property type="entry name" value="C2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0106">Calcium</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="14" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="12" id="KW-0446">Lipid-binding</keyword>
    <keyword evidence="13 20" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="10" id="KW-0735">Signal-anchor</keyword>
    <keyword evidence="6 20" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 20" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="2062"/>
            <end position="2084"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="1"/>
            <end position="101"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="197"/>
            <end position="315"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="354"/>
            <end position="490"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="1130"/>
            <end position="1256"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="1303"/>
            <end position="1431"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="1578"/>
            <end position="1696"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="1812"/>
            <end position="1960"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="127"/>
            <end position="208"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="2013"/>
            <end position="2034"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="19">
        <location>
            <begin position="127"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="172"/>
            <end position="182"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="19">
        <location>
            <begin position="188"/>
            <end position="203"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="2015"/>
            <end position="2034"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004274"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007561"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023121"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00060070"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00068801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00078385"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00079079"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015935.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2097" mass="239506" checksum="B0EC9B7185AB9C4F" modified="2019-05-08" version="1">MLRVFILHAENVQTPDTDISDSYCSVVFEGVKKRTKVIKNNVNPVWNEGFDWDLKGIPLDQNAELTVVVKDHETVGRNRFLGEARVPLQDVLVNPTLTASFNAPLLDTKRQSTGAFLNLQVSYIPPPGAAPLFPPPTPPEPTPSTAELDTVTDTAGEEETEDQIVTGDETEPSTGPSGSEQPTLPRKPPSHHVHGIKRKKSAPKKLLSNKPQDFQIRVRVIEGRQLPGVNIKSVVKVTVAGQTKRTRIRKGNGPFFDEIFFFNVFESPAELFDEPIFITVVDSRSLRTDSVIGEFQLDVGTVYAEPKHALLRKWLLLSDSEDFSAGAKGYVKVSLFVLGPGDEAPLEKKEVAEEKEDIEGNLLRPTGVTLRGAHFCLKIYRAEDLPQMDDAIMDNVKQIFGFESNKKNLVDPFVEVSFAGRTISSKILEKNANPQWNQRITVPAMFPSMCEKMRIRVMDWDRVTHNDVIGTTHLRLSQISAPGGELEDEFPVLAKPLKAAAMDDGLGFLPTFGPCYVNLYGSPREFTGFPDPYEELNMGKGEGVAFRGRLLMELETKLVEHIEQKLEDIPPDDILRVEKHLRRRKYCLFAAFYSASMLQDVDAAIQFEVSIGNYGNKFDNTCLPLASTTQYSGAVFDGCQYYYLPWGNVKPVVVLSSYWEDISHRTDMQNLLQRTADRLEAGLELVHLALKARCSSNELDSLVSQLIDDIIADCSQPLVDVSNKTASTHLDQFVYKFRSTNLHQIIQAALNLKHEDSDLQTVLDQAEDWLLRLKAMAEEPQNSLPDIVIWMLQGDKRVAYARVPAHEVLFSRTGANCCGKNCGKLQTIFLKYPQEEVMGPRIPAQVRVQLWFGLAVDEKEFNKYAEGKLSVFAETYENQVKLALVGNWGTTGLTYPKYSDVTGKIKLPKDSFRPSVGWTWAGDWFISPEKTLLYDTDAGHMSFVEEVFENQVRLPGGQWIHMADAYTDVNGEKVLPKEEIECPPGWKWEDVEWDTDLNRAVDERGWEYGIAIPPDHKPKAWVPAEKMYHTSRRRRWVRLRRRDLKQMEALKKHKQEELEGEGWEYASLFGWKFHLNYRKTDTFRRRRWRRRMEPLERTGAAAVFALEGALGGVTDDKSDDGKSVSTLSFGVNRPTVSCIFDYGSRYHLRCYIYQARDLIAMDKDSFSDPYAIVSFLHQSQKTVVVKNTLNPLWDQTLIFYEIEIFGDPQNVADVPPNIVVEIYDHDTYGADEFMGRCICKPSLERSPRLSWHPVARGNRKAGELLAAFELIQREKPAVHHLPGFESELSASLDESADSDLPYPPPQREPNIYMVPQGIKPVLQRTAIEILAWGLRNLKSYQLASITSPSLQVECGGQTVQSCVIKNVRKNPNFDVCVLFMEVRLPKEALYCPPVIMKVIDNRQFGRKPVVGQCTIQSLEEFYCDPYKEETDAPEEQPDEVSLTPRDDVLIDIDDKEPLIPVQLAEGMTSSTLINMPASRADLHKIPWEEEFIDWWSKFYASTGEREKCGSYLEKGFDTLKVYETELENMEEFERLSDFCHTFKLYRGRTQDINEDPSVVGEFKGSFKMYPLPDDPRVPVPPRQFHQLPAKGPQECLIRIYIIRAFDLQPKDSNGKCDPYVKISVGKKSINDQENYIPCTLEPVFGKMFELTCTLPLEKDLKVTLYDYDLLSKDEKIGETVIDLENRFLSKFGARCGLPQTYCISGPNQWRDQLRPSQLLHLFSLQRNCKAPIYKPDRVVFREQEYFLSEFEDGKPPNPHLGAPEERLALYTLRKQGLVPEHVETRPLYSPLQPEIEQGKLQMWVDLFPKSLGLPGPPFNITPRRAKRFFLRCIIWNTKDVILDDLSITGEKMSDIYVKGWMTGHEENKQKTDVHYRSMGGEGNFNWRFVFPFDYLAAEQVCHIAKKEHFWSLDKTDNKVPPQLILQIWDNDKFSFDDYLGSVQMNLNRMPAPTKTAEKCSLNLLDETLSSDRFVSLFEQKAVKGWWPCVVEAEEKKTIAGKLEMTLEIVTEQEQEERPAGMGRDEPNMNPKLEDPKRPETSFLWFTSPYKTMKYILWRRFKWVFLIAIVVFILLLFIGIFIYAFPNYAAMKLVKPFS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLX9</accession>
    <name>A0A452HLX9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Fatty acid synthase</fullName>
            <ecNumber evidence="5">1.1.1.100</ecNumber>
            <ecNumber evidence="2">1.3.1.39</ecNumber>
            <ecNumber evidence="8">2.3.1.38</ecNumber>
            <ecNumber evidence="9">2.3.1.39</ecNumber>
            <ecNumber evidence="7">2.3.1.41</ecNumber>
            <ecNumber evidence="4">2.3.1.85</ecNumber>
            <ecNumber evidence="3">3.1.2.14</ecNumber>
            <ecNumber evidence="6">4.2.1.59</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="73 74">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="74" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="73" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="73" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="36">Fatty acid synthetase is a multifunctional enzyme that catalyzes the de novo biosynthesis of long-chain saturated fatty acids starting from acetyl-CoA and malonyl-CoA in the presence of NADPH. This multifunctional protein contains 7 catalytic activities and a site for the binding of the prosthetic group 4'-phosphopantetheine of the acyl carrier protein ([ACP]) domain.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="43">
            <text>(2E)-butenoyl-[ACP] + NADPH + H(+) = butanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41812"/>
            <dbReference type="Rhea" id="RHEA-COMP:9627"/>
            <dbReference type="Rhea" id="RHEA-COMP:9628"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78453"/>
            <dbReference type="ChEBI" id="CHEBI:78454"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="43">
            <dbReference type="Rhea" id="RHEA:41813"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="67">
            <text>(2E)-decenoyl-[ACP] + NADPH + H(+) = decanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41864"/>
            <dbReference type="Rhea" id="RHEA-COMP:9639"/>
            <dbReference type="Rhea" id="RHEA-COMP:9640"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78467"/>
            <dbReference type="ChEBI" id="CHEBI:78468"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="67">
            <dbReference type="Rhea" id="RHEA:41865"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="50">
            <text>(2E)-dodecenoyl-[ACP] + NADPH + H(+) = dodecanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41880"/>
            <dbReference type="Rhea" id="RHEA-COMP:9643"/>
            <dbReference type="Rhea" id="RHEA-COMP:9644"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:65264"/>
            <dbReference type="ChEBI" id="CHEBI:78472"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="50">
            <dbReference type="Rhea" id="RHEA:41881"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="45">
            <text>(2E)-hexadecenoyl-[ACP] + NADPH + H(+) = hexadecanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41912"/>
            <dbReference type="Rhea" id="RHEA-COMP:9651"/>
            <dbReference type="Rhea" id="RHEA-COMP:9652"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78481"/>
            <dbReference type="ChEBI" id="CHEBI:78483"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="45">
            <dbReference type="Rhea" id="RHEA:41913"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="46">
            <text>(2E)-hexenoyl-[ACP] + NADPH + H(+) = hexanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41832"/>
            <dbReference type="Rhea" id="RHEA-COMP:9631"/>
            <dbReference type="Rhea" id="RHEA-COMP:9632"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78458"/>
            <dbReference type="ChEBI" id="CHEBI:78459"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="46">
            <dbReference type="Rhea" id="RHEA:41833"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="60">
            <text>(2E)-octadecenoyl-[ACP] + NADPH + H(+) = octadecanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41928"/>
            <dbReference type="Rhea" id="RHEA-COMP:9655"/>
            <dbReference type="Rhea" id="RHEA-COMP:9656"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78489"/>
            <dbReference type="ChEBI" id="CHEBI:78495"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="60">
            <dbReference type="Rhea" id="RHEA:41929"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="53">
            <text>(2E)-octenoyl-[ACP] + NADPH + H(+) = octanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41848"/>
            <dbReference type="Rhea" id="RHEA-COMP:9635"/>
            <dbReference type="Rhea" id="RHEA-COMP:9636"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78462"/>
            <dbReference type="ChEBI" id="CHEBI:78463"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="53">
            <dbReference type="Rhea" id="RHEA:41849"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="62">
            <text>(2E)-tetradecenoyl-[ACP] + NADPH + H(+) = tetradecanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41896"/>
            <dbReference type="Rhea" id="RHEA-COMP:9647"/>
            <dbReference type="Rhea" id="RHEA-COMP:9648"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78475"/>
            <dbReference type="ChEBI" id="CHEBI:78477"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="62">
            <dbReference type="Rhea" id="RHEA:41897"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="35">
            <text>(3R)-hydroxybutanoyl-[ACP] = (2E)-butenoyl-[ACP] + H2O</text>
            <dbReference type="Rhea" id="RHEA:41808"/>
            <dbReference type="Rhea" id="RHEA-COMP:9626"/>
            <dbReference type="Rhea" id="RHEA-COMP:9627"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:78451"/>
            <dbReference type="ChEBI" id="CHEBI:78453"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="35">
            <dbReference type="Rhea" id="RHEA:41809"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="30">
            <text>(3R)-hydroxydecanoyl-[ACP] = (2E)-decenoyl-[ACP] + H2O</text>
            <dbReference type="Rhea" id="RHEA:41860"/>
            <dbReference type="Rhea" id="RHEA-COMP:9638"/>
            <dbReference type="Rhea" id="RHEA-COMP:9639"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:78466"/>
            <dbReference type="ChEBI" id="CHEBI:78467"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="30">
            <dbReference type="Rhea" id="RHEA:41861"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="28">
            <text>(3R)-hydroxydodecanoyl-[ACP] = (2E)-dodecenoyl-[ACP] + H2O</text>
            <dbReference type="Rhea" id="RHEA:41876"/>
            <dbReference type="Rhea" id="RHEA-COMP:9642"/>
            <dbReference type="Rhea" id="RHEA-COMP:9643"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:78470"/>
            <dbReference type="ChEBI" id="CHEBI:78472"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="28">
            <dbReference type="Rhea" id="RHEA:41877"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="34">
            <text>(3R)-hydroxyhexadecanoyl-[ACP] = (2E)-hexadecenoyl-[ACP] + H2O</text>
            <dbReference type="Rhea" id="RHEA:41908"/>
            <dbReference type="Rhea" id="RHEA-COMP:9650"/>
            <dbReference type="Rhea" id="RHEA-COMP:9651"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:78480"/>
            <dbReference type="ChEBI" id="CHEBI:78481"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="34">
            <dbReference type="Rhea" id="RHEA:41909"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="29">
            <text>(3R)-hydroxyhexanoyl-[ACP] = (2E)-hexenoyl-[ACP] + H2O</text>
            <dbReference type="Rhea" id="RHEA:41828"/>
            <dbReference type="Rhea" id="RHEA-COMP:9630"/>
            <dbReference type="Rhea" id="RHEA-COMP:9631"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:78457"/>
            <dbReference type="ChEBI" id="CHEBI:78458"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="29">
            <dbReference type="Rhea" id="RHEA:41829"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="33">
            <text>(3R)-hydroxyoctadecanoyl-[ACP] = (2E)-octadecenoyl-[ACP] + H2O</text>
            <dbReference type="Rhea" id="RHEA:41924"/>
            <dbReference type="Rhea" id="RHEA-COMP:9654"/>
            <dbReference type="Rhea" id="RHEA-COMP:9655"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:78488"/>
            <dbReference type="ChEBI" id="CHEBI:78489"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="33">
            <dbReference type="Rhea" id="RHEA:41925"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="27">
            <text>(3R)-hydroxyoctanoyl-[ACP] = (2E)-octenoyl-[ACP] + H2O</text>
            <dbReference type="Rhea" id="RHEA:41844"/>
            <dbReference type="Rhea" id="RHEA-COMP:9634"/>
            <dbReference type="Rhea" id="RHEA-COMP:9635"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:78461"/>
            <dbReference type="ChEBI" id="CHEBI:78462"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="27">
            <dbReference type="Rhea" id="RHEA:41845"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="32">
            <text>(3R)-hydroxytetradecanoyl-[ACP] = (2E)-tetradecenoyl-[ACP] + H2O</text>
            <dbReference type="Rhea" id="RHEA:41892"/>
            <dbReference type="Rhea" id="RHEA-COMP:9646"/>
            <dbReference type="Rhea" id="RHEA-COMP:9647"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:78474"/>
            <dbReference type="ChEBI" id="CHEBI:78475"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="32">
            <dbReference type="Rhea" id="RHEA:41893"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="47">
            <text>3-oxobutanoyl-[ACP] + NADPH + H(+) = (3R)-hydroxybutanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41804"/>
            <dbReference type="Rhea" id="RHEA-COMP:9625"/>
            <dbReference type="Rhea" id="RHEA-COMP:9626"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78450"/>
            <dbReference type="ChEBI" id="CHEBI:78451"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="47">
            <dbReference type="Rhea" id="RHEA:41805"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="41">
            <text>3-oxodecanoyl-[ACP] + NADPH + H(+) = (3R)-hydroxydecanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41856"/>
            <dbReference type="Rhea" id="RHEA-COMP:9637"/>
            <dbReference type="Rhea" id="RHEA-COMP:9638"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78464"/>
            <dbReference type="ChEBI" id="CHEBI:78466"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="41">
            <dbReference type="Rhea" id="RHEA:41857"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="63">
            <text>3-oxododecanoyl-[ACP] + NADPH + H(+) = (3R)-hydroxydodecanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41872"/>
            <dbReference type="Rhea" id="RHEA-COMP:9641"/>
            <dbReference type="Rhea" id="RHEA-COMP:9642"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78469"/>
            <dbReference type="ChEBI" id="CHEBI:78470"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="63">
            <dbReference type="Rhea" id="RHEA:41873"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="64">
            <text>3-oxohexadecanoyl-[ACP] + NADPH + H(+) = (3R)-hydroxyhexadecanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41904"/>
            <dbReference type="Rhea" id="RHEA-COMP:9649"/>
            <dbReference type="Rhea" id="RHEA-COMP:9650"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78478"/>
            <dbReference type="ChEBI" id="CHEBI:78480"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="64">
            <dbReference type="Rhea" id="RHEA:41905"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="55">
            <text>3-oxohexanoyl-[ACP] + NADPH + H(+) = (3R)-hydroxyhexanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41824"/>
            <dbReference type="Rhea" id="RHEA-COMP:9629"/>
            <dbReference type="Rhea" id="RHEA-COMP:9630"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78456"/>
            <dbReference type="ChEBI" id="CHEBI:78457"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="55">
            <dbReference type="Rhea" id="RHEA:41825"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="38">
            <text>3-oxooctadecanoyl-[ACP] + NADPH + H(+) = (3R)-hydroxyoctadecanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41920"/>
            <dbReference type="Rhea" id="RHEA-COMP:9653"/>
            <dbReference type="Rhea" id="RHEA-COMP:9654"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78487"/>
            <dbReference type="ChEBI" id="CHEBI:78488"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="38">
            <dbReference type="Rhea" id="RHEA:41921"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="65">
            <text>3-oxooctanoyl-[ACP] + NADPH + H(+) = (3R)-hydroxyoctanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41840"/>
            <dbReference type="Rhea" id="RHEA-COMP:9633"/>
            <dbReference type="Rhea" id="RHEA-COMP:9634"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78460"/>
            <dbReference type="ChEBI" id="CHEBI:78461"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="65">
            <dbReference type="Rhea" id="RHEA:41841"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="59">
            <text>3-oxotetradecanoyl-[ACP] + NADPH + H(+) = (3R)-hydroxytetradecanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41888"/>
            <dbReference type="Rhea" id="RHEA-COMP:9645"/>
            <dbReference type="Rhea" id="RHEA-COMP:9646"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78473"/>
            <dbReference type="ChEBI" id="CHEBI:78474"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="59">
            <dbReference type="Rhea" id="RHEA:41889"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="40">
            <text>a (3R)-hydroxyacyl-[ACP] + NADP(+) = a 3-oxoacyl-[ACP] + NADPH + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17397"/>
            <dbReference type="Rhea" id="RHEA-COMP:9916"/>
            <dbReference type="Rhea" id="RHEA-COMP:9945"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78776"/>
            <dbReference type="ChEBI" id="CHEBI:78827"/>
            <dbReference type="EC" id="1.1.1.100"/>
        </reaction>
        <physiologicalReaction direction="right-to-left" evidence="40">
            <dbReference type="Rhea" id="RHEA:17399"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="31">
            <text>a (3R)-hydroxyacyl-[ACP] = a (2E)-enoyl-[ACP] + H2O</text>
            <dbReference type="Rhea" id="RHEA:13097"/>
            <dbReference type="Rhea" id="RHEA-COMP:9925"/>
            <dbReference type="Rhea" id="RHEA-COMP:9945"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:78784"/>
            <dbReference type="ChEBI" id="CHEBI:78827"/>
            <dbReference type="EC" id="4.2.1.59"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="31">
            <dbReference type="Rhea" id="RHEA:13098"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="56">
            <text>a 2,3-saturated acyl-[ACP] + NADP(+) = a (2E)-enoyl-[ACP] + NADPH + H(+)</text>
            <dbReference type="Rhea" id="RHEA:22564"/>
            <dbReference type="Rhea" id="RHEA-COMP:9925"/>
            <dbReference type="Rhea" id="RHEA-COMP:9926"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78784"/>
            <dbReference type="ChEBI" id="CHEBI:78785"/>
            <dbReference type="EC" id="1.3.1.39"/>
        </reaction>
        <physiologicalReaction direction="right-to-left" evidence="56">
            <dbReference type="Rhea" id="RHEA:22566"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="54">
            <text>a fatty acyl-[ACP] + malonyl-[ACP] + H(+) = a 3-oxoacyl-[ACP] + holo-[ACP] + CO2</text>
            <dbReference type="Rhea" id="RHEA:22836"/>
            <dbReference type="Rhea" id="RHEA-COMP:9623"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="Rhea" id="RHEA-COMP:9916"/>
            <dbReference type="Rhea" id="RHEA-COMP:14125"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78449"/>
            <dbReference type="ChEBI" id="CHEBI:78776"/>
            <dbReference type="ChEBI" id="CHEBI:138651"/>
            <dbReference type="EC" id="2.3.1.41"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="54">
            <dbReference type="Rhea" id="RHEA:22837"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="37">
            <text>acetyl-CoA + n malonyl-CoA + 2n NADPH + 2n H(+) = a long-chain fatty acid + (n+1) CoA + n CO2 + 2n NADP(+).</text>
            <dbReference type="EC" id="2.3.1.85"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="48">
            <text>acetyl-[ACP] + malonyl-[ACP] + H(+) = 3-oxobutanoyl-[ACP] + holo-[ACP] + CO2</text>
            <dbReference type="Rhea" id="RHEA:41800"/>
            <dbReference type="Rhea" id="RHEA-COMP:9621"/>
            <dbReference type="Rhea" id="RHEA-COMP:9623"/>
            <dbReference type="Rhea" id="RHEA-COMP:9625"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78446"/>
            <dbReference type="ChEBI" id="CHEBI:78449"/>
            <dbReference type="ChEBI" id="CHEBI:78450"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="48">
            <dbReference type="Rhea" id="RHEA:41801"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="66">
            <text>butanoyl-[ACP] + malonyl-[ACP] + H(+) = 3-oxohexanoyl-[ACP] + holo-[ACP] + CO2</text>
            <dbReference type="Rhea" id="RHEA:41820"/>
            <dbReference type="Rhea" id="RHEA-COMP:9623"/>
            <dbReference type="Rhea" id="RHEA-COMP:9628"/>
            <dbReference type="Rhea" id="RHEA-COMP:9629"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78449"/>
            <dbReference type="ChEBI" id="CHEBI:78454"/>
            <dbReference type="ChEBI" id="CHEBI:78456"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="66">
            <dbReference type="Rhea" id="RHEA:41821"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="61">
            <text>decanoyl-[ACP] + malonyl-[ACP] + H(+) = 3-oxododecanoyl-[ACP] + holo-[ACP] + CO2</text>
            <dbReference type="Rhea" id="RHEA:41868"/>
            <dbReference type="Rhea" id="RHEA-COMP:9623"/>
            <dbReference type="Rhea" id="RHEA-COMP:9640"/>
            <dbReference type="Rhea" id="RHEA-COMP:9641"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78449"/>
            <dbReference type="ChEBI" id="CHEBI:78468"/>
            <dbReference type="ChEBI" id="CHEBI:78469"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="61">
            <dbReference type="Rhea" id="RHEA:41869"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="44">
            <text>dodecanoyl-[ACP] + malonyl-[ACP] + H(+) = 3-oxotetradecanoyl-[ACP] + holo-[ACP] + CO2</text>
            <dbReference type="Rhea" id="RHEA:41884"/>
            <dbReference type="Rhea" id="RHEA-COMP:9623"/>
            <dbReference type="Rhea" id="RHEA-COMP:9644"/>
            <dbReference type="Rhea" id="RHEA-COMP:9645"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:65264"/>
            <dbReference type="ChEBI" id="CHEBI:78449"/>
            <dbReference type="ChEBI" id="CHEBI:78473"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="44">
            <dbReference type="Rhea" id="RHEA:41885"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="58">
            <text>hexadecanoyl-[ACP] + H2O = hexadecanoate + holo-[ACP] + H(+)</text>
            <dbReference type="Rhea" id="RHEA:41932"/>
            <dbReference type="Rhea" id="RHEA-COMP:9652"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:7896"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78483"/>
            <dbReference type="EC" id="3.1.2.14"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="58">
            <dbReference type="Rhea" id="RHEA:41933"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="49">
            <text>hexadecanoyl-[ACP] + malonyl-[ACP] + H(+) = 3-oxooctadecanoyl-[ACP] + holo-[ACP] + CO2</text>
            <dbReference type="Rhea" id="RHEA:41916"/>
            <dbReference type="Rhea" id="RHEA-COMP:9623"/>
            <dbReference type="Rhea" id="RHEA-COMP:9652"/>
            <dbReference type="Rhea" id="RHEA-COMP:9653"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78449"/>
            <dbReference type="ChEBI" id="CHEBI:78483"/>
            <dbReference type="ChEBI" id="CHEBI:78487"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="49">
            <dbReference type="Rhea" id="RHEA:41917"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="39">
            <text>hexanoyl-[ACP] + malonyl-[ACP] + H(+) = 3-oxooctanoyl-[ACP] + holo-[ACP] + CO2</text>
            <dbReference type="Rhea" id="RHEA:41836"/>
            <dbReference type="Rhea" id="RHEA-COMP:9623"/>
            <dbReference type="Rhea" id="RHEA-COMP:9632"/>
            <dbReference type="Rhea" id="RHEA-COMP:9633"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78449"/>
            <dbReference type="ChEBI" id="CHEBI:78459"/>
            <dbReference type="ChEBI" id="CHEBI:78460"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="39">
            <dbReference type="Rhea" id="RHEA:41837"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="57">
            <text>holo-[ACP] + acetyl-CoA = acetyl-[ACP] + CoA</text>
            <dbReference type="Rhea" id="RHEA:41788"/>
            <dbReference type="Rhea" id="RHEA-COMP:9621"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57288"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78446"/>
            <dbReference type="EC" id="2.3.1.38"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="57">
            <dbReference type="Rhea" id="RHEA:41789"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="52">
            <text>holo-[ACP] + malonyl-CoA = malonyl-[ACP] + CoA</text>
            <dbReference type="Rhea" id="RHEA:41792"/>
            <dbReference type="Rhea" id="RHEA-COMP:9623"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57384"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78449"/>
            <dbReference type="EC" id="2.3.1.39"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="52">
            <dbReference type="Rhea" id="RHEA:41793"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="68">
            <text>octanoyl-[ACP] + malonyl-[ACP] + H(+) = 3-oxodecanoyl-[ACP] + holo-[ACP] + CO2</text>
            <dbReference type="Rhea" id="RHEA:41852"/>
            <dbReference type="Rhea" id="RHEA-COMP:9623"/>
            <dbReference type="Rhea" id="RHEA-COMP:9636"/>
            <dbReference type="Rhea" id="RHEA-COMP:9637"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78449"/>
            <dbReference type="ChEBI" id="CHEBI:78463"/>
            <dbReference type="ChEBI" id="CHEBI:78464"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="68">
            <dbReference type="Rhea" id="RHEA:41853"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="51">
            <text>tetradecanoyl-[ACP] + H2O = tetradecanoate + holo-[ACP] + H(+)</text>
            <dbReference type="Rhea" id="RHEA:30123"/>
            <dbReference type="Rhea" id="RHEA-COMP:9648"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30807"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78477"/>
            <dbReference type="EC" id="3.1.2.14"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="51">
            <dbReference type="Rhea" id="RHEA:30124"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="42">
            <text>tetradecanoyl-[ACP] + malonyl-[ACP] + H(+) = 3-oxohexadecanoyl-[ACP] + holo-[ACP] + CO2</text>
            <dbReference type="Rhea" id="RHEA:41900"/>
            <dbReference type="Rhea" id="RHEA-COMP:9623"/>
            <dbReference type="Rhea" id="RHEA-COMP:9648"/>
            <dbReference type="Rhea" id="RHEA-COMP:9649"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78449"/>
            <dbReference type="ChEBI" id="CHEBI:78477"/>
            <dbReference type="ChEBI" id="CHEBI:78478"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="42">
            <dbReference type="Rhea" id="RHEA:41901"/>
        </physiologicalReaction>
    </comment>
    <comment type="pathway">
        <text evidence="1">Lipid metabolism; fatty acid biosynthesis.</text>
    </comment>
    <dbReference type="EC" id="1.1.1.100" evidence="5"/>
    <dbReference type="EC" id="1.3.1.39" evidence="2"/>
    <dbReference type="EC" id="2.3.1.38" evidence="8"/>
    <dbReference type="EC" id="2.3.1.39" evidence="9"/>
    <dbReference type="EC" id="2.3.1.41" evidence="7"/>
    <dbReference type="EC" id="2.3.1.85" evidence="4"/>
    <dbReference type="EC" id="3.1.2.14" evidence="3"/>
    <dbReference type="EC" id="4.2.1.59" evidence="6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018267.1">
        <property type="protein sequence ID" value="ENSGAGP00000015987.1"/>
        <property type="gene ID" value="ENSGAGG00000012026.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00094"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019171">
        <property type="term" value="F:(3R)-hydroxyacyl-[acyl-carrier-protein] dehydratase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004316">
        <property type="term" value="F:3-oxoacyl-[acyl-carrier-protein] reductase (NADPH) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004315">
        <property type="term" value="F:3-oxoacyl-[acyl-carrier-protein] synthase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004313">
        <property type="term" value="F:[acyl-carrier-protein] S-acetyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004314">
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        <property type="entry name" value="enoyl_red"/>
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        <property type="match status" value="1"/>
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        <property type="entry name" value="Medium-chain alcohol dehydrogenases, catalytic domain"/>
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        <property type="entry name" value="Vaccinia Virus protein VP39"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="ACP-like_sf"/>
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        <property type="entry name" value="Acyl_transferase_dom"/>
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        <property type="entry name" value="Acyl_Trfase/lysoPLipase"/>
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        <property type="entry name" value="ADH-like_C"/>
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    <dbReference type="InterPro" id="IPR020843">
        <property type="entry name" value="ER"/>
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        <property type="entry name" value="FAS_pseudo-KR"/>
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        <property type="entry name" value="GroES-like_sf"/>
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        <property type="entry name" value="Ketoacyl_synth_AS"/>
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        <property type="entry name" value="Ketoacyl_synth_C"/>
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        <property type="entry name" value="Ketoacyl_synth_N"/>
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        <property type="entry name" value="KR_dom"/>
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        <property type="entry name" value="Malonyl_transacylase_ACP-bd"/>
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        <property type="entry name" value="PKS_assoc"/>
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        <property type="entry name" value="PKS_Beta-ketoAc_synthase_dom"/>
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        <property type="entry name" value="PKS_dehydratase_sf"/>
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        <property type="entry name" value="PKS_DH"/>
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        <property type="entry name" value="PKS_DH_N"/>
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        <property type="entry name" value="PKS_KR"/>
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        <property type="entry name" value="PKS_mFAS_DH"/>
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        <property type="entry name" value="PKS_NRPS_Biosynth_Enz"/>
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        <property type="entry name" value="PKS_PP-bd"/>
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    <dbReference type="InterPro" id="IPR009081">
        <property type="entry name" value="PP-bd_ACP"/>
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        <property type="entry name" value="Ppantetheine_attach_site"/>
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        <property type="entry name" value="SAM-dependent_MTases_sf"/>
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    <dbReference type="InterPro" id="IPR001031">
        <property type="entry name" value="Thioesterase"/>
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    <dbReference type="InterPro" id="IPR016039">
        <property type="entry name" value="Thiolase-like"/>
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    <dbReference type="PANTHER" id="PTHR43775">
        <property type="entry name" value="FATTY ACID SYNTHASE"/>
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        <property type="entry name" value="FATTY ACID SYNTHASE"/>
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    <dbReference type="Pfam" id="PF00698">
        <property type="entry name" value="Acyl_transf_1"/>
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        <property type="entry name" value="ADH_zinc_N"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="FAS_pseudo-KR"/>
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        <property type="entry name" value="ketoacyl-synt"/>
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    <dbReference type="Pfam" id="PF02801">
        <property type="entry name" value="Ketoacyl-synt_C"/>
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        <property type="entry name" value="KR"/>
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        <property type="entry name" value="PP-binding"/>
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        <property type="entry name" value="Thioesterase"/>
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        <property type="entry name" value="PKS_AT"/>
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    <dbReference type="SMART" id="SM00826">
        <property type="entry name" value="PKS_DH"/>
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        <property type="entry name" value="PKS_ER"/>
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        <property type="entry name" value="PKS_KR"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="PKS_KS"/>
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        <property type="entry name" value="PKS_PP"/>
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        <property type="entry name" value="alpha/beta-Hydrolases"/>
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        <property type="entry name" value="FabD/lysophospholipase-like"/>
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        <property type="entry name" value="GroES-like"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="NAD(P)-binding Rossmann-fold domains"/>
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    <dbReference type="SUPFAM" id="SSF55048">
        <property type="entry name" value="Probable ACP-binding domain of malonyl-CoA ACP transacylase"/>
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    </dbReference>
    <dbReference type="SUPFAM" id="SSF53335">
        <property type="entry name" value="S-adenosyl-L-methionine-dependent methyltransferases"/>
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    <dbReference type="SUPFAM" id="SSF53901">
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    <dbReference type="PROSITE" id="PS50075">
        <property type="entry name" value="CARRIER"/>
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    <dbReference type="PROSITE" id="PS00606">
        <property type="entry name" value="KS3_1"/>
        <property type="match status" value="1"/>
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    <dbReference type="PROSITE" id="PS52004">
        <property type="entry name" value="KS3_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00012">
        <property type="entry name" value="PHOSPHOPANTETHEINE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS52019">
        <property type="entry name" value="PKS_MFAS_DH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="20" id="KW-0007">Acetylation</keyword>
    <keyword evidence="24" id="KW-0275">Fatty acid biosynthesis</keyword>
    <keyword evidence="17" id="KW-0276">Fatty acid metabolism</keyword>
    <keyword evidence="16" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="12" id="KW-0444">Lipid biosynthesis</keyword>
    <keyword evidence="23" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="25" id="KW-0456">Lyase</keyword>
    <keyword evidence="26" id="KW-0511">Multifunctional enzyme</keyword>
    <keyword evidence="22" id="KW-0520">NAD</keyword>
    <keyword evidence="18" id="KW-0521">NADP</keyword>
    <keyword evidence="21" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="11" id="KW-0596">Phosphopantetheine</keyword>
    <keyword evidence="13" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="19" id="KW-0663">Pyridoxal phosphate</keyword>
    <keyword evidence="74" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="15" id="KW-0702">S-nitrosylation</keyword>
    <keyword evidence="14" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Ketosynthase family 3 (KS3)" evidence="71">
        <location>
            <begin position="1"/>
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        </location>
    </feature>
    <feature type="domain" description="PKS/mFAS DH" evidence="72">
        <location>
            <begin position="844"/>
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        </location>
    </feature>
    <feature type="domain" description="Carrier" evidence="70">
        <location>
            <begin position="2124"/>
            <end position="2201"/>
        </location>
    </feature>
    <feature type="region of interest" description="N-terminal hotdog fold" evidence="69">
        <location>
            <begin position="844"/>
            <end position="967"/>
        </location>
    </feature>
    <feature type="region of interest" description="C-terminal hotdog fold" evidence="69">
        <location>
            <begin position="984"/>
            <end position="1111"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor; for dehydratase activity" evidence="69">
        <location>
            <position position="878"/>
        </location>
    </feature>
    <feature type="active site" description="Proton donor; for dehydratase activity" evidence="69">
        <location>
            <position position="1034"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00005194"/>
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    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00012004"/>
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    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012480"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00012873"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012948"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00013167"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00013191"/>
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    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00013256"/>
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    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00013258"/>
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    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00018769"/>
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    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022450"/>
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    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022516"/>
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    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
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    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
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    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022799"/>
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    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
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    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022832"/>
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    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00022857"/>
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    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00022898"/>
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    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
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    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
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    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00023027"/>
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    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
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    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00023160"/>
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    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00023239"/>
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    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00023268"/>
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    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="ARBA" id="ARBA00023332"/>
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    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="ARBA" id="ARBA00023351"/>
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    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="ARBA" id="ARBA00023373"/>
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    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="ARBA" id="ARBA00023388"/>
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    </evidence>
    <evidence type="ECO:0000256" key="31">
        <source>
            <dbReference type="ARBA" id="ARBA00023394"/>
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    </evidence>
    <evidence type="ECO:0000256" key="32">
        <source>
            <dbReference type="ARBA" id="ARBA00023398"/>
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    </evidence>
    <evidence type="ECO:0000256" key="33">
        <source>
            <dbReference type="ARBA" id="ARBA00023399"/>
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    </evidence>
    <evidence type="ECO:0000256" key="34">
        <source>
            <dbReference type="ARBA" id="ARBA00023401"/>
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    </evidence>
    <evidence type="ECO:0000256" key="35">
        <source>
            <dbReference type="ARBA" id="ARBA00023402"/>
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    </evidence>
    <evidence type="ECO:0000256" key="36">
        <source>
            <dbReference type="ARBA" id="ARBA00023442"/>
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    </evidence>
    <evidence type="ECO:0000256" key="37">
        <source>
            <dbReference type="ARBA" id="ARBA00044883"/>
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    </evidence>
    <evidence type="ECO:0000256" key="38">
        <source>
            <dbReference type="ARBA" id="ARBA00047300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="39">
        <source>
            <dbReference type="ARBA" id="ARBA00047394"/>
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    </evidence>
    <evidence type="ECO:0000256" key="40">
        <source>
            <dbReference type="ARBA" id="ARBA00047400"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="41">
        <source>
            <dbReference type="ARBA" id="ARBA00047440"/>
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    </evidence>
    <evidence type="ECO:0000256" key="42">
        <source>
            <dbReference type="ARBA" id="ARBA00047451"/>
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    </evidence>
    <evidence type="ECO:0000256" key="43">
        <source>
            <dbReference type="ARBA" id="ARBA00047500"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="44">
        <source>
            <dbReference type="ARBA" id="ARBA00047578"/>
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    </evidence>
    <evidence type="ECO:0000256" key="45">
        <source>
            <dbReference type="ARBA" id="ARBA00047810"/>
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    </evidence>
    <evidence type="ECO:0000256" key="46">
        <source>
            <dbReference type="ARBA" id="ARBA00047897"/>
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    </evidence>
    <evidence type="ECO:0000256" key="47">
        <source>
            <dbReference type="ARBA" id="ARBA00047953"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="48">
        <source>
            <dbReference type="ARBA" id="ARBA00047961"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="49">
        <source>
            <dbReference type="ARBA" id="ARBA00048051"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="50">
        <source>
            <dbReference type="ARBA" id="ARBA00048281"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="51">
        <source>
            <dbReference type="ARBA" id="ARBA00048289"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="52">
        <source>
            <dbReference type="ARBA" id="ARBA00048404"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="53">
        <source>
            <dbReference type="ARBA" id="ARBA00048420"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="54">
        <source>
            <dbReference type="ARBA" id="ARBA00048506"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="55">
        <source>
            <dbReference type="ARBA" id="ARBA00048571"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="56">
        <source>
            <dbReference type="ARBA" id="ARBA00048650"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="57">
        <source>
            <dbReference type="ARBA" id="ARBA00048691"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="58">
        <source>
            <dbReference type="ARBA" id="ARBA00048704"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="59">
        <source>
            <dbReference type="ARBA" id="ARBA00048935"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="60">
        <source>
            <dbReference type="ARBA" id="ARBA00049019"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="61">
        <source>
            <dbReference type="ARBA" id="ARBA00049109"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="62">
        <source>
            <dbReference type="ARBA" id="ARBA00049171"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="63">
        <source>
            <dbReference type="ARBA" id="ARBA00049263"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="64">
        <source>
            <dbReference type="ARBA" id="ARBA00049414"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="65">
        <source>
            <dbReference type="ARBA" id="ARBA00049422"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="66">
        <source>
            <dbReference type="ARBA" id="ARBA00049449"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="67">
        <source>
            <dbReference type="ARBA" id="ARBA00049521"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="68">
        <source>
            <dbReference type="ARBA" id="ARBA00049533"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="69">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01363"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="70">
        <source>
            <dbReference type="PROSITE" id="PS50075"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="71">
        <source>
            <dbReference type="PROSITE" id="PS52004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="72">
        <source>
            <dbReference type="PROSITE" id="PS52019"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="73">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015987.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="74">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2482" mass="272963" checksum="5E11BC1BFD187C14" modified="2019-05-08" version="1">MEDVVIAGMAGKLPESENMQEFWENLFNGVDMVTEDDRRWKPGLYGLPKRNGKLKDISKFDASFFGVHPKQAHTMDPQLRLMLEASYEAILDGGINPATLRGTDTGVWIGVSGSEAGEAFSQDPEQLLGYSMTGCQRAMFANRISYFFDLKGPSMAIDTACSSSLIALENAYKAICHGECNAALVGGVNLLLKPSTSVQFMKLGMLSPDGACKSFDASGNGYCRSEAVVVILMTKKSMAKRIYATIVNAGSNTDGFKEQGVTFPSGEMQQALVSSLYKESGIQPEEVEYVEAHGTGTKIGDPQELNGIANIFCQLEREPLLIGSTKSNMGHPEPASGLAALAKVILSLEHGLWAPNIHYNTPNPEIPALQDGRVQVVAKPTPVKGGLVSINSFGFGGSNVHVILRPNKKKFYPLERHNLPRLVQVCGRTQEAVEMLIEQSRRLREYSSFVSLLNDISGIPISSMPYRGYTLIGSQSDIKEVQHTRTSGRPLWYICSGMGTQWKGMGLSLMKLELFQQSILRSDEALKDTGLKVSDLLLNADENTFEDAVHAFVGLAAIQIAQIDILKAMDLQPDGIVGHSVGELACGYADNSLSHEEAILAAYWRGRCIKEAKLPPGGMAAVGLTWEECKRHCPPGVVPACHNSEDTVTVSGPQDAVNKFVAKLKKEGVFAKEVRSAGVAFHSYYMASIAPVLLSALQKVIPNAKPRSPRWISTSIPESQWQSELAQNSSAEYHVNNLVSPVLFQEGLRHVPDNAVVVEIAPHALLQAVLRRSLKPTCTILPLMKREHKNNLEFFLTHVGKIYLTGINVLSNNLFPLVEYPAPVGTPLISPYILWDHSQTWDVPKAEDFPTGSGGSSSASIYNIDMNSESSDNYLVGHCIDGRVLYPATGYLVLAWRTLARSLGIAMEQMPVVFEEVTIHQATILSNKGSTQLEVRLLPASHCFEVSGNGNLAVSGKICLLEETALNNFRNQQVDLETPPNMRSKFSLSKADIYKELRLRGYDYGPTFQGLLESNNEGNRGKVLWNGNWVTFLDTLLQMIILGETGHSLRLPTRIRSVCIDPSLHQERVHKYKDDTEAFDVVVDRCLDNIMAGGVRLSGLHATVAPRRQQEQRPPTLEKFCFVPYVESDCLSSEAHLCAHLEHCKGLICKLQEKVAFHGVKLAIPGLETVEAFGETVSMQKGLLHILAEICHLELNGNLYSELGQIVTREKMHLQDDPLLNALLDSPELKACVDIALENITSHRMKIVEALAGDGRLFSRVTGLLNTQPMLQLDYTATDQAPEILSVNESQLQETGVSIAQWDPISPPPGHLTSADLLVCNCSVSIFGNPAEILSNLAATVKEGGFILLHTLLKGETLGEIISFLTTPDLQQRGLLAQAEWEDLFRKTLLNLVAVKKSFFGSVIFLCRRQASAKIPVFLPVEETHYKWVESLKVILADASEQPVWLTATSCGNSGIVGMVNCLRQEPGGHRIRCLFVSNLNSSLAIPPTSPSAREMQKILQGDLVMNVYRDGKWGSFRHFLLQQAQPQELTDYAYVNVLTRGDLSSLRWISSPLRHFHTTNPNIQLCRVYYASLNFRDIMLATGKLPPDAIPGNWTLQQCMLGMEFSGRDPTGKRVMGLLPAKGLATVVDSDKRFLWEVPENWTLEEAASVPVVYATAYYALVVRGGMKKGESVLIHSGSGGVGQAAIAIALSMGCRVFATVGSVEKREYLQTRFPQLDANSFANSRNTSFEQHILRVTNGKGVNLVLNSLAEEKLQASLRCLARHGRFLEIGKYDLSNNSPLGMALFLKNVAFHGILLDAIFEEGNQEWEEVSDLLKMGIRDGVVKPLRTTIFNREEVEAAFRFMAQGKHIGKVMIKVHEEEKEFPARETEAVRLSAISRTSCPPTKSYIITGGLGGFGLELAQWLVERGAQKLVLTSRSGIRSGYQAKRVRGWKEMGVQVLVSTSDVGTLEGTQHLIEEASLLGPVGGIFNLAMVLKDAMLENQSPEFFHDVSKPKYAGTLHLDWVSREKCPDLDYFVAFSSISCGRGNVGQSNYGFANSTMERICEQRQHDGLPGLAVQWGAIGDVGVVLETMGSNDTVVGGTLPQRLSSCLEVLDHFLNQSHPVMSSFVLAEKVSVKSEGSSQRDLMEAVAHILGVRDVSSLNAESSLADLGLDSLMGVEVRQTLERDYDLVMTMREIRLLTINKLRELSSKSGTAEELKPSLLMKAGQAEHPELDLNNLLVNPEGPTITRLNEVQSTERPLFLIHPIEGSITVFHTLASRLSMPSYGLQCTKAAPLDSIQSLAAYYIDCIKQVQPEGPYRIAGYSFGACVAFEMCSQLQAQQNPSHSLNSLFLFDGSHSYVAAYTQSYRAKLTPGNEAEAETEALCAFIQQFTSTEYNKLLETLLPLQDLEARVNAAVDLIMLSHKNINRETLCFAAASFYYKLKAADKYVPESKYHGNVTLLRAKTHNEYEEGLGGDYKLSEVSLRNSSRYEAALS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLY0</accession>
    <name>A0A452HLY0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Dysferlin</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="17">Dystrophy-associated fer-1-like protein</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="18">Fer-1-like protein 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">Key calcium ion sensor involved in the Ca(2+)-triggered synaptic vesicle-plasma membrane fusion. Plays a role in the sarcolemma repair mechanism of both skeletal muscle and cardiomyocytes that permits rapid resealing of membranes disrupted by mechanical stress.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <location evidence="1">Sarcolemma</location>
            <topology evidence="1">Single-pass type II membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasmic vesicle membrane</location>
            <topology evidence="2">Single-pass type II membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the ferlin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018221.1">
        <property type="protein sequence ID" value="ENSGAGP00000015946.1"/>
        <property type="gene ID" value="ENSGAGG00000011993.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030659">
        <property type="term" value="C:cytoplasmic vesicle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012505">
        <property type="term" value="C:endomembrane system"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015630">
        <property type="term" value="C:microtubule cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030315">
        <property type="term" value="C:T-tubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008289">
        <property type="term" value="F:lipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002281">
        <property type="term" value="P:macrophage activation involved in immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061025">
        <property type="term" value="P:membrane fusion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002280">
        <property type="term" value="P:monocyte activation involved in immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050765">
        <property type="term" value="P:negative regulation of phagocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007009">
        <property type="term" value="P:plasma membrane organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd08373">
        <property type="entry name" value="C2A_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04011">
        <property type="entry name" value="C2B_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04018">
        <property type="entry name" value="C2C_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04017">
        <property type="entry name" value="C2D_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04037">
        <property type="entry name" value="C2E_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd08374">
        <property type="entry name" value="C2F_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000009">
        <property type="entry name" value="dysferlin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000021">
        <property type="entry name" value="dysferlin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000026">
        <property type="entry name" value="dysferlin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000033">
        <property type="entry name" value="dysferlin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000037">
        <property type="entry name" value="dysferlin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000061">
        <property type="entry name" value="dysferlin isoform X8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000008">
        <property type="entry name" value="C2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037726">
        <property type="entry name" value="C2A_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037720">
        <property type="entry name" value="C2B_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037722">
        <property type="entry name" value="C2C_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037723">
        <property type="entry name" value="C2D_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037724">
        <property type="entry name" value="C2E_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037725">
        <property type="entry name" value="C2F_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012968">
        <property type="entry name" value="FerIin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037721">
        <property type="entry name" value="Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012560">
        <property type="entry name" value="Ferlin_A-domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012561">
        <property type="entry name" value="Ferlin_B-domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032362">
        <property type="entry name" value="Ferlin_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055072">
        <property type="entry name" value="Ferlin_DSRM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006614">
        <property type="entry name" value="Peroxin/Ferlin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12546:SF44">
        <property type="entry name" value="DYSFERLIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12546">
        <property type="entry name" value="FER-1-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00168">
        <property type="entry name" value="C2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22901">
        <property type="entry name" value="dsrm_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08165">
        <property type="entry name" value="FerA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08150">
        <property type="entry name" value="FerB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08151">
        <property type="entry name" value="FerI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16165">
        <property type="entry name" value="Ferlin_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00239">
        <property type="entry name" value="C2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SMART" id="SM00694">
        <property type="entry name" value="DysFC"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00693">
        <property type="entry name" value="DysFN"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM01200">
        <property type="entry name" value="FerA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01201">
        <property type="entry name" value="FerB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01202">
        <property type="entry name" value="FerI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49562">
        <property type="entry name" value="C2 domain (Calcium/lipid-binding domain, CaLB)"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50004">
        <property type="entry name" value="C2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0106">Calcium</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="14" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="12" id="KW-0446">Lipid-binding</keyword>
    <keyword evidence="13 20" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="10" id="KW-0735">Signal-anchor</keyword>
    <keyword evidence="6 20" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 20" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="2058"/>
            <end position="2080"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="1"/>
            <end position="101"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="197"/>
            <end position="315"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="354"/>
            <end position="490"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="1130"/>
            <end position="1256"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="1303"/>
            <end position="1431"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="1574"/>
            <end position="1692"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="1808"/>
            <end position="1956"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="127"/>
            <end position="208"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="2009"/>
            <end position="2030"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="19">
        <location>
            <begin position="127"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="172"/>
            <end position="182"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="19">
        <location>
            <begin position="188"/>
            <end position="203"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="2011"/>
            <end position="2030"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004274"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007561"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023121"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00060070"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00068801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00078385"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00079079"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015946.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2093" mass="238982" checksum="3A50206F4BAC7A0D" modified="2019-05-08" version="1">MLRVFILHAENVQTPDTDISDSYCSVVFEGVKKRTKVIKNNVNPVWNEGFDWDLKGIPLDQNAELTVVVKDHETVGRNRFLGEARVPLQDVLVNPTLTASFNAPLLDTKRQSTGAFLNLQVSYIPPPGAAPLFPPPTPPEPTPSTAELDTVTDTAGEEETEDQIVTGDETEPSTGPSGSEQPTLPRKPPSHHVHGIKRKKSAPKKLLSNKPQDFQIRVRVIEGRQLPGVNIKSVVKVTVAGQTKRTRIRKGNGPFFDEIFFFNVFESPAELFDEPIFITVVDSRSLRTDSVIGEFQLDVGTVYAEPKHALLRKWLLLSDSEDFSAGAKGYVKVSLFVLGPGDEAPLEKKEVAEEKEDIEGNLLRPTGVTLRGAHFCLKIYRAEDLPQMDDAIMDNVKQIFGFESNKKNLVDPFVEVSFAGRTISSKILEKNANPQWNQRITVPAMFPSMCEKMRIRVMDWDRVTHNDVIGTTHLRLSQISAPGGELEDEFPVLAKPLKAAAMDDGLGFLPTFGPCYVNLYGSPREFTGFPDPYEELNMGKGEGVAFRGRLLMELETKLVEHIEQKLEDIPPDDILRVEKHLRRRKYCLFAAFYSASMLQDVDAAIQFEVSIGNYGNKFDNTCLPLASTTQYSGAVFDGCQYYYLPWGNVKPVVVLSSYWEDISHRTDMQNLLQRTADRLEAGLELVHLALKARCSSNELDSLVSQLIDDIIADCSQPLVDVSNKTASTHLDQFVYKFRSTNLHQIIQAALNLKHEDSDLQTVLDQAEDWLLRLKAMAEEPQNSLPDIVIWMLQGDKRVAYARVPAHEVLFSRTGANCCGKNCGKLQTIFLKYPQEEVMGPRIPAQVRVQLWFGLAVDEKEFNKYAEGKLSVFAETYENQVKLALVGNWGTTGLTYPKYSDVTGKIKLPKDSFRPSVGWTWAGDWFISPEKTLLYDTDAGHMSFVEEVFENQVRLPGGQWIHMADAYTDVNGEKVLPKEEIECPPGWKWEDVEWDTDLNRAVDERGWEYGIAIPPDHKPKAWVPAEKMYHTSRRRRWVRLRRRDLKQMEALKKHKQEELEGEGWEYASLFGWKFHLNYRKTDTFRRRRWRRRMEPLERTGAAAVFALEGALGGVTDDKSDDGKSVSTLSFGVNRPTVSCIFDYGSRYHLRCYIYQARDLIAMDKDSFSDPYAIVSFLHQSQKTVVVKNTLNPLWDQTLIFYEIEIFGDPQNVADVPPNIVVEIYDHDTYGADEFMGRCICKPSLERSPRLSWHPVARGNRKAGELLAAFELIQREKPAVHHLPGFESELSASLDESADSDLPYPPPQREPNIYMVPQGIKPVLQRTAIEILAWGLRNLKSYQLASITSPSLQVECGGQTVQSCVIKNVRKNPNFDVCVLFMEVRLPKEALYCPPVIMKVIDNRQFGRKPVVGQCTIQSLEEFYCDPYKEETDAPEEQPDEVSLTPRDDVLIDIDDKEPLIPVQLAEGMTSSTLINMPASRADLHEEEFIDWWSKFYASTGEREKCGSYLEKGFDTLKVYETELENMEEFERLSDFCHTFKLYRGRTQDINEDPSVVGEFKGSFKMYPLPDDPRVPVPPRQFHQLPAKGPQECLIRIYIIRAFDLQPKDSNGKCDPYVKISVGKKSINDQENYIPCTLEPVFGKMFELTCTLPLEKDLKVTLYDYDLLSKDEKIGETVIDLENRFLSKFGARCGLPQTYCISGPNQWRDQLRPSQLLHLFSLQRNCKAPIYKPDRVVFREQEYFLSEFEDGKPPNPHLGAPEERLALYTLRKQGLVPEHVETRPLYSPLQPEIEQGKLQMWVDLFPKSLGLPGPPFNITPRRAKRFFLRCIIWNTKDVILDDLSITGEKMSDIYVKGWMTGHEENKQKTDVHYRSMGGEGNFNWRFVFPFDYLAAEQVCHIAKKEHFWSLDKTDNKVPPQLILQIWDNDKFSFDDYLGSVQMNLNRMPAPTKTAEKCSLNLLDETLSSDRFVSLFEQKAVKGWWPCVVEAEEKKTIAGKLEMTLEIVTEQEQEERPAGMGRDEPNMNPKLEDPKRPETSFLWFTSPYKTMKYILWRRFKWVFLIAIVVFILLLFIGIFIYAFPNYAAMKLVKPFS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLY3</accession>
    <name>A0A452HLY3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">LRAT domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the H-rev107 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLY3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018224.1">
        <property type="protein sequence ID" value="ENSGAGP00000015949.1"/>
        <property type="gene ID" value="ENSGAGG00000012005.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016410">
        <property type="term" value="F:N-acyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008970">
        <property type="term" value="F:phospholipase A1 activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004623">
        <property type="term" value="F:phospholipase A2 activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070292">
        <property type="term" value="P:N-acylphosphatidylethanolamine metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1720.10">
        <property type="entry name" value="endopeptidase domain like (from Nostoc punctiforme)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051496">
        <property type="entry name" value="H-rev107_PLA/AT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007053">
        <property type="entry name" value="LRAT_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13943">
        <property type="entry name" value="HRAS-LIKE SUPPRESSOR - RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13943:SF31">
        <property type="entry name" value="PHOSPHOLIPASE A AND ACYLTRANSFERASE 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04970">
        <property type="entry name" value="LRAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51934">
        <property type="entry name" value="LRAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="4" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="LRAT" evidence="5">
        <location>
            <begin position="86"/>
            <end position="198"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS51934"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015949.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="243" mass="27087" checksum="5EC609C0444FC244" modified="2019-05-08" version="1">MEPIERQILPSPLCTLGQSEREALPGVCVTLAPLGVFPYSSEIWGRQQPAWTIQSRDLPPRCFCSELWLCAVCLGDLKACRRRDEKKPENRGLDTCRGPRHLENISCHCEIPGAGFASLMSTLTNKAMVKKERLLDVVGRHRYRVNNKHDRKFPPLPAGKIVQAAEQLVGQEMLYRVTSENCEHFVTELRYGVAKSDQVLGLAPCHPDRTPGFSPQPLGGERGLVVRAGRPVSQAFWILCSSS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="22" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLY5</accession>
    <name>A0A452HLY5_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="1">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="1 2">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="2" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="1" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="1" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000018226.1">
        <property type="protein sequence ID" value="ENSGAGP00000015951.1"/>
        <property type="gene ID" value="ENSGAGG00000012004.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000313" key="1">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015951.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="233" mass="24359" checksum="13D29118C04C6F0D" modified="2025-10-08" version="2">MXXLCCREPCRNHDLGCLNVAGKCRPSTPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRLDPCPHRPGCWASLACASGGKGRKIIWI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM62</accession>
    <name>A0A452HM62_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Fatty acid synthase</fullName>
            <ecNumber evidence="5">1.1.1.100</ecNumber>
            <ecNumber evidence="2">1.3.1.39</ecNumber>
            <ecNumber evidence="8">2.3.1.38</ecNumber>
            <ecNumber evidence="9">2.3.1.39</ecNumber>
            <ecNumber evidence="7">2.3.1.41</ecNumber>
            <ecNumber evidence="4">2.3.1.85</ecNumber>
            <ecNumber evidence="3">3.1.2.14</ecNumber>
            <ecNumber evidence="6">4.2.1.59</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="73 74">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="74" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="73" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="73" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="36">Fatty acid synthetase is a multifunctional enzyme that catalyzes the de novo biosynthesis of long-chain saturated fatty acids starting from acetyl-CoA and malonyl-CoA in the presence of NADPH. This multifunctional protein contains 7 catalytic activities and a site for the binding of the prosthetic group 4'-phosphopantetheine of the acyl carrier protein ([ACP]) domain.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="43">
            <text>(2E)-butenoyl-[ACP] + NADPH + H(+) = butanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41812"/>
            <dbReference type="Rhea" id="RHEA-COMP:9627"/>
            <dbReference type="Rhea" id="RHEA-COMP:9628"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78453"/>
            <dbReference type="ChEBI" id="CHEBI:78454"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="43">
            <dbReference type="Rhea" id="RHEA:41813"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="67">
            <text>(2E)-decenoyl-[ACP] + NADPH + H(+) = decanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41864"/>
            <dbReference type="Rhea" id="RHEA-COMP:9639"/>
            <dbReference type="Rhea" id="RHEA-COMP:9640"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78467"/>
            <dbReference type="ChEBI" id="CHEBI:78468"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="67">
            <dbReference type="Rhea" id="RHEA:41865"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="50">
            <text>(2E)-dodecenoyl-[ACP] + NADPH + H(+) = dodecanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41880"/>
            <dbReference type="Rhea" id="RHEA-COMP:9643"/>
            <dbReference type="Rhea" id="RHEA-COMP:9644"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:65264"/>
            <dbReference type="ChEBI" id="CHEBI:78472"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="50">
            <dbReference type="Rhea" id="RHEA:41881"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="45">
            <text>(2E)-hexadecenoyl-[ACP] + NADPH + H(+) = hexadecanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41912"/>
            <dbReference type="Rhea" id="RHEA-COMP:9651"/>
            <dbReference type="Rhea" id="RHEA-COMP:9652"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78481"/>
            <dbReference type="ChEBI" id="CHEBI:78483"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="45">
            <dbReference type="Rhea" id="RHEA:41913"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="46">
            <text>(2E)-hexenoyl-[ACP] + NADPH + H(+) = hexanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41832"/>
            <dbReference type="Rhea" id="RHEA-COMP:9631"/>
            <dbReference type="Rhea" id="RHEA-COMP:9632"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78458"/>
            <dbReference type="ChEBI" id="CHEBI:78459"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="46">
            <dbReference type="Rhea" id="RHEA:41833"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="60">
            <text>(2E)-octadecenoyl-[ACP] + NADPH + H(+) = octadecanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41928"/>
            <dbReference type="Rhea" id="RHEA-COMP:9655"/>
            <dbReference type="Rhea" id="RHEA-COMP:9656"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78489"/>
            <dbReference type="ChEBI" id="CHEBI:78495"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="60">
            <dbReference type="Rhea" id="RHEA:41929"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="53">
            <text>(2E)-octenoyl-[ACP] + NADPH + H(+) = octanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41848"/>
            <dbReference type="Rhea" id="RHEA-COMP:9635"/>
            <dbReference type="Rhea" id="RHEA-COMP:9636"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78462"/>
            <dbReference type="ChEBI" id="CHEBI:78463"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="53">
            <dbReference type="Rhea" id="RHEA:41849"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="62">
            <text>(2E)-tetradecenoyl-[ACP] + NADPH + H(+) = tetradecanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41896"/>
            <dbReference type="Rhea" id="RHEA-COMP:9647"/>
            <dbReference type="Rhea" id="RHEA-COMP:9648"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78475"/>
            <dbReference type="ChEBI" id="CHEBI:78477"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="62">
            <dbReference type="Rhea" id="RHEA:41897"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="35">
            <text>(3R)-hydroxybutanoyl-[ACP] = (2E)-butenoyl-[ACP] + H2O</text>
            <dbReference type="Rhea" id="RHEA:41808"/>
            <dbReference type="Rhea" id="RHEA-COMP:9626"/>
            <dbReference type="Rhea" id="RHEA-COMP:9627"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:78451"/>
            <dbReference type="ChEBI" id="CHEBI:78453"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="35">
            <dbReference type="Rhea" id="RHEA:41809"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="30">
            <text>(3R)-hydroxydecanoyl-[ACP] = (2E)-decenoyl-[ACP] + H2O</text>
            <dbReference type="Rhea" id="RHEA:41860"/>
            <dbReference type="Rhea" id="RHEA-COMP:9638"/>
            <dbReference type="Rhea" id="RHEA-COMP:9639"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:78466"/>
            <dbReference type="ChEBI" id="CHEBI:78467"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="30">
            <dbReference type="Rhea" id="RHEA:41861"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="28">
            <text>(3R)-hydroxydodecanoyl-[ACP] = (2E)-dodecenoyl-[ACP] + H2O</text>
            <dbReference type="Rhea" id="RHEA:41876"/>
            <dbReference type="Rhea" id="RHEA-COMP:9642"/>
            <dbReference type="Rhea" id="RHEA-COMP:9643"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:78470"/>
            <dbReference type="ChEBI" id="CHEBI:78472"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="28">
            <dbReference type="Rhea" id="RHEA:41877"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="34">
            <text>(3R)-hydroxyhexadecanoyl-[ACP] = (2E)-hexadecenoyl-[ACP] + H2O</text>
            <dbReference type="Rhea" id="RHEA:41908"/>
            <dbReference type="Rhea" id="RHEA-COMP:9650"/>
            <dbReference type="Rhea" id="RHEA-COMP:9651"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:78480"/>
            <dbReference type="ChEBI" id="CHEBI:78481"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="34">
            <dbReference type="Rhea" id="RHEA:41909"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="29">
            <text>(3R)-hydroxyhexanoyl-[ACP] = (2E)-hexenoyl-[ACP] + H2O</text>
            <dbReference type="Rhea" id="RHEA:41828"/>
            <dbReference type="Rhea" id="RHEA-COMP:9630"/>
            <dbReference type="Rhea" id="RHEA-COMP:9631"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:78457"/>
            <dbReference type="ChEBI" id="CHEBI:78458"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="29">
            <dbReference type="Rhea" id="RHEA:41829"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="33">
            <text>(3R)-hydroxyoctadecanoyl-[ACP] = (2E)-octadecenoyl-[ACP] + H2O</text>
            <dbReference type="Rhea" id="RHEA:41924"/>
            <dbReference type="Rhea" id="RHEA-COMP:9654"/>
            <dbReference type="Rhea" id="RHEA-COMP:9655"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:78488"/>
            <dbReference type="ChEBI" id="CHEBI:78489"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="33">
            <dbReference type="Rhea" id="RHEA:41925"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="27">
            <text>(3R)-hydroxyoctanoyl-[ACP] = (2E)-octenoyl-[ACP] + H2O</text>
            <dbReference type="Rhea" id="RHEA:41844"/>
            <dbReference type="Rhea" id="RHEA-COMP:9634"/>
            <dbReference type="Rhea" id="RHEA-COMP:9635"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:78461"/>
            <dbReference type="ChEBI" id="CHEBI:78462"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="27">
            <dbReference type="Rhea" id="RHEA:41845"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="32">
            <text>(3R)-hydroxytetradecanoyl-[ACP] = (2E)-tetradecenoyl-[ACP] + H2O</text>
            <dbReference type="Rhea" id="RHEA:41892"/>
            <dbReference type="Rhea" id="RHEA-COMP:9646"/>
            <dbReference type="Rhea" id="RHEA-COMP:9647"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:78474"/>
            <dbReference type="ChEBI" id="CHEBI:78475"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="32">
            <dbReference type="Rhea" id="RHEA:41893"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="47">
            <text>3-oxobutanoyl-[ACP] + NADPH + H(+) = (3R)-hydroxybutanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41804"/>
            <dbReference type="Rhea" id="RHEA-COMP:9625"/>
            <dbReference type="Rhea" id="RHEA-COMP:9626"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78450"/>
            <dbReference type="ChEBI" id="CHEBI:78451"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="47">
            <dbReference type="Rhea" id="RHEA:41805"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="41">
            <text>3-oxodecanoyl-[ACP] + NADPH + H(+) = (3R)-hydroxydecanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41856"/>
            <dbReference type="Rhea" id="RHEA-COMP:9637"/>
            <dbReference type="Rhea" id="RHEA-COMP:9638"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78464"/>
            <dbReference type="ChEBI" id="CHEBI:78466"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="41">
            <dbReference type="Rhea" id="RHEA:41857"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="63">
            <text>3-oxododecanoyl-[ACP] + NADPH + H(+) = (3R)-hydroxydodecanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41872"/>
            <dbReference type="Rhea" id="RHEA-COMP:9641"/>
            <dbReference type="Rhea" id="RHEA-COMP:9642"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78469"/>
            <dbReference type="ChEBI" id="CHEBI:78470"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="63">
            <dbReference type="Rhea" id="RHEA:41873"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="64">
            <text>3-oxohexadecanoyl-[ACP] + NADPH + H(+) = (3R)-hydroxyhexadecanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41904"/>
            <dbReference type="Rhea" id="RHEA-COMP:9649"/>
            <dbReference type="Rhea" id="RHEA-COMP:9650"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78478"/>
            <dbReference type="ChEBI" id="CHEBI:78480"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="64">
            <dbReference type="Rhea" id="RHEA:41905"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="55">
            <text>3-oxohexanoyl-[ACP] + NADPH + H(+) = (3R)-hydroxyhexanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41824"/>
            <dbReference type="Rhea" id="RHEA-COMP:9629"/>
            <dbReference type="Rhea" id="RHEA-COMP:9630"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78456"/>
            <dbReference type="ChEBI" id="CHEBI:78457"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="55">
            <dbReference type="Rhea" id="RHEA:41825"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="38">
            <text>3-oxooctadecanoyl-[ACP] + NADPH + H(+) = (3R)-hydroxyoctadecanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41920"/>
            <dbReference type="Rhea" id="RHEA-COMP:9653"/>
            <dbReference type="Rhea" id="RHEA-COMP:9654"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78487"/>
            <dbReference type="ChEBI" id="CHEBI:78488"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="38">
            <dbReference type="Rhea" id="RHEA:41921"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="65">
            <text>3-oxooctanoyl-[ACP] + NADPH + H(+) = (3R)-hydroxyoctanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41840"/>
            <dbReference type="Rhea" id="RHEA-COMP:9633"/>
            <dbReference type="Rhea" id="RHEA-COMP:9634"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78460"/>
            <dbReference type="ChEBI" id="CHEBI:78461"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="65">
            <dbReference type="Rhea" id="RHEA:41841"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="59">
            <text>3-oxotetradecanoyl-[ACP] + NADPH + H(+) = (3R)-hydroxytetradecanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41888"/>
            <dbReference type="Rhea" id="RHEA-COMP:9645"/>
            <dbReference type="Rhea" id="RHEA-COMP:9646"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78473"/>
            <dbReference type="ChEBI" id="CHEBI:78474"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="59">
            <dbReference type="Rhea" id="RHEA:41889"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="40">
            <text>a (3R)-hydroxyacyl-[ACP] + NADP(+) = a 3-oxoacyl-[ACP] + NADPH + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17397"/>
            <dbReference type="Rhea" id="RHEA-COMP:9916"/>
            <dbReference type="Rhea" id="RHEA-COMP:9945"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78776"/>
            <dbReference type="ChEBI" id="CHEBI:78827"/>
            <dbReference type="EC" id="1.1.1.100"/>
        </reaction>
        <physiologicalReaction direction="right-to-left" evidence="40">
            <dbReference type="Rhea" id="RHEA:17399"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="31">
            <text>a (3R)-hydroxyacyl-[ACP] = a (2E)-enoyl-[ACP] + H2O</text>
            <dbReference type="Rhea" id="RHEA:13097"/>
            <dbReference type="Rhea" id="RHEA-COMP:9925"/>
            <dbReference type="Rhea" id="RHEA-COMP:9945"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:78784"/>
            <dbReference type="ChEBI" id="CHEBI:78827"/>
            <dbReference type="EC" id="4.2.1.59"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="31">
            <dbReference type="Rhea" id="RHEA:13098"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="56">
            <text>a 2,3-saturated acyl-[ACP] + NADP(+) = a (2E)-enoyl-[ACP] + NADPH + H(+)</text>
            <dbReference type="Rhea" id="RHEA:22564"/>
            <dbReference type="Rhea" id="RHEA-COMP:9925"/>
            <dbReference type="Rhea" id="RHEA-COMP:9926"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78784"/>
            <dbReference type="ChEBI" id="CHEBI:78785"/>
            <dbReference type="EC" id="1.3.1.39"/>
        </reaction>
        <physiologicalReaction direction="right-to-left" evidence="56">
            <dbReference type="Rhea" id="RHEA:22566"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="54">
            <text>a fatty acyl-[ACP] + malonyl-[ACP] + H(+) = a 3-oxoacyl-[ACP] + holo-[ACP] + CO2</text>
            <dbReference type="Rhea" id="RHEA:22836"/>
            <dbReference type="Rhea" id="RHEA-COMP:9623"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="Rhea" id="RHEA-COMP:9916"/>
            <dbReference type="Rhea" id="RHEA-COMP:14125"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78449"/>
            <dbReference type="ChEBI" id="CHEBI:78776"/>
            <dbReference type="ChEBI" id="CHEBI:138651"/>
            <dbReference type="EC" id="2.3.1.41"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="54">
            <dbReference type="Rhea" id="RHEA:22837"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="37">
            <text>acetyl-CoA + n malonyl-CoA + 2n NADPH + 2n H(+) = a long-chain fatty acid + (n+1) CoA + n CO2 + 2n NADP(+).</text>
            <dbReference type="EC" id="2.3.1.85"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="48">
            <text>acetyl-[ACP] + malonyl-[ACP] + H(+) = 3-oxobutanoyl-[ACP] + holo-[ACP] + CO2</text>
            <dbReference type="Rhea" id="RHEA:41800"/>
            <dbReference type="Rhea" id="RHEA-COMP:9621"/>
            <dbReference type="Rhea" id="RHEA-COMP:9623"/>
            <dbReference type="Rhea" id="RHEA-COMP:9625"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78446"/>
            <dbReference type="ChEBI" id="CHEBI:78449"/>
            <dbReference type="ChEBI" id="CHEBI:78450"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="48">
            <dbReference type="Rhea" id="RHEA:41801"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="66">
            <text>butanoyl-[ACP] + malonyl-[ACP] + H(+) = 3-oxohexanoyl-[ACP] + holo-[ACP] + CO2</text>
            <dbReference type="Rhea" id="RHEA:41820"/>
            <dbReference type="Rhea" id="RHEA-COMP:9623"/>
            <dbReference type="Rhea" id="RHEA-COMP:9628"/>
            <dbReference type="Rhea" id="RHEA-COMP:9629"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78449"/>
            <dbReference type="ChEBI" id="CHEBI:78454"/>
            <dbReference type="ChEBI" id="CHEBI:78456"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="66">
            <dbReference type="Rhea" id="RHEA:41821"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="61">
            <text>decanoyl-[ACP] + malonyl-[ACP] + H(+) = 3-oxododecanoyl-[ACP] + holo-[ACP] + CO2</text>
            <dbReference type="Rhea" id="RHEA:41868"/>
            <dbReference type="Rhea" id="RHEA-COMP:9623"/>
            <dbReference type="Rhea" id="RHEA-COMP:9640"/>
            <dbReference type="Rhea" id="RHEA-COMP:9641"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78449"/>
            <dbReference type="ChEBI" id="CHEBI:78468"/>
            <dbReference type="ChEBI" id="CHEBI:78469"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="61">
            <dbReference type="Rhea" id="RHEA:41869"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="44">
            <text>dodecanoyl-[ACP] + malonyl-[ACP] + H(+) = 3-oxotetradecanoyl-[ACP] + holo-[ACP] + CO2</text>
            <dbReference type="Rhea" id="RHEA:41884"/>
            <dbReference type="Rhea" id="RHEA-COMP:9623"/>
            <dbReference type="Rhea" id="RHEA-COMP:9644"/>
            <dbReference type="Rhea" id="RHEA-COMP:9645"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:65264"/>
            <dbReference type="ChEBI" id="CHEBI:78449"/>
            <dbReference type="ChEBI" id="CHEBI:78473"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="44">
            <dbReference type="Rhea" id="RHEA:41885"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="58">
            <text>hexadecanoyl-[ACP] + H2O = hexadecanoate + holo-[ACP] + H(+)</text>
            <dbReference type="Rhea" id="RHEA:41932"/>
            <dbReference type="Rhea" id="RHEA-COMP:9652"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:7896"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78483"/>
            <dbReference type="EC" id="3.1.2.14"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="58">
            <dbReference type="Rhea" id="RHEA:41933"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="49">
            <text>hexadecanoyl-[ACP] + malonyl-[ACP] + H(+) = 3-oxooctadecanoyl-[ACP] + holo-[ACP] + CO2</text>
            <dbReference type="Rhea" id="RHEA:41916"/>
            <dbReference type="Rhea" id="RHEA-COMP:9623"/>
            <dbReference type="Rhea" id="RHEA-COMP:9652"/>
            <dbReference type="Rhea" id="RHEA-COMP:9653"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78449"/>
            <dbReference type="ChEBI" id="CHEBI:78483"/>
            <dbReference type="ChEBI" id="CHEBI:78487"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="49">
            <dbReference type="Rhea" id="RHEA:41917"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="39">
            <text>hexanoyl-[ACP] + malonyl-[ACP] + H(+) = 3-oxooctanoyl-[ACP] + holo-[ACP] + CO2</text>
            <dbReference type="Rhea" id="RHEA:41836"/>
            <dbReference type="Rhea" id="RHEA-COMP:9623"/>
            <dbReference type="Rhea" id="RHEA-COMP:9632"/>
            <dbReference type="Rhea" id="RHEA-COMP:9633"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78449"/>
            <dbReference type="ChEBI" id="CHEBI:78459"/>
            <dbReference type="ChEBI" id="CHEBI:78460"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="39">
            <dbReference type="Rhea" id="RHEA:41837"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="57">
            <text>holo-[ACP] + acetyl-CoA = acetyl-[ACP] + CoA</text>
            <dbReference type="Rhea" id="RHEA:41788"/>
            <dbReference type="Rhea" id="RHEA-COMP:9621"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57288"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78446"/>
            <dbReference type="EC" id="2.3.1.38"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="57">
            <dbReference type="Rhea" id="RHEA:41789"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="52">
            <text>holo-[ACP] + malonyl-CoA = malonyl-[ACP] + CoA</text>
            <dbReference type="Rhea" id="RHEA:41792"/>
            <dbReference type="Rhea" id="RHEA-COMP:9623"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57384"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78449"/>
            <dbReference type="EC" id="2.3.1.39"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="52">
            <dbReference type="Rhea" id="RHEA:41793"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="68">
            <text>octanoyl-[ACP] + malonyl-[ACP] + H(+) = 3-oxodecanoyl-[ACP] + holo-[ACP] + CO2</text>
            <dbReference type="Rhea" id="RHEA:41852"/>
            <dbReference type="Rhea" id="RHEA-COMP:9623"/>
            <dbReference type="Rhea" id="RHEA-COMP:9636"/>
            <dbReference type="Rhea" id="RHEA-COMP:9637"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78449"/>
            <dbReference type="ChEBI" id="CHEBI:78463"/>
            <dbReference type="ChEBI" id="CHEBI:78464"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="68">
            <dbReference type="Rhea" id="RHEA:41853"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="51">
            <text>tetradecanoyl-[ACP] + H2O = tetradecanoate + holo-[ACP] + H(+)</text>
            <dbReference type="Rhea" id="RHEA:30123"/>
            <dbReference type="Rhea" id="RHEA-COMP:9648"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30807"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78477"/>
            <dbReference type="EC" id="3.1.2.14"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="51">
            <dbReference type="Rhea" id="RHEA:30124"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="42">
            <text>tetradecanoyl-[ACP] + malonyl-[ACP] + H(+) = 3-oxohexadecanoyl-[ACP] + holo-[ACP] + CO2</text>
            <dbReference type="Rhea" id="RHEA:41900"/>
            <dbReference type="Rhea" id="RHEA-COMP:9623"/>
            <dbReference type="Rhea" id="RHEA-COMP:9648"/>
            <dbReference type="Rhea" id="RHEA-COMP:9649"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78449"/>
            <dbReference type="ChEBI" id="CHEBI:78477"/>
            <dbReference type="ChEBI" id="CHEBI:78478"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="42">
            <dbReference type="Rhea" id="RHEA:41901"/>
        </physiologicalReaction>
    </comment>
    <comment type="pathway">
        <text evidence="1">Lipid metabolism; fatty acid biosynthesis.</text>
    </comment>
    <dbReference type="EC" id="1.1.1.100" evidence="5"/>
    <dbReference type="EC" id="1.3.1.39" evidence="2"/>
    <dbReference type="EC" id="2.3.1.38" evidence="8"/>
    <dbReference type="EC" id="2.3.1.39" evidence="9"/>
    <dbReference type="EC" id="2.3.1.41" evidence="7"/>
    <dbReference type="EC" id="2.3.1.85" evidence="4"/>
    <dbReference type="EC" id="3.1.2.14" evidence="3"/>
    <dbReference type="EC" id="4.2.1.59" evidence="6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018314.1">
        <property type="protein sequence ID" value="ENSGAGP00000016031.1"/>
        <property type="gene ID" value="ENSGAGG00000012026.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00094"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019171">
        <property type="term" value="F:(3R)-hydroxyacyl-[acyl-carrier-protein] dehydratase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004316">
        <property type="term" value="F:3-oxoacyl-[acyl-carrier-protein] reductase (NADPH) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004315">
        <property type="term" value="F:3-oxoacyl-[acyl-carrier-protein] synthase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004313">
        <property type="term" value="F:[acyl-carrier-protein] S-acetyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004314">
        <property type="term" value="F:[acyl-carrier-protein] S-malonyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0141148">
        <property type="term" value="F:enoyl-[acyl-carrier-protein] reductase (NADPH) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004312">
        <property type="term" value="F:fatty acid synthase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016297">
        <property type="term" value="F:fatty acyl-[ACP] hydrolase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031177">
        <property type="term" value="F:phosphopantetheine binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006633">
        <property type="term" value="P:fatty acid biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="CDD" id="cd05195">
        <property type="entry name" value="enoyl_red"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd08954">
        <property type="entry name" value="KR_1_FAS_SDR_x"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="CDD" id="cd00833">
        <property type="entry name" value="PKS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.1200.10:FF:000013">
        <property type="entry name" value="Fatty acid synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.129.110:FF:000002">
        <property type="entry name" value="Fatty acid synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.366.10:FF:000005">
        <property type="entry name" value="Fatty acid synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1820:FF:000059">
        <property type="entry name" value="Fatty acid synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1820:FF:000105">
        <property type="entry name" value="Fatty acid synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.720:FF:000249">
        <property type="entry name" value="Fatty acid synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.180.10:FF:000015">
        <property type="entry name" value="Fatty acid synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.720:FF:000209">
        <property type="entry name" value="Polyketide synthase Pks12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.3290">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.47.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1200.10">
        <property type="entry name" value="ACP-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1820">
        <property type="entry name" value="alpha/beta hydrolase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.366.10">
        <property type="entry name" value="Malonyl-Coenzyme A Acyl Carrier Protein, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.180.10">
        <property type="entry name" value="Medium-chain alcohol dehydrogenases, catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.720">
        <property type="entry name" value="NAD(P)-binding Rossmann-like Domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.129.110">
        <property type="entry name" value="Polyketide synthase dehydratase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.150">
        <property type="entry name" value="Vaccinia Virus protein VP39"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029058">
        <property type="entry name" value="AB_hydrolase_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001227">
        <property type="entry name" value="Ac_transferase_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036736">
        <property type="entry name" value="ACP-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014043">
        <property type="entry name" value="Acyl_transferase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016035">
        <property type="entry name" value="Acyl_Trfase/lysoPLipase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013149">
        <property type="entry name" value="ADH-like_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020843">
        <property type="entry name" value="ER"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049391">
        <property type="entry name" value="FAS_pseudo-KR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011032">
        <property type="entry name" value="GroES-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018201">
        <property type="entry name" value="Ketoacyl_synth_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014031">
        <property type="entry name" value="Ketoacyl_synth_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014030">
        <property type="entry name" value="Ketoacyl_synth_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057326">
        <property type="entry name" value="KR_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016036">
        <property type="entry name" value="Malonyl_transacylase_ACP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036291">
        <property type="entry name" value="NAD(P)-bd_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032821">
        <property type="entry name" value="PKS_assoc"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020841">
        <property type="entry name" value="PKS_Beta-ketoAc_synthase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042104">
        <property type="entry name" value="PKS_dehydratase_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020807">
        <property type="entry name" value="PKS_DH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049552">
        <property type="entry name" value="PKS_DH_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013968">
        <property type="entry name" value="PKS_KR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049900">
        <property type="entry name" value="PKS_mFAS_DH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050091">
        <property type="entry name" value="PKS_NRPS_Biosynth_Enz"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020806">
        <property type="entry name" value="PKS_PP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009081">
        <property type="entry name" value="PP-bd_ACP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006162">
        <property type="entry name" value="Ppantetheine_attach_site"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029063">
        <property type="entry name" value="SAM-dependent_MTases_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001031">
        <property type="entry name" value="Thioesterase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016039">
        <property type="entry name" value="Thiolase-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43775">
        <property type="entry name" value="FATTY ACID SYNTHASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43775:SF7">
        <property type="entry name" value="FATTY ACID SYNTHASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00698">
        <property type="entry name" value="Acyl_transf_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00107">
        <property type="entry name" value="ADH_zinc_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21149">
        <property type="entry name" value="FAS_pseudo-KR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16197">
        <property type="entry name" value="KAsynt_C_assoc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00109">
        <property type="entry name" value="ketoacyl-synt"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02801">
        <property type="entry name" value="Ketoacyl-synt_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08659">
        <property type="entry name" value="KR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21089">
        <property type="entry name" value="PKS_DH_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00550">
        <property type="entry name" value="PP-binding"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00975">
        <property type="entry name" value="Thioesterase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00827">
        <property type="entry name" value="PKS_AT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00826">
        <property type="entry name" value="PKS_DH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00829">
        <property type="entry name" value="PKS_ER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00822">
        <property type="entry name" value="PKS_KR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00825">
        <property type="entry name" value="PKS_KS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00823">
        <property type="entry name" value="PKS_PP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47336">
        <property type="entry name" value="ACP-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53474">
        <property type="entry name" value="alpha/beta-Hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52151">
        <property type="entry name" value="FabD/lysophospholipase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50129">
        <property type="entry name" value="GroES-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51735">
        <property type="entry name" value="NAD(P)-binding Rossmann-fold domains"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55048">
        <property type="entry name" value="Probable ACP-binding domain of malonyl-CoA ACP transacylase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53335">
        <property type="entry name" value="S-adenosyl-L-methionine-dependent methyltransferases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53901">
        <property type="entry name" value="Thiolase-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50075">
        <property type="entry name" value="CARRIER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00606">
        <property type="entry name" value="KS3_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS52004">
        <property type="entry name" value="KS3_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00012">
        <property type="entry name" value="PHOSPHOPANTETHEINE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS52019">
        <property type="entry name" value="PKS_MFAS_DH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="20" id="KW-0007">Acetylation</keyword>
    <keyword evidence="24" id="KW-0275">Fatty acid biosynthesis</keyword>
    <keyword evidence="17" id="KW-0276">Fatty acid metabolism</keyword>
    <keyword evidence="16" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="12" id="KW-0444">Lipid biosynthesis</keyword>
    <keyword evidence="23" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="25" id="KW-0456">Lyase</keyword>
    <keyword evidence="26" id="KW-0511">Multifunctional enzyme</keyword>
    <keyword evidence="22" id="KW-0520">NAD</keyword>
    <keyword evidence="18" id="KW-0521">NADP</keyword>
    <keyword evidence="21" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="11" id="KW-0596">Phosphopantetheine</keyword>
    <keyword evidence="13" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="19" id="KW-0663">Pyridoxal phosphate</keyword>
    <keyword evidence="74" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="15" id="KW-0702">S-nitrosylation</keyword>
    <keyword evidence="14" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Ketosynthase family 3 (KS3)" evidence="71">
        <location>
            <begin position="1"/>
            <end position="275"/>
        </location>
    </feature>
    <feature type="domain" description="PKS/mFAS DH" evidence="72">
        <location>
            <begin position="713"/>
            <end position="980"/>
        </location>
    </feature>
    <feature type="domain" description="Carrier" evidence="70">
        <location>
            <begin position="1993"/>
            <end position="2070"/>
        </location>
    </feature>
    <feature type="region of interest" description="N-terminal hotdog fold" evidence="69">
        <location>
            <begin position="713"/>
            <end position="836"/>
        </location>
    </feature>
    <feature type="region of interest" description="C-terminal hotdog fold" evidence="69">
        <location>
            <begin position="853"/>
            <end position="980"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor; for dehydratase activity" evidence="69">
        <location>
            <position position="747"/>
        </location>
    </feature>
    <feature type="active site" description="Proton donor; for dehydratase activity" evidence="69">
        <location>
            <position position="903"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00005194"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00012004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012480"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00012873"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012948"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00013167"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00013191"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00013256"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00013258"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00018769"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022450"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022516"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022799"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022832"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00022857"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00022898"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00023027"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00023160"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00023239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00023268"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="ARBA" id="ARBA00023332"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="ARBA" id="ARBA00023351"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="ARBA" id="ARBA00023373"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="ARBA" id="ARBA00023388"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="31">
        <source>
            <dbReference type="ARBA" id="ARBA00023394"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="32">
        <source>
            <dbReference type="ARBA" id="ARBA00023398"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="33">
        <source>
            <dbReference type="ARBA" id="ARBA00023399"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="34">
        <source>
            <dbReference type="ARBA" id="ARBA00023401"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="35">
        <source>
            <dbReference type="ARBA" id="ARBA00023402"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="36">
        <source>
            <dbReference type="ARBA" id="ARBA00023442"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="37">
        <source>
            <dbReference type="ARBA" id="ARBA00044883"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="38">
        <source>
            <dbReference type="ARBA" id="ARBA00047300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="39">
        <source>
            <dbReference type="ARBA" id="ARBA00047394"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="40">
        <source>
            <dbReference type="ARBA" id="ARBA00047400"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="41">
        <source>
            <dbReference type="ARBA" id="ARBA00047440"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="42">
        <source>
            <dbReference type="ARBA" id="ARBA00047451"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="43">
        <source>
            <dbReference type="ARBA" id="ARBA00047500"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="44">
        <source>
            <dbReference type="ARBA" id="ARBA00047578"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="45">
        <source>
            <dbReference type="ARBA" id="ARBA00047810"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="46">
        <source>
            <dbReference type="ARBA" id="ARBA00047897"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="47">
        <source>
            <dbReference type="ARBA" id="ARBA00047953"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="48">
        <source>
            <dbReference type="ARBA" id="ARBA00047961"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="49">
        <source>
            <dbReference type="ARBA" id="ARBA00048051"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="50">
        <source>
            <dbReference type="ARBA" id="ARBA00048281"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="51">
        <source>
            <dbReference type="ARBA" id="ARBA00048289"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="52">
        <source>
            <dbReference type="ARBA" id="ARBA00048404"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="53">
        <source>
            <dbReference type="ARBA" id="ARBA00048420"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="54">
        <source>
            <dbReference type="ARBA" id="ARBA00048506"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="55">
        <source>
            <dbReference type="ARBA" id="ARBA00048571"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="56">
        <source>
            <dbReference type="ARBA" id="ARBA00048650"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="57">
        <source>
            <dbReference type="ARBA" id="ARBA00048691"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="58">
        <source>
            <dbReference type="ARBA" id="ARBA00048704"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="59">
        <source>
            <dbReference type="ARBA" id="ARBA00048935"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="60">
        <source>
            <dbReference type="ARBA" id="ARBA00049019"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="61">
        <source>
            <dbReference type="ARBA" id="ARBA00049109"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="62">
        <source>
            <dbReference type="ARBA" id="ARBA00049171"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="63">
        <source>
            <dbReference type="ARBA" id="ARBA00049263"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="64">
        <source>
            <dbReference type="ARBA" id="ARBA00049414"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="65">
        <source>
            <dbReference type="ARBA" id="ARBA00049422"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="66">
        <source>
            <dbReference type="ARBA" id="ARBA00049449"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="67">
        <source>
            <dbReference type="ARBA" id="ARBA00049521"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="68">
        <source>
            <dbReference type="ARBA" id="ARBA00049533"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="69">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01363"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="70">
        <source>
            <dbReference type="PROSITE" id="PS50075"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="71">
        <source>
            <dbReference type="PROSITE" id="PS52004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="72">
        <source>
            <dbReference type="PROSITE" id="PS52019"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="73">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016031.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="74">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2382" mass="261733" checksum="8F4D24BE79507110" modified="2019-05-08" version="1">MTGCQRAMFANRISYFFDLKGPSMAIDTACSSSLIALENAYKAICHGECNAALVGGVNLLLKPSTSVQFMKLGMLSPDGACKSFDASGNGYCRSEAVVVILMTKKSMAKRIYATIVNAGSNTDGFKEQGVTFPSGEMQQALVSSLYKESGIQPEEVEYVEAHGTGTKIGDPQELNGIANIFCQLEREPLLIGSTKSNMGHPEPASGLAALAKVILSLEHGLWAPNIHYNTPNPEIPALQDGRVQVVAKPTPVKGGLVSINSFGFGGSNVHVILRPNKKKFYPLERHNLPRLVQVCGRTQEAVEMLIEQSRRLREYSSFVSLLNDISGIPISSMPYRGYTLIGSQSDIKEVQHTRTSGRPLWYICSGMGTQWKGMGLSLMKLELFQQSILRSDEALKDTGLKVSDLLLNADENTFEDAVHAFVGLAAIQIAQIDILKAMDLQPDGIVGHSVGELACGYADNSLSHEEAILAAYWRGRCIKEAKLPPGGMAAVGLTWEECKRHCPPGVVPACHNSEDTVTVSGPQDAVNKFVAKLKKEGVFAKEVRSAGVAFHSYYMASIAPVLLSALQKVIPNAKPRSPRWISTSIPESQWQSELAQNSSAEYHVNNLVSPVLFQEGLRHVPDNAVVVEIAPHALLQAVLRRSLKPTCTILPLMKREHKNNLEFFLTHVGKIYLTGINVLSNNLFPLVEYPAPVGTPLISPYILWDHSQTWDVPKAEDFPTGSGGSSSASIYNIDMNSESSDNYLVGHCIDGRVLYPATGYLVLAWRTLARSLGIAMEQMPVVFEEVTIHQATILSNKGSTQLEVRLLPASHCFEVSGNGNLAVSGKICLLEETALNNFRNQQVDLETPPNMRSKFSLSKADIYKELRLRGYDYGPTFQGLLESNNEGNRGKVLWNGNWVTFLDTLLQMIILGETGHSLRLPTRIRSVCIDPSLHQERVHKYKDDTEAFDVVVDRCLDNIMAGGVRLSGLHATVAPRRQQEQRPPTLEKFCFVPYVESDCLSSEAHLCAHLEHCKGLICKLQEKVAFHGVKLAIPGLETVEAFGETVSMQKGLLHILAEICHLELNGNLYSELGQIVTREKMHLQDDPLLNALLDSPELKACVDIALENITSHRMKIVEALAGDGRLFSRVTGLLNTQPMLQLDYTATDQAPEILSVNESQLQETGVSIAQWDPISPPPGHLTSADLLVCNCSVSIFGNPAEILSNLAATVKEGGFILLHTLLKGETLGEIISFLTTPDLQQRGLLAQAEWEDLFRKTLLNLVAVKKSFFGSVIFLCRRQASAKIPVFLPVEETHYKWVESLKVILADASEQPVWLTATSCGNSGIVGMVNCLRQEPGGHRIRCLFVSNLNSSLAIPPTSPSAREMQKILQGDLVMNVYRDGKWGSFRHFLLQQAQPQELTDYAYVNVLTRGDLSSLRWISSPLRHFHTTNPNIQLCRVYYASLNFRDIMLATGKLPPDAIPGNWTLQQCMLGMEFSGRDPTGKRVMGLLPAKGLATVVDSDKRFLWEVPENWTLEEAASVPVVYATAYYALVVRGGMKKGESVLIHSGSGGVGQAAIAIALSMGCRVFATVGSVEKREYLQTRFPQLDANSFANSRNTSFEQHILRVTNGKGVNLVLNSLAEEKLQASLRCLARHGRFLEIGKYDLSNNSPLGMALFLKNVAFHGILLDAIFEEGNQEWEEVSDLLKMGIRDGVVKPLRTTIFNREEVEAAFRFMAQGKHIGKVMIKVHEEEKEFPARETEAVRLSAISRTSCPPTKSYIITGGLGGFGLELAQWLVERGAQKLVLTSRSGIRSGYQAKRVRGWKEMGVQVLVSTSDVGTLEGTQHLIEEASLLGPVGGIFNLAMVLKDAMLENQSPEFFHDVSKPKYAGTLHLDWVSREKCPDLDYFVAFSSISCGRGNVGQSNYGFANSTMERICEQRQHDGLPGLAVQWGAIGDVGVVLETMGSNDTVVGGTLPQRLSSCLEVLDHFLNQSHPVMSSFVLAEKVSVKSEGSSQRDLMEAVAHILGVRDVSSLNAESSLADLGLDSLMGVEVRQTLERDYDLVMTMREIRLLTINKLRELSSKSGTAEELKPSLLMKAGQAEHPELDLNNLLVNPEGPTITRLNEVQSTERPLFLIHPIEGSITVFHTLASRLSMPSYGLQCTKAAPLDSIQSLAAYYIDCIKQVQPEGPYRIAGYSFGACVAFEMCSQLQAQQNPSHSLNSLFLFDGSHSYVAAYTQSYRAKLTPGNEAEAETEALCAFIQQFTSTEYNKLLETLLPLQDLEARVNAAVDLIMLSHKNINRETLCFAAASFYYKLKAADKYVPESKYHGNVTLLRAKTHNEYEEGLGGDYKLSEVCEGKVSVHVIEGDHRTFLEGAGVESIIGIIHSSLAEPRVSVREG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM65</accession>
    <name>A0A452HM65_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Collagen alpha-1(XXI) chain</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Secreted</location>
            <location evidence="2">Extracellular space</location>
            <location evidence="2">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="11">Belongs to the fibril-associated collagens with interrupted helices (FACIT) family.</text>
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    <dbReference type="AlphaFoldDB" id="A0A452HM65"/>
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        <property type="project" value="UniProtKB-KW"/>
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        <property type="entry name" value="Collagen alpha-1(XXI) chain isoform X1"/>
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        <property type="entry name" value="ECM_Assembly_Org"/>
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        <property type="entry name" value="TSPN-like_N"/>
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        <property type="entry name" value="VWF_A"/>
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    <dbReference type="InterPro" id="IPR036465">
        <property type="entry name" value="vWFA_dom_sf"/>
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        <property type="entry name" value="COLLAGEN ALPHA"/>
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    </dbReference>
    <dbReference type="PANTHER" id="PTHR24020:SF90">
        <property type="entry name" value="COLLAGEN ALPHA-1(XXI) CHAIN"/>
        <property type="match status" value="1"/>
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    <dbReference type="Pfam" id="PF00092">
        <property type="entry name" value="VWA"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="VWFADOMAIN"/>
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        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
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        <property type="entry name" value="VWFA"/>
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    <keyword evidence="5" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="9" id="KW-0325">Glycoprotein</keyword>
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    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="6 13" id="KW-0732">Signal</keyword>
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        <source>
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    </evidence>
    <sequence length="437" mass="49415" checksum="5F7E6ED0579F5661" modified="2019-05-08" version="1">MAQNLKVLQTILILLLQDYISAEDGEIRASCRTAPADLVFILDGSYSVGPENFEIIKRWLVNITRNFDIGPKFIQVGVVQYSDYPVLEIPLGAYDSTENLIREMESIQYLGGNTRTGRAIQFSIDHLFAKSSRFLTKIAVVLTDGKSQDEVKDAAAEARKNKITLFAIGVGSEIEEDELRAIANKPSSTYVFYVEDYIAISRIREVIKQKLCEESVCPTRIPVAARDEKGFDILLGLGVKKKVKKRIQIPSSNAKAYEITSNIDLSEFTRNVFPEGLPPSYVFVSTQRFKVRKTWDLWRILSLDRRPQITVTVNGEDKTLSFTTTSLVNGTQVVTFTAPRVKILFDEGWHQIRLLVTEAHVTLYIDDQETEMKPLHPVLGIYISGLTQIGKYPGKEETVQFDVQKLRIYCDPEQNNRETACEIPGFVSMIDELSRIV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM66</accession>
    <name>A0A452HM66_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="10">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
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        <property type="gene ID" value="ENSGAGG00000012051.1"/>
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        <property type="entry name" value="Matrin/U1-C_Znf_C2H2"/>
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        <location>
            <begin position="2354"/>
            <end position="2363"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="7">
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            <begin position="2447"/>
            <end position="2461"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00022771"/>
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            <dbReference type="ARBA" id="ARBA00022833"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
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    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00176"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50102"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50171"/>
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    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016040.1"/>
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    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2461" mass="270371" checksum="80015CA566AD4579" modified="2019-05-08" version="1">MQPGDRVVASQGVISKGKRAKSVELPSQLPLDPLRRFVLFLESFAPFVSSLNLGIVSPLLLGPPPLQLAQIKTQLALQQLTSAASNNSTPPYALLNQAFLKIAMFNPRGNLHQRPRAPNPSGSKPPGAFRGGGIAGLQRKPAPGTPQGISQRFGGHENLQRTGSRRTNVQVTQHRTDPRQAKEMADLQEEQKEKVRTSRWDNNPCSTGNTSRKHPVSTRITEQNTNVQSRYTTESASSILASFGLSNEDLEELSRYPDDQLTPENMPLILRDIRMRKMSHQLPNLHSQSREKETVGGTGGSTVKSKVIDYGHTSRYGYTEDPLEVRVYNPEEPAEENRKEFQPQQTISVPSSNVTCNPVFPVEELMKQMGFHSDSSNTQSFFPVDATGKVPGLCMTPTGLPMVKPMSQPGMPPMMPPMSQPGMPPMMPPMSQPAMPPMMPPVMPPVPQSVMTPIVQQSLTQPPMAQPVMPPISHPPFSAELLAAVRHHERIQHESGTNQSNAQIGSGQKNFQTQVEAPIKSPFGIVKASWLPVFSQAAAQKVKRLPTPSMMNDYYATSPRIFPHMCSLCNMECTHMKDWIQHQNNPAHIESCRQLRQQYPDWNPEVHSSKRNEGERKENQTPKRRSNSASPGPRHSRGSGSGHILRRSRSRSRSPGRFRLPRPRSRSPRQMRRPSPRHRSRSPQRSRNPLRCSPRPQRSASNEWASRRTTRSPDKKAALEAVVKSLGPSFVAEFNKQKSFQAAGQGSSGIGKASPAQGISGVGKIHGSVKKPPSATGHHKTLKKDLSSVSGSAASKMKSETAGVQEKKEMNFDGKAAKETNVPRPAPYNRLLREKLLSCGTVLHISDLPDDGFSDQDIKKIVQPFGKVSDLLVLRSRNEAFLEMNYKEAVIAAVKYGETVPVLVNGKRVKISIAEKPKAPPGQVKVSLKKTPQNVKKATLSTKKDGNSTTTKKTKSATSAASAGKHEVTKKPAVQTDSKTKKTSTTSAKPNEKKRIDPKKSAESKKKDAKPKKVAEPKRAVETKTTIESKKAVEARKTTEPKKAGDDEAKEISKPIAVQEKITLTSTLEVESNQSMESVANKENLEAKNTVEVKTAESTRKENASKATGETTKSEEPNDPTSKELDDMCVVLISNLPDKGYSVEEVSNLTKPFGGLKDVLILSSHKKAYLEINQKSADSMVKFYTCFPMSVDGNQLCISMAPEYKNIKDEEAIFTAIIKDANSKVNTETLYNQFVHLGNLPEDGYTELEVVCVGLRFGKVDHYVVLKNKKKAILQLDSPDSAKSMCCFLKQYPYNMGENTLTCMLSPRRELAEDEVTKKELKNQEPSKGSSDLKKKPEGSGVVQTAAANPPVETSVAKEEIISNSVKAETSTVQIEASELEPERTVRDSATKPSEVETSKGESEVAVTGSETKPADDELSKVKSEEILLPPFNLQKEEVVKDNSETELLESAITTVTERETKVKPEELRVPASGSSEELSSVGKLEDVQSDSAVKPATVETAEAVSEAVPPDSAAVSVEMLSAEDIQPSNKLDTPVKNAVTGTVENKLEIPVASAVVSIEKKPEKSVTKVGTALEKKLEKSMAKIGTAMEKKLEKPVASTGVDTEKKPEKPVDNVETDVEKKPEKPVDNVGMDVEKKPEKPVDNVITDIENIEKNVLNEKNERTLIKAHQNKGPGQVKIDETSKASVSAAAFVSIKEPTSVGKTILKAVVSIPDISKSRVMMRRNETSLTKTEEQKASSKPETRSRSATEKKLSSKEVGQPRATGSRSGLPDSVTKSKLDISPVLVKGGSGRSSSQQDKDSRVESRGSSKQSQERESRSSSMKKDDNSNKIPAGRISKTSKTVSSGNVKPKEEEELFPFNLDEFVTVDEVVEETDSPLQTRRNPPRGKRKDSAKNNSSSEPVTKRKKGKSSIAHVDESELSFVTLDEIVEEEEMTAQLVGDSNLEAITDPQSLVTVDEVNEEEELISEAVKDPQSLVTLDEISEQEEFSSHDLPKDVSSSVSEEQDLLKAEPLVTVDEIGEVEELPLNEPSDLNIEETLKPKEEDDKRAIEDPGDFISSQLPEDPSTLVTVDEIQEDSDDQPLVTLDEVTEDDEDFLADFNRLKEELNFVTVDEVGEEDEEEENTFIGTNLEEEDIVAIAGPEEMEILADIGREGEDIFANSKSEEKVTLEADTKEIENETLVADKQEADEALSANTEETEKAETWNKTSEKEIKGETESEQQSAELSLSKAHESNEMGRQTKNKHQPPVKTPGKRGRPRKRPIVPSPETPIETEQEKCDKNTEETADKTELVESSTSDSNKEKIGKGTLTTPESNSLSEDSQTTTVTDSVSSSLLPDINASLLPDADIKQEGCSETPSQSSEKPVKDELGSGDTEPESKRRKVDSSADKLKTPSTPKGLDFLVPKAGYFCQICSLFYAGEMSMKNHCKTLRHQQNMEKFMAKQKDGEDEEEDEGEELSSR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM76</accession>
    <name>A0A452HM76_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="10">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018330.1">
        <property type="protein sequence ID" value="ENSGAGP00000016047.1"/>
        <property type="gene ID" value="ENSGAGG00000012051.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008380">
        <property type="term" value="P:RNA splicing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd12716">
        <property type="entry name" value="RRM1_2_NP220"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000690">
        <property type="entry name" value="Matrin/U1-C_Znf_C2H2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003604">
        <property type="entry name" value="Matrin/U1-like-C_Znf_C2H2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012677">
        <property type="entry name" value="Nucleotide-bd_a/b_plait_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035979">
        <property type="entry name" value="RBD_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000504">
        <property type="entry name" value="RRM_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033096">
        <property type="entry name" value="ZNF638_RRM1/2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15592">
        <property type="entry name" value="MATRIN 3/NUCLEAR PROTEIN 220-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00360">
        <property type="entry name" value="RRM"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00451">
        <property type="entry name" value="ZnF_U1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54928">
        <property type="entry name" value="RNA-binding domain, RBD"/>
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    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="5" id="KW-0539">Nucleus</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="4" id="KW-0862">Zinc</keyword>
    <keyword evidence="3" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="RRM" evidence="8">
        <location>
            <begin position="841"/>
            <end position="916"/>
        </location>
    </feature>
    <feature type="domain" description="Matrin-type" evidence="9">
        <location>
            <begin position="2402"/>
            <end position="2432"/>
        </location>
    </feature>
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            <begin position="109"/>
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        </location>
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            <begin position="284"/>
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        <source>
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            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2454" mass="269354" checksum="69C55980DE164427" modified="2019-05-08" version="1">MQPGDRVVASQGVISKGKRAKSVELPSQLPLDPLRRFVLFLESFAPFVSSLNLGIVSPLLLGPPPLQLAQIKTQLALQQLTSAASNNSTPPYALLNQAFLKIAMFNPRGNLHQRPRAPNPSGSKPPGAFRGGGIAGLQRKPAPGTPQGISQRFGGHENLQRTGSRRTNVQVTQHRTDPRQAKEMADLQEEQKEKVRTSRWDNNPCSTGNTSRKHPVSTRITEQNTNVQSRYTTESASSILASFGLSNEDLEELSRYPDDQLTPENMPLILRDIRMRKMSHQLPNLHSQSREKETVGGTGGSTVKSKVIDYGHTSRYGYTEDPLEVRVYNPEEPAEENRKEFQPQQTISVPSSNVTCNPVFPVEELMKQMGFHSDSSNTQSFFPVDATGKVPGLCMTPTGLPMVKPMSQPGMPPMMPPMSQPGMPPMMPPMSQPAMPPMMPPVMPPVPQSVMTPIVQQSLTQPPMAQPVMPPISHPPFSAELLAAVRHHERIQHESGTNQSNAQIGSGQKNFQTQVEAPIKSPFGIVKASWLPVFSQAAAQKVKRLPTPSMMNDYYATSPRIFPHMCSLCNMECTHMKDWIQHQNNPAHIESCRQLRQQYPDWNPEVHSSKRNEGERKENQTPKRRSNSASPGPRHSRGSGSGHILRRSRSRSRSPGRFRLPRPRSRSPRQMRRPSPRHRSRSPQRSRNPLRCSPRPQRSASNEWASRRTTRSPDKKAALEAVVKSLGPSFVAEFNKQKSFQAAGQGSSGIGKASPAQGISGVGKIHGSVKKPPSATGHHKTLKKDLSSVSGSAASKMKSETAGVQEKKEMNFDGKAAKETNVPRPAPYNRLLREKLLSCGTVLHISDLPDDGFSDQDIKKIVQPFGKVSDLLVLRSRNEAFLEMNYKEAVIAAVKYGETVPVLVNGKRVKISIAEKPKAPPGQVKVSLKKTPQNVKKATLSTKKDGNSTTTKKTKSATSAASAAKLTKAGQTTTKAVKLAGKHEVTKKPAVQTDSKTKKTSTTSAKPNEKKRIDPKKSAESKKKDAKPKKVAEPKRAVETKTTIESKKAVEARKTTEPKKAGDDEAKEISKPIAVQEKITLTSTLEVESNQSMESVANKENLEAKNTVEVKTAESTRKENASKATGETTKSEEPNDPTSKELDDMCVVLISNLPDKGYSVEEVSNLTKPFGGLKDVLILSSHKKAYLEINQKSADSMVKFYTCFPMSVDGNQLCISMAPEYKNIKDEEAIFTAIIKDANSKVNTETLYNQFVHLGNLPEDGYTELEVVCVGLRFGKVDHYVVLKNKKKAILQLDSPDSAKSMCCFLKQYPYNMGENTLTCMLSPRRELAEDEVTKKELKNQEPSKGSSDLKKKPEGSGVVQTAAANPPVETSVAKEEIISNSVKAETSTVQIEASELEPERTVRDSATKPSEVETSKGESEVAVTGSETKPADDELSKVKSEEILLPPFNLQKEEVVKDNSETELLESAITTVTERETKVKPEELRVPASGSSEELSSVGKLEDVQSDSAVKPATVETAEAVSEAVPPDSAAVSVEMLSAEDIQPSNKLDTPVKNAVTGTVENKLEIPVASAVVSIEKKPEKSVTKVGTALEKKLEKSMAKIGTAMEKKLEKPVASTGVDTEKKPEKPVDNVETDVEKKPEKPVDNVGMDVEKKPEKPVDNVITDIENIEKNVLNEKNERTLIKAHQNKGPGQVKIDETSKASVSAAAFVSIKEPTSVGKTILKAVVSIPDISKSRVMMRRNETSLTKTEEQKASSKPETRSRSATEKKLSSKEVGQPRATGSRSGLPDSVTKSKLDISPVLVKGGSGRSSSQQDKDSRVESRGSSKQSQERESRSSSMKKDDNSNKIPAGRISKTSKTVSSGNVKPKEEEELFPFNLDEFVTVDEVVEETDSPLQTRRNPPRGKRKDSAKNNSSSEPVTKRKKGKSSIAHVDESELSFVTLDEIVEEEEMTAQLVGDSNLEAITDPQSLVTVDEVNEEEELISEAVKDPQSLVTLDEISEQEEFSSHDLPKDVSSSVSEEQDLLKAEPLVTVDEIGEVEELPLNEPSDLNIEETLKPKEEDDKRAIEDPGDFISSQLPEDPSTLVTVDEIQEDSDDQPLVTLDEVTEDDEDFLADFNRLKEELNFVTVDEVGEEDEEEENTFIGTNLEEEDIVAIAGPEEMEILADIGREGEDIFANSKSEEKVTLEADTKEIENETLVADKQEADEALSANTEETEKAETWNKTSEKEIKGETESEQQSAVKTPGKRGRPRKRPIVPSPETPIETEQEKCDKNTEETADKTELVESSTSDSNKEKIGKGTLTTPESNSLSEDSQTTTVTDSVSSSLLPDINASLLPDADIKQEGCSETPSQSSEKPVKDELGSGDTEPESKRRKVDSSADKLKTPSTPKGLDFLVPKAGYFCQICSLFYAGEMSMKNHCKTLRHQQNMEKFMAKQKDGEDEEEDEGEELSSR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM78</accession>
    <name>A0A452HM78_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Fibronectin type-III domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the type I cytokine receptor family. Type 2 subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HM78"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018320.1">
        <property type="protein sequence ID" value="ENSGAGP00000016037.1"/>
        <property type="gene ID" value="ENSGAGG00000011475.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009897">
        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004896">
        <property type="term" value="F:cytokine receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000494">
        <property type="entry name" value="Leptin receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000501">
        <property type="entry name" value="Leptin receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000568">
        <property type="entry name" value="Leptin receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000613">
        <property type="entry name" value="Leptin receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003529">
        <property type="entry name" value="Hematopoietin_rcpt_Gp130_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010457">
        <property type="entry name" value="IgC2-like_lig-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041182">
        <property type="entry name" value="LEP-R_IGD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23037">
        <property type="entry name" value="CYTOKINE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23037:SF44">
        <property type="entry name" value="LEPTIN RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06328">
        <property type="entry name" value="Lep_receptor_Ig"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18589">
        <property type="entry name" value="ObR_Ig"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01353">
        <property type="entry name" value="HEMATOPO_REC_L_F2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7 12" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-0675">Receptor</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="3 12" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 12" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="823"/>
            <end position="845"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="13">
        <location>
            <begin position="222"/>
            <end position="316"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="13">
        <location>
            <begin position="523"/>
            <end position="619"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="13">
        <location>
            <begin position="724"/>
            <end position="817"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="940"/>
            <end position="959"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="940"/>
            <end position="949"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="950"/>
            <end position="959"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008921"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016037.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1147" mass="130360" checksum="173E3D5104E6F7D2" modified="2019-05-08" version="1">VYHQNFMQMAVAHCMVHQIPASSFMLPCMSPNETSVIPLTAGVVASSPGLHREHGTTETNSLMLVAEESFLCCLWSETNANCSLYTAGMETKKFTSSEVSSFAPQQTDLSWNIQCWTKGDLELFVCILKLSNKKSYLNGEFKINLLYVLSELSLEDISTGSLKGNFMVTPCNGNGSDKYECHIPSLKLNYTYITWLNIINGVTLLQSPLMSVRPINIVKPESPLNLRLEMTDKGQLKICWSNPVLTPYPLQYEVKFSGNTTQNAWQVVEIVIETSLIIGNMLVGSSYLVQVRCKSLHGTGFWSDWSTPYNLNTEDVMYFPPKILTSVGSNVSFHCLYNDKNKMILSKKIVWWLNLAEEIPERQYTLVNDRVSRVTLFNLNATKPRGKFFYNALYCCNQNRECHHRYAELYVVDVNINITCETDGYLTKMTCRWSANTNTLLVGSSLQLKYYRSSIYCSDFPSIPPKSEAKECHLQRNHYYECTFQPIFLLSGYTMWIEIKHQLGTLESPPTCVIPADVVKPFPPSNVKAEITKNIGLLNVSWTNPAFPNSDLKFQIRYSVNREEILWEIFEVSNAPTRSVMIKVLDLCVVYIVQVRCSGLDGLGYWSDWSKPAYTIVQDIQAPLRGPEFWRVINEDPIRKQKNVTLVWKPLMKNYSLCSVCGYIVKQHTSENITWTEYVTSGTTYTYPWMEDAHTITILAVNSVGASSMNFNLTLSQQMSTVNIVQSLSAYPVNSSCVILTWTLSPQMYVITSFVIEWKNLNEEEQMKWIRVAPNISKYYIYDHFILIEKYQFSLYPIFPEGVGNPKTTDEFIKDRSEKQNDAGLYVILPIVISSFVLLLGTLLISQQRMKKLFWEDVPNPKNCSWAQGVNFQKPETFEHLFIKHPEAISFGPLLLEPEIVLEDISVAKALKSEDKQDLLAVDSMFTMIQDSEHDSACSSSHFSNSLSESSHDDKVSERTMRQSNIKYATIISNCKLSGLYEEQKNLSGSFSGCFLGEDSLVIDPFSRSWEVGNQAFLILPDQHPSQASKTISLSVVSSEGFSEPSDHDDTFSDGDSPERSLYYLGLTSVKKNENDIFLTENSRVMCHLHTNGLFKDMGFLQDTSSDLNPFIKNSLKYENSVKTFVPYMPQFQATTIKLQETTETKT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM79</accession>
    <name>A0A452HM79_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="10">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018337.1">
        <property type="protein sequence ID" value="ENSGAGP00000016053.1"/>
        <property type="gene ID" value="ENSGAGG00000012051.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008380">
        <property type="term" value="P:RNA splicing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd12716">
        <property type="entry name" value="RRM1_2_NP220"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.330">
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000690">
        <property type="entry name" value="Matrin/U1-C_Znf_C2H2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003604">
        <property type="entry name" value="Matrin/U1-like-C_Znf_C2H2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012677">
        <property type="entry name" value="Nucleotide-bd_a/b_plait_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035979">
        <property type="entry name" value="RBD_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000504">
        <property type="entry name" value="RRM_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033096">
        <property type="entry name" value="ZNF638_RRM1/2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15592">
        <property type="entry name" value="MATRIN 3/NUCLEAR PROTEIN 220-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00360">
        <property type="entry name" value="RRM"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00451">
        <property type="entry name" value="ZnF_U1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54928">
        <property type="entry name" value="RNA-binding domain, RBD"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50102">
        <property type="entry name" value="RRM"/>
        <property type="match status" value="1"/>
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</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM80</accession>
    <name>A0A452HM80_9SAUR</name>
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            <fullName evidence="11">Cysteine rich transmembrane BMP regulator 1</fullName>
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        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
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        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
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        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
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        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
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        <citation type="submission" date="2025-08" db="UniProtKB">
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        <scope>IDENTIFICATION</scope>
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        <citation type="submission" date="2025-09" db="UniProtKB">
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        <scope>IDENTIFICATION</scope>
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    <keyword evidence="3" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="4" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <keyword evidence="1 5" id="KW-0732">Signal</keyword>
    <keyword evidence="4" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="5">
        <location>
            <begin position="1"/>
            <end position="37"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019220518" description="Cysteine rich transmembrane BMP regulator 1" evidence="5">
        <location>
            <begin position="38"/>
            <end position="976"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="4">
        <location>
            <begin position="881"/>
            <end position="901"/>
        </location>
    </feature>
    <feature type="domain" description="IGFBP N-terminal" evidence="8">
        <location>
            <begin position="36"/>
            <end position="115"/>
        </location>
    </feature>
    <feature type="domain" description="VWFC" evidence="6">
        <location>
            <begin position="342"/>
            <end position="398"/>
        </location>
    </feature>
    <feature type="domain" description="Antistasin-like" evidence="7">
        <location>
            <begin position="410"/>
            <end position="439"/>
        </location>
    </feature>
    <feature type="domain" description="Antistasin-like" evidence="7">
        <location>
            <begin position="446"/>
            <end position="473"/>
        </location>
    </feature>
    <feature type="domain" description="Antistasin-like" evidence="7">
        <location>
            <begin position="480"/>
            <end position="505"/>
        </location>
    </feature>
    <feature type="domain" description="Antistasin-like" evidence="7">
        <location>
            <begin position="508"/>
            <end position="533"/>
        </location>
    </feature>
    <feature type="domain" description="VWFC" evidence="6">
        <location>
            <begin position="547"/>
            <end position="604"/>
        </location>
    </feature>
    <feature type="domain" description="VWFC" evidence="6">
        <location>
            <begin position="618"/>
            <end position="676"/>
        </location>
    </feature>
    <feature type="domain" description="VWFC" evidence="6">
        <location>
            <begin position="692"/>
            <end position="750"/>
        </location>
    </feature>
    <feature type="domain" description="VWFC" evidence="6">
        <location>
            <begin position="758"/>
            <end position="815"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50184"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS51252"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51323"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016093.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="11">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="976" mass="107205" checksum="8A36506003C3257F" modified="2019-05-08" version="1">MYLAAVLAGRRRPGGSGWHLGAGWLLLVLLGESGTRALVCLPCDESKCEEPKSCPGSIVLGICGCCFMCARQRNESCGGVYGLHGACDRGLRCVIRPPLNGDSITEYEVGVCEDENWDDQLLGFEPCNENLVTGCNIINGKCECDTIRTCSNPFEFPSQDTCLSALKRIEEEKPDCSKARCEVQFSPRCPEDSILIEGYAPPGECCPLPSRCVCNPAGCLRKVCQPGYLNILVSKASGKPGECCDLYECKPVFSVDCSTVECPPVQQVVCPLDSYETQVRLTADGCCTLPTRCECLSGLCGFPMCEAGSIPQIVSHGDGTPGKCCDVFECVNDPIHPVNNPAGCYANGQIQAHGDRWREDDCTFCQCINGDPHCVATACGQSCLNPVKVPGECCPVCEEPTYITIGPSTCEILVNCTLTERDCIYSFRLDQNGCRICQCKTREELCTGLISGCSLDCSFGFQTDVHNCEICQCRSRPKKCKPIVCDKYCPFGYLKNKHGCDICRCKKCPELPCGKICPMGFQQNNHGCLLCKCREIPDSVMPPVKTGSCLSMDGRRHEKEESWHDGCRECYCHNGREMCALITCPVPNCGNPTIHPGQCCPSCPDESIVQKPELSNPSICHAPGGEYFVEGETWNIDSCTQCTCHSGRVLCETEVCPPLLCQNPIRTQDSCCPQCPDEPLQPSSSSNESMPSYCKNDEGDIFLTAESWKPNVCTSCICMDGVISCYSESCPPVSCERPVLRKGQCCPYCIEDTIPKKVVCHFSGKTYADEERWDIDSCTHCYCLQGQTLCSTVSCPPLPCVEPINVEGSCCPMCPEMYVPEPTNIPIEKTNHRDAELEAPIWPTPSENDIVHLHRDMSHLQGDFREGIGLHSSEDTSLNSVAWVTVPIMTALVLIIALLLVNQKKQWIPVSCYRAPTKPSCLNNQLVYVDCKKGTMVQVDSSQRMLRIADPDSRYSGFYSMQKQNNLQADNFYQTV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM85</accession>
    <name>A0A452HM85_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Receptor-type tyrosine-protein phosphatase</fullName>
            <ecNumber evidence="16">3.1.3.48</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">Tyrosine protein phosphatase which is involved in integrin-mediated focal adhesion formation. Following integrin engagement, specifically recruits BCAR3, BCAR1 and CRK to focal adhesions thereby promoting SRC-mediated phosphorylation of BRAC1 and the subsequent activation of PAK and small GTPase RAC1 and CDC42.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="16">
            <text>O-phospho-L-tyrosyl-[protein] + H2O = L-tyrosyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:10684"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="EC" id="3.1.3.48"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell junction</location>
            <location evidence="1">Focal adhesion</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 16">Belongs to the protein-tyrosine phosphatase family. Receptor class 4 subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.48" evidence="16"/>
    <dbReference type="AlphaFoldDB" id="A0A452HM85"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018342.1">
        <property type="protein sequence ID" value="ENSGAGP00000016058.1"/>
        <property type="gene ID" value="ENSGAGG00000010212.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005925">
        <property type="term" value="C:focal adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="CDD" id="cd14621">
        <property type="entry name" value="R-PTPc-A-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd14623">
        <property type="entry name" value="R-PTPc-A-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000007">
        <property type="entry name" value="Receptor-type tyrosine-protein phosphatase alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000021">
        <property type="entry name" value="Receptor-type tyrosine-protein phosphatase alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050348">
        <property type="entry name" value="Protein-Tyr_Phosphatase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000242">
        <property type="entry name" value="PTP_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016130">
        <property type="entry name" value="Tyr_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003595">
        <property type="entry name" value="Tyr_Pase_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016336">
        <property type="entry name" value="Tyr_Pase_rcpt_a/e-type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19134">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19134:SF433">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00102">
        <property type="entry name" value="Y_phosphatase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF002006">
        <property type="entry name" value="PTPR_alpha_epsilon"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00700">
        <property type="entry name" value="PRTYPHPHTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00194">
        <property type="entry name" value="PTPc"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00404">
        <property type="entry name" value="PTPc_motif"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00383">
        <property type="entry name" value="TYR_PHOSPHATASE_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50055">
        <property type="entry name" value="TYR_PHOSPHATASE_PTP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="11" id="KW-0965">Cell junction</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="14" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9 16" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="13" id="KW-0472">Membrane</keyword>
    <keyword evidence="5 19" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="10 16" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
    <keyword evidence="6" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="12" id="KW-1133">Transmembrane helix</keyword>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="21">
        <location>
            <begin position="130"/>
            <end position="390"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="22">
        <location>
            <begin position="310"/>
            <end position="381"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="21">
        <location>
            <begin position="422"/>
            <end position="680"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="22">
        <location>
            <begin position="596"/>
            <end position="671"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="1"/>
            <end position="53"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="20">
        <location>
            <begin position="1"/>
            <end position="27"/>
        </location>
    </feature>
    <feature type="active site" description="Phosphocysteine intermediate" evidence="17">
        <location>
            <position position="331"/>
        </location>
    </feature>
    <feature type="active site" description="Phosphocysteine intermediate" evidence="17">
        <location>
            <position position="621"/>
        </location>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="299"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <begin position="331"/>
            <end position="337"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="375"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="modified residue" description="Phosphotyrosine" evidence="19">
        <location>
            <position position="687"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007765"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00054255"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PIRNR" id="PIRNR002006"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="PIRSR" id="PIRSR002006-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="PIRSR" id="PIRSR002006-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="PIRSR" id="PIRSR002006-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016058.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="691" mass="79148" checksum="72E550348A32BFD0" modified="2019-05-08" version="1">PCPLSTPPPPNSSPFPGFPHLSNPPRPLTTTPRTSAPYPTPTPFHDAQSSRSSQPCCLAGASHAAWQERWARMLAWRNEHRCNSLCDIAEPQSMPLLARSPSTNRKYPPLPVDKLEEEINRRMADDNKLFREEFNALPACPIQATCEAASKEENKEKNRYVNILPYDHSRVHLTPIEGVPDSDYINASFINGYQEKNKFIAAQGPKEETVNDFWRMIWEQNTATIVMVTNLKERKECKCAQYWPDQGCWTYGNIRVSVEDVTVLVDYTVRKFCIQQVGDVTNKKPQRLVTQFHFTSWPDFGVPFTPIGMLKFLKKVKTCNPQYAGAIVVHCSAGVGRTGTFIVIDAMLDMMHAERKVDVYGFVSRIRAQRCQMVQTDMQYVFIYQALLEHYLYGDTELEVTSLETHLQKIYNKVPGTSSNGLEEEFKKLTSIKIQNDKMRTGNLPANMKKNRVLQIIPYEFNRVIIPVKRGEENTDYVNASFIDGYRQKDSYIASQGPLQHTIEDFWRMIWEWKSCSIVMLTELEERGQEKCAQYWPSDGSVSYGDITMELKKEEECESYTVRDLLVTNTRENKSRQIRQFHFHGWPEVGIPSDGKGMINIIAAVQKQQQQSGNHPITVHCSAGAGRTGTFCALSTVLERVKAEGILDVFQTVKSLRLQRPHMVQTLEQYEFCYKVVQEYIDAFSDYANFK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM86</accession>
    <name>A0A452HM86_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="10">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
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        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
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            <dbReference type="PubMed" id="28562605"/>
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        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
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        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
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        </citation>
        <scope>IDENTIFICATION</scope>
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    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
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                <consortium name="Ensembl"/>
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        </citation>
        <scope>IDENTIFICATION</scope>
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</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM87</accession>
    <name>A0A452HM87_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">ZZ-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
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        <property type="gene ID" value="ENSGAGG00000012063.1"/>
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        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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        <property type="term" value="C:synapse"/>
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<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
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            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
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        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
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                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
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        <scope>IDENTIFICATION</scope>
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        <scope>IDENTIFICATION</scope>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50102"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50171"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016066.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2352" mass="258538" checksum="960364F9D378A50C" modified="2019-05-08" version="1">MFNPRGNLHQRPRAPNPSGSKPPGAFRGGGIAGLQRKPAPGTPQGISQRFGGHENLQRTGSRRTNVQVTQHRTDPRQAKEMADLQEEQKEKVRTSRWDNNPCSTGNTSRKHPVSTRITEQNTNVQSRYTTESASSILASFGLSNEDLEELSRYPDDQLTPENMPLILRDIRMRKMSHQLPNLHSQSREKETVGGTGGSTVKSKVIDYGHTSRYGYTEDPLEVRVYNPEEPAEENRKEFQPQQTISVPSSNVTCNPVFPVEELMKQMGFHSDSSNTQSFFPVDATGKVPGLCMTPTGLPMVKPMSQPGMPPMMPPMSQPGMPPMMPPMSQPAMPPMMPPVMPPVPQSVMTPIVQQSLTQPPMAQPVMPPISHPPFSAELLAAVRHHERIQHESGTNQSNAQIGSGQKNFQTQVEAPIKSPFGIVKASWLPVFSQAAAQKVKRLPTPSMMNDYYATSPRIFPHMCSLCNMECTHMKDWIQHQNNPAHIESCRQLRQQYPDWNPEVHSSKRNEGERKENQTPKRRSNSASPGPRHSRGSGSGHILRRSRSRSRSPGRFRLPRPRSRSPRQMRRPSPRHRSRSPQRSRNPLRCSPRPQRSASNEWASRRTTRSPDKKAALEAVVKSLGPSFVAEFNKQKSFQAAGQGSSGIGKASPAQGISGVGKIHGSVKKPPSATGHHKTLKKDLSSVSGSAASKMKSETAGVQEKKEMNFDGKAAKETNVPRPAPYNRLLREKLLSCGTVLHISDLPDDGFSDQDIKKIVQPFGKVSDLLVLRSRNEAFLEMNYKEAVIAAVKYGETVPVLVNGKRVKISIAEKPKAPPGQVKVSLKKTPQNVKKATLSTKKDGNSTTTKKTKSATSAASAAKLTKAGQTTTKAVKLAGKHEVTKKPAVQTDSKTKKTSTTSAKPNEKKRIDPKKSAESKKKDAKPKKVAEPKRAVETKTTIESKKAVEARKTTEPKKAGDDEAKEISKPIAVQEKITLTSTLEVESNQSMESVANKEATGETTKSEEPNDPTSKELDDMCVVLISNLPDKGYSVEEVSNLTKPFGGLKDVLILSSHKKAYLEINQKSADSMVKFYTCFPMSVDGNQLCISMAPEYKNIKDEEAIFTAIIKDANSKVNTETLYNQFVHLGNLPEDGYTELEVVCVGLRFGKVDHYVVLKNKKKAILQLDSPDSAKSMCCFLKQYPYNMGENTLTCMLSPRRELAEDEVTKKELKNQEPSKGSSDLKKKPEGSGVVQTAAANPPVETSVAKEEIISNSVKAETSTVQIEASELEPERTVRDSATKPSEVETSKGESEVAVTGSETKPADDELSKVKSEEILLPPFNLQKEEVVKDNSETELLESAITTVTERETKVKPEELRVPASGSSEELSSVGKLEDVQSDSAVKPATVETAEAVSEAVPPDSAAVSVEMLSAEDIQPSNKLDTPVKNAVTGTVENKLEIPVASAVVSIEKKPEKSVTKVGTALEKKLEKSMAKIGTAMEKKLEKPVASTGVDTEKKPEKPVDNVETDVEKKPEKPVDNVGMDVEKKPEKPVDNVITDIENIEKNVLNEKNERTLIKAHQNKGPGQVKIDETSKASVSAAAFVSIKEPTSVGKTILKAVVSIPDISKSRVMMRRNETSLTKTEEQKASSKPETRSRSATEKKLSSKEVGQPRATGSRSGLPDSVTKSKLDISPVLVKGGSGRSSSQQDKDSRVESRGSSKQSQERESRSSSMKKDDNSNKIPAGRISKTSKTVSSGNVKPKEEEELFPFNLDEFVTVDEVVEETDSPLQTRRNPPRGKRKDSAKNNSSSEPVTKRKKGKSSIAHVDESELSFVTLDEIVEEEEMTAQLVGDSNLEAITDPQSLVTVDEVNEEEELISEAVKDPQSLVTLDEISEQEEFSSHDLPKDVSSSVSEEQDLLKAEPLVTVDEIGEVEELPLNEPSDLNIEETLKPKEEDDKRAIEDPGDFISSQLPEDPSTLVTVDEIQEDSDDQPLVTLDEVTEDDEDFLADFNRLKEELNFVTVDEVGEEDEEEENTFIGTNLEEEDIVAIAGPEEMEILADIGREGEDIFANSKSEEKVTLEADTKEIENETLVADKQEADEALSANTEETEKAETWNKTSEKEIKGETESEQQSAELSLSKAHESNEMGRQTKNKHQPPVKTPGKRGRPRKRPIVPSPETPIETEQEKCDKNTEETADKTELVESSTSDSNKEKIGKGTLTTPESNSLSEDSQTTTVTDSVSSSLLPDINASLLPDADIKQEGCSETPSQSSEKPVKDELGSGDTEPESKRRKVDSSADKLKTPSTPKGLDFLVPKAGYFCQICSLFYAGEMSMKNHCKTLRHQQNMEKFMAKQKDGEDEEEDEGEELSSR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM92</accession>
    <name>A0A452HM92_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">protein-tyrosine-phosphatase</fullName>
            <ecNumber evidence="3">3.1.3.48</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the protein-tyrosine phosphatase family. Receptor class 4 subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.48" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452HM92"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018339.1">
        <property type="protein sequence ID" value="ENSGAGP00000016055.1"/>
        <property type="gene ID" value="ENSGAGG00000010212.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005925">
        <property type="term" value="C:focal adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="CDD" id="cd14621">
        <property type="entry name" value="R-PTPc-A-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd14623">
        <property type="entry name" value="R-PTPc-A-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050348">
        <property type="entry name" value="Protein-Tyr_Phosphatase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000242">
        <property type="entry name" value="PTP_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016130">
        <property type="entry name" value="Tyr_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003595">
        <property type="entry name" value="Tyr_Pase_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016336">
        <property type="entry name" value="Tyr_Pase_rcpt_a/e-type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19134">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19134:SF433">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00102">
        <property type="entry name" value="Y_phosphatase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF002006">
        <property type="entry name" value="PTPR_alpha_epsilon"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00700">
        <property type="entry name" value="PRTYPHPHTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00194">
        <property type="entry name" value="PTPc"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00404">
        <property type="entry name" value="PTPc_motif"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00383">
        <property type="entry name" value="TYR_PHOSPHATASE_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50055">
        <property type="entry name" value="TYR_PHOSPHATASE_PTP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="12" id="KW-0472">Membrane</keyword>
    <keyword evidence="5 16" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="10" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
    <keyword evidence="6" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11" id="KW-1133">Transmembrane helix</keyword>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="17">
        <location>
            <begin position="69"/>
            <end position="329"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="18">
        <location>
            <begin position="249"/>
            <end position="320"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="17">
        <location>
            <begin position="361"/>
            <end position="619"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="18">
        <location>
            <begin position="535"/>
            <end position="610"/>
        </location>
    </feature>
    <feature type="active site" description="Phosphocysteine intermediate" evidence="14">
        <location>
            <position position="270"/>
        </location>
    </feature>
    <feature type="active site" description="Phosphocysteine intermediate" evidence="14">
        <location>
            <position position="560"/>
        </location>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="238"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <begin position="270"/>
            <end position="276"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="314"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="modified residue" description="Phosphotyrosine" evidence="16">
        <location>
            <position position="626"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007765"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00013064"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PIRSR" id="PIRSR002006-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PIRSR" id="PIRSR002006-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PIRSR" id="PIRSR002006-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016055.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="630" mass="72538" checksum="D007825A83F21FAE" modified="2025-10-08" version="2">MTLPLPWIPTPIQPSTNEHRCNSLCDIAEPQSMPLLARSPSTNRKYPPLPVDKLEEEINRRMADDNKLFREEFNALPACPIQATCEAASKEENKEKNRYVNILPYDHSRVHLTPIEGVPDSDYINASFINGYQEKNKFIAAQGPKEETVNDFWRMIWEQNTATIVMVTNLKERKECKCAQYWPDQGCWTYGNIRVSVEDVTVLVDYTVRKFCIQQVGDVTNKKPQRLVTQFHFTSWPDFGVPFTPIGMLKFLKKVKTCNPQYAGAIVVHCSAGVGRTGTFIVIDAMLDMMHAERKVDVYGFVSRIRAQRCQMVQTDMQYVFIYQALLEHYLYGDTELEVTSLETHLQKIYNKVPGTSSNGLEEEFKKLTSIKIQNDKMRTGNLPANMKKNRVLQIIPYEFNRVIIPVKRGEENTDYVNASFIDGYRQKDSYIASQGPLQHTIEDFWRMIWEWKSCSIVMLTELEERGQEKCAQYWPSDGSVSYGDITMELKKEEECESYTVRDLLVTNTRENKSRQIRQFHFHGWPEVGIPSDGKGMINIIAAVQKQQQQSGNHPITVHCSAGAGRTGTFCALSTVLERVKAEGILDVFQTVKSLRLQRPHMVQTLEQYEFCYKVVQEYIDAFSDYANFK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM93</accession>
    <name>A0A452HM93_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Putative malate dehydrogenase 1B</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the LDH/MDH superfamily. MDH type 2 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HM93"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018348.1">
        <property type="protein sequence ID" value="ENSGAGP00000016064.1"/>
        <property type="gene ID" value="ENSGAGG00000012069.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016615">
        <property type="term" value="F:malate dehydrogenase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016616">
        <property type="term" value="F:oxidoreductase activity, acting on the CH-OH group of donors, NAD or NADP as acceptor"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006108">
        <property type="term" value="P:malate metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006099">
        <property type="term" value="P:tricarboxylic acid cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.720:FF:000144">
        <property type="entry name" value="Malate dehydrogenase [NADP]"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.110.10:FF:000006">
        <property type="entry name" value="putative malate dehydrogenase 1B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.110.10">
        <property type="entry name" value="Lactate dehydrogenase/glycoside hydrolase, family 4, C-terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.720">
        <property type="entry name" value="NAD(P)-binding Rossmann-like Domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022383">
        <property type="entry name" value="Lactate/malate_DH_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015955">
        <property type="entry name" value="Lactate_DH/Glyco_Ohase_4_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010945">
        <property type="entry name" value="Malate_DH_type2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036291">
        <property type="entry name" value="NAD(P)-bd_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23382">
        <property type="entry name" value="MALATE DEHYDROGENASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02866">
        <property type="entry name" value="Ldh_1_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56327">
        <property type="entry name" value="LDH C-terminal domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51735">
        <property type="entry name" value="NAD(P)-binding Rossmann-fold domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0520">NAD</keyword>
    <keyword evidence="3" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0816">Tricarboxylic acid cycle</keyword>
    <feature type="domain" description="Lactate/malate dehydrogenase C-terminal" evidence="7">
        <location>
            <begin position="377"/>
            <end position="457"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="507"/>
            <end position="568"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="548"/>
            <end position="568"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00009613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022532"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023027"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00039310"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF02866"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016064.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="568" mass="63885" checksum="0FF2686A4EBC5EA7" modified="2019-05-08" version="1">MAKFVLAGKADCPYFAKAELLADYLQINLPNFKIHKITQHPDKWEQWLRDICEKNGWKHKRSPIVWRELLDRGGKGLLLGGFNDFMEHAQHYYGITSDMLSEQMLRIAKENLQTHIEIEKEEEYIKSLINPLQVWISSASAPACYNLIPLLANGEVFGMATEVSIHLLDSSHYKEILHGIVMEAEDLAFPLLRSVSMHTELDDAFVQADIVILLDDILFQHEIPSLEDCIRQVTEQCKVYGSLIEKNANSKVKVIVSGKTFVNLRALMIMTFAPSIDGQNIIAVAMFLENAAKAMLARKLNMNSAGVKDVIVWGNISGNSYVDLSKAKVYKYDSAIWGPPNFSRHLLDMIYDGSWVRSEFAFVLSSLSSWGHHCLGMSPAHVIATVLRYWYQDSPPGEIVSMGILSEGQFCIPEGIIFSMPVRFQDGSWEVITETEISEETQETLKRLAHDLIQEKQVALGEIPEMRPYKEPAFNGFNLLALLEEENVQTPPEDLDTLSNGLLQPETQEGEKVQATTDEPEAPPDGLLQSEMLEEDKVQVPIDDPETSSESLNQSETSLSNAENNENP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM95</accession>
    <name>A0A452HM95_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="3">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000018352.1">
        <property type="protein sequence ID" value="ENSGAGP00000016068.1"/>
        <property type="gene ID" value="ENSGAGG00000012070.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005814">
        <property type="term" value="C:centriole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036064">
        <property type="term" value="C:ciliary basal body"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007224">
        <property type="term" value="P:smoothened signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029246">
        <property type="entry name" value="TALPID3"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15721">
        <property type="entry name" value="KIAA0586 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15721:SF2">
        <property type="entry name" value="PROTEIN TALPID3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15324">
        <property type="entry name" value="TALPID3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="678"/>
            <end position="711"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1150"/>
            <end position="1176"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1207"/>
            <end position="1258"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1293"/>
            <end position="1319"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="249"/>
            <end position="293"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="500"/>
            <end position="527"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="572"/>
            <end position="599"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="698"/>
            <end position="711"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="1207"/>
            <end position="1219"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="1306"/>
            <end position="1319"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016068.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1634" mass="179831" checksum="6250C6390778D50B" modified="2019-05-08" version="1">MEVESNSSVSQDSMASFTAGDVLTRSTSLHTKEKPTGAWTAATDCTRKKVKISVKKLREVASPYQADDAALFRDKTSYIPPALSANKPLPNRSTVTTSSSVTQAVRSTLAERSSEHLLSVENDLEPTPLSQKDYSKEAAVKGVPLQVMEDGNKPEDGDIFISQYTTGEKEALRAVLKQRTQNPPVFKEVKVQLLENVSAEKKESTQDTRSAHSNIDSAATIAAATAAAIATTAPLLKVQSDLEAKVSSVSALLSKLQETDKQLQRVAEKQTNIKAQQQERSRCHDRVNELEKQMTVFMEQRIQHLEKLQQQQIHVQSHLISSAINTGGFQPVNIPSSNLVAKHSTNPELQQPLTNQVSSCHRNLFSTNAVPTQAYSSQFGNYGVCTQKSPLKTPAPRRYAPVPVSKNLKISQKISKKERPIAEKENVPKSAYGGVVDGGRLLEQILNSQETPLRQTESSEKSLLNNTKMAWHSERECRPPALQTDSFPAFEGPLQDYNSINKTVKKAEDVLQDLGQLKREMHGMLQEAKAWKSDMNDFVKPTNPVVAPSLAECHQVSKPSILQTVKAPKSVLKDAERILRGVQNNKKVLEENLEAIIRAKDGDAMYTFINALTTNRHVLEEIRIRKTVDEWIKAISVEIQAEMAKNDYEQTQYNQKDPKTPWIKRTQNTRDMKTNKEIKAKTKKPQGSLTKKYLSAAKPSQKQSENNFGRQSLRSYSVSETLLKKEGRAEGPLKASAVVQNEEYLSQVYGKPIYQGHRSTLKKAPYLRFNSPSPKSKPQRPKVIECVRGTKVKSARTQTNSCTQKVIISPKRQRPLSAPLKESHYLFSPSREMPDLSGPLEGHLIPMAIPLGQMQINGISPQPAGVVIGKPHPVTVTTSVLPTSPKPQPKLKKPNIAVIEMRSEKKDPPKLTVQVLPNVDIDSISNDSVTINQASTSSEAALLPVSSLIQGPEQTQYEEEELKFPGTNFVDVTDIIQDQEEEDRIPELSEPILEFNRQTEVVSPKYNGPPFPPTAPASQQPADILDEIIERRETLENRLISWVEQEVMARIISGMYPVQQERVPSVSASDSEESEAVTSDIVEATGGGGFQLFVSAGVPVNSEMISHFVNEALTETIAVMLGDREAQKAVPAANVLQSTTVLPESVVSTPLATPPHTPLPEKEPAPVKTPVSSPSVTEIGGDVGEQEQMTEKGFDIPAATTPICTPVFTPVDTPPRVTTPSPPVSEHGSDATRMGSPEPPNPWGDAELPLDEEKPSPLNEETLHPRAVVMSVAKDEEPEMLILPSPEPVRPFESLPCEPRVPSPLHTPSSGPSTHESSLTVTETETADKPISEGEVLFSYGQLVAARALAEGGLSLPNLSESLTSTLHDANEMDYDPPSEGQVMRRPHRGYHRDPMLSLFAKLNQAPVALHEVIYHSEDSDNSVGELSEGQRPRLTRAAESILMGHSVYTDQVAGLVTEQRPHRVPSPGECDRVTAELLGDAADVSHGPMSMAELESQPLLNPAPQTAQTNNRLTSLSEDLPEKQSQKVVQVEPVRSRVIHVRNKSEVTQQQDSQGDMDRTHVEPNMYLTLLLKDKAYAPGIKLTVTGRPNDSSDSYDASTRIQCASDRSQIVPTETRRKLEIEYIKQLSVQII</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM96</accession>
    <name>A0A452HM96_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Receptor-type tyrosine-protein phosphatase</fullName>
            <ecNumber evidence="16">3.1.3.48</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">Tyrosine protein phosphatase which is involved in integrin-mediated focal adhesion formation. Following integrin engagement, specifically recruits BCAR3, BCAR1 and CRK to focal adhesions thereby promoting SRC-mediated phosphorylation of BRAC1 and the subsequent activation of PAK and small GTPase RAC1 and CDC42.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="16">
            <text>O-phospho-L-tyrosyl-[protein] + H2O = L-tyrosyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:10684"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="EC" id="3.1.3.48"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell junction</location>
            <location evidence="1">Focal adhesion</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 16">Belongs to the protein-tyrosine phosphatase family. Receptor class 4 subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.48" evidence="16"/>
    <dbReference type="AlphaFoldDB" id="A0A452HM96"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018344.1">
        <property type="protein sequence ID" value="ENSGAGP00000016060.1"/>
        <property type="gene ID" value="ENSGAGG00000010212.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005925">
        <property type="term" value="C:focal adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="CDD" id="cd14621">
        <property type="entry name" value="R-PTPc-A-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd14623">
        <property type="entry name" value="R-PTPc-A-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000007">
        <property type="entry name" value="Receptor-type tyrosine-protein phosphatase alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000021">
        <property type="entry name" value="Receptor-type tyrosine-protein phosphatase alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050348">
        <property type="entry name" value="Protein-Tyr_Phosphatase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000242">
        <property type="entry name" value="PTP_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016130">
        <property type="entry name" value="Tyr_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003595">
        <property type="entry name" value="Tyr_Pase_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016336">
        <property type="entry name" value="Tyr_Pase_rcpt_a/e-type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19134">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19134:SF433">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00102">
        <property type="entry name" value="Y_phosphatase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF002006">
        <property type="entry name" value="PTPR_alpha_epsilon"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00700">
        <property type="entry name" value="PRTYPHPHTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00194">
        <property type="entry name" value="PTPc"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00404">
        <property type="entry name" value="PTPc_motif"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00383">
        <property type="entry name" value="TYR_PHOSPHATASE_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50055">
        <property type="entry name" value="TYR_PHOSPHATASE_PTP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="11" id="KW-0965">Cell junction</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="14" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9 16" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="13" id="KW-0472">Membrane</keyword>
    <keyword evidence="5 19" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="10 16" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
    <keyword evidence="6" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="12" id="KW-1133">Transmembrane helix</keyword>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="20">
        <location>
            <begin position="99"/>
            <end position="359"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="21">
        <location>
            <begin position="279"/>
            <end position="350"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="20">
        <location>
            <begin position="391"/>
            <end position="649"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="21">
        <location>
            <begin position="565"/>
            <end position="640"/>
        </location>
    </feature>
    <feature type="active site" description="Phosphocysteine intermediate" evidence="17">
        <location>
            <position position="300"/>
        </location>
    </feature>
    <feature type="active site" description="Phosphocysteine intermediate" evidence="17">
        <location>
            <position position="590"/>
        </location>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="268"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <begin position="300"/>
            <end position="306"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="344"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="modified residue" description="Phosphotyrosine" evidence="19">
        <location>
            <position position="656"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007765"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00054255"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PIRNR" id="PIRNR002006"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="PIRSR" id="PIRSR002006-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="PIRSR" id="PIRSR002006-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="PIRSR" id="PIRSR002006-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016060.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="660" mass="75527" checksum="81D13005E00DC92F" modified="2019-05-08" version="1">HPATLPGRSGGPECWRHAAGPGASLPGWELRGWAARSHGLDVVRGPNEHRCNSLCDIAEPQSMPLLARSPSTNRKYPPLPVDKLEEEINRRMADDNKLFREEFNALPACPIQATCEAASKEENKEKNRYVNILPYDHSRVHLTPIEGVPDSDYINASFINGYQEKNKFIAAQGPKEETVNDFWRMIWEQNTATIVMVTNLKERKECKCAQYWPDQGCWTYGNIRVSVEDVTVLVDYTVRKFCIQQVGDVTNKKPQRLVTQFHFTSWPDFGVPFTPIGMLKFLKKVKTCNPQYAGAIVVHCSAGVGRTGTFIVIDAMLDMMHAERKVDVYGFVSRIRAQRCQMVQTDMQYVFIYQALLEHYLYGDTELEVTSLETHLQKIYNKVPGTSSNGLEEEFKKLTSIKIQNDKMRTGNLPANMKKNRVLQIIPYEFNRVIIPVKRGEENTDYVNASFIDGYRQKDSYIASQGPLQHTIEDFWRMIWEWKSCSIVMLTELEERGQEKCAQYWPSDGSVSYGDITMELKKEEECESYTVRDLLVTNTRENKSRQIRQFHFHGWPEVGIPSDGKGMINIIAAVQKQQQQSGNHPITVHCSAGAGRTGTFCALSTVLERVKAEGILDVFQTVKSLRLQRPHMVQTLEQYEFCYKVVQEYIDAFSDYANFK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM98</accession>
    <name>A0A452HM98_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="3">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000018358.1">
        <property type="protein sequence ID" value="ENSGAGP00000016072.1"/>
        <property type="gene ID" value="ENSGAGG00000012070.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005814">
        <property type="term" value="C:centriole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036064">
        <property type="term" value="C:ciliary basal body"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007224">
        <property type="term" value="P:smoothened signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029246">
        <property type="entry name" value="TALPID3"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15721">
        <property type="entry name" value="KIAA0586 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15721:SF2">
        <property type="entry name" value="PROTEIN TALPID3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15324">
        <property type="entry name" value="TALPID3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="664"/>
            <end position="697"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1136"/>
            <end position="1162"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1193"/>
            <end position="1244"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1279"/>
            <end position="1305"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="249"/>
            <end position="293"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="558"/>
            <end position="585"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="684"/>
            <end position="697"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="1193"/>
            <end position="1205"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="1292"/>
            <end position="1305"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016072.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1620" mass="178166" checksum="21B8347673C5E703" modified="2019-05-08" version="1">MEVESNSSVSQDSMASFTAGDVLTRSTSLHTKEKPTGAWTAATDCTRKKVKISVKKLREVASPYQADDAALFRDKTSYIPPALSANKPLPNRSTVTTSSSVTQAVRSTLAERSSEHLLSVENDLEPTPLSQKDYSKEAAVKGVPLQVMEDGNKPEDGDIFISQYTTGEKEALRAVLKQRTQNPPVFKEVKVQLLENVSAEKKESTQDTRSAHSNIDSAATIAAATAAAIATTAPLLKVQSDLEAKVSSVSALLSKLQETDKQLQRVAEKQTNIKAQQQERSRCHDRVNELEKQMTVFMEQRIQHLEKLQQQQIHVQSHLISSAINTGGFQPVNIPSSNLVAKHSTNPELQQPLTNQVSSCHRNLFSTNAVPTQAYSSQFGNYGVCTQKSPLKTPAPRRYAPVPVSKNVKISQKISKKERPIAEKENVPKSAYGGVVDGGRLLEQILNSQETPLRQTESSEKSLLNNTKMAWHSERECRPPALQTDSFPAFEGPLQDYNSINKTVKKAEDVLQDLGQLKREMHGMLQPTNPVVAPSLAECHQVSKPSILQTVKAPKSVLKDAERILRGVQNNKKVLEENLEAIIRAKDGDAMYTFINALTTNRHVLEEIRIRKTVDEWIKAISVEIQAEMAKNDYEQTQYNQKDPKTPWIKRTQNTRDMKTNKEIKAKTKKPQGSLTKKYLSAAKPSQKQSENNFGRQSLRSYSVSETLLKKEGRAEGPLKASAVVQNEEYLSQVYGKPIYQGHRSTLKKAPYLRFNSPSPKSKPQRPKVIECVRGTKVKSARTQTNSCTQKVIISPKRQRPLSAPLKESHYLFSPSREMPDLSGPLEGHLIPMAIPLGQMQINGISPQPAGVVIGKPHPVTVTTSVLPTSPKPQPKLKKPNIAVIEMRSEKKDPPKLTVQVLPNVDIDSISNDSVTINQASTSSEAALLPVSSLIQGPEQTQYEEEELKFPGTNFVDVTDIIQDQEEEDRIPELSEPILEFNRQTEVVSPKYNGPPFPPTAPASQQPADILDEIIERRETLENRLISWVEQEVMARIISGMYPVQQERVPSVSASDSEESEAVTSDIVEATGGGGFQLFVSAGVPVNSEMISHFVNEALTETIAVMLGDREAQKAVPAANVLQSTTVLPESVVSTPLATPPHTPLPEKEPAPVKTPVSSPSVTEIGGDVGEQEQMTEKGFDIPAATTPICTPVFTPVDTPPRVTTPSPPVSEHGSDATRMGSPEPPNPWGDAELPLDEEKPSPLNEETLHPRAVVMSVAKDEEPEMLILPSPEPVRPFESLPCEPRVPSPLHTPSSGPSTHESSLTVTETETADKPISEGEVLFSYGQLVAARALAEGGLSLPNLSESLTSTLHDANEMDYDPPSEGQVMRRPHRGYHRDPMLSLFAKLNQAPVALHEVIYHSEDSDNSVGELSEGQRPRLTRAAESILMGHSVYTDQVAGLVTEQRPHRVPSPGECDRVTAELLGDAADVSHGPMSMAELESQPLLNPAPQTAQTNNRLTSLSEDLPEKQSQKVVQVEPVRSRVIHVRNKSEVTQQQDSQGDMDRTHVEPNMYLTLLLKDKAYAPGIKLTVTGRPNDSSDSYDASTRIQCASDRSQIVPTETRRKLEIEYIKQLSVQII</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM99</accession>
    <name>A0A452HM99_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="19">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="12">Belongs to the APP family.</text>
    </comment>
    <comment type="caution">
        <text evidence="12">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HM99"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018400.1">
        <property type="protein sequence ID" value="ENSGAGP00000016113.1"/>
        <property type="gene ID" value="ENSGAGG00000012099.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009986">
        <property type="term" value="C:cell surface"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012505">
        <property type="term" value="C:endomembrane system"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043005">
        <property type="term" value="C:neuron projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0099503">
        <property type="term" value="C:secretory vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140677">
        <property type="term" value="F:molecular function activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004867">
        <property type="term" value="F:serine-type endopeptidase inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046914">
        <property type="term" value="F:transition metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007409">
        <property type="term" value="P:axonogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007417">
        <property type="term" value="P:central nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010604">
        <property type="term" value="P:positive regulation of macromolecule metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051246">
        <property type="term" value="P:regulation of protein metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd21709">
        <property type="entry name" value="JMTM_APLP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd22607">
        <property type="entry name" value="Kunitz_ABPP-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1490.140:FF:000001">
        <property type="entry name" value="Amyloid beta (A4) protein b"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.570.10:FF:000001">
        <property type="entry name" value="Amyloid beta A4 protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.770:FF:000001">
        <property type="entry name" value="Amyloid beta A4 protein-like isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.410.10:FF:000003">
        <property type="entry name" value="amyloid-like protein 2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.770">
        <property type="entry name" value="Amyloid precursor protein, E2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1490.140">
        <property type="entry name" value="Amyloidogenic glycoprotein, copper-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.570.10">
        <property type="entry name" value="Amyloidogenic glycoprotein, heparin-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.410.10">
        <property type="entry name" value="Pancreatic trypsin inhibitor Kunitz domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Amyloid_Cu-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008155">
        <property type="entry name" value="Amyloid_glyco"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011178">
        <property type="entry name" value="Amyloid_glyco_Cu-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024329">
        <property type="entry name" value="Amyloid_glyco_E2_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008154">
        <property type="entry name" value="Amyloid_glyco_extra"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015849">
        <property type="entry name" value="Amyloid_glyco_heparin-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036454">
        <property type="entry name" value="Amyloid_glyco_heparin-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019745">
        <property type="entry name" value="Amyloid_glyco_intracell_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019543">
        <property type="entry name" value="APP_amyloid_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019744">
        <property type="entry name" value="APP_CUBD_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036176">
        <property type="entry name" value="E2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002223">
        <property type="entry name" value="Kunitz_BPTI"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036880">
        <property type="entry name" value="Kunitz_BPTI_sf"/>
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    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020901">
        <property type="entry name" value="Prtase_inh_Kunz-CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23103">
        <property type="entry name" value="ALZHEIMER'S DISEASE BETA-AMYLOID RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23103:SF14">
        <property type="entry name" value="AMYLOID BETA PRECURSOR LIKE PROTEIN 2"/>
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    <dbReference type="Pfam" id="PF10515">
        <property type="entry name" value="APP_amyloid"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="APP_Cu_bd"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Kunitz_BPTI"/>
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    <dbReference type="PRINTS" id="PR00203">
        <property type="entry name" value="AMYLOIDA4"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00759">
        <property type="entry name" value="BASICPTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00006">
        <property type="entry name" value="A4_EXTRA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00131">
        <property type="entry name" value="KU"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56491">
        <property type="entry name" value="A heparin-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF89811">
        <property type="entry name" value="Amyloid beta a4 protein copper binding domain (domain 2)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57362">
        <property type="entry name" value="BPTI-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109843">
        <property type="entry name" value="CAPPD, an extracellular domain of amyloid beta A4 protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00319">
        <property type="entry name" value="APP_CUBD"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="APP_E1"/>
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    </dbReference>
    <dbReference type="PROSITE" id="PS51870">
        <property type="entry name" value="APP_E2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00320">
        <property type="entry name" value="APP_INTRA"/>
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        <property type="entry name" value="BPTI_KUNITZ_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50279">
        <property type="entry name" value="BPTI_KUNITZ_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0186">Copper</keyword>
    <keyword evidence="10 12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9 14" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="2" id="KW-0646">Protease inhibitor</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0722">Serine protease inhibitor</keyword>
    <keyword evidence="5 15" id="KW-0732">Signal</keyword>
    <keyword evidence="3 14" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 14" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="15">
        <location>
            <begin position="1"/>
            <end position="25"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5018987577" evidence="15">
        <location>
            <begin position="26"/>
            <end position="749"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="680"/>
            <end position="702"/>
        </location>
    </feature>
    <feature type="domain" description="E1" evidence="17">
        <location>
            <begin position="40"/>
            <end position="201"/>
        </location>
    </feature>
    <feature type="domain" description="BPTI/Kunitz inhibitor" evidence="16">
        <location>
            <begin position="302"/>
            <end position="352"/>
        </location>
    </feature>
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        <location>
            <begin position="365"/>
            <end position="556"/>
        </location>
    </feature>
    <feature type="region of interest" description="GFLD subdomain" evidence="12">
        <location>
            <begin position="40"/>
            <end position="135"/>
        </location>
    </feature>
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        <location>
            <begin position="143"/>
            <end position="201"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="209"/>
            <end position="263"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="13">
        <location>
            <begin position="209"/>
            <end position="224"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="13">
        <location>
            <begin position="233"/>
            <end position="261"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="85"/>
            <end position="129"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="110"/>
            <end position="117"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
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            <begin position="145"/>
            <end position="199"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
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            <end position="186"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="170"/>
            <end position="198"/>
        </location>
    </feature>
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        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
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        <source>
            <dbReference type="ARBA" id="ARBA00022690"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
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            <dbReference type="ARBA" id="ARBA00022723"/>
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        </source>
    </evidence>
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        <source>
            <dbReference type="PROSITE" id="PS51869"/>
        </source>
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        <source>
            <dbReference type="PROSITE" id="PS51870"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016113.1"/>
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    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="749" mass="85461" checksum="EE6F67F2F7E21A8C" modified="2019-05-08" version="1">MVAGGELALLLLPLLALTGPAATLAGYIEALAANAGTGFAVAEPQIAMFCGKLNMHVNIQTGKWEPDTSGTKSCFGTKEEILQYCEEIYPELQITNVVEANQPVSIDNWCKKGKKQCKGHTHIVVPYKCLVGEFVSDVLLVPEKCRFFHKERMDVCESHQHWHTVAKEACLTEGMILHSYGMLLPCGVDQFHGTEYVCCPQTKIVDEALSQEEEDEDEDEDDDYDLYKSEFPTEPDVEEFTGTAVEEDEEEEDEEEEEVVEDRDYYYDNYKVDDYNEETPTESSNDRSLSEKEMISDMKAVCSQEAMTGPCRAVMPRWYFDLYKRKCIRFIYGGCGGNRNNFESEEYCMAVCKKMIPPTPVPTDDVDVYFETPADDNEHARFQKAKEQLEVRHHNRMDRVKKEWEEAEYESKNLPKAERQTLIQHFQAMVKSLEKEAASEKQQLVETHLARVEAMLNDRRRIALENYLAALQADPPRPHRILQALKRYVRAENKDRLHTVRHYQHVLAVDPEKAAQMKSQVMTHLHVIEERMNQSLSLLYKVPYVAEEIQDEIDELLQEQRADMDQFTSSISESQVDVRVSSEESEEIPLQEGKPFRPFQVKTFPAVPGNEGSGMADLDGLIGAEEKVINSKNKVDENVVIDETLDVKEMIFNAERVGGLMDEPDNESSLREDFSLSSSALIGLLVIAVAIATVIVISLVMLRKRQYGTISHGIVEVDPMLTPEERHLNKMQNHGYENPTYKYLEQMQI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP99</accession>
    <name>A0A452IP99_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">pantothenate kinase</fullName>
            <ecNumber evidence="5">2.7.1.33</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>(R)-pantothenate + ATP = (R)-4'-phosphopantothenate + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:16373"/>
            <dbReference type="ChEBI" id="CHEBI:10986"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29032"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.1.33"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="3">Cofactor biosynthesis; coenzyme A biosynthesis; CoA from (R)-pantothenate: step 1/5.</text>
    </comment>
    <comment type="subunit">
        <text evidence="4">Homodimer.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="13">Belongs to the type II pantothenate kinase family.</text>
    </comment>
    <dbReference type="EC" id="2.7.1.33" evidence="5"/>
    <dbReference type="AlphaFoldDB" id="A0A452IP99"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033682.1">
        <property type="protein sequence ID" value="ENSGAGP00000029664.1"/>
        <property type="gene ID" value="ENSGAGG00000021410.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004594">
        <property type="term" value="F:pantothenate kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015937">
        <property type="term" value="P:coenzyme A biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd24135">
        <property type="entry name" value="ASKHA_NBD_PanK-II_Pank1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.420.40:FF:000244">
        <property type="entry name" value="pantothenate kinase 1-like isoform X3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.420.510:FF:000001">
        <property type="entry name" value="pantothenate kinase 2, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.420.40:FF:000392">
        <property type="entry name" value="pantothenate kinase 3-like isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.420.510">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043129">
        <property type="entry name" value="ATPase_NBD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004567">
        <property type="entry name" value="Type_II_PanK"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00555">
        <property type="entry name" value="panK_eukar"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12280">
        <property type="entry name" value="PANTOTHENATE KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12280:SF23">
        <property type="entry name" value="PANTOTHENATE KINASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03630">
        <property type="entry name" value="Fumble"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53067">
        <property type="entry name" value="Actin-like ATPase domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="11" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="12" id="KW-0173">Coenzyme A biosynthesis</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="10" id="KW-0418">Kinase</keyword>
    <keyword evidence="9" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0808">Transferase</keyword>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1"/>
            <end position="44"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="66"/>
            <end position="156"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="12"/>
            <end position="25"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="97"/>
            <end position="120"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001206"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005225"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00011738"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012102"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022993"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00060870"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029664.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="521" mass="57147" checksum="F21FBBB03104BC30" modified="2019-05-08" version="1">GSRVAWGVNGRPGTSSERPGPSQSRCVPLASLRGGGRIRKEMGGEWNGGGEGACCCQHQAPPLTLQVTLNPAPPPNPPAARRQRSEAATAMEERPRAQPGAQPAAAAPSGSLLNGLLHNGFHTPPLLRQEPPVPGHGSPAKKCRLRRRLDSGRRNRPPFPWFGMDIGGTLVKLVYFEPKDITAEEEQEEVENLKSIRKYLTSNTAYGKTGIRDVHLELKNLTMCGRKGNLHFIRFPTSAMHRFIQMGSEKNFSSLHTTLCATGGGAYKFEEDFRTIADLQLHKLDELDCLIQGLLYVDSVGYNGQPECYYFENPTDPEQCQKKPYCLDNPYPMLLVNIGSGVSILAVYSKDNYKRVTGTSLGGGTFLGLCCLLTGCETFEEALEMAAKGDSTNVDKLVKDIYGGDYERFGLQGSAVASSFGHMMSKEKRDSISKEDLARATLVTITNNIGSIARMCALNENIDRVVFVGNFLRINMISMKVLAYAMDYWSKGQLKALFLEHEGYFGAVGALLELLKMPENQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IPA0</accession>
    <name>A0A452IPA0_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="7">Scavenger receptor class A member 5</fullName>
        </submittedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IPA0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033780.1">
        <property type="protein sequence ID" value="ENSGAGP00000029751.1"/>
        <property type="gene ID" value="ENSGAGG00000021461.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="F:scavenger receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006898">
        <property type="term" value="P:receptor-mediated endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031638">
        <property type="term" value="P:zymogen activation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="entry name" value="Scavenger receptor class A member 5"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="SRCR-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="SR-AI/II"/>
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    </dbReference>
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        <property type="entry name" value="SRCR-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48071:SF24">
        <property type="entry name" value="DELETED IN MALIGNANT BRAIN TUMORS 1 PROTEIN-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48071">
        <property type="entry name" value="SRCR DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00530">
        <property type="entry name" value="SRCR"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="MACSCAVRCPTR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00258">
        <property type="entry name" value="SPERACTRCPTR"/>
    </dbReference>
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        <property type="entry name" value="SR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56487">
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="SRCR_2"/>
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    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="1 3" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="2" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="SRCR" evidence="6">
        <location>
            <begin position="268"/>
            <end position="368"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="202"/>
            <end position="227"/>
        </location>
    </feature>
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            <begin position="139"/>
            <end position="184"/>
        </location>
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        <location>
            <begin position="206"/>
            <end position="215"/>
        </location>
    </feature>
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        <location>
            <begin position="293"/>
            <end position="357"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="3">
        <location>
            <begin position="306"/>
            <end position="367"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="3">
        <location>
            <begin position="337"/>
            <end position="347"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00196"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50287"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029751.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="370" mass="40457" checksum="667AD77C01A63C3E" modified="2019-05-08" version="1">ALCPPPFDTFSAGPVLGLVHWSLWVHAAGKNSLFLMTGSLQKLEGFLWSLQTQASQTDKAVASLRDSLHHQSDMAQLETYRLSVEGNSSRLLLEHHNSLLTHLAAQVESLSDQLTEVGGAVGTINRTFSYDVGIHHTRIQDLQVLISNATEDARQMRRIHIAMEQQLKQELAILNNVTEDLRLKDWEHSIALRNISVIRGKGLPGERGPLGPPGLKGDKGDLGPSGPAGMRGFKGTVQGSWRDGPAGDAHHLLCVSLLLTAAAEEGLIRLVNGSVPHEGRVEVFHDRHWGTVCDDSWDKKDGDVACRMLGFRGAEEVYRMARFGQGIGRIWMDDVACKGTEDSLFQCTFSSWGKSNCGHAEDAGVKCLTQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0X7</accession>
    <name>A0A452J0X7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">LisH domain and HEAT repeat-containing protein KIAA1468</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Golgi apparatus</location>
            <location evidence="2">trans-Golgi network</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Recycling endosome</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038522.1">
        <property type="protein sequence ID" value="ENSGAGP00000034022.1"/>
        <property type="gene ID" value="ENSGAGG00000023990.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055037">
        <property type="term" value="C:recycling endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005802">
        <property type="term" value="C:trans-Golgi network"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032367">
        <property type="term" value="P:intracellular cholesterol transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000080">
        <property type="entry name" value="lisH domain and HEAT repeat-containing protein KIAA1468 homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000218">
        <property type="entry name" value="lisH domain and HEAT repeat-containing protein KIAA1468 homolog isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021133">
        <property type="entry name" value="HEAT_type_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006594">
        <property type="entry name" value="LisH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040362">
        <property type="entry name" value="RELCH"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR32059">
        <property type="entry name" value="RAB11-BINDING PROTEIN RELCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR32059:SF0">
        <property type="entry name" value="RAB11-BINDING PROTEIN RELCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00667">
        <property type="entry name" value="LisH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50077">
        <property type="entry name" value="HEAT_REPEAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50896">
        <property type="entry name" value="LISH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7 11" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0967">Endosome</keyword>
    <keyword evidence="6" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="8" id="KW-0445">Lipid transport</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <feature type="repeat" description="HEAT" evidence="10">
        <location>
            <begin position="963"/>
            <end position="999"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1"/>
            <end position="52"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="398"/>
            <end position="423"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="11">
        <location>
            <begin position="319"/>
            <end position="353"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="12">
        <location>
            <begin position="25"/>
            <end position="35"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="403"/>
            <end position="423"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004172"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004601"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00084053"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00103"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034022.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1173" mass="130042" checksum="6D149224D23D753B" modified="2019-05-08" version="1">MAAAGGGSSVNPFLSDSEEEEPRDGGGGNDPGLGGPLSPQQHEEEEEAAVEVPGVLPGLRAAAPGEAGWAGPGAGEPPRVPVDALAAQLLRDQFLLTALELHTELLESGRELPRLRDYFSNPGNFERQSGTPPAGGAPGLWGAAGLGLLGGCELITEGGGDRAGSISTLDSLDFARYSDDGNRETDERVAENEVPLQERKNYKSSPEIQEPIKPLEKRALNFLVNEFLLKNSYKLTSITFSDESDDQDFELWDDVGLNIPKPPDLLQLYRDFGNHQVTARDVVDVAVGVEEDELEVTTPTLGSVPVFGVSQSTEQFLIVQKLEDQINALNSEKWSLTEQIRRLESEIDFLKNENFAVPVVCDLVHVPLSVPEDNGSYLDIRSSDNDSKHGKTEEISLSISAEADSKTPKDDTLNSLPSKDREKLPSCALSNKAAIHFDKPNRTLSPAFHQALLSFCRMSAESRLGSEVSRIADSEKSVMLMLGRCLPHIVPNVLLAKREELIPLILCTACLHPEPKERDQLLHILFNLIKRPDDEQRQMILTGCVAFARHVGPTRVEAELLPQCWEQINHKYPERRLLVAESCGALAPYLPKEIRSSLVLSMLQQMLMEDKADLVREAVIKSLGIIMGYIDDPDKYQQGFELLLSALGDPSERVVSATHQVFLPAYAAWTTELGNLQLHLIPTLLNKIEKLLREGEHGLDEHKLHMYLSALQSLIPSLFSLVLQNAPFTSKAKLQGEVPPIEVTRFPRPVSPLQDVATIIGSREQLAVLLQLYDYQLELEGTTGWETLLWVVNQLLPQLIEIVGKINVASTACVHEFSRFFWRLCRTFGKIFTNTKVKPQFQEILRLSEENIDSTAGNGVLTKATVPIYATGVLTCYIQEEDRKLLIGFLEDVMTMLSLSHAPLNSLKASFVELGANPAYHELLLTVLWYGVVHTSALVRCTAARMFELLVKGVHETLVAQRVVPALITYSSDPEISVRIATIPAFGTIMETVTQREXRVKMQLASFLEDPQYQDQHSLQTEIIKTFGRVGPNAEPRFRDEFVIPHLHKLALVNNQQSVDSKRLDIATHLFEAYSALSCCFISEDLMVNHFLPGLRCLRTDMEHLSPEHEVILSSMIKECEQKVENKTVQEPQGSMSIAASLVSEDTKTKFLNKMGQLTTSGAMLANVFQRKK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0Y2</accession>
    <name>A0A452J0Y2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">tRNA (32-2'-O)-methyltransferase regulator THADA</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="3">Together with methyltransferase FTSJ1, methylates the 2'-O-ribose of nucleotides at position 32 of the anticodon loop of substrate tRNAs.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the THADA family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038527.1">
        <property type="protein sequence ID" value="ENSGAGP00000034027.1"/>
        <property type="gene ID" value="ENSGAGG00000024118.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030488">
        <property type="term" value="P:tRNA methylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056843">
        <property type="entry name" value="THADA-like_TPR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056842">
        <property type="entry name" value="THADA-like_TPR_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019442">
        <property type="entry name" value="THADA/TRM732_DUF2428"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051954">
        <property type="entry name" value="tRNA_methyltransferase_THADA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14387">
        <property type="entry name" value="THADA/DEATH RECEPTOR INTERACTING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14387:SF7">
        <property type="entry name" value="THYROID ADENOMA-ASSOCIATED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10350">
        <property type="entry name" value="DUF2428"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25150">
        <property type="entry name" value="TPR_Trm732"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25151">
        <property type="entry name" value="TPR_Trm732_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0819">tRNA processing</keyword>
    <feature type="domain" description="tRNA (32-2'-O)-methyltransferase regulator THADA-like TPR repeats region" evidence="6">
        <location>
            <begin position="513"/>
            <end position="785"/>
        </location>
    </feature>
    <feature type="domain" description="DUF2428" evidence="5">
        <location>
            <begin position="929"/>
            <end position="1229"/>
        </location>
    </feature>
    <feature type="domain" description="tRNA (32-2'-O)-methyltransferase regulator THADA-like C-terminal TPR repeats region" evidence="7">
        <location>
            <begin position="1231"/>
            <end position="1394"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00010409"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022694"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00035625"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00035698"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="Pfam" id="PF10350"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="Pfam" id="PF25150"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF25151"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034027.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1792" mass="202084" checksum="6C94E878DFE530BC" modified="2019-05-08" version="1">MVLKKKKEIQVDAFVLECQELETLRSFADAEDQNLASLLLHCVQLSDGIHQIYSMKQIVTLLKKMDRNSACSPMIKCCLDIIGVMYFSLGAKNPLKKMLASSLNGLPEQLMTEAVHSFVRCLKEELKTTDLCLYRKVMDNLASCMENFSLGMIIFLQFLQKSVLEIQEENSGNSIVETRLMHDLLMAIKVSMMLLQKVQENFQGSLWRKPGSLLWQSMCGLLKSFTNFLMNEGLLPTVQTTSGLAVILFTKTMFEPVEKLPCLVSDLLLRSVKCASVPEWFVSSCECLCTEELSDSVLLSLCHGALAMLEWKNGSMSENGEKLLLDIALVLLSLASRLKESNMAVSLSRILAIWTNSALDVLESSSQNLKSSLNGNSNTTGKLLEYVYTHWEHPLDAVRHQAKLIFKNILRIHQATLKGSVVKSDPFFSMLTESLLSLEWHVKGKYASLGCLVECVGVENILTVDRTIPAQILDVMGDQSLAPYASDLLETMFINHKSHLISAFQKNIWINHWHDTWVSPLLLILCEGNPDQTTYVIDYYLPKLLKCSPDSLSYMIKILQASADANIGSCNTRGALGALMACLRTARAHGHLQFREIMRDDLLSIGYVKQGLVHQHNQVRVDALGLLCETHRSTEILTMEEMQLIQFFITYNLNSQSPAVRQQICSLLKKLFCRIQESSQVLYKLEQNKTKQELIKESAKWDHFATLQQYKDFMSSICDKLFEALFPGSSHPTRFSSLTILGSIAEIFSAPEGQTQAIFRLAQEIDPTRVQMLIQCFASTFEEVKVLAFELLMKLYDRLDLQDLENLHLLFQAAMDLSISTKPYDCVTASYLLNFLIHHKGLLNVCLKNQQVADTFQIDEDMSASTMEKNTLAVIKYLLSNLEREIFQAEKSLLQAAASFPMYGRVHCITGALQQLPLNNLTLMTEWRQMVTKLILMSYRLSSVVSPIVQSSSPEGLIPMDNDSETAGRLQMILHEIQPRDTNDYFTQAKIFKEHHRVDFAKLTDDEPVENVCTEIRGKEGQTCDVTAQMVLVCCWRSMKEISLLLGMLCKLLPLQTVSESSDGLIAVEQVKDIGDYFKHHLLQSRHRGAFELAYTGFVMLTEILSRCTNESLRTLPQQWLYNVLEEIKSCDPSSKLCATRRSAGIPFYIQALLASEPKNKTNLLKMTMNELVSLAVPVNESLNIIPQVHALNILRALFRDTRFGENIIPYIADGIQAAVLGFTSPVWAVRNSSTLLFSTLITRIFGVKRGKDENSKKNRMTGREFFSRFPTLYPFLLQQLEVVARTMDSETGELKLHPSLFLLLLILGKLYPSPMDGTYSALSMAPFVPFIMRCGHSPVYRSREMAGRALVPFVMISQIPHTVQSLLAGLPECTNPCIQQNSVHGMLLQVFHLLGSYLESKHRVNSDLQQGLVDIITCIKAKVWLAKRYNPCLVTRAAYLDVLVVLSDYLGKSRRKGIELFGFWEEISKVISASELMTGIYYFSAIPGFTQYLQSITKLVISVLSVAADPDFSNSSSPVAEQMMNSSSMVVHLLQSDFHEVRLPMLEAVLLWWKQVNSKQATEEKMGPLSLLSDLEVILLRMAMKEKHPECFCKVLEILHCMDLRNVLPRTQDSIKMNPKEFLNWAMNIISASDSIKIQSIALKLASKLVTHLVQNCQEVKRFWIQLVTSCCGDEEQTAIRLAAAEVLINITPLFLTNQKLILGLSSTLAMWKCILLLLQSEELIVRDTAADVIRVAQSQSNICNKTGTGWVCARSYLTLMDTHIETFHNTILQIITGPLCSLCLKLKDYY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0Y3</accession>
    <name>A0A452J0Y3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Pleiotrophin</fullName>
            <shortName evidence="8">PTN</shortName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="8">Secreted growth factor that mediates its signal through cell-surface proteoglycan and non-proteoglycan receptors. Regulates many processes like cell proliferation, cell survival, cell growth, cell differentiation and cell migration.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 8">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 8">Belongs to the pleiotrophin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0Y3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038545.1">
        <property type="protein sequence ID" value="ENSGAGP00000034041.1"/>
        <property type="gene ID" value="ENSGAGG00000024186.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008083">
        <property type="term" value="F:growth factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051781">
        <property type="term" value="P:positive regulation of cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.60.10:FF:000001">
        <property type="entry name" value="Pleiotrophin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.90.10:FF:000001">
        <property type="entry name" value="Pleiotrophin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.90.10">
        <property type="entry name" value="Heparin-binding Growth Factor, Midkine, Chain A- C-terminal Domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.20.60.10">
        <property type="entry name" value="Pleiotrophin/Midkine, N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000762">
        <property type="entry name" value="Midkine_heparin-bd_GF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020090">
        <property type="entry name" value="PTN/MK_C_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038130">
        <property type="entry name" value="PTN/MK_C_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020091">
        <property type="entry name" value="PTN/MK_diS_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020089">
        <property type="entry name" value="PTN/MK_N_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037122">
        <property type="entry name" value="PTN/MK_N_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020092">
        <property type="entry name" value="PTN_MK_heparin-bd_GF_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13850:SF1">
        <property type="entry name" value="PLEIOTROPHIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13850">
        <property type="entry name" value="PLEIOTROPHIN FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01091">
        <property type="entry name" value="PTN_MK_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05196">
        <property type="entry name" value="PTN_MK_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00269">
        <property type="entry name" value="PTNMIDKINE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00193">
        <property type="entry name" value="PTN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57288">
        <property type="entry name" value="Midkine"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00619">
        <property type="entry name" value="PTN_MK_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00620">
        <property type="entry name" value="PTN_MK_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6 8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="8" id="KW-0339">Growth factor</keyword>
    <keyword evidence="4 8" id="KW-0358">Heparin-binding</keyword>
    <keyword evidence="7 8" id="KW-0497">Mitogen</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 8" id="KW-0964">Secreted</keyword>
    <keyword evidence="5 8" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="8">
        <location>
            <begin position="1"/>
            <end position="30"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019584908" description="Pleiotrophin" evidence="8">
        <location>
            <begin position="31"/>
            <end position="165"/>
        </location>
    </feature>
    <feature type="domain" description="Pleiotrophin/Midkine N-terminal" evidence="11">
        <location>
            <begin position="32"/>
            <end position="93"/>
        </location>
    </feature>
    <feature type="domain" description="Pleiotrophin/Midkine C-terminal" evidence="10">
        <location>
            <begin position="94"/>
            <end position="158"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="139"/>
            <end position="165"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005403"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022674"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="RuleBase" id="RU369117"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="Pfam" id="PF01091"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF05196"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034041.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="165" mass="18675" checksum="F56A51F2F2235B2A" modified="2019-05-08" version="1">MQQQQQQRRMFTAALLAFIFILAAMGTTEAGKKEKPEKKVKKSDCGEWQWSVCVPTSGDCGLGTREGTRTGAECKQTMKTQRCKIPCNWKKQFGAECKYQFQAWGECDLNTALKTRTGNLKRALHNADCQKIVTISKPCGKLTKPKPQEPKKKKKEGKKQEKMLD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0Y4</accession>
    <name>A0A452J0Y4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Myosin motor domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="6">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <comment type="caution">
        <text evidence="6">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0Y4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038519.1">
        <property type="protein sequence ID" value="ENSGAGP00000034019.1"/>
        <property type="gene ID" value="ENSGAGG00000024156.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016459">
        <property type="term" value="C:myosin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000146">
        <property type="term" value="F:microfilament motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007015">
        <property type="term" value="P:actin filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13140">
        <property type="entry name" value="MYOSIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13140:SF313">
        <property type="entry name" value="UNCONVENTIONAL MYOSIN-VC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 6" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="2 6" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="4 6" id="KW-0505">Motor protein</keyword>
    <keyword evidence="3 6" id="KW-0518">Myosin</keyword>
    <keyword evidence="1 6" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Myosin motor" evidence="7">
        <location>
            <begin position="67"/>
            <end position="628"/>
        </location>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <begin position="161"/>
            <end position="168"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034019.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="628" mass="72124" checksum="FC6A3B18785841B9" modified="2025-10-08" version="2">MFWGSSRSCYNRVWIPDNEEVWQSAEITKSYRTGDRVLHLLLEDGTELNYPVDPAPLPPLRNPDILVGENDLTALSYLHEPAVLHNLKVRFVESKLIYTYSGIILVAMNPYKQLPIYGDAIIHAYSGQNMGDMDPHIFAVAEEAYKQMARNNRNQSVIVSGESGAGKTVSARYAMRYFATVSKSNSNVHVEDKVLASNPITEAVGNAKTTRNDNSSRFGKYTEISFDQRHQITGANMRTYLLEKSRVVFQSENERNYHIFYQLCASALQPEFKHLKLGHAEEFNYTRMGGNTVIEGVDDRADMMETQKTFTLLGLKEDFQMDVFKILAAILHLGNVQMTAVGDEKSSVSLEDKHLNIFCELLDLDNDKVAQWLCHRKIITTSETVIKPMTKLQALNARDALAKKIYSHLFDFIVERINKALHFSGKQHTFIGVLDIYGFETFDVNSFEQFCINYANEKLQQQFNLLLTCAFHNEYMKENIPWTLIDFYDNQPVIDLIEAKMGILELLDEECLLPQGTDENWLQKLYNNYINKNPFFQKPKMSNTSFIIQHFADKVEYKSEGFLEKNRDTVYEVLIDILRNSKFHLCANFFRDRPLPISPYRSTITVKSAKTVVKPPNKQFRTTVGSEV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0Y8</accession>
    <name>A0A452J0Y8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">tRNA (32-2'-O)-methyltransferase regulator THADA</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="3">Together with methyltransferase FTSJ1, methylates the 2'-O-ribose of nucleotides at position 32 of the anticodon loop of substrate tRNAs.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the THADA family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038533.1">
        <property type="protein sequence ID" value="ENSGAGP00000034032.1"/>
        <property type="gene ID" value="ENSGAGG00000024118.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030488">
        <property type="term" value="P:tRNA methylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056843">
        <property type="entry name" value="THADA-like_TPR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056842">
        <property type="entry name" value="THADA-like_TPR_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019442">
        <property type="entry name" value="THADA/TRM732_DUF2428"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051954">
        <property type="entry name" value="tRNA_methyltransferase_THADA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14387">
        <property type="entry name" value="THADA/DEATH RECEPTOR INTERACTING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14387:SF7">
        <property type="entry name" value="THYROID ADENOMA-ASSOCIATED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10350">
        <property type="entry name" value="DUF2428"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25150">
        <property type="entry name" value="TPR_Trm732"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25151">
        <property type="entry name" value="TPR_Trm732_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0819">tRNA processing</keyword>
    <feature type="domain" description="tRNA (32-2'-O)-methyltransferase regulator THADA-like TPR repeats region" evidence="6">
        <location>
            <begin position="516"/>
            <end position="788"/>
        </location>
    </feature>
    <feature type="domain" description="DUF2428" evidence="5">
        <location>
            <begin position="932"/>
            <end position="1232"/>
        </location>
    </feature>
    <feature type="domain" description="tRNA (32-2'-O)-methyltransferase regulator THADA-like C-terminal TPR repeats region" evidence="7">
        <location>
            <begin position="1234"/>
            <end position="1397"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00010409"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022694"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00035625"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00035698"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="Pfam" id="PF10350"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="Pfam" id="PF25150"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF25151"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034032.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1764" mass="198901" checksum="1CDBB31F26F915BA" modified="2019-05-08" version="1">MVLKKKKEIQVDAFVLECQELETLRSFADAEDQNLASLLLHCVQLSDGIHQIYSMKQIVTLLKKMDRNSACSPMIKCCLDIIGVMYFSLGAKNPLKKMLASSLNGLPEQLMTEAVHSFVRCLKEELKTTDLCLYRKVMDNLASCMENFSLGMIIFLQFLQKSVLEIQEENRQLSGNSIVETRLMHDLLMAIKVSMMLLQKVQENFQGSLWRKPGSLLWQSMCGLLKSFTNFLMNEGLLPTVQTTSGLAVILFTKTMFEPVEKLPCLVSDLLLRSVKCASVPEWFVSSCECLCTEELSDSVLLSLCHGALAMLEWKNGSMSENGEKLLLDIALVLLSLASRLKESNMAVSLSRILAIWTNSALDVLESSSQNLKSSLNGNSNTTGKLLEYVYTHWEHPLDAVRHQAKLIFKNILRIHQATLKGSVVKSDPFFSMLTESLLSLEWHVKGKYASLGCLVECVGVENILTVDRTIPAQILDVMGDQSLAPYASDLLETMFINHKSHLISAFQKNIWINHWHDTWVSPLLLILCEGNPDQTTYVIDYYLPKLLKCSPDSLSYMIKILQASADANIGSCNTRGALGALMACLRTARAHGHLQFREIMRDDLLSIGYVKQGLVHQHNQVRVDALGLLCETHRSTEILTMEEMQLIQFFITYNLNSQSPAVRQQICSLLKKLFCRIQESSQVLYKLEQNKTKQELIKESAKWDHFATLQQYKDFMSSICDKLFEALFPGSSHPTRFSSLTILGSIAEIFSAPEGQTQAIFRLAQEIDPTRVQMLIQCFASTFEEVKVLAFELLMKLYDRLDLQDLENLHLLFQAAMDLSISTKPYDCVTASYLLNFLIHHKGLLNVCLKNQQVADTFQIDEDMSASTMEKNTLAVIKYLLSNLEREIFQAEKSLLQAAASFPMYGRVHCITGALQQLPLNNLTLMTEWRQMVTKLILMSYRLSSVVSPIVQSSSPEGLIPMDNDSETAGRLQMILHEIQPRDTNDYFTQAKIFKEHHRVDFAKLTDDEPVENVCTEIRGKEGQTCDVTAQMVLVCCWRSMKEISLLLGMLCKLLPLQTVSESSDGLIAVEQVKDIGDYFKHHLLQSRHRGAFELAYTGFVMLTEILSRCTNESLRTLPQQWLYNVLEEIKSCDPSSKLCATRRSAGIPFYIQALLASEPKNKTNLLKMTMNELVSLAVPVNESLNIIPQVHALNILRALFRDTRFGENIIPYIADGIQAAVLGFTSPVWAVRNSSTLLFSTLITRIFGVKRGKDENSKKNRMTGREFFSRFPTLYPFLLQQLEVVARTMDSETGELKLHPSLFLLLLILGKLYPSPMDGTYSALSMAPFVPFIMRCGHSPVYRSREMAGRALVPFVMISQIPHTVQSLLAGLPECTNPCIQQNSVHGMLLQVFHLLGSYLESKHRVNSDLQQGLVDIITCIKAKVWLAKRQNPCLVTRAAYLDVLVVLSDYLGKSRRKGKSTLFGFWEEISKVISASELMTGIYYFSAIPGFTQYLQSITKLVISVLSVAADPDFSNSSSPVAEQMMNSSSMVVHLLQSDFHEVRLPMLEAVLLWWKQVNSKQATEEKMGPLSLLSDLEVILLRMAMKEKHPECFCKVLEILHCMDLRNVLPRTQDSIKMNPKEFLNWAMNIISCDPESALNCTVEYSHAFLCFQVMESEMRQWIQLVTSCCGDEEQTAIRLAAAEVLINITPLFLTNQKLILGLSSTLAMWKCILLLLQSEELIVRDTAADVIRVAQSQSNICNKTGTRLLRPQGSAAKIF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0Z0</accession>
    <name>A0A452J0Z0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Oxysterol-binding protein-related protein 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">Binds phospholipids; exhibits strong binding to phosphatidic acid and weak binding to phosphatidylinositol 3-phosphate. Stabilizes GTP-bound RAB7A on late endosomes/lysosomes and alters functional properties of late endocytic compartments via its interaction with RAB7A. Binds 25-hydroxycholesterol and cholesterol.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Late endosome</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the OSBP family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0Z0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038571.1">
        <property type="protein sequence ID" value="ENSGAGP00000034061.1"/>
        <property type="gene ID" value="ENSGAGG00000024195.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005770">
        <property type="term" value="C:late endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097038">
        <property type="term" value="C:perinuclear endoplasmic reticulum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015485">
        <property type="term" value="F:cholesterol binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006869">
        <property type="term" value="P:lipid transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd13285">
        <property type="entry name" value="PH_ORP1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000094">
        <property type="entry name" value="Oxysterol-binding protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000098">
        <property type="entry name" value="Oxysterol-binding protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000231">
        <property type="entry name" value="Oxysterol-binding protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.160.120:FF:000005">
        <property type="entry name" value="Oxysterol-binding protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.3490:FF:000003">
        <property type="entry name" value="Oxysterol-binding protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.160.120">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.3490">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037239">
        <property type="entry name" value="OSBP_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000648">
        <property type="entry name" value="Oxysterol-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10972">
        <property type="entry name" value="OXYSTEROL-BINDING PROTEIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10972:SF53">
        <property type="entry name" value="OXYSTEROL-BINDING PROTEIN-RELATED PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01237">
        <property type="entry name" value="Oxysterol_BP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00233">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF144000">
        <property type="entry name" value="Oxysterol-binding protein-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6 12" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="7 13" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0967">Endosome</keyword>
    <keyword evidence="8" id="KW-0445">Lipid transport</keyword>
    <keyword evidence="9" id="KW-0446">Lipid-binding</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <feature type="repeat" description="ANK" evidence="12">
        <location>
            <begin position="47"/>
            <end position="79"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="12">
        <location>
            <begin position="80"/>
            <end position="112"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="12">
        <location>
            <begin position="175"/>
            <end position="207"/>
        </location>
    </feature>
    <feature type="domain" description="PH" evidence="15">
        <location>
            <begin position="237"/>
            <end position="336"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="743"/>
            <end position="770"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="13">
        <location>
            <begin position="435"/>
            <end position="462"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="13">
        <location>
            <begin position="827"/>
            <end position="861"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004603"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023121"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00055419"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00074471"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034061.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="906" mass="102346" checksum="6EEB416442A98E84" modified="2019-05-08" version="1">MITEAEQQLLHDARNGNCEEVKQLLETMDKGEVVVDINCKGRSKSNLGWTPLHLACYFGHGNVVKDLLKAGAEVNVLNDMGDTPLHRAAFTGRKEVVMLLLRYDADTSIVNGGGQTARDVTQDKEIKNMLEAVERTQERKLEELLLGAAREGETAKLSALLNRPKPPDINCTDQMGNTPLHCAAYRAHKHCALKLLKKGADSKIRNKNDQTPFDLAQGAEIKQILAGSIGKVMNKRLKRYEGLLWKSLRFFGWKLYWVVLEHGVLSWYRKQADAVNNDHRQGCKHLTQAVCTVKSTDSCLFSVKCFDDSVHSFKVPKNSPPKARENWLEAIEDHSAYSTHYCTQEQLSDEEDDDTVSVAELQETLKKAQTCQQRLDKEISNFLAMIKSCSTTKGMPPSFLQKIEVVSEVSKETCEALGNCLKLFTKQEGVRNLKLEHEQEKNKILSEALETLATEHHELEQSLVKGSPPLSILSEEQFYDAVSDSESEKSLSGFETVTAYSLEDSVESNNTITSSMSEEKVCGGGGKVCGGGESLSNGIKKHRTSLPSPMFSRNDFSIWSILRKCIGMELSKITMPVIFNEPLSFLQRLTEYMEHTYLIHQASSLSGTVERMQCVAAFAVSAVASQWERTGKPFNPLLGETYELVRHDEAYTWTNPTCCVHNIIVGKLWIEQYGNVEITNHKTGEKCVLNFKPCGLFGKELHKVEGYIQDKSKRKVCSLYGKWTECLYSVDPATFEAYKKNDKKNTEEKKNSKQVGNSEEPDEMPLPDSESVHVIPGSILLWRIAPRPPNSVQMYNFTSFAMALNELDKEMESVVPKTDCRLRPDIRAMENGEIDIASEEKKRLEEKQRAVRKNRSKSEEDWKTSHLTALGPRWFHQGPNPYTGAQDWLYSGNYWDRNYFNLPDIY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0Z1</accession>
    <name>A0A452J0Z1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Myelin transcription factor 1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the MYT1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0Z1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038484.1">
        <property type="protein sequence ID" value="ENSGAGP00000033984.1"/>
        <property type="gene ID" value="ENSGAGG00000024142.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.320.30:FF:000001">
        <property type="entry name" value="Myelin transcription factor 1-like, a"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.320.30">
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013681">
        <property type="entry name" value="Myelin_TF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002515">
        <property type="entry name" value="Znf_C2H2C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036060">
        <property type="entry name" value="Znf_C2H2C_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10816:SF10">
        <property type="entry name" value="MYELIN TRANSCRIPTION FACTOR 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10816">
        <property type="entry name" value="MYELIN TRANSCRIPTION FACTOR 1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08474">
        <property type="entry name" value="MYT1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01530">
        <property type="entry name" value="zf-C2HC"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103637">
        <property type="entry name" value="CCHHC domain"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51802">
        <property type="entry name" value="ZF_CCHHC"/>
        <property type="match status" value="6"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="8" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="9" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 11" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="Myelin transcription factor 1" evidence="14">
        <location>
            <begin position="506"/>
            <end position="763"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="29"/>
            <end position="114"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="189"/>
            <end position="313"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="463"/>
            <end position="486"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="552"/>
            <end position="586"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="640"/>
            <end position="747"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="12">
        <location>
            <begin position="970"/>
            <end position="1032"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="13">
        <location>
            <begin position="53"/>
            <end position="87"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="13">
        <location>
            <begin position="98"/>
            <end position="114"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="13">
        <location>
            <begin position="192"/>
            <end position="214"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="13">
        <location>
            <begin position="222"/>
            <end position="259"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="13">
        <location>
            <begin position="273"/>
            <end position="285"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="13">
        <location>
            <begin position="295"/>
            <end position="307"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="13">
        <location>
            <begin position="556"/>
            <end position="586"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="13">
        <location>
            <begin position="655"/>
            <end position="696"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="13">
        <location>
            <begin position="732"/>
            <end position="743"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010194"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01143"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="Pfam" id="PF08474"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033984.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1098" mass="121585" checksum="65A629C9CC9F47C1" modified="2019-05-08" version="1">VPTELIGQEVSLQSCPLAKKRKLQDAEAEHLVSKRKSHPLKLALDEGYNVDSDGSEDTEVKEESVTDESEGTLEEEETETIEQEETPSPEPAEAKSSRYGQNAATSTPGPSKANYSSYQEIIANSLLNLGQIAEETLAIEGQMPENELQVSKPPVNVVHLVHEGVEEEVVEEECDKEIIIQTEDAEEVIEVTSERSSELCPEHRDNMNCEESCKLQKAMQAEAEDEEDDEEEEEEEEEEEEEEEEEEEEEEEEEEEELASEVICEETPQTSRDSQKTHCEGRFSPKPEYSVIVEVRSDDDKDDDARSQKSTVTDESEMYDMMTRGNLGLLEQAIALKAEQVKIMREPNRLPGEHVKHFQVDEKQNKPLDTIRKNYYNPSRPEKREIKCPTPGCDGTGHVTGLYPHHRSLSGCPHKDRIPPEILAMHENVLKCPTPGCTGQGHVNSNRNTHRSLSGCPIAAAEKLAKSHEKQQPQAGDPSKSSSNSDRILRPMCFVKQLEIPQYGSYRPNMVPATPRANLAKELEKYSKVTFDYASFDAQVFGKRMLAPKIQTSETSPKAFKSKPFPKASSPSHSPSSSYVKSTSSSSTGFDYSHDAEAAHMAATAILNLSTRCWEMPENLSTKQQDLPSKSMDIEVDENGTLDLSMNKHRKRESTFPSSSSCSSSPSVKSPDVSQRQNSTSATSSTMTSPQSSQTSRQDEWDGPIDYTKPNRQREEEPEESEPAAHSFASSEADEQEVSEENFEERKYPGEVTLTNFKLKFLPKDIKKELLTCPTPGCDGSGHITGNYASHRSLSGCPLADKSLRNLMAAHSADLKCPTPGCDGSGHITGNYASHRSLSGCPRAKKSGIKITPTKDDKEDPELMKCPVPGCVGLGHISGKYASHRSASGCPLAARRQKEGSLNGSSFSWKSLKNEGPTCPTPGCDGSGHANGSFLTHRSLSGCPRATFAGKKGKLSGDEILNTKFKTSDVLENDEEIKQLNKEITELNESNSEMEAAMVKLQSQISTMEKNLKNIEEENKIIEEQNEALFLELSGLSQALIQSLANIRLPHMEPISEQNFDAYVNTLTDMYTNQECYQNPENRDLLESIKQAVKGIQV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0Z3</accession>
    <name>A0A452J0Z3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">RAR related orphan receptor B</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 11">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="11">Belongs to the nuclear hormone receptor family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0Z3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038558.1">
        <property type="protein sequence ID" value="ENSGAGP00000034051.1"/>
        <property type="gene ID" value="ENSGAGG00000024178.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004879">
        <property type="term" value="F:nuclear receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd06968">
        <property type="entry name" value="NR_DBD_ROR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.50.10:FF:000003">
        <property type="entry name" value="Nuclear orphan receptor ROR-beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.50.10">
        <property type="entry name" value="Erythroid Transcription Factor GATA-1, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.565.10">
        <property type="entry name" value="Retinoid X Receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035500">
        <property type="entry name" value="NHR-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044101">
        <property type="entry name" value="NR_DBD_ROR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000536">
        <property type="entry name" value="Nucl_hrmn_rcpt_lig-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001723">
        <property type="entry name" value="Nuclear_hrmn_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003079">
        <property type="entry name" value="ROR_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001628">
        <property type="entry name" value="Znf_hrmn_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013088">
        <property type="entry name" value="Znf_NHR/GATA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45805">
        <property type="entry name" value="NUCLEAR HORMONE RECEPTOR HR3-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45805:SF7">
        <property type="entry name" value="NUCLEAR RECEPTOR ROR-BETA-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00104">
        <property type="entry name" value="Hormone_recep"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00105">
        <property type="entry name" value="zf-C4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01293">
        <property type="entry name" value="RORNUCRECPTR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00398">
        <property type="entry name" value="STRDHORMONER"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00047">
        <property type="entry name" value="STROIDFINGER"/>
    </dbReference>
    <dbReference type="SMART" id="SM00430">
        <property type="entry name" value="HOLI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00399">
        <property type="entry name" value="ZnF_C4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57716">
        <property type="entry name" value="Glucocorticoid receptor-like (DNA-binding domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48508">
        <property type="entry name" value="Nuclear receptor ligand-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51843">
        <property type="entry name" value="NR_LBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00031">
        <property type="entry name" value="NUCLEAR_REC_DBD_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51030">
        <property type="entry name" value="NUCLEAR_REC_DBD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="7 11" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3 11" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10 11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="9 11" id="KW-0675">Receptor</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8 11" id="KW-0804">Transcription</keyword>
    <keyword evidence="6 11" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="5 11" id="KW-0862">Zinc</keyword>
    <keyword evidence="4 11" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="Nuclear receptor" evidence="13">
        <location>
            <begin position="46"/>
            <end position="121"/>
        </location>
    </feature>
    <feature type="domain" description="NR LBD" evidence="14">
        <location>
            <begin position="252"/>
            <end position="486"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1"/>
            <end position="26"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="RuleBase" id="RU004334"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS51030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS51843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034051.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="17">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="498" mass="56757" checksum="62F4CA0E925510F7" modified="2019-05-08" version="1">SFQRPVMSVRDFPLDQAPGEKREPLQGISTQGHCRWAGESTPQIEVIPCKICGDKSSGIHYGVITCEGCKGFFRRSQQNNASYSCSRQRNCLIDRTNRNRCQHCRLQKCLALGMSRDAVKFGRMSKKQRDSLYAEVQKHQQNQEQNRVSKDDSEVLARVYPTSVSSGVSDLDDISLLSDGLLFDFPLTPDGGSTYYNLDLLTSAQPSPDQSSMDVSDAKFIKQESIYELMLEPGLFAHSTLEGGTDPQTSSTENRIAQNVVKSHLETCQYTTEELKRLAWMLYTQEEIRSLQNKSCETMWQQCSLQISNAIQYVVEFAKRIDGFMELCQNDQIILLKAGCLEVLLIRMIRAFNPLNNMVLFEGKYGGMQMFKSLGCDDLINAVFELGRNLCRLQLSDEEIALFTAAVLLSPDRPWLTESKKVQKLQDKIYIALQHEIQKKHSSEDKLSKLPLMKTICNLHLDKLEFFRLLHPETAMNFPPLYKEVFNSELQYSDPRES</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0Z6</accession>
    <name>A0A452J0Z6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">tRNA (32-2'-O)-methyltransferase regulator THADA</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="3">Together with methyltransferase FTSJ1, methylates the 2'-O-ribose of nucleotides at position 32 of the anticodon loop of substrate tRNAs.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the THADA family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0Z6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038537.1">
        <property type="protein sequence ID" value="ENSGAGP00000034034.1"/>
        <property type="gene ID" value="ENSGAGG00000024118.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030488">
        <property type="term" value="P:tRNA methylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056843">
        <property type="entry name" value="THADA-like_TPR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056842">
        <property type="entry name" value="THADA-like_TPR_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019442">
        <property type="entry name" value="THADA/TRM732_DUF2428"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051954">
        <property type="entry name" value="tRNA_methyltransferase_THADA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14387">
        <property type="entry name" value="THADA/DEATH RECEPTOR INTERACTING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14387:SF7">
        <property type="entry name" value="THYROID ADENOMA-ASSOCIATED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10350">
        <property type="entry name" value="DUF2428"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25150">
        <property type="entry name" value="TPR_Trm732"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25151">
        <property type="entry name" value="TPR_Trm732_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0819">tRNA processing</keyword>
    <feature type="domain" description="tRNA (32-2'-O)-methyltransferase regulator THADA-like TPR repeats region" evidence="6">
        <location>
            <begin position="232"/>
            <end position="504"/>
        </location>
    </feature>
    <feature type="domain" description="DUF2428" evidence="5">
        <location>
            <begin position="648"/>
            <end position="948"/>
        </location>
    </feature>
    <feature type="domain" description="tRNA (32-2'-O)-methyltransferase regulator THADA-like C-terminal TPR repeats region" evidence="7">
        <location>
            <begin position="950"/>
            <end position="1053"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00010409"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022694"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00035625"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00035698"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="Pfam" id="PF10350"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="Pfam" id="PF25150"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF25151"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034034.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1067" mass="120663" checksum="64D896FAD09DA363" modified="2019-05-08" version="1">VSSCECLCTEELSDSVLLSLCHGALAMLEWKNGSMSENGEKLLLDIALVLLSLASRLKESNMAVSLSRILAIWTNSALDVLESSSQNLKSSLNGNSNTTGKLLEYVYTHWEHPLDAVRHQAKLIFKNILRIHQATLKGSVVKSDPFFSMLTESLLSLEWHVKGKYASLGCLVECVGVENILTVDRTIPAQILDVMGDQSLAPYASDLLETMFINHKSHLISAFQKNIWINHWHDTWVSPLLLILCEGNPDQTTYVIDYYLPKLLKCSPDSLSYMIKILQASADTLLILSLGSCNTRGALGALMACLRTARAHGHLQFREIMRDDLLSIGYVKQGLVHQHNQVRVDALGLLCETHRSTEILTMEEMQLIQFFITYNLNSQSPAVRQQICSLLKKLFCRIQESSQVLYKLEQNKTKQELIKESAKWDHFATLQQYKDFMSSICDKLFEALFPGSSHPTRFSSLTILGSIAEIFSAPEGEVFDFHKIDPTRVQMLIQCFASTFEEVKVLAFELLMKLYDRLDLQDLENLHLLFQAAMDLSISTKPYDCVTASYLLNFLIHHKGLLNVCLKNQQVADTFQIDEDMSASTMEKNTLAVIKYLLSNLEREIFQAEKSLLQAAASFPMYGRVHCITGALQQLPLNNLTLMTEWRQMVTKLILMSYRLSSVVSPIVQSSSPEGLIPMDNDSETAGRLQMILHEIQPRDTNDYFTQAKIFKEHHRVDFAKLTDDEPVENVCTEIRGKEGQTCDVTAQMVLVCCWRSMKEISLLLGMLCKLLPLQTVSESSDGLIAVEQVKDIGDYFKHHLLQSRHRGAFELAYTGFVMLTEILSRCTNESLRTLPQQWLYNVLEEIKSCDPSSKLCATRRSAGIPFYIQALLASEPKNKTNLLKMTMNELVSLAVPVNESLNIIPQVHALNILRALFRDTRFGENIIPYIADGIQAAVLGFTSPVWAVRNSSTLLFSTLITRIFGVKRGKDENSKKNRMTGREFFSRFPTLYPFLLQQLEVVARTMDSETGELKLHPSLFLLLLILGKLYPSPMDGTYSALSMAPFVPFIMRDWEPGWEAVWYKPK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0Z9</accession>
    <name>A0A452J0Z9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">RAR related orphan receptor B</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0Z9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038564.1">
        <property type="protein sequence ID" value="ENSGAGP00000034057.1"/>
        <property type="gene ID" value="ENSGAGG00000024178.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004879">
        <property type="term" value="F:nuclear receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd06968">
        <property type="entry name" value="NR_DBD_ROR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.565.10:FF:000005">
        <property type="entry name" value="Nuclear orphan receptor ROR-beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.50.10:FF:000003">
        <property type="entry name" value="Nuclear orphan receptor ROR-beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.50.10">
        <property type="entry name" value="Erythroid Transcription Factor GATA-1, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.565.10">
        <property type="entry name" value="Retinoid X Receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035500">
        <property type="entry name" value="NHR-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044101">
        <property type="entry name" value="NR_DBD_ROR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000536">
        <property type="entry name" value="Nucl_hrmn_rcpt_lig-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001723">
        <property type="entry name" value="Nuclear_hrmn_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003079">
        <property type="entry name" value="ROR_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001628">
        <property type="entry name" value="Znf_hrmn_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013088">
        <property type="entry name" value="Znf_NHR/GATA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45805">
        <property type="entry name" value="NUCLEAR HORMONE RECEPTOR HR3-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45805:SF7">
        <property type="entry name" value="NUCLEAR RECEPTOR ROR-BETA-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00104">
        <property type="entry name" value="Hormone_recep"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00105">
        <property type="entry name" value="zf-C4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01293">
        <property type="entry name" value="RORNUCRECPTR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00398">
        <property type="entry name" value="STRDHORMONER"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00047">
        <property type="entry name" value="STROIDFINGER"/>
    </dbReference>
    <dbReference type="SMART" id="SM00430">
        <property type="entry name" value="HOLI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00399">
        <property type="entry name" value="ZnF_C4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57716">
        <property type="entry name" value="Glucocorticoid receptor-like (DNA-binding domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48508">
        <property type="entry name" value="Nuclear receptor ligand-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51843">
        <property type="entry name" value="NR_LBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51030">
        <property type="entry name" value="NUCLEAR_REC_DBD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="7" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="9" id="KW-0675">Receptor</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0804">Transcription</keyword>
    <keyword evidence="6" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <keyword evidence="4" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="Nuclear receptor" evidence="12">
        <location>
            <begin position="3"/>
            <end position="78"/>
        </location>
    </feature>
    <feature type="domain" description="NR LBD" evidence="13">
        <location>
            <begin position="211"/>
            <end position="449"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="88"/>
            <end position="108"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS51030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS51843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034057.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="16">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="461" mass="52740" checksum="E7C5F528FD93F5FA" modified="2019-05-08" version="1">MRPTSQKICGDKSSGIHYGVITCEGCKGFFRRSQQNNASYSCSRQRNCLIDRTNRNRCQHCRLQKCLALGMSRDAVKFGRMSKKQRDSLYAEVQKHQQNQEQNRVSKDDSEVLARVYPTSVSSGVSDLDDISLLSDGLLFDFPLTPDGGSTYYNLDLLTSAQPSPDQSSMDVSDAKFIKQESIYELMLEPGLFAHSTLEGSQLASDISMLEIDRIAQNVVKSHLETCQYTTEELKRLAWMLYTQEEIRSLQNKSCETMWQQCSLQISNAIQYVVEFAKRIDGFMELCQNDQIILLKAGCLEVLLIRMIRAFNPLNNMVLFEGKYGGMQMFKSLGCDDLINAVFELGRNLCRLQLSDEEIALFTAAVLLSPDRPWLTESKKVQKLQDKIYIALQHEIQKKHSSEDKLSKMISKLPLMKTICNLHLDKLEFFRLLHPETAMNFPPLYKEVFNSELQYSDPRES</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J194</accession>
    <name>A0A452J194_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="19">Glutamate receptor</fullName>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="19">Receptor for glutamate that functions as a ligand-gated ion channel in the central nervous system and plays an important role in excitatory synaptic transmission. L-glutamate acts as an excitatory neurotransmitter at many synapses in the central nervous system.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>Ca(2+)(in) = Ca(2+)(out)</text>
            <dbReference type="Rhea" id="RHEA:29671"/>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="14 19">Postsynaptic cell membrane</location>
            <topology evidence="14 19">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="19">Belongs to the glutamate-gated ion channel (TC 1.A.10.1) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J194"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038675.1">
        <property type="protein sequence ID" value="ENSGAGP00000034152.1"/>
        <property type="gene ID" value="ENSGAGG00000024221.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015276">
        <property type="term" value="F:ligand-gated monoatomic ion channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0038023">
        <property type="term" value="F:signaling receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd06382">
        <property type="entry name" value="PBP1_iGluR_Kainate"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.2300:FF:000010">
        <property type="entry name" value="Glutamate ionotropic receptor kainate type subunit 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.190.10:FF:000240">
        <property type="entry name" value="Glutamate receptor ionotropic, kainate 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.70:FF:000010">
        <property type="entry name" value="Putative glutamate receptor ionotropic kainate 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.70">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.2300">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.190.10">
        <property type="entry name" value="Periplasmic binding protein-like II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001828">
        <property type="entry name" value="ANF_lig-bd_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019594">
        <property type="entry name" value="Glu/Gly-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001508">
        <property type="entry name" value="Iono_Glu_rcpt_met"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015683">
        <property type="entry name" value="Ionotropic_Glu_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001320">
        <property type="entry name" value="Iontro_rcpt_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028082">
        <property type="entry name" value="Peripla_BP_I"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18966">
        <property type="entry name" value="IONOTROPIC GLUTAMATE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01094">
        <property type="entry name" value="ANF_receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00060">
        <property type="entry name" value="Lig_chan"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10613">
        <property type="entry name" value="Lig_chan-Glu_bd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00177">
        <property type="entry name" value="NMDARECEPTOR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00079">
        <property type="entry name" value="PBPe"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53822">
        <property type="entry name" value="Periplasmic binding protein-like I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53850">
        <property type="entry name" value="Periplasmic binding protein-like II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 19" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="8 18" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13 19" id="KW-0407">Ion channel</keyword>
    <keyword evidence="6 19" id="KW-0406">Ion transport</keyword>
    <keyword evidence="12 19" id="KW-1071">Ligand-gated ion channel</keyword>
    <keyword evidence="7 19" id="KW-0472">Membrane</keyword>
    <keyword evidence="11 19" id="KW-0628">Postsynaptic cell membrane</keyword>
    <keyword evidence="9 19" id="KW-0675">Receptor</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="19" id="KW-0732">Signal</keyword>
    <keyword evidence="5 19" id="KW-0770">Synapse</keyword>
    <keyword evidence="3 19" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 19" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="1 19" id="KW-0813">Transport</keyword>
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        <location>
            <begin position="1"/>
            <end position="19"/>
        </location>
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            <begin position="20"/>
            <end position="827"/>
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            <position position="458"/>
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            <name>L-glutamate</name>
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    <sequence length="827" mass="92617" checksum="7A3A9E79C17924A5" modified="2019-05-08" version="1">MLLFYLLASTGGIFETVENEPVNVEELAFKFAVTNINRNKTLMPNTTLTYDIQRINLFDSFEASRRACDQLALGVAALFGPSHSSSVSAVQSICNALEVPHIQTRWKHPTVDNKDTFYINLYPDYAAISRAILDLVLYYNWKIVTVVYEDSTGLIRLQELIKAPSRYNIKIKIRQLPSGNKDARPLLKEMKKGKEFYVIFDCSHETAAEILKQILSMGMMTEYYHYFFTTLDLFALDLEPYRYSGVNMTGFRLLNIDNPQVTSVIEKWSMERLQAPPKPETGLLDGMMTTEAALMYDAVYMVAVASQRASQMTVSSLQCHRHKPWRFGPRFMNLIKEARWDGLTGHITFNKTDGLRKDFDLDIISLKEEGTEKIGVWNSNSGLNMTDSNKDRSTNITDSLANRTLIVTTILVSPAQLMVQSEETDSGTSSPVSNGSYSGEDGIFSNLCSQKADLAVAPLTITYVREKVIDFSKPFMTLGISILYRKPNGTNPGVFSFLNPLSPDIWMYVLLACLGVSCVLFVIARFTPYEWYNPHPCNPDSDVVENNFTLLNSFWFGVGALMQQGSELMPKALSTRIVGGIWWFFTLIIISSYTANLAAFLTVERMESPIDSADDLAKQTKIEYGAVRDGSTMTFFKKSKISTYEKMWAFMSSRQQTALVKNNDEGIQRVLTTDYALLMESTSIEYVTQRNCNLTQIGGLIDSKGYGVGTPIGSPYRDKITIAILQLQEEGKLHMMKEKWWRGNGCPEEDSKEASALGVENIGGIFIVLAAGLVLSVFVAIGEFIYKSRKNSDIEQVSCLLRAVPLRVYSGSRAVESLPSISGTKAT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J197</accession>
    <name>A0A452J197_9SAUR</name>
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        <recommendedName>
            <fullName evidence="12">ARID domain-containing protein</fullName>
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        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
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        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
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        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
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        <citation type="submission" date="2025-08" db="UniProtKB">
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            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
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    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
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        </citation>
        <scope>IDENTIFICATION</scope>
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    <keyword evidence="8" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="2" id="KW-0488">Methylation</keyword>
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    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
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    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0805">Transcription regulation</keyword>
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        </source>
    </evidence>
    <sequence length="1343" mass="145291" checksum="3ED72A6B752E7F72" modified="2019-05-08" version="1">PPFYSAVSASGSTSSQGEQSNPAQSPFSPHASPHLSGIPGGPSPSPVGSPVGSNQSRSGPISPASIPGSQMPPQPPGSQSESSSHPALSQSPMPQDRGIQRNPQISQYGPQQTGPSMSPHPSPGGQMHPGIGSFQQSNSSGTYGPQMSQYGPQGNYSRPSTYSGVANTSYSGPGPGMGINANSQIHGQGPSQPCGTMPLGRMPSAGMQSRPFPGNMGSMTPNSPGMSLQGGPGMGPPMPTVNRKAQEAAAAVMQAAANSAQSRQGSFPGMNQSGIMGSSSPYTQPMNNSSGLMNPQAPPYSMAPNMVNSSTAPMGLADMMTPGESKLPLPLKADGKEEGPSQPESKSKKSSSSTTTGEKITKVYELGSEPERKMWVDRYLTFMEERGTPVASLPAVGKKPLDLFRLYVCVKEIGGLAQVNKNKKWRELATNLNVGTSSSAASSLKKQYIQYLFAFECKIERGEEPPPEVFSTGDTKKQAKIQPPSPANSGSLQGPQTPQSTGSNSMTEVPGDLKPPTPASTPHGQMTPMQGGRNSAVSVHDPFSDVSDSAFPKRNSMTPNAPYQQGMNMPDMMGRMPYEPNKDPFGGIRKVPGSSEPFMAPGQMPNSGIQDIYNQTPSGAMSTMSMSQRQQFSYGGGYDRRSDHGLGPEGSMGPPGQSNMVPSNSDPSMYSPNRYPGQQRHEPYGQQYPGQGPPSGQPPYGGHQPGMYPQQQVKEVESFIFMTFIFSPLESTASNIFFFSHSPVTPEAWRVMMSLKSGLLAESTWALDTINILLYDDSTVATFNLSQLSGFLELLVEYFRKCLIDIFGILMEYEVGDPGHKALDHNAVKKDDTQSLADEEEEEEDDIEKIEGEEKSTVFATPSAIADPSERPKQASKFDKLPIKIVKKNNLFVIDRSDKLGRVQEFNSGLLHWQLGGGDTTEHIQTHFESKMEIPPRRRAPPPLNSPSKKKDLEGKGEPEEQQEKSITATIDDVLSARPGALSEDSNPGSQTESSKFPFGIHQAKSHRNIKLLEDEPRSRDETPLCTITHWQDSLAKRCICVSNIVRSLSFVPGNDTEMSKHPGLVLILGKLILLHHEHPERKRAPQTYEKEEEEDKGVACSKDEWWWDCLEVLRDNTLVTLANISGQLDLSAYTESICLPILDGLLHWMVCPSAEAQDPFPTVGPNSVLSPQRLVLETLCKLSIQDNNVDLILATPPFSRQEKLYATLVRYVGDRKNPVCREMSMALLSNLAQGDMLAARAIAVQKGSIGNLISFLEDGVTMAQYQQSQHNLMHMQPPPLEPPSVDMMCRAAKALLAMARVDENRSEFLLHEGRLLDISISSVLNSLVASVICDVLFQIGQL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J198</accession>
    <name>A0A452J198_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="19">Glutamate receptor</fullName>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="19">Receptor for glutamate that functions as a ligand-gated ion channel in the central nervous system and plays an important role in excitatory synaptic transmission. L-glutamate acts as an excitatory neurotransmitter at many synapses in the central nervous system.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>Ca(2+)(in) = Ca(2+)(out)</text>
            <dbReference type="Rhea" id="RHEA:29671"/>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="14 19">Postsynaptic cell membrane</location>
            <topology evidence="14 19">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="19">Belongs to the glutamate-gated ion channel (TC 1.A.10.1) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J198"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038672.1">
        <property type="protein sequence ID" value="ENSGAGP00000034149.1"/>
        <property type="gene ID" value="ENSGAGG00000024221.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015276">
        <property type="term" value="F:ligand-gated monoatomic ion channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0038023">
        <property type="term" value="F:signaling receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd06382">
        <property type="entry name" value="PBP1_iGluR_Kainate"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.2300:FF:000010">
        <property type="entry name" value="Glutamate ionotropic receptor kainate type subunit 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.190.10:FF:000240">
        <property type="entry name" value="Glutamate receptor ionotropic, kainate 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.70:FF:000010">
        <property type="entry name" value="Putative glutamate receptor ionotropic kainate 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.70">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.2300">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.190.10">
        <property type="entry name" value="Periplasmic binding protein-like II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001828">
        <property type="entry name" value="ANF_lig-bd_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019594">
        <property type="entry name" value="Glu/Gly-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001508">
        <property type="entry name" value="Iono_Glu_rcpt_met"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015683">
        <property type="entry name" value="Ionotropic_Glu_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001320">
        <property type="entry name" value="Iontro_rcpt_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028082">
        <property type="entry name" value="Peripla_BP_I"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18966">
        <property type="entry name" value="IONOTROPIC GLUTAMATE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01094">
        <property type="entry name" value="ANF_receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00060">
        <property type="entry name" value="Lig_chan"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10613">
        <property type="entry name" value="Lig_chan-Glu_bd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00177">
        <property type="entry name" value="NMDARECEPTOR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00079">
        <property type="entry name" value="PBPe"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53822">
        <property type="entry name" value="Periplasmic binding protein-like I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53850">
        <property type="entry name" value="Periplasmic binding protein-like II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 19" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="8 18" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13 19" id="KW-0407">Ion channel</keyword>
    <keyword evidence="6 19" id="KW-0406">Ion transport</keyword>
    <keyword evidence="12 19" id="KW-1071">Ligand-gated ion channel</keyword>
    <keyword evidence="7 19" id="KW-0472">Membrane</keyword>
    <keyword evidence="11 19" id="KW-0628">Postsynaptic cell membrane</keyword>
    <keyword evidence="9 19" id="KW-0675">Receptor</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="19" id="KW-0732">Signal</keyword>
    <keyword evidence="5 19" id="KW-0770">Synapse</keyword>
    <keyword evidence="3 19" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 19" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="1 19" id="KW-0813">Transport</keyword>
    <feature type="signal peptide" evidence="19">
        <location>
            <begin position="1"/>
            <end position="19"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5041020999" description="Glutamate receptor" evidence="19">
        <location>
            <begin position="20"/>
            <end position="848"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="508"/>
            <end position="527"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="584"/>
            <end position="606"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="765"/>
            <end position="789"/>
        </location>
    </feature>
    <feature type="domain" description="Ionotropic glutamate receptor C-terminal" evidence="20">
        <location>
            <begin position="419"/>
            <end position="746"/>
        </location>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="461"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="463"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="468"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="634"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="635"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="683"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="site" description="Crucial to convey clamshell closure to channel opening" evidence="17">
        <location>
            <position position="613"/>
        </location>
    </feature>
    <feature type="site" description="Interaction with the cone snail toxin Con-ikot-ikot" evidence="17">
        <location>
            <position position="640"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="18">
        <location>
            <begin position="68"/>
            <end position="319"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="18">
        <location>
            <begin position="695"/>
            <end position="749"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023257"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00034104"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00036634"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="RuleBase" id="RU367118"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="SMART" id="SM00079"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034149.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="848" mass="95529" checksum="563079B21CD74597" modified="2019-05-08" version="1">MLLFYLLASTGGIFETVENEPVNVEELAFKFAVTNINRNKTLMPNTTLTYDIQRINLFDSFEASRRACDQLALGVAALFGPSHSSSVSAVQSICNALEVPHIQTRWKHPTVDNKDTFYINLYPDYAAISRAILDLVLYYNWKIVTVVYEDSTGLIRLQELIKAPSRYNIKIKIRQLPSGNKDARPLLKEMKKGKEFYVIFDCSHETAAEILKQILSMGMMTEYYHYFFTTLDLFALDLEPYRYSGVNMTGFRLLNIDNPQVTSVIEKWSMERLQAPPKPETGLLDGMMTTEAALMYDAVYMVAVASQRASQMTVSSLQCHRHKPWRFGPRFMNLIKEARWDGLTGHITFNKTDGLRKDFDLDIISLKEEGTEKAAGEVSKNLYKVWKKIGVWNSNSGLNMTDSNKDRSTNITDSLANRTLIVTTILVSPAVEGIKICLLNAKKMRIFSNLCSQKADLAVAPLTITYVREKVIDFSKPFMTLGISILYRKPNGTNPGVFSFLNPLSPDIWMYVLLACLGVSCVLFVIARFTPYEWYNPHPCNPDSDVVENNFTLLNSFWFGVGALMQQGSELMPKALSTRIVGGIWWFFTLIIISSYTANLAAFLTVERMESPIDSADDLAKQTKIEYGAVRDGSTMTFFKKSKISTYEKMWAFMSSRQQTALVKNNDEGIQRVLTTDYALLMESTSIEYVTQRNCNLTQIGGLIDSKGYGVGTPIGSPYRDKITIAILQLQEEGKLHMMKEKWWRGNGCPEEDSKEASALGVENIGGIFIVLAAGLVLSVFVAIGEFIYKSRKNSDIEQAFCFFYGLQCKQTHPSNSTSGTTLSTELDCGKLIREERGIRTQPSIHTV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J199</accession>
    <name>A0A452J199_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="12">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the GPSM family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J199"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038680.1">
        <property type="protein sequence ID" value="ENSGAGP00000034157.1"/>
        <property type="gene ID" value="ENSGAGG00000024269.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000038683.1">
        <property type="protein sequence ID" value="ENSGAGP00000034160.1"/>
        <property type="gene ID" value="ENSGAGG00000024269.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005938">
        <property type="term" value="C:cell cortex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001965">
        <property type="term" value="F:G-protein alpha-subunit binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005092">
        <property type="term" value="F:GDP-dissociation inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000132">
        <property type="term" value="P:establishment of mitotic spindle orientation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000043">
        <property type="entry name" value="G-protein-signaling modulator 2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.10">
        <property type="entry name" value="Tetratricopeptide repeat domain"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003109">
        <property type="entry name" value="GoLoco_motif"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052386">
        <property type="entry name" value="GPSM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011990">
        <property type="entry name" value="TPR-like_helical_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019734">
        <property type="entry name" value="TPR_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45954:SF2">
        <property type="entry name" value="G-PROTEIN-SIGNALING MODULATOR 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45954">
        <property type="entry name" value="LD33695P"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02188">
        <property type="entry name" value="GoLoco"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13374">
        <property type="entry name" value="TPR_10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13424">
        <property type="entry name" value="TPR_12"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13176">
        <property type="entry name" value="TPR_7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00390">
        <property type="entry name" value="GoLoco"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00028">
        <property type="entry name" value="TPR"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48452">
        <property type="entry name" value="TPR-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50877">
        <property type="entry name" value="GOLOCO"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50005">
        <property type="entry name" value="TPR"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="8 10" id="KW-0802">TPR repeat</keyword>
    <feature type="repeat" description="TPR" evidence="10">
        <location>
            <begin position="80"/>
            <end position="113"/>
        </location>
    </feature>
    <feature type="repeat" description="TPR" evidence="10">
        <location>
            <begin position="260"/>
            <end position="293"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="393"/>
            <end position="422"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="461"/>
            <end position="485"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="622"/>
            <end position="641"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="655"/>
            <end position="677"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="663"/>
            <end position="677"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00006600"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022803"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00339"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034157.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="677" mass="75988" checksum="749CA7218D40C7AB" modified="2019-05-08" version="1">MPWIKIDRLLQRPKSLWKFCRSCVLPASPAHRPYRLRMEASCLELALEGERLCKAGDFKAGAAFFEAAVQVGTEDLKTLSAIYSQLGNAYFYLKEYSKALEYHKHDLTLARTIGDRIGEAKASGNLGNTLKILGQFDEAIVCCQRHLEISQEQGDKVGEARALYNIGNVYHAKGKQLSWSMAQDPGSLPPEVKETLQRASEYYERNLSLVKELGDRAAQGRAYGNLGNTQYLLGNFNEATAFHKERLAIAKEFGDKAAERRAYSNLGNAHIFLGRFDISAEYYKKTLQLSRQLKDQAVEAQACYSLGNTYTLLQDYERAIDYHLKHLLIAQELGDRVGEGRACWSLGNAYVSLGNHDQALRFARKHLEISQEIGDRNGELTARMNVAQLKSALGDEDGDATHHSSSYEAQGARPKRSQRNSMDSLDLLKFPSEKEQNGDSHHMGNLKITGKDFLSLPVRSRKYQEIQSSSERKSQDSNMSPLDASDVRLQVSQSKINRAPSDEECFFDLLSKFQSSRMDDQRCPLEECQAEAAEAAATSVPALEERISQSSLIASPQTEEFFDLIASSQSRRLDDQRASVGSLPGLRITHNNLGHLRMEGDAQEPGDEFFNMLMKCQSSRIDDQRCAPPDTVPRGPTMPDEDFFSLIQRVQAKRMDEQRVDLASAQEEQQQRRPSSS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIN0</accession>
    <name>A0A452GIN0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Sodium/hydrogen exchanger</fullName>
        </recommendedName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="16">Plasma membrane Na(+)/H(+) antiporter. Exchanges intracellular H(+) ions for extracellular Na(+) in 1:1 stoichiometry, playing a key role in salt and fluid absorption and pH homeostasis. Major apical Na(+)/H(+) exchanger in kidney and intestine playing an important role in renal and intestine Na(+) absorption and blood pressure regulation.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>Na(+)(in) + H(+)(out) = Na(+)(out) + H(+)(in)</text>
            <dbReference type="Rhea" id="RHEA:29419"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Apical cell membrane</location>
            <topology evidence="2">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Early endosome membrane</location>
            <topology evidence="3">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Recycling endosome membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="18">Belongs to the monovalent cation:proton antiporter 1 (CPA1) transporter (TC 2.A.36) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIN0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001642.1">
        <property type="protein sequence ID" value="ENSGAGP00000001448.1"/>
        <property type="gene ID" value="ENSGAGG00000000884.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031526">
        <property type="term" value="C:brush border membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031901">
        <property type="term" value="C:early endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055038">
        <property type="term" value="C:recycling endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015386">
        <property type="term" value="F:potassium:proton antiporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015385">
        <property type="term" value="F:sodium:proton antiporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051453">
        <property type="term" value="P:regulation of intracellular pH"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098719">
        <property type="term" value="P:sodium ion import across plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.1330">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018422">
        <property type="entry name" value="Cation/H_exchanger_CPA1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006153">
        <property type="entry name" value="Cation/H_exchanger_TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018410">
        <property type="entry name" value="Na/H_exchanger_3/5"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004709">
        <property type="entry name" value="NaH_exchanger"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00840">
        <property type="entry name" value="b_cpa1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10110">
        <property type="entry name" value="SODIUM/HYDROGEN EXCHANGER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10110:SF90">
        <property type="entry name" value="SODIUM_HYDROGEN EXCHANGER 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00999">
        <property type="entry name" value="Na_H_Exchanger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01084">
        <property type="entry name" value="NAHEXCHNGR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01087">
        <property type="entry name" value="NAHEXCHNGR3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 18" id="KW-0050">Antiport</keyword>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="10" id="KW-0967">Endosome</keyword>
    <keyword evidence="13 18" id="KW-0406">Ion transport</keyword>
    <keyword evidence="14 20" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0732">Signal</keyword>
    <keyword evidence="12" id="KW-0915">Sodium</keyword>
    <keyword evidence="15 18" id="KW-0739">Sodium transport</keyword>
    <keyword evidence="8 18 20" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 20" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="4 18" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="21"/>
            <end position="39"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="51"/>
            <end position="68"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="80"/>
            <end position="103"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="115"/>
            <end position="137"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="144"/>
            <end position="168"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="188"/>
            <end position="212"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="305"/>
            <end position="328"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="340"/>
            <end position="361"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="373"/>
            <end position="393"/>
        </location>
    </feature>
    <feature type="domain" description="Cation/H+ exchanger transmembrane" evidence="21">
        <location>
            <begin position="4"/>
            <end position="399"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="601"/>
            <end position="699"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="19">
        <location>
            <begin position="614"/>
            <end position="629"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="656"/>
            <end position="684"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="685"/>
            <end position="699"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004195"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004424"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004520"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022449"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023053"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023201"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00045831"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00047524"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="RuleBase" id="RU003722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="Pfam" id="PF00999"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001448.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="759" mass="85586" checksum="BA393E92224F861D" modified="2019-05-08" version="1">SLILSLLFFHLSHKVTRVVPESALLIVLGLFLGGIVWAADHIASFTLTPNVFFFYLLPPIVLDAGYFMPNKLFFGNLGTILLYAVIGTVWNAATTGMSLYGVYLSGIMGQLDIGLLDFLLFGSLIAAVDPVAVLAVFEEVHVNEVLFIIVFGESLLNDAVTVVLYNVFDSFVAMGADNVTGVDCVKGIVSFFVVSLGGTFVGIVFAFLLSLVTRFTKHVRIIEPGFVFVISYLSYLTAEMLSLSAILAITFCGVCCQKYVKANICDQSSTTVRYTMKMLASGSETIIFMFLGISAVNPDIWTWNTAFILLTLVFISVYRVIGVVIQTWVLNHYRVVQLEIIDQVVMSYGGLRGAVAFALVALLDEKKVKEKNLFVSTTIIVVFFTVIFQGLTIKPLVQWLKVKKSEHREPKLNEKLHGRAFDHILSAVEDISGQIGHNYLRDKWSNFDRKILSKVLMRRSAQKSRDQILNIFHELNLKDAISYVAEGERRGSLAFIRSSSTDNMVNVDFSTPRPSTVEASVSCLLREKASSVCLDMQALEQRRKSIRDTEDTVTHHTLQQYLYKPRQEHIHLYSRHEIIQNEDEKQDKEIFHRTMRKRLESFKSTKLGINQPKKLNKSHKRDRGQKRRNSNAIPNGKIPSENPAPDFTLKEKEVSDTNNKPPVNSGYCSETSYEPVNSQDNSVDMQERLEESNPKENQEKKTLLNAQAHYRSAAARRVQSRRSFHRPKPLQLPKLQKLASFPSSCHIPPHSLEQEETQL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIN5</accession>
    <name>A0A452GIN5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Amino acid transporter transmembrane domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the amino acid/polyamine transporter 2 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIN5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001649.1">
        <property type="protein sequence ID" value="ENSGAGP00000001454.1"/>
        <property type="gene ID" value="ENSGAGG00000001158.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015179">
        <property type="term" value="F:L-amino acid transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013057">
        <property type="entry name" value="AA_transpt_TM"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22950">
        <property type="entry name" value="AMINO ACID TRANSPORTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22950:SF646">
        <property type="entry name" value="SODIUM-COUPLED NEUTRAL AMINO ACID TRANSPORTER 10-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01490">
        <property type="entry name" value="Aa_trans"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0029">Amino-acid transport</keyword>
    <keyword evidence="7 9" id="KW-0472">Membrane</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 9" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 9" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="12"/>
            <end position="31"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="37"/>
            <end position="60"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="85"/>
            <end position="106"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="126"/>
            <end position="144"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="156"/>
            <end position="175"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="227"/>
            <end position="254"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="274"/>
            <end position="292"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="347"/>
            <end position="368"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="380"/>
            <end position="397"/>
        </location>
    </feature>
    <feature type="domain" description="Amino acid transporter transmembrane" evidence="10">
        <location>
            <begin position="8"/>
            <end position="378"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="430"/>
            <end position="686"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="725"/>
            <end position="797"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="872"/>
            <end position="900"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="920"/>
            <end position="1048"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="441"/>
            <end position="453"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="467"/>
            <end position="487"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="546"/>
            <end position="598"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="634"/>
            <end position="647"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="663"/>
            <end position="686"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="725"/>
            <end position="745"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="755"/>
            <end position="778"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="876"/>
            <end position="900"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="972"/>
            <end position="983"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="992"/>
            <end position="1002"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="1006"/>
            <end position="1015"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="1019"/>
            <end position="1029"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="1033"/>
            <end position="1048"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008066"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022970"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="Pfam" id="PF01490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001454.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1156" mass="127053" checksum="1C9BFCD425E90D40" modified="2019-05-08" version="1">MASSAAAGSNWGLIMNIVNSIVGVSVLTMPFCFRQCGIVLGALLLIFCSWMTHQSCMFLVKSANLSKRRTYPGLAFHAYGKAGKMLVETSMIGLMLGTCIAFYVVIGDLGSNFFARLLGLQVTGSFRIILLFAVSLCIVLPLSLQRNMMASIQSFSAMALIFYTVFMFVIVLSSFKHGLFGGQWLKRISYMRWEGIFRCIPIFGMSFACQSQVLPTYDSLDEPSVKIMSSIFASSLNVVTTFYVTVGFFGYVSYTEAIAGNVLMNFPSNLVTEMIRVGFMMSVAVGFPMMILPCRQALNTLLFEQQQKDGTFAAGGYMPPLRFKALTLTIVFGTMVGGIMIPNVETVLGLTGATMGSLICFICPALIYRKIHKNVLCSQLILGIGLGILVISTYTTLSVTEDMPVKMEAVELADPLKEAKNVIDQAQVQIPVQDPAVEGAEDSREKPKLPNEKEEMEQPQIKVPMDVPKRDEAKAKQQEEVQLDRPDQGIALPVGEAHRHEPPVPHDAVVVDEGQDREESEENKSPPEEANANRLKPAMEKAAQLDPKREEPNLEPGKEDGAQREFDRPLEDLKNAPEVRRDGRPLYKDLEPGKRVVEPRTLAVLPGGEKRAEAAGGRENANLQQPGNVAEAVKSIEKEIDPGEKQELLPPARLDPAQAQQPEQREIRNAEDKPQENADSKLEEAGKAKALDHAVLLQVIKEQQVQQKQLLDQQAKLLAVIEEQHKEMHQQRQEEGGDEKPKQETPQEADLPLAKSKEEEGRDVKADVAAKHLPKEILEPPQPGPGNQPVDLAEQHQRNVDDLEEAVHRPRMKFLVQVSVAGPKIPEALPKGGKVPQHKANKVVKNSLENLGDPLHSDNSRQANDIGHAVQAAGNHPEKPAAKELGGGHEGKAVGDSQEKKGALEAGEVVAGAVADIQRELPRAVKGQEVAGKGPERVFGVDSKTKPANKEEPEDLELPLDSSNKLQVAENMQHKQEHLRLGNDDQADGAGSKQQQASQVNQAGKVEAKKEDFRGKTVNVEQGPQGSHLNKQKVLDEKLPKDPAEKDAFENDIALKREKGAALLADQKPDPENVKPNRDLKLQADLDLRRRRRDLPPRLEEETEASEGVIIGLNPLPDVKVNDLRSALETRLNQAAEGAQQVVHSRQIKQMAEEEV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIN7</accession>
    <name>A0A452GIN7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Armadillo repeat-containing protein 8</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIN7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001733.1">
        <property type="protein sequence ID" value="ENSGAGP00000001519.1"/>
        <property type="gene ID" value="ENSGAGG00000001203.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034657">
        <property type="term" value="C:GID complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043161">
        <property type="term" value="P:proteasome-mediated ubiquitin-dependent protein catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000061">
        <property type="entry name" value="armadillo repeat-containing protein 8 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000070">
        <property type="entry name" value="armadillo repeat-containing protein 8 isoform X1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000225">
        <property type="entry name" value="Armadillo"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038739">
        <property type="entry name" value="ARMC8/Vid28"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15651">
        <property type="entry name" value="ARMADILLO REPEAT-CONTAINING PROTEIN 8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15651:SF7">
        <property type="entry name" value="ARMADILLO REPEAT-CONTAINING PROTEIN 8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00514">
        <property type="entry name" value="Arm"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00185">
        <property type="entry name" value="ARM"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50176">
        <property type="entry name" value="ARM_REPEAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0539">Nucleus</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <feature type="repeat" description="ARM" evidence="7">
        <location>
            <begin position="61"/>
            <end position="97"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00013746"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00259"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001519.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="642" mass="71736" checksum="574FA889D4E1419C" modified="2019-05-08" version="1">MACVLETPLRMSVLSEVTASSRHYVDRLFDPDPQKVLQGVIDMKNAVIGNNKQKANLIVLGAVPRLLYLLQQETSSTELRTECAVVLGSLAMGTENNVKSLLDCHIIPALLQGLLSPELKFIEACLRCLRTIFTSPVTPEELLYTGPDHQTILFNHGAVQNIAHLLSSTSYKVRMQALKCFSVLAFENPQVSMTLVNVLVDGELLPQIFVKMLQRDKPIEMQLTSAKCLTYMCRAGAIQTDDNCIVLKALPSLVRMCSKERLLEERVEGAETLAYLIETDVELQRIASITDHLIAMLSDYFKYPSSVSAITDIKRLDHDLKHAHELRQAAFKLYASLGANDEDIRKKIIETENMMDRIVNGLSESSIKVRLSAVRCLHSLSRSVQQLRTSFQDHAVWKPLMKVLQNAPDEVLVVASSTLCNLLLEFSPSKEPILESGAIELLCSLTQSENLALRVNGIWALMNMAFQAEQKIKADILQGLSTEQLFQLLSDSDVNVLMKTLGLLRNLLSTRPHIDQIMSTHGKQIMQAVTLILEGEHNIEVKEQTLCILANIADGTTAKELIMTNDDILQKIKYYMSHSNAKLQLAAMFCISNLIWNEEEGSQERQDKLRDIGVVDILHKLSQSSDPNLCDKAKTALQQYLA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIN8</accession>
    <name>A0A452GIN8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Septin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="3 4">Belongs to the TRAFAC class TrmE-Era-EngA-EngB-Septin-like GTPase superfamily. Septin GTPase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIN8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001659.1">
        <property type="protein sequence ID" value="ENSGAGP00000001458.1"/>
        <property type="gene ID" value="ENSGAGG00000001174.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005525">
        <property type="term" value="F:GTP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="CDD" id="cd01850">
        <property type="entry name" value="CDC_Septin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000036">
        <property type="entry name" value="septin-6 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR030379">
        <property type="entry name" value="G_SEPTIN_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016491">
        <property type="entry name" value="Septin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18884">
        <property type="entry name" value="SEPTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00735">
        <property type="entry name" value="Septin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF006698">
        <property type="entry name" value="Septin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51719">
        <property type="entry name" value="G_SEPTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="2 4" id="KW-0342">GTP-binding</keyword>
    <keyword evidence="1 4" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Septin-type G" evidence="7">
        <location>
            <begin position="38"/>
            <end position="304"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="404"/>
            <end position="433"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="5">
        <location>
            <begin position="320"/>
            <end position="398"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="404"/>
            <end position="416"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PIRNR" id="PIRNR006698"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="RuleBase" id="RU004560"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS51719"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001458.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="433" mass="49485" checksum="8E964E0E62A6F22E" modified="2019-05-08" version="1">MAAAEIARQGEGCRTVPLSGHVGFDSLPDQLVNKSVNHGFCFNILCVGETGLGKSTLMDTLFNTKFEGETASHSQPGVQLKSSTYDLQESNVGLKLTVVGTVGFGDQINKEDSYKPIVEFIDAQFEAYLQEELKIKRVLHNFHDSRIHACLYFIAPTGHSLKSLDLVTMKKLDSKVNIIPIIAKSDAISKSELTKFKIKITSELVSNGVQIYQFPTDDESVAEINGTMNAHLPFAVIGSTEELKIGNKMMKARQYPWGTVQVENEAHCDFVKLREMLIRVNMEDLREQTHTRHYELYRRCKLEEMGFKDTDPDSKPFSLQETYEAKRNEFLGELQKKEEEMRQMFVQRVKEKEAELKEAEKELHEKFDRLKKLHQDEKKKLEDKKKSLDDEVNAFKQRKTAAELLQSQAQQAGGSQTLKREKERKNNPWLCTE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIN9</accession>
    <name>A0A452GIN9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Topoisomerase 6 subunit A/Spo11 TOPRIM domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="3">
            <text>ATP-dependent breakage, passage and rejoining of double-stranded DNA.</text>
            <dbReference type="EC" id="5.6.2.2"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the TOP6A family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIN9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001747.1">
        <property type="protein sequence ID" value="ENSGAGP00000001531.1"/>
        <property type="gene ID" value="ENSGAGG00000001218.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000228">
        <property type="term" value="C:nuclear chromosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003918">
        <property type="term" value="F:DNA topoisomerase type II (double strand cut, ATP-hydrolyzing) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042138">
        <property type="term" value="P:meiotic DNA double-strand break formation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000706">
        <property type="term" value="P:meiotic DNA double-strand break processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007131">
        <property type="term" value="P:reciprocal meiotic recombination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00223">
        <property type="entry name" value="TOPRIM_TopoIIB_SPO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.1360.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013048">
        <property type="entry name" value="Meiotic_Spo11"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002815">
        <property type="entry name" value="Spo11/TopoVI_A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036078">
        <property type="entry name" value="Spo11/TopoVI_A_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034136">
        <property type="entry name" value="TOPRIM_Topo6A/Spo11"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10848">
        <property type="entry name" value="MEIOTIC RECOMBINATION PROTEIN SPO11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10848:SF0">
        <property type="entry name" value="MEIOTIC RECOMBINATION PROTEIN SPO11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21180">
        <property type="entry name" value="TOP6A-Spo11_Toprim"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01551">
        <property type="entry name" value="SPO11HOMOLOG"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01550">
        <property type="entry name" value="TOP6AFAMILY"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56726">
        <property type="entry name" value="DNA topoisomerase IV, alpha subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS52041">
        <property type="entry name" value="TOPO_IIB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0413">Isomerase</keyword>
    <keyword evidence="2" id="KW-0539">Nucleus</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0799">Topoisomerase</keyword>
    <feature type="domain" description="Topoisomerase 6 subunit A/Spo11 TOPRIM" evidence="4">
        <location>
            <begin position="80"/>
            <end position="222"/>
        </location>
    </feature>
    <feature type="active site" description="O-(5'-phospho-DNA)-tyrosine intermediate" evidence="3">
        <location>
            <position position="48"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01385"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="Pfam" id="PF21180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001531.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="228" mass="26193" checksum="5C3D457826B87FB0" modified="2019-05-08" version="1">MISHYTTRKIKSDCPKSALKFALMLKILSVIYKMVQNNTYATKRDIYYSDTLLFGSQSVVDNIVNDISCMLKIPRRSLHIGKGVPDLNTRLLVRKLWDTCHIPIFTLMDADPHGIEIMCIYKYGSVSMSFEAHHLTVPAIRWLGLLPSDIERLNIRKDALIPLTKQDQNKLTSMQKRPYMACQPMWKKEMEIMAASKMKAEIQALTSLSSDYLSRVYLPSKLQFGGWI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIQ2</accession>
    <name>A0A452GIQ2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Septin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="3 4">Belongs to the TRAFAC class TrmE-Era-EngA-EngB-Septin-like GTPase superfamily. Septin GTPase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIQ2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001686.1">
        <property type="protein sequence ID" value="ENSGAGP00000001478.1"/>
        <property type="gene ID" value="ENSGAGG00000001174.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005525">
        <property type="term" value="F:GTP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="CDD" id="cd01850">
        <property type="entry name" value="CDC_Septin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000036">
        <property type="entry name" value="septin-6 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR030379">
        <property type="entry name" value="G_SEPTIN_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016491">
        <property type="entry name" value="Septin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18884">
        <property type="entry name" value="SEPTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00735">
        <property type="entry name" value="Septin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF006698">
        <property type="entry name" value="Septin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51719">
        <property type="entry name" value="G_SEPTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="2 4" id="KW-0342">GTP-binding</keyword>
    <keyword evidence="1 4" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Septin-type G" evidence="7">
        <location>
            <begin position="39"/>
            <end position="305"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="405"/>
            <end position="427"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="5">
        <location>
            <begin position="321"/>
            <end position="399"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="405"/>
            <end position="417"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PIRNR" id="PIRNR006698"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="RuleBase" id="RU004560"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS51719"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001478.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="427" mass="48741" checksum="EBB7DBAC769309B9" modified="2019-05-08" version="1">MAAAEIARQVGEGCRTVPLSGHVGFDSLPDQLVNKSVNHGFCFNILCVGETGLGKSTLMDTLFNTKFEGETASHSQPGVQLKSSTYDLQESNVGLKLTVVGTVGFGDQINKEDSYKPIVEFIDAQFEAYLQEELKIKRVLHNFHDSRIHACLYFIAPTGHSLKSLDLVTMKKLDSKVNIIPIIAKSDAISKSELTKFKIKITSELVSNGVQIYQFPTDDESVAEINGTMNAHLPFAVIGSTEELKIGNKMMKARQYPWGTVQVENEAHCDFVKLREMLIRVNMEDLREQTHTRHYELYRRCKLEEMGFKDTDPDSKPFSLQETYEAKRNEFLGELQKKEEEMRQMFVQRVKEKEAELKEAEKELHEKFDRLKKLHQDEKKKLEDKKKSLDDEVNAFKQRKTAAELLQSQAQQAGGSQTLKREKERKN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJ00</accession>
    <name>A0A452GJ00_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="26">Histone-lysine N-methyltransferase 2C</fullName>
            <ecNumber evidence="22">2.1.1.364</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="34 35">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="35" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="34" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="34" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="24">Histone methyltransferase that catalyzes methyl group transfer from S-adenosyl-L-methionine to the epsilon-amino group of 'Lys-4' of histone H3 (H3K4). Part of chromatin remodeling machinery predominantly forms H3K4me1 methylation marks at active chromatin sites where transcription and DNA repair take place. Likely plays a redundant role with KMT2D in enriching H3K4me1 mark on primed and active enhancer elements.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="23">
            <text>L-lysyl(4)-[histone H3] + S-adenosyl-L-methionine = N(6)-methyl-L-lysyl(4)-[histone H3] + S-adenosyl-L-homocysteine + H(+)</text>
            <dbReference type="Rhea" id="RHEA:60264"/>
            <dbReference type="Rhea" id="RHEA-COMP:15543"/>
            <dbReference type="Rhea" id="RHEA-COMP:15547"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29969"/>
            <dbReference type="ChEBI" id="CHEBI:57856"/>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
            <dbReference type="ChEBI" id="CHEBI:61929"/>
            <dbReference type="EC" id="2.1.1.364"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="23">
            <dbReference type="Rhea" id="RHEA:60265"/>
        </physiologicalReaction>
    </comment>
    <comment type="subunit">
        <text evidence="25">Component of the MLL3 complex (also named ASCOM complex), at least composed of catalytic subunit KMT2C/MLL3, ASH2L, RBBP5, WDR5, NCOA6, DPY30, KDM6A, PAXIP1/PTIP, PAGR1 and alpha- and beta-tubulin. Forms a core complex with the evolutionary conserved subcomplex WRAD composed of WDR5, RBBP5, ASH2L/ASH2 and DPY30 subunits; WRAD differentially stimulates the methyltransferase activity. Interacts (via WIN motif) with WDR5.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="2.1.1.364" evidence="22"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001874.1">
        <property type="protein sequence ID" value="ENSGAGP00000001649.1"/>
        <property type="gene ID" value="ENSGAGG00000001223.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0044666">
        <property type="term" value="C:MLL3/4 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016746">
        <property type="term" value="F:acyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140945">
        <property type="term" value="F:histone H3K4 monomethyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003713">
        <property type="term" value="F:transcription coactivator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032259">
        <property type="term" value="P:methylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045944">
        <property type="term" value="P:positive regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd15696">
        <property type="entry name" value="ePHD1_KMT2C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd15697">
        <property type="entry name" value="ePHD2_KMT2C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd22026">
        <property type="entry name" value="HMG-box_KMT2C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd15509">
        <property type="entry name" value="PHD1_KMT2C_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd15594">
        <property type="entry name" value="PHD2_KMT2C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd15511">
        <property type="entry name" value="PHD3_KMT2C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd15596">
        <property type="entry name" value="PHD4_KMT2C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd15513">
        <property type="entry name" value="PHD5_KMT2C_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd15600">
        <property type="entry name" value="PHD6_KMT2C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd19171">
        <property type="entry name" value="SET_KMT2C_2D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.30.10:FF:000009">
        <property type="entry name" value="Histone-lysine N-methyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.170.270.10:FF:000003">
        <property type="entry name" value="Histone-lysine N-methyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Histone-lysine N-methyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Histone-lysine N-methyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Histone-lysine N-methyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Histone-lysine N-methyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.40.10:FF:001142">
        <property type="entry name" value="Histone-lysine N-methyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.40.10:FF:000080">
        <property type="entry name" value="Histone-lysine N-methyltransferase 2C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.40.10:FF:000095">
        <property type="entry name" value="Histone-lysine N-methyltransferase 2C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.360">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.30.10">
        <property type="entry name" value="High mobility group box domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.170.270.10">
        <property type="entry name" value="SET domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034732">
        <property type="entry name" value="EPHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003889">
        <property type="entry name" value="FYrich_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003888">
        <property type="entry name" value="FYrich_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009071">
        <property type="entry name" value="HMG_box_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036910">
        <property type="entry name" value="HMG_box_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000637">
        <property type="entry name" value="HMGI/Y_DNA-bd_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041967">
        <property type="entry name" value="KMT2C_ePHD1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041968">
        <property type="entry name" value="KMT2C_ePHD2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047004">
        <property type="entry name" value="KMT2C_PHD2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047003">
        <property type="entry name" value="KMT2C_PHD4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047005">
        <property type="entry name" value="KMT2C_PHD6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037877">
        <property type="entry name" value="PHD3_KMT2C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003616">
        <property type="entry name" value="Post-SET_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001214">
        <property type="entry name" value="SET_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046341">
        <property type="entry name" value="SET_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011011">
        <property type="entry name" value="Znf_FYVE_PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001965">
        <property type="entry name" value="Znf_PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019787">
        <property type="entry name" value="Znf_PHD-finger"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001841">
        <property type="entry name" value="Znf_RING"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45888:SF1">
        <property type="entry name" value="HISTONE-LYSINE N-METHYLTRANSFERASE 2C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45888">
        <property type="entry name" value="HL01030P-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05965">
        <property type="entry name" value="FYRC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05964">
        <property type="entry name" value="FYRN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00628">
        <property type="entry name" value="PHD"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00856">
        <property type="entry name" value="SET"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13771">
        <property type="entry name" value="zf-HC5HC2H"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13832">
        <property type="entry name" value="zf-HC5HC2H_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00542">
        <property type="entry name" value="FYRC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00541">
        <property type="entry name" value="FYRN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00398">
        <property type="entry name" value="HMG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00249">
        <property type="entry name" value="PHD"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SMART" id="SM00508">
        <property type="entry name" value="PostSET"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="5"/>
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    </dbReference>
    <dbReference type="SUPFAM" id="SSF57903">
        <property type="entry name" value="FYVE/PHD zinc finger"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47095">
        <property type="entry name" value="HMG-box"/>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51805">
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        <property type="match status" value="2"/>
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    <dbReference type="PROSITE" id="PS51543">
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        <property type="entry name" value="FYRN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00354">
        <property type="entry name" value="HMGI_Y"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="POST_SET"/>
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    </dbReference>
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        <property type="entry name" value="ZF_PHD_2"/>
        <property type="match status" value="6"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="12" id="KW-0007">Acetylation</keyword>
    <keyword evidence="17" id="KW-0010">Activator</keyword>
    <keyword evidence="21" id="KW-0012">Acyltransferase</keyword>
    <keyword evidence="11" id="KW-0156">Chromatin regulator</keyword>
    <keyword evidence="14 28" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="15" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="20" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="2" id="KW-0488">Methylation</keyword>
    <keyword evidence="4" id="KW-0489">Methyltransferase</keyword>
    <keyword evidence="19" id="KW-0539">Nucleus</keyword>
    <keyword evidence="16" id="KW-0564">Palmitate</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="35" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0949">S-adenosyl-L-methionine</keyword>
    <keyword evidence="18" id="KW-0804">Transcription</keyword>
    <keyword evidence="13" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="5" id="KW-0808">Transferase</keyword>
    <keyword evidence="10" id="KW-0862">Zinc</keyword>
    <keyword evidence="9 27" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="PHD-type" evidence="33">
        <location>
            <begin position="223"/>
            <end position="327"/>
        </location>
    </feature>
    <feature type="domain" description="PHD-type" evidence="30">
        <location>
            <begin position="337"/>
            <end position="387"/>
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    </feature>
    <feature type="domain" description="PHD-type" evidence="30">
        <location>
            <begin position="384"/>
            <end position="434"/>
        </location>
    </feature>
    <feature type="domain" description="PHD-type" evidence="30">
        <location>
            <begin position="461"/>
            <end position="516"/>
        </location>
    </feature>
    <feature type="domain" description="PHD-type" evidence="30">
        <location>
            <begin position="938"/>
            <end position="991"/>
        </location>
    </feature>
    <feature type="domain" description="PHD-type" evidence="30">
        <location>
            <begin position="988"/>
            <end position="1038"/>
        </location>
    </feature>
    <feature type="domain" description="PHD-type" evidence="30">
        <location>
            <begin position="1065"/>
            <end position="1120"/>
        </location>
    </feature>
    <feature type="domain" description="PHD-type" evidence="33">
        <location>
            <begin position="4350"/>
            <end position="4458"/>
        </location>
    </feature>
    <feature type="domain" description="SET" evidence="31">
        <location>
            <begin position="4722"/>
            <end position="4838"/>
        </location>
    </feature>
    <feature type="domain" description="Post-SET" evidence="32">
        <location>
            <begin position="4846"/>
            <end position="4862"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="29">
        <location>
            <begin position="1"/>
            <end position="120"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="29">
        <location>
            <begin position="163"/>
            <end position="208"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="29">
        <location>
            <begin position="590"/>
            <end position="619"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="29">
        <location>
            <begin position="634"/>
            <end position="678"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="29">
        <location>
            <begin position="809"/>
            <end position="842"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="29">
        <location>
            <begin position="866"/>
            <end position="895"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="29">
        <location>
            <begin position="1200"/>
            <end position="1298"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="29">
        <location>
            <begin position="1357"/>
            <end position="1387"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="29">
        <location>
            <begin position="1415"/>
            <end position="1444"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="29">
        <location>
            <begin position="1551"/>
            <end position="1588"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="29">
        <location>
            <begin position="1666"/>
            <end position="1724"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="29">
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                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
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        <property type="entry name" value="GST_NTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 5" id="KW-0251">Elongation factor</keyword>
    <keyword evidence="4 5" id="KW-0648">Protein biosynthesis</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="GST N-terminal" evidence="9">
        <location>
            <begin position="3"/>
            <end position="88"/>
        </location>
    </feature>
    <feature type="domain" description="GST C-terminal" evidence="10">
        <location>
            <begin position="89"/>
            <end position="217"/>
        </location>
    </feature>
    <feature type="domain" description="EF-1-gamma C-terminal" evidence="8">
        <location>
            <begin position="271"/>
            <end position="397"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="222"/>
            <end position="260"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="222"/>
            <end position="253"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00003468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00011237"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022768"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022917"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00519"/>
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    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="RuleBase" id="RU003494"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50040"/>
        </source>
    </evidence>
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            <dbReference type="PROSITE" id="PS50404"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50405"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001645.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="397" mass="45242" checksum="933013821EF14C13" modified="2019-05-08" version="1">MAAAGTLYTYPENWRAFKALIAAQYSGAKIKVLSTPPQFHFGQTNKTPEFLKKFPIGKVPAFEGADGFCVFESNAIAHYVSNDELRGSTKEIAAQIIQWVSFADSDIVPPASTWVFPTLGIMHYNKQATEYAKEEVKRVLGILDSHLKTQTFLVGERITLADITVVCTLLWLYKQVLEPSFRQPYSNTNRWFVTCINQPQFKAVLGEVKLCEKMAQFDAKKFAENQPKRDTPKKEKPAKEEKKQEKKEEKRSEPEEELDECDQALAAEPKSKDPFAHLPKSPFVMDEFKRKYSNEDTLTVALPYFWEHFDKDGWSIWYSQYRFPEELSQTFMSCNLITGMFQRLDKLRKNAFSSVILFGTNNDSTISGIWVFRGQELAFPVSLCTVCGCAGESGCTH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJ03</accession>
    <name>A0A452GJ03_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Ig-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GJ03"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001800.1">
        <property type="protein sequence ID" value="ENSGAGP00000001581.1"/>
        <property type="gene ID" value="ENSGAGG00000001272.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="entry name" value="IgC1"/>
        <property type="match status" value="3"/>
    </dbReference>
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        <property type="entry name" value="Uncharacterized LOC100216153"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
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        <property type="entry name" value="Ig-like_dom_sf"/>
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    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
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        <property type="entry name" value="Ig/MHC_CS"/>
    </dbReference>
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        <property type="entry name" value="Ig_C1-set"/>
    </dbReference>
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        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013106">
        <property type="entry name" value="Ig_V-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050380">
        <property type="entry name" value="Immune_Resp_Modulators"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23411">
        <property type="entry name" value="TAPASIN"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="C1-set"/>
        <property type="match status" value="3"/>
    </dbReference>
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    </dbReference>
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    </dbReference>
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    </dbReference>
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    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
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        <location>
            <begin position="1"/>
            <end position="21"/>
        </location>
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    <feature type="chain" id="PRO_5019568313" description="Ig-like domain-containing protein" evidence="2">
        <location>
            <begin position="22"/>
            <end position="453"/>
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        <location>
            <begin position="7"/>
            <end position="127"/>
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            <begin position="132"/>
            <end position="227"/>
        </location>
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        <location>
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            <end position="324"/>
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        <location>
            <begin position="331"/>
            <end position="436"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001581.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="453" mass="49032" checksum="55748734327D7300" modified="2019-05-08" version="1">MPLAVAPGLLLLLLLASGTRGALVVSVPTSPVRAQPGSDLLLGCHFSVGGAVDMDALVVQWKLGGRPVAEFSEVQSYFRAGASLSLEGLRVGNASLLLPRVGGTDAGLYTCSIIHSPSRESRQVELRVEAPPRVSVSGTVVRAGEHSTVTCSVNSFYPESVNITWLRDKQIVGGPETPLAQPGPDGLYSATSLLQLVPSISHTDANYSCRVQHITLGAPIQEPFQLSVLSQPRLTLLQPLVTLEGQVTLQCLVSGYYPPDITVQWLRDGDPLVTMESSPMKETDGSFALRSSYILNHPESNAQANFACRVQHPALSTPLESTATWTVPQAPVLGEISRPQVLAPGERVTLSCQISRFYPKELSVTWHRRGRGEPAFRSLDNLHTHKIDTPDSTDRKSYSVTSQLCFTPMVPEDEGAEYRCYVEHVTLKEAEEKSTGPLELRGCPLLDGDMAIF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJ04</accession>
    <name>A0A452GJ04_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Pre-mRNA-processing factor 40 homolog A</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="15">Formin-binding protein 11</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="16">Formin-binding protein 3</fullName>
        </alternativeName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="12">Binds to WASL/N-WASP and suppresses its translocation from the nucleus to the cytoplasm, thereby inhibiting its cytoplasmic function. Plays a role in the regulation of cell morphology and cytoskeletal organization. Required in the control of cell shape and migration. May play a role in cytokinesis. May be involved in pre-mRNA splicing.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus matrix</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Nucleus speckle</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="13">Belongs to the PRPF40 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJ04"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001811.1">
        <property type="protein sequence ID" value="ENSGAGP00000001592.1"/>
        <property type="gene ID" value="ENSGAGG00000001270.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016363">
        <property type="term" value="C:nuclear matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016607">
        <property type="term" value="C:nuclear speck"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005685">
        <property type="term" value="C:U1 snRNP"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071004">
        <property type="term" value="C:U2-type prespliceosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045292">
        <property type="term" value="P:mRNA cis splicing, via spliceosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00201">
        <property type="entry name" value="WW"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.440:FF:000003">
        <property type="entry name" value="Pre-mRNA processing factor 40 homolog A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.440:FF:000002">
        <property type="entry name" value="pre-mRNA-processing factor 40 homolog A isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.440:FF:000009">
        <property type="entry name" value="pre-mRNA-processing factor 40 homolog A isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.440:FF:000011">
        <property type="entry name" value="pre-mRNA-processing factor 40 homolog A isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.70.10:FF:000029">
        <property type="entry name" value="pre-mRNA-processing factor 40 homolog A isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.70.10:FF:000031">
        <property type="entry name" value="pre-mRNA-processing factor 40 homolog A isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.440:FF:000012">
        <property type="entry name" value="pre-mRNA-processing factor 40 homolog A isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.440">
        <property type="entry name" value="FF domain"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002713">
        <property type="entry name" value="FF_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036517">
        <property type="entry name" value="FF_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039726">
        <property type="entry name" value="Prp40-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001202">
        <property type="entry name" value="WW_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036020">
        <property type="entry name" value="WW_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11864:SF20">
        <property type="entry name" value="PRE-MRNA-PROCESSING FACTOR 40 HOMOLOG A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11864">
        <property type="entry name" value="PRE-MRNA-PROCESSING PROTEIN PRP40"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01846">
        <property type="entry name" value="FF"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25432">
        <property type="entry name" value="FF_PRPF40A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00397">
        <property type="entry name" value="WW"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00441">
        <property type="entry name" value="FF"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00456">
        <property type="entry name" value="WW"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81698">
        <property type="entry name" value="FF domain"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF64518">
        <property type="entry name" value="Phase 1 flagellin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51045">
        <property type="entry name" value="WW domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51676">
        <property type="entry name" value="FF"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01159">
        <property type="entry name" value="WW_DOMAIN_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50020">
        <property type="entry name" value="WW_DOMAIN_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0007">Acetylation</keyword>
    <keyword evidence="17" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="3" id="KW-0488">Methylation</keyword>
    <keyword evidence="6" id="KW-0507">mRNA processing</keyword>
    <keyword evidence="10" id="KW-0508">mRNA splicing</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="8" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="WW" evidence="19">
        <location>
            <begin position="98"/>
            <end position="131"/>
        </location>
    </feature>
    <feature type="domain" description="WW" evidence="19">
        <location>
            <begin position="144"/>
            <end position="172"/>
        </location>
    </feature>
    <feature type="domain" description="FF" evidence="20">
        <location>
            <begin position="332"/>
            <end position="386"/>
        </location>
    </feature>
    <feature type="domain" description="FF" evidence="20">
        <location>
            <begin position="399"/>
            <end position="453"/>
        </location>
    </feature>
    <feature type="domain" description="FF" evidence="20">
        <location>
            <begin position="466"/>
            <end position="526"/>
        </location>
    </feature>
    <feature type="domain" description="FF" evidence="20">
        <location>
            <begin position="546"/>
            <end position="606"/>
        </location>
    </feature>
    <feature type="domain" description="FF" evidence="20">
        <location>
            <begin position="611"/>
            <end position="666"/>
        </location>
    </feature>
    <feature type="domain" description="FF" evidence="20">
        <location>
            <begin position="681"/>
            <end position="738"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="86"/>
            <end position="116"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="182"/>
            <end position="202"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="297"/>
            <end position="322"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="742"/>
            <end position="897"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="17">
        <location>
            <begin position="385"/>
            <end position="412"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="17">
        <location>
            <begin position="517"/>
            <end position="549"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="86"/>
            <end position="95"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="96"/>
            <end position="111"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="182"/>
            <end position="191"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="18">
        <location>
            <begin position="192"/>
            <end position="202"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="18">
        <location>
            <begin position="746"/>
            <end position="767"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="18">
        <location>
            <begin position="806"/>
            <end position="821"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="822"/>
            <end position="855"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004109"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004324"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022664"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023187"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00057440"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00061317"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00072041"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00078214"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00080815"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50020"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS51676"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001592.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="897" mass="103251" checksum="C19D46B3FCFEBE16" modified="2019-05-08" version="1">MQDEGSGNLMLPEDTGGWGHLASLPKSIKMPGMMQSMMPGMMMSHMSQAHMQPTVPPGVNSMDTQVGTTPPGTQITHQIVCAAQQTTTTNSSINEDPSKQKSMWTEHKSPDGRTYYYNTETKQSTWEKPDDLKTPAEQLLSKCPWKEYKSDSGKPYYYNSQTKESRWAKPKELEDLEAMIKAEENSSKPEESTPTSSAPAVATEITDTVTATATTATTTATTTTSDVETAATTVPAVAEAPEAEASTVAAVVENENTVTTTADEQVQQTTTTTTTTTAATTIVTPVVQEQNAEAVTNTVEETSKQEASVDVAPKKEEEDTQPVKKTYTWNTKEEAKQAFKELLKEKRVPSNASWEQAMKMIINDPRYSALAKLSEKKQAFNAYKVQTEKEEKEEARSKYKEAKESFQRFLENHEKMTSTTRYKKAEQMFGEMEVWNAISERDRLEIYEDVLFFLSKKEKEQAKQLRKRNWEALKNILDNMANVTYSTTWSEAQQYLMDNPTFAEDEELQNMDKEDALICFEEHIRALEKEEEEEKQKSLLRERRRQRKNRESFQLFLDELHEHGQLHSMSSWMELYPTISSDIRFTNMLGQPGSTALDLFKFYVEDLKARYHDEKKIIKDILKDKGFLVEVNTTFEDFVTVISSTKRATTLDAGNIKLAFNSLLEKAEAREREREKEEARKMKRKESAFKSMLKQATPPIELDAVWEDIRERFVKEPAFEDITLESERKRIFKDFIHVLEHECQHHHSKNKKHSKKSKKHHRKRSRSRSGSESDDDDSHSKKKRQRSESRSVSDRSSSAESERSYKKSKKHKKKNKKRRHKSDSPESDTEREKDKKEKERENEKDRPRQRSESKHKSPPKKRPGKDSGNWDTSGSELSEGELEKQRRTLLEQLDEDQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJ09</accession>
    <name>A0A452GJ09_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">DUF4704 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000001817.1">
        <property type="protein sequence ID" value="ENSGAGP00000001597.1"/>
        <property type="gene ID" value="ENSGAGG00000000400.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019901">
        <property type="term" value="F:protein kinase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008104">
        <property type="term" value="P:intracellular protein localization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050865">
        <property type="entry name" value="BEACH_Domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031570">
        <property type="entry name" value="NBEA/BDCP_DUF4704"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13743">
        <property type="entry name" value="BEIGE/BEACH-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13743:SF62">
        <property type="entry name" value="NEUROBEACHIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15787">
        <property type="entry name" value="DUF4704"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0853">WD repeat</keyword>
    <feature type="domain" description="DUF4704" evidence="3">
        <location>
            <begin position="6"/>
            <end position="392"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="416"/>
            <end position="439"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="684"/>
            <end position="720"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1284"/>
            <end position="1303"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="425"/>
            <end position="438"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="684"/>
            <end position="708"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="709"/>
            <end position="720"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="1288"/>
            <end position="1298"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022574"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="Pfam" id="PF15787"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001597.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1411" mass="156231" checksum="FB9C51D3E699A04E" modified="2019-05-08" version="1">MERNFSSRVHITRAVLEQFLSFAKYLDGLSHGAPLLKQLCDHILFNPAIWIHTPAKVQLSLYTYLSAEFIGTATIYTTIRRVGTVLQLMHTLKYYYWVVNPADSSGITPKGLEGPRPSQKEIISLRAFMLLFLKQLILKDRGVKEDELQSILNYLLTMHEDENIHDVLQLLVALMSEHPASMIPAFDQRNGIRVIYKLLASKSESIWVQALKVLGYFLKHLGHKRKVEIMHTHSLFTLLGERLMLHTNTVTITTYNTLYEILTEQVCTQVVHKPHPEPDSTVKIQNPMILKVVATLLKNSTPSAELMEVRRLFLSDMIKLFSNSRENRRCLLQCSVWQDWMFSLGYINPKNSEEQKITEMVYNIFRILLYHAIKYEWGGWRVWVDTLSIAHSKVTYEAHKEYLAKMYEEYQRQEEENIKKGKKGNVSTISGLSSQTTGAKGGLEIREIEDLSQSQSPESETDYPVSTDTRDLLMATKVSDEVLGSSDRPGGGVHVEVHDLLVDIKAEKVEATEVKLDDMDLSPETLVTGENGALVEVESLLDNVYCAAVEKLQNNVHGSVGIIKKSEEKDNGPLITLADEKDEPSTNNTSFLFDKIPSQEEKLLPELSSNHITIPNVQETQMHLGVSDDLGLLAHMTGSVDITCTSSIIEDKEFKIHTTSDGMSNLSERELGSSSKGLEYAEMTATTLEPESSGSKSLPNVDTGSIISDTERSDDGKEAGKEIRKIQTTTTTQAVQGRSVTQQDRDLRVDLGFRGMPMTEEQRRQFSPGPRTTMFRIPEFKWSPMHQRLLTDLLFALETDVHVWRSHSTKSVMDFVNSNENIIFVHNTIHLISQMVDNIIIACGGILPLLSAATSPTTELENIEVTQGMSAETAVTFLSRLMAMVDVLVFASSLNFSEIEAEKNMSSGGLMRQCLRLVCCVAVRNCLECRQRQRERVNKSSLISSKTQETLQGVTATALTKSPLENVPGNLSPIKDPDRLLQDVDINRLRAVVFRDVDDSKQAQFLALAVVYFISVLMVSKYRDILEPQRETARCGSQAGRNIRQEINSPTSTVVVIPSIPHPSLNHGFLAKLIPEQSFAHSFYKETPAVFPENIKEKETPTPVEDIQLESSIPHTDSGIGEEQMPSILNGTDLETSAGPDAMSELFSTLSSEVKKSQESLTESPSEILKPAPSISSISQSKGMNVKEILKSLVAAPIEIAECGPDPIPYPDPALKREAQSILPMQFHSFDRSVVVPVKKPPPGSLAVTTVGAATAGGVLPPGTTPNIFAATGATPKSMINTTGAVDSGSSSSSSSSSFVNGATSKNLPAVQTVAPMPEDSAENMSITAKLERALEKVAPLLREIFVDFAPFLSRTLLGSHGQELLIEGLVCMKSSTSVVELVMLLCSQEWQNSIQKNAGLAFIELINEGR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJ21</accession>
    <name>A0A452GJ21_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">RING-type E3 ubiquitin transferase</fullName>
            <ecNumber evidence="5">2.3.2.27</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>S-ubiquitinyl-[E2 ubiquitin-conjugating enzyme]-L-cysteine + [acceptor protein]-L-lysine = [E2 ubiquitin-conjugating enzyme]-L-cysteine + N(6)-ubiquitinyl-[acceptor protein]-L-lysine.</text>
            <dbReference type="EC" id="2.3.2.27"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="3">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the TRIM/RBCC family.</text>
    </comment>
    <dbReference type="EC" id="2.3.2.27" evidence="5"/>
    <dbReference type="AlphaFoldDB" id="A0A452GJ21"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001883.1">
        <property type="protein sequence ID" value="ENSGAGP00000001658.1"/>
        <property type="gene ID" value="ENSGAGG00000001264.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061630">
        <property type="term" value="F:ubiquitin protein ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd16598">
        <property type="entry name" value="RING-HC_TRIM26_C-IV"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12888">
        <property type="entry name" value="SPRY_PRY_TRIM7_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.920:FF:000004">
        <property type="entry name" value="Butyrophilin subfamily 1 member A1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.920">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001870">
        <property type="entry name" value="B30.2/SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043136">
        <property type="entry name" value="B30.2/SPRY_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003879">
        <property type="entry name" value="Butyrophylin_SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006574">
        <property type="entry name" value="PRY"/>
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        <property type="entry name" value="TRIM/RBCC"/>
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        <property type="entry name" value="Znf_B-box"/>
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        <property type="entry name" value="Znf_B-box_chordata"/>
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        <property type="entry name" value="Znf_PARP"/>
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        <property type="entry name" value="Znf_RING"/>
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        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
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        <property type="entry name" value="Znf_RING_CS"/>
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        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE TRIM"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="zf-B_box"/>
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        <property type="entry name" value="zf-C3HC4_4"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="BBOXZNFINGER"/>
    </dbReference>
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        <property type="entry name" value="BUTYPHLNCDUF"/>
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        <property type="match status" value="1"/>
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    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
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    </dbReference>
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        <property type="entry name" value="RING/U-box"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="B302_SPRY"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="ZF_BBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="ZF_PARP_2"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="ZF_RING_1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="ZF_RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12 14" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0808">Transferase</keyword>
    <keyword evidence="10" id="KW-0833">Ubl conjugation pathway</keyword>
    <keyword evidence="11" id="KW-0862">Zinc</keyword>
    <keyword evidence="9 13" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="RING-type" evidence="16">
        <location>
            <begin position="16"/>
            <end position="59"/>
        </location>
    </feature>
    <feature type="domain" description="PARP-type" evidence="15">
        <location>
            <begin position="43"/>
            <end position="68"/>
        </location>
    </feature>
    <feature type="domain" description="B box-type" evidence="17">
        <location>
            <begin position="81"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="domain" description="B30.2/SPRY" evidence="18">
        <location>
            <begin position="254"/>
            <end position="439"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="14">
        <location>
            <begin position="119"/>
            <end position="194"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00008518"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00024"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50064"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50119"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50188"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001658.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="439" mass="50493" checksum="01D70DB6EAE2E596" modified="2019-05-08" version="1">MASSLSGLDMREEATCPICLEYLTDPVSVDCGHNFCRGCVTQYCVEWAGRGSLTCPICRSQILKGNLRPNWQLGNLVEKMKHVAVCTRHKKTLDLFCEEDEEAVCVVCERASQHRSHTVLLIEEAAQKYKAHLKTLKEEREKQLGLKATGEGKRQEQKIVSEFQQLRQFLEEQERLLLAQLEKLDKEILRIQKENVTKLSGQISHLSELIHELEGKCQKPASEFLQVRLSLKHSRSMCRDRRAPESCPLESSSQRSQCNFSQHCFEISNKLLHMTLDPDTAHPQLILSEDRKTVRYGRLWNKLPNTTKRFDYERCVLGSEGFTMGRHCWEVEVGERGGWAVGVARESLGRKGEISFKPKGGIWAVEYWEGQYWALTDPLTPLPLSWVPRRIWIYLDYEQGQVTFFNADNESLIFTFPPASFAGERIRPLFVPKLPPPRE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJ22</accession>
    <name>A0A452GJ22_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">Diacylglycerol kinase</fullName>
            <shortName evidence="21">DAG kinase</shortName>
            <ecNumber evidence="21">2.7.1.107</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="27 28">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="28" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="27" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="27" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>1,2-di-(9Z-octadecenoyl)-sn-glycerol + ATP = 1,2-di-(9Z-octadecenoyl)-sn-glycero-3-phosphate + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:40327"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:52333"/>
            <dbReference type="ChEBI" id="CHEBI:74546"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="18">
            <dbReference type="Rhea" id="RHEA:40328"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>a 1,2-diacyl-sn-glycerol + ATP = a 1,2-diacyl-sn-glycero-3-phosphate + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10272"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:17815"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:58608"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.1.107"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="19">
            <dbReference type="Rhea" id="RHEA:10273"/>
        </physiologicalReaction>
    </comment>
    <comment type="pathway">
        <text evidence="20">Glycerolipid metabolism.</text>
    </comment>
    <comment type="pathway">
        <text evidence="3">Lipid metabolism; glycerolipid metabolism.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4 21">Belongs to the eukaryotic diacylglycerol kinase family.</text>
    </comment>
    <dbReference type="EC" id="2.7.1.107" evidence="21"/>
    <dbReference type="AlphaFoldDB" id="A0A452GJ22"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001895.1">
        <property type="protein sequence ID" value="ENSGAGP00000001669.1"/>
        <property type="gene ID" value="ENSGAGG00000001299.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00230"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004143">
        <property type="term" value="F:ATP-dependent diacylglycerol kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046982">
        <property type="term" value="F:protein heterodimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042803">
        <property type="term" value="F:protein homodimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046486">
        <property type="term" value="P:glycerolipid metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007200">
        <property type="term" value="P:phospholipase C-activating G protein-coupled receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd20847">
        <property type="entry name" value="C1_DGKdelta_rpt1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd20893">
        <property type="entry name" value="C1_DGKdelta_rpt2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd13274">
        <property type="entry name" value="PH_DGK_type2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd09575">
        <property type="entry name" value="SAM_DGK-delta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.50:FF:000021">
        <property type="entry name" value="Diacylglycerol kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Diacylglycerol kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.200.40:FF:000001">
        <property type="entry name" value="Diacylglycerol kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.60.20:FF:000002">
        <property type="entry name" value="Diacylglycerol kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.60.20:FF:000029">
        <property type="entry name" value="Diacylglycerol kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10330:FF:000001">
        <property type="entry name" value="Diacylglycerol kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.200.40">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.60.20">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.10330">
        <property type="entry name" value="Probable inorganic polyphosphate/atp-NAD kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.50">
        <property type="entry name" value="Transcription Factor, Ets-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017438">
        <property type="entry name" value="ATP-NAD_kinase_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046349">
        <property type="entry name" value="C1-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047478">
        <property type="entry name" value="C1_DGKdelta_rpt1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047477">
        <property type="entry name" value="C1_DGKdelta_rpt2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037607">
        <property type="entry name" value="DGK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037606">
        <property type="entry name" value="DGK-delta_SAM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR054474">
        <property type="entry name" value="DGKD_4H"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000756">
        <property type="entry name" value="Diacylglycerol_kin_accessory"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001206">
        <property type="entry name" value="Diacylglycerol_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016064">
        <property type="entry name" value="NAD/diacylglycerol_kinase_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002219">
        <property type="entry name" value="PKC_DAG/PE"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001660">
        <property type="entry name" value="SAM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013761">
        <property type="entry name" value="SAM/pointed_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11255">
        <property type="entry name" value="DIACYLGLYCEROL KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11255:SF30">
        <property type="entry name" value="DIACYLGLYCEROL KINASE DELTA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00130">
        <property type="entry name" value="C1_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00609">
        <property type="entry name" value="DAGK_acc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00781">
        <property type="entry name" value="DAGK_cat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22944">
        <property type="entry name" value="DGKD_4H"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00169">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07647">
        <property type="entry name" value="SAM_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00109">
        <property type="entry name" value="C1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00045">
        <property type="entry name" value="DAGKa"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00046">
        <property type="entry name" value="DAGKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00233">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00454">
        <property type="entry name" value="SAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57889">
        <property type="entry name" value="Cysteine-rich domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF111331">
        <property type="entry name" value="NAD kinase/diacylglycerol kinase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47769">
        <property type="entry name" value="SAM/Pointed domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50146">
        <property type="entry name" value="DAGK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50105">
        <property type="entry name" value="SAM_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00479">
        <property type="entry name" value="ZF_DAG_PE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50081">
        <property type="entry name" value="ZF_DAG_PE_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="15 21" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="5" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="13 21" id="KW-0418">Kinase</keyword>
    <keyword evidence="16" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="17" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11 21" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="28" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0677">Repeat</keyword>
    <keyword evidence="8 21" id="KW-0808">Transferase</keyword>
    <keyword evidence="14" id="KW-0862">Zinc</keyword>
    <keyword evidence="12" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="PH" evidence="23">
        <location>
            <begin position="1"/>
            <end position="88"/>
        </location>
    </feature>
    <feature type="domain" description="Phorbol-ester/DAG-type" evidence="24">
        <location>
            <begin position="105"/>
            <end position="155"/>
        </location>
    </feature>
    <feature type="domain" description="Phorbol-ester/DAG-type" evidence="24">
        <location>
            <begin position="177"/>
            <end position="228"/>
        </location>
    </feature>
    <feature type="domain" description="DAGKc" evidence="26">
        <location>
            <begin position="259"/>
            <end position="394"/>
        </location>
    </feature>
    <feature type="domain" description="SAM" evidence="25">
        <location>
            <begin position="1048"/>
            <end position="1111"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="497"/>
            <end position="543"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="600"/>
            <end position="623"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="22">
        <location>
            <begin position="497"/>
            <end position="506"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="22">
        <location>
            <begin position="523"/>
            <end position="537"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00009280"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023371"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023411"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00060536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="RuleBase" id="RU361128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS50081"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS50105"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="26">
        <source>
            <dbReference type="PROSITE" id="PS50146"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001669.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="28">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1117" mass="125384" checksum="A0F4993A6A6C9FB8" modified="2019-05-08" version="1">MLMKQTSSFQRWKRRYFKLRGRTLYYAKNAKSIIFDEVDLTDASVAESSTKNINNSFTVITPCRKLILCADNRKEMEDWIAALKTIQNREHFESTQYSMDHFSGMHNWYACSHARPTYCNVCREALSGVTSHGLSCEVCKFKAHKRCAVRATNNCKWTTLASIGKDIIEDEDGISMPHQWLEGNLPVSAKCMVCDKTCGSVLRLQDWRCLWCKAMVHTTCKDLLPTKCPLGLCKVSVIPPTALNSIDSDGFWKATCPPSCTSPLLVFVNSKSGDNQGVKFLRRFKQLLNPAQVFDLMNGGPHLGLRLFQKFDTFRILVCGGDGSVGWVLSEIDSLNLHKQCQLGVLPLGTGNDLARVLGWGSACDDDTQLPQILEKLERASTKMLDRWSIMVYETKLPRQSSTSTVTEDFSDDSEVQKILFYEDSVAAHLSKILTSDQHSVVISSAKVLCETVKDFVARVGKAYEKATESSEESEVMARKCSVLKEKLDSLLKTLNDESQASSSLSNPPPTIAEETEDGDGSGSACDSSSDHSMGSSCTARPQIFRPREQLMLRANSLKKAIRQIIEHAEKAVDEQNAQTQEQEGFILGLPTSEEGKELRNEDKTSLQPLHSGNYGVPKGRSHRKDIRAGMSGSLPGSSVISRLLINADPFNSEPENLECYTEKCVMNNYFGIGLDAKISLDFNNKRDEHPEKCRSRTKNMMWYGVLGTKELLHRTYKNLEQKVLLECDGRPIPLPSLQGIAVLNIPSYAGGTNFWGGTKEDDTFTAPSFDDKILEVVAVFGSMQMAVSRVINLQHHRIAQCRTVKIAILGDEGVPVQVDGEAWIQPPGYIWIVHKNRAQTLTRDRAFESTLKSWEDKQKCELPRPSSFSLHPEIMSEEESTQINQFGQAAGALIHSIREIAQSYQDMEQELAHAVNASSKSMDKVYAKSKSTEGLNCSLVVEMVNNVKALHSETELLLAGKMALQLDPPQKEQLLAALMDMDLQLRKLADIPWLWQLMEPSDDENQMLDYSKRSRSGKFRLVTKFKKEKNNKNKETRSSMGLPVHLWGTEEVAAWLEHLSLCEYKDIFIRHDVRGSELLHLERRDLKDLGVTKVGHMKRILHGIKELSRNTLVSEV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GJ28</accession>
    <name>A0A452GJ28_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Zinc finger protein 423</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="19">Olf1/EBF-associated zinc finger protein</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="20">Smad- and Olf-interacting zinc finger protein</fullName>
        </alternativeName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="17">Transcription factor that can both act as an activator or a repressor depending on the context. Plays a central role in BMP signaling and olfactory neurogenesis. Associates with SMADs in response to BMP2 leading to activate transcription of BMP target genes. Acts as a transcriptional repressor via its interaction with EBF1, a transcription factor involved in terminal olfactory receptor neurons differentiation; this interaction preventing EBF1 to bind DNA and activate olfactory-specific genes. Involved in olfactory neurogenesis by participating in a developmental switch that regulates the transition from differentiation to maturation in olfactory receptor neurons. Controls proliferation and differentiation of neural precursors in cerebellar vermis formation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GJ28"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001837.1">
        <property type="protein sequence ID" value="ENSGAGP00000001617.1"/>
        <property type="gene ID" value="ENSGAGG00000001293.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043565">
        <property type="term" value="F:sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045892">
        <property type="term" value="P:negative regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000468">
        <property type="entry name" value="B-cell lymphoma 6 protein-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000244">
        <property type="entry name" value="zinc finger protein 423"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000469">
        <property type="entry name" value="Zinc finger protein 423"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000483">
        <property type="entry name" value="Zinc finger protein 423"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000548">
        <property type="entry name" value="Zinc finger protein 423"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000760">
        <property type="entry name" value="Zinc finger protein 423"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001417">
        <property type="entry name" value="zinc finger protein 423"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002384">
        <property type="entry name" value="zinc finger protein 423"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:004063">
        <property type="entry name" value="Zinc finger protein 423"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000143">
        <property type="entry name" value="Zinc finger protein 521"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000167">
        <property type="entry name" value="Zinc finger protein 521"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000261">
        <property type="entry name" value="Zinc finger protein 521"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24379:SF128">
        <property type="entry name" value="C2H2-TYPE DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24379">
        <property type="entry name" value="KRAB AND ZINC FINGER DOMAIN-CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13912">
        <property type="entry name" value="zf-C2H2_6"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="29"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="25"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="22"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="14" id="KW-0010">Activator</keyword>
    <keyword evidence="3" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="9" id="KW-0221">Differentiation</keyword>
    <keyword evidence="13" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="6" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="16" id="KW-0539">Nucleus</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0678">Repressor</keyword>
    <keyword evidence="15" id="KW-0804">Transcription</keyword>
    <keyword evidence="12" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="10" id="KW-0862">Zinc</keyword>
    <keyword evidence="8 21" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="62"/>
            <end position="89"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="90"/>
            <end position="117"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="118"/>
            <end position="145"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="146"/>
            <end position="173"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="187"/>
            <end position="215"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="219"/>
            <end position="247"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="408"/>
            <end position="436"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="445"/>
            <end position="468"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="560"/>
            <end position="582"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="590"/>
            <end position="612"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="620"/>
            <end position="644"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="648"/>
            <end position="676"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="678"/>
            <end position="706"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="709"/>
            <end position="736"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="735"/>
            <end position="758"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="813"/>
            <end position="840"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="857"/>
            <end position="884"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="886"/>
            <end position="913"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="1047"/>
            <end position="1075"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="1155"/>
            <end position="1182"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="1186"/>
            <end position="1214"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="23">
        <location>
            <begin position="1216"/>
            <end position="1241"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="267"/>
            <end position="327"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="518"/>
            <end position="546"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="1069"/>
            <end position="1090"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="22">
        <location>
            <begin position="287"/>
            <end position="301"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="22">
        <location>
            <begin position="532"/>
            <end position="543"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="22">
        <location>
            <begin position="1075"/>
            <end position="1088"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022491"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00055444"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00067811"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00076623"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00080204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001617.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1241" mass="140495" checksum="633BC4061540B29F" modified="2019-05-08" version="1">MEIASLEAADFSKLLYDGDDDPQLSWVASSPSSKDVASPTQMIGDGCDLGIGEEEGGTGLPYPCQFCDKSFIRLSYLKRHEQIHSDKLPFKCTYCSRLFKHKRSRDRHIKLHTGDKKYHCHECEAAFSRSDHLKIHLKTHSSSKPFKCTVCKRGFSSTSSLQSHMQAHKKNKEHMAKTEKEVKKDDFMCDYCEETFSQTEDLEKHVMTRHPQLSEKADLQCIHCPEVFADENSLLSHIHQAHANKKHKCPMCPEQFSSVEEVYCHLDSHRQPDSSNHSVSPDPVLGSVASMSSATPDSSASVERGSTPDSTLKPLRGQKKIRSAEREEGQNWSKVTYSCPYCSKRDFNSLAVLEIHLKTIHADKPQQSHTCQICLDSMPTLYNLNEHVRKVHKNHAYPMMQFNNISAFHCNYCPEMFADINSLQEHIRITHCGPSATPQDGNNAFFCNQCSMGFLTEASLTEHIQQTHCNVGNSKLDSPVVQPTQSFMEVYSCPYCTNSPIFGSILKLTKHIKENHKNIPLAHNKKSKAEQSPVSSDVEVSSPKRQRLSASLNSVSNGEYPCNQCDLKFSNFESFQTHLKLHLELLLRKQSCPQCKEDFDSQESLLQHLTVHYMTTSTHYVCESCDKQFSSVDDLQKHLLDMHTFVLYHCTLCQEVFDSKVSIQVHLAVKHSNEKKMYRCTACNWDFRKEVDLQIHVKHSHLGNPTKSHKCIFCGETFSTEVELQCHITTHSKKYNCKFCSKAFHAIILLEKHLREKHCVFDAGAENGTANGMTPTNKKTEAADLQNMLMKNPDTSNSHEASEDDVDASEPMYGCDICGAAYTMEVLLQNHRLRDHNIRPGEDDCSRKKAEFIKGSHKCNICSRTFFSENGLREHMQTHRGPAKHYMCPICGERFPSLLTLTEHKVTHSKSLDTGTCRICKMPLQSEEEFIEHCQMHPDLRNSLTGFRCVVCMQTVTSTLELKIHGTFHMQKLAGNSAASSPNGQALQKLYKCALCLKEFRNKQDLVKLDVNGLPYGLCAGCMTRGTNGQSSGLTPQEVNERPCGSLRCPECSVKFESAEDLENHIQLDHRDLTPETSGQRKGTQTSPVPRKKTYQCIKCQMTFENEREIQIHVANHMIGAAQTPYGETLPSFPAPALVERRTMKNVTAEEGINHECKLCNQMFDSPAKLLCHLIEHSFEGMGGTFKCPVCFTVFVQANKLQQHIFAVHGQEDKIYDCSQCPQKFFFQTELQNHTLSQHAQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZP1</accession>
    <name>A0A452GZP1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="27">CMP-N-acetylneuraminate-beta-galactosamide-alpha-2,3-sialyltransferase 4</fullName>
            <ecNumber evidence="16">2.4.3.2</ecNumber>
            <ecNumber evidence="17">2.4.3.4</ecNumber>
            <ecNumber evidence="23">2.4.3.6</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="28">Alpha 2,3-sialyltransferase IV</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="18">Gal-beta-1,3-GalNAc-alpha-2,3-sialyltransferase</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="30">Gal-beta-1,4-GlcNAc-alpha-2,3-sialyltransferase</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="29">N-acetyllactosaminide alpha-2,3-sialyltransferase</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="31">ST3Gal IV</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="32">Sialyltransferase 4C</fullName>
        </alternativeName>
    </protein>
    <organism evidence="35 36">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="36" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="35" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="35" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="14">
            <text>a beta-D-galactosyl-(1-&gt;3)-N-acetyl-alpha-D-galactosaminyl derivative + CMP-N-acetyl-beta-neuraminate = an N-acetyl-alpha-neuraminyl-(2-&gt;3)-beta-D-galactosyl-(1-&gt;3)-N-acetyl-alpha-D-galactosaminyl derivative + CMP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:21616"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57812"/>
            <dbReference type="ChEBI" id="CHEBI:60377"/>
            <dbReference type="ChEBI" id="CHEBI:133470"/>
            <dbReference type="ChEBI" id="CHEBI:139596"/>
            <dbReference type="EC" id="2.4.3.4"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="14">
            <dbReference type="Rhea" id="RHEA:21617"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="25">
            <text>a beta-D-galactosyl-(1-&gt;3)-N-acetyl-beta-D-galactosaminyl derivative + CMP-N-acetyl-beta-neuraminate = an N-acetyl-alpha-neuraminyl-(2-&gt;3)-beta-D-galactosyl-(1-&gt;3)-N-acetyl-beta-D-galactosaminyl derivative + CMP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:52380"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57812"/>
            <dbReference type="ChEBI" id="CHEBI:60377"/>
            <dbReference type="ChEBI" id="CHEBI:136588"/>
            <dbReference type="ChEBI" id="CHEBI:136589"/>
            <dbReference type="EC" id="2.4.3.2"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="25">
            <dbReference type="Rhea" id="RHEA:52381"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="22">
            <text>a beta-D-galactosyl-(1-&gt;4)-N-acetyl-beta-D-glucosaminyl derivative + CMP-N-acetyl-beta-neuraminate = an N-acetyl-alpha-neuraminyl-(2-&gt;3)-beta-D-galactosyl-(1-&gt;4)-N-acetyl-beta-D-glucosaminyl derivative + CMP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:52316"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57812"/>
            <dbReference type="ChEBI" id="CHEBI:60377"/>
            <dbReference type="ChEBI" id="CHEBI:133507"/>
            <dbReference type="ChEBI" id="CHEBI:136545"/>
            <dbReference type="EC" id="2.4.3.6"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="22">
            <dbReference type="Rhea" id="RHEA:52317"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>a ganglioside GA1 (d18:1(4E)) + CMP-N-acetyl-beta-neuraminate = a ganglioside GM1b (d18:1(4E)) + CMP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:47560"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:27938"/>
            <dbReference type="ChEBI" id="CHEBI:57812"/>
            <dbReference type="ChEBI" id="CHEBI:60377"/>
            <dbReference type="ChEBI" id="CHEBI:78568"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="19">
            <dbReference type="Rhea" id="RHEA:47561"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="20">
            <text>a ganglioside GM1 (d18:1(4E)) + CMP-N-acetyl-beta-neuraminate = a ganglioside GD1a (d18:1(4E)) + CMP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:18021"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57812"/>
            <dbReference type="ChEBI" id="CHEBI:60377"/>
            <dbReference type="ChEBI" id="CHEBI:77709"/>
            <dbReference type="ChEBI" id="CHEBI:78445"/>
            <dbReference type="EC" id="2.4.3.2"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="20">
            <dbReference type="Rhea" id="RHEA:18022"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="26">
            <text>a ganglioside GT1c (d18:1(4E)) + CMP-N-acetyl-beta-neuraminate = a ganglioside GQ1c (d18:1(4E)) + CMP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:47588"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57812"/>
            <dbReference type="ChEBI" id="CHEBI:60377"/>
            <dbReference type="ChEBI" id="CHEBI:87789"/>
            <dbReference type="ChEBI" id="CHEBI:87791"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="26">
            <dbReference type="Rhea" id="RHEA:47589"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="24">
            <text>a neolactoside nLc4Cer + CMP-N-acetyl-beta-neuraminate = a neolactoside IV(3)-alpha-NeuAc-nLc4Cer + CMP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:65432"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57812"/>
            <dbReference type="ChEBI" id="CHEBI:60377"/>
            <dbReference type="ChEBI" id="CHEBI:90376"/>
            <dbReference type="ChEBI" id="CHEBI:90390"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="24">
            <dbReference type="Rhea" id="RHEA:65433"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="21">
            <text>a neolactoside nLc4Cer(d18:1(4E)) + CMP-N-acetyl-beta-neuraminate = a neolactoside IV(3)-alpha-NeuAc-nLc4Cer(d18:1(4E)) + CMP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:18913"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:17006"/>
            <dbReference type="ChEBI" id="CHEBI:57812"/>
            <dbReference type="ChEBI" id="CHEBI:58665"/>
            <dbReference type="ChEBI" id="CHEBI:60377"/>
            <dbReference type="EC" id="2.4.3.6"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="21">
            <dbReference type="Rhea" id="RHEA:18914"/>
        </physiologicalReaction>
    </comment>
    <comment type="pathway">
        <text evidence="2">Protein modification; protein glycosylation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Golgi apparatus membrane</location>
            <topology evidence="1">Single-pass type II membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="15">Golgi apparatus</location>
            <location evidence="15">Golgi stack membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the glycosyltransferase 29 family.</text>
    </comment>
    <dbReference type="EC" id="2.4.3.2" evidence="16"/>
    <dbReference type="EC" id="2.4.3.4" evidence="17"/>
    <dbReference type="EC" id="2.4.3.6" evidence="23"/>
    <dbReference type="AlphaFoldDB" id="A0A452GZP1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008930.1">
        <property type="protein sequence ID" value="ENSGAGP00000007742.1"/>
        <property type="gene ID" value="ENSGAGG00000006181.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032580">
        <property type="term" value="C:Golgi cisterna membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000139">
        <property type="term" value="C:Golgi membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0047288">
        <property type="term" value="F:beta-D-galactosyl-(1-&gt;3)-N-acetyl-beta-D-galactosaminide alpha-2,3- sialyltransferase"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003836">
        <property type="term" value="F:beta-galactoside (CMP) alpha-2,3-sialyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008118">
        <property type="term" value="F:N-acetyllactosaminide alpha-2,3-sialyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009247">
        <property type="term" value="P:glycolipid biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd23982">
        <property type="entry name" value="GT29_ST3GAL4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1480.20:FF:000005">
        <property type="entry name" value="ST3 beta-galactoside alpha-2,3-sialyltransferase 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1480.20">
        <property type="entry name" value="Glycosyl transferase family 29"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001675">
        <property type="entry name" value="Glyco_trans_29"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051142">
        <property type="entry name" value="Glycosyltransferase_29"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038578">
        <property type="entry name" value="GT29-like_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13713:SF57">
        <property type="entry name" value="CMP-N-ACETYLNEURAMINATE-BETA-GALACTOSAMIDE-ALPHA-2,3-SIALYLTRANSFERASE 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13713">
        <property type="entry name" value="SIALYLTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00777">
        <property type="entry name" value="Glyco_transf_29"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="4" id="KW-0328">Glycosyltransferase</keyword>
    <keyword evidence="9" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="10" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="11 34" id="KW-0472">Membrane</keyword>
    <keyword evidence="36" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0735">Signal-anchor</keyword>
    <keyword evidence="5" id="KW-0808">Transferase</keyword>
    <keyword evidence="6 34" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 34" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="34">
        <location>
            <begin position="38"/>
            <end position="57"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="33">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="33">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004323"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004922"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00006003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022676"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00036292"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00037859"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00039106"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00039107"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00042448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00043673"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00043773"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00048162"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00049294"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00049726"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00051076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00051975"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00052660"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="ARBA" id="ARBA00072813"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="ARBA" id="ARBA00075868"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="ARBA" id="ARBA00076295"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="ARBA" id="ARBA00076532"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="31">
        <source>
            <dbReference type="ARBA" id="ARBA00081234"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="32">
        <source>
            <dbReference type="ARBA" id="ARBA00082801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="33">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="34">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="35">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007742.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="36">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="361" mass="41508" checksum="D8765F129B14960F" modified="2019-05-08" version="1">MADSGNAQDSCEVTSSGRNESNSHRVAPHLLIKMINKFRWKVILGLAFFLVMAWYYISREESFSFPVQDVRMACPQGEMEKKVTLLIGNYTRDHPMFLHLNDYFWVKNRSPYELPYGTKGSEDILLRVLAITSYSVPESIQSMKCRRCAVVGNGHRLRNSSMGEAINKYDVVIRLNNAPVHGYEADVGSKTTMRLFYPESAHFDPRRENNPDTLLVLVPFKPMDFQWLEMILNDKKRIRKGFWKQPPLIWDANPEQVRILNPYFMEITAAKLLNLSMKQPRKIKQKPTTGLLAITLALHFCDLVHIAGFGYPNSANKKQSIHYYEQITLKSMAASEHNVSHEALAIKRMLELGVVKNLTHF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZP5</accession>
    <name>A0A452GZP5_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="14">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="1 12">
            <name>heme</name>
            <dbReference type="ChEBI" id="CHEBI:30413"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Microsome membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 13">Belongs to the cytochrome P450 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZP5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008720.1">
        <property type="protein sequence ID" value="ENSGAGP00000007552.1"/>
        <property type="gene ID" value="ENSGAGG00000005769.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="F:heme binding"/>
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        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005506">
        <property type="term" value="F:iron ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016712">
        <property type="term" value="F:oxidoreductase activity, acting on paired donors, with incorporation or reduction of molecular oxygen, reduced flavin or flavoprotein as one donor, and incorporation of one atom of oxygen"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046222">
        <property type="term" value="P:aflatoxin metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006082">
        <property type="term" value="P:organic acid metabolic process"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006805">
        <property type="term" value="P:xenobiotic metabolic process"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="entry name" value="cytochrome P450 2W1 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.630.10">
        <property type="entry name" value="Cytochrome P450"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Cyt_P450"/>
    </dbReference>
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        <property type="entry name" value="Cyt_P450_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002401">
        <property type="entry name" value="Cyt_P450_E_grp-I"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036396">
        <property type="entry name" value="Cyt_P450_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050182">
        <property type="entry name" value="Cytochrome_P450_fam2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24300:SF302">
        <property type="entry name" value="CYTOCHROME P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24300">
        <property type="entry name" value="CYTOCHROME P450 508A4-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00067">
        <property type="entry name" value="p450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00463">
        <property type="entry name" value="EP450I"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00385">
        <property type="entry name" value="P450"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48264">
        <property type="entry name" value="Cytochrome P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00086">
        <property type="entry name" value="CYTOCHROME_P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="4 12 13" id="KW-0349">Heme</keyword>
    <keyword evidence="9 12 13" id="KW-0408">Iron</keyword>
    <keyword evidence="11" id="KW-0472">Membrane</keyword>
    <keyword evidence="5 12 13" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7" id="KW-0492">Microsome</keyword>
    <keyword evidence="10 13" id="KW-0503">Monooxygenase</keyword>
    <keyword evidence="8 13" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <feature type="binding site" description="axial binding residue" evidence="12">
        <location>
            <position position="424"/>
        </location>
        <ligand>
            <name>heme</name>
            <dbReference type="ChEBI" id="CHEBI:30413"/>
        </ligand>
        <ligandPart>
            <name>Fe</name>
            <dbReference type="ChEBI" id="CHEBI:18248"/>
        </ligandPart>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001971"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004524"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00010617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022848"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023033"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PIRSR" id="PIRSR602401-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="RuleBase" id="RU000461"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007552.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="481" mass="54945" checksum="FB8F1D1164A3D386" modified="2019-05-08" version="1">MALADPTVILLLFVVTSVSILFVKKSWNKRTCLNFPPGPRPLPILGNLHQLNLKRPYRTLLKLSEKYGPVFSIHMGPKNMVVLSGYKMVKEALVNQADAFAERPRIPIFEKFARGNGIIFAHGENWKVMRRFTLTTLRDFGMGRRTIENKILEECDYLIKDFESHKGSSFDTMKMMNAAVSNVIVSIMLGHRFDYEDPKLLRLIHLINESVRLSGSPMVTLFNMFPALGFLPGNHKTVFKNLEELYSFTQQTFIEHLKQLDPNDQRSFIDAFLIRGECVLVSNLFTAGMETTSTTLRWGLLLMMKYPAIQKKVHEEIERVIGSAQLRYEHRTQMPYTDAVIHEIQRFANILPMSLPHETTEDVRLNGYVIPKGTYIIPLLASVLYDKTQWEKPGEFCPQHFLSTDGKFVTNEAFMPFSAGRRICAGETLARMELFLFFTSLLQRFTFHPPPGVTSSDLDLTPAVGFTTPPMPHKICAMPCR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZP7</accession>
    <name>A0A452GZP7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">UNC-45/Cro1/She4 central domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZP7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008862.1">
        <property type="protein sequence ID" value="ENSGAGP00000007678.1"/>
        <property type="gene ID" value="ENSGAGG00000006144.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051879">
        <property type="term" value="F:Hsp90 protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000087">
        <property type="entry name" value="Unc-45 myosin chaperone A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000043">
        <property type="entry name" value="Unc-45 myosin chaperone B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.30">
        <property type="entry name" value="Regulator of chromosome condensation 1/beta-lactamase-inhibitor protein II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009091">
        <property type="entry name" value="RCC1/BLIP-II"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000408">
        <property type="entry name" value="Reg_chr_condens"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024660">
        <property type="entry name" value="UCS_central_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45994">
        <property type="entry name" value="FI21225P1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45994:SF3">
        <property type="entry name" value="PROTEIN UNC-45 HOMOLOG A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00415">
        <property type="entry name" value="RCC1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11701">
        <property type="entry name" value="UNC45-central"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50985">
        <property type="entry name" value="RCC1/BLIP-II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00626">
        <property type="entry name" value="RCC1_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50012">
        <property type="entry name" value="RCC1_3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="UNC-45/Cro1/She4 central" evidence="5">
        <location>
            <begin position="267"/>
            <end position="455"/>
        </location>
    </feature>
    <feature type="repeat" description="RCC1" evidence="3">
        <location>
            <begin position="958"/>
            <end position="1009"/>
        </location>
    </feature>
    <feature type="repeat" description="RCC1" evidence="3">
        <location>
            <begin position="1010"/>
            <end position="1104"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="1"/>
            <end position="34"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="1103"/>
            <end position="1135"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="19"/>
            <end position="28"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="1126"/>
            <end position="1135"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00235"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="Pfam" id="PF11701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007678.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1135" mass="123123" checksum="CEA661A259D1CDA4" modified="2019-05-08" version="1">MCEPGAQEQGFPGGIARSGEQHAGEGKHQRSRSKAWARALEGQYSQRCWPLRGLVLPASWPRPGRVTDPSAGRGSHCRVIEAVTNLFLHLSQMKLMSCTDSKVEQMFQILLDPKETDTDKKQKAAQNLIVLAREEAGAEKIFQSDGVRLLLRMLDTGREDAMLAALRTLAGLCSGHRSRTMAILAELGAPRVSAMLGVEHEQVSLAACNLLQVMFDALKEGLQRDFRGKEEALVLDPSKELKLLIGHLLEMLNREGTSAHGRDNILNLLIKVVPRKSLRDPNNSLTLWVIDQGLKKILEVGGTVCEAPGSLPATENSRMSAAVLLSKLYSDLKCDAERENFHRLCEDYVRSWFEGHGLAGKLRAIQTASCLLQGPSEAGNQVLELEGIMESVLALCASVREVDQLVAVEALIHAADKAKRASFITANGVTLLKNIYKCSGRDSIRIRALVGLCKLGSAGGIDFSMKQFAEGSTLKLAKQCRKWLCNEAIDAGTRRWAVEGLAYLTFDADVKEEFVEDKAAVQAMFQLAKSEDKSVLFAVASTLVNCTNSYDHEEPDPQMLELAKYAKQHVPEQHPKDEPGCVRRRVRKLLAAGVVSALTCMVKSENPALTNSCRELISRVFLAVVEEAADRGSVVAQGGGKALIPLSLEGTEVGQAKAAQALAKITITSNPDMAFPGERIYEVVRPLVRLLHLHRTGLENFEGLMALTNLAGISERLRQKILKEKAVPMIEGYMFEEHELLRLAATECMCNMAMSTEVQELFLAEGSDRLKLLVLYSGEDDEKLRRAASGTLAMLTSLLPQICGKITQVTAHWLEILQALLLSPNVELQHRGAVIVLNMMVADREIAAKLMERHGRLVLGGWLAGALPGHCQDLLPSESHVLLLRGTALEAWARGAGLCQGPSWQQQLHPGEAARPLPLAPCGYMIPRPPFLCPLPPALRAQKLVLGHEHVVLLGQDGTIYTWGCGRHGQLGHGGLEPALEPRLVEALHGVPMGDVAAGGWHSVSVSEGGDLYVWGWNESGQLALPSKALAEKRAPATASVSQHEEPCPGEAELKLVNHEAALGAEAADFISIQAFPALLDLPEGSEVSKVSCGSRHTAAVTLAAPQGSQSPPRYPEGDPALCSDMPQTLQPAPR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZQ1</accession>
    <name>A0A452GZQ1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Fibrillar collagen NC1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZQ1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008944.1">
        <property type="protein sequence ID" value="ENSGAGP00000007755.1"/>
        <property type="gene ID" value="ENSGAGG00000005830.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005581">
        <property type="term" value="C:collagen trimer"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005201">
        <property type="term" value="F:extracellular matrix structural constituent"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.1000:FF:000001">
        <property type="entry name" value="Collagen alpha-1 type I chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.1000">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008160">
        <property type="entry name" value="Collagen"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050938">
        <property type="entry name" value="Collagen_Structural_Proteins"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000885">
        <property type="entry name" value="Fib_collagen_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR37456:SF6">
        <property type="entry name" value="COLLAGEN ALPHA-1(XXIII) CHAIN-LIKE ISOFORM X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR37456">
        <property type="entry name" value="SI:CH211-266K2.1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01410">
        <property type="entry name" value="COLFI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01391">
        <property type="entry name" value="Collagen"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00038">
        <property type="entry name" value="COLFI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51461">
        <property type="entry name" value="NC1_FIB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0106">Calcium</keyword>
    <keyword evidence="7" id="KW-0176">Collagen</keyword>
    <keyword evidence="8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="9" id="KW-0379">Hydroxylation</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <feature type="domain" description="Fibrillar collagen NC1" evidence="11">
        <location>
            <begin position="846"/>
            <end position="1079"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1"/>
            <end position="295"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="324"/>
            <end position="849"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="74"/>
            <end position="87"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="255"/>
            <end position="265"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="10">
        <location>
            <begin position="365"/>
            <end position="374"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="375"/>
            <end position="385"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="409"/>
            <end position="420"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="486"/>
            <end position="495"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="10">
        <location>
            <begin position="506"/>
            <end position="515"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="691"/>
            <end position="712"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="725"/>
            <end position="739"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="10">
        <location>
            <begin position="752"/>
            <end position="761"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="10">
        <location>
            <begin position="791"/>
            <end position="806"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023119"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023278"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS51461"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007755.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1079" mass="105306" checksum="B53FB7A7B626C1B0" modified="2019-05-08" version="1">PGDPGPPGHPTHSGPDGRNPFAAQMAGLDEKSGISSQMGFMPGAVGEAGPTGARGPEGPQGQRGETGPPGPAGSQGLPGSPGTSGPPGLAGPPGSPGPQGSTGPPGIRGQAVIMSLSPWKSQYLGPHGPQGPIGSVGEEGKRGPRGDPGSVGPPGPVGERGAPGNRGFPGSDGLPGPKGAQGERGPAGSSGPKGSQGDPGRTGEPGLPGARGLTGNPGVQGPEGKIGPLGAPGEDGRPGPAGSIGIRGQPGSMGLPGPKGSSGDPGKPGEAGNAGVSGQRGAPGKDGEVGPSGPVGPPVLSLASCLNSSTYTMTFPFIFFQGLPGPPGPPGEAGKPGDQVTSFGERGNPGERGEPGAAGLQGEKGMAGGHGPDGPKGNPGPTGTPGDPGPPGLQGMPGERGIAGTPGPKGDRGGVGEKGSEGTAGNDGARVSNPIALLEIALSNGGEPGPRGLVGPPGSRGNPGPDGQPGVKGEPGEPGQKGDAGSPGPQGLSGSPGPPGPHGVPGLKGGRGTQGPPGATGFPGSAGRVGPPGPTGALGPAGPIGDPGKEGPPGLRGDPGAHGRVGDRGPAGPPGGPGDKGDSGEDGQPGPDGPPGPAGTTGQRGIVGMPGQRGERGMPGLPGPGGTPGKQGSTGPPGDKGPSGPIGQPGASGPVGEPGPEVSIVGDRGEPGPAGLPGAQGGPGTPGPVGPTGEAGQRGEPGSRGPVGPPGRAGKRGLPGPQGPRGDKGDHGDRGDRGQKGHRGFTGLQGLPGPPGPPGPVGPSGKDGNPGPHGPLGPPGVRGSLGEAGPEGPPGDPGPPGPPGPPGHLTAAIGDIMGHFDESMADPLPEFTEDEAAPDDNNKTDPGVHATLKSLSSQIETMRSPDGSKKHPARTCDDLKLCHPSKKSGEYWIDPNQGCVEDALKVYCNMETGETCISANPSSIPRKMWWTSRSPDLKPVWYGLDMNRGSQFVYGDSESPNTAVTQMTFLRLLSKEASQNITYHCKNSIGYMDDQSKNLKKAVILKGANDLEIKAEGNNRFRYTVLQDSCSKRNGNIGKTVFEYRTQNVARLPIVDIAPVDIGNTDQEFGIEIGPVCFV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZQ2</accession>
    <name>A0A452GZQ2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Tissue factor pathway inhibitor</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="11">Extrinsic pathway inhibitor</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="12">Lipoprotein-associated coagulation inhibitor</fullName>
        </alternativeName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Inhibits factor X (X(a)) directly and, in a Xa-dependent way, inhibits VIIa/tissue factor activity, presumably by forming a quaternary Xa/LACI/VIIa/TF complex. It possesses an antithrombotic action and also the ability to associate with lipoproteins in plasma.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZQ2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008812.1">
        <property type="protein sequence ID" value="ENSGAGP00000007635.1"/>
        <property type="gene ID" value="ENSGAGG00000006105.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004867">
        <property type="term" value="F:serine-type endopeptidase inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd22613">
        <property type="entry name" value="Kunitz_TFPI1_1-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd22614">
        <property type="entry name" value="Kunitz_TFPI1_2-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.410.10:FF:000004">
        <property type="entry name" value="Tissue factor pathway inhibitor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.410.10:FF:000012">
        <property type="entry name" value="Tissue factor pathway inhibitor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.410.10">
        <property type="entry name" value="Pancreatic trypsin inhibitor Kunitz domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002223">
        <property type="entry name" value="Kunitz_BPTI"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036880">
        <property type="entry name" value="Kunitz_BPTI_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020901">
        <property type="entry name" value="Prtase_inh_Kunz-CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050098">
        <property type="entry name" value="TFPI/VKTCI-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10083:SF374">
        <property type="entry name" value="BPTI_KUNITZ INHIBITOR DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10083">
        <property type="entry name" value="KUNITZ-TYPE PROTEASE INHIBITOR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00014">
        <property type="entry name" value="Kunitz_BPTI"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00759">
        <property type="entry name" value="BASICPTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00131">
        <property type="entry name" value="KU"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57362">
        <property type="entry name" value="BPTI-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00280">
        <property type="entry name" value="BPTI_KUNITZ_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50279">
        <property type="entry name" value="BPTI_KUNITZ_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51257">
        <property type="entry name" value="PROKAR_LIPOPROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="8" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="3" id="KW-0646">Protease inhibitor</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="6" id="KW-0722">Serine protease inhibitor</keyword>
    <keyword evidence="4 13" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="13">
        <location>
            <begin position="1"/>
            <end position="27"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5018973391" description="Tissue factor pathway inhibitor" evidence="13">
        <location>
            <begin position="28"/>
            <end position="248"/>
        </location>
    </feature>
    <feature type="domain" description="BPTI/Kunitz inhibitor" evidence="14">
        <location>
            <begin position="53"/>
            <end position="103"/>
        </location>
    </feature>
    <feature type="domain" description="BPTI/Kunitz inhibitor" evidence="14">
        <location>
            <begin position="125"/>
            <end position="175"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022690"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00057773"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00073658"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00081110"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00081787"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50279"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007635.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="248" mass="28203" checksum="98774F28785D604E" modified="2019-05-08" version="1">MKEMKRGCFLVTTLLLLLGCVPWHVTADSKDGEDDQDIPGAALPPLKLGNSICALKADGGECKAIYTRYYFNIQTRQCEMFEYGGCHGNKNNFLTLEECQEMCVVSEFPVKKTRGRFGKEKPNFCFMEQDPGICRGYFSRYFYNKESQQCEKFKYGGCLGNQNNFKALEECQIACQDNTNSLQLDQHAEHPIIMNNSSPVAKPSEFPRLLVVQEGTRNSWEETGLSHQVFLMVFCTCTHIYFLRLGDG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZQ3</accession>
    <name>A0A452GZQ3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">UNC-45/Cro1/She4 central domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZQ3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008867.1">
        <property type="protein sequence ID" value="ENSGAGP00000007683.1"/>
        <property type="gene ID" value="ENSGAGG00000006144.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051879">
        <property type="term" value="F:Hsp90 protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="entry name" value="Unc-45 myosin chaperone B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.30">
        <property type="entry name" value="Regulator of chromosome condensation 1/beta-lactamase-inhibitor protein II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009091">
        <property type="entry name" value="RCC1/BLIP-II"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000408">
        <property type="entry name" value="Reg_chr_condens"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024660">
        <property type="entry name" value="UCS_central_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45994">
        <property type="entry name" value="FI21225P1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45994:SF3">
        <property type="entry name" value="PROTEIN UNC-45 HOMOLOG A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00415">
        <property type="entry name" value="RCC1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11701">
        <property type="entry name" value="UNC45-central"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50985">
        <property type="entry name" value="RCC1/BLIP-II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00626">
        <property type="entry name" value="RCC1_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50012">
        <property type="entry name" value="RCC1_3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="UNC-45/Cro1/She4 central" evidence="5">
        <location>
            <begin position="267"/>
            <end position="455"/>
        </location>
    </feature>
    <feature type="repeat" description="RCC1" evidence="3">
        <location>
            <begin position="906"/>
            <end position="957"/>
        </location>
    </feature>
    <feature type="repeat" description="RCC1" evidence="3">
        <location>
            <begin position="958"/>
            <end position="1052"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="1"/>
            <end position="34"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="19"/>
            <end position="28"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00235"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="Pfam" id="PF11701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007683.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1097" mass="119098" checksum="EE82CB028BE251CD" modified="2019-05-08" version="1">MCEPGAQEQGFPGGIARSGEQHAGEGKHQRSRSKAWARALEGQYSQRCWPLRGLVLPASWPRPGRVTDPSAGRGSHCRVIEAVTNLFLHLSQMKLMSCTDSKVEQMFQILLDPKETDTDKKQKAAQNLIVLAREEAGAEKIFQSDGVRLLLRMLDTGREDAMLAALRTLAGLCSGHRSRTMAILAELGAPRVSAMLGVEHEQVSLAACNLLQVMFDALKEGLQRDFRGKEEALVLDPSKELKLLIGHLLEMLNREGTSAHGRDNILNLLIKVVPRKSLRDPNNSLTLWVIDQGLKKILEVGGTVCEAPGSLPATENSRMSAAVLLSKLYSDLKCDAERENFHRLCEDYVRSWFEGHGLAGKLRAIQTASCLLQGPSEAGNQVLELEGIMESVLALCASVREVDQLVAVEALIHAADKAKRASFITANGVTLLKNIYKCSGRDSIRIRALVGLCKLGSAGGIDFSMKQFAEGSTLKLAKQCRKWLCNEAIDAGTRRWAVEGLAYLTFDADVKEEFVEDKAAVQAMFQLAKSEDKSVLFAVASTLVNCTNSYDHEEPDPQMLELAKYAKQHVPEQHPKDEPGCVRRRVRKLLAAGVVSALTCMVKSENPALTNSCRELISRVFLAVVEEAADRGSVVAQGGGKALIPLSLEGTEVGQAKAAQALAKITITSNPDMAFPGERIYEVVRPLVRLLHLHRTGLENFEGLMALTNLAGISERLRQKILKEKAVPMIEGYMFEEHELLRLAATECMCNMAMSTETAHWLEILQALLLSPNVELQHRGAVIVLNMMVADREIAAKLMERHGRLVLGGWLAGALPGHCQDLLPSESHVLLLRGTALEAWARGAGLCQGPSWQQQLHPGEAARPLPLAPCGYMIPRPPFLCPLPPALRAQKLVLGHEHVVLLGQDGTIYTWGCGRHGQLGHGGLEPALEPRLVEALHGVPMGDVAAGGWHSVSVSEGGDLYVWGWNESGQLALPSKALAEKRAPATASVSQHEEPCPGEAELKLVNHEAALGAEAADFISIQAFPALLDLPEGSEVSKVSCGSRHTAAVTRKYGQLGHNDTASSDQPRQVGYFPANGLTVEDVVCGPWNTYVGAVEK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZQ6</accession>
    <name>A0A452GZQ6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Fibrillar collagen NC1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZQ6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008949.1">
        <property type="protein sequence ID" value="ENSGAGP00000007760.1"/>
        <property type="gene ID" value="ENSGAGG00000005830.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005581">
        <property type="term" value="C:collagen trimer"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031012">
        <property type="term" value="C:extracellular matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030020">
        <property type="term" value="F:extracellular matrix structural constituent conferring tensile strength"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030198">
        <property type="term" value="P:extracellular matrix organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="entry name" value="Collagen alpha-1 type I chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.1000">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008160">
        <property type="entry name" value="Collagen"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050149">
        <property type="entry name" value="Collagen_superfamily"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000885">
        <property type="entry name" value="Fib_collagen_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24023">
        <property type="entry name" value="COLLAGEN ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24023:SF1109">
        <property type="entry name" value="COLLAGEN ALPHA-4(IV) CHAIN-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01410">
        <property type="entry name" value="COLFI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01391">
        <property type="entry name" value="Collagen"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SMART" id="SM00038">
        <property type="entry name" value="COLFI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51461">
        <property type="entry name" value="NC1_FIB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0106">Calcium</keyword>
    <keyword evidence="7" id="KW-0176">Collagen</keyword>
    <keyword evidence="8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="9" id="KW-0379">Hydroxylation</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <feature type="domain" description="Fibrillar collagen NC1" evidence="11">
        <location>
            <begin position="828"/>
            <end position="1061"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1"/>
            <end position="295"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="311"/>
            <end position="831"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="42"/>
            <end position="53"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="92"/>
            <end position="105"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="273"/>
            <end position="283"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="10">
        <location>
            <begin position="314"/>
            <end position="323"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="325"/>
            <end position="336"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="10">
        <location>
            <begin position="347"/>
            <end position="356"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="357"/>
            <end position="367"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="391"/>
            <end position="402"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="468"/>
            <end position="477"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="10">
        <location>
            <begin position="488"/>
            <end position="497"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="673"/>
            <end position="694"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="707"/>
            <end position="721"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="10">
        <location>
            <begin position="734"/>
            <end position="743"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="10">
        <location>
            <begin position="773"/>
            <end position="788"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023119"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023278"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS51461"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007760.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1061" mass="103516" checksum="1496FE35875E4922" modified="2019-05-08" version="1">PGDPGPPGHPTHSGPDGRNPFAAQMAGLDEKSGISSQMGFMPGAVGPAGARGPEGPPGKPGEDGEAGPTGARGPEGPQGQRGETGPPGPAGSQGLPGSPGTSGPPGLAGPPGSPGPQGSTGPPGIRGQAVIMSLSPWKSQYLGPHGPQGPIGSVGEEGKRGPRGDPGSVGPPGPVGERGAPGNRGFPGSDGLPGPKGAQGERGPAGSSGPKGSQGDPGRTGEPGLPGARGLTGNPGVQGPEGKIGPLGAPGEDGRPGPAGSIGIRGQPGSMGLPGPKGSSGDPGKPGEAGNAGVSGQRCFPSNLYFFSYQGVPGDPGAGGPLGPRGERGNPGERGEPGAAGLQGEKGMAGGHGPDGPKGNPGPTGTPGDPGPPGLQGMPGERGIAGTPGPKGDRGGVGEKGSEGTAGNDGARVSNPIALLEIALSNGGEPGPRGLVGPPGSRGNPGPDGQPGVKGEPGEPGQKGDAGSPGPQGLSGSPGPPGPHGVPGLKGGRGTQGPPGATGFPGSAGRVGPPGPTGALGPAGPIGDPGKEGPPGLRGDPGAHGRVGDRGPAGPPGGPGDKGDSGEDGQPGPDGPPGPAGTTGQRGIVGMPGQRGERGMPGLPGPGGTPGKQGSTGPPGDKGPSGPIGQPGASGPVGEPGPEVSIVGDRGEPGPAGLPGAQGGPGTPGPVGPTGEAGQRGEPGSRGPVGPPGRAGKRGLPGPQGPRGDKGDHGDRGDRGQKGHRGFTGLQGLPGPPGPPGPVGPSGKDGNPGPHGPLGPPGVRGSLGEAGPEGPPGDPGPPGPPGPPGHLTAAIGDIMGHFDESMADPLPEFTEDEAAPDDNNKTDPGVHATLKSLSSQIETMRSPDGSKKHPARTCDDLKLCHPSKKSGEYWIDPNQGCVEDALKVYCNMETGETCISANPSSIPRKMWWTSRSPDLKPVWYGLDMNRGSQFVYGDSESPNTAVTQMTFLRLLSKEASQNITYHCKNSIGYMDDQSKNLKKAVILKGANDLEIKAEGNNRFRYTVLQDSCSKRNGNIGKTVFEYRTQNVARLPIVDIAPVDIGNTDQEFGIEIGPVCFV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZQ8</accession>
    <name>A0A452GZQ8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Nuclear receptor subfamily 1 group I member 3</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="16">Constitutive androstane receptor</fullName>
        </alternativeName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Binds and transactivates the retinoic acid response elements that control expression of the retinoic acid receptor beta 2 and alcohol dehydrogenase 3 genes. Transactivates both the phenobarbital responsive element module of the human CYP2B6 gene and the CYP3A4 xenobiotic response element.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="17">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="17">Belongs to the nuclear hormone receptor family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZQ8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008866.1">
        <property type="protein sequence ID" value="ENSGAGP00000007682.1"/>
        <property type="gene ID" value="ENSGAGG00000006024.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004879">
        <property type="term" value="F:nuclear receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000122">
        <property type="term" value="P:negative regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045944">
        <property type="term" value="P:positive regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07156">
        <property type="entry name" value="NR_DBD_VDR_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.50.10:FF:000044">
        <property type="entry name" value="retinoic acid receptor beta isoform X4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.50.10">
        <property type="entry name" value="Erythroid Transcription Factor GATA-1, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.565.10">
        <property type="entry name" value="Retinoid X Receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035500">
        <property type="entry name" value="NHR-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000536">
        <property type="entry name" value="Nucl_hrmn_rcpt_lig-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050234">
        <property type="entry name" value="Nuclear_hormone_rcpt_NR1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001723">
        <property type="entry name" value="Nuclear_hrmn_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001628">
        <property type="entry name" value="Znf_hrmn_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013088">
        <property type="entry name" value="Znf_NHR/GATA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24082">
        <property type="entry name" value="NUCLEAR HORMONE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24082:SF231">
        <property type="entry name" value="NUCLEAR RECEPTOR SUBFAMILY 1 GROUP I MEMBER 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00104">
        <property type="entry name" value="Hormone_recep"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00105">
        <property type="entry name" value="zf-C4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00398">
        <property type="entry name" value="STRDHORMONER"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00047">
        <property type="entry name" value="STROIDFINGER"/>
    </dbReference>
    <dbReference type="SMART" id="SM00430">
        <property type="entry name" value="HOLI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00399">
        <property type="entry name" value="ZnF_C4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57716">
        <property type="entry name" value="Glucocorticoid receptor-like (DNA-binding domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48508">
        <property type="entry name" value="Nuclear receptor ligand-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51843">
        <property type="entry name" value="NR_LBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00031">
        <property type="entry name" value="NUCLEAR_REC_DBD_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51030">
        <property type="entry name" value="NUCLEAR_REC_DBD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0010">Activator</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="12" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="8 17" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="4 17" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13 17" id="KW-0539">Nucleus</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="11 17" id="KW-0675">Receptor</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10 17" id="KW-0804">Transcription</keyword>
    <keyword evidence="7 17" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6 17" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 17" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="Nuclear receptor" evidence="18">
        <location>
            <begin position="22"/>
            <end position="97"/>
        </location>
    </feature>
    <feature type="domain" description="NR LBD" evidence="19">
        <location>
            <begin position="127"/>
            <end position="370"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00037362"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00040912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00043125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="RuleBase" id="RU004334"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS51030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS51843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007682.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="370" mass="41762" checksum="EA5AC136A42DD762" modified="2019-05-08" version="1">MGPTDGEPSPGGGPGAQPEGEEKICAVCGDRATGYHFHVMTCEGCKGFFRRSISKGVCFTCPFTRNCTITKAKRRQCQACRLQKCLAVGMRKDMIMSEEALQMRRVLRKRKQQEREPAGEPAELTEEQEQLISILIEAHKRHFDSGFSQFVHCRPPVRLYIHSLSPQNPQEPRGGPMAPLQSLPGDMLPDIFSMLPHFADLSTILIQQVIKFAKEIPAFRSAWLGSHTQAAQTAWLEGEWETGLPHHPSSLSPAGFQQIYLEPLLKFHISLRKLRLHEAEYVLLLAIALFSPDHPGITQRHTVDQLQEKMALTLKSYIDQRHPLPEGRLLYAKLLLLLTELRTLKVENTRQILHIQDLTAMTPLLSEIIS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZR5</accession>
    <name>A0A452GZR5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">IMP dehydrogenase/GMP reductase domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GZR5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008962.1">
        <property type="protein sequence ID" value="ENSGAGP00000007773.1"/>
        <property type="gene ID" value="ENSGAGG00000006197.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016491">
        <property type="term" value="F:oxidoreductase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.20.20.70">
        <property type="entry name" value="Aldolase class I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013785">
        <property type="entry name" value="Aldolase_TIM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050139">
        <property type="entry name" value="GMP_reductase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43170">
        <property type="entry name" value="GMP REDUCTASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43170:SF3">
        <property type="entry name" value="GMP REDUCTASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51412">
        <property type="entry name" value="Inosine monophosphate dehydrogenase (IMPDH)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007773.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="4">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="119" mass="13323" checksum="B23F02BA73A91CC5" modified="2019-05-08" version="1">MEIAKVAPGDSPMLRWLGKPMPHVEADLKLDFHDVLLRPKRSSLKSRAEVDLIRTFTFRNSKQTYTGIPIMVANMDTVGTFEMAKCLGSLHCCCLCWILFSGPSILKVANPVSFKKTSH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZR9</accession>
    <name>A0A452GZR9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Nuclear receptor subfamily 1 group I member 3</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="16">Constitutive androstane receptor</fullName>
        </alternativeName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Binds and transactivates the retinoic acid response elements that control expression of the retinoic acid receptor beta 2 and alcohol dehydrogenase 3 genes. Transactivates both the phenobarbital responsive element module of the human CYP2B6 gene and the CYP3A4 xenobiotic response element.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="17">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="17">Belongs to the nuclear hormone receptor family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZR9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008876.1">
        <property type="protein sequence ID" value="ENSGAGP00000007692.1"/>
        <property type="gene ID" value="ENSGAGG00000006024.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004879">
        <property type="term" value="F:nuclear receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000122">
        <property type="term" value="P:negative regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045944">
        <property type="term" value="P:positive regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07156">
        <property type="entry name" value="NR_DBD_VDR_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.50.10:FF:000044">
        <property type="entry name" value="retinoic acid receptor beta isoform X4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.50.10">
        <property type="entry name" value="Erythroid Transcription Factor GATA-1, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.565.10">
        <property type="entry name" value="Retinoid X Receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035500">
        <property type="entry name" value="NHR-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000536">
        <property type="entry name" value="Nucl_hrmn_rcpt_lig-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050234">
        <property type="entry name" value="Nuclear_hormone_rcpt_NR1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001723">
        <property type="entry name" value="Nuclear_hrmn_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001628">
        <property type="entry name" value="Znf_hrmn_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013088">
        <property type="entry name" value="Znf_NHR/GATA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24082">
        <property type="entry name" value="NUCLEAR HORMONE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24082:SF231">
        <property type="entry name" value="NUCLEAR RECEPTOR SUBFAMILY 1 GROUP I MEMBER 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00104">
        <property type="entry name" value="Hormone_recep"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00105">
        <property type="entry name" value="zf-C4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00398">
        <property type="entry name" value="STRDHORMONER"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00047">
        <property type="entry name" value="STROIDFINGER"/>
    </dbReference>
    <dbReference type="SMART" id="SM00430">
        <property type="entry name" value="HOLI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00399">
        <property type="entry name" value="ZnF_C4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57716">
        <property type="entry name" value="Glucocorticoid receptor-like (DNA-binding domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48508">
        <property type="entry name" value="Nuclear receptor ligand-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51843">
        <property type="entry name" value="NR_LBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00031">
        <property type="entry name" value="NUCLEAR_REC_DBD_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51030">
        <property type="entry name" value="NUCLEAR_REC_DBD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0010">Activator</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="12" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="8 17" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="4 17" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13 17" id="KW-0539">Nucleus</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="11 17" id="KW-0675">Receptor</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10 17" id="KW-0804">Transcription</keyword>
    <keyword evidence="7 17" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6 17" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 17" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="Nuclear receptor" evidence="19">
        <location>
            <begin position="35"/>
            <end position="110"/>
        </location>
    </feature>
    <feature type="domain" description="NR LBD" evidence="20">
        <location>
            <begin position="140"/>
            <end position="370"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1"/>
            <end position="31"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="18">
        <location>
            <begin position="10"/>
            <end position="24"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00037362"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00040912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00043125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="RuleBase" id="RU004334"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS51030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS51843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007692.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="370" mass="41625" checksum="4A5D3E129943C95F" modified="2019-05-08" version="1">MLSPSDRESSSSSSSPSVLSLGAGSREDGEPEGEEKICAVCGDRATGYHFHVMTCEGCKGFFRRSISKGVCFTCPFTRNCTITKAKRRQCQACRLQKCLAVGMRKDMIMSEEALQMRRVLRKRKQQEREPAGEPAELTEEQEQLISILIEAHKRHFDSGFSQFVHCRVRTPTAASPAGTRASLPGDMLPDIFSMLPHFADLSTILIQQVIKFAKEIPAFRSAWLGSHTQAAQTAWLEGEWETGLPHHPSSLSPAGFQQIYLEPLLKFHISLRKLRLHEAEYVLLLAIALFSPDHPGITQRHTVDQLQEKMALTLKSYIDQRHPLPEGRLLYAKLLLLLTELRTLKVENTRQILHIQDLTAMTPLLSEIIS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZS5</accession>
    <name>A0A452GZS5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="22">Voltage-dependent T-type calcium channel subunit alpha</fullName>
        </recommendedName>
    </protein>
    <organism evidence="27 28">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="28" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="27" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="27" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="22">Voltage-sensitive calcium channels (VSCC) mediate the entry of calcium ions into excitable cells and are also involved in a variety of calcium-dependent processes, including muscle contraction, hormone or neurotransmitter release, gene expression, cell motility, cell division and cell death. This channel gives rise to T-type calcium currents. T-type calcium channels belong to the "low-voltage activated (LVA)" group and are strongly blocked by nickel and mibefradil. A particularity of this type of channels is an opening at quite negative potentials, and a voltage-dependent inactivation. T-type channels serve pacemaking functions in both central neurons and cardiac nodal cells and support calcium signaling in secretory cells and vascular smooth muscle. They may also be involved in the modulation of firing patterns of neurons which is important for information processing as well as in cell growth processes.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>Ca(2+)(in) = Ca(2+)(out)</text>
            <dbReference type="Rhea" id="RHEA:29671"/>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="22">Membrane</location>
            <topology evidence="22">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="19">Belongs to the calcium channel alpha-1 subunit (TC 1.A.1.11) family. CACNA1G subfamily.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000008972.1">
        <property type="protein sequence ID" value="ENSGAGP00000007783.1"/>
        <property type="gene ID" value="ENSGAGG00000006199.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005891">
        <property type="term" value="C:voltage-gated calcium channel complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008331">
        <property type="term" value="F:high voltage-gated calcium channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008332">
        <property type="term" value="F:low voltage-gated calcium channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098703">
        <property type="term" value="P:calcium ion import across plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.70:FF:000014">
        <property type="entry name" value="Voltage-dependent T-type calcium channel subunit alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.70:FF:000018">
        <property type="entry name" value="Voltage-dependent T-type calcium channel subunit alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.70:FF:000029">
        <property type="entry name" value="Voltage-dependent T-type calcium channel subunit alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.70:FF:000032">
        <property type="entry name" value="Voltage-dependent T-type calcium channel subunit alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Voltage-dependent T-type calcium channel subunit alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.350:FF:000008">
        <property type="entry name" value="Voltage-dependent T-type calcium channel subunit alpha"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Voltage-dependent T-type calcium channel subunit alpha"/>
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        <property type="entry name" value="Voltage-dependent T-type calcium channel subunit alpha"/>
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        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.350">
        <property type="entry name" value="Voltage-gated potassium channels. Chain C"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005821">
        <property type="entry name" value="Ion_trans_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050599">
        <property type="entry name" value="VDCC_alpha-1_subunit"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005445">
        <property type="entry name" value="VDCC_T_a1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002077">
        <property type="entry name" value="VDCCAlpha1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027359">
        <property type="entry name" value="Volt_channel_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45628">
        <property type="entry name" value="VOLTAGE-DEPENDENT CALCIUM CHANNEL TYPE A SUBUNIT ALPHA-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45628:SF33">
        <property type="entry name" value="VOLTAGE-DEPENDENT T-TYPE CALCIUM CHANNEL SUBUNIT ALPHA-1G"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00520">
        <property type="entry name" value="Ion_trans"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00167">
        <property type="entry name" value="CACHANNEL"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01629">
        <property type="entry name" value="TVDCCALPHA1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81324">
        <property type="entry name" value="Voltage-gated potassium channels"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="11 20 22" id="KW-0106">Calcium</keyword>
    <keyword evidence="8 22" id="KW-0107">Calcium channel</keyword>
    <keyword evidence="7 22" id="KW-0109">Calcium transport</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="23" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="16 21" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="17" id="KW-0407">Ion channel</keyword>
    <keyword evidence="14" id="KW-0406">Ion transport</keyword>
    <keyword evidence="15 25" id="KW-0472">Membrane</keyword>
    <keyword evidence="20" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="28" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0677">Repeat</keyword>
    <keyword evidence="9 25" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="13 25" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <keyword evidence="12 22" id="KW-0851">Voltage-gated channel</keyword>
    <feature type="transmembrane region" description="Helical" evidence="25">
        <location>
            <begin position="36"/>
            <end position="53"/>
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    <feature type="transmembrane region" description="Helical" evidence="25">
        <location>
            <begin position="73"/>
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    <feature type="transmembrane region" description="Helical" evidence="25">
        <location>
            <begin position="164"/>
            <end position="187"/>
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    <feature type="transmembrane region" description="Helical" evidence="25">
        <location>
            <begin position="289"/>
            <end position="310"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="25">
        <location>
            <begin position="316"/>
            <end position="341"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="25">
        <location>
            <begin position="711"/>
            <end position="729"/>
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    </feature>
    <feature type="transmembrane region" description="Helical" evidence="25">
        <location>
            <begin position="741"/>
            <end position="762"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="25">
        <location>
            <begin position="832"/>
            <end position="851"/>
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    </feature>
    <feature type="transmembrane region" description="Helical" evidence="25">
        <location>
            <begin position="908"/>
            <end position="931"/>
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    <feature type="transmembrane region" description="Helical" evidence="25">
        <location>
            <begin position="1244"/>
            <end position="1262"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="25">
        <location>
            <begin position="1282"/>
            <end position="1303"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="25">
        <location>
            <begin position="1379"/>
            <end position="1401"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="25">
        <location>
            <begin position="1480"/>
            <end position="1504"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="25">
        <location>
            <begin position="1577"/>
            <end position="1596"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="25">
        <location>
            <begin position="1608"/>
            <end position="1629"/>
        </location>
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    <feature type="transmembrane region" description="Helical" evidence="25">
        <location>
            <begin position="1678"/>
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    <feature type="transmembrane region" description="Helical" evidence="25">
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            <begin position="1711"/>
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        </location>
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    <feature type="transmembrane region" description="Helical" evidence="25">
        <location>
            <begin position="1795"/>
            <end position="1816"/>
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        <location>
            <begin position="31"/>
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        </location>
    </feature>
    <feature type="domain" description="Ion transport" evidence="26">
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            <begin position="710"/>
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        <location>
            <begin position="1242"/>
            <end position="1514"/>
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    <feature type="domain" description="Ion transport" evidence="26">
        <location>
            <begin position="1577"/>
            <end position="1827"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="24">
        <location>
            <begin position="413"/>
            <end position="499"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="24">
        <location>
            <begin position="1045"/>
            <end position="1128"/>
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    </feature>
    <feature type="region of interest" description="Disordered" evidence="24">
        <location>
            <begin position="1154"/>
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    <feature type="region of interest" description="Disordered" evidence="24">
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            <begin position="2004"/>
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    <feature type="region of interest" description="Disordered" evidence="24">
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            <begin position="2125"/>
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        <location>
            <begin position="1510"/>
            <end position="1540"/>
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        <location>
            <begin position="426"/>
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    </feature>
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            <begin position="469"/>
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            <begin position="484"/>
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            <begin position="1052"/>
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        <location>
            <position position="301"/>
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        <ligand>
            <name>Ca(2+)</name>
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        <location>
            <position position="890"/>
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        <ligand>
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        <location>
            <position position="1453"/>
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        <ligand>
            <name>Ca(2+)</name>
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        <location>
            <position position="253"/>
        </location>
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        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
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    </evidence>
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            <dbReference type="ARBA" id="ARBA00022490"/>
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    </evidence>
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            <dbReference type="ARBA" id="ARBA00022553"/>
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            <dbReference type="ARBA" id="ARBA00022568"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022673"/>
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    </evidence>
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            <dbReference type="ARBA" id="ARBA00022692"/>
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    </evidence>
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    </evidence>
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            <dbReference type="ARBA" id="ARBA00022837"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022882"/>
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    </evidence>
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            <dbReference type="ARBA" id="ARBA00022989"/>
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    </evidence>
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        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00036634"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00061006"/>
        </source>
    </evidence>
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            <dbReference type="PIRSR" id="PIRSR602077-1"/>
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        <source>
            <dbReference type="PIRSR" id="PIRSR602077-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="RuleBase" id="RU003808"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="SAM" id="Coils"/>
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    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
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            <dbReference type="SAM" id="Phobius"/>
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            <dbReference type="Pfam" id="PF00520"/>
        </source>
    </evidence>
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            <dbReference type="Ensembl" id="ENSGAGP00000007783.1"/>
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            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2384" mass="267388" checksum="DBA58DC97A43232A" modified="2019-05-08" version="1">MGEHDVSSFSRPFPACVLKGLLISLTDLLCTWFERLSMLVILLNCVTLGMFHPCEDMACDSPRCRILQSFDDFIFAFFAVEMIVKMIALGIFGKKCYLGDTWNRLDFFIVIAGMLEYSLDLQNVSFSAVRTVRVLRPLRAINRVPSMRILVTLLLDTLPMLGNVLLLCFFVFFIFGIVGVQLWAGLLRNRCFLPENFSLPYTVELERYYQTENEDENPFICSQPRENGMRYCRSIPTRREEGLECTLDFYAYNDTTNTSCVNWNQYYTNCSAGEHNPFKGAINFDNIGYAWIAIFQVITLEGWVDIMYFVMDAHSFYNFIYFILLIIVGSFFMINLCLVVIATQFSETKQRESQLMKEQRIRYLSNASTLASFSEPGSCYDELLKYLVYVARKASKQLVKGYREAGFKMGLLSSPRNKSGAERQPCRHRQRKRSSVHHLIHHHHHHHHHYHLGNGNLRAPRASPEISDVETSSLHNGTNRLMLPPTTSNPHPTPNAAASNTESVHSIYHADCHFEPIHCRSSLPQPGLSLPSPEGLPKNVVGRKVYPTVHPSTSHEMLKEKNVADSTVNPGASMLTNLNIPPGPYSTMHKLLETQSTGFFSVHSTDTGAGFPGACKSSCKISNQCGKLDSGSCNPDSCPYCIKTLARDLELTDYETADSDSEGVYEFTQDAHYSDQRDPQRGKTKGRKTSQILAFWKVVCETFRKIVDSKYFGRGIMIAILINTLSMGIEYHEQPEELTNALEISNIVFTSLFALEMLLKVLVYGPFGYIKNPYNIFDGIIVVISVWEIVGQQGGGLSVLRTFRLMRVLKLVRFMPALQRQLVVLMKTMDNVATFCMLLMLFIFIFSILGMHLFGCKFASERDGDTLPDRKNFDSLLWAIVTVFQILTQEDWNKVLYNGMASTSSWAALYFIALMTFGNYVLFNLLVAILVEGFQTEEITKREDASGQLSCIQLPVDSAVGDASKSDSEGDLFPHSLEEEGGLKKNLSNPALMVLSDLPELKKSLTPPLIIHTAATPMPMPKSAMFGDAARGYDSRRASSVSMDPNAYELKSPSSIQSSPHSPWSAASSWNSRRSSWNSLGRAPSLKRRSQSGERKSLLSGNGKESSEEGESSDEERSSRAGSVNGSLPHRMASLETKGSFDLQDTLQVPSLYRTSSMHSSRTAASEHQDCNGKSSPGVLLQQLPLDDDDGDDEGNLSKSERVKAWIRAHLPSWCKERDSWSIYIFAPHSKFRLMCNKIITHKMFDHIVLVIIFLNCITIAMERPKIEPHSAERIFLTLSNYIFTVIFLAEMTVKVVALGLCFGEKAYLKSSWNVLDGVLVLISVIDILVSMVSDSGTKILGMLRVLRLLRTLRPLRVISRAQGLKLVVETLMSSLKPIGNIVVICCAFFIIFGILGVQLFKGKFFVCQGEDTRNITNKSDCAEASYKWVRHKYNFDNLGQALMSLFVLASKDGWVDIMYDGLDAVGVDQQPVMNYNPWMLLYFISFLLIVAFFVLNMFVGVVVENFHKCRQHQEEEEAKRREEKRLRRLEKKRRNLMLDDVIMESSASAVQEAQCKPYYSDYSRFRLLIHQMCTSHYLDLFITGVIGLNVITMAMEHYQQPKVLDEALKICNYIFTVIFVLESVFKLIAFGFRRFFQDRWNQLDLAIVLLSIMGITLEEIEVNASLPINPTIIRIMRVLRIARVLKLLKMAVGMRALLDTVMQALPQVGNLGLLFMLLFFIFAALGVELFGDLECDDTHHCEGLGRHATFRNFGMAFLTLFRVSTGDNWNGIMKDTLRDCDPESTCYNTVISPMYFVSFVLTAQFVLVNVVIAVLMKHLEESNKEAKEEAELEAELEMEMKTITPGPIPASDLFAWTGGNGGERPESPRGCTNPMQFKVDSQLSLFYPMERHLFDTISLLIQESLEGELKLMDNLSGSVCHHYALPAPEYYNSEKQIPLAEMEALSLTSDILSEKSWSLALTDDSFPDDTNTLLLNALESNTDHLSRDDTLTLSPSKDLLSVRKPSVGRTHSLPNDSYMFQPPNAGSCANSQAARNASHQKSQSGSTASVQSQPANTSSLLQIPKDHFHHIRPHGNLDWESKSKIPPTVHSPSAQRLLRRQVAIRNDSLDMYCSESKENLHAEVNELSDTNVPAGSTEDPSEASESACWGQSSVRTQHSHNQYNISKHTPALCACTDSYQETLRDSIDQDVSEINSSLEPFSSGTCTALSACSVVPPLTPKRNIGNSSNVTLKDLKKYYSVDTQGLLRKPSSWLDDQRRHSIEICSMQNSPQHHSTSSSSGFISQVLSEMEGLQGARQKKKLSPPCISIDPPDGQSLLPRGSHGISSPSADICLRRRAPSCDSKDSMDIGDSLLPDSMSTSPTPKTDLLTLPSFSFDQAEMDP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZS7</accession>
    <name>A0A452GZS7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Transcription factor ETV6</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="12">Transcriptional repressor; binds to the DNA sequence 5'-CCGGAAGT-3'. Plays a role in hematopoiesis and malignant transformation.</text>
    </comment>
    <comment type="subunit">
        <text evidence="13">Can form homodimers or heterodimers with TEL2 or FLI1. Interacts with L3MBTL1 and HDAC9.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 15">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 15">Belongs to the ETS family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZS7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008764.1">
        <property type="protein sequence ID" value="ENSGAGP00000007592.1"/>
        <property type="gene ID" value="ENSGAGG00000006072.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000977">
        <property type="term" value="F:RNA polymerase II transcription regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd08535">
        <property type="entry name" value="SAM_PNT-Tel_Yan"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.50:FF:000030">
        <property type="entry name" value="transcription factor ETV6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.10:FF:000176">
        <property type="entry name" value="transcription factor ETV6 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.50">
        <property type="entry name" value="Transcription Factor, Ets-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.10">
        <property type="entry name" value="Winged helix-like DNA-binding domain superfamily/Winged helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000418">
        <property type="entry name" value="Ets_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046328">
        <property type="entry name" value="ETS_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003118">
        <property type="entry name" value="Pointed_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013761">
        <property type="entry name" value="SAM/pointed_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036388">
        <property type="entry name" value="WH-like_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036390">
        <property type="entry name" value="WH_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11849">
        <property type="entry name" value="ETS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11849:SF19">
        <property type="entry name" value="TRANSCRIPTION FACTOR ETV6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00178">
        <property type="entry name" value="Ets"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02198">
        <property type="entry name" value="SAM_PNT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00454">
        <property type="entry name" value="ETSDOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00413">
        <property type="entry name" value="ETS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00251">
        <property type="entry name" value="SAM_PNT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47769">
        <property type="entry name" value="SAM/Pointed domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46785">
        <property type="entry name" value="Winged helix' DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00346">
        <property type="entry name" value="ETS_DOMAIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50061">
        <property type="entry name" value="ETS_DOMAIN_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51433">
        <property type="entry name" value="PNT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0007">Acetylation</keyword>
    <keyword evidence="9 15" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="4" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="11 15" id="KW-0539">Nucleus</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0678">Repressor</keyword>
    <keyword evidence="10" id="KW-0804">Transcription</keyword>
    <keyword evidence="8" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="PNT" evidence="18">
        <location>
            <begin position="34"/>
            <end position="118"/>
        </location>
    </feature>
    <feature type="domain" description="ETS" evidence="17">
        <location>
            <begin position="333"/>
            <end position="412"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="150"/>
            <end position="252"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005562"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022491"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00055208"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00063414"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00067754"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="RuleBase" id="RU004019"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50061"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS51433"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007592.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="412" mass="48242" checksum="C18AB059468D9F5D" modified="2019-05-08" version="1">MSHGQQERISYTPPASPVVNYPSSSPLHVPVPRAIRMEEDAIRLPAHLRLQPVYWSRDDVAQWLKWAEKEFSLRSIDSNSFEMNGKALLLLTKEDFRYRSPHSGDVLYELLQHILKQRKARMLFSPFFHPGNSIHAQPEVILHQNHDEDNFVQRPSRPSAETVHHNPPTIELLHRSRSPITTNHRPSPDPEQRSLKSPLDNTIRRLSPADRVPGTRHHQENNHQEPYPLSVSPIENNHCPPPPEVLQKPSSPRQENTRVIQLMPSPIMHPLILNPRHSVEFKPSRLSEDGLHREVKPINLSHREELAYMNHIMVSVSPPEEHAMPIGRIADCRLLWDYVYQLLSDSRYENFIRWEDKESKIFRIVDPNGLARLWGNHKNRTNMTYEKMSRALRHYYKLNIIRKEPGQRLLFR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZS8</accession>
    <name>A0A452GZS8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Alpha-mannosidase</fullName>
            <ecNumber evidence="8">3.2.1.-</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="6">Catalyzes the first committed step in the biosynthesis of complex N-glycans. It controls conversion of high mannose to complex N-glycans; the final hydrolytic step in the N-glycan maturation pathway.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="7">
            <text>N(4)-{beta-D-GlcNAc-(1-&gt;2)-alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;6)]-alpha-D-Man-(1-&gt;6)]-beta-D-Man-(1-&gt;4)-beta-D-GlcNAc-(1-&gt;4)-beta-D-GlcNAc}-L-asparaginyl-[protein] + 2 H2O = 2 alpha-D-mannopyranose + an N(4)-{beta-D-GlcNAc-(1-&gt;2)-alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;6)]-beta-D-Man-(1-&gt;4)-beta-D-GlcNAc-(1-&gt;4)-beta-D-GlcNAc}-L-asparaginyl-[protein]</text>
            <dbReference type="Rhea" id="RHEA:56052"/>
            <dbReference type="Rhea" id="RHEA-COMP:14368"/>
            <dbReference type="Rhea" id="RHEA-COMP:14369"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:28729"/>
            <dbReference type="ChEBI" id="CHEBI:60615"/>
            <dbReference type="ChEBI" id="CHEBI:60625"/>
            <dbReference type="EC" id="3.2.1.114"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="8">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
        <text evidence="8">Binds 1 zinc ion per subunit.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1 8">Belongs to the glycosyl hydrolase 38 family.</text>
    </comment>
    <dbReference type="EC" id="3.2.1.-" evidence="8"/>
    <dbReference type="AlphaFoldDB" id="A0A452GZS8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008842.1">
        <property type="protein sequence ID" value="ENSGAGP00000007660.1"/>
        <property type="gene ID" value="ENSGAGG00000002370.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000139">
        <property type="term" value="C:Golgi membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030246">
        <property type="term" value="F:carbohydrate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004572">
        <property type="term" value="F:mannosyl-oligosaccharide 1,3-1,6-alpha-mannosidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006013">
        <property type="term" value="P:mannose metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006491">
        <property type="term" value="P:N-glycan processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1270.50:FF:000001">
        <property type="entry name" value="Alpha-mannosidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.20.110.10:FF:000003">
        <property type="entry name" value="Alpha-mannosidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.20.110.10">
        <property type="entry name" value="Glycoside hydrolase 38, N terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.50">
        <property type="entry name" value="Glycoside hydrolase family 38, central domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1180">
        <property type="entry name" value="Golgi alpha-mannosidase II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.70.98.30">
        <property type="entry name" value="Golgi alpha-mannosidase II, domain 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011013">
        <property type="entry name" value="Gal_mutarotase_sf_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011330">
        <property type="entry name" value="Glyco_hydro/deAcase_b/a-brl"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011682">
        <property type="entry name" value="Glyco_hydro_38_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015341">
        <property type="entry name" value="Glyco_hydro_38_cen"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037094">
        <property type="entry name" value="Glyco_hydro_38_cen_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000602">
        <property type="entry name" value="Glyco_hydro_38_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027291">
        <property type="entry name" value="Glyco_hydro_38_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028995">
        <property type="entry name" value="Glyco_hydro_57/38_cen_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013780">
        <property type="entry name" value="Glyco_hydro_b"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050843">
        <property type="entry name" value="Glycosyl_Hydrlase_38"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11607">
        <property type="entry name" value="ALPHA-MANNOSIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11607:SF57">
        <property type="entry name" value="ALPHA-MANNOSIDASE 2X"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09261">
        <property type="entry name" value="Alpha-mann_mid"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07748">
        <property type="entry name" value="Glyco_hydro_38C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01074">
        <property type="entry name" value="Glyco_hydro_38N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00872">
        <property type="entry name" value="Alpha-mann_mid"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF88688">
        <property type="entry name" value="Families 57/38 glycoside transferase middle domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF74650">
        <property type="entry name" value="Galactose mutarotase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF88713">
        <property type="entry name" value="Glycoside hydrolase/deacetylase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 8" id="KW-0326">Glycosidase</keyword>
    <keyword evidence="3 8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="10" id="KW-0472">Membrane</keyword>
    <keyword evidence="2 8" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="10" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="4 8" id="KW-0862">Zinc</keyword>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="7"/>
            <end position="26"/>
        </location>
    </feature>
    <feature type="domain" description="Glycoside hydrolase family 38 central" evidence="11">
        <location>
            <begin position="502"/>
            <end position="588"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="885"/>
            <end position="934"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00009792"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023295"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00059516"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00093232"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="RuleBase" id="RU361199"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="SMART" id="SM00872"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007660.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="934" mass="105198" checksum="7E465094D385692D" modified="2019-05-08" version="1">MKLKKQVTVCGAAIFCVAVFSLYLMLDRVQHDPARHQSGGNFPRSQISVLQNRIEQLEQLLEENHEIISHIKESVLELTAHVDGQPVLLPYRTPNGSWVLPPESRPSFFSISPQDCQFALGSGGQKPDLQMLAVASVLPFDNQDGGVWKQGFDITYEPHDWDTEPLQVFVVPHSHNDPGWIKTFEKYYADQTQHILNSMVLKLQEDPRRRFIWSEISFFSKWWDNISTQKRAAVRRLVGGGQLEMVTGGWVMPDEANSHYFAMIDQLIEGHQWLEKNIGVTPRSGWAVDPFGHSATMPYLLRRANLTSMLIQRVHYTIKKHFAATRSLEFMWRQTWDPDSSTDIFCHMMPFYSYDVPHTCGPDPKICCQFDFKRLPGGRINCPWKVPPRAITEANVAERAHLLLDQYRKKSKLFRSKVLLVPLGDDFRYDKPQEWDAQFLNYQRLFDFLNAHPSLHVQAQFGTLSDYFDALSKRTGIIPGLKPPGFPVLSGDFFSYADREDHYWTGYYTSRPFYKSLDRVLEAHLRGAEILYSLALSHARHAGMDSRYPLSDYALLTEARRALGLFQHHDAITGTAKEAVVVDYGVRLLHSLVNLKRIISNAAHYLVLGEKESYHYDAAGPFLGMDDMRLNQDSLPEKTIIKLDTSPRCTPSSGLGQALSLLVNSPRVRVLSEEGQPLAIQLSAQWSSATDMVPDVYQVSIMTRLPALGLSILQLYKSFDSHSTLTSSVHIHLHGRDLPVRPREVFPVRIIPATSDDFCLDNQHMQACFSGLTGLLQSVRRAGEEHRLSSEFLIYGTRSAKDKSGAYLFLPDGDAKPYAPKEPPVVRVTEGPFFSEVASYYQHVQQVVRLYNVPGVEGLSLDVSCLVDIRDHVNKEMALRLSTTIPGAAPPVPEEAASPGQLLPHASHGLHPGQAEPPDPAHSPGPGGVQPTQR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLQ0</accession>
    <name>A0A452HLQ0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Transitional endoplasmic reticulum ATPase</fullName>
            <ecNumber evidence="2">3.6.4.6</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="6">Valosin-containing protein</fullName>
        </alternativeName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="7">
            <text>ATP + H2O = ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:13065"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="3.6.4.6"/>
        </reaction>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the AAA ATPase family.</text>
    </comment>
    <dbReference type="EC" id="3.6.4.6" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452HLQ0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018186.1">
        <property type="protein sequence ID" value="ENSGAGP00000015912.1"/>
        <property type="gene ID" value="ENSGAGG00000011980.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034098">
        <property type="term" value="C:VCP-NPL4-UFD1 AAA ATPase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031593">
        <property type="term" value="F:polyubiquitin modification-dependent protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097352">
        <property type="term" value="P:autophagosome maturation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051228">
        <property type="term" value="P:mitotic spindle disassembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030970">
        <property type="term" value="P:retrograde protein transport, ER to cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd19519">
        <property type="entry name" value="RecA-like_CDC48_r1-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd19528">
        <property type="entry name" value="RecA-like_CDC48_r2-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.8.60:FF:000004">
        <property type="entry name" value="Cell division control 48"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.330.10:FF:000001">
        <property type="entry name" value="Cell division control 48"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.40.20:FF:000003">
        <property type="entry name" value="Transitional endoplasmic reticulum ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000012">
        <property type="entry name" value="Transitional endoplasmic reticulum ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000048">
        <property type="entry name" value="Transitional endoplasmic reticulum ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.8.60">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.40.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.330.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.20.150">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003593">
        <property type="entry name" value="AAA+_ATPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005938">
        <property type="entry name" value="AAA_ATPase_CDC48"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050168">
        <property type="entry name" value="AAA_ATPase_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041569">
        <property type="entry name" value="AAA_lid_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009010">
        <property type="entry name" value="Asp_de-COase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003959">
        <property type="entry name" value="ATPase_AAA_core"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003960">
        <property type="entry name" value="ATPase_AAA_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004201">
        <property type="entry name" value="Cdc48_dom2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029067">
        <property type="entry name" value="CDC48_domain_2-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003338">
        <property type="entry name" value="CDC4_N-term_subdom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01243">
        <property type="entry name" value="CDC48"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23077">
        <property type="entry name" value="AAA-FAMILY ATPASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23077:SF63">
        <property type="entry name" value="TRANSITIONAL ENDOPLASMIC RETICULUM ATPASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00004">
        <property type="entry name" value="AAA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17862">
        <property type="entry name" value="AAA_lid_3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02933">
        <property type="entry name" value="CDC48_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02359">
        <property type="entry name" value="CDC48_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00382">
        <property type="entry name" value="AAA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM01072">
        <property type="entry name" value="CDC48_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01073">
        <property type="entry name" value="CDC48_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50692">
        <property type="entry name" value="ADC-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54585">
        <property type="entry name" value="Cdc48 domain 2-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00674">
        <property type="entry name" value="AAA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="4" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="CDC48 N-terminal subdomain" evidence="10">
        <location>
            <begin position="1"/>
            <end position="63"/>
        </location>
    </feature>
    <feature type="domain" description="CDC48" evidence="9">
        <location>
            <begin position="80"/>
            <end position="146"/>
        </location>
    </feature>
    <feature type="domain" description="AAA+ ATPase" evidence="8">
        <location>
            <begin position="192"/>
            <end position="328"/>
        </location>
    </feature>
    <feature type="domain" description="AAA+ ATPase" evidence="8">
        <location>
            <begin position="465"/>
            <end position="604"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006914"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00012674"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00019970"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00031860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00048883"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="SMART" id="SM00382"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="SMART" id="SM01072"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="SMART" id="SM01073"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015912.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="738" mass="82110" checksum="F9991B566D22E7CC" modified="2019-05-08" version="1">MEELHLFRGDTVLLKGKKRRETVCIVLTDDSCHSEKIRMNRVTRNNLRVRLGDVVSIQACPDVKYGKRIHVLPIDDTIVGLTGNLFDVYLKPYFLEAYRPVHKGDIFLVRGGMRAVEFKVVEVDPSPHCIVAPDTVIHCEGGPIKREDEEESLNDVGYDDIGGCRKQMAQIKEMVELPLRHPGLFKAIGVKPPRGILLYGPPGTGKTLVARAVANETGAFFFLINGPEIMSKLAGESESNLRKAFEEAEKNAPAIIFIDELDAIAPKREKTHGEVERRIVSQLLTLMDGLKQRSHVIVMAATNRPNSVDPALRRFGRFDREIDIGIPDSVGRLEILQIHTKNMKLAEDVDLERVARETHGHVGADLAALCSEAALQAIRKKMNIIDLEDDTIDADILNAMAVTMDDFQWALSQSNPSALRETVVEVPQVCWEDIGGLQEVKRELQELVQFPVEYPDKFLKFGMTPSRGVLFYGPPGCGKTLLAKAIANECQANFVSIKGPELLTMWFGESEANVRDVFDKARQAAPCILFFDELDSIAKARGGGAGDGGGAADRVINQILTEMDGMTNKKTVFIIGATNRPDIIDPAILRPGRLDQLIYIPLPDDTSRAAILSANLRKSPVARDVDLNYLAKITHGFSGADLTEICQRACKLAIREAIETEIKMERERQSHPDAIMDDDYDPVPEIRRDHFEEAMRFARRSVSDNDIRKYEMFAQTLQQSRGFGNFRFPAGKQSGDGG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLQ2</accession>
    <name>A0A452HLQ2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">GOLD domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HLQ2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018127.1">
        <property type="protein sequence ID" value="ENSGAGP00000015865.1"/>
        <property type="gene ID" value="ENSGAGG00000011932.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.680">
        <property type="entry name" value="GOLD domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009038">
        <property type="entry name" value="GOLD_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036598">
        <property type="entry name" value="GOLD_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052269">
        <property type="entry name" value="Golgi-PI4KB_interaction"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22973">
        <property type="entry name" value="LD35087P"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22973:SF3">
        <property type="entry name" value="PROTEIN TMED8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13897">
        <property type="entry name" value="GOLD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF101576">
        <property type="entry name" value="Supernatant protein factor (SPF), C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50866">
        <property type="entry name" value="GOLD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0007">Acetylation</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="GOLD" evidence="3">
        <location>
            <begin position="177"/>
            <end position="334"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1"/>
            <end position="114"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="58"/>
            <end position="71"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="87"/>
            <end position="114"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS50866"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015865.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="336" mass="36939" checksum="8FF099CC9CE6A29C" modified="2019-05-08" version="1">MSEVLVAAAGSCFEPPAAGEPGGRPCSVPEGDGQGRRSPPVPPSETPENEDSTPKPESASQTVASSVSEASQCRPPIGSPVNVHAAEYSRKGDGTSEDQDVARQGEKLADQIQPPREEAVVRMTKYRAPQGPGDIVMIQSEHTGAVDVLSAELETTDLLDEQRKAQPPPLAPPSTWTTCKIREFKAKMGKEKNAQMVVKRGEVVTVRVPTHPNGKRVCWEFATDDYDIGFGVYFDWTPVTSTAITVQVSESSDEEDEEEEDEIEGLTPVGDVERGSKSYLRNRYGEIMPVYRRDSHRDVQAGTHDYPGEGIYLLKFDNSYSLLRNKTLFFHVYYTS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLQ3</accession>
    <name>A0A452HLQ3_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="12">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 8">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the TALE/PBX homeobox family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLQ3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018126.1">
        <property type="protein sequence ID" value="ENSGAGP00000015864.1"/>
        <property type="gene ID" value="ENSGAGG00000011919.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00086">
        <property type="entry name" value="homeodomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.60:FF:000008">
        <property type="entry name" value="Pre-B-cell leukemia transcription factor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.60">
        <property type="entry name" value="Homeodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001356">
        <property type="entry name" value="HD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017970">
        <property type="entry name" value="Homeobox_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009057">
        <property type="entry name" value="Homeodomain-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008422">
        <property type="entry name" value="KN_HD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005542">
        <property type="entry name" value="PBX_PBC_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050224">
        <property type="entry name" value="TALE_homeobox"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11850">
        <property type="entry name" value="HOMEOBOX PROTEIN TRANSCRIPTION FACTORS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05920">
        <property type="entry name" value="Homeobox_KN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03792">
        <property type="entry name" value="PBC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00389">
        <property type="entry name" value="HOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46689">
        <property type="entry name" value="Homeodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00027">
        <property type="entry name" value="HOMEOBOX_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50071">
        <property type="entry name" value="HOMEOBOX_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51978">
        <property type="entry name" value="PBC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 8" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="5 8" id="KW-0371">Homeobox</keyword>
    <keyword evidence="7 8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0804">Transcription</keyword>
    <keyword evidence="3" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="PBC" evidence="11">
        <location>
            <begin position="1"/>
            <end position="126"/>
        </location>
    </feature>
    <feature type="domain" description="Homeobox" evidence="10">
        <location>
            <begin position="125"/>
            <end position="188"/>
        </location>
    </feature>
    <feature type="DNA-binding region" description="Homeobox" evidence="8">
        <location>
            <begin position="127"/>
            <end position="189"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="218"/>
            <end position="243"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="218"/>
            <end position="228"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007601"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023155"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00108"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50071"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS51978"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015864.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="243" mass="27102" checksum="5620A448F5942F91" modified="2019-05-08" version="1">MRLDNMLLAEGVSGPEKGGGSAAAAAAAAASGGSSDNSIEHSDYRAKLTQIRQIYHTELEKYEQACNEFTTHVMNLLREQSRTRPISPKEIERMVGIIHRKFSSIQMQLKQSTCEAVMILRSRFLDARRKRRNFSKQATEILNEYFYSHLSNPYPSEEAKEELAKKCSITVSQVSNWFGNKRIRYKKNIGKFQEEANLYAAKTAVTAAHAVAAAVQNNQTNSPTTPNSGGYPASCYQSDGRIQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLR3</accession>
    <name>A0A452HLR3_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="8">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HLR3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018149.1">
        <property type="protein sequence ID" value="ENSGAGP00000015883.1"/>
        <property type="gene ID" value="ENSGAGG00000011952.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046580">
        <property type="term" value="P:negative regulation of Ras protein signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd04013">
        <property type="entry name" value="C2_SynGAP_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd13375">
        <property type="entry name" value="PH_SynGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05136">
        <property type="entry name" value="RasGAP_DAB2IP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd22265">
        <property type="entry name" value="UDM1_RNF168"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.506.10:FF:000001">
        <property type="entry name" value="Ras GTPase-activating protein nGAP isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000010">
        <property type="entry name" value="Ras GTPase-activating protein nGAP isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.506.10">
        <property type="entry name" value="GTPase Activation - p120gap, domain 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000008">
        <property type="entry name" value="C2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021887">
        <property type="entry name" value="DAB2P_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039360">
        <property type="entry name" value="Ras_GTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023152">
        <property type="entry name" value="RasGAP_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001936">
        <property type="entry name" value="RasGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057606">
        <property type="entry name" value="SynGAP1-like_PH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037779">
        <property type="entry name" value="SynGAP_PH"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10194">
        <property type="entry name" value="RAS GTPASE-ACTIVATING PROTEINS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10194:SF25">
        <property type="entry name" value="RAS_RAP GTPASE-ACTIVATING PROTEIN SYNGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00168">
        <property type="entry name" value="C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12004">
        <property type="entry name" value="DAB2P_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25321">
        <property type="entry name" value="PH_RASGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00616">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00239">
        <property type="entry name" value="C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00233">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00323">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49562">
        <property type="entry name" value="C2 domain (Calcium/lipid-binding domain, CaLB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50004">
        <property type="entry name" value="C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00509">
        <property type="entry name" value="RAS_GTPASE_ACTIV_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50018">
        <property type="entry name" value="RAS_GTPASE_ACTIV_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="1" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="PH" evidence="5">
        <location>
            <begin position="202"/>
            <end position="236"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="6">
        <location>
            <begin position="227"/>
            <end position="345"/>
        </location>
    </feature>
    <feature type="domain" description="Ras-GAP" evidence="7">
        <location>
            <begin position="405"/>
            <end position="597"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="76"/>
            <end position="116"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="879"/>
            <end position="907"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="951"/>
            <end position="1041"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="3">
        <location>
            <begin position="1089"/>
            <end position="1155"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="4">
        <location>
            <begin position="962"/>
            <end position="971"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="977"/>
            <end position="1002"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="1027"/>
            <end position="1039"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015883.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1205" mass="137575" checksum="6ACCD5A85103198B" modified="2019-05-08" version="1">MSYVPFRDVRGAPMHRMPYMHSPYERPAWNPRFCIISGNQLLMLDEDEIHPLLMRERRSEPCRTKLLRRTVSVPVEGRPEHEHHVSRSRRKSVPAGKQYSINMDAPPAPPFRPSQGFLSRRLKSSIKRTKSQPKLDRTSSFRQILPRFRSADHDRARLMQSFKESHSHESLLSPSSAAEALELNLDEDSIIKPVHSSILGQEFCFEVTTASGTKCFACRSAAERDKWIENLQRAVKPNKDNSRRVDNVLKLWIIEARELPPKKRYYCELCLDDMLYARTTSKMRTDNVFWGEHFEFNNLPAVRTIRLHVYKDTDKKRKKDKSNYVGMVSIPIASVTGRHFAEQWYPLSQPSTSKSKAGCPAVRVKSRFQTMSILPMELYKEFAEYVTNNYRMLCAVLEPVLSVKSKEEVACALVHILQSTGKAKDFLSDMAMTEVDRFMDREHLIFRENTLATKAIEEYLKLIGQKYLKDAIGEFIRALYESEENCEVDPMKCSSSTLPDHQANLRMCCELALCKVVNSHCVFPRELKEVFASWRLRCAERGREDIADRLISASLFLRFLCPAVMSPSLFGLMQEYPDEQTSRTLTLIAKVIQNLANFSKFGSKEEYMAFMNEFLELEWASMQQFLYEISNLDTLTNSSSFEGYIDLGRELSTLHALLWEVMPQLSKEALIKLGPLPRLLNDISVALRNPHIQRQPSHQGERLQTKPIILRGPSSDMQPYMMRDLNSSVDLQPYMVRGLNSSMDMTRLPSPTKEKGSKDMFFVTRPPLARSSPAYCTSSSDITEPDQKMLSVNKSVSMLDLQDSRMNSVSNLQSVGDMLNSSQASITAAMGLRPSRLSQGSGSSIAAGLRLSHMGVTTDGAAAQQLRVPLSFQNPLFHMASDGPSAAPSHRPSEGNPDTFAPFHGYSKSEDLSASKHIIHSHSYSDEFTRQGSEFTKRQLSLQENLQNMLSPPQITIGPQPQRAPPQPQPQVPLQRGKSQQLTVSAAQKPRPSSGNLLQSPEPTYGPARARQQSVTKETPSALKPTITKQPSHTPSTLNPVMPASERTVAWVSNMPHLSADIESSHIEREEYKLKEYSKSMDESRLDRVKEYEEEIHSLKERLHMSNRKLEEYERRLMSQEEQTSKILMQYQQRLEQSEKRLRQQQVEKDSQIKSIIGRLMLVEEELRRDHSAAHDVSEPRKRLLDAQLFIRRGSFPPWVQQTRV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLR6</accession>
    <name>A0A452HLR6_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="18">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="9">Endomembrane system</location>
            <topology evidence="9">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="11">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLR6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018145.1">
        <property type="protein sequence ID" value="ENSGAGP00000015879.1"/>
        <property type="gene ID" value="ENSGAGG00000011936.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012505">
        <property type="term" value="C:endomembrane system"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098839">
        <property type="term" value="C:postsynaptic density membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004222">
        <property type="term" value="F:metalloendopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd04269">
        <property type="entry name" value="ZnMc_adamalysin_II_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.390.10:FF:000014">
        <property type="entry name" value="disintegrin and metalloproteinase domain-containing protein 11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.70.10:FF:000001">
        <property type="entry name" value="Disintegrin and metalloproteinase domain-containing protein 22"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.390.10">
        <property type="entry name" value="Collagenase (Catalytic Domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.70.10">
        <property type="entry name" value="Disintegrin domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006586">
        <property type="entry name" value="ADAM_Cys-rich"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018358">
        <property type="entry name" value="Disintegrin_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001762">
        <property type="entry name" value="Disintegrin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036436">
        <property type="entry name" value="Disintegrin_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013111">
        <property type="entry name" value="EGF_extracell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024079">
        <property type="entry name" value="MetalloPept_cat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001590">
        <property type="entry name" value="Peptidase_M12B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002870">
        <property type="entry name" value="Peptidase_M12B_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034027">
        <property type="entry name" value="Reprolysin_adamalysin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11905">
        <property type="entry name" value="ADAM A DISINTEGRIN AND METALLOPROTEASE DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11905:SF14">
        <property type="entry name" value="DISINTEGRIN AND METALLOPROTEINASE DOMAIN-CONTAINING PROTEIN 22"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08516">
        <property type="entry name" value="ADAM_CR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00200">
        <property type="entry name" value="Disintegrin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07974">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01562">
        <property type="entry name" value="Pep_M12B_propep"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01421">
        <property type="entry name" value="Reprolysin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00289">
        <property type="entry name" value="DISINTEGRIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00608">
        <property type="entry name" value="ACR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00050">
        <property type="entry name" value="DISIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57552">
        <property type="entry name" value="Blood coagulation inhibitor (disintegrin)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55486">
        <property type="entry name" value="Metalloproteases ('zincins'), catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50215">
        <property type="entry name" value="ADAM_MEPRO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00427">
        <property type="entry name" value="DISINTEGRIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50214">
        <property type="entry name" value="DISINTEGRIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0165">Cleavage on pair of basic residues</keyword>
    <keyword evidence="7 11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="1 11" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="8" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6 13" id="KW-0472">Membrane</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 14" id="KW-0732">Signal</keyword>
    <keyword evidence="3 13" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 13" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="14">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019291771" evidence="14">
        <location>
            <begin position="24"/>
            <end position="930"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="13">
        <location>
            <begin position="732"/>
            <end position="755"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase M12B" evidence="17">
        <location>
            <begin position="237"/>
            <end position="436"/>
        </location>
    </feature>
    <feature type="domain" description="Disintegrin" evidence="16">
        <location>
            <begin position="442"/>
            <end position="529"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="15">
        <location>
            <begin position="672"/>
            <end position="709"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="59"/>
            <end position="79"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="780"/>
            <end position="849"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="879"/>
            <end position="930"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="788"/>
            <end position="805"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="806"/>
            <end position="824"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="12">
        <location>
            <begin position="837"/>
            <end position="848"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="883"/>
            <end position="903"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="10">
        <location>
            <begin position="501"/>
            <end position="521"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="11">
        <location>
            <begin position="699"/>
            <end position="708"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022685"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00046288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00068"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50214"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50215"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015879.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="930" mass="103928" checksum="E73FD1EB1A26606E" modified="2019-05-08" version="1">MQLAAYLLSLMFCALGTFSLVHSRQEGAALLTEMERRAENRFLERQNIVPLRLIPRPEDGTQTEHGVLNTRVSSSSDRKQSTHVARASFQVDAFGSSFILDVMLNHDLLSSEYLERHIEHGGKTMEVKGGEHCYYQGQIRGNSESFVALSTCHGLHGMFCDGNHTYLIEPEENYTSHEDFHFHSVYKSRLFEFPLDDLPSEFWQMNTTSQKFVVKPRHKRSKRQVHQISRKVEEETKYIELMIVNDHLMCKKHRLSVGHTNSYAKSVVNMADLIYKEQLNTRIVLVAMETWATDNKFTISENPLVTLREFMKYRRDFIREKSDAVHLFSGSQFQSSRSGAAYIGGICSLLKGGGVNEFGKPDLLAVTLAQSLGHNLGIFSDKRKLLTGECKCEDTWSGCIMGDTGYYLPSKFSKCDIEEYHEFLNNGGGACLFNKPSKLLDPPECGNGFVEDGEECDCGTTAECAIEGGGCCHTCTLTAGSQCSNGLCCRKCKFESKGVVCRDAVNDCDIPENCSGNSSQCSPNMHKMDGYSCDNKQGICFGGRCKTRDRQCKYIWGEKVTAADRYCYEKLNIEGTEKGNCGRGKDTWIQCNKQDVLCGYLLCSNISNVPRLGELDGEITSSVIQLGKSYNCSGGHVKLDEETDLGYVEDGTPCGPEMICFERRCLPTDSFNFSICPGTTDDKICSGHGVCSNEIKCVCDRFWVGDDCNTVYKDASIFDEEMAGKGVVSTNIIIGAIAGTILVLALVLAITGWGYKNYRRQRQIPQGDYVKKPGEADSFYSDMPPGVSTNSASSSKKRSNGLSHSWSERIPDTKHISDICENGRPRSNSWQGNIGGNRKKIRGKKYRPRSNSTEYLNPWFKREYNVAKWVEDVNKNTEGPYFRTLSPAKSPSSSTGSIASSRKYPYPMPPLPDEEKKANRPSARLWETSI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLS6</accession>
    <name>A0A452HLS6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Spermatogenesis-defective protein 39 homolog</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="7">VPS33B-interacting protein in apical-basolateral polarity regulator</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="8">VPS33B-interacting protein in polarity and apical restriction</fullName>
        </alternativeName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasmic vesicle</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Early endosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Late endosome</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLS6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018160.1">
        <property type="protein sequence ID" value="ENSGAGP00000015891.1"/>
        <property type="gene ID" value="ENSGAGG00000011955.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005769">
        <property type="term" value="C:early endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005770">
        <property type="term" value="C:late endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006886">
        <property type="term" value="P:intracellular protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007034">
        <property type="term" value="P:vacuolar transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.780">
        <property type="entry name" value="Vps16, C-terminal region"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040057">
        <property type="entry name" value="Spe-39"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038132">
        <property type="entry name" value="Vps16_C_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13364">
        <property type="entry name" value="DEFECTIVE SPERMATOGENESIS PROTEIN 39"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13364:SF6">
        <property type="entry name" value="SPERMATOGENESIS-DEFECTIVE PROTEIN 39 HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="5" id="KW-0967">Endosome</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="100"/>
            <end position="129"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="100"/>
            <end position="110"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="118"/>
            <end position="129"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004412"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004541"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004603"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00019368"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00029984"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00031270"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015891.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="485" mass="55487" checksum="0A9BD1A5EF521FCA" modified="2019-05-08" version="1">MVLCAELAGPPAHPGEEAGTLRAVGLAGGGMNRMKVDEEEYWHSSKCKAFTFDDDDDELSQLKESKRAVNSLREIVDDDDDLEKFSWSGEPVGSISWSIKETASSSSNTPEGRDSGLQKGSSSHPALPKQASSYSISSFFKGRNKLGSFQSLSDALSDTGVKSYAPELRRPKADYKDYSSDWSPKDTVRRMQRGKICSLERFRSLREKLELLDEAVSVHDGNVITAILIFLKRTLKREILFQELEVRQVALRHLIHFLKETGEQKLFLDLLRFLDRTEEVALSQYREHLSIQDAEKRREFLKGCIRLPFSSEDAVHVQDHYTLLERQIIIEANDRHLEAAGQSEIFRKYPRKASILNMPLVTTLFYSCFYHYTEGEGTFSSPTNLKKTFKIPDKQYVLTALAARAKLRAWHDVDALFTTKNWLGYTKKRAPIGFHRVVEILQRNSAPVQVLQEYVRLVEDVETRLNLAMKYKCHDVVVDQIRWKN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLT2</accession>
    <name>A0A452HLT2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">Receptor-type tyrosine-protein phosphatase alpha</fullName>
            <ecNumber evidence="4">3.1.3.48</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="16">Tyrosine protein phosphatase which is involved in integrin-mediated focal adhesion formation. Following integrin engagement, specifically recruits BCAR3, BCAR1 and CRK to focal adhesions thereby promoting SRC-mediated phosphorylation of BRAC1 and the subsequent activation of PAK and small GTPase RAC1 and CDC42.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell junction</location>
            <location evidence="1">Focal adhesion</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the protein-tyrosine phosphatase family. Receptor class 4 subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.48" evidence="4"/>
    <dbReference type="AlphaFoldDB" id="A0A452HLT2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018163.1">
        <property type="protein sequence ID" value="ENSGAGP00000015894.1"/>
        <property type="gene ID" value="ENSGAGG00000010212.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005925">
        <property type="term" value="C:focal adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="CDD" id="cd14621">
        <property type="entry name" value="R-PTPc-A-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd14623">
        <property type="entry name" value="R-PTPc-A-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000007">
        <property type="entry name" value="Receptor-type tyrosine-protein phosphatase alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000021">
        <property type="entry name" value="Receptor-type tyrosine-protein phosphatase alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050348">
        <property type="entry name" value="Protein-Tyr_Phosphatase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000242">
        <property type="entry name" value="PTP_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016130">
        <property type="entry name" value="Tyr_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003595">
        <property type="entry name" value="Tyr_Pase_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016336">
        <property type="entry name" value="Tyr_Pase_rcpt_a/e-type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19134">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19134:SF433">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00102">
        <property type="entry name" value="Y_phosphatase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF002006">
        <property type="entry name" value="PTPR_alpha_epsilon"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00700">
        <property type="entry name" value="PRTYPHPHTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00194">
        <property type="entry name" value="PTPc"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00404">
        <property type="entry name" value="PTPc_motif"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00383">
        <property type="entry name" value="TYR_PHOSPHATASE_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50055">
        <property type="entry name" value="TYR_PHOSPHATASE_PTP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12" id="KW-0965">Cell junction</keyword>
    <keyword evidence="5" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="15" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="14" id="KW-0472">Membrane</keyword>
    <keyword evidence="6 20" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="11" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0677">Repeat</keyword>
    <keyword evidence="8" id="KW-0732">Signal</keyword>
    <keyword evidence="7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="13" id="KW-1133">Transmembrane helix</keyword>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="21">
        <location>
            <begin position="64"/>
            <end position="324"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="22">
        <location>
            <begin position="244"/>
            <end position="315"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="21">
        <location>
            <begin position="356"/>
            <end position="614"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="22">
        <location>
            <begin position="530"/>
            <end position="605"/>
        </location>
    </feature>
    <feature type="active site" description="Phosphocysteine intermediate" evidence="18">
        <location>
            <position position="265"/>
        </location>
    </feature>
    <feature type="active site" description="Phosphocysteine intermediate" evidence="18">
        <location>
            <position position="555"/>
        </location>
    </feature>
    <feature type="binding site" evidence="19">
        <location>
            <position position="233"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="19">
        <location>
            <begin position="265"/>
            <end position="271"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="19">
        <location>
            <position position="309"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="modified residue" description="Phosphotyrosine" evidence="20">
        <location>
            <position position="621"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007765"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00013064"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00054255"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00074216"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="PIRSR" id="PIRSR002006-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="PIRSR" id="PIRSR002006-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="PIRSR" id="PIRSR002006-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015894.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="625" mass="71971" checksum="22C282FD43CBB12B" modified="2019-05-08" version="1">PDLAGARSPWRNEHRCNSLCDIAEPQSMPLLARSPSTNRKYPPLPVDKLEEEINRRMADDNKLFREEFNALPACPIQATCEAASKEENKEKNRYVNILPYDHSRVHLTPIEGVPDSDYINASFINGYQEKNKFIAAQGPKEETVNDFWRMIWEQNTATIVMVTNLKERKECKCAQYWPDQGCWTYGNIRVSVEDVTVLVDYTVRKFCIQQVGDVTNKKPQRLVTQFHFTSWPDFGVPFTPIGMLKFLKKVKTCNPQYAGAIVVHCSAGVGRTGTFIVIDAMLDMMHAERKVDVYGFVSRIRAQRCQMVQTDMQYVFIYQALLEHYLYGDTELEVTSLETHLQKIYNKVPGTSSNGLEEEFKKLTSIKIQNDKMRTGNLPANMKKNRVLQIIPYEFNRVIIPVKRGEENTDYVNASFIDGYRQKDSYIASQGPLQHTIEDFWRMIWEWKSCSIVMLTELEERGQEKCAQYWPSDGSVSYGDITMELKKEEECESYTVRDLLVTNTRENKSRQIRQFHFHGWPEVGIPSDGKGMINIIAAVQKQQQQSGNHPITVHCSAGAGRTGTFCALSTVLERVKAEGILDVFQTVKSLRLQRPHMVQTLEQYEFCYKVVQEYIDAFSDYANFK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="22" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLT4</accession>
    <name>A0A452HLT4_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="1">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="1 2">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="2" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="1" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="1" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HLT4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018222.1">
        <property type="protein sequence ID" value="ENSGAGP00000015947.1"/>
        <property type="gene ID" value="ENSGAGG00000012004.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000313" key="1">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015947.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="222" mass="23249" checksum="EECB8FA6D37C68A3" modified="2019-05-08" version="1">CREQCGDFTGKCRPSTPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPSAPAVLPCVPRPCRPVMHTCFQPCCFSLSLSLAEAEAEPKGWL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLT5</accession>
    <name>A0A452HLT5_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="18">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="9">Endomembrane system</location>
            <topology evidence="9">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="11">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLT5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018164.1">
        <property type="protein sequence ID" value="ENSGAGP00000015895.1"/>
        <property type="gene ID" value="ENSGAGG00000011936.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012505">
        <property type="term" value="C:endomembrane system"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098839">
        <property type="term" value="C:postsynaptic density membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004222">
        <property type="term" value="F:metalloendopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd04269">
        <property type="entry name" value="ZnMc_adamalysin_II_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.390.10:FF:000014">
        <property type="entry name" value="disintegrin and metalloproteinase domain-containing protein 11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.70.10:FF:000001">
        <property type="entry name" value="Disintegrin and metalloproteinase domain-containing protein 22"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.390.10">
        <property type="entry name" value="Collagenase (Catalytic Domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.70.10">
        <property type="entry name" value="Disintegrin domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006586">
        <property type="entry name" value="ADAM_Cys-rich"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018358">
        <property type="entry name" value="Disintegrin_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001762">
        <property type="entry name" value="Disintegrin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036436">
        <property type="entry name" value="Disintegrin_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013111">
        <property type="entry name" value="EGF_extracell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024079">
        <property type="entry name" value="MetalloPept_cat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001590">
        <property type="entry name" value="Peptidase_M12B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002870">
        <property type="entry name" value="Peptidase_M12B_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034027">
        <property type="entry name" value="Reprolysin_adamalysin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11905">
        <property type="entry name" value="ADAM A DISINTEGRIN AND METALLOPROTEASE DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11905:SF14">
        <property type="entry name" value="DISINTEGRIN AND METALLOPROTEINASE DOMAIN-CONTAINING PROTEIN 22"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08516">
        <property type="entry name" value="ADAM_CR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00200">
        <property type="entry name" value="Disintegrin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07974">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01562">
        <property type="entry name" value="Pep_M12B_propep"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01421">
        <property type="entry name" value="Reprolysin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00289">
        <property type="entry name" value="DISINTEGRIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00608">
        <property type="entry name" value="ACR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00050">
        <property type="entry name" value="DISIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57552">
        <property type="entry name" value="Blood coagulation inhibitor (disintegrin)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55486">
        <property type="entry name" value="Metalloproteases ('zincins'), catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50215">
        <property type="entry name" value="ADAM_MEPRO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00427">
        <property type="entry name" value="DISINTEGRIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50214">
        <property type="entry name" value="DISINTEGRIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0165">Cleavage on pair of basic residues</keyword>
    <keyword evidence="7 11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="1 11" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="8" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6 13" id="KW-0472">Membrane</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 14" id="KW-0732">Signal</keyword>
    <keyword evidence="3 13" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 13" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="14">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5018993257" evidence="14">
        <location>
            <begin position="24"/>
            <end position="902"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="13">
        <location>
            <begin position="732"/>
            <end position="755"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase M12B" evidence="17">
        <location>
            <begin position="237"/>
            <end position="436"/>
        </location>
    </feature>
    <feature type="domain" description="Disintegrin" evidence="16">
        <location>
            <begin position="442"/>
            <end position="529"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="15">
        <location>
            <begin position="672"/>
            <end position="709"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="59"/>
            <end position="79"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="782"/>
            <end position="902"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="12">
        <location>
            <begin position="801"/>
            <end position="812"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="834"/>
            <end position="843"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="857"/>
            <end position="875"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="10">
        <location>
            <begin position="501"/>
            <end position="521"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="11">
        <location>
            <begin position="699"/>
            <end position="708"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022685"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00046288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00068"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50214"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50215"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015895.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="902" mass="100396" checksum="752FB864828F41DD" modified="2019-05-08" version="1">MQLAAYLLSLMFCALGTFSLVHSRQEGAALLTEMERRAENRFLERQNIVPLRLIPRPEDGTQTEHGVLNTRVSSSSDRKQSTHVARASFQVDAFGSSFILDVMLNHDLLSSEYLERHIEHGGKTMEVKGGEHCYYQGQIRGNSESFVALSTCHGLHGMFCDGNHTYLIEPEENYTSHEDFHFHSVYKSRLFEFPLDDLPSEFWQMNTTSQKFVVKPRHKRSKRQVHQISRKVEEETKYIELMIVNDHLMCKKHRLSVGHTNSYAKSVVNMADLIYKEQLNTRIVLVAMETWATDNKFTISENPLVTLREFMKYRRDFIREKSDAVHLFSGSQFQSSRSGAAYIGGICSLLKGGGVNEFGKPDLLAVTLAQSLGHNLGIFSDKRKLLTGECKCEDTWSGCIMGDTGYYLPSKFSKCDIEEYHEFLNNGGGACLFNKPSKLLDPPECGNGFVEDGEECDCGTTAECAIEGGGCCHTCTLTAGSQCSNGLCCRKCKFESKGVVCRDAVNDCDIPENCSGNSSQCSPNMHKMDGYSCDNKQGICFGGRCKTRDRQCKYIWGEKVTAADRYCYEKLNIEGTEKGNCGRGKDTWIQCNKQDVLCGYLLCSNISNVPRLGELDGEITSSVIQLGKSYNCSGGHVKLDEETDLGYVEDGTPCGPEMICFERRCLPTDSFNFSICPGTTDDKICSGHGVCSNEIKCVCDRFWVGDDCNTVYKDASIFDEEMAGKGVVSTNIIIGAIAGTILVLALVLAITGWGYKNYRRQRSNGLSHSWSERIPDTKHISDICENGRPRSNSWQGNIGGNRKKIRGKKYRPRSNSTDRVSHVSEPGLSLAHQIPTQGISQGGSHHPHTGNRDHRTLSPAKSPSSSTGSIASSRKYPYPMPPLPDEEKKANRPSARLWETSI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLT7</accession>
    <name>A0A452HLT7_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="18">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="9">Endomembrane system</location>
            <topology evidence="9">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="11">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLT7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018170.1">
        <property type="protein sequence ID" value="ENSGAGP00000015901.1"/>
        <property type="gene ID" value="ENSGAGG00000011936.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012505">
        <property type="term" value="C:endomembrane system"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098839">
        <property type="term" value="C:postsynaptic density membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004222">
        <property type="term" value="F:metalloendopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd04269">
        <property type="entry name" value="ZnMc_adamalysin_II_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.390.10:FF:000014">
        <property type="entry name" value="disintegrin and metalloproteinase domain-containing protein 11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.70.10:FF:000001">
        <property type="entry name" value="Disintegrin and metalloproteinase domain-containing protein 22"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.390.10">
        <property type="entry name" value="Collagenase (Catalytic Domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.70.10">
        <property type="entry name" value="Disintegrin domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006586">
        <property type="entry name" value="ADAM_Cys-rich"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018358">
        <property type="entry name" value="Disintegrin_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001762">
        <property type="entry name" value="Disintegrin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036436">
        <property type="entry name" value="Disintegrin_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013111">
        <property type="entry name" value="EGF_extracell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024079">
        <property type="entry name" value="MetalloPept_cat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001590">
        <property type="entry name" value="Peptidase_M12B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002870">
        <property type="entry name" value="Peptidase_M12B_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034027">
        <property type="entry name" value="Reprolysin_adamalysin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11905">
        <property type="entry name" value="ADAM A DISINTEGRIN AND METALLOPROTEASE DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11905:SF14">
        <property type="entry name" value="DISINTEGRIN AND METALLOPROTEINASE DOMAIN-CONTAINING PROTEIN 22"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08516">
        <property type="entry name" value="ADAM_CR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00200">
        <property type="entry name" value="Disintegrin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07974">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01562">
        <property type="entry name" value="Pep_M12B_propep"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01421">
        <property type="entry name" value="Reprolysin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00289">
        <property type="entry name" value="DISINTEGRIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00608">
        <property type="entry name" value="ACR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00050">
        <property type="entry name" value="DISIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57552">
        <property type="entry name" value="Blood coagulation inhibitor (disintegrin)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55486">
        <property type="entry name" value="Metalloproteases ('zincins'), catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50215">
        <property type="entry name" value="ADAM_MEPRO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00427">
        <property type="entry name" value="DISINTEGRIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50214">
        <property type="entry name" value="DISINTEGRIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0165">Cleavage on pair of basic residues</keyword>
    <keyword evidence="7 11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="1 11" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="8" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6 13" id="KW-0472">Membrane</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 14" id="KW-0732">Signal</keyword>
    <keyword evidence="3 13" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 13" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="14">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019489404" evidence="14">
        <location>
            <begin position="24"/>
            <end position="901"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="13">
        <location>
            <begin position="732"/>
            <end position="755"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase M12B" evidence="17">
        <location>
            <begin position="237"/>
            <end position="436"/>
        </location>
    </feature>
    <feature type="domain" description="Disintegrin" evidence="16">
        <location>
            <begin position="442"/>
            <end position="529"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="15">
        <location>
            <begin position="672"/>
            <end position="709"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="59"/>
            <end position="79"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="780"/>
            <end position="901"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="788"/>
            <end position="805"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="806"/>
            <end position="824"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="12">
        <location>
            <begin position="837"/>
            <end position="848"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="857"/>
            <end position="872"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="10">
        <location>
            <begin position="501"/>
            <end position="521"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="11">
        <location>
            <begin position="699"/>
            <end position="708"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022685"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00046288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00068"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50214"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50215"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015901.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="901" mass="100282" checksum="6732D212DD36F93A" modified="2019-05-08" version="1">MQLAAYLLSLMFCALGTFSLVHSRQEGAALLTEMERRAENRFLERQNIVPLRLIPRPEDGTQTEHGVLNTRVSSSSDRKQSTHVARASFQVDAFGSSFILDVMLNHDLLSSEYLERHIEHGGKTMEVKGGEHCYYQGQIRGNSESFVALSTCHGLHGMFCDGNHTYLIEPEENYTSHEDFHFHSVYKSRLFEFPLDDLPSEFWQMNTTSQKFVVKPRHKRSKRQVHQISRKVEEETKYIELMIVNDHLMCKKHRLSVGHTNSYAKSVVNMADLIYKEQLNTRIVLVAMETWATDNKFTISENPLVTLREFMKYRRDFIREKSDAVHLFSGSQFQSSRSGAAYIGGICSLLKGGGVNEFGKPDLLAVTLAQSLGHNLGIFSDKRKLLTGECKCEDTWSGCIMGDTGYYLPSKFSKCDIEEYHEFLNNGGGACLFNKPSKLLDPPECGNGFVEDGEECDCGTTAECAIEGGGCCHTCTLTAGSQCSNGLCCRKCKFESKGVVCRDAVNDCDIPENCSGNSSQCSPNMHKMDGYSCDNKQGICFGGRCKTRDRQCKYIWGEKVTAADRYCYEKLNIEGTEKGNCGRGKDTWIQCNKQDVLCGYLLCSNISNVPRLGELDGEITSSVIQLGKSYNCSGGHVKLDEETDLGYVEDGTPCGPEMICFERRCLPTDSFNFSICPGTTDDKICSGHGVCSNEIKCVCDRFWVGDDCNTVYKDASIFDEEMAGKGVVSTNIIIGAIAGTILVLALVLAITGWGYKNYRRQRQIPQGDYVKKPGEADSFYSDMPPGVSTNSASSSKKRSNGLSHSWSERIPDTKHISDICENGRPRSNSWQGNIGGNRKKIRGKKYRPRSNSTETLSPAKSPSSSTGSIASSRKYPYPMPPLPDEEKKANRPSARLWETSI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLT9</accession>
    <name>A0A452HLT9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Tetraspanin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 7">Membrane</location>
            <topology evidence="1 7">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 7">Belongs to the tetraspanin (TM4SF) family.</text>
    </comment>
    <comment type="caution">
        <text evidence="7">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLT9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018227.1">
        <property type="protein sequence ID" value="ENSGAGP00000015952.1"/>
        <property type="gene ID" value="ENSGAGG00000012007.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1450.10">
        <property type="entry name" value="Tetraspanin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018499">
        <property type="entry name" value="Tetraspanin/Peripherin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000301">
        <property type="entry name" value="Tetraspanin_animals"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008952">
        <property type="entry name" value="Tetraspanin_EC2_sf"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00335">
        <property type="entry name" value="Tetraspanin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF002419">
        <property type="entry name" value="Tetraspanin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48652">
        <property type="entry name" value="Tetraspanin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5 7" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 7" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="30"/>
            <end position="48"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="60"/>
            <end position="81"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="169"/>
            <end position="195"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="6">
        <location>
            <begin position="125"/>
            <end position="155"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="6">
        <location>
            <begin position="126"/>
            <end position="143"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PIRSR" id="PIRSR002419-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU361218"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015952.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="212" mass="24282" checksum="5AB79C8A179044E2" modified="2019-05-08" version="1">MTVITRYGAHFKIISGVSYEANPYRIFHNVAFYSGICLSMILVLAALLSTAATVRESVCLMAMGFFCFAIVFCALIQAAFWRYTNNSEVEDAMMDVYDSVYEEVRRNLSSFRRQELATIHETFLCCGKQSPFGETGGIENETCPSEHMGTAREDCLQEIQDFLKKHMDFVSALLIITICFMVYGMILTSFLWFSIHFSNNLDRRGKYIFRET</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM20</accession>
    <name>A0A452HM20_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="3">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HM20"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018254.1">
        <property type="protein sequence ID" value="ENSGAGP00000015975.1"/>
        <property type="gene ID" value="ENSGAGG00000012017.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034451">
        <property type="term" value="C:centriolar satellite"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005814">
        <property type="term" value="C:centriole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005876">
        <property type="term" value="C:spindle microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007099">
        <property type="term" value="P:centriole replication"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060271">
        <property type="term" value="P:cilium assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007020">
        <property type="term" value="P:microtubule nucleation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045931">
        <property type="term" value="P:positive regulation of mitotic cell cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042481">
        <property type="entry name" value="CCDC57"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46725">
        <property type="entry name" value="COILED-COIL DOMAIN-CONTAINING PROTEIN 57"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46725:SF1">
        <property type="entry name" value="COILED-COIL DOMAIN-CONTAINING PROTEIN 57"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="606"/>
            <end position="627"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="727"/>
            <end position="762"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="5"/>
            <end position="54"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="83"/>
            <end position="110"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="137"/>
            <end position="171"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="300"/>
            <end position="327"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="378"/>
            <end position="419"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="519"/>
            <end position="546"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="739"/>
            <end position="762"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015975.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="867" mass="100862" checksum="17E8C205B6839A8D" modified="2019-05-08" version="1">MLKMEKELNELLVRKEQEWKELQARQLYFQKTTLQDTRKQLQEMHKKFNRLKEDFTYNLKVLEDRDRELERYDAMFTHLKMAENAKQAEVSDLRIQVDKLQQTLLKETRKQEALQYQYQQKLNEHQLEMERLHSSKVSDIDHHCEEYKKLKQQLERKLQEVEGELALQKQELLMEFDAEIKKREHAFQRQTDEMSNLVLSHELKVKLLTRELEALKEAGMKAAESLQIAETTNLKLEKEAKCKDWELKDLAAVKDAQIQDLECKLHLVYQTWPKEETFQRKHEVLDRFAKEKDPVLASVKEVHAEQIQKWENQIRELQISHETLEVELRRREWRHADCLKEKDTVIEKLQGELTTLKTSWDSHLAQISKETVSKDLQIQSLHEEEMKLRTELSKFQQDVERYKQQLSLAAEREQILERAKVQAELDWQWRCENAERNQYQKSEDLIQSLSTARDQVTAQLQEKEQKLCEMETVLSALTFEKDQAVQALQKHGILPEGEKQMFLGDNKGVLGKDFSATEIQRLKEQNTSMRVAIAEMRKEMEALSDQTPSSFPRKEKIQYTQQTATDAKAATVIFTPDYVQSLEEEIRKLKHKYWIMEEQLEKASKFSKKSSAPFDDPSSTQHHSAPPITQLHTHNFQETSGAICADEVFSGAPLKKLEARTAHLDSMVTQHIQKELQYAQRQRFSRLTSMITFPSSIERLSCGNVPSSGGEGAELPQIQVQMESSVRNCSPGQKKECTSSETNQSQQLNRESTSQSQQAWMSSSGVHSSLQDIWQILDMGSSPSILSPQSNIDQVEFKATHTAKFEESQHTVITGDREKAPVAELTVKGTNTEVQPKLIANKSSQTHPIKSKVSHQMQKIRNYNVKH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM22</accession>
    <name>A0A452HM22_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">PNPLA domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="caution">
        <text evidence="2">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HM22"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018266.1">
        <property type="protein sequence ID" value="ENSGAGP00000015986.1"/>
        <property type="gene ID" value="ENSGAGG00000012001.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005811">
        <property type="term" value="C:lipid droplet"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004806">
        <property type="term" value="F:triacylglycerol lipase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055088">
        <property type="term" value="P:lipid homeostasis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019433">
        <property type="term" value="P:triglyceride catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.1090.10">
        <property type="entry name" value="Cytosolic phospholipase A2 catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016035">
        <property type="entry name" value="Acyl_Trfase/lysoPLipase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033562">
        <property type="entry name" value="PLPL"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002641">
        <property type="entry name" value="PNPLA_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12406">
        <property type="entry name" value="CALCIUM-INDEPENDENT PHOSPHOLIPASE A2 IPLA2 -RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12406:SF23">
        <property type="entry name" value="OMEGA-HYDROXYCERAMIDE TRANSACYLASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01734">
        <property type="entry name" value="Patatin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52151">
        <property type="entry name" value="FabD/lysophospholipase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51635">
        <property type="entry name" value="PNPLA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="2" id="KW-0442">Lipid degradation</keyword>
    <keyword evidence="1 2" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="PNPLA" evidence="3">
        <location>
            <begin position="15"/>
            <end position="155"/>
        </location>
    </feature>
    <feature type="short sequence motif" description="GXSXG" evidence="2">
        <location>
            <begin position="50"/>
            <end position="54"/>
        </location>
    </feature>
    <feature type="short sequence motif" description="DGA/G" evidence="2">
        <location>
            <begin position="142"/>
            <end position="144"/>
        </location>
    </feature>
    <feature type="active site" description="Nucleophile" evidence="2">
        <location>
            <position position="52"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="2">
        <location>
            <position position="142"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01161"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS51635"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015986.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="248" mass="27716" checksum="BDDA41C03D82A79C" modified="2019-05-08" version="1">MAEADRNGTGTPFSLSFSGSGFLALYQIGVVQSLLELAPEILKSASKVYGASAGSLIAAAVDLKESFHEAVRDARKTILGPLSPGCNMLCTIQKRLYKMLPENSHQVASGRLHISLTRVMDGQNVMVSEFSSKEDLIQRYVDGGFTNLQPCSDLEAVITVSPFTGELDICPRDCPAIFYSFQIFNSSIQISVENLCRISYALFPPSYLFFFRGYQDAVLFLRRNSKWYSEPVKRQEAFSVWHCFPYRE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM24</accession>
    <name>A0A452HM24_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Dysferlin</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="17">Dystrophy-associated fer-1-like protein</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="18">Fer-1-like protein 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">Key calcium ion sensor involved in the Ca(2+)-triggered synaptic vesicle-plasma membrane fusion. Plays a role in the sarcolemma repair mechanism of both skeletal muscle and cardiomyocytes that permits rapid resealing of membranes disrupted by mechanical stress.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <location evidence="1">Sarcolemma</location>
            <topology evidence="1">Single-pass type II membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasmic vesicle membrane</location>
            <topology evidence="2">Single-pass type II membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the ferlin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018260.1">
        <property type="protein sequence ID" value="ENSGAGP00000015980.1"/>
        <property type="gene ID" value="ENSGAGG00000011993.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030659">
        <property type="term" value="C:cytoplasmic vesicle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012505">
        <property type="term" value="C:endomembrane system"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015630">
        <property type="term" value="C:microtubule cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030315">
        <property type="term" value="C:T-tubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008289">
        <property type="term" value="F:lipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002281">
        <property type="term" value="P:macrophage activation involved in immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061025">
        <property type="term" value="P:membrane fusion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002280">
        <property type="term" value="P:monocyte activation involved in immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050765">
        <property type="term" value="P:negative regulation of phagocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007009">
        <property type="term" value="P:plasma membrane organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd08373">
        <property type="entry name" value="C2A_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04011">
        <property type="entry name" value="C2B_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04018">
        <property type="entry name" value="C2C_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04017">
        <property type="entry name" value="C2D_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04037">
        <property type="entry name" value="C2E_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd08374">
        <property type="entry name" value="C2F_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000009">
        <property type="entry name" value="dysferlin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000021">
        <property type="entry name" value="dysferlin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000026">
        <property type="entry name" value="dysferlin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000033">
        <property type="entry name" value="dysferlin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000037">
        <property type="entry name" value="dysferlin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000061">
        <property type="entry name" value="dysferlin isoform X8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000008">
        <property type="entry name" value="C2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037726">
        <property type="entry name" value="C2A_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037720">
        <property type="entry name" value="C2B_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037722">
        <property type="entry name" value="C2C_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037723">
        <property type="entry name" value="C2D_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037724">
        <property type="entry name" value="C2E_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037725">
        <property type="entry name" value="C2F_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012968">
        <property type="entry name" value="FerIin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037721">
        <property type="entry name" value="Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012560">
        <property type="entry name" value="Ferlin_A-domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012561">
        <property type="entry name" value="Ferlin_B-domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032362">
        <property type="entry name" value="Ferlin_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055072">
        <property type="entry name" value="Ferlin_DSRM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006614">
        <property type="entry name" value="Peroxin/Ferlin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12546:SF44">
        <property type="entry name" value="DYSFERLIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12546">
        <property type="entry name" value="FER-1-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00168">
        <property type="entry name" value="C2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22901">
        <property type="entry name" value="dsrm_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08165">
        <property type="entry name" value="FerA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08150">
        <property type="entry name" value="FerB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08151">
        <property type="entry name" value="FerI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16165">
        <property type="entry name" value="Ferlin_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00239">
        <property type="entry name" value="C2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SMART" id="SM00694">
        <property type="entry name" value="DysFC"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00693">
        <property type="entry name" value="DysFN"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM01200">
        <property type="entry name" value="FerA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01201">
        <property type="entry name" value="FerB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01202">
        <property type="entry name" value="FerI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49562">
        <property type="entry name" value="C2 domain (Calcium/lipid-binding domain, CaLB)"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50004">
        <property type="entry name" value="C2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0106">Calcium</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="14" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="12" id="KW-0446">Lipid-binding</keyword>
    <keyword evidence="13 20" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="10" id="KW-0735">Signal-anchor</keyword>
    <keyword evidence="6 20" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 20" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="2037"/>
            <end position="2059"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="1"/>
            <end position="101"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="197"/>
            <end position="315"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="354"/>
            <end position="490"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="1130"/>
            <end position="1256"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="1303"/>
            <end position="1431"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="1553"/>
            <end position="1671"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="1787"/>
            <end position="1935"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="127"/>
            <end position="208"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1988"/>
            <end position="2009"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="19">
        <location>
            <begin position="127"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="172"/>
            <end position="182"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="19">
        <location>
            <begin position="188"/>
            <end position="203"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="1990"/>
            <end position="2009"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004274"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007561"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023121"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00060070"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00068801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00078385"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00079079"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015980.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2072" mass="236784" checksum="BD1D11C381DED7FB" modified="2019-05-08" version="1">MLRVFILHAENVQTPDTDISDSYCSVVFEGVKKRTKVIKNNVNPVWNEGFDWDLKGIPLDQNAELTVVVKDHETVGRNRFLGEARVPLQDVLVNPTLTASFNAPLLDTKRQSTGAFLNLQVSYIPPPGAAPLFPPPTPPEPTPSTAELDTVTDTAGEEETEDQIVTGDETEPSTGPSGSEQPTLPRKPPSHHVHGIKRKKSAPKKLLSNKPQDFQIRVRVIEGRQLPGVNIKSVVKVTVAGQTKRTRIRKGNGPFFDEIFFFNVFESPAELFDEPIFITVVDSRSLRTDSVIGEFQLDVGTVYAEPKHALLRKWLLLSDSEDFSAGAKGYVKVSLFVLGPGDEAPLEKKEVAEEKEDIEGNLLRPTGVTLRGAHFCLKIYRAEDLPQMDDAIMDNVKQIFGFESNKKNLVDPFVEVSFAGRTISSKILEKNANPQWNQRITVPAMFPSMCEKMRIRVMDWDRVTHNDVIGTTHLRLSQISAPGGELEDEFPVLAKPLKAAAMDDGLGFLPTFGPCYVNLYGSPREFTGFPDPYEELNMGKGEGVAFRGRLLMELETKLVEHIEQKLEDIPPDDILRVEKHLRRRKYCLFAAFYSASMLQDVDAAIQFEVSIGNYGNKFDNTCLPLASTTQYSGAVFDGCQYYYLPWGNVKPVVVLSSYWEDISHRTDMQNLLQRTADRLEAGLELVHLALKARCSSNELDSLVSQLIDDIIADCSQPLVDVSNKTASTHLDQFVYKFRSTNLHQIIQAALNLKHEDSDLQTVLDQAEDWLLRLKAMAEEPQNSLPDIVIWMLQGDKRVAYARVPAHEVLFSRTGANCCGKNCGKLQTIFLKYPQEEVMGPRIPAQVRVQLWFGLAVDEKEFNKYAEGKLSVFAETYENQVKLALVGNWGTTGLTYPKYSDVTGKIKLPKDSFRPSVGWTWAGDWFISPEKTLLYDTDAGHMSFVEEVFENQVRLPGGQWIHMADAYTDVNGEKVLPKEEIECPPGWKWEDVEWDTDLNRAVDERGWEYGIAIPPDHKPKAWVPAEKMYHTSRRRRWVRLRRRDLKQMEALKKHKQEELEGEGWEYASLFGWKFHLNYRKTDTFRRRRWRRRMEPLERTGAAAVFALEGALGGVTDDKSDDGKSVSTLSFGVNRPTVSCIFDYGSRYHLRCYIYQARDLIAMDKDSFSDPYAIVSFLHQSQKTVVVKNTLNPLWDQTLIFYEIEIFGDPQNVADVPPNIVVEIYDHDTYGADEFMGRCICKPSLERSPRLSWHPVARGNRKAGELLAAFELIQREKPAVHHLPGFESELSASLDESADSDLPYPPPQREPNIYMVPQGIKPVLQRTAIEILAWGLRNLKSYQLASITSPSLQVECGGQTVQSCVIKNVRKNPNFDVCVLFMEVRLPKEALYCPPVIMKVIDNRQFGRKPVVGQCTIQSLEEFYCDPYKEETDAPEEQPDEVSLTPRDDVLIDIDDKEPLIPVQEEEFIDWWSKFYASTGEREKCGSYLEKGFDTLKVYETELENMEEFERLSDFCHTFKLYRGRTQDINEDPSVVGEFKGSFKMYPLPDDPRVPVPPRQFHQLPAKGPQECLIRIYIIRAFDLQPKDSNGKCDPYVKISVGKKSINDQENYIPCTLEPVFGKMFELTCTLPLEKDLKVTLYDYDLLSKDEKIGETVIDLENRFLSKFGARCGLPQTYCISGPNQWRDQLRPSQLLHLFSLQRNCKAPIYKPDRVVFREQEYFLSEFEDGKPPNPHLGAPEERLALYTLRKQGLVPEHVETRPLYSPLQPEIEQGKLQMWVDLFPKSLGLPGPPFNITPRRAKRFFLRCIIWNTKDVILDDLSITGEKMSDIYVKGWMTGHEENKQKTDVHYRSMGGEGNFNWRFVFPFDYLAAEQVCHIAKKEHFWSLDKTDNKVPPQLILQIWDNDKFSFDDYLGSVQMNLNRMPAPTKTAEKCSLNLLDETLSSDRFVSLFEQKAVKGWWPCVVEAEEKKTIAGKLEMTLEIVTEQEQEERPAGMGRDEPNMNPKLEDPKRPETSFLWFTSPYKTMKYILWRRFKWVFLIAIVVFILLLFIGIFIYAFPNYAAMKLVKPFS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM27</accession>
    <name>A0A452HM27_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Dysferlin</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="17">Dystrophy-associated fer-1-like protein</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="18">Fer-1-like protein 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">Key calcium ion sensor involved in the Ca(2+)-triggered synaptic vesicle-plasma membrane fusion. Plays a role in the sarcolemma repair mechanism of both skeletal muscle and cardiomyocytes that permits rapid resealing of membranes disrupted by mechanical stress.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <location evidence="1">Sarcolemma</location>
            <topology evidence="1">Single-pass type II membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasmic vesicle membrane</location>
            <topology evidence="2">Single-pass type II membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the ferlin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018271.1">
        <property type="protein sequence ID" value="ENSGAGP00000015991.1"/>
        <property type="gene ID" value="ENSGAGG00000011993.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030659">
        <property type="term" value="C:cytoplasmic vesicle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012505">
        <property type="term" value="C:endomembrane system"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015630">
        <property type="term" value="C:microtubule cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030315">
        <property type="term" value="C:T-tubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008289">
        <property type="term" value="F:lipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002281">
        <property type="term" value="P:macrophage activation involved in immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061025">
        <property type="term" value="P:membrane fusion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002280">
        <property type="term" value="P:monocyte activation involved in immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050765">
        <property type="term" value="P:negative regulation of phagocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007009">
        <property type="term" value="P:plasma membrane organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd08373">
        <property type="entry name" value="C2A_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04011">
        <property type="entry name" value="C2B_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04018">
        <property type="entry name" value="C2C_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04017">
        <property type="entry name" value="C2D_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04037">
        <property type="entry name" value="C2E_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd08374">
        <property type="entry name" value="C2F_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000009">
        <property type="entry name" value="dysferlin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000021">
        <property type="entry name" value="dysferlin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000026">
        <property type="entry name" value="dysferlin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000033">
        <property type="entry name" value="dysferlin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000037">
        <property type="entry name" value="dysferlin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000061">
        <property type="entry name" value="dysferlin isoform X8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000008">
        <property type="entry name" value="C2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037726">
        <property type="entry name" value="C2A_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037720">
        <property type="entry name" value="C2B_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037722">
        <property type="entry name" value="C2C_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037723">
        <property type="entry name" value="C2D_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037724">
        <property type="entry name" value="C2E_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037725">
        <property type="entry name" value="C2F_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012968">
        <property type="entry name" value="FerIin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037721">
        <property type="entry name" value="Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012560">
        <property type="entry name" value="Ferlin_A-domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012561">
        <property type="entry name" value="Ferlin_B-domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032362">
        <property type="entry name" value="Ferlin_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055072">
        <property type="entry name" value="Ferlin_DSRM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006614">
        <property type="entry name" value="Peroxin/Ferlin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12546:SF44">
        <property type="entry name" value="DYSFERLIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12546">
        <property type="entry name" value="FER-1-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00168">
        <property type="entry name" value="C2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22901">
        <property type="entry name" value="dsrm_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08165">
        <property type="entry name" value="FerA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08150">
        <property type="entry name" value="FerB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08151">
        <property type="entry name" value="FerI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16165">
        <property type="entry name" value="Ferlin_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00239">
        <property type="entry name" value="C2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SMART" id="SM00694">
        <property type="entry name" value="DysFC"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00693">
        <property type="entry name" value="DysFN"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM01200">
        <property type="entry name" value="FerA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01201">
        <property type="entry name" value="FerB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01202">
        <property type="entry name" value="FerI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49562">
        <property type="entry name" value="C2 domain (Calcium/lipid-binding domain, CaLB)"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50004">
        <property type="entry name" value="C2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0106">Calcium</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="14" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="12" id="KW-0446">Lipid-binding</keyword>
    <keyword evidence="13 20" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="10" id="KW-0735">Signal-anchor</keyword>
    <keyword evidence="6 20" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 20" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="2034"/>
            <end position="2056"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="1"/>
            <end position="101"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="197"/>
            <end position="315"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="354"/>
            <end position="489"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="1116"/>
            <end position="1242"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="1279"/>
            <end position="1407"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="1550"/>
            <end position="1668"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="1784"/>
            <end position="1932"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="127"/>
            <end position="208"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1985"/>
            <end position="2006"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="19">
        <location>
            <begin position="127"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="172"/>
            <end position="182"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="19">
        <location>
            <begin position="188"/>
            <end position="203"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="1987"/>
            <end position="2006"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004274"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007561"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023121"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00060070"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00068801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00078385"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00079079"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015991.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2069" mass="236413" checksum="1F2FF73C53A660EE" modified="2019-05-08" version="1">MLRVFILHAENVQTPDTDISDSYCSVVFEGVKKRTKVIKNNVNPVWNEGFDWDLKGIPLDQNAELTVVVKDHETVGRNRFLGEARVPLQDVLVNPTLTASFNAPLLDTKRQSTGAFLNLQVSYIPPPGAAPLFPPPTPPEPTPSTAELDTVTDTAGEEETEDQIVTGDETEPSTGPSGSEQPTLPRKPPSHHVHGIKRKKSAPKKLLSNKPQDFQIRVRVIEGRQLPGVNIKSVVKVTVAGQTKRTRIRKGNGPFFDEIFFFNVFESPAELFDEPIFITVVDSRSLRTDSVIGEFQLDVGTVYAEPKHALLRKWLLLSDSEDFSAGAKGYVKVSLFVLGPGDEAPLEKKEVAEEKEDIEGNLLRPTGVTLRGAHFCLKIYRAEDLPQMDDAIMDNVKQIFGFESNKKNLVDPFVEVSFAGRTISSKILEKNANPQWNQRITVPAMFPSMCEKMRIRVMDWDRVTHNDVIGTTHLRLSQISAPGGELEVDDGLGFLPTFGPCYVNLYGSPREFTGFPDPYEELNMGKGEGVAFRGRLLMELETKLVEHIEQKLEDIPPDDILRVEKHLRRRKYCLFAAFYSASMLQDVDAAIQFEVSIGNYGNKFDNTCLPLASTTQYSGAVFDGCQYYYLPWGNVKPVVVLSSYWEDISHRTDMQNLLQRTADRLEAGLELVHLALKARCSSNELDSLVSQLIDDIIADCSQPLVDVSNKTASTHLDQFVYKFRSTNLHQIIQAALNLKHEDSDLQTVLDQAEDWLLRLKAMAEEPQNSLPDIVIWMLQGDKRVAYARVPAHEVLFSRTGANCCGKNCGKLQTIFLKYPQEEVMGPRIPAQVRVQLWFGLAVDEKEFNKYAEGKLSVFAETYENQVKLALVGNWGTTGLTYPKYSDVTGKIKLPKDSFRPSVGWTWAGDWFISPEKTLLYDTDAGHMSFVEEVFENQVRLPGGQWIHMADAYTDVNGEKVLPKEEIECPPGWKWEDVEWDTDLNRAVDERGWEYGIAIPPDHKPKAWVPAEKMYHTSRRRRWVRLRRRDLKQMEALKKHKQEELEGEGWEYASLFGWKFHLNYRKTDTFRRRRWRRRMEPLERTGAAAVFALEGALGGVTDDKSDDGKSVSTLSFGVNRPTVSCIFDYGSRYHLRCYIYQARDLIAMDKDSFSDPYAIVSFLHQSQKTVVVKNTLNPLWDQTLIFYEIEIFGDPQNVADVPPNIVVEIYDHDTYGADEFMGRCICKPSLERSPRLSWHPVARGNRKAGELLAAFELIQREKSELSASLDESADSDLPYPPPQREPNIYMVPQGIKPVLQRTAIEILAWGLRNLKSYQLASITSPSLQVECGGQTVQSCVIKNVRKNPNFDVCVLFMEVRLPKEALYCPPVIMKVIDNRQFGRKPVVGQCTIQSLEEFYCDPYKEETDAPEEQPDEVSLTPRDDVLIDIDDKEPLIPVQLAEGMTSSTLINMPASRADLHEEEFIDWWSKFYASTGEREKCGSYLEKGFDTLKVYETELENMEEFERLSDFCHTFKLYRGRTQDINEDPSVVGEFKGSFKMYPLPDDPRVPVPPRQFHQLPAKGPQECLIRIYIIRAFDLQPKDSNGKCDPYVKISVGKKSINDQENYIPCTLEPVFGKMFELTCTLPLEKDLKVTLYDYDLLSKDEKIGETVIDLENRFLSKFGARCGLPQTYCISGPNQWRDQLRPSQLLHLFSLQRNCKAPIYKPDRVVFREQEYFLSEFEDGKPPNPHLGAPEERLALYTLRKQGLVPEHVETRPLYSPLQPEIEQGKLQMWVDLFPKSLGLPGPPFNITPRRAKRFFLRCIIWNTKDVILDDLSITGEKMSDIYVKGWMTGHEENKQKTDVHYRSMGGEGNFNWRFVFPFDYLAAEQVCHIAKKEHFWSLDKTDNKVPPQLILQIWDNDKFSFDDYLGSVQMNLNRMPAPTKTAEKCSLNLLDETLSSDRFVSLFEQKAVKGWWPCVVEAEEKKTIAGKLEMTLEIVTEQEQEERPAGMGRDEPNMNPKLEDPKRPETSFLWFTSPYKTMKYILWRRFKWVFLIAIVVFILLLFIGIFIYAFPNYAAMKLVKPFS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM30</accession>
    <name>A0A452HM30_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="16">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HM30"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018265.1">
        <property type="protein sequence ID" value="ENSGAGP00000015985.1"/>
        <property type="gene ID" value="ENSGAGG00000012025.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001228">
        <property type="term" value="F:DNA-binding transcription activator activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd07765">
        <property type="entry name" value="KRAB_A-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001833">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002434">
        <property type="entry name" value="Zinc finger protein 1168"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000638">
        <property type="entry name" value="Zinc finger protein 184"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001823">
        <property type="entry name" value="Zinc finger protein 322"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002343">
        <property type="entry name" value="Zinc finger protein 33A"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002090">
        <property type="entry name" value="Zinc finger protein 473"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000737">
        <property type="entry name" value="Zinc finger protein 565"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000361">
        <property type="entry name" value="Zinc finger protein 658"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001909">
        <property type="entry name" value="KRAB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036051">
        <property type="entry name" value="KRAB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24393">
        <property type="entry name" value="ZINC FINGER PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24393:SF100">
        <property type="entry name" value="ZINC FINGER PROTEIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01352">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="SMART" id="SM00349">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109640">
        <property type="entry name" value="KRAB domain (Kruppel-associated box)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50805">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="11"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="13" id="KW-0732">Signal</keyword>
    <keyword evidence="9" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 11" id="KW-0863">Zinc-finger</keyword>
    <feature type="signal peptide" evidence="13">
        <location>
            <begin position="1"/>
            <end position="16"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019548014" evidence="13">
        <location>
            <begin position="17"/>
            <end position="462"/>
        </location>
    </feature>
    <feature type="domain" description="KRAB" evidence="15">
        <location>
            <begin position="14"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="14">
        <location>
            <begin position="142"/>
            <end position="169"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="14">
        <location>
            <begin position="170"/>
            <end position="197"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="14">
        <location>
            <begin position="198"/>
            <end position="225"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="14">
        <location>
            <begin position="226"/>
            <end position="253"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="14">
        <location>
            <begin position="254"/>
            <end position="281"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="14">
        <location>
            <begin position="282"/>
            <end position="309"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="14">
        <location>
            <begin position="310"/>
            <end position="337"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="14">
        <location>
            <begin position="338"/>
            <end position="365"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="14">
        <location>
            <begin position="366"/>
            <end position="393"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="14">
        <location>
            <begin position="394"/>
            <end position="421"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="14">
        <location>
            <begin position="422"/>
            <end position="449"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="92"/>
            <end position="126"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="101"/>
            <end position="119"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015985.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="462" mass="52889" checksum="CE7F194397132737" modified="2019-05-08" version="1">LVLSVGLLLFQGPVTFEEVAVYFTSEEWALLDPTQRALYWDVMQENYENVTSLGKGYCPLSSWKGKWRGKVYASPTMPLLLCPQVAKHRELSQRSKRNVSRSHEQGKSCEIQHRPEREQGNQPGEKMVLSKHQRIHTGKRPYKCSECGKSFITSSVLSQHQRIHTGERPYECLECGKSFITSSDLSQHQRIHTGERPYKCSECGKSFITSSDLSKHQRIHTGKRPYKCSECGKSFITSSVLSQHQRIHTGERPYECRECGKSFSSSSALSHHQRIHTGERPFECSDCGKTFAFRSVLSKHQRIHTGERPYKCSECGKSFTQSSQLTEHQRIHTGERPFECSDCGKTFTYRSVLSKHQRIHTGERPYKCHECGKSFTRSSHLTGHQRIHTGERPYKCHECGESFTRSSHLSGHQRIHTGERPYKCGKCGKTFIHRQELISHQRIHTGMRPNGSSECGKTLSCH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM34</accession>
    <name>A0A452HM34_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="15">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HM34"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018270.1">
        <property type="protein sequence ID" value="ENSGAGP00000015990.1"/>
        <property type="gene ID" value="ENSGAGG00000012025.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000785">
        <property type="term" value="C:chromatin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031519">
        <property type="term" value="C:PcG protein complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005667">
        <property type="term" value="C:transcription regulator complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd07765">
        <property type="entry name" value="KRAB_A-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001833">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002434">
        <property type="entry name" value="Zinc finger protein 1168"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000638">
        <property type="entry name" value="Zinc finger protein 184"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001823">
        <property type="entry name" value="Zinc finger protein 322"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002343">
        <property type="entry name" value="Zinc finger protein 33A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001498">
        <property type="entry name" value="Zinc finger protein 404"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002090">
        <property type="entry name" value="Zinc finger protein 473"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000737">
        <property type="entry name" value="Zinc finger protein 565"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002134">
        <property type="entry name" value="Zinc finger protein 616"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001909">
        <property type="entry name" value="KRAB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036051">
        <property type="entry name" value="KRAB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14003">
        <property type="entry name" value="TRANSCRIPTIONAL REPRESSOR PROTEIN YY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14003:SF23">
        <property type="entry name" value="ZINC FINGER PROTEIN 143"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01352">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="SMART" id="SM00349">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109640">
        <property type="entry name" value="KRAB domain (Kruppel-associated box)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50805">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="14"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="12" id="KW-0732">Signal</keyword>
    <keyword evidence="9" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 11" id="KW-0863">Zinc-finger</keyword>
    <feature type="signal peptide" evidence="12">
        <location>
            <begin position="1"/>
            <end position="19"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019227028" evidence="12">
        <location>
            <begin position="20"/>
            <end position="484"/>
        </location>
    </feature>
    <feature type="domain" description="KRAB" evidence="14">
        <location>
            <begin position="17"/>
            <end position="100"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="78"/>
            <end position="100"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="101"/>
            <end position="128"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="129"/>
            <end position="156"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="157"/>
            <end position="184"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="192"/>
            <end position="219"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="220"/>
            <end position="247"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="248"/>
            <end position="275"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="276"/>
            <end position="303"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="304"/>
            <end position="331"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="332"/>
            <end position="359"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="360"/>
            <end position="387"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="388"/>
            <end position="415"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="416"/>
            <end position="443"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="444"/>
            <end position="471"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015990.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="484" mass="55326" checksum="368F1673C9AD7B50" modified="2019-05-08" version="1">MGQQVLSVGLLLFQGPVTFEEVAVYFTSEEWALLDPTQRALYWDVMQENYENVTSLGKGYCPLSSWKGKWRVPRKQGCGKTFSCSSSLSVHQRIHTGERPYGCCQCGKTFTFRSNLVHHMRIHTGKRPYKCSECGKSFITSSDLSKHQRIHTGERPFECSECGKTFAFRSVLSKHQRIHTGKRPYKCILSPYECLECGKSFITSSDLSQHQRIHTGERPYKCSECGKSFITSSDLSKHQRIHTGKRPYKCSECGKSFITSSVLSQHQRIHTGERPYECRECGKSFSSSSALSHHQRIHTGERPFECSDCGKTFAFRSVLSKHQRIHTGERPYKCSECGKSFTQSSQLTEHQRIHTGERPFECSDCGKTFTYRSVLSKHQRIHTGERPYKCHECGKSFTRSSHLTGHQRIHTGERPYKCHECGESFTRSSHLSGHQRIHTGERPYKCGKCGKTFIHRQELISHQRIHTGMRPNGSSECGKTLSCH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM42</accession>
    <name>A0A452HM42_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Sema domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the semaphorin family.</text>
    </comment>
    <comment type="caution">
        <text evidence="8">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HM42"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018289.1">
        <property type="protein sequence ID" value="ENSGAGP00000016009.1"/>
        <property type="gene ID" value="ENSGAGG00000011815.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045499">
        <property type="term" value="F:chemorepellent activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0038191">
        <property type="term" value="F:neuropilin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030215">
        <property type="term" value="F:semaphorin receptor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007411">
        <property type="term" value="P:axon guidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001755">
        <property type="term" value="P:neural crest cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030335">
        <property type="term" value="P:positive regulation of cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071526">
        <property type="term" value="P:semaphorin-plexin signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05871">
        <property type="entry name" value="Ig_Sema3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd11253">
        <property type="entry name" value="Sema_3E"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:000015">
        <property type="entry name" value="Semaphorin 3B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000030">
        <property type="entry name" value="Semaphorin 3F like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1680.10:FF:000001">
        <property type="entry name" value="Semaphorin 3F like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1680.10">
        <property type="entry name" value="ligand-binding face of the semaphorins, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013151">
        <property type="entry name" value="Immunoglobulin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016201">
        <property type="entry name" value="PSI"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001627">
        <property type="entry name" value="Semap_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036352">
        <property type="entry name" value="Semap_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027231">
        <property type="entry name" value="Semaphorin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015513">
        <property type="entry name" value="Semaphorin_3E_Sema"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11036">
        <property type="entry name" value="SEMAPHORIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11036:SF22">
        <property type="entry name" value="SEMAPHORIN-3E"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00047">
        <property type="entry name" value="ig"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01403">
        <property type="entry name" value="Sema"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00423">
        <property type="entry name" value="PSI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00630">
        <property type="entry name" value="Sema"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103575">
        <property type="entry name" value="Plexin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF101912">
        <property type="entry name" value="Sema domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51004">
        <property type="entry name" value="SEMA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0964">Secreted</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <feature type="domain" description="Sema" evidence="9">
        <location>
            <begin position="26"/>
            <end position="510"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009492"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00352"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016009.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="769" mass="88949" checksum="CFF422029DDBFE7A" modified="2019-05-08" version="1">MEFRWHFGRCLSTPMVLHLAPARRKRLGITALELWDLNRTSVFHSPSGYLGLHTMLLDEYQERLFVGGRDLVYSLSLDRVSENYREIHWPSTPQQAEECTIKGRDADECANYIRVLHRYNRTHLLACGTGAFDPLCTFIRVGHPSEDHLFQLDWHKFERGRGRCPFDPSSSFISTLIGGELFAGLYNDYWGRDAAVLRTMGRMPHIRTEPDDERLLKEPKFVGSYMIPDNEDRDDNKVYFFFTEKALEAETGAHAIYARVGRVCANDMGGQRMLVNKWSTFLKTRLICSVPGMNGINTHFDELEDVFLLQTRDNKNPVVFGLFNTTSNIFRGYAICVYHIASIRAAFNGPYAHKEGPEYHWSLYEGKVPYPRPGSCASKINGGLYGTTKDYPDEAVRFARSHPLMYQPIKPVHKRPILVKTDGKYNLKQIAVDRVEAEDGQYDVLFIGTDNGIVLKVITIYNQDTESMEEVILEEMQVFKVPVPIISMEISSKRQQLYVGSESAVAQVKFHQCDMYGTACADCCLARDPYCAWDGISCSRYYPTGMQAKRRFRRQDVRHGNAAQQCFGQQFIGEALEKTEERIVYGIEYNSTLLECTPRTLQAKVIWFVQRAHEARKEEVKTDDRIVKMDLGLLFLRLHRLDAGTYFCQTVEHSIVHTVRKITLEVVEEGRVDEMFNRDYEEDIPHKMPCPAQSSIPQGSKPWYKEFLQLIGYSNFQRVEEYCEKVWCTDKRRKKLKMSPSKWKYANPQEKKARIRPEHYRLPRNIVDS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM54</accession>
    <name>A0A452HM54_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Putative malate dehydrogenase 1B</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the LDH/MDH superfamily. MDH type 2 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HM54"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018351.1">
        <property type="protein sequence ID" value="ENSGAGP00000016067.1"/>
        <property type="gene ID" value="ENSGAGG00000012069.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016615">
        <property type="term" value="F:malate dehydrogenase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016616">
        <property type="term" value="F:oxidoreductase activity, acting on the CH-OH group of donors, NAD or NADP as acceptor"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006108">
        <property type="term" value="P:malate metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006099">
        <property type="term" value="P:tricarboxylic acid cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.720:FF:000144">
        <property type="entry name" value="Malate dehydrogenase [NADP]"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.110.10:FF:000006">
        <property type="entry name" value="putative malate dehydrogenase 1B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.110.10">
        <property type="entry name" value="Lactate dehydrogenase/glycoside hydrolase, family 4, C-terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.720">
        <property type="entry name" value="NAD(P)-binding Rossmann-like Domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022383">
        <property type="entry name" value="Lactate/malate_DH_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015955">
        <property type="entry name" value="Lactate_DH/Glyco_Ohase_4_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010945">
        <property type="entry name" value="Malate_DH_type2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036291">
        <property type="entry name" value="NAD(P)-bd_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23382">
        <property type="entry name" value="MALATE DEHYDROGENASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02866">
        <property type="entry name" value="Ldh_1_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56327">
        <property type="entry name" value="LDH C-terminal domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51735">
        <property type="entry name" value="NAD(P)-binding Rossmann-fold domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0520">NAD</keyword>
    <keyword evidence="3" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0816">Tricarboxylic acid cycle</keyword>
    <feature type="domain" description="Lactate/malate dehydrogenase C-terminal" evidence="7">
        <location>
            <begin position="279"/>
            <end position="359"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="409"/>
            <end position="470"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="450"/>
            <end position="470"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00009613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022532"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023027"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00039310"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF02866"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016067.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="470" mass="52430" checksum="54C6A329C1F565CD" modified="2019-05-08" version="1">MLSEQMLRIAKENLQTHIEIEKEEEYIKSLINPLQVWISSASAPACYNLIPLLANGEVFGMATEVSIHLLDSSHYKEILHGIVMEAEDLAFPLLRSVSMHTELDDAFVQADIVILLDDILFQHEIPSLEDCIRQVTEQCKVYGSLIEKNANSKVKVIVSGKTFVNLRALMIMTFAPSIDGQNIIAVAMFLENAAKAMLARKLNMNSAGVKDVIVWGNISGNSYVDLSKAKVYKYDSAIWGPPNFSRHLLDMIYDGSWVRSEFAFVLSSLSSWGHHCLGMSPAHVIATVLRYWYQDSPPGEIVSMGILSEGQFCIPEGIIFSMPVRFQDGSWEVITETEISEETQETLKRLAHDLIQEKQVALGEIPEMRPYKEPAFNGFNLLALLEEENVQTPPEDLDTLSNGLLQPETQEGEKVQATTDEPEAPPDGLLQSEMLEEDKVQVPIDDPETSSESLNQSETSLSNAENNENP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM55</accession>
    <name>A0A452HM55_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Protein transport protein SEC23</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="12">Component of the coat protein complex II (COPII) which promotes the formation of transport vesicles from the endoplasmic reticulum (ER). The coat has two main functions, the physical deformation of the endoplasmic reticulum membrane into vesicles and the selection of cargo molecules for their transport to the Golgi complex.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="13">Cytoplasmic vesicle</location>
            <location evidence="13">COPII-coated vesicle membrane</location>
            <topology evidence="13">Peripheral membrane protein</topology>
            <orientation evidence="13">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1 13">Endoplasmic reticulum membrane</location>
            <topology evidence="1 13">Peripheral membrane protein</topology>
            <orientation evidence="1 13">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="13">Cytoplasm</location>
            <location evidence="13">Cytosol</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 13">Belongs to the SEC23/SEC24 family. SEC23 subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HM55"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018304.1">
        <property type="protein sequence ID" value="ENSGAGP00000016021.1"/>
        <property type="gene ID" value="ENSGAGG00000006201.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030127">
        <property type="term" value="C:COPII vesicle coat"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070971">
        <property type="term" value="C:endoplasmic reticulum exit site"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090110">
        <property type="term" value="P:COPII-coated vesicle cargo loading"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006886">
        <property type="term" value="P:intracellular protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd01478">
        <property type="entry name" value="Sec23-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd11287">
        <property type="entry name" value="Sec23_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.730:FF:000003">
        <property type="entry name" value="Protein transport protein SEC23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.380:FF:000001">
        <property type="entry name" value="Protein transport protein SEC23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1670:FF:000006">
        <property type="entry name" value="Protein transport protein SEC23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.20.10:FF:000003">
        <property type="entry name" value="Protein transport protein SEC23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.410:FF:000011">
        <property type="entry name" value="Protein transport protein SEC23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1670">
        <property type="entry name" value="beta-sandwich domain of Sec23/24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.730">
        <property type="entry name" value="Sec23/Sec24 helical domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.20.10">
        <property type="entry name" value="Severin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.410">
        <property type="entry name" value="von Willebrand factor, type A domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.380">
        <property type="entry name" value="Zn-finger domain of Sec23/24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029006">
        <property type="entry name" value="ADF-H/Gelsolin-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012990">
        <property type="entry name" value="Beta-sandwich_Sec23_24"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007123">
        <property type="entry name" value="Gelsolin-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036180">
        <property type="entry name" value="Gelsolin-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037364">
        <property type="entry name" value="Sec23"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006900">
        <property type="entry name" value="Sec23/24_helical_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036175">
        <property type="entry name" value="Sec23/24_helical_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006896">
        <property type="entry name" value="Sec23/24_trunk_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037550">
        <property type="entry name" value="Sec23_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036465">
        <property type="entry name" value="vWFA_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006895">
        <property type="entry name" value="Znf_Sec23_Sec24"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036174">
        <property type="entry name" value="Znf_Sec23_Sec24_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11141">
        <property type="entry name" value="PROTEIN TRANSPORT PROTEIN SEC23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11141:SF10">
        <property type="entry name" value="PROTEIN TRANSPORT PROTEIN SEC23B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00626">
        <property type="entry name" value="Gelsolin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08033">
        <property type="entry name" value="Sec23_BS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04815">
        <property type="entry name" value="Sec23_helical"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04811">
        <property type="entry name" value="Sec23_trunk"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04810">
        <property type="entry name" value="zf-Sec23_Sec24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81995">
        <property type="entry name" value="beta-sandwich domain of Sec23/24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF82754">
        <property type="entry name" value="C-terminal, gelsolin-like domain of Sec23/24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81811">
        <property type="entry name" value="Helical domain of Sec23/24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53300">
        <property type="entry name" value="vWA-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF82919">
        <property type="entry name" value="Zn-finger domain of Sec23/24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 13" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11 13" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="6 13" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="8 13" id="KW-0931">ER-Golgi transport</keyword>
    <keyword evidence="10 13" id="KW-0472">Membrane</keyword>
    <keyword evidence="5 13" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9 13" id="KW-0653">Protein transport</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 13" id="KW-0813">Transport</keyword>
    <keyword evidence="7 13" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="Zinc finger Sec23/Sec24-type" evidence="15">
        <location>
            <begin position="58"/>
            <end position="98"/>
        </location>
    </feature>
    <feature type="domain" description="Sec23/Sec24 trunk" evidence="16">
        <location>
            <begin position="127"/>
            <end position="380"/>
        </location>
    </feature>
    <feature type="domain" description="Sec23/Sec24 beta-sandwich" evidence="18">
        <location>
            <begin position="391"/>
            <end position="494"/>
        </location>
    </feature>
    <feature type="domain" description="Sec23/Sec24 helical" evidence="17">
        <location>
            <begin position="508"/>
            <end position="606"/>
        </location>
    </feature>
    <feature type="domain" description="Gelsolin-like" evidence="14">
        <location>
            <begin position="622"/>
            <end position="708"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004397"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009210"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022892"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00037370"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="RuleBase" id="RU365030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="Pfam" id="PF00626"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="Pfam" id="PF04810"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF04811"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF04815"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF08033"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016021.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="751" mass="84983" checksum="66449AF412334889" modified="2019-05-08" version="1">MATYLEFIQQNEERDGVRFSWNVWPSSRLEATRMVVPLACLLTPLKERLDLPPVQYEPVLCSRQTCKAVLNPLCQVDYRAKLWACNFCFQRNQFPPAYAGISEINQPAELMPQFSTIEYIVQVNASWLKEDLQALKESLQMSLSLLPPEALVGLITFGRMVQVHELSCEGISKSYVFRGTKDLTAKQIQDMLGLTRPVVPVQQGRPLQPQEQPPISSRFLQPVHKIDMNLTDLLGELQRDPWPVTQGKRPLRSTGVALSIAVGLLEGTFPNTGARIMLFTGGPPTQGPGMVVGDELKTPIRSWHDIEKDNARFMKKATKHYETLANRTAANGHCIDIYACALDQTGLLEMKCCANLTGGHMMMGDSFNTSLFKQTFQRVFSKDFNGEFRMAFGATLEVKTSRELKISGAIGPCVSLNAKGPCVSENELGIGGTCQWKICSLDPSSTLAIYFEVVNQHNAPIPQGGRGAVQFVTQYQHSSTQRRIRVTTIARNWADAQSQLQHIEAAFDQEVAAVLMARLGVYRAESEEGPDVLRWLDRQLIRLCQKFGQYNKDDPNSFRLSDSFSLYPQFMFHLRRSPFLQVFNNSPDESSFYRHHFSRQDLTQSLIMIQPILYSYSFSGPPEPVLLDSSSILADRILLMDTFFQIVIYHGETIAQWQKAGYQDMPEYENFKHLLQAPLDDAQEILQTRFPMPRYINTEHGSSQARFLLSKVNPSQTHNNLYAWGQVSNTFSATLFESHWIRSLVKAFSFP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM56</accession>
    <name>A0A452HM56_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="16">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="9">Endomembrane system</location>
            <topology evidence="9">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="11">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HM56"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018313.1">
        <property type="protein sequence ID" value="ENSGAGP00000016030.1"/>
        <property type="gene ID" value="ENSGAGG00000012038.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012505">
        <property type="term" value="C:endomembrane system"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042734">
        <property type="term" value="C:presynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004222">
        <property type="term" value="F:metalloendopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd04269">
        <property type="entry name" value="ZnMc_adamalysin_II_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.390.10:FF:000014">
        <property type="entry name" value="disintegrin and metalloproteinase domain-containing protein 11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.70.10:FF:000001">
        <property type="entry name" value="Disintegrin and metalloproteinase domain-containing protein 22"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.390.10">
        <property type="entry name" value="Collagenase (Catalytic Domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.70.10">
        <property type="entry name" value="Disintegrin domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006586">
        <property type="entry name" value="ADAM_Cys-rich"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018358">
        <property type="entry name" value="Disintegrin_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001762">
        <property type="entry name" value="Disintegrin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036436">
        <property type="entry name" value="Disintegrin_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024079">
        <property type="entry name" value="MetalloPept_cat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001590">
        <property type="entry name" value="Peptidase_M12B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002870">
        <property type="entry name" value="Peptidase_M12B_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034027">
        <property type="entry name" value="Reprolysin_adamalysin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11905">
        <property type="entry name" value="ADAM A DISINTEGRIN AND METALLOPROTEASE DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11905:SF13">
        <property type="entry name" value="DISINTEGRIN AND METALLOPROTEINASE DOMAIN-CONTAINING PROTEIN 23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08516">
        <property type="entry name" value="ADAM_CR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00200">
        <property type="entry name" value="Disintegrin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01562">
        <property type="entry name" value="Pep_M12B_propep"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01421">
        <property type="entry name" value="Reprolysin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00608">
        <property type="entry name" value="ACR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00050">
        <property type="entry name" value="DISIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57552">
        <property type="entry name" value="Blood coagulation inhibitor (disintegrin)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55486">
        <property type="entry name" value="Metalloproteases ('zincins'), catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50215">
        <property type="entry name" value="ADAM_MEPRO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00427">
        <property type="entry name" value="DISINTEGRIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50214">
        <property type="entry name" value="DISINTEGRIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0165">Cleavage on pair of basic residues</keyword>
    <keyword evidence="7 11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="1 11" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="8" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6 12" id="KW-0472">Membrane</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="3 12" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 12" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="736"/>
            <end position="758"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase M12B" evidence="15">
        <location>
            <begin position="241"/>
            <end position="439"/>
        </location>
    </feature>
    <feature type="domain" description="Disintegrin" evidence="14">
        <location>
            <begin position="445"/>
            <end position="530"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="13">
        <location>
            <begin position="674"/>
            <end position="711"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="10">
        <location>
            <begin position="502"/>
            <end position="522"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="11">
        <location>
            <begin position="701"/>
            <end position="710"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022685"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00046288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00068"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50214"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50215"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016030.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="759" mass="84842" checksum="B23B2CF9DFE52EE4" modified="2019-05-08" version="1">MAGYFSSSALHWNETAGGIAGALSDKENTFLQSNSSSSKNNNQSNEIQKEVTLPSRLVYYINRDSESPYRILDTRARHQQKHNKAVHLAQASFQIEAFGSKFILDLTLNNDLLSSDYVEIHYEDGKPQFSKGGEHCYYHGNIRGVQNSKAALSTCNGLNGMFEDSTYVYFIEPMDLTHSAKSTVRPHIIQRTLGTQTSKQMKNVETESSSEWSFLSELQWLRKRRRKRAVSASRGIFEEMKYLELMIVNDHKMFKKHRSIHAYTNNFAKSVVNLVDAIYKEQLNTRVVLVAVETWTDRDRINIRPDPLQMLHDFSRYRQHYIKQHTDAVHLLSNVTFHYKRSSLSYFGGVCSVTRGVGVNEYALPLAMAQELSQSLAQNLGIQWEPASRKPKCDCIESWGGCIMEETGVYHSRKFSRCSIAEYKEFLLRGGGACLFNRPTKLFETPECGNGYVEAGEECDCGFRMECYENCCKKCSLSNGAHCSDGPCCNTSCLFFPRGYDCRYAVNECDITEYCTGDSGQCPPNLHKQDGYACDSNQGRCYNGECKTRDNQCKYIWGIKATGSDKFCYEKLNTEGTKKGNCGKDGDQWIQCSKHDVFCGFLLCTNLTRVPRIGHLQGEITPTSFYHQGRVVDCSGAHVLLDDNTDLGYVEDGAPCGPHMMCVDRKCLPIQSLNISSCPVGSDGKVCSDHGVCSNEATCICNFSWAGTDCSIDDPIRDTSNKKVEGPKVSMATNRLIGAVAGTILALGVIFGGTGWGIE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM57</accession>
    <name>A0A452HM57_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="15">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="9">Endomembrane system</location>
            <topology evidence="9">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="11">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HM57"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018307.1">
        <property type="protein sequence ID" value="ENSGAGP00000016024.1"/>
        <property type="gene ID" value="ENSGAGG00000012038.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012505">
        <property type="term" value="C:endomembrane system"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042734">
        <property type="term" value="C:presynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004222">
        <property type="term" value="F:metalloendopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd04269">
        <property type="entry name" value="ZnMc_adamalysin_II_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.390.10:FF:000014">
        <property type="entry name" value="disintegrin and metalloproteinase domain-containing protein 11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.70.10:FF:000001">
        <property type="entry name" value="Disintegrin and metalloproteinase domain-containing protein 22"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.390.10">
        <property type="entry name" value="Collagenase (Catalytic Domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.70.10">
        <property type="entry name" value="Disintegrin domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006586">
        <property type="entry name" value="ADAM_Cys-rich"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018358">
        <property type="entry name" value="Disintegrin_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001762">
        <property type="entry name" value="Disintegrin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036436">
        <property type="entry name" value="Disintegrin_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024079">
        <property type="entry name" value="MetalloPept_cat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001590">
        <property type="entry name" value="Peptidase_M12B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002870">
        <property type="entry name" value="Peptidase_M12B_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034027">
        <property type="entry name" value="Reprolysin_adamalysin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11905">
        <property type="entry name" value="ADAM A DISINTEGRIN AND METALLOPROTEASE DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11905:SF13">
        <property type="entry name" value="DISINTEGRIN AND METALLOPROTEINASE DOMAIN-CONTAINING PROTEIN 23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08516">
        <property type="entry name" value="ADAM_CR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00200">
        <property type="entry name" value="Disintegrin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01562">
        <property type="entry name" value="Pep_M12B_propep"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01421">
        <property type="entry name" value="Reprolysin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00608">
        <property type="entry name" value="ACR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00050">
        <property type="entry name" value="DISIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57552">
        <property type="entry name" value="Blood coagulation inhibitor (disintegrin)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55486">
        <property type="entry name" value="Metalloproteases ('zincins'), catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50215">
        <property type="entry name" value="ADAM_MEPRO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00427">
        <property type="entry name" value="DISINTEGRIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50214">
        <property type="entry name" value="DISINTEGRIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0165">Cleavage on pair of basic residues</keyword>
    <keyword evidence="7 11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="1 11" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="8" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6" id="KW-0472">Membrane</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="3" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5" id="KW-1133">Transmembrane helix</keyword>
    <feature type="domain" description="Peptidase M12B" evidence="14">
        <location>
            <begin position="241"/>
            <end position="439"/>
        </location>
    </feature>
    <feature type="domain" description="Disintegrin" evidence="13">
        <location>
            <begin position="445"/>
            <end position="530"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="12">
        <location>
            <begin position="674"/>
            <end position="711"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="10">
        <location>
            <begin position="502"/>
            <end position="522"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="11">
        <location>
            <begin position="701"/>
            <end position="710"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022685"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00046288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00068"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50214"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50215"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016024.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="747" mass="83919" checksum="379A1FFD056F1D82" modified="2019-05-08" version="1">MAGYFSSSALHWNETAGGIAGALSDKENTFLQSNSSSSKNNNQSNEIQKEVTLPSRLVYYINRDSESPYRILDTRARHQQKHNKAVHLAQASFQIEAFGSKFILDLTLNNDLLSSDYVEIHYEDGKPQFSKGGEHCYYHGNIRGVQNSKAALSTCNGLNGMFEDSTYVYFIEPMDLTHSAKSTVRPHIIQRTLGTQTSKQMKNVETESSSEWSFLSELQWLRKRRRKRAVSASRGIFEEMKYLELMIVNDHKMFKKHRSIHAYTNNFAKSVVNLVDAIYKEQLNTRVVLVAVETWTDRDRINIRPDPLQMLHDFSRYRQHYIKQHTDAVHLLSNVTFHYKRSSLSYFGGVCSVTRGVGVNEYALPLAMAQELSQSLAQNLGIQWEPASRKPKCDCIESWGGCIMEETGVYHSRKFSRCSIAEYKEFLLRGGGACLFNRPTKLFETPECGNGYVEAGEECDCGFRMECYENCCKKCSLSNGAHCSDGPCCNTSCLFFPRGYDCRYAVNECDITEYCTGDSGQCPPNLHKQDGYACDSNQGRCYNGECKTRDNQCKYIWGIKATGSDKFCYEKLNTEGTKKGNCGKDGDQWIQCSKHDVFCGFLLCTNLTRVPRIGHLQGEITPTSFYHQGRVVDCSGAHVLLDDNTDLGYVEDGAPCGPHMMCVDRKCLPIQSLNISSCPVGSDGKVCSDHGVCSNEATCICNFSWAGTDCSIDDPIRDTSNKKVEGPKEMSKREGMIQISKVLSEGT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP52</accession>
    <name>A0A452IP52_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">PH domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IP52"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033638.1">
        <property type="protein sequence ID" value="ENSGAGP00000029624.1"/>
        <property type="gene ID" value="ENSGAGG00000021320.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="CDD" id="cd13271">
        <property type="entry name" value="PH2_TAPP1_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000049">
        <property type="entry name" value="pleckstrin homology domain-containing family A member 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051707">
        <property type="entry name" value="PI-Interact_SigTrans_Reg"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14336:SF4">
        <property type="entry name" value="PLECKSTRIN HOMOLOGY DOMAIN-CONTAINING FAMILY A MEMBER 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14336">
        <property type="entry name" value="TANDEM PH DOMAIN CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00169">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00233">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="PH" evidence="2">
        <location>
            <begin position="1"/>
            <end position="41"/>
        </location>
    </feature>
    <feature type="domain" description="PH" evidence="2">
        <location>
            <begin position="122"/>
            <end position="220"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="297"/>
            <end position="318"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="301"/>
            <end position="318"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029624.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="318" mass="36148" checksum="EB237080BD62B7B4" modified="2019-05-08" version="1">LKLSFPNKPTKKLTIVRKYFLQANDQQDLVEWVNVLNKAIKITVPKQSDLLCQPDNPNRQVENPGGKKQVSYRTEIVGGVPIITPTQREELNECGEGGDRNYLKRTQSHLPYFSAKPPPDNAVIKAGYCVKQGAVMKNWKRRYFQLDENTIGYFKSEQEKEPLRIIPLKEVHKVQECKQSDIMMRDNLFEIVTTSRTFYVQADSPEDMHSWIKAISGAIVAQRGPGRSAASMRQARRLSNPCIQRSIPALLQNPAMLSILPTPQPPHIPQPLTTTLWSQTLTPNALPWRSEDFTNLLPRPSQGTSRSRLSLQENQLPK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP56</accession>
    <name>A0A452IP56_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Glycosyl hydrolase family 31 C-terminal domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IP56"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033718.1">
        <property type="protein sequence ID" value="ENSGAGP00000029697.1"/>
        <property type="gene ID" value="ENSGAGG00000021438.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000033719.1">
        <property type="protein sequence ID" value="ENSGAGP00000029698.1"/>
        <property type="gene ID" value="ENSGAGG00000021438.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000033722.1">
        <property type="protein sequence ID" value="ENSGAGP00000029701.1"/>
        <property type="gene ID" value="ENSGAGG00000021438.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.20.20.80">
        <property type="entry name" value="Glycosidases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1180">
        <property type="entry name" value="Golgi alpha-mannosidase II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050985">
        <property type="entry name" value="Alpha-glycosidase_related"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017853">
        <property type="entry name" value="GH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048395">
        <property type="entry name" value="Glyco_hydro_31_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013780">
        <property type="entry name" value="Glyco_hydro_b"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43053">
        <property type="entry name" value="GLYCOSIDASE FAMILY 31"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43053:SF6">
        <property type="entry name" value="SITS-BINDING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21365">
        <property type="entry name" value="Glyco_hydro_31_3rd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51445">
        <property type="entry name" value="(Trans)glycosidases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="1" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="1">
        <location>
            <begin position="7"/>
            <end position="27"/>
        </location>
    </feature>
    <feature type="domain" description="Glycosyl hydrolase family 31 C-terminal" evidence="2">
        <location>
            <begin position="547"/>
            <end position="602"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF21365"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029697.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="603" mass="68121" checksum="C401AB5DECA8CDA8" modified="2019-05-08" version="1">MNETWKGAIACLGVAVFFVMTIGIIYWQVVDQPNKNWILRGSISGLIWERRSHSLVLQTLREEKTLVEINVGNFPDMEQPFVKNLCWLNKTEFCYTWDAMADLEISLQPGPSTGTECYRINWTPLHCKITLKDCFSMTNVSWYGGASIRAQRWPLNNVNTKSQPFVISNLPKNPTGYGSVLEKYFLGSTGVTVTVAPDIPISLSVESNRHFCLESPSSIAMVPLQYMVCISQNITAAHQEVGSQLSELERRLPNTDILGLPIWRHYKAADSAAKMERGLRSFFNRLKRHHLGEGLIALNEHSTILLSNADHMRFPMALRHTRGLSLIKPLKFSITLSPYTSIKSQLFLSSLQEGKENYWLSLQSKFDSHPTPLLTKWKGQFSVRLNVTNDAARSWYMDRVSLLWQRLGAQYVTFEGAEGNTFAEQAIQPPRELEGDKYIGIFALMAATLGNSTIISAITRSNHLPLFIQMTPLQSDWSYAGLKGIIPSVLHYSLLGYNFFIPDAVGGTLANEFLTEEELYIRWLQLVTFLPVMSFSTPPWVCCDDWVLIAPITEQGRVQRDIYLPGEGHKWMDTNTAQVFDGGTLIRNYSVSLVEVPVFVKAS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP62</accession>
    <name>A0A452IP62_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="7">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000033697.1">
        <property type="protein sequence ID" value="ENSGAGP00000029676.1"/>
        <property type="gene ID" value="ENSGAGG00000021419.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031410">
        <property type="term" value="C:cytoplasmic vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032483">
        <property type="term" value="P:regulation of Rab protein signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000042">
        <property type="entry name" value="C-myc promoter-binding protein isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.100.10.50:FF:000001">
        <property type="entry name" value="DENN domain containing 4C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.100.10.50">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.11500">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.10">
        <property type="entry name" value="Tetratricopeptide repeat domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001194">
        <property type="entry name" value="cDENN_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005112">
        <property type="entry name" value="dDENN_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043153">
        <property type="entry name" value="DENN_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051696">
        <property type="entry name" value="DENN_Domain_GEFs"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023341">
        <property type="entry name" value="MABP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002885">
        <property type="entry name" value="PPR_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011990">
        <property type="entry name" value="TPR-like_helical_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037516">
        <property type="entry name" value="Tripartite_DENN"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005113">
        <property type="entry name" value="uDENN_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12296">
        <property type="entry name" value="DENN DOMAIN-CONTAINING PROTEIN 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12296:SF17">
        <property type="entry name" value="DENN DOMAIN-CONTAINING PROTEIN 4C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03455">
        <property type="entry name" value="dDENN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02141">
        <property type="entry name" value="DENN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03456">
        <property type="entry name" value="uDENN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00801">
        <property type="entry name" value="dDENN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00799">
        <property type="entry name" value="DENN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00800">
        <property type="entry name" value="uDENN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50211">
        <property type="entry name" value="DENN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51498">
        <property type="entry name" value="MABP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51375">
        <property type="entry name" value="PPR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0344">Guanine-nucleotide releasing factor</keyword>
    <keyword evidence="1" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="MABP" evidence="6">
        <location>
            <begin position="40"/>
            <end position="199"/>
        </location>
    </feature>
    <feature type="domain" description="UDENN" evidence="5">
        <location>
            <begin position="191"/>
            <end position="641"/>
        </location>
    </feature>
    <feature type="repeat" description="PPR" evidence="3">
        <location>
            <begin position="819"/>
            <end position="853"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="946"/>
            <end position="1040"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="1083"/>
            <end position="1148"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="1237"/>
            <end position="1331"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="1421"/>
            <end position="1467"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="1585"/>
            <end position="1613"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="994"/>
            <end position="1006"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="1021"/>
            <end position="1030"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="1084"/>
            <end position="1095"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="1096"/>
            <end position="1112"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="1293"/>
            <end position="1309"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="1317"/>
            <end position="1331"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="1421"/>
            <end position="1433"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="1453"/>
            <end position="1464"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="1585"/>
            <end position="1600"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022658"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00708"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50211"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS51498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029676.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1863" mass="205720" checksum="3DA9EFF79021291B" modified="2019-05-08" version="1">MIEDKGPRVADYFVVAGLTDTSTPLDQEVNRFETKSAGPKAPITDLAVIIKSAGETVPEGYTCVEATPTALQANLNYGSLKSPELFLCYKRGRDKPPLTDVGVLYEGKERVILGCEVIQATPYGRCANVNNSSATSQRIFITYRRAPPVRPQNSLAVTDICVIVTSKGETPPHTFCKVDKNLNCGMWGSSVYLCYKKSMPASNSIAYKAGLIFRYPEDDYESFPLSESVPLFCLPMGATIECWDPQTKYPLPVFSTFVLTGSSAEKVYGAAIQFYEPYSRELLTEKQCVQLGLLTAVERKVVTSKSINSNTCICLLSHWPFFEAFRKFLMFIYKLSVSGPHPLPIEKHISHFMHNIPFPSPQRPRILVQLSAHDALILSQPVSTPLPLSGANFSTLLMNLGPENCATLLLFVLLEGKILLHSLRPAVLTGVAEAVVAMIFPFQWQCPYIPLCPLSLAAVLSAPVPFIVGVDSRYFDLYDPPPDVVCIDLDTNMVYLSDDKKSVNWKQLPKKPCKSLLNTLRKLHPQLASVHRKTQEGSAVEMTPIEADFSWQKKMTELEMEIQEAFLRFMASILKGYRTFLKPITQAPSNKTTAADSLFDLQGFLKSRDRAYTKFYMLLTKTQIFSRFIEECSFVSDKDTSLAFFDDCIEKLFPDKGTEKGEKVDVDSAEDTRLIELDESQKSEHTVFIMPPEPPPDDGQARAPKYSYKNFPRLDFELFDRPQELKLSFNRHPAGGNISNSPALMAKRTKQEIKTAHKLAKKYYTNPPLWAKCLFSHCYSLWFICLPAYIRVSHPKVRALQQAYDILIKMRKTDVEPLDEVCYRVVMQLCGLCGHPVQAVRVLFEMKNAGVQPNAITYGYYNKAVLESHWPSSNRSGIFLWTKVQNVVRGVAQFRQCLKKATRSSQVSVISGGQSDQGYGSKDELVRDDGDNHLAAEMLAEKDSSCKMKDLTGEVGGTGEKHQQSAEPHSSADPPAWGSSIVKVPSGIFDSTGRKNSSGTASSPLFITQDPVEDSVFNDASPKKTGEKGQKRQKLFSERSCSFSTESRAGMLLKKGSLDLSSSEIAIMMGADAKILTAALSGAKTSPPHTSKSSSFDSMGDQQVSGLSGTRTHVSEGAQELELKSSPVEEPEEGQVHLENGGEDSGARADNVHAKLLDGNQPPNQKTDARDAVTKRNSFYGVIKPIEREDVEVGLDPLSLLATESIEPKPSDPEEKIVSPVVARNLADEIESYMNLKSPLGSKSSSMELHKQESREAAASSTLSHSQERRSSLPSDSVPPAPEKSENRRSPLVSRSKTFTGRLKQQTHSRVQKERSPSLTVLGRSSQHGSLGSVVTSLSNLKLDNILSGPKIDVLKSGMKQAANVASRMWDSVTSQYNYLDDEEETHRPDFTFPPGFEDHILGDNLSPKSGIPGLVTSELTQSTTSLGSSNSSGDVGKQHYTTGEVSFPRSMKGLDSEKSEHGSSHNTSLSSIFQNCAMEVLMSSCSQCRVCGALVYDEEIMAGWAADDSNLNTTCPFCKNTFLPLLNVEFKDLRGSASFFLKPSTSGDSLHSGNIPLTADPLEQKPMSSPAVADLISFSDHLKPNQTIPEETSSATGQSDVPEEQSEGPRAMKTSVINSSKRGASLTRSHSVGGPLQNIDLSHRPSHGVSTVSLPNSLQEVVDPLGKRPNPLPLSVPYLSPLVLRKELESLLENEGEQVIHTSTFINQHPIIFWNLVWYFRRLDLPSNLPGLILTSEHCNDGVQLPLSSLSQDSKLVYIQLLWDNINLHQEPGEPLYITWRNLNSVEKKSSIASEDQQATSTLVENIKLSIQHNNVVEPINLLLQKVKPNIKRQRSFYREILFLSLVSLGRENIDIGKLKIL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP63</accession>
    <name>A0A452IP63_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">XRCC3/RAD51 homolog 2 helix-hairpin-helix domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IP63"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033732.1">
        <property type="protein sequence ID" value="ENSGAGP00000029710.1"/>
        <property type="gene ID" value="ENSGAGG00000021444.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033063">
        <property type="term" value="C:Rad51B-Rad51C-Rad51D-XRCC2 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005657">
        <property type="term" value="C:replication fork"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008094">
        <property type="term" value="F:ATP-dependent activity, acting on DNA"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003690">
        <property type="term" value="F:double-stranded DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000400">
        <property type="term" value="F:four-way junction DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003697">
        <property type="term" value="F:single-stranded DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000724">
        <property type="term" value="P:double-strand break repair via homologous recombination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058766">
        <property type="entry name" value="HHH_XRCC3_RAD51B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR030548">
        <property type="entry name" value="RAD51B"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46456">
        <property type="entry name" value="DNA REPAIR PROTEIN RAD51 HOMOLOG 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46456:SF1">
        <property type="entry name" value="DNA REPAIR PROTEIN RAD51 HOMOLOG 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26169">
        <property type="entry name" value="HHH_XRCC3_RpoA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0227">DNA damage</keyword>
    <keyword evidence="4" id="KW-0233">DNA recombination</keyword>
    <keyword evidence="3" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="5" id="KW-0539">Nucleus</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="XRCC3/RAD51 homolog 2 helix-hairpin-helix" evidence="6">
        <location>
            <begin position="6"/>
            <end position="60"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023172"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="Pfam" id="PF26169"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029710.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="128" mass="14078" checksum="CDA661FBBEB97BC4" modified="2019-05-08" version="1">MATKKLRRAGLSQELCDRLSRHQIITCRDFLCLSLLELMKVTGKSYWNVQKLLCKVSLACAPKMQTAYEMKMKTAINPSSAFLSTTLNGLDKALHGGVACGSLTEVKLIFTVFTAVVPKGHNLSWGSI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP68</accession>
    <name>A0A452IP68_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="23">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 16">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000033950.1">
        <property type="protein sequence ID" value="ENSGAGP00000029897.1"/>
        <property type="gene ID" value="ENSGAGG00000021570.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005884">
        <property type="term" value="C:actin filament"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030027">
        <property type="term" value="C:lamellipodium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016459">
        <property type="term" value="C:myosin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001726">
        <property type="term" value="C:ruffle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000146">
        <property type="term" value="F:microfilament motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030048">
        <property type="term" value="P:actin filament-based movement"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072673">
        <property type="term" value="P:lamellipodium morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007266">
        <property type="term" value="P:Rho protein signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd01385">
        <property type="entry name" value="MYSc_Myo9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd17217">
        <property type="entry name" value="RA_Myosin-IXb"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04407">
        <property type="entry name" value="RhoGAP_myosin_IXB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.720:FF:000003">
        <property type="entry name" value="Putative unconventional myosin-IXa"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000008">
        <property type="entry name" value="Putative unconventional myosin-IXa"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.820:FF:000003">
        <property type="entry name" value="unconventional myosin-IXa isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.555.10:FF:000009">
        <property type="entry name" value="unconventional myosin-IXa isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.530:FF:000005">
        <property type="entry name" value="unconventional myosin-IXa isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000013">
        <property type="entry name" value="unconventional myosin-IXa isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.190:FF:000013">
        <property type="entry name" value="unconventional myosin-IXa isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.530:FF:000009">
        <property type="entry name" value="unconventional myosin-IXb isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.190">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.60.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.20.240.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.555.10">
        <property type="entry name" value="Rho GTPase activation protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046349">
        <property type="entry name" value="C1-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000048">
        <property type="entry name" value="IQ_motif_EF-hand-BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046987">
        <property type="entry name" value="Myo9"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036023">
        <property type="entry name" value="MYSc_Myo9"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002219">
        <property type="entry name" value="PKC_DAG/PE"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000159">
        <property type="entry name" value="RA_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046989">
        <property type="entry name" value="RA_Myosin-IXb"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000198">
        <property type="entry name" value="RhoGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028557">
        <property type="entry name" value="RhoGAP_myosin_IXB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029071">
        <property type="entry name" value="Ubiquitin-like_domsf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46184:SF2">
        <property type="entry name" value="UNCONVENTIONAL MYOSIN-IXB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46184">
        <property type="entry name" value="UNCONVENTIONAL MYOSIN-IXB-LIKE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00612">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00788">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00620">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00109">
        <property type="entry name" value="C1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00015">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00314">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00324">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57889">
        <property type="entry name" value="Cysteine-rich domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54236">
        <property type="entry name" value="Ubiquitin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50200">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50238">
        <property type="entry name" value="RHOGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00479">
        <property type="entry name" value="ZF_DAG_PE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50081">
        <property type="entry name" value="ZF_DAG_PE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="15 16" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="11 16" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="12 17" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="3" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="6" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="14 16" id="KW-0505">Motor protein</keyword>
    <keyword evidence="13 16" id="KW-0518">Myosin</keyword>
    <keyword evidence="8 16" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="10" id="KW-0862">Zinc</keyword>
    <keyword evidence="9" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="Ras-associating" evidence="20">
        <location>
            <begin position="15"/>
            <end position="113"/>
        </location>
    </feature>
    <feature type="domain" description="Myosin motor" evidence="22">
        <location>
            <begin position="145"/>
            <end position="978"/>
        </location>
    </feature>
    <feature type="domain" description="Phorbol-ester/DAG-type" evidence="19">
        <location>
            <begin position="1683"/>
            <end position="1732"/>
        </location>
    </feature>
    <feature type="domain" description="Rho-GAP" evidence="21">
        <location>
            <begin position="1751"/>
            <end position="1939"/>
        </location>
    </feature>
    <feature type="region of interest" description="Actin-binding" evidence="16">
        <location>
            <begin position="860"/>
            <end position="882"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1125"/>
            <end position="1178"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1220"/>
            <end position="1259"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1276"/>
            <end position="1394"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1504"/>
            <end position="1532"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1656"/>
            <end position="1677"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1966"/>
            <end position="1997"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="2018"/>
            <end position="2053"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="17">
        <location>
            <begin position="1066"/>
            <end position="1093"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="1148"/>
            <end position="1173"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="1220"/>
            <end position="1233"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="1276"/>
            <end position="1287"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="1303"/>
            <end position="1313"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="1342"/>
            <end position="1354"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="1380"/>
            <end position="1394"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="1517"/>
            <end position="1528"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="18">
        <location>
            <begin position="1664"/>
            <end position="1674"/>
        </location>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <begin position="238"/>
            <end position="245"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50081"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50200"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50238"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029897.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2201" mass="253021" checksum="2805C7CE8F9A4B76" modified="2019-05-08" version="1">MSLKDADSAVCQEKATYNLHIYPQLSTESTTCCRVTATKDSTTSDVIKDVIQILDLDVSKHYVLVEVKESGGEEWVLDLNDSPVHRVLLWPRRAQDEHPQNDGYYFLLQERDTDGTIKYVHMQLVSQEKDARRLVERGFLPWHQENFDDLCNLSNLTETTLLENLKRRFLKHKIYTYAGSILIAINPFKFLPIYNPKYVKMYENHQLGKLEPHIFAIADVAYHTMLKKHVNQCIVISGESGSGKTQSTNFLIHCFTALSQKGYASGVERTILGAGPVLEAFGNAKTAHNNNSSRFGKFIQVNYLENGIVRGAVVEKYLLEKSRLVSQEKDERNYHVFYYLLLGVSEEERKEFHLKQPEDYFYLNQHNLKIEDGEDLRHDFERLKQAMEMVGFLPATKKQIFSVLSAILYLGNVTYKKRATGRDEGLEVGPTEVLDTLSQLLKVKREVLVEVLTKRKTVTANDKLILPYSLSEAITARDSMAKSLYSALFDWIVLRINHALLNKKDMEESVSCLSIGVLDIFGFEDFETNSFEQFCINYANEQLQYYFNQHIFKLEQEEYQSEGIAWHNIDYTDNVTCIHLISKKPTGLFYLLDEESNFPHATNQTLLAKFKQQHEENKFFVGTPVKEPAFIIRHFAGKVKYQIKDFREKNMDYMRPDIVALLRSSDSAYVRELIGMDPVAVFRWAILRAAIRAMAVFAEAGRQRAQKTAGVVRHGPRVPLGELQRANTPIEKVYRCSVLDFSFDCSEDFDINAFEDIIAFYESKKDLHDQIIKSIKGLPWQGEDPCRLLRSLSRLQQPHVHFLKSRGTKQKQIIPKNLLDSKSLKLIVSMTLHDRTTKSLLHLHKKKKPPSISAQFQNSLNKLLETLGKAEPFFIRCIRSNAEKRELFFDEGLVLQQLRYTGMLETVRIRRSGYSAKYTFQDFIDQFQVLLPQNAKASKEEISAHLNKLKLDKNNYQIGKTKVFMKEAERQMLQDTLHKEVIRKIVLLQSWLRMVLERRHFLRMRQATITLQTCWRSYRIRRALERTNAAIYIQSVWRRYRERESYHQQRRRISLVQALVRGHLQRKRFQQMIVEKQQEMQQAQEGEDSVSQEDGNELVLEQLAVKPEMQLDQGIEQMAAGVDEAQNDGDEKLSQATEKAVPSQKSITDGHEKLTSSREKRESRRQRGLEHNKLQNKHVLFPLKEQSLVCQEEETSAKETPETVQEPNKFTAYKTVLPKGTEEERIPVEENKTLSDTQVPGEILDSDVSFPKQSNESKQEILDDKIIDKLKIQDNQKNQVKGSQSFTCPERPSDFSLHIKKNLTPTRSFQTGDENYGNKNKHRISKVDKDLDSPPSFQIQRYLHDPGELRDKREKWKGKRQSEAGQNDVLSESLDERNRTGQSPQTLLSKKGTTSSLDDIANLAQSVTMIQPSPETMAEEKSNKKQTLQRKSNEQLSLLDLAVSVQLPSQPMDAKSTFKSPLRKLLGKKPDKKYVRESPDVMEEGENLPLLSCILISESVESQKLSEAPSGHPRRPQAGERPVKETSKGKRNRTIKIKKISAVSEKWPASVIREITNANELKYLDEFLLNKINDLNSQKSATECLFFEATEKFRGNIKTMYSVPNGQIHVGYKDLIENYQLLVINLAKEREEKDVKLVLNLFQSLLDEFSRGYAKREESEQHKQSKTQKKKRKQERAIEEHNGHVFTSYQVSIRQSCEQCSSYIWPMEKACLCSTCKLICHKKCMSKTQSCCTSPCGKKNEQGVDSRHFGVCVSSLISDKNSVPVVMEKLLEYVEMHGLYTEGIYRKSGSANRMRELKQLLQTDPNSVKLENYPIHTITGILKQWLRELPDPLMTFAQYSDFLRAVELPEKQEQLIAIYSVLEQLPQANHNTLERLIFHLVKVALIEDVNRMSPNALAIVFAPCLLRCPDNSDPLTSMKDVSKTTMCVEMLIKEQMRKYKIKMDEINQLEAAESIAFRRLSLLRQNTSKSPQDKGNDDSFPELGSLQEEDEVSDVDNREKEILIERIQSIKEEKEDITYRLPELDQQGSDEENLDSETSVSTESLPEDRTGRLDTEGTMLSEFQCHGQCPSVPAKDTCGGPSISLSLASTFSSVTRRRPSSVLVKMKVPCRNPVIPTANIKLPYGIPKLTESQRRTSAEESQPVVRRRDRLARRTDKIHSVYIAQGSVVAQTQDPLDEYEPTAKVKRRFSDPYSHISCIDE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP71</accession>
    <name>A0A452IP71_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Adhesion G-protein coupled receptor V1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="20">G-protein coupled receptor 98</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="19">Very large G-protein coupled receptor 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cell membrane</location>
            <topology evidence="3">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cell projection</location>
            <location evidence="1">Stereocilium membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Photoreceptor inner segment</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the G-protein coupled receptor 2 family. Adhesion G-protein coupled receptor (ADGR) subfamily.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000033654.1">
        <property type="protein sequence ID" value="ENSGAGP00000029639.1"/>
        <property type="gene ID" value="ENSGAGG00000021364.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001917">
        <property type="term" value="C:photoreceptor inner segment"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060171">
        <property type="term" value="C:stereocilium membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010855">
        <property type="term" value="F:adenylate cyclase inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004930">
        <property type="term" value="F:G protein-coupled receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001965">
        <property type="term" value="F:G-protein alpha-subunit binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016787">
        <property type="term" value="F:hydrolase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048513">
        <property type="term" value="P:animal organ development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007166">
        <property type="term" value="P:cell surface receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071277">
        <property type="term" value="P:cellular response to calcium ion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007605">
        <property type="term" value="P:sensory perception of sound"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007601">
        <property type="term" value="P:visual perception"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.2030:FF:000017">
        <property type="entry name" value="Adhesion G protein-coupled receptor V1"/>
        <property type="match status" value="4"/>
    </dbReference>
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        <property type="entry name" value="Adhesion G protein-coupled receptor V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.2030:FF:000021">
        <property type="entry name" value="Adhesion G protein-coupled receptor V1"/>
        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.2030:FF:000007">
        <property type="entry name" value="Adhesion G-protein coupled receptor V1"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.2030:FF:000009">
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.2030:FF:000012">
        <property type="entry name" value="Adhesion G-protein coupled receptor V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.2030:FF:000013">
        <property type="entry name" value="Adhesion G-protein coupled receptor V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.2030:FF:000014">
        <property type="entry name" value="Adhesion G-protein coupled receptor V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.2030:FF:000028">
        <property type="entry name" value="Adhesion G-protein coupled receptor V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.200">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.220.50">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.2030">
        <property type="match status" value="34"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1070.10">
        <property type="entry name" value="Rhodopsin 7-helix transmembrane proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026919">
        <property type="entry name" value="ADGRV1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038081">
        <property type="entry name" value="CalX-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003644">
        <property type="entry name" value="Calx_beta"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009039">
        <property type="entry name" value="EAR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005492">
        <property type="entry name" value="EPTP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057244">
        <property type="entry name" value="GAIN_B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046338">
        <property type="entry name" value="GAIN_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017981">
        <property type="entry name" value="GPCR_2-like_7TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000832">
        <property type="entry name" value="GPCR_2_secretin-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46682">
        <property type="entry name" value="ADHESION G-PROTEIN COUPLED RECEPTOR V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46682:SF1">
        <property type="entry name" value="ADHESION G-PROTEIN COUPLED RECEPTOR V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00002">
        <property type="entry name" value="7tm_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03160">
        <property type="entry name" value="Calx-beta"/>
        <property type="match status" value="33"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03736">
        <property type="entry name" value="EPTP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13385">
        <property type="entry name" value="Laminin_G_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00237">
        <property type="entry name" value="Calx_beta"/>
        <property type="match status" value="18"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF75011">
        <property type="entry name" value="3-carboxy-cis,cis-mucoante lactonizing enzyme"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF141072">
        <property type="entry name" value="CalX-like"/>
        <property type="match status" value="37"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50912">
        <property type="entry name" value="EAR"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50261">
        <property type="entry name" value="G_PROTEIN_RECEP_F2_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50221">
        <property type="entry name" value="GAIN_B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0106">Calcium</keyword>
    <keyword evidence="5" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="17" id="KW-0966">Cell projection</keyword>
    <keyword evidence="14" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="12" id="KW-0297">G-protein coupled receptor</keyword>
    <keyword evidence="9" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="13 21" id="KW-0472">Membrane</keyword>
    <keyword evidence="15" id="KW-0675">Receptor</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
    <keyword evidence="16" id="KW-0807">Transducer</keyword>
    <keyword evidence="6 21" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 21" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="27"/>
            <end position="46"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="5841"/>
            <end position="5861"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="5873"/>
            <end position="5892"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="5912"/>
            <end position="5932"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="5944"/>
            <end position="5970"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="5990"/>
            <end position="6013"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="6034"/>
            <end position="6054"/>
        </location>
    </feature>
    <feature type="domain" description="GAIN-B" evidence="22">
        <location>
            <begin position="5677"/>
            <end position="5836"/>
        </location>
    </feature>
    <feature type="domain" description="G-protein coupled receptors family 2 profile 2" evidence="23">
        <location>
            <begin position="5838"/>
            <end position="6013"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004289"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004437"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00007343"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023224"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00070037"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00078072"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00083929"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS50221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS50261"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029639.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="6094" mass="669952" checksum="8745ABF9CFBB71FF" modified="2019-05-08" version="1">[encoded data, 6094 characters]</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP73</accession>
    <name>A0A452IP73_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">Copine-3</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="18">Copine III</fullName>
        </alternativeName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">Calcium-dependent phospholipid-binding protein that plays a role in ERBB2-mediated tumor cell migration in response to growth factor heregulin stimulation.</text>
    </comment>
    <comment type="subunit">
        <text evidence="16">Monomer. Interacts with ERBB2 (preferentially with the tyrosine phosphorylated form); this interaction occurs at the cell membrane and is increased in a growth factor heregulin-dependent manner. Interacts with SHC1; this interaction may mediate the binding of CPNE3 with ERBB2. Interacts with RACK1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cell junction</location>
            <location evidence="3">Focal adhesion</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the copine family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IP73"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033816.1">
        <property type="protein sequence ID" value="ENSGAGP00000029779.1"/>
        <property type="gene ID" value="ENSGAGG00000021499.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005925">
        <property type="term" value="C:focal adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005544">
        <property type="term" value="F:calcium-dependent phospholipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071277">
        <property type="term" value="P:cellular response to calcium ion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903265">
        <property type="term" value="P:positive regulation of tumor necrosis factor-mediated signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043122">
        <property type="term" value="P:regulation of canonical NF-kappaB signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd04048">
        <property type="entry name" value="C2A_Copine"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04047">
        <property type="entry name" value="C2B_Copine"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd01459">
        <property type="entry name" value="vWA_copine_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000042">
        <property type="entry name" value="Copine 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000099">
        <property type="entry name" value="Copine 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000008">
        <property type="entry name" value="C2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037768">
        <property type="entry name" value="C2B_Copine"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045052">
        <property type="entry name" value="Copine"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010734">
        <property type="entry name" value="Copine_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002035">
        <property type="entry name" value="VWF_A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036465">
        <property type="entry name" value="vWFA_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10857">
        <property type="entry name" value="COPINE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10857:SF2">
        <property type="entry name" value="COPINE-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00168">
        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07002">
        <property type="entry name" value="Copine"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00239">
        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00327">
        <property type="entry name" value="VWA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49562">
        <property type="entry name" value="C2 domain (Calcium/lipid-binding domain, CaLB)"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53300">
        <property type="entry name" value="vWA-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50004">
        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50234">
        <property type="entry name" value="VWFA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="11" id="KW-0106">Calcium</keyword>
    <keyword evidence="12" id="KW-0965">Cell junction</keyword>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="13" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="14" id="KW-0539">Nucleus</keyword>
    <keyword evidence="8" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="C2" evidence="19">
        <location>
            <begin position="1"/>
            <end position="114"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="19">
        <location>
            <begin position="123"/>
            <end position="245"/>
        </location>
    </feature>
    <feature type="domain" description="VWFA" evidence="20">
        <location>
            <begin position="289"/>
            <end position="509"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00009048"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00058857"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00065466"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00074834"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00076171"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50234"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029779.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="533" mass="59444" checksum="AB3B44C32EBD391C" modified="2019-05-08" version="1">MAQCVTKVELSISCENLIDKDVGSKSDPLCVLLQNVGGDQWAELDRTEKIKNCQKPEFSKKLLIDYYFEKVQKLKFGVYDIDNKSYDLNDDDYLGGVECTLGQIVSSKTFTRPLELKKGKPAGKGTITISAEEIKDTRIVSLEIEAQNLDKKDFLGKSDPFLEFYKPSDGGKWQLIYRSEVIKNNLNPCWRKFSVPLQTFCGGDFNKPIKVSVSDMDESTSSDLIGEFTCITAKLLEARDHVVEFECIHPGKKQKKKSYKNSGIVRIKSCKIETEYSFLDYIMGGCQINFTVGIDFTGSNGDPKSPDSLHYISPNGINEYLTAIWSVGNVVQDYDTDKLFPAFGFGAQVPPNWQVSHEFALNFNPNNPYCQGIQGIVDAYRQALPQVRLYGPTNFSPIINHVARFAAHSAQQGTAAQYFVLLIITDGEITDLDQTRQAIVNASKLPMSVIIVGVGNADFKAMEFLDGDNGVLKSLTGEPAVRDIVQFVPFRQFLNAPREALSQTVLAEVPKQLVSYYKMQGWAPVKLPGAKQE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP74</accession>
    <name>A0A452IP74_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IP74"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033707.1">
        <property type="protein sequence ID" value="ENSGAGP00000029686.1"/>
        <property type="gene ID" value="ENSGAGG00000021428.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="145"/>
            <end position="170"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="145"/>
            <end position="167"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029686.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="192" mass="22569" checksum="795473743AE7D5B9" modified="2019-05-08" version="1">IFPAKNKAFRVQNKVFHKDNQALRAENQALSEDNKILQERKMDYFEEKKVLEKQEKALRKQNKALWEEIKALQEQEIAFQMEEKALQEETVALWEENKAFQVQIKALQKDEIALQEEKKALQKQGKALQEEKIALREENRALHEENKALQNEKKALDEENKPRKVEGGGKQQVGLLQGHPLEWHAAHHFCRI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP76</accession>
    <name>A0A452IP76_9SAUR</name>
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        <submittedName>
            <fullName evidence="8">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
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            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IP76"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033752.1">
        <property type="protein sequence ID" value="ENSGAGP00000029725.1"/>
        <property type="gene ID" value="ENSGAGG00000021459.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007265">
        <property type="term" value="P:Ras protein signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd17210">
        <property type="entry name" value="RA_RGL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00155">
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    </dbReference>
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        <property type="entry name" value="REM"/>
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    </dbReference>
    <dbReference type="FunFam" id="1.10.840.10:FF:000005">
        <property type="entry name" value="Ral guanine nucleotide dissociation stimulator isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.870.10:FF:000003">
        <property type="entry name" value="Ral guanine nucleotide dissociation stimulator isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Ral guanine nucleotide dissociation stimulator isoform 1"/>
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    </dbReference>
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        <property type="entry name" value="Phosphatidylinositol 3-kinase Catalytic Subunit, Chain A, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.870.10">
        <property type="entry name" value="Son of sevenless (SoS) protein Chain: S domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="RA_dom"/>
    </dbReference>
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        <property type="entry name" value="Ras-like_GEF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000651">
        <property type="entry name" value="Ras-like_Gua-exchang_fac_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019804">
        <property type="entry name" value="Ras_G-nucl-exch_fac_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023578">
        <property type="entry name" value="Ras_GEF_dom_sf"/>
    </dbReference>
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    </dbReference>
    <dbReference type="InterPro" id="IPR036964">
        <property type="entry name" value="RASGEF_cat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR030748">
        <property type="entry name" value="RGL1_RA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029071">
        <property type="entry name" value="Ubiquitin-like_domsf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23113">
        <property type="entry name" value="GUANINE NUCLEOTIDE EXCHANGE FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23113:SF199">
        <property type="entry name" value="RAL GUANINE NUCLEOTIDE DISSOCIATION STIMULATOR-LIKE 1"/>
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    </dbReference>
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    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
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    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
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    </dbReference>
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        <property type="match status" value="1"/>
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    <keyword evidence="2 3" id="KW-0344">Guanine-nucleotide releasing factor</keyword>
    <keyword evidence="1" id="KW-0597">Phosphoprotein</keyword>
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    <feature type="domain" description="N-terminal Ras-GEF" evidence="7">
        <location>
            <begin position="65"/>
            <end position="196"/>
        </location>
    </feature>
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        <location>
            <begin position="232"/>
            <end position="501"/>
        </location>
    </feature>
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        <location>
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            <end position="735"/>
        </location>
    </feature>
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            <end position="622"/>
        </location>
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        <location>
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            <end position="561"/>
        </location>
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    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="586"/>
            <end position="602"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="603"/>
            <end position="621"/>
        </location>
    </feature>
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        <source>
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        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022658"/>
        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
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        <source>
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        </source>
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        </source>
    </evidence>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029725.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="768" mass="86985" checksum="FDF184322819C46A" modified="2019-05-08" version="1">MKLLWKAKMSSIQDGGEEVEEGAVYHVTLKRVQIQQAANKGARWLGVEGDQLPPGHTVSQYETCKIRTIKAGTLEKLVENLLTAFGDNDFTYISIFLSTYRGFASTKEVLERLLDRYGNLETSNCEEDGSQNSSESKTVLRNAIASILRAWLDQCSEDFREPPNYPCLQKLLDYLKRIMPGSDPERRAQNLLEQLQKQEVKNENGFHSTFTFSFDEEEEMETGGPEEFSFFPEDLVAEQLTYMDAKLFKKVVPHHCLGCIWSRRDKKENKHLAPTIRATISQFNAVTKCVVSTILKGKDLKTQQRAKIIEKWIHIAHECRILKNFSSLRAIVSALQSNSIYRLKKTWASVPKDRMLMFEELSDIFSDHDNYLTSRELLMKEGTSKFANLDSSVKENQKRTQRRLQLQKDMGVMQGTVPYLGTFLTDLTMLDTALQDYIEGGLINFEKRRREFEVIAQIKLLQSSCNSYCMTPDQKFIRWFRRQRLLTEEESYGLSCEIEAAADTNTTSPKPRKSMVKRLSLLFLGADVITNSTPTKEQPKSTPSGSSGESMDSVSVSSCESNHFESEEVCITPIDTPDEPQKKLSESSSSCSSIHSMDTSSSGVSSLTNTISPPPSLTSNPKIHKRSISVTSITSAVLPPVYNQQNEDMCIIRISIEDNNGNMYKSIMLTSQDKTPAVIQRAMSKHNLEADPVEDYELVQVIAEDKELVIPDSANVFYAMNSQVNFDFILRKKTSLDGQVKMRSRSSLTLPRTAKRGCWTNRHSKITL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP79</accession>
    <name>A0A452IP79_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Kelch like family member 20</fullName>
        </recommendedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
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            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IP79"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033712.1">
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        <property type="gene ID" value="ENSGAGG00000021434.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="FunFam" id="2.120.10.80:FF:000021">
        <property type="entry name" value="kelch-like protein 1 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.120.10.80">
        <property type="entry name" value="Kelch-type beta propeller"/>
        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
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    </dbReference>
    <dbReference type="PANTHER" id="PTHR45632:SF13">
        <property type="entry name" value="KELCH-LIKE PROTEIN 26"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45632">
        <property type="entry name" value="LD33804P"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01344">
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        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00612">
        <property type="entry name" value="Kelch"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF117281">
        <property type="entry name" value="Kelch motif"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0880">Kelch repeat</keyword>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022441"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029691.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="4">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="145" mass="16104" checksum="19B96F9B74763758" modified="2019-05-08" version="1">MIKDLRDIDGHSLEREENFRRMEKVYPGTYIQTKGCVGVATCDGFLYAVGGHDAPASNHCSRLLDYVERYDPKTDTWTMVAPLSMPRDAVGVCLLGDKLYAVGGYDGQTYLNTMESYDPQTNEWTLMASLNIGRAGACIVVIKQP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP81</accession>
    <name>A0A452IP81_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="7">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IP81"/>
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        <property type="protein sequence ID" value="ENSGAGP00000029731.1"/>
        <property type="gene ID" value="ENSGAGG00000021459.1"/>
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        <property type="component" value="Unassembled WGS sequence"/>
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        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
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        <property type="project" value="UniProtKB-KW"/>
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        <property type="term" value="P:Ras protein signal transduction"/>
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        <property type="match status" value="1"/>
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        <property type="entry name" value="Ral guanine nucleotide dissociation stimulator isoform 1"/>
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    </dbReference>
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        <property type="entry name" value="Phosphatidylinositol 3-kinase Catalytic Subunit, Chain A, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Ras guanine-nucleotide exchange factors catalytic domain"/>
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    </dbReference>
    <dbReference type="Gene3D" id="1.20.870.10">
        <property type="entry name" value="Son of sevenless (SoS) protein Chain: S domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000159">
        <property type="entry name" value="RA_dom"/>
    </dbReference>
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        <property type="entry name" value="Ras-like_GEF"/>
    </dbReference>
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        <property type="entry name" value="Ras-like_Gua-exchang_fac_N"/>
    </dbReference>
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        <property type="entry name" value="Ras_G-nucl-exch_fac_CS"/>
    </dbReference>
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        <property type="entry name" value="Ras_GEF_dom_sf"/>
    </dbReference>
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        <property type="entry name" value="RASGEF_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036964">
        <property type="entry name" value="RASGEF_cat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR030748">
        <property type="entry name" value="RGL1_RA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029071">
        <property type="entry name" value="Ubiquitin-like_domsf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23113">
        <property type="entry name" value="GUANINE NUCLEOTIDE EXCHANGE FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23113:SF199">
        <property type="entry name" value="RAL GUANINE NUCLEOTIDE DISSOCIATION STIMULATOR-LIKE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00788">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="RasGEF"/>
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    </dbReference>
    <dbReference type="Pfam" id="PF00618">
        <property type="entry name" value="RasGEF_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00314">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00147">
        <property type="entry name" value="RasGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00229">
        <property type="entry name" value="RasGEFN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48366">
        <property type="entry name" value="Ras GEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54236">
        <property type="entry name" value="Ubiquitin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50200">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00720">
        <property type="entry name" value="RASGEF"/>
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    <dbReference type="PROSITE" id="PS50009">
        <property type="entry name" value="RASGEF_CAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50212">
        <property type="entry name" value="RASGEF_NTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1 2" id="KW-0344">Guanine-nucleotide releasing factor</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="N-terminal Ras-GEF" evidence="6">
        <location>
            <begin position="65"/>
            <end position="196"/>
        </location>
    </feature>
    <feature type="domain" description="Ras-GEF" evidence="4">
        <location>
            <begin position="163"/>
            <end position="429"/>
        </location>
    </feature>
    <feature type="domain" description="Ras-associating" evidence="5">
        <location>
            <begin position="576"/>
            <end position="663"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="459"/>
            <end position="550"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="3">
        <location>
            <begin position="469"/>
            <end position="489"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="3">
        <location>
            <begin position="514"/>
            <end position="530"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="531"/>
            <end position="549"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022658"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
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            <dbReference type="PROSITE-ProRule" id="PRU00168"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50009"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50200"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50212"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029731.1"/>
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    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="696" mass="78662" checksum="8016F3F8126432FF" modified="2019-05-08" version="1">MKLLWKAKMSSIQDGGEEVEEGAVYHVTLKRVQIQQAANKGARWLGVEGDQLPPGHTVSQYETCKIRTIKAGTLEKLVENLLTAFGDNDFTYISIFLSTYRGFASTKEVLERLLDRYGNLETSNCEEDGSQNSSESKTVLRNAIASILRAWLDQCSEDFREPPNYPCLQKLLDYLKRIMPGSDPERRAQNLLEQLQKQEVKNENGFHSTFTFSFDEEEEMETGGPEEFSFFPEDLVAEQLTYMDAECRILKNFSSLRAIVSALQSNSIYRLKKTWASVPKDRMLMFEELSDIFSDHDNYLTSRELLMKEGTSKFANLDSSVKENQKRTQRRLQLQKDMGVMQGTVPYLGTFLTDLTMLDTALQDYIEGGLINFEKRRREFEVIAQIKLLQSSCNSYCMTPDQKFIRWFRRQRLLTEEESYGLSCEIEAAADTNTTSPKPRKSMVKRLSLLFLGADVITNSTPTKEQPKSTPSGSSGESMDSVSVSSCESNHFESEEVCITPIDTPDEPQKKLSESSSSCSSIHSMDTSSSGVSSLTNTISPPPSLTSNPKIHKRSISVTSITSAVLPPVYNQQNEDMCIIRISIEDNNGNMYKSIMLTSQDKTPAVIQRAMSKHNLEADPVEDYELVQVIAEDKELVIPDSANVFYAMNSQVNFDFILRKKTSLDGQVKMRSRSSLTLPRTAKRGCWTNRHSKITL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP82</accession>
    <name>A0A452IP82_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Amine oxidase</fullName>
            <ecNumber evidence="14">1.4.3.-</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="12">
            <text>a secondary aliphatic amine + O2 + H2O = a primary amine + an aldehyde + H2O2</text>
            <dbReference type="Rhea" id="RHEA:26414"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15379"/>
            <dbReference type="ChEBI" id="CHEBI:16240"/>
            <dbReference type="ChEBI" id="CHEBI:17478"/>
            <dbReference type="ChEBI" id="CHEBI:58855"/>
            <dbReference type="ChEBI" id="CHEBI:65296"/>
            <dbReference type="EC" id="1.4.3.4"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 14">
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Mitochondrion outer membrane</location>
            <topology evidence="2">Single-pass type IV membrane protein</topology>
            <orientation evidence="2">Cytoplasmic side</orientation>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 14">Belongs to the flavin monoamine oxidase family.</text>
    </comment>
    <dbReference type="EC" id="1.4.3.-" evidence="14"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033664.1">
        <property type="protein sequence ID" value="ENSGAGP00000029649.1"/>
        <property type="gene ID" value="ENSGAGG00000021218.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005741">
        <property type="term" value="C:mitochondrial outer membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050660">
        <property type="term" value="F:flavin adenine dinucleotide binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097621">
        <property type="term" value="F:monoamine oxidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008131">
        <property type="term" value="F:primary methylamine oxidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009308">
        <property type="term" value="P:amine metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.405.10:FF:000005">
        <property type="entry name" value="Amine oxidase [flavin-containing]"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.660.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.130">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.50.50.60">
        <property type="entry name" value="FAD/NAD(P)-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.405.10">
        <property type="entry name" value="Guanine Nucleotide Dissociation Inhibitor, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002937">
        <property type="entry name" value="Amino_oxidase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036188">
        <property type="entry name" value="FAD/NAD-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001613">
        <property type="entry name" value="Flavin_amine_oxidase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050703">
        <property type="entry name" value="Flavin_MAO"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43563">
        <property type="entry name" value="AMINE OXIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43563:SF16">
        <property type="entry name" value="AMINE OXIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01593">
        <property type="entry name" value="Amino_oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00757">
        <property type="entry name" value="AMINEOXDASEF"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54373">
        <property type="entry name" value="FAD-linked reductases, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51905">
        <property type="entry name" value="FAD/NAD(P)-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7 14" id="KW-0274">FAD</keyword>
    <keyword evidence="4 14" id="KW-0285">Flavoprotein</keyword>
    <keyword evidence="11 14" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="6" id="KW-1000">Mitochondrion outer membrane</keyword>
    <keyword evidence="9 14" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 14" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 14" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="511"/>
            <end position="532"/>
        </location>
    </feature>
    <feature type="domain" description="Amine oxidase" evidence="15">
        <location>
            <begin position="39"/>
            <end position="470"/>
        </location>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <begin position="57"/>
            <end position="58"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <position position="258"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <position position="368"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <position position="446"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001974"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004362"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005995"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022630"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022787"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022827"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00048448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PIRSR" id="PIRSR601613-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="RuleBase" id="RU362067"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="Pfam" id="PF01593"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029649.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="540" mass="60942" checksum="8242DE42B9617872" modified="2019-05-08" version="1">MTPLTKYLGALTTDCLEATQATESTVWSCSVLFLICPLGLSAAKLLTESGLNVVVLEARDRVGGRTFTARNKQVQYVDLGGAYVGPTQNRILRLSKELGIETYKVNEVEHLIHHVKGKSYPFKGAFPTMWNPFVILDFNNLWRTMDEMGKEIPNEAPWKAPHAEEWDKMSMQDFIDKVCWTNASKRFATLFVNVDVTSEPHEVSALWFLWYVKQCGGTGRIFSTTNGGQERKFVGGSGQISEKLMERLGGRVKLEKPVVRIDQSGENVIVETLDHESYEVKYVVSAIPPVLNLKIHFNPPLPPMRNQLISRVPMGSVIKCMVYYKEPFWRKKDYCGTMLIEDEEAVIGMTLDDTKPNCSVPDXCIVFFLVFKSLHFSHRKKRICELYAKVLGSEEALHPVHYEEKNWCEEQYSGGCYTAYFPPGIMTQFGRIIRQPVGRIYFAGTETATEWSGYMEGAVQAGERAAREILCSMGKISEGEIWKSEPESVDVPARPITTTFWERNLPSVPGLLRLVGFSTFFTSVAAVGLFAYKKGLLVRN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP84</accession>
    <name>A0A452IP84_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">3-phosphoinositide-dependent protein kinase 1</fullName>
            <ecNumber evidence="6">2.7.11.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="28 29">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="29" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="28" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="28" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="24">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="23">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cell junction</location>
            <location evidence="3">Focal adhesion</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Peripheral membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the protein kinase superfamily. AGC Ser/Thr protein kinase family. PDPK1 subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="6"/>
    <dbReference type="AlphaFoldDB" id="A0A452IP84"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033826.1">
        <property type="protein sequence ID" value="ENSGAGP00000029789.1"/>
        <property type="gene ID" value="ENSGAGG00000021510.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005925">
        <property type="term" value="C:focal adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004676">
        <property type="term" value="F:3-phosphoinositide-dependent protein kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035556">
        <property type="term" value="P:intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043066">
        <property type="term" value="P:negative regulation of apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd01262">
        <property type="entry name" value="PH_PDK1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05581">
        <property type="entry name" value="STKc_PDK1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000126">
        <property type="entry name" value="3-phosphoinositide dependent protein kinase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000163">
        <property type="entry name" value="3-phosphoinositide-dependent protein kinase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000257">
        <property type="entry name" value="3-phosphoinositide-dependent protein kinase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033931">
        <property type="entry name" value="PDK1-typ_PH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039046">
        <property type="entry name" value="PDPK1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050236">
        <property type="entry name" value="Ser_Thr_kinase_AGC"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24356:SF163">
        <property type="entry name" value="3-PHOSPHOINOSITIDE-DEPENDENT PROTEIN KINASE 1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24356">
        <property type="entry name" value="SERINE/THREONINE-PROTEIN KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14593">
        <property type="entry name" value="PH_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="18" id="KW-0007">Acetylation</keyword>
    <keyword evidence="15 25" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="17" id="KW-0965">Cell junction</keyword>
    <keyword evidence="8" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="9" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="14" id="KW-0418">Kinase</keyword>
    <keyword evidence="20" id="KW-0472">Membrane</keyword>
    <keyword evidence="13 25" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="22" id="KW-0539">Nucleus</keyword>
    <keyword evidence="11" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="29" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="21" id="KW-0804">Transcription</keyword>
    <keyword evidence="19" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="12" id="KW-0808">Transferase</keyword>
    <keyword evidence="16" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="Protein kinase" evidence="27">
        <location>
            <begin position="58"/>
            <end position="318"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="26">
        <location>
            <begin position="1"/>
            <end position="54"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="26">
        <location>
            <begin position="1"/>
            <end position="26"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="26">
        <location>
            <begin position="32"/>
            <end position="46"/>
        </location>
    </feature>
    <feature type="binding site" evidence="25">
        <location>
            <position position="87"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004202"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00010006"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00012513"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00018538"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00047899"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00048679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="27">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="28">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029789.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="29">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="528" mass="60366" checksum="AA5DDDB3C54BA92A" modified="2019-05-08" version="1">MVRNQTDSSTPSVITTCGSRKGSNMEGTAAESRSSSNSLQQHSGQQPPQPRKKRRENFKFGKILGEGSFSTVVLARELATSREYAIKILEKRHIIKENKVPYVTRERDVMSRLDHPFFVKLYFTFQDDEKLYFGLSYAKNGELLKYIRKIGSFDETCTRFYTAEIVSALEYLHGKGIIHRDLKPENILLNEEMHIQITDFGTAKVLSADSRQARANSFVGTAQYVSPELLTEKSACKSSDLWALGCIIYQLVAGLPPFRAGNEYLIFQKIIKLEYDFPEKFFPKAKDLVEKLLVLDATKRLGCEEMGGYGPLKAHPFFESITWENLHHQTPPKLTAYLPAMSEDDEDCYGNYDNLLSQFGCMQVSGSASSPSLSAPETSHPQTSGNIEQYIHDLDNNSFELDLQFSEDEKRLLLAKQAGGNPWHQFVENNLILKMGPVDKRKGLFARRRQLLLTEGPHLYYVDPVNKVLKGEIPWSLELRPEAKNFKTFFVHTPNRTYYLMDPSGNAHKWCKKIHEVWRHRYHQNAAK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP88</accession>
    <name>A0A452IP88_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Protein Wnt</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="11">Ligand for members of the frizzled family of seven transmembrane receptors.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 11">Secreted</location>
            <location evidence="1 11">Extracellular space</location>
            <location evidence="1 11">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 11">Belongs to the Wnt family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IP88"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033764.1">
        <property type="protein sequence ID" value="ENSGAGP00000029736.1"/>
        <property type="gene ID" value="ENSGAGG00000021458.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005125">
        <property type="term" value="F:cytokine activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005109">
        <property type="term" value="F:frizzled binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060070">
        <property type="term" value="P:canonical Wnt signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045165">
        <property type="term" value="P:cell fate commitment"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030182">
        <property type="term" value="P:neuron differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd19336">
        <property type="entry name" value="Wnt_Wnt4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.2460.20:FF:000001">
        <property type="entry name" value="Wnt homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.2460.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005817">
        <property type="entry name" value="Wnt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009142">
        <property type="entry name" value="Wnt4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043158">
        <property type="entry name" value="Wnt_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018161">
        <property type="entry name" value="Wnt_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12027:SF104">
        <property type="entry name" value="PROTEIN WNT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12027">
        <property type="entry name" value="WNT RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00110">
        <property type="entry name" value="wnt"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01844">
        <property type="entry name" value="WNT4PROTEIN"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01349">
        <property type="entry name" value="WNTPROTEIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00097">
        <property type="entry name" value="WNT1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00246">
        <property type="entry name" value="WNT1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 11" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="9" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
    <keyword evidence="6 11" id="KW-0879">Wnt signaling pathway</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005683"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022687"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="RuleBase" id="RU003500"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029736.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="346" mass="38390" checksum="4AE5513E911B620F" modified="2019-05-08" version="1">KRARKQQPELCLFAVAMRVAGQKRGISSLEEYLAKQPSLRGTLQDSSACEGLKGLAEEQVRICQRQVEAMDSVKHGAELAIEECQHQFHNRRWNCSTLQGLQVFGTRESAFIHAISSAGVAFAVTQACSRGDLPASWTLSLSFPGFQWSGCSDNLSYGIAFSQAFVDNPERSRGVSSSRALMNLHNNEAGRKVKLVPCVECKCHGVSGSCEVRTCWKAMPPFRKVGNVLKEKFEGATEVHPKRVGSRKLLVPKSSRFKPYTAHDLVYLVASPDFCDRDPHSGVFGTSGRQCNRTSHAMDGCELLCCGRGFRTAQAERVERCSCKFHWCCSVKCKQCHHLVEVHTCR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP89</accession>
    <name>A0A452IP89_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">1-alkyl-2-acetylglycerophosphocholine esterase</fullName>
            <ecNumber evidence="1">3.1.1.47</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="EC" id="3.1.1.47" evidence="1"/>
    <dbReference type="AlphaFoldDB" id="A0A452IP89"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033973.1">
        <property type="protein sequence ID" value="ENSGAGP00000029915.1"/>
        <property type="gene ID" value="ENSGAGG00000021591.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003847">
        <property type="term" value="F:1-alkyl-2-acetylglycerophosphocholine esterase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016042">
        <property type="term" value="P:lipid catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1820">
        <property type="entry name" value="alpha/beta hydrolase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029058">
        <property type="entry name" value="AB_hydrolase_fold"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10272">
        <property type="entry name" value="PLATELET-ACTIVATING FACTOR ACETYLHYDROLASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10272:SF6">
        <property type="entry name" value="PLATELET-ACTIVATING FACTOR ACETYLHYDROLASE 2, CYTOPLASMIC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03403">
        <property type="entry name" value="PAF-AH_p_II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53474">
        <property type="entry name" value="alpha/beta-Hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="3" id="KW-0442">Lipid degradation</keyword>
    <keyword evidence="4" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="17"/>
            <end position="74"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00013201"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022963"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029915.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="495" mass="53924" checksum="58889A9E4C2A46D1" modified="2019-05-08" version="1">MHWAALPPLFPWGGGSPGRCRRCSSGPGGPGSRPALRTDRLRPGAPPAGSAELPGAAGGVLSPAQEGMGGAQSLRLPHGNGPHQVGCTDVMVGPNRKGLFFRLFYPCLPQEEAEQPLWIPRREYCDGLADYINLKWCTPLLNVTFGSYRVPVSWNGPFKACGSRYPLIIFSHGLGAFRDHSACATYFCKAEAGGSNSLEAALQEEWIPFRRVQEGQKEFHFRNPQVHQRANECVRGLRLVQEISCGKAVTNVLHKGFDLSALKESVDLTKVAVMGHSFGGATAILALSKEDHFSCAVALDAWMFPLKNDLYHKVTKPVLFINTEKFQTAESVAKMKRLSSRNRQTKIITILGSVHLSQTDFTFLAGNLLNRLFQTRGTLDPYAGLDITNRAALAFLQKHLSTALGRKMFFIGMVQWGPLSTQTSELRLSPPWPGAADSLEEAMLAKGFLSRVGGDHSSQCVGMVFRGEPPSCPIHRAGQDSQKPRGGVVTCLVLC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP92</accession>
    <name>A0A452IP92_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Vacuolar protein sorting-associated protein 33A</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Plays a role in vesicle-mediated protein trafficking to lysosomal compartments including the endocytic membrane transport and autophagic pathways. Believed to act as a core component of the putative HOPS and CORVET endosomal tethering complexes which are proposed to be involved in the Rab5-to-Rab7 endosome conversion probably implicating MON1A/B, and via binding SNAREs and SNARE complexes to mediate tethering and docking events during SNARE-mediated membrane fusion. The HOPS complex is proposed to be recruited to Rab7 on the late endosomal membrane and to regulate late endocytic, phagocytic and autophagic traffic towards lysosomes. The CORVET complex is proposed to function as a Rab5 effector to mediate early endosome fusion probably in specific endosome subpopulations. Required for fusion of endosomes and autophagosomes with lysosomes; the function is dependent on its association with VPS16 but not VIPAS39. The function in autophagosome-lysosome fusion implicates STX17 but not UVRAG.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cytoplasmic vesicle</location>
            <location evidence="3">Autophagosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasmic vesicle</location>
            <location evidence="1">Clathrin-coated vesicle</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Early endosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Late endosome membrane</location>
            <topology evidence="4">Peripheral membrane protein</topology>
            <orientation evidence="4">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="5">Lysosome membrane</location>
            <topology evidence="5">Peripheral membrane protein</topology>
            <orientation evidence="5">Cytoplasmic side</orientation>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the STXBP/unc-18/SEC1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IP92"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033839.1">
        <property type="protein sequence ID" value="ENSGAGP00000029801.1"/>
        <property type="gene ID" value="ENSGAGG00000021422.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005776">
        <property type="term" value="C:autophagosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030136">
        <property type="term" value="C:clathrin-coated vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005769">
        <property type="term" value="C:early endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031902">
        <property type="term" value="C:late endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005765">
        <property type="term" value="C:lysosomal membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016192">
        <property type="term" value="P:vesicle-mediated transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.850:FF:000002">
        <property type="entry name" value="Vacuolar protein sorting-associated protein 33A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.830.10:FF:000006">
        <property type="entry name" value="Vacuolar protein sorting-associated protein 33A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1910:FF:000005">
        <property type="entry name" value="vacuolar protein sorting-associated protein 33A isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1910:FF:000008">
        <property type="entry name" value="vacuolar protein sorting-associated protein 33A isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.850">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1910">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.2060">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.830.10">
        <property type="entry name" value="Syntaxin Binding Protein 1, Chain A, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043154">
        <property type="entry name" value="Sec-1-like_dom1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043127">
        <property type="entry name" value="Sec-1-like_dom3a"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001619">
        <property type="entry name" value="Sec1-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027482">
        <property type="entry name" value="Sec1-like_dom2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036045">
        <property type="entry name" value="Sec1-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043155">
        <property type="entry name" value="VPS33_dom3b"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11679">
        <property type="entry name" value="VESICLE PROTEIN SORTING-ASSOCIATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00995">
        <property type="entry name" value="Sec1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56815">
        <property type="entry name" value="Sec1/munc18-like (SM) proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="8" id="KW-0967">Endosome</keyword>
    <keyword evidence="11" id="KW-0458">Lysosome</keyword>
    <keyword evidence="10" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-0653">Protein transport</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0813">Transport</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004132"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004412"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004419"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004492"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004630"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00009884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023228"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00039493"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00054535"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029801.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="539" mass="61318" checksum="431243384EDBEF7E" modified="2019-05-08" version="1">MFTLKAGRLPPADVKNIIFLVRPRLELMDIIAENVLSEDRVRSPLRDFHILFVPRRSLLCEQRLKDQGVLGSFIHREQYSLDLIPFDGDLLSMESESAFKECYLENDQTSLYQAAKGLMTLQALYGTIPQIFGKGDCARHVANMMIRMKREFPGGQNSIFPIFDTLLLLDRNVDLLSPLATQLTYEGLIDEIYGIQNTYVKLPPEKFAPKKQGEGGKDLPTEPKKLQLNSAEELYAEIRDKNFNAVGSVLSKKAKIISAAFEERHHAKTVGEIKQFVSQLPHMQAARSSLANHTSIAELIKDITTSEDFFDNLTVEQEFMSGIDTDKVNNYTEDCIAQKHPLIKILRLVCLQSVCNSGLKQKVLDHYKREILQTYGYEHILTLNNLEKAGLLKLQTSSRNNYPTIRKTMRLWMDDVNEQNPNDISYVYSGYAPLSVRLAQLLARPGWRSIEEVLKMLPGPHFEERQQLPTGLQKKRQQGENRVTLVFFLGGVTYAEIAALRFLSQMEDGGTEYVIATTKLINGTNWIKSLMEKLEPPPF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0T2</accession>
    <name>A0A452J0T2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">LisH domain and HEAT repeat-containing protein KIAA1468</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Golgi apparatus</location>
            <location evidence="2">trans-Golgi network</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Recycling endosome</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038510.1">
        <property type="protein sequence ID" value="ENSGAGP00000034010.1"/>
        <property type="gene ID" value="ENSGAGG00000023990.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055037">
        <property type="term" value="C:recycling endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005802">
        <property type="term" value="C:trans-Golgi network"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032367">
        <property type="term" value="P:intracellular cholesterol transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000080">
        <property type="entry name" value="lisH domain and HEAT repeat-containing protein KIAA1468 homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000218">
        <property type="entry name" value="lisH domain and HEAT repeat-containing protein KIAA1468 homolog isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021133">
        <property type="entry name" value="HEAT_type_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006594">
        <property type="entry name" value="LisH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040362">
        <property type="entry name" value="RELCH"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR32059">
        <property type="entry name" value="RAB11-BINDING PROTEIN RELCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR32059:SF0">
        <property type="entry name" value="RAB11-BINDING PROTEIN RELCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00667">
        <property type="entry name" value="LisH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50077">
        <property type="entry name" value="HEAT_REPEAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50896">
        <property type="entry name" value="LISH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7 11" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0967">Endosome</keyword>
    <keyword evidence="6" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="8" id="KW-0445">Lipid transport</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <feature type="repeat" description="HEAT" evidence="10">
        <location>
            <begin position="987"/>
            <end position="1023"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1"/>
            <end position="51"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="422"/>
            <end position="447"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="11">
        <location>
            <begin position="182"/>
            <end position="216"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="11">
        <location>
            <begin position="343"/>
            <end position="377"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="12">
        <location>
            <begin position="25"/>
            <end position="35"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="427"/>
            <end position="447"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004172"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004601"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00084053"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00103"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034010.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1197" mass="132706" checksum="57DBA01014DBD399" modified="2019-05-08" version="1">MAAAGGGSSVNPFLSDSEEEEPRDGGGGNDPGLGGPLSPQQHEEEEEAAVEVPGVLPGLRAAAPGEAGWAGPGAGEPPRVPVDALAAQLLRDQFLLTALELHTELLESGRELPRLRDYFSNPGNFERQSGTPPAGGAPGLWGAAGLGLLGGCELITEGGGDRAGSISTLDSLDFARYSDDGNRETDERVAVLEFELRKAKETIQALRANLTQAAENEVPLQERKNYKSSPEIQEPIKPLEKRALNFLVNEFLLKNSYKLTSITFSDESDDQDFELWDDVGLNIPKPPDLLQLYRDFGNHQVTARDVVDVAVGVEEDELEVTTPTLGSVPVFGVSQSTEQFLIVQKLEDQINALNSEKWSLTEQIRRLESEIDFLKNENFAVPVVCDLVHVPLSVPEDNGSYLDIRSSDNDSKHGKTEEISLSISAEADSKTPKDDTLNSLPSKDREKLPSCALSNKAAIHFDKPNRTLSPAFHQALLSFCRMSAESRLGSEVSRIADSEKSVMLMLGRCLPHIVPNVLLAKREELIPLILCTACLHPEPKERDQLLHILFNLIKRPDDEQRQMILTGCVAFARHVGPTRVEAELLPQCWEQINHKYPERRLLVAESCGALAPYLPKEIRSSLVLSMLQQMLMEDKADLVREAVIKSLGIIMGYIDDPDKYQQGFELLLSALGDPSERVVSATHQVFLPAYAAWTTELGNLQLHLIPTLLNKIEKLLREGEHGLDEHKLHMYLSALQSLIPSLFSLVLQNAPFTSKAKLQGEVPPIEVTRFPRPVSPLQDVATIIGSREQLAVLLQLYDYQLELEGTTGWETLLWVVNQLLPQLIEIVGKINVASTACVHEFSRFFWRLCRTFGKIFTNTKVKPQFQEILRLSEENIDSTAGNGVLTKATVPIYATGVLTCYIQEEDRKLLIGFLEDVMTMLSLSHAPLNSLKASFVELGANPAYHELLLTVLWYGVVHTSALVRCTAARMFELTLRGMSEALVDKRVAPALVTLSSDPEFSVRIATIPAFGTIMETVTQREXRVKMQLASFLEDPQYQDQHSLQTEIIKTFGRVGPNAEPRFRDEFVIPHLHKLALVNNQQSVDSKRLDIATHLFEAYSALSCCFISEDLMVNHFLPGLRCLRTDMEHLSPEHEVILSSMIKECEQKVENKTVQEPQGSMSIAASLVSEDTKTKFLNKMGQLTTSGAMLANVFQRKK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0T3</accession>
    <name>A0A452J0T3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">Myosin motor domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">Muscle contraction.</text>
    </comment>
    <comment type="subunit">
        <text evidence="14">Muscle myosin is a hexameric protein that consists of 2 heavy chain subunits (MHC), 2 alkali light chain subunits (MLC) and 2 regulatory light chain subunits (MLC-2).</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Myofibril</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 15">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038421.1">
        <property type="protein sequence ID" value="ENSGAGP00000033926.1"/>
        <property type="gene ID" value="ENSGAGG00000020688.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030016">
        <property type="term" value="C:myofibril"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032982">
        <property type="term" value="C:myosin filament"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016460">
        <property type="term" value="C:myosin II complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000146">
        <property type="term" value="F:microfilament motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000003">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000004">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000006">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000013">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000007">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000008">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.4820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.530:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000024">
        <property type="entry name" value="Myosin heavy chain, isoform J"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.720:FF:000001">
        <property type="entry name" value="Myosin heavy chain, muscle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.340">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.370">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.4820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.2420">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002928">
        <property type="entry name" value="Myosin_tail"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014751">
        <property type="entry name" value="XRCC4-like_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615">
        <property type="entry name" value="MYOSIN HEAVY CHAIN, NON-MUSCLE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615:SF69">
        <property type="entry name" value="MYOSIN-6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01576">
        <property type="entry name" value="Myosin_tail_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90257">
        <property type="entry name" value="Myosin rod fragments"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57997">
        <property type="entry name" value="Tropomyosin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12 15" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="7 15" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="8 16" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0488">Methylation</keyword>
    <keyword evidence="10 15" id="KW-0505">Motor protein</keyword>
    <keyword evidence="11" id="KW-0514">Muscle protein</keyword>
    <keyword evidence="9 15" id="KW-0518">Myosin</keyword>
    <keyword evidence="6 15" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0787">Thick filament</keyword>
    <feature type="domain" description="Myosin motor" evidence="17">
        <location>
            <begin position="73"/>
            <end position="760"/>
        </location>
    </feature>
    <feature type="region of interest" description="Actin-binding" evidence="15">
        <location>
            <begin position="637"/>
            <end position="659"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="16">
        <location>
            <begin position="822"/>
            <end position="1138"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="16">
        <location>
            <begin position="1182"/>
            <end position="1908"/>
        </location>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <begin position="166"/>
            <end position="173"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004657"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022433"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023179"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00037488"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00038612"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033926.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1922" mass="222257" checksum="5D8A926A08F1A28B" modified="2019-05-08" version="1">AKMMDMSEFGEAAPFLRKSDKEIMILQTVAFDGKTGYVFLCDSSCQQCYTCLSLQTVTVKEDDVQPMNPPKFDMIEDMAMLTHLNEASVLHNLNRRYSHWMIYTYSGLFCVTINPYKWLPVYNPEVVNAYKGKRRSEAPPHIFSIADNAYNDMLRNRENQSMLITGESGAGKTVNTKRVIQYFATVAALGEPGKKSGTLEDQIIQANPALEAFGNAKTLRNDNSSRFGKFIRIHFGTTGKLSSADIEIYLLEKSRVIFQQPGERSYHIFYQILSGKKAELQDMLLVSTNPYDYHFSSQGAVTVDNLDDGEELMATDQAMDILGFIPDEKYGAYKLTGSIMHFGNMKFKQRPREEQAETDGTESTDKAAYLMGLNSSDLAKGLLHPRVKVGNEYVTKGQSVEQVVYAVGALSKAVYDRMFKWLVIRINKTLDTRLPRQFFIGVLDIAGFEIFDYNSFEQLCINYTNEKLQQFFNHHMFVLEQEEYKKEGIEWVFIDFGLDLQACIDLIEKPLGILSILEEECMFPKATDMTFKAKLYDNHLGKSPNLQKPRIDKKRKYEAHFELLHYAGAVPYNIIGWLEKNKDPLNETVVGIFQKSANKLLASLFENYVSADSGNTEKKRKKGASFQTVSSLHKENLNKLMTNLRSTAPHFVRCIIPNETKTPGAMDAYLVLHQLRCNGVLEGIRICRKGFPNRVLYADFKQRYRILNPAAIPEDKFVDSRKASEKLLASLDIDQTQYRFGHTKVFFKAGLLGHLEEMRDHRLSKILTMIQARARGKLMRIEFQKIVERRDALLVIQWNIRAFMAVKNWPWMKLFFKIKPLLKSAETEKEMANMKEEFLKLKEALEKSEARRKELEEKQVSLVQEKNDLLLQLQAEQDTLADAEERCDLLIKSKIQMEAKVKELTERVEDEEEMNSELTSKKRKLEDECSELKKDIDDLEITLAKVEKEKHATENKVKNLTEEMASLDETISKLTKEKKSLQEAHQQALDDLQAEEDKVNTLSKTKLKLEQQVDDLEGSLEQEKKIRMDLERVKRKLEGDLKLTQESLMDLENDKLQLDEKLKKKEFEINQLNSKLEDEQTIVMQLQKKIKELQARIEELEEELEAERAARAKVEKQRSDLARELEELSERLEEAGGATAAQLEMNKKREAEFLKVRRDLEEATLHYEATAATLRKKHADSMAELGEQVDNLQRVKQKLEKEKSELKLEVDDLLSNMEQMVKGKSNAEKLCRTYEDNLNETKTKLDEMTRLMNDLTTQKGKLQTENGEFARQLEEKESLISQLSRGKTSFTQQIEELRRLLEEETKSKNALAHALQAARHDCDLLREQFEEEQEAKAELQRALSKGNAEVAQWRTKYETDAIQRTEELEEAKKKLAARLQEAEEAIEAANAKCSSLEKTKHRLQNELEDIVIDLERANSAAAALDKKQRNFDKLISDWKQKYEETQAELEASQKEARSLSTELFKLKNAYEETLDNLETVKRENKNLQEEISDLTDQISEGNKSLHEIEKIKKQVEQEKSEVQLALEEAEGALEHEESKSLRFQLEISQIKAEFERKLAEKDEEMENIRRNQQRTIDSLQSTLDSEAKSRNEAIRLKKKMEGDLNEMEIQLSHANRHASEATNKHEKSGRGWSITGDLQVQLDDSEHLNEDLKEQLAVTDRRNNLIQSELDELRALLDQTERGRKLAEQELLEATERVNLLHTQNTSLINQKKKLETDISQMQNEVEEAIQECRNAEEKAKKAITDAAMMAEELKKEQDTSAHLERMKKNMEQTIKDLQKRLDEAEQIALKGGKKQIQKLESRVRELENDLENELRRNTESQKGVRKYERRIKELTYQAEEDKKNLTRMQELIDKLQLKVKSYKHQAEEAEAQSNQYLAKYRKQQHELDDAEERAEIAESQVNKLRTKTRDIGIKKVHEEEE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0U0</accession>
    <name>A0A452J0U0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">tRNA (32-2'-O)-methyltransferase regulator THADA</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="3">Together with methyltransferase FTSJ1, methylates the 2'-O-ribose of nucleotides at position 32 of the anticodon loop of substrate tRNAs.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the THADA family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038515.1">
        <property type="protein sequence ID" value="ENSGAGP00000034015.1"/>
        <property type="gene ID" value="ENSGAGG00000024118.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030488">
        <property type="term" value="P:tRNA methylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056843">
        <property type="entry name" value="THADA-like_TPR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056842">
        <property type="entry name" value="THADA-like_TPR_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019442">
        <property type="entry name" value="THADA/TRM732_DUF2428"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051954">
        <property type="entry name" value="tRNA_methyltransferase_THADA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14387">
        <property type="entry name" value="THADA/DEATH RECEPTOR INTERACTING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14387:SF7">
        <property type="entry name" value="THYROID ADENOMA-ASSOCIATED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10350">
        <property type="entry name" value="DUF2428"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25150">
        <property type="entry name" value="TPR_Trm732"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25151">
        <property type="entry name" value="TPR_Trm732_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0819">tRNA processing</keyword>
    <feature type="domain" description="tRNA (32-2'-O)-methyltransferase regulator THADA-like TPR repeats region" evidence="6">
        <location>
            <begin position="524"/>
            <end position="681"/>
        </location>
    </feature>
    <feature type="domain" description="DUF2428" evidence="5">
        <location>
            <begin position="896"/>
            <end position="1196"/>
        </location>
    </feature>
    <feature type="domain" description="tRNA (32-2'-O)-methyltransferase regulator THADA-like C-terminal TPR repeats region" evidence="7">
        <location>
            <begin position="1198"/>
            <end position="1361"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00010409"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022694"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00035625"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00035698"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="Pfam" id="PF10350"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="Pfam" id="PF25150"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF25151"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034015.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1738" mass="195432" checksum="7F0786E521E4CCBC" modified="2019-05-08" version="1">MVLKKKKEIQVDAFVLECQELETLRSFADAEDQNLASLLLHCVQLSDGIHQIYSMKQIVTLLKKMDRNSACSPMIKCCLDIIGVMYFSLGAKNPLKKMLASSLNGLPEQLMTEAVHSFVRCLKEELKTTDLCLYRKVMDNLASCMENFSLGMIICNIIFHIFNLFSCDLELGIVYSVKLNLSGNSIVETRLMHDLLMAIKVSMMLLQKVQENFQGSLWRKPGSLLWQSMCGLLKSFTNFLMNEGLLPTVQTTSGLAVILFTKTMFEPVEKLPCLVSDLLLRSVKCASVPEWFVSSCECLCTEELSDSVLLSLCHGALAMLEWKNGSMSENGEKLLLDIALVLLSLASRLKESNMAVSLSRILAIWTNSALDVLESSSQNLKSSLNGNSNTTGKLLEYVYTHWEHPLDAVRHQAKLIFKNILRIHQATLKGSVVKSDPFFSMLTESLLSLEWHVKGKYASLGCLVECVGVENILTVDRTIPAQILDVMGDQSLAPYASDLLETMFINHKSHLISAFQKNIWINHWHDTWVSPLLLILCEGNPDQTTYVIDYYLPKLLKCSPDSLSYMIKILQASADANIGSCNTRGALGALMACLRTARAHGHLQFREIMRDDLLSIGYVKQGLVHQHNQVRVDALGLLCETHRSTEILTMEEMQLIQFFITYNLNSQSPAVRQQICSLLKKDFMSSICDKLFEALFPGSSHPTRFSSLTILGSIAEIFSAPEGEVFDFHSGIDPTRVQMLIQCFASTFEEVKVLAFELLMKLYDRLDLQDLENLHLLFQAAMDLSISTKPYDCVTASYLLNFLIHHKGLLNVCLKNQQVADTFQIDEDMSASTMEKNTLAVIKYLLSNLEREIFQAEKSLLQAAASFPMYGRVHCITGALQQLPLNNLTLMTEWRQMVTKLILMSYRLSSVVSPIVQSSSPEGLIPMDNDSETAGRLQMILHEIQPRDTNDYFTQAKIFKEHHRVDFAKLTDDEPVENVCTEIRGKEGQTCDVTAQMVLVCCWRSMKEISLLLGMLCKLLPLQTVSESSDGLIAVEQVKDIGDYFKHHLLQSRHRGAFELAYTGFVMLTEILSRCTNESLRTLPQQWLYNVLEEIKSCDPSSKLCATRRSAGIPFYIQALLASEPKNKTNLLKMTMNELVSLAVPVNESLNIIPQVHALNILRALFRDTRFGENIIPYIADGIQAAVLGFTSPVWAVRNSSTLLFSTLITRIFGVKRGKDENSKKNRMTGREFFSRFPTLYPFLLQQLEVVARTMDSETGELKLHPSLFLLLLILGKLYPSPMDGTYSALSMAPFVPFIMRCGHSPVYRSREMAGRALVPFVMISQIPHTVQSLLAGLPECTNPCIQQNSVHGMLLQVFHLLGSYLESKHRVNSDLQQGLVDIITCIKAKVWLAKRQNPCLVTRAAYLDVLVVLSDYLGKSRRKELFGFWEEISKVISASELMTGIYYFSAIPGFTQYLQSITKLVISVLSVAADPDFSNSSSPVAEQMMNSSSMVVHLLQSDFHEVRLPMLEAVLLWWKQVNSKQATEEKMGPLSLLSDLEVILLRMAMKEKHPECFCKVGAIHFSQDLKYDGKMYCIFMMCSLTYFYSIKIQSIALKLASKLVTHLVQNCQEVMESEMRQWIQLVTSCCGDEEQTAIRLAAAEVLINITPLFLTNQKLILGLSSTLAMWKCILLLLQSEELIVRDTAADVIRVAQGPQDPDSLSLTTKENKPFPFPLLPPGVPSLGSWRDIQIQTP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0U1</accession>
    <name>A0A452J0U1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">C3H1-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the ZC3H12 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0U1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038474.1">
        <property type="protein sequence ID" value="ENSGAGP00000033974.1"/>
        <property type="gene ID" value="ENSGAGG00000024154.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036464">
        <property type="term" value="C:cytoplasmic ribonucleoprotein granule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016787">
        <property type="term" value="F:hydrolase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003729">
        <property type="term" value="F:mRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004521">
        <property type="term" value="F:RNA endonuclease activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd18729">
        <property type="entry name" value="PIN_Zc3h12-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.11980:FF:000001">
        <property type="entry name" value="ZC3H12A isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.11980">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040546">
        <property type="entry name" value="Rege-1_UBA-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021869">
        <property type="entry name" value="RNase_Zc3h12_NYN"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051101">
        <property type="entry name" value="ZC3H12/N4BP1_RNase_Reg"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000571">
        <property type="entry name" value="Znf_CCCH"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12876">
        <property type="entry name" value="N4BP1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12876:SF36">
        <property type="entry name" value="RIBONUCLEASE ZC3H12C-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11977">
        <property type="entry name" value="RNase_Zc3h12a"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18039">
        <property type="entry name" value="UBA_6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50103">
        <property type="entry name" value="ZF_C3H1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0255">Endonuclease</keyword>
    <keyword evidence="7" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="9" id="KW-0460">Magnesium</keyword>
    <keyword evidence="4 10" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="3" id="KW-0540">Nuclease</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8 10" id="KW-0862">Zinc</keyword>
    <keyword evidence="6 10" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C3H1-type" evidence="12">
        <location>
            <begin position="413"/>
            <end position="438"/>
        </location>
    </feature>
    <feature type="zinc finger region" description="C3H1-type" evidence="10">
        <location>
            <begin position="413"/>
            <end position="438"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="66"/>
            <end position="103"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="457"/>
            <end position="504"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="527"/>
            <end position="549"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="66"/>
            <end position="82"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="534"/>
            <end position="545"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010922"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022759"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50103"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033974.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="972" mass="109841" checksum="7587DA88EC2805F1" modified="2019-05-08" version="1">MLLAFKEYGTLYIQEYRKNSKVESNTCSSFMGLKDHLGHDLGHLYVERTDTQINAVGSWSMVEKPTMDKVNSRKEDADKEASEETGSSSCDSEESTNSDNESERLSNIIAESHLLPKTHRQLCRSPCLQPHVLKRNEILQDLKIEEAQIGSKEVKKPPDVVKEYQTKLEFALKLGYSEEQIQLVLNKLGTDALINDILGELVKLGNKTETDQPVNNVNTNIMHEASSIESQRSGSPLQEDVTEDGDNLRPIVIDGSNVAMSHGNKEVFSCRGIKLAVDWFLERHHKDVTVFVPAWRKEQSRPDALITDQEILRKLEKEKILVFTPSRRVQGRRVVCYDDRFIVKLAFESDGIIVSNDNYRDLANEKPEWKKFIDERLLMYSFVNDKFMPPDDPLGRHGPSLDNFLRKKPIVPEHKKQPCPYGKKCTYGHKCKYYHPERGNQPQRSVADELRAMSRSTAAKAASEGGLVKSNSVPCSTKIDGTSDLKRAAPKRQSDPSIRTQVYQDLEEKLPTKNKLETRSVPSLVSLPNSSVVKPQSTAPLSNGLPSGVHFPPQDQRPQGQYPPMMMATKNHGTPMPYEQYPKCESPVDVGYYSMLNAYSNLSISGPRSPERRFSLDTDYRISSVASDCSSEGSMSCGSSDSYVGYNDRSYMSSPDPQLEENLKCQHVHPHSRLNSQPFLQSYHDPLTRVQSYSHEEPKHHHKPPTPYMSVHLQHPTVGARSSCPNDYSALQSLQHSKTVHLGRALVSTRIDSISDSRLYDNSPLRQRKPYSSQDGLGSWDRQSYGIDAYGYRQTYSLPSNPTQPCYEQFAFQSLPEQQDQTWCIPYCGIPQDPPSEQLVGMAASICESSTQALRSKSSIWTDAKWIQSRMENFSGQIEARTDHLLWGMERRKKHNCSSRCVAQVRSILGVCVPMCRVRDFGLSSICRVEYGALWSATPMHWHIRHNRPAPSQFLLAGNSDRGAGGRVMEWT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0U4</accession>
    <name>A0A452J0U4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">LisH domain and HEAT repeat-containing protein KIAA1468</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Golgi apparatus</location>
            <location evidence="2">trans-Golgi network</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Recycling endosome</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038488.1">
        <property type="protein sequence ID" value="ENSGAGP00000033988.1"/>
        <property type="gene ID" value="ENSGAGG00000023990.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055037">
        <property type="term" value="C:recycling endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005802">
        <property type="term" value="C:trans-Golgi network"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032367">
        <property type="term" value="P:intracellular cholesterol transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000080">
        <property type="entry name" value="lisH domain and HEAT repeat-containing protein KIAA1468 homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000218">
        <property type="entry name" value="lisH domain and HEAT repeat-containing protein KIAA1468 homolog isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021133">
        <property type="entry name" value="HEAT_type_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006594">
        <property type="entry name" value="LisH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040362">
        <property type="entry name" value="RELCH"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR32059">
        <property type="entry name" value="RAB11-BINDING PROTEIN RELCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR32059:SF0">
        <property type="entry name" value="RAB11-BINDING PROTEIN RELCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00667">
        <property type="entry name" value="LisH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50077">
        <property type="entry name" value="HEAT_REPEAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50896">
        <property type="entry name" value="LISH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7 11" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0967">Endosome</keyword>
    <keyword evidence="6" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="8" id="KW-0445">Lipid transport</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <feature type="repeat" description="HEAT" evidence="10">
        <location>
            <begin position="947"/>
            <end position="983"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1"/>
            <end position="71"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="130"/>
            <end position="154"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="172"/>
            <end position="202"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="383"/>
            <end position="407"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="11">
        <location>
            <begin position="303"/>
            <end position="337"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="12">
        <location>
            <begin position="46"/>
            <end position="56"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="132"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="172"/>
            <end position="196"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="387"/>
            <end position="407"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004172"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004601"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00084053"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00103"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033988.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1157" mass="128638" checksum="78677D6A8A087B3A" modified="2019-05-08" version="1">AARPALVPALRGWRGAATAGAGGVAGPASSVNPFLSDSEEEEPRDGGGGNDPGLGGPLSPQQHEEEEEAAVEVPGVLPGLRAAAPGEPPRVPVDALAAQLLRDQFLLTALELHTELLESGRELPRLRDYFSNPGNFERQSGTPPAGGAPGLWGAADRAGSISTLDSLDFARYSDDGNRETDERVAENEVPLQERKNYKSSPEIQEPIKPLEKRALNFLVNEFLLKNSYKLTSITFSDESDDQDFELWDDVGLNIPKPPDLLQLYRDFGNHQVTARDVVDVAVGVEEDELEVTTPTLGSQFLIVQKLEDQINALNSEKWSLTEQIRRLESEIDFLKNENFAVPVVCDLVHVPLSVPEDNGSYLDIRSSDNDSKHGKTEEISLSISAEADSKTPKDDTLNSLPSKDREKLPSCALSNKAAIHFDKPNRTLSPAFHQALLSFCRMSAESRLGSEVSRIADSEKSVMLMLGRCLPHIVPNVLLAKREELIPLILCTACLHPEPKERDQLLHILFNLIKRPDDEQRQMILTGCVAFARHVGPTRVEAELLPQCWEQINHKYPERRLLVAESCGALAPYLPKEIRSSLVLSMLQQMLMEDKADLVREAVIKSLGIIMGYIDDPDKYQQGFELLLSALGDPSERVVSATHQVFLPAYAAWTTELGNLQLHLIPTLLNKIEKLLREGEHGLDEHKLHMYLSALQSLIPSLFSLVLQNAPFTSKAKLQGEVPPIEVTRFPRPVSPLQDVATIIGSREQLAVLLQLYDYQLELEGTTGWETLLWVVNQLLPQLIEIVGKINVASTACVHEFSRFFWRLCRTFGKIFTNTKVKPQFQEILRLSEENIDSTAGNGVLTKATVPIYATGVLTCYIQEEDRKLLIGFLEDVMTMLSLSHAPLNSLKASFVELGANPAYHELLLTVLWYGVVHTSALVRCTAARMFELLVKGVHETLVAQRVVPALITYSSDPEISVRIATIPAFGTIMETVTQREXRVKMQLASFLEDPQYQDQHSLQTEIIKTFGRVGPNAEPRFRDEFVIPHLHKLALVNNQQSVDSKRLDIATHLFEAYSALSCCFISEDLMVNHFLPGLRCLRTDMEHLSPEHEVILSSMIKECEQKVENKTVQEPQGSMSIAASLVSEDTKTKFLNKMGQLTTSGAMLANVFQRKK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0U7</accession>
    <name>A0A452J0U7_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="18">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
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    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
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        <scope>IDENTIFICATION</scope>
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    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
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    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
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    <comment type="similarity">
        <text evidence="2 12">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
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        <property type="component" value="Unassembled WGS sequence"/>
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        <property type="term" value="C:cytoplasm"/>
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        <property type="project" value="UniProtKB-SubCell"/>
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        <property type="project" value="UniProtKB-UniRule"/>
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        <ligand>
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        <source>
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        <source>
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        </source>
    </evidence>
    <sequence length="2127" mass="244969" checksum="7FC26C6520084C82" modified="2019-05-08" version="1">MDLKTGREFDVPIGAVVKLCDSGQIQVVDDEGNEHWIFPHNASHIKPMHPTSIHGVEDMIRLGDLNEAGILRNLLIRYREHLIYTYTGSILVAVNPYQLLPIYSPEQIRLYTNKKIGEMPPHIFAIADNCYYNMQRNNRDQCCIISGESGAGKTESTKLILQFLAAISGQHSWIEQQVLEANPILEAFGNAKTIRNDNSSRFGKYIDIHFNKRGAIEGAKIEQYLLEKSRVCRQAHDERNYHVFYCMLKGMTPEQKKKLGLGKATDYNYLAMGNCTTCDGRDDSKEYANIRSAMKVLMFTDTENWEISKLLAAILHMGNLRYEARMYDNLDACEVVRSASLITAASLLEVDPQDVMNCLTSRTIITRGETVSTPLSMEQALDVRDAFVKGIYGRLFVWIVEKINAAIYRPPSHELQSVRRSIGLLDIFGFENFTVNSFEQLCINFANENLQQFFVRHVFKLEQEEYNLENINWQHIEFTDNQDALDMIAIKPMNIISLIDEESKFPKGTDTTMLHKLNSQHKLNTNYIPPKNNHETQFGINHFAGVVYYETKGFLEKNRDTLHGDIIQLVHSSKNKFIKQIFQADVAMFLCGYASSTFGQSPAPTPKGAETRKRSPTLSSQFKRSLELLMRTLSGCQPFFVRCIKPNEYKKPMLFDRELCVRQLRYSGMMETIRIRRAGYPIRYSFVEFVDRYRVLMPGVKPAYKQGDLRGTCQRIAEAVLGKDDDWQIGKTKIFLKDHHDMLLEIERDKAITDKVILIQKVVRGFKNRSNFLKVRHSALMIQRYWRGHNCRKNYGAMRTGFLRLQALCRSRKLHTQYHMARRRIIEFQARCRGCLVRRAFRHRLWAVFTIQAYARGMIARRLYKRLKGEYHRRLEAEKLRLAEEERLRKEMSAKKAKEEAEKKHQVRLAQLAREDAEREVKEKEEARRKKELLDKMEKARKEPVNDSDMVDKMFGFLGTTSSLPGQEGQAPNGFEDLERAQKELEEEDLDAALPLPEEEEEDLSEYKFAKFAATYFQGTTTHTYIRRPLKQPLLYHEDEGDQLSAHIDGHKKNSVRHKLVSLTLKKKSKLTEEVTKRLHDGESTLQGNSMLEDRPTSNLEKLHFIIGNGILRPALRDEIYCQICKQLTQNPSKSSHARGWILMSLCVGCFAPSEKFVKYLRNFINGGPPGYAPYCEERLRRTFANGTRTQPPSWLELQATKSKKPIMLPVTFMDGTTKTLLTDSATTAKELCNSLADKISLKDRFGFSLYIALFDKVSSLGSGNDHVMDAVSQCEQYAKEQGAQERNAPWRLFFRKEIFTPWHNPSEDSVATNLIYQQIVRGVKFGEYRCDKEEDLAELASQQYYVDYGATMVLERLLNLIPSYIPDREITASKTVEKWAQLIIAAHKKGIYTQKRADPKKVKEEVVDFARFKWPLLFSRFYEAFKFSGPSLPKNDVIVAVNWTGVYFVDEQEQVLLELSFPEITAVSSSRGGKLQGQSFTLATIKGDEYTFTSNNAEDIRDLVVTFLEGLRKRSKYVVTLQDNPNPVGEESGFLSFHKGDLIVMDQDTGEHVMNSGWANGVNERTKQRGDFPTDCVYVLPTVTMPPLEIVALVTMTPDQRQDVIRMSQLAISDSEERVKPYTLEEFSYDYFRPPPKHTLSRVMITKNRGKDKLWCYTREPIKQPLLKKILGSEELSQEACMAFIAVLKYMGDYPSKRTRSVNELTDQIFEGALKAEPLKDEIYCQILKQLTDNHIKYSEEKGWELLWLCTGLFPPSNVLLPHVQRFLQSRKHHPLATDCIQRLQKALRNGSRKYPPHLVEVEAIQHKTTQIFHKVYFPDDTDEAFEVESSTKAKDFCQNISSRLLLKSSEGFSLFVKISDKVISVPEGDFFFDFVRHLTDWIKKARPAKDGIVPSLTYQVFFMKKLWTNTIPGKDSMADSIFHYYQELPKYLRGYHKCTREEVLQLAALIYRVKFEDDKSYFPSIPKLLKELVPQDLLRQLSPDDWKRSIVAYFNKHAGKTREEAKLAFLKIVFKWPTFGSAFFEVKQTTEPNYPEILLIAINKHGVSLIDPKTKDILTTHPFTKISNWSSGNTYFHITIGNLVRGSKLLCETSLGYKMDDLLTSYISQMLTAMSKQRSAKGSSK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0U8</accession>
    <name>A0A452J0U8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">LisH domain and HEAT repeat-containing protein KIAA1468</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Golgi apparatus</location>
            <location evidence="2">trans-Golgi network</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Recycling endosome</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038520.1">
        <property type="protein sequence ID" value="ENSGAGP00000034020.1"/>
        <property type="gene ID" value="ENSGAGG00000023990.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055037">
        <property type="term" value="C:recycling endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005802">
        <property type="term" value="C:trans-Golgi network"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032367">
        <property type="term" value="P:intracellular cholesterol transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000080">
        <property type="entry name" value="lisH domain and HEAT repeat-containing protein KIAA1468 homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000218">
        <property type="entry name" value="lisH domain and HEAT repeat-containing protein KIAA1468 homolog isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021133">
        <property type="entry name" value="HEAT_type_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006594">
        <property type="entry name" value="LisH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040362">
        <property type="entry name" value="RELCH"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR32059">
        <property type="entry name" value="RAB11-BINDING PROTEIN RELCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR32059:SF0">
        <property type="entry name" value="RAB11-BINDING PROTEIN RELCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00667">
        <property type="entry name" value="LisH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50077">
        <property type="entry name" value="HEAT_REPEAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50896">
        <property type="entry name" value="LISH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7 11" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0967">Endosome</keyword>
    <keyword evidence="6" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="8" id="KW-0445">Lipid transport</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <feature type="repeat" description="HEAT" evidence="10">
        <location>
            <begin position="977"/>
            <end position="1013"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1"/>
            <end position="76"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="119"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="412"/>
            <end position="437"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="11">
        <location>
            <begin position="175"/>
            <end position="209"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="11">
        <location>
            <begin position="333"/>
            <end position="367"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="12">
        <location>
            <begin position="25"/>
            <end position="35"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="130"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="417"/>
            <end position="437"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004172"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004601"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00084053"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00103"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034020.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1187" mass="131732" checksum="3EC81CCC1761A2F3" modified="2019-05-08" version="1">MAAAGGGSSVNPFLSDSEEEEPRDGGGGNDPGLGGPLSPQQHEEEEEAAVEVPGVLPGLRAAAPGEAGWAGPGAGEPPRVPVDALAAQLLRDQFLLTALELHTELLESGRELPRLRDYFSNPGNFERQSGTPPAGGAPGLWGAAGAAPGGAAANRAGSISTLDSLDFARYSDDGNRETDERVAVLEFELRKAKETIQALRANLTQAAENEVPLQERKNYKSSPEIQEPIKPLEKRALNFLVNEFLLKNSYKLTSITFSDESDDQDFELWDDVGLNIPKPPDLLQLYRDFGNHQVTARDVVDVAVGVEEDELEVTTPTLGSVPTLLCRFQFLIVQKLEDQINALNSEKWSLTEQIRRLESEIDFLKNENFAVPVVCDLVHVPLSVPEDNGSYLDIRSSDNDSKHGKTEEISLSISAEADSKTPKDDTLNSLPSKDREKLPSCALSNKAAIHFDKPNRTLSPAFHQALLSFCRMSAESRLGSEVSRIADSEKSVMLMLGRCLPHIVPNVLLAKREELIPLILCTACLHPEPKERDQLLHILFNLIKRPDDEQRQMILTGCVAFARHVGPTRVEAELLPQCWEQINHKYPERRLLVAESCGALAPYLPKEIRSSLVLSMLQQMLMEDKADLVREAVIKSLGIIMGYIDDPDKYQQGFELLLSALGDPSERVVSATHQVFLPAYAAWTTELGNLQLHLIPTLLNKIEKLLREGEHGLDEHKLHMYLSALQSLIPSLFSLVLQNAPFTSKAKLQGEVPPIEVTRFPRPVSPLQDVATIIGSREQLAVLLQLYDYQLELEGTTGWETLLWVVNQLLPQLIEIVGKINVASTACVHEFSRFFWRLCRTFGKIFTNTKVKPQFQEILRLSEENIDSTAGNGVLTKATVPIYATGVLTCYIQEEDRKLLIGFLEDVMTMLSLSHAPLNSLKASFVELGANPAYHELLLTVLWYGVVHTSALVRCTAARMFELLVKGVHETLVAQRVVPALITYSSDPEISVRIATIPAFGTIMETVTQREXRVKMQLASFLEDPQYQDQHSLQTEIIKTFGRVGPNAEPRFRDEFVIPHLHKLALVNNQQSVDSKRLDIATHLFEAYSALSCCFISEDLMVNHFLPGLRCLRTDMEHLSPEHEVILSSMIKECEQKVENKTVQEPQGSMSIAASLVSEDTKTKFLNKMGQLTTSGAMLANVFQRKK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0U9</accession>
    <name>A0A452J0U9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">E3 ubiquitin-protein ligase RNF130</fullName>
            <ecNumber evidence="5">2.3.2.27</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="20">Goliath homolog</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="19">RING finger protein 130</fullName>
        </alternativeName>
    </protein>
    <organism evidence="25 26">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="26" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="25" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="25" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>S-ubiquitinyl-[E2 ubiquitin-conjugating enzyme]-L-cysteine + [acceptor protein]-L-lysine = [E2 ubiquitin-conjugating enzyme]-L-cysteine + N(6)-ubiquitinyl-[acceptor protein]-L-lysine.</text>
            <dbReference type="EC" id="2.3.2.27"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="4">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Membrane</location>
            <topology evidence="2">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="2.3.2.27" evidence="5"/>
    <dbReference type="AlphaFoldDB" id="A0A452J0U9"/>
    <dbReference type="STRING" id="38772.ENSGAGP00000033943"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038440.1">
        <property type="protein sequence ID" value="ENSGAGP00000033943.1"/>
        <property type="gene ID" value="ENSGAGG00000024136.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061630">
        <property type="term" value="F:ubiquitin protein ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006915">
        <property type="term" value="P:apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045893">
        <property type="term" value="P:positive regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016567">
        <property type="term" value="P:protein ubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006511">
        <property type="term" value="P:ubiquitin-dependent protein catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd02122">
        <property type="entry name" value="PA_GRAIL_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.40.10:FF:000009">
        <property type="entry name" value="E3 ubiquitin-protein ligase RNF130"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.50.30.30:FF:000011">
        <property type="entry name" value="E3 ubiquitin-protein ligase RNF130"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.50.30.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046450">
        <property type="entry name" value="PA_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003137">
        <property type="entry name" value="PA_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051834">
        <property type="entry name" value="RING_finger_E3_ligase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001841">
        <property type="entry name" value="Znf_RING"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45931:SF21">
        <property type="entry name" value="RING FINGER PROTEIN 130"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45931">
        <property type="entry name" value="SI:CH211-59O9.10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02225">
        <property type="entry name" value="PA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13639">
        <property type="entry name" value="zf-RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00184">
        <property type="entry name" value="RING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52025">
        <property type="entry name" value="PA domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57850">
        <property type="entry name" value="RING/U-box"/>
        <property type="match status" value="1"/>
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    <dbReference type="PROSITE" id="PS50089">
        <property type="entry name" value="ZF_RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="9" id="KW-0053">Apoptosis</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="17" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="16 22" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="26" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="11 23" id="KW-0732">Signal</keyword>
    <keyword evidence="7" id="KW-0808">Transferase</keyword>
    <keyword evidence="8 22" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="15 22" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="13" id="KW-0833">Ubl conjugation pathway</keyword>
    <keyword evidence="14" id="KW-0862">Zinc</keyword>
    <keyword evidence="12 21" id="KW-0863">Zinc-finger</keyword>
    <feature type="signal peptide" evidence="23">
        <location>
            <begin position="1"/>
            <end position="32"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019349368" description="E3 ubiquitin-protein ligase RNF130" evidence="23">
        <location>
            <begin position="33"/>
            <end position="424"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="22">
        <location>
            <begin position="200"/>
            <end position="223"/>
        </location>
    </feature>
    <feature type="domain" description="RING-type" evidence="24">
        <location>
            <begin position="269"/>
            <end position="310"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022703"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00067476"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00080460"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00082289"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS50089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033943.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="424" mass="47307" checksum="C085E89C0619D15B" modified="2019-05-08" version="1">MMLAKRTCQCWKRLSSLPVLVLVNCCVLLASADNTSQAYYTALINVTVLNPDRVSPTLLRIDRGRYGQDSPKVEVKGLLLAPVPINGVVDRLGCDPRTRFHVPPNTKQWIALLQRGNCTFREKILRAASHNATAVVIYNNISSEEPVTMTHQGTGDIVAVMITESKVKEILNYLEKNISVLMAIAVGARVPPKNFSRGSLVFVSISFIVLMIISSAWLIFYFIQKIRYTNARDRNQRRLGDAAKKAISKLTTRTVKKGDKETDPDFDHCAVCIECYKQNDVVRILPCKHVFHKACVDPWLSEHCTCPMCKLNILKALGIVPSVPCTDNVAFDMERLTRGQPANRRSALGDFANDNSLSLEPLRTSGMSQLPQDGEITPRTGEINIAVTSGHFFHHNSLSPRSLVYESEFSTIQPALDMYDENKT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0V2</accession>
    <name>A0A452J0V2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Myosin VIIA</fullName>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 12">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038485.1">
        <property type="protein sequence ID" value="ENSGAGP00000033985.1"/>
        <property type="gene ID" value="ENSGAGG00000024151.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045177">
        <property type="term" value="C:apical part of cell"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005902">
        <property type="term" value="C:microvillus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016459">
        <property type="term" value="C:myosin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003779">
        <property type="term" value="F:actin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003774">
        <property type="term" value="F:cytoskeletal motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009887">
        <property type="term" value="P:animal organ morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030182">
        <property type="term" value="P:neuron differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007423">
        <property type="term" value="P:sensory organ development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007605">
        <property type="term" value="P:sensory perception of sound"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd17092">
        <property type="entry name" value="FERM1_F1_Myosin-VII"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd17093">
        <property type="entry name" value="FERM2_F1_Myosin-VII"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd14473">
        <property type="entry name" value="FERM_B-lobe"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="CDD" id="cd13198">
        <property type="entry name" value="FERM_C1_MyoVII"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd13199">
        <property type="entry name" value="FERM_C2_MyoVII"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd01381">
        <property type="entry name" value="MYSc_Myo7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.80.10:FF:000012">
        <property type="entry name" value="Myosin VIIA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.530:FF:000004">
        <property type="entry name" value="Myosin VIIA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000007">
        <property type="entry name" value="Myosin VIIA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.80.10:FF:000013">
        <property type="entry name" value="Unconventional myosin-VIIa"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000075">
        <property type="entry name" value="unconventional myosin-VIIa"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000079">
        <property type="entry name" value="unconventional myosin-VIIa"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.40:FF:000113">
        <property type="entry name" value="unconventional myosin-VIIa"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.20.90:FF:000036">
        <property type="entry name" value="Unconventional myosin-VIIa"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.20.90:FF:000051">
        <property type="entry name" value="Unconventional myosin-VIIa"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.190">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.80.10">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.20.240.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.530">
        <property type="entry name" value="MyTH4 domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.20.90">
        <property type="entry name" value="Phosphatidylinositol 3-kinase Catalytic Subunit, Chain A, domain 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019749">
        <property type="entry name" value="Band_41_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014352">
        <property type="entry name" value="FERM/acyl-CoA-bd_prot_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035963">
        <property type="entry name" value="FERM_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019748">
        <property type="entry name" value="FERM_central"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000299">
        <property type="entry name" value="FERM_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000048">
        <property type="entry name" value="IQ_motif_EF-hand-BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002404">
        <property type="entry name" value="IRS_PTB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057130">
        <property type="entry name" value="Myosin_VII_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041793">
        <property type="entry name" value="MyoVII_FERM_C1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041794">
        <property type="entry name" value="MyoVII_FERM_C2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036106">
        <property type="entry name" value="MYSc_Myo7"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000857">
        <property type="entry name" value="MyTH4_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038185">
        <property type="entry name" value="MyTH4_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029071">
        <property type="entry name" value="Ubiquitin-like_domsf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051567">
        <property type="entry name" value="Unconventional_Myosin_ATPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22692">
        <property type="entry name" value="MYOSIN VII, XV"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22692:SF34">
        <property type="entry name" value="MYOSIN VIIA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21998">
        <property type="entry name" value="FERM_C1_MyoVII"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00373">
        <property type="entry name" value="FERM_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00612">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02174">
        <property type="entry name" value="IRS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24123">
        <property type="entry name" value="Myosin_VII_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00784">
        <property type="entry name" value="MyTH4"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21989">
        <property type="entry name" value="RA_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00295">
        <property type="entry name" value="B41"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00015">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00139">
        <property type="entry name" value="MyTH4"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00326">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47031">
        <property type="entry name" value="Second domain of FERM"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54236">
        <property type="entry name" value="Ubiquitin-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50057">
        <property type="entry name" value="FERM_3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51016">
        <property type="entry name" value="MYTH4"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10 12" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="7 12" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="13" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9 12" id="KW-0505">Motor protein</keyword>
    <keyword evidence="8 12" id="KW-0518">Myosin</keyword>
    <keyword evidence="6 12" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="3 11" id="KW-0728">SH3 domain</keyword>
    <feature type="domain" description="Myosin motor" evidence="17">
        <location>
            <begin position="54"/>
            <end position="730"/>
        </location>
    </feature>
    <feature type="domain" description="MyTH4" evidence="16">
        <location>
            <begin position="1006"/>
            <end position="1242"/>
        </location>
    </feature>
    <feature type="domain" description="FERM" evidence="15">
        <location>
            <begin position="1247"/>
            <end position="1556"/>
        </location>
    </feature>
    <feature type="domain" description="SH3" evidence="14">
        <location>
            <begin position="1554"/>
            <end position="1623"/>
        </location>
    </feature>
    <feature type="domain" description="MyTH4" evidence="16">
        <location>
            <begin position="1698"/>
            <end position="1847"/>
        </location>
    </feature>
    <feature type="domain" description="FERM" evidence="15">
        <location>
            <begin position="1853"/>
            <end position="2156"/>
        </location>
    </feature>
    <feature type="region of interest" description="Actin-binding" evidence="12">
        <location>
            <begin position="607"/>
            <end position="629"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="13">
        <location>
            <begin position="856"/>
            <end position="924"/>
        </location>
    </feature>
    <feature type="binding site" evidence="12">
        <location>
            <begin position="147"/>
            <end position="154"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50057"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS51016"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033985.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="20">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="2167" mass="249638" checksum="9AE143E0181DF2F9" modified="2019-05-08" version="1">MDLKTGREFDVPIGAVVKLCDSGQIQVVDDEGNEHWIFPHNASHIKPMHPTSIHGVEDMIRLGDLNEAGILRNLLIRYREHLIYTYTGSILVAVNPYQLLPIYSPEQIRLYTNKKIGEMPPHIFAIADNCYYNMQRNNRDQCCIISGESGAGKTESTKLILQFLAAISGQHSWIEQQVLEANPILEAFGNAKTIRNDNSSRFGKYIDIHFNKRGAIEGAKIEQYLLEKSRVCRQAHDERNYHVFYCMLKGMTPEQKKKLGLGKATDYNYLAMGNCTTCDGRDDSKEYANIRSAMKVLMFTDTENWEISKLLAAILHMGNLRYEARMYDNLDACEVVRSASLITAASLLEVDPQDVMNCLTSRTIITRGETVSTPLSMEQALDVRDAFVKGIYGRLFVWIVEKINAAIYRPPSHELQSVRRSIGLLDIFGFENFTVNSFEQLCINFANENLQQFFVRHVFKLEQEEYNLENINWQHIEFTDNQDALDMIAIKPMNIISLIDEESKFPKGTDTTMLHKLNSQHKLNTNYIPPKNNHETQFGINHFAGVVYYETKGFLEKNRDTLHGDIIQLVHSSKNKFIKQIFQADVAMGAETRKRSPTLSSQFKRSLELLMRTLSGCQPFFVRCIKPNEYKKPMLFDRELCVRQLRYSGMMETIRIRRAGYPIRYSFVEFVDRYRVLMPGVKPAYKQGDLRGTCQRIAEAVLGKDDDWQIGKTKIFLKDHHDMLLEIERDKAITDKVILIQKVVRGFKNRSNFLKVRHSALMIQRYWRGHNCRKNYGAMRTGFLRLQALCRSRKLHTQYHMARRRIIEFQARCRGCLVRRAFRHRLWAVFTIQAYARGMIARRLYKRLKGEYHRRLEAEKLRLAEEERLRKEMSAKKAKEEAEKKHQVRLAQLAREDAEREVKEKEEARRKKELLDKMEKARKEPVNDSDMVDKMFGFLGTTSSLPGQEGQAPNGFEDLERAQKELEEEDLDAALPLPEEEEEDLSEYKFAKFAATYFQGTTTHTYIRRPLKQPLLYHEDEGDQLAALAVWITILRFMGDLPEPKYHTAMSDGSEKIPVMTKIYETLGKKTYKKELQALQGEGESAHIDGHKKNSVRHKLVSLTLKKKSKLTEEVTKRLHDGESTLQGNSMLEDRPTSNLEKLHFIIGNGILRPALRDEIYCQICKQLTQNPSKSSHARGWILMSLCVGCFAPSEKFVKYLRNFINGGPPGYAPYCEERLRRTFANGTRTQPPSWLELQATKSKKPIMLPVTFMDGTTKTLLTDSATTAKELCNSLADKISLKDRFGFSLYIALFDKVSSLGSGNDHVMDAVSQCEQYAKEQGAQERNAPWRLFFRKEIFTPWHNPSEDSVATNLIYQQIVRGVKFGEYRCDKEEDLAELASQQYYVDYGATMVLERLLNLIPSYIPDREITASKTVEKWAQLIIAAHKKGIYTQKRADPKKVKEEVVDFARFKWPLLFSRFYEAFKFSGPSLPKNDVIVAVNWTGVYFVDEQEQVLLELSFPEITAVSSSRGGKLQGQSFTLATIKGDEYTFTSNNAEDIRDLVVTFLEGLRKRSKYVVTLQDNPNPVGEESGFLSFHKGDLIVMDQDTGEHVMNSGWANGVNERTKQRGDFPTDCVYVLPTVTMPPLEIVALVTMTPDQRQDVIRMSQLAISDSEERVKPYTLEEFSYDYFRPPPKHTLSRVMITKNRGKDKLWCYTREPIKQPLLKKILGSEELSQEACMAFIAVLKYMGDYPSKRTRSVNELTDQIFEGALKAEPLKDEIYCQILKQLTDNHIKYSEEKGWELLWLCTGLFPPSNVLLPHVQRFLQSRKHHPLATDCIQRLQKALRNGSRKYPPHLVEVEAIQHKTTQIFHKVYFPDDTDEAFEVESSTKAKDFCQNISSRLLLKSSEGFSLFVKISDKVISVPEGDFFFDFVRHLTDWIKKARPAKDGIVPSLTYQVFFMKKLWTNTIPGKDSMADSIFHYYQELPKYLRGYHKCTREEVLQLAALIYRVKFEDDKSYFPSIPKLLKELVPQDLLRQLSPDDWKRSIVAYFNKHAGKTREEAKLAFLKIVFKWPTFGSAFFEVKQTTEPNYPEILLIAINKHGVSLIDPKTKDILTTHPFTKISNWSSGNTYFHITIGNLVRGSKLLCETSLGYKMDDLLTSYISQMLTAMSKQRSAKGSSK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0V3</accession>
    <name>A0A452J0V3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">tRNA (32-2'-O)-methyltransferase regulator THADA</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="3">Together with methyltransferase FTSJ1, methylates the 2'-O-ribose of nucleotides at position 32 of the anticodon loop of substrate tRNAs.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the THADA family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038506.1">
        <property type="protein sequence ID" value="ENSGAGP00000034006.1"/>
        <property type="gene ID" value="ENSGAGG00000024118.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030488">
        <property type="term" value="P:tRNA methylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056843">
        <property type="entry name" value="THADA-like_TPR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056842">
        <property type="entry name" value="THADA-like_TPR_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019442">
        <property type="entry name" value="THADA/TRM732_DUF2428"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051954">
        <property type="entry name" value="tRNA_methyltransferase_THADA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14387">
        <property type="entry name" value="THADA/DEATH RECEPTOR INTERACTING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14387:SF7">
        <property type="entry name" value="THYROID ADENOMA-ASSOCIATED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10350">
        <property type="entry name" value="DUF2428"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25150">
        <property type="entry name" value="TPR_Trm732"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25151">
        <property type="entry name" value="TPR_Trm732_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0819">tRNA processing</keyword>
    <feature type="domain" description="tRNA (32-2'-O)-methyltransferase regulator THADA-like TPR repeats region" evidence="6">
        <location>
            <begin position="533"/>
            <end position="805"/>
        </location>
    </feature>
    <feature type="domain" description="DUF2428" evidence="5">
        <location>
            <begin position="949"/>
            <end position="1249"/>
        </location>
    </feature>
    <feature type="domain" description="tRNA (32-2'-O)-methyltransferase regulator THADA-like C-terminal TPR repeats region" evidence="7">
        <location>
            <begin position="1251"/>
            <end position="1414"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00010409"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022694"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00035625"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00035698"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="Pfam" id="PF10350"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="Pfam" id="PF25150"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF25151"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034006.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1807" mass="203612" checksum="1192B6CF66AE342A" modified="2019-05-08" version="1">MVLKKKKEIQVDAFVLECQELETLRSFADAEDQNLASLLLHCVQLSDGIHQIYSMKQIVTLLKKMDRNSACSPMIKCCLDIIGVMYFSLGAKNPLKKMLASSLNGLPEQLMTEAVHSFVRCLKEELKTTDLCLYRKVMDNLASCMENFSLGMIICNIIFHIFNLFSCDLELVLQFLQKSVLEIQEENRQLSGNSIVETRLMHDLLMAIKVSMMLLQKVQENFQGSLWRKPGSLLWQSMCGLLKSFTNFLMNEGLLPTVQTTSGLAVILFTKTMFEPVEKLPCLVSDLLLRSVKCASVPEWFVSSCECLCTEELSDSVLLSLCHGALAMLEWKNGSMSENGEKLLLDIALVLLSLASRLKESNMAVSLSRILAIWTNSALDVLESSSQNLKSSLNGNSNTTGKLLEYVYTHWEHPLDAVRHQAKLIFKNILRIHQATLKGSVVKSDPFFSMLTESLLSLEWHVKGKYASLGCLVECVGVENILTVDRTIPAQILDVMGDQSLAPYASDLLETMFINHKSHLISAFQKNIWINHWHDTWVSPLLLILCEGNPDQTTYVIDYYLPKLLKCSPDSLSYMIKILQASADANIGSCNTRGALGALMACLRTARAHGHLQFREIMRDDLLSIGYVKQGLVHQHNQVRVDALGLLCETHRSTEILTMEEMQLIQFFITYNLNSQSPAVRQQICSLLKKLFCRIQESSQVLYKLEQNKTKQELIKESAKWDHFATLQQYKDFMSSICDKLFEALFPGSSHPTRFSSLTILGSIAEIFSAPEGQTQAIFRLAQEIDPTRVQMLIQCFASTFEEVKVLAFELLMKLYDRLDLQDLENLHLLFQAAMDLSISTKPYDCVTASYLLNFLIHHKGLLNVCLKNQQVADTFQIDEDMSASTMEKNTLAVIKYLLSNLEREIFQAEKSLLQAAASFPMYGRVHCITGALQQLPLNNLTLMTEWRQMVTKLILMSYRLSSVVSPIVQSSSPEGLIPMDNDSETAGRLQMILHEIQPRDTNDYFTQAKIFKEHHRVDFAKLTDDEPVENVCTEIRGKEGQTCDVTAQMVLVCCWRSMKEISLLLGMLCKLLPLQTVSESSDGLIAVEQVKDIGDYFKHHLLQSRHRGAFELAYTGFVMLTEILSRCTNESLRTLPQQWLYNVLEEIKSCDPSSKLCATRRSAGIPFYIQALLASEPKNKTNLLKMTMNELVSLAVPVNESLNIIPQVHALNILRALFRDTRFGENIIPYIADGIQAAVLGFTSPVWAVRNSSTLLFSTLITRIFGVKRGKDENSKKNRMTGREFFSRFPTLYPFLLQQLEVVARTMDSETGELKLHPSLFLLLLILGKLYPSPMDGTYSALSMAPFVPFIMRCGHSPVYRSREMAGRALVPFVMISQIPHTVQSLLAGLPECTNPCIQQNSVHGMLLQVFHLLGSYLESKHRVNSDLQQGLVDIITCIKAKVWLAKRQNPCLVTRAAYLDVLVVLSDYLGKSRRKGIELFGFWEEISKVISASELMTGIYYFSAIPGFTQYLQSITKLVISVLSVAADPDFSNSSSPVAEQMMNSSSMVVHLLQSDFHEVRLPMLEAVLLWWKQVNSKQATEEKMGPLSLLSDLEVILLRMAMKEKHPECFCKVLEILHCMDLRNVLPRTQDSIKMNPKEFLNWAMNIISASDSIKIQSIALKLASKLVMESEMRQWIQLVTSCCGDEEQTAIRLAAAEVLINITPLFLTNQKLILGLSSTLAMWKCILLLLQSEELIVRDTAADVIRVAQSQSNICNKTGTVWGPQDPDSLSLTTKENKPFPFPLLPPGVPSLGSWRDIQIQTP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0V4</accession>
    <name>A0A452J0V4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Rho guanine nucleotide exchange factor 4</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038525.1">
        <property type="protein sequence ID" value="ENSGAGP00000034025.1"/>
        <property type="gene ID" value="ENSGAGG00000024175.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035556">
        <property type="term" value="P:intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
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    <dbReference type="CDD" id="cd01224">
        <property type="entry name" value="PH_Collybistin_ASEF"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="RhoGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="SH3_ASEF"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Rho guanine nucleotide exchange factor 9"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Rho guanine nucleotide exchange factor 9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.900.10">
        <property type="entry name" value="Dbl homology (DH) domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="DBL_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000219">
        <property type="entry name" value="DH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001331">
        <property type="entry name" value="GDS_CDC24_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055251">
        <property type="entry name" value="SOS1_NGEF_PH"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47544">
        <property type="entry name" value="RHO GUANINE NUCLEOTIDE EXCHANGE FACTOR 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47544:SF2">
        <property type="entry name" value="RHO GUANINE NUCLEOTIDE EXCHANGE FACTOR 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00621">
        <property type="entry name" value="RhoGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00018">
        <property type="entry name" value="SH3_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22697">
        <property type="entry name" value="SOS1_NGEF_PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00233">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00325">
        <property type="entry name" value="RhoGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00326">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48065">
        <property type="entry name" value="DBL homology domain (DH-domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="1"/>
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    <dbReference type="PROSITE" id="PS00741">
        <property type="entry name" value="DH_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50010">
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0344">Guanine-nucleotide releasing factor</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 5" id="KW-0728">SH3 domain</keyword>
    <feature type="domain" description="SH3" evidence="7">
        <location>
            <begin position="1466"/>
            <end position="1525"/>
        </location>
    </feature>
    <feature type="domain" description="DH" evidence="9">
        <location>
            <begin position="1562"/>
            <end position="1746"/>
        </location>
    </feature>
    <feature type="domain" description="PH" evidence="8">
        <location>
            <begin position="1777"/>
            <end position="1884"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="36"/>
            <end position="66"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="175"/>
            <end position="197"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="442"/>
            <end position="470"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="932"/>
            <end position="967"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="1087"/>
            <end position="1110"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="1131"/>
            <end position="1172"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="1251"/>
            <end position="1293"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="47"/>
            <end position="63"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="185"/>
            <end position="196"/>
        </location>
    </feature>
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            <end position="457"/>
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    </feature>
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            <begin position="932"/>
            <end position="941"/>
        </location>
    </feature>
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            <begin position="958"/>
            <end position="967"/>
        </location>
    </feature>
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            <end position="1160"/>
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        <location>
            <begin position="1251"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
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        </source>
    </evidence>
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        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
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        </source>
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            <dbReference type="PROSITE" id="PS50010"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034025.1"/>
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    </evidence>
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        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="12">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1927" mass="214677" checksum="E8FB9A710DA8FF9D" modified="2019-05-08" version="1">MSQLELSQKAQLELSSDLKVAPSRSEFWIQTTKLRREEAGEEELILSSDPSNTGSGTSGGDTISKTETKLVVDGLQSEAKSENNKVQSCVSSEPAEDAIARSKKSPAGLSIACLKANSESCLTFLETQQFPLQPNVGSSHSLHNINFETNRTDSKRNSEAFEFHYELDSRSNIKCSRRNSDNNEETTATKTRNRTPGATRKMDSLLYQPCLKSKFQLPHMRAEIKESKTWHSAPENINAGGQRDYLGNCIQLRDKLRLSCSRIHLEGLGFEYTWKNIERVGEGSEKAIKVEGKFGIKGSLLNTCLESNNSQLSHPAGSQQDAEANLHTESETCGLNIEADMCADGLSSAQLQLRDTFEWDSIERETPSALASEPMSINAEGEVRKAKCKSERGFKAHGFSHSINSESGISEDQATCLVGVPAIAENKGDNYLANYGPAGKTQLNSLEESSPESPSTEACVGERGSEDSSELNLDWKSLETALNSGGQDIAGEQDVSMLQNLNCKNNALQGHSVLVTMNGHFADAGKKDESDGVDACSVEIATKPHYMHQGKKLMKLSELKPALIIISVEEKGLQSKMLSDDCPAEIITGTLGELANEDLGSHPIASKGTDESQGDLNAYSSCEEQTLPVQPISGNEEVTAAGITGQEESEIVQQQVERTEPDLSLKLSCSDFPVTTIKNESSGVNLHLEDAIVPLKEKADACTGAHLKSPGIISGGISQVEPCRTNAGVQSETELLKKSLFETRDSKVTKENEFNTNQSLEQDTCGTIATALEKNRISGLCQGDDADGHTFGSIISCVHSLEKKKTESYNISKDQIGIGEDSSSRELKLSSTVEELVSLSKDNEQNPNVTDAFNCIVKTESQEASALGLDNISTSSANSEEMDENSFHFLGSNSSLYKAVQKQTKTGKPSKFLVFSKMASFKKTKTAAAENQGSSNFFGSKTKTEELDAGKEDEDTESLQSFKSSSPSSAFRRKESYTSEYSDDDDLFYERPAGLFSRISLRKASGSGRILMDNIDTNFTSPSLHVKHSEARVLESVENNDSEVPQYFGVRKSLSENEYKGNKNPEGKKFRTRLALAHKSFSSFFESKSLEKENPEQSPKGSLKHEKEKAKLRQSSWKAFLKSREADGLKKPTLASLSSSQESPKTTRGYQQGPGNVTRRTSTEKQESSNGQAFFSSYESNATLMDANHSDTSVESIEVXNMRRKRVLSYDLTIKCPEYPDYAAQQETMVNSLDTSEEYWLRSPFSPSDLQSPFTQLSPSCPQLSTYEGKDMPCRPMSPKPQSPRPSSQRKSFRYPGRISATSMISLGNSSAIYSNLEAPERPKTLKPRGTLLLSVHSLDNEYLREDSGISSQSQISLNTASSVSDILRDEARMSLSITSPVAFEILPLKLHPFSQSTPTGLDCVGWKRHISFPVITDGALDKAALADDVGSEEDLYEDFRSSNHRYGHPGGGGEQLAINELISDGSVVYAEALWDHVTMDDQELGFKAGDVIEVMDATNKEWWWGRILDSEGWFPASFVRLRVNQDEPMEDYPMKAEDGKEEDASIAARRYGVGQISKDQMRTNVINEIISTERDYIKHLKDICEGYIKQCRKRADMFTEEQLKTIFGNIEDIYKCQKKFVKALEKKFNKDHPHLSEVGSCFLEYQTEFQIYSEYCNNHPNACMELSRLTKVNKYVYFFEACRLLQKMIDISLDGFLLTPVQKICKYPLQLAELLKYTNPQHRDFKDVEAALNAMKNVARLINERKRRLENIDKIAQWQSSIEDWEGEDVLVRSSELIYSGELTKISQPQAKSQQRMFFLFDHQLVCCKKDLLRRDILYYKSRINMDDMEIVDMEDGKDKDFNISVKNAFKLLCRDPEEVHLFCAKKPEQKQRWLKAFENERKQVQLDQETGFSITEVQKKQAMLNANKQHHTGKPKGECGPFFFF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0V5</accession>
    <name>A0A452J0V5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">E3 ubiquitin-protein ligase RNF130</fullName>
            <ecNumber evidence="5">2.3.2.27</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="20">Goliath homolog</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="19">RING finger protein 130</fullName>
        </alternativeName>
    </protein>
    <organism evidence="25 26">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="26" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="25" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="25" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>S-ubiquitinyl-[E2 ubiquitin-conjugating enzyme]-L-cysteine + [acceptor protein]-L-lysine = [E2 ubiquitin-conjugating enzyme]-L-cysteine + N(6)-ubiquitinyl-[acceptor protein]-L-lysine.</text>
            <dbReference type="EC" id="2.3.2.27"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="4">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Membrane</location>
            <topology evidence="2">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="2.3.2.27" evidence="5"/>
    <dbReference type="AlphaFoldDB" id="A0A452J0V5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038445.1">
        <property type="protein sequence ID" value="ENSGAGP00000033948.1"/>
        <property type="gene ID" value="ENSGAGG00000024136.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061630">
        <property type="term" value="F:ubiquitin protein ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006915">
        <property type="term" value="P:apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045893">
        <property type="term" value="P:positive regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016567">
        <property type="term" value="P:protein ubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006511">
        <property type="term" value="P:ubiquitin-dependent protein catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd02122">
        <property type="entry name" value="PA_GRAIL_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.40.10:FF:000009">
        <property type="entry name" value="E3 ubiquitin-protein ligase RNF130"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.50.30.30:FF:000011">
        <property type="entry name" value="E3 ubiquitin-protein ligase RNF130"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.50.30.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046450">
        <property type="entry name" value="PA_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003137">
        <property type="entry name" value="PA_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051834">
        <property type="entry name" value="RING_finger_E3_ligase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001841">
        <property type="entry name" value="Znf_RING"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45931:SF21">
        <property type="entry name" value="RING FINGER PROTEIN 130"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45931">
        <property type="entry name" value="SI:CH211-59O9.10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02225">
        <property type="entry name" value="PA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13639">
        <property type="entry name" value="zf-RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00184">
        <property type="entry name" value="RING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52025">
        <property type="entry name" value="PA domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57850">
        <property type="entry name" value="RING/U-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50089">
        <property type="entry name" value="ZF_RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="9" id="KW-0053">Apoptosis</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="17" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="16 22" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="26" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="11 23" id="KW-0732">Signal</keyword>
    <keyword evidence="7" id="KW-0808">Transferase</keyword>
    <keyword evidence="8 22" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="15 22" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="13" id="KW-0833">Ubl conjugation pathway</keyword>
    <keyword evidence="14" id="KW-0862">Zinc</keyword>
    <keyword evidence="12 21" id="KW-0863">Zinc-finger</keyword>
    <feature type="signal peptide" evidence="23">
        <location>
            <begin position="1"/>
            <end position="32"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019275161" description="E3 ubiquitin-protein ligase RNF130" evidence="23">
        <location>
            <begin position="33"/>
            <end position="420"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="22">
        <location>
            <begin position="200"/>
            <end position="223"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="22">
        <location>
            <begin position="391"/>
            <end position="410"/>
        </location>
    </feature>
    <feature type="domain" description="RING-type" evidence="24">
        <location>
            <begin position="269"/>
            <end position="310"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022703"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00067476"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00080460"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00082289"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS50089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033948.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="420" mass="46655" checksum="740DB76D11599774" modified="2019-05-08" version="1">MMLAKRTCQCWKRLSSLPVLVLVNCCVLLASADNTSQAYYTALINVTVLNPDRVSPTLLRIDRGRYGQDSPKVEVKGLLLAPVPINGVVDRLGCDPRTRFHVPPNTKQWIALLQRGNCTFREKILRAASHNATAVVIYNNISSEEPVTMTHQGTGDIVAVMITESKVKEILNYLEKNISVLMAIAVGARVPPKNFSRGSLVFVSISFIVLMIISSAWLIFYFIQKIRYTNARDRNQRRLGDAAKKAISKLTTRTVKKGDKETDPDFDHCAVCIECYKQNDVVRILPCKHVFHKACVDPWLSEHCTCPMCKLNILKALGIVPSVPCTDNVAFDMERLTRGQPANRRSALGDFANDNSLSLEPLRTSGMSQLPQDGEITPRTGEINIAVTKEWFIIASFGLLSALTLCYMIIKATASLNADE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0V9</accession>
    <name>A0A452J0V9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">tRNA (32-2'-O)-methyltransferase regulator THADA</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="3">Together with methyltransferase FTSJ1, methylates the 2'-O-ribose of nucleotides at position 32 of the anticodon loop of substrate tRNAs.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the THADA family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038490.1">
        <property type="protein sequence ID" value="ENSGAGP00000033990.1"/>
        <property type="gene ID" value="ENSGAGG00000024118.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030488">
        <property type="term" value="P:tRNA methylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056843">
        <property type="entry name" value="THADA-like_TPR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056842">
        <property type="entry name" value="THADA-like_TPR_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019442">
        <property type="entry name" value="THADA/TRM732_DUF2428"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051954">
        <property type="entry name" value="tRNA_methyltransferase_THADA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14387">
        <property type="entry name" value="THADA/DEATH RECEPTOR INTERACTING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14387:SF7">
        <property type="entry name" value="THYROID ADENOMA-ASSOCIATED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10350">
        <property type="entry name" value="DUF2428"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25150">
        <property type="entry name" value="TPR_Trm732"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25151">
        <property type="entry name" value="TPR_Trm732_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0819">tRNA processing</keyword>
    <feature type="domain" description="tRNA (32-2'-O)-methyltransferase regulator THADA-like TPR repeats region" evidence="6">
        <location>
            <begin position="561"/>
            <end position="833"/>
        </location>
    </feature>
    <feature type="domain" description="DUF2428" evidence="5">
        <location>
            <begin position="977"/>
            <end position="1277"/>
        </location>
    </feature>
    <feature type="domain" description="tRNA (32-2'-O)-methyltransferase regulator THADA-like C-terminal TPR repeats region" evidence="7">
        <location>
            <begin position="1279"/>
            <end position="1442"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00010409"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022694"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00035625"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00035698"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="Pfam" id="PF10350"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="Pfam" id="PF25150"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF25151"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033990.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1758" mass="198000" checksum="527887A8CD7C72B8" modified="2019-05-08" version="1">MIEPRRGAEAVWVGGQKAAAEAGCRASVPLGLHFNHTKMVLKKKKEIQVDAFVLECQELETLRSFADAEDQNLASLLLHCVQLSDGIHQIYSMKQIVTLLKKMDRNSACSPMIKCCLDIIGVMYFSLGAKNPLKKMLASSLNGLPEQLMTEAVHSFVRCLKEELKTTDLCLYRKVMDNLASCMENFSLGKASINNLFKEVLQFLQKSVLEIQEENRKLSGNSIVETRLMHDLLMAIKVSMMLLQKVQENFQGSLWRKPGSLLWQSMCGLLKSFTNFLMNEGLLPTVQTTSGLAVILFTKTMFEPVEKLPCLVSDLLLRSVKCASVPEWFVSSCECLCTEELSDSVLLSLCHGALAMLEWKNGSMSENGEKLLLDIALVLLSLASRLKESNMAVSLSRILAIWTNSALDVLESSSQNLKSSLNGNSNTTGKLLEYVYTHWEHPLDAVRHQAKLIFKNILRIHQATLKGSVVKSDPFFSMLTESLLSLEWHVKGKYASLGCLVECVGVENILTVDRTIPAQILDVMGDQSLAPYASDLLETMFINHKSHLISAFQKNIWINHWHDTWVSPLLLILCEGNPDQTTYVIDYYLPKLLKCSPDSLSYMIKILQASADANIGSCNTRGALGALMACLRTARAHGHLQFREIMRDDLLSIGYVKQGLVHQHNQVRVDALGLLCETHRSTEILTMEEMQLIQFFITYNLNSQSPAVRQQICSLLKKLFCRIQESSQVLYKLEQNKTKQELIKESAKWDHFATLQQYKDFMSSICDKLFEALFPGSSHPTRFSSLTILGSIAEIFSAPEGQTQAIFRLAQEIDPTRVQMLIQCFASTFEEVKVLAFELLMKLYDRLDLQDLENLHLLFQAAMDLSISTKPYDCVTASYLLNFLIHHKGLLNVCLKNQQVADTFQIDEDMSASTMEKNTLAVIKYLLSNLEREIFQAEKSLLQAAASFPMYGRVHCITGALQQLPLNNLTLMTEWRQMVTKLILMSYRLSSVVSPIVQSSSPEGLIPMDNDSETAGRLQMILHEIQPRDTNDYFTQAKIFKEHHRVDFAKLTDDEPVENVCTEIRGKEGQTCDVTAQMVLVCCWRSMKEISLLLGMLCKLLPLQTVSESSDGLIAVEQVKDIGDYFKHHLLQSRHRGAFELAYTGFVMLTEILSRCTNESLRTLPQQWLYNVLEEIKSCDPSSKLCATRRSAGIPFYIQALLASEPKNKTNLLKMTMNELVSLAVPVNESLNIIPQVHALNILRALFRDTRFGENIIPYIADGIQAAVLGFTSPVWAVRNSSTLLFSTLITRIFGVKRGKDENSKKNRMTGREFFSRFPTLYPFLLQQLEVVARTMDSETGELKLHPSLFLLLLILGKLYPSPMDGTYSALSMAPFVPFIMRCGHSPVYRSREMAGRALVPFVMISQIPHTVQSLLAGLPECTNPCIQQNSVHGMLLQVFHLLGSYLESKHRVNSDLQQGLVDIITCIKAKVWLAKRQNPCLVTRAAYLDVLVVLSDYLGKSRRKGIELFGFWEEISKVISASELMTGIYYFSAIPGFTQYLQSITKLVISVLSVAADPDFSNSSSPVAEQMMNSSSMVVHLLQSDFHEVRLPMLEAVLLWWKQVNSKQATEEKMGPLSLLSDLEVILLRMAMKEKHPECFCKVLEILHCMDLRNVLPRTQDSIKMNPKEFLNWAMNIISASDSIKIQSIALKLASKLVTHLVQNCQEVMESEMRQWIQLVTSCCGDEEQTAIRLAAAEVLINITPLFLTNQKLILGE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0W0</accession>
    <name>A0A452J0W0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Myosin motor domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1 10">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038516.1">
        <property type="protein sequence ID" value="ENSGAGP00000034016.1"/>
        <property type="gene ID" value="ENSGAGG00000024156.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016459">
        <property type="term" value="C:myosin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000146">
        <property type="term" value="F:microfilament motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007015">
        <property type="term" value="P:actin filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd23767">
        <property type="entry name" value="IQCD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd01380">
        <property type="entry name" value="MYSc_Myo5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.530:FF:000002">
        <property type="entry name" value="Class V myosin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000089">
        <property type="entry name" value="Myosin VC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.190:FF:000001">
        <property type="entry name" value="unconventional myosin-Va"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.1590:FF:000005">
        <property type="entry name" value="unconventional myosin-Vc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.190">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.1590">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.220.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000048">
        <property type="entry name" value="IQ_motif_EF-hand-BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036103">
        <property type="entry name" value="MYSc_Myo5"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13140">
        <property type="entry name" value="MYOSIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13140:SF313">
        <property type="entry name" value="UNCONVENTIONAL MYOSIN-VC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00612">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00015">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9 10" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="4 10" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="5" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="6 11" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="8 10" id="KW-0505">Motor protein</keyword>
    <keyword evidence="7 10" id="KW-0518">Myosin</keyword>
    <keyword evidence="3 10" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="Myosin motor" evidence="12">
        <location>
            <begin position="62"/>
            <end position="748"/>
        </location>
    </feature>
    <feature type="region of interest" description="Actin-binding" evidence="10">
        <location>
            <begin position="627"/>
            <end position="649"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="11">
        <location>
            <begin position="879"/>
            <end position="1093"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="11">
        <location>
            <begin position="1195"/>
            <end position="1222"/>
        </location>
    </feature>
    <feature type="binding site" evidence="10">
        <location>
            <begin position="156"/>
            <end position="163"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00008314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034016.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1298" mass="151909" checksum="584B9C6E2AD252D7" modified="2019-05-08" version="1">MSICELQVWIPDNEEVWQSAEITKSYRTGDRVLHLLLEDGTELNYPVDPAPLPPLRNPDILVGENDLTALSYLHEPAVLHNLKVRFVESKLIYTYSGIILVAMNPYKQLPIYGDAIIHAYSGQNMGDMDPHIFAVAEEAYKQMARNNRNQSVIVSGESGAGKTVSARYAMRYFATVSKSNSNVHVEDKVLASNPITEAVGNAKTTRNDNSSRFGKYTEISFDQRHQITGANMRTYLLEKSRVVFQSENERNYHIFYQLCASALQPEFKHLKLGHAEEFNYTRMGGNTVIEGVDDRADMMETQKTFTLLGLKEDFQMDVFKILAAILHLGNVQMTAVGDEKSSVSLEDKHLNIFCELLDLDNDKVAQWLCHRKIITTSETVIKPMTKLQALNARDALAKKIYSHLFDFIVERINKALHFSGKQHTFIGVLDIYGFETFDVNSFEQFCINYANEKLQQQFNLHVFKLEQEEYMKENIPWTLIDFYDNQPVIDLIEAKMGILELLDEECLLPQGTDENWLQKLYNNYINKNPFFQKPKMSNTSFIIQHFADKVEYKSEGFLEKNRDTVYEVLIDILRNSKFHLCANFFRDRPLPISPYRSTITVKSAKTVVKPPNKQFRTTVGSEFRSSLYLLMETLNATTPHYVRCIKPNDEKLPFEFDSKRVVQQLRACGVLETIRISAQSYPSRWTYIEFFSRYSILMTQHELSLSDKKKICKTVLQRLIKDPNQYQFGRTKIFFRAGQVAYLEKLRSDKLRQACIVIQKNIRSWHQRKKFLRIKQAAVTIQQYFRGQRAVRQAITATALKQAWAAIVIQKYCRGYIVRKLCQLIHVAAITIQAYVRGFLARKKFRKMLKEHKAVILQKYARAWLARRRFQNIRRFVLNIQLSYRVQRLQKKLEDQNKENHGLLERLTHLASTHANDLDTIQKLELELEKLATQKKTSEEKGKKYKEDTEQKLSKLEKQNAELQEQKTFLEQKLQEETKEMKGTIRKQIKVIKMLEKKFQNEKQDYEKETESLKGEIKTLKEEKIKLQGQIEEEHIIQDDLKLEVGQLKKQAKMIPELQKEIELLQTQKMDVEKQVQSQKREMREKVSEVAEQFIESYDVEAVRSRLSIEDLEHLNEDGELWFAYEGLKKATRVLESQFQSQKENYEKEIEGLNFKVEHLSQEIIHLQKLFREENDINDSIRLEVSRLTSENMMIPDFKQQISELQKQKMDLEQHLQEQAAKTKGKEPMNKLFVISRTLEEQTETHTKEEKQIVTIQEIQETNNHLKEQHEIENEVKSKIKQKAFHLTLENMVRNEII</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0W2</accession>
    <name>A0A452J0W2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Secretory carrier-associated membrane protein</fullName>
            <shortName evidence="10">Secretory carrier membrane protein</shortName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Probably involved in membrane protein trafficking.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 10">Membrane</location>
            <topology evidence="1 10">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 10">Belongs to the SCAMP family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0W2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038507.1">
        <property type="protein sequence ID" value="ENSGAGP00000034007.1"/>
        <property type="gene ID" value="ENSGAGG00000024172.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055038">
        <property type="term" value="C:recycling endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032588">
        <property type="term" value="C:trans-Golgi network membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007273">
        <property type="entry name" value="SCAMP"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10687:SF11">
        <property type="entry name" value="SECRETORY CARRIER-ASSOCIATED MEMBRANE PROTEIN 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10687">
        <property type="entry name" value="SECRETORY CARRIER-ASSOCIATED MEMBRANE PROTEIN SCAMP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04144">
        <property type="entry name" value="SCAMP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 10" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="6" id="KW-0653">Protein transport</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 10" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 10" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="3 10" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="43"/>
            <end position="63"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="69"/>
            <end position="90"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="102"/>
            <end position="125"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="145"/>
            <end position="170"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="185"/>
            <end position="229"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="185"/>
            <end position="194"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="203"/>
            <end position="215"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010482"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00037350"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="RuleBase" id="RU363122"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034007.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="229" mass="26105" checksum="CB218ACE5A912E07" modified="2019-05-08" version="1">MAEKVNNFPPLPKFIPLKPCFYQNFADEIPIDYQSLVKRIYHLWIFYCVTLVVNLIGCLAWWIGGGYGVNFGLAILWLFLFSPCSYICWFRPAYKGFRSDSSFNFMAFFFIFGAQFILTVIQAIGISNWGACGWLAATTFFSTSVAAAVFMLFPAIMFTMSAVAMSFYLLRVHKIYRGTGGSFQKAQDEWNSGSWRDPPSREAQYNNFSGNSLPQYPTVPNYPTGNQWP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0W3</accession>
    <name>A0A452J0W3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Unconventional myosin-Va</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="14">Dilute myosin heavy chain, non-muscle</fullName>
        </alternativeName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="12">Processive actin-based motor that can move in large steps approximating the 36-nm pseudo-repeat of the actin filament. Can hydrolyze ATP in the presence of actin, which is essential for its function as a motor protein. Involved in melanosome transport. Also mediates the transport of vesicles to the plasma membrane. May also be required for some polarization process involved in dendrite formation.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>ATP + H2O = ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:13065"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="11">
            <dbReference type="Rhea" id="RHEA:13066"/>
        </physiologicalReaction>
    </comment>
    <comment type="similarity">
        <text evidence="1 15">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038539.1">
        <property type="protein sequence ID" value="ENSGAGP00000034035.1"/>
        <property type="gene ID" value="ENSGAGG00000024176.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016459">
        <property type="term" value="C:myosin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000146">
        <property type="term" value="F:microfilament motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007015">
        <property type="term" value="P:actin filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd01380">
        <property type="entry name" value="MYSc_Myo5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.530:FF:000002">
        <property type="entry name" value="Class V myosin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.190:FF:000006">
        <property type="entry name" value="Myosin VA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000089">
        <property type="entry name" value="Myosin VC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.1590:FF:000003">
        <property type="entry name" value="Myosin-Va isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.190:FF:000001">
        <property type="entry name" value="unconventional myosin-Va"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.190">
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.1590">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002710">
        <property type="entry name" value="Dilute_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000048">
        <property type="entry name" value="IQ_motif_EF-hand-BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058662">
        <property type="entry name" value="Myo5a/b_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036103">
        <property type="entry name" value="MYSc_Myo5"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004009">
        <property type="entry name" value="SH3_Myosin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13140">
        <property type="entry name" value="MYOSIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13140:SF273">
        <property type="entry name" value="UNCONVENTIONAL MYOSIN-VA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01843">
        <property type="entry name" value="DIL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00612">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25966">
        <property type="entry name" value="Myo5a"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM01132">
        <property type="entry name" value="DIL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00015">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51126">
        <property type="entry name" value="DILUTE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51844">
        <property type="entry name" value="SH3_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10 15" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="5 15" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="6" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="7 16" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="9 15" id="KW-0505">Motor protein</keyword>
    <keyword evidence="8 15" id="KW-0518">Myosin</keyword>
    <keyword evidence="4 15" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="Myosin N-terminal SH3-like" evidence="20">
        <location>
            <begin position="8"/>
            <end position="60"/>
        </location>
    </feature>
    <feature type="domain" description="Myosin motor" evidence="19">
        <location>
            <begin position="69"/>
            <end position="764"/>
        </location>
    </feature>
    <feature type="domain" description="Dilute" evidence="18">
        <location>
            <begin position="1506"/>
            <end position="1782"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="602"/>
            <end position="635"/>
        </location>
    </feature>
    <feature type="region of interest" description="Actin-binding" evidence="15">
        <location>
            <begin position="644"/>
            <end position="666"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1107"/>
            <end position="1148"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1170"/>
            <end position="1203"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="16">
        <location>
            <begin position="916"/>
            <end position="1099"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="16">
        <location>
            <begin position="1322"/>
            <end position="1411"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="602"/>
            <end position="614"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="1117"/>
            <end position="1131"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="17">
        <location>
            <begin position="1137"/>
            <end position="1148"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="17">
        <location>
            <begin position="1170"/>
            <end position="1197"/>
        </location>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <begin position="163"/>
            <end position="170"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00008314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00048778"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00054098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00068036"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00076270"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS51126"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS51844"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034035.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1827" mass="212495" checksum="158A56D0D4D3FAA6" modified="2019-05-08" version="1">MAVSELYTKYARVWIPDPEEVWKSAELLKDYKQGDKVIQLRLEEGKELEYGLDPKTKELPPLRNPDILVGENDLTALSYLHEPAVLHNLKVRFIDSKLIYTYCGIVLVAINPYEQLPIYGEDIINAYSGQNMGDMDPHIFAVAEEAYKQMARDERNQSIIVSGESGAGKTVSAKYAMRYFATVSGSASEANVEEKVLASNPIMESIGNAKTTRNDNSSRFGKYIEIGFDKRYRIIGANMRTYLLEKSRVVFQAEEERNYHIFYQLCASAALPEFKTLRLGNANYFHYTKQGGSPVIDGIDDAKEMVHTRQACTLLGINDLYQMGIFRILAGILHLGNVGFTSRDADSCTIPHKHEPLTIFCDLMGVEYQEMSHWLCHRKLATASETYIKPISKLQAINARDALAKHIYAKLFNWIVDHVNKALHSTIKQHSFIGVLDIYGFETFEINSFEQFCINYANEKLQQQFNMHVFKLEQEEYMKEQIPWTLIDFYDNQPCINLIEAKLGILDLLDEECKMPKGSDDSWAQKLYNTHLKKCALFEKPRMSNKAFIIQHFADKVEYQCEGFLEKNKDTVYEEQIKVLKSSKQFKMLLELFQDEEKVLSPTSATPSGRTLLSRTAVKPAKAKPGQASKEHKKTVGHQFRNSLHLLMETLNATTPHYVRCIKPNDFKFPFTFDEKRAVQQLRACGVLETIRISAAGFPSRWTYQEFFSRYRVLMKQKDVLSDRKQTCKNLLENLILNKDKYQFGKTKIFFRAGQVAYLEKIRADKLRVACIRIQKTIRGWLMRKKYLRMRKAAITIQRYVRGYQARCYAKFLRRTRAAIIIQKFQRMYVVRKRYHLMRAATIVLQSLLRGYIARNKYQRMLREHKAIIIQKHVRGWLARVCYHRSLKAIVYLQCCYRRMMAKRELKKLKIEARSVERYKKLNIGMENKIMQLQRKVDEQNKDYKSLLEKLTNLETTYNTETEKLRSDVERLRVSEEEAKNATNRVNTLQEEITRLRKELHQTQSEKKTIEEWADKYKQETEQLVSELKEQNTLLKKEKEELNRRIHDQAKEMTETMEKKLVEETKQLELDLNDERLRYQNLLNEFSRLEERYEDLRDEVNLMVSIPKPGHKRTDSTHSSNESEYTFSSEITETEDLPLRMEEPSEKKAPLDMSLFLKLQKRVTELEQEKQSLQDELDRKEEQALRTKTREEEERPPMRGAELEYESLKRQELESENKKLKNELNELQKALLEKSSPEVTAPGAPAYRVLLEQLTSVSEELEVRKEEVLILRSQLVSQKEAIQPKNTMTDSTILLEDVQKMKDKGEIAQAYIGLKETNRLLESQLQSQKKSHDNELEALRGEIQSLKEENNRQQQLLAQNLQLPPEARIEASLQHEITRLTNENLDLMEQLEKQDKTVRKLKKQLKVFAKKIGELEVGQMENISPGQIIDEPIRPVNIPRKEKDFQGMLEYKKDDEPKLVKYLILDLKPRGVAVNLIPGLPAYILFMCVRHADYLNDDQKVRSLLTSTINGIKKVLKKRGDDFETVSFWLSNTCRFLHCLKQYSGEEGFMKHNTPRQNEHCLTNFDLAEYRQVLSDLAIQIYQQLIRVLENILQPMIVSGMLEHETIQGVSGVKPPGLRKRTSSIADEGTYTLDSIIRQLNTFHFVMCQHGMDPELIKQVIKQTFYITGAVTLNNLLLRKDMCSWSKGMQIRYNVSQLEEWLRDKNLMNSGAKETLEPLIQAAQLLQVKKKTDEDAEAICSMCNALTTAQIVKVLNLYTPVNEFEERVSISFIRTIQMRLRDRKDSPQLLMDAKHIFPVTFPFNPSSLALETIQIPASLGLAFISRV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0W5</accession>
    <name>A0A452J0W5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">tRNA (32-2'-O)-methyltransferase regulator THADA</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="3">Together with methyltransferase FTSJ1, methylates the 2'-O-ribose of nucleotides at position 32 of the anticodon loop of substrate tRNAs.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the THADA family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038521.1">
        <property type="protein sequence ID" value="ENSGAGP00000034021.1"/>
        <property type="gene ID" value="ENSGAGG00000024118.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030488">
        <property type="term" value="P:tRNA methylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056843">
        <property type="entry name" value="THADA-like_TPR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056842">
        <property type="entry name" value="THADA-like_TPR_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019442">
        <property type="entry name" value="THADA/TRM732_DUF2428"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051954">
        <property type="entry name" value="tRNA_methyltransferase_THADA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14387">
        <property type="entry name" value="THADA/DEATH RECEPTOR INTERACTING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14387:SF7">
        <property type="entry name" value="THYROID ADENOMA-ASSOCIATED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10350">
        <property type="entry name" value="DUF2428"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25150">
        <property type="entry name" value="TPR_Trm732"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25151">
        <property type="entry name" value="TPR_Trm732_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0819">tRNA processing</keyword>
    <feature type="domain" description="tRNA (32-2'-O)-methyltransferase regulator THADA-like TPR repeats region" evidence="6">
        <location>
            <begin position="533"/>
            <end position="805"/>
        </location>
    </feature>
    <feature type="domain" description="DUF2428" evidence="5">
        <location>
            <begin position="949"/>
            <end position="1249"/>
        </location>
    </feature>
    <feature type="domain" description="tRNA (32-2'-O)-methyltransferase regulator THADA-like C-terminal TPR repeats region" evidence="7">
        <location>
            <begin position="1251"/>
            <end position="1414"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00010409"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022694"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00035625"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00035698"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="Pfam" id="PF10350"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="Pfam" id="PF25150"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF25151"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034021.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1805" mass="203850" checksum="844E06BF12307B79" modified="2019-05-08" version="1">MVLKKKKEIQVDAFVLECQELETLRSFADAEDQNLASLLLHCVQLSDGIHQIYSMKQIVTLLKKMDRNSACSPMIKCCLDIIGVMYFSLGAKNPLKKMLASSLNGLPEQLMTEAVHSFVRCLKEELKTTDLCLYRKVMDNLASCMENFSLGMIICNIIFHIFNLFSCDLELVLQFLQKSVLEIQEENRQLSGNSIVETRLMHDLLMAIKVSMMLLQKVQENFQGSLWRKPGSLLWQSMCGLLKSFTNFLMNEGLLPTVQTTSGLAVILFTKTMFEPVEKLPCLVSDLLLRSVKCASVPEWFVSSCECLCTEELSDSVLLSLCHGALAMLEWKNGSMSENGEKLLLDIALVLLSLASRLKESNMAVSLSRILAIWTNSALDVLESSSQNLKSSLNGNSNTTGKLLEYVYTHWEHPLDAVRHQAKLIFKNILRIHQATLKGSVVKSDPFFSMLTESLLSLEWHVKGKYASLGCLVECVGVENILTVDRTIPAQILDVMGDQSLAPYASDLLETMFINHKSHLISAFQKNIWINHWHDTWVSPLLLILCEGNPDQTTYVIDYYLPKLLKCSPDSLSYMIKILQASADTLLILSLGSCNTRGALGALMACLRTARAHGHLQFREIMRDDLLSIGYVKQGLVHQHNQVRVDALGLLCETHRSTEILTMEEMQLIQFFITYNLNSQSPAVRQQICSLLKKLFCRIQESSQVLYKLEQNKTKQELIKESAKWDHFATLQQYKDFMSSICDKLFEALFPGSSHPTRFSSLTILGSIAEIFSAPEGEVFDFHKIDPTRVQMLIQCFASTFEEVKVLAFELLMKLYDRLDLQDLENLHLLFQAAMDLSISTKPYDCVTASYLLNFLIHHKGLLNVCLKNQQVADTFQIDEDMSASTMEKNTLAVIKYLLSNLEREIFQAEKSLLQAAASFPMYGRVHCITGALQQLPLNNLTLMTEWRQMVTKLILMSYRLSSVVSPIVQSSSPEGLIPMDNDSETAGRLQMILHEIQPRDTNDYFTQAKIFKEHHRVDFAKLTDDEPVENVCTEIRGKEGQTCDVTAQMVLVCCWRSMKEISLLLGMLCKLLPLQTVSESSDGLIAVEQVKDIGDYFKHHLLQSRHRGAFELAYTGFVMLTEILSRCTNESLRTLPQQWLYNVLEEIKSCDPSSKLCATRRSAGIPFYIQALLASEPKNKTNLLKMTMNELVSLAVPVNESLNIIPQVHALNILRALFRDTRFGENIIPYIADGIQAAVLGFTSPVWAVRNSSTLLFSTLITRIFGVKRGKDENSKKNRMTGREFFSRFPTLYPFLLQQLEVVARTMDSETGELKLHPSLFLLLLILGKLYPSPMDGTYSALSMAPFVPFIMRCGHSPVYRSREMAGRALVPFVMISQIPHTVQSLLAGLPECTNPCIQQNSVHGMLLQVFHLLGSYLESKHRVNSDLQQGLVDIITCIKAKVWLAKRQNPCLVTRAAYLDVLVVLSDYLGKSRRKELFGFWEEISKVISASELMTGIYYFSAIPGFTQYLQSITKLVISVLSVAADPDFSNSSSPVAEQMMNSSSMVVHLLQSDFHEVRLPMLEAVLLWWKQVNSKQATEEKMGPLSLLSDLEVILLRMAMKEKHPECFCKVLEILHCMDLRNVLPRTQDSIKMNPKEFLNWAMNIISASDSIKIQSIALKLASKLVTHLVQNCQEVMESEMRQWIQLVTSCCGDEEQTAIRLAAAEVLINITPLFLTNQKLILGLSSTLAMWKCILLLLQSEELIVRDTAADVIRVAQSQSNICNKTGTVCLLLNLVIFLLLQKHYLKRHLFCLVHQALLQP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0W6</accession>
    <name>A0A452J0W6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">Myosin motor domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">Muscle contraction.</text>
    </comment>
    <comment type="subunit">
        <text evidence="14">Muscle myosin is a hexameric protein that consists of 2 heavy chain subunits (MHC), 2 alkali light chain subunits (MLC) and 2 regulatory light chain subunits (MLC-2).</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Myofibril</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 15">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038456.1">
        <property type="protein sequence ID" value="ENSGAGP00000033958.1"/>
        <property type="gene ID" value="ENSGAGG00000020688.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030016">
        <property type="term" value="C:myofibril"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032982">
        <property type="term" value="C:myosin filament"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016460">
        <property type="term" value="C:myosin II complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000146">
        <property type="term" value="F:microfilament motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000003">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000004">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000013">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000007">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000008">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.4820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.530:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000024">
        <property type="entry name" value="Myosin heavy chain, isoform J"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.720:FF:000001">
        <property type="entry name" value="Myosin heavy chain, muscle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.340">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.370">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.4820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.2420">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002928">
        <property type="entry name" value="Myosin_tail"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014751">
        <property type="entry name" value="XRCC4-like_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615">
        <property type="entry name" value="MYOSIN HEAVY CHAIN, NON-MUSCLE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615:SF69">
        <property type="entry name" value="MYOSIN-6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01576">
        <property type="entry name" value="Myosin_tail_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90257">
        <property type="entry name" value="Myosin rod fragments"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57997">
        <property type="entry name" value="Tropomyosin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12 15" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="7 15" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="8 16" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0488">Methylation</keyword>
    <keyword evidence="10 15" id="KW-0505">Motor protein</keyword>
    <keyword evidence="11" id="KW-0514">Muscle protein</keyword>
    <keyword evidence="9 15" id="KW-0518">Myosin</keyword>
    <keyword evidence="6 15" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0787">Thick filament</keyword>
    <feature type="domain" description="Myosin motor" evidence="17">
        <location>
            <begin position="83"/>
            <end position="772"/>
        </location>
    </feature>
    <feature type="region of interest" description="Actin-binding" evidence="15">
        <location>
            <begin position="649"/>
            <end position="671"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="16">
        <location>
            <begin position="834"/>
            <end position="1107"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="16">
        <location>
            <begin position="1151"/>
            <end position="1874"/>
        </location>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <begin position="176"/>
            <end position="183"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004657"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022433"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023179"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00037488"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00038612"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033958.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1889" mass="218155" checksum="EFA745EC18C25711" modified="2019-05-08" version="1">LAKMMDMSEFGEAAPFLRKSDKEIMILQTVAFDGKTGYVFLCLVCSVERQGDSSCQQCYTCLSLQTVTVKEDDVQPMNPPKFDMIEDMAMLTHLNEASVLHNLNRRYSHWMIYTYSGLFCVTINPYKWLPVYNPEVVNAYKGKRRSEAPPHIFSIADNAYNDMLRNRENQSMLITGESGAGKTVNTKRVIQYFATVAALGEPGKKSGTLEDQIIQANPALEAFGNAKTLRNDNSSRFGKFIRIHFGTTGKLSSADIEIYLLEKSRVIFQQPGERSYHIFYQILSGKKAELQDMLLVSTNPYDYHFSSQGAVTVDNLDDGEELMATDQAMDILGFIPDEKYGAYKLTGSIMHFGNMKFKQRPREEQAETDGTESTDKAAYLMGLNSSDLAKGLLHPRVKVGNEYVTKGQSVEQVVYAVGALSKAVYDRMFKWLVIRINKTLDTRLPRQFFIGVLDIAGFEIFDYNSFEQLCINYTNEKLQQFFNHHMFVLEQEEYKKEGIEWVFIDFGLDLQACIDLIEKPLGILSILEEECMFPKATDMTFKAKLYDNHLGKSPNLQKPRIDKKRKYEAHFELLHYAGAVPYNIIGWLEKNKDPLNETVVGIFQKSANKLLASLFENYVSADSVDHSTEKKRKKGASFQTVSSLHKENLNKLMTNLRSTAPHFVRCIIPNETKTPGAMDAYLVLHQLRCNGVLEGIRICRKGFPNRVLYADFKQRYRILNPAAIPEDKFVDSRKASEKLLASLDIDQTQYRFGHTKVFFKAGLLGHLEEMRDHRLSKILTMIQARARGKLMRIEFQKIVERRDALLVIQWNIRAFMAVKNWPWMKLFFKIKPLLKSAETEKEMANMKEEFLKLKEALEKSEARRKELEEKQVSLVQEKNDLLLQLQAEQDTLADAEERCDLLIKSKIQMEAKVKELTERVEDEEEMNSELTSKKRKLEDECSELKKDIDDLEITLAKVEKEKHATENKVKNLTEEMASLDETISKLTKEKKSLQEAHQQALDDLQAEEDKVNTLSKTKLKLEQQVDDLEGSLEQEKKIRMDLERVKRKLEGDLKLTQENTVCIARIEELEEELEAERAARAKVEKQRSDLARELEELSERLEEAGGATAAQLEMNKKREAEFLKVRRDLEEATLHYEATAATLRKKHADSMAELGEQVDNLQRVKQKLEKEKSELKLEVDDLLSNMEQMVKGKSNAEKLCRTYEDNLNETKTKLDEMTRLMNDLTTQKGKLQTENGEFARQLEEKESLISQLSRGKTSFTQQIEELRRLLEEETKSKNALAHALQAARHDCDLLREQFEEEQEAKAELQRALSKGNAEVAQWRTKYETDAIQRTEELEEAKKKLAARLQEAEEAIEAANAKCSSLEKTKHRLQNELEDIVIDLERANSAAAALDKKQRNFDKLISDWKQKYEETQAELEASQKEARSLSTELFKLKNAYEETLDNLETVKRENKNLQEEISDLTDQISEGNKSLHEIEKIKKQVEQEKSEVQLALEEAEGALEHEESKSLRFQLEISQIKAEFERKLAEKDEEMENIRRNQQRTIDSLQSTLDSEAKSRNEAIRLKKKMEGDLNEMEIQLSHANRHASEATKSARILQAQIKDLQVQLDDSEHLNEDLKEQLAVTDRRNNLIQSELDELRALLDQTERGRKLAEQELLEATERVNLLHTQNTSLINQKKKLETDISQMQNEVEEAIQECRNAEEKAKKAITDAAMMAEELKKEQDTSAHLERMKKNMEQTIKDLQKRLDEAEQIALKGGKKQIQKLESRVRELENDLENELRRNTESQKGVRKYERRIKELTYQAEEDKKNLTRMQELIDKLQLKVKSYKHQAEEAEAQSNQYLAKYRKQQHELDDAEERAEIAESQVNKLRTKTRDIGIKKVWCYSAE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0W7</accession>
    <name>A0A452J0W7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Dynein light chain roadblock</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Acts as one of several non-catalytic accessory components of the cytoplasmic dynein 1 complex that are thought to be involved in linking dynein to cargos and to adapter proteins that regulate dynein function. Cytoplasmic dynein 1 acts as a motor for the intracellular retrograde motility of vesicles and organelles along microtubules.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 10">Cytoplasm</location>
            <location evidence="1 10">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 10">Belongs to the GAMAD family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0W7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038512.1">
        <property type="protein sequence ID" value="ENSGAGP00000034012.1"/>
        <property type="gene ID" value="ENSGAGG00000024173.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005868">
        <property type="term" value="C:cytoplasmic dynein complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005874">
        <property type="term" value="C:microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045505">
        <property type="term" value="F:dynein intermediate chain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007018">
        <property type="term" value="P:microtubule-based movement"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.450.30:FF:000002">
        <property type="entry name" value="Dynein light chain roadblock"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.450.30">
        <property type="entry name" value="Dynein light chain 2a, cytoplasmic"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016561">
        <property type="entry name" value="DYNLRB1/2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004942">
        <property type="entry name" value="Roadblock/LAMTOR2_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10779">
        <property type="entry name" value="DYNEIN LIGHT CHAIN ROADBLOCK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03259">
        <property type="entry name" value="Robl_LC7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF009998">
        <property type="entry name" value="DLC7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00960">
        <property type="entry name" value="Robl_LC7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103196">
        <property type="entry name" value="Roadblock/LC7 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 10" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8 10" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="6 10" id="KW-0243">Dynein</keyword>
    <keyword evidence="5 10" id="KW-0493">Microtubule</keyword>
    <keyword evidence="7 10" id="KW-0505">Motor protein</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 10" id="KW-0813">Transport</keyword>
    <feature type="domain" description="Roadblock/LAMTOR2" evidence="11">
        <location>
            <begin position="4"/>
            <end position="92"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007191"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023017"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00025362"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PIRNR" id="PIRNR009998"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="SMART" id="SM00960"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034012.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="96" mass="10866" checksum="FB657814F659B59A" modified="2019-05-08" version="1">MAEVEETLKRIQSQKGVQGIIVVNSEGIPIKSTVDNSTTVQYAGLMHSFIMKARSTVRDIDPQNDLTFLRIRSKKNEIMIAPDKDYFLIVIQNPTE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0W9</accession>
    <name>A0A452J0W9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="3">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000038526.1">
        <property type="protein sequence ID" value="ENSGAGP00000034026.1"/>
        <property type="gene ID" value="ENSGAGG00000023990.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055037">
        <property type="term" value="C:recycling endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005802">
        <property type="term" value="C:trans-Golgi network"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032367">
        <property type="term" value="P:intracellular cholesterol transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000080">
        <property type="entry name" value="lisH domain and HEAT repeat-containing protein KIAA1468 homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006594">
        <property type="entry name" value="LisH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040362">
        <property type="entry name" value="RELCH"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR32059">
        <property type="entry name" value="RAB11-BINDING PROTEIN RELCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR32059:SF0">
        <property type="entry name" value="RAB11-BINDING PROTEIN RELCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00667">
        <property type="entry name" value="LisH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50896">
        <property type="entry name" value="LISH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1"/>
            <end position="51"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="422"/>
            <end position="447"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="182"/>
            <end position="216"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="343"/>
            <end position="377"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="2">
        <location>
            <begin position="25"/>
            <end position="35"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="427"/>
            <end position="447"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034026.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1028" mass="113958" checksum="658A6C6753883C2E" modified="2019-05-08" version="1">MAAAGGGSSVNPFLSDSEEEEPRDGGGGNDPGLGGPLSPQQHEEEEEAAVEVPGVLPGLRAAAPGEAGWAGPGAGEPPRVPVDALAAQLLRDQFLLTALELHTELLESGRELPRLRDYFSNPGNFERQSGTPPAGGAPGLWGAAGLGLLGGCELITEGGGDRAGSISTLDSLDFARYSDDGNRETDERVAVLEFELRKAKETIQALRANLTQAAENEVPLQERKNYKSSPEIQEPIKPLEKRALNFLVNEFLLKNSYKLTSITFSDESDDQDFELWDDVGLNIPKPPDLLQLYRDFGNHQVTARDVVDVAVGVEEDELEVTTPTLGSVPVFGVSQSTEQFLIVQKLEDQINALNSEKWSLTEQIRRLESEIDFLKNENFAVPVVCDLVHVPLSVPEDNGSYLDIRSSDNDSKHGKTEEISLSISAEADSKTPKDDTLNSLPSKDREKLPSCALSNKAAIHFDKPNRTLSPAFHQALLSFCRMSAESRLGSEVSRIADSEKSVMLMLGRCLPHIVPNVLLAKREELIPLILCTACLHPEPKERDQLLHILFNLIKRPDDEQRQMILTGCVAFARHVGPTRVEAELLPQCWEQINHKYPERRLLVAESCGALAPYLPKEIRSSLVLSMLQQMLMEDKADLVREAVIKSLGIIMGYIDDPDKYQQGFELLLSALGDPSERVVSATHQVFLPAYAAWTTELGNLQLHLIPTLLNKIEKLLREGEHGLDEHKLHMYLSALQSLIPSLFSLVLQNAPFTSKAKLQGEVPPIEVTRFPRPVSPLQDVATIIGSREQLAVLLQLYDYQLELEGTTGWETLLWVVNQLLPQLIEIVGKINVASTACVHEFSRFFWRLCRTFGKIFTNTKVKPQFQEILRLSEENIDSTAGNGVLTKATVPIYATGVLTCYIQEEDRKLLIGFLEDVMTMLSLSHAPLNSLKASFVELGANPAYHELLLTVLWYGVVHTSALVRCTAARMFELCKDSHNSGFWYHHGNSYSERXKSKNAVSVFPGRSSVSRPTFFTDRDHKNVWKSWT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0X2</accession>
    <name>A0A452J0X2_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="13">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasmic vesicle</location>
            <location evidence="2">Autophagosome membrane</location>
            <topology evidence="2">Lipid-anchor</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 12">Belongs to the ATG8 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0X2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038517.1">
        <property type="protein sequence ID" value="ENSGAGP00000034017.1"/>
        <property type="gene ID" value="ENSGAGG00000024174.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000421">
        <property type="term" value="C:autophagosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031410">
        <property type="term" value="C:cytoplasmic vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005874">
        <property type="term" value="C:microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006914">
        <property type="term" value="P:autophagy"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.20.90:FF:000059">
        <property type="entry name" value="Microtubule-associated proteins 1A/1B light chain 3B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.20.90">
        <property type="entry name" value="Phosphatidylinositol 3-kinase Catalytic Subunit, Chain A, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004241">
        <property type="entry name" value="Atg8-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029071">
        <property type="entry name" value="Ubiquitin-like_domsf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10969">
        <property type="entry name" value="MICROTUBULE-ASSOCIATED PROTEINS 1A/1B LIGHT CHAIN 3-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02991">
        <property type="entry name" value="ATG8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54236">
        <property type="entry name" value="Ubiquitin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6 12" id="KW-0072">Autophagy</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="10" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="8" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="9 11" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="7" id="KW-0472">Membrane</keyword>
    <keyword evidence="5" id="KW-0493">Microtubule</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <feature type="lipid moiety-binding region" description="Phosphatidylserine amidated glycine; alternate" evidence="11">
        <location>
            <position position="120"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004512"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007293"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023006"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PIRSR" id="PIRSR604241-50"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="RuleBase" id="RU004384"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034017.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="121" mass="14345" checksum="748DAD260D552399" modified="2019-05-08" version="1">MPSDRPFKQRRSFVDRSKEVQQIREQHPSKIPVIIERYKGEKQLPVLDKTKFLVPDHVNMSELVKIIRRRLQLNPTQAFFLLVNQHSMVSVSTPISEIYEQEKDEDGFLYMVYASQETFGY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J151</accession>
    <name>A0A452J151_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="17">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
            <location evidence="1">Basement membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="12">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J151"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038616.1">
        <property type="protein sequence ID" value="ENSGAGP00000034104.1"/>
        <property type="gene ID" value="ENSGAGG00000024224.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005604">
        <property type="term" value="C:basement membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009887">
        <property type="term" value="P:animal organ morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007411">
        <property type="term" value="P:axon guidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009888">
        <property type="term" value="P:tissue development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00055">
        <property type="entry name" value="EGF_Lam"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000067">
        <property type="entry name" value="Laminin subunit gamma 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000441">
        <property type="entry name" value="Laminin subunit gamma 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000533">
        <property type="entry name" value="Laminin subunit gamma 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000174">
        <property type="entry name" value="Laminin subunit gamma-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050440">
        <property type="entry name" value="Laminin/Netrin_ECM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000034">
        <property type="entry name" value="Laminin_IV"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002049">
        <property type="entry name" value="LE_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10574:SF313">
        <property type="entry name" value="LAMININ SUBUNIT GAMMA-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10574">
        <property type="entry name" value="NETRIN/LAMININ-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00053">
        <property type="entry name" value="EGF_laminin"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00052">
        <property type="entry name" value="Laminin_B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00011">
        <property type="entry name" value="EGFLAMININ"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00180">
        <property type="entry name" value="EGF_Lam"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SMART" id="SM00281">
        <property type="entry name" value="LamB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01248">
        <property type="entry name" value="EGF_LAM_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50027">
        <property type="entry name" value="EGF_LAM_2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51115">
        <property type="entry name" value="LAMININ_IVA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0084">Basement membrane</keyword>
    <keyword evidence="7" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="8 13" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="9 12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11 12" id="KW-0424">Laminin EGF-like domain</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="4 14" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="14">
        <location>
            <begin position="1"/>
            <end position="19"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019506221" evidence="14">
        <location>
            <begin position="20"/>
            <end position="1155"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="15">
        <location>
            <begin position="77"/>
            <end position="123"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="15">
        <location>
            <begin position="127"/>
            <end position="174"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin IV type A" evidence="16">
        <location>
            <begin position="201"/>
            <end position="369"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="15">
        <location>
            <begin position="504"/>
            <end position="559"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="13">
        <location>
            <begin position="624"/>
            <end position="651"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="13">
        <location>
            <begin position="834"/>
            <end position="875"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="13">
        <location>
            <begin position="1002"/>
            <end position="1029"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="13">
        <location>
            <begin position="1083"/>
            <end position="1138"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="77"/>
            <end position="89"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="97"/>
            <end position="106"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="145"/>
            <end position="154"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="528"/>
            <end position="537"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004302"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022869"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023292"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00460"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50027"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS51115"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034104.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1155" mass="127343" checksum="4921E74ABA571D31" modified="2019-05-08" version="1">LPKLLRSLCLTVAIISVLSVCDCNGKSSQCIFDRDLLRETGNGYRCLNCIDNTDGIHCERCKEEFYRQRDGESCLPCNCSPRGSLGLQCDNYGRCSCKPGVMGDKCDRCQWGFHSFSEAGCQEQRPCDCDPAGSAGHCVSGRCVCKASVSGERCDRCKQGYYHLEAGNPEGCSQCFCYGHSAMCASAENYNIYKITSTFQQDSEGWRGVYVNGSPVQLQWSPRHRDVFVTARQSEPTYFLAPARFLGNQQVSYGQTLSFDYRMDRGGRRPSPQDVVLEGAGLRVTASLVPHGKMLPCGISKTYTFRLDEHPRSGWSPKLSYFEYRRLLGNLTALWIRATYGEYSTGYIDNVTLVSAQLTSGAPASWVEQCVCPPGYQGQFCERCAPGYKRDASNLGPFSTCVLCNCQGGGNCDPDTGDCYSGDENMDPNANCPFGFYRDPWNLQSCQLCPCRNGQGCSVAPGRKEVICNNCPLGVTGANCERCADGYFGNPLGDNGHVRPCQPCQCNNNVDPSAVGNCNRLTGECLKCLYNTAGFACDQCRDGFFGNPLALNPAEKCQACNCNSVGSDPLKCRSDGSCVCKPGFEGPNCEHTQCPACYSHVKTQVDQYLQQLQHLEVLVSRLQLGSGAEANEELEGKMQQAEETLREILRETLTSDKSLESRMSKMKGQESSYQSRLADIRVMVQRLQVLGHQYQAQVQDIRRLIEKAHLDLGQSKVTLSGLVSSPQDFSAQALQTSNIPGHCCRGQRNPLLLTSQGGSGLCVSWAVKHEVSVLAYDAFLFGRYDEAKLLASELETDTSRSALDADRAYQGSLVLLSSMSRFSKIDTGSFQKEANRLRQEADALMGLVETYMAEYRRLQSNTRTWEEEIKQLLQSGESKRLVSSQLLSRANLAKSRAQQALSAGNATFYEVELILKNLRGFHLQVDGRRREAEEAMRRLPLISNMVTNANDKTRRAEGALGSAATDAKTARSMAGEAKEITGGIQQEIRRLNLEANRTADGVLALEKGVITLLNEAREAERELTRKALEVDMDATTAQKTGQEAQGAHASARTAEAAVQETLHALEDFLHLMDQPGPVDEQALRLLELNLTAARTKNSQLKEQMAELEKTASLQRLRVQTLERSVNEILADIRNLEEIRNNLPPGCYNTNSIEKP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J157</accession>
    <name>A0A452J157_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Oxysterol-binding protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">Binds phospholipids; exhibits strong binding to phosphatidic acid and weak binding to phosphatidylinositol 3-phosphate. Stabilizes GTP-bound RAB7A on late endosomes/lysosomes and alters functional properties of late endocytic compartments via its interaction with RAB7A. Binds 25-hydroxycholesterol and cholesterol.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Late endosome</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 12">Belongs to the OSBP family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J157"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038561.1">
        <property type="protein sequence ID" value="ENSGAGP00000034054.1"/>
        <property type="gene ID" value="ENSGAGG00000024195.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005770">
        <property type="term" value="C:late endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097038">
        <property type="term" value="C:perinuclear endoplasmic reticulum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015485">
        <property type="term" value="F:cholesterol binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006869">
        <property type="term" value="P:lipid transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd13285">
        <property type="entry name" value="PH_ORP1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000094">
        <property type="entry name" value="Oxysterol-binding protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000098">
        <property type="entry name" value="Oxysterol-binding protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000231">
        <property type="entry name" value="Oxysterol-binding protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.160.120:FF:000005">
        <property type="entry name" value="Oxysterol-binding protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.3490:FF:000003">
        <property type="entry name" value="Oxysterol-binding protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.160.120">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.3490">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037239">
        <property type="entry name" value="OSBP_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000648">
        <property type="entry name" value="Oxysterol-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018494">
        <property type="entry name" value="Oxysterol-bd_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10972">
        <property type="entry name" value="OXYSTEROL-BINDING PROTEIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10972:SF53">
        <property type="entry name" value="OXYSTEROL-BINDING PROTEIN-RELATED PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01237">
        <property type="entry name" value="Oxysterol_BP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00233">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF144000">
        <property type="entry name" value="Oxysterol-binding protein-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01013">
        <property type="entry name" value="OSBP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6 11" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="7 14" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0967">Endosome</keyword>
    <keyword evidence="8 13" id="KW-0445">Lipid transport</keyword>
    <keyword evidence="9" id="KW-0446">Lipid-binding</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="3 13" id="KW-0813">Transport</keyword>
    <feature type="repeat" description="ANK" evidence="11">
        <location>
            <begin position="47"/>
            <end position="79"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="11">
        <location>
            <begin position="80"/>
            <end position="112"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="11">
        <location>
            <begin position="175"/>
            <end position="207"/>
        </location>
    </feature>
    <feature type="domain" description="PH" evidence="16">
        <location>
            <begin position="237"/>
            <end position="336"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="804"/>
            <end position="831"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="14">
        <location>
            <begin position="435"/>
            <end position="462"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="14">
        <location>
            <begin position="888"/>
            <end position="922"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004603"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023121"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00055419"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="RuleBase" id="RU003844"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="RuleBase" id="RU003845"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034054.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="959" mass="108441" checksum="D696519E7B867D5F" modified="2019-05-08" version="1">MITEAEQQLLHDARNGNCEEVKQLLETMDKGEVVVDINCKGRSKSNLGWTPLHLACYFGHGNVVKDLLKAGAEVNVLNDMGDTPLHRAAFTGRKEVVMLLLRYDADTSIVNGGGQTARDVTQDKEIKNMLEAVERTQERKLEELLLGAAREGETAKLSALLNRPKPPDINCTDQMGNTPLHCAAYRAHKHCALKLLKKGADSKIRNKNDQTPFDLAQGAEIKQILAGSIGKVMNKRLKRYEGLLWKSLRFFGWKLYWVVLEHGVLSWYRKQADAVNNDHRQGCKHLTQAVCTVKSTDSCLFSVKCFDDSVHSFKVPKNSPPKARENWLEAIEDHSAYSTHYCTQEQLSDEEDDDTVSVAELQETLKKAQTCQQRLDKEISNFLAMIKSCSTTKGMPPSFLQKIEVVSEVSKETCEALGNCLKLFTKQEGVRNLKLEHEQEKNKILSEALETLATEHHELEQSLVKGSPPLSILSEEQFYDAVSDSESEKSLSGFETVTAYSLEDSVESNNTITSSMSEEKVCGGGGKVCGGGESLSNGIKKHRTSLPSPMFSRNDFSIWSILRKCIGMELSKITMPVIFNEPLSFLQRLTEYMEHTYLIHQASSLSGTVERMQCVAAFAVSAVASQWERTGKPFNPLLGETYELVRDDLGFRLLSEQVSHHPPISAFYAEGLNNDFVFHGSIYPKLKFWGKSVEAEPKGTITLELLEHDEAYTWTNPTCCVHNIIVGKLWIEQYGNVEITNHKTGEKCVLNFKPCGLFGKELHKVEGYIQDKSKRKVCSLYGKWTECLYSVDPATFEAYKKNDKKNTEEKKNSKQVGNSEEPDEMPLPDSESVHVIPGSILLWRIAPRPPNSVQMYNFTSFAMALNELDKEMESVVPKTDCRLRPDIRAMENGEIDIASEEKKRLEEKQRAVRKNRSKSEEDWKTRWFHQGPNPYTGAQDWLYSGNYWDRNYFNLPDIY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J159</accession>
    <name>A0A452J159_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">E3 ubiquitin-protein ligase CHFR</fullName>
            <ecNumber evidence="5">2.3.2.27</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="17">Checkpoint with forkhead and RING finger domains protein</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="16">RING-type E3 ubiquitin transferase CHFR</fullName>
        </alternativeName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>S-ubiquitinyl-[E2 ubiquitin-conjugating enzyme]-L-cysteine + [acceptor protein]-L-lysine = [E2 ubiquitin-conjugating enzyme]-L-cysteine + N(6)-ubiquitinyl-[acceptor protein]-L-lysine.</text>
            <dbReference type="EC" id="2.3.2.27"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="3">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
            <location evidence="2">PML body</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the CHFR family.</text>
    </comment>
    <dbReference type="EC" id="2.3.2.27" evidence="5"/>
    <dbReference type="AlphaFoldDB" id="A0A452J159"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038613.1">
        <property type="protein sequence ID" value="ENSGAGP00000034101.1"/>
        <property type="gene ID" value="ENSGAGG00000024231.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00143"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016605">
        <property type="term" value="C:PML body"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061630">
        <property type="term" value="F:ubiquitin protein ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016567">
        <property type="term" value="P:protein ubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006511">
        <property type="term" value="P:ubiquitin-dependent protein catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd22672">
        <property type="entry name" value="FHA_CHFR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd16503">
        <property type="entry name" value="RING-HC_CHFR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.40.10:FF:000203">
        <property type="entry name" value="E3 ubiquitin-protein ligase CHFR isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.40.140:FF:000001">
        <property type="entry name" value="E3 ubiquitin-protein ligase CHFR isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.200.20:FF:000022">
        <property type="entry name" value="E3 ubiquitin-protein ligase CHFR isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.200.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040909">
        <property type="entry name" value="CHFR_Znf-CRD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052256">
        <property type="entry name" value="E3_ubiquitin-ligase_CHFR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000253">
        <property type="entry name" value="FHA_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008984">
        <property type="entry name" value="SMAD_FHA_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001841">
        <property type="entry name" value="Znf_RING"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017907">
        <property type="entry name" value="Znf_RING_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16079:SF4">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE CHFR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16079">
        <property type="entry name" value="UBIQUITIN LIGASE PROTEIN CHFR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00498">
        <property type="entry name" value="FHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13923">
        <property type="entry name" value="zf-C3HC4_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17979">
        <property type="entry name" value="zf-CRD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00240">
        <property type="entry name" value="FHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00184">
        <property type="entry name" value="RING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57850">
        <property type="entry name" value="RING/U-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49879">
        <property type="entry name" value="SMAD/FHA domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50006">
        <property type="entry name" value="FHA_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00518">
        <property type="entry name" value="ZF_RING_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50089">
        <property type="entry name" value="ZF_RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="15" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="7" id="KW-0132">Cell division</keyword>
    <keyword evidence="9" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-0498">Mitosis</keyword>
    <keyword evidence="14" id="KW-0539">Nucleus</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0808">Transferase</keyword>
    <keyword evidence="12" id="KW-0833">Ubl conjugation pathway</keyword>
    <keyword evidence="13" id="KW-0862">Zinc</keyword>
    <keyword evidence="10 18" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="FHA" evidence="20">
        <location>
            <begin position="38"/>
            <end position="89"/>
        </location>
    </feature>
    <feature type="domain" description="RING-type" evidence="21">
        <location>
            <begin position="304"/>
            <end position="343"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="145"/>
            <end position="268"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="390"/>
            <end position="414"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="145"/>
            <end position="156"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="157"/>
            <end position="166"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="175"/>
            <end position="193"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="220"/>
            <end position="233"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="234"/>
            <end position="265"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="19">
        <location>
            <begin position="400"/>
            <end position="414"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004322"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00005797"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00017908"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022776"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00029800"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00031332"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50006"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034101.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="628" mass="70351" checksum="2B51A00C4B95A350" modified="2019-05-08" version="1">MEHSEGGDQNQQQQPWGKLIRLGTDEAEPHVLLLKREWTIGRKKGCDLSFPGNKLVSGDHCKITVDKESGQASLEDTSTNGTVINKLKVVKKQTCPLQTGDVIYVVYRKNEPENNVAYLYESFNAKQDVTQEPVEANVENTCHLTKDTSHIGRSDDGTQITSSLPATESCYEEPQPSTSTSNLFNASSTSLIEQASAERDNPSTSEPRYPGASHEEQGQLYANSETSTITSNIIDKERNESFTSEPQREKEEDLEPAKKKMKGDENASPYLQMTAPNQCIIKAGSEGAKPTNVKPDKMEETLTCIICQELLHDCVSLQPCMHTFCAACYSGWMERSSLCPTCRCPVERICKNHILNNLVEAYLIQHPDKCRNEEDVRSMDARNKITQDMLQPKVRRSFSDEEGSSEDLLELSDVDSESSDISQPYIVCRQCPGYRRHSIPPLPCPGQEGEAGGLQALGDAPSTSADFPATVQEYVCPSQGSHVICTCCFQPMPDRRAEREQNPNIAPQQCTVCLQPFCHLYWGCTRMACFGCLAPFCEINLGDKCLDGILNNNHYESDILKDYLASRGFTWKNMLNESLLALQREVFMLSVTVTSRPDCYWGRNCRTQVKAHHAMKFNHICEQTRFKN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J162</accession>
    <name>A0A452J162_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="17">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
            <location evidence="1">Basement membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="12">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J162"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038618.1">
        <property type="protein sequence ID" value="ENSGAGP00000034106.1"/>
        <property type="gene ID" value="ENSGAGG00000024224.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005604">
        <property type="term" value="C:basement membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009887">
        <property type="term" value="P:animal organ morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007411">
        <property type="term" value="P:axon guidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009888">
        <property type="term" value="P:tissue development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00055">
        <property type="entry name" value="EGF_Lam"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000067">
        <property type="entry name" value="Laminin subunit gamma 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000441">
        <property type="entry name" value="Laminin subunit gamma 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000533">
        <property type="entry name" value="Laminin subunit gamma 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000174">
        <property type="entry name" value="Laminin subunit gamma-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050440">
        <property type="entry name" value="Laminin/Netrin_ECM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000034">
        <property type="entry name" value="Laminin_IV"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002049">
        <property type="entry name" value="LE_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10574:SF313">
        <property type="entry name" value="LAMININ SUBUNIT GAMMA-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10574">
        <property type="entry name" value="NETRIN/LAMININ-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00053">
        <property type="entry name" value="EGF_laminin"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00052">
        <property type="entry name" value="Laminin_B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00011">
        <property type="entry name" value="EGFLAMININ"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SMART" id="SM00180">
        <property type="entry name" value="EGF_Lam"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SMART" id="SM00281">
        <property type="entry name" value="LamB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01248">
        <property type="entry name" value="EGF_LAM_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50027">
        <property type="entry name" value="EGF_LAM_2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51115">
        <property type="entry name" value="LAMININ_IVA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0084">Basement membrane</keyword>
    <keyword evidence="7" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="8 13" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="9 12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11 12" id="KW-0424">Laminin EGF-like domain</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="4 14" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="14">
        <location>
            <begin position="1"/>
            <end position="19"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019149071" evidence="14">
        <location>
            <begin position="20"/>
            <end position="1113"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="15">
        <location>
            <begin position="77"/>
            <end position="123"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="15">
        <location>
            <begin position="132"/>
            <end position="179"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin IV type A" evidence="16">
        <location>
            <begin position="206"/>
            <end position="374"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="15">
        <location>
            <begin position="509"/>
            <end position="564"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="13">
        <location>
            <begin position="629"/>
            <end position="663"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="13">
        <location>
            <begin position="809"/>
            <end position="836"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="13">
        <location>
            <begin position="963"/>
            <end position="990"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="13">
        <location>
            <begin position="1041"/>
            <end position="1096"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="77"/>
            <end position="89"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="97"/>
            <end position="106"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="150"/>
            <end position="159"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="533"/>
            <end position="542"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004302"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022869"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023292"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00460"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50027"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS51115"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034106.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1113" mass="122138" checksum="DC64C92F7B7669CE" modified="2019-05-08" version="1">LPKLLRSLCLTVAIISVLSVCDCNGKSSQCIFDRDLLRETGNGYRCLNCIDNTDGIHCERCKEEFYRQRDGESCLPCNCSPRGSLGLQCDNYGRCSCKPGVMGDKCDRCQWGFHSFSEAGCQAGGQVSSAQCDCDPAGSAGHCVSGRCVCKASVSGERCDRCKQGYYHLEAGNPEGCSQCFCYGHSAMCASAENYNIYKITSTFQQDSEGWRGVYVNGSPVQLQWSPRHRDVFVTARQSEPTYFLAPARFLGNQQVSYGQTLSFDYRMDRGGRRPSPQDVVLEGAGLRVTASLVPHGKMLPCGISKTYTFRLDEHPRSGWSPKLSYFEYRRLLGNLTALWIRATYGEYSTGYIDNVTLVSAQLTSGAPASWVEQCVCPPGYQGQFCERCAPGYKRDASNLGPFSTCVLCNCQGGGNCDPDTGDCYSGDENMDPNANCPFGFYRDPWNLQSCQLCPCRNGQGCSVAPGRKEVICNNCPLGVTGANCERCADGYFGNPLGDNGHVRPCQPCQCNNNVDPSAVGNCNRLTGECLKCLYNTAGFACDQCRDGFFGNPLALNPAEKCQACNCNSVGSDPLKCRSDGSCVCKPGFEGPNCEHTQCPACYSHVKTQVDQYLQQLQHLEVLVSRLQLGSGAEANEELEGKMQQAEETLREILRETLSLQGNARHLPLDGSRMSKMKGQESSYQSRLADIRVMVQRLQVLGHQYQAQVQDIRRLIEKAHLDLGQSKVTLSGLVSSPQDFSAQALQTTLLFSSHVQSANAIEQAVRAAVDDSGQALELVRSAVSRGGILASSVQGLLRKQEADALMGLVETYMAEYRRLQSNTRTWEEEIKQLLQSGESKRLVSSQLLSRANLAKSRAQQALSAGNATFYEVELILKNLRGFHLQVDGRRREAEEAMRRLPLISNMVTNANDKTRRAEGALGSAATDAKTARSMAGEAKEITGGIQQEIRRLNLEANRTADGVLALEKGVITLLNEAREAERELTRKALEVDMDATTAQKEAQGAHASARTAEAAVQETLHALEDFLHLMDQPGPVDEQALRLLELNLTAARTKNSQLKEQMAELEKTASLQRLRVQTLERSVNEILADIRNLEEIRNNLPPGCYNTNSIEKP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J164</accession>
    <name>A0A452J164_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Xylose isomerase-like TIM barrel domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J164"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038635.1">
        <property type="protein sequence ID" value="ENSGAGP00000034120.1"/>
        <property type="gene ID" value="ENSGAGG00000024243.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008903">
        <property type="term" value="F:hydroxypyruvate isomerase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046487">
        <property type="term" value="P:glyoxylate metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.20.20.150">
        <property type="entry name" value="Divalent-metal-dependent TIM barrel enzymes"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050417">
        <property type="entry name" value="Sugar_Epim/Isomerase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036237">
        <property type="entry name" value="Xyl_isomerase-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013022">
        <property type="entry name" value="Xyl_isomerase-like_TIM-brl"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43489:SF6">
        <property type="entry name" value="HYDROXYPYRUVATE ISOMERASE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43489">
        <property type="entry name" value="ISOMERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01261">
        <property type="entry name" value="AP_endonuc_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51658">
        <property type="entry name" value="Xylose isomerase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0413">Isomerase</keyword>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Xylose isomerase-like TIM barrel" evidence="2">
        <location>
            <begin position="2"/>
            <end position="174"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00023235"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF01261"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034120.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="202" mass="22619" checksum="06AB5B9A5CD9F295" modified="2019-05-08" version="1">AAFREGLALAVKYAKTLGCPRIHLMAGRLPAGTKQAAMALEMEATFIENLRYAADILAQEDMTGLLEPMNSRITDPRYFLTTPHQAAAILEKVGRPNLRLQLDIFHCQIMDGNLTRNLEMYFPLIGHIQVGQVPGRHEPDSPGEVNFPYLFQLLESLGYEGYVGCEYTPQGEWAVRSSWCQGDQDHPWVSISNPFSVKHTCP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J165</accession>
    <name>A0A452J165_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">Anaphase-promoting complex subunit 4</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J165"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038651.1">
        <property type="protein sequence ID" value="ENSGAGP00000034134.1"/>
        <property type="gene ID" value="ENSGAGG00000024170.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005680">
        <property type="term" value="C:anaphase-promoting complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034399">
        <property type="term" value="C:nuclear periphery"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031145">
        <property type="term" value="P:anaphase-promoting complex-dependent catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070979">
        <property type="term" value="P:protein K11-linked ubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024789">
        <property type="entry name" value="APC4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024977">
        <property type="entry name" value="Apc4-like_WD40_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024790">
        <property type="entry name" value="APC4_long_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036322">
        <property type="entry name" value="WD40_repeat_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13260">
        <property type="entry name" value="ANAPHASE PROMOTING COMPLEX SUBUNIT 4 APC4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13260:SF0">
        <property type="entry name" value="ANAPHASE-PROMOTING COMPLEX SUBUNIT 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12896">
        <property type="entry name" value="ANAPC4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12894">
        <property type="entry name" value="ANAPC4_WD40"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50978">
        <property type="entry name" value="WD40 repeat-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="2" id="KW-0132">Cell division</keyword>
    <keyword evidence="6" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0498">Mitosis</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="4" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="393"/>
            <end position="411"/>
        </location>
    </feature>
    <feature type="domain" description="Anaphase-promoting complex subunit 4-like WD40" evidence="7">
        <location>
            <begin position="19"/>
            <end position="95"/>
        </location>
    </feature>
    <feature type="domain" description="Anaphase-promoting complex subunit 4 long" evidence="8">
        <location>
            <begin position="198"/>
            <end position="388"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00016067"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022776"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF12894"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF12896"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034134.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="498" mass="56134" checksum="FFE90F32DC912CE5" modified="2019-05-08" version="1">ILFVKNSPVYQWFSICGPQTHVLLHRLANFQRVWSLPPNENTGKEVTALAWRPDGKILAFGLADTKRVILCDVEKPESLHSFSVDAPVTYMHWMEVTEESSVLTSFYNAEDESNLLLPKLPALPKNNFCDLFVIRLNALVLGGSSGFIEVYAYGMYKIATVTGVAGSCLGLCFSSDLKSLSVITEVQVSPDSAPEITYFQLDTSLLSVYLPEVTRMARKFTHISTLLQYIKLSLTCMCEAWEEILMQMDSRLTKFVQEKNTTTSVQDEFMQLLLWGKASLELQALLMNQLTVKGLKKLGQSIESSYSSIQKLVISHLQSGSEALLYHLSELKGMASWKQKYESLGLDAAGIEGAITAVGSFILKANELLQVIDSSMKNFKAFFRWLYVGKLNYILSISYLLYILIICFGIMTQKDITFVADFLTEHFNEAPDLYNRKGKYFNVERVGQYLKDEDDDLVSPPNTEGNQWFNFLQNSTHLKGISCLLQTVCILGQQLLLF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J176</accession>
    <name>A0A452J176_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">Anaphase-promoting complex subunit 4</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J176"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038638.1">
        <property type="protein sequence ID" value="ENSGAGP00000034123.1"/>
        <property type="gene ID" value="ENSGAGG00000024170.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005680">
        <property type="term" value="C:anaphase-promoting complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034399">
        <property type="term" value="C:nuclear periphery"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031145">
        <property type="term" value="P:anaphase-promoting complex-dependent catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070979">
        <property type="term" value="P:protein K11-linked ubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030071">
        <property type="term" value="P:regulation of mitotic metaphase/anaphase transition"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024789">
        <property type="entry name" value="APC4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024977">
        <property type="entry name" value="Apc4-like_WD40_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056358">
        <property type="entry name" value="APC4_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024790">
        <property type="entry name" value="APC4_long_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017169">
        <property type="entry name" value="APC4_metazoa"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036322">
        <property type="entry name" value="WD40_repeat_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13260">
        <property type="entry name" value="ANAPHASE PROMOTING COMPLEX SUBUNIT 4 APC4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13260:SF0">
        <property type="entry name" value="ANAPHASE-PROMOTING COMPLEX SUBUNIT 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12896">
        <property type="entry name" value="ANAPC4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12894">
        <property type="entry name" value="ANAPC4_WD40"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23405">
        <property type="entry name" value="WD40_APC4_C-half"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF037303">
        <property type="entry name" value="APC4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50978">
        <property type="entry name" value="WD40 repeat-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="2" id="KW-0132">Cell division</keyword>
    <keyword evidence="3" id="KW-0498">Mitosis</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="Anaphase-promoting complex subunit 4-like WD40" evidence="7">
        <location>
            <begin position="19"/>
            <end position="97"/>
        </location>
    </feature>
    <feature type="domain" description="Anaphase-promoting complex subunit 4 long" evidence="8">
        <location>
            <begin position="197"/>
            <end position="395"/>
        </location>
    </feature>
    <feature type="domain" description="Anaphase-promoting complex subunit 4 C-terminal half WD40" evidence="9">
        <location>
            <begin position="526"/>
            <end position="696"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="699"/>
            <end position="754"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="6">
        <location>
            <begin position="699"/>
            <end position="714"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00016067"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022776"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF12894"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF12896"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF23405"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034123.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="754" mass="85627" checksum="D9EC3109B3CCB8A6" modified="2019-05-08" version="1">SQFLFSLFPSLPPQHILFVKNSPVLLHRLANFQRVWSLPPNENTGKEVTALAWRPDGKILAFGLADTKRVILCDVEKPESLHSFSVDAPVTYMHWMEVTEESSVLTSFYNAEDESNLLLPKLPALPKNNFCDLFVIRLNALVLGGSSGFIEVYAYGMYKIATVAGSCLGLCFSSDLKSLSVITEVQVSPDSAPEITYFQLDTSLLSVYLPEVTRMARKFTHISTLLQYIKLSLTCMCEAWEEILMQMDSRLTKFVQEKNTTTSVQDEFMQLLLWGKASLELQALLMNQLTVKGLKKLGQSIESSYSSIQKLVISHLQSGSEALLYHLSELKGMASWKQKYESLGLDAAGIEGAITAVGSFILKANELLQVIDSSMKNFKAFFRWLYVAMLRMSEDHVLPELNKMTQKDITFVADFLTEHFNEAPDLYNRKGKYFNVERVGQYLKDEDDDLVSPPNTEGNQWFNFLQNSTHLKESPLLFPYYPQKSLHFVKRRMEGIIDQCLQKPAVSPVCYKRCDSTPRLFKLPFMWNDKSSNLHYVLFTMLESSLSKIYILRRHTDVSRSANNGLLSVDFGNFLNNSINESSDTGFYSCVDAHFYDDETITVVLKENAEQEGKERILAQLPLSSVYVDEDQEREFSLNSNKRLDEQSDDIPTRTVFLESQWRVLENMKAQYVAVNGIRKVSCVLSSNLRHVRVFEMDVEDDGEAEEEEEEEETTPIVGDEHEELTLSAVNHNSMCGGSAEADETEEHESSLDS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J178</accession>
    <name>A0A452J178_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Ran-specific GTPase-activating protein</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="8">Ran-binding protein 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="4">Plays a role in RAN-dependent nucleocytoplasmic transport. Alleviates the TNPO1-dependent inhibition of RAN GTPase activity and mediates the dissociation of RAN from proteins involved in transport into the nucleus. Induces a conformation change in the complex formed by XPO1 and RAN that triggers the release of the nuclear export signal of cargo proteins. Promotes the disassembly of the complex formed by RAN and importin beta. Promotes dissociation of RAN from a complex with KPNA2 and CSE1L. Required for normal mitotic spindle assembly and normal progress through mitosis via its effect on RAN. Does not increase the RAN GTPase activity by itself, but increases GTP hydrolysis mediated by RANGAP1. Inhibits RCC1-dependent exchange of RAN-bound GDP by GTP.</text>
    </comment>
    <comment type="subunit">
        <text evidence="6">Interacts with RAN (via C-terminus of GTP-bound form) but not with GDP-bound RAN. Identified in a complex composed of RAN, RANGAP1 and RANBP1. Identified in a complex that contains TNPO1, RAN and RANBP1. Identified in a complex that contains CSE1L, KPNA2, RAN and RANBP1. Identified in a complex with nucleotide-free RAN and RCC1.</text>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the RANBP1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J178"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038652.1">
        <property type="protein sequence ID" value="ENSGAGP00000034135.1"/>
        <property type="gene ID" value="ENSGAGG00000024254.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005643">
        <property type="term" value="C:nuclear pore"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006913">
        <property type="term" value="P:nucleocytoplasmic transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046604">
        <property type="term" value="P:positive regulation of mitotic centrosome separation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd13179">
        <property type="entry name" value="RanBD_RanBP1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000824">
        <property type="entry name" value="Ran-specific GTPase-activating protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000156">
        <property type="entry name" value="Ran_bind_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045255">
        <property type="entry name" value="RanBP1-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045256">
        <property type="entry name" value="RanBP1_RanBD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23138">
        <property type="entry name" value="RAN BINDING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23138:SF94">
        <property type="entry name" value="RAN BINDING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00638">
        <property type="entry name" value="Ran_BP1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00160">
        <property type="entry name" value="RanBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50196">
        <property type="entry name" value="RANBD1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0007">Acetylation</keyword>
    <keyword evidence="1" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="RanBD1" evidence="10">
        <location>
            <begin position="26"/>
            <end position="164"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="1"/>
            <end position="33"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="162"/>
            <end position="210"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="1"/>
            <end position="26"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00056716"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00061276"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00066150"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00067380"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00081162"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50196"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034135.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="210" mass="24444" checksum="A95E009FE47565F9" modified="2019-05-08" version="1">MAETKETHEEHDTSTENADDSNHDPQFEPIVSLPEQEIKTLEEDEEELFKMRAKLFRFASENDLPEWKERGTGDVKLLKHREKGTIRLLMRRDKTLKICANHYITPLMELKPNAGSDRAWVWNTHADFADESPKPELLAIRFLNAENAQKFKAKFEECRNEVDKKAKKAGTEKNDSADKVAEKLEELSVKEESKKSEKKEETKEKTEEKQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J179</accession>
    <name>A0A452J179_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">Anaphase-promoting complex subunit 4</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J179"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038673.1">
        <property type="protein sequence ID" value="ENSGAGP00000034150.1"/>
        <property type="gene ID" value="ENSGAGG00000024170.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005680">
        <property type="term" value="C:anaphase-promoting complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034399">
        <property type="term" value="C:nuclear periphery"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031145">
        <property type="term" value="P:anaphase-promoting complex-dependent catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070979">
        <property type="term" value="P:protein K11-linked ubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024789">
        <property type="entry name" value="APC4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056358">
        <property type="entry name" value="APC4_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024790">
        <property type="entry name" value="APC4_long_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13260">
        <property type="entry name" value="ANAPHASE PROMOTING COMPLEX SUBUNIT 4 APC4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13260:SF0">
        <property type="entry name" value="ANAPHASE-PROMOTING COMPLEX SUBUNIT 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12896">
        <property type="entry name" value="ANAPC4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23405">
        <property type="entry name" value="WD40_APC4_C-half"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="2" id="KW-0132">Cell division</keyword>
    <keyword evidence="3" id="KW-0498">Mitosis</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="Anaphase-promoting complex subunit 4 long" evidence="7">
        <location>
            <begin position="108"/>
            <end position="306"/>
        </location>
    </feature>
    <feature type="domain" description="Anaphase-promoting complex subunit 4 C-terminal half WD40" evidence="8">
        <location>
            <begin position="450"/>
            <end position="620"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="623"/>
            <end position="678"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="6">
        <location>
            <begin position="623"/>
            <end position="638"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00016067"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022776"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF12896"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF23405"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034150.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="678" mass="76833" checksum="C07B2E18E5128D30" modified="2019-05-08" version="1">MHWMEVTEESSVLTSFYNAEDESNLLLPKLPALPKNYSTTAKIFRLNALVLGGSSGFIEVYAYGMYKIATVTGVAGSCLGLCFSSDLKSLSVITEVQVSPDSAPEITYFQLDTSLLSVYLPEVTRMARKFTHISTLLQYIKLSLTCMCEAWEEILMQMDSRLTKFVQEKNTTTSVQDEFMQLLLWGKASLELQALLMNQLTVKGLKKLGQSIESSYSSIQKLVISHLQSGSEALLYHLSELKGMASWKQKYESLGLDAAGIEGAITAVGSFILKANELLQVIDSSMKNFKAFFRWLYVAMLRMSEDHVLPELNKMTQKDITFVADFLTEHFNEAPDLYNRKGKYFNVERVGQYLKDEDDDLVSPPNTEGNQWFNFLQNSTHLKESPLLFPYYPQKSLHFVKRRMEGIIDQCLQKPADVIGKSVHQAVCMCLYKGSKSEDSTPRLFKLPFMWNDKSSNLHYVLFTMLESSLSKIYILRRHTDVSRSANNGLLSVDFGNFLNNSINESSDTGFYSCVDAHFYDDETITVVLKENAEQEGKERILAQLPLSSVYVDEDQEREFSLNSNKRLDEQSDDIPTRTVFLESQWRVLENMKAQYVAVNGIRKVSCVLSSNLRHVRVFEMDVEDDGEAEEEEEEEETTPIVGDEHEELTLSAVNHNSMCGGSAEADETEEHESSLDS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J185</accession>
    <name>A0A452J185_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">Anaphase-promoting complex subunit 4</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J185"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038650.1">
        <property type="protein sequence ID" value="ENSGAGP00000034133.1"/>
        <property type="gene ID" value="ENSGAGG00000024170.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005680">
        <property type="term" value="C:anaphase-promoting complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034399">
        <property type="term" value="C:nuclear periphery"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031145">
        <property type="term" value="P:anaphase-promoting complex-dependent catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070979">
        <property type="term" value="P:protein K11-linked ubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030071">
        <property type="term" value="P:regulation of mitotic metaphase/anaphase transition"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024789">
        <property type="entry name" value="APC4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024977">
        <property type="entry name" value="Apc4-like_WD40_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056358">
        <property type="entry name" value="APC4_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024790">
        <property type="entry name" value="APC4_long_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017169">
        <property type="entry name" value="APC4_metazoa"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036322">
        <property type="entry name" value="WD40_repeat_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13260">
        <property type="entry name" value="ANAPHASE PROMOTING COMPLEX SUBUNIT 4 APC4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13260:SF0">
        <property type="entry name" value="ANAPHASE-PROMOTING COMPLEX SUBUNIT 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12896">
        <property type="entry name" value="ANAPC4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12894">
        <property type="entry name" value="ANAPC4_WD40"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23405">
        <property type="entry name" value="WD40_APC4_C-half"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF037303">
        <property type="entry name" value="APC4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50978">
        <property type="entry name" value="WD40 repeat-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="2" id="KW-0132">Cell division</keyword>
    <keyword evidence="7" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0498">Mitosis</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="4" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="391"/>
            <end position="411"/>
        </location>
    </feature>
    <feature type="domain" description="Anaphase-promoting complex subunit 4-like WD40" evidence="8">
        <location>
            <begin position="19"/>
            <end position="95"/>
        </location>
    </feature>
    <feature type="domain" description="Anaphase-promoting complex subunit 4 long" evidence="9">
        <location>
            <begin position="198"/>
            <end position="388"/>
        </location>
    </feature>
    <feature type="domain" description="Anaphase-promoting complex subunit 4 C-terminal half WD40" evidence="10">
        <location>
            <begin position="533"/>
            <end position="703"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="706"/>
            <end position="761"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="6">
        <location>
            <begin position="706"/>
            <end position="721"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00016067"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022776"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF12894"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF12896"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="Pfam" id="PF23405"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034133.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="761" mass="86190" checksum="CE5DFDE6717C3597" modified="2019-05-08" version="1">ILFVKNSPVYQWFSICGPQTHVLLHRLANFQRVWSLPPNENTGKEVTALAWRPDGKILAFGLADTKRVILCDVEKPESLHSFSVDAPVTYMHWMEVTEESSVLTSFYNAEDESNLLLPKLPALPKNNFCDLFVIRLNALVLGGSSGFIEVYAYGMYKIATVTGVAGSCLGLCFSSDLKSLSVITEVQVSPDSAPEITYFQLDTSLLSVYLPEVTRMARKFTHISTLLQYIKLSLTCMCEAWEEILMQMDSRLTKFVQEKNTTTSVQDEFMQLLLWGKASLELQALLMNQLTVKGLKKLGQSIESSYSSIQKLVISHLQSGSEALLYHLSELKGMASWKQKYESLGLDAAGIEGAITAVGSFILKANELLQVIDSSMKNFKAFFRWLYVGKLNYILSISYLLYILIICFGIMTQKDITFVADFLTEHFNEAPDLYNRKGKYFNVERVGQYLKDEDDDLVSPPNTEGNQWFNFLQNSTHLKESPLLFPYYPQKSLHFVKRRMEGIIDQCLQKPADVIGKSVHQAVCMCLYKGSKRWNDKSSNLHYVLFTMLESSLSKIYILRRHTDVSRSANNGLLSVDFGNFLNNSINESSDTGFYSCVDAHFYDDETITVVLKENAEQEGKERILAQLPLSSVYVDEDQEREFSLNSNKRLDEQSDDIPTRTVFLESQWRVLENMKAQYVAVNGIRKVSCVLSSNLRHVRVFEMDVEDDGEAEEEEEEEETTPIVGDEHEELTLSAVNHNSMCGGSAEADETEEHESSLDS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J187</accession>
    <name>A0A452J187_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12 14">Uracil-DNA glycosylase</fullName>
            <shortName evidence="12">UDG</shortName>
            <ecNumber evidence="12 14">3.2.2.27</ecNumber>
        </recommendedName>
    </protein>
    <gene>
        <name evidence="12" type="primary">UNG</name>
        <name evidence="12" type="synonym">UNG1</name>
    </gene>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="12 14">Excises uracil residues from the DNA which can arise as a result of misincorporation of dUMP residues by DNA polymerase or due to deamination of cytosine.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="12 14">
            <text>Hydrolyzes single-stranded DNA or mismatched double-stranded DNA and polynucleotides, releasing free uracil.</text>
            <dbReference type="EC" id="3.2.2.27"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="9">
            <text>a 2'-deoxyuridine in double-stranded DNA + H2O = a 2'-deoxyribose 5'-monophosphate in double-stranded DNA + uracil</text>
            <dbReference type="Rhea" id="RHEA:81455"/>
            <dbReference type="Rhea" id="RHEA-COMP:14231"/>
            <dbReference type="Rhea" id="RHEA-COMP:17071"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:17568"/>
            <dbReference type="ChEBI" id="CHEBI:133902"/>
            <dbReference type="ChEBI" id="CHEBI:139095"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="9">
            <dbReference type="Rhea" id="RHEA:81456"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="10">
            <text>a 2'-deoxyuridine in single-stranded DNA + H2O = a 2'-deoxyribose 5'-monophosphate in single-stranded DNA + uracil</text>
            <dbReference type="Rhea" id="RHEA:81459"/>
            <dbReference type="Rhea" id="RHEA-COMP:12847"/>
            <dbReference type="Rhea" id="RHEA-COMP:19684"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:17568"/>
            <dbReference type="ChEBI" id="CHEBI:133902"/>
            <dbReference type="ChEBI" id="CHEBI:139095"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="10">
            <dbReference type="Rhea" id="RHEA:81460"/>
        </physiologicalReaction>
    </comment>
    <comment type="subunit">
        <text evidence="11">Interacts with RPA2 subunit of the RPA trimer; this interaction mediates UNG2 recruitment to RPA-coated single-stranded DNA at stalled replication forks. Interacts with PCNA; this interaction mediates UNG2 recruitment to S-phase replication foci. Interacts (via N-terminus) with FAM72A.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="12">Mitochondrion</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="12">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="1 12 14">Belongs to the uracil-DNA glycosylase (UDG) superfamily. UNG family.</text>
    </comment>
    <dbReference type="EC" id="3.2.2.27" evidence="12 14"/>
    <dbReference type="AlphaFoldDB" id="A0A452J187"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038592.1">
        <property type="protein sequence ID" value="ENSGAGP00000034080.1"/>
        <property type="gene ID" value="ENSGAGG00000024220.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005654">
        <property type="term" value="C:nucleoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004844">
        <property type="term" value="F:uracil DNA N-glycosylase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097510">
        <property type="term" value="P:base-excision repair, AP site formation via deaminated base removal"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd10027">
        <property type="entry name" value="UDG-F1-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.470.10:FF:000004">
        <property type="entry name" value="Uracil-DNA glycosylase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.470.10">
        <property type="entry name" value="Uracil-DNA glycosylase-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="HAMAP" id="MF_00148">
        <property type="entry name" value="UDG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002043">
        <property type="entry name" value="UDG_fam1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018085">
        <property type="entry name" value="Ura-DNA_Glyclase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005122">
        <property type="entry name" value="Uracil-DNA_glycosylase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036895">
        <property type="entry name" value="Uracil-DNA_glycosylase-like_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF003588">
        <property type="entry name" value="PRK05254.1-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF003589">
        <property type="entry name" value="PRK05254.1-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF003591">
        <property type="entry name" value="PRK05254.1-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF003592">
        <property type="entry name" value="PRK05254.1-5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00628">
        <property type="entry name" value="ung"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11264">
        <property type="entry name" value="URACIL-DNA GLYCOSYLASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11264:SF0">
        <property type="entry name" value="URACIL-DNA GLYCOSYLASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03167">
        <property type="entry name" value="UDG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00986">
        <property type="entry name" value="UDG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00987">
        <property type="entry name" value="UreE_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52141">
        <property type="entry name" value="Uracil-DNA glycosylase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00130">
        <property type="entry name" value="U_DNA_GLYCOSYLASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0007">Acetylation</keyword>
    <keyword evidence="3 12 14" id="KW-0227">DNA damage</keyword>
    <keyword evidence="7 12 14" id="KW-0234">DNA repair</keyword>
    <keyword evidence="4 12 14" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="6 12" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="8 12" id="KW-0539">Nucleus</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Uracil-DNA glycosylase-like" evidence="16">
        <location>
            <begin position="127"/>
            <end position="288"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="1"/>
            <end position="61"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="12 13">
        <location>
            <position position="142"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00008184"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00052069"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00052828"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00064140"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="HAMAP-Rule" id="MF_03166"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10072"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="RuleBase" id="RU003780"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="SMART" id="SM00986"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034080.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="301" mass="33854" checksum="3C6AFD8C07AFACEC" modified="2019-05-08" version="1">MIGQKTLHSFFRAAPPRKRERSPEQGGDAEASFAKKLKLVGDESGKASPRSPPLSPEQQERIRRNKEAALQRLASRSVPAGFGEGWRRQLAGEFTKPYFSQLMAFVADERKRHTIYPPAQQVFTWTQMCDITDVKVVILGQDPYHGPGQAHGLCFSVQRPVPPPPSLENIYKELSADIEDFTHPGHGDLTGWAKQGVLLLNAVLTVRAHQPNSHKEKGWEQFTDAVVSWLNNNLDGLVFMLWGAYAQKKGSSIDRKRHHILQTVHPSPLSVHRGFFGCQHFSKTNELLKKSGKKPIDWRAL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J188</accession>
    <name>A0A452J188_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">Anaphase-promoting complex subunit 4</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J188"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038669.1">
        <property type="protein sequence ID" value="ENSGAGP00000034146.1"/>
        <property type="gene ID" value="ENSGAGG00000024170.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005680">
        <property type="term" value="C:anaphase-promoting complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034399">
        <property type="term" value="C:nuclear periphery"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031145">
        <property type="term" value="P:anaphase-promoting complex-dependent catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070979">
        <property type="term" value="P:protein K11-linked ubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024789">
        <property type="entry name" value="APC4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056358">
        <property type="entry name" value="APC4_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024790">
        <property type="entry name" value="APC4_long_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13260">
        <property type="entry name" value="ANAPHASE PROMOTING COMPLEX SUBUNIT 4 APC4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13260:SF0">
        <property type="entry name" value="ANAPHASE-PROMOTING COMPLEX SUBUNIT 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12896">
        <property type="entry name" value="ANAPC4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23405">
        <property type="entry name" value="WD40_APC4_C-half"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="2" id="KW-0132">Cell division</keyword>
    <keyword evidence="3" id="KW-0498">Mitosis</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="Anaphase-promoting complex subunit 4 long" evidence="6">
        <location>
            <begin position="123"/>
            <end position="321"/>
        </location>
    </feature>
    <feature type="domain" description="Anaphase-promoting complex subunit 4 C-terminal half WD40" evidence="7">
        <location>
            <begin position="446"/>
            <end position="605"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00016067"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022776"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="Pfam" id="PF12896"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF23405"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034146.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="635" mass="72126" checksum="E825DC677ACCD572" modified="2019-05-08" version="1">MHWMEVTEESSVLTSFYNAEDESNLLLPKLPALPKNYSTTAKIFSEEKSDEIMKLLGDVRLNALVLGGSSGFIEVYAYGMYKIATVTGVAGSCLGLCFSSDLKSLSVITEVQVSPDSAPEITYFQLDTSLLSVYLPEVTRMARKFTHISTLLQYIKLSLTCMCEAWEEILMQMDSRLTKFVQEKNTTTSVQDEFMQLLLWGKASLELQALLMNQLTVKGLKKLGQSIESSYSSIQKLVISHLQSGSEALLYHLSELKGMASWKQKYESLGLDAAGIEGAITAVGSFILKANELLQVIDSSMKNFKAFFRWLYVAMLRMSEDHVLPELNKMTQKDITFVADFLTEHFNEYLKDEDDDLVSPPNTEGNQWFNFLQNSTHLKESPLLFPYYPQKSLHFVKRRMEGIIDQCLQKPADVIGKSVHQAVCMCLYKGSKSEDSTPRLFKLPFMWNDKSSNLHYVLFTMLESSLSKIYILRRHTDVSRSANNGLLSVDFGNFLNNSINESSDTGFYSCVDAHFYDDETITVVLKENAEQEGKERILAQLPLSSVYVDEDQEREFSLNSNKRLDEQSDDIPTRTVFLESQWRVLENMKAQYVAVNGIRKVSCVILRKKEGDNLHLLSRVEKIIIIQLKYLLASV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J189</accession>
    <name>A0A452J189_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="19">Glutamate receptor</fullName>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="19">Receptor for glutamate that functions as a ligand-gated ion channel in the central nervous system and plays an important role in excitatory synaptic transmission. L-glutamate acts as an excitatory neurotransmitter at many synapses in the central nervous system.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>Ca(2+)(in) = Ca(2+)(out)</text>
            <dbReference type="Rhea" id="RHEA:29671"/>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="14 19">Postsynaptic cell membrane</location>
            <topology evidence="14 19">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="19">Belongs to the glutamate-gated ion channel (TC 1.A.10.1) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J189"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038670.1">
        <property type="protein sequence ID" value="ENSGAGP00000034147.1"/>
        <property type="gene ID" value="ENSGAGG00000024221.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015276">
        <property type="term" value="F:ligand-gated monoatomic ion channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0038023">
        <property type="term" value="F:signaling receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd06382">
        <property type="entry name" value="PBP1_iGluR_Kainate"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.2300:FF:000010">
        <property type="entry name" value="Glutamate ionotropic receptor kainate type subunit 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.190.10:FF:000240">
        <property type="entry name" value="Glutamate receptor ionotropic, kainate 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.70:FF:000010">
        <property type="entry name" value="Putative glutamate receptor ionotropic kainate 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.70">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.2300">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.190.10">
        <property type="entry name" value="Periplasmic binding protein-like II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001828">
        <property type="entry name" value="ANF_lig-bd_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019594">
        <property type="entry name" value="Glu/Gly-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001508">
        <property type="entry name" value="Iono_Glu_rcpt_met"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015683">
        <property type="entry name" value="Ionotropic_Glu_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001320">
        <property type="entry name" value="Iontro_rcpt_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028082">
        <property type="entry name" value="Peripla_BP_I"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18966">
        <property type="entry name" value="IONOTROPIC GLUTAMATE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01094">
        <property type="entry name" value="ANF_receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00060">
        <property type="entry name" value="Lig_chan"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10613">
        <property type="entry name" value="Lig_chan-Glu_bd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00177">
        <property type="entry name" value="NMDARECEPTOR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00079">
        <property type="entry name" value="PBPe"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53822">
        <property type="entry name" value="Periplasmic binding protein-like I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53850">
        <property type="entry name" value="Periplasmic binding protein-like II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 19" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="8 18" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13 19" id="KW-0407">Ion channel</keyword>
    <keyword evidence="6 19" id="KW-0406">Ion transport</keyword>
    <keyword evidence="12 19" id="KW-1071">Ligand-gated ion channel</keyword>
    <keyword evidence="7 19" id="KW-0472">Membrane</keyword>
    <keyword evidence="11 19" id="KW-0628">Postsynaptic cell membrane</keyword>
    <keyword evidence="9 19" id="KW-0675">Receptor</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="19" id="KW-0732">Signal</keyword>
    <keyword evidence="5 19" id="KW-0770">Synapse</keyword>
    <keyword evidence="3 19" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 19" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="1 19" id="KW-0813">Transport</keyword>
    <feature type="signal peptide" evidence="19">
        <location>
            <begin position="1"/>
            <end position="19"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5027143585" description="Glutamate receptor" evidence="19">
        <location>
            <begin position="20"/>
            <end position="847"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="505"/>
            <end position="524"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="581"/>
            <end position="603"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="762"/>
            <end position="786"/>
        </location>
    </feature>
    <feature type="domain" description="Ionotropic glutamate receptor C-terminal" evidence="20">
        <location>
            <begin position="404"/>
            <end position="743"/>
        </location>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="458"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="460"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="465"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="631"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="632"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="680"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="site" description="Crucial to convey clamshell closure to channel opening" evidence="17">
        <location>
            <position position="610"/>
        </location>
    </feature>
    <feature type="site" description="Interaction with the cone snail toxin Con-ikot-ikot" evidence="17">
        <location>
            <position position="637"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="18">
        <location>
            <begin position="68"/>
            <end position="319"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="18">
        <location>
            <begin position="692"/>
            <end position="746"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023257"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00034104"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00036634"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="RuleBase" id="RU367118"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="SMART" id="SM00079"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034147.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="847" mass="95377" checksum="8CD9C5F1EBFB18C9" modified="2019-05-08" version="1">MLLFYLLASTGGIFETVENEPVNVEELAFKFAVTNINRNKTLMPNTTLTYDIQRINLFDSFEASRRACDQLALGVAALFGPSHSSSVSAVQSICNALEVPHIQTRWKHPTVDNKDTFYINLYPDYAAISRAILDLVLYYNWKIVTVVYEDSTGLIRLQELIKAPSRYNIKIKIRQLPSGNKDARPLLKEMKKGKEFYVIFDCSHETAAEILKQILSMGMMTEYYHYFFTTLDLFALDLEPYRYSGVNMTGFRLLNIDNPQVTSVIEKWSMERLQAPPKPETGLLDGMMTTEAALMYDAVYMVAVASQRASQMTVSSLQCHRHKPWRFGPRFMNLIKEARWDGLTGHITFNKTDGLRKDFDLDIISLKEEGTEKIGVWNSNSGLNMTDSNKDRSTNITDSLANRTLIVTTILVSPAQLMVQSEETDSGTSSPVSNGSYSGEDGIFSNLCSQKADLAVAPLTITYVREKVIDFSKPFMTLGISILYRKPNGTNPGVFSFLNPLSPDIWMYVLLACLGVSCVLFVIARFTPYEWYNPHPCNPDSDVVENNFTLLNSFWFGVGALMQQGSELMPKALSTRIVGGIWWFFTLIIISSYTANLAAFLTVERMESPIDSADDLAKQTKIEYGAVRDGSTMTFFKKSKISTYEKMWAFMSSRQQTALVKNNDEGIQRVLTTDYALLMESTSIEYVTQRNCNLTQIGGLIDSKGYGVGTPIGSPYRDKITIAILQLQEEGKLHMMKEKWWRGNGCPEEDSKEASALGVENIGGIFIVLAAGLVLSVFVAIGEFIYKSRKNSDIEQCLSFNAIMEELGISLRHQKKLKKRSRTKGRSSFTSILTCHQRRTLRKETMA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J190</accession>
    <name>A0A452J190_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="19">Glutamate receptor</fullName>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="19">Receptor for glutamate that functions as a ligand-gated ion channel in the central nervous system and plays an important role in excitatory synaptic transmission. L-glutamate acts as an excitatory neurotransmitter at many synapses in the central nervous system.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>Ca(2+)(in) = Ca(2+)(out)</text>
            <dbReference type="Rhea" id="RHEA:29671"/>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="14 19">Postsynaptic cell membrane</location>
            <topology evidence="14 19">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="19">Belongs to the glutamate-gated ion channel (TC 1.A.10.1) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J190"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038655.1">
        <property type="protein sequence ID" value="ENSGAGP00000034138.1"/>
        <property type="gene ID" value="ENSGAGG00000024221.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015276">
        <property type="term" value="F:ligand-gated monoatomic ion channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0038023">
        <property type="term" value="F:signaling receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd06382">
        <property type="entry name" value="PBP1_iGluR_Kainate"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.2300:FF:000010">
        <property type="entry name" value="Glutamate ionotropic receptor kainate type subunit 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.190.10:FF:000240">
        <property type="entry name" value="Glutamate receptor ionotropic, kainate 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.70:FF:000010">
        <property type="entry name" value="Putative glutamate receptor ionotropic kainate 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.70">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.2300">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.190.10">
        <property type="entry name" value="Periplasmic binding protein-like II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001828">
        <property type="entry name" value="ANF_lig-bd_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019594">
        <property type="entry name" value="Glu/Gly-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001508">
        <property type="entry name" value="Iono_Glu_rcpt_met"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015683">
        <property type="entry name" value="Ionotropic_Glu_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001320">
        <property type="entry name" value="Iontro_rcpt_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028082">
        <property type="entry name" value="Peripla_BP_I"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18966">
        <property type="entry name" value="IONOTROPIC GLUTAMATE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01094">
        <property type="entry name" value="ANF_receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00060">
        <property type="entry name" value="Lig_chan"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10613">
        <property type="entry name" value="Lig_chan-Glu_bd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00177">
        <property type="entry name" value="NMDARECEPTOR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00079">
        <property type="entry name" value="PBPe"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53822">
        <property type="entry name" value="Periplasmic binding protein-like I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53850">
        <property type="entry name" value="Periplasmic binding protein-like II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 19" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="8 18" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13 19" id="KW-0407">Ion channel</keyword>
    <keyword evidence="6 19" id="KW-0406">Ion transport</keyword>
    <keyword evidence="12 19" id="KW-1071">Ligand-gated ion channel</keyword>
    <keyword evidence="7 19" id="KW-0472">Membrane</keyword>
    <keyword evidence="11 19" id="KW-0628">Postsynaptic cell membrane</keyword>
    <keyword evidence="9 19" id="KW-0675">Receptor</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 19" id="KW-0770">Synapse</keyword>
    <keyword evidence="3 19" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 19" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="1 19" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="510"/>
            <end position="529"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="586"/>
            <end position="608"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="767"/>
            <end position="791"/>
        </location>
    </feature>
    <feature type="domain" description="Ionotropic glutamate receptor C-terminal" evidence="20">
        <location>
            <begin position="421"/>
            <end position="748"/>
        </location>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="463"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="465"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="470"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="636"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="637"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="685"/>
        </location>
        <ligand>
            <name>L-glutamate</name>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
        </ligand>
    </feature>
    <feature type="site" description="Crucial to convey clamshell closure to channel opening" evidence="17">
        <location>
            <position position="615"/>
        </location>
    </feature>
    <feature type="site" description="Interaction with the cone snail toxin Con-ikot-ikot" evidence="17">
        <location>
            <position position="642"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="18">
        <location>
            <begin position="85"/>
            <end position="336"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="18">
        <location>
            <begin position="697"/>
            <end position="751"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023257"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00034104"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00036634"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="PIRSR" id="PIRSR601508-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="RuleBase" id="RU367118"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="SMART" id="SM00079"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034138.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="814" mass="91806" checksum="80B7F8F4C8D148A6" modified="2019-05-08" version="1">MDLSSMNLSNSFLKPLILLAITTSRTLRGIFETVENEPVNVEELAFKFAVTNINRNKTLMPNTTLTYDIQRINLFDSFEASRRACDQLALGVAALFGPSHSSSVSAVQSICNALEVPHIQTRWKHPTVDNKDTFYINLYPDYAAISRAILDLVLYYNWKIVTVVYEDSTGLIRLQELIKAPSRYNIKIKIRQLPSGNKDARPLLKEMKKGKEFYVIFDCSHETAAEILKQILSMGMMTEYYHYFFTTLDLFALDLEPYRYSGVNMTGFRLLNIDNPQVTSVIEKWSMERLQAPPKPETGLLDGMMTTEAALMYDAVYMVAVASQRASQMTVSSLQCHRHKPWRFGPRFMNLIKEARWDGLTGHITFNKTDGLRKDFDLDIISLKEEGTEKIGVWNSNSGLNMTDSNKDRSTNITDSLANRTLIVTTILVSPAVEGIKICLLNAKKMRIFSNLCSQKADLAVAPLTITYVREKVIDFSKPFMTLGISILYRKPNGTNPGVFSFLNPLSPDIWMYVLLACLGVSCVLFVIARFTPYEWYNPHPCNPDSDVVENNFTLLNSFWFGVGALMQQGSELMPKALSTRIVGGIWWFFTLIIISSYTANLAAFLTVERMESPIDSADDLAKQTKIEYGAVRDGSTMTFFKKSKISTYEKMWAFMSSRQQTALVKNNDEGIQRVLTTDYALLMESTSIEYVTQRNCNLTQIGGLIDSKGYGVGTPIGSPYRDKITIAILQLQEEGKLHMMKEKWWRGNGCPEEDSKEASALGVENIGGIFIVLAAGLVLSVFVAIGEFIYKSRKNSDIEQVSCLLPACFWFVF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIG2</accession>
    <name>A0A452GIG2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">NF-kappa-B-repressing factor</fullName>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">Enhances the ATPase activity of DHX15 by acting like a brace that tethers mobile sections of DHX15 together, stabilizing a functional conformation with high RNA affinity of DHX15. Involved in the constitutive silencing of the interferon beta promoter, independently of the virus-induced signals, and in the inhibition of the basal and cytokine-induced iNOS promoter activity. Also involved in the regulation of IL-8 transcription. May also act as a DNA-binding transcription regulator: interacts with a specific negative regulatory element (NRE) 5'-AATTCCTCTGA-3' to mediate transcriptional repression of certain NK-kappa-B responsive genes.</text>
    </comment>
    <comment type="subunit">
        <text evidence="11">Interacts with NF-kappa-B. Interacts with XRN2. Interacts (via G-patch domain) with DHX15; promoting the RNA helicase activity of DHX15.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
            <location evidence="1">Nucleolus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIG2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001616.1">
        <property type="protein sequence ID" value="ENSGAGP00000001429.1"/>
        <property type="gene ID" value="ENSGAGG00000001146.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000001626.1">
        <property type="protein sequence ID" value="ENSGAGP00000001438.1"/>
        <property type="gene ID" value="ENSGAGG00000001146.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005730">
        <property type="term" value="C:nucleolus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005654">
        <property type="term" value="C:nucleoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd02640">
        <property type="entry name" value="R3H_NRF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1370.50:FF:000004">
        <property type="entry name" value="NFKB repressing factor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.20:FF:000030">
        <property type="entry name" value="NFKB repressing factor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.20:FF:000034">
        <property type="entry name" value="NFKB repressing factor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.20">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1370.50">
        <property type="entry name" value="R3H-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014720">
        <property type="entry name" value="dsRBD_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058828">
        <property type="entry name" value="DSRM_CARF/NKRF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000467">
        <property type="entry name" value="G_patch_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001374">
        <property type="entry name" value="R3H_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036867">
        <property type="entry name" value="R3H_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034071">
        <property type="entry name" value="R3H_NRF"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16148:SF15">
        <property type="entry name" value="NF-KAPPA-B-REPRESSING FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16148">
        <property type="entry name" value="NF-KAPPA-B-REPRESSING FACTOR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00035">
        <property type="entry name" value="dsrm"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26535">
        <property type="entry name" value="DSRM_CARF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01585">
        <property type="entry name" value="G-patch"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01424">
        <property type="entry name" value="R3H"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00358">
        <property type="entry name" value="DSRM"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00443">
        <property type="entry name" value="G_patch"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00393">
        <property type="entry name" value="R3H"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54768">
        <property type="entry name" value="dsRNA-binding domain-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF82708">
        <property type="entry name" value="R3H domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50174">
        <property type="entry name" value="G_PATCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51061">
        <property type="entry name" value="R3H"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="9" id="KW-0539">Nucleus</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0678">Repressor</keyword>
    <keyword evidence="8" id="KW-0804">Transcription</keyword>
    <keyword evidence="6" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="5" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="G-patch" evidence="14">
        <location>
            <begin position="561"/>
            <end position="606"/>
        </location>
    </feature>
    <feature type="domain" description="R3H" evidence="15">
        <location>
            <begin position="610"/>
            <end position="674"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="1"/>
            <end position="93"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="13">
        <location>
            <begin position="67"/>
            <end position="76"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="13">
        <location>
            <begin position="78"/>
            <end position="93"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004604"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022491"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00055813"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00062272"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00068533"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50174"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS51061"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001438.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="700" mass="78266" checksum="F70D8175DA955D0B" modified="2019-05-08" version="1">MEKVLKMAEGIDIGEMPSFELVPPAKPPKRPYSPSCSPYPNPEPPKKVLPRFRVRPRFEPVHFVASSEKDERKEDPSDNQTQESNKDANTNSTAQSVENYVDFVFNSFNTQEADPQFSNSVGFGYASSQTTATSVVLNSMDTTQSGAPQLSTSSSVLQSSSDTFPQSVITAKQYFIDKLSAAIWKNLANPDANTGTDKINYTYLLTRSIQACKTNPEYIYAPLKEIAPADLPKNKKLPTDGFACEVRCQNVYLTTGYAGSKNGSRDRATELAVKLLQKSVEVKVAQRKFKHTYREDLIVCESGTCPQELPPALKQLDDFLAANKDFSSGQSGSDPLQGSSSSTKHWTNFVLTENASDAIGILNNSASFNKMSVEYKYDLMSNRSWRCRVYLQDHCLAEGYGSKKTSKHAAADEALKILQKMQSNVSPIKTTQVQKVGCSSRSSGKKKDLKDLVIYENSENPVCTLNDTAQFNKMTVEYVFERMTGMRWKCKVMLENEFIAEAVGVKKSVKHEAAEEAVKILKKTQPTVVNNLKKGTIEDVISRNEIRGRSAEEAFKQKIKEDNIGNQILRKMGWTGGGLGKAGEGIREPIAVKEQFKREGLGLDVERVNKIAKRDIEQIIRNYARSDSHVDLTFSTELTNDERKQIHQIAQKYGLKSKSHGQGHDRFLVVSRKRRKEDLLDQLKQEGHVGHYELIMPQAN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIG4</accession>
    <name>A0A452GIG4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Sodium-dependent multivitamin transporter</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="21">Solute carrier family 5 member 6</fullName>
        </alternativeName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="18">Sodium-dependent multivitamin transporter that mediates the electrogenic transport of pantothenate, biotin, lipoate and iodide. Functions as a Na(+)-coupled substrate symporter where the stoichiometry of Na(+):substrate is 2:1, creating an electrochemical Na(+) gradient used as driving force for substrate uptake. Required for biotin and pantothenate uptake in the intestine across the brush border membrane. Plays a role in the maintenance of intestinal mucosa integrity, by providing the gut mucosa with biotin. Contributes to the luminal uptake of biotin and pantothenate into the brain across the blood-brain barrier.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="16">
            <text>(R)-lipoate(out) + 2 Na(+)(out) = (R)-lipoate(in) + 2 Na(+)(in)</text>
            <dbReference type="Rhea" id="RHEA:73379"/>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
            <dbReference type="ChEBI" id="CHEBI:83088"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>(R)-pantothenate(out) + 2 Na(+)(out) = (R)-pantothenate(in) + 2 Na(+)(in)</text>
            <dbReference type="Rhea" id="RHEA:73371"/>
            <dbReference type="ChEBI" id="CHEBI:29032"/>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>biotin(out) + 2 Na(+)(out) = biotin(in) + 2 Na(+)(in)</text>
            <dbReference type="Rhea" id="RHEA:73375"/>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
            <dbReference type="ChEBI" id="CHEBI:57586"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="14">
            <text>iodide(out) + 2 Na(+)(out) = iodide(in) + 2 Na(+)(in)</text>
            <dbReference type="Rhea" id="RHEA:71207"/>
            <dbReference type="ChEBI" id="CHEBI:16382"/>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
        </reaction>
    </comment>
    <comment type="subunit">
        <text evidence="19">Interacts with PDZD11.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Apical cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 22">Belongs to the sodium:solute symporter (SSF) (TC 2.A.21) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIG4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001552.1">
        <property type="protein sequence ID" value="ENSGAGP00000001377.1"/>
        <property type="gene ID" value="ENSGAGG00000001097.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016324">
        <property type="term" value="C:apical plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015075">
        <property type="term" value="F:monoatomic ion transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015293">
        <property type="term" value="F:symporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090482">
        <property type="term" value="F:vitamin transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098660">
        <property type="term" value="P:inorganic ion transmembrane transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015887">
        <property type="term" value="P:pantothenate transmembrane transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006814">
        <property type="term" value="P:sodium ion transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1730.10:FF:000011">
        <property type="entry name" value="sodium-dependent multivitamin transporter isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1730.10">
        <property type="entry name" value="Sodium/glucose cotransporter"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038377">
        <property type="entry name" value="Na/Glc_symporter_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001734">
        <property type="entry name" value="Na/solute_symporter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051163">
        <property type="entry name" value="Sodium:Solute_Symporter_SSF"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00813">
        <property type="entry name" value="sss"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR42985">
        <property type="entry name" value="SODIUM-COUPLED MONOCARBOXYLATE TRANSPORTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR42985:SF2">
        <property type="entry name" value="SODIUM-DEPENDENT MULTIVITAMIN TRANSPORTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00474">
        <property type="entry name" value="SSF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50283">
        <property type="entry name" value="NA_SOLUT_SYMP_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="13" id="KW-0092">Biotin</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9" id="KW-0406">Ion transport</keyword>
    <keyword evidence="10 23" id="KW-0472">Membrane</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0915">Sodium</keyword>
    <keyword evidence="12" id="KW-0739">Sodium transport</keyword>
    <keyword evidence="6" id="KW-0769">Symport</keyword>
    <keyword evidence="5 23" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 23" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="23">
        <location>
            <begin position="6"/>
            <end position="27"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="23">
        <location>
            <begin position="47"/>
            <end position="67"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="23">
        <location>
            <begin position="79"/>
            <end position="101"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="23">
        <location>
            <begin position="122"/>
            <end position="143"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="23">
        <location>
            <begin position="155"/>
            <end position="175"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="23">
        <location>
            <begin position="187"/>
            <end position="206"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="23">
        <location>
            <begin position="234"/>
            <end position="252"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="23">
        <location>
            <begin position="273"/>
            <end position="298"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="23">
        <location>
            <begin position="333"/>
            <end position="356"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="23">
        <location>
            <begin position="376"/>
            <end position="396"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="23">
        <location>
            <begin position="402"/>
            <end position="427"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="23">
        <location>
            <begin position="434"/>
            <end position="455"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="23">
        <location>
            <begin position="508"/>
            <end position="531"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004424"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006434"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022847"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023053"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023201"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023267"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00036099"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00050243"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00050457"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00052729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00058802"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00061728"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00073170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00078601"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="RuleBase" id="RU362091"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001377.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="576" mass="62403" checksum="94683B5B3D69DF73" modified="2019-05-08" version="1">MAFTVADYTVFVLLLVLSSAIGLFYALSGGRQRTVQEFLLANRSMSFLPVALSLLATFQSAVAILGVPAEIYRFGTEYWFLGCCYFLGLLIPAHVFIPVFYRLHLTSTYEYLELRFNKAVRVCGTVTFIFQMVIYMGVVLYAPALALNAVTQFDLWGAVLTMGLVCTLYTTLGGLKAVIWTDVFQTLVMFAGQLAVIVVGTVKVGSMGRVWQLAADQGKISGIDLNPDPFERHTFWTLSVGGVFMMLSLYGVNQAQVQRYLSSRTEQEAVLSCYAVFPCQQVVLCLSCLTGLVMFAYYQEHPLTAAQSQASSDQLVLYFVMDVLKDLPGLPGLFVACLFSGSLSTISSAFNSLATVTMEDLIHPYFPGLRESRATLLSKLLALSYGLLCLGMAYLSSMMGPMLQAAISIFGMVGGPLLGLFCLGMFFPCANPTGAIAGLAAGLAMAFWIGIGSLVSNLRAAPSALPLSNITDFPQASSLSTAIATTLLTSTPAPKRLSGLQKFYNLSYMWYSAHNSTTVILVGLLVSLLTGPTKGATVNPRTIYPVLPRLLCCLPQRYRERLSCGVVCSAEGDQSR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIJ2</accession>
    <name>A0A452GIJ2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Rho GTPase-activating protein 39</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
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    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
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            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
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        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
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        <property type="gene ID" value="ENSGAGG00000001177.1"/>
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</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIJ7</accession>
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        <recommendedName>
            <fullName evidence="3">SCAN box domain-containing protein</fullName>
        </recommendedName>
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        <name type="scientific">Gopherus agassizii</name>
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            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
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            <taxon>Gopherus</taxon>
        </lineage>
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        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
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                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
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            <dbReference type="PubMed" id="28562605"/>
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        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
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        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
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        <scope>IDENTIFICATION</scope>
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        <citation type="submission" date="2025-09" db="UniProtKB">
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                <consortium name="Ensembl"/>
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        <scope>IDENTIFICATION</scope>
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            <end position="1916"/>
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            <end position="1242"/>
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        <location>
            <begin position="1199"/>
            <end position="1223"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="1224"/>
            <end position="1235"/>
        </location>
    </feature>
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    </evidence>
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    </evidence>
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    </evidence>
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    </evidence>
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        <source>
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        </source>
    </evidence>
    <sequence length="1925" mass="215605" checksum="F26267B1DAE213FD" modified="2019-05-08" version="1">ACARRPRYPVSRPQEAVGSGQSRRPTASGSGAVVLPAGMINPSVPIRNIRMKFAVLIGLIQVGEVSNRDIVETVLNLLVGGEFDLEMNFIIQDAESITCMSELLEHCDVTCQAEIWSMFTAILRKSVRNLQTSTEVGLIEQVLLKMSTVDDMIADLLVDMLGVLASYSITVKELKLLFSMLRGENGIWPRHAIKLLSVLNQMPQRHGPDTFFNFPGCSAAAIALPPIAKWPYQNGFTLNTWFRMDPLNNINVDKDKPYLYCFRTSKGVGYSAHFVGNCLIVTSLKSKGKGFQHCVKYDFQPRKWYMISIVHIYNRWRNSEIRCYVNGQLVSYGDMAWHVNTNDSYDKCFLGSSETADANRVFCGQLGAVYVFTEALNPAQIFAIHQLGPGYKSTFKFKSESDIHLAEHHKQVLYDGKLASSIAFTYNAKATDAQLCLESSPKENPSIFVHSPHALMLQDVKAIVTHSIHSAIHSIGGIQVLFPLFAHMSYRMYMYLSDFILVFKNKNVTFLLVLSKYCPSSNFMQSSSLTLKEESSSRVHITRAVLEQFLSFAKYLDGLSHGAPLLKQLCDHILFNPAIWIHTPAKVQLSLYTYLSAEFIGTATIYTTIRRVGTVLQLMHTLKYYYWVVNPADSSGITPKGLEGPRPSQKEIISLRAFMLLFLKQLILKDRGVKEDELQSILNYLLTMHEDENIHDVLQLLVALMSEHPASMIPAFDQRNGIRVIYKLLASKSESIWVQALKVLGYFLKHLGHKRKVEIMHTHSLFTLLGERLMLHTNTVTITTYNTLYEILTEQVCTQVVHKPHPEPDSTVKIQNPMILKVVATLLKNSTPSAELMEVRRLFLSDMIKLFSNSRENRRCLLQCSVWQDWMFSLGYINPKNSEEQKITEMVYNIFRILLYHAIKYEWGGWRVWVDTLSIAHSKVTYEAHKEYLAKMYEEYQSTISGLSSQTTGAKGGLEIREIEDLSQSQSPESETDYPVSTDTRDLLMATKVSDEVLGSSDRPGGGVHVEVHDLLVDIKAEKVEATEVKLDDMDLSPETLVTGENGALVEVESLLDNVYCAAVEKLQNNVHGSVGIIKKSEEKDNGPLITLADEKDEPSTNNTSFLFDKIPSQEEKLLPELSSNHITIPNVQETQMHLGVSDDLGLLAHMTGSVDITCTSSIIEDKEFKIHTTSDGMSNLSERELGSSSKGLEYAEMTATTLEPESSGSKSLPNVDTGSIISDTERSDDGKEAVQGRSVTQQDRDLRVDLGFRGMPMTEEQRRQFSPGPRTTMFRIPEFKWSPMHQRLLTDLLFALETDVHVWRSHSTKSVMDFVNSNENIIFVHNTIHLISQMVDNIIIACGGILPLLSAATSPTTELENIEVTQGMSAETAVTFLSRLMAMVDVLVFASSLNFSEIEAEKNMSSGGLMRQCLRLVCCVAVRNCLECRQRQRERVNKSSLISSKTQETLQGVTATSPLENVPGNLSPIKDPDRLLQDVDINRLRAVVFRDVDDSKQAQFLALAVVYFISVLMVSKYRDILEPQRETARCGSQQGLLTLLSPAVFLFWVCIEKPLELESSIPHTDSGIGEEQMPSILNGTDLETSAGPDAMSELFSTLSSEVKKSQESLTESPSEILKPAPSISSISQSKGMNVKEILKSLVAAPIEIAECGPDPIPYPDPALKREAQSILPMQFHSFDRYMAELIILYCCNAEIWSIYLLTCAVDSGSSSSSSSSSFVNGATSKNLPAVQTVAPMPEDSAENMSITAKLERALEKVAPLLREIFVDFAPFLSRTLLGSHGQELLIEGLVCMKSSTSVVELVMLLCSQEWQNSIQKNAGLAFIELINEGRSGLTTAVRAQRYHEWRYQEDKTPRSQLYDLIHLAQKWLQPESWSPEEILEVLVIDRYMRALPPDLCKWVGQNDPSTYDEMITLVERRMTARELT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIJ8</accession>
    <name>A0A452GIJ8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Ig-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GIJ8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001599.1">
        <property type="protein sequence ID" value="ENSGAGP00000001414.1"/>
        <property type="gene ID" value="ENSGAGG00000001083.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00099">
        <property type="entry name" value="IgV"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000604">
        <property type="entry name" value="immunoglobulin superfamily member 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000674">
        <property type="entry name" value="Immunoglobulin superfamily member 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000689">
        <property type="entry name" value="Immunoglobulin superfamily member 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000932">
        <property type="entry name" value="Immunoglobulin superfamily member 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000491">
        <property type="entry name" value="Immunoglobulin superfamily, member 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013106">
        <property type="entry name" value="Ig_V-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051102">
        <property type="entry name" value="IgSF_V-set/TM_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12207:SF21">
        <property type="entry name" value="IMMUNOGLOBULIN SUPERFAMILY MEMBER 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12207">
        <property type="entry name" value="V-SET AND TRANSMEMBRANE DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07686">
        <property type="entry name" value="V-set"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00406">
        <property type="entry name" value="IGv"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="7"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="4" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="5" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0732">Signal</keyword>
    <keyword evidence="5" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="12"/>
            <end position="30"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="979"/>
            <end position="1002"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="6">
        <location>
            <begin position="31"/>
            <end position="136"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="6">
        <location>
            <begin position="183"/>
            <end position="274"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="6">
        <location>
            <begin position="294"/>
            <end position="427"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="6">
        <location>
            <begin position="447"/>
            <end position="550"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="6">
        <location>
            <begin position="564"/>
            <end position="691"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="6">
        <location>
            <begin position="701"/>
            <end position="827"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="6">
        <location>
            <begin position="839"/>
            <end position="960"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001414.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1049" mass="118351" checksum="6ED8808BE2FC30D8" modified="2019-05-08" version="1">MLYGICSHGPGIVFVFSHLLPVFILFIAVIPDSLRVTSVPQTLNKVERDSLELKCEVSKSTAQHSHLSVSWYRQKVGEPPVEVISLSRDFVLRAGSTYAQRQATGNVRLDKVGETTSRLTVYNLQPSDQGEFYCEAAEWIQDPDKSWYAMTRKRSEGTTVNIQATDKEFSVRLDTEKRTYTVGEPVEFRCILEAQNVPERYFSISWAFNSSLIASLGPNAVPVLNSDFAQREALGQLKVAKESDNVFVLKIYRLRLEDSGKYNCRVTEREKTVTGDFIDKESKRPKNTPITVLPLKTSISVEMTNNASNVLEGESLRFGCNVHSGFGPQSHLSVTWQLVDKQNRRSDVVRLDRDGTLQPGPSYSERSSYGGIQMEQVRPGSFTLEIFNSVKADEGHYECHVTEWTRTQDGEWQMIGERHASMPVSITALEAGFAVTAISRTPGVTYNESFDLQCIIKPHYPSWVPVSVTWRFQPAGSTEFHDLVTFTRDGGVQWGDRYMGFRTRTAIEKAESSNNVRLSISRASDTEAGKYQCVAELWRKNYNSSWTRLADRTSNLLEIRVLRPVTKLQVSKSKRSITLVENRPIPLNCSVKSQTSPDSHFAVLWYVHKPADADGKLILKTTHSSAFEYGTYAEEEGLRGRLQFERSSSGGLFSLTVQRAEVRDSGSYYCHVEEWLLSPNHAWYKLAEEVSGRTEVTVKQPDNRLQVNQTHKNLTVLENQWVTLECAVTNRTSPASQLSVEWYVSKSGHPEKEAVARLNRQATLQYGDLATLSDLKGRLHLESPSPGLYLLTLQNVMVQDSGAYECRVEEWLLDPSNQWYKRAEDLSGLTTVTVKQPDPTLQVDTTISNLTVLEKESFPLDCSILSRSSQESRRVSHGMEEEEEREALLSVGLDAVFGQEAGRFEGRLRFQRLSSVLYRLTVLQAGAADAGNYSCRVEEWLPDPKRVWYRLAEEESGTITIHVQDTGSTLQSIICSNDALFYFVFFYPFPIFGILIITILLVRFKSRNSSKNSEGKNGVPLLWIKEPHLNYSPTCLEPPVLSIHPGTID</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIJ9</accession>
    <name>A0A452GIJ9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Isoaspartyl peptidase/L-asparaginase</fullName>
            <ecNumber evidence="3">3.4.19.5</ecNumber>
            <ecNumber evidence="4">3.5.1.1</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="9">Asparaginase-like protein 1</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="12">Beta-aspartyl-peptidase</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="10">Isoaspartyl dipeptidase</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="11">L-asparagine amidohydrolase</fullName>
        </alternativeName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>Cleavage of a beta-linked Asp residue from the N-terminus of a polypeptide.</text>
            <dbReference type="EC" id="3.4.19.5"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>L-asparagine + H2O = L-aspartate + NH4(+)</text>
            <dbReference type="Rhea" id="RHEA:21016"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:28938"/>
            <dbReference type="ChEBI" id="CHEBI:29991"/>
            <dbReference type="ChEBI" id="CHEBI:58048"/>
            <dbReference type="EC" id="3.5.1.1"/>
        </reaction>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the Ntn-hydrolase family.</text>
    </comment>
    <dbReference type="EC" id="3.4.19.5" evidence="3"/>
    <dbReference type="EC" id="3.5.1.1" evidence="4"/>
    <dbReference type="AlphaFoldDB" id="A0A452GIJ9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001701.1">
        <property type="protein sequence ID" value="ENSGAGP00000001491.1"/>
        <property type="gene ID" value="ENSGAGG00000001194.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000001705.1">
        <property type="protein sequence ID" value="ENSGAGP00000001495.1"/>
        <property type="gene ID" value="ENSGAGG00000001194.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004067">
        <property type="term" value="F:asparaginase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008798">
        <property type="term" value="F:beta-aspartyl-peptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033345">
        <property type="term" value="P:L-asparagine catabolic process via L-aspartate"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd04702">
        <property type="entry name" value="ASRGL1_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.60.20.30:FF:000001">
        <property type="entry name" value="Isoaspartyl peptidase/L-asparaginase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.60.20.30">
        <property type="entry name" value="(Glycosyl)asparaginase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033844">
        <property type="entry name" value="ASRGL1_meta"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029055">
        <property type="entry name" value="Ntn_hydrolases_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000246">
        <property type="entry name" value="Peptidase_T2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10188:SF41">
        <property type="entry name" value="ISOASPARTYL PEPTIDASE_L-ASPARAGINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10188">
        <property type="entry name" value="L-ASPARAGINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01112">
        <property type="entry name" value="Asparaginase_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56235">
        <property type="entry name" value="N-terminal nucleophile aminohydrolases (Ntn hydrolases)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0068">Autocatalytic cleavage</keyword>
    <keyword evidence="7" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="6" id="KW-0645">Protease</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <feature type="active site" description="Nucleophile" evidence="14">
        <location>
            <position position="166"/>
        </location>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <begin position="194"/>
            <end position="197"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <begin position="217"/>
            <end position="220"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="site" description="Cleavage; by autolysis" evidence="16">
        <location>
            <begin position="165"/>
            <end position="166"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010872"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012879"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00012920"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022280"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022813"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00029701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00029780"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00030414"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00030667"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00049366"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PIRSR" id="PIRSR600246-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PIRSR" id="PIRSR600246-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PIRSR" id="PIRSR600246-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001495.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="312" mass="32973" checksum="4C9743D1E42ACA78" modified="2019-05-08" version="1">MKPVIVVHGGAGRIFKDREEGARSGVTRAAMKGYSILKQGGNALDAVEEAVTSMEDDPHFNAGCGSVLNEKGEVEMDAIIMDGKSLASGAVSAVKCIANPIKLARLVMEKTEHMLLTDHGANLFAKAMGVPEIPGEKLITERSRERWSKNLEPDSNPQEFKKDPGTVGAVAIDNEGNVACATSTGGLTNKLVGRVGDSACIGSGGYADNHVGAISTTGHGESMMKVVLARLILYHMEQGMSPMEAADTALNYMKTRVGGLGGVIVVSNSGDWAARFSTKQMSWATVKDDQLQYGIYTGEIHTSSVAEALESK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIK6</accession>
    <name>A0A452GIK6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Protein kinase domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GIK6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001604.1">
        <property type="protein sequence ID" value="ENSGAGP00000001418.1"/>
        <property type="gene ID" value="ENSGAGG00000001107.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035556">
        <property type="term" value="P:intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043408">
        <property type="term" value="P:regulation of MAPK cascade"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050285">
        <property type="entry name" value="STE20_Ser/Thr_kinase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48015">
        <property type="entry name" value="SERINE/THREONINE-PROTEIN KINASE TAO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48015:SF23">
        <property type="entry name" value="SERINE_THREONINE-PROTEIN KINASE 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="1" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Protein kinase" evidence="2">
        <location>
            <begin position="27"/>
            <end position="89"/>
        </location>
    </feature>
    <feature type="binding site" evidence="1">
        <location>
            <position position="56"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001418.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="89" mass="10188" checksum="40E204885097A8DD" modified="2019-05-08" version="1">LLQPPAPLHKLKKLSEDSLTKQPEEVFDVLEKLGEGSYGSVFKAIHKESGQVVAIKQVPVESDLQEIIKEISIMQQCDRFFLFCVFFFF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIL6</accession>
    <name>A0A452GIL6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Armadillo repeat-containing protein 8</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIL6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001715.1">
        <property type="protein sequence ID" value="ENSGAGP00000001504.1"/>
        <property type="gene ID" value="ENSGAGG00000001203.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034657">
        <property type="term" value="C:GID complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043161">
        <property type="term" value="P:proteasome-mediated ubiquitin-dependent protein catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000061">
        <property type="entry name" value="armadillo repeat-containing protein 8 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000070">
        <property type="entry name" value="armadillo repeat-containing protein 8 isoform X1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000225">
        <property type="entry name" value="Armadillo"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038739">
        <property type="entry name" value="ARMC8/Vid28"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15651">
        <property type="entry name" value="ARMADILLO REPEAT-CONTAINING PROTEIN 8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15651:SF7">
        <property type="entry name" value="ARMADILLO REPEAT-CONTAINING PROTEIN 8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00185">
        <property type="entry name" value="ARM"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0539">Nucleus</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00013746"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001504.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="669" mass="75082" checksum="0EF9140CBE7F297B" modified="2019-05-08" version="1">MACVLETPLRMSVLSEVTASSRHYVDRLFDPDPQKVLQGVMLLYLLQQETSSTELRTECAVVLGSLAMGTENNVKSLLDCHIIPALLQGLLSPELKFIEACLRCLRTIFTSPVTPEELLYTDATVIPHLMALLSRSRYTQEYICQIFSHCCKLRHWHSCGFKSSPQPLLCANGPDHQTILFNHGAVQNIAHLLSSTSYKVRMQALKCFSVLAFENPQVSMTLVNVLVDGELLPQIFVKMLQRDKPIEMQLTSAKCLTYMCRAGAIQTDDNCIVLKALPSLVRMCSKERLLEERVEGAETLAYLIETDVELQRIASITDHLIAMLSDYFKYPSSVSAITDIKRLDHDLKHAHELRQAAFKLYASLGANDEDIRKKIIETENMMDRIVNGLSESSIKVRLSAVRCLHSLSRSVQQLRTSFQDHAVWKPLMKVLQNAPDEVLVVASSTLCNLLLEFSPSKEPILESGAIELLCSLTQSENLALRVNGIWALMNMAFQAEQKIKADILQGLSTEQLFQLLSDSDVNVLMKTLGLLRNLLSTRPHIDQIMSTHGKQIMQAVTLILEGEHNIEVKEQTLCILANIADGTTAKELIMTNDDILQKIKYYMSHSNAKLQLAAMFCISNLIWNEEEGSQERQDKLRDIGVVDILHKLSQSSDPNLCDKAKTALQQYLA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIL9</accession>
    <name>A0A452GIL9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Opioid growth factor receptor (OGFr) conserved domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the opioid growth factor receptor family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIL9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001624.1">
        <property type="protein sequence ID" value="ENSGAGP00000001436.1"/>
        <property type="gene ID" value="ENSGAGG00000001133.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140625">
        <property type="term" value="F:opioid growth factor receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006757">
        <property type="entry name" value="OGF_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039574">
        <property type="entry name" value="OGFr"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14015">
        <property type="entry name" value="OPIOID GROWTH FACTOR RECEPTOR OGFR ZETA-TYPE OPIOID RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14015:SF0">
        <property type="entry name" value="OPIOID GROWTH FACTOR RECEPTOR-LIKE PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04664">
        <property type="entry name" value="OGFr_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Opioid growth factor receptor (OGFr) conserved" evidence="3">
        <location>
            <begin position="112"/>
            <end position="315"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1"/>
            <end position="83"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="306"/>
            <end position="438"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="2">
        <location>
            <begin position="15"/>
            <end position="31"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="48"/>
            <end position="63"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="64"/>
            <end position="74"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="328"/>
            <end position="337"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="348"/>
            <end position="358"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="359"/>
            <end position="372"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="373"/>
            <end position="402"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="403"/>
            <end position="429"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00010365"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="Pfam" id="PF04664"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001436.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="455" mass="52437" checksum="3BD2FA7CE2EC8B0C" modified="2019-05-08" version="1">MGSLLSGVSVKEPATVEDCDSTWETDSEPELQDPAPEPPAPLPAASARPEETRQEGEEVEGGREAAAAAAPGPEQIDDATELTTKPKRSFYAARDLYKYRHQYPNFKDLRYQNDLCNLRFYKNKIPFKPDGVYIEEVLNKWKGDYEKLEHNHTYIQWLFPLREQGLNFYAKELTTYEIEEFKKTKEAIRRFLLAYKMMLEFFGIKLIDKTGNVARAANWQERFQHLNESQHNYLRITRILKSLGELGYESFKSPLVKFILHEALVEDTIPNIKQSALEYFVYTIRDRRERRQLLRFAQQRYTPSEHFIWGPPRKQKLEGNKTGKKPTSPVSSHNSYISKHKKWRDSKNTSITSSSSSKTAEEKKVELTRNGEKNNQCGTETNCEAAKQSNSEKNSDTESQNFKSEEVLSLSDKKENVSHSKKDKGKDGENLSQDCGNPTVTDTEDLILQCRHILC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIM2</accession>
    <name>A0A452GIM2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="35">Serine/threonine-protein kinase TOR</fullName>
            <ecNumber evidence="35">2.7.11.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="40 41">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="41" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="40" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="40" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="33">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="32">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="34">
            <text>L-tyrosyl-[protein] + ATP = O-phospho-L-tyrosyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10596"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.10.2"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="34">
            <dbReference type="Rhea" id="RHEA:10597"/>
        </physiologicalReaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Peripheral membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasmic vesicle</location>
            <location evidence="3">Phagosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="5">Endoplasmic reticulum membrane</location>
            <topology evidence="5">Peripheral membrane protein</topology>
            <orientation evidence="5">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Golgi apparatus membrane</location>
            <topology evidence="2">Peripheral membrane protein</topology>
            <orientation evidence="2">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="8">Lysosome membrane</location>
            <topology evidence="8">Peripheral membrane protein</topology>
            <orientation evidence="8">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="6">Microsome membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="7">Mitochondrion outer membrane</location>
            <topology evidence="7">Peripheral membrane protein</topology>
            <orientation evidence="7">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Nucleus</location>
            <location evidence="4">PML body</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="9 35">Belongs to the PI3/PI4-kinase family.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="35"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001714.1">
        <property type="protein sequence ID" value="ENSGAGP00000001503.1"/>
        <property type="gene ID" value="ENSGAGG00000000429.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000139">
        <property type="term" value="C:Golgi membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005765">
        <property type="term" value="C:lysosomal membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005741">
        <property type="term" value="C:mitochondrial outer membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045335">
        <property type="term" value="C:phagocytic vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016605">
        <property type="term" value="C:PML body"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031931">
        <property type="term" value="C:TORC1 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031932">
        <property type="term" value="C:TORC2 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004715">
        <property type="term" value="F:non-membrane spanning protein tyrosine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0106310">
        <property type="term" value="F:protein serine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="RHEA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0044877">
        <property type="term" value="F:protein-containing complex binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016242">
        <property type="term" value="P:negative regulation of macroautophagy"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009891">
        <property type="term" value="P:positive regulation of biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045597">
        <property type="term" value="P:positive regulation of cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051240">
        <property type="term" value="P:positive regulation of multicellular organismal process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008361">
        <property type="term" value="P:regulation of cell size"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019216">
        <property type="term" value="P:regulation of lipid metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051896">
        <property type="term" value="P:regulation of phosphatidylinositol 3-kinase/protein kinase B signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0038202">
        <property type="term" value="P:TORC1 signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05169">
        <property type="entry name" value="PIKKc_TOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.1070.11:FF:000074">
        <property type="entry name" value="DJ576K7.1 (FK506 binding protein 12-rapamycin associated protein 1)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.1070.11:FF:000058">
        <property type="entry name" value="MTOR isoform 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000060">
        <property type="entry name" value="Serine/threonine-protein kinase mTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000094">
        <property type="entry name" value="Serine/threonine-protein kinase mTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000532">
        <property type="entry name" value="Serine/threonine-protein kinase mTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.150:FF:000001">
        <property type="entry name" value="Serine/threonine-protein kinase TOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000083">
        <property type="entry name" value="Serine/threonine-protein kinase TOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1010.10:FF:000004">
        <property type="entry name" value="Serine/threonine-protein kinase TOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.150">
        <property type="entry name" value="FKBP12-rapamycin binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1070.11">
        <property type="entry name" value="Phosphatidylinositol 3-/4-kinase, catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1010.10">
        <property type="entry name" value="Phosphatidylinositol 3-kinase Catalytic Subunit, Chain A, domain 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.10">
        <property type="entry name" value="Tetratricopeptide repeat domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050517">
        <property type="entry name" value="DDR_Repair_Kinase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003152">
        <property type="entry name" value="FATC_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009076">
        <property type="entry name" value="FRB_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036738">
        <property type="entry name" value="FRB_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057564">
        <property type="entry name" value="HEAT_ATR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024585">
        <property type="entry name" value="mTOR_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000403">
        <property type="entry name" value="PI3/4_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036940">
        <property type="entry name" value="PI3/4_kinase_cat_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018936">
        <property type="entry name" value="PI3/4_kinase_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003151">
        <property type="entry name" value="PIK-rel_kinase_FAT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014009">
        <property type="entry name" value="PIK_FAT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026683">
        <property type="entry name" value="TOR_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011990">
        <property type="entry name" value="TPR-like_helical_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11139">
        <property type="entry name" value="ATAXIA TELANGIECTASIA MUTATED ATM -RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11139:SF9">
        <property type="entry name" value="SERINE_THREONINE-PROTEIN KINASE MTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02259">
        <property type="entry name" value="FAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02260">
        <property type="entry name" value="FATC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08771">
        <property type="entry name" value="FRB_dom"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23593">
        <property type="entry name" value="HEAT_ATR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11865">
        <property type="entry name" value="mTOR_dom"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00454">
        <property type="entry name" value="PI3_PI4_kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01346">
        <property type="entry name" value="DUF3385"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01343">
        <property type="entry name" value="FATC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00146">
        <property type="entry name" value="PI3Kc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01345">
        <property type="entry name" value="Rapamycin_bind"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47212">
        <property type="entry name" value="FKBP12-rapamycin-binding domain of FKBP-rapamycin-associated protein (FRAP)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51189">
        <property type="entry name" value="FAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51190">
        <property type="entry name" value="FATC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00915">
        <property type="entry name" value="PI3_4_KINASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00916">
        <property type="entry name" value="PI3_4_KINASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50290">
        <property type="entry name" value="PI3_4_KINASE_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="25" id="KW-0007">Acetylation</keyword>
    <keyword evidence="22 35" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="10" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="31" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="21" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="26" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="12" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="18 35" id="KW-0418">Kinase</keyword>
    <keyword evidence="29" id="KW-0458">Lysosome</keyword>
    <keyword evidence="28" id="KW-0472">Membrane</keyword>
    <keyword evidence="24" id="KW-0492">Microsome</keyword>
    <keyword evidence="27" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="19" id="KW-1000">Mitochondrion outer membrane</keyword>
    <keyword evidence="17 35" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="30" id="KW-0539">Nucleus</keyword>
    <keyword evidence="14" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="41" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="16" id="KW-0677">Repeat</keyword>
    <keyword evidence="13 35" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="20" id="KW-0802">TPR repeat</keyword>
    <keyword evidence="15 35" id="KW-0808">Transferase</keyword>
    <keyword evidence="23" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="FAT" evidence="38">
        <location>
            <begin position="1374"/>
            <end position="1960"/>
        </location>
    </feature>
    <feature type="domain" description="PI3K/PI4K catalytic" evidence="37">
        <location>
            <begin position="2134"/>
            <end position="2447"/>
        </location>
    </feature>
    <feature type="domain" description="FATC" evidence="39">
        <location>
            <begin position="2495"/>
            <end position="2527"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="36">
        <location>
            <begin position="1804"/>
            <end position="1844"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="36">
        <location>
            <begin position="1815"/>
            <end position="1838"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004202"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004255"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004262"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004322"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004397"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00004524"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00004570"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00004630"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00011031"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00022787"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00022803"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00022848"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="ARBA" id="ARBA00023228"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="31">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="32">
        <source>
            <dbReference type="ARBA" id="ARBA00047899"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="33">
        <source>
            <dbReference type="ARBA" id="ARBA00048679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="34">
        <source>
            <dbReference type="ARBA" id="ARBA00051942"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="35">
        <source>
            <dbReference type="RuleBase" id="RU364109"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="36">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="37">
        <source>
            <dbReference type="PROSITE" id="PS50290"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="38">
        <source>
            <dbReference type="PROSITE" id="PS51189"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="39">
        <source>
            <dbReference type="PROSITE" id="PS51190"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="40">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001503.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="41">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2527" mass="287268" checksum="2A01FE3A6626960C" modified="2019-05-08" version="1">RFFAGPAAAAGEETPGPNTVSILQQFASGLKSRNEEIRAKAAKDLQHYVTMELREMSQEESTRFYDQLNHHIFELVSSSDANERKGGILAIASLIGVEGGNATRIGRFANYLRNLLPSNDPVVMEMASKAIGRLALAGDTFTAEYVEFEVKRALEWLGADRNEGRRHAAVLVLRELAISVPTFFFQQVQPFFDNIFVAVWDPKQAIREGAVSALRACLILTTQREPKEMQKPQWYRHTYEEAEKGFDETLAKEKGMNRDDRIHGALLILNELVRISSMEGERLREEMEEITQQQMVHDKYCKDLLGFGTKPRHITPFTSFQSVQPPQSNALVGLLGYSPHQGIMGFGASPIPAKSALVESRCCRDLMEEKFDQVCLWVLKCRTSKNSLIQMTILNLLPRLAAFRPSAFTADQYLPDTMNHILSCVKKEKERTAAFQALGLLSVAVRSEFQAYLPKVLEIIKAALPPKDFAHKRQKSMQVDATVFTCISMLARAMGPCIQQDIKELLEPMLAVGLSPALTAVLYDLSRQIPQLKKDIQDGLLKMLSLVLMHKPLRHPGMPKGLAHQLASPSLTNIPEASDVGSITLALRTLGSFEFEGHSLTQFVRHCADHFLNSEHKEIRMEAARTCSRLLTPSIHLISGHAHVVSQTAVQVVADVLSKLLVVGITDPDPDIRYCVLASLDERFDAHLAQAENLQALFVALNDQVFEIRELAICTVGRLSSMNPAFVMPFLRKMLIQILTELEHSGVGRIKEQSARMLGHLVSNAPRLIRPYMEPILKALIVKLKDPDPDPNPGVINNVLATIGELAQVSGLEMRKWVDELFIIIMDMLQDSSLLAKRQVALWTLGQLVASTGYVVEPYRKYPTLLDVLLNFLKTEQNQGTRREAIRVLGLLGALDPYKHKVNIGMIDQSRDASAVSLSESKSSQDSCKQRETGERMCTLHFSMVALMRIFRDQSLSHHHTMVVQAITFIFKSLGLKCVQFLPQVMPTFLNVIRVCDGAIREFLFQQLGMLVSFVRSHIRPYMDEIVTLMRDFWVMNNSIQSTIILLIEQIVVALGGEFKLYLPQLIPHMLRVFMHDSSPGRNVSVKLLNAIQLFGANLDDYLHLLLPPIVKLFDAPDVPLVARKAALETVDRLTESLDFTDYASRIIHPIVRTLDLSPELRPTAMDTLSSLVFQLGKKYQIFIPMVNKVLMRHRINHQRYDVLICRIVKGYTLADEEEDPLIYQHRMLRSNQGEMLTSGPVETGPMKKLHVSTINLQKAWGAARRVSKDDWLEWLRRLSLELLKDSSSPSLRSCWALAQAYNPMARDLFNAAFVSCWSELNEDQQDELIRSIELALTSQDIAEVTQTLLNLAEFMEHSDKGPLPLRDDNGIVLLGERAAKCRAYAKALHYKELEFQKGPTPAILESLISINNKLQQPEAAAGVLEYAMKHFGELEIQATWYEKLHEWEDALVAYDKKMDTNKDDPELMLGRMRCLEALGEWGQLHQQCCEKWTLVNDETQAKMARMAAAAAWGLGQWDSMEEYTCMIPRDTHDGAFYRAVLALHQDLFSLAQQCIDKARDLLDAELTAMAGESYSRAYGAMVSCQMLSELEEVIQYKLVPERREIIRQIWWERLQGCQRIVEDWQRILMVRSLVVNPHEDMRTWLKYASLCGKSGRLALAHKTLVLLLGVDPSRQLDHPLPTVHPQVTYAYMKHMWKSARKIDAFQHMQHFVQTMQEQAQHAIATEDQQHKQELHKLMARCFLKLGEWQLNLQGINESTIPKVLQYYSASTEHDGNWYKAWHAWAVMNFEAVLHYKHQNQARDEKKKLRHASGASITSANTEGSNSDSEAESTENSPIPSPVQKKVTEDLSRTLLLYTVPAVQGFFRSISLSRGNNLQDTLRVLTLWFDYGHWPDVNEALVEGVKAIQIDTWLQVIPQLIARIDTPRPLVGRLIHQLLTDIGRYHPQALIYPLTVASKSTTTARHNAANKILKNMCEHSNTLVQQAMMVSEELIRVAILWHEMWHEGLEEASRLYFGERNVKGMFEVLEPLHAMMERGPQTLKETSFNQAYGRDLMEAQEWCRKYMKSGNVKDLTQAWDLYYHVFRRISKQLPQLTSLELQYVSPKLLMCRDLELAVPGTYDPNQPIIRIQSIAPSLQVITSKQRPRKLTLMGSNGHEFVFLLKGHEDLRQDERVMQLFGLVNTLLANDPTSLRKNLSIQRYAVIPLSTNSGLIGWVPHCDTLHALIRDYREKKKILLNIEHRIMLRMAPDYDHLTLMQKVEVFEHAVNNTAGDDLAKLLWLKSPSSEVWFDRRTNYTRSLAVMSMVGYILGLGDRHPSNLMLDRLSGKILHIDFGDCFEVAMTREKFPEKIPFRLTRMLTNAMEVTGLDGNYRITCHTVMEVLREHKDSVMAVLEAFVYDPLLNWRLMDTNTKGNKRSRTRTDSYSASQSVEMLDNVELGETAHKKTGTTVPESIHSFIGDGLVKPEALNKKAIQIINRVRDKLTGRDFSHDETLDVPTQVELLIKQATSHENLCQCYIGWCPFW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIM8</accession>
    <name>A0A452GIM8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">AP complex subunit sigma</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Part of the AP-3 complex, an adaptor-related complex which is not clathrin-associated. The complex is associated with the Golgi region as well as more peripheral structures. It facilitates the budding of vesicles from the Golgi membrane and may be directly involved in trafficking to lysosomes. In concert with the BLOC-1 complex, AP-3 is required to target cargos into vesicles assembled at cell bodies for delivery into neurites and nerve terminals.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasmic vesicle membrane</location>
            <topology evidence="1">Peripheral membrane protein</topology>
            <orientation evidence="1">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Golgi apparatus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 10">Belongs to the adaptor complexes small subunit family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIM8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001640.1">
        <property type="protein sequence ID" value="ENSGAGP00000001446.1"/>
        <property type="gene ID" value="ENSGAGG00000001155.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030123">
        <property type="term" value="C:AP-3 adaptor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030659">
        <property type="term" value="C:cytoplasmic vesicle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006896">
        <property type="term" value="P:Golgi to vacuole transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006886">
        <property type="term" value="P:intracellular protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="CDD" id="cd14834">
        <property type="entry name" value="AP3_sigma"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.450.60:FF:000001">
        <property type="entry name" value="AP complex subunit sigma"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.450.60">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016635">
        <property type="entry name" value="AP_complex_ssu"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022775">
        <property type="entry name" value="AP_mu_sigma_su"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027155">
        <property type="entry name" value="APS3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000804">
        <property type="entry name" value="Clathrin_sm-chain_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011012">
        <property type="entry name" value="Longin-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11753">
        <property type="entry name" value="ADAPTOR COMPLEXES SMALL SUBUNIT FAMILY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01217">
        <property type="entry name" value="Clat_adaptor_s"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF015588">
        <property type="entry name" value="AP_complex_sigma"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF64356">
        <property type="entry name" value="SNARE-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00989">
        <property type="entry name" value="CLAT_ADAPTOR_S"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="6" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="7 10" id="KW-0472">Membrane</keyword>
    <keyword evidence="5 10" id="KW-0653">Protein transport</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 10" id="KW-0813">Transport</keyword>
    <feature type="domain" description="AP complex mu/sigma subunit" evidence="11">
        <location>
            <begin position="1"/>
            <end position="147"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004555"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00006972"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00025605"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PIRNR" id="PIRNR015588"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF01217"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001446.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="162" mass="18847" checksum="4D78AEAB608F63FC" modified="2019-05-08" version="1">MIKAILIFNNHGKPRLSKFYQRYSEDTQQQIIRETFHLVSKRDENVCNFLEGGLLIGGSDNKLIYRHYATLYFVFCVDSSESELGILDLIQVFVETLDKCFENVCELDLIFHVDKVHNILSEMVMGGMVLETNMNEIVTQIDAQNKLEKSETFIFQSPRQDR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZI5</accession>
    <name>A0A452GZI5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">RRM domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZI5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008721.1">
        <property type="protein sequence ID" value="ENSGAGP00000007553.1"/>
        <property type="gene ID" value="ENSGAGG00000006037.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005847">
        <property type="term" value="C:mRNA cleavage and polyadenylation specificity factor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003729">
        <property type="term" value="F:mRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031124">
        <property type="term" value="P:mRNA 3'-end processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd12671">
        <property type="entry name" value="RRM_CSTF2_CSTF2T"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.20.70:FF:000001">
        <property type="entry name" value="Cleavage stimulation factor subunit 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.630:FF:000001">
        <property type="entry name" value="Cleavage stimulation factor subunit 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.330:FF:000061">
        <property type="entry name" value="cleavage stimulation factor subunit 2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.630">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.330">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.20.70">
        <property type="entry name" value="Transcription termination and cleavage factor, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025742">
        <property type="entry name" value="CSTF2_hinge"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026896">
        <property type="entry name" value="CSTF_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038192">
        <property type="entry name" value="CSTF_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012677">
        <property type="entry name" value="Nucleotide-bd_a/b_plait_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035979">
        <property type="entry name" value="RBD_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000504">
        <property type="entry name" value="RRM_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45735">
        <property type="entry name" value="CLEAVAGE STIMULATION FACTOR SUBUNIT 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45735:SF2">
        <property type="entry name" value="CLEAVAGE STIMULATION FACTOR SUBUNIT 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14327">
        <property type="entry name" value="CSTF2_hinge"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14304">
        <property type="entry name" value="CSTF_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00076">
        <property type="entry name" value="RRM_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00360">
        <property type="entry name" value="RRM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54928">
        <property type="entry name" value="RNA-binding domain, RBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50102">
        <property type="entry name" value="RRM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0507">mRNA processing</keyword>
    <keyword evidence="6" id="KW-0539">Nucleus</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="5 7" id="KW-0694">RNA-binding</keyword>
    <feature type="domain" description="RRM" evidence="9">
        <location>
            <begin position="16"/>
            <end position="94"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="198"/>
            <end position="520"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="8">
        <location>
            <begin position="199"/>
            <end position="222"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="223"/>
            <end position="242"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="8">
        <location>
            <begin position="279"/>
            <end position="295"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="354"/>
            <end position="364"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="366"/>
            <end position="379"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="401"/>
            <end position="480"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022664"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00176"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50102"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007553.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="606" mass="63686" checksum="D63D0D984CB87333" modified="2019-05-08" version="1">MAGLAVRDPAVDRSLRSVFVGNIPYEATEEQLKDIFSEVGPVVSFRLVYDRETGKPKGYGFCEYQDQETALSAMRNLNGREFSGRALRVDNAASEKNKEELKSLGTGAPIIESPYGDPINPEDAPESISRAVASLPPEQMFELMKQMKLCVQNSPQEARNMLLQNPQLAYALLQAQVVMRIVDPEIALKILHRQTSVPPLIPGNPQPVPGPGPGPGPGPGPGPNAQMNQQNPPSSQPQPIGGMHVNGAPPLMQPPMQGGVPAPGQMPASVAGPGPGPLAPGGGMQPQVGMPGGGPVPLERGQVPMPDPRAPMQRGPLPASGPPPRGLLGDAPNDPRGGTLLSVTGEVEPRGYLGPPHQGPPMHHMPGHDNRGPPPHEMRGGPMGEPRPLMGEPRGPLMDARGGRDPRGMEPRGMEPRGMEPRGMEPRGMEPRGMEPRGMEPRGMEPRGMEPRGMEPRGMEPRGMEPRGMDARGMEPRGMEPRGPGPNPRGPMPGGIQGPGPLNLGASGPQGPRQVPNMPGAGIQGGGMPGAGIQAGAMPGAVVQGAGQPGGFSPGQNQVTPQDHEKAALIMQVLQLTADQIAMLPPEQRQSILILKEQIQKSTGAP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZL1</accession>
    <name>A0A452GZL1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Nuclear receptor domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the nuclear hormone receptor family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZL1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008897.1">
        <property type="protein sequence ID" value="ENSGAGP00000007712.1"/>
        <property type="gene ID" value="ENSGAGG00000006024.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004879">
        <property type="term" value="F:nuclear receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000122">
        <property type="term" value="P:negative regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045944">
        <property type="term" value="P:positive regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.50.10:FF:000030">
        <property type="entry name" value="Nuclear Hormone Receptor family"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.50.10">
        <property type="entry name" value="Erythroid Transcription Factor GATA-1, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050234">
        <property type="entry name" value="Nuclear_hormone_rcpt_NR1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001628">
        <property type="entry name" value="Znf_hrmn_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013088">
        <property type="entry name" value="Znf_NHR/GATA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24082">
        <property type="entry name" value="NUCLEAR HORMONE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24082:SF231">
        <property type="entry name" value="NUCLEAR RECEPTOR SUBFAMILY 1 GROUP I MEMBER 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00105">
        <property type="entry name" value="zf-C4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00047">
        <property type="entry name" value="STROIDFINGER"/>
    </dbReference>
    <dbReference type="SMART" id="SM00399">
        <property type="entry name" value="ZnF_C4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57716">
        <property type="entry name" value="Glucocorticoid receptor-like (DNA-binding domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00031">
        <property type="entry name" value="NUCLEAR_REC_DBD_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51030">
        <property type="entry name" value="NUCLEAR_REC_DBD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0010">Activator</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="5" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13" id="KW-0539">Nucleus</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="12" id="KW-0675">Receptor</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="11" id="KW-0804">Transcription</keyword>
    <keyword evidence="8" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="7" id="KW-0862">Zinc</keyword>
    <keyword evidence="6" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="Nuclear receptor" evidence="14">
        <location>
            <begin position="6"/>
            <end position="81"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005993"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS51030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007712.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="117" mass="12665" checksum="8BCB377EEB6D12FE" modified="2019-05-08" version="1">MPEGEEKICAVCGDRATGYHFHVMTCEGCKGFFRRSISKGVCFTCPFTRNCTITKAKRRQCQACRLQKCLAVGMRKDSKVPPGRALGWAGASAVQGSRLARHRAGLVPCSGSWLAGP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZL3</accession>
    <name>A0A452GZL3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">Receptor-type tyrosine-protein phosphatase F</fullName>
            <ecNumber evidence="3">3.1.3.48</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>O-phospho-L-tyrosyl-[protein] + H2O = L-tyrosyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:10684"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="EC" id="3.1.3.48"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the protein-tyrosine phosphatase family. Receptor class 2A subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.48" evidence="3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008759.1">
        <property type="protein sequence ID" value="ENSGAGP00000007588.1"/>
        <property type="gene ID" value="ENSGAGG00000005093.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
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    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="CDD" id="cd05738">
        <property type="entry name" value="IgI_2_RPTP_IIa_LAR_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05739">
        <property type="entry name" value="IgI_3_RPTP_IIa_LAR_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000010">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000027">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X1"/>
        <property type="match status" value="1"/>
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    <dbReference type="FunFam" id="2.60.40.10:FF:000036">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X1"/>
        <property type="match status" value="1"/>
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    <dbReference type="FunFam" id="2.60.40.10:FF:000066">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000144">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000015">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000082">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000128">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000098">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase F isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000353">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase F isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013098">
        <property type="entry name" value="Ig_I-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050713">
        <property type="entry name" value="RTP_Phos/Ushers"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46957">
        <property type="entry name" value="CYTOKINE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46957:SF9">
        <property type="entry name" value="PROTEIN-TYROSINE-PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07679">
        <property type="entry name" value="I-set"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00014">
        <property type="entry name" value="FNTYPEIII"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="13" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="15" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="4" id="KW-0358">Heparin-binding</keyword>
    <keyword evidence="8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="16" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="12 20" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="14" id="KW-0675">Receptor</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="5 20" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 20" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="1168"/>
            <end position="1191"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="21">
        <location>
            <begin position="51"/>
            <end position="146"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="21">
        <location>
            <begin position="154"/>
            <end position="236"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="243"/>
            <end position="334"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="339"/>
            <end position="433"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="437"/>
            <end position="526"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="531"/>
            <end position="628"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="633"/>
            <end position="732"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="733"/>
            <end position="827"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="831"/>
            <end position="923"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="927"/>
            <end position="1011"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="613"/>
            <end position="634"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1333"/>
            <end position="1361"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004167"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010504"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00013064"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022674"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
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            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
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        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
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        </source>
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        </source>
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        </source>
    </evidence>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
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            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007588.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1398" mass="154431" checksum="4BC572F1F3944E4B" modified="2019-05-08" version="1">EQPRCWWRAWPGTLAGAGGRQQQPGTPLWCWSGICCMLFPVAEDHLPAGFPTIDMGPQLKVVEKARTATMLCAASGNPDPEISWFKDFLPVDTATSNGRIKQLRSGGTPIRGALQIENSEESDQGKYECVATNSAGTRYSAPANLYVRVRRVAPRFSIPPSNHEVMPGGSVNLTCVAVGAPMPYVKWMAGVEELTKEDEMPVGRNVLELNNIVQSASYTCVAMSSLGMIEATAQITVKALPKPPTEPLVTETTATSVTLTWDSGNSDPISYYVIQYKPKSWEEGLFQEVDGVATTRYSIGGLSPFSEYEFRIIAVNNIGRGPPSELVEARTGEQAPSSPPLKVQARMLSASTMLVQWEQPEEPNGQIRGYRVYYTTDPHLPLSMWQKHNTDDSHLTTVGSLITGTTYSIRVLAFTSVGDGPPSDIIQVKTQQGVPAQPADFQAEAESDTRILLTWLPALQERITKYELLYWEGEDGSQRKVNFDPSSSYAVEGLKPDTLYKFQLGARSELGLGVYTPVLEARTAQSTPSAPPQEVECVSTSSTTIRVSWVPPPAQSRNGVITQYSIAYQAVDAEDGAKHVVDGIGRERSSWEIKDLEKWTEYKVWVRAHTDVGPGPESSPVHVRTDEDVPSAPPRKVEVESLNSTAIRVSWKLPLSNKQHGQIRGYQVTYVKLENNEPRGQPVIKDVMLAEAQETIIGGLLPETTYSITVAAYTTKGDGARSKPKVVTTTGAVPGKPSMLISTTAMNTALIQWHPPKEMVGELLGYRLQYKRTDEEKMNIMDFEKNVHHYTVTSLHKGATYIFKLSAKNRAGPGEEFEKEITTTEDVPSGFPQNLRVVGLTTSTTEVAWDPPVLAERNGRITNYTVMYRDINSQQELTNITRETSIVLTNLRPDTTYDIKVRACNSKGVGPLSPSIQSRTMPVEQVFAKNFRVNAVMKTSVLLSWEVPDSYKSAVPFKILYNSQSVEVDGHSMKKLISDLQPDTEYSFVLMNRGSSAGGLQHRVSIHTAPDVLQSKPISTNKYMQEGKFTLTLPKVQTSAPVRWYYIVVVPVDQSTSSRTARWRTPDEMELDQLLEAISQGRRQRRQADRLKPYIAALVDMLPETFTLGDEKTYKGFYNKPLSEDLSYRCFVLASLEDGDTKRYAASPYSDEIVMEVSSAKQQDEPEMLWVMGPVLAVILIIIIVIAILLFKRKRANSPSSKDEHSIGLKDSLLAHSSDPVEMRRLNYQTPGMRDHPPIPVTDLADNIDRLKANDGLRFSQEYEVSCREGDRCAWHCPEPGVAGRRWAMAASPRGGSVDSFHAHSFSVHLPRGYPLSWLVPQLQPPAASLVTRRSCPQAPRPGLQRPGVPHXGSPYSCSADTPQSHLASPCPPTHTLALLIPLSPALPLPGLPLVLLC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZL4</accession>
    <name>A0A452GZL4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Kinesin-like protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="8 9">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Kinesin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZL4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008829.1">
        <property type="protein sequence ID" value="ENSGAGP00000007648.1"/>
        <property type="gene ID" value="ENSGAGG00000006119.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005874">
        <property type="term" value="C:microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008017">
        <property type="term" value="F:microtubule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003777">
        <property type="term" value="F:microtubule motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007018">
        <property type="term" value="P:microtubule-based movement"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056524">
        <property type="entry name" value="KIF6/9_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027640">
        <property type="entry name" value="Kinesin-like_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019821">
        <property type="entry name" value="Kinesin_motor_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001752">
        <property type="entry name" value="Kinesin_motor_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47968">
        <property type="entry name" value="CENTROMERE PROTEIN E"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47968:SF62">
        <property type="entry name" value="KINESIN FAMILY MEMBER 5A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23735">
        <property type="entry name" value="KIF9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00225">
        <property type="entry name" value="Kinesin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00380">
        <property type="entry name" value="KINESINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00129">
        <property type="entry name" value="KISc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00411">
        <property type="entry name" value="KINESIN_MOTOR_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50067">
        <property type="entry name" value="KINESIN_MOTOR_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 8 9" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="6" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="3 9" id="KW-0493">Microtubule</keyword>
    <keyword evidence="8 9" id="KW-0505">Motor protein</keyword>
    <keyword evidence="4 8 9" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Kinesin motor" evidence="11">
        <location>
            <begin position="7"/>
            <end position="263"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="419"/>
            <end position="505"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="453"/>
            <end position="464"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="466"/>
            <end position="498"/>
        </location>
    </feature>
    <feature type="binding site" evidence="8">
        <location>
            <begin position="94"/>
            <end position="101"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00283"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="RuleBase" id="RU000394"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50067"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007648.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="738" mass="83736" checksum="E882EFD0B6B3B0A6" modified="2019-05-08" version="1">MSAKERKVHAFVRVKPTANFPQDMIKYGPDNKSIDIYIRRDVKKGVINNKQTDWSFKLDGVLHNASQDSVYDAVARDLVSQALDGYNGTIMCYGQTGAGKTYTMTGATENYKHRGIIPRAIQQVFRAVEERIDQSITVRMSYLEIYNETLFDLLSTMPHDMSGDTHMAVVEYPQGVFVKGLSMHPASHEEDALNLLFEGETNRIIGEHTLNKNSSRSHCIFTIYIESHSRILSDAKYTASKINLVDLAGSERLGKTGLSSLRFATRMKWVTTEPAINEKYDPERMVKNLEKEIILLKQELAMHDSLVNRSLVNYDPLNEIQIAEINSQVRRYLEGATDDIDIMNIRQIQEVFKQFKVILSQQEQEVEARLRNKYTLIDKSDFAAITAAQKAGLVDAEGHLVGEVDGQGFGIGVVPLSKPGGKKVKARKAKEQLSPTTRKEGLGSPVLGKDLDSVSPSRSQLVSSTKDLDVKEGLLREQDAASIETHRSESVPKEESHRPSSPPSKLVAFEEFKNERGSEINRIFKENKAILSDRKKKASEITHRLNLIKQEMDITKEALGAQKKEREEQGEYVNEEGQIIIDEEEFLLVMKLKDLKKQYRSDFDELRDLKAEVQYCQHLVDQCRNRLISEFEIWYNESFLIPEDVQEAIKPGGNIRPGMIPMNRVLCLDEDEQERFDRLQGEVLPACPDSTSFYNAKMKTDQKHKYFRAMAALQQMHKKPGLITATVKNKPPAALHVI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZL6</accession>
    <name>A0A452GZL6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Myosin VB</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="4">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <comment type="caution">
        <text evidence="4">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZL6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008904.1">
        <property type="protein sequence ID" value="ENSGAGP00000007717.1"/>
        <property type="gene ID" value="ENSGAGG00000005542.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016459">
        <property type="term" value="C:myosin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003779">
        <property type="term" value="F:actin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003774">
        <property type="term" value="F:cytoskeletal motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051020">
        <property type="term" value="F:GTPase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd23767">
        <property type="entry name" value="IQCD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd15477">
        <property type="entry name" value="Myo5b_CBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.190:FF:000006">
        <property type="entry name" value="Myosin VA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.190:FF:000001">
        <property type="entry name" value="unconventional myosin-Va"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.190">
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.1590">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002710">
        <property type="entry name" value="Dilute_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000048">
        <property type="entry name" value="IQ_motif_EF-hand-BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058662">
        <property type="entry name" value="Myo5a/b_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037990">
        <property type="entry name" value="Myo5b_CBD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052072">
        <property type="entry name" value="Vascular_dev_regulator"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16027:SF6">
        <property type="entry name" value="DILUTE DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16027">
        <property type="entry name" value="DILUTE DOMAIN-CONTAINING PROTEIN YPR089W"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01843">
        <property type="entry name" value="DIL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00612">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25966">
        <property type="entry name" value="Myo5a"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01132">
        <property type="entry name" value="DIL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00015">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51126">
        <property type="entry name" value="DILUTE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="2" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="3 5" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-0505">Motor protein</keyword>
    <keyword evidence="4" id="KW-0518">Myosin</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0677">Repeat</keyword>
    <keyword evidence="7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="12"/>
            <end position="33"/>
        </location>
    </feature>
    <feature type="domain" description="Myosin motor" evidence="9">
        <location>
            <begin position="51"/>
            <end position="114"/>
        </location>
    </feature>
    <feature type="domain" description="Dilute" evidence="8">
        <location>
            <begin position="849"/>
            <end position="1126"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="447"/>
            <end position="477"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="5">
        <location>
            <begin position="501"/>
            <end position="616"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="5">
        <location>
            <begin position="663"/>
            <end position="763"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="456"/>
            <end position="476"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51126"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007717.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="12">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1171" mass="137032" checksum="9893288F9819EC8D" modified="2019-05-08" version="1">MAETIMVYIVKVSIVQFAAYPCFCGVVMVYQVAVGGAHIPVFLKLGSLLVWSYHDFFNRYRVLMKKREISKNDKKLICKTLLENLIKDPDKFQFGRTKIFFRAGQVAYLEKLRADKFRAATIMIQKTVRGWLQRVKYRRMRNAAITIQRYTRGHLARRLADHLRKTRAAIIFQKQYRMLRIFHAFQKIRKATITIQAFTRGMFVRKIYHKILLEHKATIIQKHARAWLARKQFLHFKCAAVVIQCYFRRMKARQELKKLKIEARSAQHLKKLNIGMENKVVQLQRKIDDQNKEYMHLNEQLSALTSAYATDVEKLKKELEQYRQNQGDTNQLVNLKKEIESLRLELEKSHSERKIIEDTYDKEKDLLRKRVSDLEEENALLKQEKEELNNRILCQSEDEFAQNTVKENLLTKKELEEERSRYQNLVKEYSRLEQRYDNLRDEMTIIKQTPGHRRNPSNQSSLESDSNCPSVSTSEIGDTEDAIQQVEDVGLDKAAMDMTLFLKLQKRVRELEQERKKLQTQLEKKKSQVIETKNEMDSDRDTDLAYNNLKRQELESENKKLKNELNELRKTIVDRVTQNNSSSDVQDSYTLLLNQLKSANEELEVRKEEVLILRAQIIKAAQQKDIGKNMDSNINTYTSWPNSDKHVDAEDAIEAYHGVCQTNRLLEAQLQDQRREHEEEVETLKTQMETLREEMEKQQQAFFQTLQLPPEAQVEFGIQQEIARLTNENLDLKELLEKLEKNEKKLKKQLKIYMKKVQDFEATQATAQTERRRHELTRQVTVQRKEKDFQGMLEYYKEDEPLLIRNLITDLKPQTVSATVPCLPAYILYMCIRHADYINDDQKVHSLLTSTINGIKKVLKKHNDDFQMTSFWLANTCRLLHCLKQYSGDTGFMTQNTAKQNEHCLKNFDLTEYRQVLSDLSIQIYQQLFKIAEGMLQPMIVSAMLESESIQGLSGVKPMGYRNRSSSMVDGDNSYSLDAVIRQLNTFHSIMCDQGLDPEIIQQVFKQLFYMINAVTLNNLLLRKDVCSWSTGMQLRYNISQLEEWLRGKNLQQSGAAETVEPLIQAAQLLQLKKKTSEDAEAICSLCTSLSTQQIVKILNLYTPVNEFEERVTVAFIRNIQANLQERNDPPQLLLDYKYMFPVLFPFSPSSITMDSIHIPASLNLEFLNEV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZL9</accession>
    <name>A0A452GZL9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">TRPM SLOG domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GZL9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008687.1">
        <property type="protein sequence ID" value="ENSGAGP00000007524.1"/>
        <property type="gene ID" value="ENSGAGG00000006018.1"/>
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        <property type="component" value="Unassembled WGS sequence"/>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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        <property type="term" value="F:calcium-activated cation channel activity"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0099604">
        <property type="term" value="F:ligand-gated calcium channel activity"/>
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        <property type="project" value="TreeGrafter"/>
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        <property type="entry name" value="TRPM"/>
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        <property type="entry name" value="TRPM_SLOG"/>
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    <dbReference type="PANTHER" id="PTHR13800:SF6">
        <property type="entry name" value="TRANSIENT RECEPTOR POTENTIAL CATION CHANNEL SUBFAMILY M MEMBER 4"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="TRANSIENT RECEPTOR POTENTIAL CATION CHANNEL, SUBFAMILY M, MEMBER 6"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="LSDAT_euk"/>
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    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="TRPM SLOG" evidence="2">
        <location>
            <begin position="5"/>
            <end position="119"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="135"/>
            <end position="155"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF18139"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007524.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="155" mass="16047" checksum="21C5A0B766C48C06" modified="2019-05-08" version="1">MGLRPFIRLSDSSDPALAYNLVTEHWKISPPNLVVSVVGGDGEAPVRAWLRDLLRKGLVKAAQSTGAWIMTGGLQAGIGQYVGEAVRDHGTASTSPCARVVAMGIAPWGVVANREALSCTGASPRPRCCTSCWSGARGGPSPPTTSLWSAPPAAP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZM1</accession>
    <name>A0A452GZM1_9SAUR</name>
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        <recommendedName>
            <fullName evidence="8">Chromo domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
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            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
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            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
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        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZM1"/>
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        <property type="gene ID" value="ENSGAGG00000006170.1"/>
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        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
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        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
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    </dbReference>
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        <property type="entry name" value="CSD_HP1alpha_Cbx5"/>
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        <property type="entry name" value="Chromobox protein homolog 1"/>
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    </dbReference>
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    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0832">Ubl conjugation</keyword>
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            <end position="152"/>
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        </location>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
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        </source>
    </evidence>
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        <source>
            <dbReference type="PROSITE" id="PS50013"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007722.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="264" mass="29700" checksum="B610770414D8B779" modified="2019-05-08" version="1">MCRRTREGAHWPRMRGAGAQAPARWGGCFGGRRALRRSGLSSEGLREGGVGAVPDPLRTQELGGLGALGGTSCAMGKKTKRTADSSSSEDEEEYVVEKVLDRRVVKGQVEYLLKWKGFSEEHNTWEPEKNLDCPELISEFMKKYKKMKEGENNKSREKSESTKRKSSLSNSAEDIKTKKKRESNDIARGFERGLEPEKIIGATDSCGDLMFLMKWKDTDEADLVLAKEANVKCPQIVIAFYEERLTWHAYPEDTENKEREAAKS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZM3</accession>
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        <recommendedName>
            <fullName evidence="3">Sm domain-containing protein</fullName>
        </recommendedName>
    </protein>
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        <name type="scientific">Gopherus agassizii</name>
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            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the ataxin-2 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZM3"/>
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        <property type="gene ID" value="ENSGAGG00000006078.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003729">
        <property type="term" value="F:mRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="P:stress granule assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045117">
        <property type="entry name" value="ATXN2-like"/>
    </dbReference>
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        <property type="entry name" value="PAM2_motif"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047575">
        <property type="entry name" value="Sm"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025852">
        <property type="entry name" value="SM_dom_ATX"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12854">
        <property type="entry name" value="ATAXIN 2-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12854:SF8">
        <property type="entry name" value="ATAXIN-2-LIKE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06741">
        <property type="entry name" value="LsmAD"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="PAM2"/>
        <property type="match status" value="1"/>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS52002">
        <property type="entry name" value="SM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
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        <location>
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            <end position="135"/>
        </location>
    </feature>
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    </feature>
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        </location>
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        <location>
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        </location>
    </feature>
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        </location>
    </feature>
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        <location>
            <begin position="963"/>
            <end position="972"/>
        </location>
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        <source>
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        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
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    </evidence>
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        <source>
            <dbReference type="PROSITE" id="PS52002"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007598.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="979" mass="105066" checksum="704C35ECED1D61B3" modified="2019-05-08" version="1">MGVAVLVGAQSRQWDSDIQLSLSQIHLGGGFPAVLSLPSHWAMCWGCWVFEGVYNNSRMLHFLTAVVGSTCDVKVKNGSTFEGIFKTLSSKFELAVDAVHRKTPDQLVGPKREDIVDTMVFKPTDVMLVHFRNVDFNYATKDKFTDSAIAMNSKVNGEHKEKVLQRWEGGDSNSDDYDLESDMSNGWDPNEMFKFNEENYGVKTTYDSSLSSYTVPLEKDNSEEFRQREARAAQLAREIESSPQYRLRIAMENDDGRTEEEKHSSVQRQVSGRDSPSLASREGKYVPLPQRVREGSRGGIRCSSSRGGRPGVGSMPPRGSAHHPDSNSSPAPEQRGINGGPSRMSPKSQRPIRGAKTMSSPSSRPVEVSAAPPAVGRMYPPRSPKSASATPASSQDSPVGSAVSAAPASPSEARTGLESSVSPNPPSPKVAAPVLATAVEGKEAPTSVAKDPGRTLEATGSCELSKAANKAGLQNEQKRTQLEELRKFGAQFKLQPSSSPETSLESFSARPKEPLEGKEKSLEPGICEGPEEAPPVSMVPKPNGTSLELLPESGKEEKGLCEVAEQQATNPPGKGEPEDKEEGPVSEQVKKSTLNPNAKEFNPSKTLLSVNKSTSTPTSPGPRAHSTPSIPVITAGQTGMYSPQYISYIPQIHMNPAVQAPQMYPYPVSSSVPGQQGKYRGAKGSLPPQRTDQHQPTSAPPIMQAAAAAGPPLVAATPYSSYISYNPQQFTGQPTMMQPMAHYPSQPVFAPMLQSNPRMMTSGSHPQAIVSSSTPQYPPSEQPTPQTLYATVHQSYPHHATQLHPHQPQPATTPTGNQQQAQHAAPSPVQHQAGQPPHLASAQQQQNLYHTATLTATPPSITPGPSAQSPQSSFPQQAVYAIHAHQQLQHSYTNMAHVTQAHVQTGITAAPPPHPGAPHPPQVMLLHPTQTHGGPPQGGVPQSGVPTLSASTPSPYTYIGHHQVQSHPSQQLPFHPPGN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZM8</accession>
    <name>A0A452GZM8_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="6">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GZM8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008777.1">
        <property type="protein sequence ID" value="ENSGAGP00000007603.1"/>
        <property type="gene ID" value="ENSGAGG00000006083.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060170">
        <property type="term" value="C:ciliary membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097730">
        <property type="term" value="C:non-motile cilium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005525">
        <property type="term" value="F:GTP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003924">
        <property type="term" value="F:GTPase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:1905515">
        <property type="term" value="P:non-motile cilium assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097500">
        <property type="term" value="P:receptor localization to non-motile cilium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000415">
        <property type="entry name" value="ADP-ribosylation factor-like GTPase 13B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051995">
        <property type="entry name" value="Ciliary_GTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006689">
        <property type="entry name" value="Small_GTPase_ARF/SAR"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46090:SF4">
        <property type="entry name" value="ADP RIBOSYLATION FACTOR LIKE GTPASE 13A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46090">
        <property type="entry name" value="ADP-RIBOSYLATION FACTOR-LIKE PROTEIN 13B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00025">
        <property type="entry name" value="Arf"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00328">
        <property type="entry name" value="SAR1GTPBP"/>
    </dbReference>
    <dbReference type="SMART" id="SM00177">
        <property type="entry name" value="ARF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00178">
        <property type="entry name" value="SAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51417">
        <property type="entry name" value="ARF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2 3" id="KW-0342">GTP-binding</keyword>
    <keyword evidence="4" id="KW-0460">Magnesium</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="1 3" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="219"/>
            <end position="241"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="220"/>
            <end position="232"/>
        </location>
    </feature>
    <feature type="binding site" evidence="3">
        <location>
            <begin position="42"/>
            <end position="49"/>
        </location>
        <ligand>
            <name>GTP</name>
            <dbReference type="ChEBI" id="CHEBI:37565"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="4">
        <location>
            <position position="49"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="4">
        <location>
            <position position="66"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="3">
        <location>
            <begin position="144"/>
            <end position="147"/>
        </location>
        <ligand>
            <name>GTP</name>
            <dbReference type="ChEBI" id="CHEBI:37565"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PIRSR" id="PIRSR606689-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PIRSR" id="PIRSR606689-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007603.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="241" mass="27365" checksum="AFC9C7FE6D1F8295" modified="2019-05-08" version="1">MKVQRLEVLPSLQNMFQLLSHCWSRLQAIQEPIRKVTLLVMGLDNTGKTSLITEIQRVLSCEVLPTTKPNQTELRVDQFEVSLVDLSRGQRSWGTWRNHYGDAHGIIFVLDSSNVLRMEEAKRTLGHLLAHPRISGKPLLLLANKQDKVDALLPCELIECLSLEKLVNENKSLCRIEPSSATKSLHKSQSRTILQGLRWLLHTIAINYSVLSARVQQDSLDQKAPREQEAPRRATRAHSQT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZN0</accession>
    <name>A0A452GZN0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Spondin-1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="11">F-spondin</fullName>
        </alternativeName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZN0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008834.1">
        <property type="protein sequence ID" value="ENSGAGP00000007652.1"/>
        <property type="gene ID" value="ENSGAGG00000005723.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031012">
        <property type="term" value="C:extracellular matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000026">
        <property type="entry name" value="Spondin 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000013">
        <property type="entry name" value="Spondin 1a"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000024">
        <property type="entry name" value="Spondin-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000034">
        <property type="entry name" value="Spondin-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000081">
        <property type="entry name" value="Spondin-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.2130">
        <property type="entry name" value="F-spondin domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.20.100.10">
        <property type="entry name" value="Thrombospondin type-1 (TSP1) repeat"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051418">
        <property type="entry name" value="Spondin/Thrombospondin_T1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009465">
        <property type="entry name" value="Spondin_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038678">
        <property type="entry name" value="Spondin_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000884">
        <property type="entry name" value="TSP1_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036383">
        <property type="entry name" value="TSP1_rpt_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044004">
        <property type="entry name" value="TSP1_spondin_dom"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF038123">
        <property type="entry name" value="NF038123_dom"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11311">
        <property type="entry name" value="SPONDIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11311:SF16">
        <property type="entry name" value="SPONDIN-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06468">
        <property type="entry name" value="Spond_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19028">
        <property type="entry name" value="TSP1_spondin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00090">
        <property type="entry name" value="TSP_1"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00209">
        <property type="entry name" value="TSP1"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF82895">
        <property type="entry name" value="TSP-1 type 1 repeat"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51020">
        <property type="entry name" value="SPONDIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50092">
        <property type="entry name" value="TSP1"/>
        <property type="match status" value="6"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="9" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="4" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="5" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0964">Secreted</keyword>
    <keyword evidence="6 13" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="13">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019578587" description="Spondin-1" evidence="13">
        <location>
            <begin position="23"/>
            <end position="527"/>
        </location>
    </feature>
    <feature type="domain" description="Spondin" evidence="14">
        <location>
            <begin position="1"/>
            <end position="108"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="451"/>
            <end position="470"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="451"/>
            <end position="466"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00019594"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00030964"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS51020"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007652.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="527" mass="59640" checksum="38056423A98C615E" modified="2019-05-08" version="1">AHQTITGNLIFLFLICRRAAPSAEFSVDRTRHLMSFLTMLGPSPDWNVGLSAEDLCTKDCGWVQKVIQDLIPWDAGTDSGVTYESPNKPTIPQEKIRPLTSLDHPQSPFYDPEGGSIAVVARVVIERIARKGEQCNIIPDNVDDIVADLVPEEKEEDDTPETCIYSNWSPWSACSSSTCDKGKRMRQRMLKAQLDLSVPCPDTQDFQPCMGPGCSDEDGSTCMMSEWITWSPCTVSCGMGMRSRERYVKQFPDDGSMCKVPTEETEKCIVNEECSPSSCLVTEWAEWNECSASCGIGMKKRHRMIKMTPADGSMCNTETTEVEKCMMPECHTIPCMLSPWSEWSDCSVTCGKGMRTRQRMLKSAAELGDCNEELEQVEKCMLPECPIDCELTEWSKWSECNKSCGKGHIIRTRMIKMEPQFGGAPCPETVQRKKCRIRKCLRGPSLEKRRWKEAREKRRSEHTKEDVEGEPYPGCKMRPWTAWSECTKLCGGGIQERFMTVKKRFKSTQFTSCKDKKEIRACNVHPC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZN5</accession>
    <name>A0A452GZN5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="27">CMP-N-acetylneuraminate-beta-galactosamide-alpha-2,3-sialyltransferase 4</fullName>
            <ecNumber evidence="16">2.4.3.2</ecNumber>
            <ecNumber evidence="17">2.4.3.4</ecNumber>
            <ecNumber evidence="23">2.4.3.6</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="28">Alpha 2,3-sialyltransferase IV</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="18">Gal-beta-1,3-GalNAc-alpha-2,3-sialyltransferase</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="30">Gal-beta-1,4-GlcNAc-alpha-2,3-sialyltransferase</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="29">N-acetyllactosaminide alpha-2,3-sialyltransferase</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="31">ST3Gal IV</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="32">Sialyltransferase 4C</fullName>
        </alternativeName>
    </protein>
    <organism evidence="35 36">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="36" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="35" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="35" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="14">
            <text>a beta-D-galactosyl-(1-&gt;3)-N-acetyl-alpha-D-galactosaminyl derivative + CMP-N-acetyl-beta-neuraminate = an N-acetyl-alpha-neuraminyl-(2-&gt;3)-beta-D-galactosyl-(1-&gt;3)-N-acetyl-alpha-D-galactosaminyl derivative + CMP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:21616"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57812"/>
            <dbReference type="ChEBI" id="CHEBI:60377"/>
            <dbReference type="ChEBI" id="CHEBI:133470"/>
            <dbReference type="ChEBI" id="CHEBI:139596"/>
            <dbReference type="EC" id="2.4.3.4"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="14">
            <dbReference type="Rhea" id="RHEA:21617"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="25">
            <text>a beta-D-galactosyl-(1-&gt;3)-N-acetyl-beta-D-galactosaminyl derivative + CMP-N-acetyl-beta-neuraminate = an N-acetyl-alpha-neuraminyl-(2-&gt;3)-beta-D-galactosyl-(1-&gt;3)-N-acetyl-beta-D-galactosaminyl derivative + CMP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:52380"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57812"/>
            <dbReference type="ChEBI" id="CHEBI:60377"/>
            <dbReference type="ChEBI" id="CHEBI:136588"/>
            <dbReference type="ChEBI" id="CHEBI:136589"/>
            <dbReference type="EC" id="2.4.3.2"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="25">
            <dbReference type="Rhea" id="RHEA:52381"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="22">
            <text>a beta-D-galactosyl-(1-&gt;4)-N-acetyl-beta-D-glucosaminyl derivative + CMP-N-acetyl-beta-neuraminate = an N-acetyl-alpha-neuraminyl-(2-&gt;3)-beta-D-galactosyl-(1-&gt;4)-N-acetyl-beta-D-glucosaminyl derivative + CMP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:52316"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57812"/>
            <dbReference type="ChEBI" id="CHEBI:60377"/>
            <dbReference type="ChEBI" id="CHEBI:133507"/>
            <dbReference type="ChEBI" id="CHEBI:136545"/>
            <dbReference type="EC" id="2.4.3.6"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="22">
            <dbReference type="Rhea" id="RHEA:52317"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>a ganglioside GA1 (d18:1(4E)) + CMP-N-acetyl-beta-neuraminate = a ganglioside GM1b (d18:1(4E)) + CMP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:47560"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:27938"/>
            <dbReference type="ChEBI" id="CHEBI:57812"/>
            <dbReference type="ChEBI" id="CHEBI:60377"/>
            <dbReference type="ChEBI" id="CHEBI:78568"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="19">
            <dbReference type="Rhea" id="RHEA:47561"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="20">
            <text>a ganglioside GM1 (d18:1(4E)) + CMP-N-acetyl-beta-neuraminate = a ganglioside GD1a (d18:1(4E)) + CMP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:18021"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57812"/>
            <dbReference type="ChEBI" id="CHEBI:60377"/>
            <dbReference type="ChEBI" id="CHEBI:77709"/>
            <dbReference type="ChEBI" id="CHEBI:78445"/>
            <dbReference type="EC" id="2.4.3.2"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="20">
            <dbReference type="Rhea" id="RHEA:18022"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="26">
            <text>a ganglioside GT1c (d18:1(4E)) + CMP-N-acetyl-beta-neuraminate = a ganglioside GQ1c (d18:1(4E)) + CMP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:47588"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57812"/>
            <dbReference type="ChEBI" id="CHEBI:60377"/>
            <dbReference type="ChEBI" id="CHEBI:87789"/>
            <dbReference type="ChEBI" id="CHEBI:87791"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="26">
            <dbReference type="Rhea" id="RHEA:47589"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="24">
            <text>a neolactoside nLc4Cer + CMP-N-acetyl-beta-neuraminate = a neolactoside IV(3)-alpha-NeuAc-nLc4Cer + CMP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:65432"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57812"/>
            <dbReference type="ChEBI" id="CHEBI:60377"/>
            <dbReference type="ChEBI" id="CHEBI:90376"/>
            <dbReference type="ChEBI" id="CHEBI:90390"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="24">
            <dbReference type="Rhea" id="RHEA:65433"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="21">
            <text>a neolactoside nLc4Cer(d18:1(4E)) + CMP-N-acetyl-beta-neuraminate = a neolactoside IV(3)-alpha-NeuAc-nLc4Cer(d18:1(4E)) + CMP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:18913"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:17006"/>
            <dbReference type="ChEBI" id="CHEBI:57812"/>
            <dbReference type="ChEBI" id="CHEBI:58665"/>
            <dbReference type="ChEBI" id="CHEBI:60377"/>
            <dbReference type="EC" id="2.4.3.6"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="21">
            <dbReference type="Rhea" id="RHEA:18914"/>
        </physiologicalReaction>
    </comment>
    <comment type="pathway">
        <text evidence="2">Protein modification; protein glycosylation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Golgi apparatus membrane</location>
            <topology evidence="1">Single-pass type II membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="15">Golgi apparatus</location>
            <location evidence="15">Golgi stack membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the glycosyltransferase 29 family.</text>
    </comment>
    <dbReference type="EC" id="2.4.3.2" evidence="16"/>
    <dbReference type="EC" id="2.4.3.4" evidence="17"/>
    <dbReference type="EC" id="2.4.3.6" evidence="23"/>
    <dbReference type="AlphaFoldDB" id="A0A452GZN5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008925.1">
        <property type="protein sequence ID" value="ENSGAGP00000007737.1"/>
        <property type="gene ID" value="ENSGAGG00000006181.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032580">
        <property type="term" value="C:Golgi cisterna membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000139">
        <property type="term" value="C:Golgi membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0047288">
        <property type="term" value="F:beta-D-galactosyl-(1-&gt;3)-N-acetyl-beta-D-galactosaminide alpha-2,3- sialyltransferase"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003836">
        <property type="term" value="F:beta-galactoside (CMP) alpha-2,3-sialyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008118">
        <property type="term" value="F:N-acetyllactosaminide alpha-2,3-sialyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009247">
        <property type="term" value="P:glycolipid biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd23982">
        <property type="entry name" value="GT29_ST3GAL4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1480.20:FF:000005">
        <property type="entry name" value="ST3 beta-galactoside alpha-2,3-sialyltransferase 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1480.20">
        <property type="entry name" value="Glycosyl transferase family 29"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001675">
        <property type="entry name" value="Glyco_trans_29"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051142">
        <property type="entry name" value="Glycosyltransferase_29"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038578">
        <property type="entry name" value="GT29-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012163">
        <property type="entry name" value="Sialyl_trans"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13713:SF57">
        <property type="entry name" value="CMP-N-ACETYLNEURAMINATE-BETA-GALACTOSAMIDE-ALPHA-2,3-SIALYLTRANSFERASE 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13713">
        <property type="entry name" value="SIALYLTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00777">
        <property type="entry name" value="Glyco_transf_29"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF005557">
        <property type="entry name" value="Sialyl_trans"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="4" id="KW-0328">Glycosyltransferase</keyword>
    <keyword evidence="9" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="10" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="11 34" id="KW-0472">Membrane</keyword>
    <keyword evidence="36" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0735">Signal-anchor</keyword>
    <keyword evidence="5" id="KW-0808">Transferase</keyword>
    <keyword evidence="6 34" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 34" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="34">
        <location>
            <begin position="7"/>
            <end position="24"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="33">
        <location>
            <begin position="115"/>
            <end position="268"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004323"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004922"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00006003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022676"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00036292"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00037859"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00039106"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00039107"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00042448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00043673"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00043773"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00048162"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00049294"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00049726"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00051076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00051975"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00052660"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="ARBA" id="ARBA00072813"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="ARBA" id="ARBA00075868"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="ARBA" id="ARBA00076295"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="ARBA" id="ARBA00076532"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="31">
        <source>
            <dbReference type="ARBA" id="ARBA00081234"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="32">
        <source>
            <dbReference type="ARBA" id="ARBA00082801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="33">
        <source>
            <dbReference type="PIRSR" id="PIRSR005557-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="34">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="35">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007737.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="36">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="328" mass="38014" checksum="E47299AA5F4A60A6" modified="2019-05-08" version="1">MINKFRWKVILGLAFFLVMAWYYISREESFSFPVQDVRMACPQGEMEKKVTLLIGNYTRDHPMFLHLNDYFWVKNRSPYELPYGTKGSEDILLRVLAITSYSVPESIQSMKCRRCAVVGNGHRLRNSSMGEAINKYDVVIRLNNAPVHGYEADVGSKTTMRLFYPESAHFDPRRENNPDTLLVLVPFKPMDFQWLEMILNDKKRIRKGFWKQPPLIWDANPEQVRILNPYFMEITAAKLLNLSMKQPRKIKQKPTTGLLAITLALHFCDLVHIAGFGYPNSANKKQSIHYYEQITLKSMAASEHNVSHEALAIKRMLELGVVKNLTHF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLJ8</accession>
    <name>A0A452HLJ8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Serine/threonine-protein phosphatase</fullName>
            <ecNumber evidence="12">3.1.3.16</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="10">
            <text>O-phospho-L-seryl-[protein] + H2O = L-seryl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:20629"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="EC" id="3.1.3.16"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>O-phospho-L-threonyl-[protein] + H2O = L-threonyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:47004"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="EC" id="3.1.3.16"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="2">
            <name>Fe(3+)</name>
            <dbReference type="ChEBI" id="CHEBI:29034"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the PPP phosphatase family. PP-2B subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.16" evidence="12"/>
    <dbReference type="AlphaFoldDB" id="A0A452HLJ8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018057.1">
        <property type="protein sequence ID" value="ENSGAGP00000015812.1"/>
        <property type="gene ID" value="ENSGAGG00000011890.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033192">
        <property type="term" value="F:calmodulin-dependent protein phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097720">
        <property type="term" value="P:calcineurin-mediated signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd07416">
        <property type="entry name" value="MPP_PP2B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.60.21.10:FF:000002">
        <property type="entry name" value="Serine/threonine-protein phosphatase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.60.21.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004843">
        <property type="entry name" value="Calcineurin-like_PHP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029052">
        <property type="entry name" value="Metallo-depent_PP-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041751">
        <property type="entry name" value="MPP_PP2B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043360">
        <property type="entry name" value="PP2B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006186">
        <property type="entry name" value="Ser/Thr-sp_prot-phosphatase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45673">
        <property type="entry name" value="SERINE/THREONINE-PROTEIN PHOSPHATASE 2B CATALYTIC SUBUNIT 1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00149">
        <property type="entry name" value="Metallophos"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00114">
        <property type="entry name" value="STPHPHTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00156">
        <property type="entry name" value="PP2Ac"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56300">
        <property type="entry name" value="Metallo-dependent phosphatases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00125">
        <property type="entry name" value="SER_THR_PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="5 12" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="9" id="KW-0408">Iron</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="Serine/threonine specific protein phosphatases" evidence="14">
        <location>
            <begin position="138"/>
            <end position="143"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="477"/>
            <end position="506"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="13">
        <location>
            <begin position="490"/>
            <end position="506"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00001965"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00009905"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00047761"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00048336"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="RuleBase" id="RU004273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS00125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015812.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="506" mass="57232" checksum="22259485D7700C44" modified="2019-05-08" version="1">MCIPFIQGATAVPFPPTHRLTSEEVFDIDGKPRVDILKNHLVKEGRVDEEIALRIINEGAAILRREKTMIEVEAPITVCGDIHGQFFDLMKLFEVGGSPANTRYLFLGDYVDRGYFSIECVLYLWVLKILYPNTLFLLRGNHECRHLTEYFTFKQECKIKYSERVYDACMEAFDCLPLAALLNQQFLCVHGGLSPEINTLDDIKRLDRFKEPPAFGPMCDLLWSDPSEDFGNEKSQEHFSHNTVRGCSYFYSYPAVCEFLQNNNLLSIIRAHEAQDAGYRMYRKSQTTGFPSLITIFSAPNYLDVYNNKAAVLKYENNVMNIRQFNCSPHPYWLPNFMDVFTWSLPFVGEKVTEMLVNVLSICSDDELMTEGEDQFDGTAAARKEIIRNKIRAIGKMARVFSVLREESESVLTLKGLTPTGMLPSGVLAGGRQTLQSATVEAIEAEKAIRGFSPQHKICSFEEAKGLDRINERMPPRKDALQQDGINSVNTTNANGNNGTGNNNVQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLM0</accession>
    <name>A0A452HLM0_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="7">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLM0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018148.1">
        <property type="protein sequence ID" value="ENSGAGP00000015882.1"/>
        <property type="gene ID" value="ENSGAGG00000011948.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043998">
        <property type="term" value="F:histone H2A acetyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010485">
        <property type="term" value="F:histone H4 acetyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:1990189">
        <property type="term" value="F:protein N-terminal-serine acetyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.630.30:FF:000346">
        <property type="entry name" value="N(alpha)-acetyltransferase 40, NatD catalytic subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.630.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016181">
        <property type="entry name" value="Acyl_CoA_acyltransferase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039949">
        <property type="entry name" value="NAA40"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR20531">
        <property type="entry name" value="N-ALPHA-ACETYLTRANSFERASE 40"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR20531:SF1">
        <property type="entry name" value="N-ALPHA-ACETYLTRANSFERASE 40"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55729">
        <property type="entry name" value="Acyl-CoA N-acyltransferases (Nat)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0012">Acyltransferase</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="5" id="KW-0539">Nucleus</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0808">Transferase</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023315"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015882.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="167" mass="19500" checksum="8938E47F5D2A6D4E" modified="2019-05-08" version="1">MGRKSSKAKEKKQKRLEERAAMDAVCAKVEAANKLGDPLEAFPVFKKYDRNGLNVSIECKRVSGMNQATVDWAFELTKTNMQTLYEQSEWGWKDREKREEMTDDRAWYLIAWEDSSIPVAFSHFRFDVECGDEVLYCYEVQLESKVRRKGLGKFLIQILQLMANRDL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLM5</accession>
    <name>A0A452HLM5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Receptor-type tyrosine-protein phosphatase</fullName>
            <ecNumber evidence="16">3.1.3.48</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">Tyrosine protein phosphatase which is involved in integrin-mediated focal adhesion formation. Following integrin engagement, specifically recruits BCAR3, BCAR1 and CRK to focal adhesions thereby promoting SRC-mediated phosphorylation of BRAC1 and the subsequent activation of PAK and small GTPase RAC1 and CDC42.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="24">
            <text>O-phospho-L-tyrosyl-[protein] + H2O = L-tyrosyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:10684"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="EC" id="3.1.3.48"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell junction</location>
            <location evidence="1">Focal adhesion</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 16">Belongs to the protein-tyrosine phosphatase family. Receptor class 4 subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.48" evidence="16"/>
    <dbReference type="AlphaFoldDB" id="A0A452HLM5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018153.1">
        <property type="protein sequence ID" value="ENSGAGP00000015887.1"/>
        <property type="gene ID" value="ENSGAGG00000010212.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005925">
        <property type="term" value="C:focal adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="CDD" id="cd14621">
        <property type="entry name" value="R-PTPc-A-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd14623">
        <property type="entry name" value="R-PTPc-A-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000007">
        <property type="entry name" value="Receptor-type tyrosine-protein phosphatase alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000021">
        <property type="entry name" value="Receptor-type tyrosine-protein phosphatase alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050348">
        <property type="entry name" value="Protein-Tyr_Phosphatase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000242">
        <property type="entry name" value="PTP_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016130">
        <property type="entry name" value="Tyr_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003595">
        <property type="entry name" value="Tyr_Pase_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016336">
        <property type="entry name" value="Tyr_Pase_rcpt_a/e-type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19134">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19134:SF433">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00102">
        <property type="entry name" value="Y_phosphatase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF002006">
        <property type="entry name" value="PTPR_alpha_epsilon"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00700">
        <property type="entry name" value="PRTYPHPHTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00194">
        <property type="entry name" value="PTPc"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00404">
        <property type="entry name" value="PTPc_motif"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00383">
        <property type="entry name" value="TYR_PHOSPHATASE_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50055">
        <property type="entry name" value="TYR_PHOSPHATASE_PTP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="11" id="KW-0965">Cell junction</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="14" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9 16" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="13" id="KW-0472">Membrane</keyword>
    <keyword evidence="5 19" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="10 16" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
    <keyword evidence="6" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="12" id="KW-1133">Transmembrane helix</keyword>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="20">
        <location>
            <begin position="83"/>
            <end position="343"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="21">
        <location>
            <begin position="263"/>
            <end position="334"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="20">
        <location>
            <begin position="375"/>
            <end position="633"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="21">
        <location>
            <begin position="549"/>
            <end position="624"/>
        </location>
    </feature>
    <feature type="active site" description="Phosphocysteine intermediate" evidence="17">
        <location>
            <position position="284"/>
        </location>
    </feature>
    <feature type="active site" description="Phosphocysteine intermediate" evidence="17">
        <location>
            <position position="574"/>
        </location>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="252"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <begin position="284"/>
            <end position="290"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="328"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="modified residue" description="Phosphotyrosine" evidence="19">
        <location>
            <position position="640"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007765"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00054255"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PIRNR" id="PIRNR002006"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="PIRSR" id="PIRSR002006-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="PIRSR" id="PIRSR002006-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="PIRSR" id="PIRSR002006-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015887.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="644" mass="74192" checksum="22D85A9C9594CAE7" modified="2019-05-08" version="1">PDCLAPYPHPHLLIAPPSLDSHTYPTLHWRNEHRCNSLCDIAEPQSMPLLARSPSTNRKYPPLPVDKLEEEINRRMADDNKLFREEFNALPACPIQATCEAASKEENKEKNRYVNILPYDHSRVHLTPIEGVPDSDYINASFINGYQEKNKFIAAQGPKEETVNDFWRMIWEQNTATIVMVTNLKERKECKCAQYWPDQGCWTYGNIRVSVEDVTVLVDYTVRKFCIQQVGDVTNKKPQRLVTQFHFTSWPDFGVPFTPIGMLKFLKKVKTCNPQYAGAIVVHCSAGVGRTGTFIVIDAMLDMMHAERKVDVYGFVSRIRAQRCQMVQTDMQYVFIYQALLEHYLYGDTELEVTSLETHLQKIYNKVPGTSSNGLEEEFKKLTSIKIQNDKMRTGNLPANMKKNRVLQIIPYEFNRVIIPVKRGEENTDYVNASFIDGYRQKDSYIASQGPLQHTIEDFWRMIWEWKSCSIVMLTELEERGQEKCAQYWPSDGSVSYGDITMELKKEEECESYTVRDLLVTNTRENKSRQIRQFHFHGWPEVGIPSDGKGMINIIAAVQKQQQQSGNHPITVHCSAGAGRTGTFCALSTVLERVKAEGILDVFQTVKSLRLQRPHMVQTLEQYEFCYKVVQEYIDAFSDYANFK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLM9</accession>
    <name>A0A452HLM9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">G-protein coupled receptors family 1 profile domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLM9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018088.1">
        <property type="protein sequence ID" value="ENSGAGP00000015834.1"/>
        <property type="gene ID" value="ENSGAGG00000011898.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045202">
        <property type="term" value="C:synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="GOC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001591">
        <property type="term" value="F:dopamine neurotransmitter receptor activity, coupled via Gi/Go"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004930">
        <property type="term" value="F:G protein-coupled receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007195">
        <property type="term" value="P:adenylate cyclase-inhibiting dopamine receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051481">
        <property type="term" value="P:negative regulation of cytosolic calcium ion concentration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051967">
        <property type="term" value="P:negative regulation of synaptic transmission, glutamatergic"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060158">
        <property type="term" value="P:phospholipase C-activating dopamine receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0014059">
        <property type="term" value="P:regulation of dopamine secretion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043266">
        <property type="term" value="P:regulation of potassium ion transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1070.10">
        <property type="entry name" value="Rhodopsin 7-helix transmembrane proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002185">
        <property type="entry name" value="Dopamine_D4_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000929">
        <property type="entry name" value="Dopamine_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000276">
        <property type="entry name" value="GPCR_Rhodpsn"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017452">
        <property type="entry name" value="GPCR_Rhodpsn_7TM"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24248">
        <property type="entry name" value="ADRENERGIC RECEPTOR-RELATED G-PROTEIN COUPLED RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24248:SF143">
        <property type="entry name" value="D(4) DOPAMINE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00001">
        <property type="entry name" value="7tm_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00569">
        <property type="entry name" value="DOPAMINED4R"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00242">
        <property type="entry name" value="DOPAMINER"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00237">
        <property type="entry name" value="GPCRRHODOPSN"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81321">
        <property type="entry name" value="Family A G protein-coupled receptor-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50262">
        <property type="entry name" value="G_PROTEIN_RECEP_F1_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="7" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5" id="KW-0297">G-protein coupled receptor</keyword>
    <keyword evidence="6 10" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-0675">Receptor</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0807">Transducer</keyword>
    <keyword evidence="3 10" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 10" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="32"/>
            <end position="51"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="71"/>
            <end position="98"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="191"/>
            <end position="212"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="224"/>
            <end position="247"/>
        </location>
    </feature>
    <feature type="domain" description="G-protein coupled receptors family 1 profile" evidence="11">
        <location>
            <begin position="13"/>
            <end position="244"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023224"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50262"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015834.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="263" mass="29988" checksum="0DE2BA0162F45EE6" modified="2019-05-08" version="1">MDVETAAQTPIMFIAVSIPLNYNRQQIDLRQLILISTTWIFAFAVASPVIFGLNNVPDRDPTLCQLEDNNYIVYSSICSFFIPCPVMLVLYCAMFQGLKRWEEARKAKLKSSVYAGGQKLCCPPPLLERKQAEIKLQECSLYDRSECSFPRGYMMNSGIQTVSCPHLKYSHPAHKQKQAKINGRERKAMKVLPVVVGAFLCCWTPFFVVHITRALCKSCTIPSLVTSIVTWLGYVNSALNPIIYTIFNTEFRKFFRKALHFFC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLN5</accession>
    <name>A0A452HLN5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">DnaJ homolog subfamily C member 21</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HLN5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018166.1">
        <property type="protein sequence ID" value="ENSGAGP00000015897.1"/>
        <property type="gene ID" value="ENSGAGG00000011966.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003676">
        <property type="term" value="F:nucleic acid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd06257">
        <property type="entry name" value="DnaJ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.110:FF:000046">
        <property type="entry name" value="dnaJ homolog subfamily C member 21"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.110">
        <property type="entry name" value="DnaJ domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051964">
        <property type="entry name" value="Chaperone_stress_response"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001623">
        <property type="entry name" value="DnaJ_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018253">
        <property type="entry name" value="DnaJ_domain_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036869">
        <property type="entry name" value="J_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003604">
        <property type="entry name" value="Matrin/U1-like-C_Znf_C2H2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022755">
        <property type="entry name" value="Znf_C2H2_jaz"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR054076">
        <property type="entry name" value="ZUO1-like_ZHD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR44029">
        <property type="entry name" value="DNAJ HOMOLOG SUBFAMILY C MEMBER 21"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR44029:SF1">
        <property type="entry name" value="DNAJ HOMOLOG SUBFAMILY C MEMBER 21"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00226">
        <property type="entry name" value="DnaJ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12171">
        <property type="entry name" value="zf-C2H2_jaz"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21884">
        <property type="entry name" value="ZUO1-like_ZHD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00625">
        <property type="entry name" value="JDOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00271">
        <property type="entry name" value="DnaJ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00451">
        <property type="entry name" value="ZnF_U1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46565">
        <property type="entry name" value="Chaperone J-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00636">
        <property type="entry name" value="DNAJ_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50076">
        <property type="entry name" value="DNAJ_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0862">Zinc</keyword>
    <keyword evidence="2" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="J" evidence="6">
        <location>
            <begin position="3"/>
            <end position="69"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="276"/>
            <end position="304"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="348"/>
            <end position="479"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="514"/>
            <end position="537"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="5">
        <location>
            <begin position="281"/>
            <end position="291"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="292"/>
            <end position="304"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="361"/>
            <end position="370"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="5">
        <location>
            <begin position="382"/>
            <end position="393"/>
        </location>
    </feature>
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        <location>
            <begin position="394"/>
            <end position="404"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="405"/>
            <end position="416"/>
        </location>
    </feature>
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        <location>
            <begin position="417"/>
            <end position="427"/>
        </location>
    </feature>
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        <location>
            <begin position="428"/>
            <end position="442"/>
        </location>
    </feature>
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        <location>
            <begin position="444"/>
            <end position="457"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="5">
        <location>
            <begin position="459"/>
            <end position="468"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="5">
        <location>
            <begin position="528"/>
            <end position="537"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00074367"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015897.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="537" mass="63199" checksum="DE92B4AACA8F0B64" modified="2019-05-08" version="1">MKCHYEVLGLARDASDEELKRAYRRLALRWHPDKNLENADEAAEQFKLIQAAYDVLSDPQERAWYDNHREALLKGGVDGEYQDDSLDLLRYFTVNCYSGYDDDEEGFFAVYRQVFENIAKEEMEYISQEDIEEFPTFGYSQSDYDKVVHPFYAYWQSFCTQKNFAWKEEYDTRQASNRWEKRAMEKENKKTRDKARKERNELVRQLVAFIRKRDKRVQAHRKLIEEQNAEKARKAEELRRQQKLKQAKLGEQYKEQSWIAMSDLERELQQMEAQYEKEFGDGSDDEEEREELEPREGIDDKLSDEAEGVEFYDDLYCPACDKSLKTEKAMKNHEKSKKHREMVSLLRQQLEEEEKEFSVSTDDKNEVNTREDEEMEETPKQKLSKKKKKKHKTMTTYEDSFNNKTTDDATDEQKEADQEDSITEELENVSQRNTASEKTSAAKTVDEVNEPKSEAKSIPKTKGKRTKDTKKSVKVTSELQDKNEDIPIHCITCNCEFQSRNKLFEHLKATGHAKAMQAPAANGAGNSRSKKEKRKNR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLN8</accession>
    <name>A0A452HLN8_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="12">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLN8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018108.1">
        <property type="protein sequence ID" value="ENSGAGP00000015848.1"/>
        <property type="gene ID" value="ENSGAGG00000011916.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003779">
        <property type="term" value="F:actin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000083">
        <property type="entry name" value="Uveal autoantigen with coiled-coil domains and ankyrin repeats"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042420">
        <property type="entry name" value="RAI14/UACA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24129">
        <property type="entry name" value="ANKYCORBIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24129:SF0">
        <property type="entry name" value="ANKYCORBIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00023">
        <property type="entry name" value="Ank"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01415">
        <property type="entry name" value="ANKYRIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="5"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5 9" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="6 10" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <feature type="repeat" description="ANK" evidence="9">
        <location>
            <begin position="52"/>
            <end position="84"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="9">
        <location>
            <begin position="85"/>
            <end position="117"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="9">
        <location>
            <begin position="118"/>
            <end position="150"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="9">
        <location>
            <begin position="151"/>
            <end position="183"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="9">
        <location>
            <begin position="184"/>
            <end position="216"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="244"/>
            <end position="310"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="362"/>
            <end position="399"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="10">
        <location>
            <begin position="456"/>
            <end position="748"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="10">
        <location>
            <begin position="777"/>
            <end position="915"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="9"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="245"/>
            <end position="259"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="282"/>
            <end position="291"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="300"/>
            <end position="310"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015848.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="950" mass="107767" checksum="BA45F5EAD1193B1D" modified="2019-05-08" version="1">MKSLKAKFRKSDTNEWNKNDDRLLQAVENGDPEKVASLLGKKGANATKQDSEGKTAFHLAAAKGHAECLRIMMTHGVDVTAQDGTGHTALHLAAKNSHLDCIKRLLQSKCPADSTDNSGKTALHYAASCGCLQVVQLLCEHKCPINVKDLDGNIPLLLAVQNGHAEICKHLLDHGADINSRDKNGRTALMLACEAGGLNIVETLIRKGADFSLVDALGHDALHYSKLSENAGIQNLLLSNISPDADMKSPTKPKQHDQVSKLSSERSGTPKKRKAPPPPISPIQEEISSIQRDNKDRLSDSTTGADSLLDVSSETDQQDLLVLLQAKIASLTLHNKELQDKLQEKAPRETEMDSTLNSYHSTQTEFDQSVDKQSQTSAQELRSSSLNATQTQEKSTNSEVEIKHLQEDLKDVQRKLDNSEAKRKYLETQLQSRIPETIHLNSTDISENSCDLSQKLKETQCQYEEAMKEVLNVQKQMKLGLVASENKDTYADVQELRVTCEEIEVLKQELKKALEESERHKEKVRELQEKLEEREQNVASRMSVEECEEIKSSYCSIIENINQEKALLIERYKEGQEEIKRLQDKLNQLQLESSEEPREMKEARNRMIDDLHRQVSELSRLYKEAQTELEDYRNRKALEDVALEYIPRDEHEKLMRVANSLKDKAEDALSDMKSQYTQVLNEAAQLKQLIDTQKQNSVPVAEHLEVVTALRCTSKEMEEEINELREELINKETELRSLQKELLEEKAAINEAMVPRASYEKLQSSLEREVSILSFKLKDLIKEKENVSLDAAQLRNEVLQLKGEKEGAQTLLEAKEQEINGLYRKYHQAQEDLLDMKRRAESSSKLEEDKDKKINEMSKEVSKLKEALNSLSQLSYSTSAPKRQSQQLEALQQQVKQLQNQLAETKKQHQEIVSVYRMHLLYAVQGQMDEDVQKVLKQILTMCKSQSQKK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLN9</accession>
    <name>A0A452HLN9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="12">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 8">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the TALE/PBX homeobox family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLN9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018106.1">
        <property type="protein sequence ID" value="ENSGAGP00000015846.1"/>
        <property type="gene ID" value="ENSGAGG00000011919.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00086">
        <property type="entry name" value="homeodomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.60:FF:000008">
        <property type="entry name" value="Pre-B-cell leukemia transcription factor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.60">
        <property type="entry name" value="Homeodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001356">
        <property type="entry name" value="HD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017970">
        <property type="entry name" value="Homeobox_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009057">
        <property type="entry name" value="Homeodomain-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008422">
        <property type="entry name" value="KN_HD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005542">
        <property type="entry name" value="PBX_PBC_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050224">
        <property type="entry name" value="TALE_homeobox"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11850">
        <property type="entry name" value="HOMEOBOX PROTEIN TRANSCRIPTION FACTORS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05920">
        <property type="entry name" value="Homeobox_KN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03792">
        <property type="entry name" value="PBC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00389">
        <property type="entry name" value="HOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46689">
        <property type="entry name" value="Homeodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00027">
        <property type="entry name" value="HOMEOBOX_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50071">
        <property type="entry name" value="HOMEOBOX_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51978">
        <property type="entry name" value="PBC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 8" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="5 8" id="KW-0371">Homeobox</keyword>
    <keyword evidence="7 8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0804">Transcription</keyword>
    <keyword evidence="3" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="PBC" evidence="11">
        <location>
            <begin position="1"/>
            <end position="126"/>
        </location>
    </feature>
    <feature type="domain" description="Homeobox" evidence="10">
        <location>
            <begin position="125"/>
            <end position="188"/>
        </location>
    </feature>
    <feature type="DNA-binding region" description="Homeobox" evidence="8">
        <location>
            <begin position="127"/>
            <end position="189"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="218"/>
            <end position="241"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="295"/>
            <end position="326"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="218"/>
            <end position="233"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="295"/>
            <end position="314"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007601"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023155"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00108"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50071"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS51978"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015846.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="326" mass="35508" checksum="67C681A7E95B1E57" modified="2019-05-08" version="1">MRLDNMLLAEGVSGPEKGGGSAAAAAAAAASGGSSDNSIEHSDYRAKLTQIRQIYHTELEKYEQACNEFTTHVMNLLREQSRTRPISPKEIERMVGIIHRKFSSIQMQLKQSTCEAVMILRSRFLDARRKRRNFSKQATEILNEYFYSHLSNPYPSEEAKEELAKKCSITVSQVSNWFGNKRIRYKKNIGKFQEEANLYAAKTAVTAAHAVAAAVQNNQTNSPTTPNSGSSGSFNLPNSGDMFMNMQSLNGDSYQGSQVGANVQSQVDTLRHVINQTGGYSDALGGGSLYSPHNLNANGGWQDATTPSSVTSPTEGPGSVHSDTSN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLP1</accession>
    <name>A0A452HLP1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Zinc finger E-box-binding homeobox 2</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="21">Smad-interacting protein 1</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="22">Zinc finger homeobox protein 1b</fullName>
        </alternativeName>
    </protein>
    <organism evidence="26 27">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="27" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="26" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="26" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="19">Transcriptional inhibitor that binds to DNA sequence 5'-CACCT-3' in different promoters. Represses transcription of E-cadherin. Represses expression of MEOX2.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Chromosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the delta-EF1/ZFH-1 C2H2-type zinc-finger family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLP1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018113.1">
        <property type="protein sequence ID" value="ENSGAGP00000015852.1"/>
        <property type="gene ID" value="ENSGAGG00000011928.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005694">
        <property type="term" value="C:chromosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005654">
        <property type="term" value="C:nucleoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000122">
        <property type="term" value="P:negative regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048023">
        <property type="term" value="P:positive regulation of melanin biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045944">
        <property type="term" value="P:positive regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030177">
        <property type="term" value="P:positive regulation of Wnt signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000013">
        <property type="entry name" value="Putative zinc finger E-box-binding homeobox 2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000082">
        <property type="entry name" value="Putative zinc finger E-box-binding homeobox 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.60:FF:000105">
        <property type="entry name" value="Zinc finger E-box binding homeobox 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000744">
        <property type="entry name" value="zinc finger E-box-binding homeobox 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000117">
        <property type="entry name" value="Zinc finger E-box-binding homeobox 2 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000145">
        <property type="entry name" value="Zinc finger protein 574"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.60">
        <property type="entry name" value="Homeodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008598">
        <property type="entry name" value="Di19_Zn-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001356">
        <property type="entry name" value="HD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009057">
        <property type="entry name" value="Homeodomain-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051574">
        <property type="entry name" value="ZnF_E-box_Homeobox"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24391">
        <property type="entry name" value="HISTONE H4 TRANSCRIPTION FACTOR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24391:SF11">
        <property type="entry name" value="ZINC FINGER E-BOX-BINDING HOMEOBOX 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05605">
        <property type="entry name" value="zf-Di19"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00389">
        <property type="entry name" value="HOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46689">
        <property type="entry name" value="Homeodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="6"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="13" id="KW-0007">Acetylation</keyword>
    <keyword evidence="4" id="KW-0158">Chromosome</keyword>
    <keyword evidence="15" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="16" id="KW-0371">Homeobox</keyword>
    <keyword evidence="6" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="8" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="18" id="KW-0539">Nucleus</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="27" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0678">Repressor</keyword>
    <keyword evidence="17" id="KW-0804">Transcription</keyword>
    <keyword evidence="14" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="12" id="KW-0832">Ubl conjugation</keyword>
    <keyword evidence="11" id="KW-0862">Zinc</keyword>
    <keyword evidence="10 23" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="25">
        <location>
            <begin position="211"/>
            <end position="239"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="25">
        <location>
            <begin position="241"/>
            <end position="263"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="25">
        <location>
            <begin position="282"/>
            <end position="309"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="25">
        <location>
            <begin position="999"/>
            <end position="1026"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="25">
        <location>
            <begin position="1027"/>
            <end position="1054"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="25">
        <location>
            <begin position="1055"/>
            <end position="1083"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="24">
        <location>
            <begin position="1"/>
            <end position="26"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="24">
        <location>
            <begin position="55"/>
            <end position="115"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="24">
        <location>
            <begin position="720"/>
            <end position="742"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="24">
        <location>
            <begin position="770"/>
            <end position="858"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="24">
        <location>
            <begin position="1117"/>
            <end position="1214"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="24">
        <location>
            <begin position="12"/>
            <end position="24"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="24">
        <location>
            <begin position="55"/>
            <end position="74"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="24">
        <location>
            <begin position="89"/>
            <end position="98"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="24">
        <location>
            <begin position="720"/>
            <end position="730"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="24">
        <location>
            <begin position="781"/>
            <end position="809"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="24">
        <location>
            <begin position="840"/>
            <end position="855"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="24">
        <location>
            <begin position="1127"/>
            <end position="1155"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="24">
        <location>
            <begin position="1156"/>
            <end position="1167"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="24">
        <location>
            <begin position="1168"/>
            <end position="1179"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="24">
        <location>
            <begin position="1186"/>
            <end position="1205"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00009867"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022454"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022491"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023155"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00058733"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00067197"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00080819"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00082894"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015852.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1214" mass="136328" checksum="20F7DA003E06D912" modified="2019-05-08" version="1">MKQQIMADGPRYKRRKQANPRRKNVVNYENVVETGSETDEEDKLHIAEDDSIINTLDQETSPASVPNHESSPHVSQALLPRDEEEDEMRESGVDHTWHNSEILQASVDGPEEMKEEYDTMGPEAAIQTTGNNGTVKNANCTSDFEEYFAKRKLEEGDGHAVSIAEYLQRSDTAIIYPEAPEELSRLGTPEANGQEENDLPPGTPDAFAQLLTCPYCDRGYKRLTSLKEHIKYRHEKNEENFSCPLCSYTFAYRTQLERHMVTHKPGTDQHQMLTQGAGNRKFKCTECGKAFKYKHHLKEHLRIHSGEKPYECPNCKKRFSHSGSYSSHISSKKCIGLISVNGRMRNNIKTGSSPNSVSSSPTNSAITQLRNKLENGKPLSMSEQTGLLKIKTEPLDFNEYKVLMATHGFSGASPFMNGGLGATSPLGVHASAQSPMQHLSVGMEAPLLGFPAIGSNLSEVQKVLQIVDNTVSRQKMDCKAEEISKLKGYHMKDPCPQTEEQGVTSPNIPPVGLPVVSHNGATKSIIDYTLEKVNEAKACLQSLTTDSRRQLSNIKKEKLRTLIDLVTEDKMIDNHNISTPFSCQFCKESFPGPIPLHQHERYLCKMNEEIKAVLQPHENMVPNKPGVFVDKQALLLSSVLSEKGMTSPINSYKDHMSVLKAYYAMNMEPNSDELLKISIAVGLPQEFVKEWFEQRKFYQYSSSRSPSLERTSAKAVLAATNTPTKDSLSARSPIKPVDSVTSPSIAELHNSITNCDTPLRLTKPTHFTNIKPVDKLDHSRSNTPSPLNLSSTSSKNSHSSSYTPNSFSSEELQAEPLDLSLPKQMKEPKSVIATKNKTKSSSINLDHNSVSSSSENSDEPLNLTFIKKEFSNSNNLDKSTNPVFGLNPFSAKPLYTTLPPQSAFPPATFMPPVQTSIPGLRPYPGLDQMSFLPHMAYTYPTGAATFADMQQRRKFQRKQGFQGELLDGTPDYMSGLDDMTDSDSCLSRKKIKKTESGMYACDLCDKTFQKSSSLLRHKYEHTGKRPHQCQICKKAFKHKHHLIEHSRLHSGEKPYQCDKCGKRFSHSGSYSQHMNHRYSYCKREAEEREAAEREAREKGHLEPTELLMNRAYLQSITPQGYSDSEERESMPRDGESEKEHEKEGEDGYEKLGRQDGDEEFEEEEEESENKSMDTDPDTIRDEEETGEHSMDDSSEDGKMETKSDHEEDNMEDGM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLP3</accession>
    <name>A0A452HLP3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Zinc finger E-box-binding homeobox 2</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="21">Smad-interacting protein 1</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="22">Zinc finger homeobox protein 1b</fullName>
        </alternativeName>
    </protein>
    <organism evidence="26 27">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="27" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="26" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="26" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="19">Transcriptional inhibitor that binds to DNA sequence 5'-CACCT-3' in different promoters. Represses transcription of E-cadherin. Represses expression of MEOX2.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Chromosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the delta-EF1/ZFH-1 C2H2-type zinc-finger family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLP3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018115.1">
        <property type="protein sequence ID" value="ENSGAGP00000015854.1"/>
        <property type="gene ID" value="ENSGAGG00000011928.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005694">
        <property type="term" value="C:chromosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005654">
        <property type="term" value="C:nucleoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000122">
        <property type="term" value="P:negative regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048023">
        <property type="term" value="P:positive regulation of melanin biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045944">
        <property type="term" value="P:positive regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030177">
        <property type="term" value="P:positive regulation of Wnt signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000013">
        <property type="entry name" value="Putative zinc finger E-box-binding homeobox 2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000082">
        <property type="entry name" value="Putative zinc finger E-box-binding homeobox 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.60:FF:000105">
        <property type="entry name" value="Zinc finger E-box binding homeobox 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000744">
        <property type="entry name" value="zinc finger E-box-binding homeobox 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000117">
        <property type="entry name" value="Zinc finger E-box-binding homeobox 2 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000145">
        <property type="entry name" value="Zinc finger protein 574"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.60">
        <property type="entry name" value="Homeodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008598">
        <property type="entry name" value="Di19_Zn-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001356">
        <property type="entry name" value="HD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009057">
        <property type="entry name" value="Homeodomain-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051574">
        <property type="entry name" value="ZnF_E-box_Homeobox"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24391">
        <property type="entry name" value="HISTONE H4 TRANSCRIPTION FACTOR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24391:SF11">
        <property type="entry name" value="ZINC FINGER E-BOX-BINDING HOMEOBOX 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05605">
        <property type="entry name" value="zf-Di19"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00389">
        <property type="entry name" value="HOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46689">
        <property type="entry name" value="Homeodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="6"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="13" id="KW-0007">Acetylation</keyword>
    <keyword evidence="4" id="KW-0158">Chromosome</keyword>
    <keyword evidence="15" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="16" id="KW-0371">Homeobox</keyword>
    <keyword evidence="6" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="8" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="18" id="KW-0539">Nucleus</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="27" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0678">Repressor</keyword>
    <keyword evidence="17" id="KW-0804">Transcription</keyword>
    <keyword evidence="14" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="12" id="KW-0832">Ubl conjugation</keyword>
    <keyword evidence="11" id="KW-0862">Zinc</keyword>
    <keyword evidence="10 23" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="25">
        <location>
            <begin position="148"/>
            <end position="176"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="25">
        <location>
            <begin position="178"/>
            <end position="200"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="25">
        <location>
            <begin position="219"/>
            <end position="246"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="25">
        <location>
            <begin position="936"/>
            <end position="963"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="25">
        <location>
            <begin position="964"/>
            <end position="991"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="25">
        <location>
            <begin position="992"/>
            <end position="1020"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="24">
        <location>
            <begin position="1"/>
            <end position="26"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="24">
        <location>
            <begin position="56"/>
            <end position="116"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="24">
        <location>
            <begin position="657"/>
            <end position="679"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="24">
        <location>
            <begin position="707"/>
            <end position="795"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="24">
        <location>
            <begin position="1054"/>
            <end position="1151"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="24">
        <location>
            <begin position="12"/>
            <end position="24"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="24">
        <location>
            <begin position="56"/>
            <end position="74"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="24">
        <location>
            <begin position="89"/>
            <end position="98"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="24">
        <location>
            <begin position="657"/>
            <end position="667"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="24">
        <location>
            <begin position="718"/>
            <end position="746"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="24">
        <location>
            <begin position="777"/>
            <end position="792"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="24">
        <location>
            <begin position="1064"/>
            <end position="1092"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="24">
        <location>
            <begin position="1093"/>
            <end position="1104"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="24">
        <location>
            <begin position="1105"/>
            <end position="1116"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="24">
        <location>
            <begin position="1123"/>
            <end position="1142"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00009867"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022454"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022491"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023155"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00058733"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00067197"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00080819"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00082894"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015854.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1151" mass="129313" checksum="04B5707250A14408" modified="2019-05-08" version="1">MKQQIMADGPRYKRRKQANPRRKNVVNYENVVETGSETDEEDKLHIAEDDSIINTLDQETSPASVPNHESSPHVSQALLPRDEEEDEMRESGVDHTWHNSEILQASVDGPEEMKEEYDTMGPEAAIQTTGNNGTDLPPGTPDAFAQLLTCPYCDRGYKRLTSLKEHIKYRHEKNEENFSCPLCSYTFAYRTQLERHMVTHKPGTDQHQMLTQGAGNRKFKCTECGKAFKYKHHLKEHLRIHSGEKPYECPNCKKRFSHSGSYSSHISSKKCIGLISVNGRMRNNIKTGSSPNSVSSSPTNSAITQLRNKLENGKPLSMSEQTGLLKIKTEPLDFNEYKVLMATHGFSGASPFMNGGLGATSPLGVHASAQSPMQHLSVGMEAPLLGFPAIGSNLSEVQKVLQIVDNTVSRQKMDCKAEEISKLKGYHMKDPCPQTEEQGVTSPNIPPVGLPVVSHNGATKSIIDYTLEKVNEAKACLQSLTTDSRRQLSNIKKEKLRTLIDLVTEDKMIDNHNISTPFSCQFCKESFPGPIPLHQHERYLCKMNEEIKAVLQPHENMVPNKPGVFVDKQALLLSSVLSEKGMTSPINSYKDHMSVLKAYYAMNMEPNSDELLKISIAVGLPQEFVKEWFEQRKFYQYSSSRSPSLERTSAKAVLAATNTPTKDSLSARSPIKPVDSVTSPSIAELHNSITNCDTPLRLTKPTHFTNIKPVDKLDHSRSNTPSPLNLSSTSSKNSHSSSYTPNSFSSEELQAEPLDLSLPKQMKEPKSVIATKNKTKSSSINLDHNSVSSSSENSDEPLNLTFIKKEFSNSNNLDKSTNPVFGLNPFSAKPLYTTLPPQSAFPPATFMPPVQTSIPGLRPYPGLDQMSFLPHMAYTYPTGAATFADMQQRRKFQRKQGFQGELLDGTPDYMSGLDDMTDSDSCLSRKKIKKTESGMYACDLCDKTFQKSSSLLRHKYEHTGKRPHQCQICKKAFKHKHHLIEHSRLHSGEKPYQCDKCGKRFSHSGSYSQHMNHRYSYCKREAEEREAAEREAREKGHLEPTELLMNRAYLQSITPQGYSDSEERESMPRDGESEKEHEKEGEDGYEKLGRQDGDEEFEEEEEESENKSMDTDPDTIRDEEETGEHSMDDSSEDGKMETKSDHEEDNMEDGM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLP4</accession>
    <name>A0A452HLP4_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="12">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLP4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018114.1">
        <property type="protein sequence ID" value="ENSGAGP00000015853.1"/>
        <property type="gene ID" value="ENSGAGG00000011916.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003779">
        <property type="term" value="F:actin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000083">
        <property type="entry name" value="Uveal autoantigen with coiled-coil domains and ankyrin repeats"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042420">
        <property type="entry name" value="RAI14/UACA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24129">
        <property type="entry name" value="ANKYCORBIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24129:SF0">
        <property type="entry name" value="ANKYCORBIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00023">
        <property type="entry name" value="Ank"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01415">
        <property type="entry name" value="ANKYRIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="5"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5 9" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="6 10" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <feature type="repeat" description="ANK" evidence="9">
        <location>
            <begin position="52"/>
            <end position="84"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="9">
        <location>
            <begin position="85"/>
            <end position="117"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="9">
        <location>
            <begin position="118"/>
            <end position="150"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="9">
        <location>
            <begin position="151"/>
            <end position="183"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="9">
        <location>
            <begin position="184"/>
            <end position="216"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="247"/>
            <end position="270"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="290"/>
            <end position="310"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="362"/>
            <end position="399"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="10">
        <location>
            <begin position="456"/>
            <end position="748"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="10">
        <location>
            <begin position="777"/>
            <end position="915"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="9"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="252"/>
            <end position="269"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="290"/>
            <end position="299"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="300"/>
            <end position="310"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015853.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="950" mass="107632" checksum="88CF22898CB46014" modified="2019-05-08" version="1">MKSLKAKFRKSDTNEWNKNDDRLLQAVENGDPEKVASLLGKKGANATKQDSEGKTAFHLAAAKGHAECLRIMMTHGVDVTAQDGTGHTALHLAAKNSHLDCIKRLLQSKCPADSTDNSGKTALHYAASCGCLQVVQLLCEHKCPINVKDLDGNIPLLLAVQNGHAEICKHLLDHGADINSRDKNGRTALMLACEAGGLNIVETLIRKGADFSLVDALGHDALHYSKLSENAGIQNLLLSNISPDADMKSPTKPKQFSDLSSPRSATSTPIAGKGQAFFAEQVCKEEISSIQRDNKDRLSDSTTGADSLLDVSSETDQQDLLVLLQAKIASLTLHNKELQDKLQEKAPRETEMDSTLNSYHSTQTEFDQSVDKQSQTSAQELRSSSLNATQTQEKSTNSEVEIKHLQEDLKDVQRKLDNSEAKRKYLETQLQSRIPETIHLNSTDISENSCDLSQKLKETQCQYEEAMKEVLNVQKQMKLGLVASENKDTYADVQELRVTCEEIEVLKQELKKALEESERHKEKVRELQEKLEEREQNVASRMSVEECEEIKSSYCSIIENINQEKALLIERYKEGQEEIKRLQDKLNQLQLESSEEPREMKEARNRMIDDLHRQVSELSRLYKEAQTELEDYRNRKALEDVALEYIPRDEHEKLMRVANSLKDKAEDALSDMKSQYTQVLNEAAQLKQLIDTQKQNSVPVAEHLEVVTALRCTSKEMEEEINELREELINKETELRSLQKELLEEKAAINEAMVPRASYEKLQSSLEREVSILSFKLKDLIKEKENVSLDAAQLRNEVLQLKGEKEGAQTLLEAKEQEINGLYRKYHQAQEDLLDMKRRAESSSKLEEDKDKKINEMSKEVSKLKEALNSLSQLSYSTSAPKRQSQQLEALQQQVKQLQNQLAETKKQHQEIVSVYRMHLLYAVQGQMDEDVQKVLKQILTMCKSQSQKK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLP5</accession>
    <name>A0A452HLP5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="30">Serine/threonine-protein kinase MARK2</fullName>
            <ecNumber evidence="8">2.7.11.1</ecNumber>
            <ecNumber evidence="7">2.7.11.26</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="31">ELKL motif kinase 1</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="32">MAP/microtubule affinity-regulating kinase 2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="38 39">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="39" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="38" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="38" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="28">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="27">
            <text>L-seryl-[tau protein] + ATP = O-phospho-L-seryl-[tau protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:12801"/>
            <dbReference type="Rhea" id="RHEA-COMP:13701"/>
            <dbReference type="Rhea" id="RHEA-COMP:13702"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.26"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="26">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="29">
            <text>L-threonyl-[tau protein] + ATP = O-phospho-L-threonyl-[tau protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:53904"/>
            <dbReference type="Rhea" id="RHEA-COMP:13703"/>
            <dbReference type="Rhea" id="RHEA-COMP:13704"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.26"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="5">Cell projection</location>
            <location evidence="5">Dendrite</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cytoplasm</location>
            <location evidence="4">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Endomembrane system</location>
            <topology evidence="3">Peripheral membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Lateral cell membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the protein kinase superfamily. CAMK Ser/Thr protein kinase family. SNF1 subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="8"/>
    <dbReference type="EC" id="2.7.11.26" evidence="7"/>
    <dbReference type="AlphaFoldDB" id="A0A452HLP5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018178.1">
        <property type="protein sequence ID" value="ENSGAGP00000015907.1"/>
        <property type="gene ID" value="ENSGAGG00000011970.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030425">
        <property type="term" value="C:dendrite"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012505">
        <property type="term" value="C:endomembrane system"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016328">
        <property type="term" value="C:lateral plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008289">
        <property type="term" value="F:lipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0106310">
        <property type="term" value="F:protein serine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="RHEA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050321">
        <property type="term" value="F:tau-protein kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035556">
        <property type="term" value="P:intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000226">
        <property type="term" value="P:microtubule cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016055">
        <property type="term" value="P:Wnt signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd14072">
        <property type="entry name" value="STKc_MARK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd14406">
        <property type="entry name" value="UBA_MARK2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:001032">
        <property type="entry name" value="KP78b, isoform A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.8.10:FF:000013">
        <property type="entry name" value="Non-specific serine/threonine protein kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000003">
        <property type="entry name" value="Non-specific serine/threonine protein kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.310.80:FF:000001">
        <property type="entry name" value="Non-specific serine/threonine protein kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.8.10">
        <property type="entry name" value="DNA helicase RuvA subunit, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.310.80">
        <property type="entry name" value="Kinase associated domain 1, KA1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028375">
        <property type="entry name" value="KA1/Ssp2_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001772">
        <property type="entry name" value="KA1_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049508">
        <property type="entry name" value="MARK1-4_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015940">
        <property type="entry name" value="UBA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24346">
        <property type="entry name" value="MAP/MICROTUBULE AFFINITY-REGULATING KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24346:SF56">
        <property type="entry name" value="SERINE_THREONINE-PROTEIN KINASE MARK2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02149">
        <property type="entry name" value="KA1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00627">
        <property type="entry name" value="UBA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00165">
        <property type="entry name" value="UBA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103243">
        <property type="entry name" value="KA1-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50032">
        <property type="entry name" value="KA1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50030">
        <property type="entry name" value="UBA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="20 33" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="10" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="25" id="KW-0966">Cell projection</keyword>
    <keyword evidence="11" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="24" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="9" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="19" id="KW-0221">Differentiation</keyword>
    <keyword evidence="18" id="KW-0418">Kinase</keyword>
    <keyword evidence="22" id="KW-0446">Lipid-binding</keyword>
    <keyword evidence="21" id="KW-0460">Magnesium</keyword>
    <keyword evidence="23" id="KW-0472">Membrane</keyword>
    <keyword evidence="16" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="17 33" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="13" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="39" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="12" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="14" id="KW-0808">Transferase</keyword>
    <keyword evidence="15" id="KW-0879">Wnt signaling pathway</keyword>
    <feature type="domain" description="Protein kinase" evidence="35">
        <location>
            <begin position="54"/>
            <end position="305"/>
        </location>
    </feature>
    <feature type="domain" description="UBA" evidence="36">
        <location>
            <begin position="324"/>
            <end position="363"/>
        </location>
    </feature>
    <feature type="domain" description="KA1" evidence="37">
        <location>
            <begin position="723"/>
            <end position="772"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="34">
        <location>
            <begin position="1"/>
            <end position="50"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="34">
        <location>
            <begin position="382"/>
            <end position="589"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="34">
        <location>
            <begin position="607"/>
            <end position="672"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="34">
        <location>
            <begin position="19"/>
            <end position="46"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="34">
        <location>
            <begin position="417"/>
            <end position="431"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="34">
        <location>
            <begin position="432"/>
            <end position="444"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="34">
        <location>
            <begin position="466"/>
            <end position="485"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="34">
        <location>
            <begin position="493"/>
            <end position="505"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="34">
        <location>
            <begin position="506"/>
            <end position="525"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="34">
        <location>
            <begin position="546"/>
            <end position="560"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="34">
        <location>
            <begin position="642"/>
            <end position="672"/>
        </location>
    </feature>
    <feature type="binding site" evidence="33">
        <location>
            <position position="83"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004124"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004184"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004279"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00006234"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00012407"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00012513"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022687"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00023121"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00047899"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="ARBA" id="ARBA00048291"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="ARBA" id="ARBA00048679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="ARBA" id="ARBA00048878"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="ARBA" id="ARBA00070360"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="31">
        <source>
            <dbReference type="ARBA" id="ARBA00079159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="32">
        <source>
            <dbReference type="ARBA" id="ARBA00083985"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="33">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="34">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="35">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="36">
        <source>
            <dbReference type="PROSITE" id="PS50030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="37">
        <source>
            <dbReference type="PROSITE" id="PS50032"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="38">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015907.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="39">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="772" mass="86398" checksum="C9B37E5A58AA81C8" modified="2019-05-08" version="1">MSSSGRSPLPTVNERDTEQPTLGHTESKASSKSNMLRGRNSATSTDEQPHIGNYRLLKTIGKGNFAKVKLARHVLTGKEVAVKIIDKTQLNSSSLQKLFREVRIMKVLNHPNIVKLFEVIETEKTLYLVMEYASGGEVFDYLVAHGRMKEKEARAKFRQIVSAVQYCHQKFIVHRDLKAENLLLDADMNIKIADFGFSNEFTFGNKLDTFCGSPPYAAPELFQGKKYDGPEVDVWSLGVILYTLVSGSLPFDGQNLKELRERVLRGKYRIPFYMSTDCENLLKKFLILNPSKRGTLEQIMKDRWMNVGHEDDELKPYMEPVPDYKDPRRTELMISMGYTREEIQESLIGQKYNEVMATYLLLGYKNSELDNDNITLKPRPAAELANSSAPSPSHKVQRSVSANPKQRRFSDQVPAIPTSTSYSKKTQSNNAENKRPEEEREAGRKASSTVKVPASPLTGLERKKSTPTPSTNSVLSTSTNRSRNSPMLERTGLGQNSIQNGKDSLTTPGSRASTASASAAVGSARPRQHQKSMSASVHPNKPTLPPTENNCEAQRPSTAPQRVPGASPSAHNISSAIPERTNFPRGISSRSTFHAGQLRQVRDQQNLPYNVPPASPSGNSQGRRGASGSIFSKFTSKFVRRNPHEPESKDRVETLRPHMVGGDKERDENRDAKPRSLRFTWSMKTTSSMEPNEMMKEIRKVLDANSCQCELQEKYMLLCMHGAPGHDSFVQWEMEVCKLPRLSLNGVRFKRITGTSMAFKNIASKVANELKL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLP9</accession>
    <name>A0A452HLP9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="12">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLP9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018120.1">
        <property type="protein sequence ID" value="ENSGAGP00000015859.1"/>
        <property type="gene ID" value="ENSGAGG00000011916.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003779">
        <property type="term" value="F:actin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000083">
        <property type="entry name" value="Uveal autoantigen with coiled-coil domains and ankyrin repeats"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042420">
        <property type="entry name" value="RAI14/UACA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24129">
        <property type="entry name" value="ANKYCORBIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24129:SF0">
        <property type="entry name" value="ANKYCORBIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00023">
        <property type="entry name" value="Ank"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01415">
        <property type="entry name" value="ANKYRIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="5"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5 9" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="6 10" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <feature type="repeat" description="ANK" evidence="9">
        <location>
            <begin position="52"/>
            <end position="84"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="9">
        <location>
            <begin position="85"/>
            <end position="117"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="9">
        <location>
            <begin position="118"/>
            <end position="150"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="9">
        <location>
            <begin position="151"/>
            <end position="183"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="9">
        <location>
            <begin position="184"/>
            <end position="216"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="242"/>
            <end position="281"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="333"/>
            <end position="370"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="10">
        <location>
            <begin position="427"/>
            <end position="719"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="10">
        <location>
            <begin position="748"/>
            <end position="886"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="9"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="245"/>
            <end position="270"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="271"/>
            <end position="281"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015859.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="921" mass="104619" checksum="9433076AC0A4F26B" modified="2019-05-08" version="1">MKSLKAKFRKSDTNEWNKNDDRLLQAVENGDPEKVASLLGKKGANATKQDSEGKTAFHLAAAKGHAECLRIMMTHGVDVTAQDGTGHTALHLAAKNSHLDCIKRLLQSKCPADSTDNSGKTALHYAASCGCLQVVQLLCEHKCPINVKDLDGNIPLLLAVQNGHAEICKHLLDHGADINSRDKNGRTALMLACEAGGLNIVETLIRKGADFSLVDALGHDALHYSKLSENAGIQNLLLSNISPDADMKSPTKPKQEEISSIQRDNKDRLSDSTTGADSLLDVSSETDQQDLLVLLQAKIASLTLHNKELQDKLQEKAPRETEMDSTLNSYHSTQTEFDQSVDKQSQTSAQELRSSSLNATQTQEKSTNSEVEIKHLQEDLKDVQRKLDNSEAKRKYLETQLQSRIPETIHLNSTDISENSCDLSQKLKETQCQYEEAMKEVLNVQKQMKLGLVASENKDTYADVQELRVTCEEIEVLKQELKKALEESERHKEKVRELQEKLEEREQNVASRMSVEECEEIKSSYCSIIENINQEKALLIERYKEGQEEIKRLQDKLNQLQLESSEEPREMKEARNRMIDDLHRQVSELSRLYKEAQTELEDYRNRKALEDVALEYIPRDEHEKLMRVANSLKDKAEDALSDMKSQYTQVLNEAAQLKQLIDTQKQNSVPVAEHLEVVTALRCTSKEMEEEINELREELINKETELRSLQKELLEEKAAINEAMVPRASYEKLQSSLEREVSILSFKLKDLIKEKENVSLDAAQLRNEVLQLKGEKEGAQTLLEAKEQEINGLYRKYHQAQEDLLDMKRRAESSSKLEEDKDKKINEMSKEVSKLKEALNSLSQLSYSTSAPKRQSQQLEALQQQVKQLQNQLAETKKQHQEIVSVYRMHLLYAVQGQMDEDVQKVLKQILTMCKSQSQKK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM00</accession>
    <name>A0A452HM00_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Dysferlin</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="17">Dystrophy-associated fer-1-like protein</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="18">Fer-1-like protein 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">Key calcium ion sensor involved in the Ca(2+)-triggered synaptic vesicle-plasma membrane fusion. Plays a role in the sarcolemma repair mechanism of both skeletal muscle and cardiomyocytes that permits rapid resealing of membranes disrupted by mechanical stress.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <location evidence="1">Sarcolemma</location>
            <topology evidence="1">Single-pass type II membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasmic vesicle membrane</location>
            <topology evidence="2">Single-pass type II membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the ferlin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018246.1">
        <property type="protein sequence ID" value="ENSGAGP00000015969.1"/>
        <property type="gene ID" value="ENSGAGG00000011993.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030659">
        <property type="term" value="C:cytoplasmic vesicle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012505">
        <property type="term" value="C:endomembrane system"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015630">
        <property type="term" value="C:microtubule cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030315">
        <property type="term" value="C:T-tubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008289">
        <property type="term" value="F:lipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002281">
        <property type="term" value="P:macrophage activation involved in immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061025">
        <property type="term" value="P:membrane fusion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002280">
        <property type="term" value="P:monocyte activation involved in immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050765">
        <property type="term" value="P:negative regulation of phagocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007009">
        <property type="term" value="P:plasma membrane organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd08373">
        <property type="entry name" value="C2A_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04011">
        <property type="entry name" value="C2B_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04018">
        <property type="entry name" value="C2C_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04017">
        <property type="entry name" value="C2D_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04037">
        <property type="entry name" value="C2E_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd08374">
        <property type="entry name" value="C2F_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000009">
        <property type="entry name" value="dysferlin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000021">
        <property type="entry name" value="dysferlin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000026">
        <property type="entry name" value="dysferlin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000033">
        <property type="entry name" value="dysferlin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000037">
        <property type="entry name" value="dysferlin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000061">
        <property type="entry name" value="dysferlin isoform X8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000008">
        <property type="entry name" value="C2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037726">
        <property type="entry name" value="C2A_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037720">
        <property type="entry name" value="C2B_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037722">
        <property type="entry name" value="C2C_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037723">
        <property type="entry name" value="C2D_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037724">
        <property type="entry name" value="C2E_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037725">
        <property type="entry name" value="C2F_Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012968">
        <property type="entry name" value="FerIin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037721">
        <property type="entry name" value="Ferlin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012560">
        <property type="entry name" value="Ferlin_A-domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012561">
        <property type="entry name" value="Ferlin_B-domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032362">
        <property type="entry name" value="Ferlin_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055072">
        <property type="entry name" value="Ferlin_DSRM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006614">
        <property type="entry name" value="Peroxin/Ferlin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12546:SF44">
        <property type="entry name" value="DYSFERLIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12546">
        <property type="entry name" value="FER-1-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00168">
        <property type="entry name" value="C2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22901">
        <property type="entry name" value="dsrm_Ferlin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08165">
        <property type="entry name" value="FerA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08150">
        <property type="entry name" value="FerB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08151">
        <property type="entry name" value="FerI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16165">
        <property type="entry name" value="Ferlin_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00239">
        <property type="entry name" value="C2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SMART" id="SM00694">
        <property type="entry name" value="DysFC"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00693">
        <property type="entry name" value="DysFN"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM01200">
        <property type="entry name" value="FerA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01201">
        <property type="entry name" value="FerB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01202">
        <property type="entry name" value="FerI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49562">
        <property type="entry name" value="C2 domain (Calcium/lipid-binding domain, CaLB)"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50004">
        <property type="entry name" value="C2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0106">Calcium</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="14" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="12" id="KW-0446">Lipid-binding</keyword>
    <keyword evidence="13 20" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="10" id="KW-0735">Signal-anchor</keyword>
    <keyword evidence="6 20" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 20" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="2048"/>
            <end position="2070"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="1"/>
            <end position="101"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="197"/>
            <end position="315"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="354"/>
            <end position="489"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="1116"/>
            <end position="1242"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="1289"/>
            <end position="1417"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="1564"/>
            <end position="1682"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="21">
        <location>
            <begin position="1798"/>
            <end position="1946"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="127"/>
            <end position="208"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1999"/>
            <end position="2020"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="19">
        <location>
            <begin position="127"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="172"/>
            <end position="182"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="19">
        <location>
            <begin position="188"/>
            <end position="203"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="2001"/>
            <end position="2020"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004274"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007561"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023121"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00060070"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00068801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00078385"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00079079"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015969.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2083" mass="238023" checksum="C4968BD16E3BA5D0" modified="2019-05-08" version="1">MLRVFILHAENVQTPDTDISDSYCSVVFEGVKKRTKVIKNNVNPVWNEGFDWDLKGIPLDQNAELTVVVKDHETVGRNRFLGEARVPLQDVLVNPTLTASFNAPLLDTKRQSTGAFLNLQVSYIPPPGAAPLFPPPTPPEPTPSTAELDTVTDTAGEEETEDQIVTGDETEPSTGPSGSEQPTLPRKPPSHHVHGIKRKKSAPKKLLSNKPQDFQIRVRVIEGRQLPGVNIKSVVKVTVAGQTKRTRIRKGNGPFFDEIFFFNVFESPAELFDEPIFITVVDSRSLRTDSVIGEFQLDVGTVYAEPKHALLRKWLLLSDSEDFSAGAKGYVKVSLFVLGPGDEAPLEKKEVAEEKEDIEGNLLRPTGVTLRGAHFCLKIYRAEDLPQMDDAIMDNVKQIFGFESNKKNLVDPFVEVSFAGRTISSKILEKNANPQWNQRITVPAMFPSMCEKMRIRVMDWDRVTHNDVIGTTHLRLSQISAPGGELEVDDGLGFLPTFGPCYVNLYGSPREFTGFPDPYEELNMGKGEGVAFRGRLLMELETKLVEHIEQKLEDIPPDDILRVEKHLRRRKYCLFAAFYSASMLQDVDAAIQFEVSIGNYGNKFDNTCLPLASTTQYSGAVFDGCQYYYLPWGNVKPVVVLSSYWEDISHRTDMQNLLQRTADRLEAGLELVHLALKARCSSNELDSLVSQLIDDIIADCSQPLVDVSNKTASTHLDQFVYKFRSTNLHQIIQAALNLKHEDSDLQTVLDQAEDWLLRLKAMAEEPQNSLPDIVIWMLQGDKRVAYARVPAHEVLFSRTGANCCGKNCGKLQTIFLKYPQEEVMGPRIPAQVRVQLWFGLAVDEKEFNKYAEGKLSVFAETYENQVKLALVGNWGTTGLTYPKYSDVTGKIKLPKDSFRPSVGWTWAGDWFISPEKTLLYDTDAGHMSFVEEVFENQVRLPGGQWIHMADAYTDVNGEKVLPKEEIECPPGWKWEDVEWDTDLNRAVDERGWEYGIAIPPDHKPKAWVPAEKMYHTSRRRRWVRLRRRDLKQMEALKKHKQEELEGEGWEYASLFGWKFHLNYRKTDTFRRRRWRRRMEPLERTGAAAVFALEGALGGVTDDKSDDGKSVSTLSFGVNRPTVSCIFDYGSRYHLRCYIYQARDLIAMDKDSFSDPYAIVSFLHQSQKTVVVKNTLNPLWDQTLIFYEIEIFGDPQNVADVPPNIVVEIYDHDTYGADEFMGRCICKPSLERSPRLSWHPVARGNRKAGELLAAFELIQREKPAVHHLPGFESELSASLDESADSDLPYPPPQREPNIYMVPQGIKPVLQRTAIEILAWGLRNLKSYQLASITSPSLQVECGGQTVQSCVIKNVRKNPNFDVCVLFMEVRLPKEALYCPPVIMKVIDNRQFGRKPVVGQCTIQSLEEFYCDPYKEETDAPEEQPDEVSLTPRDDVLIDIDDKEPLIPVQLAEGMTSSTLINMPASRADLHKIPWEEEFIDWWSKFYASTGEREKCGSYLEKGFDTLKVYETELENMEEFERLSDFCHTFKLYRGRTQDINEDPSVVGEFKGSFKMYPLPDDPRVPVPPRQFHQLPAKGPQECLIRIYIIRAFDLQPKDSNGKCDPYVKISVGKKSINDQENYIPCTLEPVFGKMFELTCTLPLEKDLKVTLYDYDLLSKDEKIGETVIDLENRFLSKFGARCGLPQTYCISGPNQWRDQLRPSQLLHLFSLQRNCKAPIYKPDRVVFREQEYFLSEFEDGKPPNPHLGAPEERLALYTLRKQGLVPEHVETRPLYSPLQPEIEQGKLQMWVDLFPKSLGLPGPPFNITPRRAKRFFLRCIIWNTKDVILDDLSITGEKMSDIYVKGWMTGHEENKQKTDVHYRSMGGEGNFNWRFVFPFDYLAAEQVCHIAKKEHFWSLDKTDNKVPPQLILQIWDNDKFSFDDYLGSVQMNLNRMPAPTKTAEKCSLNLLDETLSSDRFVSLFEQKAVKGWWPCVVEAEEKKTIAGKLEMTLEIVTEQEQEERPAGMGRDEPNMNPKLEDPKRPETSFLWFTSPYKTMKYILWRRFKWVFLIAIVVFILLLFIGIFIYAFPNYAAMKLVKPFS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="22" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM01</accession>
    <name>A0A452HM01_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HM01"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018242.1">
        <property type="protein sequence ID" value="ENSGAGP00000015966.1"/>
        <property type="gene ID" value="ENSGAGG00000012007.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="1" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="1">
        <location>
            <begin position="64"/>
            <end position="85"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="1">
        <location>
            <begin position="122"/>
            <end position="141"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015966.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="152" mass="17053" checksum="1A048E53502ACD45" modified="2019-05-08" version="1">MVRVTSPLDFIPNSSSLLLSRVKISPLYSQFYCYDPVVGSVKAAESPVWSCGELSLQKGEHRGAVLSLGFFCFAIVFCALIQAAFWRYTNNSEFLCCGKQSPFGETGGIENETCPSEHMGTVYGMILTSFLWFSIHFSNNLDRRGKYIFRET</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM05</accession>
    <name>A0A452HM05_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Fibronectin type-III domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the type I cytokine receptor family. Type 2 subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HM05"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018258.1">
        <property type="protein sequence ID" value="ENSGAGP00000015978.1"/>
        <property type="gene ID" value="ENSGAGG00000011475.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009897">
        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004896">
        <property type="term" value="F:cytokine receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000494">
        <property type="entry name" value="Leptin receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000501">
        <property type="entry name" value="Leptin receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000568">
        <property type="entry name" value="Leptin receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000613">
        <property type="entry name" value="Leptin receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003529">
        <property type="entry name" value="Hematopoietin_rcpt_Gp130_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010457">
        <property type="entry name" value="IgC2-like_lig-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041182">
        <property type="entry name" value="LEP-R_IGD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23037">
        <property type="entry name" value="CYTOKINE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23037:SF44">
        <property type="entry name" value="LEPTIN RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06328">
        <property type="entry name" value="Lep_receptor_Ig"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18589">
        <property type="entry name" value="ObR_Ig"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01353">
        <property type="entry name" value="HEMATOPO_REC_L_F2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7 12" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-0675">Receptor</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="3 12" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 12" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="807"/>
            <end position="829"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="13">
        <location>
            <begin position="209"/>
            <end position="300"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="13">
        <location>
            <begin position="507"/>
            <end position="603"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="13">
        <location>
            <begin position="708"/>
            <end position="801"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="924"/>
            <end position="943"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="924"/>
            <end position="933"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="934"/>
            <end position="943"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008921"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015978.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1131" mass="128446" checksum="331ECCC3F53F8ED2" modified="2019-05-08" version="1">ESCPSFCFHIMDYIPASSFMLPCMSPNETSVIPLTAGVVASSPGLHREHGTTETNSLMLVAEESFLCCLWSETNANCSLYTAGMETKKFTSSEVSSFAPQQTGSWNIQCWTKGDLELFVCILKLSNKKSYLNGEFKINLLYVLLLSGSLKGNFMVTPCNGNGSDKYECHIPSLKLNYTYITWLNIINGVTLLQSPLMSVRPINIVKPESPLNLRLEMTDKGQLKICWSNPVLTPYPLQYEVKFSGNTTQNVVEIVIETSLIIGNMLVGSSYLVQVRCKSLHGTGFWSDWSTPYNLNTEDVMYFPPKILTSVGSNVSFHCLYNDKNKMILSKKIVWWLNLAEEIPERQYTLVNDRVSRVTLFNLNATKPRGKFFYNALYCCNQNRECHHRYAELYVVDVNINITCETDGYLTKMTCRWSANTNTLLVGSSLQLKYYRSSIYCSDFPSIPPKSEAKECHLQRNHYYECTFQPIFLLSGYTMWIEIKHQLGTLESPPTCVIPADVVKPFPPSNVKAEITKNIGLLNVSWTNPAFPNSDLKFQIRYSVNREEILWEIFEVSNAPTRSVMIKVLDLCVVYIVQVRCSGLDGLGYWSDWSKPAYTIVQDIQAPLRGPEFWRVINEDPIRKQKNVTLVWKPLMKNYSLCSVCGYIVKQHTSENITWTEYVTSGTTYTYPWMEDAHTITILAVNSVGASSMNFNLTLSQQMSTVNIVQSLSAYPVNSSCVILTWTLSPQMYVITSFVIEWKNLNEEEQMKWIRVAPNISKYYIYDHFILIEKYQFSLYPIFPEGVGNPKTTDEFIKDRSEKQNDAGLYVILPIVISSFVLLLGTLLISQQRMKKLFWEDVPNPKNCSWAQGVNFQKPETFEHLFIKHPEAISFGPLLLEPEIVLEDISVAKALKSEDKQDLLAVDSMFTMIQDSEHDSACSSSHFSNSLSESSHDDKVSERTMRQSNIKYATIISNCKLSGLYEEQKNLSGSFSGCFLGEDSLVIDPFSRSWEVGNQAFLILPDQHPSQASKTISLSVVSSEGFSEPSDHDDTFSDGDSPERSLYYLGLTSVKKNENDIFLTENSRVMCHLHTNGLFKDMGFLQDTSSDLNPFIKNSLKYENSVKTFVPYMPQFQATTIKLQETTETKT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM08</accession>
    <name>A0A452HM08_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">ABC transporter domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the ABC transporter superfamily. ABCF family. EF3 subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HM08"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018299.1">
        <property type="protein sequence ID" value="ENSGAGP00000016017.1"/>
        <property type="gene ID" value="ENSGAGG00000012040.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd03221">
        <property type="entry name" value="ABCF_EF-3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000467">
        <property type="entry name" value="ATP-binding cassette sub-family F member 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:001092">
        <property type="entry name" value="ATP-binding cassette sub-family F member 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:004204">
        <property type="entry name" value="Multidrug resistance protein, putative"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003593">
        <property type="entry name" value="AAA+_ATPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032781">
        <property type="entry name" value="ABC_tran_Xtn"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003439">
        <property type="entry name" value="ABC_transporter-like_ATP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017871">
        <property type="entry name" value="ABC_transporter-like_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050611">
        <property type="entry name" value="ABCF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19211:SF15">
        <property type="entry name" value="ATP-BINDING CASSETTE SUB-FAMILY F MEMBER 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19211">
        <property type="entry name" value="ATP-BINDING TRANSPORT PROTEIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00005">
        <property type="entry name" value="ABC_tran"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12848">
        <property type="entry name" value="ABC_tran_Xtn"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00382">
        <property type="entry name" value="AAA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00211">
        <property type="entry name" value="ABC_TRANSPORTER_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50893">
        <property type="entry name" value="ABC_TRANSPORTER_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0007">Acetylation</keyword>
    <keyword evidence="5" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="ABC transporter" evidence="9">
        <location>
            <begin position="86"/>
            <end position="325"/>
        </location>
    </feature>
    <feature type="domain" description="ABC transporter" evidence="9">
        <location>
            <begin position="396"/>
            <end position="570"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1"/>
            <end position="51"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="162"/>
            <end position="209"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00011054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50893"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016017.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="580" mass="66278" checksum="1774C1DAEFE60EEF" modified="2019-05-08" version="1">MPSDLAKKKAAKKKEAAKARQRPKKTPEENGDAEIEQQEMKNEEANGTEGSEIDALTKELEDFELKKAAARAVTGVLASHPNSTDVHIINLSLTFHGQELLSDTKLELNSGRRYGLIGLNGIGKSMLLSAIGKREVPIPEHIDIYHLTREMPPSEKTPLQCVMEVDTERAMLEREAERLAHEDAECEKLMELYERLEELDADKAEVRASRILYGLGFTPAMQRKKLKDFSGGWRMRVALARALFIRPFMLLLDEPTNHLDLDACVWLEEELKTFKRILVLISHSQDFLNGICTNIIHMHNRKLKYYTGNYDQYVKTRLELEENQMKRFHWEQDQIAHMKNYIARFGHGSAKLARQAQSKEKTLQKMMASGLTEKVVNDKTLSFYFPPCGKIPPPVIMVQNVSFKYMKDGPWIYNNLEFGIDLDTRVALVGPNGAGKSTLLKLLTGELLPTDGMIRKHSHVKIGRYHQVSPIRNLSDGQKCRVCFAWLAWQNPHMLFLDEPTNHLDIETIDALADAINEFEGGMMLVSHDFRLIQQVAQEIWVCEKQTIAKWQGDILAYKEHLKSKLVDEEPQLTKKTHNV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM09</accession>
    <name>A0A452HM09_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="12">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HM09"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018263.1">
        <property type="protein sequence ID" value="ENSGAGP00000015983.1"/>
        <property type="gene ID" value="ENSGAGG00000012025.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001228">
        <property type="term" value="F:DNA-binding transcription activator activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd07765">
        <property type="entry name" value="KRAB_A-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001833">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001639">
        <property type="entry name" value="Si:dkey-7i4.21"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002434">
        <property type="entry name" value="Zinc finger protein 1168"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000638">
        <property type="entry name" value="Zinc finger protein 184"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001823">
        <property type="entry name" value="Zinc finger protein 322"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002343">
        <property type="entry name" value="Zinc finger protein 33A"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002090">
        <property type="entry name" value="Zinc finger protein 473"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000737">
        <property type="entry name" value="Zinc finger protein 565"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002134">
        <property type="entry name" value="Zinc finger protein 616"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001909">
        <property type="entry name" value="KRAB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036051">
        <property type="entry name" value="KRAB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24376">
        <property type="entry name" value="ZINC FINGER PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24376:SF238">
        <property type="entry name" value="ZINC FINGER PROTEIN 850"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01352">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="SMART" id="SM00349">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109640">
        <property type="entry name" value="KRAB domain (Kruppel-associated box)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50805">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="14"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="2" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <keyword evidence="4 8" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="KRAB" evidence="11">
        <location>
            <begin position="28"/>
            <end position="99"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="10">
        <location>
            <begin position="131"/>
            <end position="158"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="10">
        <location>
            <begin position="159"/>
            <end position="186"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="10">
        <location>
            <begin position="187"/>
            <end position="214"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="10">
        <location>
            <begin position="215"/>
            <end position="242"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="10">
        <location>
            <begin position="243"/>
            <end position="270"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="10">
        <location>
            <begin position="271"/>
            <end position="298"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="10">
        <location>
            <begin position="299"/>
            <end position="326"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="10">
        <location>
            <begin position="327"/>
            <end position="354"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="10">
        <location>
            <begin position="355"/>
            <end position="382"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="10">
        <location>
            <begin position="383"/>
            <end position="410"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="10">
        <location>
            <begin position="411"/>
            <end position="438"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="10">
        <location>
            <begin position="439"/>
            <end position="466"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="10">
        <location>
            <begin position="467"/>
            <end position="494"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="10">
        <location>
            <begin position="495"/>
            <end position="522"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="85"/>
            <end position="113"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015983.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="535" mass="61537" checksum="7CFED658030F5B28" modified="2019-05-08" version="1">FYRWALYLPPSSLLESPPVGLLLFQGPVTFEEVAVYFTSEEWALLDPTQRALYWDVMQENYENVTSLGFPVSKPHIISQLEQGAEPWKEEQNSQQGNVEQVAKHRELSQRSKRNVSRREKMGKFISWERPYGCCQCGKTFTFRSNLVHHMRIHTGKRPYKCSECGKSFITSSDLSKHQRIHTGERPFECSECGKTFAFRSVLSKHQRIHTGKRPYKCSECGKSFITSSVLSQHQRIHTGERPYECLECGKSFITSSDLSQHQRIHTGERPYKCSECGKSFITSSDLSKHQRIHTGKRPYKCSECGKSFITSSVLSQHQRIHTGERPYECRECGKSFSSSSALSHHQRIHTGERPFECSDCGKTFAFRSVLSKHQRIHTGERPYKCSECGKSFTQSSQLTEHQRIHTGERPFECSDCGKTFTYRSVLSKHQRIHTGERPYKCHECGKSFTRSSHLTGHQRIHTGERPYKCHECGESFTRSSHLSGHQRIHTGERPYKCGKCGKTFIHRQELISHQRIHTGMRPNGSSECGKTLSCH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="22" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM10</accession>
    <name>A0A452HM10_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HM10"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018241.1">
        <property type="protein sequence ID" value="ENSGAGP00000015965.1"/>
        <property type="gene ID" value="ENSGAGG00000012007.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="1" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="1">
        <location>
            <begin position="64"/>
            <end position="88"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="1">
        <location>
            <begin position="109"/>
            <end position="134"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015965.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="170" mass="18860" checksum="4D755676F1D64A17" modified="2019-05-08" version="1">MVRVTSPLDFIPNSSSLLLSRVKISPLYSQFYCYDPVVGSVKAAESPVWSCGELSLQKGEHRGAVLSLGFFCFAIVFCALIQAAFWRYTNNSEVSVFLLRNTKNQSPDGDPIAVTASLSCLLLFLPLTAGAVLLQGKQGVYGMILTSFLWFSIHFSNNLDRRGKYIFRET</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM13</accession>
    <name>A0A452HM13_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">PNPLA domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="caution">
        <text evidence="2">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HM13"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018264.1">
        <property type="protein sequence ID" value="ENSGAGP00000015984.1"/>
        <property type="gene ID" value="ENSGAGG00000012001.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005811">
        <property type="term" value="C:lipid droplet"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004806">
        <property type="term" value="F:triacylglycerol lipase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055088">
        <property type="term" value="P:lipid homeostasis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019433">
        <property type="term" value="P:triglyceride catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.1090.10:FF:000016">
        <property type="entry name" value="Patatin like phospholipase domain containing 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.1090.10">
        <property type="entry name" value="Cytosolic phospholipase A2 catalytic domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016035">
        <property type="entry name" value="Acyl_Trfase/lysoPLipase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033562">
        <property type="entry name" value="PLPL"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002641">
        <property type="entry name" value="PNPLA_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12406">
        <property type="entry name" value="CALCIUM-INDEPENDENT PHOSPHOLIPASE A2 IPLA2 -RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12406:SF23">
        <property type="entry name" value="OMEGA-HYDROXYCERAMIDE TRANSACYLASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01734">
        <property type="entry name" value="Patatin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52151">
        <property type="entry name" value="FabD/lysophospholipase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51635">
        <property type="entry name" value="PNPLA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="2" id="KW-0442">Lipid degradation</keyword>
    <keyword evidence="1 2" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="PNPLA" evidence="3">
        <location>
            <begin position="15"/>
            <end position="177"/>
        </location>
    </feature>
    <feature type="short sequence motif" description="GXSXG" evidence="2">
        <location>
            <begin position="50"/>
            <end position="54"/>
        </location>
    </feature>
    <feature type="short sequence motif" description="DGA/G" evidence="2">
        <location>
            <begin position="164"/>
            <end position="166"/>
        </location>
    </feature>
    <feature type="active site" description="Nucleophile" evidence="2">
        <location>
            <position position="52"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="2">
        <location>
            <position position="164"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01161"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS51635"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015984.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="270" mass="30088" checksum="D831E241C9FA5590" modified="2019-05-08" version="1">MAEADRNGTGTPFSLSFSGSGFLALYQIGVVQSLLELAPEILKSASKVYGASAGSLIAAAVDLKESFHEAVRDARKTILGPLSPGCNMLCTIQKRLYKMLPENSHQVASGRLHISLTRVMDGQNVMVSEFSSKEDLIQALICSCFVPIYCGFIPPSFRGVRYVDGGFTNLQPCSDLEAVITVSPFTGELDICPRDCPAIFYSFQIFNSSIQISVENLCRISYALFPPSYLFFFRGYQDAVLFLRRNSKWYSEPVKRQEAFSVWHCFPYRE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM14</accession>
    <name>A0A452HM14_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="14">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HM14"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018268.1">
        <property type="protein sequence ID" value="ENSGAGP00000015988.1"/>
        <property type="gene ID" value="ENSGAGG00000012025.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000785">
        <property type="term" value="C:chromatin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031519">
        <property type="term" value="C:PcG protein complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005667">
        <property type="term" value="C:transcription regulator complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd07765">
        <property type="entry name" value="KRAB_A-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001833">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002434">
        <property type="entry name" value="Zinc finger protein 1168"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000638">
        <property type="entry name" value="Zinc finger protein 184"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001823">
        <property type="entry name" value="Zinc finger protein 322"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002343">
        <property type="entry name" value="Zinc finger protein 33A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001498">
        <property type="entry name" value="Zinc finger protein 404"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002090">
        <property type="entry name" value="Zinc finger protein 473"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000737">
        <property type="entry name" value="Zinc finger protein 565"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002134">
        <property type="entry name" value="Zinc finger protein 616"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001909">
        <property type="entry name" value="KRAB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036051">
        <property type="entry name" value="KRAB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14003">
        <property type="entry name" value="TRANSCRIPTIONAL REPRESSOR PROTEIN YY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14003:SF23">
        <property type="entry name" value="ZINC FINGER PROTEIN 143"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01352">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="SMART" id="SM00349">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109640">
        <property type="entry name" value="KRAB domain (Kruppel-associated box)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50805">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="14"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="9" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 11" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="KRAB" evidence="13">
        <location>
            <begin position="10"/>
            <end position="93"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="71"/>
            <end position="93"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="94"/>
            <end position="121"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="122"/>
            <end position="149"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="150"/>
            <end position="177"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="185"/>
            <end position="212"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="213"/>
            <end position="240"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="241"/>
            <end position="268"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="269"/>
            <end position="296"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="297"/>
            <end position="324"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="325"/>
            <end position="352"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="353"/>
            <end position="380"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="381"/>
            <end position="408"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="409"/>
            <end position="436"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="437"/>
            <end position="464"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015988.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="477" mass="54497" checksum="144D6C74AC1C87D9" modified="2019-05-08" version="1">DASVSARGPVTFEEVAVYFTSEEWALLDPTQRALYWDVMQENYENVTSLGKGYCPLSSWKGKWRVPRKQGCGKTFSCSSSLSVHQRIHTGERPYGCCQCGKTFTFRSNLVHHMRIHTGKRPYKCSECGKSFITSSDLSKHQRIHTGERPFECSECGKTFAFRSVLSKHQRIHTGKRPYKCILSPYECLECGKSFITSSDLSQHQRIHTGERPYKCSECGKSFITSSDLSKHQRIHTGKRPYKCSECGKSFITSSVLSQHQRIHTGERPYECRECGKSFSSSSALSHHQRIHTGERPFECSDCGKTFAFRSVLSKHQRIHTGERPYKCSECGKSFTQSSQLTEHQRIHTGERPFECSDCGKTFTYRSVLSKHQRIHTGERPYKCHECGKSFTRSSHLTGHQRIHTGERPYKCHECGESFTRSSHLSGHQRIHTGERPYKCGKCGKTFIHRQELISHQRIHTGMRPNGSSECGKTLSCH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM17</accession>
    <name>A0A452HM17_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="13">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HM17"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018261.1">
        <property type="protein sequence ID" value="ENSGAGP00000015981.1"/>
        <property type="gene ID" value="ENSGAGG00000012025.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043565">
        <property type="term" value="F:sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd07765">
        <property type="entry name" value="KRAB_A-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001833">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002434">
        <property type="entry name" value="Zinc finger protein 1168"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000638">
        <property type="entry name" value="Zinc finger protein 184"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001823">
        <property type="entry name" value="Zinc finger protein 322"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002343">
        <property type="entry name" value="Zinc finger protein 33A"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002090">
        <property type="entry name" value="Zinc finger protein 473"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000052">
        <property type="entry name" value="zinc finger protein 546 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000737">
        <property type="entry name" value="Zinc finger protein 565"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002134">
        <property type="entry name" value="Zinc finger protein 616"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001909">
        <property type="entry name" value="KRAB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036051">
        <property type="entry name" value="KRAB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24394">
        <property type="entry name" value="ZINC FINGER PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24394:SF48">
        <property type="entry name" value="ZINC FINGER PROTEIN 771"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01352">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="SMART" id="SM00349">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109640">
        <property type="entry name" value="KRAB domain (Kruppel-associated box)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50805">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="14"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="2" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9" id="KW-0539">Nucleus</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="8" id="KW-0804">Transcription</keyword>
    <keyword evidence="6" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <keyword evidence="4 10" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="KRAB" evidence="12">
        <location>
            <begin position="18"/>
            <end position="94"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="11">
        <location>
            <begin position="89"/>
            <end position="116"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="11">
        <location>
            <begin position="117"/>
            <end position="144"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="11">
        <location>
            <begin position="145"/>
            <end position="172"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="11">
        <location>
            <begin position="173"/>
            <end position="200"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="11">
        <location>
            <begin position="201"/>
            <end position="228"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="11">
        <location>
            <begin position="229"/>
            <end position="256"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="11">
        <location>
            <begin position="257"/>
            <end position="284"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="11">
        <location>
            <begin position="285"/>
            <end position="312"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="11">
        <location>
            <begin position="313"/>
            <end position="340"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="11">
        <location>
            <begin position="341"/>
            <end position="368"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="11">
        <location>
            <begin position="369"/>
            <end position="396"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="11">
        <location>
            <begin position="397"/>
            <end position="424"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="11">
        <location>
            <begin position="425"/>
            <end position="452"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="11">
        <location>
            <begin position="453"/>
            <end position="480"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
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    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015981.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="493" mass="56328" checksum="782191EE7BBA693B" modified="2019-05-08" version="1">FLQEGEGHVGLLLFQGPVTFEEVAVYFTSEEWALLDPTQRALYWDVMQENYENVTSLGKGYCPLSSWKGKWRGKETTTQQEILMGKRKDTCTECGKTFSCSSSLSVHQRIHTGERPYGCCQCGKTFTFRSNLVHHMRIHTGKRPYKCSECGKSFITSSDLSKHQRIHTGERPFECSECGKTFAFRSVLSKHQRIHTGKRPYKCSECGKSFITSSVLSQHQRIHTGERPYECLECGKSFITSSDLSQHQRIHTGERPYKCSECGKSFITSSDLSKHQRIHTGKRPYKCSECGKSFITSSVLSQHQRIHTGERPYECRECGKSFSSSSALSHHQRIHTGERPFECSDCGKTFAFRSVLSKHQRIHTGERPYKCSECGKSFTQSSQLTEHQRIHTGERPFECSDCGKTFTYRSVLSKHQRIHTGERPYKCHECGKSFTRSSHLTGHQRIHTGERPYKCGKCGKTFIHRQELISHQRIHTGMRPNGSSECGKTLSCH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM18</accession>
    <name>A0A452HM18_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Fatty acid synthase</fullName>
            <ecNumber evidence="5">1.1.1.100</ecNumber>
            <ecNumber evidence="2">1.3.1.39</ecNumber>
            <ecNumber evidence="8">2.3.1.38</ecNumber>
            <ecNumber evidence="9">2.3.1.39</ecNumber>
            <ecNumber evidence="7">2.3.1.41</ecNumber>
            <ecNumber evidence="4">2.3.1.85</ecNumber>
            <ecNumber evidence="3">3.1.2.14</ecNumber>
            <ecNumber evidence="6">4.2.1.59</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="73 74">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="74" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="73" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="73" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="36">Fatty acid synthetase is a multifunctional enzyme that catalyzes the de novo biosynthesis of long-chain saturated fatty acids starting from acetyl-CoA and malonyl-CoA in the presence of NADPH. This multifunctional protein contains 7 catalytic activities and a site for the binding of the prosthetic group 4'-phosphopantetheine of the acyl carrier protein ([ACP]) domain.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="43">
            <text>(2E)-butenoyl-[ACP] + NADPH + H(+) = butanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41812"/>
            <dbReference type="Rhea" id="RHEA-COMP:9627"/>
            <dbReference type="Rhea" id="RHEA-COMP:9628"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78453"/>
            <dbReference type="ChEBI" id="CHEBI:78454"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="43">
            <dbReference type="Rhea" id="RHEA:41813"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="67">
            <text>(2E)-decenoyl-[ACP] + NADPH + H(+) = decanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41864"/>
            <dbReference type="Rhea" id="RHEA-COMP:9639"/>
            <dbReference type="Rhea" id="RHEA-COMP:9640"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78467"/>
            <dbReference type="ChEBI" id="CHEBI:78468"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="67">
            <dbReference type="Rhea" id="RHEA:41865"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="50">
            <text>(2E)-dodecenoyl-[ACP] + NADPH + H(+) = dodecanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41880"/>
            <dbReference type="Rhea" id="RHEA-COMP:9643"/>
            <dbReference type="Rhea" id="RHEA-COMP:9644"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:65264"/>
            <dbReference type="ChEBI" id="CHEBI:78472"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="50">
            <dbReference type="Rhea" id="RHEA:41881"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="45">
            <text>(2E)-hexadecenoyl-[ACP] + NADPH + H(+) = hexadecanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41912"/>
            <dbReference type="Rhea" id="RHEA-COMP:9651"/>
            <dbReference type="Rhea" id="RHEA-COMP:9652"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78481"/>
            <dbReference type="ChEBI" id="CHEBI:78483"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="45">
            <dbReference type="Rhea" id="RHEA:41913"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="46">
            <text>(2E)-hexenoyl-[ACP] + NADPH + H(+) = hexanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41832"/>
            <dbReference type="Rhea" id="RHEA-COMP:9631"/>
            <dbReference type="Rhea" id="RHEA-COMP:9632"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78458"/>
            <dbReference type="ChEBI" id="CHEBI:78459"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="46">
            <dbReference type="Rhea" id="RHEA:41833"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="60">
            <text>(2E)-octadecenoyl-[ACP] + NADPH + H(+) = octadecanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41928"/>
            <dbReference type="Rhea" id="RHEA-COMP:9655"/>
            <dbReference type="Rhea" id="RHEA-COMP:9656"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78489"/>
            <dbReference type="ChEBI" id="CHEBI:78495"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="60">
            <dbReference type="Rhea" id="RHEA:41929"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="53">
            <text>(2E)-octenoyl-[ACP] + NADPH + H(+) = octanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41848"/>
            <dbReference type="Rhea" id="RHEA-COMP:9635"/>
            <dbReference type="Rhea" id="RHEA-COMP:9636"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78462"/>
            <dbReference type="ChEBI" id="CHEBI:78463"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="53">
            <dbReference type="Rhea" id="RHEA:41849"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="62">
            <text>(2E)-tetradecenoyl-[ACP] + NADPH + H(+) = tetradecanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41896"/>
            <dbReference type="Rhea" id="RHEA-COMP:9647"/>
            <dbReference type="Rhea" id="RHEA-COMP:9648"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78475"/>
            <dbReference type="ChEBI" id="CHEBI:78477"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="62">
            <dbReference type="Rhea" id="RHEA:41897"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="35">
            <text>(3R)-hydroxybutanoyl-[ACP] = (2E)-butenoyl-[ACP] + H2O</text>
            <dbReference type="Rhea" id="RHEA:41808"/>
            <dbReference type="Rhea" id="RHEA-COMP:9626"/>
            <dbReference type="Rhea" id="RHEA-COMP:9627"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:78451"/>
            <dbReference type="ChEBI" id="CHEBI:78453"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="35">
            <dbReference type="Rhea" id="RHEA:41809"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="30">
            <text>(3R)-hydroxydecanoyl-[ACP] = (2E)-decenoyl-[ACP] + H2O</text>
            <dbReference type="Rhea" id="RHEA:41860"/>
            <dbReference type="Rhea" id="RHEA-COMP:9638"/>
            <dbReference type="Rhea" id="RHEA-COMP:9639"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:78466"/>
            <dbReference type="ChEBI" id="CHEBI:78467"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="30">
            <dbReference type="Rhea" id="RHEA:41861"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="28">
            <text>(3R)-hydroxydodecanoyl-[ACP] = (2E)-dodecenoyl-[ACP] + H2O</text>
            <dbReference type="Rhea" id="RHEA:41876"/>
            <dbReference type="Rhea" id="RHEA-COMP:9642"/>
            <dbReference type="Rhea" id="RHEA-COMP:9643"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:78470"/>
            <dbReference type="ChEBI" id="CHEBI:78472"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="28">
            <dbReference type="Rhea" id="RHEA:41877"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="34">
            <text>(3R)-hydroxyhexadecanoyl-[ACP] = (2E)-hexadecenoyl-[ACP] + H2O</text>
            <dbReference type="Rhea" id="RHEA:41908"/>
            <dbReference type="Rhea" id="RHEA-COMP:9650"/>
            <dbReference type="Rhea" id="RHEA-COMP:9651"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:78480"/>
            <dbReference type="ChEBI" id="CHEBI:78481"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="34">
            <dbReference type="Rhea" id="RHEA:41909"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="29">
            <text>(3R)-hydroxyhexanoyl-[ACP] = (2E)-hexenoyl-[ACP] + H2O</text>
            <dbReference type="Rhea" id="RHEA:41828"/>
            <dbReference type="Rhea" id="RHEA-COMP:9630"/>
            <dbReference type="Rhea" id="RHEA-COMP:9631"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:78457"/>
            <dbReference type="ChEBI" id="CHEBI:78458"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="29">
            <dbReference type="Rhea" id="RHEA:41829"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="33">
            <text>(3R)-hydroxyoctadecanoyl-[ACP] = (2E)-octadecenoyl-[ACP] + H2O</text>
            <dbReference type="Rhea" id="RHEA:41924"/>
            <dbReference type="Rhea" id="RHEA-COMP:9654"/>
            <dbReference type="Rhea" id="RHEA-COMP:9655"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:78488"/>
            <dbReference type="ChEBI" id="CHEBI:78489"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="33">
            <dbReference type="Rhea" id="RHEA:41925"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="27">
            <text>(3R)-hydroxyoctanoyl-[ACP] = (2E)-octenoyl-[ACP] + H2O</text>
            <dbReference type="Rhea" id="RHEA:41844"/>
            <dbReference type="Rhea" id="RHEA-COMP:9634"/>
            <dbReference type="Rhea" id="RHEA-COMP:9635"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:78461"/>
            <dbReference type="ChEBI" id="CHEBI:78462"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="27">
            <dbReference type="Rhea" id="RHEA:41845"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="32">
            <text>(3R)-hydroxytetradecanoyl-[ACP] = (2E)-tetradecenoyl-[ACP] + H2O</text>
            <dbReference type="Rhea" id="RHEA:41892"/>
            <dbReference type="Rhea" id="RHEA-COMP:9646"/>
            <dbReference type="Rhea" id="RHEA-COMP:9647"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:78474"/>
            <dbReference type="ChEBI" id="CHEBI:78475"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="32">
            <dbReference type="Rhea" id="RHEA:41893"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="47">
            <text>3-oxobutanoyl-[ACP] + NADPH + H(+) = (3R)-hydroxybutanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41804"/>
            <dbReference type="Rhea" id="RHEA-COMP:9625"/>
            <dbReference type="Rhea" id="RHEA-COMP:9626"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78450"/>
            <dbReference type="ChEBI" id="CHEBI:78451"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="47">
            <dbReference type="Rhea" id="RHEA:41805"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="41">
            <text>3-oxodecanoyl-[ACP] + NADPH + H(+) = (3R)-hydroxydecanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41856"/>
            <dbReference type="Rhea" id="RHEA-COMP:9637"/>
            <dbReference type="Rhea" id="RHEA-COMP:9638"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78464"/>
            <dbReference type="ChEBI" id="CHEBI:78466"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="41">
            <dbReference type="Rhea" id="RHEA:41857"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="63">
            <text>3-oxododecanoyl-[ACP] + NADPH + H(+) = (3R)-hydroxydodecanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41872"/>
            <dbReference type="Rhea" id="RHEA-COMP:9641"/>
            <dbReference type="Rhea" id="RHEA-COMP:9642"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78469"/>
            <dbReference type="ChEBI" id="CHEBI:78470"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="63">
            <dbReference type="Rhea" id="RHEA:41873"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="64">
            <text>3-oxohexadecanoyl-[ACP] + NADPH + H(+) = (3R)-hydroxyhexadecanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41904"/>
            <dbReference type="Rhea" id="RHEA-COMP:9649"/>
            <dbReference type="Rhea" id="RHEA-COMP:9650"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78478"/>
            <dbReference type="ChEBI" id="CHEBI:78480"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="64">
            <dbReference type="Rhea" id="RHEA:41905"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="55">
            <text>3-oxohexanoyl-[ACP] + NADPH + H(+) = (3R)-hydroxyhexanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41824"/>
            <dbReference type="Rhea" id="RHEA-COMP:9629"/>
            <dbReference type="Rhea" id="RHEA-COMP:9630"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78456"/>
            <dbReference type="ChEBI" id="CHEBI:78457"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="55">
            <dbReference type="Rhea" id="RHEA:41825"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="38">
            <text>3-oxooctadecanoyl-[ACP] + NADPH + H(+) = (3R)-hydroxyoctadecanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41920"/>
            <dbReference type="Rhea" id="RHEA-COMP:9653"/>
            <dbReference type="Rhea" id="RHEA-COMP:9654"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78487"/>
            <dbReference type="ChEBI" id="CHEBI:78488"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="38">
            <dbReference type="Rhea" id="RHEA:41921"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="65">
            <text>3-oxooctanoyl-[ACP] + NADPH + H(+) = (3R)-hydroxyoctanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41840"/>
            <dbReference type="Rhea" id="RHEA-COMP:9633"/>
            <dbReference type="Rhea" id="RHEA-COMP:9634"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78460"/>
            <dbReference type="ChEBI" id="CHEBI:78461"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="65">
            <dbReference type="Rhea" id="RHEA:41841"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="59">
            <text>3-oxotetradecanoyl-[ACP] + NADPH + H(+) = (3R)-hydroxytetradecanoyl-[ACP] + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:41888"/>
            <dbReference type="Rhea" id="RHEA-COMP:9645"/>
            <dbReference type="Rhea" id="RHEA-COMP:9646"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78473"/>
            <dbReference type="ChEBI" id="CHEBI:78474"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="59">
            <dbReference type="Rhea" id="RHEA:41889"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="40">
            <text>a (3R)-hydroxyacyl-[ACP] + NADP(+) = a 3-oxoacyl-[ACP] + NADPH + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17397"/>
            <dbReference type="Rhea" id="RHEA-COMP:9916"/>
            <dbReference type="Rhea" id="RHEA-COMP:9945"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78776"/>
            <dbReference type="ChEBI" id="CHEBI:78827"/>
            <dbReference type="EC" id="1.1.1.100"/>
        </reaction>
        <physiologicalReaction direction="right-to-left" evidence="40">
            <dbReference type="Rhea" id="RHEA:17399"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="31">
            <text>a (3R)-hydroxyacyl-[ACP] = a (2E)-enoyl-[ACP] + H2O</text>
            <dbReference type="Rhea" id="RHEA:13097"/>
            <dbReference type="Rhea" id="RHEA-COMP:9925"/>
            <dbReference type="Rhea" id="RHEA-COMP:9945"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:78784"/>
            <dbReference type="ChEBI" id="CHEBI:78827"/>
            <dbReference type="EC" id="4.2.1.59"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="31">
            <dbReference type="Rhea" id="RHEA:13098"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="56">
            <text>a 2,3-saturated acyl-[ACP] + NADP(+) = a (2E)-enoyl-[ACP] + NADPH + H(+)</text>
            <dbReference type="Rhea" id="RHEA:22564"/>
            <dbReference type="Rhea" id="RHEA-COMP:9925"/>
            <dbReference type="Rhea" id="RHEA-COMP:9926"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:78784"/>
            <dbReference type="ChEBI" id="CHEBI:78785"/>
            <dbReference type="EC" id="1.3.1.39"/>
        </reaction>
        <physiologicalReaction direction="right-to-left" evidence="56">
            <dbReference type="Rhea" id="RHEA:22566"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="54">
            <text>a fatty acyl-[ACP] + malonyl-[ACP] + H(+) = a 3-oxoacyl-[ACP] + holo-[ACP] + CO2</text>
            <dbReference type="Rhea" id="RHEA:22836"/>
            <dbReference type="Rhea" id="RHEA-COMP:9623"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="Rhea" id="RHEA-COMP:9916"/>
            <dbReference type="Rhea" id="RHEA-COMP:14125"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78449"/>
            <dbReference type="ChEBI" id="CHEBI:78776"/>
            <dbReference type="ChEBI" id="CHEBI:138651"/>
            <dbReference type="EC" id="2.3.1.41"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="54">
            <dbReference type="Rhea" id="RHEA:22837"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="37">
            <text>acetyl-CoA + n malonyl-CoA + 2n NADPH + 2n H(+) = a long-chain fatty acid + (n+1) CoA + n CO2 + 2n NADP(+).</text>
            <dbReference type="EC" id="2.3.1.85"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="48">
            <text>acetyl-[ACP] + malonyl-[ACP] + H(+) = 3-oxobutanoyl-[ACP] + holo-[ACP] + CO2</text>
            <dbReference type="Rhea" id="RHEA:41800"/>
            <dbReference type="Rhea" id="RHEA-COMP:9621"/>
            <dbReference type="Rhea" id="RHEA-COMP:9623"/>
            <dbReference type="Rhea" id="RHEA-COMP:9625"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78446"/>
            <dbReference type="ChEBI" id="CHEBI:78449"/>
            <dbReference type="ChEBI" id="CHEBI:78450"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="48">
            <dbReference type="Rhea" id="RHEA:41801"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="66">
            <text>butanoyl-[ACP] + malonyl-[ACP] + H(+) = 3-oxohexanoyl-[ACP] + holo-[ACP] + CO2</text>
            <dbReference type="Rhea" id="RHEA:41820"/>
            <dbReference type="Rhea" id="RHEA-COMP:9623"/>
            <dbReference type="Rhea" id="RHEA-COMP:9628"/>
            <dbReference type="Rhea" id="RHEA-COMP:9629"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78449"/>
            <dbReference type="ChEBI" id="CHEBI:78454"/>
            <dbReference type="ChEBI" id="CHEBI:78456"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="66">
            <dbReference type="Rhea" id="RHEA:41821"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="61">
            <text>decanoyl-[ACP] + malonyl-[ACP] + H(+) = 3-oxododecanoyl-[ACP] + holo-[ACP] + CO2</text>
            <dbReference type="Rhea" id="RHEA:41868"/>
            <dbReference type="Rhea" id="RHEA-COMP:9623"/>
            <dbReference type="Rhea" id="RHEA-COMP:9640"/>
            <dbReference type="Rhea" id="RHEA-COMP:9641"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78449"/>
            <dbReference type="ChEBI" id="CHEBI:78468"/>
            <dbReference type="ChEBI" id="CHEBI:78469"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="61">
            <dbReference type="Rhea" id="RHEA:41869"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="44">
            <text>dodecanoyl-[ACP] + malonyl-[ACP] + H(+) = 3-oxotetradecanoyl-[ACP] + holo-[ACP] + CO2</text>
            <dbReference type="Rhea" id="RHEA:41884"/>
            <dbReference type="Rhea" id="RHEA-COMP:9623"/>
            <dbReference type="Rhea" id="RHEA-COMP:9644"/>
            <dbReference type="Rhea" id="RHEA-COMP:9645"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:65264"/>
            <dbReference type="ChEBI" id="CHEBI:78449"/>
            <dbReference type="ChEBI" id="CHEBI:78473"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="44">
            <dbReference type="Rhea" id="RHEA:41885"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="58">
            <text>hexadecanoyl-[ACP] + H2O = hexadecanoate + holo-[ACP] + H(+)</text>
            <dbReference type="Rhea" id="RHEA:41932"/>
            <dbReference type="Rhea" id="RHEA-COMP:9652"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:7896"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78483"/>
            <dbReference type="EC" id="3.1.2.14"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="58">
            <dbReference type="Rhea" id="RHEA:41933"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="49">
            <text>hexadecanoyl-[ACP] + malonyl-[ACP] + H(+) = 3-oxooctadecanoyl-[ACP] + holo-[ACP] + CO2</text>
            <dbReference type="Rhea" id="RHEA:41916"/>
            <dbReference type="Rhea" id="RHEA-COMP:9623"/>
            <dbReference type="Rhea" id="RHEA-COMP:9652"/>
            <dbReference type="Rhea" id="RHEA-COMP:9653"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78449"/>
            <dbReference type="ChEBI" id="CHEBI:78483"/>
            <dbReference type="ChEBI" id="CHEBI:78487"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="49">
            <dbReference type="Rhea" id="RHEA:41917"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="39">
            <text>hexanoyl-[ACP] + malonyl-[ACP] + H(+) = 3-oxooctanoyl-[ACP] + holo-[ACP] + CO2</text>
            <dbReference type="Rhea" id="RHEA:41836"/>
            <dbReference type="Rhea" id="RHEA-COMP:9623"/>
            <dbReference type="Rhea" id="RHEA-COMP:9632"/>
            <dbReference type="Rhea" id="RHEA-COMP:9633"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78449"/>
            <dbReference type="ChEBI" id="CHEBI:78459"/>
            <dbReference type="ChEBI" id="CHEBI:78460"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="39">
            <dbReference type="Rhea" id="RHEA:41837"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="57">
            <text>holo-[ACP] + acetyl-CoA = acetyl-[ACP] + CoA</text>
            <dbReference type="Rhea" id="RHEA:41788"/>
            <dbReference type="Rhea" id="RHEA-COMP:9621"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57288"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78446"/>
            <dbReference type="EC" id="2.3.1.38"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="57">
            <dbReference type="Rhea" id="RHEA:41789"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="52">
            <text>holo-[ACP] + malonyl-CoA = malonyl-[ACP] + CoA</text>
            <dbReference type="Rhea" id="RHEA:41792"/>
            <dbReference type="Rhea" id="RHEA-COMP:9623"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57384"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78449"/>
            <dbReference type="EC" id="2.3.1.39"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="52">
            <dbReference type="Rhea" id="RHEA:41793"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="68">
            <text>octanoyl-[ACP] + malonyl-[ACP] + H(+) = 3-oxodecanoyl-[ACP] + holo-[ACP] + CO2</text>
            <dbReference type="Rhea" id="RHEA:41852"/>
            <dbReference type="Rhea" id="RHEA-COMP:9623"/>
            <dbReference type="Rhea" id="RHEA-COMP:9636"/>
            <dbReference type="Rhea" id="RHEA-COMP:9637"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78449"/>
            <dbReference type="ChEBI" id="CHEBI:78463"/>
            <dbReference type="ChEBI" id="CHEBI:78464"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="68">
            <dbReference type="Rhea" id="RHEA:41853"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="51">
            <text>tetradecanoyl-[ACP] + H2O = tetradecanoate + holo-[ACP] + H(+)</text>
            <dbReference type="Rhea" id="RHEA:30123"/>
            <dbReference type="Rhea" id="RHEA-COMP:9648"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30807"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78477"/>
            <dbReference type="EC" id="3.1.2.14"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="51">
            <dbReference type="Rhea" id="RHEA:30124"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="42">
            <text>tetradecanoyl-[ACP] + malonyl-[ACP] + H(+) = 3-oxohexadecanoyl-[ACP] + holo-[ACP] + CO2</text>
            <dbReference type="Rhea" id="RHEA:41900"/>
            <dbReference type="Rhea" id="RHEA-COMP:9623"/>
            <dbReference type="Rhea" id="RHEA-COMP:9648"/>
            <dbReference type="Rhea" id="RHEA-COMP:9649"/>
            <dbReference type="Rhea" id="RHEA-COMP:9685"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:64479"/>
            <dbReference type="ChEBI" id="CHEBI:78449"/>
            <dbReference type="ChEBI" id="CHEBI:78477"/>
            <dbReference type="ChEBI" id="CHEBI:78478"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="42">
            <dbReference type="Rhea" id="RHEA:41901"/>
        </physiologicalReaction>
    </comment>
    <comment type="pathway">
        <text evidence="1">Lipid metabolism; fatty acid biosynthesis.</text>
    </comment>
    <dbReference type="EC" id="1.1.1.100" evidence="5"/>
    <dbReference type="EC" id="1.3.1.39" evidence="2"/>
    <dbReference type="EC" id="2.3.1.38" evidence="8"/>
    <dbReference type="EC" id="2.3.1.39" evidence="9"/>
    <dbReference type="EC" id="2.3.1.41" evidence="7"/>
    <dbReference type="EC" id="2.3.1.85" evidence="4"/>
    <dbReference type="EC" id="3.1.2.14" evidence="3"/>
    <dbReference type="EC" id="4.2.1.59" evidence="6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018273.1">
        <property type="protein sequence ID" value="ENSGAGP00000015993.1"/>
        <property type="gene ID" value="ENSGAGG00000012026.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00094"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019171">
        <property type="term" value="F:(3R)-hydroxyacyl-[acyl-carrier-protein] dehydratase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004316">
        <property type="term" value="F:3-oxoacyl-[acyl-carrier-protein] reductase (NADPH) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004315">
        <property type="term" value="F:3-oxoacyl-[acyl-carrier-protein] synthase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004313">
        <property type="term" value="F:[acyl-carrier-protein] S-acetyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004314">
        <property type="term" value="F:[acyl-carrier-protein] S-malonyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0141148">
        <property type="term" value="F:enoyl-[acyl-carrier-protein] reductase (NADPH) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004312">
        <property type="term" value="F:fatty acid synthase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016297">
        <property type="term" value="F:fatty acyl-[ACP] hydrolase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031177">
        <property type="term" value="F:phosphopantetheine binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006633">
        <property type="term" value="P:fatty acid biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="CDD" id="cd05195">
        <property type="entry name" value="enoyl_red"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd08954">
        <property type="entry name" value="KR_1_FAS_SDR_x"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="CDD" id="cd00833">
        <property type="entry name" value="PKS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.1200.10:FF:000013">
        <property type="entry name" value="Fatty acid synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.129.110:FF:000002">
        <property type="entry name" value="Fatty acid synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.366.10:FF:000005">
        <property type="entry name" value="Fatty acid synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.47.10:FF:000033">
        <property type="entry name" value="Fatty acid synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1820:FF:000059">
        <property type="entry name" value="Fatty acid synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.180.10:FF:000015">
        <property type="entry name" value="Fatty acid synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.720:FF:000209">
        <property type="entry name" value="Polyketide synthase Pks12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.3290">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.47.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1200.10">
        <property type="entry name" value="ACP-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1820">
        <property type="entry name" value="alpha/beta hydrolase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.366.10">
        <property type="entry name" value="Malonyl-Coenzyme A Acyl Carrier Protein, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.180.10">
        <property type="entry name" value="Medium-chain alcohol dehydrogenases, catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.720">
        <property type="entry name" value="NAD(P)-binding Rossmann-like Domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.129.110">
        <property type="entry name" value="Polyketide synthase dehydratase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.150">
        <property type="entry name" value="Vaccinia Virus protein VP39"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029058">
        <property type="entry name" value="AB_hydrolase_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001227">
        <property type="entry name" value="Ac_transferase_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036736">
        <property type="entry name" value="ACP-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014043">
        <property type="entry name" value="Acyl_transferase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016035">
        <property type="entry name" value="Acyl_Trfase/lysoPLipase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013149">
        <property type="entry name" value="ADH-like_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020843">
        <property type="entry name" value="ER"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049391">
        <property type="entry name" value="FAS_pseudo-KR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011032">
        <property type="entry name" value="GroES-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018201">
        <property type="entry name" value="Ketoacyl_synth_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014031">
        <property type="entry name" value="Ketoacyl_synth_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014030">
        <property type="entry name" value="Ketoacyl_synth_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057326">
        <property type="entry name" value="KR_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016036">
        <property type="entry name" value="Malonyl_transacylase_ACP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036291">
        <property type="entry name" value="NAD(P)-bd_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032821">
        <property type="entry name" value="PKS_assoc"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020841">
        <property type="entry name" value="PKS_Beta-ketoAc_synthase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042104">
        <property type="entry name" value="PKS_dehydratase_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020807">
        <property type="entry name" value="PKS_DH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049552">
        <property type="entry name" value="PKS_DH_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013968">
        <property type="entry name" value="PKS_KR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049900">
        <property type="entry name" value="PKS_mFAS_DH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050091">
        <property type="entry name" value="PKS_NRPS_Biosynth_Enz"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020806">
        <property type="entry name" value="PKS_PP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009081">
        <property type="entry name" value="PP-bd_ACP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006162">
        <property type="entry name" value="Ppantetheine_attach_site"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029063">
        <property type="entry name" value="SAM-dependent_MTases_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001031">
        <property type="entry name" value="Thioesterase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016039">
        <property type="entry name" value="Thiolase-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43775">
        <property type="entry name" value="FATTY ACID SYNTHASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43775:SF7">
        <property type="entry name" value="FATTY ACID SYNTHASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00698">
        <property type="entry name" value="Acyl_transf_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00107">
        <property type="entry name" value="ADH_zinc_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21149">
        <property type="entry name" value="FAS_pseudo-KR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16197">
        <property type="entry name" value="KAsynt_C_assoc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00109">
        <property type="entry name" value="ketoacyl-synt"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02801">
        <property type="entry name" value="Ketoacyl-synt_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08659">
        <property type="entry name" value="KR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21089">
        <property type="entry name" value="PKS_DH_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00550">
        <property type="entry name" value="PP-binding"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00975">
        <property type="entry name" value="Thioesterase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00827">
        <property type="entry name" value="PKS_AT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00826">
        <property type="entry name" value="PKS_DH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00829">
        <property type="entry name" value="PKS_ER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00822">
        <property type="entry name" value="PKS_KR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00825">
        <property type="entry name" value="PKS_KS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00823">
        <property type="entry name" value="PKS_PP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47336">
        <property type="entry name" value="ACP-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53474">
        <property type="entry name" value="alpha/beta-Hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52151">
        <property type="entry name" value="FabD/lysophospholipase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50129">
        <property type="entry name" value="GroES-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51735">
        <property type="entry name" value="NAD(P)-binding Rossmann-fold domains"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55048">
        <property type="entry name" value="Probable ACP-binding domain of malonyl-CoA ACP transacylase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53335">
        <property type="entry name" value="S-adenosyl-L-methionine-dependent methyltransferases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53901">
        <property type="entry name" value="Thiolase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50075">
        <property type="entry name" value="CARRIER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00606">
        <property type="entry name" value="KS3_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS52004">
        <property type="entry name" value="KS3_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00012">
        <property type="entry name" value="PHOSPHOPANTETHEINE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS52019">
        <property type="entry name" value="PKS_MFAS_DH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="20" id="KW-0007">Acetylation</keyword>
    <keyword evidence="24" id="KW-0275">Fatty acid biosynthesis</keyword>
    <keyword evidence="17" id="KW-0276">Fatty acid metabolism</keyword>
    <keyword evidence="16" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="12" id="KW-0444">Lipid biosynthesis</keyword>
    <keyword evidence="23" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="25" id="KW-0456">Lyase</keyword>
    <keyword evidence="26" id="KW-0511">Multifunctional enzyme</keyword>
    <keyword evidence="22" id="KW-0520">NAD</keyword>
    <keyword evidence="18" id="KW-0521">NADP</keyword>
    <keyword evidence="21" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="11" id="KW-0596">Phosphopantetheine</keyword>
    <keyword evidence="13" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="19" id="KW-0663">Pyridoxal phosphate</keyword>
    <keyword evidence="74" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="15" id="KW-0702">S-nitrosylation</keyword>
    <keyword evidence="14" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Ketosynthase family 3 (KS3)" evidence="71">
        <location>
            <begin position="1"/>
            <end position="406"/>
        </location>
    </feature>
    <feature type="domain" description="PKS/mFAS DH" evidence="72">
        <location>
            <begin position="844"/>
            <end position="1111"/>
        </location>
    </feature>
    <feature type="domain" description="Carrier" evidence="70">
        <location>
            <begin position="2043"/>
            <end position="2120"/>
        </location>
    </feature>
    <feature type="region of interest" description="N-terminal hotdog fold" evidence="69">
        <location>
            <begin position="844"/>
            <end position="967"/>
        </location>
    </feature>
    <feature type="region of interest" description="C-terminal hotdog fold" evidence="69">
        <location>
            <begin position="984"/>
            <end position="1111"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor; for dehydratase activity" evidence="69">
        <location>
            <position position="878"/>
        </location>
    </feature>
    <feature type="active site" description="Proton donor; for dehydratase activity" evidence="69">
        <location>
            <position position="1034"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00005194"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00012004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012480"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00012873"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012948"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00013167"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00013191"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00013256"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00013258"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00018769"/>
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    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022450"/>
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    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022516"/>
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    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
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    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
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    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022799"/>
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    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
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    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022832"/>
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    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00022857"/>
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    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00022898"/>
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    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
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    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
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    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00023027"/>
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    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
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    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00023160"/>
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    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00023239"/>
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    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00023268"/>
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    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="ARBA" id="ARBA00023332"/>
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    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="ARBA" id="ARBA00023351"/>
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    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="ARBA" id="ARBA00023373"/>
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    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="ARBA" id="ARBA00023388"/>
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    </evidence>
    <evidence type="ECO:0000256" key="31">
        <source>
            <dbReference type="ARBA" id="ARBA00023394"/>
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    </evidence>
    <evidence type="ECO:0000256" key="32">
        <source>
            <dbReference type="ARBA" id="ARBA00023398"/>
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    </evidence>
    <evidence type="ECO:0000256" key="33">
        <source>
            <dbReference type="ARBA" id="ARBA00023399"/>
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    </evidence>
    <evidence type="ECO:0000256" key="34">
        <source>
            <dbReference type="ARBA" id="ARBA00023401"/>
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    </evidence>
    <evidence type="ECO:0000256" key="35">
        <source>
            <dbReference type="ARBA" id="ARBA00023402"/>
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    </evidence>
    <evidence type="ECO:0000256" key="36">
        <source>
            <dbReference type="ARBA" id="ARBA00023442"/>
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    </evidence>
    <evidence type="ECO:0000256" key="37">
        <source>
            <dbReference type="ARBA" id="ARBA00044883"/>
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    </evidence>
    <evidence type="ECO:0000256" key="38">
        <source>
            <dbReference type="ARBA" id="ARBA00047300"/>
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    </evidence>
    <evidence type="ECO:0000256" key="39">
        <source>
            <dbReference type="ARBA" id="ARBA00047394"/>
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    </evidence>
    <evidence type="ECO:0000256" key="40">
        <source>
            <dbReference type="ARBA" id="ARBA00047400"/>
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    </evidence>
    <evidence type="ECO:0000256" key="41">
        <source>
            <dbReference type="ARBA" id="ARBA00047440"/>
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    </evidence>
    <evidence type="ECO:0000256" key="42">
        <source>
            <dbReference type="ARBA" id="ARBA00047451"/>
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    </evidence>
    <evidence type="ECO:0000256" key="43">
        <source>
            <dbReference type="ARBA" id="ARBA00047500"/>
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    </evidence>
    <evidence type="ECO:0000256" key="44">
        <source>
            <dbReference type="ARBA" id="ARBA00047578"/>
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    </evidence>
    <evidence type="ECO:0000256" key="45">
        <source>
            <dbReference type="ARBA" id="ARBA00047810"/>
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    </evidence>
    <evidence type="ECO:0000256" key="46">
        <source>
            <dbReference type="ARBA" id="ARBA00047897"/>
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    </evidence>
    <evidence type="ECO:0000256" key="47">
        <source>
            <dbReference type="ARBA" id="ARBA00047953"/>
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    </evidence>
    <evidence type="ECO:0000256" key="48">
        <source>
            <dbReference type="ARBA" id="ARBA00047961"/>
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    </evidence>
    <evidence type="ECO:0000256" key="49">
        <source>
            <dbReference type="ARBA" id="ARBA00048051"/>
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    </evidence>
    <evidence type="ECO:0000256" key="50">
        <source>
            <dbReference type="ARBA" id="ARBA00048281"/>
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    </evidence>
    <evidence type="ECO:0000256" key="51">
        <source>
            <dbReference type="ARBA" id="ARBA00048289"/>
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    </evidence>
    <evidence type="ECO:0000256" key="52">
        <source>
            <dbReference type="ARBA" id="ARBA00048404"/>
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    </evidence>
    <evidence type="ECO:0000256" key="53">
        <source>
            <dbReference type="ARBA" id="ARBA00048420"/>
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    </evidence>
    <evidence type="ECO:0000256" key="54">
        <source>
            <dbReference type="ARBA" id="ARBA00048506"/>
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    </evidence>
    <evidence type="ECO:0000256" key="55">
        <source>
            <dbReference type="ARBA" id="ARBA00048571"/>
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    </evidence>
    <evidence type="ECO:0000256" key="56">
        <source>
            <dbReference type="ARBA" id="ARBA00048650"/>
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    </evidence>
    <evidence type="ECO:0000256" key="57">
        <source>
            <dbReference type="ARBA" id="ARBA00048691"/>
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    </evidence>
    <evidence type="ECO:0000256" key="58">
        <source>
            <dbReference type="ARBA" id="ARBA00048704"/>
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    </evidence>
    <evidence type="ECO:0000256" key="59">
        <source>
            <dbReference type="ARBA" id="ARBA00048935"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="60">
        <source>
            <dbReference type="ARBA" id="ARBA00049019"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="61">
        <source>
            <dbReference type="ARBA" id="ARBA00049109"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="62">
        <source>
            <dbReference type="ARBA" id="ARBA00049171"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="63">
        <source>
            <dbReference type="ARBA" id="ARBA00049263"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="64">
        <source>
            <dbReference type="ARBA" id="ARBA00049414"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="65">
        <source>
            <dbReference type="ARBA" id="ARBA00049422"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="66">
        <source>
            <dbReference type="ARBA" id="ARBA00049449"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="67">
        <source>
            <dbReference type="ARBA" id="ARBA00049521"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="68">
        <source>
            <dbReference type="ARBA" id="ARBA00049533"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="69">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01363"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="70">
        <source>
            <dbReference type="PROSITE" id="PS50075"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="71">
        <source>
            <dbReference type="PROSITE" id="PS52004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="72">
        <source>
            <dbReference type="PROSITE" id="PS52019"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="73">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015993.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="74">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2401" mass="264432" checksum="E0A33BB76CA7FF8E" modified="2019-05-08" version="1">MEDVVIAGMAGKLPESENMQEFWENLFNGVDMVTEDDRRWKPGLYGLPKRNGKLKDISKFDASFFGVHPKQAHTMDPQLRLMLEASYEAILDGGINPATLRGTDTGVWIGVSGSEAGEAFSQDPEQLLGYSMTGCQRAMFANRISYFFDLKGPSMAIDTACSSSLIALENAYKAICHGECNAALVGGVNLLLKPSTSVQFMKLGMLSPDGACKSFDASGNGYCRSEAVVVILMTKKSMAKRIYATIVNAGSNTDGFKEQGVTFPSGEMQQALVSSLYKESGIQPEEVEYVEAHGTGTKIGDPQELNGIANIFCQLEREPLLIGSTKSNMGHPEPASGLAALAKVILSLEHGLWAPNIHYNTPNPEIPALQDGRVQVVAKPTPVKGGLVSINSFGFGGSNVHVILRPNKKKFYPLERHNLPRLVQVCGRTQEAVEMLIEQSRRLREYSSFVSLLNDISGIPISSMPYRGYTLIGSQSDIKEVQHTRTSGRPLWYICSGMGTQWKGMGLSLMKLELFQQSILRSDEALKDTGLKVSDLLLNADENTFEDAVHAFVGLAAIQIAQIDILKAMDLQPDGIVGHSVGELACGYADNSLSHEEAILAAYWRGRCIKEAKLPPGGMAAVGLTWEECKRHCPPGVVPACHNSEDTVTVSGPQDAVNKFVAKLKKEGVFAKEVRSAGVAFHSYYMASIAPVLLSALQKVIPNAKPRSPRWISTSIPESQWQSELAQNSSAEYHVNNLVSPVLFQEGLRHVPDNAVVVEIAPHALLQAVLRRSLKPTCTILPLMKREHKNNLEFFLTHVGKIYLTGINVLSNNLFPLVEYPAPVGTPLISPYILWDHSQTWDVPKAEDFPTGSGGSSSASIYNIDMNSESSDNYLVGHCIDGRVLYPATGYLVLAWRTLARSLGIAMEQMPVVFEEVTIHQATILSNKGSTQLEVRLLPASHCFEVSGNGNLAVSGKICLLEETALNNFRNQQVDLETPPNMRSKFSLSKADIYKELRLRGYDYGPTFQGLLESNNEGNRGKVLWNGNWVTFLDTLLQMIILGETGHSLRLPTRIRSVCIDPSLHQERVHKYKDDTEAFDVVVDRCLDNIMAGGVRLSGLHATVAPRRQQEQRPPTLEKFCFVPYVESDCLSSEAHLCAHLEHCKGLICKLQEKVAFHGVKLAIPGLETVEAFGETVSMQKGLLHILAEICHLELNGNLYSELGQIVTREKMHLQDDPLLNALLDSPELKACVDIALENITSHRMKIVEALAGDGRLFSRVTGLLNTQPMLQLDYTATDQAPEILSVNESQLQETGVSIAQWDPISPPPGHLTSADLLVCNCSVSIFGNPAEILSNLAATVKEGGFILLHTLLKGETLGEIISFLTTPDLQQRGLLAQAEWEDLFRKTLLNLVAVKKSFFGSVIFLCRRQASAKIPVFLPVEETHYKWVESLKVILADASEQPVWLTATSCGNSGIVGMVNCLRQEPGGHRIRCLFVSNLNSSLAIPPTSPSAREMQKILQGDLVMNVYRDGKWGSFRHFLLQQAQPQELTDYAYVNVLTRGDLSSLRWISSPLRHFHTTNPNIQLCRVYYASLNFRDIMLATGKLPPDAIPGNWTLQQCMLGMEFSGRDPTGKRVMGLLPAKGLATVVDSDKRFLWEVPENWTLEEAASVPVVYATAYYALVVRGGMKKGESVLIHSGSGGVGQAAIAIALSMGCRVFATVGSVEKREYLQTRFPQLDANSFANSRNTSFEQHILRVTNGKGVNLVLNSLAEEKLQASLRCLARHGRFLEIGKYDLSNNSPLGMALFLKNVAFHGILLDAIFEEGNQEWEEVSDLLKMGIRDGVVKPLRTTIFNREEVEAAFRFMAQGKHIGKVMIKVHEEEKEFPARETEAVRLSAISRTSCPPTKSYIITGGLGGFGLELAQWLVERGAQKLVLTSRSGIRSGYQAKRVRGWKEMGVQVLVSTSDVGTLEGTQHLIEEASLLGPVGGIFNLAMVLKDAMLENQSPEFFHDVSKPKYAGTLHLDWVSREKCPDLDYFVAFSSISCGRGNVGQSNYGFANSTMERICEQRQHDGLPGVRDVSSLNAESSLADLGLDSLMGVEVRQTLERDYDLVMTMREIRLLTINKLRELSSKSGTAEELKPSLLMKAGQAEHPELDLNNLLVNPEGPTITRLNEVQSTERPLFLIHPIEGSITVFHTLASRLSMPSYGLQCTKAAPLDSIQSLAAYYIDCIKQVQPEGPYRIAGYSFGACVAFEMCSQLQAQQNPSHSLNSLFLFDGSHSYVAAYTQSYRAKLTPGNEAEAETEALCAFIQQFTSTEYNKLLETLLPLQDLEARVNAAVDLIMLSHKNINRETLCFAAASFYYKLKAADKYVPESKYHGNVTLLRAKTHNEYEEGLGGDYKLSEVSLRNSSRYEAALS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HM19</accession>
    <name>A0A452HM19_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Mitotic spindle organizing protein 2B</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
            <location evidence="2">Microtubule organizing center</location>
            <location evidence="2">Centrosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Spindle</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the MOZART2 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018310.1">
        <property type="protein sequence ID" value="ENSGAGP00000016027.1"/>
        <property type="gene ID" value="ENSGAGG00000012048.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005819">
        <property type="term" value="C:spindle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024332">
        <property type="entry name" value="MOZART2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR28578:SF2">
        <property type="entry name" value="MITOTIC-SPINDLE ORGANIZING PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR28578">
        <property type="entry name" value="MITOTIC-SPINDLE ORGANIZING PROTEIN 2A-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12926">
        <property type="entry name" value="MOZART2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="5" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="140"/>
            <end position="207"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="159"/>
            <end position="168"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="194"/>
            <end position="207"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004186"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000016027.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="9">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="207" mass="22127" checksum="EC6073EFAD9387BA" modified="2019-05-08" version="1">MLSRSSVPAVGERVSGLCRECPTRARPVAGRTRGWMRLWAKWGPRCGPGGSCRRRRRRSCSSWRRRPGSASTRKCSGGGGXXXXXXXXXXXXXXXXXXXARAGRAADPAPSPRRLLLDLLRMNVAPLAVFQMLKSLCAGQRPAPSGPAETGPEARGRNKTNPSVSGTQGLAERCSREGSVQRMPRQPSTSRLQKTGTSGKSSGGSST</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP11</accession>
    <name>A0A452IP11_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">small monomeric GTPase</fullName>
            <ecNumber evidence="5">3.6.5.2</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>GTP + H2O = GDP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:19669"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:37565"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:58189"/>
            <dbReference type="EC" id="3.6.5.2"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="13">
            <dbReference type="Rhea" id="RHEA:19670"/>
        </physiologicalReaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="pathway">
        <text evidence="3">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Membrane</location>
            <topology evidence="2">Lipid-anchor</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the small GTPase superfamily. Rab family.</text>
    </comment>
    <dbReference type="EC" id="3.6.5.2" evidence="5"/>
    <dbReference type="AlphaFoldDB" id="A0A452IP11"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033881.1">
        <property type="protein sequence ID" value="ENSGAGP00000029841.1"/>
        <property type="gene ID" value="ENSGAGG00000021542.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00143"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003925">
        <property type="term" value="F:G protein activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005525">
        <property type="term" value="F:GTP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035556">
        <property type="term" value="P:intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016567">
        <property type="term" value="P:protein ubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="CDD" id="cd03742">
        <property type="entry name" value="SOCS_Rab40"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:001506">
        <property type="entry name" value="Ras-related protein Rab-40C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005225">
        <property type="entry name" value="Small_GTP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001806">
        <property type="entry name" value="Small_GTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050305">
        <property type="entry name" value="Small_GTPase_Rab"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001496">
        <property type="entry name" value="SOCS_box"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036036">
        <property type="entry name" value="SOCS_box-like_dom_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00231">
        <property type="entry name" value="small_GTP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47980">
        <property type="entry name" value="LD44762P"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00071">
        <property type="entry name" value="Ras"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07525">
        <property type="entry name" value="SOCS_box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00449">
        <property type="entry name" value="RASTRNSFRMNG"/>
    </dbReference>
    <dbReference type="SMART" id="SM00175">
        <property type="entry name" value="RAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00173">
        <property type="entry name" value="RAS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00174">
        <property type="entry name" value="RHO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00253">
        <property type="entry name" value="SOCS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00969">
        <property type="entry name" value="SOCS_box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF158235">
        <property type="entry name" value="SOCS box-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51419">
        <property type="entry name" value="RAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51421">
        <property type="entry name" value="RAS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50225">
        <property type="entry name" value="SOCS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0342">GTP-binding</keyword>
    <keyword evidence="11" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="8" id="KW-0460">Magnesium</keyword>
    <keyword evidence="6" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="10" id="KW-0564">Palmitate</keyword>
    <keyword evidence="12" id="KW-0636">Prenylation</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="SOCS box" evidence="15">
        <location>
            <begin position="167"/>
            <end position="220"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="236"/>
            <end position="273"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="262"/>
            <end position="273"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004635"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00006270"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00011984"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023139"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023289"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00047660"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50225"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029841.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="273" mass="31058" checksum="1BCC0A27F7896894" modified="2019-05-08" version="1">MLGKEKSWRAFKMERRNPLTPIAMLQHYVTWAPSLFQALGIDYKTTTILLDGRRVKLELWDTSGQGRFCTIFRSYSRGAQGILLVYDITNRWSFDGIDRWIKEIDEHAPGVPRILVGNRLHLAFKRQVPTEQARSYAEKNCMTFFEVSPLCNFNVTESFTELSRIVLMRHGMEKIWRPNRVFSLQDLCCRAIVSCTPVHLIDKLPLPVTIKSHLKSFSMANGMNAVMMHGRSYSLASSGGGSSSKGNSLKRSKSIRPPQSPPQNCSRNNCKIS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP12</accession>
    <name>A0A452IP12_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">Ras-GAP domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000033661.1">
        <property type="protein sequence ID" value="ENSGAGP00000029646.1"/>
        <property type="gene ID" value="ENSGAGG00000019812.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005938">
        <property type="term" value="C:cell cortex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903479">
        <property type="term" value="P:mitotic actomyosin contractile ring assembly actin filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05131">
        <property type="entry name" value="RasGAP_IQGAP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00201">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.506.10:FF:000004">
        <property type="entry name" value="IQ motif containing GTPase activating protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.190">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.506.10">
        <property type="entry name" value="GTPase Activation - p120gap, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000048">
        <property type="entry name" value="IQ_motif_EF-hand-BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000593">
        <property type="entry name" value="RasGAP_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023152">
        <property type="entry name" value="RasGAP_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001936">
        <property type="entry name" value="RasGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001202">
        <property type="entry name" value="WW_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149">
        <property type="entry name" value="RAS GTPASE-ACTIVATING PROTEIN WITH IQ MOTIF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149:SF12">
        <property type="entry name" value="RAS GTPASE-ACTIVATING-LIKE PROTEIN IQGAP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00612">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00616">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03836">
        <property type="entry name" value="RasGAP_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00015">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00323">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00456">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF143885">
        <property type="entry name" value="RGC domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00509">
        <property type="entry name" value="RAS_GTPASE_ACTIV_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50018">
        <property type="entry name" value="RAS_GTPASE_ACTIV_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01159">
        <property type="entry name" value="WW_DOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Ras-GAP" evidence="1">
        <location>
            <begin position="622"/>
            <end position="855"/>
        </location>
    </feature>
    <evidence type="ECO:0000259" key="1">
        <source>
            <dbReference type="PROSITE" id="PS50018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029646.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1282" mass="146606" checksum="C0B29DA8C75A7C23" modified="2019-05-08" version="1">MAAYDVMLGYCLPTQHLKKKIDHINTAIRNCDANKTLVALMKPEAQLPVVHPFAAAVYQTELFNLQQQNARYLAHDELSIAVEMLSAVVLLNQALGSKDILAIKNHLRNPSVGFNNVEDENFQRYADTLLSIKSEASSQGQDYLSWNDVQNCIDMVNMQVQEENDRIVAVSYINEAIDQGNPGKTLETLLLPTVKLHDVNPTNARHYQDVLHDAKVQKCKESHDESAVLWLDEIQKGINDANSNIEKAANLALGTSMINNCLEKHDSQPVLDILQSPKFGLRVVPECAETYYKNLLEAKNLKTKEDSSEGPWLKLIMKNRYDYYYNVETGESTFVPPEGFIPKTSWLTSEEIQTIVGQVTADYNREQLWFANENLILQLQALARGFLVRKSYEERKGFLQKQEPSVIKIQASWKGYKQRKSYTDRLRVLQGNVAAIVKVSCSMLSKVFSLWSFFQNEEIVKIQAFLRANKARDDYRTLIGAENPPLNVLRKFAYLLDQSDLDFQEELEVTRLREEVVTKIRSNQQLEKDLNLMDIKIGLLVKNRISLQDVVLHSKKLNKKSKNQVEEMVTGDKQGIKGLSKERRKQLEAYQHLFYLLQTNPMYLAKLIFQMPQNKSTKFMDTVIFTLYNYASNQREEYLLLKLFKTALEEEINSKVDQIQDIVTGNPTVIKMVVSFNRGARGQNSLRQLLAPVVKEIMDDKSLLINTSPVEVYKAWVNQLETQTGEASKLPYDVTTEQALEHPEVVNRLESSIQSLRTVTDKVLVSIFSSLSMMPYGMRYIAKVLKNSLHEKFPDATEDELLKIVGNLLYYRYMNPAIVAPDGFDIIDMTAGGQIHSDQRRNLGCVAKVLQHAASNKLFEGESAHLSSMNTYLSQTYQKFRNFFQIACDVPEPEEKFNIDEYSDMVTLSKPVIYISIEEIINTHSLLLEHQDAIAPEQNDLLNELLKGLGAVPDVESFLGEGAVDPNDPNKENTLSQLAKTEISLSLTSKYELQEGEDQDLKGLMIKTKRLVVDVIRTQPGDTLLEILEIPASAQQESEHLKVVEKRAILDSKTPEKMKHSQSMLEDGQLPIEQKKRKIQRNLRTLEQAGLVSSANKYQEIINEIAKDIRNQRRYRHHRKAELVKLQQTLSALQSKTAFYEDQVNYYNTYIKTCLDNLTRKNSRRSIKLDGKVEEKGSKKLKHTSLKYTAARLHEKGVILEIEDLQTNQFKNVMFDITPGEEMGDFEIKAKFLGVQMEKVELHFQDLLQMQYEGVAVMKMFDKAKVNVNLLIFLLNKKFYGK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP14</accession>
    <name>A0A452IP14_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">PWWP domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IP14"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033599.1">
        <property type="protein sequence ID" value="ENSGAGP00000029589.1"/>
        <property type="gene ID" value="ENSGAGG00000021370.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="CDD" id="cd20150">
        <property type="entry name" value="PWWP_HDGFL3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.140:FF:000017">
        <property type="entry name" value="hepatoma-derived growth factor isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000313">
        <property type="entry name" value="PWWP_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12550:SF82">
        <property type="entry name" value="HDGF LIKE 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12550">
        <property type="entry name" value="HEPATOMA-DERIVED GROWTH FACTOR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00855">
        <property type="entry name" value="PWWP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00293">
        <property type="entry name" value="PWWP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF63748">
        <property type="entry name" value="Tudor/PWWP/MBT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50812">
        <property type="entry name" value="PWWP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="PWWP" evidence="2">
        <location>
            <begin position="11"/>
            <end position="68"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="97"/>
            <end position="203"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="100"/>
            <end position="109"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="1">
        <location>
            <begin position="121"/>
            <end position="130"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="1">
        <location>
            <begin position="135"/>
            <end position="160"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="169"/>
            <end position="178"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="188"/>
            <end position="203"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="PROSITE" id="PS50812"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029589.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="203" mass="22592" checksum="3D660BBD5A55BF62" modified="2019-05-08" version="1">MARPRPREYKAGDLVFAKMKGYPHWPARIDELPEGAVKPPANKYPIFFFGTHETAFLGPKDLFPYKEYKDKFGKSNKRKGFNEGLWEIENNPGVKFTGYQAIQQQSSSETEGEGGNTADASSEEEGDRVEEDGKGKRKNEKAGSKRKKSYSSKKSSKQSRKSPGDEDDKDCKEEENKSSSDAGDAGNDTRNTSSDLQKTSEGT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP15</accession>
    <name>A0A452IP15_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Mitogen-activated protein kinase kinase kinase 7</fullName>
            <ecNumber evidence="5">2.7.11.25</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="30 31">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="31" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="30" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="30" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="25">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.25"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="24">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.25"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Peripheral membrane protein</topology>
            <orientation evidence="2">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the protein kinase superfamily. STE Ser/Thr protein kinase family. MAP kinase kinase kinase subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.25" evidence="5"/>
    <dbReference type="AlphaFoldDB" id="A0A452IP15"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033639.1">
        <property type="protein sequence ID" value="ENSGAGP00000029625.1"/>
        <property type="gene ID" value="ENSGAGG00000021391.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000287">
        <property type="term" value="F:magnesium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004709">
        <property type="term" value="F:MAP kinase kinase kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019901">
        <property type="term" value="F:protein kinase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005102">
        <property type="term" value="F:signaling receptor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006915">
        <property type="term" value="P:apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071560">
        <property type="term" value="P:cellular response to transforming growth factor beta stimulus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006955">
        <property type="term" value="P:immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007254">
        <property type="term" value="P:JNK cascade"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043123">
        <property type="term" value="P:positive regulation of canonical NF-kappaB signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043410">
        <property type="term" value="P:positive regulation of MAPK cascade"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009893">
        <property type="term" value="P:positive regulation of metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd14058">
        <property type="entry name" value="STKc_TAK1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000143">
        <property type="entry name" value="Mitogen-activated protein kinase kinase kinase 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000152">
        <property type="entry name" value="Mitogen-activated protein kinase kinase kinase 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049637">
        <property type="entry name" value="MAP3K7"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001245">
        <property type="entry name" value="Ser-Thr/Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46716">
        <property type="entry name" value="MITOGEN-ACTIVATED PROTEIN KINASE KINASE KINASE 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46716:SF1">
        <property type="entry name" value="MITOGEN-ACTIVATED PROTEIN KINASE KINASE KINASE 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07714">
        <property type="entry name" value="PK_Tyr_Ser-Thr"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF038168">
        <property type="entry name" value="MAPKKK7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00109">
        <property type="entry name" value="TYRKINASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="13" id="KW-0053">Apoptosis</keyword>
    <keyword evidence="17 26" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="7" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="27" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="8" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="16" id="KW-0418">Kinase</keyword>
    <keyword evidence="18" id="KW-0460">Magnesium</keyword>
    <keyword evidence="22" id="KW-0472">Membrane</keyword>
    <keyword evidence="14" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="15 26" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="11" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="31" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="21" id="KW-0346">Stress response</keyword>
    <keyword evidence="23" id="KW-0804">Transcription</keyword>
    <keyword evidence="20" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="12" id="KW-0808">Transferase</keyword>
    <keyword evidence="19" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="Protein kinase" evidence="29">
        <location>
            <begin position="26"/>
            <end position="281"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="28">
        <location>
            <begin position="291"/>
            <end position="333"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="28">
        <location>
            <begin position="348"/>
            <end position="397"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="28">
        <location>
            <begin position="423"/>
            <end position="476"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="27">
        <location>
            <begin position="514"/>
            <end position="541"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="28">
        <location>
            <begin position="296"/>
            <end position="322"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="28">
        <location>
            <begin position="427"/>
            <end position="442"/>
        </location>
    </feature>
    <feature type="binding site" evidence="26">
        <location>
            <position position="53"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004413"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00006529"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012406"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00017660"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022703"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00023016"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00047559"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00048329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="29">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="30">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029625.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="31">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="587" mass="65469" checksum="58DE820DA2B0D51E" modified="2019-05-08" version="1">MSAATSAEMIETPPVLNFEEIVYKEIEVEEVVGRGAFGVVCKAKWRGKDVAIKQIESESERKAFIVELRQLSRVNHPNIVKLYGACQNPVCLVMEYAEGGSLYNVLHGAEPLPYYTAAHAMSWCLQCSQGVAYLHSMKPKALIHRDLKPPNLLLVAGGTVLKICDFGTACDIQTHMTNNKGSAAWMAPEVFEGSNYSEKCDVFSWGIILWEVITRRKPFDEIGGPAFRIMWAVHNGTRPPLIKNLPKPIESLMTRCWSKDPSQRPSMEEIVKIMTHLMRYFPGADEPLQYPCQYSDEGQSNSATSTGSFMDMTSTNTSNKSDANMEPSDFQGTATNDTIKRLESKLAQQMKNSAKQSGDPGRLSLPPSRGSSVESLSDVRGRPQSTLVSGEGKRMSADMSEIEARITAGNGQQRRRSIQDLSVAVTESNQESRNSSRSSSPSVRMITTPGPTSEKPARSLPWTPDESADTNGSDNSIPMAYLTLDHQLQPLAPCPNSKESMAVFEQHCKMAQEYMKVQTEIALLLQRRQELIAELDQDEKDQQNTSRLVQEHKKLLDENKSLSTYYQQCKKQLEVIRNQQQKRQGTS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP16</accession>
    <name>A0A452IP16_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Tuberin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Catalytic component of the TSC-TBC complex, a multiprotein complex that acts as a negative regulator of the canonical mTORC1 complex, an evolutionarily conserved central nutrient sensor that stimulates anabolic reactions and macromolecule biosynthesis to promote cellular biomass generation and growth. Within the TSC-TBC complex, TSC2 acts as a GTPase-activating protein (GAP) for the small GTPase RHEB, a direct activator of the protein kinase activity of mTORC1. In absence of nutrients, the TSC-TBC complex inhibits mTORC1, thereby preventing phosphorylation of ribosomal protein S6 kinase (RPS6KB1 and RPS6KB2) and EIF4EBP1 (4E-BP1) by the mTORC1 signaling. The TSC-TBC complex is inactivated in response to nutrients, relieving inhibition of mTORC1. Involved in microtubule-mediated protein transport via its ability to regulate mTORC1 signaling. Also stimulates the intrinsic GTPase activity of the Ras-related proteins RAP1A and RAB5.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytosol</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="8">Lysosome membrane</location>
            <topology evidence="8">Peripheral membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000033665.1">
        <property type="protein sequence ID" value="ENSGAGP00000029651.1"/>
        <property type="gene ID" value="ENSGAGG00000021398.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005765">
        <property type="term" value="C:lysosomal membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045202">
        <property type="term" value="C:synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033596">
        <property type="term" value="C:TSC1-TSC2 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046627">
        <property type="term" value="P:negative regulation of insulin receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051898">
        <property type="term" value="P:negative regulation of phosphatidylinositol 3-kinase/protein kinase B signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032007">
        <property type="term" value="P:negative regulation of TOR signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030178">
        <property type="term" value="P:negative regulation of Wnt signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051726">
        <property type="term" value="P:regulation of cell cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016241">
        <property type="term" value="P:regulation of macroautophagy"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051056">
        <property type="term" value="P:regulation of small GTPase mediated signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.11210:FF:000004">
        <property type="entry name" value="Tuberin isoform X3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.11210">
        <property type="entry name" value="Rap/Ran-GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035974">
        <property type="entry name" value="Rap/Ran-GAP_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000331">
        <property type="entry name" value="Rap/Ran_GAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003913">
        <property type="entry name" value="Tuberin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018515">
        <property type="entry name" value="Tuberin-type_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027107">
        <property type="entry name" value="Tuberin/Ral-act_asu"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024584">
        <property type="entry name" value="Tuberin_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10063">
        <property type="entry name" value="TUBERIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10063:SF0">
        <property type="entry name" value="TUBERIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11864">
        <property type="entry name" value="DUF3384"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02145">
        <property type="entry name" value="Rap_GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03542">
        <property type="entry name" value="Tuberin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01431">
        <property type="entry name" value="TUBERIN"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF111347">
        <property type="entry name" value="Rap/Ran-GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50085">
        <property type="entry name" value="RAPGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="2" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="7" id="KW-0458">Lysosome</keyword>
    <keyword evidence="6" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="Rap-GAP" evidence="12">
        <location>
            <begin position="1503"/>
            <end position="1730"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1"/>
            <end position="34"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="931"/>
            <end position="964"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1044"/>
            <end position="1123"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1294"/>
            <end position="1320"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1369"/>
            <end position="1458"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1745"/>
            <end position="1765"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="1"/>
            <end position="16"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="17"/>
            <end position="30"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="936"/>
            <end position="964"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="1369"/>
            <end position="1385"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004514"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023228"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023765"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00054764"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00070662"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50085"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029651.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1781" mass="199125" checksum="0912D2756DCC0830" modified="2019-05-08" version="1">MAKQQNKDPGLKEKFKSLLGLGPSRPSSKSSEGKQTEFIITAEILKELSIECGLNNRIRAISQICEVAKTKKFEEHAVEAIWKVVADMLQPELPAEARHAVLHLLKAIIQGQGERLGILRAHFFKVIKDYPSNEDLHERLAVFKALTDNGRSITHLEEELAEFVLQWMDVGLSSEFLLVLVNLVKFNSCYLEDYVADMVHMICLLCIQTSSSVDIEVSLQVLDAVVCYNCLPSESLPVFIITLCRTINVKELCEPCWKLMRNLLGTHLGHSAIYNMCRIMEDRAYMTDAALLRGAVFFVGMALWGAHRLYSLKNSPTSVLPSFLKAMTCPNAVVSYEIVLSITRLIKKYGKELQAVTWDILLDIIERLLQQLQSIENQDLKSIVHALLTTVEELCDQNEFHGSEERFFELVERCADQRPESSLLNLITYKAQSIHPAKDGWIHNLKVLMERFFRNECRSAVRIKVLDVLSFVLSINRQFYEEELINVVVISQLAHIPEDKDHQVRKLATQLLVDLAEGCNTHHFNSLLDIIEKVAARSLWAPVDLEERDFLSHSAALEDVKTAVLGLLVILQTKLYSLPASHATRVYEMLINHIELHYKNVYCLPIASSIRLQAFDFLLMLRADSLHRLGLTNKDGAVRFSPYCLCDLVEPEKRVSEKKPAGTLSPPSGSPSVSSQNATLRMGYLPYSLLFRVLLQCLKQETDWKVLKLVLNKLPESLRYKVLFLTSPCKLDQLCSALCSMLTDKKTAERLHGTPDGFSRNDLHLAVVPVLTALISYHNYLDKAKQREMVYCLEHGLIYRCANQCVVALSVCSVEMPDIIIKALPVLIVKLTHISATANMAIPLLEFLSTLARLPHLYRNFAAEQYASVFAISLPYTNPSKFNQYIVCLAHHVIAMWFIRCRLPFRKDFVPYITKGLRSNVLLSFDDTPEKDSFRARSTSLNERPKSRIQTSITSSSLGSADENSMAQADDNLKNLHLELTETCLDMMARYVFSNFTAVPKRSPVGEFLLAGGRTKTWLVGNKLVTITTSVGTGTRSLLGLDSGELQSSTESSPDPVLQVRQTKEAPAKLESQAGQQVCRGSRNRVRSMSGGHALRVGALDSSAPQGLRSPAASAPRTEKATPGAHALLQKEKTNLAAYVPLLTQGWAEILVRRPTGNTSWLMSLENPLSPFSSDINNMPLQELSNALMAAERFKEHRETALYKSLSVPSSSLATGTSKPSLLQRSNTVASFSSMYQSSCQGKLHRSISWAESAVVLEEGSQLEVDLKETDSLVSQEFEDFKVVCSEQAMAEEKFSKPHNAYSRSSSTSSQEEKSLKSNELVAGGIPIGRVVTSEDGRTLEELSFQPSQPLSKSSSSPELQTLQEVLKDTNAKEATGRLSTELKSKSQSGNLEGEGASSWLSRGDDAGITGTDRLDCNVPSSSPRSPSGHRPRGYTISDSAPSRRGKRIERDAFKSRTAASNAEKVPGINPSFVFLQLYHSPFFGDESNKPLLLPNETFESSVQLLDQIPSYDTHKIAVLYVGEGQSNNEIAILSNEHGSYRYTEFLTGLGKLIELKDCQPDKIYLGGLDVCGEDGQFTYCWHDDIMQAIFHIATLMPTKDLDKYRCDKKRHLGNDFVSIVYNDSGEDFKLGTIKGQFNFVHVIITPLDYDCNLVTLQCRKDMEGLVDTSVAKIVSDKNLPFVARQMALHANMASQVHHSRSNPTDTYPSKWIARLRHIKRLRHRIREETQYQTPGFPLMQMHPSALTKPPPQVPQDPASTYETGQRKRLISSVDDFTEFV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP17</accession>
    <name>A0A452IP17_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IP17"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033889.1">
        <property type="protein sequence ID" value="ENSGAGP00000029846.1"/>
        <property type="gene ID" value="ENSGAGG00000021549.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000033893.1">
        <property type="protein sequence ID" value="ENSGAGP00000029850.1"/>
        <property type="gene ID" value="ENSGAGG00000021549.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005783">
        <property type="term" value="C:endoplasmic reticulum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006506">
        <property type="term" value="P:GPI anchor biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007720">
        <property type="entry name" value="PigQ/GPI1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21329:SF3">
        <property type="entry name" value="PHOSPHATIDYLINOSITOL N-ACETYLGLUCOSAMINYLTRANSFERASE SUBUNIT Q"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21329">
        <property type="entry name" value="PHOSPHATIDYLINOSITOL N-ACETYLGLUCOSAMINYLTRANSFERASE SUBUNIT Q-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05024">
        <property type="entry name" value="Gpi1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="1" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="1">
        <location>
            <begin position="203"/>
            <end position="222"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="1">
        <location>
            <begin position="273"/>
            <end position="292"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="1">
        <location>
            <begin position="338"/>
            <end position="357"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="1">
        <location>
            <begin position="364"/>
            <end position="391"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="1">
        <location>
            <begin position="397"/>
            <end position="419"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="1">
        <location>
            <begin position="440"/>
            <end position="464"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="1">
        <location>
            <begin position="470"/>
            <end position="495"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029846.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="576" mass="66166" checksum="8E7695B532CB4DCF" modified="2019-05-08" version="1">MVLKVFFPSCCATAESGLLIGRWIPEQNSAVVLAVIHFPFIPGQVKQYLSEVQRVTQVNMCVLGSWSNSKQEKEEALDKFLEDLSTIFSHEPWIQLSKEGDSKFWSCCIFQKYPSSPKEHEFILVYYDQRKVMLSQLHPTDSLTRSADHKGNDASKLGAVFDTVAKSKVLFMTDRYDDGPIKLTHWQSEGVEASIIVELMKQASVPVCVLLTFLLSLVSSVCKSRVLKFWPLSFIWSKLSTCEQLGHRLEHLQIISSNRKAQNQTQLMRKANIFVSLLIDVALGILLMSWLYRENRIGQLADTLIPVADHIAKELQDLLQWLMGVPAGLKMNRALDQVLGRFFLYHIHLWISYIHLMSPFIEMILWYVGLSACLGLTVALSILSDIIALLTFHIYCFYVYGARLYCLKIYGLSSLWRLFRGKKWNVLRQRVDSCSYDLDQLFIGTLLFTILLFLLPTTALYYLVFTLLRLLVVVVQGLIHLLVDLIDSLPLYSVILRLCRSYRLAASVKFRVLEQKPGKPLRLLMQINPLSYGSVVQTYRLTSYACYPKDSWSSLCKKLFVGELIYPWKHKEEKQD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP19</accession>
    <name>A0A452IP19_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">E3 ubiquitin-protein ligase RNF170</fullName>
            <ecNumber evidence="4">2.3.2.27</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="16">RING finger protein 170</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="15">RING-type E3 ubiquitin transferase RNF170</fullName>
        </alternativeName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>S-ubiquitinyl-[E2 ubiquitin-conjugating enzyme]-L-cysteine + [acceptor protein]-L-lysine = [E2 ubiquitin-conjugating enzyme]-L-cysteine + N(6)-ubiquitinyl-[acceptor protein]-L-lysine.</text>
            <dbReference type="EC" id="2.3.2.27"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="3">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Endoplasmic reticulum membrane</location>
            <topology evidence="2">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="2.3.2.27" evidence="4"/>
    <dbReference type="AlphaFoldDB" id="A0A452IP19"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033643.1">
        <property type="protein sequence ID" value="ENSGAGP00000029629.1"/>
        <property type="gene ID" value="ENSGAGG00000021394.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000033644.1">
        <property type="protein sequence ID" value="ENSGAGP00000029630.1"/>
        <property type="gene ID" value="ENSGAGG00000021394.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061630">
        <property type="term" value="F:ubiquitin protein ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd16553">
        <property type="entry name" value="RING-HC_RNF170"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010652">
        <property type="entry name" value="DUF1232"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038896">
        <property type="entry name" value="RNF170"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027370">
        <property type="entry name" value="Znf-RING_euk"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001841">
        <property type="entry name" value="Znf_RING"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017907">
        <property type="entry name" value="Znf_RING_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22894:SF1">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE RNF170"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22894">
        <property type="entry name" value="RING-TYPE DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06803">
        <property type="entry name" value="DUF1232"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13445">
        <property type="entry name" value="zf-RING_UBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00184">
        <property type="entry name" value="RING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57850">
        <property type="entry name" value="RING/U-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00518">
        <property type="entry name" value="ZF_RING_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50089">
        <property type="entry name" value="ZF_RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="11" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="14 18" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0808">Transferase</keyword>
    <keyword evidence="7 18" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="13 18" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="10" id="KW-0833">Ubl conjugation pathway</keyword>
    <keyword evidence="12" id="KW-0862">Zinc</keyword>
    <keyword evidence="9 17" id="KW-0863">Zinc-finger</keyword>
    <feature type="transmembrane region" description="Helical" evidence="18">
        <location>
            <begin position="26"/>
            <end position="45"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="18">
        <location>
            <begin position="187"/>
            <end position="211"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="18">
        <location>
            <begin position="223"/>
            <end position="246"/>
        </location>
    </feature>
    <feature type="domain" description="RING-type" evidence="19">
        <location>
            <begin position="87"/>
            <end position="130"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004477"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00012483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00014068"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00030110"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00031107"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029630.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="256" mass="29527" checksum="256058A470413848" modified="2019-05-08" version="1">MASQQAEVQSLKLDNDSVIEGVSDQVLVAVVLSFTLIAALVYMLLRNEHQNIHPDNQELVRALRQQLQTDQDASAGDRHRFYTDMSCPVCLQQATFPIETNCGHLFCGACIIAYWRYGSWLGAIRCPICRQTVTLFLPLFREDQQDAAQVLQDVNDYNRRFSGQPRSIMERIMDLPTLLRHAFREMFTVGGLFWMFRLRIFLCLLGALLYLASPLDFLPEALFGILGFLDDFFVIFLLLIYISIMYREVITQRLNR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP20</accession>
    <name>A0A452IP20_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Amino acid transporter transmembrane domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IP20"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033672.1">
        <property type="protein sequence ID" value="ENSGAGP00000029656.1"/>
        <property type="gene ID" value="ENSGAGG00000021403.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015179">
        <property type="term" value="F:L-amino acid transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013057">
        <property type="entry name" value="AA_transpt_TM"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22950">
        <property type="entry name" value="AMINO ACID TRANSPORTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22950:SF222">
        <property type="entry name" value="SODIUM-COUPLED NEUTRAL AMINO ACID TRANSPORTER 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01490">
        <property type="entry name" value="Aa_trans"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4 5" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 5" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="3 5" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="105"/>
            <end position="127"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="152"/>
            <end position="173"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="194"/>
            <end position="215"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="221"/>
            <end position="242"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="367"/>
            <end position="392"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="412"/>
            <end position="434"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="454"/>
            <end position="471"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="477"/>
            <end position="499"/>
        </location>
    </feature>
    <feature type="domain" description="Amino acid transporter transmembrane" evidence="6">
        <location>
            <begin position="74"/>
            <end position="245"/>
        </location>
    </feature>
    <feature type="domain" description="Amino acid transporter transmembrane" evidence="6">
        <location>
            <begin position="325"/>
            <end position="505"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="Pfam" id="PF01490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029656.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="520" mass="58008" checksum="EE7DD74B8AD31E39" modified="2019-05-08" version="1">MDRMELQNVNIKLDEESNSGESIEDSYTDVIDPEKATISSQFANDDAESQKFLPNGFLGKKKLADYQEEYNPGTTSFGMSAFNLSNAIMGSGILGLSYAMANTGIILFVILLLSIAILSLYSVHLLLKTSKEGGSLIYEKLGEKAFGWPGKYGAFLSIIMQNIGAMSSYLFIIKYELPEVIRAFLHLEENSGEWYLNGNYLVILVSVGIILPLSLLKNLGYLGYTSGFSLTCMVFFLTVVIYKKSQIPCPLPVLENSIGNWTYNNSVPMHLVILPNESVNSRVNFMMDNTPGHTTDFEEAKDSLHTSGVEYEAHSDSDNMCQPKYFVFNSQTAYTIPILAFAFVCHPEVLPIYSELKDRSRKRMQNVSNISIAGMLIMYLLAALFGYLTFFGEVEDELLHTYTKVYTFDTPLLLVRLAVLVAVTLTVPIVLFPIRSSVIALLFPKRPFNWIRHFLIAAVILAFTNTLVIFVPAIKDIFGFIGASAATMLIFILPAAFYLRLVKKEPLRSAQKIGNLSSFT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP22</accession>
    <name>A0A452IP22_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Transformation/transcription domain associated protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="1 2">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="2" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="1" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="1" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IP22"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033894.1">
        <property type="protein sequence ID" value="ENSGAGP00000029851.1"/>
        <property type="gene ID" value="ENSGAGG00000021426.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035267">
        <property type="term" value="C:NuA4 histone acetyltransferase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000124">
        <property type="term" value="C:SAGA complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006281">
        <property type="term" value="P:DNA repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006355">
        <property type="term" value="P:regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050517">
        <property type="entry name" value="DDR_Repair_Kinase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11139">
        <property type="entry name" value="ATAXIA TELANGIECTASIA MUTATED ATM -RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11139:SF1">
        <property type="entry name" value="TRANSFORMATION_TRANSCRIPTION DOMAIN-ASSOCIATED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000313" key="1">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029851.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="3">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="267" mass="30970" checksum="DD831B2B7AAA6328" modified="2019-05-08" version="1">MAFVSTQGPTVVDQTTLMKKYLQFVAALTDVNTPDETKLKMMQEVSENFENVTSSPQYSTFLEHIIPRFLTFLQDGEVQFLQEKPAQQLRKLVLEIIHRIPTNEHLRPHTKNVLSVMFRFLETENEENVLICLRIIIELHKQFRPPITQEIHHFLDFVKQIYKELPKVVNRYFENPQVIPENTVPSPEMVGMITTIVVKVNPERDDSETRTHSIIPRGSLSLKVLAELPIIVVLMYQLYKLNIHNVVAEFVPLIMNTIIIQVSSQAR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP23</accession>
    <name>A0A452IP23_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Vacuolar protein sorting-associated protein 33A</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Plays a role in vesicle-mediated protein trafficking to lysosomal compartments including the endocytic membrane transport and autophagic pathways. Believed to act as a core component of the putative HOPS and CORVET endosomal tethering complexes which are proposed to be involved in the Rab5-to-Rab7 endosome conversion probably implicating MON1A/B, and via binding SNAREs and SNARE complexes to mediate tethering and docking events during SNARE-mediated membrane fusion. The HOPS complex is proposed to be recruited to Rab7 on the late endosomal membrane and to regulate late endocytic, phagocytic and autophagic traffic towards lysosomes. The CORVET complex is proposed to function as a Rab5 effector to mediate early endosome fusion probably in specific endosome subpopulations. Required for fusion of endosomes and autophagosomes with lysosomes; the function is dependent on its association with VPS16 but not VIPAS39. The function in autophagosome-lysosome fusion implicates STX17 but not UVRAG.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cytoplasmic vesicle</location>
            <location evidence="3">Autophagosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasmic vesicle</location>
            <location evidence="1">Clathrin-coated vesicle</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Early endosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Late endosome membrane</location>
            <topology evidence="4">Peripheral membrane protein</topology>
            <orientation evidence="4">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="5">Lysosome membrane</location>
            <topology evidence="5">Peripheral membrane protein</topology>
            <orientation evidence="5">Cytoplasmic side</orientation>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the STXBP/unc-18/SEC1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IP23"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033753.1">
        <property type="protein sequence ID" value="ENSGAGP00000029726.1"/>
        <property type="gene ID" value="ENSGAGG00000021422.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005776">
        <property type="term" value="C:autophagosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030136">
        <property type="term" value="C:clathrin-coated vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005769">
        <property type="term" value="C:early endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031902">
        <property type="term" value="C:late endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005765">
        <property type="term" value="C:lysosomal membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016192">
        <property type="term" value="P:vesicle-mediated transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.850:FF:000002">
        <property type="entry name" value="Vacuolar protein sorting-associated protein 33A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.830.10:FF:000006">
        <property type="entry name" value="Vacuolar protein sorting-associated protein 33A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1910:FF:000005">
        <property type="entry name" value="vacuolar protein sorting-associated protein 33A isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1910:FF:000008">
        <property type="entry name" value="vacuolar protein sorting-associated protein 33A isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.850">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1910">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.2060">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.830.10">
        <property type="entry name" value="Syntaxin Binding Protein 1, Chain A, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043154">
        <property type="entry name" value="Sec-1-like_dom1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043127">
        <property type="entry name" value="Sec-1-like_dom3a"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001619">
        <property type="entry name" value="Sec1-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027482">
        <property type="entry name" value="Sec1-like_dom2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036045">
        <property type="entry name" value="Sec1-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043155">
        <property type="entry name" value="VPS33_dom3b"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11679">
        <property type="entry name" value="VESICLE PROTEIN SORTING-ASSOCIATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00995">
        <property type="entry name" value="Sec1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56815">
        <property type="entry name" value="Sec1/munc18-like (SM) proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="8" id="KW-0967">Endosome</keyword>
    <keyword evidence="11" id="KW-0458">Lysosome</keyword>
    <keyword evidence="10" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-0653">Protein transport</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0813">Transport</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004132"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004412"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004419"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004492"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004630"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00009884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023228"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00039493"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00054535"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029726.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="598" mass="67626" checksum="AB589A081E3B480A" modified="2019-05-08" version="1">VATSVGCGRQSPACPSHCSRSVAFDVSSLGQAIVWDEYLTGPFGLIAQYSLLKEHEVEKMFTLKAGRLPPADVKNIIFLVRPRLELMDIIAENVLSEDRVRSPLRDFHILFVPRRSLLCEQRLKDQGVLGSFIHREQYSLDLIPFDGDLLSMESESAFKECYLENDQTSLYQAAKGLMTLQALYGTIPQIFGKGDCARHVANMMIRMKREFPGGQNSIFPIFDTLLLLDRNVDLLSPLATQLTYEGLIDEIYGIQNTYVKLPPEKFAPKKQGEGGKDLPTEPKKLQLNSAEELYAEIRDKNFNAVGSVLSKKAKIISAAFEERHHAKTVGEIKQFVSQLPHMQAARSSLANHTSIAELIKDITTSEDFFDNLTVEQEFMSGIDTDKVNNYTEDCIAQKHPLIKILRLVCLQSVCNSGLKQKVLDHYKREILQTYGYEHILTLNNLEKAGLLKLQTSSRNNYPTIRKTMRLWMDDVNEQNPNDISYVYSGYAPLSVRLAQLLARPGWRSIEEVLKMLPGPHFEERQQLPTGLQKKRQQGENRVTLVFFLGGVTYAEIAALRFLSQMEDGGTEYVIATTKLINGTNWIKSLMEKLEPPPF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP25</accession>
    <name>A0A452IP25_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Adhesion G-protein coupled receptor V1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="20">G-protein coupled receptor 98</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="19">Very large G-protein coupled receptor 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cell membrane</location>
            <topology evidence="3">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cell projection</location>
            <location evidence="1">Stereocilium membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Photoreceptor inner segment</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the G-protein coupled receptor 2 family. Adhesion G-protein coupled receptor (ADGR) subfamily.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000033611.1">
        <property type="protein sequence ID" value="ENSGAGP00000029599.1"/>
        <property type="gene ID" value="ENSGAGG00000021364.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001917">
        <property type="term" value="C:photoreceptor inner segment"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060171">
        <property type="term" value="C:stereocilium membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010855">
        <property type="term" value="F:adenylate cyclase inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004930">
        <property type="term" value="F:G protein-coupled receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001965">
        <property type="term" value="F:G-protein alpha-subunit binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016787">
        <property type="term" value="F:hydrolase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048513">
        <property type="term" value="P:animal organ development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007166">
        <property type="term" value="P:cell surface receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071277">
        <property type="term" value="P:cellular response to calcium ion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007605">
        <property type="term" value="P:sensory perception of sound"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007601">
        <property type="term" value="P:visual perception"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.2030:FF:000017">
        <property type="entry name" value="Adhesion G protein-coupled receptor V1"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.2030:FF:000020">
        <property type="entry name" value="Adhesion G protein-coupled receptor V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.2030:FF:000021">
        <property type="entry name" value="Adhesion G protein-coupled receptor V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.2030:FF:000023">
        <property type="entry name" value="Adhesion G protein-coupled receptor V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.2030:FF:000031">
        <property type="entry name" value="Adhesion G protein-coupled receptor V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.2030:FF:000046">
        <property type="entry name" value="Adhesion G protein-coupled receptor V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.220.50:FF:000020">
        <property type="entry name" value="Adhesion G-protein coupled receptor V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.2030:FF:000007">
        <property type="entry name" value="Adhesion G-protein coupled receptor V1"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.2030:FF:000009">
        <property type="entry name" value="adhesion G-protein coupled receptor V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.2030:FF:000012">
        <property type="entry name" value="Adhesion G-protein coupled receptor V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.2030:FF:000013">
        <property type="entry name" value="Adhesion G-protein coupled receptor V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.2030:FF:000014">
        <property type="entry name" value="Adhesion G-protein coupled receptor V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.2030:FF:000028">
        <property type="entry name" value="Adhesion G-protein coupled receptor V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.200">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.220.50">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.2030">
        <property type="match status" value="34"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1070.10">
        <property type="entry name" value="Rhodopsin 7-helix transmembrane proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026919">
        <property type="entry name" value="ADGRV1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038081">
        <property type="entry name" value="CalX-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003644">
        <property type="entry name" value="Calx_beta"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009039">
        <property type="entry name" value="EAR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005492">
        <property type="entry name" value="EPTP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057244">
        <property type="entry name" value="GAIN_B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046338">
        <property type="entry name" value="GAIN_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017981">
        <property type="entry name" value="GPCR_2-like_7TM"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46682">
        <property type="entry name" value="ADHESION G-PROTEIN COUPLED RECEPTOR V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46682:SF1">
        <property type="entry name" value="ADHESION G-PROTEIN COUPLED RECEPTOR V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03160">
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        <property type="match status" value="2"/>
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        <property type="match status" value="1"/>
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        <property type="match status" value="37"/>
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        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="EAR"/>
        <property type="match status" value="5"/>
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        <property type="match status" value="1"/>
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        <property type="entry name" value="GAIN_B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0106">Calcium</keyword>
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    <keyword evidence="12" id="KW-0297">G-protein coupled receptor</keyword>
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    <keyword evidence="13 21" id="KW-0472">Membrane</keyword>
    <keyword evidence="15" id="KW-0675">Receptor</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
    <keyword evidence="16" id="KW-0807">Transducer</keyword>
    <keyword evidence="6 21" id="KW-0812">Transmembrane</keyword>
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        <location>
            <begin position="27"/>
            <end position="46"/>
        </location>
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        <location>
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    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="5951"/>
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        <location>
            <begin position="5997"/>
            <end position="6018"/>
        </location>
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    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="6025"/>
            <end position="6049"/>
        </location>
    </feature>
    <feature type="domain" description="GAIN-B" evidence="22">
        <location>
            <begin position="5677"/>
            <end position="5836"/>
        </location>
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    <feature type="domain" description="G-protein coupled receptors family 2 profile 2" evidence="23">
        <location>
            <begin position="5885"/>
            <end position="6050"/>
        </location>
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    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004289"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004437"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00007343"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
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    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
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    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023224"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00070037"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00078072"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00083929"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS50221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS50261"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029599.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="6200" mass="680887" checksum="3A4FE43E8916F137" modified="2019-05-08" version="1">[encoded data, 6200 characters]</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP28</accession>
    <name>A0A452IP28_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Protein Wnt</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
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        <text evidence="11">Ligand for members of the frizzled family of seven transmembrane receptors.</text>
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        <subcellularLocation>
            <location evidence="1 11">Secreted</location>
            <location evidence="1 11">Extracellular space</location>
            <location evidence="1 11">Extracellular matrix</location>
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    </comment>
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        <text evidence="2 11">Belongs to the Wnt family.</text>
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        <property type="project" value="TreeGrafter"/>
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    </dbReference>
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        <property type="term" value="P:canonical Wnt signaling pathway"/>
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        <property type="project" value="TreeGrafter"/>
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        <property type="term" value="P:cell fate commitment"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="P:neuron differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd19336">
        <property type="entry name" value="Wnt_Wnt4"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Wnt homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
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        <property type="entry name" value="Wnt4"/>
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        <property type="entry name" value="Wnt_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12027:SF104">
        <property type="entry name" value="PROTEIN WNT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12027">
        <property type="entry name" value="WNT RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="WNT4PROTEIN"/>
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        <property type="entry name" value="WNTPROTEIN"/>
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        <property type="entry name" value="WNT1"/>
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        <property type="entry name" value="WNT1"/>
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    </dbReference>
    <proteinExistence type="inferred from homology"/>
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    <keyword evidence="8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="9" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
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        </source>
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        </source>
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        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="RuleBase" id="RU003500"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029732.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="373" mass="41126" checksum="3F4DD2DBC8EF63DA" modified="2019-05-08" version="1">MKPLESKASCVSLEWNCCRGRKARGSDVDSARIGTEEEERTTGIRPSDPTACICLSLIQPSLRGTLQDSSACEGLKGLAEEQVRICQRQVEAMDSVKHGAELAIEECQHQFHNRRWNCSTLQGLQVFGKVAIQGTRESAFIHAISSAGVAFAVTQACSRGDSLPASWTLSLSFPGFQWSGCSDNLSYGIAFSQAFVDNPERSRGVSSSRALMNLHNNEAGRKVKLVECKCHGVSGSCEVRTCWKAMPPFRKVGNVLKEKFEGATEVHPKRVGSRKLLVPKSSRFKPYTAHDLVYLVASPDFCDRDPHSGVFGTSGRQCNRTSHAMDGCELLCCGRGFRTAQAERVERCSCKFHWCCSVKCKQCHHLVEVHTCR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP29</accession>
    <name>A0A452IP29_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Adhesion G-protein coupled receptor V1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="20">G-protein coupled receptor 98</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="19">Very large G-protein coupled receptor 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cell membrane</location>
            <topology evidence="3">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cell projection</location>
            <location evidence="1">Stereocilium membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Photoreceptor inner segment</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the G-protein coupled receptor 2 family. Adhesion G-protein coupled receptor (ADGR) subfamily.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000033688.1">
        <property type="protein sequence ID" value="ENSGAGP00000029667.1"/>
        <property type="gene ID" value="ENSGAGG00000021364.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001917">
        <property type="term" value="C:photoreceptor inner segment"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060171">
        <property type="term" value="C:stereocilium membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010855">
        <property type="term" value="F:adenylate cyclase inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004930">
        <property type="term" value="F:G protein-coupled receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001965">
        <property type="term" value="F:G-protein alpha-subunit binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016787">
        <property type="term" value="F:hydrolase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
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        <property type="term" value="P:animal organ development"/>
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    </dbReference>
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        <property type="entry name" value="Laminin_G_3"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Calx_beta"/>
        <property type="match status" value="18"/>
    </dbReference>
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    </dbReference>
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        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
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    </dbReference>
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    </dbReference>
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    </dbReference>
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    <keyword evidence="5" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="17" id="KW-0966">Cell projection</keyword>
    <keyword evidence="14" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="12" id="KW-0297">G-protein coupled receptor</keyword>
    <keyword evidence="9" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="13 21" id="KW-0472">Membrane</keyword>
    <keyword evidence="15" id="KW-0675">Receptor</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
    <keyword evidence="16" id="KW-0807">Transducer</keyword>
    <keyword evidence="6 21" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 21" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="27"/>
            <end position="46"/>
        </location>
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    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="5841"/>
            <end position="5863"/>
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        <location>
            <begin position="5677"/>
            <end position="5836"/>
        </location>
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        </source>
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        </source>
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        </source>
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        <source>
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        </source>
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        </source>
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        </source>
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        </source>
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        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
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        <source>
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        </source>
    </evidence>
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        </source>
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        </source>
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        <source>
            <dbReference type="ARBA" id="ARBA00070037"/>
        </source>
    </evidence>
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        <source>
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        </source>
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        <source>
            <dbReference type="ARBA" id="ARBA00083929"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="PROSITE" id="PS50221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029667.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="5891" mass="646472" checksum="9AC3109436D5090E" modified="2019-05-08" version="1">[encoded data, 5891 characters]</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP30</accession>
    <name>A0A452IP30_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">CYRIA/CYRIB Rac1 binding domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the CYFIP family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IP30"/>
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    </dbReference>
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    </dbReference>
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    </dbReference>
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        <property type="entry name" value="CYTOFMRPINTP"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="CYRIA/CYRIB Rac1 binding" evidence="2">
        <location>
            <begin position="64"/>
            <end position="275"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
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        </source>
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        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="430" mass="49738" checksum="07D3BB612CAE1A18" modified="2019-05-08" version="1">MTAQVTLEDALSNVDLLEELPLPDQQPCIEPPPSSLLYQPNFNTNFEDRNAFVTGIARYIEQATVHSNMNEMLEDGQEYAVMLYTWRSCSRAIPQVKCNEQPNRVEIYEKTVEVLEPEVTKLMNFMYFQRNAIERFCGEVKRLCHAERRKDFVSEAYLITLGKFINMFAVLDELKNMKCSVKNDHSAYKRAAQFLRKMADPQSIQESQNLSMFLANHNKITQSLQQQLEVISGYEELLADIVNLCVDYYENKMYLTPSEKHMLLKVMGFGLYLMDGSVSNIYKLDAKKRINLTKIDKFFKQLQVVPLFGDMQIELARYIKTSAHYEENKSRWTCTSSSSSPQYNICEQMIQIREDHMRFISELARYSNSEVVTGSGRQEAQKTDAEYRKLFDLSLQGLQLLSQWSAHVMEVVCYLGSKTRKGGSLGRRDS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP31</accession>
    <name>A0A452IP31_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Peptidase S1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000033907.1">
        <property type="protein sequence ID" value="ENSGAGP00000029862.1"/>
        <property type="gene ID" value="ENSGAGG00000021548.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001867">
        <property type="term" value="P:complement activation, lectin pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00190">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.10:FF:000062">
        <property type="entry name" value="mannan-binding lectin serine protease 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.10:FF:000068">
        <property type="entry name" value="transmembrane protease serine 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.10.10">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009003">
        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043504">
        <property type="entry name" value="Peptidase_S1_PA_chymotrypsin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001314">
        <property type="entry name" value="Peptidase_S1A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001254">
        <property type="entry name" value="Trypsin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033116">
        <property type="entry name" value="TRYPSIN_SER"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24255">
        <property type="entry name" value="COMPLEMENT COMPONENT 1, S SUBCOMPONENT-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24255:SF13">
        <property type="entry name" value="MANNAN-BINDING LECTIN SERINE PROTEASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00089">
        <property type="entry name" value="Trypsin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00722">
        <property type="entry name" value="CHYMOTRYPSIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00020">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50494">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50240">
        <property type="entry name" value="TRYPSIN_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00135">
        <property type="entry name" value="TRYPSIN_SER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 4" id="KW-0732">Signal</keyword>
    <keyword evidence="1" id="KW-0768">Sushi</keyword>
    <feature type="signal peptide" evidence="4">
        <location>
            <begin position="1"/>
            <end position="26"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019120223" description="Peptidase S1 domain-containing protein" evidence="4">
        <location>
            <begin position="27"/>
            <end position="294"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase S1" evidence="5">
        <location>
            <begin position="39"/>
            <end position="280"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022659"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029862.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="294" mass="32667" checksum="2FBF508CB2300682" modified="2019-05-08" version="1">ISRTLQKTLLAPVRLTVLSFLSPVCGQPVRPLPGIIKRIIGGRNAELGFFPWQALIVVEDMSRVPNDKWFGSGALLSDSWVLTAAHVLRSQRRXAPSAGTTLSSRNKKDAVNRTVEEIILHVGFDIQNYNHDIALVKLKEKVTMDAYVMPVCLPQFEHELEGPHPNTLGLVAGWGISNPNITYVKLPVVLHDECKTSYESRSGNYSVTENMFCAGYYEGGKDTCLGDSGGAFVIQDPRTQRWVAQGLVSWGGPEECGSKQVYGVYTKVSNYVDWVEEKMGSSHKWTFLDPELNR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP32</accession>
    <name>A0A452IP32_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">RNA guanine-7 methyltransferase activating subunit</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the RAM family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IP32"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033616.1">
        <property type="protein sequence ID" value="ENSGAGP00000029604.1"/>
        <property type="gene ID" value="ENSGAGG00000021379.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000033617.1">
        <property type="protein sequence ID" value="ENSGAGP00000029605.1"/>
        <property type="gene ID" value="ENSGAGG00000021379.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031533">
        <property type="term" value="C:mRNA capping enzyme complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0106005">
        <property type="term" value="P:RNA 5'-cap (guanine-N7)-methylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028271">
        <property type="entry name" value="RAMAC"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48168">
        <property type="entry name" value="RNA GUANINE-7 METHYLTRANSFERASE-ACTIVATING SUBUNIT-LIKE (PSEUDOGENE)-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48168:SF1">
        <property type="entry name" value="RNA GUANINE-N7 METHYLTRANSFERASE ACTIVATING SUBUNIT-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15320">
        <property type="entry name" value="RAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0539">Nucleus</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="22"/>
            <end position="119"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="52"/>
            <end position="72"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="75"/>
            <end position="100"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00034716"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029604.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="7">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="119" mass="14495" checksum="5285291F009B4B53" modified="2019-05-08" version="1">MTSSTEVPLNYEEMFSHRFTADDEEYQEYLKRPTDPPPLVEEWKNRSGGNQRNRDWFQDGRQFRGRGDRHDWQGGNRSNQWPGRSWGNSYQQHRQGQSYYPQYDYGYNSYNPRPHYGRY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP34</accession>
    <name>A0A452IP34_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">HAUS augmin-like complex subunit 6 N-terminal domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IP34"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033660.1">
        <property type="protein sequence ID" value="ENSGAGP00000029645.1"/>
        <property type="gene ID" value="ENSGAGG00000021397.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070652">
        <property type="term" value="C:HAUS complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:1990498">
        <property type="term" value="C:mitotic spindle microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008017">
        <property type="term" value="F:microtubule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051225">
        <property type="term" value="P:spindle assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026797">
        <property type="entry name" value="HAUS_6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028163">
        <property type="entry name" value="HAUS_6_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16151">
        <property type="entry name" value="HAUS AUGMIN-LIKE COMPLEX SUBUNIT 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16151:SF2">
        <property type="entry name" value="HAUS AUGMIN-LIKE COMPLEX SUBUNIT 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14661">
        <property type="entry name" value="HAUS6_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="HAUS augmin-like complex subunit 6 N-terminal" evidence="2">
        <location>
            <begin position="1"/>
            <end position="196"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="164"/>
            <end position="191"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF14661"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029645.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="930" mass="106201" checksum="6AF655D068878E22" modified="2019-05-08" version="1">MFDKPNKDAFHVVARFLFTKLDHPRSIEIFRFCCPPADKKGDAEFRKQCCEWLKKISDECGNSFPPVVASLFLSPGGPKFIHLMYHFARYVIVQHMKKDSEDTNIHFPEAVNSRPHDSYKAAARYRVACIRFLQSIQREDFVIREYQRKAQLLTKQIRDPRAECAELHKLLQKMEQNSQKQTDQAERVQKVRSRWASVMETLTCLEKEKEVVDSVVDGHVDQYALDGTNVTISVPRLLLDKIENQMYKLHIGNVYEAGKLNLLTVIQLLNEALKILRYERQQVDHEALKLDLQYIEGKTKFQNKMLLGLKSMRHKLKREDHVSIVESIAEKQRDWDMKWENFLGQSPFSLIKDQNPVLDLLPAMSPLSFDPATEEAYKSSVFCKYPASVPDTPKKDCQANDKKEDAAFESIMDTTSTGRMLSHLQSARQSFLTAEPSSIVLEKDVHTTPPKRKEQSVSQILKYTVHKPRPIEAWKNGDTHVLQSPIAVKREDPIKKAQEQLAEEVADVVVSESTQGDGGKGMELDDLIGSLASNPFLTRKQIPRTPENLITEIRSSWRKAIQTEDPSDIELAQTEMVTKEAPLDLESAICSRRDSSMACFMSSSHMSDLNVSNLLERKSAVSWMTPAPKKQVVVSHVSESPTQIPATRMPDNERSWELGSKHTVSSKSLTENLEESIFPTVDKSMNPLDNSAERDIKTDALSPFFCQNSSVHTTLSWDASQMVSSTSSDSNEVIQFGILHETLPEELGNISLNSSESLETDDEIECNSRDNKSLENIVKNEWAAQECKQDLQSIRNRYEALKKTVFRNEEDSFQSPRKQFLRHRSERNLIPGCKETKEVFSPLGKLYALDAEYIKTPSHMSVFERRHTLSPLIAFSPVQERLRSMTQKKPGDFLHNLKEEENLKEKLDAKESSSDIKTRKDEAVSHLIEL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP40</accession>
    <name>A0A452IP40_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Protein Wnt</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Ligand for members of the frizzled family of seven transmembrane receptors.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 9">Secreted</location>
            <location evidence="1 9">Extracellular space</location>
            <location evidence="1 9">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 9">Belongs to the Wnt family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IP40"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033698.1">
        <property type="protein sequence ID" value="ENSGAGP00000029677.1"/>
        <property type="gene ID" value="ENSGAGG00000021418.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005125">
        <property type="term" value="F:cytokine activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005109">
        <property type="term" value="F:frizzled binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060070">
        <property type="term" value="P:canonical Wnt signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045165">
        <property type="term" value="P:cell fate commitment"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030182">
        <property type="term" value="P:neuron differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd19354">
        <property type="entry name" value="Wnt_Wnt9b"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.2460.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005817">
        <property type="entry name" value="Wnt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043158">
        <property type="entry name" value="Wnt_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018161">
        <property type="entry name" value="Wnt_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12027:SF84">
        <property type="entry name" value="PROTEIN WNT-9B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12027">
        <property type="entry name" value="WNT RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00110">
        <property type="entry name" value="wnt"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01349">
        <property type="entry name" value="WNTPROTEIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00097">
        <property type="entry name" value="WNT1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00246">
        <property type="entry name" value="WNT1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 9" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="7" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="8" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="6 9" id="KW-0879">Wnt signaling pathway</keyword>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="251"/>
            <end position="270"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="252"/>
            <end position="270"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005683"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022687"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="RuleBase" id="RU003500"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029677.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="349" mass="38915" checksum="F8A10A7A6F079DC9" modified="2025-10-08" version="2">MALLLRAVGNVMPCVMCLTGKEGLTHLPSLGPSANPAQGKAHLKQCDLLKLSRKQKRLCQREPGLAETLRDAIRLSIMECQYQFRYERWNCSLDGRTSLLKRGFKETAFLYAVSSAALTHSLARACSAGRMERCTCDDSPDLENRKAWQWGVCGDNLKYSTKFLKNFLGQKKIGKDLRAKVDIHNTNVGIKAVKNGLKTTCKCHGVSGSCAVRTCWKQLSPFHETGKILKLRFDDAVKIFSATNDAVGHSELVSTQRRSHSPKSPATPRSTDLVYVEDSPSFCRPSKYSLGTAGRTCSREASCDSLCCGRGYNTQSRMVTFSCHCQVQWCCYVECQQCMQEEVIYSCKQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP44</accession>
    <name>A0A452IP44_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Scavenger receptor class A member 5</fullName>
        </recommendedName>
    </protein>
    <gene>
        <name evidence="9" type="primary">SCARA5</name>
    </gene>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Ferritin receptor that mediates non-transferrin-dependent delivery of iron. Mediates cellular uptake of ferritin-bound iron by stimulating ferritin endocytosis from the cell surface with consequent iron delivery within the cell. Delivery of iron to cells by ferritin is required for the development of specific cell types, suggesting the existence of cell type-specific mechanisms of iron traffic in organogenesis, which alternatively utilize transferrin or non-transferrin iron delivery pathways.</text>
    </comment>
    <comment type="subunit">
        <text evidence="9">Homotrimer.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="9">Cell membrane</location>
            <topology evidence="9">Single-pass type II membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type II membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="9">Belongs to the SCARA5 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IP44"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033776.1">
        <property type="protein sequence ID" value="ENSGAGP00000029747.1"/>
        <property type="gene ID" value="ENSGAGG00000021461.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070287">
        <property type="term" value="F:ferritin receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006897">
        <property type="term" value="P:endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006879">
        <property type="term" value="P:intracellular iron ion homeostasis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034755">
        <property type="term" value="P:iron ion transmembrane transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031638">
        <property type="term" value="P:zymogen activation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000011">
        <property type="entry name" value="Scavenger receptor class A member 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.250.10">
        <property type="entry name" value="SRCR-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="HAMAP" id="MF_03070">
        <property type="entry name" value="SCARA5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008160">
        <property type="entry name" value="Collagen"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034726">
        <property type="entry name" value="SCARA5"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001190">
        <property type="entry name" value="SRCR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036772">
        <property type="entry name" value="SRCR-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48071:SF24">
        <property type="entry name" value="DELETED IN MALIGNANT BRAIN TUMORS 1 PROTEIN-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48071">
        <property type="entry name" value="SRCR DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01391">
        <property type="entry name" value="Collagen"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00530">
        <property type="entry name" value="SRCR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00258">
        <property type="entry name" value="SPERACTRCPTR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00202">
        <property type="entry name" value="SR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56487">
        <property type="entry name" value="SRCR-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00420">
        <property type="entry name" value="SRCR_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50287">
        <property type="entry name" value="SRCR_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="6 10" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="8" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9" id="KW-0406">Ion transport</keyword>
    <keyword evidence="9" id="KW-0408">Iron</keyword>
    <keyword evidence="9" id="KW-0410">Iron transport</keyword>
    <keyword evidence="5 9 12" id="KW-0472">Membrane</keyword>
    <keyword evidence="7 9" id="KW-0675">Receptor</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 9" id="KW-0735">Signal-anchor</keyword>
    <keyword evidence="2 9 12" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 9 12" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="9" id="KW-0813">Transport</keyword>
    <feature type="topological domain" description="Cytoplasmic" evidence="9">
        <location>
            <begin position="1"/>
            <end position="58"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="58"/>
            <end position="80"/>
        </location>
    </feature>
    <feature type="topological domain" description="Extracellular" evidence="9">
        <location>
            <begin position="80"/>
            <end position="485"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="13">
        <location>
            <begin position="383"/>
            <end position="483"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="301"/>
            <end position="373"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="243"/>
            <end position="288"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="320"/>
            <end position="344"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="362"/>
            <end position="373"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="10">
        <location>
            <begin position="408"/>
            <end position="472"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="10">
        <location>
            <begin position="421"/>
            <end position="482"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="10">
        <location>
            <begin position="452"/>
            <end position="462"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004606"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="HAMAP-Rule" id="MF_03070"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00196"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50287"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029747.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="485" mass="53583" checksum="1E9A53879F2847FF" modified="2019-05-08" version="1">MENKAMYLHTFSERENGSIFEESFDGRNLSKLNLCEDGKRAALIWRCGWLGSITALKYAVVGLYLLVFLILVGFFILAVSRPQSSPEDLKALTGNVNRLNESIRDMQLKLLQLPLKGDILDHIWRLQDLLQNHSDSLFLMTGSLQKLEGFLWSLQTQASQTDKAVASLRDSLHHQSDMAQLETYRLSVEGNSSRLLLEHHNSLLTHLAAQVESLSDQLTEVGGAVGTINRTFSYDVGIHHTRIQDLQVLISNATEDARQMRRIHIAMEQQLKQELAILNNVTEDLRLKDWEHSIALRNISVIRGPPGPKGDQGNEGMEGEPGVPGLPGLRGLPGERGPLGPPGLKGDKGDLGPSGPAGMRGFKGDRGQKGEKGDRAAAEEGLIRLVNGSVPHEGRVEVFHDRHWGTVCDDSWDKKDGDVACRMLGFRGAEEVYRMARFGQGIGRIWMDDVACKGTEDSLFQCTFSSWGKSNCGHAEDAGVKCLTQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP45</accession>
    <name>A0A452IP45_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">thioredoxin-disulfide reductase (NADPH)</fullName>
            <ecNumber evidence="2">1.8.1.9</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="11">
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </cofactor>
        <text evidence="11">Binds 1 FAD per subunit.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1 13">Belongs to the class-I pyridine nucleotide-disulfide oxidoreductase family.</text>
    </comment>
    <dbReference type="EC" id="1.8.1.9" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452IP45"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033703.1">
        <property type="protein sequence ID" value="ENSGAGP00000029682.1"/>
        <property type="gene ID" value="ENSGAGG00000021420.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050660">
        <property type="term" value="F:flavin adenine dinucleotide binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004362">
        <property type="term" value="F:glutathione-disulfide reductase (NADPH) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004791">
        <property type="term" value="F:thioredoxin-disulfide reductase (NADPH) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045454">
        <property type="term" value="P:cell redox homeostasis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034599">
        <property type="term" value="P:cellular response to oxidative stress"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006749">
        <property type="term" value="P:glutathione metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.50.50.60:FF:000190">
        <property type="entry name" value="Thioredoxin reductase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.390.30:FF:000004">
        <property type="entry name" value="Thioredoxin reductase 1, cytoplasmic"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.390.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.50.50.60">
        <property type="entry name" value="FAD/NAD(P)-binding domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036188">
        <property type="entry name" value="FAD/NAD-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023753">
        <property type="entry name" value="FAD/NAD-binding_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016156">
        <property type="entry name" value="FAD/NAD-linked_Rdtase_dimer_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046952">
        <property type="entry name" value="GSHR/TRXR-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001100">
        <property type="entry name" value="Pyr_nuc-diS_OxRdtase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004099">
        <property type="entry name" value="Pyr_nucl-diS_OxRdtase_dimer"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012999">
        <property type="entry name" value="Pyr_OxRdtase_I_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006338">
        <property type="entry name" value="Thioredoxin/glutathione_Rdtase"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01438">
        <property type="entry name" value="TGR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR42737">
        <property type="entry name" value="GLUTATHIONE REDUCTASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR42737:SF7">
        <property type="entry name" value="THIOREDOXIN-DISULFIDE REDUCTASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07992">
        <property type="entry name" value="Pyr_redox_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02852">
        <property type="entry name" value="Pyr_redox_dim"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000350">
        <property type="entry name" value="Mercury_reductase_MerA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00368">
        <property type="entry name" value="FADPNR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00411">
        <property type="entry name" value="PNDRDTASEI"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51905">
        <property type="entry name" value="FAD/NAD(P)-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55424">
        <property type="entry name" value="FAD/NAD-linked reductases, dimerisation (C-terminal) domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00076">
        <property type="entry name" value="PYRIDINE_REDOX_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="4 11 13" id="KW-0274">FAD</keyword>
    <keyword evidence="3 13" id="KW-0285">Flavoprotein</keyword>
    <keyword evidence="11" id="KW-0520">NAD</keyword>
    <keyword evidence="5" id="KW-0521">NADP</keyword>
    <keyword evidence="11" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="7 13" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="9 13" id="KW-0676">Redox-active center</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0712">Selenocysteine</keyword>
    <feature type="domain" description="FAD/NAD(P)-binding" evidence="15">
        <location>
            <begin position="7"/>
            <end position="328"/>
        </location>
    </feature>
    <feature type="domain" description="Pyridine nucleotide-disulphide oxidoreductase dimerisation" evidence="14">
        <location>
            <begin position="348"/>
            <end position="459"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="10">
        <location>
            <position position="450"/>
        </location>
    </feature>
    <feature type="binding site" evidence="11">
        <location>
            <position position="48"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="11">
        <location>
            <position position="112"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="11">
        <location>
            <begin position="178"/>
            <end position="185"/>
        </location>
        <ligand>
            <name>NAD(+)</name>
            <dbReference type="ChEBI" id="CHEBI:57540"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="11">
        <location>
            <position position="270"/>
        </location>
        <ligand>
            <name>NAD(+)</name>
            <dbReference type="ChEBI" id="CHEBI:57540"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="11">
        <location>
            <position position="312"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="disulfide bond" description="Redox-active" evidence="12">
        <location>
            <begin position="39"/>
            <end position="44"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007532"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00012610"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022630"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022827"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022857"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022933"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023284"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PIRSR" id="PIRSR000350-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PIRSR" id="PIRSR000350-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PIRSR" id="PIRSR000350-4"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="RuleBase" id="RU003691"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="Pfam" id="PF02852"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="Pfam" id="PF07992"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029682.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="475" mass="52077" checksum="4CCF374687F88AF9" modified="2019-05-08" version="1">MIALLVFCFEAAQFGKKVAVLDYVEPSPKGTKWGLGGTCVNVGCIPKKLMHQASLLGSALKDAQHYGWNISHPVRHDWPVMAEAIQNYVKSLNWGHRVQLQDKKVKYFNLTGSLMDAHTVRGLTNAGKETIITAENIVIATGGRPKYPAHIAGALEYGITSDDIFWLKESPGKTLVVGASYVSLECAGFLTGIGQDTTVMVRSIPLRGFDQQMSFLVTDYMESCGTRFLKKCTPSKVEKLKSGKLQVTWKHADSGKEEMDDFDTVMWAVGRAPDTKTLNLETIGVKISSETGKIIVDASEATSVPHIYAIGDITEGRPELTPTAIAAGKLLAQRLFGQSSELMDYDNVPTTVFTPLEYGCVGLSEEKAVEHYGLDNIEVYHAYYKPLEFTVAERDATQCYIKMVCLRKKEQRILGLHFIGPNAGEVIQGFALGMKCGATYAQMMKTVGIHPTCAEEVTKLRITKRSGLDAKVTGC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP48</accession>
    <name>A0A452IP48_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="6">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>pyridoxal 5'-phosphate</name>
            <dbReference type="ChEBI" id="CHEBI:597326"/>
        </cofactor>
    </comment>
    <comment type="similarity">
        <text evidence="2 5">Belongs to the class-III pyridoxal-phosphate-dependent aminotransferase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IP48"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033676.1">
        <property type="protein sequence ID" value="ENSGAGP00000029660.1"/>
        <property type="gene ID" value="ENSGAGG00000021408.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016829">
        <property type="term" value="F:lyase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030170">
        <property type="term" value="F:pyridoxal phosphate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008483">
        <property type="term" value="F:transaminase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00610">
        <property type="entry name" value="OAT_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.640.10:FF:000058">
        <property type="entry name" value="ethanolamine-phosphate phospho-lyase isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1150.10">
        <property type="entry name" value="Aspartate Aminotransferase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.640.10">
        <property type="entry name" value="Type I PLP-dependent aspartate aminotransferase-like (Major domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005814">
        <property type="entry name" value="Aminotrans_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049704">
        <property type="entry name" value="Aminotrans_3_PPA_site"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015424">
        <property type="entry name" value="PyrdxlP-dep_Trfase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015421">
        <property type="entry name" value="PyrdxlP-dep_Trfase_major"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015422">
        <property type="entry name" value="PyrdxlP-dep_Trfase_small"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45688">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45688:SF6">
        <property type="entry name" value="5-PHOSPHOHYDROXY-L-LYSINE PHOSPHO-LYASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00202">
        <property type="entry name" value="Aminotran_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000521">
        <property type="entry name" value="Transaminase_4ab_Lys_Orn"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53383">
        <property type="entry name" value="PLP-dependent transferases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00600">
        <property type="entry name" value="AA_TRANSFER_CLASS_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0456">Lyase</keyword>
    <keyword evidence="3 5" id="KW-0663">Pyridoxal phosphate</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001933"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008954"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022898"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="RuleBase" id="RU003560"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029660.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="445" mass="50113" checksum="01F8D88B14AE210F" modified="2019-05-08" version="1">MQARPETLALRRQLIGSSCKLFFPEEPVKIVRAKGQYIYDEHGREYLDCINNVSHVGHCHPDVVKAAHEQNQLLNTNSRYLHDNLVDYAKRLSKTLPEKLCTFYFLNSGSEANDLALRLARQYTKHKDVIVLDHAYHGHLTSLIDISPYKFRNLEGQKEWVHVAPLPDTYRGIYREDYKDSVLAYANEVKKIIEQAQKRGRKIAAFFIESLPSVGGQIIPPEGYFQMIAEHIHKAGGVFVADEIQVGFGRVGKCFWAFQLQGEDFVPDIVTMGKPMGNGHPIACVATTKEIAEAFADTGVEYFNTFGGNPVSCAIGLAVLDVIEKEHLQAHATQVGNFLMELLNQQKVKHPIIGDVRGVGLFIGVDLIKDQEKRTPATKEAEYLITRLKREYILLSTDGPGKNVLKFKPPLCFTMENAKSVVDKMDEILTDMEREKTCEPVGNCH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP49</accession>
    <name>A0A452IP49_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Transcriptional regulator</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="3">Transcription factor that binds DNA in a non-specific manner, yet also specifically recognizes the core sequence 5'-CAC[GA]TG-3'. Activates the transcription of growth-related genes.</text>
    </comment>
    <comment type="subunit">
        <text evidence="3">Efficient DNA binding requires dimerization with another bHLH protein.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IP49"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033781.1">
        <property type="protein sequence ID" value="ENSGAGP00000029752.1"/>
        <property type="gene ID" value="ENSGAGG00000021479.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003700">
        <property type="term" value="F:DNA-binding transcription factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046983">
        <property type="term" value="F:protein dimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd11458">
        <property type="entry name" value="bHLHzip_c-Myc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.280.10:FF:000019">
        <property type="entry name" value="Myc proto-oncogene protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.280.10">
        <property type="entry name" value="Helix-loop-helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011598">
        <property type="entry name" value="bHLH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036638">
        <property type="entry name" value="HLH_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003327">
        <property type="entry name" value="Myc-LZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050433">
        <property type="entry name" value="Myc_transcription_factors"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002418">
        <property type="entry name" value="Tscrpt_reg_Myc"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012682">
        <property type="entry name" value="Tscrpt_reg_Myc_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45851">
        <property type="entry name" value="MYC PROTO-ONCOGENE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00010">
        <property type="entry name" value="HLH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02344">
        <property type="entry name" value="Myc-LZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01056">
        <property type="entry name" value="Myc_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF001705">
        <property type="entry name" value="Myc_protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00044">
        <property type="entry name" value="LEUZIPPRMYC"/>
    </dbReference>
    <dbReference type="SMART" id="SM00353">
        <property type="entry name" value="HLH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47459">
        <property type="entry name" value="HLH, helix-loop-helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50888">
        <property type="entry name" value="BHLH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0010">Activator</keyword>
    <keyword evidence="4" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="1 3" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0539">Nucleus</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="BHLH" evidence="6">
        <location>
            <begin position="323"/>
            <end position="375"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="144"/>
            <end position="172"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="198"/>
            <end position="263"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="308"/>
            <end position="328"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="4">
        <location>
            <begin position="379"/>
            <end position="406"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="5">
        <location>
            <begin position="154"/>
            <end position="166"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="5">
        <location>
            <begin position="198"/>
            <end position="225"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="5">
        <location>
            <begin position="226"/>
            <end position="239"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PIRNR" id="PIRNR001705"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50888"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029752.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="408" mass="45681" checksum="099610BCE1D18AA6" modified="2019-05-08" version="1">MPLTSTLPSKNYDYDYDSVQPYFYFEEEEENFYLAAQQRGSELQPPAPSEDIWKKFELLPTPPLSPSRRSSLAAAYFPSNADQLEMVTELLGGDMVNQSFICDPDDESFVKSIIIQDCMWSGFSAAAKLEKVVSEKLASYQAARREGGSGSRLCPPPPPSQPPSLAPSPASASTYLHDLGAAASDCIDPSVVFPYPLSERSSKPGSPSSSPASLLGDDTPPTTSSDSEEEPEEDEEEIDVVTTSESESITQPTEEHSKPHHSPLVLKRCHVPIHQHNYAAPPSTKLEYPSAKRIKLDNGRVLKQISNNRKCSSPRTSDTEENDKRRTHNVLERQRRNELKLSFFALRDQIPEVANNEKAPKVVILKKATEYVLSIQTDEHRLITEKEQLRKRREQLKHKLEQLRNSCA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0M5</accession>
    <name>A0A452J0M5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3 15">Coatomer subunit beta'</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">The coatomer is a cytosolic protein complex that binds to dilysine motifs and reversibly associates with Golgi non-clathrin-coated vesicles, which further mediate biosynthetic protein transport from the ER, via the Golgi up to the trans Golgi network. Coatomer complex is required for budding from Golgi membranes, and is essential for the retrograde Golgi-to-ER transport of dilysine-tagged proteins. In mammals, the coatomer can only be recruited by membranes associated to ADP-ribosylation factors (ARFs), which are small GTP-binding proteins; the complex also influences the Golgi structural integrity, as well as the processing, activity, and endocytic recycling of LDL receptors.</text>
    </comment>
    <comment type="function">
        <text evidence="14">This coatomer complex protein, essential for Golgi budding and vesicular trafficking, is a selective binding protein (RACK) for protein kinase C, epsilon type. It binds to Golgi membranes in a GTP-dependent manner.</text>
    </comment>
    <comment type="subunit">
        <text evidence="15">Oligomeric complex that consists of at least the alpha, beta, beta', gamma, delta, epsilon and zeta subunits.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 15">Cytoplasmic vesicle</location>
            <location evidence="1 15">COPI-coated vesicle membrane</location>
            <topology evidence="1 15">Peripheral membrane protein</topology>
            <orientation evidence="1 15">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="15">Golgi apparatus membrane</location>
            <topology evidence="15">Peripheral membrane protein</topology>
            <orientation evidence="15">Cytoplasmic side</orientation>
        </subcellularLocation>
        <text evidence="15">The coatomer is cytoplasmic or polymerized on the cytoplasmic side of the Golgi, as well as on the vesicles/buds originating from it.</text>
    </comment>
    <comment type="similarity">
        <text evidence="2 15">Belongs to the WD repeat COPB2 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0M5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038447.1">
        <property type="protein sequence ID" value="ENSGAGP00000033950.1"/>
        <property type="gene ID" value="ENSGAGG00000024137.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030126">
        <property type="term" value="C:COPI vesicle coat"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000139">
        <property type="term" value="C:Golgi membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005198">
        <property type="term" value="F:structural molecule activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006888">
        <property type="term" value="P:endoplasmic reticulum to Golgi vesicle-mediated transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006891">
        <property type="term" value="P:intra-Golgi vesicle-mediated transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006886">
        <property type="term" value="P:intracellular protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006890">
        <property type="term" value="P:retrograde vesicle-mediated transport, Golgi to endoplasmic reticulum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd22947">
        <property type="entry name" value="Coatomer_WDAD_beta-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00200">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.470:FF:000001">
        <property type="entry name" value="Coatomer subunit beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:000008">
        <property type="entry name" value="Coatomer subunit beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.470">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006692">
        <property type="entry name" value="Beta-prop_COPA/B_2nd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050844">
        <property type="entry name" value="Coatomer_complex_subunit"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016453">
        <property type="entry name" value="COPB2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056176">
        <property type="entry name" value="TPR_COPA_B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020472">
        <property type="entry name" value="WD40_PAC1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036322">
        <property type="entry name" value="WD40_repeat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001680">
        <property type="entry name" value="WD40_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19876">
        <property type="entry name" value="COATOMER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19876:SF2">
        <property type="entry name" value="COATOMER SUBUNIT BETA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04053">
        <property type="entry name" value="B-prop_COPA_B_2nd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23953">
        <property type="entry name" value="TPR_COPA_B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00400">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF005567">
        <property type="entry name" value="Coatomer_beta'_subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00320">
        <property type="entry name" value="GPROTEINBRPT"/>
    </dbReference>
    <dbReference type="SMART" id="SM00320">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50978">
        <property type="entry name" value="WD40 repeat-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50082">
        <property type="entry name" value="WD_REPEATS_2"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50294">
        <property type="entry name" value="WD_REPEATS_REGION"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 15" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="12 15" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="8 15" id="KW-0931">ER-Golgi transport</keyword>
    <keyword evidence="10 15" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="11 15" id="KW-0472">Membrane</keyword>
    <keyword evidence="9 15" id="KW-0653">Protein transport</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="4 15" id="KW-0813">Transport</keyword>
    <keyword evidence="6 16" id="KW-0853">WD repeat</keyword>
    <feature type="repeat" description="WD" evidence="16">
        <location>
            <begin position="11"/>
            <end position="52"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="16">
        <location>
            <begin position="95"/>
            <end position="127"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="16">
        <location>
            <begin position="138"/>
            <end position="180"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="16">
        <location>
            <begin position="181"/>
            <end position="224"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="16">
        <location>
            <begin position="225"/>
            <end position="266"/>
        </location>
    </feature>
    <feature type="domain" description="COPA/B second beta-propeller" evidence="17">
        <location>
            <begin position="319"/>
            <end position="577"/>
        </location>
    </feature>
    <feature type="domain" description="COPA/B TPR" evidence="18">
        <location>
            <begin position="594"/>
            <end position="774"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004347"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010844"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00021083"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022574"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022892"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00024791"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00025693"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PIRNR" id="PIRNR005567"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF04053"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF23953"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033950.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="882" mass="99660" checksum="6E11A178C3527DC2" modified="2019-05-08" version="1">MPLRLDIKRKLTARSDRVKSVDLHPTEPWMLASLYNGSVCVWNHETQTLVKTFEVCDLPVRAAKFVARKNWVVTGADDMQIRVFNYNTLERVHLFEAHSDYIRCIAVHPTQPFILTSSDDMLIKLWDWDKKWSCSQVFEGHTHYVMQIVINPKDNNQFASASLDRTIKVWQLGSSSPNFTLEGHEKGVNCIDYYSGGDKPYLISGADDRLVKIWDYQNKTCVQTLEGHAQNVSCVSFHPELPIIITGSEDGTVRIWHSSTYRLESTLNYGMERVWCVASLRGSNNVALGYDEGSIIVKLGREEPAMSMDSNGKIIWAKHSEVQQANLKAMGDAEIKDGERLPLAVKDMGSCEIYPQTIQHNPNGRFVVVCGDGEYIIYTAMALRNKSFGSAQEFVWAHDSSEYAIRESNSVVKIFKNFKEKKSFKPDFGAEGIYGGFLLGVRSVNGLAFYDWDNTELIRRIEIQPKHIFWSDSGELVCIATEESFFILKYLSEKVAAAQETHEGVTEDGIEDAFEVLGEIQEIVKTGLWVGDCFIYTSSVNRLNYYVGGEIVTIAHLDRTMYLLGYIPKDNRLYLGDKELNIVSYSLLVSVLEYQTAVMRRDFGMADKVLPTIPKEQRTRVAHFLEKQGFKQQALAVSTDPEHRFELALQLGELKIAYQLAVEAESEQKWKQLAELAISKCQFGLAQECLHHAQDYGGLLLLATASGNANMVNKLAEGAEKDGKNNVAFMSYFLQGKVDCCLELLIKTGRLPEAAFLARTYLPSQVSRVVKMWRENLSKVNQKAAESLADPTEYENLFPGLKEAFVVEDYVKETHTDLRPAKEYPLVTPNEERNLLEEAQGFKPSGVVVSQALLDLDDDLDNLDLEDIDTTDINLDEEILDD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0M8</accession>
    <name>A0A452J0M8_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="20">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 16">Membrane</location>
            <topology evidence="1 16">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 16">Belongs to the integrin alpha chain family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0M8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038401.1">
        <property type="protein sequence ID" value="ENSGAGP00000033907.1"/>
        <property type="gene ID" value="ENSGAGG00000024116.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000038406.1">
        <property type="protein sequence ID" value="ENSGAGP00000033912.1"/>
        <property type="gene ID" value="ENSGAGG00000024116.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009897">
        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008305">
        <property type="term" value="C:integrin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033627">
        <property type="term" value="P:cell adhesion mediated by integrin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098609">
        <property type="term" value="P:cell-cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007160">
        <property type="term" value="P:cell-matrix adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007229">
        <property type="term" value="P:integrin-mediated signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1460:FF:000007">
        <property type="entry name" value="Integrin subunit alpha 9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.130:FF:000006">
        <property type="entry name" value="Integrin, alpha 9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1510:FF:000013">
        <property type="entry name" value="Integrin, alpha 9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1530:FF:000005">
        <property type="entry name" value="Integrin, alpha 9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.130">
        <property type="entry name" value="Integrin alpha, N-terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1460">
        <property type="entry name" value="Integrin domains. Chain A, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1510">
        <property type="entry name" value="ntegrin, alpha v. Chain A, domain 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1530">
        <property type="entry name" value="ntegrin, alpha v. Chain A, domain 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013517">
        <property type="entry name" value="FG-GAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013519">
        <property type="entry name" value="Int_alpha_beta-p"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000413">
        <property type="entry name" value="Integrin_alpha"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013649">
        <property type="entry name" value="Integrin_alpha_Ig-like_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048285">
        <property type="entry name" value="Integrin_alpha_Ig-like_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048286">
        <property type="entry name" value="Integrin_alpha_Ig-like_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028994">
        <property type="entry name" value="Integrin_alpha_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032695">
        <property type="entry name" value="Integrin_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23220">
        <property type="entry name" value="INTEGRIN ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23220:SF69">
        <property type="entry name" value="INTEGRIN ALPHA-9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01839">
        <property type="entry name" value="FG-GAP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08441">
        <property type="entry name" value="Integrin_A_Ig_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20805">
        <property type="entry name" value="Integrin_A_Ig_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20806">
        <property type="entry name" value="Integrin_A_Ig_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01185">
        <property type="entry name" value="INTEGRINA"/>
    </dbReference>
    <dbReference type="SMART" id="SM00191">
        <property type="entry name" value="Int_alpha"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69318">
        <property type="entry name" value="Integrin alpha N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69179">
        <property type="entry name" value="Integrin domains"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51470">
        <property type="entry name" value="FG_GAP"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0106">Calcium</keyword>
    <keyword evidence="8 16" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="14" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10 16" id="KW-0401">Integrin</keyword>
    <keyword evidence="11" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13 16" id="KW-0675">Receptor</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5 16" id="KW-0732">Signal</keyword>
    <keyword evidence="3" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="16">
        <location>
            <begin position="1"/>
            <end position="27"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5044516220" evidence="16">
        <location>
            <begin position="28"/>
            <end position="993"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="29"/>
            <end position="94"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="288"/>
            <end position="347"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="349"/>
            <end position="406"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="409"/>
            <end position="472"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha first immunoglubulin-like" evidence="17">
        <location>
            <begin position="457"/>
            <end position="616"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha second immunoglobulin-like" evidence="18">
        <location>
            <begin position="618"/>
            <end position="762"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha third immunoglobulin-like" evidence="19">
        <location>
            <begin position="770"/>
            <end position="951"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023037"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00803"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="RuleBase" id="RU003762"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF08441"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF20805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="Pfam" id="PF20806"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033912.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="993" mass="110740" checksum="E156131DEA71227D" modified="2019-05-08" version="1">MPGPQWKVKRSWFKVLVGLIAVALAYSYNVDLDHPVVYQGPNGSFFGYSVLEHYYDNTRWVLVGAPKADSKYSASVKSPGAVFKCRVHTNPERRCTELDMGRGNTRGTLCGKTCKEDRDDEWMGVSLARQPRADGNVLACAHRWKNIYYETEYVLPHGFCNIIPPNLHSNGRKLLPCYEEYKKKYGEEHGSCQAGIAGFFTEELVIMGAPGSFYWTGTVKVLNLTDNTYFKLNDDAVIARRYTYLGYAVTAGHFSQPTTTDVVGGAPQDGGIGKVYIFRTDRGSGSLIKIFQASGKKMGSYFGSSLCAVDLNSDGLSDLLVGAPMFSEIRDEGQVTVYINRGNGVLEEQLVLNGDNAYNAHFGESMASLGDIDDDGYSDVAIGSPKEDDYVGAVYIYHGDAGGIVPQYSMKLSGQTINPMLRMFGQSISGGVDMDGNGYPDVTIGAFMSDNVVLLRSRPVITMDISIFLPASINITAPQCQDGLQQVNCLNVTVCFLFNGKRVPGEIGLNYILSADVAKKEKGQQPRVYFVSSGETVGQIVEQLQLSYMREKCDHYLAYVKRRVKDVISPIVFEAAYSLGEHVMGKEEKELPALKPILRWKKGEKIAQRNQTIFERNCQSDDCAADLKLQGRLLLSSVDDKTPYLALGAVKNISLNISISNIGDDAYDTNVFFNFSRELFFIKMWQKEELGIACELLESDFLKCSVGFPFMRSKSKYEFSVIFDTSHLSGEEEFLTFLVTAQSGNLEHSESLHDNTLTLSVPLMHEVDSSITGTVSPTSFFYGGSVEASNFVQLENYECLFQPLNFTLQVYNTGPSTLPESFVDILFPNRLSASGAETFHIQQMMVAQEKGNCFFQKNPNPCVIPQENDNIFHTIFAFFTKSGRKVLDCERPGRSCLIVRCNLSSLAKAESRSIDVYMLLNTEILKKDSSSVIQFVTRAKVRVDTTLRIVEVSNGNPEDATMGFFRRRYKEIIEAEKNRKENEDSWDWVQKNQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0M9</accession>
    <name>A0A452J0M9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Tektin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">Sperm-specific microtubule inner protein (MIP) part of the dynein-decorated doublet microtubules (DMTs) in flagellar axoneme. Forms an extensive interaction network in different conformations that reinforces the helix bundle composed by other tektin proteins (TEKT1 to TEKT4) and MIPs to anchor the tektin bundle onto the tubulin wall of A-tubule of the sperm flagellum.</text>
    </comment>
    <comment type="subunit">
        <text evidence="11">Microtubule inner protein component of sperm flagellar doublet microtubules. Interacts with TEKT3.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="12">Cytoplasm</location>
            <location evidence="12">Cytoskeleton</location>
            <location evidence="12">Cilium axoneme</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Flagellum axoneme</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 12">Belongs to the tektin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0M9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038354.1">
        <property type="protein sequence ID" value="ENSGAGP00000033863.1"/>
        <property type="gene ID" value="ENSGAGG00000024073.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005930">
        <property type="term" value="C:axoneme"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015630">
        <property type="term" value="C:microtubule cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036126">
        <property type="term" value="C:sperm flagellum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060271">
        <property type="term" value="P:cilium assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060294">
        <property type="term" value="P:cilium movement involved in cell motility"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048256">
        <property type="entry name" value="Tektin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000435">
        <property type="entry name" value="Tektins"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19960">
        <property type="entry name" value="TEKTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19960:SF23">
        <property type="entry name" value="TEKTIN-5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03148">
        <property type="entry name" value="Tektin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00511">
        <property type="entry name" value="TEKTIN"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9 12" id="KW-0966">Cell projection</keyword>
    <keyword evidence="7 12" id="KW-0969">Cilium</keyword>
    <keyword evidence="6" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="5 12" id="KW-0282">Flagellum</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0832">Ubl conjugation</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004611"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007209"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022846"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023069"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00046061"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00047089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="RuleBase" id="RU367040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033863.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="467" mass="53869" checksum="79AFD960CC7CE7CE" modified="2019-05-08" version="1">MEFLGTTQTASYCGPKKSYLFPEPVSTSTTQDSYQSYYLPGYRYLNSWRPSLFYRVSAVPTNSGDQGSMRISAPLRPPTILPSVRSALYARYSPHDWYQANTVQYKGSESCRHWAGRMNADSLRLMQDKNQLTQQTQDSTGRSLGQRLSDIDFWRSELTYELERLLTETRALETAKRRLECAADEAQRPLKVGQRYLILNVLVCLRVRLYYLTDKGHSQLPSGINRDAQHALERDLSDKNSAHFIDEKCYNLRNTSDSISFYHGVEKIDGTISVPETWAKFSDDNIRHSQNARANSIKLCEETENTLENVSEEMWKQFTDTNLAFTNRIAETADTKNKLQTHLAKTLQEIFQTENTIMLLERAIMAKECPLKVAQTRLEGRTRRPNVELCRDIPQFKLVNEVFTIDDTLQTLKLRLREAHDTLQLLVLNKARLEHDLSVKANSLFIDRKCMDMRKTFPSTPRLVGYT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0P1</accession>
    <name>A0A452J0P1_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="4">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0P1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038463.1">
        <property type="protein sequence ID" value="ENSGAGP00000033965.1"/>
        <property type="gene ID" value="ENSGAGG00000024148.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030515">
        <property type="term" value="F:snoRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.79.10">
        <property type="entry name" value="Nucleoside Triphosphate Pyrophosphohydrolase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR054754">
        <property type="entry name" value="NudT16"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31699">
        <property type="entry name" value="NUDIX T16 FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31699:SF6">
        <property type="entry name" value="TUDOR-INTERACTING REPAIR REGULATOR PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22327">
        <property type="entry name" value="Nudt16-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0539">Nucleus</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0694">RNA-binding</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033965.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="197" mass="20442" checksum="185467705000F9D2" modified="2019-05-08" version="1">MAALTGSAGAAGLLPSLSVPGVTELKPLSRYEAMRLGPGWSHSCHAMLYAPNPGMLFGRIPLRYAVLVMGMVRVPLYTQKDRMGGLPNFLGNSFIGTAKFQLLFALKVLNMVPEEKLAEAVAATQKQKKPAAEPAAVTANVTTANVTTTTAAVTAAVTANQIVAKPANELANQPLAVSMAEPAAEPTAEPAAEAMAE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="35" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0P3</accession>
    <name>A0A452J0P3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Vascular endothelial growth factor receptor 3</fullName>
            <ecNumber evidence="4">2.7.10.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="35 36">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="36" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="35" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="25">
            <text>L-tyrosyl-[protein] + ATP = O-phospho-L-tyrosyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10596"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.10.1"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="30">Membrane</location>
            <topology evidence="30">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="30">Belongs to the protein kinase superfamily. Tyr protein kinase family. CSF-1/PDGF receptor subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.10.1" evidence="4"/>
    <dbReference type="AlphaFoldDB" id="A0A452J0P3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038422.1">
        <property type="protein sequence ID" value="ENSGAGP00000033927.1"/>
        <property type="gene ID" value="ENSGAGG00000024090.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000038430.1">
        <property type="protein sequence ID" value="ENSGAGP00000033934.1"/>
        <property type="gene ID" value="ENSGAGG00000024090.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043235">
        <property type="term" value="C:receptor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019838">
        <property type="term" value="F:growth factor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004714">
        <property type="term" value="F:transmembrane receptor protein tyrosine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001525">
        <property type="term" value="P:angiogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030335">
        <property type="term" value="P:positive regulation of cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043408">
        <property type="term" value="P:regulation of MAPK cascade"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048010">
        <property type="term" value="P:vascular endothelial growth factor receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05863">
        <property type="entry name" value="IgI_VEGFR-3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05102">
        <property type="entry name" value="PTKc_VEGFR3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000077">
        <property type="entry name" value="Vascular endothelial growth factor receptor 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000532">
        <property type="entry name" value="Vascular endothelial growth factor receptor 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000041">
        <property type="entry name" value="Vascular endothelial growth factor receptor 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000143">
        <property type="entry name" value="Vascular endothelial growth factor receptor 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000247">
        <property type="entry name" value="Vascular endothelial growth factor receptor 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000411">
        <property type="entry name" value="Vascular endothelial growth factor receptor 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000479">
        <property type="entry name" value="Vascular endothelial growth factor receptor 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013098">
        <property type="entry name" value="Ig_I-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050122">
        <property type="entry name" value="RTK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001245">
        <property type="entry name" value="Ser-Thr/Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008266">
        <property type="entry name" value="Tyr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020635">
        <property type="entry name" value="Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001824">
        <property type="entry name" value="Tyr_kinase_rcpt_3_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041348">
        <property type="entry name" value="VEGFR-2_TMD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055229">
        <property type="entry name" value="VEGFR1-3_5th"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416">
        <property type="entry name" value="TYROSINE-PROTEIN KINASE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416:SF49">
        <property type="entry name" value="VASCULAR ENDOTHELIAL GROWTH FACTOR RECEPTOR 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07679">
        <property type="entry name" value="I-set"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13927">
        <property type="entry name" value="Ig_3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22971">
        <property type="entry name" value="Ig_VEGFR-1-like_5th"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07714">
        <property type="entry name" value="PK_Tyr_Ser-Thr"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17988">
        <property type="entry name" value="VEGFR-2_TMD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000615">
        <property type="entry name" value="TyrPK_CSF1-R"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01832">
        <property type="entry name" value="VEGFRECEPTOR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01835">
        <property type="entry name" value="VEGFRECEPTR3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00219">
        <property type="entry name" value="TyrKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00109">
        <property type="entry name" value="PROTEIN_KINASE_TYR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00240">
        <property type="entry name" value="RECEPTOR_TYR_KIN_III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0037">Angiogenesis</keyword>
    <keyword evidence="16 27 29" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="20" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="22" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="24 30" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="15" id="KW-0418">Kinase</keyword>
    <keyword evidence="28" id="KW-0460">Magnesium</keyword>
    <keyword evidence="18 32" id="KW-0472">Membrane</keyword>
    <keyword evidence="28" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="14 27 29" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="23" id="KW-0539">Nucleus</keyword>
    <keyword evidence="8" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="21 30" id="KW-0675">Receptor</keyword>
    <keyword evidence="36" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="13" id="KW-0677">Repeat</keyword>
    <keyword evidence="12" id="KW-0732">Signal</keyword>
    <keyword evidence="10" id="KW-0808">Transferase</keyword>
    <keyword evidence="11 30 32" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="17 32" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="19" id="KW-0829">Tyrosine-protein kinase</keyword>
    <feature type="transmembrane region" description="Helical" evidence="32">
        <location>
            <begin position="591"/>
            <end position="615"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="34">
        <location>
            <begin position="40"/>
            <end position="137"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="34">
        <location>
            <begin position="142"/>
            <end position="226"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="34">
        <location>
            <begin position="233"/>
            <end position="368"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="34">
        <location>
            <begin position="371"/>
            <end position="486"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="34">
        <location>
            <begin position="493"/>
            <end position="579"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="33">
        <location>
            <begin position="660"/>
            <end position="987"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="31">
        <location>
            <begin position="291"/>
            <end position="312"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="31">
        <location>
            <begin position="998"/>
            <end position="1026"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="31">
        <location>
            <begin position="1153"/>
            <end position="1184"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="31">
        <location>
            <begin position="297"/>
            <end position="309"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="26">
        <location>
            <position position="851"/>
        </location>
    </feature>
    <feature type="binding site" evidence="27">
        <location>
            <begin position="667"/>
            <end position="674"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="27 29">
        <location>
            <position position="694"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="27">
        <location>
            <position position="855"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="28">
        <location>
            <position position="856"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="28">
        <location>
            <position position="869"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00011902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022258"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022657"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023137"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00051243"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="PIRSR" id="PIRSR000615-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="PIRSR" id="PIRSR000615-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="PIRSR" id="PIRSR000615-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="RuleBase" id="RU000311"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="31">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="32">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="33">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="34">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="35">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033927.1"/>
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    </evidence>
    <evidence type="ECO:0000313" key="36">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1184" mass="134901" checksum="75F4692675467D8C" modified="2019-05-08" version="1">MLVPTHLIRDSLFVQCETIIDKKLFRSNYFIVHVTGNELYDIQLFPKKAMELLVGEKLVLNCTVWAEFNAGVNFQWDYPGKQMHRAVTELPERRSQQTHTELSSILIIQNVSQQDLGKYMCMASNGEQTFKESIDVIVHEKPFINVEWRKGSVIEATAGDEVVKLPVKVVAYPPPEFQWYKDGKLIPNKQSPYSMQIKDVSEQNSGTYTLVLWNRLAGLEERISLQLIVNVPPHIHEKEASSPSIYSRRSHQALTCTVYGIPAPEVIQWQWRPWSPCRMFSRRSLSKHRAARRHQRDRMPECKDWKDVSPQDAENPIESIDTWTESVEGRNKTVSKLVIQEANVSVMYKCVASNKVGRDERLIYFYVTTIPDGFEIESQPSEEPIEGQDLRLSCNADNYTYENLQWYRLNLSKLHHEEGNPLILDCKNVHHYATKMQGELHFKPDSNDAALLLTIPNISLREEGDYVCEVQNRKTREKHCHKKYISVQALEVPRLKQNLTDIWVNVSDSIEMRCKVDGAHIPDISWYKDEKLVEEVSGIDLADSNQRLSIQRVREEDAGLYLCSVCNAKGCVNSSASVSVEGSDDRTNVEIVILIGTGVIAVFFWILLILIFCNIKRPAHADIKTGYLSIIMDPGEVPLEEQCEYLPYDSSKWEFPRERLRLGKVLGHGAFGKVVEASAFGINKSNSCETVAVKMLKEGATASEHKALMSELKILIHIGNHLNVVNLLGACTKPNGPLMVIVEFCKYGNLSNYLRTKREGFSPYREKSPRLRIQVRSLVEAVRADRRSRFGTSDSAIFNRLLMNKSQTAQPIQEVDDLWQSPLTMEDLICYSFQVARGMEFLASRKCIHRDLAARNILLSENNVVKICDFGLARDIYKDPDYVRKGSARLPLKWMAPESIFDKVYTTQSDVWSFGVLLWEIFSLGASPYPGVQINEEFCQRLKDGTRMRAPEYATAEIYRIMLSCWHGDPKERPTFSDLVEILGDLLQENVQQEGKDYIPLNDSQSSEDDGFSQVPSSAQQNSDEEEFDMRLRCHSLAARYYNCVSFPGCLTGGNQIRCPSRIKTFEEFPMTHTTYKTHPDNQTDSGMVLASEEFERIENRHRKEGGFSSKGPSRNAELAVEQLAGVDLRDRCRPLYQSQVGGQTFYNSEYGELSEHSEDGSYTPPGEGASPSAVHASFFSEQY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0P4</accession>
    <name>A0A452J0P4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">Myosin heavy chain 6</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">Muscle contraction.</text>
    </comment>
    <comment type="subunit">
        <text evidence="14">Muscle myosin is a hexameric protein that consists of 2 heavy chain subunits (MHC), 2 alkali light chain subunits (MLC) and 2 regulatory light chain subunits (MLC-2).</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Myofibril</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 15">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038426.1">
        <property type="protein sequence ID" value="ENSGAGP00000033930.1"/>
        <property type="gene ID" value="ENSGAGG00000020688.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030016">
        <property type="term" value="C:myofibril"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032982">
        <property type="term" value="C:myosin filament"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016460">
        <property type="term" value="C:myosin II complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000146">
        <property type="term" value="F:microfilament motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd01377">
        <property type="entry name" value="MYSc_class_II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000003">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000004">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000006">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000013">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000007">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000008">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.4820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.530:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.360:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000024">
        <property type="entry name" value="Myosin heavy chain, isoform J"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.720:FF:000001">
        <property type="entry name" value="Myosin heavy chain, muscle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.340">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.370">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.4820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.2420">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.360">
        <property type="entry name" value="Myosin S1 fragment, N-terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008989">
        <property type="entry name" value="Myosin_S1_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002928">
        <property type="entry name" value="Myosin_tail"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004009">
        <property type="entry name" value="SH3_Myosin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014751">
        <property type="entry name" value="XRCC4-like_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615">
        <property type="entry name" value="MYOSIN HEAVY CHAIN, NON-MUSCLE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615:SF69">
        <property type="entry name" value="MYOSIN-6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02736">
        <property type="entry name" value="Myosin_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01576">
        <property type="entry name" value="Myosin_tail_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90257">
        <property type="entry name" value="Myosin rod fragments"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57997">
        <property type="entry name" value="Tropomyosin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51844">
        <property type="entry name" value="SH3_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12 15" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="7 15" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="8 16" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0488">Methylation</keyword>
    <keyword evidence="10 15" id="KW-0505">Motor protein</keyword>
    <keyword evidence="11" id="KW-0514">Muscle protein</keyword>
    <keyword evidence="9 15" id="KW-0518">Myosin</keyword>
    <keyword evidence="6 15" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0787">Thick filament</keyword>
    <feature type="domain" description="Myosin N-terminal SH3-like" evidence="18">
        <location>
            <begin position="10"/>
            <end position="59"/>
        </location>
    </feature>
    <feature type="domain" description="Myosin motor" evidence="17">
        <location>
            <begin position="63"/>
            <end position="754"/>
        </location>
    </feature>
    <feature type="region of interest" description="Actin-binding" evidence="15">
        <location>
            <begin position="631"/>
            <end position="653"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="16">
        <location>
            <begin position="816"/>
            <end position="1132"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="16">
        <location>
            <begin position="1176"/>
            <end position="1899"/>
        </location>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <begin position="156"/>
            <end position="163"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004657"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022433"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023179"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00037488"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00038612"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS51844"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033930.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="21">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1913" mass="221216" checksum="77AB6F9D984E3BB3" modified="2019-05-08" version="1">MVNYYLFDSIGKRRCWIPDEKKAYTEGDIKESSGGKVTVETVDGKTVTVKEDDVQPMNPPKFDMIEDMAMLTHLNEASVLHNLNRRYSHWMIYTYSGLFCVTINPYKWLPVYNPEVVNAYKGKRRSEAPPHIFSIADNAYNDMLRNRENQSMLITGESGAGKTVNTKRVIQYFATVAALGEPGKKSSPAAGTLEDQIIQANPALEAFGNAKTLRNDNSSRFGKFIRIHFGTTGKLSSADIEIYLLEKSRVIFQQPGERSYHIFYQILSGKKAELQDMLLVSTNPYDYHFSSQGAVTVDNLDDGEELMATDQAMDILGFIPDEKYGAYKLTGSIMHFGNMKFKQRPREEQAETDGTESTDKAAYLMGLNSSDLAKGLLHPRVKVGNEYVTKGQSVEQVVYAVGALSKAVYDRMFKWLVIRINKTLDTRLPRQFFIGVLDIAGFEIFDYNSFEQLCINYTNEKLQQFFNHHMFVLEQEEYKKEGIEWVFIDFGLDLQACIDLIEKPLGILSILEEECMFPKATDMTFKAKLYDNHLGKSPNLQKPRIDKKRKYEAHFELLHYAGAVPYNIIGWLEKNKDPLNETVVGIFQKSANKLLASLFENYVSADSGNTEKKRKKGASFQTVSSLHKENLNKLMTNLRSTAPHFVRCIIPNETKTPGAMDAYLVLHQLRCNGVLEGIRICRKGFPNRVLYADFKQRYRILNPAAIPEDKFVDSRKASEKLLASLDIDQTQYRFGHTKVFFKAGLLGHLEEMRDHRLSKILTMIQARARGKLMRIEFQKIVERRDALLVIQWNIRAFMAVKNWPWMKLFFKIKPLLKSAETEKEMANMKEEFLKLKEALEKSEARRKELEEKQVSLVQEKNDLLLQLQAEQDTLADAEERCDLLIKSKIQMEAKVKELTERVEDEEEMNSELTSKKRKLEDECSELKKDIDDLEITLAKVEKEKHATENKVKNLTEEMASLDETISKLTKEKKSLQEAHQQALDDLQAEEDKVNTLSKTKLKLEQQVDDLEGSLEQEKKIRMDLERVKRKLEGDLKLTQESLMDLENDKLQLDEKLKKKEFEINQLNSKLEDEQTIVMQLQKKIKELQARIEELEEELEAERAARAKVEKQRSDLARELEELSERLEEAGGATAAQLEMNKKREAEFLKVRRDLEEATLHYEATAATLRKKHADSMAELGEQVDNLQRVKQKLEKEKSELKLEVDDLLSNMEQMVKGKSNAEKLCRTYEDNLNETKTKLDEMTRLMNDLTTQKGKLQTENGEFARQLEEKESLISQLSRGKTSFTQQIEELRRLLEEETKSKNALAHALQAARHDCDLLREQFEEEQEAKAELQRALSKGNAEVAQWRTKYETDAIQRTEELEEAKKKLAARLQEAEEAIEAANAKCSSLEKTKHRLQNELEDIVIDLERANSAAAALDKKQRNFDKLISDWKQKYEETQAELEASQKEARSLSTELFKLKNAYEETLDNLETVKRENKNLQEEISDLTDQISEGNKSLHEIEKIKKQVEQEKSEVQLALEEAEGALEHEESKSLRFQLEISQIKAEFERKLAEKDEEMENIRRNQQRTIDSLQSTLDSEAKSRNEAIRLKKKMEGDLNEMEIQLSHANRHASEATKSARILQAQIKDLQVQLDDSEHLNEDLKEQLAVTDRRNNLIQSELDELRALLDQTERGRKLAEQELLEATERVNLLHTQNTSLINQKKKLETDISQMQNEVEEAIQECRNAEEKAKKAITDAAMMAEELKKEQDTSAHLERMKKNMEQTIKDLQKRLDEAEQIALKGGKKQIQKLESRVRELENDLENELRRNTESQKGVRKYERRIKELTYQAEEDKKNLTRMQELIDKLQLKVKSYKHQAEEAEAQSNQYLAKYRKQQHELDDAEERAEIAESQVNKLRTKTRDIGIKKVHEEEE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0P5</accession>
    <name>A0A452J0P5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">E3 ubiquitin-protein ligase</fullName>
            <ecNumber evidence="8">2.3.2.27</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="8">RING-type E3 ubiquitin transferase</fullName>
        </alternativeName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="8">E3 ubiquitin ligase.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>S-ubiquitinyl-[E2 ubiquitin-conjugating enzyme]-L-cysteine + [acceptor protein]-L-lysine = [E2 ubiquitin-conjugating enzyme]-L-cysteine + N(6)-ubiquitinyl-[acceptor protein]-L-lysine.</text>
            <dbReference type="EC" id="2.3.2.27"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="8">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="2.3.2.27" evidence="8"/>
    <dbReference type="AlphaFoldDB" id="A0A452J0P5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038470.1">
        <property type="protein sequence ID" value="ENSGAGP00000033970.1"/>
        <property type="gene ID" value="ENSGAGG00000024152.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005769">
        <property type="term" value="C:early endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061630">
        <property type="term" value="F:ubiquitin protein ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008333">
        <property type="term" value="P:endosome to lysosome transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043951">
        <property type="term" value="P:negative regulation of cAMP-mediated signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045744">
        <property type="term" value="P:negative regulation of G protein-coupled receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016567">
        <property type="term" value="P:protein ubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="CDD" id="cd16816">
        <property type="entry name" value="mRING-HC-C3HC5_MGRN1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.40.10:FF:000013">
        <property type="entry name" value="E3 ubiquitin-protein ligase MGRN1 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045194">
        <property type="entry name" value="MGRN1/RNF157-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058981">
        <property type="entry name" value="MGRN1/RNF157-like_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001841">
        <property type="entry name" value="Znf_RING"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22996:SF2">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE MGRN1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22996">
        <property type="entry name" value="MAHOGUNIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26192">
        <property type="entry name" value="RNF157-like_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13920">
        <property type="entry name" value="zf-C3HC4_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00184">
        <property type="entry name" value="RING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57850">
        <property type="entry name" value="RING/U-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50089">
        <property type="entry name" value="ZF_RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="3 8" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 8" id="KW-0808">Transferase</keyword>
    <keyword evidence="5 8" id="KW-0833">Ubl conjugation pathway</keyword>
    <keyword evidence="6 8" id="KW-0862">Zinc</keyword>
    <keyword evidence="4 7 8" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="RING-type" evidence="10">
        <location>
            <begin position="279"/>
            <end position="318"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="404"/>
            <end position="491"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="9">
        <location>
            <begin position="442"/>
            <end position="453"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="RuleBase" id="RU369081"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033970.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="536" mass="59223" checksum="628932E25F37D218" modified="2019-05-08" version="1">MGSLLSRRIAGVEDIDIQANSAYRYPPKSGNYFASHFFMGGEKFDTPHPEGYLFGENMDLNFLGNRPVQFPYVTPAPHEPVKTLRSLVNIRKDSLRLVRYKDDLDSPTEENGKQKVLYSLEFTFDADARVAITIYCQATEEFISGMAVYSTKNPSLQSETVHYKRGVSQQFSLPSFKIDFSDWKDDELNFDLDRGVFPVVIQAVVDEGDVVVEVTGHAHVLLAAFEKHVDGSFSVKPLKQKQIVDRVSYLLQEIYGIENKNNQETKPSDDENSDNSNECVVCLSDLRDTLILPCRHLCLCNSCADTLRYQANNCPICRLPFRALLQIRAVRKKPGALSPVSFSTVLAQTMDHDEHSSSDNIPPGYEPISLLEALNGLRSISPSIPSAPLYEEISYTGMVDGLPPAGRPLAGIDRATESSPQKSKRSKSPDSALRSPSSPIHEEDEEKLSEDSDSQPALSGAELGLRENSSPGSMVAEEVEEISSLQQGSYLPSMEDILPENICSEHRSLTQSESDSAVNVYLPALGPDSCSIGIEE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0P6</accession>
    <name>A0A452J0P6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Protein arginine methyltransferase NDUFAF7</fullName>
            <ecNumber evidence="7">2.1.1.320</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="7">Arginine methyltransferase involved in the assembly or stability of mitochondrial NADH:ubiquinone oxidoreductase complex (complex I).</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="6 7">
            <text>L-arginyl-[protein] + 2 S-adenosyl-L-methionine = N(omega),N(omega)'-dimethyl-L-arginyl-[protein] + 2 S-adenosyl-L-homocysteine + 2 H(+)</text>
            <dbReference type="Rhea" id="RHEA:48108"/>
            <dbReference type="Rhea" id="RHEA-COMP:10532"/>
            <dbReference type="Rhea" id="RHEA-COMP:11992"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29965"/>
            <dbReference type="ChEBI" id="CHEBI:57856"/>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
            <dbReference type="ChEBI" id="CHEBI:88221"/>
            <dbReference type="EC" id="2.1.1.320"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 7">Mitochondrion</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 7">Belongs to the NDUFAF7 family.</text>
    </comment>
    <dbReference type="EC" id="2.1.1.320" evidence="7"/>
    <dbReference type="AlphaFoldDB" id="A0A452J0P6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038373.1">
        <property type="protein sequence ID" value="ENSGAGP00000033881.1"/>
        <property type="gene ID" value="ENSGAGG00000024102.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035243">
        <property type="term" value="F:protein-arginine omega-N symmetric methyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032259">
        <property type="term" value="P:methylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032981">
        <property type="term" value="P:mitochondrial respiratory chain complex I assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.12710">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003788">
        <property type="entry name" value="NDUFAF7"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038375">
        <property type="entry name" value="NDUFAF7_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029063">
        <property type="entry name" value="SAM-dependent_MTases_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12049">
        <property type="entry name" value="PROTEIN ARGININE METHYLTRANSFERASE NDUFAF7, MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12049:SF7">
        <property type="entry name" value="PROTEIN ARGININE METHYLTRANSFERASE NDUFAF7, MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02636">
        <property type="entry name" value="Methyltransf_28"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53335">
        <property type="entry name" value="S-adenosyl-L-methionine-dependent methyltransferases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 7" id="KW-0489">Methyltransferase</keyword>
    <keyword evidence="5 7" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0732">Signal</keyword>
    <keyword evidence="4 7" id="KW-0808">Transferase</keyword>
    <feature type="signal peptide" evidence="8">
        <location>
            <begin position="1"/>
            <end position="25"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019214530" description="Protein arginine methyltransferase NDUFAF7" evidence="8">
        <location>
            <begin position="26"/>
            <end position="415"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004173"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005891"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022603"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00048612"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU364114"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033881.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="415" mass="46784" checksum="6743C77FE190F20A" modified="2019-05-08" version="1">MARLPARRALLAAVLGVSWQQRLLGHHVVPAAWRRRRFSAGSRAEDSSPVTPMLKHLMFKIKSTGPITVAEYMREVLTNPVKGYYMHHDMLGERGDFVTSPEISQIFGEVFNQLGSLLNKCDISIHLVEVSPKLSEIQAFTLTGGKIQPELDVTSPVYMKGISKAGIPIFWYRELQDVPQGTSFYLAHEFFDALPIHKFQRTEKGWREVLIDIDPEAPDQLRFVLAPSTTPATELFIQRKEVRDHVEVCPDAGILIQRLACRIQEDGGAALIADYGHNGNKTDTFRSFRSHELHDALTGPGTADLTADVDFSYLRTMTQGRVATLGPLNQHEFLKNMGIDVRLQVLLQNSSDKAMHKHLLQGYDMLMNPKKMGDRFHFFALLPHQRLAHSDQKVNLRHSRSLSATVAGFGELVWK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0Q2</accession>
    <name>A0A452J0Q2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">Myosin motor domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">Muscle contraction.</text>
    </comment>
    <comment type="subunit">
        <text evidence="14">Muscle myosin is a hexameric protein that consists of 2 heavy chain subunits (MHC), 2 alkali light chain subunits (MLC) and 2 regulatory light chain subunits (MLC-2).</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Myofibril</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 15">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038443.1">
        <property type="protein sequence ID" value="ENSGAGP00000033946.1"/>
        <property type="gene ID" value="ENSGAGG00000020688.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030016">
        <property type="term" value="C:myofibril"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032982">
        <property type="term" value="C:myosin filament"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016460">
        <property type="term" value="C:myosin II complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000146">
        <property type="term" value="F:microfilament motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000003">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000004">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000006">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000013">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000007">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000008">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.4820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.530:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000024">
        <property type="entry name" value="Myosin heavy chain, isoform J"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.720:FF:000001">
        <property type="entry name" value="Myosin heavy chain, muscle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.340">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.370">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.4820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.2420">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002928">
        <property type="entry name" value="Myosin_tail"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014751">
        <property type="entry name" value="XRCC4-like_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615">
        <property type="entry name" value="MYOSIN HEAVY CHAIN, NON-MUSCLE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615:SF69">
        <property type="entry name" value="MYOSIN-6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01576">
        <property type="entry name" value="Myosin_tail_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90257">
        <property type="entry name" value="Myosin rod fragments"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57997">
        <property type="entry name" value="Tropomyosin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12 15" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="7 15" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="8 16" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0488">Methylation</keyword>
    <keyword evidence="10 15" id="KW-0505">Motor protein</keyword>
    <keyword evidence="11" id="KW-0514">Muscle protein</keyword>
    <keyword evidence="9 15" id="KW-0518">Myosin</keyword>
    <keyword evidence="6 15" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0787">Thick filament</keyword>
    <feature type="domain" description="Myosin motor" evidence="17">
        <location>
            <begin position="84"/>
            <end position="776"/>
        </location>
    </feature>
    <feature type="region of interest" description="Actin-binding" evidence="15">
        <location>
            <begin position="653"/>
            <end position="675"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="16">
        <location>
            <begin position="838"/>
            <end position="1154"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="16">
        <location>
            <begin position="1198"/>
            <end position="1921"/>
        </location>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <begin position="177"/>
            <end position="184"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004657"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022433"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023179"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00037488"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00038612"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033946.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1933" mass="223322" checksum="F2F11DDF42D243E4" modified="2019-05-08" version="1">ELAKMMDMSEFGEAAPFLRKSDKEIMILQTVAFDGKTGYVFLCLVCSVERQGDSSCQQCYTCLSLQTVTVKEDDVQPMNPPKFDMIEDMAMLTHLNEASVLHNLNRRYSHWMIYTYSGLFCVTINPYKWLPVYNPEVVNAYKGKRRSEAPPHIFSIADNAYNDMLRNRENQSMLITGESGAGKTVNTKRVIQYFATVAALGEPGKKSVSMQGGTLEDQIIQANPALEAFGNAKTLRNDNSSRFGKFIRIHFGTTGKLSSADIEIYLLEKSRVIFQQPGERSYHIFYQILSGKKAELQDMLLVSTNPYDYHFSSQGAVTVDNLDDGEELMATDQAMDILGFIPDEKYGAYKLTGSIMHFGNMKFKQRPREEQAETDGTESTDKAAYLMGLNSSDLAKGLLHPRVKVGNEYVTKGQSVEQVVYAVGALSKAVYDRMFKWLVIRINKTLDTRLPRQFFIGVLDIAGFEIFDYNSFEQLCINYTNEKLQQFFNHHMFVLEQEEYKKEGIEWVFIDFGLDLQACIDLIEKPLGILSILEEECMFPKATDMTFKAKLYDNHLGKSPNLQKPRIDKKRKYEAHFELLHYAGAVPYNIIGWLEKNKDPLNETVVGIFQKSANKLLASLFENYVSADSGNTEKKRKKGASFQTVSSLHKENLNKLMTNLRSTAPHFVRCIIPNETKTPGAMDAYLVLHQLRCNGVLEGIRICRKGFPNRVLYADFKQRYRILNPAAIPEDKFVDSRKASEKLLASLDIDQTQYRFGHTKVFFKAGLLGHLEEMRDHRLSKILTMIQARARGKLMRIEFQKIVERRDALLVIQWNIRAFMAVKNWPWMKLFFKIKPLLKSAETEKEMANMKEEFLKLKEALEKSEARRKELEEKQVSLVQEKNDLLLQLQAEQDTLADAEERCDLLIKSKIQMEAKVKELTERVEDEEEMNSELTSKKRKLEDECSELKKDIDDLEITLAKVEKEKHATENKVKNLTEEMASLDETISKLTKEKKSLQEAHQQALDDLQAEEDKVNTLSKTKLKLEQQVDDLEGSLEQEKKIRMDLERVKRKLEGDLKLTQESLMDLENDKLQLDEKLKKKEFEINQLNSKLEDEQTIVMQLQKKIKELQARIEELEEELEAERAARAKVEKQRSDLARELEELSERLEEAGGATAAQLEMNKKREAEFLKVRRDLEEATLHYEATAATLRKKHADSMAELGEQVDNLQRVKQKLEKEKSELKLEVDDLLSNMEQMVKGKSNAEKLCRTYEDNLNETKTKLDEMTRLMNDLTTQKGKLQTENGEFARQLEEKESLISQLSRGKTSFTQQIEELRRLLEEETKSKNALAHALQAARHDCDLLREQFEEEQEAKAELQRALSKGNAEVAQWRTKYETDAIQRTEELEEAKKKLAARLQEAEEAIEAANAKCSSLEKTKHRLQNELEDIVIDLERANSAAAALDKKQRNFDKLISDWKQKYEETQAELEASQKEARSLSTELFKLKNAYEETLDNLETVKRENKNLQEEISDLTDQISEGNKSLHEIEKIKKQVEQEKSEVQLALEEAEGALEHEESKSLRFQLEISQIKAEFERKLAEKDEEMENIRRNQQRTIDSLQSTLDSEAKSRNEAIRLKKKMEGDLNEMEIQLSHANRHASEATKSARILQAQIKDLQVQLDDSEHLNEDLKEQLAVTDRRNNLIQSELDELRALLDQTERGRKLAEQELLEATERVNLLHTQNTSLINQKKKLETDISQMQNEVEEAIQECRNAEEKAKKAITDAAMMAEELKKEQDTSAHLERMKKNMEQTIKDLQKRLDEAEQIALKGGKKQIQKLESRVRELENDLENELRRNTESQKGVRKYERRIKELTYQAEEDKKNLTRMQELIDKLQLKVKSYKHQAEEAEAQSNQYLAKYRKQQHELDDAEERAEIAESQVNKLRTKTRDIGIKKVCAQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0Q4</accession>
    <name>A0A452J0Q4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">Myosin heavy chain 6</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">Muscle contraction.</text>
    </comment>
    <comment type="subunit">
        <text evidence="14">Muscle myosin is a hexameric protein that consists of 2 heavy chain subunits (MHC), 2 alkali light chain subunits (MLC) and 2 regulatory light chain subunits (MLC-2).</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Myofibril</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 15">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038435.1">
        <property type="protein sequence ID" value="ENSGAGP00000033939.1"/>
        <property type="gene ID" value="ENSGAGG00000020688.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030016">
        <property type="term" value="C:myofibril"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032982">
        <property type="term" value="C:myosin filament"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016460">
        <property type="term" value="C:myosin II complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000146">
        <property type="term" value="F:microfilament motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000003">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000004">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000006">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000013">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000007">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000008">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.4820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.530:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.360:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000024">
        <property type="entry name" value="Myosin heavy chain, isoform J"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.720:FF:000001">
        <property type="entry name" value="Myosin heavy chain, muscle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.340">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.370">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.4820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.2420">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.360">
        <property type="entry name" value="Myosin S1 fragment, N-terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008989">
        <property type="entry name" value="Myosin_S1_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002928">
        <property type="entry name" value="Myosin_tail"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004009">
        <property type="entry name" value="SH3_Myosin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014751">
        <property type="entry name" value="XRCC4-like_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615">
        <property type="entry name" value="MYOSIN HEAVY CHAIN, NON-MUSCLE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615:SF69">
        <property type="entry name" value="MYOSIN-6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02736">
        <property type="entry name" value="Myosin_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01576">
        <property type="entry name" value="Myosin_tail_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90257">
        <property type="entry name" value="Myosin rod fragments"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57997">
        <property type="entry name" value="Tropomyosin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51844">
        <property type="entry name" value="SH3_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12 15" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="7 15" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="8 16" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0488">Methylation</keyword>
    <keyword evidence="10 15" id="KW-0505">Motor protein</keyword>
    <keyword evidence="11" id="KW-0514">Muscle protein</keyword>
    <keyword evidence="9 15" id="KW-0518">Myosin</keyword>
    <keyword evidence="6 15" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0787">Thick filament</keyword>
    <feature type="domain" description="Myosin N-terminal SH3-like" evidence="18">
        <location>
            <begin position="10"/>
            <end position="59"/>
        </location>
    </feature>
    <feature type="domain" description="Myosin motor" evidence="17">
        <location>
            <begin position="63"/>
            <end position="753"/>
        </location>
    </feature>
    <feature type="region of interest" description="Actin-binding" evidence="15">
        <location>
            <begin position="630"/>
            <end position="652"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="16">
        <location>
            <begin position="815"/>
            <end position="1131"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="16">
        <location>
            <begin position="1175"/>
            <end position="1898"/>
        </location>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <begin position="156"/>
            <end position="163"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004657"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022433"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023179"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00037488"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00038612"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS51844"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033939.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="21">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1912" mass="221170" checksum="94E9669447313A4D" modified="2019-05-08" version="1">MVNYYLFDSIGKRRCWIPDEKKAYTEGDIKESSGGKVTVETVDGKTVTVKEDDVQPMNPPKFDMIEDMAMLTHLNEASVLHNLNRRYSHWMIYTYSGLFCVTINPYKWLPVYNPEVVNAYKGKRRSEAPPHIFSIADNAYNDMLRNRENQSMLITGESGAGKTVNTKRVIQYFATVAALGEPGKKNVPGGTLEDQIIQANPALEAFGNAKTLRNDNSSRFGKFIRIHFGTTGKLSSADIEIYLLEKSRVIFQQPGERSYHIFYQILSGKKAELQDMLLVSTNPYDYHFSSQGAVTVDNLDDGEELMATDQAMDILGFIPDEKYGAYKLTGSIMHFGNMKFKQRPREEQAETDGTESTDKAAYLMGLNSSDLAKGLLHPRVKVGNEYVTKGQSVEQVVYAVGALSKAVYDRMFKWLVIRINKTLDTRLPRQFFIGVLDIAGFEIFDYNSFEQLCINYTNEKLQQFFNHHMFVLEQEEYKKEGIEWVFIDFGLDLQACIDLIEKPLGILSILEEECMFPKATDMTFKAKLYDNHLGKSPNLQKPRIDKKRKYEAHFELLHYAGAVPYNIIGWLEKNKDPLNETVVGIFQKSANKLLASLFENYVSADSGNTEKKRKKGASFQTVSSLHKENLNKLMTNLRSTAPHFVRCIIPNETKTPGAMDAYLVLHQLRCNGVLEGIRICRKGFPNRVLYADFKQRYRILNPAAIPEDKFVDSRKASEKLLASLDIDQTQYRFGHTKVFFKAGLLGHLEEMRDHRLSKILTMIQARARGKLMRIEFQKIVERRDALLVIQWNIRAFMAVKNWPWMKLFFKIKPLLKSAETEKEMANMKEEFLKLKEALEKSEARRKELEEKQVSLVQEKNDLLLQLQAEQDTLADAEERCDLLIKSKIQMEAKVKELTERVEDEEEMNSELTSKKRKLEDECSELKKDIDDLEITLAKVEKEKHATENKVKNLTEEMASLDETISKLTKEKKSLQEAHQQALDDLQAEEDKVNTLSKTKLKLEQQVDDLEGSLEQEKKIRMDLERVKRKLEGDLKLTQESLMDLENDKLQLDEKLKKKEFEINQLNSKLEDEQTIVMQLQKKIKELQARIEELEEELEAERAARAKVEKQRSDLARELEELSERLEEAGGATAAQLEMNKKREAEFLKVRRDLEEATLHYEATAATLRKKHADSMAELGEQVDNLQRVKQKLEKEKSELKLEVDDLLSNMEQMVKGKSNAEKLCRTYEDNLNETKTKLDEMTRLMNDLTTQKGKLQTENGEFARQLEEKESLISQLSRGKTSFTQQIEELRRLLEEETKSKNALAHALQAARHDCDLLREQFEEEQEAKAELQRALSKGNAEVAQWRTKYETDAIQRTEELEEAKKKLAARLQEAEEAIEAANAKCSSLEKTKHRLQNELEDIVIDLERANSAAAALDKKQRNFDKLISDWKQKYEETQAELEASQKEARSLSTELFKLKNAYEETLDNLETVKRENKNLQEEISDLTDQISEGNKSLHEIEKIKKQVEQEKSEVQLALEEAEGALEHEESKSLRFQLEISQIKAEFERKLAEKDEEMENIRRNQQRTIDSLQSTLDSEAKSRNEAIRLKKKMEGDLNEMEIQLSHANRHASEATKSARILQAQIKDLQVQLDDSEHLNEDLKEQLAVTDRRNNLIQSELDELRALLDQTERGRKLAEQELLEATERVNLLHTQNTSLINQKKKLETDISQMQNEVEEAIQECRNAEEKAKKAITDAAMMAEELKKEQDTSAHLERMKKNMEQTIKDLQKRLDEAEQIALKGGKKQIQKLESRVRELENDLENELRRNTESQKGVRKYERRIKELTYQAEEDKKNLTRMQELIDKLQLKVKSYKHQAEEAEAQSNQYLAKYRKQQHELDDAEERAEIAESQVNKLRTKTRDIGIKKVHEEEE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0Q5</accession>
    <name>A0A452J0Q5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Myelin transcription factor 1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the MYT1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0Q5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038481.1">
        <property type="protein sequence ID" value="ENSGAGP00000033981.1"/>
        <property type="gene ID" value="ENSGAGG00000024142.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.320.30:FF:000001">
        <property type="entry name" value="Myelin transcription factor 1-like, a"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.320.30">
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013681">
        <property type="entry name" value="Myelin_TF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002515">
        <property type="entry name" value="Znf_C2H2C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036060">
        <property type="entry name" value="Znf_C2H2C_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10816:SF10">
        <property type="entry name" value="MYELIN TRANSCRIPTION FACTOR 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10816">
        <property type="entry name" value="MYELIN TRANSCRIPTION FACTOR 1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08474">
        <property type="entry name" value="MYT1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01530">
        <property type="entry name" value="zf-C2HC"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103637">
        <property type="entry name" value="CCHHC domain"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51802">
        <property type="entry name" value="ZF_CCHHC"/>
        <property type="match status" value="7"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="8" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="9" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 11" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="Myelin transcription factor 1" evidence="14">
        <location>
            <begin position="541"/>
            <end position="596"/>
        </location>
    </feature>
    <feature type="domain" description="Myelin transcription factor 1" evidence="14">
        <location>
            <begin position="597"/>
            <end position="771"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="83"/>
            <end position="128"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="248"/>
            <end position="347"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="498"/>
            <end position="521"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="648"/>
            <end position="755"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="12">
        <location>
            <begin position="978"/>
            <end position="1040"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="13">
        <location>
            <begin position="90"/>
            <end position="124"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="13">
        <location>
            <begin position="255"/>
            <end position="292"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="13">
        <location>
            <begin position="306"/>
            <end position="318"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="13">
        <location>
            <begin position="328"/>
            <end position="340"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="13">
        <location>
            <begin position="663"/>
            <end position="704"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="13">
        <location>
            <begin position="740"/>
            <end position="751"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010194"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01143"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="Pfam" id="PF08474"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033981.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1106" mass="122712" checksum="BA8F3BCE0E299151" modified="2019-05-08" version="1">SSHPIPVRELRGRAGQFRGPDVARGLYCPTPGCNGSGHIRGKYARHRSLQSCPLAKKRKLQDAEAEHLVSKRKSHPLKLALDEGYNVDSDGSEDTEVKEESVTDESEGTLEEEETETIEQEETPSPEPAEGALSLLIADTPGPSKANYSSYQEIIANSLLNLGQIAEETLAIEGQMPENELQVSKPPVNVVHLVHEGVEEEVVEEECDKEIIIQTEDAEEVIEVTSERSSELCPEHRDNMNCEESCKLQKAMQAEAEDEEDDEEEEEEEEEEEEEEEEEEEEEEEEEEEELASEVICEETPQTSRDSQKTHCEGRFSPKPEYSVIVEVRSDDDKDDDARSQKSTVTDESEMYDMMTRGNLGLLEQAIALKAEQVKIMREPNRLPGEHVKHFQVDEKQNKPLDTIRKNYYSKDPSRPEKREIKCPTPGCDGTGHVTGLYPHHRSLSGCPHKDRIPPEILAMHENVLKCPTPGCTGQGHVNSNRNTHRSLSGCPIAAAEKLAKSHEKQQPQAGDPSKSSSNSDRILRPMCFVKQLEIPQYGSYRPNMVPATPRANLAKELEKYSKVTFDYASFDAQVFGKRMLAPKIQTSETSPKAFKCFDYSHDAEAAHMAATAILNLSTRCWEMPENLSTKQQDLPSKSMDIEVDENGTLDLSMNKHRKRESTFPSSSSCSSSPSVKSPDVSQRQNSTSATSSTMTSPQSSQTSRQDEWDGPIDYTKPNRQREEEPEESEPAAHSFASSEADEQEVSEENFEERKYPGEVTLTNFKLKFLPKDIKKELLTCPTPGCDGSGHITGNYASHRSLSGCPLADKSLRNLMAAHSADLKCPTPGCDGSGHITGNYASHRSLSGCPRAKKSGIKITPTKDDKEDPELMKCPVPGCVGLGHISGKYASHRSASGCPLAARRQKEGSLNGSSFSWKSLKNEGPTCPTPGCDGSGHANGSFLTHRSLSGCPRATFAGKKGKLSGDEILNTKFKTSDVLENDEEIKQLNKEITELNESNSEMEAAMVKLQSQISTMEKNLKNIEEENKIIEEQNEALFLELSGLSQALIQSLANIRLPHMEPISEQNFDAYVNTLTDMYTNQECYQNPENRDLLESIKQAVKGIQV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0Q6</accession>
    <name>A0A452J0Q6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Cortexin 3</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0Q6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038383.1">
        <property type="protein sequence ID" value="ENSGAGP00000033891.1"/>
        <property type="gene ID" value="ENSGAGG00000024108.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020066">
        <property type="entry name" value="Cortexin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16736">
        <property type="entry name" value="CORTEXIN-1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16736:SF4">
        <property type="entry name" value="CORTEXIN-2-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11057">
        <property type="entry name" value="Cortexin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4 5" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 5" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="3 5" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="30"/>
            <end position="50"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004167"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033891.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="8">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="79" mass="8894" checksum="92F147CC5568FE62" modified="2019-05-08" version="1">FDPGQTATALPVPGVALPIPTLSLTLEQKAAFVFIFFLFIFLGLLIVRCFQILLDPYRSMPSSTRMDYMEKDTSDYCLP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0R0</accession>
    <name>A0A452J0R0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Myelin transcription factor 1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the MYT1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0R0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038486.1">
        <property type="protein sequence ID" value="ENSGAGP00000033986.1"/>
        <property type="gene ID" value="ENSGAGG00000024142.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.320.30:FF:000001">
        <property type="entry name" value="Myelin transcription factor 1-like, a"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.320.30">
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013681">
        <property type="entry name" value="Myelin_TF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002515">
        <property type="entry name" value="Znf_C2H2C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036060">
        <property type="entry name" value="Znf_C2H2C_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10816:SF10">
        <property type="entry name" value="MYELIN TRANSCRIPTION FACTOR 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10816">
        <property type="entry name" value="MYELIN TRANSCRIPTION FACTOR 1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08474">
        <property type="entry name" value="MYT1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01530">
        <property type="entry name" value="zf-C2HC"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103637">
        <property type="entry name" value="CCHHC domain"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51802">
        <property type="entry name" value="ZF_CCHHC"/>
        <property type="match status" value="7"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="8" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="9" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 11" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="Myelin transcription factor 1" evidence="14">
        <location>
            <begin position="553"/>
            <end position="608"/>
        </location>
    </feature>
    <feature type="domain" description="Myelin transcription factor 1" evidence="14">
        <location>
            <begin position="609"/>
            <end position="783"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="67"/>
            <end position="159"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="234"/>
            <end position="359"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="510"/>
            <end position="533"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="660"/>
            <end position="767"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="12">
        <location>
            <begin position="990"/>
            <end position="1052"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="13">
        <location>
            <begin position="94"/>
            <end position="128"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="13">
        <location>
            <begin position="139"/>
            <end position="159"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="13">
        <location>
            <begin position="237"/>
            <end position="259"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="13">
        <location>
            <begin position="267"/>
            <end position="304"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="13">
        <location>
            <begin position="318"/>
            <end position="330"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="13">
        <location>
            <begin position="340"/>
            <end position="352"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="13">
        <location>
            <begin position="675"/>
            <end position="716"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="13">
        <location>
            <begin position="752"/>
            <end position="763"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010194"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01143"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="Pfam" id="PF08474"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033986.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1118" mass="124172" checksum="8BC2AECCA785C7D4" modified="2019-05-08" version="1">KAVEKEREREKWVNVAKKGEPLTNHGALCHSCPTPGCNGSGHIRGKYARHRSLQSCPLAKKRKLQDAEAEHLVSKRKSHPLKLALDEGYNVDSDGSEDTEVKEESVTDESEGTLEEEETETIEQEETPSPEPAEVRSPAKSSRYGQNAATSTPGPSKANYSSYQEIIANSLLNLGQIAEETLAIEGQMPENELQVSKPPVNVVHLVHEGVEEEVVEEECDKEIIIQTEDAEEVIEVTSERSSELCPEHRDNMNCEESCKLQKAMQAEAEDEEDDEEEEEEEEEEEEEEEEEEEEEEEEEEEELASEVICEETPQTSRDSQKTHCEGRFSPKPEYSVIVEVRSDDDKDDDARSQKSTVTDESEMYDMMTRGNLGLLEQAIALKAEQVKIMREPNRLPGEHVKHFQVDEKQNKPLDTIRKNYYSKDPSRPEKREIKCPTPGCDGTGHVTGLYPHHRSLSGCPHKDRIPPEILAMHENVLKCPTPGCTGQGHVNSNRNTHRSLSGCPIAAAEKLAKSHEKQQPQAGDPSKSSSNSDRILRPMCFVKQLEIPQYGSYRPNMVPATPRANLAKELEKYSKVTFDYASFDAQVFGKRMLAPKIQTSETSPKAFKCFDYSHDAEAAHMAATAILNLSTRCWEMPENLSTKQQDLPSKSMDIEVDENGTLDLSMNKHRKRESTFPSSSSCSSSPSVKSPDVSQRQNSTSATSSTMTSPQSSQTSRQDEWDGPIDYTKPNRQREEEPEESEPAAHSFASSEADEQEVSEENFEERKYPGEVTLTNFKLKFLPKDIKKELLTCPTPGCDGSGHITGNYASHRSLSGCPLADKSLRNLMAAHSADLKCPTPGCDGSGHITGNYASHRSLSGCPRAKKSGIKITPTKDDKEDPELMKCPVPGCVGLGHISGKYASHRSASGCPLAARRQKEGSLNGSSFSWKSLKNEGPTCPTPGCDGSGHANGSFLTHRSLSGCPRATFAGKKGKLSGDEILNTKFKTSDVLENDEEIKQLNKEITELNESNSEMEAAMVKLQSQISTMEKNLKNIEEENKIIEEQNEALFLELSGLSQALIQSLANIRLPHMEPISEQNFDAYVNTLTDMYTNQECYQNPENRDLLESIKQAVKGIQV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0R3</accession>
    <name>A0A452J0R3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">Myosin motor domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">Muscle contraction.</text>
    </comment>
    <comment type="subunit">
        <text evidence="14">Muscle myosin is a hexameric protein that consists of 2 heavy chain subunits (MHC), 2 alkali light chain subunits (MLC) and 2 regulatory light chain subunits (MLC-2).</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Myofibril</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 15">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038454.1">
        <property type="protein sequence ID" value="ENSGAGP00000033956.1"/>
        <property type="gene ID" value="ENSGAGG00000020688.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030016">
        <property type="term" value="C:myofibril"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032982">
        <property type="term" value="C:myosin filament"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016460">
        <property type="term" value="C:myosin II complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000146">
        <property type="term" value="F:microfilament motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000003">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000004">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000006">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000013">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000007">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000008">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.4820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.530:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000024">
        <property type="entry name" value="Myosin heavy chain, isoform J"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.720:FF:000001">
        <property type="entry name" value="Myosin heavy chain, muscle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.340">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.370">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.4820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.2420">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002928">
        <property type="entry name" value="Myosin_tail"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014751">
        <property type="entry name" value="XRCC4-like_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615">
        <property type="entry name" value="MYOSIN HEAVY CHAIN, NON-MUSCLE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615:SF69">
        <property type="entry name" value="MYOSIN-6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01576">
        <property type="entry name" value="Myosin_tail_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90257">
        <property type="entry name" value="Myosin rod fragments"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57997">
        <property type="entry name" value="Tropomyosin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12 15" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="7 15" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="8 16" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0488">Methylation</keyword>
    <keyword evidence="10 15" id="KW-0505">Motor protein</keyword>
    <keyword evidence="11" id="KW-0514">Muscle protein</keyword>
    <keyword evidence="9 15" id="KW-0518">Myosin</keyword>
    <keyword evidence="6 15" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0787">Thick filament</keyword>
    <feature type="domain" description="Myosin motor" evidence="17">
        <location>
            <begin position="83"/>
            <end position="775"/>
        </location>
    </feature>
    <feature type="region of interest" description="Actin-binding" evidence="15">
        <location>
            <begin position="652"/>
            <end position="674"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="16">
        <location>
            <begin position="837"/>
            <end position="1153"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="16">
        <location>
            <begin position="1197"/>
            <end position="1920"/>
        </location>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <begin position="176"/>
            <end position="183"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004657"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022433"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023179"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00037488"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00038612"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033956.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1935" mass="223630" checksum="C84F0197673341CF" modified="2019-05-08" version="1">LAKMMDMSEFGEAAPFLRKSDKEIMILQTVAFDGKTGYVFLCLVCSVERQGDSSCQQCYTCLSLQTVTVKEDDVQPMNPPKFDMIEDMAMLTHLNEASVLHNLNRRYSHWMIYTYSGLFCVTINPYKWLPVYNPEVVNAYKGKRRSEAPPHIFSIADNAYNDMLRNRENQSMLITGESGAGKTVNTKRVIQYFATVAALGEPGKKSVSMQGGTLEDQIIQANPALEAFGNAKTLRNDNSSRFGKFIRIHFGTTGKLSSADIEIYLLEKSRVIFQQPGERSYHIFYQILSGKKAELQDMLLVSTNPYDYHFSSQGAVTVDNLDDGEELMATDQAMDILGFIPDEKYGAYKLTGSIMHFGNMKFKQRPREEQAETDGTESTDKAAYLMGLNSSDLAKGLLHPRVKVGNEYVTKGQSVEQVVYAVGALSKAVYDRMFKWLVIRINKTLDTRLPRQFFIGVLDIAGFEIFDYNSFEQLCINYTNEKLQQFFNHHMFVLEQEEYKKEGIEWVFIDFGLDLQACIDLIEKPLGILSILEEECMFPKATDMTFKAKLYDNHLGKSPNLQKPRIDKKRKYEAHFELLHYAGAVPYNIIGWLEKNKDPLNETVVGIFQKSANKLLASLFENYVSADSGNTEKKRKKGASFQTVSSLHKENLNKLMTNLRSTAPHFVRCIIPNETKTPGAMDAYLVLHQLRCNGVLEGIRICRKGFPNRVLYADFKQRYRILNPAAIPEDKFVDSRKASEKLLASLDIDQTQYRFGHTKVFFKAGLLGHLEEMRDHRLSKILTMIQARARGKLMRIEFQKIVERRDALLVIQWNIRAFMAVKNWPWMKLFFKIKPLLKSAETEKEMANMKEEFLKLKEALEKSEARRKELEEKQVSLVQEKNDLLLQLQAEQDTLADAEERCDLLIKSKIQMEAKVKELTERVEDEEEMNSELTSKKRKLEDECSELKKDIDDLEITLAKVEKEKHATENKVKNLTEEMASLDETISKLTKEKKSLQEAHQQALDDLQAEEDKVNTLSKTKLKLEQQVDDLEGSLEQEKKIRMDLERVKRKLEGDLKLTQESLMDLENDKLQLDEKLKKKEFEINQLNSKLEDEQTIVMQLQKKIKELQARIEELEEELEAERAARAKVEKQRSDLARELEELSERLEEAGGATAAQLEMNKKREAEFLKVRRDLEEATLHYEATAATLRKKHADSMAELGEQVDNLQRVKQKLEKEKSELKLEVDDLLSNMEQMVKGKSNAEKLCRTYEDNLNETKTKLDEMTRLMNDLTTQKGKLQTENGEFARQLEEKESLISQLSRGKTSFTQQIEELRRLLEEETKSKNALAHALQAARHDCDLLREQFEEEQEAKAELQRALSKGNAEVAQWRTKYETDAIQRTEELEEAKKKLAARLQEAEEAIEAANAKCSSLEKTKHRLQNELEDIVIDLERANSAAAALDKKQRNFDKLISDWKQKYEETQAELEASQKEARSLSTELFKLKNAYEETLDNLETVKRENKNLQEEISDLTDQISEGNKSLHEIEKIKKQVEQEKSEVQLALEEAEGALEHEESKSLRFQLEISQIKAEFERKLAEKDEEMENIRRNQQRTIDSLQSTLDSEAKSRNEAIRLKKKMEGDLNEMEIQLSHANRHASEATKSARILQAQIKDLQVQLDDSEHLNEDLKEQLAVTDRRNNLIQSELDELRALLDQTERGRKLAEQELLEATERVNLLHTQNTSLINQKKKLETDISQMQNEVEEAIQECRNAEEKAKKAITDAAMMAEELKKEQDTSAHLERMKKNMEQTIKDLQKRLDEAEQIALKGGKKQIQKLESRVRELENDLENELRRNTESQKGVRKYERRIKELTYQAEEDKKNLTRMQELIDKLQLKVKSYKHQAEEAEAQSNQYLAKYRKQQHELDDAEERAEIAESQVNKLRTKTRDIGIKKVWCYSAE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0R4</accession>
    <name>A0A452J0R4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">heme oxygenase (biliverdin-producing)</fullName>
            <ecNumber evidence="2">1.14.14.18</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the heme oxygenase family.</text>
    </comment>
    <dbReference type="EC" id="1.14.14.18" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452J0R4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038491.1">
        <property type="protein sequence ID" value="ENSGAGP00000033991.1"/>
        <property type="gene ID" value="ENSGAGG00000024161.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0020037">
        <property type="term" value="F:heme binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004392">
        <property type="term" value="F:heme oxygenase (decyclizing) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042167">
        <property type="term" value="P:heme catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006788">
        <property type="term" value="P:heme oxidation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006979">
        <property type="term" value="P:response to oxidative stress"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd19165">
        <property type="entry name" value="HemeO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.910.10:FF:000001">
        <property type="entry name" value="Heme oxygenase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.910.10">
        <property type="entry name" value="Heme oxygenase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002051">
        <property type="entry name" value="Haem_Oase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016053">
        <property type="entry name" value="Haem_Oase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016084">
        <property type="entry name" value="Haem_Oase-like_multi-hlx"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018207">
        <property type="entry name" value="Haem_oxygenase_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10720">
        <property type="entry name" value="HEME OXYGENASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10720:SF2">
        <property type="entry name" value="HEME OXYGENASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01126">
        <property type="entry name" value="Heme_oxygenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00088">
        <property type="entry name" value="HAEMOXYGNASE"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48613">
        <property type="entry name" value="Heme oxygenase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00593">
        <property type="entry name" value="HEME_OXYGENASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0349">Heme</keyword>
    <keyword evidence="7" id="KW-0408">Iron</keyword>
    <keyword evidence="8" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="6" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="8" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="281"/>
            <end position="299"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00012360"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033991.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="302" mass="34461" checksum="007B936FBC8C7F55" modified="2019-05-08" version="1">MFPISCPFRISVLHSPTDLSEMLKEGTKESHDRAENTQFVKDFLKGRIKKELFKLATVALYYTYSALEEEMDCNKDHPAFAPLYFPLELHRKEALAKDMEYFYGENWKEKIQCSEATRCYVDRIHYIGQYEPELLVAHAYTRYMGDLSGGQILKKVAQRALKLPSTGEGIQFYVFDNISNAQQFKQFYRARINALDLDKKAKERIVEEANKAFEFNMQVFDELDKIGALLTEEAQDGGLPVHDGKGDLRKCPYYATKLGKGDPGCPYHIGMAMLKQPTMQFVLAACIALAAGIATWYVMDGH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0S0</accession>
    <name>A0A452J0S0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">tRNA (32-2'-O)-methyltransferase regulator THADA</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="3">Together with methyltransferase FTSJ1, methylates the 2'-O-ribose of nucleotides at position 32 of the anticodon loop of substrate tRNAs.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the THADA family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038498.1">
        <property type="protein sequence ID" value="ENSGAGP00000033998.1"/>
        <property type="gene ID" value="ENSGAGG00000024118.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030488">
        <property type="term" value="P:tRNA methylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056843">
        <property type="entry name" value="THADA-like_TPR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056842">
        <property type="entry name" value="THADA-like_TPR_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019442">
        <property type="entry name" value="THADA/TRM732_DUF2428"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051954">
        <property type="entry name" value="tRNA_methyltransferase_THADA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14387">
        <property type="entry name" value="THADA/DEATH RECEPTOR INTERACTING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14387:SF7">
        <property type="entry name" value="THYROID ADENOMA-ASSOCIATED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10350">
        <property type="entry name" value="DUF2428"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25150">
        <property type="entry name" value="TPR_Trm732"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25151">
        <property type="entry name" value="TPR_Trm732_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0819">tRNA processing</keyword>
    <feature type="domain" description="tRNA (32-2'-O)-methyltransferase regulator THADA-like TPR repeats region" evidence="6">
        <location>
            <begin position="665"/>
            <end position="781"/>
        </location>
    </feature>
    <feature type="domain" description="DUF2428" evidence="5">
        <location>
            <begin position="925"/>
            <end position="1225"/>
        </location>
    </feature>
    <feature type="domain" description="tRNA (32-2'-O)-methyltransferase regulator THADA-like C-terminal TPR repeats region" evidence="7">
        <location>
            <begin position="1227"/>
            <end position="1390"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00010409"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022694"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00035625"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00035698"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="Pfam" id="PF10350"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="Pfam" id="PF25150"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF25151"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033998.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1706" mass="192007" checksum="2A9C6FAB677B548D" modified="2019-05-08" version="1">MIEPRRGAEAVWVGGQKAAAEAGCRASVPLGLHFNHTKMVLKKKKEIQVDAFVLECQELETLRSFADAEDQNLASLLLHCVQLSDGIHQIYSMKQIVTLLKKMDRNSACSPMIKCCLDIIGVMYFSLGAKNPLKKMLASSLNGLPEQLMTEAVHSFVRCLKEELKTTDLCLYRKVMDNLASCMENFSLGKASINNLFKEVLQFLQKSVLEIQEENRKLSGNSIVETRLMHDLLMAIKVSMMLLQKVQENFQGSLWRKPGSLLWQSMCGLLKSFTNFLMNEGLLPTVQTTSGLAVILFTKTMFEPVEKLPCLVSDLLLRSVKCASVPEWFVSSCECLCTEELSDSVLLSLCHGALAMLEWKNGSMSENGEKLLLDIALVLLSLASRLKESNMAVSLSRILAIWTNSALDVLESSSQNLKSSLNGNSNTTGKLLEYVYTHWEHPLDAVRHQAKLIFKNILRIHQATLKGSVVKSDPFFSMLTESLLSLEWHVKGKYASLGCLVECVGVENILTVDRTIPAQILDVMGDQSLAPYASDLLETMFINHKSHLISAFQKNIWINHWHDTWVSPLLLILCEGNPDQTTYVIDYYLPKLLKCSPDSLSYMIKILQASADANIGSCNTRGALGALMACLRTARAHGHLQFREIMRDDLLSIGYVKQGLVHQHNQLFCRIQESSQVLYKLEQNKTKQELIKESAKWDHFATLQQYKDFMSSICDKLFEALFPGSSHPTRFSSLTILGSIAEIFSAPEGQTQAIFRLAQEIDPTRVQMLIQCFASTFEEVKVLAFELLMKLYDRLDLQDLENLHLLFQAAMDLSISTKPYDCVTASYLLNFLIHHKGLLNVCLKNQQVADTFQIDEDMSASTMEKNTLAVIKYLLSNLEREIFQAEKSLLQAAASFPMYGRVHCITGALQQLPLNNLTLMTEWRQMVTKLILMSYRLSSVVSPIVQSSSPEGLIPMDNDSETAGRLQMILHEIQPRDTNDYFTQAKIFKEHHRVDFAKLTDDEPVENVCTEIRGKEGQTCDVTAQMVLVCCWRSMKEISLLLGMLCKLLPLQTVSESSDGLIAVEQVKDIGDYFKHHLLQSRHRGAFELAYTGFVMLTEILSRCTNESLRTLPQQWLYNVLEEIKSCDPSSKLCATRRSAGIPFYIQALLASEPKNKTNLLKMTMNELVSLAVPVNESLNIIPQVHALNILRALFRDTRFGENIIPYIADGIQAAVLGFTSPVWAVRNSSTLLFSTLITRIFGVKRGKDENSKKNRMTGREFFSRFPTLYPFLLQQLEVVARTMDSETGELKLHPSLFLLLLILGKLYPSPMDGTYSALSMAPFVPFIMRCGHSPVYRSREMAGRALVPFVMISQIPHTVQSLLAGLPECTNPCIQQNSVHGMLLQVFHLLGSYLESKHRVNSDLQQGLVDIITCIKAKVWLAKRQNPCLVTRAAYLDVLVVLSDYLGKSRRKGIELFGFWEEISKVISASELMTGIYYFSAIPGFTQYLQSITKLVISVLSVAADPDFSNSSSPVAEQMMNSSSMVVHLLQSDFHEVRLPMLEAVLLWWKQVNSKQATEEKMGPLSLLSDLEVILLRMAMKEKHPECFCKVLEILHCMDLRNVLPRTQDSIKMNPKEFLNWAMNIISASDSIKIQSIALKLASKLVTHLVQNCQEVMESEMRQWIQLVTSCCGDEEQTAIRLAAAEVLINITPLFLTNQKLILGE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0S1</accession>
    <name>A0A452J0S1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">LisH domain and HEAT repeat-containing protein KIAA1468</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Golgi apparatus</location>
            <location evidence="2">trans-Golgi network</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Recycling endosome</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038460.1">
        <property type="protein sequence ID" value="ENSGAGP00000033962.1"/>
        <property type="gene ID" value="ENSGAGG00000023990.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055037">
        <property type="term" value="C:recycling endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005802">
        <property type="term" value="C:trans-Golgi network"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032367">
        <property type="term" value="P:intracellular cholesterol transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000080">
        <property type="entry name" value="lisH domain and HEAT repeat-containing protein KIAA1468 homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000218">
        <property type="entry name" value="lisH domain and HEAT repeat-containing protein KIAA1468 homolog isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021133">
        <property type="entry name" value="HEAT_type_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006594">
        <property type="entry name" value="LisH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040362">
        <property type="entry name" value="RELCH"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR32059">
        <property type="entry name" value="RAB11-BINDING PROTEIN RELCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR32059:SF0">
        <property type="entry name" value="RAB11-BINDING PROTEIN RELCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00667">
        <property type="entry name" value="LisH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50077">
        <property type="entry name" value="HEAT_REPEAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50896">
        <property type="entry name" value="LISH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7 11" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0967">Endosome</keyword>
    <keyword evidence="6" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="8" id="KW-0445">Lipid transport</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <feature type="repeat" description="HEAT" evidence="10">
        <location>
            <begin position="976"/>
            <end position="1012"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1"/>
            <end position="71"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="142"/>
            <end position="164"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="414"/>
            <end position="436"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="11">
        <location>
            <begin position="187"/>
            <end position="221"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="11">
        <location>
            <begin position="337"/>
            <end position="364"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="12">
        <location>
            <begin position="46"/>
            <end position="56"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="142"/>
            <end position="152"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="416"/>
            <end position="436"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004172"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004601"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00084053"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00103"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033962.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1186" mass="131675" checksum="851A640D9007EC94" modified="2019-05-08" version="1">AARPALVPALRGWRGAATAGAGGVAGPASSVNPFLSDSEEEEPRDGGGGNDPGLGGPLSPQQHEEEEEAAVEVPGVLPGLRAAAPGEAGWAGPGAGEPPRVPVDALAAQLLRDQFLLTALELHTELLESGRELPRLRDYFSNPGNFERQSGTPPAGGAPGLWGAADRAGSISTLDSLDFARYSDDGNRETDERVAVLEFELRKAKETIQALRANLTQAAENEVPLQERKNYKSSPEIQEPIKPLEKRALNFLVNEFLLKNSYKLTSITFSDESDDQDFELWDDVGLNIPKPPDLLQLYRDFGNHQVTARDVVDVAVGVEEDELEVTTPTLGSQFLIVQKLEDQINALNSEKWSLTEQIRRLERYNENFAVPVVCDLVHVPLSVPEDNGSYLDIRSSDNDSKHGKTEEISLSISAEADSKTPKDDTLNSLPSKDREKLPSCALSNKAAIHFDKPNRTLSPAFHQALLSFCRMSAESRLGSEVSRIADSEKSVMLMLGRCLPHIVPNVLLAKREELIPLILCTACLHPEPKERDQLLHILFNLIKRPDDEQRQMILTGCVAFARHVGPTRVEAELLPQCWEQINHKYPERRLLVAESCGALAPYLPKEIRSSLVLSMLQQMLMEDKADLVREAVIKSLGIIMGYIDDPDKYQQGFELLLSALGDPSERVVSATHQVFLPAYAAWTTELGNLQLHLIPTLLNKIEKLLREGEHGLDEHKLHMYLSALQSLIPSLFSLVLQNAPFTSKAKLQGEVPPIEVTRFPRPVSPLQDVATIIGSREQLAVLLQLYDYQLELEGTTGWETLLWVVNQLLPQLIEIVGKINVASTACVHEFSRFFWRLCRTFGKIFTNTKVKPQFQEILRLSEENIDSTAGNGVLTKATVPIYATGVLTCYIQEEDRKLLIGFLEDVMTMLSLSHAPLNSLKASFVELGANPAYHELLLTVLWYGVVHTSALVRCTAARMFELLVKGVHETLVAQRVVPALITYSSDPEISVRIATIPAFGTIMETVTQREXRVKMQLASFLEDPQYQDQHSLQTEIIKTFGRVGPNAEPRFRDEFVIPHLHKLALVNNQQSVDSKRLDIATHLFEAYSALSCCFISEDLMVNHFLPGLRCLRTDMEHLSPEHEVILSSMIKECEQKVENKTVQEPQGSMSIAASLVSEDTKTKFLNKMGQLTTSGAMLANVFQRKK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0S2</accession>
    <name>A0A452J0S2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Fork-head domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 6">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0S2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038464.1">
        <property type="protein sequence ID" value="ENSGAGP00000033966.1"/>
        <property type="gene ID" value="ENSGAGG00000024149.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000038466.1">
        <property type="protein sequence ID" value="ENSGAGP00000033968.1"/>
        <property type="gene ID" value="ENSGAGG00000024149.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd20023">
        <property type="entry name" value="FH_FOXJ1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.10:FF:000030">
        <property type="entry name" value="Forkhead box protein K2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.10">
        <property type="entry name" value="Winged helix-like DNA-binding domain superfamily/Winged helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047512">
        <property type="entry name" value="FH_FOXJ1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001766">
        <property type="entry name" value="Fork_head_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047513">
        <property type="entry name" value="FOXJ1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR030456">
        <property type="entry name" value="TF_fork_head_CS_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036388">
        <property type="entry name" value="WH-like_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036390">
        <property type="entry name" value="WH_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46805">
        <property type="entry name" value="FORKHEAD BOX PROTEIN J1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46805:SF3">
        <property type="entry name" value="FORKHEAD BOX PROTEIN J1-B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00250">
        <property type="entry name" value="Forkhead"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00053">
        <property type="entry name" value="FORKHEAD"/>
    </dbReference>
    <dbReference type="SMART" id="SM00339">
        <property type="entry name" value="FH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46785">
        <property type="entry name" value="Winged helix' DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00658">
        <property type="entry name" value="FORK_HEAD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50039">
        <property type="entry name" value="FORK_HEAD_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3 6" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="5 6" id="KW-0539">Nucleus</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0804">Transcription</keyword>
    <keyword evidence="2" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="Fork-head" evidence="8">
        <location>
            <begin position="127"/>
            <end position="221"/>
        </location>
    </feature>
    <feature type="DNA-binding region" description="Fork-head" evidence="6">
        <location>
            <begin position="127"/>
            <end position="221"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="54"/>
            <end position="108"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="62"/>
            <end position="73"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="98"/>
            <end position="107"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50039"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033966.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="429" mass="47131" checksum="AF804EFFA7D37B6C" modified="2019-05-08" version="1">MPILISPDIAARFKEKWLLLHPEDQDNVGGAVTLDDSLTSLQWLQDFSIVTTDPERPPVPSCPLQQPPQQLFQGSEAPASPPAGDTAAKGMPPRLGKPTSTAISPGTSYLPGLGATEIDYKSNPRVKPPYSYATLICLAVRASKQAKVTLSAIYSWIMENFCYYRHAEPSWQNSIRHNLSLNKCFRKVPRQKDEPGKGGFWEIDPQYADMFSDGVVKRRRPAAAPYSPPWQRPAVPALEPKCSLLSPVPHLGDGQPGHTPHQATCCLVHQHSQRCPLLDAPSPLPKRSCPAARTAQSPLLPSSGRDAEALRGEFDWADAFDDVFRGNGSNFEDLDINAALSSLSTEIDLAMQGRYIPPAGKWGAPAPSHPSPGASHWEDFTPFADTPQHPWEEAKGEALSNPWGFEQGFNFCDGFLSEMQPWDKVDTFM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0S5</accession>
    <name>A0A452J0S5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">E3 ubiquitin-protein ligase</fullName>
            <ecNumber evidence="20">2.3.2.26</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="27 28">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="28" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="27" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="27" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1 20">
            <text>S-ubiquitinyl-[E2 ubiquitin-conjugating enzyme]-L-cysteine + [acceptor protein]-L-lysine = [E2 ubiquitin-conjugating enzyme]-L-cysteine + N(6)-ubiquitinyl-[acceptor protein]-L-lysine.</text>
            <dbReference type="EC" id="2.3.2.26"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="5 20">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="subunit">
        <text evidence="19">(Microbial infection) Interacts with Epstein-Barr virus LMP2A.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Peripheral membrane protein</topology>
            <orientation evidence="2">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Early endosome membrane</location>
            <topology evidence="3">Peripheral membrane protein</topology>
            <orientation evidence="3">Cytoplasmic side</orientation>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="2.3.2.26" evidence="20"/>
    <dbReference type="AlphaFoldDB" id="A0A452J0S5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038505.1">
        <property type="protein sequence ID" value="ENSGAGP00000034005.1"/>
        <property type="gene ID" value="ENSGAGG00000024168.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00143"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031901">
        <property type="term" value="C:early endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061630">
        <property type="term" value="F:ubiquitin protein ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006915">
        <property type="term" value="P:apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051607">
        <property type="term" value="P:defense response to virus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045087">
        <property type="term" value="P:innate immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050687">
        <property type="term" value="P:negative regulation of defense response to virus"/>
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        <property type="project" value="UniProtKB-ARBA"/>
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    <dbReference type="GO" id="GO:0002683">
        <property type="term" value="P:negative regulation of immune system process"/>
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        <property type="project" value="UniProtKB-ARBA"/>
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    <dbReference type="GO" id="GO:1902532">
        <property type="term" value="P:negative regulation of intracellular signal transduction"/>
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        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051241">
        <property type="term" value="P:negative regulation of multicellular organismal process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009896">
        <property type="term" value="P:positive regulation of catabolic process"/>
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        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002684">
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010604">
        <property type="term" value="P:positive regulation of macromolecule metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043161">
        <property type="term" value="P:proteasome-mediated ubiquitin-dependent protein catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035519">
        <property type="term" value="P:protein K29-linked ubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070936">
        <property type="term" value="P:protein K48-linked ubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070534">
        <property type="term" value="P:protein K63-linked ubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd04021">
        <property type="entry name" value="C2_E3_ubiquitin_ligase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00078">
        <property type="entry name" value="HECTc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00201">
        <property type="entry name" value="WW"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.70.10:FF:000005">
        <property type="entry name" value="E3 ubiquitin-protein ligase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.70.10:FF:000009">
        <property type="entry name" value="E3 ubiquitin-protein ligase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.70.10:FF:000039">
        <property type="entry name" value="E3 ubiquitin-protein ligase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000092">
        <property type="entry name" value="E3 ubiquitin-protein ligase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1750.10:FF:000002">
        <property type="entry name" value="E3 ubiquitin-protein ligase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.2410.10:FF:000002">
        <property type="entry name" value="E3 ubiquitin-protein ligase HECW2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.70.10:FF:000063">
        <property type="entry name" value="E3 ubiquitin-protein ligase NEDD4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.20.70.10">
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.2160.10">
        <property type="entry name" value="Hect, E3 ligase catalytic domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.2410.10">
        <property type="entry name" value="Hect, E3 ligase catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1750.10">
        <property type="entry name" value="Hect, E3 ligase catalytic domains"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000008">
        <property type="entry name" value="C2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024928">
        <property type="entry name" value="E3_ub_ligase_SMURF1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050409">
        <property type="entry name" value="E3_ubiq-protein_ligase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000569">
        <property type="entry name" value="HECT_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035983">
        <property type="entry name" value="Hect_E3_ubiquitin_ligase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001202">
        <property type="entry name" value="WW_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036020">
        <property type="entry name" value="WW_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11254:SF66">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE ITCHY HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11254">
        <property type="entry name" value="HECT DOMAIN UBIQUITIN-PROTEIN LIGASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00168">
        <property type="entry name" value="C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00632">
        <property type="entry name" value="HECT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00397">
        <property type="entry name" value="WW"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF001569">
        <property type="entry name" value="E3_ub_ligase_SMURF1"/>
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    </dbReference>
    <dbReference type="SMART" id="SM00239">
        <property type="entry name" value="C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00119">
        <property type="entry name" value="HECTc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00456">
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    </dbReference>
    <dbReference type="SUPFAM" id="SSF49562">
        <property type="entry name" value="C2 domain (Calcium/lipid-binding domain, CaLB)"/>
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    </dbReference>
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        <property type="entry name" value="Hect, E3 ligase catalytic domain"/>
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    </dbReference>
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        <property type="entry name" value="WW domain"/>
        <property type="match status" value="4"/>
    </dbReference>
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        <property type="entry name" value="C2"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="WW_DOMAIN_1"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50020">
        <property type="entry name" value="WW_DOMAIN_2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="16" id="KW-0007">Acetylation</keyword>
    <keyword evidence="17" id="KW-0051">Antiviral defense</keyword>
    <keyword evidence="11" id="KW-0053">Apoptosis</keyword>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="13" id="KW-0967">Endosome</keyword>
    <keyword evidence="15" id="KW-0391">Immunity</keyword>
    <keyword evidence="9" id="KW-0399">Innate immunity</keyword>
    <keyword evidence="18" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="28" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="12" id="KW-0677">Repeat</keyword>
    <keyword evidence="10 20" id="KW-0808">Transferase</keyword>
    <keyword evidence="14 20 22" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="C2" evidence="24">
        <location>
            <begin position="1"/>
            <end position="117"/>
        </location>
    </feature>
    <feature type="domain" description="WW" evidence="25">
        <location>
            <begin position="303"/>
            <end position="336"/>
        </location>
    </feature>
    <feature type="domain" description="WW" evidence="25">
        <location>
            <begin position="335"/>
            <end position="368"/>
        </location>
    </feature>
    <feature type="domain" description="WW" evidence="25">
        <location>
            <begin position="414"/>
            <end position="447"/>
        </location>
    </feature>
    <feature type="domain" description="WW" evidence="25">
        <location>
            <begin position="454"/>
            <end position="487"/>
        </location>
    </feature>
    <feature type="domain" description="HECT" evidence="26">
        <location>
            <begin position="545"/>
            <end position="832"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="23">
        <location>
            <begin position="149"/>
            <end position="311"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="23">
        <location>
            <begin position="152"/>
            <end position="165"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="23">
        <location>
            <begin position="187"/>
            <end position="201"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="23">
        <location>
            <begin position="209"/>
            <end position="223"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="23">
        <location>
            <begin position="234"/>
            <end position="270"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="23">
        <location>
            <begin position="277"/>
            <end position="289"/>
        </location>
    </feature>
    <feature type="active site" description="Glycyl thioester intermediate" evidence="21 22">
        <location>
            <position position="800"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000885"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004413"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004469"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022588"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022703"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022859"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023118"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00064638"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="PIRNR" id="PIRNR001569"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="PIRSR" id="PIRSR001569-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00104"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS50004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS50020"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="26">
        <source>
            <dbReference type="PROSITE" id="PS50237"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034005.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="28">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="832" mass="95080" checksum="BF966EFE47215C14" modified="2019-05-08" version="1">MAGSGPRSGTVSGFAMKSQLQITVISAKLKENKNKWFGPSPYVEVSVDGQSKKTEKCNNTNSPKWKQHLTVIVTPVSKLNFRVWSHQTLKSDILLGSATLEVHETLKSNNMKLEQVVVTLQLVGDREPAEVIGDLSVCLDGMQVDPELLTNGDATSIRSPTQNDGSRARNDTRANANGLENYEDAGSSESRTVNGSDSPLLTNGVFKPARPPRPSRPPPPTPRRPSSAAGCVNGPSSMSIENDESSAGISPSTTPNAPSEQSPEGATAGTTVPAASALTTPPVSRQVQPVNPPSQPHSTVSQGPLPPGWEQRVDQHGRVYYVDHVEKRTTWDRPEPLPPGWERRVDNMGRIYYVDHFTRTTTWQRPTLESVRNFEQWQLQRSQLQGAMQQFNQRFIYGNQDVTAAQNAEFDPLGTLPHGWEKRTDSNGRVYFVNHNTRITQWEDPRNQGQLNEKPLPEGWEMRFTVDGIPYFVDHNRRTTTYIDPRTGKSALDNGPQIAYVRDFKAKVQYFRFWCQQLAMPQHTKITVSRKTLFEDSFQQIMSFSPQDLRRRLWVIFPGEEGLDYGGVAREWFFLLSHEVLNPMYCLFEYVGKDNYCLQINPASYINPDHLKYFRFIGRFIAMALFHVRFIDTGFSLPFYKRILNKPTGLKDLESVDPEFYNSLIWVKLVAEWRLSRGVEEQTQAFFEGFNEILPQQYLQYFDAKELEVLLCGMQEIDLTDWQRHTIYRHYTRTSKQIMWFWQFVKEIDNEKRMRLLQFVTGTCRLPVGGFADLMGSNGPQKFCIEKVGKENWLPRSHTCFNRLDLPPYKSYEQLKEKLLFAIEETEGFGQE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0S6</accession>
    <name>A0A452J0S6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="22">Myosin heavy chain 6</fullName>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Muscle contraction.</text>
    </comment>
    <comment type="subunit">
        <text evidence="15">Muscle myosin is a hexameric protein that consists of 2 heavy chain subunits (MHC), 2 alkali light chain subunits (MLC) and 2 regulatory light chain subunits (MLC-2).</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Myofibril</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 16">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038465.1">
        <property type="protein sequence ID" value="ENSGAGP00000033967.1"/>
        <property type="gene ID" value="ENSGAGG00000020688.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030016">
        <property type="term" value="C:myofibril"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032982">
        <property type="term" value="C:myosin filament"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016460">
        <property type="term" value="C:myosin II complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000146">
        <property type="term" value="F:microfilament motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000003">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000004">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000006">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000013">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000007">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000008">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.4820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.530:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.360:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.720:FF:000001">
        <property type="entry name" value="Myosin heavy chain, muscle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000101">
        <property type="entry name" value="Slow myosin heavy chain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.340">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.370">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.4820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.2420">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.360">
        <property type="entry name" value="Myosin S1 fragment, N-terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008989">
        <property type="entry name" value="Myosin_S1_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002928">
        <property type="entry name" value="Myosin_tail"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004009">
        <property type="entry name" value="SH3_Myosin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014751">
        <property type="entry name" value="XRCC4-like_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615">
        <property type="entry name" value="MYOSIN HEAVY CHAIN, NON-MUSCLE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615:SF69">
        <property type="entry name" value="MYOSIN-6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02736">
        <property type="entry name" value="Myosin_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01576">
        <property type="entry name" value="Myosin_tail_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90257">
        <property type="entry name" value="Myosin rod fragments"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57997">
        <property type="entry name" value="Tropomyosin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51844">
        <property type="entry name" value="SH3_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="13 16" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="7 16" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="8" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="9 17" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0488">Methylation</keyword>
    <keyword evidence="11 16" id="KW-0505">Motor protein</keyword>
    <keyword evidence="12" id="KW-0514">Muscle protein</keyword>
    <keyword evidence="10 16" id="KW-0518">Myosin</keyword>
    <keyword evidence="6 16" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0787">Thick filament</keyword>
    <feature type="domain" description="Myosin N-terminal SH3-like" evidence="19">
        <location>
            <begin position="30"/>
            <end position="79"/>
        </location>
    </feature>
    <feature type="domain" description="Myosin motor" evidence="18">
        <location>
            <begin position="83"/>
            <end position="771"/>
        </location>
    </feature>
    <feature type="region of interest" description="Actin-binding" evidence="16">
        <location>
            <begin position="648"/>
            <end position="670"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="17">
        <location>
            <begin position="833"/>
            <end position="1149"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="17">
        <location>
            <begin position="1193"/>
            <end position="1916"/>
        </location>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <begin position="168"/>
            <end position="175"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004657"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022433"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023179"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00037488"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00038612"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS51844"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033967.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="22">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1931" mass="223156" checksum="6DEDAF3F8905E079" modified="2019-05-08" version="1">MMDMSEFGEAAPFLRKSDKEIMILQTVAFDGKRRCWIPDEKKAYTEGDIKESSGGKVTVETVDGKTVTVKEDDVQPMNPPKFDMIEDMAMLTHLNEASVLHNLNRRYSHWMIYTYSGLFCVTINPYKWLPVYNPEVVNAYKGKRRSEAPPHIFSIADNAYNDMLRSNGESGAGKTVNTKRVIQYFATVAALGEPGKKSQPATRTGGTLEDQIIQANPALEAFGNAKTLRNDNSSRFGKFIRIHFGTTGKLSSADIEIYLLEKSRVIFQQPGERSYHIFYQILSGKKAELQDMLLVSTNPYDYHFSSQGAVTVDNLDDGEELMATDQAMDILGFIPDEKYGAYKLTGSIMHFGNMKFKQRPREEQAETDGTESTDKAAYLMGLNSSDLAKGLLHPRVKVGNEYVTKGQSVEQVVYAVGALSKAVYDRMFKWLVIRINKTLDTRLPRQFFIGVLDIAGFEIFDYNSFEQLCINYTNEKLQQFFNHHMFVLEQEEYKKEGIEWVFIDFGLDLQACIDLIEKPLGILSILEEECMFPKATDMTFKAKLYDNHLGKSPNLQKPRIDKKRKYEAHFELLHYAGAVPYNIIGWLEKNKDPLNETVVGIFQKSANKLLASLFENYVSADSVDHSTEKKRKKGASFQTVSSLHKENLNKLMTNLRSTAPHFVRCIIPNETKTPGAMDAYLVLHQLRCNGVLEGIRICRKGFPNRVLYADFKQRYRILNPAAIPEDKFVDSRKASEKLLASLDIDQTQYRFGHTKVFFKAGLLGHLEEMRDHRLSKILTMIQARARGKLMRIEFQKIVERRDALLVIQWNIRAFMAVKNWPWMKLFFKIKPLLKSAETEKEMANMKEEFLKLKEALEKSEARRKELEEKQVSLVQEKNDLLLQLQAEQDTLADAEERCDLLIKSKIQMEAKVKELTERVEDEEEMNSELTSKKRKLEDECSELKKDIDDLEITLAKVEKEKHATENKVKNLTEEMASLDETISKLTKEKKSLQEAHQQALDDLQAEEDKVNTLSKTKLKLEQQVDDLEGSLEQEKKIRMDLERVKRKLEGDLKLTQESLMDLENDKLQLDEKLKKKEFEINQLNSKLEDEQTIVMQLQKKIKELQARIEELEEELEAERAARAKVEKQRSDLARELEELSERLEEAGGATAAQLEMNKKREAEFLKVRRDLEEATLHYEATAATLRKKHADSMAELGEQVDNLQRVKQKLEKEKSELKLEVDDLLSNMEQMVKGKSNAEKLCRTYEDNLNETKTKLDEMTRLMNDLTTQKGKLQTENGEFARQLEEKESLISQLSRGKTSFTQQIEELRRLLEEETKSKNALAHALQAARHDCDLLREQFEEEQEAKAELQRALSKGNAEVAQWRTKYETDAIQRTEELEEAKKKLAARLQEAEEAIEAANAKCSSLEKTKHRLQNELEDIVIDLERANSAAAALDKKQRNFDKLISDWKQKYEETQAELEASQKEARSLSTELFKLKNAYEETLDNLETVKRENKNLQEEISDLTDQISEGNKSLHEIEKIKKQVEQEKSEVQLALEEAEGALEHEESKSLRFQLEISQIKAEFERKLAEKDEEMENIRRNQQRTIDSLQSTLDSEAKSRNEAIRLKKKMEGDLNEMEIQLSHANRHASEATKSARILQAQIKDLQVQLDDSEHLNEDLKEQLAVTDRRNNLIQSELDELRALLDQTERGRKLAEQELLEATERVNLLHTQNTSLINQKKKLETDISQMQNEVEEAIQECRNAEEKAKKAITDAAMMAEELKKEQDTSAHLERMKKNMEQTIKDLQKRLDEAEQIALKGGKKQIQKLESRVRELENDLENELRRNTESQKGVRKYERRIKELTYQAEEDKKNLTRMQELIDKLQLKVKSYKHQAEEAEAQSNQYLAKYRKQQHELDDAEERAEIAESQVNKLRTKTRDIGIKKVWCYSAE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J111</accession>
    <name>A0A452J111_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">RAR related orphan receptor B</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J111"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038556.1">
        <property type="protein sequence ID" value="ENSGAGP00000034049.1"/>
        <property type="gene ID" value="ENSGAGG00000024178.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004879">
        <property type="term" value="F:nuclear receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd06968">
        <property type="entry name" value="NR_DBD_ROR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.50.10:FF:000003">
        <property type="entry name" value="Nuclear orphan receptor ROR-beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.50.10">
        <property type="entry name" value="Erythroid Transcription Factor GATA-1, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.565.10">
        <property type="entry name" value="Retinoid X Receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035500">
        <property type="entry name" value="NHR-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044101">
        <property type="entry name" value="NR_DBD_ROR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000536">
        <property type="entry name" value="Nucl_hrmn_rcpt_lig-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001723">
        <property type="entry name" value="Nuclear_hrmn_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003079">
        <property type="entry name" value="ROR_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001628">
        <property type="entry name" value="Znf_hrmn_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013088">
        <property type="entry name" value="Znf_NHR/GATA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45805">
        <property type="entry name" value="NUCLEAR HORMONE RECEPTOR HR3-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45805:SF7">
        <property type="entry name" value="NUCLEAR RECEPTOR ROR-BETA-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00104">
        <property type="entry name" value="Hormone_recep"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00105">
        <property type="entry name" value="zf-C4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01293">
        <property type="entry name" value="RORNUCRECPTR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00398">
        <property type="entry name" value="STRDHORMONER"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00047">
        <property type="entry name" value="STROIDFINGER"/>
    </dbReference>
    <dbReference type="SMART" id="SM00430">
        <property type="entry name" value="HOLI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00399">
        <property type="entry name" value="ZnF_C4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57716">
        <property type="entry name" value="Glucocorticoid receptor-like (DNA-binding domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48508">
        <property type="entry name" value="Nuclear receptor ligand-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51843">
        <property type="entry name" value="NR_LBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51030">
        <property type="entry name" value="NUCLEAR_REC_DBD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="7" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="9" id="KW-0675">Receptor</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0804">Transcription</keyword>
    <keyword evidence="6" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <keyword evidence="4" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="Nuclear receptor" evidence="12">
        <location>
            <begin position="3"/>
            <end position="78"/>
        </location>
    </feature>
    <feature type="domain" description="NR LBD" evidence="13">
        <location>
            <begin position="211"/>
            <end position="449"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="88"/>
            <end position="108"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS51030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS51843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034049.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="16">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="459" mass="52205" checksum="094142BF62944AB7" modified="2019-05-08" version="1">MRPTSQKICGDKSSGIHYGVITCEGCKGFFRRSQQNNASYSCSRQRNCLIDRTNRNRCQHCRLQKCLALGMSRDAVKFGRMSKKQRDSLYAEVQKHQQNQEQNRVSKDDSEVLARVYPTSVSSGVSDLDDISLLSDGLLFDFPLTPDGGSTYYNLDLLTSAQPSPDQSSMDVSDAKFIKQESIYELMLEPGLFAHSTLEGSQLASDISMLEIDRIAQNVVKSHLETCQYTTEELKRLAWMLYTQEEIRSLQNKSCETMWQQCSLQISNAIQYVVEFAKRIDGFMELCQNDQIILLKAGCLEVLLIRMIRAFNPLNNMVLFEGKYGGMQMFKSLGCDDLINAVFELGRNLCRLQLSDEEIALFTAAVLLSPDRPWLTESKKVQKLQDKIYIALQHEIQKKHSSEDKLSKMISKLPLMKTICNLHLDKLEFFRLLHPETAMNFPPLYKEAPGSVTGLSYSP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J114</accession>
    <name>A0A452J114_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">LisH domain and HEAT repeat-containing protein KIAA1468</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Golgi apparatus</location>
            <location evidence="2">trans-Golgi network</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Recycling endosome</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038513.1">
        <property type="protein sequence ID" value="ENSGAGP00000034013.1"/>
        <property type="gene ID" value="ENSGAGG00000023990.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055037">
        <property type="term" value="C:recycling endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005802">
        <property type="term" value="C:trans-Golgi network"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032367">
        <property type="term" value="P:intracellular cholesterol transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000080">
        <property type="entry name" value="lisH domain and HEAT repeat-containing protein KIAA1468 homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000218">
        <property type="entry name" value="lisH domain and HEAT repeat-containing protein KIAA1468 homolog isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021133">
        <property type="entry name" value="HEAT_type_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006594">
        <property type="entry name" value="LisH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040362">
        <property type="entry name" value="RELCH"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR32059">
        <property type="entry name" value="RAB11-BINDING PROTEIN RELCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR32059:SF0">
        <property type="entry name" value="RAB11-BINDING PROTEIN RELCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00667">
        <property type="entry name" value="LisH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50077">
        <property type="entry name" value="HEAT_REPEAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50896">
        <property type="entry name" value="LISH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7 11" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0967">Endosome</keyword>
    <keyword evidence="6" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="8" id="KW-0445">Lipid transport</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <feature type="repeat" description="HEAT" evidence="10">
        <location>
            <begin position="985"/>
            <end position="1021"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1"/>
            <end position="76"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="121"/>
            <end position="164"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="420"/>
            <end position="445"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="11">
        <location>
            <begin position="180"/>
            <end position="214"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="11">
        <location>
            <begin position="341"/>
            <end position="375"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="12">
        <location>
            <begin position="25"/>
            <end position="35"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="130"/>
            <end position="155"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="425"/>
            <end position="445"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004172"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004601"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00084053"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00103"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034013.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1195" mass="132432" checksum="750AE905FD69C2B3" modified="2019-05-08" version="1">MAAAGGGSSVNPFLSDSEEEEPRDGGGGNDPGLGGPLSPQQHEEEEEAAVEVPGVLPGLRAAAPGEAGWAGPGAGEPPRVPVDALAAQLLRDQFLLTALELHTELLESGRELPRLRDYFSNPGNFERQSGTPPAGGAPGLWGAAGAAPGGAAASGKEPDRAGSISTLDSLDFARYSDDGNRETDERVAVLEFELRKAKETIQALRANLTQAAENEVPLQERKNYKSSPEIQEPIKPLEKRALNFLVNEFLLKNSYKLTSITFSDESDDQDFELWDDVGLNIPKPPDLLQLYRDFGNHQVTARDVVDVAVGVEEDELEVTTPTLGSVPVFGVSQSTEQFLIVQKLEDQINALNSEKWSLTEQIRRLESEIDFLKNENFAVPVVCDLVHVPLSVPEDNGSYLDIRSSDNDSKHGKTEEISLSISAEADSKTPKDDTLNSLPSKDREKLPSCALSNKAAIHFDKPNRTLSPAFHQALLSFCRMSAESRLGSEVSRIADSEKSVMLMLGRCLPHIVPNVLLAKREELIPLILCTACLHPEPKERDQLLHILFNLIKRPDDEQRQMILTGCVAFARHVGPTRVEAELLPQCWEQINHKYPERRLLVAESCGALAPYLPKEIRSSLVLSMLQQMLMEDKADLVREAVIKSLGIIMGYIDDPDKYQQGFELLLSALGDPSERVVSATHQVFLPAYAAWTTELGNLQLHLIPTLLNKIEKLLREGEHGLDEHKLHMYLSALQSLIPSLFSLVLQNAPFTSKAKLQGEVPPIEVTRFPRPVSPLQDVATIIGSREQLAVLLQLYDYQLELEGTTGWETLLWVVNQLLPQLIEIVGKINVASTACVHEFSRFFWRLCRTFGKIFTNTKVKPQFQEILRLSEENIDSTAGNGVLTKATVPIYATGVLTCYIQEEDRKLLIGFLEDVMTMLSLSHAPLNSLKASFVELGANPAYHELLLTVLWYGVVHTSALVRCTAARMFELLVKGVHETLVAQRVVPALITYSSDPEISVRIATIPAFGTIMETVTQREXRVKMQLASFLEDPQYQDQHSLQTEIIKTFGRVGPNAEPRFRDEFVIPHLHKLALVNNQQSVDSKRLDIATHLFEAYSALSCCFISEDLMVNHFLPGLRCLRTDMEHLSPEHEVILSSMIKECEQKVENKTVQEPQGSMSIAASLVSEDTKTKFLNKMGQLTTSGAMLANVFQRKK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J118</accession>
    <name>A0A452J118_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Inactive rhomboid protein</fullName>
            <shortName evidence="11">iRhom</shortName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="11">Rhomboid family member</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="11">Rhomboid veinlet-like protein</fullName>
        </alternativeName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1 11">Regulates ADAM17 protease, a sheddase of the epidermal growth factor (EGF) receptor ligands and TNF, thereby plays a role in sleep, cell survival, proliferation, migration and inflammation. Does not exhibit any protease activity on its own.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2 11">Endoplasmic reticulum membrane</location>
            <topology evidence="2 11">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Golgi apparatus membrane</location>
            <topology evidence="3">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4 11">Belongs to the peptidase S54 family.</text>
    </comment>
    <comment type="caution">
        <text evidence="11">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J118"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038599.1">
        <property type="protein sequence ID" value="ENSGAGP00000034087.1"/>
        <property type="gene ID" value="ENSGAGG00000024222.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000139">
        <property type="term" value="C:Golgi membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050709">
        <property type="term" value="P:negative regulation of protein secretion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042058">
        <property type="term" value="P:regulation of epidermal growth factor receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1540.10:FF:000001">
        <property type="entry name" value="Putative inactive rhomboid protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1540.10">
        <property type="entry name" value="Rhomboid-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051512">
        <property type="entry name" value="Inactive_Rhomboid"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022241">
        <property type="entry name" value="iRhom1_2_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022764">
        <property type="entry name" value="Peptidase_S54_rhomboid_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035952">
        <property type="entry name" value="Rhomboid-like_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45965">
        <property type="entry name" value="INACTIVE RHOMBOID PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45965:SF4">
        <property type="entry name" value="INACTIVE RHOMBOID PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12595">
        <property type="entry name" value="iRhom1-2_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01694">
        <property type="entry name" value="Rhomboid"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF144091">
        <property type="entry name" value="Rhomboid-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6 11" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="8" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="9 11" id="KW-0472">Membrane</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 11" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 11" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="656"/>
            <end position="680"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="692"/>
            <end position="711"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="717"/>
            <end position="735"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="747"/>
            <end position="767"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="773"/>
            <end position="792"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="804"/>
            <end position="825"/>
        </location>
    </feature>
    <feature type="domain" description="Inactive rhomboid protein 1/2 N-terminal" evidence="13">
        <location>
            <begin position="92"/>
            <end position="285"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase S54 rhomboid" evidence="12">
        <location>
            <begin position="651"/>
            <end position="787"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00002661"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004477"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004653"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00009045"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="RuleBase" id="RU369051"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF01694"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="Pfam" id="PF12595"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034087.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="855" mass="97711" checksum="6097E377B7889158" modified="2019-05-08" version="1">MSEARHDSTSSLQRKKPPWLKLDIPVPVSVAMPEEPTFVQPFKRQAFLRSVSMPAENTRIPSPPNEVRRPMLQRQTSITQTIRRGTADWFGVSKDSDATQKWQRKSLRHCSQRYGKLKPQVIREMDLPSQDNISLTSTETPPPLYVGPCQLGMQKIVDPLARGRAFRMADDNDGYSIPHTPITPGATSLCSFTSSRSGFNRLPRRRKRESVAKMSFRAAAALMKGRSVKDSTLRRAQRRSFTPASFLEEDTVDFPDELDTSFFAREGVLHEELSTYPDEVFESPSEAAIKDSEVKPQDQADLTGGALDKNELERSHLMLPLERGWRKQKEGTPPQPKVRLRQEVVSVSPQRRGQRIAVPVRKLFAKEKRPYGLGMVGKLTNRTYRKRIDSYVKRQIEDMDDHRPFFTYWVTFVHSLITILAVCIYGIAPVGFSQHETVDSVLMNRGIYENVKYVQQENFWIGPSSEALIHLGAKFSPCMRQDQQVHNFISAMREKEKHSACCVRNDKSGCVQTSEEECSSTLAVWVKWPHHSSTPTLAGGKRQFGSVCHQDPRVCEQPASMDPHEWPDDITKWPICTKNSAGNYTNHLHMDCVITGRPCCIGTKGRCEITSREYCDFMRGYFHEEATLCSQVHCMDDVCGLLPFLNPEIPDQFYRLWLSLFLHAGILHCLVSVCFQMTILRDLEKLAGWHRISIIYLLSGITGNLASAIFLPYRAEVGPAGSQFGILACLFVELFQSWQILARPWRAFFKLLAVVLFLFAFGLLPWIDNFAHISGFISGLFLSFAFLPYISFGKFDLYRKRCQIIVFQLIFIGLFSGLVILFYFYPIKCEWCEFLTCIPFTDKFCEKYDLDAQLH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J121</accession>
    <name>A0A452J121_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Oxysterol-binding protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="5">Belongs to the OSBP family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J121"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038579.1">
        <property type="protein sequence ID" value="ENSGAGP00000034068.1"/>
        <property type="gene ID" value="ENSGAGG00000024195.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097038">
        <property type="term" value="C:perinuclear endoplasmic reticulum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015485">
        <property type="term" value="F:cholesterol binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006869">
        <property type="term" value="P:lipid transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd13285">
        <property type="entry name" value="PH_ORP1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000098">
        <property type="entry name" value="Oxysterol-binding protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000231">
        <property type="entry name" value="Oxysterol-binding protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.160.120:FF:000005">
        <property type="entry name" value="Oxysterol-binding protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.160.120">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.3490">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037239">
        <property type="entry name" value="OSBP_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000648">
        <property type="entry name" value="Oxysterol-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018494">
        <property type="entry name" value="Oxysterol-bd_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10972">
        <property type="entry name" value="OXYSTEROL-BINDING PROTEIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10972:SF53">
        <property type="entry name" value="OXYSTEROL-BINDING PROTEIN-RELATED PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00023">
        <property type="entry name" value="Ank"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01237">
        <property type="entry name" value="Oxysterol_BP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00233">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF144000">
        <property type="entry name" value="Oxysterol-binding protein-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01013">
        <property type="entry name" value="OSBP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="2 6" id="KW-0445">Lipid transport</keyword>
    <keyword evidence="3" id="KW-0446">Lipid-binding</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1 6" id="KW-0813">Transport</keyword>
    <feature type="repeat" description="ANK" evidence="4">
        <location>
            <begin position="41"/>
            <end position="73"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="4">
        <location>
            <begin position="136"/>
            <end position="168"/>
        </location>
    </feature>
    <feature type="domain" description="PH" evidence="9">
        <location>
            <begin position="198"/>
            <end position="297"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="765"/>
            <end position="792"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="396"/>
            <end position="423"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="849"/>
            <end position="883"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023121"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="RuleBase" id="RU003844"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="RuleBase" id="RU003845"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034068.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="963" mass="109112" checksum="0F3179A326E2B4B1" modified="2019-05-08" version="1">MMLEMETVKRLNNYWKPWTKEKWLLTSTAKAGAEVNVLNDMGDTPLHRAAFTGRKEVVMLLLRYDADTSIVNGGGQTARDVTQDKEIKNMLEAVERTQERKLEELLLGAAREGETAKLSALLNRPKPPDINCTDQMGNTPLHCAAYRAHKHCALKLLKKGADSKIRNKNDQTPFDLAQGAEIKQILAGSIGKVMNKRLKRYEGLLWKSLRFFGWKLYWVVLEHGVLSWYRKQADAVNNDHRQGCKHLTQAVCTVKSTDSCLFSVKCFDDSVHSFKVPKNSPPKARENWLEAIEDHSAYSTHYCTQEQLSDEEDDDTVSVAELQETLKKAQTCQQRLDKEISNFLAMIKSCSTTKGMPPSFLQKIEVVSEVSKETCEALGNCLKLFTKQEGVRNLKLEHEQEKNKILSEALETLATEHHELEQSLVKGSPPLSILSEEQFYDAVSDSESEKSLSGFETVTAYSLEDSVESNNTITSSMSEEKVCGGGGKVCGGGESLSNGIKKHRTSLPSPMFSRNDFSIWSILRKCIGMELSKITMPVIFNEPLSFLQRLTEYMEHTYLIHQASSLSGTVERMQCVAAFAVSAVASQWERTGKPFNPLLGETYELVRDDLGFRLLSEQVSHHPPISAFYAEGLNNDFVFHGSIYPKLKFWGKSVEAEPKGTITLELLEHDEAYTWTNPTCCVHNIIVGKLWIEQYGNVEITNHKTGEKCVLNFKPCGLFGKELHKVEGYIQDKSKRKVCSLYGKWTECLYSVDPATFEAYKKNDKKNTEEKKNSKQVGNSEEPDEMPLPDSESVHVIPGSILLWRIAPRPPNSVQMYNFTSFAMALNELDKEMESVVPKTDCRLRPDIRAMENGEIDIASEEKKRLEEKQRAVRKNRSKSEEDWKTRSVHLIKFMAVQIRCLSTHRLFSITRRVDDHSCSAITILFEILIRVFVMPGLVRNVQGSLQHQPCFIFLLNIPLPHL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J127</accession>
    <name>A0A452J127_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Myosin motor domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1 9">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <comment type="caution">
        <text evidence="9">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J127"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038523.1">
        <property type="protein sequence ID" value="ENSGAGP00000034023.1"/>
        <property type="gene ID" value="ENSGAGG00000024156.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016459">
        <property type="term" value="C:myosin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000146">
        <property type="term" value="F:microfilament motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007015">
        <property type="term" value="P:actin filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.530:FF:000002">
        <property type="entry name" value="Class V myosin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000089">
        <property type="entry name" value="Myosin VC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13140">
        <property type="entry name" value="MYOSIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13140:SF313">
        <property type="entry name" value="UNCONVENTIONAL MYOSIN-VC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 9" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="4 9" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="5" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7 9" id="KW-0505">Motor protein</keyword>
    <keyword evidence="6 9" id="KW-0518">Myosin</keyword>
    <keyword evidence="3 9" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="Myosin motor" evidence="10">
        <location>
            <begin position="63"/>
            <end position="594"/>
        </location>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <begin position="157"/>
            <end position="164"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00008314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034023.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="594" mass="68636" checksum="943E4E38004E6044" modified="2019-05-08" version="1">MHNNTCRKVWIPDNEEVWQSAEITKSYRTGDRVLHLLLEDGTELNYPVDPAPLPPLRNPDILVGENDLTALSYLHEPAVLHNLKVRFVESKLIYTYSGIILVAMNPYKQLPIYGDAIIHAYSGQNMGDMDPHIFAVAEEAYKQMARNNRNQSVIVSGESGAGKTVSARYAMRYFATVSKSNSNVHVEDKVLASNPITEAVGNAKTTRNDNSSRFGKYTEISFDQRHQITGANMRTYLLEKSRVVFQSENERNYHIFYQLCASALQPEFKHLKLGHAEEFNYTRMGGNTVIEGVDDRADMMETQKTFTLLGLKEDFQMDVFKILAAILHLGNVNVYFSSLKLEDKHLNIFCELLDLDNDKVAQWLCHRKIITTSETVIKPMTKLQALNARDALAKKIYSHLFDFIVERINKALHFSGKQHTFIGVLDIYGFETFDVNSFEQFCINYANEKLQQQFNLHVFKLEQEEYMKENIPWTLIDFYDNQPVIDLIEAKMGILELLDEECLLPQGTDENWLQKLYNNYINKNPFFQKPKMSNTSFIIQHFADKVEYKSEGFLEKNRDTVYEVLIDILRNSKVSKFIMANYSTRDTKILVEYE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J129</accession>
    <name>A0A452J129_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">D-amino-acid oxidase</fullName>
            <ecNumber evidence="17">1.4.3.3</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="33 34">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="34" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="33" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="25">
            <text>D-alanine + O2 + H2O = pyruvate + H2O2 + NH4(+)</text>
            <dbReference type="Rhea" id="RHEA:22688"/>
            <dbReference type="ChEBI" id="CHEBI:15361"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15379"/>
            <dbReference type="ChEBI" id="CHEBI:16240"/>
            <dbReference type="ChEBI" id="CHEBI:28938"/>
            <dbReference type="ChEBI" id="CHEBI:57416"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="25">
            <dbReference type="Rhea" id="RHEA:22689"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="27">
            <text>D-cysteine + O2 + H2O = 2-oxo-3-sulfanylpropanoate + H2O2 + NH4(+)</text>
            <dbReference type="Rhea" id="RHEA:78791"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15379"/>
            <dbReference type="ChEBI" id="CHEBI:16240"/>
            <dbReference type="ChEBI" id="CHEBI:28938"/>
            <dbReference type="ChEBI" id="CHEBI:35236"/>
            <dbReference type="ChEBI" id="CHEBI:57678"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="27">
            <dbReference type="Rhea" id="RHEA:78792"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="21">
            <text>D-dopa + O2 + H2O = 3-(3,4-dihydroxyphenyl)pyruvate + H2O2 + NH4(+)</text>
            <dbReference type="Rhea" id="RHEA:70971"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15379"/>
            <dbReference type="ChEBI" id="CHEBI:16240"/>
            <dbReference type="ChEBI" id="CHEBI:28938"/>
            <dbReference type="ChEBI" id="CHEBI:29055"/>
            <dbReference type="ChEBI" id="CHEBI:149689"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="21">
            <dbReference type="Rhea" id="RHEA:70972"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="29">
            <text>D-leucine + O2 + H2O = 4-methyl-2-oxopentanoate + H2O2 + NH4(+)</text>
            <dbReference type="Rhea" id="RHEA:78211"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15379"/>
            <dbReference type="ChEBI" id="CHEBI:16240"/>
            <dbReference type="ChEBI" id="CHEBI:17865"/>
            <dbReference type="ChEBI" id="CHEBI:28938"/>
            <dbReference type="ChEBI" id="CHEBI:143079"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="29">
            <dbReference type="Rhea" id="RHEA:78212"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="26">
            <text>D-lysine + O2 + H2O = 6-amino-2-oxohexanoate + H2O2 + NH4(+)</text>
            <dbReference type="Rhea" id="RHEA:37583"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15379"/>
            <dbReference type="ChEBI" id="CHEBI:16240"/>
            <dbReference type="ChEBI" id="CHEBI:28938"/>
            <dbReference type="ChEBI" id="CHEBI:32557"/>
            <dbReference type="ChEBI" id="CHEBI:58183"/>
            <dbReference type="EC" id="1.4.3.3"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="26">
            <dbReference type="Rhea" id="RHEA:37584"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="23">
            <text>D-methionine + O2 + H2O = 4-methylsulfanyl-2-oxobutanoate + H2O2 + NH4(+)</text>
            <dbReference type="Rhea" id="RHEA:78207"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15379"/>
            <dbReference type="ChEBI" id="CHEBI:16240"/>
            <dbReference type="ChEBI" id="CHEBI:16723"/>
            <dbReference type="ChEBI" id="CHEBI:28938"/>
            <dbReference type="ChEBI" id="CHEBI:57932"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="23">
            <dbReference type="Rhea" id="RHEA:78208"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="22">
            <text>D-phenylalanine + O2 + H2O = 3-phenylpyruvate + H2O2 + NH4(+)</text>
            <dbReference type="Rhea" id="RHEA:70963"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15379"/>
            <dbReference type="ChEBI" id="CHEBI:16240"/>
            <dbReference type="ChEBI" id="CHEBI:18005"/>
            <dbReference type="ChEBI" id="CHEBI:28938"/>
            <dbReference type="ChEBI" id="CHEBI:57981"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="22">
            <dbReference type="Rhea" id="RHEA:70964"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>D-proline + O2 = 1-pyrroline-2-carboxylate + H2O2</text>
            <dbReference type="Rhea" id="RHEA:78259"/>
            <dbReference type="ChEBI" id="CHEBI:15379"/>
            <dbReference type="ChEBI" id="CHEBI:16240"/>
            <dbReference type="ChEBI" id="CHEBI:39785"/>
            <dbReference type="ChEBI" id="CHEBI:57726"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="19">
            <dbReference type="Rhea" id="RHEA:78260"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="24">
            <text>D-serine + O2 + H2O = 3-hydroxypyruvate + H2O2 + NH4(+)</text>
            <dbReference type="Rhea" id="RHEA:70951"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15379"/>
            <dbReference type="ChEBI" id="CHEBI:16240"/>
            <dbReference type="ChEBI" id="CHEBI:17180"/>
            <dbReference type="ChEBI" id="CHEBI:28938"/>
            <dbReference type="ChEBI" id="CHEBI:35247"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="24">
            <dbReference type="Rhea" id="RHEA:70952"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="20">
            <text>D-tryptophan + O2 + H2O = indole-3-pyruvate + H2O2 + NH4(+)</text>
            <dbReference type="Rhea" id="RHEA:78247"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15379"/>
            <dbReference type="ChEBI" id="CHEBI:16240"/>
            <dbReference type="ChEBI" id="CHEBI:17640"/>
            <dbReference type="ChEBI" id="CHEBI:28938"/>
            <dbReference type="ChEBI" id="CHEBI:57719"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="20">
            <dbReference type="Rhea" id="RHEA:78248"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="28">
            <text>D-valine + O2 + H2O = 3-methyl-2-oxobutanoate + H2O2 + NH4(+)</text>
            <dbReference type="Rhea" id="RHEA:78203"/>
            <dbReference type="ChEBI" id="CHEBI:11851"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15379"/>
            <dbReference type="ChEBI" id="CHEBI:16240"/>
            <dbReference type="ChEBI" id="CHEBI:28938"/>
            <dbReference type="ChEBI" id="CHEBI:74338"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="28">
            <dbReference type="Rhea" id="RHEA:78204"/>
        </physiologicalReaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 30">
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Cytosol</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Peroxisome matrix</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="16">Presynaptic active zone</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the DAMOX/DASOX family.</text>
    </comment>
    <dbReference type="EC" id="1.4.3.3" evidence="17"/>
    <dbReference type="AlphaFoldDB" id="A0A452J129"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038609.1">
        <property type="protein sequence ID" value="ENSGAGP00000034097.1"/>
        <property type="gene ID" value="ENSGAGG00000024232.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000038611.1">
        <property type="protein sequence ID" value="ENSGAGP00000034099.1"/>
        <property type="gene ID" value="ENSGAGG00000024232.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000038612.1">
        <property type="protein sequence ID" value="ENSGAGP00000034100.1"/>
        <property type="gene ID" value="ENSGAGG00000024232.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005782">
        <property type="term" value="C:peroxisomal matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048786">
        <property type="term" value="C:presynaptic active zone"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003884">
        <property type="term" value="F:D-amino-acid oxidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071949">
        <property type="term" value="F:FAD binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055130">
        <property type="term" value="P:D-alanine catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036088">
        <property type="term" value="P:D-serine catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006562">
        <property type="term" value="P:L-proline catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.9.10:FF:000004">
        <property type="entry name" value="D-amino-acid oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.9.10">
        <property type="entry name" value="D-Amino Acid Oxidase, subunit A, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.720">
        <property type="entry name" value="NAD(P)-binding Rossmann-like Domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006181">
        <property type="entry name" value="D-amino_acid_oxidase_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023209">
        <property type="entry name" value="DAO"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006076">
        <property type="entry name" value="FAD-dep_OxRdtase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11530">
        <property type="entry name" value="D-AMINO ACID OXIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11530:SF15">
        <property type="entry name" value="D-AMINO-ACID OXIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01266">
        <property type="entry name" value="DAO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000189">
        <property type="entry name" value="D-aa_oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54373">
        <property type="entry name" value="FAD-linked reductases, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51971">
        <property type="entry name" value="Nucleotide-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00677">
        <property type="entry name" value="DAO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="15" id="KW-0966">Cell projection</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11 30" id="KW-0274">FAD</keyword>
    <keyword evidence="9" id="KW-0285">Flavoprotein</keyword>
    <keyword evidence="12" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="14" id="KW-0576">Peroxisome</keyword>
    <keyword evidence="8" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="34" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0702">S-nitrosylation</keyword>
    <keyword evidence="7" id="KW-0964">Secreted</keyword>
    <keyword evidence="31" id="KW-0732">Signal</keyword>
    <keyword evidence="13" id="KW-0770">Synapse</keyword>
    <feature type="signal peptide" evidence="31">
        <location>
            <begin position="1"/>
            <end position="16"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5044604608" description="D-amino-acid oxidase" evidence="31">
        <location>
            <begin position="17"/>
            <end position="347"/>
        </location>
    </feature>
    <feature type="domain" description="FAD dependent oxidoreductase" evidence="32">
        <location>
            <begin position="2"/>
            <end position="328"/>
        </location>
    </feature>
    <feature type="binding site" evidence="30">
        <location>
            <begin position="37"/>
            <end position="38"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="30">
        <location>
            <begin position="44"/>
            <end position="45"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="30">
        <location>
            <begin position="49"/>
            <end position="51"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="30">
        <location>
            <position position="53"/>
        </location>
        <ligand>
            <name>D-dopa</name>
            <dbReference type="ChEBI" id="CHEBI:149689"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="30">
        <location>
            <position position="182"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="30">
        <location>
            <position position="217"/>
        </location>
        <ligand>
            <name>D-dopa</name>
            <dbReference type="ChEBI" id="CHEBI:149689"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="30">
        <location>
            <position position="228"/>
        </location>
        <ligand>
            <name>D-serine</name>
            <dbReference type="ChEBI" id="CHEBI:35247"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="30">
        <location>
            <position position="283"/>
        </location>
        <ligand>
            <name>D-serine</name>
            <dbReference type="ChEBI" id="CHEBI:35247"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="30">
        <location>
            <begin position="312"/>
            <end position="317"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="30">
        <location>
            <position position="313"/>
        </location>
        <ligand>
            <name>D-serine</name>
            <dbReference type="ChEBI" id="CHEBI:35247"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001974"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004253"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004514"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00006730"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022630"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022799"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022827"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023140"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00034101"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00039101"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00039751"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00044716"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00047579"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00048079"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00048252"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00048401"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00048643"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00048711"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00048747"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="ARBA" id="ARBA00049123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="ARBA" id="ARBA00049182"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="ARBA" id="ARBA00049287"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="PIRSR" id="PIRSR000189-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="31">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="32">
        <source>
            <dbReference type="Pfam" id="PF01266"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="33">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034097.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="34">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="347" mass="39321" checksum="5681FE5DE4B52FEA" modified="2019-05-08" version="1">MRVAVIGAGVIGLSTALCIYDQHHSVVQPLEIEVYADRYTPLTTSDGAAGLWQPYVDDKRNAQETLWNKETFDYLCGHLNTPEAEEMGLFPISGYNLFTQPVPDPPWKDIVLGFRNLTAKELELFPGYSYGWFNTALILEGKSYLPWLTKRLKQRRVKFFHKKIESFHEMAAKGVEVIINCTGVRAGELQPDPGLMPGRGQIIKVLAPWVKHFIMTHDLESGIYNSPYIVPGSKLVTLGGIFQLGNWNEENSPQDHKTIWENCCKLLPSLQKAKIVDEWSGFRPVRYTVRLEREAIRHGPLKSEVIHNYGHGGFGLTIHWGCAMEAARIFGRILEEKKLARPPQSHL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J130</accession>
    <name>A0A452J130_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">ARID domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038591.1">
        <property type="protein sequence ID" value="ENSGAGP00000034079.1"/>
        <property type="gene ID" value="ENSGAGG00000024184.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035060">
        <property type="term" value="C:brahma complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
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    <dbReference type="GO" id="GO:0071565">
        <property type="term" value="C:nBAF complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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    <dbReference type="GO" id="GO:0005654">
        <property type="term" value="C:nucleoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
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    <dbReference type="GO" id="GO:0016514">
        <property type="term" value="C:SWI/SNF complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
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    <dbReference type="GO" id="GO:0031491">
        <property type="term" value="F:nucleosome binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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    <dbReference type="GO" id="GO:0006338">
        <property type="term" value="P:chromatin remodeling"/>
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        <property type="project" value="InterPro"/>
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    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045893">
        <property type="term" value="P:positive regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd16877">
        <property type="entry name" value="ARID_ARID1B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.60:FF:000002">
        <property type="entry name" value="AT-rich interactive domain-containing protein 1B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.60">
        <property type="entry name" value="ARID DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038040">
        <property type="entry name" value="ARID_ARID1B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001606">
        <property type="entry name" value="ARID_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036431">
        <property type="entry name" value="ARID_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021906">
        <property type="entry name" value="BAF250/Osa"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033388">
        <property type="entry name" value="BAF250_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12656:SF11">
        <property type="entry name" value="AT-RICH INTERACTIVE DOMAIN-CONTAINING PROTEIN 1B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12656">
        <property type="entry name" value="BRG-1 ASSOCIATED FACTOR 250 BAF250"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01388">
        <property type="entry name" value="ARID"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12031">
        <property type="entry name" value="BAF250_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01014">
        <property type="entry name" value="ARID"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00501">
        <property type="entry name" value="BRIGHT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46774">
        <property type="entry name" value="ARID-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51011">
        <property type="entry name" value="ARID"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0007">Acetylation</keyword>
    <keyword evidence="4" id="KW-0156">Chromatin regulator</keyword>
    <keyword evidence="8" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="12" id="KW-0472">Membrane</keyword>
    <keyword evidence="2" id="KW-0488">Methylation</keyword>
    <keyword evidence="5" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="12" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="12" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="83"/>
            <end position="106"/>
        </location>
    </feature>
    <feature type="domain" description="ARID" evidence="13">
        <location>
            <begin position="518"/>
            <end position="609"/>
        </location>
    </feature>
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        <location>
            <begin position="1"/>
            <end position="20"/>
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            <begin position="34"/>
            <end position="53"/>
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            <begin position="964"/>
            <end position="1006"/>
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            <begin position="1059"/>
            <end position="1131"/>
        </location>
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    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="128"/>
            <end position="142"/>
        </location>
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    <feature type="compositionally biased region" description="Polar residues" evidence="11">
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            <begin position="197"/>
            <end position="209"/>
        </location>
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        <location>
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            <end position="221"/>
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        </location>
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        <location>
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            <end position="290"/>
        </location>
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            <begin position="358"/>
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            <begin position="429"/>
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        <location>
            <begin position="1114"/>
            <end position="1125"/>
        </location>
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        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
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        <source>
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            <dbReference type="ARBA" id="ARBA00022902"/>
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    </evidence>
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            <dbReference type="ARBA" id="ARBA00022990"/>
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    </evidence>
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            <dbReference type="ARBA" id="ARBA00023015"/>
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    </evidence>
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            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00023242"/>
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    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="PROSITE" id="PS51011"/>
        </source>
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            <dbReference type="Ensembl" id="ENSGAGP00000034079.1"/>
        </source>
    </evidence>
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            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1473" mass="158853" checksum="507D1520A3A5E388" modified="2019-05-08" version="1">YDRERGSPQLSLPAAAPGLVGRDASPAMAALGLRLQDRRPSPSRSGLGLGGGAPAEPGAKLPCPGCAWVCAPQPWQVWAAAQLGLGFASAVPWVLVLTHTLFFFFFSQDLSGSIDDLPTGTEATLSSAVSASGSTSSQGEQSNPAQSPFSPHASPHLSGIPGGPSPSPVGSPVGSNQSRSGPISPASIPGFMAGIQRNPQISQYGPQQTGPSMSPHPSPGGQMHPGIGSFQQSNSSGTYGPQMSQYGPQGNYSRPSTYSGVANTSYSGPGPGMGINANSQIHGQGPSQPCGTMPLGRMPSAGMQSRPFPGNMGSMTPNSPGMSLQGGPGMGPPMPTVNRKAQEAAAAVMQAAANSAQSRQGSFPGMNQSGIMGSSSPYTQPMNNSSGLMNPQAPPYSMAPNMVNSSTAPMGLADMMTPGESKLPLPLKADGKEEGPSQPESKSKDSYSSQGISQPPTPGNLPVPSPMSPSSASISSFHGDESDSISSPGWPKTPSSPKSSSSTTTGEKITKVYELGSEPERKMWVDRYLTFMEERGTPVASLPAVGKKPLDLFRLYVCVKEIGGLAQVNKNKKWRELATNLNVGTSSSAASSLKKQYIQYLFAFECKIERGEEPPPEVFSTGDTKKQAKIQPPSPANSGSLQGPQTPQSTGSNSMTEVPGDLKPPTPASTPHGQMTPMQGGRNSAVSVHDPFSDVSDSAFPKRNSMTPNAPYQQGMNMPDMMGRMPYEPNKDPFGGIRKVPGSSEPFMAPGQMPNSGIQDIYNQTPSGAMSTMSMSQRQQFSYGGGYDRRHEPYGQQYPGQGPPSGQPPYGGHQPGMYPQQQVKEVESFIFMTFILXQPVLKPRRKITSKDIVTPEAWRVMMSLKSGLLAESTWALDTINILLYDDSTVATFNLSQLSGFLELLVEYFRKCLIDIFGILMEYEVGDPGHKALDHNAVKKDDTQSLAVDTGKEEENDECIDYFDEEEEDEEEEEEDDIEKIEGEEKSTVFATPSAIADPSERPKQASKFDKLPIKIVKKNNLFVIDRSDKLGRVQEFNSGLLHWQLGGGDTTEHIQTHFESKMEIPPRRRAPPPLNSPSKKKDLEGKGEPEEQQEKSITATIDDVLSARPGALSEDSNPGSQTESSKFPFGIHQAKSHRNIKLLEDEPRSRDETPLCTITHWQDSLAKRCICVSNIVRSLSFVPGNDTEMSKHPGLVLILGKLILLHHEHPERKRAPQTYEKEEEEDKGVACSKDEWWWDCLEVLRDNTLVTLANISGQLDLSAYTESICLPILDGLLHWMVCPSAEAQDPFPTVGPNSVLSPQRLVLETLCKLSIQDNNVDLILATPPFSRQEKLYATLVRYVGDRKNPVCREMSMALLSNLAQGDMLAARAIAVQKGSIGNLISFLEDGVTMAQYQQSQHNLMHMQPPPLEPPSVDMMCRAAKALLAMARVDENRSEFLLHEGRLLDISISSVLNSLVASVICDVLFQIGQL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J131</accession>
    <name>A0A452J131_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Ubiquitin carboxyl-terminal hydrolase</fullName>
            <ecNumber evidence="7">3.4.19.12</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1 7">
            <text>Thiol-dependent hydrolysis of ester, thioester, amide, peptide and isopeptide bonds formed by the C-terminal Gly of ubiquitin (a 76-residue protein attached to proteins as an intracellular targeting signal).</text>
            <dbReference type="EC" id="3.4.19.12"/>
        </reaction>
    </comment>
    <comment type="similarity">
        <text evidence="2 7">Belongs to the peptidase C19 family.</text>
    </comment>
    <dbReference type="EC" id="3.4.19.12" evidence="7"/>
    <dbReference type="AlphaFoldDB" id="A0A452J131"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038600.1">
        <property type="protein sequence ID" value="ENSGAGP00000034088.1"/>
        <property type="gene ID" value="ENSGAGG00000024225.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004843">
        <property type="term" value="F:cysteine-type deubiquitinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016579">
        <property type="term" value="P:protein deubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd02662">
        <property type="entry name" value="Peptidase_C19F"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.70.10">
        <property type="entry name" value="Cysteine proteinases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038765">
        <property type="entry name" value="Papain-like_cys_pep_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050164">
        <property type="entry name" value="Peptidase_C19"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001394">
        <property type="entry name" value="Peptidase_C19_UCH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028889">
        <property type="entry name" value="USP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018200">
        <property type="entry name" value="USP_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24006">
        <property type="entry name" value="UBIQUITIN CARBOXYL-TERMINAL HYDROLASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24006:SF888">
        <property type="entry name" value="UBIQUITIN CARBOXYL-TERMINAL HYDROLASE 30"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00443">
        <property type="entry name" value="UCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54001">
        <property type="entry name" value="Cysteine proteinases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00972">
        <property type="entry name" value="USP_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00973">
        <property type="entry name" value="USP_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50235">
        <property type="entry name" value="USP_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 7" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="8" id="KW-0472">Membrane</keyword>
    <keyword evidence="3 7" id="KW-0645">Protease</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6 7" id="KW-0788">Thiol protease</keyword>
    <keyword evidence="8" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="4 7" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="6"/>
            <end position="25"/>
        </location>
    </feature>
    <feature type="domain" description="USP" evidence="9">
        <location>
            <begin position="37"/>
            <end position="466"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000707"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009085"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022807"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU366025"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50235"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034088.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="481" mass="54703" checksum="3F741686B9497417" modified="2019-05-08" version="1">MKNWGVVGGIAAALAAGVYVLWGPITERKKRRKGLVPGLLNLGNTCFMNSLLQGLSACPSFIKWLEDFTAQYKTDQNESSEHQYLSLTLLCLLKALSCQEVTEDDVLDASCLLEVLRLYRWQISSFEEQDAHELFHVLTSSLEDERERQPRITHLFDVHSLEQPEITQKQISCRTTGSLHPMSSHWKSQHPFHGRLTSNMLCKHCKRQSPVRCDTFDSLSLSIPAAMWGRPMTLDHCLHHFISSESVKDVVCDNCTKIQAEGTLNGQTTENQRTTFVKQLKLGKLPQCLCIHLQRLSWSDQGMPLKHHEHVQFNEFLIMDIYKYHLPVHKSSQHDLNQKGNPRMLSETKDGSAVKASDLEQTCSTKSFFMNGACSPSLLTSPITFPLAVVSDCSSPVYLYRLMAVVVHHGDMHSGHFVTYRRSPPSPKTTLSVSNQWLWISDDTVRKASLQEVLSSSAYLLFYERVHSRMQHQTQDFRAEE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J132</accession>
    <name>A0A452J132_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Rho guanine nucleotide exchange factor 4</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038528.1">
        <property type="protein sequence ID" value="ENSGAGP00000034028.1"/>
        <property type="gene ID" value="ENSGAGG00000024175.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035556">
        <property type="term" value="P:intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd01224">
        <property type="entry name" value="PH_Collybistin_ASEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00160">
        <property type="entry name" value="RhoGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd11973">
        <property type="entry name" value="SH3_ASEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.900.10:FF:000002">
        <property type="entry name" value="Rho guanine nucleotide exchange factor 9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000015">
        <property type="entry name" value="Rho guanine nucleotide exchange factor 9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.900.10">
        <property type="entry name" value="Dbl homology (DH) domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035899">
        <property type="entry name" value="DBL_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000219">
        <property type="entry name" value="DH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001331">
        <property type="entry name" value="GDS_CDC24_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055251">
        <property type="entry name" value="SOS1_NGEF_PH"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47544">
        <property type="entry name" value="RHO GUANINE NUCLEOTIDE EXCHANGE FACTOR 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47544:SF2">
        <property type="entry name" value="RHO GUANINE NUCLEOTIDE EXCHANGE FACTOR 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00621">
        <property type="entry name" value="RhoGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00018">
        <property type="entry name" value="SH3_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22697">
        <property type="entry name" value="SOS1_NGEF_PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00233">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00325">
        <property type="entry name" value="RhoGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00326">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48065">
        <property type="entry name" value="DBL homology domain (DH-domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00741">
        <property type="entry name" value="DH_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50010">
        <property type="entry name" value="DH_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0344">Guanine-nucleotide releasing factor</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 5" id="KW-0728">SH3 domain</keyword>
    <feature type="domain" description="SH3" evidence="7">
        <location>
            <begin position="1376"/>
            <end position="1435"/>
        </location>
    </feature>
    <feature type="domain" description="DH" evidence="9">
        <location>
            <begin position="1472"/>
            <end position="1656"/>
        </location>
    </feature>
    <feature type="domain" description="PH" evidence="8">
        <location>
            <begin position="1687"/>
            <end position="1794"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="36"/>
            <end position="66"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="175"/>
            <end position="197"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="442"/>
            <end position="470"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="932"/>
            <end position="967"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="1087"/>
            <end position="1110"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="1131"/>
            <end position="1172"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="1251"/>
            <end position="1293"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="47"/>
            <end position="63"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="185"/>
            <end position="196"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="447"/>
            <end position="457"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="932"/>
            <end position="941"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="958"/>
            <end position="967"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="1135"/>
            <end position="1160"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="1251"/>
            <end position="1266"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022658"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50010"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034028.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="12">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1837" mass="204956" checksum="E6E3A98167AA0F2F" modified="2019-05-08" version="1">MSQLELSQKAQLELSSDLKVAPSRSEFWIQTTKLRREEAGEEELILSSDPSNTGSGTSGGDTISKTETKLVVDGLQSEAKSENNKVQSCVSSEPAEDAIARSKKSPAGLSIACLKANSESCLTFLETQQFPLQPNVGSSHSLHNINFETNRTDSKRNSEAFEFHYELDSRSNIKCSRRNSDNNEETTATKTRNRTPGATRKMDSLLYQPCLKSKFQLPHMRAEIKESKTWHSAPENINAGGQRDYLGNCIQLRDKLRLSCSRIHLEGLGFEYTWKNIERVGEGSEKAIKVEGKFGIKGSLLNTCLESNNSQLSHPAGSQQDAEANLHTESETCGLNIEADMCADGLSSAQLQLRDTFEWDSIERETPSALASEPMSINAEGEVRKAKCKSERGFKAHGFSHSINSESGISEDQATCLVGVPAIAENKGDNYLANYGPAGKTQLNSLEESSPESPSTEACVGERGSEDSSELNLDWKSLETALNSGGQDIAGEQDVSMLQNLNCKNNALQGHSVLVTMNGHFADAGKKDESDGVDACSVEIATKPHYMHQGKKLMKLSELKPALIIISVEEKGLQSKMLSDDCPAEIITGTLGELANEDLGSHPIASKGTDESQGDLNAYSSCEEQTLPVQPISGNEEVTAAGITGQEESEIVQQQVERTEPDLSLKLSCSDFPVTTIKNESSGVNLHLEDAIVPLKEKADACTGAHLKSPGIISGGISQVEPCRTNAGVQSETELLKKSLFETRDSKVTKENEFNTNQSLEQDTCGTIATALEKNRISGLCQGDDADGHTFGSIISCVHSLEKKKTESYNISKDQIGIGEDSSSRELKLSSTVEELVSLSKDNEQNPNVTDAFNCIVKTESQEASALGLDNISTSSANSEEMDENSFHFLGSNSSLYKAVQKQTKTGKPSKFLVFSKMASFKKTKTAAAENQGSSNFFGSKTKTEELDAGKEDEDTESLQSFKSSSPSSAFRRKESYTSEYSDDDDLFYERPAGLFSRISLRKASGSGRILMDNIDTNFTSPSLHVKHSEARVLESVENNDSEVPQYFGVRKSLSENEYKGNKNPEGKKFRTRLALAHKSFSSFFESKSLEKENPEQSPKGSLKHEKEKAKLRQSSWKAFLKSREADGLKKPTLASLSSSQESPKTTRGYQQGPGNVTRRTSTEKQESSNGQAFFSSYESNATLMDANHSDTSVESIEVXNMRRKRVLSYDLTIKCPEYPDYAAQQETMVNSLDTSEEYWLRSPFSPSDLQSPFTQLSPSCPQLSTYEGKDMPCRPMSPKPQSPRPSSQRKSFRYPGRISATSMISLGNSSAIYSNLEAPERPKTLKPRGTLLLSVHSLDNEYLREDSGISSQSQISLNTASSVSDILRDELISDGSVVYAEALWDHVTMDDQELGFKAGDVIEVMDATNKEWWWGRILDSEGWFPASFVRLRVNQDEPMEDYPMKAEDGKEEDASIAARRYGVGQISKDQMRTNVINEIISTERDYIKHLKDICEGYIKQCRKRADMFTEEQLKTIFGNIEDIYKCQKKFVKALEKKFNKDHPHLSEVGSCFLEYQTEFQIYSEYCNNHPNACMELSRLTKVNKYVYFFEACRLLQKMIDISLDGFLLTPVQKICKYPLQLAELLKYTNPQHRDFKDVEAALNAMKNVARLINERKRRLENIDKIAQWQSSIEDWEGEDVLVRSSELIYSGELTKISQPQAKSQQRMFFLFDHQLVCCKKDLLRRDILYYKSRINMDDMEIVDMEDGKDKDFNISVKNAFKLLCRDPEEVHLFCAKKPEQKQRWLKAFENERKQVQLDQETGFSITEVQKKQAMLNANKQHHTGKPKGECGPFFFF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J135</accession>
    <name>A0A452J135_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">ARID domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038596.1">
        <property type="protein sequence ID" value="ENSGAGP00000034084.1"/>
        <property type="gene ID" value="ENSGAGG00000024184.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035060">
        <property type="term" value="C:brahma complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071565">
        <property type="term" value="C:nBAF complex"/>
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        <property type="match status" value="1"/>
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        <property type="entry name" value="ARID DNA-binding domain"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Leucine-rich Repeat Variant"/>
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    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0805">Transcription regulation</keyword>
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        <location>
            <begin position="548"/>
            <end position="639"/>
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    <sequence length="1503" mass="161861" checksum="8808578E88FAFDD1" modified="2019-05-08" version="1">RERGSPQLSLPAAAPGLVGRDASPAMAALGLRLQDRRPSPSRSGLGLGGGAPAEPGAKLPCPGCAWVCAPQPWQVWAAAQLGLGFASAVPWVLVLTHTLFFFFFSQDLSGSIDDLPTGTEATLSSAVSASGSTSSQGEQSNPAQSPFSPHASPHLSGIPGGPSPSPVGSPVGSNQSRSGPISPASIPGSQMPPQPPGSQSESSSHPALSQSPMPQDRGFMAGIQRNPQISQYGPQQTGPSMSPHPSPGGQMHPGIGSFQQSNSSGTYGPQMSQYGPQGNYSRPSTYSGVANTSYSGPGPGMGINANSQIHGQGPSQPCGTMPLGRMPSAGMQSRPFPGNMGSMTPNSPGMSLQGGPGMGPPMPTVNRKAQEAAAAVMQAAANSAQSRQGSFPGMNQSGIMGSSSPYTQPMNNSSGLMNPQAPPYSMAPNMVNSSTGNAPMGLADMMTPGESKLPLPLKADGKEEGPSQPESKSKDSYSSQGISQPPTPGNLPVPSPMSPSSASISSFHGDESDSISSPGWPKTPSSPKSSSSTTTGEKITKVYELGSEPERKMWVDRYLTFMEERGTPVASLPAVGKKPLDLFRLYVCVKEIGGLAQVNKNKKWRELATNLNVGTSSSAASSLKKQYIQYLFAFECKIERGEEPPPEVFSTGDTKKQAKIQPPSPANSGSLQGPQTPQSTGSNSMTEVPGDLKPPTPASTPHGQMTPMQGGRNSAVSVHDPFSDVSDSAFPKRNSMTPNAPYQQGMNMPDMMGRMPYEPNKDPFGGIRKVPGSSEPFMAPGQMPNSGIQDIYNQTPSGAMSTMSMSQRQQFSYGGGYDRRHEPYGQQYPGQGPPSGQPPYGGHQPGMYPQQQVKEVESFIFMTFILXQPVLKPRRKITSKDIVTPEAWRVMMSLKSGLLAESTWALDTINILLYDDSTVATFNLSQLSGFLELLVEYFRKCLIDIFGILMEYEVGDPGHKALDHNAVKKDDTQSLAVDTGKEEENDECIDYFDEEEEDEEEEEEDDIEKIEGEEKSTVFATPSAIADPSERPKQASKFDKLPIKIVKKNNLFVIDRSDKLGRVQEFNSGLLHWQLGGGDTTEHIQTHFESKMEIPPRRRAPPPLNSPSKKKDLEGKGEPEEQQEKSITATIDDVLSARPGALSEDSNPGSQTESSKFPFGIHQAKSHRNIKLLEDEPRSRDETPLCTITHWQDSLAKRCICVSNIVRSLSFVPGNDTEMSKHPGLVLILGKLILLHHEHPERKRAPQTYEKEEEEDKGVACSKDEWWWDCLEVLRDNTLVTLANISGQLDLSAYTESICLPILDGLLHWMVCPSAEAQDPFPTVGPNSVLSPQRLVLETLCKLSIQDNNVDLILATPPFSRQEKLYATLVRYVGDRKNPVCREMSMALLSNLAQGDMLAARAIAVQKGSIGNLISFLEDGVTMAQYQQSQHNLMHMQPPPLEPPSVDMMCRAAKALLAMARVDENRSEFLLHEGRLLDISISSVLNSLVASVICDVLFQIGQL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J136</accession>
    <name>A0A452J136_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">ARID domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038605.1">
        <property type="protein sequence ID" value="ENSGAGP00000034093.1"/>
        <property type="gene ID" value="ENSGAGG00000024184.1"/>
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        <property type="component" value="Unassembled WGS sequence"/>
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        <property type="project" value="UniProtKB-ARBA"/>
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        <property type="entry name" value="Leucine-rich Repeat Variant"/>
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        <property type="entry name" value="ARID"/>
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    </dbReference>
    <proteinExistence type="predicted"/>
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    <sequence length="1354" mass="147398" checksum="921FAEA4DFC3034B" modified="2019-05-08" version="1">MPPQPPGSQSESSSHPALSQSPMPQDRGFMAGIQRNPQISQYGPQQTGPSMSPHPSPGGQMHPGIGSFQQSNSSGTYGPQMSQYGPQGNYSRPSTYSGVANTSYSGPGPGMGINANSQIHGQGPSQPCGTMPLGRMPSAGMQSRPFPGNMGSMTPNSPGMSLQGGPGMGPPMPTVNRKAQEAAAAVMQAAANSAQSRPPYIRSPAYPSQSGAGGRPMFSSQHPNYGNAQAPMMHQPDQYGQGSFPGMNQSGIMGSSSPYTQPMNNSSGLMNPQAPPYSMAPNMVNSSTAPMGLADMMTPGESKLPLPLKADGKEEGPSQPESKSKDSYSSQGISQPPTPGNLPVPSPMSPSSASISSFHGDESDSISSPGWPKTPSSPKSSSSTTTGEKITKVYELGSEPERKMWVDRYLTFMEERGTPVASLPAVGKKPLDLFRLYVCVKEIGGLAQVNKNKKWRELATNLNVGTSSSAASSLKKQYIQYLFAFECKIERGEEPPPEVFSTGDTKKQAKIQPPSPANSGSLQGPQTPQSTGSNSMTEVPGDLKPPTPASTPHGQMTPMQGGRNSAVSVHDPFSDVSDSAFPKRNSMTPNAPYQQGMNMPDMMGRMPYEPNKDPFGGIRKVPGSSEPFMAPGQMPNSGIQDIYNQTPSGAMSTMSMSQRQQFSYGGGYDRRHEPYGQQYPGQGPPSGQPPYGGHQPGMYPQQQVKEVESFIFMTFILXQPVLKPRRKITSKDIVTPEAWRVMMSLKSGLLAESTWALDTINILLYDDSTVATFNLSQLSGFLELLVEYFRKCLIDIFGILMEYEVGDPGHKALDHNAVKKDDTQSLAVDTGKEEENDECIDYFDEEEEDEEEEEEDDIEKIEGEEKSTVFATPSAIADPSERPKQASKFDKLPIKIVKKNNLFVIDRSDKLGRVQEFNSGLLHWQLGGGDTTEHIQTHFESKMEIPPRRRAPPPLNSPSKKKDLEGKGEPEEQQEKSITATIDDVLSARPGALSEDSNPGSQTESSKFPFGIHQAKSHRNIKLLEDEPRSRDETPLCTITHWQDSLAKRCICVSNIVRSLSFVPGNDTEMSKHPGLVLILGKLILLHHEHPERKRAPQTYEKEEEEDKGVACSKDEWWWDCLEVLRDNTLVTLANISGQLDLSAYTESICLPILDGLLHWMVCPSAEAQDPFPTVGPNSVLSPQRLVLETLCKLSIQDNNVDLILATPPFSRQEKLYATLVRYVGDRKNPVCREMSMALLSNLAQGDMLAARAIAVQKGSIGNLISFLEDGVTMAQYQQSQHNLMHMQPPPLEPPSVDMMCRAAKALLAMARVDENRSEFLLHEGRLLDISISSVLNSLVASVICDVLFQIGQL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J145</accession>
    <name>A0A452J145_9SAUR</name>
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        <recommendedName>
            <fullName evidence="12">ARID domain-containing protein</fullName>
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        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
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        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
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        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
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        <citation type="submission" date="2025-08" db="UniProtKB">
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        <scope>IDENTIFICATION</scope>
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        <citation type="submission" date="2025-09" db="UniProtKB">
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        <scope>IDENTIFICATION</scope>
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    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS51011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034094.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1251" mass="136464" checksum="B2D65D93A34183BB" modified="2019-05-08" version="1">MPPQPPGSQSESSSHPALSQSPMPQDRGNYSRPSTYSGVANTSYSGPGPGMGINANSQIHGQGPSQPCGTMPLGRMPSAGMQSRPFPGNMGSMTPNSPGMSLQGGPGMGPPMPTVNRKAQEAAAAVMQAAANSAQSRQGSFPGMNQSGIMGSSSPYTQPMNNSSGLMNPQAPPYSMAPNMVNSSTAPMGLADMMTPGESKLPLPLKADGKEEGPSQPESKSKDSYSSQGISQPPTPGNLPVPSPMSPSSASISSFHGDESDSISSPGWPKTPSSPKSSSSTTTGEKITKVYELGSEPERKMWVDRYLTFMEERGTPVASLPAVGKKPLDLFRLYVCVKEIGGLAQVNKNKKWRELATNLNVGTSSSAASSLKKQYIQYLFAFECKIERGEEPPPEVFSTGDTKKQAKIQPPSPANSGSLQGPQTPQSTGSNSMTEVPGDLKPPTPASTPHGQMTPMQGGRNSAVSVHDPFSDVSDSAFPKRNSMTPNAPYQQGMNMPDMMGRMPYEPNKDPFGGIRKVPGSSEPFMAPGQMPNSGIQDIYNQTPSGAMSTMSMSQRQQFSYGGGYDRRHEPYGQQYPGQGPPSGQPPYGGHQPGMYPQQQVKEVESFIFMTFILXQPVLKPRRKITSKDIVTPEAWRVMMSLKSGLLAESTWALDTINILLYDDSTVATFNLSQLSGFLELLVEYFRKCLIDIFGILMEYEVGDPGHKALDHNAVKKDDTQSLAVDTGKEEENDECIDYFDEEEEDEEEEEEDDIEKIEGEEKSTVFATPSAIADPSERPKQASKFDKLPIKIVKKNNLFVIDRSDKLGRVQEFNSGLLHWQLGGGDTTEHIQTHFESKMEIPPRRRAPPPLNSPSKKKDLEGKGEPEEQQEKSITATIDDVLSARPGALSEDSNPGSQTESSKFPFGIHQAKSHRNIKLLEDEPRSRDETPLCTITHWQDSLAKRCICVSNIVRSLSFVPGNDTEMSKHPGLVLILGKLILLHHEHPERKRAPQTYEKEEEEDKGVACSKDEWWWDCLEVLRDNTLVTLANISGQLDLSAYTESICLPILDGLLHWMVCPSAEAQDPFPTVGPNSVLSPQRLVLETLCKLSIQDNNVDLILATPPFSRQEKLYATLVRYVGDRKNPVCREMSMALLSNLAQGDMLAARAIAVQKGSIGNLISFLEDGVTMAQYQQSQHNLMHMQPPPLEPPSVDMMCRAAKALLAMARVDENRSEFLLHEGRLLDISISSVLNSLVASVICDVLFQIGQL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J147</accession>
    <name>A0A452J147_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">RNA helicase</fullName>
            <ecNumber evidence="3">3.6.4.13</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="9">
            <text>ATP + H2O = ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:13065"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="3.6.4.13"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the DNA2/NAM7 helicase family. SDE3 subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.6.4.13" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452J147"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038634.1">
        <property type="protein sequence ID" value="ENSGAGP00000034119.1"/>
        <property type="gene ID" value="ENSGAGG00000024226.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016787">
        <property type="term" value="F:hydrolase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003724">
        <property type="term" value="F:RNA helicase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="CDD" id="cd18078">
        <property type="entry name" value="DEXXQc_Mov10L1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18808">
        <property type="entry name" value="SF1_C_Upf1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000999">
        <property type="entry name" value="Mov10 like RISC complex RNA helicase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041679">
        <property type="entry name" value="DNA2/NAM7-like_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041677">
        <property type="entry name" value="DNA2/NAM7_AAA_11"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049080">
        <property type="entry name" value="MOV-10-like_beta-barrel"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047187">
        <property type="entry name" value="SF1_C_Upf1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45418">
        <property type="entry name" value="CANCER/TESTIS ANTIGEN 55"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45418:SF1">
        <property type="entry name" value="CANCER_TESTIS ANTIGEN 55"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13086">
        <property type="entry name" value="AAA_11"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13087">
        <property type="entry name" value="AAA_12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21634">
        <property type="entry name" value="MOV-10_beta-barrel"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-0347">Helicase</keyword>
    <keyword evidence="6" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="5" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="10">
        <location>
            <begin position="1"/>
            <end position="18"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019055587" description="RNA helicase" evidence="10">
        <location>
            <begin position="19"/>
            <end position="1070"/>
        </location>
    </feature>
    <feature type="domain" description="Helicase MOV-10-like beta-barrel" evidence="13">
        <location>
            <begin position="528"/>
            <end position="599"/>
        </location>
    </feature>
    <feature type="domain" description="DNA2/NAM7 helicase helicase" evidence="11">
        <location>
            <begin position="676"/>
            <end position="762"/>
        </location>
    </feature>
    <feature type="domain" description="DNA2/NAM7 helicase helicase" evidence="11">
        <location>
            <begin position="787"/>
            <end position="856"/>
        </location>
    </feature>
    <feature type="domain" description="DNA2/NAM7 helicase-like C-terminal" evidence="12">
        <location>
            <begin position="875"/>
            <end position="1039"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005601"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012552"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022806"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00047984"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF13086"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF13087"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="Pfam" id="PF21634"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034119.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1070" mass="119820" checksum="2D9549CB4AA2E28C" modified="2019-05-08" version="1">MLQLLSRVLAYFWRRAEGAGPGVQRPAATGTLKTVQGVVTRFCHDYGMIDDLICFTSDAVMGGVPLNVGQKVIAIVEEDKTSNGLKAIRVEAVSDKWENYSSPTYDSSELKTKMLIGNITSLTEDGGYINKTTSFSMAQVCEGFEPCKGDWVQAEYFINPTTWSSEAVSVKPLRYKRVDKVRISSVCGRIGVVDDSIFFILDYLKLPDGYSPRRHDLVNVVVVESNQSCYIWRALCMAPSDKNGDKGGLKVSRMTNFGTLKLGESKNMVIWIENKGNVPHSLISCRLAGWEKEKQFSFQLPEKGQKTPTAFLSSSVPVKQENLSKGSISSLNDSREPMCQPLNNAFVTNNGTPPGIGENGRNVEEQPTVETEASGEIFIPPDGKTSIVIICTAVNPGLCKELLLLCFSDFIIGRYIEVTVINEEETLIAATEPFSLRKHKVIPELQRMKTTVTVTEHKRSCRRHLPSFLPHYPIPDRLRKCVERKLDILTFQPLLAERLNLSNYKANLSTLLWLEEIHAEMEIKEFSMSGVILKKNGNFLVLEVPGLAEGRPSLNAGDKVILKSQDYSEHIIEYIAYVTEIHDEDATLRLNPEFEQVYNFEPMDVEFALNRILFFLSFPFLKPKVLFPDTLVLQSPQVVKTWNNMGRRTVAADHEHIVNGIMTPEQRDCEFFNPVLNEHQKLAVKRILSGECRPTPYVLFGPPGTGKTVTMIEAILQVHYTLPDSRILVCAPSNSATDLVCLRLHDSNFLKPGVMVRVNASCRCEETISDTVKLYCKDGEDIWKASRFRIIITTCSSAGMFYQIGVRLGHFTHVFVDEAGQASEPECLIPLGLISEVNGQIVLAGDPMQLGPVIKSTLALAYGLNISMLERLMSRSLYQRNEDTFGAYGSYNPLLVTKLLKNYRSHSTLLALPSKLFYHKELEVCADTSVVTSLLGWEKLPRKGFPLIFHGMRGTETREGHNPSWFNPTEAVQVMRYCCLLAKHVNNAVSVTDIGVITPYRKQVEKIRFLLRNVDLMDIKVGSVEEFQGQEYLVIILSTDPCFSALLEYSLANGVYVGCDLPSELKSLQQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIB0</accession>
    <name>A0A452GIB0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">ALMS motif domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Microtubule organizing center</location>
            <location evidence="1">Centrosome</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000001563.1">
        <property type="protein sequence ID" value="ENSGAGP00000001387.1"/>
        <property type="gene ID" value="ENSGAGG00000001108.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005814">
        <property type="term" value="C:centriole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046599">
        <property type="term" value="P:regulation of centriole replication"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029299">
        <property type="entry name" value="ALMS_motif"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21553:SF26">
        <property type="entry name" value="ALMS MOTIF DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21553">
        <property type="entry name" value="ALMS1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15309">
        <property type="entry name" value="ALMS_motif"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="3" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="ALMS motif" evidence="6">
        <location>
            <begin position="1777"/>
            <end position="1832"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="370"/>
            <end position="394"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="698"/>
            <end position="733"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="1260"/>
            <end position="1281"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="1804"/>
            <end position="1836"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="4">
        <location>
            <begin position="108"/>
            <end position="150"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="4">
        <location>
            <begin position="222"/>
            <end position="280"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="4">
        <location>
            <begin position="496"/>
            <end position="542"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="4">
        <location>
            <begin position="918"/>
            <end position="974"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="370"/>
            <end position="389"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="5">
        <location>
            <begin position="708"/>
            <end position="722"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="5">
        <location>
            <begin position="1261"/>
            <end position="1276"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="1804"/>
            <end position="1830"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="Pfam" id="PF15309"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001387.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1836" mass="209825" checksum="69290F0AE5DDCE19" modified="2019-05-08" version="1">MYAKMKRKVAKGGRLRLSPNEEALLLKEEYERRRRLRLQQVREQERYIAVQIRQEVKQRRDEQLHQLAEELKAEWWKAQNQKIKALEKLYLSSLRAVGEGHRQAKENEPDLVALAKEAEERKQRAEKRHKEALKEQKNQKEKLLKEQTWRTNARKHAMYVEKERAAKIAGLPPPPPHPFENFEVKRLPTVKVYNADNFSVTHHHLLEPYVDREMDTHQPDARLVAEEEAKRLDKLQKEEERDRREQLEKAHLRGSHALKIVHLAQDRERLMKELEQMQNMDLARRRQIVAQMPPQLFERAYRRMEIKEDWQRELEFAFEDMYSGDRKMKGDMILHLEPQPLPIPSDRSQDEDLDISLEPDFAYETQPKLDDVTEKESQHPNEPENHEVRVAPQPQSKLVLKKLLNKIQSQKNQWTSKCEAEIQSEIETIESGTLASEERRLCDSDPGHEQHTDSVSEAIESPEILEHTIIAGNSVLIHPQEQAAKIRMEVETQKQIERIEQQKQQQLILLKQIEEKKICLEAEFLKVQMKAQLEEIKKKEQEEKTVSVELRPDPLMNQQREVEHKTETAAVPGTSSSNEENHIQMIRNYQQCLLLQNRLHKQSVEEARKQLQEYQNKLKQRYPSISMPLGSAITLQSRFINLKPTTCESVLQMRGPTVFHRDHPSANQSSVQDHVQEYAPPLTTHSRTHFVLEKPFEQRDEQRHNMGSSRQIQSIEASQSEQRQFHLPQQNYPSQEQMEILGPSDNQVRDLSESQKLPGSVMMTSQDASVMMQTEVHIQRDSSALPTEESSESSRTLGLDKKSVMSQKMPNLLIETEAQKQPQLIVLDTPAAEGSLIAQAASFDSSEYQQVPTESSIKTTFEQTIPQAFQHLTSLFPDESETGAIQESLASKNGGASLLNYSDILKLRERVLASSESIQAQQDHLKELQEQLDVQREALLSRQKMQEELLLRKQAKLKEQMQRQQEALKDFLNKQAEDSNTYREMTETQKCDHFNLMTYLFRDAENKSQEANCGDVNKYDENRLLSQSISRAEQIEQSQRILGREQRWRTSKPPLAKIKLGLDLEQHELSVIPELDTPRSGKVSFTDKTDSVGEEASLISSVEELVSGMCNSESVHEERDLSIANCKEQGSSENPRYTNQLSSSWHERLMVDASSSHEPANPKASVVMDQDLPIYAADIGRGVITDSGPSFRPDSTVALSKGWTTSFAPQACAREVACDYLSSTTISTGSFLTSEKLDLSPANTELSSSNSAEDRILRETASSPWDSSASPAPATALQRQENSSGVCEFHLPAEENFIFNRSQIQQIIEKYTKDLNWLPENSTSFRAPAVGLDFSEIERNFPDFQHQLFQPLEPSPDFDISSSLSQHRISEDSKDFGKSFPFSIRSQDTSVFPEDTSSSLKARSYNLHSFLNTKKLNNTTSQAEESIKANVTLGSEESFHPLHLESTLNEHSQSTEQPIGLRIVFEQNSKIKKSENENYVSPVGDQDEGLARNIENTNLQSSLENLRSPTLSSLEELGSFYQLITTQITENETLLTEHSVKEVGELRKENLSFEELPVVTPNKHRDLTETPVLKNEANRIVEGDHSPCSTNILEEEQSILKVQMSSRMPRAIDSEIYVSPSQATGQQNGGLPEIPQNQNFNSVLTISSGSHSLQSRVPVWETESGHGIMEEPELTLISSNDISVAESDSEHSSQEKKRKNKFDSLAYAGQSELKSCSEGREFLPLDPEADYSVFTLSDYAAIAQSPNEDQSPLHQSAGVVVSQLKKVGEVKVCSREDKRSADVQMQQRTLRLYNQLAEVKIRKEEKSRQETYAKNREKAKEFQKKTLEKLRAKKTH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIB1</accession>
    <name>A0A452GIB1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">DNA ligase</fullName>
            <ecNumber evidence="15">6.5.1.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">DNA ligase that seals nicks in double-stranded during DNA repair. Also involved in DNA replication and DNA recombination.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13 15">
            <text>ATP + (deoxyribonucleotide)n-3'-hydroxyl + 5'-phospho-(deoxyribonucleotide)m = (deoxyribonucleotide)n+m + AMP + diphosphate.</text>
            <dbReference type="EC" id="6.5.1.1"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 16">Belongs to the ATP-dependent DNA ligase family.</text>
    </comment>
    <dbReference type="EC" id="6.5.1.1" evidence="15"/>
    <dbReference type="AlphaFoldDB" id="A0A452GIB1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001475.1">
        <property type="protein sequence ID" value="ENSGAGP00000001316.1"/>
        <property type="gene ID" value="ENSGAGG00000000751.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003910">
        <property type="term" value="F:DNA ligase (ATP) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071897">
        <property type="term" value="P:DNA biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006310">
        <property type="term" value="P:DNA recombination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006281">
        <property type="term" value="P:DNA repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903461">
        <property type="term" value="P:Okazaki fragment processing involved in mitotic DNA replication"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07900">
        <property type="entry name" value="Adenylation_DNA_ligase_I_Euk"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd07969">
        <property type="entry name" value="OBF_DNA_ligase_I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.3260.10:FF:000001">
        <property type="entry name" value="DNA ligase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.50.140:FF:000062">
        <property type="entry name" value="DNA ligase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.470.30:FF:000016">
        <property type="entry name" value="DNA ligase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1490.70">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.3260.10">
        <property type="entry name" value="DNA ligase, ATP-dependent, N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.470.30">
        <property type="entry name" value="DNA ligase/mRNA capping enzyme"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.50.140">
        <property type="entry name" value="Nucleic acid-binding proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050191">
        <property type="entry name" value="ATP-dep_DNA_ligase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000977">
        <property type="entry name" value="DNA_ligase_ATP-dep"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012309">
        <property type="entry name" value="DNA_ligase_ATP-dep_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012310">
        <property type="entry name" value="DNA_ligase_ATP-dep_cent"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016059">
        <property type="entry name" value="DNA_ligase_ATP-dep_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012308">
        <property type="entry name" value="DNA_ligase_ATP-dep_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036599">
        <property type="entry name" value="DNA_ligase_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012340">
        <property type="entry name" value="NA-bd_OB-fold"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00574">
        <property type="entry name" value="dnl1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45674:SF4">
        <property type="entry name" value="DNA LIGASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45674">
        <property type="entry name" value="DNA LIGASE 1/3 FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04679">
        <property type="entry name" value="DNA_ligase_A_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01068">
        <property type="entry name" value="DNA_ligase_A_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04675">
        <property type="entry name" value="DNA_ligase_A_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF117018">
        <property type="entry name" value="ATP-dependent DNA ligase DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56091">
        <property type="entry name" value="DNA ligase/mRNA capping enzyme, catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50249">
        <property type="entry name" value="Nucleic acid-binding proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00697">
        <property type="entry name" value="DNA_LIGASE_A1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00333">
        <property type="entry name" value="DNA_LIGASE_A2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50160">
        <property type="entry name" value="DNA_LIGASE_A3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 15" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="12" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="4" id="KW-0132">Cell division</keyword>
    <keyword evidence="7 15" id="KW-0227">DNA damage</keyword>
    <keyword evidence="9 15" id="KW-0233">DNA recombination</keyword>
    <keyword evidence="10 15" id="KW-0234">DNA repair</keyword>
    <keyword evidence="5" id="KW-0235">DNA replication</keyword>
    <keyword evidence="3 15" id="KW-0436">Ligase</keyword>
    <keyword evidence="6 15" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="ATP-dependent DNA ligase family profile" evidence="18">
        <location>
            <begin position="578"/>
            <end position="714"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="21"/>
            <end position="61"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="17">
        <location>
            <begin position="45"/>
            <end position="59"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007572"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022598"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022705"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023172"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00034003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00058910"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="RuleBase" id="RU000617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="RuleBase" id="RU004196"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50160"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001316.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="840" mass="93227" checksum="9B56089A0EE68F88" modified="2019-05-08" version="1">SEAAAMQRNGLSLCLDSPVKSAKRPLRRVLESDSDEEEAVLTARAEMEPTQDKSLERASSKQVSSAHPLSLPWNSHGASLGSCIPPAPTGRIPANIRPGLHPGLCAHRVSCDSAASPMHALLQLGACLIHAIIVPVATCAGLPSLGVLVQANCWETLRNTSAVGPVYWTWWLPISLWLCDPSSFSKPVVPLDPMEYNPSKSSYHPVQDACWGHGQRVPYLAVARTFERIEEVSARLKNIETLSNLFRSVVALSPSDILPCIYLCLNRLGPAYKGLELGIGETILMKAVAQAMGRQLDQIKAEAQEKGDLGLVAESSRSNQRTMFTPPKLSAAGVFSKLQEIARMTGSASMNKKIDIIKGLFVACRHSEARYITRSLGGKLRIGLAEQSVLSAIAHAVSLTPPGQGFPPEVLDAGQSKSAEARKAWLEEQTQILKQTFCELPSYDIIVPILLEHGVESLPQHCKITPGIPLKPMLAQPTKGIGEVLKRFEEAAFTCEYKYDGERAQIHILENGDVHIYSRNQEDNTSKYPDIISRIPKVKKDSVQSCILDAEAVAWDPEKRQIQPFQVLTTRKRKDVEAAEIKVQVCLYAFDILYLNGESLVKEPFSRRRQLLHESFVETEGEFLFATSMDTCSLDEISEFLDQSIKDSCEGLMVKTLEVDATYEIAKRSHKWLKLKKDYLEGVGDTLDLVVIGAYLGKGKRVGMYGGFLLACYDEESEEYQSICKVGSVGSLHRAGNLQKPRPYYRWDSSTEPNHWLAAVQVWEVKCADLSISPVHKAAIGLVDEEKGISLRFPRFLRVRDDKKPEEATTSSQVAELYRRQQQIQNQQLVEKAEEEEDFY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIB3</accession>
    <name>A0A452GIB3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">Exocyst complex component EXOC6/Sec15 N-terminal domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GIB3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001488.1">
        <property type="protein sequence ID" value="ENSGAGP00000001328.1"/>
        <property type="gene ID" value="ENSGAGG00000001025.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000145">
        <property type="term" value="C:exocyst"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006893">
        <property type="term" value="P:Golgi to plasma membrane transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006886">
        <property type="term" value="P:intracellular protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090522">
        <property type="term" value="P:vesicle tethering involved in exocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007225">
        <property type="entry name" value="EXOC6/Sec15"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048359">
        <property type="entry name" value="EXOC6_Sec15_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12702:SF2">
        <property type="entry name" value="EXOCYST COMPLEX COMPONENT 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12702">
        <property type="entry name" value="SEC15"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20651">
        <property type="entry name" value="EXOC6_Sec15_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Exocyst complex component EXOC6/Sec15 N-terminal" evidence="1">
        <location>
            <begin position="44"/>
            <end position="146"/>
        </location>
    </feature>
    <evidence type="ECO:0000259" key="1">
        <source>
            <dbReference type="Pfam" id="PF20651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001328.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="155" mass="18012" checksum="4650DFD5C38BC152" modified="2019-05-08" version="1">MAESLAEHERILQEIESTDTACVGPTLRSVYDDQPNAHQRFMEKLDACIRNHDREIEKMCNFHHQGFVDAITELLKVRADAGKLKVQVTDTNRRLQDAGKEVIAQTEEIIRCRIQQRNITTVVEKLQLCLPVLEMYSKLKEQMNVKRQQVRCFSD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIC0</accession>
    <name>A0A452GIC0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Eukaryotic translation initiation factor 4 gamma 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="11">Appears to play a role in the switch from cap-dependent to IRES-mediated translation during mitosis, apoptosis and viral infection. Cleaved by some caspases and viral proteases.</text>
    </comment>
    <comment type="subunit">
        <text evidence="13">Interacts with the serine/threonine protein kinases MKNK1 and MKNK2. Binds EIF4A and EIF3. Interacts with MIF4GD. Interacts with DAZAP2.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the eukaryotic initiation factor 4G family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIC0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001576.1">
        <property type="protein sequence ID" value="ENSGAGP00000001397.1"/>
        <property type="gene ID" value="ENSGAGG00000000987.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016281">
        <property type="term" value="C:eukaryotic translation initiation factor 4F complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003729">
        <property type="term" value="F:mRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003743">
        <property type="term" value="F:translation initiation factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006446">
        <property type="term" value="P:regulation of translational initiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd11559">
        <property type="entry name" value="W2_eIF4G1_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.180:FF:000007">
        <property type="entry name" value="Eukaryotic translation initiation factor 4 gamma 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.180:FF:000011">
        <property type="entry name" value="Eukaryotic translation initiation factor 4 gamma 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.180:FF:000017">
        <property type="entry name" value="Eukaryotic translation initiation factor 4 gamma 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.180">
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003891">
        <property type="entry name" value="Initiation_fac_eIF4g_MI"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003890">
        <property type="entry name" value="MIF4G-like_typ-3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003307">
        <property type="entry name" value="W2_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23253">
        <property type="entry name" value="EUKARYOTIC TRANSLATION INITIATION FACTOR 4 GAMMA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23253:SF9">
        <property type="entry name" value="EUKARYOTIC TRANSLATION INITIATION FACTOR 4 GAMMA 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02847">
        <property type="entry name" value="MA3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02854">
        <property type="entry name" value="MIF4G"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02020">
        <property type="entry name" value="W2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00515">
        <property type="entry name" value="eIF5C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00544">
        <property type="entry name" value="MA3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00543">
        <property type="entry name" value="MIF4G"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51366">
        <property type="entry name" value="MI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51363">
        <property type="entry name" value="W2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0007">Acetylation</keyword>
    <keyword evidence="5" id="KW-0396">Initiation factor</keyword>
    <keyword evidence="4" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="2" id="KW-0488">Methylation</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="9" id="KW-0648">Protein biosynthesis</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0678">Repressor</keyword>
    <keyword evidence="8" id="KW-0810">Translation regulation</keyword>
    <keyword evidence="7" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="MI" evidence="16">
        <location>
            <begin position="508"/>
            <end position="631"/>
        </location>
    </feature>
    <feature type="domain" description="W2" evidence="15">
        <location>
            <begin position="685"/>
            <end position="869"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="17"/>
            <end position="74"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="40"/>
            <end position="56"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="64"/>
            <end position="74"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00005775"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022491"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022540"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022845"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022917"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00037759"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00040449"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00046720"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS51363"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS51366"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001397.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="872" mass="98420" checksum="6E101DE7528D6551" modified="2019-05-08" version="1">ILRCQAAKVESAIAEGGASRFSASSGGGGSRGAPQHYPKTASNSEFLGKTPGQNAQKWIPSRSTRRDDDSANDKERHDAIFRKVRGILNKLTPEKFDKLCLELLNVGVESKLILKGIILLIVDKALEEPKYSSLYAQLCLRLAEDAPNFDGPSAECHPGQKQSTTFRRLLISKLQDEFENRTRNVDIYDKRDGPLLPEEEEQRAIAKIKMLGNIKFIGELGKLDLIHESILHKCIKTLLEKKKRVQLKDMGEDLECLCQIMRTVGPRLDHAKAKSLMDQYFARMRSLMLSKELPARIRFLLQDTVELREHNWVPRKAFLDNGPKTINQIRQDAVKDLGVFIPAPMAQGMRSDFFLEGPFMPPRMKLDRDPLGGLADMFGQMPGSGIGTGPGVIQDRFSPTMGRHRSNQLFNGHGGHLMPPAQSQFGDLGKSFLKNQISLRPAQSFLMNKNQVPKLQPQITMIPPSAQPPRTQTPPLGQPPQLGLKTNPPLIQEKPAKISKKPPPSKEELLKLTEAVVTDYLNNGNVNDAISGVREMRAPKPFIPEMLSKVIIQSLDRSDEDKEKASALISLLKQEGIATSDNFMQAFLNVLDQCPKLEVDIPLVKSYLAQFAARAIISELVSISELAQPLESGTHFPLFLLCLQQLAKLQDREWLTELFQQSKVNMQKMLPEIDQNKDRMLEILEGKGLSFLFPLLKLEKELLKQIKLDPSPQAIYKWIKDNISPKLHVDKGFVNILMTSFLQYISSEVSPPNDEPDSSSAPSKEQLEQEKQLLLSFKPVMQKFLHDHVDLQVSALYALQVHCYNNNFPKGMLLHFFVHFYDMEIIEEEAFLAWKEDISQEFPGKGKALFQVNQWLTWLETAEEEESEEEAD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIC5</accession>
    <name>A0A452GIC5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="30">Thromboxane-A synthase</fullName>
            <ecNumber evidence="7">4.2.1.152</ecNumber>
            <ecNumber evidence="29">5.3.99.5</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="31">Cytochrome P450 5A1</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="25">Hydroperoxy icosatetraenoate dehydratase</fullName>
        </alternativeName>
    </protein>
    <organism evidence="36 37">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="37" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="36" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="36" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="32">Catalyzes the conversion of prostaglandin H2 (PGH2) to thromboxane A2 (TXA2), a potent inducer of blood vessel constriction and platelet aggregation. Also cleaves PGH2 to 12-hydroxy-heptadecatrienoicacid (12-HHT) and malondialdehyde, which is known to act as a mediator of DNA damage. 12-HHT and malondialdehyde are formed stoichiometrically in the same amounts as TXA2. Additionally, displays dehydratase activity, toward (15S)-hydroperoxy-(5Z,8Z,11Z,13E)-eicosatetraenoate (15(S)-HPETE) producing 15-KETE and 15-HETE.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="26">
            <text>(15S)-hydroperoxy-(5Z,8Z,11Z,13E)-eicosatetraenoate + AH2 = (15S)-hydroxy-(5Z,8Z,11Z,13E)-eicosatetraenoate + A + H2O</text>
            <dbReference type="Rhea" id="RHEA:48856"/>
            <dbReference type="ChEBI" id="CHEBI:13193"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:17499"/>
            <dbReference type="ChEBI" id="CHEBI:57409"/>
            <dbReference type="ChEBI" id="CHEBI:57446"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="26">
            <dbReference type="Rhea" id="RHEA:48857"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>(15S)-hydroperoxy-(5Z,8Z,11Z,13E)-eicosatetraenoate = 15-oxo-(5Z,8Z,11Z,13E)-eicosatetraenoate + H2O</text>
            <dbReference type="Rhea" id="RHEA:48636"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:57410"/>
            <dbReference type="ChEBI" id="CHEBI:57446"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="1">
            <dbReference type="Rhea" id="RHEA:48637"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="2">
            <text>a hydroperoxyeicosatetraenoate = an oxoeicosatetraenoate + H2O</text>
            <dbReference type="Rhea" id="RHEA:55556"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:59720"/>
            <dbReference type="ChEBI" id="CHEBI:131859"/>
            <dbReference type="EC" id="4.2.1.152"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="2">
            <dbReference type="Rhea" id="RHEA:55557"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="27">
            <text>prostaglandin H2 = (12S)-hydroxy-(5Z,8E,10E)-heptadecatrienoate + malonaldehyde</text>
            <dbReference type="Rhea" id="RHEA:48644"/>
            <dbReference type="ChEBI" id="CHEBI:57405"/>
            <dbReference type="ChEBI" id="CHEBI:90694"/>
            <dbReference type="ChEBI" id="CHEBI:566274"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="28">
            <text>prostaglandin H2 = thromboxane A2</text>
            <dbReference type="Rhea" id="RHEA:17137"/>
            <dbReference type="ChEBI" id="CHEBI:57405"/>
            <dbReference type="ChEBI" id="CHEBI:57445"/>
            <dbReference type="EC" id="5.3.99.5"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="28">
            <dbReference type="Rhea" id="RHEA:17138"/>
        </physiologicalReaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="3 33">
            <name>heme</name>
            <dbReference type="ChEBI" id="CHEBI:30413"/>
        </cofactor>
    </comment>
    <comment type="subunit">
        <text evidence="6">Monomer.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="4">Endoplasmic reticulum membrane</location>
            <topology evidence="4">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5 34">Belongs to the cytochrome P450 family.</text>
    </comment>
    <dbReference type="EC" id="4.2.1.152" evidence="7"/>
    <dbReference type="EC" id="5.3.99.5" evidence="29"/>
    <dbReference type="AlphaFoldDB" id="A0A452GIC5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001585.1">
        <property type="protein sequence ID" value="ENSGAGP00000001402.1"/>
        <property type="gene ID" value="ENSGAGG00000001072.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0020037">
        <property type="term" value="F:heme binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0106256">
        <property type="term" value="F:hydroperoxy icosatetraenoate dehydratase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005506">
        <property type="term" value="F:iron ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016705">
        <property type="term" value="F:oxidoreductase activity, acting on paired donors, with incorporation or reduction of molecular oxygen"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008395">
        <property type="term" value="F:steroid hydroxylase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004796">
        <property type="term" value="F:thromboxane-A synthase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001516">
        <property type="term" value="P:prostaglandin biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.630.10:FF:000003">
        <property type="entry name" value="cytochrome P450 3A12-like isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.630.10">
        <property type="entry name" value="Cytochrome P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001128">
        <property type="entry name" value="Cyt_P450"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017972">
        <property type="entry name" value="Cyt_P450_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002401">
        <property type="entry name" value="Cyt_P450_E_grp-I"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036396">
        <property type="entry name" value="Cyt_P450_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050705">
        <property type="entry name" value="Cytochrome_P450_3A"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24302:SF47">
        <property type="entry name" value="CYTOCHROME P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24302">
        <property type="entry name" value="CYTOCHROME P450 FAMILY 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00067">
        <property type="entry name" value="p450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00463">
        <property type="entry name" value="EP450I"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00385">
        <property type="entry name" value="P450"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48264">
        <property type="entry name" value="Cytochrome P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00086">
        <property type="entry name" value="CYTOCHROME_P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="14" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="22" id="KW-0275">Fatty acid biosynthesis</keyword>
    <keyword evidence="15" id="KW-0276">Fatty acid metabolism</keyword>
    <keyword evidence="11 33 34" id="KW-0349">Heme</keyword>
    <keyword evidence="18 33 34" id="KW-0408">Iron</keyword>
    <keyword evidence="23" id="KW-0413">Isomerase</keyword>
    <keyword evidence="9" id="KW-0444">Lipid biosynthesis</keyword>
    <keyword evidence="20" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="24" id="KW-0456">Lyase</keyword>
    <keyword evidence="21 35" id="KW-0472">Membrane</keyword>
    <keyword evidence="13 33 34" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="19 34" id="KW-0503">Monooxygenase</keyword>
    <keyword evidence="17 34" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="10" id="KW-0643">Prostaglandin biosynthesis</keyword>
    <keyword evidence="8" id="KW-0644">Prostaglandin metabolism</keyword>
    <keyword evidence="37" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="12 35" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="16 35" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="35">
        <location>
            <begin position="203"/>
            <end position="221"/>
        </location>
    </feature>
    <feature type="binding site" description="axial binding residue" evidence="33">
        <location>
            <position position="460"/>
        </location>
        <ligand>
            <name>heme</name>
            <dbReference type="ChEBI" id="CHEBI:30413"/>
        </ligand>
        <ligandPart>
            <name>Fe</name>
            <dbReference type="ChEBI" id="CHEBI:18248"/>
        </ligandPart>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001143"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00001719"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00001971"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004477"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00010617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00011245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00013084"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022501"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022516"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022585"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022832"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023033"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00023160"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00023235"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00023239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00033404"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00036380"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="ARBA" id="ARBA00036424"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="ARBA" id="ARBA00036475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="ARBA" id="ARBA00038872"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="ARBA" id="ARBA00040834"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="31">
        <source>
            <dbReference type="ARBA" id="ARBA00042726"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="32">
        <source>
            <dbReference type="ARBA" id="ARBA00054825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="33">
        <source>
            <dbReference type="PIRSR" id="PIRSR602401-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="34">
        <source>
            <dbReference type="RuleBase" id="RU000461"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="35">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="36">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001402.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="37">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="514" mass="58722" checksum="AEDA853D5C5B8855" modified="2019-05-08" version="1">ELEPGVCFVPIVYSTSSFSRLKRVGIRHPAPLPFIGNLLFFRQGFWENHTKLITEYGPVCGYYIGRRMYVLVSEPEMIKHILVKDFSNFTNRMTPGLVTKPVIDSILCLRDQRWKEVRSILTPAFSSAKMKEMTPLINQACDVLLSNLKVYADSGKAFDIQRSYGCFTLDVVASVAFGTHVDSQKNPDDTFVKNCKKFFEFSMLRPILILIIAFPFLMIPLSRILPNKRRDEMNGFFIKAIKNMIALRDQQDANERRRDFLQLILDARSSTNYISVEHFDIVNQADVSIQDNKPSVDKALPKKVQKTLSEDEIVGQAFIFLIGGYETTTNALSFATYLLATNPECQEKLLQEIDEFSGKHDVPDYQNIQELPYLDMVIAETLRMYPPAFRFTRETSKDCVVMGQHIPAGAVIEIAVGPLHYNPEFWPEPEKFIPERFTMEAKLQQHPFAYLPFGAGPRSCIGMRMALLEIKMTLLRILQKFRFETSPKTQIPLQLTSVSSLAPKNGVYVKIVPR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIE2</accession>
    <name>A0A452GIE2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4 11">Ubiquitin-like protein ATG12</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="11">Ubiquitin-like protein involved in autophagic vesicle formation.</text>
    </comment>
    <comment type="subunit">
        <text evidence="11">Forms a conjugate with ATG5.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Preautophagosomal structure membrane</location>
            <topology evidence="2">Peripheral membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 11">Belongs to the ATG12 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIE2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001602.1">
        <property type="protein sequence ID" value="ENSGAGP00000001416.1"/>
        <property type="gene ID" value="ENSGAGG00000001143.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034274">
        <property type="term" value="C:Atg12-Atg5-Atg16 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000421">
        <property type="term" value="C:autophagosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034045">
        <property type="term" value="C:phagophore assembly site membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019776">
        <property type="term" value="F:Atg8-family ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000045">
        <property type="term" value="P:autophagosome assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097352">
        <property type="term" value="P:autophagosome maturation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000422">
        <property type="term" value="P:autophagy of mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061723">
        <property type="term" value="P:glycophagy"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034727">
        <property type="term" value="P:piecemeal microautophagy of the nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd01612">
        <property type="entry name" value="Ubl_ATG12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.20.90:FF:000117">
        <property type="entry name" value="Ubiquitin-like protein ATG12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.20.90">
        <property type="entry name" value="Phosphatidylinositol 3-kinase Catalytic Subunit, Chain A, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007242">
        <property type="entry name" value="Atg12"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029071">
        <property type="entry name" value="Ubiquitin-like_domsf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13385">
        <property type="entry name" value="AUTOPHAGY PROTEIN 12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13385:SF0">
        <property type="entry name" value="UBIQUITIN-LIKE PROTEIN ATG12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04110">
        <property type="entry name" value="APG12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54236">
        <property type="entry name" value="Ubiquitin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0007">Acetylation</keyword>
    <keyword evidence="9 11" id="KW-0072">Autophagy</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6 11" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="10" id="KW-0472">Membrane</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7 11" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1"/>
            <end position="52"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="27"/>
            <end position="42"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004623"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007778"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00015875"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023006"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="RuleBase" id="RU361201"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001416.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="142" mass="15300" checksum="D87644CB9C3A8395" modified="2019-05-08" version="1">MAEVEEQLPLSPQDGGRGECGEEPSDGGAAVGVAEAVPPSAGSPVTEEPAEDTKKKIDVLLKAVGDTPIMKTKKWSVERTRTIQGLVDFIKKFLKLVASEQLFIYVNQSFAPSPDQEVGTLYECFGSDGKLVLHYCKSQAWG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIE5</accession>
    <name>A0A452GIE5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">dolichyl-phosphate-mannose--protein mannosyltransferase</fullName>
            <ecNumber evidence="5">2.4.1.109</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="pathway">
        <text evidence="3">Protein modification; protein glycosylation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Endoplasmic reticulum</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the TMTC family.</text>
    </comment>
    <dbReference type="EC" id="2.4.1.109" evidence="5"/>
    <dbReference type="AlphaFoldDB" id="A0A452GIE5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001529.1">
        <property type="protein sequence ID" value="ENSGAGP00000001359.1"/>
        <property type="gene ID" value="ENSGAGG00000000661.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00378"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004169">
        <property type="term" value="F:dolichyl-phosphate-mannose-protein mannosyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.10">
        <property type="entry name" value="Tetratricopeptide repeat domain"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013618">
        <property type="entry name" value="TMTC_DUF1736"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052384">
        <property type="entry name" value="TMTC_O-mannosyltransferase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011990">
        <property type="entry name" value="TPR-like_helical_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019734">
        <property type="entry name" value="TPR_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR44216">
        <property type="entry name" value="PROTEIN O-MANNOSYL-TRANSFERASE TMTC2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR44216:SF3">
        <property type="entry name" value="PROTEIN O-MANNOSYL-TRANSFERASE TMTC2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08409">
        <property type="entry name" value="TMTC_DUF1736"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13424">
        <property type="entry name" value="TPR_12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13432">
        <property type="entry name" value="TPR_16"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13181">
        <property type="entry name" value="TPR_8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00028">
        <property type="entry name" value="TPR"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48452">
        <property type="entry name" value="TPR-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50005">
        <property type="entry name" value="TPR"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50293">
        <property type="entry name" value="TPR_REGION"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="12 14" id="KW-0472">Membrane</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="9 13" id="KW-0802">TPR repeat</keyword>
    <keyword evidence="6" id="KW-0808">Transferase</keyword>
    <keyword evidence="7 14" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 14" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="74"/>
            <end position="91"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="125"/>
            <end position="145"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="210"/>
            <end position="234"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="300"/>
            <end position="317"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="382"/>
            <end position="406"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="412"/>
            <end position="432"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="439"/>
            <end position="457"/>
        </location>
    </feature>
    <feature type="domain" description="DUF1736" evidence="15">
        <location>
            <begin position="237"/>
            <end position="309"/>
        </location>
    </feature>
    <feature type="repeat" description="TPR" evidence="13">
        <location>
            <begin position="482"/>
            <end position="515"/>
        </location>
    </feature>
    <feature type="repeat" description="TPR" evidence="13">
        <location>
            <begin position="516"/>
            <end position="549"/>
        </location>
    </feature>
    <feature type="repeat" description="TPR" evidence="13">
        <location>
            <begin position="550"/>
            <end position="583"/>
        </location>
    </feature>
    <feature type="repeat" description="TPR" evidence="13">
        <location>
            <begin position="595"/>
            <end position="628"/>
        </location>
    </feature>
    <feature type="repeat" description="TPR" evidence="13">
        <location>
            <begin position="666"/>
            <end position="699"/>
        </location>
    </feature>
    <feature type="repeat" description="TPR" evidence="13">
        <location>
            <begin position="768"/>
            <end position="801"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004922"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00007882"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012839"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022803"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00339"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="Pfam" id="PF08409"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001359.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="825" mass="93232" checksum="5880331AA2B79A4D" modified="2025-10-08" version="2">MVLENTFSTSSGDYVLCGSIKTNQDLLPETPWTHIFYNDFWGTILTHSGSHKSYRPLCTLSFRINHAIGGMDPWSYHLVNVLLHTAVTGLFTNFSRILFGDGYWTLIAGLLFASHPIHTEAVAGIVGRADVGACFFFLLSLMCYVKHCSTRSYSTSTWGWILGTGLCAGCSMLWKEQGVTVLAVSAVYDVFVFHRLKMNQIIPSIYKRKNLSLFFSIGLLTFWGTALLGVRLYWMGNKPPSFSNSDNPAADSDSFLTRTLTFFYLPTKNLWLLLCPDTLSFDWSMDAVPLLKTASDWRNLHTVAFYAGLLLLAYSSLKSSSNKRECNGKTVMNGKQNTNGHSCHSDMEYKSLELKSSFASKEENGIKKRVIQRPQLPSTENIVLLSLSLLIVPFVPATNLFFYVGFVIAERVLYIPSMGFCLLITVGARSLYVKAQKRFLKNFIFCATAALILFYGLKTVVRNEDWQNEEMLYRSGIKVNPAKAWGNLGNVLKSQSKISEAESAYRNALYYRSNMADMLYNLGLLLQENSRFSEALHYYKLAIGSRPTLASAYLNTGIILMTQGKTEEARRTFLKCSEIPDENLKDPHAHKSSVTSCLYNLGKLFHEQGHYEDALMVYKEAIQKMPRQFAPQSLYNMMGEAYMRLSKLPEAEHWYMESLRSKTDHIPAHLTYGKLLALTGRKSEAEKYFLKAIQLDPTKGNGYMHYGQFLLEESRLIEAAEMAKKAAELDNTEFDVVFNAAHMLRQASLNEAAEKYYEMAAGLRPNYPAALMNLGAILHLNGKLRDAEANYLRALQFKPDDVITQSNLRKLWNIMEKQGLKTSKT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIE7</accession>
    <name>A0A452GIE7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Phosphatase and actin regulator</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="5">Binds PPP1CA and actin.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1 5">Belongs to the phosphatase and actin regulator family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIE7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001523.1">
        <property type="protein sequence ID" value="ENSGAGP00000001354.1"/>
        <property type="gene ID" value="ENSGAGG00000001084.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003779">
        <property type="term" value="F:actin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004864">
        <property type="term" value="F:protein phosphatase inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030036">
        <property type="term" value="P:actin cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.1750">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.2130">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004018">
        <property type="entry name" value="RPEL_repeat"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12751:SF7">
        <property type="entry name" value="PHOSPHATASE AND ACTIN REGULATOR 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12751">
        <property type="entry name" value="PHOSPHATASE AND ACTIN REGULATOR PHACTR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02755">
        <property type="entry name" value="RPEL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00707">
        <property type="entry name" value="RPEL"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51073">
        <property type="entry name" value="RPEL"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 5" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="6" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 5" id="KW-0677">Repeat</keyword>
    <feature type="repeat" description="RPEL" evidence="4">
        <location>
            <begin position="52"/>
            <end position="77"/>
        </location>
    </feature>
    <feature type="repeat" description="RPEL" evidence="4">
        <location>
            <begin position="360"/>
            <end position="385"/>
        </location>
    </feature>
    <feature type="repeat" description="RPEL" evidence="4">
        <location>
            <begin position="398"/>
            <end position="423"/>
        </location>
    </feature>
    <feature type="repeat" description="RPEL" evidence="4">
        <location>
            <begin position="436"/>
            <end position="461"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="82"/>
            <end position="158"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="171"/>
            <end position="326"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="409"/>
            <end position="436"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="92"/>
            <end position="115"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="116"/>
            <end position="126"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="130"/>
            <end position="143"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="7">
        <location>
            <begin position="188"/>
            <end position="197"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="251"/>
            <end position="285"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="293"/>
            <end position="323"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00009795"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00401"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="RuleBase" id="RU301113"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001354.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="518" mass="58767" checksum="9EB63997D05251F2" modified="2019-05-08" version="1">MEQSPAMHSEYLVSGIRTPPVRRNSKLATLGRIFKPWKWRKKKNEKLKQTSAVLEKKITARPGRDELIKKGLLEMMEQDAGGKACAGDGATRSMQSDPSSPVCQALTSEEMQQESTAAAAAATNATSLGEELHSEELKDDAAKKPSVSTEDLEDACLPVTEETPQALLISESTDSPQKPMERNLAQLPSPPSLPIPLPKAISKVKSVTGQAALFHPSGQKGSEPHVRGQLPTPTGSPHPTAVHLPLPPSRVIEELHRALATKHRQDSFHSRESKGSPKKRVDVRPSRTSSIEGNKEKEENALSHSNDPENKRNSVKESDENKENLIMNSELKDNLVFYQDEEVLNDSIISGTLPRKCKKELLAVKLRNRPSKQELEDRNIFPRRTDEERQEIRQQIEMKLSKRLSQRPAVEELERRNILKQRNDQTEQEERREIKQRLTRKLNQRPTVDELRDRKILIRFSDYVEVAKAQDYDRRADKPWTRLSAADKAAIRKELNEYKSNEMEVHASSKHLTRFHRP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIE9</accession>
    <name>A0A452GIE9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Ig-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GIE9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001606.1">
        <property type="protein sequence ID" value="ENSGAGP00000001420.1"/>
        <property type="gene ID" value="ENSGAGG00000001083.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00099">
        <property type="entry name" value="IgV"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000604">
        <property type="entry name" value="immunoglobulin superfamily member 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000674">
        <property type="entry name" value="Immunoglobulin superfamily member 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000689">
        <property type="entry name" value="Immunoglobulin superfamily member 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000932">
        <property type="entry name" value="Immunoglobulin superfamily member 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000491">
        <property type="entry name" value="Immunoglobulin superfamily, member 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013106">
        <property type="entry name" value="Ig_V-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051102">
        <property type="entry name" value="IgSF_V-set/TM_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12207:SF21">
        <property type="entry name" value="IMMUNOGLOBULIN SUPERFAMILY MEMBER 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12207">
        <property type="entry name" value="V-SET AND TRANSMEMBRANE DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07686">
        <property type="entry name" value="V-set"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00406">
        <property type="entry name" value="IGv"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="7"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="4" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="5" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1 6" id="KW-0732">Signal</keyword>
    <keyword evidence="5" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="6">
        <location>
            <begin position="1"/>
            <end position="26"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019134057" description="Ig-like domain-containing protein" evidence="6">
        <location>
            <begin position="27"/>
            <end position="1040"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="970"/>
            <end position="993"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="7">
        <location>
            <begin position="22"/>
            <end position="127"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="7">
        <location>
            <begin position="174"/>
            <end position="265"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="7">
        <location>
            <begin position="285"/>
            <end position="418"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="7">
        <location>
            <begin position="438"/>
            <end position="541"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="7">
        <location>
            <begin position="555"/>
            <end position="682"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="7">
        <location>
            <begin position="692"/>
            <end position="818"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="7">
        <location>
            <begin position="830"/>
            <end position="951"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001420.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1040" mass="117478" checksum="11784DAF1E37C99A" modified="2019-05-08" version="1">SLRVFVFSHLLPVFILFIAVIPDSLRVTSVPQTLNKVERDSLELKCEVSKSTAQHSHLSVSWYRQKVGEPPVEVISLSRDFVLRAGSTYAQRQATGNVRLDKVGETTSRLTVYNLQPSDQGEFYCEAAEWIQDPDKSWYAMTRKRSEGTTVNIQATDKEFSVRLDTEKRTYTVGEPVEFRCILEAQNVPERYFSISWAFNSSLIASLGPNAVPVLNSDFAQREALGQLKVAKESDNVFVLKIYRLRLEDSGKYNCRVTEREKTVTGDFIDKESKRPKNTPITVLPLKTSISVEMTNNASNVLEGESLRFGCNVHSGFGPQSHLSVTWQLVDKQNRRSDVVRLDRDGTLQPGPSYSERSSYGGIQMEQVRPGSFTLEIFNSVKADEGHYECHVTEWTRTQDGEWQMIGERHASMPVSITALEAGFAVTAISRTPGVTYNESFDLQCIIKPHYPSWVPVSVTWRFQPAGSTEFHDLVTFTRDGGVQWGDRYMGFRTRTAIEKAESSNNVRLSISRASDTEAGKYQCVAELWRKNYNSSWTRLADRTSNLLEIRVLRPVTKLQVSKSKRSITLVENRPIPLNCSVKSQTSPDSHFAVLWYVHKPADADGKLILKTTHSSAFEYGTYAEEEGLRGRLQFERSSSGGLFSLTVQRAEVRDSGSYYCHVEEWLLSPNHAWYKLAEEVSGRTEVTVKQPDNRLQVNQTHKNLTVLENQWVTLECAVTNRTSPASQLSVEWYVSKSGHPEKEAVARLNRQATLQYGDLATLSDLKGRLHLESPSPGLYLLTLQNVMVQDSGAYECRVEEWLLDPSNQWYKRAEDLSGLTTVTVKQPDPTLQVDTTISNLTVLEKESFPLDCSILSRSSQESRRVSHGMEEEEEREALLSVGLDAVFGQEAGRFEGRLRFQRLSSVLYRLTVLQAGAADAGNYSCRVEEWLPDPKRVWYRLAEEESGTITIHVQDTGSTLQSIICSNDALFYFVFFYPFPIFGILIITILLVRFKSRNSSKNSEGKNGVPLLWIKEPHLNYSPTCLEPPVLSIHPGTID</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIF1</accession>
    <name>A0A452GIF1_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="14">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>all-trans-retinoate + reduced [NADPH--hemoprotein reductase] + O2 = all-trans-4-hydroxyretinoate + oxidized [NADPH--hemoprotein reductase] + H2O + H(+)</text>
            <dbReference type="Rhea" id="RHEA:51984"/>
            <dbReference type="Rhea" id="RHEA-COMP:11964"/>
            <dbReference type="Rhea" id="RHEA-COMP:11965"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:15379"/>
            <dbReference type="ChEBI" id="CHEBI:35291"/>
            <dbReference type="ChEBI" id="CHEBI:57618"/>
            <dbReference type="ChEBI" id="CHEBI:58210"/>
            <dbReference type="ChEBI" id="CHEBI:134178"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="11">
            <dbReference type="Rhea" id="RHEA:51985"/>
        </physiologicalReaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 12">
            <name>heme</name>
            <dbReference type="ChEBI" id="CHEBI:30413"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 13">Belongs to the cytochrome P450 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000001532.1">
        <property type="protein sequence ID" value="ENSGAGP00000001360.1"/>
        <property type="gene ID" value="ENSGAGG00000001074.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0020037">
        <property type="term" value="F:heme binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005506">
        <property type="term" value="F:iron ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004497">
        <property type="term" value="F:monooxygenase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016705">
        <property type="term" value="F:oxidoreductase activity, acting on paired donors, with incorporation or reduction of molecular oxygen"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034653">
        <property type="term" value="P:retinoic acid catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016125">
        <property type="term" value="P:sterol metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd20636">
        <property type="entry name" value="CYP26C1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.630.10:FF:000009">
        <property type="entry name" value="Cytochrome P450 26B1 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.630.10">
        <property type="entry name" value="Cytochrome P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001128">
        <property type="entry name" value="Cyt_P450"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017972">
        <property type="entry name" value="Cyt_P450_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002403">
        <property type="entry name" value="Cyt_P450_E_grp-IV"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036396">
        <property type="entry name" value="Cyt_P450_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24286">
        <property type="entry name" value="CYTOCHROME P450 26"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24286:SF100">
        <property type="entry name" value="CYTOCHROME P450 26C1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00067">
        <property type="entry name" value="p450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00465">
        <property type="entry name" value="EP450IV"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00385">
        <property type="entry name" value="P450"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48264">
        <property type="entry name" value="Cytochrome P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00086">
        <property type="entry name" value="CYTOCHROME_P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 12 13" id="KW-0349">Heme</keyword>
    <keyword evidence="7 12 13" id="KW-0408">Iron</keyword>
    <keyword evidence="9" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="10" id="KW-0472">Membrane</keyword>
    <keyword evidence="5 12 13" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8 13" id="KW-0503">Monooxygenase</keyword>
    <keyword evidence="6 13" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <feature type="binding site" description="axial binding residue" evidence="12">
        <location>
            <position position="438"/>
        </location>
        <ligand>
            <name>heme</name>
            <dbReference type="ChEBI" id="CHEBI:30413"/>
        </ligand>
        <ligandPart>
            <name>Fe</name>
            <dbReference type="ChEBI" id="CHEBI:18248"/>
        </ligandPart>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001971"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004370"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00010617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023033"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00048965"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PIRSR" id="PIRSR602403-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="RuleBase" id="RU000461"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001360.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="499" mass="55907" checksum="6AE7D2AF0ED85ACB" modified="2019-05-08" version="1">SVFSTATRTTALAALLALGLLLTVSQHLWTLRWTLSRDRGSALPLPKGSMGWPFFGETLHWLVQGSSFHSSRREKYGHVFKTHLLGKPVIRVTGAENIRKILLGEHSLVSTQWPQSTQIILGSNTLLNSIGELHRQRRKILARVFSRAALESYLPGIQKVVSWELRGWCREPGSVAVYSSAKTLTFRVAARVLLGLSLEEKQFKDLAKTFEQLVENLFSLPLNIPFSGLRKGIKARDLLHEYMEKAIMEKLQGKNSEGYSDALDFLINSAKEHGKEFTMQELKESAIELIFAAFFTTASASTSLILLLLKHPSAVEKIRQELVSHELHRHCHCLSPAPEMNLDKLSRLRYLDCVVKEVLRVLPPVSGGYRTALQTFELDGYQIPKGWSVMYSIRDTHETAAVYQSPPERFDPDRFGSARQKGAGRFHYIPFGGGLRSCIGQELAKAILKLLAIELVSTARWELATPGFPEMQTVPIVHPVDGLQLFFHPLRPGGAXREA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIF3</accession>
    <name>A0A452GIF3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Peptidase S1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GIF3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001538.1">
        <property type="protein sequence ID" value="ENSGAGP00000001364.1"/>
        <property type="gene ID" value="ENSGAGG00000000827.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00190">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.10:FF:000005">
        <property type="entry name" value="Serine protease 37"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.10.10">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009003">
        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043504">
        <property type="entry name" value="Peptidase_S1_PA_chymotrypsin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001314">
        <property type="entry name" value="Peptidase_S1A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001254">
        <property type="entry name" value="Trypsin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018114">
        <property type="entry name" value="TRYPSIN_HIS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271:SF81">
        <property type="entry name" value="GRANZYME B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271">
        <property type="entry name" value="KALLIKREIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00089">
        <property type="entry name" value="Trypsin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00722">
        <property type="entry name" value="CHYMOTRYPSIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00020">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50494">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50240">
        <property type="entry name" value="TRYPSIN_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00134">
        <property type="entry name" value="TRYPSIN_HIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1 4" id="KW-0732">Signal</keyword>
    <keyword evidence="2" id="KW-0865">Zymogen</keyword>
    <feature type="signal peptide" evidence="4">
        <location>
            <begin position="1"/>
            <end position="19"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019051295" description="Peptidase S1 domain-containing protein" evidence="4">
        <location>
            <begin position="20"/>
            <end position="248"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase S1" evidence="5">
        <location>
            <begin position="22"/>
            <end position="246"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023145"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001364.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="248" mass="27257" checksum="3553ECBD7E783457" modified="2019-05-08" version="1">MLLLLLLPTAFLLPLGAQAGDIIGGREARPHSRPYMAFVETKMGPESTTECGGFLVREDFVLTAAHCDEEGSGDAPTVVILGAHDLNRKERTQQEIQVRRKIPHEDFDRTTLENDIMLLQVVSLTTVSHVLKPIRLPPANFLLKPEATCSVAGWGRTGVHMVRPATKLQEVTLKMADDKQCELFFDSVHGLHGEHQNEAVIGDTGGPLVCNGFAQSIVSYGKDSGLPSAVYTRMSAFIPWLRKKMGRE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIF4</accession>
    <name>A0A452GIF4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Sodium/hydrogen exchanger</fullName>
        </recommendedName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="16">Plasma membrane Na(+)/H(+) antiporter. Exchanges intracellular H(+) ions for extracellular Na(+) in 1:1 stoichiometry, playing a key role in salt and fluid absorption and pH homeostasis. Major apical Na(+)/H(+) exchanger in kidney and intestine playing an important role in renal and intestine Na(+) absorption and blood pressure regulation.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>Na(+)(in) + H(+)(out) = Na(+)(out) + H(+)(in)</text>
            <dbReference type="Rhea" id="RHEA:29419"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Apical cell membrane</location>
            <topology evidence="2">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Early endosome membrane</location>
            <topology evidence="3">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Recycling endosome membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="18">Belongs to the monovalent cation:proton antiporter 1 (CPA1) transporter (TC 2.A.36) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIF4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001627.1">
        <property type="protein sequence ID" value="ENSGAGP00000001439.1"/>
        <property type="gene ID" value="ENSGAGG00000000884.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031526">
        <property type="term" value="C:brush border membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031901">
        <property type="term" value="C:early endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055038">
        <property type="term" value="C:recycling endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015386">
        <property type="term" value="F:potassium:proton antiporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015385">
        <property type="term" value="F:sodium:proton antiporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051453">
        <property type="term" value="P:regulation of intracellular pH"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098719">
        <property type="term" value="P:sodium ion import across plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.1330">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018422">
        <property type="entry name" value="Cation/H_exchanger_CPA1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006153">
        <property type="entry name" value="Cation/H_exchanger_TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018410">
        <property type="entry name" value="Na/H_exchanger_3/5"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004709">
        <property type="entry name" value="NaH_exchanger"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00840">
        <property type="entry name" value="b_cpa1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10110">
        <property type="entry name" value="SODIUM/HYDROGEN EXCHANGER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10110:SF90">
        <property type="entry name" value="SODIUM_HYDROGEN EXCHANGER 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00999">
        <property type="entry name" value="Na_H_Exchanger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01084">
        <property type="entry name" value="NAHEXCHNGR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01087">
        <property type="entry name" value="NAHEXCHNGR3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 18" id="KW-0050">Antiport</keyword>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="10" id="KW-0967">Endosome</keyword>
    <keyword evidence="13 18" id="KW-0406">Ion transport</keyword>
    <keyword evidence="14 20" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0732">Signal</keyword>
    <keyword evidence="12" id="KW-0915">Sodium</keyword>
    <keyword evidence="15 18" id="KW-0739">Sodium transport</keyword>
    <keyword evidence="8 18 20" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 20" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="4 18" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="7"/>
            <end position="27"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="39"/>
            <end position="56"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="68"/>
            <end position="91"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="103"/>
            <end position="125"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="132"/>
            <end position="156"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="176"/>
            <end position="200"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="260"/>
            <end position="281"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="293"/>
            <end position="316"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="328"/>
            <end position="349"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="361"/>
            <end position="381"/>
        </location>
    </feature>
    <feature type="domain" description="Cation/H+ exchanger transmembrane" evidence="21">
        <location>
            <begin position="9"/>
            <end position="387"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="589"/>
            <end position="688"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="701"/>
            <end position="725"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="19">
        <location>
            <begin position="602"/>
            <end position="617"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="643"/>
            <end position="673"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="674"/>
            <end position="688"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="19">
        <location>
            <begin position="707"/>
            <end position="717"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004195"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004424"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004520"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022449"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023053"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023201"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00045831"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00047524"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="RuleBase" id="RU003722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="Pfam" id="PF00999"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001439.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="748" mass="84152" checksum="B9FAF9C209454B7A" modified="2019-05-08" version="1">LGLAKTLVAIALLIVLGLFLGGIVWAADHIASFTLTPNVFFFYLLPPIVLDAGYFMPNKLFFGNLGTILLYAVIGTVWNAATTGMSLYGVYLSGIMGQLDIGLLDFLLFGSLIAAVDPVAVLAVFEEVHVNEVLFIIVFGESLLNDAVTVVLYNVFDSFVAMGADNVTGVDCVKGIVSFFVVSLGGTFVGIVFAFLLSLVTRFTKHVRIIEPGFVFVISYLSYLTAEMLSLSAILAITFCGVCCQKYVKANICDQSSTTVRYTMKMLASGSETIIFMFLGISAVNPDIWTWNTAFILLTLVFISVYRVIGVVIQTWVLNHYRVVQLEIIDQVVMSYGGLRGAVAFALVALLDEKKVKEKNLFVSTTIIVVFFTVIFQGLTIKPLVQWLKVKKSEHREPKLNEKLHGRAFDHILSAVEDISGQIGHNYLRDKWSNFDRKILSKVLMRRSAQKSRDQILNIFHELNLKDAISYVAEGERRGSLAFIRSSSTDNMVNVDFSTPRPSTVEASVSCLLREKASSVCLDMQALEQRRKSIRDTEDTVTHHTLQQYLYKPRQEHIHLYSRHEIIQNEDEKQDKEIFHRTMRKRLESFKSTKLGINQPKKLNKSHKRDRGQKRRNSNAIPNGKIPSENPAPDFTLKEKGNFSDTNNKPPVNSGYCSETSYEPVNSQDNSVDMQERLEESNPKENQEKKTLLNAQAHYRSAAARRVQSRRSFHRPKPLQLPKLQKLASFPSSCHIPPHSLEQEETQL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIF7</accession>
    <name>A0A452GIF7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Sodium/hydrogen exchanger</fullName>
        </recommendedName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="16">Plasma membrane Na(+)/H(+) antiporter. Exchanges intracellular H(+) ions for extracellular Na(+) in 1:1 stoichiometry, playing a key role in salt and fluid absorption and pH homeostasis. Major apical Na(+)/H(+) exchanger in kidney and intestine playing an important role in renal and intestine Na(+) absorption and blood pressure regulation.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>Na(+)(in) + H(+)(out) = Na(+)(out) + H(+)(in)</text>
            <dbReference type="Rhea" id="RHEA:29419"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Apical cell membrane</location>
            <topology evidence="2">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Early endosome membrane</location>
            <topology evidence="3">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Recycling endosome membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="18">Belongs to the monovalent cation:proton antiporter 1 (CPA1) transporter (TC 2.A.36) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIF7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001612.1">
        <property type="protein sequence ID" value="ENSGAGP00000001425.1"/>
        <property type="gene ID" value="ENSGAGG00000000884.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031526">
        <property type="term" value="C:brush border membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031901">
        <property type="term" value="C:early endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055038">
        <property type="term" value="C:recycling endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015386">
        <property type="term" value="F:potassium:proton antiporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015385">
        <property type="term" value="F:sodium:proton antiporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051453">
        <property type="term" value="P:regulation of intracellular pH"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098719">
        <property type="term" value="P:sodium ion import across plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.1330">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018422">
        <property type="entry name" value="Cation/H_exchanger_CPA1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006153">
        <property type="entry name" value="Cation/H_exchanger_TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018410">
        <property type="entry name" value="Na/H_exchanger_3/5"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004709">
        <property type="entry name" value="NaH_exchanger"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00840">
        <property type="entry name" value="b_cpa1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10110">
        <property type="entry name" value="SODIUM/HYDROGEN EXCHANGER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10110:SF90">
        <property type="entry name" value="SODIUM_HYDROGEN EXCHANGER 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00999">
        <property type="entry name" value="Na_H_Exchanger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01084">
        <property type="entry name" value="NAHEXCHNGR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01087">
        <property type="entry name" value="NAHEXCHNGR3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 18" id="KW-0050">Antiport</keyword>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="10" id="KW-0967">Endosome</keyword>
    <keyword evidence="13 18" id="KW-0406">Ion transport</keyword>
    <keyword evidence="14 20" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0732">Signal</keyword>
    <keyword evidence="12" id="KW-0915">Sodium</keyword>
    <keyword evidence="15 18" id="KW-0739">Sodium transport</keyword>
    <keyword evidence="8 18 20" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 20" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="4 18" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="12"/>
            <end position="33"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="45"/>
            <end position="62"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="74"/>
            <end position="97"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="109"/>
            <end position="131"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="138"/>
            <end position="162"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="182"/>
            <end position="206"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="266"/>
            <end position="287"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="299"/>
            <end position="322"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="334"/>
            <end position="355"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="367"/>
            <end position="387"/>
        </location>
    </feature>
    <feature type="domain" description="Cation/H+ exchanger transmembrane" evidence="21">
        <location>
            <begin position="6"/>
            <end position="393"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="595"/>
            <end position="655"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="19">
        <location>
            <begin position="608"/>
            <end position="623"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004195"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004424"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004520"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022449"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023053"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023201"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00045831"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00047524"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="RuleBase" id="RU003722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="Pfam" id="PF00999"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001425.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="728" mass="82109" checksum="0B840977C9743C16" modified="2019-05-08" version="1">EVFHLSHKVTRVVPESALLIVLGLFLGGIVWAADHIASFTLTPNVFFFYLLPPIVLDAGYFMPNKLFFGNLGTILLYAVIGTVWNAATTGMSLYGVYLSGIMGQLDIGLLDFLLFGSLIAAVDPVAVLAVFEEVHVNEVLFIIVFGESLLNDAVTVVLYNVFDSFVAMGADNVTGVDCVKGIVSFFVVSLGGTFVGIVFAFLLSLVTRFTKHVRIIEPGFVFVISYLSYLTAEMLSLSAILAITFCGVCCQKYVKANICDQSSTTVRYTMKMLASGSETIIFMFLGISAVNPDIWTWNTAFILLTLVFISVYRVIGVVIQTWVLNHYRVVQLEIIDQVVMSYGGLRGAVAFALVALLDEKKVKEKNLFVSTTIIVVFFTVIFQGLTIKPLVQWLKVKKSEHREPKLNEKLHGRAFDHILSAVEDISGQIGHNYLRDKWSNFDRKILSKVLMRRSAQKSRDQILNIFHELNLKDAISYVAEGERRGSLAFIRSSSTDNMVNVDFSTPRPSTVEASVSCLLREKASSVCLDMQALEQRRKSIRDTEDTVTHHTLQQYLYKPRQEHIHLYSRHEIIQNEDEKQDKEIFHRTMRKRLESFKSTKLGINQPKKLNKSHKRDRGQKRRNSNAIPNGKIPSENPAPDFTLKEKGIDNPAFSADDDQSIYMKFPPWLTDEDSVVPSQRARVQLPYSPNNFRRLTPFRLSNKSVDSFLLADGPEERPRAFLPETTHM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIG1</accession>
    <name>A0A452GIG1_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="5">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000001617.1">
        <property type="protein sequence ID" value="ENSGAGP00000001430.1"/>
        <property type="gene ID" value="ENSGAGG00000000400.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019901">
        <property type="term" value="F:protein kinase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008104">
        <property type="term" value="P:intracellular protein localization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.200:FF:000010">
        <property type="entry name" value="neurobeachin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.200">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050865">
        <property type="entry name" value="BEACH_Domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031570">
        <property type="entry name" value="NBEA/BDCP_DUF4704"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046852">
        <property type="entry name" value="Neurobeachin_a-sol"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13743">
        <property type="entry name" value="BEIGE/BEACH-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13743:SF62">
        <property type="entry name" value="NEUROBEACHIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15787">
        <property type="entry name" value="DUF4704"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13385">
        <property type="entry name" value="Laminin_G_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20425">
        <property type="entry name" value="Neurobeachin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0853">WD repeat</keyword>
    <feature type="domain" description="Neurobeachin alpha-solenoid region" evidence="4">
        <location>
            <begin position="53"/>
            <end position="187"/>
        </location>
    </feature>
    <feature type="domain" description="DUF4704" evidence="3">
        <location>
            <begin position="434"/>
            <end position="913"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="937"/>
            <end position="960"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="946"/>
            <end position="959"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022574"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="Pfam" id="PF15787"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="Pfam" id="PF20425"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001430.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1789" mass="199915" checksum="98FD5D656BCA5EB4" modified="2019-05-08" version="1">ACARRPRYPVSRPASGSGAVVLPAGMINPSVPIRNIRMKFAVLIGLIQVGEVSNRDIVETVLNLLVGGEFDLEMNFIIQDAESITCMSELLEHCDVTCQAEIWSMFTAILRKSVRNLQTSTEVGLIEQVLLKMSTVDDMIADLLVDMLGVLASYSITVKELKLLFSMLRGENGIWPRHAIKLLSVLNQMPQRHGPDTFFNFPGCSAAAIALPPIAKWPYQNGFTLNTWFRMDPLNNINVDKDKPYLYCFRTSKGVGYSAHFVGNCLIVTSLKSKGKGFQHCVKYDFQPRKWYMISIVHIYNRWRNSEIRCYVNGQLVSYGDMAWHVNTNDSYDKCFLGSSETADANRVFCGQLGAVYVFTEALNPAQIFAIHQLGPGYKSTFKFKSESDIHLAEHHKQVLYDGKLASSIAFTYNAKATDAQLCLESSPKENPSIFVHSPHALMLQDVKAIVTHSIHSAIHSIGGIQVLFPLFAQLDNRQLNDSQMETTVCATLLAFLVELLKSSVAMQEQMLGGKGFLVIGYLLEKSSRVHITRAVLEQFLSFAKYLDGLSHGAPLLKQLCDHILFNPAIWIHTPAKVQLSLYTYLSAEFIGTATIYTTIRRVGTVLQLMHTLKYYYWVVNPADSSGITPKGLEGPRPSQKEIISLRAFMLLFLKQLILKDRGVKEDELQSILNYLLTMHEDENIHDVLQLLVALMSEHPASMIPAFDQRNGIRVIYKLLASKSESIWVQALKVLGYFLKHLGHKRKVEIMHTHSLFTLLGERLMLHTNTVTITTYNTLYEILTEQVCTQVVHKPHPEPDSTVKIQNPMILKVVATLLKNSTPSAELMEVRRLFLSDMIKLFSNSRENRRCLLQCSVWQDWMFSLGYINPKNSEEQKITEMVYNIFRILLYHAIKYEWGGWRVWVDTLSIAHSKVTYEAHKEYLAKMYEEYQRQEEENIKKGKKGNVSTISGLSSQTTGAKGGLEIREIEDLSQSQSPESETDYPVSTDTRDLLMATKVSDEVLGSSDRPGGGVHVEVHDLLVDIKAEKVEATEVKLDDMDLSPETLVTGENGALVEVESLLDNVYCAAVEKLQNNVHGSVGIIKKSEEKDNGPLITLADEKDEPSTNNTSFLFDKIPSQEEKLLPELSSNHITIPNVQETQMHLDLRVDLGFRGMPMTEEQRRQFSPGPRTTMFRIPEFKWSPMHQRLLTDLLFALETDVHVWRSHSTKSVMDFVNSNENIIFVHNTIHLISQMVDNIIIACGGILPLLSAATSPTTELENIEVTQGMSAETAVTFLSRLMAMVDVLVFASSLNFSEIEAEKNMSSGGLMRQCLRLVCCVAVRNCLECRQRQRERVNKSSLISSKTQETLQGVTATSPLENVPGNLSPIKDPDRLLQDVDINRLRAVVFRDVDDSKQAQFLALAVVYFISVLMVSKYRDILEPQRETARCGSQADLGDSTLLTLLPHFDLSPSVLLESSIPHTDSGIGEEQMPSILNGTDLETSAGPDAMSELFSTLSSEVKKSQESLTESPSEILKPAPSISSISQSKGMNVKEILKSLVAAPIEIAECGPDPIPYPDPALKREAQSILPMQFHSFDRYMAELIILYCCNAEIWSIYLLTCAVDSGSSSSSSSSSFVNGATSKNLPAVQTVAPMPEDSAENMSITAKLERALEKVAPLLREIFVDFAPFLSRTLLGSHGQELLIEGLVCMKSSTSVVELVMLLCSQEWQNSIQKNAGLADGHKQTSEKLPTRKVSRRLALAMMVISRWKKQKLAPGSSGRLVLGERFSGGTRQRTLDMIMSGKRCLR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIH2</accession>
    <name>A0A452GIH2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Sodium/hydrogen exchanger</fullName>
        </recommendedName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="16">Plasma membrane Na(+)/H(+) antiporter. Exchanges intracellular H(+) ions for extracellular Na(+) in 1:1 stoichiometry, playing a key role in salt and fluid absorption and pH homeostasis. Major apical Na(+)/H(+) exchanger in kidney and intestine playing an important role in renal and intestine Na(+) absorption and blood pressure regulation.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>Na(+)(in) + H(+)(out) = Na(+)(out) + H(+)(in)</text>
            <dbReference type="Rhea" id="RHEA:29419"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Apical cell membrane</location>
            <topology evidence="2">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Early endosome membrane</location>
            <topology evidence="3">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Recycling endosome membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="18">Belongs to the monovalent cation:proton antiporter 1 (CPA1) transporter (TC 2.A.36) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIH2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001645.1">
        <property type="protein sequence ID" value="ENSGAGP00000001451.1"/>
        <property type="gene ID" value="ENSGAGG00000000884.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031526">
        <property type="term" value="C:brush border membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031901">
        <property type="term" value="C:early endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055038">
        <property type="term" value="C:recycling endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015386">
        <property type="term" value="F:potassium:proton antiporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015385">
        <property type="term" value="F:sodium:proton antiporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051453">
        <property type="term" value="P:regulation of intracellular pH"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098719">
        <property type="term" value="P:sodium ion import across plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.1330">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018422">
        <property type="entry name" value="Cation/H_exchanger_CPA1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006153">
        <property type="entry name" value="Cation/H_exchanger_TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018410">
        <property type="entry name" value="Na/H_exchanger_3/5"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004709">
        <property type="entry name" value="NaH_exchanger"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00840">
        <property type="entry name" value="b_cpa1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10110">
        <property type="entry name" value="SODIUM/HYDROGEN EXCHANGER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10110:SF90">
        <property type="entry name" value="SODIUM_HYDROGEN EXCHANGER 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00999">
        <property type="entry name" value="Na_H_Exchanger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01084">
        <property type="entry name" value="NAHEXCHNGR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01087">
        <property type="entry name" value="NAHEXCHNGR3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 18" id="KW-0050">Antiport</keyword>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="10" id="KW-0967">Endosome</keyword>
    <keyword evidence="13 18" id="KW-0406">Ion transport</keyword>
    <keyword evidence="14 20" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0732">Signal</keyword>
    <keyword evidence="12" id="KW-0915">Sodium</keyword>
    <keyword evidence="15 18" id="KW-0739">Sodium transport</keyword>
    <keyword evidence="8 18 20" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 20" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="4 18" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="21"/>
            <end position="39"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="51"/>
            <end position="68"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="80"/>
            <end position="103"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="115"/>
            <end position="137"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="144"/>
            <end position="168"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="188"/>
            <end position="212"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="305"/>
            <end position="328"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="340"/>
            <end position="361"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="373"/>
            <end position="393"/>
        </location>
    </feature>
    <feature type="domain" description="Cation/H+ exchanger transmembrane" evidence="21">
        <location>
            <begin position="4"/>
            <end position="399"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="601"/>
            <end position="661"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="19">
        <location>
            <begin position="614"/>
            <end position="629"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004195"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004424"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004520"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022449"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023053"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023201"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00045831"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00047524"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="RuleBase" id="RU003722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="Pfam" id="PF00999"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001451.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="734" mass="82768" checksum="359BDF0ED757EDDE" modified="2019-05-08" version="1">SLILSLLFFHLSHKVTRVVPESALLIVLGLFLGGIVWAADHIASFTLTPNVFFFYLLPPIVLDAGYFMPNKLFFGNLGTILLYAVIGTVWNAATTGMSLYGVYLSGIMGQLDIGLLDFLLFGSLIAAVDPVAVLAVFEEVHVNEVLFIIVFGESLLNDAVTVVLYNVFDSFVAMGADNVTGVDCVKGIVSFFVVSLGGTFVGIVFAFLLSLVTRFTKHVRIIEPGFVFVISYLSYLTAEMLSLSAILAITFCGVCCQKYVKANICDQSSTTVRYTMKMLASGSETIIFMFLGISAVNPDIWTWNTAFILLTLVFISVYRVIGVVIQTWVLNHYRVVQLEIIDQVVMSYGGLRGAVAFALVALLDEKKVKEKNLFVSTTIIVVFFTVIFQGLTIKPLVQWLKVKKSEHREPKLNEKLHGRAFDHILSAVEDISGQIGHNYLRDKWSNFDRKILSKVLMRRSAQKSRDQILNIFHELNLKDAISYVAEGERRGSLAFIRSSSTDNMVNVDFSTPRPSTVEASVSCLLREKASSVCLDMQALEQRRKSIRDTEDTVTHHTLQQYLYKPRQEHIHLYSRHEIIQNEDEKQDKEIFHRTMRKRLESFKSTKLGINQPKKLNKSHKRDRGQKRRNSNAIPNGKIPSENPAPDFTLKEKGIDNPAFSADDDQSIYMKFPPWLTDEDSVVPSQRARVQLPYSPNNFRRLTPFRLSNKSVDSFLLADGPEERPRAFLPETTHM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIH4</accession>
    <name>A0A452GIH4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="30">Thromboxane-A synthase</fullName>
            <ecNumber evidence="7">4.2.1.152</ecNumber>
            <ecNumber evidence="29">5.3.99.5</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="31">Cytochrome P450 5A1</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="25">Hydroperoxy icosatetraenoate dehydratase</fullName>
        </alternativeName>
    </protein>
    <organism evidence="36 37">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="37" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="36" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="36" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="32">Catalyzes the conversion of prostaglandin H2 (PGH2) to thromboxane A2 (TXA2), a potent inducer of blood vessel constriction and platelet aggregation. Also cleaves PGH2 to 12-hydroxy-heptadecatrienoicacid (12-HHT) and malondialdehyde, which is known to act as a mediator of DNA damage. 12-HHT and malondialdehyde are formed stoichiometrically in the same amounts as TXA2. Additionally, displays dehydratase activity, toward (15S)-hydroperoxy-(5Z,8Z,11Z,13E)-eicosatetraenoate (15(S)-HPETE) producing 15-KETE and 15-HETE.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="26">
            <text>(15S)-hydroperoxy-(5Z,8Z,11Z,13E)-eicosatetraenoate + AH2 = (15S)-hydroxy-(5Z,8Z,11Z,13E)-eicosatetraenoate + A + H2O</text>
            <dbReference type="Rhea" id="RHEA:48856"/>
            <dbReference type="ChEBI" id="CHEBI:13193"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:17499"/>
            <dbReference type="ChEBI" id="CHEBI:57409"/>
            <dbReference type="ChEBI" id="CHEBI:57446"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="26">
            <dbReference type="Rhea" id="RHEA:48857"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>(15S)-hydroperoxy-(5Z,8Z,11Z,13E)-eicosatetraenoate = 15-oxo-(5Z,8Z,11Z,13E)-eicosatetraenoate + H2O</text>
            <dbReference type="Rhea" id="RHEA:48636"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:57410"/>
            <dbReference type="ChEBI" id="CHEBI:57446"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="1">
            <dbReference type="Rhea" id="RHEA:48637"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="2">
            <text>a hydroperoxyeicosatetraenoate = an oxoeicosatetraenoate + H2O</text>
            <dbReference type="Rhea" id="RHEA:55556"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:59720"/>
            <dbReference type="ChEBI" id="CHEBI:131859"/>
            <dbReference type="EC" id="4.2.1.152"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="2">
            <dbReference type="Rhea" id="RHEA:55557"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="27">
            <text>prostaglandin H2 = (12S)-hydroxy-(5Z,8E,10E)-heptadecatrienoate + malonaldehyde</text>
            <dbReference type="Rhea" id="RHEA:48644"/>
            <dbReference type="ChEBI" id="CHEBI:57405"/>
            <dbReference type="ChEBI" id="CHEBI:90694"/>
            <dbReference type="ChEBI" id="CHEBI:566274"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="28">
            <text>prostaglandin H2 = thromboxane A2</text>
            <dbReference type="Rhea" id="RHEA:17137"/>
            <dbReference type="ChEBI" id="CHEBI:57405"/>
            <dbReference type="ChEBI" id="CHEBI:57445"/>
            <dbReference type="EC" id="5.3.99.5"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="28">
            <dbReference type="Rhea" id="RHEA:17138"/>
        </physiologicalReaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="3 33">
            <name>heme</name>
            <dbReference type="ChEBI" id="CHEBI:30413"/>
        </cofactor>
    </comment>
    <comment type="subunit">
        <text evidence="6">Monomer.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="4">Endoplasmic reticulum membrane</location>
            <topology evidence="4">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5 34">Belongs to the cytochrome P450 family.</text>
    </comment>
    <dbReference type="EC" id="4.2.1.152" evidence="7"/>
    <dbReference type="EC" id="5.3.99.5" evidence="29"/>
    <dbReference type="AlphaFoldDB" id="A0A452GIH4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001565.1">
        <property type="protein sequence ID" value="ENSGAGP00000001389.1"/>
        <property type="gene ID" value="ENSGAGG00000001072.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0020037">
        <property type="term" value="F:heme binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0106256">
        <property type="term" value="F:hydroperoxy icosatetraenoate dehydratase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005506">
        <property type="term" value="F:iron ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016705">
        <property type="term" value="F:oxidoreductase activity, acting on paired donors, with incorporation or reduction of molecular oxygen"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008395">
        <property type="term" value="F:steroid hydroxylase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004796">
        <property type="term" value="F:thromboxane-A synthase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001516">
        <property type="term" value="P:prostaglandin biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.630.10:FF:000003">
        <property type="entry name" value="cytochrome P450 3A12-like isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.630.10">
        <property type="entry name" value="Cytochrome P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001128">
        <property type="entry name" value="Cyt_P450"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017972">
        <property type="entry name" value="Cyt_P450_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002401">
        <property type="entry name" value="Cyt_P450_E_grp-I"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036396">
        <property type="entry name" value="Cyt_P450_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050705">
        <property type="entry name" value="Cytochrome_P450_3A"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24302:SF47">
        <property type="entry name" value="CYTOCHROME P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24302">
        <property type="entry name" value="CYTOCHROME P450 FAMILY 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00067">
        <property type="entry name" value="p450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00463">
        <property type="entry name" value="EP450I"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00385">
        <property type="entry name" value="P450"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48264">
        <property type="entry name" value="Cytochrome P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00086">
        <property type="entry name" value="CYTOCHROME_P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="14" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="22" id="KW-0275">Fatty acid biosynthesis</keyword>
    <keyword evidence="15" id="KW-0276">Fatty acid metabolism</keyword>
    <keyword evidence="11 33 34" id="KW-0349">Heme</keyword>
    <keyword evidence="18 33 34" id="KW-0408">Iron</keyword>
    <keyword evidence="23" id="KW-0413">Isomerase</keyword>
    <keyword evidence="9" id="KW-0444">Lipid biosynthesis</keyword>
    <keyword evidence="20" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="24" id="KW-0456">Lyase</keyword>
    <keyword evidence="21 35" id="KW-0472">Membrane</keyword>
    <keyword evidence="13 33 34" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="19 34" id="KW-0503">Monooxygenase</keyword>
    <keyword evidence="17 34" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="10" id="KW-0643">Prostaglandin biosynthesis</keyword>
    <keyword evidence="8" id="KW-0644">Prostaglandin metabolism</keyword>
    <keyword evidence="37" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="12 35" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="16 35" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="35">
        <location>
            <begin position="199"/>
            <end position="217"/>
        </location>
    </feature>
    <feature type="binding site" description="axial binding residue" evidence="33">
        <location>
            <position position="456"/>
        </location>
        <ligand>
            <name>heme</name>
            <dbReference type="ChEBI" id="CHEBI:30413"/>
        </ligand>
        <ligandPart>
            <name>Fe</name>
            <dbReference type="ChEBI" id="CHEBI:18248"/>
        </ligandPart>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001143"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00001719"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00001971"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004477"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00010617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00011245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00013084"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022501"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022516"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022585"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022832"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023033"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00023160"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00023235"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00023239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00033404"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00036380"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="ARBA" id="ARBA00036424"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="ARBA" id="ARBA00036475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="ARBA" id="ARBA00038872"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="ARBA" id="ARBA00040834"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="31">
        <source>
            <dbReference type="ARBA" id="ARBA00042726"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="32">
        <source>
            <dbReference type="ARBA" id="ARBA00054825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="33">
        <source>
            <dbReference type="PIRSR" id="PIRSR602401-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="34">
        <source>
            <dbReference type="RuleBase" id="RU000461"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="35">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="36">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001389.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="37">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="510" mass="58395" checksum="8E63CCEBB7444708" modified="2019-05-08" version="1">IFHLGLFKYSTSSFSRLKRVGIRHPAPLPFIGNLLFFRQGFWENHTKLITEYGPVCGYYIGRRMYVLVSEPEMIKHILVKDFSNFTNRMTPGLVTKPVIDSILCLRDQRWKEVRSILTPAFSSAKMKEMTPLINQACDVLLSNLKVYADSGKAFDIQRSYGCFTLDVVASVAFGTHVDSQKNPDDTFVKNCKKFFEFSMLRPILILIIAFPFLMIPLSRILPNKRRDEMNGFFIKAIKNMIALRDQQDANERRRDFLQLILDARSSTNYISVEHFDIVNQADVSIQDNKPSVDKALPKKVQKTLSEDEIVGQAFIFLIGGYETTTNALSFATYLLATNPECQEKLLQEIDEFSGKHDVPDYQNIQELPYLDMVIAETLRMYPPAFRFTRETSKDCVVMGQHIPAGAVIEIAVGPLHYNPEFWPEPEKFIPERFTMEAKLQQHPFAYLPFGAGPRSCIGMRMALLEIKMTLLRILQKFRFETSPKTQIPLQLTSVSSLAPKNGVYVKIVPR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIH6</accession>
    <name>A0A452GIH6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Opioid growth factor receptor (OGFr) conserved domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the opioid growth factor receptor family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIH6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001643.1">
        <property type="protein sequence ID" value="ENSGAGP00000001449.1"/>
        <property type="gene ID" value="ENSGAGG00000001133.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140625">
        <property type="term" value="F:opioid growth factor receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006757">
        <property type="entry name" value="OGF_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039574">
        <property type="entry name" value="OGFr"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14015">
        <property type="entry name" value="OPIOID GROWTH FACTOR RECEPTOR OGFR ZETA-TYPE OPIOID RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14015:SF0">
        <property type="entry name" value="OPIOID GROWTH FACTOR RECEPTOR-LIKE PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04664">
        <property type="entry name" value="OGFr_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Opioid growth factor receptor (OGFr) conserved" evidence="3">
        <location>
            <begin position="87"/>
            <end position="290"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1"/>
            <end position="52"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="281"/>
            <end position="457"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="2">
        <location>
            <begin position="15"/>
            <end position="31"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="32"/>
            <end position="48"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="303"/>
            <end position="312"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="323"/>
            <end position="333"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="334"/>
            <end position="347"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="348"/>
            <end position="377"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="378"/>
            <end position="404"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="405"/>
            <end position="417"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="424"/>
            <end position="438"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00010365"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="Pfam" id="PF04664"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001449.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="457" mass="52781" checksum="85644C6721245242" modified="2019-05-08" version="1">MGSLLSGVSVKEPATVEDCDSTWETDSEPELQDPGGDQQQQQEAIGGEEPIDDATELTTKPKRSFYAARDLYKYRHQYPNFKDLRYQNDLCNLRFYKNKIPFKPDGVYIEEVLNKWKGDYEKLEHNHTYIQWLFPLREQGLNFYAKELTTYEIEEFKKTKEAIRRFLLAYKMMLEFFGIKLIDKTGNVARAANWQERFQHLNESQHNYLRITRILKSLGELGYESFKSPLVKFILHEALVEDTIPNIKQSALEYFVYTIRDRRERRQLLRFAQQRYTPSEHFIWGPPRKQKLEGNKTGKKPTSPVSSHNSYISKHKKWRDSKNTSITSSSSSKTAEEKKVELTRNGEKNNQCGTETNCEAAKQSNSEKNSDTESQNFKSEEVLSLSDKKENVSHSKKDKGKDGENLSQDCGNPTVTDTEPCDSENCSNNVSADVQDSLGQDKPPVESTIKNKDEIKS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIH8</accession>
    <name>A0A452GIH8_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="11">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1 7">Belongs to the peptidase M16 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIH8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001569.1">
        <property type="protein sequence ID" value="ENSGAGP00000001393.1"/>
        <property type="gene ID" value="ENSGAGG00000000237.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005782">
        <property type="term" value="C:peroxisomal matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004222">
        <property type="term" value="F:metalloendopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050435">
        <property type="term" value="P:amyloid-beta metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042447">
        <property type="term" value="P:hormone catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043171">
        <property type="term" value="P:peptide catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051603">
        <property type="term" value="P:proteolysis involved in protein catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.830.10:FF:000004">
        <property type="entry name" value="Putative insulin-degrading enzyme"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.830.10:FF:000006">
        <property type="entry name" value="Putative insulin-degrading enzyme"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.830.10">
        <property type="entry name" value="Metalloenzyme, LuxS/M16 peptidase-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011249">
        <property type="entry name" value="Metalloenz_LuxS/M16"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011765">
        <property type="entry name" value="Pept_M16_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001431">
        <property type="entry name" value="Pept_M16_Zn_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050626">
        <property type="entry name" value="Peptidase_M16"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007863">
        <property type="entry name" value="Peptidase_M16_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032632">
        <property type="entry name" value="Peptidase_M16_M"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43690:SF18">
        <property type="entry name" value="INSULIN-DEGRADING ENZYME-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43690">
        <property type="entry name" value="NARDILYSIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00675">
        <property type="entry name" value="Peptidase_M16"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05193">
        <property type="entry name" value="Peptidase_M16_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16187">
        <property type="entry name" value="Peptidase_M16_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF63411">
        <property type="entry name" value="LuxS/MPP-like metallohydrolase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00143">
        <property type="entry name" value="INSULINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="6" id="KW-0482">Metalloprotease</keyword>
    <keyword evidence="2" id="KW-0645">Protease</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="Peptidase M16 N-terminal" evidence="8">
        <location>
            <begin position="69"/>
            <end position="203"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase M16 C-terminal" evidence="9">
        <location>
            <begin position="230"/>
            <end position="408"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase M16 middle/third" evidence="10">
        <location>
            <begin position="414"/>
            <end position="544"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007261"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023049"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU004447"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF00675"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF05193"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="Pfam" id="PF16187"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001393.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="565" mass="64837" checksum="39965BEEBD503272" modified="2019-05-08" version="1">LKYFSIGLLPAQGARGKQQWGPLPRVRSNTHNKMSNPAIKRLANKIIKSPEDKREYRGLELENGIKAMLICDPTTDKSSAALDVHIGSLSDPLNIAGLSHFCEHMLFLGTKKYPKENEYSQFLSEHAGSSNAFTSGEHTNYYFDVSHEHLEGALDRFAQFFLCPLFDESCKEREVNAVDSEHEKNLMNDAWRLFQLEKATGNPNHPFSKFGTGNKFTLETRPTKEGIDVRQELLKFHSTYYSSNLMAICVLGRESLDELTGLVVKLFSEVENKNVPLPEFPEHPFQEQHLRQLYKVVPIKDIRNLYVTFPIPDLQKYYKSNPGHYLGHLIGHEGPGSLLSELKAKGWVNTLVGGQKEGARGFMFFIINVDLTEEGLLHVEDIILHMFQYIQKLRADGPQEWVFQECKDLNAVAFRFKDKERPRGYTSKLGGILHYYPIEEVLAAEYLLEEFRPDLIEMVLDKLRPENVRVAIVSKSFEGKTDRTEEWYGTQYKQEAISDEVIQKWQNADLNGKFKLPMKNEFIPTNFEILPLEKDAPQYPALIKVRICITYIKWKLCQPASLLEG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GII3</accession>
    <name>A0A452GII3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">non-specific serine/threonine protein kinase</fullName>
            <ecNumber evidence="2">2.7.11.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the protein kinase superfamily. STE Ser/Thr protein kinase family. STE20 subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452GII3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001578.1">
        <property type="protein sequence ID" value="ENSGAGP00000001398.1"/>
        <property type="gene ID" value="ENSGAGG00000001107.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd06612">
        <property type="entry name" value="STKc_MST1_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000075">
        <property type="entry name" value="Serine/threonine-protein kinase 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000410">
        <property type="entry name" value="Serine/threonine-protein kinase 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050629">
        <property type="entry name" value="STE20/SPS1-PAK"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48012:SF2">
        <property type="entry name" value="STERILE20-LIKE KINASE, ISOFORM B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48012">
        <property type="entry name" value="STERILE20-LIKE KINASE, ISOFORM B-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000654">
        <property type="entry name" value="Integrin-linked_kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6 7" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="5" id="KW-0418">Kinase</keyword>
    <keyword evidence="4 7" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="5" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Protein kinase" evidence="8">
        <location>
            <begin position="23"/>
            <end position="274"/>
        </location>
    </feature>
    <feature type="binding site" evidence="7">
        <location>
            <position position="52"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00008874"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00012513"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001398.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="307" mass="34633" checksum="07A590ABAC85B2B9" modified="2019-05-08" version="1">PIEAAPLKKLSEDSLTKQPEEVFDVLEKLGEGSYGSVFKAIHKESGQVVAIKQVPVESDLQEIIKEISIMQQCDSPYVVKYYGSYFKNTDLWIVMEYCGAGSVSDIIRLRNKTLTEDEIATILKSTLKGLEYLHFMRKIHRDIKAGNILLNTEGHAKLADFGVAGQLTDTMAKRNTVIGTPFWMAPEVIQEIGYNCVADIWSLGITSIEMAEGKPPYADIHPMRAIFMIPTNPPPTFRKPELWSDEFTDFVKKCLVKNPEQRATATQLLQHTFIKNAKPVSILRDLITEAMEIKSLYSDEQRSIPFC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GII4</accession>
    <name>A0A452GII4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Rho GTPase-activating protein 39</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GII4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001684.1">
        <property type="protein sequence ID" value="ENSGAGP00000001477.1"/>
        <property type="gene ID" value="ENSGAGG00000001177.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
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        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd04389">
        <property type="entry name" value="RhoGAP_KIAA1688"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.70.10:FF:000022">
        <property type="entry name" value="Rho GTPase activating protein 39"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.555.10:FF:000011">
        <property type="entry name" value="Rho GTPase-activating protein 39"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.530:FF:000003">
        <property type="entry name" value="Rho GTPase-activating protein 39"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.20.70.10">
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="MyTH4 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.555.10">
        <property type="entry name" value="Rho GTPase activation protein"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="MyTH4_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038185">
        <property type="entry name" value="MyTH4_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000198">
        <property type="entry name" value="RhoGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001202">
        <property type="entry name" value="WW_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036020">
        <property type="entry name" value="WW_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45876">
        <property type="entry name" value="FI04035P"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45876:SF1">
        <property type="entry name" value="RHO GTPASE-ACTIVATING PROTEIN 39"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00784">
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="MyTH4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00324">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00456">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
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    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0007">Acetylation</keyword>
    <keyword evidence="2" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="6" id="KW-0539">Nucleus</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
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        <location>
            <begin position="22"/>
            <end position="56"/>
        </location>
    </feature>
    <feature type="domain" description="MyTH4" evidence="11">
        <location>
            <begin position="696"/>
            <end position="853"/>
        </location>
    </feature>
    <feature type="domain" description="Rho-GAP" evidence="10">
        <location>
            <begin position="864"/>
            <end position="1052"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="70"/>
            <end position="138"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
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            <begin position="184"/>
            <end position="223"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
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            <end position="282"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="300"/>
            <end position="337"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="375"/>
            <end position="401"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="420"/>
            <end position="485"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="493"/>
            <end position="512"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="545"/>
            <end position="573"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="70"/>
            <end position="91"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="112"/>
            <end position="121"/>
        </location>
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            <end position="199"/>
        </location>
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        </location>
    </feature>
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        <location>
            <begin position="430"/>
            <end position="459"/>
        </location>
    </feature>
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        </location>
    </feature>
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        </source>
    </evidence>
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        </source>
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        </source>
    </evidence>
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            <dbReference type="Ensembl" id="ENSGAGP00000001477.1"/>
        </source>
    </evidence>
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    </evidence>
    <sequence length="1057" mass="120458" checksum="D75EA7F0E6437E2B" modified="2019-05-08" version="1">MYANLITGECVWDPPAGVRVKRTSENQWWELFDPNTSRFYYYNATTQRTVWHRPQNCDIIPLAKLQTLKQNTESPRASTENSPGRSSNVSREGSTSSSLEQELEGSEKVQDQTRSNRQSAQFAVVKEEPESSSPPGVFEKEYEIYRDYNAEGQLLHYRTSSLRWNTGTKERLLIKVTDREPSFLMHQGNGYTTDSQQGIRSRRPSGGQQSPNLQTFAPDTDNSVFFPERKQSPFLKRADHVGSCSPLLGRGDSFCSPLQSQHRKHSSESQPSSPRYAYEPPLYEEPPMEYQAPIYDEPPVDMQYEASGNYKAGSPQKSPIRKPHPFLQSPKQGPNSPYQQLVLTKQKCPERFMSLEYSPAGKEYVRQLVYVEQSGSSPKLKTGLRHKYSPNPIGGSYSLQHSPCLVRDQRLEIKSGDYSSMEGADFRHSQPVSQASQGEDSMSWSSQQDTMSSTGYSPSTRKRKSRKPSLSHDPNTSSTDGSGDREALNEQLMAEERPMPSTYRPQMNRTESKSIEAEAVRGFPEPFLAQARLAWEAQQAHYHMKQRSSWDSQKDGSGYESDGALPLPMPGPVVRAFSEDEALAQQESKHWKRNTFEKLGFPQILLEKSVSVQTNLASPEPYLHPSQSEDLGACAQFESSRQNRNMMPSASCVFPTFTLRKPSSETDIENWASKHFNKHTQGLFRRKVSIANMLAWSSESIKKPMIMTTDRNVKKEACEIFKLIQMYMGDRRAKADQLSVALEIATKGWSMQGLRDELYIQLCRQTTENFRYESLARGWELMAICLAFFPPTPKFHSYLEGYIYRHMDPVNDTKGVAISTYAKYCYNKLQKAALTGAKKGLKKPNIEEIRHAKNAVFSPSMFGSSLQEIMNMQKERYPDRQLPWVQTRLSEEVLALNGDQTEGIFRVPGDIDEVNALKLQVDQWKIPTGLEDPHVPASLLKLWYRELEEPLIPHEFYEQCITHYENPEAAIAVIHSLPRINKMVLCYLIRFLQVFVQPANVAVTKMDVNNLAMVMAPNCLRCQSDDPRIIFENTRKEMSFIRVLIQHLDTSFMEGVL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GII5</accession>
    <name>A0A452GII5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">UBC core domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="4">Belongs to the ubiquitin-conjugating enzyme family.</text>
    </comment>
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        <property type="protein sequence ID" value="ENSGAGP00000001399.1"/>
        <property type="gene ID" value="ENSGAGG00000001114.1"/>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
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        <property type="project" value="UniProtKB-KW"/>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Ubiquitin Conjugating Enzyme"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Ub_conjugating_enzyme"/>
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    </dbReference>
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        <property type="entry name" value="UBQ-conjugating_AS"/>
    </dbReference>
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        <property type="entry name" value="UBQ-conjugating_enzyme/RWD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24067">
        <property type="entry name" value="UBIQUITIN-CONJUGATING ENZYME E2"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="UQ_con"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="UBCc"/>
        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
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    </dbReference>
    <proteinExistence type="inferred from homology"/>
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    <keyword evidence="4" id="KW-0547">Nucleotide-binding</keyword>
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    <keyword evidence="1" id="KW-0808">Transferase</keyword>
    <keyword evidence="2 4" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="UBC core" evidence="6">
        <location>
            <begin position="4"/>
            <end position="112"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="1"/>
            <end position="30"/>
        </location>
    </feature>
    <feature type="active site" description="Glycyl thioester intermediate" evidence="3">
        <location>
            <position position="86"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="RuleBase" id="RU362109"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50127"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001399.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="112" mass="12707" checksum="371D88723F46BDB5" modified="2019-05-08" version="1">MSTPARRRLMRDFKRLQEDPPAGVSGAPSENNIMVWNAXTPEGTPFEDGTFKLTIEFTEEYPNKPPTVRFVSKMFHPNVYADGSICLDILQNRWSPTYDVSSILTSIQVVPF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GII6</accession>
    <name>A0A452GII6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Sodium/hydrogen exchanger</fullName>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="16">Plasma membrane Na(+)/H(+) antiporter. Exchanges intracellular H(+) ions for extracellular Na(+) in 1:1 stoichiometry, playing a key role in salt and fluid absorption and pH homeostasis. Major apical Na(+)/H(+) exchanger in kidney and intestine playing an important role in renal and intestine Na(+) absorption and blood pressure regulation.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>Na(+)(in) + H(+)(out) = Na(+)(out) + H(+)(in)</text>
            <dbReference type="Rhea" id="RHEA:29419"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Apical cell membrane</location>
            <topology evidence="2">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Early endosome membrane</location>
            <topology evidence="3">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Recycling endosome membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="18">Belongs to the monovalent cation:proton antiporter 1 (CPA1) transporter (TC 2.A.36) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GII6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001675.1">
        <property type="protein sequence ID" value="ENSGAGP00000001468.1"/>
        <property type="gene ID" value="ENSGAGG00000000884.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031526">
        <property type="term" value="C:brush border membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031901">
        <property type="term" value="C:early endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055038">
        <property type="term" value="C:recycling endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015386">
        <property type="term" value="F:potassium:proton antiporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015385">
        <property type="term" value="F:sodium:proton antiporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051453">
        <property type="term" value="P:regulation of intracellular pH"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098719">
        <property type="term" value="P:sodium ion import across plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.1330">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018422">
        <property type="entry name" value="Cation/H_exchanger_CPA1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006153">
        <property type="entry name" value="Cation/H_exchanger_TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018410">
        <property type="entry name" value="Na/H_exchanger_3/5"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004709">
        <property type="entry name" value="NaH_exchanger"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00840">
        <property type="entry name" value="b_cpa1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10110">
        <property type="entry name" value="SODIUM/HYDROGEN EXCHANGER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10110:SF90">
        <property type="entry name" value="SODIUM_HYDROGEN EXCHANGER 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00999">
        <property type="entry name" value="Na_H_Exchanger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01084">
        <property type="entry name" value="NAHEXCHNGR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01087">
        <property type="entry name" value="NAHEXCHNGR3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 18" id="KW-0050">Antiport</keyword>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="10" id="KW-0967">Endosome</keyword>
    <keyword evidence="13 18" id="KW-0406">Ion transport</keyword>
    <keyword evidence="14 19" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0732">Signal</keyword>
    <keyword evidence="12" id="KW-0915">Sodium</keyword>
    <keyword evidence="15 18" id="KW-0739">Sodium transport</keyword>
    <keyword evidence="8 18 19" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 19" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="4 18" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="25"/>
            <end position="46"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="58"/>
            <end position="75"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="87"/>
            <end position="110"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="122"/>
            <end position="144"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="151"/>
            <end position="175"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="195"/>
            <end position="219"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="279"/>
            <end position="300"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="312"/>
            <end position="335"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="347"/>
            <end position="368"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="380"/>
            <end position="400"/>
        </location>
    </feature>
    <feature type="domain" description="Cation/H+ exchanger transmembrane" evidence="20">
        <location>
            <begin position="12"/>
            <end position="406"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004195"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004424"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004520"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022449"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023053"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023201"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00045831"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00047524"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="RuleBase" id="RU003722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="Pfam" id="PF00999"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001468.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="762" mass="85949" checksum="566AAAAA812AF725" modified="2019-05-08" version="1">QSFSAKLYMSSAVLIFHLSHKVTRVVPESALLIVLGLFLGGIVWAADHIASFTLTPNVFFFYLLPPIVLDAGYFMPNKLFFGNLGTILLYAVIGTVWNAATTGMSLYGVYLSGIMGQLDIGLLDFLLFGSLIAAVDPVAVLAVFEEVHVNEVLFIIVFGESLLNDAVTVVLYNVFDSFVAMGADNVTGVDCVKGIVSFFVVSLGGTFVGIVFAFLLSLVTRFTKHVRIIEPGFVFVISYLSYLTAEMLSLSAILAITFCGVCCQKYVKANICDQSSTTVRYTMKMLASGSETIIFMFLGISAVNPDIWTWNTAFILLTLVFISVYRVIGVVIQTWVLNHYRVVQLEIIDQVVMSYGGLRGAVAFALVALLDEKKVKEKNLFVSTTIIVVFFTVIFQGLTIKPLVQWLKVKKSEHREPKLNEKLHGRAFDHILSAVEDISGQIGHNYLRDKWSNFDRKILSKVLMRRSAQKSRDQILNIFHELNLKDAISYVAEGERRGSLAFIRSSSTDNMVNVDFSTPRPSTVEASVSCLLREKASSVCLDMQALEQRRKSIRDTEDTVTHHTLQQYLYKPRQEHIHLYSRHEIIQNEDEKQDKEIFHRTMRKRLESFKSTKLGINQPKKLNKSHKRDRGQKRVRGLPFTQTKDVEFSDPEENYESLHMDKGVDFLANVTKQNCTDATSGIDNPAFSADDDQSIYMKFPPWLTDEDSVVPSQRARVQLPYSPNNFRRLTPFRLSNKSVDSFLLADGPEERPRAFLPETTHM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GII8</accession>
    <name>A0A452GII8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Nuclear receptor 2C2-associated protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the NR2C2AP family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GII8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001691.1">
        <property type="protein sequence ID" value="ENSGAGP00000001483.1"/>
        <property type="gene ID" value="ENSGAGG00000001192.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.260:FF:000070">
        <property type="entry name" value="Nuclear receptor 2C2-associated protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.260">
        <property type="entry name" value="Galactose-binding domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008979">
        <property type="entry name" value="Galactose-bd-like_sf"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49785">
        <property type="entry name" value="Galactose-binding domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0539">Nucleus</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009556"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00019956"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001483.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="246" mass="26979" checksum="32D77B08B69E22D2" modified="2019-05-08" version="1">MVPRVARLAPASSGKIMPNKSSAAAAPGADRGSGVSVTAAFDAVLHTRLIREQTENRSSEPCNSAAAPLPGALHRRLPLGALALNLISGVCWRLPWLERSDTFSSDMPTGPLICNKTVSRVSSVLNRDAKQFGKKHMFDGSEETCWNSDQGSSQWVMLEFPQPVKVSQLQIQFQGGFASQLCTLEGGRKGEELVKISDFYPEDINALQSFRTEETMLDKLKITFENSTDFFGRIIVYRLSVLGEKL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZC1</accession>
    <name>A0A452GZC1_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="11">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Lipid droplet</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the LDAF1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZC1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008706.1">
        <property type="protein sequence ID" value="ENSGAGP00000007541.1"/>
        <property type="gene ID" value="ENSGAGG00000006028.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005811">
        <property type="term" value="C:lipid droplet"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029709">
        <property type="entry name" value="LDAF1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14275:SF0">
        <property type="entry name" value="LIPID DROPLET ASSEMBLY FACTOR 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14275">
        <property type="entry name" value="PROMETHIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16015">
        <property type="entry name" value="Promethin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="4" id="KW-0551">Lipid droplet</keyword>
    <keyword evidence="8 10" id="KW-0472">Membrane</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 10" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 10" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="84"/>
            <end position="108"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="115"/>
            <end position="139"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="145"/>
            <end position="165"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="13"/>
            <end position="44"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="13"/>
            <end position="28"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004477"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004502"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022677"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007541.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="189" mass="19119" checksum="BBA456BF041D6799" modified="2019-05-08" version="1">MCLGGGLSPMTAAAPTNPAGGSGGSSSPAPQPVAAPVPGAASASQAEAGGGAQAAARRWNGESGSEVVSFMNSRVGQYLDDHPFVALSLLVFIAVSAIPVAFFLIFVISTTIVACMGVIVIEGVVVSVGGIVLLCALSGLGVLSLAVSGVLSICYIALSTLINYWHTPNYGRGGRNSYMQLYNALWLLM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZC5</accession>
    <name>A0A452GZC5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="19">RecQ-like DNA helicase BLM</fullName>
            <ecNumber evidence="17">5.6.2.4</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="18">DNA 3'-5' helicase BLM</fullName>
        </alternativeName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="16">
            <text>Couples ATP hydrolysis with the unwinding of duplex DNA by translocating in the 3'-5' direction.</text>
            <dbReference type="EC" id="5.6.2.4"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the helicase family. RecQ subfamily.</text>
    </comment>
    <dbReference type="EC" id="5.6.2.4" evidence="17"/>
    <dbReference type="AlphaFoldDB" id="A0A452GZC5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008564.1">
        <property type="protein sequence ID" value="ENSGAGP00000007415.1"/>
        <property type="gene ID" value="ENSGAGG00000005383.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005694">
        <property type="term" value="C:chromosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043138">
        <property type="term" value="F:3'-5' DNA helicase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009378">
        <property type="term" value="F:four-way junction helicase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016818">
        <property type="term" value="F:hydrolase activity, acting on acid anhydrides, in phosphorus-containing anhydrides"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006260">
        <property type="term" value="P:DNA replication"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000724">
        <property type="term" value="P:double-strand break repair via homologous recombination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000723">
        <property type="term" value="P:telomere maintenance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd18016">
        <property type="entry name" value="DEXHc_RecQ2_BLM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18794">
        <property type="entry name" value="SF2_C_RecQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000537">
        <property type="entry name" value="Bloom syndrome RecQ-like helicase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000340">
        <property type="entry name" value="Bloom syndrome, RecQ helicase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.80">
        <property type="entry name" value="HRDC domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.10">
        <property type="entry name" value="Winged helix-like DNA-binding domain superfamily/Winged helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012532">
        <property type="entry name" value="BDHCT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032439">
        <property type="entry name" value="BDHCT_assoc"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011545">
        <property type="entry name" value="DEAD/DEAH_box_helicase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002464">
        <property type="entry name" value="DNA/RNA_helicase_DEAH_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004589">
        <property type="entry name" value="DNA_helicase_ATP-dep_RecQ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014001">
        <property type="entry name" value="Helicase_ATP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001650">
        <property type="entry name" value="Helicase_C-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044876">
        <property type="entry name" value="HRDC_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032284">
        <property type="entry name" value="RecQ_Zn-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018982">
        <property type="entry name" value="RQC_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036388">
        <property type="entry name" value="WH-like_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036390">
        <property type="entry name" value="WH_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00614">
        <property type="entry name" value="recQ_fam"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13710">
        <property type="entry name" value="DNA HELICASE RECQ FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13710:SF153">
        <property type="entry name" value="RECQ-LIKE DNA HELICASE BLM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08072">
        <property type="entry name" value="BDHCT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16204">
        <property type="entry name" value="BDHCT_assoc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00270">
        <property type="entry name" value="DEAD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00271">
        <property type="entry name" value="Helicase_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16124">
        <property type="entry name" value="RecQ_Zn_bind"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09382">
        <property type="entry name" value="RQC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00487">
        <property type="entry name" value="DEXDc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00490">
        <property type="entry name" value="HELICc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00956">
        <property type="entry name" value="RQC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46785">
        <property type="entry name" value="Winged helix' DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00690">
        <property type="entry name" value="DEAH_ATP_HELICASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51192">
        <property type="entry name" value="HELICASE_ATP_BIND_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51194">
        <property type="entry name" value="HELICASE_CTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="11" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="7" id="KW-0227">DNA damage</keyword>
    <keyword evidence="13" id="KW-0234">DNA repair</keyword>
    <keyword evidence="4" id="KW-0235">DNA replication</keyword>
    <keyword evidence="12" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="9" id="KW-0347">Helicase</keyword>
    <keyword evidence="8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="14" id="KW-0413">Isomerase</keyword>
    <keyword evidence="5" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="6" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="15" id="KW-0539">Nucleus</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="Helicase ATP-binding" evidence="21">
        <location>
            <begin position="418"/>
            <end position="593"/>
        </location>
    </feature>
    <feature type="domain" description="Helicase C-terminal" evidence="22">
        <location>
            <begin position="619"/>
            <end position="766"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="1"/>
            <end position="37"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="20">
        <location>
            <begin position="1"/>
            <end position="16"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="20">
        <location>
            <begin position="17"/>
            <end position="29"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005446"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022705"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022806"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023235"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00034617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00034808"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00044542"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00073450"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS51192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS51194"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007415.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1003" mass="113052" checksum="519D2ED037655D67" modified="2019-05-08" version="1">FSSVSALSKKQYNGKQKITEGEKNSHSPKELAIGWDETPDYNGIEEDDYLDFIPPSPEEELLSSSSSVKNHLFSVMEDICKLVDSIPIHELKSLSCGNDLLQQRNLRRELLANHIDSNANKSDLGLVFASNWKSHPGHVTSMDDSIPVQSGQKYSLNSEGNSFRFTKVPSKLSVAVECQGNLNMSTFSTTKLQTLDTSGASDFSQYTALDTNITKNAGLSFSKMDVKERTSLSFHSEISSFNSSLCEKPRMEKDGSCFFSEMPVTSTTLKVENKSSSSLNNVGGDDSLGARDIDFDLDNFVLDDDFDERWDNPRCVSADEKTSTPLYQPIREGEPAKSLLKSFSTVKETSPIYNPVPKPNILLSAKSCSDPLLKKSPSKNPALARFRGCTFSHSNDMMRIFHKKFGLHHFRTNQLEAINASLLGEDCFILMPTGGGKSLCYQLPACLSSGVTIVISPLRSLILDQVEKLMTLDISATYLTGDRTDTEASKIYIQLSKKDPQIKLLYVTPEKVCSSGRLISALKNLYERDLLARFVIDEAHCVSQWGHDFRQDYKRLNMLRQKFPSVPMMALTATANPRVQKDILNQLEMLKPQVFTMSFNRHNLKYDVLPKKPKRVALDCLEWIKKYHPHDSGIIYCLSRQDCNKTADILQEKGLAALAYHAGLSDTVRDLVQRKWINQDGCQVICATIAFGMGIDKPDVRYVIHASLPKSVEGYYQESGRAGRDGEVSHCLLFYSYSDVTRLRRLIQMEKDGNSQTRKTHFDNLYSMVHYCENVHECRRIQLLAYFGETGFNLNFCKEHPEVICDNCCRKKDYKSRNVTDEVKSIIRFVQEHCGQMGRRNSGSGRYTLNMMIDIFLGTKSARIQSGIFGKGAAYSRHNVERLFRKLVLDKILEEELYITAAEQAVAYVTVGEKAQAVLNGYLQVEFYETENASSIRKQRASVTKMSQREEMVKKCLVELTDVCKTLGKVFDVHYFNIFNTATLKRISRIHPGVGLHWGGGTL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZC7</accession>
    <name>A0A452GZC7_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="7">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the TMEM45 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZC7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008713.1">
        <property type="protein sequence ID" value="ENSGAGP00000007546.1"/>
        <property type="gene ID" value="ENSGAGG00000005952.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006904">
        <property type="entry name" value="DUF716"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042127">
        <property type="entry name" value="TMEM45"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16007">
        <property type="entry name" value="EPIDIDYMAL MEMBRANE PROTEIN E9-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16007:SF21">
        <property type="entry name" value="TRANSMEMBRANE PROTEIN 45A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04819">
        <property type="entry name" value="DUF716"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 6" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 6" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 6" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="6"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="91"/>
            <end position="112"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="124"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="154"/>
            <end position="173"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="227"/>
            <end position="249"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006948"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007546.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="262" mass="30347" checksum="2DA152F8FA86A2C9" modified="2019-05-08" version="1">MGNFKGHALPGSFFLLFGLWWSVKYPLKYVCRKNKNACYLGSRAGFQRLEFIEGIIKAVFALIGMIAEQFVPDGPHLKLYNYEEKHWDHLMNWQHATMYLFYGISGLVDVVAHGTNALPVALDRMMLSLAVFIEGFLFYYHIHGRDMLDFHVHQLLLVAIFGAALCILLEVFFRGSIVLEMLRTSLCILQGSWLWQIGFVLYPPNGRLEWNQMDHNNMMFLTMCYCWHYAFAFLILAVNYTIVSCTLLLKNCLLSHSVCMKF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZC8</accession>
    <name>A0A452GZC8_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="10">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Lipid droplet</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the LDAF1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZC8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008699.1">
        <property type="protein sequence ID" value="ENSGAGP00000007534.1"/>
        <property type="gene ID" value="ENSGAGG00000006028.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005811">
        <property type="term" value="C:lipid droplet"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029709">
        <property type="entry name" value="LDAF1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14275:SF0">
        <property type="entry name" value="LIPID DROPLET ASSEMBLY FACTOR 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14275">
        <property type="entry name" value="PROMETHIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16015">
        <property type="entry name" value="Promethin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="4" id="KW-0551">Lipid droplet</keyword>
    <keyword evidence="8 9" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 9" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 9" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="42"/>
            <end position="66"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="73"/>
            <end position="97"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="103"/>
            <end position="123"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004477"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004502"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022677"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007534.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="158" mass="17139" checksum="2BEC3543399828A3" modified="2019-05-08" version="1">MSKEMQELQKQWHSMMQTIHTNSNVVSFMNSRVGQYLDDHPFVALSLLVFIAVSAIPVAFFLIFVISTTIVACMGVIVIEGVVVSVGGIVLLCALSGLGVLSLAVSGVLSICYIALSTLINYWHTPNSLVKKQEAIENSLLQSNPPVMDPSNKNTKSE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZD1</accession>
    <name>A0A452GZD1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">IRF tryptophan pentad repeat domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZD1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008813.1">
        <property type="protein sequence ID" value="ENSGAGP00000007636.1"/>
        <property type="gene ID" value="ENSGAGG00000006103.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002376">
        <property type="term" value="P:immune system process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045944">
        <property type="term" value="P:positive regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd00103">
        <property type="entry name" value="IRF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.10:FF:000041">
        <property type="entry name" value="Interferon regulatory factor 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.10">
        <property type="entry name" value="Winged helix-like DNA-binding domain superfamily/Winged helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019817">
        <property type="entry name" value="Interferon_reg_fac_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001346">
        <property type="entry name" value="Interferon_reg_fact_DNA-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036388">
        <property type="entry name" value="WH-like_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036390">
        <property type="entry name" value="WH_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11949">
        <property type="entry name" value="INTERFERON REGULATORY FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11949:SF7">
        <property type="entry name" value="INTERFERON REGULATORY FACTOR 8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00605">
        <property type="entry name" value="IRF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00267">
        <property type="entry name" value="INTFRNREGFCT"/>
    </dbReference>
    <dbReference type="SMART" id="SM00348">
        <property type="entry name" value="IRF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46785">
        <property type="entry name" value="Winged helix' DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00601">
        <property type="entry name" value="IRF_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51507">
        <property type="entry name" value="IRF_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0010">Activator</keyword>
    <keyword evidence="3" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="6" id="KW-0539">Nucleus</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0804">Transcription</keyword>
    <keyword evidence="2" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="IRF tryptophan pentad repeat" evidence="7">
        <location>
            <begin position="57"/>
            <end position="164"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS51507"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007636.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="188" mass="21837" checksum="AE014970FE94B9F8" modified="2019-05-08" version="1">MRGRGGGVPTFAYPLWPVSAVLSEPAARSRWVHLGEQRVGRKPILQTADRMCDRNGGRRLRQWLIEQIDSGMYPGLIWENEEKSMFRIPWKHAGKQDYNQEVDASIFKAWAVFKGKFKEGDKAEPATWKTRLRCALNKSPDFEEVTDRSQLDISEPYKVYRIVPEEEQKCKCPDKLSDLFHSHLPPEY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZD3</accession>
    <name>A0A452GZD3_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="7">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the TMEM45 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZD3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008705.1">
        <property type="protein sequence ID" value="ENSGAGP00000007540.1"/>
        <property type="gene ID" value="ENSGAGG00000005952.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006904">
        <property type="entry name" value="DUF716"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042127">
        <property type="entry name" value="TMEM45"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16007">
        <property type="entry name" value="EPIDIDYMAL MEMBRANE PROTEIN E9-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16007:SF21">
        <property type="entry name" value="TRANSMEMBRANE PROTEIN 45A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04819">
        <property type="entry name" value="DUF716"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 6" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 6" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 6" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="6"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="51"/>
            <end position="71"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="91"/>
            <end position="112"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="124"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="154"/>
            <end position="173"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="185"/>
            <end position="203"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="218"/>
            <end position="243"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006948"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007540.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="266" mass="30623" checksum="3735B48AA083F6A6" modified="2019-05-08" version="1">MGNFKGHALPGSFFLLFGLWWSVKYPLKYVCRKNKNACYLGSRAGFQRLEFIEGIIKAVFALIGMIAEQFVPDGPHLKLYNYEEKHWDHLMNWQHATMYLFYGISGLVDVVAHGTNALPVALDRMMLSLAVFIEGFLFYYHIHGRDMLDFHVHQLLLVAIFGAALCILLEVFFRGSIVLEMLRTSLCILQGSWLWQIGFVLYPPNGRLEWNQMDHNNMMFLTMCYCWHYAFAFLILAVNYTIVRLEGTSRGHRVQSPALLAGPNTV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZE1</accession>
    <name>A0A452GZE1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Rho GTPase-activating protein 7</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="15">Rho-type GTPase-activating protein 7</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="16">START domain-containing protein 12</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="20">START domain-containing protein 13</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="14">StAR-related lipid transfer protein 12</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="19">StAR-related lipid transfer protein 13</fullName>
        </alternativeName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="18">Homodimer. Interacts with TAX1BP1.</text>
    </comment>
    <comment type="subunit">
        <text evidence="17">Interacts with EF1A1, facilitates EF1A1 distribution to the membrane periphery and ruffles upon growth factor stimulation and suppresses cell migration. Interacts with tensin TNS1 (via N-terminus); the interaction is decreased by phosphorylation of TNS1. Interacts with TNS3 and PTEN; in resting cells, interacts with TNS3 (via C2 tensin-type domain) but, following growth factor stimulation, TNS3 and PTEN are phosphorylated which leads to weakened interaction with TNS3 and enhanced interaction with PTEN. Interacts (via C-terminus) with tensin TNS4 (via SH2 domain); the interaction is independent of tyrosine phosphorylation of DLC1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell junction</location>
            <location evidence="1">Focal adhesion</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Lipid droplet</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Mitochondrion membrane</location>
            <topology evidence="2">Peripheral membrane protein</topology>
            <orientation evidence="2">Cytoplasmic side</orientation>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZE1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008731.1">
        <property type="protein sequence ID" value="ENSGAGP00000007563.1"/>
        <property type="gene ID" value="ENSGAGG00000006046.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005925">
        <property type="term" value="C:focal adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005811">
        <property type="term" value="C:lipid droplet"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045121">
        <property type="term" value="C:membrane raft"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031966">
        <property type="term" value="C:mitochondrial membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008289">
        <property type="term" value="F:lipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030036">
        <property type="term" value="P:actin cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035023">
        <property type="term" value="P:regulation of Rho protein signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd04375">
        <property type="entry name" value="RhoGAP_DLC1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd09591">
        <property type="entry name" value="SAM_DLC1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.555.10:FF:000007">
        <property type="entry name" value="rho GTPase-activating protein 7 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.530.20:FF:000009">
        <property type="entry name" value="StAR related lipid transfer domain containing 13"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.2070:FF:000001">
        <property type="entry name" value="StAR-related lipid transfer domain-containing 13"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.2070">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.530.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.555.10">
        <property type="entry name" value="Rho GTPase activation protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000198">
        <property type="entry name" value="RhoGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001660">
        <property type="entry name" value="SAM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013761">
        <property type="entry name" value="SAM/pointed_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023393">
        <property type="entry name" value="START-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002913">
        <property type="entry name" value="START_lipid-bd_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12659:SF2">
        <property type="entry name" value="RHO GTPASE-ACTIVATING PROTEIN 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12659">
        <property type="entry name" value="RHO-TYPE GTPASE ACTIVATING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00620">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07647">
        <property type="entry name" value="SAM_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01852">
        <property type="entry name" value="START"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00324">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00234">
        <property type="entry name" value="START"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55961">
        <property type="entry name" value="Bet v1-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47769">
        <property type="entry name" value="SAM/Pointed domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50238">
        <property type="entry name" value="RHOGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50848">
        <property type="entry name" value="START"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="11" id="KW-0007">Acetylation</keyword>
    <keyword evidence="10" id="KW-0965">Cell junction</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="9" id="KW-0551">Lipid droplet</keyword>
    <keyword evidence="13" id="KW-0472">Membrane</keyword>
    <keyword evidence="12" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="8" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Rho-GAP" evidence="22">
        <location>
            <begin position="642"/>
            <end position="848"/>
        </location>
    </feature>
    <feature type="domain" description="START" evidence="23">
        <location>
            <begin position="878"/>
            <end position="1086"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="21">
        <location>
            <begin position="152"/>
            <end position="174"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="21">
        <location>
            <begin position="293"/>
            <end position="325"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="21">
        <location>
            <begin position="380"/>
            <end position="439"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="21">
        <location>
            <begin position="492"/>
            <end position="558"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="21">
        <location>
            <begin position="156"/>
            <end position="166"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="21">
        <location>
            <begin position="295"/>
            <end position="325"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="21">
        <location>
            <begin position="384"/>
            <end position="401"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="21">
        <location>
            <begin position="412"/>
            <end position="426"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="21">
        <location>
            <begin position="500"/>
            <end position="512"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="21">
        <location>
            <begin position="521"/>
            <end position="536"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004346"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004502"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00014465"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022677"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00030542"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00030675"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00032733"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00046839"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00065039"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00067490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00076865"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS50238"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS50848"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007563.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1086" mass="122137" checksum="1DA79DB11E1092E8" modified="2019-05-08" version="1">MCQAPPETMILTQIEAKEACDWLRAAGFPQYAQLYEDLLFPIDITLVKREHDFLDRDAIEALCRRLNTLNKCAVMKLEISPHRKRSEDSDEDEPCAISGKWTFQRDSKRWSRLEEFDVFPPKQDLNPSSPEAPLLTNAASHESVLTDLSERQEVASIHSTSSTSSHQVTLQSEATTTRTNSVVSVCSSGHFITNDDSFSSLPSPKELSSFSVNMKANEKNARSKTKSLLKRMESLKIKSSHHGKNKAPSKLGLIISGPILKEGLDEDKLKQFNCMEISTLNGNHINVPMVRKRSVSNSTQTSSNGSQSETSSTVSTPSPVTRTRSLSAYNKRVGMYLEGFDPFNQSTFGDVMEQNFKNRGSFAEDTVFFIPEDHKPGTFPKALSNGNFSPSESNSSVNWRTGSFHGHGHLSLRKENSAESSKELSTGKRRNSSSSVGSRLSIYDNVPGSILYSSTGDLADLENEDIFPELDDILYHVKGMQKIVNQWSEKFSDEGDSDSALDSVSPCPSSPKQIHLDVNNDRTTPSDLDSTGNSLTETEEPSGMQDRRDSGVGASLTRSNRQRLRCHSFQSSHGLSLSSVSLQINSQSVAQMNLLQKYSLLKLTALLEKYTPSNKHGFSWAVPKFMKRIKVPDYKDRNVFGVPLPINVQRTGQPLPQSIQQAMWYLRSHCLDQVGLFRKSGVKSRIQALRQMNESATDSVNYEGQSAYDVADMLKQYFRDLPEPLMTNKLSETFLQIYQYVPKDQRLQAIKAAVMLLPDENREVLQILLYFLGDVTAAVKENQMTPTNLAVCLAPSLFHLNTLKRENSSPRVMQRKQSLGKPDQKDLNENLAATQGLAHMIAECKKLFQIPEEMSRCRNSYTEQDLRPLSLEELGRINSMELSDYHCYLQDCMDDLLKEIKDKFKGWVSYSTSEQAELAYKKVCEGPPLRLWKATIEVPAVPEEVLNRLLKEQHLWDEDLLDSKVIETLDSQTDVYQYVQNSMAPHPARDYVILRTWRTNLSKGACVLLASSVDHDHAPVLGVRVNVLLSRYLIEPCGSGKSKLTYMCRIDLRGHMPEWYTKSFGHLCASEVAKIRDSFNKVVKST</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZG1</accession>
    <name>A0A452GZG1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Neurotensin/neuromedin N</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">Neurotensin may play an endocrine or paracrine role in the regulation of fat metabolism. It causes contraction of smooth muscle.</text>
    </comment>
    <comment type="subunit">
        <text evidence="11">Interacts with NTSR1. Interacts with SORT1. Interacts with SORL1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasmic vesicle</location>
            <location evidence="1">Secretory vesicle</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the neurotensin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZG1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008757.1">
        <property type="protein sequence ID" value="ENSGAGP00000007586.1"/>
        <property type="gene ID" value="ENSGAGG00000006071.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043679">
        <property type="term" value="C:axon terminus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030133">
        <property type="term" value="C:transport vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005184">
        <property type="term" value="F:neuropeptide hormone activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097746">
        <property type="term" value="P:blood vessel diameter maintenance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008055">
        <property type="entry name" value="NeurotensiN"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15356">
        <property type="entry name" value="NEUROTENSIN/NEUROMEDIN N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15356:SF0">
        <property type="entry name" value="NEUROTENSIN_NEUROMEDIN N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07421">
        <property type="entry name" value="Pro-NT_NN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01668">
        <property type="entry name" value="NEUROTENSIN"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0165">Cleavage on pair of basic residues</keyword>
    <keyword evidence="9" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0964">Secreted</keyword>
    <keyword evidence="7 12" id="KW-0732">Signal</keyword>
    <keyword evidence="8" id="KW-0838">Vasoactive</keyword>
    <feature type="signal peptide" evidence="12">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019489818" description="Neurotensin/neuromedin N" evidence="12">
        <location>
            <begin position="23"/>
            <end position="165"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004398"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00009827"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00016213"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022685"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022858"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00025449"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00046937"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007586.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="165" mass="19405" checksum="2F297E3416360C16" modified="2019-05-08" version="1">MARMRIQLVCMILLAFTSWSLCSDSEEEMKALEADLLTNMYTSKPYRVKLPYWKMILLNVCSVINNINNEAEETAESEEDDLLPRGQFPAALDGSSLEVMLTIYQLQKVCHSRAFQHWEMLQQDVFNQEDSNQDIMKRKTPYILKRQLHGNKARRPYILKRSSYY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZG5</accession>
    <name>A0A452GZG5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase</fullName>
            <ecNumber evidence="10">2.7.4.24</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">Bifunctional inositol kinase that acts in concert with the IP6K kinases to synthesize the diphosphate group-containing inositol pyrophosphates diphosphoinositol pentakisphosphate, PP-InsP5, and bis-diphosphoinositol tetrakisphosphate, (PP)2-InsP4. PP-InsP5 and (PP)2-InsP4, also respectively called InsP7 and InsP8, may regulate a variety of cellular processes, including apoptosis, vesicle trafficking, cytoskeletal dynamics, and exocytosis. Phosphorylates inositol hexakisphosphate (InsP6).</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="9">
            <text>1D-myo-inositol hexakisphosphate + ATP = 1-diphospho-1D-myo-inositol 2,3,4,5,6-pentakisphosphate + ADP</text>
            <dbReference type="Rhea" id="RHEA:37459"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:58130"/>
            <dbReference type="ChEBI" id="CHEBI:74946"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.4.24"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="9">
            <dbReference type="Rhea" id="RHEA:37460"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="8">
            <text>5-diphospho-1D-myo-inositol 1,2,3,4,6-pentakisphosphate + ATP + H(+) = 1,5-bis(diphospho)-1D-myo-inositol 2,3,4,6-tetrakisphosphate + ADP</text>
            <dbReference type="Rhea" id="RHEA:10276"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:58628"/>
            <dbReference type="ChEBI" id="CHEBI:77983"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.4.24"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="8">
            <dbReference type="Rhea" id="RHEA:10277"/>
        </physiologicalReaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 10">Cytoplasm</location>
            <location evidence="1 10">Cytosol</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 10">Belongs to the histidine acid phosphatase family. VIP1 subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.4.24" evidence="10"/>
    <dbReference type="AlphaFoldDB" id="A0A452GZG5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008614.1">
        <property type="protein sequence ID" value="ENSGAGP00000007458.1"/>
        <property type="gene ID" value="ENSGAGG00000005969.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033857">
        <property type="term" value="F:5-diphosphoinositol pentakisphosphate 1-kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0052723">
        <property type="term" value="F:inositol hexakisphosphate 1-kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="RHEA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006020">
        <property type="term" value="P:inositol metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032958">
        <property type="term" value="P:inositol phosphate biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07061">
        <property type="entry name" value="HP_HAP_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.470.20:FF:000003">
        <property type="entry name" value="Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.11950:FF:000003">
        <property type="entry name" value="Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1240:FF:000013">
        <property type="entry name" value="Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.11950">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.470.20">
        <property type="entry name" value="ATP-grasp fold, B domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1240">
        <property type="entry name" value="Phosphoglycerate mutase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033379">
        <property type="entry name" value="Acid_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000560">
        <property type="entry name" value="His_Pase_clade-2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037446">
        <property type="entry name" value="His_Pase_VIP1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029033">
        <property type="entry name" value="His_PPase_superfam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040557">
        <property type="entry name" value="VIP1_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12750">
        <property type="entry name" value="DIPHOSPHOINOSITOL PENTAKISPHOSPHATE KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12750:SF10">
        <property type="entry name" value="INOSITOL HEXAKISPHOSPHATE AND DIPHOSPHOINOSITOL-PENTAKISPHOSPHATE KINASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00328">
        <property type="entry name" value="His_Phos_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18086">
        <property type="entry name" value="PPIP5K2_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56059">
        <property type="entry name" value="Glutathione synthetase ATP-binding domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53254">
        <property type="entry name" value="Phosphoglycerate mutase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00616">
        <property type="entry name" value="HIS_ACID_PHOSPHAT_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7 10" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="3 10" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6 10" id="KW-0418">Kinase</keyword>
    <keyword evidence="5 10" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 10" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="VIP1 N-terminal" evidence="12">
        <location>
            <begin position="1"/>
            <end position="83"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="848"/>
            <end position="932"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="949"/>
            <end position="968"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1117"/>
            <end position="1151"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="865"/>
            <end position="895"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="11">
        <location>
            <begin position="910"/>
            <end position="921"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="1119"/>
            <end position="1129"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="1138"/>
            <end position="1151"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004514"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005609"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00033696"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00034629"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="RuleBase" id="RU365032"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF18086"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007458.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1151" mass="130138" checksum="C40FA4C58BF759E6" modified="2019-05-08" version="1">MAKKSKSKPMNEILERLSMFKYITVVIFEEDVILNEPVENWPLCDCLISFHSKGFPLDKAVAYAKLRNPFVINDLNMQYHIQDRREVYSILKAEGILLPRYAILNRDPNNPKECSLIEGEDQVEVNGEVFQKPFVEKPVSAEDHNVYIYYPTSAGGGSQRLFRKIGSRSSVYSPESNVRKTGSYIYEEFMPTDGTDVKVYTVGPDYAHAEARKSPALDGKVERDSEGKEVRYPVILNAREKLIAWKVCLAFKQTVCGFDLLRANGQSYVCDVNGFSFVKNSMKYYDDCAKILGNIIMRELAPQFQIPWSIPLEAEDIPIVPTTSGTMMELRCVIAVIRHGDRTPKQKMKMEVKHQKFFDLFEKCDGYKSGKLKLKKPKQLQEVLDIARQLLIELGQNNDSEIEESKAKLEQLKTVLEMYGHFSGINRKVQLTYLPHGCQKTSSEEEDNRRDEPSLLLVLKWGGELTPAGRVQAEELGRAFRCMYPGGQGDYAGFPGCGLLRLHSTYRHDLKIYASDEGRVQMTAAAFAKGLLALEGELTPILVQMVKSANMNGLLDSDSDSLSSCQHRVKARLHEILQRDREFMPEDYEKLTPSGSISLMKSMKVIRNPVRTCDKVYSLIQSLTSQIRQRMEDPKSADIQLYHSETLELMLRRWAKLEKDFKTKNGRYDISKIPDIYDCIKYDVQHNGSLKLENTMELYRLSKALADIVIPQEYGISKVEKLEIAKGYCTPLVRKIRSDLQRTQDDDTVNKLHPLYSRGVMSPERHVRTRLYFTSESHVHSLLSTLRYGALCDESKDEQWKRAMDYLNIVSELNYMTQIVIMLYEDPNKELSSEERFHVELHFSPGAKGCEEDKNLPAGFGYRPASRENEGSKKTSHKNDSDEESHTSKRDETDRSIVVFKPMVSDPIHIHRKSPLPRSRKIGSVEEESPLSVSSPECIGTWLHYTSGVGTGRRRRRSGEQITSSPVSPKSLAFTSSIFGSWQQVLSDNNSNLRTPRTILEQKQSGLDTTRGSAVKRFSISFARHPTNGFELYSMVPSICPLETLHNSLSLKQVDEFLASIAAPSSDNLPEISTALCASSAYLSSLSGRKCSLNTYNPAKIQPTPLAALPERLAIEKPTLSTSGPSNHISDVAPSPKESVKCGDETLTEKK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZH8</accession>
    <name>A0A452GZH8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">DCN1-like protein</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="3">Defective in cullin neddylation protein 1-like protein</fullName>
        </alternativeName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZH8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008767.1">
        <property type="protein sequence ID" value="ENSGAGP00000007595.1"/>
        <property type="gene ID" value="ENSGAGG00000006077.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000151">
        <property type="term" value="C:ubiquitin ligase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097602">
        <property type="term" value="F:cullin family protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031624">
        <property type="term" value="F:ubiquitin conjugating enzyme binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032182">
        <property type="term" value="F:ubiquitin-like protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045116">
        <property type="term" value="P:protein neddylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.238.10:FF:000041">
        <property type="entry name" value="DCN1-like protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.238.200:FF:000002">
        <property type="entry name" value="DCN1-like protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.238.200">
        <property type="entry name" value="Cullin, PONY binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.238.10">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014764">
        <property type="entry name" value="DCN-prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042460">
        <property type="entry name" value="DCN1-like_PONY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005176">
        <property type="entry name" value="PONY_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12281:SF8">
        <property type="entry name" value="DCN1-LIKE PROTEIN 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12281">
        <property type="entry name" value="RP42 RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03556">
        <property type="entry name" value="Cullin_binding"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51229">
        <property type="entry name" value="DCUN1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0539">Nucleus</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="DCUN1" evidence="5">
        <location>
            <begin position="101"/>
            <end position="287"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="40"/>
            <end position="79"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="47"/>
            <end position="58"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="RuleBase" id="RU363131"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS51229"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007595.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="292" mass="33985" checksum="FD58B925CAB6A726" modified="2019-05-08" version="1">MHSDAAAVNFQLNSHLSTLANIHKIYHTLNRLNLTEDVGQDDHQTGSLRSCSSSDCFSKVMPPRKKRRPATGDDLSAKKSRHDGMYRKYDSTRIKTEEEVFSSKRCLEWFYEYAGTDDVVGPEGMEKFCEDIGVEPENVVMLVLAWKLDAQNMGYFTLQEWLKGMTSLQCDVTEKLRNSLDYLRSLLNEPTNFKLIYRYAFDFAREKDQRSLDINTAKCMLGLLLGKTWPLFPVFHQFLEQSKYKVINKDQWCNVLEFSRTINLDLSNYDEDGAWPVLLDEFVEWYKGKQMT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZH9</accession>
    <name>A0A452GZH9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Alpha-mannosidase</fullName>
            <ecNumber evidence="16">3.2.1.-</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Catalyzes the first committed step in the biosynthesis of complex N-glycans. It controls conversion of high mannose to complex N-glycans; the final hydrolytic step in the N-glycan maturation pathway.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>N(4)-{beta-D-GlcNAc-(1-&gt;2)-alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;6)]-alpha-D-Man-(1-&gt;6)]-beta-D-Man-(1-&gt;4)-beta-D-GlcNAc-(1-&gt;4)-beta-D-GlcNAc}-L-asparaginyl-[protein] + 2 H2O = 2 alpha-D-mannopyranose + an N(4)-{beta-D-GlcNAc-(1-&gt;2)-alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;6)]-beta-D-Man-(1-&gt;4)-beta-D-GlcNAc-(1-&gt;4)-beta-D-GlcNAc}-L-asparaginyl-[protein]</text>
            <dbReference type="Rhea" id="RHEA:56052"/>
            <dbReference type="Rhea" id="RHEA-COMP:14368"/>
            <dbReference type="Rhea" id="RHEA-COMP:14369"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:28729"/>
            <dbReference type="ChEBI" id="CHEBI:60615"/>
            <dbReference type="ChEBI" id="CHEBI:60625"/>
            <dbReference type="EC" id="3.2.1.114"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="16">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
        <text evidence="16">Binds 1 zinc ion per subunit.</text>
    </comment>
    <comment type="pathway">
        <text evidence="2">Protein modification; protein glycosylation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Golgi apparatus membrane</location>
            <topology evidence="1">Single-pass type II membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 16">Belongs to the glycosyl hydrolase 38 family.</text>
    </comment>
    <dbReference type="EC" id="3.2.1.-" evidence="16"/>
    <dbReference type="AlphaFoldDB" id="A0A452GZH9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008715.1">
        <property type="protein sequence ID" value="ENSGAGP00000007548.1"/>
        <property type="gene ID" value="ENSGAGG00000002370.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000139">
        <property type="term" value="C:Golgi membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030246">
        <property type="term" value="F:carbohydrate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004572">
        <property type="term" value="F:mannosyl-oligosaccharide 1,3-1,6-alpha-mannosidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006013">
        <property type="term" value="P:mannose metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006491">
        <property type="term" value="P:N-glycan processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1270.50:FF:000001">
        <property type="entry name" value="Alpha-mannosidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.70.98.30:FF:000002">
        <property type="entry name" value="Alpha-mannosidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.20.110.10:FF:000003">
        <property type="entry name" value="Alpha-mannosidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.20.110.10">
        <property type="entry name" value="Glycoside hydrolase 38, N terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.50">
        <property type="entry name" value="Glycoside hydrolase family 38, central domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1180">
        <property type="entry name" value="Golgi alpha-mannosidase II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.70.98.30">
        <property type="entry name" value="Golgi alpha-mannosidase II, domain 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011013">
        <property type="entry name" value="Gal_mutarotase_sf_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011330">
        <property type="entry name" value="Glyco_hydro/deAcase_b/a-brl"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011682">
        <property type="entry name" value="Glyco_hydro_38_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015341">
        <property type="entry name" value="Glyco_hydro_38_cen"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037094">
        <property type="entry name" value="Glyco_hydro_38_cen_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000602">
        <property type="entry name" value="Glyco_hydro_38_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027291">
        <property type="entry name" value="Glyco_hydro_38_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028995">
        <property type="entry name" value="Glyco_hydro_57/38_cen_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013780">
        <property type="entry name" value="Glyco_hydro_b"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050843">
        <property type="entry name" value="Glycosyl_Hydrlase_38"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11607">
        <property type="entry name" value="ALPHA-MANNOSIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11607:SF57">
        <property type="entry name" value="ALPHA-MANNOSIDASE 2X"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09261">
        <property type="entry name" value="Alpha-mann_mid"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07748">
        <property type="entry name" value="Glyco_hydro_38C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01074">
        <property type="entry name" value="Glyco_hydro_38N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00872">
        <property type="entry name" value="Alpha-mann_mid"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF88688">
        <property type="entry name" value="Families 57/38 glycoside transferase middle domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF74650">
        <property type="entry name" value="Galactose mutarotase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF88713">
        <property type="entry name" value="Glycoside hydrolase/deacetylase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="13 16" id="KW-0326">Glycosidase</keyword>
    <keyword evidence="10" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="6 16" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="11 17" id="KW-0472">Membrane</keyword>
    <keyword evidence="5 16" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0735">Signal-anchor</keyword>
    <keyword evidence="4 17" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9 17" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="7 16" id="KW-0862">Zinc</keyword>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="7"/>
            <end position="26"/>
        </location>
    </feature>
    <feature type="domain" description="Glycoside hydrolase family 38 central" evidence="18">
        <location>
            <begin position="502"/>
            <end position="588"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004323"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004922"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00009792"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023295"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00059516"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00093232"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="RuleBase" id="RU361199"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="SMART" id="SM00872"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007548.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1129" mass="127472" checksum="26962E3CF722E93B" modified="2019-05-08" version="1">MKLKKQVTVCGAAIFCVAVFSLYLMLDRVQHDPARHQSGGNFPRSQISVLQNRIEQLEQLLEENHEIISHIKESVLELTAHVDGQPVLLPYRTPNGSWVLPPESRPSFFSISPQDCQFALGSGGQKPDLQMLAVASVLPFDNQDGGVWKQGFDITYEPHDWDTEPLQVFVVPHSHNDPGWIKTFEKYYADQTQHILNSMVLKLQEDPRRRFIWSEISFFSKWWDNISTQKRAAVRRLVGGGQLEMVTGGWVMPDEANSHYFAMIDQLIEGHQWLEKNIGVTPRSGWAVDPFGHSATMPYLLRRANLTSMLIQRVHYTIKKHFAATRSLEFMWRQTWDPDSSTDIFCHMMPFYSYDVPHTCGPDPKICCQFDFKRLPGGRINCPWKVPPRAITEANVAERAHLLLDQYRKKSKLFRSKVLLVPLGDDFRYDKPQEWDAQFLNYQRLFDFLNAHPSLHVQAQFGTLSDYFDALSKRTGIIPGLKPPGFPVLSGDFFSYADREDHYWTGYYTSRPFYKSLDRVLEAHLRGAEILYSLALSHARHAGMDSRYPLSDYALLTEARRALGLFQHHDAITGTAKEAVVVDYGVRLLHSLVNLKRIISNAAHYLVLGEKESYHYDAAGPFLGMDDMRLNQDSLPEKTIIKLDTSPRCTPSSGLGQALSLLVNSPRVRVLSEEGQPLAIQLSAQWSSATDMVPDVYQVSIMTRLPALGLSILQLYKSFDSHSTLTSSVHIHLHGRDLPVRPREVFPVRIIPATSDDFCLDNQHMQACFSGLTGLLQSVRRAGEEHRLSSEFLIYGTRSAKDKSGAYLFLPDGDAKPYAPKEPPVVRVTEGPFFSEVASYYQHVQQVVRLYNVPGVEGLSLDVSCLVDIRDHVNKEMALRLSTTIASEDTFFTDLNGFQIQPRRFLKKLPLQANFYPMPAMAYIQDKQSRLTLHTAQALGVSSLHSGEGGQCPRAICNWLQRLLEPRTHHCSQSKQGQDNRPISFPSLLSHMTSMHLNAEVLAMPVAREKPAPPALRSFRPLSATVPCDFHILNLRTLQAEDDLLPSAEIALILHRKGFDCGLEAKHLGFNCTTSQGKFSLGSLFHGLELGSLQPTSLTLMYPLGAASPNSTVIHMDPMEIATFRIHFE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZI8</accession>
    <name>A0A452GZI8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Tripartite motif containing 10</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GZI8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008645.1">
        <property type="protein sequence ID" value="ENSGAGP00000007483.1"/>
        <property type="gene ID" value="ENSGAGG00000005976.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd16594">
        <property type="entry name" value="RING-HC_TRIM7-like_C-IV"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12888">
        <property type="entry name" value="SPRY_PRY_TRIM7_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.920:FF:000004">
        <property type="entry name" value="Butyrophilin subfamily 1 member A1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.920">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001870">
        <property type="entry name" value="B30.2/SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043136">
        <property type="entry name" value="B30.2/SPRY_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003879">
        <property type="entry name" value="Butyrophylin_SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006574">
        <property type="entry name" value="PRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003877">
        <property type="entry name" value="SPRY_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050143">
        <property type="entry name" value="TRIM/RBCC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000315">
        <property type="entry name" value="Znf_B-box"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001841">
        <property type="entry name" value="Znf_RING"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017907">
        <property type="entry name" value="Znf_RING_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24103">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE TRIM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13765">
        <property type="entry name" value="PRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00622">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00643">
        <property type="entry name" value="zf-B_box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15227">
        <property type="entry name" value="zf-C3HC4_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01407">
        <property type="entry name" value="BUTYPHLNCDUF"/>
    </dbReference>
    <dbReference type="SMART" id="SM00589">
        <property type="entry name" value="PRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00184">
        <property type="entry name" value="RING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00449">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57845">
        <property type="entry name" value="B-box zinc-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57850">
        <property type="entry name" value="RING/U-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50188">
        <property type="entry name" value="B302_SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50119">
        <property type="entry name" value="ZF_BBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00518">
        <property type="entry name" value="ZF_RING_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50089">
        <property type="entry name" value="ZF_RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="1" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0862">Zinc</keyword>
    <keyword evidence="2 4" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="RING-type" evidence="7">
        <location>
            <begin position="16"/>
            <end position="57"/>
        </location>
    </feature>
    <feature type="domain" description="B box-type" evidence="8">
        <location>
            <begin position="48"/>
            <end position="105"/>
        </location>
    </feature>
    <feature type="domain" description="B30.2/SPRY" evidence="9">
        <location>
            <begin position="237"/>
            <end position="425"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="210"/>
            <end position="247"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="5">
        <location>
            <begin position="157"/>
            <end position="206"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="237"/>
            <end position="247"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00024"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50119"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50188"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007483.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="12">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="425" mass="47838" checksum="17A3CA886376B010" modified="2019-05-08" version="1">MAAENPVESLQEEATCPICLEYFTEPVTLECGHNFCRACIAQCWEGSDTDISCPQCRETVQQRNLRPNRQLANVALKLFCEEDQSPICVICRESRAHCAHTVVPIQEAAQEYKEKIQAYLKALREEREKLLGFKVTEKGKCQEYLRQKIVSEFQELREFLEEQERLLLAQLEKLDEEIVRIQNENVSKLSEQISRLSELISEMEGKCQKPGNRVRLQPLPGAGPRSTSPEALAPGQARSQRPSSSPNALILSVPANVTLDPDTAQLRLVLSEDRKSVRYGDTRQDLPDNPQRFDTDFCVLGCEGFTSGRHCWEVEVEDGQCWAVGVARESVGRKGGISYSPEGGIWAVLRWMGQFQVLTSPVTPLPLRRVPSRIQVCLDCDRGQVTFIDAGDEAPIFTFPPGSIPGERIRPWLCVGYRSRLCLCP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZJ0</accession>
    <name>A0A452GZJ0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">Receptor-type tyrosine-protein phosphatase F</fullName>
            <ecNumber evidence="3">3.1.3.48</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>O-phospho-L-tyrosyl-[protein] + H2O = L-tyrosyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:10684"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="EC" id="3.1.3.48"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the protein-tyrosine phosphatase family. Receptor class 2A subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.48" evidence="3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008727.1">
        <property type="protein sequence ID" value="ENSGAGP00000007559.1"/>
        <property type="gene ID" value="ENSGAGG00000005093.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="CDD" id="cd05738">
        <property type="entry name" value="IgI_2_RPTP_IIa_LAR_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05739">
        <property type="entry name" value="IgI_3_RPTP_IIa_LAR_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000010">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000027">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000036">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000066">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000144">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000015">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000082">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000128">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase delta isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000098">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase F isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000353">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase F isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013098">
        <property type="entry name" value="Ig_I-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050713">
        <property type="entry name" value="RTP_Phos/Ushers"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46957">
        <property type="entry name" value="CYTOKINE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46957:SF9">
        <property type="entry name" value="PROTEIN-TYROSINE-PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07679">
        <property type="entry name" value="I-set"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00014">
        <property type="entry name" value="FNTYPEIII"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="13" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="15" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="4" id="KW-0358">Heparin-binding</keyword>
    <keyword evidence="8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="16" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="12 20" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="14" id="KW-0675">Receptor</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="5 20" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 20" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="1151"/>
            <end position="1174"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="21">
        <location>
            <begin position="40"/>
            <end position="129"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="21">
        <location>
            <begin position="137"/>
            <end position="219"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="226"/>
            <end position="317"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="322"/>
            <end position="416"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="420"/>
            <end position="509"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="514"/>
            <end position="611"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="616"/>
            <end position="715"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="716"/>
            <end position="810"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="814"/>
            <end position="906"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="910"/>
            <end position="994"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="596"/>
            <end position="617"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1316"/>
            <end position="1344"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004167"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010504"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00013064"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022674"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00044158"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00051722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007559.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1381" mass="152402" checksum="F9706797CC7928E5" modified="2019-05-08" version="1">MSSLPTNGQKARSLDPVLLCGGFCCMLFPVAEDHLPAGFPTIDMGPQLKVVEKARTATMLCAASGNPDPEISWFKDFLPVDTATSNGRIKQLRSGALQIENSEESDQGKYECVATNSAGTRYSAPANLYVRVRRVAPRFSIPPSNHEVMPGGSVNLTCVAVGAPMPYVKWMAGVEELTKEDEMPVGRNVLELNNIVQSASYTCVAMSSLGMIEATAQITVKALPKPPTEPLVTETTATSVTLTWDSGNSDPISYYVIQYKPKSWEEGLFQEVDGVATTRYSIGGLSPFSEYEFRIIAVNNIGRGPPSELVEARTGEQAPSSPPLKVQARMLSASTMLVQWEQPEEPNGQIRGYRVYYTTDPHLPLSMWQKHNTDDSHLTTVGSLITGTTYSIRVLAFTSVGDGPPSDIIQVKTQQGVPAQPADFQAEAESDTRILLTWLPALQERITKYELLYWEGEDGSQRKVNFDPSSSYAVEGLKPDTLYKFQLGARSELGLGVYTPVLEARTAQSTPSAPPQEVECVSTSSTTIRVSWVPPPAQSRNGVITQYSIAYQAVDAEDGAKHVVDGIGRERSSWEIKDLEKWTEYKVWVRAHTDVGPGPESSPVHVRTDEDVPSAPPRKVEVESLNSTAIRVSWKLPLSNKQHGQIRGYQVTYVKLENNEPRGQPVIKDVMLAEAQETIIGGLLPETTYSITVAAYTTKGDGARSKPKVVTTTGAVPGKPSMLISTTAMNTALIQWHPPKEMVGELLGYRLQYKRTDEEKMNIMDFEKNVHHYTVTSLHKGATYIFKLSAKNRAGPGEEFEKEITTTEDVPSGFPQNLRVVGLTTSTTEVAWDPPVLAERNGRITNYTVMYRDINSQQELTNITRETSIVLTNLRPDTTYDIKVRACNSKGVGPLSPSIQSRTMPVEQVFAKNFRVNAVMKTSVLLSWEVPDSYKSAVPFKILYNSQSVEVDGHSMKKLISDLQPDTEYSFVLMNRGSSAGGLQHRVSIHTAPDVLQSKPISTNKYMQEGKFTLTLPKVQTSAPVRWYYIVVVPVDQSTSSRTARWRTPDEMELDQLLEAISQGRRQRRQADRLKPYIAALVDMLPETFTLGDEKTYKGFYNKPLSEDLSYRCFVLASLEDGDTKRYAASPYSDEIVMEVSSAKQQDEPEMLWVMGPVLAVILIIIIVIAILLFKRKRANSPSSKDEHSIGLKDSLLAHSSDPVEMRRLNYQTPGMRDHPPIPVTDLADNIDRLKANDGLRFSQEYEVSCREGDRCAWHCPEPGVAGRRWAMAASPRGGSVDSFHAHSFSVHLPRGYPLSWLVPQLQPPAASLVTRRSCPQAPRPGLQRPGVPHXGSPYSCSADTPQSHLASPCPPTHTLALLIPLSPALPLPGLPLVLLC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZJ3</accession>
    <name>A0A452GZJ3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Alpha-mannosidase</fullName>
            <ecNumber evidence="16">3.2.1.-</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Catalyzes the first committed step in the biosynthesis of complex N-glycans. It controls conversion of high mannose to complex N-glycans; the final hydrolytic step in the N-glycan maturation pathway.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>N(4)-{beta-D-GlcNAc-(1-&gt;2)-alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;6)]-alpha-D-Man-(1-&gt;6)]-beta-D-Man-(1-&gt;4)-beta-D-GlcNAc-(1-&gt;4)-beta-D-GlcNAc}-L-asparaginyl-[protein] + 2 H2O = 2 alpha-D-mannopyranose + an N(4)-{beta-D-GlcNAc-(1-&gt;2)-alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;6)]-beta-D-Man-(1-&gt;4)-beta-D-GlcNAc-(1-&gt;4)-beta-D-GlcNAc}-L-asparaginyl-[protein]</text>
            <dbReference type="Rhea" id="RHEA:56052"/>
            <dbReference type="Rhea" id="RHEA-COMP:14368"/>
            <dbReference type="Rhea" id="RHEA-COMP:14369"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:28729"/>
            <dbReference type="ChEBI" id="CHEBI:60615"/>
            <dbReference type="ChEBI" id="CHEBI:60625"/>
            <dbReference type="EC" id="3.2.1.114"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="16">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
        <text evidence="16">Binds 1 zinc ion per subunit.</text>
    </comment>
    <comment type="pathway">
        <text evidence="2">Protein modification; protein glycosylation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Golgi apparatus membrane</location>
            <topology evidence="1">Single-pass type II membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 16">Belongs to the glycosyl hydrolase 38 family.</text>
    </comment>
    <dbReference type="EC" id="3.2.1.-" evidence="16"/>
    <dbReference type="AlphaFoldDB" id="A0A452GZJ3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008733.1">
        <property type="protein sequence ID" value="ENSGAGP00000007565.1"/>
        <property type="gene ID" value="ENSGAGG00000002370.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000139">
        <property type="term" value="C:Golgi membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030246">
        <property type="term" value="F:carbohydrate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004572">
        <property type="term" value="F:mannosyl-oligosaccharide 1,3-1,6-alpha-mannosidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006013">
        <property type="term" value="P:mannose metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006491">
        <property type="term" value="P:N-glycan processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd11667">
        <property type="entry name" value="GH38N_Man2A2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1270.50:FF:000001">
        <property type="entry name" value="Alpha-mannosidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.70.98.30:FF:000002">
        <property type="entry name" value="Alpha-mannosidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.20.110.10:FF:000003">
        <property type="entry name" value="Alpha-mannosidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.20.110.10">
        <property type="entry name" value="Glycoside hydrolase 38, N terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.50">
        <property type="entry name" value="Glycoside hydrolase family 38, central domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1180">
        <property type="entry name" value="Golgi alpha-mannosidase II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.70.98.30">
        <property type="entry name" value="Golgi alpha-mannosidase II, domain 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011013">
        <property type="entry name" value="Gal_mutarotase_sf_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011330">
        <property type="entry name" value="Glyco_hydro/deAcase_b/a-brl"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011682">
        <property type="entry name" value="Glyco_hydro_38_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015341">
        <property type="entry name" value="Glyco_hydro_38_cen"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037094">
        <property type="entry name" value="Glyco_hydro_38_cen_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000602">
        <property type="entry name" value="Glyco_hydro_38_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027291">
        <property type="entry name" value="Glyco_hydro_38_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028995">
        <property type="entry name" value="Glyco_hydro_57/38_cen_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013780">
        <property type="entry name" value="Glyco_hydro_b"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050843">
        <property type="entry name" value="Glycosyl_Hydrlase_38"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11607">
        <property type="entry name" value="ALPHA-MANNOSIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11607:SF57">
        <property type="entry name" value="ALPHA-MANNOSIDASE 2X"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09261">
        <property type="entry name" value="Alpha-mann_mid"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07748">
        <property type="entry name" value="Glyco_hydro_38C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01074">
        <property type="entry name" value="Glyco_hydro_38N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00872">
        <property type="entry name" value="Alpha-mann_mid"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF88688">
        <property type="entry name" value="Families 57/38 glycoside transferase middle domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF74650">
        <property type="entry name" value="Galactose mutarotase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF88713">
        <property type="entry name" value="Glycoside hydrolase/deacetylase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="13 16" id="KW-0326">Glycosidase</keyword>
    <keyword evidence="10" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="6 16" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="11 17" id="KW-0472">Membrane</keyword>
    <keyword evidence="5 16" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0735">Signal-anchor</keyword>
    <keyword evidence="4 17" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9 17" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="7 16" id="KW-0862">Zinc</keyword>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="7"/>
            <end position="26"/>
        </location>
    </feature>
    <feature type="domain" description="Glycoside hydrolase family 38 central" evidence="18">
        <location>
            <begin position="502"/>
            <end position="588"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004323"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004922"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00009792"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022968"/>
        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023295"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00059516"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00093232"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="RuleBase" id="RU361199"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="SMART" id="SM00872"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007565.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1125" mass="127126" checksum="749D5E47FB0BDCA0" modified="2019-05-08" version="1">MKLKKQVTVCGAAIFCVAVFSLYLMLDRVQHDPARHQSGGNFPRSQISVLQNRIEQLEQLLEENHEIISHIKESVLELTAHVDGQPVLLPYRTPNGSWVLPPESRPSFFSISPQDCQFALGSGGQKPDLQMLAVASVLPFDNQDGGVWKQGFDITYEPHDWDTEPLQVFVVPHSHNDPGWIKTFEKYYADQTQHILNSMVLKLQEDPRRRFIWSEISFFSKWWDNISTQKRAAVRRLVGGGQLEMVTGGWVMPDEANSHYFAMIDQLIEGHQWLEKNIGVTPRSGWAVDPFGHSATMPYLLRRANLTSMLIQRVHYTIKKHFAATRSLEFMWRQTWDPDSSTDIFCHMMPFYSYDVPHTCGPDPKICCQFDFKRLPGGRINCPWKVPPRAITEANVAERAHLLLDQYRKKSKLFRSKVLLVPLGDDFRYDKPQEWDAQFLNYQRLFDFLNAHPSLHVQAQFGTLSDYFDALSKRTGIIPGLKPPGFPVLSGDFFSYADREDHYWTGYYTSRPFYKSLDRVLEAHLRGAEILYSLALSHARHAGMDSRYPLSDYALLTEARRALGLFQHHDAITGTAKEAVVVDYGVRLLHSLVNLKRIISNAAHYLVLGEKESYHYDAAGPFLGMVSPALCPGPGPDLALSLLVNSPRVRVLSEEGQPLAIQLSAQWSSATDMVPDVYQVSIMTRLPALGLSILQLYKSFDSHSTLTSSVHIHLHGRDLPVRPREVFPVRIIPATSDDFCLDNQHMQACFSGLTGLLQSVRRAGEEHRLSSEFLIYGTRSAKDKSGAYLFLPDGDAKPYAPKEPPVVRVTEGPFFSEVASYYQHVQQVVRLYNVPGVEGLSLDVSCLVDIRDHVNKEMALRLSTTIASEDTFFTDLNGFQIQPRRFLKKLPLQANFYPMPAMAYIQDKQSRLTLHTAQALGVSSLHSGQLEVILDRRLMQDDNRGLGQGLKDNKRTCNRFRLLLERRTTANRVQDNRPISFPSLLSHMTSMHLNAEVLAMPVAREKPAPPALRSFRPLSATVPCDFHILNLRTLQAEDDLLPSAEIALILHRKGFDCGLEAKHLGFNCTTSQGKFSLGSLFHGLELGSLQPTSLTLMYPLGAASPNSTVIHMDPMEIATFRIHFE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZJ4</accession>
    <name>A0A452GZJ4_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="3">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the TAFA family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZJ4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008650.1">
        <property type="protein sequence ID" value="ENSGAGP00000007488.1"/>
        <property type="gene ID" value="ENSGAGG00000005995.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048018">
        <property type="term" value="F:receptor ligand activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020350">
        <property type="entry name" value="Chemokine-like_TAFA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051743">
        <property type="entry name" value="TAFA_chemokine-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31770">
        <property type="entry name" value="CHEMOKINE-LIKE PROTEIN TAFA FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31770:SF1">
        <property type="entry name" value="CHEMOKINE-LIKE PROTEIN TAFA-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12020">
        <property type="entry name" value="TAFA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0732">Signal</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006101"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007488.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="131" mass="14536" checksum="566A89297C370F2C" modified="2019-05-08" version="1">MNKRYLQKATKGKLLIIIFVVTLWGKLASGANHHKAHHVKTGTCEVVALHRCCNKNKIEERSQTVKCSCFPGQVAGTTRAAPSCVDASIVEQKWWCHMQPCLEGEECKVLPDRKGWSCSSGNKVKTTRANV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZJ5</accession>
    <name>A0A452GZJ5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Sm domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the ataxin-2 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZJ5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008790.1">
        <property type="protein sequence ID" value="ENSGAGP00000007616.1"/>
        <property type="gene ID" value="ENSGAGG00000006078.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010494">
        <property type="term" value="C:cytoplasmic stress granule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003729">
        <property type="term" value="F:mRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034063">
        <property type="term" value="P:stress granule assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045117">
        <property type="entry name" value="ATXN2-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009604">
        <property type="entry name" value="LsmAD_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009818">
        <property type="entry name" value="PAM2_motif"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047575">
        <property type="entry name" value="Sm"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025852">
        <property type="entry name" value="SM_dom_ATX"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12854">
        <property type="entry name" value="ATAXIN 2-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12854:SF8">
        <property type="entry name" value="ATAXIN-2-LIKE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06741">
        <property type="entry name" value="LsmAD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07145">
        <property type="entry name" value="PAM2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14438">
        <property type="entry name" value="SM-ATX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01272">
        <property type="entry name" value="LsmAD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS52002">
        <property type="entry name" value="SM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Sm" evidence="3">
        <location>
            <begin position="58"/>
            <end position="135"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="253"/>
            <end position="461"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="522"/>
            <end position="659"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="696"/>
            <end position="726"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="781"/>
            <end position="813"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="825"/>
            <end position="903"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="253"/>
            <end position="264"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="266"/>
            <end position="278"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="299"/>
            <end position="319"/>
        </location>
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            <begin position="384"/>
            <end position="413"/>
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        <location>
            <begin position="522"/>
            <end position="537"/>
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            <end position="887"/>
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    </feature>
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    </feature>
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        </source>
    </evidence>
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    </evidence>
    <evidence type="ECO:0000259" key="3">
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            <dbReference type="PROSITE" id="PS52002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007616.1"/>
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    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1000" mass="107410" checksum="13D0E7D632783C87" modified="2019-05-08" version="1">MGVAVLVGAQSRQWDSDIQLSLSQIHLGGGFPAVLSLPSHWAMCWGCWVFEGVYNNSRMLHFLTAVVGSTCDVKVKNGSTFEGIFKTLSSKFELAVDAVHRKTPDQLVGPKREDIVDTMVFKPTDVMLVHFRNVDFNYATKDKFTDSAIAMNSKVNGEHKEKVLQRWEGGDSNSDDYDLESDMSNGWDPNEMFKFNEENYGVKTTYDSSLSSYTVPLEKDNSEEFRQREARAAQLAREIESSPQYRLRIAMENDDGRTEEEKHSSVQRQVSGRDSPSLASREGKYVPLPQRVREGSRGGIRCSSSRGGRPGVGSMPPRGSAHHPDSNSSPAPEQRGINGGPSRMSPKSQRPIRGAKTMSSPSSRPVEVSAAPPAVGRMYPPRSPKSASATPASSQDSPVGSAVSAAPASPSEARTGLESSVSPNPPSPKVAAPVLATAVEGKEAPTSVAKDPGRTLEATGSCELSKAANKAGLQNEQKRTQLEELRKFGAQFKVRPGDCKEGGGEWAYLGWAHHEFLAPLQLQPSSSPETSLESFSARPKEPLEGKEKSLEPGICEGPEEAPPVSMVPKPNGTSLELLPESGKEEKGLCEVAEQQATNPPGKGEPEDKEEGPVSEQVKKSTLNPNAKEFNPSKTLLSVNKSTSTPTSPGPRAHSTPSIPVITAGQTGMYSPQYISYIPQIHMNPAVQAPQMYPYPVSSSVPGQQGKYRGAKGSLPPQRTDQHQPTSAPPIMQAAAAAGPPLVAATPYSSYISYNPQQFTGQPTMMQPMAHYPSQPVFAPMLQSNPRMMTSGSHPQAIVSSSTPQYPPSEQPTPQTLYATVHQSYPHHATQLHPHQPQPATTPTGNQQQAQHAAPSPVQHQAGQPPHLASAQQQQNLYHTATLTATPPSITPGPSAQSPQSSFPQQAVYAIHAHQQLQHSYTNMAHVTQAHVQTGITAAPPPHPGAPHPPQVMLLHPTQTHGGPPQGGVPQSGVPTLSASTPSPYTYIGHHQGQCWGWGDV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZJ7</accession>
    <name>A0A452GZJ7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Hypoxia-inducible factor 3-alpha</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="19">HIF3-alpha-1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Mitochondrion</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZJ7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008881.1">
        <property type="protein sequence ID" value="ENSGAGP00000007696.1"/>
        <property type="gene ID" value="ENSGAGG00000006155.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
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    <dbReference type="GO" id="GO:0005634">
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        <property type="project" value="UniProtKB-SubCell"/>
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    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046983">
        <property type="term" value="F:protein dimerization activity"/>
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        <property type="project" value="InterPro"/>
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        <property type="term" value="F:RNA polymerase II transcription regulatory region sequence-specific DNA binding"/>
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        <property type="project" value="TreeGrafter"/>
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    <dbReference type="GO" id="GO:0001525">
        <property type="term" value="P:angiogenesis"/>
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        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006915">
        <property type="term" value="P:apoptotic process"/>
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        <property type="project" value="UniProtKB-KW"/>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="entry name" value="bHLH-PAS_HIF3a_PASD7"/>
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        <property type="entry name" value="Helix-loop-helix DNA-binding domain"/>
        <property type="match status" value="1"/>
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        <property type="match status" value="2"/>
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        <property type="entry name" value="HIF_alpha-like"/>
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        <property type="entry name" value="HLH_DNA-bd_sf"/>
    </dbReference>
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    </dbReference>
    <dbReference type="InterPro" id="IPR013767">
        <property type="entry name" value="PAS_fold"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00229">
        <property type="entry name" value="sensory_box"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23043">
        <property type="entry name" value="HYPOXIA-INDUCIBLE FACTOR 1 ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23043:SF18">
        <property type="entry name" value="HYPOXIA-INDUCIBLE FACTOR 3-ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23171">
        <property type="entry name" value="bHLH_HIF1A"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14598">
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00353">
        <property type="entry name" value="HLH"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="PAC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00091">
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        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47459">
        <property type="entry name" value="HLH, helix-loop-helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55785">
        <property type="entry name" value="PYP-like sensor domain (PAS domain)"/>
        <property type="match status" value="2"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50112">
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        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8" id="KW-0037">Angiogenesis</keyword>
    <keyword evidence="9" id="KW-0053">Apoptosis</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="17" id="KW-0379">Hydroxylation</keyword>
    <keyword evidence="7" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="14" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="16" id="KW-0539">Nucleus</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0678">Repressor</keyword>
    <keyword evidence="13" id="KW-0346">Stress response</keyword>
    <keyword evidence="15" id="KW-0804">Transcription</keyword>
    <keyword evidence="12" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="11" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="BHLH" evidence="22">
        <location>
            <begin position="67"/>
            <end position="120"/>
        </location>
    </feature>
    <feature type="domain" description="PAS" evidence="21">
        <location>
            <begin position="140"/>
            <end position="195"/>
        </location>
    </feature>
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        <location>
            <begin position="295"/>
            <end position="346"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="403"/>
            <end position="434"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="478"/>
            <end position="519"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
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            <begin position="568"/>
            <end position="669"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="774"/>
            <end position="795"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="20">
        <location>
            <begin position="404"/>
            <end position="415"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="20">
        <location>
            <begin position="593"/>
            <end position="605"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="20">
        <location>
            <begin position="628"/>
            <end position="637"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004173"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022491"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022657"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022703"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023016"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023278"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00068506"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00077359"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50112"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS50888"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007696.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="795" mass="87330" checksum="977965A3752EA433" modified="2019-05-08" version="1">MHSWLRLSSPGPPLITHCPPPLEVPPHLPLPLPPVPTPNINVCRDPMRDLALSNPAAAEGCRSTTELRKEKSRDAARCRRSKETEVFYQLAHTLPFARGVSAHLDKASIMRLTISYLRMHKLVTSEEWRGQPEQVDACYLKALDGFVMVLTEEGDMAYLSENVSKHLGLTQLELIGHSIFDFIHPCDQEELQDVLSPRQGFSKKEVKTERSFSLRMKSTLTSRGRTVNLKSATWKVLHCAGHMRSYTPAREGPEEAEGGYAEPPLRCLVLICEAIPHPANIETPLDSSTFLSQHSMDMKFTYCDERIAEVAGYTPDELLGCSIYEYIHALDSDAVSKSIHTLLSKGQAVTGQYRFLAKSGGYLWAQTQATVISNSKNSQPESVVCVHFVLSQVEETGMVLSLEQTERQGEGRRLPPEPNPAHQANGSPEELEPDTGETILNLSFERHGPRLLAFLRPAHVSEVELQRDPRRFCSPDLQRLLGPIFDPPGARTNREGAPRARSSPHPPKAGGPPQDSAKSTAPAELLFDMENVQKLFASTQEAVETVLQDYEALDLEMLAPYISMDDDFQLSSTDQPPWLAEKRGDPAVARGTGPPPPSASPPPRPRSSSFHGVSAREPDLPGLPRWGSETSLSQPRSLQPPEEEPMELEGPGSPRTTPPGDNRRDHTASHLGARKRALELNLEEEGADFLEAAPLKRAHGSEPDGFLLPSLNLGFLLSVEEGLDAGVPRGGMVLGRKLLALEEPMALLGDMLPFVVDGPALSQLALYDAEDEALEQSGGHFQPSEELLGELDQAT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZK0</accession>
    <name>A0A452GZK0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Interferon regulatory factor 8</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="11">Interferon consensus sequence-binding protein</fullName>
        </alternativeName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZK0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008809.1">
        <property type="protein sequence ID" value="ENSGAGP00000007632.1"/>
        <property type="gene ID" value="ENSGAGG00000006103.1"/>
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        <property type="component" value="Unassembled WGS sequence"/>
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        <property type="project" value="UniProtKB-SubCell"/>
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        <property type="term" value="C:nucleus"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="project" value="TreeGrafter"/>
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        <property type="term" value="P:defense response to other organism"/>
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        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
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        <property type="term" value="P:immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
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        <property type="term" value="P:negative regulation of transcription by RNA polymerase II"/>
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        <property type="project" value="UniProtKB-ARBA"/>
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        <property type="term" value="P:positive regulation of transcription by RNA polymerase II"/>
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        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
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        <property type="entry name" value="IRF"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Interferon regulatory factor 4"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Interferon regulatory factor 8"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.10">
        <property type="entry name" value="Winged helix-like DNA-binding domain superfamily/Winged helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Interferon_reg_fac_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001346">
        <property type="entry name" value="Interferon_reg_fact_DNA-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019471">
        <property type="entry name" value="Interferon_reg_factor-3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017855">
        <property type="entry name" value="SMAD-like_dom_sf"/>
    </dbReference>
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        <property type="entry name" value="SMAD_FHA_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036388">
        <property type="entry name" value="WH-like_DNA-bd_sf"/>
    </dbReference>
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        <property type="entry name" value="WH_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11949">
        <property type="entry name" value="INTERFERON REGULATORY FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11949:SF7">
        <property type="entry name" value="INTERFERON REGULATORY FACTOR 8"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="IRF"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="IRF-3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00267">
        <property type="entry name" value="INTFRNREGFCT"/>
    </dbReference>
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        <property type="entry name" value="IRF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01243">
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49879">
        <property type="entry name" value="SMAD/FHA domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46785">
        <property type="entry name" value="Winged helix' DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="IRF_1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="IRF_2"/>
        <property type="match status" value="1"/>
    </dbReference>
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    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="9" id="KW-0539">Nucleus</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0678">Repressor</keyword>
    <keyword evidence="8" id="KW-0804">Transcription</keyword>
    <keyword evidence="5" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="IRF tryptophan pentad repeat" evidence="12">
        <location>
            <begin position="57"/>
            <end position="164"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022491"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00067364"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00082311"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS51507"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007632.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="462" mass="52996" checksum="FEBEBAF1358BD32D" modified="2019-05-08" version="1">MRGRGGGVPTFAYPLWPVSAVLSEPAARSRWVHLGEQRVGRKPILQTADRMCDRNGGRRLRQWLIEQIDSGMYPGLIWENEEKSMFRIPWKHAGKQDYNQEVDASIFKAWAVFKGKFKEGDKAEPATWKTRLRCALNKSPDFEEVTDRSQLDISEPYKVYRIVPEEEQKCKAGIANGGSLNEATDMDCSPSAIDDLMKESPVVDEYLGIIKRSPSPPQENCRNPPIPDWWMQQPSPVYSQMVISFYYGGKLVGHTTTSYHEGCRISLSHAPAHCDKLYGPDSLEHVQFPSADAIPNDRQKHITKKLFGHLDRGVLLHSNKQGIFIKRLCQGRVFWSGNNVLYKDRPNKLDRDEVVKIFDSNQFVRELQQFYSSQGRFPESRVLLCFGEEFPDTVPLRSKLILVQVEQLWVRQLVEEAGKVYSSGPMLPVADETQHDQIFRIFQDICGTHQRPLFRENQQITV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZK5</accession>
    <name>A0A452GZK5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">Nuclear receptor subfamily 1 group I member 3</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="18">Constitutive androstane receptor</fullName>
        </alternativeName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="16">Binds and transactivates the retinoic acid response elements that control expression of the retinoic acid receptor beta 2 and alcohol dehydrogenase 3 genes. Transactivates both the phenobarbital responsive element module of the human CYP2B6 gene and the CYP3A4 xenobiotic response element.</text>
    </comment>
    <comment type="function">
        <text evidence="1">Nuclear hormone receptor that can act as a repressor or activator of transcription. High affinity receptor for thyroid hormones, including triiodothyronine and thyroxine.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the nuclear hormone receptor family. NR1 subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZK5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008892.1">
        <property type="protein sequence ID" value="ENSGAGP00000007707.1"/>
        <property type="gene ID" value="ENSGAGG00000006024.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004879">
        <property type="term" value="F:nuclear receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000122">
        <property type="term" value="P:negative regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045944">
        <property type="term" value="P:positive regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.50.10">
        <property type="entry name" value="Erythroid Transcription Factor GATA-1, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.565.10">
        <property type="entry name" value="Retinoid X Receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035500">
        <property type="entry name" value="NHR-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000536">
        <property type="entry name" value="Nucl_hrmn_rcpt_lig-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050234">
        <property type="entry name" value="Nuclear_hormone_rcpt_NR1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001723">
        <property type="entry name" value="Nuclear_hrmn_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001728">
        <property type="entry name" value="ThyrH_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001628">
        <property type="entry name" value="Znf_hrmn_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013088">
        <property type="entry name" value="Znf_NHR/GATA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24082">
        <property type="entry name" value="NUCLEAR HORMONE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24082:SF231">
        <property type="entry name" value="NUCLEAR RECEPTOR SUBFAMILY 1 GROUP I MEMBER 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00105">
        <property type="entry name" value="zf-C4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00398">
        <property type="entry name" value="STRDHORMONER"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00546">
        <property type="entry name" value="THYROIDHORMR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00399">
        <property type="entry name" value="ZnF_C4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57716">
        <property type="entry name" value="Glucocorticoid receptor-like (DNA-binding domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48508">
        <property type="entry name" value="Nuclear receptor ligand-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51843">
        <property type="entry name" value="NR_LBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51030">
        <property type="entry name" value="NUCLEAR_REC_DBD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="11" id="KW-0010">Activator</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="14" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="10" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="6" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="15" id="KW-0539">Nucleus</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="13" id="KW-0675">Receptor</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="12" id="KW-0804">Transcription</keyword>
    <keyword evidence="9" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="8" id="KW-0862">Zinc</keyword>
    <keyword evidence="7" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="Nuclear receptor" evidence="20">
        <location>
            <begin position="1"/>
            <end position="62"/>
        </location>
    </feature>
    <feature type="domain" description="NR LBD" evidence="21">
        <location>
            <begin position="92"/>
            <end position="281"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="248"/>
            <end position="281"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00003008"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00008092"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00037362"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00040912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00043125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS51030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS51843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007707.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="281" mass="31519" checksum="FE9C1063CAA7EC1B" modified="2019-05-08" version="1">MGVSPAPQTLPLFCRRSISKGVCFTCPFTRNCTITKAKRRQCQACRLQKCLAVGMRKDMIMSEEALQMRRVLRKRKQQEREPAGEPAELTEEQEQLISILIEAHKRHFDSGFSQFVHCRPPVRLYIHRDMLPDIFSMLPHFADLSTILIQQVIKFAKEIPAFRSPHLAASLLYPSALPCCIPPAPHPTQPCSAASIQHPAPSALPTGFQQIYLEPLLKFHISLRKLRLHEAEYVLLLAIALFSPGEPSPSPHHGLRDTPPTLPPRWALISPSPHHGLGGPP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZK6</accession>
    <name>A0A452GZK6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="19">Septin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="5">Chromosome</location>
            <location evidence="5">Centromere</location>
            <location evidence="5">Kinetochore</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cleavage furrow</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Cytoskeleton</location>
            <location evidence="3">Cilium axoneme</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Spindle</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Midbody</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="19 20">Belongs to the TRAFAC class TrmE-Era-EngA-EngB-Septin-like GTPase superfamily. Septin GTPase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZK6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008665.1">
        <property type="protein sequence ID" value="ENSGAGP00000007502.1"/>
        <property type="gene ID" value="ENSGAGG00000006002.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005930">
        <property type="term" value="C:axoneme"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032154">
        <property type="term" value="C:cleavage furrow"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000776">
        <property type="term" value="C:kinetochore"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030496">
        <property type="term" value="C:midbody"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031105">
        <property type="term" value="C:septin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005819">
        <property type="term" value="C:spindle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005525">
        <property type="term" value="F:GTP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd01850">
        <property type="entry name" value="CDC_Septin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000162">
        <property type="entry name" value="septin-7 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR030379">
        <property type="entry name" value="G_SEPTIN_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016491">
        <property type="entry name" value="Septin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008115">
        <property type="entry name" value="Septin7"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18884">
        <property type="entry name" value="SEPTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00735">
        <property type="entry name" value="Septin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF006698">
        <property type="entry name" value="Septin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01742">
        <property type="entry name" value="SEPTIN7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51719">
        <property type="entry name" value="G_SEPTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="17" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="8" id="KW-0132">Cell division</keyword>
    <keyword evidence="16" id="KW-0966">Cell projection</keyword>
    <keyword evidence="18" id="KW-0137">Centromere</keyword>
    <keyword evidence="6" id="KW-0158">Chromosome</keyword>
    <keyword evidence="13" id="KW-0969">Cilium</keyword>
    <keyword evidence="12 21" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="15" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="14 20" id="KW-0342">GTP-binding</keyword>
    <keyword evidence="11" id="KW-0995">Kinetochore</keyword>
    <keyword evidence="10" id="KW-0498">Mitosis</keyword>
    <keyword evidence="9 20" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Septin-type G" evidence="22">
        <location>
            <begin position="27"/>
            <end position="296"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="21">
        <location>
            <begin position="316"/>
            <end position="401"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004186"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004214"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004430"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004626"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004629"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022454"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022776"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022838"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023069"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023328"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="PIRNR" id="PIRNR006698"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="RuleBase" id="RU004560"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS51719"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007502.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="422" mass="49172" checksum="80EC9D61675BDDDC" modified="2019-05-08" version="1">MGQQKNLEGYVGFANLPNQVYRKSVKRGFEFTLMVVGESGLGKSTLINSLFLTDLYSPDYPGPSHRIKKTVQVEQSKVLIKEGGVQLLLTIVDTPGFGDAVDNSNCWQPVIDYIDSKFEDYLNAESRVNRRQMPDNRVQCCLYFIAPSGHGLKPLDIEFMKRLHEKVNIIPLIAKADTLTPEECQQFKKQIMKEIQEHKIKIYEFPETDDEEENKLVKKIKDRLPLAVVGSNTIIEVNGKRVRGRQYPWGVAEVENGEHCDFTILRNMLIRTHMQDLKDVTNNVHYENYRSRKLAAVTYNGVDNNKNKGQLTKSPLAQMEEERREHVAKMKKMEMEMEQVFEMKVKEKVQKLKDSEAELQRRHEQMKKNLEAQHKELEEKRRQFEDEKANWETQQRILEQQNSSRYVGFLLKLFVSNVPHVL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLE0</accession>
    <name>A0A452HLE0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Carbonic anhydrase</fullName>
            <ecNumber evidence="13">4.2.1.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">Reversible hydration of carbon dioxide. Its role in saliva is unknown.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>hydrogencarbonate + H(+) = CO2 + H2O</text>
            <dbReference type="Rhea" id="RHEA:10748"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:17544"/>
            <dbReference type="EC" id="4.2.1.1"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 13">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 13">Belongs to the alpha-carbonic anhydrase family.</text>
    </comment>
    <comment type="caution">
        <text evidence="12">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="EC" id="4.2.1.1" evidence="13"/>
    <dbReference type="AlphaFoldDB" id="A0A452HLE0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017982.1">
        <property type="protein sequence ID" value="ENSGAGP00000015751.1"/>
        <property type="gene ID" value="ENSGAGG00000011854.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004089">
        <property type="term" value="F:carbonate dehydratase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="CDD" id="cd03125">
        <property type="entry name" value="alpha_CA_VI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.200.10:FF:000003">
        <property type="entry name" value="Carbonic anhydrase 12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.200">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.200.10">
        <property type="entry name" value="Alpha carbonic anhydrase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001148">
        <property type="entry name" value="CA_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036398">
        <property type="entry name" value="CA_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023561">
        <property type="entry name" value="Carbonic_anhydrase_a-class"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018338">
        <property type="entry name" value="Carbonic_anhydrase_a-class_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001759">
        <property type="entry name" value="PTX_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18952">
        <property type="entry name" value="CARBONIC ANHYDRASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18952:SF110">
        <property type="entry name" value="CARBONIC ANHYDRASE 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00194">
        <property type="entry name" value="Carb_anhydrase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00354">
        <property type="entry name" value="Pentaxin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00895">
        <property type="entry name" value="PENTAXIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM01057">
        <property type="entry name" value="Carb_anhydrase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00159">
        <property type="entry name" value="PTX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51069">
        <property type="entry name" value="Carbonic anhydrase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00162">
        <property type="entry name" value="ALPHA_CA_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51144">
        <property type="entry name" value="ALPHA_CA_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51828">
        <property type="entry name" value="PTX_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="8" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9 13" id="KW-0456">Lyase</keyword>
    <keyword evidence="5 13" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="6 13" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="Alpha-carbonic anhydrase" evidence="14">
        <location>
            <begin position="1"/>
            <end position="247"/>
        </location>
    </feature>
    <feature type="domain" description="Pentraxin (PTX)" evidence="15">
        <location>
            <begin position="292"/>
            <end position="489"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00010718"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00025355"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00048348"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01172"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="RuleBase" id="RU367011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS51144"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS51828"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015751.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="490" mass="54990" checksum="C183D55A8E8528DB" modified="2019-05-08" version="1">MHWGKHFAACAGKHQSPIDIQKRKVRYNPQLMQLELSGYDGPLQGNFTMTNNGHSVQIDLPPTLRITKGLPSLFTAVQMHLHWGGLDLETSGSEHTVDGMRYFAELHIVHYNSDNYSSFEEAKDKWNGLAVLAFLYVDGNFENTYYSEFISNLAKIRFAGQSAKLSSLDVQAMLPENLSHFYRYQGSLTTPPCSENVIWTVFDSPIILSHTQIKLLENTLLDWQNNTLRNDYHHAQPLNDRVVESSFKAKGTKELCLPEELSSKLEQIQTQLQEMKKHLLAGVGLTGLKSSAFPAFYFPAENIASYVAVQPLRDMKLQAFTLCFWIQTQNRGSQTILAYRTQERDNELVVTVGSAVGLWLGGHFVEFPLHHKSEEWLHYCVAWASHSGAANLWLNGAVGREKHLQRGYVSQAGGTLVLGKDRDELLGIFSNGFSGWMSHVSLWSWVISHADIRVLALCRCSELKGDVIAWGETPMSLSGGVVLESDPSCQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLE1</accession>
    <name>A0A452HLE1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">phosphopyruvate hydratase</fullName>
            <ecNumber evidence="4">4.2.1.11</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="10">2-phospho-D-glycerate hydro-lyase</fullName>
        </alternativeName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>(2R)-2-phosphoglycerate = phosphoenolpyruvate + H2O</text>
            <dbReference type="Rhea" id="RHEA:10164"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:58289"/>
            <dbReference type="ChEBI" id="CHEBI:58702"/>
            <dbReference type="EC" id="4.2.1.11"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="14">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
        <text evidence="14">Mg(2+) is required for catalysis and for stabilizing the dimer.</text>
    </comment>
    <comment type="pathway">
        <text evidence="2">Carbohydrate degradation; glycolysis; pyruvate from D-glyceraldehyde 3-phosphate: step 4/5.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the enolase family.</text>
    </comment>
    <dbReference type="EC" id="4.2.1.11" evidence="4"/>
    <dbReference type="AlphaFoldDB" id="A0A452HLE1"/>
    <dbReference type="STRING" id="38772.ENSGAGP00000015726"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017956.1">
        <property type="protein sequence ID" value="ENSGAGP00000015726.1"/>
        <property type="gene ID" value="ENSGAGG00000011847.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000017971.1">
        <property type="protein sequence ID" value="ENSGAGP00000015741.1"/>
        <property type="gene ID" value="ENSGAGG00000011847.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000017974.1">
        <property type="protein sequence ID" value="ENSGAGP00000015743.1"/>
        <property type="gene ID" value="ENSGAGG00000011847.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00109">
        <property type="reaction ID" value="UER00187"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000015">
        <property type="term" value="C:phosphopyruvate hydratase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000287">
        <property type="term" value="F:magnesium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004634">
        <property type="term" value="F:phosphopyruvate hydratase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006096">
        <property type="term" value="P:glycolytic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="CDD" id="cd03313">
        <property type="entry name" value="enolase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.390.10:FF:000001">
        <property type="entry name" value="Enolase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.20.20.120:FF:000002">
        <property type="entry name" value="Enolase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.20.20.120">
        <property type="entry name" value="Enolase-like C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.390.10">
        <property type="entry name" value="Enolase-like, N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="HAMAP" id="MF_00318">
        <property type="entry name" value="Enolase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000941">
        <property type="entry name" value="Enolase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036849">
        <property type="entry name" value="Enolase-like_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029017">
        <property type="entry name" value="Enolase-like_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020810">
        <property type="entry name" value="Enolase_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020809">
        <property type="entry name" value="Enolase_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020811">
        <property type="entry name" value="Enolase_N"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01060">
        <property type="entry name" value="eno"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11902:SF12">
        <property type="entry name" value="ALPHA-ENOLASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11902">
        <property type="entry name" value="ENOLASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00113">
        <property type="entry name" value="Enolase_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03952">
        <property type="entry name" value="Enolase_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF001400">
        <property type="entry name" value="Enolase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00148">
        <property type="entry name" value="ENOLASE"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDF00002">
        <property type="entry name" value="enolase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDG00178">
        <property type="entry name" value="enolase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDS00001">
        <property type="entry name" value="Enolase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01192">
        <property type="entry name" value="Enolase_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01193">
        <property type="entry name" value="Enolase_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51604">
        <property type="entry name" value="Enolase C-terminal domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54826">
        <property type="entry name" value="Enolase N-terminal domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00164">
        <property type="entry name" value="ENOLASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0324">Glycolysis</keyword>
    <keyword evidence="9" id="KW-0456">Lyase</keyword>
    <keyword evidence="7 14" id="KW-0460">Magnesium</keyword>
    <keyword evidence="6 14" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Enolase N-terminal" evidence="16">
        <location>
            <begin position="3"/>
            <end position="134"/>
        </location>
    </feature>
    <feature type="domain" description="Enolase C-terminal TIM barrel" evidence="15">
        <location>
            <begin position="142"/>
            <end position="431"/>
        </location>
    </feature>
    <feature type="active site" description="Proton donor" evidence="12">
        <location>
            <position position="210"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="12">
        <location>
            <position position="343"/>
        </location>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <position position="158"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <position position="167"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="245"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="293"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <position position="293"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="318"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <position position="318"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <begin position="370"/>
            <end position="373"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <position position="394"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005031"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00009604"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00012058"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023152"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00031125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00048333"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PIRSR" id="PIRSR001400-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PIRSR" id="PIRSR001400-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PIRSR" id="PIRSR001400-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="SMART" id="SM01192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="SMART" id="SM01193"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015743.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="434" mass="47301" checksum="80FA1B65D4CCCF73" modified="2019-05-08" version="1">MSILKISAREIFDSRGNPTVEVDLFTNKGLFRAAVPSGASTGIYEALELRDNDKTRFLGKGVSKAVGHVNKTIAPALISKNINVVEQEKIDKLMLEMDGSENKSKFGANAILGVSLAVCKAGAAEKGVPLYRHIADLAGNPEVILPVPAFNVINGGSHAGNKLAMQEFMILPVGAESFKEAMRIGAEVYHNLKNVIKEKYGKDATNVGDEGGFAPNILENKEALELLKTAISKAGYTDKVVIGMDVAASEFYRDGKYDLDFKSPDDPSRYITPDQLADLYKSFIKSYPLVSVEDPFDQDDWSAWKKFTASVDVQVVGDDLTVTNPKRIAKAVEEKSCNCLLLKVNQIGSVTESLQACKLAQSNGWGVMVSHRSGETEDTFIADLVVGLCTGQIKTGAPCRSERLAKYNQLLRIEEELGSKARFAGRNFRNPRIN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLE2</accession>
    <name>A0A452HLE2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Dedicator of cytokinesis 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the DOCK family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLE2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017988.1">
        <property type="protein sequence ID" value="ENSGAGP00000015757.1"/>
        <property type="gene ID" value="ENSGAGG00000011372.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031267">
        <property type="term" value="F:small GTPase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007520">
        <property type="term" value="P:myoblast fusion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007264">
        <property type="term" value="P:small GTPase-mediated signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd08694">
        <property type="entry name" value="C2_Dock-A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000044">
        <property type="entry name" value="dedicator of cytokinesis protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1270.350:FF:000001">
        <property type="entry name" value="dedicator of cytokinesis protein 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.350">
        <property type="entry name" value="Dedicator of cytokinesis N-terminal subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027007">
        <property type="entry name" value="C2_DOCK-type_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026791">
        <property type="entry name" value="DOCK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047026">
        <property type="entry name" value="DOCK1_C2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032376">
        <property type="entry name" value="DOCK_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042455">
        <property type="entry name" value="DOCK_N_sub1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056372">
        <property type="entry name" value="TPR_DOCK"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45653">
        <property type="entry name" value="DEDICATOR OF CYTOKINESIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45653:SF1">
        <property type="entry name" value="DEDICATOR OF CYTOKINESIS PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14429">
        <property type="entry name" value="DOCK-C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16172">
        <property type="entry name" value="DOCK_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00018">
        <property type="entry name" value="SH3_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23554">
        <property type="entry name" value="TPR_DOCK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00452">
        <property type="entry name" value="SH3DOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00326">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51650">
        <property type="entry name" value="C2_DOCK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 4" id="KW-0728">SH3 domain</keyword>
    <feature type="domain" description="SH3" evidence="6">
        <location>
            <begin position="7"/>
            <end position="68"/>
        </location>
    </feature>
    <feature type="domain" description="C2 DOCK-type" evidence="7">
        <location>
            <begin position="423"/>
            <end position="607"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00983"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS51650"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015757.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="10">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="711" mass="81406" checksum="CDE8AF196FBC064D" modified="2019-05-08" version="1">NPCWAPLNYFLFLAFYNYDARGADELSLQIGDTVHILETYEGWYRGYTLRKKSKKGIFPASYIQLKEAIVEGKGQHETVIPSELPLIQEVTTTLREWSIIWRQLYVQDNRDMFHSVRHMIYDLIEWRSQILSGTLPQDELKELKKKVTAKIDYGNRILDLDLVVRDEDGNILDPEQTSTISLFRAHEIASKQVEERLLEEKSQKQNIDINRQAKFAATPSFALFVNLKNVVCKIGEDAEVLMSLYDPLESKFISENYLVRWSSSGLPKDIDRLHNLRAVFTDLGSKDLKREKISFVCQIVRVGRMELRDNNTRKLTSGLRRPFGVAVMDVTDIINGKVDDEDKQHFIPFQPVAGENDFLQTVINKVIAAKEVNHKGQGLWVTLKLLPGDIHQIRKEFPHLVDRSTAVARKMGFPEIIMPGDVRNDIYVTLVQGDFDKGSKTTAKNVEVTVSVYDEDGKRLENVIFPGAGDDAISEYKSVIYYQVKQPRWFETVKVAIPIEDVNRSHLRFTFRHRSSQDSKDKSEKIFALAFVKLMRYDGTTLRDGEHDLIVYKAEAKKLEDASTYLSLPSTKIELEEKGHATTGKSMQNLGSCTISKDSFQISTLVCSTKLTQNVDLLGLLKWRSNTNLLQQNLRQLMKVDGGEVVKFLQDTLDALFNIMMENSESETFDTLVFDALVFIIGLIADRKFQHFNPVLETYIKKHFSATLAYT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLE3</accession>
    <name>A0A452HLE3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">C2 DOCK-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the DOCK family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLE3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017993.1">
        <property type="protein sequence ID" value="ENSGAGP00000015761.1"/>
        <property type="gene ID" value="ENSGAGG00000011372.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031267">
        <property type="term" value="F:small GTPase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007520">
        <property type="term" value="P:myoblast fusion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007264">
        <property type="term" value="P:small GTPase-mediated signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd08694">
        <property type="entry name" value="C2_Dock-A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000044">
        <property type="entry name" value="dedicator of cytokinesis protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1270.350:FF:000001">
        <property type="entry name" value="dedicator of cytokinesis protein 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.350">
        <property type="entry name" value="Dedicator of cytokinesis N-terminal subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027007">
        <property type="entry name" value="C2_DOCK-type_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026791">
        <property type="entry name" value="DOCK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047026">
        <property type="entry name" value="DOCK1_C2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032376">
        <property type="entry name" value="DOCK_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042455">
        <property type="entry name" value="DOCK_N_sub1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056372">
        <property type="entry name" value="TPR_DOCK"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45653">
        <property type="entry name" value="DEDICATOR OF CYTOKINESIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45653:SF1">
        <property type="entry name" value="DEDICATOR OF CYTOKINESIS PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14429">
        <property type="entry name" value="DOCK-C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16172">
        <property type="entry name" value="DOCK_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23554">
        <property type="entry name" value="TPR_DOCK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51650">
        <property type="entry name" value="C2_DOCK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="C2 DOCK-type" evidence="4">
        <location>
            <begin position="367"/>
            <end position="551"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00983"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS51650"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015761.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="623" mass="71163" checksum="3BB19FE71C835BE4" modified="2019-05-08" version="1">KKSKKVNVLFPVQIHFLLQHETVIPSELPLIQEVTTTLREWSIIWRQLYVVSKIHMFHSVRHMIYDLIEWRSQILSGTLPQDELKELKKKVTAKIDYGNRILDLDLVVRDEDGNILDPEQTSTISLFRAHEIASKQVEERLLEEKSQKQNIDINRQAKFAATPSFALFVNLKNVVCKIGEDAEVLMSLYDPLESKFISENYLVRWSSSGLPKDIDRLHNLRAVFTDLGSKDLKREKISFVCQIVRVGRMELRDNNTRKLTSGLRRPFGVAVMDVTDIINGKVDDEDKQHFIPFQPVAGENDFLQTVINKVIAAKEVNHKGQGLWVTLKLLPGDIHQIRKEFPHLVDRSTAVARKMGFPEIIMPGDVRNDIYVTLVQGDFDKGSKTTAKNVEVTVSVYDEDGKRLENVIFPGAGDDAISEYKSVIYYQVKQPRWFETVKVAIPIEDVNRSHLRFTFRHRSSQDSKDKSEKIFALAFVKLMRYDGTTLRDGEHDLIVYKAEAKKLEDASTYLSLPSTKIELEEKGHATTGKSMQNLGSCTISKDSFQISTLVCSTKLTQNVDLLGLLKWRSNTNLLQQNLRQLMKVDGGEVVKFLQDTLDALFNIMMENSESETFDTLVFDALVS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLE4</accession>
    <name>A0A452HLE4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Solute carrier family 2, facilitated glucose transporter member 5</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="13">Fructose transporter</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="12">Glucose transporter type 5, small intestine</fullName>
        </alternativeName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>D-fructose(out) = D-fructose(in)</text>
            <dbReference type="Rhea" id="RHEA:60372"/>
            <dbReference type="ChEBI" id="CHEBI:37721"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <location evidence="2">Sarcolemma</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cell membrane</location>
            <topology evidence="3">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the major facilitator superfamily. Sugar transporter (TC 2.A.1.1) family. Glucose transporter subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLE4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017987.1">
        <property type="protein sequence ID" value="ENSGAGP00000015756.1"/>
        <property type="gene ID" value="ENSGAGG00000011859.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042383">
        <property type="term" value="C:sarcolemma"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055056">
        <property type="term" value="F:D-glucose transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005353">
        <property type="term" value="F:fructose transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046323">
        <property type="term" value="P:D-glucose import"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070837">
        <property type="term" value="P:dehydroascorbic acid transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1990539">
        <property type="term" value="P:fructose import across plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd17432">
        <property type="entry name" value="MFS_GLUT_Class2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1250.20:FF:001511">
        <property type="entry name" value="Solute carrier family 2, facilitated glucose transporter member 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1250.20">
        <property type="entry name" value="MFS general substrate transporter like domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045263">
        <property type="entry name" value="GLUT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020846">
        <property type="entry name" value="MFS_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005828">
        <property type="entry name" value="MFS_sugar_transport-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036259">
        <property type="entry name" value="MFS_trans_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003663">
        <property type="entry name" value="Sugar/inositol_transpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005829">
        <property type="entry name" value="Sugar_transporter_CS"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00879">
        <property type="entry name" value="SP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23503">
        <property type="entry name" value="SOLUTE CARRIER FAMILY 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23503:SF32">
        <property type="entry name" value="SOLUTE CARRIER FAMILY 2, FACILITATED GLUCOSE TRANSPORTER MEMBER 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00083">
        <property type="entry name" value="Sugar_tr"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00171">
        <property type="entry name" value="SUGRTRNSPORT"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103473">
        <property type="entry name" value="MFS general substrate transporter"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50850">
        <property type="entry name" value="MFS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00216">
        <property type="entry name" value="SUGAR_TRANSPORT_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00217">
        <property type="entry name" value="SUGAR_TRANSPORT_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11 15" id="KW-0472">Membrane</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0762">Sugar transport</keyword>
    <keyword evidence="9 15" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="10 15" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="6 14" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="21"/>
            <end position="41"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="76"/>
            <end position="97"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="109"/>
            <end position="129"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="135"/>
            <end position="157"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="169"/>
            <end position="187"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="199"/>
            <end position="220"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="285"/>
            <end position="306"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="321"/>
            <end position="343"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="350"/>
            <end position="372"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="378"/>
            <end position="396"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="417"/>
            <end position="438"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="444"/>
            <end position="467"/>
        </location>
    </feature>
    <feature type="domain" description="Major facilitator superfamily (MFS) profile" evidence="16">
        <location>
            <begin position="28"/>
            <end position="471"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000590"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004135"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00007004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00015973"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022597"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00029961"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00031099"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="RuleBase" id="RU003346"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50850"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015756.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="519" mass="57076" checksum="66B6375067692F68" modified="2019-05-08" version="1">MPAETGPEPSQTCTPRTGKMTLLLAGVTLVSAFGSSFQYGYNVSVINSPAVFMQEFYNHTYLDRKGIPMESSFQTLLWSLTVSMYPLGGFFGSLMVGPLVNKCGRKGTLLINNIFSIVPAILMGTSKVAKTFEAIILSRIIVGICAGLSSNVVPMYLGEMSPKNLRGAVGVVPQLFITIGILIAQILGLTDILGNVEGWPVLLGLTGIPAVVELLLLPFFPESPRYLLIQKGNEEQARQALKRLRGWDDVDDEIEEMRREDQSEKLEGRLSVFNLFTFRALRWQLISIIVMMMGQQLSGINAIFYYANDIFRSAGVNENNVQYVTVGIGAVNVFMTCVAVFIIESLGRRLLLLAGFGICCVSSAVLTVALNLQATVSWMQYLSIACVIVYIIGHAIGASPIPSVMITEMFLQSSRPAAFMVGGSVHWLCNFTVGLVFLYMVEGLGAYCFLVFCAICLATVIYIFIIVPETKGKTFMEINQSMAKRNKVEIQMDKEKLNDFQPAFETLKKNEEGYSITGL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLF4</accession>
    <name>A0A452HLF4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">C2 DOCK-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the DOCK family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000018001.1">
        <property type="protein sequence ID" value="ENSGAGP00000015768.1"/>
        <property type="gene ID" value="ENSGAGG00000011372.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031267">
        <property type="term" value="F:small GTPase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007520">
        <property type="term" value="P:myoblast fusion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007264">
        <property type="term" value="P:small GTPase-mediated signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd08694">
        <property type="entry name" value="C2_Dock-A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000044">
        <property type="entry name" value="dedicator of cytokinesis protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1270.350:FF:000001">
        <property type="entry name" value="dedicator of cytokinesis protein 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.350">
        <property type="entry name" value="Dedicator of cytokinesis N-terminal subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027007">
        <property type="entry name" value="C2_DOCK-type_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026791">
        <property type="entry name" value="DOCK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047026">
        <property type="entry name" value="DOCK1_C2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032376">
        <property type="entry name" value="DOCK_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042455">
        <property type="entry name" value="DOCK_N_sub1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45653">
        <property type="entry name" value="DEDICATOR OF CYTOKINESIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45653:SF1">
        <property type="entry name" value="DEDICATOR OF CYTOKINESIS PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14429">
        <property type="entry name" value="DOCK-C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16172">
        <property type="entry name" value="DOCK_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51650">
        <property type="entry name" value="C2_DOCK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="C2 DOCK-type" evidence="4">
        <location>
            <begin position="438"/>
            <end position="622"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00983"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS51650"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015768.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="672" mass="76878" checksum="B0EB44D718D6D7AA" modified="2019-05-08" version="1">NPCWAPLNYFLFLAFYNYDARGADELSLQIGDTVHILETYEAITNRVYKLDYINHFTILPKFVFANFRQHETVIPSELPLIQEVTTTLREWSIIWRQLYVVNNRDMFHSVRHMIYDLIEWRSQILSGTLPQDELKELKKKVTAKIDYGNRILDLDLVVRDEDGNILDPEQTSTISLFRAHEIASKQVEERLLEEKSQKQNIDINRQAKFAATPSFALFVNLKNVVCKIGEDAEVLMSLYDPLESKFISENYLVRWSSSGLPKDIDRLHNLRAVFTDLGSKDLKREKISFVCQIVRVGRMELRDNNTRKLTSGLRRPFGVAVMDVTDIINGKVDDEDKQHFIPFQPLALDDAIRHKQLNISSRFSPRVAGENDFLQTVINKVIAAKEVNHKGQGLWVTLKLLPGDIHQIRKEFPHLVDRSTAVARKMGFPEIIMPGDVRNDIYVTLVQGDFDKGSKTTAKNVEVTVSVYDEDGKRLENVIFPGAGDDAISEYKSVIYYQVKQPRWFETVKVAIPIEDVNRSHLRFTFRHRSSQDSKDKSEKIFALAFVKLMRYDGTTLRDGEHDLIVYKAEAKKLEDASTYLSLPSTKIELEEKGHATTGKSMQNLGSCTISKDSFQISTLVCSTKLTQNGELLEEMFLHHMCPLNFPPFITVPDQMKGLLXGRAAATPCLLS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLF9</accession>
    <name>A0A452HLF9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="8">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLF9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018006.1">
        <property type="protein sequence ID" value="ENSGAGP00000015773.1"/>
        <property type="gene ID" value="ENSGAGG00000011846.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042995">
        <property type="term" value="C:cell projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005332">
        <property type="term" value="F:gamma-aminobutyric acid:sodium:chloride symporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000175">
        <property type="entry name" value="Na/ntran_symport"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037272">
        <property type="entry name" value="SNS_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11616:SF124">
        <property type="entry name" value="SODIUM- AND CHLORIDE-DEPENDENT GABA TRANSPORTER 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11616">
        <property type="entry name" value="SODIUM/CHLORIDE DEPENDENT TRANSPORTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00209">
        <property type="entry name" value="SNF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00176">
        <property type="entry name" value="NANEUSMPORT"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF161070">
        <property type="entry name" value="SNF-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50267">
        <property type="entry name" value="NA_NEUROTRAN_SYMP_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5 7" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0915">Sodium</keyword>
    <keyword evidence="3 7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 7" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="2" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="20"/>
            <end position="40"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="72"/>
            <end position="93"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="105"/>
            <end position="131"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="164"/>
            <end position="196"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="208"/>
            <end position="233"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="239"/>
            <end position="261"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="281"/>
            <end position="303"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="323"/>
            <end position="344"/>
        </location>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="79"/>
        </location>
        <ligand>
            <name>Na(+)</name>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="111"/>
        </location>
        <ligand>
            <name>Na(+)</name>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="176"/>
        </location>
        <ligand>
            <name>Na(+)</name>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="179"/>
        </location>
        <ligand>
            <name>Na(+)</name>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="180"/>
        </location>
        <ligand>
            <name>Na(+)</name>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
            <label>1</label>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PIRSR" id="PIRSR600175-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015773.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="402" mass="45216" checksum="D392900BAD20BAD3" modified="2019-05-08" version="1">MTSQGPFHLLHRLGAQDWRYFCVVYVTATFPYVMLLILLVRGVTLPGASEGIKFYLYPDLSRLSDPQVWVDAGTQIFFSYAICLGCLTALGSYNNYNNNCYRDCIMLCCLNSGTSFVAGFAIFSVLGFMAYEQGVPIAEVAESGPGLAFIAYPKAVTMMPLSPLWAALFFMMLIFLGLDSQFVCVESLVTAVVDMYPKIFRRGYRRELLILALSIVSYFVGLIMLTEGGMYIFQLFDSYAASGMCLLFVAIFECICIGWIYGSNRFYDNIEDMIGYRPLSLIKWCWMVLTPGICAGIFIFFLVKYKPLKYNNVYTYPDWGYGIGWMMALSSMVCIPLWICIKLWKTEGTVLEKFRKLVIPSPDLKMRGKFGAGAYAATVNDCDAKLKGDGNISAITEKETHF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLH8</accession>
    <name>A0A452HLH8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">MHC class I-like antigen recognition-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="10">Belongs to the MHC class I family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLH8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018025.1">
        <property type="protein sequence ID" value="ENSGAGP00000015789.1"/>
        <property type="gene ID" value="ENSGAGG00000011875.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009897">
        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042612">
        <property type="term" value="C:MHC class I protein complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002474">
        <property type="term" value="P:antigen processing and presentation of peptide antigen via MHC class I"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006955">
        <property type="term" value="P:immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.500.10:FF:000001">
        <property type="entry name" value="H-2 class I histocompatibility antigen, alpha chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.500.10">
        <property type="entry name" value="MHC class I-like antigen recognition-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050208">
        <property type="entry name" value="MHC_class-I_related"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011161">
        <property type="entry name" value="MHC_I-like_Ag-recog"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037055">
        <property type="entry name" value="MHC_I-like_Ag-recog_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011162">
        <property type="entry name" value="MHC_I/II-like_Ag-recog"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001039">
        <property type="entry name" value="MHC_I_a_a1/a2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16675:SF242">
        <property type="entry name" value="MAJOR HISTOCOMPATIBILITY COMPLEX CLASS I-RELATED GENE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16675">
        <property type="entry name" value="MHC CLASS I-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00129">
        <property type="entry name" value="MHC_I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01638">
        <property type="entry name" value="MHCCLASSI"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54452">
        <property type="entry name" value="MHC antigen-recognition domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="9" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="5" id="KW-0391">Immunity</keyword>
    <keyword evidence="7" id="KW-0472">Membrane</keyword>
    <keyword evidence="2" id="KW-0490">MHC I</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="3" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6" id="KW-1133">Transmembrane helix</keyword>
    <feature type="domain" description="MHC class I-like antigen recognition-like" evidence="11">
        <location>
            <begin position="30"/>
            <end position="204"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022451"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022859"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="RuleBase" id="RU004439"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF00129"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015789.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="232" mass="27008" checksum="FBEE1BE6FC390151" modified="2019-05-08" version="1">PVILARRSLGSLAGSLSLIHISPPTPPGFHSRRVLDTVVSEPGPGLPWYSRVVYVDDQVIYVYTSGMQRAEPRPVWMAQNEGPEFWDERTWWAQRHQKWFKASLNTLGQLYNQSKGFHIIQMKYGCDLREDNTTWGFYQDSYDGRDFLTFDTESMTLGAADIGAQITKRRWDAEVFDNQQWKRYLEERCIPWLRSSLEYGKETLQRKGKMGDKPYPEGHYKLYLHSPAPVPK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLH9</accession>
    <name>A0A452HLH9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">receptor protein-tyrosine kinase</fullName>
            <ecNumber evidence="3">2.7.10.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="27 28">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="28" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="27" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="27" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>L-tyrosyl-[protein] + ATP = O-phospho-L-tyrosyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10596"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.10.1"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the protein kinase superfamily. CAMK Ser/Thr protein kinase family.</text>
    </comment>
    <dbReference type="EC" id="2.7.10.1" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452HLH9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018026.1">
        <property type="protein sequence ID" value="ENSGAGP00000015790.1"/>
        <property type="gene ID" value="ENSGAGG00000010943.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043235">
        <property type="term" value="C:receptor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004714">
        <property type="term" value="F:transmembrane receptor protein tyrosine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007169">
        <property type="term" value="P:cell surface receptor protein tyrosine kinase signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006909">
        <property type="term" value="P:phagocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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    </dbReference>
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        <property type="entry name" value="MER proto-oncogene, tyrosine kinase"/>
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        <property type="entry name" value="Tyrosine-protein kinase receptor TYRO3"/>
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        <property type="entry name" value="Tyrosine-protein kinase receptor TYRO3"/>
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        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
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    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Ig-like_fold"/>
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    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
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        <property type="entry name" value="Immunoglobulin_dom"/>
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    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
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    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050122">
        <property type="entry name" value="RTK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001245">
        <property type="entry name" value="Ser-Thr/Tyr_kinase_cat_dom"/>
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    <dbReference type="InterPro" id="IPR008266">
        <property type="entry name" value="Tyr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020635">
        <property type="entry name" value="Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416:SF257">
        <property type="entry name" value="TYROSINE-PROTEIN KINASE MER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416">
        <property type="entry name" value="TYROSINE-PROTEIN KINASE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
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        <property type="match status" value="2"/>
    </dbReference>
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        <property type="entry name" value="ig"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Ig_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07714">
        <property type="entry name" value="PK_Tyr_Ser-Thr"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000615">
        <property type="entry name" value="TyrPK_CSF1-R"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00109">
        <property type="entry name" value="TYRKINASE"/>
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        <property type="match status" value="2"/>
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        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00219">
        <property type="entry name" value="TyrKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
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    </dbReference>
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        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
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    </dbReference>
    <dbReference type="PROSITE" id="PS00109">
        <property type="entry name" value="PROTEIN_KINASE_TYR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10 20 22" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="14" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="16" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="17" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="9" id="KW-0418">Kinase</keyword>
    <keyword evidence="21" id="KW-0460">Magnesium</keyword>
    <keyword evidence="12 23" id="KW-0472">Membrane</keyword>
    <keyword evidence="21" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8 20 22" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="15" id="KW-0675">Receptor</keyword>
    <keyword evidence="28" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0808">Transferase</keyword>
    <keyword evidence="6 23" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 23" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="13" id="KW-0829">Tyrosine-protein kinase</keyword>
    <feature type="transmembrane region" description="Helical" evidence="23">
        <location>
            <begin position="457"/>
            <end position="480"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="25">
        <location>
            <begin position="48"/>
            <end position="140"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="25">
        <location>
            <begin position="152"/>
            <end position="234"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="26">
        <location>
            <begin position="241"/>
            <end position="336"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="26">
        <location>
            <begin position="341"/>
            <end position="438"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="24">
        <location>
            <begin position="542"/>
            <end position="768"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="19">
        <location>
            <position position="678"/>
        </location>
    </feature>
    <feature type="binding site" evidence="22">
        <location>
            <position position="574"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="20">
        <location>
            <position position="682"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="21">
        <location>
            <position position="683"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="21">
        <location>
            <position position="696"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00011902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023137"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00051243"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="PIRSR" id="PIRSR000615-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="PIRSR" id="PIRSR000615-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="PIRSR" id="PIRSR000615-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="26">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015790.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="28">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="768" mass="85615" checksum="2CB36B9F60930966" modified="2019-05-08" version="1">RTAAWPCRTDHSSKYNCSCHTRVSLGSARTNVHVTSAAHSLQQVKFNPTVGHILINEHKDFKFNCSIKGPQFLFNQDSAVISLWKDGKELQEPGRMSTRHYQVTDDVETTMVSTFSITNAQRSDNGSYSCKLKIHNEEIESDPIFVQMEGLPHFLRQPQSLNITRNRPFNLTCEAMGPPEPVQIYWFRNSSRVNDQPYISPSVLTLPGLSETAIFSCEAHNRKGLTASNEVQINIKRTPAAPADVRVLNRTAHSLVVSWVPGFDGYSPLHSCRLQVREAAALNKGSFLIWNTSVPPHRYHVQQLQPLAEYSIRVSCRNEVGWSSFSPWTLASTTEGAPSTPPLNVTVFFNESSSSLEVRWAKPHLGRKHGELQGYRLLHTWQNSKITQNRSSEAQQNATVTTVAIVVTNATYAVRVAAVTKGGIGPFSDPVVVFIPGDGFMASAPSSTPAPGNNHSFAIVLGFILGIVAIGLILYFSVIVRKRWMETKFGTAFHSDDSEHVVSYTAKKSYSRRAVELTLASLGVREELQQKLQDVVIDRDSLALGKILGEGEFGSVMEGHLSQPSGTVQKVAVKTMKLEDFSQREIEEFLSEAACMKDFDHPNVIRLLGVCIELSSRQVPKPMVVLPFMKYGDLHSFLLRSRFETGTRHVPLQTLLKFMVDIALGMEYLSTRQFLHRDLAARNCMLRDDMTVCVADFGLSKKIYSGDYYRQGRIAKMPVKWIAIESLADRVYTTKSDVVRTPGAGSPTLKSELQSMVEVCNTASLWKP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLI0</accession>
    <name>A0A452HLI0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">non-specific serine/threonine protein kinase</fullName>
            <ecNumber evidence="4">2.7.11.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="15 22">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="14 22">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the protein kinase superfamily. AGC Ser/Thr protein kinase family.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018022.1">
        <property type="protein sequence ID" value="ENSGAGP00000015788.1"/>
        <property type="gene ID" value="ENSGAGG00000011873.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000287">
        <property type="term" value="F:magnesium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007010">
        <property type="term" value="P:cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032502">
        <property type="term" value="P:developmental process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035556">
        <property type="term" value="P:intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd23076">
        <property type="entry name" value="PDZ_MAST4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05609">
        <property type="entry name" value="STKc_MAST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:001045">
        <property type="entry name" value="Microtubule-associated serine/threonine kinase 1a"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000012">
        <property type="entry name" value="microtubule-associated serine/threonine-protein kinase 2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1480.20:FF:000001">
        <property type="entry name" value="microtubule-associated serine/threonine-protein kinase 4 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="microtubule-associated serine/threonine-protein kinase 4 isoform X2"/>
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        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
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        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
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        <property type="entry name" value="Kinase-like_dom_sf"/>
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        <property type="entry name" value="MAST"/>
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        <property type="entry name" value="MAST_pre-PK_dom_sf"/>
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        <property type="entry name" value="PDZ"/>
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        <property type="entry name" value="Prot_kinase_dom"/>
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        <property type="entry name" value="Ser/Thr_kinase_AS"/>
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        <property type="entry name" value="Ser_Thr_kinase_AGC"/>
    </dbReference>
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        <property type="entry name" value="MICROTUBULE-ASSOCIATED SERINE_THREONINE-PROTEIN KINASE 4"/>
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        <property type="entry name" value="SERINE/THREONINE-PROTEIN KINASE"/>
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        <property type="entry name" value="PROTEIN_KINASE_ST"/>
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    </dbReference>
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    <keyword evidence="12" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11" id="KW-0418">Kinase</keyword>
    <keyword evidence="13" id="KW-0460">Magnesium</keyword>
    <keyword evidence="9" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="8" id="KW-0808">Transferase</keyword>
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        <location>
            <begin position="462"/>
            <end position="735"/>
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            <begin position="736"/>
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            <begin position="2416"/>
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    <sequence length="2501" mass="275964" checksum="606F40692D46DB7B" modified="2019-05-08" version="1">MLFRKSSNPEVSSGPGKSFKFKRQLSEDGRPFRRGSLGGALTGRYLLPNTIAQQSWQPAETSNLVRMRSQALGQSAPSLTASLEGPLQKSHFSSTRHRQRLVEPAATKKELSLPRRGSFCRTSNRKSLIGNGQSPALPRPHSPLSAHAGSSPQDSPRNFSPSASAHFSFARRTDGRRWSLASLPSSGYGTNTPSSTVSSSCSSQEKLHQLPYQPTPDELHFLSKHFCTTESIASDNRCRTTPMRPRSRSLSPGRSPACCDHEIIMMNHVYKERFPKATAQMEERLQDIITNHSPDNILPLADGVLSFTHHQIIELARDCLDKSHQGLITSRYFFELQHKLDKLLQEGQERSESGELAFIKQLVRKILIVIARPARLLECLEFDPEEFYYLLEAAEGHAKEGHVIKTDIPRYIISQLGLNKDPLEEIAQLANYDSGTAETPETDESSSNTSVTLRRKPRESDFETIKLISNGAYGAVYFVRHKETRQRFAMKKINKQNLILRNQIQQAFVERDILTFAENPFVVSMYCSFETRRHLCMVMEYVEGGDCATLMKNMGPLPVDMARMYFAETVLALEYLHNYGIVHRDLKPDNLLVTSMGHIKLTDFGLSKVGLMSMTTNLYEGHIEKDAREFLDKQVCGTPEYIAPEVILRQGYGKPVDWWAMGIILYEFLVGCVPFFGDTPEELFGQVISDEINWPEKDEAPPPDSQDLITLLLRQNPMERLGTGGAYEVKQHQFFRSLDWNSLLRQKAEFIPQLESEDDTSYFDTRSEKYHHMETEEEDDTNDEDFNLEIRQFSSCSHRFSKVFSSIDRITRNQGEEKEDTGDKTKNVTLPSVESLSWSSEYSELQQLSTSNSSDTESTRLRHSSGLLPKLAISSEGDQDETTCSEGSREEQEKAASNEEAAQDEPEVTTPASTISSSTLSVGSFSEHLDQINGRNESVDSTDNSSKPSSEPASHMARQRLESTEKKKISGKVTKSLSASALSLMIPGDVFGVSPLGSPMSPHSLSSDLSSSRDSSPSRDSSVASATPHQPILIHSSGKKYGFTIRAIRVYVGDSDIYTVHHIVWNVEEGSPAYQAGLKAGDLITHINGEPVHGLVHTEVIELLLKSGNKVSITTTPFENTSIKTGPARRNSYRSRMVRRSKKTKKKESLERRRSLFKKLAKQPSPLLHTSRSFSCLNRSLSSGESLPGSPTHSLSPRSPTPSYRSTPDFPSGTNSSQSSSPSSSGPNSPAGSGHIRPSTLHGLGPKLGGQRYRSGRRKSAGSIPLSPLARTPSPTPQPPSPQRSPSPLLGHSIGNSKIAQAFPSKMHSPPTIVRHIVRPKSAEPPRSPLLKRVQSEEKLAPSYGCDKKHLCSRKHSLEVTQEEVNRELSQREVTLQSLEENVCDVPSLTRVRPAEQGCLKRPVCRKLGRQESVDELDREKLKVKIVMKKQDLAEKQDCLQKASTIHEPSGESENPAILRLEERDKKSFQKALERSNHFETKITALETPSVGGILKDTLHKNANMRPNDCIPLDTHMPCHGPPLNELSHLSYDFKRISPPCTLQDVLPHSSDRLLNGKGESTDKNAPTKEFVKFERLDGKLANIDYLRKKMSFEEKDDSLCPVLKPKLTSNVHECLQLNTVRPINVQQDSAAVADSRAFISSAHAAQISSVSFVPLKALNGRVETGLEKSALISTESPVRKSPSEYKLEGRSVSCLKPIEGTLDIALLSGPQASKTELPSCLLPESQNTSNEVGSSMPLASQCNGVKEETSCQKEVVSPHSKSSKSNLLEPQFSQSSKSFQNAPATPVTAELETGKKISSPDAACNISVREGDQKREDSHSKQKDHATEAKLHTVHNQCLHTALASATPSTAQGTPEKTAGKESQTQKELPVKYPVTQRNYESFSVKAEGIRDSSVKKSIPDAFKTSDSKTIQRLSADFAIPLQTHRTTRVQTAGSKVESKDGKDLPKQFAKDDITPAGSSVERDMNKQEVMDSKSEMAFPKRPVQQSVESNTKSEKSLVCSSAQKDNSKDLRKKDQKHSSNIQLHIIERKLPNLITRQQFSPSCFPSVRETVNRNSVIDAHVHLGVQEQVKTGSAYMESSNNKSRQINETCSSKSKHSDKFTFAQKQCTANIKEEETVIQPAAGSRKDGKHANKSVQDSHSFVLISQRKLNNEYIQVEKPLGLEHGSAPSIQINSSTPETKPKSSCNGQAPAGSENSKHFLSQETAGFSETADQKQLYVNEKQNMPPKFSTVKDCLLLNKLADKDQSNQSITADKRPEGQKCTEALYVQNDSGKLEPSLFLNCDLGKVAEKAPIVSKGSPDSKGKGQVSQKQLTDANKQITAMLLSTATVQSSCHSAVTPLKQLNCPPSFPESRQEASPLSSAKVKTGSPELPDTSCKELKKQATSSTGDGKREITKSVSLLTLHSSAIPVEALHPASNLGSKPGNQEKPLSASNTVDVKGEDPIQAQPPAAKKQNVGKDFPRSVTTSDRPTALSSDKDSVRQRRGKDASRSSPHKKSSQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLI1</accession>
    <name>A0A452HLI1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">non-specific serine/threonine protein kinase</fullName>
            <ecNumber evidence="4">2.7.11.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="15 22">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="14 22">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the protein kinase superfamily. AGC Ser/Thr protein kinase family.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018038.1">
        <property type="protein sequence ID" value="ENSGAGP00000015797.1"/>
        <property type="gene ID" value="ENSGAGG00000011873.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000287">
        <property type="term" value="F:magnesium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007010">
        <property type="term" value="P:cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032502">
        <property type="term" value="P:developmental process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035556">
        <property type="term" value="P:intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd23076">
        <property type="entry name" value="PDZ_MAST4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05609">
        <property type="entry name" value="STKc_MAST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:001045">
        <property type="entry name" value="Microtubule-associated serine/threonine kinase 1a"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000012">
        <property type="entry name" value="microtubule-associated serine/threonine-protein kinase 2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1480.20:FF:000001">
        <property type="entry name" value="microtubule-associated serine/threonine-protein kinase 4 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000008">
        <property type="entry name" value="microtubule-associated serine/threonine-protein kinase 4 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.42.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1480.20">
        <property type="entry name" value="MAST3 pre-PK domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000961">
        <property type="entry name" value="AGC-kinase_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037711">
        <property type="entry name" value="MAST"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015022">
        <property type="entry name" value="MAST_pre-PK_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023142">
        <property type="entry name" value="MAST_pre-PK_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001478">
        <property type="entry name" value="PDZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041489">
        <property type="entry name" value="PDZ_6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036034">
        <property type="entry name" value="PDZ_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050236">
        <property type="entry name" value="Ser_Thr_kinase_AGC"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24356:SF224">
        <property type="entry name" value="MICROTUBULE-ASSOCIATED SERINE_THREONINE-PROTEIN KINASE 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24356">
        <property type="entry name" value="SERINE/THREONINE-PROTEIN KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08926">
        <property type="entry name" value="DUF1908"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17820">
        <property type="entry name" value="PDZ_6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00228">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF140482">
        <property type="entry name" value="MAST3 pre-PK domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50156">
        <property type="entry name" value="PDZ domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51285">
        <property type="entry name" value="AGC_KINASE_CTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50106">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11" id="KW-0418">Kinase</keyword>
    <keyword evidence="13" id="KW-0460">Magnesium</keyword>
    <keyword evidence="9" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="8" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Protein kinase" evidence="17">
        <location>
            <begin position="366"/>
            <end position="639"/>
        </location>
    </feature>
    <feature type="domain" description="AGC-kinase C-terminal" evidence="19">
        <location>
            <begin position="640"/>
            <end position="712"/>
        </location>
    </feature>
    <feature type="domain" description="PDZ" evidence="18">
        <location>
            <begin position="935"/>
            <end position="1023"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1"/>
            <end position="110"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="339"/>
            <end position="359"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="715"/>
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        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="751"/>
            <end position="876"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
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            <begin position="903"/>
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    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
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            <begin position="1022"/>
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    </feature>
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    <feature type="region of interest" description="Disordered" evidence="16">
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    <feature type="region of interest" description="Disordered" evidence="16">
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        <location>
            <begin position="1832"/>
            <end position="1924"/>
        </location>
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    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="2070"/>
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    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="2250"/>
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        <location>
            <begin position="2320"/>
            <end position="2405"/>
        </location>
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        <location>
            <begin position="45"/>
            <end position="55"/>
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    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="16">
        <location>
            <begin position="56"/>
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        <source>
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        </source>
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        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
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    </evidence>
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    </evidence>
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    </evidence>
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    </evidence>
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    </evidence>
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            <dbReference type="ARBA" id="ARBA00047899"/>
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    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00048679"/>
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    <evidence type="ECO:0000256" key="16">
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            <dbReference type="SAM" id="MobiDB-lite"/>
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    <evidence type="ECO:0000259" key="17">
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            <dbReference type="PROSITE" id="PS50011"/>
        </source>
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    <evidence type="ECO:0000259" key="18">
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        </source>
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    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2405" mass="265483" checksum="67D430FE628EBDDA" modified="2019-05-08" version="1">MKAQRERLQIPGLTLDCRTSNRKSLIGNGQSPALPRPHSPLSAHAGSSPQDSPRNFSPSASAHFSFARSHGHRNDRTDGRRWSLASLPSSGYGTNTPSSTVSSSCSSQEKLHQLPYQPTPDELHFLSKHFCTTESIASDNRCRTTPMRPRSRSLSPGRSPACCDHEIIMMNHVYKERFPKATAQMEERLQDIITNHSPDNILPLADGVLSFTHHQIIELARDCLDKSHQGLITSRYFFELQHKLDKLLQEGQERSESGELAFIKQLVRKILIVIARPARLLECLEFDPEEFYYLLEAAEGHAKEGHVIKTDIPRYIISQLGLNKDPLEEIAQLANYDSGTAETPETDESSSNTSVTLRRKPRESDFETIKLISNGAYGAVYFVRHKETRQRFAMKKINKQNLILRNQIQQAFVERDILTFAENPFVVSMYCSFETRRHLCMVMEYVEGGDCATLMKNMGPLPVDMARMYFAETVLALEYLHNYGIVHRDLKPDNLLVTSMGHIKLTDFGLSKVGLMSMTTNLYEGHIEKDAREFLDKQVCGTPEYIAPEVILRQGYGKPVDWWAMGIILYEFLVGCVPFFGDTPEELFGQVISDEINWPEKDEAPPPDSQDLITLLLRQNPMERLGTGGAYEVKQHQFFRSLDWNSLLRQKAEFIPQLESEDDTSYFDTRSEKYHHMETEEEDDTNDEDFNLEIRQFSSCSHRFSKVFSSIDRITRNQGEEKEDTGDKTKNVTLPSVESLSWSSEYSELQQLSTSNSSDTESTRLRHSSGLLPKLAISSEGDQDETTCSEGSREEQEKAASNEEAAQDEPEVTTPASTISSSTLSVGSFSEHLDQINGRNESVDSTDNSSKPSSEPASHMARQRLESTEKKKISGKVTKSLSASALSLMIPGDVFGVSPLGSPMSPHSLSSDLSSSRDSSPSRDSSVASATPHQPILIHSSGKKYGFTIRAIRVYVGDSDIYTVHHIVWNVEEGSPAYQAGLKAGDLITHINGEPVHGLVHTEVIELLLKSGNKVSITTTPFENTSIKTGPARRNSYRSRMVRRSKKTKKKESLERRRSLFKKLAKQPSPLLHTSRSFSCLNRSLSSGESLPGSPTHSLSPRSPTPSYRSTPDFPSGTNSSQSSSPSSSGPNSPAGSGHIRPSTLHGLGPKLGGQRYRSGRRKSAGSIPLSPLARTPSPTPQPPSPQRSPSPLLGHSIGNSKIAQAFPSKMHSPPTIVRHIVRPKSAEPPRSPLLKRVQSEEKLAPSYGCDKKHLCSRKHSLEVTQEEVNRELSQREVTLQSLEENVCDVPSLTRVRPAEQGCLKRPVCRKLGRQESVDELDREKLKVKIVMKKQDLAEKQDCLQKASTIHEPSGESENPAILRLEERDKKSFQKALERSNHFETKITALETPSVGGILKDTLHKNANMRPNDCIPLDTHMPCHGPPLNELSHLSYDFKRISPPCTLQDVLPHSSDRLLNGKGESTDKNAPTKEFVKFERLDGKLANIDYLRKKMSFEEKDDSLCPVLKPKLTSNVHECLQLNTVRPINVQQDSAAVADSRAFISSAHAAQISSVSFVPLKALNGRVETGLEKSALISTESPVRKSPSEYKLEGRSVSCLKPIEGTLDIALLSGPQASKTELPSCLLPESQNTSNEVGSSMPLASQCNGVKEETSCQKEVVSPHSKSSKSNLLEPQFSQSSKSFQNAPATPVTAELETGKKISSPDAACNISVREGDQKREDSHSKQKDHATEAKLHTVHNQCLHTALASATPSTAQGTPEKTAGKESQTQKELPVKYPVTQRNYESFSVKAEGIRDSSVKKSIPDAFKTSDSKTIQRLSADFAIPLQTHRTTRVQTAGSKVESKDGKDLPKQFAKDDITPAGSSVERDMNKQEVMDSKSEMAFPKRPVQQSVESNTKSEKSLVCSSAQKDNSKDLRKKDQKHSSNIQLHIIERKLPNLITRQQFSPSCFPSVRETVNRNSVIDAHVHLGVQEQVKTGSAYMESSNNKSRQINETCSSKSKHSDKFTFAQKQCTANIKEEETVIQPAAGSRKDGKHANKSVQDSHSFVLISQRKLNNEYIQVEKPLGLEHGSAPSIQINSSTPETKPKSSCNGQAPAGSENSKHFLSQETAGFSETADQKQLYVNEKQNMPPKFSTVKDCLLLNKLADKDQSNQSITADKRPEGQKCTEALYVQNDSGKLEPSLFLNCDLGKVAEKAPIVSKGSPDSKGKGQVSQKQLTDANKQITAMLLSTATVQSSCHSAVTPLKQLNCPPSFPESRQEASPLSSAKVKTGSPELPDTSCKELKKQATSSTGDGKREITKSVSLLTLHSSAIPVEALHPASNLGSKPGNQEKPLSASNTVDVKGEDPIQAQPPAAKKQNVGKDFPRSVTTSDRPTALSSDKDSVRQRRGKDASRSSPHKKSSQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLI5</accession>
    <name>A0A452HLI5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">non-specific serine/threonine protein kinase</fullName>
            <ecNumber evidence="4">2.7.11.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="15 22">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="14 22">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the protein kinase superfamily. AGC Ser/Thr protein kinase family.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018030.1">
        <property type="protein sequence ID" value="ENSGAGP00000015793.1"/>
        <property type="gene ID" value="ENSGAGG00000011873.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000287">
        <property type="term" value="F:magnesium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007010">
        <property type="term" value="P:cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032502">
        <property type="term" value="P:developmental process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035556">
        <property type="term" value="P:intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd23076">
        <property type="entry name" value="PDZ_MAST4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05609">
        <property type="entry name" value="STKc_MAST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:001045">
        <property type="entry name" value="Microtubule-associated serine/threonine kinase 1a"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000012">
        <property type="entry name" value="microtubule-associated serine/threonine-protein kinase 2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1480.20:FF:000001">
        <property type="entry name" value="microtubule-associated serine/threonine-protein kinase 4 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000008">
        <property type="entry name" value="microtubule-associated serine/threonine-protein kinase 4 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.42.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1480.20">
        <property type="entry name" value="MAST3 pre-PK domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000961">
        <property type="entry name" value="AGC-kinase_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037711">
        <property type="entry name" value="MAST"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015022">
        <property type="entry name" value="MAST_pre-PK_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023142">
        <property type="entry name" value="MAST_pre-PK_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001478">
        <property type="entry name" value="PDZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041489">
        <property type="entry name" value="PDZ_6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036034">
        <property type="entry name" value="PDZ_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050236">
        <property type="entry name" value="Ser_Thr_kinase_AGC"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24356:SF224">
        <property type="entry name" value="MICROTUBULE-ASSOCIATED SERINE_THREONINE-PROTEIN KINASE 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24356">
        <property type="entry name" value="SERINE/THREONINE-PROTEIN KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08926">
        <property type="entry name" value="DUF1908"/>
        <property type="match status" value="1"/>
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    <dbReference type="Pfam" id="PF17820">
        <property type="entry name" value="PDZ_6"/>
        <property type="match status" value="1"/>
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    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00228">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF140482">
        <property type="entry name" value="MAST3 pre-PK domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50156">
        <property type="entry name" value="PDZ domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51285">
        <property type="entry name" value="AGC_KINASE_CTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50106">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11" id="KW-0418">Kinase</keyword>
    <keyword evidence="13" id="KW-0460">Magnesium</keyword>
    <keyword evidence="9" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="8" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Protein kinase" evidence="17">
        <location>
            <begin position="444"/>
            <end position="717"/>
        </location>
    </feature>
    <feature type="domain" description="AGC-kinase C-terminal" evidence="19">
        <location>
            <begin position="718"/>
            <end position="790"/>
        </location>
    </feature>
    <feature type="domain" description="PDZ" evidence="18">
        <location>
            <begin position="1013"/>
            <end position="1101"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1"/>
            <end position="39"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="92"/>
            <end position="188"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="417"/>
            <end position="437"/>
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    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="793"/>
            <end position="814"/>
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    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="829"/>
            <end position="954"/>
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    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
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    <feature type="region of interest" description="Disordered" evidence="16">
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    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
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    <feature type="region of interest" description="Disordered" evidence="16">
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    <feature type="region of interest" description="Disordered" evidence="16">
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    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1910"/>
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    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="2148"/>
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    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="2328"/>
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    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="2398"/>
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    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1"/>
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    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
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            <begin position="123"/>
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    </feature>
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        <location>
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    <feature type="compositionally biased region" description="Pro residues" evidence="16">
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        <location>
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    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
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        <source>
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    </evidence>
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        <source>
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        <source>
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        <source>
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    </evidence>
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        <source>
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        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
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        <source>
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        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
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        <source>
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        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
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    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00047899"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00048679"/>
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    <evidence type="ECO:0000256" key="16">
        <source>
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        <source>
            <dbReference type="PROSITE" id="PS50011"/>
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        <source>
            <dbReference type="PROSITE" id="PS50106"/>
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        <source>
            <dbReference type="PROSITE" id="PS51285"/>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015793.1"/>
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    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2483" mass="273924" checksum="2180F34564F43A97" modified="2019-05-08" version="1">MLFRKSSNPEVSSGPGKSFKFKRQLSEDGRPFRRGSLGGALTGRYLLPNTIAQQSWQPAETSNLVRMRSQALGQSAPSLTASLKELSLPRRGSFCRTSNRKSLIGNGQSPALPRPHSPLSAHAGSSPQDSPRNFSPSASAHFSFARSHGHRNDRTDGRRWSLASLPSSGYGTNTPSSTVSSSCSSQEKLHQLPYQPTPDELHFLSKHFCTTESIASDNRCRTTPMRPRSRSLSPGRSPACCDHEIIMMNHVYKERFPKATAQMEERLQDIITNHSPDNILPLADGVLSFTHHQIIELARDCLDKSHQGLITSRYFFELQHKLDKLLQEGQERSESGELAFIKQLVRKILIVIARPARLLECLEFDPEEFYYLLEAAEGHAKEGHVIKTDIPRYIISQLGLNKDPLEEIAQLANYDSGTAETPETDESSSNTSVTLRRKPRESDFETIKLISNGAYGAVYFVRHKETRQRFAMKKINKQNLILRNQIQQAFVERDILTFAENPFVVSMYCSFETRRHLCMVMEYVEGGDCATLMKNMGPLPVDMARMYFAETVLALEYLHNYGIVHRDLKPDNLLVTSMGHIKLTDFGLSKVGLMSMTTNLYEGHIEKDAREFLDKQVCGTPEYIAPEVILRQGYGKPVDWWAMGIILYEFLVGCVPFFGDTPEELFGQVISDEINWPEKDEAPPPDSQDLITLLLRQNPMERLGTGGAYEVKQHQFFRSLDWNSLLRQKAEFIPQLESEDDTSYFDTRSEKYHHMETEEEDDTNDEDFNLEIRQFSSCSHRFSKVFSSIDRITRNQGEEKEDTGDKTKNVTLPSVESLSWSSEYSELQQLSTSNSSDTESTRLRHSSGLLPKLAISSEGDQDETTCSEGSREEQEKAASNEEAAQDEPEVTTPASTISSSTLSVGSFSEHLDQINGRNESVDSTDNSSKPSSEPASHMARQRLESTEKKKISGKVTKSLSASALSLMIPGDVFGVSPLGSPMSPHSLSSDLSSSRDSSPSRDSSVASATPHQPILIHSSGKKYGFTIRAIRVYVGDSDIYTVHHIVWNVEEGSPAYQAGLKAGDLITHINGEPVHGLVHTEVIELLLKSGNKVSITTTPFENTSIKTGPARRNSYRSRMVRRSKKTKKKESLERRRSLFKKLAKQPSPLLHTSRSFSCLNRSLSSGESLPGSPTHSLSPRSPTPSYRSTPDFPSGTNSSQSSSPSSSGPNSPAGSGHIRPSTLHGLGPKLGGQRYRSGRRKSAGSIPLSPLARTPSPTPQPPSPQRSPSPLLGHSIGNSKIAQAFPSKMHSPPTIVRHIVRPKSAEPPRSPLLKRVQSEEKLAPSYGCDKKHLCSRKHSLEVTQEEVNRELSQREVTLQSLEENVCDVPSLTRVRPAEQGCLKRPVCRKLGRQESVDELDREKLKVKIVMKKQDLAEKQDCLQKASTIHEPSGESENPAILRLEERDKKSFQKALERSNHFETKITALETPSVGGILKDTLHKNANMRPNDCIPLDTHMPCHGPPLNELSHLSYDFKRISPPCTLQDVLPHSSDRLLNGKGESTDKNAPTKEFVKFERLDGKLANIDYLRKKMSFEEKDDSLCPVLKPKLTSNVHECLQLNTVRPINVQQDSAAVADSRAFISSAHAAQISSVSFVPLKALNGRVETGLEKSALISTESPVRKSPSEYKLEGRSVSCLKPIEGTLDIALLSGPQASKTELPSCLLPESQNTSNEVGSSMPLASQCNGVKEETSCQKEVVSPHSKSSKSNLLEPQFSQSSKSFQNAPATPVTAELETGKKISSPDAACNISVREGDQKREDSHSKQKDHATEAKLHTVHNQCLHTALASATPSTAQGTPEKTAGKESQTQKELPVKYPVTQRNYESFSVKAEGIRDSSVKKSIPDAFKTSDSKTIQRLSADFAIPLQTHRTTRVQTAGSKVESKDGKDLPKQFAKDDITPAGSSVERDMNKQEVMDSKSEMAFPKRPVQQSVESNTKSEKSLVCSSAQKDNSKDLRKKDQKHSSNIQLHIIERKLPNLITRQQFSPSCFPSVRETVNRNSVIDAHVHLGVQEQVKTGSAYMESSNNKSRQINETCSSKSKHSDKFTFAQKQCTANIKEEETVIQPAAGSRKDGKHANKSVQDSHSFVLISQRKLNNEYIQVEKPLGLEHGSAPSIQINSSTPETKPKSSCNGQAPAGSENSKHFLSQETAGFSETADQKQLYVNEKQNMPPKFSTVKDCLLLNKLADKDQSNQSITADKRPEGQKCTEALYVQNDSGKLEPSLFLNCDLGKVAEKAPIVSKGSPDSKGKGQVSQKQLTDANKQITAMLLSTATVQSSCHSAVTPLKQLNCPPSFPESRQEASPLSSAKVKTGSPELPDTSCKELKKQATSSTGDGKREITKSVSLLTLHSSAIPVEALHPASNLGSKPGNQEKPLSASNTVDVKGEDPIQAQPPAAKKQNVGKDFPRSVTTSDRPTALSSDKDSVRQRRGKDASRSSPHKKSSQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLI6</accession>
    <name>A0A452HLI6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Maleylacetoacetate isomerase</fullName>
            <ecNumber evidence="6">2.5.1.18</ecNumber>
            <ecNumber evidence="7">5.2.1.2</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="16">GSTZ1-1</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="15">Glutathione S-transferase zeta 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>4-maleylacetoacetate = 4-fumarylacetoacetate</text>
            <dbReference type="Rhea" id="RHEA:14817"/>
            <dbReference type="ChEBI" id="CHEBI:17105"/>
            <dbReference type="ChEBI" id="CHEBI:18034"/>
            <dbReference type="EC" id="5.2.1.2"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>RX + glutathione = an S-substituted glutathione + a halide anion + H(+)</text>
            <dbReference type="Rhea" id="RHEA:16437"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16042"/>
            <dbReference type="ChEBI" id="CHEBI:17792"/>
            <dbReference type="ChEBI" id="CHEBI:57925"/>
            <dbReference type="ChEBI" id="CHEBI:90779"/>
            <dbReference type="EC" id="2.5.1.18"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="2">
            <name>glutathione</name>
            <dbReference type="ChEBI" id="CHEBI:57925"/>
        </cofactor>
    </comment>
    <comment type="pathway">
        <text evidence="4">Amino-acid degradation; L-phenylalanine degradation; acetoacetate and fumarate from L-phenylalanine: step 5/6.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the GST superfamily. Zeta family.</text>
    </comment>
    <dbReference type="EC" id="2.5.1.18" evidence="6"/>
    <dbReference type="EC" id="5.2.1.2" evidence="7"/>
    <dbReference type="AlphaFoldDB" id="A0A452HLI6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018080.1">
        <property type="protein sequence ID" value="ENSGAGP00000015832.1"/>
        <property type="gene ID" value="ENSGAGG00000011906.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00139">
        <property type="reaction ID" value="UER00340"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004364">
        <property type="term" value="F:glutathione transferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016034">
        <property type="term" value="F:maleylacetoacetate isomerase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006749">
        <property type="term" value="P:glutathione metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006559">
        <property type="term" value="P:L-phenylalanine catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006572">
        <property type="term" value="P:L-tyrosine catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd03191">
        <property type="entry name" value="GST_C_Zeta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd03042">
        <property type="entry name" value="GST_N_Zeta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1050.10:FF:000010">
        <property type="entry name" value="Maleylacetoacetate isomerase isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.30.10:FF:000041">
        <property type="entry name" value="Maleylacetoacetate isomerase isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1050.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.30.10">
        <property type="entry name" value="Glutaredoxin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010987">
        <property type="entry name" value="Glutathione-S-Trfase_C-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036282">
        <property type="entry name" value="Glutathione-S-Trfase_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040079">
        <property type="entry name" value="Glutathione_S-Trfase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004045">
        <property type="entry name" value="Glutathione_S-Trfase_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004046">
        <property type="entry name" value="GST_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005955">
        <property type="entry name" value="GST_Zeta"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034330">
        <property type="entry name" value="GST_Zeta_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034333">
        <property type="entry name" value="GST_Zeta_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036249">
        <property type="entry name" value="Thioredoxin-like_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01262">
        <property type="entry name" value="maiA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR42673">
        <property type="entry name" value="MALEYLACETOACETATE ISOMERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR42673:SF4">
        <property type="entry name" value="MALEYLACETOACETATE ISOMERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14497">
        <property type="entry name" value="GST_C_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13417">
        <property type="entry name" value="GST_N_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDS00019">
        <property type="entry name" value="Glutathione_Transferase_(cytos"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDG00358">
        <property type="entry name" value="Main_(cytGST)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47616">
        <property type="entry name" value="GST C-terminal domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52833">
        <property type="entry name" value="Thioredoxin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50405">
        <property type="entry name" value="GST_CTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50404">
        <property type="entry name" value="GST_NTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="12" id="KW-0413">Isomerase</keyword>
    <keyword evidence="11" id="KW-0585">Phenylalanine catabolism</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0808">Transferase</keyword>
    <keyword evidence="10" id="KW-0828">Tyrosine catabolism</keyword>
    <feature type="domain" description="GST N-terminal" evidence="17">
        <location>
            <begin position="5"/>
            <end position="88"/>
        </location>
    </feature>
    <feature type="domain" description="GST C-terminal" evidence="18">
        <location>
            <begin position="93"/>
            <end position="211"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001622"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00001955"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004671"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00010007"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00012452"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00013199"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022878"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023232"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023235"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00047960"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00068155"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00080669"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00082080"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50404"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50405"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015832.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="215" mass="24256" checksum="A5293B0D55DFBD3F" modified="2019-05-08" version="1">MDSSGRPVLYSYFRSSCSWRVRIALALKGIAYDQEPVNLLKDGGQQFSTEFQAVNPMQQVPALKIDGIILSQSLAIIQYLEETRPNPRLLPQDLKKRAQARMISDHIVSGIQPLQNLTFLKQVGEKMEWAQHNITCGFKALEQILQHTAGRYCIGDEVSMADLCLVPQVHNAERFKVDLAPYPTITRINKALLELEAFQVSHPCRQPDTPAELRA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLJ1</accession>
    <name>A0A452HLJ1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Annexin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="6">Calcium/phospholipid-binding protein which promotes membrane fusion and is involved in exocytosis.</text>
    </comment>
    <comment type="domain">
        <text evidence="7">A pair of annexin repeats may form one binding site for calcium and phospholipid.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1 7">Belongs to the annexin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLJ1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018039.1">
        <property type="protein sequence ID" value="ENSGAGP00000015798.1"/>
        <property type="gene ID" value="ENSGAGG00000011880.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034704">
        <property type="term" value="C:calcium channel complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012506">
        <property type="term" value="C:vesicle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005262">
        <property type="term" value="F:calcium channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005544">
        <property type="term" value="F:calcium-dependent phospholipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001786">
        <property type="term" value="F:phosphatidylserine binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048513">
        <property type="term" value="P:animal organ development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055074">
        <property type="term" value="P:calcium ion homeostasis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009888">
        <property type="term" value="P:tissue development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.220.10:FF:000001">
        <property type="entry name" value="Annexin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.220.10:FF:000002">
        <property type="entry name" value="Annexin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.220.10:FF:000003">
        <property type="entry name" value="Annexin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.220.10:FF:000004">
        <property type="entry name" value="Annexin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.220.10">
        <property type="entry name" value="Annexin"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001464">
        <property type="entry name" value="Annexin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018502">
        <property type="entry name" value="Annexin_repeat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018252">
        <property type="entry name" value="Annexin_repeat_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037104">
        <property type="entry name" value="Annexin_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10502">
        <property type="entry name" value="ANNEXIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10502:SF239">
        <property type="entry name" value="ANNEXIN A7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00191">
        <property type="entry name" value="Annexin"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00196">
        <property type="entry name" value="ANNEXIN"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01871">
        <property type="entry name" value="ANNEXINVII"/>
    </dbReference>
    <dbReference type="SMART" id="SM00335">
        <property type="entry name" value="ANX"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47874">
        <property type="entry name" value="Annexin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00223">
        <property type="entry name" value="ANNEXIN_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51897">
        <property type="entry name" value="ANNEXIN_2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 7" id="KW-0041">Annexin</keyword>
    <keyword evidence="3 7" id="KW-0106">Calcium</keyword>
    <keyword evidence="5 7" id="KW-0111">Calcium/phospholipid-binding</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 7" id="KW-0677">Repeat</keyword>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="69"/>
            <end position="155"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="138"/>
            <end position="155"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007831"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023216"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023302"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00037210"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU003540"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015798.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="466" mass="49991" checksum="0AB1F296E1E960DB" modified="2019-05-08" version="1">MSYPGYPPSGYPAFPGYPPTGQESVYPPAGQYPYPSVPGGFPPVGGGAYPAAPPSGGFPGAAGYPAPGSYPGAPQPGGGYPGAPQPGGMPSYPGAPTGPGFGVPPNGSSFGGYPQPSSQSYVGGGPAQIPVGYPGGQVPSSVPSQPAAMTQSTQGTIRAAPNFDAGRDAEILRKAMKGFGTDEQAIINVVSNRSSDQRQKIKAAFKTMYGKDLIKDLKSELSGNMEELILALFMPPTYYDAWSLRHAMKGAGTQERVLIEILCTRTNQEIRDIVRTYKTEFGRDIEQDIRADTSGHFERLLISMCQGNRDENQSVNYQKAQEDAQRLYQAGEGKLGTDESCFNMILATRSFPQLKATVETYSKIANRDLLSSIGREFSGNVENGLKTIVQCALNRPAFFAERLYYSMKGAGTDDSTLVRIIVTRSEIDLVQIKQLFTQMYQKTLATMISSDTSGDYRKLLLAIVGQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLK1</accession>
    <name>A0A452HLK1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Serine/threonine-protein phosphatase</fullName>
            <ecNumber evidence="12">3.1.3.16</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="10 17">
            <text>O-phospho-L-seryl-[protein] + H2O = L-seryl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:20629"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="EC" id="3.1.3.16"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="11 17">
            <text>O-phospho-L-threonyl-[protein] + H2O = L-threonyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:47004"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="EC" id="3.1.3.16"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="2">
            <name>Fe(3+)</name>
            <dbReference type="ChEBI" id="CHEBI:29034"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the PPP phosphatase family. PP-2B subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.16" evidence="12"/>
    <dbReference type="AlphaFoldDB" id="A0A452HLK1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018054.1">
        <property type="protein sequence ID" value="ENSGAGP00000015809.1"/>
        <property type="gene ID" value="ENSGAGG00000011890.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033192">
        <property type="term" value="F:calmodulin-dependent protein phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097720">
        <property type="term" value="P:calcineurin-mediated signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd07416">
        <property type="entry name" value="MPP_PP2B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.60.21.10:FF:000002">
        <property type="entry name" value="Serine/threonine-protein phosphatase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.60.21.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004843">
        <property type="entry name" value="Calcineurin-like_PHP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029052">
        <property type="entry name" value="Metallo-depent_PP-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041751">
        <property type="entry name" value="MPP_PP2B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043360">
        <property type="entry name" value="PP2B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006186">
        <property type="entry name" value="Ser/Thr-sp_prot-phosphatase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45673">
        <property type="entry name" value="SERINE/THREONINE-PROTEIN PHOSPHATASE 2B CATALYTIC SUBUNIT 1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00149">
        <property type="entry name" value="Metallophos"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00114">
        <property type="entry name" value="STPHPHTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00156">
        <property type="entry name" value="PP2Ac"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56300">
        <property type="entry name" value="Metallo-dependent phosphatases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00125">
        <property type="entry name" value="SER_THR_PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="5 12" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="9" id="KW-0408">Iron</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="Serine/threonine specific protein phosphatases" evidence="14">
        <location>
            <begin position="150"/>
            <end position="155"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="1"/>
            <end position="21"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="479"/>
            <end position="508"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="13">
        <location>
            <begin position="492"/>
            <end position="508"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00001965"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00009905"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00047761"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00048336"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="RuleBase" id="RU004273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS00125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015809.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="508" mass="57346" checksum="2CFE3A0F06DE420C" modified="2019-05-08" version="1">MSVKEGSKGPPPAHGTSDRAVKAVPFPPTHRLTSEEVFDIDGKPRVDILKNHLVKEGRVDEEIALRIINEGAAILRREKTMIEVEAPITVCGDIHGQFFDLMKLFEVGGSPANTRYLFLGDYVDRGYFSIECVLYLWVLKILYPNTLFLLRGNHECRHLTEYFTFKQECKIKYSERVYDACMEAFDCLPLAALLNQQFLCVHGGLSPEINTLDDIKRLDRFKEPPAFGPMCDLLWSDPSEDFGNEKSQEHFSHNTVRGCSYFYSYPAVCEFLQNNNLLSIIRAHEAQDAGYRMYRKSQTTGFPSLITIFSAPNYLDVYNNKAAVLKYENNVMNIRQFNCSPHPYWLPNFMDVFTWSLPFVGEKVTEMLVNVLSICSDDELMTEGEDQFDGTAAARKEIIRNKIRAIGKMARVFSVLREESESVLTLKGLTPTGMLPSGVLAGGRQTLQSAIRGFSPQHKICSFEEAKGLDRINERMPPRKDALQQDGINSVNTTNANGNNGTGNNNVQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLK3</accession>
    <name>A0A452HLK3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">EGF-like calcium-binding domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HLK3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018053.1">
        <property type="protein sequence ID" value="ENSGAGP00000015808.1"/>
        <property type="gene ID" value="ENSGAGG00000011881.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007413">
        <property type="term" value="P:axonal fasciculation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027039">
        <property type="entry name" value="Crtac1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001881">
        <property type="entry name" value="EGF-like_Ca-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018097">
        <property type="entry name" value="EGF_Ca-bd_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013517">
        <property type="entry name" value="FG-GAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028994">
        <property type="entry name" value="Integrin_alpha_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049883">
        <property type="entry name" value="NOTCH1_EGF-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011519">
        <property type="entry name" value="UnbV_ASPIC"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16026">
        <property type="entry name" value="CARTILAGE ACIDIC PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16026:SF0">
        <property type="entry name" value="CARTILAGE ACIDIC PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07645">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13517">
        <property type="entry name" value="FG-GAP_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07593">
        <property type="entry name" value="UnbV_ASPIC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00179">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69318">
        <property type="entry name" value="Integrin alpha N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01187">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="1" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0732">Signal</keyword>
    <feature type="domain" description="EGF-like calcium-binding" evidence="4">
        <location>
            <begin position="446"/>
            <end position="493"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="SMART" id="SM00179"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015808.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="532" mass="57586" checksum="F47C46419B05F669" modified="2019-05-08" version="1">AGPCRAEAMWLSPWICLPDTPNYPACSCQRLATYTDKLFKLRAGRWEDLLSDEVNRAVASRFAGRSVACVDRAGSGRYAIYIANYANSNLGPHVLLEMDVAASDPARGIVVLTDVAAEAGVNKYTGGRGVAVGPIVSASASDIFCDNENGPNFLFHNQGDGSFVDVAASAGLDDPYQHGRGVALADFNRDGKVDIVYGNWNGPHRLYLQMNTSGRVHFRDIATPKFSMPSPVRTVLAADFDNNQELEVFFNNIAYRGSSANRLFRIIRREHLDPVIEELNPGDAMEPEGRGTGGAVTDFDGDGLLDLILSHGESMAQPLSVFRGTQGAGNNWLRVIPRTRFGAFARGARVVLFTRHSGAHLRIIDGGSGYLCEMEPVAHFGLGRDEASSLEVTWPDGRVVTRAVASGETNSVLEIPYPQGAEGQLPPTSLECGQGFSQHENGRCVDTDECAEFPFVCPREKPLCSNTYGGYRCHSSRRCSRGFEPSEDGTACVAQVAFFGGYPSAGSRLWAWRPPSSLLSLGLGLCLHFYAL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLK8</accession>
    <name>A0A452HLK8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Protein O-mannosyl-transferase 2</fullName>
            <ecNumber evidence="4">2.4.1.109</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="14">
            <text>a di-trans,poly-cis-dolichyl beta-D-mannosyl phosphate + L-seryl-[protein] = 3-O-(alpha-D-mannosyl)-L-seryl-[protein] + a di-trans,poly-cis-dolichyl phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17377"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:13546"/>
            <dbReference type="Rhea" id="RHEA-COMP:19498"/>
            <dbReference type="Rhea" id="RHEA-COMP:19501"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:57683"/>
            <dbReference type="ChEBI" id="CHEBI:58211"/>
            <dbReference type="ChEBI" id="CHEBI:137321"/>
            <dbReference type="EC" id="2.4.1.109"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>a di-trans,poly-cis-dolichyl beta-D-mannosyl phosphate + L-threonyl-[protein] = 3-O-(alpha-D-mannosyl)-L-threonyl-[protein] + a di-trans,poly-cis-dolichyl phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:53396"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:13547"/>
            <dbReference type="Rhea" id="RHEA-COMP:19498"/>
            <dbReference type="Rhea" id="RHEA-COMP:19501"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:57683"/>
            <dbReference type="ChEBI" id="CHEBI:58211"/>
            <dbReference type="ChEBI" id="CHEBI:137323"/>
            <dbReference type="EC" id="2.4.1.109"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="2">Protein modification; protein glycosylation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the glycosyltransferase 39 family.</text>
    </comment>
    <dbReference type="EC" id="2.4.1.109" evidence="4"/>
    <dbReference type="AlphaFoldDB" id="A0A452HLK8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018068.1">
        <property type="protein sequence ID" value="ENSGAGP00000015822.1"/>
        <property type="gene ID" value="ENSGAGG00000011816.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00378"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004169">
        <property type="term" value="F:dolichyl-phosphate-mannose-protein mannosyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="CDD" id="cd23282">
        <property type="entry name" value="beta-trefoil_MIR_POMT2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.80.10.50:FF:000026">
        <property type="entry name" value="Blast:Protein O-mannosyl-transferase 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.80.10.50">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003342">
        <property type="entry name" value="ArnT-like_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036300">
        <property type="entry name" value="MIR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016093">
        <property type="entry name" value="MIR_motif"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027005">
        <property type="entry name" value="PMT-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032421">
        <property type="entry name" value="PMT_4TMC"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10050">
        <property type="entry name" value="DOLICHYL-PHOSPHATE-MANNOSE--PROTEIN MANNOSYLTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10050:SF46">
        <property type="entry name" value="PROTEIN O-MANNOSYL-TRANSFERASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02815">
        <property type="entry name" value="MIR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02366">
        <property type="entry name" value="PMT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16192">
        <property type="entry name" value="PMT_4TMC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00472">
        <property type="entry name" value="MIR"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF82109">
        <property type="entry name" value="MIR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50919">
        <property type="entry name" value="MIR"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="5" id="KW-0328">Glycosyltransferase</keyword>
    <keyword evidence="11 15" id="KW-0472">Membrane</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0808">Transferase</keyword>
    <keyword evidence="7 15" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="10 15" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="51"/>
            <end position="72"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="79"/>
            <end position="101"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="107"/>
            <end position="123"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="135"/>
            <end position="168"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="188"/>
            <end position="214"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="506"/>
            <end position="529"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="545"/>
            <end position="564"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="576"/>
            <end position="594"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="606"/>
            <end position="624"/>
        </location>
    </feature>
    <feature type="domain" description="MIR" evidence="16">
        <location>
            <begin position="243"/>
            <end position="299"/>
        </location>
    </feature>
    <feature type="domain" description="MIR" evidence="16">
        <location>
            <begin position="313"/>
            <end position="369"/>
        </location>
    </feature>
    <feature type="domain" description="MIR" evidence="16">
        <location>
            <begin position="374"/>
            <end position="431"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004477"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004922"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007222"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00012839"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022676"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00039583"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00045085"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00045102"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50919"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015822.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="625" mass="70308" checksum="C9B4F1AD0C20EC2F" modified="2019-05-08" version="1">MGSYYINRTFFFDVHPPLGKMLIGLAGYLSGYNGTFPFQKPGDRYGHHNYIGMRGFCAFLGSCLVPFAYLIVLELSKSLSAALLAASILIFDTGCITLSQYILLDPILMFFLMGAVLSMVKYNSCAVRPFSAPWWFWLSLTGVNLAGAIGVKFVGLFVVLLVGLNTVYDLWELLGDLSLSLVTLGKHLLARVFCLILLPLVLYTAQFAVHFAVLNKSGPGDGFFSSAFQSRLIGNNLHNVSVPEHLAYGSVITMKNLRMAGGYLHSHWHLYPEGVGARQQQVTAYLHKDLNNLWVIKKHDSNTADHLDPSCSVEFVRHGDVIRLEHKETSRNLHSHQHEAPMTRKHFQVTGYGINGTGDANDFWRIEVVGRKSGKRIKVLRSQIRLTHLATGCILGSSGKTLPKWGWEQVEVTCTPYVKETPNALWNVEDHINPKLPNISLDVLKPSFPEILLESHMVMIRGNSGLKPKENEVTSKPWHWPINYQGLRFSGVNETDYRVYLLGNPVVWWLNLVTIGLYLLMATCTIVAIKRGAQLTVEFKELSEVLLHSGGKIMLGWLLHYLPFFMMGRVLYFHHYFPAMLFSSMLTGITWDTLLKFCAGCLSCSMGKRVYTYGMLALMLLIVYR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLL0</accession>
    <name>A0A452HLL0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">EGF-like calcium-binding domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HLL0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018060.1">
        <property type="protein sequence ID" value="ENSGAGP00000015815.1"/>
        <property type="gene ID" value="ENSGAGG00000011881.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007413">
        <property type="term" value="P:axonal fasciculation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00054">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027039">
        <property type="entry name" value="Crtac1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001881">
        <property type="entry name" value="EGF-like_Ca-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018097">
        <property type="entry name" value="EGF_Ca-bd_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013517">
        <property type="entry name" value="FG-GAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028994">
        <property type="entry name" value="Integrin_alpha_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049883">
        <property type="entry name" value="NOTCH1_EGF-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011519">
        <property type="entry name" value="UnbV_ASPIC"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16026">
        <property type="entry name" value="CARTILAGE ACIDIC PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16026:SF0">
        <property type="entry name" value="CARTILAGE ACIDIC PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07645">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13517">
        <property type="entry name" value="FG-GAP_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07593">
        <property type="entry name" value="UnbV_ASPIC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00179">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69318">
        <property type="entry name" value="Integrin alpha N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01187">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="1" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0732">Signal</keyword>
    <feature type="domain" description="EGF-like calcium-binding" evidence="4">
        <location>
            <begin position="411"/>
            <end position="458"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="SMART" id="SM00179"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015815.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="497" mass="53730" checksum="51091A563EF95E35" modified="2019-05-08" version="1">AWLFKLRAGRWEDLLSDEVNRAVASRFAGRSVACVDRAGSGRYAIYIANYANSNLGPHVLLEMDVAASDPARGIVVLTDVAAEAGVNKYTGGRGVAVGPIVSASASDIFCDNENGPNFLFHNQGDGSFVDVAASAGLDDPYQHGRGVALADFNRDGKVDIVYGNWNGPHRLYLQMNTSGRVHFRDIATPKFSMPSPVRTVLAADFDNNQELEVFFNNIAYRGSSANRLFRIIRREHLDPVIEELNPGDAMEPEGRGTGGAVTDFDGDGLLDLILSHGESMAQPLSVFRGTQGAGNNWLRVIPRTRFGAFARGARVVLFTRHSGAHLRIIDGGSGYLCEMEPVAHFGLGRDEASSLEVTWPDGRVVTRAVASGETNSVLEIPYPQGAEGQLPPTSLECGQGFSQHENGRCVDTDECAEFPFVCPREKPLCSNTYGGYRCHSSRRCSRGFEPSEDGTACVAQVAFFGGYPSAGSRLWAWRPPSSLLSLGLGLCLHFYAL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLL1</accession>
    <name>A0A452HLL1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2 10">Ubiquitin-like modifier-activating enzyme ATG7</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="10">Autophagy-related protein 7</fullName>
        </alternativeName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">E1-like activating enzyme involved in the 2 ubiquitin-like systems required for cytoplasm to vacuole transport (Cvt) and autophagy. Activates ATG12 for its conjugation with ATG5 as well as the ATG8 family proteins for their conjugation with phosphatidylethanolamine. Both systems are needed for the ATG8 association to Cvt vesicles and autophagosomes membranes. Required for autophagic death induced by caspase-8 inhibition. Required for mitophagy which contributes to regulate mitochondrial quantity and quality by eliminating the mitochondria to a basal level to fulfill cellular energy requirements and preventing excess ROS production. Modulates p53/TP53 activity to regulate cell cycle and survival during metabolic stress.</text>
    </comment>
    <comment type="subunit">
        <text evidence="10">Homodimer.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="10">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="10">Preautophagosomal structure</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="1 10">Belongs to the ATG7 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLL1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018065.1">
        <property type="protein sequence ID" value="ENSGAGP00000015819.1"/>
        <property type="gene ID" value="ENSGAGG00000011896.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000407">
        <property type="term" value="C:phagophore assembly site"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019778">
        <property type="term" value="F:Atg12 activating enzyme activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019779">
        <property type="term" value="F:Atg8 activating enzyme activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000045">
        <property type="term" value="P:autophagosome assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000422">
        <property type="term" value="P:autophagy of mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006995">
        <property type="term" value="P:cellular response to nitrogen starvation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034727">
        <property type="term" value="P:piecemeal microautophagy of the nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009896">
        <property type="term" value="P:positive regulation of catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032446">
        <property type="term" value="P:protein modification by small protein conjugation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042981">
        <property type="term" value="P:regulation of apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060255">
        <property type="term" value="P:regulation of macromolecule metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0080090">
        <property type="term" value="P:regulation of primary metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048511">
        <property type="term" value="P:rhythmic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd01486">
        <property type="entry name" value="Apg7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.140.100:FF:000001">
        <property type="entry name" value="Ubiquitin-like modifier-activating enzyme ATG7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.140.70:FF:000001">
        <property type="entry name" value="Ubiquitin-like modifier-activating enzyme atg7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.720:FF:000156">
        <property type="entry name" value="Ubiquitin-like modifier-activating enzyme ATG7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.720">
        <property type="entry name" value="NAD(P)-binding Rossmann-like Domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.140.100">
        <property type="entry name" value="Ubiquitin-like modifier-activating enzyme ATG7 C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.140.70">
        <property type="entry name" value="Ubiquitin-like modifier-activating enzyme ATG7 N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006285">
        <property type="entry name" value="Atg7"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032197">
        <property type="entry name" value="Atg7_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042522">
        <property type="entry name" value="Atg7_N_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042523">
        <property type="entry name" value="Atg7_N_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045886">
        <property type="entry name" value="ThiF/MoeB/HesA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000594">
        <property type="entry name" value="ThiF_NAD_FAD-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035985">
        <property type="entry name" value="Ubiquitin-activating_enz"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01381">
        <property type="entry name" value="E1_like_apg7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10953">
        <property type="entry name" value="UBIQUITIN-ACTIVATING ENZYME E1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10953:SF3">
        <property type="entry name" value="UBIQUITIN-LIKE MODIFIER-ACTIVATING ENZYME ATG7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16420">
        <property type="entry name" value="ATG7_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00899">
        <property type="entry name" value="ThiF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69572">
        <property type="entry name" value="Activating enzymes of the ubiquitin-like proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7 10" id="KW-0072">Autophagy</keyword>
    <keyword evidence="8" id="KW-0090">Biological rhythms</keyword>
    <keyword evidence="11" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4 10" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6 10" id="KW-0653">Protein transport</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 10" id="KW-0813">Transport</keyword>
    <keyword evidence="5 10" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="Ubiquitin-like modifier-activating enzyme Atg7 N-terminal" evidence="13">
        <location>
            <begin position="19"/>
            <end position="329"/>
        </location>
    </feature>
    <feature type="domain" description="THIF-type NAD/FAD binding fold" evidence="12">
        <location>
            <begin position="344"/>
            <end position="606"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="11">
        <location>
            <begin position="450"/>
            <end position="477"/>
        </location>
    </feature>
    <feature type="active site" description="Glycyl thioester intermediate" evidence="9">
        <location>
            <position position="578"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00010931"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00017647"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023006"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023108"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PIRSR" id="PIRSR606285-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="RuleBase" id="RU366022"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF00899"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="Pfam" id="PF16420"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015819.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="691" mass="77049" checksum="AA13C666E6115FF1" modified="2019-05-08" version="1">MATADSETHNPLEPRIPKLQFVPFTSALDAGFWHELTQKKLNEYRLDETPKGIKGYYYNGDPFGLPARLTLEFSAFDMNAATPAHCCPAVGTLYNTNTFESFKSCDKKSLLEKAANEIWETIKSGTALENPMLLNKFLLLTFADLKKYRFYYWFCYPALCFPDGIPIVQQAVCLGERFSPTQIQALQKAYDELCQREGVTALPYFLIKYNDKSVLVSLLKNWNDFFQDQEEKVTVGVYDPCNFTQYPGWPLRNLLVMAAHRWGSHLHSVEVLCFRDRTMQGVRDIAHSSIFEIKLPQMPQSPDCPKAVGWEKNQKGGMGPRMVNLSECMDPKRLAESSVDLNLKLMCWRLVPTLDLEKVVSSKCLLLGAGTLGCCVARTLMGWGVRQITFVDNAKISYSNPVRQPLYEFEDCLGGGKPKALAAADRLQKIFPGVNAEGFNMSIPMPGHPVNFSEVTMEQAQKDVAQLERLVDAHDIIFLLMDTRESRWLPAVIAASKRKLVINAALGFDTFVVMRHGLKKPKQQESGDSCSSNASGSADLLGSSLFSNIPGYKLGCYFCNDVVAPGDSTRDRTLDQQCTVSRPGLAMIAGALAVELMASVLQHPEGGYAVASSSDDRMNEPPTSLGLVPHQIRGFLSRFDNVLPVSLAFDKCTACSPKEGWETDKEEGQQRTNTSSLLKTSICYFEHSLRR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLL2</accession>
    <name>A0A452HLL2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Protein O-mannosyl-transferase 2</fullName>
            <ecNumber evidence="4">2.4.1.109</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="14">
            <text>a di-trans,poly-cis-dolichyl beta-D-mannosyl phosphate + L-seryl-[protein] = 3-O-(alpha-D-mannosyl)-L-seryl-[protein] + a di-trans,poly-cis-dolichyl phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17377"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:13546"/>
            <dbReference type="Rhea" id="RHEA-COMP:19498"/>
            <dbReference type="Rhea" id="RHEA-COMP:19501"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:57683"/>
            <dbReference type="ChEBI" id="CHEBI:58211"/>
            <dbReference type="ChEBI" id="CHEBI:137321"/>
            <dbReference type="EC" id="2.4.1.109"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>a di-trans,poly-cis-dolichyl beta-D-mannosyl phosphate + L-threonyl-[protein] = 3-O-(alpha-D-mannosyl)-L-threonyl-[protein] + a di-trans,poly-cis-dolichyl phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:53396"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:13547"/>
            <dbReference type="Rhea" id="RHEA-COMP:19498"/>
            <dbReference type="Rhea" id="RHEA-COMP:19501"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:57683"/>
            <dbReference type="ChEBI" id="CHEBI:58211"/>
            <dbReference type="ChEBI" id="CHEBI:137323"/>
            <dbReference type="EC" id="2.4.1.109"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="2">Protein modification; protein glycosylation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the glycosyltransferase 39 family.</text>
    </comment>
    <dbReference type="EC" id="2.4.1.109" evidence="4"/>
    <dbReference type="AlphaFoldDB" id="A0A452HLL2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018064.1">
        <property type="protein sequence ID" value="ENSGAGP00000015818.1"/>
        <property type="gene ID" value="ENSGAGG00000011816.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00378"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004169">
        <property type="term" value="F:dolichyl-phosphate-mannose-protein mannosyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="CDD" id="cd23282">
        <property type="entry name" value="beta-trefoil_MIR_POMT2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.80.10.50:FF:000026">
        <property type="entry name" value="Blast:Protein O-mannosyl-transferase 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.80.10.50">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003342">
        <property type="entry name" value="ArnT-like_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036300">
        <property type="entry name" value="MIR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016093">
        <property type="entry name" value="MIR_motif"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027005">
        <property type="entry name" value="PMT-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032421">
        <property type="entry name" value="PMT_4TMC"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10050">
        <property type="entry name" value="DOLICHYL-PHOSPHATE-MANNOSE--PROTEIN MANNOSYLTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10050:SF46">
        <property type="entry name" value="PROTEIN O-MANNOSYL-TRANSFERASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02815">
        <property type="entry name" value="MIR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02366">
        <property type="entry name" value="PMT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16192">
        <property type="entry name" value="PMT_4TMC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00472">
        <property type="entry name" value="MIR"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF82109">
        <property type="entry name" value="MIR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50919">
        <property type="entry name" value="MIR"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="5" id="KW-0328">Glycosyltransferase</keyword>
    <keyword evidence="11 16" id="KW-0472">Membrane</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0808">Transferase</keyword>
    <keyword evidence="7 16" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="10 16" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="175"/>
            <end position="196"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="202"/>
            <end position="220"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="227"/>
            <end position="246"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="258"/>
            <end position="291"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="312"/>
            <end position="333"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="622"/>
            <end position="645"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="666"/>
            <end position="686"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="692"/>
            <end position="714"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="726"/>
            <end position="746"/>
        </location>
    </feature>
    <feature type="domain" description="MIR" evidence="17">
        <location>
            <begin position="429"/>
            <end position="485"/>
        </location>
    </feature>
    <feature type="domain" description="MIR" evidence="17">
        <location>
            <begin position="490"/>
            <end position="547"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="52"/>
            <end position="80"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004477"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004922"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007222"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00012839"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022676"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00039583"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00045085"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00045102"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50919"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015818.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="775" mass="86614" checksum="195692C10E47B8CE" modified="2019-05-08" version="1">MPCEGTRLRFPPCPTARTPFPLMPCAVVPGCRRLPRVGAQMTLAVGCCQAESGPRCGGLRQRRPRRPAGEPSRPSPPPRAWAPLEAPVGVWAPLASITLLSFASRFYRLSEPPHVCWDETHFGKMGSYYINRTFFFDVHPPLGKMLIGLAGYLSGYNGTFPFQKPGDRYGHHNYIGMRGFCAFLGSCLVPFAYLIVLELSKSLSAALLAASILIFDTGCITLSQYILLDPILMFFLMGAVLSMVKYNSCAVRPFSAPWWFWLSLTGVNLAGAIGVKFVGLFVVLLVGLNTVYDLWELLGDLSLSLVTLGKHLLARVFCLILLPLVLYTAQFAVHFAVLNKSGPGDGFFSSAFQSRLIGNNLHNVSVPEHLAYGSVITMKNLRMAGGYLHSHWHLYPEGVGARQQQVSVSLYHSLSLLPASDHLDPSCSVEFVRHGDVIRLEHKETSRNLHSHQHEAPMTRKHFQVTGYGINGTGDANDFWRIEVVGRKSGKRIKVLRSQIRLTHLATGCILGSSGKTLPKWGWEQVEVTCTPYVKETPNALWNVEDHINPKLPNISLDVLKPSFPEILLESHMVMIRGNSGLKPKENEVTSKPWHWPINYQGLRFSGVNETDYRVYLLGNPVVWWLNLVTIGLYLLMATCTIVAIKRGAQLTVEFKELSEVLLHSGGKIMLGWLLHYLPFFMMGRVLYFHHYFPAMLFSSMLTGITWDTLLKFCAGCLSCSMGKRVYTYGMLALMLLIVYSFYLFHPLSYGMVGPMASDPNSPMAGLRWLESWEF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLL4</accession>
    <name>A0A452HLL4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">N-alpha-acetyltransferase 40</fullName>
            <ecNumber evidence="4">2.3.1.257</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="14">N-acetyltransferase 11</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="15">N-alpha-acetyltransferase D</fullName>
        </alternativeName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="12">
            <text>N-terminal L-seryl-[histone H2A] + acetyl-CoA = N-terminal N(alpha)-acetyl-L-seryl-[histone H2A] + CoA + H(+)</text>
            <dbReference type="Rhea" id="RHEA:50600"/>
            <dbReference type="Rhea" id="RHEA-COMP:12742"/>
            <dbReference type="Rhea" id="RHEA-COMP:12744"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57288"/>
            <dbReference type="ChEBI" id="CHEBI:64738"/>
            <dbReference type="ChEBI" id="CHEBI:83690"/>
            <dbReference type="EC" id="2.3.1.257"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>N-terminal L-seryl-[histone H4] + acetyl-CoA = N-terminal N(alpha)-acetyl-L-seryl-[histone H4] + CoA + H(+)</text>
            <dbReference type="Rhea" id="RHEA:50596"/>
            <dbReference type="Rhea" id="RHEA-COMP:12740"/>
            <dbReference type="Rhea" id="RHEA-COMP:12743"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57288"/>
            <dbReference type="ChEBI" id="CHEBI:64738"/>
            <dbReference type="ChEBI" id="CHEBI:83690"/>
            <dbReference type="EC" id="2.3.1.257"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the acetyltransferase family. NAA40 subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.3.1.257" evidence="4"/>
    <dbReference type="AlphaFoldDB" id="A0A452HLL4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018141.1">
        <property type="protein sequence ID" value="ENSGAGP00000015876.1"/>
        <property type="gene ID" value="ENSGAGG00000011948.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043998">
        <property type="term" value="F:histone H2A acetyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010485">
        <property type="term" value="F:histone H4 acetyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:1990189">
        <property type="term" value="F:protein N-terminal-serine acetyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.630.30:FF:000033">
        <property type="entry name" value="N-alpha-acetyltransferase 40 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.630.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016181">
        <property type="entry name" value="Acyl_CoA_acyltransferase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000182">
        <property type="entry name" value="GNAT_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039949">
        <property type="entry name" value="NAA40"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR20531">
        <property type="entry name" value="N-ALPHA-ACETYLTRANSFERASE 40"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR20531:SF1">
        <property type="entry name" value="N-ALPHA-ACETYLTRANSFERASE 40"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00583">
        <property type="entry name" value="Acetyltransf_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55729">
        <property type="entry name" value="Acyl-CoA N-acyltransferases (Nat)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51186">
        <property type="entry name" value="GNAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="11" id="KW-0012">Acyltransferase</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="10" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="8" id="KW-0519">Myristate</keyword>
    <keyword evidence="9" id="KW-0539">Nucleus</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="N-acetyltransferase" evidence="16">
        <location>
            <begin position="42"/>
            <end position="195"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00008870"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00012950"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00015043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022707"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023315"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00047821"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00049524"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00079213"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00082154"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS51186"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015876.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="216" mass="24847" checksum="70A254174ECB94F5" modified="2019-05-08" version="1">MDAVCAKVEAANKLGDPLEAFPVFKKYDRNGLNVSIECKRVSGMNQATVDWAFELTKTNMQTLYEQSEWGWKDREKREEMTDDRAWYLIAWEDSSIPVAFSHFRFDVECGDEVLYCYEVQLESKVRRKGLGKFLIQILQLMANSTQMKKVMLTVFKHNHGAYQFFREALQFEIDDNSPSMSGCCGDDCSYEILSRRTKFGESQHSHSGSHCGGCCH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLL6</accession>
    <name>A0A452HLL6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2 10">Ubiquitin-like modifier-activating enzyme ATG7</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="10">Autophagy-related protein 7</fullName>
        </alternativeName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">E1-like activating enzyme involved in the 2 ubiquitin-like systems required for cytoplasm to vacuole transport (Cvt) and autophagy. Activates ATG12 for its conjugation with ATG5 as well as the ATG8 family proteins for their conjugation with phosphatidylethanolamine. Both systems are needed for the ATG8 association to Cvt vesicles and autophagosomes membranes. Required for autophagic death induced by caspase-8 inhibition. Required for mitophagy which contributes to regulate mitochondrial quantity and quality by eliminating the mitochondria to a basal level to fulfill cellular energy requirements and preventing excess ROS production. Modulates p53/TP53 activity to regulate cell cycle and survival during metabolic stress.</text>
    </comment>
    <comment type="subunit">
        <text evidence="10">Homodimer.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="10">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="10">Preautophagosomal structure</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="1 10">Belongs to the ATG7 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLL6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018073.1">
        <property type="protein sequence ID" value="ENSGAGP00000015826.1"/>
        <property type="gene ID" value="ENSGAGG00000011896.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000407">
        <property type="term" value="C:phagophore assembly site"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019778">
        <property type="term" value="F:Atg12 activating enzyme activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019779">
        <property type="term" value="F:Atg8 activating enzyme activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000045">
        <property type="term" value="P:autophagosome assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000422">
        <property type="term" value="P:autophagy of mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006995">
        <property type="term" value="P:cellular response to nitrogen starvation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034727">
        <property type="term" value="P:piecemeal microautophagy of the nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009896">
        <property type="term" value="P:positive regulation of catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032446">
        <property type="term" value="P:protein modification by small protein conjugation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042981">
        <property type="term" value="P:regulation of apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060255">
        <property type="term" value="P:regulation of macromolecule metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0080090">
        <property type="term" value="P:regulation of primary metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048511">
        <property type="term" value="P:rhythmic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd01486">
        <property type="entry name" value="Apg7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.140.100:FF:000001">
        <property type="entry name" value="Ubiquitin-like modifier-activating enzyme ATG7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.140.70:FF:000001">
        <property type="entry name" value="Ubiquitin-like modifier-activating enzyme atg7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.720:FF:000156">
        <property type="entry name" value="Ubiquitin-like modifier-activating enzyme ATG7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.720">
        <property type="entry name" value="NAD(P)-binding Rossmann-like Domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.140.100">
        <property type="entry name" value="Ubiquitin-like modifier-activating enzyme ATG7 C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.140.70">
        <property type="entry name" value="Ubiquitin-like modifier-activating enzyme ATG7 N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006285">
        <property type="entry name" value="Atg7"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032197">
        <property type="entry name" value="Atg7_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042522">
        <property type="entry name" value="Atg7_N_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042523">
        <property type="entry name" value="Atg7_N_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045886">
        <property type="entry name" value="ThiF/MoeB/HesA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000594">
        <property type="entry name" value="ThiF_NAD_FAD-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035985">
        <property type="entry name" value="Ubiquitin-activating_enz"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01381">
        <property type="entry name" value="E1_like_apg7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10953">
        <property type="entry name" value="UBIQUITIN-ACTIVATING ENZYME E1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10953:SF3">
        <property type="entry name" value="UBIQUITIN-LIKE MODIFIER-ACTIVATING ENZYME ATG7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16420">
        <property type="entry name" value="ATG7_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00899">
        <property type="entry name" value="ThiF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69572">
        <property type="entry name" value="Activating enzymes of the ubiquitin-like proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7 10" id="KW-0072">Autophagy</keyword>
    <keyword evidence="8" id="KW-0090">Biological rhythms</keyword>
    <keyword evidence="11" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4 10" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6 10" id="KW-0653">Protein transport</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 10" id="KW-0813">Transport</keyword>
    <keyword evidence="5 10" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="Ubiquitin-like modifier-activating enzyme Atg7 N-terminal" evidence="13">
        <location>
            <begin position="19"/>
            <end position="329"/>
        </location>
    </feature>
    <feature type="domain" description="THIF-type NAD/FAD binding fold" evidence="12">
        <location>
            <begin position="344"/>
            <end position="606"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="11">
        <location>
            <begin position="450"/>
            <end position="477"/>
        </location>
    </feature>
    <feature type="active site" description="Glycyl thioester intermediate" evidence="9">
        <location>
            <position position="578"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00010931"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00017647"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023006"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023108"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PIRSR" id="PIRSR606285-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="RuleBase" id="RU366022"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF00899"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="Pfam" id="PF16420"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015826.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="690" mass="76730" checksum="446C753394110AFC" modified="2019-05-08" version="1">MATADSETHNPLEPRIPKLQFVPFTSALDAGFWHELTQKKLNEYRLDETPKGIKGYYYNGDPFGLPARLTLEFSAFDMNAATPAHCCPAVGTLYNTNTFESFKSCDKKSLLEKAANEIWETIKSGTALENPMLLNKFLLLTFADLKKYRFYYWFCYPALCFPDGIPIVQQAVCLGERFSPTQIQALQKAYDELCQREGVTALPYFLIKYNDKSVLVSLLKNWNDFFQDQEEKVTVGVYDPCNFTQYPGWPLRNLLVMAAHRWGSHLHSVEVLCFRDRTMQGVRDIAHSSIFEIKLPQMPQSPDCPKAVGWEKNQKGGMGPRMVNLSECMDPKRLAESSVDLNLKLMCWRLVPTLDLEKVVSSKCLLLGAGTLGCCVARTLMGWGVRQITFVDNAKISYSNPVRQPLYEFEDCLGGGKPKALAAADRLQKIFPGVNAEGFNMSIPMPGHPVNFSEVTMEQAQKDVAQLERLVDAHDIIFLLMDTRESRWLPAVIAASKRKLVINAALGFDTFVVMRHGLKKPKQQESGDSCSSNASGSADLLGSSLFSNIPGYKLGCYFCNDVVAPGDSTRDRTLDQQCTVSRPGLAMIAGALAVELMASVLQHPEGGYAVASSSDDRMNEPPTSLGLVPHQIRGFLSRFDNVLPVSLAFDKCTACSPKDLGSNPILIPMFWLQIKSFWSASPLPTLFSGF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLL7</accession>
    <name>A0A452HLL7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2 10">Ubiquitin-like modifier-activating enzyme ATG7</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="10">Autophagy-related protein 7</fullName>
        </alternativeName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">E1-like activating enzyme involved in the 2 ubiquitin-like systems required for cytoplasm to vacuole transport (Cvt) and autophagy. Activates ATG12 for its conjugation with ATG5 as well as the ATG8 family proteins for their conjugation with phosphatidylethanolamine. Both systems are needed for the ATG8 association to Cvt vesicles and autophagosomes membranes. Required for autophagic death induced by caspase-8 inhibition. Required for mitophagy which contributes to regulate mitochondrial quantity and quality by eliminating the mitochondria to a basal level to fulfill cellular energy requirements and preventing excess ROS production. Modulates p53/TP53 activity to regulate cell cycle and survival during metabolic stress.</text>
    </comment>
    <comment type="subunit">
        <text evidence="10">Homodimer.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="10">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="10">Preautophagosomal structure</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="1 10">Belongs to the ATG7 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLL7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018071.1">
        <property type="protein sequence ID" value="ENSGAGP00000015824.1"/>
        <property type="gene ID" value="ENSGAGG00000011896.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000407">
        <property type="term" value="C:phagophore assembly site"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019778">
        <property type="term" value="F:Atg12 activating enzyme activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019779">
        <property type="term" value="F:Atg8 activating enzyme activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000045">
        <property type="term" value="P:autophagosome assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000422">
        <property type="term" value="P:autophagy of mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006995">
        <property type="term" value="P:cellular response to nitrogen starvation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034727">
        <property type="term" value="P:piecemeal microautophagy of the nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009896">
        <property type="term" value="P:positive regulation of catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032446">
        <property type="term" value="P:protein modification by small protein conjugation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042981">
        <property type="term" value="P:regulation of apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060255">
        <property type="term" value="P:regulation of macromolecule metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0080090">
        <property type="term" value="P:regulation of primary metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048511">
        <property type="term" value="P:rhythmic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd01486">
        <property type="entry name" value="Apg7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.140.100:FF:000001">
        <property type="entry name" value="Ubiquitin-like modifier-activating enzyme ATG7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.140.70:FF:000001">
        <property type="entry name" value="Ubiquitin-like modifier-activating enzyme atg7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.720:FF:000156">
        <property type="entry name" value="Ubiquitin-like modifier-activating enzyme ATG7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.720">
        <property type="entry name" value="NAD(P)-binding Rossmann-like Domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.140.100">
        <property type="entry name" value="Ubiquitin-like modifier-activating enzyme ATG7 C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.140.70">
        <property type="entry name" value="Ubiquitin-like modifier-activating enzyme ATG7 N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006285">
        <property type="entry name" value="Atg7"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032197">
        <property type="entry name" value="Atg7_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042522">
        <property type="entry name" value="Atg7_N_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042523">
        <property type="entry name" value="Atg7_N_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045886">
        <property type="entry name" value="ThiF/MoeB/HesA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000594">
        <property type="entry name" value="ThiF_NAD_FAD-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035985">
        <property type="entry name" value="Ubiquitin-activating_enz"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01381">
        <property type="entry name" value="E1_like_apg7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10953">
        <property type="entry name" value="UBIQUITIN-ACTIVATING ENZYME E1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10953:SF3">
        <property type="entry name" value="UBIQUITIN-LIKE MODIFIER-ACTIVATING ENZYME ATG7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16420">
        <property type="entry name" value="ATG7_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00899">
        <property type="entry name" value="ThiF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69572">
        <property type="entry name" value="Activating enzymes of the ubiquitin-like proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7 10" id="KW-0072">Autophagy</keyword>
    <keyword evidence="8" id="KW-0090">Biological rhythms</keyword>
    <keyword evidence="11" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4 10" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6 10" id="KW-0653">Protein transport</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 10" id="KW-0813">Transport</keyword>
    <keyword evidence="5 10" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="Ubiquitin-like modifier-activating enzyme Atg7 N-terminal" evidence="13">
        <location>
            <begin position="19"/>
            <end position="329"/>
        </location>
    </feature>
    <feature type="domain" description="THIF-type NAD/FAD binding fold" evidence="12">
        <location>
            <begin position="344"/>
            <end position="606"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="11">
        <location>
            <begin position="450"/>
            <end position="477"/>
        </location>
    </feature>
    <feature type="active site" description="Glycyl thioester intermediate" evidence="9">
        <location>
            <position position="578"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00010931"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00017647"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023006"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023108"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PIRSR" id="PIRSR606285-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="RuleBase" id="RU366022"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF00899"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="Pfam" id="PF16420"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015824.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="666" mass="73988" checksum="6B5BA5E95EDCCCFE" modified="2019-05-08" version="1">MATADSETHNPLEPRIPKLQFVPFTSALDAGFWHELTQKKLNEYRLDETPKGIKGYYYNGDPFGLPARLTLEFSAFDMNAATPAHCCPAVGTLYNTNTFESFKSCDKKSLLEKAANEIWETIKSGTALENPMLLNKFLLLTFADLKKYRFYYWFCYPALCFPDGIPIVQQAVCLGERFSPTQIQALQKAYDELCQREGVTALPYFLIKYNDKSVLVSLLKNWNDFFQDQEEKVTVGVYDPCNFTQYPGWPLRNLLVMAAHRWGSHLHSVEVLCFRDRTMQGVRDIAHSSIFEIKLPQMPQSPDCPKAVGWEKNQKGGMGPRMVNLSECMDPKRLAESSVDLNLKLMCWRLVPTLDLEKVVSSKCLLLGAGTLGCCVARTLMGWGVRQITFVDNAKISYSNPVRQPLYEFEDCLGGGKPKALAAADRLQKIFPGVNAEGFNMSIPMPGHPVNFSEVTMEQAQKDVAQLERLVDAHDIIFLLMDTRESRWLPAVIAASKRKLVINAALGFDTFVVMRHGLKKPKQQESGDSCSSNASGSADLLGSSLFSNIPGYKLGCYFCNDVVAPGDSTRDRTLDQQCTVSRPGLAMIAGALAVELMASVLQHPEGGYAVASSSDDRMNEPPTSLGLVPHQIRGFLSRFDNVLPVSLAFDKCTACSPKSPKVRIGI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLL8</accession>
    <name>A0A452HLL8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">EGF-like calcium-binding domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HLL8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018084.1">
        <property type="protein sequence ID" value="ENSGAGP00000015833.1"/>
        <property type="gene ID" value="ENSGAGG00000011881.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007413">
        <property type="term" value="P:axonal fasciculation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00054">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027039">
        <property type="entry name" value="Crtac1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001881">
        <property type="entry name" value="EGF-like_Ca-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018097">
        <property type="entry name" value="EGF_Ca-bd_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013517">
        <property type="entry name" value="FG-GAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028994">
        <property type="entry name" value="Integrin_alpha_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049883">
        <property type="entry name" value="NOTCH1_EGF-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011519">
        <property type="entry name" value="UnbV_ASPIC"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16026">
        <property type="entry name" value="CARTILAGE ACIDIC PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16026:SF0">
        <property type="entry name" value="CARTILAGE ACIDIC PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07645">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13517">
        <property type="entry name" value="FG-GAP_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07593">
        <property type="entry name" value="UnbV_ASPIC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00179">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69318">
        <property type="entry name" value="Integrin alpha N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01187">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="1" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0732">Signal</keyword>
    <feature type="domain" description="EGF-like calcium-binding" evidence="4">
        <location>
            <begin position="397"/>
            <end position="444"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="SMART" id="SM00179"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015833.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="483" mass="52005" checksum="E484F0E29F762012" modified="2025-10-08" version="2">MSDEVNRAVASRFAGRSVACVDRAGSGRYAIYIANYANSNLGPHVLLEMDVAASDPARGIVVLTDVAAEAGVNKYTGGRGVAVGPIVSASASDIFCDNENGPNFLFHNQGDGSFVDVAASAGLDDPYQHGRGVALADFNRDGKVDIVYGNWNGPHRLYLQMNTSGRVHFRDIATPKFSMPSPVRTVLAADFDNNQELEVFFNNIAYRGSSANRLFRIIRREHLDPVIEELNPGDAMEPEGRGTGGAVTDFDGDGLLDLILSHGESMAQPLSVFRGTQGAGNNWLRVIPRTRFGAFARGARVVLFTRHSGAHLRIIDGGSGYLCEMEPVAHFGLGRDEASSLEVTWPDGRVVTRAVASGETNSVLEIPYPQGAEGQLPPTSLECGQGFSQHENGRCVDTDECAEFPFVCPREKPLCSNTYGGYRCHSSRRCSRGFEPSEDGTACVAQVAFFGGYPSAGSRLWAWRPPSSLLSLGLGLCLHFYAL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLL9</accession>
    <name>A0A452HLL9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">G-protein coupled receptors family 1 profile domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="10">Belongs to the G-protein coupled receptor 1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLL9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018069.1">
        <property type="protein sequence ID" value="ENSGAGP00000015823.1"/>
        <property type="gene ID" value="ENSGAGG00000011898.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045202">
        <property type="term" value="C:synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="GOC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001591">
        <property type="term" value="F:dopamine neurotransmitter receptor activity, coupled via Gi/Go"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004930">
        <property type="term" value="F:G protein-coupled receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007195">
        <property type="term" value="P:adenylate cyclase-inhibiting dopamine receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051481">
        <property type="term" value="P:negative regulation of cytosolic calcium ion concentration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051967">
        <property type="term" value="P:negative regulation of synaptic transmission, glutamatergic"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060158">
        <property type="term" value="P:phospholipase C-activating dopamine receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0014059">
        <property type="term" value="P:regulation of dopamine secretion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043266">
        <property type="term" value="P:regulation of potassium ion transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1070.10">
        <property type="entry name" value="Rhodopsin 7-helix transmembrane proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002185">
        <property type="entry name" value="Dopamine_D4_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000929">
        <property type="entry name" value="Dopamine_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000276">
        <property type="entry name" value="GPCR_Rhodpsn"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017452">
        <property type="entry name" value="GPCR_Rhodpsn_7TM"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24248">
        <property type="entry name" value="ADRENERGIC RECEPTOR-RELATED G-PROTEIN COUPLED RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24248:SF143">
        <property type="entry name" value="D(4) DOPAMINE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00001">
        <property type="entry name" value="7tm_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00569">
        <property type="entry name" value="DOPAMINED4R"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00242">
        <property type="entry name" value="DOPAMINER"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00237">
        <property type="entry name" value="GPCRRHODOPSN"/>
    </dbReference>
    <dbReference type="SMART" id="SM01381">
        <property type="entry name" value="7TM_GPCR_Srsx"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81321">
        <property type="entry name" value="Family A G protein-coupled receptor-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00237">
        <property type="entry name" value="G_PROTEIN_RECEP_F1_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50262">
        <property type="entry name" value="G_PROTEIN_RECEP_F1_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="7" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5 10" id="KW-0297">G-protein coupled receptor</keyword>
    <keyword evidence="6 11" id="KW-0472">Membrane</keyword>
    <keyword evidence="8 10" id="KW-0675">Receptor</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9 10" id="KW-0807">Transducer</keyword>
    <keyword evidence="3 10 11" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 11" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="42"/>
            <end position="61"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="81"/>
            <end position="108"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="201"/>
            <end position="222"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="234"/>
            <end position="257"/>
        </location>
    </feature>
    <feature type="domain" description="G-protein coupled receptors family 1 profile" evidence="12">
        <location>
            <begin position="1"/>
            <end position="254"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023224"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="RuleBase" id="RU000688"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50262"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015823.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="273" mass="31117" checksum="4F21DB3D7DAFF82A" modified="2019-05-08" version="1">MTMDVMLCTASIFNLCAISVDRFIAVSIPLNYNRQQIDLRQLILISTTWIFAFAVASPVIFGLNNVPDRDPTLCQLEDNNYIVYSSICSFFIPCPVMLVLYCAMFQGLKRWEEARKAKLKSSVYAGGQKLCCPPPLLERKQAEIKLQECSLYDRSECSFPRGYMMNSGIQTVSCPHLKYSHPAHKQKQAKINGRERKAMKVLPVVVGAFLCCWTPFFVVHITRALCKSCTIPSLVTSIVTWLGYVNSALNPIIYTIFNTEFRKFFRKALHFFC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP01</accession>
    <name>A0A452IP01_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">UDP-glucose glycoprotein glucosyltransferase 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Recognizes glycoproteins with minor folding defects. Reglucosylates single N-glycans near the misfolded part of the protein, thus providing quality control for protein folding in the endoplasmic reticulum. Reglucosylated proteins are recognized by calreticulin for recycling to the endoplasmic reticulum and refolding or degradation.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="10">
            <text>N(4)-(alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;6)]-alpha-D-Man-(1-&gt;6)]-beta-D-Man-(1-&gt;4)-beta-D-GlcNAc-(1-&gt;4)-beta-D-GlcNAc)-L-asparaginyl-[protein] (N-glucan mannose isomer 9A1,2,3B1,2,3) + UDP-alpha-D-glucose = N(4)-(alpha-D-Glc-(1-&gt;3)-alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;6)]-alpha-D-Man-(1-&gt;6)]-beta-D-Man-(1-&gt;4)-beta-D-GlcNAc-(1-&gt;4)-beta-D-GlcNAc)-L-asparaginyl-[protein] + UDP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:61304"/>
            <dbReference type="Rhea" id="RHEA-COMP:14356"/>
            <dbReference type="Rhea" id="RHEA-COMP:14357"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:58223"/>
            <dbReference type="ChEBI" id="CHEBI:58885"/>
            <dbReference type="ChEBI" id="CHEBI:59080"/>
            <dbReference type="ChEBI" id="CHEBI:139493"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </cofactor>
    </comment>
    <comment type="pathway">
        <text evidence="3">Protein modification; protein glycosylation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Endoplasmic reticulum lumen</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the glycosyltransferase 8 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000033869.1">
        <property type="protein sequence ID" value="ENSGAGP00000029829.1"/>
        <property type="gene ID" value="ENSGAGG00000021377.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00378"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005788">
        <property type="term" value="C:endoplasmic reticulum lumen"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003980">
        <property type="term" value="F:UDP-glucose:glycoprotein glucosyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051082">
        <property type="term" value="F:unfolded protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036503">
        <property type="term" value="P:ERAD pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0018279">
        <property type="term" value="P:protein N-linked glycosylation via asparagine"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.550.10">
        <property type="entry name" value="Spore Coat Polysaccharide Biosynthesis Protein SpsA, Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040497">
        <property type="entry name" value="Glyco_transf_24"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029044">
        <property type="entry name" value="Nucleotide-diphossugar_trans"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009448">
        <property type="entry name" value="UDP-g_GGtrans"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040693">
        <property type="entry name" value="UGGT_TRXL_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040694">
        <property type="entry name" value="UGGT_TRXL_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040692">
        <property type="entry name" value="UGGT_TRXL_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040525">
        <property type="entry name" value="UGGT_TRXL_4"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11226">
        <property type="entry name" value="UDP-GLUCOSE GLYCOPROTEIN:GLUCOSYLTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11226:SF3">
        <property type="entry name" value="UDP-GLUCOSE:GLYCOPROTEIN GLUCOSYLTRANSFERASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18404">
        <property type="entry name" value="Glyco_transf_24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18400">
        <property type="entry name" value="Thioredoxin_12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18401">
        <property type="entry name" value="Thioredoxin_13"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18402">
        <property type="entry name" value="Thioredoxin_14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18403">
        <property type="entry name" value="Thioredoxin_15"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06427">
        <property type="entry name" value="UDP-g_GGTase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53448">
        <property type="entry name" value="Nucleotide-diphospho-sugar transferases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="8" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6 12" id="KW-0732">Signal</keyword>
    <keyword evidence="5" id="KW-0808">Transferase</keyword>
    <feature type="signal peptide" evidence="12">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019183272" description="UDP-glucose glycoprotein glucosyltransferase 1" evidence="12">
        <location>
            <begin position="24"/>
            <end position="1397"/>
        </location>
    </feature>
    <feature type="domain" description="UGGT thioredoxin-like" evidence="13">
        <location>
            <begin position="36"/>
            <end position="216"/>
        </location>
    </feature>
    <feature type="domain" description="UGGT thioredoxin-like" evidence="14">
        <location>
            <begin position="290"/>
            <end position="421"/>
        </location>
    </feature>
    <feature type="domain" description="UGGT thioredoxin-like" evidence="15">
        <location>
            <begin position="429"/>
            <end position="677"/>
        </location>
    </feature>
    <feature type="domain" description="UDP-glucose:glycoprotein glucosyltransferase thioredoxin-like" evidence="16">
        <location>
            <begin position="707"/>
            <end position="933"/>
        </location>
    </feature>
    <feature type="domain" description="Glucosyltransferase 24 catalytic" evidence="17">
        <location>
            <begin position="1235"/>
            <end position="1342"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1370"/>
            <end position="1397"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001913"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004922"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00006351"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00045874"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00048456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="Pfam" id="PF18400"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="Pfam" id="PF18401"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="Pfam" id="PF18402"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF18403"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF18404"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029829.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="20">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1397" mass="158462" checksum="828CE49100092318" modified="2019-05-08" version="1">MKVWKAVCTYFVLSISSLAKADSKAVTTSLTTKWLSTPLLLEASEFLAEEGQEKFWTFVEASQDIRSSDHGTNYSSYHAMLKAACQNLSPLQQNLLKFSLSLRSYSATIQAFQQIAADEPPPVGCNSFFAVHGKKTCEFDTLEILLQTASERLKPFLFKGDHRFPLSNPESPVVILYAEIGTEEFSKFHNLLVSKASAGDITYILRHYVANPSKEKVYLSGYGVELAIKSTEYKAKDDTQVKGMEVNATMIGENDPIDEVQGFLFGKLRQLHPDLKEELKELRKHLIESTNEMAPLKVWQLQDLSFQTAARILAAPTVDALMVMKDLSQNFPTKARAITKTVVSSELRTEIEENQKYFKGTLGLQSGDSAFFINGLHIDLDTQDIFSLFDVLRNEARVMEGLHSLGIEGLSLHNVLKLNIQPSDSDYAVDIRSPAISWINNLEVDSRYNSWPSSVQELLRPTFPGVIRQIRKNFHNFVLIVDPSHESTAELLNVAEMFFSNHIPLRIGLVFVVNDSEDVDGLQDAGVALLRAFNYVAQEVDNNYAFQTLISIYNKVKTGDQLQVEHVVSVLEKQYPYVEMNSILGIDSVYDQNRKEGRGYYEQTGVGPLPIVLFNGMPFQKEQLDPDDLETITMQKILETTSLFQRAVYLGELSSDQDVVEYIMNQPNVVPRINSRILMSDREYLDLTATNNFFVDDYARFTFLNSKDKTAAVANSMTYLTKKGMSSKEIYDDSFVRPVTFWIVGDFDKPSGRQLLYDAIKHQKSSNHVRISMINNPSEDPTSKNTVITRAIWAALQTQTSNNAKNFITKMAKEETAKALEAGADITEFAVGGMDVNTFKEAFESPKVDFILSHAMYCRDVLKLQKGQRAVISNGRIIGPLEDNELFNQDDFHLLENIILKTSGQKIKSHIQQLGVEEDLASDLVMKVDALLSAQPKGEARIEYQFFEDRYSAVKLRPKEGETYFDVVAIVDPATRDAQRLAPLLMVLNQLINMNLRVFMNCQSKLSDMPLKSFYRYVLEPEISFTADDNFAPGPIAKFLDMPQTPLFTLNLNTPESWMVESVRTPYDLDNIYLEEVDNRVAAEYELEYLLLEGHCYDITTGQPPRGLQFTLGTSANPVIVDTIVMANLGYFQLKANPGAWVLRLRKGRSEDIYRIYSHDGTDSPPDANEVTVVLNNFKSKIIKVKVQKKLDMMNEDLLSDGTSENESGFWESLKWGFTGGQKNDEMKQDKDDVINIFSVASGHLYERFLRIMMLSVLKHTKTPVKFWFLKNYLSPMFKEFIPYMAEKYNFQYELVQYKWPRWLHQQTEKQRIIWGYKILFLDVLFPLVVDKFLFVDADQVSYLMALGSLAGITRSAYIAEEGSALQQMWQGSPCPSLPTDWRGTEPSRQGASGRAR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP03</accession>
    <name>A0A452IP03_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">C2H2-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IP03"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033589.1">
        <property type="protein sequence ID" value="ENSGAGP00000029579.1"/>
        <property type="gene ID" value="ENSGAGG00000021361.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006356">
        <property type="term" value="P:regulation of transcription by RNA polymerase I"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040436">
        <property type="entry name" value="Disconnected-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15021">
        <property type="entry name" value="DISCONNECTED-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15021:SF1">
        <property type="entry name" value="ZINC FINGER PROTEIN BASONUCLIN-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12874">
        <property type="entry name" value="zf-met"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0862">Zinc</keyword>
    <keyword evidence="1" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="3">
        <location>
            <begin position="327"/>
            <end position="355"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="3">
        <location>
            <begin position="693"/>
            <end position="721"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="3">
        <location>
            <begin position="901"/>
            <end position="924"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="228"/>
            <end position="247"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="268"/>
            <end position="320"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="372"/>
            <end position="393"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="944"/>
            <end position="967"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="230"/>
            <end position="247"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="290"/>
            <end position="299"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="300"/>
            <end position="316"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029579.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="967" mass="108917" checksum="BF24C97C870FB018" modified="2019-05-08" version="1">FAIRCTLVNCSCLCFKPGKINQRQCDQCRHGWVAHALSKLRIPNLYPTSQVEIVQSNVVFDISSLMLYGTQAIPVRLKILLDRLFSVLKQEEVIQILHALDWTLQDYIRGYVLQDASGKVLDHWSIMTNEEELATLQQFLRFGETKSIVELMAIQEKEGQSIIIPTTTTNLDIRAFIESCNQRSPSLSSSVDKMSPANIHHFENIVNNMAFMLPFQFFSPVPPPLIGSPPERHLLEQGQDRSNETKQDVQIPFSESSFLISSSTSFQVENERSINSPDATTKTEDDAPLSDSSSHNTITKLERTELSPENKMKSIEKNGIGPRKGRVFCTACEKTFYDKGTLKIHYNAVHLKIKHKCTIEGCNMVFSSLRSRNRHSANPNPRLHMPMNRNNRDKDLRNSLTIAGPEETKRMDFTILTPDSRPITSYVSHCTDSKVQSSFPSIGQNGVLFPNLKTVQPVLPFYRSPVTPAELANTPGILPSLPLISSSIPEQLVAKGLPFDALPKKKSRKSSMPIKIEKEAVETANENSNLASLEDDTPLQVVSDGELETCEHRIEKQMNDRVEKHPISGNSWKSLSGVEGPKYFDSVIEPNNKYIKETSENELDHSKREVTPEENQALKIASHEMVSEDPEHHHSDHRIPNNDCTELPHHLLTGGLFSALSNRGAAIPCLEDSKDMDHISQHTLGIQKAESRFHCDICKKTFKNPYSVKMHYKNVHLKEMHICTVEGCNAAFPSRRSRDRHSSNLNLHHKLLTKDTLEFKDNYFNATYLLKDMAKEVCQDVSLKQHVGQTSVIFKGMNRTGSLVFPLSKIREPCSENYGYDPMNDGAVLDLSTTSSIKSESSAHSSWDSDGGSEECIMPLDDSDESCEGPSLIPNEELYPDCSVIEKANQSFTNLPSNLPITCHICQKNYSNKGTFRAHYKTVHLRQLHKCKVPGCNTMFSSVRSRNRHSQNPNLHRSLTRSPNSLQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP04</accession>
    <name>A0A452IP04_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="19">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="14">Synaptic cell membrane</location>
            <topology evidence="14">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="15">Belongs to the ligand-gated ion channel (TC 1.A.9) family.</text>
    </comment>
    <comment type="caution">
        <text evidence="15">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IP04"/>
    <dbReference type="STRING" id="38772.ENSGAGP00000029615"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033629.1">
        <property type="protein sequence ID" value="ENSGAGP00000029615.1"/>
        <property type="gene ID" value="ENSGAGG00000021387.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0022848">
        <property type="term" value="F:acetylcholine-gated monoatomic cation-selective channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004888">
        <property type="term" value="F:transmembrane signaling receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd19026">
        <property type="entry name" value="LGIC_ECD_nAChR_B3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd19064">
        <property type="entry name" value="LGIC_TM_nAChR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.70.170.10:FF:000005">
        <property type="entry name" value="Neuronal nicotinic acetylcholine receptor alpha4 subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.390:FF:000001">
        <property type="entry name" value="Neuronal nicotinic acetylcholine receptor subunit 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.70.170.10">
        <property type="entry name" value="Neurotransmitter-gated ion-channel ligand-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.390">
        <property type="entry name" value="Neurotransmitter-gated ion-channel transmembrane domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006202">
        <property type="entry name" value="Neur_chan_lig-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036734">
        <property type="entry name" value="Neur_chan_lig-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006201">
        <property type="entry name" value="Neur_channel"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036719">
        <property type="entry name" value="Neuro-gated_channel_TM_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038050">
        <property type="entry name" value="Neuro_actylchol_rec"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006029">
        <property type="entry name" value="Neurotrans-gated_channel_TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018000">
        <property type="entry name" value="Neurotransmitter_ion_chnl_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002394">
        <property type="entry name" value="Nicotinic_acetylcholine_rcpt"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00860">
        <property type="entry name" value="LIC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18945">
        <property type="entry name" value="NEUROTRANSMITTER GATED ION CHANNEL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02931">
        <property type="entry name" value="Neur_chan_LBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02932">
        <property type="entry name" value="Neur_chan_memb"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00254">
        <property type="entry name" value="NICOTINICR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00252">
        <property type="entry name" value="NRIONCHANNEL"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90112">
        <property type="entry name" value="Neurotransmitter-gated ion-channel transmembrane pore"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF63712">
        <property type="entry name" value="Nicotinic receptor ligand binding domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00236">
        <property type="entry name" value="NEUROTR_ION_CHANNEL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="9" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13 15" id="KW-0407">Ion channel</keyword>
    <keyword evidence="7 15" id="KW-0406">Ion transport</keyword>
    <keyword evidence="12" id="KW-1071">Ligand-gated ion channel</keyword>
    <keyword evidence="8 15" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0675">Receptor</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 16" id="KW-0732">Signal</keyword>
    <keyword evidence="6" id="KW-0770">Synapse</keyword>
    <keyword evidence="3 15" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 15" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="1 15" id="KW-0813">Transport</keyword>
    <feature type="signal peptide" evidence="16">
        <location>
            <begin position="1"/>
            <end position="15"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019259119" evidence="16">
        <location>
            <begin position="16"/>
            <end position="431"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="236"/>
            <end position="259"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="271"/>
            <end position="289"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="301"/>
            <end position="322"/>
        </location>
    </feature>
    <feature type="domain" description="Neurotransmitter-gated ion-channel ligand-binding" evidence="17">
        <location>
            <begin position="32"/>
            <end position="234"/>
        </location>
    </feature>
    <feature type="domain" description="Neurotransmitter-gated ion-channel transmembrane" evidence="18">
        <location>
            <begin position="242"/>
            <end position="362"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00034099"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="RuleBase" id="RU000687"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF02931"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF02932"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029615.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="431" mass="49202" checksum="35B9CD2C1E6CC605" modified="2019-05-08" version="1">MLLLLDLTLVPLILCVSNSAVAGLSLIAENEDALLRHLFHGYQKWVRPVENSNDTIKVLFGLKISQLVDVDEKNQLMTTNVWLKQEWTDHKLCWNPEDYGGITAIRVPSESLWLPDIVLFENADGRFEGSLMTKVIVKYNGLVTWTPPASYKSSCTMDVTFFPFDRQNCSMKFGSWTYDGNMVDLILVDENVDRKDFFDNGEWEILKAKGMKGSRKDGLHSYPFITYSFVLRRLPLFYTLFLIVPCLGLSFLTVLVFYLPSDEGEKLSLSTSVLVSLTVFLLVIEEIIPSSSKVIPLIGEYLLFIMIFVTLSIIVTVFVINVHHRSSATYHPMAPWVKRLFLQKLPRLLCLKGHVDRYTFSEAAGTIPVLKSKLSGKKKQKQVKDGEKIVIAFLEKAADSIRYISGHVKKEHFIRQVMLTHEEYMDVLTEK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP07</accession>
    <name>A0A452IP07_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Hypoxanthine phosphoribosyltransferase</fullName>
            <ecNumber evidence="5">2.4.2.8</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="5">
            <text>IMP + diphosphate = hypoxanthine + 5-phospho-alpha-D-ribose 1-diphosphate</text>
            <dbReference type="Rhea" id="RHEA:17973"/>
            <dbReference type="ChEBI" id="CHEBI:17368"/>
            <dbReference type="ChEBI" id="CHEBI:33019"/>
            <dbReference type="ChEBI" id="CHEBI:58017"/>
            <dbReference type="ChEBI" id="CHEBI:58053"/>
            <dbReference type="EC" id="2.4.2.8"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="5">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="pathway">
        <text evidence="5">Purine metabolism; IMP biosynthesis via salvage pathway; IMP from hypoxanthine: step 1/1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="5">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="1 5">Belongs to the purine/pyrimidine phosphoribosyltransferase family.</text>
    </comment>
    <dbReference type="EC" id="2.4.2.8" evidence="5"/>
    <dbReference type="AlphaFoldDB" id="A0A452IP07"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033656.1">
        <property type="protein sequence ID" value="ENSGAGP00000029641.1"/>
        <property type="gene ID" value="ENSGAGG00000021401.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00591">
        <property type="reaction ID" value="UER00648"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004422">
        <property type="term" value="F:hypoxanthine phosphoribosyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000166">
        <property type="term" value="F:nucleotide binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032264">
        <property type="term" value="P:IMP salvage"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006166">
        <property type="term" value="P:purine ribonucleoside salvage"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd06223">
        <property type="entry name" value="PRTases_typeI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.2020:FF:000038">
        <property type="entry name" value="Hypoxanthine phosphoribosyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.2020">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050408">
        <property type="entry name" value="HGPRT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005904">
        <property type="entry name" value="Hxn_phspho_trans"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029057">
        <property type="entry name" value="PRTase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000836">
        <property type="entry name" value="PRTase_dom"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01203">
        <property type="entry name" value="HGPRTase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43340">
        <property type="entry name" value="HYPOXANTHINE-GUANINE PHOSPHORIBOSYLTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43340:SF5">
        <property type="entry name" value="PHOSPHORIBOSYLTRANSFERASE DOMAIN-CONTAINING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00156">
        <property type="entry name" value="Pribosyltran"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53271">
        <property type="entry name" value="PRTase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="5" id="KW-0328">Glycosyltransferase</keyword>
    <keyword evidence="4 5" id="KW-0460">Magnesium</keyword>
    <keyword evidence="2 5" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="3 5" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="5" id="KW-0660">Purine salvage</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Phosphoribosyltransferase" evidence="6">
        <location>
            <begin position="48"/>
            <end position="197"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00008391"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="RuleBase" id="RU364099"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="Pfam" id="PF00156"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029641.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="220" mass="25161" checksum="260E5983B2684C72" modified="2019-05-08" version="1">GLLRYSSPHRDISDDWPGYSLDLFTYPQHYYGDLEYVLIPHGVIVDRTECLAKDIMQDIGDNNIVVLCVLKGGYKFCADLVEYIETLSRNSERSISMKVDFVRLKSYHNDQSTQDMQIIGGEDLSKLTGKSVLIVEDIIGTGRTMKALLNNIEKYKPKMIKVASLLVKRTSQPDGYRPDYYGFEIPNVFVVGYALEYNEHFRDLNHICVINDNGKGKYKV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IP09</accession>
    <name>A0A452IP09_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">pantothenate kinase</fullName>
            <ecNumber evidence="5">2.7.1.33</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>(R)-pantothenate + ATP = (R)-4'-phosphopantothenate + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:16373"/>
            <dbReference type="ChEBI" id="CHEBI:10986"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29032"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.1.33"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="3">Cofactor biosynthesis; coenzyme A biosynthesis; CoA from (R)-pantothenate: step 1/5.</text>
    </comment>
    <comment type="subunit">
        <text evidence="4">Homodimer.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="13">Belongs to the type II pantothenate kinase family.</text>
    </comment>
    <dbReference type="EC" id="2.7.1.33" evidence="5"/>
    <dbReference type="AlphaFoldDB" id="A0A452IP09"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033733.1">
        <property type="protein sequence ID" value="ENSGAGP00000029711.1"/>
        <property type="gene ID" value="ENSGAGG00000021410.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004594">
        <property type="term" value="F:pantothenate kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015937">
        <property type="term" value="P:coenzyme A biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd24135">
        <property type="entry name" value="ASKHA_NBD_PanK-II_Pank1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.420.40:FF:000244">
        <property type="entry name" value="pantothenate kinase 1-like isoform X3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.420.510:FF:000001">
        <property type="entry name" value="pantothenate kinase 2, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.420.40:FF:000392">
        <property type="entry name" value="pantothenate kinase 3-like isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.420.40">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.420.510">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043129">
        <property type="entry name" value="ATPase_NBD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004567">
        <property type="entry name" value="Type_II_PanK"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00555">
        <property type="entry name" value="panK_eukar"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12280">
        <property type="entry name" value="PANTOTHENATE KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12280:SF23">
        <property type="entry name" value="PANTOTHENATE KINASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03630">
        <property type="entry name" value="Fumble"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53067">
        <property type="entry name" value="Actin-like ATPase domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="11" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="12" id="KW-0173">Coenzyme A biosynthesis</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="10" id="KW-0418">Kinase</keyword>
    <keyword evidence="14" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0808">Transferase</keyword>
    <keyword evidence="14" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="14" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="59"/>
            <end position="77"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001206"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005225"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00011738"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012102"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022993"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00060870"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029711.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="436" mass="48985" checksum="50E2801213AE7E70" modified="2019-05-08" version="1">RSWWWGRERRKLIVGDVVVLLPLLLNCCWQRHCLQSWAAGEWRLLAGIPVLKAEVPSSIFIAPLSFFFPPFLAFPWFGMDIGGTLVKLVYFEPKDITAEEEQEEVENLKSIRKYLTSNTAYGKTGIRDVHLELKNLTMCGRKGNLHFIRFPTSAMHRFIQMGSEKNFSSLHTTLCATGGGAYKFEEDFRTIADLQLHKLDELDCLIQGLLYVDSVGYNGQPECYYFENPTDPEQCQKKPYCLDNPYPMLLVNIGSGVSILAVYSKDNYKRVTGTSLGGGTFLGLCCLLTGCETFEEALEMAAKGDSTNVDKLVKDIYGGDYERFGLQGSAVASSFGHMMSKEKRDSISKEDLARATLVTITNNIGSIARMCALNENIDRVVFVGNFLRINMISMKVLAYAMDYWSKGQLKALFLEHEGYFGAVGALLELLKMPENQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0H0</accession>
    <name>A0A452J0H0_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="14">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038342.1">
        <property type="protein sequence ID" value="ENSGAGP00000033852.1"/>
        <property type="gene ID" value="ENSGAGG00000024077.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003714">
        <property type="term" value="F:transcription corepressor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006351">
        <property type="term" value="P:DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="6.10.140.2220:FF:000001">
        <property type="entry name" value="CBFA2/RUNX1 translocation partner 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.1110:FF:000001">
        <property type="entry name" value="RUNX1 translocation partner 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.2220">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.230">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.1110">
        <property type="entry name" value="TAFH/NHR1 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013289">
        <property type="entry name" value="CBFA2T1/2/3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013291">
        <property type="entry name" value="MTGR1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014896">
        <property type="entry name" value="NHR2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037249">
        <property type="entry name" value="TAFH/NHR1_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003894">
        <property type="entry name" value="TAFH_NHR1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002893">
        <property type="entry name" value="Znf_MYND"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10379">
        <property type="entry name" value="MTG8 ETO EIGHT TWENTY ONE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10379:SF13">
        <property type="entry name" value="PROTEIN CBFA2T2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08788">
        <property type="entry name" value="NHR2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07531">
        <property type="entry name" value="TAFH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01753">
        <property type="entry name" value="zf-MYND"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01875">
        <property type="entry name" value="ETOFAMILY"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01877">
        <property type="entry name" value="MTGR1PROTEIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00549">
        <property type="entry name" value="TAFH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF144232">
        <property type="entry name" value="HIT/MYND zinc finger-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF158553">
        <property type="entry name" value="TAFH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51119">
        <property type="entry name" value="TAFH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01360">
        <property type="entry name" value="ZF_MYND_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50865">
        <property type="entry name" value="ZF_MYND_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="10" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0678">Repressor</keyword>
    <keyword evidence="7" id="KW-0804">Transcription</keyword>
    <keyword evidence="6" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <keyword evidence="4 9" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="TAFH" evidence="13">
        <location>
            <begin position="99"/>
            <end position="194"/>
        </location>
    </feature>
    <feature type="domain" description="MYND-type" evidence="12">
        <location>
            <begin position="492"/>
            <end position="528"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="17"/>
            <end position="93"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="219"/>
            <end position="242"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="382"/>
            <end position="413"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="551"/>
            <end position="586"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="10">
        <location>
            <begin position="431"/>
            <end position="473"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="11">
        <location>
            <begin position="32"/>
            <end position="45"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="50"/>
            <end position="65"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="222"/>
            <end position="236"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="394"/>
            <end position="404"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="565"/>
            <end position="580"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022491"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50865"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS51119"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033852.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="586" mass="64642" checksum="DE69301A497A58B8" modified="2019-05-08" version="1">MKGVPCISFEKRVPVMPGSPVEVKIQSRSSPPSMPPLPPVNPGGPRPLSFTPTALPNGINHSPPTLNGAPSPPQRFSNGPASSSSSSLTNQQLPATCGARQLSKLKRFLTTLQQFGNDISPEIGEKVRTLVLALVNSTVTIEEFHCKLQEATNFPLRPFVIPFLKANLPLLQRELLHCARAAKQTPSQYLAQHEHILLNTNTTSPADSSELLMEVNGNGKRHXADGKENSFERDPLPSEPPAKRVCTISPAPRHSPALALPLMNPAGQFHPTPPPLQHYTLEDIATSHLYRDPNKLLEQRDMRDRHSNLGLNGGYQDELVDHRLTEREWADEWKHLDHALNCIMEMVEKTRRSMAVLRRCQEADREELNYWKRRCSESAETRKGGSEFISRQHSPGSSDSISSDSQRDFSSRSGTGYVTEEIWKKAEEAVNEVKRQAMSEVQKAVAEAEQKAFEMIASERARMEQTIADAKRQATEDAFLVINEQEESTESCWNCGRKASETCSGCNIARYCGSFCQHKDWERHHRICGQGLHSQAKPLSLPAGRLAAAAAKPADGVPSPALEKTSATTSRSSTPASVTAIDTNGL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0H1</accession>
    <name>A0A452J0H1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">GRIP domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000038386.1">
        <property type="protein sequence ID" value="ENSGAGP00000033893.1"/>
        <property type="gene ID" value="ENSGAGG00000024101.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031267">
        <property type="term" value="F:small GTPase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048193">
        <property type="term" value="P:Golgi vesicle transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.220.60">
        <property type="entry name" value="GRIP domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000237">
        <property type="entry name" value="GRIP_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19327">
        <property type="entry name" value="GOLGIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19327:SF0">
        <property type="entry name" value="GOLGIN SUBFAMILY A MEMBER 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01465">
        <property type="entry name" value="GRIP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00755">
        <property type="entry name" value="Grip"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF101283">
        <property type="entry name" value="GRIP domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50913">
        <property type="entry name" value="GRIP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="GRIP" evidence="3">
        <location>
            <begin position="2226"/>
            <end position="2273"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1"/>
            <end position="62"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="82"/>
            <end position="127"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="139"/>
            <end position="197"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="281"/>
            <end position="319"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="197"/>
            <end position="245"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="323"/>
            <end position="697"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="753"/>
            <end position="1027"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="1062"/>
            <end position="1192"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="1353"/>
            <end position="1412"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="1454"/>
            <end position="1488"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="1521"/>
            <end position="1862"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="1898"/>
            <end position="2076"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="2107"/>
            <end position="2159"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="25"/>
            <end position="49"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="89"/>
            <end position="124"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="140"/>
            <end position="168"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="169"/>
            <end position="178"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="284"/>
            <end position="295"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS50913"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033893.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2285" mass="268047" checksum="90B33B94531D62AA" modified="2019-05-08" version="1">MFKKLRQKIVEEQTQPRSPGGRAPQQPQTYSKSSPPSPGNRSRTSSLTEQSDEGTLTPDKELLAGMIAEPAFLSEYTIFALDPKRPKPQSDSVNVSKQTTPKSTDSNGSEPPSPQLSDTQTFAQKLQLRVPSMESLFRSPVKESLFRSSSKESLVRTSSRESLNRLDFDTPGPTFDPPSDVESEAEESLGNVDSLGKEQLLQRLRRMERSLGSYRGKYSELVTAYQTVQREKKKLQELQMDQQAKKHLQEEFDASLEEKDQLISVLQTQVSLLKQRLQNGQISTELPETNTETEPQSPAKEISAENVLEPGGTEGDGDSAKTLETLIKRVKRQENLLQRCKEMIRSHKERSAQLTNEKEALQEQLEERLQELEKMKELHMTEKTKLITQLRDAKNLIEQLEQDKGMVIAETKRQMHETMEMKEEEIAQLRAHIKQITTQGEELREQKEKSEKAAFEELEKALSTAQKVEEARKKLQAEMDEKIKAIEKTSEEERVTLQQELTRVKQEVVEIMKKSSEERVAELEKLHKRELATKEQELNERLQVQGKEFQEKMRAALEKSQSEFLQTVQEKEQQESLALEELELQKKAIQSECEKKLQDMQQEIQTARTRILELESSLAKYSQDDKRQLEELSTLIESEKTKYNKEIIVMVEKHREELENVKQQEEELWTEKLQILKQQQQSTIEKLREKHEQEMDIMLKEKETVFRAHVEEMNAKTLEKLDVKQTELEALSFELSETLQIRHDLEKDGDKIKQESETKLEQQRKQYQEKMDMMVKEHDISVQGIQKAHTEETNQLKVQLEEKEKHLEELKAQEHKLKESLEISEAELKQVSAKLGDLCQSHQNNSSEQARVYEGQLTELQQKLTNMEAEKSHLKEQIVRAESQLNEVRTELDTYISRVRDLRQQLEEQSSENMQKITSLKQKCESQLKDLSTETDQAKRDLLERENQLEQLKKQQNQQVEELRQKLSAKEERISALQVEYENSLKDQGTKMEKIKQKAKEMQEMFKRKLSEQEAKLKKEFENKQLEFSQKEKHFNAKMLEMAHASSTGINDAVSKLEMNQKEQLESLAEAHRWELEEVAQTWGKKLNQQVEELKEKHEMELHKKEKEVGDLKQKLATFSAEKEGTNIEITLLKEEQVKRDESWNELQEQLKQLLAQVNTLSQSETDLKAQLEKLEDDLSHSLKDRAVLQEQLREQKTMEEKHKIKVTELTDMLKTSDEKLQILESSHCKEREDYEKKLEDKHLASQQKQIEFERLSSELAAQLDIYWKEADTLLQTKTNEVIEKCTEKINLVISRIAHCQHQTTKIKKAILIKNCKIPELETQLRQITEHQTTVSSSLQESTQQLQDKENVIILMKADIERLVTEKELLQKEGGHQQQAATEKETCITQLRKELSENINVVTSMREELKEKESEICALNKLVNELNVKLENSIGLTEKEAAISLLSKQHQEDQLQLLNQVQELSSRVESLSQEKASALEQVDHWMNKLSEWKKKAQLRFTQNHNTIKELQNKLELNNIQVGGKDEEIHQLKEELDKQSRNLDSLKDVMEQKQRRREKEEGKLTAELKIQTARITELEEHIARKTAENDSLMEELKRHNQQRDTEQKELVQQLQQTEVVVVEKDNRLKEAEEKVLSLEKQMDSMKAELDTKQREFEQMKTAMMKSKEEELKMLEDRMNAENTTKLADLKKKAEQKIASVKKQLTSQMEEKEQQFKLAMENQLKDLEQKMQEREAKIMDLEEKNRLIRDSADLEREMMQKVENEKASLEQEKASVLENVRQMYEEKISEIQKDLIDKDGLLNKYENEQQESNDSKLKMKSKQEELLKKLECTEKKLKEEQSVIERLQKEIEEKNKKYSLLLDQHTQDGDDLASIREELKAQVQKHYDMENVIGDFQKQFQEKDKVSQSLEQKIRELEDDIVKKKEVCKIEMAEVSLKYEEKLQGLQQQLDERNRQLKIFEENAEEKTRSDLELQKLLGDMQTQQKDLQAKLKEAETEKQKLRKEISKVQKDLRTLRKEHQQELDIMKKESLEEMEQKIRIEQEDIELKHNSTLKQLMREFNTQLSQKERELETAVHETISKAQDVETELIETHHAETTQLHRKIAEKDDDLKRTVKKYEEILEAREEEMTAKVNELQTQLEELQKEYKQKLADEENRRREEVTITELQIQLAQKTTIVNDSKLKEQEFIEQIHILKDQLKNYEKNVYVTAVGSPYRDGNLCHTDVSLFGEPTEFEYLRKVLFEYMMGRETKTMAKVITTVLKFPADQTQKILEREDARPLSWLRPS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="36" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0H2</accession>
    <name>A0A452J0H2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="26">Histone-lysine N-methyltransferase NSD3</fullName>
            <ecNumber evidence="24">2.1.1.370</ecNumber>
            <ecNumber evidence="25">2.1.1.371</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="27">Nuclear SET domain-containing protein 3</fullName>
        </alternativeName>
    </protein>
    <organism evidence="36 37">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="37" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="36" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="36" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="22">Histone methyltransferase. Preferentially dimethylates 'Lys-4' and 'Lys-27' of histone H3 forming H3K4me2 and H3K27me2. H3 'Lys-4' methylation represents a specific tag for epigenetic transcriptional activation, while 'Lys-27' is a mark for transcriptional repression.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="20">
            <text>L-lysyl(27)-[histone H3] + 2 S-adenosyl-L-methionine = N(6),N(6)-dimethyl-L-lysyl(27)-[histone H3] + 2 S-adenosyl-L-homocysteine + 2 H(+)</text>
            <dbReference type="Rhea" id="RHEA:64452"/>
            <dbReference type="Rhea" id="RHEA-COMP:15539"/>
            <dbReference type="Rhea" id="RHEA-COMP:15548"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29969"/>
            <dbReference type="ChEBI" id="CHEBI:57856"/>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
            <dbReference type="ChEBI" id="CHEBI:61976"/>
            <dbReference type="EC" id="2.1.1.371"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="21">
            <text>L-lysyl(4)-[histone H3] + 2 S-adenosyl-L-methionine = N(6),N(6)-dimethyl-L-lysyl(4)-[histone H3] + 2 S-adenosyl-L-homocysteine + 2 H(+)</text>
            <dbReference type="Rhea" id="RHEA:64448"/>
            <dbReference type="Rhea" id="RHEA-COMP:15540"/>
            <dbReference type="Rhea" id="RHEA-COMP:15547"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29969"/>
            <dbReference type="ChEBI" id="CHEBI:57856"/>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
            <dbReference type="ChEBI" id="CHEBI:61976"/>
            <dbReference type="EC" id="2.1.1.370"/>
        </reaction>
    </comment>
    <comment type="subunit">
        <text evidence="23">Interacts with BRD4. Interacts (via KIKL motif) with BRD3 (via NET domain).</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Chromosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="2.1.1.370" evidence="24"/>
    <dbReference type="EC" id="2.1.1.371" evidence="25"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038345.1">
        <property type="protein sequence ID" value="ENSGAGP00000033854.1"/>
        <property type="gene ID" value="ENSGAGG00000024078.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005694">
        <property type="term" value="C:chromosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140952">
        <property type="term" value="F:histone H3K27 dimethyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140946">
        <property type="term" value="F:histone H3K4 dimethyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032259">
        <property type="term" value="P:methylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006355">
        <property type="term" value="P:regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd15649">
        <property type="entry name" value="PHD1_NSD3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd15652">
        <property type="entry name" value="PHD2_NSD3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd15655">
        <property type="entry name" value="PHD3_NSD3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd15658">
        <property type="entry name" value="PHD4_NSD3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd15661">
        <property type="entry name" value="PHD5_NSD3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd20163">
        <property type="entry name" value="PWWP_NSD3_rpt1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd20166">
        <property type="entry name" value="PWWP_NSD3_rpt2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd19212">
        <property type="entry name" value="SET_NSD3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.170.270.10:FF:000002">
        <property type="entry name" value="Histone-lysine N-methyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.140:FF:000004">
        <property type="entry name" value="Histone-lysine N-methyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.140:FF:000030">
        <property type="entry name" value="Histone-lysine N-methyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.40.10:FF:000025">
        <property type="entry name" value="Histone-lysine N-methyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.40.10:FF:000106">
        <property type="entry name" value="Histone-lysine N-methyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.40.10:FF:000205">
        <property type="entry name" value="Histone-lysine N-methyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.40.10:FF:001336">
        <property type="entry name" value="Histone-lysine N-methyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.140">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.170.270.10">
        <property type="entry name" value="SET domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006560">
        <property type="entry name" value="AWS_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041306">
        <property type="entry name" value="C5HCH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055198">
        <property type="entry name" value="NSD_PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR059153">
        <property type="entry name" value="NSD_PHD-1st"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047456">
        <property type="entry name" value="PHD2_NSD3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047458">
        <property type="entry name" value="PHD4_NSD3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047527">
        <property type="entry name" value="PHD5_NSD3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055197">
        <property type="entry name" value="PHDvar_NSD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003616">
        <property type="entry name" value="Post-SET_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000313">
        <property type="entry name" value="PWWP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047451">
        <property type="entry name" value="PWWP_NSD3_rpt1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047453">
        <property type="entry name" value="PWWP_NSD3_rpt2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050777">
        <property type="entry name" value="SET2_Histone-Lys_MeTrsfase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001214">
        <property type="entry name" value="SET_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046341">
        <property type="entry name" value="SET_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047461">
        <property type="entry name" value="SET_NSD3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019786">
        <property type="entry name" value="Zinc_finger_PHD-type_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011011">
        <property type="entry name" value="Znf_FYVE_PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001965">
        <property type="entry name" value="Znf_PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019787">
        <property type="entry name" value="Znf_PHD-finger"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22884">
        <property type="entry name" value="SET DOMAIN PROTEINS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17907">
        <property type="entry name" value="AWS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17982">
        <property type="entry name" value="C5HCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23011">
        <property type="entry name" value="PHD-1st_NSD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22908">
        <property type="entry name" value="PHD_NSD"/>
        <property type="match status" value="1"/>
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    <dbReference type="Pfam" id="PF23004">
        <property type="entry name" value="PHDvar_NSD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00855">
        <property type="entry name" value="PWWP"/>
        <property type="match status" value="2"/>
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    <dbReference type="Pfam" id="PF00856">
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        <property type="match status" value="1"/>
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    <dbReference type="SMART" id="SM00570">
        <property type="entry name" value="AWS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00249">
        <property type="entry name" value="PHD"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00508">
        <property type="entry name" value="PostSET"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00293">
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        <property type="match status" value="2"/>
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    <dbReference type="SMART" id="SM00317">
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        <property type="match status" value="1"/>
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    <dbReference type="SUPFAM" id="SSF57903">
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        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF82199">
        <property type="entry name" value="SET domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF63748">
        <property type="entry name" value="Tudor/PWWP/MBT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51215">
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        <property type="match status" value="1"/>
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    <dbReference type="PROSITE" id="PS50868">
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    <dbReference type="PROSITE" id="PS50280">
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        <property type="match status" value="1"/>
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    <dbReference type="PROSITE" id="PS01359">
        <property type="entry name" value="ZF_PHD_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50016">
        <property type="entry name" value="ZF_PHD_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="15" id="KW-0007">Acetylation</keyword>
    <keyword evidence="14" id="KW-0156">Chromatin regulator</keyword>
    <keyword evidence="3" id="KW-0158">Chromosome</keyword>
    <keyword evidence="17 29" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="9" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="6" id="KW-0489">Methyltransferase</keyword>
    <keyword evidence="19" id="KW-0539">Nucleus</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="37" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0677">Repeat</keyword>
    <keyword evidence="8" id="KW-0949">S-adenosyl-L-methionine</keyword>
    <keyword evidence="18" id="KW-0804">Transcription</keyword>
    <keyword evidence="16" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="7" id="KW-0808">Transferase</keyword>
    <keyword evidence="13" id="KW-0832">Ubl conjugation</keyword>
    <keyword evidence="12" id="KW-0862">Zinc</keyword>
    <keyword evidence="11 28" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="PWWP" evidence="33">
        <location>
            <begin position="271"/>
            <end position="334"/>
        </location>
    </feature>
    <feature type="domain" description="PHD-type" evidence="31">
        <location>
            <begin position="703"/>
            <end position="750"/>
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    </feature>
    <feature type="domain" description="PHD-type" evidence="31">
        <location>
            <begin position="864"/>
            <end position="908"/>
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        <location>
            <begin position="913"/>
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            <begin position="1046"/>
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    <feature type="domain" description="SET" evidence="32">
        <location>
            <begin position="1098"/>
            <end position="1215"/>
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            <begin position="1222"/>
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    </feature>
    <feature type="region of interest" description="Disordered" evidence="30">
        <location>
            <begin position="123"/>
            <end position="154"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="30">
        <location>
            <begin position="190"/>
            <end position="241"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="30">
        <location>
            <begin position="345"/>
            <end position="368"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="30">
        <location>
            <begin position="406"/>
            <end position="468"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="30">
        <location>
            <begin position="545"/>
            <end position="700"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="29">
        <location>
            <begin position="987"/>
            <end position="1014"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="29">
        <location>
            <begin position="1248"/>
            <end position="1275"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="30">
        <location>
            <begin position="131"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="30">
        <location>
            <begin position="205"/>
            <end position="232"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="30">
        <location>
            <begin position="428"/>
            <end position="446"/>
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    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="30">
        <location>
            <begin position="545"/>
            <end position="565"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="30">
        <location>
            <begin position="585"/>
            <end position="597"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="30">
        <location>
            <begin position="673"/>
            <end position="693"/>
        </location>
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        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
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    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004286"/>
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    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022454"/>
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    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
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    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022603"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022691"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00050891"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00052270"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00056539"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00065976"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00066818"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00066819"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00068930"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="ARBA" id="ARBA00081740"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00146"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="31">
        <source>
            <dbReference type="PROSITE" id="PS50016"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="32">
        <source>
            <dbReference type="PROSITE" id="PS50280"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="33">
        <source>
            <dbReference type="PROSITE" id="PS50812"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="34">
        <source>
            <dbReference type="PROSITE" id="PS50868"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="35">
        <source>
            <dbReference type="PROSITE" id="PS51215"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="36">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033854.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="37">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1390" mass="156314" checksum="E470C36AD6B5621D" modified="2019-05-08" version="1">MDFSFSFMQGIMGNTIQQPPQLIDSANIRQEDAFDASSDIAEDGGQTPYETALQQGFQYPTPTTEDLPPITNGYPPTINMYEPQAKYQTYSQYPNGSANGFGAVRNFSPPEYYQMENSNMRPHEVLEKSSPPQPPPPPPPSVPQTVIPKKTGSPEIKLKITKTIQNGRELFESSLCGDLLNEVQASEYAKMKHEGRKEKRKKSSKHDSSRSEERKSHKIPKLEPEEQNRPNERLSVLSERPREDTLLEETPVQQFSPSLPTQVTTDIKFQVGDLVWSKVGTYPWWPCMVSSDPQLEVHTKINTRGAREYHVQFFSNQPERAWVHEKRVREYKGHKQYEQLLAEATKQASNHSEKQKIRKPRPQRERAQWDIGIAHAEKALKMTREERIEQYTFIYIDKEPEEALSKAKKTAAPKSEAKKNRRPKPALNAQTEQTKVVSASPSLSNTEVRRQSQRRQSSVEEEEPPPPVKIAWKTAAARKSLPASITMHKGSLDLQKCNMSPVVKIEQVFALQNAAGDGKFIDQFVYSTKGVGNKTEISVKGQDKLISTSNQKSERATAQNASSPETTSGSTGSVEKKQQRRSIRTRSESEKSTEVVPKKKIKKEQVEMVPQTAVKTGLQKGASEISDSCKPLKKRSRASTDVELTSSAYRDTSDSDSRGLNDLQGSFGKHSDSPSATADADASDVQSVDSSLSRRGTGTNKKDTVCQICESSGESLVSCEAECCRVFHMECLGLKSIPDGKFICMECKNGQHTCFSCKLPGKDVKRCSVNACGKFYHEACVRKFATAVFESRGFRCPQHCCTACSTDKDIHKASKGRMVRCLRCPVAYHSGDGCIAAGSLFVSSHILICSNHSKRNHSSSAVNVGFCFVCARGGKLLCCESCPASFHPECLNIEMPEGCWNCNDCKAGKKLRYKQIVWVKLGNYRWWPAEICNPRSVPLNIQGLKHDIGDFPVFFFGSHDYYWVHQGRVFPYVEGDKSFAEGQTSINKTFKKALEEAAKRFQELKAQRESKEALEIERNSRKPPPYKHIKSNKVIGKVQIQVADLSEIPRCNCKPSDENPCGLESECLNRMLQYECHPQVCPAGERCQNQCFTKRLYPDAEIIKTDRRGWGLRTKRNIKKGEFVNEYVGELIDEEECRLRIKRAHENSITNFYMLTVTKDRIIDAGPKGNFSRFMNHSCNPNCETQKWTVNGDVRVGLFALRDIPAGMELTFNYNLDCLGNGRTECHCGAENCSGFLGVRPKTAFATANEEKAKNAKLKQKKRKIKTEQKQMHEDNCFQCGDGGELVMCDKKDCPKAYHLLCLNLTQPPFGKWECPWHQCDICCNPAISFCEFCPRSFCKDHEKGALVASALDGCLCCSEHNPKSPMAPEYWSKIKCKLQPQNPGEEAKE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0H6</accession>
    <name>A0A452J0H6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Spastin</fullName>
            <ecNumber evidence="10">5.6.1.1</ecNumber>
        </recommendedName>
    </protein>
    <gene>
        <name evidence="10" type="primary">SPAST</name>
        <name evidence="10" type="synonym">SPG4</name>
    </gene>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">ATP-dependent microtubule severing protein that specifically recognizes and cuts microtubules that are polyglutamylated. Preferentially recognizes and acts on microtubules decorated with short polyglutamate tails: severing activity increases as the number of glutamates per tubulin rises from one to eight, but decreases beyond this glutamylation threshold. Microtubule severing promotes reorganization of cellular microtubule arrays and the release of microtubules from the centrosome following nucleation. Required for membrane traffic from the endoplasmic reticulum (ER) to the Golgi and for completion of the abscission stage of cytokinesis. Also plays a role in axon growth and the formation of axonal branches.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>n ATP + n H2O + a microtubule = n ADP + n phosphate + (n+1) alpha/beta tubulin heterodimers.</text>
            <dbReference type="EC" id="5.6.1.1"/>
        </reaction>
    </comment>
    <comment type="subunit">
        <text evidence="10">Homohexamer. The homohexamer is stabilized by ATP-binding. The homohexamer may adopt a ring conformation through which microtubules pass prior to being severed. Interacts with microtubules.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="10">Membrane</location>
            <topology evidence="10">Peripheral membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="10">Cytoplasm</location>
            <location evidence="10">Cytoskeleton</location>
            <location evidence="10">Microtubule organizing center</location>
            <location evidence="10">Centrosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="10">Cytoplasm</location>
            <location evidence="10">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="10">Cytoplasm</location>
            <location evidence="10">Perinuclear region</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="10">Nucleus</location>
        </subcellularLocation>
        <text evidence="10">Forms an intramembrane hairpin-like structure in the membrane.</text>
    </comment>
    <comment type="similarity">
        <text evidence="10">Belongs to the AAA ATPase family. Spastin subfamily.</text>
    </comment>
    <dbReference type="EC" id="5.6.1.1" evidence="10"/>
    <dbReference type="AlphaFoldDB" id="A0A452J0H6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038391.1">
        <property type="protein sequence ID" value="ENSGAGP00000033898.1"/>
        <property type="gene ID" value="ENSGAGG00000024111.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005874">
        <property type="term" value="C:microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048471">
        <property type="term" value="C:perinuclear region of cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005819">
        <property type="term" value="C:spindle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008017">
        <property type="term" value="F:microtubule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008568">
        <property type="term" value="F:microtubule severing ATPase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007409">
        <property type="term" value="P:axonogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032506">
        <property type="term" value="P:cytokinetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006888">
        <property type="term" value="P:endoplasmic reticulum to Golgi vesicle-mediated transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051013">
        <property type="term" value="P:microtubule severing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031117">
        <property type="term" value="P:positive regulation of microtubule depolymerization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034214">
        <property type="term" value="P:protein hexamerization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="CDD" id="cd02679">
        <property type="entry name" value="MIT_spastin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd19524">
        <property type="entry name" value="RecA-like_spastin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000093">
        <property type="entry name" value="Fidgetin-like 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.8.60:FF:000036">
        <property type="entry name" value="Spastin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.80:FF:000006">
        <property type="entry name" value="Spastin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.8.60">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.80">
        <property type="entry name" value="Phosphotransferase system, lactose/cellobiose-type IIA subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="HAMAP" id="MF_03021">
        <property type="entry name" value="Spastin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003593">
        <property type="entry name" value="AAA+_ATPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041569">
        <property type="entry name" value="AAA_lid_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003959">
        <property type="entry name" value="ATPase_AAA_core"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003960">
        <property type="entry name" value="ATPase_AAA_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007330">
        <property type="entry name" value="MIT_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050304">
        <property type="entry name" value="MT-severing_AAA_ATPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015415">
        <property type="entry name" value="Spast_Vps4_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017179">
        <property type="entry name" value="Spastin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23074">
        <property type="entry name" value="AAA DOMAIN-CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23074:SF86">
        <property type="entry name" value="SPASTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00004">
        <property type="entry name" value="AAA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17862">
        <property type="entry name" value="AAA_lid_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09336">
        <property type="entry name" value="Vps4_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00382">
        <property type="entry name" value="AAA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00745">
        <property type="entry name" value="MIT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00674">
        <property type="entry name" value="AAA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 10 11" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="10" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="2 10" id="KW-0132">Cell division</keyword>
    <keyword evidence="1 10" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7 10" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="10" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="10" id="KW-0221">Differentiation</keyword>
    <keyword evidence="8 10" id="KW-0413">Isomerase</keyword>
    <keyword evidence="6 10 13" id="KW-0472">Membrane</keyword>
    <keyword evidence="3 10" id="KW-0493">Microtubule</keyword>
    <keyword evidence="10" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="4 10 11" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="13" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="13" id="KW-1133">Transmembrane helix</keyword>
    <feature type="topological domain" description="Cytoplasmic" evidence="10">
        <location>
            <begin position="1"/>
            <end position="61"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="13">
        <location>
            <begin position="62"/>
            <end position="83"/>
        </location>
    </feature>
    <feature type="intramembrane region" description="Helical" evidence="10">
        <location>
            <begin position="62"/>
            <end position="82"/>
        </location>
    </feature>
    <feature type="topological domain" description="Cytoplasmic" evidence="10">
        <location>
            <begin position="83"/>
            <end position="584"/>
        </location>
    </feature>
    <feature type="domain" description="MIT" evidence="15">
        <location>
            <begin position="114"/>
            <end position="192"/>
        </location>
    </feature>
    <feature type="domain" description="AAA+ ATPase" evidence="14">
        <location>
            <begin position="342"/>
            <end position="478"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1"/>
            <end position="45"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="248"/>
            <end position="277"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="13"/>
            <end position="29"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="248"/>
            <end position="274"/>
        </location>
    </feature>
    <feature type="binding site" evidence="10">
        <location>
            <begin position="350"/>
            <end position="357"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023235"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00036378"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="HAMAP-Rule" id="MF_03021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="RuleBase" id="RU003651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="SMART" id="SM00382"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="SMART" id="SM00745"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033898.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="584" mass="63250" checksum="F6BFB61203009DB2" modified="2019-05-08" version="1">MNSPGGRGKKKGSAGSSSPPQAPGAFPSSPAGPAPPLQQQAAAGAAAASPHKRNLYYFSYPLFAAFALLRFVAFQLGLLFAWLCERLSRGALMAAKGRAGAGDAPEPGGAPETVRACHKRAFECISVALRIDEEERAGQKEQAVEWYKKGIEELEKGIAVLVIGQGDQCERARRLQSKMTTNLAMAKDRLQLLECGAVPKKKDPLTHTSNSLPRSKTVAVAKTGSTGLSGPHRTPSYSGISTFSVSRQGVNPAASTHKATSKNSRTNKPSTPTTAARKKKDLKIFRNVDSNLANLILNEIVDNGPAVKFDDIAGQELAKQALQEIVILPSLRPELFTGLRAPPRGLLLFGPPGNGKTMLAKAVAAESNATFFNISAASLTSKYVGEGEKLVRALFAVARELQPSIIFIDEVDSLLCERREGEHDASRRLKTEFLIEFDGVQSSGEDRILVMGATNRPQELDDAVLRRFTKRVYVSLPNEETRLILLKNLLSKQGSPLTQKELAQLARMTDGYSGSDLTALAKDAALGPIRELKPEQVKNMSASEMRNIRLSDFTESLKKIKRSLSPQTLDAYIRWNKDFGDTTV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0H9</accession>
    <name>A0A452J0H9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">G protein-coupled receptor kinase</fullName>
            <ecNumber evidence="11">2.7.11.-</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>[G-protein-coupled receptor] + ATP = [G-protein-coupled receptor]-phosphate + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:12008"/>
            <dbReference type="Rhea" id="RHEA-COMP:11260"/>
            <dbReference type="Rhea" id="RHEA-COMP:11261"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43176"/>
            <dbReference type="ChEBI" id="CHEBI:68546"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.16"/>
        </reaction>
    </comment>
    <comment type="similarity">
        <text evidence="2 11">Belongs to the protein kinase superfamily. AGC Ser/Thr protein kinase family. GPRK subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.-" evidence="11"/>
    <dbReference type="AlphaFoldDB" id="A0A452J0H9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038302.1">
        <property type="protein sequence ID" value="ENSGAGP00000033813.1"/>
        <property type="gene ID" value="ENSGAGG00000024060.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004703">
        <property type="term" value="F:G protein-coupled receptor kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009966">
        <property type="term" value="P:regulation of signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd05605">
        <property type="entry name" value="STKc_GRK4_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.167.10:FF:000009">
        <property type="entry name" value="G protein-coupled receptor kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000074">
        <property type="entry name" value="G protein-coupled receptor kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:001100">
        <property type="entry name" value="G protein-coupled receptor kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.167.10">
        <property type="entry name" value="Regulator of G-protein Signalling 4, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000961">
        <property type="entry name" value="AGC-kinase_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000239">
        <property type="entry name" value="GPCR_kinase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016137">
        <property type="entry name" value="RGS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036305">
        <property type="entry name" value="RGS_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044926">
        <property type="entry name" value="RGS_subdomain_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24355:SF26">
        <property type="entry name" value="G PROTEIN-COUPLED RECEPTOR KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24355">
        <property type="entry name" value="G PROTEIN-COUPLED RECEPTOR KINASE/RIBOSOMAL PROTEIN S6 KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00615">
        <property type="entry name" value="RGS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00717">
        <property type="entry name" value="GPCRKINASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00315">
        <property type="entry name" value="RGS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00133">
        <property type="entry name" value="S_TK_X"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48097">
        <property type="entry name" value="Regulator of G-protein signaling, RGS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51285">
        <property type="entry name" value="AGC_KINASE_CTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50132">
        <property type="entry name" value="RGS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 10 11" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="7 11" id="KW-0418">Kinase</keyword>
    <keyword evidence="6 10 11" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 11" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="5 11" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="RGS" evidence="13">
        <location>
            <begin position="64"/>
            <end position="176"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="12">
        <location>
            <begin position="191"/>
            <end position="453"/>
        </location>
    </feature>
    <feature type="domain" description="AGC-kinase C-terminal" evidence="14">
        <location>
            <begin position="454"/>
            <end position="519"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="9">
        <location>
            <position position="316"/>
        </location>
    </feature>
    <feature type="binding site" evidence="10">
        <location>
            <position position="229"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001256"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009793"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PIRSR" id="PIRSR600239-51"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="RuleBase" id="RU000308"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50132"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS51285"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033813.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="602" mass="69370" checksum="C9215EE78D0FC837" modified="2019-05-08" version="1">MVERCSLAVSRHWRSGLGLVLPGGGGKRKGRSKKWKEILRFPHISQCDDLRKGLEREYSSLCDKQPIGRLLFRQFCERRPELQRCIHFLDAVIDYELSSDERRKEMGDEIIRRFFKSESSDYMPEISQALMNQCMENLQKSACKDLFSNCLHPMHEYLSGAPFADYRDSMYFDRFLQWKYLERQSVTKDTFRQYRILGKGGFGEVCACQVRATGKMYACKKLEKKRIKKRKGEAMALNEKQILEKVNSRFVVSLAYAYETKDALCLVLTIMNGGDLKFHIYNMGNPGFEDERVIFYAAEICCGLQHLHQEGIVYRDLKPENILLDDDGHIRISDLGLAIKIPEGEMIRGRVGTVGYMAPEVINNERYTFSPDWWGLGCLIYEMIEGQSPFRARKERVKREEVEKRVQEEQELYSDKFTEDTKAICKMLLTKDPKQRLGCGEDGATQVKQHPFFRTINFKRLEAGVIKPSFVPDPRAVYCKDVLDIEQFSTVKGVNLDQTDSDFYAKFATGSVSIPWQNEMIETECFKDLNVFGPNGMRSPDLDWKQLPEPPKRSLLQRLFRRHPADYTISTNTHSNLTAGVNSHPPTANSVCQGRAMAQVPS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0I1</accession>
    <name>A0A452J0I1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Transmembrane protein 161A</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the TMEM161 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0I1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038397.1">
        <property type="protein sequence ID" value="ENSGAGP00000033903.1"/>
        <property type="gene ID" value="ENSGAGG00000024114.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019395">
        <property type="entry name" value="Transmembrane_161A/B"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13624">
        <property type="entry name" value="RE42071P"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13624:SF4">
        <property type="entry name" value="TRANSMEMBRANE PROTEIN 161A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10268">
        <property type="entry name" value="Tmemb_161AB"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6 9" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="3 9" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 9" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="102"/>
            <end position="123"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="135"/>
            <end position="156"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="168"/>
            <end position="184"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="226"/>
            <end position="248"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="268"/>
            <end position="288"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="326"/>
            <end position="351"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="415"/>
            <end position="437"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009706"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00040182"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033903.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="444" mass="50266" checksum="531D6B7A34E2171A" modified="2019-05-08" version="1">MGVQLVVSLLTASIMQKMSPHCSFARWLLCNGSLYRYKHPSEEELCALAGKQKPKAKRDRRPNGVAEDKPLSVPKDINLHLETTPVTTVDALVLRYFLEYQWFVDFAVYSTLIYLFTEGYYCLVDPQKEMNVGILWCLLTVVFSFKIFFMVMHHYFRSEEGGERSVCLTFAFFFLLIAMVVLVVREDYLEFGLDSGLANVSDNLEIFLKQQGWEWTVPITKLTFKLGLVALCSFIGACLTFPGLRLAQTHLDALKMAADRPMTHMSDSAYNSTRLWTIVALCLLRLAMTRHHLQAYLQLAERWVEQMKREAGRITALEIQRKVARIYCYITVVSLQYLGPIILTLHCTLLLKSLGDYSWGLFPEPTPVSPRVGAAPARPAVPSAEEEGGEDMQATVAQITGALGAILTPLFYRGLFAFLTWWVAVCQVITSLFGLYFHQYLAAS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0I5</accession>
    <name>A0A452J0I5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Tektin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">Sperm-specific microtubule inner protein (MIP) part of the dynein-decorated doublet microtubules (DMTs) in flagellar axoneme. Forms an extensive interaction network in different conformations that reinforces the helix bundle composed by other tektin proteins (TEKT1 to TEKT4) and MIPs to anchor the tektin bundle onto the tubulin wall of A-tubule of the sperm flagellum.</text>
    </comment>
    <comment type="subunit">
        <text evidence="11">Microtubule inner protein component of sperm flagellar doublet microtubules. Interacts with TEKT3.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="12">Cytoplasm</location>
            <location evidence="12">Cytoskeleton</location>
            <location evidence="12">Cilium axoneme</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Flagellum axoneme</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 12">Belongs to the tektin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0I5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038358.1">
        <property type="protein sequence ID" value="ENSGAGP00000033867.1"/>
        <property type="gene ID" value="ENSGAGG00000024073.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005930">
        <property type="term" value="C:axoneme"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015630">
        <property type="term" value="C:microtubule cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036126">
        <property type="term" value="C:sperm flagellum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060271">
        <property type="term" value="P:cilium assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060294">
        <property type="term" value="P:cilium movement involved in cell motility"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048256">
        <property type="entry name" value="Tektin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000435">
        <property type="entry name" value="Tektins"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19960">
        <property type="entry name" value="TEKTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19960:SF23">
        <property type="entry name" value="TEKTIN-5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03148">
        <property type="entry name" value="Tektin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00511">
        <property type="entry name" value="TEKTIN"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9 12" id="KW-0966">Cell projection</keyword>
    <keyword evidence="7 12" id="KW-0969">Cilium</keyword>
    <keyword evidence="6" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="5 12" id="KW-0282">Flagellum</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0832">Ubl conjugation</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004611"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007209"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022846"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023069"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00046061"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00047089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="RuleBase" id="RU367040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033867.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="457" mass="52696" checksum="E15B00F89F1CA179" modified="2019-05-08" version="1">MEFLGTTQTASYCGPKKSYLFPEPVSTSTTQDSYQSYYLPGYRYLNSWRPSLFYRVSAVPTNSGDQGSMRISAPLRPPTILPSVRSALYARYSPHDWYQANTVQYKGSESCRHWAGRMNADSLRLMQDKNQLTQQTQDSTGRSLGQRLSDIDFWRSELTYELERLLTETRALETAKRRLECAADEAQRPLKVGQRYLILNVLVCLRVQLVLCSINRDAQHALERDLSDKNSAHFIDEKCYNLRNTSDSISFYHGVEKIDGTISVPETWAKFSDDNIRHSQNARANSIKLCEETENTLENVSEEMWKQFTDTNLAFTNRIAETADTKNKLQTHLAKTLQEIFQTENTIMLLERAIMAKECPLKVAQTRLEGRTRRPNVELCRDIPQFKLVNEVFTIDDTLQTLKLRLREAHDTLQLLVLNKARLEHDLSVKANSLFIDRKCMDMRKTFPSTPRLVGYT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0I6</accession>
    <name>A0A452J0I6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Amino acid transporter</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 8">Membrane</location>
            <topology evidence="1 8">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="8">Belongs to the dicarboxylate/amino acid:cation symporter (DAACS) (TC 2.A.23) family.</text>
    </comment>
    <comment type="caution">
        <text evidence="8">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0I6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038402.1">
        <property type="protein sequence ID" value="ENSGAGP00000033908.1"/>
        <property type="gene ID" value="ENSGAGG00000024115.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015501">
        <property type="term" value="F:glutamate:sodium symporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005313">
        <property type="term" value="F:L-glutamate transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015175">
        <property type="term" value="F:neutral L-amino acid transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.3860.10">
        <property type="entry name" value="Sodium:dicarboxylate symporter"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050746">
        <property type="entry name" value="DAACS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001991">
        <property type="entry name" value="Na-dicarboxylate_symporter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018107">
        <property type="entry name" value="Na-dicarboxylate_symporter_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036458">
        <property type="entry name" value="Na:dicarbo_symporter_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11958:SF102">
        <property type="entry name" value="AMINO ACID TRANSPORTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11958">
        <property type="entry name" value="SODIUM/DICARBOXYLATE SYMPORTER-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00375">
        <property type="entry name" value="SDF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00173">
        <property type="entry name" value="EDTRNSPORT"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF118215">
        <property type="entry name" value="Proton glutamate symport protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00713">
        <property type="entry name" value="NA_DICARBOXYL_SYMP_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00714">
        <property type="entry name" value="NA_DICARBOXYL_SYMP_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6 8" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 8" id="KW-0769">Symport</keyword>
    <keyword evidence="3 8" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 8" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="2 8" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="133"/>
            <end position="153"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="173"/>
            <end position="194"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="206"/>
            <end position="232"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="244"/>
            <end position="269"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022847"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="RuleBase" id="RU361216"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033908.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="443" mass="47952" checksum="2F1939B4FCE6B037" modified="2019-05-08" version="1">VIMVPSVCICPLFTSCQMILLPDQSLFFSQEKQYFSFPGELLMRMLKMLILPLITSRNMFPSNLIEASFQQYRTVLVPVVKSSGFPKIPGKPLSFIYFAPDDENPEIQRPVLLEITPSPEMTYRILPGTSNEMNVLGIVIFSATIGLLLGKMGERGTPLVNVCQCLNEAVMKIVSMAVWYFPFGIVFLIAGKILEMEDPVVIGKKLGLYAITVVTGLVIHGLILLPLLFTLITRKNPFAFIRGILQALLIALATSSSSATLPITLKCLLENNGIDRRVARFVLPVGATINMDGTALYEAVAAIFIAQVNEYDLDLGQIITISITATAASIGAAGIPQSGLVTMVIVLTSVGLPTDDITLIIAVDWALDRLRTMTNVLGDALAAGIIAHVCQKDFAPKPEKVQASGNSPLEACNARQLPVSRESIWSGKIYCGGCCDASSQSNH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0K2</accession>
    <name>A0A452J0K2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">BTB domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J0K2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038417.1">
        <property type="protein sequence ID" value="ENSGAGP00000033923.1"/>
        <property type="gene ID" value="ENSGAGG00000024103.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
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        <property type="match status" value="1"/>
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        <property type="entry name" value="BTB1_POZ_LZTR1"/>
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        <property type="match status" value="1"/>
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        <property type="entry name" value="Leucine-zipper-like transcriptional regulator 1"/>
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        <property type="entry name" value="Kelch-type beta propeller"/>
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        <property type="entry name" value="Potassium Channel Kv1.1, Chain A"/>
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    <keyword evidence="1" id="KW-0880">Kelch repeat</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
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        <location>
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    </feature>
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            <begin position="499"/>
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    </evidence>
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        <source>
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    </evidence>
    <sequence length="672" mass="76646" checksum="77A226E530AEC7E1" modified="2019-05-08" version="1">KRISSEGASSMFLAPAVGSTQTKMTPHQDVLTSCSNFLLNDMWTIGLQDRELTCWEEIEQSGEIPPSCCNFPVAVCKDKMFVFSGQSGAKITNNLFQFEFKEKIWTRIPTEHLLRGSPPPPQRRYGHTMVAFDRHLYVFGGAADNTLPNELHCYDVDSQTWEVIQPSPDSEVAHPEVAERVAAPEEAPSFSSEERGLKKSRDVFGLDFGITTVKQQPTLPSGRLFHAAAVISDAMYIFGGTVDNNIRSGEMYRFQFSCYPKCTLHEDYGRLWENRQFSDLEFVLGEKEDRVRGHAAIVTARCKWLRKKIIQARERLKQATLWVPRSGICGNVKLCCLQPLLEVTIREAEAQPFEVLMQFLYTDKIKYPRKGHVQDVLLIMDVYRLALNFKLSRLEQLCLQYIEASVDLQNVLIVCENANKLQLEQLKEHCLNFVVKESHFNQVIMMKEFEHLSSSLIVEIVRRKQQPPVRTHSDQPVDIGTSLIQDMKAYLEGAGIEFCDIILLLDGHPRPAHKAILAARSSYFEAMFRSFMPEDGQVNISIGEMVPSKQAFESMLRYIYYGEVNMPPEDSLYLFAAPYYYGFYNNRLQAYCKQNLEMNVTVENVLQILEAADKTQALDMKKHCLHIIVHQFTKVSKLPNLRSLSQPLLLDIIESLANHISDKQCAELGSDI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0K3</accession>
    <name>A0A452J0K3_9SAUR</name>
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        <recommendedName>
            <fullName evidence="2">Coiled-coil domain-containing protein 22</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
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    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the CCDC22 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0K3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038379.1">
        <property type="protein sequence ID" value="ENSGAGP00000033887.1"/>
        <property type="gene ID" value="ENSGAGG00000024099.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097602">
        <property type="term" value="F:cullin family protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:2000060">
        <property type="term" value="P:positive regulation of ubiquitin-dependent protein catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="entry name" value="CCDC22"/>
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    <dbReference type="InterPro" id="IPR048348">
        <property type="entry name" value="CCDC22_CC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048349">
        <property type="entry name" value="CCDC22_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15668:SF4">
        <property type="entry name" value="COILED-COIL DOMAIN-CONTAINING PROTEIN 22"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15668">
        <property type="entry name" value="JM1 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05667">
        <property type="entry name" value="CCDC22_CC"/>
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        <property type="entry name" value="CCDC22_N"/>
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    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="CCDC22 N-terminal" evidence="5">
        <location>
            <begin position="1"/>
            <end position="109"/>
        </location>
    </feature>
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            <begin position="188"/>
            <end position="410"/>
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    </feature>
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            <begin position="158"/>
            <end position="187"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="415"/>
            <end position="445"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033887.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="475" mass="53092" checksum="2FC06CDA69C0646B" modified="2019-05-08" version="1">MDEVDKILIHSLRLSGTQVPEEVQSLQEFTTELIVESVVRCLRVIHPSLGTSLSHVLPPGMSARFRIGTSLAQACQELGYQGEVGYQTFLYSSEPEIRRLLLFLVEKLPRDDAEDARQPAGKSAGLLRAIAAKIKEELGTPWVPPACRTPRLQRLQVGPPPPQIQTHVSSQRALYPPHSRPSPRAGLRQQRLALGGREQTLQVKSRALELLPQAENNLAKLQLLVESSAQRLVHLAAQWETRRGPLMDQYRHLRAARHSCQLESTRRVSEIQALHERGRSAATEARRKEELCRQLLTELESLPKDVSRAAYTQRILEIVGNIRKQKEEITKILSATRALQKEINSLTGKLDRTFAVTDELVFKDAKRDESVRKAYKYLAALHENCSQLIQTIEDAGAIQREIRDLEEQVRVAGVGPGGWGAPRGPPSSSTEGVNMPPQIESEGAKKTLANLERILSDYRALRQENAALLGRSRDP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0K5</accession>
    <name>A0A452J0K5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">GRIP domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000038382.1">
        <property type="protein sequence ID" value="ENSGAGP00000033890.1"/>
        <property type="gene ID" value="ENSGAGG00000024101.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031267">
        <property type="term" value="F:small GTPase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048193">
        <property type="term" value="P:Golgi vesicle transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.220.60">
        <property type="entry name" value="GRIP domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000237">
        <property type="entry name" value="GRIP_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19327">
        <property type="entry name" value="GOLGIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19327:SF0">
        <property type="entry name" value="GOLGIN SUBFAMILY A MEMBER 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01465">
        <property type="entry name" value="GRIP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00755">
        <property type="entry name" value="Grip"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF101283">
        <property type="entry name" value="GRIP domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50913">
        <property type="entry name" value="GRIP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="GRIP" evidence="3">
        <location>
            <begin position="2237"/>
            <end position="2284"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1"/>
            <end position="162"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="292"/>
            <end position="330"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="165"/>
            <end position="213"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="334"/>
            <end position="708"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="764"/>
            <end position="1038"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="1073"/>
            <end position="1203"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="1364"/>
            <end position="1423"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="1465"/>
            <end position="1499"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="1532"/>
            <end position="1873"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="1909"/>
            <end position="2087"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="2118"/>
            <end position="2170"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="25"/>
            <end position="49"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="64"/>
            <end position="92"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="108"/>
            <end position="136"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="137"/>
            <end position="146"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="295"/>
            <end position="306"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS50913"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033890.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2299" mass="269520" checksum="42EEF586707148C3" modified="2019-05-08" version="1">MFKKLRQKIVEEQTQPRSPGGRAPQQPQTYSKSSPPSPGNRSRTSSLTEQSDEGTLTPDKENVSKQTTPKSTDSNGSEPPSPQLSDTQTFAQKLQLRVPSMESLFRSPVKESLFRSSSKESLVRTSSRESLNRLDFDTPGPTFDPPSDVESEAEESLGNVDSLGKEQLLQRLRRMERSLGSYRGKYSELVTAYQTVQREKKKLQGILSQSQDKALRRIGELREAVAVLKADIREGGFSYDIWSSSMKELQMDQQAKKHLQEEFDASLEEKDQLISVLQTQVSLLKQRLQNGQISTELPETNTETEPQSPAKEISAENVLEPGGTEGDGDSAKTLETLIKRVKRQENLLQRCKEMIRSHKERSAQLTNEKEALQEQLEERLQELEKMKELHMTEKTKLITQLRDAKNLIEQLEQDKGMVIAETKRQMHETMEMKEEEIAQLRAHIKQITTQGEELREQKEKSEKAAFEELEKALSTAQKVEEARKKLQAEMDEKIKAIEKTSEEERVTLQQELTRVKQEVVEIMKKSSEERVAELEKLHKRELATKEQELNERLQVQGKEFQEKMRAALEKSQSEFLQTVQEKEQQESLALEELELQKKAIQSECEKKLQDMQQEIQTARTRILELESSLAKYSQDDKRQLEELSTLIESEKTKYNKEIIVMVEKHREELENVKQQEEELWTEKLQILKQQQQSTIEKLREKHEQEMDIMLKEKETVFRAHVEEMNAKTLEKLDVKQTELEALSFELSETLQIRHDLEKDGDKIKQESETKLEQQRKQYQEKMDMMVKEHDISVQGIQKAHTEETNQLKVQLEEKEKHLEELKAQEHKLKESLEISEAELKQVSAKLGDLCQSHQNNSSEQARVYEGQLTELQQKLTNMEAEKSHLKEQIVRAESQLNEVRTELDTYISRVRDLRQQLEEQSSENMQKITSLKQKCESQLKDLSTETDQAKRDLLERENQLEQLKKQQNQQVEELRQKLSAKEERISALQVEYENSLKDQGTKMEKIKQKAKEMQEMFKRKLSEQEAKLKKEFENKQLEFSQKEKHFNAKMLEMAHASSTGINDAVSKLEMNQKEQLESLAEAHRWELEEVAQTWGKKLNQQVEELKEKHEMELHKKEKEVGDLKQKLATFSAEKEGTNIEITLLKEEQVKRDESWNELQEQLKQLLAQVNTLSQSETDLKAQLEKLEDDLSHSLKDRAVLQEQLREQKTMEEKHKIKVTELTDMLKTSDEKLQILESSHCKEREDYEKKLEDKHLASQQKQIEFERLSSELAAQLDIYWKEADTLLQTKTNEVIEKCTEKINLVISRIAHCQHQTTKIKKAILIKNCKIPELETQLRQITEHQTTVSSSLQESTQQLQDKENVIILMKADIERLVTEKELLQKEGGHQQQAATEKETCITQLRKELSENINVVTSMREELKEKESEICALNKLVNELNVKLENSIGLTEKEAAISLLSKQHQEDQLQLLNQVQELSSRVESLSQEKASALEQVDHWMNKLSEWKKKAQLRFTQNHNTIKELQNKLELNNIQVGGKDEEIHQLKEELDKQSRNLDSLKDVMEQKQRRREKEEGKLTAELKIQTARITELEEHIARKTAENDSLMEELKRHNQQRDTEQKELVQQLQQTEVVVVEKDNRLKEAEEKVLSLEKQMDSMKAELDTKQREFEQMKTAMMKSKEEELKMLEDRMNAENTTKLADLKKKAEQKIASVKKQLTSQMEEKEQQFKLAMENQLKDLEQKMQEREAKIMDLEEKNRLIRDSADLEREMMQKVENEKASLEQEKASVLENVRQMYEEKISEIQKDLIDKDGLLNKYENEQQESNDSKLKMKSKQEELLKKLECTEKKLKEEQSVIERLQKEIEEKNKKYSLLLDQHTQDGDDLASIREELKAQVQKHYDMENVIGDFQKQFQEKDKVSQSLEQKIRELEDDIVKKKEVCKIEMAEVSLKYEEKLQGLQQQLDERNRQLKIFEENAEEKTRSDLELQKLLGDMQTQQKDLQAKLKEAETEKQKLRKEISKVQKDLRTLRKEHQQELDIMKKESLEEMEQKIRIEQEDIELKHNSTLKQLMREFNTQLSQKERELETAVHETISKAQDVETELIETHHAETTQLHRKIAEKDDDLKRTVKKYEEILEAREEEMTAKVNELQTQLEELQKEYKQKLADEENRRREEVTITELQIQLAQKTTIVNDSKLKEQEFIEQIHILKDQLKNYEKNVYVTAVGSPYRDGNLCHTDVSLFGEPTEFEYLRKVLFEYMMGRETKTMAKVITTVLKFPADQTQKILEREDARPLSTSSRSGIF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0K6</accession>
    <name>A0A452J0K6_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="7">Spastin</fullName>
        </submittedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the AAA ATPase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0K6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038392.1">
        <property type="protein sequence ID" value="ENSGAGP00000033899.1"/>
        <property type="gene ID" value="ENSGAGG00000024111.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015630">
        <property type="term" value="C:microtubule cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000226">
        <property type="term" value="P:microtubule cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.8.60:FF:000022">
        <property type="entry name" value="Fidgetin like 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.8.60">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041569">
        <property type="entry name" value="AAA_lid_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050304">
        <property type="entry name" value="MT-severing_AAA_ATPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015415">
        <property type="entry name" value="Spast_Vps4_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23074">
        <property type="entry name" value="AAA DOMAIN-CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23074:SF86">
        <property type="entry name" value="SPASTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17862">
        <property type="entry name" value="AAA_lid_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09336">
        <property type="entry name" value="Vps4_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="4" id="KW-0472">Membrane</keyword>
    <keyword evidence="2" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="AAA ATPase AAA+ lid" evidence="6">
        <location>
            <begin position="93"/>
            <end position="121"/>
        </location>
    </feature>
    <feature type="domain" description="Spastin/Vps4 C-terminal" evidence="5">
        <location>
            <begin position="132"/>
            <end position="166"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006914"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="Pfam" id="PF09336"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="Pfam" id="PF17862"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033899.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="170" mass="19379" checksum="106028541B6F0F6E" modified="2019-05-08" version="1">MSLSRCNMLRMNCTHFGSYEKHHFFPVSLCKRVLFTLFSHHLDLLLKHADQHCAWGRETGNKGKLLPTQPLSKPSLSQLTKEITCSTFCVHFRMTDGYSGSDLTALAKDAALGPIRELKPEQVKNMSASEMRNIRLSDFTESLKKIKRSLSPQTLDAYIRWNKDFGDTTV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0K8</accession>
    <name>A0A452J0K8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">BTB domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J0K8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038423.1">
        <property type="protein sequence ID" value="ENSGAGP00000033928.1"/>
        <property type="gene ID" value="ENSGAGG00000024103.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006310">
        <property type="term" value="P:DNA recombination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd18505">
        <property type="entry name" value="BACK1_LZTR1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18506">
        <property type="entry name" value="BACK2_LZTR1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18308">
        <property type="entry name" value="BTB1_POZ_LZTR1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18309">
        <property type="entry name" value="BTB2_POZ_LZTR1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.710.10:FF:000024">
        <property type="entry name" value="Leucine-zipper-like transcriptional regulator 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.710.10:FF:000049">
        <property type="entry name" value="leucine-zipper-like transcriptional regulator 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.120.10.80">
        <property type="entry name" value="Kelch-type beta propeller"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.710.10">
        <property type="entry name" value="Potassium Channel Kv1.1, Chain A"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000210">
        <property type="entry name" value="BTB/POZ_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015915">
        <property type="entry name" value="Kelch-typ_b-propeller"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051568">
        <property type="entry name" value="LZTR1/Attractin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004321">
        <property type="entry name" value="RAG2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011333">
        <property type="entry name" value="SKP1/BTB/POZ_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46376">
        <property type="entry name" value="LEUCINE-ZIPPER-LIKE TRANSCRIPTIONAL REGULATOR 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46376:SF1">
        <property type="entry name" value="LEUCINE-ZIPPER-LIKE TRANSCRIPTIONAL REGULATOR 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00651">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24681">
        <property type="entry name" value="Kelch_KLHDC2_KLHL20_DRC7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03089">
        <property type="entry name" value="RAG2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00225">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF117281">
        <property type="entry name" value="Kelch motif"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54695">
        <property type="entry name" value="POZ domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50097">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0880">Kelch repeat</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="BTB" evidence="3">
        <location>
            <begin position="251"/>
            <end position="349"/>
        </location>
    </feature>
    <feature type="domain" description="BTB" evidence="3">
        <location>
            <begin position="479"/>
            <end position="548"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022441"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS50097"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033928.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="652" mass="74518" checksum="C5C0F29D28BFEE9D" modified="2019-05-08" version="1">KRISSEGASSMLNDMWTIGLQDRELTCWEEIEQSGEIPPSCCNFPVAVCKDKMFVFSGQSGAKITNNLFQFEFKEKIWTRIPTEHLLRGSPPPPQRRYGHTMVAFDRHLYVFGGAADNTLPNELHCYDVDSQTWEVIQPSPDSEVAHPEVAERVAAPEEAPSFSSEERGLKKSRDVFGLDFGITTVKQQPTLPSGRLFHAAAVISDAMYIFGGTVDNNIRSGEMYRFQFSCYPKCTLHEDYGRLWENRQFSDLEFVLGEKEDRVRGHAAIVTARCKWLRKKIIQARERLKQATNQPFSTPVPGHTQTFASLFTLCCLQPLLEVTIREAEAQPFEVLMQFLYTDKIKYPRKGHVQDVLLIMDVYRLALNFKLSRLEQLCLQYIEASVDLQNVLIVCENANKLQLEQLKEHCLNFVVKESHFNQVIMMKEFEHLSSSLIVEIVRRKQQPPVRTHSDQPVDIGTSLIQDMKAYLEGAGIEFCDIILLLDGHPRPAHKAILAARSSYFEAMFRSFMPEDGQVNISIGEMVPSKQAFESMLRYIYYGEVNMPPEDSLYLFAAPYYYGFYNNRLQAYCKQNLEMNVTVENVLQILEAADKTQALDMKKHCLHIIVHQFTKVSKLPNLRSLSQPLLLDIIESLANHISDKQCAELGSDI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0L1</accession>
    <name>A0A452J0L1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">GRIP domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000038390.1">
        <property type="protein sequence ID" value="ENSGAGP00000033897.1"/>
        <property type="gene ID" value="ENSGAGG00000024101.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031267">
        <property type="term" value="F:small GTPase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048193">
        <property type="term" value="P:Golgi vesicle transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.220.60">
        <property type="entry name" value="GRIP domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000237">
        <property type="entry name" value="GRIP_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19327">
        <property type="entry name" value="GOLGIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19327:SF0">
        <property type="entry name" value="GOLGIN SUBFAMILY A MEMBER 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01465">
        <property type="entry name" value="GRIP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00755">
        <property type="entry name" value="Grip"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF101283">
        <property type="entry name" value="GRIP domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50913">
        <property type="entry name" value="GRIP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="GRIP" evidence="3">
        <location>
            <begin position="2225"/>
            <end position="2272"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1"/>
            <end position="62"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="82"/>
            <end position="127"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="139"/>
            <end position="194"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="324"/>
            <end position="362"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="197"/>
            <end position="245"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="366"/>
            <end position="696"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="752"/>
            <end position="1026"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="1061"/>
            <end position="1191"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="1352"/>
            <end position="1411"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="1453"/>
            <end position="1487"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="1520"/>
            <end position="1861"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="1897"/>
            <end position="2075"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="2106"/>
            <end position="2158"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="25"/>
            <end position="49"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="89"/>
            <end position="124"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="140"/>
            <end position="168"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="169"/>
            <end position="178"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="327"/>
            <end position="338"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS50913"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033897.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2284" mass="267590" checksum="8E62026BBB556346" modified="2019-05-08" version="1">MFKKLRQKIVEEQTQPRSPGGRAPQQPQTYSKSSPPSPGNRSRTSSLTEQSDEGTLTPDKELLAGMIAEPAFLSEYTIFALDPKRPKPQSDSVNVSKQTTPKSTDSNGSEPPSPQLSDTQTFAQKLQLRVPSMESLFRSPVKESLFRSSSKESLVRTSSRESLNRLDFDTPGPTFDPPSDVESEAEESLGNVDSLGKEQLLQRLRRMERSLGSYRGKYSELVTAYQTVQREKKKLQGILSQSQDKALRRIGELREAVAVLKADIREGGFSYDIWSSSMKELQMDQQAKKHLQEEFDASLEEKDQLISVLQTQVSLLKQRLQNGQISTELPETNTETEPQSPAKEISAENVLEPGGTEGDGDSAKTLETLIKRVKRQENLLQRCKEMIRSHKERSAQLTNEKEALQEQLEERLQELEKMKELHMTEKTKLITQLRDAKNLIEQLEQDKGMVIAETKRQMHETMEMKEEEIAQLRAHIKQITTQGEELREQKEKSEKAAFEELEKALSTAQKVEEARKKLQAEMDEKIKAIEKTSEEERVTLQQELTRVKQEVVEIMKEKSQSEFLQTVQEKEQQESLALEELELQKKAIQSECEKKLQDMQQEIQTARTRILELESSLAKYSQDDKRQLEELSTLIESEKTKYNKEIIVMVEKHREELENVKQQEEELWTEKLQILKQQQQSTIEKLREKHEQEMDIMLKEKETVFRAHVEEMNAKTLEKLDVKQTELEALSFELSETLQIRHDLEKDGDKIKQESETKLEQQRKQYQEKMDMMVKEHDISVQGIQKAHTEETNQLKVQLEEKEKHLEELKAQEHKLKESLEISEAELKQVSAKLGDLCQSHQNNSSEQARVYEGQLTELQQKLTNMEAEKSHLKEQIVRAESQLNEVRTELDTYISRVRDLRQQLEEQSSENMQKITSLKQKCESQLKDLSTETDQAKRDLLERENQLEQLKKQQNQQVEELRQKLSAKEERISALQVEYENSLKDQGTKMEKIKQKAKEMQEMFKRKLSEQEAKLKKEFENKQLEFSQKEKHFNAKMLEMAHASSTGINDAVSKLEMNQKEQLESLAEAHRWELEEVAQTWGKKLNQQVEELKEKHEMELHKKEKEVGDLKQKLATFSAEKEGTNIEITLLKEEQVKRDESWNELQEQLKQLLAQVNTLSQSETDLKAQLEKLEDDLSHSLKDRAVLQEQLREQKTMEEKHKIKVTELTDMLKTSDEKLQILESSHCKEREDYEKKLEDKHLASQQKQIEFERLSSELAAQLDIYWKEADTLLQTKTNEVIEKCTEKINLVISRIAHCQHQTTKIKKAILIKNCKIPELETQLRQITEHQTTVSSSLQESTQQLQDKENVIILMKADIERLVTEKELLQKEGGHQQQAATEKETCITQLRKELSENINVVTSMREELKEKESEICALNKLVNELNVKLENSIGLTEKEAAISLLSKQHQEDQLQLLNQVQELSSRVESLSQEKASALEQVDHWMNKLSEWKKKAQLRFTQNHNTIKELQNKLELNNIQVGGKDEEIHQLKEELDKQSRNLDSLKDVMEQKQRRREKEEGKLTAELKIQTARITELEEHIARKTAENDSLMEELKRHNQQRDTEQKELVQQLQQTEVVVVEKDNRLKEAEEKVLSLEKQMDSMKAELDTKQREFEQMKTAMMKSKEEELKMLEDRMNAENTTKLADLKKKAEQKIASVKKQLTSQMEEKEQQFKLAMENQLKDLEQKMQEREAKIMDLEEKNRLIRDSADLEREMMQKVENEKASLEQEKASVLENVRQMYEEKISEIQKDLIDKDGLLNKYENEQQESNDSKLKMKSKQEELLKKLECTEKKLKEEQSVIERLQKEIEEKNKKYSLLLDQHTQDGDDLASIREELKAQVQKHYDMENVIGDFQKQFQEKDKVSQSLEQKIRELEDDIVKKKEVCKIEMAEVSLKYEEKLQGLQQQLDERNRQLKIFEENAEEKTRSDLELQKLLGDMQTQQKDLQAKLKEAETEKQKLRKEISKVQKDLRTLRKEHQQELDIMKKESLEEMEQKIRIEQEDIELKHNSTLKQLMREFNTQLSQKERELETAVHETISKAQDVETELIETHHAETTQLHRKIAEKDDDLKRTVKKYEEILEAREEEMTAKVNELQTQLEELQKEYKQKLADEENRRREEVTITELQIQLAQKTTIVNDSKLKEQEFIEQIHILKDQLKNYEKNVYVTAVGSPYRDGNLCHTDVSLFGEPTEFEYLRKVLFEYMMGRETKTMAKVITTVLKFPADQTQKILEREDARPLSWLRPS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0L2</accession>
    <name>A0A452J0L2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">BTB domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J0L2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038429.1">
        <property type="protein sequence ID" value="ENSGAGP00000033933.1"/>
        <property type="gene ID" value="ENSGAGG00000024103.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd18505">
        <property type="entry name" value="BACK1_LZTR1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18506">
        <property type="entry name" value="BACK2_LZTR1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18308">
        <property type="entry name" value="BTB1_POZ_LZTR1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18309">
        <property type="entry name" value="BTB2_POZ_LZTR1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.120.10.80:FF:000065">
        <property type="entry name" value="Leucine-zipper-like transcription regulator 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.710.10:FF:000024">
        <property type="entry name" value="Leucine-zipper-like transcriptional regulator 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.710.10:FF:000049">
        <property type="entry name" value="leucine-zipper-like transcriptional regulator 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.420">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.120.10.80">
        <property type="entry name" value="Kelch-type beta propeller"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.710.10">
        <property type="entry name" value="Potassium Channel Kv1.1, Chain A"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000210">
        <property type="entry name" value="BTB/POZ_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015915">
        <property type="entry name" value="Kelch-typ_b-propeller"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051568">
        <property type="entry name" value="LZTR1/Attractin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011333">
        <property type="entry name" value="SKP1/BTB/POZ_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46376">
        <property type="entry name" value="LEUCINE-ZIPPER-LIKE TRANSCRIPTIONAL REGULATOR 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46376:SF1">
        <property type="entry name" value="LEUCINE-ZIPPER-LIKE TRANSCRIPTIONAL REGULATOR 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00651">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24681">
        <property type="entry name" value="Kelch_KLHDC2_KLHL20_DRC7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00225">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF117281">
        <property type="entry name" value="Kelch motif"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54695">
        <property type="entry name" value="POZ domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50097">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0880">Kelch repeat</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="BTB" evidence="3">
        <location>
            <begin position="215"/>
            <end position="315"/>
        </location>
    </feature>
    <feature type="domain" description="BTB" evidence="3">
        <location>
            <begin position="445"/>
            <end position="514"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022441"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS50097"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033933.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="618" mass="71123" checksum="C81ACA53AC6DC704" modified="2019-05-08" version="1">FYDSVILSLGYDWFHLSLHAVVLNDMWTIGLQDRELTCWEEIEQSGEIPPSCCNFPVAVCKDKMFVFSGQSGAKITNNLFQFEFKEKIWTRIPTEHLLRGSPPPPQRRYGHTMVAFDRHLYVFGGAADNTLPNELHCYDVDSQTWEVIQPSPDSELPSGRLFHAAAVISDAMYIFGGTVDNNIRSGEMYRFQFSCYPKCTLHEDYGRLWENRQFSDLEFVLGEKEDRVRGHAAIVTARCKWLRKKIIQARERLKQKSKQEIEEEGQALSQNEGICGNVKLCCLQPLLEVTIREAEAQPFEVLMQFLYTDKIKYPRKGHVQDVLLIMDVYRLALNFKLSRLEQLCLQYIEASVDLQNVLIVCENANKLQLEQLKEHCLNFVVKESHFNQVIMMKEFEHLSSSLIVEIVRRKQQPPVRTHSDQPVDIGTSLIQDMKAYLEGAGIEFCDIILLLDGHPRPAHKAILAARSSYFEAMFRSFMPEDGQVNISIGEMVPSKQAFESMLRYIYYGEVNMPPEDSLYLFAAPYYYGFYNNRLQAYCKQNLEMNVTVENVLQILEAADKTQALDMKKHCLHIIVHQFTKVSKLPNLRSLSQPLLLDIIESLANHISDKQCAELGSDI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="35" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0L3</accession>
    <name>A0A452J0L3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Vascular endothelial growth factor receptor 3</fullName>
            <ecNumber evidence="4">2.7.10.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="36 37">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="37" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="36" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="36" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="25">
            <text>L-tyrosyl-[protein] + ATP = O-phospho-L-tyrosyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10596"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.10.1"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="30">Membrane</location>
            <topology evidence="30">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="30">Belongs to the protein kinase superfamily. Tyr protein kinase family. CSF-1/PDGF receptor subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.10.1" evidence="4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038398.1">
        <property type="protein sequence ID" value="ENSGAGP00000033904.1"/>
        <property type="gene ID" value="ENSGAGG00000024090.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043235">
        <property type="term" value="C:receptor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019838">
        <property type="term" value="F:growth factor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004714">
        <property type="term" value="F:transmembrane receptor protein tyrosine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001525">
        <property type="term" value="P:angiogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030335">
        <property type="term" value="P:positive regulation of cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043408">
        <property type="term" value="P:regulation of MAPK cascade"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048010">
        <property type="term" value="P:vascular endothelial growth factor receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05863">
        <property type="entry name" value="IgI_VEGFR-3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05102">
        <property type="entry name" value="PTKc_VEGFR3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000077">
        <property type="entry name" value="Vascular endothelial growth factor receptor 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000532">
        <property type="entry name" value="Vascular endothelial growth factor receptor 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000041">
        <property type="entry name" value="Vascular endothelial growth factor receptor 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000143">
        <property type="entry name" value="Vascular endothelial growth factor receptor 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000247">
        <property type="entry name" value="Vascular endothelial growth factor receptor 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000411">
        <property type="entry name" value="Vascular endothelial growth factor receptor 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000479">
        <property type="entry name" value="Vascular endothelial growth factor receptor 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000548">
        <property type="entry name" value="vascular endothelial growth factor receptor 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000949">
        <property type="entry name" value="vascular endothelial growth factor receptor 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013098">
        <property type="entry name" value="Ig_I-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050122">
        <property type="entry name" value="RTK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001245">
        <property type="entry name" value="Ser-Thr/Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008266">
        <property type="entry name" value="Tyr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020635">
        <property type="entry name" value="Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001824">
        <property type="entry name" value="Tyr_kinase_rcpt_3_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041348">
        <property type="entry name" value="VEGFR-2_TMD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055229">
        <property type="entry name" value="VEGFR1-3_5th"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055238">
        <property type="entry name" value="VEGFR1-3_N_Ig-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416">
        <property type="entry name" value="TYROSINE-PROTEIN KINASE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416:SF49">
        <property type="entry name" value="VASCULAR ENDOTHELIAL GROWTH FACTOR RECEPTOR 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07679">
        <property type="entry name" value="I-set"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13927">
        <property type="entry name" value="Ig_3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22971">
        <property type="entry name" value="Ig_VEGFR-1-like_5th"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07714">
        <property type="entry name" value="PK_Tyr_Ser-Thr"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21339">
        <property type="entry name" value="VEGFR-1-like_Ig-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17988">
        <property type="entry name" value="VEGFR-2_TMD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22854">
        <property type="entry name" value="VEGFR1-3_N_Ig-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000615">
        <property type="entry name" value="TyrPK_CSF1-R"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01832">
        <property type="entry name" value="VEGFRECEPTOR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01835">
        <property type="entry name" value="VEGFRECEPTR3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00219">
        <property type="entry name" value="TyrKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00109">
        <property type="entry name" value="PROTEIN_KINASE_TYR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00240">
        <property type="entry name" value="RECEPTOR_TYR_KIN_III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0037">Angiogenesis</keyword>
    <keyword evidence="16 27 29" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="20" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="22" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="24 30" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="15" id="KW-0418">Kinase</keyword>
    <keyword evidence="28" id="KW-0460">Magnesium</keyword>
    <keyword evidence="18 32" id="KW-0472">Membrane</keyword>
    <keyword evidence="28" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="14 27 29" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="23" id="KW-0539">Nucleus</keyword>
    <keyword evidence="8" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="21 30" id="KW-0675">Receptor</keyword>
    <keyword evidence="37" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="13" id="KW-0677">Repeat</keyword>
    <keyword evidence="12 33" id="KW-0732">Signal</keyword>
    <keyword evidence="10" id="KW-0808">Transferase</keyword>
    <keyword evidence="11 30 32" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="17 32" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="19" id="KW-0829">Tyrosine-protein kinase</keyword>
    <feature type="signal peptide" evidence="33">
        <location>
            <begin position="1"/>
            <end position="24"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019232544" description="Vascular endothelial growth factor receptor 3" evidence="33">
        <location>
            <begin position="25"/>
            <end position="1376"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="32">
        <location>
            <begin position="783"/>
            <end position="807"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="35">
        <location>
            <begin position="30"/>
            <end position="137"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="35">
        <location>
            <begin position="240"/>
            <end position="337"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="35">
        <location>
            <begin position="342"/>
            <end position="426"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="35">
        <location>
            <begin position="433"/>
            <end position="560"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="35">
        <location>
            <begin position="563"/>
            <end position="678"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="35">
        <location>
            <begin position="685"/>
            <end position="771"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="34">
        <location>
            <begin position="852"/>
            <end position="1179"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="31">
        <location>
            <begin position="1190"/>
            <end position="1218"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="31">
        <location>
            <begin position="1345"/>
            <end position="1376"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="26">
        <location>
            <position position="1043"/>
        </location>
    </feature>
    <feature type="binding site" evidence="27">
        <location>
            <begin position="859"/>
            <end position="866"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="27 29">
        <location>
            <position position="886"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="27">
        <location>
            <position position="1047"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="28">
        <location>
            <position position="1048"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="28">
        <location>
            <position position="1061"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00011902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022258"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022657"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023137"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00051243"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="PIRSR" id="PIRSR000615-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="PIRSR" id="PIRSR000615-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="PIRSR" id="PIRSR000615-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="RuleBase" id="RU000311"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="31">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="32">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="33">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="34">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="35">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="36">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033904.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="37">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1376" mass="156786" checksum="342E656DA48C5CBB" modified="2019-05-08" version="1">GVDQIYRISTLLLLFFLSPDLVRGYSLTPPTLSITEEEHTINSKDTLTVTCRGQHPLEWSWPGDLVDPDQGEKENETVVLVNLSDGRTIREEDCEGTNTKPYCKVLVLTGTQANDTGYYRCYYKYISAKIEGTTAISIYVFVRDFEQPFINRPETLLINKKENTWVPCLVSIPDLNITLFLPSSIIYPDGRRILWDNKKGMLVPTHLIRDSLFVQCETIIDKKLFRSNYFIVHVTGNELYDIQLFPKKAMELLVGEKLVLNCTVWAEFNAGVNFQWDYPGKQMHRAVTELPERRSQQTHTELSSILIIQNVSQQDLGKYMCMASNGEQTFKESIDVIVHEKPFINVEWRKGSVIEATAGDEVVKLPVKVVAYPPPEFQWYKDGKLIPNKQSPYSMQIKDVSEQNSGTYTLVLWNRLAGLEERISLQLIVNVPPHIHEKEASSPSIYSRRSHQALTCTVYGIPAPEVIQWQWRPWSPCRMFSRPRRHQRDRMPECKDWKDVSPQDAENPIESIDTWTESVEGRNKTVSKLVIQEANVSVMYKCVASNKVGRDERLIYFYVTTIPDGFEIESQPSEEPIEGQDLRLSCNADNYTYENLQWYRLNLSKLHHEEGNPLILDCKNVHHYATKMQGELHFKPDSNDAALLLTIPNISLREEGDYVCEVQNRKTREKHCHKKYISVQALEVPRLKQNLTDIWVNVSDSIEMRCKVDGAHIPDISWYKDEKLVEEVSGIDLADSNQRLSIQRVREEDAGLYLCSVCNAKGCVNSSASVSVEGSDDRTNVEIVILIGTGVIAVFFWILLILIFCNIKRPAHADIKTGYLSIIMDPGEVPLEEQCEYLPYDSSKWEFPRERLRLGKVLGHGAFGKVVEASAFGINKSNSCETVAVKMLKEGATASEHKALMSELKILIHIGNHLNVVNLLGACTKPNGPLMVIVEFCKYGNLSNYLRTKREGFSPYREKSPRLRIQVRSLVEAVRADRRSRFGTSDSAIFNRLLMNKSQTAQPIQEVDDLWQSPLTMEDLICYSFQVARGMEFLASRKCIHRDLAARNILLSENNVVKICDFGLARDIYKDPDYVRKGSARLPLKWMAPESIFDKVYTTQSDVWSFGVLLWEIFSLGASPYPGVQINEEFCQRLKDGTRMRAPEYATAEIYRIMLSCWHGDPKERPTFSDLVEILGDLLQENVQQEGKDYIPLNDSQSSEDDGFSQVPSSAQQNSDEEEFDMRLRCHSLAARYYNCVSFPGCLTGGNQIRCPSRIKTFEEFPMTHTTYKTHPDNQTDSGMVLASEEFERIENRHRKEGGFSSKGPSRNAELAVEQLAGVDLRDRCRPLYQSQVGGQTFYNSEYGELSEHSEDGSYTPPGEGASPSAVHASFFSEQY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0L4</accession>
    <name>A0A452J0L4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Tektin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">Sperm-specific microtubule inner protein (MIP) part of the dynein-decorated doublet microtubules (DMTs) in flagellar axoneme. Forms an extensive interaction network in different conformations that reinforces the helix bundle composed by other tektin proteins (TEKT1 to TEKT4) and MIPs to anchor the tektin bundle onto the tubulin wall of A-tubule of the sperm flagellum.</text>
    </comment>
    <comment type="subunit">
        <text evidence="11">Microtubule inner protein component of sperm flagellar doublet microtubules. Interacts with TEKT3.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="12">Cytoplasm</location>
            <location evidence="12">Cytoskeleton</location>
            <location evidence="12">Cilium axoneme</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Flagellum axoneme</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 12">Belongs to the tektin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0L4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038337.1">
        <property type="protein sequence ID" value="ENSGAGP00000033848.1"/>
        <property type="gene ID" value="ENSGAGG00000024073.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005930">
        <property type="term" value="C:axoneme"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015630">
        <property type="term" value="C:microtubule cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036126">
        <property type="term" value="C:sperm flagellum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060271">
        <property type="term" value="P:cilium assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060294">
        <property type="term" value="P:cilium movement involved in cell motility"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048256">
        <property type="entry name" value="Tektin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000435">
        <property type="entry name" value="Tektins"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19960">
        <property type="entry name" value="TEKTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19960:SF23">
        <property type="entry name" value="TEKTIN-5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03148">
        <property type="entry name" value="Tektin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00511">
        <property type="entry name" value="TEKTIN"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9 12" id="KW-0966">Cell projection</keyword>
    <keyword evidence="7 12" id="KW-0969">Cilium</keyword>
    <keyword evidence="6" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="5 12" id="KW-0282">Flagellum</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0832">Ubl conjugation</keyword>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004611"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007209"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022846"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023069"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00046061"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00047089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="RuleBase" id="RU367040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033848.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="353" mass="40768" checksum="FE48C0E7B19E840E" modified="2019-05-08" version="1">MRMISAVPTNSGDQGSMRISAPLRPPTILPSVRSALYARYSPHDWYQANTVQYKGSESCRHWAGRMNADSLRLMQDKNQLTQQTQDSTGRSLGQRLSDIDFWRSELTYELERLLTETRALETAKRRLECAADEAQRPLKVGQRTSFFSSQFKLFSSRISVPETWAKFSDDNIRHSQNARANSIKLCEETENTLENVSEEMWKQFTDTNLAFTNRIAETADTKNKLQTHLAKTLQEIFQTENTIMLLERAIMAKECPLKVAQTRLEGRTRRPNVELCRDIPQFKLVNEVFTIDDTLQTLKLRLREAHDTLQLLVLNKARLEHDLSVKANSLFIDRKCMDMRKTFPSTPRLVGYT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0L5</accession>
    <name>A0A452J0L5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Transmembrane protein 161A</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the TMEM161 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0L5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038394.1">
        <property type="protein sequence ID" value="ENSGAGP00000033900.1"/>
        <property type="gene ID" value="ENSGAGG00000024114.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019395">
        <property type="entry name" value="Transmembrane_161A/B"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13624">
        <property type="entry name" value="RE42071P"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13624:SF4">
        <property type="entry name" value="TRANSMEMBRANE PROTEIN 161A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10268">
        <property type="entry name" value="Tmemb_161AB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6 9" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="3 9" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 9" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="133"/>
            <end position="153"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="165"/>
            <end position="184"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="222"/>
            <end position="244"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="265"/>
            <end position="286"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="364"/>
            <end position="389"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="434"/>
            <end position="457"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009706"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00040182"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033900.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="458" mass="51703" checksum="2DE0A63DAF95FFFD" modified="2019-05-08" version="1">MGVQLVVSLLTASIMQKMSPHCSFARWLLCNGSLYRYKHPSEEELCALAGKQKPKAKRDRRPNGVAEDKPLSVPKDINLHLETTPVTTVDALVLRYFLEYQWFVDFAVYSTLIYLFTEGYYCLVDPQKEMNVGILWCLLTVVFSFKIFFMVMHHYFRSEEGGERSVCLTFAFFFLLIAMVVLVVREDYLEFGLDSGLANVSDNLEIFLKQQGWEWTVPITKLTFKLGLVALCSFIGACLTFPGLRLAQTHLDALKMAADRPMTQILLHASFLAPLVVVLMWVKPISRDLLLHAPMGKQTVQIMSDSAYNSTRLWTIVALCLLRLAMTRHHLQAYLQLAERWVEQMKREAGRITALEIQRKVARIYCYITVVSLQYLGPIILTLHCTLLLKSLGDYSWGLFPEPTPVSPRVGAAPARPAVPSAEEEGGEDMQATVAQITGALGAILTPLFYRGLFAFLT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="35" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0L7</accession>
    <name>A0A452J0L7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Vascular endothelial growth factor receptor 3</fullName>
            <ecNumber evidence="4">2.7.10.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="35 36">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="36" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="35" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="35" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="25">
            <text>L-tyrosyl-[protein] + ATP = O-phospho-L-tyrosyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10596"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.10.1"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="30">Membrane</location>
            <topology evidence="30">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="30">Belongs to the protein kinase superfamily. Tyr protein kinase family. CSF-1/PDGF receptor subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.10.1" evidence="4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038403.1">
        <property type="protein sequence ID" value="ENSGAGP00000033909.1"/>
        <property type="gene ID" value="ENSGAGG00000024090.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043235">
        <property type="term" value="C:receptor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019838">
        <property type="term" value="F:growth factor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004714">
        <property type="term" value="F:transmembrane receptor protein tyrosine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001525">
        <property type="term" value="P:angiogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030335">
        <property type="term" value="P:positive regulation of cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043408">
        <property type="term" value="P:regulation of MAPK cascade"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048010">
        <property type="term" value="P:vascular endothelial growth factor receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05863">
        <property type="entry name" value="IgI_VEGFR-3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000077">
        <property type="entry name" value="Vascular endothelial growth factor receptor 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000532">
        <property type="entry name" value="Vascular endothelial growth factor receptor 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000041">
        <property type="entry name" value="Vascular endothelial growth factor receptor 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000143">
        <property type="entry name" value="Vascular endothelial growth factor receptor 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000247">
        <property type="entry name" value="Vascular endothelial growth factor receptor 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000411">
        <property type="entry name" value="Vascular endothelial growth factor receptor 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000479">
        <property type="entry name" value="Vascular endothelial growth factor receptor 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000548">
        <property type="entry name" value="vascular endothelial growth factor receptor 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000949">
        <property type="entry name" value="vascular endothelial growth factor receptor 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013098">
        <property type="entry name" value="Ig_I-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050122">
        <property type="entry name" value="RTK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001245">
        <property type="entry name" value="Ser-Thr/Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008266">
        <property type="entry name" value="Tyr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020635">
        <property type="entry name" value="Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001824">
        <property type="entry name" value="Tyr_kinase_rcpt_3_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041348">
        <property type="entry name" value="VEGFR-2_TMD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055229">
        <property type="entry name" value="VEGFR1-3_5th"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055238">
        <property type="entry name" value="VEGFR1-3_N_Ig-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416">
        <property type="entry name" value="TYROSINE-PROTEIN KINASE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416:SF49">
        <property type="entry name" value="VASCULAR ENDOTHELIAL GROWTH FACTOR RECEPTOR 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07679">
        <property type="entry name" value="I-set"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13927">
        <property type="entry name" value="Ig_3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22971">
        <property type="entry name" value="Ig_VEGFR-1-like_5th"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07714">
        <property type="entry name" value="PK_Tyr_Ser-Thr"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21339">
        <property type="entry name" value="VEGFR-1-like_Ig-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17988">
        <property type="entry name" value="VEGFR-2_TMD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22854">
        <property type="entry name" value="VEGFR1-3_N_Ig-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000615">
        <property type="entry name" value="TyrPK_CSF1-R"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01832">
        <property type="entry name" value="VEGFRECEPTOR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01835">
        <property type="entry name" value="VEGFRECEPTR3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00219">
        <property type="entry name" value="TyrKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00109">
        <property type="entry name" value="PROTEIN_KINASE_TYR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00240">
        <property type="entry name" value="RECEPTOR_TYR_KIN_III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0037">Angiogenesis</keyword>
    <keyword evidence="16 27 29" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="20" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="22" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="24 30" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="15" id="KW-0418">Kinase</keyword>
    <keyword evidence="28" id="KW-0460">Magnesium</keyword>
    <keyword evidence="18 32" id="KW-0472">Membrane</keyword>
    <keyword evidence="28" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="14 27 29" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="23" id="KW-0539">Nucleus</keyword>
    <keyword evidence="8" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="21 30" id="KW-0675">Receptor</keyword>
    <keyword evidence="36" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="13" id="KW-0677">Repeat</keyword>
    <keyword evidence="12" id="KW-0732">Signal</keyword>
    <keyword evidence="10" id="KW-0808">Transferase</keyword>
    <keyword evidence="11 30 32" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="17 32" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="19" id="KW-0829">Tyrosine-protein kinase</keyword>
    <feature type="transmembrane region" description="Helical" evidence="32">
        <location>
            <begin position="764"/>
            <end position="788"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="34">
        <location>
            <begin position="17"/>
            <end position="124"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="34">
        <location>
            <begin position="227"/>
            <end position="317"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="34">
        <location>
            <begin position="322"/>
            <end position="406"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="34">
        <location>
            <begin position="413"/>
            <end position="541"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="34">
        <location>
            <begin position="544"/>
            <end position="659"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="34">
        <location>
            <begin position="666"/>
            <end position="752"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="33">
        <location>
            <begin position="833"/>
            <end position="1145"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="31">
        <location>
            <begin position="1156"/>
            <end position="1184"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="31">
        <location>
            <begin position="1311"/>
            <end position="1342"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="26">
        <location>
            <position position="1009"/>
        </location>
    </feature>
    <feature type="binding site" evidence="27">
        <location>
            <begin position="840"/>
            <end position="847"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="27 29">
        <location>
            <position position="867"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="27">
        <location>
            <position position="1013"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="28">
        <location>
            <position position="1014"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="28">
        <location>
            <position position="1027"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00011902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022258"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022657"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023137"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00051243"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="PIRSR" id="PIRSR000615-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="PIRSR" id="PIRSR000615-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="PIRSR" id="PIRSR000615-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="RuleBase" id="RU000311"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="31">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="32">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="33">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="34">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="35">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033909.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="36">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1342" mass="152836" checksum="1EB60E5214989548" modified="2019-05-08" version="1">LILSPGLICSRYSLTPPTLSITEEEHTINSKDTLTVTCRGQHPLEWSWPGDLVDPDQGEKENETVVLVNLSDGRTIREEDCEGTNTKPYCKVLVLTGTQANDTGYYRCYYKYISAKIEGTTAISIYVFVRDFEQPFINRPETLLINKKENTWVPCLVSIPDLNITLFLPSSIIYPDGRRILWDNKKGMLVPTHLIRDSLFVQCETIIDKKLFRSNYFIVHVTGNELYDIQLFPKKAMELLVGEKLVLNCTVWAEFNAGVNFQWDYPGKQLPERRSQQTHTELSSILIIQNVSQQDLGKYMCMASNGEQTFKESIDVIVHEKPFINVEWRKGSVIEATAGDEVVKLPVKVVAYPPPEFQWYKDGKLIPNKQSPYSMQIKDVSEQNSGTYTLVLWNRLAGLEERISLQLIVNVPPHIHEKEASSPSIYSRRSHQALTCTVYGIPAPEVIQWQWRPWSPCRMFSRRSLRHQRDRMPECKDWKDVSPQDAENPIESIDTWTESVEGRNKTVSKLVIQEANVSVMYKCVASNKVGRDERLIYFYVTTIPDGFEIESQPSEEPIEGQDLRLSCNADNYTYENLQWYRLNLSKLHHEEGNPLILDCKNVHHYATKMQGELHFKPDSNDAALLLTIPNISLREEGDYVCEVQNRKTREKHCHKKYISVQALEVPRLKQNLTDIWVNVSDSIEMRCKVDGAHIPDISWYKDEKLVEEVSGIDLADSNQRLSIQRVREEDAGLYLCSVCNAKGCVNSSASVSVEGSDDRTNVEIVILIGTGVIAVFFWILLILIFCNIKRPAHADIKTGYLSIIMDPGEVPLEEQCEYLPYDSSKWEFPRERLRLGKVLGHGAFGKVVEASAFGINKSNSCETVAVKMLKEGATASEHKALMSELKILIHIGNHLNVVNLLGACTKPNGPLMVIVEFCKYGNLSNYLRTKREGFSPYRVCSLLHLVSSLRAFPMISERRRISLITFFLSLWTVDDLWQSPLTMEDLICYSFQVARGMEFLASRKCIHRDLAARNILLSENNVVKICDFGLARDIYKDPDYVRKGSARLPLKWMAPESIFDKVYTTQSDVWSFGVLLWEIFSLGASPYPGVQINEEFCQRLKDGTRMRAPEYATAEIYRIMLSCWHGDPKERPTFSDLVEILGDLLQENVQQEGKDYIPLNDSQSSEDDGFSQVPSSAQQNSDEEEFDMRLRCHSLAARYYNCVSFPGCLTGGNQIRCPSRIKTFEEFPMTHTTYKTHPDNQTDSGMVLASEEFERIENRHRKEGGFSSKGPSRNAELAVEQLAGVDLRDRCRPLYQSQVGGQTFYNSEYGELSEHSEDGSYTPPGEGASPSAVHASFFSEQY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="35" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0L8</accession>
    <name>A0A452J0L8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Vascular endothelial growth factor receptor 3</fullName>
            <ecNumber evidence="4">2.7.10.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="35 36">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="36" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="35" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="35" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="25">
            <text>L-tyrosyl-[protein] + ATP = O-phospho-L-tyrosyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10596"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.10.1"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="30">Membrane</location>
            <topology evidence="30">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="30">Belongs to the protein kinase superfamily. Tyr protein kinase family. CSF-1/PDGF receptor subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.10.1" evidence="4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038396.1">
        <property type="protein sequence ID" value="ENSGAGP00000033902.1"/>
        <property type="gene ID" value="ENSGAGG00000024090.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043235">
        <property type="term" value="C:receptor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019838">
        <property type="term" value="F:growth factor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004714">
        <property type="term" value="F:transmembrane receptor protein tyrosine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001525">
        <property type="term" value="P:angiogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030335">
        <property type="term" value="P:positive regulation of cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043408">
        <property type="term" value="P:regulation of MAPK cascade"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048010">
        <property type="term" value="P:vascular endothelial growth factor receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05863">
        <property type="entry name" value="IgI_VEGFR-3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000077">
        <property type="entry name" value="Vascular endothelial growth factor receptor 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000532">
        <property type="entry name" value="Vascular endothelial growth factor receptor 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000041">
        <property type="entry name" value="Vascular endothelial growth factor receptor 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000143">
        <property type="entry name" value="Vascular endothelial growth factor receptor 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000247">
        <property type="entry name" value="Vascular endothelial growth factor receptor 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000411">
        <property type="entry name" value="Vascular endothelial growth factor receptor 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000479">
        <property type="entry name" value="Vascular endothelial growth factor receptor 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000548">
        <property type="entry name" value="vascular endothelial growth factor receptor 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000949">
        <property type="entry name" value="vascular endothelial growth factor receptor 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013098">
        <property type="entry name" value="Ig_I-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050122">
        <property type="entry name" value="RTK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001245">
        <property type="entry name" value="Ser-Thr/Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008266">
        <property type="entry name" value="Tyr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020635">
        <property type="entry name" value="Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001824">
        <property type="entry name" value="Tyr_kinase_rcpt_3_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041348">
        <property type="entry name" value="VEGFR-2_TMD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055229">
        <property type="entry name" value="VEGFR1-3_5th"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055238">
        <property type="entry name" value="VEGFR1-3_N_Ig-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416">
        <property type="entry name" value="TYROSINE-PROTEIN KINASE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416:SF49">
        <property type="entry name" value="VASCULAR ENDOTHELIAL GROWTH FACTOR RECEPTOR 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07679">
        <property type="entry name" value="I-set"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13927">
        <property type="entry name" value="Ig_3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22971">
        <property type="entry name" value="Ig_VEGFR-1-like_5th"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07714">
        <property type="entry name" value="PK_Tyr_Ser-Thr"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21339">
        <property type="entry name" value="VEGFR-1-like_Ig-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17988">
        <property type="entry name" value="VEGFR-2_TMD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22854">
        <property type="entry name" value="VEGFR1-3_N_Ig-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000615">
        <property type="entry name" value="TyrPK_CSF1-R"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00109">
        <property type="entry name" value="TYRKINASE"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01832">
        <property type="entry name" value="VEGFRECEPTOR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01835">
        <property type="entry name" value="VEGFRECEPTR3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00219">
        <property type="entry name" value="TyrKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00109">
        <property type="entry name" value="PROTEIN_KINASE_TYR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00240">
        <property type="entry name" value="RECEPTOR_TYR_KIN_III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0037">Angiogenesis</keyword>
    <keyword evidence="16 27 29" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="20" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="22" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="24 30" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="15" id="KW-0418">Kinase</keyword>
    <keyword evidence="28" id="KW-0460">Magnesium</keyword>
    <keyword evidence="18 32" id="KW-0472">Membrane</keyword>
    <keyword evidence="28" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="14 27 29" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="23" id="KW-0539">Nucleus</keyword>
    <keyword evidence="8" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="21 30" id="KW-0675">Receptor</keyword>
    <keyword evidence="36" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="13" id="KW-0677">Repeat</keyword>
    <keyword evidence="12" id="KW-0732">Signal</keyword>
    <keyword evidence="10" id="KW-0808">Transferase</keyword>
    <keyword evidence="11 30 32" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="17 32" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="19" id="KW-0829">Tyrosine-protein kinase</keyword>
    <feature type="transmembrane region" description="Helical" evidence="32">
        <location>
            <begin position="772"/>
            <end position="796"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="34">
        <location>
            <begin position="26"/>
            <end position="133"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="34">
        <location>
            <begin position="236"/>
            <end position="326"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="34">
        <location>
            <begin position="331"/>
            <end position="415"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="34">
        <location>
            <begin position="422"/>
            <end position="549"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="34">
        <location>
            <begin position="552"/>
            <end position="667"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="34">
        <location>
            <begin position="674"/>
            <end position="760"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="33">
        <location>
            <begin position="841"/>
            <end position="1160"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="31">
        <location>
            <begin position="1171"/>
            <end position="1199"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="31">
        <location>
            <begin position="1326"/>
            <end position="1357"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="26">
        <location>
            <position position="1024"/>
        </location>
    </feature>
    <feature type="binding site" evidence="27">
        <location>
            <begin position="848"/>
            <end position="855"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="27 29">
        <location>
            <position position="875"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="27">
        <location>
            <position position="1028"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="28">
        <location>
            <position position="1029"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="28">
        <location>
            <position position="1042"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00011902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022258"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022657"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023137"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00051243"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="PIRSR" id="PIRSR000615-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="PIRSR" id="PIRSR000615-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="PIRSR" id="PIRSR000615-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="RuleBase" id="RU000311"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="31">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="32">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="33">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="34">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="35">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033902.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="36">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1357" mass="154447" checksum="394A5BAFB11C6511" modified="2019-05-08" version="1">NERGGRQCGRVCVCLYLVRGYSLTPPTLSITEEEHTINSKDTLTVTCRGQHPLEWSWPGDLVDPDQGEKENETVVLVNLSDGRTIREEDCEGTNTKPYCKVLVLTGTQANDTGYYRCYYKYISAKIEGTTAISIYVFVRDFEQPFINRPETLLINKKENTWVPCLVSIPDLNITLFLPSSIIYPDGRRILWDNKKGMLVPTHLIRDSLFVQCETIIDKKLFRSNYFIVHVTGNELYDIQLFPKKAMELLVGEKLVLNCTVWAEFNAGVNFQWDYPGKQLPERRSQQTHTELSSILIIQNVSQQDLGKYMCMASNGEQTFKESIDVIVHEKPFINVEWRKGSVIEATAGDEVVKLPVKVVAYPPPEFQWYKDGKLIPNKQSPYSMQIKDVSEQNSGTYTLVLWNRLAGLEERISLQLIVNVPPHIHEKEASSPSIYSRRSHQALTCTVYGIPAPEVIQWQWRPWSPCRMFSRPRRHQRDRMPECKDWKDVSPQDAENPIESIDTWTESVEGRNKTVSKLVIQEANVSVMYKCVASNKVGRDERLIYFYVTTIPDGFEIESQPSEEPIEGQDLRLSCNADNYTYENLQWYRLNLSKLHHEEGNPLILDCKNVHHYATKMQGELHFKPDSNDAALLLTIPNISLREEGDYVCEVQNRKTREKHCHKKYISVQALEVPRLKQNLTDIWVNVSDSIEMRCKVDGAHIPDISWYKDEKLVEEVSGIDLADSNQRLSIQRVREEDAGLYLCSVCNAKGCVNSSASVSVEGSDDRTNVEIVILIGTGVIAVFFWILLILIFCNIKRPAHADIKTGYLSIIMDPGEVPLEEQCEYLPYDSSKWEFPRERLRLGKVLGHGAFGKVVEASAFGINKSNSCETVAVKMLKEGATASEHKALMSELKILIHIGNHLNVVNLLGACTKPNGKYARPPRQLAVKRKALSPPCWEKSPRLRIQVRSLVEAVRADRRSRFGTSDSAIFNRLLMNKSQTAQPIQEVDDLWQSPLTMEDLICYSFQVARGMEFLASRKCIHRDLAARNILLSENNVVKICDFGLARDIYKDPDYVRKGSARLPLKWMAPESIFDKVYTTQSDVWSFGVLLWEIFSLGASPYPGVQINEEFCQRLKDGTRMRAPEYATAEIYRIMLSCWHGDPKERPTFSDLVEILGDLLQENVQQEGKDYIPLNDSQSSEDDGFSQVPSSAQQNSDEEEFDMRLRCHSLAARYYNCVSFPGCLTGGNQIRCPSRIKTFEEFPMTHTTYKTHPDNQTDSGMVLASEEFERIENRHRKEGGFSSKGPSRNAELAVEQLAGVDLRDRCRPLYQSQVGGQTFYNSEYGELSEHSEDGSYTPPGEGASPSAVHASFFSEQY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0M1</accession>
    <name>A0A452J0M1_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="13">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
            <location evidence="2">Cilium axoneme</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Cilium basal body</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="8">Belongs to the TRAF3IP1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0M1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038343.1">
        <property type="protein sequence ID" value="ENSGAGP00000033853.1"/>
        <property type="gene ID" value="ENSGAGG00000024079.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005930">
        <property type="term" value="C:axoneme"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036064">
        <property type="term" value="C:ciliary basal body"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030992">
        <property type="term" value="C:intraciliary transport particle B"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008017">
        <property type="term" value="F:microtubule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060271">
        <property type="term" value="P:cilium assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042073">
        <property type="term" value="P:intraciliary transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070507">
        <property type="term" value="P:regulation of microtubule cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.418.50">
        <property type="entry name" value="Microtubule-binding protein MIP-T3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018799">
        <property type="entry name" value="TRAF3IP1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041476">
        <property type="entry name" value="TRAF3IP1_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040468">
        <property type="entry name" value="TRAF3IP1_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042576">
        <property type="entry name" value="TRAF3IP1_N_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31363">
        <property type="entry name" value="TRAF3-INTERACTING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31363:SF0">
        <property type="entry name" value="TRAF3-INTERACTING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10243">
        <property type="entry name" value="MIP-T3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17749">
        <property type="entry name" value="MIP-T3_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0966">Cell projection</keyword>
    <keyword evidence="4" id="KW-0970">Cilium biogenesis/degradation</keyword>
    <keyword evidence="5 9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="TRAF3-interacting protein 1 N-terminal" evidence="11">
        <location>
            <begin position="1"/>
            <end position="68"/>
        </location>
    </feature>
    <feature type="domain" description="TRAF3-interacting protein 1 C-terminal" evidence="12">
        <location>
            <begin position="479"/>
            <end position="630"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="83"/>
            <end position="452"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="473"/>
            <end position="519"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="549"/>
            <end position="608"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="98"/>
            <end position="304"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="343"/>
            <end position="365"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="479"/>
            <end position="497"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="504"/>
            <end position="519"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004120"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004430"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022794"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00043971"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF10243"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF17749"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033853.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="636" mass="72807" checksum="C4FA9F3AB3599A3C" modified="2019-05-08" version="1">MNGLYTDFEMKSDNVKDKDAKISFLQKAIDVVIMVTGEPLSVKPARIVAGHEPERTNEFLQAIGKCCLNKLSSDDAVKRVLAGEKADIKGKPPSTSKSQDKENRESRAEEQKSHKDKEGRGDNEIKDRSTSRDRKPREELKEEEKKQKEKERDKHKDKEDRHKDLEADNFREGEKHEREKSKNRTSKQGRETEKSRVKEKREMEREKDLEHDKGQEKERKHEGGKEKEKLKERDKEKGRDKDRERDKDRGKERERDRRRERGKDGERLKEQGTDKAEKKSTGSEDTLPKKTERSSKDTKTEPGKESGSPARIPRQFSTKGPRQRVKPGAEGILETNSPADSVLDEKAATILHAKAEPRPAIKHQGESACDAEGEAGNRASEKPIVSENDEVSNDLPPHVTQRRPPRPNSARPAPPRIKRQESTEVLLPERNGSGKAVSNVIIDKQNSDDEDDQFVVEAALQLPEMPEMETEPIVELDGDEKHGGLVKRILETKRDYETSQQSKSTEKEKPLLSEAARRKEKDLVSKEIEKFRASIQTLCRSALPLGKIMDYIQEDMDAMKNELQMWQHENKQHAEALQKEQSITDSAVEPLKAELAELEQLIKDQQDKICTVKANILKKEEKIRKMVLNINLSTRR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0N0</accession>
    <name>A0A452J0N0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">Myosin motor domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">Muscle contraction.</text>
    </comment>
    <comment type="subunit">
        <text evidence="14">Muscle myosin is a hexameric protein that consists of 2 heavy chain subunits (MHC), 2 alkali light chain subunits (MLC) and 2 regulatory light chain subunits (MLC-2).</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Myofibril</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 15">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038453.1">
        <property type="protein sequence ID" value="ENSGAGP00000033955.1"/>
        <property type="gene ID" value="ENSGAGG00000020688.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030016">
        <property type="term" value="C:myofibril"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032982">
        <property type="term" value="C:myosin filament"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016460">
        <property type="term" value="C:myosin II complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000146">
        <property type="term" value="F:microfilament motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000003">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000004">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000013">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000007">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000008">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.4820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.530:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000024">
        <property type="entry name" value="Myosin heavy chain, isoform J"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.720:FF:000001">
        <property type="entry name" value="Myosin heavy chain, muscle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.340">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.370">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.4820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.2420">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002928">
        <property type="entry name" value="Myosin_tail"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014751">
        <property type="entry name" value="XRCC4-like_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615">
        <property type="entry name" value="MYOSIN HEAVY CHAIN, NON-MUSCLE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615:SF69">
        <property type="entry name" value="MYOSIN-6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01576">
        <property type="entry name" value="Myosin_tail_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90257">
        <property type="entry name" value="Myosin rod fragments"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57997">
        <property type="entry name" value="Tropomyosin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12 15" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="7 15" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="8 16" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0488">Methylation</keyword>
    <keyword evidence="10 15" id="KW-0505">Motor protein</keyword>
    <keyword evidence="11" id="KW-0514">Muscle protein</keyword>
    <keyword evidence="9 15" id="KW-0518">Myosin</keyword>
    <keyword evidence="6 15" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0787">Thick filament</keyword>
    <feature type="domain" description="Myosin motor" evidence="17">
        <location>
            <begin position="77"/>
            <end position="769"/>
        </location>
    </feature>
    <feature type="region of interest" description="Actin-binding" evidence="15">
        <location>
            <begin position="646"/>
            <end position="668"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="16">
        <location>
            <begin position="831"/>
            <end position="1127"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="16">
        <location>
            <begin position="1171"/>
            <end position="1894"/>
        </location>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <begin position="170"/>
            <end position="177"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004657"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022433"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023179"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00037488"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00038612"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033955.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1906" mass="220092" checksum="324E1310A6A47674" modified="2019-05-08" version="1">LAKMMDMSEFGEAAPFLRKSDKEIMILQTVAFDGKTGYVFLCLVCDSSCQQCYTCLSLQTVTVKEDDVQPMNPPKFDMIEDMAMLTHLNEASVLHNLNRRYSHWMIYTYSGLFCVTINPYKWLPVYNPEVVNAYKGKRRSEAPPHIFSIADNAYNDMLRNRENQSMLITGESGAGKTVNTKRVIQYFATVAALGEPGKKSVSMQGGTLEDQIIQANPALEAFGNAKTLRNDNSSRFGKFIRIHFGTTGKLSSADIEIYLLEKSRVIFQQPGERSYHIFYQILSGKKAELQDMLLVSTNPYDYHFSSQGAVTVDNLDDGEELMATDQAMDILGFIPDEKYGAYKLTGSIMHFGNMKFKQRPREEQAETDGTESTDKAAYLMGLNSSDLAKGLLHPRVKVGNEYVTKGQSVEQVVYAVGALSKAVYDRMFKWLVIRINKTLDTRLPRQFFIGVLDIAGFEIFDYNSFEQLCINYTNEKLQQFFNHHMFVLEQEEYKKEGIEWVFIDFGLDLQACIDLIEKPLGILSILEEECMFPKATDMTFKAKLYDNHLGKSPNLQKPRIDKKRKYEAHFELLHYAGAVPYNIIGWLEKNKDPLNETVVGIFQKSANKLLASLFENYVSADSGNTEKKRKKGASFQTVSSLHKENLNKLMTNLRSTAPHFVRCIIPNETKTPGAMDAYLVLHQLRCNGVLEGIRICRKGFPNRVLYADFKQRYRILNPAAIPEDKFVDSRKASEKLLASLDIDQTQYRFGHTKVFFKAGLLGHLEEMRDHRLSKILTMIQARARGKLMRIEFQKIVERRDALLVIQWNIRAFMAVKNWPWMKLFFKIKPLLKSAETEKEMANMKEEFLKLKEALEKSEARRKELEEKQVSLVQEKNDLLLQLQAEQDTLADAEERCDLLIKSKIQMEAKVKELTERVEDEEEMNSELTSKKRKLEDECSELKKDIDDLEITLAKVEKEKHATENKVKNLTEEMASLDETISKLTKEKKSLQEAHQQALDDLQAEEDKVNTLSKTKLKLEQQVDDLEGSLEQEKKIRMDLERVKRKLEGDLKLTQENTVCILNSKLEDEQTIVMQLQKKIKELQARIEELEEELEAERAARAKVEKQRSDLARELEELSERLEEAGGATAAQLEMNKKREAEFLKVRRDLEEATLHYEATAATLRKKHADSMAELGEQVDNLQRVKQKLEKEKSELKLEVDDLLSNMEQMVKGKSNAEKLCRTYEDNLNETKTKLDEMTRLMNDLTTQKGKLQTENGEFARQLEEKESLISQLSRGKTSFTQQIEELRRLLEEETKSKNALAHALQAARHDCDLLREQFEEEQEAKAELQRALSKGNAEVAQWRTKYETDAIQRTEELEEAKKKLAARLQEAEEAIEAANAKCSSLEKTKHRLQNELEDIVIDLERANSAAAALDKKQRNFDKLISDWKQKYEETQAELEASQKEARSLSTELFKLKNAYEETLDNLETVKRENKNLQEEISDLTDQISEGNKSLHEIEKIKKQVEQEKSEVQLALEEAEGALEHEESKSLRFQLEISQIKAEFERKLAEKDEEMENIRRNQQRTIDSLQSTLDSEAKSRNEAIRLKKKMEGDLNEMEIQLSHANRHASEATKSARILQAQIKDLQVQLDDSEHLNEDLKEQLAVTDRRNNLIQSELDELRALLDQTERGRKLAEQELLEATERVNLLHTQNTSLINQKKKLETDISQMQNEVEEAIQECRNAEEKAKKAITDAAMMAEELKKEQDTSAHLERMKKNMEQTIKDLQKRLDEAEQIALKGGKKQIQKLESRVRELENDLENELRRNTESQKGVRKYERRIKELTYQAEEDKKNLTRMQELIDKLQLKVKSYKHQAEEAEAQSNQYLAKYRKQQHELDDAEERAEIAESQVNKLRTKTRDIGIKKSENK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0N2</accession>
    <name>A0A452J0N2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">Myosin motor domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">Muscle contraction.</text>
    </comment>
    <comment type="subunit">
        <text evidence="14">Muscle myosin is a hexameric protein that consists of 2 heavy chain subunits (MHC), 2 alkali light chain subunits (MLC) and 2 regulatory light chain subunits (MLC-2).</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Myofibril</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 15">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038409.1">
        <property type="protein sequence ID" value="ENSGAGP00000033915.1"/>
        <property type="gene ID" value="ENSGAGG00000020688.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030016">
        <property type="term" value="C:myofibril"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032982">
        <property type="term" value="C:myosin filament"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016460">
        <property type="term" value="C:myosin II complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000146">
        <property type="term" value="F:microfilament motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000003">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000004">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000013">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000007">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000008">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.4820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.530:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000024">
        <property type="entry name" value="Myosin heavy chain, isoform J"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.720:FF:000001">
        <property type="entry name" value="Myosin heavy chain, muscle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.340">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.370">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.4820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.2420">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002928">
        <property type="entry name" value="Myosin_tail"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014751">
        <property type="entry name" value="XRCC4-like_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615">
        <property type="entry name" value="MYOSIN HEAVY CHAIN, NON-MUSCLE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615:SF69">
        <property type="entry name" value="MYOSIN-6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01576">
        <property type="entry name" value="Myosin_tail_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90257">
        <property type="entry name" value="Myosin rod fragments"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57997">
        <property type="entry name" value="Tropomyosin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12 15" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="7 15" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="8 16" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0488">Methylation</keyword>
    <keyword evidence="10 15" id="KW-0505">Motor protein</keyword>
    <keyword evidence="11" id="KW-0514">Muscle protein</keyword>
    <keyword evidence="9 15" id="KW-0518">Myosin</keyword>
    <keyword evidence="6 15" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0787">Thick filament</keyword>
    <feature type="domain" description="Myosin motor" evidence="17">
        <location>
            <begin position="81"/>
            <end position="773"/>
        </location>
    </feature>
    <feature type="region of interest" description="Actin-binding" evidence="15">
        <location>
            <begin position="650"/>
            <end position="672"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="16">
        <location>
            <begin position="835"/>
            <end position="1131"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="16">
        <location>
            <begin position="1175"/>
            <end position="1898"/>
        </location>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <begin position="174"/>
            <end position="181"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004657"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022433"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023179"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00037488"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00038612"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033915.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1912" mass="220857" checksum="E9DDDAC9DB0CC927" modified="2019-05-08" version="1">ELAKMMDMSEFGEAAPFLRKSDKEIMILQTVAFDGKTGYVFLCSVERQGDSSCQQCYTCLSLQTVTVKEDDVQPMNPPKFDMIEDMAMLTHLNEASVLHNLNRRYSHWMIYTYSGLFCVTINPYKWLPVYNPEVVNAYKGKRRSEAPPHIFSIADNAYNDMLRNRENQSMLITGESGAGKTVNTKRVIQYFATVAALGEPGKKSVSMQGGTLEDQIIQANPALEAFGNAKTLRNDNSSRFGKFIRIHFGTTGKLSSADIEIYLLEKSRVIFQQPGERSYHIFYQILSGKKAELQDMLLVSTNPYDYHFSSQGAVTVDNLDDGEELMATDQAMDILGFIPDEKYGAYKLTGSIMHFGNMKFKQRPREEQAETDGTESTDKAAYLMGLNSSDLAKGLLHPRVKVGNEYVTKGQSVEQVVYAVGALSKAVYDRMFKWLVIRINKTLDTRLPRQFFIGVLDIAGFEIFDYNSFEQLCINYTNEKLQQFFNHHMFVLEQEEYKKEGIEWVFIDFGLDLQACIDLIEKPLGILSILEEECMFPKATDMTFKAKLYDNHLGKSPNLQKPRIDKKRKYEAHFELLHYAGAVPYNIIGWLEKNKDPLNETVVGIFQKSANKLLASLFENYVSADSGNTEKKRKKGASFQTVSSLHKENLNKLMTNLRSTAPHFVRCIIPNETKTPGAMDAYLVLHQLRCNGVLEGIRICRKGFPNRVLYADFKQRYRILNPAAIPEDKFVDSRKASEKLLASLDIDQTQYRFGHTKVFFKAGLLGHLEEMRDHRLSKILTMIQARARGKLMRIEFQKIVERRDALLVIQWNIRAFMAVKNWPWMKLFFKIKPLLKSAETEKEMANMKEEFLKLKEALEKSEARRKELEEKQVSLVQEKNDLLLQLQAEQDTLADAEERCDLLIKSKIQMEAKVKELTERVEDEEEMNSELTSKKRKLEDECSELKKDIDDLEITLAKVEKEKHATENKVKNLTEEMASLDETISKLTKEKKSLQEAHQQALDDLQAEEDKVNTLSKTKLKLEQQVDDLEGSLEQEKKIRMDLERVKRKLEGDLKLTQENTVCILNSKLEDEQTIVMQLQKKIKELQARIEELEEELEAERAARAKVEKQRSDLARELEELSERLEEAGGATAAQLEMNKKREAEFLKVRRDLEEATLHYEATAATLRKKHADSMAELGEQVDNLQRVKQKLEKEKSELKLEVDDLLSNMEQMVKGKSNAEKLCRTYEDNLNETKTKLDEMTRLMNDLTTQKGKLQTENGEFARQLEEKESLISQLSRGKTSFTQQIEELRRLLEEETKSKNALAHALQAARHDCDLLREQFEEEQEAKAELQRALSKGNAEVAQWRTKYETDAIQRTEELEEAKKKLAARLQEAEEAIEAANAKCSSLEKTKHRLQNELEDIVIDLERANSAAAALDKKQRNFDKLISDWKQKYEETQAELEASQKEARSLSTELFKLKNAYEETLDNLETVKRENKNLQEEISDLTDQISEGNKSLHEIEKIKKQVEQEKSEVQLALEEAEGALEHEESKSLRFQLEISQIKAEFERKLAEKDEEMENIRRNQQRTIDSLQSTLDSEAKSRNEAIRLKKKMEGDLNEMEIQLSHANRHASEATKSARILQAQIKDLQVQLDDSEHLNEDLKEQLAVTDRRNNLIQSELDELRALLDQTERGRKLAEQELLEATERVNLLHTQNTSLINQKKKLETDISQMQNEVEEAIQECRNAEEKAKKAITDAAMMAEELKKEQDTSAHLERMKKNMEQTIKDLQKRLDEAEQIALKGGKKQIQKLESRVRELENDLENELRRNTESQKGVRKYERRIKELTYQAEEDKKNLTRMQELIDKLQLKVKSYKHQAEEAEAQSNQYLAKYRKQQHELDDAEERAEIAESQVNKLRTKTRDIGIKKVHEEEE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0N8</accession>
    <name>A0A452J0N8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Vascular endothelial growth factor receptor 3</fullName>
            <ecNumber evidence="4">2.7.10.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="34 35">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="35" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="34" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="34" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="25">
            <text>L-tyrosyl-[protein] + ATP = O-phospho-L-tyrosyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10596"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.10.1"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="30">Membrane</location>
            <topology evidence="30">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="30">Belongs to the protein kinase superfamily. Tyr protein kinase family. CSF-1/PDGF receptor subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.10.1" evidence="4"/>
    <dbReference type="AlphaFoldDB" id="A0A452J0N8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038416.1">
        <property type="protein sequence ID" value="ENSGAGP00000033922.1"/>
        <property type="gene ID" value="ENSGAGG00000024090.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043235">
        <property type="term" value="C:receptor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019838">
        <property type="term" value="F:growth factor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004714">
        <property type="term" value="F:transmembrane receptor protein tyrosine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001525">
        <property type="term" value="P:angiogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030335">
        <property type="term" value="P:positive regulation of cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043408">
        <property type="term" value="P:regulation of MAPK cascade"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048010">
        <property type="term" value="P:vascular endothelial growth factor receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000077">
        <property type="entry name" value="Vascular endothelial growth factor receptor 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000532">
        <property type="entry name" value="Vascular endothelial growth factor receptor 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000041">
        <property type="entry name" value="Vascular endothelial growth factor receptor 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000143">
        <property type="entry name" value="Vascular endothelial growth factor receptor 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000411">
        <property type="entry name" value="Vascular endothelial growth factor receptor 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000548">
        <property type="entry name" value="vascular endothelial growth factor receptor 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013098">
        <property type="entry name" value="Ig_I-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050122">
        <property type="entry name" value="RTK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001245">
        <property type="entry name" value="Ser-Thr/Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008266">
        <property type="entry name" value="Tyr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020635">
        <property type="entry name" value="Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001824">
        <property type="entry name" value="Tyr_kinase_rcpt_3_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041348">
        <property type="entry name" value="VEGFR-2_TMD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055229">
        <property type="entry name" value="VEGFR1-3_5th"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416">
        <property type="entry name" value="TYROSINE-PROTEIN KINASE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416:SF49">
        <property type="entry name" value="VASCULAR ENDOTHELIAL GROWTH FACTOR RECEPTOR 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07679">
        <property type="entry name" value="I-set"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13927">
        <property type="entry name" value="Ig_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22971">
        <property type="entry name" value="Ig_VEGFR-1-like_5th"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07714">
        <property type="entry name" value="PK_Tyr_Ser-Thr"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21339">
        <property type="entry name" value="VEGFR-1-like_Ig-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17988">
        <property type="entry name" value="VEGFR-2_TMD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000615">
        <property type="entry name" value="TyrPK_CSF1-R"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00109">
        <property type="entry name" value="TYRKINASE"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01832">
        <property type="entry name" value="VEGFRECEPTOR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01835">
        <property type="entry name" value="VEGFRECEPTR3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00219">
        <property type="entry name" value="TyrKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00109">
        <property type="entry name" value="PROTEIN_KINASE_TYR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00240">
        <property type="entry name" value="RECEPTOR_TYR_KIN_III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0037">Angiogenesis</keyword>
    <keyword evidence="16 27 29" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="20" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="22" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="24 30" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="15" id="KW-0418">Kinase</keyword>
    <keyword evidence="28" id="KW-0460">Magnesium</keyword>
    <keyword evidence="18 31" id="KW-0472">Membrane</keyword>
    <keyword evidence="28" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="14 27 29" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="23" id="KW-0539">Nucleus</keyword>
    <keyword evidence="8" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="21 30" id="KW-0675">Receptor</keyword>
    <keyword evidence="35" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="13" id="KW-0677">Repeat</keyword>
    <keyword evidence="12" id="KW-0732">Signal</keyword>
    <keyword evidence="10" id="KW-0808">Transferase</keyword>
    <keyword evidence="11 30 31" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="17 31" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="19" id="KW-0829">Tyrosine-protein kinase</keyword>
    <feature type="transmembrane region" description="Helical" evidence="31">
        <location>
            <begin position="12"/>
            <end position="29"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="31">
        <location>
            <begin position="578"/>
            <end position="599"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="33">
        <location>
            <begin position="242"/>
            <end position="358"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="33">
        <location>
            <begin position="373"/>
            <end position="473"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="33">
        <location>
            <begin position="480"/>
            <end position="566"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="32">
        <location>
            <begin position="647"/>
            <end position="937"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="26">
        <location>
            <position position="801"/>
        </location>
    </feature>
    <feature type="binding site" evidence="27">
        <location>
            <begin position="654"/>
            <end position="661"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="27 29">
        <location>
            <position position="681"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="27">
        <location>
            <position position="805"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="28">
        <location>
            <position position="806"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="28">
        <location>
            <position position="819"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00011902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022258"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022657"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023137"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
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        </source>
    </evidence>
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    </evidence>
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        <source>
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    </evidence>
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    </evidence>
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        <source>
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    </evidence>
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    </evidence>
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    </evidence>
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        </source>
    </evidence>
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        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
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        <source>
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    </evidence>
    <evidence type="ECO:0000313" key="35">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
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    </evidence>
    <sequence length="1027" mass="116962" checksum="17E6A859D5E2732E" modified="2019-05-08" version="1">MPEIHLFPVSPPSVLLLPLFILSFLYFSFSDFEQPFINRPETLLINKKENTWVPCLVSIPDLNITLFLPSSIIYPDGRRILWDNKKGMLVPTHLIRDSLFVQCETIIDKKLFRSNYFIVHVTGNELYDIQLFPKKAMELLVGEKLVLNCTVWAEFNAGVNFQWDYPGKQVHQLDRSLSASLPSTPFCLVTCLRGPAFTEKPFINVEWRKGSVIEATAGDEVVKLPVKVVITNIVILLISVPPHIHEKEASSPSIYSRRSHQALTCTVYGIPAPEVIQWQWRPRHQRDRMPECKDWKDVSPQDAENPIESIDTWTESVEGRNKTVSKLVIQEANVSVMYKCVASNKVGRDERLIYFYVTNGFEIESQPSEEPIEGQDLRLSCNADNYTYENLQWYRLNLSKLHHEEGNPLILDCKNVHHYATKMQGELHFKPDSNDAALLLTIPNISLREEGDYVCEVQNRKTREKHCHKKYISVQALEVPRLKQNLTDIWVNVSDSIEMRCKVDGAHIPDISWYKDEKLVEEVSGIDLADSNQRLSIQRVREEDAGLYLCSVCNAKGCVNSSASVSVEGSDDRTNVEIVILIGTGVIAVFFWILLILIFCNIKRPAHADIKTGYLSIIMDPGEVPLEEQCEYLPYDSSKWEFPRERLRLGKVLGHGAFGKVVEASAFGINKSNSCETVAVKMLKEGATASEHKALMSELKILIHIGNHLNVVNLLGACTKPNGPLMVIVEFCKYGNLSNYLRTKREGFSPYRVCEITFFLSLWTVDDLWQSPLTMEDLICYSFQVARGMEFLASRKCIHRDLAARNILLSENNVVKICDFGLARDIYKDPDYVRKGSARLPLKWMAPESIFDKVYTTQSDVWSFGVLLWEIFSLGASPYPGVQINEEFCQRLKDGTRMRAPEYATAEIYRIMLSCWHGDPKERPTFSDLVEILGDLLQENVQQVRFSCTVIIQHQTALGQFSSNSLFFLSLLAIPLDNQTDSGMVLASEEFERIENRHRKEGGFRYLLNSLPWTTLVVHSSACCEAP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J100</accession>
    <name>A0A452J100_9SAUR</name>
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        <submittedName>
            <fullName evidence="3">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J100"/>
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        <property type="protein sequence ID" value="ENSGAGP00000034071.1"/>
        <property type="gene ID" value="ENSGAGG00000024210.1"/>
    </dbReference>
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    </dbReference>
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    </dbReference>
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        <property type="entry name" value="CENTROSOMAL PROTEIN OF 128 KDA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46657:SF1">
        <property type="entry name" value="CENTROSOMAL PROTEIN OF 128 KDA"/>
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    </dbReference>
    <proteinExistence type="predicted"/>
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            <begin position="55"/>
            <end position="88"/>
        </location>
    </feature>
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    </feature>
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        <location>
            <begin position="828"/>
            <end position="862"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
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    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
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        <source>
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    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="896" mass="106672" checksum="22E5F608F12A7446" modified="2019-05-08" version="1">MLGQYREYNNEQTEAIATLKETLEQSIGQLRSQRLSRNSGMRSASLSSLYTSDLDGEAATGSHHFQPTSPLRDYGDSQSTRRRRSRSANVRFVDEANNLDQLHSLHHSLRDLSSEQVRLGDDISRELSRRYRTDAETKKILTDLSEKLTESQRQETVSERVERRLQEIEQEMRTERQVVEGRQDQLGHMSMQLQELEQELERSQRKLDQSEDSREALLHQIEDLHSQLLRAEADRVDLQHQVSQVAMHHQSCHDEQEDDRRIRSVVERSEREKQELEKQILELRAKLNHSAVMSKVEELKRCIEHKDKEKAQLAMHIQVLTSDLENREKQQQRMLDQLKEIQSCYKACESERRQTELQTAELAQQLEESAKEADRYLAEFRQSETLRLETEKKKEELKVKAQESIRHWKLKYKKLERDMVKQNETISQLMDKSDQVLKEKDDLKAQLLSAIHQIENLRKELSDVLTKRAQQEEELHCKEVKLSETKSQQIVLEQEIREVHDKANKLESELQKQILVQNQMRNEKEHLEEELETANMINEKDQERFLEMQEDIKNLSAIRAELTNRIAEEEKVKKEILKNFSDLQKHQESKQEEMTTTSWQLKMERDVHQQELADLRSELKNVKTKHERNVQELMKLFRQEKDETENHIRRLKTELLDEKNVVKAQRRQVEKMKTECDKMTEELTQSEQENTKLRRKYELVKQELEEKDKQINSEDDHLRRMEEARLQLRDQLRCLETEQESILTMIGSEIDAACEILSGDSVEKFKAISLTPGLQNDPHRWLAETKTKLQWMCEEVKERESKERKLRRHLLQSREQLKHLTLSKECELQSLFEQIKKQEHVLEEVNREKRDLLEKTHRREEEMAALQVYLLIRLEKLYFLVTFLLNAFLLNPHRGN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J106</accession>
    <name>A0A452J106_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">RAR related orphan receptor B</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 11">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="11">Belongs to the nuclear hormone receptor family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J106"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038560.1">
        <property type="protein sequence ID" value="ENSGAGP00000034053.1"/>
        <property type="gene ID" value="ENSGAGG00000024178.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004879">
        <property type="term" value="F:nuclear receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd06968">
        <property type="entry name" value="NR_DBD_ROR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.565.10:FF:000005">
        <property type="entry name" value="Nuclear orphan receptor ROR-beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.50.10:FF:000003">
        <property type="entry name" value="Nuclear orphan receptor ROR-beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.50.10">
        <property type="entry name" value="Erythroid Transcription Factor GATA-1, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.565.10">
        <property type="entry name" value="Retinoid X Receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035500">
        <property type="entry name" value="NHR-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044101">
        <property type="entry name" value="NR_DBD_ROR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000536">
        <property type="entry name" value="Nucl_hrmn_rcpt_lig-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001723">
        <property type="entry name" value="Nuclear_hrmn_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003079">
        <property type="entry name" value="ROR_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001628">
        <property type="entry name" value="Znf_hrmn_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013088">
        <property type="entry name" value="Znf_NHR/GATA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45805">
        <property type="entry name" value="NUCLEAR HORMONE RECEPTOR HR3-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45805:SF7">
        <property type="entry name" value="NUCLEAR RECEPTOR ROR-BETA-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00104">
        <property type="entry name" value="Hormone_recep"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00105">
        <property type="entry name" value="zf-C4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01293">
        <property type="entry name" value="RORNUCRECPTR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00398">
        <property type="entry name" value="STRDHORMONER"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00047">
        <property type="entry name" value="STROIDFINGER"/>
    </dbReference>
    <dbReference type="SMART" id="SM00430">
        <property type="entry name" value="HOLI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00399">
        <property type="entry name" value="ZnF_C4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57716">
        <property type="entry name" value="Glucocorticoid receptor-like (DNA-binding domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48508">
        <property type="entry name" value="Nuclear receptor ligand-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51843">
        <property type="entry name" value="NR_LBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00031">
        <property type="entry name" value="NUCLEAR_REC_DBD_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51030">
        <property type="entry name" value="NUCLEAR_REC_DBD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="7 11" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3 11" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10 11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="9 11" id="KW-0675">Receptor</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8 11" id="KW-0804">Transcription</keyword>
    <keyword evidence="6 11" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="5 11" id="KW-0862">Zinc</keyword>
    <keyword evidence="4 11" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="Nuclear receptor" evidence="13">
        <location>
            <begin position="18"/>
            <end position="93"/>
        </location>
    </feature>
    <feature type="domain" description="NR LBD" evidence="14">
        <location>
            <begin position="226"/>
            <end position="464"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="104"/>
            <end position="123"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="RuleBase" id="RU004334"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS51030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS51843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034053.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="17">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="476" mass="54517" checksum="6A6E3963ED909225" modified="2019-05-08" version="1">MEHKPCEQRLKELAQIEVIPCKICGDKSSGIHYGVITCEGCKGFFRRSQQNNASYSCSRQRNCLIDRTNRNRCQHCRLQKCLALGMSRDAVKFGRMSKKQRDSLYAEVQKHQQNQEQNRVSKDDSEVLARVYPTSVSSGVSDLDDISLLSDGLLFDFPLTPDGGSTYYNLDLLTSAQPSPDQSSMDVSDAKFIKQESIYELMLEPGLFAHSTLEGSQLASDISMLEIDRIAQNVVKSHLETCQYTTEELKRLAWMLYTQEEIRSLQNKSCETMWQQCSLQISNAIQYVVEFAKRIDGFMELCQNDQIILLKAGCLEVLLIRMIRAFNPLNNMVLFEGKYGGMQMFKSLGCDDLINAVFELGRNLCRLQLSDEEIALFTAAVLLSPDRPWLTESKKVQKLQDKIYIALQHEIQKKHSSEDKLSKMISKLPLMKTICNLHLDKLEFFRLLHPETAMNFPPLYKEVFNSELQYSDPRES</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J109</accession>
    <name>A0A452J109_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">LisH domain and HEAT repeat-containing protein KIAA1468</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Golgi apparatus</location>
            <location evidence="2">trans-Golgi network</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Recycling endosome</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038508.1">
        <property type="protein sequence ID" value="ENSGAGP00000034008.1"/>
        <property type="gene ID" value="ENSGAGG00000023990.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055037">
        <property type="term" value="C:recycling endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005802">
        <property type="term" value="C:trans-Golgi network"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032367">
        <property type="term" value="P:intracellular cholesterol transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000080">
        <property type="entry name" value="lisH domain and HEAT repeat-containing protein KIAA1468 homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000218">
        <property type="entry name" value="lisH domain and HEAT repeat-containing protein KIAA1468 homolog isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021133">
        <property type="entry name" value="HEAT_type_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006594">
        <property type="entry name" value="LisH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040362">
        <property type="entry name" value="RELCH"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR32059">
        <property type="entry name" value="RAB11-BINDING PROTEIN RELCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR32059:SF0">
        <property type="entry name" value="RAB11-BINDING PROTEIN RELCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00667">
        <property type="entry name" value="LisH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50077">
        <property type="entry name" value="HEAT_REPEAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50896">
        <property type="entry name" value="LISH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7 11" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0967">Endosome</keyword>
    <keyword evidence="6" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="8" id="KW-0445">Lipid transport</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <feature type="repeat" description="HEAT" evidence="10">
        <location>
            <begin position="987"/>
            <end position="1023"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1"/>
            <end position="51"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="422"/>
            <end position="447"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="11">
        <location>
            <begin position="182"/>
            <end position="216"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="11">
        <location>
            <begin position="343"/>
            <end position="377"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="12">
        <location>
            <begin position="25"/>
            <end position="35"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="427"/>
            <end position="447"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004172"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004601"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00084053"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00103"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000034008.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1197" mass="132738" checksum="358D977AB43F590E" modified="2019-05-08" version="1">MAAAGGGSSVNPFLSDSEEEEPRDGGGGNDPGLGGPLSPQQHEEEEEAAVEVPGVLPGLRAAAPGEAGWAGPGAGEPPRVPVDALAAQLLRDQFLLTALELHTELLESGRELPRLRDYFSNPGNFERQSGTPPAGGAPGLWGAAGLGLLGGCELITEGGGDRAGSISTLDSLDFARYSDDGNRETDERVAVLEFELRKAKETIQALRANLTQAAENEVPLQERKNYKSSPEIQEPIKPLEKRALNFLVNEFLLKNSYKLTSITFSDESDDQDFELWDDVGLNIPKPPDLLQLYRDFGNHQVTARDVVDVAVGVEEDELEVTTPTLGSVPVFGVSQSTEQFLIVQKLEDQINALNSEKWSLTEQIRRLESEIDFLKNENFAVPVVCDLVHVPLSVPEDNGSYLDIRSSDNDSKHGKTEEISLSISAEADSKTPKDDTLNSLPSKDREKLPSCALSNKAAIHFDKPNRTLSPAFHQALLSFCRMSAESRLGSEVSRIADSEKSVMLMLGRCLPHIVPNVLLAKREELIPLILCTACLHPEPKERDQLLHILFNLIKRPDDEQRQMILTGCVAFARHVGPTRVEAELLPQCWEQINHKYPERRLLVAESCGALAPYLPKEIRSSLVLSMLQQMLMEDKADLVREAVIKSLGIIMGYIDDPDKYQQGFELLLSALGDPSERVVSATHQVFLPAYAAWTTELGNLQLHLIPTLLNKIEKLLREGEHGLDEHKLHMYLSALQSLIPSLFSLVLQNAPFTSKAKLQGEVPPIEVTRFPRPVSPLQDVATIIGSREQLAVLLQLYDYQLELEGTTGWETLLWVVNQLLPQLIEIVGKINVASTACVHEFSRFFWRLCRTFGKIFTNTKVKPQFQEILRLSEENIDSTAGNGVLTKATVPIYATGVLTCYIQEEDRKLLIGFLEDVMTMLSLSHAPLNSLKASFVELGANPAYHELLLTVLWYGVVHTSALVRCTAARMFELLVKGVHETLVAQRVVPALITYSSDPEISVRIATIPAFGTIMETVTQREXRVKMQLASFLEDPQYQDQHSLQTEIIKTFGRVGPNAEPRFRDEFVIPHLHKLALVNNQQSVDSKRLDIATHLFEAYSALSCCFISEDLMVNHFLPGLRCLRTDMEHLSPEHEVILSSMIKECEQKVENKTVQEPQGSMSIAASLVSEDTKTKFLNKMGQLTTSGAMLANVFQRKK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHV0</accession>
    <name>A0A452GHV0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Nephronectin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">Functional ligand of integrin alpha-8/beta-1 in kidney development. Regulates the expression of GDNF with integrin alpha-8/beta-1 which is essential for kidney development. May also play a role in the development and function of various tissues, regulating cell adhesion, spreading and survival through the binding of several integrins.</text>
    </comment>
    <comment type="subunit">
        <text evidence="14">Homodimer and homotrimer.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the nephronectin family.</text>
    </comment>
    <comment type="caution">
        <text evidence="16">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHV0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001353.1">
        <property type="protein sequence ID" value="ENSGAGP00000001205.1"/>
        <property type="gene ID" value="ENSGAGG00000000964.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00054">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="CDD" id="cd06263">
        <property type="entry name" value="MAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000038">
        <property type="entry name" value="Fibrillin 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000187">
        <property type="entry name" value="nephronectin isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000476">
        <property type="entry name" value="nephronectin isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.200:FF:000133">
        <property type="entry name" value="nephronectin isoform X1"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="nephronectin isoform X2"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="nephronectin isoform X2"/>
        <property type="match status" value="1"/>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
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        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001881">
        <property type="entry name" value="EGF-like_Ca-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000152">
        <property type="entry name" value="EGF-type_Asp/Asn_hydroxyl_site"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018097">
        <property type="entry name" value="EGF_Ca-bd_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009030">
        <property type="entry name" value="Growth_fac_rcpt_cys_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000998">
        <property type="entry name" value="MAM_dom"/>
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    <dbReference type="InterPro" id="IPR052235">
        <property type="entry name" value="Nephronectin_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049883">
        <property type="entry name" value="NOTCH1_EGF-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24050:SF19">
        <property type="entry name" value="NEPHRONECTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24050">
        <property type="entry name" value="PA14 DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07645">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00629">
        <property type="entry name" value="MAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00179">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00137">
        <property type="entry name" value="MAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57184">
        <property type="entry name" value="Growth factor receptor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00010">
        <property type="entry name" value="ASX_HYDROXYL"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01186">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01187">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50060">
        <property type="entry name" value="MAM_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0106">Calcium</keyword>
    <keyword evidence="11" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="3" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="9" id="KW-0221">Differentiation</keyword>
    <keyword evidence="12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6 16" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="5" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="7 18" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="18">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019037789" description="Nephronectin" evidence="18">
        <location>
            <begin position="21"/>
            <end position="625"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="19">
        <location>
            <begin position="121"/>
            <end position="160"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="19">
        <location>
            <begin position="201"/>
            <end position="245"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="19">
        <location>
            <begin position="246"/>
            <end position="286"/>
        </location>
    </feature>
    <feature type="domain" description="MAM" evidence="20">
        <location>
            <begin position="476"/>
            <end position="619"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="343"/>
            <end position="421"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="17">
        <location>
            <begin position="346"/>
            <end position="355"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="17">
        <location>
            <begin position="376"/>
            <end position="389"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="399"/>
            <end position="421"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009738"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00053701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00064366"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00069631"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50060"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001205.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="625" mass="69202" checksum="DBD91B60459322BC" modified="2019-05-08" version="1">MDLFFLRLLAVAWLHLEAAAEFDGRWPRQIVSSMGLCRFGSRVDCCWGWARQSWGQCQPICQPRCKHGECIGPNKCKCLPGYTGKTCNQAEHLYTSPTDRHSEPPLFRSLDHHATNVPSRDLNECGLKPRPCKHRCMNTYGSYKCYCLNGYMLMPDGSCSNALSCSMANCQYGCDVVKGEVRCHCPSPGLQLGPDGRTCIDVDECATGRVVCPRFRKCVNTFGSYICKCHKGFELMYIGGKYQCHDIDECSIGQYQCGSFSRCYNTQGTYKCKCKEGYRGDGINCLPIPKVMIEPSRPNNIFKGNTTLPKGSSGMTNRIPDIGGTRQPPRPPHVPAIITVRPAPKPTVQPTTKPTIWRTPKPTTRFVPKPMTTRPPLKPATRPPTPPQPTTSKPAPTARSPQLTTKDSSLVPTETTSPQGFTVDNRIQIEPQKPRGDVFIPRQPGVSNNLFEIFEIERGISADEDEANDDPGILVHSCNFDSGLCGWIKDKDDNLHWEPLRDPSGGQYMTISEPKGKEGKVARLVLPLGQLAHSGDLCLSFRHKVSGLHSGLLQVFLRKNGAHGPAVWGRNGGHGWRQTQITLQDSGIKSVIFKGEKGKGRTGDIGLDDVSLRKGRCSEEHEQHH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHV3</accession>
    <name>A0A452GHV3_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="9">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the protein-tyrosine phosphatase family. Non-receptor class dual specificity subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHV3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001351.1">
        <property type="protein sequence ID" value="ENSGAGP00000001203.1"/>
        <property type="gene ID" value="ENSGAGG00000000928.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017017">
        <property type="term" value="F:MAP kinase tyrosine/serine/threonine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001706">
        <property type="term" value="P:endoderm formation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043409">
        <property type="term" value="P:negative regulation of MAPK cascade"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd14640">
        <property type="entry name" value="DSP_DUSP4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000015">
        <property type="entry name" value="Dual specificity phosphatase 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000340">
        <property type="entry name" value="Dual-sp_phosphatase_cat-dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008343">
        <property type="entry name" value="MKP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016130">
        <property type="entry name" value="Tyr_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003595">
        <property type="entry name" value="Tyr_Pase_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020422">
        <property type="entry name" value="TYR_PHOSPHATASE_DUAL_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10159">
        <property type="entry name" value="DUAL SPECIFICITY PROTEIN PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10159:SF111">
        <property type="entry name" value="DUAL SPECIFICITY PROTEIN PHOSPHATASE 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00782">
        <property type="entry name" value="DSPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01764">
        <property type="entry name" value="MAPKPHPHTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00195">
        <property type="entry name" value="DSPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00404">
        <property type="entry name" value="PTPc_motif"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00383">
        <property type="entry name" value="TYR_PHOSPHATASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50054">
        <property type="entry name" value="TYR_PHOSPHATASE_DUAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="5" id="KW-0539">Nucleus</keyword>
    <keyword evidence="4" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="7">
        <location>
            <begin position="22"/>
            <end position="163"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="8">
        <location>
            <begin position="90"/>
            <end position="141"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008601"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001203.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="200" mass="22274" checksum="B686AB3A134F1604" modified="2019-05-08" version="1">FSARRHSNLGKDCSSPESEGGGPVEILPFLYLGSAYHAARRDMLDTLGITALLNVSSDCPNHFEGHYQYKCIPVEDNHKADISSWFMEAIEYIDLVKDCHGRVLVHCQAGISRSATICLAYLMMKKRVKLEEAFEFVKQRRSIISPNFSFMGQLLQFESQVLATSCAAEAASPSGTLRERGKLCRHSGLGWSLGSEPWEL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHV4</accession>
    <name>A0A452GHV4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Olduvai domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Golgi apparatus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000001306.1">
        <property type="protein sequence ID" value="ENSGAGP00000001164.1"/>
        <property type="gene ID" value="ENSGAGG00000000835.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060090">
        <property type="term" value="F:molecular adaptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007098">
        <property type="term" value="P:centrosome cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090063">
        <property type="term" value="P:positive regulation of microtubule nucleation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903358">
        <property type="term" value="P:regulation of Golgi organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056273">
        <property type="entry name" value="CDK5RAP2_MYOME_CC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012943">
        <property type="entry name" value="Cnn_1N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052593">
        <property type="entry name" value="MT-associated_AKAP9-binding"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010630">
        <property type="entry name" value="Olduvai_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46501">
        <property type="entry name" value="MYOMEGALIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46501:SF2">
        <property type="entry name" value="MYOMEGALIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23246">
        <property type="entry name" value="CC_CDK5RAP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07989">
        <property type="entry name" value="Cnn_1N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01148">
        <property type="entry name" value="DUF1220"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51316">
        <property type="entry name" value="ODV"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="5" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Olduvai" evidence="9">
        <location>
            <begin position="1483"/>
            <end position="1574"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="924"/>
            <end position="971"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1243"/>
            <end position="1264"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1324"/>
            <end position="1350"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1518"/>
            <end position="1632"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1946"/>
            <end position="1987"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="2173"/>
            <end position="2220"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="41"/>
            <end position="68"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="94"/>
            <end position="150"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="236"/>
            <end position="291"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="350"/>
            <end position="516"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="545"/>
            <end position="630"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="732"/>
            <end position="874"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="992"/>
            <end position="1019"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="1185"/>
            <end position="1212"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="1293"/>
            <end position="1320"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="1765"/>
            <end position="1845"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="2113"/>
            <end position="2147"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="954"/>
            <end position="963"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="1245"/>
            <end position="1256"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="1526"/>
            <end position="1543"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="1567"/>
            <end position="1578"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="1582"/>
            <end position="1602"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="1609"/>
            <end position="1624"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="1946"/>
            <end position="1955"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="1966"/>
            <end position="1984"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="2197"/>
            <end position="2209"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004555"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51316"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001164.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2239" mass="251529" checksum="1F96AA7EDF5B882F" modified="2019-05-08" version="1">MSNGYRTLSQHLNDLKKENFSLKLRIYFLEERMQQKYEASREDVYKRNIELKVEVESLKQELQERQQTLDKAWAAAENLTSCNEAEFQRRYEERQQEKEHVSELLENKIQLLQEEARLAKSEAERTAALVEAEKGRCLELTKKLKELTKEEEKRGAQVLRDDHCAALAQKDKHIEELIQELSAREQLIELLATEKQNLLQRLEEPREIEVQNLPEDHQQLTQCDPVASGRVSESHAAELQDKIQQYDATNKLLQEKLNEMNFELKSIQEASQRQDRTIQSLNEALKSKESETEELYHVIEGQNETMGKLRDMLHRSQLGHLQTSEGPSPSQQQQHMALLDLQNTLFCTQLEMQKLKRAQWQKEHQLAEARRTTQLLEATVQEEQQLKEVSWKHNQELRGVVQQVQAELQGKVQELWALEGEKCCEIRAQEQRVQRLSQRLAHKEQLLQESWELLQRRQSLEKNPVVTDTMLEKLQQRIKHRDAALERAIDEKFCAVEEKEQELQQLHISVREREHDLERLRSVLSSNEATIHSMESLLKAKTLELEQVTATCQNLQWLKEEVEAKSNCWQKEQEGIIQQLQASLHDRNKEVEELTATLLCKLGPGQSEIAEELCLRLQRKERMLQDLLSDRNRQAGEHETELRELLQAMSIREQRSRVTAEKMVQALAEKSDEVQFLRQRLVGNEPGKNPVADGRLLLQKDKQPFQELLPGGCGNATATRISKGEDSKYRMERGLLETAAELEKELINAREELELMARKERESRLELSSLQSVVTAQEEELQVQASDIESLTRNIQSKEDLIKDLQMQLVDPEEIPAVERLTQEVLVLREKVATAESRGQEATGNRQQQLLLMLEGLVAEKSRLNEALQAERQLYSSLVKFHAHPDSPERGQTLQAELEGAQALRGRLEESLGRSMEHLRRLETLGAFGGQPGRDDAKDGSTDSIEGEAPPRTVNHQTSQGLPANSPEAKAAQPDLLTLAPLERDSSLQAELLHVRAKNQQLLEQKAKVEGELWELKALMEEAGFSSLSSIRKVLLSLCLETAELKEQLGGSPLSEGWEDKAEARAALCTQNSKLQSSETVVTLLKEQLVLNCPEGESKFPPQLVTGMATEIDRLWTEMGVASGKRPALEGQLQEDPAKRPCPRSQAVELGPAQAPRGMSQQMEACPAGSWQRLVEPRVGPSGQLVQCRQRNQELQDKLAVSEATVRAQAEQLEQYRTLLSEPLVQQDNKQVQVDLQDLGYETCGRSENEADREETPSPECEEPDVFSKAGLMEELSRPGPWGGLCWGQSEDVAVLQQHVRELQLQLQNSNKALQRLQRRTRSFSSTSDYASGAERLHKPVPSATDEDEGWQSDGVGAFCPPALQASRDLEWLVHRVSLLEAQLHGRWAGGVLPEELRSAVWPGKYDSLIQAQARELSHLRQKMREGRSVCHLLTQHLRDTVKSFEELLRGTDIDYYMGQSFREHLAQGGQLAERLSSKLSSRDRSDVEDKTGHEFVALRLSKELREKERMIETLQAKLQERCETPSSSRALSESPRSNSSASFLSDGPEACSDGDATSEYSQLQGDRSEQPSRRLTDVTHPAALASNPSPAAAPHQASAESIRGYPTGPSSQHPPMDSGQTRTGGPLAAPPVKPLPLTSDFGEASTSRCLHVPHLTPQSCPLQSSTQLDRKAGAAFLESNPLWAQPPMGALYGHLPSGYLASHKLTGADLLEGHLAEIRSLRQRLEESICINDRLREQLEKRLATTTKGNGYPTNIYIQGLESASSLSSETQALREENQSLQLQFAHLSRELEQLRETTRCRLQKAETENRTLAQELAEQRQSGRQLREERLALQENNHRLECTVPLLQQQCQENQQLCQALCAELRIHETLSSSSQAALSACSGDVHRRPPSSSELGALLAEVRTLRRKLLRGIQVNSALRQQLARQQPEGSAAPLFAAHQATPNWQPFQDSSPSPPVRDVGMSSPAPLSPPALLSGPAPLAHQLKEPPVDDALVRRSAPAPAGDAPSSSSASKGGCQVIGHVDDYYTLKQLILEGKTLVHQMASLLRPALAMPSLEPRGTKALDRGSVQQLLSTASALHQLLEQCSSLLTTFWRGALPAAPSPAQPQTAEQAMKDELLALRARLAEQENLLQRAAERLQDTCRLKDNMEHFIVSHLTQTHAVLRKARTNLEKSKPQGSSVKSPPFIGAGEISVGLSQHPSHVIPTNQEHRRKKSCPDTLKWQEDQSWPCFVHLPSS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHX9</accession>
    <name>A0A452GHX9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">DNA ligase</fullName>
            <ecNumber evidence="15">6.5.1.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">DNA ligase that seals nicks in double-stranded during DNA repair. Also involved in DNA replication and DNA recombination.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13 15">
            <text>ATP + (deoxyribonucleotide)n-3'-hydroxyl + 5'-phospho-(deoxyribonucleotide)m = (deoxyribonucleotide)n+m + AMP + diphosphate.</text>
            <dbReference type="EC" id="6.5.1.1"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 16">Belongs to the ATP-dependent DNA ligase family.</text>
    </comment>
    <dbReference type="EC" id="6.5.1.1" evidence="15"/>
    <dbReference type="AlphaFoldDB" id="A0A452GHX9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001390.1">
        <property type="protein sequence ID" value="ENSGAGP00000001238.1"/>
        <property type="gene ID" value="ENSGAGG00000000751.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003910">
        <property type="term" value="F:DNA ligase (ATP) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071897">
        <property type="term" value="P:DNA biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006310">
        <property type="term" value="P:DNA recombination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006281">
        <property type="term" value="P:DNA repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903461">
        <property type="term" value="P:Okazaki fragment processing involved in mitotic DNA replication"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07900">
        <property type="entry name" value="Adenylation_DNA_ligase_I_Euk"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd07969">
        <property type="entry name" value="OBF_DNA_ligase_I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.3260.10:FF:000001">
        <property type="entry name" value="DNA ligase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.50.140:FF:000062">
        <property type="entry name" value="DNA ligase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.470.30:FF:000016">
        <property type="entry name" value="DNA ligase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1490.70">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.3260.10">
        <property type="entry name" value="DNA ligase, ATP-dependent, N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.470.30">
        <property type="entry name" value="DNA ligase/mRNA capping enzyme"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.50.140">
        <property type="entry name" value="Nucleic acid-binding proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050191">
        <property type="entry name" value="ATP-dep_DNA_ligase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000977">
        <property type="entry name" value="DNA_ligase_ATP-dep"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012309">
        <property type="entry name" value="DNA_ligase_ATP-dep_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012310">
        <property type="entry name" value="DNA_ligase_ATP-dep_cent"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016059">
        <property type="entry name" value="DNA_ligase_ATP-dep_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012308">
        <property type="entry name" value="DNA_ligase_ATP-dep_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036599">
        <property type="entry name" value="DNA_ligase_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012340">
        <property type="entry name" value="NA-bd_OB-fold"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00574">
        <property type="entry name" value="dnl1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45674:SF4">
        <property type="entry name" value="DNA LIGASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45674">
        <property type="entry name" value="DNA LIGASE 1/3 FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04679">
        <property type="entry name" value="DNA_ligase_A_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01068">
        <property type="entry name" value="DNA_ligase_A_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04675">
        <property type="entry name" value="DNA_ligase_A_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF117018">
        <property type="entry name" value="ATP-dependent DNA ligase DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56091">
        <property type="entry name" value="DNA ligase/mRNA capping enzyme, catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50249">
        <property type="entry name" value="Nucleic acid-binding proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00697">
        <property type="entry name" value="DNA_LIGASE_A1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00333">
        <property type="entry name" value="DNA_LIGASE_A2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50160">
        <property type="entry name" value="DNA_LIGASE_A3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 15" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="12" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="4" id="KW-0132">Cell division</keyword>
    <keyword evidence="7 15" id="KW-0227">DNA damage</keyword>
    <keyword evidence="9 15" id="KW-0233">DNA recombination</keyword>
    <keyword evidence="10 15" id="KW-0234">DNA repair</keyword>
    <keyword evidence="5" id="KW-0235">DNA replication</keyword>
    <keyword evidence="3 15" id="KW-0436">Ligase</keyword>
    <keyword evidence="6 15" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="ATP-dependent DNA ligase family profile" evidence="17">
        <location>
            <begin position="562"/>
            <end position="698"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007572"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022598"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022705"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023172"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00034003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00058910"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="RuleBase" id="RU000617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="RuleBase" id="RU004196"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50160"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001238.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="830" mass="92524" checksum="F9027BFD3F9540F1" modified="2019-05-08" version="1">LSDSQLRYRLSEAAAMQRNGLSLCLDSPVKSAKRPLRRVLESDSDEEEAVLTARAEMEPTQDKSLERASSKQVSSAHPLSLPWNSHGASLGSLSCDSAASPMHALLQLGACLIHAIIVPVATCAGLPSLGVLVQANCWETLRNTSAVGPVYWTWWLPISLWLCDPSSFSKPVVPLDPMEYNPSKSSYHPVQDACWGHGQRVPYLAVARTFERIEEVSARLKNIETLSNLFRSVVALSPSDILPCIYLCLNRLGPAYKGLELGIGETILMKAVAQAMGRQLDQIKAEAQEKGDLGLVAESSRSNQRTMFTPPKLSAAGVFSKLQEIARMTGSASMNKKIDIIKGLFVACRHSEARYITRSLGGKLRIGLAEQSVLSAIAHAVSLTPPGQGFPPEVLDAGQSKSAEARKAWLEEQTQILKQTFCELPSYDIIVPILLEHGVESLPQHCKITPGIPLKPMLAQPTKGIGEVLKRFEEAAFTCEYKYDGERAQIHILENGDVHIYSRNQEDNTSKYPDIISRIPKVKKDSVQSCILDAEAVAWDPEKRQIQPFQVLTTRKRKDVEAAEIKVQVCLYAFDILYLNGESLVKEPFSRRRQLLHESFVETEGEFLFATSMDTCSLDEISEFLDQSIKDSCEGLMVKTLEVDATYEIAKRSHKWLKLKKDYLEGVGDTLDLVVIGAYLGKGKRVGMYGGFLLACYDEESEEYQSICKVGSVGSLHRLDAHLQQTARPKPRPYYRWDSSTEPNHWLAAVQVWEVKCADLSISPVHKAAIGLVDEEKGISLRFPRFLRVRDDKKPEEATTSSQVAELYRRQQQIQNQQLVEKAEEEEDFY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHY0</accession>
    <name>A0A452GHY0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Reticulon</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="6">Inhibits amyloid precursor protein processing, probably by blocking BACE1 activity.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 7">Endoplasmic reticulum membrane</location>
            <topology evidence="1 7">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHY0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001336.1">
        <property type="protein sequence ID" value="ENSGAGP00000001190.1"/>
        <property type="gene ID" value="ENSGAGG00000000953.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043005">
        <property type="term" value="C:neuron projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0014069">
        <property type="term" value="C:postsynaptic density"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007420">
        <property type="term" value="P:brain development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071787">
        <property type="term" value="P:endoplasmic reticulum tubular network formation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030182">
        <property type="term" value="P:neuron differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.2480:FF:000001">
        <property type="entry name" value="Reticulon"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.2480">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003388">
        <property type="entry name" value="Reticulon"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046964">
        <property type="entry name" value="RTN1-4"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45799:SF5">
        <property type="entry name" value="RETICULON-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45799">
        <property type="entry name" value="RETICULON-LIKE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02453">
        <property type="entry name" value="Reticulon"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50845">
        <property type="entry name" value="RETICULON"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3 7" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="5 7" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 7" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="35"/>
            <end position="64"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="150"/>
            <end position="171"/>
        </location>
    </feature>
    <feature type="domain" description="Reticulon" evidence="8">
        <location>
            <begin position="21"/>
            <end position="208"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004477"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00037306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU210713"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50845"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001190.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="208" mass="23687" checksum="13EE866BF86EA08B" modified="2019-05-08" version="1">MQASADSTKMDCLWSNWKCQAIDLLYWRDIKQTGIVFGSILLLLFSLTQFSVVSVIAYLALAALSATISFRIYKSVLQAVQKTDEGHPFKAYLEMEMSLSQEQIQKYTDCLQLYVNSTVRELRRLFLVQDLVDSLKFAVLMWLLTYVGALFNGLTLMIMAVVSMFTLPVVYDKYQAQIDQYLGLVRTHINTVMAKIQAKIPGAKRKAE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHY5</accession>
    <name>A0A452GHY5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15 17">Protoporphyrinogen oxidase</fullName>
            <ecNumber evidence="5 17">1.3.3.4</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1 17">Catalyzes the 6-electron oxidation of protoporphyrinogen-IX to form protoporphyrin-IX.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="16 17">
            <text>protoporphyrinogen IX + 3 O2 = protoporphyrin IX + 3 H2O2</text>
            <dbReference type="Rhea" id="RHEA:25576"/>
            <dbReference type="ChEBI" id="CHEBI:15379"/>
            <dbReference type="ChEBI" id="CHEBI:16240"/>
            <dbReference type="ChEBI" id="CHEBI:57306"/>
            <dbReference type="ChEBI" id="CHEBI:57307"/>
            <dbReference type="EC" id="1.3.3.4"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="17">
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </cofactor>
        <text evidence="17">Binds 1 FAD per subunit.</text>
    </comment>
    <comment type="pathway">
        <text evidence="3 17">Porphyrin-containing compound metabolism; protoporphyrin-IX biosynthesis; protoporphyrin-IX from protoporphyrinogen-IX: step 1/1.</text>
    </comment>
    <comment type="subunit">
        <text evidence="14">Monomer. Homodimer.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Mitochondrion inner membrane</location>
            <topology evidence="2">Peripheral membrane protein</topology>
            <orientation evidence="2">Intermembrane side</orientation>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4 17">Belongs to the protoporphyrinogen/coproporphyrinogen oxidase family. Protoporphyrinogen oxidase subfamily.</text>
    </comment>
    <dbReference type="EC" id="1.3.3.4" evidence="5 17"/>
    <dbReference type="AlphaFoldDB" id="A0A452GHY5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001337.1">
        <property type="protein sequence ID" value="ENSGAGP00000001191.1"/>
        <property type="gene ID" value="ENSGAGG00000000960.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00251">
        <property type="reaction ID" value="UER00324"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005743">
        <property type="term" value="C:mitochondrial inner membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005758">
        <property type="term" value="C:mitochondrial intermembrane space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004729">
        <property type="term" value="F:oxygen-dependent protoporphyrinogen oxidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006785">
        <property type="term" value="P:heme B biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006782">
        <property type="term" value="P:protoporphyrinogen IX biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="FunFam" id="3.50.50.60:FF:000133">
        <property type="entry name" value="Protoporphyrinogen oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.50.50.60">
        <property type="entry name" value="FAD/NAD(P)-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002937">
        <property type="entry name" value="Amino_oxidase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036188">
        <property type="entry name" value="FAD/NAD-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004572">
        <property type="entry name" value="Protoporphyrinogen_oxidase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050464">
        <property type="entry name" value="Zeta_carotene_desat/Oxidored"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00562">
        <property type="entry name" value="proto_IX_ox"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR42923">
        <property type="entry name" value="PROTOPORPHYRINOGEN OXIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR42923:SF3">
        <property type="entry name" value="PROTOPORPHYRINOGEN OXIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01593">
        <property type="entry name" value="Amino_oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54373">
        <property type="entry name" value="FAD-linked reductases, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51905">
        <property type="entry name" value="FAD/NAD(P)-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51257">
        <property type="entry name" value="PROKAR_LIPOPROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 17" id="KW-0274">FAD</keyword>
    <keyword evidence="6 17" id="KW-0285">Flavoprotein</keyword>
    <keyword evidence="11 17" id="KW-0350">Heme biosynthesis</keyword>
    <keyword evidence="12" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="7" id="KW-0999">Mitochondrion inner membrane</keyword>
    <keyword evidence="9 17" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="13 17" id="KW-0627">Porphyrin biosynthesis</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Amine oxidase" evidence="18">
        <location>
            <begin position="12"/>
            <end position="472"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00002600"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004137"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005073"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00010551"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012867"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022630"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022792"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022827"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023133"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023244"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00024380"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00044160"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00047554"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="RuleBase" id="RU367069"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF01593"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001191.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="476" mass="51492" checksum="DABED82066E2C15D" modified="2019-05-08" version="1">MQRTVAVVGGGISGLTACYHLARSPNAPKIILLEGSGRLGGWLHSTRTEDGAIFEHGPRGIRPAGTVGKNTLLMVSELGLDADVLPVPGDHPASKNRYLYVSRALHRLPSGIGGVLRTVPPFTRPLLWSGLRELAAPRGTQPDETIHSFVSRRFGKELADIAIDSLCRGVFAGDCRALSVRSCFPALFEAEQRRRSVILGMVLGGGKGPALDSALSRRACDERWSQWSLRGGMETLPEALQDFLRQRGVEMHCHARLRRLERTAAECWQITLEDGSVRADHVISAIPARALAVLLPGWAEPLAQELRSIEAVSVGVVNLQYNGATLPITGFGHLVPSFEDSSLLGIVYDSVAFPEQNGSCGPSVRLTVMLGGAWFKQGFGDPDTVPQAVLLSRAKEAVRAHLGIVAEPSRTIVKVLKACIPQYTLGHWQRTETMSRYLEQEGLPLSLIGASYQGVSVNDCIYSAKMAVTRLLDQAC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHY6</accession>
    <name>A0A452GHY6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4 11">Protoporphyrinogen oxidase</fullName>
            <ecNumber evidence="4 11">1.3.3.4</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1 11">Catalyzes the 6-electron oxidation of protoporphyrinogen-IX to form protoporphyrin-IX.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="10 11">
            <text>protoporphyrinogen IX + 3 O2 = protoporphyrin IX + 3 H2O2</text>
            <dbReference type="Rhea" id="RHEA:25576"/>
            <dbReference type="ChEBI" id="CHEBI:15379"/>
            <dbReference type="ChEBI" id="CHEBI:16240"/>
            <dbReference type="ChEBI" id="CHEBI:57306"/>
            <dbReference type="ChEBI" id="CHEBI:57307"/>
            <dbReference type="EC" id="1.3.3.4"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="11">
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </cofactor>
        <text evidence="11">Binds 1 FAD per subunit.</text>
    </comment>
    <comment type="pathway">
        <text evidence="2 11">Porphyrin-containing compound metabolism; protoporphyrin-IX biosynthesis; protoporphyrin-IX from protoporphyrinogen-IX: step 1/1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="11">Mitochondrion inner membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 11">Belongs to the protoporphyrinogen/coproporphyrinogen oxidase family. Protoporphyrinogen oxidase subfamily.</text>
    </comment>
    <dbReference type="EC" id="1.3.3.4" evidence="4 11"/>
    <dbReference type="AlphaFoldDB" id="A0A452GHY6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001344.1">
        <property type="protein sequence ID" value="ENSGAGP00000001196.1"/>
        <property type="gene ID" value="ENSGAGG00000000960.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00251">
        <property type="reaction ID" value="UER00324"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005743">
        <property type="term" value="C:mitochondrial inner membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004729">
        <property type="term" value="F:oxygen-dependent protoporphyrinogen oxidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006782">
        <property type="term" value="P:protoporphyrinogen IX biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.50.50.60">
        <property type="entry name" value="FAD/NAD(P)-binding domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002937">
        <property type="entry name" value="Amino_oxidase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036188">
        <property type="entry name" value="FAD/NAD-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004572">
        <property type="entry name" value="Protoporphyrinogen_oxidase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050464">
        <property type="entry name" value="Zeta_carotene_desat/Oxidored"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00562">
        <property type="entry name" value="proto_IX_ox"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR42923">
        <property type="entry name" value="PROTOPORPHYRINOGEN OXIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR42923:SF3">
        <property type="entry name" value="PROTOPORPHYRINOGEN OXIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01593">
        <property type="entry name" value="Amino_oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54373">
        <property type="entry name" value="FAD-linked reductases, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51905">
        <property type="entry name" value="FAD/NAD(P)-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51257">
        <property type="entry name" value="PROKAR_LIPOPROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6 11" id="KW-0274">FAD</keyword>
    <keyword evidence="5 11" id="KW-0285">Flavoprotein</keyword>
    <keyword evidence="8 11" id="KW-0350">Heme biosynthesis</keyword>
    <keyword evidence="7 11" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="9 11" id="KW-0627">Porphyrin biosynthesis</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Amine oxidase" evidence="12">
        <location>
            <begin position="12"/>
            <end position="323"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00002600"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005073"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00010551"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00012867"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022630"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022827"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023133"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023244"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00047554"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="RuleBase" id="RU367069"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF01593"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001196.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="438" mass="47527" checksum="72321916F33AA29D" modified="2019-05-08" version="1">MQRTVAVVGGGISGLTACYHLARSPNAPKIILLEGSGRLGGWLHSTRTEDGAIFEHGPRGIRPAGTVGKNTLLMVSELGLDADVLPVPGDHPASKNRYLYVSRALHRLPSGIGGVLRTVPPFTRPLLWSGLRELAAPRGTQPDETIHSFVSRRFGKELADIAIDSLCRGVFAGDCRALSVRSCFPALFEAEQRRRSVILGMVLGGGKGPALDSALSRRACDERWSQWSLRGGMETLPEALQDFLRQRGVEMHCHARLRRLERTAAECWQITLEDGSVRADHVISAIPARALAVLLPGWAEPLAQELRSIEAVSVGVVNLQYNGATLPITVMLGGAWFKQGFGDPDTVPQAVLLSRAKEAVRAHLGIVAEPSRTIVKVLKACIPQYTLGHWQRTETMSRYLEQEGLPLSLIGASYQGVSVNDCIYSAKMAVTRLLDQAC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHY8</accession>
    <name>A0A452GHY8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Glutathione S-transferase C-terminal domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the GSTCD family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHY8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001394.1">
        <property type="protein sequence ID" value="ENSGAGP00000001241.1"/>
        <property type="gene ID" value="ENSGAGG00000001002.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000001406.1">
        <property type="protein sequence ID" value="ENSGAGP00000001253.1"/>
        <property type="gene ID" value="ENSGAGG00000001002.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.150:FF:000125">
        <property type="entry name" value="Glutathione S-transferase C-terminal domain-containing protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.150">
        <property type="entry name" value="Vaccinia Virus protein VP39"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036282">
        <property type="entry name" value="Glutathione-S-Trfase_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025714">
        <property type="entry name" value="Methyltranfer_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029063">
        <property type="entry name" value="SAM-dependent_MTases_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13369">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13369:SF0">
        <property type="entry name" value="GLUTATHIONE S-TRANSFERASE C-TERMINAL DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13679">
        <property type="entry name" value="Methyltransf_32"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47616">
        <property type="entry name" value="GST C-terminal domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53335">
        <property type="entry name" value="S-adenosyl-L-methionine-dependent methyltransferases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Methyltransferase" evidence="5">
        <location>
            <begin position="421"/>
            <end position="543"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008797"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022165"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="Pfam" id="PF13679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001241.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="599" mass="67458" checksum="3AF247E486EFC891" modified="2019-05-08" version="1">MKGGVTEEHLYLAYSHQFEDCIFPLHTSVSLFLLSYCDCKLFKVFLVPASDSTDDQLFIKAQPGDLEVHFISRCQLPLVVQSCCLPAIVEENGKFCRAGLSVVLRHIIQKTYEVDPSKKLVLELLGFKKTCLKACAEVSQWTRLCEISIPQAVESLLNASPDQCQVIPPEVLLLEKKLGEPVRVHNDDKIRRQKLQQQKAGYKTAAPALTKRATKNLSPEEGTIVEIKDQEPASLELKVAFSKLVVQEIPAATNREPFHIRKTKTSDLPPLEHVFAEGLYFTLADIVLLPCIHQFLVFIKKHNKLLDLPLISSWYHRVQEVPGVKSAAAKCNMLFLQFPDLLSSPDEQLRDFYTIPDDLEEEHEDYQFIGGPRPTMTKLMESGIEAKFSLHPCPRWTLDWNSLPAAVSPGEGKMSSDRALRKQQQLNNLVSMVTKLSKPGDVIVDFCSGGGHVGIVLAHMLPSCQVVLIENKELSLVRAKDRSDELGLHNIWFIQANLDYFKGAFNIGVALHACGVATDMVIEHCIKAHAAFVICPCCYGFIQNTVKFTFPRSHQFKEVLSYKEHMILCRFADQTAVQLPPKRRLIEENACDHPFVMEK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHZ0</accession>
    <name>A0A452GHZ0_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GHZ0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001395.1">
        <property type="protein sequence ID" value="ENSGAGP00000001242.1"/>
        <property type="gene ID" value="ENSGAGG00000000985.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="16"/>
            <end position="85"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="127"/>
            <end position="236"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001242.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="248" mass="29491" checksum="F5D76F63A919A466" modified="2019-05-08" version="1">MFKPLSYPSPSVVSICKKNRKYYNILQAELQELEKQMESTLLETKATERQIHQQDDDIETTKYHCESLESQVRSLYAEKIKLKLDTEAAQEEFEMMLARNGAYHEKIMAHKKLYWEAESKMPVMLELAKKQDMVKELKTKKEELMNDLQNPEGQVIKQVQEEITHLIEEVTIVKESINEKKKLLEEEKKVHAKLRKEIEVQNKRCDAILKRLHCQLNKLQLNRRQWHWNIQQMEKKAAELRKCLGVTE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHZ1</accession>
    <name>A0A452GHZ1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">C2H2-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHZ1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001349.1">
        <property type="protein sequence ID" value="ENSGAGP00000001201.1"/>
        <property type="gene ID" value="ENSGAGG00000000943.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002434">
        <property type="entry name" value="Zinc finger protein 1168"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002343">
        <property type="entry name" value="Zinc finger protein 33A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000200">
        <property type="entry name" value="zinc finger protein 510 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23226:SF416">
        <property type="entry name" value="FI01424P"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23226">
        <property type="entry name" value="ZINC FINGER AND SCAN DOMAIN-CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9" id="KW-0539">Nucleus</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="8" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 10" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="74"/>
            <end position="101"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="102"/>
            <end position="129"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="130"/>
            <end position="157"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1"/>
            <end position="65"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="49"/>
            <end position="65"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001201.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="193" mass="21835" checksum="A9B51DA38161EBF7" modified="2019-05-08" version="1">MVVGGVSAQASTHCSGDAMVSENEEQKSQQEDPEQVESHSALLQSSKGDVSRSHEQGTAHEIQHRDLKDTTRPYVCCECRKAFSQSFELTRHQRIHTGERPYGCCECGKSFIQHSHLIRHQRNHTGERPYKCGECGKSFTQSSDLIIHQRMHTGERPYCQGKTCHFSLSDTGRRWGDSEIPSFPITPEMFTPL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHZ9</accession>
    <name>A0A452GHZ9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Reticulon</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="6">Inhibits amyloid precursor protein processing, probably by blocking BACE1 activity.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 7">Endoplasmic reticulum membrane</location>
            <topology evidence="1 7">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHZ9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001350.1">
        <property type="protein sequence ID" value="ENSGAGP00000001202.1"/>
        <property type="gene ID" value="ENSGAGG00000000953.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043005">
        <property type="term" value="C:neuron projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0014069">
        <property type="term" value="C:postsynaptic density"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007420">
        <property type="term" value="P:brain development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071787">
        <property type="term" value="P:endoplasmic reticulum tubular network formation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030182">
        <property type="term" value="P:neuron differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.2480:FF:000001">
        <property type="entry name" value="Reticulon"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.2480">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003388">
        <property type="entry name" value="Reticulon"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046964">
        <property type="entry name" value="RTN1-4"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45799:SF5">
        <property type="entry name" value="RETICULON-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45799">
        <property type="entry name" value="RETICULON-LIKE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02453">
        <property type="entry name" value="Reticulon"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50845">
        <property type="entry name" value="RETICULON"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3 7" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="5 7" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 7" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="30"/>
            <end position="59"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="145"/>
            <end position="166"/>
        </location>
    </feature>
    <feature type="domain" description="Reticulon" evidence="8">
        <location>
            <begin position="16"/>
            <end position="203"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004477"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00037306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU210713"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50845"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001202.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="203" mass="22957" checksum="17E02D38F0204856" modified="2019-05-08" version="1">MATSLQTSFDICPSCAIDLLYWRDIKQTGIVFGSILLLLFSLTQFSVVSVIAYLALAALSATISFRIYKSVLQAVQKTDEGHPFKAYLEMEMSLSQEQIQKYTDCLQLYVNSTVRELRRLFLVQDLVDSLKFAVLMWLLTYVGALFNGLTLMIMAVVSMFTLPVVYDKYQAQIDQYLGLVRTHINTVMAKIQAKIPGAKRKAE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GI75</accession>
    <name>A0A452GI75_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">DNA ligase</fullName>
            <ecNumber evidence="15">6.5.1.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">DNA ligase that seals nicks in double-stranded during DNA repair. Also involved in DNA replication and DNA recombination.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13 15">
            <text>ATP + (deoxyribonucleotide)n-3'-hydroxyl + 5'-phospho-(deoxyribonucleotide)m = (deoxyribonucleotide)n+m + AMP + diphosphate.</text>
            <dbReference type="EC" id="6.5.1.1"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 16">Belongs to the ATP-dependent DNA ligase family.</text>
    </comment>
    <dbReference type="EC" id="6.5.1.1" evidence="15"/>
    <dbReference type="AlphaFoldDB" id="A0A452GI75"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001501.1">
        <property type="protein sequence ID" value="ENSGAGP00000001341.1"/>
        <property type="gene ID" value="ENSGAGG00000000751.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003910">
        <property type="term" value="F:DNA ligase (ATP) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071897">
        <property type="term" value="P:DNA biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006310">
        <property type="term" value="P:DNA recombination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006281">
        <property type="term" value="P:DNA repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903461">
        <property type="term" value="P:Okazaki fragment processing involved in mitotic DNA replication"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07900">
        <property type="entry name" value="Adenylation_DNA_ligase_I_Euk"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd07969">
        <property type="entry name" value="OBF_DNA_ligase_I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.3260.10:FF:000001">
        <property type="entry name" value="DNA ligase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.50.140:FF:000062">
        <property type="entry name" value="DNA ligase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.470.30:FF:000016">
        <property type="entry name" value="DNA ligase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1490.70">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.3260.10">
        <property type="entry name" value="DNA ligase, ATP-dependent, N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.470.30">
        <property type="entry name" value="DNA ligase/mRNA capping enzyme"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.50.140">
        <property type="entry name" value="Nucleic acid-binding proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050191">
        <property type="entry name" value="ATP-dep_DNA_ligase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000977">
        <property type="entry name" value="DNA_ligase_ATP-dep"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012309">
        <property type="entry name" value="DNA_ligase_ATP-dep_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012310">
        <property type="entry name" value="DNA_ligase_ATP-dep_cent"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016059">
        <property type="entry name" value="DNA_ligase_ATP-dep_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012308">
        <property type="entry name" value="DNA_ligase_ATP-dep_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036599">
        <property type="entry name" value="DNA_ligase_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012340">
        <property type="entry name" value="NA-bd_OB-fold"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00574">
        <property type="entry name" value="dnl1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45674:SF4">
        <property type="entry name" value="DNA LIGASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45674">
        <property type="entry name" value="DNA LIGASE 1/3 FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04679">
        <property type="entry name" value="DNA_ligase_A_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01068">
        <property type="entry name" value="DNA_ligase_A_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04675">
        <property type="entry name" value="DNA_ligase_A_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF117018">
        <property type="entry name" value="ATP-dependent DNA ligase DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56091">
        <property type="entry name" value="DNA ligase/mRNA capping enzyme, catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50249">
        <property type="entry name" value="Nucleic acid-binding proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00697">
        <property type="entry name" value="DNA_LIGASE_A1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00333">
        <property type="entry name" value="DNA_LIGASE_A2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50160">
        <property type="entry name" value="DNA_LIGASE_A3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 15" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="12" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="4" id="KW-0132">Cell division</keyword>
    <keyword evidence="7 15" id="KW-0227">DNA damage</keyword>
    <keyword evidence="9 15" id="KW-0233">DNA recombination</keyword>
    <keyword evidence="10 15" id="KW-0234">DNA repair</keyword>
    <keyword evidence="5" id="KW-0235">DNA replication</keyword>
    <keyword evidence="3 15" id="KW-0436">Ligase</keyword>
    <keyword evidence="6 15" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="ATP-dependent DNA ligase family profile" evidence="18">
        <location>
            <begin position="535"/>
            <end position="671"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1"/>
            <end position="50"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="17">
        <location>
            <begin position="1"/>
            <end position="28"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007572"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022598"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022705"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023172"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00034003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00058910"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="RuleBase" id="RU000617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="RuleBase" id="RU004196"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50160"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001341.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="806" mass="89746" checksum="BDE4D98E7853D6C3" modified="2019-05-08" version="1">MQTPQSDISADSLSPSSSHSAGDSPALSDSPAISHAGIPRRRTGKDPRIPANIRPGLHPGLCAHRVSCDSAASPMHALLQLGACLIHAIIVPVATCAGLPSLGVLVQANCWETLRNTSAVGPVYWTWWLPISLWLCDPSSFSKPVVPLDPMEYNPSKSSYHPVQDACWGHGQRVPYLAVARTFERIEEVSARLKNIETLSNLFRSVVALSPSDILPCIYLCLNRLGPAYKGLELGIGETILMKAVAQAMGRQLDQIKAEAQEKGDLGLVAESSRSNQRTMFTPPKLSAAGVFSKLQEIARMTGSASMNKKIDIIKGLFVACRHSEARYITRSLGGKLRIGLAEQSVLSAIAHAVSLTPPGQGFPPEVLDAGQSKSAEARKAWLEEQTQILKQTFCELPSYDIIVPILLEHGVESLPQHCKITPGIPLKPMLAQPTKGIGEVLKRFEEAAFTCEYKYDGERAQIHILENGDVHIYSRNQEDNTSKYPDIISRIPKVKKDSVQSCILDAEAVAWDPEKRQIQPFQVLTTRKRKDVEAAEIKVQVCLYAFDILYLNGESLVKEPFSRRRQLLHESFVETEGEFLFATSMDTCSLDEISEFLDQSIKDSCEGLMVKTLEVDATYEIAKRSHKWLKLKKDYLEGVGDTLDLVVIGAYLGKGKRVGMYGGFLLACYDEESEEYQSICKIGTGFTEESLEKHYSFLKDHVLEKPRPYYRWDSSTEPNHWLAAVQVWEVKCADLSISPVHKAAIGLVDEEKGISLRFPRFLRVRDDKKPEEATTSSQVAELYRRQQQIQNQQLVEKAEEEEDFY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIA1</accession>
    <name>A0A452GIA1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2 11">Zinc finger Ran-binding domain-containing protein 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="11">May interfere with constitutive 5'-splice site selection.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 11">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="10 11">Belongs to the ZRANB2 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIA1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001474.1">
        <property type="protein sequence ID" value="ENSGAGP00000001315.1"/>
        <property type="gene ID" value="ENSGAGG00000001059.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001530">
        <property type="term" value="F:lipopolysaccharide binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006396">
        <property type="term" value="P:RNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.1060.10:FF:000004">
        <property type="entry name" value="Zinc finger Ran-binding domain-containing protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.1060.10">
        <property type="entry name" value="Zinc finger, RanBP2-type"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001876">
        <property type="entry name" value="Znf_RanBP2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036443">
        <property type="entry name" value="Znf_RanBP2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017337">
        <property type="entry name" value="ZRANB2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12999:SF17">
        <property type="entry name" value="ZINC FINGER RAN-BINDING DOMAIN-CONTAINING PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12999">
        <property type="entry name" value="ZINC FINGER RAN-BINDING DOMAIN-CONTAINING PROTEIN 2 ZRANB2-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00641">
        <property type="entry name" value="Zn_ribbon_RanBP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF037956">
        <property type="entry name" value="UCP037956_ZnF_Ran"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00547">
        <property type="entry name" value="ZnF_RBZ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90209">
        <property type="entry name" value="Ran binding protein zinc finger-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01358">
        <property type="entry name" value="ZF_RANBP2_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50199">
        <property type="entry name" value="ZF_RANBP2_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 11" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9 11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="8 11" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="7 11" id="KW-0862">Zinc</keyword>
    <keyword evidence="6 12" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="RanBP2-type" evidence="14">
        <location>
            <begin position="57"/>
            <end position="86"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="109"/>
            <end position="326"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="13">
        <location>
            <begin position="142"/>
            <end position="156"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="13">
        <location>
            <begin position="188"/>
            <end position="202"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="13">
        <location>
            <begin position="203"/>
            <end position="216"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="13">
        <location>
            <begin position="245"/>
            <end position="267"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="13">
        <location>
            <begin position="268"/>
            <end position="277"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="13">
        <location>
            <begin position="294"/>
            <end position="310"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="13">
        <location>
            <begin position="311"/>
            <end position="326"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00017543"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00025731"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PIRNR" id="PIRNR037956"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00322"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50199"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001315.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="326" mass="36803" checksum="20DA257C3F8E868F" modified="2019-05-08" version="1">MAPLDALASLRCGNVNFARRTSCNRCGREKTTEAKMMKAGGTEIGKTLAEKSRGLFSANDWQCKTCGNVNWARRSECNMCNTPKYAKLEERTGYGGGFNERENVEYIEREESDGEYDEFGRKKKKYRGKPVGPASILKEVEDKESEGEEEEDEDEDLSKYKLDEDEDEDEADLSKYNLDASEEEDANKKKKSNRRSRSKSRSSHSQSSSRSSSHSSSRSRSRSHSRSSSSSRSRSRSTSREHSRSRGSKSRSSSRSHRGSSTPRKRSYSSSRSSSSPERGKKRSRSRSSSPGDRKKRRSRSRSPERRRRSSSGSSHSGSRTSSKKK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIA2</accession>
    <name>A0A452GIA2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Peptidase S1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GIA2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001549.1">
        <property type="protein sequence ID" value="ENSGAGP00000001374.1"/>
        <property type="gene ID" value="ENSGAGG00000000827.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00190">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.10:FF:000005">
        <property type="entry name" value="Serine protease 37"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.10.10">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009003">
        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043504">
        <property type="entry name" value="Peptidase_S1_PA_chymotrypsin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001314">
        <property type="entry name" value="Peptidase_S1A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001254">
        <property type="entry name" value="Trypsin_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271:SF81">
        <property type="entry name" value="GRANZYME B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271">
        <property type="entry name" value="KALLIKREIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00089">
        <property type="entry name" value="Trypsin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00722">
        <property type="entry name" value="CHYMOTRYPSIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00020">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50494">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50240">
        <property type="entry name" value="TRYPSIN_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1 4" id="KW-0732">Signal</keyword>
    <keyword evidence="2" id="KW-0865">Zymogen</keyword>
    <feature type="signal peptide" evidence="4">
        <location>
            <begin position="1"/>
            <end position="19"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019441759" description="Peptidase S1 domain-containing protein" evidence="4">
        <location>
            <begin position="20"/>
            <end position="246"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase S1" evidence="5">
        <location>
            <begin position="22"/>
            <end position="244"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023145"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001374.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="246" mass="26937" checksum="D7CF33830E97540B" modified="2019-05-08" version="1">MLLLLLLPTAFLLPLGAQAGDIIGGREARPHSRPYMAFVETKMGPESTTECGGFLVREDFVLTACCSAGSGDAPTVVILGAHDLNRKERTQQEIQVRRKIPHEDFDRTTLENDIMLLQVVSLTTVSHVLKPIRLPPANFLLKPEATCSVAGWGRTGVHMVRPATKLQEVTLKMADDKQCELFFDSVHGLHGEHQNEAVIGDTGGPLVCNGFAQSIVSYGKDSGLPSAVYTRMSAFIPWLRKKMGRE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIA3</accession>
    <name>A0A452GIA3_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="11">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1 7">Belongs to the peptidase M16 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIA3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001548.1">
        <property type="protein sequence ID" value="ENSGAGP00000001373.1"/>
        <property type="gene ID" value="ENSGAGG00000000237.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005782">
        <property type="term" value="C:peroxisomal matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004222">
        <property type="term" value="F:metalloendopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050435">
        <property type="term" value="P:amyloid-beta metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042447">
        <property type="term" value="P:hormone catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043171">
        <property type="term" value="P:peptide catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051603">
        <property type="term" value="P:proteolysis involved in protein catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.830.10:FF:000004">
        <property type="entry name" value="Putative insulin-degrading enzyme"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.830.10:FF:000006">
        <property type="entry name" value="Putative insulin-degrading enzyme"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.830.10">
        <property type="entry name" value="Metalloenzyme, LuxS/M16 peptidase-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011249">
        <property type="entry name" value="Metalloenz_LuxS/M16"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011765">
        <property type="entry name" value="Pept_M16_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001431">
        <property type="entry name" value="Pept_M16_Zn_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050626">
        <property type="entry name" value="Peptidase_M16"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007863">
        <property type="entry name" value="Peptidase_M16_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032632">
        <property type="entry name" value="Peptidase_M16_M"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43690:SF18">
        <property type="entry name" value="INSULIN-DEGRADING ENZYME-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43690">
        <property type="entry name" value="NARDILYSIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00675">
        <property type="entry name" value="Peptidase_M16"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05193">
        <property type="entry name" value="Peptidase_M16_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16187">
        <property type="entry name" value="Peptidase_M16_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF63411">
        <property type="entry name" value="LuxS/MPP-like metallohydrolase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00143">
        <property type="entry name" value="INSULINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="6" id="KW-0482">Metalloprotease</keyword>
    <keyword evidence="2" id="KW-0645">Protease</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="Peptidase M16 N-terminal" evidence="8">
        <location>
            <begin position="73"/>
            <end position="207"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase M16 C-terminal" evidence="9">
        <location>
            <begin position="234"/>
            <end position="412"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase M16 middle/third" evidence="10">
        <location>
            <begin position="418"/>
            <end position="548"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007261"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023049"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU004447"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF00675"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF05193"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="Pfam" id="PF16187"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001373.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="569" mass="65585" checksum="CE99F56D7553B8A0" modified="2019-05-08" version="1">LPCLCFPICKTELTHSFRKEFEFLLFPSRLQKNTHNKMSNPAIKRLANKIIKSPEDKREYRGLELENGIKAMLICDPTTDKSSAALDVHIGSLSDPLNIAGLSHFCEHMLFLGTKKYPKENEYSQFLSEHAGSSNAFTSGEHTNYYFDVSHEHLEGALDRFAQFFLCPLFDESCKEREVNAVDSEHEKNLMNDAWRLFQLEKATGNPNHPFSKFGTGNKFTLETRPTKEGIDVRQELLKFHSTYYSSNLMAICVLGRESLDELTGLVVKLFSEVENKNVPLPEFPEHPFQEQHLRQLYKVVPIKDIRNLYVTFPIPDLQKYYKSNPGHYLGHLIGHEGPGSLLSELKAKGWVNTLVGGQKEGARGFMFFIINVDLTEEGLLHVEDIILHMFQYIQKLRADGPQEWVFQECKDLNAVAFRFKDKERPRGYTSKLGGILHYYPIEEVLAAEYLLEEFRPDLIEMVLDKLRPENVRVAIVSKSFEGKTDRTEEWYGTQYKQEAISDEVIQKWQNADLNGKFKLPMKNEFIPTNFEILPLEKDAPQYPALIKVRICITYIKWKLCQPASLLEG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIA5</accession>
    <name>A0A452GIA5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="30">Thromboxane-A synthase</fullName>
            <ecNumber evidence="7">4.2.1.152</ecNumber>
            <ecNumber evidence="29">5.3.99.5</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="31">Cytochrome P450 5A1</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="25">Hydroperoxy icosatetraenoate dehydratase</fullName>
        </alternativeName>
    </protein>
    <organism evidence="36 37">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="37" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="36" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="36" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="32">Catalyzes the conversion of prostaglandin H2 (PGH2) to thromboxane A2 (TXA2), a potent inducer of blood vessel constriction and platelet aggregation. Also cleaves PGH2 to 12-hydroxy-heptadecatrienoicacid (12-HHT) and malondialdehyde, which is known to act as a mediator of DNA damage. 12-HHT and malondialdehyde are formed stoichiometrically in the same amounts as TXA2. Additionally, displays dehydratase activity, toward (15S)-hydroperoxy-(5Z,8Z,11Z,13E)-eicosatetraenoate (15(S)-HPETE) producing 15-KETE and 15-HETE.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="26">
            <text>(15S)-hydroperoxy-(5Z,8Z,11Z,13E)-eicosatetraenoate + AH2 = (15S)-hydroxy-(5Z,8Z,11Z,13E)-eicosatetraenoate + A + H2O</text>
            <dbReference type="Rhea" id="RHEA:48856"/>
            <dbReference type="ChEBI" id="CHEBI:13193"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:17499"/>
            <dbReference type="ChEBI" id="CHEBI:57409"/>
            <dbReference type="ChEBI" id="CHEBI:57446"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="26">
            <dbReference type="Rhea" id="RHEA:48857"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>(15S)-hydroperoxy-(5Z,8Z,11Z,13E)-eicosatetraenoate = 15-oxo-(5Z,8Z,11Z,13E)-eicosatetraenoate + H2O</text>
            <dbReference type="Rhea" id="RHEA:48636"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:57410"/>
            <dbReference type="ChEBI" id="CHEBI:57446"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="1">
            <dbReference type="Rhea" id="RHEA:48637"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="2">
            <text>a hydroperoxyeicosatetraenoate = an oxoeicosatetraenoate + H2O</text>
            <dbReference type="Rhea" id="RHEA:55556"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:59720"/>
            <dbReference type="ChEBI" id="CHEBI:131859"/>
            <dbReference type="EC" id="4.2.1.152"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="2">
            <dbReference type="Rhea" id="RHEA:55557"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="27">
            <text>prostaglandin H2 = (12S)-hydroxy-(5Z,8E,10E)-heptadecatrienoate + malonaldehyde</text>
            <dbReference type="Rhea" id="RHEA:48644"/>
            <dbReference type="ChEBI" id="CHEBI:57405"/>
            <dbReference type="ChEBI" id="CHEBI:90694"/>
            <dbReference type="ChEBI" id="CHEBI:566274"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="28">
            <text>prostaglandin H2 = thromboxane A2</text>
            <dbReference type="Rhea" id="RHEA:17137"/>
            <dbReference type="ChEBI" id="CHEBI:57405"/>
            <dbReference type="ChEBI" id="CHEBI:57445"/>
            <dbReference type="EC" id="5.3.99.5"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="28">
            <dbReference type="Rhea" id="RHEA:17138"/>
        </physiologicalReaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="3 33">
            <name>heme</name>
            <dbReference type="ChEBI" id="CHEBI:30413"/>
        </cofactor>
    </comment>
    <comment type="subunit">
        <text evidence="6">Monomer.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="4">Endoplasmic reticulum membrane</location>
            <topology evidence="4">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5 34">Belongs to the cytochrome P450 family.</text>
    </comment>
    <dbReference type="EC" id="4.2.1.152" evidence="7"/>
    <dbReference type="EC" id="5.3.99.5" evidence="29"/>
    <dbReference type="AlphaFoldDB" id="A0A452GIA5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001557.1">
        <property type="protein sequence ID" value="ENSGAGP00000001381.1"/>
        <property type="gene ID" value="ENSGAGG00000001072.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0020037">
        <property type="term" value="F:heme binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0106256">
        <property type="term" value="F:hydroperoxy icosatetraenoate dehydratase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005506">
        <property type="term" value="F:iron ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016705">
        <property type="term" value="F:oxidoreductase activity, acting on paired donors, with incorporation or reduction of molecular oxygen"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008395">
        <property type="term" value="F:steroid hydroxylase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004796">
        <property type="term" value="F:thromboxane-A synthase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001516">
        <property type="term" value="P:prostaglandin biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.630.10:FF:000003">
        <property type="entry name" value="cytochrome P450 3A12-like isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.630.10">
        <property type="entry name" value="Cytochrome P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001128">
        <property type="entry name" value="Cyt_P450"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017972">
        <property type="entry name" value="Cyt_P450_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002401">
        <property type="entry name" value="Cyt_P450_E_grp-I"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036396">
        <property type="entry name" value="Cyt_P450_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050705">
        <property type="entry name" value="Cytochrome_P450_3A"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24302:SF47">
        <property type="entry name" value="CYTOCHROME P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24302">
        <property type="entry name" value="CYTOCHROME P450 FAMILY 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00067">
        <property type="entry name" value="p450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00463">
        <property type="entry name" value="EP450I"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00385">
        <property type="entry name" value="P450"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48264">
        <property type="entry name" value="Cytochrome P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00086">
        <property type="entry name" value="CYTOCHROME_P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="14" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="22" id="KW-0275">Fatty acid biosynthesis</keyword>
    <keyword evidence="15" id="KW-0276">Fatty acid metabolism</keyword>
    <keyword evidence="11 33 34" id="KW-0349">Heme</keyword>
    <keyword evidence="18 33 34" id="KW-0408">Iron</keyword>
    <keyword evidence="23" id="KW-0413">Isomerase</keyword>
    <keyword evidence="9" id="KW-0444">Lipid biosynthesis</keyword>
    <keyword evidence="20" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="24" id="KW-0456">Lyase</keyword>
    <keyword evidence="21 35" id="KW-0472">Membrane</keyword>
    <keyword evidence="13 33 34" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="19 34" id="KW-0503">Monooxygenase</keyword>
    <keyword evidence="17 34" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="10" id="KW-0643">Prostaglandin biosynthesis</keyword>
    <keyword evidence="8" id="KW-0644">Prostaglandin metabolism</keyword>
    <keyword evidence="37" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="12 35" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="16 35" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="35">
        <location>
            <begin position="217"/>
            <end position="235"/>
        </location>
    </feature>
    <feature type="binding site" description="axial binding residue" evidence="33">
        <location>
            <position position="474"/>
        </location>
        <ligand>
            <name>heme</name>
            <dbReference type="ChEBI" id="CHEBI:30413"/>
        </ligand>
        <ligandPart>
            <name>Fe</name>
            <dbReference type="ChEBI" id="CHEBI:18248"/>
        </ligandPart>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001143"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00001719"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00001971"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004477"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00010617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00011245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00013084"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022501"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022516"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022585"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022832"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023033"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00023160"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00023235"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00023239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00033404"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00036380"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="ARBA" id="ARBA00036424"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="ARBA" id="ARBA00036475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="ARBA" id="ARBA00038872"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="ARBA" id="ARBA00040834"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="31">
        <source>
            <dbReference type="ARBA" id="ARBA00042726"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="32">
        <source>
            <dbReference type="ARBA" id="ARBA00054825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="33">
        <source>
            <dbReference type="PIRSR" id="PIRSR602401-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="34">
        <source>
            <dbReference type="RuleBase" id="RU000461"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="35">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="36">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001381.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="37">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="528" mass="60251" checksum="35861DD28B8109B6" modified="2019-05-08" version="1">GFGFLPWAPENLTVTLLDGPPEICSRYSTSSFSRLKRVGIRHPAPLPFIGNLLFFRQGFWENHTKLITEYGPVCGYYIGRRMYVLVSEPEMIKHILVKDFSNFTNRMTPGLVTKPVIDSILCLRDQRWKEVRSILTPAFSSAKMKEMTPLINQACDVLLSNLKVYADSGKAFDIQRSYGCFTLDVVASVAFGTHVDSQKNPDDTFVKNCKKFFEFSMLRPILILIIAFPFLMIPLSRILPNKRRDEMNGFFIKAIKNMIALRDQQDANERRRDFLQLILDARSSTNYISVEHFDIVNQADVSIQDNKPSVDKALPKKVQKTLSEDEIVGQAFIFLIGGYETTTNALSFATYLLATNPECQEKLLQEIDEFSGKHDVPDYQNIQELPYLDMVIAETLRMYPPAFRFTRETSKDCVVMGQHIPAGAVIEIAVGPLHYNPEFWPEPEKFIPERFTMEAKLQQHPFAYLPFGAGPRSCIGMRMALLEIKMTLLRILQKFRFETSPKTQIPLQLTSVSSLAPKNGVYVKIVPR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GIA8</accession>
    <name>A0A452GIA8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Exocyst complex component</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="4">Component of the exocyst complex involved in the docking of exocytic vesicles with fusion sites on the plasma membrane.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1 4">Belongs to the SEC15 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GIA8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001470.1">
        <property type="protein sequence ID" value="ENSGAGP00000001311.1"/>
        <property type="gene ID" value="ENSGAGG00000001025.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000145">
        <property type="term" value="C:exocyst"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006893">
        <property type="term" value="P:Golgi to plasma membrane transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006886">
        <property type="term" value="P:intracellular protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090522">
        <property type="term" value="P:vesicle tethering involved in exocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.357.30:FF:000001">
        <property type="entry name" value="Exocyst complex component"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.670:FF:000001">
        <property type="entry name" value="Exocyst complex component"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.670">
        <property type="entry name" value="Dsl1p vesicle tethering complex, Tip20p subunit, domain D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.357.30">
        <property type="entry name" value="Exocyst complex subunit Sec15 C-terminal domain, N-terminal subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007225">
        <property type="entry name" value="EXOC6/Sec15"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046361">
        <property type="entry name" value="EXOC6/Sec15_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042045">
        <property type="entry name" value="EXOC6/Sec15_C_dom1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048359">
        <property type="entry name" value="EXOC6_Sec15_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042044">
        <property type="entry name" value="EXOC6PINT-1/Sec15/Tip20_C_dom2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12702:SF2">
        <property type="entry name" value="EXOCYST COMPLEX COMPONENT 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12702">
        <property type="entry name" value="SEC15"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20651">
        <property type="entry name" value="EXOC6_Sec15_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04091">
        <property type="entry name" value="Sec15_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF025007">
        <property type="entry name" value="Sec15"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 4" id="KW-0268">Exocytosis</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 4" id="KW-0813">Transport</keyword>
    <feature type="domain" description="Exocyst complex component EXOC6/Sec15 N-terminal" evidence="6">
        <location>
            <begin position="44"/>
            <end position="213"/>
        </location>
    </feature>
    <feature type="domain" description="Exocyst complex subunit EXOC6/Sec15 C-terminal" evidence="5">
        <location>
            <begin position="407"/>
            <end position="765"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007944"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PIRNR" id="PIRNR025007"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="Pfam" id="PF04091"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="Pfam" id="PF20651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001311.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="808" mass="93723" checksum="4352C7033A72DA18" modified="2019-05-08" version="1">MAESLAEHERILQEIESTDTACVGPTLRSVYDDQPNAHQRFMEKLDACIRNHDREIEKMCNFHHQGFVDAITELLKVRADAGKLKVQVTDTNRRLQDAGKEVIAQTEEIIRCRIQQRNITTVVEKLQLCLPVLEMYSKLKEQMNVKRYYSALKTMEQLENTYLPRVSQYRFCQIMIENLPKLREEIKEISMSDLKDFLESIRKHSDKIGETAMKQAQQQKTFNTVLQKQNNVNYGKNMYVINGRVPEPKYEITSKRTLEEEDEHEEEVLTAHDLVDFSPVYRCLHIYSVLGDGETFENYYRKQRRKQARLVLQPQSSMHETVEGYRRYFSQIVGFFVVEDHILHVTQGLVTRAYTDELWNMALSKIIAVLRTHSSYCTDPDLVLELKNLIVVFADTLQGYGFPVNRLFDLLFEIRDQYNETLLKKWTGLFRDIFEADNYSPIPVANEEEYKIVISKFPFQDPELDKQSFPKKLPMSQSVPQIYIQVKEFIYASLKFSESLHRSSTEIDDMLRKSTNLLLTRTLSSCLQNLIKKPHIGLTELVQIIINTTHLEQACKYLEDFITNITNISQETVHTARLYGLSTFKDARHAAEGEIYTKLNQKIDEFIQLADYDWTMSEPAGRASGYLMDLINFLRSTFQVFTHLPNNNNDHAAMSGKVAQTACMSACQHLSTSLMQMLLDSELKQISMGAVQQFNLDVIQCELFASSEPVPGFHGDTLQLAFIDLRQLLDLFMVWDWSTYLADYGQPTSKYLRVNPSTALTLLEKMKETSKKNNIFAQFRKNDRDKQKLIETVVKQLRSLVNGMSQHT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GID1</accession>
    <name>A0A452GID1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">IPT/TIG domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="11">Forms either a homodimer or a heterodimer with a related family member.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 12">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 12">Belongs to the COE family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GID1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001590.1">
        <property type="protein sequence ID" value="ENSGAGP00000001405.1"/>
        <property type="gene ID" value="ENSGAGG00000001112.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003700">
        <property type="term" value="F:DNA-binding transcription factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045893">
        <property type="term" value="P:positive regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd11606">
        <property type="entry name" value="COE_DBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd01175">
        <property type="entry name" value="IPT_COE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.4280:FF:000001">
        <property type="entry name" value="transcription factor COE1 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.3180:FF:000001">
        <property type="entry name" value="transcription factor COE1 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:001696">
        <property type="entry name" value="Transcription factor COE3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.4280">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.3180">
        <property type="entry name" value="Transcription factor COE1, DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032200">
        <property type="entry name" value="COE_DBD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038173">
        <property type="entry name" value="COE_DBD_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032201">
        <property type="entry name" value="COE_HLH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038006">
        <property type="entry name" value="COE_IPT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014756">
        <property type="entry name" value="Ig_E-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002909">
        <property type="entry name" value="IPT_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003523">
        <property type="entry name" value="Transcription_factor_COE"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018350">
        <property type="entry name" value="Transcription_factor_COE_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10747">
        <property type="entry name" value="TRANSCRIPTION FACTOR COE FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16422">
        <property type="entry name" value="COE1_DBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16423">
        <property type="entry name" value="COE1_HLH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01833">
        <property type="entry name" value="TIG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00429">
        <property type="entry name" value="IPT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81296">
        <property type="entry name" value="E set domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01345">
        <property type="entry name" value="COE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0010">Activator</keyword>
    <keyword evidence="12" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="7 12" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3 12" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10 12" id="KW-0539">Nucleus</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9 12" id="KW-0804">Transcription</keyword>
    <keyword evidence="6 12" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="5 12" id="KW-0862">Zinc</keyword>
    <keyword evidence="4 12" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="IPT/TIG" evidence="14">
        <location>
            <begin position="253"/>
            <end position="337"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="442"/>
            <end position="473"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="557"/>
            <end position="580"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="13">
        <location>
            <begin position="568"/>
            <end position="580"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010340"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00065252"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="RuleBase" id="RU004489"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="SMART" id="SM00429"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001405.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="580" mass="63325" checksum="B8FA3329AD75A118" modified="2019-05-08" version="1">MFGIQENIPRGGTTMKEEPLGSGMNPVRSWMHTAGVVDANTAAQSGVGLARAHFEKQPPSNLRKSNFFHFVLALYDRQGQPVEIERTAFVDFVEKEKEPNNEKTNNGIHYKLQLLYSNGVRTEQDLYVRLIDSMTKQAIVYEGQDKNPEMCRVLLTHEIMCSRCCDKKSCGNRNETPSDPVIIDRFFLKFFLKCNQNCLKNAGNPRDMRRFQVVVSTTVNVDGHVLAVSDNMFVHNNSKHGRRARRLDPSEATPCIKAISPSEGWTTGGATVIIIGDNFFDGLQVVFGTMLVWSELITPHAIRVQTPPRHIPGVVEVTLSYKSKQFCKGAPGRFVYTALNEPTIDYGFQRLQKVIPRHPGDPERLPKEVLLKRAADLVEALYGMPHNNQEIILKRAADIAEALYSVPRNHNQIPTLANTPAHTGMMGVNSFSSQLAVNVSETSQANDQVGYSRNTSSVSPRGYVPSSTPQQSNYNTVSNSMNGYGNAGMPNLGVPGSPGFLNGSSANSPYGIVPSSPTMAASSVTLPSNCSSTHGIFSFSPANVISAVKQKSAFAPVVRPQASPPPSCTSANGNGLQVSP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GID2</accession>
    <name>A0A452GID2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Retinoblastoma-like protein 2</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="15">130 kDa retinoblastoma-associated protein</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="14">Retinoblastoma-related protein 2</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="16">pRb2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="12">Key regulator of entry into cell division. Directly involved in heterochromatin formation by maintaining overall chromatin structure and, in particular, that of constitutive heterochromatin by stabilizing histone methylation. Recruits and targets histone methyltransferases KMT5B and KMT5C, leading to epigenetic transcriptional repression. Controls histone H4 'Lys-20' trimethylation. Probably acts as a transcription repressor by recruiting chromatin-modifying enzymes to promoters. Potent inhibitor of E2F-mediated trans-activation, associates preferentially with E2F5. Binds to cyclins A and E. Binds to and may be involved in the transforming capacity of the adenovirus E1A protein. May act as a tumor suppressor.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the retinoblastoma protein (RB) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GID2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001513.1">
        <property type="protein sequence ID" value="ENSGAGP00000001348.1"/>
        <property type="gene ID" value="ENSGAGG00000001075.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000785">
        <property type="term" value="C:chromatin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005667">
        <property type="term" value="C:transcription regulator complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000977">
        <property type="term" value="F:RNA polymerase II transcription regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006325">
        <property type="term" value="P:chromatin organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:2000134">
        <property type="term" value="P:negative regulation of G1/S transition of mitotic cell cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd20606">
        <property type="entry name" value="CYCLIN_RBL2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00043">
        <property type="entry name" value="CYCLIN_SF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.472.10:FF:000048">
        <property type="entry name" value="Retinoblastoma-like 2, isoform CRA_a"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.472.10:FF:000049">
        <property type="entry name" value="Retinoblastoma-like 2, isoform CRA_a"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.472.140:FF:000001">
        <property type="entry name" value="Retinoblastoma-like 2, isoform CRA_a"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.472.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.472.10">
        <property type="entry name" value="Cyclin-like"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013763">
        <property type="entry name" value="Cyclin-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036915">
        <property type="entry name" value="Cyclin-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002720">
        <property type="entry name" value="RB_A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002719">
        <property type="entry name" value="RB_B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015030">
        <property type="entry name" value="RB_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028309">
        <property type="entry name" value="RB_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024599">
        <property type="entry name" value="RB_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13742">
        <property type="entry name" value="RETINOBLASTOMA-ASSOCIATED PROTEIN RB -RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13742:SF8">
        <property type="entry name" value="RETINOBLASTOMA-LIKE PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11934">
        <property type="entry name" value="DUF3452"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01858">
        <property type="entry name" value="RB_A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01857">
        <property type="entry name" value="RB_B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00385">
        <property type="entry name" value="CYCLIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01367">
        <property type="entry name" value="DUF3452"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01368">
        <property type="entry name" value="RB_A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01369">
        <property type="entry name" value="Rb_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47954">
        <property type="entry name" value="Cyclin-like"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="11" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="5" id="KW-0156">Chromatin regulator</keyword>
    <keyword evidence="7" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="9" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0678">Repressor</keyword>
    <keyword evidence="8" id="KW-0804">Transcription</keyword>
    <keyword evidence="6" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="Retinoblastoma-associated protein N-terminal" evidence="19">
        <location>
            <begin position="79"/>
            <end position="221"/>
        </location>
    </feature>
    <feature type="domain" description="Retinoblastoma-associated protein A-box" evidence="20">
        <location>
            <begin position="399"/>
            <end position="591"/>
        </location>
    </feature>
    <feature type="domain" description="Cyclin-like" evidence="18">
        <location>
            <begin position="833"/>
            <end position="919"/>
        </location>
    </feature>
    <feature type="domain" description="Retinoblastoma-associated protein C-terminal" evidence="21">
        <location>
            <begin position="1006"/>
            <end position="1125"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1"/>
            <end position="25"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="636"/>
            <end position="682"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="793"/>
            <end position="813"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="912"/>
            <end position="985"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="17">
        <location>
            <begin position="1"/>
            <end position="10"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="642"/>
            <end position="660"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="793"/>
            <end position="812"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="17">
        <location>
            <begin position="912"/>
            <end position="925"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="17">
        <location>
            <begin position="927"/>
            <end position="936"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="17">
        <location>
            <begin position="948"/>
            <end position="957"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="17">
        <location>
            <begin position="961"/>
            <end position="971"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022491"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00056129"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00071619"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00076028"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00077146"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00081550"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="SMART" id="SM00385"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="SMART" id="SM01367"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="SMART" id="SM01368"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="SMART" id="SM01369"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001348.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1125" mass="127632" checksum="2C85CDBA8D542783" modified="2019-05-08" version="1">MPGEEEEEAEAGVAGGASSPPAEEWGTRQRYEELCSSLNMDERARAEAWLSYQGMKRNYTLEGNDLHWLACALYVACRKAVPTVSRGTVEGNYVSLTRILRCSEQSLIEFFNKMKKWEDMANLPLQFRERTERLERNFTVSAVIFKKYEPIFQDIFTYPQDDQPRQQRGRKQRRQPCTVSEVFQFCWVLFVHAKGNFPMISDDLVNSYHLLLCALDLVYGNALQCPNRRELLNPAFKGLPEDFHSKDYKPSSDPPCIIEKLCSLHDGLVLDAKGIKEHFWKPYIRKLFDKKILKGKEENLTGFLDPGSFGDSFKAINKAYEEYVLSVGNLDERIFLGEDADEEIGTLRRCLNTASGMETAERVQVKYNLQQHFDRSKSLRITTPLTGRKYIKESNPNVTPVSIATYNLSRLHTMLTGLKNAPSENLEQILRACSRDPSQSIAKRVKEMYEVYCQCTQSEGEFSNFSKDNAGKHFRLAEVLYYKVLESVIEQERRRLGDTDLSAILEQDVFHRSLLACCLEIITFSYKPPGTFPFITEIFDVPVYHFYKVIEIFIRAEDGLCREVVKHLNHIEEQILESMAWKCESPLWDRIRDNENKVPTCGEVMPPHYFERSNGNSLVGSPLTPRRINEVRAETGGLGKGLSSSPTTLYDRYSSPTTNPTRRRLFVDNDNPPDSGTPVRVSQQPVVNTVPVQNVTPETMSVTPVPGQTLVTVATATVTANNGQTVTIPVQGIANENGGITFFPVQVNVGGQAQAVSGSIQPLSAQTLAGGTLNTQMSGTTLQLPGQVAVQQISPSGDQQRQNQQFSTTTTSKPRKTGSLYLFFRKVYHLASVRLRDLCVKLDISDELRKKIWTCFEFSLVQCPELMMDRHLDQLLMCAIYVMAKVTKEDKSFQNIMRCYRTQPQARSHVYRTVRIKGRRQRRRSGSSDSSIQQNSPTERSKERTSRDSSPVMRSSSTLPVPQPNSAPPTPTRLTGANSDIEEEEERGDLIRFYNNIYIEQMKEFALKYSSTNATDAPPLSPYPFVRIGSPRRVQLSQNHSIYISPHKNESTLSPREKIFYYFSSSPSKRLSEINSMIRSGETPTKKRGILLEDGTESPAKRICQENHTALLRRLQDVANDRGSH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GID6</accession>
    <name>A0A452GID6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Synaptotagmin-like mitochondrial and lipid-binding domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GID6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001507.1">
        <property type="protein sequence ID" value="ENSGAGP00000001344.1"/>
        <property type="gene ID" value="ENSGAGG00000001064.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098592">
        <property type="term" value="C:cytoplasmic side of apical plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035091">
        <property type="term" value="F:phosphatidylinositol binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008526">
        <property type="term" value="F:phosphatidylinositol transfer activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035774">
        <property type="term" value="P:positive regulation of insulin secretion involved in cellular response to glucose stimulus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039934">
        <property type="entry name" value="C2CD2/C2CD2L"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040885">
        <property type="entry name" value="SMP_C2CD2L"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21119">
        <property type="entry name" value="C2 DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21119:SF8">
        <property type="entry name" value="PHOSPHOLIPID TRANSFER PROTEIN C2CD2L"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18696">
        <property type="entry name" value="SMP_C2CD2L"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49562">
        <property type="entry name" value="C2 domain (Calcium/lipid-binding domain, CaLB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Synaptotagmin-like mitochondrial and lipid-binding" evidence="2">
        <location>
            <begin position="90"/>
            <end position="178"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="530"/>
            <end position="585"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="606"/>
            <end position="637"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="648"/>
            <end position="667"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="557"/>
            <end position="569"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF18696"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001344.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="667" mass="72223" checksum="0022B0C5E36D9BE0" modified="2019-05-08" version="1">MELELGWAALVLLFAASLLTVSAWLLQYSRGVWSGRGKPETSGGALREAGIWKSLLRLRVARDGAPEEAVAGARGLLASLFAFRSFRENWQRAWVRALNEQACRHGSSIQITFEESPQLPPAVNISHVTCTDQADHSMVLHCHLSADAVKFPVSVTQQSPAAVAMDTYHVTLALLQAQKNDGKVDLSTIKDLIEDSVISTQPAMMVNLKAWTAGSNMMSCEKLSQESSSNIVASTVSKLLLRHMQVLNLNCEKEGGGGKLCCVAELDSPLQQKWTKPARASISSAAAAVEWNEELMLELGPRSKELKLRVLGNSSGGENVLLGYTAVSLDSCTKQPSGRLVYSLSPGAGQLLTPAATLTLELLYHESPLSPNPQYATSLRTSITPTKKIEMDRTIMPDGTIVTTVTTVQSRPKVDCKLDSPSRSPSKVEVTEKMTTRLSESSCPSTALASNSWDSHMSNGLDPVAETAIRQLTETNNKSAKKTPTKRSTLIISGVSKVPIAQDEMALSLGYAASMEATMHGYSIMAGMCDHPQSSTEPTQLLEASFSGQRASQELDETTRSDISERPSMEDVESETGSTGALETRSLKDHKVSFLRSGTKLIFRRKNKQKEAGLSQSHDDLSNASTSSATRKKASSFSRRLIKRFSFKSKSKPKASGSTTSMGENYL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GID7</accession>
    <name>A0A452GID7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">non-specific serine/threonine protein kinase</fullName>
            <ecNumber evidence="2">2.7.11.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="9">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="8">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the protein kinase superfamily. STE Ser/Thr protein kinase family. STE20 subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452GID7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001595.1">
        <property type="protein sequence ID" value="ENSGAGP00000001410.1"/>
        <property type="gene ID" value="ENSGAGG00000001107.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.4270:FF:000001">
        <property type="entry name" value="Serine/threonine-protein kinase 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000410">
        <property type="entry name" value="Serine/threonine-protein kinase 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.4270">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050629">
        <property type="entry name" value="STE20/SPS1-PAK"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48012:SF10">
        <property type="entry name" value="FI20177P1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48012">
        <property type="entry name" value="STERILE20-LIKE KINASE, ISOFORM B-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7 10" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="6" id="KW-0418">Kinase</keyword>
    <keyword evidence="5 10" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="4" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Protein kinase" evidence="12">
        <location>
            <begin position="27"/>
            <end position="348"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="190"/>
            <end position="235"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="260"/>
            <end position="282"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="288"/>
            <end position="307"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="11">
        <location>
            <begin position="198"/>
            <end position="210"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="215"/>
            <end position="235"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="271"/>
            <end position="280"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="288"/>
            <end position="298"/>
        </location>
    </feature>
    <feature type="binding site" evidence="10">
        <location>
            <position position="56"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00008874"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00012513"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00047899"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00048679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001410.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="348" mass="39981" checksum="0282B8F74508C7A1" modified="2019-05-08" version="1">LLQPPAPLHKLKKLSEDSLTKQPEEVFDVLEKLGEGSYGSVFKAIHKESGQVVAIKQVPVESDLQEIIKEISIMQQCDSPYVVKYYGSYFKNTDLWIVMEYCGAGSVSDIIRLRNKTAIFMIPTNPPPTFRKPELWSDEFTDFVKKCLVKNPEQRATATQLLQHTFIKNAKPVSILRDLITEAMEIKAKRHEEQQRELEEEEENSEEDELDSHTMVKTNSESAGTMRATSTMSESAQTMIEHNSTMLESDLGTMVINSDDDEEEEDGTMKRNATSPQVQRPSFMDYFDKQDSKNKSHENCNQNMPEPYHISKYPDNWKVPQDGDFDFVSKFLMFTSLHAVYFMYAKIN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GID8</accession>
    <name>A0A452GID8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">DH domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GID8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001522.1">
        <property type="protein sequence ID" value="ENSGAGP00000001353.1"/>
        <property type="gene ID" value="ENSGAGG00000001047.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.900.10">
        <property type="entry name" value="Dbl homology (DH) domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035899">
        <property type="entry name" value="DBL_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000219">
        <property type="entry name" value="DH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051492">
        <property type="entry name" value="Dynamin-Rho_GEF"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22834">
        <property type="entry name" value="NUCLEAR FUSION PROTEIN FUS2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22834:SF17">
        <property type="entry name" value="RHO GUANINE NUCLEOTIDE EXCHANGE FACTOR 38"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00621">
        <property type="entry name" value="RhoGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48065">
        <property type="entry name" value="DBL homology domain (DH-domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50010">
        <property type="entry name" value="DH_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="DH" evidence="2">
        <location>
            <begin position="97"/>
            <end position="173"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="39"/>
            <end position="82"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="39"/>
            <end position="54"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="55"/>
            <end position="68"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="PROSITE" id="PS50010"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001353.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="189" mass="21512" checksum="BE15B7D0FBA002FD" modified="2019-05-08" version="1">METKEAPGKEGMGAKRKNLTFLKHRLYMLERRKTDTVVENSVSVDQGNSGTLRRSQSDRTEYSQKLQEKMAPQAGSLSPVTSLQENEEQLTRRMMAKRVKIIAELVQTEKDYLHDLELCIREVVQPLRHKQIVRFDVDGLFSNIESVHQISAKLLSLLEEATADMDPRMQVIGMFTANYGNASLWNSGV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2025-10-08" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYZ0</accession>
    <name>A0A452GYZ0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">DNA-repair protein Xrcc1 N-terminal domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000008498.1">
        <property type="protein sequence ID" value="ENSGAGP00000007353.1"/>
        <property type="gene ID" value="ENSGAGG00000005905.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000008503.1">
        <property type="protein sequence ID" value="ENSGAGP00000007358.1"/>
        <property type="gene ID" value="ENSGAGG00000005905.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003684">
        <property type="term" value="F:damaged DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006284">
        <property type="term" value="P:base-excision repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000012">
        <property type="term" value="P:single strand break repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.260:FF:000025">
        <property type="entry name" value="DNA repair protein XRCC1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.260">
        <property type="entry name" value="Galactose-binding domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008979">
        <property type="entry name" value="Galactose-bd-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002706">
        <property type="entry name" value="Xrcc1_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11370">
        <property type="entry name" value="DNA-REPAIR PROTEIN XRCC1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11370:SF4">
        <property type="entry name" value="DNA-REPAIR PROTEIN XRCC1 N-TERMINAL DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01834">
        <property type="entry name" value="XRCC1_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49785">
        <property type="entry name" value="Galactose-binding domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="DNA-repair protein Xrcc1 N-terminal" evidence="2">
        <location>
            <begin position="1"/>
            <end position="149"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="189"/>
            <end position="219"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="288"/>
            <end position="316"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="371"/>
            <end position="396"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF01834"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007358.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="396" mass="43514" checksum="91AFF0FF2D749AC7" modified="2019-05-08" version="1">MAPVKINHVVSFTSQDPRYPVENLLRDDGIHPWLNSLQDRSRQLKVELQLERASSIGYIDIGNCGCAFIQVDVGRSFWPLEQPYVTMVPSTTLMTPADSKLDRNRSGVRMFKEGDFLAAALGEKWDRVRITCSQPFNKQAQFGLSFIHIRTPLDPDRSHAAPAPPQTGLEPAASPWLASPAFLRTFFPEVQPSMPRPGTQAGRARSRGGRPSKLDDDVEAGGEMGTCPICAGCFSTDLLPVHASTCGEDGSLPDTAWPSPPWENPQHEETPETWVLCPICEXAPSARSRSAQQRWKGTPAPVGSRQEPWRPPTPGCGWNRADRARPCCRGTTLAYAPLFPHQPRIEVSLCLSLRATLLSWVGTAFGLKGHRRHGDTGTQTPCHGHPGMRSALGQLH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="22" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYZ1</accession>
    <name>A0A452GYZ1_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="1">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="1 2">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="2" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="1" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="1" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GYZ1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008638.1">
        <property type="protein sequence ID" value="ENSGAGP00000007480.1"/>
        <property type="gene ID" value="ENSGAGG00000005968.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021748">
        <property type="entry name" value="DUF3314"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR36292">
        <property type="entry name" value="UPF0575 PROTEIN C19ORF67"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR36292:SF1">
        <property type="entry name" value="UPF0575 PROTEIN C19ORF67"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11771">
        <property type="entry name" value="DUF3314"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000313" key="1">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007480.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="296" mass="33714" checksum="3BD0055A039039C1" modified="2019-05-08" version="1">CFDFSCLVLEHEPLAKSPTHENLALEELLTPVTEKLKYLLKKAEDFQTYLLYSRDRMQKEQFAKAVPTFLQMCQPYFEYLESTARSYNSGLGALQTSVRKRLLEISEQLALRLEQLVLMYASFSFVSLEDTDPFNVSCFFCGRFWLSEWRQLSVFRFCISAPYTAAHLPCNLYKKMRWNLDILEEGVGAGGRRRGYFLCFRDTGRENAVNMQKLWSIGRWVPLDPDAEHSSDVLVLCRQPTGDFQPLLTIGFEEPSHTLATDLLVQILSAPACAPLPCPGTSREPLHWGSSGGQGQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYZ2</accession>
    <name>A0A452GYZ2_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="3">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GYZ2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008532.1">
        <property type="protein sequence ID" value="ENSGAGP00000007384.1"/>
        <property type="gene ID" value="ENSGAGG00000005929.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042319">
        <property type="entry name" value="GINM1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR28549">
        <property type="entry name" value="GLYCOPROTEIN INTEGRAL MEMBRANE PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR28549:SF1">
        <property type="entry name" value="GLYCOPROTEIN INTEGRAL MEMBRANE PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0732">Signal</keyword>
    <keyword evidence="1" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="1" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="2">
        <location>
            <begin position="1"/>
            <end position="25"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019354182" evidence="2">
        <location>
            <begin position="26"/>
            <end position="323"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="1">
        <location>
            <begin position="266"/>
            <end position="285"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007384.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="323" mass="36026" checksum="F67E7EAE601F19A7" modified="2019-05-08" version="1">MEAAPGPGGLLLLLLVLGLLGPAAPQPAAPPQFLSQENIRVNVTMLKTSEESQEAQVVFNITYNSGQIYVNDFPMKRGVTRIKCQTLILETGNSDNLLDQRHWGIVSVRILFREWPLASRSNLQLIVIQEEVTEIDGREVQQEEVTEINILIKGLRVLRHSNHTVPLKESMLYSIPRDNDVLFTLPNLSGKVDVQGPLQPTSHYLLRQAETTVDEETLPGKLPETPLRTEPPSSYKVMCQWVEYLRKELCGFWLTSFPVFFKLMEVIGTGVVGAALVLGILKVLFPSCEHKGVLHLEDINTTAGLVINISSDIPEKTDNLVEK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYZ4</accession>
    <name>A0A452GYZ4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">cyclin-dependent kinase</fullName>
            <ecNumber evidence="3">2.7.11.22</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="13">Galactosyltransferase-associated protein kinase p58/GTA</fullName>
        </alternativeName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="12">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.22"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.22"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the protein kinase superfamily. CMGC Ser/Thr protein kinase family. CDC2/CDKX subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.22" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452GYZ4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008408.1">
        <property type="protein sequence ID" value="ENSGAGP00000007270.1"/>
        <property type="gene ID" value="ENSGAGG00000005840.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004693">
        <property type="term" value="F:cyclin-dependent protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007346">
        <property type="term" value="P:regulation of mitotic cell cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07843">
        <property type="entry name" value="STKc_CDC2L1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000054">
        <property type="entry name" value="Cyclin-dependent kinase 11B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000124">
        <property type="entry name" value="cyclin-dependent kinase 11B isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="CDK"/>
    </dbReference>
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        <property type="entry name" value="CDK11/PITSLRE_STKc"/>
    </dbReference>
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        <property type="entry name" value="Kinase-like_dom_sf"/>
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        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
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        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24056">
        <property type="entry name" value="CELL DIVISION PROTEIN KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24056:SF107">
        <property type="entry name" value="CYCLIN-DEPENDENT KINASE 11A-RELATED"/>
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    </dbReference>
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        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="S_TKc"/>
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        <property type="entry name" value="Protein kinase-like (PK-like)"/>
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    </dbReference>
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        <property type="entry name" value="PROTEIN_KINASE_ST"/>
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    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="10" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="8" id="KW-0418">Kinase</keyword>
    <keyword evidence="7" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="6" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Protein kinase" evidence="15">
        <location>
            <begin position="390"/>
            <end position="675"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="18"/>
            <end position="369"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="685"/>
            <end position="747"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="18"/>
            <end position="60"/>
        </location>
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    <feature type="compositionally biased region" description="Basic residues" evidence="14">
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            <begin position="95"/>
            <end position="113"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
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            <begin position="114"/>
            <end position="131"/>
        </location>
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            <begin position="138"/>
            <end position="227"/>
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    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="238"/>
            <end position="252"/>
        </location>
    </feature>
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        <location>
            <begin position="257"/>
            <end position="292"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="293"/>
            <end position="305"/>
        </location>
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    <feature type="compositionally biased region" description="Acidic residues" evidence="14">
        <location>
            <begin position="306"/>
            <end position="317"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="325"/>
            <end position="334"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006485"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012425"/>
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    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
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    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
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            <dbReference type="ARBA" id="ARBA00047811"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00048367"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00079859"/>
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    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007270.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
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    </evidence>
    <sequence length="747" mass="87540" checksum="B2DF9B81A9DFD239" modified="2019-05-08" version="1">MGDEKDSWKVKTLDEILQEKKRRKEQEEKAEIKRIKNSDDRDSKRDSLEEGELRDHRMEITIRNSPYRREDSMEDRGEEDDSLAIKPPQQTSRKEKAHHRKDEKRKEKRRHRSHSAEGGKHARVKEKEREHERRKRHREEQDKARREWERQKRREMAREHSRRERDRLEQLERKRERERKMREQQKEQRELKERERRAEERRKEREARREVSAHHRTVREDYGDKVKIRTWSRSPLRQQREKLEQGDGRKPESGSGSEEEEEESSSGESEEEEEEEEEEEEEEEEEEEEEETGSNSEEVSEQSAEVSEEEMSEEEERENGNHIPIESRFDRDSAGSEGEEEEVGEETPHSNAMTEGDYVPDSPASSPVELKQELPKYLPALQGCRSVEEFQCLNRIEEGTYGVVYRAKDKKTDEIVALKRLKMEKEKEGFPITSLREINTILKAQHPNIVTVREIVVGSNMDKIYIVMNYVEHDLKSLMETMKQPFLPGEVKTLMIQLLRGVKHLHDNWILHRDLKTSNLLLSHKGILKVGDFGLAREYGSPLKPYTPVVVTLWYRAPELLLGAKEYSTAIDMWSVGCIFGELLTQKPLFPGKSEIDQINKVFKDLGTPSEKIWPGYNELPAVKKMTFTEYPYNNLRKRFGALLSDPGFDLMNKFLTYYPGRRITAEDGLKHEYFRETPLPIDPSMFPTWPAKSEQQRVKRGTSPRPPEGGLGYSQLGDDDLKDTGFHLTTTNQGASAAGPGFSLKF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYZ7</accession>
    <name>A0A452GYZ7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Lysosomal-trafficking regulator</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="8">Interacts with CPAP, LIP8 and ZNF521.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000008483.1">
        <property type="protein sequence ID" value="ENSGAGP00000007339.1"/>
        <property type="gene ID" value="ENSGAGG00000005878.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098542">
        <property type="term" value="P:defense response to other organism"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006955">
        <property type="term" value="P:immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002449">
        <property type="term" value="P:lymphocyte mediated immunity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043473">
        <property type="term" value="P:pigmentation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd06071">
        <property type="entry name" value="Beach"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="PH_BEACH"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Lysosomal trafficking regulator"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Lysosomal trafficking regulator"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="neurobeachin isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1540.10">
        <property type="entry name" value="BEACH domain"/>
        <property type="match status" value="1"/>
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    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
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    </dbReference>
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        <property type="entry name" value="BEACH_dom"/>
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        <property type="entry name" value="BEACH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050865">
        <property type="entry name" value="BEACH_Domain"/>
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        <property type="entry name" value="PH-BEACH_dom"/>
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        <property type="entry name" value="PH-like_dom_sf"/>
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        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
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    <dbReference type="InterPro" id="IPR019775">
        <property type="entry name" value="WD40_repeat_CS"/>
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    <dbReference type="InterPro" id="IPR036322">
        <property type="entry name" value="WD40_repeat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001680">
        <property type="entry name" value="WD40_rpt"/>
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        <property type="entry name" value="BEIGE/BEACH-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="LYSOSOMAL-TRAFFICKING REGULATOR"/>
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    </dbReference>
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        <property type="entry name" value="PH_BEACH"/>
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        <property type="entry name" value="WD40"/>
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    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81837">
        <property type="entry name" value="BEACH domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50978">
        <property type="entry name" value="WD40 repeat-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50197">
        <property type="entry name" value="BEACH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51783">
        <property type="entry name" value="PH_BEACH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00678">
        <property type="entry name" value="WD_REPEATS_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50082">
        <property type="entry name" value="WD_REPEATS_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50294">
        <property type="entry name" value="WD_REPEATS_REGION"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="11" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="7" id="KW-0653">Protein transport</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0813">Transport</keyword>
    <keyword evidence="5 10" id="KW-0853">WD repeat</keyword>
    <feature type="domain" description="BEACH-type PH" evidence="14">
        <location>
            <begin position="3003"/>
            <end position="3109"/>
        </location>
    </feature>
    <feature type="domain" description="BEACH" evidence="13">
        <location>
            <begin position="3114"/>
            <end position="3416"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="10">
        <location>
            <begin position="3606"/>
            <end position="3640"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="156"/>
            <end position="176"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="913"/>
            <end position="932"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1218"/>
            <end position="1247"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="11">
        <location>
            <begin position="2176"/>
            <end position="2205"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="163"/>
            <end position="176"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="12">
        <location>
            <begin position="1218"/>
            <end position="1232"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="1233"/>
            <end position="1244"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022574"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00064710"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00067676"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50197"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS51783"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007339.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="3795" mass="430026" checksum="9A02AF11C2C3C6E3" modified="2019-05-08" version="1">MSSASNSLAREFLIDVNRLCNAVVQRAEAREEEEEETHMAALGQYLVQGRGFTLLTTLNSNMDQELTCREELLTLLLSLLPLVWKIPVQEEKAINFNLPFTVDVCLTKETNSNSLRSAHEKQNLEGIVHPSFQTPVKLNSSFRKIRRQRKITHRYSVRDARKSQLSTSDSEGNSDNKTTIMIKHRRSQILQPFVTHLPRDDYHAEVVRSFNPSPISLENSRQIIPTQESKSEVLNESAAAAVVSSMDNSPFDLCHVLLSLLEKVCKFDITLNHNSALAATIVPSLTEFLSGFGDCCNVSGDAESEVVSAGWTEEPVALVQRMLFRTVLHLLSIDITNAETMPDNLKKNLTDLLRAALKIRTCLEKQSDPFVPRHKKTLQEVQDNFVFSKYRHRVLLLPELLEGVLQILICCLQSAASNPFYFSQAVDLVHEFIQHQGFKLLEITVLQMEWLVSKDEEITPEASEHLKALISSIMKIISTVKKVKSEQLHQSMCTRKRHRRCEYSHFMHHHRDLSGLPVSAFKNQVSRNPFEEVVDGEVHYPERCCCIAVCAHQCFHLLQQVSLSSTCAQILSGVHNVGICCCMDPKSVITPLLQASKLPVLKNFQQHILSILNKFILDQLGGAEVSQKIIQAACNICTIDCDQLAELEDTLQSNSTDIGLASNFSCPSYRFQGILPSSGSEDMLLKWDALEAYQDFAFEEDKLRSVQIASHICHLIQKGNVVVQWKLYNYIYNPVLQRGVELAHRSQQLGITAVCNHVCSCHSLCLPQEVLQIYLQTLPALFKSRIIQNLFLSCNGVNQMTELNYLDSVRSHSLKVFETLIFCLGDQQKDQAMPELEGMDNEQKESILDLVTSFRSQQMISDVPQSLSKFYAGLRKAYPKKKKCVSQEVHVNLIHLFLCVAFLCVSKEEECDRDSANDSEDTSGYDSTASEPLSHKLPCLSLESLTLPSPEHIHRAADIWSMCRWIYLCSSVFQRQFHRLGGFEVCHRLIIMIIQKLAKNKEKEQEKKERPLSENNKCLNASPLNQPVFKDDSYYLTVNKQLTQSELNSLDENPDASGQPVSECISCDHSLKVEQISVAMVELEELKKSVKQENEWSLQSIRLLEALLTICLHSANSTQQKAEVELLHQSVCLEDVLLEMRDQLAQSRAIETCLAKPLLDSLLRVALGNYSADLDHSEEKTEKILHTEKASVPQPEDLSEEAEESQCCSFKLFAEEEGYEADSESNPDDSESKDEGTDTKREAECMVPSSYSSDLTENISQGELIYPEICMLELKLLSAGNPSLDVLAHVFQSFLKVIRQKKKNATFLMQQGAVKSLLGGFLNILIQKDSSFHACQIVLVELLVSLMSSRTCAEELTLLLRIFLEETPCTEILLKGILKIVETNIYMSPLQYLSFPLLKSPNLNSSSSQKSTSALNSKTVGLLKRAKFSQNKKEADGENLTHQLLSSWHVAPIHLPLVGQNCWPHMSEGFSVSLWFHVECVQETESPTEKGKKMKKRSKPLIVRESSFDGTEGSKSVRMEYLNLGDKLVEDGCIHIISLGSKALMIQVWADLNTGAFIFRMCMDPNDEMKAGLLAQAESPENIFLVGKWEHLAITYLQQPEGKKNIHGELLLWVSGQRKCEISLDYTLPRKTTLSDSNKTFCMIGHCTSSQEEPLQLAGKWDLGNLLLFNGAKIGPEEAFYLYACGPDFTAIMPCKYGKTMTDYSKYITKDILQCERIKEFFMKKKEADVGPLIESLAVVYTTCCPAQYTIYEPVIRLKGQVKTVPSQRPFSSKEVQSNFLDPYHLKMLQPVEYQTLQGILHEIGGTGAFVFLFARVIDITNCEETQALALQVILSLTKYNQQRIQEMENCNGYSMIHQVLIKPKCIVGFHILKTLLEGCFSDEILNINESGQFMLDVDSTAVIQDIKLLEDLLLDWKIWSKAQQGVWETLLAALEILIRANHHQQMFNIKQLLKARVVHHFLLTCQVLQEQKEGHLTSLPREVCRSFVKIIEEVLGSPPDLELLTLIFNFLLAVHPPTNTYVCHNPTNFYFSLHIDGKIFQEKVQSIKYLRNSSIGGKSVDSPSLTTSPAGFTASPLGGSIISKIIIPQMMISQALQRRCKSLPASPTSSPQVHPKRLPGSLGRSIDNLQEVWESDSVPQEKNIFVESSETLKRIQEELFVSSCESAKTVCDSELTSAEALVDISKEELELSENKELNTELKSNENGAAVESLLRRPDNLKGLPSFQKSQNNFAGLGLAFSAQSGSSSVACWPSLADRSALPEDWESLTFSSVSEQTFKRSERPDYRSTEDCLVLICCGLYDLLRGVLLILPDIMLEDVIDKLIQPDTLIVLVNHPSPLIQERVIKLLDAYFNRATREQKEQFLKNRGFSLLANQLYLHQGTQELVECFLEMLFGRSVGLDEEFDREDVKNAGLFQKWCVIPVLGLIENSLYDNVLLHNSFCLLLQIINSCSKMADMLLDNGLLYVLCNTLAALNGLETNIPLNDYKLLLCDIQQLLVAVTIHACSSLGSQYFRVIEDLIVLLGHLQNNKNEKTQNMAIALQFRVLQAAIELIKATANQDSQDLANSLRLPATHHHAMFQKRKSIADSRKFSRAQSDTLVMKMRLVASDELNMMMQRRMSQENPIRATENEFIQRLQRLTVLAVNRFIYQEPTEDCFDVLGTPETTIQCRLSTSQLEASEEEDQRELPSNTNSFLKEMFKMMIEGIKVSIATSRISTPRQQWRRILWSCKDTFRMQLGRLLVHILSPAQPLQERKQALEIVCETNHQEIIRDCLSVPLQHGPKLALYLYELMHDHKDELTREEQVSSGVFMNALKFCGYKCISPSAPPKPDLLNAMKEDRKKFEAEECVNKTVWEKTVINNQQNLSQRLDAKSKDISKIAADITQAVSLSQGMERKKVIQHIRGMYKTDLSASRHWQELVQQLTHDRTLWYDPVYYPTSWQLDPTEGPNRERRRLRRCYLTIPNKYLLRDRQKQEEVIKPPLSYLFEDKTHSSHSSTVKDKAASENIRVNRRCINVAPSRETAGELLLGKYGMYFVEDNASDTIESSSFHGETEPASFSWTYEEIKEVHKRWWQLRDNAVEIFLTNGRTLLLAFDNTKVRDDVYHNILMNNLPNLLEYGNITALTHLWYTGQITNFEYLTHLNKHAGRSFNDLMQYPVFPFILSDYTSETLDLNDPSVYRNLAKPIAVQSKEKEDRYVDTYKYLEEEYRKGAREDDPMPPVQPYHYGSHYSNSGTVLHFLVRMPPFTKMFLAYQDQSFDIPDRTFHSTHTTWRLSSYESMTDVKELIPEFFYLPEFLVNREGFDFGLRQNGERVNHVNLPPWARNDPRLFILIHRQALESDHVSQTICNWIDLVFGYKQKGKASVQAINVFHPATYFGMDVSAVEDPVQRRALETMIKTYGQTPRQLFHTAHVSRPGSRLTIEGELPAAMGLLVQFAFRETREHAKEIVYPSPLPWIKGLKWGEYVGSPSAPDPVVCFSQPHGERFGSLQALPTRAICGLSQKFCLLMVYSKEQGVRSMHSTDIQWSAILSWGYADNILRLKSKQSEPPINFIQSSQLHQVTSCAWVPDSCQLFTGSKSGVITAYMNRFSSNMHSEIEMESQVHLYGHTAEITSLFVCKPYSIMISVSKDGTCIIWDLNRLCYVQSLAGHKSPVTAVSASEMTGDIATVCDSVGGGSDLRLWTVNGDLVGHVHCREIICSVAFSNQPEGVSINVIAGGLENGVVRLWSTWDLKPVREITFPKSNKPIVSLTFSCDGHHLYTANSEGTVIAWCRKDQQRLKLPMFYSFLSSYAAG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZA1</accession>
    <name>A0A452GZA1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Oligomycin sensitivity conferral protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the ATPase delta chain family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZA1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008609.1">
        <property type="protein sequence ID" value="ENSGAGP00000007454.1"/>
        <property type="gene ID" value="ENSGAGG00000005893.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005743">
        <property type="term" value="C:mitochondrial inner membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045259">
        <property type="term" value="C:proton-transporting ATP synthase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046933">
        <property type="term" value="F:proton-transporting ATP synthase activity, rotational mechanism"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.520.20">
        <property type="entry name" value="N-terminal domain of the delta subunit of the F1F0-ATP synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026015">
        <property type="entry name" value="ATP_synth_OSCP/delta_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000711">
        <property type="entry name" value="ATPase_OSCP/dsu"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01145">
        <property type="entry name" value="ATP_synt_delta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11910">
        <property type="entry name" value="ATP SYNTHASE DELTA CHAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00213">
        <property type="entry name" value="OSCP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00125">
        <property type="entry name" value="ATPASEDELTA"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47928">
        <property type="entry name" value="N-terminal domain of the delta subunit of the F1F0-ATP synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0066">ATP synthesis</keyword>
    <keyword evidence="4" id="KW-0375">Hydrogen ion transport</keyword>
    <keyword evidence="5" id="KW-0406">Ion transport</keyword>
    <keyword evidence="6" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004370"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007046"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022781"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023310"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00033369"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007454.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="137" mass="15067" checksum="C1994164B8CC08A6" modified="2025-10-08" version="2">MRFFSTSLARPASQLVKPPIQVYGLEGRYATALYSAASKQKKLDQVEKELSRVSTLLKDPKLSSVVVNPHVKSVIKQKTINDALVKEKLSPITINFMKLLAENGRLQQTPDVISAFGKIMSAYRGEVLCSVTTANVS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZA4</accession>
    <name>A0A452GZA4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Zinc finger protein RFP-like</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GZA4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008615.1">
        <property type="protein sequence ID" value="ENSGAGP00000007459.1"/>
        <property type="gene ID" value="ENSGAGG00000005976.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd19762">
        <property type="entry name" value="Bbox2_TRIM7-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd16594">
        <property type="entry name" value="RING-HC_TRIM7-like_C-IV"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12888">
        <property type="entry name" value="SPRY_PRY_TRIM7_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.920">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001870">
        <property type="entry name" value="B30.2/SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043136">
        <property type="entry name" value="B30.2/SPRY_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003879">
        <property type="entry name" value="Butyrophylin_SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006574">
        <property type="entry name" value="PRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003877">
        <property type="entry name" value="SPRY_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050143">
        <property type="entry name" value="TRIM/RBCC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000315">
        <property type="entry name" value="Znf_B-box"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001841">
        <property type="entry name" value="Znf_RING"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017907">
        <property type="entry name" value="Znf_RING_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24103">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE TRIM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13765">
        <property type="entry name" value="PRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00622">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00643">
        <property type="entry name" value="zf-B_box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15227">
        <property type="entry name" value="zf-C3HC4_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01407">
        <property type="entry name" value="BUTYPHLNCDUF"/>
    </dbReference>
    <dbReference type="SMART" id="SM00336">
        <property type="entry name" value="BBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00589">
        <property type="entry name" value="PRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00184">
        <property type="entry name" value="RING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00449">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57845">
        <property type="entry name" value="B-box zinc-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57850">
        <property type="entry name" value="RING/U-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50188">
        <property type="entry name" value="B302_SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50119">
        <property type="entry name" value="ZF_BBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00518">
        <property type="entry name" value="ZF_RING_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50089">
        <property type="entry name" value="ZF_RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="1" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0862">Zinc</keyword>
    <keyword evidence="2 4" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="RING-type" evidence="6">
        <location>
            <begin position="17"/>
            <end position="58"/>
        </location>
    </feature>
    <feature type="domain" description="B box-type" evidence="7">
        <location>
            <begin position="95"/>
            <end position="132"/>
        </location>
    </feature>
    <feature type="domain" description="B30.2/SPRY" evidence="8">
        <location>
            <begin position="241"/>
            <end position="429"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="5">
        <location>
            <begin position="180"/>
            <end position="229"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
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        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00024"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50089"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="PROSITE" id="PS50119"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="PROSITE" id="PS50188"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007459.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="11">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="429" mass="48216" checksum="A98C3B6C21DA01E0" modified="2025-10-08" version="2">MNSGKNPVESLQEEATCPICLEYFTEPVTLECGHNFCRACIAQCWEGSDTDISCPQCRETVQQRNLRPNRQLANVVEIVKRLSLQAAKGAGGDGVCGEHQEALKLFCEEDQSPICVICRESRAHCAHTVVPIQEAAQEYKLSMKGPQVPFISLTSPFHIFIQKQTQTERQKIVSEFQELREFLEEQERLLLAQLEKLDEEIVRIQNENVSKLSEQISRLSELISEMEGKCQKPGSEFLKVRLREKQSSSPNALILSVPANVTLDPDTAQLRLVLSEDRKSVRYGDTRQDLPDNPQRFDTDFCVLGCEGFTSGRHCWEVEVEDGQCWAVGVARESVGRKGGISYSPEGGIWAVLRWMGQFQVLTSPVTPLPLRRVPSRIQVCLDCDRGQVTFIDAGDEAPIFTFPPGSIPGERIRPWLCVGYRSRLCLCP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZB2</accession>
    <name>A0A452GZB2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">POF1B helix-loop-helix domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
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            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GZB2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008540.1">
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        <property type="gene ID" value="ENSGAGG00000005876.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="term" value="C:actin filament"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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    </dbReference>
    <dbReference type="GO" id="GO:0005923">
        <property type="term" value="C:bicellular tight junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
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        <property type="term" value="P:actin filament organization"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="P:bicellular tight junction assembly"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="P:epithelial cell morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="entry name" value="POF1B"/>
    </dbReference>
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    </dbReference>
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        <property type="entry name" value="PREMATURE OVARIAN FAILURE, 1B"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="POF1B_HlH"/>
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    </dbReference>
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    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="POF1B helix-loop-helix" evidence="2">
        <location>
            <begin position="40"/>
            <end position="110"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="131"/>
            <end position="158"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="194"/>
            <end position="302"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF24617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007392.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="347" mass="40943" checksum="DCC856452740E5FB" modified="2019-05-08" version="1">MTRIWPLQSSTVRTVIEQRLQRKYSNIQYLQETNGQLDSRYFGELLAEVNRKSNDLYSCLLQHVEKIGRRYYIEDLIPKGLSELTKQQIRYLLQMRVTSDKSLRLVLSTFGSLREELCHLQDDLGKLETDKMLLEKDLAFKESQVKEYETLLASVRENNRQQQQGLRDSASKCRSLEEQIISLRYAEGEKDCRLKELEYCKRALEQEIQNLRMQTCSNPVIQTTTDELSSRYVEMINNLREDKDREIRNLRSQINQFQQDISRRDGSNSDFQMRLHELTSMLEEKETFIKQQQEELFRLKQERLSCNQSTGVTKSIITKKYMNQYPILGLLSDDCMASSPKHVSYPC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZB3</accession>
    <name>A0A452GZB3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">RHD domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZB3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008796.1">
        <property type="protein sequence ID" value="ENSGAGP00000007620.1"/>
        <property type="gene ID" value="ENSGAGG00000006099.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007249">
        <property type="term" value="P:canonical NF-kappaB signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033554">
        <property type="term" value="P:cellular response to stress"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006954">
        <property type="term" value="P:inflammatory response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045087">
        <property type="term" value="P:innate immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030098">
        <property type="term" value="P:lymphocyte differentiation"/>
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        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0038061">
        <property type="term" value="P:non-canonical NF-kappaB signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="P:positive regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034097">
        <property type="term" value="P:response to cytokine"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048511">
        <property type="term" value="P:rhythmic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd01177">
        <property type="entry name" value="IPT_NFkappaB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd07886">
        <property type="entry name" value="RHD-n_RelB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000046">
        <property type="entry name" value="Nuclear factor NF-kappa-B p105 subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.340:FF:000005">
        <property type="entry name" value="RELB proto-oncogene, NF-kB subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.340">
        <property type="entry name" value="Rel homology domain (RHD), DNA-binding domain"/>
        <property type="match status" value="1"/>
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    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014756">
        <property type="entry name" value="Ig_E-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002909">
        <property type="entry name" value="IPT_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033926">
        <property type="entry name" value="IPT_NFkappaB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000451">
        <property type="entry name" value="NFkB/Dor"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008967">
        <property type="entry name" value="p53-like_TF_DNA-bd_sf"/>
    </dbReference>
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        <property type="entry name" value="RelB_RHD_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032400">
        <property type="entry name" value="RelB_transact"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR030492">
        <property type="entry name" value="RHD_CS"/>
    </dbReference>
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        <property type="entry name" value="RHD_dimer"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011539">
        <property type="entry name" value="RHD_DNA_bind_dom"/>
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        <property type="entry name" value="RHD_DNA_bind_dom_sf"/>
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        <property type="entry name" value="NUCLEAR FACTOR NF-KAPPA-B PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="TRANSCRIPTION FACTOR RELB"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00554">
        <property type="entry name" value="RHD_DNA_bind"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00057">
        <property type="entry name" value="NFKBTNSCPFCT"/>
    </dbReference>
    <dbReference type="SMART" id="SM00429">
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="E set domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49417">
        <property type="entry name" value="p53-like transcription factors"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
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    <keyword evidence="4" id="KW-0010">Activator</keyword>
    <keyword evidence="3" id="KW-0090">Biological rhythms</keyword>
    <keyword evidence="6" id="KW-0539">Nucleus</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0804">Transcription</keyword>
    <keyword evidence="2" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="RHD" evidence="8">
        <location>
            <begin position="159"/>
            <end position="336"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1"/>
            <end position="28"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="46"/>
            <end position="69"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="128"/>
            <end position="147"/>
        </location>
    </feature>
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        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50254"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007620.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="593" mass="65181" checksum="623202E6A1D2B2AB" modified="2019-05-08" version="1">MAHTAQASGKGAAISDEDELGEWTLGNGGTATSEIIEELIKEDGFQPQAPPFLSKSSPKGTAKLVPRGSNPAVPTGMLRLMEPTGPSPLQGGLASFGSSRPTAACVLLPDSQRPLLSQSLSSLQLTPRLVPRGTTPPPASRPHWASPIRSPKEQLEDMLEPPTLVITEQPKQRGMRFRYECEGRSAGSILGESSTEASKTLPAIELLNCQKIPEVKVIACLVWKDWPYRIHPHSLVGKDCNNGLCEVTLRPQANPKHSFNNLGIQCVKKKEIEAAIEKKLQLGIDPFKAGSLKNHQEVDMNVVRICFQASYQEPSGQIRHFSPVLSEPIFDKKSTNTSELRICRMNKECGPCTGGEELYLLCDKVQKEDIAVVFRRDAWESKADFSQADVHRQIAIVFKTPPYQHLDILEPVEVEVYLQRLTDSVCSEPFHFTYLPKDHDAYGVHVKRKRGMPDVLEELSASDPHGIEAKKRKKKPGYMDHFLAPQAAEEGFALHPSLEHITVPEPFDLCLGPDYASYSCLNQRDMLPTACSPMELPGCLGAEFLLDTYSSAQSNPAPFLAPGDSSTGDIEGTASLVGSNMFPSQYKEAYERA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZB5</accession>
    <name>A0A452GZB5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Protein NPAT C-terminal domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000008625.1">
        <property type="protein sequence ID" value="ENSGAGP00000007469.1"/>
        <property type="gene ID" value="ENSGAGG00000005980.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003712">
        <property type="term" value="F:transcription coregulator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006594">
        <property type="entry name" value="LisH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031442">
        <property type="entry name" value="NPAT_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052850">
        <property type="entry name" value="NPAT_LisH"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15087">
        <property type="entry name" value="PROTEIN NPAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15087:SF14">
        <property type="entry name" value="PROTEIN NPAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15712">
        <property type="entry name" value="NPAT_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50896">
        <property type="entry name" value="LISH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Protein NPAT C-terminal" evidence="2">
        <location>
            <begin position="732"/>
            <end position="1402"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="179"/>
            <end position="241"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="354"/>
            <end position="387"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="403"/>
            <end position="443"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="489"/>
            <end position="542"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="595"/>
            <end position="641"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="946"/>
            <end position="1086"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1127"/>
            <end position="1212"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1236"/>
            <end position="1256"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1339"/>
            <end position="1378"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="179"/>
            <end position="189"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="228"/>
            <end position="237"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="366"/>
            <end position="377"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="409"/>
            <end position="420"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="499"/>
            <end position="513"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="525"/>
            <end position="536"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="619"/>
            <end position="640"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="957"/>
            <end position="996"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="1008"/>
            <end position="1023"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="1036"/>
            <end position="1052"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="1069"/>
            <end position="1078"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="1127"/>
            <end position="1147"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="1165"/>
            <end position="1183"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="1">
        <location>
            <begin position="1184"/>
            <end position="1194"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="1203"/>
            <end position="1212"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="1345"/>
            <end position="1364"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF15712"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007469.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1402" mass="151693" checksum="6B80254DAC71B165" modified="2019-05-08" version="1">MLLPSDVARLVLGYLQQEKLLATCREFILESSDLKEYAEHYTEEGFVPACLLSLFGKNLTTILNEYVAMKTKETTNEVPAIMSSLWKKLDYTLSQIRSMQNSAGFSANQRTRTRNGIAEMKRQRILQSAPPTSGLLSIAHQSGQLNSSCMAPTQIIHRPTVNQNVSQTRLNMLFAHQSQTQENNVSTGDSIHILVPGPQERKFNSSLLSPGRRKSESQKKKSIASPGPHSTSRSFQDSSVVVVEEESEPLQELIDGNFPNEIHMSEEAIQDILEQTESDPAFQALFDLFEPGKNKTNKNAPHAISNQNGVEGTMLVDDHNLETIASSLETQEASDHSREAQSCTDNHSEEVSCALKNSNDGDDTTNKSVTCEQSHGSSKQKKQTDNLKTVNDEQIGELEIAFESVPDLTDSNQRENSGTEQSKHFVGSYSNKESAELVSKSQTAMETEKDALNHNGQNTAGLEYDHSDIPQSSLASLDEDTITIENRTYPGTSECHLSPDTSLSGKTNISPSHRSPDHSVMLKKSNASKCSQSGDSGENEKDITEETPMLPVTLQQNSNHHSNCQDLDKMVLDCVERSVNISDLTRTELQLEVVDTSNKPHSGDQFTLDKPSTKDLSFHSGSSDSGRTQVEMQEPSSSTETDAETIFFSSGDDGCTEISVVSTESNPTVSEACHIPPLETACLTEESSTEFKSIGGGSSSSQPVDVDPSSIVSLKIIISDDPFISSDTELNNAVSSITGENLPTIILSSPAKTTGLSKCVTSEESDRAGDSASAEQNLLVLRPKDPVVSTANIQNEDCTVFSVAGATHLSKDGGFIQLMPATSTAFGNSNNLYIATCVTDPTTLGTTVTPSNVVVLPGNSVPLAAQAPATQQLQTPPRSNMFTMNQTVSPNFSQGSAIIIASPVQPVLPGMVGMIPVSVVGQSGNTFSAPPRQVIHMPIANPMCNRSIPKIPIPPKSQKTAGARNKSNTGKLVTSTAKPVSHPSSRVQRTGNSDRNASADLGRKLEESSLTVPAEPTSSNSKPNESHRRVLCFDSALSTPGGNTQVKNTKSSLQKERNENPLYAVDSPNAASASVKIQSNKREKDKTLPRILCKAEIGGSRNAAVKDAQPERRAVAAGLALDSFHKTTANKENELRRDVDEKQKNQDAAKLSNGQKNVGLWNEKTVASVQEQNKKQGSLSNVNSKTSGSVSLSSKEPKRDQTKPSNHSLSLASPLTKQAVEMLHGIQWHSPASKVVENGDLPVPRTPSGVGDTHTEDIADSVRTPTCRRYTEESTTPRIMVPPATPDLPACSPASETGSENSVSMAAHTLMILSRAAIARTNTMTPLKDNIQQFRSVRSTAKKRKLEDLNECERNSRSASKKELQSSTTPVKKKKIKKKKLPHCFPAGMDVDKFLLSLHYDE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZC0</accession>
    <name>A0A452GZC0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Zinc finger protein RFP-like</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GZC0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008632.1">
        <property type="protein sequence ID" value="ENSGAGP00000007474.1"/>
        <property type="gene ID" value="ENSGAGG00000005976.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd19762">
        <property type="entry name" value="Bbox2_TRIM7-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd16594">
        <property type="entry name" value="RING-HC_TRIM7-like_C-IV"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12888">
        <property type="entry name" value="SPRY_PRY_TRIM7_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.920:FF:000004">
        <property type="entry name" value="Butyrophilin subfamily 1 member A1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.920">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001870">
        <property type="entry name" value="B30.2/SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043136">
        <property type="entry name" value="B30.2/SPRY_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003879">
        <property type="entry name" value="Butyrophylin_SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006574">
        <property type="entry name" value="PRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003877">
        <property type="entry name" value="SPRY_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050143">
        <property type="entry name" value="TRIM/RBCC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000315">
        <property type="entry name" value="Znf_B-box"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001841">
        <property type="entry name" value="Znf_RING"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017907">
        <property type="entry name" value="Znf_RING_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24103">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE TRIM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13765">
        <property type="entry name" value="PRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00622">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00643">
        <property type="entry name" value="zf-B_box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15227">
        <property type="entry name" value="zf-C3HC4_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01407">
        <property type="entry name" value="BUTYPHLNCDUF"/>
    </dbReference>
    <dbReference type="SMART" id="SM00336">
        <property type="entry name" value="BBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00589">
        <property type="entry name" value="PRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00184">
        <property type="entry name" value="RING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00449">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57845">
        <property type="entry name" value="B-box zinc-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57850">
        <property type="entry name" value="RING/U-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50188">
        <property type="entry name" value="B302_SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50119">
        <property type="entry name" value="ZF_BBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00518">
        <property type="entry name" value="ZF_RING_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50089">
        <property type="entry name" value="ZF_RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="1" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0862">Zinc</keyword>
    <keyword evidence="2 4" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="RING-type" evidence="6">
        <location>
            <begin position="16"/>
            <end position="57"/>
        </location>
    </feature>
    <feature type="domain" description="B box-type" evidence="7">
        <location>
            <begin position="94"/>
            <end position="131"/>
        </location>
    </feature>
    <feature type="domain" description="B30.2/SPRY" evidence="8">
        <location>
            <begin position="243"/>
            <end position="431"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="5">
        <location>
            <begin position="131"/>
            <end position="227"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00024"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50119"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50188"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007474.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="11">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="431" mass="48523" checksum="2B3395A136E3565C" modified="2019-05-08" version="1">MAAENPVESLQEEATCPICLEYFTEPVTLECGHNFCRACIAQCWEGSDTDISCPQCRETVQQRNLRPNRQLANVVEIVKRLSLQAAKGAGGDGVCGEHQEALKLFCEEDQSPICVICRESRAHCAHTVVPIQEAAQEYKEKIQAYLKALREEREKLLGFKKQTQTERQKIVSEFQELREFLEEQERLLLAQLEKLDEEIVRIQNENVSKLSEQISRLSELISEMEGKCQKPGSEFLKFEKWSQPSLGPSSSPNALILSVPANVTLDPDTAQLRLVLSEDRKSVRYGDTRQDLPDNPQRFDTDFCVLGCEGFTSGRHCWEVEVEDGQCWAVGVARESVGRKGGISYSPEGGIWAVLRWMGQFQVLTSPVTPLPLRRVPSRIQVCLDCDRGQVTFIDAGDEAPIFTFPPGSIPGERIRPWLCVGYRSRLCLCP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZE5</accession>
    <name>A0A452GZE5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Mothers against decapentaplegic homolog</fullName>
            <shortName evidence="9">MAD homolog</shortName>
            <shortName evidence="9">Mothers against DPP homolog</shortName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="9">SMAD family member</fullName>
        </alternativeName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="9">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="9">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="1 9">Belongs to the dwarfin/SMAD family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZE5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008666.1">
        <property type="protein sequence ID" value="ENSGAGP00000007503.1"/>
        <property type="gene ID" value="ENSGAGG00000006006.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071144">
        <property type="term" value="C:heteromeric SMAD protein complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070411">
        <property type="term" value="F:I-SMAD binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032924">
        <property type="term" value="P:activin receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009653">
        <property type="term" value="P:anatomical structure morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045944">
        <property type="term" value="P:positive regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060395">
        <property type="term" value="P:SMAD protein signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007179">
        <property type="term" value="P:transforming growth factor beta receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd10491">
        <property type="entry name" value="MH1_SMAD_2_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd10985">
        <property type="entry name" value="MH2_SMAD_2_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.200.10:FF:000001">
        <property type="entry name" value="Mothers against decapentaplegic homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.520.10:FF:000001">
        <property type="entry name" value="Mothers against decapentaplegic homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.200.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.520.10">
        <property type="entry name" value="SMAD MH1 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013790">
        <property type="entry name" value="Dwarfin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003619">
        <property type="entry name" value="MAD_homology1_Dwarfin-type"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013019">
        <property type="entry name" value="MAD_homology_MH1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017855">
        <property type="entry name" value="SMAD-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001132">
        <property type="entry name" value="SMAD_dom_Dwarfin-type"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008984">
        <property type="entry name" value="SMAD_FHA_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036578">
        <property type="entry name" value="SMAD_MH1_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13703:SF42">
        <property type="entry name" value="MOTHERS AGAINST DECAPENTAPLEGIC HOMOLOG 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13703">
        <property type="entry name" value="SMAD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03165">
        <property type="entry name" value="MH1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03166">
        <property type="entry name" value="MH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00523">
        <property type="entry name" value="DWA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00524">
        <property type="entry name" value="DWB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56366">
        <property type="entry name" value="SMAD MH1 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49879">
        <property type="entry name" value="SMAD/FHA domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51075">
        <property type="entry name" value="MH1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51076">
        <property type="entry name" value="MH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 9" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8 9" id="KW-0539">Nucleus</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7 9" id="KW-0804">Transcription</keyword>
    <keyword evidence="5 9" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="4" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="MH1" evidence="11">
        <location>
            <begin position="10"/>
            <end position="146"/>
        </location>
    </feature>
    <feature type="domain" description="MH2" evidence="12">
        <location>
            <begin position="244"/>
            <end position="437"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="177"/>
            <end position="222"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="177"/>
            <end position="187"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="203"/>
            <end position="213"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00005545"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="RuleBase" id="RU361195"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS51075"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS51076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007503.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="437" mass="48913" checksum="0DC32049E3BDDE3C" modified="2019-05-08" version="1">MSSILPFTPPVVKRLLGWKKSAGGSGGAGGGEQNGQEEKWCEKAVKSLVKKLKKTGQLDELEKAITTQNCNTKCVTIPRSLDGRLQVSHRKGLPHVIYCRLWRWPDLHSHHELKAIENCEYAFNLKKDEVCVNPYHYQRVETPVLPPVLVPRHTEILTELPPLDDYTHSIPENTNFPAGIEPQSNYIPETPPPGYISEDGETSDQQLNQSMDTGSPAELSPSTLSPVNHSLDLQPVTYSEPAFWCSIAYYELNQRVGETFHASQPSLTVDGFTDPSNSERFCLGLLSNVNRNATVEMTRRHIGRGVRLYYIGGEVFAECLSDSAIFVQSPNCNQRYGWHPATVCKIPPGCNLKIFNNQEFAALLAQSVNQGFEAVYQLTRMCTIRMSFVKGWGAEYRRQTVTSTPCWIELHLNGPLQWLDKVLTQMGSPSVRCSSMS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZE9</accession>
    <name>A0A452GZE9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="19">Phospholipid-transporting ATPase</fullName>
            <ecNumber evidence="19">7.6.2.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="15 19">
            <text>ATP + H2O + phospholipidSide 1 = ADP + phosphate + phospholipidSide 2.</text>
            <dbReference type="EC" id="7.6.2.1"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 18">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Golgi apparatus</location>
            <location evidence="2">trans-Golgi network membrane</location>
            <topology evidence="2">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="19">Membrane</location>
            <topology evidence="19">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 19">Belongs to the cation transport ATPase (P-type) (TC 3.A.3) family. Type IV subfamily.</text>
    </comment>
    <comment type="caution">
        <text evidence="19">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="EC" id="7.6.2.1" evidence="19"/>
    <dbReference type="AlphaFoldDB" id="A0A452GZE9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008595.1">
        <property type="protein sequence ID" value="ENSGAGP00000007440.1"/>
        <property type="gene ID" value="ENSGAGG00000000721.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005768">
        <property type="term" value="C:endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005802">
        <property type="term" value="C:trans-Golgi network"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140326">
        <property type="term" value="F:ATPase-coupled intramembrane lipid transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000287">
        <property type="term" value="F:magnesium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006897">
        <property type="term" value="P:endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045332">
        <property type="term" value="P:phospholipid translocation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006890">
        <property type="term" value="P:retrograde vesicle-mediated transport, Golgi to endoplasmic reticulum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.1110.10:FF:000008">
        <property type="entry name" value="Phospholipid-transporting ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1000:FF:000009">
        <property type="entry name" value="Phospholipid-transporting ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.1110.10">
        <property type="entry name" value="Calcium-transporting ATPase, cytoplasmic domain N"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.70.150.10">
        <property type="entry name" value="Calcium-transporting ATPase, cytoplasmic transduction domain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1110.10">
        <property type="entry name" value="Calcium-transporting ATPase, transmembrane domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1000">
        <property type="entry name" value="HAD superfamily/HAD-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023299">
        <property type="entry name" value="ATPase_P-typ_cyto_dom_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018303">
        <property type="entry name" value="ATPase_P-typ_P_site"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023298">
        <property type="entry name" value="ATPase_P-typ_TM_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036412">
        <property type="entry name" value="HAD-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023214">
        <property type="entry name" value="HAD_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006539">
        <property type="entry name" value="P-type_ATPase_IV"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001757">
        <property type="entry name" value="P_typ_ATPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032630">
        <property type="entry name" value="P_typ_ATPase_c"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044492">
        <property type="entry name" value="P_typ_ATPase_HD_dom"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01652">
        <property type="entry name" value="ATPase-Plipid"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01494">
        <property type="entry name" value="ATPase_P-type"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24092:SF50">
        <property type="entry name" value="PHOSPHOLIPID-TRANSPORTING ATPASE IIB-RELATED"/>
        <property type="match status" value="1"/>
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    <dbReference type="PANTHER" id="PTHR24092">
        <property type="entry name" value="PROBABLE PHOSPHOLIPID-TRANSPORTING ATPASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00702">
        <property type="entry name" value="Hydrolase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16212">
        <property type="entry name" value="PhoLip_ATPase_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00119">
        <property type="entry name" value="CATATPASE"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDG00002">
        <property type="entry name" value="C1.7:_P-type_atpase_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDS00003">
        <property type="entry name" value="Haloacid_Dehalogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDF00027">
        <property type="entry name" value="p-type_atpase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81665">
        <property type="entry name" value="Calcium ATPase, transmembrane domain M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56784">
        <property type="entry name" value="HAD-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81660">
        <property type="entry name" value="Metal cation-transporting ATPase, ATP-binding domain N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00154">
        <property type="entry name" value="ATPASE_E1_E2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 17 19" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="12" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="13" id="KW-0445">Lipid transport</keyword>
    <keyword evidence="9 18 19" id="KW-0460">Magnesium</keyword>
    <keyword evidence="14 19" id="KW-0472">Membrane</keyword>
    <keyword evidence="6 18" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7 17 19" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10 19" id="KW-1278">Translocase</keyword>
    <keyword evidence="5 19" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 19" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="4" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="107"/>
            <end position="127"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="666"/>
            <end position="686"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="692"/>
            <end position="713"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="720"/>
            <end position="740"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="746"/>
            <end position="772"/>
        </location>
    </feature>
    <feature type="domain" description="P-type ATPase C-terminal" evidence="20">
        <location>
            <begin position="601"/>
            <end position="781"/>
        </location>
    </feature>
    <feature type="active site" description="4-aspartylphosphate intermediate" evidence="16">
        <location>
            <position position="192"/>
        </location>
    </feature>
    <feature type="binding site" evidence="17">
        <location>
            <position position="192"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="192"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
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        <location>
            <position position="193"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
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        <location>
            <position position="194"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
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        <location>
            <position position="194"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
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        </ligand>
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        <location>
            <position position="296"/>
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        <ligand>
            <name>ATP</name>
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        <location>
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        <ligand>
            <name>ATP</name>
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        <location>
            <position position="362"/>
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        <location>
            <position position="391"/>
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            <name>ATP</name>
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        </ligand>
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        <location>
            <position position="471"/>
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            <name>ATP</name>
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        <location>
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        <location>
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        <location>
            <position position="583"/>
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    <sequence length="789" mass="88366" checksum="64AA7405F0405C8E" modified="2019-05-08" version="1">LKVAVSCTQRLPALGDLFSISAYVYAQKPQLDIHSFEGTFTREDSDPTVHESLSIENTLWASTVVASGTVIGVVIYTGKETRSVMNTSNPKNKVGLLDLELNQLTKALFLALVALSIVMVTLQGFVGPWYRNLFRFLLLFSYIIPISLRVNLDMGKAAYGWMIMKDDNIPGTVVRTSTIPEELGRLVYLLTDKTGTLTQNEMIFKRLHLGTVSYGTDTMDEIQNHIISSYSQSSAPKVRKTVSSRIHEAVKAIALCHNVTPVYESHAGVNGETEYAEVDQNFSDENRTYQASSPDEVALVQWTESVGLTLVNRDLTSMQLKSPGGQILTYYILQVFPFTSESKRMGIIVRDESTGEITFYMKGADVAMSTIVQYNDWLEEECGNMAREGLRTLVVAKKSLTEEQHQDFENRYNQAKLSIHDRALKVAAVVESLEREMELLCLTGVEDQLQADVRPTLEMLRNAGIKIWMLTGDKLETATCIAKSSHLVSRTQDIHIFRPVISRGEAHLELNAFRRKHDCALVISGDSLEVCLKYYEHEFVELACQCPAVVCCRCSPTQKAHIVKLLQHHTGKRTCAIGDGGNDVSMIQAADCGIGIEGKAVFSSVFYFASIPLYQGFLMVGYATIYTMFPVFSLVLDQDVKPEMALLYPELYKDLTKGRSLSFKTFLIWVLISIYQGGILMYGALLLFESEFVHVVAISFTALILTELLMVALTIRTWHWLMVVAEFFSLGCYVASLAFLNEYFDVAFITTVTFLWKVSAITVVSCLPLYILKYLKRKFSPPSYSKLTS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZF3</accession>
    <name>A0A452GZF3_9SAUR</name>
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        <recommendedName>
            <fullName evidence="14">Contactin 6</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZF3"/>
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        <property type="protein sequence ID" value="ENSGAGP00000007562.1"/>
        <property type="gene ID" value="ENSGAGG00000005911.1"/>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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        <property type="project" value="UniProtKB-SubCell"/>
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        <property type="term" value="F:cell-cell adhesion mediator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007411">
        <property type="term" value="P:axon guidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="P:brain development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="match status" value="4"/>
    </dbReference>
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        <property type="entry name" value="Contactin 1"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Contactin 1"/>
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        <property type="match status" value="1"/>
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        <property type="entry name" value="DCC isoform 1"/>
        <property type="match status" value="2"/>
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        <property type="entry name" value="Neuronal cell adhesion molecule"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="8"/>
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        <property type="entry name" value="FN3_dom"/>
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        <property type="entry name" value="FN3_sf"/>
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        <property type="entry name" value="Ig-like_dom"/>
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        <property type="entry name" value="Ig-like_dom_sf"/>
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        <property type="entry name" value="Ig-like_fold"/>
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        <property type="entry name" value="Ig_I-set"/>
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        <property type="entry name" value="Ig_sub"/>
    </dbReference>
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        <property type="entry name" value="Ig_sub2"/>
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        <property type="entry name" value="CONTACTIN 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR44170">
        <property type="entry name" value="PROTEIN SIDEKICK"/>
        <property type="match status" value="1"/>
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    <dbReference type="Pfam" id="PF00041">
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        <property type="match status" value="1"/>
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        <property type="match status" value="4"/>
    </dbReference>
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        <property type="entry name" value="IG"/>
        <property type="match status" value="4"/>
    </dbReference>
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        <property type="entry name" value="IGc2"/>
        <property type="match status" value="4"/>
    </dbReference>
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        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="2"/>
    </dbReference>
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        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="7" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="8" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="6" id="KW-0472">Membrane</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0732">Signal</keyword>
    <feature type="domain" description="Ig-like" evidence="10">
        <location>
            <begin position="64"/>
            <end position="132"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="10">
        <location>
            <begin position="142"/>
            <end position="226"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="10">
        <location>
            <begin position="232"/>
            <end position="326"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="10">
        <location>
            <begin position="338"/>
            <end position="411"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="11">
        <location>
            <begin position="424"/>
            <end position="522"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="11">
        <location>
            <begin position="527"/>
            <end position="624"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="11">
        <location>
            <begin position="629"/>
            <end position="725"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="11">
        <location>
            <begin position="726"/>
            <end position="823"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007562.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="14">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="855" mass="94436" checksum="88E9276305884E86" modified="2019-05-08" version="1">MKAKVHDLIVNRGTNIARFLVDGLFAAELVQLYVSEIVGRWVKSRGTGLWGANIRDAYLVNPLCTRKTSSIEAPCTPLPVPSISWKRSDGHPLARKIKHYKSKGVLEIPDFQQEDEGFYECIAGNIRGRNIARGQLIIYALPEWDKKIQNAYLSLYDSLFWECKASGKPSPSYSWLKNGQPLMPEERIQIENGTLTITMLNVSDSGLYQCVAENKYNTIYTNAELRVIASAPDFSKNPVKQLSVAQVGGEVTIECKPNASPRAVINWKKGTNLLQQNKRIFMLDDGSLKIYNITKTDAGVYTCIAMNQFGVARNSGNLIVKERTVITVPPSKMDVTVGESIVLPCQVSHDPSAEVVFTWSFNGDVIEFKRGVNHFERIGGESVGDLMIRNIQLNHSGKYVCMVQTTIDSLSATADVIVRGPPGPPQDVRVEHISSTTALLCWRPGTDNNSPVHIYSVQTRTPFSVGWQAVSTVPEVLNGRTYSATVTGLNPWVEYEFRVVAGNNIGIGEPSRPSELLRTKASIPLVAPTNVSGGGGSRSELVITWESVPEELQNGEGFGYIIMFRPLGSTSWTKAVVASIEASKYVYRNESITPLSPFEVKVGVYNNEGEGTLSSISIVYSGEDEPQIAPKGVSVQSFSASEVEVSWNAIAWNRHTGRVLGYEVLYWTDDSKETAAGKVRVSGNVTARNITGLKANTIYFATIRAYNTAGTGPSSTPVNITTKKSPPSQPPVNIAWKLTNSKICLNWEHVKTMENESEVLGYKISYRQNRQSKTHILETNNTSAELLVPFEEDYLIEIRTVSDGGDGSSSEEIRIPKISTTANIRDDYTIEGWECTLYFIKNVNINIALLKYIKI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZF4</accession>
    <name>A0A452GZF4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Protein kinase N1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GZF4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008683.1">
        <property type="protein sequence ID" value="ENSGAGP00000007519.1"/>
        <property type="gene ID" value="ENSGAGG00000006005.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd05589">
        <property type="entry name" value="STKc_PKN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000058">
        <property type="entry name" value="Putative serine/threonine-protein kinase N2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000038">
        <property type="entry name" value="serine/threonine-protein kinase N2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000961">
        <property type="entry name" value="AGC-kinase_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017892">
        <property type="entry name" value="Pkinase_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24351">
        <property type="entry name" value="RIBOSOMAL PROTEIN S6 KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00433">
        <property type="entry name" value="Pkinase_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00133">
        <property type="entry name" value="S_TK_X"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51285">
        <property type="entry name" value="AGC_KINASE_CTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6 7" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="5" id="KW-0418">Kinase</keyword>
    <keyword evidence="4 7" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="3" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Protein kinase" evidence="9">
        <location>
            <begin position="178"/>
            <end position="435"/>
        </location>
    </feature>
    <feature type="domain" description="AGC-kinase C-terminal" evidence="10">
        <location>
            <begin position="436"/>
            <end position="503"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="116"/>
            <end position="154"/>
        </location>
    </feature>
    <feature type="binding site" evidence="7">
        <location>
            <position position="207"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS51285"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007519.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="13">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="503" mass="56119" checksum="DEACFCE53BB5D3C3" modified="2019-05-08" version="1">MGSLECGLWWPHCTGGWELPHSPWEHSSVAAGVGGCSDLTLSLPSPSCPGPASHPGLLLPQVTFFDPVIERIPKLQRQKKIFSKQQGKAFLRARQMNIDIATWVRLLRRLLPTASSTGAYSPSAHVPTGPDSEGHHSLRPGQLADGGERPERGALPQRCPVLTLTLHPRTTPLTLDEFRFLAVLGRGHFGKVLLSEYSHTGELYAIKALKKGDIVARDEVESLMCEKRIFETVNSSRHPFLVNLFACFQTPEHVCFVMEYTAGGDLMMHIHTDVFTEPRATFYAACVVLGLQFLHERKIVPHPVKGGLLNAWGGPQGLTLSPPRSGMGYGDRTSTFCGTPEFLAPEVLTDTSYTRTVDWWGLGVLIYEMLVGESPFPGDDEEEVFDSIVNDEVRYPRFLSTEAIGVMRRLLRRNPERRLGSSERDAEDVKKQPFFRVIDWEALLARQLPPPFVPIIKGREDISNFDEEFTAEAPALTPPREPRPLTAAEQEAFRHFDYLADTC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZF5</accession>
    <name>A0A452GZF5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Cytochrome P450 family 2 subfamily W member 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="1 13">
            <name>heme</name>
            <dbReference type="ChEBI" id="CHEBI:30413"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Endoplasmic reticulum membrane</location>
            <topology evidence="3">Peripheral membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Microsome membrane</location>
            <topology evidence="2">Peripheral membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4 14">Belongs to the cytochrome P450 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZF5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008600.1">
        <property type="protein sequence ID" value="ENSGAGP00000007445.1"/>
        <property type="gene ID" value="ENSGAGG00000005769.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0020037">
        <property type="term" value="F:heme binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005506">
        <property type="term" value="F:iron ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016712">
        <property type="term" value="F:oxidoreductase activity, acting on paired donors, with incorporation or reduction of molecular oxygen, reduced flavin or flavoprotein as one donor, and incorporation of one atom of oxygen"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006082">
        <property type="term" value="P:organic acid metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006805">
        <property type="term" value="P:xenobiotic metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.630.10:FF:000238">
        <property type="entry name" value="Cytochrome P450 2A6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.630.10">
        <property type="entry name" value="Cytochrome P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001128">
        <property type="entry name" value="Cyt_P450"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017972">
        <property type="entry name" value="Cyt_P450_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002401">
        <property type="entry name" value="Cyt_P450_E_grp-I"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036396">
        <property type="entry name" value="Cyt_P450_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050182">
        <property type="entry name" value="Cytochrome_P450_fam2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24300:SF302">
        <property type="entry name" value="CYTOCHROME P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24300">
        <property type="entry name" value="CYTOCHROME P450 508A4-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00067">
        <property type="entry name" value="p450"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00463">
        <property type="entry name" value="EP450I"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00385">
        <property type="entry name" value="P450"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48264">
        <property type="entry name" value="Cytochrome P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00086">
        <property type="entry name" value="CYTOCHROME_P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="5 13 14" id="KW-0349">Heme</keyword>
    <keyword evidence="10 13 14" id="KW-0408">Iron</keyword>
    <keyword evidence="12 15" id="KW-0472">Membrane</keyword>
    <keyword evidence="6 13 14" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8" id="KW-0492">Microsome</keyword>
    <keyword evidence="11 14" id="KW-0503">Monooxygenase</keyword>
    <keyword evidence="9 14" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="15" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="15" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="6"/>
            <end position="26"/>
        </location>
    </feature>
    <feature type="binding site" description="axial binding residue" evidence="13">
        <location>
            <position position="368"/>
        </location>
        <ligand>
            <name>heme</name>
            <dbReference type="ChEBI" id="CHEBI:30413"/>
        </ligand>
        <ligandPart>
            <name>Fe</name>
            <dbReference type="ChEBI" id="CHEBI:18248"/>
        </ligandPart>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001971"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004174"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004406"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00010617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022848"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023033"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PIRSR" id="PIRSR602401-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="RuleBase" id="RU000461"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007445.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="18">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="425" mass="48303" checksum="D61BA6749665DDCD" modified="2019-05-08" version="1">SSKMALADPTVILLLFVVTSVSILFVKKSWNKRTCLNFPPGPRPLPILGNLHQLNLKRPYRTLLKKYGPVFSIHMGPKNMVVLSGYKMVKEALVNQADLPSSLATILGVANRMQQRSSFDTMKMMNAAVSNVIVSIMLGHRFDYEDPKLLRLIHLINESVRLSGSPMVTNLEELYSFTQQTFIEHLKQLDPNDQRSFIDAFLIRQQEEKSNPNTHFKNENLQSLVSNLFTAGMETTSTTLRWGLLLMMKYPAIQKKVHEEIERVIGSAQLRYEHRTQMPYTDAVIHEIQRFANILPMSLPHETTEDVRLNGYVIPKGTYIIPLLASVLYDKTQWEKPGEFCPQHFLSTDGKFVTNEAFMPFSAGRRICAGETLARMELFLFFTSLLQRFTFHPPPGVTSSDLDLTPAVGFTTPPMPHKICAMPCR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZF7</accession>
    <name>A0A452GZF7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">RBR-type E3 ubiquitin transferase</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000008736.1">
        <property type="protein sequence ID" value="ENSGAGP00000007567.1"/>
        <property type="gene ID" value="ENSGAGG00000006049.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071797">
        <property type="term" value="C:LUBAC complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036435">
        <property type="term" value="F:K48-linked polyubiquitin modification-dependent protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070530">
        <property type="term" value="F:K63-linked polyubiquitin modification-dependent protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1990450">
        <property type="term" value="F:linear polyubiquitin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061630">
        <property type="term" value="F:ubiquitin protein ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097039">
        <property type="term" value="P:protein linear polyubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd19815">
        <property type="entry name" value="Bbox1_HOIP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.2190">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.1100">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.8.10">
        <property type="entry name" value="DNA helicase RuvA subunit, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047543">
        <property type="entry name" value="Bbox1_RNF31-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036339">
        <property type="entry name" value="PUB-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018997">
        <property type="entry name" value="PUB_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026254">
        <property type="entry name" value="RNF31-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032065">
        <property type="entry name" value="RNF31-UBA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040641">
        <property type="entry name" value="RNF31_PUB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000315">
        <property type="entry name" value="Znf_B-box"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001876">
        <property type="entry name" value="Znf_RanBP2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036443">
        <property type="entry name" value="Znf_RanBP2_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16004:SF5">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE RNF31"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16004">
        <property type="entry name" value="RING FINGER PROTEIN 31-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09409">
        <property type="entry name" value="PUB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18486">
        <property type="entry name" value="PUB_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16678">
        <property type="entry name" value="UBA_HOIP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00643">
        <property type="entry name" value="zf-B_box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00547">
        <property type="entry name" value="ZnF_RBZ"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF143503">
        <property type="entry name" value="PUG domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90209">
        <property type="entry name" value="Ran binding protein zinc finger-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50119">
        <property type="entry name" value="ZF_BBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01358">
        <property type="entry name" value="ZF_RANBP2_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50199">
        <property type="entry name" value="ZF_RANBP2_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0862">Zinc</keyword>
    <keyword evidence="2 4" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="B box-type" evidence="6">
        <location>
            <begin position="211"/>
            <end position="257"/>
        </location>
    </feature>
    <feature type="domain" description="RanBP2-type" evidence="7">
        <location>
            <begin position="296"/>
            <end position="326"/>
        </location>
    </feature>
    <feature type="domain" description="RanBP2-type" evidence="7">
        <location>
            <begin position="339"/>
            <end position="368"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="420"/>
            <end position="456"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00322"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50119"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50199"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007567.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="10">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="631" mass="68588" checksum="2608C3358E7BEF34" modified="2019-05-08" version="1">MAQAVPPLGEPQFLQLREELVQALARSPVDLHPETVQQLVEAPLPPHIKYREIDAQGILRANMQGQSPACLRTISTALNILEKYGRNLLNPLKPRYWRGVKFNNPVFRSTVDAIQGGRDVLRLYGYTEQQPDGLSFAEGLEEPDARRVASVTVDVVLLRAELNLLLSNNHPNPDTLQRMLAVGTGSRGALARGHEGLGCGVIPLSHPPGPPPRATCLLCGLEPASLHCPECNQSLCPECDRLFHKHPARAQHQRLPVGEPGLGAGPPAQSPPFPQGLGLSMVPSEPPVARPALPPKPRLPWRCGACHAPNEARAVLCVGCDRPRGCPTPPNPPPDGDPLRGAWACQTCTFLNTGPTVLCAVCERPRLAGRPGAPDPQHPPDGPPVPMQGAPGWQCEHCTFWNQLPGRVCEVCNRTXGAPGWQTSQLAGPPQEEKGPRRPGKPHPLSPEEVERRRQEKLREDGMKLVAMIREAELVGVPPEEVGAALRYSGTELPLPWLRSELPAVLDAVVEQATERGAGAELGAITQQEAHAAWTQSQGDLDEAVSRCLSTRRCKVRELAALGFAERGPVLQALYQNGGDMWGALTELQRLQLEPFHRRMWDLAEPEIDFHAPDRQVRARGRGTWGKSGGG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="22" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZF9</accession>
    <name>A0A452GZF9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">GAL3A protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="1 2">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="2" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="1" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="1" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GZF9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008843.1">
        <property type="protein sequence ID" value="ENSGAGP00000007661.1"/>
        <property type="gene ID" value="ENSGAGG00000006133.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="CDD" id="cd03133">
        <property type="entry name" value="GATase1_ES1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.880">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029062">
        <property type="entry name" value="Class_I_gatase-like"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF008747">
        <property type="entry name" value="PRK11780.1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10224">
        <property type="entry name" value="ES1 PROTEIN HOMOLOG, MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10224:SF5">
        <property type="entry name" value="GLUTAMINE AMIDOTRANSFERASE LIKE CLASS 1 DOMAIN CONTAINING 3A-LIKE1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52317">
        <property type="entry name" value="Class I glutamine amidotransferase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000313" key="1">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007661.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="3">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="225" mass="23915" checksum="72CC9351A48BF500" modified="2019-05-08" version="1">MGKRVALVLAGCGVFDGSEVHEASAALVHLSRGGAQVRIFAPNIEQMHVVDHFRGSPTEEKRNVLVESARLARGNIEDLAELKVGEFDALIFPGGFGVAKNLCSWAVDGKNCTVNELVKSTLQAFHSAKKPIGLCCISPVLAAKVFPGCEVTVGQDKNVDGRFPDAETASAIVELGCKHICKDVAESHVDATNKIVTTCAFMCKAPLHEIFDGIGTMVAEVLKLS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKY1</accession>
    <name>A0A452HKY1_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the TNFAIP8 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKY1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017806.1">
        <property type="protein sequence ID" value="ENSGAGP00000015590.1"/>
        <property type="gene ID" value="ENSGAGG00000011757.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042981">
        <property type="term" value="P:regulation of apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1440.160:FF:000001">
        <property type="entry name" value="Tumor necrosis factor alpha-induced protein 8-like 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1440.160">
        <property type="entry name" value="Tumor necrosis factor alpha-induced protein 8-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008477">
        <property type="entry name" value="TNFAIP8-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038355">
        <property type="entry name" value="TNFAIP8_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12757:SF5">
        <property type="entry name" value="TUMOR NECROSIS FACTOR ALPHA-INDUCED PROTEIN 8-LIKE PROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12757">
        <property type="entry name" value="TUMOR NECROSIS FACTOR INDUCED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05527">
        <property type="entry name" value="TNFAIP8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00038267"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015590.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="204" mass="23499" checksum="880E7C5F24F8F14E" modified="2019-05-08" version="1">MDSDSAELSEGELLAPAGPDCFSSKNLALQAQKKLLSKMATKTMANMLIDDTSSEIFDELYKVTREHTRNKKEAHKIMKDLIKVAIKIGILYRNNQFNQEELEIVDKFRKKLNQTAMTIVSFYEVEYTFDRNVLAELLNECKDLVHELVERHLTPRSHGRINHVFNHFADVEFLTALYSLDGDCRPHLKKICDGINKLLDEKVL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKY2</accession>
    <name>A0A452HKY2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12 16">Tyrosine-protein kinase</fullName>
            <ecNumber evidence="12 16">2.7.10.2</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="11 21">
            <text>L-tyrosyl-[protein] + ATP = O-phospho-L-tyrosyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10596"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.10.2"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endomembrane system</location>
            <topology evidence="1">Peripheral membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="12">Belongs to the protein kinase superfamily. Tyr protein kinase family. JAK subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.10.2" evidence="12 16"/>
    <dbReference type="AlphaFoldDB" id="A0A452HKY2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017811.1">
        <property type="protein sequence ID" value="ENSGAGP00000015594.1"/>
        <property type="gene ID" value="ENSGAGG00000011755.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012505">
        <property type="term" value="C:endomembrane system"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005131">
        <property type="term" value="F:growth hormone receptor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004715">
        <property type="term" value="F:non-membrane spanning protein tyrosine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019221">
        <property type="term" value="P:cytokine-mediated signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060397">
        <property type="term" value="P:growth hormone receptor signaling pathway via JAK-STAT"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035556">
        <property type="term" value="P:intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050865">
        <property type="term" value="P:regulation of cell activation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0022407">
        <property type="term" value="P:regulation of cell-cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd05080">
        <property type="entry name" value="PTKc_Tyk2_rpt2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000110">
        <property type="entry name" value="Tyrosine-protein kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000084">
        <property type="entry name" value="Tyrosine-protein kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000135">
        <property type="entry name" value="Tyrosine-protein kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000114">
        <property type="entry name" value="Tyrosine-protein kinase JAK2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.505.10">
        <property type="entry name" value="SH2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019749">
        <property type="entry name" value="Band_41_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035963">
        <property type="entry name" value="FERM_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000299">
        <property type="entry name" value="FERM_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041155">
        <property type="entry name" value="FERM_F1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041046">
        <property type="entry name" value="FERM_F2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051286">
        <property type="entry name" value="JAK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041381">
        <property type="entry name" value="JAK1-3/TYK2_PHL_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001245">
        <property type="entry name" value="Ser-Thr/Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000980">
        <property type="entry name" value="SH2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036860">
        <property type="entry name" value="SH2_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008266">
        <property type="entry name" value="Tyr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020635">
        <property type="entry name" value="Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016251">
        <property type="entry name" value="Tyr_kinase_non-rcpt_Jak/Tyk2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016045">
        <property type="entry name" value="Tyr_kinase_non-rcpt_TYK2_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45807:SF6">
        <property type="entry name" value="NON-RECEPTOR TYROSINE-PROTEIN KINASE TYK2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45807">
        <property type="entry name" value="TYROSINE-PROTEIN KINASE HOPSCOTCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18379">
        <property type="entry name" value="FERM_F1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18377">
        <property type="entry name" value="FERM_F2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17887">
        <property type="entry name" value="Jak1_Phl"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07714">
        <property type="entry name" value="PK_Tyr_Ser-Thr"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21990">
        <property type="entry name" value="SH2_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000636">
        <property type="entry name" value="TyrPK_Jak"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01823">
        <property type="entry name" value="JANUSKINASE"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00109">
        <property type="entry name" value="TYRKINASE"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01827">
        <property type="entry name" value="YKINASETYK2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00295">
        <property type="entry name" value="B41"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00252">
        <property type="entry name" value="SH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00219">
        <property type="entry name" value="TyrKc"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47031">
        <property type="entry name" value="Second domain of FERM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55550">
        <property type="entry name" value="SH2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50057">
        <property type="entry name" value="FERM_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00109">
        <property type="entry name" value="PROTEIN_KINASE_TYR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7 12 14" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="6 12 16" id="KW-0418">Kinase</keyword>
    <keyword evidence="9" id="KW-0472">Membrane</keyword>
    <keyword evidence="5 12 14" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="8" id="KW-0727">SH2 domain</keyword>
    <keyword evidence="3 12 16" id="KW-0808">Transferase</keyword>
    <keyword evidence="10 12 16" id="KW-0829">Tyrosine-protein kinase</keyword>
    <feature type="domain" description="FERM" evidence="18">
        <location>
            <begin position="1"/>
            <end position="380"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="17">
        <location>
            <begin position="538"/>
            <end position="820"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="17">
        <location>
            <begin position="842"/>
            <end position="1121"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="13">
        <location>
            <position position="968"/>
        </location>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <begin position="848"/>
            <end position="856"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="14 15">
        <location>
            <position position="875"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004184"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022999"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023137"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00051245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PIRNR" id="PIRNR000636"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PIRSR" id="PIRSR000636-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PIRSR" id="PIRSR000636-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="RuleBase" id="RU362096"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50057"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015594.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1132" mass="129052" checksum="4897C7E1CCCF6437" modified="2019-05-08" version="1">MHPCHGTGILLSPAGITPLCYSLFALCDVQARVWLPPNHVFEISKDTNLNLLFRMRYYFRNWHAMSDKEPAVYRNVPRQSDALEDKLQGGALLDKSSFEYLFEQGKYEFINDVVSLKDLQSEQEIHRFKNESLGMAVLHLSHMALKRGVSLEEVAKKNSFKGCIPRSFCRQIQQNSYLTKFRMRNVFRKFVQRFQRHTVSAGKLTEQDIMYKYLATLENLAPRFGSELFPVLSLETASEGEKVLLYVNGGHTQLEDGCASSPRDRSTTHEVLVTGTDGIQWRTVPVESAESLPHRGYFGRKTRAKELDLKGPCRPVERNEAKWVRFCDFQEITHIALEDCKVSINRQDNKCLELVLPSYETALSLVSLVDGYFRLTADSSHYLCHEVAPPRLVMSVLNGIHGPMQEEFVFAKLKREEQDGLYVIRWSILDFNRLILSVVKRDRGQATPKYRQFRIQKRGSSFVLEGWEREFSTVRELMDVLKGCTLKSGDESFAVRRCCPPKPTEISDLIIARRVKDSTRQILNLTQLSFHQIRKNEITQRAHLGQGTRTNIYEGVLHVCGGAGGDDEAEYFSTEQNNNSREMHVVLKVLDPSHRDIALAFFETASLMSQVSHIHLAFVHGVCVRGSENIMVEEYIEHGPLDVLLRKEKGRVTVGWKITVAKQLASALSYLEDKNLVHGNVCAKNILLARKGLENGLLPFVKLSDPGVSFTVLSREERVDRIPWIAPECVQDVGSLSTAADKWSFGTTLLEICFDADVPLKERTPSEKERFYEKKHRLPEPSCKELAALISQCLTYAPSERPSFRTILRDLTQLQPHNLVDVTSVSPHFPVSDPTVFQKRYLKKIRELGEGHFGKVSLYCYDPTNDGTGEMVAVKSLKSGCSQQLLTSWKREIEILKTLYHENIVKYKGCCSEQGEKIVQLIMEYVPLGSLRDYLPKHNVSLAHILLFAQQICEGMAYLHSLHYIHRDLAARNVLLENENVVKIGDFGLAKAIPEGHEYYRVCEDGDSPVFWYAMECLKECKFYYASDVWSFGVTLYELLTHCDSSQSPPVKFIEMIGATQGQMTVLRLIELLDRGKRLPSPKDCPCEIYCLMKNCWESEASFRPAFKNLVPILKNFHEKYKAQAPSVFSVC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLA1</accession>
    <name>A0A452HLA1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="22">Tyrosine-protein kinase Mer</fullName>
            <ecNumber evidence="3">2.7.10.1</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="23">Proto-oncogene c-Mer</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="24">Receptor tyrosine kinase MerTK</fullName>
        </alternativeName>
    </protein>
    <organism evidence="30 31">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="31" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="30" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="30" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="20">
            <text>L-tyrosyl-[protein] + ATP = O-phospho-L-tyrosyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10596"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.10.1"/>
        </reaction>
    </comment>
    <comment type="subunit">
        <text evidence="21">Interacts (upon activation) with TNK2; stimulates TNK2 autophosphorylation. Interacts (via N-terminus) with extracellular ligands LGALS3, TUB, TULP1 and GAS6. Interacts with VAV1 in a phosphotyrosine-independent manner. Interacts with TIMD4; this interaction enhances TIMD4-mediated efferocytosis.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the protein kinase superfamily. CAMK Ser/Thr protein kinase family.</text>
    </comment>
    <dbReference type="EC" id="2.7.10.1" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452HLA1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017943.1">
        <property type="protein sequence ID" value="ENSGAGP00000015714.1"/>
        <property type="gene ID" value="ENSGAGG00000010943.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043235">
        <property type="term" value="C:receptor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004714">
        <property type="term" value="F:transmembrane receptor protein tyrosine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007169">
        <property type="term" value="P:cell surface receptor protein tyrosine kinase signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006909">
        <property type="term" value="P:phagocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="CDD" id="cd00096">
        <property type="entry name" value="Ig"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000902">
        <property type="entry name" value="MER proto-oncogene, tyrosine kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:001050">
        <property type="entry name" value="MER proto-oncogene, tyrosine kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000834">
        <property type="entry name" value="Proto-oncogene tyrosine-protein kinase MER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000089">
        <property type="entry name" value="Tyrosine-protein kinase receptor TYRO3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000296">
        <property type="entry name" value="Tyrosine-protein kinase receptor TYRO3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000111">
        <property type="entry name" value="Tyrosine-protein kinase receptor TYRO3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013151">
        <property type="entry name" value="Immunoglobulin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050122">
        <property type="entry name" value="RTK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001245">
        <property type="entry name" value="Ser-Thr/Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008266">
        <property type="entry name" value="Tyr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020635">
        <property type="entry name" value="Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416:SF257">
        <property type="entry name" value="TYROSINE-PROTEIN KINASE MER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416">
        <property type="entry name" value="TYROSINE-PROTEIN KINASE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00047">
        <property type="entry name" value="ig"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13927">
        <property type="entry name" value="Ig_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07714">
        <property type="entry name" value="PK_Tyr_Ser-Thr"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00109">
        <property type="entry name" value="TYRKINASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00219">
        <property type="entry name" value="TyrKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00109">
        <property type="entry name" value="PROTEIN_KINASE_TYR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12 25" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="16" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="18" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="19" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="11" id="KW-0418">Kinase</keyword>
    <keyword evidence="14 26" id="KW-0472">Membrane</keyword>
    <keyword evidence="10 25" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="17" id="KW-0675">Receptor</keyword>
    <keyword evidence="31" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0677">Repeat</keyword>
    <keyword evidence="8" id="KW-0732">Signal</keyword>
    <keyword evidence="6" id="KW-0808">Transferase</keyword>
    <keyword evidence="7 26" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="13 26" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="15" id="KW-0829">Tyrosine-protein kinase</keyword>
    <feature type="transmembrane region" description="Helical" evidence="26">
        <location>
            <begin position="503"/>
            <end position="526"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="28">
        <location>
            <begin position="94"/>
            <end position="186"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="28">
        <location>
            <begin position="198"/>
            <end position="280"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="29">
        <location>
            <begin position="287"/>
            <end position="382"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="29">
        <location>
            <begin position="387"/>
            <end position="484"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="27">
        <location>
            <begin position="588"/>
            <end position="859"/>
        </location>
    </feature>
    <feature type="binding site" evidence="25">
        <location>
            <position position="620"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00011902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023137"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00051243"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00066004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00070212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00077110"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00082356"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="27">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="28">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="29">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="30">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015714.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="31">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="994" mass="111342" checksum="DDFC1978E59DC8DD" modified="2019-05-08" version="1">MPQSPGYFFSLAQCIYTKCNSFSWRDQPIPEQPRTGIGNLWHAARQGKHPGRPGWFVYLPLPQVRLIAAPRSRPMGAVGSTSAAHSLQQVKFNPTVGHILINEHKDFKFNCSIKGPQFLFNQDSAVISLWKDGKELQEPGRMSTRHYQVTDDVETTMVSTFSITNAQRSDNGSYSCKLKIHNEEIESDPIFVQMEGLPHFLRQPQSLNITRNRPFNLTCEAMGPPEPVQIYWFRNSSRVNDQPYISPSVLTLPGLSETAIFSCEAHNRKGLTASNEVQINIKRTPAAPADVRVLNRTAHSLVVSWVPGFDGYSPLHSCRLQVREAAALNKGSFLIWNTSVPPHRYHVQQLQPLAEYSIRVSCRNEVGWSSFSPWTLASTTEGAPSTPPLNVTVFFNESSSSLEVRWAKPHLGRKHGELQGYRLLHTWQNSKITQNRSSEAQQNATVTTVAIVVTNATYAVRVAAVTKGGIGPFSDPVVVFIPGDGFMASAPSSTPAPGNNHSFAIVLGFILGIVAIGLILYFSVIVRKRWMETKFGTAFHSDDSEHVVSYTAKKSYSRRAVELTLASLGVREELQQKLQDVVIDRDSLALGKILGEGEFGSVMEGHLSQPSGTVQKVAVKTMKLEDFSQREIEEFLSEAACMKDFDHPNVIRLLGVCIELSSRQVPKPMVVLPFMKYGDLHSFLLRSRFETGTRHVPLQTLLKFMVDIALGMEYLSTRQFLHRDLAARNCMLRDDMTVCVADFGLSKKIYSGDYYRQGRIAKMPVKWIAIESLADRVYTTKSDVWAFGVTMWEIATRGMTPYPGVQNHEIYDYLLHGHRLKQPEDCLDELYEVMYSCWRADPHERPAFPELKLHLEKLLESLPKVSGSEDVIYINTSLPEKSEELTEESEFPQTDMAFNRNYVTVSSSPKPDTTVVTAEVHESDQREERYILKGETDEQTAGSLTLEQGHAPLLQGTITRNGATWGPASMLLGNSLADELLYADDCSEDSEVLL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLA5</accession>
    <name>A0A452HLA5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Activation-induced cytidine deaminase AID domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HLA5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017934.1">
        <property type="protein sequence ID" value="ENSGAGP00000015705.1"/>
        <property type="gene ID" value="ENSGAGG00000011839.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004126">
        <property type="term" value="F:cytidine deaminase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="project" value="TreeGrafter"/>
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        <property type="project" value="InterPro"/>
    </dbReference>
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        <property type="term" value="P:cytidine to uridine editing"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="entry name" value="Cytidine Deaminase, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
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        <property type="entry name" value="MRNA EDITING ENZYME"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="SINGLE-STRANDED DNA CYTOSINE DEAMINASE"/>
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    </dbReference>
    <dbReference type="Pfam" id="PF08210">
        <property type="entry name" value="APOBEC_N"/>
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    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="1" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Activation-induced cytidine deaminase AID" evidence="3">
        <location>
            <begin position="11"/>
            <end position="53"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="Pfam" id="PF08210"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015705.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="69" mass="8264" checksum="F2BB04BE981CB064" modified="2019-05-08" version="1">MDSLLMKQKKFLYNFKNLRWARGRHETYLCYVVKRRDSATSCSLDFGYLRNKPLDEVDDLRDAFKILGL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLA7</accession>
    <name>A0A452HLA7_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="16">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000017948.1">
        <property type="protein sequence ID" value="ENSGAGP00000015719.1"/>
        <property type="gene ID" value="ENSGAGG00000011232.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003682">
        <property type="term" value="F:chromatin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043565">
        <property type="term" value="F:sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003714">
        <property type="term" value="F:transcription corepressor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd04709">
        <property type="entry name" value="BAH_MTA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd11661">
        <property type="entry name" value="SANT_MTA3_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00202">
        <property type="entry name" value="ZnF_GATA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.60:FF:000052">
        <property type="entry name" value="Arginine-glutamic acid dipeptide (RE) repeats"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.490:FF:000005">
        <property type="entry name" value="Arginine-glutamic acid dipeptide (RE) repeats"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.60">
        <property type="entry name" value="Homeodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002951">
        <property type="entry name" value="Atrophin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001025">
        <property type="entry name" value="BAH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043151">
        <property type="entry name" value="BAH_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000949">
        <property type="entry name" value="ELM2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009057">
        <property type="entry name" value="Homeodomain-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001005">
        <property type="entry name" value="SANT/Myb"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017884">
        <property type="entry name" value="SANT_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000679">
        <property type="entry name" value="Znf_GATA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13859:SF12">
        <property type="entry name" value="ARGININE-GLUTAMIC ACID DIPEPTIDE REPEATS PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13859">
        <property type="entry name" value="ATROPHIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03154">
        <property type="entry name" value="Atrophin-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01426">
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00320">
        <property type="entry name" value="GATA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00249">
        <property type="entry name" value="Myb_DNA-binding"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="ZnF_GATA"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Glucocorticoid receptor-like (DNA-binding domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="ELM2"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8" id="KW-0007">Acetylation</keyword>
    <keyword evidence="2" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0804">Transcription</keyword>
    <keyword evidence="9" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="7" id="KW-0832">Ubl conjugation</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="BAH" evidence="13">
        <location>
            <begin position="103"/>
            <end position="256"/>
        </location>
    </feature>
    <feature type="domain" description="ELM2" evidence="14">
        <location>
            <begin position="264"/>
            <end position="312"/>
        </location>
    </feature>
    <feature type="domain" description="SANT" evidence="15">
        <location>
            <begin position="316"/>
            <end position="368"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1"/>
            <end position="90"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
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            <end position="420"/>
        </location>
    </feature>
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            <end position="1057"/>
        </location>
    </feature>
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        <location>
            <begin position="1100"/>
            <end position="1164"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="1"/>
            <end position="36"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="12">
        <location>
            <begin position="74"/>
            <end position="85"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="390"/>
            <end position="399"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
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            <end position="413"/>
        </location>
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            <end position="565"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
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        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
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        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
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        </location>
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    <feature type="compositionally biased region" description="Low complexity" evidence="12">
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        </location>
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        <location>
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        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="12">
        <location>
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        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
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            <end position="765"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="12">
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        </location>
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        <location>
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        </location>
    </feature>
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        <location>
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        </location>
    </feature>
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        </location>
    </feature>
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        <location>
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            <end position="926"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
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            <end position="950"/>
        </location>
    </feature>
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        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
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        </location>
    </feature>
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    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
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        </location>
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        <source>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS51038"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS51156"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS51293"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015719.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1490" mass="165834" checksum="78097C37DC048981" modified="2019-05-08" version="1">MTADKEKDKDKEKDRDRDRDRERDKREKVRESENSRPRRSCTLEGGAKNYAESDHSEDEDNDNNSATTEESTKKNKKKPPKKKSRYERTDSGEITSFITEDDVVYRPGDCVYIESRRPNTPYFICSIQDFKLVSSFLFSNALDLLFQSKRDHLLMNVKWYYRQSEVPDSVYQHLVQDRHNENDSGRELVITDPVIKNRELFISDYVDTYHAAALRGKCNISHFSDIFAAREFKARMDSFFYILGYNPETRRLNSTQGEIRVGPTFAGMCDGGSTEDGCVAASRDDTTLNALNTLHESNYDAGKALQRLVKKPVPKLIEKCWTEDEVKRFIKGLRQYGKNFFRIRKELLPNKETGELITFYYYWKKTPEAASSRAHRRHRRQAVFRRIKTRTASTPVNTPSRPPSSEFLDLSSASEDDFDSEDSEQELKGYACRHCFTTTSKDWHHGGRENILLCTDCRIHFKKYGELPPIEKPVDPPPFMFKPVKEEDDGLSGKHSMRTRRSRGSMSTLRSGRKKQPASPDGRASPTNEDIRSSGRNSPSAASTSSNDSKADSVKKSTKKIKEEASSPLKSSKRQREKTASDTDEPERTTAKKSKTQEISRPNSPSEGEGESSDSRSVNDDGSSDPKDIDQDNRSTSPSIPSPQDNESDSDSSAQQQMLQAQPQVLQAQNVPAQAPPSTLPVSTQLPTPIPTASSTPTAPLQVSLSVSQPPSHPPAPPPAHSHIQQAPALHPQRLPSPHPPLQPLTVSQSQPQGPASSQPHSQPPLHGQVQPVPHNLQAQPLLPHPVPPQPFPLPSQPSQSQVPLQTQAASHPHAALQVQAVQPALQAQAPLQQPQPPREQPLPPAPMAMPHIKPPPTTPIPQLPTPQSHKHPPHLSGPSPFSMNSNLPPPPALKPLSSLSTHHPPSAHPPPLQLMPQSQPLQSSPAQPPVLTQSQSLPPAASHPPSSLHQVSSQAPFSQHPFVPGGPPSITPPSCPSTSTPSSVPGVPSQTSVATSAASNGNIPVVTSCAIPPIQIKEEAPDEAEEPESPPPPPRSPSPEPTVVDTPSHASQSARFYKHLDRGYNSCARTDLYFMPLAGSKLAKKREEAVEKAKREAEQKAREEREREKEKEKEREREREREREAERAAKVSSSSHEGRLGEPQLTGPTHMRPSFEPPPTTIAAVPPYIGPDTPALRTLSEYARPHVMSPTNRNHPFFVPLNPTDPLLAYHMPGLYNVDPTIRERELREREIREREIRERELRERMKPGFEVKPPELDALHPATNPMEHFARHGALTIPPTAGPHPFASFHPGLNPLERERLALAGPQLRPEMSYPDRLAAERIHAERMASLTNDPLARLQMFNVTPHHHQHSHIHSHLHLHQQDPLHQGSAGPVHPLVDPLAAGPHLARFPYPPGTIPNPLLGQPPHEHEMLRHPVFGTPYPRDLPGAIPPPMSAAHQLQAMHAQSAELQRLAMEQQWLHGHPHMHGGHLPSQEDYYSRLKKEGDKQL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLA8</accession>
    <name>A0A452HLA8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">FAD-binding PCMH-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="1 15">
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="16">
            <name>Mo-molybdopterin</name>
            <dbReference type="ChEBI" id="CHEBI:71302"/>
        </cofactor>
        <text evidence="16">Binds 1 Mo-molybdopterin (Mo-MPT) cofactor per subunit.</text>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="13">
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="16">
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:49601"/>
        </cofactor>
        <text evidence="16">Binds 2 [2Fe-2S] clusters.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the xanthine dehydrogenase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLA8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017968.1">
        <property type="protein sequence ID" value="ENSGAGP00000015738.1"/>
        <property type="gene ID" value="ENSGAGG00000010622.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051537">
        <property type="term" value="F:2 iron, 2 sulfur cluster binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071949">
        <property type="term" value="F:FAD binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005506">
        <property type="term" value="F:iron ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016491">
        <property type="term" value="F:oxidoreductase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.120:FF:000001">
        <property type="entry name" value="Aldehyde oxidase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.365.10:FF:000003">
        <property type="entry name" value="Aldehyde oxidase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.365.10:FF:000025">
        <property type="entry name" value="Aldehyde oxidase 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.365.10:FF:000001">
        <property type="entry name" value="Xanthine dehydrogenase oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.365.10:FF:000004">
        <property type="entry name" value="Xanthine dehydrogenase oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.390.50:FF:000001">
        <property type="entry name" value="Xanthine dehydrogenase oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.43.10:FF:000001">
        <property type="entry name" value="Xanthine dehydrogenase/oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.465.10:FF:000004">
        <property type="entry name" value="Xanthine dehydrogenase/oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.20.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.465.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.120">
        <property type="entry name" value="[2Fe-2S]-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1170.50">
        <property type="entry name" value="Aldehyde oxidase/xanthine dehydrogenase, a/b hammerhead"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.365.10">
        <property type="entry name" value="Aldehyde oxidase/xanthine dehydrogenase, molybdopterin binding domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.390.50">
        <property type="entry name" value="CO dehydrogenase flavoprotein, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.43.10">
        <property type="entry name" value="Uridine Diphospho-n-acetylenolpyruvylglucosamine Reductase, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002888">
        <property type="entry name" value="2Fe-2S-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036884">
        <property type="entry name" value="2Fe-2S-bd_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036010">
        <property type="entry name" value="2Fe-2S_ferredoxin-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001041">
        <property type="entry name" value="2Fe-2S_ferredoxin-type"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006058">
        <property type="entry name" value="2Fe2S_fd_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000674">
        <property type="entry name" value="Ald_Oxase/Xan_DH_a/b"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036856">
        <property type="entry name" value="Ald_Oxase/Xan_DH_a/b_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016208">
        <property type="entry name" value="Ald_Oxase/xanthine_DH-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008274">
        <property type="entry name" value="AldOxase/xan_DH_MoCoBD1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046867">
        <property type="entry name" value="AldOxase/xan_DH_MoCoBD2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037165">
        <property type="entry name" value="AldOxase/xan_DH_Mopterin-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012675">
        <property type="entry name" value="Beta-grasp_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005107">
        <property type="entry name" value="CO_DH_flav_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036683">
        <property type="entry name" value="CO_DH_flav_C_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016166">
        <property type="entry name" value="FAD-bd_PCMH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036318">
        <property type="entry name" value="FAD-bd_PCMH-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016167">
        <property type="entry name" value="FAD-bd_PCMH_sub1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016169">
        <property type="entry name" value="FAD-bd_PCMH_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002346">
        <property type="entry name" value="Mopterin_DH_FAD-bd"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45444">
        <property type="entry name" value="XANTHINE DEHYDROGENASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45444:SF3">
        <property type="entry name" value="XANTHINE DEHYDROGENASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03450">
        <property type="entry name" value="CO_deh_flav_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00941">
        <property type="entry name" value="FAD_binding_5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00111">
        <property type="entry name" value="Fer2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01799">
        <property type="entry name" value="Fer2_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02738">
        <property type="entry name" value="MoCoBD_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20256">
        <property type="entry name" value="MoCoBD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000127">
        <property type="entry name" value="Xanthine_DH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01008">
        <property type="entry name" value="Ald_Xan_dh_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01092">
        <property type="entry name" value="CO_deh_flav_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54292">
        <property type="entry name" value="2Fe-2S ferredoxin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55447">
        <property type="entry name" value="CO dehydrogenase flavoprotein C-terminal domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47741">
        <property type="entry name" value="CO dehydrogenase ISP C-domain like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54665">
        <property type="entry name" value="CO dehydrogenase molybdoprotein N-domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56176">
        <property type="entry name" value="FAD-binding/transporter-associated domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56003">
        <property type="entry name" value="Molybdenum cofactor-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00197">
        <property type="entry name" value="2FE2S_FER_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51387">
        <property type="entry name" value="FAD_PCMH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7 16" id="KW-0001">2Fe-2S</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9 15" id="KW-0274">FAD</keyword>
    <keyword evidence="6" id="KW-0285">Flavoprotein</keyword>
    <keyword evidence="11 16" id="KW-0408">Iron</keyword>
    <keyword evidence="12 16" id="KW-0411">Iron-sulfur</keyword>
    <keyword evidence="8 16" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="5 16" id="KW-0500">Molybdenum</keyword>
    <keyword evidence="10" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="FAD-binding PCMH-type" evidence="17">
        <location>
            <begin position="212"/>
            <end position="397"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="14">
        <location>
            <position position="1217"/>
        </location>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="38"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="43"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="46"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="68"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="108"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>2</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="111"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>2</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="143"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>2</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="145"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>2</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <begin position="240"/>
            <end position="247"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="343"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="405"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="723"/>
        </location>
        <ligand>
            <name>Mo-molybdopterin</name>
            <dbReference type="ChEBI" id="CHEBI:71302"/>
        </ligand>
        <ligandPart>
            <name>Mo</name>
            <dbReference type="ChEBI" id="CHEBI:28685"/>
        </ligandPart>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="754"/>
        </location>
        <ligand>
            <name>Mo-molybdopterin</name>
            <dbReference type="ChEBI" id="CHEBI:71302"/>
        </ligand>
        <ligandPart>
            <name>Mo</name>
            <dbReference type="ChEBI" id="CHEBI:28685"/>
        </ligandPart>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="868"/>
        </location>
        <ligand>
            <name>Mo-molybdopterin</name>
            <dbReference type="ChEBI" id="CHEBI:71302"/>
        </ligand>
        <ligandPart>
            <name>Mo</name>
            <dbReference type="ChEBI" id="CHEBI:28685"/>
        </ligandPart>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="870"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="1035"/>
        </location>
        <ligand>
            <name>Mo-molybdopterin</name>
            <dbReference type="ChEBI" id="CHEBI:71302"/>
        </ligand>
        <ligandPart>
            <name>Mo</name>
            <dbReference type="ChEBI" id="CHEBI:28685"/>
        </ligandPart>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001974"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00006849"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022505"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022630"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022714"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022827"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023014"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00034078"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PIRSR" id="PIRSR000127-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PIRSR" id="PIRSR000127-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PIRSR" id="PIRSR000127-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS51387"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015738.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1273" mass="141094" checksum="8D8C9728BDA6CB5D" modified="2019-05-08" version="1">APLHGERDRHHSNNTENQQLFSLTAAFAVRLSGTKYGCGIGGCGACTVMISTYEPTSKKIMHYPANSCLLPICSLHSSAVTTVEGVGSTKTRVHPIQERLAKCHGSQCGFCTPGMVMSMYALLRNHPEPSMEQITAALDGNLCRCTGYRPIIDTYKSFTGVGRKTNRSPFSKMHSGLCNPEEFQPVDPTQEFIFPPELMRMAQEQQKRTLVFRGERTTWISPTNLRELLELKAMYPKAPLIVGNTSVGLEMKRDGVHHPVFIHPTRIPDLHVVSNTNNGLVIGAACSLAQLRDILMKVVPELPEEKTKIFRALLQQLKTLAGEQIRSLASLGGHIVSRGSTWDLNPVLAVGNSVLSLASKDGTRQIPLNDQFLAGYENADLKPKEIIVSVFIPYSKKDDFVSAFRQAERRRNAVSVVNSGMRVLFKPNTDIIMDMNILYGGTGPITVSARKSCQRLIGRQWNEQMLDEACKLVLGEISLSPSAPGGKVEYRRTLLVSFLFKFYLEVLYGLNKMYPAKFPDLSKKNLSALGAFHSEVPQGMQTYQDVDPVQPPQDPVGRPIMHQSGVKHATGEAVYCNDIRAVDEELFLAVVTINKRDVRKMLQVFTSLYVLGVLQVICVGQIVCAVVAESAVQAKQGVEKMKIEYEDLEPILTIEDAIKHNSYFGKEKKIEQGNIEEGFKSADEIIEGEIHIGGQEHFYLETNSVLVVPRGEDNEMDVHVSTQDAARVQEFVALALDVQSSKIMCHTKRVGGAFGGKITKPAIFAVVAAVAANKTGRPIRFTLERDEDMLITGGRHPFFGKYKVGFTNDGRITAADIEPYINAGCTLDDSEMVIEYVVLKTDNAYKIPNLRIRGRACKTNLPSNTAFRGFGFPQSGLFTESWITTVATQTGLPQEKIREINMYKEVNLTPFKEELDATNLVKCWKECMEKSEYYSRKTAVEEFNKQNYWKKKGIAIIPMKFSVGFNATQYHQAAALVHIYTDGSVLVTHGGIELGQGIHTKMLQIASRELKVPLSYVHFCETSTITVPNAIVTAGSIGSDVNGKAVQNACQILRKRLEPIINKNPEGKWEDWITEAYDQSVSLSATGYFRGYMVNMNWDTGEGQAFPYFVFGAACSEVEIDCLTGDHKNIRTDIVMDACFSINPAVDIGQIEGAFVQGVGLYTMEEIKFSPEGEQYTLGPDTYKIPAVCDIPEQFRVYLLPNSRNPIAIYSAKGIGEAGLFLGCSVFFAIWDAVAAVRKERGLTGTFTLNSPLTIERIRMACADNFTEMVMVF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLA9</accession>
    <name>A0A452HLA9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="12">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLA9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017953.1">
        <property type="protein sequence ID" value="ENSGAGP00000015723.1"/>
        <property type="gene ID" value="ENSGAGG00000011232.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043565">
        <property type="term" value="F:sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003714">
        <property type="term" value="F:transcription corepressor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd11661">
        <property type="entry name" value="SANT_MTA3_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00202">
        <property type="entry name" value="ZnF_GATA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.60:FF:000052">
        <property type="entry name" value="Arginine-glutamic acid dipeptide (RE) repeats"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.1240.50">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.60">
        <property type="entry name" value="Homeodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002951">
        <property type="entry name" value="Atrophin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000949">
        <property type="entry name" value="ELM2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009057">
        <property type="entry name" value="Homeodomain-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001005">
        <property type="entry name" value="SANT/Myb"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017884">
        <property type="entry name" value="SANT_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000679">
        <property type="entry name" value="Znf_GATA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13859:SF12">
        <property type="entry name" value="ARGININE-GLUTAMIC ACID DIPEPTIDE REPEATS PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13859">
        <property type="entry name" value="ATROPHIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03154">
        <property type="entry name" value="Atrophin-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00320">
        <property type="entry name" value="GATA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00249">
        <property type="entry name" value="Myb_DNA-binding"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01217">
        <property type="entry name" value="PRICHEXTENSN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00717">
        <property type="entry name" value="SANT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00401">
        <property type="entry name" value="ZnF_GATA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57716">
        <property type="entry name" value="Glucocorticoid receptor-like (DNA-binding domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46689">
        <property type="entry name" value="Homeodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51156">
        <property type="entry name" value="ELM2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51293">
        <property type="entry name" value="SANT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0007">Acetylation</keyword>
    <keyword evidence="2" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0804">Transcription</keyword>
    <keyword evidence="6" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="4" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="ELM2" evidence="10">
        <location>
            <begin position="1"/>
            <end position="58"/>
        </location>
    </feature>
    <feature type="domain" description="SANT" evidence="11">
        <location>
            <begin position="62"/>
            <end position="114"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="135"/>
            <end position="166"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="213"/>
            <end position="803"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="846"/>
            <end position="910"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="136"/>
            <end position="145"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="150"/>
            <end position="159"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="280"/>
            <end position="294"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="295"/>
            <end position="311"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="323"/>
            <end position="344"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="359"/>
            <end position="379"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="380"/>
            <end position="391"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="400"/>
            <end position="419"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="437"/>
            <end position="446"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="9">
        <location>
            <begin position="457"/>
            <end position="466"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="494"/>
            <end position="511"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="9">
        <location>
            <begin position="529"/>
            <end position="542"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="543"/>
            <end position="554"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="561"/>
            <end position="579"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="9">
        <location>
            <begin position="580"/>
            <end position="611"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="641"/>
            <end position="651"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="661"/>
            <end position="672"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="680"/>
            <end position="696"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="9">
        <location>
            <begin position="711"/>
            <end position="722"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="723"/>
            <end position="740"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="9">
        <location>
            <begin position="776"/>
            <end position="787"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="846"/>
            <end position="876"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS51156"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS51293"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015723.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1236" mass="136062" checksum="222834EB2CEB9B9C" modified="2019-05-08" version="1">RREYNESMAAFAGMCDGGSTEDGCVAASRDDTTLNALNTLHESNYDAGKALQRLVKKPVPKLIEKCWTEDEVKRFIKGLRQYGKNFFRIRKELLPNKETGELITFYYYWKKTPEAASSRAHRRHRRQAVFRRIKTRTASTPVNTPSRPPSSEFLDLSSASEDDFDSEDSEQELKGYACRHCFTTTSKDWHHGGRENILLCTDCRIHFKKYGELPPIEKPVDPPPFMFKPVKEEDDGLSGKHSMRTRRSRGSMSTLRSGRKKQPASPDGRASPTNEDIRSSGRNSPSAASTSSNDSKADSVKKSTKKIKEEASSPLKSSKRQREKTASDTDEPERTTAKKSKTQEISRPNSPSEGEGESSDSRSVNDDGSSDPKDIDQDNRSTSPSIPSPQDNESDSDSSAQQQMLQAQPQVLQAQNVPAQAPPSTLPVSTQLPTPIPTASSTPTAPLQVSLSVSQPPSHPPAPPPAHSHIQQAPALHPQRLPSPHPPLQPLTVSQSQPQGPASSQPHSQPPLHGQVQPVPHNLQAQPLLPHPVPPQPFPLPSQPSQSQVPLQTQAASHPHAALQVQAVQPALQAQAPLQQPQPPREQPLPPAPMAMPHIKPPPTTPIPQLPTPQSHKHPPHLSGPSPFSMNSNLPPPPALKPLSSLSTHHPPSAHPPPLQLMPQSQPLQSSPAQPPVLTQSQSLPPAASHPPSSLHQVSSQAPFSQHPFVPGGPPSITPPSCPSTSTPSSVPGVPSQTSVATSAASNGNIPVVTSCAIPPIQIKEEAPDEAEEPESPPPPPRSPSPEPTVVDTPSHASQSARFYKHLDRGYNSCARTDLYFMPLAGSKLAKKREEAVEKAKREAEQKAREEREREKEKEKEREREREREREAERAAKVSSSSHEGRLGEPQLTGPTHMRPSFEPPPTTIAAVPPYIGPDTPALRTLSEYARPHVMSPTNRNHPFFVPLNPTDPLLAYHMPGLYNVDPTIRERELREREIREREIRERELRERMKPGFEVKPPELDALHPATNPMEHFARHGALTIPPTAGPHPFASFHPGLNPLERERLALAGPQLRPEMSYPDRLAAERIHAERMASLTNDPLARLQMFNVTPHHHQHSHIHSHLHLHQQDPLHQGSAGPVHPLVDPLAAGPHLARFPYPPGTIPNPLLGQPPHEHEMLRHPVFGTPYPRDLPGAIPPPMSAAHQLQAMHAQSAELQRLAMEQQWLHGHPHMHGGHLPSQEDYYSRLKKEGDKQL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLB2</accession>
    <name>A0A452HLB2_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="16">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
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    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
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                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
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        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
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        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
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        <scope>IDENTIFICATION</scope>
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        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
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    <sequence length="1171" mass="129752" checksum="8E512B645436275C" modified="2019-05-08" version="1">MTADKEKDKDKEKDRDRDRDRERDKREKVRESENSRPRRSCTLEGGAKNYAESDHSEDEDNDNNSATTEESTKKNKKKPPKKKSRYERTDSGEITSFITEDDVVYRPGDCVYIESRRPNTPYFICSIQDFKLVSTCTLPWSRGPPGISFLFSNALDLLFQSKRDHLLMNVKWYYRQSEVPDSVYQHLVQDRHNENDSGRELVITDPVIKNRELFISDYVDTYHAAALRGKCNISHFSDIFAAREFKARMDSFFYILGYNPETRRLNSTQGEIRVGPSHQVKDLIVFIDDFMAAFAGMCDGGSTEDGCVAASRDDTTLNALNTLHESNYDAGKALQRLVKKPVPKLIEKCWTEDEVKRFIKGLRQYGKNFFRIRKELLPNKETGELITFYYYWKKTPEAASSRAHRRHRRQAVFRRIKTRTASTPVNTPSRPPSSEFLDLSSASEDDFDSEDSEQELKGYACRHCFTTTSKDWHHGGRENILLCTDCRIHFKKYGELPPIEKPVDPPPFMFKPVKEEDDGLSGKHSMRTRRSRGSMSTLRSGRKKQPASPDGRASPTNEDIRSSGRNSPSAASTSSNDSKADSVKKSTKKIKEEASSPLKSSKRQREKTASDTDEPERTTAKKSKTQEISRPNSPSEGEGESSDSRSVNDDGSSDPKDIDQDNRSTSPSIPSPQDNESDSDSSAQQQMLQAQPQVLQAQNVPAQAPPSTLPVSTQLPTPIPTASSTPTAPLQVSLSVSQPPSHPPAPPPAHSHIQQAPALHPQRLPSPHPPLQPLTVSQSQPQGPASSQPHSQPPLHGQVQPVPHNLQAQPLLPHPVPPQPFPLPSQPSQSQVPLQTQAASHPHAALQVQAVQPALQAQAPLQQPQPPREQPLPPAPMAMPHIKPPPTTPIPQLPTPQSHKHPPHLSGPSPFSMNSNLPPPPALKPLSSLSTHHPPSAHPPPLQLMPQSQPLQSSPAQPPVLTQSQSLPPAASHPPSSLHQVSSQAPFSQHPFVPGGPPSITPPSCPSTSTPSSVPGVPSQTSVATSAASNGNIPVVTSCAIPPIQIKEEAPDEAEEPESPPPPPRSPSPEPTVVDTPSHASQSARFYKHLDRGYNSCARTDLYFMPLAGSKLAKKREEAVEKAKREAEQKAREEREREKEKEKEREREREREREAERAAVGLSVCRAGWVR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLB3</accession>
    <name>A0A452HLB3_9SAUR</name>
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        <recommendedName>
            <fullName evidence="20">Tyrosine-protein kinase Mer</fullName>
            <ecNumber evidence="3">2.7.10.1</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="21">Proto-oncogene c-Mer</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="22">Receptor tyrosine kinase MerTK</fullName>
        </alternativeName>
    </protein>
    <organism evidence="31 32">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="32" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="31" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="31" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>L-tyrosyl-[protein] + ATP = O-phospho-L-tyrosyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10596"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.10.1"/>
        </reaction>
    </comment>
    <comment type="subunit">
        <text evidence="19">Interacts (upon activation) with TNK2; stimulates TNK2 autophosphorylation. Interacts (via N-terminus) with extracellular ligands LGALS3, TUB, TULP1 and GAS6. Interacts with VAV1 in a phosphotyrosine-independent manner. Interacts with TIMD4; this interaction enhances TIMD4-mediated efferocytosis.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the protein kinase superfamily. CAMK Ser/Thr protein kinase family.</text>
    </comment>
    <dbReference type="EC" id="2.7.10.1" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452HLB3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017955.1">
        <property type="protein sequence ID" value="ENSGAGP00000015725.1"/>
        <property type="gene ID" value="ENSGAGG00000010943.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
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    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043235">
        <property type="term" value="C:receptor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004714">
        <property type="term" value="F:transmembrane receptor protein tyrosine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007169">
        <property type="term" value="P:cell surface receptor protein tyrosine kinase signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006909">
        <property type="term" value="P:phagocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="CDD" id="cd00096">
        <property type="entry name" value="Ig"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000902">
        <property type="entry name" value="MER proto-oncogene, tyrosine kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:001050">
        <property type="entry name" value="MER proto-oncogene, tyrosine kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000834">
        <property type="entry name" value="Proto-oncogene tyrosine-protein kinase MER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000089">
        <property type="entry name" value="Tyrosine-protein kinase receptor TYRO3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000296">
        <property type="entry name" value="Tyrosine-protein kinase receptor TYRO3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000509">
        <property type="entry name" value="Tyrosine-protein kinase receptor UFO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
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    <dbReference type="InterPro" id="IPR013151">
        <property type="entry name" value="Immunoglobulin_dom"/>
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    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050122">
        <property type="entry name" value="RTK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001245">
        <property type="entry name" value="Ser-Thr/Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008266">
        <property type="entry name" value="Tyr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020635">
        <property type="entry name" value="Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416:SF257">
        <property type="entry name" value="TYROSINE-PROTEIN KINASE MER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416">
        <property type="entry name" value="TYROSINE-PROTEIN KINASE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00047">
        <property type="entry name" value="ig"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13927">
        <property type="entry name" value="Ig_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07714">
        <property type="entry name" value="PK_Tyr_Ser-Thr"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000615">
        <property type="entry name" value="TyrPK_CSF1-R"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00109">
        <property type="entry name" value="TYRKINASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00219">
        <property type="entry name" value="TyrKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00109">
        <property type="entry name" value="PROTEIN_KINASE_TYR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10 24 26" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="14" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="16" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="17" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="9" id="KW-0418">Kinase</keyword>
    <keyword evidence="25" id="KW-0460">Magnesium</keyword>
    <keyword evidence="12 27" id="KW-0472">Membrane</keyword>
    <keyword evidence="25" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8 24 26" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="15" id="KW-0675">Receptor</keyword>
    <keyword evidence="32" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0808">Transferase</keyword>
    <keyword evidence="6 27" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 27" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="13" id="KW-0829">Tyrosine-protein kinase</keyword>
    <feature type="transmembrane region" description="Helical" evidence="27">
        <location>
            <begin position="459"/>
            <end position="482"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="27">
        <location>
            <begin position="606"/>
            <end position="628"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="29">
        <location>
            <begin position="50"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="29">
        <location>
            <begin position="154"/>
            <end position="236"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="30">
        <location>
            <begin position="243"/>
            <end position="338"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="30">
        <location>
            <begin position="343"/>
            <end position="440"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="28">
        <location>
            <begin position="544"/>
            <end position="801"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="23">
        <location>
            <position position="666"/>
        </location>
    </feature>
    <feature type="binding site" evidence="26">
        <location>
            <position position="576"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="24">
        <location>
            <position position="670"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="25">
        <location>
            <position position="671"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="25">
        <location>
            <position position="684"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00011902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023137"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00051243"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00066004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00070212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00077110"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00082356"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="PIRSR" id="PIRSR000615-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="PIRSR" id="PIRSR000615-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="PIRSR" id="PIRSR000615-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="28">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="29">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="30">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="31">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015725.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="32">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="936" mass="104749" checksum="57CE9A2FDEEE7C77" modified="2019-05-08" version="1">MSKVVNLGLILQRTLRRGRRGGHPTCCHPRKHSRWTASAAHSLQQVKFNPTVGHILINEHKDFKFNCSIKGPQFLFNQDSAVISLWKDGKELQEPGRMSTRHYQVTDDVETTMVSTFSITNAQRSDNGSYSCKLKIHNEEIESDPIFVQMEGLPHFLRQPQSLNITRNRPFNLTCEAMGPPEPVQIYWFRNSSRVNDQPYISPSVLTLPGLSETAIFSCEAHNRKGLTASNEVQINIKRTPAAPADVRVLNRTAHSLVVSWVPGFDGYSPLHSCRLQVREAAALNKGSFLIWNTSVPPHRYHVQQLQPLAEYSIRVSCRNEVGWSSFSPWTLASTTEGAPSTPPLNVTVFFNESSSSLEVRWAKPHLGRKHGELQGYRLLHTWQNSKITQNRSSEAQQNATVTTVAIVVTNATYAVRVAAVTKGGIGPFSDPVVVFIPGDGFMASAPSSTPAPGNNHSFAIVLGFILGIVAIGLILYFSVIVRKRWMETKFGTAFHSDDSEHVVSYTAKKSYSRRAVELTLASLGVREELQQKLQDVVIDRDSLALGKILGEGEFGSVMEGHLSQPSGTVQKVAVKTMKLEDFSQREIEEFLSEAACMKDFDHPNVIRLLVALTPFGLHGTFTCFLYVQTSSLSSQHVPLQTLLKFMVDIALGMEYLSTRQFLHRDLAARNCMLRDDMTVCVADFGLSKKIYSGDYYRQGRIAKMPVKWIAIESLADRVYTTKSDVWAFGVTMWEIATRGMTPYPGVQNHEIYDYLLHGHRLKQPEDCLDELYEVMYSCWRADPHERPAFPELKLHLEKLLESLPKVSGSEDVIYINTSLPEKSEELTEESEFPQTDMAFNRNYVTVSSSPKPDTTVVTAEVHESDQREERYILKGETDEQTAGSLTLEQGHAPLLQGTITRNGATWGPASMLLGNSLADELLYADDCSEDSEVLL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLB7</accession>
    <name>A0A452HLB7_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="17">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the immunoglobulin superfamily. L1/neurofascin/NgCAM family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLB7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017959.1">
        <property type="protein sequence ID" value="ENSGAGP00000015729.1"/>
        <property type="gene ID" value="ENSGAGG00000006819.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030424">
        <property type="term" value="C:axon"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098632">
        <property type="term" value="F:cell-cell adhesion mediator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007411">
        <property type="term" value="P:axon guidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007420">
        <property type="term" value="P:brain development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="CDD" id="cd00096">
        <property type="entry name" value="Ig"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05845">
        <property type="entry name" value="IgI_2_L1-CAM_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000057">
        <property type="entry name" value="neural cell adhesion molecule L1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000658">
        <property type="entry name" value="Neural cell adhesion molecule L1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000367">
        <property type="entry name" value="Neural cell adhesion molecule L1-like protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000005">
        <property type="entry name" value="Neuronal cell adhesion molecule"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000038">
        <property type="entry name" value="Neuronal cell adhesion molecule"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000347">
        <property type="entry name" value="Neuronal cell adhesion molecule"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013098">
        <property type="entry name" value="Ig_I-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013151">
        <property type="entry name" value="Immunoglobulin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026966">
        <property type="entry name" value="Neurofascin/L1/NrCAM_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR44170:SF44">
        <property type="entry name" value="L1 CELL ADHESION MOLECULE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR44170">
        <property type="entry name" value="PROTEIN SIDEKICK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13882">
        <property type="entry name" value="Bravo_FIGEY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07679">
        <property type="entry name" value="I-set"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00047">
        <property type="entry name" value="ig"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13927">
        <property type="entry name" value="Ig_3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="5"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="10" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="12" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="9 14" id="KW-0472">Membrane</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <keyword evidence="4 14" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 14" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="865"/>
            <end position="886"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="15">
        <location>
            <begin position="54"/>
            <end position="141"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="15">
        <location>
            <begin position="155"/>
            <end position="243"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="15">
        <location>
            <begin position="248"/>
            <end position="330"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="15">
        <location>
            <begin position="335"/>
            <end position="419"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="15">
        <location>
            <begin position="428"/>
            <end position="518"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="16">
        <location>
            <begin position="518"/>
            <end position="613"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="16">
        <location>
            <begin position="615"/>
            <end position="711"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="16">
        <location>
            <begin position="736"/>
            <end position="831"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="598"/>
            <end position="626"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="810"/>
            <end position="852"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="896"/>
            <end position="935"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="965"/>
            <end position="995"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="13">
        <location>
            <begin position="896"/>
            <end position="915"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="13">
        <location>
            <begin position="981"/>
            <end position="995"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008588"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015729.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="995" mass="109466" checksum="A832183EA3B481D5" modified="2019-05-08" version="1">EAGVTVVEGSGTLYVNNSNAGRHQGVYRCYASNDLGTAMGPEIRVIAESTPQWPKEKVLPLEVEEGESVILPCNPPASAMPPKIYWLNSRIVHIAQDERVSMGQDGYLYFANVRRSDSHPDYICHAHYLGPRTIIQKEPIELRVRPTNSVKFRQPRLMVPPGESSSHVALRGSSLALECIAEGFPTPTVEWIRLNGRLPAERVALENSQKTLRIRDVVEEDDGEYQCLARNTQGTARHTYTVTVEAAPYWAQQPHSRVYGPGENVRLDCSVYGKPNPLVSWRINGLAPDPTRTLRSGALILSQVQPNDSLVTQCEAVNHHGRLLANAYIYVIQLPARILTPDKQRYAVVENQTASLDCQTFGAPMPSVEWFTGRMEPALQDERAFVHTNGSLRLPRAQRQDTGTYVCLAQNDQTNVTISAWLEVKGATRIELGPQSAVVKKSSNVTFQCLVRFDETLQERGVEWWRDGRPIRESDDSDKYTIEPGWLRISGVDYTDQGVYSCRAHTELDAVEESAELRVVELQVSEPQERWVKLSWTPGDDHNSPIEKFVVEAEDGLRDPGTWRDVSTVPGHQPWAQLHLVPYGDFRFRVRAANRFGRGEPSAASAPVRTSPAAPERYPEGVAGEGNETNNMLITWKPLNRSDWNAPDLSYRVQWRRLGKGGGWSEETVSGPPLLVTRTPTFVPYEIKVQAVNPIGKGPEPPTVVGYSGEDGEWGGGLSVTLPRLPPSLCPPVPSPPSSLLLERVSDTVLALEWTAPQFPNGALTEYLLQYQQGESRGRWAGGHKGWNRGSYGWGGLAIGQVQDGYWLDPTPSGEQCSRRGKLRHTQASSPRVLGPGAGSLGNAARRSAPPVLTTDQRSFATEGWFIGFICAVVLLLLLLLILCFVKRSKGGKYSVKDKEDIQVDSEARPMKDETFGEYSDNEEKPFGSSQPSLNGALPALGSEDSLADYGGSVDVQFNEDGSFIGQYSGHTGKEPAGNNSSGATSPTTATAALE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLB8</accession>
    <name>A0A452HLB8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">C2 DOCK-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the DOCK family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLB8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017960.1">
        <property type="protein sequence ID" value="ENSGAGP00000015730.1"/>
        <property type="gene ID" value="ENSGAGG00000011372.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031267">
        <property type="term" value="F:small GTPase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007520">
        <property type="term" value="P:myoblast fusion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007264">
        <property type="term" value="P:small GTPase-mediated signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd08694">
        <property type="entry name" value="C2_Dock-A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000044">
        <property type="entry name" value="dedicator of cytokinesis protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1270.350:FF:000001">
        <property type="entry name" value="dedicator of cytokinesis protein 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.350">
        <property type="entry name" value="Dedicator of cytokinesis N-terminal subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027007">
        <property type="entry name" value="C2_DOCK-type_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026791">
        <property type="entry name" value="DOCK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047026">
        <property type="entry name" value="DOCK1_C2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032376">
        <property type="entry name" value="DOCK_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042455">
        <property type="entry name" value="DOCK_N_sub1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056372">
        <property type="entry name" value="TPR_DOCK"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45653">
        <property type="entry name" value="DEDICATOR OF CYTOKINESIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45653:SF1">
        <property type="entry name" value="DEDICATOR OF CYTOKINESIS PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14429">
        <property type="entry name" value="DOCK-C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16172">
        <property type="entry name" value="DOCK_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23554">
        <property type="entry name" value="TPR_DOCK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00326">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51650">
        <property type="entry name" value="C2_DOCK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0728">SH3 domain</keyword>
    <feature type="domain" description="C2 DOCK-type" evidence="6">
        <location>
            <begin position="418"/>
            <end position="602"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00983"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS51650"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015730.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="987" mass="113558" checksum="31DAA562F78CCBC0" modified="2019-05-08" version="1">NPCWAPLNYFLFLAFYNYDARGADELSLQIGDTVHILETYEDKLFFPPAKGIFPASYIQLKEAIVEGKGQHETVIPSELPLIQEVTTTLREWSIIWRQLYVQDNRDMFHSVRHMIYDLIEWRSQILSGTLPQDELKELKKKVTAKIDYGNRILDLDLVVRDEDGNILDPEQTSTISLFRAHEIASKQVEERLLEEKSQKQNIDINRQAKFAATPSFALFVNLKNVVCKIGEDAEVLMSLYDPLESKFISENYLVRWSSSGLPKDIDRLHNLRAVFTDLGSKDLKREKISFVCQIVRVGRMELRDNNTRKLTSGLRRPFGVAVMDVTDIINGKVDDEDKQHFIPFQPVAGENDFLQTVINKVIAAKEVNHKGQGLWVTLKLLPGDIHQIRKEFPHLVDRSTAVARKMGFPEIIMPGDVRNDIYVTLVQGDFDKGSKTTAKNVEVTVSVYDEDGKRLENVIFPGAGDDAISEYKSVIYYQVKQPRWFETVKVAIPIEDVNRSHLRFTFRHRSSQDSKDKSEKIFALAFVKLMRYDGTTLRDGEHDLIVYKAEAKKLEDASTYLSLPSTKIELEEKGHATTGKSMQNLGSCTISKDSFQISTLVCSTKLTQNVDLLGLLKWRSNTNLLQQNLRQLMKVDGGEVVKFLQDTLDALFNIMMENSESETFDTLVFDALVFIIGLIADRKFQHFNPVLETYIKKHFSATLAYTKLTKVLRNYVDNAEKPGVTDQLFKAMKALEYIFKFIVRSRILFNQLYENKGEADFMESLLQLFKSINEMMSSVSDQTVMVKGAALKYLPTIVNDVKLVFDPKELSKLFTDFILNVPTGRLTIQKLYCLIEIVHSDLFTQHDCREILLPMMTDQLKYHLERQEDLEACCQLLSNILEVLYRKDVGPTQRHVQIIMEKLLRTVNRTVISMGRDSELIGNFVASMTAILRQMEDYHYAHLIKTFGKMRSDVVDFLMETFIMFKNLIGKNVYPFDWVIMNMMQNK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLB9</accession>
    <name>A0A452HLB9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="6">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the zygin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLB9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017985.1">
        <property type="protein sequence ID" value="ENSGAGP00000015754.1"/>
        <property type="gene ID" value="ENSGAGG00000011856.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030424">
        <property type="term" value="C:axon"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011680">
        <property type="entry name" value="FEZ"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12394:SF4">
        <property type="entry name" value="FASCICULATION AND ELONGATION PROTEIN ZETA-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12394">
        <property type="entry name" value="ZYGIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07763">
        <property type="entry name" value="FEZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 4" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="132"/>
            <end position="160"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="175"/>
            <end position="198"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="4">
        <location>
            <begin position="229"/>
            <end position="294"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="136"/>
            <end position="152"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="5">
        <location>
            <begin position="180"/>
            <end position="194"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006788"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015754.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="370" mass="42975" checksum="189ACA4A1ED9A7BC" modified="2019-05-08" version="1">MEAPLVSLDEEFEDLRPCYVDDREEKPHCFYGSSPQHLEDPSLSELENFSSEIISFKSMEDLVNEFDEKLNVCFRNYNAKTENLAPVKNHFQIQEEEEHLQDEEVWDALTDNYISSVTEDWRDPDIEVLNGNISDSEIHEKEEEELHEKSENDSGINEEPLLTADQVIEEIEEMMQNSPDPEEEEENLEEDDGGETSSQADSVLLQEMQLLTQTFNNNWSYEGLEQMSVSELAELLDHVESAIRDYSEELVQQLARRDELEFEKEVKNSFITVLMEVQNKQKEHRELMKRRRKEKGLSLQSSRIERGTQMPLKRFSMEGISNILQTGIRQTFGNSATDKQYLNTVIPYEKKGSPPSVEDLQMLTNMLCPT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLC1</accession>
    <name>A0A452HLC1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Dedicator of cytokinesis 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the DOCK family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLC1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017950.1">
        <property type="protein sequence ID" value="ENSGAGP00000015721.1"/>
        <property type="gene ID" value="ENSGAGG00000011372.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031267">
        <property type="term" value="F:small GTPase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007520">
        <property type="term" value="P:myoblast fusion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007264">
        <property type="term" value="P:small GTPase-mediated signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd08694">
        <property type="entry name" value="C2_Dock-A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.350">
        <property type="entry name" value="Dedicator of cytokinesis N-terminal subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027007">
        <property type="entry name" value="C2_DOCK-type_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026791">
        <property type="entry name" value="DOCK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047026">
        <property type="entry name" value="DOCK1_C2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032376">
        <property type="entry name" value="DOCK_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042455">
        <property type="entry name" value="DOCK_N_sub1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056372">
        <property type="entry name" value="TPR_DOCK"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45653">
        <property type="entry name" value="DEDICATOR OF CYTOKINESIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45653:SF1">
        <property type="entry name" value="DEDICATOR OF CYTOKINESIS PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14429">
        <property type="entry name" value="DOCK-C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16172">
        <property type="entry name" value="DOCK_N"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23554">
        <property type="entry name" value="TPR_DOCK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00326">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51650">
        <property type="entry name" value="C2_DOCK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 5" id="KW-0728">SH3 domain</keyword>
    <feature type="domain" description="SH3" evidence="8">
        <location>
            <begin position="42"/>
            <end position="98"/>
        </location>
    </feature>
    <feature type="domain" description="C2 DOCK-type" evidence="9">
        <location>
            <begin position="438"/>
            <end position="622"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1"/>
            <end position="30"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00983"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51650"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015721.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="12">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1062" mass="121721" checksum="53E07B986AAA267E" modified="2025-10-08" version="2">MTPPERSPRIPSRFGGCGRGSPAPDGRRASPCAMTRWVPTKREEKYGVAFYNYDARGADELSLQIGDTVHILETYEDKLFFPPAKGIFPASYIQLKEAIVEGKGQHETVIPSELPLIQEVTTTLREWSIIWRQLYVQDNRDMFHSVRHMIYDLIEWRSQILSGTLPQDELKELKKKVTAKIDYGNRILDLDLVVRDEDGNILDPEQTSTISLFRAHEIASKQVEERLLEEKSQKQNIDINRQAKFAATPSFALFVNLKNVVCKIGEDAEVLMSLYDPLESKFIRLVSSVLKPEMNRILIIKDLKREKISFVCQIVRVGRMELRDNNTRKLTSGLRRPFGVAVMDVTDIINGKVDDEDKQHFIPFQPVAGENDFLQTVINKVIAAKEVNHKGQGLWVTLKLLPGDIHQIRKEFPHLVDRSTAVARKMGFPEIIMPGDVRNDIYVTLVQGDFDKGSKTTAKNVEVTVSVYDEDGKRLENVIFPGAGDDAISEYKSVIYYQVKQPRWFETVKVAIPIEDVNRSHLRFTFRHRSSQDSKDKSEKIFALAFVKLMRYDGTTLRDGEHDLIVYKAEAKKLEDASTYLSLPSTKIELEEKGHATTGKSMQNLGSCTISKDSFQISTLVCSTKLTQNVDLLGLLKWRSNTNLLQQNLRQLMKVDGGEVVKFLQDTLDALFNIMMENSESETFDTLVFDALVFIIGLIADRKFQHFNPVLETYIKKHFSATLAYTKLTKVLRNYVDNAEKPGVTDQLFKAMKALEYIFKFIVRSRILFNQLYENKGEADFMESLLQLFKSINEMMSSVSDQTVMVKGAALKYLPTIVNDVKLVFDPKELSKLFTDFILNVPTGRLTIQKLYCLIEIVHSDLFTQHDCREILLPMMTDQLKYHLERQEDLEACCQLLSNILEVLYRKDVGPTQRHVQIIMEKLLRTVNRTVISMGRDSELIGNFVASMTAILRQMEDYHYAHLIKTFGKMRSDVVDFLMETFIMFKNLIGKNVYPFDWVIMNMMQNNGLIRTQLLTLVPELGQLRLAAFLEQTLKPSSLIIMEQRSILVPTTLEPMQLQKIS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLC3</accession>
    <name>A0A452HLC3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">C2 DOCK-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the DOCK family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLC3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017965.1">
        <property type="protein sequence ID" value="ENSGAGP00000015735.1"/>
        <property type="gene ID" value="ENSGAGG00000011372.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031267">
        <property type="term" value="F:small GTPase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007520">
        <property type="term" value="P:myoblast fusion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007264">
        <property type="term" value="P:small GTPase-mediated signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd08694">
        <property type="entry name" value="C2_Dock-A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.350">
        <property type="entry name" value="Dedicator of cytokinesis N-terminal subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027007">
        <property type="entry name" value="C2_DOCK-type_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026791">
        <property type="entry name" value="DOCK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047026">
        <property type="entry name" value="DOCK1_C2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032376">
        <property type="entry name" value="DOCK_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042455">
        <property type="entry name" value="DOCK_N_sub1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056372">
        <property type="entry name" value="TPR_DOCK"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45653">
        <property type="entry name" value="DEDICATOR OF CYTOKINESIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45653:SF1">
        <property type="entry name" value="DEDICATOR OF CYTOKINESIS PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14429">
        <property type="entry name" value="DOCK-C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16172">
        <property type="entry name" value="DOCK_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23554">
        <property type="entry name" value="TPR_DOCK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51650">
        <property type="entry name" value="C2_DOCK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="C2 DOCK-type" evidence="4">
        <location>
            <begin position="364"/>
            <end position="548"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00983"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS51650"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015735.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="772" mass="88630" checksum="D57B23B9A85DB200" modified="2025-10-08" version="2">MDYINHFTILPKFVFANFRQHETVIPSELPLIQEVTTTLREWSIIWRQLYVVSKIHMFHSVRHMIYDLIEWRSQILSGTLPQDELKELKKKVTAKIDYGNRILDLDLVVRDEDGNILDPEQTSTISLFRAHEIASKQVEERLLEEKVRYTIPNSKFAATPSFALFVNLKNVVCKIGEDAEVLMSLYDPLESKFISENYLVRWSSSGLPKDIDRLHNLRAVFTDLGSKDLKREKISFVCQIVRVGRMELRDNNTRKLTSGLRRPFGVAVMDVTDIINGKVDDEDKQHFIPFQPVAGENDFLQTVINKVIAAKEVNHKGQGLWVTLKLLPGDIHQIRKEFPHLVDRSTAVARKMGFPEIIMPGDVRNDIYVTLVQGDFDKGSKTTAKNVEVTVSVYDEDGKRLENVIFPGAGDDAISEYKSVIYYQVKQPRWFETVKVAIPIEDVNRSHLRFTFRHRSSQDSKDKSEKIFALAFVKLMRYDGTTLRDGEHDLIVYKAEAKKLEDASTYLSLPSTKIELEEKGHATTGKSMQNLGSCTISKDSFQISTLVCSTKLTQNVDLLGLLKWRSNTNLLQQNLRQLMKVDGGEVVKFLQDTLDALFNIMMENSESETFDTLVFDALVFIIGLIADRKFQHFNPVLETYIKKHFSATLAYTKLTKVLRNYVDNAEKPGVTDQLFKAMKALEYIFKFIVRSRILFNQDPLALEECSADVSALQRGPCDLVLYILIYKQYVEVHNLSQKSVITNVNKYKKNKNKKTPELVQTKKPTNITQKLY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLC5</accession>
    <name>A0A452HLC5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">C2 DOCK-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the DOCK family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLC5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017969.1">
        <property type="protein sequence ID" value="ENSGAGP00000015739.1"/>
        <property type="gene ID" value="ENSGAGG00000011372.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031267">
        <property type="term" value="F:small GTPase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007520">
        <property type="term" value="P:myoblast fusion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007264">
        <property type="term" value="P:small GTPase-mediated signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd08694">
        <property type="entry name" value="C2_Dock-A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000044">
        <property type="entry name" value="dedicator of cytokinesis protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1270.350:FF:000001">
        <property type="entry name" value="dedicator of cytokinesis protein 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.350">
        <property type="entry name" value="Dedicator of cytokinesis N-terminal subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027007">
        <property type="entry name" value="C2_DOCK-type_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026791">
        <property type="entry name" value="DOCK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047026">
        <property type="entry name" value="DOCK1_C2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032376">
        <property type="entry name" value="DOCK_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042455">
        <property type="entry name" value="DOCK_N_sub1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056372">
        <property type="entry name" value="TPR_DOCK"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45653">
        <property type="entry name" value="DEDICATOR OF CYTOKINESIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45653:SF1">
        <property type="entry name" value="DEDICATOR OF CYTOKINESIS PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14429">
        <property type="entry name" value="DOCK-C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16172">
        <property type="entry name" value="DOCK_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23554">
        <property type="entry name" value="TPR_DOCK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51650">
        <property type="entry name" value="C2_DOCK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="C2 DOCK-type" evidence="4">
        <location>
            <begin position="375"/>
            <end position="559"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00983"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS51650"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015739.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="708" mass="80527" checksum="A7711F5EA6F30F9F" modified="2019-05-08" version="1">RPVGAAGSGGQYVPQPAPLPAAPIGLEQRTAASRIIPSELPLIQEVTTTLREWSIIWRQLYVVSKIHMFHSVRHMIYDLIEWRSQILSGTLPQDELKELKKKVTAKIDYGNRILDLDLVVRDEDGNILDPEQTSTISLFRAHEIASKQVEERLLEEKVRYTIPNSKFAATPSFALFVNLKNVVCKIGEDAEVLMSLYDPLESKFISENYLVRWSSSGLPKDIDRLHNLRAVFTDLGSKDLKREKISFVCQIVRVGRMELRDNNTRKLTSGLRRPFGVAVMDVTDIINGKVDDEDKQHFIPFQPVAGENDFLQTVINKVIAAKEVNHKGQGLWVTLKLLPGDIHQIRKEFPHLVDRSTAVARKMGFPEIIMPGDVRNDIYVTLVQGDFDKGSKTTAKNVEVTVSVYDEDGKRLENVIFPGAGDDAISEYKSVIYYQVKQPRWFETVKVAIPIEDVNRSHLRFTFRHRSSQDSKDKSEKIFALAFVKLMRYDGTTLRDGEHDLIVYKAEAKKLEDASTYLSLPSTKIELEEKGHATTGKSMQNLGSCTISKDSFQISTLVCSTKLTQNVDLLGLLKWRSNTNLLQQNLRQLMKVDGGEVVKFLQDTLDALFNIMMENSESETFDTLVFDALVFIIGLIADRKFQHFNPVLETYIKKHFSATLAYTKLTKVLRNYVDNAEKPGVTDQLFKAMKALEYIFKFIVRSRILFNQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLC8</accession>
    <name>A0A452HLC8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Homeobox domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="5">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the TALE/MEIS homeobox family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLC8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017976.1">
        <property type="protein sequence ID" value="ENSGAGP00000015745.1"/>
        <property type="gene ID" value="ENSGAGG00000011836.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006355">
        <property type="term" value="P:regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00086">
        <property type="entry name" value="homeodomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.60:FF:000004">
        <property type="entry name" value="Meis2 homeobox isoform 2c"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.60">
        <property type="entry name" value="Homeodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001356">
        <property type="entry name" value="HD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009057">
        <property type="entry name" value="Homeodomain-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008422">
        <property type="entry name" value="KN_HD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032453">
        <property type="entry name" value="PKNOX/Meis_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050224">
        <property type="entry name" value="TALE_homeobox"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11850">
        <property type="entry name" value="HOMEOBOX PROTEIN TRANSCRIPTION FACTORS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05920">
        <property type="entry name" value="Homeobox_KN"/>
        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
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        <property type="entry name" value="HOX"/>
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    </dbReference>
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        <property type="entry name" value="Homeodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50071">
        <property type="entry name" value="HOMEOBOX_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 5" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3 5" id="KW-0371">Homeobox</keyword>
    <keyword evidence="4 5" id="KW-0539">Nucleus</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Homeobox" evidence="7">
        <location>
            <begin position="172"/>
            <end position="235"/>
        </location>
    </feature>
    <feature type="DNA-binding region" description="Homeobox" evidence="5">
        <location>
            <begin position="174"/>
            <end position="236"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="237"/>
            <end position="257"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="309"/>
            <end position="358"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="6">
        <location>
            <begin position="247"/>
            <end position="257"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="6">
        <location>
            <begin position="315"/>
            <end position="340"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00009661"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023155"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00108"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50071"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015745.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="358" mass="39524" checksum="94F6D73B43958B14" modified="2019-05-08" version="1">SAVSCSTQGGVFSHPSCSAMVKAIQVLRIHLLELEKVNELCKDFCNRYITCLKTKMHSDNLLRNDLGGPYSPNQPSINLHPQDLLQNSPNSMTAVANNPQGIVVPASALQPGNISMTTVNSQVVSGGALYQPVTMVTSQGQVVTQAIPQGAIQIQNTQVNLDLTSLLDNEDKKSKNKRGVLPKHATNIMRSWLFQHLMHPYPTEDEKRQIAAQTNLTLLQVNNWFINARRRILQPMLDASNPDPAPKAKKIKSQHRPTQRFWPNSIAAGVLQQQGVNPTTNPDGSLIMDNLQSLSSDNATIAMQQAMMAAHDDSLDGTEDEEEDEMEEEEEEELEEDNDELQTTNVSDLGLEHSDSLE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLD0</accession>
    <name>A0A452HLD0_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="8">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLD0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017999.1">
        <property type="protein sequence ID" value="ENSGAGP00000015766.1"/>
        <property type="gene ID" value="ENSGAGG00000011846.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042995">
        <property type="term" value="C:cell projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005332">
        <property type="term" value="F:gamma-aminobutyric acid:sodium:chloride symporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000175">
        <property type="entry name" value="Na/ntran_symport"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037272">
        <property type="entry name" value="SNS_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11616:SF124">
        <property type="entry name" value="SODIUM- AND CHLORIDE-DEPENDENT GABA TRANSPORTER 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11616">
        <property type="entry name" value="SODIUM/CHLORIDE DEPENDENT TRANSPORTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00209">
        <property type="entry name" value="SNF"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="NANEUSMPORT"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50267">
        <property type="entry name" value="NA_NEUROTRAN_SYMP_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5 7" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0915">Sodium</keyword>
    <keyword evidence="3 7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 7" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="2" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="23"/>
            <end position="48"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="68"/>
            <end position="93"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="105"/>
            <end position="131"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="164"/>
            <end position="196"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="208"/>
            <end position="233"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="239"/>
            <end position="261"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="281"/>
            <end position="303"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="323"/>
            <end position="344"/>
        </location>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="79"/>
        </location>
        <ligand>
            <name>Na(+)</name>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="111"/>
        </location>
        <ligand>
            <name>Na(+)</name>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
            <label>1</label>
        </ligand>
    </feature>
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        <location>
            <position position="176"/>
        </location>
        <ligand>
            <name>Na(+)</name>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
            <label>1</label>
        </ligand>
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        <location>
            <position position="179"/>
        </location>
        <ligand>
            <name>Na(+)</name>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
            <label>1</label>
        </ligand>
    </feature>
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        <location>
            <position position="180"/>
        </location>
        <ligand>
            <name>Na(+)</name>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
            <label>1</label>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PIRSR" id="PIRSR600175-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015766.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="402" mass="44883" checksum="EA1C203FBC6A2700" modified="2019-05-08" version="1">LLPSGSVALLDTLQPSTPRPLRVVYVTATFPYVMLLILLVRGVTLPGASEGIKFYLYPDLSRLSDPQVWVDAGTQIFFSYAICLGCLTALGSYNNYNNNCYRDCIMLCCLNSGTSFVAGFAIFSVLGFMAYEQGVPIAEVAESGPGLAFIAYPKAVTMMPLSPLWAALFFMMLIFLGLDSQFVCVESLVTAVVDMYPKIFRRGYRRELLILALSIVSYFVGLIMLTEGGMYIFQLFDSYAASGMCLLFVAIFECICIGWIYGSNRFYDNIEDMIGYRPLSLIKWCWMVLTPGICAGIFIFFLVKYKPLKYNNVYTYPDWGYGIGWMMALSSMVCIPLWICIKLWKTEGTVLEKFRKLVIPSPDLKMRGKFGAGAYAATVNDCDAKLKGDGNISAITEKETHF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLD3</accession>
    <name>A0A452HLD3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Homeobox domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="5">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the TALE/MEIS homeobox family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLD3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017977.1">
        <property type="protein sequence ID" value="ENSGAGP00000015746.1"/>
        <property type="gene ID" value="ENSGAGG00000011836.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006355">
        <property type="term" value="P:regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00086">
        <property type="entry name" value="homeodomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.60:FF:000004">
        <property type="entry name" value="Meis2 homeobox isoform 2c"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.60">
        <property type="entry name" value="Homeodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001356">
        <property type="entry name" value="HD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009057">
        <property type="entry name" value="Homeodomain-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008422">
        <property type="entry name" value="KN_HD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032453">
        <property type="entry name" value="PKNOX/Meis_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050224">
        <property type="entry name" value="TALE_homeobox"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11850">
        <property type="entry name" value="HOMEOBOX PROTEIN TRANSCRIPTION FACTORS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05920">
        <property type="entry name" value="Homeobox_KN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16493">
        <property type="entry name" value="Meis_PKNOX_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00389">
        <property type="entry name" value="HOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46689">
        <property type="entry name" value="Homeodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50071">
        <property type="entry name" value="HOMEOBOX_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 5" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3 5" id="KW-0371">Homeobox</keyword>
    <keyword evidence="4 5" id="KW-0539">Nucleus</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Homeobox" evidence="7">
        <location>
            <begin position="176"/>
            <end position="239"/>
        </location>
    </feature>
    <feature type="DNA-binding region" description="Homeobox" evidence="5">
        <location>
            <begin position="178"/>
            <end position="240"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="241"/>
            <end position="261"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="313"/>
            <end position="362"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="6">
        <location>
            <begin position="251"/>
            <end position="261"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="6">
        <location>
            <begin position="319"/>
            <end position="344"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00009661"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023155"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00108"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50071"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015746.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="362" mass="40036" checksum="67F385ADC9B909D0" modified="2019-05-08" version="1">KGLPPTIAPAASSGQLMARLLKCMVKAIQVLRIHLLELEKVNELCKDFCNRYITCLKTKMHSDNLLRNDLGGPYSPNQPSINLHPQDLLQNSPNSMTAVANNPQGIVVPASALQPGNISMTTVNSQVVSGGALYQPVTMVTSQGQVVTQAIPQGAIQIQNTQVNLDLTSLLDNEDKKSKNKRGVLPKHATNIMRSWLFQHLMHPYPTEDEKRQIAAQTNLTLLQVNNWFINARRRILQPMLDASNPDPAPKAKKIKSQHRPTQRFWPNSIAAGVLQQQGVNPTTNPDGSLIMDNLQSLSSDNATIAMQQAMMAAHDDSLDGTEDEEEDEMEEEEEEELEEDNDELQTTNVSDLGLEHSDSLE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLD8</accession>
    <name>A0A452HLD8_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="6">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the zygin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLD8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017986.1">
        <property type="protein sequence ID" value="ENSGAGP00000015755.1"/>
        <property type="gene ID" value="ENSGAGG00000011856.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030424">
        <property type="term" value="C:axon"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011680">
        <property type="entry name" value="FEZ"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12394:SF4">
        <property type="entry name" value="FASCICULATION AND ELONGATION PROTEIN ZETA-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12394">
        <property type="entry name" value="ZYGIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07763">
        <property type="entry name" value="FEZ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 4" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="145"/>
            <end position="169"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="4">
        <location>
            <begin position="200"/>
            <end position="265"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="5">
        <location>
            <begin position="151"/>
            <end position="165"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006788"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015755.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="363" mass="42175" checksum="ACF2782563597F34" modified="2019-05-08" version="1">MEAPLVSLDEEFEDLRPCYVDDREEKPHCFYGSSPQHLEDPSLSELENFSSEIISFKSMEDLVNEFDEKLNVCFRNYNAKTENLAPVKNHFQIQEEEEHLQDEEVWDALTDNYISSVTEDWRDPDIEVLNGNISDSEVIEEIEEMMQNSPDPEEEEENLEEDDGGETSSQADSVLLQEMQLLTQTFNNNWSYEGLEQMSVSELAELLDHVESAIRDYSEELVQQLARRDELEFEKEVKNSFITVLMEVQNKQKEHRELMKRRRKEKGLSLQSSRIERGTQMPLKRFSMEGISNILQTGIRQTFGNSATDKQYLNTVIPYEKKGSPPSVEDLQMLTNILFAMKEDNEKVPTLLTDYILKVLCPT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLD9</accession>
    <name>A0A452HLD9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">receptor protein-tyrosine kinase</fullName>
            <ecNumber evidence="2">2.7.10.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="26 27">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="27" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="26" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="26" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>L-tyrosyl-[protein] + ATP = O-phospho-L-tyrosyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10596"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.10.1"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="2.7.10.1" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452HLD9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018009.1">
        <property type="protein sequence ID" value="ENSGAGP00000015776.1"/>
        <property type="gene ID" value="ENSGAGG00000010943.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043235">
        <property type="term" value="C:receptor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004714">
        <property type="term" value="F:transmembrane receptor protein tyrosine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007169">
        <property type="term" value="P:cell surface receptor protein tyrosine kinase signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006909">
        <property type="term" value="P:phagocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:001050">
        <property type="entry name" value="MER proto-oncogene, tyrosine kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000834">
        <property type="entry name" value="Proto-oncogene tyrosine-protein kinase MER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000089">
        <property type="entry name" value="Tyrosine-protein kinase receptor TYRO3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000111">
        <property type="entry name" value="Tyrosine-protein kinase receptor TYRO3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050122">
        <property type="entry name" value="RTK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001245">
        <property type="entry name" value="Ser-Thr/Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008266">
        <property type="entry name" value="Tyr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020635">
        <property type="entry name" value="Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416:SF257">
        <property type="entry name" value="TYROSINE-PROTEIN KINASE MER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416">
        <property type="entry name" value="TYROSINE-PROTEIN KINASE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07714">
        <property type="entry name" value="PK_Tyr_Ser-Thr"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00109">
        <property type="entry name" value="TYRKINASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00219">
        <property type="entry name" value="TyrKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00109">
        <property type="entry name" value="PROTEIN_KINASE_TYR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="9 19 21" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="13" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="15" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="16" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="8" id="KW-0418">Kinase</keyword>
    <keyword evidence="20" id="KW-0460">Magnesium</keyword>
    <keyword evidence="11 22" id="KW-0472">Membrane</keyword>
    <keyword evidence="20" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7 19 21" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="14" id="KW-0675">Receptor</keyword>
    <keyword evidence="27" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0808">Transferase</keyword>
    <keyword evidence="5 22" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="10 22" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="12" id="KW-0829">Tyrosine-protein kinase</keyword>
    <feature type="transmembrane region" description="Helical" evidence="22">
        <location>
            <begin position="365"/>
            <end position="388"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="24">
        <location>
            <begin position="73"/>
            <end position="159"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="25">
        <location>
            <begin position="154"/>
            <end position="249"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="25">
        <location>
            <begin position="254"/>
            <end position="346"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="23">
        <location>
            <begin position="450"/>
            <end position="721"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="18">
        <location>
            <position position="586"/>
        </location>
    </feature>
    <feature type="binding site" evidence="21">
        <location>
            <position position="482"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="19">
        <location>
            <position position="590"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="20">
        <location>
            <position position="591"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="20">
        <location>
            <position position="604"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00011902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023137"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00051243"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="PIRSR" id="PIRSR000615-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="PIRSR" id="PIRSR000615-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="PIRSR" id="PIRSR000615-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015776.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="856" mass="95806" checksum="E6343A78E462C711" modified="2019-05-08" version="1">MPFLWVILRSSVKVFQHFLSFVPVINLKVNPFLSPGSITNAQRSDNGSYSCKLKIHNEEIESDPIFVQMEGLPHFLRQPQSLNITRNRPFNLTCEAMGPPEPVQIYWFRNSSRVNDQPYISPSVLTLPGECRPGAVSACRADQRWLSHVGTPAAPADVRVLNRTAHSLVVSWVPGFDGYSPLHSCRLQVREAAALNKGSFLIWNTSVPPHRYHVQQLQPLAEYSIRVSCRNEVGWSSFSPWTLASTTEGAPSTPPLNVTVFFNESSSSLEVRWAKPHLGRKHGELQGYRLLHTWQNSKITVTQQNATVTTVAIVVTNATYAVRVAAVTKGGIGPFSDPVVVFIPGDGFMASAPSSTPAPGNNHSFAIVLGFILGIVAIGLILYFSVIVRKRWMETKFGTAFHSDDSEHVVSYTAKKSYSRRAVELTLASLGVREELQQKLQDVVIDRDSLALGKILGEGEFGSVMEGHLSQPSGTVQKVAVKTMKLEDFSQREIEEFLSEAACMKDFDHPNVIRLLGVCIELSSRQVPKPMVVLPFMKYGDLHSFLLRSRFETGTRHVPLQTLLKFMVDIALGMEYLSTRQFLHRDLAARNCMLRDDMTVCVADFGLSKKIYSGDYYRQGRIAKMPVKWIAIESLADRVYTTKSDVWAFGVTMWEIATRGMTPYPGVQNHEIYDYLLHGHRLKQPEDCLDELYEVMYSCWRADPHERPAFPELKLHLEKLLESLPKVSGSEDVIYINTSLPEKSEELTEESEFPQTDMAFNRNYVTVSSSPKPDTTVVTAEVHESDQREERYILKGETDEQTAGSLTLEQGHAPLLQGTITRNGATWGPASMLLGNSLADELLYADDCSEDSEVLL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLG3</accession>
    <name>A0A452HLG3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Aminopeptidase</fullName>
            <ecNumber evidence="20">3.4.11.-</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="18 20">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
        <text evidence="18 20">Binds 1 zinc ion per subunit.</text>
    </comment>
    <comment type="subunit">
        <text evidence="3">Homodimer.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type II membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 20">Belongs to the peptidase M1 family.</text>
    </comment>
    <dbReference type="EC" id="3.4.11.-" evidence="20"/>
    <dbReference type="AlphaFoldDB" id="A0A452HLG3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018011.1">
        <property type="protein sequence ID" value="ENSGAGP00000015778.1"/>
        <property type="gene ID" value="ENSGAGG00000011694.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070006">
        <property type="term" value="F:metalloaminopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042277">
        <property type="term" value="F:peptide binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043171">
        <property type="term" value="P:peptide catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd09601">
        <property type="entry name" value="M1_APN-Q_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1910:FF:000005">
        <property type="entry name" value="Aminopeptidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.50.20:FF:000012">
        <property type="entry name" value="Aminopeptidase N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.390.10:FF:000016">
        <property type="entry name" value="Glutamyl aminopeptidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1730:FF:000001">
        <property type="entry name" value="Leucyl-cystinyl aminopeptidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.50.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1910">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.390.10">
        <property type="entry name" value="Neutral Protease Domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1730">
        <property type="entry name" value="tricorn interacting facor f3 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045357">
        <property type="entry name" value="Aminopeptidase_N-like_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042097">
        <property type="entry name" value="Aminopeptidase_N-like_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024571">
        <property type="entry name" value="ERAP1-like_C_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034016">
        <property type="entry name" value="M1_APN-typ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001930">
        <property type="entry name" value="Peptidase_M1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050344">
        <property type="entry name" value="Peptidase_M1_aminopeptidases"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014782">
        <property type="entry name" value="Peptidase_M1_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027268">
        <property type="entry name" value="Peptidase_M4/M1_CTD_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11533:SF172">
        <property type="entry name" value="AMINOPEPTIDASE N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11533">
        <property type="entry name" value="PROTEASE M1 ZINC METALLOPROTEASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11838">
        <property type="entry name" value="ERAP1_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01433">
        <property type="entry name" value="Peptidase_M1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17900">
        <property type="entry name" value="Peptidase_M1_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00756">
        <property type="entry name" value="ALADIPTASE"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF63737">
        <property type="entry name" value="Leukotriene A4 hydrolase N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55486">
        <property type="entry name" value="Metalloproteases ('zincins'), catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 20" id="KW-0031">Aminopeptidase</keyword>
    <keyword evidence="5" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="15" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="16" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9 20" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="14 20" id="KW-0472">Membrane</keyword>
    <keyword evidence="8 18 20" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13 20" id="KW-0482">Metalloprotease</keyword>
    <keyword evidence="6 20" id="KW-0645">Protease</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="11" id="KW-0735">Signal-anchor</keyword>
    <keyword evidence="7 20" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="12 20" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="10 18 20" id="KW-0862">Zinc</keyword>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="12"/>
            <end position="33"/>
        </location>
    </feature>
    <feature type="domain" description="Aminopeptidase N-like N-terminal" evidence="23">
        <location>
            <begin position="79"/>
            <end position="271"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase M1 membrane alanine aminopeptidase" evidence="21">
        <location>
            <begin position="307"/>
            <end position="534"/>
        </location>
    </feature>
    <feature type="domain" description="ERAP1-like C-terminal" evidence="22">
        <location>
            <begin position="612"/>
            <end position="939"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="17">
        <location>
            <position position="380"/>
        </location>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="379"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <note>catalytic</note>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="383"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <note>catalytic</note>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="402"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <note>catalytic</note>
        </ligand>
    </feature>
    <feature type="site" description="Transition state stabilizer" evidence="19">
        <location>
            <position position="468"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004401"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00011738"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022438"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023049"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="PIRSR" id="PIRSR634016-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="PIRSR" id="PIRSR634016-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="PIRSR" id="PIRSR634016-4"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="RuleBase" id="RU364040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="Pfam" id="PF01433"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="Pfam" id="PF11838"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="Pfam" id="PF17900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015778.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="961" mass="107881" checksum="CC8A2FA1F12C9C9A" modified="2019-05-08" version="1">MAAGFFISKTLGIVSIILGLGAVATIIALSVVYSQEKEKNAASFLKATPANVTNTTPLASTAAPANPWNNLRLPESLIPEHYAVTLQPFLTPDASSGLYIFKGNSTVRFSCQNATDLILIHSKQLNYTQQSGFLVSLQGVSGSSPPPITKTWLETETQYLVVQLQGQLEKGKSYELVSSFTGELADDLAGFYRSEYVEDGETRVVATTQMQAPDARKAFPCFDEPAMKANFTVTLIHSPNHLALSNMPQTIDGTTWNVTEFQTTPKMSTYLLAFIISQFEAVEAMQDNVLIRIWGRPQAIAAGQGDYALNVTGPILKFFENHYNTSYPLPKSDQVALPDFNAGAMENWGLVTYRENSLLYDPLFSSIGNKERVVTVIAHELAHQWFGNLVTLRWWNDLWLNEGFASYVEYLGADSAEPSWHIKDLIVQSEMYRVMKVDALASSHPLSFLESEINTPAQISEVFDAIAYSKGASVLRMLSEFLTEDSFKKGLQSYLHAFSYGNTVYDDLWTHLQKVVVEDKIQLPSTIKDIMDRWTLQMGFPMVTVDTKVGSISQEHFLLDTESIVERPSAFNYTWIVPVTWMRQQDGAGQIYWLSQNSANNPSFMVTGTHNWLLLNINVTGYFRVNYDQDNWNKLLTQLDTDHQAIPVINRAQIIDDAFNLARAKHISIVLALNTTTYLGKEQEYLPWSAALDNLGYFRLMFDRSEVYGPMQKYIQKQVTPVFQYFGNVTSNWTRIPDGLMEQYSEINTISTACSNGVTECEQLAAAFFNQWKQNPSNNPIPPNLRSSIYCSAIKTGSEADWDFLWQMFKNATVVAEADKLRSAMACSNKQPWILSRYLQYCLDPTKIRKQDATSTINSIASNVVGQSLAWDFIRGNWKTLFSQYGGGSFSFSSLIQAVTQRFASEFELQQLEQFKVDNADVGFGSGTRALEQALEQTKANIKWVAENKQTVLTWFETASS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLG7</accession>
    <name>A0A452HLG7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">non-specific serine/threonine protein kinase</fullName>
            <ecNumber evidence="4">2.7.11.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
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        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
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        <citation type="submission" date="2025-08" db="UniProtKB">
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        </source>
    </evidence>
    <sequence length="2413" mass="266497" checksum="0FF73082C6E4AC51" modified="2019-05-08" version="1">MDMSDPNIWTVLSNYTLPHLRNGNRLRRTQSCRTSNRKSLIGNGQSPALPRPHSPLSAHAGSSPQDSPRNFSPSASAHFSFARRTDGRRWSLASLPSSGYGTNTPSSTVSSSCSSQEKLHQLPYQPTPDELHFLSKHFCTTESIASDNRCRTTPMRPRSRSLSPGRSPACCDHEIIMMNHVYKERFPKATAQMEERLQDIITNHSPDNILPLADGVLSFTHHQIIELARDCLDKSHQGLITSRYFFELQHKLDKLLQEGQERSESGELAFIKQLVRKILIVIARPARLLECLEFDPEEFYYLLEAAEGHAKEGHVIKTDIPRYIISQLGLNKDPLEEIAQLANYDSGTAETPETDESSSNTSVTLRRKPRESDFETIKLISNGAYGAVYFVRHKETRQRFAMKKINKQNLILRNQIQQAFVERDILTFAENPFVVSMYCSFETRRHLCMVMEYVEGGDCATLMKNMGPLPVDMARMYFAETVLALEYLHNYGIVHRDLKPDNLLVTSMGHIKLTDFGLSKVGLMSMTTNLYEGHIEKDAREFLDKQVCGTPEYIAPEVILRQGYGKPVDWWAMGIILYEFLVGCVPFFGDTPEELFGQVISDEINWPEKDEAPPPDSQDLITLLLRQNPMERLGTGGAYEVKQHQFFRSLDWNSLLRQKAEFIPQLESEDDTSYFDTRSEKYHHMETEEEDDTNDEDFNLEIRQFSSCSHRFSKVFSSIDRITRNQGEEKEDTGDKTKNVTLPSVESLSWSSEYSELQQLSTSNSSDTESTRLRHSSGLLPKLAISSEGDQDETTCSEGSREEQEKAASNEEAAQDEPEVTTPASTISSSTLSVGSFSEHLDQINGRNESVDSTDNSSKPSSEPASHMARQRLESTEKKKISGKVTKSLSASALSLMIPGDVFGVSPLGSPMSPHSLSSDLSSSRDSSPSRDSSVASATPHQPILIHSSGKKYGFTIRAIRVYVGDSDIYTVHHIVWNVEEGSPAYQAGLKAGDLITHINGEPVHGLVHTEVIELLLKSGNKVSITTTPFENTSIKTGPARRNSYRSRMVRRSKKTKKKESLERRRSLFKKLAKQPSPLLHTSRSFSCLNRSLSSGESLPGSPTHSLSPRSPTPSYRSTPDFPSGTNSSQSSSPSSSGPNSPAGSGHIRPSTLHGLGPKLGGQRYRSGRRKSAGSIPLSPLARTPSPTPQPPSPQRSPSPLLGHSIGNSKIAQAFPSKMHSPPTIVRHIVRPKSAEPPRSPLLKRVQSEEKLAPSYGCDKKHLCSRKHSLEVTQEEVNRELSQREVTLQSLEENVCDVPSLTRVRPAEQGCLKRPVCRKLGRQESVDELDREKLKVKIVMKKQDLAEKQDCLQKASTIHEPSGESENPAILRLEERDKKSFQKALERSNHFETKITALETPSVGGILKDTLHKNANMRPNDCIPLDTHMPCHGPPLNELSHLSYDFKRISPPCTLQDVLPHSSDRLLNGKGESTDKNAPTKEFVKFERLDGKLANIDYLRKKMSFEEKDDSLCPVLKPKLTSNVHECLQLNTVRPINVQQDSAAVADSRAFISSAHAAQISSVSFVPLKALNGRVETGLEKSALISTESPVRKSPSEYKLEGRSVSCLKPIEGTLDIALLSGPQASKTELPSCLLPESQNTSNEVGSSMPLASQCNGVKEETSCQKEVVSPHSKSSKSNLLEPQFSQSSKSFQNAPATPVTAELETGKKISSPDAACNISVREGDQKREDSHSKQKDHATEAKLHTVHNQCLHTALASATPSTAQGTPEKTAGKESQTQKELPVKYPVTQRNYESFSVKAEGIRDSSVKKSIPDAFKTSDSKTIQRLSADFAIPLQTHRTTRVQTAGSKVESKDGKDLPKQFAKDDITPAGSSVERDMNKQEVMDSKSEMAFPKRPVQQSVESNTKSEKSLVCSSAQKDNSKDLRKKDQKHSSNIQLHIIERKLPNLITRQQFSPSCFPSVRETVNRNSVIDAHVHLGVQEQVKTGSAYMESSNNKSRQINETCSSKSKHSDKFTFAQKQCTANIKEEETVIQPAAGSRKDGKHANKSVQDSHSFVLISQRKLNNEYIQVEKPLGLEHGSAPSIQINSSTPETKPKSSCNGQAPAGSENSKHFLSQETAGFSETADQKQLYVNEKQNMPPKFSTVKDCLLLNKLADKDQSNQSITADKRPEGQKCTEALYVQNDSGKLEPSLFLNCDLGKVAEKAPIVSKGSPDSKGKGQVSQKQLTDANKQITAMLLSTATVQSSCHSAVTPLKQLNCPPSFPESRQEASPLSSAKVKTGSPELPDTSCKELKKQATSSTGDGKREITKSVSLLTLHSSAIPVEALHPASNLGSKPGNQEKPLSASNTVDVKGEDPIQAQPPAAKKQNVGKDFPRSVTTSDRPTALSSDKDSVRQRRGKDASRSSPHKKSSQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLG8</accession>
    <name>A0A452HLG8_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="8">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HLG8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018008.1">
        <property type="protein sequence ID" value="ENSGAGP00000015775.1"/>
        <property type="gene ID" value="ENSGAGG00000011846.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042995">
        <property type="term" value="C:cell projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005332">
        <property type="term" value="F:gamma-aminobutyric acid:sodium:chloride symporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000175">
        <property type="entry name" value="Na/ntran_symport"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037272">
        <property type="entry name" value="SNS_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11616:SF124">
        <property type="entry name" value="SODIUM- AND CHLORIDE-DEPENDENT GABA TRANSPORTER 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11616">
        <property type="entry name" value="SODIUM/CHLORIDE DEPENDENT TRANSPORTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00209">
        <property type="entry name" value="SNF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00176">
        <property type="entry name" value="NANEUSMPORT"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF161070">
        <property type="entry name" value="SNF-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50267">
        <property type="entry name" value="NA_NEUROTRAN_SYMP_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5 7" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0915">Sodium</keyword>
    <keyword evidence="3 7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 7" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="2" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="12"/>
            <end position="34"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="66"/>
            <end position="87"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="99"/>
            <end position="125"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="158"/>
            <end position="190"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="202"/>
            <end position="227"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="233"/>
            <end position="255"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="275"/>
            <end position="297"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="317"/>
            <end position="338"/>
        </location>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="73"/>
        </location>
        <ligand>
            <name>Na(+)</name>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="105"/>
        </location>
        <ligand>
            <name>Na(+)</name>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="170"/>
        </location>
        <ligand>
            <name>Na(+)</name>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="173"/>
        </location>
        <ligand>
            <name>Na(+)</name>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="174"/>
        </location>
        <ligand>
            <name>Na(+)</name>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
            <label>1</label>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PIRSR" id="PIRSR600175-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015775.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="396" mass="44460" checksum="E9F73572FBBB1EE3" modified="2019-05-08" version="1">GMILCYKTCVELQMYIVVYVTATFPYVMLLILLVRGVTLPGASEGIKFYLYPDLSRLSDPQVWVDAGTQIFFSYAICLGCLTALGSYNNYNNNCYRDCIMLCCLNSGTSFVAGFAIFSVLGFMAYEQGVPIAEVAESGPGLAFIAYPKAVTMMPLSPLWAALFFMMLIFLGLDSQFVCVESLVTAVVDMYPKIFRRGYRRELLILALSIVSYFVGLIMLTEGGMYIFQLFDSYAASGMCLLFVAIFECICIGWIYGSNRFYDNIEDMIGYRPLSLIKWCWMVLTPGICAGIFIFFLVKYKPLKYNNVYTYPDWGYGIGWMMALSSMVCIPLWICIKLWKTEGTVLEKFRKLVIPSPDLKMRGKFGAGAYAATVNDCDAKLKGDGNISAITEKETHF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HLH0</accession>
    <name>A0A452HLH0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">EGF-like calcium-binding domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HLH0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000018050.1">
        <property type="protein sequence ID" value="ENSGAGP00000015806.1"/>
        <property type="gene ID" value="ENSGAGG00000011881.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007413">
        <property type="term" value="P:axonal fasciculation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027039">
        <property type="entry name" value="Crtac1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001881">
        <property type="entry name" value="EGF-like_Ca-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018097">
        <property type="entry name" value="EGF_Ca-bd_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013517">
        <property type="entry name" value="FG-GAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028994">
        <property type="entry name" value="Integrin_alpha_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049883">
        <property type="entry name" value="NOTCH1_EGF-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011519">
        <property type="entry name" value="UnbV_ASPIC"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16026">
        <property type="entry name" value="CARTILAGE ACIDIC PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16026:SF0">
        <property type="entry name" value="CARTILAGE ACIDIC PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07645">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13517">
        <property type="entry name" value="FG-GAP_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07593">
        <property type="entry name" value="UnbV_ASPIC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00179">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69318">
        <property type="entry name" value="Integrin alpha N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01187">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="1" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0732">Signal</keyword>
    <feature type="domain" description="EGF-like calcium-binding" evidence="4">
        <location>
            <begin position="499"/>
            <end position="546"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="SMART" id="SM00179"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015806.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="585" mass="62968" checksum="86FDAEE0460C71FB" modified="2019-05-08" version="1">MALPLIPWPCPSLVPRGVALAWCCPIPWSCPSLAPRSVAPPSIPWPCPILAPRGVMLLPVSWLCPSLAPWGVAPPPIPCPSSPGLATYTDKLFKLRAGRWEDLLSDEVNRAVASRFAGRSVACVDRGSGRYAIYIANYANSNLGPHVLLEMDVAASDPARGIVVLTDVAAEAGVNKYTGGRGVAVGPIVSASASDIFCDNENGPNFLFHNQGDGSFVDVAASAGLDDPYQHGRGVALADFNRDGKVDIVYGNWNGPHRLYLQMNTSGRVHFRDIATPKFSMPSPVRTVLAADFDNNQELEVFFNNIAYRGSSANRLFRIIRREHLDPVIEELNPGDAMEPEGRGTGGAVTDFDGDGLLDLILSHGESMAQPLSVFRGTQGAGNNWLRVIPRTRFGAFARGARVVLFTRHSGAHLRIIDGGSGYLCEMEPVAHFGLGRDEASSLEVTWPDGRVVTRAVASGETNSVLEIPYPQGAEGQLPPTSLECGQGFSQHENGRCVDTDECAEFPFVCPREKPLCSNTYGGYRCHSSRRCSRGFEPSEDGTACVAQVAFFGGYPSAGSRLWAWRPPSSLLSLGLGLCLHFYAL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0A1</accession>
    <name>A0A452J0A1_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="21">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 16">Membrane</location>
            <topology evidence="1 16">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 16">Belongs to the integrin alpha chain family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0A1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038309.1">
        <property type="protein sequence ID" value="ENSGAGP00000033820.1"/>
        <property type="gene ID" value="ENSGAGG00000023825.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009897">
        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008305">
        <property type="term" value="C:integrin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001525">
        <property type="term" value="P:angiogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033627">
        <property type="term" value="P:cell adhesion mediated by integrin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098609">
        <property type="term" value="P:cell-cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007160">
        <property type="term" value="P:cell-matrix adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007229">
        <property type="term" value="P:integrin-mediated signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.130:FF:000003">
        <property type="entry name" value="Integrin alpha V"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1510:FF:000001">
        <property type="entry name" value="Integrin alpha V"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1530:FF:000002">
        <property type="entry name" value="integrin alpha-V isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1460:FF:000001">
        <property type="entry name" value="Integrin, alpha V"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.130">
        <property type="entry name" value="Integrin alpha, N-terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1460">
        <property type="entry name" value="Integrin domains. Chain A, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1510">
        <property type="entry name" value="ntegrin, alpha v. Chain A, domain 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1530">
        <property type="entry name" value="ntegrin, alpha v. Chain A, domain 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013517">
        <property type="entry name" value="FG-GAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013519">
        <property type="entry name" value="Int_alpha_beta-p"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000413">
        <property type="entry name" value="Integrin_alpha"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018184">
        <property type="entry name" value="Integrin_alpha_C_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013649">
        <property type="entry name" value="Integrin_alpha_Ig-like_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048285">
        <property type="entry name" value="Integrin_alpha_Ig-like_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048286">
        <property type="entry name" value="Integrin_alpha_Ig-like_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028994">
        <property type="entry name" value="Integrin_alpha_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032695">
        <property type="entry name" value="Integrin_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23220">
        <property type="entry name" value="INTEGRIN ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23220:SF4">
        <property type="entry name" value="INTEGRIN ALPHA-V"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01839">
        <property type="entry name" value="FG-GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08441">
        <property type="entry name" value="Integrin_A_Ig_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20805">
        <property type="entry name" value="Integrin_A_Ig_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20806">
        <property type="entry name" value="Integrin_A_Ig_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00357">
        <property type="entry name" value="Integrin_alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01185">
        <property type="entry name" value="INTEGRINA"/>
    </dbReference>
    <dbReference type="SMART" id="SM00191">
        <property type="entry name" value="Int_alpha"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69318">
        <property type="entry name" value="Integrin alpha N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69179">
        <property type="entry name" value="Integrin domains"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51470">
        <property type="entry name" value="FG_GAP"/>
        <property type="match status" value="6"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0106">Calcium</keyword>
    <keyword evidence="8 16" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="14" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10 16" id="KW-0401">Integrin</keyword>
    <keyword evidence="11" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13 16" id="KW-0675">Receptor</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5 16" id="KW-0732">Signal</keyword>
    <keyword evidence="3" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="16">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5018807530" evidence="16">
        <location>
            <begin position="21"/>
            <end position="997"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="22"/>
            <end position="87"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="98"/>
            <end position="159"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="222"/>
            <end position="276"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="277"/>
            <end position="342"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="343"/>
            <end position="397"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="401"/>
            <end position="464"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha first immunoglubulin-like" evidence="18">
        <location>
            <begin position="449"/>
            <end position="607"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha second immunoglobulin-like" evidence="19">
        <location>
            <begin position="608"/>
            <end position="736"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha third immunoglobulin-like" evidence="20">
        <location>
            <begin position="754"/>
            <end position="960"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="975"/>
            <end position="997"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="17">
        <location>
            <begin position="975"/>
            <end position="991"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023037"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00803"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="RuleBase" id="RU003762"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF08441"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="Pfam" id="PF20805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="Pfam" id="PF20806"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033820.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="997" mass="110411" checksum="0CD09BC8F6AD0651" modified="2019-05-08" version="1">MARLRLLLLLLGRWLALGGAFNLDADGPAVYSGSEGSYFGFAVDFFAPDSSSIFLLVGAPKANTTQPDIVEGGQVLKCNWNSNKNCQPVIFDSTGDRNFAPNDPLEFKSHQWFGASVRSKNDKILACAPLYHWRTEVKQEREPVGTCYLHDGSKAVEYAPCRSTIIDADGQGFCQGGFSIDFTKQLSILMGFFPAGQLISDRVAEIVSKYNSKVYSIKYDDQLATRSASAAFDDSYLGYSVAVGEFSGDGIEDFVSGVPRAARTLGMVSIYNGKNMSSLYNFTGEQMAAYFGYSVAATDINGDNYTDLFIGAPLFMERGSDGKLQEVGQVSICLQQASGGFQIIKLNGFEIFARFSSSIAPLGDLDQDGFNVAAPYGGEDKRGVVYIYNGRATGLNAVPSQILEGQWAARTMPPSFGYSLKGATDVDKNGYPDLIVGAFGVDTAVLYRARPVISVNAGLEVNPTILNPEEKICELPDTRVKVSCFNVKFCLTADGKGKLPNTLTFQVELLLDKLKQKGALRRALFLHNKQPGHSKNMTVVKGGTMRCEELQAFLRDESEFRDKLTPITVFMEYRLDYKTAKDSAGLQPILNQFTPANISRQTHILLDCGEDNICKPKLQVAVHSDQKQIYIGDDNPLTLIVDAQNQGEGAYEAELFVIIPPQADFIGVVRNNEALARLSCAFKTENQTRLVVCDLGNPMKAGTKLLAGLRFSVNQQSEMDTSVKFDLQIRSSNLFDNLSPTTSYQADLAILASVEIRGVSSPDHIFLPIANWQPQENPETEEDIGPLVQHIYELRNNGPSAFSKVMLNLQWPYKYNNNTLLYIMQYEIDGPMNCTSDMEINPLNIKISVPKADEKNETIRDEDNRDHHINRRDITAIEGDVHTLGCGTAECLKIVCQVGRLERGKSAILYVKSRLWTQTFMNKENQNHSYSLKSSASFNVIEFPYKNLSFEDIHNSTVVSHLCLNMMGFFKRVRPPQEEQEREQLQPHENGEGTSEA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0A2</accession>
    <name>A0A452J0A2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Set1/Ash2 histone methyltransferase complex subunit ASH2</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="14">ASH2-like protein</fullName>
        </alternativeName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="12">Transcriptional regulator. Component or associated component of some histone methyltransferase complexes which regulates transcription through recruitment of those complexes to gene promoters. Component of the Set1/Ash2 histone methyltransferase (HMT) complex, a complex that specifically methylates 'Lys-4' of histone H3, but not if the neighboring 'Lys-9' residue is already methylated. As part of the MLL1/MLL complex it is involved in methylation and dimethylation at 'Lys-4' of histone H3. May play a role in hematopoiesis. In association with RBBP5 and WDR5, stimulates the histone methyltransferase activities of KMT2A, KMT2B, KMT2C, KMT2D, SETD1A and SETD1B.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0A2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038268.1">
        <property type="protein sequence ID" value="ENSGAGP00000033786.1"/>
        <property type="gene ID" value="ENSGAGG00000024046.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071339">
        <property type="term" value="C:MLL1 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048188">
        <property type="term" value="C:Set1C/COMPASS complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000976">
        <property type="term" value="F:transcription cis-regulatory region binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006325">
        <property type="term" value="P:chromatin organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010557">
        <property type="term" value="P:positive regulation of macromolecule biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010468">
        <property type="term" value="P:regulation of gene expression"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd15583">
        <property type="entry name" value="PHD_ash2p_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12872">
        <property type="entry name" value="SPRY_Ash2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.980.20:FF:000002">
        <property type="entry name" value="Ash2 like, histone lysine methyltransferase complex subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.920:FF:000008">
        <property type="entry name" value="Set1/Ash2 histone methyltransferase complex subunit ASH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.920">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.980.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037353">
        <property type="entry name" value="ASH2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049455">
        <property type="entry name" value="ASH2-like_PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR053835">
        <property type="entry name" value="ASH2L-like_WH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001870">
        <property type="entry name" value="B30.2/SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043136">
        <property type="entry name" value="B30.2/SPRY_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003877">
        <property type="entry name" value="SPRY_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10598">
        <property type="entry name" value="SET1/ASH2 HISTONE METHYLTRANSFERASE COMPLEX SUBUNIT ASH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10598:SF0">
        <property type="entry name" value="SET1_ASH2 HISTONE METHYLTRANSFERASE COMPLEX SUBUNIT ASH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21198">
        <property type="entry name" value="ASH2L-like_WH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21257">
        <property type="entry name" value="PHD_ash2p_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00622">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00449">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50188">
        <property type="entry name" value="B302_SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0156">Chromatin regulator</keyword>
    <keyword evidence="9" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="2" id="KW-0488">Methylation</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0804">Transcription</keyword>
    <keyword evidence="8" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="B30.2/SPRY" evidence="16">
        <location>
            <begin position="386"/>
            <end position="609"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="1"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="285"/>
            <end position="357"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="15">
        <location>
            <begin position="41"/>
            <end position="77"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="15">
        <location>
            <begin position="303"/>
            <end position="321"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00056054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00068996"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00075872"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50188"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033786.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="654" mass="71825" checksum="6C66A949ABB23AC9" modified="2019-05-08" version="1">MQGRYPSAMGPSAGITPYGRKGKSRESPSQSLVRLRGRKMAAAAGATGAPAPEEGEVAAPPEGAAGDPPGPPEQGAGDSETGGDTTLVDVTAGLETESSNGKDPLEGVGDSSEVLDTQTGSVDEENGRQLGEIELQCGICTKWFTADTFGIDTTSCLPFMTNYSFHCNVCHHSGNTYFLRKQANLKEMCLSALANLTWQSRTQDEHPKTMFSKDKDIIPFIDKYWECMTTRQRPGKMTWPNNIVKTMCKERDVFLVKEHPDPGSKDPEDDYPKFGLLDQDLANIGPAYDNQKQSNAVSTSGSLNGGASLGGGIPAGSSGKGRGAKRKQQDGGTTGTAKKTRSDPLFSAQRLPPHGYPLEHPFNKDGYRYILAEPDPHAPDPEKLELDCWAGKPIPGDLYRACLYERVLLALHDRAPQLKISDDRLTVIGEKGYSMVRASHGVRKGDWYFEISVDEMPPDTAARLGWSQPLGNLQAPLGYDKFSYSWRSKKGTKFHQSIGKHYSAGYGQGDVLGFYISLPEDTETAKSLPDTYKDKALIKFKSYLYFEEKDFVDKAEKSLKQTPGSQIIFYKNGVSQGVAFKEIFEGVYFPAISLYKGCTVSVNFGPYFKYPPRNITYRPMSDMGWGAVVEHTLADVLYHVETEVDGRRSPPWEP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0A4</accession>
    <name>A0A452J0A4_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J0A4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038269.1">
        <property type="protein sequence ID" value="ENSGAGP00000033787.1"/>
        <property type="gene ID" value="ENSGAGG00000024042.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031440">
        <property type="entry name" value="DUF4670"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21937">
        <property type="entry name" value="CCDC66 DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21937:SF5">
        <property type="entry name" value="GENE 973-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="231"/>
            <end position="254"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="294"/>
            <end position="326"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="459"/>
            <end position="501"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="236"/>
            <end position="254"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="303"/>
            <end position="326"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033787.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="666" mass="75012" checksum="710DBD5985D750D9" modified="2019-05-08" version="1">MPNLSLLSRGTGQVVKKTQEKLEVHFEPEDYLNWKSHEGYGHVSRLLNERQFMKGCWGLYLPKTYSTKTGALVLYSEDLAEPSWKQRGGMSGQQGLRHRRNKLQIELHTLQDLTRAILAYGGRQKDQKETAWHPHLHFLKKPDSQIDRQIRPGYSAKRYLFRLSQTWDPSIIYRLQCAGYIRDPLLLQNSLNTPNHRIGQQDLSAVPKKYHLLPAFPSFWAELSQHIHPGQSYSVQKEDENREKQVEGDEVETRGQRRARVPVWVSVRRLSLELRPSPAKEATWSQNAACTKVPQGTEEDHQDVEHSMEGQWHGEKPHQTESQEPHEKLNQTVKSSVGSATIDTSQLLSERSYMTFYGGYFPGRKKPYSVKQGHLNNQDGKEGHLIETGFFPPIQSAMDSEQGVVRVEHRKQAPETLKFPSISEESPRVQVPHRKQFRSSDPPKELLILPLLIQLQTQPETKAKNQTAASCDKSRSEADDDCEKLLAPSPHDLISGPEGTTEQHQMVIGENPEIHQGSFFLPPITNQKFTLEGKKAGTRGVNAGKEGNYKRDGDLFTQNALPGLDLLPPKYGKKGPGNQSSKTDFKMFSSSTMTSSTSKTLPIGITRGTLPEELRECYNGTSLGSLIMGPDGEIVCLSLLGSVQDAEVPAQFSFVPDKAKGIQEKF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0A7</accession>
    <name>A0A452J0A7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">BAG domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J0A7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038314.1">
        <property type="protein sequence ID" value="ENSGAGP00000033825.1"/>
        <property type="gene ID" value="ENSGAGG00000024066.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000774">
        <property type="term" value="F:adenyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051087">
        <property type="term" value="F:protein-folding chaperone binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050821">
        <property type="term" value="P:protein stabilization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.120">
        <property type="entry name" value="BAG domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039773">
        <property type="entry name" value="BAG_chaperone_regulator"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036533">
        <property type="entry name" value="BAG_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003103">
        <property type="entry name" value="BAG_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12329:SF10">
        <property type="entry name" value="BAG FAMILY MOLECULAR CHAPERONE REGULATOR 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12329">
        <property type="entry name" value="BCL2-ASSOCIATED ATHANOGENE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02179">
        <property type="entry name" value="BAG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00264">
        <property type="entry name" value="BAG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF63491">
        <property type="entry name" value="BAG domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51035">
        <property type="entry name" value="BAG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0143">Chaperone</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="BAG" evidence="3">
        <location>
            <begin position="354"/>
            <end position="431"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1"/>
            <end position="263"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="2">
        <location>
            <begin position="53"/>
            <end position="70"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="111"/>
            <end position="127"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="2">
        <location>
            <begin position="164"/>
            <end position="173"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="2">
        <location>
            <begin position="183"/>
            <end position="195"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="240"/>
            <end position="252"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00023186"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS51035"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033825.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="432" mass="47251" checksum="84D2C6A0ABF8AA1C" modified="2019-05-08" version="1">MSALRRLLYGPEAERARPEPEPPPSPWGMEPRACPQHRGAGPAPAWPGDSYCPGGGSGWERSGRGGGGGPQQPPYPGYGSNYWNSAVQPRAPYPSPYPAGPELQSQGLESPYTNGAYGAPGAAPHYANLPQSNTYYSSGHPRASYPAESLSMYRPPSPASHWHYPPPDCPPEGPSLRRQAPGYSPPPTPGMPVPHYPYGGDSNPGIPQQGPPPQPRPQDESWASSGIYGMQPRYAWPAASTPQGSPFISESSPFWPGNGAPSPHPPAHYPISACYQGANHHNYFPEVNQQPSGTMNDHKSTQFNTRPSHFNPKVQYSAQPQMYNNVPRKPPLNQDAGFKPSDQPLSNSLGIQPGIQRVMQVMEEVEQLEQEVDEFVGKKTDKAYRLLEEMLTKELLELDSIETHGQDNVRQARKEAVHRIQAILEKLEKKGL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0A9</accession>
    <name>A0A452J0A9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Set1/Ash2 histone methyltransferase complex subunit ASH2</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="14">ASH2-like protein</fullName>
        </alternativeName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="12">Transcriptional regulator. Component or associated component of some histone methyltransferase complexes which regulates transcription through recruitment of those complexes to gene promoters. Component of the Set1/Ash2 histone methyltransferase (HMT) complex, a complex that specifically methylates 'Lys-4' of histone H3, but not if the neighboring 'Lys-9' residue is already methylated. As part of the MLL1/MLL complex it is involved in methylation and dimethylation at 'Lys-4' of histone H3. May play a role in hematopoiesis. In association with RBBP5 and WDR5, stimulates the histone methyltransferase activities of KMT2A, KMT2B, KMT2C, KMT2D, SETD1A and SETD1B.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0A9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038274.1">
        <property type="protein sequence ID" value="ENSGAGP00000033792.1"/>
        <property type="gene ID" value="ENSGAGG00000024046.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071339">
        <property type="term" value="C:MLL1 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048188">
        <property type="term" value="C:Set1C/COMPASS complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000976">
        <property type="term" value="F:transcription cis-regulatory region binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006325">
        <property type="term" value="P:chromatin organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010557">
        <property type="term" value="P:positive regulation of macromolecule biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010468">
        <property type="term" value="P:regulation of gene expression"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd15583">
        <property type="entry name" value="PHD_ash2p_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12872">
        <property type="entry name" value="SPRY_Ash2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.980.20:FF:000002">
        <property type="entry name" value="Ash2 like, histone lysine methyltransferase complex subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.920:FF:000008">
        <property type="entry name" value="Set1/Ash2 histone methyltransferase complex subunit ASH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.920">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.980.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037353">
        <property type="entry name" value="ASH2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049455">
        <property type="entry name" value="ASH2-like_PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR053835">
        <property type="entry name" value="ASH2L-like_WH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001870">
        <property type="entry name" value="B30.2/SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043136">
        <property type="entry name" value="B30.2/SPRY_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003877">
        <property type="entry name" value="SPRY_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10598">
        <property type="entry name" value="SET1/ASH2 HISTONE METHYLTRANSFERASE COMPLEX SUBUNIT ASH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10598:SF0">
        <property type="entry name" value="SET1_ASH2 HISTONE METHYLTRANSFERASE COMPLEX SUBUNIT ASH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21198">
        <property type="entry name" value="ASH2L-like_WH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21257">
        <property type="entry name" value="PHD_ash2p_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00622">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00449">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50188">
        <property type="entry name" value="B302_SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0156">Chromatin regulator</keyword>
    <keyword evidence="9" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="2" id="KW-0488">Methylation</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0804">Transcription</keyword>
    <keyword evidence="8" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="B30.2/SPRY" evidence="16">
        <location>
            <begin position="378"/>
            <end position="601"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="1"/>
            <end position="118"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="277"/>
            <end position="349"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="15">
        <location>
            <begin position="33"/>
            <end position="69"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="15">
        <location>
            <begin position="295"/>
            <end position="313"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00056054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00068996"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00075872"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50188"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033792.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="646" mass="70933" checksum="3989441852D7A4BF" modified="2019-05-08" version="1">MGPSAGITPYGRKGKSRESPSQSLVRLRGRKMAAAAGATGAPAPEEGEVAAPPEGAAGDPPGPPEQGAGDSETGGDTTLVDVTAGLETESSNGKDPLEGVGDSSEVLDTQTGSVDEENGRQLGEIELQCGICTKWFTADTFGIDTTSCLPFMTNYSFHCNVCHHSGNTYFLRKQANLKEMCLSALANLTWQSRTQDEHPKTMFSKDKDIIPFIDKYWECMTTRQRPGKMTWPNNIVKTMCKERDVFLVKEHPDPGSKDPEDDYPKFGLLDQDLANIGPAYDNQKQSNAVSTSGSLNGGASLGGGIPAGSSGKGRGAKRKQQDGGTTGTAKKTRSDPLFSAQRLPPHGYPLEHPFNKDGYRYILAEPDPHAPDPEKLELDCWAGKPIPGDLYRACLYERVLLALHDRAPQLKISDDRLTVIGEKGYSMVRASHGVRKGDWYFEISVDEMPPDTAARLGWSQPLGNLQAPLGYDKFSYSWRSKKGTKFHQSIGKHYSAGYGQGDVLGFYISLPEDTETAKSLPDTYKDKALIKFKSYLYFEEKDFVDKAEKSLKQTPGSQIIFYKNGVSQGVAFKEIFEGVYFPAISLYKGCTVSVNFGPYFKYPPRNITYRPMSDMGWGAVVEHTLADVLYHVETEVDGRRSPPWEP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0B2</accession>
    <name>A0A452J0B2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Calponin-homology (CH) domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the Nav/unc-53 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038319.1">
        <property type="protein sequence ID" value="ENSGAGP00000033830.1"/>
        <property type="gene ID" value="ENSGAGG00000024067.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0022008">
        <property type="term" value="P:neurogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00009">
        <property type="entry name" value="AAA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000316">
        <property type="entry name" value="Putative neuron navigator 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.418.10">
        <property type="entry name" value="Calponin-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003593">
        <property type="entry name" value="AAA+_ATPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003959">
        <property type="entry name" value="ATPase_AAA_core"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001715">
        <property type="entry name" value="CH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036872">
        <property type="entry name" value="CH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057568">
        <property type="entry name" value="CortBP2_NAV1-like_AAA_lid"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039041">
        <property type="entry name" value="Nav/unc-53"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057126">
        <property type="entry name" value="NAV1-like_ubiquitin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12784:SF18">
        <property type="entry name" value="NEURON NAVIGATOR 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12784">
        <property type="entry name" value="STEERIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00004">
        <property type="entry name" value="AAA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25408">
        <property type="entry name" value="AAA_lid_NAV1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23092">
        <property type="entry name" value="Ubiquitin_6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00382">
        <property type="entry name" value="AAA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47576">
        <property type="entry name" value="Calponin-homology domain, CH-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50021">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 3" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Calponin-homology (CH)" evidence="5">
        <location>
            <begin position="1"/>
            <end position="47"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="66"/>
            <end position="459"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="505"/>
            <end position="524"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="743"/>
            <end position="1185"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="1220"/>
            <end position="1245"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="1276"/>
            <end position="1323"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="1522"/>
            <end position="1606"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="1676"/>
            <end position="1707"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="3">
        <location>
            <begin position="1419"/>
            <end position="1502"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="66"/>
            <end position="106"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="121"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="168"/>
            <end position="191"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="198"/>
            <end position="215"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="216"/>
            <end position="226"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="290"/>
            <end position="313"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="328"/>
            <end position="354"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="384"/>
            <end position="398"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="420"/>
            <end position="445"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="508"/>
            <end position="524"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="748"/>
            <end position="776"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="777"/>
            <end position="791"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="792"/>
            <end position="802"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="848"/>
            <end position="857"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="884"/>
            <end position="896"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="943"/>
            <end position="961"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="1026"/>
            <end position="1039"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="1056"/>
            <end position="1065"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="1075"/>
            <end position="1096"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="1122"/>
            <end position="1132"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="1133"/>
            <end position="1158"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="4">
        <location>
            <begin position="1165"/>
            <end position="1174"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="1220"/>
            <end position="1244"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="1276"/>
            <end position="1290"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="1291"/>
            <end position="1300"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="1301"/>
            <end position="1311"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="1312"/>
            <end position="1323"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="1527"/>
            <end position="1544"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="1692"/>
            <end position="1707"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006255"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033830.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2192" mass="235111" checksum="2AC45A57A98A671F" modified="2019-05-08" version="1">MIENVDVCLSFLAARGVNVQGLSAEEIRNGNLKAILGLFFSLSRYKQQQHHQQQYYQSLVELQQHVSHSSPPAEISQSKTQQDMQSSLRARYATQSNHSGIATSQKKTSRLPGPSRVPAAGSSTKVQGASNLNRRSQSFNSIDKNKPPQYANSNEKDSPKGPHPSAGMNGNVQHPISTGQQPVSAIPSPSASKPWRSKSMNVKHSATSTMLSVKQPSPATSPTPSSDRLKPPLSEGVKPPSSGQKSMLEKFKLVNARTALRPPLSSSSGPNDSGREDDTFSECGEMDGFSSGLNSGGSTSSSPKVSPKLTPPKAGSKNLSNKKSLLQPKDKEEKNRDKNKVCTEKTVKEEKDQVPETSTKKSSKIASLIPKGSKTTAAKKESLIPSSSGIPKPGSKVPTAKQSASPVCAGSKESEKFRTTKGNQSQSTTKSQANEKASTSCNLASSEGKESSPALTPGSSAALSVSMAASSGQVTGNGIVQLPQQQQYSHPNTATVAPFIYRTHSENDSTSLPPADSCTSPTKMDLSYSKTAKQCLEEISGEDPETRRMRTVKNIADLRQNLEETMSSLRGTQISHSTLETTFDSTVTTEVNGRSIPGLTSRSTPVTWRLGQASPRLQAGDAPSLGSGYPRSSAGRFIHTDPSRFMYTTPLRRAAVSRLGNMSQIDMNEKGSSDLDMSSEVDVGGYMSDGDILGKSLRTDDISSGYMTDGGLNLYTRSLNRIPDLAASRDVIQRGVHDVTVDADSWDDSSSVSSGLSDTLDNLSTDDLNTTSSVSSYSNITASSRKNTHTQLKTDSEKRFVTDSENWGSTEELKKPEEDFDSNVDGSGKWKGLPSGLSEDSEKGGQKASLSVSQTGSWRRGMTAQVGTTSSRHKAGTSALKTPGKTDDAKASEKVKTPLKGTSIQRSPSDAGKSSGDEGKKPPSGIGRSTATGSFGFKKAGVGSSTIITTSGATITSGSATLGKMPKSTAIGGKSNAGRKTSLDGSQNQDDGVLHVNSKTTLQYRSLPRPSKSSTSGIPGRGGHRSSTSSIDSNLSSKSAGAATTKLREPTKIGSGRSSPITINQTDKEKEKVAVSDSESVSLSSSPKSSPTSASACGTPGLRQPGSKYPDIASPTFRRLFGAKAGGKAASAPNTESVKPTSAMPSPSTTLARQGSLESPSSGTGSMGSAGGQSGSSSPLFSKPSELNTDVVSLSHSLASSPASVHSFTSGGLVWAANLSSSSAGSKDTPSYQSMTSLHTSSESIDLPLSHHSSLSGLTTGTQEVQSLLMRTGSVRSTLSERYTPSSRQTSQEESKEWLRSHSTGGLQDTGSQSPIVSPSAMSSSATGKYHFSNLVSPTNLSQFNLPGPSMMRSNSIPAQDSSFDLYDDSQLCGSATSLEERPRAISHSGSFRDSMEEVHGSSLSLVSSTSSLYSTAEEKAHSEQIHKLRRELVASQEKVATLTSQLSANAHLVAAFEKSLGNMTGRLQSLTMTAEQKESELIELRETIEMLKAQNSAAQAAIQGALNGPDHAHKDLLIRRQHSSESVSSINSATSHSSIGSGNDADSKKKKKKNWLRSSFKQAFGKKKSTKPPSSHSDIEELTDSSLPSSPKLPHNSGDCGTLSMKPSQSASAICECTEAEAEIILQLKSELREKELKLTDIRLEALSSAHHLDQIREAMNRMQNEIEILKAENDRLKAETGNTGKPARPSSESSSSTSSSSSRQSLGLSLNNLNITDSVTSDILLDDVTDGALPKEGHSVKIIVTISKGYGRSKDQKTQAYLIGSIGVSGKTKWDVLDGVIRRLFKEYIFRVDPATSLGLSSDCIASYCIGDLIRVHSLEVPELLPCGYLVGDNNIITVNLKGVEENSLDSFVFDTLIPKPITQRYFNLLMEHHRIILSGPSGTGKTYLANRLAEYIITKLDRKKTEDAIATFNVDHKSSKDLRQYLANLAEQCSADNSDADLPVVIILDNLHHIGSLSDIFNGFLNCKYNTCPYIIGTMNQGVTSSPNLELHHNFRWVLCANHTEPVKGFLGRYLRRKLIETEIERNMRNNDLIKIIDWIPKTWHHLNSFLETHSSSDVTIGPRLFLPCPMDVDGSRVWFTDLWNYSLVPYLLEAVREGLQMYGKRAPWEDPSKWVADTYPWSSAPVQHEWPSLLQLRPEDVGFEGYASAQEGTTSKHIPQSDTEGDPLVRMKLFAFLCSLCFNLEFET</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0B5</accession>
    <name>A0A452J0B5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">NACHT domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J0B5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038276.1">
        <property type="protein sequence ID" value="ENSGAGP00000033794.1"/>
        <property type="gene ID" value="ENSGAGG00000024047.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045345">
        <property type="term" value="P:positive regulation of MHC class I biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045348">
        <property type="term" value="P:positive regulation of MHC class II biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045944">
        <property type="term" value="P:positive regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:001028">
        <property type="entry name" value="Class II major histocompatibility complex transactivator"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001611">
        <property type="entry name" value="Leu-rich_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025875">
        <property type="entry name" value="Leu-rich_rpt_4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008095">
        <property type="entry name" value="MHC_II_transact"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007111">
        <property type="entry name" value="NACHT_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041267">
        <property type="entry name" value="NLRP_HD2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47189">
        <property type="entry name" value="MHC CLASS II TRANSACTIVATOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47189:SF1">
        <property type="entry name" value="MHC CLASS II TRANSACTIVATOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12799">
        <property type="entry name" value="LRR_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13516">
        <property type="entry name" value="LRR_6"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05729">
        <property type="entry name" value="NACHT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17776">
        <property type="entry name" value="NLRC4_HD2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01719">
        <property type="entry name" value="MHCIIACTVATR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00368">
        <property type="entry name" value="LRR_RI"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52047">
        <property type="entry name" value="RNI-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51450">
        <property type="entry name" value="LRR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50837">
        <property type="entry name" value="NACHT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="1" id="KW-0433">Leucine-rich repeat</keyword>
    <keyword evidence="3" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="NACHT" evidence="5">
        <location>
            <begin position="471"/>
            <end position="606"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022614"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033794.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1200" mass="135786" checksum="C89E862A0CD56B8D" modified="2019-05-08" version="1">MLEKDVISKEYHQTLLQEKDREDLARKISLTVVEKWDLCLNTLVPLRCLKLYTKMPQDITDNESSLNRGIAASKRQTMDSAVLTENSFLDLLHSNIDPLHLYSLLDTRLSDEESELSTDPEIDTINHDQFSNLDVFYTMESNESVEESIFSNAGEAYSRIAGLAEYVLKDQQERQMEDIFGKLILDEMANETTEGFTEMKKQKCQKRSFVSTAESYSDASKPKCRRPDAARRMTEQVCTEGSRAVGDVRASCQKRVSSGNSNPDCLLVGAKGIQVISGFAVLPQQIINLPVGNGASNIIIYPVSSSPTETLQISPAFPTTLAAQVDLAVQSILGSPDKNTSIGPFAYMPETAAVTDLTDEEEISNEISQIPVVPSEQILKRPKPVEAFCTSLKDYFQDVCKFMAMDREVTLDHLYIDGALMQGQTETKSGKNVIKVMEKELVIYNLEEKEKAAIERSQIFQVLGGKELETKVIVVLGKAGMGKSILVQKICQDWSNGKFSQFEFVFWFECRRLSLPEKQYSLKDLLLELFVKPREGRDQVFEYILQNPNKVLLIFDGFEELQDHDGFIHYPDCQSYKELYSIRELFAGLFQKKILSGCTLLLTARPKDKFNQYLSKVDKIIEMIGFSPQQVELYIARYFEGLPYSDDAVKLITNCQYLFSHCYSPVICRFICFLCEMMFEMGDKDLPPTLTSLFVKYLQQKLIPTQTDVTAIGNQQHVTRLAQLAWYLGEKNQNALKSSLFPSKEVKEFALKYGLVLPFTFPKHSGNGEEEFGSAFSDFVIQNFLSALHLLLAEDIKDKSLTKYLSLPARKKKPHNWLDLVPRFLPGLLYLQHDPFLCSLSDEHVMKIITKKQKTLLKYIKKIQISDLYPEKLLELLHCVYETQDSYLLQHVALRIKAELSFLGTLLTPPDVYVLHFILKRSPKEFSLDLRNSSINLQGLKHLVGLKNVTSFRASLSDTVRLWESLERTKEYELLKISVEKFDIDPFKAETMKDIHDLSDLVQIQEKMIHCMEEAFGSCYQIPAIKNLRKLEFALGPACGLQGFLKLVEILAAFPSLQHLDLDALSENEIGDEGTKSLCEVLSELTSLETLNLSQNKITDLGAEQLATALPSLSSLKTLSLYNNYICDAGAENIAKILPAMASLRVLHVQCNKITDAGAQKLTDSLRKCPHIKSVAMWNPTIPYGVLEHLQLLDSRISLL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0B8</accession>
    <name>A0A452J0B8_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="21">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 16">Membrane</location>
            <topology evidence="1 16">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 16">Belongs to the integrin alpha chain family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0B8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038324.1">
        <property type="protein sequence ID" value="ENSGAGP00000033835.1"/>
        <property type="gene ID" value="ENSGAGG00000023825.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009897">
        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008305">
        <property type="term" value="C:integrin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001525">
        <property type="term" value="P:angiogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033627">
        <property type="term" value="P:cell adhesion mediated by integrin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098609">
        <property type="term" value="P:cell-cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007160">
        <property type="term" value="P:cell-matrix adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007229">
        <property type="term" value="P:integrin-mediated signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1510:FF:000001">
        <property type="entry name" value="Integrin alpha V"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1530:FF:000002">
        <property type="entry name" value="integrin alpha-V isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.930:FF:000001">
        <property type="entry name" value="Integrin subunit alpha V"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1460:FF:000001">
        <property type="entry name" value="Integrin, alpha V"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.930">
        <property type="entry name" value="Bicelle-embedded integrin alpha(iib) transmembrane segment"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.130">
        <property type="entry name" value="Integrin alpha, N-terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1460">
        <property type="entry name" value="Integrin domains. Chain A, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1510">
        <property type="entry name" value="ntegrin, alpha v. Chain A, domain 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1530">
        <property type="entry name" value="ntegrin, alpha v. Chain A, domain 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013517">
        <property type="entry name" value="FG-GAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013519">
        <property type="entry name" value="Int_alpha_beta-p"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000413">
        <property type="entry name" value="Integrin_alpha"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018184">
        <property type="entry name" value="Integrin_alpha_C_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013649">
        <property type="entry name" value="Integrin_alpha_Ig-like_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048285">
        <property type="entry name" value="Integrin_alpha_Ig-like_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048286">
        <property type="entry name" value="Integrin_alpha_Ig-like_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028994">
        <property type="entry name" value="Integrin_alpha_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032695">
        <property type="entry name" value="Integrin_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23220">
        <property type="entry name" value="INTEGRIN ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23220:SF4">
        <property type="entry name" value="INTEGRIN ALPHA-V"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01839">
        <property type="entry name" value="FG-GAP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08441">
        <property type="entry name" value="Integrin_A_Ig_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20805">
        <property type="entry name" value="Integrin_A_Ig_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20806">
        <property type="entry name" value="Integrin_A_Ig_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00357">
        <property type="entry name" value="Integrin_alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01185">
        <property type="entry name" value="INTEGRINA"/>
    </dbReference>
    <dbReference type="SMART" id="SM00191">
        <property type="entry name" value="Int_alpha"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69318">
        <property type="entry name" value="Integrin alpha N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69179">
        <property type="entry name" value="Integrin domains"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51470">
        <property type="entry name" value="FG_GAP"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00242">
        <property type="entry name" value="INTEGRIN_ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0106">Calcium</keyword>
    <keyword evidence="8 16" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="14" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10 16" id="KW-0401">Integrin</keyword>
    <keyword evidence="11 16" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13 16" id="KW-0675">Receptor</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <keyword evidence="3 16" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9 16" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="797"/>
            <end position="819"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="45"/>
            <end position="99"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="100"/>
            <end position="165"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="166"/>
            <end position="223"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="15">
        <location>
            <begin position="227"/>
            <end position="290"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha first immunoglubulin-like" evidence="18">
        <location>
            <begin position="275"/>
            <end position="433"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha second immunoglobulin-like" evidence="19">
        <location>
            <begin position="434"/>
            <end position="562"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha third immunoglobulin-like" evidence="20">
        <location>
            <begin position="580"/>
            <end position="786"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="830"/>
            <end position="851"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="17">
        <location>
            <begin position="830"/>
            <end position="845"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023037"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00803"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="RuleBase" id="RU003762"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF08441"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="Pfam" id="PF20805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="Pfam" id="PF20806"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033835.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="851" mass="94354" checksum="00D2001215F5004B" modified="2019-05-08" version="1">ALNADSAQLSILMGFFPAGQLISDRVAEIVSKYNSKVYSIKYDDQLATRSASAAFDDSYLGYSVAVGEFSGDGIEDFVSGVPRAARTLGMVSIYNGKNMSSLYNFTGEQMAAYFGYSVAATDINGDNYTDLFIGAPLFMERGSDGKLQEVGQVSICLQQASGGFQIIKLNGFEIFARFSSSIAPLGDLDQDGFNDIAVAAPYGGEDKRGVVYIYNGRATGLNAVPSQILEGQWAARTMPPSFGYSLKGATDVDKNGYPDLIVGAFGVDTAVLYRARPVISVNAGLEVNPTILNPEEKICELPDTRVKVSCFNVKFCLTADGKGKLPNTLTFQVELLLDKLKQKGALRRALFLHNKQPGHSKNMTVVKGGTMRCEELQAFLRDESEFRDKLTPITVFMEYRLDYKTAKDSAGLQPILNQFTPANISRQTHILLDCGEDNICKPKLQVAVHSDQKQIYIGDDNPLTLIVDAQNQGEGAYEAELFVIIPPQADFIGVVRNNEALARLSCAFKTENQTRLVVCDLGNPMKAGTKLLAGLRFSVNQQSEMDTSVKFDLQIRSSNLFDNLSPTTSYQADLAILASVEIRGVSSPDHIFLPIANWQPQENPETEEDIGPLVQHIYELRNNGPSAFSKVMLNLQWPYKYNNNTLLYIMQYEIDGPMNCTSDMEINPLNIKVNEKNETIRDEDNRDHHINRRDITAIEGDVHTLGCGTAECLKIVCQVGRLERGKSAILYVKSRLWTQTFMNKENQNHSYSLKSSASFNVIEFPYKNLSFEDIHNSTVVTTNITWGIQPQPMPVPVWVIILAVLAGLLLLAVLVFVMYRMGFFKRVRPPQEEQEREQLQPHENGEGTSEA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0C0</accession>
    <name>A0A452J0C0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="27">Bone morphogenetic protein receptor type-2</fullName>
            <ecNumber evidence="5">2.7.11.30</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="28">Bone morphogenetic protein receptor type II</fullName>
        </alternativeName>
    </protein>
    <organism evidence="34 35">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="35" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="34" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="34" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="25">On ligand binding, forms a receptor complex consisting of two type II and two type I transmembrane serine/threonine kinases. Type II receptors phosphorylate and activate type I receptors which autophosphorylate, then bind and activate SMAD transcriptional regulators. Can also mediate signaling through the activation of the p38MAPK cascade. Binds to BMP7, BMP2 and, less efficiently, BMP4. Binding is weak but enhanced by the presence of type I receptors for BMPs. Mediates induction of adipogenesis by GDF6. Promotes signaling also by binding to activin A/INHBA.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="23">
            <text>L-seryl-[receptor-protein] + ATP = O-phospho-L-seryl-[receptor-protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:18673"/>
            <dbReference type="Rhea" id="RHEA-COMP:11022"/>
            <dbReference type="Rhea" id="RHEA-COMP:11023"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.30"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="24">
            <text>L-threonyl-[receptor-protein] + ATP = O-phospho-L-threonyl-[receptor-protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:44880"/>
            <dbReference type="Rhea" id="RHEA-COMP:11024"/>
            <dbReference type="Rhea" id="RHEA-COMP:11025"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.30"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="2">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29035"/>
        </cofactor>
    </comment>
    <comment type="subunit">
        <text evidence="26">Interacts with GDF5. Interacts with BMP4. Interacts with SCUBE3. Interacts with TSC22D1/TSC-22. Interacts with activin A/INHBA.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cell membrane</location>
            <topology evidence="3">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the protein kinase superfamily. TKL Ser/Thr protein kinase family. TGFB receptor subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.30" evidence="5"/>
    <dbReference type="AlphaFoldDB" id="A0A452J0C0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038287.1">
        <property type="protein sequence ID" value="ENSGAGP00000033799.1"/>
        <property type="gene ID" value="ENSGAGG00000024058.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030054">
        <property type="term" value="C:cell junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043226">
        <property type="term" value="C:organelle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043235">
        <property type="term" value="C:receptor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036122">
        <property type="term" value="F:BMP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005024">
        <property type="term" value="F:transforming growth factor beta receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001568">
        <property type="term" value="P:blood vessel development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030509">
        <property type="term" value="P:BMP signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030324">
        <property type="term" value="P:lung development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008285">
        <property type="term" value="P:negative regulation of cell population proliferation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001649">
        <property type="term" value="P:osteoblast differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045597">
        <property type="term" value="P:positive regulation of cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0040017">
        <property type="term" value="P:positive regulation of locomotion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045778">
        <property type="term" value="P:positive regulation of ossification"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045944">
        <property type="term" value="P:positive regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001558">
        <property type="term" value="P:regulation of cell growth"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0014916">
        <property type="term" value="P:regulation of lung blood pressure"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:2000026">
        <property type="term" value="P:regulation of multicellular organismal development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009888">
        <property type="term" value="P:tissue development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd14054">
        <property type="entry name" value="STKc_BMPR2_AMHR2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd23614">
        <property type="entry name" value="TFP_LU_ECD_BMPR2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000180">
        <property type="entry name" value="Receptor protein serine/threonine kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.60.10:FF:000006">
        <property type="entry name" value="Receptor protein serine/threonine kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000174">
        <property type="entry name" value="Receptor protein serine/threonine kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.60.10">
        <property type="entry name" value="CD59"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000472">
        <property type="entry name" value="Activin_recp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045860">
        <property type="entry name" value="Snake_toxin-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000333">
        <property type="entry name" value="TGFB_receptor"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23255:SF63">
        <property type="entry name" value="BONE MORPHOGENETIC PROTEIN RECEPTOR TYPE-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23255">
        <property type="entry name" value="TRANSFORMING GROWTH FACTOR-BETA RECEPTOR TYPE I AND II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01064">
        <property type="entry name" value="Activin_recp"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57302">
        <property type="entry name" value="Snake toxin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="15 29" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="19" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="21" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="14" id="KW-0418">Kinase</keyword>
    <keyword evidence="16" id="KW-0460">Magnesium</keyword>
    <keyword evidence="22" id="KW-0464">Manganese</keyword>
    <keyword evidence="18 31" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13 29" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="8" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="20" id="KW-0675">Receptor</keyword>
    <keyword evidence="35" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="12 32" id="KW-0732">Signal</keyword>
    <keyword evidence="9" id="KW-0808">Transferase</keyword>
    <keyword evidence="10 31" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="17 31" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="32">
        <location>
            <begin position="1"/>
            <end position="24"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019143792" description="Bone morphogenetic protein receptor type-2" evidence="32">
        <location>
            <begin position="25"/>
            <end position="1005"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="31">
        <location>
            <begin position="155"/>
            <end position="176"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="33">
        <location>
            <begin position="207"/>
            <end position="508"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="30">
        <location>
            <begin position="596"/>
            <end position="630"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="30">
        <location>
            <begin position="750"/>
            <end position="778"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="30">
        <location>
            <begin position="875"/>
            <end position="916"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="30">
        <location>
            <begin position="930"/>
            <end position="980"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="30">
        <location>
            <begin position="607"/>
            <end position="630"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="30">
        <location>
            <begin position="887"/>
            <end position="900"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="30">
        <location>
            <begin position="901"/>
            <end position="912"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="30">
        <location>
            <begin position="930"/>
            <end position="951"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="30">
        <location>
            <begin position="952"/>
            <end position="967"/>
        </location>
    </feature>
    <feature type="binding site" evidence="29">
        <location>
            <position position="234"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001936"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00009605"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012401"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00023211"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00047681"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00048773"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00057734"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00063997"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="ARBA" id="ARBA00073640"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="ARBA" id="ARBA00077874"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="31">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="32">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="33">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="34">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033799.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="35">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1005" mass="112014" checksum="0F6400ABEE2009AC" modified="2019-05-08" version="1">MSASLQHLLLVLLLSSATLLLSTAAATQSEERLCAFKDPYQQDHGISENRISQENGTILCVKGTSCYGLWEKTREGDIHLVKQGCWSHIGDLQECQYEECIVTTTPSLIQNGTYRFCCCSTDLCNVNFTENFPPPDPTDTPLFNSSNSFHRDETIVIALASVSVLAVLIAAIFFGYRMLAGDRKQGLHSMNMMEAAASEPSLDLDNLKLLELIGRGRYGAVYKGSLDEHPVAVKVFSFANRQNFINERNIYRVPLMDHDNIARFIVGDERFTADGRMEYLLVMEYYPNGSLCKYLSLHTSDWVSSCRLAHSVTRGLAYLHTELPRGDHYKPAISHRDLNSRNVLVKNDGTCVISDFGLSMRLTGNRLVRPGEEDNAAISEVGTIRYMAPEVLEGAVNLRDCESALKQVDMYALGLIYWEIFMRCTDLFPGESVPEYQMAFQTEVGNHPTFEDMQVLVSREKQRPKFPEAWKENSLAVRSLKETIEDCWDQDAEARLTAQCAEERMAELMMIWERNKSVSPTVNPMSTAMQNERNLSHHRRVPKIRPYPDYSSSSYIEDSIHHTDSIVKNISSEHSVSTTPLTSGEKNRNSINYERQQAQARIPSPETSVTSLSTNTTTTNTTGLTPSTGMTTISEMPYSDETNQHATNVMQPVGPTPVCLQLTEEDLETNKLDPKEVDKNLKESSDENLMEHSLKQFSGPDPLSSTSSSLLYPLIKLAAEVTGQQDFTQAGNGQACLIPDVPSTQVYPLPKQQNLPKRPTSLPLNTKNSTKEPRLKFGGKHKSNLKQVETGVAKMNTINAAEPHVVTVTMNGVVGRSQSINSHGVTTQYANGAVPSGQTTSSVAHRAQEMLQNQFSGEDSRLNINSSPDEHEPLLRREQQVGQDEGVLDRLVDRRERPLDNGRTNANNNNSNPCLMQDVTTLGTQRMTTYPGQTQTRRAQRPNSLDLSATNSLDSSSLQLGDSSQDGKPGSGEKIKKRVKTPYSLKRWRPSTWVISTEPLDCEET</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0C1</accession>
    <name>A0A452J0C1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Glucose-6-phosphate exchanger SLC37A2</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="13">Solute carrier family 37 member 2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>D-glucose 6-phosphate(in) + phosphate(out) = D-glucose 6-phosphate(out) + phosphate(in)</text>
            <dbReference type="Rhea" id="RHEA:71535"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:61548"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the major facilitator superfamily. Organophosphate:Pi antiporter (OPA) (TC 2.A.1.4) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0C1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038244.1">
        <property type="protein sequence ID" value="ENSGAGP00000033763.1"/>
        <property type="gene ID" value="ENSGAGG00000024013.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061513">
        <property type="term" value="F:glucose 6-phosphate:phosphate antiporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035435">
        <property type="term" value="P:phosphate ion transmembrane transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd17344">
        <property type="entry name" value="MFS_SLC37A1_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1250.20:FF:000050">
        <property type="entry name" value="glucose-6-phosphate exchanger SLC37A2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1250.20:FF:000028">
        <property type="entry name" value="Sugar phosphate exchanger 3 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1250.20">
        <property type="entry name" value="MFS general substrate transporter like domains"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011701">
        <property type="entry name" value="MFS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020846">
        <property type="entry name" value="MFS_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036259">
        <property type="entry name" value="MFS_trans_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044740">
        <property type="entry name" value="SLC37A1_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000849">
        <property type="entry name" value="Sugar_P_transporter"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43184:SF9">
        <property type="entry name" value="GLUCOSE-6-PHOSPHATE EXCHANGER SLC37A2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43184">
        <property type="entry name" value="MAJOR FACILITATOR SUPERFAMILY TRANSPORTER 16, ISOFORM B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07690">
        <property type="entry name" value="MFS_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF002808">
        <property type="entry name" value="Hexose_phosphate_transp"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103473">
        <property type="entry name" value="MFS general substrate transporter"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50850">
        <property type="entry name" value="MFS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0050">Antiport</keyword>
    <keyword evidence="7" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9 15" id="KW-0472">Membrane</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0762">Sugar transport</keyword>
    <keyword evidence="6 15" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 15" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="20"/>
            <end position="38"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="114"/>
            <end position="136"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="182"/>
            <end position="201"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="207"/>
            <end position="228"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="283"/>
            <end position="304"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="324"/>
            <end position="341"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="348"/>
            <end position="365"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="377"/>
            <end position="402"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="414"/>
            <end position="436"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="448"/>
            <end position="468"/>
        </location>
    </feature>
    <feature type="domain" description="Major facilitator superfamily (MFS) profile" evidence="16">
        <location>
            <begin position="27"/>
            <end position="471"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="244"/>
            <end position="265"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004477"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009598"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022449"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022597"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00034251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00039218"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00042041"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50850"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033763.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="496" mass="53718" checksum="EBBA9AF98CCCF12E" modified="2019-05-08" version="1">MGYSSLPTPGSWGAHQLYVVYRGLTLVLTFVFYASFHLSRKPISIVKSELHTNCSSLEKIPPNDTNSTTWCDWAPFDQANYNELFGALDNAFLVAYAIGMFISGIFGERLPLRYYLSGGMVLSGLFTSLFGLAYFWEIHVLWYYIIIQICNGLVQTTGWPSVVTCVGNWFGKGKRGLIMGIWNSHTSVGNILGSLIAGVWVSSAWGLSFIVPGIIIAIMGIVCFLFLVECECELEEKDPEVTLSADDGSCSRENTIDSSEYPKEPAEQPGAISFLGALRIPGVVEFSLCLLFAKLVSYTFLYWLPLYIVNVAHFSAKEAGDLSTLFDVGGILGGILAGLISDYTDGRATTCCVMLIVAAPMLFLYNHVGQNGIGSSIAMLIVCGALVNGPYALITTAVSADLGTHECLKGNAKALSTVTAIIDGTGSVGAALGPLLAGLISPTGWNNVFYMLITADILACLFLSRVVYKEIKGWCGCYTRKRGYVLLKSQVCRVFL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0C5</accession>
    <name>A0A452J0C5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Myosin motor domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="12">Muscle contraction.</text>
    </comment>
    <comment type="subunit">
        <text evidence="13">Muscle myosin is a hexameric protein that consists of 2 heavy chain subunits (MHC), 2 alkali light chain subunits (MLC) and 2 regulatory light chain subunits (MLC-2).</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Myofibril</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 14">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038292.1">
        <property type="protein sequence ID" value="ENSGAGP00000033804.1"/>
        <property type="gene ID" value="ENSGAGG00000020256.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030016">
        <property type="term" value="C:myofibril"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032982">
        <property type="term" value="C:myosin filament"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016460">
        <property type="term" value="C:myosin II complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000146">
        <property type="term" value="F:microfilament motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd01377">
        <property type="entry name" value="MYSc_class_II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000003">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000004">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000006">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000003">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000007">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000008">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.530:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Myosin heavy chain, isoform J"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Myosin heavy chain, muscle"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.370">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.4820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.2420">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.1160">
        <property type="entry name" value="Vasodilator-stimulated phosphoprotein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002928">
        <property type="entry name" value="Myosin_tail"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014751">
        <property type="entry name" value="XRCC4-like_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615">
        <property type="entry name" value="MYOSIN HEAVY CHAIN, NON-MUSCLE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615:SF69">
        <property type="entry name" value="MYOSIN-6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01576">
        <property type="entry name" value="Myosin_tail_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90257">
        <property type="entry name" value="Myosin rod fragments"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57997">
        <property type="entry name" value="Tropomyosin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="11 14" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="6 14" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9 14" id="KW-0505">Motor protein</keyword>
    <keyword evidence="10" id="KW-0514">Muscle protein</keyword>
    <keyword evidence="8 14" id="KW-0518">Myosin</keyword>
    <keyword evidence="5 14" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0787">Thick filament</keyword>
    <feature type="domain" description="Myosin motor" evidence="16">
        <location>
            <begin position="69"/>
            <end position="731"/>
        </location>
    </feature>
    <feature type="region of interest" description="Actin-binding" evidence="14">
        <location>
            <begin position="608"/>
            <end position="630"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="1205"/>
            <end position="1227"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="15">
        <location>
            <begin position="1205"/>
            <end position="1224"/>
        </location>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <begin position="162"/>
            <end position="169"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004657"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022433"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023179"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00037488"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00038612"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033804.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1873" mass="216519" checksum="6664D2125222888E" modified="2019-05-08" version="1">MGDAHMAEFGAAAPYLRKSDKERLEAQTRIFDLRSDCFVSDEQEEFVKGKITVTVKESDVHQQNPPKFDKIEDMAMLTFLHEPAVLYNLKERYAAWMIYTYSGLFCVTVNPYKWLPVYNAEVVAAYRGKKRSEAPPHIFSISDNAYQNMLTNRENQSILITGESGAGKTVNTKRVIQYFASIAAISDRKKEGTSDQIIQANPALEAFGNAKTVRNDNSSRFGKFIRIHFGATGKLASADIETYLLEKSRVIFQLKAERNYHIFYQILSNQKPELLGESLTLHPSPPLPPPGPQSAFDVLGFTPEEKVGVYKLTGAIMHFGNMKFKQKQREEQAEPDGTEDADKSAYLMGLNSADLLKGLCHPRVKVGNEYVTKGQNVQQVYYSIGALAKSVYEKMFSWMVTRINSTLETKQPRQYFIGVLDIAGFEIFDFNSFEQLCINFTNEKLQQFFNHHMFVLEQEEYKKEGIEWVFIDFGMDLQACIDLIEKPMGIMSILEEECMFPKATDMTFKAKLFDNHLGKSANFSKPRNIKGKPEAHFSLIHYAGTVDYNILGWLQKNKDPLNETVVGLYQKSSLKLLALLFANYAAAESGEKTRQGSYYTPTPSRFLTLALMANLKTTHPHFVRCIIPNERKAPGVMDNPLVMHQLRCNGVLEGIRICRKGFPNRILYGDFRQRYRILNPAAIPEGQFIDSRKGAEKLLSSLDIDPNQYKFGHTKVFFKAGLLGLLEEMRDERLSRIITRIQAQSRGQLMRIEFKKILERRDALLVIQWNIRAFMGVKNWPWMKLYFKIKPLLKSAETEKEMQTMKEEFGRLKEALEKSEARRKELEEKMVSLLQEKNDLQLQVQAEQDNLNDAEERCDQLIKNKIQLEAKVKELTERMEDEEEMNAELTAKKRKLEDECSELKKDIDDLELTLAKVEKEKHATENKVKNLTEEMAGLDEIIAKLTKEKKSLQEAHQQALDDLQVEEDKVNSLTKAKLKLEQQADDLEGSLEQEKKIRMDLERAKRKLEGDLKLTQENIMDLENDKQQLEEKLKKKEFDINQQNSKIEDEQALGLQLQKKLKELQARIEELEEELEAERTARAKVEKLRSDLSRELEEISERLEEAGGATSVQIELNKKREAEFQKMRRDLEEATLQHEATAATLRKKHADSVAELGEQIDNLQRVKQKLEKEKSELKLELDDISSNMEQLLKAKHRAKLEETQRAMNDTSTQRAKLQTENGELSRQLEEKEALISQLTRGKQSYTQQMEDLKRQLEEETKAKNALAHALQSARHDCDLLREQYEEETEAKAELQRSLSKANSEVAQWRTKYETDAIQRTEELEEAKKKLAQRLQEAEEAVEAVNAKCSSLEKTKHRLQNEIEDLMVDVERSNAAAAALDKKQRNFDKILSEWKQKFEESQTELESSQKEARSLSTELFKLKNAYEESLEHLETFKRENKNLQEEISDLTEQLGEGGKNLHELEKVRKQLEAEKMELQAALEEAEASLEHEEGKILRAQLEFNQFKADFERKLAEKDEEMEQAKRNHLRVVDSLQTSLDAETRSRNEALRVKKKMEGDLNEMEIQLSHANRMAAEAQSHLKATQTQLKDSQLQLDDFIRANEDLKENIAIVERRNSLLQAELEELRAVVEQTERTRRLAEQELIEASERVQLLHTQNTSLINQKKKMEADLSQLQTEVEEAVQECRNAEEKAKKAITDAAMMAEELKKEQDTSAHLERMKKNMEQTIKDLQLRLDEAEQLALKGGKKQLQKLEARVRELENELESEQKRNAESIKGLRKSERRIKELTYQTEEDKKNLMRLQDLVDKLQMKVKAYKRQAEEAEEQANTNLAKFRKVQHELDEAEERADIAESQVNKLRAKSRDTTFYCMREDN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0D0</accession>
    <name>A0A452J0D0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Glucose-6-phosphate exchanger SLC37A2</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="13">Solute carrier family 37 member 2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>D-glucose 6-phosphate(in) + phosphate(out) = D-glucose 6-phosphate(out) + phosphate(in)</text>
            <dbReference type="Rhea" id="RHEA:71535"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:61548"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the major facilitator superfamily. Organophosphate:Pi antiporter (OPA) (TC 2.A.1.4) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0D0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038254.1">
        <property type="protein sequence ID" value="ENSGAGP00000033773.1"/>
        <property type="gene ID" value="ENSGAGG00000024013.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061513">
        <property type="term" value="F:glucose 6-phosphate:phosphate antiporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035435">
        <property type="term" value="P:phosphate ion transmembrane transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd17344">
        <property type="entry name" value="MFS_SLC37A1_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1250.20:FF:000050">
        <property type="entry name" value="glucose-6-phosphate exchanger SLC37A2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1250.20:FF:000028">
        <property type="entry name" value="Sugar phosphate exchanger 3 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1250.20">
        <property type="entry name" value="MFS general substrate transporter like domains"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011701">
        <property type="entry name" value="MFS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020846">
        <property type="entry name" value="MFS_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036259">
        <property type="entry name" value="MFS_trans_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044740">
        <property type="entry name" value="SLC37A1_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000849">
        <property type="entry name" value="Sugar_P_transporter"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43184:SF9">
        <property type="entry name" value="GLUCOSE-6-PHOSPHATE EXCHANGER SLC37A2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43184">
        <property type="entry name" value="MAJOR FACILITATOR SUPERFAMILY TRANSPORTER 16, ISOFORM B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07690">
        <property type="entry name" value="MFS_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF002808">
        <property type="entry name" value="Hexose_phosphate_transp"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103473">
        <property type="entry name" value="MFS general substrate transporter"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50850">
        <property type="entry name" value="MFS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0050">Antiport</keyword>
    <keyword evidence="7" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9 15" id="KW-0472">Membrane</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0762">Sugar transport</keyword>
    <keyword evidence="6 15" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 15" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="39"/>
            <end position="56"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="63"/>
            <end position="85"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="91"/>
            <end position="119"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="131"/>
            <end position="150"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="156"/>
            <end position="177"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="246"/>
            <end position="267"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="287"/>
            <end position="304"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="316"/>
            <end position="343"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="404"/>
            <end position="424"/>
        </location>
    </feature>
    <feature type="domain" description="Major facilitator superfamily (MFS) profile" evidence="16">
        <location>
            <begin position="1"/>
            <end position="427"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="188"/>
            <end position="228"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004477"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009598"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022449"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022597"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00034251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00039218"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00042041"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50850"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033773.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="452" mass="48885" checksum="C32FDF330636F4A6" modified="2019-05-08" version="1">TNCSSLEKIPPNDTNSTTWCDWAPFDQANYNELFGALDNAFLVAYAIGMFISGIFGERLPLRYYLSGGMVLSGLFTSLFGLAYFWEIHVLWYYIIIQICNGLVQTTGWPSVVTCVGNWFGKGKRGLIMGIWNSHTSVGNILGSLIAGVWVSSAWGLSFIVPGIIIAIMGIVCFLFLVEYPEDVDCSPPLHHTESEESEEKDPEVTLSADDGSCSRENTIDSSEYPKEPAEQPGAISFLGALRIPGVVEFSLCLLFAKLVSYTFLYWLPLYIVNVAHFSAKEAGDLSTLFDVGGILGGILAGLISDYTDGRATTCCVMLIVAAPMVLLILFLFAAMLIVCGALVNGPYALITTAVSADLGTHECLKGNAKALSTVTAIIDGTGSVGAALGPLLAGLISPTGWNNVFYMLITADILACLFLSRVVYKEIKGWCGCYTRKRGYVLLKSQVCRVFL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0D1</accession>
    <name>A0A452J0D1_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="19">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">Muscle contraction.</text>
    </comment>
    <comment type="subunit">
        <text evidence="14">Muscle myosin is a hexameric protein that consists of 2 heavy chain subunits (MHC), 2 alkali light chain subunits (MLC) and 2 regulatory light chain subunits (MLC-2).</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Myofibril</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 15">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038297.1">
        <property type="protein sequence ID" value="ENSGAGP00000033809.1"/>
        <property type="gene ID" value="ENSGAGG00000020256.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030016">
        <property type="term" value="C:myofibril"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032982">
        <property type="term" value="C:myosin filament"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016460">
        <property type="term" value="C:myosin II complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000146">
        <property type="term" value="F:microfilament motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd01377">
        <property type="entry name" value="MYSc_class_II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000003">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000004">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000006">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000013">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000003">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000007">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000008">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.4820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.530:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.360:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000024">
        <property type="entry name" value="Myosin heavy chain, isoform J"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.720:FF:000001">
        <property type="entry name" value="Myosin heavy chain, muscle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.340">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.370">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.4820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.2420">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.360">
        <property type="entry name" value="Myosin S1 fragment, N-terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008989">
        <property type="entry name" value="Myosin_S1_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002928">
        <property type="entry name" value="Myosin_tail"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004009">
        <property type="entry name" value="SH3_Myosin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014751">
        <property type="entry name" value="XRCC4-like_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615">
        <property type="entry name" value="MYOSIN HEAVY CHAIN, NON-MUSCLE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615:SF69">
        <property type="entry name" value="MYOSIN-6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02736">
        <property type="entry name" value="Myosin_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01576">
        <property type="entry name" value="Myosin_tail_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90257">
        <property type="entry name" value="Myosin rod fragments"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51844">
        <property type="entry name" value="SH3_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12 15" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="7 15" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="8" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0488">Methylation</keyword>
    <keyword evidence="10 15" id="KW-0505">Motor protein</keyword>
    <keyword evidence="11" id="KW-0514">Muscle protein</keyword>
    <keyword evidence="9 15" id="KW-0518">Myosin</keyword>
    <keyword evidence="6 15" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0787">Thick filament</keyword>
    <feature type="domain" description="Myosin N-terminal SH3-like" evidence="18">
        <location>
            <begin position="32"/>
            <end position="81"/>
        </location>
    </feature>
    <feature type="domain" description="Myosin motor" evidence="17">
        <location>
            <begin position="85"/>
            <end position="765"/>
        </location>
    </feature>
    <feature type="region of interest" description="Actin-binding" evidence="15">
        <location>
            <begin position="642"/>
            <end position="664"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1894"/>
            <end position="1929"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1919"/>
            <end position="1929"/>
        </location>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <begin position="178"/>
            <end position="185"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004657"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022433"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023179"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00037488"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00038612"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS51844"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033809.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1929" mass="222443" checksum="A4FD5DABD10BDBBE" modified="2019-05-08" version="1">MGDAHMAEFGAAAPYLRKSDKERLEAQTRIFDLRSDCFVSDEQEEFVKGKIVSRDGERLTVETENGKTVTVKESDVHQQNPPKFDKIEDMAMLTFLHEPAVLYNLKERYAAWMIYTYSGLFCVTVNPYKWLPVYNAEVVAAYRGKKRSEAPPHIFSISDNAYQNMLTNRENQSILITGESGAGKTVNTKRVIQYFASIAAISDRKKEGTSANKGTLEDQIIQANPALEAFGNAKTVRNDNSSRFGKFIRIHFGATGKLASADIETYLLEKSRVIFQLKAERNYHIFYQILSNQKPELLEMMLVTNNPYDYSFISQGEVTVASINDSEELIATDSAFDVLGFTPEEKVGVYKLTGAIMHFGNMKFKQKQREEQAEPDGTEDADKSAYLMGLNSADLLKGLCHPRVKVGNEYVTKGQNVQQVYYSIGALAKSVYEKMFSWMVTRINSTLETKQPRQYFIGVLDIAGFEIFDFNSFEQLCINFTNEKLQQFFNHHMFVLEQEEYKKEGIEWVFIDFGMDLQACIDLIEKPMGIMSILEEECMFPKATDMTFKAKLFDNHLGKSANFSKPRNIKGKPEAHFSLIHYAGTVDYNILGWLQKNKDPLNETVVGLYQKSSLKLLALLFANYAAAESGSYYTPTPSRFLTLALMANLKTTHPHFVRCIIPNERKAPGVMDNPLVMHQLRCNGVLEGIRICRKGFPNRILYGDFRQRYRILNPAAIPEGQFIDSRKGAEKLLSSLDIDPNQYKFGHTKVFFKAGLLGLLEEMRDERLSRIITRIQAQSRGQLMRIEFKKILERRDALLVIQWNIRAFMGVKNWPWMKLYFKIKPLLKSAETEKEMQTMKEEFGRLKEALEKSEARRKELEEKMVSLLQEKNDLQLQVQAEQDNLNDAEERCDQLIKNKIQLEAKVKELTERMEDEEEMNAELTAKKRKLEDECSELKKDIDDLELTLAKVEKEKHATENKVKNLTEEMAGLDEIIAKLTKEKKSLQEAHQQALDDLQVEEDKVNSLTKAKLKLEQQADDLEGSLEQEKKIRMDLERAKRKLEGDLKLTQENIMDLENDKQQLEEKLKKKEFDINQQNSKIEDEQALGLQLQKKLKELQARIEELEEELEAERTARAKVEKLRSDLSRELEEISERLEEAGGATSVQIELNKKREAEFQKMRRDLEEATLQHEATAATLRKKHADSVAELGEQIDNLQRVKQKLEKEKSELKLELDDISSNMEQLLKAKVSLEKLSRTLEDQTAEHRAKLEETQRAMNDTSTQRAKLQTENGELSRQLEEKEALISQLTRGKQSYTQQMEDLKRQLEEETKAKNALAHALQSARHDCDLLREQYEEETEAKAELQRSLSKANSEVAQWRTKYETDAIQRTEELEEAKKKLAQRLQEAEEAVEAVNAKCSSLEKTKHRLQNEIEDLMVDVERSNAAAAALDKKQRNFDKILSEWKQKFEESQTELESSQKEARSLSTELFKLKNAYEESLEHLETFKRENKNLQEEISDLTEQLGEGGKNLHELEKVRKQLEAEKMELQAALEEAEASLEHEEGKILRAQLEFNQFKADFERKLAEKDEEMEQAKRNHLRVVDSLQTSLDAETRSRNEALRVKKKMEGDLNEMEIQLSHANRMAAEAQSHLKATQTQLKDSQLQLDDFIRANEDLKENIAIVERRNSLLQAELEELRAVVEQTERTRRLAEQELIEASERVQLLHTQNTSLINQKKKMEADLSQLQTEVEEAVQECRNAEEKAKKAITDAAMMAEELKKEQDTSAHLERMKKNMEQTIKDLQLRLDEAEQLALKGGKKQLQKLEARVRELENELESEQKRNAESIKGLRKSERRIKELTYQTEEDKKNLMRLQDLVDKLQMKVKAYKRQAEEAEEQANTNLAKFRKVQHELDEAEERADIAESQVNKLRAKSRDAGGKTGLNNVTLTASH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0D2</accession>
    <name>A0A452J0D2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Thrombospondin-like N-terminal domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="12">Belongs to the multiplexin collagen family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038305.1">
        <property type="protein sequence ID" value="ENSGAGP00000033816.1"/>
        <property type="gene ID" value="ENSGAGG00000024061.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005581">
        <property type="term" value="C:collagen trimer"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031012">
        <property type="term" value="C:extracellular matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030020">
        <property type="term" value="F:extracellular matrix structural constituent conferring tensile strength"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030198">
        <property type="term" value="P:extracellular matrix organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00247">
        <property type="entry name" value="Endostatin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000008">
        <property type="entry name" value="collagen alpha-1(XVIII) chain isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.1620.70:FF:000003">
        <property type="entry name" value="Collagen type XVIII alpha 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.200:FF:000039">
        <property type="entry name" value="Collagen XV alpha 1 chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.200">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.1620.70">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.100.10">
        <property type="entry name" value="Mannose-Binding Protein A, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016186">
        <property type="entry name" value="C-type_lectin-like/link_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008160">
        <property type="entry name" value="Collagen"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050149">
        <property type="entry name" value="Collagen_superfamily"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010515">
        <property type="entry name" value="Collagenase_NC10/endostatin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016187">
        <property type="entry name" value="CTDL_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010363">
        <property type="entry name" value="DUF959_COL18_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048287">
        <property type="entry name" value="TSPN-like_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045463">
        <property type="entry name" value="XV/XVIII_trimerization_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24023:SF1112">
        <property type="entry name" value="COL_CUTICLE_N DOMAIN-CONTAINING PROTEIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24023">
        <property type="entry name" value="COLLAGEN ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01391">
        <property type="entry name" value="Collagen"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20010">
        <property type="entry name" value="Collagen_trimer"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06121">
        <property type="entry name" value="DUF959"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06482">
        <property type="entry name" value="Endostatin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00210">
        <property type="entry name" value="TSPN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56436">
        <property type="entry name" value="C-type lectin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="8" id="KW-0176">Collagen</keyword>
    <keyword evidence="9" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11" id="KW-0379">Hydroxylation</keyword>
    <keyword evidence="7" id="KW-0654">Proteoglycan</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <feature type="domain" description="Thrombospondin-like N-terminal" evidence="14">
        <location>
            <begin position="340"/>
            <end position="528"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="250"/>
            <end position="273"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="535"/>
            <end position="1082"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="1108"/>
            <end position="1334"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="1388"/>
            <end position="1470"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="13">
        <location>
            <begin position="616"/>
            <end position="634"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="13">
        <location>
            <begin position="649"/>
            <end position="658"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="13">
        <location>
            <begin position="711"/>
            <end position="721"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="13">
        <location>
            <begin position="726"/>
            <end position="742"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="13">
        <location>
            <begin position="795"/>
            <end position="807"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="13">
        <location>
            <begin position="832"/>
            <end position="841"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="13">
        <location>
            <begin position="876"/>
            <end position="885"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="13">
        <location>
            <begin position="886"/>
            <end position="899"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="13">
        <location>
            <begin position="901"/>
            <end position="919"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="13">
        <location>
            <begin position="1151"/>
            <end position="1160"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="13">
        <location>
            <begin position="1221"/>
            <end position="1239"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="13">
        <location>
            <begin position="1292"/>
            <end position="1305"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="13">
        <location>
            <begin position="1315"/>
            <end position="1327"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="13">
        <location>
            <begin position="1432"/>
            <end position="1448"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="13">
        <location>
            <begin position="1452"/>
            <end position="1463"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022974"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023119"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023278"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00061275"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="SMART" id="SM00210"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033816.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1658" mass="170455" checksum="8064AA9062534C41" modified="2019-05-08" version="1">MASEWVWLRNKWQNASPECETRVQCCWLPGRSCGRVWAKDKTPGVLRSQFPVGVLPCRGRRAGNRRGWLVGSRAWQGLWLATPPFSFVHALRSCRGQTNVSAPGCCLLQDTFPGTVFFMAPAEQRLCLLLLLICCCSLAETRFLDWLWGTQKNPTVTTLPPTESQTTELLTSPVATTTQAAVPATAQPGDQVWSTLSVTRQAPAVETAVPALAKPAMPQGKEENIAGVGAEILDVAEGIRSLVQLWDERTTQRTGNTGDPTAVAPLTNPPTSLAAPNSAMGLADHGNMTANSTGNGHSLLGTGQPEASLPTPAGPLPAWNRTQALLQKPAAALPEHFSTEVSLLQLIGDPPPEQITQVNGPDNTHAYVFGPDANTGQVARYHLPSPFYRDFSLLFHVQPTTNKAGVLFAITDASQTVIYVGVKLSAMQDRKQQIIFYYTEPGSESSYAAATFSVPSLANEWTRFAISVEDDEVILYLNCEEFHRVRFERSPDEMELEDGSGLFVAQAGGADPDKYQGMIADLKVTGDHQAADLQCEEQEEDTDVASGDFGSGAEERPHLSGRKQGIPAVSRLPEPPPVTSPPTAGETVGKVSVGSQLPTDQIKVEGLQQVSTEARVGTKGEKGAPGEKGEHGPKGDSGSGRVLATGSTKGDKGEKGDLGVKGSAGFGYPGSKGQKGEPGAQGSPGPAGLPGPPGSTLQRLDGSVVEQVAGPPGPTGPPGPPGKDGLPGKDGEPGDPGEDGKPGDVGPQGFPGTPGEPGLKGQKGEPGVGARGPPGLPGPPGTPGLSSKQDKLGPRGPPGPPGPPGVPGLPGEPGRFGMNRTELPGPPGLPGLPGRDGVPGAQGPPGPPGKSGLAGQPGLQGERGDPGDLGLPGAPGPKGGKGEMGLAGAPGETGLAGLPGPMGPRGPPGPPGPAGPPGPGYEAGFGDMEGSGMPVLSAVPGERGPKGPQGPPGLPGLKGDAGSPGLPGLPGQKGDHGAPGVDGRPGLEGFPGPQGPKGDKGSQGAKGERGQDGVGSPGPPGPPGQVIYALGEEKALASLPGPEGKEGHAGFPGPMGPKGDLGDPGFPGTPGPKGEKGEPGVIIGPDGTGISAGTKGDKGEQGLVGPMGPAGPHGRPGQKGEIGFPGRPGRPSMNGLKGEKGDPGGLGLQGPPGPPGPPGAPGNVASIYDNNAFSESGPPGPPGLPGYQGAPGQKGERGETGAPGPPGQFPYGLNEFGSTFRGERGDRGDPGLKGEKGEPGEGGLYGPSVSGPPGPQGYPGLPGPKGDSIRGLPGPPGPQGPPGIGYEGRQGPPGPPGPPGPPSFPGPHRQTVSIPGPPGPPGPPGPPGTSGSLTSYGLRILPAYQSMLSTAQEVPEGWLIFVQDREELYIRVRSGFRRVRLEEHTSISSPGLDNEVYDKPPSVHYSHGGTASSGFLHPRREHNPSSTAQPWRADDSIANHHRLPDHLPHGAQPQQEPLDSFSPNRRGESIPSAVHRHHDFQPALHLVALNAPLSGSMRGIRGADFQCFQQARAVGLTGTFRAFLSSRLQDLYSIVRRADRSGMPIVNLRDEVLFNNWEALFSGAEAQFRAGVRILSFDGRDVLRDSAWPQKNVWHGSDAKGHRLTESYCETWRTDDSVVTGQASSLASGKLLEQKVSSCRNAFIVLCIENSFMTSSKK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0D5</accession>
    <name>A0A452J0D5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Eukaryotic translation initiation factor 4E binding protein 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the eIF4E-binding protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0D5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038259.1">
        <property type="protein sequence ID" value="ENSGAGP00000033778.1"/>
        <property type="gene ID" value="ENSGAGG00000024040.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008190">
        <property type="term" value="F:eukaryotic initiation factor 4E binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045947">
        <property type="term" value="P:negative regulation of translational initiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008606">
        <property type="entry name" value="EIF4EBP"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12669">
        <property type="entry name" value="EUKARYOTIC TRANSLATION INITIATION FACTOR 4E-BINDING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12669:SF14">
        <property type="entry name" value="EUKARYOTIC TRANSLATION INITIATION FACTOR 4E-BINDING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05456">
        <property type="entry name" value="eIF_4EBP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0652">Protein synthesis inhibitor</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0810">Translation regulation</keyword>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="1"/>
            <end position="46"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="1"/>
            <end position="10"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="35"/>
            <end position="46"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00005480"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022845"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023193"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033778.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="7">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="114" mass="12282" checksum="1A36DFC77808AF0F" modified="2019-05-08" version="1">MSGSCRQGHTLSRAIPAKRLSLPDGAPLPPGDYSTTPGGTVFGTTPGGTRIIYDRKFLMECRNSPVAKTSPCDLPDIPGVTSPNVEELKIENNHVHNYLEKVGIGEEAQFDMDI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0D6</accession>
    <name>A0A452J0D6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Integrin alpha-V</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="19">Vitronectin receptor subunit alpha</fullName>
        </alternativeName>
    </protein>
    <organism evidence="26 27">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="27" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="26" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="26" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell junction</location>
            <location evidence="1">Focal adhesion</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2 21">Membrane</location>
            <topology evidence="2 21">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 21">Belongs to the integrin alpha chain family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0D6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038303.1">
        <property type="protein sequence ID" value="ENSGAGP00000033814.1"/>
        <property type="gene ID" value="ENSGAGG00000023825.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009897">
        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005925">
        <property type="term" value="C:focal adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008305">
        <property type="term" value="C:integrin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001525">
        <property type="term" value="P:angiogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033627">
        <property type="term" value="P:cell adhesion mediated by integrin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098609">
        <property type="term" value="P:cell-cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007160">
        <property type="term" value="P:cell-matrix adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007229">
        <property type="term" value="P:integrin-mediated signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.130:FF:000003">
        <property type="entry name" value="Integrin alpha V"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1510:FF:000001">
        <property type="entry name" value="Integrin alpha V"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1530:FF:000002">
        <property type="entry name" value="integrin alpha-V isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.930:FF:000001">
        <property type="entry name" value="Integrin subunit alpha V"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1460:FF:000001">
        <property type="entry name" value="Integrin, alpha V"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.930">
        <property type="entry name" value="Bicelle-embedded integrin alpha(iib) transmembrane segment"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.130">
        <property type="entry name" value="Integrin alpha, N-terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1460">
        <property type="entry name" value="Integrin domains. Chain A, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1510">
        <property type="entry name" value="ntegrin, alpha v. Chain A, domain 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1530">
        <property type="entry name" value="ntegrin, alpha v. Chain A, domain 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013517">
        <property type="entry name" value="FG-GAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013519">
        <property type="entry name" value="Int_alpha_beta-p"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000413">
        <property type="entry name" value="Integrin_alpha"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018184">
        <property type="entry name" value="Integrin_alpha_C_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013649">
        <property type="entry name" value="Integrin_alpha_Ig-like_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048285">
        <property type="entry name" value="Integrin_alpha_Ig-like_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048286">
        <property type="entry name" value="Integrin_alpha_Ig-like_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028994">
        <property type="entry name" value="Integrin_alpha_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032695">
        <property type="entry name" value="Integrin_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23220">
        <property type="entry name" value="INTEGRIN ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23220:SF4">
        <property type="entry name" value="INTEGRIN ALPHA-V"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01839">
        <property type="entry name" value="FG-GAP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08441">
        <property type="entry name" value="Integrin_A_Ig_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20805">
        <property type="entry name" value="Integrin_A_Ig_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20806">
        <property type="entry name" value="Integrin_A_Ig_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00357">
        <property type="entry name" value="Integrin_alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01185">
        <property type="entry name" value="INTEGRINA"/>
    </dbReference>
    <dbReference type="SMART" id="SM00191">
        <property type="entry name" value="Int_alpha"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69318">
        <property type="entry name" value="Integrin alpha N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Integrin domains"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51470">
        <property type="entry name" value="FG_GAP"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00242">
        <property type="entry name" value="INTEGRIN_ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0106">Calcium</keyword>
    <keyword evidence="10 21" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="11" id="KW-0965">Cell junction</keyword>
    <keyword evidence="4" id="KW-0165">Cleavage on pair of basic residues</keyword>
    <keyword evidence="15" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="17" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13 21" id="KW-0401">Integrin</keyword>
    <keyword evidence="14 21" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="16 21" id="KW-0675">Receptor</keyword>
    <keyword evidence="27" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="7 21" id="KW-0732">Signal</keyword>
    <keyword evidence="5 21" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="12 21" id="KW-1133">Transmembrane helix</keyword>
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        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
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        <location>
            <begin position="21"/>
            <end position="1029"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="975"/>
            <end position="997"/>
        </location>
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        <location>
            <begin position="22"/>
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        </location>
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        <location>
            <begin position="98"/>
            <end position="159"/>
        </location>
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        <location>
            <begin position="222"/>
            <end position="276"/>
        </location>
    </feature>
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        <location>
            <begin position="277"/>
            <end position="342"/>
        </location>
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            <begin position="343"/>
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        </location>
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        <location>
            <begin position="404"/>
            <end position="467"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha first immunoglubulin-like" evidence="23">
        <location>
            <begin position="452"/>
            <end position="610"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha second immunoglobulin-like" evidence="24">
        <location>
            <begin position="611"/>
            <end position="739"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha third immunoglobulin-like" evidence="25">
        <location>
            <begin position="757"/>
            <end position="964"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="1008"/>
            <end position="1029"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="22">
        <location>
            <begin position="1008"/>
            <end position="1023"/>
        </location>
    </feature>
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        <source>
            <dbReference type="ARBA" id="ARBA00004246"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00008054"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022685"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
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        </source>
    </evidence>
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    </evidence>
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    </evidence>
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        </source>
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    </evidence>
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        <source>
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    </evidence>
    <sequence length="1029" mass="113945" checksum="47BCACC98BA00872" modified="2019-05-08" version="1">MARLRLLLLLLGRWLALGGAFNLDADGPAVYSGSEGSYFGFAVDFFAPDSSSIFLLVGAPKANTTQPDIVEGGQVLKCNWNSNKNCQPVIFDSTGDRNFAPNDPLEFKSHQWFGASVRSKNDKILACAPLYHWRTEVKQEREPVGTCYLHDGSKAVEYAPCRSTIIDADGQGFCQGGFSIDFTKQLSILMGFFPAGQLISDRVAEIVSKYNSKVYSIKYDDQLATRSASAAFDDSYLGYSVAVGEFSGDGIEDFVSGVPRAARTLGMVSIYNGKNMSSLYNFTGEQMAAYFGYSVAATDINGDNYTDLFIGAPLFMERGSDGKLQEVGQVSICLQQASGGFQIIKLNGFEIFARFSSSIAPLGDLDQDGFNDIAVAAPYGGEDKRGVVYIYNGRATGLNAVPSQILEGQWAARTMPPSFGYSLKGATDVDKNGYPDLIVGAFGVDTAVLYRARPVISVNAGLEVNPTILNPEEKICELPDTRVKVSCFNVKFCLTADGKGKLPNTLTFQVELLLDKLKQKGALRRALFLHNKQPGHSKNMTVVKGGTMRCEELQAFLRDESEFRDKLTPITVFMEYRLDYKTAKDSAGLQPILNQFTPANISRQTHILLDCGEDNICKPKLQVAVHSDQKQIYIGDDNPLTLIVDAQNQGEGAYEAELFVIIPPQADFIGVVRNNEALARLSCAFKTENQTRLVVCDLGNPMKAGTKLLAGLRFSVNQQSEMDTSVKFDLQIRSSNLFDNLSPTTSYQADLAILASVEIRGVSSPDHIFLPIANWQPQENPETEEDIGPLVQHIYELRNNGPSAFSKVMLNLQWPYKYNNNTLLYIMQYEIDGPMNCTSDMEINPLNIKVTDEKNETIRDEDNRDHHINRRDITAIEGDVHTLGCGTAECLKIVCQVGRLERGKSAILYVKSRLWTQTFMNKENQNHSYSLKSSASFNVIEFPYKNLSFEDIHNSTVVTTNITWGIQPQPMPVPVWVIILAVLAGLLLLAVLVFVMYRMGFFKRVRPPQEEQEREQLQPHENGEGTSEA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0E0</accession>
    <name>A0A452J0E0_9SAUR</name>
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        <submittedName>
            <fullName evidence="7">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
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            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J0E0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038308.1">
        <property type="protein sequence ID" value="ENSGAGP00000033819.1"/>
        <property type="gene ID" value="ENSGAGG00000024064.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
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        <property type="project" value="UniProtKB-KW"/>
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    <keyword evidence="2" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="3" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022485"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023014"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033819.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="157" mass="17743" checksum="1BFDC3E07860B86D" modified="2019-05-08" version="1">LKHCFYPQVHLSGSGWSPVYAEENLSVMSVGFLLSGPDDAVIWRRPQKMVCRMIKQFLCDVQWGEIDYLIVDTPSGTSDEHLSIVEVLLQIGFVCPKCKVRLSKNILTHMMCQNLNISFLGKVPLDPQIGKVGKLHNKKQQCACHYFRVIFGISAMC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0E2</accession>
    <name>A0A452J0E2_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="21">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 16">Membrane</location>
            <topology evidence="1 16">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 16">Belongs to the integrin alpha chain family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0E2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038315.1">
        <property type="protein sequence ID" value="ENSGAGP00000033826.1"/>
        <property type="gene ID" value="ENSGAGG00000023825.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009897">
        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008305">
        <property type="term" value="C:integrin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001525">
        <property type="term" value="P:angiogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033627">
        <property type="term" value="P:cell adhesion mediated by integrin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="project" value="TreeGrafter"/>
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        <property type="term" value="P:integrin-mediated signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
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        <property type="entry name" value="Integrin alpha V"/>
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        <property type="entry name" value="integrin alpha-V isoform X2"/>
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        <property type="entry name" value="Integrin subunit alpha V"/>
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        <property type="entry name" value="Bicelle-embedded integrin alpha(iib) transmembrane segment"/>
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        <property type="entry name" value="Integrin alpha, N-terminal"/>
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        <property type="entry name" value="Integrin domains. Chain A, domain 2"/>
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        <property type="entry name" value="ntegrin, alpha v. Chain A, domain 3"/>
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        <property type="entry name" value="ntegrin, alpha v. Chain A, domain 4"/>
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        <property type="entry name" value="Int_alpha_beta-p"/>
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        <property type="entry name" value="Integrin_alpha_C_CS"/>
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        <property type="entry name" value="Integrin_alpha_Ig-like_1"/>
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        <property type="entry name" value="Integrin_alpha_Ig-like_2"/>
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        <property type="entry name" value="Integrin_alpha_N"/>
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        <property type="entry name" value="Integrin_dom_sf"/>
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        <property type="entry name" value="INTEGRIN ALPHA-V"/>
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        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
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            <begin position="21"/>
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            <begin position="910"/>
            <end position="932"/>
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        </location>
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    </evidence>
    <sequence length="964" mass="107086" checksum="06C9614458D6A629" modified="2019-05-08" version="1">MARLRLLLLLLGRWLALGGAFNLDADGPAVYSGSEGSYFGFAVDFFAPDSSSIFLLVGAPKANTTQPDIVEGGQVLKCNWNSNKNCQPVIFDSTGDRNFAPNDPLEFKSHQWFGASVRSKNDKILACAPLYHWRTEVKQEREPVGTCYLHDGSKAVEYAPCRSTIIDADGQGFCQGGFSIDFTKQLSILMGFFPAGQLISDRVAEIVSKYNSKVYSIKYDDQLATRSASAAFDDSYLGYSVAVGEFSGDGIEDFVSGVPRAARTLGMVSIYNGKNMSSLYNFTGEQMAAYFGYSVAATDINGDNYTDLFIGAPLFMERGSDGKLQEVGQVSICLQQASGGFQIIKLNGFEIFARFSSSIAPLGDLDQDGFNDLIVGAFGVDTAVLYRARPVISVNAGLEVNPTILNPEEKICELPDTRVKVSCFNVKFCLTADGKGKLPNTLTFQVELLLDKLKQKGALRRALFLHNKQPGHSKNMTVVKGGTMRCEELQAFLRDESEFRDKLTPITVFMEYRLDYKTAKDSAGLQPILNQFTPANISRQTHILLDCGEDNICKPKLQVAVHSDQKQIYIGDDNPLTLIVDAQNQGEGAYEAELFVIIPPQADFIGVVRNNEALARLSCAFKTENQTRLVVCDLGNPMKAGTKLLAGLRFSVNQQSEMDTSVKFDLQIRSSNLFDNLSPTTSYQADLAILASVEIRGVSSPDHIFLPIANWQPQENPETEEDIGPLVQHIYELRNNGPSAFSKVMLNLQWPYKYNNNTLLYIMQYEIDGPMNCTSDMEINPLNIKVNEKNETIRDEDNRDHHINRRDITAIEGDVHTLGCGTAECLKIVCQVGRLERGKSAILYVKSRLWTQTFMNKENQNHSYSLKSSASFNVIEFPYKNLSFEDIHNSTVVTTNITWGIQPQPMPVPVWVIILAVLAGLLLLAVLVFVMYRMGFFKRVRPPQEEQEREQLQPHENGEGTSEA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0E3</accession>
    <name>A0A452J0E3_9SAUR</name>
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        <submittedName>
            <fullName evidence="5">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
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        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J0E3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038351.1">
        <property type="protein sequence ID" value="ENSGAGP00000033860.1"/>
        <property type="gene ID" value="ENSGAGG00000024085.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="term" value="C:endoplasmic reticulum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000137">
        <property type="term" value="C:Golgi cis cisterna"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="F:calcium ion binding"/>
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        <property type="project" value="InterPro"/>
    </dbReference>
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        <property type="term" value="P:regulation of amyloid precursor protein biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="ABM_dom"/>
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        <property type="entry name" value="Dimeric_a/b-barrel"/>
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        <property type="entry name" value="EF-hand-dom_pair"/>
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        <property type="entry name" value="EF_Hand_1_Ca_BS"/>
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        <property type="entry name" value="EF_hand_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039862">
        <property type="entry name" value="NECAB1/2/3"/>
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    <dbReference type="PANTHER" id="PTHR12178">
        <property type="entry name" value="EF-HAND DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
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    <dbReference type="PANTHER" id="PTHR12178:SF3">
        <property type="entry name" value="N-TERMINAL EF-HAND CALCIUM-BINDING PROTEIN 3"/>
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        <property type="entry name" value="Dimeric alpha+beta barrel"/>
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    </dbReference>
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    </dbReference>
    <proteinExistence type="predicted"/>
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    <keyword evidence="1" id="KW-0479">Metal-binding</keyword>
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    <feature type="domain" description="EF-hand" evidence="3">
        <location>
            <begin position="24"/>
            <end position="52"/>
        </location>
    </feature>
    <feature type="domain" description="ABM" evidence="4">
        <location>
            <begin position="244"/>
            <end position="332"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
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            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS50222"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS51725"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033860.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="344" mass="39418" checksum="4F02553D9353B73D" modified="2019-05-08" version="1">MAGPRRRSFAEAIGVGCIWSKTFQVFRRADKNDDGKLSFEEFKNYFADGILSSEELWELFSGIDRHHSDNLETEKLCDYFSEHLGEYRHVLSALEALNTAVLAAMDKTKLEYESSSKMEQFVTRFLLRETRTQLQTLQASLECASDTIEEQAGWERIDLKKSEVFIGPRSGRRCGRRAQKTVCVSPTDPYSGMLSTGVCVEADSQWVAQINRLQQLIDKLECRSPRLEPLKEEIVCKGRNAHILVAQRQLSVTESGLEGFHQALQSYTEVTARQSHCLHVSSRKLTSEACFILYEFWQDVTSWRSHLQSSCSKTFQWAIIDFLESPELVTTMLFPASWWIVNNN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0E4</accession>
    <name>A0A452J0E4_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="19">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">Muscle contraction.</text>
    </comment>
    <comment type="subunit">
        <text evidence="14">Muscle myosin is a hexameric protein that consists of 2 heavy chain subunits (MHC), 2 alkali light chain subunits (MLC) and 2 regulatory light chain subunits (MLC-2).</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Myofibril</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 15">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038316.1">
        <property type="protein sequence ID" value="ENSGAGP00000033827.1"/>
        <property type="gene ID" value="ENSGAGG00000020256.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030016">
        <property type="term" value="C:myofibril"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032982">
        <property type="term" value="C:myosin filament"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016460">
        <property type="term" value="C:myosin II complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000146">
        <property type="term" value="F:microfilament motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd01377">
        <property type="entry name" value="MYSc_class_II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000003">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000004">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000006">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000013">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000003">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000007">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000008">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.4820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.530:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.360:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000024">
        <property type="entry name" value="Myosin heavy chain, isoform J"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.720:FF:000001">
        <property type="entry name" value="Myosin heavy chain, muscle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.340">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.370">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.4820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.2420">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.360">
        <property type="entry name" value="Myosin S1 fragment, N-terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008989">
        <property type="entry name" value="Myosin_S1_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002928">
        <property type="entry name" value="Myosin_tail"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004009">
        <property type="entry name" value="SH3_Myosin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014751">
        <property type="entry name" value="XRCC4-like_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615">
        <property type="entry name" value="MYOSIN HEAVY CHAIN, NON-MUSCLE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615:SF69">
        <property type="entry name" value="MYOSIN-6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02736">
        <property type="entry name" value="Myosin_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01576">
        <property type="entry name" value="Myosin_tail_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90257">
        <property type="entry name" value="Myosin rod fragments"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57997">
        <property type="entry name" value="Tropomyosin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51844">
        <property type="entry name" value="SH3_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12 15" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="7 15" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="8" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0488">Methylation</keyword>
    <keyword evidence="10 15" id="KW-0505">Motor protein</keyword>
    <keyword evidence="11" id="KW-0514">Muscle protein</keyword>
    <keyword evidence="9 15" id="KW-0518">Myosin</keyword>
    <keyword evidence="6 15" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0787">Thick filament</keyword>
    <feature type="domain" description="Myosin N-terminal SH3-like" evidence="18">
        <location>
            <begin position="32"/>
            <end position="81"/>
        </location>
    </feature>
    <feature type="domain" description="Myosin motor" evidence="17">
        <location>
            <begin position="85"/>
            <end position="778"/>
        </location>
    </feature>
    <feature type="region of interest" description="Actin-binding" evidence="15">
        <location>
            <begin position="655"/>
            <end position="677"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1906"/>
            <end position="1936"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="16">
        <location>
            <begin position="1923"/>
            <end position="1936"/>
        </location>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <begin position="178"/>
            <end position="185"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004657"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022433"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023179"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00037488"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00038612"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS51844"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033827.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1936" mass="223125" checksum="F27CD925071C980E" modified="2019-05-08" version="1">MGDAHMAEFGAAAPYLRKSDKERLEAQTRIFDLRSDCFVSDEQEEFVKGKIVSRDGERLTVETENGKTVTVKESDVHQQNPPKFDKIEDMAMLTFLHEPAVLYNLKERYAAWMIYTYSGLFCVTVNPYKWLPVYNAEVVAAYRGKKRSEAPPHIFSISDNAYQNMLTNRENQSILITGESGAGKTVNTKRVIQYFASIAAINPPCCELSSPSGTLEDQIIQANPALEAFGNAKTVRNDNSSRFGKFIRIHFGATGKLASADIETYLLEKSRVIFQLKAERNYHIFYQILSNQKPELLEMMLVTNNPYDYSFISQGEVTVASINDSEELIATDSAFDVLGFTPEEKVGVYKLTGAIMHFGNMKFKQKQREEQAEPDGTEDADKSAYLMGLNSADLLKGLCHPRVKVGNEYVTKGQNVQQVYYSIGALAKSVYEKMFSWMVTRINSTLETKQPRQYFIGVLDIAGFEIFDFNSFEQLCINFTNEKLQQFFNHHMFVLEQEEYKKEGIEWVFIDFGMDLQACIDLIEKPMGIMSILEEECMFPKATDMTFKAKLFDNHLGKSANFSKPRNIKGKPEAHFSLIHYAGTVDYNILGWLQKNKDPLNETVVGLYQKSSLKLLALLFANYAAAESGDGGKGKGAKKKGSSFQTVSALHRENLNKLMANLKTTHPHFVRCIIPNERKAPGVMDNPLVMHQLRCNGVLEGIRICRKGFPNRILYGDFRQRYRILNPAAIPEGQFIDSRKGAEKLLSSLDIDPNQYKFGHTKVFFKAGLLGLLEEMRDERLSRIITRIQAQSRGQLMRIEFKKILERRDALLVIQWNIRAFMGVKNWPWMKLYFKIKPLLKSAETEKEMQTMKEEFGRLKEALEKSEARRKELEEKMVSLLQEKNDLQLQVQAEQDNLNDAEERCDQLIKNKIQLEAKVKELTERMEDEEEMNAELTAKKRKLEDECSELKKDIDDLELTLAKVEKEKHATENKVKNLTEEMAGLDEIIAKLTKEKKSLQEAHQQALDDLQVEEDKVNSLTKAKLKLEQQADDLEGSLEQEKKIRMDLERAKRKLEGDLKLTQENIMDLENDKQQLEEKLKKKEFDINQQNSKIEDEQALGLQLQKKLKELQARIEELEEELEAERTARAKVEKLRSDLSRELEEISERLEEAGGATSVQIELNKKREAEFQKMRRDLEEATLQHEATAATLRKKHADSVAELGEQIDNLQRVKQKLEKEKSELKLELDDISSNMEQLLKAKVSLEKLSRTLEDQTAEHRAKLEETQRAMNDTSTQRAKLQTENGELSRQLEEKEALISQLTRGKQSYTQQMEDLKRQLEEETKAKNALAHALQSARHDCDLLREQYEEETEAKAELQRSLSKANSEVAQWRTKYETDAIQRTEELEEAKKKLAQRLQEAEEAVEAVNAKCSSLEKTKHRLQNEIEDLMVDVERSNAAAAALDKKQRNFDKILSEWKQKFEESQTELESSQKEARSLSTELFKLKNAYEESLEHLETFKRENKNLQEEISDLTEQLGEGGKNLHELEKVRKQLEAEKMELQAALEEAEASLEHEEGKILRAQLEFNQFKADFERKLAEKDEEMEQAKRNHLRVVDSLQTSLDAETRSRNEALRVKKKMEGDLNEMEIQLSHANRMAAEAQSHLKATQTQLKDSQLQLDDFIRANEDLKENIAIVERRNSLLQAELEELRAVVEQTERTRRLAEQELIEASERVQLLHTQNTSLINQKKKMEADLSQLQTEVEEAVQECRNAEEKAKKAITDAAMMAEELKKEQDTSAHLERMKKNMEQTIKDLQLRLDEAEQLALKGGKKQLQKLEARVRELENELESEQKRNAESIKGLRKSERRIKELTYQTEEDKKNLMRLQDLVDKLQMKVKAYKRQAEEAEEQANTNLAKFRKVQHELDEAEERADIAESQVNKLRAKSRDAGGKKLHDEE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0E7</accession>
    <name>A0A452J0E7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Tektin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">Sperm-specific microtubule inner protein (MIP) part of the dynein-decorated doublet microtubules (DMTs) in flagellar axoneme. Forms an extensive interaction network in different conformations that reinforces the helix bundle composed by other tektin proteins (TEKT1 to TEKT4) and MIPs to anchor the tektin bundle onto the tubulin wall of A-tubule of the sperm flagellum.</text>
    </comment>
    <comment type="subunit">
        <text evidence="11">Microtubule inner protein component of sperm flagellar doublet microtubules. Interacts with TEKT3.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="12">Cytoplasm</location>
            <location evidence="12">Cytoskeleton</location>
            <location evidence="12">Cilium axoneme</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Flagellum axoneme</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 12">Belongs to the tektin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0E7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038356.1">
        <property type="protein sequence ID" value="ENSGAGP00000033865.1"/>
        <property type="gene ID" value="ENSGAGG00000024073.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005930">
        <property type="term" value="C:axoneme"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015630">
        <property type="term" value="C:microtubule cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036126">
        <property type="term" value="C:sperm flagellum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060271">
        <property type="term" value="P:cilium assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060294">
        <property type="term" value="P:cilium movement involved in cell motility"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048256">
        <property type="entry name" value="Tektin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000435">
        <property type="entry name" value="Tektins"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19960">
        <property type="entry name" value="TEKTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19960:SF23">
        <property type="entry name" value="TEKTIN-5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03148">
        <property type="entry name" value="Tektin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00511">
        <property type="entry name" value="TEKTIN"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9 12" id="KW-0966">Cell projection</keyword>
    <keyword evidence="7 12" id="KW-0969">Cilium</keyword>
    <keyword evidence="6" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="5 12" id="KW-0282">Flagellum</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0832">Ubl conjugation</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004611"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007209"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022846"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023069"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00046061"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00047089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="RuleBase" id="RU367040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033865.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="459" mass="52989" checksum="8749BC81539E1648" modified="2019-05-08" version="1">MEFLGTTQTASYCGPKKSYLFPEPVSTSTTQDSYQSYYLPGYRYLNSWRPSLFYRVSAVPTNSGDQGSMRISAPLRPPTILPSVRSALYARYSPHDWYQANTVQYKGSESCRHWAGRMNADSLRLMQDKNQLTQQTQDSTGRSLGQRLSDIDFWRSELTYELERLLTETRALETAKRRLECAADEAQRPLKVGQRYLILNVLVCLRVRLYYLRSAAPEDAQHALERDLSDKNSAHFIDEKCYNLRNTSDSISFYHGVEKIDGTISVPETWAKFSDDNIRHSQNARANSIKLCEETENTLENVSEEMWKQFTDTNLAFTNRIAETADTKNKLQTHLAKTLQEIFQTENTIMLLERAIMAKECPLKVAQTRLEGRTRRPNVELCRDIPQFKLVNEVFTIDDTLQTLKLRLREAHDTLQLLVLNKARLEHDLSVKANSLFIDRKCMDMRKTFPSTPRLVGYT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0E8</accession>
    <name>A0A452J0E8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Set1/Ash2 histone methyltransferase complex subunit ASH2</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="14">ASH2-like protein</fullName>
        </alternativeName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="12">Transcriptional regulator. Component or associated component of some histone methyltransferase complexes which regulates transcription through recruitment of those complexes to gene promoters. Component of the Set1/Ash2 histone methyltransferase (HMT) complex, a complex that specifically methylates 'Lys-4' of histone H3, but not if the neighboring 'Lys-9' residue is already methylated. As part of the MLL1/MLL complex it is involved in methylation and dimethylation at 'Lys-4' of histone H3. May play a role in hematopoiesis. In association with RBBP5 and WDR5, stimulates the histone methyltransferase activities of KMT2A, KMT2B, KMT2C, KMT2D, SETD1A and SETD1B.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0E8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038270.1">
        <property type="protein sequence ID" value="ENSGAGP00000033788.1"/>
        <property type="gene ID" value="ENSGAGG00000024046.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071339">
        <property type="term" value="C:MLL1 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048188">
        <property type="term" value="C:Set1C/COMPASS complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000976">
        <property type="term" value="F:transcription cis-regulatory region binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006325">
        <property type="term" value="P:chromatin organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010557">
        <property type="term" value="P:positive regulation of macromolecule biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010468">
        <property type="term" value="P:regulation of gene expression"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd15583">
        <property type="entry name" value="PHD_ash2p_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12872">
        <property type="entry name" value="SPRY_Ash2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.980.20:FF:000002">
        <property type="entry name" value="Ash2 like, histone lysine methyltransferase complex subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.920:FF:000008">
        <property type="entry name" value="Set1/Ash2 histone methyltransferase complex subunit ASH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.920">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.980.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037353">
        <property type="entry name" value="ASH2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049455">
        <property type="entry name" value="ASH2-like_PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR053835">
        <property type="entry name" value="ASH2L-like_WH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001870">
        <property type="entry name" value="B30.2/SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043136">
        <property type="entry name" value="B30.2/SPRY_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003877">
        <property type="entry name" value="SPRY_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10598">
        <property type="entry name" value="SET1/ASH2 HISTONE METHYLTRANSFERASE COMPLEX SUBUNIT ASH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10598:SF0">
        <property type="entry name" value="SET1_ASH2 HISTONE METHYLTRANSFERASE COMPLEX SUBUNIT ASH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21198">
        <property type="entry name" value="ASH2L-like_WH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21257">
        <property type="entry name" value="PHD_ash2p_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00622">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00449">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50188">
        <property type="entry name" value="B302_SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0156">Chromatin regulator</keyword>
    <keyword evidence="9" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="2" id="KW-0488">Methylation</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="16" id="KW-0732">Signal</keyword>
    <keyword evidence="10" id="KW-0804">Transcription</keyword>
    <keyword evidence="8" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5" id="KW-0863">Zinc-finger</keyword>
    <feature type="signal peptide" evidence="16">
        <location>
            <begin position="1"/>
            <end position="32"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019399349" description="Set1/Ash2 histone methyltransferase complex subunit ASH2" evidence="16">
        <location>
            <begin position="33"/>
            <end position="651"/>
        </location>
    </feature>
    <feature type="domain" description="B30.2/SPRY" evidence="17">
        <location>
            <begin position="383"/>
            <end position="606"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="48"/>
            <end position="117"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="282"/>
            <end position="354"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="15">
        <location>
            <begin position="57"/>
            <end position="79"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="15">
        <location>
            <begin position="300"/>
            <end position="318"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00056054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00068996"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00075872"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50188"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033788.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="651" mass="72616" checksum="B35810A4D4DBCEE3" modified="2019-05-08" version="1">MDPNTRETLVSLVNWTFVTVCITAALQGPCHDEECHCTEEDTNIKKRTVQAPSTLQPEKEQEASGQHGIDKADRNKNARGDTTLVDVTAGLETESSNGKDPLEGVGDSSEVLDTQTGSVDEENGRQLGEIELQCGICTKWFTADTFGIDTTSCLPFMTNYSFHCNVCHHSGNTYFLRKQANLKEMCLSALANLTWQSRTQDEHPKTMFSKDKDIIPFIDKYWECMTTRQRPGKMTWPNNIVKTMCKERDVFLVKEHPDPGSKDPEDDYPKFGLLDQDLANIGPAYDNQKQSNAVSTSGSLNGGASLGGGIPAGSSGKGRGAKRKQQDGGTTGTAKKTRSDPLFSAQRLPPHGYPLEHPFNKDGYRYILAEPDPHAPDPEKLELDCWAGKPIPGDLYRACLYERVLLALHDRAPQLKISDDRLTVIGEKGYSMVRASHGVRKGDWYFEISVDEMPPDTAARLGWSQPLGNLQAPLGYDKFSYSWRSKKGTKFHQSIGKHYSAGYGQGDVLGFYISLPEDTETAKSLPDTYKDKALIKFKSYLYFEEKDFVDKAEKSLKQTPGSQIIFYKNGVSQGVAFKEIFEGVYFPAISLYKGCTVSVNFGPYFKYPPRNITYRPMSDMGWGAVVEHTLADVLYHVETEVDGRRSPPWEP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0E9</accession>
    <name>A0A452J0E9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Calponin-homology (CH) domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the Nav/unc-53 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038321.1">
        <property type="protein sequence ID" value="ENSGAGP00000033832.1"/>
        <property type="gene ID" value="ENSGAGG00000024067.1"/>
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        <property type="entry name" value="CortBP2_NAV1-like_AAA_lid"/>
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        <property type="entry name" value="Nav/unc-53"/>
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        <property type="entry name" value="NEURON NAVIGATOR 3"/>
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        <property type="entry name" value="Ubiquitin_6"/>
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            <begin position="1"/>
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    <sequence length="2183" mass="234220" checksum="BCCF19928A8428FC" modified="2019-05-08" version="1">MIENVDVCLSFLAARGVNVQGLSAEEIRNGNLKAILGLFFSLSRYKQQQHHQQQYYQSLVELQQHVSHSSPPAEISQSKTQQDMQSRLPGPSRVPAAGSSTKVQGASNLNRRSQSFNSIDKNKPPQYANSNEKDSPKGPHPSAGMNGNVQHPISTGQQPVSAIPSPSASKPWRSKSMNVKHSATSTMLSVKQPSPATSPTPSSDRLKPPLSEGVKPPSSGQKSMLEKFKLVNARTALRPPLSSSSGPNDSGREDDTFSECGEMDGFSSGLNSGGSTSSSPKVSPKLTPPKAGSKNLSNKKSLLQPKDKEEKNRDKNKVCTEKTVKEEKDQVPETSTKKSSKIASLIPKGSKTTAAKKESLIPSSSGIPKPGSKVPTAKQSASPVCAGSKESEKFRTTKGNQSQSTTKSQANEKASTSCNLASSEGKESSPALTPGSSAALSVSMAASSGQVTGNGIVQLPQQQQYSHPNTATVAPFIYRTHSENDSTSLPPADSCTSPTKMDLSYSKTAKQCLEEISGEDPETRRMRTVKNIADLRQNLEETMSSLRGTQISHSTLETTFDSTVTTEVNGRSIPGLTSRSTPVTWRLGQASPRLQAGDAPSLGSGYPRSSAGRFIHTDPSRFMYTTPLRRAAVSRLGNMSQIDMNEKGSSDLDMSSEVDVGGYMSDGDILGKSLRTDDISSGYMTDGGLNLYTRSLNRIPDLAASRDVIQRGVHDVTVDADSWDDSSSVSSGLSDTLDNLSTDDLNTTSSVSSYSNITASSRKNTHTQLKTDSEKRFVTDSENWGSTEELKKPEEDFDSNVDGSGKWKGLPSGLSEDSEKGGQKASLSVSQTGSWRRGMTAQVGTTSSRHKAGTSALKTPGKTDDAKASEKVKTPLKGTSIQRSPSDAGKSSGDEGKKPPSGIGRSTATGSFGFKKAGVGSSTIITTSGATITSGSATLGKMPKSTAIGGKSNAGRKTSLDGSQNQDDGVLHVNSKTTLQYRSLPRPSKSSTSGIPGRGGHRSSTSSIDSNLSSKSAGAATTKLREPTKIGSGRSSPITINQTDKEKEKVAVSDSESVSLSSSPKSSPTSASACGTPGLRQPGSKYPDIASPTFRRLFGAKAGGKAASAPNTESVKPTSAMPSPSTTLARQGSLESPSSGTGSMGSAGGQSGSSSPLFSKPSELNTDVVSLSHSLASSPASVHSFTSGGLVWAANLSSSSAGSKDTPSYQSMTSLHTSSESIDLPLSHHSSLSGLTTGTQEVQSLLMRTGSVRSTLSESMQLDRNTLPKKGLRYTPSSRQTSQEESKEWLRSHSTGGLQDTGSQSPIVSPSAMSSSATGKYHFSNLVSPTNLSQFNLPGPSMMRSNSIPAQDSSFDLYDDSQLCGSATSLEERPRAISHSGSFRDSMEEVHGSSLSLVSSTSSLYSTAEEKAHSEQIHKLRRELVASQEKVATLTSQLSANAHLVAAFEKSLGNMTGRLQSLTMTAEQKESELIELRETIEMLKAQNSAAQAAIQGALNGPDHAHKDLLIRRQHSSESVSSINSATSHSSIGSGNDADSKKKKKKNWLRSSFKQAFGKKKSTKPPSSHSDIEELTDSSLPSSPKLPHNSGDCGTLSMKPSQSASAICECTEAEAEIILQLKSELREKELKLTDIRLEALSSAHHLDQIREAMNRMQNEIEILKAENDRLKAETGNTGKPARPSSESSSSTSSSSSRQSLGLSLNNLNITDSVTSDILLDDVTDGALPKEGHSVKIIVTISKGYGRSKDQKTQAYLIGSIGVSGKTKWDVLDGVIRRLFKEYIFRVDPATSLGLSSDCIASYCIGDLIRVHSLEVPELLPCGYLVGDNNIITVNLKGVEENSLDSFVFDTLIPKPITQRYFNLLMEHHRIILSGPSGTGKTYLANRLAEYIITKLDRKKTEDAIATFNVDHKSSKDLRQYLANLAEQCSADNSDADLPVVIILDNLHHIGSLSDIFNGFLNCKYNTCPYIIGTMNQGVTSSPNLELHHNFRWVLCANHTEPVKGFLGRYLRRKLIETEIERNMRNNDLIKIIDWIPKTWHHLNSFLETHSSSDVTIGPRLFLPCPMDVDGSRVWFTDLWNYSLVPYLLEAVREGLQMYGKRAPWEDPSKWVADTYPWSSAPVQHEWPSLLQLRPEDVGFEGYASAQEGTTSKHIPQSDTEGDPLVRMKLFAFLCSLCFNLEFET</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0F3</accession>
    <name>A0A452J0F3_9SAUR</name>
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        <submittedName>
            <fullName evidence="5">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the PA-PLA1 family.</text>
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    <dbReference type="AlphaFoldDB" id="A0A452J0F3"/>
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        <property type="protein sequence ID" value="ENSGAGP00000033837.1"/>
        <property type="gene ID" value="ENSGAGG00000024071.1"/>
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        <property type="project" value="TreeGrafter"/>
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        <property type="term" value="F:metal ion binding"/>
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        <property type="term" value="F:phospholipase activity"/>
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        <property type="project" value="TreeGrafter"/>
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        <property type="term" value="F:triacylglycerol lipase activity"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="entry name" value="ankyrin repeat and SAM domain-containing protein 4B"/>
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        <property type="entry name" value="Transcription Factor, Ets-1"/>
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    </dbReference>
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        <property type="entry name" value="AB_hydrolase_fold"/>
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        <property type="entry name" value="WWE_SEC23-DDH2"/>
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        <property type="entry name" value="PA-PL1 PHOSPHOLIPASE FAMILY"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="PHOSPHOLIPASE DDHD2"/>
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    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
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        <location>
            <begin position="25"/>
            <end position="107"/>
        </location>
    </feature>
    <feature type="domain" description="DDHD" evidence="4">
        <location>
            <begin position="452"/>
            <end position="656"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1"/>
            <end position="21"/>
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        <location>
            <begin position="576"/>
            <end position="603"/>
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        <location>
            <begin position="1"/>
            <end position="10"/>
        </location>
    </feature>
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        <location>
            <begin position="11"/>
            <end position="20"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="576"/>
            <end position="601"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00038464"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS50918"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS51043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033837.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="667" mass="76517" checksum="F50B4483905EA583" modified="2019-05-08" version="1">MSAVEQQQAEPSPSSNSSSSFEVVEADTANKYEAVVPHWFYCKVIDSRERWIPFSQQDSEKLEEAHNSRKDDEQLVVPTEGGRYDVHLKKRLRCAMYWEEEVSEVRRCTWFYKGDKDNKYIPYSESFSEELEETYMIAITLDEWKKRLESPTREIIILHNPKLMVHYHPVTTSDDWGSTPTEQGRPRTVKRGVENISADIPCGEPLQIDHLVFVVHGIGPACDIRFRSIVQCVNDFRNVSLSMLQAHFKKAQEQQQIGRVEFLPVNWHSPLHSTGVDVDLERITLPSINRLRHFINDTILDVFFYNSSTYCQTIVDTVASEMNRLYRLFLQRNPDFKGGVSIAGHSLGSLILFDLLTNQKDSSEDEDDNKEGSQMGAYGQGDIGGVKEILKKLELSEYYSVFEKERMDTQALALCTEKDLKDMGIPLGPRTKILHYVRTKDETQVSANYPQLIYKPDIFFAFGSPIGMFLTVRGVKRIDPNYSLPTCKGFFNIFHPFDPVAYRIEPMIIPDLEFEPMLLPHHKGRKRMHLELKEGLTRMGMDLKNNLLGSLRIAWQSFTRSPLLAVEAAATEKEAETSLEKQTVHSDMKPEEPAAIGKEDTPPINVGMLNGGHRIDYVLQEKPIESFNEYLFALQGHLCYWESEDTVLLILKEIYQTQGITLDQPLQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0F6</accession>
    <name>A0A452J0F6_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="19">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">Muscle contraction.</text>
    </comment>
    <comment type="subunit">
        <text evidence="14">Muscle myosin is a hexameric protein that consists of 2 heavy chain subunits (MHC), 2 alkali light chain subunits (MLC) and 2 regulatory light chain subunits (MLC-2).</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Myofibril</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 15">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038330.1">
        <property type="protein sequence ID" value="ENSGAGP00000033841.1"/>
        <property type="gene ID" value="ENSGAGG00000020256.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030016">
        <property type="term" value="C:myofibril"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032982">
        <property type="term" value="C:myosin filament"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016460">
        <property type="term" value="C:myosin II complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000146">
        <property type="term" value="F:microfilament motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd01377">
        <property type="entry name" value="MYSc_class_II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000003">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000004">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000006">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000013">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000003">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000007">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000008">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.4820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.530:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.360:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000024">
        <property type="entry name" value="Myosin heavy chain, isoform J"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.720:FF:000001">
        <property type="entry name" value="Myosin heavy chain, muscle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.340">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.370">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.4820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.2420">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.360">
        <property type="entry name" value="Myosin S1 fragment, N-terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008989">
        <property type="entry name" value="Myosin_S1_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002928">
        <property type="entry name" value="Myosin_tail"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004009">
        <property type="entry name" value="SH3_Myosin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014751">
        <property type="entry name" value="XRCC4-like_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615">
        <property type="entry name" value="MYOSIN HEAVY CHAIN, NON-MUSCLE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615:SF69">
        <property type="entry name" value="MYOSIN-6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02736">
        <property type="entry name" value="Myosin_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01576">
        <property type="entry name" value="Myosin_tail_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90257">
        <property type="entry name" value="Myosin rod fragments"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57997">
        <property type="entry name" value="Tropomyosin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51844">
        <property type="entry name" value="SH3_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12 15" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="7 15" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="8" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0488">Methylation</keyword>
    <keyword evidence="10 15" id="KW-0505">Motor protein</keyword>
    <keyword evidence="11" id="KW-0514">Muscle protein</keyword>
    <keyword evidence="9 15" id="KW-0518">Myosin</keyword>
    <keyword evidence="6 15" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0787">Thick filament</keyword>
    <feature type="domain" description="Myosin N-terminal SH3-like" evidence="18">
        <location>
            <begin position="32"/>
            <end position="81"/>
        </location>
    </feature>
    <feature type="domain" description="Myosin motor" evidence="17">
        <location>
            <begin position="85"/>
            <end position="779"/>
        </location>
    </feature>
    <feature type="region of interest" description="Actin-binding" evidence="15">
        <location>
            <begin position="656"/>
            <end position="678"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1907"/>
            <end position="1952"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="16">
        <location>
            <begin position="1924"/>
            <end position="1937"/>
        </location>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <begin position="178"/>
            <end position="185"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004657"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022433"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023179"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00037488"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00038612"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS51844"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033841.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1952" mass="224731" checksum="C7F4AF1C8F1EF69A" modified="2019-05-08" version="1">MGDAHMAEFGAAAPYLRKSDKERLEAQTRIFDLRSDCFVSDEQEEFVKGKIVSRDGERLTVETENGKTVTVKESDVHQQNPPKFDKIEDMAMLTFLHEPAVLYNLKERYAAWMIYTYSGLFCVTVNPYKWLPVYNAEVVAAYRGKKRSEAPPHIFSISDNAYQNMLTNRENQSILITGESGAGKTVNTKRVIQYFASIAAISDRKKEGTSANKGTLEDQIIQANPALEAFGNAKTVRNDNSSRFGKFIRIHFGATGKLASADIETYLLEKSRVIFQLKAERNYHIFYQILSNQKPELLEMMLVTNNPYDYSFISQGEVTVASINDSEELIATDSAFDVLGFTPEEKVGVYKLTGAIMHFGNMKFKQKQREEQAEPDGTEDADKSAYLMGLNSADLLKGLCHPRVKVGNEYVTKGQNVQQVYYSIGALAKSVYEKMFSWMVTRINSTLETKQPRQYFIGVLDIAGFEIFDFNSFEQLCINFTNEKLQQFFNHHMFVLEQEEYKKEGIEWVFIDFGMDLQACIDLIEKPMGIMSILEEECMFPKATDMTFKAKLFDNHLGKSANFSKPRNIKGKPEAHFSLIHYAGTVDYNILGWLQKNKDPLNETVVGLYQKSSLKLLALLFANYAAAESGDGGKGKGAKKKGSSFQTVSALHRENLNKLMANLKTTHPHFVRCIIPNERKAPGVMDNPLVMHQLRCNGVLEGIRICRKGFPNRILYGDFRQRYRILNPAAIPEGQFIDSRKGAEKLLSSLDIDPNQYKFGHTKVFFKAGLLGLLEEMRDERLSRIITRIQAQSRGQLMRIEFKKILERRDALLVIQWNIRAFMGVKNWPWMKLYFKIKPLLKSAETEKEMQTMKEEFGRLKEALEKSEARRKELEEKMVSLLQEKNDLQLQVQAEQDNLNDAEERCDQLIKNKIQLEAKVKELTERMEDEEEMNAELTAKKRKLEDECSELKKDIDDLELTLAKVEKEKHATENKVKNLTEEMAGLDEIIAKLTKEKKSLQEAHQQALDDLQVEEDKVNSLTKAKLKLEQQADDLEGSLEQEKKIRMDLERAKRKLEGDLKLTQENIMDLENDKQQLEEKLKKKEFDINQQNSKIEDEQALGLQLQKKLKELQARIEELEEELEAERTARAKVEKLRSDLSRELEEISERLEEAGGATSVQIELNKKREAEFQKMRRDLEEATLQHEATAATLRKKHADSVAELGEQIDNLQRVKQKLEKEKSELKLELDDISSNMEQLLKAKVSLEKLSRTLEDQTAEHRAKLEETQRAMNDTSTQRAKLQTENGELSRQLEEKEALISQLTRGKQSYTQQMEDLKRQLEEETKAKNALAHALQSARHDCDLLREQYEEETEAKAELQRSLSKANSEVAQWRTKYETDAIQRTEELEEAKKKLAQRLQEAEEAVEAVNAKCSSLEKTKHRLQNEIEDLMVDVERSNAAAAALDKKQRNFDKILSEWKQKFEESQTELESSQKEARSLSTELFKLKNAYEESLEHLETFKRENKNLQEEISDLTEQLGEGGKNLHELEKVRKQLEAEKMELQAALEEAEASLEHEEGKILRAQLEFNQFKADFERKLAEKDEEMEQAKRNHLRVVDSLQTSLDAETRSRNEALRVKKKMEGDLNEMEIQLSHANRMAAEAQSHLKATQTQLKDSQLQLDDFIRANEDLKENIAIVERRNSLLQAELEELRAVVEQTERTRRLAEQELIEASERVQLLHTQNTSLINQKKKMEADLSQLQTEVEEAVQECRNAEEKAKKAITDAAMMAEELKKEQDTSAHLERMKKNMEQTIKDLQLRLDEAEQLALKGGKKQLQKLEARVRELENELESEQKRNAESIKGLRKSERRIKELTYQTEEDKKNLMRLQDLVDKLQMKVKAYKRQAEEAEEQANTNLAKFRKVQHELDEAEERADIAESQVNKLRAKSRDAGGKVSDRDGGPALRPAGPTQRLQP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0F7</accession>
    <name>A0A452J0F7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">CCAAT/enhancer-binding protein zeta</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="8">CCAAT-box-binding transcription factor</fullName>
        </alternativeName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Stimulates transcription from the HSP70 promoter.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the CBF/MAK21 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0F7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038370.1">
        <property type="protein sequence ID" value="ENSGAGP00000033878.1"/>
        <property type="gene ID" value="ENSGAGG00000024100.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000805">
        <property type="entry name" value="Similar to transcription factor CBF/MAK21"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005612">
        <property type="entry name" value="CCAAT-binding_factor"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040155">
        <property type="entry name" value="CEBPZ/Mak21-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12048">
        <property type="entry name" value="CCAAT-BINDING FACTOR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12048:SF0">
        <property type="entry name" value="CCAAT_ENHANCER-BINDING PROTEIN ZETA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03914">
        <property type="entry name" value="CBF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0010">Activator</keyword>
    <keyword evidence="7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0804">Transcription</keyword>
    <keyword evidence="4" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="CCAAT-binding factor" evidence="12">
        <location>
            <begin position="529"/>
            <end position="734"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="103"/>
            <end position="156"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="636"/>
            <end position="661"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="870"/>
            <end position="952"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1014"/>
            <end position="1034"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="103"/>
            <end position="136"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="11">
        <location>
            <begin position="888"/>
            <end position="938"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007797"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00031941"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00058879"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00073389"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF03914"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033878.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1034" mass="117831" checksum="00E928C2D6D438D0" modified="2019-05-08" version="1">MLGVAAAACSAVMAGPAGLRELDETDEEDGSESGFMLEEVLRLGGTKQDYIMLAALDETEDLVDGGKQGAIDDLGQGELEEFIQSLGVCKYAKKIMVVEEKEKKDKPAKKEKIGKKDTSAPSLEGKKENQVKENKGKVQINNGKDGKKKTAQRNEEHIPTVPVVKKEKQEEVFEFHARQVLLIKPGGKWYDLEYTNEFSSEQQNQVFVSKYKALAQRLYQQETDLYKSKTKLQKGASSAWMNNVVSSGTLTDRMAAMTLLIQDGSVHALQYVETLVNLIKKKGSRRQSLMALDTFKELLLSDLLPDNRKLHTFSQHPFNKLEKLSSGNRDSRDRRLILWYFEHQLKHLVAEFVQVLETLTHDSLVATKARALAVAYELLCNKPEEEKALLVQLVNKLGDPQNKIATKASYLLETLLHKHPNMKGVVCNEVERLLYRSNISPKAQYYAVCFLNQIVLSHEENDLANKLITLYFCFFRSCIKKKEVDSKMLSALLTGVNRAYPYAQIGDEKVKEQMNTLFKVLHLVNFSTSVQALMLLFQVMDSHQTVSDRYYTALYKKLLDPGLAACSKQSMFLNLVYKSVKADVVLRRVKAFVKRLLQVTCGQMPPFICGALYLVSEIMKIKPGLRVQLQDHVESEEEEHFHDLEEAEEDEEKFTDADKETAAERKNMMESSVKGDNTNSAASWVHHQNMSGGKNSSCYDPLHRNPLFCGAESTSLWELKKLSEHFHPSVALFAKTILEGDYIQYAGDPLQDFTLMRFLDRFVYRNPKLHKGKENTGSVVMQPKKKQLMNDVRSLAVNSKEFLAQDESKIPVDEVFFHRFYKKLDKEKRKRPVEEESVEDVDDDEFERILDSFEGDSYFDTVSKDDIDFAGNIKNKPKGGKKSKRDEECSDDLEDSDDEDEFNEMDDEVVSLGSMDEEFRDDIDEEGGAFMDVSEDDNSPGSKQQKKRKLKDTSMLAAAEEFGTLLDENVTSKFDNIGMNAMANKDNASIKQLKWEAERDDWLHNRDVKTIIKKKKQFRHKGLKNKHKGKKSRK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0F8</accession>
    <name>A0A452J0F8_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="19">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">Muscle contraction.</text>
    </comment>
    <comment type="subunit">
        <text evidence="14">Muscle myosin is a hexameric protein that consists of 2 heavy chain subunits (MHC), 2 alkali light chain subunits (MLC) and 2 regulatory light chain subunits (MLC-2).</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Myofibril</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 15">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038328.1">
        <property type="protein sequence ID" value="ENSGAGP00000033839.1"/>
        <property type="gene ID" value="ENSGAGG00000020256.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030016">
        <property type="term" value="C:myofibril"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032982">
        <property type="term" value="C:myosin filament"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016460">
        <property type="term" value="C:myosin II complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000146">
        <property type="term" value="F:microfilament motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd01377">
        <property type="entry name" value="MYSc_class_II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000003">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000004">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000006">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000013">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000003">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000007">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000008">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.4820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.530:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.360:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000024">
        <property type="entry name" value="Myosin heavy chain, isoform J"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.720:FF:000001">
        <property type="entry name" value="Myosin heavy chain, muscle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.340">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.370">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.4820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.2420">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.360">
        <property type="entry name" value="Myosin S1 fragment, N-terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008989">
        <property type="entry name" value="Myosin_S1_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002928">
        <property type="entry name" value="Myosin_tail"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004009">
        <property type="entry name" value="SH3_Myosin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014751">
        <property type="entry name" value="XRCC4-like_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615">
        <property type="entry name" value="MYOSIN HEAVY CHAIN, NON-MUSCLE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615:SF69">
        <property type="entry name" value="MYOSIN-6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02736">
        <property type="entry name" value="Myosin_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01576">
        <property type="entry name" value="Myosin_tail_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90257">
        <property type="entry name" value="Myosin rod fragments"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57997">
        <property type="entry name" value="Tropomyosin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51844">
        <property type="entry name" value="SH3_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12 15" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="7 15" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="8" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0488">Methylation</keyword>
    <keyword evidence="10 15" id="KW-0505">Motor protein</keyword>
    <keyword evidence="11" id="KW-0514">Muscle protein</keyword>
    <keyword evidence="9 15" id="KW-0518">Myosin</keyword>
    <keyword evidence="6 15" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0787">Thick filament</keyword>
    <feature type="domain" description="Myosin N-terminal SH3-like" evidence="18">
        <location>
            <begin position="38"/>
            <end position="87"/>
        </location>
    </feature>
    <feature type="domain" description="Myosin motor" evidence="17">
        <location>
            <begin position="91"/>
            <end position="767"/>
        </location>
    </feature>
    <feature type="region of interest" description="Actin-binding" evidence="15">
        <location>
            <begin position="644"/>
            <end position="666"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1895"/>
            <end position="1940"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="16">
        <location>
            <begin position="1912"/>
            <end position="1925"/>
        </location>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <begin position="184"/>
            <end position="191"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004657"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022433"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023179"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00037488"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00038612"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS51844"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033839.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1940" mass="223611" checksum="AB2AC2A86EA68293" modified="2019-05-08" version="1">PSASPHMGDAHMAEFGAAAPYLRKSDKERLEAQTRIFDLRSDCFVSDEQEEFVKGKIVSRDGERLTVETENGKTVTVKESDVHQQNPPKFDKIEDMAMLTFLHEPAVLYNLKERYAAWMIYTYSGLFCVTVNPYKWLPVYNAEVVAAYRGKKRSEAPPHIFSISDNAYQNMLTNRENQSILITGESGAGKTVNTKRVIQYFASIAAISDRKKEGTSGTLEDQIIQANPALEAFGNAKTVRNDNSSRFGKFIRIHFGATGKLASADIETYLLEKSRVIFQLKAERNYHIFYQILSNQKPELLEMMLVTNNPYDYSFISQGEVTVASINDSEELIATDSAFDVLGFTPEEKVGVYKLTGAIMHFGNMKFKQKQREEQAEPDGTEDADKSAYLMGLNSADLLKGLCHPRVKVGNEYVTKGQNVQQVYYSIGALAKSVYEKMFSWMVTRINSTLETKQPRQYFIGVLDIAGFEIFDFNSFEQLCINFTNEKLQQFFNHHMFVLEQEEYKKEGIEWVFIDFGMDLQACIDLIEKPMGIMSILEEECMFPKATDMTFKAKLFDNHLGKSANFSKPRNIKGKPEAHFSLIHYAGTVDYNILGWLQKNKDPLNETVVGLYQKSSLKLLALLFANYAAAESSYYTPTPSRFLTLALMANLKTTHPHFVRCIIPNERKAPGVMDNPLVMHQLRCNGVLEGIRICRKGFPNRILYGDFRQRYRILNPAAIPEGQFIDSRKGAEKLLSSLDIDPNQYKFGHTKVFFKAGLLGLLEEMRDERLSRIITRIQAQSRGQLMRIEFKKILERRDALLVIQWNIRAFMGVKNWPWMKLYFKIKPLLKSAETEKEMQTMKEEFGRLKEALEKSEARRKELEEKMVSLLQEKNDLQLQVQAEQDNLNDAEERCDQLIKNKIQLEAKVKELTERMEDEEEMNAELTAKKRKLEDECSELKKDIDDLELTLAKVEKEKHATENKVKNLTEEMAGLDEIIAKLTKEKKSLQEAHQQALDDLQVEEDKVNSLTKAKLKLEQQADDLEGSLEQEKKIRMDLERAKRKLEGDLKLTQENIMDLENDKQQLEEKLKKKEFDINQQNSKIEDEQALGLQLQKKLKELQARIEELEEELEAERTARAKVEKLRSDLSRELEEISERLEEAGGATSVQIELNKKREAEFQKMRRDLEEATLQHEATAATLRKKHADSVAELGEQIDNLQRVKQKLEKEKSELKLELDDISSNMEQLLKAKVSLEKLSRTLEDQTAEHRAKLEETQRAMNDTSTQRAKLQTENGELSRQLEEKEALISQLTRGKQSYTQQMEDLKRQLEEETKAKNALAHALQSARHDCDLLREQYEEETEAKAELQRSLSKANSEVAQWRTKYETDAIQRTEELEEAKKKLAQRLQEAEEAVEAVNAKCSSLEKTKHRLQNEIEDLMVDVERSNAAAAALDKKQRNFDKILSEWKQKFEESQTELESSQKEARSLSTELFKLKNAYEESLEHLETFKRENKNLQEEISDLTEQLGEGGKNLHELEKVRKQLEAEKMELQAALEEAEASLEHEEGKILRAQLEFNQFKADFERKLAEKDEEMEQAKRNHLRVVDSLQTSLDAETRSRNEALRVKKKMEGDLNEMEIQLSHANRMAAEAQSHLKATQTQLKDSQLQLDDFIRANEDLKENIAIVERRNSLLQAELEELRAVVEQTERTRRLAEQELIEASERVQLLHTQNTSLINQKKKMEADLSQLQTEVEEAVQECRNAEEKAKKAITDAAMMAEELKKEQDTSAHLERMKKNMEQTIKDLQLRLDEAEQLALKGGKKQLQKLEARVRELENELESEQKRNAESIKGLRKSERRIKELTYQTEEDKKNLMRLQDLVDKLQMKVKAYKRQAEEAEEQANTNLAKFRKVQHELDEAEERADIAESQVNKLRAKSRDAGGKVSDRDGGPALRPAGPTQRLQP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0F9</accession>
    <name>A0A452J0F9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">AAA+ ATPase domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the Nav/unc-53 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038331.1">
        <property type="protein sequence ID" value="ENSGAGP00000033842.1"/>
        <property type="gene ID" value="ENSGAGG00000024067.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0022008">
        <property type="term" value="P:neurogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00009">
        <property type="entry name" value="AAA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000316">
        <property type="entry name" value="Putative neuron navigator 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003593">
        <property type="entry name" value="AAA+_ATPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003959">
        <property type="entry name" value="ATPase_AAA_core"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057568">
        <property type="entry name" value="CortBP2_NAV1-like_AAA_lid"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039041">
        <property type="entry name" value="Nav/unc-53"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057126">
        <property type="entry name" value="NAV1-like_ubiquitin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12784:SF18">
        <property type="entry name" value="NEURON NAVIGATOR 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12784">
        <property type="entry name" value="STEERIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00004">
        <property type="entry name" value="AAA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25408">
        <property type="entry name" value="AAA_lid_NAV1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23092">
        <property type="entry name" value="Ubiquitin_6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00382">
        <property type="entry name" value="AAA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 3" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="AAA+ ATPase" evidence="5">
        <location>
            <begin position="1529"/>
            <end position="1683"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="86"/>
            <end position="141"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="384"/>
            <end position="826"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="861"/>
            <end position="886"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="920"/>
            <end position="978"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="1177"/>
            <end position="1261"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="1331"/>
            <end position="1362"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="3">
        <location>
            <begin position="1074"/>
            <end position="1157"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="102"/>
            <end position="115"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="389"/>
            <end position="417"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="418"/>
            <end position="432"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="433"/>
            <end position="443"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="489"/>
            <end position="498"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="525"/>
            <end position="537"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="584"/>
            <end position="602"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="667"/>
            <end position="680"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="697"/>
            <end position="706"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="716"/>
            <end position="737"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="763"/>
            <end position="773"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="774"/>
            <end position="799"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="4">
        <location>
            <begin position="806"/>
            <end position="815"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="861"/>
            <end position="885"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="946"/>
            <end position="955"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="956"/>
            <end position="966"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="967"/>
            <end position="978"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="1182"/>
            <end position="1199"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="1347"/>
            <end position="1362"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006255"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="SMART" id="SM00382"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033842.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1847" mass="199891" checksum="180CEC1E27E3BC21" modified="2019-05-08" version="1">MQRPAELSVFKGKAVRRGNARLQKQKSLTNLTLISEGEKRRYSYRENWFGNASKSSQNCHQREAEVEDRISLSKLLSQSVHSLYHTNSATSEKTEVLPPAGSNKSSRRSGGQSTDEGFEVDNEERDKSDDENIPCKSNFTSRNWAEEEEEGCLREGTAQLDEDVIMTMLGDLEQVLYTDILGEDPETRRMRTVKNIADLRQNLEETMSSLRGTQISHSTLETTFDSTVTTEVNGRSIPGLTSRSTPVTWRLGQASPRLQAGDAPSLGSGYPRSSAGRFIHTDPSRFMYTTPLRRAAVSRLGNMSQIDMNEKGSSDLDMSSEVDVGGYMSDGDILGKSLRTDDISSGYMTDGGLNLYTRSLNRIPDLAASRDVIQRGVHDVTVDADSWDDSSSVSSGLSDTLDNLSTDDLNTTSSVSSYSNITASSRKNTHTQLKTDSEKRFVTDSENWGSTEELKKPEEDFDSNVDGSGKWKGLPSGLSEDSEKGGQKASLSVSQTGSWRRGMTAQVGTTSSRHKAGTSALKTPGKTDDAKASEKVKTPLKGTSIQRSPSDAGKSSGDEGKKPPSGIGRSTATGSFGFKKAGVGSSTIITTSGATITSGSATLGKMPKSTAIGGKSNAGRKTSLDGSQNQDDGVLHVNSKTTLQYRSLPRPSKSSTSGIPGRGGHRSSTSSIDSNLSSKSAGAATTKLREPTKIGSGRSSPITINQTDKEKEKVAVSDSESVSLSSSPKSSPTSASACGTPGLRQPGSKYPDIASPTFRRLFGAKAGGKAASAPNTESVKPTSAMPSPSTTLARQGSLESPSSGTGSMGSAGGQSGSSSPLFSKPSELNTDVVSLSHSLASSPASVHSFTSGGLVWAANLSSSSAGSKDTPSYQSMTSLHTSSESIDLPLSHHSSLSGLTTGTQEVQSLLMRTGSVRSTLSESMQLDRNTLPKKGLRYTPSSRQTSQEESKEWLRSHSTGGLQDTGSQSPIVSPSAMSSSATGKYHFSNLVSPTNLSQFNLPGPSMMRSNSIPAQDSSFDLYDDSQLCGSATSLEERPRAISHSGSFRDSMEEVHGSSLSLVSSTSSLYSTAEEKAHSEQIHKLRRELVASQEKVATLTSQLSANAHLVAAFEKSLGNMTGRLQSLTMTAEQKESELIELRETIEMLKAQNSAAQAAIQGALNGPDHAHKDLLIRRQHSSESVSSINSATSHSSIGSGNDADSKKKKKKNWLRSSFKQAFGKKKSTKPPSSHSDIEELTDSSLPSSPKLPHNSGDCGTLSMKPSQSASAICECTEAEAEIILQLKSELREKELKLTDIRLEALSSAHHLDQIREAMNRMQNEIEILKAENDRLKAETGNTGKPARPSSESSSSTSSSSSRQSLGLSLNNLNITDSVTSDILLDDVTDGALPKEGHSVKIIVTISKGYGRSKDQKTQAYLIGSIGVSGKTKWDVLDGVIRRLFKEYIFRVDPATSLGLSSDCIASYCIGDLIRVHSLEVPELLPCGYLVGDNNIITVNLKGVEENSLDSFVFDTLIPKPITQRYFNLLMEHHRIILSGPSGTGKTYLANRLAEYIITKLDRKKTEDAIATFNVDHKSSKDLRQYLANLAEQCSADNSDADLPVVIILDNLHHIGSLSDIFNGFLNCKYNTCPYIIGTMNQGVTSSPNLELHHNFRWVLCANHTEPVKGFLGRYLRRKLIETEIERNMRNNDLIKIIDWIPKTWHHLNSFLETHSSSDVTIGPRLFLPCPMDVDGSRVWFTDLWNYSLVPYLLEAVREGLQMYGKRAPWEDPSKWVADTYPWSSAPVQHEWPSLLQLRPEDVGFEGYASAQEGTTSKHIPQSDTEGDPLVRMKLFAFLCSLCFNLEFET</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0G0</accession>
    <name>A0A452J0G0_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="3">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J0G0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038335.1">
        <property type="protein sequence ID" value="ENSGAGP00000033846.1"/>
        <property type="gene ID" value="ENSGAGG00000024076.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000151">
        <property type="term" value="C:ubiquitin ligase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016567">
        <property type="term" value="P:protein ubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006511">
        <property type="term" value="P:ubiquitin-dependent protein catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24173:SF27">
        <property type="entry name" value="ANKYRIN REPEAT AND SOCS BOX PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24173">
        <property type="entry name" value="ANKYRIN REPEAT CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00023">
        <property type="entry name" value="Ank"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0677">Repeat</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033846.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="225" mass="24748" checksum="53424B199EE958E8" modified="2019-05-08" version="1">MAEASYPNRPGYLQRETEAGLEAQSSDSHGATLGATDGNAGRNLKEWLREQFCDHPLEHCEDTRLHDAAYVGDLDTLRSLLQEESFQRYGADVDVNHHLPSRVPSPSSRPLTSLVVCPLYISAAYHNLQCFKLLLQAGANPDFNCCGPVNIQGFSRGSPVCVMDAVLRHGCEPAFVNLLIDFGANLNLVKAEGLDTESTGRVKVNPEALQMFKEARSKWLASPRH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="36" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0G5</accession>
    <name>A0A452J0G5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="26">Histone-lysine N-methyltransferase NSD3</fullName>
            <ecNumber evidence="24">2.1.1.370</ecNumber>
            <ecNumber evidence="25">2.1.1.371</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="27">Nuclear SET domain-containing protein 3</fullName>
        </alternativeName>
    </protein>
    <organism evidence="36 37">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="37" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="36" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="36" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="22">Histone methyltransferase. Preferentially dimethylates 'Lys-4' and 'Lys-27' of histone H3 forming H3K4me2 and H3K27me2. H3 'Lys-4' methylation represents a specific tag for epigenetic transcriptional activation, while 'Lys-27' is a mark for transcriptional repression.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="20">
            <text>L-lysyl(27)-[histone H3] + 2 S-adenosyl-L-methionine = N(6),N(6)-dimethyl-L-lysyl(27)-[histone H3] + 2 S-adenosyl-L-homocysteine + 2 H(+)</text>
            <dbReference type="Rhea" id="RHEA:64452"/>
            <dbReference type="Rhea" id="RHEA-COMP:15539"/>
            <dbReference type="Rhea" id="RHEA-COMP:15548"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29969"/>
            <dbReference type="ChEBI" id="CHEBI:57856"/>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
            <dbReference type="ChEBI" id="CHEBI:61976"/>
            <dbReference type="EC" id="2.1.1.371"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="21">
            <text>L-lysyl(4)-[histone H3] + 2 S-adenosyl-L-methionine = N(6),N(6)-dimethyl-L-lysyl(4)-[histone H3] + 2 S-adenosyl-L-homocysteine + 2 H(+)</text>
            <dbReference type="Rhea" id="RHEA:64448"/>
            <dbReference type="Rhea" id="RHEA-COMP:15540"/>
            <dbReference type="Rhea" id="RHEA-COMP:15547"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29969"/>
            <dbReference type="ChEBI" id="CHEBI:57856"/>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
            <dbReference type="ChEBI" id="CHEBI:61976"/>
            <dbReference type="EC" id="2.1.1.370"/>
        </reaction>
    </comment>
    <comment type="subunit">
        <text evidence="23">Interacts with BRD4. Interacts (via KIKL motif) with BRD3 (via NET domain).</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Chromosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="2.1.1.370" evidence="24"/>
    <dbReference type="EC" id="2.1.1.371" evidence="25"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038340.1">
        <property type="protein sequence ID" value="ENSGAGP00000033851.1"/>
        <property type="gene ID" value="ENSGAGG00000024078.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005694">
        <property type="term" value="C:chromosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140952">
        <property type="term" value="F:histone H3K27 dimethyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140946">
        <property type="term" value="F:histone H3K4 dimethyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032259">
        <property type="term" value="P:methylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006355">
        <property type="term" value="P:regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd15649">
        <property type="entry name" value="PHD1_NSD3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd15652">
        <property type="entry name" value="PHD2_NSD3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd15655">
        <property type="entry name" value="PHD3_NSD3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd15658">
        <property type="entry name" value="PHD4_NSD3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd15661">
        <property type="entry name" value="PHD5_NSD3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd20163">
        <property type="entry name" value="PWWP_NSD3_rpt1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd20166">
        <property type="entry name" value="PWWP_NSD3_rpt2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd19212">
        <property type="entry name" value="SET_NSD3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.170.270.10:FF:000002">
        <property type="entry name" value="Histone-lysine N-methyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.140:FF:000004">
        <property type="entry name" value="Histone-lysine N-methyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.140:FF:000030">
        <property type="entry name" value="Histone-lysine N-methyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.40.10:FF:000025">
        <property type="entry name" value="Histone-lysine N-methyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.40.10:FF:000106">
        <property type="entry name" value="Histone-lysine N-methyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.40.10:FF:000205">
        <property type="entry name" value="Histone-lysine N-methyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.40.10:FF:001336">
        <property type="entry name" value="Histone-lysine N-methyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.140">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.170.270.10">
        <property type="entry name" value="SET domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006560">
        <property type="entry name" value="AWS_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041306">
        <property type="entry name" value="C5HCH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055198">
        <property type="entry name" value="NSD_PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR059153">
        <property type="entry name" value="NSD_PHD-1st"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047456">
        <property type="entry name" value="PHD2_NSD3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047458">
        <property type="entry name" value="PHD4_NSD3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047527">
        <property type="entry name" value="PHD5_NSD3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055197">
        <property type="entry name" value="PHDvar_NSD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003616">
        <property type="entry name" value="Post-SET_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000313">
        <property type="entry name" value="PWWP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047451">
        <property type="entry name" value="PWWP_NSD3_rpt1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047453">
        <property type="entry name" value="PWWP_NSD3_rpt2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050777">
        <property type="entry name" value="SET2_Histone-Lys_MeTrsfase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001214">
        <property type="entry name" value="SET_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046341">
        <property type="entry name" value="SET_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047461">
        <property type="entry name" value="SET_NSD3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019786">
        <property type="entry name" value="Zinc_finger_PHD-type_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011011">
        <property type="entry name" value="Znf_FYVE_PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001965">
        <property type="entry name" value="Znf_PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019787">
        <property type="entry name" value="Znf_PHD-finger"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22884">
        <property type="entry name" value="SET DOMAIN PROTEINS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17907">
        <property type="entry name" value="AWS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17982">
        <property type="entry name" value="C5HCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23011">
        <property type="entry name" value="PHD-1st_NSD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22908">
        <property type="entry name" value="PHD_NSD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23004">
        <property type="entry name" value="PHDvar_NSD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00855">
        <property type="entry name" value="PWWP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00856">
        <property type="entry name" value="SET"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00570">
        <property type="entry name" value="AWS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00249">
        <property type="entry name" value="PHD"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00508">
        <property type="entry name" value="PostSET"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00293">
        <property type="entry name" value="PWWP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00317">
        <property type="entry name" value="SET"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57903">
        <property type="entry name" value="FYVE/PHD zinc finger"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF82199">
        <property type="entry name" value="SET domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF63748">
        <property type="entry name" value="Tudor/PWWP/MBT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51215">
        <property type="entry name" value="AWS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50868">
        <property type="entry name" value="POST_SET"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50812">
        <property type="entry name" value="PWWP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50280">
        <property type="entry name" value="SET"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01359">
        <property type="entry name" value="ZF_PHD_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50016">
        <property type="entry name" value="ZF_PHD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="15" id="KW-0007">Acetylation</keyword>
    <keyword evidence="14" id="KW-0156">Chromatin regulator</keyword>
    <keyword evidence="3" id="KW-0158">Chromosome</keyword>
    <keyword evidence="17 29" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="9" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="6" id="KW-0489">Methyltransferase</keyword>
    <keyword evidence="19" id="KW-0539">Nucleus</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="37" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0677">Repeat</keyword>
    <keyword evidence="8" id="KW-0949">S-adenosyl-L-methionine</keyword>
    <keyword evidence="18" id="KW-0804">Transcription</keyword>
    <keyword evidence="16" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="7" id="KW-0808">Transferase</keyword>
    <keyword evidence="13" id="KW-0832">Ubl conjugation</keyword>
    <keyword evidence="12" id="KW-0862">Zinc</keyword>
    <keyword evidence="11 28" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="PWWP" evidence="33">
        <location>
            <begin position="248"/>
            <end position="311"/>
        </location>
    </feature>
    <feature type="domain" description="PHD-type" evidence="31">
        <location>
            <begin position="680"/>
            <end position="727"/>
        </location>
    </feature>
    <feature type="domain" description="PWWP" evidence="33">
        <location>
            <begin position="940"/>
            <end position="1002"/>
        </location>
    </feature>
    <feature type="domain" description="AWS" evidence="35">
        <location>
            <begin position="1073"/>
            <end position="1123"/>
        </location>
    </feature>
    <feature type="domain" description="SET" evidence="32">
        <location>
            <begin position="1125"/>
            <end position="1242"/>
        </location>
    </feature>
    <feature type="domain" description="Post-SET" evidence="34">
        <location>
            <begin position="1249"/>
            <end position="1265"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="30">
        <location>
            <begin position="123"/>
            <end position="154"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="30">
        <location>
            <begin position="190"/>
            <end position="231"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="30">
        <location>
            <begin position="322"/>
            <end position="345"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="30">
        <location>
            <begin position="383"/>
            <end position="445"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="30">
        <location>
            <begin position="522"/>
            <end position="677"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="29">
        <location>
            <begin position="1014"/>
            <end position="1041"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="29">
        <location>
            <begin position="1275"/>
            <end position="1302"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="30">
        <location>
            <begin position="131"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="30">
        <location>
            <begin position="205"/>
            <end position="224"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="30">
        <location>
            <begin position="405"/>
            <end position="423"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="30">
        <location>
            <begin position="522"/>
            <end position="542"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="30">
        <location>
            <begin position="562"/>
            <end position="574"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="30">
        <location>
            <begin position="650"/>
            <end position="670"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022454"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022603"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022691"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00050891"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00052270"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00056539"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00065976"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00066818"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00066819"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00068930"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="ARBA" id="ARBA00081740"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00146"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="31">
        <source>
            <dbReference type="PROSITE" id="PS50016"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="32">
        <source>
            <dbReference type="PROSITE" id="PS50280"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="33">
        <source>
            <dbReference type="PROSITE" id="PS50812"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="34">
        <source>
            <dbReference type="PROSITE" id="PS50868"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="35">
        <source>
            <dbReference type="PROSITE" id="PS51215"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="36">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033851.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="37">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1417" mass="159214" checksum="0EA399C6F9E4CC89" modified="2019-05-08" version="1">MDFSFSFMQGIMGNTIQQPPQLIDSANIRQEDAFDASSDIAEDGGQTPYETALQQGFQYPTPTTEDLPPITNGYPPTINMYEPQAKYQTYSQYPNGSANGFGAVRNFSPPEYYQMENSNMRPHEVLEKSSPPQPPPPPPPSVPQTVIPKKTGSPEIKLKITKTIQNGRELFESSLCGDLLNEVQASEYAKMKHEGRKEKRKKSSKHDSSRSEERKSHKIPKLEPEEQNVQQFSPSLPTQVTTDIKFQVGDLVWSKVGTYPWWPCMVSSDPQLEVHTKINTRGAREYHVQFFSNQPERAWVHEKRVREYKGHKQYEQLLAEATKQASNHSEKQKIRKPRPQRERAQWDIGIAHAEKALKMTREERIEQYTFIYIDKEPEEALSKAKKTAAPKSEAKKNRRPKPALNAQTEQTKVVSASPSLSNTEVRRQSQRRQSSVEEEEPPPPVKIAWKTAAARKSLPASITMHKGSLDLQKCNMSPVVKIEQVFALQNAAGDGKFIDQFVYSTKGVGNKTEISVKGQDKLISTSNQKSERATAQNASSPETTSGSTGSVEKKQQRRSIRTRSESEKSTEVVPKKKIKKEQVEMVPQTAVKTGLQKGASEISDSCKPLKKRSRASTDVELTSSAYRDTSDSDSRGLNDLQGSFGKHSDSPSATADADASDVQSVDSSLSRRGTGTNKKDTVCQICESSGESLVSCEAECCRVFHMECLGLKSIPDGKFICMECKNGQHTCFSCKLPGKDVKRCSVNACGKFYHEACVRKFATAVFESRGFRCPQHCCTACSTDKDIHKASKGRMVRCLRCPVAYHSGDGCIAAGSLFVSSHILICSNHSKRNHSSSAVNVGFCFVCARGLVVQDHSDPLFSSYAYKSHYLLNESNRAELMKLPMIPPPSSASKKKCEKGGKLLCCESCPASFHPECLNIEMPEGCWNCNDCKAGKKLRYKQIVWVKLGNYRWWPAEICNPRSVPLNIQGLKHDIGDFPVFFFGSHDYYWVHQGRVFPYVEGDKSFAEGQTSINKTFKKALEEAAKRFQELKAQRESKEALEIERNSRKPPPYKHIKSNKVIGKVQIQVADLSEIPRCNCKPSDENPCGLESECLNRMLQYECHPQVCPAGERCQNQCFTKRLYPDAEIIKTDRRGWGLRTKRNIKKGEFVNEYVGELIDEEECRLRIKRAHENSITNFYMLTVTKDRIIDAGPKGNFSRFMNHSCNPNCETQKWTVNGDVRVGLFALRDIPAGMELTFNYNLDCLGNGRTECHCGAENCSGFLGVRPKTAFATANEEKAKNAKLKQKKRKIKTEQKQMHEDNCFQCGDGGELVMCDKKDCPKAYHLLCLNLTQPPFGKWECPWHQCDICCNPAISFCEFCPRSFCKDHEKGALVASALDGCLCCSEHNPKSPMAPEYWSKIKCKLQPQNPGEEAKE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0G7</accession>
    <name>A0A452J0G7_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="19">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">Muscle contraction.</text>
    </comment>
    <comment type="subunit">
        <text evidence="14">Muscle myosin is a hexameric protein that consists of 2 heavy chain subunits (MHC), 2 alkali light chain subunits (MLC) and 2 regulatory light chain subunits (MLC-2).</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Myofibril</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 15">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038338.1">
        <property type="protein sequence ID" value="ENSGAGP00000033849.1"/>
        <property type="gene ID" value="ENSGAGG00000020256.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030016">
        <property type="term" value="C:myofibril"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032982">
        <property type="term" value="C:myosin filament"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016460">
        <property type="term" value="C:myosin II complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000146">
        <property type="term" value="F:microfilament motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd01377">
        <property type="entry name" value="MYSc_class_II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000003">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000004">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000006">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000013">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000003">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000007">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000008">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.4820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.530:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000024">
        <property type="entry name" value="Myosin heavy chain, isoform J"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.720:FF:000001">
        <property type="entry name" value="Myosin heavy chain, muscle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.340">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.370">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.4820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.2420">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.360">
        <property type="entry name" value="Myosin S1 fragment, N-terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008989">
        <property type="entry name" value="Myosin_S1_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002928">
        <property type="entry name" value="Myosin_tail"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004009">
        <property type="entry name" value="SH3_Myosin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014751">
        <property type="entry name" value="XRCC4-like_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615">
        <property type="entry name" value="MYOSIN HEAVY CHAIN, NON-MUSCLE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615:SF69">
        <property type="entry name" value="MYOSIN-6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02736">
        <property type="entry name" value="Myosin_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01576">
        <property type="entry name" value="Myosin_tail_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90257">
        <property type="entry name" value="Myosin rod fragments"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57997">
        <property type="entry name" value="Tropomyosin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51844">
        <property type="entry name" value="SH3_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12 15" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="7 15" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="8" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0488">Methylation</keyword>
    <keyword evidence="10 15" id="KW-0505">Motor protein</keyword>
    <keyword evidence="11" id="KW-0514">Muscle protein</keyword>
    <keyword evidence="9 15" id="KW-0518">Myosin</keyword>
    <keyword evidence="6 15" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0787">Thick filament</keyword>
    <feature type="domain" description="Myosin N-terminal SH3-like" evidence="18">
        <location>
            <begin position="32"/>
            <end position="83"/>
        </location>
    </feature>
    <feature type="domain" description="Myosin motor" evidence="17">
        <location>
            <begin position="83"/>
            <end position="776"/>
        </location>
    </feature>
    <feature type="region of interest" description="Actin-binding" evidence="15">
        <location>
            <begin position="653"/>
            <end position="675"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1897"/>
            <end position="1942"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="16">
        <location>
            <begin position="1914"/>
            <end position="1927"/>
        </location>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <begin position="176"/>
            <end position="183"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004657"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022433"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023179"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00037488"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00038612"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS51844"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033849.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1942" mass="223487" checksum="80A76677FFF07CD4" modified="2019-05-08" version="1">MGDAHMAEFGAAAPYLRKSDKERLEAQTRIFDLRSDCFVSDEQEEFVKGKIVSRDGERLTVETENGKVRLGTAESGQNPPKFDKIEDMAMLTFLHEPAVLYNLKERYAAWMIYTYSGLFCVTVNPYKWLPVYNAEVVAAYRGKKRSEAPPHIFSISDNAYQNMLTNRENQSILITGESGAGKTVNTKRVIQYFASIAAISDRKKEGTSANKGTLEDQIIQANPALEAFGNAKTVRNDNSSRFGKFIRIHFGATGKLASADIETYLLEKSRVIFQLKAERNYHIFYQILSNQKPELLEMMLVTNNPYDYSFISQGEVTVASINDSEELIATDSAFDVLGFTPEEKVGVYKLTGAIMHFGNMKFKQKQREEQAEPDGTEDADKSAYLMGLNSADLLKGLCHPRVKVGNEYVTKGQNVQQVYYSIGALAKSVYEKMFSWMVTRINSTLETKQPRQYFIGVLDIAGFEIFDFNSFEQLCINFTNEKLQQFFNHHMFVLEQEEYKKEGIEWVFIDFGMDLQACIDLIEKPMGIMSILEEECMFPKATDMTFKAKLFDNHLGKSANFSKPRNIKGKPEAHFSLIHYAGTVDYNILGWLQKNKDPLNETVVGLYQKSSLKLLALLFANYAAAESGEKTRQGERKKGSSFQTVSALHRENLNKLMANLKTTHPHFVRCIIPNERKAPGVMDNPLVMHQLRCNGVLEGIRICRKGFPNRILYGDFRQRYRILNPAAIPEGQFIDSRKGAEKLLSSLDIDPNQYKFGHTKVFFKAGLLGLLEEMRDERLSRIITRIQAQSRGQLMQGARDALLVIQWNIRAFMGVKNWPWMKLYFKIKPLLKSAETEKEMQVRSKEFGRLKEALEKSEARRKELEEKMVSLLQEKNDLQLQVQAEQDNLNDAEERCDQLIKNKIQLEAKVKELTERMEDEEEMNAELTAKKRKLEDECSELKKDIDDLELTLAKVEKEKHATENKVKNLTEEMAGLDEIIAKLTKEKKSLQEAHQQALDDLQVEEDKVNSLTKAKLKLEQQADDLEGSLEQEKKIRMDLERAKRKLEGDLKLTQENIMDLENDKQQLEEKLKKKEFDINQQNSKIEDEQALGLQLQKKLKELQARIEELEEELEAERTARAKVEKLRSDLSRELEEISERLEEAGGATSVQIELNKKREAEFQKMRRDLEEATLQHEATAATLRKKHADSVAELGEQIDNLQRVKQKLEKEKSELKLELDDISSNMEQLLKAKVSLEKLSRTLEDQTAEHRAKLEETQRAMNDTSTQRAKLQTENGELSRQLEEKEALISQLTRGKQSYTQQMEDLKRQLEEETKAKNALAHALQSARHDCDLLREQYEEETEAKAELQRSLSKANSEVAQWRTKYETDAIQRTEELEEAKKKLAQRLQEAEEAVEAVNAKCSSLEKTKHRLQNEIEDLMVDVERSNAAAAALDKKQRNFDKILSEWKQKFEESQTELESSQKEARSLSTELFKLKNAYEESLEHLETFKRENKNLQEEISDLTEQLGEGGKNLHELEKVRKQLEAEKMELQAALEEAEASLEHEEGKILRAQLEFNQFKADFERKLAEKDEEMEQAKRNHLRVVDSLQTSLDAETRSRNEALRVKKKMEGDLNEMEIQLSHANRMAAEAQSHLKATQTQLKDSQLQLDDFIRANEDLKENIAIVERRNSLLQAELEELRAVVEQTERTRRLAEQELIEASERVQLLHTQNTSLINQKKKMEADLSQLQTEVEEAVQECRNAEEKAKKAITDAAMMAEELKKEQDTSAHLERMKKNMEQTIKDLQLRLDEAEQLALKGGKKQLQKLEARVRELENELESEQKRNAESIKGLRKSERRIKELTYQTEEDKKNLMRLQDLVDKLQMKVKAYKRQAEEAEEQANTNLAKFRKVQHELDEAEERADIAESQVNKLRAKSRDAGGKVSDRDGGPALRPAGPTQRLQP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0G9</accession>
    <name>A0A452J0G9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Integrin alpha-V</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="19">Vitronectin receptor subunit alpha</fullName>
        </alternativeName>
    </protein>
    <organism evidence="26 27">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="27" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="26" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="26" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell junction</location>
            <location evidence="1">Focal adhesion</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2 21">Membrane</location>
            <topology evidence="2 21">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 21">Belongs to the integrin alpha chain family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0G9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038291.1">
        <property type="protein sequence ID" value="ENSGAGP00000033803.1"/>
        <property type="gene ID" value="ENSGAGG00000023825.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009897">
        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005925">
        <property type="term" value="C:focal adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008305">
        <property type="term" value="C:integrin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001525">
        <property type="term" value="P:angiogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033627">
        <property type="term" value="P:cell adhesion mediated by integrin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098609">
        <property type="term" value="P:cell-cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007160">
        <property type="term" value="P:cell-matrix adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007229">
        <property type="term" value="P:integrin-mediated signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.130:FF:000003">
        <property type="entry name" value="Integrin alpha V"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1510:FF:000001">
        <property type="entry name" value="Integrin alpha V"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1530:FF:000002">
        <property type="entry name" value="integrin alpha-V isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.930:FF:000001">
        <property type="entry name" value="Integrin subunit alpha V"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1460:FF:000001">
        <property type="entry name" value="Integrin, alpha V"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.930">
        <property type="entry name" value="Bicelle-embedded integrin alpha(iib) transmembrane segment"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.130">
        <property type="entry name" value="Integrin alpha, N-terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1460">
        <property type="entry name" value="Integrin domains. Chain A, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1510">
        <property type="entry name" value="ntegrin, alpha v. Chain A, domain 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1530">
        <property type="entry name" value="ntegrin, alpha v. Chain A, domain 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013517">
        <property type="entry name" value="FG-GAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013519">
        <property type="entry name" value="Int_alpha_beta-p"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000413">
        <property type="entry name" value="Integrin_alpha"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018184">
        <property type="entry name" value="Integrin_alpha_C_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013649">
        <property type="entry name" value="Integrin_alpha_Ig-like_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048285">
        <property type="entry name" value="Integrin_alpha_Ig-like_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048286">
        <property type="entry name" value="Integrin_alpha_Ig-like_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028994">
        <property type="entry name" value="Integrin_alpha_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032695">
        <property type="entry name" value="Integrin_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23220">
        <property type="entry name" value="INTEGRIN ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23220:SF4">
        <property type="entry name" value="INTEGRIN ALPHA-V"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01839">
        <property type="entry name" value="FG-GAP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08441">
        <property type="entry name" value="Integrin_A_Ig_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20805">
        <property type="entry name" value="Integrin_A_Ig_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20806">
        <property type="entry name" value="Integrin_A_Ig_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00357">
        <property type="entry name" value="Integrin_alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01185">
        <property type="entry name" value="INTEGRINA"/>
    </dbReference>
    <dbReference type="SMART" id="SM00191">
        <property type="entry name" value="Int_alpha"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69318">
        <property type="entry name" value="Integrin alpha N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69179">
        <property type="entry name" value="Integrin domains"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51470">
        <property type="entry name" value="FG_GAP"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00242">
        <property type="entry name" value="INTEGRIN_ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0106">Calcium</keyword>
    <keyword evidence="10 21" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="11" id="KW-0965">Cell junction</keyword>
    <keyword evidence="4" id="KW-0165">Cleavage on pair of basic residues</keyword>
    <keyword evidence="15" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="17" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13 21" id="KW-0401">Integrin</keyword>
    <keyword evidence="14 21" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="16 21" id="KW-0675">Receptor</keyword>
    <keyword evidence="27" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="7 21" id="KW-0732">Signal</keyword>
    <keyword evidence="5 21" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="12 21" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="21">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5018817853" description="Integrin alpha-V" evidence="21">
        <location>
            <begin position="21"/>
            <end position="1033"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="979"/>
            <end position="1001"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="20">
        <location>
            <begin position="22"/>
            <end position="87"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="20">
        <location>
            <begin position="98"/>
            <end position="159"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="20">
        <location>
            <begin position="222"/>
            <end position="276"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="20">
        <location>
            <begin position="277"/>
            <end position="342"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="20">
        <location>
            <begin position="343"/>
            <end position="400"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="20">
        <location>
            <begin position="404"/>
            <end position="467"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha first immunoglubulin-like" evidence="23">
        <location>
            <begin position="452"/>
            <end position="610"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha second immunoglobulin-like" evidence="24">
        <location>
            <begin position="611"/>
            <end position="739"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha third immunoglobulin-like" evidence="25">
        <location>
            <begin position="757"/>
            <end position="968"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="1012"/>
            <end position="1033"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="22">
        <location>
            <begin position="1012"/>
            <end position="1027"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00008054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022685"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023037"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00068422"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00082120"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00803"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="RuleBase" id="RU003762"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="Pfam" id="PF08441"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="Pfam" id="PF20805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="Pfam" id="PF20806"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033803.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1033" mass="114341" checksum="65B516E0F0F02774" modified="2019-05-08" version="1">MARLRLLLLLLGRWLALGGAFNLDADGPAVYSGSEGSYFGFAVDFFAPDSSSIFLLVGAPKANTTQPDIVEGGQVLKCNWNSNKNCQPVIFDSTGDRNFAPNDPLEFKSHQWFGASVRSKNDKILACAPLYHWRTEVKQEREPVGTCYLHDGSKAVEYAPCRSTIIDADGQGFCQGGFSIDFTKQLSILMGFFPAGQLISDRVAEIVSKYNSKVYSIKYDDQLATRSASAAFDDSYLGYSVAVGEFSGDGIEDFVSGVPRAARTLGMVSIYNGKNMSSLYNFTGEQMAAYFGYSVAATDINGDNYTDLFIGAPLFMERGSDGKLQEVGQVSICLQQASGGFQIIKLNGFEIFARFSSSIAPLGDLDQDGFNDIAVAAPYGGEDKRGVVYIYNGRATGLNAVPSQILEGQWAARTMPPSFGYSLKGATDVDKNGYPDLIVGAFGVDTAVLYRARPVISVNAGLEVNPTILNPEEKICELPDTRVKVSCFNVKFCLTADGKGKLPNTLTFQVELLLDKLKQKGALRRALFLHNKQPGHSKNMTVVKGGTMRCEELQAFLRDESEFRDKLTPITVFMEYRLDYKTAKDSAGLQPILNQFTPANISRQTHILLDCGEDNICKPKLQVAVHSDQKQIYIGDDNPLTLIVDAQNQGEGAYEAELFVIIPPQADFIGVVRNNEALARLSCAFKTENQTRLVVCDLGNPMKAGTKLLAGLRFSVNQQSEMDTSVKFDLQIRSSNLFDNLSPTTSYQADLAILASVEIRGVSSPDHIFLPIANWQPQENPETEEDIGPLVQHIYELRNNGPSAFSKVMLNLQWPYKYNNNTLLYIMQYEIDGPMNCTSDMEINPLNIKISVPKADEKNETIRDEDNRDHHINRRDITAIEGDVHTLGCGTAECLKIVCQVGRLERGKSAILYVKSRLWTQTFMNKENQNHSYSLKSSASFNVIEFPYKNLSFEDIHNSTVVTTNITWGIQPQPMPVPVWVIILAVLAGLLLLAVLVFVMYRMGFFKRVRPPQEEQEREQLQPHENGEGTSEA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0J7</accession>
    <name>A0A452J0J7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Vascular endothelial growth factor receptor 3</fullName>
            <ecNumber evidence="4">2.7.10.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="34 35">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="35" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="34" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="34" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="25">
            <text>L-tyrosyl-[protein] + ATP = O-phospho-L-tyrosyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10596"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.10.1"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="30">Membrane</location>
            <topology evidence="30">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="30">Belongs to the protein kinase superfamily. Tyr protein kinase family. CSF-1/PDGF receptor subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.10.1" evidence="4"/>
    <dbReference type="AlphaFoldDB" id="A0A452J0J7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038412.1">
        <property type="protein sequence ID" value="ENSGAGP00000033918.1"/>
        <property type="gene ID" value="ENSGAGG00000024090.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043235">
        <property type="term" value="C:receptor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019838">
        <property type="term" value="F:growth factor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004714">
        <property type="term" value="F:transmembrane receptor protein tyrosine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001525">
        <property type="term" value="P:angiogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030335">
        <property type="term" value="P:positive regulation of cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043408">
        <property type="term" value="P:regulation of MAPK cascade"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048010">
        <property type="term" value="P:vascular endothelial growth factor receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00096">
        <property type="entry name" value="Ig"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="CDD" id="cd05863">
        <property type="entry name" value="IgI_VEGFR-3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000077">
        <property type="entry name" value="Vascular endothelial growth factor receptor 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000532">
        <property type="entry name" value="Vascular endothelial growth factor receptor 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000041">
        <property type="entry name" value="Vascular endothelial growth factor receptor 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000143">
        <property type="entry name" value="Vascular endothelial growth factor receptor 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000247">
        <property type="entry name" value="Vascular endothelial growth factor receptor 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000479">
        <property type="entry name" value="Vascular endothelial growth factor receptor 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000548">
        <property type="entry name" value="vascular endothelial growth factor receptor 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013098">
        <property type="entry name" value="Ig_I-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050122">
        <property type="entry name" value="RTK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001245">
        <property type="entry name" value="Ser-Thr/Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008266">
        <property type="entry name" value="Tyr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020635">
        <property type="entry name" value="Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001824">
        <property type="entry name" value="Tyr_kinase_rcpt_3_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041348">
        <property type="entry name" value="VEGFR-2_TMD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416">
        <property type="entry name" value="TYROSINE-PROTEIN KINASE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416:SF49">
        <property type="entry name" value="VASCULAR ENDOTHELIAL GROWTH FACTOR RECEPTOR 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07679">
        <property type="entry name" value="I-set"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13927">
        <property type="entry name" value="Ig_3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07714">
        <property type="entry name" value="PK_Tyr_Ser-Thr"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21339">
        <property type="entry name" value="VEGFR-1-like_Ig-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17988">
        <property type="entry name" value="VEGFR-2_TMD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000615">
        <property type="entry name" value="TyrPK_CSF1-R"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00109">
        <property type="entry name" value="TYRKINASE"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01832">
        <property type="entry name" value="VEGFRECEPTOR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01835">
        <property type="entry name" value="VEGFRECEPTR3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00219">
        <property type="entry name" value="TyrKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00109">
        <property type="entry name" value="PROTEIN_KINASE_TYR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00240">
        <property type="entry name" value="RECEPTOR_TYR_KIN_III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0037">Angiogenesis</keyword>
    <keyword evidence="16 27 29" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="20" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="22" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="24 30" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="15" id="KW-0418">Kinase</keyword>
    <keyword evidence="28" id="KW-0460">Magnesium</keyword>
    <keyword evidence="18 31" id="KW-0472">Membrane</keyword>
    <keyword evidence="28" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="14 27 29" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="23" id="KW-0539">Nucleus</keyword>
    <keyword evidence="8" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="21 30" id="KW-0675">Receptor</keyword>
    <keyword evidence="35" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="13" id="KW-0677">Repeat</keyword>
    <keyword evidence="12" id="KW-0732">Signal</keyword>
    <keyword evidence="10" id="KW-0808">Transferase</keyword>
    <keyword evidence="11 30 31" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="17 31" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="19" id="KW-0829">Tyrosine-protein kinase</keyword>
    <feature type="transmembrane region" description="Helical" evidence="31">
        <location>
            <begin position="623"/>
            <end position="647"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="33">
        <location>
            <begin position="127"/>
            <end position="219"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="33">
        <location>
            <begin position="224"/>
            <end position="308"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="33">
        <location>
            <begin position="315"/>
            <end position="410"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="33">
        <location>
            <begin position="495"/>
            <end position="611"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="32">
        <location>
            <begin position="692"/>
            <end position="982"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="26">
        <location>
            <position position="846"/>
        </location>
    </feature>
    <feature type="binding site" evidence="27">
        <location>
            <begin position="699"/>
            <end position="706"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="27 29">
        <location>
            <position position="726"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="27">
        <location>
            <position position="850"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="28">
        <location>
            <position position="851"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="28">
        <location>
            <position position="864"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00011902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022258"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022657"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023137"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00051243"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="PIRSR" id="PIRSR000615-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="PIRSR" id="PIRSR000615-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="PIRSR" id="PIRSR000615-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="RuleBase" id="RU000311"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="31">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="32">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="33">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="34">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033918.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="35">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1072" mass="122224" checksum="D6094054FACCDC94" modified="2019-05-08" version="1">MPEIHLFPVSPPSVLLLPLFILSFLYFSFSDFEQPFINRPETLLINKKENTWVPCLVSIPDLNITLFLPSSIIYPDGRRILWDNKKGMLVPTHLIRDSLFVQCETIIDKKLFRSNYFIVHVTGNELYDIQLFPKKAMELLVGEKLVLNCTVWAEFNAGVNFQWDYPGKQVHLPERRSQQTHTELSSILIIQNVSQQDLGKYMCMASNGEQTFKESIDVIVHEKPFINVEWRKGSVIEATAGDEVVKLPVKVVAYPPPEFQWYKDGKLIPNKQSPYSMQIKDVSEQNSGTYTLVLWNRLAGLEERISLQLIVNVPPHIHEKEASSPSIYSRRSHQALTCTVYGIPAPEVIQWQWSREDPVSPLTISHSDIHFLYQTVSKLVIQEANVSVMYKCVASNKVGRDERLIYFYVTNGFEIESQPSEEPIEGQDLRLSCNADNYTYENLQWYRLNLSKLHHEEGNPLILDCKNVHHYATKMQGELHFKPDSNDAALLLTIPNISLREEGDYVCEVQNRKTREKHCHKKYISVQGEERGNIWVNVSDSIEMRCKVDGAHIPDISWYKDEKLVEEVSGIDLADSNQRLSIQRVREEDAGLYLCSVCNAKGCVNSSASVSVEGSDDRTNVEIVILIGTGVIAVFFWILLILIFCNIKRPAHADIKTGYLSIIMDPGEVPLEEQCEYLPYDSSKWEFPRERLRLGKVLGHGAFGKVVEASAFGINKSNSCETVAVKMLKEGATASEHKALMSELKILIHIGNHLNVVNLLGACTKPNGPLMVIVEFCKYGNLSNYLRTKREGFSPYRVCEITFFLSLWTVDDLWQSPLTMEDLICYSFQVARGMEFLASRKCIHRDLAARNILLSENNVVKICDFGLARDIYKDPDYVRKGSARLPLKWMAPESIFDKVYTTQSDVWSFGVLLWEIFSLGASPYPGVQINEEFCQRLKDGTRMRAPEYATAEIYRIMLSCWHGDPKERPTFSDLVEILGDLLQENVQQVRFSCTVIIQHQTALGQFSSNSLFFLSLLAIPLDNQTDSGMVLASEEFERIENRHRKEGGFRYLLNSLPWTTLVVHSSACCEAP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0J8</accession>
    <name>A0A452J0J8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">GRIP domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000038374.1">
        <property type="protein sequence ID" value="ENSGAGP00000033882.1"/>
        <property type="gene ID" value="ENSGAGG00000024101.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031267">
        <property type="term" value="F:small GTPase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048193">
        <property type="term" value="P:Golgi vesicle transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.220.60">
        <property type="entry name" value="GRIP domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000237">
        <property type="entry name" value="GRIP_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19327">
        <property type="entry name" value="GOLGIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19327:SF0">
        <property type="entry name" value="GOLGIN SUBFAMILY A MEMBER 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01465">
        <property type="entry name" value="GRIP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00755">
        <property type="entry name" value="Grip"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF101283">
        <property type="entry name" value="GRIP domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50913">
        <property type="entry name" value="GRIP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="GRIP" evidence="3">
        <location>
            <begin position="2269"/>
            <end position="2316"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1"/>
            <end position="62"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="82"/>
            <end position="127"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="139"/>
            <end position="194"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="324"/>
            <end position="362"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="197"/>
            <end position="245"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="366"/>
            <end position="740"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="796"/>
            <end position="1070"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="1105"/>
            <end position="1235"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="1396"/>
            <end position="1455"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="1497"/>
            <end position="1531"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="1564"/>
            <end position="1905"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="1941"/>
            <end position="2119"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="2150"/>
            <end position="2202"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="25"/>
            <end position="49"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="89"/>
            <end position="124"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="140"/>
            <end position="168"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="169"/>
            <end position="178"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="327"/>
            <end position="338"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS50913"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033882.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2328" mass="272811" checksum="4083B1BD2B5C7BCB" modified="2019-05-08" version="1">MFKKLRQKIVEEQTQPRSPGGRAPQQPQTYSKSSPPSPGNRSRTSSLTEQSDEGTLTPDKELLAGMIAEPAFLSEYTIFALDPKRPKPQSDSVNVSKQTTPKSTDSNGSEPPSPQLSDTQTFAQKLQLRVPSMESLFRSPVKESLFRSSSKESLVRTSSRESLNRLDFDTPGPTFDPPSDVESEAEESLGNVDSLGKEQLLQRLRRMERSLGSYRGKYSELVTAYQTVQREKKKLQGILSQSQDKALRRIGELREAVAVLKADIREGGFSYDIWSSSMKELQMDQQAKKHLQEEFDASLEEKDQLISVLQTQVSLLKQRLQNGQISTELPETNTETEPQSPAKEISAENVLEPGGTEGDGDSAKTLETLIKRVKRQENLLQRCKEMIRSHKERSAQLTNEKEALQEQLEERLQELEKMKELHMTEKTKLITQLRDAKNLIEQLEQDKGMVIAETKRQMHETMEMKEEEIAQLRAHIKQITTQGEELREQKEKSEKAAFEELEKALSTAQKVEEARKKLQAEMDEKIKAIEKTSEEERVTLQQELTRVKQEVVEIMKKSSEERVAELEKLHKRELATKEQELNERLQVQGKEFQEKMRAALEKSQSEFLQTVQEKEQQESLALEELELQKKAIQSECEKKLQDMQQEIQTARTRILELESSLAKYSQDDKRQLEELSTLIESEKTKYNKEIIVMVEKHREELENVKQQEEELWTEKLQILKQQQQSTIEKLREKHEQEMDIMLKEKETVFRAHVEEMNAKTLEKLDVKQTELEALSFELSETLQIRHDLEKDGDKIKQESETKLEQQRKQYQEKMDMMVKEHDISVQGIQKAHTEETNQLKVQLEEKEKHLEELKAQEHKLKESLEISEAELKQVSAKLGDLCQSHQNNSSEQARVYEGQLTELQQKLTNMEAEKSHLKEQIVRAESQLNEVRTELDTYISRVRDLRQQLEEQSSENMQKITSLKQKCESQLKDLSTETDQAKRDLLERENQLEQLKKQQNQQVEELRQKLSAKEERISALQVEYENSLKDQGTKMEKIKQKAKEMQEMFKRKLSEQEAKLKKEFENKQLEFSQKEKHFNAKMLEMAHASSTGINDAVSKLEMNQKEQLESLAEAHRWELEEVAQTWGKKLNQQVEELKEKHEMELHKKEKEVGDLKQKLATFSAEKEGTNIEITLLKEEQVKRDESWNELQEQLKQLLAQVNTLSQSETDLKAQLEKLEDDLSHSLKDRAVLQEQLREQKTMEEKHKIKVTELTDMLKTSDEKLQILESSHCKEREDYEKKLEDKHLASQQKQIEFERLSSELAAQLDIYWKEADTLLQTKTNEVIEKCTEKINLVISRIAHCQHQTTKIKKAILIKNCKIPELETQLRQITEHQTTVSSSLQESTQQLQDKENVIILMKADIERLVTEKELLQKEGGHQQQAATEKETCITQLRKELSENINVVTSMREELKEKESEICALNKLVNELNVKLENSIGLTEKEAAISLLSKQHQEDQLQLLNQVQELSSRVESLSQEKASALEQVDHWMNKLSEWKKKAQLRFTQNHNTIKELQNKLELNNIQVGGKDEEIHQLKEELDKQSRNLDSLKDVMEQKQRRREKEEGKLTAELKIQTARITELEEHIARKTAENDSLMEELKRHNQQRDTEQKELVQQLQQTEVVVVEKDNRLKEAEEKVLSLEKQMDSMKAELDTKQREFEQMKTAMMKSKEEELKMLEDRMNAENTTKLADLKKKAEQKIASVKKQLTSQMEEKEQQFKLAMENQLKDLEQKMQEREAKIMDLEEKNRLIRDSADLEREMMQKVENEKASLEQEKASVLENVRQMYEEKISEIQKDLIDKDGLLNKYENEQQESNDSKLKMKSKQEELLKKLECTEKKLKEEQSVIERLQKEIEEKNKKYSLLLDQHTQDGDDLASIREELKAQVQKHYDMENVIGDFQKQFQEKDKVSQSLEQKIRELEDDIVKKKEVCKIEMAEVSLKYEEKLQGLQQQLDERNRQLKIFEENAEEKTRSDLELQKLLGDMQTQQKDLQAKLKEAETEKQKLRKEISKVQKDLRTLRKEHQQELDIMKKESLEEMEQKIRIEQEDIELKHNSTLKQLMREFNTQLSQKERELETAVHETISKAQDVETELIETHHAETTQLHRKIAEKDDDLKRTVKKYEEILEAREEEMTAKVNELQTQLEELQKEYKQKLADEENRRREEVTITELQIQLAQKTTIVNDSKLKEQEFIEQIHILKDQLKNYEKNVYVTAVGSPYRDGNLCHTDVSLFGEPTEFEYLRKVLFEYMMGRETKTMAKVITTVLKFPADQTQKILEREDARPLSWLRPS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0J9</accession>
    <name>A0A452J0J9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="19">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 16">Membrane</location>
            <topology evidence="1 16">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 16">Belongs to the integrin alpha chain family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J0J9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038322.1">
        <property type="protein sequence ID" value="ENSGAGP00000033833.1"/>
        <property type="gene ID" value="ENSGAGG00000023825.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009897">
        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008305">
        <property type="term" value="C:integrin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001525">
        <property type="term" value="P:angiogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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    <sequence length="1004" mass="111006" checksum="3EDD0BA55CDC3C4E" modified="2019-05-08" version="1">MARLRLLLLLLGRWLALGGAFNLDADGPAVYSGSEGSYFGFAVDFFAPDSSSIFLLVGAPKANTTQPDIVEGGQVLKCNWNSNKNCQPVIFDSTGDRNFAPNDPLEFKSHQWFGASVRSKNDKILACAPLYHWRTEVKQEREPVGTCYLHDGSKAVEYAPCRSTIIDADGQGFCQGGFSIDFTKQLSILMGFFPAGQLISDRVAEIVSKYNSKVYSIKYDDQLATRSASAAFDDSYLGYSVAVGEFSGDGIEDFVSGVPRAARTLGMVSIYNGKNMSSLYNFTGEQMAAYFGYSVAATDINGDNYTDLFIGAPLFMERGSDGKLQEVGQVSICLQQASGGFQIIKLNGFEIFARFSSSIAPLGDLDQDGFNDIAVAAPYGGEDKRGVVYIYNGRATGLNAVPSQILEGQWAARTMPPSFGYSLKGATDVDKNGYPDLIVGAFGVDTAVLYRARPVISVNAGLEVNPTILNPEEKICELPDTRVKVSCFNVKFCLTADGKGKLPNTLTFQVELLLDKLKQKGALRRALFLHNKQPGHSKNMTVVKGGTMRCEELQAFLRDESEFRDKLTPITVFMEYRLDYKTAKDSAGLQPILNQFTPANISRQTHILLDCGEDNICKPKLQVAVHSDQKQIYIGDDNPLTLIVDAQNQGEGAYEAELFVIIPPQADFIGVVRNNEVTRLSCAFKTENQTRLVVCDLGNPMKAGTKVSQIHAGLRFSVNQQSEMDTSVKFDLQIRSVSSPDHIFLPIANWQPQENPETEEDIGPLVQHIYELRNNGPSAFSKVMLNLQWPYKYNNNTLLYIMQYEIDGPMNCTSDMEINPLNIKISVPKADEKNETIRDEDNRDHHINRRDITAIEGDVHTLGCGTAECLKIVCQVGRLERGKSAILYVKSRLWTQTFMNKENQNHSYSLKSSASFNVIEFPYKNLSFEDIHNSTVVTTNITWGIQPQPMPVPVWVIILAVLAGLLLLAVLVFVMYRVSIIPWGRGGHTLGAGRGVPLLMYYCK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J0K0</accession>
    <name>A0A452J0K0_9SAUR</name>
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        <recommendedName>
            <fullName evidence="3">GRIP domain-containing protein</fullName>
        </recommendedName>
    </protein>
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        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
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        <property type="gene ID" value="ENSGAGG00000024101.1"/>
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    </dbReference>
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        <property type="entry name" value="GOLGIN SUBFAMILY A MEMBER 4"/>
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    <keyword evidence="1" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
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        <location>
            <begin position="2245"/>
            <end position="2292"/>
        </location>
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            <begin position="25"/>
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        </location>
    </feature>
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    </feature>
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        </location>
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        <location>
            <begin position="303"/>
            <end position="314"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="PROSITE" id="PS50913"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033885.1"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2307" mass="270395" checksum="65F8DE1471B85D35" modified="2019-05-08" version="1">MFKKLRQKIVEEQTQPRSPGGRAPQQPQTYSKSSPPSPGNRSRTSSLTEQSDEGTLTPDKELLAGMIAEPAFLSEYTIFALDPKRPKPQSDSVNVSKQTTPKSTDSNGSEPPSPQLSDTQTFAQKLQLRVPSMESLFRSPVKESLFRSSSKESLVRTSSRESLNRLDFDTPGPTFDPPSDVESEAEESLGNVDSLGKEQLLQRLRRMERSLGSYRGKYSELVTAYQTVQREKKKLQGILSQSQDKALRRIGELREELQMDQQAKKHLQEEFDASLEEKDQLISVLQTQVSLLKQRLQNGQISTELPETNTETEPQSPAKEISAENVLEPGGTEGDGDSAKTLETLIKRVKRQENLLQRCKEMIRSHKERSAQLTNEKEALQEQLEERLQELEKMKELHMTEKTKLITQLRDAKNLIEQLEQDKGMVIAETKRQMHETMEMKEEEIAQLRAHIKQITTQGEELREQKEKSEKAAFEELEKALSTAQKVEEARKKLQAEMDEKIKAIEKTSEEERVTLQQELTRVKQEVVEIMKKSSEERVAELEKLHKRELATKEQELNERLQVQGKEFQEKMRAALEKSQSEFLQTVQEKEQQESLALEELELQKKAIQSECEKKLQDMQQEIQTARTRILELESSLAKYSQDDKRQLEELSTLIESEKTKYNKEIIVMVEKHREELENVKQQEEELWTEKLQILKQQQQSTIEKLREKHEQEMDIMLKEKETVFRAHVEEMNAKTLEKLDVKQTELEALSFELSETLQIRHDLEKDGDKIKQESETKLEQQRKQYQEKMDMMVKEHDISVQGIQKAHTEETNQLKVQLEEKEKHLEELKAQEHKLKESLEISEAELKQVSAKLGDLCQSHQNNSSEQARVYEGQLTELQQKLTNMEAEKSHLKEQIVRAESQLNEVRTELDTYISRVRDLRQQLEEQSSENMQKITSLKQKCESQLKDLSTETDQAKRDLLERENQLEQLKKQQNQQVEELRQKLSAKEERISALQVEYENSLKDQGTKMEKIKQKAKEMQEMFKRKLSEQEAKLKKEFENKQLEFSQKEKHFNAKMLEMAHASSTGINDAVSKLEMNQKEQLESLAEAHRWELEEVAQTWGKKLNQQVEELKEKHEMELHKKEKEVGDLKQKLATFSAEKEGTNIEITLLKEEQVKRDESWNELQEQLKQLLAQVNTLSQSETDLKAQLEKLEDDLSHSLKDRAVLQEQLREQKTMEEKHKIKVTELTDMLKTSDEKLQILESSHCKEREDYEKKLEDKHLASQQKQIEFERLSSELAAQLDIYWKEADTLLQTKTNEVIEKCTEKINLVISRIAHCQHQTTKIKKAILIKNCKIPELETQLRQITEHQTTVSSSLQESTQQLQDKENVIILMKADIERLVTEKELLQKEGGHQQQAATEKETCITQLRKELSENINVVTSMREELKEKESEICALNKLVNELNVKLENSIGLTEKEAAISLLSKQHQEDQLQLLNQVQELSSRVESLSQEKASALEQVDHWMNKLSEWKKKAQLRFTQNHNTIKELQNKLELNNIQVGGKDEEIHQLKEELDKQSRNLDSLKDVMEQKQRRREKEEGKLTAELKIQTARITELEEHIARKTAENDSLMEELKRHNQQRDTEQKELVQQLQQTEVVVVEKDNRLKEAEEKVLSLEKQMDSMKAELDTKQREFEQMKTAMMKSKEEELKMLEDRMNAENTTKLADLKKKAEQKIASVKKQLTSQMEEKEQQFKLAMENQLKDLEQKMQEREAKIMDLEEKNRLIRDSADLEREMMQKVENEKASLEQEKASVLENVRQMYEEKISEIQKDLIDKDGLLNKYENEQQESNDSKLKMKSKQEELLKKLECTEKKLKEEQSVIERLQKEIEEKNKKYSLLLDQHTQDGDDLASIREELKAQVQKHYDMENVIGDFQKQFQEKDKVSQSLEQKIRELEDDIVKKKEVCKIEMAEVSLKYEEKLQGLQQQLDERNRQLKIFEENAEEKTRSDLELQKLLGDMQTQQKDLQAKLKEAETEKQKLRKEISKVQKDLRTLRKEHQQELDIMKKESLEEMEQKIRIEQEDIELKHNSTLKQLMREFNTQLSQKERELETAVHETISKAQDVETELIETHHAETTQLHRKIAEKDDDLKRTVKKYEEILEAREEEMTAKVNELQTQLEELQKEYKQKLADEENRRREEVTITELQIQLAQKTTIVNDSKLKEQEFIEQIHILKDQLKNYEKNVYVTAVGSPYRDGNLCHTDVSLFGEPTEFEYLRKVLFEYMMGRETKTMAKVITTVLKFPADQTQKILEREDARPLSTSSRSGIF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHQ2</accession>
    <name>A0A452GHQ2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">ELK domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Microtubule organizing center</location>
            <location evidence="1">Centrosome</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000001289.1">
        <property type="protein sequence ID" value="ENSGAGP00000001150.1"/>
        <property type="gene ID" value="ENSGAGG00000000844.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060090">
        <property type="term" value="F:molecular adaptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028745">
        <property type="entry name" value="AKAP9/Pericentrin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005539">
        <property type="entry name" value="ELK_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019528">
        <property type="entry name" value="PACT_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR44981:SF3">
        <property type="entry name" value="PERICENTRIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR44981">
        <property type="entry name" value="PERICENTRIN-LIKE PROTEIN, ISOFORM F"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10495">
        <property type="entry name" value="PACT_coil_coil"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01188">
        <property type="entry name" value="ELK"/>
        <property type="match status" value="9"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4 6" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="5" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="ELK" evidence="8">
        <location>
            <begin position="314"/>
            <end position="335"/>
        </location>
    </feature>
    <feature type="domain" description="ELK" evidence="8">
        <location>
            <begin position="470"/>
            <end position="491"/>
        </location>
    </feature>
    <feature type="domain" description="ELK" evidence="8">
        <location>
            <begin position="528"/>
            <end position="549"/>
        </location>
    </feature>
    <feature type="domain" description="ELK" evidence="8">
        <location>
            <begin position="596"/>
            <end position="619"/>
        </location>
    </feature>
    <feature type="domain" description="ELK" evidence="8">
        <location>
            <begin position="733"/>
            <end position="754"/>
        </location>
    </feature>
    <feature type="domain" description="ELK" evidence="8">
        <location>
            <begin position="762"/>
            <end position="780"/>
        </location>
    </feature>
    <feature type="domain" description="ELK" evidence="8">
        <location>
            <begin position="903"/>
            <end position="922"/>
        </location>
    </feature>
    <feature type="domain" description="ELK" evidence="8">
        <location>
            <begin position="1056"/>
            <end position="1078"/>
        </location>
    </feature>
    <feature type="domain" description="ELK" evidence="8">
        <location>
            <begin position="3017"/>
            <end position="3035"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1"/>
            <end position="47"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="70"/>
            <end position="102"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="3314"/>
            <end position="3339"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="117"/>
            <end position="195"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="220"/>
            <end position="340"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="376"/>
            <end position="441"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="487"/>
            <end position="668"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="695"/>
            <end position="729"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="758"/>
            <end position="981"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="1194"/>
            <end position="1295"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="1338"/>
            <end position="1462"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="1668"/>
            <end position="1695"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="1752"/>
            <end position="1936"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="1979"/>
            <end position="2017"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="2064"/>
            <end position="2139"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="2166"/>
            <end position="2249"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="2313"/>
            <end position="2417"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="2494"/>
            <end position="2570"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="2946"/>
            <end position="3093"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="3140"/>
            <end position="3174"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="3499"/>
            <end position="3574"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="1"/>
            <end position="13"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="72"/>
            <end position="87"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="SMART" id="SM01188"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001150.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="3803" mass="443389" checksum="6E1D2E4E34D9549C" modified="2019-05-08" version="1">MDEERRRKLEAGRAKLAHFRQRKTKGDNTQSQKKAAKRKGSSDYMQNIPKEECVIVAQDSDVLTDLNISVESKTEETETRSENKADVDLSNSNPNLVDDRASEEYDQLDVQMCRTRIMELEKRLLDKQEATEKLTAEVDDLLEQLAQHSEATHLQELETAVQERNEIIAQLTSNLQQARQNRDDIQEEASLLADQIHGLQLQLHEANELLKSRIPGKSELSQAQEQMAVFQNRLKEQSSQLQIMHQKAHDLELQLESSQKIEKLNAEIKDKVEKQQNLEEERKEQIGLIKKVHEREHDREISELIIKHEDEMKELRAELIKKQQQLLDELQQQMEGRHKTEVQHAQLQSQTLHNLELEALRLSLTNMHTSQLELMQSNLRKEKETALVELREMLNDKRAQEVAILQSRHQFELEHIDEQHLKEKEEIALKHQQELDEKKKIELEMEKEHIQTLETLNGDWALKTETCLKNVREELSEKHQTEMGELQKTLKLELDRVKEELKSLSLEKEQVEMKYENLENEHQKAITKLREQLQTEHDQCLENLKRKSKEREQEMQQEMEKLQATYEELKTQSQEEIKQLWSQLDSTRASRQELSELKEQLLARTSHVDEIERLKQDFEQQQQQRRNEHETELEQLRIYFEEKLGAAEENYREELTLLHQRLQELKEYSLLESEISQDQPLDISSSVTLLEEASEKERRDTLDQLTQQLEQHKEELTCLRLQMEEKHRHELDSLRSSLALQYKEDLMKMKMDLSDRYISEIEELKKKHCLDLEQLRAKLSEEHIKEITKLRLQSAQDAARQVEAEVAERVCVLDNEHKAKLSLLQSEKQHIAELQGEIQHLEREITKLKDIGVQHETQLKEEMEKIKCKLVDNHKNELKRAREEVQQMEQLHKEKEEEWKCEREALRIKAEEKLSQLRTELEDKAEYEKQTLQKKFELREAEMNQLQDQQAARIVELEKSLKEQQNSLQQLEDSLASMQTTQKAQAQYDSELQAARTLMEKDLEKATFELQEECAAKLMEAQNKFMEEHKVMTQKFTTEQEVLLQELKGKHADELELQSQQLQEKHKQQILSLTVELETKHQAETETLKSTLQSKQQAQLEAHIAELQAKHQAQIDELEAKHLSNLDSLESSYLSEIQIIRDEHKQVVEELQMHFTEQLQEKDKANQAILTQELERLKLKHSEELQLSQDNLKIELATIHIEKLKAMAAELEEAHKEDLNTALQNQRCMLEEESHKALDVLREEVLQMEEHHKKALKELQDLHVTEVQKQKEEYSWQLQEELEKLKAQHEHEEKMYASASTSEVETVRAALLAQFEEVKLKQQLQFQDEIELLKCQSEVLLEQQITQLKEEFEAERKTAWRSKEQVLIQETEKAQAHYKKEKEELSVQIQEKTNIIIQLEKKVESLNHELEETNCELETLLQRRDRENQEGEYLVAMLRSDIELSQNERKKLQESYQQILKLFVEMVKATIATEDLICKKVGLCLDDSLASGDSRESRSMMEEVGFIQHFKAREKHDLEKSDLLDETLTEHNQISAVTDEGSELSQHLCESVFASPDMAFENEEMILKICRRLRTAVEKLLELVTESTKQLEKTHEIHAHFEEEFSRRTQETAQVVNQHHDLMECLNEESEAKNQLALELHKAEGIIEGYVVEKAALEEAMSLKEESERRLVLELESLQEQFQKLTQEQAILREERDMLISQKKALAANVGEIEVGVPVANVAKKEDSGLLKEVEFLAKEKLELQCQAEKDHSTLRSQMKALEVELEEQMDQNQKLAKKSLEVTDLRQQIQVLEKQLRSQRQFMDEQAIEREHERDEFQQEIQKLEDQLKLSGKSQTSGEPKQYGIFELTLQIESLQAEIKDKTDDYNKLLLEKEQKYREVTVQDKEIEELAARIRELELTNTEASKTVSRLEQELQKMKKRETELTQDKEALQQQQYNNLIQISALQSKLDEARHRVPSEGSSDHGLKEQLQAEQEALLLKEREILSLLEQLEQFKENLINKNEEILRLNLQLEMQKNLAAGIGQLQVENVHLKEDIAKLHMMQSQNVGNSESAVLQFPQALLEEKNQEIDHLNEQIERLQRELEGATDNKVVENGKSEIDELRLLVEHLRGDQERLHRDKAEEVEQLHGVIEKLQKELVQLEPMYHEVSDSQDNLNLLWLEKQAENLQNELKKGSLDFQEHSDKNKKAVLLHSKLKELQEELELVSTAREALQQLLEEKESQFKMEVEILEQNLQNVQESSRQHLAELTSLKVQHDAFQEEYSLLKTLLSQRDGEIGIMSTRIQELEDTLRAREALLLESDLQIKTVAEQRVVEAAELQHLTEKIARLEDELLKKDLHQKTEAEEVQTLQSEVSRLDFKVQTLNQKEVVYQREIDELQANAIKLKDQIQEFLKELQALRSERDDLYSQLESYKEKDQSNDQEAKLHKSVFKQEEVLVHTNGMERLGEERGANADQSFAASVSQSDKLILVQLEATNTSENHKDLITKMTLHQEELKSELACVKKELELSEEQTGKILEDIKEKDAAIADLKTHSRNLKTQIKQLQETLLTQEAEMTDKARELQDLKKQCQQILEFHQTPDSMKCKLNNLSQNTEHDKESPKDFKCLVMDMTSWDSPEIVRKQETSIELQPSLHLTPFSEVDSVHSTDFEMKHNKSSVVQGDTPGLLGYQGLYANTNEEAVVRLHLKSPAFSESTYSIPEYQNTDKRIPVRDVDNFTLTPSDSQDDLRSTMSGLEGASDGYVSNASSDLAAKLNQELETSEHLDASFLAYLQYCGIFQDAMDPVTEKETISPQLKSVLKMVYEDSRRILALSEHPFPLSDQKSIQQSAAAEGWRKEGLILLDAIQSLRDYLSKVEDRGDKESSDTSFDWRGELLQAVQSVLEKERNMFQIDLQSHLCNPGSGDEGSLVEKLEHIVKRQEEQQRTVLEHLLSSDRNSLLSEIQDLRAQLRMTHLQNQEKLQQLQETLTNAEDRGSKQEHQLRRKVELLEYKLQQEKTIANDLQNSLSEEQERASEMHELLKQERSVVSDLKSELSESKEENESLQTFLQKLQREILQLRSELENKEKDLTVALQDLQNEHSKQKELRNIFEEQHLQHKLREDEKTKALEELQAALELQCIQNNQMSVALEHEKSANDNLRKELQIEHSRCEALLSQERNKLTELQSNLEVEKNHSLELLNALNHERVLTEQLSMRGNEDSSCQHKESLVEQTFVQELQAQLDAERSRAMELAAIIEKTHQQAIQSKRQLEAEVQMCCEETQKEREISTKLRATLESLQSQKQEVIRSLEIQREREAKLKVDWEQLQTLFRTMKEQEKRKKEEREKERREEQRAEFEKRTEWQKDKERVHELELQHQRDEHRIKELQQTLAELEKQERNLASQKSQRELSSCPIKNDYNANLSLTTETEALHLQQQQLEKIRQQLLFAAVHLNEFIYKTVDKTVNDWSASNDEAIASLLHTLKELKSELLTSSTPLKPLQGSVSLVDLLLKENGELTKSVTTLTEEKLELRAVISQLEKNLQQHHRRGISNRQIRSTDDAHSVQESERASWQREKTILQNALKQAESELAKATVEIENKPVMETSNPKLQKLYKKYLRAESFRKALVYQKKYLLLLLGGFQDCEQATLSLIARMGVYPSPADLQVSESRSRPFTKFRSAVRVVIAVSRLKFLVKKWHKVNRKGAQGENISHSIGHNPVSGARTEVLRQQQLPSVNVNSPPTRDIGSCHRTSSARFVNHSPKSSYWLHNRLHPSTSSVSEKSLVSTQDPERSLTEYIHRLEVIQQRLGGAQIGSTPGRPHQRNIK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHQ7</accession>
    <name>A0A452GHQ7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Uridylate-specific endoribonuclease</fullName>
            <ecNumber evidence="11">4.6.1.-</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>ribonucleotidyl-uridine-RNA = a 5'-end dephospho-uridine-RNA + a 3'-end 2',3'-cyclophospho-ribonucleotide-RNA</text>
            <dbReference type="Rhea" id="RHEA:67792"/>
            <dbReference type="Rhea" id="RHEA-COMP:10464"/>
            <dbReference type="Rhea" id="RHEA-COMP:17354"/>
            <dbReference type="Rhea" id="RHEA-COMP:17356"/>
            <dbReference type="ChEBI" id="CHEBI:83064"/>
            <dbReference type="ChEBI" id="CHEBI:173117"/>
            <dbReference type="ChEBI" id="CHEBI:173224"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 11">
            <name>Mn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29035"/>
        </cofactor>
    </comment>
    <comment type="subunit">
        <text evidence="3 11">Monomer.</text>
    </comment>
    <comment type="similarity">
        <text evidence="2 11">Belongs to the ENDOU family.</text>
    </comment>
    <dbReference type="EC" id="4.6.1.-" evidence="11"/>
    <dbReference type="AlphaFoldDB" id="A0A452GHQ7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001294.1">
        <property type="protein sequence ID" value="ENSGAGP00000001155.1"/>
        <property type="gene ID" value="ENSGAGG00000000910.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016787">
        <property type="term" value="F:hydrolase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016829">
        <property type="term" value="F:lyase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004521">
        <property type="term" value="F:RNA endonuclease activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="CDD" id="cd21159">
        <property type="entry name" value="XendoU"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039787">
        <property type="entry name" value="ENDOU"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037227">
        <property type="entry name" value="EndoU-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018998">
        <property type="entry name" value="EndoU_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12439">
        <property type="entry name" value="PLACENTAL PROTEIN 11-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12439:SF11">
        <property type="entry name" value="URIDYLATE-SPECIFIC ENDORIBONUCLEASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09412">
        <property type="entry name" value="XendoU"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF142877">
        <property type="entry name" value="EndoU-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51959">
        <property type="entry name" value="ENDOU"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6 11" id="KW-0255">Endonuclease</keyword>
    <keyword evidence="7 11" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="10" id="KW-0456">Lyase</keyword>
    <keyword evidence="9 11" id="KW-0464">Manganese</keyword>
    <keyword evidence="5 11" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="4 11" id="KW-0540">Nuclease</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8 11" id="KW-0694">RNA-binding</keyword>
    <feature type="domain" description="EndoU" evidence="12">
        <location>
            <begin position="10"/>
            <end position="287"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001936"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010168"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00011245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022759"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023211"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="RuleBase" id="RU367085"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS51959"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001155.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="294" mass="34227" checksum="C6648D1C2C7D44F1" modified="2019-05-08" version="1">MLEGIQDRELNHELSKLFNELWDADVNRFVPGKDYTIDLQEKAGFVQQSSGVARDSASEPLFRYVNEERLKSIKTFAAFVSLLDNYETSTGVAEVVTPQEMAENNRFLDAILETEVMKLTHQYLVRKNWSKPSLNDFKSQLYDIWFQLYTRETGKGPDSCGFEHVFVGETKHGKEILGLHNWVQFYLQEKHKQVDYKGYIARRKKSRPDEDDQVLSLQFSWKGLVKPVGSTFIGVSPEFEFALYTIIFLQSQEKVTREMVRLEEYELQIVVYRHGKYIGTAYPVLLSSNNEDLY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHR0</accession>
    <name>A0A452GHR0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Protein AMBP</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">In the N-terminal section; belongs to the calycin superfamily. Lipocalin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHR0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001252.1">
        <property type="protein sequence ID" value="ENSGAGP00000001117.1"/>
        <property type="gene ID" value="ENSGAGG00000000433.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004867">
        <property type="term" value="F:serine-type endopeptidase inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd22596">
        <property type="entry name" value="Kunitz_bikunin_1-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd22597">
        <property type="entry name" value="Kunitz_bikunin_2-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.410.10:FF:000010">
        <property type="entry name" value="Alpha1-microglobulin/bikunin (AMBP)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.128.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.410.10">
        <property type="entry name" value="Pancreatic trypsin inhibitor Kunitz domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002968">
        <property type="entry name" value="A1-microglobln"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029856">
        <property type="entry name" value="AMBP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012674">
        <property type="entry name" value="Calycin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002223">
        <property type="entry name" value="Kunitz_BPTI"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036880">
        <property type="entry name" value="Kunitz_BPTI_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000566">
        <property type="entry name" value="Lipocln_cytosolic_FA-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020901">
        <property type="entry name" value="Prtase_inh_Kunz-CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46676">
        <property type="entry name" value="PROTEIN AMBP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46676:SF1">
        <property type="entry name" value="PROTEIN AMBP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00014">
        <property type="entry name" value="Kunitz_BPTI"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00061">
        <property type="entry name" value="Lipocalin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01215">
        <property type="entry name" value="A1MCGLOBULIN"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00759">
        <property type="entry name" value="BASICPTASE"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00179">
        <property type="entry name" value="LIPOCALIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00131">
        <property type="entry name" value="KU"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57362">
        <property type="entry name" value="BPTI-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50814">
        <property type="entry name" value="Lipocalins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00280">
        <property type="entry name" value="BPTI_KUNITZ_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50279">
        <property type="entry name" value="BPTI_KUNITZ_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0157">Chromophore</keyword>
    <keyword evidence="5" id="KW-0165">Cleavage on pair of basic residues</keyword>
    <keyword evidence="11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6" id="KW-0646">Protease inhibitor</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="9" id="KW-0722">Serine protease inhibitor</keyword>
    <keyword evidence="7 13" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="13">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019367664" description="Protein AMBP" evidence="13">
        <location>
            <begin position="21"/>
            <end position="341"/>
        </location>
    </feature>
    <feature type="domain" description="BPTI/Kunitz inhibitor" evidence="14">
        <location>
            <begin position="220"/>
            <end position="270"/>
        </location>
    </feature>
    <feature type="domain" description="BPTI/Kunitz inhibitor" evidence="14">
        <location>
            <begin position="276"/>
            <end position="326"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008238"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00018905"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022685"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022690"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50279"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001117.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="341" mass="38066" checksum="43700467D52EBC0A" modified="2019-05-08" version="1">MKFQEFFFLLFLAYLPTARGSPTGSQVQDIQVQENFEPERVYGKWYDIATGTTCKWMKQYKDKFNMGTLVLGPGMTSNQISTTITQNTGTQLGYTVEVPTGGCVCISFSAGWDVTINSYVVHTNYEEYAIILMQKKSSFGLTITAKLYGRSPELREGLIVDFRQFAQEMGIPEDSIFIMINKGNPAMMRVTRDRRAVLPEQEGSAEGPLLPLISKKEDFCRLNKDVGPCMGMNVRFFYNSSSMACETFHYGGCLGNGNNFLSEKKCLQTCRTEAACRLPIVPGPCQQPVILWAFDATQGKCTSFNYGGCQGNGNKFYTEKECREYCGVPADGMVRPVLSQS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHT4</accession>
    <name>A0A452GHT4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Paired domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHT4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001335.1">
        <property type="protein sequence ID" value="ENSGAGP00000001189.1"/>
        <property type="gene ID" value="ENSGAGG00000000896.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00131">
        <property type="entry name" value="PAX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.10:FF:000013">
        <property type="entry name" value="Paired box 8 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.10:FF:000003">
        <property type="entry name" value="Paired box protein Pax-6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.10">
        <property type="entry name" value="Winged helix-like DNA-binding domain superfamily/Winged helix DNA-binding domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009057">
        <property type="entry name" value="Homeodomain-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043182">
        <property type="entry name" value="PAIRED_DNA-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001523">
        <property type="entry name" value="Paired_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043565">
        <property type="entry name" value="PAX_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036388">
        <property type="entry name" value="WH-like_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45636:SF20">
        <property type="entry name" value="PAIRED BOX PROTEIN PAX-5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45636">
        <property type="entry name" value="PAIRED BOX PROTEIN PAX-6-RELATED-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00292">
        <property type="entry name" value="PAX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00027">
        <property type="entry name" value="PAIREDBOX"/>
    </dbReference>
    <dbReference type="SMART" id="SM00351">
        <property type="entry name" value="PAX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46689">
        <property type="entry name" value="Homeodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00034">
        <property type="entry name" value="PAIRED_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51057">
        <property type="entry name" value="PAIRED_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="5" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="3" id="KW-0563">Paired box</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0804">Transcription</keyword>
    <keyword evidence="4" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="Paired" evidence="9">
        <location>
            <begin position="7"/>
            <end position="133"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="134"/>
            <end position="153"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="172"/>
            <end position="209"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="247"/>
            <end position="295"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="134"/>
            <end position="150"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022724"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51057"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001189.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="306" mass="33453" checksum="3DD4E9ABF6027763" modified="2019-05-08" version="1">QDLNKNRHGGVNQLGGVFVNGRPLPDVVRQRIVELAHQGVRPCDISRQLRVSHGCVSKILGRYYETGSIKPGVIGGSKPKVATPKVVEKIAEYKRQNPTMFAWEIRDRLLAERVCDNDTVPSVSSINRIIRTKVQQPPNQQVPASSHSIASTGSVTQVSTVSTDSAGSSYSISGILGITSPSAESNKRKRDEGIQESPVPNGHSLPGRDFLRKQMRGDLFTQQQLEVLDRVFERQHYSDIFTTTEPIKPEQLPAPAAIADGAHLGPPKRGPPSPADMKPPEMGSVSGSSLRPKEINYKQNIEDINC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHT5</accession>
    <name>A0A452GHT5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Peptidase S1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GHT5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001281.1">
        <property type="protein sequence ID" value="ENSGAGP00000001142.1"/>
        <property type="gene ID" value="ENSGAGG00000000827.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00190">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.10:FF:000005">
        <property type="entry name" value="Serine protease 37"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.10.10">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009003">
        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043504">
        <property type="entry name" value="Peptidase_S1_PA_chymotrypsin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001314">
        <property type="entry name" value="Peptidase_S1A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001254">
        <property type="entry name" value="Trypsin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018114">
        <property type="entry name" value="TRYPSIN_HIS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271:SF81">
        <property type="entry name" value="GRANZYME B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271">
        <property type="entry name" value="KALLIKREIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00089">
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00722">
        <property type="entry name" value="CHYMOTRYPSIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00020">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50494">
        <property type="entry name" value="Trypsin-like serine proteases"/>
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    </dbReference>
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        <property type="entry name" value="TRYPSIN_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00134">
        <property type="entry name" value="TRYPSIN_HIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0732">Signal</keyword>
    <keyword evidence="2" id="KW-0865">Zymogen</keyword>
    <feature type="domain" description="Peptidase S1" evidence="4">
        <location>
            <begin position="17"/>
            <end position="241"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023145"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001142.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="243" mass="26991" checksum="9D29BA83E474EDCF" modified="2019-05-08" version="1">MPYLGSSPRYVLAKGDIIGGREARPHSRPYMAFVETKMGPESTTECGGFLVREDFVLTAAHCDEEGSGDAPTVVILGAHDLNRKERTQQEIQVRRKIPHEDFDRTTLENDIMLLQLELRAEWNNAVKPIRLPPANFLLKPEATCSVAGWGRTGVHMVRPATKLQEVTLKMADDKQCELFFDSVHGLHGEHQNEAVIGDTGGPLVCNGFAQSIVSYGKDSGLPSAVYTRMSAFIPWLRKKMGRE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHT7</accession>
    <name>A0A452GHT7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Reticulon</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="6">Inhibits amyloid precursor protein processing, probably by blocking BACE1 activity.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 7">Endoplasmic reticulum membrane</location>
            <topology evidence="1 7">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHT7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001333.1">
        <property type="protein sequence ID" value="ENSGAGP00000001187.1"/>
        <property type="gene ID" value="ENSGAGG00000000953.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043005">
        <property type="term" value="C:neuron projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0014069">
        <property type="term" value="C:postsynaptic density"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007420">
        <property type="term" value="P:brain development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071787">
        <property type="term" value="P:endoplasmic reticulum tubular network formation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030182">
        <property type="term" value="P:neuron differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.2480:FF:000001">
        <property type="entry name" value="Reticulon"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.2480">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003388">
        <property type="entry name" value="Reticulon"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046964">
        <property type="entry name" value="RTN1-4"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45799:SF5">
        <property type="entry name" value="RETICULON-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45799">
        <property type="entry name" value="RETICULON-LIKE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02453">
        <property type="entry name" value="Reticulon"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50845">
        <property type="entry name" value="RETICULON"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3 7" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="5 7" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 7" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="77"/>
            <end position="106"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="192"/>
            <end position="213"/>
        </location>
    </feature>
    <feature type="domain" description="Reticulon" evidence="9">
        <location>
            <begin position="63"/>
            <end position="250"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1"/>
            <end position="52"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="13"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="24"/>
            <end position="37"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004477"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00037306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU210713"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50845"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001187.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="250" mass="27548" checksum="34A64413E1E15417" modified="2019-05-08" version="1">MSAPPDRQGLPLSGSGARWAGAGAEERGAGAQRRDGDGAQQPQLPAGSAREHPPVTMATASTAIDLLYWRDIKQTGIVFGSILLLLFSLTQFSVVSVIAYLALAALSATISFRIYKSVLQAVQKTDEGHPFKAYLEMEMSLSQEQIQKYTDCLQLYVNSTVRELRRLFLVQDLVDSLKFAVLMWLLTYVGALFNGLTLMIMAVVSMFTLPVVYDKYQAQIDQYLGLVRTHINTVMAKIQAKIPGAKRKAE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHU3</accession>
    <name>A0A452GHU3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Olduvai domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Golgi apparatus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000001282.1">
        <property type="protein sequence ID" value="ENSGAGP00000001143.1"/>
        <property type="gene ID" value="ENSGAGG00000000835.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060090">
        <property type="term" value="F:molecular adaptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007098">
        <property type="term" value="P:centrosome cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090063">
        <property type="term" value="P:positive regulation of microtubule nucleation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903358">
        <property type="term" value="P:regulation of Golgi organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056273">
        <property type="entry name" value="CDK5RAP2_MYOME_CC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012943">
        <property type="entry name" value="Cnn_1N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052593">
        <property type="entry name" value="MT-associated_AKAP9-binding"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010630">
        <property type="entry name" value="Olduvai_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46501">
        <property type="entry name" value="MYOMEGALIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46501:SF2">
        <property type="entry name" value="MYOMEGALIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23246">
        <property type="entry name" value="CC_CDK5RAP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07989">
        <property type="entry name" value="Cnn_1N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01148">
        <property type="entry name" value="DUF1220"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51316">
        <property type="entry name" value="ODV"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="5" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Olduvai" evidence="9">
        <location>
            <begin position="1430"/>
            <end position="1521"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="871"/>
            <end position="918"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1190"/>
            <end position="1211"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1271"/>
            <end position="1297"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1465"/>
            <end position="1586"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1957"/>
            <end position="1998"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="2183"/>
            <end position="2231"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="41"/>
            <end position="68"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="94"/>
            <end position="150"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="236"/>
            <end position="291"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="350"/>
            <end position="386"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="418"/>
            <end position="463"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="492"/>
            <end position="577"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="679"/>
            <end position="821"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="939"/>
            <end position="966"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="1132"/>
            <end position="1159"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="1240"/>
            <end position="1267"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="1776"/>
            <end position="1856"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="2124"/>
            <end position="2158"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="901"/>
            <end position="910"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="1192"/>
            <end position="1203"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="1473"/>
            <end position="1490"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="1514"/>
            <end position="1525"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="1529"/>
            <end position="1549"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="1556"/>
            <end position="1571"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="1957"/>
            <end position="1966"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="1977"/>
            <end position="1995"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="2208"/>
            <end position="2220"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004555"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51316"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001143.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2250" mass="252066" checksum="7F03F855772118B2" modified="2019-05-08" version="1">MSNGYRTLSQHLNDLKKENFSLKLRIYFLEERMQQKYEASREDVYKRNIELKVEVESLKQELQERQQTLDKAWAAAENLTSCNEAEFQRRYEERQQEKEHVSELLENKIQLLQEEARLAKSEAERTAALVEAEKGRCLELTKKLKELTKEEEKRGAQVLRDDHCAALAQKDKHIEELIQELSAREQLIELLATEKQNLLQRLEEPREIEVQNLPEDHQQLTQCDPVASGRVSESHAAELQDKIQQYDATNKLLQEKLNEMNFELKSIQEASQRQDRTIQSLNEALKSKESETEELYHVIEGQNETMGKLRDMLHRSQLGHLQTSEGPSPSQQQQHMALLDLQNTLFCTQLEMQKLKRAQWQKEHQLAEARRTTQLLEATVQEEQQLKEVSWKHNQESWELLQRRQSLEKNPVVTDTMLEKLQQRIKHRDAALERAIDEKFCAVEEKEQELQQLHISVREREHDLERLRSVLSSNEATIHSMESLLKAKTLELEQVTATCQNLQWLKEEVEAKSNCWQKEQEGIIQQLQASLHDRNKEVEELTATLLCKLGPGQSEIAEELCLRLQRKERMLQDLLSDRNRQAGEHETELRELLQAMSIREQRSRVTAEKMVQALAEKSDEVQFLRQRLVGNEPGKNPVADGRLLLQKDKQPFQELLPGGCGNATATRISKGEDSKYRMERGLLETAAELEKELINAREELELMARKERESRLELSSLQSVVTAQEEELQVQASDIESLTRNIQSKEDLIKDLQMQLVDPEEIPAVERLTQEVLVLREKVATAESRGQEATGNRQQQLLLMLEGLVAEKSRLNEALQAERQLYSSLVKFHAHPDSPERGQTLQAELEGAQALRGRLEESLGRSMEHLRRLETLGAFGGQPGRDDAKDGSTDSIEGEAPPRTVNHQTSQGLPANSPEAKAAQPDLLTLAPLERDSSLQAELLHVRAKNQQLLEQKAKVEGELWELKALMEEAGFSSLSSIRKVLLSLCLETAELKEQLGGSPLSEGWEDKAEARAALCTQNSKLQSSETVVTLLKEQLVLNCPEGESKFPPQLVTGMATEIDRLWTEMGVASGKRPALEGQLQEDPAKRPCPRSQAVELGPAQAPRGMSQQMEACPAGSWQRLVEPRVGPSGQLVQCRQRNQELQDKLAVSEATVRAQAEQLEQYRTLLSEPLVQQDNKQVQVDLQDLGYETCGRSENEADREETPSPECEEPDVFSKAGLMEELSRPGPWGGLCWGQSEDVAVLQQHVRELQLQLQNSNKALQRLQRRTRSFSSTSDYASGAERLHKPVPSATDEDEGWQSDGVGAFCPPALQASRDLEWLVHRVSLLEAQLHGRWAGGVLPEELRSAVWPGKYDSLIQAQARELSHLRQKMREGRSVCHLLTQHLRDTVKSFEELLRGTDIDYYMGQSFREHLAQGGQLAERLSSKLSSRDRSDVEDKTGHEFVALRLSKELREKERMIETLQAKLQERCETPSSSRALSESPRSNSSASFLSDGPEACSDGDATSEYSQLQGDRSEQPSRRLTDVTHPAALASNPSPAAAPHQASAESIRGYPTGPSSQHPPMDSGQTRTGLHFDSLSKPLSVPLTPAPGPSAFLPFGPPPPTPSALLGCCGTPVFSLAEVQQELHMLQKQLGERGPLAAPPVKPLPLTSDFGEASTSRCLHVPHLTPQSCPLQSSTQLDRKAGAAFLESNPLWAQPPMGALYGHLPSGYLASHKLTGADLLEGHLAEIRSLRQRLEESICINDRLREQLEKRLATTTKGNGYPTNIYIQGLESASSLSSETQALREENQSLQLQFAHLSRELEQLRETTRCRLQKAETENRTLAQELAEQRQSGRQLREERLALQENNHRLECTVPLLQQQCQENQQLCQALCAELRIHETLSSSSQAALSACSGDVHRRPPSSSELGALLAEVRTLRRKLLRGIQVNSALRQQLARQQPEGSAAPLFAAHQATPNWQPFQDSSPSPPVRDVGMSSPAPLSPPALLSGPAPLAHQLKEPPVDDALVRRSAPAPAGDAPSSSSASKGGCQVIGHVDDYYTLKQLILEGKTLVHQMASLLRPALAMPSLEPRGTKALDRGSVQQLLSTASALHQLLEQCSSLLTTFWRGALPAAPSPAQPQTAEQAMKDELLALRARLAEQENLLQRAAERLQDTCRLKDNMEHFIVSHLTQTHAVLRKARTNLEKSKPQGSSVKSPPFIGAGEISVGLSQHPSHVIPTNQEHRRKKSCPDTLKWQEDQSWPCFVHLPSS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHU7</accession>
    <name>A0A452GHU7_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="1">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="1 2">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="2" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="1" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="1" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GHU7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001346.1">
        <property type="protein sequence ID" value="ENSGAGP00000001198.1"/>
        <property type="gene ID" value="ENSGAGG00000000959.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043247">
        <property type="entry name" value="CCDC183"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47115">
        <property type="entry name" value="COILED-COIL DOMAIN-CONTAINING PROTEIN 183"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47115:SF1">
        <property type="entry name" value="COILED-COIL DOMAIN-CONTAINING PROTEIN 183"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000313" key="1">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001198.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="206" mass="23991" checksum="164B81953F72341C" modified="2019-05-08" version="1">MNVHNVLLYELKRRGGMVEGLQRKLQHLHSHVPGSPVTHEPSLLQVIRQLENNIEKMLMKIRTGEKIYYLYLKMVDFLKDELAQLPLQLNVLQHMVEVYQVELKGMRLMAVDATEATEAAKMDMTNMETEFIAERRFRENSLNFQKKQIDRIRVKDASERHRRVVSPRELRGPDLLGWLCPEASLAWRGPGASFPPMLFYDSLGCF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHU9</accession>
    <name>A0A452GHU9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Uridylate-specific endoribonuclease</fullName>
            <ecNumber evidence="11">4.6.1.-</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>ribonucleotidyl-uridine-RNA = a 5'-end dephospho-uridine-RNA + a 3'-end 2',3'-cyclophospho-ribonucleotide-RNA</text>
            <dbReference type="Rhea" id="RHEA:67792"/>
            <dbReference type="Rhea" id="RHEA-COMP:10464"/>
            <dbReference type="Rhea" id="RHEA-COMP:17354"/>
            <dbReference type="Rhea" id="RHEA-COMP:17356"/>
            <dbReference type="ChEBI" id="CHEBI:83064"/>
            <dbReference type="ChEBI" id="CHEBI:173117"/>
            <dbReference type="ChEBI" id="CHEBI:173224"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 11">
            <name>Mn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29035"/>
        </cofactor>
    </comment>
    <comment type="subunit">
        <text evidence="3 11">Monomer.</text>
    </comment>
    <comment type="similarity">
        <text evidence="2 11">Belongs to the ENDOU family.</text>
    </comment>
    <dbReference type="EC" id="4.6.1.-" evidence="11"/>
    <dbReference type="AlphaFoldDB" id="A0A452GHU9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001297.1">
        <property type="protein sequence ID" value="ENSGAGP00000001158.1"/>
        <property type="gene ID" value="ENSGAGG00000000910.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016787">
        <property type="term" value="F:hydrolase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016829">
        <property type="term" value="F:lyase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
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    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004521">
        <property type="term" value="F:RNA endonuclease activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
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        <property type="entry name" value="XendoU"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039787">
        <property type="entry name" value="ENDOU"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037227">
        <property type="entry name" value="EndoU-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018998">
        <property type="entry name" value="EndoU_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12439">
        <property type="entry name" value="PLACENTAL PROTEIN 11-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12439:SF11">
        <property type="entry name" value="URIDYLATE-SPECIFIC ENDORIBONUCLEASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09412">
        <property type="entry name" value="XendoU"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF142877">
        <property type="entry name" value="EndoU-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51959">
        <property type="entry name" value="ENDOU"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6 11" id="KW-0255">Endonuclease</keyword>
    <keyword evidence="7 11" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="10" id="KW-0456">Lyase</keyword>
    <keyword evidence="9 11" id="KW-0464">Manganese</keyword>
    <keyword evidence="5 11" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="4 11" id="KW-0540">Nuclease</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8 11" id="KW-0694">RNA-binding</keyword>
    <feature type="domain" description="EndoU" evidence="12">
        <location>
            <begin position="5"/>
            <end position="282"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001936"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010168"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00011245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022759"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023211"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="RuleBase" id="RU367085"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS51959"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001158.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="289" mass="33431" checksum="AC16B2DBF572AD74" modified="2019-05-08" version="1">MDLNCSKGNLGWTLKKLPSDVNRFVPGKDYTIDLQEKAGFVQQSSGVARDSASEPLFRYVNEERLKSIKTFAAFVSLLDNYETSTGVAEVVTPQEMAENNRFLDAILETEVMKLTHQYLVRKNWSKPSLNDFKSQLYDIWFQLYTRETGKGPDSCGFEHVFVGETKHGKEILGLHNWVQFYLQEKHKQVDYKGYIARRKKSRPDEDDQVLSLQFSWKGLVKPVGSTFIGVSPEFEFALYTIIFLQSQEKVTREMVRLEEYELQIVVYRHGKYIGTAYPVLLSSNNEDLY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHW1</accession>
    <name>A0A452GHW1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Protein AMBP</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">In the N-terminal section; belongs to the calycin superfamily. Lipocalin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHW1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001307.1">
        <property type="protein sequence ID" value="ENSGAGP00000001165.1"/>
        <property type="gene ID" value="ENSGAGG00000000433.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004867">
        <property type="term" value="F:serine-type endopeptidase inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd22596">
        <property type="entry name" value="Kunitz_bikunin_1-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd22597">
        <property type="entry name" value="Kunitz_bikunin_2-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.410.10:FF:000010">
        <property type="entry name" value="Alpha1-microglobulin/bikunin (AMBP)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.128.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.410.10">
        <property type="entry name" value="Pancreatic trypsin inhibitor Kunitz domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002968">
        <property type="entry name" value="A1-microglobln"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029856">
        <property type="entry name" value="AMBP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012674">
        <property type="entry name" value="Calycin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002223">
        <property type="entry name" value="Kunitz_BPTI"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036880">
        <property type="entry name" value="Kunitz_BPTI_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000566">
        <property type="entry name" value="Lipocln_cytosolic_FA-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020901">
        <property type="entry name" value="Prtase_inh_Kunz-CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46676">
        <property type="entry name" value="PROTEIN AMBP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46676:SF1">
        <property type="entry name" value="PROTEIN AMBP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00014">
        <property type="entry name" value="Kunitz_BPTI"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00061">
        <property type="entry name" value="Lipocalin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01215">
        <property type="entry name" value="A1MCGLOBULIN"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00759">
        <property type="entry name" value="BASICPTASE"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00179">
        <property type="entry name" value="LIPOCALIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00131">
        <property type="entry name" value="KU"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57362">
        <property type="entry name" value="BPTI-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50814">
        <property type="entry name" value="Lipocalins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00280">
        <property type="entry name" value="BPTI_KUNITZ_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50279">
        <property type="entry name" value="BPTI_KUNITZ_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0157">Chromophore</keyword>
    <keyword evidence="5" id="KW-0165">Cleavage on pair of basic residues</keyword>
    <keyword evidence="11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6" id="KW-0646">Protease inhibitor</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="9" id="KW-0722">Serine protease inhibitor</keyword>
    <keyword evidence="7 13" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="13">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019466698" description="Protein AMBP" evidence="13">
        <location>
            <begin position="21"/>
            <end position="324"/>
        </location>
    </feature>
    <feature type="domain" description="BPTI/Kunitz inhibitor" evidence="14">
        <location>
            <begin position="203"/>
            <end position="253"/>
        </location>
    </feature>
    <feature type="domain" description="BPTI/Kunitz inhibitor" evidence="14">
        <location>
            <begin position="259"/>
            <end position="309"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008238"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00018905"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022685"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022690"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50279"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001165.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="324" mass="36112" checksum="CE31E14F751A6CF0" modified="2019-05-08" version="1">MKFQEFFFLLFLAYLPTARGSPTGSQVQDIQVQENFEPERVYGKWYDIATGTTCKWMKQYKDKFNMGTLVLGPGMTSNQISTTITQNTGTQLGYTVHPGDSAAAVSSWDVTINSYVVHTNYEEYAIILMQKKSSFGLTITAKLYGRSPELREGLIVDFRQFAQEMGIPEDSIFIMINKGECVPSETQPEPKVSSTLLMCSDFCRLNKDVGPCMGMNVRFFYNSSSMACETFHYGGCLGNGNNFLSEKKCLQTCRTEAACRLPIVPGPCQQPVILWAFDATQGKCTSFNYGGCQGNGNKFYTEKECREYCGVPADGMVRPVLSQS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHW2</accession>
    <name>A0A452GHW2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Nephronectin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">Functional ligand of integrin alpha-8/beta-1 in kidney development. Regulates the expression of GDNF with integrin alpha-8/beta-1 which is essential for kidney development. May also play a role in the development and function of various tissues, regulating cell adhesion, spreading and survival through the binding of several integrins.</text>
    </comment>
    <comment type="subunit">
        <text evidence="14">Homodimer and homotrimer.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the nephronectin family.</text>
    </comment>
    <comment type="caution">
        <text evidence="16">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHW2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001361.1">
        <property type="protein sequence ID" value="ENSGAGP00000001213.1"/>
        <property type="gene ID" value="ENSGAGG00000000964.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00054">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="CDD" id="cd06263">
        <property type="entry name" value="MAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000038">
        <property type="entry name" value="Fibrillin 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000187">
        <property type="entry name" value="nephronectin isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000476">
        <property type="entry name" value="nephronectin isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.200:FF:000133">
        <property type="entry name" value="nephronectin isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000184">
        <property type="entry name" value="nephronectin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000268">
        <property type="entry name" value="nephronectin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.200">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001881">
        <property type="entry name" value="EGF-like_Ca-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000152">
        <property type="entry name" value="EGF-type_Asp/Asn_hydroxyl_site"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018097">
        <property type="entry name" value="EGF_Ca-bd_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009030">
        <property type="entry name" value="Growth_fac_rcpt_cys_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000998">
        <property type="entry name" value="MAM_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052235">
        <property type="entry name" value="Nephronectin_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049883">
        <property type="entry name" value="NOTCH1_EGF-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24050:SF19">
        <property type="entry name" value="NEPHRONECTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24050">
        <property type="entry name" value="PA14 DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07645">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00629">
        <property type="entry name" value="MAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00179">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00137">
        <property type="entry name" value="MAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57184">
        <property type="entry name" value="Growth factor receptor domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00010">
        <property type="entry name" value="ASX_HYDROXYL"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01186">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01187">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50060">
        <property type="entry name" value="MAM_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0106">Calcium</keyword>
    <keyword evidence="11" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="3" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="9" id="KW-0221">Differentiation</keyword>
    <keyword evidence="12 16" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6 16" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="5" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="7 18" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="18">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019282131" description="Nephronectin" evidence="18">
        <location>
            <begin position="21"/>
            <end position="611"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="19">
        <location>
            <begin position="74"/>
            <end position="105"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="19">
        <location>
            <begin position="107"/>
            <end position="146"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="19">
        <location>
            <begin position="187"/>
            <end position="231"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="19">
        <location>
            <begin position="232"/>
            <end position="272"/>
        </location>
    </feature>
    <feature type="domain" description="MAM" evidence="20">
        <location>
            <begin position="462"/>
            <end position="605"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="329"/>
            <end position="407"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="17">
        <location>
            <begin position="332"/>
            <end position="341"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="17">
        <location>
            <begin position="362"/>
            <end position="375"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="385"/>
            <end position="407"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="95"/>
            <end position="104"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009738"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00053701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00064366"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00069631"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50060"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001213.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="611" mass="67768" checksum="C52F04CD7817603F" modified="2019-05-08" version="1">MDLFFLRLLAVAWLHLEAAAEFDGRWPRQIVSSMGLCRFGSRVDCCWGWARQSWGQCQPFYVLRQRIARIRCQLKAICQPRCKHGECIGPNKCKCLPGYTGKTCNQDLNECGLKPRPCKHRCMNTYGSYKCYCLNGYMLMPDGSCSNALSCSMANCQYGCDVVKGEVRCHCPSPGLQLGPDGRTCIDVDECATGRVVCPRFRKCVNTFGSYICKCHKGFELMYIGGKYQCHDIDECSIGQYQCGSFSRCYNTQGTYKCKCKEGYRGDGINCLPIPKVMIEPSRPNNIFKGNTTLPKGSSGMTNRIPDIGGTRQPPRPPHVPAIITVRPAPKPTVQPTTKPTIWRTPKPTTRFVPKPMTTRPPLKPATRPPTPPQPTTSKPAPTARSPQLTTKDSSLVPTETTSPQGFTVDNRIQIEPQKPRGDVFIPRQPGVSNNLFEIFEIERGISADEDEANDDPGILVHSCNFDSGLCGWIKDKDDNLHWEPLRDPSGGQYMTISEPKGKEGKVARLVLPLGQLAHSGDLCLSFRHKVSGLHSGLLQVFLRKNGAHGPAVWGRNGGHGWRQTQITLQDSGIKSVIFKGEKGKGRTGDIGLDDVSLRKGRCSEEHEQHH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHW3</accession>
    <name>A0A452GHW3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">ELK domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Microtubule organizing center</location>
            <location evidence="1">Centrosome</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000001314.1">
        <property type="protein sequence ID" value="ENSGAGP00000001169.1"/>
        <property type="gene ID" value="ENSGAGG00000000844.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060090">
        <property type="term" value="F:molecular adaptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028745">
        <property type="entry name" value="AKAP9/Pericentrin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005539">
        <property type="entry name" value="ELK_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019528">
        <property type="entry name" value="PACT_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR44981:SF3">
        <property type="entry name" value="PERICENTRIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR44981">
        <property type="entry name" value="PERICENTRIN-LIKE PROTEIN, ISOFORM F"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10495">
        <property type="entry name" value="PACT_coil_coil"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01188">
        <property type="entry name" value="ELK"/>
        <property type="match status" value="9"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4 6" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="5" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="ELK" evidence="8">
        <location>
            <begin position="259"/>
            <end position="280"/>
        </location>
    </feature>
    <feature type="domain" description="ELK" evidence="8">
        <location>
            <begin position="415"/>
            <end position="436"/>
        </location>
    </feature>
    <feature type="domain" description="ELK" evidence="8">
        <location>
            <begin position="473"/>
            <end position="494"/>
        </location>
    </feature>
    <feature type="domain" description="ELK" evidence="8">
        <location>
            <begin position="541"/>
            <end position="564"/>
        </location>
    </feature>
    <feature type="domain" description="ELK" evidence="8">
        <location>
            <begin position="678"/>
            <end position="699"/>
        </location>
    </feature>
    <feature type="domain" description="ELK" evidence="8">
        <location>
            <begin position="707"/>
            <end position="725"/>
        </location>
    </feature>
    <feature type="domain" description="ELK" evidence="8">
        <location>
            <begin position="848"/>
            <end position="867"/>
        </location>
    </feature>
    <feature type="domain" description="ELK" evidence="8">
        <location>
            <begin position="1001"/>
            <end position="1023"/>
        </location>
    </feature>
    <feature type="domain" description="ELK" evidence="8">
        <location>
            <begin position="2962"/>
            <end position="2980"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1"/>
            <end position="46"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="69"/>
            <end position="102"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="3259"/>
            <end position="3284"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="117"/>
            <end position="285"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="321"/>
            <end position="386"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="432"/>
            <end position="613"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="640"/>
            <end position="674"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="703"/>
            <end position="926"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="1139"/>
            <end position="1240"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="1283"/>
            <end position="1407"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="1613"/>
            <end position="1640"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="1697"/>
            <end position="1881"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="1924"/>
            <end position="1962"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="2009"/>
            <end position="2084"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="2111"/>
            <end position="2194"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="2258"/>
            <end position="2362"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="2439"/>
            <end position="2515"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="2891"/>
            <end position="3038"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="3085"/>
            <end position="3119"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="3444"/>
            <end position="3519"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="1"/>
            <end position="13"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="72"/>
            <end position="87"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="SMART" id="SM01188"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001169.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="3748" mass="437074" checksum="5B158BC37F56FA89" modified="2019-05-08" version="1">MDEERRRKLEAGRAKLAHFRQRKTKGDNTQSQKKAAKRKGSSDYMQNIPKEECVIVAQDSDVLTDLNISVESKTEETETRSENKADVDLSNSNPNLVDDRASEEYDQLDVQMCRTRIMELEKRLLDKQEATEKLTAEVDDLLEQLAQHSEATHLQELETAVQERNEIIAQLTSNLQQARQNRDDIQEEASLLADQIHGLQLQLHEIEKLNAEIKDKVEKQQNLEEERKEQIGLIKKVHEREHDREISELIIKHEDEMKELRAELIKKQQQLLDELQQQMEGRHKTEVQHAQLQSQTLHNLELEALRLSLTNMHTSQLELMQSNLRKEKETALVELREMLNDKRAQEVAILQSRHQFELEHIDEQHLKEKEEIALKHQQELDEKKKIELEMEKEHIQTLETLNGDWALKTETCLKNVREELSEKHQTEMGELQKTLKLELDRVKEELKSLSLEKEQVEMKYENLENEHQKAITKLREQLQTEHDQCLENLKRKSKEREQEMQQEMEKLQATYEELKTQSQEEIKQLWSQLDSTRASRQELSELKEQLLARTSHVDEIERLKQDFEQQQQQRRNEHETELEQLRIYFEEKLGAAEENYREELTLLHQRLQELKEYSLLESEISQDQPLDISSSVTLLEEASEKERRDTLDQLTQQLEQHKEELTCLRLQMEEKHRHELDSLRSSLALQYKEDLMKMKMDLSDRYISEIEELKKKHCLDLEQLRAKLSEEHIKEITKLRLQSAQDAARQVEAEVAERVCVLDNEHKAKLSLLQSEKQHIAELQGEIQHLEREITKLKDIGVQHETQLKEEMEKIKCKLVDNHKNELKRAREEVQQMEQLHKEKEEEWKCEREALRIKAEEKLSQLRTELEDKAEYEKQTLQKKFELREAEMNQLQDQQAARIVELEKSLKEQQNSLQQLEDSLASMQTTQKAQAQYDSELQAARTLMEKDLEKATFELQEECAAKLMEAQNKFMEEHKVMTQKFTTEQEVLLQELKGKHADELELQSQQLQEKHKQQILSLTVELETKHQAETETLKSTLQSKQQAQLEAHIAELQAKHQAQIDELEAKHLSNLDSLESSYLSEIQIIRDEHKQVVEELQMHFTEQLQEKDKANQAILTQELERLKLKHSEELQLSQDNLKIELATIHIEKLKAMAAELEEAHKEDLNTALQNQRCMLEEESHKALDVLREEVLQMEEHHKKALKELQDLHVTEVQKQKEEYSWQLQEELEKLKAQHEHEEKMYASASTSEVETVRAALLAQFEEVKLKQQLQFQDEIELLKCQSEVLLEQQITQLKEEFEAERKTAWRSKEQVLIQETEKAQAHYKKEKEELSVQIQEKTNIIIQLEKKVESLNHELEETNCELETLLQRRDRENQEGEYLVAMLRSDIELSQNERKKLQESYQQILKLFVEMVKATIATEDLICKKVGLCLDDSLASGDSRESRSMMEEVGFIQHFKAREKHDLEKSDLLDETLTEHNQISAVTDEGSELSQHLCESVFASPDMAFENEEMILKICRRLRTAVEKLLELVTESTKQLEKTHEIHAHFEEEFSRRTQETAQVVNQHHDLMECLNEESEAKNQLALELHKAEGIIEGYVVEKAALEEAMSLKEESERRLVLELESLQEQFQKLTQEQAILREERDMLISQKKALAANVGEIEVGVPVANVAKKEDSGLLKEVEFLAKEKLELQCQAEKDHSTLRSQMKALEVELEEQMDQNQKLAKKSLEVTDLRQQIQVLEKQLRSQRQFMDEQAIEREHERDEFQQEIQKLEDQLKLSGKSQTSGEPKQYGIFELTLQIESLQAEIKDKTDDYNKLLLEKEQKYREVTVQDKEIEELAARIRELELTNTEASKTVSRLEQELQKMKKRETELTQDKEALQQQQYNNLIQISALQSKLDEARHRVPSEGSSDHGLKEQLQAEQEALLLKEREILSLLEQLEQFKENLINKNEEILRLNLQLEMQKNLAAGIGQLQVENVHLKEDIAKLHMMQSQNVGNSESAVLQFPQALLEEKNQEIDHLNEQIERLQRELEGATDNKVVENGKSEIDELRLLVEHLRGDQERLHRDKAEEVEQLHGVIEKLQKELVQLEPMYHEVSDSQDNLNLLWLEKQAENLQNELKKGSLDFQEHSDKNKKAVLLHSKLKELQEELELVSTAREALQQLLEEKESQFKMEVEILEQNLQNVQESSRQHLAELTSLKVQHDAFQEEYSLLKTLLSQRDGEIGIMSTRIQELEDTLRAREALLLESDLQIKTVAEQRVVEAAELQHLTEKIARLEDELLKKDLHQKTEAEEVQTLQSEVSRLDFKVQTLNQKEVVYQREIDELQANAIKLKDQIQEFLKELQALRSERDDLYSQLESYKEKDQSNDQEAKLHKSVFKQEEVLVHTNGMERLGEERGANADQSFAASVSQSDKLILVQLEATNTSENHKDLITKMTLHQEELKSELACVKKELELSEEQTGKILEDIKEKDAAIADLKTHSRNLKTQIKQLQETLLTQEAEMTDKARELQDLKKQCQQILEFHQTPDSMKCKLNNLSQNTEHDKESPKDFKCLVMDMTSWDSPEIVRKQETSIELQPSLHLTPFSEVDSVHSTDFEMKHNKSSVVQGDTPGLLGYQGLYANTNEEAVVRLHLKSPAFSESTYSIPEYQNTDKRIPVRDVDNFTLTPSDSQDDLRSTMSGLEGASDGYVSNASSDLAAKLNQELETSEHLDASFLAYLQYCGIFQDAMDPVTEKETISPQLKSVLKMVYEDSRRILALSEHPFPLSDQKSIQQSAAAEGWRKEGLILLDAIQSLRDYLSKVEDRGDKESSDTSFDWRGELLQAVQSVLEKERNMFQIDLQSHLCNPGSGDEGSLVEKLEHIVKRQEEQQRTVLEHLLSSDRNSLLSEIQDLRAQLRMTHLQNQEKLQQLQETLTNAEDRGSKQEHQLRRKVELLEYKLQQEKTIANDLQNSLSEEQERASEMHELLKQERSVVSDLKSELSESKEENESLQTFLQKLQREILQLRSELENKEKDLTVALQDLQNEHSKQKELRNIFEEQHLQHKLREDEKTKALEELQAALELQCIQNNQMSVALEHEKSANDNLRKELQIEHSRCEALLSQERNKLTELQSNLEVEKNHSLELLNALNHERVLTEQLSMRGNEDSSCQHKESLVEQTFVQELQAQLDAERSRAMELAAIIEKTHQQAIQSKRQLEAEVQMCCEETQKEREISTKLRATLESLQSQKQEVIRSLEIQREREAKLKVDWEQLQTLFRTMKEQEKRKKEEREKERREEQRAEFEKRTEWQKDKERVHELELQHQRDEHRIKELQQTLAELEKQERNLASQKSQRELSSCPIKNDYNANLSLTTETEALHLQQQQLEKIRQQLLFAAVHLNEFIYKTVDKTVNDWSASNDEAIASLLHTLKELKSELLTSSTPLKPLQGSVSLVDLLLKENGELTKSVTTLTEEKLELRAVISQLEKNLQQHHRRGISNRQIRSTDDAHSVQESERASWQREKTILQNALKQAESELAKATVEIENKPVMETSNPKLQKLYKKYLRAESFRKALVYQKKYLLLLLGGFQDCEQATLSLIARMGVYPSPADLQVSESRSRPFTKFRSAVRVVIAVSRLKFLVKKWHKVNRKGAQGENISHSIGHNPVSGARTEVLRQQQLPSVNVNSPPTRDIGSCHRTSSARFVNHSPKSSYWLHNRLHPSTSSVSEKSLVSTQDPERSLTEYIHRLEVIQQRLGGAQIGSTPGRPHQRNIK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHW4</accession>
    <name>A0A452GHW4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Protein AMBP</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">In the N-terminal section; belongs to the calycin superfamily. Lipocalin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHW4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001315.1">
        <property type="protein sequence ID" value="ENSGAGP00000001170.1"/>
        <property type="gene ID" value="ENSGAGG00000000433.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004867">
        <property type="term" value="F:serine-type endopeptidase inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd22596">
        <property type="entry name" value="Kunitz_bikunin_1-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd22597">
        <property type="entry name" value="Kunitz_bikunin_2-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.410.10:FF:000010">
        <property type="entry name" value="Alpha1-microglobulin/bikunin (AMBP)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.128.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.410.10">
        <property type="entry name" value="Pancreatic trypsin inhibitor Kunitz domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002968">
        <property type="entry name" value="A1-microglobln"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029856">
        <property type="entry name" value="AMBP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012674">
        <property type="entry name" value="Calycin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002223">
        <property type="entry name" value="Kunitz_BPTI"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036880">
        <property type="entry name" value="Kunitz_BPTI_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000566">
        <property type="entry name" value="Lipocln_cytosolic_FA-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020901">
        <property type="entry name" value="Prtase_inh_Kunz-CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46676">
        <property type="entry name" value="PROTEIN AMBP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46676:SF1">
        <property type="entry name" value="PROTEIN AMBP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00014">
        <property type="entry name" value="Kunitz_BPTI"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00061">
        <property type="entry name" value="Lipocalin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01215">
        <property type="entry name" value="A1MCGLOBULIN"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00759">
        <property type="entry name" value="BASICPTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00131">
        <property type="entry name" value="KU"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57362">
        <property type="entry name" value="BPTI-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50814">
        <property type="entry name" value="Lipocalins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00280">
        <property type="entry name" value="BPTI_KUNITZ_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50279">
        <property type="entry name" value="BPTI_KUNITZ_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0157">Chromophore</keyword>
    <keyword evidence="5" id="KW-0165">Cleavage on pair of basic residues</keyword>
    <keyword evidence="11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6" id="KW-0646">Protease inhibitor</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="9" id="KW-0722">Serine protease inhibitor</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
    <feature type="domain" description="BPTI/Kunitz inhibitor" evidence="13">
        <location>
            <begin position="204"/>
            <end position="254"/>
        </location>
    </feature>
    <feature type="domain" description="BPTI/Kunitz inhibitor" evidence="13">
        <location>
            <begin position="260"/>
            <end position="310"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008238"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00018905"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022685"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022690"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50279"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001170.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="325" mass="35846" checksum="C66EA62CD6DE4C4C" modified="2019-05-08" version="1">MYLSRPVVPKYFTSGPPLTLSVPLPPVPGPGVYGKWYDIATGTTCKWMKQYKDKFNMGTLVLGPGMTSNQISTTSTRLRQGVCTQVSGEYQKTDIPGKYTYYNPSWDVTINSYVVHTNYEEYAIILMQKKSSFGLTITAKLYGTCLCLNLLRGLGVRLWPRGDGSEGRGHGHVAPGCHHHMPCISVALKRNSLGSTLLMCSDFCRLNKDVGPCMGMNVRFFYNSSSMACETFHYGGCLGNGNNFLSEKKCLQTCRTEAACRLPIVPGPCQQPVILWAFDATQGKCTSFNYGGCQGNGNKFYTEKECREYCGVPADGMEEFLGLSN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHW5</accession>
    <name>A0A452GHW5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Kazal-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHW5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001362.1">
        <property type="protein sequence ID" value="ENSGAGP00000001214.1"/>
        <property type="gene ID" value="ENSGAGG00000000956.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005518">
        <property type="term" value="F:collagen binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050840">
        <property type="term" value="F:extracellular matrix binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00104">
        <property type="entry name" value="KAZAL_FS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.238.10:FF:000053">
        <property type="entry name" value="Putative testican-3 isoform 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.60.30:FF:000003">
        <property type="entry name" value="SPARC/osteonectin, cwcv and kazal-like domains proteoglycan 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.60.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.238.10">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011992">
        <property type="entry name" value="EF-hand-dom_pair"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002350">
        <property type="entry name" value="Kazal_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036058">
        <property type="entry name" value="Kazal_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019577">
        <property type="entry name" value="SPARC/Testican_Ca-bd-dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13866">
        <property type="entry name" value="SPARC OSTEONECTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13866:SF17">
        <property type="entry name" value="TESTICAN-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07648">
        <property type="entry name" value="Kazal_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10591">
        <property type="entry name" value="SPARC_Ca_bdg"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00280">
        <property type="entry name" value="KAZAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47473">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF100895">
        <property type="entry name" value="Kazal-type serine protease inhibitors"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51465">
        <property type="entry name" value="KAZAL_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="7" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="8" id="KW-0357">Heparan sulfate</keyword>
    <keyword evidence="5" id="KW-0654">Proteoglycan</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <feature type="domain" description="Kazal-like" evidence="9">
        <location>
            <begin position="71"/>
            <end position="122"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022974"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023207"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51465"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001214.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="297" mass="33240" checksum="838FEB8502C06279" modified="2019-05-08" version="1">MEWKSVSSTIVSSLSNDDLYSLDPSKDPCLKVKCSPHKVCVTHDYETAICVSRKHLWQKKGNLAPKHWIGPASLVKCKPCPVMHASPVCGSDGHTYTSKCKLEFHACTSGKTTAARCEGPCPCLPGQESPKHKAEKTACTDKELRSLASRLKDWFGALHEDANRVIKPTSSDTAPGRFDTSILPICKDSLGWMFNKLDMNYDLLLDHSEINAIYLDKYEPCIKPLFNSCDSFKDGKLSNNEWCYCFQKPGGLPCQNEMNRIQKLSRGKSLLGFGSESHLWNQSIVLVSYWLGREMLN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHW7</accession>
    <name>A0A452GHW7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Nephronectin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="7">Functional ligand of integrin alpha-8/beta-1 in kidney development. Regulates the expression of GDNF with integrin alpha-8/beta-1 which is essential for kidney development. May also play a role in the development and function of various tissues, regulating cell adhesion, spreading and survival through the binding of several integrins.</text>
    </comment>
    <comment type="subunit">
        <text evidence="8">Homodimer and homotrimer.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the nephronectin family.</text>
    </comment>
    <comment type="caution">
        <text evidence="10">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHW7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001369.1">
        <property type="protein sequence ID" value="ENSGAGP00000001218.1"/>
        <property type="gene ID" value="ENSGAGG00000000964.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00054">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd06263">
        <property type="entry name" value="MAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000038">
        <property type="entry name" value="Fibrillin 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000187">
        <property type="entry name" value="nephronectin isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.200:FF:000133">
        <property type="entry name" value="nephronectin isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000184">
        <property type="entry name" value="nephronectin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000268">
        <property type="entry name" value="nephronectin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.200">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001881">
        <property type="entry name" value="EGF-like_Ca-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000152">
        <property type="entry name" value="EGF-type_Asp/Asn_hydroxyl_site"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018097">
        <property type="entry name" value="EGF_Ca-bd_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009030">
        <property type="entry name" value="Growth_fac_rcpt_cys_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000998">
        <property type="entry name" value="MAM_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052235">
        <property type="entry name" value="Nephronectin_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049883">
        <property type="entry name" value="NOTCH1_EGF-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24050:SF19">
        <property type="entry name" value="NEPHRONECTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24050">
        <property type="entry name" value="PA14 DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07645">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00629">
        <property type="entry name" value="MAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00179">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00137">
        <property type="entry name" value="MAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57184">
        <property type="entry name" value="Growth factor receptor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00010">
        <property type="entry name" value="ASX_HYDROXYL"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01186">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01187">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50060">
        <property type="entry name" value="MAM_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="6 10" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="2 10" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="3 12" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="12">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019476959" description="Nephronectin" evidence="12">
        <location>
            <begin position="21"/>
            <end position="571"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="13">
        <location>
            <begin position="74"/>
            <end position="105"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="13">
        <location>
            <begin position="147"/>
            <end position="191"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="13">
        <location>
            <begin position="192"/>
            <end position="232"/>
        </location>
    </feature>
    <feature type="domain" description="MAM" evidence="14">
        <location>
            <begin position="422"/>
            <end position="565"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="289"/>
            <end position="367"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="292"/>
            <end position="301"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="11">
        <location>
            <begin position="322"/>
            <end position="335"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="345"/>
            <end position="367"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="10">
        <location>
            <begin position="95"/>
            <end position="104"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00009738"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00053701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00064366"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00069631"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50060"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001218.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="571" mass="63234" checksum="632CB2EC698FD3F5" modified="2019-05-08" version="1">MDLFFLRLLAVAWLHLEAAAEFDGRWPRQIVSSMGLCRFGSRVDCCWGWARQSWGQCQPFYVLRQRIARIRCQLKAICQPRCKHGECIGPNKCKCLPGYTGKTCNQDALSCSMANCQYGCDVVKGEVRCHCPSPGLQLGPDGRTCIDVDECATGRVVCPRFRKCVNTFGSYICKCHKGFELMYIGGKYQCHDIDECSIGQYQCGSFSRCYNTQGTYKCKCKEGYRGDGINCLPIPKVMIEPSRPNNIFKGNTTLPKGSSGMTNRIPDIGGTRQPPRPPHVPAIITVRPAPKPTVQPTTKPTIWRTPKPTTRFVPKPMTTRPPLKPATRPPTPPQPTTSKPAPTARSPQLTTKDSSLVPTETTSPQGFTVDNRIQIEPQKPRGDVFIPRQPGVSNNLFEIFEIERGISADEDEANDDPGILVHSCNFDSGLCGWIKDKDDNLHWEPLRDPSGGQYMTISEPKGKEGKVARLVLPLGQLAHSGDLCLSFRHKVSGLHSGLLQVFLRKNGAHGPAVWGRNGGHGWRQTQITLQDSGIKSVIFKGEKGKGRTGDIGLDDVSLRKGRCSEEHEQHH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHX0</accession>
    <name>A0A452GHX0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Paired domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHX0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001326.1">
        <property type="protein sequence ID" value="ENSGAGP00000001180.1"/>
        <property type="gene ID" value="ENSGAGG00000000896.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00131">
        <property type="entry name" value="PAX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.10:FF:000013">
        <property type="entry name" value="Paired box 8 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.10:FF:000003">
        <property type="entry name" value="Paired box protein Pax-6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.10">
        <property type="entry name" value="Winged helix-like DNA-binding domain superfamily/Winged helix DNA-binding domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009057">
        <property type="entry name" value="Homeodomain-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043182">
        <property type="entry name" value="PAIRED_DNA-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001523">
        <property type="entry name" value="Paired_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022130">
        <property type="entry name" value="Pax2_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043565">
        <property type="entry name" value="PAX_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036388">
        <property type="entry name" value="WH-like_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45636:SF20">
        <property type="entry name" value="PAIRED BOX PROTEIN PAX-5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45636">
        <property type="entry name" value="PAIRED BOX PROTEIN PAX-6-RELATED-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00292">
        <property type="entry name" value="PAX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12403">
        <property type="entry name" value="Pax2_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00027">
        <property type="entry name" value="PAIREDBOX"/>
    </dbReference>
    <dbReference type="SMART" id="SM00351">
        <property type="entry name" value="PAX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46689">
        <property type="entry name" value="Homeodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00034">
        <property type="entry name" value="PAIRED_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51057">
        <property type="entry name" value="PAIRED_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="5" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="3" id="KW-0563">Paired box</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0804">Transcription</keyword>
    <keyword evidence="4" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="Paired" evidence="9">
        <location>
            <begin position="7"/>
            <end position="133"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="134"/>
            <end position="153"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="173"/>
            <end position="209"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="134"/>
            <end position="150"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022724"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51057"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001180.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="394" mass="42702" checksum="C95442C1D9517E0E" modified="2019-05-08" version="1">QDLNKNRHGGVNQLGGVFVNGRPLPDVVRQRIVELAHQGVRPCDISRQLRVSHGCVSKILGRYYETGSIKPGVIGGSKPKVATPKVVEKIAEYKRQNPTMFAWEIRDRLLAERVCDNDTVPSVSSINRIIRTKVQQPPNQQVPASSHSIASTGSVTQVSTVSTDSAGSSYSISGILGITSPSAESNKRKRDEGIQESPVPNGHSLPGRDFLRKQMRGDLFTQQQLEVLDRVFERQHYSDIFTTTEPIKPEQTTEYSAMASLAGGLDDMKANLTSPTSAELGASVPGPQSYPIVTGRELASTTLPGYPPHVPPAGQGSYSAPTLTGMVPGSEFSGSPYSHPQYSTYNDSWRFPNPGLLGEFSYPSFYRNSLLTAAGACPVGGISPHWMLGQLWPL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHX3</accession>
    <name>A0A452GHX3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Kazal-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHX3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001370.1">
        <property type="protein sequence ID" value="ENSGAGP00000001220.1"/>
        <property type="gene ID" value="ENSGAGG00000000956.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005518">
        <property type="term" value="F:collagen binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050840">
        <property type="term" value="F:extracellular matrix binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00104">
        <property type="entry name" value="KAZAL_FS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.60.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.238.10">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011992">
        <property type="entry name" value="EF-hand-dom_pair"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002350">
        <property type="entry name" value="Kazal_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036058">
        <property type="entry name" value="Kazal_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019577">
        <property type="entry name" value="SPARC/Testican_Ca-bd-dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13866">
        <property type="entry name" value="SPARC OSTEONECTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13866:SF17">
        <property type="entry name" value="TESTICAN-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07648">
        <property type="entry name" value="Kazal_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10591">
        <property type="entry name" value="SPARC_Ca_bdg"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00280">
        <property type="entry name" value="KAZAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47473">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF100895">
        <property type="entry name" value="Kazal-type serine protease inhibitors"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51465">
        <property type="entry name" value="KAZAL_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="3" id="KW-0732">Signal</keyword>
    <feature type="domain" description="Kazal-like" evidence="6">
        <location>
            <begin position="62"/>
            <end position="113"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS51465"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001220.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="289" mass="32282" checksum="75020C92AB5EADBC" modified="2025-10-08" version="2">MVYFRNDYFVPSLDPSKDPCLKVKCSPHKVCVTHDYETAICVSRKHLWQKKGNLAPKHWIGPASLVKCKPCPVMHASPVCGSDGHTYTSKCKLEFHACTSGKTTAARCEGPCPCLPGQESPKHKAEKTACTDKELRSLASRLKDWFGALHEDANRVIKPTSSDTAPGRFDTSILPICKDSLGWMFNKLDMNYDLLLDHSEINAIYLDKYEPCIKPLFNSCDSFKDGKLSNNEWCYCFQKPGGLPCQNEMNRIQKLSRGKSLLGKFDSALFFIPGGKGHSIAMADTVDYM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHX4</accession>
    <name>A0A452GHX4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Reticulon</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="6">Inhibits amyloid precursor protein processing, probably by blocking BACE1 activity.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 7">Endoplasmic reticulum membrane</location>
            <topology evidence="1 7">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHX4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001331.1">
        <property type="protein sequence ID" value="ENSGAGP00000001185.1"/>
        <property type="gene ID" value="ENSGAGG00000000953.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043005">
        <property type="term" value="C:neuron projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0014069">
        <property type="term" value="C:postsynaptic density"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007420">
        <property type="term" value="P:brain development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071787">
        <property type="term" value="P:endoplasmic reticulum tubular network formation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030182">
        <property type="term" value="P:neuron differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.2480:FF:000001">
        <property type="entry name" value="Reticulon"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.2480">
        <property type="match status" value="1"/>
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    </dbReference>
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    </dbReference>
    <dbReference type="PANTHER" id="PTHR45799:SF5">
        <property type="entry name" value="RETICULON-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45799">
        <property type="entry name" value="RETICULON-LIKE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02453">
        <property type="entry name" value="Reticulon"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50845">
        <property type="entry name" value="RETICULON"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3 7" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="5 7" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 7" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="333"/>
            <end position="362"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="448"/>
            <end position="469"/>
        </location>
    </feature>
    <feature type="domain" description="Reticulon" evidence="9">
        <location>
            <begin position="319"/>
            <end position="506"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1"/>
            <end position="68"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
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            <begin position="272"/>
            <end position="314"/>
        </location>
    </feature>
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        <location>
            <begin position="13"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="24"/>
            <end position="37"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004477"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00037306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU210713"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50845"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001185.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="506" mass="55226" checksum="771C760F76D380DC" modified="2019-05-08" version="1">MSAPPDRQGLPLSGSGARWAGAGAEERGAGAQRRDGDGAQQPQLPAGSAREHPPVTMATASTGDAEASDLFDYSSTKTSKGGDRSFYTSLIADDRYTSPPDSTYFSGILQKENSPVTCSDSTEGFNTTGHSIQDVMGFESRGLFSSDSGIEMTPAEPTDVNKTLADPMKVEAYKYMDISRSEEVKYQERCNTSLEDKNTSFIDKCPETAAKLEHISGSKKPAFEETKVAKGQNLFEESTVPSCAEEIFGDQHSASLITAPVKITLTEIENIGETANKESPPNKQETGLKPSHDVVPTVTVSEPEDDSPGSLTPPSSGTAIDLLYWRDIKQTGIVFGSILLLLFSLTQFSVVSVIAYLALAALSATISFRIYKSVLQAVQKTDEGHPFKAYLEMEMSLSQEQIQKYTDCLQLYVNSTVRELRRLFLVQDLVDSLKFAVLMWLLTYVGALFNGLTLMIMAVVSMFTLPVVYDKYQAQIDQYLGLVRTHINTVMAKIQAKIPGAKRKAE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHX6</accession>
    <name>A0A452GHX6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Olduvai domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Golgi apparatus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000001327.1">
        <property type="protein sequence ID" value="ENSGAGP00000001181.1"/>
        <property type="gene ID" value="ENSGAGG00000000835.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060090">
        <property type="term" value="F:molecular adaptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007098">
        <property type="term" value="P:centrosome cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090063">
        <property type="term" value="P:positive regulation of microtubule nucleation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903358">
        <property type="term" value="P:regulation of Golgi organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056273">
        <property type="entry name" value="CDK5RAP2_MYOME_CC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012943">
        <property type="entry name" value="Cnn_1N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052593">
        <property type="entry name" value="MT-associated_AKAP9-binding"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010630">
        <property type="entry name" value="Olduvai_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46501">
        <property type="entry name" value="MYOMEGALIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46501:SF2">
        <property type="entry name" value="MYOMEGALIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23246">
        <property type="entry name" value="CC_CDK5RAP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07989">
        <property type="entry name" value="Cnn_1N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01148">
        <property type="entry name" value="DUF1220"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51316">
        <property type="entry name" value="ODV"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="5" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Olduvai" evidence="9">
        <location>
            <begin position="1483"/>
            <end position="1574"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="924"/>
            <end position="971"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1243"/>
            <end position="1264"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1324"/>
            <end position="1350"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1518"/>
            <end position="1639"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1927"/>
            <end position="1968"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="2153"/>
            <end position="2201"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="41"/>
            <end position="68"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="94"/>
            <end position="150"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="236"/>
            <end position="291"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="350"/>
            <end position="516"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="545"/>
            <end position="630"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="732"/>
            <end position="874"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="992"/>
            <end position="1019"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="1185"/>
            <end position="1212"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="1293"/>
            <end position="1320"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="1672"/>
            <end position="1720"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="1746"/>
            <end position="1826"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="2094"/>
            <end position="2128"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="954"/>
            <end position="963"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="1245"/>
            <end position="1256"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="1526"/>
            <end position="1543"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="1567"/>
            <end position="1578"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="1582"/>
            <end position="1602"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="1609"/>
            <end position="1624"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="1927"/>
            <end position="1936"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="1947"/>
            <end position="1965"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="2178"/>
            <end position="2190"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004555"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51316"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001181.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2220" mass="249523" checksum="478CCC108803AE2A" modified="2019-05-08" version="1">MSNGYRTLSQHLNDLKKENFSLKLRIYFLEERMQQKYEASREDVYKRNIELKVEVESLKQELQERQQTLDKAWAAAENLTSCNEAEFQRRYEERQQEKEHVSELLENKIQLLQEEARLAKSEAERTAALVEAEKGRCLELTKKLKELTKEEEKRGAQVLRDDHCAALAQKDKHIEELIQELSAREQLIELLATEKQNLLQRLEEPREIEVQNLPEDHQQLTQCDPVASGRVSESHAAELQDKIQQYDATNKLLQEKLNEMNFELKSIQEASQRQDRTIQSLNEALKSKESETEELYHVIEGQNETMGKLRDMLHRSQLGHLQTSEGPSPSQQQQHMALLDLQNTLFCTQLEMQKLKRAQWQKEHQLAEARRTTQLLEATVQEEQQLKEVSWKHNQELRGVVQQVQAELQGKVQELWALEGEKCCEIRAQEQRVQRLSQRLAHKEQLLQESWELLQRRQSLEKNPVVTDTMLEKLQQRIKHRDAALERAIDEKFCAVEEKEQELQQLHISVREREHDLERLRSVLSSNEATIHSMESLLKAKTLELEQVTATCQNLQWLKEEVEAKSNCWQKEQEGIIQQLQASLHDRNKEVEELTATLLCKLGPGQSEIAEELCLRLQRKERMLQDLLSDRNRQAGEHETELRELLQAMSIREQRSRVTAEKMVQALAEKSDEVQFLRQRLVGNEPGKNPVADGRLLLQKDKQPFQELLPGGCGNATATRISKGEDSKYRMERGLLETAAELEKELINAREELELMARKERESRLELSSLQSVVTAQEEELQVQASDIESLTRNIQSKEDLIKDLQMQLVDPEEIPAVERLTQEVLVLREKVATAESRGQEATGNRQQQLLLMLEGLVAEKSRLNEALQAERQLYSSLVKFHAHPDSPERGQTLQAELEGAQALRGRLEESLGRSMEHLRRLETLGAFGGQPGRDDAKDGSTDSIEGEAPPRTVNHQTSQGLPANSPEAKAAQPDLLTLAPLERDSSLQAELLHVRAKNQQLLEQKAKVEGELWELKALMEEAGFSSLSSIRKVLLSLCLETAELKEQLGGSPLSEGWEDKAEARAALCTQNSKLQSSETVVTLLKEQLVLNCPEGESKFPPQLVTGMATEIDRLWTEMGVASGKRPALEGQLQEDPAKRPCPRSQAVELGPAQAPRGMSQQMEACPAGSWQRLVEPRVGPSGQLVQCRQRNQELQDKLAVSEATVRAQAEQLEQYRTLLSEPLVQQDNKQVQVDLQDLGYETCGRSENEADREETPSPECEEPDVFSKAGLMEELSRPGPWGGLCWGQSEDVAVLQQHVRELQLQLQNSNKALQRLQRRTRSFSSTSDYASGAERLHKPVPSATDEDEGWQSDGVGAFCPPALQASRDLEWLVHRVSLLEAQLHGRWAGGVLPEELRSAVWPGKYDSLIQAQARELSHLRQKMREGRSVCHLLTQHLRDTVKSFEELLRGTDIDYYMGQSFREHLAQGGQLAERLSSKLSSRDRSDVEDKTGHEFVALRLSKELREKERMIETLQAKLQERCETPSSSRALSESPRSNSSASFLSDGPEACSDGDATSEYSQLQGDRSEQPSRRLTDVTHPAALASNPSPAAAPHQASAESIRGYPTGPSSQHPPMDSGQTRTGLHFDSLSKPLSVPLTPAPGPSAFLPFGPPPPTPSALLGCCGTPVFSLAEVQQELHMLQKQLGESADLLEGHLAEIRSLRQRLEESICINDRLREQLEKRLATTTKGNGYPTNIYIQGLESASSLSSETQALREENQSLQLQFAHLSRELEQLRETTRCRLQKAETENRTLAQELAEQRQSGRQLREERLALQENNHRLECTVPLLQQQCQENQQLCQALCAELRIHETLSSSSQAALSACSGDVHRRPPSSSELGALLAEVRTLRRKLLRGIQVNSALRQQLARQQPEGSAAPLFAAHQATPNWQPFQDSSPSPPVRDVGMSSPAPLSPPALLSGPAPLAHQLKEPPVDDALVRRSAPAPAGDAPSSSSASKGGCQVIGHVDDYYTLKQLILEGKTLVHQMASLLRPALAMPSLEPRGTKALDRGSVQQLLSTASALHQLLEQCSSLLTTFWRGALPAAPSPAQPQTAEQAMKDELLALRARLAEQENLLQRAAERLQDTCRLKDNMEHFIVSHLTQTHAVLRKARTNLEKSKPQGSSVKSPPFIGAGEISVGLSQHPSHVIPTNQEHRRKKSCPDTLKWQEDQSWPCFVHLPSS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GI22</accession>
    <name>A0A452GI22_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Mediator of RNA polymerase II transcription subunit 17</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="7">Mediator complex subunit 17</fullName>
        </alternativeName>
    </protein>
    <gene>
        <name evidence="7" type="primary">MED17</name>
    </gene>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="6 7">Component of the Mediator complex, a coactivator involved in the regulated transcription of nearly all RNA polymerase II-dependent genes. Mediator functions as a bridge to convey information from gene-specific regulatory proteins to the basal RNA polymerase II transcription machinery. Mediator is recruited to promoters by direct interactions with regulatory proteins and serves as a scaffold for the assembly of a functional preinitiation complex with RNA polymerase II and the general transcription factors.</text>
    </comment>
    <comment type="subunit">
        <text evidence="7">Component of the Mediator complex.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 7">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 7">Belongs to the Mediator complex subunit 17 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GI22"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001449.1">
        <property type="protein sequence ID" value="ENSGAGP00000001291.1"/>
        <property type="gene ID" value="ENSGAGG00000001036.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070847">
        <property type="term" value="C:core mediator complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016592">
        <property type="term" value="C:mediator complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003712">
        <property type="term" value="F:transcription coregulator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019313">
        <property type="entry name" value="Mediator_Med17"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13114">
        <property type="entry name" value="MEDIATOR OF RNA POLYMERASE II TRANSCRIPTION SUBUNIT 17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13114:SF7">
        <property type="entry name" value="MEDIATOR OF RNA POLYMERASE II TRANSCRIPTION SUBUNIT 17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10156">
        <property type="entry name" value="Med17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0010">Activator</keyword>
    <keyword evidence="5 7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 7" id="KW-0804">Transcription</keyword>
    <keyword evidence="3 7" id="KW-0805">Transcription regulation</keyword>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="51"/>
            <end position="77"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005635"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00025687"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU364140"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001291.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="599" mass="67042" checksum="313DEA6D0F8790B7" modified="2019-05-08" version="1">MSGAPAVRISIESASEKQVQEVGLDGSETYLQPLSMSQNLARLAQRIDFSQGSGSEEDEPPPGAQARDWAAPGDAEDEEGLVKFQPSLWPWDSVRNNLRGALTEMCVLYDVLSIVKDKKFMTLDPVVQDPLPPKQNPQTLQLISKKKSLAGAAQILLKGAERLSKSVAENQENKRQRDFNSELLRLRQHWKLRKVGDKILGDLSYRSAGSLFPHHGTFEVIKNTDIDLDKKIPEDYCPLDVQIPSDLEGSAYIKVSIQKQAPDIGDLGTVNLFKRPLPKSKPGAPHWQTKLEAAQNVLLCKEIFAQLSREAVQIKSQIPHIVVKNQIISQPFPGLQLSISLCHSSNDKKSQKAASEKQSPEDHLYVLEHNLHQLIREFHKQTLSSTMMPHPASAPFGHKRMRMAGPQAFDKNDISSLQPNEGLLEKIIKQAKHIFLRRRSIQLQLNIGVEQIRVVHRDGRVITLSHQEQELQDFLLSQMSQHQVHAVQQLAKVMGWHVLSFSNHVGLGPVESIGNASAITVASPSGDYAISVRNGPESGSKVMVQFPRCQCKDLPKSDVLQDSKWNHLRGPFREVQWNKMEGRNFVYKLELLMAALTPC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GI27</accession>
    <name>A0A452GI27_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Mediator of RNA polymerase II transcription subunit 17</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="7">Mediator complex subunit 17</fullName>
        </alternativeName>
    </protein>
    <gene>
        <name evidence="7" type="primary">MED17</name>
    </gene>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="6 7">Component of the Mediator complex, a coactivator involved in the regulated transcription of nearly all RNA polymerase II-dependent genes. Mediator functions as a bridge to convey information from gene-specific regulatory proteins to the basal RNA polymerase II transcription machinery. Mediator is recruited to promoters by direct interactions with regulatory proteins and serves as a scaffold for the assembly of a functional preinitiation complex with RNA polymerase II and the general transcription factors.</text>
    </comment>
    <comment type="subunit">
        <text evidence="7">Component of the Mediator complex.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 7">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 7">Belongs to the Mediator complex subunit 17 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GI27"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001455.1">
        <property type="protein sequence ID" value="ENSGAGP00000001297.1"/>
        <property type="gene ID" value="ENSGAGG00000001036.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070847">
        <property type="term" value="C:core mediator complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016592">
        <property type="term" value="C:mediator complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003712">
        <property type="term" value="F:transcription coregulator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019313">
        <property type="entry name" value="Mediator_Med17"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13114">
        <property type="entry name" value="MEDIATOR OF RNA POLYMERASE II TRANSCRIPTION SUBUNIT 17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13114:SF7">
        <property type="entry name" value="MEDIATOR OF RNA POLYMERASE II TRANSCRIPTION SUBUNIT 17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10156">
        <property type="entry name" value="Med17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0010">Activator</keyword>
    <keyword evidence="5 7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 7" id="KW-0804">Transcription</keyword>
    <keyword evidence="3 7" id="KW-0805">Transcription regulation</keyword>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="51"/>
            <end position="77"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005635"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00025687"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU364140"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001297.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="588" mass="65707" checksum="39B50FAAFD2CC761" modified="2019-05-08" version="1">MSGAPAVRISIESASEKQVQEVGLDGSETYLQPLSMSQNLARLAQRIDFSQGSGSEEDEPPPGAQARDWAAPGDAEDEEGLVKFQPSLWPWDSVRNNLRGALTEMCVLYDVLSIVKDKKFMTLDPVVQDPLPPKQNPQTLQLISKKKSLAGAAQILLKGAERLSKSVAENQENKRQRDFNSELLRLRQHWKLRKVGDKILGDLSYRSAGSLFPHHGTFEVIKNTDIDLDKKIPEDYCPLDVQIPSDLEGSAYIKVSIQKQAPDIGDLGTVNLFKRPLPKSKPGAPHWQTKLEAAQNVLLCKEIFAQLSREAVQIKSQIPHIVVKNQIISQPFPGLQLSISLCHSSNDKKSQKAASEKQSPEDHLYVLEHNLHQLIREFHKQTLSSTMMPHPASAPFGHKRMRMAGPQAFDKNDISSLQPNEGLLEKIIKQAKHIFLRRRTARTIDSLASRIEDPQIQAHWSNINDVYESSVKVLITSQGYEQICKSIQLQLNIGVEQIRVVHRDGRVITLSHQEQELQDFLLSQMSQHQVHAVQQLAKVMGWHVLSFSNHVGLGPVESIGNASAITVASPSGDYAISGRILYDYNRNV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GI31</accession>
    <name>A0A452GI31_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">Exocyst complex component</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">Component of the exocyst complex involved in the docking of exocytic vesicles with fusion sites on the plasma membrane.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the SEC15 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GI31"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001460.1">
        <property type="protein sequence ID" value="ENSGAGP00000001302.1"/>
        <property type="gene ID" value="ENSGAGG00000001025.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000145">
        <property type="term" value="C:exocyst"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006893">
        <property type="term" value="P:Golgi to plasma membrane transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006886">
        <property type="term" value="P:intracellular protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090522">
        <property type="term" value="P:vesicle tethering involved in exocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.357.30:FF:000001">
        <property type="entry name" value="Exocyst complex component"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.357.30">
        <property type="entry name" value="Exocyst complex subunit Sec15 C-terminal domain, N-terminal subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007225">
        <property type="entry name" value="EXOC6/Sec15"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046361">
        <property type="entry name" value="EXOC6/Sec15_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042045">
        <property type="entry name" value="EXOC6/Sec15_C_dom1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048359">
        <property type="entry name" value="EXOC6_Sec15_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12702:SF2">
        <property type="entry name" value="EXOCYST COMPLEX COMPONENT 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12702">
        <property type="entry name" value="SEC15"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20651">
        <property type="entry name" value="EXOC6_Sec15_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04091">
        <property type="entry name" value="Sec15_C"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF025007">
        <property type="entry name" value="Sec15"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="1" id="KW-0268">Exocytosis</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0813">Transport</keyword>
    <feature type="domain" description="Exocyst complex component EXOC6/Sec15 N-terminal" evidence="3">
        <location>
            <begin position="57"/>
            <end position="226"/>
        </location>
    </feature>
    <feature type="domain" description="Exocyst complex subunit EXOC6/Sec15 C-terminal" evidence="2">
        <location>
            <begin position="420"/>
            <end position="602"/>
        </location>
    </feature>
    <feature type="domain" description="Exocyst complex subunit EXOC6/Sec15 C-terminal" evidence="2">
        <location>
            <begin position="609"/>
            <end position="718"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="PIRNR" id="PIRNR025007"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF04091"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="Pfam" id="PF20651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001302.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="761" mass="88523" checksum="43EFE35F805624B5" modified="2019-05-08" version="1">MARKSKLNLLELYSEENKDQIYENVFAEIESFELPLGTSLRSVYDDQPNAHQRFMEKLDACIRNHDREIEKMCNFHHQGFVDAITELLKVRADAGKLKVQVTDTNRRLQDAGKEVIAQTEEIIRCRIQQRNITTVVEKLQLCLPVLEMYSKLKEQMNVKRYYSALKTMEQLENTYLPRVSQYRFCQIMIENLPKLREEIKEISMSDLKDFLESIRKHSDKIGETAMKQAQQQKTFNTVLQKQNNVNYGKNMYVINGRVPEPKYEITSKRTLEEEDEHEEEVLTAHDLVDFSPVYRCLHIYSVLGDGETFENYYRKQRRKQARLVLQPQSSMHETVEGYRRYFSQIVGFFVVEDHILHVTQGLVTRAYTDELWNMALSKIIAVLRTHSSYCTDPDLVLELKNLIVVFADTLQGYGFPVNRLFDLLFEIRDQYNETLLKKWTGLFRDIFEADNYSPIPVANEEEYKIVISKFPFQDPELDKQSFPKKLPMSQSVPQIYIQVKEFIYASLKFSESLHRSSTEIDDMLRKSTNLLLTRTLSSCLQNLIKKPHIGLTELVQIIINTTHLEQACKYLEDFITNITNISQETVHTARLYGLSTFKNNNNDHAAMSGKVAQTACMSACQHLSTSLMQMLLDSELKQISMGAVQQFNLDVIQCELFASSEPVPGFHGDTLQLAFIDLRQLLDLFMVWDWSTYLADYGQPTSKYLRVNPSTALTLLEKMKETSKKNNIFAQFRKNDRDKQKLIETVVKQLRSLVNGMSQHT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GI58</accession>
    <name>A0A452GI58_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">DNA ligase</fullName>
            <ecNumber evidence="15">6.5.1.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">DNA ligase that seals nicks in double-stranded during DNA repair. Also involved in DNA replication and DNA recombination.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13 15">
            <text>ATP + (deoxyribonucleotide)n-3'-hydroxyl + 5'-phospho-(deoxyribonucleotide)m = (deoxyribonucleotide)n+m + AMP + diphosphate.</text>
            <dbReference type="EC" id="6.5.1.1"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 16">Belongs to the ATP-dependent DNA ligase family.</text>
    </comment>
    <dbReference type="EC" id="6.5.1.1" evidence="15"/>
    <dbReference type="AlphaFoldDB" id="A0A452GI58"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001492.1">
        <property type="protein sequence ID" value="ENSGAGP00000001332.1"/>
        <property type="gene ID" value="ENSGAGG00000000751.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003910">
        <property type="term" value="F:DNA ligase (ATP) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071897">
        <property type="term" value="P:DNA biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006310">
        <property type="term" value="P:DNA recombination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006281">
        <property type="term" value="P:DNA repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903461">
        <property type="term" value="P:Okazaki fragment processing involved in mitotic DNA replication"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07900">
        <property type="entry name" value="Adenylation_DNA_ligase_I_Euk"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd07969">
        <property type="entry name" value="OBF_DNA_ligase_I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.3260.10:FF:000001">
        <property type="entry name" value="DNA ligase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.50.140:FF:000062">
        <property type="entry name" value="DNA ligase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.470.30:FF:000016">
        <property type="entry name" value="DNA ligase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1490.70">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.3260.10">
        <property type="entry name" value="DNA ligase, ATP-dependent, N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.470.30">
        <property type="entry name" value="DNA ligase/mRNA capping enzyme"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.50.140">
        <property type="entry name" value="Nucleic acid-binding proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050191">
        <property type="entry name" value="ATP-dep_DNA_ligase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000977">
        <property type="entry name" value="DNA_ligase_ATP-dep"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012309">
        <property type="entry name" value="DNA_ligase_ATP-dep_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012310">
        <property type="entry name" value="DNA_ligase_ATP-dep_cent"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016059">
        <property type="entry name" value="DNA_ligase_ATP-dep_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012308">
        <property type="entry name" value="DNA_ligase_ATP-dep_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036599">
        <property type="entry name" value="DNA_ligase_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012340">
        <property type="entry name" value="NA-bd_OB-fold"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00574">
        <property type="entry name" value="dnl1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45674:SF4">
        <property type="entry name" value="DNA LIGASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45674">
        <property type="entry name" value="DNA LIGASE 1/3 FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04679">
        <property type="entry name" value="DNA_ligase_A_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01068">
        <property type="entry name" value="DNA_ligase_A_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04675">
        <property type="entry name" value="DNA_ligase_A_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF117018">
        <property type="entry name" value="ATP-dependent DNA ligase DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56091">
        <property type="entry name" value="DNA ligase/mRNA capping enzyme, catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50249">
        <property type="entry name" value="Nucleic acid-binding proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00697">
        <property type="entry name" value="DNA_LIGASE_A1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00333">
        <property type="entry name" value="DNA_LIGASE_A2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50160">
        <property type="entry name" value="DNA_LIGASE_A3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 15" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="12" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="4" id="KW-0132">Cell division</keyword>
    <keyword evidence="7 15" id="KW-0227">DNA damage</keyword>
    <keyword evidence="9 15" id="KW-0233">DNA recombination</keyword>
    <keyword evidence="10 15" id="KW-0234">DNA repair</keyword>
    <keyword evidence="5" id="KW-0235">DNA replication</keyword>
    <keyword evidence="3 15" id="KW-0436">Ligase</keyword>
    <keyword evidence="6 15" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="ATP-dependent DNA ligase family profile" evidence="18">
        <location>
            <begin position="573"/>
            <end position="709"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="20"/>
            <end position="60"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="17">
        <location>
            <begin position="44"/>
            <end position="58"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007572"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022598"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022705"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023172"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00034003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00058910"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="RuleBase" id="RU000617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="RuleBase" id="RU004196"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50160"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001332.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="844" mass="94099" checksum="67DFC89433549624" modified="2019-05-08" version="1">EAAAMQRNGLSLCLDSPVKSAKRPLRRVLESDSDEEEAVLTARAEMEPTQDKSLERASSKQVSSAHPLSLPWNSHGASLGSCIPPRIPANIRPGLHPGLCAHRVSCDSAASPMHALLQLGACLIHAIIVPVATCAGLPSLGVLVQANCWETLRNTSAVGPVYWTWWLPISLWLCDPSSFSKPVVPLDPMEYNPSKSSYHPVQDACWGHGQRVPYLAVARTFERIEEVSARLKNIETLSNLFRSVVALSPSDILPCIYLCLNRLGPAYKGLELGIGETILMKAVAQAMGRQLDQIKAEAQEKGDLGLVAESSRSNQRTMFTPPKLSAAGVFSKLQEIARMTGSASMNKKIDIIKGLFVACRHSEARYITRSLGGKLRIGLAEQSVLSAIAHAVSLTPPGQGFPPEVLDAGQSKSAEARKAWLEEQTQILKQTFCELPSYDIIVPILLEHGVESLPQHCKITPGIPLKPMLAQPTKGIGEVLKRFEEAAFTCEYKYDGERAQIHILENGDVHIYSRNQEDNTSKYPDIISRIPKVKKDSVQSCILDAEAVAWDPEKRQIQPFQVLTTRKRKDVEAAEIKVQVCLYAFDILYLNGESLVKEPFSRRRQLLHESFVETEGEFLFATSMDTCSLDEISEFLDQSIKDSCEGLMVKTLEVDATYEIAKRSHKWLKLKKDYLEGVGDTLDLVVIGAYLGKGKRVGMYGGFLLACYDEESEEYQSICKIGTGFTEESLEKHYSFLKDHVLEKPRPYYRWDSSTEPNHWLAAVQVWEVKCADLSISPVHKAAIGLVDEEKGISLRFPRFLRVRDDKKPEEATTSSQVAELYRRQQQIQNQQLVEKAEEEEDFY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GI60</accession>
    <name>A0A452GI60_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Eukaryotic translation initiation factor 4 gamma 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="11">Appears to play a role in the switch from cap-dependent to IRES-mediated translation during mitosis, apoptosis and viral infection. Cleaved by some caspases and viral proteases.</text>
    </comment>
    <comment type="subunit">
        <text evidence="13">Interacts with the serine/threonine protein kinases MKNK1 and MKNK2. Binds EIF4A and EIF3. Interacts with MIF4GD. Interacts with DAZAP2.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the eukaryotic initiation factor 4G family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GI60"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001485.1">
        <property type="protein sequence ID" value="ENSGAGP00000001325.1"/>
        <property type="gene ID" value="ENSGAGG00000000987.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016281">
        <property type="term" value="C:eukaryotic translation initiation factor 4F complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003729">
        <property type="term" value="F:mRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003743">
        <property type="term" value="F:translation initiation factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006446">
        <property type="term" value="P:regulation of translational initiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd11559">
        <property type="entry name" value="W2_eIF4G1_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.180:FF:000007">
        <property type="entry name" value="Eukaryotic translation initiation factor 4 gamma 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.180:FF:000011">
        <property type="entry name" value="Eukaryotic translation initiation factor 4 gamma 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.180:FF:000017">
        <property type="entry name" value="Eukaryotic translation initiation factor 4 gamma 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.180">
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003891">
        <property type="entry name" value="Initiation_fac_eIF4g_MI"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003890">
        <property type="entry name" value="MIF4G-like_typ-3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003307">
        <property type="entry name" value="W2_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23253">
        <property type="entry name" value="EUKARYOTIC TRANSLATION INITIATION FACTOR 4 GAMMA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23253:SF9">
        <property type="entry name" value="EUKARYOTIC TRANSLATION INITIATION FACTOR 4 GAMMA 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02847">
        <property type="entry name" value="MA3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02854">
        <property type="entry name" value="MIF4G"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02020">
        <property type="entry name" value="W2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00515">
        <property type="entry name" value="eIF5C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00544">
        <property type="entry name" value="MA3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00543">
        <property type="entry name" value="MIF4G"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51366">
        <property type="entry name" value="MI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51363">
        <property type="entry name" value="W2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0007">Acetylation</keyword>
    <keyword evidence="5" id="KW-0396">Initiation factor</keyword>
    <keyword evidence="4" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="2" id="KW-0488">Methylation</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="9" id="KW-0648">Protein biosynthesis</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0678">Repressor</keyword>
    <keyword evidence="8" id="KW-0810">Translation regulation</keyword>
    <keyword evidence="7" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="MI" evidence="16">
        <location>
            <begin position="533"/>
            <end position="656"/>
        </location>
    </feature>
    <feature type="domain" description="W2" evidence="15">
        <location>
            <begin position="710"/>
            <end position="894"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="39"/>
            <end position="61"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="51"/>
            <end position="61"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00005775"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022491"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022540"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022845"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022917"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00037759"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00040449"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00046720"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS51363"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS51366"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001325.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="897" mass="101550" checksum="D82BF89C9B99A99E" modified="2019-05-08" version="1">EPVSCGGAGAAASTFSFLSLLPLFISFNKYSEFLGKTPGQNAQKWIPSRSTRRDDDSANDKERHDAIFRKVRGILNKLTPEKFDKLCLELLNVGVESKLILKGIILLIVDKALEEPKYSSLYAQLCLRLAEDAPNFDGPSAECHPGQKQSTTFRRLLISKLQDEFENRTRNVDIYDKRDGPLLPEEEEQRAIAKIKMLGNIKFIGELGKLDLIHESILHKCIKTLLEKKKRVQLKDMGEDLECLCQIMRTVGPRLDHAKAKSLMDQYFARMRSLMLSKELPARIRFLLQDTVELREHNWVPRKAFLDNGPKTINQIRQDAVKDLGVFIPAPMAQGMRSDFFLEGPFMPPRMKLDRDPLGGLADMFGQMPGSGIGTGPGVIQDRFSPTMGRHRSNQLFNGHGGHLMPPAQSQFGDLGKSFLKNQGQSQLYHNQNQGLLSQQQGQSKDMPPRFSKKGQLNADEISLRPAQSFLMNKNQVPKLQPQITMIPPSAQPPRTQTPPLGQPPQLGLKTNPPLIQEKPAKISKKPPPSKEELLKLTEAVVTDYLNNGNVNDAISGVREMRAPKPFIPEMLSKVIIQSLDRSDEDKEKASALISLLKQEGIATSDNFMQAFLNVLDQCPKLEVDIPLVKSYLAQFAARAIISELVSISELAQPLESGTHFPLFLLCLQQLAKLQDREWLTELFQQSKVNMQKMLPEIDQNKDRMLEILEGKGLSFLFPLLKLEKELLKQIKLDPSPQAIYKWIKDNISPKLHVDKGFVNILMTSFLQYISSEVSPPNDEPDSSSAPSKEQLEQEKQLLLSFKPVMQKFLHDHVDLQVSALYALQVHCYNNNFPKGMLLHFFVHFYDMEIIEEEAFLAWKEDISQEFPGKGKALFQVNQWLTWLETAEEEESEEEAD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GI62</accession>
    <name>A0A452GI62_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Homeobox domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 7 8">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the paired homeobox family. Bicoid subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GI62"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001497.1">
        <property type="protein sequence ID" value="ENSGAGP00000001337.1"/>
        <property type="gene ID" value="ENSGAGG00000001053.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00086">
        <property type="entry name" value="homeodomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.60:FF:000142">
        <property type="entry name" value="homeobox protein OTX2 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.60">
        <property type="entry name" value="Homeodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001356">
        <property type="entry name" value="HD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017970">
        <property type="entry name" value="Homeobox_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009057">
        <property type="entry name" value="Homeodomain-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003022">
        <property type="entry name" value="Otx2_TF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003025">
        <property type="entry name" value="Otx_TF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013851">
        <property type="entry name" value="Otx_TF_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45793">
        <property type="entry name" value="HOMEOBOX PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45793:SF2">
        <property type="entry name" value="HOMEOBOX PROTEIN OTX2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00046">
        <property type="entry name" value="Homeodomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03529">
        <property type="entry name" value="TF_Otx"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01257">
        <property type="entry name" value="OTX2HOMEOBOX"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01255">
        <property type="entry name" value="OTXHOMEOBOX"/>
    </dbReference>
    <dbReference type="SMART" id="SM00389">
        <property type="entry name" value="HOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46689">
        <property type="entry name" value="Homeodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00027">
        <property type="entry name" value="HOMEOBOX_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50071">
        <property type="entry name" value="HOMEOBOX_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="4 7 8" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="5 7 8" id="KW-0371">Homeobox</keyword>
    <keyword evidence="6 7 8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Homeobox" evidence="10">
        <location>
            <begin position="36"/>
            <end position="96"/>
        </location>
    </feature>
    <feature type="DNA-binding region" description="Homeobox" evidence="7">
        <location>
            <begin position="38"/>
            <end position="97"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="93"/>
            <end position="143"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="132"/>
            <end position="143"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006503"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023155"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00108"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="RuleBase" id="RU000682"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50071"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001337.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="289" mass="31608" checksum="05E42DE03461CE81" modified="2019-05-08" version="1">MMSYLKQPPYAVNGLSLTTSGMDLLHPSVGYPATPRKQRRERTTFTRAQLDVLEALFAKTRYPDIFMREEVALKINLPESRVQVWFKNRRAKCRQQQQQQQNGGQSKVRPAKKKSSPAREVSSESGTNGQFTPPSSTSVPAISSSSAPVSIWSPASISPLSDPLSTSSSCMQRSYPMTYTQASGYSQGYAGSTSYFGGMDCGSYLTPMHHQLPGPGATLSPMGTNAVTSHLNQSPASLSTQGYGASSLGFNSTTDCLDYKDQTASWKLNFNADCLDYKDQTSSWKFQVL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GI78</accession>
    <name>A0A452GI78_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="8">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GI78"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001509.1">
        <property type="protein sequence ID" value="ENSGAGP00000001346.1"/>
        <property type="gene ID" value="ENSGAGG00000001047.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00160">
        <property type="entry name" value="RhoGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12141">
        <property type="entry name" value="SH3_DNMBP_C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.40:FF:000066">
        <property type="entry name" value="dynamin-binding protein isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.40:FF:000193">
        <property type="entry name" value="Rho guanine nucleotide exchange factor 38"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.60">
        <property type="entry name" value="Arfaptin homology (AH) domain/BAR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.900.10">
        <property type="entry name" value="Dbl homology (DH) domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027267">
        <property type="entry name" value="AH/BAR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035899">
        <property type="entry name" value="DBL_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000219">
        <property type="entry name" value="DH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051492">
        <property type="entry name" value="Dynamin-Rho_GEF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22834">
        <property type="entry name" value="NUCLEAR FUSION PROTEIN FUS2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22834:SF17">
        <property type="entry name" value="RHO GUANINE NUCLEOTIDE EXCHANGE FACTOR 38"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00621">
        <property type="entry name" value="RhoGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07653">
        <property type="entry name" value="SH3_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14604">
        <property type="entry name" value="SH3_9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00325">
        <property type="entry name" value="RhoGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00326">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103657">
        <property type="entry name" value="BAR/IMD domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48065">
        <property type="entry name" value="DBL homology domain (DH-domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50010">
        <property type="entry name" value="DH_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0344">Guanine-nucleotide releasing factor</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1 3" id="KW-0728">SH3 domain</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="5">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019343314" evidence="5">
        <location>
            <begin position="24"/>
            <end position="729"/>
        </location>
    </feature>
    <feature type="domain" description="DH" evidence="7">
        <location>
            <begin position="38"/>
            <end position="226"/>
        </location>
    </feature>
    <feature type="domain" description="SH3" evidence="6">
        <location>
            <begin position="497"/>
            <end position="560"/>
        </location>
    </feature>
    <feature type="domain" description="SH3" evidence="6">
        <location>
            <begin position="665"/>
            <end position="728"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="588"/>
            <end position="623"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022658"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50010"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001346.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="729" mass="84085" checksum="24048CB7B558159D" modified="2019-05-08" version="1">MSGHIHISPALWILSGSLSPVTSLQENEEQLTRRMMAKRVKIIAELVQTEKDYLHDLELCIREVVQPLRHKQVRKFDVDGLFSNIESVHQISAKLLSLLEEATADMDPLFLEIKGPLEDTYKIYCYHHDDAHTMLESYEKNEELKQHLKHWVAILHDSLFVCINNRGKPNLMDMGSLMIKPIQRVMKYPLLLCELLNSTPASHPDHKALQDAFAAVKDINVNINELKRRKDLVLKYKKNDEEESLKEKFSRLNIHSISKKSKRVTSHLKILTGGEPQVKDQTFNKEEKLFRSLEKTVKFCVKNISFSLQHLQVGISHTHTYMSYALIKRICEQFNTHFSQRKYYLSAGDELDLAKKDYEALNAQLVEELQVFNRTAKTIVMNCLHCFITLLRDMMSSAFQLELLFLTSLSYVMYMRYNLTVYLQLSSSSIYEVQNQVLEEFHNLNFVKENSSATFIERKLSFEKKKPAPTQVSIMPLSPESPRQTESQRSKLLSTYSTESLYQAKRKCNATQEFDIDLYEGELVAVVEQKDPFGSTSRWRVDTGIHKGYVYSSFLKPYNPAKVQKDIAETSFCDDDFDNISLFVSSRPTNDSSARNLGHKRSDSSSSNNSLCEKPAINGTETDHDVDDQHVSIIHAAYLSISVFPIPNIKVVHYCLQFKGETLVYKRTTFYAVYAFQARSHQELSLQEYQRVRILQFSDLSGNKEWWLAEAKGQKGYVPANYLGKMTYA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GI81</accession>
    <name>A0A452GI81_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Type I cytokine receptor cytokine-binding domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="1 2">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="2" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="1" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="1" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GI81"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001451.1">
        <property type="protein sequence ID" value="ENSGAGP00000001293.1"/>
        <property type="gene ID" value="ENSGAGG00000001038.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000313" key="1">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001293.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="3">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="200" mass="23054" checksum="09B8A1C2E4422F44" modified="2019-05-08" version="1">MQSYTLRNTECSHIYRMRDSILENSDSEKDIGLMLKTSCQCNTVPQVCLHSNREVQFLVQVHIHTLALLVLMLKNSSKNGTSVENFSCVIYNISFMNCTWEAVFIGSKNFQDVKDIKNTVYFLVNGSSSKSEIQFYDDDYFILEKLTPPLNITVSCSEDRAECIVQWKQPQLSHMERNGDGCFEYQIDIRNKKTNVSVTF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GI83</accession>
    <name>A0A452GI83_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Peptidase S1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GI83"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001518.1">
        <property type="protein sequence ID" value="ENSGAGP00000001350.1"/>
        <property type="gene ID" value="ENSGAGG00000000827.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00190">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.10:FF:000005">
        <property type="entry name" value="Serine protease 37"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.10.10">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009003">
        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043504">
        <property type="entry name" value="Peptidase_S1_PA_chymotrypsin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001314">
        <property type="entry name" value="Peptidase_S1A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001254">
        <property type="entry name" value="Trypsin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018114">
        <property type="entry name" value="TRYPSIN_HIS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271:SF81">
        <property type="entry name" value="GRANZYME B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271">
        <property type="entry name" value="KALLIKREIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00089">
        <property type="entry name" value="Trypsin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00722">
        <property type="entry name" value="CHYMOTRYPSIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00020">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50494">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50240">
        <property type="entry name" value="TRYPSIN_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00134">
        <property type="entry name" value="TRYPSIN_HIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0732">Signal</keyword>
    <keyword evidence="2" id="KW-0865">Zymogen</keyword>
    <feature type="domain" description="Peptidase S1" evidence="4">
        <location>
            <begin position="14"/>
            <end position="238"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023145"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001350.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="240" mass="26661" checksum="9BB2F8CC377684D4" modified="2019-05-08" version="1">MQSLSCLVSSEGDIIGGREARPHSRPYMAFVETKMGPESTTECGGFLVREDFVLTAAHCDEEGSGDAPTVVILGAHDLNRKERTQQEIQVRRKIPHEDFDRTTLENDIMLLQLELRAEWNNAVKPIRLPPANFLLKPEATCSVAGWGRTGVHMVRPATKLQEVTLKMADDKQCELFFDSGSYYPSSPSRLFTFGDTGGPLVCNGFAQSIVSYGKDSGLPSAVYTRMSAFIPWLRKKMGRE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GI86</accession>
    <name>A0A452GI86_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Protein fantom</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="16">Nephrocystin-8</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="17">RPGR-interacting protein 1-like protein</fullName>
        </alternativeName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Negatively regulates signaling through the G-protein coupled thromboxane A2 receptor (TBXA2R). May be involved in mechanisms like programmed cell death, craniofacial development, patterning of the limbs, and formation of the left-right axis. Involved in the organization of apical junctions; the function is proposed to implicate a NPHP1-4-8 module. Does not seem to be strictly required for ciliogenesis. Involved in establishment of planar cell polarity such as in cochlear sensory epithelium and is proposed to implicate stabilization of disheveled proteins. Involved in regulation of proteasomal activity at the primary cilium probably implicating association with PSDM2.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="4">Cell junction</location>
            <location evidence="4">Tight junction</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Cytoskeleton</location>
            <location evidence="3">Cilium axoneme</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Cilium basal body</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
            <location evidence="2">Microtubule organizing center</location>
            <location evidence="2">Centrosome</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the RPGRIP1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GI86"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001456.1">
        <property type="protein sequence ID" value="ENSGAGP00000001298.1"/>
        <property type="gene ID" value="ENSGAGG00000001041.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005930">
        <property type="term" value="C:axoneme"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005923">
        <property type="term" value="C:bicellular tight junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036064">
        <property type="term" value="C:ciliary basal body"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032391">
        <property type="term" value="C:photoreceptor connecting cilium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031870">
        <property type="term" value="F:thromboxane A2 receptor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1905515">
        <property type="term" value="P:non-motile cilium assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009966">
        <property type="term" value="P:regulation of signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046548">
        <property type="term" value="P:retinal rod cell development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00030">
        <property type="entry name" value="C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000196">
        <property type="entry name" value="Protein fantom"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000073">
        <property type="entry name" value="protein fantom isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000075">
        <property type="entry name" value="protein fantom isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021656">
        <property type="entry name" value="C2-C2_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000008">
        <property type="entry name" value="C2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041091">
        <property type="entry name" value="RPGRIP1_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031139">
        <property type="entry name" value="RPGRIP1_fam"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14240:SF4">
        <property type="entry name" value="PROTEIN FANTOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14240">
        <property type="entry name" value="RETINITIS PIGMENTOSA GTPASE REGULATOR-INTERACTING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00168">
        <property type="entry name" value="C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11618">
        <property type="entry name" value="C2-C2_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18111">
        <property type="entry name" value="RPGR1_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00239">
        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49562">
        <property type="entry name" value="C2 domain (Calcium/lipid-binding domain, CaLB)"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50004">
        <property type="entry name" value="C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0965">Cell junction</keyword>
    <keyword evidence="13" id="KW-0966">Cell projection</keyword>
    <keyword evidence="11" id="KW-0969">Cilium</keyword>
    <keyword evidence="10 18" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="12" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0796">Tight junction</keyword>
    <feature type="domain" description="C2" evidence="20">
        <location>
            <begin position="768"/>
            <end position="894"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="974"/>
            <end position="1004"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1027"/>
            <end position="1077"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="18">
        <location>
            <begin position="105"/>
            <end position="139"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="18">
        <location>
            <begin position="200"/>
            <end position="255"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="18">
        <location>
            <begin position="326"/>
            <end position="457"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="18">
        <location>
            <begin position="494"/>
            <end position="549"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="974"/>
            <end position="986"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="1027"/>
            <end position="1044"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="1045"/>
            <end position="1061"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="19">
        <location>
            <begin position="1062"/>
            <end position="1074"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004120"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004430"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004435"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00006042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022427"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023069"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00058876"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00071893"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00078503"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00082462"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001298.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1274" mass="146576" checksum="CA42118E6B58C07A" modified="2019-05-08" version="1">MSAPADETAADLPVRDVGLNLAGIGGLQESSAAQSVKARQAVSRISREELEDRFLRLHDENILLKQHAHKQEDKIKRMATKLIRLVNDKKRSEQIGGGPKRLGQDVELEEMIEQLQGKVRELEKQNEGIRKRLISTKQQLQVQGHRHTPYSHVQSRINTGLRKVNEAVAIQDHTRKGMRFQDVELRCPQPMLPRYGHSLLEEARAEIRNLENVIESQKGQIEELEHTGEVLRSQIRKKEKEFEESVLQLKEQQATGQRLSIRDNVEMIKLHKQLMEKSNALSAMEGKFLQLQESQRNLKASHDALMANGNELNLQLKEQRLKCLHLEKELQSMTFSEQRIEELQDRINDLEKERDLLKENYDKLYNSAFSVSHEQQWKLKEQQLKLQIAQLETALKSDLADKSEILDRMKLERDQNEKLTQENKELQLHYLEHKQQLDELKNRMKFFTKESDTEMAELSEALMLIKVRKQQKNGDLTFLEKIEDDIHKDTEYAMRDLQATHAETVQELEKTRNMLIVQHKINKDYQTEVEAVTQKMEGLQKDYELKLEQYVHLLDIRAARIRKLEAQLKDIAYGTKQYKFRPEILPEDPADEFDETVHLERGENLFEIHISKVIFSTEAVHAFGDQEPVTFCTYAFYDFELQTTPVVHGLNPSYNFTSQYLVRVDDFFLQYIQKSTITLEVHHALGTDYETAAACQLRFQEILENNGRIYSTAILVGTGGHIPNYGTVDYWVRLRVPMDQAIRLYKERSKALGYITSNLKEHEKAPQLKQPIHRTAQVSTSADGDLNELHITIKCCSNLKSRKKHLQPNPYVVYKFFDFADHDTPIIHSSNSPQFDDHMCFPVPMNTDLDQYLKSESLTFYVFDDGETEEGLYVGKAIVPLISLAHDRYISGTFELTDSEGHSSGTINVDLKWRFAYLPPSGSAVAADLVSYIQNEKPMATKLPVKEEVPTLTLTPSFTSSMTKFPEKELMEIQTSGEHERKKDTQRPNFEAPKPKPRQRTALADKKVSFMDVSILQIPVIQERVADDKVKETSGEIQQEKDDLSQLSEGQLADQSSATSGDETEITEELEPEDQDNRLATDAMESIITDSDDCIVPSPVSKNIKQSIEKIRIEIISLSLTDSRVAMDDTIQRLFVECRFYSFLAEETPVSLPKPKSGQWVHYNYSNVIHVDKVNNCARREFLKSVLQESELNTGSIRFTVVSDPPEDEQDLECEDIGFAYVSLTEIFQKEKDIIEQDIDVFDSQDDSAVIGKLKVTVEALHALRSVYEEHKDD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GI95</accession>
    <name>A0A452GI95_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">inorganic diphosphatase</fullName>
            <ecNumber evidence="4">3.6.1.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subunit">
        <text evidence="3">Homodimer.</text>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the PPase family.</text>
    </comment>
    <dbReference type="EC" id="3.6.1.1" evidence="4"/>
    <dbReference type="AlphaFoldDB" id="A0A452GI95"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001545.1">
        <property type="protein sequence ID" value="ENSGAGP00000001370.1"/>
        <property type="gene ID" value="ENSGAGG00000001100.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004427">
        <property type="term" value="F:inorganic diphosphate phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000287">
        <property type="term" value="F:magnesium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006796">
        <property type="term" value="P:phosphate-containing compound metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00412">
        <property type="entry name" value="pyrophosphatase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.80.10:FF:000005">
        <property type="entry name" value="Pyrophosphatase (inorganic) 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.80.10">
        <property type="entry name" value="Inorganic pyrophosphatase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008162">
        <property type="entry name" value="Pyrophosphatase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036649">
        <property type="entry name" value="Pyrophosphatase_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10286">
        <property type="entry name" value="INORGANIC PYROPHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00719">
        <property type="entry name" value="Pyrophosphatase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50324">
        <property type="entry name" value="Inorganic pyrophosphatase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00387">
        <property type="entry name" value="PPASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0007">Acetylation</keyword>
    <keyword evidence="6" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="7" id="KW-0460">Magnesium</keyword>
    <keyword evidence="5" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="9">
        <location>
            <begin position="1"/>
            <end position="24"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019106811" description="inorganic diphosphatase" evidence="9">
        <location>
            <begin position="25"/>
            <end position="345"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006220"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00011738"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00012146"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001370.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="345" mass="39498" checksum="0C44DC3D4BD703A5" modified="2019-05-08" version="1">MRAELGQALRGLLLLSARSCSCSGAGRRRWLVPPLPRCSWLSTMAQCSTEQRGRPHSPEYRLYFKNAEGKYISPFHDIPLLAGSKEDKEVPVKRSKTNGSEVVFNMVVEVPRWSNAKMEIATKEPLNPIKQDIKKGKLRYVANIFPHKGYIWNYGALPQTWEDPHHKDKDTGCCGDNDPIDVCEIGTKVHSPGEIIQVKVLGVMALIDEGETDWKIIAISVDDPEAQKIHDIDDVKKHKPGYLEATAEWFRLYKVPDDKPENQFAFNGEFKDKDFAVEIIKSTHEYWKALLHKKADPGAIKWHHLLRVEILFLQKLMCGTFLTSDHRNVLKSHHQILNHPFFPEF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GI96</accession>
    <name>A0A452GI96_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="11">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1 7">Belongs to the peptidase M16 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GI96"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001543.1">
        <property type="protein sequence ID" value="ENSGAGP00000001368.1"/>
        <property type="gene ID" value="ENSGAGG00000000237.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005782">
        <property type="term" value="C:peroxisomal matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004222">
        <property type="term" value="F:metalloendopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050435">
        <property type="term" value="P:amyloid-beta metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042447">
        <property type="term" value="P:hormone catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043171">
        <property type="term" value="P:peptide catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051603">
        <property type="term" value="P:proteolysis involved in protein catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.830.10:FF:000004">
        <property type="entry name" value="Putative insulin-degrading enzyme"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.830.10:FF:000006">
        <property type="entry name" value="Putative insulin-degrading enzyme"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.830.10">
        <property type="entry name" value="Metalloenzyme, LuxS/M16 peptidase-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011249">
        <property type="entry name" value="Metalloenz_LuxS/M16"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011765">
        <property type="entry name" value="Pept_M16_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001431">
        <property type="entry name" value="Pept_M16_Zn_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050626">
        <property type="entry name" value="Peptidase_M16"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007863">
        <property type="entry name" value="Peptidase_M16_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032632">
        <property type="entry name" value="Peptidase_M16_M"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43690:SF18">
        <property type="entry name" value="INSULIN-DEGRADING ENZYME-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43690">
        <property type="entry name" value="NARDILYSIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00675">
        <property type="entry name" value="Peptidase_M16"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05193">
        <property type="entry name" value="Peptidase_M16_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16187">
        <property type="entry name" value="Peptidase_M16_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF63411">
        <property type="entry name" value="LuxS/MPP-like metallohydrolase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00143">
        <property type="entry name" value="INSULINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="6" id="KW-0482">Metalloprotease</keyword>
    <keyword evidence="2" id="KW-0645">Protease</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="Peptidase M16 N-terminal" evidence="8">
        <location>
            <begin position="68"/>
            <end position="202"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase M16 C-terminal" evidence="9">
        <location>
            <begin position="229"/>
            <end position="407"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase M16 middle/third" evidence="10">
        <location>
            <begin position="413"/>
            <end position="543"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007261"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023049"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU004447"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF00675"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF05193"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="Pfam" id="PF16187"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001368.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="564" mass="64984" checksum="27EBCE907B8EDC9F" modified="2019-05-08" version="1">PGEQERLHLFLSLPEFEFLLFPSRLQKNTHNKMSNPAIKRLANKIIKSPEDKREYRGLELENGIKAMLICDPTTDKSSAALDVHIGSLSDPLNIAGLSHFCEHMLFLGTKKYPKENEYSQFLSEHAGSSNAFTSGEHTNYYFDVSHEHLEGALDRFAQFFLCPLFDESCKEREVNAVDSEHEKNLMNDAWRLFQLEKATGNPNHPFSKFGTGNKFTLETRPTKEGIDVRQELLKFHSTYYSSNLMAICVLGRESLDELTGLVVKLFSEVENKNVPLPEFPEHPFQEQHLRQLYKVVPIKDIRNLYVTFPIPDLQKYYKSNPGHYLGHLIGHEGPGSLLSELKAKGWVNTLVGGQKEGARGFMFFIINVDLTEEGLLHVEDIILHMFQYIQKLRADGPQEWVFQECKDLNAVAFRFKDKERPRGYTSKLGGILHYYPIEEVLAAEYLLEEFRPDLIEMVLDKLRPENVRVAIVSKSFEGKTDRTEEWYGTQYKQEAISDEVIQKWQNADLNGKFKLPMKNEFIPTNFEILPLEKDAPQYPALIKVRICITYIKWKLCQPASLLEG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYS0</accession>
    <name>A0A452GYS0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Protein spinster homolog 1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="12">Spns1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>a 1-O-(1Z-alkenyl)-sn-glycero-3-phosphocholine(out) + H(+)(out) = a 1-O-(1Z-alkenyl)-sn-glycero-3-phosphocholine(in) + H(+)(in)</text>
            <dbReference type="Rhea" id="RHEA:74447"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:77287"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="14">
            <text>a 1-O-(1Z-alkenyl)-sn-glycero-3-phosphoethanolamine(out) + H(+)(out) = a 1-O-(1Z-alkenyl)-sn-glycero-3-phosphoethanolamine(in) + H(+)(in)</text>
            <dbReference type="Rhea" id="RHEA:74455"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:77288"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="9">
            <text>a 1-acyl-sn-glycero-3-phosphocholine(out) + H(+)(out) = a 1-acyl-sn-glycero-3-phosphocholine(in) + H(+)(in)</text>
            <dbReference type="Rhea" id="RHEA:74435"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:58168"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="10">
            <text>a 1-acyl-sn-glycero-3-phosphoethanolamine(out) + H(+)(out) = a 1-acyl-sn-glycero-3-phosphoethanolamine(in) + H(+)(in)</text>
            <dbReference type="Rhea" id="RHEA:74439"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:64381"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Lysosome membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="8">Belongs to the major facilitator superfamily. Spinster (TC 2.A.1.49) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYS0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008411.1">
        <property type="protein sequence ID" value="ENSGAGP00000007273.1"/>
        <property type="gene ID" value="ENSGAGG00000005846.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005765">
        <property type="term" value="C:lysosomal membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0022857">
        <property type="term" value="F:transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006869">
        <property type="term" value="P:lipid transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1250.20">
        <property type="entry name" value="MFS general substrate transporter like domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011701">
        <property type="entry name" value="MFS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020846">
        <property type="entry name" value="MFS_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044770">
        <property type="entry name" value="MFS_spinster-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036259">
        <property type="entry name" value="MFS_trans_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23505:SF13">
        <property type="entry name" value="PROTEIN SPINSTER HOMOLOG 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23505">
        <property type="entry name" value="SPINSTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07690">
        <property type="entry name" value="MFS_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103473">
        <property type="entry name" value="MFS general substrate transporter"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50850">
        <property type="entry name" value="MFS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0445">Lipid transport</keyword>
    <keyword evidence="7" id="KW-0458">Lysosome</keyword>
    <keyword evidence="6 16" id="KW-0472">Membrane</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 16" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 16" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="2" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="53"/>
            <end position="73"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="93"/>
            <end position="112"/>
        </location>
    </feature>
    <feature type="domain" description="Major facilitator superfamily (MFS) profile" evidence="17">
        <location>
            <begin position="55"/>
            <end position="199"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="1"/>
            <end position="37"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="155"/>
            <end position="199"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="15">
        <location>
            <begin position="166"/>
            <end position="175"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004155"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023228"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00024338"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00035932"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00036238"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00039482"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00041462"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00047765"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00048915"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50850"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007273.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="199" mass="21120" checksum="D02D2BE921CB5A71" modified="2019-05-08" version="1">MATRHGPHPDAAPFLSQADETEADAAEGGSRADGGQPEVGVHSVTGISYSRSVMIVAVLCYINLLNYMDRFTVAGILPDIESFFRIGDSSSGLLQTVFMSSYMVLAPIFGYLGDRYHRKRLLCLGIAFWSAVTLGSSFIPQEVPAATSAAACWPSSTSPSPSAGEGRWGAQGRPGRGWESQFSPPLPSPQQAGGECKEC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYS9</accession>
    <name>A0A452GYS9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Ig-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYS9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008424.1">
        <property type="protein sequence ID" value="ENSGAGP00000007283.1"/>
        <property type="gene ID" value="ENSGAGG00000005772.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="CDD" id="cd05763">
        <property type="entry name" value="IgI_LRIG1-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000161">
        <property type="entry name" value="Leucine rich repeats and immunoglobulin like domains 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000040">
        <property type="entry name" value="Leucine rich repeats and immunoglobulin like domains 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000150">
        <property type="entry name" value="Leucine rich repeats and immunoglobulin like domains 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000224">
        <property type="entry name" value="Leucine rich repeats and immunoglobulin like domains 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000023">
        <property type="entry name" value="Leucine rich repeats and immunoglobulin like domains 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000483">
        <property type="entry name" value="Cys-rich_flank_reg_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013098">
        <property type="entry name" value="Ig_I-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001611">
        <property type="entry name" value="Leu-rich_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003591">
        <property type="entry name" value="Leu-rich_rpt_typical-subtyp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050467">
        <property type="entry name" value="LRFN"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45842:SF16">
        <property type="entry name" value="LEUCINE-RICH REPEATS AND IMMUNOGLOBULIN-LIKE DOMAINS 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45842">
        <property type="entry name" value="SYNAPTIC ADHESION-LIKE MOLECULE SALM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07679">
        <property type="entry name" value="I-set"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13927">
        <property type="entry name" value="Ig_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13855">
        <property type="entry name" value="LRR_8"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01463">
        <property type="entry name" value="LRRCT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00019">
        <property type="entry name" value="LEURICHRPT"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00365">
        <property type="entry name" value="LRR_SD22"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SMART" id="SM00369">
        <property type="entry name" value="LRR_TYP"/>
        <property type="match status" value="12"/>
    </dbReference>
    <dbReference type="SMART" id="SM00082">
        <property type="entry name" value="LRRCT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52058">
        <property type="entry name" value="L domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51450">
        <property type="entry name" value="LRR"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="9" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="3" id="KW-0433">Leucine-rich repeat</keyword>
    <keyword evidence="8 12" id="KW-0472">Membrane</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5 13" id="KW-0732">Signal</keyword>
    <keyword evidence="4 12" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 12" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="13">
        <location>
            <begin position="1"/>
            <end position="28"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019360708" description="Ig-like domain-containing protein" evidence="13">
        <location>
            <begin position="29"/>
            <end position="938"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="790"/>
            <end position="816"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="14">
        <location>
            <begin position="484"/>
            <end position="583"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="14">
        <location>
            <begin position="588"/>
            <end position="677"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="14">
        <location>
            <begin position="682"/>
            <end position="768"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022614"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007283.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="938" mass="103447" checksum="469354A51805443D" modified="2019-05-08" version="1">MSCPLRAAPAAALGLLLLLLGRLAGSSAAGGPGSPSSAAGGPGSPSCPAPCRCFGDLLDCSRLKLISGGRMHSFFMLQQTFFFDLNRKLNNNELETIPNLGPVSANITLLSLTSNKIGGILPEHLKPFQSLETLDLSNNNISELKTASFPFLQLKYLYINSNRITSMEPGVFDSLSSTLQVLKLNRNKISVIPQKMFKLSHLQHLELNRNKIKKIDGLTFQGLTALKSLKMQRNGVNRLMDGAFWGLTNMEVLQLDHNNLTEITKGWLYGLLMLQQLHLSQNAISRISPDAWEFCQKLSELDLTFNHLARLDDSSFVGLSLLVGLYIGNNKVNYIADCAFRGLSSLKTLDLKNNEISWTIEDMNGAFSGLDKLRILILQGNRIRSITKKAFSGLDALEHLDLSNNAIMSVQGNAFSQMKKLEELHLNTSSLLCDCQLKWLPQWISENNFQSFVNASCAHPPLLKGRSIFAVSPDEFVCDDFPKPQITVQPETQSAIKGSNLSFVCSAASSSDSPMTFAWKKDNELLHDAEMENYAHLRARGGEVMEYTTILRLRNVEFSNEGKYQCVISNHFGSSYSVKAKLTVNMLPSFTKMPMDLSIRAGATARLECAAVGHPVPQIAWQKDGGTDFPAARKRRMHVMPEDDVFFIVDVKIEDTGVYSCTAQNTAGSISANATLTVLETPSFLRPLLDRTVTKGETAVLQCIAGGSPPPRLNWTKDDSPLVVTERHFFAAGNQLLIIVDTDIEDAGKYTCEMSNTLGTERGNIRLNVIPTPTCDSPQNIAPSLDDDGWATVGIVIIAVVCCVVGTSLVWVVIIYHTRRRNEDCSITNTDETNLPADIPSYLSSQGTLAERQDGYGSSENGSHHQFITSSMGGYFLQQRDSNGKYFKYKTWHCFWLLEIRHCGRCGDLFLISNCRVTFVAPLSSNSAQSCLFNPKPD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYV1</accession>
    <name>A0A452GYV1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Dihydropyrimidinase-related protein 1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="12">Collapsin response mediator protein 1</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="13">Inactive dihydropyrimidinase</fullName>
        </alternativeName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Necessary for signaling by class 3 semaphorins and subsequent remodeling of the cytoskeleton. Plays a role in axon guidance. During the axon guidance process, acts downstream of SEMA3A to promote FLNA dissociation from F-actin which results in the rearrangement of the actin cytoskeleton and the collapse of the growth cone. Involved in invasive growth and cell migration. May participate in cytokinesis.</text>
    </comment>
    <comment type="subunit">
        <text evidence="10">Homotetramer, and heterotetramer with DPYSL2, DPYSL3, DPYSL4 or DPYSL5. Interacts with PLXNA1. Interacts with FLNA (via calponin-homology (CH) domain 1 and filamin repeat 24); the interaction alters FLNA ternary structure and thus promotes FLNA dissociation from F-actin.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
            <location evidence="2">Microtubule organizing center</location>
            <location evidence="2">Centrosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Spindle</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Perikaryon</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the metallo-dependent hydrolases superfamily. Hydantoinase/dihydropyrimidinase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYV1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008586.1">
        <property type="protein sequence ID" value="ENSGAGP00000007432.1"/>
        <property type="gene ID" value="ENSGAGG00000005962.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043204">
        <property type="term" value="C:perikaryon"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005819">
        <property type="term" value="C:spindle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016812">
        <property type="term" value="F:hydrolase activity, acting on carbon-nitrogen (but not peptide) bonds, in cyclic amides"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd01314">
        <property type="entry name" value="D-HYD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.20.20.140:FF:000217">
        <property type="entry name" value="Dihydropyrimidinase-related protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.40.10:FF:000021">
        <property type="entry name" value="Dihydropyrimidinase-related protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.40.10:FF:000022">
        <property type="entry name" value="Dihydropyrimidinase-related protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.20.20.140">
        <property type="entry name" value="Metal-dependent hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.40.10">
        <property type="entry name" value="Urease, subunit C, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006680">
        <property type="entry name" value="Amidohydro-rel"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011778">
        <property type="entry name" value="Hydantoinase/dihydroPyrase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011059">
        <property type="entry name" value="Metal-dep_hydrolase_composite"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032466">
        <property type="entry name" value="Metal_Hydrolase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050378">
        <property type="entry name" value="Metallo-dep_Hydrolases_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR02033">
        <property type="entry name" value="D-hydantoinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11647:SF54">
        <property type="entry name" value="DIHYDROPYRIMIDINASE-RELATED PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11647">
        <property type="entry name" value="HYDRANTOINASE/DIHYDROPYRIMIDINASE FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01979">
        <property type="entry name" value="Amidohydro_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51338">
        <property type="entry name" value="Composite domain of metallo-dependent hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51556">
        <property type="entry name" value="Metallo-dependent hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="7" id="KW-0944">Nitration</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Amidohydrolase-related" evidence="15">
        <location>
            <begin position="64"/>
            <end position="453"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="515"/>
            <end position="572"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="534"/>
            <end position="543"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004186"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004484"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00008829"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023074"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00059281"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00061880"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00071519"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00079257"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00081558"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="Pfam" id="PF01979"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007432.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="572" mass="62498" checksum="CF841266A866BA7F" modified="2019-05-08" version="1">MSYQGKKSIPHITSDRLLIKGGRIINDDYSFYADIYLEDGLIKQIGENLIVPGGVKTIEANGRMVIPGGIDVNTYLQKPYQGMTAVDDFYQGTKAALAGGTTMIIDHVLPEAGTSLLTAFEKWHEAADTKSCCDYSLHVDITSWYDGIREELDILVQDKGVNSFQVYMAYKDLYQMSDSQLYEAFTFLKGLGAVIMVHAENGDLIAQEQKRILEMGITGPEGHALSRPEELEAEAVFRAITIASRINSPVYITKVMSKSAADIIALARKRGPLVFGEPITASLGTDGTYYWSKNWAKAAAFVTSPPLSPDPTTPDHLNSLLACGDLQVTGSGHCPYSTAQKAIGKDNFTMIPEGVNGIEERMTVVWDKSVAIGKMDENQFVAVTSTNAAKIFNLYPRKGRIAVGSDADVVIWDPDKVKTITAKSHKSAVEYNIFEGMECHGAPLVVISQGKIVFEDGNLHVNKGMGRFIPRKPFPEYLYQRIKIRNKVTGLQGVSRGMYDGPVYEIPATPKYATPALSAKSSPAKNQPPPIRNLHQSNFSLSGAQVDDNNPRRTGHRIVAPPGGRSNITSLG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYV6</accession>
    <name>A0A452GYV6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="19">Phospholipid-transporting ATPase</fullName>
            <ecNumber evidence="19">7.6.2.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="15 19">
            <text>ATP + H2O + phospholipidSide 1 = ADP + phosphate + phospholipidSide 2.</text>
            <dbReference type="EC" id="7.6.2.1"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 18">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Golgi apparatus</location>
            <location evidence="2">trans-Golgi network membrane</location>
            <topology evidence="2">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="19">Membrane</location>
            <topology evidence="19">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 19">Belongs to the cation transport ATPase (P-type) (TC 3.A.3) family. Type IV subfamily.</text>
    </comment>
    <dbReference type="EC" id="7.6.2.1" evidence="19"/>
    <dbReference type="AlphaFoldDB" id="A0A452GYV6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008592.1">
        <property type="protein sequence ID" value="ENSGAGP00000007437.1"/>
        <property type="gene ID" value="ENSGAGG00000000721.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005768">
        <property type="term" value="C:endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005802">
        <property type="term" value="C:trans-Golgi network"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140326">
        <property type="term" value="F:ATPase-coupled intramembrane lipid transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000287">
        <property type="term" value="F:magnesium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006897">
        <property type="term" value="P:endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045332">
        <property type="term" value="P:phospholipid translocation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006890">
        <property type="term" value="P:retrograde vesicle-mediated transport, Golgi to endoplasmic reticulum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.1110.10:FF:000008">
        <property type="entry name" value="Phospholipid-transporting ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1000:FF:000009">
        <property type="entry name" value="Phospholipid-transporting ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.1110.10">
        <property type="entry name" value="Calcium-transporting ATPase, cytoplasmic domain N"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.70.150.10">
        <property type="entry name" value="Calcium-transporting ATPase, cytoplasmic transduction domain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1110.10">
        <property type="entry name" value="Calcium-transporting ATPase, transmembrane domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1000">
        <property type="entry name" value="HAD superfamily/HAD-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023299">
        <property type="entry name" value="ATPase_P-typ_cyto_dom_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023298">
        <property type="entry name" value="ATPase_P-typ_TM_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036412">
        <property type="entry name" value="HAD-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023214">
        <property type="entry name" value="HAD_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006539">
        <property type="entry name" value="P-type_ATPase_IV"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001757">
        <property type="entry name" value="P_typ_ATPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032630">
        <property type="entry name" value="P_typ_ATPase_c"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044492">
        <property type="entry name" value="P_typ_ATPase_HD_dom"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01652">
        <property type="entry name" value="ATPase-Plipid"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01494">
        <property type="entry name" value="ATPase_P-type"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24092:SF50">
        <property type="entry name" value="PHOSPHOLIPID-TRANSPORTING ATPASE IIB-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24092">
        <property type="entry name" value="PROBABLE PHOSPHOLIPID-TRANSPORTING ATPASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00702">
        <property type="entry name" value="Hydrolase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16212">
        <property type="entry name" value="PhoLip_ATPase_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00119">
        <property type="entry name" value="CATATPASE"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDG00002">
        <property type="entry name" value="C1.7:_P-type_atpase_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDS00003">
        <property type="entry name" value="Haloacid_Dehalogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDF00027">
        <property type="entry name" value="p-type_atpase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81665">
        <property type="entry name" value="Calcium ATPase, transmembrane domain M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56784">
        <property type="entry name" value="HAD-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81660">
        <property type="entry name" value="Metal cation-transporting ATPase, ATP-binding domain N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 17 19" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="12" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="13" id="KW-0445">Lipid transport</keyword>
    <keyword evidence="9 18 19" id="KW-0460">Magnesium</keyword>
    <keyword evidence="14 19" id="KW-0472">Membrane</keyword>
    <keyword evidence="6 18" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7 17 19" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10 19" id="KW-1278">Translocase</keyword>
    <keyword evidence="5 19" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 19" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="4" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="107"/>
            <end position="127"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="699"/>
            <end position="719"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="749"/>
            <end position="769"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="775"/>
            <end position="796"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="803"/>
            <end position="822"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="842"/>
            <end position="866"/>
        </location>
    </feature>
    <feature type="domain" description="P-type ATPase C-terminal" evidence="21">
        <location>
            <begin position="637"/>
            <end position="875"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="249"/>
            <end position="271"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="20">
        <location>
            <begin position="249"/>
            <end position="259"/>
        </location>
    </feature>
    <feature type="active site" description="4-aspartylphosphate intermediate" evidence="16">
        <location>
            <position position="192"/>
        </location>
    </feature>
    <feature type="binding site" evidence="17">
        <location>
            <position position="192"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="192"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="17">
        <location>
            <position position="193"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="17">
        <location>
            <position position="194"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="194"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="17">
        <location>
            <position position="327"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="17">
        <location>
            <position position="369"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="17">
        <location>
            <position position="393"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="17">
        <location>
            <position position="422"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="17">
        <location>
            <position position="502"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="17">
        <location>
            <position position="503"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="17">
        <location>
            <position position="504"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="17">
        <location>
            <position position="584"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="17">
        <location>
            <position position="590"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="610"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="17">
        <location>
            <position position="613"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="17">
        <location>
            <position position="614"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="614"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004166"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00008109"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022967"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00034036"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PIRSR" id="PIRSR606539-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="PIRSR" id="PIRSR606539-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="PIRSR" id="PIRSR606539-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="RuleBase" id="RU362033"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="Pfam" id="PF16212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007437.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="883" mass="98573" checksum="64E4D91D08FEC440" modified="2019-05-08" version="1">LKVAVSCTQRLPALGDLFSISAYVYAQKPQLDIHSFEGTFTREDSDPTVHESLSIENTLWASTVVASGTVIGVVIYTGKETRSVMNTSNPKNKVGLLDLELNQLTKALFLALVALSIVMVTLQGFVGPWYRNLFRFLLLFSYIIPISLRVNLDMGKAAYGWMIMKDDNIPGTVVRTSTIPEELGRLVYLLTDKTGASPVALLYSPFRTLTQNEMIFKRLHLGTVSYGTDTMDEIQNHIISSYSQMYSQSSGNSTSSTPSRKVQSSAPKVRKTVSSRIHEAVKAIALCHNVTPVYESHAGVNGETEYAEVDQNFSDENRTYQASSPDEVALVQWTESVGLTLVNRDLTSMQLKSPGGQILTYYILQVFPFTSESKRMGIIVRDESTGEITFYMKGADVAMSTIVQYNDWLEEECGNMAREGLRTLVVAKKSLTEEQHQDFENRYNQAKLSIHDRALKVAAVVESLEREMELLCLTGVEDQLQADVRPTLEMLRNAGIKIWMLTGDKLETATCIAKSSHLVSRTQDIHIFRPVISRGEAHLELNAFRRKHDCALVISGDSLEVCLKYYEHEFVELACQCPAVVCCRCSPTQKAHIVKLLQHHTGKRTCAIGDGGNDVSMIQAADCGIGIEGKEGKQASLAADFSITQFKHIGRLLMVHGRNSYKRSAALGQFVMHRGLIISTMQAVFSSVFYFASIPLYQGFLMVGYATIYTMFPVFSLVLDQDVKPEMALLYPELYKDLTKGRSLSFKTFLIWVLISIYQGGILMYGALLLFESEFVHVVAISFTALILTELLMVALTIRTWHWLMVVAEFFSLGCYVASLAFLNEYFGIGRVSFGAFLDVAFITTVTFLWKVSAITVVSCLPLYILKYLKRKFSPPSYSKLTS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYV8</accession>
    <name>A0A452GYV8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Cytochrome c oxidase assembly homolog COX15</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>Fe(II)-heme o + 2 A + H2O = Fe(II)-heme a + 2 AH2</text>
            <dbReference type="Rhea" id="RHEA:63388"/>
            <dbReference type="ChEBI" id="CHEBI:13193"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:17499"/>
            <dbReference type="ChEBI" id="CHEBI:60530"/>
            <dbReference type="ChEBI" id="CHEBI:61715"/>
            <dbReference type="EC" id="1.17.99.9"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="11">
            <dbReference type="Rhea" id="RHEA:63389"/>
        </physiologicalReaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>heme b</name>
            <dbReference type="ChEBI" id="CHEBI:60344"/>
        </cofactor>
    </comment>
    <comment type="pathway">
        <text evidence="10">Porphyrin-containing compound metabolism; heme A biosynthesis; heme A from heme O: step 1/1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Membrane</location>
            <topology evidence="2">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYV8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008437.1">
        <property type="protein sequence ID" value="ENSGAGP00000007296.1"/>
        <property type="gene ID" value="ENSGAGG00000005857.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005743">
        <property type="term" value="C:mitochondrial inner membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0120547">
        <property type="term" value="F:heme A synthase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016653">
        <property type="term" value="F:oxidoreductase activity, acting on NAD(P)H, heme protein as acceptor"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006784">
        <property type="term" value="P:heme A biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="HAMAP" id="MF_01665">
        <property type="entry name" value="HemeA_synth_type2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003780">
        <property type="entry name" value="COX15/CtaA_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023754">
        <property type="entry name" value="HemeA_Synthase_type2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009003">
        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23289">
        <property type="entry name" value="CYTOCHROME C OXIDASE ASSEMBLY PROTEIN COX15"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23289:SF2">
        <property type="entry name" value="CYTOCHROME C OXIDASE ASSEMBLY PROTEIN COX15 HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02628">
        <property type="entry name" value="COX15-CtaA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50494">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0350">Heme biosynthesis</keyword>
    <keyword evidence="7" id="KW-0408">Iron</keyword>
    <keyword evidence="9 13" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="6" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 13" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 13" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="13">
        <location>
            <begin position="76"/>
            <end position="96"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="13">
        <location>
            <begin position="158"/>
            <end position="175"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="13">
        <location>
            <begin position="187"/>
            <end position="206"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="13">
        <location>
            <begin position="226"/>
            <end position="250"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="13">
        <location>
            <begin position="271"/>
            <end position="294"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="13">
        <location>
            <begin position="328"/>
            <end position="347"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="13">
        <location>
            <begin position="359"/>
            <end position="383"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="13">
        <location>
            <begin position="389"/>
            <end position="406"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="41"/>
            <end position="61"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="41"/>
            <end position="59"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001970"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023133"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00044501"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00048044"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007296.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="16">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="414" mass="46229" checksum="C323C773E65C9D4A" modified="2019-05-08" version="1">MVCIIKQRPLSAQLITLRIPLSGEPPPIPGRWPRLCPARPWRQPQQRRAASEAAAHPGAVAPPVPGSAAERLVGRWLLVCSGTVAGAVVLGGVTRLTESGLSMVDWHLVREMKPPQTQQEWEAEFQKYQQFPEFKILNHDMTLREFKFIWYMEYSHRMWGRVVGLAYVLPAVYFWRKGWLSSSMKGCVLALCGLVCFQGLLGWYMVKSGLEEKPDSHDIPRVSQYRLAAHLGSALVLYCASLWSGLSLLLPQHKLPETYQLLRLRRFAHGTTGLIFLTALSGAFVAGLDAGLVYNSFPKMGERWIPDDLLSFSPVLKNIFENPTTVQFNHRILGITSVTAITALYLLSRKIPLPRRTKIAVASLMAVAYMQVGLGISTLLLYVPTPLAAIHQSGSLALLTVALWLMNELRRVPK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYV9</accession>
    <name>A0A452GYV9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">VLRF1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="domain">
        <text evidence="11">The VLRF1 domain mediates binding to the 60S ribosomal subunit.</text>
    </comment>
    <comment type="similarity">
        <text evidence="2 11">Belongs to the ANKZF1/VMS1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYV9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008375.1">
        <property type="protein sequence ID" value="ENSGAGP00000007244.1"/>
        <property type="gene ID" value="ENSGAGG00000005817.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004519">
        <property type="term" value="F:endonuclease activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016787">
        <property type="term" value="F:hydrolase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036503">
        <property type="term" value="P:ERAD pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047139">
        <property type="entry name" value="ANKZ1/VMS1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041175">
        <property type="entry name" value="VLRF1/Vms1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16036">
        <property type="entry name" value="ANKYRIN REPEAT AND ZINC FINGER DOMAIN-CONTAINING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16036:SF2">
        <property type="entry name" value="TRNA ENDONUCLEASE ANKZF1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00023">
        <property type="entry name" value="Ank"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18826">
        <property type="entry name" value="bVLRF1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS52044">
        <property type="entry name" value="VLRF1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 10" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3 11" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6 11" id="KW-0255">Endonuclease</keyword>
    <keyword evidence="7 11" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="4 11" id="KW-0540">Nuclease</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="VLRF1" evidence="13">
        <location>
            <begin position="199"/>
            <end position="342"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="10">
        <location>
            <begin position="549"/>
            <end position="581"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="107"/>
            <end position="155"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="373"/>
            <end position="474"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="501"/>
            <end position="536"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="630"/>
            <end position="662"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="704"/>
            <end position="752"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="12">
        <location>
            <begin position="414"/>
            <end position="425"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="426"/>
            <end position="439"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="443"/>
            <end position="457"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="643"/>
            <end position="662"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="12">
        <location>
            <begin position="739"/>
            <end position="752"/>
        </location>
    </feature>
    <feature type="active site" evidence="11">
        <location>
            <position position="242"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009262"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022759"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01389"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS52044"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007244.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="752" mass="81660" checksum="ACF5C2ABA673851B" modified="2019-05-08" version="1">MQPLEVRSVFEAAQDPTFLHGLSLVNCSSEDLPSTDCNAITQEKPVVADGEERAQGVPEISDRMCCSSCSRVFDSREEQTEHYRLDWHRFNLKQRLLGRQALTAEEFEEKTRAGDVSSISGSDSDDSDSGSESDLLPPREHTDPGSSQQSLRPGTRSAKVLFRNSQGQLLSAYRCVLGSKKGGGEQQAELVASLQSMGTGTCWVILMTGGGHFAGAVFRGPEVLDHKTFHRYTVRARRGTVQGVRDAQGQASMPKSAGASLRRYNEAALLKDIQDLLASWVQHLQGAQRIFLRVPCANRALLFSGRNPPLHKDDPRICSIPFSTRRATFREVLRVHAALATLQVYGKDTVVADLAGFPRKGWRKVPCARVEEPQQGDASFLSEEENEESQAGELEMVDVTLSTLHLRESEVTPKRSRKKRRKKDKVGRGARAEGPRGQEEPEGSSSQPGAETVTDPQEGSGAEPAPWRKGETPLGQLRDALFTACKTGDVGTLRRLLGVAESGAPPAGSEDGAMGQSQGHPGRVQSEGSEPNHAVPGLLSLLSEPIDGTGFTLLHVAAAAGKGAAVRLLLEAGADPALRDRQERPPYCVSADKSTRNVFRKFMVDHPAKYDYGRAKVPGPLTAEMEARRLEKQRAQKTQRKQRAQELREQQQRLAQEQEEKRRFAALPDREKALLAMWGVPAGPDPLPLPRLLLLLYGLPADSPTGPRRPPLVPAPLGRQGVATHKGACVETDSRLLPAGGGGRGAGHSRGG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYW2</accession>
    <name>A0A452GYW2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Glycolipid transfer protein domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="7">Accelerates the intermembrane transfer of various glycolipids. Catalyzes the transfer of various glycosphingolipids between membranes but does not catalyze the transfer of phospholipids. May be involved in the intracellular translocation of glucosylceramides.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the GLTP family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYW2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008459.1">
        <property type="protein sequence ID" value="ENSGAGP00000007318.1"/>
        <property type="gene ID" value="ENSGAGG00000005875.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1902387">
        <property type="term" value="F:ceramide 1-phosphate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1902388">
        <property type="term" value="F:ceramide 1-phosphate transfer activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.3520.10">
        <property type="entry name" value="Glycolipid transfer protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036497">
        <property type="entry name" value="GLTP_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014830">
        <property type="entry name" value="Glycolipid_transfer_prot_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10219:SF97">
        <property type="entry name" value="GLYCOLIPID TRANSFER PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10219">
        <property type="entry name" value="GLYCOLIPID TRANSFER PROTEIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08718">
        <property type="entry name" value="GLTP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF110004">
        <property type="entry name" value="Glycolipid transfer protein, GLTP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0445">Lipid transport</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <feature type="domain" description="Glycolipid transfer protein" evidence="8">
        <location>
            <begin position="15"/>
            <end position="147"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007148"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00037246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF08718"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007318.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="190" mass="21766" checksum="1F633C2A77F87395" modified="2019-05-08" version="1">MLLLLFRSRRKNPQDCLGTPIVYSPVKADLAGNIKKIRAVYESNPTKFKTLKNILEVEKELYGPAWPKTGATLALMWLKRGLKFIQVLLQSLSDGERDEENPNLIRVNALKAYEIALKKYHGWMLQKLFSGSVYALPYKSDLLKALEKGKEVNEEETIEKIHQFLAKATPVLDAIYDMYTKMNAELNYKA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYW4</accession>
    <name>A0A452GYW4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Transcription factor IIIA</fullName>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYW4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008605.1">
        <property type="protein sequence ID" value="ENSGAGP00000007450.1"/>
        <property type="gene ID" value="ENSGAGG00000005891.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003712">
        <property type="term" value="F:transcription coregulator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042254">
        <property type="term" value="P:ribosome biogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000032">
        <property type="entry name" value="Krueppel-like factor 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001102">
        <property type="entry name" value="Transcription factor IIIA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001998">
        <property type="entry name" value="Transcription factor IIIA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000145">
        <property type="entry name" value="Zinc finger protein 574"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR054599">
        <property type="entry name" value="TFIIIA_Zfn-C2H2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051061">
        <property type="entry name" value="Zinc_finger_trans_reg"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46179:SF1">
        <property type="entry name" value="TRANSCRIPTION FACTOR IIIA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46179">
        <property type="entry name" value="ZINC FINGER PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22110">
        <property type="entry name" value="TFIIIA_zf-C2H2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="6"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="9" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0690">Ribosome biogenesis</keyword>
    <keyword evidence="7" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="10" id="KW-0804">Transcription</keyword>
    <keyword evidence="8" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 13" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="19"/>
            <end position="48"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="49"/>
            <end position="78"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="102"/>
            <end position="128"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="158"/>
            <end position="185"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="187"/>
            <end position="217"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="218"/>
            <end position="247"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="286"/>
            <end position="305"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022517"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00040434"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007450.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="305" mass="35176" checksum="65337DFD5ECA8253" modified="2019-05-08" version="1">MAASFLLRSRPPADCAKRFICSFPDCEATFSKAWRLNAHLCRHTGERPFICNYNGCGKGFTRDFHLTRHFLTHSGEKPFDKSFRKHQQLKVHHCQHTSEPFFKCSHEACGKHFLTPSRLKRHEKIHEGYACKKDNCSFVGKTWTEHLKHMKDCHAEPVICGVCSKTFKRKDYLRQHQKTHAEEREVCRCPREGCGRTYTTVFNLQSHILSFHEEKKPFFCDYAGCGKVFAMKQSLARHAVVHDPDKGKLNLKAKQTRPKRSLASRLSGYIPPKAQRRKLEAATENKALNKPAESELPTVETLTLH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYW6</accession>
    <name>A0A452GYW6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Lysosomal-trafficking regulator</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="8">Interacts with CPAP, LIP8 and ZNF521.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="STRING" id="38772.ENSGAGP00000007324"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008466.1">
        <property type="protein sequence ID" value="ENSGAGP00000007324.1"/>
        <property type="gene ID" value="ENSGAGG00000005878.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098542">
        <property type="term" value="P:defense response to other organism"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006955">
        <property type="term" value="P:immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002449">
        <property type="term" value="P:lymphocyte mediated immunity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043473">
        <property type="term" value="P:pigmentation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd06071">
        <property type="entry name" value="Beach"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd01201">
        <property type="entry name" value="PH_BEACH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:000217">
        <property type="entry name" value="Lysosomal trafficking regulator"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:000292">
        <property type="entry name" value="Lysosomal trafficking regulator"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.1540.10:FF:000001">
        <property type="entry name" value="neurobeachin isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1540.10">
        <property type="entry name" value="BEACH domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000409">
        <property type="entry name" value="BEACH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036372">
        <property type="entry name" value="BEACH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050865">
        <property type="entry name" value="BEACH_Domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023362">
        <property type="entry name" value="PH-BEACH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019775">
        <property type="entry name" value="WD40_repeat_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036322">
        <property type="entry name" value="WD40_repeat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001680">
        <property type="entry name" value="WD40_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13743">
        <property type="entry name" value="BEIGE/BEACH-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13743:SF86">
        <property type="entry name" value="LYSOSOMAL-TRAFFICKING REGULATOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02138">
        <property type="entry name" value="Beach"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14844">
        <property type="entry name" value="PH_BEACH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00400">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01026">
        <property type="entry name" value="Beach"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00320">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81837">
        <property type="entry name" value="BEACH domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50978">
        <property type="entry name" value="WD40 repeat-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50197">
        <property type="entry name" value="BEACH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51783">
        <property type="entry name" value="PH_BEACH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00678">
        <property type="entry name" value="WD_REPEATS_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50082">
        <property type="entry name" value="WD_REPEATS_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50294">
        <property type="entry name" value="WD_REPEATS_REGION"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="11" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="7" id="KW-0653">Protein transport</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0813">Transport</keyword>
    <keyword evidence="5 10" id="KW-0853">WD repeat</keyword>
    <feature type="domain" description="BEACH-type PH" evidence="14">
        <location>
            <begin position="3003"/>
            <end position="3109"/>
        </location>
    </feature>
    <feature type="domain" description="BEACH" evidence="13">
        <location>
            <begin position="3114"/>
            <end position="3416"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="10">
        <location>
            <begin position="3606"/>
            <end position="3640"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="156"/>
            <end position="176"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="913"/>
            <end position="932"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1218"/>
            <end position="1247"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="11">
        <location>
            <begin position="2176"/>
            <end position="2205"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="163"/>
            <end position="176"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="12">
        <location>
            <begin position="1218"/>
            <end position="1232"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="1233"/>
            <end position="1244"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022574"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00064710"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00067676"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50197"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS51783"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007324.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="3795" mass="430056" checksum="1244671E2B3E1F0C" modified="2019-05-08" version="1">MSSASNSLAREFLIDVNRLCNAVVQRAEAREEEEEETHMAALGQYLVQGRGFTLLTTLNSNMDQELTCREELLTLLLSLLPLVWKIPVQEEKAINFNLPFTVDVCLTKETNSNSLRSAHEKQNLEGIVHPSFQTPVKLNSSFRKIRRQRKITHRYSVRDARKSQLSTSDSEGNSDNKTTIMIKHRRSQILQPFVTHLPRDDYHAEVVRSFNPSPISLENSRQIIPTQESKSEVLNESAAAAVVSSMDNSPFDLCHVLLSLLEKVCKFDITLNHNSALAATIVPSLTEFLSGFGDCCNVSGDAESEVVSAGWTEEPVALVQRMLFRTVLHLLSIDITNAETMPDNLKKNLTDLLRAALKIRTCLEKQSDPFVPRHKKTLQEVQDNFVFSKYRHRVLLLPELLEGVLQILICCLQSAASNPFYFSQAVDLVHEFIQHQGFKLLEITVLQMEWLVSKDEEITPEASEHLKALISSIMKIISTVKKVKSEQLHQSMCTRKRHRRCEYSHFMHHHRDLSGLPVSAFKNQVSRNPFEEVVDGEVHYPERCCCIAVCAHQCFHLLQQVSLSSTCAQILSGVHNVGICCCMDPKSVITPLLQASKLPVLKNFQQHILSILNKFILDQLGGAEVSQKIIQAACNICTIDCDQLAELEDTLQSNSTDIGLASNFSCPSYRFQGILPSSGSEDMLLKWDALEAYQDFAFEEDKLRSVQIASHICHLIQKGNVVVQWKLYNYIYNPVLQRGVELAHRSQQLGITAVCNHVCSCHSLCLPQEVLQIYLQTLPALFKSRIIQNLFLSCNGVNQMTELNYLDSVRSHSLKVFETLIFCLGDQQKDQAMPELEGMDNEQKESILDLVTSFRSQQMISDVPQSLSKFYAGLRKAYPKKKKCVSQEVHVNLIHLFLCVAFLCVSKEEECDRDSANDSEDTSGYDSTASEPLSHKLPCLSLESLTLPSPEHIHRAADIWSMCRWIYLCSSVFQRQFHRLGGFEVCHRLIIMIIQKLAKNKEKEQEKKERPLSENNKCLNASPLNQPVFKDDSYYLTVNKQLTQSELNSLDENPDASGQPVSECISCDHSLKVEQISVAMVELEELKKSVKQENEWSLQSIRLLEALLTICLHSANSTQQKAEVELLHQSVCLEDVLLEMRDQLAQSRAIETCLAKPLLDSLLRVALGNYSADLDHSEEKTEKILHTEKASVPQPEDLSEEAEESQCCSFKLFAEEEGYEADSESNPDDSESKDEGTDTKREAECMVPSSYSSDLTENISQGELIYPEICMLELKLLSAGNPSLDVLAHVFQSFLKVIRQKKKNATFLMQQGAVKSLLGGFLNILIQKDSSFHACQIVLVELLVSLMSSRTCAEELTLLLRIFLEETPCTEILLKGILKIVETNIYMSPLQYLSFPLLKSPNLNSSSSQKSTSALNSKTVGLLKRAKFSQNKKEADGENLTHQLLSSWHVAPIHLPLVGQNCWPHMSEGFSVSLWFHVECVQETESPTEKGKKMKKRSKPLIVRESSFDGTEGSKSVRMEYLNLGDKLVEDGCIHIISLGSKALMIQVWADLNTGAFIFRMCMDPNDEMKAGLLAQAESPENIFLVGKWEHLAITYLQQPEGKKNIHGELLLWVSGQRKCEISLDYTLPRKTTLSDSNKTFCMIGHCTSSQEEPLQLAGKWDLGNLLLFNGAKIGPEEAFYLYACGPDFTAIMPCKYGKTMTDYSKYITKDILQCERIKEFFMKKKEADVGPLIESLAVVYTTCCPAQYTIYEPVIRLKGQVKTVPSQRPFSSKEVQSNFLDPYHLKMLQPVEYQTLQGILHEIGGTGAFVFLFARVIDITNCEETQALALQVILSLTKYNQQRIQEMENCNGYSMIHQVLIKPKCIVGFHILKTLLEGCFSDEILNINESGQFMLDVDSTAVIQDIKLLEDLLLDWKIWSKAQQGVWETLLAALEILIRANHHQQMFNIKQLLKARVVHHFLLTCQVLQEQKEGHLTSLPREVCRSFVKIIEEVLGSPPDLELLTLIFNFLLAVHPPTNTYVCHNPTNFYFSLHIDGKIFQEKVQSIKYLRNSSIGGKSVDSPSLTTSPAGFTTSPLGGSIISKIIIPQMMISQALQRRCKSLPASPTSSPQVHPKRLPGSLGRSIDNLQEVWESDSVPQEKNIFVESSETLKRIQEELFVSSCESAKTVCDSELTSAEALVDISKEELELSENKELNTELKSNENGAAVESLLRRPDNLKGLPSFQKSQNNFAGLGLAFSAQSGSSSVACWPSLADRSALPEDWESLTFSSVSEQTFKRSERPDYRSTEDCLVLICCGLYDLLRGVLLILPDIMLEDVIDKLIQPDTLIVLVNHPSPLIQERVIKLLDAYFNRATREQKEQFLKNRGFSLLANQLYLHQGTQELVECFLEMLFGRSVGLDEEFDREDVKNAGLFQKWCVIPVLGLIENSLYDNVLLHNSFCLLLQIINSCSKMADMLLDNGLLYVLCNTLAALNGLETNIPLNDYKLLLCDIQQLLVAVTIHACSSLGSQYFRVIEDLIVLLGHLQNNKNEKTQNMAIALQFRVLQAAIELIKATANQDSQDLANSLRLPATHHHAMFQKRKSIADSRKFSRAQSDTLVMKMRLVASDELNMMMQRRMSQENPIRATENEFIQRLQRLTVLAVNRFIYQEPTEDCFDVLGTPETTIQCRLSTSQLEASEEEDQRELPSNTNSFLKEMFKMMIEGIKVSIATSRISTPRQQWRRILWSCKDTFRMQLGRLLVHILSPAQPLQERKQALEIVCETNHQEIIRDCLSVPLQHGPKLALYLYELMHDHKDELTREEQVSSGVFMNALKFCGYKCISPSAPPKPDLLNAMKEDRKKFEAEECVNKTVWEKTVINNQQNLSQRLDAKSKDISKIAADITQAVSLSQGMERKKVIQHIRGMYKTDLSASRHWQELVQQLTHDRTLWYDPVYYPTSWQLDPTEGPNRERRRLRRCYLTIPNKYLLRDRQKQEEVIKPPLSYLFEDKTHSSHSSTVKDKAASENIRVNRRCINVAPSRETAGELLLGKYGMYFVEDNASDTIESSSFHGETEPASFSWTYEEIKEVHKRWWQLRDNAVEIFLTNGRTLLLAFDNTKVRDDVYHNILMNNLPNLLEYGNITALTHLWYTGQITNFEYLTHLNKHAGRSFNDLMQYPVFPFILSDYTSETLDLNDPSVYRNLAKPIAVQSKEKEDRYVDTYKYLEEEYRKGAREDDPMPPVQPYHYGSHYSNSGTVLHFLVRMPPFTKMFLAYQDQSFDIPDRTFHSTHTTWRLSSYESMTDVKELIPEFFYLPEFLVNREGFDFGLRQNGERVNHVNLPPWARNDPRLFILIHRQALESDHVSQTICNWIDLVFGYKQKGKASVQAINVFHPATYFGMDVSAVEDPVQRRALETMIKTYGQTPRQLFHTAHVSRPGSRLTIEGELPAAMGLLVQFAFRETREHAKEIVYPSPLPWIKGLKWGEYVGSPSAPDPVVCFSQPHGERFGSLQALPTRAICGLSQKFCLLMVYSKEQGVRSMHSTDIQWSAILSWGYADNILRLKSKQSEPPINFIQSSQLHQVTSCAWVPDSCQLFTGSKSGVITAYMNRFSSNMHSEIEMESQVHLYGHTAEITSLFVCKPYSIMISVSKDGTCIIWDLNRLCYVQSLAGHKSPVTAVSASEMTGDIATVCDSVGGGSDLRLWTVNGDLVGHVHCREIICSVAFSNQPEGVSINVIAGGLENGVVRLWSTWDLKPVREITFPKSNKPIVSLTFSCDGHHLYTANSEGTVIAWCRKDQQRLKLPMFYSFLSSYAAG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYW7</accession>
    <name>A0A452GYW7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Lysosomal-trafficking regulator</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="8">Interacts with CPAP, LIP8 and ZNF521.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000008504.1">
        <property type="protein sequence ID" value="ENSGAGP00000007359.1"/>
        <property type="gene ID" value="ENSGAGG00000005878.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098542">
        <property type="term" value="P:defense response to other organism"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006955">
        <property type="term" value="P:immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002449">
        <property type="term" value="P:lymphocyte mediated immunity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043473">
        <property type="term" value="P:pigmentation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd06071">
        <property type="entry name" value="Beach"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd01201">
        <property type="entry name" value="PH_BEACH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:000217">
        <property type="entry name" value="Lysosomal trafficking regulator"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:000292">
        <property type="entry name" value="Lysosomal trafficking regulator"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.1540.10:FF:000001">
        <property type="entry name" value="neurobeachin isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1540.10">
        <property type="entry name" value="BEACH domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000409">
        <property type="entry name" value="BEACH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036372">
        <property type="entry name" value="BEACH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050865">
        <property type="entry name" value="BEACH_Domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023362">
        <property type="entry name" value="PH-BEACH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019775">
        <property type="entry name" value="WD40_repeat_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036322">
        <property type="entry name" value="WD40_repeat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001680">
        <property type="entry name" value="WD40_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13743">
        <property type="entry name" value="BEIGE/BEACH-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13743:SF86">
        <property type="entry name" value="LYSOSOMAL-TRAFFICKING REGULATOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02138">
        <property type="entry name" value="Beach"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14844">
        <property type="entry name" value="PH_BEACH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00400">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01026">
        <property type="entry name" value="Beach"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00320">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81837">
        <property type="entry name" value="BEACH domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50978">
        <property type="entry name" value="WD40 repeat-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50197">
        <property type="entry name" value="BEACH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51783">
        <property type="entry name" value="PH_BEACH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00678">
        <property type="entry name" value="WD_REPEATS_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50082">
        <property type="entry name" value="WD_REPEATS_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50294">
        <property type="entry name" value="WD_REPEATS_REGION"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="11" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="7" id="KW-0653">Protein transport</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0813">Transport</keyword>
    <keyword evidence="5 10" id="KW-0853">WD repeat</keyword>
    <feature type="domain" description="BEACH-type PH" evidence="14">
        <location>
            <begin position="2926"/>
            <end position="3032"/>
        </location>
    </feature>
    <feature type="domain" description="BEACH" evidence="13">
        <location>
            <begin position="3037"/>
            <end position="3339"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="10">
        <location>
            <begin position="3529"/>
            <end position="3563"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="156"/>
            <end position="176"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="913"/>
            <end position="932"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1218"/>
            <end position="1247"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="11">
        <location>
            <begin position="2176"/>
            <end position="2205"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="163"/>
            <end position="176"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="12">
        <location>
            <begin position="1218"/>
            <end position="1232"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="1233"/>
            <end position="1244"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022574"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00064710"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00067676"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50197"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS51783"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007359.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="3718" mass="421457" checksum="FDE5567108432963" modified="2019-05-08" version="1">MSSASNSLAREFLIDVNRLCNAVVQRAEAREEEEEETHMAALGQYLVQGRGFTLLTTLNSNMDQELTCREELLTLLLSLLPLVWKIPVQEEKAINFNLPFTVDVCLTKETNSNSLRSAHEKQNLEGIVHPSFQTPVKLNSSFRKIRRQRKITHRYSVRDARKSQLSTSDSEGNSDNKTTIMIKHRRSQILQPFVTHLPRDDYHAEVVRSFNPSPISLENSRQIIPTQESKSEVLNESAAAAVVSSMDNSPFDLCHVLLSLLEKVCKFDITLNHNSALAATIVPSLTEFLSGFGDCCNVSGDAESEVVSAGWTEEPVALVQRMLFRTVLHLLSIDITNAETMPDNLKKNLTDLLRAALKIRTCLEKQSDPFVPRHKKTLQEVQDNFVFSKYRHRVLLLPELLEGVLQILICCLQSAASNPFYFSQAVDLVHEFIQHQGFKLLEITVLQMEWLVSKDEEITPEASEHLKALISSIMKIISTVKKVKSEQLHQSMCTRKRHRRCEYSHFMHHHRDLSGLPVSAFKNQVSRNPFEEVVDGEVHYPERCCCIAVCAHQCFHLLQQVSLSSTCAQILSGVHNVGICCCMDPKSVITPLLQASKLPVLKNFQQHILSILNKFILDQLGGAEVSQKIIQAACNICTIDCDQLAELEDTLQSNSTDIGLASNFSCPSYRFQGILPSSGSEDMLLKWDALEAYQDFAFEEDKLRSVQIASHICHLIQKGNVVVQWKLYNYIYNPVLQRGVELAHRSQQLGITAVCNHVCSCHSLCLPQEVLQIYLQTLPALFKSRIIQNLFLSCNGVNQMTELNYLDSVRSHSLKVFETLIFCLGDQQKDQAMPELEGMDNEQKESILDLVTSFRSQQMISDVPQSLSKFYAGLRKAYPKKKKCVSQEVHVNLIHLFLCVAFLCVSKEEECDRDSANDSEDTSGYDSTASEPLSHKLPCLSLESLTLPSPEHIHRAADIWSMCRWIYLCSSVFQRQFHRLGGFEVCHRLIIMIIQKLAKNKEKEQEKKERPLSENNKCLNASPLNQPVFKDDSYYLTVNKQLTQSELNSLDENPDASGQPVSECISCDHSLKVEQISVAMVELEELKKSVKQENEWSLQSIRLLEALLTICLHSANSTQQKAEVELLHQSVCLEDVLLEMRDQLAQSRAIETCLAKPLLDSLLRVALGNYSADLDHSEEKTEKILHTEKASVPQPEDLSEEAEESQCCSFKLFAEEEGYEADSESNPDDSESKDEGTDTKREAECMVPSSYSSDLTENISQGELIYPEICMLELKLLSAGNPSLDVLAHVFQSFLKVIRQKKKNATFLMQQGAVKSLLGGFLNILIQKDSSFHACQIVLVELLVSLMSSRTCAEELTLLLRIFLEETPCTEILLKGILKIVETNIYMSPLQYLSFPLLKSPNLNSSSSQKSTSALNSKTVGLLKRAKFSQNKKEADGENLTHQLLSSWHVAPIHLPLVGQNCWPHMSEGFSVSLWFHVECVQETESPTEKGKKMKKRSKPLIVRESSFDGTEGSKSVRMEYLNLGDKLVEDGCIHIISLGSKALMIQVWADLNTGAFIFRMCMDPNDEMKAGLLAQAESPENIFLVGKWEHLAITYLQQPEGKKNIHGELLLWVSGQRKCEISLDYTLPRKTTLSDSNKTFCMIGHCTSSQEEPLQLAGKWDLGNLLLFNGAKIGPEEAFYLYACGPDFTAIMPCKYGKTMTDYSKYITKDILQCERIKEFFMKKKEADVGPLIESLAVVYTTCCPAQYTIYEPVIRLKGQVKTVPSQRPFSSKEVQSNFLDPYHLKMLQPVEYQTLQGILHEIGGTGAFVFLFARVIDITNCEETQALALQVILSLTKYNQQRIQEMENCNGYSMIHQVLIKPKCIVGFHILKTLLEGCFSDEILNINESGQFMLDVDSTAVIQDIKLLEDLLLDWKIWSKAQQGVWETLLAALEILIRANHHQQMFNIKQLLKARVVHHFLLTCQVLQEQKEGHLTSLPREVCRSFVKIIEEVLGSPPDLELLTLIFNFLLAVHPPTNTYVCHNPTNFYFSLHIDGKIFQEKVQSIKYLRNSSIGGKSVDSPSLTTSPAGFTTSPLGGSIISKIIIPQMMISQALQRRCKSLPASPTSSPQVHPKRLPGSLGRSIDNLQEVWESDSVPQEKNIFVESSETLKRIQEELFVSSCESAKTVCDSELTSAEALVDISKEELELSENKELNTELKSNENGAAVESLLRRPDNLKGLPSFQKSQNNFAGLGLAFSAQSGSSSVACWPSLADRSALPEDWESLTFSSVSEQTFKRSERPDYRSTEDCLVLICCGLYDLLRGVLLILPDIMLEDVIDKLIQPDTLIVLVNHPSPLIQERVIKLLDAYFNRATREQKEQFLKNRGFSLLANQLYLHQGTQELVECFLEMLFGRSVGLDEDIPLNDYKLLLCDIQQLLVAVTIHACSSLGSQYFRVIEDLIVLLGHLQNNKNEKTQNMAIALQFRVLQAAIELIKATANQDSQDLANSLRLPATHHHAMFQKRKSIADSRKFSRAQSDTLVMKMRLVASDELNMMMQRRMSQENPIRATENEFIQRLQRLTVLAVNRFIYQEPTEDCFDVLGTPETTIQCRLSTSQLEASEEEDQRELPSNTNSFLKEMFKMMIEGIKVSIATSRISTPRQQWRRILWSCKDTFRMQLGRLLVHILSPAQPLQERKQALEIVCETNHQEIIRDCLSVPLQHGPKLALYLYELMHDHKDELTREEQVSSGVFMNALKFCGYKCISPSAPPKPDLLNAMKEDRKKFEAEECVNKTVWEKTVINNQQNLSQRLDAKSKDISKIAADITQAVSLSQGMERKKVIQHIRGMYKTDLSASRHWQELVQQLTHDRTLWYDPVYYPTSWQLDPTEGPNRERRRLRRCYLTIPNKYLLRDRQKQEEVIKPPLSYLFEDKTHSSHSSTVKDKAASENIRVNRRCINVAPSRETAGELLLGKYGMYFVEDNASDTIESSSFHGETEPASFSWTYEEIKEVHKRWWQLRDNAVEIFLTNGRTLLLAFDNTKVRDDVYHNILMNNLPNLLEYGNITALTHLWYTGQITNFEYLTHLNKHAGRSFNDLMQYPVFPFILSDYTSETLDLNDPSVYRNLAKPIAVQSKEKEDRYVDTYKYLEEEYRKGAREDDPMPPVQPYHYGSHYSNSGTVLHFLVRMPPFTKMFLAYQDQSFDIPDRTFHSTHTTWRLSSYESMTDVKELIPEFFYLPEFLVNREGFDFGLRQNGERVNHVNLPPWARNDPRLFILIHRQALESDHVSQTICNWIDLVFGYKQKGKASVQAINVFHPATYFGMDVSAVEDPVQRRALETMIKTYGQTPRQLFHTAHVSRPGSRLTIEGELPAAMGLLVQFAFRETREHAKEIVYPSPLPWIKGLKWGEYVGSPSAPDPVVCFSQPHGERFGSLQALPTRAICGLSQKFCLLMVYSKEQGVRSMHSTDIQWSAILSWGYADNILRLKSKQSEPPINFIQSSQLHQVTSCAWVPDSCQLFTGSKSGVITAYMNRFSSNMHSEIEMESQVHLYGHTAEITSLFVCKPYSIMISVSKDGTCIIWDLNRLCYVQSLAGHKSPVTAVSASEMTGDIATVCDSVGGGSDLRLWTVNGDLVGHVHCREIICSVAFSNQPEGVSINVIAGGLENGVVRLWSTWDLKPVREITFPKSNKPIVSLTFSCDGHHLYTANSEGTVIAWCRKDQQRLKLPMFYSFLSSYAAG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYX4</accession>
    <name>A0A452GYX4_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="12">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1 7">Belongs to the glycosyl hydrolase 35 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYX4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008387.1">
        <property type="protein sequence ID" value="ENSGAGP00000007254.1"/>
        <property type="gene ID" value="ENSGAGG00000005826.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004565">
        <property type="term" value="F:beta-galactosidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005975">
        <property type="term" value="P:carbohydrate metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.260:FF:000021">
        <property type="entry name" value="Beta-galactosidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.20.20.80:FF:000017">
        <property type="entry name" value="Beta-galactosidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.260">
        <property type="entry name" value="Galactose-binding domain-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.20.20.80">
        <property type="entry name" value="Glycosidases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026283">
        <property type="entry name" value="B-gal_1-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048912">
        <property type="entry name" value="BetaGal1-like_ABD1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048913">
        <property type="entry name" value="BetaGal_gal-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008979">
        <property type="entry name" value="Galactose-bd-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017853">
        <property type="entry name" value="GH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031330">
        <property type="entry name" value="Gly_Hdrlase_35_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001944">
        <property type="entry name" value="Glycoside_Hdrlase_35"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23421">
        <property type="entry name" value="BETA-GALACTOSIDASE RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21317">
        <property type="entry name" value="BetaGal_ABD_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21467">
        <property type="entry name" value="BetaGal_gal-bd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01301">
        <property type="entry name" value="Glyco_hydro_35"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF006336">
        <property type="entry name" value="B-gal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00742">
        <property type="entry name" value="GLHYDRLASE35"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51445">
        <property type="entry name" value="(Trans)glycosidases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49785">
        <property type="entry name" value="Galactose-binding domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="5" id="KW-0326">Glycosidase</keyword>
    <keyword evidence="3" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 8" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="8">
        <location>
            <begin position="1"/>
            <end position="25"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5018983601" evidence="8">
        <location>
            <begin position="26"/>
            <end position="639"/>
        </location>
    </feature>
    <feature type="domain" description="Glycoside hydrolase 35 catalytic" evidence="9">
        <location>
            <begin position="38"/>
            <end position="355"/>
        </location>
    </feature>
    <feature type="domain" description="Beta-galactosidase 1-like first all-beta" evidence="10">
        <location>
            <begin position="402"/>
            <end position="514"/>
        </location>
    </feature>
    <feature type="domain" description="Beta-galactosidase galactose-binding" evidence="11">
        <location>
            <begin position="539"/>
            <end position="596"/>
        </location>
    </feature>
    <feature type="active site" description="Proton donor" evidence="6">
        <location>
            <position position="185"/>
        </location>
    </feature>
    <feature type="active site" description="Nucleophile" evidence="6">
        <location>
            <position position="265"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00009809"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023295"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PIRSR" id="PIRSR006336-1"/>
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    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU003679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF01301"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="Pfam" id="PF21317"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF21467"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007254.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="639" mass="70730" checksum="E8660C67097D1A81" modified="2019-05-08" version="1">MGPAGRVLLLAAALLALLALEVCAGARSFQVDYAHDCFRKDGTPFRYISGSIHYARVPRPAWRDRLLKMYMSGLNAVQVYVPWNYHEPLPGVYDFSGDRDLGHFLDLTAELGLLVILRPGPYICAEWEMGGLPAWLLWKPDIVLRSSDPDYLQAVDSWLDVLLPRIKPRLYQNGGNIISIQVENEYGSYSACDYDYLRHLLAAFRTRLGPDVLLFTTDGNSAAELRCGTLQGLYTTVDFGPGPNVTAAFAPQRLYEPKGPLVNSEYYTGWLDYWGEPHASSSPVQVARGLQDMLQLGANVNMYMFQGGTNFGYWSGADYKGCYKPVTTSYDYDAPLSEAGDPTEKLFAIRTVIGKFQPLPEGPMPPATPKYAYGHVPLRKQGEVLELLDVLCPGGPIRSRFPRTFEAVKQAHGFVLYRTRLPRDVWDPAPLSTPPNSVCDRAYVLLNGEYQGRLERDGQMTLNLTGQAGARLDLLVENMGRINFGVNASDFKGLLQNLSLDSALLSDWLIYPLDVDSAVTRGWPLPTTQAEGSRAGLGPALYTGAFQTPGVAWDTFVKFPGWSKGQLWINGFNVGRYWPVRGPQQTLFVPGSLLSASAPNNITLLELEGAPPRPFVLFLDRPLFNQTVSARAGTELRPP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYX7</accession>
    <name>A0A452GYX7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="22">Platelet endothelial aggregation receptor 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="11">Belongs to the MEGF family.</text>
    </comment>
    <comment type="caution">
        <text evidence="12">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYX7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008515.1">
        <property type="protein sequence ID" value="ENSGAGP00000007369.1"/>
        <property type="gene ID" value="ENSGAGG00000005918.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="CDD" id="cd00055">
        <property type="entry name" value="EGF_Lam"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.170.300.10:FF:000002">
        <property type="entry name" value="Multiple epidermal growth factor-like domains 10"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.170.300.10:FF:000007">
        <property type="entry name" value="multiple epidermal growth factor-like domains protein 10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000114">
        <property type="entry name" value="Multiple epidermal growth factor-like domains protein 11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.170.300.10:FF:000041">
        <property type="entry name" value="Tyrosine protein kinase receptor tie-1, putative"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.170.300.10">
        <property type="entry name" value="Tie2 ligand-binding domain superfamily"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013111">
        <property type="entry name" value="EGF_extracell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057138">
        <property type="entry name" value="EGF_PEAR1L-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011489">
        <property type="entry name" value="EMI_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002049">
        <property type="entry name" value="LE_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052485">
        <property type="entry name" value="MEGF_diff_regulators"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24052">
        <property type="entry name" value="DELTA-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24052:SF12">
        <property type="entry name" value="PLATELET ENDOTHELIAL AGGREGATION RECEPTOR 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07974">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00053">
        <property type="entry name" value="EGF_laminin"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23301">
        <property type="entry name" value="EGF_PEAR1L"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00011">
        <property type="entry name" value="EGFLAMININ"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="16"/>
    </dbReference>
    <dbReference type="SMART" id="SM00180">
        <property type="entry name" value="EGF_Lam"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01186">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50027">
        <property type="entry name" value="EGF_LAM_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51041">
        <property type="entry name" value="EMI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="9 12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3 12" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13" id="KW-0424">Laminin EGF-like domain</keyword>
    <keyword evidence="8 15" id="KW-0472">Membrane</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5 16" id="KW-0732">Signal</keyword>
    <keyword evidence="4 15" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 15" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="16">
        <location>
            <begin position="1"/>
            <end position="18"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019057365" description="Platelet endothelial aggregation receptor 1" evidence="16">
        <location>
            <begin position="19"/>
            <end position="1072"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="800"/>
            <end position="823"/>
        </location>
    </feature>
    <feature type="domain" description="EMI" evidence="19">
        <location>
            <begin position="23"/>
            <end position="101"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="17">
        <location>
            <begin position="181"/>
            <end position="216"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="17">
        <location>
            <begin position="224"/>
            <end position="260"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="17">
        <location>
            <begin position="273"/>
            <end position="303"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="17">
        <location>
            <begin position="316"/>
            <end position="346"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="18">
        <location>
            <begin position="363"/>
            <end position="406"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="17">
        <location>
            <begin position="405"/>
            <end position="435"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="17">
        <location>
            <begin position="486"/>
            <end position="521"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="18">
        <location>
            <begin position="537"/>
            <end position="584"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="17">
        <location>
            <begin position="577"/>
            <end position="607"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="17">
        <location>
            <begin position="615"/>
            <end position="650"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1013"/>
            <end position="1072"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="206"/>
            <end position="215"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="250"/>
            <end position="259"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="293"/>
            <end position="302"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="336"/>
            <end position="345"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="13">
        <location>
            <begin position="382"/>
            <end position="391"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="425"/>
            <end position="434"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="511"/>
            <end position="520"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="13">
        <location>
            <begin position="554"/>
            <end position="563"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="597"/>
            <end position="606"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="640"/>
            <end position="649"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004162"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00038377"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00460"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50027"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS51041"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007369.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="22">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1072" mass="114460" checksum="F5CB4EFF08735627" modified="2019-05-08" version="1">MKLHVVLLLGLQMHLATSLNPSDPNVCSYWESFTTATKESYAHPYAQASKDSCDGTWSFLKPCTQHKIVYKTAYRQAVKIDYRKRYRCCQGYYESADVCVPRCTKECVHGRCVAPDQCQCEQGWRGTDCSSVCNGQSWGPHCENPCQCGDGGACDPLTGACVCSPGYKDSMCKVPCDPGTYGEGCQLACPCKNTDRCHGETGACLCQPGFTGTYCELLCLNSSDGLHCPAYCPCQNGGICHPPNTTHCVCPPGWMGTICSIPCPQGQYGSGCLGECQCHNNGLCDPVTGRCQCALGYTGERCREECPIGKYGQDCGQPCDCANGGRCFHINGACLCEAGFYGPRCEERKCLDGLYGLACHLPCLCNPQNSQSCHPMSGECTCKPGWAGLHCNETCPHGYYGANCQEPCLCLNGGTCDGATGLCHCAPGYTGDHCSSHCPTDTYGVNCSMQCACKNAFACSPIDGTCVCKEGWHGGDCSRPCPTGTWGPGCNETCRCANGAACSPVSGACSCAAGWHGSRCQEPCTNGAYGPGCSSQCDCSHADGCSPVTGQCRCLPGWTGPRCNLVCDDGFWGRNCNQPCACKNGAACSSEDGSCSCAPGYRGPACQRPCQPGRYGKKCAMLCRCSNHSTCHPVDGSCDCFPGWTGSDCSQPCPPGTWGEACGHTCRCPNGGWCSTVDGTCSCPAGLTGRRCSEGCPPGSYGENCAQACQCQNMAQCDPATGRCLCSSGYTGAHCETKCFPGTFGPGCLQVCDCLHNSTCHHVTGSCQCELGRIGARCEKWSPDHPFTMVPAVPVGYSSLGAIIGIIILVILLVALLVFFLCYRHWQKGKENRHLSVAYTTGRTDSSEYAVPDVPPSYTHYYSNPSYHTLSRCTPTSPAPSSQDRASSLKVPCNQHFSNTKSMAYGPDCNASLPADWKHHGAPALKPRERGGGPMDRSYSYSNSLGKYYSNAYIKEEALRASNSSLNSENPYATIKDLPELMAKPSEGSYMEMKSPVKREMSYAEIRLFEEPAAGSRAEETCPPGAEGVSPAAPSSPPNHYDSPKNSHIPSHYDMPPVRHYPPSPPLRRQDR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYX8</accession>
    <name>A0A452GYX8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Protein sel-1 homolog 1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="12">Suppressor of lin-12-like protein 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">Plays a role in the endoplasmic reticulum quality control (ERQC) system also called ER-associated degradation (ERAD) involved in ubiquitin-dependent degradation of misfolded endoplasmic reticulum proteins. Enhances SYVN1 stability. Plays a role in LPL maturation and secretion. Required for normal differentiation of the pancreas epithelium, and for normal exocrine function and survival of pancreatic cells. May play a role in Notch signaling.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="9">Belongs to the sel-1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYX8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008485.1">
        <property type="protein sequence ID" value="ENSGAGP00000007341.1"/>
        <property type="gene ID" value="ENSGAGG00000004194.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036503">
        <property type="term" value="P:ERAD pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007219">
        <property type="term" value="P:Notch signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000208">
        <property type="entry name" value="Protein sel-1 homolog 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000193">
        <property type="entry name" value="protein sel-1 homolog 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000200">
        <property type="entry name" value="protein sel-1 homolog 1 precursor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.10">
        <property type="entry name" value="Tetratricopeptide repeat domain"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006597">
        <property type="entry name" value="Sel1-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050767">
        <property type="entry name" value="Sel1_AlgK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011990">
        <property type="entry name" value="TPR-like_helical_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11102">
        <property type="entry name" value="SEL-1-LIKE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11102:SF147">
        <property type="entry name" value="SEL1L ADAPTOR SUBUNIT OF ERAD E3 UBIQUITIN LIGASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08238">
        <property type="entry name" value="Sel1"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="SMART" id="SM00671">
        <property type="entry name" value="SEL1"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81901">
        <property type="entry name" value="HCP-like"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="8" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7 13" id="KW-0472">Membrane</keyword>
    <keyword evidence="5" id="KW-0914">Notch signaling pathway</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 14" id="KW-0732">Signal</keyword>
    <keyword evidence="2 13" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 13" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="14">
        <location>
            <begin position="1"/>
            <end position="21"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019207463" description="Protein sel-1 homolog 1" evidence="14">
        <location>
            <begin position="22"/>
            <end position="624"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="13">
        <location>
            <begin position="575"/>
            <end position="593"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004115"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022976"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00038101"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00054921"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00068339"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00076627"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007341.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="624" mass="69934" checksum="FBC410D61274F106" modified="2019-05-08" version="1">SKTSLLHLIKPLIFFLFLAEEQSNKRRQMQEAEDVYQTGMKIFNESSKKAQKKEAYQYLLKAADMNHTKAMEKVSYALLFGDYLKQNIQSAKELFEKLTEEGSPKGQTVSVRVAVSYYAILSSVLLLTAGYRYWGGIGVLQSCESALTHYRLVANHVASDISLTGGTVVQRIRLADEVENPGMASGMLEEDLIQYYQFLAEKGDVQAQVGLGQLHLHGGRGVEQNHQRAFEYFNQAANAGNSHAMAFLGKLYSEGSDIIPQSNETALQYFKKAADMGNPVGQSGLGMAYLYGRGVPVNYELALKYFQKAAEQGWVDGQLQLGSMYYNGIGVKRDYKQALKYFNLASQGGHILAFYNLAQMHATGTGVMRSCHTAVELFKNVCERGRWSERLMTAYNSYKDGDSNAAVVQYLLLAEQGYEVAQSNAAFILDQKEASIVGENETYPQALLHWNRAASQGYTVARIKLGDYHFYGFGTDVDYETAFIHYRLASEQQHSAQAMFNLGYMHEKGLGIKQDIHLAKRFYDMAAEASPDAQVPVFLALCKLGIVYSLQYVRETNIREIVSHLDMDQLLGPEWDLYLMTIIALLLGTVIAYRQRQHQAILRPVGLRPAVPQQEPPPEHQPPQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYY4</accession>
    <name>A0A452GYY4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Zinc finger protein RFP-like</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GYY4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008626.1">
        <property type="protein sequence ID" value="ENSGAGP00000007470.1"/>
        <property type="gene ID" value="ENSGAGG00000005976.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd19762">
        <property type="entry name" value="Bbox2_TRIM7-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd16594">
        <property type="entry name" value="RING-HC_TRIM7-like_C-IV"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12888">
        <property type="entry name" value="SPRY_PRY_TRIM7_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.920:FF:000004">
        <property type="entry name" value="Butyrophilin subfamily 1 member A1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.920">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001870">
        <property type="entry name" value="B30.2/SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043136">
        <property type="entry name" value="B30.2/SPRY_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003879">
        <property type="entry name" value="Butyrophylin_SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006574">
        <property type="entry name" value="PRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003877">
        <property type="entry name" value="SPRY_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050143">
        <property type="entry name" value="TRIM/RBCC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000315">
        <property type="entry name" value="Znf_B-box"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001841">
        <property type="entry name" value="Znf_RING"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017907">
        <property type="entry name" value="Znf_RING_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24103">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE TRIM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13765">
        <property type="entry name" value="PRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00622">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00643">
        <property type="entry name" value="zf-B_box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15227">
        <property type="entry name" value="zf-C3HC4_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01407">
        <property type="entry name" value="BUTYPHLNCDUF"/>
    </dbReference>
    <dbReference type="SMART" id="SM00336">
        <property type="entry name" value="BBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00589">
        <property type="entry name" value="PRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00184">
        <property type="entry name" value="RING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00449">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57845">
        <property type="entry name" value="B-box zinc-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57850">
        <property type="entry name" value="RING/U-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50188">
        <property type="entry name" value="B302_SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50119">
        <property type="entry name" value="ZF_BBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00518">
        <property type="entry name" value="ZF_RING_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50089">
        <property type="entry name" value="ZF_RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="1" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0862">Zinc</keyword>
    <keyword evidence="2 4" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="RING-type" evidence="6">
        <location>
            <begin position="16"/>
            <end position="57"/>
        </location>
    </feature>
    <feature type="domain" description="B box-type" evidence="7">
        <location>
            <begin position="94"/>
            <end position="131"/>
        </location>
    </feature>
    <feature type="domain" description="B30.2/SPRY" evidence="8">
        <location>
            <begin position="261"/>
            <end position="449"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="5">
        <location>
            <begin position="190"/>
            <end position="239"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00024"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50119"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50188"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007470.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="11">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="449" mass="50873" checksum="DCE0A19F5C041235" modified="2019-05-08" version="1">MAAENPVESLQEEATCPICLEYFTEPVTLECGHNFCRACIAQCWEGSDTDISCPQCRETVQQRNLRPNRQLANVVEIVKRLSLQAAKGAGGDGVCGEHQEALKLFCEEDQSPICVICRESRAHCAHTVVPIQEAAQEYKEKIQFSLLHAQLSMKGPQVPFISLTSPFHIFIQKQTQTERQKIVSEFQELREFLEEQERLLLAQLEKLDEEIVRIQNENVSKLSEQISRLSELISEMEGKCQKPGSEFLKVRLREKQRRRLHGGLRKTIVLLEALRKFKANVTLDPDTAQLRLVLSEDRKSVRYGDTRQDLPDNPQRFDTDFCVLGCEGFTSGRHCWEVEVEDGQCWAVGVARESVGRKGGISYSPEGGIWAVLRWMGQFQVLTSPVTPLPLRRVPSRIQVCLDCDRGQVTFIDAGDEAPIFTFPPGSIPGERIRPWLCVGYRSRLCLCP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYY9</accession>
    <name>A0A452GYY9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Ion transport domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYY9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008526.1">
        <property type="protein sequence ID" value="ENSGAGP00000007379.1"/>
        <property type="gene ID" value="ENSGAGG00000005923.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005227">
        <property type="term" value="F:calcium-activated cation channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0099604">
        <property type="term" value="F:ligand-gated calcium channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005821">
        <property type="entry name" value="Ion_trans_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050927">
        <property type="entry name" value="TRPM"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13800:SF6">
        <property type="entry name" value="TRANSIENT RECEPTOR POTENTIAL CATION CHANNEL SUBFAMILY M MEMBER 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13800">
        <property type="entry name" value="TRANSIENT RECEPTOR POTENTIAL CATION CHANNEL, SUBFAMILY M, MEMBER 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00520">
        <property type="entry name" value="Ion_trans"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4 5" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="2 5" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="3 5" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="6">
        <location>
            <begin position="1"/>
            <end position="17"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019120394" description="Ion transport domain-containing protein" evidence="6">
        <location>
            <begin position="18"/>
            <end position="249"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="37"/>
            <end position="59"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="79"/>
            <end position="96"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="150"/>
            <end position="175"/>
        </location>
    </feature>
    <feature type="domain" description="Ion transport" evidence="7">
        <location>
            <begin position="45"/>
            <end position="187"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF00520"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007379.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="249" mass="28321" checksum="1AF610B90F5C0542" modified="2019-05-08" version="1">MPPRITLAFFFFSLCCAGGLDPPSSHSSPPRMFPRSYASGRTVLCLDFMVFTLRLIHIFAVNKQLGPKMIIVGKMMKDVFLFLFFLAVWLVAYGVTTEGLLHPSDQRLAWIFRRVFYRPYLQIFGQIPLNEIDGAGRGSPLEAEPCTNTYANWLVLVLLVIFLLVANILLLNLLIAMFSYTFAKVQGNSDTYWKSQRYSLILEYHNRPALAPPFIIIRHHGLPDTPCCEPRRPGTMDCPVGLEFRSLLL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ40</accession>
    <name>A0A452GZ40_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase</fullName>
            <ecNumber evidence="11">2.7.4.24</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="11">Bifunctional inositol kinase that acts in concert with the IP6K kinases to synthesize the diphosphate group-containing inositol pyrophosphates diphosphoinositol pentakisphosphate, PP-InsP5, and bis-diphosphoinositol tetrakisphosphate, (PP)2-InsP4. PP-InsP5 and (PP)2-InsP4, also respectively called InsP7 and InsP8, may regulate a variety of cellular processes, including apoptosis, vesicle trafficking, cytoskeletal dynamics, and exocytosis. Phosphorylates inositol hexakisphosphate (InsP6).</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="10">
            <text>1D-myo-inositol hexakisphosphate + ATP = 1-diphospho-1D-myo-inositol 2,3,4,5,6-pentakisphosphate + ADP</text>
            <dbReference type="Rhea" id="RHEA:37459"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:58130"/>
            <dbReference type="ChEBI" id="CHEBI:74946"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.4.24"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="10">
            <dbReference type="Rhea" id="RHEA:37460"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="9">
            <text>5-diphospho-1D-myo-inositol 1,2,3,4,6-pentakisphosphate + ATP + H(+) = 1,5-bis(diphospho)-1D-myo-inositol 2,3,4,6-tetrakisphosphate + ADP</text>
            <dbReference type="Rhea" id="RHEA:10276"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:58628"/>
            <dbReference type="ChEBI" id="CHEBI:77983"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.4.24"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="9">
            <dbReference type="Rhea" id="RHEA:10277"/>
        </physiologicalReaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 11">Cytoplasm</location>
            <location evidence="1 11">Cytosol</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 11">Belongs to the histidine acid phosphatase family. VIP1 subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.4.24" evidence="11"/>
    <dbReference type="AlphaFoldDB" id="A0A452GZ40"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008599.1">
        <property type="protein sequence ID" value="ENSGAGP00000007444.1"/>
        <property type="gene ID" value="ENSGAGG00000005969.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033857">
        <property type="term" value="F:5-diphosphoinositol pentakisphosphate 1-kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0052723">
        <property type="term" value="F:inositol hexakisphosphate 1-kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="RHEA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006020">
        <property type="term" value="P:inositol metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032958">
        <property type="term" value="P:inositol phosphate biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07061">
        <property type="entry name" value="HP_HAP_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.470.20:FF:000003">
        <property type="entry name" value="Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.11950:FF:000001">
        <property type="entry name" value="Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.11950:FF:000003">
        <property type="entry name" value="Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1240:FF:000013">
        <property type="entry name" value="Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.11950">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.470.20">
        <property type="entry name" value="ATP-grasp fold, B domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1240">
        <property type="entry name" value="Phosphoglycerate mutase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033379">
        <property type="entry name" value="Acid_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000560">
        <property type="entry name" value="His_Pase_clade-2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037446">
        <property type="entry name" value="His_Pase_VIP1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029033">
        <property type="entry name" value="His_PPase_superfam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040557">
        <property type="entry name" value="VIP1_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12750">
        <property type="entry name" value="DIPHOSPHOINOSITOL PENTAKISPHOSPHATE KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12750:SF10">
        <property type="entry name" value="INOSITOL HEXAKISPHOSPHATE AND DIPHOSPHOINOSITOL-PENTAKISPHOSPHATE KINASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00328">
        <property type="entry name" value="His_Phos_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18086">
        <property type="entry name" value="PPIP5K2_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56059">
        <property type="entry name" value="Glutathione synthetase ATP-binding domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53254">
        <property type="entry name" value="Phosphoglycerate mutase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00616">
        <property type="entry name" value="HIS_ACID_PHOSPHAT_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 11" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="3 11" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7 11" id="KW-0418">Kinase</keyword>
    <keyword evidence="6 11" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 11" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="VIP1 N-terminal" evidence="13">
        <location>
            <begin position="54"/>
            <end position="143"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1"/>
            <end position="21"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="908"/>
            <end position="958"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1159"/>
            <end position="1193"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="925"/>
            <end position="955"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="1161"/>
            <end position="1171"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="1180"/>
            <end position="1193"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004514"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005609"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00033696"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00034629"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="RuleBase" id="RU365032"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="Pfam" id="PF18086"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007444.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1193" mass="134620" checksum="E0A9940680FDCAF6" modified="2019-05-08" version="1">MSVSATESDPPRFFVGGEDGDELLDSSRSADFDHFYDHAEEEDEEYDSPPERLIVVGICSMAKKSKSKPMNEILERLSMFKYITVVIFEEDVILNEPVENWPLCDCLISFHSKGFPLDKAVAYAKLRNPFVINDLNMQYHIQDRREVYSILKAEGILLPRYAILNRDPNNPKECSLIEGEDQVEVNGEVFQKPFVEKPVSAEDHNVYIYYPTSAGGGSQRLFRKIGSRSSVYSPESNVRKTGSYIYEEFMPTDGTDVKVYTVGPDYAHAEARKSPALDGKVERDSEGKEVRYPVILNAREKLIAWKVCLAFKQTVCGFDLLRANGQSYVCDVNGFSFVKNSMKYYDDCAKILGNIIMRELAPQFQIPWSIPLEAEDIPIVPTTSGTMMELRCVIAVIRHGDRTPKQKMKMEVKHQKFFDLFEKCDGYKSGKLKLKKPKQLQEVLDIARQLLIELGQNNDSEIEESKAKLEQLKTVLEMYGHFSGINRKVQLTYLPHGCQKTSSEEEDNRRDEPSLLLVLKWGGELTPAGRVQAEELGRAFRCMYPGGQGDYAGFPGCGLLRLHSTYRHDLKIYASDEGRVQMTAAAFAKGLLALEGELTPILVQMVKSANMNGLLDSDSDSLSSCQHRVKARLHEILQRDREFMPEDYEKLTPSGSISLMKSMKVIRNPVRTCDKVYSLIQSLTSQIRQRMEDPKSADIQLYHSETLELMLRRWAKLEKDFKTKNGRYDISKIPDIYDCIKYDVQHNGSLKLENTMELYRLSKALADIVIPQEYGISKVEKLEIAKGYCTPLVRKIRSDLQRTQDDDTVNKLHPLYSRGVMSPERHVRTRLYFTSESHVHSLLSTLRYGALCDESKDEQWKRAMDYLNIVSELNYMTQIVIMLYEDPNKELSSEERFHVELHFSPGAKGCEEDKNLPAGFGYRPASRENEGSKKTSHKNDSDEESHTSKRDETDRSIVVFKPMVSDPIHIHRKSPLPRSRKIGSVEVLSDNNSNLRTPRTILEQKQSGLGSHCAGLFSTSVLAGSSSAPNLQDYARTQRKKLTSSGCLDDTTRGSAVKRFSISFARHPTNGFELYSMVPSICPLETLHNSLSLKQVDEFLASIAAPSSDNLPEISTALCASSAYLSSLSGRKCSLNTYNPAKIQPTPLAALPERLAIEKPTLSTSGPSNHISDVAPSPKESVKCGDETLTEKK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ43</accession>
    <name>A0A452GZ43_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Alpha-mannosidase</fullName>
            <ecNumber evidence="16">3.2.1.-</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Catalyzes the first committed step in the biosynthesis of complex N-glycans. It controls conversion of high mannose to complex N-glycans; the final hydrolytic step in the N-glycan maturation pathway.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>N(4)-{beta-D-GlcNAc-(1-&gt;2)-alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;6)]-alpha-D-Man-(1-&gt;6)]-beta-D-Man-(1-&gt;4)-beta-D-GlcNAc-(1-&gt;4)-beta-D-GlcNAc}-L-asparaginyl-[protein] + 2 H2O = 2 alpha-D-mannopyranose + an N(4)-{beta-D-GlcNAc-(1-&gt;2)-alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;6)]-beta-D-Man-(1-&gt;4)-beta-D-GlcNAc-(1-&gt;4)-beta-D-GlcNAc}-L-asparaginyl-[protein]</text>
            <dbReference type="Rhea" id="RHEA:56052"/>
            <dbReference type="Rhea" id="RHEA-COMP:14368"/>
            <dbReference type="Rhea" id="RHEA-COMP:14369"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:28729"/>
            <dbReference type="ChEBI" id="CHEBI:60615"/>
            <dbReference type="ChEBI" id="CHEBI:60625"/>
            <dbReference type="EC" id="3.2.1.114"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="16">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
        <text evidence="16">Binds 1 zinc ion per subunit.</text>
    </comment>
    <comment type="pathway">
        <text evidence="2">Protein modification; protein glycosylation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Golgi apparatus membrane</location>
            <topology evidence="1">Single-pass type II membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 16">Belongs to the glycosyl hydrolase 38 family.</text>
    </comment>
    <dbReference type="EC" id="3.2.1.-" evidence="16"/>
    <dbReference type="AlphaFoldDB" id="A0A452GZ43"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008709.1">
        <property type="protein sequence ID" value="ENSGAGP00000007544.1"/>
        <property type="gene ID" value="ENSGAGG00000002370.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000139">
        <property type="term" value="C:Golgi membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030246">
        <property type="term" value="F:carbohydrate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004572">
        <property type="term" value="F:mannosyl-oligosaccharide 1,3-1,6-alpha-mannosidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006013">
        <property type="term" value="P:mannose metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006491">
        <property type="term" value="P:N-glycan processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd11667">
        <property type="entry name" value="GH38N_Man2A2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1270.50:FF:000001">
        <property type="entry name" value="Alpha-mannosidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.70.98.30:FF:000002">
        <property type="entry name" value="Alpha-mannosidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.20.110.10:FF:000003">
        <property type="entry name" value="Alpha-mannosidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.20.110.10">
        <property type="entry name" value="Glycoside hydrolase 38, N terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.50">
        <property type="entry name" value="Glycoside hydrolase family 38, central domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1180">
        <property type="entry name" value="Golgi alpha-mannosidase II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.70.98.30">
        <property type="entry name" value="Golgi alpha-mannosidase II, domain 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011013">
        <property type="entry name" value="Gal_mutarotase_sf_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011330">
        <property type="entry name" value="Glyco_hydro/deAcase_b/a-brl"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011682">
        <property type="entry name" value="Glyco_hydro_38_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015341">
        <property type="entry name" value="Glyco_hydro_38_cen"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037094">
        <property type="entry name" value="Glyco_hydro_38_cen_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000602">
        <property type="entry name" value="Glyco_hydro_38_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027291">
        <property type="entry name" value="Glyco_hydro_38_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028995">
        <property type="entry name" value="Glyco_hydro_57/38_cen_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013780">
        <property type="entry name" value="Glyco_hydro_b"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050843">
        <property type="entry name" value="Glycosyl_Hydrlase_38"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11607">
        <property type="entry name" value="ALPHA-MANNOSIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11607:SF57">
        <property type="entry name" value="ALPHA-MANNOSIDASE 2X"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09261">
        <property type="entry name" value="Alpha-mann_mid"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07748">
        <property type="entry name" value="Glyco_hydro_38C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01074">
        <property type="entry name" value="Glyco_hydro_38N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00872">
        <property type="entry name" value="Alpha-mann_mid"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF88688">
        <property type="entry name" value="Families 57/38 glycoside transferase middle domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF74650">
        <property type="entry name" value="Galactose mutarotase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF88713">
        <property type="entry name" value="Glycoside hydrolase/deacetylase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="13 16" id="KW-0326">Glycosidase</keyword>
    <keyword evidence="10" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="6 16" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="11 17" id="KW-0472">Membrane</keyword>
    <keyword evidence="5 16" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0735">Signal-anchor</keyword>
    <keyword evidence="4 17" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9 17" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="7 16" id="KW-0862">Zinc</keyword>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="7"/>
            <end position="26"/>
        </location>
    </feature>
    <feature type="domain" description="Glycoside hydrolase family 38 central" evidence="18">
        <location>
            <begin position="502"/>
            <end position="588"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004323"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004922"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00009792"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023295"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00059516"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00093232"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="RuleBase" id="RU361199"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="SMART" id="SM00872"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007544.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1144" mass="129421" checksum="EC0E9AF7BACBC482" modified="2019-05-08" version="1">MKLKKQVTVCGAAIFCVAVFSLYLMLDRVQHDPARHQSGGNFPRSQISVLQNRIEQLEQLLEENHEIISHIKESVLELTAHVDGQPVLLPYRTPNGSWVLPPESRPSFFSISPQDCQFALGSGGQKPDLQMLAVASVLPFDNQDGGVWKQGFDITYEPHDWDTEPLQVFVVPHSHNDPGWIKTFEKYYADQTQHILNSMVLKLQEDPRRRFIWSEISFFSKWWDNISTQKRAAVRRLVGGGQLEMVTGGWVMPDEANSHYFAMIDQLIEGHQWLEKNIGVTPRSGWAVDPFGHSATMPYLLRRANLTSMLIQRVHYTIKKHFAATRSLEFMWRQTWDPDSSTDIFCHMMPFYSYDVPHTCGPDPKICCQFDFKRLPGGRINCPWKVPPRAITEANVAERAHLLLDQYRKKSKLFRSKVLLVPLGDDFRYDKPQEWDAQFLNYQRLFDFLNAHPSLHVQAQFGTLSDYFDALSKRTGIIPGLKPPGFPVLSGDFFSYADREDHYWTGYYTSRPFYKSLDRVLEAHLRGAEILYSLALSHARHAGMDSRYPLSDYALLTEARRALGLFQHHDAITGTAKEAVVVDYGVRLLHSLVNLKRIISNAAHYLVLGEKESYHYDAAGPFLGMDDMRLNQDSLPEKTIIKLDTSPRCTPSSGLGQALSLLVNSPRVRVLSEEGQPLAIQLSAQWSSATDMVPDVYQVSIMTRLPALGLSILQLYKSFDSHSTLTSSVHIHLHGRDLPVRPREVFPVRIIPATSDDFCLDNQHMQACFSGLTGLLQSVRRAGEEHRLSSEFLIYGTRSAKDKSGAYLFLPDGDAKPYAPKEPPVVRVTEGPFFSEVASYYQHVQQVVRLYNVPGVEGLSLDVSCLVDIRDHVNKEMALRLSTTIASEDTFFTDLNGFQIQPRRFLKKLPLQANFYPMPAMAYIQDKQSRLTLHTAQALGVSSLHSGQLEVILDRRLMQDDNRGLGQGLKDNKRTCNRFRLLLERRTTANRVQDNRPISFPSLLSHMTSMHLNAEVLAMPVAREKPAPPALRSFRPLSATVPCDFHILNLRTLQAEDDLLPSAEIALILHRKGFDCGLEAKHLGFNCTTSQGKFSLGSLFHGLELGSLQPTSLTLMYPLGAASPNSTVIHMDPMEIATFRIHFE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ44</accession>
    <name>A0A452GZ44_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="7">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the NIPA family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZ44"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008578.1">
        <property type="protein sequence ID" value="ENSGAGP00000007428.1"/>
        <property type="gene ID" value="ENSGAGG00000005959.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015095">
        <property type="term" value="F:magnesium ion transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008521">
        <property type="entry name" value="Mg_trans_NIPA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12570">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12570:SF16">
        <property type="entry name" value="NIPA-LIKE PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05653">
        <property type="entry name" value="Mg_trans_NIPA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 6" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 6" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 6" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="42"/>
            <end position="62"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="82"/>
            <end position="102"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="109"/>
            <end position="132"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="152"/>
            <end position="173"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="185"/>
            <end position="209"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="221"/>
            <end position="239"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="251"/>
            <end position="272"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="278"/>
            <end position="300"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007230"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007428.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="338" mass="37572" checksum="8A07A0AC04E93669" modified="2019-05-08" version="1">MAGEMHVLTTLDHLPWNSTSAEQGARNGSQSHFWYNTNKTQLLGVLLAITGNFLISISLNIQKYAHLRLACQANQKPYYASKLWWCGIILMGLGELGNAAAYGFAPATLIAPLGCVSVIGGTLTIAGTYLLVTFAPNVTQEFTATKIQNYLISWQFLIYVILEIIIFCILLYFYERKDVKHIVVLLMMVALLASLTIISVKAVAGMITFSAKGKMQLTYPIFYIMFIIMVTSCVFQVKFLNQAMQLYNATVVVPINFVFFTTSAIIAGVIFYQEFQGAALLSVFMFLFGCLLSFLGVFVITRNREEEHLQIPYIDCGHIPGKHGFGCSLSNILPVIFV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ46</accession>
    <name>A0A452GZ46_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Gamma-tubulin complex component</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="6">Component of the gamma-tubulin ring complex (gTuRC) which mediates microtubule nucleation. The gTuRC regulates the minus-end nucleation of alpha-beta tubulin heterodimers that grow into microtubule protafilaments, a critical step in centrosome duplication and spindle formation.</text>
    </comment>
    <comment type="subunit">
        <text evidence="7">Component of the gamma-tubulin ring complex (gTuRC) consisting of TUBGCP2, TUBGCP3, TUBGCP4, TUBGCP5 and TUBGCP6 and gamma-tubulin TUBG1 or TUBG2. TUBGCP2, TUBGCP3, TUBGCP4, TUBGCP5 and TUBGCP6 assemble in a 5:5:2:1:1 stoichiometry; each is associated with a gamma-tubulin, thereby arranging 14 gamma-tubulins in a helical manner. Gamma-tubulin at the first position is blocked by TUBGCP3 at the last position, allowing 13 protafilaments to grow into a microtubule. The gTuRC (via TUBGCP3 and TUBGCP6) interacts with ACTB and MZT1; the interactions form a luminal bridge that stabilizes the initial structure during complex assembly. The gTuRC (via TUBGCP2) interacts with MZT2A/MZT2B and CDK5RAP2 (via CM1 motif); the interactions play a role in gTuRC activation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Microtubule organizing center</location>
            <location evidence="1">Centrosome</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 8">Belongs to the TUBGCP family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZ46"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008461.1">
        <property type="protein sequence ID" value="ENSGAGP00000007320.1"/>
        <property type="gene ID" value="ENSGAGG00000004413.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000930">
        <property type="term" value="C:gamma-tubulin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005874">
        <property type="term" value="C:microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000922">
        <property type="term" value="C:spindle pole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043015">
        <property type="term" value="F:gamma-tubulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051011">
        <property type="term" value="F:microtubule minus-end binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031122">
        <property type="term" value="P:cytoplasmic microtubule organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051321">
        <property type="term" value="P:meiotic cell cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007020">
        <property type="term" value="P:microtubule nucleation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000278">
        <property type="term" value="P:mitotic cell cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051225">
        <property type="term" value="P:spindle assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd22572">
        <property type="entry name" value="GCP5_NTD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.1900:FF:000005">
        <property type="entry name" value="Gamma-tubulin complex component"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.1900">
        <property type="entry name" value="Gamma-tubulin complex, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007259">
        <property type="entry name" value="GCP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR059169">
        <property type="entry name" value="GCP5_N_ext"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040457">
        <property type="entry name" value="GCP_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042241">
        <property type="entry name" value="GCP_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041470">
        <property type="entry name" value="GCP_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19302">
        <property type="entry name" value="GAMMA TUBULIN COMPLEX PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19302:SF33">
        <property type="entry name" value="GAMMA-TUBULIN COMPLEX COMPONENT 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04130">
        <property type="entry name" value="GCP_C_terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17681">
        <property type="entry name" value="GCP_N_terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 8" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="5 8" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="4 8" id="KW-0493">Microtubule</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Gamma tubulin complex component protein N-terminal" evidence="11">
        <location>
            <begin position="265"/>
            <end position="614"/>
        </location>
    </feature>
    <feature type="domain" description="Gamma tubulin complex component C-terminal" evidence="10">
        <location>
            <begin position="713"/>
            <end position="1014"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="151"/>
            <end position="196"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="522"/>
            <end position="541"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="9">
        <location>
            <begin position="152"/>
            <end position="166"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="182"/>
            <end position="191"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="528"/>
            <end position="540"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010337"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00093416"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00093551"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="RuleBase" id="RU363050"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="Pfam" id="PF04130"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF17681"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007320.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1021" mass="118279" checksum="0DF393F16A20B07C" modified="2019-05-08" version="1">MAGQPPRWSPLEQAQDRDVRALIRRLTGLREAQGGEEPGGRFQTALIFAWSNFRFHRFLDVSSHKVERTIEGIYEKLIVHSDLSKAASWKRLIEKFLNSPLPSTEETKVCNIFFFSFRPEISFVFLLEKEEKFDWGKYLMEGEEIYFGPDIDTPDWSEESEGEEDAQQPLSREDSGIQVDRTPQEEQDQNKKTVSQVTWKVGEPDARSWLEQHVVPQYWTGRAPRFSHSFHLHSNLAAVWDQHLYSSDPLYMPEERTLVTETQIIRETLWLLSGVKKLFIFQLNEGKVTVRNDIIVTHLTHNCLRSVLEQIAAYGQVVFRLQKFIDEVMGHSSESVIPGTISTPKKTTEPPFRTYQAFMWALYKYFISFKEELTEIEKCVINKDKTVTLSIVVDKLSPRLAQLKVLHKVFSTGIAEVPQVPPDTRNVVRASHLLNTLYKAILEYDNVGEASEQTVSLLFALWVETVRPYLQTVDEWIVHGNLFDPAKEFIIQRNKNVPVNHRDFWYATYTLYSVSEKMENEEKMSDNASASSGSDQAPSSRQHTMVSFLKPVLKQIIMAGKSMQLLKNLQSKDDSPRQAASRDAERKSLYTLFLESVQSRLRHGEESVPDIITEQQATKQSLIKMQSIAERHLELDDVHDPLLAINFARLYLEQSDFHEKFTGGDVCVDRSSESVTCQTFELTLRSCLYPHIDKQYLECCGNLMQTLKKDYRLVEYLQAMRNFFLLEAGDTMYDFYTSIFDKIREKETWQNVSFLNVQLQEAVGQRYPEDSSRLSISFESIDTAKKKLPVHTLDGLTLSYKVPWPVDIVISLECQKIYNQVFLLLLQIKWAKYSLDVLRFDELVNATEKPQLKEGPQLEQETVPQFGSQTELIKQQIHRMFLVRVKLMHFVNSLHNYIMTRILHSTGLEFQHQVEEAKDLDQLIKIHYRYLSTIHDRCLLREKVSFVKEAIMKVLNLVLIFADRWQAGLGAWKMESIEKMESDFKNCHMFLVTVLNKAVCRGSFPHLESLAVSLMAGMEQS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ49</accession>
    <name>A0A452GZ49_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">ATP synthase peripheral stalk subunit OSCP, mitochondrial</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="14">ATP synthase subunit O</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="11">Oligomycin sensitivity conferral protein</fullName>
        </alternativeName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="12">Component of the ATP synthase complex composed at least of ATP5F1A/subunit alpha, ATP5F1B/subunit beta, ATP5MC1/subunit c (homooctomer), MT-ATP6/subunit a, MT-ATP8/subunit 8, ATP5ME/subunit e, ATP5MF/subunit f, ATP5MG/subunit g, ATP5MK/subunit k, ATP5MJ/subunit j, ATP5F1C/subunit gamma, ATP5F1D/subunit delta, ATP5F1E/subunit epsilon, ATP5PF/subunit F6, ATP5PB/subunit b, ATP5PD/subunit d, ATP5PO/subunit OSCP. ATP synthase complex consists of a soluble F(1) head domain (subunits alpha(3) and beta(3)) - the catalytic core - and a membrane F(0) domain - the membrane proton channel (subunits c, a, 8, e, f, g, k and j). These two domains are linked by a central stalk (subunits gamma, delta, and epsilon) rotating inside the F1 region and a stationary peripheral stalk (subunits F6, b, d, and OSCP).</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Mitochondrion inner membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the ATPase delta chain family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZ49"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008587.1">
        <property type="protein sequence ID" value="ENSGAGP00000007433.1"/>
        <property type="gene ID" value="ENSGAGG00000005893.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005743">
        <property type="term" value="C:mitochondrial inner membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045259">
        <property type="term" value="C:proton-transporting ATP synthase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046933">
        <property type="term" value="F:proton-transporting ATP synthase activity, rotational mechanism"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.520.20:FF:000002">
        <property type="entry name" value="ATP synthase subunit O, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.520.20">
        <property type="entry name" value="N-terminal domain of the delta subunit of the F1F0-ATP synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026015">
        <property type="entry name" value="ATP_synth_OSCP/delta_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000711">
        <property type="entry name" value="ATPase_OSCP/dsu"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01145">
        <property type="entry name" value="ATP_synt_delta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11910">
        <property type="entry name" value="ATP SYNTHASE DELTA CHAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00213">
        <property type="entry name" value="OSCP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00125">
        <property type="entry name" value="ATPASEDELTA"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47928">
        <property type="entry name" value="N-terminal domain of the delta subunit of the F1F0-ATP synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0066">ATP synthesis</keyword>
    <keyword evidence="4" id="KW-0375">Hydrogen ion transport</keyword>
    <keyword evidence="7" id="KW-0406">Ion transport</keyword>
    <keyword evidence="9" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="5" id="KW-0999">Mitochondrion inner membrane</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0809">Transit peptide</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007046"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022781"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022792"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023310"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00033369"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00064647"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00073432"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00078525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007433.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="140" mass="15445" checksum="9C2F0CB0900B3D9B" modified="2019-05-08" version="1">MLQTVTNLTSFFFKPPIQVYGLEGRYATALYSAASKQKKLDQVEKELSRVSTLLKDPKLSSVVVNPHVKSVIKQKTINDALVKEKLSPITINFMKLLAENGRLQQTPDVISAFGKIMSAYRGEVLCSVTTANVNINTVVS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ72</accession>
    <name>A0A452GZ72_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Tyrosine-protein kinase receptor</fullName>
            <ecNumber evidence="16">2.7.10.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="14">
            <text>L-tyrosyl-[protein] + ATP = O-phospho-L-tyrosyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10596"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.10.1"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="16">Belongs to the protein kinase superfamily. Tyr protein kinase family. Insulin receptor subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.10.1" evidence="16"/>
    <dbReference type="AlphaFoldDB" id="A0A452GZ72"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008571.1">
        <property type="protein sequence ID" value="ENSGAGP00000007421.1"/>
        <property type="gene ID" value="ENSGAGG00000005944.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030424">
        <property type="term" value="C:axon"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005899">
        <property type="term" value="C:insulin receptor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005009">
        <property type="term" value="F:insulin receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043560">
        <property type="term" value="F:insulin receptor substrate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="CDD" id="cd05032">
        <property type="entry name" value="PTKc_InsR_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.220.10">
        <property type="entry name" value="Hormone Receptor, Insulin-like Growth Factor Receptor 1, Chain A, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.20.20">
        <property type="entry name" value="Receptor L-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000494">
        <property type="entry name" value="Rcpt_L-dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036941">
        <property type="entry name" value="Rcpt_L-dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050122">
        <property type="entry name" value="RTK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001245">
        <property type="entry name" value="Ser-Thr/Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008266">
        <property type="entry name" value="Tyr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020635">
        <property type="entry name" value="Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002011">
        <property type="entry name" value="Tyr_kinase_rcpt_2_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416:SF338">
        <property type="entry name" value="INSULIN RECEPTOR-RELATED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416">
        <property type="entry name" value="TYROSINE-PROTEIN KINASE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07714">
        <property type="entry name" value="PK_Tyr_Ser-Thr"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01030">
        <property type="entry name" value="Recep_L_domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00109">
        <property type="entry name" value="TYRKINASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00219">
        <property type="entry name" value="TyrKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52058">
        <property type="entry name" value="L domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00109">
        <property type="entry name" value="PROTEIN_KINASE_TYR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00239">
        <property type="entry name" value="RECEPTOR_TYR_KIN_II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 15" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7" id="KW-0418">Kinase</keyword>
    <keyword evidence="10" id="KW-0472">Membrane</keyword>
    <keyword evidence="6 15" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="2 16" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="12 16" id="KW-0675">Receptor</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0808">Transferase</keyword>
    <keyword evidence="4 16" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="11" id="KW-0829">Tyrosine-protein kinase</keyword>
    <feature type="domain" description="Fibronectin type-III" evidence="19">
        <location>
            <begin position="183"/>
            <end position="303"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="19">
        <location>
            <begin position="523"/>
            <end position="618"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="18">
        <location>
            <begin position="680"/>
            <end position="955"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="431"/>
            <end position="456"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="431"/>
            <end position="442"/>
        </location>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="714"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023137"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00051243"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="RuleBase" id="RU000312"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007421.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1029" mass="116092" checksum="238FCE285D6979C8" modified="2025-10-08" version="2">MSECPSGYTRNDSSIFCHKCEGLCPKECKVGTKTIDSTRAAQELAGCTYVEGNLIVNIRRGYNLASELQNSLGLIETITGFLKIKHSFALVSLSFFKNLKLIRGDSMVDGNYTLYVLDNQNLQQLWDWSHHTLSIPVGKVYFAFNPKLCLSEIYRMEEVTGTKGRQNKAEINPRTNGDRASCESRTLRFVSNITESDRIFLKWERYRPPEYRDLLSFIVYYKESTFQNVTEYVGQDACGANSWNVVDVELPLNNDQDPGVALLNLKPWTQYAIFVRAITLTTAEEGRNYGAQSEVVYVRTMPAAPTVPRDVLSMSNSSSHIIVRWKPPSQRNGNITYYLVLWQQLAEDLELYVNDYCHKGEAGGPGEGLLLPFVGGEDRREERCCPCHRDSGQLQMEGEAVSFQKKFENFLHNSIIIPRTRKRRRDILAMTSSGHPLSNASSLEPPGKPPSLNLTGAGPKAQTDFRIFEDKVVRDRTVISQLRHFTEYRIDIHACNHAAHTVGCSAATFVFARTMPELEADNIPGNVTWEPASKNSVLLRWEEPKNPNGLILKYEIKYSRENEVTTVVCVSRHRYAKYRGVHLALLQPGNYSAKVRATSLAGNGSWTGLIKFYILGPGMYLGWCSSISPHEALTFVLLFPPRRNNDGYPSGTLYASVNPEYFSTSDMYIPDEWEVSRDKITVIRELGQGSFGMVYEGIAQGIEKEDEETKVALKTVNELATMRERIEFLNEASVMKAFKCHHVVRLLGVVSQGQPALVIMELMTRGDLKSYLRSLRPDAENNPGLLPPSLKNMIQMAGEIADGMAYLNAKKFVHRDLAARNCMVSEDFTVKIGDFGMTRDIYETDYYRKGGKGLLPVRWMSPEALKDGIFNTHSDVWSFGVVLWEIATLAEQPYQGLSNEQVLRFVMDNGILKRPENCPEKLHELMSWCWQQNPRQRPSFTQLLESIKDDMDPTFRNISFFYSAENKRRGSCEPSDVETDNLLEEEPSTALAALRLPLATNGSPGPAVAPQKDRSCSRNCCHLNGAPYL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ73</accession>
    <name>A0A452GZ73_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="14">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="8">This is one of the enzymes that catalyzes the last step of the mitochondrial beta-oxidation pathway, an aerobic process breaking down fatty acids into acetyl-CoA. Using free coenzyme A/CoA, catalyzes the thiolytic cleavage of medium- to long-chain 3-oxoacyl-CoAs into acetyl-CoA and a fatty acyl-CoA shortened by two carbon atoms. The activity of the enzyme is reversible and it can also catalyze the condensation of two acetyl-CoA molecules into acetoacetyl-CoA. Thereby, it plays a major role in ketone body metabolism.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="9">
            <text>propanoyl-CoA + acetyl-CoA = 2-methyl-3-oxobutanoyl-CoA + CoA</text>
            <dbReference type="Rhea" id="RHEA:30719"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57288"/>
            <dbReference type="ChEBI" id="CHEBI:57335"/>
            <dbReference type="ChEBI" id="CHEBI:57392"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="9">
            <dbReference type="Rhea" id="RHEA:30720"/>
        </physiologicalReaction>
        <physiologicalReaction direction="right-to-left" evidence="9">
            <dbReference type="Rhea" id="RHEA:30721"/>
        </physiologicalReaction>
    </comment>
    <comment type="pathway">
        <text evidence="1">Lipid metabolism; fatty acid beta-oxidation.</text>
    </comment>
    <comment type="similarity">
        <text evidence="2 11">Belongs to the thiolase-like superfamily. Thiolase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZ73"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008617.1">
        <property type="protein sequence ID" value="ENSGAGP00000007461.1"/>
        <property type="gene ID" value="ENSGAGG00000005978.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003985">
        <property type="term" value="F:acetyl-CoA C-acetyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006635">
        <property type="term" value="P:fatty acid beta-oxidation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00751">
        <property type="entry name" value="thiolase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.47.10:FF:000007">
        <property type="entry name" value="acetyl-CoA acetyltransferase, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.47.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002155">
        <property type="entry name" value="Thiolase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016039">
        <property type="entry name" value="Thiolase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020615">
        <property type="entry name" value="Thiolase_acyl_enz_int_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020617">
        <property type="entry name" value="Thiolase_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020616">
        <property type="entry name" value="Thiolase_N"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01930">
        <property type="entry name" value="AcCoA-C-Actrans"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18919:SF156">
        <property type="entry name" value="ACETYL-COA ACETYLTRANSFERASE, MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18919">
        <property type="entry name" value="ACETYL-COA C-ACYLTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02803">
        <property type="entry name" value="Thiolase_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00108">
        <property type="entry name" value="Thiolase_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000429">
        <property type="entry name" value="Ac-CoA_Ac_transf"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53901">
        <property type="entry name" value="Thiolase-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00098">
        <property type="entry name" value="THIOLASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7 11" id="KW-0012">Acyltransferase</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="6" id="KW-0630">Potassium</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 11" id="KW-0808">Transferase</keyword>
    <keyword evidence="5" id="KW-0809">Transit peptide</keyword>
    <feature type="domain" description="Thiolase N-terminal" evidence="12">
        <location>
            <begin position="33"/>
            <end position="289"/>
        </location>
    </feature>
    <feature type="domain" description="Thiolase C-terminal" evidence="13">
        <location>
            <begin position="298"/>
            <end position="380"/>
        </location>
    </feature>
    <feature type="binding site" evidence="10">
        <location>
            <position position="210"/>
        </location>
        <ligand>
            <name>CoA</name>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="10">
        <location>
            <begin position="249"/>
            <end position="251"/>
        </location>
        <ligand>
            <name>CoA</name>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="10">
        <location>
            <position position="254"/>
        </location>
        <ligand>
            <name>CoA</name>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="10">
        <location>
            <position position="275"/>
        </location>
        <ligand>
            <name>CoA</name>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00005005"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010982"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022958"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023315"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00045244"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00048502"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PIRSR" id="PIRSR000429-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="RuleBase" id="RU003557"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF00108"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="Pfam" id="PF02803"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007461.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="407" mass="43572" checksum="603BC189BBAD7820" modified="2019-05-08" version="1">MAAMLGGREVARLLQVLKYTNRGYASQRTLNEVVIASAARTPIGSFQGSLSSLRATELGSIAIKGAIERAGIPAEEVKEVYMGNVLQAGQGQAPARQALLGAGLPISTPATTINKVCASGMKSVMMAAQNLMCGNQDVMVAGGMESMSNVPYTMNRGATPYGGVKLEDLIVKDGLTDVYNKIHMGNCAENTAKKLTVSREEQDTYAINSYTRSRTAWESGELTKEVVPVTISQKGKPDIEVKEDEEYKRVDFSKVPKLKTVFQKENGTVTAANASTLNDGAAALVVMTAEAAKRLNVTPLARIVAFADAAVDPIDFPIAPAYAVPKVLSDAGLKKEDIAMWEINEAFSVVVLANIKMLDIDPQKVNIHGGAVSLGHPIGEIAIYEFLEGFTFLFHYHHTGVLKTSTC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ76</accession>
    <name>A0A452GZ76_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">BTB domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GZ76"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008641.1">
        <property type="protein sequence ID" value="ENSGAGP00000007482.1"/>
        <property type="gene ID" value="ENSGAGG00000005975.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00143"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050801">
        <property type="term" value="P:monoatomic ion homeostasis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070936">
        <property type="term" value="P:protein K48-linked ubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070294">
        <property type="term" value="P:renal sodium ion absorption"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="entry name" value="Kelch-like family member 12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.710.10:FF:000001">
        <property type="entry name" value="Kelch-like family member 20"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.420">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.710.10">
        <property type="entry name" value="Potassium Channel Kv1.1, Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011705">
        <property type="entry name" value="BACK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000210">
        <property type="entry name" value="BTB/POZ_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011333">
        <property type="entry name" value="SKP1/BTB/POZ_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24412">
        <property type="entry name" value="KELCH PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24412:SF179">
        <property type="entry name" value="KELCH-LIKE PROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07707">
        <property type="entry name" value="BACK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00651">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00875">
        <property type="entry name" value="BACK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00225">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54695">
        <property type="entry name" value="POZ domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50097">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0880">Kelch repeat</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="BTB" evidence="4">
        <location>
            <begin position="51"/>
            <end position="118"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="1"/>
            <end position="29"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022441"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50097"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007482.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="304" mass="34631" checksum="B7FD9F94537C58C6" modified="2019-05-08" version="1">MNGEERKGLSSQSPDEAEDKEKDQRTVTVNPSHMGKAFKVMNELRSKRLLCDVVIVAEAIEMEAHRVVLAACSPYFCAMFTGDMTERKAKKIEIKDVDGQTLSKLIDYIYTAEIEVTEENVQVLLPAASLLQLMDVRQNCCDFLQSQLHPTNCLGIRAFADVHTCTELLQQANAYAEQHFPEVMLGEEFLNLSLDQVCSLISSDKLTVSSEEKVFEAVISWINYEKESRLEHMAELMEHVRLPLLPRDYLVQTVEEEALIKNNNTCKDFLIEAMKYHLLPLDQRQLIKNPRTKPRTPVSLPKVS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ78</accession>
    <name>A0A452GZ78_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">ATP-dependent DNA helicase</fullName>
            <ecNumber evidence="17">5.6.2.4</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="23">
            <text>ATP + H2O = ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:13065"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="16 17">
            <text>Couples ATP hydrolysis with the unwinding of duplex DNA by translocating in the 3'-5' direction.</text>
            <dbReference type="EC" id="5.6.2.4"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2 17">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 17">Belongs to the helicase family. RecQ subfamily.</text>
    </comment>
    <dbReference type="EC" id="5.6.2.4" evidence="17"/>
    <dbReference type="AlphaFoldDB" id="A0A452GZ78"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008505.1">
        <property type="protein sequence ID" value="ENSGAGP00000007360.1"/>
        <property type="gene ID" value="ENSGAGG00000005383.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005694">
        <property type="term" value="C:chromosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043138">
        <property type="term" value="F:3'-5' DNA helicase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="RHEA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009378">
        <property type="term" value="F:four-way junction helicase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006260">
        <property type="term" value="P:DNA replication"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000724">
        <property type="term" value="P:double-strand break repair via homologous recombination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000723">
        <property type="term" value="P:telomere maintenance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd18016">
        <property type="entry name" value="DEXHc_RecQ2_BLM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18794">
        <property type="entry name" value="SF2_C_RecQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000537">
        <property type="entry name" value="Bloom syndrome RecQ-like helicase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000340">
        <property type="entry name" value="Bloom syndrome, RecQ helicase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.80">
        <property type="entry name" value="HRDC domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.10">
        <property type="entry name" value="Winged helix-like DNA-binding domain superfamily/Winged helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032437">
        <property type="entry name" value="BLM_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011545">
        <property type="entry name" value="DEAD/DEAH_box_helicase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002464">
        <property type="entry name" value="DNA/RNA_helicase_DEAH_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004589">
        <property type="entry name" value="DNA_helicase_ATP-dep_RecQ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014001">
        <property type="entry name" value="Helicase_ATP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001650">
        <property type="entry name" value="Helicase_C-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044876">
        <property type="entry name" value="HRDC_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032284">
        <property type="entry name" value="RecQ_Zn-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018982">
        <property type="entry name" value="RQC_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036388">
        <property type="entry name" value="WH-like_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036390">
        <property type="entry name" value="WH_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00614">
        <property type="entry name" value="recQ_fam"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13710">
        <property type="entry name" value="DNA HELICASE RECQ FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13710:SF153">
        <property type="entry name" value="RECQ-LIKE DNA HELICASE BLM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16202">
        <property type="entry name" value="BLM_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00270">
        <property type="entry name" value="DEAD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00271">
        <property type="entry name" value="Helicase_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16124">
        <property type="entry name" value="RecQ_Zn_bind"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09382">
        <property type="entry name" value="RQC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00487">
        <property type="entry name" value="DEXDc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00490">
        <property type="entry name" value="HELICc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00956">
        <property type="entry name" value="RQC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46785">
        <property type="entry name" value="Winged helix' DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00690">
        <property type="entry name" value="DEAH_ATP_HELICASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51192">
        <property type="entry name" value="HELICASE_ATP_BIND_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51194">
        <property type="entry name" value="HELICASE_CTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="11 17" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="7" id="KW-0227">DNA damage</keyword>
    <keyword evidence="13" id="KW-0234">DNA repair</keyword>
    <keyword evidence="4" id="KW-0235">DNA replication</keyword>
    <keyword evidence="12" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="9 17" id="KW-0347">Helicase</keyword>
    <keyword evidence="8 17" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="14" id="KW-0413">Isomerase</keyword>
    <keyword evidence="5" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="6 17" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="15 17" id="KW-0539">Nucleus</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="Helicase ATP-binding" evidence="19">
        <location>
            <begin position="369"/>
            <end position="544"/>
        </location>
    </feature>
    <feature type="domain" description="Helicase C-terminal" evidence="20">
        <location>
            <begin position="570"/>
            <end position="717"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="117"/>
            <end position="144"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="206"/>
            <end position="228"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="265"/>
            <end position="284"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="206"/>
            <end position="215"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="218"/>
            <end position="228"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="265"/>
            <end position="276"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005446"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022705"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022806"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023235"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00034617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="RuleBase" id="RU364117"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS51192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS51194"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007360.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="963" mass="108759" checksum="6B91B3DA183BCA06" modified="2019-05-08" version="1">MAALPQNNLQKQLELHSAKGTHNKLALPKHKPGGFTFKKKSAPGNAVSINCFSVTKSSALKEKDVNTSQTKSVASCPTSKDKQIKINDLFEKTSKQHELEQLHPNWVSLNSFECSQQVSNDLPPRKEENVSKKPGNSASQKVDHNCSDDAIITIEDDWDDIDDFDISGKQKNSKPLVLSPKNLIVKPSKTLQISSSCTVRLSKASRTVNSEQQLSKHCPSENDEKSLNDKQPALANRSLICLDVSVPNDCKRLLGEDLQEKKLSTETVLSDKKEETPTGNASSMVAFSEMPPVPCAGPGRQAELQSLNAWIRRWCFPFLLDPLLKKSPSKNPALARFRGCTFSHSNDMMRIFHKKFGLHHFRTNQLEAINASLLGEDCFILMPTGGGKSLCYQLPACLSSGVTIVISPLRSLILDQVEKLMTLDISATYLTGDRTDTEASKIYIQLSKKDPQIKLLYVTPEKVCSSGRLISALKNLYERDLLARFVIDEAHCVSQWGHDFRQDYKRLNMLRQKFPSVPMMALTATANPRVQKDILNQLEMLKPQVFTMSFNRHNLKYDVLPKKPKRVALDCLEWIKKYHPHDSGIIYCLSRQDCNKTADILQEKGLAALAYHAGLSDTVRDLVQRKWINQDGCQVICATIAFGMGIDKPDVRYVIHASLPKSVEGYYQESGRAGRDGEVSHCLLFYSYSDVTRLRRLIQMEKDGNSQTRKTHFDNLYSMVHYCENVHECRRIQLLAYFGETGFNLNFCKEHPEVICDNCCRKKDYKSRNVTDEVKSIIRFVQEHCGQMGRRNSGSGRYTLNMMIDIFLGTKSARIQSGIFGKGAAYSRHNVERLFRKLVLDKILEEELYITAAEQAVAYVTVGEKAQAVLNGYLQVEFYETENASSIRKQRASVTKMSQREEMVKKCLVELTDVCKTLGKVFDVHYFNIFNTATLKRIAVGYFWFRAGFIPGWAYTGGGGRFE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ81</accession>
    <name>A0A452GZ81_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="19">Phospholipid-transporting ATPase</fullName>
            <ecNumber evidence="19">7.6.2.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="15 19">
            <text>ATP + H2O + phospholipidSide 1 = ADP + phosphate + phospholipidSide 2.</text>
            <dbReference type="EC" id="7.6.2.1"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 18">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Golgi apparatus</location>
            <location evidence="2">trans-Golgi network membrane</location>
            <topology evidence="2">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="19">Membrane</location>
            <topology evidence="19">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 19">Belongs to the cation transport ATPase (P-type) (TC 3.A.3) family. Type IV subfamily.</text>
    </comment>
    <dbReference type="EC" id="7.6.2.1" evidence="19"/>
    <dbReference type="AlphaFoldDB" id="A0A452GZ81"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008649.1">
        <property type="protein sequence ID" value="ENSGAGP00000007487.1"/>
        <property type="gene ID" value="ENSGAGG00000000721.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005768">
        <property type="term" value="C:endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005802">
        <property type="term" value="C:trans-Golgi network"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140326">
        <property type="term" value="F:ATPase-coupled intramembrane lipid transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000287">
        <property type="term" value="F:magnesium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006897">
        <property type="term" value="P:endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045332">
        <property type="term" value="P:phospholipid translocation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006890">
        <property type="term" value="P:retrograde vesicle-mediated transport, Golgi to endoplasmic reticulum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.1110.10:FF:000008">
        <property type="entry name" value="Phospholipid-transporting ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1000:FF:000009">
        <property type="entry name" value="Phospholipid-transporting ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.1110.10">
        <property type="entry name" value="Calcium-transporting ATPase, cytoplasmic domain N"/>
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        <property type="entry name" value="Calcium-transporting ATPase, cytoplasmic transduction domain A"/>
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        <property type="entry name" value="Calcium-transporting ATPase, transmembrane domain"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="HAD superfamily/HAD-like"/>
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    <dbReference type="InterPro" id="IPR023299">
        <property type="entry name" value="ATPase_P-typ_cyto_dom_N"/>
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        <property type="entry name" value="ATPase_P-typ_P_site"/>
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    <dbReference type="InterPro" id="IPR036412">
        <property type="entry name" value="HAD-like_sf"/>
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        <property type="entry name" value="HAD_sf"/>
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    <dbReference type="InterPro" id="IPR006539">
        <property type="entry name" value="P-type_ATPase_IV"/>
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        <property type="entry name" value="P_typ_ATPase_c"/>
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    <dbReference type="InterPro" id="IPR044492">
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        <property type="entry name" value="Hydrolase"/>
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        <property type="entry name" value="PhoLip_ATPase_C"/>
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    <dbReference type="PRINTS" id="PR00119">
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        <property type="entry name" value="Haloacid_Dehalogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
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    </dbReference>
    <dbReference type="PROSITE" id="PS00154">
        <property type="entry name" value="ATPASE_E1_E2"/>
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    </dbReference>
    <proteinExistence type="inferred from homology"/>
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    <keyword evidence="12" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="13" id="KW-0445">Lipid transport</keyword>
    <keyword evidence="9 18 19" id="KW-0460">Magnesium</keyword>
    <keyword evidence="14 19" id="KW-0472">Membrane</keyword>
    <keyword evidence="6 18" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7 17 19" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10 19" id="KW-1278">Translocase</keyword>
    <keyword evidence="5 19" id="KW-0812">Transmembrane</keyword>
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    <keyword evidence="4" id="KW-0813">Transport</keyword>
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        <location>
            <begin position="12"/>
            <end position="33"/>
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    <feature type="transmembrane region" description="Helical" evidence="19">
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            <begin position="91"/>
            <end position="111"/>
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    <feature type="transmembrane region" description="Helical" evidence="19">
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            <end position="136"/>
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            <end position="649"/>
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            <end position="675"/>
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            <end position="725"/>
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            <begin position="731"/>
            <end position="752"/>
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            <end position="779"/>
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        <location>
            <begin position="785"/>
            <end position="811"/>
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            <begin position="593"/>
            <end position="820"/>
        </location>
    </feature>
    <feature type="active site" description="4-aspartylphosphate intermediate" evidence="16">
        <location>
            <position position="176"/>
        </location>
    </feature>
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        <location>
            <position position="176"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
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        <location>
            <position position="176"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
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    <sequence length="828" mass="93167" checksum="4FCC029A53E38DFF" modified="2019-05-08" version="1">IGDFFSFSAALLWVPWLVYRIVITCRFLILLSYSSSTHLVKIHNQPFLFSPGTVIGVVIYTGKETRSVMNTSNPKNKVGLLDLELNQLTKALFLALVALSIVMVTLQGFVGPWYRNLFRFLLLFSYIIPISLRVNLDMGKAAYGWMIMKDDNIPGTVVRTSTIPEELGRLVYLLTDKTGTLTQNEMIFKRLHLGTVSYGTDTMDEIQNHIISSYSQQAPFQSPKVRKTVSSRIHEAVKAIALCHNVTPVYESHAGVNGETEYAEVDQNFSDENRTYQASSPDEVALVQWTESVGLTLVNRDLTSMQLKSPGGQILTYYILQVFPFTSESKRMGIIVRDESTGEITFYMKGADVAMSTIVQYNDWLEEECGNMAREGLRTLVVAKKSLTEEQHQDFENRYNQAKLSIHDRALKVAAVVESLEREMELLCLTGVEDQLQADVRPTLEMLRNAGIKIWMLTGDKLETATCIAKSSHLVSRTQDIHIFRPVISRGEAHLELNAFRRKHDCALVISGDSLEVCLKYYEHEFVELACQCPAVVCCRCSPTQKAHIVKLLQHHTGKRTCAIGDGGNDVSMIQAADCGIGIEGKEGKQASLAADFSITQFKHIGRLLMVHGRNSYKRSAALGQFVMHRGLIISTMQAVFSSVFYFASIPLYQGFLMVGYATIYTMFPVFSLVLDQDVKPEMALLYPELYKDLTKGRSLSFKTFLIWVLISIYQGGILMYGALLLFESEFVHVVAISFTALILTELLMVALTIRTWHWLMVVAEFFSLGCYVASLAFLNEYFDVAFITTVTFLWKVSAITVVSCLPLYILKYLKRKFSPPSYSKLTS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ83</accession>
    <name>A0A452GZ83_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="7">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the TMEM45 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZ83"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008627.1">
        <property type="protein sequence ID" value="ENSGAGP00000007471.1"/>
        <property type="gene ID" value="ENSGAGG00000005952.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006904">
        <property type="entry name" value="DUF716"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042127">
        <property type="entry name" value="TMEM45"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16007">
        <property type="entry name" value="EPIDIDYMAL MEMBRANE PROTEIN E9-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16007:SF21">
        <property type="entry name" value="TRANSMEMBRANE PROTEIN 45A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04819">
        <property type="entry name" value="DUF716"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 6" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 6" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 6" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="12"/>
            <end position="31"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="59"/>
            <end position="79"/>
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            <begin position="99"/>
            <end position="120"/>
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            <begin position="132"/>
            <end position="150"/>
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            <begin position="162"/>
            <end position="181"/>
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            <begin position="193"/>
            <end position="211"/>
        </location>
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        <location>
            <begin position="226"/>
            <end position="250"/>
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        <source>
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        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
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    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
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    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Phobius"/>
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    </evidence>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007471.1"/>
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        </source>
    </evidence>
    <sequence length="259" mass="30003" checksum="E28B5CC861F7A126" modified="2019-05-08" version="1">ELGFLKQDMGNFKGHALPGSFFLLFGLWWSVKYPLKYVCRKNKNACYLGSRAGFQRLEFIEGIIKAVFALIGMIAEQFVPDGPHLKLYNYEEKHWDHLMNWQHATMYLFYGISGLVDVVAHGTNALPVALDRMMLSLAVFIEGFLFYYHIHGRDMLDFHVHQLLLVAIFGAALCILLEVFFRGSIVLEMLRTSLCILQGSWLWQIGFVLYPPNGRLEWNQMDHNNMMFLTMCYCWHYAFAFLILAVNYTIVSCLEQLSS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ87</accession>
    <name>A0A452GZ87_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="19">Phospholipid-transporting ATPase</fullName>
            <ecNumber evidence="19">7.6.2.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="15 19">
            <text>ATP + H2O + phospholipidSide 1 = ADP + phosphate + phospholipidSide 2.</text>
            <dbReference type="EC" id="7.6.2.1"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 18">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Golgi apparatus</location>
            <location evidence="2">trans-Golgi network membrane</location>
            <topology evidence="2">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="19">Membrane</location>
            <topology evidence="19">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 19">Belongs to the cation transport ATPase (P-type) (TC 3.A.3) family. Type IV subfamily.</text>
    </comment>
    <dbReference type="EC" id="7.6.2.1" evidence="19"/>
    <dbReference type="AlphaFoldDB" id="A0A452GZ87"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008590.1">
        <property type="protein sequence ID" value="ENSGAGP00000007436.1"/>
        <property type="gene ID" value="ENSGAGG00000000721.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005768">
        <property type="term" value="C:endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005802">
        <property type="term" value="C:trans-Golgi network"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140326">
        <property type="term" value="F:ATPase-coupled intramembrane lipid transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000287">
        <property type="term" value="F:magnesium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006897">
        <property type="term" value="P:endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045332">
        <property type="term" value="P:phospholipid translocation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006890">
        <property type="term" value="P:retrograde vesicle-mediated transport, Golgi to endoplasmic reticulum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.1110.10:FF:000008">
        <property type="entry name" value="Phospholipid-transporting ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1000:FF:000009">
        <property type="entry name" value="Phospholipid-transporting ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.1110.10">
        <property type="entry name" value="Calcium-transporting ATPase, cytoplasmic domain N"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.70.150.10">
        <property type="entry name" value="Calcium-transporting ATPase, cytoplasmic transduction domain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1110.10">
        <property type="entry name" value="Calcium-transporting ATPase, transmembrane domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1000">
        <property type="entry name" value="HAD superfamily/HAD-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023299">
        <property type="entry name" value="ATPase_P-typ_cyto_dom_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018303">
        <property type="entry name" value="ATPase_P-typ_P_site"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023298">
        <property type="entry name" value="ATPase_P-typ_TM_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008250">
        <property type="entry name" value="ATPase_P-typ_transduc_dom_A_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036412">
        <property type="entry name" value="HAD-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023214">
        <property type="entry name" value="HAD_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006539">
        <property type="entry name" value="P-type_ATPase_IV"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001757">
        <property type="entry name" value="P_typ_ATPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032630">
        <property type="entry name" value="P_typ_ATPase_c"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044492">
        <property type="entry name" value="P_typ_ATPase_HD_dom"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01652">
        <property type="entry name" value="ATPase-Plipid"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01494">
        <property type="entry name" value="ATPase_P-type"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24092:SF50">
        <property type="entry name" value="PHOSPHOLIPID-TRANSPORTING ATPASE IIB-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24092">
        <property type="entry name" value="PROBABLE PHOSPHOLIPID-TRANSPORTING ATPASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00702">
        <property type="entry name" value="Hydrolase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16212">
        <property type="entry name" value="PhoLip_ATPase_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00119">
        <property type="entry name" value="CATATPASE"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDG00002">
        <property type="entry name" value="C1.7:_P-type_atpase_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDS00003">
        <property type="entry name" value="Haloacid_Dehalogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDF00027">
        <property type="entry name" value="p-type_atpase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81653">
        <property type="entry name" value="Calcium ATPase, transduction domain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81665">
        <property type="entry name" value="Calcium ATPase, transmembrane domain M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56784">
        <property type="entry name" value="HAD-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81660">
        <property type="entry name" value="Metal cation-transporting ATPase, ATP-binding domain N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00154">
        <property type="entry name" value="ATPASE_E1_E2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 17 19" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="12" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="13" id="KW-0445">Lipid transport</keyword>
    <keyword evidence="9 18 19" id="KW-0460">Magnesium</keyword>
    <keyword evidence="14 19" id="KW-0472">Membrane</keyword>
    <keyword evidence="6 18" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7 17 19" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10 19" id="KW-1278">Translocase</keyword>
    <keyword evidence="5 19" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 19" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="4" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="68"/>
            <end position="87"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="116"/>
            <end position="136"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="696"/>
            <end position="716"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="746"/>
            <end position="766"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="772"/>
            <end position="793"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="800"/>
            <end position="819"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="839"/>
            <end position="863"/>
        </location>
    </feature>
    <feature type="domain" description="P-type ATPase C-terminal" evidence="21">
        <location>
            <begin position="634"/>
            <end position="872"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="246"/>
            <end position="268"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="20">
        <location>
            <begin position="246"/>
            <end position="256"/>
        </location>
    </feature>
    <feature type="active site" description="4-aspartylphosphate intermediate" evidence="16">
        <location>
            <position position="201"/>
        </location>
    </feature>
    <feature type="binding site" evidence="17">
        <location>
            <position position="201"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="201"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="17">
        <location>
            <position position="202"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="17">
        <location>
            <position position="203"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="203"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="17">
        <location>
            <position position="324"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="17">
        <location>
            <position position="366"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="17">
        <location>
            <position position="390"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="17">
        <location>
            <position position="419"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="17">
        <location>
            <position position="499"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="17">
        <location>
            <position position="500"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="17">
        <location>
            <position position="501"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="17">
        <location>
            <position position="581"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="17">
        <location>
            <position position="587"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="607"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="17">
        <location>
            <position position="610"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="17">
        <location>
            <position position="611"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18">
        <location>
            <position position="611"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004166"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00008109"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022967"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00034036"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PIRSR" id="PIRSR606539-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="PIRSR" id="PIRSR606539-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="PIRSR" id="PIRSR606539-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="RuleBase" id="RU362033"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="Pfam" id="PF16212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007436.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="880" mass="98363" checksum="08A43FBA437B9541" modified="2019-05-08" version="1">FLDGETDWKLKVAVSCTQRLPALGDLFSISAYVYAQKPQLDIHSFEGTFTREDSDPTVHESLSIENTLWASTVVASGTVIGVVIYTGKETRSVMNTSNPKNKVGLLDLELNQLTKALFLALVALSIVMVTLQGFVGPWYRNLFRFLLLFSYIIPISLRVNLDMGKAAYGWMIMKDDNIPGTVVRTSTIPEELGRLVYLLTDKTGTLTQNEMIFKRLHLGTVSYGTDTMDEIQNHIISSYSQMYSQSSGNSTSSTPSRKVQSSAPKVRKTVSSRIHEAVKAIALCHNVTPVYESHAGVNGETEYAEVDQNFSDENRTYQASSPDEVALVQWTESVGLTLVNRDLTSMQLKSPGGQILTYYILQVFPFTSESKRMGIIVRDESTGEITFYMKGADVAMSTIVQYNDWLEEECGNMAREGLRTLVVAKKSLTEEQHQDFENRYNQAKLSIHDRALKVAAVVESLEREMELLCLTGVEDQLQADVRPTLEMLRNAGIKIWMLTGDKLETATCIAKSSHLVSRTQDIHIFRPVISRGEAHLELNAFRRKHDCALVISGDSLEVCLKYYEHEFVELACQCPAVVCCRCSPTQKAHIVKLLQHHTGKRTCAIGDGGNDVSMIQAADCGIGIEGKEGKQASLAADFSITQFKHIGRLLMVHGRNSYKRSAALGQFVMHRGLIISTMQAVFSSVFYFASIPLYQGFLMVGYATIYTMFPVFSLVLDQDVKPEMALLYPELYKDLTKGRSLSFKTFLIWVLISIYQGGILMYGALLLFESEFVHVVAISFTALILTELLMVALTIRTWHWLMVVAEFFSLGCYVASLAFLNEYFGIGRVSFGAFLDVAFITTVTFLWKVSAITVVSCLPLYILKYLKRKFSPPSYSKLTS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ88</accession>
    <name>A0A452GZ88_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Peptidase M14 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 13">Belongs to the peptidase M14 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZ88"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008634.1">
        <property type="protein sequence ID" value="ENSGAGP00000007476.1"/>
        <property type="gene ID" value="ENSGAGG00000005982.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004181">
        <property type="term" value="F:metallocarboxypeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd03870">
        <property type="entry name" value="M14_CPA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.630.10:FF:000132">
        <property type="entry name" value="Carboxypeptidase A1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.340:FF:000001">
        <property type="entry name" value="Carboxypeptidase A5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.340">
        <property type="entry name" value="Metallocarboxypeptidase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.630.10">
        <property type="entry name" value="Zn peptidases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057246">
        <property type="entry name" value="CARBOXYPEPT_ZN_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057247">
        <property type="entry name" value="CARBOXYPEPT_ZN_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034248">
        <property type="entry name" value="CPA_M14_CPD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036990">
        <property type="entry name" value="M14A-like_propep"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003146">
        <property type="entry name" value="M14A_act_pep"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000834">
        <property type="entry name" value="Peptidase_M14"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11705:SF71">
        <property type="entry name" value="CARBOXYPEPTIDASE A2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11705">
        <property type="entry name" value="PROTEASE FAMILY M14 CARBOXYPEPTIDASE A,B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00246">
        <property type="entry name" value="Peptidase_M14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02244">
        <property type="entry name" value="Propep_M14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00765">
        <property type="entry name" value="CRBOXYPTASEA"/>
    </dbReference>
    <dbReference type="SMART" id="SM00631">
        <property type="entry name" value="Zn_pept"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54897">
        <property type="entry name" value="Protease propeptides/inhibitors"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53187">
        <property type="entry name" value="Zn-dependent exopeptidases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00132">
        <property type="entry name" value="CARBOXYPEPT_ZN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00133">
        <property type="entry name" value="CARBOXYPEPT_ZN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS52035">
        <property type="entry name" value="PEPTIDASE_M14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0121">Carboxypeptidase</keyword>
    <keyword evidence="12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="9" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-0482">Metalloprotease</keyword>
    <keyword evidence="6" id="KW-0645">Protease</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="8 14" id="KW-0732">Signal</keyword>
    <keyword evidence="10" id="KW-0862">Zinc</keyword>
    <feature type="signal peptide" evidence="14">
        <location>
            <begin position="1"/>
            <end position="18"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019475861" description="Peptidase M14 domain-containing protein" evidence="14">
        <location>
            <begin position="19"/>
            <end position="390"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase M14" evidence="15">
        <location>
            <begin position="92"/>
            <end position="385"/>
        </location>
    </feature>
    <feature type="active site" description="Proton donor/acceptor" evidence="13">
        <location>
            <position position="351"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005988"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022645"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023049"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01379"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS52035"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007476.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="390" mass="43670" checksum="49BDB8ED0D67A035" modified="2019-05-08" version="1">SNMRLILFFSALAGASWCLETFLDFWLHPSNPALPVDIRVPFPSLQSVKVFLESHHIEYSILLEDLQVDKKKQEMVNNQRRERDTSNFNYGAYHPLESIYAALDGLAAEYPSLVSKQEIGKSYEKRPMYILKFSTGGTNRPAIWLDAGIHSREWITQATALWTAQKIASEYGNNPSITSLLNRMDIFLLVVTNPDGYAFTHTNNRMWRKTRSKNSGSLCIGVDPNRNWDAGFGGPGASTNPCSESYHGPSANSEVEVKNVVNFIKSHGKIKAFLTLHSYSQLLLFPFGYKCTKPADYNELDTVGKAAASALASLYGTTYKVGTICSTIYQASGGSIDWSYDYGIKYSFAFELRDTGRYGFLLPASQIIPTAEETWLGLLKIMEHVRDNQS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ92</accession>
    <name>A0A452GZ92_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Cytochrome P450 family 2 subfamily W member 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="1 13">
            <name>heme</name>
            <dbReference type="ChEBI" id="CHEBI:30413"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Endoplasmic reticulum membrane</location>
            <topology evidence="3">Peripheral membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Microsome membrane</location>
            <topology evidence="2">Peripheral membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4 14">Belongs to the cytochrome P450 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZ92"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008596.1">
        <property type="protein sequence ID" value="ENSGAGP00000007441.1"/>
        <property type="gene ID" value="ENSGAGG00000005769.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0020037">
        <property type="term" value="F:heme binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005506">
        <property type="term" value="F:iron ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016712">
        <property type="term" value="F:oxidoreductase activity, acting on paired donors, with incorporation or reduction of molecular oxygen, reduced flavin or flavoprotein as one donor, and incorporation of one atom of oxygen"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006082">
        <property type="term" value="P:organic acid metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006805">
        <property type="term" value="P:xenobiotic metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.630.10:FF:000238">
        <property type="entry name" value="Cytochrome P450 2A6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.630.10">
        <property type="entry name" value="Cytochrome P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001128">
        <property type="entry name" value="Cyt_P450"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017972">
        <property type="entry name" value="Cyt_P450_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002401">
        <property type="entry name" value="Cyt_P450_E_grp-I"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036396">
        <property type="entry name" value="Cyt_P450_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050182">
        <property type="entry name" value="Cytochrome_P450_fam2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24300:SF302">
        <property type="entry name" value="CYTOCHROME P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24300">
        <property type="entry name" value="CYTOCHROME P450 508A4-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00067">
        <property type="entry name" value="p450"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00463">
        <property type="entry name" value="EP450I"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00385">
        <property type="entry name" value="P450"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48264">
        <property type="entry name" value="Cytochrome P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00086">
        <property type="entry name" value="CYTOCHROME_P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="5 13 14" id="KW-0349">Heme</keyword>
    <keyword evidence="10 13 14" id="KW-0408">Iron</keyword>
    <keyword evidence="12 15" id="KW-0472">Membrane</keyword>
    <keyword evidence="6 13 14" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8" id="KW-0492">Microsome</keyword>
    <keyword evidence="11 14" id="KW-0503">Monooxygenase</keyword>
    <keyword evidence="9 14" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="15" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="15" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="6"/>
            <end position="26"/>
        </location>
    </feature>
    <feature type="binding site" description="axial binding residue" evidence="13">
        <location>
            <position position="369"/>
        </location>
        <ligand>
            <name>heme</name>
            <dbReference type="ChEBI" id="CHEBI:30413"/>
        </ligand>
        <ligandPart>
            <name>Fe</name>
            <dbReference type="ChEBI" id="CHEBI:18248"/>
        </ligandPart>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001971"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004174"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004406"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00010617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022848"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023033"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PIRSR" id="PIRSR602401-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="RuleBase" id="RU000461"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007441.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="18">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="426" mass="48525" checksum="713D6F9C6FF1BAFF" modified="2019-05-08" version="1">SSKMALADPTVILLLFVVTSVSILFVKKSWNKRTCLNFPPGPRPLPILGNLHQLNLKRPYRTLLKKYGPVFSIHMGPKNMVVLSGYKMVKEALVNQADAFAERPRIPIFEKFARGNGSSFDTMKMMNAAVSNVIVSIMLGHRFDYEDPKLLRLIHLINESVRLSGSPMVTNLEELYSFTQQTFIEHLKQLDPNDQRSFIDAFLIRQQEEKSNPNTHFKNENLQSLVSNLFTAGMETTSTTLRWGLLLMMKYPAIQKKVHEEIERVIGSAQLRYEHRTQMPYTDAVIHEIQRFANILPMSLPHETTEDVRLNGYVIPKGTYIIPLLASVLYDKTQWEKPGEFCPQHFLSTDGKFVTNEAFMPFSAGRRICAGETLARMELFLFFTSLLQRFTFHPPPGVTSSDLDLTPAVGFTTPPMPHKICAMPCR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKT2</accession>
    <name>A0A452HKT2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Mortality factor 4-like protein 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKT2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017743.1">
        <property type="protein sequence ID" value="ENSGAGP00000015531.1"/>
        <property type="gene ID" value="ENSGAGG00000011716.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035267">
        <property type="term" value="C:NuA4 histone acetyltransferase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006325">
        <property type="term" value="P:chromatin organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006310">
        <property type="term" value="P:DNA recombination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006281">
        <property type="term" value="P:DNA repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006355">
        <property type="term" value="P:regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd18983">
        <property type="entry name" value="CBD_MSL3_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.274.30:FF:000001">
        <property type="entry name" value="Mortality factor 4-like protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.140:FF:000024">
        <property type="entry name" value="Mortality factor 4-like protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.274.30">
        <property type="entry name" value="MRG domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016197">
        <property type="entry name" value="Chromo-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000953">
        <property type="entry name" value="Chromo/chromo_shadow_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008676">
        <property type="entry name" value="MRG"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038217">
        <property type="entry name" value="MRG_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026541">
        <property type="entry name" value="MRG_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025995">
        <property type="entry name" value="Tudor-knot"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10880:SF48">
        <property type="entry name" value="MORTALITY FACTOR 4 LIKE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10880">
        <property type="entry name" value="MORTALITY FACTOR 4-LIKE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05712">
        <property type="entry name" value="MRG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11717">
        <property type="entry name" value="Tudor-knot"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF038133">
        <property type="entry name" value="HAT_Nua4_EAF3/MRG15"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00298">
        <property type="entry name" value="CHROMO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54160">
        <property type="entry name" value="Chromo domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51640">
        <property type="entry name" value="MRG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0007">Acetylation</keyword>
    <keyword evidence="4" id="KW-0156">Chromatin regulator</keyword>
    <keyword evidence="3" id="KW-0227">DNA damage</keyword>
    <keyword evidence="8" id="KW-0233">DNA recombination</keyword>
    <keyword evidence="9" id="KW-0234">DNA repair</keyword>
    <keyword evidence="2" id="KW-0341">Growth regulation</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0804">Transcription</keyword>
    <keyword evidence="6" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="Chromo" evidence="13">
        <location>
            <begin position="58"/>
            <end position="111"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1"/>
            <end position="34"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="110"/>
            <end position="132"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="162"/>
            <end position="199"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="12">
        <location>
            <begin position="9"/>
            <end position="18"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022604"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023172"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00071326"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="SMART" id="SM00298"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015531.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="377" mass="43419" checksum="2ABFAC5F20F8B20E" modified="2019-05-08" version="1">MRRSWRGPAGRGGVGGGRVTYKWRPSRTPSPNSRRDCQDFLFLRLRSERVLCFHGPLLYEAKCVKVAIKDKQVKYFIHYSGWNKNWDEWVPESRVLKYVDTNLQKQKELQKANQEQYAEGKMRGAAPGKKTSGLQQKNVEVFFRRDGAHTVCCLETPTISTRKTKKNKQKTPGIGEGSSTSETPQPPRKKRARVDPTVESEETFMNRVEVKVKIPEELKPWLVDDWDLITRQKQLFYLPAKKNVDSILEDYASYKKSRGNTDNKEYAVNEVVAGIKEYFNVMLGTQLLYKFERPQYAEILADHPDAPMSQVYGAPHLLRLFVRIGAMLAYTPLDEKSLALLLNYLHDFLKYLAKNASTLFSASDYEVAPPEYHRKAV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKT4</accession>
    <name>A0A452HKT4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">Neural cell adhesion molecule L1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">Neural cell adhesion molecule involved in the dynamics of cell adhesion and in the generation of transmembrane signals at tyrosine kinase receptors. During brain development, critical in multiple processes, including neuronal migration, axonal growth and fasciculation, and synaptogenesis. In the mature brain, plays a role in the dynamics of neuronal structure and function, including synaptic plasticity.</text>
    </comment>
    <comment type="subunit">
        <text evidence="16">Interacts with SHTN1; the interaction occurs in axonal growth cones. Interacts with isoform 2 of BSG.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cell projection</location>
            <location evidence="2">Growth cone</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the immunoglobulin superfamily. L1/neurofascin/NgCAM family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000017758.1">
        <property type="protein sequence ID" value="ENSGAGP00000015545.1"/>
        <property type="gene ID" value="ENSGAGG00000006819.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009986">
        <property type="term" value="C:cell surface"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030426">
        <property type="term" value="C:growth cone"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098632">
        <property type="term" value="F:cell-cell adhesion mediator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007411">
        <property type="term" value="P:axon guidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007420">
        <property type="term" value="P:brain development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="CDD" id="cd05845">
        <property type="entry name" value="IgI_2_L1-CAM_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000057">
        <property type="entry name" value="neural cell adhesion molecule L1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000367">
        <property type="entry name" value="Neural cell adhesion molecule L1-like protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000005">
        <property type="entry name" value="Neuronal cell adhesion molecule"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000028">
        <property type="entry name" value="Neuronal cell adhesion molecule"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000038">
        <property type="entry name" value="Neuronal cell adhesion molecule"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000078">
        <property type="entry name" value="Neuronal cell adhesion molecule"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000347">
        <property type="entry name" value="Neuronal cell adhesion molecule"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013098">
        <property type="entry name" value="Ig_I-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013151">
        <property type="entry name" value="Immunoglobulin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026966">
        <property type="entry name" value="Neurofascin/L1/NrCAM_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR44170:SF44">
        <property type="entry name" value="L1 CELL ADHESION MOLECULE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR44170">
        <property type="entry name" value="PROTEIN SIDEKICK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13882">
        <property type="entry name" value="Bravo_FIGEY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
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        <property type="entry name" value="I-set"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="ig"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Ig_3"/>
        <property type="match status" value="2"/>
    </dbReference>
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        <property type="match status" value="4"/>
    </dbReference>
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        <property type="entry name" value="IG"/>
        <property type="match status" value="5"/>
    </dbReference>
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        <property type="entry name" value="IGc2"/>
        <property type="match status" value="4"/>
    </dbReference>
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        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="5"/>
    </dbReference>
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        <property type="match status" value="4"/>
    </dbReference>
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        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="5"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="13" id="KW-0966">Cell projection</keyword>
    <keyword evidence="11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="14" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="10 19" id="KW-0472">Membrane</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="5 19" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9 19" id="KW-1133">Transmembrane helix</keyword>
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        <location>
            <begin position="941"/>
            <end position="962"/>
        </location>
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            <begin position="1"/>
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            <begin position="435"/>
            <end position="519"/>
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            <begin position="528"/>
            <end position="626"/>
        </location>
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    <feature type="domain" description="Fibronectin type-III" evidence="21">
        <location>
            <begin position="628"/>
            <end position="724"/>
        </location>
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    <feature type="domain" description="Fibronectin type-III" evidence="21">
        <location>
            <begin position="729"/>
            <end position="831"/>
        </location>
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    <feature type="domain" description="Fibronectin type-III" evidence="21">
        <location>
            <begin position="835"/>
            <end position="930"/>
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        <location>
            <begin position="40"/>
            <end position="72"/>
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    </feature>
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        <location>
            <begin position="611"/>
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        <location>
            <begin position="973"/>
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        <location>
            <begin position="1041"/>
            <end position="1071"/>
        </location>
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        <location>
            <begin position="63"/>
            <end position="72"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="973"/>
            <end position="991"/>
        </location>
    </feature>
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        <location>
            <begin position="1057"/>
            <end position="1071"/>
        </location>
    </feature>
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        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00004624"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00008588"/>
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    </evidence>
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            <dbReference type="ARBA" id="ARBA00022475"/>
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    </evidence>
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    </evidence>
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            <dbReference type="Proteomes" id="UP000291020"/>
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    <sequence length="1071" mass="118400" checksum="F219010EE9AED8A7" modified="2019-05-08" version="1">MGEXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXPPPRVSTDLLCLPTPHSSPPRPLKPPVFRWTKDGRWFDPRQEAGVTVVEGSGTLYVNNSNAGRHQGVYRCYASNDLGTAMGPEIRVIAESTPQWPKEKVLPLEVEEGESVILPCNPPASAMPPKIYWLNSRIVHIAQDERVSMGQDGYLYFANVRRSDSHPDYICHAHYLGPRTIIQKEPIELRVRPSKLGANSVKFRQPRLMVPPGESSSHVALRGSSLALECIAEGFPTPTVEWIRLNGRLPAERVALENSQKTLRIRDVVEEDDGEYQCLARNTQGTARHTYTVTVEAAPYWAQQPHSRVYGPGENVRLDCSVYGKPNPLVSWRINGVPWGELAPDPTRTLRSGALILSQVQPNDSLVTQCEAVNHHGRLLANAYIYVIRECPLPARILTPDKQRYAVVENQTASLDCQTFGAPMPSVEWFTGRMEPALQDERAFVHTNGSLRLPRAQRQDTGTYVCLAQNDQTNVTISAWLEVKGGTPAPPSMQPVSEPQERWVKLSWTPGDDHNSPIEKFVVEAEDGLRDPGTWRDVSTVPGHQPWAQLHLVPYGDFRFRVRAANRFGRGEPSAASAPVRTSPAAPERYPEGVAGEGNETNNMLITWKPLNRSDWNAPDLSYRVQWRRLGKGGGWSEETVSGPPLLVTRTPTFVPYEIKVQAVNPIGKGPEPPTVVGYSGEDVPQVNPENVGVEIFNSTSVLVSWTLPNRDQLRGHLRGFRVFYWRLGSVGERSRRXPPPLVLTVEGEASWARLGGLRPWSWYRLYVVVFNGRSDGPHSEDIDFSTPEGVPSPPSSLLLERVSDTVLALEWTAPQFPNGALTEYLLQYQQVNGSELGPLHEISFPVSQLNGTLARLDPRAWYRFQLRALTRAGQGEPLVQEGSTVPEPGKGRSFATEGWFIGFICAVVLLLLLLLILCFVKRSKGGKYSVKDKEDIQVDSEARPMKDETFGEYSDNEEKPFGSSQPSLNGALPALGSEDSLADYGGSVDVQFNEDGSFIGQYSGHTGKEPAGNNSSGATSPTTATAALE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKT6</accession>
    <name>A0A452HKT6_9SAUR</name>
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        <recommendedName>
            <fullName evidence="2">Nascent polypeptide-associated complex subunit alpha-like UBA domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
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            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HKT6"/>
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        <property type="protein sequence ID" value="ENSGAGP00000015538.1"/>
        <property type="gene ID" value="ENSGAGG00000011722.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
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        <property type="term" value="P:negative regulation of apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="P:protein stabilization"/>
        <property type="evidence" value="ECO:0007669"/>
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    </dbReference>
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        <property type="entry name" value="UBA_HYPK"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="DNA helicase RuvA subunit, C-terminal domain"/>
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    </dbReference>
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        <property type="entry name" value="NAC-like_UBA"/>
    </dbReference>
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        <property type="entry name" value="HUNTINGTIN-INTERACTING PROTEIN K"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31184">
        <property type="entry name" value="HUNTINGTIN-INTERACTING PROTEIN K FAMILY MEMBER"/>
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    <proteinExistence type="predicted"/>
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            <end position="128"/>
        </location>
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        </location>
    </feature>
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            <begin position="68"/>
            <end position="80"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF19026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015538.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="129" mass="14106" checksum="B201265A09F9503C" modified="2019-05-08" version="1">MAAEGDVELELETEPNGAGGGADGAGGRPAEKPRKHDSGAADLERVTDYAEEKEIQSSNLETAMSVIGDRRSREQKAKQEREKELAKVTIKKEDLELIMNEMEISRAAAERSLREHMGNVVEALITLTN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKU1</accession>
    <name>A0A452HKU1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Latent transforming growth factor beta binding protein 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="9">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000017763.1">
        <property type="protein sequence ID" value="ENSGAGP00000015550.1"/>
        <property type="gene ID" value="ENSGAGG00000008780.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019838">
        <property type="term" value="F:growth factor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00054">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="12"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="18"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.290.10">
        <property type="entry name" value="TGF-beta binding (TB) domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026823">
        <property type="entry name" value="cEGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001881">
        <property type="entry name" value="EGF-like_Ca-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013032">
        <property type="entry name" value="EGF-like_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000152">
        <property type="entry name" value="EGF-type_Asp/Asn_hydroxyl_site"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018097">
        <property type="entry name" value="EGF_Ca-bd_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009030">
        <property type="entry name" value="Growth_fac_rcpt_cys_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049883">
        <property type="entry name" value="NOTCH1_EGF-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017878">
        <property type="entry name" value="TB_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036773">
        <property type="entry name" value="TB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052080">
        <property type="entry name" value="vWF_C/EGF_Fibrillin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47333:SF4">
        <property type="entry name" value="EGF-LIKE DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47333">
        <property type="entry name" value="VON WILLEBRAND FACTOR C AND EGF DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12662">
        <property type="entry name" value="cEGF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07645">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12661">
        <property type="entry name" value="hEGF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00683">
        <property type="entry name" value="TB"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="19"/>
    </dbReference>
    <dbReference type="SMART" id="SM00179">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="17"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57184">
        <property type="entry name" value="Growth factor receptor domain"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57581">
        <property type="entry name" value="TB module/8-cys domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00010">
        <property type="entry name" value="ASX_HYDROXYL"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01186">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="11"/>
    </dbReference>
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        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="7"/>
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    <dbReference type="PROSITE" id="PS51364">
        <property type="entry name" value="TB"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7 9" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="4 9" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="3" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="8" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <feature type="domain" description="EGF-like" evidence="11">
        <location>
            <begin position="59"/>
            <end position="91"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="11">
        <location>
            <begin position="265"/>
            <end position="297"/>
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            <begin position="412"/>
            <end position="455"/>
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        <location>
            <begin position="532"/>
            <end position="584"/>
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        <location>
            <begin position="692"/>
            <end position="733"/>
        </location>
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    <feature type="domain" description="EGF-like" evidence="11">
        <location>
            <begin position="734"/>
            <end position="770"/>
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    <feature type="domain" description="EGF-like" evidence="11">
        <location>
            <begin position="777"/>
            <end position="816"/>
        </location>
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    <feature type="domain" description="EGF-like" evidence="11">
        <location>
            <begin position="857"/>
            <end position="897"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="11">
        <location>
            <begin position="982"/>
            <end position="1024"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="11">
        <location>
            <begin position="1055"/>
            <end position="1095"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="11">
        <location>
            <begin position="1140"/>
            <end position="1181"/>
        </location>
    </feature>
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        <location>
            <begin position="1246"/>
            <end position="1298"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="11">
        <location>
            <begin position="1320"/>
            <end position="1358"/>
        </location>
    </feature>
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        <location>
            <begin position="1422"/>
            <end position="1474"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="11">
        <location>
            <begin position="1611"/>
            <end position="1655"/>
        </location>
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    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1"/>
            <end position="30"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="382"/>
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        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="605"/>
            <end position="628"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="7"/>
            <end position="17"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="382"/>
            <end position="406"/>
        </location>
    </feature>
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        <location>
            <begin position="63"/>
            <end position="73"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="9">
        <location>
            <begin position="81"/>
            <end position="90"/>
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    </feature>
    <feature type="disulfide bond" evidence="9">
        <location>
            <begin position="269"/>
            <end position="279"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="9">
        <location>
            <begin position="287"/>
            <end position="296"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
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            <dbReference type="ARBA" id="ARBA00004498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
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    </evidence>
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    </evidence>
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        <source>
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    </evidence>
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    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
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    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS51364"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015550.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="15">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1667" mass="180403" checksum="FAA3CC6F0C753363" modified="2025-10-08" version="2">MSAPIHRSQSGASSQPCSPAGAGQARDPTPQRARLKCLEDQIQPLGWSHVSLFTPFHVPPAVCDPPCQNKGSCSRPQLCICRSGFQGAQCEEVVPERVYHPPGSVAALQPPAGSLPKRAGSAEREGMRPCRLRSLPTFLQPQSELSPAGRTQGRIVFPVSRHTGSSRTVRRYPAGNTQLMSNALPNGNGHEQSSSGPHAAYQEHTLSLWGANLTEKIRKIKIVFTPTICKQTCRSGHCYNSCQKGDTTTLYSQGGQNHDPKSGFRIYFCQIPCLNGGRCVGRDECWCPSNSTGKFCHLPVPKLEKNQVGRVPKTLASGSIKQSMYTLPLSNQLASLNPSLVNVHIKHPPEATVQIHQVARVRGDLGQSEENSVEAVLLSQLPTGPQHSYSNGNSNSITTESGQQQRPPELMGRCFRETLHGQCANPLEGLAKFEDCCGSVGLFWGVNRCIPCPPRPAHPVIENRQVECPQGYKRLNRSHCQDINECLMLGLCKDTECVNTRGSFLCTCKPGTMLDPSRSRCVSDKAVSMEQGLCYRSAAEGVCALPLAQSITKQICCCSRVGKGWGRNCEKCPMLGSETFKEICPAGHGYTYSSSDIRLSMRKAEAEELPFSSEEQGESSDQTPTQAAKRLQLQEALSGGVTGTLARAGTSAAEGVLGCDATLVAGTADATQEPAPHPGQWPVTSPCLSLSEMDRCASVPDPCGPGTCVNLPDRYSCICGPGYQLHPGLPQCIDDDECMRDPCAGKGHCINSVGSYSCLCYPGYTLLASQGTQTCQDLNECEQPSMCQGGQCTNTPGSYHCECARGYIMGHTGQCEDIDECADPSTCPRGRCVNALGSYECLSCAAGYQPMNGRCVDDDECFTVGICAHGLCINLAGSFRCSCYHGYEVTADEKSCQDINECATHNACPMGLCINSEGSFSCAACGAGYTVSRDGSMCEDIDECSSATACPLGICTNTDGSFACRACDAGYMLSSNRFTCEDMNECLASEICSPNGECLNSHGSYFCICAPGFSNAAGGVTKCALDPGYRPGLSLGPVYTHRTASVLIFILSLPDVDECADKSRCLRGQCFNTEGSYRCLCENGFRHSQETDDCVDVDECVDFGDTVCGTWRCQNTPGSYRCVMGCQPGFHRTPLGDCIDIDECANETLCGSHAFCDNTDGSFRCLCDSGYENLPPGQDCVDVNECELMVAVCGTALCENVEGSFLCLCLSDHEEYNTEAGECRPRAGMGDPERPIAPRPAGSERKECYYHISDVQLCDSVLAKNTTKEECCCTEGAAWGDNCETHPCPILVEYSEICPSGKGYIPVEGTLLFGQTSYTDADECEMFGPEICRNGHCVNTVPGYKCFCRSGYLYDSSRLECVDQDECENEMSCVNGECLNTAGSFHCFCGLPLVLDVTRNRCINVSSRAGEWDPEEHEIHLDICWQRVSNYICNQPLRGRRTTYTECCCRYGEAWSQDCALCPHRASEDFAHLCNVARGETEREPGLQERPGYEYGPALEDSHYGLYSPDLGPYYNYLSPEYGAPDASFPQRELSSEVGGSTFLRTPQLPPAQSQPRYLPSQTGDFEGLQAEECGILNGCENGRCVRVREGYTCDCFDGFQLDMTHMACVDINECEEVSSSGPLCEGGTCENTEGSYRCNCSPGFVVQAEPHRCAPAPAQRPEAAQL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKW5</accession>
    <name>A0A452HKW5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Ribonucleoprotein PTB-binding 1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="12">Protein raver-1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Cooperates with PTBP1 to modulate regulated alternative splicing events. Promotes exon skipping. Cooperates with PTBP1 to modulate switching between mutually exclusive exons during maturation of the TPM1 pre-mRNA.</text>
    </comment>
    <comment type="subunit">
        <text evidence="10">Interacts with PTBP1, RAVER2, VCL and ACTN1. Part of a complex containing RAVER1, VCL and ACTN1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKW5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017795.1">
        <property type="protein sequence ID" value="ENSGAGP00000015579.1"/>
        <property type="gene ID" value="ENSGAGG00000011748.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003729">
        <property type="term" value="F:mRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd12663">
        <property type="entry name" value="RRM1_RAVER1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12665">
        <property type="entry name" value="RRM2_RAVER1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12667">
        <property type="entry name" value="RRM3_RAVER1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.330:FF:000100">
        <property type="entry name" value="Putative ribonucleoprotein PTB-binding 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.330:FF:000116">
        <property type="entry name" value="Putative ribonucleoprotein PTB-binding 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.330:FF:000125">
        <property type="entry name" value="Putative ribonucleoprotein PTB-binding 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.330">
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050502">
        <property type="entry name" value="Euk_RNA-bind_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012677">
        <property type="entry name" value="Nucleotide-bd_a/b_plait_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034633">
        <property type="entry name" value="RAVER1_RRM1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034635">
        <property type="entry name" value="RAVER1_RRM3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035979">
        <property type="entry name" value="RBD_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000504">
        <property type="entry name" value="RRM_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48025">
        <property type="entry name" value="OS02G0815200 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48025:SF1">
        <property type="entry name" value="RRM DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00076">
        <property type="entry name" value="RRM_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00360">
        <property type="entry name" value="RRM"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54928">
        <property type="entry name" value="RNA-binding domain, RBD"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50102">
        <property type="entry name" value="RRM"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0007">Acetylation</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="6 13" id="KW-0694">RNA-binding</keyword>
    <feature type="domain" description="RRM" evidence="15">
        <location>
            <begin position="55"/>
            <end position="126"/>
        </location>
    </feature>
    <feature type="domain" description="RRM" evidence="15">
        <location>
            <begin position="128"/>
            <end position="206"/>
        </location>
    </feature>
    <feature type="domain" description="RRM" evidence="15">
        <location>
            <begin position="217"/>
            <end position="295"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1"/>
            <end position="47"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="677"/>
            <end position="711"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="1"/>
            <end position="13"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="38"/>
            <end position="47"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00058259"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00066243"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00072395"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00076009"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00176"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50102"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015579.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="733" mass="77660" checksum="25FEF3E6E08F3709" modified="2019-05-08" version="1">MAAVSVSGAASVPSPEPPGPEPGPDRVPEEELPSLDPAEVRSRLERSSHQFRNRRKVLIRGLPGDVTNQEVHDLLSDYELKYCFVDKYKGTAFVTLLNGEQAESAIQKFHLTKLRDKEISVQLQPTDALLCIANLPRLYTQQQFEELVRPFGNLERCFLVYSEKTGHSKGYGFVEYMKKDSAARAKSELLGKQLGTQTLYVHWTDVNQLTVDLVHSKCLCVGKLPHNYSDLEELRQVFSAISAPTFCQLAYGQDGQLKGFGVLEYESAEIAEMVQQATDGLPLAGNHIRVSFCAPGPPGRSMLAALIAAQATALNRGKGLLPEPNILQILNSLGNPASIQLLLNPLLHGAVGGKQGILGAPPSMPLVTNPALSAALLQLALQNQTQAQQKPGILGDSPLSSLQHSTLGLATAPAQPGSSLLGELASGGPLPADMTQGHVKSPIVPSGGMPLASFLGPVGTERENSAMGTQVSQLTSSPGTPSPLRGLTTSILGSVISGLQKPKQVENGASLLGEPPKDFKIPLNPYLNLHTLLPANSLGGGANKAFTLKAGVLGNVSNPRIPQPPMADPPLPSPGLAGDNYAFDYQPDLGSRMYAQSRDHAGPILGGFGHSRHKMSSSPGFERSSLGSPLPPFYSGSPTSYFTSGLQAGLKQSHLNKAVGMPPVGTGDNILGLGPNSHSHGSHSKTPIGGQKRAFSHLLPSPEPSPEGGYVGQHSQGLGGHYADSYLKRKRIF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKW9</accession>
    <name>A0A452HKW9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="13">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="8">Belongs to the BET family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000017803.1">
        <property type="protein sequence ID" value="ENSGAGP00000015587.1"/>
        <property type="gene ID" value="ENSGAGG00000011754.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000785">
        <property type="term" value="C:chromatin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006338">
        <property type="term" value="P:chromatin remodeling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006355">
        <property type="term" value="P:regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05497">
        <property type="entry name" value="Bromo_Brdt_I_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05498">
        <property type="entry name" value="Bromo_Brdt_II_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.920.10:FF:000003">
        <property type="entry name" value="Bromodomain-containing protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1270.220:FF:000001">
        <property type="entry name" value="bromodomain-containing protein 2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.920.10:FF:000002">
        <property type="entry name" value="Bromodomain-containing protein 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.220">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.920.10">
        <property type="entry name" value="Bromodomain-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031354">
        <property type="entry name" value="BRD4_CDT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043508">
        <property type="entry name" value="Bromo_Brdt_I"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043509">
        <property type="entry name" value="Bromo_Brdt_II"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050935">
        <property type="entry name" value="Bromo_chromatin_reader"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001487">
        <property type="entry name" value="Bromodomain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036427">
        <property type="entry name" value="Bromodomain-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018359">
        <property type="entry name" value="Bromodomain_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027353">
        <property type="entry name" value="NET_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038336">
        <property type="entry name" value="NET_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22880:SF143">
        <property type="entry name" value="BROMODOMAIN-CONTAINING PROTEIN 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22880">
        <property type="entry name" value="FALZ-RELATED BROMODOMAIN-CONTAINING PROTEINS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17035">
        <property type="entry name" value="BET"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17105">
        <property type="entry name" value="BRD4_CDT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00439">
        <property type="entry name" value="Bromodomain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00503">
        <property type="entry name" value="BROMODOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00297">
        <property type="entry name" value="BROMO"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47370">
        <property type="entry name" value="Bromodomain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00633">
        <property type="entry name" value="BROMODOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50014">
        <property type="entry name" value="BROMODOMAIN_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51525">
        <property type="entry name" value="NET"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 9" id="KW-0103">Bromodomain</keyword>
    <keyword evidence="3" id="KW-0156">Chromatin regulator</keyword>
    <keyword evidence="7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0804">Transcription</keyword>
    <keyword evidence="4" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="Bromo" evidence="11">
        <location>
            <begin position="75"/>
            <end position="147"/>
        </location>
    </feature>
    <feature type="domain" description="Bromo" evidence="11">
        <location>
            <begin position="366"/>
            <end position="438"/>
        </location>
    </feature>
    <feature type="domain" description="NET" evidence="12">
        <location>
            <begin position="600"/>
            <end position="682"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1"/>
            <end position="58"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="176"/>
            <end position="346"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="459"/>
            <end position="613"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="673"/>
            <end position="899"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="924"/>
            <end position="1295"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="23"/>
            <end position="40"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="200"/>
            <end position="213"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="214"/>
            <end position="226"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="232"/>
            <end position="241"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="10">
        <location>
            <begin position="242"/>
            <end position="264"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="317"/>
            <end position="333"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="335"/>
            <end position="346"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="479"/>
            <end position="495"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="10">
        <location>
            <begin position="533"/>
            <end position="551"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="552"/>
            <end position="568"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="699"/>
            <end position="712"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="10">
        <location>
            <begin position="724"/>
            <end position="744"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="10">
        <location>
            <begin position="752"/>
            <end position="774"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="10">
        <location>
            <begin position="854"/>
            <end position="863"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="10">
        <location>
            <begin position="873"/>
            <end position="882"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="928"/>
            <end position="959"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="10">
        <location>
            <begin position="960"/>
            <end position="980"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="1123"/>
            <end position="1144"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="1159"/>
            <end position="1170"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="1171"/>
            <end position="1235"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="1236"/>
            <end position="1264"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="1268"/>
            <end position="1285"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023117"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00044509"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00035"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50014"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS51525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015587.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1295" mass="145700" checksum="4D76E6258B87F003" modified="2019-05-08" version="1">MSLESGPGTRLRHLPVMGDGLETTQMSTTQAQTQPQQGNTVGTNPPPPETSNPNKPKRQTNQLQYLLKVVLKTLWKHQFAWPFHHPVDAVKLNLPDYYKIIKTPMDMGTIRKRLENNYYWNAQECIQDFNTMFTNCYIYNKPGDDIVLMAEALEKLFLQKISEMTEEETEIVIVQAKGRGRGRKEAVTPKPGASTVQNVTQASSPAQTQASQQNLQSVQTTHSFPSVTPDLIVQPPVMTVVPPQPPAPPPPPPPPSVPPTPAPQPIQAHPPVIPTAAQPVKTKKGVKRKADTTTPTTIDAIHESSLPTEPKSAKMGPRRESSRPVKPPKKDVPDSQQHVVEKSNSNKVSEQLKYCSGIIKEMFAKKHAAYAWPFYKPVDVEALGLHDYCDIIKHPMDLSTIKSKLENREYRDAQEFAADVRLMFSNCYKYNPADHEVVAMARKLQDVFEMRFAKMPDEPEEPMIPASSPVVVPPPTKVAPPSSSDSSSDSSSDSDSSSDDSEEERAQRLAELQEQLKAVHEQLAALSQPQQSKPKKKEKDKKEKKKEKHKKKEEVEENKKSKTKEPPLKKAKKNNSNSSNSSSKKEPVMVKNSKPPPVYESEEEDKCKPMSYEEKRQLSLDINKLPGEKLGRVVHIIQSREPSLKNSNPDEIEIDFETLKPSTLRELERYVTSCLRKKRKPQAEKVDVIAGSSKMKGFSSSESESTSESSSSDSEDSETEMAPKLKKKGHSGREQKKHHHHHHQQVQQQMQPLPPQPVPQQAPPPMKSSPPPFVPAQVPIMEHPLPGNMFDTITHFTQPIMHLPQPEMPPHLPQQPEHSTPPHLNQHAVVPPPALHNALPQQPSRPSNRAAALPPKPSRPPATSPAISQQPMLPQPPLPQPPQVLLEDEEPSTPHHSLPLSTSQMQLYLQQLQKVQPQTPLLPSVKVQSQPQASLQPPAHPSVQQLQQQQQRPMHMQPMQFPPHIPQPQPPPQQHPPPQPQSAKPQQVIQHHPSPRHHKPDPYSTGHVREAPSPLMMHSPQMPQFQGLGHQSPPQQNIQPKKQELRAPSVVQSQPMAAKEEKMHSPVIRNETFSPTLRQEPPKHPESIKPPAHIQQRTEMKPMEGGRPVIRPPEQNVPPPGVQDKDRQKQELKTPVAPKKDLKIKNMGSWASLVQKPLTTPSSTAKSSSDSFEHFKRAAREKEEREKALKAQAEQVEKEKERLRREQERMRSRDDEDALEQARRVQEEVRRRQEQQHQQQQQQQVSAAASAPQAQSSQPQSSQSMLDQQREMARKREQERRRREAVSGGGGIWGL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKX0</accession>
    <name>A0A452HKX0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Rho family-interacting cell polarization regulator 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Apical cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cell projection</location>
            <location evidence="4">Filopodium</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cell projection</location>
            <location evidence="3">Stereocilium membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the RIPOR family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKX0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017798.1">
        <property type="protein sequence ID" value="ENSGAGP00000015582.1"/>
        <property type="gene ID" value="ENSGAGG00000011582.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000017800.1">
        <property type="protein sequence ID" value="ENSGAGP00000015584.1"/>
        <property type="gene ID" value="ENSGAGG00000011582.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016324">
        <property type="term" value="C:apical plasma membrane"/>
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        <property type="entry name" value="RHO family interacting cell polarization regulator 2"/>
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        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="ARM-like"/>
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        <property type="entry name" value="ARM-type_fold"/>
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        <property type="entry name" value="FAM65_N"/>
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        <property type="entry name" value="RIPOR3"/>
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        <property type="entry name" value="PROTEIN KINASE PKN/PRK1, EFFECTOR"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="RHO FAMILY-INTERACTING CELL POLARIZATION REGULATOR 2"/>
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        <property type="entry name" value="PL48"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="ARM repeat"/>
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    <proteinExistence type="inferred from homology"/>
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    <keyword evidence="7" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="18" id="KW-0966">Cell projection</keyword>
    <keyword evidence="9" id="KW-0145">Chemotaxis</keyword>
    <keyword evidence="15" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="8" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="17" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="13" id="KW-0221">Differentiation</keyword>
    <keyword evidence="12" id="KW-1009">Hearing</keyword>
    <keyword evidence="16" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0517">Myogenesis</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="11" id="KW-0734">Signal transduction inhibitor</keyword>
    <feature type="domain" description="FAM65 N-terminal" evidence="20">
        <location>
            <begin position="1"/>
            <end position="235"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
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            <end position="400"/>
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            <begin position="544"/>
            <end position="580"/>
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            <begin position="349"/>
            <end position="359"/>
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    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="360"/>
            <end position="389"/>
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        <location>
            <begin position="566"/>
            <end position="580"/>
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    <evidence type="ECO:0000313" key="22">
        <source>
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    </evidence>
    <sequence length="938" mass="103919" checksum="5BF53E3A49B7BCC9" modified="2019-05-08" version="1">MRRLEFHISMVDELYEAYCIQRRLCDGASKMKQAFAMSPASKAARESLSEINRSYKEYTENMCTIEGELENLLGEFCIKMKGLAGFARLCPGDQYEIFMRYGRQRWKLKGKIEVNGKQSWDGEEMVFLPLIVGLISIKVTEVKGLATHTLVGSVTCETKDLFAARPQVVAVDINDLGTIKLNLEITWYPFDVEDLTPSTGNVNKASALQRRMSTYSQGTPETPTFKDYSFFKWLHPPQDRPQLSVLDALQDTFFDRLQRSRSFSDLPSLRLRPKAGLEQYSNSPDDIFENGMAGTEKRPISFSFGDIPNGDCASPPITVDSASALDPSSPEITVTPPEHKSSKNPAMHSSSTNSSLDSTPESKDLKSQPEHTPVHENKKSSRVASEVHQKPPSPGSDNLFIDSSIPVSLLQETDDVSELKPVELDTYEGNITKQLVKRLTSAEAPMTPERLQCEGSISGESEGYKSYLDGSIEDAFQGLLLALEPHKEQYKEFQDLDQQVMHLDDILKCKPAVNRSRSSSSISLTVESALESFDFLNTSDFDDEDGGGDEVCNGGGGADSVFSDTETEKTSYRSDHPEARGHLSEALTEDTGVGTSVAGSPLPLTTGNESLDITIVKHLQCCTQLIQQIVFSSKTPFVVRNLLDKLSKEIVVLENLAEISIENMGNIISVTEAIPEFHKKLSLLSFWTKCTGPTGVYHTSTDKMIKQLDVTFATAVNEECPGLAETVFRILLSQILDRTEPVLSSSLPSEILTVFQYYNYFASNGVSDLGSYLLQLAKEASVVQKLQTLKDEKLLQNIAELTSSSLPPQQEVLRSLALLLTENKNEIGEAVTLLLSASAKNRHFREKALIYYCEVLTQPNLHLQKAACMALKCLQATESIKMLVTLCQSDNEEIRKVASETLLSLGEDGRLAYEQLDKFPREFVKVGGRWGTEVATAF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKX2</accession>
    <name>A0A452HKX2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Ribonucleoprotein PTB-binding 1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="12">Protein raver-1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Cooperates with PTBP1 to modulate regulated alternative splicing events. Promotes exon skipping. Cooperates with PTBP1 to modulate switching between mutually exclusive exons during maturation of the TPM1 pre-mRNA.</text>
    </comment>
    <comment type="subunit">
        <text evidence="10">Interacts with PTBP1, RAVER2, VCL and ACTN1. Part of a complex containing RAVER1, VCL and ACTN1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKX2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017796.1">
        <property type="protein sequence ID" value="ENSGAGP00000015580.1"/>
        <property type="gene ID" value="ENSGAGG00000011748.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003729">
        <property type="term" value="F:mRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd12663">
        <property type="entry name" value="RRM1_RAVER1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12665">
        <property type="entry name" value="RRM2_RAVER1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12667">
        <property type="entry name" value="RRM3_RAVER1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.330:FF:000100">
        <property type="entry name" value="Putative ribonucleoprotein PTB-binding 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.330:FF:000116">
        <property type="entry name" value="Putative ribonucleoprotein PTB-binding 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.330:FF:000125">
        <property type="entry name" value="Putative ribonucleoprotein PTB-binding 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.330">
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050502">
        <property type="entry name" value="Euk_RNA-bind_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012677">
        <property type="entry name" value="Nucleotide-bd_a/b_plait_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034633">
        <property type="entry name" value="RAVER1_RRM1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034635">
        <property type="entry name" value="RAVER1_RRM3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035979">
        <property type="entry name" value="RBD_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000504">
        <property type="entry name" value="RRM_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48025">
        <property type="entry name" value="OS02G0815200 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48025:SF1">
        <property type="entry name" value="RRM DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00076">
        <property type="entry name" value="RRM_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00360">
        <property type="entry name" value="RRM"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54928">
        <property type="entry name" value="RNA-binding domain, RBD"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50102">
        <property type="entry name" value="RRM"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0007">Acetylation</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="6 13" id="KW-0694">RNA-binding</keyword>
    <feature type="domain" description="RRM" evidence="15">
        <location>
            <begin position="55"/>
            <end position="126"/>
        </location>
    </feature>
    <feature type="domain" description="RRM" evidence="15">
        <location>
            <begin position="128"/>
            <end position="206"/>
        </location>
    </feature>
    <feature type="domain" description="RRM" evidence="15">
        <location>
            <begin position="217"/>
            <end position="295"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1"/>
            <end position="47"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="558"/>
            <end position="597"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="1"/>
            <end position="13"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="38"/>
            <end position="47"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00058259"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00066243"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00072395"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00076009"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00176"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50102"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015580.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="614" mass="65929" checksum="AF70418990C06985" modified="2019-05-08" version="1">MAAVSVSGAASVPSPEPPGPEPGPDRVPEEELPSLDPAEVRSRLERSSHQFRNRRKVLIRGLPGDVTNQEVHDLLSDYELKYCFVDKYKGTAFVTLLNGEQAESAIQKFHLTKLRDKEISVQLQPTDALLCIANLPRLYTQQQFEELVRPFGNLERCFLVYSEKTGHSKGYGFVEYMKKDSAARAKSELLGKQLGTQTLYVHWTDVNQLTVDLVHSKCLCVGKLPHNYSDLEELRQVFSAISAPTFCQLAYGQDGQLKGFGVLEYESAEIAEMVQQATDGLPLAGNHIRVSFCAPGPPGRSMLAALIAAQATALNRGKGLLPEPNILQILNSLGNPASIQLLLNPLLHGAVGGKQGILGAPPSMPLVTNPALSAALLQLALQNQTQAQQASLLGEPPKDFKIPLNPYLNLHTLLPANSLGGGANKAFTLKAGVLGNVSNPRIPQPPMADPPLPSPGLAGDNYAFDYQPDLGSRMYAQSRDHAGPILGGFGHSRHKMSSSPGFERSSLGSPLPPFYSGSPTSYFTSGLQAGLKQSHLNKAVGMPPVGTGDNILGLGPNSHSHGSHSKTPIGGQKRAFSHLLPSPEPSPEGGYVGQHSQGLGGHYADSYLKRKRIF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKX3</accession>
    <name>A0A452HKX3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Rho family-interacting cell polarization regulator 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Apical cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cell projection</location>
            <location evidence="4">Filopodium</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cell projection</location>
            <location evidence="3">Stereocilium membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the RIPOR family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKX3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017794.1">
        <property type="protein sequence ID" value="ENSGAGP00000015578.1"/>
        <property type="gene ID" value="ENSGAGG00000011582.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016324">
        <property type="term" value="C:apical plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030175">
        <property type="term" value="C:filopodium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060171">
        <property type="term" value="C:stereocilium membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006935">
        <property type="term" value="P:chemotaxis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007517">
        <property type="term" value="P:muscle organ development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009968">
        <property type="term" value="P:negative regulation of signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007605">
        <property type="term" value="P:sensory perception of sound"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000191">
        <property type="entry name" value="RHO family interacting cell polarization regulator 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031780">
        <property type="entry name" value="FAM65_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026136">
        <property type="entry name" value="RIPOR3"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15829">
        <property type="entry name" value="PROTEIN KINASE PKN/PRK1, EFFECTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15829:SF2">
        <property type="entry name" value="RHO FAMILY-INTERACTING CELL POLARIZATION REGULATOR 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15903">
        <property type="entry name" value="PL48"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="14" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="7" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="18" id="KW-0966">Cell projection</keyword>
    <keyword evidence="9" id="KW-0145">Chemotaxis</keyword>
    <keyword evidence="15" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="8" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="17" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="13" id="KW-0221">Differentiation</keyword>
    <keyword evidence="12" id="KW-1009">Hearing</keyword>
    <keyword evidence="16" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0517">Myogenesis</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="11" id="KW-0734">Signal transduction inhibitor</keyword>
    <feature type="domain" description="FAM65 N-terminal" evidence="20">
        <location>
            <begin position="5"/>
            <end position="308"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1"/>
            <end position="25"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="326"/>
            <end position="427"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="571"/>
            <end position="607"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="1"/>
            <end position="21"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="19">
        <location>
            <begin position="376"/>
            <end position="386"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="387"/>
            <end position="416"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="593"/>
            <end position="607"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004289"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004486"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00005744"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00013627"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022500"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022541"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022700"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022740"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="Pfam" id="PF15903"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015578.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="965" mass="107039" checksum="F615D86EB174C222" modified="2019-05-08" version="1">MSILQKTHNLGHKNSTTSKEPQPQRMEEVYRALKNGLDEYLEVHQTELDKLTAQLKDMKRNSRVLCYLHRSRVTFSRYRRKRGNKGVDELYEAYCIQRRLCDGASKMKQAFAMSPASKAARESLSEINRSYKEYTENMCTIEGELENLLGEFCIKMKGLAGFARLCPGDQYEIFMRYGRQRWKLKGKIEVNGKQSWDGEEMVFLPLIVGLISIKVTEVKGLATHTLVGSVTCETKDLFAARPQVVAVDINDLGTIKLNLEITWYPFDVEDLTPSTGNVNKASALQRRMSTYSQGTPETPTFKDYSFFSNSPDDIFENGMAGTEKRPISFSFGDIPNGDCASPPITVDSASALDPSSPEITVTPPEHKSSKNPAMHSSSTNSSLDSTPESKDLKSQPEHTPVHENKKSSRVASEVHQKPPSPGSDNLFIDSSIPVSLLQETDDVSELKPVELDTYEGNITKQLVKRLTSAEAPMTPERLQCEGSISGESEGYKSYLDGSIEDAFQGLLLALEPHKEQYKEFQDLDQQVMHLDDILKCKPAVNRSRSSSSISLTVESALESFDFLNTSDFDDEDGGGDEVCNGGGGADSVFSDTETEKTSYRSDHPEARGHLSEALTEDTGVGTSVAGSPLPLTTGNESLDITIVKHLQCCTQLIQQIVFSSKTPFVVRNLLDKLSKEIVVLENLAEISIENMGNIISVTEAIPEFHKKLSLLSFWTKCTGPTGVYHTSTDKMIKQLDVTFATAVNEECPGLAETVFRILLSQILDRTEPVLSSSLPSEILTVFQYYNYFASNGVSDLGSYLLQLAKEASVVQKLQTLKDEKLLQNIAELTSSSLPPQQEVLRSLALLLTENKNEIGEAVTLLLSASAKNRHFREKALIYYCEVLTQPNLHLQKAACMALKCLQATESIKMLVTLCQSDNEEIRKVASETLLSLGEDGRLAYEQLDKFPREFVKVGGRWGTEVATAF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKX4</accession>
    <name>A0A452HKX4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Rho family-interacting cell polarization regulator 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Apical cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cell projection</location>
            <location evidence="4">Filopodium</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cell projection</location>
            <location evidence="3">Stereocilium membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the RIPOR family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKX4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017808.1">
        <property type="protein sequence ID" value="ENSGAGP00000015592.1"/>
        <property type="gene ID" value="ENSGAGG00000011582.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016324">
        <property type="term" value="C:apical plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030175">
        <property type="term" value="C:filopodium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060171">
        <property type="term" value="C:stereocilium membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006935">
        <property type="term" value="P:chemotaxis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007517">
        <property type="term" value="P:muscle organ development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009968">
        <property type="term" value="P:negative regulation of signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007605">
        <property type="term" value="P:sensory perception of sound"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000191">
        <property type="entry name" value="RHO family interacting cell polarization regulator 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031780">
        <property type="entry name" value="FAM65_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026136">
        <property type="entry name" value="RIPOR3"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15829">
        <property type="entry name" value="PROTEIN KINASE PKN/PRK1, EFFECTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15829:SF2">
        <property type="entry name" value="RHO FAMILY-INTERACTING CELL POLARIZATION REGULATOR 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15903">
        <property type="entry name" value="PL48"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="14" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="7" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="18" id="KW-0966">Cell projection</keyword>
    <keyword evidence="9" id="KW-0145">Chemotaxis</keyword>
    <keyword evidence="15" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="8" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="17" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="13" id="KW-0221">Differentiation</keyword>
    <keyword evidence="12" id="KW-1009">Hearing</keyword>
    <keyword evidence="16" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0517">Myogenesis</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="11" id="KW-0734">Signal transduction inhibitor</keyword>
    <feature type="domain" description="FAM65 N-terminal" evidence="20">
        <location>
            <begin position="1"/>
            <end position="232"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="250"/>
            <end position="351"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="495"/>
            <end position="531"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="19">
        <location>
            <begin position="300"/>
            <end position="310"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="311"/>
            <end position="340"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="517"/>
            <end position="531"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004289"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004486"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00005744"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00013627"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022500"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022541"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022700"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022740"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="Pfam" id="PF15903"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015592.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="889" mass="98114" checksum="D6E397A085624A9F" modified="2019-05-08" version="1">MRRLEFHISMVDELYEAYCIQRRLCDGASKMKQAFAMSPASKAARESLSEINRSYKEYTENMCTIEGELENLLGEFCIKMKGLAGFARLCPGDQYEIFMRYGRQRWKLKGKIEVNGKQSWDGEEMVFLPLIVGLISIKVTEVKGLATHTLVGSVTCETKDLFAARPQVVAVDINDLGTIKLNLEITWYPFDVEDLTPSTGNVNKASALQRRMSTYSQGTPETPTFKDYSFFSNSPDDIFENGMAGTEKRPISFSFGDIPNGDCASPPITVDSASALDPSSPEITVTPPEHKSSKNPAMHSSSTNSSLDSTPESKDLKSQPEHTPVHENKKSSRVASEVHQKPPSPGSDNLFIDSSIPVSLLQETDDVSELKPVELDTYEGNITKQLVKRLTSAEAPMTPERLQCEGSISGESEGYKSYLDGSIEDAFQGLLLALEPHKEQYKEFQDLDQQVMHLDDILKCKPAVNRSRSSSSISLTVESALESFDFLNTSDFDDEDGGGDEVCNGGGGADSVFSDTETEKTSYRSDHPEARGHLSEALTEDTGVGTSVAGSPLPLTTGNESLDITIVKHLQCCTQLIQQIVFSSKTPFVVRNLLDKLSKEIVVLENLAEISIENMGNIISVTEAIPEFHKKLSLLSFWTKCTGPTGVYHTSTDKMIKQLDVTFATAVNEECPGLAETVFRILLSQILDRTEPVLSSSLPSEILTVFQYYNYFASNGVSDLGSYLLQLAKEASVVQKLQTLKDEKLLQNIAELTSSSLPPQQEVLRSLALLLTENKNEIGEAVTLLLSASAKNRHFREKALIYYCEVLTQPNLHLQKAACMALKCLQATESIKMLVTLCQSDNEEIRKVASETLLSLGEDGRLAYEQLDKFPREFVKVGGRWGTEVATAF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKX5</accession>
    <name>A0A452HKX5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Ribonucleoprotein PTB-binding 1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="12">Protein raver-1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Cooperates with PTBP1 to modulate regulated alternative splicing events. Promotes exon skipping. Cooperates with PTBP1 to modulate switching between mutually exclusive exons during maturation of the TPM1 pre-mRNA.</text>
    </comment>
    <comment type="subunit">
        <text evidence="10">Interacts with PTBP1, RAVER2, VCL and ACTN1. Part of a complex containing RAVER1, VCL and ACTN1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKX5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017792.1">
        <property type="protein sequence ID" value="ENSGAGP00000015576.1"/>
        <property type="gene ID" value="ENSGAGG00000011748.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003729">
        <property type="term" value="F:mRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd12663">
        <property type="entry name" value="RRM1_RAVER1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12665">
        <property type="entry name" value="RRM2_RAVER1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12667">
        <property type="entry name" value="RRM3_RAVER1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.330:FF:000100">
        <property type="entry name" value="Putative ribonucleoprotein PTB-binding 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.330:FF:000116">
        <property type="entry name" value="Putative ribonucleoprotein PTB-binding 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.330:FF:000125">
        <property type="entry name" value="Putative ribonucleoprotein PTB-binding 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.330">
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050502">
        <property type="entry name" value="Euk_RNA-bind_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012677">
        <property type="entry name" value="Nucleotide-bd_a/b_plait_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034633">
        <property type="entry name" value="RAVER1_RRM1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034635">
        <property type="entry name" value="RAVER1_RRM3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035979">
        <property type="entry name" value="RBD_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000504">
        <property type="entry name" value="RRM_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48025">
        <property type="entry name" value="OS02G0815200 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48025:SF1">
        <property type="entry name" value="RRM DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00076">
        <property type="entry name" value="RRM_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00360">
        <property type="entry name" value="RRM"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54928">
        <property type="entry name" value="RNA-binding domain, RBD"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50102">
        <property type="entry name" value="RRM"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0007">Acetylation</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="6 13" id="KW-0694">RNA-binding</keyword>
    <feature type="domain" description="RRM" evidence="15">
        <location>
            <begin position="55"/>
            <end position="126"/>
        </location>
    </feature>
    <feature type="domain" description="RRM" evidence="15">
        <location>
            <begin position="128"/>
            <end position="206"/>
        </location>
    </feature>
    <feature type="domain" description="RRM" evidence="15">
        <location>
            <begin position="217"/>
            <end position="295"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1"/>
            <end position="47"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="678"/>
            <end position="717"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="1"/>
            <end position="13"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="38"/>
            <end position="47"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00058259"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00066243"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00072395"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00076009"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00176"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50102"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015576.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="734" mass="77869" checksum="8255EE8117FC6EC7" modified="2019-05-08" version="1">MAAVSVSGAASVPSPEPPGPEPGPDRVPEEELPSLDPAEVRSRLERSSHQFRNRRKVLIRGLPGDVTNQEVHDLLSDYELKYCFVDKYKGTAFVTLLNGEQAESAIQKFHLTKLRDKEISVQLQPTDALLCIANLPRLYTQQQFEELVRPFGNLERCFLVYSEKTGHSKGYGFVEYMKKDSAARAKSELLGKQLGTQTLYVHWTDVNQLTVDLVHSKCLCVGKLPHNYSDLEELRQVFSAISAPTFCQLAYGQDGQLKGFGVLEYESAEIAEMVQQATDGLPLAGNHIRVSFCAPGPPGRSMLAALIAAQATALNRGKGLLPEPNILQILNSLGNPASIQLLLNPLLHGAVGGKQGILGAPPSMPLVTNPALSAALLQLALQNQTQAQQALLGSSLLLQNQVWTQLLQNKENQAVLHKPGILGDSPLSSLQHSTLGLATAPAQPGSSLLGELASGGPLPADMTQGHVKSPIVPSGGMPLASFLGPVGTERENSAMGTQVSQLTSSPGTPSPLRGLTTSILGSVISGLQKPKQVENGASLLGEPPKDFKIPLNPYLNLHTLLPANSLGGGANKAFTLKAGVLGNVSNPRIPQPPMADPPLPSPGLAGDNYAFDYQPMSSSPGFERSSLGSPLPPFYSGSPTSYFTSGLQAGLKQSHLNKAVGMPPVGTGDNILGLGPNSHSHGSHSKTPIGGQKRAFSHLLPSPEPSPEGGYVGQHSQGLGGHYADSYLKRKRIF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKZ1</accession>
    <name>A0A452HKZ1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Atypical kinase COQ8A, mitochondrial</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="16">Chaperone activity of bc1 complex-like</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="17">Coenzyme Q protein 8A</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="15">aarF domain-containing protein kinase 3</fullName>
        </alternativeName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="18">Atypical kinase involved in the biosynthesis of coenzyme Q, also named ubiquinone, an essential lipid-soluble electron transporter for aerobic cellular respiration. Its substrate specificity is still unclear: may act as a protein kinase that mediates phosphorylation of COQ3. According to other reports, acts as a small molecule kinase, possibly a lipid kinase that phosphorylates a prenyl lipid in the ubiquinone biosynthesis pathway, as suggested by its ability to bind coenzyme Q lipid intermediates. However, the small molecule kinase activity was not confirmed by another publication. Shows an unusual selectivity for binding ADP over ATP.</text>
    </comment>
    <comment type="pathway">
        <text evidence="2">Cofactor biosynthesis; ubiquinone biosynthesis.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Mitochondrion membrane</location>
            <topology evidence="1">Single-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the protein kinase superfamily. ADCK protein kinase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKZ1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017807.1">
        <property type="protein sequence ID" value="ENSGAGP00000015591.1"/>
        <property type="gene ID" value="ENSGAGG00000011753.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00232"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031966">
        <property type="term" value="C:mitochondrial membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016301">
        <property type="term" value="F:kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006744">
        <property type="term" value="P:ubiquinone biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="CDD" id="cd13970">
        <property type="entry name" value="ABC1_ADCK3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004147">
        <property type="entry name" value="ABC1_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034646">
        <property type="entry name" value="ADCK3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051409">
        <property type="entry name" value="Atypical_kinase_ADCK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43851">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43851:SF1">
        <property type="entry name" value="ATYPICAL KINASE COQ8A, MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03109">
        <property type="entry name" value="ABC1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="9" id="KW-0418">Kinase</keyword>
    <keyword evidence="14" id="KW-0472">Membrane</keyword>
    <keyword evidence="13" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="8" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0808">Transferase</keyword>
    <keyword evidence="11" id="KW-0809">Transit peptide</keyword>
    <keyword evidence="7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="12" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="6" id="KW-0831">Ubiquinone biosynthesis</keyword>
    <feature type="domain" description="ABC1 atypical kinase-like" evidence="20">
        <location>
            <begin position="300"/>
            <end position="526"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="72"/>
            <end position="114"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="182"/>
            <end position="203"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="79"/>
            <end position="97"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004304"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004749"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00009670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00018535"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022688"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00031775"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00032726"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00033204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00058956"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="Pfam" id="PF03109"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015591.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="621" mass="69078" checksum="77F0796052B32F65" modified="2019-05-08" version="1">MAGDAIMLIRGLAKLSKAIIETQVGQLRQGIGGGDAAILARTFQTTVEEQFSAAMGKMQDLGKQQENLTDLGEDFGSDYNFSGPESSNATKDYSSPGQPDKGNGTEGPAYTYATNGPFRNVSEIGDTSPGQIPFSAKVNAKLFGGFRDPGNPFAAAIGQNRAFHQDHSSVSGLTAEDIDKARQAKTDPEHKPHKQMLSERARERKVPVTRLGRLANFGGLAVGLGIGALAEVAKKSLRHEERNGKKAVMDSSPFLSEANAERIVRTLCKVRGAALKLGQMLSIQDNAFINPHLQKIFERVRQSADFMPIKQMMKTLNDDLGPNWRDKLEFFEERPFAAASIGQVHLARLKNGKEVAMKIQYPGVAQSIDSDVSNLMTVLSMSNVLPEGLFHEHMMEVLSRELAMECDYKREAACAKKFKELLKDHPFFYVPDVVDELCSQHVLTTDLVSGFPLDQAEGLSQEIRNEICSNILILCLRELFEFRFMQTDPNWSNFFYDPQLHKVALLDFGASRGFGEDFTDLYIEEMENAHLEAVLILGEAFASEEPFDFGTQSTTEKIHGLIPVMLKHRLVPPPEETYSLHRKMGGSFLICSKLKAKIACKNMFQEAYSKYWSKRAKEQEQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKZ2</accession>
    <name>A0A452HKZ2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Scaffolding anchor of CK1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the FAM83 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000017822.1">
        <property type="protein sequence ID" value="ENSGAGP00000015604.1"/>
        <property type="gene ID" value="ENSGAGG00000011761.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019901">
        <property type="term" value="F:protein kinase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.870.10:FF:000004">
        <property type="entry name" value="protein FAM83H isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.870.10">
        <property type="entry name" value="Endonuclease Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050944">
        <property type="entry name" value="FAM83"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012461">
        <property type="entry name" value="SACK1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16181">
        <property type="entry name" value="PROTEIN FAM83A-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16181:SF29">
        <property type="entry name" value="PROTEIN FAM83A-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07894">
        <property type="entry name" value="SACK1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56024">
        <property type="entry name" value="Phospholipase D/nuclease"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Scaffolding anchor of CK1" evidence="5">
        <location>
            <begin position="16"/>
            <end position="297"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="74"/>
            <end position="96"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="319"/>
            <end position="371"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="398"/>
            <end position="459"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="83"/>
            <end position="96"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="426"/>
            <end position="440"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006937"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="Pfam" id="PF07894"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015604.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="481" mass="53686" checksum="2A3CCCEADE82C1D1" modified="2019-05-08" version="1">MWXLSLRLEELKKLWWRKPIPLVLQHSETARLAVDAFLEQGEPGYLQAIAEEQELPFLSPLDMDYMSQHRSQSILEPNARKGHQTEPKEGDSEDRASLLSELSSGTYFPLLSDMQPPELELGWPVIPLATGYGQTQASVYFQRNKANSIKDLLRCLISRAKTVIAVVMDLFTDMEILCDLMEASSRRRVSVYLILDEKYLKYFIEMCSKMGLNKDHFPNMRVRCVSGDTYYSKGGKKFEGQVLEKFVLIDCDHVLAGTYSFTWLCSQVHTCLVTHFRGKIVEDFDREFQRLYVESKPVPGFCIPEAGTKPGLRCAPGGSKAGLGARGRSAVPRGFSSGEAAASSRLPSAGRGRSAQPPKPRPRPRCAATETRVGAALPARFRWLLAAWTXARAALRARRQQGRAGGPRAQRGSPRILQRRSSRVLPPAQRGAGPQRAASQAPPPAALCGHRDAGRRRSTRALPVVAGRLDRKLLRARRRCR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKZ5</accession>
    <name>A0A452HKZ5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">AB hydrolase-1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="2">Belongs to the AB hydrolase superfamily. Epoxide hydrolase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKZ5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017823.1">
        <property type="protein sequence ID" value="ENSGAGP00000015605.1"/>
        <property type="gene ID" value="ENSGAGG00000011762.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004301">
        <property type="term" value="F:epoxide hydrolase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1820">
        <property type="entry name" value="alpha/beta hydrolase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000073">
        <property type="entry name" value="AB_hydrolase_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029058">
        <property type="entry name" value="AB_hydrolase_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000639">
        <property type="entry name" value="Epox_hydrolase-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43329">
        <property type="entry name" value="EPOXIDE HYDROLASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00561">
        <property type="entry name" value="Abhydrolase_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00111">
        <property type="entry name" value="ABHYDROLASE"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00412">
        <property type="entry name" value="EPOXHYDRLASE"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53474">
        <property type="entry name" value="alpha/beta-Hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="1" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="3" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="3" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="3">
        <location>
            <begin position="39"/>
            <end position="60"/>
        </location>
    </feature>
    <feature type="domain" description="AB hydrolase-1" evidence="4">
        <location>
            <begin position="88"/>
            <end position="321"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00038334"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="Pfam" id="PF00561"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015605.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="337" mass="38178" checksum="62101BF509D9A4C2" modified="2019-05-08" version="1">HTCYKQLGFPPPAILTASLTRAQASSVAFLAQVLIQDTFRAVTCTFVLLTLTYLFMTPLLSLQSSGLRLHYVTAGQEEVQLMLCLFAWRYQLQEFKRQFRVVALDLRGYGGSDAPRGKQHYQMETLLVDVHGVIQVLGFTKCILVGHDWGGAIGWEFAMNHPDMVEKLIIMNATYSSILKEYIAGHPCQLLRSSYIFMFQLQRLPELLLSLGDFQLLKDTLAGRKGGIQNPKWHLTAQELEAYIFGLSKPGALTPPLDYYRNIFSRSPMQCKGMLVPMLVICGEKDVYLEAGMIPLLEQYVCKRCRVGRIPEASHFVSEDQPDKVNQLIWSFLQESD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKZ9</accession>
    <name>A0A452HKZ9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Atypical kinase COQ8A, mitochondrial</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="16">Chaperone activity of bc1 complex-like</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="17">Coenzyme Q protein 8A</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="15">aarF domain-containing protein kinase 3</fullName>
        </alternativeName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="18">Atypical kinase involved in the biosynthesis of coenzyme Q, also named ubiquinone, an essential lipid-soluble electron transporter for aerobic cellular respiration. Its substrate specificity is still unclear: may act as a protein kinase that mediates phosphorylation of COQ3. According to other reports, acts as a small molecule kinase, possibly a lipid kinase that phosphorylates a prenyl lipid in the ubiquinone biosynthesis pathway, as suggested by its ability to bind coenzyme Q lipid intermediates. However, the small molecule kinase activity was not confirmed by another publication. Shows an unusual selectivity for binding ADP over ATP.</text>
    </comment>
    <comment type="pathway">
        <text evidence="2">Cofactor biosynthesis; ubiquinone biosynthesis.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Mitochondrion membrane</location>
            <topology evidence="1">Single-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the protein kinase superfamily. ADCK protein kinase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKZ9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017813.1">
        <property type="protein sequence ID" value="ENSGAGP00000015596.1"/>
        <property type="gene ID" value="ENSGAGG00000011753.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00232"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031966">
        <property type="term" value="C:mitochondrial membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016301">
        <property type="term" value="F:kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006744">
        <property type="term" value="P:ubiquinone biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="CDD" id="cd13970">
        <property type="entry name" value="ABC1_ADCK3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004147">
        <property type="entry name" value="ABC1_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034646">
        <property type="entry name" value="ADCK3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051409">
        <property type="entry name" value="Atypical_kinase_ADCK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43851">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43851:SF1">
        <property type="entry name" value="ATYPICAL KINASE COQ8A, MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03109">
        <property type="entry name" value="ABC1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="9" id="KW-0418">Kinase</keyword>
    <keyword evidence="14" id="KW-0472">Membrane</keyword>
    <keyword evidence="13" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="8" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0808">Transferase</keyword>
    <keyword evidence="11" id="KW-0809">Transit peptide</keyword>
    <keyword evidence="7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="12" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="6" id="KW-0831">Ubiquinone biosynthesis</keyword>
    <feature type="domain" description="ABC1 atypical kinase-like" evidence="20">
        <location>
            <begin position="259"/>
            <end position="498"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="72"/>
            <end position="114"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="182"/>
            <end position="203"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="79"/>
            <end position="97"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004304"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004749"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00009670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00018535"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022688"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00031775"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00032726"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00033204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00058956"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="Pfam" id="PF03109"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015596.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="609" mass="67999" checksum="48166FE173AF832F" modified="2019-05-08" version="1">MAGDAIMLIRGLAKLSKAIIETQVGQLRQGIGGGDAAILARTFQTTVEEQFSAAMGKMQDLGKQQENLTDLGEDFGSDYNFSGPESSNATKDYSSPGQPDKGNGTEGPAYTYATNGPFRNVSEIGDTSPGQIPFSAKVNAKLFGGFRDPGNPFAAAIGQNRAFHQDHSSVSGLTAEDIDKARQAKTDPEHKPHKQMLSERARERKVPVTRLGRLANFGGLAVGLGIGALAEVAKKSLRHEERNDNAFINPHLQKIFERVRQSADFMPIKQMMKTLNDDLGPNWRDKLEFFEERPFAAASIGQVHLARLKNGKEVAMKIQYPGVAQSIDSDVSNLMTVLSMSNVLPEGLFHEHMMEVLSRELAMECDYKREAACAKKFKELLKDHPFFYVPDVVDELCSQHVLTTDLVSGFPLDQAEGLSQEIRNEICSNILILCLRELFEFRFMQTDPNWSNFFYDPQLHKVALLDFGASRGFGEDFTDLYIEVIKAAADQDRETVLRKSIEMKFLTGYEVKEMENAHLEAVLILGEAFASEEPFDFGTQSTTEKIHGLIPVMLKHRLVPPPEETYSLHRKMGGSFLICSKLKAKIACKNMFQEAYSKYWSKRAKEQEQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HL52</accession>
    <name>A0A452HL52_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Fibronectin type-III domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HL52"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017887.1">
        <property type="protein sequence ID" value="ENSGAGP00000015659.1"/>
        <property type="gene ID" value="ENSGAGG00000011789.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098982">
        <property type="term" value="C:GABA-ergic synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098978">
        <property type="term" value="C:glutamatergic synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045296">
        <property type="term" value="F:cadherin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017147">
        <property type="term" value="F:Wnt-protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007411">
        <property type="term" value="P:axon guidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090090">
        <property type="term" value="P:negative regulation of canonical Wnt signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072112">
        <property type="term" value="P:podocyte differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003093">
        <property type="term" value="P:regulation of glomerular filtration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000488">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase O isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000415">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase O isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058859">
        <property type="entry name" value="Fn3_R-PTP-O"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042996">
        <property type="entry name" value="PTPRO"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47028">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE O"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47028:SF1">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE O"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26586">
        <property type="entry name" value="Fn3_R-PTP-O"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0472">Membrane</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0732">Signal</keyword>
    <keyword evidence="1" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="1" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="2">
        <location>
            <begin position="1"/>
            <end position="26"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019494044" description="Fibronectin type-III domain-containing protein" evidence="2">
        <location>
            <begin position="27"/>
            <end position="845"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="1">
        <location>
            <begin position="812"/>
            <end position="836"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="3">
        <location>
            <begin position="426"/>
            <end position="523"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="3">
        <location>
            <begin position="623"/>
            <end position="716"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="3">
        <location>
            <begin position="717"/>
            <end position="809"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015659.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="845" mass="94828" checksum="033A01689CE08CA7" modified="2019-05-08" version="1">MKSQIPPGFLLLLWLLLFSPFQVTVRDDNCIVVSLEESDVVSSSFVYVVKITGESKNYFFQFEEFNSTLPSPVVFNASYHGLYYIVTLMVVNSNMVSKPVRSITVLTKPLPISSVSIYDYKPSPETGVLFEIQYPEKYNVFTRVNISYWEGKDYRTMLYKDFFKGKTVFNHWLPGICYSNITFQLVSEATFNKSTLVEYSGVRHEPKQHRTVPYPPRNISVQIVHMNKNNWEEHSGNFLEESFMGPQDVIGQDKLIHFSNGPTEIPSANTSDAWPDYHYNSTDYTTTTQPYWWTNEADSSESEDEFVNVVPSEYENSSTASMSGKLTPETPSFLPVQMVLSWLPPKPPTAFDGFHINIEREGKISATLSEDTHEFIAELKEPGKYKLSVTTFSSSGSCEIRRSQSAKTLSFYISPTGEWIEELTEKPQHVSVTVLSSTTALMSWTSSQDSSSNNTIVSVVSLTCQKQKESQRLEKHYCTEVNSSNSIIENLVPGAQYQVVIYLRKGPLVGPPSDPVTFAIVPTGIKDLMLYPLGPTAVILSWNRPYLGVFRKYVVEMFYFNPSTMTSEWMTYYEIAATVSLTASVKIANLLPAWYYNFRVTMVTWGDPELSCCDSSTISYITAPVSPEITSVEYFNNLLYVSWTYGDNGTDLSHSRMLHWMIIAEGKKKIKKSVTRNVMTTVLSLPPGDIYNLSVTACTERGSNASTPQIVKLEPAPPKSLFAVNKTQTSVTLLWVEEGVADFFEVFCQQAGSGQETKLQEPVTVSSHVVTISSLLPATSYNCSVITFSHNSPSVPTFIAVSTMVTEMNPNIVVISVLAILSILLIGLLLVTLVVLRRKHLQMAR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HL55</accession>
    <name>A0A452HL55_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Receptor-type tyrosine-protein phosphatase O</fullName>
            <ecNumber evidence="2">3.1.3.48</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="17">Glomerular epithelial protein 1</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="18">Protein tyrosine phosphatase U2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Possesses tyrosine phosphatase activity. Plays a role in regulating the glomerular pressure/filtration rate relationship through an effect on podocyte structure and function.</text>
    </comment>
    <comment type="subunit">
        <text evidence="15">Interacts (phosphorylated form) with FYN and GRB2.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="13">Belongs to the protein-tyrosine phosphatase family. Receptor class 3 subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.48" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452HL55"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017894.1">
        <property type="protein sequence ID" value="ENSGAGP00000015666.1"/>
        <property type="gene ID" value="ENSGAGG00000011789.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000017895.1">
        <property type="protein sequence ID" value="ENSGAGP00000015667.1"/>
        <property type="gene ID" value="ENSGAGG00000011789.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098982">
        <property type="term" value="C:GABA-ergic synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098978">
        <property type="term" value="C:glutamatergic synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045296">
        <property type="term" value="F:cadherin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017147">
        <property type="term" value="F:Wnt-protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007411">
        <property type="term" value="P:axon guidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090090">
        <property type="term" value="P:negative regulation of canonical Wnt signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072112">
        <property type="term" value="P:podocyte differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003093">
        <property type="term" value="P:regulation of glomerular filtration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="CDD" id="cd14614">
        <property type="entry name" value="R-PTPc-O"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000488">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase O isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000039">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase O isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000415">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase O isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058859">
        <property type="entry name" value="Fn3_R-PTP-O"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000242">
        <property type="entry name" value="PTP_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042996">
        <property type="entry name" value="PTPRO"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016130">
        <property type="entry name" value="Tyr_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003595">
        <property type="entry name" value="Tyr_Pase_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47028">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE O"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47028:SF1">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE O"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26586">
        <property type="entry name" value="Fn3_R-PTP-O"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00102">
        <property type="entry name" value="Y_phosphatase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00700">
        <property type="entry name" value="PRTYPHPHTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00194">
        <property type="entry name" value="PTPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00404">
        <property type="entry name" value="PTPc_motif"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00383">
        <property type="entry name" value="TYR_PHOSPHATASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50055">
        <property type="entry name" value="TYR_PHOSPHATASE_PTP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="10 19" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="8" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="11" id="KW-0675">Receptor</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <keyword evidence="4 19" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9 19" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="726"/>
            <end position="750"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="786"/>
            <end position="807"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="234"/>
            <end position="330"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="340"/>
            <end position="437"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="537"/>
            <end position="630"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="631"/>
            <end position="723"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="20">
        <location>
            <begin position="844"/>
            <end position="1101"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="21">
        <location>
            <begin position="1018"/>
            <end position="1092"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00013064"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00025789"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00056314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00063538"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00073607"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00075446"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00080595"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015666.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1122" mass="127586" checksum="19631930D777E5EF" modified="2019-05-08" version="1">MVVNSNMVSKPVRSITVLTKPLPISSVSIYDYKPSPETGVLFEIQYPEKYNVFTRVNISYWEGKDYRTMLYKDFFKGKTVFNHWLPGICYSNITFQLVSEATFNKSTLVEYSGVRHEPKQHRTVPYPPRNISVQIVHMNKNNWEEHSGNFLEESFMGPQDVIGQDKLIHFSNGPTEIPSANTSDAWPDYHYNSTDYTTTTQPYWWTNEADSSESEDEFVNVVPSEYENSSTASMSGKLTPETPSFLPVQMVLSWLPPKPPTAFDGFHINIEREEKTTEFLTVGEDTHEFIAELKEPGKYKLSVTTFSSSGSCEIRRSQSAKTLSFYISPTGEWIEELTEKPQHVSVTVLSSTTALMSWTSSQDSSSNNTIVSVVSLTCQKQKESQRLEKHYCTEVNSSNSIIENLVPGAQYQVVIYLRKGPLVGPPSDPVTFAIVPTGIKDLMLYPLGPTAVILSWNRPYLGVFRKYVVEMFYFNPSTMTSEWMTYYEIAATVSLTASVKIANLLPAWYYNFRVTMVTWGDPELSCCDSSTISYITAPVSPEITSVEYFNNLLYVSWTYGDNGTDLSHSRMLHWMIIAEGKKKIKKSVTRNVMTTVLSLPPGDIYNLSVTACTERGSNASTPQIVKLEPAPPKSLFAVNKTQTSVTLLWVEEGVADFFEVFCQQAGSGQETKLQEPVTVSSHVVTISSLLPATSYNCSVITFSHNSPSVPTFIAVSTMVTEMNPNIVVISVLAILSILLIGLLLVTLVVLRRKHLQMARECGAGTFVNFASLERDGKLPYNWRRSVFAFLTLLPSCLWTDYLLAFYINPWSKNGLKKRKLTNPVQLDDFDGYIKDMAKDSDYKFSLQFEELKLIGLDIPHFAADLPMNRCKNRYTNILPYDFSRVRLVSMNEEEGSDYINANYIPGYNSPQEYIATQGPLPETRNDFWKMVLQQKSQIIVMLTQCNEKRRVKCDHYWPFTEEPVAYGDITVEMLSEEEQMDWSYRNFRISYADEVQDVMHFNYTAWPDHGVPTTNAAESILQFVQMVRQQSAKSKGPMVIHCSAGVGRTGTFIALDRLLQHIRDHEFVDILGLVSDMRSYRMSMVQTEEQYIFIHQCVQLMWLKKKQQFCISDVIYENVSKS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HL61</accession>
    <name>A0A452HL61_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Receptor-type tyrosine-protein phosphatase O</fullName>
            <ecNumber evidence="2">3.1.3.48</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="17">Glomerular epithelial protein 1</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="18">Protein tyrosine phosphatase U2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Possesses tyrosine phosphatase activity. Plays a role in regulating the glomerular pressure/filtration rate relationship through an effect on podocyte structure and function.</text>
    </comment>
    <comment type="subunit">
        <text evidence="15">Interacts (phosphorylated form) with FYN and GRB2.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="13">Belongs to the protein-tyrosine phosphatase family. Receptor class 3 subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.48" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452HL61"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017898.1">
        <property type="protein sequence ID" value="ENSGAGP00000015670.1"/>
        <property type="gene ID" value="ENSGAGG00000011789.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098982">
        <property type="term" value="C:GABA-ergic synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098978">
        <property type="term" value="C:glutamatergic synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045296">
        <property type="term" value="F:cadherin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017147">
        <property type="term" value="F:Wnt-protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007411">
        <property type="term" value="P:axon guidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090090">
        <property type="term" value="P:negative regulation of canonical Wnt signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072112">
        <property type="term" value="P:podocyte differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003093">
        <property type="term" value="P:regulation of glomerular filtration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="CDD" id="cd14614">
        <property type="entry name" value="R-PTPc-O"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000488">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase O isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000039">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase O isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000415">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase O isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058859">
        <property type="entry name" value="Fn3_R-PTP-O"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000242">
        <property type="entry name" value="PTP_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042996">
        <property type="entry name" value="PTPRO"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016130">
        <property type="entry name" value="Tyr_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003595">
        <property type="entry name" value="Tyr_Pase_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47028">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE O"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47028:SF1">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE O"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26586">
        <property type="entry name" value="Fn3_R-PTP-O"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00102">
        <property type="entry name" value="Y_phosphatase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00700">
        <property type="entry name" value="PRTYPHPHTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00194">
        <property type="entry name" value="PTPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00404">
        <property type="entry name" value="PTPc_motif"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00383">
        <property type="entry name" value="TYR_PHOSPHATASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50055">
        <property type="entry name" value="TYR_PHOSPHATASE_PTP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="10 19" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="8" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="11" id="KW-0675">Receptor</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <keyword evidence="4 19" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9 19" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="726"/>
            <end position="750"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="234"/>
            <end position="330"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="340"/>
            <end position="437"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="537"/>
            <end position="630"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="631"/>
            <end position="723"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="20">
        <location>
            <begin position="816"/>
            <end position="1073"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="21">
        <location>
            <begin position="990"/>
            <end position="1064"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00013064"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00025789"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00056314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00063538"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00073607"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00075446"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00080595"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015670.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1094" mass="124197" checksum="A11A3ABFB377B45A" modified="2019-05-08" version="1">MVVNSNMVSKPVRSITVLTKPLPISSVSIYDYKPSPETGVLFEIQYPEKYNVFTRVNISYWEGKDYRTMLYKDFFKGKTVFNHWLPGICYSNITFQLVSEATFNKSTLVEYSGVRHEPKQHRTVPYPPRNISVQIVHMNKNNWEEHSGNFLEESFMGPQDVIGQDKLIHFSNGPTEIPSANTSDAWPDYHYNSTDYTTTTQPYWWTNEADSSESEDEFVNVVPSEYENSSTASMSGKLTPETPSFLPVQMVLSWLPPKPPTAFDGFHINIEREEKTTEFLTVGEDTHEFIAELKEPGKYKLSVTTFSSSGSCEIRRSQSAKTLSFYISPTGEWIEELTEKPQHVSVTVLSSTTALMSWTSSQDSSSNNTIVSVVSLTCQKQKESQRLEKHYCTEVNSSNSIIENLVPGAQYQVVIYLRKGPLVGPPSDPVTFAIVPTGIKDLMLYPLGPTAVILSWNRPYLGVFRKYVVEMFYFNPSTMTSEWMTYYEIAATVSLTASVKIANLLPAWYYNFRVTMVTWGDPELSCCDSSTISYITAPVSPEITSVEYFNNLLYVSWTYGDNGTDLSHSRMLHWMIIAEGKKKIKKSVTRNVMTTVLSLPPGDIYNLSVTACTERGSNASTPQIVKLEPAPPKSLFAVNKTQTSVTLLWVEEGVADFFEVFCQQAGSGQETKLQEPVTVSSHVVTISSLLPATSYNCSVITFSHNSPSVPTFIAVSTMVTEMNPNIVVISVLAILSILLIGLLLVTLVVLRRKHLQMARECGAGTFVNFASLERDGKLPYNWSKNGLKKRKLTNPVQLDDFDGYIKDMAKDSDYKFSLQFEELKLIGLDIPHFAADLPMNRCKNRYTNILPYDFSRVRLVSMNEEEGSDYINANYIPGYNSPQEYIATQGPLPETRNDFWKMVLQQKSQIIVMLTQCNEKRRVKCDHYWPFTEEPVAYGDITVEMLSEEEQMDWSYRNFRISYADEVQDVMHFNYTAWPDHGVPTTNAAESILQFVQMVRQQSAKSKGPMVIHCSAGVGRTGTFIALDRLLQHIRDHEFVDILGLVSDMRSYRMSMVQTEEQYIFIHQCVQLMWLKKKQQFCISDVIYENVSKS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HL62</accession>
    <name>A0A452HL62_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Receptor-type tyrosine-protein phosphatase O</fullName>
            <ecNumber evidence="2">3.1.3.48</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="17">Glomerular epithelial protein 1</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="18">Protein tyrosine phosphatase U2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Possesses tyrosine phosphatase activity. Plays a role in regulating the glomerular pressure/filtration rate relationship through an effect on podocyte structure and function.</text>
    </comment>
    <comment type="subunit">
        <text evidence="15">Interacts (phosphorylated form) with FYN and GRB2.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="13">Belongs to the protein-tyrosine phosphatase family. Receptor class 3 subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.48" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452HL62"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017901.1">
        <property type="protein sequence ID" value="ENSGAGP00000015672.1"/>
        <property type="gene ID" value="ENSGAGG00000011789.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098982">
        <property type="term" value="C:GABA-ergic synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098978">
        <property type="term" value="C:glutamatergic synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045296">
        <property type="term" value="F:cadherin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017147">
        <property type="term" value="F:Wnt-protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007411">
        <property type="term" value="P:axon guidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090090">
        <property type="term" value="P:negative regulation of canonical Wnt signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072112">
        <property type="term" value="P:podocyte differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003093">
        <property type="term" value="P:regulation of glomerular filtration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="CDD" id="cd14614">
        <property type="entry name" value="R-PTPc-O"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000488">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase O isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000039">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase O isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000415">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase O isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058859">
        <property type="entry name" value="Fn3_R-PTP-O"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000242">
        <property type="entry name" value="PTP_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042996">
        <property type="entry name" value="PTPRO"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016130">
        <property type="entry name" value="Tyr_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003595">
        <property type="entry name" value="Tyr_Pase_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47028">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE O"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47028:SF1">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE O"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26586">
        <property type="entry name" value="Fn3_R-PTP-O"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00102">
        <property type="entry name" value="Y_phosphatase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00700">
        <property type="entry name" value="PRTYPHPHTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00194">
        <property type="entry name" value="PTPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00404">
        <property type="entry name" value="PTPc_motif"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00383">
        <property type="entry name" value="TYR_PHOSPHATASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50055">
        <property type="entry name" value="TYR_PHOSPHATASE_PTP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="10 19" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="8" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="11" id="KW-0675">Receptor</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <keyword evidence="4 19" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9 19" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="672"/>
            <end position="696"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="732"/>
            <end position="753"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="286"/>
            <end position="383"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="483"/>
            <end position="576"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="577"/>
            <end position="669"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="20">
        <location>
            <begin position="790"/>
            <end position="1047"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="21">
        <location>
            <begin position="964"/>
            <end position="1038"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00013064"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00025789"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00056314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00063538"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00073607"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00075446"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00080595"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015672.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1068" mass="121495" checksum="C2609378C6809B89" modified="2019-05-08" version="1">MVVNSNMVSKPVRSITVLTKPLPISSVSIYDYKPSPETGVLFEIQYPEKYNVFTRVNISYWEGKDYRTMLYKDFFKGKTVFNHWLPGICYSNITFQLVSEATFNKSTLVEYSGVRHEPKQHRTVPYPPRNISVQIVHMNKNNWEEHSGNFLEESFMGPQDVIGQDKLIHFSNGPTEIPSANTSDAWPDYHYNSTDYTTTTQPYWWTNEADSSESEDEFVNVVPSEYENSSTASMSGKLTPETPSFLPVQMVLSWLPPKPPTAFDGFHINIEREGPTGEWIEELTEKPQHVSVTVLSSTTALMSWTSSQDSSSNNTIVSVVSLTCQKQKESQRLEKHYCTEVNSSNSIIENLVPGAQYQVVIYLRKGPLVGPPSDPVTFAIVPTGIKDLMLYPLGPTAVILSWNRPYLGVFRKYVVEMFYFNPSTMTSEWMTYYEIAATVSLTASVKIANLLPAWYYNFRVTMVTWGDPELSCCDSSTISYITAPVSPEITSVEYFNNLLYVSWTYGDNGTDLSHSRMLHWMIIAEGKKKIKKSVTRNVMTTVLSLPPGDIYNLSVTACTERGSNASTPQIVKLEPAPPKSLFAVNKTQTSVTLLWVEEGVADFFEVFCQQAGSGQETKLQEPVTVSSHVVTISSLLPATSYNCSVITFSHNSPSVPTFIAVSTMVTEMNPNIVVISVLAILSILLIGLLLVTLVVLRRKHLQMARECGAGTFVNFASLERDGKLPYNWRRSVFAFLTLLPSCLWTDYLLAFYINPWSKNGLKKRKLTNPVQLDDFDGYIKDMAKDSDYKFSLQFEELKLIGLDIPHFAADLPMNRCKNRYTNILPYDFSRVRLVSMNEEEGSDYINANYIPGYNSPQEYIATQGPLPETRNDFWKMVLQQKSQIIVMLTQCNEKRRVKCDHYWPFTEEPVAYGDITVEMLSEEEQMDWSYRNFRISYADEVQDVMHFNYTAWPDHGVPTTNAAESILQFVQMVRQQSAKSKGPMVIHCSAGVGRTGTFIALDRLLQHIRDHEFVDILGLVSDMRSYRMSMVQTEEQYIFIHQCVQLMWLKKKQQFCISDVIYENVSKS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HL86</accession>
    <name>A0A452HL86_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">CMP/dCMP-type deaminase domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the cytidine and deoxycytidylate deaminase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HL86"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017928.1">
        <property type="protein sequence ID" value="ENSGAGP00000015699.1"/>
        <property type="gene ID" value="ENSGAGG00000011837.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004126">
        <property type="term" value="F:cytidine deaminase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016554">
        <property type="term" value="P:cytidine to uridine editing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd01283">
        <property type="entry name" value="cytidine_deaminase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.140.10">
        <property type="entry name" value="Cytidine Deaminase, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041547">
        <property type="entry name" value="APOBEC1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050610">
        <property type="entry name" value="APOBEC_Cyt_Deaminase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002125">
        <property type="entry name" value="CMP_dCMP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016193">
        <property type="entry name" value="Cytidine_deaminase-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13857:SF26">
        <property type="entry name" value="C-U-EDITING ENZYME APOBEC-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13857">
        <property type="entry name" value="MRNA EDITING ENZYME"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18774">
        <property type="entry name" value="APOBEC4_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53927">
        <property type="entry name" value="Cytidine deaminase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51747">
        <property type="entry name" value="CYT_DCMP_DEAMINASES_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="6" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="CMP/dCMP-type deaminase" evidence="9">
        <location>
            <begin position="5"/>
            <end position="130"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00006576"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51747"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015699.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="239" mass="28540" checksum="C7A590E861278227" modified="2019-05-08" version="1">CTGHKGPKARWQIQPKEFRAIFDPSAFPKVTYLLYEIKWGSSTKFWRKWCLNTPTQHAEINCLENDFKAIHHGPSVHCSIIWFLSWSPCGLCCRLIIHFLRAHPKVTLQIYIGRLFRHRDERNRQGLRDLVSSGVNVYIMSLPAYDYCWRTFVDHRVREDDIYWPCYFTPWMIFYMLEFQCILQGLPPCLQIWLKNYTGSPVFQLVLQESHRKILWQSLHTANSPQQRLIFRPRPSTEH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HL88</accession>
    <name>A0A452HL88_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="22">Tyrosine-protein kinase Mer</fullName>
            <ecNumber evidence="3">2.7.10.1</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="23">Proto-oncogene c-Mer</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="24">Receptor tyrosine kinase MerTK</fullName>
        </alternativeName>
    </protein>
    <organism evidence="30 31">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="31" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="30" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="30" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="20">
            <text>L-tyrosyl-[protein] + ATP = O-phospho-L-tyrosyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10596"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.10.1"/>
        </reaction>
    </comment>
    <comment type="subunit">
        <text evidence="21">Interacts (upon activation) with TNK2; stimulates TNK2 autophosphorylation. Interacts (via N-terminus) with extracellular ligands LGALS3, TUB, TULP1 and GAS6. Interacts with VAV1 in a phosphotyrosine-independent manner. Interacts with TIMD4; this interaction enhances TIMD4-mediated efferocytosis.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the protein kinase superfamily. CAMK Ser/Thr protein kinase family.</text>
    </comment>
    <dbReference type="EC" id="2.7.10.1" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452HL88"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017946.1">
        <property type="protein sequence ID" value="ENSGAGP00000015717.1"/>
        <property type="gene ID" value="ENSGAGG00000010943.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043235">
        <property type="term" value="C:receptor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004714">
        <property type="term" value="F:transmembrane receptor protein tyrosine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007169">
        <property type="term" value="P:cell surface receptor protein tyrosine kinase signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006909">
        <property type="term" value="P:phagocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000834">
        <property type="entry name" value="Proto-oncogene tyrosine-protein kinase MER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000089">
        <property type="entry name" value="Tyrosine-protein kinase receptor TYRO3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000296">
        <property type="entry name" value="Tyrosine-protein kinase receptor TYRO3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000111">
        <property type="entry name" value="Tyrosine-protein kinase receptor TYRO3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050122">
        <property type="entry name" value="RTK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001245">
        <property type="entry name" value="Ser-Thr/Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008266">
        <property type="entry name" value="Tyr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020635">
        <property type="entry name" value="Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416:SF257">
        <property type="entry name" value="TYROSINE-PROTEIN KINASE MER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416">
        <property type="entry name" value="TYROSINE-PROTEIN KINASE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13927">
        <property type="entry name" value="Ig_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07714">
        <property type="entry name" value="PK_Tyr_Ser-Thr"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00109">
        <property type="entry name" value="TYRKINASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00219">
        <property type="entry name" value="TyrKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00109">
        <property type="entry name" value="PROTEIN_KINASE_TYR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12 25" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="16" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="18" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="19" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="11" id="KW-0418">Kinase</keyword>
    <keyword evidence="14 26" id="KW-0472">Membrane</keyword>
    <keyword evidence="10 25" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="17" id="KW-0675">Receptor</keyword>
    <keyword evidence="31" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0677">Repeat</keyword>
    <keyword evidence="8" id="KW-0732">Signal</keyword>
    <keyword evidence="6" id="KW-0808">Transferase</keyword>
    <keyword evidence="7 26" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="13 26" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="15" id="KW-0829">Tyrosine-protein kinase</keyword>
    <feature type="transmembrane region" description="Helical" evidence="26">
        <location>
            <begin position="451"/>
            <end position="474"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="28">
        <location>
            <begin position="164"/>
            <end position="246"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="29">
        <location>
            <begin position="253"/>
            <end position="348"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="27">
        <location>
            <begin position="536"/>
            <end position="807"/>
        </location>
    </feature>
    <feature type="binding site" evidence="25">
        <location>
            <position position="568"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00011902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023137"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00051243"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00066004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00070212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00077110"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00082356"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="27">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="28">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="29">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="30">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015717.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="31">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="942" mass="105344" checksum="6EE6A7876D8AE186" modified="2019-05-08" version="1">SADCENTEARQERRTSHLLPSTQAFSVDWLAAPRRPRSLQTTTHDTPKANARTTPVELYLDQANASLLDEASAAHSLQQVKFNPTVGHILINEHKDFKFNCSIKGPQFLFNQDSAVISLWKDGKELQEPVWEAGANAHDACAESRRCIVFLRCLVSVPFCVGLPHFLRQPQSLNITRNRPFNLTCEAMGPPEPVQIYWFRNSSRVNDQPYISPSVLTLPGLSETAIFSCEAHNRKGLTASNEVQINIKRTPAAPADVRVLNRTAHSLVVSWVPGFDGYSPLHSCRLQVREAAALNKGSFLIWNTSVPPHRYHVQQLQPLAEYSIRVSCRNEVGWSSFSPWTLASTTEGAPSTPPLNVTVFFNESSSSLEVRWAKPHLGRKHGELQGYRLLHTWQNSKITVITNATYAVRVAAVTKGGIGPFSDPVVVFIPGDGFMASAPSSTPAPGNNHSFAIVLGFILGIVAIGLILYFSVIVRKRWMETKFGTAFHSDDSEHVVSYTAKKSYSRRAVELTLASLGVREELQQKLQDVVIDRDSLALGKILGEGEFGSVMEGHLSQPSGTVQKVAVKTMKLEDFSQREIEEFLSEAACMKDFDHPNVIRLLGVCIELSSRQVPKPMVVLPFMKYGDLHSFLLRSRFETGTRVRNSATLLKFMVDIALGMEYLSTRQFLHRDLAARNCMLRDDMTVCVADFGLSKKIYSGDYYRQGRIAKMPVKWIAIESLADRVYTTKSDVWAFGVTMWEIATRGMTPYPGVQNHEIYDYLLHGHRLKQPEDCLDELYEVMYSCWRADPHERPAFPELKLHLEKLLESLPKVSGSEDVIYINTSLPEKSEELTEESEFPQTDMAFNRNYVTVSSSPKPDTTVVTAEVHESDQREERYILKGETDEQTAGSLTLEQGHAPLLQGTITRNGATWGPASMLLGNSLADELLYADDCSEDSEVLL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HL92</accession>
    <name>A0A452HL92_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Activation-induced cytidine deaminase AID domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HL92"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017932.1">
        <property type="protein sequence ID" value="ENSGAGP00000015703.1"/>
        <property type="gene ID" value="ENSGAGG00000011838.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004126">
        <property type="term" value="F:cytidine deaminase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016554">
        <property type="term" value="P:cytidine to uridine editing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.140.10">
        <property type="entry name" value="Cytidine Deaminase, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013158">
        <property type="entry name" value="AID"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050610">
        <property type="entry name" value="APOBEC_Cyt_Deaminase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13857">
        <property type="entry name" value="MRNA EDITING ENZYME"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13857:SF10">
        <property type="entry name" value="SINGLE-STRANDED DNA CYTOSINE DEAMINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08210">
        <property type="entry name" value="APOBEC_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="1" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Activation-induced cytidine deaminase AID" evidence="3">
        <location>
            <begin position="13"/>
            <end position="55"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="Pfam" id="PF08210"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015703.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="71" mass="8496" checksum="A545BC839D84EB57" modified="2019-05-08" version="1">MFPCSLLMKQKKFLYNFKNLRWARGRHETYLCYVVKRRDSATSCSLDFGYLRNKPLDEVDDLRDAFKILGL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HL93</accession>
    <name>A0A452HL93_9SAUR</name>
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        <submittedName>
            <fullName evidence="16">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
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    <dbReference type="Ensembl" id="ENSGAGT00000017951.1">
        <property type="protein sequence ID" value="ENSGAGP00000015722.1"/>
        <property type="gene ID" value="ENSGAGG00000011232.1"/>
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        <property type="component" value="Unassembled WGS sequence"/>
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        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
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        <property type="term" value="F:chromatin binding"/>
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        <property type="project" value="InterPro"/>
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        <property type="term" value="F:sequence-specific DNA binding"/>
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        <property type="project" value="InterPro"/>
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        <property type="term" value="F:transcription corepressor activity"/>
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        <property type="project" value="TreeGrafter"/>
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        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
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        <property type="entry name" value="SANT_MTA3_like"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="ZnF_GATA"/>
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        <property type="entry name" value="Arginine-glutamic acid dipeptide (RE) repeats"/>
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        <property type="entry name" value="Arginine-glutamic acid dipeptide (RE) repeats"/>
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    <dbReference type="InterPro" id="IPR000679">
        <property type="entry name" value="Znf_GATA"/>
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    <dbReference type="PANTHER" id="PTHR13859:SF12">
        <property type="entry name" value="ARGININE-GLUTAMIC ACID DIPEPTIDE REPEATS PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13859">
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        <property type="match status" value="1"/>
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    </dbReference>
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        <property type="entry name" value="ZnF_GATA"/>
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    </dbReference>
    <dbReference type="SUPFAM" id="SSF57716">
        <property type="entry name" value="Glucocorticoid receptor-like (DNA-binding domain)"/>
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    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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    <keyword evidence="8" id="KW-0007">Acetylation</keyword>
    <keyword evidence="2" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0804">Transcription</keyword>
    <keyword evidence="9" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="7" id="KW-0832">Ubl conjugation</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5" id="KW-0863">Zinc-finger</keyword>
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        <location>
            <begin position="103"/>
            <end position="241"/>
        </location>
    </feature>
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        <location>
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            <end position="297"/>
        </location>
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        </location>
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        </location>
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    </evidence>
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    </evidence>
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</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HL97</accession>
    <name>A0A452HL97_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="10">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the MFAP family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HL97"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017937.1">
        <property type="protein sequence ID" value="ENSGAGP00000015708.1"/>
        <property type="gene ID" value="ENSGAGG00000011840.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001527">
        <property type="term" value="C:microfibril"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048048">
        <property type="term" value="P:embryonic eye morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008673">
        <property type="entry name" value="MAGP"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16485">
        <property type="entry name" value="MICROFIBRILLAR-ASSOCIATED PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16485:SF6">
        <property type="entry name" value="MICROFIBRILLAR-ASSOCIATED PROTEIN 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05507">
        <property type="entry name" value="MAGP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="4" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="7" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0964">Secreted</keyword>
    <keyword evidence="5 9" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="9">
        <location>
            <begin position="1"/>
            <end position="19"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019578314" evidence="9">
        <location>
            <begin position="20"/>
            <end position="140"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="114"/>
            <end position="140"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005317"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015708.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="140" mass="15980" checksum="B6A5C3030DCC18B5" modified="2019-05-08" version="1">MMTFGARALLCIIALSVLPADWFLLMVNCQEVPTTPAAIVNTTEIPECREEQFPCTRLYSVHKPVKQCISYLCVTSVRRMYIINKEVCSRVVCKEDEVMQDEICRQLAGLPPRRLRRSGQPRGPPCRWGQNKELNKPDAL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HL98</accession>
    <name>A0A452HL98_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">C2 DOCK-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the DOCK family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HL98"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017957.1">
        <property type="protein sequence ID" value="ENSGAGP00000015727.1"/>
        <property type="gene ID" value="ENSGAGG00000011372.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031267">
        <property type="term" value="F:small GTPase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007520">
        <property type="term" value="P:myoblast fusion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007264">
        <property type="term" value="P:small GTPase-mediated signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd08694">
        <property type="entry name" value="C2_Dock-A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.350">
        <property type="entry name" value="Dedicator of cytokinesis N-terminal subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027007">
        <property type="entry name" value="C2_DOCK-type_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026791">
        <property type="entry name" value="DOCK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047026">
        <property type="entry name" value="DOCK1_C2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032376">
        <property type="entry name" value="DOCK_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042455">
        <property type="entry name" value="DOCK_N_sub1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056372">
        <property type="entry name" value="TPR_DOCK"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45653">
        <property type="entry name" value="DEDICATOR OF CYTOKINESIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45653:SF1">
        <property type="entry name" value="DEDICATOR OF CYTOKINESIS PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14429">
        <property type="entry name" value="DOCK-C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16172">
        <property type="entry name" value="DOCK_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23554">
        <property type="entry name" value="TPR_DOCK"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51650">
        <property type="entry name" value="C2_DOCK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="C2 DOCK-type" evidence="4">
        <location>
            <begin position="368"/>
            <end position="552"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00983"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS51650"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015727.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="921" mass="105953" checksum="13AAAB802687B2CF" modified="2025-10-08" version="2">MDYINHFTILPKFVFANFRQHETVIPSELPLIQEVTTTLREWSIIWRQLYVVSKIHMFHSVRHMIYDLIEWRSQILSGTLPQDELKELKKKVTAKIDYGNRILDLDLVVRDEDGNILDPEQTSTISLFRAHEIASKQVEERLLEEKSQKQNIDINRQAKFAATPSFALFVNLKNVVCKIGEDAEVLMSLYDPLESKFISENYLVRWSSSGLPKDIDRLHNLRAVFTDLGSKDLKREKISFVCQIVRVGRMELRDNNTRKLTSGLRRPFGVAVMDVTDIINGKVDDEDKQHFIPFQPVAGENDFLQTVINKVIAAKEVNHKGQGLWVTLKLLPGDIHQIRKEFPHLVDRSTAVARKMGFPEIIMPGDVRNDIYVTLVQGDFDKGSKTTAKNVEVTVSVYDEDGKRLENVIFPGAGDDAISEYKSVIYYQVKQPRWFETVKVAIPIEDVNRSHLRFTFRHRSSQDSKDKSEKIFALAFVKLMRYDGTTLRDGEHDLIVYKAEAKKLEDASTYLSLPSTKIELEEKGHATTGKSMQNLGSCTISKDSFQISTLVCSTKLTQNVDLLGLLKWRSNTNLLQQNLRQLMKVDGGEVVKFLQDTLDALFNIMMENSESETFDTLVFDALVFIIGLIADRKFQHFNPVLETYIKKHFSATLAYTKLTKVLRNYVDNAEKPGVTDQLFKAMKALEYIFKFIVRSRILFNQLYENKGEADFMESLLQLFKSINEMMSSVSDQTVMVKVGVSKKNVLALSNNYAVDTIILSLQGPTQRHVQIIMEKLLRTVNRTVISMGRDSELIVSGNFVASMTAILRQMEDYHYAHLIKTFGKMRSDVVDFLMETFIMFKNLIGKNVYPFDWVIMNMMQNKFIIRRDLYGSLHRICYPTVLAEILKISLPPFIFLGAPSFSEELRQHWKEFYLQPIGPTY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HL99</accession>
    <name>A0A452HL99_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="22">Tyrosine-protein kinase Mer</fullName>
            <ecNumber evidence="3">2.7.10.1</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="23">Proto-oncogene c-Mer</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="24">Receptor tyrosine kinase MerTK</fullName>
        </alternativeName>
    </protein>
    <organism evidence="33 34">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="34" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="33" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="20">
            <text>L-tyrosyl-[protein] + ATP = O-phospho-L-tyrosyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10596"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.10.1"/>
        </reaction>
    </comment>
    <comment type="subunit">
        <text evidence="21">Interacts (upon activation) with TNK2; stimulates TNK2 autophosphorylation. Interacts (via N-terminus) with extracellular ligands LGALS3, TUB, TULP1 and GAS6. Interacts with VAV1 in a phosphotyrosine-independent manner. Interacts with TIMD4; this interaction enhances TIMD4-mediated efferocytosis.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the protein kinase superfamily. CAMK Ser/Thr protein kinase family.</text>
    </comment>
    <dbReference type="EC" id="2.7.10.1" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452HL99"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017929.1">
        <property type="protein sequence ID" value="ENSGAGP00000015700.1"/>
        <property type="gene ID" value="ENSGAGG00000010943.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000017941.1">
        <property type="protein sequence ID" value="ENSGAGP00000015712.1"/>
        <property type="gene ID" value="ENSGAGG00000010943.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043235">
        <property type="term" value="C:receptor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004714">
        <property type="term" value="F:transmembrane receptor protein tyrosine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007169">
        <property type="term" value="P:cell surface receptor protein tyrosine kinase signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006909">
        <property type="term" value="P:phagocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:001050">
        <property type="entry name" value="MER proto-oncogene, tyrosine kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000834">
        <property type="entry name" value="Proto-oncogene tyrosine-protein kinase MER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000089">
        <property type="entry name" value="Tyrosine-protein kinase receptor TYRO3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000296">
        <property type="entry name" value="Tyrosine-protein kinase receptor TYRO3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000111">
        <property type="entry name" value="Tyrosine-protein kinase receptor TYRO3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050122">
        <property type="entry name" value="RTK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001245">
        <property type="entry name" value="Ser-Thr/Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008266">
        <property type="entry name" value="Tyr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020635">
        <property type="entry name" value="Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416:SF257">
        <property type="entry name" value="TYROSINE-PROTEIN KINASE MER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416">
        <property type="entry name" value="TYROSINE-PROTEIN KINASE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13927">
        <property type="entry name" value="Ig_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07714">
        <property type="entry name" value="PK_Tyr_Ser-Thr"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000615">
        <property type="entry name" value="TyrPK_CSF1-R"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00109">
        <property type="entry name" value="TYRKINASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00219">
        <property type="entry name" value="TyrKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00109">
        <property type="entry name" value="PROTEIN_KINASE_TYR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12 26 28" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="16" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="18" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="19" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="11" id="KW-0418">Kinase</keyword>
    <keyword evidence="27" id="KW-0460">Magnesium</keyword>
    <keyword evidence="14 29" id="KW-0472">Membrane</keyword>
    <keyword evidence="27" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10 26 28" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="17" id="KW-0675">Receptor</keyword>
    <keyword evidence="34" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0677">Repeat</keyword>
    <keyword evidence="8" id="KW-0732">Signal</keyword>
    <keyword evidence="6" id="KW-0808">Transferase</keyword>
    <keyword evidence="7 29" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="13 29" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="15" id="KW-0829">Tyrosine-protein kinase</keyword>
    <feature type="transmembrane region" description="Helical" evidence="29">
        <location>
            <begin position="362"/>
            <end position="385"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="31">
        <location>
            <begin position="57"/>
            <end position="139"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="32">
        <location>
            <begin position="146"/>
            <end position="241"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="32">
        <location>
            <begin position="246"/>
            <end position="343"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="30">
        <location>
            <begin position="447"/>
            <end position="718"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="25">
        <location>
            <position position="583"/>
        </location>
    </feature>
    <feature type="binding site" evidence="28">
        <location>
            <position position="479"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="26">
        <location>
            <position position="587"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="27">
        <location>
            <position position="588"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="27">
        <location>
            <position position="601"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00011902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023137"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00051243"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00066004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00070212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00077110"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00082356"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="PIRSR" id="PIRSR000615-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="PIRSR" id="PIRSR000615-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="PIRSR" id="PIRSR000615-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="30">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="31">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="32">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="33">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015712.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="34">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="853" mass="95501" checksum="237A4694DF4D606C" modified="2019-05-08" version="1">MSTRHYQVTDDVETTMVSTFSITNAQRSDNGSYSCKLKIHNEEIESDPIFVQMEGLPHFLRQPQSLNITRNRPFNLTCEAMGPPEPVQIYWFRNSSRVNDQPYISPSVLTLPGLSETAIFSCEAHNRKGLTASNEVQINIKRTPAAPADVRVLNRTAHSLVVSWVPGFDGYSPLHSCRLQVREAAALNKGSFLIWNTSVPPHRYHVQQLQPLAEYSIRVSCRNEVGWSSFSPWTLASTTEGAPSTPPLNVTVFFNESSSSLEVRWAKPHLGRKHGELQGYRLLHTWQNSKITQNRSSEAQQNATVTTVAIVVTNATYAVRVAAVTKGGIGPFSDPVVVFIPGDGFMASAPSSTPAPGNNHSFAIVLGFILGIVAIGLILYFSVIVRKRWMETKFGTAFHSDDSEHVVSYTAKKSYSRRAVELTLASLGVREELQQKLQDVVIDRDSLALGKILGEGEFGSVMEGHLSQPSGTVQKVAVKTMKLEDFSQREIEEFLSEAACMKDFDHPNVIRLLGVCIELSSRQVPKPMVVLPFMKYGDLHSFLLRSRFETGTRHVPLQTLLKFMVDIALGMEYLSTRQFLHRDLAARNCMLRDDMTVCVADFGLSKKIYSGDYYRQGRIAKMPVKWIAIESLADRVYTTKSDVWAFGVTMWEIATRGMTPYPGVQNHEIYDYLLHGHRLKQPEDCLDELYEVMYSCWRADPHERPAFPELKLHLEKLLESLPKVSGSEDVIYINTSLPEKSEELTEESEFPQTDMAFNRNYVTVSSSPKPDTTVVTAEVHESDQREERYILKGETDEQTAGSLTLEQGHAPLLQGTITRNGATWGPASMLLGNSLADELLYADDCSEDSEVLL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INW3</accession>
    <name>A0A452INW3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="22">Cell division cycle 14B</fullName>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>O-phospho-L-seryl-[protein] + H2O = L-seryl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:20629"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="EC" id="3.1.3.16"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="16">
            <text>O-phospho-L-threonyl-[protein] + H2O = L-threonyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:47004"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="EC" id="3.1.3.16"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="14">Cell projection</location>
            <location evidence="14">Kinocilium</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
            <location evidence="2">Microtubule organizing center</location>
            <location evidence="2">Centrosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Cytoskeleton</location>
            <location evidence="3">Spindle pole</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the protein-tyrosine phosphatase family. Non-receptor class CDC14 subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452INW3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033576.1">
        <property type="protein sequence ID" value="ENSGAGP00000029568.1"/>
        <property type="gene ID" value="ENSGAGG00000021351.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060091">
        <property type="term" value="C:kinocilium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000922">
        <property type="term" value="C:spindle pole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004722">
        <property type="term" value="F:protein serine/threonine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007605">
        <property type="term" value="P:sensory perception of sound"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd14499">
        <property type="entry name" value="CDC14_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd17657">
        <property type="entry name" value="CDC14_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000032">
        <property type="entry name" value="dual specificity protein phosphatase CDC14A isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000006">
        <property type="entry name" value="Dual specificity protein phosphatase CDC14B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044506">
        <property type="entry name" value="CDC14_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029260">
        <property type="entry name" value="DSPn"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000340">
        <property type="entry name" value="Dual-sp_phosphatase_cat-dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050561">
        <property type="entry name" value="PTP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016130">
        <property type="entry name" value="Tyr_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020422">
        <property type="entry name" value="TYR_PHOSPHATASE_DUAL_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23339">
        <property type="entry name" value="TYROSINE SPECIFIC PROTEIN PHOSPHATASE AND DUAL SPECIFICITY PROTEIN PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00782">
        <property type="entry name" value="DSPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14671">
        <property type="entry name" value="DSPn"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00195">
        <property type="entry name" value="DSPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00383">
        <property type="entry name" value="TYR_PHOSPHATASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50054">
        <property type="entry name" value="TYR_PHOSPHATASE_DUAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="13" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="7" id="KW-0132">Cell division</keyword>
    <keyword evidence="12" id="KW-0966">Cell projection</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="10" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="9" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="18">
        <location>
            <begin position="178"/>
            <end position="337"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="19">
        <location>
            <begin position="260"/>
            <end position="322"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="418"/>
            <end position="459"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="418"/>
            <end position="431"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="17">
        <location>
            <begin position="436"/>
            <end position="446"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004647"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00007315"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00037822"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00047761"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00048336"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029568.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="22">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="483" mass="55178" checksum="CF567EA2C0592784" modified="2019-05-08" version="1">MSVRKESRPSVEFVKGRLYFAILCQKPKSGVANTHYFCIDDELVYENFYADFGPLNLAMVYRYCCKLNKKLKSFTLIRKKIIHYTGFDQKKQANAAFLIGSYAIIYLKESPEDVYRLLLAGNASYLPFRDASFGTCSFHLTLLDCFHAINKALRYGFLDFSAFDVNEYEHYEKAENGDFNWIIPNKFIAFSGPHSRSKTENGYPHHAPEAYFPYFRRHKVTTIIRLNRKMYDAKRFADAGFEHYDLFFADGSIPSDTIVKTFLNICENAEGVIAVHCKAGLGRTGTLIACYIMKHYRMTAAETIAWIRINRPGSVIGPQQHFLVEKQTELWREGDVFRAKLKENRKIAVTKILTGVDDISINDAKNRRANRKDTELYSDEDDANCVTQGDKLRALKSKRQAKAAATVPLTVILQSSVQKSKTSEPSISGSADINKRTTRSAARKSSLKSLIPQKEKRKRNALQQMRLMKLCGHSIPRTRTVLR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INW4</accession>
    <name>A0A452INW4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1 10">Catechol O-methyltransferase</fullName>
            <ecNumber evidence="1 10">2.1.1.6</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">Catalyzes the O-methylation, and thereby the inactivation, of catecholamine neurotransmitters and catechol hormones.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="10">
            <text>a catechol + S-adenosyl-L-methionine = a guaiacol + S-adenosyl-L-homocysteine + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17877"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:33566"/>
            <dbReference type="ChEBI" id="CHEBI:57856"/>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
            <dbReference type="ChEBI" id="CHEBI:134251"/>
            <dbReference type="EC" id="2.1.1.6"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="10 13">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
        <text evidence="10 13">Binds 1 Mg(2+) ion per subunit.</text>
    </comment>
    <comment type="similarity">
        <text evidence="9 10">Belongs to the class I-like SAM-binding methyltransferase superfamily. Cation-dependent O-methyltransferase family.</text>
    </comment>
    <dbReference type="EC" id="2.1.1.6" evidence="1 10"/>
    <dbReference type="AlphaFoldDB" id="A0A452INW4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033678.1">
        <property type="protein sequence ID" value="ENSGAGP00000029662.1"/>
        <property type="gene ID" value="ENSGAGG00000021411.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030424">
        <property type="term" value="C:axon"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030425">
        <property type="term" value="C:dendrite"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016206">
        <property type="term" value="F:catechol O-methyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000287">
        <property type="term" value="F:magnesium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042424">
        <property type="term" value="P:catecholamine catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032502">
        <property type="term" value="P:developmental process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042417">
        <property type="term" value="P:dopamine metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032259">
        <property type="term" value="P:methylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="CDD" id="cd02440">
        <property type="entry name" value="AdoMet_MTases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.150:FF:000054">
        <property type="entry name" value="Catechol O-methyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.150">
        <property type="entry name" value="Vaccinia Virus protein VP39"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017128">
        <property type="entry name" value="Catechol_O-MeTrfase_euk"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029063">
        <property type="entry name" value="SAM-dependent_MTases_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002935">
        <property type="entry name" value="SAM_O-MeTrfase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43836:SF3">
        <property type="entry name" value="CATECHOL O-METHYLTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43836">
        <property type="entry name" value="CATECHOL O-METHYLTRANSFERASE 1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01596">
        <property type="entry name" value="Methyltransf_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF037177">
        <property type="entry name" value="Catechol_O-mtfrase_euk"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53335">
        <property type="entry name" value="S-adenosyl-L-methionine-dependent methyltransferases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51682">
        <property type="entry name" value="SAM_OMT_I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 10" id="KW-0128">Catecholamine metabolism</keyword>
    <keyword evidence="6 10 13" id="KW-0460">Magnesium</keyword>
    <keyword evidence="5 10 13" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="2 10" id="KW-0489">Methyltransferase</keyword>
    <keyword evidence="7 10" id="KW-0531">Neurotransmitter degradation</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 10 11" id="KW-0949">S-adenosyl-L-methionine</keyword>
    <keyword evidence="3 10" id="KW-0808">Transferase</keyword>
    <feature type="binding site" evidence="11">
        <location>
            <position position="90"/>
        </location>
        <ligand>
            <name>S-adenosyl-L-methionine</name>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="11">
        <location>
            <position position="120"/>
        </location>
        <ligand>
            <name>S-adenosyl-L-methionine</name>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="11">
        <location>
            <position position="138"/>
        </location>
        <ligand>
            <name>S-adenosyl-L-methionine</name>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <position position="189"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="11">
        <location>
            <position position="189"/>
        </location>
        <ligand>
            <name>S-adenosyl-L-methionine</name>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="12">
        <location>
            <position position="192"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <position position="217"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <position position="218"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="12">
        <location>
            <position position="218"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="12">
        <location>
            <position position="247"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00012880"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022603"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022691"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022867"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022939"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023453"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PIRNR" id="PIRNR037177"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PIRSR" id="PIRSR037177-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PIRSR" id="PIRSR037177-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PIRSR" id="PIRSR037177-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029662.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="262" mass="29848" checksum="8CFF9B2117F95E82" modified="2019-05-08" version="1">MLESYPILLSIVTSLLFIVLLLAVLIRNNGWAAILWNEIIMQKINNFIMAQSKEERILNFVLQNAVRGDPQSVVDTIDKYCSQKEWAMNVGAEKGLILDKILEETKPSVLLELGTYCGYSAVRIARLLKPSARLLTIEFNPDFAAIARQMIEFAGVQDKVKIVEGSSETIIPQLKKKHEVDTLDFVFLDHWKDRYLPDTILLLDQGLLRKGSVILADNVIFPGAPEFIKYIRNNNRFECTNYPSYVEYMKVKDAMEKAVFVG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INW5</accession>
    <name>A0A452INW5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">NACHT domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Inflammasome</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the NLRP family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452INW5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033827.1">
        <property type="protein sequence ID" value="ENSGAGP00000029790.1"/>
        <property type="gene ID" value="ENSGAGG00000021504.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006954">
        <property type="term" value="P:inflammatory response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045087">
        <property type="term" value="P:innate immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001611">
        <property type="entry name" value="Leu-rich_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007111">
        <property type="entry name" value="NACHT_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041267">
        <property type="entry name" value="NLRP_HD2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050637">
        <property type="entry name" value="NLRP_innate_immun_reg"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041075">
        <property type="entry name" value="NOD1/2_WH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45690">
        <property type="entry name" value="NACHT, LRR AND PYD DOMAINS-CONTAINING PROTEIN 12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45690:SF19">
        <property type="entry name" value="NACHT, LRR AND PYD DOMAINS-CONTAINING PROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13516">
        <property type="entry name" value="LRR_6"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05729">
        <property type="entry name" value="NACHT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17776">
        <property type="entry name" value="NLRC4_HD2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17779">
        <property type="entry name" value="WHD_NOD2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00368">
        <property type="entry name" value="LRR_RI"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52047">
        <property type="entry name" value="RNI-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50837">
        <property type="entry name" value="NACHT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9" id="KW-1271">Inflammasome</keyword>
    <keyword evidence="8" id="KW-0395">Inflammatory response</keyword>
    <keyword evidence="5" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="7" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="NACHT" evidence="10">
        <location>
            <begin position="25"/>
            <end position="157"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004110"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008665"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023198"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023233"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029790.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="833" mass="95398" checksum="48C01ED57EF55FF2" modified="2019-05-08" version="1">MTERASSVITMGTLFKPYKDGQTPQIVVLVGAAGIGKTITSKKIMLDWAAGELYKEMFDYVFYINCREMNLLTKEVSVADMILKHCPNNKTSINHILGQSDKLLFIIDELRFSFDQSEDNLCSERWEKKPVEIILSSLFRKTVLPESYLLITTRPAALEKLGRCERYAEILGFSEADRKEYFHKFFGNEKQARQAYRFVRENEILFTMCFVPIMCWIICTVLKQQLEKGEDLAQASKTATGMYVLYLSSLVNPPGSNSKPCKKASLRGLCSLAADGIWKQKILFEEEEIKMLGLDQADALPLFLNENIFQKDIDTECLYSFIHLSFQEFFAALFYVLEEDETMEDSGTPKKDVKTLLENYGNSRNYLMLTVRFLYGLLNEKSIKDMEEKFGCKISPKIKLDLIKWIQTYKQTVLSVPSVYDKIICQLEEFRCLYEIQDENFMKNALEPFTKIVICRNNLTPMDQIVLSFCVNNCHRLVSLEMYACTFRFEDRGYFNIQEIELKSVSRDQDRTFLFSLSRDQHQDELKHSRVFLLCQALKDPNCKLKKLYRTDRTAHFLICASISCSLRSCKLTADCCGDLSSVLSTNQFLTDLDLTANEVRNTGVQLLCEGLKHQNCKLQRLRLNNCNITHVSCKDLAAVLKTSRRLIELDVGTNELQEAGVQLLCEALKHPNCRLRHWLMGLAVKNTAQVTSQTLAELNLQDNRLEDSGVQQLCEALKHPNCKLQKIKLQDCDLTDVCCGDLSSVLSTSQTLSELNLKHNKLEDSGVQLLCEGLKHPRCKLQKLNLKIGKEIEKVGGWGKEEEKDKEMKQIQSLVVTIYRAPQVLCPSLLFS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INW6</accession>
    <name>A0A452INW6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Homer scaffold protein 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the UPF0235 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452INW6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033607.1">
        <property type="protein sequence ID" value="ENSGAGP00000029596.1"/>
        <property type="gene ID" value="ENSGAGG00000021378.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1200.10">
        <property type="entry name" value="YggU-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="HAMAP" id="MF_00634">
        <property type="entry name" value="UPF0235"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003746">
        <property type="entry name" value="DUF167"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036591">
        <property type="entry name" value="YggU-like_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00251">
        <property type="entry name" value="DUF167 family protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13420">
        <property type="entry name" value="UPF0235 PROTEIN C15ORF40"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13420:SF7">
        <property type="entry name" value="UPF0235 PROTEIN C15ORF40"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02594">
        <property type="entry name" value="DUF167"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01152">
        <property type="entry name" value="DUF167"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69786">
        <property type="entry name" value="YggU-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="25"/>
            <end position="53"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00010364"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029596.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="5">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="148" mass="15674" checksum="CE60C8D79B1E1AA1" modified="2019-05-08" version="1">MLVALAGVRRCSGWRQGPVRAGVRSWSGAMPRRGKEKCKDSVKPPVPSGPVVTDKSGSLTIAIHAKPGSKQNAITDVTTEAVGVAIAAPPSEGEANAELCRYLSKVLEVKKSEVVLDKGGKSREKVVKILVSMTPDEVLEKLKKEATS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INW7</accession>
    <name>A0A452INW7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">Ras-GAP domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452INW7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033584.1">
        <property type="protein sequence ID" value="ENSGAGP00000029575.1"/>
        <property type="gene ID" value="ENSGAGG00000019812.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005938">
        <property type="term" value="C:cell cortex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903479">
        <property type="term" value="P:mitotic actomyosin contractile ring assembly actin filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05131">
        <property type="entry name" value="RasGAP_IQGAP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00201">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.506.10:FF:000004">
        <property type="entry name" value="IQ motif containing GTPase activating protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.190">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.506.10">
        <property type="entry name" value="GTPase Activation - p120gap, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000048">
        <property type="entry name" value="IQ_motif_EF-hand-BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000593">
        <property type="entry name" value="RasGAP_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023152">
        <property type="entry name" value="RasGAP_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001936">
        <property type="entry name" value="RasGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001202">
        <property type="entry name" value="WW_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149">
        <property type="entry name" value="RAS GTPASE-ACTIVATING PROTEIN WITH IQ MOTIF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149:SF12">
        <property type="entry name" value="RAS GTPASE-ACTIVATING-LIKE PROTEIN IQGAP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00612">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00616">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03836">
        <property type="entry name" value="RasGAP_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00015">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00323">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00456">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF143885">
        <property type="entry name" value="RGC domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00509">
        <property type="entry name" value="RAS_GTPASE_ACTIV_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50018">
        <property type="entry name" value="RAS_GTPASE_ACTIV_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01159">
        <property type="entry name" value="WW_DOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Ras-GAP" evidence="1">
        <location>
            <begin position="600"/>
            <end position="833"/>
        </location>
    </feature>
    <evidence type="ECO:0000259" key="1">
        <source>
            <dbReference type="PROSITE" id="PS50018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029575.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1260" mass="144485" checksum="9FFFCB3FBD623916" modified="2019-05-08" version="1">LTHKTIRSHNWCNGKNPINFLYFPPELVAVDHINTAIRNCDANKTLVALMKPEAQLPVVHPFAAAVYQTELFNLQQQNAVRYLAHDELSIAVEMLSAVVLLNQALGSKDILAIKNHLRNPSVGFNNVEDENFQRYADTLLSIKSEASSQGQDYLSWNDVQNCIDMVNMQVQEENDRIVAVSYINEAIDQGNPGKTLETLLLPTVKLHDVNPTNARHYQDVLHDAKVQKCKVEPQLNCICVYVKYKMSDVEMQFCCLVALGTSMINNCLEKHDSQPVLDILQSPKFGLRVVPECAETYYKNLLEAKNLKTKEDSSEGPWLKLIMKNRYDYYYNVETGESTFVPPEGFIPKTSWLTSEEIQTIVGQVTADYNREQLWFANENLILQLQALARGFLVRKSYEERKGFLQKQEPSVIKIQVLLCNIKLFYILWSFFQNEEIVKIQAFLRANKARDDYRTLIGAENPPLNVLRKFAYLLDQSDLDFQEELEVTRLREEVVTKIRSNQQLEKDLNLMDIKIGLLVKNRISLQDVVLHSKKLNKKSKNQVEEMVTGDKQGIKGLSKERRKQLEAYQHLFYLLQTNPMYLAKLIFQMPQNKSTKFMDTVIFTLYNYASNQREEYLLLKLFKTALEEEINSKVDQIQDIVTGNPTVIKMVVSFNRGARGQNSLRQLLAPVVKEIMDDKSLLINTSPVEVYKAWVNQLETQTGEASKLPYDVTTEQALEHPEVVNRLESSIQSLRTVTDKVLVSIFSSLSMMPYGMRYIAKVLKNSLHEKFPDATEDELLKIVGNLLYYRYMNPAIVAPDGFDIIDMTAGGQIHSDQRRNLGCVAKVLQHAASNKLFEGESAHLSSMNTYLSQTYQKFRNFFQIACDVPEPEEKFNIDEYSDMVTLSKPVIYISIEEIINTHSLLLEHQDAIAPEQNDLLNELLKGLGAVPDVESFLGEGAVDPNDPNKENTLSQLAKTEISLSLTSKYELQEGEDQDLKGLMIKTKRLVVDVIRTQPGDTLLEILEIPASAQQESEHLKVVEKRAILDSKTPEKMKHSQSMLEDGQLPIEQKKRKIQRNLRTLEQAGLVSSANKYQEIINEIAKDIRNQRRYRHHRKAELVKLQQTLSALQSKTAFYEDQVNYYNTYIKTCLDNLTRKNSRRSIKLDGKVEEKGSKKLKHTSLKYTAARLHEKGVILEIEDLQTNQFKNVMFDITPGEEMGDFEIKAKFLGVQMEKVELHFQDLLQMQYEGVAVMKMFDKAKVNVNLLIFLLNKKFYGK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INW9</accession>
    <name>A0A452INW9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Protein Wnt</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Ligand for members of the frizzled family of seven transmembrane receptors.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 9">Secreted</location>
            <location evidence="1 9">Extracellular space</location>
            <location evidence="1 9">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 9">Belongs to the Wnt family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452INW9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033689.1">
        <property type="protein sequence ID" value="ENSGAGP00000029668.1"/>
        <property type="gene ID" value="ENSGAGG00000021418.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005125">
        <property type="term" value="F:cytokine activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005109">
        <property type="term" value="F:frizzled binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060070">
        <property type="term" value="P:canonical Wnt signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045165">
        <property type="term" value="P:cell fate commitment"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030182">
        <property type="term" value="P:neuron differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005817">
        <property type="entry name" value="Wnt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018161">
        <property type="entry name" value="Wnt_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12027:SF84">
        <property type="entry name" value="PROTEIN WNT-9B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12027">
        <property type="entry name" value="WNT RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00110">
        <property type="entry name" value="wnt"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01349">
        <property type="entry name" value="WNTPROTEIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00097">
        <property type="entry name" value="WNT1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00246">
        <property type="entry name" value="WNT1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 9" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="7" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="8" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="6 9" id="KW-0879">Wnt signaling pathway</keyword>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="261"/>
            <end position="280"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="262"/>
            <end position="280"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005683"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022687"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="RuleBase" id="RU003500"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029668.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="352" mass="38944" checksum="55CAACEEE62E1D39" modified="2019-05-08" version="1">PRPPPRTSLWPRCLSGLISLPALSPLSLTGKEGLTHLPSLGPSANPAQGKAHLKQCDLLKLSRKQKRLCQREPGLAETLRDAIRLSIMECQYQFRYERWNCSLDGRTSLLKRGFKETAFLYAVSSAALTHSLARACSAGRMERCTCDDSPDLENRKAWQWGVCGDNLKYSTKFLKNFLGQKKIGKDLRAKVDIHNTNVGIKAVKNGLKTTCKCHGVSGSCAVRTCWKQLSPFHETGKILKLRFDDAVKIFSATNDAVGHSELVSTQRRSHSPKSPATPRSTDLVYVEDSPSFCRPSKYSLGTAGRTCSREASCDISSILPQSLPEGKLASVTHHNILCLWYPPASSWTTERV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INX3</accession>
    <name>A0A452INX3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Tubby-like protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 6">Belongs to the TUB family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452INX3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033559.1">
        <property type="protein sequence ID" value="ENSGAGP00000029552.1"/>
        <property type="gene ID" value="ENSGAGG00000021348.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005929">
        <property type="term" value="C:cilium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061512">
        <property type="term" value="P:protein localization to cilium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.20.90.10:FF:000001">
        <property type="entry name" value="Tubby-like protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.20.90.10">
        <property type="entry name" value="Tubby Protein, Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025659">
        <property type="entry name" value="Tubby-like_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000007">
        <property type="entry name" value="Tubby_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018066">
        <property type="entry name" value="Tubby_C_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005398">
        <property type="entry name" value="Tubby_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16517:SF20">
        <property type="entry name" value="TUBBY PROTEIN HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16517">
        <property type="entry name" value="TUBBY-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01167">
        <property type="entry name" value="Tub"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16322">
        <property type="entry name" value="Tub_N"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01573">
        <property type="entry name" value="SUPERTUBBY"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01574">
        <property type="entry name" value="TUBBYPROTEIN"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54518">
        <property type="entry name" value="Tubby C-terminal domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01200">
        <property type="entry name" value="TUB_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01201">
        <property type="entry name" value="TUB_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0964">Secreted</keyword>
    <feature type="domain" description="Tubby N-terminal" evidence="9">
        <location>
            <begin position="29"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="domain" description="Tubby N-terminal" evidence="9">
        <location>
            <begin position="145"/>
            <end position="211"/>
        </location>
    </feature>
    <feature type="domain" description="Tubby C-terminal" evidence="8">
        <location>
            <begin position="229"/>
            <end position="472"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="36"/>
            <end position="67"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="112"/>
            <end position="217"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="145"/>
            <end position="155"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="7">
        <location>
            <begin position="171"/>
            <end position="181"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="195"/>
            <end position="216"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007129"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="RuleBase" id="RU361125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF01167"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF16322"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029552.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="477" mass="53633" checksum="93F45E21DBDEE823" modified="2019-05-08" version="1">MTSKHQADWIPYSVLDDEGSNLRQQKLDRQRALLEQKQKKKRQEPLMVQSNVDSRTRTRRMKHSEEQAPLVESYLNSNSSTIFHGIDGPAAFQDDVQEVGSQIQILTVGQSSHNEDDGEMMASGQQQSRQDLRVTMQKKDPHANTSWSTSTSQSSLVALDHVPGISSSMNFDEEDEDDEDSSSSSQLNSNTRPGSATSKKSNKEAASAPSPSANEPVIDVDDLEEFAVRPAPQGVTVKCRITRDKKGMDRGMYPTYYLHLEREDGKKVFLLAGRKRKKSKTSNYLISIDPTDLSRGGESFIGKLRSNLMGTKFTVYDNGVNPMKTTSSLEASNLRQELAAICYETNVLGFKGPRKMSVVIPGMNMDHERVSIRPRNEHETLLARWQNKNTESVIELHNKTPVWNDDTQSYVLNFHGRVTQASVKNFQIIHDNDPDYIVMQFGRVAEDVFTMDYNYPMCALQAFAIALSSFDSKLACE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INZ8</accession>
    <name>A0A452INZ8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Mitogen-activated protein kinase kinase kinase 7</fullName>
            <ecNumber evidence="5">2.7.11.25</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="30 31">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="31" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="30" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="30" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="25">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.25"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="24">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
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            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
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            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.25"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Peripheral membrane protein</topology>
            <orientation evidence="2">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the protein kinase superfamily. STE Ser/Thr protein kinase family. MAP kinase kinase kinase subfamily.</text>
    </comment>
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    <dbReference type="Ensembl" id="ENSGAGT00000033645.1">
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        <property type="project" value="UniProtKB-SubCell"/>
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        <property type="term" value="F:enzyme binding"/>
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        <property type="project" value="UniProtKB-ARBA"/>
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        <property type="term" value="F:magnesium ion binding"/>
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        <property type="project" value="InterPro"/>
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        <property type="project" value="UniProtKB-EC"/>
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    <dbReference type="GO" id="GO:0006915">
        <property type="term" value="P:apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006955">
        <property type="term" value="P:immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007254">
        <property type="term" value="P:JNK cascade"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043123">
        <property type="term" value="P:positive regulation of canonical NF-kappaB signal transduction"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043410">
        <property type="term" value="P:positive regulation of MAPK cascade"/>
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        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009893">
        <property type="term" value="P:positive regulation of metabolic process"/>
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        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd14058">
        <property type="entry name" value="STKc_TAK1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000143">
        <property type="entry name" value="Mitogen-activated protein kinase kinase kinase 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049637">
        <property type="entry name" value="MAP3K7"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001245">
        <property type="entry name" value="Ser-Thr/Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46716">
        <property type="entry name" value="MITOGEN-ACTIVATED PROTEIN KINASE KINASE KINASE 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46716:SF1">
        <property type="entry name" value="MITOGEN-ACTIVATED PROTEIN KINASE KINASE KINASE 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07714">
        <property type="entry name" value="PK_Tyr_Ser-Thr"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF038168">
        <property type="entry name" value="MAPKKK7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00109">
        <property type="entry name" value="TYRKINASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="13" id="KW-0053">Apoptosis</keyword>
    <keyword evidence="17 26" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="7" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="27" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="8" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="16" id="KW-0418">Kinase</keyword>
    <keyword evidence="18" id="KW-0460">Magnesium</keyword>
    <keyword evidence="22" id="KW-0472">Membrane</keyword>
    <keyword evidence="14" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="15 26" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="11" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="31" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="21" id="KW-0346">Stress response</keyword>
    <keyword evidence="23" id="KW-0804">Transcription</keyword>
    <keyword evidence="20" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="12" id="KW-0808">Transferase</keyword>
    <keyword evidence="19" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="Protein kinase" evidence="29">
        <location>
            <begin position="1"/>
            <end position="259"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="28">
        <location>
            <begin position="269"/>
            <end position="311"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="28">
        <location>
            <begin position="326"/>
            <end position="376"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="28">
        <location>
            <begin position="399"/>
            <end position="454"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="27">
        <location>
            <begin position="491"/>
            <end position="518"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="28">
        <location>
            <begin position="274"/>
            <end position="300"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="28">
        <location>
            <begin position="404"/>
            <end position="419"/>
        </location>
    </feature>
    <feature type="binding site" evidence="26">
        <location>
            <position position="53"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004413"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00006529"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012406"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00017660"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022703"/>
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    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00023016"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
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        </source>
    </evidence>
    <evidence type="ECO:0000313" key="30">
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    </evidence>
    <evidence type="ECO:0000313" key="31">
        <source>
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        </source>
    </evidence>
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</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J088</accession>
    <name>A0A452J088_9SAUR</name>
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        <recommendedName>
            <fullName evidence="11">G protein-coupled receptor kinase</fullName>
            <ecNumber evidence="11">2.7.11.-</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>[G-protein-coupled receptor] + ATP = [G-protein-coupled receptor]-phosphate + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:12008"/>
            <dbReference type="Rhea" id="RHEA-COMP:11260"/>
            <dbReference type="Rhea" id="RHEA-COMP:11261"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
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            <dbReference type="ChEBI" id="CHEBI:43176"/>
            <dbReference type="ChEBI" id="CHEBI:68546"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.16"/>
        </reaction>
    </comment>
    <comment type="similarity">
        <text evidence="2 11">Belongs to the protein kinase superfamily. AGC Ser/Thr protein kinase family. GPRK subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.-" evidence="11"/>
    <dbReference type="AlphaFoldDB" id="A0A452J088"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038293.1">
        <property type="protein sequence ID" value="ENSGAGP00000033805.1"/>
        <property type="gene ID" value="ENSGAGG00000024060.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
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        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004703">
        <property type="term" value="F:G protein-coupled receptor kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009966">
        <property type="term" value="P:regulation of signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd05605">
        <property type="entry name" value="STKc_GRK4_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.167.10:FF:000009">
        <property type="entry name" value="G protein-coupled receptor kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000074">
        <property type="entry name" value="G protein-coupled receptor kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:001100">
        <property type="entry name" value="G protein-coupled receptor kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.167.10">
        <property type="entry name" value="Regulator of G-protein Signalling 4, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000961">
        <property type="entry name" value="AGC-kinase_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000239">
        <property type="entry name" value="GPCR_kinase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016137">
        <property type="entry name" value="RGS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036305">
        <property type="entry name" value="RGS_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044926">
        <property type="entry name" value="RGS_subdomain_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24355:SF26">
        <property type="entry name" value="G PROTEIN-COUPLED RECEPTOR KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24355">
        <property type="entry name" value="G PROTEIN-COUPLED RECEPTOR KINASE/RIBOSOMAL PROTEIN S6 KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00615">
        <property type="entry name" value="RGS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00717">
        <property type="entry name" value="GPCRKINASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00315">
        <property type="entry name" value="RGS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00133">
        <property type="entry name" value="S_TK_X"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48097">
        <property type="entry name" value="Regulator of G-protein signaling, RGS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51285">
        <property type="entry name" value="AGC_KINASE_CTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50132">
        <property type="entry name" value="RGS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 10 11" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="7 11" id="KW-0418">Kinase</keyword>
    <keyword evidence="6 10 11" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 11" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="5 11" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="RGS" evidence="13">
        <location>
            <begin position="59"/>
            <end position="171"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="12">
        <location>
            <begin position="186"/>
            <end position="448"/>
        </location>
    </feature>
    <feature type="domain" description="AGC-kinase C-terminal" evidence="14">
        <location>
            <begin position="449"/>
            <end position="514"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="9">
        <location>
            <position position="311"/>
        </location>
    </feature>
    <feature type="binding site" evidence="10">
        <location>
            <position position="224"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001256"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009793"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PIRSR" id="PIRSR600239-51"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="RuleBase" id="RU000308"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50132"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS51285"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033805.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="608" mass="70010" checksum="9DC0668972D98998" modified="2019-05-08" version="1">MELENIVANTVLLKAREGGGGKRKGRSKKWKEILRFPHISQCDDLRKGLEREYSSLCDKQPIGRLLFRQFCERRPELQRCIHFLDAVIDYELSSDERRKEMGDEIIRRFFKSESSDYMPEISQALMNQCMENLQKSACKDLFSNCLHPMHEYLSGAPFADYRDSMYFDRFLQWKYLERQSVTKDTFRQYRILGKGGFGEVCACQVRATGKMYACKKLEKKRIKKRKGEAMALNEKQILEKVNSRFVVSLAYAYETKDALCLVLTIMNGGDLKFHIYNMGNPGFEDERVIFYAAEICCGLQHLHQEGIVYRDLKPENILLDDDGHIRISDLGLAIKIPEGEMIRGRVGTVGYMAPEVINNERYTFSPDWWGLGCLIYEMIEGQSPFRARKERVKREEVEKRVQEEQELYSDKFTEDTKAICKMLLTKDPKQRLGCGEDGATQVKQHPFFRTINFKRLEAGVIKPSFVPDPRAVYCKDVLDIEQFSTVKGVNLDQTDSDFYAKFATGSVSIPWQNEMIETECFKDLNVFGPNGMRSPDLDWKQLPEPPKRSLLQRLFRRHPADYTISTNTHSNLTAGVNSHPPTANSVCQGRAMAQVSSCQRDSSLETGP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J089</accession>
    <name>A0A452J089_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="6">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the CLEC16A/gop-1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J089"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038252.1">
        <property type="protein sequence ID" value="ENSGAGP00000033771.1"/>
        <property type="gene ID" value="ENSGAGG00000024031.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005770">
        <property type="term" value="C:late endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006914">
        <property type="term" value="P:autophagy"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016197">
        <property type="term" value="P:endosomal transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1901096">
        <property type="term" value="P:regulation of autophagosome maturation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007034">
        <property type="term" value="P:vacuolar transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039272">
        <property type="entry name" value="CLEC16A/TT9"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045820">
        <property type="entry name" value="CLEC16A/TT9_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019155">
        <property type="entry name" value="CLEC16A/TT9_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21481">
        <property type="entry name" value="PROTEIN CLEC16A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21481:SF0">
        <property type="entry name" value="PROTEIN CLEC16A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19439">
        <property type="entry name" value="CLEC16A_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09758">
        <property type="entry name" value="FPL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0072">Autophagy</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="FPL" evidence="4">
        <location>
            <begin position="51"/>
            <end position="198"/>
        </location>
    </feature>
    <feature type="domain" description="CLEC16A/TT9 C-terminal" evidence="5">
        <location>
            <begin position="243"/>
            <end position="876"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="379"/>
            <end position="429"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="972"/>
            <end position="996"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="3">
        <location>
            <begin position="379"/>
            <end position="390"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="985"/>
            <end position="996"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006441"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023006"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="Pfam" id="PF09758"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="Pfam" id="PF19439"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033771.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="996" mass="112800" checksum="023BB26E6DF0C7E1" modified="2019-05-08" version="1">MFGRSRSWVGGGHGKAARSIHSLDHLKYMYHVLTKNTTVTDHNRNLLVETIRSITEILIWGDQNDSSVFDFFLEKNMFVFFLNILRQKSGRYVCVQLLQTLNILFENISHETSLYYLLSNNYVNSIIVHKFDFSDEEIMAYYISFLKTLSLKLNNHTVHFFYNEHTNDFALYTEAIKFFNHPESMVRIAVRTITLNVYKVDNQPMLHYIRDKTAVPYFSNLVWFIGSHVIELDNCVQTDEEHRNRGKLSDLVAEHLDHLHYLNDILIINCEFLNDVLTDHLLNRLFLPLYVYSLVNQDKGGERPKISLQVSLYLLSQVFLIIHYAPLVNSLAEVILNGDLSVFSSKVEQDIQRSSAKSSIRCFIKPTETLERSLEINKHKGKKRLQKRPNYKNVGEEEEEERGLENTQEDPDKAKGTEGSSKGIKASGENEEIEMVIMERCKMSELSISVVTEQNTTDEEKSAAASGSEMTNWSRPFLDMVYNAQDCAEDDYYALFVLCLLYAMSQNKGIDPVKLERIQLPAQHAEEKSSYNHVLAERLIRIMNYAAQPDGKVRLATLELGCLLLKQLVLSKSGSIIKDVHLACLEGAREESVHLLRRFYKGEEIFLDMFEDEYRSMTSKPMNVEYLMMDASILLPPTGTPLTGIDFVKRLPCGDVERTRRAIRVFFMLRSLSLHLRDEPETQLPLTREEDLIKTDDVLDLNNSDLIACTVITKDGGQVQRFLAVDIYQMSLVEPDVARLGWGVVKFAGLLQDMQVTGVEDDSRALNIIIHKPASSPHSKPFPILQATFIFSDHIRCIIAKQRLAKGRIQARRMKMQRIAALLDLPVQPSTEVMGFGHSSASAAQHLPFRFYDQSRRGSCDPAVQRSVFASVDKVPVDDSSAPEQPPIIFSSQMMMVDETSSAASKSGKNPVKNPEVETANLSPSLIPAQQPTISLLSDDNADALSVESLTLVPPIDPHSIHTFSCIPQSSLPSEVCGVTEGEGSDQSQQADYDEQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J090</accession>
    <name>A0A452J090_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Glucose-6-phosphate exchanger SLC37A2</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="13">Solute carrier family 37 member 2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>D-glucose 6-phosphate(in) + phosphate(out) = D-glucose 6-phosphate(out) + phosphate(in)</text>
            <dbReference type="Rhea" id="RHEA:71535"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:61548"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the major facilitator superfamily. Organophosphate:Pi antiporter (OPA) (TC 2.A.1.4) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J090"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038250.1">
        <property type="protein sequence ID" value="ENSGAGP00000033769.1"/>
        <property type="gene ID" value="ENSGAGG00000024013.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061513">
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    <keyword evidence="9 14" id="KW-0472">Membrane</keyword>
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    <keyword evidence="5" id="KW-0762">Sugar transport</keyword>
    <keyword evidence="6 14" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 14" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="15"/>
            <end position="34"/>
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    <feature type="transmembrane region" description="Helical" evidence="14">
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    <sequence length="483" mass="52119" checksum="A450C31ED743B3DD" modified="2019-05-08" version="1">MPCLSLVPFLVLPPLRYRGLTLVLTFVFYASFHLSRKPISIVKVSAVPGIFLLLLLLFNMLASACIVTCFPDQANYNELFGALDNAFLVAYAIGMFISGIFGERLPLRYYLSGGMVLSGLFTSLFGLAYFWEIHVLWYYIIIQICNGLVQTTGWPSVVTCVGNWFGKGKRGLIMGIWNSHTSVGNILGSLIAGVWVSSAWGLSFIVPGIIIAIMGIVCFLFLVEYPEVTLSADDGSCSRENTIDSSEYPKEPAEQPGAISFLGALRIPGVVEFSLCLLFAKLVSYTFLYWLPLYIVNVAHFSAKEAGDLSTLFDVGGILGGILAGLISDYTDGRATTCCVMLIVAAPMLFLYNHVGQNGIGSSIAMLIVCGALVNGPYALITTAVSADLGTHECLKGNAKALSTVTAIIDGTGSVGAALGPLLAGLISPTGWNNVFYMLITADILACLFLSRVVYKEIKGWCGCYTRKRGYVLLKSQVCRVFL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J091</accession>
    <name>A0A452J091_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">G protein-coupled receptor kinase</fullName>
            <ecNumber evidence="11">2.7.11.-</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>[G-protein-coupled receptor] + ATP = [G-protein-coupled receptor]-phosphate + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:12008"/>
            <dbReference type="Rhea" id="RHEA-COMP:11260"/>
            <dbReference type="Rhea" id="RHEA-COMP:11261"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
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            <dbReference type="ChEBI" id="CHEBI:43176"/>
            <dbReference type="ChEBI" id="CHEBI:68546"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.16"/>
        </reaction>
    </comment>
    <comment type="similarity">
        <text evidence="2 11">Belongs to the protein kinase superfamily. AGC Ser/Thr protein kinase family. GPRK subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.-" evidence="11"/>
    <dbReference type="AlphaFoldDB" id="A0A452J091"/>
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        <property type="protein sequence ID" value="ENSGAGP00000033810.1"/>
        <property type="gene ID" value="ENSGAGG00000024060.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004703">
        <property type="term" value="F:G protein-coupled receptor kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009966">
        <property type="term" value="P:regulation of signal transduction"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd05605">
        <property type="entry name" value="STKc_GRK4_like"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="G protein-coupled receptor kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
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        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
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    </dbReference>
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    </dbReference>
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        <property type="entry name" value="GPCR_kinase"/>
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        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
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        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
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        <property type="entry name" value="Protein_kinase_ATP_BS"/>
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        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
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        <property type="entry name" value="G PROTEIN-COUPLED RECEPTOR KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24355">
        <property type="entry name" value="G PROTEIN-COUPLED RECEPTOR KINASE/RIBOSOMAL PROTEIN S6 KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00717">
        <property type="entry name" value="GPCRKINASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00315">
        <property type="entry name" value="RGS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00133">
        <property type="entry name" value="S_TK_X"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48097">
        <property type="entry name" value="Regulator of G-protein signaling, RGS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51285">
        <property type="entry name" value="AGC_KINASE_CTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
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    </dbReference>
    <dbReference type="PROSITE" id="PS50132">
        <property type="entry name" value="RGS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 10 11" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="7 11" id="KW-0418">Kinase</keyword>
    <keyword evidence="6 10 11" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 11" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="5 11" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="RGS" evidence="13">
        <location>
            <begin position="48"/>
            <end position="160"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="12">
        <location>
            <begin position="175"/>
            <end position="437"/>
        </location>
    </feature>
    <feature type="domain" description="AGC-kinase C-terminal" evidence="14">
        <location>
            <begin position="438"/>
            <end position="503"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="9">
        <location>
            <position position="300"/>
        </location>
    </feature>
    <feature type="binding site" evidence="10">
        <location>
            <position position="213"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
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        <source>
            <dbReference type="ARBA" id="ARBA00001256"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00009793"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
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        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PIRSR" id="PIRSR600239-51"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="RuleBase" id="RU000308"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033810.1"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="597" mass="68692" checksum="4440E5D92F7CD9AE" modified="2019-05-08" version="1">MASAVFGGGGKRKGRSKKWKEILRFPHISQCDDLRKGLEREYSSLCDKQPIGRLLFRQFCERRPELQRCIHFLDAVIDYELSSDERRKEMGDEIIRRFFKSESSDYMPEISQALMNQCMENLQKSACKDLFSNCLHPMHEYLSGAPFADYRDSMYFDRFLQWKYLERQSVTKDTFRQYRILGKGGFGEVCACQVRATGKMYACKKLEKKRIKKRKGEAMALNEKQILEKVNSRFVVSLAYAYETKDALCLVLTIMNGGDLKFHIYNMGNPGFEDERVIFYAAEICCGLQHLHQEGIVYRDLKPENILLDDDGHIRISDLGLAIKIPEGEMIRGRVGTVGYMAPEVINNERYTFSPDWWGLGCLIYEMIEGQSPFRARKERVKREEVEKRVQEEQELYSDKFTEDTKAICKMLLTKDPKQRLGCGEDGATQVKQHPFFRTINFKRLEAGVIKPSFVPDPRAVYCKDVLDIEQFSTVKGVNLDQTDSDFYAKFATGSVSIPWQNEMIETECFKDLNVFGPNGMRSPDLDWKQLPEPPKRSLLQRLFRRHPADYTISTNTHSNLTAGVNSHPPTANSVCQGRAMAQVSSCQRDSSLETGP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J092</accession>
    <name>A0A452J092_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000038212.1">
        <property type="protein sequence ID" value="ENSGAGP00000033733.1"/>
        <property type="gene ID" value="ENSGAGG00000024011.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051018">
        <property type="term" value="F:protein kinase A binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008382">
        <property type="entry name" value="SPHK1-interactor_AKAP_110"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10226">
        <property type="entry name" value="A KINASE ANCHOR PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10226:SF7">
        <property type="entry name" value="A-KINASE ANCHOR PROTEIN SPHKAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="208"/>
            <end position="231"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="345"/>
            <end position="375"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="517"/>
            <end position="536"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="744"/>
            <end position="772"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1217"/>
            <end position="1247"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1379"/>
            <end position="1416"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1499"/>
            <end position="1534"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="211"/>
            <end position="221"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="354"/>
            <end position="363"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="517"/>
            <end position="534"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="1">
        <location>
            <begin position="747"/>
            <end position="764"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="1393"/>
            <end position="1407"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="1508"/>
            <end position="1523"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033733.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1657" mass="184106" checksum="043655723E9D8263" modified="2019-05-08" version="1">MYEVSEHQVSGTDSSESSLGSSVTACKKVLCSNSLLESTDYWLQNQRTPCRIGFVEDKSENKCASVCFVNLDANKNDCSEEHLKQRLINVSPNLPKLISSMNVQPPKENEIVLLSGLTSGNLQADFEVSQCPWLADVCLVQCARGNRKNSIGCIIFEINKFLIGLELVQERQLQIETSILKPEDDTNCSVSSIEEDFLTASEHLEDENEADEYKNGHDNIHGTEPASDIKKHKGKGLENLHYRKTTLPFALDDNCFNKENLASAKNSTDSEEALNIVSEDSILQTVDKSIQQIQWKNGLAGRVRSSPNTNSLERSLTDKAENSYPICSSEGVSLTRMAKEELASQCDTTAKQNNKLDTEERTSERKHHPPLQNEQATTGQYATNLAESVLQDAFIRLSQSQPTFTKEAAISISVGNSFSSATCTTEDITSRSWNELPKIVIVQSPDSCENTSEWPGSGFPNLCHWSESESSAEISDYLQEENSKGHTQNVLEVALACAAKVIGTISSPHAAERLKRELATDSKSKTVDNEEEQIKSSQVLDNGADIEYSFPSALCGMTQVASAVAVCGLGETKEDKYSATSRGLLSAAETSAAITLHCSIAIGSSMEKLNRNIAETLLKEASIILTKPNTYRNVGDFMESINGRIIETATRPRISPLDEVICDELAQNVSNVILRHSVEEVRKKRQLHPGSDNNLDFSTHDMFMETANELLFNVIYFTCKKMRDIAQVGECSPLFSEDKDRWRVTEAESQATQTTATQPSQQAPDHTNEPFSTFYSANAGKDLVNMTNTKKDNEEGVPNTMESATLHSDLLCSASGHYSLVAKTSPKKRYLKRATRDCYRSPYQSKKDLRSFSDRENTLTVNECRHGVQEQLSSDAIVNTENQSKHKCDALLNNEVQVSVPLLGNHILLPSQPVLQVKHPRDTYCVTDFAEELAETVVSMATEIAAICLENSNGKQPWFCAWKRGNEYLVTQGLSCRTIKRKKESQPNGSVVRKHRPPRLSEIKRKTDEHPELKERLMNRVVDESINLEDTPESVSIFANEVAAKIMNLTELSMVDNVWQGPNHPRNRLHCERWGRAKGSSCESIPEEDLDSKGSVNTLGPMNMLGQPGSQTSSVSKQSSCESITDEFSRFMVNQMENEGRGFDLLLDYYAGKNASSILTSALQQVARKNCHLNVRPSCPSKQSSTESITEEFYKYMLREIEKENKDDASSIRSSREWSSSLLSPSPRSPFCFRQSSMPDSRSSGSRLTVNAPVKAKSLDGFAHNSHRDSLSIQQVSTVSSLGLCKSDSCLYQRCRTDQVTDMLIHETWASSIESLMRKNKIIGDEAEVTDADQFPSDSPLHVEQYANRLAANIVESGKTLIVVHQDSFDYTNREHVAEGRSLPSVSQSQSKPTRDRINLDGKKEPLTTHGSLQGSHIGSSAFLREVPLIQIETDQREELNKDLEPLTSSIIPAGETKEHLNKEKPPAAYFGKHIVSSSVLNTSVDTRSKPDAELVVETKTAEDFPNPLNSSDESTGSWSQLVNDEDNPDDTSSYLQLSERSMSNGNSSTTSSLGIMDLEIYQENMPSSPMINELAKEKAFLKEQPENIEESTSELSVETANCQKDLLVINFDLEPECPDAELRATLQWIAASELGIRTIYFQRSQENRIEKVLFTM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J093</accession>
    <name>A0A452J093_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="6">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the CLEC16A/gop-1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J093"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038257.1">
        <property type="protein sequence ID" value="ENSGAGP00000033776.1"/>
        <property type="gene ID" value="ENSGAGG00000024031.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005770">
        <property type="term" value="C:late endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006914">
        <property type="term" value="P:autophagy"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016197">
        <property type="term" value="P:endosomal transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1901096">
        <property type="term" value="P:regulation of autophagosome maturation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007034">
        <property type="term" value="P:vacuolar transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039272">
        <property type="entry name" value="CLEC16A/TT9"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045820">
        <property type="entry name" value="CLEC16A/TT9_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019155">
        <property type="entry name" value="CLEC16A/TT9_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21481">
        <property type="entry name" value="PROTEIN CLEC16A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21481:SF0">
        <property type="entry name" value="PROTEIN CLEC16A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19439">
        <property type="entry name" value="CLEC16A_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09758">
        <property type="entry name" value="FPL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0072">Autophagy</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="FPL" evidence="4">
        <location>
            <begin position="51"/>
            <end position="198"/>
        </location>
    </feature>
    <feature type="domain" description="CLEC16A/TT9 C-terminal" evidence="5">
        <location>
            <begin position="243"/>
            <end position="932"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="379"/>
            <end position="429"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="886"/>
            <end position="951"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="3">
        <location>
            <begin position="379"/>
            <end position="390"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="3">
        <location>
            <begin position="910"/>
            <end position="928"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006441"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023006"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="Pfam" id="PF09758"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="Pfam" id="PF19439"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033776.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="951" mass="107768" checksum="E9F03A80E5D6CA16" modified="2019-05-08" version="1">MFGRSRSWVGGGHGKAARSIHSLDHLKYMYHVLTKNTTVTDHNRNLLVETIRSITEILIWGDQNDSSVFDFFLEKNMFVFFLNILRQKSGRYVCVQLLQTLNILFENISHETSLYYLLSNNYVNSIIVHKFDFSDEEIMAYYISFLKTLSLKLNNHTVHFFYNEHTNDFALYTEAIKFFNHPESMVRIAVRTITLNVYKVDNQPMLHYIRDKTAVPYFSNLVWFIGSHVIELDNCVQTDEEHRNRGKLSDLVAEHLDHLHYLNDILIINCEFLNDVLTDHLLNRLFLPLYVYSLVNQDKGGERPKISLQVSLYLLSQVFLIIHYAPLVNSLAEVILNGDLSVFSSKVEQDIQRSSAKSSIRCFIKPTETLERSLEINKHKGKKRLQKRPNYKNVGEEEEEERGLENTQEDPDKAKGTEGSSKGIKASGENEEIEMVIMERCKMSELSISVVTEQNTTDEEKSAAASGSEMTNWSRPFLDMVYNAQDCAEDDYYALFVLCLLYAMSQNKGIDPVKLERIQLPAQHAEEKSSYNHVLAERLIRIMNYAAQPDGKVRLATLELGCLLLKQLVLSKSGSIIKDVHLACLEGAREESVHLLRRFYKGEEIFLDMFEDEYRSMTSKPMNVEYLMMDASILLPPTGTPLTGIDFVKRLPCGDVERTRRAIRVFFMLRSLSLHLRDEPETQLPLTREEDLIKTDDVLDLNNSDLIACTVITKDGGQVQRFLAVDIYQMSLVEPDVARLGWGVVKFAGLLQDMQVTGVEDDSRALNIIIHKPASSPHSKPFPILQATFIFSDHIRCIIAKQRLAKGRIQARRMKMQRIAALLDLPVQPSTEVMGFGHSSASAAQHLPFRFYDQSRRGSCDPAVQRSVFASVDKVPGFAVAQYINQPNPSPLSSPSPSSSGSGSTGHCDSVTASSTSTPSAAHSPSVRSGRKGRRRVFSLSEADSHGGHWV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHL0</accession>
    <name>A0A452GHL0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Olduvai domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Golgi apparatus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000001196.1">
        <property type="protein sequence ID" value="ENSGAGP00000001062.1"/>
        <property type="gene ID" value="ENSGAGG00000000835.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060090">
        <property type="term" value="F:molecular adaptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007098">
        <property type="term" value="P:centrosome cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090063">
        <property type="term" value="P:positive regulation of microtubule nucleation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903358">
        <property type="term" value="P:regulation of Golgi organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056273">
        <property type="entry name" value="CDK5RAP2_MYOME_CC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012943">
        <property type="entry name" value="Cnn_1N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052593">
        <property type="entry name" value="MT-associated_AKAP9-binding"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010630">
        <property type="entry name" value="Olduvai_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46501">
        <property type="entry name" value="MYOMEGALIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46501:SF2">
        <property type="entry name" value="MYOMEGALIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23246">
        <property type="entry name" value="CC_CDK5RAP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07989">
        <property type="entry name" value="Cnn_1N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01148">
        <property type="entry name" value="DUF1220"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51316">
        <property type="entry name" value="ODV"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="5" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Olduvai" evidence="9">
        <location>
            <begin position="1594"/>
            <end position="1685"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="23"/>
            <end position="46"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1035"/>
            <end position="1082"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1354"/>
            <end position="1375"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1435"/>
            <end position="1461"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1629"/>
            <end position="1750"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="2121"/>
            <end position="2162"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="2347"/>
            <end position="2395"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="165"/>
            <end position="228"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="254"/>
            <end position="310"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="396"/>
            <end position="451"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="510"/>
            <end position="676"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="705"/>
            <end position="790"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="843"/>
            <end position="985"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="1103"/>
            <end position="1130"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="1296"/>
            <end position="1323"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="1404"/>
            <end position="1431"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="1940"/>
            <end position="2020"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="2288"/>
            <end position="2322"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="1065"/>
            <end position="1074"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="1356"/>
            <end position="1367"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="1637"/>
            <end position="1654"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="1678"/>
            <end position="1689"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="1693"/>
            <end position="1713"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="1720"/>
            <end position="1735"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="2121"/>
            <end position="2130"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="2141"/>
            <end position="2159"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="2372"/>
            <end position="2384"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004555"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51316"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001062.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2414" mass="269850" checksum="96384B985572460E" modified="2019-05-08" version="1">MLADAGLEYAGLSRGRWLRSRLPRRRTSGARLPEQPMRSPVGAGTVQAPASHGPGCCCCCSREAPGLDPAEMCTPARGDYCDGDGKEMLPGASHVTAVLCDAPEQRNLDISDEDSLRQTPTSGLFETAHHPVPAQTLPGLEQSPLMQTHVLSDLEMGPVVQTQTLRDFEKHLNDLKKENFSLKLRIYFLEERMQQKYEASREDVYKRNIELKVEVESLKQELQERQQTLDKAWAAAENLTSCNEAEFQRRYEERQQEKEHVSELLENKIQLLQEEARLAKSEAERTAALVEAEKGRCLELTKKLKELTKEEEKRGAQVLRDDHCAALAQKDKHIEELIQELSAREQLIELLATEKQNLLQRLEEPREIEVQNLPEDHQQLTQCDPVASGRVSESHAAELQDKIQQYDATNKLLQEKLNEMNFELKSIQEASQRQDRTIQSLNEALKSKESETEELYHVIEGQNETMGKLRDMLHRSQLGHLQTSEGPSPSQQQQHMALLDLQNTLFCTQLEMQKLKRAQWQKEHQLAEARRTTQLLEATVQEEQQLKEVSWKHNQELRGVVQQVQAELQGKVQELWALEGEKCCEIRAQEQRVQRLSQRLAHKEQLLQESWELLQRRQSLEKNPVVTDTMLEKLQQRIKHRDAALERAIDEKFCAVEEKEQELQQLHISVREREHDLERLRSVLSSNEATIHSMESLLKAKTLELEQVTATCQNLQWLKEEVEAKSNCWQKEQEGIIQQLQASLHDRNKEVEELTATLLCKLGPGQSEIAEELCLRLQRKERMLQDLLSDRNRQAGEHETELRELLQAMSIREQRSRELLPGGCGNATATRISKGEDSKYRMERGLLETAAELEKELINAREELELMARKERESRLELSSLQSVVTAQEEELQVQASDIESLTRNIQSKEDLIKDLQMQLVDPEEIPAVERLTQEVLVLREKVATAESRGQEATGNRQQQLLLMLEGLVAEKSRLNEALQAERQLYSSLVKFHAHPDSPERGQTLQAELEGAQALRGRLEESLGRSMEHLRRLETLGAFGGQPGRDDAKDGSTDSIEGEAPPRTVNHQTSQGLPANSPEAKAAQPDLLTLAPLERDSSLQAELLHVRAKNQQLLEQKAKVEGELWELKALMEEAGFSSLSSIRKVLLSLCLETAELKEQLGGSPLSEGWEDKAEARAALCTQNSKLQSSETVVTLLKEQLVLNCPEGESKFPPQLVTGMATEIDRLWTEMGVASGKRPALEGQLQEDPAKRPCPRSQAVELGPAQAPRGMSQQMEACPAGSWQRLVEPRVGPSGQLVQCRQRNQELQDKLAVSEATVRAQAEQLEQYRTLLSEPLVQQDNKQVQVDLQDLGYETCGRSENEADREETPSPECEEPDVFSKAGLMEELSRPGPWGGLCWGQSEDVAVLQQHVRELQLQLQNSNKALQRLQRRTRSFSSTSDYASGAERLHKPVPSATDEDEGWQSDGVGAFCPPALQASRDLEWLVHRVSLLEAQLHGRWAGGVLPEELRSAVWPGKYDSLIQAQARELSHLRQKMREGRSVCHLLTQHLRDTVKSFEELLRGTDIDYYMGQSFREHLAQGGQLAERLSSKLSSRDRSDVEDKTGHEFVALRLSKELREKERMIETLQAKLQERCETPSSSRALSESPRSNSSASFLSDGPEACSDGDATSEYSQLQGDRSEQPSRRLTDVTHPAALASNPSPAAAPHQASAESIRGYPTGPSSQHPPMDSGQTRTGLHFDSLSKPLSVPLTPAPGPSAFLPFGPPPPTPSALLGCCGTPVFSLAEVQQELHMLQKQLGERGPLAAPPVKPLPLTSDFGEASTSRCLHVPHLTPQSCPLQSSTQLDRKAGAAFLESNPLWAQPPMGALYGHLPSGYLASHKLTGADLLEGHLAEIRSLRQRLEESICINDRLREQLEKRLATTTKGNGYPTNIYIQGLESASSLSSETQALREENQSLQLQFAHLSRELEQLRETTRCRLQKAETENRTLAQELAEQRQSGRQLREERLALQENNHRLECTVPLLQQQCQENQQLCQALCAELRIHETLSSSSQAALSACSGDVHRRPPSSSELGALLAEVRTLRRKLLRGIQVNSALRQQLARQQPEGSAAPLFAAHQATPNWQPFQDSSPSPPVRDVGMSSPAPLSPPALLSGPAPLAHQLKEPPVDDALVRRSAPAPAGDAPSSSSASKGGCQVIGHVDDYYTLKQLILEGKTLVHQMASLLRPALAMPSLEPRGTKALDRGSVQQLLSTASALHQLLEQCSSLLTTFWRGALPAAPSPAQPQTAEQAMKDELLALRARLAEQENLLQRAAERLQDTCRLKDNMEHFIVSHLTQTHAVLRKARTNLEKSKPQGSSVKSPPFIGAGEISVGLSQHPSHVIPTNQEHRRKKSCPDTLKWQEDQSWPCFVHLPSS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHM1</accession>
    <name>A0A452GHM1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">WH2 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Early endosome membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Recycling endosome membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the WASH1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHM1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001206.1">
        <property type="protein sequence ID" value="ENSGAGP00000001072.1"/>
        <property type="gene ID" value="ENSGAGG00000000812.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="GOC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031901">
        <property type="term" value="C:early endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055038">
        <property type="term" value="C:recycling endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071203">
        <property type="term" value="C:WASH complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003779">
        <property type="term" value="F:actin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043014">
        <property type="term" value="F:alpha-tubulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043015">
        <property type="term" value="F:gamma-tubulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034314">
        <property type="term" value="P:Arp2/3 complex-mediated actin nucleation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032456">
        <property type="term" value="P:endocytic recycling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006887">
        <property type="term" value="P:exocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042147">
        <property type="term" value="P:retrograde transport, endosome to Golgi"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028290">
        <property type="entry name" value="WASH1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021854">
        <property type="entry name" value="WASH1_WAHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003124">
        <property type="entry name" value="WH2_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23331">
        <property type="entry name" value="CXYORF1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23331:SF5">
        <property type="entry name" value="WAS PROTEIN FAMILY HOMOLOG 2-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11945">
        <property type="entry name" value="WASH_WAHD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51082">
        <property type="entry name" value="WH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="5" id="KW-0967">Endosome</keyword>
    <keyword evidence="6" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0813">Transport</keyword>
    <feature type="domain" description="WH2" evidence="9">
        <location>
            <begin position="335"/>
            <end position="357"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="398"/>
            <end position="445"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="400"/>
            <end position="419"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="8">
        <location>
            <begin position="436"/>
            <end position="445"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004146"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004565"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005602"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51082"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001072.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="445" mass="48314" checksum="3FCA95F6A025D0A5" modified="2019-05-08" version="1">MAAVTQKHFLEGQTYSVPLIQPDLRREEAVHQVADALQYLQKVSSDIFNRISQRVEASRMQLQAIGERVSLAQAKIEKIKGSKKAIKVFSSAKYPAPERLQEYSSIFAGAEDPATQKWPRHKIQSKHRPLDEKSLQEKLKYFPVCVSTKVHQEDDAEEGLGSLPRNISSLSSLLLFNTTENLYKKYVFLDPLAGAVTKTHVTLGTETEEKLFDAPLSITKRGQLDRQVAENYFYVPDLGQVPEIDVPSYLPDLPGIAADLMYSADLGPGIAPSASSSAIPELPTFNTESVENFRPGLQDGVLTSDGVGSSGLTPFCIPAAPVQGAPKEVVNPSTGRATLLESIRQAGGIGKANLRSVKERKLEKKKQKEQEKVITTAQGGDLMSDLFNKLLLRRKGISGKGPAAAGSADAPSAPAGAFARMSDSIPPLPAPQPPLGDEEEDDWES</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHM4</accession>
    <name>A0A452GHM4_9SAUR</name>
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        <recommendedName>
            <fullName evidence="1">FHOD1 N-terminal GTPase-binding domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
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        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
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        <property type="gene ID" value="ENSGAGG00000000900.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="project" value="TreeGrafter"/>
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        <property type="entry name" value="Leucine-rich Repeat Variant"/>
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    </dbReference>
    <dbReference type="PANTHER" id="PTHR45920:SF3">
        <property type="entry name" value="FH1_FH2 DOMAIN-CONTAINING PROTEIN 3"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Formin_GBD_N"/>
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    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="FHOD1 N-terminal GTPase-binding" evidence="1">
        <location>
            <begin position="15"/>
            <end position="97"/>
        </location>
    </feature>
    <evidence type="ECO:0000259" key="1">
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            <dbReference type="Pfam" id="PF18382"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001126.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="121" mass="13597" checksum="B19901EB3B61AB83" modified="2019-05-08" version="1">RGGGDDPGVLAALKQVQFLDDTDPFNSTNFPEPTRPPLYTFREDIALVTQLAGVHRLLRAPHKLDDCTLQLSHNGTYLDLESTLAEQRDELEGFQEDKVDREVLFTPSHKGPPDEISRQQV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHM7</accession>
    <name>A0A452GHM7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Olduvai domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Golgi apparatus</location>
        </subcellularLocation>
    </comment>
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        <property type="protein sequence ID" value="ENSGAGP00000001115.1"/>
        <property type="gene ID" value="ENSGAGG00000000835.1"/>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="project" value="TreeGrafter"/>
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        <property type="project" value="UniProtKB-SubCell"/>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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        <property type="project" value="TreeGrafter"/>
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        <property type="term" value="P:positive regulation of microtubule nucleation"/>
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        <property type="project" value="TreeGrafter"/>
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        <property type="term" value="P:regulation of Golgi organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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        <property type="entry name" value="Cnn_1N"/>
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        <property type="entry name" value="MT-associated_AKAP9-binding"/>
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        <property type="entry name" value="Olduvai_dom"/>
    </dbReference>
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        <property type="entry name" value="MYOMEGALIN"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="MYOMEGALIN"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="CC_CDK5RAP2"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Cnn_1N"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="DUF1220"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="ODV"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="5" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Olduvai" evidence="9">
        <location>
            <begin position="1574"/>
            <end position="1665"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
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            <end position="1062"/>
        </location>
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    <feature type="region of interest" description="Disordered" evidence="8">
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            <end position="1355"/>
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        <location>
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            <end position="1441"/>
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    </feature>
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            <end position="1730"/>
        </location>
    </feature>
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        <location>
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        </location>
    </feature>
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            <begin position="2327"/>
            <end position="2375"/>
        </location>
    </feature>
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        <location>
            <begin position="96"/>
            <end position="159"/>
        </location>
    </feature>
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        <location>
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        </location>
    </feature>
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            <end position="1303"/>
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        </location>
    </feature>
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        <location>
            <begin position="1920"/>
            <end position="2000"/>
        </location>
    </feature>
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            <begin position="2268"/>
            <end position="2302"/>
        </location>
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            <begin position="1045"/>
            <end position="1054"/>
        </location>
    </feature>
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            <begin position="1336"/>
            <end position="1347"/>
        </location>
    </feature>
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        <location>
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        </location>
    </feature>
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            <end position="2139"/>
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        <location>
            <begin position="2352"/>
            <end position="2364"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00004555"/>
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    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="PROSITE" id="PS51316"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001115.1"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2394" mass="268163" checksum="43CCA140BF6C7601" modified="2019-05-08" version="1">MEGAATRKRDYCDGDGKEMLPGASHVTAVLCDAPEQRNLDISDEDSLRQTPTSGLFETAHHPVPAQTLPGLEQSPLMQTHVLSDLEMGPVVQTQTLRDFEKHLNDLKKENFSLKLRIYFLEERMQQKYEASREDVYKRNIELKVEVESLKQELQERQQTLDKAWAAAENLTSCNEAEFQRRYEERQQEKEHVSELLENKIQLLQEEARLAKSEAERTAALVEAEKGRCLELTKKLKELTKEEEKRGAQVLRDDHCAALAQKDKHIEELIQELSAREQLIELLATEKQNLLQRLEEPREIEVQNLPEDHQQLTQCDPVASGRVSESHAAELQDKIQQYDATNKLLQEKLNEMNFELKSIQEASQRQDRTIQSLNEALKSKESETEELYHVIEGQNETMGKLRDMLHRSQLGHLQTSEGPSPSQQQQHMALLDLQNTLFCTQLEMQKLKRAQWQKEHQLAEARRTTQLLEATVQEEQQLKEVSWKHNQELRGVVQQVQAELQGKVQELWALEGEKCCEIRAQEQRVQRLSQRLAHKEQLLQESWELLQRRQSLEKNPVVTDTMLEKLQQRIKHRDAALERAIDEKFCAVEEKEQELQQLHISVREREHDLERLRSVLSSNEATIHSMESLLKAKTLELEQVTATCQNLQWLKEEVEAKSNCWQKEQEGIIQQLQASLHDRNKEVEELTATLLCKLGPGQSEIAEELCLRLQRKERMLQDLLSDRNRQAGEHETELRELLQAMSIREQRSRVTAEKMVQALAEKSDEVQFLRQRLVGNEPGKNPVADGRLLLQKDKQPFQELLPGGCGNATATRISKGEDSKYRMERGLLETAAELEKELINAREELELMARKERESRLELSSLQSVVTAQEEELQVQASDIESLTRNIQSKEDLIKDLQMQLVDPEEIPAVERLTQEVLVLREKVATAESRGQEATGNRQQQLLLMLEGLVAEKSRLNEALQAERQLYSSLVKFHAHPDSPERGQTLQAELEGAQALRGRLEESLGRSMEHLRRLETLGAFGGQPGRDDAKDGSTDSIEGEAPPRTVNHQTSQGLPANSPEAKAAQPDLLTLAPLERDSSLQAELLHVRAKNQQLLEQKAKVEGELWELKALMEEAGFSSLSSIRKVLLSLCLETAELKEQLGGSPLSEGWEDKAEARAALCTQNSKLQSSETVVTLLKEQLVLNCPEGESKFPPQLVTGMATEIDRLWTEMGVASGKRPALEGQLQEDPAKRPCPRSQAVELGPAQAPRGMSQQMEACPAGSWQRLVEPRVGPSGQLVQCRQRNQELQDKLAVSEATVRAQAEQLEQYRTLLSEPLVQQDNKQVQVDLQDLGYETCGRSENEADREETPSPECEEPDVFSKAGLMEELSRPGPWGGLCWGQSEDVAVLQQHVRELQLQLQNSNKALQRLQRRTRSFSSTSDYASGAERLHKPVPSATDEDEGWQSDGVGAFCPPALQASRDLEWLVHRVSLLEAQLHGRWAGGVLPEELRSAVWPGKYDSLIQAQARELSHLRQKMREGRSVCHLLTQHLRDTVKSFEELLRGTDIDYYMGQSFREHLAQGGQLAERLSSKLSSRDRSDVEDKTGHEFVALRLSKELREKERMIETLQAKLQERCETPSSSRALSESPRSNSSASFLSDGPEACSDGDATSEYSQLQGDRSEQPSRRLTDVTHPAALASNPSPAAAPHQASAESIRGYPTGPSSQHPPMDSGQTRTGLHFDSLSKPLSVPLTPAPGPSAFLPFGPPPPTPSALLGCCGTPVFSLAEVQQELHMLQKQLGERGPLAAPPVKPLPLTSDFGEASTSRCLHVPHLTPQSCPLQSSTQLDRKAGAAFLESNPLWAQPPMGALYGHLPSGYLASHKLTGADLLEGHLAEIRSLRQRLEESICINDRLREQLEKRLATTTKGNGYPTNIYIQGLESASSLSSETQALREENQSLQLQFAHLSRELEQLRETTRCRLQKAETENRTLAQELAEQRQSGRQLREERLALQENNHRLECTVPLLQQQCQENQQLCQALCAELRIHETLSSSSQAALSACSGDVHRRPPSSSELGALLAEVRTLRRKLLRGIQVNSALRQQLARQQPEGSAAPLFAAHQATPNWQPFQDSSPSPPVRDVGMSSPAPLSPPALLSGPAPLAHQLKEPPVDDALVRRSAPAPAGDAPSSSSASKGGCQVIGHVDDYYTLKQLILEGKTLVHQMASLLRPALAMPSLEPRGTKALDRGSVQQLLSTASALHQLLEQCSSLLTTFWRGALPAAPSPAQPQTAEQAMKDELLALRARLAEQENLLQRAAERLQDTCRLKDNMEHFIVSHLTQTHAVLRKARTNLEKSKPQGSSVKSPPFIGAGEISVGLSQHPSHVIPTNQEHRRKKSCPDTLKWQEDQSWPCFVHLPSS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHM8</accession>
    <name>A0A452GHM8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Arrestin C-terminal-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the arrestin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHM8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001259.1">
        <property type="protein sequence ID" value="ENSGAGP00000001124.1"/>
        <property type="gene ID" value="ENSGAGG00000000891.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005768">
        <property type="term" value="C:endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.640">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014752">
        <property type="entry name" value="Arrestin-like_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011021">
        <property type="entry name" value="Arrestin-like_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011022">
        <property type="entry name" value="Arrestin_C-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050357">
        <property type="entry name" value="Arrestin_domain-protein"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014756">
        <property type="entry name" value="Ig_E-set"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11188">
        <property type="entry name" value="ARRESTIN DOMAIN CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11188:SF172">
        <property type="entry name" value="ARRESTIN DOMAIN-CONTAINING PROTEIN 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02752">
        <property type="entry name" value="Arrestin_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00339">
        <property type="entry name" value="Arrestin_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01017">
        <property type="entry name" value="Arrestin_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81296">
        <property type="entry name" value="E set domains"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Arrestin C-terminal-like" evidence="2">
        <location>
            <begin position="168"/>
            <end position="305"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00005298"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="SMART" id="SM01017"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001124.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="310" mass="35012" checksum="A980D3647F1B03A0" modified="2019-05-08" version="1">MSVVKSISLVLPEEKVYLAGSSVAGQVILTLNNSLINPTVKVELVGRGYLQWNEETNADKDYSRNIICNNKADYVNKTKTFYIEGNWLASGTHIFDFHFNLPDRLPSTFTSNIARVSYFLKASCATRELILAKQKKYLLVQGTSSSLPTNKIESQVSVRVQSYHCCFKHGLVILRISLEKNTFAPGEQIIFMTEIDNQTGRSIRKVVFALYSIILYLGYTVRSEQRSLEDRNEVMRLESRTKAAPFEVTRITSMFSLPKLLSVSSMLIANEIMEVRYELIGTVYLPWSMNTIVAKVPIIIRTTAAGIPED</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHM9</accession>
    <name>A0A452GHM9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Protein AMBP</fullName>
        </recommendedName>
    </protein>
    <organism evidence="34 35">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="35" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="34" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="34" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="28">Kunitz-type serine protease inhibitor. Has high catalytic efficiency for F10/blood coagulation factor Xa and may act as an anticoagulant by inhibiting prothrombin activation. Inhibits trypsin and mast cell CMA1/chymase and tryptase proteases.</text>
    </comment>
    <comment type="subunit">
        <text evidence="30">I-alpha-I plasma protease inhibitors are assembled from one or two heavy chains (HC) and one light chain, bikunin. Inter-alpha-inhibitor (I-alpha-I) is composed of ITIH1/HC1, ITIH2/HC2 and bikunin, and pre-alpha-inhibitor (P-alpha-I) of ITIH3/HC3 and bikunin. Interacts with TNFAIP6 (via Link domain).</text>
    </comment>
    <comment type="subunit">
        <text evidence="29">Monomer. Also occurs as a complex with tryptase in mast cells.</text>
    </comment>
    <comment type="subunit">
        <text evidence="31">Monomer. Homodimer. In plasma, it occurs as a monomer or dimer and in covalently-linked complexes with immunoglobulin A (IgA), ALB/albumin and F2/prothrombin. Chromophore-bound alpha-1-microglobulin interacts with the constant region of immunoglobulin A. Chromophore-bound alpha-1-microglobulin interacts with ALB with molar ratio 2:1 and 1:1; this interaction does not prevent fatty acid binding to ALB. Interacts with F2/prothrombin (via N-terminus) with molar ratio 2:1 and 1:1; this interaction does not prevent the activation of prothrombin to thrombin. Interacts with NDUFAB1, a subunit of mitochondrial complex I. Interacts with FN1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Peripheral membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cytoplasm</location>
            <location evidence="4">Cytosol</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Endoplasmic reticulum</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="6">Mitochondrion inner membrane</location>
            <topology evidence="6">Peripheral membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="5">Nucleus membrane</location>
            <topology evidence="5">Peripheral membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Secreted</location>
            <location evidence="3">Extracellular space</location>
            <location evidence="3">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="7">In the N-terminal section; belongs to the calycin superfamily. Lipocalin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHM9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001266.1">
        <property type="protein sequence ID" value="ENSGAGP00000001131.1"/>
        <property type="gene ID" value="ENSGAGG00000000433.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005783">
        <property type="term" value="C:endoplasmic reticulum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005743">
        <property type="term" value="C:mitochondrial inner membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031965">
        <property type="term" value="C:nuclear membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016491">
        <property type="term" value="F:oxidoreductase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004867">
        <property type="term" value="F:serine-type endopeptidase inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd22596">
        <property type="entry name" value="Kunitz_bikunin_1-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd22597">
        <property type="entry name" value="Kunitz_bikunin_2-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.128.20:FF:000007">
        <property type="entry name" value="Alpha-1-microglobulin/bikunin precursor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.410.10:FF:000010">
        <property type="entry name" value="Alpha1-microglobulin/bikunin (AMBP)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.128.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.410.10">
        <property type="entry name" value="Pancreatic trypsin inhibitor Kunitz domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002968">
        <property type="entry name" value="A1-microglobln"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029856">
        <property type="entry name" value="AMBP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012674">
        <property type="entry name" value="Calycin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002223">
        <property type="entry name" value="Kunitz_BPTI"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036880">
        <property type="entry name" value="Kunitz_BPTI_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000566">
        <property type="entry name" value="Lipocln_cytosolic_FA-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020901">
        <property type="entry name" value="Prtase_inh_Kunz-CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46676">
        <property type="entry name" value="PROTEIN AMBP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46676:SF1">
        <property type="entry name" value="PROTEIN AMBP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00014">
        <property type="entry name" value="Kunitz_BPTI"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00061">
        <property type="entry name" value="Lipocalin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01215">
        <property type="entry name" value="A1MCGLOBULIN"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00759">
        <property type="entry name" value="BASICPTASE"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00179">
        <property type="entry name" value="LIPOCALIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00131">
        <property type="entry name" value="KU"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57362">
        <property type="entry name" value="BPTI-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50814">
        <property type="entry name" value="Lipocalins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00280">
        <property type="entry name" value="BPTI_KUNITZ_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50279">
        <property type="entry name" value="BPTI_KUNITZ_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="21" id="KW-0157">Chromophore</keyword>
    <keyword evidence="13" id="KW-0165">Cleavage on pair of basic residues</keyword>
    <keyword evidence="10" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="25" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="18" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="12" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="26" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="24" id="KW-0472">Membrane</keyword>
    <keyword evidence="23" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="17" id="KW-0999">Mitochondrion inner membrane</keyword>
    <keyword evidence="27" id="KW-0539">Nucleus</keyword>
    <keyword evidence="22" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="14" id="KW-0646">Protease inhibitor</keyword>
    <keyword evidence="20" id="KW-0654">Proteoglycan</keyword>
    <keyword evidence="35" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="16" id="KW-0677">Repeat</keyword>
    <keyword evidence="11" id="KW-0964">Secreted</keyword>
    <keyword evidence="19" id="KW-0722">Serine protease inhibitor</keyword>
    <keyword evidence="15 32" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="32">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
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        <location>
            <begin position="21"/>
            <end position="333"/>
        </location>
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    <feature type="domain" description="BPTI/Kunitz inhibitor" evidence="33">
        <location>
            <begin position="212"/>
            <end position="262"/>
        </location>
    </feature>
    <feature type="domain" description="BPTI/Kunitz inhibitor" evidence="33">
        <location>
            <begin position="268"/>
            <end position="318"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004202"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00004240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004514"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00004637"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00008238"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00018905"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022685"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022690"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022900"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022974"/>
        </source>
    </evidence>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00029383"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00029474"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="ARBA" id="ARBA00046983"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00047015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="32">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="33">
        <source>
            <dbReference type="PROSITE" id="PS50279"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="34">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001131.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="35">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="333" mass="37354" checksum="0E38B35893C477FD" modified="2019-05-08" version="1">MKFQEFFFLLFLAYLPTARGSPTGSQVQDIQVQENFEPERVYGKWYDIATGTTCKWMKQYKDKFNMGTLVLGPGMTSNQISTTSTRLRQGVCTQVSGEYQKTDIPGKYTYYNPSWDVTINSYVVHTNYEEYAIILMQKKSSFGLTITAKLYGRSPELREGLIVDFRQFAQEMGIPEDSIFIMINKGNPAMMRVTAGPPPPLGSTLLMCSDFCRLNKDVGPCMGMNVRFFYNSSSMACETFHYGGCLGNGNNFLSEKKCLQTCRTEAACRLPIVPGPCQQPVILWAFDATQGKCTSFNYGGCQGNGNKFYTEKECREYCGVPADGMVRPVLSQS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHP0</accession>
    <name>A0A452GHP0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Solute carrier family 6 member 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHP0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001265.1">
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        <property type="gene ID" value="ENSGAGG00000000898.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="P:amino acid transport"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="P:sodium ion transmembrane transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000175">
        <property type="entry name" value="Na/ntran_symport"/>
    </dbReference>
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        <property type="entry name" value="SNS_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11616">
        <property type="entry name" value="SODIUM/CHLORIDE DEPENDENT TRANSPORTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11616:SF316">
        <property type="entry name" value="SOLUTE CARRIER FAMILY 6 MEMBER 1"/>
        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5 6" id="KW-0472">Membrane</keyword>
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    <keyword evidence="4 6" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="2" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
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            <end position="57"/>
        </location>
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    <feature type="transmembrane region" description="Helical" evidence="6">
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    <feature type="transmembrane region" description="Helical" evidence="6">
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            <end position="126"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
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            <end position="167"/>
        </location>
    </feature>
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        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001130.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="9">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="208" mass="23575" checksum="11230D797EE39EF5" modified="2019-05-08" version="1">GADFHGAHIKILASPGLAFLAYPQVVTQLPMSPLWSILFFSMLMMLGLDSQGGIYVFKLFDYYSASGMCLLFLVFFETISISWCYGVNRFYRNIEEMIGYQPCSWWKLCWAFFTPLVCVGVFFFSVVEMTPLTLGKYVYPSWGQGIGWLMALSSMVLIPGYMLYYFFTSKGTIRQVKATFQLLPLLDYLSRGNFPPTSLNGRVSDAAV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHP5</accession>
    <name>A0A452GHP5_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="10">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GHP5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001271.1">
        <property type="protein sequence ID" value="ENSGAGP00000001135.1"/>
        <property type="gene ID" value="ENSGAGG00000000203.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031594">
        <property type="term" value="C:neuromuscular junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098831">
        <property type="term" value="C:presynaptic active zone cytoplasmic component"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042734">
        <property type="term" value="C:presynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043195">
        <property type="term" value="C:terminal bouton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019992">
        <property type="term" value="F:diacylglycerol binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005543">
        <property type="term" value="F:phospholipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017075">
        <property type="term" value="F:syntaxin-1 binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061789">
        <property type="term" value="P:dense core granule priming"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0099525">
        <property type="term" value="P:presynaptic dense core vesicle exocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035249">
        <property type="term" value="P:synaptic transmission, glutamatergic"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016081">
        <property type="term" value="P:synaptic vesicle docking"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016082">
        <property type="term" value="P:synaptic vesicle priming"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd04027">
        <property type="entry name" value="C2B_Munc13"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd08395">
        <property type="entry name" value="C2C_Munc13"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.357.50:FF:000001">
        <property type="entry name" value="Protein unc-13 homolog B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000002">
        <property type="entry name" value="Protein unc-13 homolog B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000014">
        <property type="entry name" value="protein unc-13 homolog B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.357.50">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.1100">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000008">
        <property type="entry name" value="C2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010439">
        <property type="entry name" value="MUN_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014770">
        <property type="entry name" value="Munc13_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014772">
        <property type="entry name" value="Munc13_dom-2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027080">
        <property type="entry name" value="Unc-13"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037302">
        <property type="entry name" value="Unc-13_C2B"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10480">
        <property type="entry name" value="PROTEIN UNC-13 HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10480:SF2">
        <property type="entry name" value="PROTEIN UNC-13 HOMOLOG C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00168">
        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06292">
        <property type="entry name" value="MUN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00360">
        <property type="entry name" value="C2DOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00239">
        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM01145">
        <property type="entry name" value="DUF1041"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49562">
        <property type="entry name" value="C2 domain (Calcium/lipid-binding domain, CaLB)"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50004">
        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51258">
        <property type="entry name" value="MHD1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51259">
        <property type="entry name" value="MHD2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0106">Calcium</keyword>
    <keyword evidence="1" id="KW-0268">Exocytosis</keyword>
    <keyword evidence="2" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <keyword evidence="4" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2" evidence="7">
        <location>
            <begin position="102"/>
            <end position="226"/>
        </location>
    </feature>
    <feature type="domain" description="MHD1" evidence="8">
        <location>
            <begin position="532"/>
            <end position="675"/>
        </location>
    </feature>
    <feature type="domain" description="MHD2" evidence="9">
        <location>
            <begin position="781"/>
            <end position="908"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="7">
        <location>
            <begin position="922"/>
            <end position="1048"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51258"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51259"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001135.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1093" mass="125661" checksum="CB758CDB60A2F190" modified="2019-05-08" version="1">YSELKNKPIHYQTPKTLLCYSSTICSDEAASQQYFHQIERRGLYNGVVRAAEKSSKHGAEDKTQNIIIAMKERMKIRERNRPEVFEVIQEMFQISKEDFVQYTKAAKQCVLDGTSKWSAKITITVVCAQGLQAKDKTGSSDPYVTVQVGKTKRRTKTIFGNLNPVWDEKFYFECHNSTDRIKVRVWDEDDDIKSRVKQHFKKESDDFLGQTIIEVRTLSGEMDVWYNLEKRTDKSAVSGAIRLKINVEIKGEEKVAPYHVQYTYLHENLFHYLTEVKSNGSVKIPEVRGDEAWKVYFDAAAQEIVDEFAMRYGIEYIYQAMTHFSCLSSKYMCSGVPAVMSTLLANINAFYAHTIATTNVSASDRFAATNFGREKFIKLLDQLHNSLRIDLSKYRDNFPASNSERLQDLKSTVDLLTSITFFRMKVLELQSPPRASMVVKDCVRACLDSTYKYIFDNCYELYSQLMDQKQEVPREEQGPTTKNLDFWAQLITLMVTIIDEDKTAYTPVLNQFPQELNMGKVSAEIMWTLFAQDMKYALEEHEKQRLCKSTDYMNLHFKVKWFYNEYVRELPALQDAVPEYSLWFEPFVIQWLDENEDVSMEFLHGALERDKKDGFQQTSDHALFSCSVVDVFTQLNQSFEIIKKLECPNPEALSHLMRRFAKTINKVLLQYAAVISNDFNSYCDKEMLPCILMNNIQQLRVQLEKMFESMGGKELDPDASVVLKELQVKLSNVLDELSIAYGASFQLVVEECVRQMSFELNQMRGNGNISANKNNAAIDAEIVLRPLMDFLDKILSLSAKICEKTVLKRVLKELWKSVLNKIEKQIVLPPLTDQTVSVQDHVTREEARSLTLRQCAIMEVALSTIKQYFHAGGNGLKKNFLEKSPDLQSLRYALSLYTQTTDALIKKFIATQTSQSHSTNNTVGEISVQVDVFTHPGTGEHRVTVKVVAINNLNWQTTAMFRPFVEVCILGPNLSDKKRKHGTKTKSNTWSPKYNETFHILSNEEGLRAYELHLSVKDYCFAREDRIVGMTVIQLQNTAEKGSCASWYPLLKNIFMDDTGLTILRILSQRTNDEVAKEFVRLKSETRSTEETV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHP7</accession>
    <name>A0A452GHP7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Protein AMBP</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">In the N-terminal section; belongs to the calycin superfamily. Lipocalin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHP7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001283.1">
        <property type="protein sequence ID" value="ENSGAGP00000001144.1"/>
        <property type="gene ID" value="ENSGAGG00000000433.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004867">
        <property type="term" value="F:serine-type endopeptidase inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd22596">
        <property type="entry name" value="Kunitz_bikunin_1-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd22597">
        <property type="entry name" value="Kunitz_bikunin_2-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.410.10:FF:000010">
        <property type="entry name" value="Alpha1-microglobulin/bikunin (AMBP)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.128.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.410.10">
        <property type="entry name" value="Pancreatic trypsin inhibitor Kunitz domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002968">
        <property type="entry name" value="A1-microglobln"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029856">
        <property type="entry name" value="AMBP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012674">
        <property type="entry name" value="Calycin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002223">
        <property type="entry name" value="Kunitz_BPTI"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036880">
        <property type="entry name" value="Kunitz_BPTI_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000566">
        <property type="entry name" value="Lipocln_cytosolic_FA-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020901">
        <property type="entry name" value="Prtase_inh_Kunz-CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46676">
        <property type="entry name" value="PROTEIN AMBP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46676:SF1">
        <property type="entry name" value="PROTEIN AMBP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00014">
        <property type="entry name" value="Kunitz_BPTI"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00061">
        <property type="entry name" value="Lipocalin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01215">
        <property type="entry name" value="A1MCGLOBULIN"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00759">
        <property type="entry name" value="BASICPTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00131">
        <property type="entry name" value="KU"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57362">
        <property type="entry name" value="BPTI-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50814">
        <property type="entry name" value="Lipocalins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00280">
        <property type="entry name" value="BPTI_KUNITZ_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50279">
        <property type="entry name" value="BPTI_KUNITZ_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0157">Chromophore</keyword>
    <keyword evidence="5" id="KW-0165">Cleavage on pair of basic residues</keyword>
    <keyword evidence="11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6" id="KW-0646">Protease inhibitor</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="9" id="KW-0722">Serine protease inhibitor</keyword>
    <keyword evidence="7 13" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="13">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019194159" description="Protein AMBP" evidence="13">
        <location>
            <begin position="21"/>
            <end position="325"/>
        </location>
    </feature>
    <feature type="domain" description="BPTI/Kunitz inhibitor" evidence="14">
        <location>
            <begin position="204"/>
            <end position="254"/>
        </location>
    </feature>
    <feature type="domain" description="BPTI/Kunitz inhibitor" evidence="14">
        <location>
            <begin position="260"/>
            <end position="310"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008238"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00018905"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022685"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022690"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50279"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001144.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="325" mass="35887" checksum="8C146D3D503D5C85" modified="2019-05-08" version="1">MKFQEFFFLLFLAYLPTARGSPTGSQVQDIQVQENFEPERVYGKWYDIATGTTCKWMKQYKDKFNMGTLVLGPGMTSNQISTTITQNTGTQLGYTVHPGDSAAAVSSWDVTINSYVVHTNYEEYAIILMQKKSSFGLTITAKLYGTCLNLLRGLGVRLWPRGDGSEGRGHGHVAPGCHHHMPCISVALKRNSLGSTLLMCSDFCRLNKDVGPCMGMNVRFFYNSSSMACETFHYGGCLGNGNNFLSEKKCLQTCRTEAACRLPIVPGPCQQPVILWAFDATQGKCTSFNYGGCQGNGNKFYTEKECREYCGVPADGMVRPVLSQS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHQ3</accession>
    <name>A0A452GHQ3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Paired domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHQ3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001296.1">
        <property type="protein sequence ID" value="ENSGAGP00000001157.1"/>
        <property type="gene ID" value="ENSGAGG00000000896.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00131">
        <property type="entry name" value="PAX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.10:FF:000013">
        <property type="entry name" value="Paired box 8 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.10:FF:000003">
        <property type="entry name" value="Paired box protein Pax-6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.10">
        <property type="entry name" value="Winged helix-like DNA-binding domain superfamily/Winged helix DNA-binding domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009057">
        <property type="entry name" value="Homeodomain-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043182">
        <property type="entry name" value="PAIRED_DNA-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001523">
        <property type="entry name" value="Paired_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022130">
        <property type="entry name" value="Pax2_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043565">
        <property type="entry name" value="PAX_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036388">
        <property type="entry name" value="WH-like_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45636:SF20">
        <property type="entry name" value="PAIRED BOX PROTEIN PAX-5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45636">
        <property type="entry name" value="PAIRED BOX PROTEIN PAX-6-RELATED-RELATED"/>
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    <sequence length="367" mass="39991" checksum="E9392233D18BCCD6" modified="2019-05-08" version="1">AEIHLSSNPSKRARHGGVNQLGGVFVNGRPLPDVVRQRIVELAHQGVRPCDISRQLRVSHGCVSKILGRYYETGSIKPGVIGGSKPKVATPKVVEKIAEYKRQNPTMFAWEIRDRLLAERVCDNDTVPSVSSINRIIRTKVQQPPNQQVPASSHSIASTGSVTQVSTVSTDSAGSSYSISGILGITSPSAESNKRKRDEGIQESPVPNGHSLPGRDFLRKQMRGDLFTQQQLEVLDRVFERQHYSDIFTTTEPIKPEQTTEYSAMASLAGGLDDMKANLTSPTSAELGASVPGPQSYPIVTGSEFSGSPYSHPQYSTYNDSWRFPNPGLLGEFSYPSFYRNSLLTAAGACPVGGISPHWMLGQLWPL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHR4</accession>
    <name>A0A452GHR4_9SAUR</name>
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        <recommendedName>
            <fullName evidence="8">ELK domain-containing protein</fullName>
        </recommendedName>
    </protein>
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        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
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            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
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        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
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                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
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            <dbReference type="PubMed" id="28562605"/>
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        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
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    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
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        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
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        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001118.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="4614" mass="538349" checksum="850AA6DC11EF024C" modified="2019-05-08" version="1">[encoded data, 4614 characters]</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHS3</accession>
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        <submittedName>
            <fullName evidence="16">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
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        <name type="scientific">Gopherus agassizii</name>
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            <taxon>Archelosauria</taxon>
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            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
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                <person name="Cornelius J.A."/>
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                <person name="Edwards T."/>
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                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
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    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
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                <consortium name="Ensembl"/>
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        <scope>IDENTIFICATION</scope>
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                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
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            <text>O-phospho-L-tyrosyl-[protein] + H2O = L-tyrosyl-[protein] + phosphate</text>
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            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
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            <dbReference type="ChEBI" id="CHEBI:61978"/>
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        </reaction>
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        </subcellularLocation>
    </comment>
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        <text evidence="4">Belongs to the protein-tyrosine phosphatase family.</text>
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    </dbReference>
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        <property type="entry name" value="Tyr_Pase_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020422">
        <property type="entry name" value="TYR_PHOSPHATASE_DUAL_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23339">
        <property type="entry name" value="TYROSINE SPECIFIC PROTEIN PHOSPHATASE AND DUAL SPECIFICITY PROTEIN PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22784">
        <property type="entry name" value="PTP-SAK"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="PTPc_motif"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50054">
        <property type="entry name" value="TYR_PHOSPHATASE_DUAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="8" id="KW-0967">Endosome</keyword>
    <keyword evidence="9" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="12" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="10" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-0488">Methylation</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="14">
        <location>
            <begin position="1"/>
            <end position="145"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="15">
        <location>
            <begin position="66"/>
            <end position="132"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004177"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
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    </evidence>
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    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00009580"/>
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    </evidence>
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        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
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    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00051722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001179.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="154" mass="17528" checksum="AB98DC0F2FF9C705" modified="2019-05-08" version="1">MRFLITHNPTNATLNKFTEELKKYGVTTLVRVCDATYDKAPVEKEGIQVLDWPFDDGAPPPNQIVDDWLNLLKTKFREEPGCCVAVHCVAGLGRAPVLVALALIECGMKYEDAVQFIRQKRRGAFNSKQLLYLEKYRPKMRLRSKDTNGHCCVQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHS8</accession>
    <name>A0A452GHS8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Metaxin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 8">Mitochondrion outer membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 8">Belongs to the metaxin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHS8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001330.1">
        <property type="protein sequence ID" value="ENSGAGP00000001184.1"/>
        <property type="gene ID" value="ENSGAGG00000000954.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="term" value="C:SAM complex"/>
        <property type="evidence" value="ECO:0007669"/>
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    </dbReference>
    <dbReference type="GO" id="GO:0007005">
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        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
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        <property type="entry name" value="GST_C_Metaxin1_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd03078">
        <property type="entry name" value="GST_N_Metaxin1_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036282">
        <property type="entry name" value="Glutathione-S-Trfase_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040079">
        <property type="entry name" value="Glutathione_S-Trfase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017410">
        <property type="entry name" value="Metaxin1/3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033468">
        <property type="entry name" value="Metaxin_GST"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050931">
        <property type="entry name" value="Mito_Protein_Transport_Metaxin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019564">
        <property type="entry name" value="Sam37/metaxin_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12289">
        <property type="entry name" value="METAXIN RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12289:SF30">
        <property type="entry name" value="METAXIN-3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17171">
        <property type="entry name" value="GST_C_6"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDS00019">
        <property type="entry name" value="Glutathione_Transferase_(cytos"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDG01180">
        <property type="entry name" value="SUF1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="GST C-terminal domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="4 8" id="KW-1000">Mitochondrion outer membrane</keyword>
    <keyword evidence="5" id="KW-0653">Protein transport</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <feature type="domain" description="Mitochondrial outer membrane transport complex Sam37/metaxin N-terminal" evidence="9">
        <location>
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            <end position="142"/>
        </location>
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            <end position="236"/>
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        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
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        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PIRNR" id="PIRNR038150"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF10568"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="Pfam" id="PF17171"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001184.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="252" mass="28358" checksum="9B1D4C6AFD3D931E" modified="2019-05-08" version="1">MASPMELSCWGGDWGLPSVHTESLIVMAYARFSGAPVKVNAIDNSWRAPKGDVPVLISEDTVIAQPAKILTFLRKQKYNADYELSAKQGADTLAYIALLEEKLLPAVLHTFWVEAENYCTVTKPWFASRIPFPLSLYLPGKMSREALNRILLTKGEPPLYSLSEVEAQIYRDGKECLNLLSNRLGTSQFFFGNTPTTLDAFVFGFLAPLYKVHFPRVQLQEHLKQLSNLCRFCDDILSGYFRLNVTDGRQPS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GI11</accession>
    <name>A0A452GI11_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">ELK domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Microtubule organizing center</location>
            <location evidence="1">Centrosome</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000001360.1">
        <property type="protein sequence ID" value="ENSGAGP00000001212.1"/>
        <property type="gene ID" value="ENSGAGG00000000844.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060090">
        <property type="term" value="F:molecular adaptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028745">
        <property type="entry name" value="AKAP9/Pericentrin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005539">
        <property type="entry name" value="ELK_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR44981:SF3">
        <property type="entry name" value="PERICENTRIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR44981">
        <property type="entry name" value="PERICENTRIN-LIKE PROTEIN, ISOFORM F"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01188">
        <property type="entry name" value="ELK"/>
        <property type="match status" value="10"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3 5" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="ELK" evidence="7">
        <location>
            <begin position="369"/>
            <end position="390"/>
        </location>
    </feature>
    <feature type="domain" description="ELK" evidence="7">
        <location>
            <begin position="1138"/>
            <end position="1159"/>
        </location>
    </feature>
    <feature type="domain" description="ELK" evidence="7">
        <location>
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            <end position="1315"/>
        </location>
    </feature>
    <feature type="domain" description="ELK" evidence="7">
        <location>
            <begin position="1352"/>
            <end position="1373"/>
        </location>
    </feature>
    <feature type="domain" description="ELK" evidence="7">
        <location>
            <begin position="1420"/>
            <end position="1443"/>
        </location>
    </feature>
    <feature type="domain" description="ELK" evidence="7">
        <location>
            <begin position="1557"/>
            <end position="1578"/>
        </location>
    </feature>
    <feature type="domain" description="ELK" evidence="7">
        <location>
            <begin position="1586"/>
            <end position="1604"/>
        </location>
    </feature>
    <feature type="domain" description="ELK" evidence="7">
        <location>
            <begin position="1727"/>
            <end position="1746"/>
        </location>
    </feature>
    <feature type="domain" description="ELK" evidence="7">
        <location>
            <begin position="1880"/>
            <end position="1902"/>
        </location>
    </feature>
    <feature type="domain" description="ELK" evidence="7">
        <location>
            <begin position="3841"/>
            <end position="3859"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="1"/>
            <end position="47"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="69"/>
            <end position="102"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="4138"/>
            <end position="4163"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="5">
        <location>
            <begin position="117"/>
            <end position="195"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="5">
        <location>
            <begin position="220"/>
            <end position="275"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="5">
        <location>
            <begin position="368"/>
            <end position="395"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="5">
        <location>
            <begin position="423"/>
            <end position="538"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="5">
        <location>
            <begin position="732"/>
            <end position="804"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="5">
        <location>
            <begin position="828"/>
            <end position="1164"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="5">
        <location>
            <begin position="1200"/>
            <end position="1265"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="5">
        <location>
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        <location>
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            <end position="1805"/>
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        <location>
            <begin position="2018"/>
            <end position="2119"/>
        </location>
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        <location>
            <begin position="2162"/>
            <end position="2286"/>
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    <feature type="coiled-coil region" evidence="5">
        <location>
            <begin position="2492"/>
            <end position="2519"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="5">
        <location>
            <begin position="2576"/>
            <end position="2760"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="5">
        <location>
            <begin position="2803"/>
            <end position="2841"/>
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    <feature type="coiled-coil region" evidence="5">
        <location>
            <begin position="2888"/>
            <end position="2963"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="5">
        <location>
            <begin position="2990"/>
            <end position="3073"/>
        </location>
    </feature>
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        <location>
            <begin position="3137"/>
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        </location>
    </feature>
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        <location>
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        <location>
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        <location>
            <begin position="4323"/>
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    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="1"/>
            <end position="13"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="72"/>
            <end position="87"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
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    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="SMART" id="SM01188"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001212.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="4377" mass="511643" checksum="B8E0CB07D736011E" modified="2019-05-08" version="1">[encoded data, 4377 characters]</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GI15</accession>
    <name>A0A452GI15_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Dual specificity protein phosphatase</fullName>
            <ecNumber evidence="6">3.1.3.16</ecNumber>
            <ecNumber evidence="6">3.1.3.48</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="12 13">
            <text>O-phospho-L-threonyl-[protein] + H2O = L-threonyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:47004"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="EC" id="3.1.3.16"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>O-phospho-L-tyrosyl-[protein] + H2O = L-tyrosyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:10684"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="EC" id="3.1.3.48"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 6">Belongs to the protein-tyrosine phosphatase family. Non-receptor class dual specificity subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.16" evidence="6"/>
    <dbReference type="EC" id="3.1.3.48" evidence="6"/>
    <dbReference type="AlphaFoldDB" id="A0A452GI15"/>
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        <property type="protein sequence ID" value="ENSGAGP00000001272.1"/>
        <property type="gene ID" value="ENSGAGG00000000928.1"/>
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    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
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        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017017">
        <property type="term" value="F:MAP kinase tyrosine/serine/threonine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004722">
        <property type="term" value="F:protein serine/threonine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
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    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
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    <dbReference type="GO" id="GO:0001706">
        <property type="term" value="P:endoderm formation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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        <property type="term" value="P:negative regulation of MAPK cascade"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd14640">
        <property type="entry name" value="DSP_DUSP4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd01446">
        <property type="entry name" value="DSP_MapKP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000015">
        <property type="entry name" value="Dual specificity phosphatase 4"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Rhodanese-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000340">
        <property type="entry name" value="Dual-sp_phosphatase_cat-dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008343">
        <property type="entry name" value="MKP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036873">
        <property type="entry name" value="Rhodanese-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016130">
        <property type="entry name" value="Tyr_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003595">
        <property type="entry name" value="Tyr_Pase_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020422">
        <property type="entry name" value="TYR_PHOSPHATASE_DUAL_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10159">
        <property type="entry name" value="DUAL SPECIFICITY PROTEIN PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10159:SF111">
        <property type="entry name" value="DUAL SPECIFICITY PROTEIN PHOSPHATASE 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00782">
        <property type="entry name" value="DSPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000939">
        <property type="entry name" value="MAPK_Ptase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01764">
        <property type="entry name" value="MAPKPHPHTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00195">
        <property type="entry name" value="DSPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00404">
        <property type="entry name" value="PTPc_motif"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52821">
        <property type="entry name" value="Rhodanese/Cell cycle control phosphatase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00383">
        <property type="entry name" value="TYR_PHOSPHATASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50054">
        <property type="entry name" value="TYR_PHOSPHATASE_DUAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 6" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="5" id="KW-0539">Nucleus</keyword>
    <keyword evidence="4 6" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="8">
        <location>
            <begin position="127"/>
            <end position="268"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="9">
        <location>
            <begin position="195"/>
            <end position="246"/>
        </location>
    </feature>
    <feature type="active site" description="Phosphocysteine intermediate" evidence="7">
        <location>
            <position position="212"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008601"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PIRNR" id="PIRNR000939"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PIRSR" id="PIRSR000939-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001272.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="281" mass="31482" checksum="E0F98B35D74A2A37" modified="2019-05-08" version="1">GALSVRCNSIVRRRAKGAVTLEQILPAEAEVRARLRAGLYSAVVLYDERSARAESLREDSTVWLVVRAVTLHACLSLIAGGYERFYSEYPEFCAKTKSLSSISPPTSMEPVELGCSSCGTPLHDQGGPVEILPFLYLGSAYHAARRDMLDTLGITALLNVSSDCPNHFEGHYQYKCIPVEDNHKADISSWFMEAIEYIDLVKDCHGRVLVHCQAGISRSATICLAYLMMKKRVKLEEAFEFVKQRRSIISPNFSFMGQLLQFESQVLCHCPICRVRRPFPL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GI16</accession>
    <name>A0A452GI16_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">3-ketoacyl-[acyl-carrier-protein] reductase beta subunit</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="12">Quinone reductase CBR4</fullName>
        </alternativeName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="pathway">
        <text evidence="2">Lipid metabolism; fatty acid biosynthesis.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Mitochondrion matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the short-chain dehydrogenases/reductases (SDR) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GI16"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001429.1">
        <property type="protein sequence ID" value="ENSGAGP00000001273.1"/>
        <property type="gene ID" value="ENSGAGG00000001019.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005759">
        <property type="term" value="C:mitochondrial matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016616">
        <property type="term" value="F:oxidoreductase activity, acting on the CH-OH group of donors, NAD or NADP as acceptor"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048038">
        <property type="term" value="F:quinone binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006633">
        <property type="term" value="P:fatty acid biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.720:FF:000285">
        <property type="entry name" value="Carbonyl reductase family member 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.720">
        <property type="entry name" value="NAD(P)-binding Rossmann-like Domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057326">
        <property type="entry name" value="KR_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036291">
        <property type="entry name" value="NAD(P)-bd_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020904">
        <property type="entry name" value="Sc_DH/Rdtase_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002347">
        <property type="entry name" value="SDR_fam"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR42760:SF133">
        <property type="entry name" value="3-OXOACYL-[ACYL-CARRIER-PROTEIN] REDUCTASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR42760">
        <property type="entry name" value="SHORT-CHAIN DEHYDROGENASES/REDUCTASES FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13561">
        <property type="entry name" value="adh_short_C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00081">
        <property type="entry name" value="GDHRDH"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00080">
        <property type="entry name" value="SDRFAMILY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00822">
        <property type="entry name" value="PKS_KR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51735">
        <property type="entry name" value="NAD(P)-binding Rossmann-fold domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00061">
        <property type="entry name" value="ADH_SHORT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="11" id="KW-0275">Fatty acid biosynthesis</keyword>
    <keyword evidence="5" id="KW-0276">Fatty acid metabolism</keyword>
    <keyword evidence="4" id="KW-0444">Lipid biosynthesis</keyword>
    <keyword evidence="9" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="10" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="8" id="KW-0520">NAD</keyword>
    <keyword evidence="6" id="KW-0521">NADP</keyword>
    <keyword evidence="7" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Ketoreductase" evidence="14">
        <location>
            <begin position="3"/>
            <end position="183"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004305"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005194"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00006484"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022516"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022832"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022857"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023027"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023160"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00041580"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00041707"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="SMART" id="SM00822"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001273.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="237" mass="25335" checksum="4D1103E3AEFE2CF3" modified="2019-05-08" version="1">MAKVCAVFGGSRGIGKAVAQLLAWKGYRLAIIARNLEVAQTTVNRLGAGHMALSCDVAMEQDVQNTLQEVERNLGQVNYLVNAAGVNRDGLLLRTTSEDMVSQINTNLLGTMLTCKAAVRHMLQQQGGAIVNIGSIVGLRGNSGQSVYSASKAGLVGFSRSLAKEVARKKIRVNMVAPGFVHTEMTAHLKEDQLKKTIPLGRFGEPDEVAQAVVFLLESPYITGHILVVDGGLQLVI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GI19</accession>
    <name>A0A452GI19_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">TFIIS central domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000001366.1">
        <property type="protein sequence ID" value="ENSGAGP00000001217.1"/>
        <property type="gene ID" value="ENSGAGG00000000965.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006351">
        <property type="term" value="P:DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.472.30">
        <property type="entry name" value="Transcription elongation factor S-II, central domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012921">
        <property type="entry name" value="SPOC_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003618">
        <property type="entry name" value="TFIIS_cen_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036575">
        <property type="entry name" value="TFIIS_cen_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11477:SF18">
        <property type="entry name" value="SPOC DOMAIN-CONTAINING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11477">
        <property type="entry name" value="TRANSCRIPTION FACTOR S-II ZINC FINGER DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07744">
        <property type="entry name" value="SPOC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07500">
        <property type="entry name" value="TFIIS_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00510">
        <property type="entry name" value="TFS2M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46942">
        <property type="entry name" value="Elongation factor TFIIS domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51321">
        <property type="entry name" value="TFIIS_CENTRAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="TFIIS central" evidence="2">
        <location>
            <begin position="963"/>
            <end position="1079"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="17"/>
            <end position="42"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="386"/>
            <end position="420"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="651"/>
            <end position="676"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="926"/>
            <end position="949"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1155"/>
            <end position="1180"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1619"/>
            <end position="1646"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="17"/>
            <end position="29"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="400"/>
            <end position="415"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="1619"/>
            <end position="1644"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="PROSITE" id="PS51321"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001217.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1658" mass="184365" checksum="419C1395378D9E8F" modified="2019-05-08" version="1">MEHQKFHGEQSKFCLNVNSKNMETSNTEAGSLGSGGPDRDINPEEEIFKKRAVVAGEAFEIPKINCVDNDDNDKGSRTVQSETWFGVGDDLDENCIKELSSTLKIPALEALGENEKSHSMKSYLKVQNQEESVTSQGARLTGVDAVKAEFPLNNEGTKLKELLSQMDFSGNLSNSWDQGTHSKSPIIPDLKEEKELFSFHLPACSEQVTGKRVGETYASDHVSNEEQSEQSTVTLNVVNQNKVSSESEVRVSGDGSSYSRSPYVKEIKAKETDFREKMIVSGGPSESPKQNCDNDERGSRTILIKPCNIGLKDLKDELKEKCKDSLISLDIETQSLKMKNESKASFSDVFQVPLHKNNVKKQNKIKIQNIKAPLENEKLNNTEKKEIKSHGEAVTSRGSDSMDTKIHHNSEETQGKRSFTQMDSPDLSDMIGDGIFSRLSLVEKEYKRSTINLSNVEEEGSRTKRLQSDSRPLSSMIPEVEKEKSREVITPVAKFRKFWNNPVTQLGKEEISYSYLVMNPNRKGFNSRLSERLFTSNISQSGRIILAPENTLQTKDLQGIPVEGTSTYLSASKTENVFGEGKSHNRSPVLPDQSDGIEDIHVTVCEPSCISPPAVEKFGTNQKGELSEMKNQMPRVEGLVQYFLEESAEEVSQGQLDVEEQKSLNQGNFKSNDESDDSSWFLMEIAEDLDNVDVANVPDGQNKAVEQETRGEHLNPSKVTLQSESMEVPVSGRHKAECYSCSNDINKIKHNDIKKSTAFCCLESDRDGQSSCVGKVAAPDSAKETCLIGEEESPLVHQSHGVQIVYYKMSGPVVELLGVPSQACKPRRLQRLVELEDSSDDEMCVCCQPPKKMSRLVLGVSKSKACKPCKMRRVEEMEDLSEDVYVDFQVTQEIKDSDYHPTLYKREMKQLLKPYAAHVRWQKPLNSDYSNEKKRRGRTPASKEASASVHGLQLSEEQSRIKVIDSLRGMLQKRLEESPDLDVHEDTILRLANNVEKEIFNLFLCVDQRYKNKYRSLLFNLKSPKNKLLFHQVVLGEVSPQSLVQMNSLELAPKELAEWRAKESKHVLEVIEKQQREPPRCCPTKLTHKGEIEIHREVDEDFTLEDLYGSALCMDKKSISQPAALSKRDTTEQHRSHLLDLDCLICTGRMAPDGESGFNPLQSKPTSKKKAEDSSRRLHSTGVYSYKERKHLEKDRPASVDVLKKSPRPKKQVRDTVLWEGFIQMFSIKQFAAKAYPVSGYGTCLIQEMSMIRFCPAMTKDFNTYNMLYSYLNNKQRYGIVNSNQMEMFMVPLPAFQPVPAKFHPLGGPGLDASHSCLLLGLILPKKVSRDPSTSSMGPSLLRETKRKTVTFKNNLVTKCFSPPLDSSGSTKLAEPFQQVPSRAFTGKRLSRNEYPPAVFHSQEPCNNMFALNTAVGEVIHGGTCHAASSSHCQNRKEETNQSMQDCAVHSLSSFSWMGGQSYSTAQESLGRQQNMIGGHTEFQADAGPAMTQNWNHGPLSSCSLCSHECGDNVALDVNMLDIQQLSSMLYPGVASSGGVSQVPSVAHLVSNEPPTASDLQNHAATGDSNPTIPVGEALSLIQHLEALVQLNSQIQNQTLFPLPSLDLSAILQATTPGESQIVGGSTSPPAQQESGVDPQSQLTLPPFCGSIYPPSGC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GI33</accession>
    <name>A0A452GI33_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">5-hydroxytryptamine receptor 7</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="14">Serotonin receptor 7</fullName>
        </alternativeName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">G-protein coupled receptor for 5-hydroxytryptamine (serotonin), a biogenic hormone that functions as a neurotransmitter, a hormone and a mitogen. Ligand binding causes a conformation change that triggers signaling via guanine nucleotide-binding proteins (G proteins) and modulates the activity of downstream effectors. HTR7 is coupled to G(s) G alpha proteins and mediates activation of adenylate cyclase activity.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="16">Belongs to the G-protein coupled receptor 1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GI33"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001391.1">
        <property type="protein sequence ID" value="ENSGAGP00000001239.1"/>
        <property type="gene ID" value="ENSGAGG00000001000.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045202">
        <property type="term" value="C:synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="GOC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004993">
        <property type="term" value="F:G protein-coupled serotonin receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071880">
        <property type="term" value="P:adenylate cyclase-activating adrenergic receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007268">
        <property type="term" value="P:chemical synaptic transmission"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007623">
        <property type="term" value="P:circadian rhythm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043410">
        <property type="term" value="P:positive regulation of MAPK cascade"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006939">
        <property type="term" value="P:smooth muscle contraction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042310">
        <property type="term" value="P:vasoconstriction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd15329">
        <property type="entry name" value="7tmA_5-HT7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1070.10:FF:000071">
        <property type="entry name" value="5-hydroxytryptamine (serotonin) receptor 7a"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1070.10">
        <property type="entry name" value="Rhodopsin 7-helix transmembrane proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001069">
        <property type="entry name" value="5HT_7_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000276">
        <property type="entry name" value="GPCR_Rhodpsn"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017452">
        <property type="entry name" value="GPCR_Rhodpsn_7TM"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24248">
        <property type="entry name" value="ADRENERGIC RECEPTOR-RELATED G-PROTEIN COUPLED RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24248:SF199">
        <property type="entry name" value="IP13425P-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00001">
        <property type="entry name" value="7tm_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00652">
        <property type="entry name" value="5HT7RECEPTR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00237">
        <property type="entry name" value="GPCRRHODOPSN"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81321">
        <property type="entry name" value="Family A G protein-coupled receptor-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00237">
        <property type="entry name" value="G_PROTEIN_RECEP_F1_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50262">
        <property type="entry name" value="G_PROTEIN_RECEP_F1_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="9" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6 16" id="KW-0297">G-protein coupled receptor</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="7 17" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-0564">Palmitate</keyword>
    <keyword evidence="10 16" id="KW-0675">Receptor</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="12 16" id="KW-0807">Transducer</keyword>
    <keyword evidence="4 16 17" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 17" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="77"/>
            <end position="101"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="113"/>
            <end position="132"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="144"/>
            <end position="173"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="194"/>
            <end position="214"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="234"/>
            <end position="255"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="325"/>
            <end position="345"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="365"/>
            <end position="383"/>
        </location>
    </feature>
    <feature type="domain" description="G-protein coupled receptors family 1 profile" evidence="18">
        <location>
            <begin position="93"/>
            <end position="380"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00015306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023139"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023224"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00031930"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00045776"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="RuleBase" id="RU000688"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50262"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001239.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="438" mass="48913" checksum="9B6FE5237F6D7DEB" modified="2019-05-08" version="1">MVMDINSSHVSGHLRAFALAAPRELSSSGMVAGSLPPRWLKIIQDVTASPSPASSLAAENDTRCGEQILSYGNMEKVVIGAILSLITLLTIAGNCLVVISVCFVKKLRQPSNYLIVSLALADLSVAVAVMPFVSVTDLIGGEWIFGHVFCNVFIAMDVMCCTASIMTLCVISIDRYLGITRPLTYPVRQNGKCMAKMILCVWLLSASITIPPLFGWAQNVNDEKVCLISQDFGYTIYSTAVAFYIPMSVMLFMYYQIYKAAKRSAAKHKFAGFPRPEENEGMVSVNGVIKLHKESEECTNFSRLLKHDRKNISIFKREQKAATTLGIIVGAFTVCWLPFFILSTARPFICGTACSCIPFWVERTFLWLGYANSLINPFIYAFFNRDLRTTYRNLLQCRYRNINRKLSAAGMHEALKLAEKPEFVLRNSDYSRKKSHDL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GI34</accession>
    <name>A0A452GI34_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Exocyst complex component</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="4">Component of the exocyst complex involved in the docking of exocytic vesicles with fusion sites on the plasma membrane.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1 4">Belongs to the SEC15 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GI34"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001453.1">
        <property type="protein sequence ID" value="ENSGAGP00000001295.1"/>
        <property type="gene ID" value="ENSGAGG00000001025.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000145">
        <property type="term" value="C:exocyst"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006893">
        <property type="term" value="P:Golgi to plasma membrane transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006886">
        <property type="term" value="P:intracellular protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090522">
        <property type="term" value="P:vesicle tethering involved in exocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.357.30:FF:000001">
        <property type="entry name" value="Exocyst complex component"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.670:FF:000001">
        <property type="entry name" value="Exocyst complex component"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.670">
        <property type="entry name" value="Dsl1p vesicle tethering complex, Tip20p subunit, domain D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.357.30">
        <property type="entry name" value="Exocyst complex subunit Sec15 C-terminal domain, N-terminal subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007225">
        <property type="entry name" value="EXOC6/Sec15"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046361">
        <property type="entry name" value="EXOC6/Sec15_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042045">
        <property type="entry name" value="EXOC6/Sec15_C_dom1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048359">
        <property type="entry name" value="EXOC6_Sec15_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042044">
        <property type="entry name" value="EXOC6PINT-1/Sec15/Tip20_C_dom2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12702:SF2">
        <property type="entry name" value="EXOCYST COMPLEX COMPONENT 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12702">
        <property type="entry name" value="SEC15"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20651">
        <property type="entry name" value="EXOC6_Sec15_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04091">
        <property type="entry name" value="Sec15_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF025007">
        <property type="entry name" value="Sec15"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 4" id="KW-0268">Exocytosis</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 4" id="KW-0813">Transport</keyword>
    <feature type="domain" description="Exocyst complex component EXOC6/Sec15 N-terminal" evidence="6">
        <location>
            <begin position="57"/>
            <end position="226"/>
        </location>
    </feature>
    <feature type="domain" description="Exocyst complex subunit EXOC6/Sec15 C-terminal" evidence="5">
        <location>
            <begin position="420"/>
            <end position="768"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007944"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PIRNR" id="PIRNR025007"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="Pfam" id="PF04091"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="Pfam" id="PF20651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001295.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="811" mass="94373" checksum="77BDE48710D4BCB0" modified="2019-05-08" version="1">MARKSKLNLLELYSEENKDQIYENVFAEIESFELPLGTSLRSVYDDQPNAHQRFMEKLDACIRNHDREIEKMCNFHHQGFVDAITELLKVRADAGKLKVQVTDTNRRLQDAGKEVIAQTEEIIRCRIQQRNITTVVEKLQLCLPVLEMYSKLKEQMNVKRYYSALKTMEQLENTYLPRVSQYRFCQIMIENLPKLREEIKEISMSDLKDFLESIRKHSDKIGETAMKQAQQQKTFNTVLQKQNNVNYGKNMYVINGRVPEPKYEITSKRTLEEEDEHEEEVLTAHDLVDFSPVYRCLHIYSVLGDGETFENYYRKQRRKQARLVLQPQSSMHETVEGYRRYFSQIVGFFVVEDHILHVTQGLVTRAYTDELWNMALSKIIAVLRTHSSYCTDPDLVLELKNLIVVFADTLQGYGFPVNRLFDLLFEIRDQYNETLLKKWTGLFRDIFEADNYSPIPVANEEEYKIVISKFPFQDPELDKQSFPKKLPMSQSVPQIYIQVKEFIYASLKFSESLHRSSTEIDDMLRKSTNLLLTRTLSSCLQNLIKKPHIGLTELVQIIINTTHLEQACKYLEDFITNITNISQETVHTARLYGLSTFKDARHAAEGEIYTKLNQKIDEFIQLADYDWTMSEPAGRASGYLMDLINFLRSTFQVFTHLPGKVAQTACMSACQHLSTSLMQMLLDSELKQISMGAVQQFNLDVIQCELFASSEPVPGFHGDTLQLAFIDLRQLLDLFMVWDWSTYLADYGQPTSKYLRVNPSTALTLLEKMKETSKKNNIFAQFRKNDRDKQKLIETVVKQLRSLVNGMSQHT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GI42</accession>
    <name>A0A452GI42_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">Gamma-aminobutyric acid receptor subunit gamma-3</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="22">GABA(A) receptor subunit gamma-3</fullName>
        </alternativeName>
    </protein>
    <organism evidence="27 28">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="28" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="27" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="27" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="18">Gamma subunit of the heteropentameric ligand-gated chloride channel gated by gamma-aminobutyric acid (GABA), a major inhibitory neurotransmitter in the brain. GABA-gated chloride channels, also named GABA(A) receptors (GABAAR), consist of five subunits arranged around a central pore and contain GABA active binding site(s) located at the alpha and beta subunit interface(s). When activated by GABA, GABAARs selectively allow the flow of chloride across the cell membrane down their electrochemical gradient.</text>
    </comment>
    <comment type="subunit">
        <text evidence="20">Heteropentamer, formed by a combination of alpha (GABRA1-6), beta (GABRB1-3), gamma (GABRG1-3), delta (GABRD), epsilon (GABRE), rho (GABRR1-3), pi (GABRP) and theta (GABRQ) chains, each subunit exhibiting distinct physiological and pharmacological properties.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="17">Postsynaptic cell membrane</location>
            <topology evidence="17">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="19">Belongs to the ligand-gated ion channel (TC 1.A.9) family. Gamma-aminobutyric acid receptor (TC 1.A.9.5) subfamily. GABRG3 sub-subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GI42"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001392.1">
        <property type="protein sequence ID" value="ENSGAGP00000001240.1"/>
        <property type="gene ID" value="ENSGAGG00000000999.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034707">
        <property type="term" value="C:chloride channel complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005254">
        <property type="term" value="F:chloride channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005230">
        <property type="term" value="F:extracellular ligand-gated monoatomic ion channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004890">
        <property type="term" value="F:GABA-A receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007214">
        <property type="term" value="P:gamma-aminobutyric acid signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd19054">
        <property type="entry name" value="LGIC_TM_GABAAR_gamma"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.390:FF:000006">
        <property type="entry name" value="Putative gamma-aminobutyric acid receptor subunit gamma-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.70.170.10:FF:000003">
        <property type="entry name" value="Putative gamma-aminobutyric acid receptor subunit gamma-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.70.170.10">
        <property type="entry name" value="Neurotransmitter-gated ion-channel ligand-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.390">
        <property type="entry name" value="Neurotransmitter-gated ion-channel transmembrane domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006028">
        <property type="entry name" value="GABAA/Glycine_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005437">
        <property type="entry name" value="GABRG-1/4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006202">
        <property type="entry name" value="Neur_chan_lig-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036734">
        <property type="entry name" value="Neur_chan_lig-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006201">
        <property type="entry name" value="Neur_channel"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036719">
        <property type="entry name" value="Neuro-gated_channel_TM_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038050">
        <property type="entry name" value="Neuro_actylchol_rec"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006029">
        <property type="entry name" value="Neurotrans-gated_channel_TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018000">
        <property type="entry name" value="Neurotransmitter_ion_chnl_CS"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00860">
        <property type="entry name" value="LIC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18945">
        <property type="entry name" value="NEUROTRANSMITTER GATED ION CHANNEL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02931">
        <property type="entry name" value="Neur_chan_LBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02932">
        <property type="entry name" value="Neur_chan_memb"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00253">
        <property type="entry name" value="GABAARECEPTR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01620">
        <property type="entry name" value="GABAARGAMMA"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00252">
        <property type="entry name" value="NRIONCHANNEL"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90112">
        <property type="entry name" value="Neurotransmitter-gated ion-channel transmembrane pore"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF63712">
        <property type="entry name" value="Nicotinic receptor ligand binding domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00236">
        <property type="entry name" value="NEUROTR_ION_CHANNEL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="13" id="KW-0868">Chloride</keyword>
    <keyword evidence="11" id="KW-0869">Chloride channel</keyword>
    <keyword evidence="10 23" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="16 24" id="KW-0407">Ion channel</keyword>
    <keyword evidence="7 24" id="KW-0406">Ion transport</keyword>
    <keyword evidence="15" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="8 24" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-0564">Palmitate</keyword>
    <keyword evidence="14" id="KW-0628">Postsynaptic cell membrane</keyword>
    <keyword evidence="28" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="6" id="KW-0770">Synapse</keyword>
    <keyword evidence="3 24" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 24" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="1 24" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="24">
        <location>
            <begin position="236"/>
            <end position="260"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="24">
        <location>
            <begin position="266"/>
            <end position="286"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="24">
        <location>
            <begin position="293"/>
            <end position="315"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="24">
        <location>
            <begin position="414"/>
            <end position="433"/>
        </location>
    </feature>
    <feature type="domain" description="Neurotransmitter-gated ion-channel ligand-binding" evidence="25">
        <location>
            <begin position="31"/>
            <end position="236"/>
        </location>
    </feature>
    <feature type="domain" description="Neurotransmitter-gated ion-channel transmembrane" evidence="26">
        <location>
            <begin position="243"/>
            <end position="354"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="23">
        <location>
            <begin position="154"/>
            <end position="168"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023139"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023173"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023214"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023257"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00034104"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00058949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00061119"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00064898"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00073032"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00078610"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="PIRSR" id="PIRSR605437-50"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="RuleBase" id="RU000687"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="Pfam" id="PF02931"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="26">
        <source>
            <dbReference type="Pfam" id="PF02932"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001240.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="28">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="434" mass="50944" checksum="A1284956BDF8AE29" modified="2019-05-08" version="1">MTESMEDYDYDYLSVNKTWVLTPKTQETDATQILNSLLKNYDNKLRPDIGIKPTFIDVDIYVNSIGPVSVIQMEYTIDIFFAQTWYDRRLRFNSTLKALTLNTNMVSRIWIPDTFFRNSKRADSHWITTPNQLLRIWNDGKVLYTLRLTIEAECLLQLQNFPMDTHSCPLVFSSYGYPREEIIYRWRRYSIEVSDQRTWRLYQFDFTGLRNTSEVLRTGAGDYMVMTVSFDLSRRMGYFAIQTYIPCILTVVLSWVSFWIKRDSTPARTSLGITTVLTMTTLSTISRKHLPRVSYITAMDLFVSVCFIFVFAALMEYATLNYLVGNKKPPEHNSKKPRLPPANAHMMPSFTTININNLMHWPPEVEEEEELGAPCLEGKECERFFCCIEDCQTGTWREGRIRIHISRLDSYSRVFFPTAFLLFNIVYWIAYLYL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GI45</accession>
    <name>A0A452GI45_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Protein fantom</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="16">Nephrocystin-8</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="17">RPGR-interacting protein 1-like protein</fullName>
        </alternativeName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Negatively regulates signaling through the G-protein coupled thromboxane A2 receptor (TBXA2R). May be involved in mechanisms like programmed cell death, craniofacial development, patterning of the limbs, and formation of the left-right axis. Involved in the organization of apical junctions; the function is proposed to implicate a NPHP1-4-8 module. Does not seem to be strictly required for ciliogenesis. Involved in establishment of planar cell polarity such as in cochlear sensory epithelium and is proposed to implicate stabilization of disheveled proteins. Involved in regulation of proteasomal activity at the primary cilium probably implicating association with PSDM2.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="4">Cell junction</location>
            <location evidence="4">Tight junction</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Cytoskeleton</location>
            <location evidence="3">Cilium axoneme</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Cilium basal body</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
            <location evidence="2">Microtubule organizing center</location>
            <location evidence="2">Centrosome</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the RPGRIP1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GI45"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001467.1">
        <property type="protein sequence ID" value="ENSGAGP00000001307.1"/>
        <property type="gene ID" value="ENSGAGG00000001041.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005930">
        <property type="term" value="C:axoneme"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005923">
        <property type="term" value="C:bicellular tight junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036064">
        <property type="term" value="C:ciliary basal body"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032391">
        <property type="term" value="C:photoreceptor connecting cilium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031870">
        <property type="term" value="F:thromboxane A2 receptor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1905515">
        <property type="term" value="P:non-motile cilium assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009966">
        <property type="term" value="P:regulation of signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046548">
        <property type="term" value="P:retinal rod cell development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00030">
        <property type="entry name" value="C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000196">
        <property type="entry name" value="Protein fantom"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000073">
        <property type="entry name" value="protein fantom isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000075">
        <property type="entry name" value="protein fantom isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021656">
        <property type="entry name" value="C2-C2_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000008">
        <property type="entry name" value="C2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041091">
        <property type="entry name" value="RPGRIP1_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031139">
        <property type="entry name" value="RPGRIP1_fam"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14240:SF4">
        <property type="entry name" value="PROTEIN FANTOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14240">
        <property type="entry name" value="RETINITIS PIGMENTOSA GTPASE REGULATOR-INTERACTING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00168">
        <property type="entry name" value="C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11618">
        <property type="entry name" value="C2-C2_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18111">
        <property type="entry name" value="RPGR1_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00239">
        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49562">
        <property type="entry name" value="C2 domain (Calcium/lipid-binding domain, CaLB)"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50004">
        <property type="entry name" value="C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0965">Cell junction</keyword>
    <keyword evidence="13" id="KW-0966">Cell projection</keyword>
    <keyword evidence="11" id="KW-0969">Cilium</keyword>
    <keyword evidence="10 18" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="12" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0796">Tight junction</keyword>
    <feature type="domain" description="C2" evidence="20">
        <location>
            <begin position="768"/>
            <end position="894"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1023"/>
            <end position="1076"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="18">
        <location>
            <begin position="105"/>
            <end position="139"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="18">
        <location>
            <begin position="200"/>
            <end position="255"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="18">
        <location>
            <begin position="326"/>
            <end position="457"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="18">
        <location>
            <begin position="494"/>
            <end position="549"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="1023"/>
            <end position="1043"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="1044"/>
            <end position="1060"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="19">
        <location>
            <begin position="1061"/>
            <end position="1073"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004120"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004430"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004435"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00006042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022427"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023069"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00058876"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00071893"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00078503"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00082462"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001307.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1273" mass="146130" checksum="4D7FD81DA16B0C48" modified="2019-05-08" version="1">MSAPADETAADLPVRDVGLNLAGIGGLQESSAAQSVKARQAVSRISREELEDRFLRLHDENILLKQHAHKQEDKIKRMATKLIRLVNDKKRSEQIGGGPKRLGQDVELEEMIEQLQGKVRELEKQNEGIRKRLISTKQQLQVQGHRHTPYSHVQSRINTGLRKVNEAVAIQDHTRKGMRFQDVELRCPQPMLPRYGHSLLEEARAEIRNLENVIESQKGQIEELEHTGEVLRSQIRKKEKEFEESVLQLKEQQATGQRLSIRDNVEMIKLHKQLMEKSNALSAMEGKFLQLQESQRNLKASHDALMANGNELNLQLKEQRLKCLHLEKELQSMTFSEQRIEELQDRINDLEKERDLLKENYDKLYNSAFSVSHEQQWKLKEQQLKLQIAQLETALKSDLADKSEILDRMKLERDQNEKLTQENKELQLHYLEHKQQLDELKNRMKFFTKESDTEMAELSEALMLIKVRKQQKNGDLTFLEKIEDDIHKDTEYAMRDLQATHAETVQELEKTRNMLIVQHKINKDYQTEVEAVTQKMEGLQKDYELKLEQYVHLLDIRAARIRKLEAQLKDIAYGTKQYKFRPEILPEDPADEFDETVHLERGENLFEIHISKVIFSTEAVHAFGDQEPVTFCTYAFYDFELQTTPVVHGLNPSYNFTSQYLVRVDDFFLQYIQKSTITLEVHHALGTDYETAAACQLRFQEILENNGRIYSTAILVGTGGHIPNYGTVDYWVRLRVPMDQAIRLYKERSKALGYITSNLKEHEKAPQLKQPIHRTAQVSTSADGDLNELHITIKCCSNLKSRKKHLQPNPYVVYKFFDFADHDTPIIHSSNSPQFDDHMCFPVPMNTDLDQYLKSESLTFYVFDDGETEEGLYVGKAIVPLISLAHDRYISGTFELTDSEGHSSGTINVDLKWRFAYLPPSGSAVAADLVSYIQNEKPMATKLPVKEEVPTLTLTPSFTSSMTAPKPKPRQRTALADKKVSFMDVSILQIPSPIADEDNEEISQKIKADTRTVSNVPEVIQERVADDKVKETSGEIQQEKDDLSQLSEGQLADQSSATSGDETEITEELEPEDQDNRLATDAMESIITDSDDCIVPSPVSKNIKQSIEKIRIEIISLSLTDSRVAMDDTIQRLFVECRFYSFLAEETPVSLPKPKSGQWVHYNYSNVIHVDKVNNCARREFLKSVLQESELNTGSIRFTVVSDPPEDEQDLECEDIGFAYVSLTEIFQKEKDIIEQDIDVFDSQDDSAVIGKLKVTVEALHALRSVYEEHKDD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GI47</accession>
    <name>A0A452GI47_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">5-hydroxytryptamine receptor 7</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="14">Serotonin receptor 7</fullName>
        </alternativeName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">G-protein coupled receptor for 5-hydroxytryptamine (serotonin), a biogenic hormone that functions as a neurotransmitter, a hormone and a mitogen. Ligand binding causes a conformation change that triggers signaling via guanine nucleotide-binding proteins (G proteins) and modulates the activity of downstream effectors. HTR7 is coupled to G(s) G alpha proteins and mediates activation of adenylate cyclase activity.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="16">Belongs to the G-protein coupled receptor 1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GI47"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001399.1">
        <property type="protein sequence ID" value="ENSGAGP00000001246.1"/>
        <property type="gene ID" value="ENSGAGG00000001000.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045202">
        <property type="term" value="C:synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="GOC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004993">
        <property type="term" value="F:G protein-coupled serotonin receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071880">
        <property type="term" value="P:adenylate cyclase-activating adrenergic receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007268">
        <property type="term" value="P:chemical synaptic transmission"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007623">
        <property type="term" value="P:circadian rhythm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043410">
        <property type="term" value="P:positive regulation of MAPK cascade"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006939">
        <property type="term" value="P:smooth muscle contraction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042310">
        <property type="term" value="P:vasoconstriction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd15329">
        <property type="entry name" value="7tmA_5-HT7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1070.10:FF:000071">
        <property type="entry name" value="5-hydroxytryptamine (serotonin) receptor 7a"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1070.10">
        <property type="entry name" value="Rhodopsin 7-helix transmembrane proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001069">
        <property type="entry name" value="5HT_7_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000276">
        <property type="entry name" value="GPCR_Rhodpsn"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017452">
        <property type="entry name" value="GPCR_Rhodpsn_7TM"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24248">
        <property type="entry name" value="ADRENERGIC RECEPTOR-RELATED G-PROTEIN COUPLED RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24248:SF199">
        <property type="entry name" value="IP13425P-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00001">
        <property type="entry name" value="7tm_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00652">
        <property type="entry name" value="5HT7RECEPTR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00237">
        <property type="entry name" value="GPCRRHODOPSN"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81321">
        <property type="entry name" value="Family A G protein-coupled receptor-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00237">
        <property type="entry name" value="G_PROTEIN_RECEP_F1_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50262">
        <property type="entry name" value="G_PROTEIN_RECEP_F1_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="9" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6 16" id="KW-0297">G-protein coupled receptor</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="7 17" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-0564">Palmitate</keyword>
    <keyword evidence="10 16" id="KW-0675">Receptor</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="12 16" id="KW-0807">Transducer</keyword>
    <keyword evidence="4 16 17" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 17" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="77"/>
            <end position="101"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="113"/>
            <end position="132"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="144"/>
            <end position="173"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="194"/>
            <end position="214"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="234"/>
            <end position="255"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="325"/>
            <end position="345"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="365"/>
            <end position="383"/>
        </location>
    </feature>
    <feature type="domain" description="G-protein coupled receptors family 1 profile" evidence="18">
        <location>
            <begin position="93"/>
            <end position="380"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00015306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023139"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023224"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00031930"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00045776"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="RuleBase" id="RU000688"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50262"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001246.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="425" mass="47325" checksum="0537A01BE038A6FF" modified="2019-05-08" version="1">MVMDINSSHVSGHLRAFALAAPRELSSSGMVAGSLPPRWLKIIQDVTASPSPASSLAAENDTRCGEQILSYGNMEKVVIGAILSLITLLTIAGNCLVVISVCFVKKLRQPSNYLIVSLALADLSVAVAVMPFVSVTDLIGGEWIFGHVFCNVFIAMDVMCCTASIMTLCVISIDRYLGITRPLTYPVRQNGKCMAKMILCVWLLSASITIPPLFGWAQNVNDEKVCLISQDFGYTIYSTAVAFYIPMSVMLFMYYQIYKAAKRSAAKHKFAGFPRPEENEGMVSVNGVIKLHKESEECTNFSRLLKHDRKNISIFKREQKAATTLGIIVGAFTVCWLPFFILSTARPFICGTACSCIPFWVERTFLWLGYANSLINPFIYAFFNRDLRTTYRNLLQCRYRNINRKLSAAGMHEALKLAEKPEFVL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GI53</accession>
    <name>A0A452GI53_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Peptidase S1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GI53"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001486.1">
        <property type="protein sequence ID" value="ENSGAGP00000001326.1"/>
        <property type="gene ID" value="ENSGAGG00000000827.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00190">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.10:FF:000005">
        <property type="entry name" value="Serine protease 37"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.10.10">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009003">
        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043504">
        <property type="entry name" value="Peptidase_S1_PA_chymotrypsin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001314">
        <property type="entry name" value="Peptidase_S1A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001254">
        <property type="entry name" value="Trypsin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018114">
        <property type="entry name" value="TRYPSIN_HIS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271:SF81">
        <property type="entry name" value="GRANZYME B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271">
        <property type="entry name" value="KALLIKREIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00089">
        <property type="entry name" value="Trypsin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00722">
        <property type="entry name" value="CHYMOTRYPSIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00020">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50494">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50240">
        <property type="entry name" value="TRYPSIN_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00134">
        <property type="entry name" value="TRYPSIN_HIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0732">Signal</keyword>
    <keyword evidence="2" id="KW-0865">Zymogen</keyword>
    <feature type="domain" description="Peptidase S1" evidence="4">
        <location>
            <begin position="14"/>
            <end position="226"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023145"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001326.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="244" mass="26358" checksum="24F8132ABA1F5808" modified="2019-05-08" version="1">MQSLSCLVSSEGDIIGGREARPHSRPYMAFVETKMGPESTTECGGFLVREDFVLTAAHCDEEGSGDAPTVVILGAHDLNRKERTQQEIQVRRKIPHEDFDRTTLENDIMLLQVVSLTTVSHVLKPIRLPPANFLLKPEATCSVAGWGRTGVHMVRPATKLQEVTLKMADDKQCELFFDSDSSSSGGAPYRVSHPGDTGGPLVCNGFAQSIVSYGKDSGLPSAVYTRGPFQPGHNLLLPSPSPGP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYR0</accession>
    <name>A0A452GYR0_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="3">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GYR0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008399.1">
        <property type="protein sequence ID" value="ENSGAGP00000007262.1"/>
        <property type="gene ID" value="ENSGAGG00000005834.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021684">
        <property type="entry name" value="WBP1-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051994">
        <property type="entry name" value="WW_domain-binding"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16209">
        <property type="entry name" value="VESICULAR, OVEREXPRESSED IN CANCER, PROSURVIVAL PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16209:SF4">
        <property type="entry name" value="WW DOMAIN BINDING PROTEIN 1-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11669">
        <property type="entry name" value="WBP-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="2" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="43"/>
            <end position="61"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="136"/>
            <end position="213"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="136"/>
            <end position="150"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="176"/>
            <end position="198"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007262.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="347" mass="39119" checksum="2EBE16B828318EA4" modified="2019-05-08" version="1">MPFLLGLRQDKQTCLGVNNQSYICETGHCCGQSQCCNYYYELWWFWLVWTIIIILSCCCVCHHRRAKHRLQAQQRQHEINLIAYREAHNYSALPFYFRFLPNYLLPPYEEVVNRPPTPPPPYSALHQQCVPAGNSSTILDTPRNLQPAQTSSPPPPGGNNNSTDSTGSLSVRDPESSMTLVEQTVTKMQATEPGNSSPRAELEETADPVSFPDKDFECKEELLKDYSCESLDHNSAIPDARDKTPGRHRRFTGDSGIEVCVCNRGHHDDDLKELDGLIDDTLGGPLDFCDSCSGRLHGDEEEGLFSASEEQHREHNHHHLPRQPVCVLLNTINEQDSPNSHTNHSPS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYR2</accession>
    <name>A0A452GYR2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Protein sel-1 homolog 1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="14">Suppressor of lin-12-like protein 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="12">Plays a role in the endoplasmic reticulum quality control (ERQC) system also called ER-associated degradation (ERAD) involved in ubiquitin-dependent degradation of misfolded endoplasmic reticulum proteins. Enhances SYVN1 stability. Plays a role in LPL maturation and secretion. Required for normal differentiation of the pancreas epithelium, and for normal exocrine function and survival of pancreatic cells. May play a role in Notch signaling.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="11">Belongs to the sel-1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYR2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008440.1">
        <property type="protein sequence ID" value="ENSGAGP00000007299.1"/>
        <property type="gene ID" value="ENSGAGG00000004194.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031012">
        <property type="term" value="C:extracellular matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036503">
        <property type="term" value="P:ERAD pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007219">
        <property type="term" value="P:Notch signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd00062">
        <property type="entry name" value="FN2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.10.10:FF:000001">
        <property type="entry name" value="Fibronectin 1a isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000208">
        <property type="entry name" value="Protein sel-1 homolog 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000193">
        <property type="entry name" value="protein sel-1 homolog 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000200">
        <property type="entry name" value="protein sel-1 homolog 1 precursor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.10.10">
        <property type="entry name" value="Fibronectin, type II, collagen-binding"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.10">
        <property type="entry name" value="Tetratricopeptide repeat domain"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000562">
        <property type="entry name" value="FN_type2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036943">
        <property type="entry name" value="FN_type2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013806">
        <property type="entry name" value="Kringle-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006597">
        <property type="entry name" value="Sel1-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050767">
        <property type="entry name" value="Sel1_AlgK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011990">
        <property type="entry name" value="TPR-like_helical_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11102">
        <property type="entry name" value="SEL-1-LIKE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11102:SF147">
        <property type="entry name" value="SEL1L ADAPTOR SUBUNIT OF ERAD E3 UBIQUITIN LIGASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00040">
        <property type="entry name" value="fn2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08238">
        <property type="entry name" value="Sel1"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00013">
        <property type="entry name" value="FNTYPEII"/>
    </dbReference>
    <dbReference type="SMART" id="SM00059">
        <property type="entry name" value="FN2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00671">
        <property type="entry name" value="SEL1"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81901">
        <property type="entry name" value="HCP-like"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57440">
        <property type="entry name" value="Kringle-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00023">
        <property type="entry name" value="FN2_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51092">
        <property type="entry name" value="FN2_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9 15" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="8 17" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-0914">Notch signaling pathway</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0732">Signal</keyword>
    <keyword evidence="2 17" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 17" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="682"/>
            <end position="700"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-II" evidence="18">
        <location>
            <begin position="78"/>
            <end position="126"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1"/>
            <end position="41"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1"/>
            <end position="10"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="16">
        <location>
            <begin position="19"/>
            <end position="41"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="83"/>
            <end position="109"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="97"/>
            <end position="124"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004115"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022976"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00038101"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00054921"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00068339"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00076627"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS51092"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007299.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="731" mass="82138" checksum="2F05E2E69966E511" modified="2019-05-08" version="1">EQVGDRSTGTRVVAGQIFHESEESDSHTEDDESFRSPEEEKESMLEELNSLEMSNLLNKNLEDVEIQKPVLTAIGGTADSEPCHFPFLFLEKEYAECTADGREDGRLWCATTYDYKRDQKWGFCETEEQSNKRRQMQEAEDVYQTGMKIFNESSKKAQKKEAYQYLLKAADMNHTKAMEKVSYALLFGDYLKQNIQSAKELFEKLTEEGSPKGQTVSVRVAVSYYAILSSVLLLTAGYRYWGGIGVLQSCESALTHYRLVANHVASDISLTGGTVVQRIRLADEVENPGMASGMLEEDLIQYYQFLAEKGDVQAQVGLGQLHLHGGRGVEQNHQRAFEYFNQAANAGNSHAMAFLGKLYSEGSDIIPQSNETALQYFKKAADMGNPVGQSGLGMAYLYGRGVPVNYELALKYFQKAAEQGWVDGQLQLGSMYYNGIGVKRDYKQALKYFNLASQGGHILAFYNLAQMHATGTGVMRSCHTAVELFKNVCERGRWSERLMTAYNSYKDGDSNAAVVQYLLLAEQGYEVAQSNAAFILDQKEASIVGENETYPQALLHWNRAASQGYTVARIKLGDYHFYGFGTDVDYETAFIHYRLASEQQHSAQAMFNLGYMHEKGLGIKQDIHLAKRFYDMAAEASPDAQVPVFLALCKLGIVYSLQYVRETNIREIVSHLDMDQLLGPEWDLYLMTIIALLLGTVIAYRQRQHQAILRPVGLRPAVPQQEPPPEHQPPQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYR7</accession>
    <name>A0A452GYR7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Zinc finger protein 711</fullName>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">Transcription regulator required for brain development. Probably acts as a transcription factor that binds to the promoter of target genes and recruits PHF8 histone demethylase, leading to activated expression of genes involved in neuron development, such as KDM5C. May compete with transcription factor ARX for activation of expression of KDM5C.</text>
    </comment>
    <comment type="subunit">
        <text evidence="14">Interacts with PHF8.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYR7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008445.1">
        <property type="protein sequence ID" value="ENSGAGP00000007304.1"/>
        <property type="gene ID" value="ENSGAGG00000005866.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001228">
        <property type="term" value="F:DNA-binding transcription activator activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903706">
        <property type="term" value="P:regulation of hemopoiesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000054">
        <property type="entry name" value="Zinc finger protein 711"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000179">
        <property type="entry name" value="Zinc finger protein 711"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000209">
        <property type="entry name" value="Zinc finger protein 711"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000170">
        <property type="entry name" value="Zinc finger protein 711 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006794">
        <property type="entry name" value="Transcrp_activ_Zfx/Zfy-dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24393">
        <property type="entry name" value="ZINC FINGER PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24393:SF140">
        <property type="entry name" value="ZINC FINGER Y-CHROMOSOMAL PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04704">
        <property type="entry name" value="Zfx_Zfy_act"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="12"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="10"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="12" id="KW-0539">Nucleus</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="11" id="KW-0804">Transcription</keyword>
    <keyword evidence="9" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="8" id="KW-0832">Ubl conjugation</keyword>
    <keyword evidence="7" id="KW-0862">Zinc</keyword>
    <keyword evidence="6 16" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="17">
        <location>
            <begin position="381"/>
            <end position="408"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="17">
        <location>
            <begin position="474"/>
            <end position="502"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="17">
        <location>
            <begin position="503"/>
            <end position="530"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="17">
        <location>
            <begin position="531"/>
            <end position="559"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="17">
        <location>
            <begin position="560"/>
            <end position="587"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="17">
        <location>
            <begin position="588"/>
            <end position="616"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="17">
        <location>
            <begin position="617"/>
            <end position="644"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="17">
        <location>
            <begin position="674"/>
            <end position="701"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="17">
        <location>
            <begin position="702"/>
            <end position="730"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="17">
        <location>
            <begin position="731"/>
            <end position="758"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00059295"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00061745"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00069176"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007304.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="759" mass="85328" checksum="7BC109691E3CD72D" modified="2019-05-08" version="1">MDPGGGSLGLQTQESKMPHTMIMQDFVAGMAGTAHIDGDHIVVSVPEAVLVSDVVTDDGITLDHGLAAEVVQGPDIITETDVVTEGVIVPESVLEADVAIEDALDTSDHVLTSDLITETVRVPDQVFVADLVTGPDGHLEHVVQDSVSGADSPTMVSEEVLVTNSDSETVIQAASTVPGSTVTIKTEDDDDGKSTSEDYLMISLDDVGEKLDHIGSTPLKISTEVSHDDVSKDDGFGSEVIKVYIFKAEAEDDVEIGGTEIVTESDFHNGHSVAGVIEQGSVGRMQREKMVYMAVKDSSQEDEDISCAEIADEVYMEVIVGEEEATSLPDTQLEDSGVNKTFVPVAWAAAYGDERRLPRRYEDCQAAAVIIGPDGQPLTVYPCHICGKKFKSRGFLKRHMKNHPDHMIKKKYQCTDCDFTTNKKISFHNHLESHKLINKVDKTHEFTEYTRRYREASPLSSNKLILRDKEPKLHKCKYCDYETAEQGLLNRHLLAVHSKNFPHVCVECGKGFRHPSELKKHMRTHTGEKPYQCQHCVFRCADQSNLKTHIKTKHGTDLPFKCEHCPQAFTDEKELLQHTELFQGHKTHQCPHCDHKSTNSSDLKRHIISVHTKDFPHKCEVCEKGFHRPSELKKHSETHKGKKIHQCRHCDFKTSDPFVLSGHILSVHTKDLPFKCKRCKRGFRQQNELKKHMKTHSGRKVYQCQYCEYSTTDASGFKRHVISIHTKDYPHRCEYCKKGFRRPSEKNQHIMRHHKEAIM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYR9</accession>
    <name>A0A452GYR9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">POF1B helix-loop-helix domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GYR9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008543.1">
        <property type="protein sequence ID" value="ENSGAGP00000007394.1"/>
        <property type="gene ID" value="ENSGAGG00000005876.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005884">
        <property type="term" value="C:actin filament"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005912">
        <property type="term" value="C:adherens junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005923">
        <property type="term" value="C:bicellular tight junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007015">
        <property type="term" value="P:actin filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070830">
        <property type="term" value="P:bicellular tight junction assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003382">
        <property type="term" value="P:epithelial cell morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026186">
        <property type="entry name" value="POF1B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056240">
        <property type="entry name" value="POF1B_HlH"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22546">
        <property type="entry name" value="PREMATURE OVARIAN FAILURE, 1B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22546:SF0">
        <property type="entry name" value="PROTEIN POF1B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24617">
        <property type="entry name" value="POF1B_HlH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="POF1B helix-loop-helix" evidence="2">
        <location>
            <begin position="41"/>
            <end position="116"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="137"/>
            <end position="164"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="200"/>
            <end position="308"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF24617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007394.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="361" mass="42482" checksum="4A757830A4CFE49C" modified="2019-05-08" version="1">GNETQVFRTVMKVNPCLFIYSFREKLPVHSLQETNGQLDSRYFGELLAEVNRKSNDLYSCLLQHVEKIGRRYYQQETYIEDLIPKGLSELTKQQIRYLLQMRVTSDKSLRLVLSTFGSLREELCHLQDDLGKLETDKMLLEKDLAFKESQVKEYETLLASVRENNRQQQQGLRDSASKCRSLEEQIISLRYAEGEKDCRLKELEYCKRALEQEIQNLRMQTCSNPVIQTTTDELSSRYVEMINNLREDKDREIRNLRSQINQFQQDISRRDGSNSDFQMRLHELTSMLEEKETFIKQQQEELFRLKQERLSCNQSTGVTKSIITKKYMNQYPILGLLSDDCMASSPVRQMETIVIEKTEAM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYT1</accession>
    <name>A0A452GYT1_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="7">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1 5">Belongs to the GDA1/CD39 NTPase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYT1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008460.1">
        <property type="protein sequence ID" value="ENSGAGP00000007319.1"/>
        <property type="gene ID" value="ENSGAGG00000005862.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004382">
        <property type="term" value="F:GDP phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017111">
        <property type="term" value="F:ribonucleoside triphosphate phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045134">
        <property type="term" value="F:UDP phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009134">
        <property type="term" value="P:nucleoside diphosphate catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.420.40:FF:000068">
        <property type="entry name" value="Ectonucleoside triphosphate diphosphohydrolase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.420.40">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.420.150">
        <property type="entry name" value="Exopolyphosphatase. Domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000407">
        <property type="entry name" value="GDA1_CD39_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11782">
        <property type="entry name" value="ADENOSINE/GUANOSINE DIPHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11782:SF33">
        <property type="entry name" value="ECTONUCLEOSIDE TRIPHOSPHATE DIPHOSPHOHYDROLASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01150">
        <property type="entry name" value="GDA1_CD39"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01238">
        <property type="entry name" value="GDA1_CD39_NTPASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="2 5" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="4" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="6">
        <location>
            <begin position="1"/>
            <end position="38"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019280519" evidence="6">
        <location>
            <begin position="39"/>
            <end position="399"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="3">
        <location>
            <position position="169"/>
        </location>
    </feature>
    <feature type="binding site" evidence="4">
        <location>
            <begin position="208"/>
            <end position="212"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00009283"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PIRSR" id="PIRSR600407-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PIRSR" id="PIRSR600407-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="RuleBase" id="RU003833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007319.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="399" mass="43902" checksum="5091379DDAAC7374" modified="2019-05-08" version="1">MAAHQAHWEHLVGLGGSRIHLLLLLRAGLCASLSFSLSRPVPQYGIVLDAGSSHTAMFVYKWPADKENDTGIVSQHSMCDVEGGGISSYAENPPAAGKSLKQCLDQAVSDVPKERHPLTPLYLGATAGMRLLNLTDPQASDRVLSAVTSTLKSFPFDFRGAKILSGEDEGVFGWVTANYLLENFIKRVWHMLWFRPKKPTLGAMDLGGASTQITFETQKPIEDPRNEVTLRLYGQTYKVYTHSFLCYGRDQILRRMLSKADGYKTNVSNPCWPKGYQRNFKLQDVYDSPCTAAEKPDSYDPQRSIDMSGSGDTAQCRLHVDSLFSFTSCSFTSCSFDGVFQPGLSGNFIVSAFSAFFYTVDFIQTVMKRRVASPDDLNTAAEAICSASWTEVSALAAEG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYT4</accession>
    <name>A0A452GYT4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">N-acetyltransferase domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the acetyltransferase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYT4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008429.1">
        <property type="protein sequence ID" value="ENSGAGP00000007288.1"/>
        <property type="gene ID" value="ENSGAGG00000005856.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00188">
        <property type="reaction ID" value="UER00363"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004145">
        <property type="term" value="F:diamine N-acetyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019809">
        <property type="term" value="F:spermidine binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009447">
        <property type="term" value="P:putrescine catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032918">
        <property type="term" value="P:spermidine acetylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd04301">
        <property type="entry name" value="NAT_SF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.630.30:FF:000064">
        <property type="entry name" value="GNAT family acetyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.630.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016181">
        <property type="entry name" value="Acyl_CoA_acyltransferase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051016">
        <property type="entry name" value="Diverse_Substrate_AcTransf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000182">
        <property type="entry name" value="GNAT_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10545">
        <property type="entry name" value="DIAMINE N-ACETYLTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10545:SF63">
        <property type="entry name" value="SPERMIDINE_SPERMINE N(1)-ACETYLTRANSFERASE-LIKE PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00583">
        <property type="entry name" value="Acetyltransf_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55729">
        <property type="entry name" value="Acyl-CoA N-acyltransferases (Nat)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51186">
        <property type="entry name" value="GNAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0012">Acyltransferase</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="N-acetyltransferase" evidence="4">
        <location>
            <begin position="10"/>
            <end position="150"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00008694"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023315"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS51186"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007288.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="155" mass="17845" checksum="A25E39BC912CDD7A" modified="2019-05-08" version="1">FKDFKVHPPASDPKLAINSQVLILDLLEDGFGEHPFYHCLIAEVPKEGQIGSSCIVGYAMYYFTYDPWIGKLLYLEDFYVMEPYRGLGIGSEILKRLSQIAIKTNCSCMHFLVVIWNKPSIEYYVRRGALDLSTEEGWHLFRIDKENLLRMASGE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYT6</accession>
    <name>A0A452GYT6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">DnaJ homolog subfamily B member 9</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="8">Endoplasmic reticulum DNA J domain-containing protein 4</fullName>
        </alternativeName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Co-chaperone for Hsp70 protein HSPA5/BiP that acts as a key repressor of the ERN1/IRE1-mediated unfolded protein response (UPR). J domain-containing co-chaperones stimulate the ATPase activity of Hsp70 proteins and are required for efficient substrate recognition by Hsp70 proteins. In the unstressed endoplasmic reticulum, interacts with the luminal region of ERN1/IRE1 and selectively recruits HSPA5/BiP: HSPA5/BiP disrupts the dimerization of the active ERN1/IRE1 luminal region, thereby inactivating ERN1/IRE1. Also involved in endoplasmic reticulum-associated degradation (ERAD) of misfolded proteins. Required for survival of B-cell progenitors and normal antibody production.</text>
    </comment>
    <comment type="subunit">
        <text evidence="10">Interacts with HSPA5/BiP; interaction is direct. Interacts with ERN1/IRE1 (via the luminal region). Interacts with DERL1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYT6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008467.1">
        <property type="protein sequence ID" value="ENSGAGP00000007325.1"/>
        <property type="gene ID" value="ENSGAGG00000005883.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005783">
        <property type="term" value="C:endoplasmic reticulum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051787">
        <property type="term" value="F:misfolded protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051087">
        <property type="term" value="F:protein-folding chaperone binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036503">
        <property type="term" value="P:ERAD pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006986">
        <property type="term" value="P:response to unfolded protein"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd06257">
        <property type="entry name" value="DnaJ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.110:FF:000054">
        <property type="entry name" value="dnaJ homolog subfamily B member 9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.110">
        <property type="entry name" value="DnaJ domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001623">
        <property type="entry name" value="DnaJ_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018253">
        <property type="entry name" value="DnaJ_domain_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051948">
        <property type="entry name" value="Hsp70_co-chaperone_J-domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036869">
        <property type="entry name" value="J_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR44360">
        <property type="entry name" value="DNAJ HOMOLOG SUBFAMILY B MEMBER 9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR44360:SF1">
        <property type="entry name" value="DNAJ HOMOLOG SUBFAMILY B MEMBER 9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00226">
        <property type="entry name" value="DnaJ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00625">
        <property type="entry name" value="JDOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00271">
        <property type="entry name" value="DnaJ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46565">
        <property type="entry name" value="Chaperone J-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00636">
        <property type="entry name" value="DNAJ_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50076">
        <property type="entry name" value="DNAJ_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0143">Chaperone</keyword>
    <keyword evidence="4" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 11" id="KW-0732">Signal</keyword>
    <keyword evidence="6" id="KW-0834">Unfolded protein response</keyword>
    <feature type="signal peptide" evidence="11">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019256464" description="DnaJ homolog subfamily B member 9" evidence="11">
        <location>
            <begin position="24"/>
            <end position="217"/>
        </location>
    </feature>
    <feature type="domain" description="J" evidence="12">
        <location>
            <begin position="26"/>
            <end position="90"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023186"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023230"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00040158"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00041533"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00045428"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00046365"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007325.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="217" mass="25060" checksum="4A31C63361D2FD35" modified="2019-05-08" version="1">MATTQSVFTFALCILMITELILARESYYDVLGVPKNASDRQIKKAFHKLAMKYHPDKNKSPGAEAKFREIAEAYETLSDENKRKEYDQFGHGGGQGNNGSPFQQSFNFNFDDLFKDFDFFGQNQNSRSKKHFENHFRGHREAHSRQRRSFQDFSFGGGLFDDVFEDMEKMFSFSGFDSAQQHTVRTDSRFHGSSKHCRTVTQRRGNMVTTYTDCSGQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYU5</accession>
    <name>A0A452GYU5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">G-protein coupled receptors family 1 profile domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 10">Cell membrane</location>
            <topology evidence="1 10">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="10">Belongs to the G-protein coupled receptor 1 family. Vasopressin/oxytocin receptor subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYU5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008481.1">
        <property type="protein sequence ID" value="ENSGAGP00000007337.1"/>
        <property type="gene ID" value="ENSGAGG00000005889.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042277">
        <property type="term" value="F:peptide binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005000">
        <property type="term" value="F:vasopressin receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032870">
        <property type="term" value="P:cellular response to hormone stimulus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045907">
        <property type="term" value="P:positive regulation of vasoconstriction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001992">
        <property type="term" value="P:regulation of systemic arterial blood pressure by vasopressin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd15388">
        <property type="entry name" value="7tmA_V2R"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1070.10:FF:000205">
        <property type="entry name" value="[Arg8]-vasotocin receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1070.10">
        <property type="entry name" value="Rhodopsin 7-helix transmembrane proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000276">
        <property type="entry name" value="GPCR_Rhodpsn"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017452">
        <property type="entry name" value="GPCR_Rhodpsn_7TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001817">
        <property type="entry name" value="Vasoprsn_rcpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24241:SF74">
        <property type="entry name" value="ARGININE VASOPRESSIN RECEPTOR 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24241">
        <property type="entry name" value="NEUROPEPTIDE RECEPTOR-RELATED G-PROTEIN COUPLED RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00001">
        <property type="entry name" value="7tm_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00237">
        <property type="entry name" value="GPCRRHODOPSN"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00896">
        <property type="entry name" value="VASOPRESSINR"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81321">
        <property type="entry name" value="Family A G protein-coupled receptor-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00237">
        <property type="entry name" value="G_PROTEIN_RECEP_F1_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50262">
        <property type="entry name" value="G_PROTEIN_RECEP_F1_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="5 10" id="KW-0297">G-protein coupled receptor</keyword>
    <keyword evidence="8 10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6 10" id="KW-0472">Membrane</keyword>
    <keyword evidence="7 10" id="KW-0675">Receptor</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9 10" id="KW-0807">Transducer</keyword>
    <keyword evidence="3 10" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 10" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="41"/>
            <end position="63"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="75"/>
            <end position="95"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="115"/>
            <end position="133"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="153"/>
            <end position="175"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="200"/>
            <end position="219"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="268"/>
            <end position="289"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="301"/>
            <end position="324"/>
        </location>
    </feature>
    <feature type="domain" description="G-protein coupled receptors family 1 profile" evidence="11">
        <location>
            <begin position="54"/>
            <end position="321"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023224"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="RuleBase" id="RU046427"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50262"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007337.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="361" mass="41282" checksum="C91E6FE436709A9E" modified="2019-05-08" version="1">FLLAKTTTTESLFPQLVLNLTNKSDPVGRQERDEQLAQLEIAVLGVIFMTASVGNFILILVLWRRRKKLSRMYVFMLHLSIADLVVAFFQVLPQLIWDITDVFIGPDALCRTIKYLQLLGMFASAYMIVVMTVDRYQAVRYPMVTFQKKKALWNAAICTSWSISLLFSLPQIFIFSKTEISPDIFECWGEFIQPWGLKAYVTWIFVAIFFIPTAILTICQVKICKIILMNIHVKKRNEFEAINQKQVLPFQASSINCISKAMIKTVKMTVVTVIAYVLCWAPFFIVQLWSVWYPSGITEGAAFTIIMLLGNLNSCANPWIYMYFCGHIPYCAKKRSKNISAQEESVITGSINLHKDTKSNL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ01</accession>
    <name>A0A452GZ01_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Lysosomal-trafficking regulator</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="8">Interacts with CPAP, LIP8 and ZNF521.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000008520.1">
        <property type="protein sequence ID" value="ENSGAGP00000007373.1"/>
        <property type="gene ID" value="ENSGAGG00000005878.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098542">
        <property type="term" value="P:defense response to other organism"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006955">
        <property type="term" value="P:immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002449">
        <property type="term" value="P:lymphocyte mediated immunity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043473">
        <property type="term" value="P:pigmentation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd06071">
        <property type="entry name" value="Beach"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd01201">
        <property type="entry name" value="PH_BEACH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:000217">
        <property type="entry name" value="Lysosomal trafficking regulator"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:000292">
        <property type="entry name" value="Lysosomal trafficking regulator"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.1540.10:FF:000001">
        <property type="entry name" value="neurobeachin isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1540.10">
        <property type="entry name" value="BEACH domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000409">
        <property type="entry name" value="BEACH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036372">
        <property type="entry name" value="BEACH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050865">
        <property type="entry name" value="BEACH_Domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023362">
        <property type="entry name" value="PH-BEACH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019775">
        <property type="entry name" value="WD40_repeat_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036322">
        <property type="entry name" value="WD40_repeat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001680">
        <property type="entry name" value="WD40_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13743">
        <property type="entry name" value="BEIGE/BEACH-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13743:SF86">
        <property type="entry name" value="LYSOSOMAL-TRAFFICKING REGULATOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02138">
        <property type="entry name" value="Beach"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14844">
        <property type="entry name" value="PH_BEACH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00400">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01026">
        <property type="entry name" value="Beach"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00320">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81837">
        <property type="entry name" value="BEACH domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50978">
        <property type="entry name" value="WD40 repeat-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50197">
        <property type="entry name" value="BEACH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51783">
        <property type="entry name" value="PH_BEACH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00678">
        <property type="entry name" value="WD_REPEATS_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50082">
        <property type="entry name" value="WD_REPEATS_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50294">
        <property type="entry name" value="WD_REPEATS_REGION"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="11" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="7" id="KW-0653">Protein transport</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0813">Transport</keyword>
    <keyword evidence="5 10" id="KW-0853">WD repeat</keyword>
    <feature type="domain" description="BEACH-type PH" evidence="14">
        <location>
            <begin position="2921"/>
            <end position="3027"/>
        </location>
    </feature>
    <feature type="domain" description="BEACH" evidence="13">
        <location>
            <begin position="3032"/>
            <end position="3334"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="10">
        <location>
            <begin position="3524"/>
            <end position="3558"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="156"/>
            <end position="176"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="913"/>
            <end position="932"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1218"/>
            <end position="1247"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="11">
        <location>
            <begin position="2094"/>
            <end position="2123"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="163"/>
            <end position="176"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="12">
        <location>
            <begin position="1218"/>
            <end position="1232"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="1233"/>
            <end position="1244"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022574"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00064710"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00067676"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50197"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS51783"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007373.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="3713" mass="420794" checksum="ACBB08C90B07AC62" modified="2019-05-08" version="1">MSSASNSLAREFLIDVNRLCNAVVQRAEAREEEEEETHMAALGQYLVQGRGFTLLTTLNSNMDQELTCREELLTLLLSLLPLVWKIPVQEEKAINFNLPFTVDVCLTKETNSNSLRSAHEKQNLEGIVHPSFQTPVKLNSSFRKIRRQRKITHRYSVRDARKSQLSTSDSEGNSDNKTTIMIKHRRSQILQPFVTHLPRDDYHAEVVRSFNPSPISLENSRQIIPTQESKSEVLNESAAAAVVSSMDNSPFDLCHVLLSLLEKVCKFDITLNHNSALAATIVPSLTEFLSGFGDCCNVSGDAESEVVSAGWTEEPVALVQRMLFRTVLHLLSIDITNAETMPDNLKKNLTDLLRAALKIRTCLEKQSDPFVPRHKKTLQEVQDNFVFSKYRHRVLLLPELLEGVLQILICCLQSAASNPFYFSQAVDLVHEFIQHQGFKLLEITVLQMEWLVSKDEEITPEASEHLKALISSIMKIISTVKKVKSEQLHQSMCTRKRHRRCEYSHFMHHHRDLSGLPVSAFKNQVSRNPFEEVVDGEVHYPERCCCIAVCAHQCFHLLQQVSLSSTCAQILSGVHNVGICCCMDPKSVITPLLQASKLPVLKNFQQHILSILNKFILDQLGGAEVSQKIIQAACNICTIDCDQLAELEDTLQSNSTDIGLASNFSCPSYRFQGILPSSGSEDMLLKWDALEAYQDFAFEEDKLRSVQIASHICHLIQKGNVVVQWKLYNYIYNPVLQRGVELAHRSQQLGITAVCNHVCSCHSLCLPQEVLQIYLQTLPALFKSRIIQNLFLSCNGVNQMTELNYLDSVRSHSLKVFETLIFCLGDQQKDQAMPELEGMDNEQKESILDLVTSFRSQQMISDVPQSLSKFYAGLRKAYPKKKKCVSQEVHVNLIHLFLCVAFLCVSKEEECDRDSANDSEDTSGYDSTASEPLSHKLPCLSLESLTLPSPEHIHRAADIWSMCRWIYLCSSVFQRQFHRLGGFEVCHRLIIMIIQKLAKNKEKEQEKKERPLSENNKCLNASPLNQPVFKDDSYYLTVNKQLTQSELNSLDENPDASGQPVSECISCDHSLKVEQISVAMVELEELKKSVKQENEWSLQSIRLLEALLTICLHSANSTQQKAEVELLHQSVCLEDVLLEMRDQLAQSRAIETCLAKPLLDSLLRVALGNYSADLDHSEEKTEKILHTEKASVPQPEDLSEEAEESQCCSFKLFAEEEGYEADSESNPDDSESKDEGTDTKREAECMVPSSYSSDLTENISQGELIYPEICMLELKLLSAGNPSLDVLAHVFQSFLKVIRQKKKNATFLMQQGAVKSLLGGFLNILIQKDSSFHACQIVLVELLVSLMSSRTCAEELTLLLRIFLEETPCTEILLKGILKIVETNIYMSPLQYLSFPLLKSPNLNSSSSQKSTSALNSKTVGLLKRAKFSQNKKEADGENLTHQLLSSWHVAPIHLPLVGQNCWPHMSEGFSVSLWFHVECVQETESPTEKGKKMKKRSKPLIVRESSFDGTEGSKSVRMEYLNLGDKLVEDGCIHIISLGSKALMIQVWADLNTGAFIFRMCMDPNDEMKAGLLAQAESPENIFLVGKWEHLAITYLQQPEGKKNIHGELLLWVSGQRKCEISLDYTLPRKTTLSDSNKTFCMIGHCTSSQEEPLQLAGKWDLGNLLLFNGAKIGPEEAFYLYACGPDFTAIMPCKYGKTMTDYSKYITKDILQCERIKEFFMKKKEADVGPLIVIDITNCEETQALALQVILSLTKYNQQRIQEMENCNGYSMIHQVLIKPKCIVGFHILKTLLEGCFSDEILNINESGQFMLDVDSTAVIQDIKLLEDLLLDWKIWSKAQQGVWETLLAALEILIRANHHQQMFNIKQLLKARVVHHFLLTCQVLQEQKEGHLTSLPREVCRSFVKIIEEVLGSPPDLELLTLIFNFLLAVHPPTNTYVCHNPTNFYFSLHIDGKIFQEKVQSIKYLRNSSIGGKSVDSPSLTTSPAGFTTSPLGGSIISKIIIPQMMISQALQRRCKSLPASPTSSPQVHPKRLPGSLGRSIDNLQEVWESDSVPQEKNIFVESSETLKRIQEELFVSSCESAKTVCDSELTSAEALVDISKEELELSENKELNTELKSNENGAAVESLLRRPDNLKGLPSFQKSQNNFAGLGLAFSAQSGSSSVACWPSLADRSALPEDWESLTFSSVSEQTFKRSERPDYRSTEDCLVLICCGLYDLLRGVLLILPDIMLEDVIDKLIQPDTLIVLVNHPSPLIQERVIKLLDAYFNRATREQKEQFLKNRGFSLLANQLYLHQGTQELVECFLEMLFGRSVGLDEEFDREDVKNAGLFQKWCVIPVLGLIENSLYDNVLLHNSFCLLLQIINSCSKMADMLLDNGLLYVLCNTLAALNGLETNIPLNDYKLLLCDIQQLLVAVTIHACSSLGSQYFRVIEDLIVLLGHLQNNKNEKTQNMAIALQFRVLQAAIELIKATANQDSQDLANSLRLPATHHHAMFQKRKSIADSRKFSRAQSDTLVMKMRLVASDELNMMMQRRMSQENPIRATENEFIQRLQRLTVLAVNRFIYQEPTEDCFDVLGTPETTIQCRLSTSQLEASEEEDQRELPSNTNSFLKEMFKMMIEGIKVSIATSRISTPRQQWRRILWSCKDTFRMQLGRLLVHILSPAQPLQERKQALEIVCETNHQEIIRDCLSVPLQHGPKLALYLYELMHDHKDELTREEQVSSGVFMNALKFCGYKCISPSAPPKPDLLNAMKEDRKKFEAEECVNKTVWEKTVINNQQNLSQRLDAKSKDISKIAADITQAVSLSQGMERKKVIQHIRGMYKTDLSASRHWQELVQQLTHDRTLWYDPVYYPTSWQLDPTEGPNRERRRLRRCYLTIPNKYLLRDRQKQEEVIKPPLSYLFEDKTHSSHSSTVKDKAASENIRVNRRCINVAPSRETAGELLLGKYGMYFVEDNASDTIESSSFHGETEPASFSWTYEEIKEVHKRWWQLRDNAVEIFLTNGRTLLLAFDNTKVRDDVYHNILMNNLPNLLEYGNITALTHLWYTGQITNFEYLTHLNKHAGRSFNDLMQYPVFPFILSDYTSETLDLNDPSVYRNLAKPIAVQSKEKEDRYVDTYKYLEEEYRKGAREDDPMPPVQPYHYGSHYSNSGTVLHFLVRMPPFTKMFLAYQDQSFDIPDRTFHSTHTTWRLSSYESMTDVKELIPEFFYLPEFLVNREGFDFGLRQNGERVNHVNLPPWARNDPRLFILIHRQALESDHVSQTICNWIDLVFGYKQKGKASVQAINVFHPATYFGMDVSAVEDPVQRRALETMIKTYGQTPRQLFHTAHVSRPGSRLTIEGELPAAMGLLVQFAFRETREHAKEIVYPSPLPWIKGLKWGEYVGSPSAPDPVVCFSQPHGERFGSLQALPTRAICGLSQKFCLLMVYSKEQGVRSMHSTDIQWSAILSWGYADNILRLKSKQSEPPINFIQSSQLHQVTSCAWVPDSCQLFTGSKSGVITAYMNRFSSNMHSEIEMESQVHLYGHTAEITSLFVCKPYSIMISVSKDGTCIIWDLNRLCYVQSLAGHKSPVTAVSASEMTGDIATVCDSVGGGSDLRLWTVNGDLVGHVHCREIICSVAFSNQPEGVSINVIAGGLENGVVRLWSTWDLKPVREITFPKSNKPIVSLTFSCDGHHLYTANSEGTVIAWCRKDQQRLKLPMFYSFLSSYAAG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ05</accession>
    <name>A0A452GZ05_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Gamma-tubulin complex component</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="6">Component of the gamma-tubulin ring complex (gTuRC) which mediates microtubule nucleation. The gTuRC regulates the minus-end nucleation of alpha-beta tubulin heterodimers that grow into microtubule protafilaments, a critical step in centrosome duplication and spindle formation.</text>
    </comment>
    <comment type="subunit">
        <text evidence="7">Component of the gamma-tubulin ring complex (gTuRC) consisting of TUBGCP2, TUBGCP3, TUBGCP4, TUBGCP5 and TUBGCP6 and gamma-tubulin TUBG1 or TUBG2. TUBGCP2, TUBGCP3, TUBGCP4, TUBGCP5 and TUBGCP6 assemble in a 5:5:2:1:1 stoichiometry; each is associated with a gamma-tubulin, thereby arranging 14 gamma-tubulins in a helical manner. Gamma-tubulin at the first position is blocked by TUBGCP3 at the last position, allowing 13 protafilaments to grow into a microtubule. The gTuRC (via TUBGCP3 and TUBGCP6) interacts with ACTB and MZT1; the interactions form a luminal bridge that stabilizes the initial structure during complex assembly. The gTuRC (via TUBGCP2) interacts with MZT2A/MZT2B and CDK5RAP2 (via CM1 motif); the interactions play a role in gTuRC activation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Microtubule organizing center</location>
            <location evidence="1">Centrosome</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 8">Belongs to the TUBGCP family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZ05"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008420.1">
        <property type="protein sequence ID" value="ENSGAGP00000007280.1"/>
        <property type="gene ID" value="ENSGAGG00000004413.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000930">
        <property type="term" value="C:gamma-tubulin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005874">
        <property type="term" value="C:microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000922">
        <property type="term" value="C:spindle pole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043015">
        <property type="term" value="F:gamma-tubulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051011">
        <property type="term" value="F:microtubule minus-end binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031122">
        <property type="term" value="P:cytoplasmic microtubule organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051321">
        <property type="term" value="P:meiotic cell cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007020">
        <property type="term" value="P:microtubule nucleation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000278">
        <property type="term" value="P:mitotic cell cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051225">
        <property type="term" value="P:spindle assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd22572">
        <property type="entry name" value="GCP5_NTD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.1900:FF:000005">
        <property type="entry name" value="Gamma-tubulin complex component"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.1900">
        <property type="entry name" value="Gamma-tubulin complex, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007259">
        <property type="entry name" value="GCP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR059169">
        <property type="entry name" value="GCP5_N_ext"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040457">
        <property type="entry name" value="GCP_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042241">
        <property type="entry name" value="GCP_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041470">
        <property type="entry name" value="GCP_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19302">
        <property type="entry name" value="GAMMA TUBULIN COMPLEX PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19302:SF33">
        <property type="entry name" value="GAMMA-TUBULIN COMPLEX COMPONENT 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04130">
        <property type="entry name" value="GCP_C_terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17681">
        <property type="entry name" value="GCP_N_terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 8" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="5 8" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="4 8" id="KW-0493">Microtubule</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Gamma tubulin complex component protein N-terminal" evidence="11">
        <location>
            <begin position="270"/>
            <end position="611"/>
        </location>
    </feature>
    <feature type="domain" description="Gamma tubulin complex component C-terminal" evidence="10">
        <location>
            <begin position="709"/>
            <end position="1010"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="155"/>
            <end position="200"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="527"/>
            <end position="546"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="9">
        <location>
            <begin position="157"/>
            <end position="171"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="187"/>
            <end position="196"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="533"/>
            <end position="545"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010337"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00093416"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00093551"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="RuleBase" id="RU363050"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="Pfam" id="PF04130"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF17681"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007280.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1017" mass="117572" checksum="48DEC58FAD470E18" modified="2019-05-08" version="1">LAHAPLEQAQDRDVRALIRRLTGLREAQGGEEPGGRFQTALIFAWSNFRFHRFLDVSSHKVERTIEGIYEKLIVHSDLSKAASWKRLIEKFLNSPLPSTEETKTDAHYGILSLLLCLSDSPSNTNYVEKPRDKEEEKFDWGKYLMEGEEIYFGPDIDTPDWSEESEGEEDAQQPLSREDSGIQVDRTPQEEQDQNKKTVSQVTWKVGEPDARSWLEQHVVPQYWTGRAPRFSHSFHLHSNLAAVWDQHLYSSDPLYMPEERTLVTETQIIRETLWLLSGVKKLFIFQLNEGKVTVRNDIIVTHLTHNCLRSVLEQIAAYGQVVFRLQKFIDEVMGHSSESVIPGTISTPKKTTEPPFRTYQAFMWALYKYFISFKEELTEIEKCVINKDKTVTLSIVVDKLSPRLAQLKVLHKVFSTGIAEVPQVPPDTRNVVRASHLLNTLYKAILEYDNVGEASEQTVSLLFALWVETVRPYLQTVDEWIVHGNLFDPAKEFIIQRNKNVPVNHRDFWYATYTLYSVSEKMENEEKMSDNASASSGSDQAPSSRQHTMVSFLKPVLKQIIMAGKSMQLLKNLQSKDDSPRKSLYTLFLESVQSRLRHGEESVPDIITEQQATKQSLIKMQSIAERHLELDDVHDPLLAINFARLYLEQSDFHEKFTGGDVCVDRSSESVTCQTFELTLRSCLYPHIDKQYLECCGNLMQTLKKDYRLVEYLQAMRNFFLLEAGDTMYDFYTSIFDKIREKETWQNVSFLNVQLQEAVGQRYPEDSSRLSISFESIDTAKKKLPVHTLDGLTLSYKVPWPVDIVISLECQKIYNQVFLLLLQIKWAKYSLDVLRFDELVNATEKPQLKEGPQLEQETVPQFGSQTELIKQQIHRMFLVRVKLMHFVNSLHNYIMTRILHSTGLEFQHQVEEAKDLDQLIKIHYRYLSTIHDRCLLREKVSFVKEAIMKVLNLVLIFADRWQAGLGAWKMESIEKMESDFKNCHMFLVTVLNKAVCRGSFPHLESLAVSLMAGMEQS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ09</accession>
    <name>A0A452GZ09_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Ig-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZ09"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008426.1">
        <property type="protein sequence ID" value="ENSGAGP00000007285.1"/>
        <property type="gene ID" value="ENSGAGG00000005772.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="CDD" id="cd05763">
        <property type="entry name" value="IgI_LRIG1-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000161">
        <property type="entry name" value="Leucine rich repeats and immunoglobulin like domains 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000040">
        <property type="entry name" value="Leucine rich repeats and immunoglobulin like domains 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000150">
        <property type="entry name" value="Leucine rich repeats and immunoglobulin like domains 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000224">
        <property type="entry name" value="Leucine rich repeats and immunoglobulin like domains 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000023">
        <property type="entry name" value="Leucine rich repeats and immunoglobulin like domains 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000483">
        <property type="entry name" value="Cys-rich_flank_reg_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013098">
        <property type="entry name" value="Ig_I-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001611">
        <property type="entry name" value="Leu-rich_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003591">
        <property type="entry name" value="Leu-rich_rpt_typical-subtyp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050467">
        <property type="entry name" value="LRFN"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45842:SF16">
        <property type="entry name" value="LEUCINE-RICH REPEATS AND IMMUNOGLOBULIN-LIKE DOMAINS 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45842">
        <property type="entry name" value="SYNAPTIC ADHESION-LIKE MOLECULE SALM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07679">
        <property type="entry name" value="I-set"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13927">
        <property type="entry name" value="Ig_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13855">
        <property type="entry name" value="LRR_8"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01463">
        <property type="entry name" value="LRRCT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00019">
        <property type="entry name" value="LEURICHRPT"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00365">
        <property type="entry name" value="LRR_SD22"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SMART" id="SM00369">
        <property type="entry name" value="LRR_TYP"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="SMART" id="SM00082">
        <property type="entry name" value="LRRCT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52058">
        <property type="entry name" value="L domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51450">
        <property type="entry name" value="LRR"/>
        <property type="match status" value="5"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="9" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="3" id="KW-0433">Leucine-rich repeat</keyword>
    <keyword evidence="8 12" id="KW-0472">Membrane</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <keyword evidence="4 12" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 12" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="740"/>
            <end position="766"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="13">
        <location>
            <begin position="434"/>
            <end position="533"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="13">
        <location>
            <begin position="538"/>
            <end position="627"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="13">
        <location>
            <begin position="632"/>
            <end position="718"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022614"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007285.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="888" mass="98754" checksum="C34536395E73FC12" modified="2019-05-08" version="1">MRASKNIPDISKVTSHLFLLSSIKTSSLSHLHSLREIKLNNNELETIPNLGPVSANITLLSLTSNKIGGILPEHLKPFQSLETLDLSNNNISELKTASFPFLQLKYLYINSNRITSMEPGVFDSLSSTLQVLKLNRNKISVIPQKMFKLSHLQHLELNRNKIKKIDGLTFQGLTALKSLKMQRNGVNRLMDGAFWGLTNMEVLQLDHNNLTEITKGWLYGLLMLQQLHLSQNAISRISPDAWEFCQKLSELDLTFNHLARLDDSSFVGLSLLVGLYIGNNKVNYIADCAFRGLSSLKTLDLKNNEISWTIEDMNGAFSGLDKLRILILQGNRIRSITKKAFSGLDALEHLDLSNNAIMSVQGNAFSQMKKLEELHLNTSSLLCDCQLKWLPQWISENNFQSFVNASCAHPPLLKGRSIFAVSPDEFVCDDFPKPQITVQPETQSAIKGSNLSFVCSAASSSDSPMTFAWKKDNELLHDAEMENYAHLRARGGEVMEYTTILRLRNVEFSNEGKYQCVISNHFGSSYSVKAKLTVNMLPSFTKMPMDLSIRAGATARLECAAVGHPVPQIAWQKDGGTDFPAARKRRMHVMPEDDVFFIVDVKIEDTGVYSCTAQNTAGSISANATLTVLETPSFLRPLLDRTVTKGETAVLQCIAGGSPPPRLNWTKDDSPLVVTERHFFAAGNQLLIIVDTDIEDAGKYTCEMSNTLGTERGNIRLNVIPTPTCDSPQNIAPSLDDDGWATVGIVIIAVVCCVVGTSLVWVVIIYHTRRRNEDCSITNTDETNLPADIPSYLSSQGTLAERQDGYGSSENGSHHQFITSSMGGYFLQQRDSNGKYFKYKTWHCFWLLEIRHCGRCGDLFLISNCRVTFVAPLSSNSAQSCLFNPKPD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ30</accession>
    <name>A0A452GZ30_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="19">Phospholipid-transporting ATPase</fullName>
            <ecNumber evidence="19">7.6.2.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="15 19">
            <text>ATP + H2O + phospholipidSide 1 = ADP + phosphate + phospholipidSide 2.</text>
            <dbReference type="EC" id="7.6.2.1"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 18">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Golgi apparatus</location>
            <location evidence="2">trans-Golgi network membrane</location>
            <topology evidence="2">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="19">Membrane</location>
            <topology evidence="19">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 19">Belongs to the cation transport ATPase (P-type) (TC 3.A.3) family. Type IV subfamily.</text>
    </comment>
    <dbReference type="EC" id="7.6.2.1" evidence="19"/>
    <dbReference type="AlphaFoldDB" id="A0A452GZ30"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008588.1">
        <property type="protein sequence ID" value="ENSGAGP00000007434.1"/>
        <property type="gene ID" value="ENSGAGG00000000721.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005768">
        <property type="term" value="C:endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005802">
        <property type="term" value="C:trans-Golgi network"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140326">
        <property type="term" value="F:ATPase-coupled intramembrane lipid transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000287">
        <property type="term" value="F:magnesium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006897">
        <property type="term" value="P:endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045332">
        <property type="term" value="P:phospholipid translocation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006890">
        <property type="term" value="P:retrograde vesicle-mediated transport, Golgi to endoplasmic reticulum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.1110.10:FF:000008">
        <property type="entry name" value="Phospholipid-transporting ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1000:FF:000009">
        <property type="entry name" value="Phospholipid-transporting ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.1110.10">
        <property type="entry name" value="Calcium-transporting ATPase, cytoplasmic domain N"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.70.150.10">
        <property type="entry name" value="Calcium-transporting ATPase, cytoplasmic transduction domain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1110.10">
        <property type="entry name" value="Calcium-transporting ATPase, transmembrane domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1000">
        <property type="entry name" value="HAD superfamily/HAD-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023299">
        <property type="entry name" value="ATPase_P-typ_cyto_dom_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018303">
        <property type="entry name" value="ATPase_P-typ_P_site"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023298">
        <property type="entry name" value="ATPase_P-typ_TM_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008250">
        <property type="entry name" value="ATPase_P-typ_transduc_dom_A_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036412">
        <property type="entry name" value="HAD-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023214">
        <property type="entry name" value="HAD_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006539">
        <property type="entry name" value="P-type_ATPase_IV"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001757">
        <property type="entry name" value="P_typ_ATPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032630">
        <property type="entry name" value="P_typ_ATPase_c"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044492">
        <property type="entry name" value="P_typ_ATPase_HD_dom"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01652">
        <property type="entry name" value="ATPase-Plipid"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01494">
        <property type="entry name" value="ATPase_P-type"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24092:SF50">
        <property type="entry name" value="PHOSPHOLIPID-TRANSPORTING ATPASE IIB-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24092">
        <property type="entry name" value="PROBABLE PHOSPHOLIPID-TRANSPORTING ATPASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00702">
        <property type="entry name" value="Hydrolase"/>
        <property type="match status" value="1"/>
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    <dbReference type="Pfam" id="PF16212">
        <property type="entry name" value="PhoLip_ATPase_C"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="CATATPASE"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDG00002">
        <property type="entry name" value="C1.7:_P-type_atpase_like"/>
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    <dbReference type="SFLD" id="SFLDS00003">
        <property type="entry name" value="Haloacid_Dehalogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDF00027">
        <property type="entry name" value="p-type_atpase"/>
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    </dbReference>
    <dbReference type="SUPFAM" id="SSF81653">
        <property type="entry name" value="Calcium ATPase, transduction domain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81665">
        <property type="entry name" value="Calcium ATPase, transmembrane domain M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56784">
        <property type="entry name" value="HAD-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81660">
        <property type="entry name" value="Metal cation-transporting ATPase, ATP-binding domain N"/>
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    </dbReference>
    <dbReference type="PROSITE" id="PS00154">
        <property type="entry name" value="ATPASE_E1_E2"/>
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    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 17 19" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="12" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="13" id="KW-0445">Lipid transport</keyword>
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    <keyword evidence="6 18" id="KW-0479">Metal-binding</keyword>
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    <keyword evidence="4" id="KW-0813">Transport</keyword>
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        <location>
            <begin position="75"/>
            <end position="94"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="123"/>
            <end position="143"/>
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            <begin position="699"/>
            <end position="716"/>
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    <feature type="transmembrane region" description="Helical" evidence="19">
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            <begin position="746"/>
            <end position="766"/>
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    <feature type="transmembrane region" description="Helical" evidence="19">
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            <begin position="772"/>
            <end position="793"/>
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            <begin position="800"/>
            <end position="819"/>
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    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="839"/>
            <end position="863"/>
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    <feature type="domain" description="P-type ATPase C-terminal" evidence="21">
        <location>
            <begin position="641"/>
            <end position="872"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="253"/>
            <end position="275"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="20">
        <location>
            <begin position="253"/>
            <end position="263"/>
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    </feature>
    <feature type="active site" description="4-aspartylphosphate intermediate" evidence="16">
        <location>
            <position position="208"/>
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        <location>
            <position position="208"/>
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        <ligand>
            <name>ATP</name>
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        <location>
            <position position="208"/>
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        <ligand>
            <name>Mg(2+)</name>
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            <name>ATP</name>
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        <location>
            <position position="397"/>
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            <name>ATP</name>
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            <position position="426"/>
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            <name>ATP</name>
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            <name>ATP</name>
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            <dbReference type="ARBA" id="ARBA00023055"/>
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    </evidence>
    <evidence type="ECO:0000313" key="23">
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    <sequence length="880" mass="98537" checksum="D76CA26795EC68F7" modified="2019-05-08" version="1">MSENLKLLLDGETDWKLKVAVSCTQRLPALGDLFSISAYVYAQKPQLDIHSFEGTFTREDSDPTVHESLSIENTLWASTVVASGTVIGVVIYTGKETRSVMNTSNPKNKVGLLDLELNQLTKALFLALVALSIVMVTLQGFVGPWYRNLFRFLLLFSYIIPISLRVNLDMGKAAYGWMIMKDDNIPGTVVRTSTIPEELGRLVYLLTDKTGTLTQNEMIFKRLHLGTVSYGTDTMDEIQNHIISSYSQMYSQSSGNSTSSTPSRKVQSSAPKVRKTVSSRIHEAVKAIALCHNVTPVYESHAGVNGETEYAEVDQNFSDENRTYQASSPDEVALVQWTESVGLTLVNRDLTSMQLKSPGGQILTYYILQVFPFTSESKRMGIIVRDESTGEITFYMKGADVAMSTIVQYNDWLEEECGNMAREGLRTLVVAKKSLTEEQHQDFENRYNQAKLSIHDRALKVAAVVESLEREMELLCLTGVEDQLQADVRPTLEMLRNAGIKIWMLTGDKLETATCIAKSSHLVSRTQDIHIFRPVISRGEAHLELNAFRRKHDCALVISGDSLEVCLKYYEHEFVELACQCPAVVCCRCSPTQKAHIVKLLQHHTGKRTCAIGDGGNDVSMIQAADCGIGIEGKEGKQASLAADFSITQFKHIGRLLMVHGRNSYKRSAALGQFVMHRGLIISTMQVFKLLCLYNTYKMSLYATIYTMFPVFSLVLDQDVKPEMALLYPELYKDLTKGRSLSFKTFLIWVLISIYQGGILMYGALLLFESEFVHVVAISFTALILTELLMVALTIRTWHWLMVVAEFFSLGCYVASLAFLNEYFGIGRVSFGAFLDVAFITTVTFLWKVSAITVVSCLPLYILKYLKRKFSPPSYSKLTS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ36</accession>
    <name>A0A452GZ36_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">Platelet endothelial aggregation receptor 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="11">Belongs to the MEGF family.</text>
    </comment>
    <comment type="caution">
        <text evidence="12">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZ36"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008523.1">
        <property type="protein sequence ID" value="ENSGAGP00000007376.1"/>
        <property type="gene ID" value="ENSGAGG00000005918.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="CDD" id="cd00055">
        <property type="entry name" value="EGF_Lam"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.170.300.10:FF:000002">
        <property type="entry name" value="Multiple epidermal growth factor-like domains 10"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="FunFam" id="2.170.300.10:FF:000007">
        <property type="entry name" value="multiple epidermal growth factor-like domains protein 10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000114">
        <property type="entry name" value="Multiple epidermal growth factor-like domains protein 11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.170.300.10">
        <property type="entry name" value="Tie2 ligand-binding domain superfamily"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013111">
        <property type="entry name" value="EGF_extracell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057138">
        <property type="entry name" value="EGF_PEAR1L-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011489">
        <property type="entry name" value="EMI_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002049">
        <property type="entry name" value="LE_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052485">
        <property type="entry name" value="MEGF_diff_regulators"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24052">
        <property type="entry name" value="DELTA-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24052:SF12">
        <property type="entry name" value="PLATELET ENDOTHELIAL AGGREGATION RECEPTOR 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07974">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00053">
        <property type="entry name" value="EGF_laminin"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23301">
        <property type="entry name" value="EGF_PEAR1L"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00011">
        <property type="entry name" value="EGFLAMININ"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="SMART" id="SM00180">
        <property type="entry name" value="EGF_Lam"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="12"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01186">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50027">
        <property type="entry name" value="EGF_LAM_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51041">
        <property type="entry name" value="EMI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="9 12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3 12" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13" id="KW-0424">Laminin EGF-like domain</keyword>
    <keyword evidence="8 15" id="KW-0472">Membrane</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <keyword evidence="4 15" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 15" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="801"/>
            <end position="824"/>
        </location>
    </feature>
    <feature type="domain" description="EMI" evidence="18">
        <location>
            <begin position="67"/>
            <end position="145"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="16">
        <location>
            <begin position="225"/>
            <end position="260"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="16">
        <location>
            <begin position="268"/>
            <end position="304"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="16">
        <location>
            <begin position="317"/>
            <end position="347"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="16">
        <location>
            <begin position="360"/>
            <end position="390"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="17">
        <location>
            <begin position="407"/>
            <end position="450"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="16">
        <location>
            <begin position="449"/>
            <end position="479"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="16">
        <location>
            <begin position="530"/>
            <end position="565"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="17">
        <location>
            <begin position="581"/>
            <end position="628"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="16">
        <location>
            <begin position="621"/>
            <end position="651"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="16">
        <location>
            <begin position="659"/>
            <end position="694"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1014"/>
            <end position="1073"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="250"/>
            <end position="259"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="294"/>
            <end position="303"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="337"/>
            <end position="346"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="380"/>
            <end position="389"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="13">
        <location>
            <begin position="426"/>
            <end position="435"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="469"/>
            <end position="478"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="555"/>
            <end position="564"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="13">
        <location>
            <begin position="598"/>
            <end position="607"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="641"/>
            <end position="650"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="684"/>
            <end position="693"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004162"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00038377"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00460"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50027"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS51041"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007376.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="21">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1073" mass="114623" checksum="FB5AD7FF53D54CF1" modified="2019-05-08" version="1">MHWDKDGPMSRGPQQAEVLGSIPSSAMDSLCKILGQSFRADVSRMKLHVVLLLGLQMHLATSLNPSDPNVCSYWESFTTATKESYAHPYAQASKDSCDGTWSFLKPCTQHKIVYKTAYRQAVKIDYRKRYRCCQGYYESADVCVPRCTKECVHGRCVAPDQCQCEQGWRGTDCSSVCNGQSWGPHCENPCQCGDGGACDPLTGACVCSPGYKDSMCKVPCDPGTYGEGCQLACPCKNTDRCHGETGACLCQPGFTGTYCELLCLNSSDGLHCPAYCPCQNGGICHPPNTTHCVCPPGWMGTICSIPCPQGQYGSGCLGECQCHNNGLCDPVTGRCQCALGYTGERCREECPIGKYGQDCGQPCDCANGGRCFHINGACLCEAGFYGPRCEERKCLDGLYGLACHLPCLCNPQNSQSCHPMSGECTCKPGWAGLHCNETCPHGYYGANCQEPCLCLNGGTCDGATGLCHCAPGYTGDHCSSHCPTDTYGVNCSMQCACKNAFACSPIDGTCVCKEGWHGGDCSRPCPTGTWGPGCNETCRCANGAACSPVSGACSCAAGWHGSRCQEPCTNGAYGPGCSSQCDCSHADGCSPVTGQCRCLPGWTGPRCNLVCDDGFWGRNCNQPCACKNGAACSSEDGSCSCAPGYRGPACQRPCQPGRYGKKCAMLCRCSNHSTCHPVDGSCDCFPGWTGSDCSQPCPPGTWGEACGHTCRCPNGGWCSTVDGTCSCPAGLTGRRCSEGCPPGSYGENCAQACQCQNMAQCDPATGRCLCSSGYTGAHCETRSPDHPFTMVPAVPVGYSSLGAIIGIIILVILLVALLVFFLCYRHWQKGKENRHLSVAYTTGRTDSSEYAVPDVPPSYTHYYSNPSYHTLSRCTPTSPAPSSQDRASSLKVPCNQHFSNTKSMAYGPDCNASLPADWKHHGAPALKPRERGGGPMDRSYSYSNSLGKYYSNAYIKEEALRASNSSLNSENPYATIKDLPELMAKPSEGSYMEMKSPVKREMSYAEIRLFEEPAAGSRAEETCPPGAEGVSPAAPSSPPNHYDSPKNSHIPSHYDMPPVRHYPPSPPLRRQDR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ37</accession>
    <name>A0A452GZ37_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Zinc finger protein RFP-like</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GZ37"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008451.1">
        <property type="protein sequence ID" value="ENSGAGP00000007310.1"/>
        <property type="gene ID" value="ENSGAGG00000005871.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd19762">
        <property type="entry name" value="Bbox2_TRIM7-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd16594">
        <property type="entry name" value="RING-HC_TRIM7-like_C-IV"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12888">
        <property type="entry name" value="SPRY_PRY_TRIM7_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.920:FF:000004">
        <property type="entry name" value="Butyrophilin subfamily 1 member A1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.920">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001870">
        <property type="entry name" value="B30.2/SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043136">
        <property type="entry name" value="B30.2/SPRY_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003879">
        <property type="entry name" value="Butyrophylin_SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006574">
        <property type="entry name" value="PRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003877">
        <property type="entry name" value="SPRY_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050143">
        <property type="entry name" value="TRIM/RBCC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000315">
        <property type="entry name" value="Znf_B-box"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001841">
        <property type="entry name" value="Znf_RING"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017907">
        <property type="entry name" value="Znf_RING_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24103">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE TRIM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13765">
        <property type="entry name" value="PRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00622">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00643">
        <property type="entry name" value="zf-B_box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15227">
        <property type="entry name" value="zf-C3HC4_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01407">
        <property type="entry name" value="BUTYPHLNCDUF"/>
    </dbReference>
    <dbReference type="SMART" id="SM00336">
        <property type="entry name" value="BBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00589">
        <property type="entry name" value="PRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00184">
        <property type="entry name" value="RING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00449">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57845">
        <property type="entry name" value="B-box zinc-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57850">
        <property type="entry name" value="RING/U-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50188">
        <property type="entry name" value="B302_SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50119">
        <property type="entry name" value="ZF_BBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00518">
        <property type="entry name" value="ZF_RING_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50089">
        <property type="entry name" value="ZF_RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="1" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0862">Zinc</keyword>
    <keyword evidence="2 4" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="RING-type" evidence="7">
        <location>
            <begin position="16"/>
            <end position="57"/>
        </location>
    </feature>
    <feature type="domain" description="B box-type" evidence="8">
        <location>
            <begin position="86"/>
            <end position="127"/>
        </location>
    </feature>
    <feature type="domain" description="B30.2/SPRY" evidence="9">
        <location>
            <begin position="216"/>
            <end position="420"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="220"/>
            <end position="241"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="382"/>
            <end position="420"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="5">
        <location>
            <begin position="155"/>
            <end position="189"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00024"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50119"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50188"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007310.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="12">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="420" mass="46794" checksum="A98C8D535AAEFA32" modified="2019-05-08" version="1">MAAANPAKTLQDEVTCPICLEYFKDPVSLDCDHSFCRACITQCWGGFHTDISCPQCREIFPQRNLRLNRQLRNIVEAARELRLQPEPESLCEKHREPLKLFCKEDEILICLVCDRSKEHRVHTVIPTEEAAEEFQVGNNQLFLKLKIVSEFQQLQEFLEEQKRLLLGQLAELNEEIVKIQNENITKLSKEISQTNDVATIKQVRNCSDISAQSQVLKTEAGPWDPGLSQPRQGSNVTLDPDTAHPSLILSEDRKSVRWGDTRQDLPDRAERQISQGCVLGCEGFTSGRHCWEVEVEVGDALFWAVGVVRESVGRKGWISCSSGGGIWAVQWCRGRFQVLTSLVISLPLSRIPSRIWVCLDCDQGQVTFIDAGDEAPIFIFPPGSVPGRESDPGSGWGGEPSSDCVPEIQGGGRGNIPLGS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ39</accession>
    <name>A0A452GZ39_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Zinc transporter ZIP13</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="9">Solute carrier family 39 member 13</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="10">Zrt- and Irt-like protein 13</fullName>
        </alternativeName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="7">
            <text>Zn(2+)(in) = Zn(2+)(out)</text>
            <dbReference type="Rhea" id="RHEA:29351"/>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the ZIP transporter (TC 2.A.5) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZ39"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008573.1">
        <property type="protein sequence ID" value="ENSGAGP00000007423.1"/>
        <property type="gene ID" value="ENSGAGG00000005958.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005385">
        <property type="term" value="F:zinc ion transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006882">
        <property type="term" value="P:intracellular zinc ion homeostasis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003689">
        <property type="entry name" value="ZIP"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16950">
        <property type="entry name" value="ZINC TRANSPORTER SLC39A7 HISTIDINE-RICH MEMBRANE PROTEIN KE4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16950:SF16">
        <property type="entry name" value="ZINC TRANSPORTER ZIP13"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02535">
        <property type="entry name" value="Zip"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0406">Ion transport</keyword>
    <keyword evidence="6 11" id="KW-0472">Membrane</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="12" id="KW-0732">Signal</keyword>
    <keyword evidence="3 11" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 11" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="4" id="KW-0813">Transport</keyword>
    <keyword evidence="4" id="KW-0862">Zinc</keyword>
    <keyword evidence="4" id="KW-0864">Zinc transport</keyword>
    <feature type="signal peptide" evidence="12">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019588355" description="Zinc transporter ZIP13" evidence="12">
        <location>
            <begin position="24"/>
            <end position="364"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="58"/>
            <end position="79"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="99"/>
            <end position="116"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="136"/>
            <end position="154"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006939"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00034634"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00040592"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00042542"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00042972"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007423.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="364" mass="39017" checksum="3BB346AE8EC8EAFA" modified="2019-05-08" version="1">MRGQRFLLHGMLALFLLSIPCEAQELQVDKATTGSSPLCEKEANSWGTLFSGERLDAWIFPLIGSVMVGLSGVFPLLVVPLETGAALRSEVGSNRLKQLLSFAIGGLLGNVFLHLLPEAWAYTCSATAGGGQSLQQQKLLGLWVIIGFLTFLALEKMFLESEKQEWPSVGNNSKAPSGGIPNGNSFSLQKVAGTVQRAETNIAQCNGSSLSSCPPDRRIKISGYLNLLANTIDNFTHGLAVAASFLVSRKVGLLTTMAILLHEIPHEVGDFAILLRAGFDRWSAAKMQLSTALGGILGTCFAICSQSPKETGETVAWILPFTSGGFLYIALVNVVPDLLEEKNPWQLKDCPEKGYTSLSGLANC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ51</accession>
    <name>A0A452GZ51_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Zinc finger protein RFP-like</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GZ51"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008469.1">
        <property type="protein sequence ID" value="ENSGAGP00000007326.1"/>
        <property type="gene ID" value="ENSGAGG00000005881.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd16594">
        <property type="entry name" value="RING-HC_TRIM7-like_C-IV"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12888">
        <property type="entry name" value="SPRY_PRY_TRIM7_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.920:FF:000004">
        <property type="entry name" value="Butyrophilin subfamily 1 member A1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.920">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001870">
        <property type="entry name" value="B30.2/SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043136">
        <property type="entry name" value="B30.2/SPRY_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003879">
        <property type="entry name" value="Butyrophylin_SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006574">
        <property type="entry name" value="PRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003877">
        <property type="entry name" value="SPRY_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050143">
        <property type="entry name" value="TRIM/RBCC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001841">
        <property type="entry name" value="Znf_RING"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017907">
        <property type="entry name" value="Znf_RING_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24103">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE TRIM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13765">
        <property type="entry name" value="PRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00622">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15227">
        <property type="entry name" value="zf-C3HC4_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01407">
        <property type="entry name" value="BUTYPHLNCDUF"/>
    </dbReference>
    <dbReference type="SMART" id="SM00589">
        <property type="entry name" value="PRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00184">
        <property type="entry name" value="RING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00449">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57850">
        <property type="entry name" value="RING/U-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50188">
        <property type="entry name" value="B302_SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00518">
        <property type="entry name" value="ZF_RING_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50089">
        <property type="entry name" value="ZF_RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0862">Zinc</keyword>
    <keyword evidence="2 4" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="RING-type" evidence="5">
        <location>
            <begin position="8"/>
            <end position="49"/>
        </location>
    </feature>
    <feature type="domain" description="B30.2/SPRY" evidence="6">
        <location>
            <begin position="64"/>
            <end position="256"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50188"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007326.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="9">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="256" mass="28776" checksum="4398C417A4B1708E" modified="2019-05-08" version="1">MAGTEATCPVCLEYFTEPVTVECGHNFCRDCISQCWEGSDTDISCPQCRETVQQRNLRPNRQLANMVEITKHMEKLKKLQTMNVTLDPDTAHPQLVLSEDRKSVRWGDTQQRLPDNPERFDYDPCVLGCEGFTLGRHCWEVEVGDGGCWALGVARESVGRKGQISCSPERGIWAVEWLGQFQALTSPVTPLCLNQAPSRIRVCLDCDQGQVTFIDAGDEAPIFTFPLGSVPGERIRPWLWAWLSDISVLLLFVITQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ53</accession>
    <name>A0A452GZ53_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="19">Phospholipid-transporting ATPase</fullName>
            <ecNumber evidence="19">7.6.2.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="15 19">
            <text>ATP + H2O + phospholipidSide 1 = ADP + phosphate + phospholipidSide 2.</text>
            <dbReference type="EC" id="7.6.2.1"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 18">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Golgi apparatus</location>
            <location evidence="2">trans-Golgi network membrane</location>
            <topology evidence="2">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="19">Membrane</location>
            <topology evidence="19">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 19">Belongs to the cation transport ATPase (P-type) (TC 3.A.3) family. Type IV subfamily.</text>
    </comment>
    <dbReference type="EC" id="7.6.2.1" evidence="19"/>
    <dbReference type="AlphaFoldDB" id="A0A452GZ53"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008593.1">
        <property type="protein sequence ID" value="ENSGAGP00000007438.1"/>
        <property type="gene ID" value="ENSGAGG00000000721.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005768">
        <property type="term" value="C:endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005802">
        <property type="term" value="C:trans-Golgi network"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140326">
        <property type="term" value="F:ATPase-coupled intramembrane lipid transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000287">
        <property type="term" value="F:magnesium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006897">
        <property type="term" value="P:endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045332">
        <property type="term" value="P:phospholipid translocation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006890">
        <property type="term" value="P:retrograde vesicle-mediated transport, Golgi to endoplasmic reticulum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.1110.10:FF:000008">
        <property type="entry name" value="Phospholipid-transporting ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1000:FF:000009">
        <property type="entry name" value="Phospholipid-transporting ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.1110.10">
        <property type="entry name" value="Calcium-transporting ATPase, cytoplasmic domain N"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1110.10">
        <property type="entry name" value="Calcium-transporting ATPase, transmembrane domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1000">
        <property type="entry name" value="HAD superfamily/HAD-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023299">
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        <property type="entry name" value="ATPase_P-typ_P_site"/>
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        <property type="entry name" value="ATPase_P-typ_TM_dom_sf"/>
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        <property type="entry name" value="HAD-like_sf"/>
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        <property type="entry name" value="HAD_sf"/>
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    <dbReference type="InterPro" id="IPR006539">
        <property type="entry name" value="P-type_ATPase_IV"/>
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    <dbReference type="InterPro" id="IPR001757">
        <property type="entry name" value="P_typ_ATPase"/>
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        <property type="entry name" value="P_typ_ATPase_c"/>
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    <dbReference type="InterPro" id="IPR044492">
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        <property type="entry name" value="ATPase-Plipid"/>
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        <property type="entry name" value="PHOSPHOLIPID-TRANSPORTING ATPASE IIB-RELATED"/>
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        <property type="entry name" value="Hydrolase"/>
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        <property type="entry name" value="PhoLip_ATPase_C"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="CATATPASE"/>
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    <dbReference type="SFLD" id="SFLDG00002">
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        <property type="entry name" value="Haloacid_Dehalogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="p-type_atpase"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Calcium ATPase, transmembrane domain M"/>
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    </dbReference>
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        <property type="entry name" value="HAD-like"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Metal cation-transporting ATPase, ATP-binding domain N"/>
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        <property type="entry name" value="ATPASE_E1_E2"/>
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    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 17 19" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="12" id="KW-0333">Golgi apparatus</keyword>
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    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
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        <location>
            <begin position="56"/>
            <end position="76"/>
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    <feature type="transmembrane region" description="Helical" evidence="19">
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            <begin position="82"/>
            <end position="101"/>
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            <end position="611"/>
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            <end position="714"/>
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            <end position="741"/>
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            <end position="773"/>
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            <begin position="555"/>
            <end position="782"/>
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            <position position="141"/>
        </location>
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        <location>
            <position position="141"/>
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        <ligand>
            <name>ATP</name>
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        </ligand>
    </feature>
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        <location>
            <position position="141"/>
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        <ligand>
            <name>Mg(2+)</name>
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    <sequence length="790" mass="88705" checksum="F86095ADFD24A166" modified="2019-05-08" version="1">LKVAVSCTQRLPALGDLFSISAYVYAQKPQLDIHSFEGTFTRVGLLDLELNQLTKALFLALVALSIVMVTLQGFVGPWYRNLFRFLLLFSYIIPISLRVNLDMGKAAYGWMIMKDDNIPGTVVRTSTIPEELGRLVYLLTDKTGTLTQNEMIFKRLHLGTVSYGTDTMDEIQNHIISSYSQSSAPKVRKTVSSRIHEAVKAIALCHNVTPVYESHAGVNGETEYAEVDQNFSDENRTYQASSPDEVALVQWTESVGLTLVNRDLTSMQLKSPGGQILTYYILQVFPFTSESKRMGIIVRDESTGEITFYMKGADVAMSTIVQYNDWLEEECGNMAREGLRTLVVAKKSLTEEQHQDFENRYNQAKLSIHDRALKVAAVVESLEREMELLCLTGVEDQLQADVRPTLEMLRNAGIKIWMLTGDKLETATCIAKSSHLVSRTQDIHIFRPVISRGEAHLELNAFRRKHDCALVISGDSLEVCLKYYEHEFVELACQCPAVVCCRCSPTQKAHIVKLLQHHTGKRTCAIGDGGNDVSMIQAADCGIGIEGKEGKQASLAADFSITQFKHIGRLLMVHGRNSYKRSAALGQFVMHRGLIISTMQAVFSSVFYFASIPLYQGFLMVGYATIYTMFPVFSLVLDQDVKPEMALLYPELYKDLTKGRSLSFKTFLIWVLISIYQGGILMYGALLLFESEFVHVVAISFTALILTELLMVALTIRTWHWLMVVAEFFSLGCYVASLAFLNEYFDVAFITTVTFLWKVSAITVVSCLPLYILKYLKRKFSPPSYSKLTS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ56</accession>
    <name>A0A452GZ56_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Cytochrome P450 family 2 subfamily B member 6</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="1 6">
            <name>heme</name>
            <dbReference type="ChEBI" id="CHEBI:30413"/>
        </cofactor>
    </comment>
    <comment type="similarity">
        <text evidence="2 7">Belongs to the cytochrome P450 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZ56"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008552.1">
        <property type="protein sequence ID" value="ENSGAGP00000007403.1"/>
        <property type="gene ID" value="ENSGAGG00000005769.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0020037">
        <property type="term" value="F:heme binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005506">
        <property type="term" value="F:iron ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016712">
        <property type="term" value="F:oxidoreductase activity, acting on paired donors, with incorporation or reduction of molecular oxygen, reduced flavin or flavoprotein as one donor, and incorporation of one atom of oxygen"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006082">
        <property type="term" value="P:organic acid metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006805">
        <property type="term" value="P:xenobiotic metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="entry name" value="Cytochrome P450, family 2, subfamily k, polypeptide 20"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Uncharacterized protein"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Cytochrome P450"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Cyt_P450"/>
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    <dbReference type="InterPro" id="IPR017972">
        <property type="entry name" value="Cyt_P450_CS"/>
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    <dbReference type="InterPro" id="IPR002401">
        <property type="entry name" value="Cyt_P450_E_grp-I"/>
    </dbReference>
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        <property type="entry name" value="Cyt_P450_sf"/>
    </dbReference>
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        <property type="entry name" value="Cytochrome_P450_fam2"/>
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        <property type="entry name" value="CYTOCHROME P450"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="CYTOCHROME P450 508A4-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00067">
        <property type="entry name" value="p450"/>
        <property type="match status" value="2"/>
    </dbReference>
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        <property type="entry name" value="EP450I"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00385">
        <property type="entry name" value="P450"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48264">
        <property type="entry name" value="Cytochrome P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00086">
        <property type="entry name" value="CYTOCHROME_P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 6 7" id="KW-0349">Heme</keyword>
    <keyword evidence="5 6 7" id="KW-0408">Iron</keyword>
    <keyword evidence="8" id="KW-0472">Membrane</keyword>
    <keyword evidence="4 6 7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7" id="KW-0503">Monooxygenase</keyword>
    <keyword evidence="7" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="6"/>
            <end position="26"/>
        </location>
    </feature>
    <feature type="binding site" description="axial binding residue" evidence="6">
        <location>
            <position position="431"/>
        </location>
        <ligand>
            <name>heme</name>
            <dbReference type="ChEBI" id="CHEBI:30413"/>
        </ligand>
        <ligandPart>
            <name>Fe</name>
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        </source>
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            <dbReference type="ARBA" id="ARBA00010617"/>
        </source>
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            <dbReference type="ARBA" id="ARBA00022617"/>
        </source>
    </evidence>
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    </evidence>
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        <source>
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        </source>
    </evidence>
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        <source>
            <dbReference type="PIRSR" id="PIRSR602401-1"/>
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    </evidence>
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        <source>
            <dbReference type="RuleBase" id="RU000461"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007403.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="11">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="488" mass="55673" checksum="6EB333333187BB35" modified="2019-05-08" version="1">SSKMALADPTVILLLFVVTSVSILFVKKSWNKRTCLNFPPGPRPLPILGNLHQLNLKRPYRTLLKKYGPVFSIHMGPKNMVVLSGYKMVKEALVNQADAFAERPRIPIFEKFARGNGIIFAHGENWKVMRRFTLTTLRDFGMGRRTIENKILEECDYLIKDFESHKGSSFDTMKMMNAAVSNVIVSIMLGHRFDYEDPKLLRLIHLINESVRLSGSPMVTNLEELYSFTQQTFIEHLKQLDPNDQRSFIDAFLIRQQEEKSNPNTHFKNENLQSLVSNLFTAGMETTSTTLRWGLLLMMKYPASSGVLCPVLGPTLHERVHEEIERVIGSAQLRYEHRTQMPYTDAVIHEIQRFANILPMSLPHETTEDVRLNGYVIPKGTYIIPLLASVLYDKTQWEKPGEFCPQHFLSTDGKFVTNEAFMPFSAGRRICAGETLARMELFLFFTSLLQRFTFHPPPGVTSSDLDLTPAVGFTTPPMPHKICAMPCR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ60</accession>
    <name>A0A452GZ60_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="4">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GZ60"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008623.1">
        <property type="protein sequence ID" value="ENSGAGP00000007467.1"/>
        <property type="gene ID" value="ENSGAGG00000005981.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027955">
        <property type="entry name" value="DUF4636"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31822">
        <property type="entry name" value="SERINE-RICH SINGLE-PASS MEMBRANE PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31822:SF1">
        <property type="entry name" value="SERINE-RICH SINGLE-PASS MEMBRANE PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15468">
        <property type="entry name" value="DUF4636"/>
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    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0472">Membrane</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0732">Signal</keyword>
    <keyword evidence="2" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="2" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="3">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019399109" evidence="3">
        <location>
            <begin position="21"/>
            <end position="234"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="40"/>
            <end position="58"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="197"/>
            <end position="234"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="1">
        <location>
            <begin position="222"/>
            <end position="234"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007467.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="234" mass="26744" checksum="9B5453CBE38E2EF1" modified="2019-05-08" version="1">MGFPFLYFWWLYLHTGPVEAITSQEPDYRECDGDDEDFCGVVGQILLSYCIAIFLLLVPRDSVCMRHSSFNVWDFLSKFFNGQNKKHTCQSADPAFYSEVAAAWAYLESKQARFVSKLSQQTVCDSDFSDINTEDLDSGASSGKEGETEYIPTTASLKQRKTELLKQRNTGSHWIREQPCLHCKAKRTRKWLSQHFFNPMPPPHGKSISHGFDSKPSSGPKFTKKWTKNSSKRS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ63</accession>
    <name>A0A452GZ63_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Transcription factor IIIA</fullName>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
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        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZ63"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008606.1">
        <property type="protein sequence ID" value="ENSGAGP00000007451.1"/>
        <property type="gene ID" value="ENSGAGG00000005891.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003712">
        <property type="term" value="F:transcription coregulator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042254">
        <property type="term" value="P:ribosome biogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000032">
        <property type="entry name" value="Krueppel-like factor 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001102">
        <property type="entry name" value="Transcription factor IIIA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001998">
        <property type="entry name" value="Transcription factor IIIA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000145">
        <property type="entry name" value="Zinc finger protein 574"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR054599">
        <property type="entry name" value="TFIIIA_Zfn-C2H2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051061">
        <property type="entry name" value="Zinc_finger_trans_reg"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46179:SF1">
        <property type="entry name" value="TRANSCRIPTION FACTOR IIIA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46179">
        <property type="entry name" value="ZINC FINGER PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22110">
        <property type="entry name" value="TFIIIA_zf-C2H2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="6"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="9" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0690">Ribosome biogenesis</keyword>
    <keyword evidence="7" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="10" id="KW-0804">Transcription</keyword>
    <keyword evidence="8" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 13" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="33"/>
            <end position="62"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="63"/>
            <end position="92"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="116"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="172"/>
            <end position="199"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="201"/>
            <end position="231"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="232"/>
            <end position="261"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="300"/>
            <end position="319"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022517"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00040434"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007451.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="319" mass="36693" checksum="DE62D5D6F4994A2F" modified="2019-05-08" version="1">AAPRITAHHVSRRLPVRGLMGESRPPADCAKRFICSFPDCEATFSKAWRLNAHLCRHTGERPFICNYNGCGKGFTRDFHLTRHFLTHSGEKPFDKSFRKHQQLKVHHCQHTSEPFFKCSHEACGKHFLTPSRLKRHEKIHEGYACKKDNCSFVGKTWTEHLKHMKDCHAEPVICGVCSKTFKRKDYLRQHQKTHAEEREVCRCPREGCGRTYTTVFNLQSHILSFHEEKKPFFCDYAGCGKVFAMKQSLARHAVVHDPDKGKLNLKAKQTRPKRSLASRLSGYIPPKAQRRKLEAATENKALNKPAESELPTVETLTLH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ67</accession>
    <name>A0A452GZ67_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Tyrosine-protein kinase receptor</fullName>
            <ecNumber evidence="20">2.7.10.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="17">Receptor with tyrosine-protein kinase activity. Functions as a pH sensing receptor which is activated by increased extracellular pH. Activates an intracellular signaling pathway that involves IRS1 and AKT1/PKB.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="16">
            <text>L-tyrosyl-[protein] + ATP = O-phospho-L-tyrosyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10596"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.10.1"/>
        </reaction>
    </comment>
    <comment type="subunit">
        <text evidence="18">Probable tetramer of 2 alpha and 2 beta chains linked by disulfide bonds. The alpha chains contribute to the formation of the ligand-binding domain, while the beta chains carry the kinase domain.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="20">Belongs to the protein kinase superfamily. Tyr protein kinase family. Insulin receptor subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.10.1" evidence="20"/>
    <dbReference type="AlphaFoldDB" id="A0A452GZ67"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008565.1">
        <property type="protein sequence ID" value="ENSGAGP00000007416.1"/>
        <property type="gene ID" value="ENSGAGG00000005944.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030424">
        <property type="term" value="C:axon"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005899">
        <property type="term" value="C:insulin receptor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005009">
        <property type="term" value="F:insulin receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043560">
        <property type="term" value="F:insulin receptor substrate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="CDD" id="cd00064">
        <property type="entry name" value="FU"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05032">
        <property type="entry name" value="PTKc_InsR_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000050">
        <property type="entry name" value="Tyrosine-protein kinase receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.220.10:FF:000014">
        <property type="entry name" value="Tyrosine-protein kinase receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000087">
        <property type="entry name" value="Tyrosine-protein kinase receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000108">
        <property type="entry name" value="Tyrosine-protein kinase receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000026">
        <property type="entry name" value="Tyrosine-protein kinase receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.20.20:FF:000001">
        <property type="entry name" value="Tyrosine-protein kinase receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.220.10">
        <property type="entry name" value="Hormone Receptor, Insulin-like Growth Factor Receptor 1, Chain A, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.20.20">
        <property type="entry name" value="Receptor L-domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006211">
        <property type="entry name" value="Furin-like_Cys-rich_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006212">
        <property type="entry name" value="Furin_repeat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009030">
        <property type="entry name" value="Growth_fac_rcpt_cys_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000494">
        <property type="entry name" value="Rcpt_L-dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036941">
        <property type="entry name" value="Rcpt_L-dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050122">
        <property type="entry name" value="RTK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001245">
        <property type="entry name" value="Ser-Thr/Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008266">
        <property type="entry name" value="Tyr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020635">
        <property type="entry name" value="Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002011">
        <property type="entry name" value="Tyr_kinase_rcpt_2_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416:SF338">
        <property type="entry name" value="INSULIN RECEPTOR-RELATED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416">
        <property type="entry name" value="TYROSINE-PROTEIN KINASE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00757">
        <property type="entry name" value="Furin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07714">
        <property type="entry name" value="PK_Tyr_Ser-Thr"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01030">
        <property type="entry name" value="Recep_L_domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00109">
        <property type="entry name" value="TYRKINASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00261">
        <property type="entry name" value="FU"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00219">
        <property type="entry name" value="TyrKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57184">
        <property type="entry name" value="Growth factor receptor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52058">
        <property type="entry name" value="L domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00109">
        <property type="entry name" value="PROTEIN_KINASE_TYR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00239">
        <property type="entry name" value="RECEPTOR_TYR_KIN_II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9 19" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="13" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="15" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="8" id="KW-0418">Kinase</keyword>
    <keyword evidence="11" id="KW-0472">Membrane</keyword>
    <keyword evidence="7 19" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="2 20" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="14 20" id="KW-0675">Receptor</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <keyword evidence="3" id="KW-0808">Transferase</keyword>
    <keyword evidence="4 20" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="10" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="12" id="KW-0829">Tyrosine-protein kinase</keyword>
    <feature type="domain" description="Fibronectin type-III" evidence="23">
        <location>
            <begin position="333"/>
            <end position="453"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="23">
        <location>
            <begin position="680"/>
            <end position="775"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="22">
        <location>
            <begin position="824"/>
            <end position="1099"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="21">
        <location>
            <begin position="582"/>
            <end position="616"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="21">
        <location>
            <begin position="583"/>
            <end position="599"/>
        </location>
    </feature>
    <feature type="binding site" evidence="19">
        <location>
            <position position="858"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023137"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00051243"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00055220"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00065424"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="RuleBase" id="RU000312"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007416.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1136" mass="128617" checksum="582A6E4E4C6EC4E0" modified="2019-05-08" version="1">MMHCSHAATMMHCLPSSKGSIMGDVVCPGSLAKQGKWELLPAVLREKPYNERNSSPRNGLCPCGDGIACTAAGECCHPECLGGCSRPWDSRACVACRHFHFNGHCLTSCPTRTYRYEGWRCVTAEYCASLRKVSENPRDASEFVIHHERCLSECPSGYTRNDSSIFCHKCEGLCPKECKVGTKTIDSTRAAQELAGCTYVEGNLIVNIRRGYNLASELQNSLGLIETITGFLKIKHSFALVSLSFFKNLKLIRGDSMVDGNYTLYVLDNQNLQQLWDWSHHTLSIPVGKVYFAFNPKLCLSEIYRMEEVTGTKGRQNKAEINPRTNGDRASCESRTLRFVSNITESDRIFLKWERYRPPEYRDLLSFIVYYKESTFQNVTEYVGQDACGANSWNVVDVELPLNNDQDPGVALLNLKPWTQYAIFVRAITLTTAEEGRNYGAQSEVVYVRTMPAAPTVPRDVLSMSNSSSHIIVRWKPPSQRNGNITYYLVLWQQLAEDLELYVNDYCHKGLKLPTSSADTRFNVDDSGGPELEQDTEERCCPCHRDSGQLQMEGEAVSFQKKFENFLHNSIIIPRPPWKVTSVKKNPQSSGHPLSNASSLEPPGKPPSLNLTGAGPKAQTDFRIFEDKVVRDRTVISQLRHFTEYRIDIHACNHAAHTVGCSAATFVFARTMPELEADNIPGNVTWEPASKNSVLLRWEEPKNPNGLILKYEIKYSRENEVTTVVCVSRHRYAKYRGVHLALLQPGNYSAKVRATSLAGNGSWTGLIKFYILGPAEEESGGFYVLLTIMPIVLMVVITFWEGGNHWLWAWMYIPDEWEVSRDKITVIRELGQGSFGMVYEGIAQGIEKEDEETKVALKTVNELATMRERIEFLNEASVMKAFKCHHVVRLLGVVSQGQPALVIMELMTRGDLKSYLRSLRPDAENNPGLLPPSLKNMIQMAGEIADGMAYLNAKKFVHRDLAARNCMVSEDFTVKIGDFGMTRDIYETDYYRKGGKGLLPVRWMSPEALKDGIFNTHSDVWSFGVVLWEIATLAEQPYQGLSNEQVLRFVMDNGILKRPENCPEKLHELMSWCWQQNPRQRPSFTQLLESIKDDMDPTFRNISFFYSAENKRRGSCDHPYHHPAPRDFCPWDGWED</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ68</accession>
    <name>A0A452GZ68_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Rho guanine nucleotide exchange factor 11</fullName>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">May play a role in the regulation of RhoA GTPase by guanine nucleotide-binding alpha-12 (GNA12) and alpha-13 (GNA13). Acts as guanine nucleotide exchange factor (GEF) for RhoA GTPase and may act as GTPase-activating protein (GAP) for GNA12 and GNA13. Involved in neurotrophin-induced neurite outgrowth.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Membrane</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000008495.1">
        <property type="protein sequence ID" value="ENSGAGP00000007350.1"/>
        <property type="gene ID" value="ENSGAGG00000005896.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001664">
        <property type="term" value="F:G protein-coupled receptor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
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        <property type="term" value="F:GTPase activator activity"/>
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    <sequence length="1611" mass="177103" checksum="1A83BC1EEE1159C0" modified="2019-05-08" version="1">MSVRPPQTALDRLSSLSSLGDSTTGRRSPGHRRQPSDCTETTGLVQRCVIIQKDQDGFGFTVSGDRIVLVQSVRPGGAAMKAGVQEGDRIVKVNGTMVTNSSHLEVVKLIKSGAYVALTLLGSPPPSVVLSSPQQDVSVSGAPGIAPLCPPPPPPPPLPPPQRITGPKPLQDPEVQKHATQILRNMLRQEEAELQRFCEVYSRNPATTLEERIEGARRRVSQLQLKIQQETGSSVDLGRPCCNSSSASLRGTEGRLSLDSQDGDSGLDSGTERFPSVSEISLNRNSVLSDPGLDSPRTSPVITSKMFQHHRRQGSDTPFLPIAEQGVDRTPRPLIIGPEEDYDPGYFNNEQCDSLFQDLGKLKSRPAHLGVFLRYIFSQADPSPLLFYLCAEVCQQMNSKDSRALGKEIWNIFLEKNAPLRIKVSEQLLADIESRLRNGEDVRSAFSEAQDAAMPEIQEQIQDYRTKRTMGLGSLYGENDLLDLDGDPQKERQVAEKQLAQLGDILSKYEEDRSSPMAFALSTYMSHTGIRMREARPASTSEKSQAFPDKDKWLPFFPKAKKSSSTKKDKDVMEDKKRNPILKYIGKPKSSSQSIKPGNVRNIIQHFENNQHSESQEPGTQRLSTGSFPEELLDSDSSRSEVKLGRSESLKGREEMKKSRKAENVPRSRSDVDMDAAAEATRLHQSASSSASSLSTRSLENPTPPYTPKMGRRSIESPSLGFGTDSFLPHLLEDDLGQLSDLEPEPDSQNWQHTVGREVMADLTHREIDRQEVINELFATEASHLRILRVLDVLFYQRMKRENLLSREELGLLFPNLPDLIDIHNSLSESMKKLREEGPIIKEIGDLMLSRFDGPAREEIQQVAADFCSYQSIALELIKTKQRKETRFQIFMQEAESNPQCRRLQLKDLIISEMQRLTKYPLLLENVLKHTDGGTPEHKKLCRARDQCREILRYVNEAVKRAENRHRLEAYQKRLDATSLERTSNPLAAEFKNLDLTSRCMIHEGALSWRISKEKTLDLHVLLLEDLLVLLQKQDEKLVLKFHSKTAPGSSDTKQTFSPVLKLNSMLIRSVATDKRALFIICTSELGPQIYELVALTPSEKNTWMELLEEAVQSSMRNATSPPKRRARELPIHRPAPSSLVLQDPDVSPILSQADSSRTEVDDVSSADDNPPDFLGKGKHPVLPEEPEAEGSEEEEEEEGEEAPAAPVPAQSSAELDEAGVSEEPGPSMHLPLPGLVLSKGLAESALEDVENLRHLILRILLPCRDPEPEDDLTPTPSVAGGAAQPWDSVLSSQDSGCQEQSVEHSGRGGPTTDRAGLPGHSPSGRPEPELESESRPSAGDVEETGWSSDGGREASQEVERPPAAEGQGGYKVVRKAEVEGTKDATPLADSSQSETELHEGGGANVDGNYFYVSMPAGPAESSTEPPAPKPPASPVEQPPSPPCPALGSQQHNQDSTLGPGSLDGAPPPEPPAPSLVIQDVDMIFRTIEQLTLKLTRLKDMELAHRELLRSLGRGSSAETTPVGVTALEMGRWLECPPCPASDHCPSPARRGCRPQQPASLAAEPPRRSPSAPVPCQGDLPKDGWVAGVAGHPLDPPNLGKDPWEVGKPEC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ69</accession>
    <name>A0A452GZ69_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase</fullName>
            <ecNumber evidence="11">2.7.4.24</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="11">Bifunctional inositol kinase that acts in concert with the IP6K kinases to synthesize the diphosphate group-containing inositol pyrophosphates diphosphoinositol pentakisphosphate, PP-InsP5, and bis-diphosphoinositol tetrakisphosphate, (PP)2-InsP4. PP-InsP5 and (PP)2-InsP4, also respectively called InsP7 and InsP8, may regulate a variety of cellular processes, including apoptosis, vesicle trafficking, cytoskeletal dynamics, and exocytosis. Phosphorylates inositol hexakisphosphate (InsP6).</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="10">
            <text>1D-myo-inositol hexakisphosphate + ATP = 1-diphospho-1D-myo-inositol 2,3,4,5,6-pentakisphosphate + ADP</text>
            <dbReference type="Rhea" id="RHEA:37459"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:58130"/>
            <dbReference type="ChEBI" id="CHEBI:74946"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.4.24"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="10">
            <dbReference type="Rhea" id="RHEA:37460"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="9">
            <text>5-diphospho-1D-myo-inositol 1,2,3,4,6-pentakisphosphate + ATP + H(+) = 1,5-bis(diphospho)-1D-myo-inositol 2,3,4,6-tetrakisphosphate + ADP</text>
            <dbReference type="Rhea" id="RHEA:10276"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:58628"/>
            <dbReference type="ChEBI" id="CHEBI:77983"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.4.24"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="9">
            <dbReference type="Rhea" id="RHEA:10277"/>
        </physiologicalReaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 11">Cytoplasm</location>
            <location evidence="1 11">Cytosol</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 11">Belongs to the histidine acid phosphatase family. VIP1 subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.4.24" evidence="11"/>
    <dbReference type="AlphaFoldDB" id="A0A452GZ69"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008603.1">
        <property type="protein sequence ID" value="ENSGAGP00000007448.1"/>
        <property type="gene ID" value="ENSGAGG00000005969.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000008612.1">
        <property type="protein sequence ID" value="ENSGAGP00000007456.1"/>
        <property type="gene ID" value="ENSGAGG00000005969.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033857">
        <property type="term" value="F:5-diphosphoinositol pentakisphosphate 1-kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000828">
        <property type="term" value="F:inositol hexakisphosphate kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006020">
        <property type="term" value="P:inositol metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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    <dbReference type="GO" id="GO:0032958">
        <property type="term" value="P:inositol phosphate biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07061">
        <property type="entry name" value="HP_HAP_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.470.20:FF:000003">
        <property type="entry name" value="Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.11950:FF:000001">
        <property type="entry name" value="Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1240:FF:000013">
        <property type="entry name" value="Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.470.20">
        <property type="entry name" value="ATP-grasp fold, B domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1240">
        <property type="entry name" value="Phosphoglycerate mutase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033379">
        <property type="entry name" value="Acid_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000560">
        <property type="entry name" value="His_Pase_clade-2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037446">
        <property type="entry name" value="His_Pase_VIP1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029033">
        <property type="entry name" value="His_PPase_superfam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040557">
        <property type="entry name" value="VIP1_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12750">
        <property type="entry name" value="DIPHOSPHOINOSITOL PENTAKISPHOSPHATE KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12750:SF10">
        <property type="entry name" value="INOSITOL HEXAKISPHOSPHATE AND DIPHOSPHOINOSITOL-PENTAKISPHOSPHATE KINASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00328">
        <property type="entry name" value="His_Phos_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18086">
        <property type="entry name" value="PPIP5K2_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56059">
        <property type="entry name" value="Glutathione synthetase ATP-binding domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53254">
        <property type="entry name" value="Phosphoglycerate mutase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00616">
        <property type="entry name" value="HIS_ACID_PHOSPHAT_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 11" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="3 11" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7 11" id="KW-0418">Kinase</keyword>
    <keyword evidence="6 11" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 11" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="VIP1 N-terminal" evidence="13">
        <location>
            <begin position="54"/>
            <end position="143"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1"/>
            <end position="21"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="908"/>
            <end position="992"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1009"/>
            <end position="1028"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1156"/>
            <end position="1190"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="925"/>
            <end position="955"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="12">
        <location>
            <begin position="970"/>
            <end position="981"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="1158"/>
            <end position="1168"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="1177"/>
            <end position="1190"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004514"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005609"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00033696"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00034629"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="RuleBase" id="RU365032"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="Pfam" id="PF18086"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007456.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1190" mass="134488" checksum="2F9D0E99C2FD4379" modified="2019-05-08" version="1">MSVSATESDPPRFFVGGEDGDELLDSSRSADFDHFYDHAEEEDEEYDSPPERLIVVGICSMAKKSKSKPMNEILERLSMFKYITVVIFEEDVILNEPVENWPLCDCLISFHSKGFPLDKAVAYAKLRNPFVINDLNMQYHIQDRREVYSILKAEGILLPRYAILNRDPNNPKECSLIEGEDQVEVNGEVFQKPFVEKPVSAEDHNVYIYYPTSAGGGSQRLFRKIGSRSSVYSPESNVRKTGSYIYEEFMPTDGTDVKVYTVGPDYAHAEARKSPALDGKVERDSEGKEVRYPVILNAREKLIAWKVCLAFKQTVCGFDLLRANGQSYVCDVNGFSFVKNSMKYYDDCAKILGNIIMRELAPQFQIPWSIPLEAEDIPIVPTTSGTMMELRCVIAVIRHGDRTPKQKMKMEVKHQKFFDLFEKCDGYKSGKLKLKKPKQLQEVLDIARQLLIELGQNNDSEIEESKAKLEQLKTVLEMYGHFSGINRKVQLTYLPHGCQKTSSEEEDNRRDEPSLLLVLKWGGELTPAGRVQAEELGRAFRCMYPGGQGDYAGFPGCGLLRLHSTYRHDLKIYASDEGRVQMTAAAFAKGLLALEGELTPILVQMVKSANMNGLLDSDSDSLSSCQHRVKARLHEILQRDREFMPEDYEKLTPSGSISLMKSMKVIRNPVRTCDKVYSLIQSLTSQIRQRMEDPKSADIQLYHSETLELMLRRWAKLEKDFKTKNGRYDISKIPDIYDCIKYDVQHNGSLKLENTMELYRLSKALADIVIPQEYGISKVEKLEIAKGYCTPLVRKIRSDLQRTQDDDTVNKLHPLYSRGVMSPERHVRTRLYFTSESHVHSLLSTLRYGALCDESKDEQWKRAMDYLNIVSELNYMTQIVIMLYEDPNKELSSEERFHVELHFSPGAKGCEEDKNLPAGFGYRPASRENEGSKKTSHKNDSDEESHTSKRDETDRSIVVFKPMVSDPIHIHRKSPLPRSRKIGSVEEESPLSVSSPECIGTWLHYTSGVGTGRRRRRSGEQITSSPVSPKSLAFTSSIFGSWQQVLSDNNSNLRTPRTILEQKQSGLGFELYSMVPSICPLETLHNSLSLKQVDEFLASIAAPSSDNLPEISTALCASSAYLSSLSGRKCSLNTYNPAKIQPTPLAALPERLAIEKPTLSTSGPSNHISDVAPSPKESVKCGDETLTEKK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKP2</accession>
    <name>A0A452HKP2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">CDK5 regulatory subunit-associated protein 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Golgi apparatus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000017700.1">
        <property type="protein sequence ID" value="ENSGAGP00000015496.1"/>
        <property type="gene ID" value="ENSGAGG00000011684.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035371">
        <property type="term" value="C:microtubule plus-end"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097431">
        <property type="term" value="C:mitotic spindle pole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000242">
        <property type="term" value="C:pericentriolar material"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043015">
        <property type="term" value="F:gamma-tubulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008017">
        <property type="term" value="F:microtubule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007099">
        <property type="term" value="P:centriole replication"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007059">
        <property type="term" value="P:chromosome segregation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000132">
        <property type="term" value="P:establishment of mitotic spindle orientation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001578">
        <property type="term" value="P:microtubule bundle formation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046600">
        <property type="term" value="P:negative regulation of centriole replication"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090266">
        <property type="term" value="P:regulation of mitotic cell cycle spindle assembly checkpoint"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042791">
        <property type="entry name" value="CDK5RAP2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056273">
        <property type="entry name" value="CDK5RAP2_MYOME_CC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012943">
        <property type="entry name" value="Cnn_1N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46930">
        <property type="entry name" value="CDK5 REGULATORY SUBUNIT-ASSOCIATED PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46930:SF1">
        <property type="entry name" value="CDK5 REGULATORY SUBUNIT-ASSOCIATED PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23246">
        <property type="entry name" value="CC_CDK5RAP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07989">
        <property type="entry name" value="Cnn_1N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="7" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Centrosomin N-terminal motif 1" evidence="11">
        <location>
            <begin position="57"/>
            <end position="130"/>
        </location>
    </feature>
    <feature type="domain" description="CDK5 regulatory subunit-associated protein 2/Myomegalin coiled coil" evidence="12">
        <location>
            <begin position="501"/>
            <end position="620"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="928"/>
            <end position="965"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="58"/>
            <end position="128"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="155"/>
            <end position="182"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="276"/>
            <end position="454"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="491"/>
            <end position="618"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="659"/>
            <end position="696"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="1454"/>
            <end position="1485"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="1552"/>
            <end position="1586"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="1656"/>
            <end position="1694"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="931"/>
            <end position="940"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004555"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00020479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF07989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF23246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015496.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1920" mass="221960" checksum="C718E8F143AFF84A" modified="2019-05-08" version="1">MDSLPGEDLTLPVEFSGSVSISRLSDVSRDEPRIGLGISGALLDVAEDAISPTRARTMKDYENQITDLKKENFNLKLRIYFLEERMQQKFDGPTEDIYKINIELKVEVESLKRDLQEREKLLIKASKAVESLAQGGDAEIQRVKEEARKKVQQVEEILTSRINLLEENLKAAQEEVEKAFAVTEREKVLRMAVEQKLSGVTNTHLKELDSITELPEKDRLIEQLTLSLKSKEAIIQHLEEEKSQSESCDGNLPAEKIRELTVALRHEKDSEIEAIKMELRNERNCFEKKVQSLQEELREREDELSIEKKNGLKRDKTIQGLTLALKTKEKENEELNSEIENLNVSLAKAREATHKAQIQKFKGAEDYQALLMEKETLLAELRSENLTKDTENRKLQRRIKRTDQELNDLNLEREKMERELDEAQLQKSRGDKTINDLRNQLEKLHGEMTEREKAVEHHYNILLSESNQKLQSQELVIKRLTDGVSQKDLLLQKFDAAIKEKDVELQELIKKCKNLLKAKEDLELKNEDLIKEKCASESQQSIIKIVKELEEKKESEYEELILVLRKEQHIYSTLVKTLKDSDSINSLQTELNNIFLLRKQLEEDILANRNLRKVLEDQIKNTKNREDETFSFCGDQTSYMSICLGEHDHLNLQVDHLSLEELKKKVAELILMVKDLHSVNQELKKKQLRLSRSETQGKETRKIINQNEFLERTEETMTQAGDFDNKNAESCQCSERTSEACPQLPLDHPLDLDMQIPPSDYSDGVCKDRHYRNPNKPWFEHDIAGKHFGRIEMGEHEVEEMSLLTSLLNENGTSVLESRQEQMKIVIALLDHLSTTEQDCSSEDIENKDEKDLRQMIIQLRAELRHFKQVNKILKQQVELKSTSDGKEKLYPDTAPQVNKDIELLKVELKDTATQTMTIEGNVMRLKHEGKRGASEEKTKYSRLNAEPQQENAYKKGEQHESLSVTSRTNKSDFIYPFKKSRLPIPLKSSRSLGNIPLTSCTQKPDMHLQHRLMALDEGLKERKIENKQLHKELSLCGSVIETVQEKAEEERVLVRAESLVRMDCKEIPVDSQAAGFDTHDKNENKMVTDQRTSQETETPHNISDMSILHSLKELNKEFYSMENQDIDSATTNSGSKSLLSLKLSVPSTDAFDDYEVIDDLEELRQRIKDMKSELEKYKMFIFHLQPTNPFLSSDIFNIVLNDAELPIERHVIQMQDTDGQETQTFSTVHQQVDYEIRIENLNSQSAETPDEQTSQNLKQLLLENEAELEREQIANMHLLDEVYRLQNKLRETSPSKYDSLVHSQARELSFQRQQIKEIHSICVTYHQHLTSLIKAFEELLQASDVDYYVAEGFREQLNQSVQLFEKLEKKFLYGESLDAEVAKLYELAQRDNETTTTMHQQLKDESPIPSAFHSLSDFEMSEKSSIGSPEPKQDLVEEDKLALVPSKFPPALLMEHLQEIRSLRQRLEDSIKTNERLRLQLERQVTDMELNQGSANIFIHGSEQHNSLTSEIHFLRKQNKVLNVMLAKGSRDKQKENEKLRESLSKKNAIIEHLHQDYECFKKENEKLQKQIGKKEDENRHLTHEIYSSHNELNRLQTEMNVKQHQLSENDKLLQSLRVELKVYEKLDEARRRKTDILDPGHEEFQKDQNNPLDLHELLTEIQSLRVQLERSIETNKTLHEKLEEQLSKEKREEAASVSAVNINYLLKRDSQHYTRMNEVGHKFSDPSSNVCELQNCGHGTSPGKAGTPKDKNIINKQMFCIPETQLAAKDDLDSSSRCSSSSSTGTSCTPRLVPGHRMWADKNGRHILGLIEDYNALRKQISEGQKLVSEMEAPLREITGMKNLEPGVKDEGMKAEILRLHKKLSEQEKKLHSTVKRLHSTNQLKENMEKVIIDQLALTHDVLKKARGNLEVSKICLH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKP5</accession>
    <name>A0A452HKP5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">Ubiquitin-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HKP5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017693.1">
        <property type="protein sequence ID" value="ENSGAGP00000015489.1"/>
        <property type="gene ID" value="ENSGAGG00000011692.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="CDD" id="cd17051">
        <property type="entry name" value="Ubl2_ANKUB1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042788">
        <property type="entry name" value="ANKUB1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000626">
        <property type="entry name" value="Ubiquitin-like_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46885">
        <property type="entry name" value="PROTEIN ANKUB1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46885:SF1">
        <property type="entry name" value="PROTEIN ANKUB1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50053">
        <property type="entry name" value="UBIQUITIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Ubiquitin-like" evidence="1">
        <location>
            <begin position="25"/>
            <end position="58"/>
        </location>
    </feature>
    <evidence type="ECO:0000259" key="1">
        <source>
            <dbReference type="PROSITE" id="PS50053"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015489.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="498" mass="56578" checksum="01D4CC3DB76A389B" modified="2019-05-08" version="1">MWKVRQDYFDIPLSEDKQGRRYLELIYAGAILKDSWILADIGISVSSTIKCVVKEEDKPAFYVYNAVTQEKVPIKGSFYLLATKVSLLKTLVTLKCGFPNSVYCLRTPEGREMYDCNTLNDYQLDIGTTLRLDVWDGWKEFLTGCLLGNKHTVQCFLSDEEPVLKYQKRVALYMAAFFGHLELAAWILKQGVRPNAAVGVHPYREWCQETHHPDVTKCPVHAAAEAGQLIILKAFVNYSVLCLECQNPAGQIPLNMCIKHKHKDCVLYLVTKMWSAVSYPNFSLPMKIYIKLKQWLLRAQSHILTTKWLNQAAVFRTRVGDTVIVDGFTKSNMTSKARIKAAGNRDYKLPNLTDQTPHGKESCPLTVTKGEQKVIKLKLPPWEDANVYRLHQKKNIRKKAAQPRKDESMDQNMCLAKVPLPPISSLRNSHPPFYYSTPNAKLLLNSSSECFSEHNGRTPRDNAIYCLAIASAFKEKPWLQQLGIARTLAKKSVCKPVC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKP7</accession>
    <name>A0A452HKP7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">CDK5 regulatory subunit-associated protein 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Golgi apparatus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000017707.1">
        <property type="protein sequence ID" value="ENSGAGP00000015502.1"/>
        <property type="gene ID" value="ENSGAGG00000011684.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035371">
        <property type="term" value="C:microtubule plus-end"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097431">
        <property type="term" value="C:mitotic spindle pole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000242">
        <property type="term" value="C:pericentriolar material"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043015">
        <property type="term" value="F:gamma-tubulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008017">
        <property type="term" value="F:microtubule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007099">
        <property type="term" value="P:centriole replication"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007059">
        <property type="term" value="P:chromosome segregation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000132">
        <property type="term" value="P:establishment of mitotic spindle orientation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001578">
        <property type="term" value="P:microtubule bundle formation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046600">
        <property type="term" value="P:negative regulation of centriole replication"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090266">
        <property type="term" value="P:regulation of mitotic cell cycle spindle assembly checkpoint"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042791">
        <property type="entry name" value="CDK5RAP2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056273">
        <property type="entry name" value="CDK5RAP2_MYOME_CC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012943">
        <property type="entry name" value="Cnn_1N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46930">
        <property type="entry name" value="CDK5 REGULATORY SUBUNIT-ASSOCIATED PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46930:SF1">
        <property type="entry name" value="CDK5 REGULATORY SUBUNIT-ASSOCIATED PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23246">
        <property type="entry name" value="CC_CDK5RAP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07989">
        <property type="entry name" value="Cnn_1N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="7" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Centrosomin N-terminal motif 1" evidence="11">
        <location>
            <begin position="57"/>
            <end position="130"/>
        </location>
    </feature>
    <feature type="domain" description="CDK5 regulatory subunit-associated protein 2/Myomegalin coiled coil" evidence="12">
        <location>
            <begin position="501"/>
            <end position="620"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="928"/>
            <end position="965"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="58"/>
            <end position="128"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="155"/>
            <end position="182"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="276"/>
            <end position="454"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="491"/>
            <end position="618"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="659"/>
            <end position="696"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="1454"/>
            <end position="1485"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="1552"/>
            <end position="1586"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="1656"/>
            <end position="1694"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="931"/>
            <end position="940"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004555"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00020479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF07989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF23246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015502.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1867" mass="215775" checksum="DEC58B6A64666CCB" modified="2019-05-08" version="1">MDSLPGEDLTLPVEFSGSVSISRLSDVSRDEPRIGLGISGALLDVAEDAISPTRARTMKDYENQITDLKKENFNLKLRIYFLEERMQQKFDGPTEDIYKINIELKVEVESLKRDLQEREKLLIKASKAVESLAQGGDAEIQRVKEEARKKVQQVEEILTSRINLLEENLKAAQEEVEKAFAVTEREKVLRMAVEQKLSGVTNTHLKELDSITELPEKDRLIEQLTLSLKSKEAIIQHLEEEKSQSESCDGNLPAEKIRELTVALRHEKDSEIEAIKMELRNERNCFEKKVQSLQEELREREDELSIEKKNGLKRDKTIQGLTLALKTKEKENEELNSEIENLNVSLAKAREATHKAQIQKFKGAEDYQALLMEKETLLAELRSENLTKDTENRKLQRRIKRTDQELNDLNLEREKMERELDEAQLQKSRGDKTINDLRNQLEKLHGEMTEREKAVEHHYNILLSESNQKLQSQELVIKRLTDGVSQKDLLLQKFDAAIKEKDVELQELIKKCKNLLKAKEDLELKNEDLIKEKCASESQQSIIKIVKELEEKKESEYEELILVLRKEQHIYSTLVKTLKDSDSINSLQTELNNIFLLRKQLEEDILANRNLRKVLEDQIKNTKNREDETFSFCGDQTSYMSICLGEHDHLNLQVDHLSLEELKKKVAELILMVKDLHSVNQELKKKQLRLSRSETQGKETRKIINQNEFLERTEETMTQAGDFDNKNAESCQCSERTSEACPQLPLDHPLDLDMQIPPSDYSDGVCKDRHYRNPNKPWFEHDIAGKHFGRIEMGEHEVEEMSLLTSLLNENGTSVLESRQEQMKIVIALLDHLSTTEQDCSSEDIENKDEKDLRQMIIQLRAELRHFKQVNKILKQQVELKSTSDGKEKLYPDTAPQVNKDIELLKVELKDTATQTMTIEGNVMRLKHEGKRGASEEKTKYSRLNAEPQQENAYKKGEQHESLSVTSRTNKSDFIYPFKKSRLPIPLKSSRSLGNIPLTSCTQKPDMHLQHRLMALDEGLKERKIENKQLHKELSLCGSVIETVQEKAEEERVLVRAESLVRMDCKEIPVDSQAAGFDTHDKNENKMVTDQRTSQETETPHNISDMSILHSLKELNKEFYSMENQDIDSATTNSGSKSLLSLKLSVPSTDAFDDYEVIDDLEELRQRIKDMKSELEKYKMFIFHLQPTNPFLSSDIFNIVLNDAELPIERHVIQMQDTDGQETQTFSTVHQQVDYEIRIENLNSQSAETPDEQTSQNLKQLLLENEAELEREQIANMHLLDEVYRLQNKLRETSPSKYDSLVHSQARELSFQRQQIKEIHSICVTYHQHLTSLIKAFEELLQASDVDYYVAEGFREQLNQSVQLFEKLEKKFLYGESLDAEVAKLYELAQRDNETTTTMHQQLKDESPIPSAFHSLSDFEMSEKSSIGSPEPKQDLVEEDKLALVPSKFPPALLMEHLQEIRSLRQRLEDSIKTNERLRLQLERQVTDMELNQGSANIFIHGSEQHNSLTSEIHFLRKQNKVLNVMLAKGSRDKQKENEKLRESLSKKNAIIEHLHQDYECFKKENEKLQKQIGKKEDENRHLTHEIYSSHNELNRLQTEMNVKQHQLSENDKLLQSLRVELKVYEKLDEARRRKTDILDPGHEEFQKDQNNPLDLHELLTEIQSLRVQLERSIETNKTLHEKLEEQLSKEKREEAASVSAVNINYLLKRDSQHYTRMNEVGHKFSDPSSNVCELQNCGHGTSPGKAGTPKDKNIINKQMFCIPETQLAAKDDLDSSSRCSSSSSTGTSCTPRLVPGHRMWADKNGRHILGLIEDYNALRKQISEGQKLVSEMEAPLREITGMKNLEPGVKVKLLLQNRSSRPGITE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKP8</accession>
    <name>A0A452HKP8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Semaphorin-6A</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="18">Semaphorin VIA</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="17">Semaphorin-6A-1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Cell surface receptor for PLXNA2 that plays an important role in cell-cell signaling. Required for normal granule cell migration in the developing cerebellum. Promotes reorganization of the actin cytoskeleton and plays an important role in axon guidance in the developing central nervous system. Can act as repulsive axon guidance cue. Has repulsive action towards migrating granular neurons. May play a role in channeling sympathetic axons into the sympathetic chains and controlling the temporal sequence of sympathetic target innervation.</text>
    </comment>
    <comment type="subunit">
        <text evidence="15">Active as a homodimer or oligomer. The SEMA6A homodimer interacts with a PLXNA2 homodimer, giving rise to a heterotetramer. Interacts with EVL.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the semaphorin family.</text>
    </comment>
    <comment type="caution">
        <text evidence="19">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKP8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017721.1">
        <property type="protein sequence ID" value="ENSGAGP00000015514.1"/>
        <property type="gene ID" value="ENSGAGG00000011703.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045499">
        <property type="term" value="F:chemorepellent activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030215">
        <property type="term" value="F:semaphorin receptor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007411">
        <property type="term" value="P:axon guidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001755">
        <property type="term" value="P:neural crest cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:2001224">
        <property type="term" value="P:positive regulation of neuron migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071526">
        <property type="term" value="P:semaphorin-plexin signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd11266">
        <property type="entry name" value="Sema_6A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:000028">
        <property type="entry name" value="semaphorin-6A isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1680.10:FF:000007">
        <property type="entry name" value="semaphorin-6A isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1680.10">
        <property type="entry name" value="ligand-binding face of the semaphorins, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002165">
        <property type="entry name" value="Plexin_repeat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001627">
        <property type="entry name" value="Semap_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036352">
        <property type="entry name" value="Semap_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027231">
        <property type="entry name" value="Semaphorin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11036">
        <property type="entry name" value="SEMAPHORIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11036:SF12">
        <property type="entry name" value="SEMAPHORIN-6A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01437">
        <property type="entry name" value="PSI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01403">
        <property type="entry name" value="Sema"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00630">
        <property type="entry name" value="Sema"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103575">
        <property type="entry name" value="Plexin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF101912">
        <property type="entry name" value="Sema domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51004">
        <property type="entry name" value="SEMA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="3" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="8" id="KW-0221">Differentiation</keyword>
    <keyword evidence="12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11 21" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7 22" id="KW-0732">Signal</keyword>
    <keyword evidence="6 21" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="10 21" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="22">
        <location>
            <begin position="1"/>
            <end position="18"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019215222" description="Semaphorin-6A" evidence="22">
        <location>
            <begin position="19"/>
            <end position="989"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="607"/>
            <end position="635"/>
        </location>
    </feature>
    <feature type="domain" description="Sema" evidence="23">
        <location>
            <begin position="24"/>
            <end position="512"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="717"/>
            <end position="741"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="821"/>
            <end position="861"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="879"/>
            <end position="926"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="952"/>
            <end position="989"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="20">
        <location>
            <begin position="881"/>
            <end position="892"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="20">
        <location>
            <begin position="893"/>
            <end position="913"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="20">
        <location>
            <begin position="976"/>
            <end position="989"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009492"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00053495"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00063062"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00074111"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00079635"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00080740"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00352"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS51004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015514.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="989" mass="110296" checksum="DF9513ED170132B6" modified="2019-05-08" version="1">MRSEALLLYFTLLHFAGAGFPEDSEPISISHGNYTKQYPVFVGHKPGRNATQRHRLDIQLIMIMNRTLYIAARDHIYTVDMDTSHTEEIYFIKKLTWKSRQADVDTCRMKGKHKDECHNFIKVLLKRNDDTLFICGTNAFNPSCRNYKMETLEFLGDEFSGMARCPYDAKHANVALFAEGKLYSATVTDFLAIDAVIYRSLGDSPTLRTVKHDSKWLKEPYFVQAVDYGDYIYFFFREIAVEYNSMGKVVFPRVAQVCKNDMGGSQRVLEKQWTSFLKARLNCSVPGDSHFYFNILQAVTDVIHINGHDIVLATFSTPYNSIPGSAVCAYDMLDIANVFTGRFKEQKSPDSTWTPVPDERVPKPRPGCCAGSVSLEKYITSNEFPDDTLNFIKTHPLMDEAVPSIVSRPWFLRTMVRYRLTKIAVDTAAGPHQNHTVVFLGSEKGIILKFLARTGNSGFLNDSLFLEEMNVYNPEKCSYDGVEDKRIMGMQLDKHSRSLYVAFSTCVIKIPLGRCERHGKCKKACIASRDPYCGWVKEGGVCIQLLSGVKLAFEQDIEHGNTDGLGDCQNSFVALNAISTASPAHEISYNTVYGVIRESYLKGHDQLVPVTLLAIAVILAFVMGAVFSGIVVYCICDHRRRDVAVVQRKEKELAHSRRGSMSSVTKLSGLFGDTQSKEPKPEAILTPLMHNGKLATPGSTAKMLIKADQHHLDLAALPTPESTPTLQQKRKPSRGSRDWERNQNLINACTKDIPAMSSPVIPTDLPLRASPSHIPSVVVLPICQQGYQHEYVDQPKMSDMAQMAMEDQNATLEYKTIKEHLSSKSPNHGVNLVENLDSMPPKVPQREASLGPPGSSLAQSGLGKRLEMHSSAYNVDYKRNYPTNSLTRNHQTSTLKRNNTNSSNSSHLSRNQSFGRGDNPPPAPQRVDSIQVHAAQAQAVTVSRQPSLTAYNSLTRSGLKRTPSLKPDVPPKPSFTPLSTSMKPNDACT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKP9</accession>
    <name>A0A452HKP9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="11">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000017717.1">
        <property type="protein sequence ID" value="ENSGAGP00000015510.1"/>
        <property type="gene ID" value="ENSGAGG00000011700.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003779">
        <property type="term" value="F:actin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000083">
        <property type="entry name" value="Uveal autoantigen with coiled-coil domains and ankyrin repeats"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.1490">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1170.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042420">
        <property type="entry name" value="RAI14/UACA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24129">
        <property type="entry name" value="ANKYCORBIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24129:SF1">
        <property type="entry name" value="UVEAL AUTOANTIGEN WITH COILED-COIL DOMAINS AND ANKYRIN REPEATS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00023">
        <property type="entry name" value="Ank"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13637">
        <property type="entry name" value="Ank_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01415">
        <property type="entry name" value="ANKYRIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="5"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5 9" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="6 10" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <feature type="repeat" description="ANK" evidence="9">
        <location>
            <begin position="54"/>
            <end position="86"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="9">
        <location>
            <begin position="87"/>
            <end position="119"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="9">
        <location>
            <begin position="120"/>
            <end position="152"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="9">
        <location>
            <begin position="153"/>
            <end position="185"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="9">
        <location>
            <begin position="186"/>
            <end position="218"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="10">
        <location>
            <begin position="282"/>
            <end position="354"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="10">
        <location>
            <begin position="419"/>
            <end position="492"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="10">
        <location>
            <begin position="519"/>
            <end position="588"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="10">
        <location>
            <begin position="616"/>
            <end position="711"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="10">
        <location>
            <begin position="740"/>
            <end position="1078"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="10">
        <location>
            <begin position="1111"/>
            <end position="1141"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="10">
        <location>
            <begin position="1166"/>
            <end position="1200"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="10">
        <location>
            <begin position="1282"/>
            <end position="1358"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015510.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1377" mass="158542" checksum="D8067144ECB69A73" modified="2019-05-08" version="1">MMSCWISCAARNTLNTDWSKYDDRLMKAAERGDVEKLSSVLAKKGVNPTKLDVEGRSAFHVVASKGNLDCLNTILVHGVDIVASDAAGRNALHLAAKYGHALCLQKLLQYNCPTENLDLHGRTALHDAAMSDCSSSIQLLCDHGASVNAKDGDGRTPLVLATQMCRPTVCQLLIDRGADLNARDKQNRTALMLGCEYGCKDAVEVLLKNGADISLVDALGHDCSYYARIGDNIDILALIKTAIEDSSKAKELMKKGQPEYKVTNVSQKRNQLYAQEVNFKQYQREQLSMQELEAENEDLKNRLREIQQEHRILLNRVSVLELQLNEEQMVADDLESEKDELRRLLTAKEKQQEESLRSIETLKSKLRHYERDYLGSGSDSGKEDMLLKQGQVFATESQTKSMLRPLELSLSCQSSSSENEILKRELDNMRISYGTAKEEISKLQKELSHKVAECKALASECERTKEESDRQIKQLEDALKDVQKRMFDSEGKVKQMQTHFLVLKDHLTNEAASGSNKLTEDLKDQLKDMKAKYEGVSAEVGKLRNQIKQNELLVVEFKRDEGRLVEENKRLQKELSMLEMKREKSGRNVLEVQGQLKEMAAKLVLSVPVEKFENMKSLLSTEVNEKAKKLVEMEREYEKLQVELRLLKREFESQKAKFSQYVRPEEHEQMRGRFERKTEELEKTISELSKNQTLQKEVERVHMDNKLLKQQIHTLTTEIKSQYVPLKLNEEMKKTSDLTIGDLTKKLAEVTQRYNENKAEAEKLMAEKTSLSETVSHFQAVYLSPEKHEKEVAALKSTVVELKRQLAELHQKYDAEQARVSKLVSENTTLRDSLNDQYVPAITHEEIKTVLNSTLEKTNRELSDLKEKTEDIKREFMRVNEENGTLKKKLRLLQTHIQTEYICVKDHESKVSTLNKSMQELQDNNTETLAKYKRGQEEILQLHAEIEAQKKELDTIQECIKSKYAPVASFEEREQNFEAVVKELKKQLLEQAQRCREWEEETQKYKQENEKLKNRILSIENDLQQNYILAEKSHEMEKMFTIRMEELNKQLRELLQKYTEIKHEKDKLQEESAKQISEALAVQTRMQGHYVPVQQFEALKRTLGHTIEDLKEDLNSKKACYDRELEKVRELQQELADLHNSSISLAEHAQMKEAFEKEIVAVKTSLKKEVEENQAKSEEISKLQSEIQSTKQALTELESKEVVDMLEYKSMKSALETQVASIAGNLASLNRKYEEMCEEALQAKKEELSAKDEKELLQSRSFCIEQEIKDQKERCDKSLTTITDLQKRIQESAKQVEAKDNKITELLNDVERLKQALNGLSQLTYTCGVPSKRQNQQVEMLQHQVKTLQQQLADADRQHQEVISIYRTHLLSAVQVC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKS1</accession>
    <name>A0A452HKS1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Mortality factor 4-like protein 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKS1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017746.1">
        <property type="protein sequence ID" value="ENSGAGP00000015534.1"/>
        <property type="gene ID" value="ENSGAGG00000011716.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035267">
        <property type="term" value="C:NuA4 histone acetyltransferase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006325">
        <property type="term" value="P:chromatin organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006310">
        <property type="term" value="P:DNA recombination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006281">
        <property type="term" value="P:DNA repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006355">
        <property type="term" value="P:regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd18983">
        <property type="entry name" value="CBD_MSL3_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.274.30:FF:000001">
        <property type="entry name" value="Mortality factor 4-like protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.140:FF:000024">
        <property type="entry name" value="Mortality factor 4-like protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.274.30">
        <property type="entry name" value="MRG domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016197">
        <property type="entry name" value="Chromo-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000953">
        <property type="entry name" value="Chromo/chromo_shadow_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008676">
        <property type="entry name" value="MRG"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038217">
        <property type="entry name" value="MRG_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026541">
        <property type="entry name" value="MRG_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025995">
        <property type="entry name" value="Tudor-knot"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10880:SF48">
        <property type="entry name" value="MORTALITY FACTOR 4 LIKE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10880">
        <property type="entry name" value="MORTALITY FACTOR 4-LIKE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05712">
        <property type="entry name" value="MRG"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11717">
        <property type="entry name" value="Tudor-knot"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF038133">
        <property type="entry name" value="HAT_Nua4_EAF3/MRG15"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00298">
        <property type="entry name" value="CHROMO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54160">
        <property type="entry name" value="Chromo domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51640">
        <property type="entry name" value="MRG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0007">Acetylation</keyword>
    <keyword evidence="4" id="KW-0156">Chromatin regulator</keyword>
    <keyword evidence="3" id="KW-0227">DNA damage</keyword>
    <keyword evidence="8" id="KW-0233">DNA recombination</keyword>
    <keyword evidence="9" id="KW-0234">DNA repair</keyword>
    <keyword evidence="2" id="KW-0341">Growth regulation</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0804">Transcription</keyword>
    <keyword evidence="6" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="Chromo" evidence="13">
        <location>
            <begin position="10"/>
            <end position="78"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="77"/>
            <end position="144"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022604"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023172"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00071326"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="SMART" id="SM00298"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015534.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="345" mass="39699" checksum="894099892362044B" modified="2019-05-08" version="1">MAPKQDPKPKFQEGERVLCFHGPLLYEAKCVKVAIKDKQVKYFIHYSGWNKNWDEWVPESRVLKYVDTNLQKQKELQKANQEQYAEGKMRGAAPGKKTSGLQQKNVEVKTKKNKQKTPGIGEGSSTSETPQPPRKKRARVDPTVESEETFMNRVEVKVKIPEELKPWLVDDWDLITRQKQIIIYFFPGIFTQAVLPIPHLKPLFYLPAKKNVDSILEDYASYKKSRGNTDNKEYAVNEVVAGIKEYFNVMLGTQLLYKFERPQYAEILADHPDAPMSQVYGAPHLLRLFVRIGAMLAYTPLDEKSLALLLNYLHDFLKYLAKNASTLFSASDYEVAPPEYHRKAV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKS3</accession>
    <name>A0A452HKS3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">protein-lysine 6-oxidase</fullName>
            <ecNumber evidence="15">1.4.3.13</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="16">
            <text>L-lysyl-[protein] + O2 + H2O = (S)-2-amino-6-oxohexanoyl-[protein] + H2O2 + NH4(+)</text>
            <dbReference type="Rhea" id="RHEA:24544"/>
            <dbReference type="Rhea" id="RHEA-COMP:9752"/>
            <dbReference type="Rhea" id="RHEA-COMP:12448"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15379"/>
            <dbReference type="ChEBI" id="CHEBI:16240"/>
            <dbReference type="ChEBI" id="CHEBI:28938"/>
            <dbReference type="ChEBI" id="CHEBI:29969"/>
            <dbReference type="ChEBI" id="CHEBI:131803"/>
            <dbReference type="EC" id="1.4.3.13"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Cu cation</name>
            <dbReference type="ChEBI" id="CHEBI:23378"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="2">
            <name>lysine tyrosylquinone residue</name>
            <dbReference type="ChEBI" id="CHEBI:20489"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Secreted</location>
            <location evidence="3">Extracellular space</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the lysyl oxidase family.</text>
    </comment>
    <comment type="caution">
        <text evidence="17">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="EC" id="1.4.3.13" evidence="15"/>
    <dbReference type="AlphaFoldDB" id="A0A452HKS3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017740.1">
        <property type="protein sequence ID" value="ENSGAGP00000015528.1"/>
        <property type="gene ID" value="ENSGAGG00000011707.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005507">
        <property type="term" value="F:copper ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003909">
        <property type="term" value="F:DNA ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004720">
        <property type="term" value="F:protein-lysine 6-oxidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030199">
        <property type="term" value="P:collagen fibril organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000001">
        <property type="entry name" value="Lysyl oxidase 4 isoform X1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000008">
        <property type="entry name" value="Lysyl oxidase homolog 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.250.10">
        <property type="entry name" value="SRCR-like domain"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016059">
        <property type="entry name" value="DNA_ligase_ATP-dep_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050912">
        <property type="entry name" value="LOX-like_protein"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001695">
        <property type="entry name" value="Lysyl_oxidase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001190">
        <property type="entry name" value="SRCR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036772">
        <property type="entry name" value="SRCR-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45817:SF5">
        <property type="entry name" value="LYSYL OXIDASE HOMOLOG 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45817">
        <property type="entry name" value="LYSYL OXIDASE-LIKE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01186">
        <property type="entry name" value="Lysyl_oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00530">
        <property type="entry name" value="SRCR"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00258">
        <property type="entry name" value="SPERACTRCPTR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00202">
        <property type="entry name" value="SR"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56487">
        <property type="entry name" value="SRCR-like"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00697">
        <property type="entry name" value="DNA_LIGASE_A1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00420">
        <property type="entry name" value="SRCR_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50287">
        <property type="entry name" value="SRCR_2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12" id="KW-0186">Copper</keyword>
    <keyword evidence="13 17" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="14" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="5" id="KW-0886">LTQ</keyword>
    <keyword evidence="7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0964">Secreted</keyword>
    <keyword evidence="8 19" id="KW-0732">Signal</keyword>
    <keyword evidence="10" id="KW-0801">TPQ</keyword>
    <feature type="signal peptide" evidence="19">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5018993490" description="protein-lysine 6-oxidase" evidence="19">
        <location>
            <begin position="23"/>
            <end position="656"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="20">
        <location>
            <begin position="29"/>
            <end position="130"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="20">
        <location>
            <begin position="146"/>
            <end position="246"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="20">
        <location>
            <begin position="260"/>
            <end position="368"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="567"/>
            <end position="586"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="596"/>
            <end position="634"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="55"/>
            <end position="119"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="68"/>
            <end position="129"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="99"/>
            <end position="109"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="171"/>
            <end position="235"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="184"/>
            <end position="245"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="215"/>
            <end position="225"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="17">
        <location>
            <begin position="336"/>
            <end position="346"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001935"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00001943"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00007492"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022477"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022772"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023008"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00038869"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00047861"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00196"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50287"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015528.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="656" mass="69505" checksum="B8AD6AB8D3C272F8" modified="2019-05-08" version="1">MLAGPIPTLLPLLLCLATPSWQQPLQIQVRLAGSRNQAGEGRLEVLYDGQWGTVCDDDFDIHVASVACRELGFESAVTWAHSAKYGQGEGPIWLDNLRCAGTERSLAECDSNGWGVSDCNHGEDAGIVCSAQHPPARSDSNPGPQVRLRAGAQVGEGRVEVLRHGQWGTVCDEKWDLLAASVVCRQLGYGTARQALAGAQLGQGLGPIHMTKVQCTGYEHLLHECSFQEAAQSGCQHEADAAVRCAMGYSGAERPGGRSVRLAGGRSPEEGVVEVLVPVGGALKWGAVCGERWGLNEAMVVCRQLGLGFASHALQETWYWQGSPEAGEVVMSGVHCTGTELSIQQCQRHGPVHCPAGGGRFSAGVSCTNSDPPHSQRPALHPTAPLLGSPLTASALLCTPQRPLLGAPQPAPCPFPTAPPAGTGWAWRSWNPPGRTPRKSSPVCEGVHPVPSPSLSPGLQRRFACANFGEQGVSAGCWDTYRHDIDCQWVDISDVGPGSYVLQVVVNPKHEVAESDFSNNVMRCQCKYDGHRIWLHGCRTGTGWGVRAGGAPVGWGGAGRAVLVPVPETGPAEGSRRPSRPSPGLRAAYSAVHTPLGRNRGHQASRGQALCPSPARPGRGVWVPHSRGSLLSPGDQYGADAVQELEQQQRLATNLI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKT1</accession>
    <name>A0A452HKT1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Rho family-interacting cell polarization regulator 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Apical cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cell projection</location>
            <location evidence="4">Filopodium</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cell projection</location>
            <location evidence="3">Stereocilium membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the RIPOR family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKT1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017760.1">
        <property type="protein sequence ID" value="ENSGAGP00000015547.1"/>
        <property type="gene ID" value="ENSGAGG00000011582.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016324">
        <property type="term" value="C:apical plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030175">
        <property type="term" value="C:filopodium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060171">
        <property type="term" value="C:stereocilium membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006935">
        <property type="term" value="P:chemotaxis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007517">
        <property type="term" value="P:muscle organ development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009968">
        <property type="term" value="P:negative regulation of signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007605">
        <property type="term" value="P:sensory perception of sound"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000191">
        <property type="entry name" value="RHO family interacting cell polarization regulator 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031780">
        <property type="entry name" value="FAM65_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026136">
        <property type="entry name" value="RIPOR3"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15829">
        <property type="entry name" value="PROTEIN KINASE PKN/PRK1, EFFECTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15829:SF2">
        <property type="entry name" value="RHO FAMILY-INTERACTING CELL POLARIZATION REGULATOR 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15903">
        <property type="entry name" value="PL48"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="14" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="7" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="18" id="KW-0966">Cell projection</keyword>
    <keyword evidence="9" id="KW-0145">Chemotaxis</keyword>
    <keyword evidence="15" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="8" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="17" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="13" id="KW-0221">Differentiation</keyword>
    <keyword evidence="12" id="KW-1009">Hearing</keyword>
    <keyword evidence="16" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0517">Myogenesis</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="11" id="KW-0734">Signal transduction inhibitor</keyword>
    <feature type="domain" description="FAM65 N-terminal" evidence="20">
        <location>
            <begin position="23"/>
            <end position="361"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="47"/>
            <end position="79"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="379"/>
            <end position="467"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="611"/>
            <end position="647"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="19">
        <location>
            <begin position="55"/>
            <end position="66"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="67"/>
            <end position="76"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="410"/>
            <end position="426"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="427"/>
            <end position="456"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="633"/>
            <end position="647"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004289"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004486"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00005744"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00013627"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022500"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022541"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022700"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022740"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="Pfam" id="PF15903"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015547.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1005" mass="111331" checksum="DCB1D9EB2AFBCDF4" modified="2019-05-08" version="1">IDRLPEIMSVGSHSFSPGGPNGIIRSQSFAGFSGLQERRSRCNSFVENSSALKKPQAKVKKTHNLGHKNSTTSKEPQPQRMEEVYRALKNGLDEYLEVHQTELDKLTAQLKDMKRNSRLVCQIKAIERYMRRLEFHISMVDELYEAYCIQRRLCDGASKMKQAFAMSPASKAARESLSEINRSYKEYTENMCTIEGELENLLGEFCIKMKGLAGFARLCPGDQYEIFMRYGRQRWKLKGKIEVNGKQSWDGEEMVFLPLIVGLISIKVTEVKGLATHTLVGSVTCETKDLFAARPQVVAVDINDLGTIKLNLEITWYPFDVEDLTPSTGNVNKASALQRRMSTYSQGTPETPTFKDYSFFSNSPDDIFENGMAGTEKRPISFSFGDIPNGDCASPPITVDSASALDPSSPEITVTPPDTNSSLDSTPESKDLKSQPEHTPVHENKKSSRVASEVHQKPPSPGSDNLFIDSSIPVSLLQETDDVSELKPVELDTYEGNITKQLVKRLTSAEAPMTPERLQCEGSISGESEGYKSYLDGSIEDAFQGLLLALEPHKEQYKEFQDLDQQVMHLDDILKCKPAVNRSRSSSSISLTVESALESFDFLNTSDFDDEDGGGDEVCNGGGGADSVFSDTETEKTSYRSDHPEARGHLSEALTEDTGVGTSVAGSPLPLTTGNESLDITIVKHLQCCTQLIQQIVFSSKTPFVVRNLLDKLSKEIVVLENLAEISIENMGNIISVTEAIPEFHKKLSLLSFWTKCTGPTGVYHTSTDKMIKQLDVTFATAVNEECPGLAETVFRILLSQILDRTEPVLSSSLPSEILTVFQYYNYFASNGVSDLGSYLLQLAKEASVVQKLQTLKDEKLLQNIAELTSSSLPPQQEVLRSLALLLTENKNEIGEAVTLLLSASAKNRHFREKALIYYCEVLTQPNLHLQKAACMALKCLQATESIKMLVTLCQSDNEEIRKVASETLLSLGEDGRLAYEQLDKFPREFVKVGGRWGTEVATAF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKU3</accession>
    <name>A0A452HKU3_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="7">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the TDE1 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000017761.1">
        <property type="protein sequence ID" value="ENSGAGP00000015548.1"/>
        <property type="gene ID" value="ENSGAGG00000011726.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005016">
        <property type="entry name" value="TDE1/TMS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10383">
        <property type="entry name" value="SERINE INCORPORATOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10383:SF5">
        <property type="entry name" value="SERINE INCORPORATOR 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03348">
        <property type="entry name" value="Serinc"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 6" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 6" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 6" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="44"/>
            <end position="62"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="98"/>
            <end position="120"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="132"/>
            <end position="152"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="164"/>
            <end position="182"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="203"/>
            <end position="228"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="240"/>
            <end position="258"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="356"/>
            <end position="378"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="398"/>
            <end position="420"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006665"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015548.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="451" mass="50133" checksum="F8B12421C26EDCCA" modified="2019-05-08" version="1">PGHRPTCVTPLSFIPQICCCCGPAPCGFCCSCCPSIKVSTGSRILYTLFHVLASTVCCLMLSRTVSEAVKQNVSATWGAWLLHLPRGMDCDILVGYSAVYRVCFGTACFYLMQAVFLINVKSSSNFRALLHNGFWFLKLLILVGLCAAAFFIPDDGFLQGTTWHYVGICGGFAFILVQLVLITAFAHTWNKNWLLGASQDKRWYLAVLLATLAFYTVASTAVSFLYTYYTHPAGCLLNKALLTLNVSLCGLLSFLSITPCVRLQQPRSSLLQASIISCYVMYLTFSALSSRPPERGQCSLGLGLGXLWAWAWAGRVTGGGAVGNRALSLDAERVCQQEEGPAGGQRIIYNEHDHVVYSYSTFHFVFFLASLYVMMTLTNCYENAVLETTFTHGSWSTFWVKVASCWACILLYFWLLLGPLCRSDLRQRWRGSPPLRILRRRRPLQRISVST</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKU6</accession>
    <name>A0A452HKU6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Mortality factor 4-like protein 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKU6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017766.1">
        <property type="protein sequence ID" value="ENSGAGP00000015553.1"/>
        <property type="gene ID" value="ENSGAGG00000011716.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035267">
        <property type="term" value="C:NuA4 histone acetyltransferase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006325">
        <property type="term" value="P:chromatin organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006310">
        <property type="term" value="P:DNA recombination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006281">
        <property type="term" value="P:DNA repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006355">
        <property type="term" value="P:regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd18983">
        <property type="entry name" value="CBD_MSL3_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.274.30:FF:000001">
        <property type="entry name" value="Mortality factor 4-like protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.140:FF:000024">
        <property type="entry name" value="Mortality factor 4-like protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.274.30">
        <property type="entry name" value="MRG domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016197">
        <property type="entry name" value="Chromo-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000953">
        <property type="entry name" value="Chromo/chromo_shadow_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008676">
        <property type="entry name" value="MRG"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038217">
        <property type="entry name" value="MRG_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026541">
        <property type="entry name" value="MRG_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025995">
        <property type="entry name" value="Tudor-knot"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10880:SF48">
        <property type="entry name" value="MORTALITY FACTOR 4 LIKE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10880">
        <property type="entry name" value="MORTALITY FACTOR 4-LIKE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05712">
        <property type="entry name" value="MRG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11717">
        <property type="entry name" value="Tudor-knot"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00298">
        <property type="entry name" value="CHROMO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54160">
        <property type="entry name" value="Chromo domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51640">
        <property type="entry name" value="MRG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0007">Acetylation</keyword>
    <keyword evidence="4" id="KW-0156">Chromatin regulator</keyword>
    <keyword evidence="3" id="KW-0227">DNA damage</keyword>
    <keyword evidence="8" id="KW-0233">DNA recombination</keyword>
    <keyword evidence="9" id="KW-0234">DNA repair</keyword>
    <keyword evidence="2" id="KW-0341">Growth regulation</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0804">Transcription</keyword>
    <keyword evidence="6" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="Chromo" evidence="13">
        <location>
            <begin position="57"/>
            <end position="124"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="123"/>
            <end position="145"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="175"/>
            <end position="212"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022604"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023172"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00071326"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="SMART" id="SM00298"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015553.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="390" mass="44651" checksum="C5E0F1D90C9226E0" modified="2019-05-08" version="1">MHFVSETSDLWTGGALKKASKLSYDPWRWHILAGGDESEWPCSMDQGGWFPAGISLLQKGERVLCFHGPLLYEAKCVKVAIKDKQVKYFIHYSGWNKNWDEWVPESRVLKYVDTNLQKQKELQKANQEQYAEGKMRGAAPGKKTSGLQQKNVEVFFRRDGAHTVCCLETPTISTRKTKKNKQKTPGIGEGSSTSETPQPPRKKRARVDPTVESEETFMNRVEVKVKIPEELKPWLVDDWDLITRQKQLFYLPAKKNVDSILEDYASYKKSRGNTDNKEYAVNEVVAGIKEYFNVMLGTQLLYKFERPQYAEILADHPDAPMSQVYGAPHLLRLFVRIGAMLAYTPLDEKSLALLLNYLHDFLKYLAKNASTLFSASDYEVAPPEYHRKAV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKU8</accession>
    <name>A0A452HKU8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Transforming acidic coiled-coil-containing protein C-terminal domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the TACC family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKU8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017759.1">
        <property type="protein sequence ID" value="ENSGAGP00000015546.1"/>
        <property type="gene ID" value="ENSGAGG00000011728.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0021987">
        <property type="term" value="P:cerebral cortex development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007052">
        <property type="term" value="P:mitotic spindle organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007097">
        <property type="term" value="P:nuclear migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.1700:FF:000001">
        <property type="entry name" value="Transforming acidic coiled-coil-containing protein 1 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.1700">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039915">
        <property type="entry name" value="TACC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007707">
        <property type="entry name" value="TACC_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13924">
        <property type="entry name" value="TRANSFORMING ACIDIC COILED-COIL CONTAINING PROTEIN 1/2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13924:SF12">
        <property type="entry name" value="TRANSFORMING ACIDIC COILED-COIL-CONTAINING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05010">
        <property type="entry name" value="TACC_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Transforming acidic coiled-coil-containing protein C-terminal" evidence="9">
        <location>
            <begin position="600"/>
            <end position="799"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1"/>
            <end position="46"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="103"/>
            <end position="157"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="185"/>
            <end position="231"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="256"/>
            <end position="276"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="304"/>
            <end position="385"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="500"/>
            <end position="525"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="548"/>
            <end position="572"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="615"/>
            <end position="688"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="717"/>
            <end position="801"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="104"/>
            <end position="114"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="133"/>
            <end position="144"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="8">
        <location>
            <begin position="197"/>
            <end position="210"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="213"/>
            <end position="224"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="256"/>
            <end position="274"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="8">
        <location>
            <begin position="305"/>
            <end position="321"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="364"/>
            <end position="374"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="550"/>
            <end position="563"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009423"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF05010"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015546.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="806" mass="88596" checksum="CD9D6D35E0F7FDE7" modified="2019-05-08" version="1">MGGAQSQRQREPPAAGQRPPGREETSSDSEGNFETPEAETPIRSPVKEICDPLLGLAVAEARTQELREDDEQLVAEISETYVPQKHSEDEVQSHLAVLQANKNSKAETDCKLDEDSSVTRVAPETEPPLDISSPFTKETDSPSSVVLAPDQTLPSNVPTATQEWVMSEEDMVVTLEGLADAESKAETLALESVPSKSKLKKVKSSSRGKKTAGETPETEAPTEGIPVPKASYQFSPAEYDEKINPFVMAGSKIQNSPSAAQQTFMASNSDTNDSGVELAEEPRGHALKLEFDFTDEVENGEVKKALPKKIGRKSASKLTPKRQREAANKPVSADGLEKAPAEVPLPKALYSVDPSQWDDPNFNPFGSSSNMQNSPPLPKASYHFDPNNFDTVDPFKPTKSLANTAADSCPTADNNLNEILESQTLEVQDEELTVGKASPKKAKSRLITTTEQVKFLCFLLSGCKVKKYEMQSLVLDVCTQDEGALVSKIPDVANCDGHATDEEKLASTSSSQKPSGPEVKGEPEDDLEYFECSNVPVSTINHAFSSSEAGFEKETSKQKEKDGPSSILGDCGLYSKDKAPGAMTTGASGSESPLESICLSESDKTAVLTLIREEIITKEIEANEWKKKYEESRQEVLEMRKIVAEYEKTIAQMIEDEQRTNMASQKSLQQLTMEKEQALADLNSVERSLSDLFRRYENLKGILEGFKKNEEALKKCAQDYLARVKQEEQRYQALKIHAEEKLDKANEEIAQVRTKAKAESAALHAGLRKEQMKVESLERALQQKNQEIEELTKICDELIAKMGKTD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKV4</accession>
    <name>A0A452HKV4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">BRCT domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Chromosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Cytoskeleton</location>
            <location evidence="3">Microtubule organizing center</location>
            <location evidence="3">Centrosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cytoplasm</location>
            <location evidence="4">Cytoskeleton</location>
            <location evidence="4">Spindle pole</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="14">Belongs to the TOPBP1 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000017788.1">
        <property type="protein sequence ID" value="ENSGAGP00000015572.1"/>
        <property type="gene ID" value="ENSGAGG00000011744.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005694">
        <property type="term" value="C:chromosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000922">
        <property type="term" value="C:spindle pole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006281">
        <property type="term" value="P:DNA repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006270">
        <property type="term" value="P:DNA replication initiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033314">
        <property type="term" value="P:mitotic DNA replication checkpoint signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007095">
        <property type="term" value="P:mitotic G2 DNA damage checkpoint signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd17737">
        <property type="entry name" value="BRCT_TopBP1_rpt1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd17731">
        <property type="entry name" value="BRCT_TopBP1_rpt2_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd17718">
        <property type="entry name" value="BRCT_TopBP1_rpt3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd17749">
        <property type="entry name" value="BRCT_TopBP1_rpt4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18434">
        <property type="entry name" value="BRCT_TopBP1_rpt5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd17727">
        <property type="entry name" value="BRCT_TopBP1_rpt6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd17738">
        <property type="entry name" value="BRCT_TopBP1_rpt7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd17728">
        <property type="entry name" value="BRCT_TopBP1_rpt8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10190:FF:000018">
        <property type="entry name" value="DNA topoisomerase 2-binding protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10190:FF:000028">
        <property type="entry name" value="DNA topoisomerase 2-binding protein 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10190:FF:000010">
        <property type="entry name" value="DNA topoisomerase II binding protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10190:FF:000020">
        <property type="entry name" value="DNA topoisomerase II binding protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10190:FF:000021">
        <property type="entry name" value="DNA topoisomerase II binding protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10190:FF:000022">
        <property type="entry name" value="DNA topoisomerase II binding protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10190:FF:000023">
        <property type="entry name" value="DNA topoisomerase II binding protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10190:FF:000029">
        <property type="entry name" value="DNA topoisomerase II binding protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10190:FF:000033">
        <property type="entry name" value="DNA topoisomerase II binding protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.10190">
        <property type="entry name" value="BRCT domain"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR059215">
        <property type="entry name" value="BRCT2_TopBP1-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001357">
        <property type="entry name" value="BRCT_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036420">
        <property type="entry name" value="BRCT_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057595">
        <property type="entry name" value="TopB1_SLF1_BRCT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049542">
        <property type="entry name" value="TopBP1-like_BRCT0"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044737">
        <property type="entry name" value="TopBP1_BRCT_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049936">
        <property type="entry name" value="TopBP1_BRCT_8"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13561">
        <property type="entry name" value="DNA REPLICATION REGULATOR DPB11-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13561:SF20">
        <property type="entry name" value="DNA TOPOISOMERASE 2-BINDING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00533">
        <property type="entry name" value="BRCT"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23294">
        <property type="entry name" value="BRCT_TopB1_SLF1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12738">
        <property type="entry name" value="PTCB-BRCT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21298">
        <property type="entry name" value="TopBP1_BRCT0"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00292">
        <property type="entry name" value="BRCT"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52113">
        <property type="entry name" value="BRCT domain"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50172">
        <property type="entry name" value="BRCT"/>
        <property type="match status" value="6"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0158">Chromosome</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="12" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="9" id="KW-0227">DNA damage</keyword>
    <keyword evidence="11" id="KW-0234">DNA repair</keyword>
    <keyword evidence="10" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="13" id="KW-0539">Nucleus</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="BRCT" evidence="16">
        <location>
            <begin position="100"/>
            <end position="171"/>
        </location>
    </feature>
    <feature type="domain" description="BRCT" evidence="16">
        <location>
            <begin position="193"/>
            <end position="282"/>
        </location>
    </feature>
    <feature type="domain" description="BRCT" evidence="16">
        <location>
            <begin position="353"/>
            <end position="443"/>
        </location>
    </feature>
    <feature type="domain" description="BRCT" evidence="16">
        <location>
            <begin position="631"/>
            <end position="728"/>
        </location>
    </feature>
    <feature type="domain" description="BRCT" evidence="16">
        <location>
            <begin position="836"/>
            <end position="927"/>
        </location>
    </feature>
    <feature type="domain" description="BRCT" evidence="16">
        <location>
            <begin position="1225"/>
            <end position="1290"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="285"/>
            <end position="307"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="1023"/>
            <end position="1052"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="15">
        <location>
            <begin position="290"/>
            <end position="299"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="15">
        <location>
            <begin position="1023"/>
            <end position="1051"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004647"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022454"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00061360"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50172"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015572.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1462" mass="163597" checksum="3FEB3AEFFBC539EA" modified="2019-05-08" version="1">MKSDKEPFFVKFIKSSGNSEYFFKALESIKEFQSEEYLQILEEVPALNIKENDKSLYICDPFSGIVFSHLKKLGCRIVGPQVVIFCMQHQRCVPRAEYPVHNMAMADITISCTSLEKETREEVHKYVQMMGGRVYRDLNASVTHLIAGEVGSKKYLVAAALKKPILLPTWVKALWEKSQEGMIRYTDISMEDFICPLFRGCTICVTGLSSVDRKEVQRLTTEHGGQYTGQLKMNECTHLIVQEPKGQKYECAKKWNVQCVSIQWFFNSIENGFCQDEAMYKTESVQKQNTTPSTSTPTSQASKPDSCTLSDVSHISNISLNGINETACSSAVNSRLDPPPDDLENLDISSFQAPDDLLDGCRIYLCGFSGRKLDKMRRLINCGGGVRFNQLNEDVTHVIVGEYNGELKQFLNKTTHRPHAVTAKWLLECFCKGYLLPVEQYIPLNFQPIENTILEQPGIKRTVPKSNSLTKKEGVKLVQHQNADEDDLLSQYVNNDSTLIEVEKLETGHFSDAAHITIQEEKRSSICHSSLTETSTVVEEGLFGRKRFLLLGFGEEDESCIVEIIKENSGKVLPPQNRAIADYAVVPLLGCKVESTVGDVVTNTWLIMCVEQQLLLNPQSNPLFTPVSVMEGSTPLEDCVLSFSQFTGAERDSLVFLAGLLGARVQEFFVRRANPKKGMFASTHLVVKEPDGSKYEAAKKWNLPAVTMAWLLESARTGKKADESKFPVENVNTGDKEESFVSQSNKTETTIMSSDIPDHPSNLLETGKKTAVMPLDVSRFQSKAFHSVISHHIGQKPTLPAEGRLLQKEPSLHLDTPSKFLSKDKLFKPSFDVKVEDSKPLADLVICVSKKLSKKQSELNAVAASLGADYRWCFDETVTHFIYQGRQNDSSKEYKSVKERGIHIVSEHWLLESAQEYKRLPESLFPHTYNPKMSLDISAVQDGRLSSSRLSSTGKTAKEAESISVDENDTADITTNQVKEAVTVREEQIAANEAKGVLTQTLEMRENFQKQLQEIMSATSIVKPQAQRGSLSRNGFDSSPTTPDGTRSVRNGRSRVLEALRQSWQAVMDINTEPSQNEQIVWDDPTAREERARLVSNLQWPNSPSQCSEQIQTDVNNMNDSPFRDSFVNREVAELAVHDSGDTGVAKIPKHPIYSDPETPVKDDQVIPTPQAPSIAFPLANPPVAPQPKEKAITVDEKVDEEPEKQCKFQLSSLNPQERIDYCHLIEELGGLVLEKQCFDPSCTHIIVGHPLRNEKYLASMAAGKWVLHRSYLEACRAAGCFVQEEDYEWGSNSILNVLSGINVNQKKLALAAMRWRKKIHKKRQETGIIEGAFSGWKVILNVDQTKEAGFKRLLQSGGAKVLPGHSLLLYKEATHLFADFSKLKPDDARVNVLEAAAQGVNCLKPEYIADYLIQELPPPMECYCLPEAVSYLQNSKELGIGLSQKRKAPGETSKVKRSRIH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKV7</accession>
    <name>A0A452HKV7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Transforming acidic coiled-coil-containing protein C-terminal domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the TACC family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKV7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017770.1">
        <property type="protein sequence ID" value="ENSGAGP00000015556.1"/>
        <property type="gene ID" value="ENSGAGG00000011728.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0021987">
        <property type="term" value="P:cerebral cortex development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007052">
        <property type="term" value="P:mitotic spindle organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007097">
        <property type="term" value="P:nuclear migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.1700:FF:000001">
        <property type="entry name" value="Transforming acidic coiled-coil-containing protein 1 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.1700">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039915">
        <property type="entry name" value="TACC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007707">
        <property type="entry name" value="TACC_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13924">
        <property type="entry name" value="TRANSFORMING ACIDIC COILED-COIL CONTAINING PROTEIN 1/2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13924:SF12">
        <property type="entry name" value="TRANSFORMING ACIDIC COILED-COIL-CONTAINING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05010">
        <property type="entry name" value="TACC_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Transforming acidic coiled-coil-containing protein C-terminal" evidence="9">
        <location>
            <begin position="616"/>
            <end position="815"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="22"/>
            <end position="74"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="129"/>
            <end position="190"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="213"/>
            <end position="259"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="284"/>
            <end position="304"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="332"/>
            <end position="416"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="512"/>
            <end position="543"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="561"/>
            <end position="610"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="631"/>
            <end position="704"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="733"/>
            <end position="817"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="47"/>
            <end position="60"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="132"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="161"/>
            <end position="172"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="180"/>
            <end position="190"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="8">
        <location>
            <begin position="225"/>
            <end position="238"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="241"/>
            <end position="252"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="284"/>
            <end position="302"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="8">
        <location>
            <begin position="333"/>
            <end position="349"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="392"/>
            <end position="402"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="566"/>
            <end position="579"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009423"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF05010"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015556.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="822" mass="90251" checksum="A8C9D3BD6B957E48" modified="2019-05-08" version="1">MAFSPWQILSPVQWAKWTWSAVRGGGAGDGEDGGVKQEIEEDDSQAETKSLSFSSDSEGNFETPEAETPIRSPVKEICDPLLGLAVAEARTQELREDDEQLVAEISETYVPQKHSEDEVQSHLAVLQANKNSKAETDCKLDEDSSVTRVAPETEPPLDISSPFTKETDSPSSVVLAPDQTLPSNVPTATQEWVMSEEDMVVTLEGLADAESKAETLALESVPSKSKLKKVKSSSRGKKTAGETPETEAPTEGIPVPKASYQFSPAEYDEKINPFVMAGSKIQNSPSAAQQTFMASNSDTNDSGVELAEEPRGHALKLEFDFTDEVENGEVKKALPKKIGRKSASKLTPKRQREAANKPVSADGLEKAPAEVPLPKALYSVDPSQWDDPNFNPFGSSSNMQNSPPLPKASYHFDPNNFDTVDPFKPTKSLANTAADSCPTADNNLNEILESQTLEVQDEELTVGKASPKKAKSRLITSGCKVKKYEMQSLVLDVCTQDEGALVSKIPDVANCDGHATDEEKLASTSSSQKPSGPEVKGEPEDDLEYFECSNVPVSTINHAFSSSEAGFEKETSKQKEKDGPSSILGDCGLYSKDKAPGAMTTGASGSESPLESICLSESDKTAVLTLIREEIITKEIEANEWKKKYEESRQEVLEMRKIVAEYEKTIAQMIEDEQRTNMASQKSLQQLTMEKEQALADLNSVERSLSDLFRRYENLKGILEGFKKNEEALKKCAQDYLARVKQEEQRYQALKIHAEEKLDKANEEIAQVRTKAKAESAALHAGLRKEQMKVESLERALQQKNQEIEELTKICDELIAKMGKTD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKV8</accession>
    <name>A0A452HKV8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Secretogranin-3</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="9">Secretogranin III</fullName>
        </alternativeName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasmic vesicle</location>
            <location evidence="1">Secretory vesicle membrane</location>
            <topology evidence="1">Peripheral membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Secreted</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKV8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017793.1">
        <property type="protein sequence ID" value="ENSGAGP00000015577.1"/>
        <property type="gene ID" value="ENSGAGG00000011750.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030667">
        <property type="term" value="C:secretory granule membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030658">
        <property type="term" value="C:transport vesicle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033366">
        <property type="term" value="P:protein localization to secretory granule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026197">
        <property type="entry name" value="SCG3"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR17388">
        <property type="entry name" value="SECRETOGRANIN III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR17388:SF2">
        <property type="entry name" value="SECRETOGRANIN-3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15467">
        <property type="entry name" value="SGIII"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0165">Cleavage on pair of basic residues</keyword>
    <keyword evidence="8" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="7" id="KW-0472">Membrane</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="6 11" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="11">
        <location>
            <begin position="1"/>
            <end position="21"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019405946" description="Secretogranin-3" evidence="11">
        <location>
            <begin position="22"/>
            <end position="471"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="82"/>
            <end position="119"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="126"/>
            <end position="145"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="208"/>
            <end position="238"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="354"/>
            <end position="406"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="84"/>
            <end position="96"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="133"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="360"/>
            <end position="394"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004268"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00013655"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022685"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00033446"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015577.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="471" mass="53654" checksum="2B6E59ED835C6D6F" modified="2019-05-08" version="1">MVFSWATLVQVLALIVSQVHGFPKPEGSRDKGIYNRELSEERPLEEQIAEAGAGKIRKTVSTENKPEFRNYSFPDDLNLLKSVTGKERNEKERESIRSSLYDHQQPAPDDADSTKNRRLVDDYDSTKSGLDYKFQDDPDGLHQLDGTPLTAEDIVQKIATRIYEENDRGVFDKIVSKLLNLGLITESQAYTLEDEVAEVLQQLIANEAKDREKESEDLDYPPSRTDSDTEEKQEEKMTPNVADQVGVMNGEIDDTLDNTWPSSSVLERRNELPSEDNFEDLQYFPNFYGLLKSLNSEEAKEKETLITIMKTLVDFVKMMVKYGTITPEEGVLYLENLDAMIALQTKSKLESPLTSTKNKLLADKSNEEADSTKEEAAKMEKQYETLKDSTKVKEQNTAGNNEEPKGKAEAYLEAIRKNIEWLKKHNKQGNKEDYDLSKLRDFIDQQADAYVDKGILDKEEADVIKRIYGSL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKW0</accession>
    <name>A0A452HKW0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Rho family-interacting cell polarization regulator 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Apical cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cell projection</location>
            <location evidence="4">Filopodium</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cell projection</location>
            <location evidence="3">Stereocilium membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the RIPOR family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKW0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017784.1">
        <property type="protein sequence ID" value="ENSGAGP00000015568.1"/>
        <property type="gene ID" value="ENSGAGG00000011582.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016324">
        <property type="term" value="C:apical plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030175">
        <property type="term" value="C:filopodium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060171">
        <property type="term" value="C:stereocilium membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006935">
        <property type="term" value="P:chemotaxis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007517">
        <property type="term" value="P:muscle organ development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009968">
        <property type="term" value="P:negative regulation of signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007605">
        <property type="term" value="P:sensory perception of sound"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000191">
        <property type="entry name" value="RHO family interacting cell polarization regulator 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031780">
        <property type="entry name" value="FAM65_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026136">
        <property type="entry name" value="RIPOR3"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15829">
        <property type="entry name" value="PROTEIN KINASE PKN/PRK1, EFFECTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15829:SF2">
        <property type="entry name" value="RHO FAMILY-INTERACTING CELL POLARIZATION REGULATOR 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15903">
        <property type="entry name" value="PL48"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="14" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="7" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="18" id="KW-0966">Cell projection</keyword>
    <keyword evidence="9" id="KW-0145">Chemotaxis</keyword>
    <keyword evidence="15" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="8" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="17" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="13" id="KW-0221">Differentiation</keyword>
    <keyword evidence="12" id="KW-1009">Hearing</keyword>
    <keyword evidence="16" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0517">Myogenesis</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="11" id="KW-0734">Signal transduction inhibitor</keyword>
    <feature type="domain" description="FAM65 N-terminal" evidence="20">
        <location>
            <begin position="16"/>
            <end position="346"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="39"/>
            <end position="73"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="364"/>
            <end position="465"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="609"/>
            <end position="645"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="19">
        <location>
            <begin position="48"/>
            <end position="59"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="60"/>
            <end position="69"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="19">
        <location>
            <begin position="414"/>
            <end position="424"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="425"/>
            <end position="454"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="631"/>
            <end position="645"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004289"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004486"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00005744"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00013627"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022500"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022541"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022700"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022740"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="Pfam" id="PF15903"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015568.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1003" mass="110874" checksum="AFB4EFDCB3C61375" modified="2019-05-08" version="1">MSVGSHSFSPGGPNGIIRSQSFAGFSGLQERRSRCNSFVENSSALKKPQAKVKKTHNLGHKNSTTSKEPQPQRMEEVYRALKNGLDEYLEVHQTELDKLTAQLKDMKRNSRVLCYLHRSRVTFSRYRRKRGNKGVDELYEAYCIQRRLCDGASKMKQAFAMSPASKAARESLSEINRSYKEYTEVCAVLCSPVLHCLAGFARLCPGDQYEIFMRYGRQRWKLKGKIEVNGKQSWDGEEMVFLPLIVGLISIKVTEVKGLATHTLVGSVTCETKDLFAARPQVVAVDINDLGTIKLNLEITWYPFDVEDLTPSTGNVNKASALQRRMSTYSQGTPETPTFKDYSFFSNSPDDIFENGMAGTEKRPISFSFGDIPNGDCASPPITVDSASALDPSSPEITVTPPEHKSSKNPAMHSSSTNSSLDSTPESKDLKSQPEHTPVHENKKSSRVASEVHQKPPSPGSDNLFIDSSIPVSLLQETDDVSELKPVELDTYEGNITKQLVKRLTSAEAPMTPERLQCEGSISGESEGYKSYLDGSIEDAFQGLLLALEPHKEQYKEFQDLDQQVMHLDDILKCKPAVNRSRSSSSISLTVESALESFDFLNTSDFDDEDGGGDEVCNGGGGADSVFSDTETEKTSYRSDHPEARGHLSEALTEDTGVGTSVAGSPLPLTTGNESLDITIVKHLQCCTQLIQQIVFSSKTPFVVRNLLDKLSKEIVVLENLAEISIENMGNIISVTEAIPEFHKKLSLLSFWTKCTGPTGVYHTSTDKMIKQLDVTFATAVNEECPGLAETVFRILLSQILDRTEPVLSSSLPSEILTVFQYYNYFASNGVSDLGSYLLQLAKEASVVQKLQTLKDEKLLQNIAELTSSSLPPQQEVLRSLALLLTENKNEIGEAVTLLLSASAKNRHFREKALIYYCEVLTQPNLHLQKAACMALKCLQATESIKMLVTLCQSDNEEIRKVASETLLSLGEDGRLAYEQLDKFPREFVKVGGRWGTEVATAF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKW2</accession>
    <name>A0A452HKW2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Rho family-interacting cell polarization regulator 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Apical cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cell projection</location>
            <location evidence="4">Filopodium</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cell projection</location>
            <location evidence="3">Stereocilium membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the RIPOR family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKW2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017790.1">
        <property type="protein sequence ID" value="ENSGAGP00000015574.1"/>
        <property type="gene ID" value="ENSGAGG00000011582.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016324">
        <property type="term" value="C:apical plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030175">
        <property type="term" value="C:filopodium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060171">
        <property type="term" value="C:stereocilium membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006935">
        <property type="term" value="P:chemotaxis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007517">
        <property type="term" value="P:muscle organ development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009968">
        <property type="term" value="P:negative regulation of signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007605">
        <property type="term" value="P:sensory perception of sound"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031780">
        <property type="entry name" value="FAM65_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026136">
        <property type="entry name" value="RIPOR3"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15829">
        <property type="entry name" value="PROTEIN KINASE PKN/PRK1, EFFECTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15829:SF2">
        <property type="entry name" value="RHO FAMILY-INTERACTING CELL POLARIZATION REGULATOR 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15903">
        <property type="entry name" value="PL48"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="14" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="7" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="18" id="KW-0966">Cell projection</keyword>
    <keyword evidence="9" id="KW-0145">Chemotaxis</keyword>
    <keyword evidence="15" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="8" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="17" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="13" id="KW-0221">Differentiation</keyword>
    <keyword evidence="12" id="KW-1009">Hearing</keyword>
    <keyword evidence="16" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0517">Myogenesis</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="11" id="KW-0734">Signal transduction inhibitor</keyword>
    <feature type="domain" description="FAM65 N-terminal" evidence="20">
        <location>
            <begin position="16"/>
            <end position="357"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="39"/>
            <end position="73"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="421"/>
            <end position="522"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="666"/>
            <end position="702"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="19">
        <location>
            <begin position="48"/>
            <end position="59"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="60"/>
            <end position="69"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="19">
        <location>
            <begin position="471"/>
            <end position="481"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="482"/>
            <end position="511"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="688"/>
            <end position="702"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004289"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004486"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00005744"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00013627"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022500"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022541"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022700"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022740"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="Pfam" id="PF15903"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015574.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1057" mass="117410" checksum="5D0984401C916F59" modified="2019-05-08" version="1">MSVGSHSFSPGGPNGIIRSQSFAGFSGLQERRSRCNSFVENSSALKKPQAKVKKTHNLGHKNSTTSKEPQPQRMEEVYRALKNGLDEYLEVHQTELDKLTAQLKDMKRNSRLVCQIKAIERYMRRLEFHISMVDELYEAYCIQRRLCDGASKMKQAFAMSPASKAARESLSEINRSYKEYTENMCTIEGELENLLGEFCIKMKGLAGFARLCPGDQYEIFMRYGRQRWKLKGKIEVNGKQSWDGEEMVFLPLIVGLISIKVTEVKGLATHTLVGSVTCETKDLFAARPQVVAVDINDLGTIKLNLEITWYPFDVEDLTPSTGNVNKASALQRRMSTYSQGTPETPTFKDYSFFKWLHPPQDRPQLSVLDALQDTFFDRLQRSRSFSDLPSLRLRPKAGLEQYSNSPDDIFENGMAGTEKRPISFSFGDIPNGDCASPPITVDSASALDPSSPEITVTPPEHKSSKNPAMHSSSTNSSLDSTPESKDLKSQPEHTPVHENKKSSRVASEVHQKPPSPGSDNLFIDSSIPVSLLQETDDVSELKPVELDTYEGNITKQLVKRLTSAEAPMTPERLQCEGSISGESEGYKSYLDGSIEDAFQGLLLALEPHKEQYKEFQDLDQQVMHLDDILKCKPAVNRSRSSSSISLTVESALESFDFLNTSDFDDEDGGGDEVCNGGGGADSVFSDTETEKTSYRSDHPEARGHLSEALTEDTGVGTSVAGSPLPLTTGNESLDITIVKHLQCCTQLIQQIVFSSKTPFVVRNLLDKLSKEIVVLENLAEISIENMGNIISVTEAIPEFHKKLSLLSFWTKCTGPTGVYHTSTDKMIKQLDVTFATAVNEECPGLAETVFRILLSQILDRTEPVLSSSLPSEILTVFQYYNYFASNGVSDLGSYLLQLAKEGRVLSTSEDLNIAELTSSSLPPQQEVLRSLALLLTENKNEIGEAVTLLLSASAKNRHFREKALIYYCEVLTQPNLHLQKAACMALKCLQATESIKMLVTLCQSDNEEIRKVASETLLSLGELDQLAAFVNSCQGDLRDWHQSMDQLLRDLDPTYIF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HL04</accession>
    <name>A0A452HL04_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Scaffolding anchor of CK1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the FAM83 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000017820.1">
        <property type="protein sequence ID" value="ENSGAGP00000015601.1"/>
        <property type="gene ID" value="ENSGAGG00000011761.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019901">
        <property type="term" value="F:protein kinase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.870.10:FF:000004">
        <property type="entry name" value="protein FAM83H isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.870.10">
        <property type="entry name" value="Endonuclease Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050944">
        <property type="entry name" value="FAM83"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012461">
        <property type="entry name" value="SACK1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16181">
        <property type="entry name" value="PROTEIN FAM83A-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16181:SF29">
        <property type="entry name" value="PROTEIN FAM83A-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07894">
        <property type="entry name" value="SACK1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56024">
        <property type="entry name" value="Phospholipase D/nuclease"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Scaffolding anchor of CK1" evidence="5">
        <location>
            <begin position="28"/>
            <end position="302"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="79"/>
            <end position="101"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="324"/>
            <end position="376"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="403"/>
            <end position="464"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="88"/>
            <end position="101"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="431"/>
            <end position="445"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006937"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="Pfam" id="PF07894"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015601.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="486" mass="53830" checksum="68E5BAAB6D767C31" modified="2019-05-08" version="1">HASQDLKALEHRLXSCGWEPPGKPIPLVLQHSETARLAVDAFLEQGEPGYLQAIAEEQELPFLSPLDMDYMSQHRSQSILEPNARKGHQTEPKEGDSEDRASLLSELSSGTYFPLLSDMQPPELELGWPVIPLATGYGQTQASVYFQRNKANSIKDLLRCLISRAKTVIAVVMDLFTDMEILCDLMEASSRRRVSVYLILDEKYLKYFIEMCSKMGLNKDHFPNMRVRCVSGDTYYSKGGKKFEGQVLEKFVLIDCDHVLAGTYSFTWLCSQVHTCLVTHFRGKIVEDFDREFQRLYVESKPVPGFCIPEAGTKPGLRCAPGGSKAGLGARGRSAVPRGFSSGEAAASSRLPSAGRGRSAQPPKPRPRPRCAATETRVGAALPARFRWLLAAWTXARAALRARRQQGRAGGPRAQRGSPRILQRRSSRVLPPAQRGAGPQRAASQAPPPAALCGHRDAGRRRSTRALPVVAGRLDRKLLRARRRCR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HL16</accession>
    <name>A0A452HL16_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Chemokine interleukin-8-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HL16"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017844.1">
        <property type="protein sequence ID" value="ENSGAGP00000015624.1"/>
        <property type="gene ID" value="ENSGAGG00000011773.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048020">
        <property type="term" value="F:CCR chemokine receptor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008009">
        <property type="term" value="F:chemokine activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061844">
        <property type="term" value="P:antimicrobial humoral immune response mediated by antimicrobial peptide"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070098">
        <property type="term" value="P:chemokine-mediated signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048245">
        <property type="term" value="P:eosinophil chemotaxis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006954">
        <property type="term" value="P:inflammatory response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030335">
        <property type="term" value="P:positive regulation of cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00272">
        <property type="entry name" value="Chemokine_CC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.50.40">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039809">
        <property type="entry name" value="Chemokine_b/g/d"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001811">
        <property type="entry name" value="Chemokine_IL8-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036048">
        <property type="entry name" value="Interleukin_8-like_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12015:SF103">
        <property type="entry name" value="C-C MOTIF CHEMOKINE 4-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12015">
        <property type="entry name" value="SMALL INDUCIBLE CYTOKINE A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00048">
        <property type="entry name" value="IL8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00199">
        <property type="entry name" value="SCY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54117">
        <property type="entry name" value="Interleukin 8-like chemokines"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0145">Chemotaxis</keyword>
    <keyword evidence="2" id="KW-0202">Cytokine</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 4" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="4">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019458703" description="Chemokine interleukin-8-like domain-containing protein" evidence="4">
        <location>
            <begin position="23"/>
            <end position="88"/>
        </location>
    </feature>
    <feature type="domain" description="Chemokine interleukin-8-like" evidence="5">
        <location>
            <begin position="30"/>
            <end position="86"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022500"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022514"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="SMART" id="SM00199"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015624.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="88" mass="9582" checksum="7423F49DAD54F3D8" modified="2019-05-08" version="1">MKVSVAALAVLLIAAFCSLASSAPIGSDPPTACCFTYTSRKIPRGLVVDYYDTNIQISDPETEIFSGREVCANPKEDWVQEYSFKRAG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HL45</accession>
    <name>A0A452HL45_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Receptor-type tyrosine-protein phosphatase O</fullName>
            <ecNumber evidence="2">3.1.3.48</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="17">Glomerular epithelial protein 1</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="18">Protein tyrosine phosphatase U2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Possesses tyrosine phosphatase activity. Plays a role in regulating the glomerular pressure/filtration rate relationship through an effect on podocyte structure and function.</text>
    </comment>
    <comment type="subunit">
        <text evidence="15">Interacts (phosphorylated form) with FYN and GRB2.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="13">Belongs to the protein-tyrosine phosphatase family. Receptor class 3 subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.48" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452HL45"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017884.1">
        <property type="protein sequence ID" value="ENSGAGP00000015656.1"/>
        <property type="gene ID" value="ENSGAGG00000011789.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098982">
        <property type="term" value="C:GABA-ergic synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098978">
        <property type="term" value="C:glutamatergic synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045296">
        <property type="term" value="F:cadherin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017147">
        <property type="term" value="F:Wnt-protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007411">
        <property type="term" value="P:axon guidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090090">
        <property type="term" value="P:negative regulation of canonical Wnt signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072112">
        <property type="term" value="P:podocyte differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003093">
        <property type="term" value="P:regulation of glomerular filtration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="CDD" id="cd14614">
        <property type="entry name" value="R-PTPc-O"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000488">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase O isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000039">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase O isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000415">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase O isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058859">
        <property type="entry name" value="Fn3_R-PTP-O"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000242">
        <property type="entry name" value="PTP_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042996">
        <property type="entry name" value="PTPRO"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016130">
        <property type="entry name" value="Tyr_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003595">
        <property type="entry name" value="Tyr_Pase_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47028">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE O"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47028:SF1">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE O"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26586">
        <property type="entry name" value="Fn3_R-PTP-O"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00102">
        <property type="entry name" value="Y_phosphatase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00700">
        <property type="entry name" value="PRTYPHPHTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00194">
        <property type="entry name" value="PTPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00404">
        <property type="entry name" value="PTPc_motif"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00383">
        <property type="entry name" value="TYR_PHOSPHATASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50055">
        <property type="entry name" value="TYR_PHOSPHATASE_PTP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="10 19" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="8" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="11" id="KW-0675">Receptor</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <keyword evidence="4 19" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9 19" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="807"/>
            <end position="831"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="867"/>
            <end position="888"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="315"/>
            <end position="411"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="421"/>
            <end position="518"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="618"/>
            <end position="711"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="712"/>
            <end position="804"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="20">
        <location>
            <begin position="925"/>
            <end position="1182"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="21">
        <location>
            <begin position="1099"/>
            <end position="1173"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00013064"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00025789"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00056314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00063538"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00073607"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00075446"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00080595"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015656.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1203" mass="136877" checksum="C957C23937EAD2F4" modified="2019-05-08" version="1">ESDYLFYKLTGIATYFTVRDDNCIVVSLEESDVVSSSFVYVVKITGESKNYFFQFEEFNSTLPSPVVFNASYHGLYYIVTLMVVNSNMVSKPVRSITVLTKPLPISSVSIYDYKPSPETGVLFEIQYPEKYNVFTRVNISYWEGKDYRTMLYKDFFKGKTVFNHWLPGICYSNITFQLVSEATFNKSTLVEYSGVRHEPKQHRTVPYPPRNISVQIVHMNKNNWEEHSGNFLEESFMGPQDVIGQDKLIHFSNGPTEIPSANTSDAWPDYHYNSTDYTTTTQPYWWTNEADSSESEDEFVNVVPSEYENSSTASMSGKLTPETPSFLPVQMVLSWLPPKPPTAFDGFHINIEREEKTTEFLTVGEDTHEFIAELKEPGKYKLSVTTFSSSGSCEIRRSQSAKTLSFYISPTGEWIEELTEKPQHVSVTVLSSTTALMSWTSSQDSSSNNTIVSVVSLTCQKQKESQRLEKHYCTEVNSSNSIIENLVPGAQYQVVIYLRKGPLVGPPSDPVTFAIVPTGIKDLMLYPLGPTAVILSWNRPYLGVFRKYVVEMFYFNPSTMTSEWMTYYEIAATVSLTASVKIANLLPAWYYNFRVTMVTWGDPELSCCDSSTISYITAPVSPEITSVEYFNNLLYVSWTYGDNGTDLSHSRMLHWMIIAEGKKKIKKSVTRNVMTTVLSLPPGDIYNLSVTACTERGSNASTPQIVKLEPAPPKSLFAVNKTQTSVTLLWVEEGVADFFEVFCQQAGSGQETKLQEPVTVSSHVVTISSLLPATSYNCSVITFSHNSPSVPTFIAVSTMVTEMNPNIVVISVLAILSILLIGLLLVTLVVLRRKHLQMARECGAGTFVNFASLERDGKLPYNWRRSVFAFLTLLPSCLWTDYLLAFYINPWSKNGLKKRKLTNPVQLDDFDGYIKDMAKDSDYKFSLQFEELKLIGLDIPHFAADLPMNRCKNRYTNILPYDFSRVRLVSMNEEEGSDYINANYIPGYNSPQEYIATQGPLPETRNDFWKMVLQQKSQIIVMLTQCNEKRRVKCDHYWPFTEEPVAYGDITVEMLSEEEQMDWSYRNFRISYADEVQDVMHFNYTAWPDHGVPTTNAAESILQFVQMVRQQSAKSKGPMVIHCSAGVGRTGTFIALDRLLQHIRDHEFVDILGLVSDMRSYRMSMVQTEEQYIFIHQCVQLMWLKKKQQFCISDVIYENVSKS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HL48</accession>
    <name>A0A452HL48_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Ubiquinol-cytochrome c chaperone domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the CBP3 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HL48"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017872.1">
        <property type="protein sequence ID" value="ENSGAGP00000015647.1"/>
        <property type="gene ID" value="ENSGAGG00000011771.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034551">
        <property type="term" value="P:mitochondrial respiratory chain complex III assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021150">
        <property type="entry name" value="Ubiq_cyt_c_chap"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007129">
        <property type="entry name" value="Ubiqinol_cyt_c_chaperone_CPB3"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12184">
        <property type="entry name" value="UBIQUINOL-CYTOCHROME C REDUCTASE COMPLEX ASSEMBLY FACTOR 1 FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12184:SF1">
        <property type="entry name" value="UBIQUINOL-CYTOCHROME-C REDUCTASE COMPLEX ASSEMBLY FACTOR 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03981">
        <property type="entry name" value="Ubiq_cyt_C_chap"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Ubiquinol-cytochrome c chaperone" evidence="2">
        <location>
            <begin position="45"/>
            <end position="180"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006407"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF03981"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015647.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="208" mass="24267" checksum="8147DD61DEC639D1" modified="2019-05-08" version="1">LPTPDQTQEKLTQTVGKITEKIKIAALRMYTCCVEKTDYEEFFHRCHMPDTLNSWFLIAQLHVWMCLVRMKQEGRAGKYMCRYIVHSMWEDVEQRGKVMGIDSFTLKQSMRSMTENFYAAIFGYDEGILSDDHVLAAALWRNLFNKHCEDPRQLELLVEYVRKQMQYLDAINGEDLLLTGEVNWRPLVESNAQSILKPTSPAYNDEGL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HL50</accession>
    <name>A0A452HL50_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Receptor-type tyrosine-protein phosphatase O</fullName>
            <ecNumber evidence="2">3.1.3.48</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="17">Glomerular epithelial protein 1</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="18">Protein tyrosine phosphatase U2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Possesses tyrosine phosphatase activity. Plays a role in regulating the glomerular pressure/filtration rate relationship through an effect on podocyte structure and function.</text>
    </comment>
    <comment type="subunit">
        <text evidence="15">Interacts (phosphorylated form) with FYN and GRB2.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="13">Belongs to the protein-tyrosine phosphatase family. Receptor class 3 subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.48" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452HL50"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017888.1">
        <property type="protein sequence ID" value="ENSGAGP00000015660.1"/>
        <property type="gene ID" value="ENSGAGG00000011789.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098982">
        <property type="term" value="C:GABA-ergic synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098978">
        <property type="term" value="C:glutamatergic synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045296">
        <property type="term" value="F:cadherin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017147">
        <property type="term" value="F:Wnt-protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007411">
        <property type="term" value="P:axon guidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090090">
        <property type="term" value="P:negative regulation of canonical Wnt signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072112">
        <property type="term" value="P:podocyte differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003093">
        <property type="term" value="P:regulation of glomerular filtration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="CDD" id="cd14614">
        <property type="entry name" value="R-PTPc-O"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000488">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase O isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000039">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase O isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000415">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase O isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058859">
        <property type="entry name" value="Fn3_R-PTP-O"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000242">
        <property type="entry name" value="PTP_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042996">
        <property type="entry name" value="PTPRO"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016130">
        <property type="entry name" value="Tyr_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003595">
        <property type="entry name" value="Tyr_Pase_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47028">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE O"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47028:SF1">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE O"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26586">
        <property type="entry name" value="Fn3_R-PTP-O"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00102">
        <property type="entry name" value="Y_phosphatase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00700">
        <property type="entry name" value="PRTYPHPHTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00194">
        <property type="entry name" value="PTPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00404">
        <property type="entry name" value="PTPc_motif"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00383">
        <property type="entry name" value="TYR_PHOSPHATASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50055">
        <property type="entry name" value="TYR_PHOSPHATASE_PTP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="10 19" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="8" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="11" id="KW-0675">Receptor</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <keyword evidence="4 19" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9 19" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="811"/>
            <end position="835"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="871"/>
            <end position="892"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="318"/>
            <end position="415"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="425"/>
            <end position="522"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="622"/>
            <end position="715"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="22">
        <location>
            <begin position="716"/>
            <end position="808"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="20">
        <location>
            <begin position="929"/>
            <end position="1186"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="21">
        <location>
            <begin position="1103"/>
            <end position="1177"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00013064"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00025789"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00056314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00063538"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00073607"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00075446"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00080595"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015660.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1207" mass="137143" checksum="F3CB6F63B3CBC23B" modified="2019-05-08" version="1">DGPLPSHRYRGSNLTPFQVTVRDDNCIVVSLEESDVVSSSFVYVVKITGESKNYFFQFEEFNSTLPSPVVFNASYHGLYYIVTLMVVNSNMVSKPVRSITVLTKPLPISSVSIYDYKPSPETGVLFEIQYPEKYNVFTRVNISYWEGKDYRTMLYKDFFKGKTVFNHWLPGICYSNITFQLVSEATFNKSTLVEYSGVRHEPKQHRTVPYPPRNISVQIVHMNKNNWEEHSGNFLEESFMGPQDVIGQDKLIHFSNGPTEIPSANTSDAWPDYHYNSTDYTTTTQPYWWTNEADSSESEDEFVNVVPSEYENSSTASMSGKLTPETPSFLPVQMVLSWLPPKPPTAFDGFHINIEREGKKTTEFLTVGEDTHEFIAELKEPGKYKLSVTTFSSSGSCEIRRSQSAKTLSFYISPTGEWIEELTEKPQHVSVTVLSSTTALMSWTSSQDSSSNNTIVSVVSLTCQKQKESQRLEKHYCTEVNSSNSIIENLVPGAQYQVVIYLRKGPLVGPPSDPVTFAIVPTGIKDLMLYPLGPTAVILSWNRPYLGVFRKYVVEMFYFNPSTMTSEWMTYYEIAATVSLTASVKIANLLPAWYYNFRVTMVTWGDPELSCCDSSTISYITAPVSPEITSVEYFNNLLYVSWTYGDNGTDLSHSRMLHWMIIAEGKKKIKKSVTRNVMTTVLSLPPGDIYNLSVTACTERGSNASTPQIVKLEPAPPKSLFAVNKTQTSVTLLWVEEGVADFFEVFCQQAGSGQETKLQEPVTVSSHVVTISSLLPATSYNCSVITFSHNSPSVPTFIAVSTMVTEMNPNIVVISVLAILSILLIGLLLVTLVVLRRKHLQMARECGAGTFVNFASLERDGKLPYNWRRSVFAFLTLLPSCLWTDYLLAFYINPWSKNGLKKRKLTNPVQLDDFDGYIKDMAKDSDYKFSLQFEELKLIGLDIPHFAADLPMNRCKNRYTNILPYDFSRVRLVSMNEEEGSDYINANYIPGYNSPQEYIATQGPLPETRNDFWKMVLQQKSQIIVMLTQCNEKRRVKCDHYWPFTEEPVAYGDITVEMLSEEEQMDWSYRNFRISYADEVQDVMHFNYTAWPDHGVPTTNAAESILQFVQMVRQQSAKSKGPMVIHCSAGVGRTGTFIALDRLLQHIRDHEFVDILGLVSDMRSYRMSMVQTEEQYIFIHQCVQLMWLKKKQQFCISDVIYENVSKS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HL68</accession>
    <name>A0A452HL68_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Kinesin-like protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5 6">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Kinesin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HL68"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017896.1">
        <property type="protein sequence ID" value="ENSGAGP00000015668.1"/>
        <property type="gene ID" value="ENSGAGG00000011818.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005874">
        <property type="term" value="C:microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008017">
        <property type="term" value="F:microtubule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003777">
        <property type="term" value="F:microtubule motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007018">
        <property type="term" value="P:microtubule-based movement"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056524">
        <property type="entry name" value="KIF6/9_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027640">
        <property type="entry name" value="Kinesin-like_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019821">
        <property type="entry name" value="Kinesin_motor_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001752">
        <property type="entry name" value="Kinesin_motor_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47968">
        <property type="entry name" value="CENTROMERE PROTEIN E"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47968:SF67">
        <property type="entry name" value="KINESIN MOTOR DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23735">
        <property type="entry name" value="KIF9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00225">
        <property type="entry name" value="Kinesin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00380">
        <property type="entry name" value="KINESINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00129">
        <property type="entry name" value="KISc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00411">
        <property type="entry name" value="KINESIN_MOTOR_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50067">
        <property type="entry name" value="KINESIN_MOTOR_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 5 6" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="6" id="KW-0493">Microtubule</keyword>
    <keyword evidence="5 6" id="KW-0505">Motor protein</keyword>
    <keyword evidence="2 5 6" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Kinesin motor" evidence="7">
        <location>
            <begin position="1"/>
            <end position="306"/>
        </location>
    </feature>
    <feature type="binding site" evidence="5">
        <location>
            <begin position="58"/>
            <end position="65"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00283"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="RuleBase" id="RU000394"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50067"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015668.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="663" mass="74790" checksum="54423AF98CDB4110" modified="2019-05-08" version="1">VPRDLADGFVNNKRESYKFKFQKIFDQEVKQDAVFESIAKPVAECALAGYNGTIFAYGQTGSGKTFTITGGAERYSDRGIIPRTLSYIFQQLEKDSSKVYTTHVSYLEIYNECGYDLLDPRHEASRLEDLPKVTIMEDPDQNIHLKNLSLQQATNEEEALNLLFLGDTNRMIAETPMNQASTRSHCIFTVHISSKEPGSATIRRSKLHLVDLAGSERVAKTGVGGQLLTEAKYINLSLHYLEQVIIALAEKHRSHIPYRNSVMTSVLRDSLGGNCMTTMIATLSLDKRNIDESISTCRFAQRVALIKNEAVLNEEVDPRLMIIQLKKVIQELKDELAMVTGVQRTEELTQEELLQLDEIIKTFLEDPDPESTLDVGADMRKIKHCFNYVKQLINHTNIPDRSLLSPDITGNEDTNTSEKKEELKKLRDLLHQRDNEINILVSMLKKEKKKAQDALFQLSAGSSGLAVSESSLSISLEEGQRPSACVNVKDTQDFSSSVHRSAFLPGKTGSGGEMSLGRQEAFEIFKRDYVDSITIEDNKQLLKKSYKTMFNRLKGLKIEIEHLQLLMEKAKVKLQKDFEVWWSEEASNLQPLNLPFFQTEKKGNGNREHGIKMFTHLNIPAYFRTPQSSSSSIPLTGDSQTDADILAFIKARQNLLQKKGKPC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HL69</accession>
    <name>A0A452HL69_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">Epidermal growth factor receptor kinase substrate 8</fullName>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell projection</location>
            <location evidence="2">Growth cone</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cell projection</location>
            <location evidence="3">Ruffle membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cell projection</location>
            <location evidence="4">Stereocilium</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cell cortex</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="16">Synapse</location>
            <location evidence="16">Synaptosome</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the EPS8 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HL69"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017909.1">
        <property type="protein sequence ID" value="ENSGAGP00000015680.1"/>
        <property type="gene ID" value="ENSGAGG00000011825.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000017912.1">
        <property type="protein sequence ID" value="ENSGAGP00000015683.1"/>
        <property type="gene ID" value="ENSGAGG00000011825.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005938">
        <property type="term" value="C:cell cortex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098978">
        <property type="term" value="C:glutamatergic synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030426">
        <property type="term" value="C:growth cone"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0014069">
        <property type="term" value="C:postsynaptic density"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032587">
        <property type="term" value="C:ruffle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032420">
        <property type="term" value="C:stereocilium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031982">
        <property type="term" value="C:vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003779">
        <property type="term" value="F:actin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:1900029">
        <property type="term" value="P:positive regulation of ruffle assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035023">
        <property type="term" value="P:regulation of Rho protein signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007266">
        <property type="term" value="P:Rho protein signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd01210">
        <property type="entry name" value="PTB_EPS8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd09540">
        <property type="entry name" value="SAM_EPS8-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd11764">
        <property type="entry name" value="SH3_Eps8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.50:FF:000023">
        <property type="entry name" value="Epidermal growth factor receptor kinase substrate 8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000174">
        <property type="entry name" value="epidermal growth factor receptor kinase substrate 8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.40:FF:000071">
        <property type="entry name" value="Epidermal growth factor receptor kinase substrate 8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.50">
        <property type="entry name" value="Transcription Factor, Ets-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039801">
        <property type="entry name" value="EPS8-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055093">
        <property type="entry name" value="EPS8_2nd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033928">
        <property type="entry name" value="EPS8_PTB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035462">
        <property type="entry name" value="Eps8_SH3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013625">
        <property type="entry name" value="PTB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006020">
        <property type="entry name" value="PTB/PI_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013761">
        <property type="entry name" value="SAM/pointed_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041418">
        <property type="entry name" value="SAM_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12287:SF21">
        <property type="entry name" value="EPIDERMAL GROWTH FACTOR RECEPTOR KINASE SUBSTRATE 8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12287">
        <property type="entry name" value="EPIDERMAL GROWTH FACTOR RECEPTOR KINASE SUBSTRATE EPS8-RELATED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22975">
        <property type="entry name" value="EPS8_2nd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08416">
        <property type="entry name" value="PTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18016">
        <property type="entry name" value="SAM_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00018">
        <property type="entry name" value="SH3_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00462">
        <property type="entry name" value="PTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00326">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="14" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="7" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="15" id="KW-0966">Cell projection</keyword>
    <keyword evidence="8" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="13" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6 18" id="KW-0728">SH3 domain</keyword>
    <keyword evidence="12" id="KW-0770">Synapse</keyword>
    <keyword evidence="10" id="KW-0771">Synaptosome</keyword>
    <keyword evidence="11" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="SH3" evidence="20">
        <location>
            <begin position="499"/>
            <end position="558"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="204"/>
            <end position="224"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="299"/>
            <end position="320"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="584"/>
            <end position="656"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="19">
        <location>
            <begin position="209"/>
            <end position="221"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="19">
        <location>
            <begin position="299"/>
            <end position="309"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="19">
        <location>
            <begin position="587"/>
            <end position="620"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="19">
        <location>
            <begin position="621"/>
            <end position="638"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="642"/>
            <end position="656"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004544"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004624"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004632"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004645"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00006197"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022599"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00034102"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00068628"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015680.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="793" mass="88370" checksum="AB0C958296237B34" modified="2019-05-08" version="1">MNGHISNTSPSVYGNYSSQINGYGSPTFAQAEREQSSKASAKAIYEQRKSYARDSASSISETSQYHVEHLTTFVLDRKEAMITVDDGIRKLKLLDAKGKVWTQDMILQVDDKAVSLVDLESKNELENFPLSTIQHCQAVMNTCNYDSILALVCKEPAQNKPDLHLFQCDEVKASFIHEDIESAISDSKGGKQKKRLDTLRMISKADSSIPPPPKAPAPVPPGTVTQVDVRSRVAAWSAWAAEQGDFEKPRQFHEHEETAEMMAARIDRDVQILNHILDDIEYFVTKLQKAAEAFAELSKRKKTKKNKKKGPGEGVLTLRAKPPPPDEFVDCFQKFKHGFNLLAKLKSHIQNPSAPELVHFLFTPLNMVVQATGGPELAGTVLSPLLTKDTIDFLRYTISNEEGQLWMSLGDTWTKARVDWPKDQFIPPYIPRFRNGWEPPLLSFMGAPNEQELNQLAESVANVAEHQRKQEMKRMSAEHPSVPDYPPSDGNYEAHSKAQPKRYAKSKYDFVARNHSELSVLKDDILEILDDRKQWWKVQNKNGLAGFVPNNILDPLRPQEPGLGHSEPAYSHTIQKQRMDYVLKQPDPSPAAPSPPPTPAPVPVPLPPIAPAPIPVPVPKAPGTISRQSSTSSESGGTIMRDSQRHKQLPVDRRKSQMEEVQDELIHRLTIGRSAAQKKFHVPRPNIPVVNITYDSSAEDVKMWLESKGFSPVTVNSLGVLTGAQLFSLNKDELKTVCPEGSRVYSQITVQKAALEANNGSSELQEIMRRRQEKISAAASDSGVESFDEGSSH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HL77</accession>
    <name>A0A452HL77_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">CMP/dCMP-type deaminase domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the cytidine and deoxycytidylate deaminase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HL77"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017930.1">
        <property type="protein sequence ID" value="ENSGAGP00000015701.1"/>
        <property type="gene ID" value="ENSGAGG00000011837.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004126">
        <property type="term" value="F:cytidine deaminase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016554">
        <property type="term" value="P:cytidine to uridine editing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd01283">
        <property type="entry name" value="cytidine_deaminase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.140.10">
        <property type="entry name" value="Cytidine Deaminase, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050610">
        <property type="entry name" value="APOBEC_Cyt_Deaminase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002125">
        <property type="entry name" value="CMP_dCMP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016193">
        <property type="entry name" value="Cytidine_deaminase-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13857:SF26">
        <property type="entry name" value="C-U-EDITING ENZYME APOBEC-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13857">
        <property type="entry name" value="MRNA EDITING ENZYME"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18769">
        <property type="entry name" value="APOBEC1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53927">
        <property type="entry name" value="Cytidine deaminase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51747">
        <property type="entry name" value="CYT_DCMP_DEAMINASES_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="CMP/dCMP-type deaminase" evidence="5">
        <location>
            <begin position="13"/>
            <end position="121"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006576"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS51747"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015701.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="187" mass="22051" checksum="5EF23B06A3A2A877" modified="2019-05-08" version="1">IFILSAIAFKVPMNQTSCIRAGWQLHSGSSGVSTKFWRKWCLNTPTQHAEINCLENDFKAIHHGPSVHCSIIWFLSWSPCGLCCRLIIHFLRAHPKVTLQIYIGRLFRHRDERNRQGLRDLVSSGVNVYIMSLPAYDYCWRTFVDHRVREDDIYWPCYFTPWMIFYMLEFQCILQVGRCLMLGWGEN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HL80</accession>
    <name>A0A452HL80_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">cyclin-dependent kinase</fullName>
            <ecNumber evidence="2">2.7.11.22</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="10">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.22"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="9">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.22"/>
        </reaction>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the protein kinase superfamily. CMGC Ser/Thr protein kinase family. CDC2/CDKX subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.22" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452HL80"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017920.1">
        <property type="protein sequence ID" value="ENSGAGP00000015691.1"/>
        <property type="gene ID" value="ENSGAGG00000011830.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004693">
        <property type="term" value="F:cyclin-dependent protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000007">
        <property type="entry name" value="Cyclin-dependent kinase 14, putative"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000061">
        <property type="entry name" value="Putative cyclin-dependent kinase 17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050108">
        <property type="entry name" value="CDK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24056">
        <property type="entry name" value="CELL DIVISION PROTEIN KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24056:SF52">
        <property type="entry name" value="CYCLIN-DEPENDENT KINASE 18"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 11 12" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="13" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-0418">Kinase</keyword>
    <keyword evidence="6 11 12" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 12" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="5" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Protein kinase" evidence="15">
        <location>
            <begin position="143"/>
            <end position="403"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="57"/>
            <end position="76"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="13">
        <location>
            <begin position="18"/>
            <end position="45"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="62"/>
            <end position="75"/>
        </location>
    </feature>
    <feature type="binding site" evidence="11">
        <location>
            <position position="172"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006485"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00012425"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00047811"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00048367"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="RuleBase" id="RU000304"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015691.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="452" mass="51896" checksum="448EE8A19192E24B" modified="2019-05-08" version="1">MNKMKNFKRRFSLSVPRTETIEESLAEFTEQFNQLNNRKNEDLAQGHLQMGHLGRGFRAEPTSISPSEVEGQSPTAVRYRRNNQRRFSMEDVSKRLSLPVDIRLPPEFLQKLQMDNPEFPKTFSRMSRRASLSDIGFGKLETYVKLEKLGEGTYATVFKGRSKLTGNLVALKEIRLEHEEGAPCTAIREVSLLKNLKHANIVTLHDFIHTEHSLTLVFEYLIFLFQLLRGLSYCHKSKILHRDLKPQNLLINERGELKLADFGLARAKSIPTKTYSNEVVTLWYRPPDVLLGSTEYSTPIDMWGVGCIHYEMVTGRPMFPGSTVKEELHLIFRLLGTPTEDSWPGITSNEEFKGYHFTQYQAQPLINHAPRLDTEGIDLLTSLLLYEAKHRISAEEALRHRYFRALGDRVQLLPDNASVFSLKEIQLQKDPGYRGSAFQQSARGKGRRQSIF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HL84</accession>
    <name>A0A452HL84_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Alpha-2-macroglobulin like 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the protease inhibitor I39 (alpha-2-macroglobulin) family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000017940.1">
        <property type="protein sequence ID" value="ENSGAGP00000015711.1"/>
        <property type="gene ID" value="ENSGAGG00000011842.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004867">
        <property type="term" value="F:serine-type endopeptidase inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd02897">
        <property type="entry name" value="A2M_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.50.10.20:FF:000001">
        <property type="entry name" value="CD109 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1930:FF:000001">
        <property type="entry name" value="CD109 isoform 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.50.10.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.20.130.20">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.1540">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1930">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1940">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.690">
        <property type="entry name" value="Alpha-macroglobulin, receptor-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009048">
        <property type="entry name" value="A-macroglobulin_rcpt-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036595">
        <property type="entry name" value="A-macroglobulin_rcpt-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050473">
        <property type="entry name" value="A2M/Complement_sys"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011625">
        <property type="entry name" value="A2M_N_BRD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041813">
        <property type="entry name" value="A2M_TED"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047565">
        <property type="entry name" value="Alpha-macroglob_thiol-ester_cl"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011626">
        <property type="entry name" value="Alpha-macroglobulin_TED"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014756">
        <property type="entry name" value="Ig_E-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001599">
        <property type="entry name" value="Macroglobln_a2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019742">
        <property type="entry name" value="MacrogloblnA2_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002890">
        <property type="entry name" value="MG2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041555">
        <property type="entry name" value="MG3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040839">
        <property type="entry name" value="MG4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008930">
        <property type="entry name" value="Terpenoid_cyclase/PrenylTrfase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11412:SF182">
        <property type="entry name" value="ALPHA-2-MACROGLOBULIN-LIKE PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11412">
        <property type="entry name" value="MACROGLOBULIN / COMPLEMENT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00207">
        <property type="entry name" value="A2M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07703">
        <property type="entry name" value="A2M_BRD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07677">
        <property type="entry name" value="A2M_recep"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01835">
        <property type="entry name" value="MG2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17791">
        <property type="entry name" value="MG3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17789">
        <property type="entry name" value="MG4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07678">
        <property type="entry name" value="TED_complement"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01360">
        <property type="entry name" value="A2M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01359">
        <property type="entry name" value="A2M_N_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01361">
        <property type="entry name" value="A2M_recep"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01419">
        <property type="entry name" value="Thiol-ester_cl"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49410">
        <property type="entry name" value="Alpha-macroglobulin receptor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81296">
        <property type="entry name" value="E set domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48239">
        <property type="entry name" value="Terpenoid cyclases/Protein prenyltransferases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00477">
        <property type="entry name" value="ALPHA_2_MACROGLOBULIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="8" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="4" id="KW-0646">Protease inhibitor</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0964">Secreted</keyword>
    <keyword evidence="6" id="KW-0722">Serine protease inhibitor</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <feature type="domain" description="Alpha-2-macroglobulin bait region" evidence="9">
        <location>
            <begin position="412"/>
            <end position="562"/>
        </location>
    </feature>
    <feature type="domain" description="Alpha-2-macroglobulin" evidence="10">
        <location>
            <begin position="705"/>
            <end position="795"/>
        </location>
    </feature>
    <feature type="domain" description="Alpha-macroglobulin receptor-binding" evidence="11">
        <location>
            <begin position="1336"/>
            <end position="1423"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010952"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022690"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="SMART" id="SM01359"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="SMART" id="SM01360"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="SMART" id="SM01361"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015711.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="14">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1428" mass="159385" checksum="E9C215618040C0B5" modified="2019-05-08" version="1">MDLSNVKSSMLLTITLETKIKTYKLLDRFVWERRQLKCFSFLVPPPSGGTEEVTTIQLSVRGGKSKITEKKQVLIQRANNSTFVQTDKPLYMPGQLVRFRIVTLNNNFIPLNDKYPLVELKDPENNRIGQWLNVKPARGIVQLSFQLAAEPPLGTYTINVDNGKAYSIFSVEEYVLPKFKVVVTEPTQIFTSDTSFPLKVCGKYTYGKPVQGKVQVSLCQTAYTWYIDRTESEPMDICQNVTGQTDKTGCFSVSVDLAFFSLGSPGYSHSINVVATLVEAGTGVEANATHVAFMSEQAGSMSFEDSDNYYHPGYPYSGKIKALDREGSILKKQPVFLVINAGNTQSYHNLTTDSTGVATFTLDTAVWNRSEVSLEGRFRLPDLVDEPTRIRQYYLNAFLHLKPYYSTTKSYLKIQRLQGVLPCGQKQVVQVDYRIDRMDLGDKAREILFSFYVVGKQKLIEDGQKRVDIKKNGELMGSFSIPLTFTFHFAPDPSLVVYAMFPDGGVVADKVQFTVSMCFKNQVNLEFSQQQDLPGATVNLQLRAAPNSLCAIRAVDQSILLLRPEGELSNSSVYGMLDFSNGGYPYQVDEYDDCYISRPFPMPVIDPLPVDRSRRSIILRPWFDQGADFFSFLKEMGLKILSNGKIKRPQDCRLYPIWYYKENILMKAPMPSIMEDSGRVPAVESQPSSVVKSSESQVREFFPETWIWELYPIESSGKKQVPVTLPDAITEWKASMFCTSESDGFGISPTVGLMAFKPFFVDLTLPYSVIRGESFTLTATVFNYLKQCIRIQITLAKSEKYQVKSCKGCEYTSCLCADEARTYYWNVTATKLGEVNITITTVALPSSELCGGRNVTVLNKGRSDTLIKPLLVRPEGVLMEKSHSSLPCPEGNSASDSVSLQLPEDVVLDSARAQVSVLGDIMGTALQNLDRLVQMPSGCGEQNMVLFAPIIYVLQYLEKTGQLTPEIRXEELKTRAVGFLQTGYQRQLQYKHRDGSYSAFGESDGEGNTWLTAFVTKCFIQAKQHIFIDDKNIEDALKWIAENQQANGCYANVGKLFHTAMKGGINDDISLAAYITAALLEAGKPLTDPMVKNSLQCLRNASSNATDIYTQALLAYVYSLAGDTVTRQLLLTKLDRQAIKSGGQIHWSRKPAQPPTEYSWSQPKSMDVELTAYAALALLTTEKVSKEEIASATGIVAWLARQQNAYGGFASTQDTVVALQALARYGAVTFIKSGNVSVTVKSGKNFQQSFHVEATNRLVLQQQPLPDIPGNYTIQAAGTGCVYVQTVLRYNVPPSKNAGTFTLSVRAGEDECEKQTPPHFLSLNISVSYVGSRGTSNMVIIEVKILSGFYPVRGTNYFLQQQPFVKRVEIRTEDLTIYLDQLNKETQTYSFTMSQDILVKDLKPAVVKVYDYYQPDEQATVEYTDPCQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INR3</accession>
    <name>A0A452INR3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Cytoplasmic FMR1 interacting protein 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="1 2">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="2" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="1" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="1" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452INR3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033516.1">
        <property type="protein sequence ID" value="ENSGAGP00000029516.1"/>
        <property type="gene ID" value="ENSGAGG00000020525.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031267">
        <property type="term" value="F:small GTPase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030833">
        <property type="term" value="P:regulation of actin filament polymerization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008081">
        <property type="entry name" value="Cytoplasmic_FMR1-int"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12195">
        <property type="entry name" value="CYTOPLASMIC FMR1-INTERACTING PROTEIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05994">
        <property type="entry name" value="FragX_IP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF008153">
        <property type="entry name" value="FMR1_interacting"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01698">
        <property type="entry name" value="CYTOFMRPINTP"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000313" key="1">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029516.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="3">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1004" mass="116305" checksum="90113C9F08BD2F3D" modified="2019-05-08" version="1">MTAQVTLEDALSNVDLLEELPLPDQQPCIEPPPSSLLYQPNFNTNFEDRNAFVTGIARYIEQATVHSNMNEMLEDGQEYAVMLYTWRSCSRAIPQVKCNEQPNRVEIYEKTVEVLEPEVTKLMNFMYFQVMGFGLYLMDGSVSNIYKLDAKKRINLTKIDKFFKQLQVVPLFGDMQIELARYIKTSAHYEENKSRWTCTSSSSSPQYNICEQMIQIREDHMRFISELARYSNSEVVTGSGRQEAQKTDAEYRKLFDLSLQGLQLLSQWSAHVMEVYSWKLVHPTDKYSNKDCPDNAEEYERATRYNYTSEEKFALVEVIAMIKGLQVLMGRMESVFNHAIRHTIYAALQDFSQVTLREPLRQAIKKKKNVIQSVLQAIRKTVCDWEAGHEPFNDPALRGEKDPKSGFDIKVPRYVLYSLDLYNDSAHYALTKFKKQFLYDEIEAEVNLCFDQFVYKLADQIFAYYKIMAGSLLLDKRLRSECKNQGATIHLPPSNRYETLLKQRHVQLLGRSIDLNRLITQRISAAMYKSMELAIGRFESEDLTSIVELDGLVEINKMTHKLLSRYMTLDSFDAMFREANHNVSAPYGRITLHVFWELNYDFLPNYCYNGSTNRFVRTVLPFSQEFQRDKQPNAQPQYLHGSKALNLTYSSIYSNYRNFVGPPHFKVICRLLGYQGIAVVMEELLKVVKSLLQGTILQYVKTLMEVMPKICRLPRHEYGSPGILEFFHHQLKDIVEYAELKTVCFQNLREVGNAVLFCLLIEQSLSLEEVCDLLHAAPFQNILPRVHVKEGERLDAKMKRLESKYAPLHLVPLIERLGTPQQIAIAREGDLLTKERLCCGLSMFEVILTRIRTFLDDPIWRGPLPSNGVMHVDECVEFHRLWSAMQFVYCIPVGTHEFTVEQCFGDGLHWAGCMIIVLLGQQRRFDVLDFCYHLLKVQKHDGKDEVIKNVPLKKMVERIRKFQILNNEIITILDKYMKSGDGEGTPVEHVRCFQPPIHQSLASN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INR5</accession>
    <name>A0A452INR5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">Ras-GAP domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452INR5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033620.1">
        <property type="protein sequence ID" value="ENSGAGP00000029607.1"/>
        <property type="gene ID" value="ENSGAGG00000019812.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005938">
        <property type="term" value="C:cell cortex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903479">
        <property type="term" value="P:mitotic actomyosin contractile ring assembly actin filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05131">
        <property type="entry name" value="RasGAP_IQGAP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00201">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.506.10:FF:000004">
        <property type="entry name" value="IQ motif containing GTPase activating protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.506.10">
        <property type="entry name" value="GTPase Activation - p120gap, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000048">
        <property type="entry name" value="IQ_motif_EF-hand-BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000593">
        <property type="entry name" value="RasGAP_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023152">
        <property type="entry name" value="RasGAP_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001936">
        <property type="entry name" value="RasGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001202">
        <property type="entry name" value="WW_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149">
        <property type="entry name" value="RAS GTPASE-ACTIVATING PROTEIN WITH IQ MOTIF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149:SF12">
        <property type="entry name" value="RAS GTPASE-ACTIVATING-LIKE PROTEIN IQGAP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00612">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00616">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03836">
        <property type="entry name" value="RasGAP_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00015">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00323">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00456">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF143885">
        <property type="entry name" value="RGC domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00509">
        <property type="entry name" value="RAS_GTPASE_ACTIV_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50018">
        <property type="entry name" value="RAS_GTPASE_ACTIV_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01159">
        <property type="entry name" value="WW_DOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Ras-GAP" evidence="1">
        <location>
            <begin position="615"/>
            <end position="843"/>
        </location>
    </feature>
    <evidence type="ECO:0000259" key="1">
        <source>
            <dbReference type="PROSITE" id="PS50018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029607.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1262" mass="144168" checksum="7FF18546723C2355" modified="2019-05-08" version="1">SVSVLHGKNPINFLYFPPELVAVDHINTAIRNCDANKTLVALMKPEAQLPVVHPFAAAVYQTELFNLQQQNRYLAHDELSIAVEMLSAVVLLNQALGSKDILAIKNHLRNPSVGFNNVEDENFQRYADTLLSIKSEASSQGQDYLSWNDVQNCIDMVNMQVQEENDRIVAVSYINEAIDQGNPGKTLETLLLPTVKLHDVNPTNARHYQDVLHDAKVQKCKESHDESAVLWLDEIQKGINDANSNIEKAANLALGTSMINNCLEKHDSQPVLDILQSPKFGLRVVPECAETYYKNLLEAKNLKTKEDSSEGPWLKLIMKNRYDYYYNVETGESTFVPPEGFIPKTSWLTSEEIQTIVGQVTADYNREQLWFANENLILQLQALARGFLVRKSYEERKGFLQKQEPSVIKIQVLLCNIKAQSLLLEHDLLFEVSCSMLSKVFSLWSFFQNEEIVKIQAFLRANKARDDYRTLIGAENPPLNVLRKFAYLLDQSDLDFQEELEVTRLREEVVTKIRSNQQLEKDLNLMDIKIGLLVKNRISLQDVVLHSKKLNKKSKNQVEEMVTGDKQGIKGLSKERRKQLEAYQHLFYLLQTNPMYLAKLIFQMPQNKSTKFMDTVIFTLYNYASNQREEYLLLKLFKTALEEEINSKVDQIQDIVTGNPTVIKMVVSFNRGARGQNSLRQLLAPVVKEIMDDKSLLINTSPVEVYKAWVNQLETQTGEASKLPYDVTTEQALEHPEVVNRLESSIQSLRTVTDKVLVSIFSSYGMRYIAKVLKNSLHEKFPDATEDELLKIVGNLLYYRYMNPAIVAPDGFDIIDMTAGGQIHSDQRRNLGCVAKVLQHAASNKLFEGESAHLSSMNTYLSQTYQKFRNFFQIACDVPEPEEKFNIDEYSDMVTLSKPVIYISIEEIINTHSLLLEHQDAIAPEQNDLLNELLKGLGAVPDVESFLGEGAVDPNDPNKENTLSQLAKTEISLSLTSKYELQEGEDQDLKGTKRLVVDVIRTQPGDTLLEILEIPASAQQESEHLKVVEKRAILDSKTPEKMKHSQSMLEDGQLPIEQKKRKIQRNLRTLEQAGLVSSANKYQEIINEIAKDIRNQRRYRHHRKAELVKLQQTLSALQSKTAFYEDQVNYYNTYIKTCLDNLTRKSIKLDGKVEEKGSKKLKHTSLKYTAARLHEKGVILEIEDLQTNQFKNVMFDITPGEEMGDFEIKAKFLGVQMEKVELHFQDLLQMQYEGVAVMKMFDKAKVNVNLLIFLLNKKFYGK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INR6</accession>
    <name>A0A452INR6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Interferon regulatory factor</fullName>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1 11">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="11">Belongs to the IRF family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452INR6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033767.1">
        <property type="protein sequence ID" value="ENSGAGP00000029738.1"/>
        <property type="gene ID" value="ENSGAGG00000021467.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002376">
        <property type="term" value="P:immune system process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00103">
        <property type="entry name" value="IRF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.10:FF:000065">
        <property type="entry name" value="Interferon regulatory factor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.10">
        <property type="entry name" value="Winged helix-like DNA-binding domain superfamily/Winged helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019817">
        <property type="entry name" value="Interferon_reg_fac_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001346">
        <property type="entry name" value="Interferon_reg_fact_DNA-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017431">
        <property type="entry name" value="IRF1/IRF2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036388">
        <property type="entry name" value="WH-like_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036390">
        <property type="entry name" value="WH_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11949">
        <property type="entry name" value="INTERFERON REGULATORY FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11949:SF3">
        <property type="entry name" value="INTERFERON REGULATORY FACTOR 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00605">
        <property type="entry name" value="IRF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF038196">
        <property type="entry name" value="IFN_RF1/2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00267">
        <property type="entry name" value="INTFRNREGFCT"/>
    </dbReference>
    <dbReference type="SMART" id="SM00348">
        <property type="entry name" value="IRF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46785">
        <property type="entry name" value="Winged helix' DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00601">
        <property type="entry name" value="IRF_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51507">
        <property type="entry name" value="IRF_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 11" id="KW-0010">Activator</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7 11" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="4" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="10 11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9 11" id="KW-0804">Transcription</keyword>
    <keyword evidence="6 11" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="5" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="IRF tryptophan pentad repeat" evidence="15">
        <location>
            <begin position="5"/>
            <end position="113"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="118"/>
            <end position="163"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="125"/>
            <end position="147"/>
        </location>
    </feature>
    <feature type="modified residue" description="N6-acetyllysine" evidence="12">
        <location>
            <position position="75"/>
        </location>
    </feature>
    <feature type="modified residue" description="N6-acetyllysine" evidence="12">
        <location>
            <position position="78"/>
        </location>
    </feature>
    <feature type="cross-link" description="Glycyl lysine isopeptide (Lys-Gly) (interchain with G-Cter in SUMO2)" evidence="13">
        <location>
            <position position="253"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PIRNR" id="PIRNR038196"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PIRSR" id="PIRSR038196-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PIRSR" id="PIRSR038196-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS51507"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029738.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="294" mass="33696" checksum="84D8FABE271A6583" modified="2019-05-08" version="1">MPVTRMRMRPWLELQINSNQIPGLVWINKDKMMFQIPWKHAAKHGWDIDKDACLFRSWAIHTGRYKVGEKEPDPKTWKANFRCAMNSLPDIEEVKDKSINKGCSAVRVYRMLPPLTKLQKKERKSKSSRESKNRSKRKSYEDQRMDEAVEAQNSSSLPDDHSGYTVHSYAEQEVEIEDASITLDLSSCDMNTMTDWRSSMEIAMADSTNDLYQLQVSPLASSSEVTDDDAEEMKEQLYKLFEQDWHATSVGGKGYLTNEPGTQNICSNFSYKEQDSEIDTTSGTNPLISEAKLL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INR8</accession>
    <name>A0A452INR8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Histone deacetylase complex subunit SAP130 C-terminal domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the SAP130 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452INR8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033521.1">
        <property type="protein sequence ID" value="ENSGAGP00000029521.1"/>
        <property type="gene ID" value="ENSGAGG00000021322.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070822">
        <property type="term" value="C:Sin3-type complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000122">
        <property type="term" value="P:negative regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024137">
        <property type="entry name" value="His_deAcase_cplx_SAP130"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031963">
        <property type="entry name" value="SAP130_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13497">
        <property type="entry name" value="HISTONE DEACETYLASE COMPLEX SUBUNIT SAP130"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13497:SF3">
        <property type="entry name" value="HISTONE DEACETYLASE COMPLEX SUBUNIT SAP130"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16014">
        <property type="entry name" value="SAP130_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0539">Nucleus</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0678">Repressor</keyword>
    <keyword evidence="5" id="KW-0804">Transcription</keyword>
    <keyword evidence="4" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="Histone deacetylase complex subunit SAP130 C-terminal" evidence="8">
        <location>
            <begin position="821"/>
            <end position="1174"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1"/>
            <end position="78"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="140"/>
            <end position="235"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="455"/>
            <end position="481"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="599"/>
            <end position="619"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="719"/>
            <end position="758"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="793"/>
            <end position="828"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="966"/>
            <end position="1014"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="7">
        <location>
            <begin position="15"/>
            <end position="26"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="7">
        <location>
            <begin position="61"/>
            <end position="78"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="7">
        <location>
            <begin position="145"/>
            <end position="155"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="161"/>
            <end position="174"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="190"/>
            <end position="207"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="208"/>
            <end position="221"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="455"/>
            <end position="467"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="794"/>
            <end position="811"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007859"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022491"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF16014"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029521.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1191" mass="127069" checksum="4042A9BE53EE89BD" modified="2019-05-08" version="1">MGTRATHRTGQSSSARHRSATLKKVQRYREPVSSLSPPPRNAHLARSPHYHSGSKLYFRPAPRRHGLGQSVRKSRPRLLPHSSRVVSAALGLNYFSQKPLLRLRFSLSLRRLGGIRQCGRDQPEPIRWRVAAAVEGELGSEHRWQREKRIRKRLRQAPSRAGQQVEMSSQQFSRSGAPPAGLGQAPPPMSTSGSTGLINPTTTAVSDESNRESEIAPREHMGPGGSLQSREEKQEPVVVRPYPQVQMLAQHHTVQSGAAVTVTAPPAHLTPAVPLSFSEGLMKPPMKPTMPSRPIAPAPPSTMSAPTKVPGQVTVTMESSIPQASAIPVATISGQQGHPSNLHHIMATNVQMSIIRSSAPGPPLHIGASHLPRGAAAAAVMSSSKVTTVLRPASQLPNAATAQPAVQHIIHQPIQSRPPVTTSSTIPPAVIATVSATRAQSPVITTTAAHATESTLSRPTLSIQQHPPSAAISIQRPAQPRDTATRITLPSHPAIGTPKQQLHTMAQKTIFSTGTPVAAATVAPILATNTIASATTAGSVSHTQAPTSTIVTMTMPSHSSHATAVTTSNIPVAKVVPQQITHTSPRIQSDYTGERSNLIPISGHRASPNPVAMETRSDNRQSVPVQFQYFLPTYPPSAYPLAAHTYTPITSSVSTIRQYPVSAQAPNSAITAQTGVGVASTVHLNPMQLMTVDASHARHIQGIQPAPISAQGIQPAPISAQGIQPAPIGTQGLHPAAPIGTQGLQPAPINAQQPQTETKTSAVVLADGATIVANPISNTFNAAPAATTVVQTHSQSASASAPAQGSSPRPSILRKKPATDGAKPKPEIHVSMATPVTVSMEAVSNQTSEQPTIAVPPTSQQPPSAIQTIIAATSPPSQPTAPLSAIPGAVPAAPATSTTIIAAPPPPSTMSGALSTVLGPPAPEIKIKEEMEPMDIMRPVSAVPPLTTNTMSPSLALLANNLSMPPNDLLPGASPRKKPRKQQHVISTEEGDMMETNSTDDEKSTAKSLLVKAEKRKSPPKEYIDEEGVRYVPVRPRPPITLLRHYRNPWKAAYHHFQRYSDVRVKEEKKAMLQEIANQKGVSCRAQGWKVHLCAAQLLQLTNLEHDVYERLTSLQEGIIPKKKAATDDDLHRINELIQGNMQRCKLVMDQISEARDSMLKVLDHKERVLKLLNKNGTVKKVSKLKRKEKV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INR9</accession>
    <name>A0A452INR9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="6">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="1 3">Belongs to the ETS family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452INR9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033625.1">
        <property type="protein sequence ID" value="ENSGAGP00000029612.1"/>
        <property type="gene ID" value="ENSGAGG00000021383.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043565">
        <property type="term" value="F:sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd08542">
        <property type="entry name" value="SAM_PNT-ETS-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.50:FF:000014">
        <property type="entry name" value="Protein c-ets-1 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.50">
        <property type="entry name" value="Transcription Factor, Ets-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.10">
        <property type="entry name" value="Winged helix-like DNA-binding domain superfamily/Winged helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045688">
        <property type="entry name" value="Ets1_N_flank"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000418">
        <property type="entry name" value="Ets_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046328">
        <property type="entry name" value="ETS_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003118">
        <property type="entry name" value="Pointed_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013761">
        <property type="entry name" value="SAM/pointed_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041886">
        <property type="entry name" value="SAM_PNT-ETS-1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036388">
        <property type="entry name" value="WH-like_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036390">
        <property type="entry name" value="WH_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11849">
        <property type="entry name" value="ETS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11849:SF209">
        <property type="entry name" value="ETS TRANSLOCATION VARIANT 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00178">
        <property type="entry name" value="Ets"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19525">
        <property type="entry name" value="Ets1_N_flank"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02198">
        <property type="entry name" value="SAM_PNT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00413">
        <property type="entry name" value="ETS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00251">
        <property type="entry name" value="SAM_PNT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47769">
        <property type="entry name" value="SAM/Pointed domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46785">
        <property type="entry name" value="Winged helix' DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00345">
        <property type="entry name" value="ETS_DOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50061">
        <property type="entry name" value="ETS_DOMAIN_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51433">
        <property type="entry name" value="PNT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 3" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0539">Nucleus</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="PNT" evidence="5">
        <location>
            <begin position="51"/>
            <end position="136"/>
        </location>
    </feature>
    <feature type="domain" description="ETS" evidence="4">
        <location>
            <begin position="335"/>
            <end position="371"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00005562"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="RuleBase" id="RU004019"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50061"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS51433"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029612.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="390" mass="44366" checksum="644217A57398C057" modified="2019-05-08" version="1">MKAAVDLKPTLTIIKTEKVDVDFFPSPDMECADVPLLTPSSKEMMSQALKATFSGFTKEQQRLGIPKDPQQWTETHVRDWVMWAVNEFSLKDVDFQKFCMNGAALCALGKDCFLELAPDFVGDILWEHLEILQKDEIKPYSANGVNPAYPESRYTSDYFISYGIEHAQCVPPSEFSEPSFITESYQTLHPISSEELFSLKYENDYPSVILRDPVQTDSLQTDYFTIKQEVVTPDNMCMGRTSRGKLGGQDSFESIESYDSCDRLTQSWNSQSSFQSLQRVPSYDSFDSEDYPAALPNHKPKGTFKDYVRDRANMNKDKPVIPAAALAGYTGSGPIQLWQFLLELLTDKSCQSFISWTGDGWEFKLSDPDEVWPIVTDQRPLKMLRRIPKS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INS3</accession>
    <name>A0A452INS3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Histone deacetylase complex subunit SAP130 C-terminal domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the SAP130 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452INS3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033526.1">
        <property type="protein sequence ID" value="ENSGAGP00000029526.1"/>
        <property type="gene ID" value="ENSGAGG00000021322.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070822">
        <property type="term" value="C:Sin3-type complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000122">
        <property type="term" value="P:negative regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024137">
        <property type="entry name" value="His_deAcase_cplx_SAP130"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031963">
        <property type="entry name" value="SAP130_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13497">
        <property type="entry name" value="HISTONE DEACETYLASE COMPLEX SUBUNIT SAP130"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13497:SF3">
        <property type="entry name" value="HISTONE DEACETYLASE COMPLEX SUBUNIT SAP130"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16014">
        <property type="entry name" value="SAP130_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0539">Nucleus</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0678">Repressor</keyword>
    <keyword evidence="5" id="KW-0804">Transcription</keyword>
    <keyword evidence="4" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="Histone deacetylase complex subunit SAP130 C-terminal" evidence="8">
        <location>
            <begin position="637"/>
            <end position="1043"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1"/>
            <end position="69"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="289"/>
            <end position="315"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="433"/>
            <end position="453"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="553"/>
            <end position="592"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="627"/>
            <end position="697"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="835"/>
            <end position="883"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="24"/>
            <end position="41"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="42"/>
            <end position="55"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="289"/>
            <end position="301"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="628"/>
            <end position="645"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="668"/>
            <end position="685"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007859"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022491"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF16014"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029526.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1060" mass="111673" checksum="AF32970899ED5C1D" modified="2019-05-08" version="1">MSSQQFSRSGAPPAGLGQAPPPMSTSGSTGLINPTTTAVSDESNRESEIAPREHMGPGGSLQSREEKQEPVVVRPYPQVQMLAQHHTVQSGAAVTVTAPPAHLTPAVPLSFSEGLMKPPMKPTMPSRPIAPAPPSTMSAPTKVPGQVTVTMESSIPQASAIPVATISGQQGHPSNLHHIMATNVQMSIIRSSAPGPPLHIGASHLPRGAAAAAVMSSSKVTTVLRPASQLPNAATAQPAVQHIIHQPIQSRPPVTTSSTIPPAVIATVSATRAQSPVITTTAAHATESTLSRPTLSIQQHPPSAAISIQRPAQPRDTATRITLPSHPAIGTPKQQLHTMAQKTIFSTGTPVAAATVAPILATNTIASATTAGSVSHTQAPTSTIVTMTMPSHSSHATAVTTSNIPVAKVVPQQITHTSPRIQSDYTGERSNLIPISGHRASPNPVAMETRSDNRQSVPVQFQYFLPTYPPSAYPLAAHTYTPITSSVSTIRQYPVSAQAPNSAITAQTGVGVASTVHLNPMQLMTVDASHARHIQGIQPAPISAQGIQPAPISAQGIQPAPIGTQGLHPAAPIGTQGLQPAPINAQQPQTETKTSAVVLADGATIVANPISNTFNAAPAATTVVQTHSQSASASAPAQGSSPRPSILRKKPATDGLAVRKSLIPPQPSEVASTRVESSVRSTSGSPRPAGAKPKPEIHVSMATPVTVSMEAVSNQTSEQPTIAVPPTSQQPPSAIQTIIAATSPPSQPTAPLSAIPGAVPAAPATSTTIIAAPPPPSTMSGALSTVLGPPAPEIKIKEEMEPMDIMRPVSAVPPLTTNTMSPSLALLANNLSMPPNDLLPGASPRKKPRKQQHVISTEEGDMMETNSTDDEKSTAKSLLVKAEKRKSPPKEYIDEEGVRYVPVRPRPPITLLRHYRNPWKAAYHHFQRYSDVRVKEEKKAMLQEIANQKGVSCRAQGWKVHLCAAQLLQLTNLEHDVYERLTSLQEGIIPKKKAATDDDLHRINELIQGNMQRCKLVMDQISEARDSMLKVLDHKERVLKLLNKNGTVKKVSKLKRKEKV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INS5</accession>
    <name>A0A452INS5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">HECT-type E3 ubiquitin transferase</fullName>
            <ecNumber evidence="4">2.3.2.26</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>S-ubiquitinyl-[E2 ubiquitin-conjugating enzyme]-L-cysteine + [acceptor protein]-L-lysine = [E2 ubiquitin-conjugating enzyme]-L-cysteine + N(6)-ubiquitinyl-[acceptor protein]-L-lysine.</text>
            <dbReference type="EC" id="2.3.2.26"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="3">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="2.3.2.26" evidence="4"/>
    <dbReference type="AlphaFoldDB" id="A0A452INS5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033556.1">
        <property type="protein sequence ID" value="ENSGAGP00000029549.1"/>
        <property type="gene ID" value="ENSGAGG00000021328.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00143"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019871">
        <property type="term" value="F:sodium channel inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061630">
        <property type="term" value="F:ubiquitin protein ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016567">
        <property type="term" value="P:protein ubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048814">
        <property type="term" value="P:regulation of dendrite morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051049">
        <property type="term" value="P:regulation of transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006511">
        <property type="term" value="P:ubiquitin-dependent protein catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00078">
        <property type="entry name" value="HECTc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00201">
        <property type="entry name" value="WW"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1750.10:FF:000026">
        <property type="entry name" value="E3 ubiquitin-protein ligase HACE1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.2160.10:FF:000001">
        <property type="entry name" value="E3 ubiquitin-protein ligase NEDD4-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.2410.10:FF:000001">
        <property type="entry name" value="E3 ubiquitin-protein ligase NEDD4-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1750.10:FF:000001">
        <property type="entry name" value="E3 ubiquitin-protein ligase NEDD4-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.70.10:FF:000006">
        <property type="entry name" value="E3 ubiquitin-protein ligase NEDD4-like protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.70.10:FF:000008">
        <property type="entry name" value="E3 ubiquitin-protein ligase NEDD4-like protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.70.10:FF:000025">
        <property type="entry name" value="E3 ubiquitin-protein ligase NEDD4-like protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.20.70.10">
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.2160.10">
        <property type="entry name" value="Hect, E3 ligase catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.2410.10">
        <property type="entry name" value="Hect, E3 ligase catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1750.10">
        <property type="entry name" value="Hect, E3 ligase catalytic domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050409">
        <property type="entry name" value="E3_ubiq-protein_ligase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000569">
        <property type="entry name" value="HECT_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035983">
        <property type="entry name" value="Hect_E3_ubiquitin_ligase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001202">
        <property type="entry name" value="WW_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036020">
        <property type="entry name" value="WW_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11254:SF440">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE NEDD-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11254">
        <property type="entry name" value="HECT DOMAIN UBIQUITIN-PROTEIN LIGASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00632">
        <property type="entry name" value="HECT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00397">
        <property type="entry name" value="WW"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00119">
        <property type="entry name" value="HECTc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00456">
        <property type="entry name" value="WW"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56204">
        <property type="entry name" value="Hect, E3 ligase catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51045">
        <property type="entry name" value="WW domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50237">
        <property type="entry name" value="HECT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01159">
        <property type="entry name" value="WW_DOMAIN_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50020">
        <property type="entry name" value="WW_DOMAIN_2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="7" id="KW-0808">Transferase</keyword>
    <keyword evidence="10" id="KW-0832">Ubl conjugation</keyword>
    <keyword evidence="9 11" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="WW" evidence="13">
        <location>
            <begin position="483"/>
            <end position="516"/>
        </location>
    </feature>
    <feature type="domain" description="WW" evidence="13">
        <location>
            <begin position="655"/>
            <end position="688"/>
        </location>
    </feature>
    <feature type="domain" description="WW" evidence="13">
        <location>
            <begin position="767"/>
            <end position="800"/>
        </location>
    </feature>
    <feature type="domain" description="WW" evidence="13">
        <location>
            <begin position="818"/>
            <end position="842"/>
        </location>
    </feature>
    <feature type="domain" description="HECT" evidence="14">
        <location>
            <begin position="929"/>
            <end position="1263"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="130"/>
            <end position="159"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="177"/>
            <end position="199"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="312"/>
            <end position="335"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="362"/>
            <end position="390"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="445"/>
            <end position="490"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="534"/>
            <end position="628"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="696"/>
            <end position="765"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="133"/>
            <end position="148"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="318"/>
            <end position="334"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="379"/>
            <end position="389"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="454"/>
            <end position="465"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="571"/>
            <end position="580"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="730"/>
            <end position="741"/>
        </location>
    </feature>
    <feature type="active site" description="Glycyl thioester intermediate" evidence="11">
        <location>
            <position position="1231"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000885"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00012485"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00104"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50020"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50237"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029549.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1264" mass="144004" checksum="37AA380774D5BE7A" modified="2019-05-08" version="1">MAHRLRFHFGSGRSNTAPESEILDQEREDDFFMAFHTLPRRNSPHPFSQHGADYGGGDLQEVGSLKRSTSMFIPQLLNNIDVRPTRSSSMQISLQRKAVNGCADECGAPESPEETLSCKFSDFREHYTCPVDSHSSSPSSPQVTSENSPPRMPEETGQRLNGAVSCNHRIFCTIPSNNQNDDVATTAEEDEASEAASGLNISGVRIQHRASSADMHQVMLSPFGSEVQDNALASEPRRWSLQHLPDTSANSGKKLFVFQLQQSQASGTGSDYNFGFTGTKGNRLVRYPRIRLERSTSYPVQPRPEQISLVEDEGNSELHGNSRNGSEISRSNSVERIPQGQECLFKIRPDQNTRQQHFRILVTRGTEEQNQVSDEDSPDTSGPTVTSATTRDDFLGQVDVPLNHLPTEDPTMERPYTFKDFLLRPRSHKSRVKGFLRLKMAYMPKNGGQEEETTEQREESERAWEVVDSNDSVSHHQEELPPPPLPPGWEEKVDNLGRTYYVNHNNRTTQWHRPSLIDVASDSDNNIRQINQEAAHRRFRSRRHISEDLEPEPSESGDASEPWETISEEASISGDSLSLSLPPPPASPVSRTSPQELSDELNRRLQVTPDSNGEQFGSLIQRDPSSRLRSCSVTDAVAEQSHLSLPSVAYVHTTPGLPSGWEERKDAKGRTYYVNHNNRTTTWTRPIMQLAEDGAVGSAANSNNHLSEPQIRRPRSLSSPTVTLSAPLEFAKDSPVRRAVKDTLSNPQSPQPSPYNSPKPQHKVTQSFLPPGWEMRIAPNGRPFFIDHNTKTTTWEDPRLKFPVHLRSKASLNPNDLGPLPPGWEERIHLDGRTFYIDHRSQVLICGDIDTRDLVPSYDSKITQWEDPRLQNPAITGPAVPYSREFKQKYDYFRKKLKKPADIPNRFEMKLHRNNIFEESYRRIMSVKRSDVLKARLWIEFESEKGLDYGGVAREWFFLLSKEMFNPYYGLFEYSATDNYTLQINPNSGLCNEDHLSYFTFIGRVAGLAVFHGKLLDGFFIRPFYKMMLGKQITLKDMESVDSEYYNSLKWILENDPTELDLMFCIDEENFGQTYQVDLKPNGSEIMVTNENKREYIDLVIQWRFVNRVQKQMNAFLEGFTELLPIDLIKIFDENELELLMCGLGDVDVNDWRQHTIYKNGYCPNHPVIQWYWKAVLLMDAEKRIRLLQFVTGTSRVPMNGFAELYGSNGPQLFTIEQWGSPEKLPRAHTCFNRLDLPPYESFEDLREKLLMAVENAQGFEGVD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INS9</accession>
    <name>A0A452INS9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Cytoplasmic FMR1-interacting protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="1 2">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="2" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="1" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="1" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452INS9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033515.1">
        <property type="protein sequence ID" value="ENSGAGP00000029515.1"/>
        <property type="gene ID" value="ENSGAGG00000020525.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031267">
        <property type="term" value="F:small GTPase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030833">
        <property type="term" value="P:regulation of actin filament polymerization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008081">
        <property type="entry name" value="Cytoplasmic_FMR1-int"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12195">
        <property type="entry name" value="CYTOPLASMIC FMR1-INTERACTING PROTEIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05994">
        <property type="entry name" value="FragX_IP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF008153">
        <property type="entry name" value="FMR1_interacting"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01698">
        <property type="entry name" value="CYTOFMRPINTP"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000313" key="1">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029515.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="3">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1114" mass="129096" checksum="2EF9A9268E2ED3A7" modified="2019-05-08" version="1">MTAQVTLEDALSNVDLLEELPLPDQQPCIEPPPSSLLYQPNFNTNFEDRNAFVTGIARYIEQATVHSNMVKCNEQPNRVEIYEKTVEVLEPEVTKLMNFMYFQRNAIERFCGEVKRLCHAERRKDFVSEAYLITLGKFINMFAVLDELKNMKCSVKNDHSAYKRAAQFLRKMADPQSIQESQNLSMFLANHNKITQSLQQQLEVISGYEELLADIVNLCVDYYENKMYLTPSEKHMLLKVMGFGLYLMDGSVSNIYKLDAKKRINLTKIDKFFKQLQVVPLFGDMQIELARYIKTSAHYEENKSRWTCTSSSSSPQYNICEQMIQIREDHMRFISELARYSNSEVVTGSGRQEAQKTDAEYRKLFDLSLQGLQLLSQWSAHVMEVYSWKLVHPTDKYSNKDCPDNAEEYERATRYNYTSEEKFALVEVIAMIKGLQVLMGRMESVFNHAIRHTIYAALQDFSQVTLREPLRQAIKKKKNVIQSVLQAIRKTVCDWEAGHEPFNDPALRGEKDPKSGFDIKVPRYVLYSLDLYNDSAHYALTKFKKQFLYDEIEAEVNLCFDQFVYKLADQIFAYYKIMAGSLLLDKRLRSECKNQGATIHLPPSNRYETLLKQRHVQLLGRSIDLNRLITQRISAAMYKSMELAIGRFESEDLTSIVELDGLVEINKMTHKLLSRYMTLDSFDAMFREANHNVSAPYGRITLHVFWELNYDFLPNYCYNGSTNRFVRTVLPFSQEFQRDKQPNAQPQYLHGSKALNLTYSSIYSNYRNFVGPPHFKVICRLLGYQGIAVVMEELLKVVKSLLQGTILQYVKTLMEVMPKICRLPRHEYGSPGILEFFHHQLKDIVEYAELKTVCFQNLREVGNAVLFCLLIEQSLSLEEVCDLLHAAPFQNILPRVHVKEGERLDAKMKRLESKYAPLHLVPLIERLGTPQQIAIAREGDLLTKERLCCGLSMFEVILTRIRTFLDDPIWRGPLPSNGVMHVDECVEFHRLWSAMQFVYCIPVGTHEFTVEQCFGDGLHWAGCMIIVLLGQQRRFDVLDFCYHLLKVQKHDGKDEVIKNVPLKKMVERIRKFQILNNEIITILDKYMKSGDGEGTPVEHVRCFQPPIHQSLASN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INV2</accession>
    <name>A0A452INV2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">C2H2-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452INV2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033591.1">
        <property type="protein sequence ID" value="ENSGAGP00000029581.1"/>
        <property type="gene ID" value="ENSGAGG00000021361.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006356">
        <property type="term" value="P:regulation of transcription by RNA polymerase I"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040436">
        <property type="entry name" value="Disconnected-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15021">
        <property type="entry name" value="DISCONNECTED-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15021:SF1">
        <property type="entry name" value="ZINC FINGER PROTEIN BASONUCLIN-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12874">
        <property type="entry name" value="zf-met"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0862">Zinc</keyword>
    <keyword evidence="1" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="3">
        <location>
            <begin position="334"/>
            <end position="362"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="3">
        <location>
            <begin position="700"/>
            <end position="728"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="3">
        <location>
            <begin position="908"/>
            <end position="931"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="235"/>
            <end position="254"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="275"/>
            <end position="327"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="379"/>
            <end position="400"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="951"/>
            <end position="974"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="237"/>
            <end position="254"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="297"/>
            <end position="306"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="307"/>
            <end position="323"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029581.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="974" mass="109708" checksum="F2300C285E63BC45" modified="2019-05-08" version="1">KEKCFKSSAIRCTLVNCSCLCFKPGKINQRQCDQCRHGWVAHALSKLRIPNLYPTSQVEIVQSNVVFDISSLMLYGTQAIPVRLKILLDRLFSVLKQEEVIQILHALDWTLQDYIRGYVLQDASGKVLDHWSIMTNEEELATLQQFLRFGETKSIVELMAIQEKEGQSIIIPTTTTNLDIRAFIESCNQRSPSLSSSVDKMSPANIHHFENIVNNMAFMLPFQFFSPVPPPLIGSPPERHLLEQGQDRSNETKQDVQIPFSESSFLISSSTSFQVENERSINSPDATTKTEDDAPLSDSSSHNTITKLERTELSPENKMKSIEKNGIGPRKGRVFCTACEKTFYDKGTLKIHYNAVHLKIKHKCTIEGCNMVFSSLRSRNRHSANPNPRLHMPMNRNNRDKDLRNSLTIAGPEETKRMDFTILTPDSRPITSYVSHCTDSKVQSSFPSIGQNGVLFPNLKTVQPVLPFYRSPVTPAELANTPGILPSLPLISSSIPEQLVAKGLPFDALPKKKSRKSSMPIKIEKEAVETANENSNLASLEDDTPLQVVSDGELETCEHRIEKQMNDRVEKHPISGNSWKSLSGVEGPKYFDSVIEPNNKYIKETSENELDHSKREVTPEENQALKIASHEMVSEDPEHHHSDHRIPNNDCTELPHHLLTGGLFSALSNRGAAIPCLEDSKDMDHISQHTLGIQKAESRFHCDICKKTFKNPYSVKMHYKNVHLKEMHICTVEGCNAAFPSRRSRDRHSSNLNLHHKLLTKDTLEFKDNYFNATYLLKDMAKEVCQDVSLKQHVGQTSVIFKGMNRTGSLVFPLSKIREPCSENYGYDPMNDGAVLDLSTTSSIKSESSAHSSWDSDGGSEECIMPLDDSDESCEGPSLIPNEELYPDCSVIEKANQSFTNLPSNLPITCHICQKNYSNKGTFRAHYKTVHLRQLHKCKVPGCNTMFSSVRSRNRHSQNPNLHRSLTRSPNSLQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INV4</accession>
    <name>A0A452INV4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="22">Cell division cycle 14B</fullName>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="15 23">
            <text>O-phospho-L-seryl-[protein] + H2O = L-seryl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:20629"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="EC" id="3.1.3.16"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="16 23">
            <text>O-phospho-L-threonyl-[protein] + H2O = L-threonyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:47004"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="EC" id="3.1.3.16"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="14">Cell projection</location>
            <location evidence="14">Kinocilium</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
            <location evidence="2">Microtubule organizing center</location>
            <location evidence="2">Centrosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Cytoskeleton</location>
            <location evidence="3">Spindle pole</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the protein-tyrosine phosphatase family. Non-receptor class CDC14 subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452INV4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033566.1">
        <property type="protein sequence ID" value="ENSGAGP00000029558.1"/>
        <property type="gene ID" value="ENSGAGG00000021351.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060091">
        <property type="term" value="C:kinocilium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000922">
        <property type="term" value="C:spindle pole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004722">
        <property type="term" value="F:protein serine/threonine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007605">
        <property type="term" value="P:sensory perception of sound"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd14499">
        <property type="entry name" value="CDC14_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd17657">
        <property type="entry name" value="CDC14_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000032">
        <property type="entry name" value="dual specificity protein phosphatase CDC14A isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000006">
        <property type="entry name" value="Dual specificity protein phosphatase CDC14B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044506">
        <property type="entry name" value="CDC14_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029260">
        <property type="entry name" value="DSPn"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000340">
        <property type="entry name" value="Dual-sp_phosphatase_cat-dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050561">
        <property type="entry name" value="PTP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016130">
        <property type="entry name" value="Tyr_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020422">
        <property type="entry name" value="TYR_PHOSPHATASE_DUAL_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23339">
        <property type="entry name" value="TYROSINE SPECIFIC PROTEIN PHOSPHATASE AND DUAL SPECIFICITY PROTEIN PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00782">
        <property type="entry name" value="DSPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14671">
        <property type="entry name" value="DSPn"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00195">
        <property type="entry name" value="DSPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00383">
        <property type="entry name" value="TYR_PHOSPHATASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50054">
        <property type="entry name" value="TYR_PHOSPHATASE_DUAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="13" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="7" id="KW-0132">Cell division</keyword>
    <keyword evidence="12" id="KW-0966">Cell projection</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="10" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="9" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="18">
        <location>
            <begin position="212"/>
            <end position="371"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="19">
        <location>
            <begin position="294"/>
            <end position="356"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1"/>
            <end position="44"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="452"/>
            <end position="495"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="17">
        <location>
            <begin position="1"/>
            <end position="13"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="452"/>
            <end position="465"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="17">
        <location>
            <begin position="470"/>
            <end position="480"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004647"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00007315"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00037822"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00047761"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00048336"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029558.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="22">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="495" mass="55789" checksum="077BBD4A588AA646" modified="2019-05-08" version="1">MKRKNWAGARRHSAPAPPAVKKTRSAAAAAAGVEEEPPPSGPESCMEITGRLYFAILCQKPKSGVANTHYFCIDDELVYENFYADFGPLNLAMVYRYCCKLNKKLKSFTLIRKKIIHYTGFDQKKQANAAFLIGSYAIIYLKESPEDVYRLLLAGNASYLPFRDASFGTCSFHLTLLDCFHAINKALRYGFLDFSAFDVNEYEHYEKAENGDFNWIIPNKFIAFSGPHSRSKTENGYPHHAPEAYFPYFRRHKVTTIIRLNRKMYDAKRFADAGFEHYDLFFADGSIPSDTIVKTFLNICENAEGVIAVHCKAGLGRTGTLIACYIMKHYRMTAAETIAWIRINRPGSVIGPQQHFLVEKQTELWREGDVFRAKLKENRKIAVTKILTGVDDISINDAKNRRANRKDTELYSDEDDANCVTQGDKLRALKSKRQAKAAATVPLTVILQSSVQKSKTSEPSISGSADINKRTTRSAARKSSLKSGHSIPRTRTVLR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INV8</accession>
    <name>A0A452INV8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="22">Cell division cycle 14B</fullName>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="15 23">
            <text>O-phospho-L-seryl-[protein] + H2O = L-seryl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:20629"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="EC" id="3.1.3.16"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="16 23">
            <text>O-phospho-L-threonyl-[protein] + H2O = L-threonyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:47004"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="EC" id="3.1.3.16"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="14">Cell projection</location>
            <location evidence="14">Kinocilium</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
            <location evidence="2">Microtubule organizing center</location>
            <location evidence="2">Centrosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Cytoskeleton</location>
            <location evidence="3">Spindle pole</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the protein-tyrosine phosphatase family. Non-receptor class CDC14 subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452INV8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033571.1">
        <property type="protein sequence ID" value="ENSGAGP00000029563.1"/>
        <property type="gene ID" value="ENSGAGG00000021351.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000033573.1">
        <property type="protein sequence ID" value="ENSGAGP00000029565.1"/>
        <property type="gene ID" value="ENSGAGG00000021351.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060091">
        <property type="term" value="C:kinocilium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000922">
        <property type="term" value="C:spindle pole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004722">
        <property type="term" value="F:protein serine/threonine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007605">
        <property type="term" value="P:sensory perception of sound"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd14499">
        <property type="entry name" value="CDC14_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd17657">
        <property type="entry name" value="CDC14_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000032">
        <property type="entry name" value="dual specificity protein phosphatase CDC14A isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000006">
        <property type="entry name" value="Dual specificity protein phosphatase CDC14B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044506">
        <property type="entry name" value="CDC14_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029260">
        <property type="entry name" value="DSPn"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000340">
        <property type="entry name" value="Dual-sp_phosphatase_cat-dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050561">
        <property type="entry name" value="PTP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016130">
        <property type="entry name" value="Tyr_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020422">
        <property type="entry name" value="TYR_PHOSPHATASE_DUAL_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23339">
        <property type="entry name" value="TYROSINE SPECIFIC PROTEIN PHOSPHATASE AND DUAL SPECIFICITY PROTEIN PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00782">
        <property type="entry name" value="DSPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14671">
        <property type="entry name" value="DSPn"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00195">
        <property type="entry name" value="DSPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00383">
        <property type="entry name" value="TYR_PHOSPHATASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50054">
        <property type="entry name" value="TYR_PHOSPHATASE_DUAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="13" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="7" id="KW-0132">Cell division</keyword>
    <keyword evidence="12" id="KW-0966">Cell projection</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="10" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="9" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="18">
        <location>
            <begin position="212"/>
            <end position="371"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="19">
        <location>
            <begin position="294"/>
            <end position="356"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1"/>
            <end position="44"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="17">
        <location>
            <begin position="1"/>
            <end position="13"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004647"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00007315"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00037822"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00047761"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00048336"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029563.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="22">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="456" mass="51672" checksum="3AEE96194F5388C9" modified="2019-05-08" version="1">MKRKNWAGARRHSAPAPPAVKKTRSAAAAAAGVEEEPPPSGPESCMEITGRLYFAILCQKPKSGVANTHYFCIDDELVYENFYADFGPLNLAMVYRYCCKLNKKLKSFTLIRKKIIHYTGFDQKKQANAAFLIGSYAIIYLKESPEDVYRLLLAGNASYLPFRDASFGTCSFHLTLLDCFHAINKALRYGFLDFSAFDVNEYEHYEKAENGDFNWIIPNKFIAFSGPHSRSKTENGYPHHAPEAYFPYFRRHKVTTIIRLNRKMYDAKRFADAGFEHYDLFFADGSIPSDTIVKTFLNICENAEGVIAVHCKAGLGRTGTLIACYIMKHYRMTAAETIAWIRINRPGSVIGPQQHFLVEKQTELWREGDVFRAKLKENRKIAVTKILTGVDDISINDAKNRRANRKDTELYSDEDDANCVTQGDKLRALKSKRQAKAAATVPLTGHSIPRTRTVLR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INV9</accession>
    <name>A0A452INV9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">CYRIA/CYRIB Rac1 binding domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the CYFIP family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452INV9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033547.1">
        <property type="protein sequence ID" value="ENSGAGP00000029542.1"/>
        <property type="gene ID" value="ENSGAGG00000020525.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031267">
        <property type="term" value="F:small GTPase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030833">
        <property type="term" value="P:regulation of actin filament polymerization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009828">
        <property type="entry name" value="CYRIA/CYRIB_Rac1-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008081">
        <property type="entry name" value="Cytoplasmic_FMR1-int"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12195">
        <property type="entry name" value="CYTOPLASMIC FMR1-INTERACTING PROTEIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07159">
        <property type="entry name" value="CYRIA-B_Rac1-bd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05994">
        <property type="entry name" value="FragX_IP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF008153">
        <property type="entry name" value="FMR1_interacting"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01698">
        <property type="entry name" value="CYTOFMRPINTP"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="CYRIA/CYRIB Rac1 binding" evidence="2">
        <location>
            <begin position="64"/>
            <end position="275"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00025790"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF07159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029542.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="796" mass="92537" checksum="0F8D68CCFE0BC5C3" modified="2019-05-08" version="1">MTAQVTLEDALSNVDLLEELPLPDQQPCIEPPPSSLLYQPNFNTNFEDRNAFVTGIARYIEQATVHSNMNEMLEDGQEYAVMLYTWRSCSRAIPQVKCNEQPNRVEIYEKTVEVLEPEVTKLMNFMYFQRNAIERFCGEVKRLCHAERRKDFVSEAYLITLGKFINMFAVLDELKNMKCSVKNDHSAYKRAAQFLRKMADPQSIQESQNLSMFLANHNKITQSLQQQLEVISGYEELLADIVNLCVDYYENKMYLTPSEKHMLLKVMGFGLYLMDGSVSNIYKLDAKKRINLTKIDKFFKQLQVVPLFGDMQIELARYIKTSAHYEENKSRWTCTSSSSSPQYNICEQMIQIREDHMRFISELARYSNSEVVTGSGRQEAQKTDAEYRKLFDLSLQGLQLLSQWSAHVMEVYSWKLVHPTDKYSNKDCPDNAEEYERATRYNYTSEEKFALVEVIAMIKGLQVLMGRMESVFNHAIRHTIYAALQDFSQVTLREPLRQAIKKKKNVIQSVLQAIRKTVCDWEAGHEPFNDPALRGEKDPKSGFDIKVPRYVLYSLDLYNDSAHYALTKFKKQFLYDEIEAEVNLCFDQFVYKLADQIFAYYKIMAGSLLLDKRLRSECKNQGATIHLPPSNRYETLLKQRHVQLLGRSIDLNRLITQRISAAMYKSMELAIGRFESEDLTSIVELDGLVEINKMTHKLLSRYMTLDSFDAMFREANHNVSAPYGRITLHVFWELNYDFLPNYCYNGSTNRFVRTVLPFSQEFQRDKQPNAQKYSSEVLHVRSQGNDCICKMCASKT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INX7</accession>
    <name>A0A452INX7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">Ras-GAP domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452INX7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033595.1">
        <property type="protein sequence ID" value="ENSGAGP00000029585.1"/>
        <property type="gene ID" value="ENSGAGG00000019812.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005938">
        <property type="term" value="C:cell cortex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903479">
        <property type="term" value="P:mitotic actomyosin contractile ring assembly actin filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05131">
        <property type="entry name" value="RasGAP_IQGAP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00201">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.506.10:FF:000004">
        <property type="entry name" value="IQ motif containing GTPase activating protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.506.10">
        <property type="entry name" value="GTPase Activation - p120gap, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000048">
        <property type="entry name" value="IQ_motif_EF-hand-BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000593">
        <property type="entry name" value="RasGAP_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023152">
        <property type="entry name" value="RasGAP_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001936">
        <property type="entry name" value="RasGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001202">
        <property type="entry name" value="WW_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149">
        <property type="entry name" value="RAS GTPASE-ACTIVATING PROTEIN WITH IQ MOTIF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149:SF12">
        <property type="entry name" value="RAS GTPASE-ACTIVATING-LIKE PROTEIN IQGAP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00612">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00616">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03836">
        <property type="entry name" value="RasGAP_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00015">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00323">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00456">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF143885">
        <property type="entry name" value="RGC domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00509">
        <property type="entry name" value="RAS_GTPASE_ACTIV_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50018">
        <property type="entry name" value="RAS_GTPASE_ACTIV_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01159">
        <property type="entry name" value="WW_DOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Ras-GAP" evidence="1">
        <location>
            <begin position="586"/>
            <end position="819"/>
        </location>
    </feature>
    <evidence type="ECO:0000259" key="1">
        <source>
            <dbReference type="PROSITE" id="PS50018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029585.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1246" mass="142835" checksum="CA148680553E8496" modified="2019-05-08" version="1">LTHKTIRSHNWCNGKNPINFLYFPPELVAVDHINTAIRNCDANKTLVALMKPEAQLPVVHPFAAAVYQTELFNLQQQNAVRYLAHDELSIAVEMLSAVVLLNQALGSKDILAIKNHLRNPSVGFNNVEDENFQRYADTLLSIKSEASSQGQDYLSWNDVQNCIDMVNMQVQEENDRIVAVSYINEAIDQGNPGKTLETLLLPTVKLHDVNPTNARHYQDVLHDAKVQKCKVEPQLSAVLWLDEIQKGINDANSNIEKAANCKYQPESTLKYHFFQDSSEGPWLKLIMKNRYDYYYNVETGESTFVPPEGFIPKTSWLTSEEIQTIVGQVTADYNREQLWFANENLILQLQALARGFLVRKSYEERKGFLQKQEPSVIKIQVLLCNIKYSSLSLFIEHDLLFEVSCSMLSKVFSLWSFFQNEEIVKIQAFLRANKARDDYRTLIGAENPPLNVLRKFAYLLDQSDLDFQEELEVTRLREEVVTKIRSNQQLEKDLNLMDIKIGLLVKNRISLQDVVLHSKKLNKKSKNQVEEMVTGDKQGIKGLSKERRKQLEAYQHLFYLLQTNPMYLAKLIFQMPQNKSTKFMDTVIFTLYNYASNQREEYLLLKLFKTALEEEINSKVDQIQDIVTGNPTVIKMVVSFNRGARGQNSLRQLLAPVVKEIMDDKSLLINTSPVEVYKAWVNQLETQTGEASKLPYDVTTEQALEHPEVVNRLESSIQSLRTVTDKVLVSIFSSLSMMPYGMRYIAKVLKNSLHEKFPDATEDELLKIVGNLLYYRYMNPAIVAPDGFDIIDMTAGGQIHSDQRRNLGCVAKVLQHAASNKLFEGESAHLSSMNTYLSQTYQKFRNFFQIACDVPEPEEKFNIDEYSDMVTLSKPVIYISIEEIINTHSLLLEHQDAIAPEQNDLLNELLKGLGAVPDVESFLGEGAVDPNDPNKENTLSQLAKTEISLSLTSKYELQEGEDQDLKGLMIKTKRLVVDVIRTQPGDTLLEILEIPASAQQESEHLKVVEKRAILDSKTPEKMKHSQSMLEDGQLPIEQKKRKIQRNLRTLEQAGLVSSANKYQEIINEIAKDIRNQRRYRHHRKAELVKLQQTLSALQSKTAFYEDQVNYYNTYIKTCLDNLTRKNSRRSIKLDGKVEEKGSKKLKHTSLKYTAARLHEKGVILEIEDLQTNQFKNVMFDITPGEEMGDFEIKAKFLGVQMEKVELHFQDLLQMQYEGVAVMKMFDKAKVNVNLLIFLLNKKFYGK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INX9</accession>
    <name>A0A452INX9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">UDP-glucose glycoprotein glucosyltransferase 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="pathway">
        <text evidence="2">Protein modification; protein glycosylation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum lumen</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the glycosyltransferase 8 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452INX9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033846.1">
        <property type="protein sequence ID" value="ENSGAGP00000029807.1"/>
        <property type="gene ID" value="ENSGAGG00000021377.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00378"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005788">
        <property type="term" value="C:endoplasmic reticulum lumen"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003980">
        <property type="term" value="F:UDP-glucose:glycoprotein glucosyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051082">
        <property type="term" value="F:unfolded protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036503">
        <property type="term" value="P:ERAD pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0018279">
        <property type="term" value="P:protein N-linked glycosylation via asparagine"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040497">
        <property type="entry name" value="Glyco_transf_24"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009448">
        <property type="entry name" value="UDP-g_GGtrans"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040693">
        <property type="entry name" value="UGGT_TRXL_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040694">
        <property type="entry name" value="UGGT_TRXL_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040692">
        <property type="entry name" value="UGGT_TRXL_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040525">
        <property type="entry name" value="UGGT_TRXL_4"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11226">
        <property type="entry name" value="UDP-GLUCOSE GLYCOPROTEIN:GLUCOSYLTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11226:SF3">
        <property type="entry name" value="UDP-GLUCOSE:GLYCOPROTEIN GLUCOSYLTRANSFERASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18404">
        <property type="entry name" value="Glyco_transf_24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18400">
        <property type="entry name" value="Thioredoxin_12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18401">
        <property type="entry name" value="Thioredoxin_13"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18402">
        <property type="entry name" value="Thioredoxin_14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18403">
        <property type="entry name" value="Thioredoxin_15"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06427">
        <property type="entry name" value="UDP-g_GGTase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="7" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 8" id="KW-0732">Signal</keyword>
    <keyword evidence="4" id="KW-0808">Transferase</keyword>
    <feature type="signal peptide" evidence="8">
        <location>
            <begin position="1"/>
            <end position="17"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019086318" description="UDP-glucose glycoprotein glucosyltransferase 1" evidence="8">
        <location>
            <begin position="18"/>
            <end position="1273"/>
        </location>
    </feature>
    <feature type="domain" description="UGGT thioredoxin-like" evidence="9">
        <location>
            <begin position="34"/>
            <end position="215"/>
        </location>
    </feature>
    <feature type="domain" description="UGGT thioredoxin-like" evidence="10">
        <location>
            <begin position="289"/>
            <end position="420"/>
        </location>
    </feature>
    <feature type="domain" description="UGGT thioredoxin-like" evidence="11">
        <location>
            <begin position="428"/>
            <end position="676"/>
        </location>
    </feature>
    <feature type="domain" description="UDP-glucose:glycoprotein glucosyltransferase thioredoxin-like" evidence="12">
        <location>
            <begin position="706"/>
            <end position="924"/>
        </location>
    </feature>
    <feature type="domain" description="Glucosyltransferase 24 catalytic" evidence="13">
        <location>
            <begin position="1226"/>
            <end position="1270"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004922"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00006351"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF18400"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="Pfam" id="PF18401"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF18402"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF18403"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="Pfam" id="PF18404"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029807.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="16">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1273" mass="143960" checksum="0FDCC930A6E7869D" modified="2019-05-08" version="1">MGIFIALLGIWFSSSLAKADSKAVTTSLTTKWLSTPLLLEASEFLAEEGQEKFWTFVEASQDIRSSDHGGTNYSSYHAMLKAACQNLSPLQQNLLKFSLSLRSYSATIQAFQQIAADEPPPVGCNSFFAVHGKKTCEFDTLEILLQTASERLKPFLFKGDHRFPLSNPESPVVILYAEIGTEEFSKFHNLLVSKASAGDITYILRHYVANPSKEKVYLSGYGVELAIKSTEYKAKDDTQVKGMEVNATMIGENDPIDEVQGFLFGKLRQLHPDLKEELKELRKHLIESTNEMAPLKVWQLQDLSFQTAARILAAPTVDALMVMKDLSQNFPTKARAITKTVVSSELRTEIEENQKYFKGTLGLQSGDSAFFINGLHIDLDTQDIFSLFDVLRNEARVMEGLHSLGIEGLSLHNVLKLNIQPSDSDYAVDIRSPAISWINNLEVDSRYNSWPSSVQELLRPTFPGVIRQIRKNFHNFVLIVDPSHESTAELLNVAEMFFSNHIPLRIGLVFVVNDSEDVDGLQDAGVALLRAFNYVAQEVDNNYAFQTLISIYNKVKTGDQLQVEHVVSVLEKQYPYVEMNSILGIDSVYDQNRKEGRGYYEQTGVGPLPIVLFNGMPFQKEQLDPDDLETITMQKILETTSLFQRAVYLGELSSDQDVVEYIMNQPNVVPRINSRILMSDREYLDLTATNNFFVDDYARFTFLNSKDKTAAVANSMTYLTKKDDSFVRPVTFWIVGDFDKPSGRQLLYDAIKHQKSSNHVRISMINNPSEDPTSKNTVITRAIWAALQTQTSNNAKNFITKMAKEETAKALEAGADITEFAVGGMDVNTFKEAFESPKVDFILSHAMYCRDVLKLQKGQRAVISNGRIIGPLEDNELFNQDDFHLLENIILKTSGQKIKSHIQQLGVEEDLASDLVMKVDALLSAQPKGEARIEYQFFEDRYSAVKLRPKEGETYFDVVAIVDPATRDAQRLAPLLMVLNQLINMNLRVFMNCQSKLSDMPLKSFYRYVLEPEISFTADDNFAPGPIAKFLDMPQTPLFTLNLNTPESWMVESVRTPYDLDNIYLEEVDNRVAAEYELEYLLLEGHCYDITTGQPPRGLQFTLGTSANPVIVDTIVMANLGYFQLKANPGAWVLRLRKGRSEDIYRIYSHDGTDSPPDANEVTVVLNNFKSKIIKVKVQKKLDMMNEDLLSDGTSENESGFWESLKWGFTGGQKNDEMKQDKDDVINIFSVASGHLYERFLRIMMLSVLKHTKTPVKFWFLKNYLSPMFKVCY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INY5</accession>
    <name>A0A452INY5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">NACHT domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452INY5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033851.1">
        <property type="protein sequence ID" value="ENSGAGP00000029812.1"/>
        <property type="gene ID" value="ENSGAGG00000021527.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006954">
        <property type="term" value="P:inflammatory response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045087">
        <property type="term" value="P:innate immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029495">
        <property type="entry name" value="NACHT-assoc"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007111">
        <property type="entry name" value="NACHT_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041267">
        <property type="entry name" value="NLRP_HD2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050637">
        <property type="entry name" value="NLRP_innate_immun_reg"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041075">
        <property type="entry name" value="NOD1/2_WH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45690">
        <property type="entry name" value="NACHT, LRR AND PYD DOMAINS-CONTAINING PROTEIN 12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45690:SF19">
        <property type="entry name" value="NACHT, LRR AND PYD DOMAINS-CONTAINING PROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14484">
        <property type="entry name" value="FISNA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05729">
        <property type="entry name" value="NACHT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17776">
        <property type="entry name" value="NLRC4_HD2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17779">
        <property type="entry name" value="WHD_NOD2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01288">
        <property type="entry name" value="FISNA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52047">
        <property type="entry name" value="RNI-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50837">
        <property type="entry name" value="NACHT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-0395">Inflammatory response</keyword>
    <keyword evidence="4" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="NACHT" evidence="8">
        <location>
            <begin position="105"/>
            <end position="310"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023198"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029812.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="571" mass="66356" checksum="9A378DFE090190CB" modified="2019-05-08" version="1">MLEVVPLWLEYYGEDVMEVTPKKYREHILEEYQVIEDRNARLGEYVSLDKRYTKLLIVKEHWQREEKEHEIITRGRKHAELMAHQTSLTTVSVLFDPDRNRESPRTVVLLGAAGIGKTMTARKIMLDWAAGKLYQKKFDYVFYINCRKINLVTAQQSMADLLLNNFPDGVRPTKEIFEKPEKLLFIIDGFDELNSVLSMEEVSLWSDPFEKKPVKTVLCSLLRKEVLQKSFLLITSRPTALEKLKQNLKFPCYAEIRGFSGDDRKEYFHKFFGDEKKATQAFNFVKENEMLFTMCFVPIVCWIICTVMKQQMERKEDLAQTSKTTTSVYLLFLTSLLTDHFTDSVQQSKTTLWKLCSLAADGILERKILFEEGDLKKHNLSLSTIHTLFLNKSIFQKDTECESVYSFIHLSFQEFFATMKKSLSSKKDVNSLLKIYGPSKNNYLMLTVRFLFGLLNKERLNDMEKKLHCKTSSAIKLDLLKWFEAELELFHCLYESQEEEFAKSAMDNFQEIRLEHIRLSTMDEIALSFCVKNCCSKQSLYLCNCTVVLPPPSSGGEEGRGPTMHPHLPPS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INY7</accession>
    <name>A0A452INY7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">Ras-GAP domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452INY7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033608.1">
        <property type="protein sequence ID" value="ENSGAGP00000029595.1"/>
        <property type="gene ID" value="ENSGAGG00000019812.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005938">
        <property type="term" value="C:cell cortex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903479">
        <property type="term" value="P:mitotic actomyosin contractile ring assembly actin filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05131">
        <property type="entry name" value="RasGAP_IQGAP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00201">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.506.10:FF:000004">
        <property type="entry name" value="IQ motif containing GTPase activating protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.506.10">
        <property type="entry name" value="GTPase Activation - p120gap, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000048">
        <property type="entry name" value="IQ_motif_EF-hand-BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000593">
        <property type="entry name" value="RasGAP_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023152">
        <property type="entry name" value="RasGAP_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001936">
        <property type="entry name" value="RasGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001202">
        <property type="entry name" value="WW_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149">
        <property type="entry name" value="RAS GTPASE-ACTIVATING PROTEIN WITH IQ MOTIF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149:SF12">
        <property type="entry name" value="RAS GTPASE-ACTIVATING-LIKE PROTEIN IQGAP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00612">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00616">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03836">
        <property type="entry name" value="RasGAP_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00015">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00323">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00456">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF143885">
        <property type="entry name" value="RGC domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00509">
        <property type="entry name" value="RAS_GTPASE_ACTIV_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50018">
        <property type="entry name" value="RAS_GTPASE_ACTIV_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01159">
        <property type="entry name" value="WW_DOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Ras-GAP" evidence="1">
        <location>
            <begin position="625"/>
            <end position="853"/>
        </location>
    </feature>
    <evidence type="ECO:0000259" key="1">
        <source>
            <dbReference type="PROSITE" id="PS50018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029595.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1272" mass="145309" checksum="ABED5A64E36547EF" modified="2019-05-08" version="1">TSPVLKEEEYSKSDGKNPINFLYFPPELVAVDHINTAIRNCDANKTLVALMKPEAQLPVVHPFAAAVYQTELFNLQQQNAVRYLAHDELSIAVEMLSAVVLLNQALGSKDILAIKNHLRNPSVGFNNVEDENFQRYADTLLSIKSEASSQGQDYLSWNDVQNCIDMVNMQVQEENDRIVAVSYINEAIDQGNPGKTLETLLLPTVKLHDVNPTNARHYQDVLHDAKVQKCKESHDESAVLWLDEIQKGINDANSNIEKAANLALGTSMINNCLEKHDSQPVLDILQSPKFGLRVVPECAETYYKNLLEAKNLKTKEDSSEGPWLKLIMKNRYDYYYNVETGESTFVPPEGFIPKTSWLTSEEIQTIVGQVTADYNREQLWFANENLILQLQALARGFLVRKSYEERKGFLQKQEPSVIKIQVLLCNIKAQSLLLEHDLLFEVSCSMLSKVFSLWSFFQNEEIVKIQAFLRANKARDDYRTLIGAENPPLNVLRKFAYLLDQSDLDFQEELEVTRLREEVVTKIRSNQQLEKDLNLMDIKIGLLVKNRISLQDVVLHSKKLNKKSKNQVEEMVTGDKQGIKGLSKERRKQLEAYQHLFYLLQTNPMYLAKLIFQMPQNKSTKFMDTVIFTLYNYASNQREEYLLLKLFKTALEEEINSKVDQIQDIVTGNPTVIKMVVSFNRGARGQNSLRQLLAPVVKEIMDDKSLLINTSPVEVYKAWVNQLETQTGEASKLPYDVTTEQALEHPEVVNRLESSIQSLRTVTDKVLVSIFSSYGMRYIAKVLKNSLHEKFPDATEDELLKIVGNLLYYRYMNPAIVAPDGFDIIDMTAGGQIHSDQRRNLGCVAKVLQHAASNKLFEGESAHLSSMNTYLSQTYQKFRNFFQIACDVPEPEEKFNIDEYSDMVTLSKPVIYISIEEIINTHSLLLEHQDAIAPEQNDLLNELLKGLGAVPDVESFLGEGAVDPNDPNKENTLSQLAKTEISLSLTSKYELQEGEDQDLKGTKRLVVDVIRTQPGDTLLEILEIPASAQQESEHLKVVEKRAILDSKTPEKMKHSQSMLEDGQLPIEQKKRKIQRNLRTLEQAGLVSSANKYQEIINEIAKDIRNQRRYRHHRKAELVKLQQTLSALQSKTAFYEDQVNYYNTYIKTCLDNLTRKSIKLDGKVEEKGSKKLKHTSLKYTAARLHEKGVILEIEDLQTNQFKNVMFDITPGEEMGDFEIKAKFLGVQMEKVELHFQDLLQMQYEGVAVMKMFDKAKVNVNLLIFLLNKKFYGK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INZ1</accession>
    <name>A0A452INZ1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">NACHT domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452INZ1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033856.1">
        <property type="protein sequence ID" value="ENSGAGP00000029817.1"/>
        <property type="gene ID" value="ENSGAGG00000021527.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006954">
        <property type="term" value="P:inflammatory response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045087">
        <property type="term" value="P:innate immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029495">
        <property type="entry name" value="NACHT-assoc"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007111">
        <property type="entry name" value="NACHT_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041267">
        <property type="entry name" value="NLRP_HD2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050637">
        <property type="entry name" value="NLRP_innate_immun_reg"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041075">
        <property type="entry name" value="NOD1/2_WH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45690">
        <property type="entry name" value="NACHT, LRR AND PYD DOMAINS-CONTAINING PROTEIN 12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45690:SF19">
        <property type="entry name" value="NACHT, LRR AND PYD DOMAINS-CONTAINING PROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14484">
        <property type="entry name" value="FISNA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05729">
        <property type="entry name" value="NACHT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17776">
        <property type="entry name" value="NLRC4_HD2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17779">
        <property type="entry name" value="WHD_NOD2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01288">
        <property type="entry name" value="FISNA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52047">
        <property type="entry name" value="RNI-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50837">
        <property type="entry name" value="NACHT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-0395">Inflammatory response</keyword>
    <keyword evidence="4" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="NACHT" evidence="8">
        <location>
            <begin position="96"/>
            <end position="414"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023198"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029817.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="592" mass="69323" checksum="80DFC0CF97D39A24" modified="2019-05-08" version="1">MLHAVIIELDYQKKYREHILEEYQVIEDRNARLGEYVSLDKRYTKLLIVKEHWQREEKEHEIITRGRKHAELMAHQTSLTTVSVLFDPDRNRESPRTVVLLGAAGIGKTMTARKIMLDWAAGKLYQKKFDYVFYINCRKINLVTAQQSMADLLLNNFPDGVRPTKEIFEKPEKLLFIIDGFDELNSVLSMEEVSLWSDPFEKKPVKTVLCSLLRKEVLQKSFLLITSRPTALEKLKQNLKFPCYAEIRGFSGDDRKEYFHKFFGDEKKATQAFNFVKENEMLFTMCFVPIVCWIICTVMKQQMERKEDLAQTSKTTTSVYLLFLTSLLTDHFTDSVQQSKTTLWKLCSLAADGILERKILFEEGDLKKHNLSLSTIHTLFLNKSIFQKDTECESVYSFIHLSFQEFFAALFYVLEEDEKTMKKSLSSKKDVNIRFLFGLLNKERLNDMEKKLHCKTSSAIKLDLLKWALRNCLNQLELFHCLYESQEEEFAKSAMDNFQEIRLEHIRLSTMDEIALSFCVKNCCSKQSLYLCNCTLGIGEREEGWIAKLRKWLFPLEPHCWCTLGSLLHHRHCPKAGLIHKPRCQWLQFCSV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INZ3</accession>
    <name>A0A452INZ3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Adhesion G-protein coupled receptor V1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="20">G-protein coupled receptor 98</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="19">Very large G-protein coupled receptor 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cell membrane</location>
            <topology evidence="3">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cell projection</location>
            <location evidence="1">Stereocilium membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Photoreceptor inner segment</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the G-protein coupled receptor 2 family. Adhesion G-protein coupled receptor (ADGR) subfamily.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000033640.1">
        <property type="protein sequence ID" value="ENSGAGP00000029626.1"/>
        <property type="gene ID" value="ENSGAGG00000021364.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001917">
        <property type="term" value="C:photoreceptor inner segment"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060171">
        <property type="term" value="C:stereocilium membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010855">
        <property type="term" value="F:adenylate cyclase inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004930">
        <property type="term" value="F:G protein-coupled receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001965">
        <property type="term" value="F:G-protein alpha-subunit binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016787">
        <property type="term" value="F:hydrolase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048513">
        <property type="term" value="P:animal organ development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007166">
        <property type="term" value="P:cell surface receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071277">
        <property type="term" value="P:cellular response to calcium ion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007605">
        <property type="term" value="P:sensory perception of sound"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007601">
        <property type="term" value="P:visual perception"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.2030:FF:000017">
        <property type="entry name" value="Adhesion G protein-coupled receptor V1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.2030:FF:000020">
        <property type="entry name" value="Adhesion G protein-coupled receptor V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.2030:FF:000021">
        <property type="entry name" value="Adhesion G protein-coupled receptor V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.2030:FF:000023">
        <property type="entry name" value="Adhesion G protein-coupled receptor V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.2030:FF:000031">
        <property type="entry name" value="Adhesion G protein-coupled receptor V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.2030:FF:000046">
        <property type="entry name" value="Adhesion G protein-coupled receptor V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1070.10:FF:000178">
        <property type="entry name" value="Adhesion G-protein coupled receptor V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.220.50:FF:000020">
        <property type="entry name" value="Adhesion G-protein coupled receptor V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.2030:FF:000007">
        <property type="entry name" value="Adhesion G-protein coupled receptor V1"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.2030:FF:000009">
        <property type="entry name" value="adhesion G-protein coupled receptor V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.2030:FF:000012">
        <property type="entry name" value="Adhesion G-protein coupled receptor V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.2030:FF:000013">
        <property type="entry name" value="Adhesion G-protein coupled receptor V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.2030:FF:000014">
        <property type="entry name" value="Adhesion G-protein coupled receptor V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.2030:FF:000028">
        <property type="entry name" value="Adhesion G-protein coupled receptor V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.200">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.220.50">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.2030">
        <property type="match status" value="34"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1070.10">
        <property type="entry name" value="Rhodopsin 7-helix transmembrane proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026919">
        <property type="entry name" value="ADGRV1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038081">
        <property type="entry name" value="CalX-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003644">
        <property type="entry name" value="Calx_beta"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009039">
        <property type="entry name" value="EAR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005492">
        <property type="entry name" value="EPTP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057244">
        <property type="entry name" value="GAIN_B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046338">
        <property type="entry name" value="GAIN_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017981">
        <property type="entry name" value="GPCR_2-like_7TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000832">
        <property type="entry name" value="GPCR_2_secretin-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46682">
        <property type="entry name" value="ADHESION G-PROTEIN COUPLED RECEPTOR V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46682:SF1">
        <property type="entry name" value="ADHESION G-PROTEIN COUPLED RECEPTOR V1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00002">
        <property type="entry name" value="7tm_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03160">
        <property type="entry name" value="Calx-beta"/>
        <property type="match status" value="32"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03736">
        <property type="entry name" value="EPTP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13385">
        <property type="entry name" value="Laminin_G_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00237">
        <property type="entry name" value="Calx_beta"/>
        <property type="match status" value="17"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF75011">
        <property type="entry name" value="3-carboxy-cis,cis-mucoante lactonizing enzyme"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF141072">
        <property type="entry name" value="CalX-like"/>
        <property type="match status" value="37"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50912">
        <property type="entry name" value="EAR"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50261">
        <property type="entry name" value="G_PROTEIN_RECEP_F2_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50221">
        <property type="entry name" value="GAIN_B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0106">Calcium</keyword>
    <keyword evidence="5" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="17" id="KW-0966">Cell projection</keyword>
    <keyword evidence="14" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="12" id="KW-0297">G-protein coupled receptor</keyword>
    <keyword evidence="9" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="13 21" id="KW-0472">Membrane</keyword>
    <keyword evidence="15" id="KW-0675">Receptor</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
    <keyword evidence="16" id="KW-0807">Transducer</keyword>
    <keyword evidence="6 21" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 21" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="27"/>
            <end position="46"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="5781"/>
            <end position="5801"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="5813"/>
            <end position="5832"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="5852"/>
            <end position="5872"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="5884"/>
            <end position="5910"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="5930"/>
            <end position="5956"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="5977"/>
            <end position="5997"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="6003"/>
            <end position="6028"/>
        </location>
    </feature>
    <feature type="domain" description="GAIN-B" evidence="22">
        <location>
            <begin position="5617"/>
            <end position="5776"/>
        </location>
    </feature>
    <feature type="domain" description="G-protein coupled receptors family 2 profile 2" evidence="23">
        <location>
            <begin position="5778"/>
            <end position="6029"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004289"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004437"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00007343"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023224"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00070037"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00078072"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00083929"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS50221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS50261"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029626.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="6179" mass="678961" checksum="6742A5767C0DE22A" modified="2019-05-08" version="1">[encoded data, 6179 characters]</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J034</accession>
    <name>A0A452J034_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Nucleoprotein TPR</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the TPR family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038192.1">
        <property type="protein sequence ID" value="ENSGAGP00000033716.1"/>
        <property type="gene ID" value="ENSGAGG00000023969.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000776">
        <property type="term" value="C:kinetochore"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031965">
        <property type="term" value="C:nuclear membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034399">
        <property type="term" value="C:nuclear periphery"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0044615">
        <property type="term" value="C:nuclear pore nuclear basket"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005819">
        <property type="term" value="C:spindle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017056">
        <property type="term" value="F:structural constituent of nuclear pore"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006406">
        <property type="term" value="P:mRNA export from nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090316">
        <property type="term" value="P:positive regulation of intracellular protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046824">
        <property type="term" value="P:positive regulation of nucleocytoplasmic transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006606">
        <property type="term" value="P:protein import into nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:1901673">
        <property type="term" value="P:regulation of mitotic spindle assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.1490">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057974">
        <property type="entry name" value="NUA/TPR/MLP1-2-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012929">
        <property type="entry name" value="Nucleoprot-TPR/MLP1-2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057577">
        <property type="entry name" value="Nucleoprot-TPR/MLP1_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18898:SF2">
        <property type="entry name" value="NUCLEOPROTEIN TPR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18898">
        <property type="entry name" value="NUCLEOPROTEIN TPR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25481">
        <property type="entry name" value="Nucleoprot-TPR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25785">
        <property type="entry name" value="TPR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07926">
        <property type="entry name" value="TPR_MLP1_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 6" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0539">Nucleus</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Nucleoprotein TPR/MPL1" evidence="9">
        <location>
            <begin position="231"/>
            <end position="310"/>
        </location>
    </feature>
    <feature type="domain" description="NUA/TPR/MLP1-2-like" evidence="10">
        <location>
            <begin position="531"/>
            <end position="628"/>
        </location>
    </feature>
    <feature type="domain" description="Nucleoprotein TPR/MLP1-2" evidence="8">
        <location>
            <begin position="1101"/>
            <end position="1228"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1"/>
            <end position="30"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="971"/>
            <end position="996"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1634"/>
            <end position="1656"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1682"/>
            <end position="1759"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1786"/>
            <end position="1822"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1834"/>
            <end position="2188"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="2291"/>
            <end position="2325"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="2337"/>
            <end position="2427"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="88"/>
            <end position="231"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="278"/>
            <end position="425"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="486"/>
            <end position="572"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="602"/>
            <end position="655"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="1056"/>
            <end position="1231"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="1283"/>
            <end position="1496"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="16"/>
            <end position="30"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="971"/>
            <end position="993"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="1636"/>
            <end position="1656"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="1682"/>
            <end position="1694"/>
        </location>
    </feature>
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        <location>
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            <end position="1715"/>
        </location>
    </feature>
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        <location>
            <begin position="1805"/>
            <end position="1822"/>
        </location>
    </feature>
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        <location>
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            <end position="1844"/>
        </location>
    </feature>
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        <location>
            <begin position="1864"/>
            <end position="1890"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="7">
        <location>
            <begin position="1931"/>
            <end position="1944"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="1947"/>
            <end position="1956"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="1987"/>
            <end position="1999"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="7">
        <location>
            <begin position="2010"/>
            <end position="2052"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="7">
        <location>
            <begin position="2062"/>
            <end position="2083"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="2091"/>
            <end position="2127"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="2291"/>
            <end position="2307"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="2308"/>
            <end position="2320"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="7">
        <location>
            <begin position="2348"/>
            <end position="2362"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="2363"/>
            <end position="2380"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="7">
        <location>
            <begin position="2416"/>
            <end position="2427"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005274"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00019789"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF07926"/>
        </source>
    </evidence>
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            <dbReference type="Pfam" id="PF25481"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="Pfam" id="PF25785"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033716.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2427" mass="273592" checksum="638E4DA7B46BFEDC" modified="2025-10-08" version="2">MPQAGQIPMSKGPGRAPSAEPASLPAPLPTATAASPAYWAAAPVSLTAPPHAQHRLPPNMAAVLQQVLERAELAKLPKPVQGKLERFLADQQSEIDGLRARHERFKVDSEQQYFEVEKRLAQSQERLVNETQECQTLRQELNKLNEQLKSLHEKNKELEAAHDRNATIQSQLTRAKEELEGEKRDLVRTSERRSQEIEHLNEDVKRLNEKLTEANSAKVELQLKLDELQTSDVSVKYREKRLEQEKELLQNQNTWLHTELKAKTDELLLISREKGNEILELKCSLENKKEEVSRMEEQVNSLKLSNENLQKHVEDLLIKLKEAKEQQAGMEERFHNELNAHIKLSNLYKSAADDSEAKSNELTRAVEELHKLLQEASEANKATEDHLAEMEESKTAMEKELGEKISKLEKELENANDLLSATKRKGAILSEEELTAMSPTAAAVAKVVKPGMKLTELYNAYVETQDQLLLEKRENKRINKYLDEIVQEVEAKAPILKRQREEYERSQKAVASLSAKLEQAMKEIQRLQEDTDKANKHASVLERENQRLEIQVKDLSQQVRVLLMELEEARGNHVIRDEEVSSADISSSSEVISQHLVSYRNIEELQQQNQRLLVALRELGEAREKEEQETTSSRISELQSQLEEAIHELEKLREARHVQLQLVDSIVRQRDMYRVLLAQTTGVVIPVQASSMLPEEISFTSTPKRSTIPQAMSTPAPVSMTESAEAVEAKAALKQLREVFENYRKEKAENDKLLNEQNEKLQEQVTDLRSQNAKMSTQLEFASKRYEMLQDNVEGYRREITSLQERTQKLTATTQKQEQIINTMTQDLRGANEKLAMAEVRAENLKKEKDILKMAEARLTQQRESLLAEQRGQNLLLTNLRTIQVILERSETETKQRLNNQVEKLEREISQLKKKLENEVEQRHVLTKNQDVHLLDTKRQLETEMNRHLNTKELLKNAQKETATLKQQLNNTEAQLASQTSQQAPGKGQSSANEDVDDLVSRLRQAEEQVNDLKERLKTSSSNVEQYRAMVLSLEESLNKEKQVTEEVRATVEARLKDSSEYQAQLEKKLMEAEKEKQELQDEKRKAIESMEQQLSELKKSLSNVQSEVQEALQRAGTAINNEQQARRDCQEQAKIASEAQNKYERELMLHAADVEALQAAKSQVAKNSAIRQQLEEATQKAESALLECKASWEERERMLKDEASKLGSRCEDLEKQNRLLHEQLENLSNKMVTSVKESVPSALNVSLNEEGKSQEQILEILRFIRREKEIAETRFEVAQVESLRYRQRVDHLERELQDLHDSLNAEREKVQITAKTIAQHEELMKKTETMNVLIETNKMLREEKERLEQELQQMQAKVRKLEADILPLQESNAELSEKSGMLQAEKKLLEEDVKRWKARTQHLLSQQKDTDLEEYRKLLSEKEANTKRIQQMSEETGRLKAEIARSNASLTTSQNLVQTLRDELTKTRTEKDTLQKELENKVADIQEKVKTITQVKKIGRRYKTQYEELKAQYDKVVAETSTQSYAEEQEQQASVQEIQELKESLSQAEVKTKALENQIENLQKTIAEKESEARSLQEQISQLQSELSRLHQDLQERTTQEEQLRQQITEKEEKTRKTLLAAKQKIAQLAGTKEQLTKENDEWKQKSSALEEQKSELEVRMSALKSQYEGRICRLERELREQQERHHEQRDEPPESTNKVPEQQRQITLKSTPASGERGIASTSDPPTANIKPTPVVSTPSKVTATAIPGSKSTPRASIRPMVTPATVTNPTTTPTATVMPTTQVETQEAMQSEGPVEHVPVFGSTSGSVRSTSPNVQTSLSQPILTVQQQTQATAFVQPTQQSHPQIEPATQEPPPTIVEVVQSSQIERPSTSTAVFGTVSATPSSSLPKRPREEEEDSTVENAEQISEETVDIPLPKKLRSVQRVGPEEEITAEESTDGEVETQIYNQDSQDSIGEGVTQGEYAPMEDSEETSQSLPIDLGPLQSDQQNTSSSQDGQSKRDDVIVIDSDDEEDDEEENEGEQEDYEDEEEEDEDDDEDTGMGDEGDDSNEGTGSADGNDGYEADDAEGADGTDPCTETEESMGGGESNQRAADSQNSGEGSTSAAESTFSHESSREQQPTSASERQASRLPQSPRRPPHPLPPRLTIHAPPQELGPPVQRIQMTRRQSVGRGLQLTPGIGGMQQHFFDDEDRTVPSTPTLVVPHRTDGFAEAIHSPQVAGVPRFRFGPPEDMPQTSSSHSDLGQLASQGGLGMYETPLFLAHEEESGGRSVPTTPLQVAAPVTVFTESTTADPSEHASQSVPMVTTSTGSLSTTTESGVGDDGDEVFVEAESEGITSEAGLEIDSQQEEESVQAPDESDLPSTSQDPPSSSSADTSSNQPKPFRRVRLQPPTLRTGVRGRQFNRQRGVTHAMGGRGGLNRGNIS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J040</accession>
    <name>A0A452J040_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Angiomotin C-terminal domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell junction</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the angiomotin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J040"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038205.1">
        <property type="protein sequence ID" value="ENSGAGP00000033727.1"/>
        <property type="gene ID" value="ENSGAGG00000024003.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005923">
        <property type="term" value="C:bicellular tight junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031410">
        <property type="term" value="C:cytoplasmic vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030036">
        <property type="term" value="P:actin cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001525">
        <property type="term" value="P:angiogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003365">
        <property type="term" value="P:establishment of cell polarity involved in ameboidal cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035329">
        <property type="term" value="P:hippo signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030334">
        <property type="term" value="P:regulation of cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009114">
        <property type="entry name" value="Angiomotin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051747">
        <property type="entry name" value="Angiomotin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024646">
        <property type="entry name" value="Angiomotin_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14826">
        <property type="entry name" value="ANGIOMOTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14826:SF12">
        <property type="entry name" value="ANGIOMOTIN-LIKE PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12240">
        <property type="entry name" value="Angiomotin_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01807">
        <property type="entry name" value="ANGIOMOTIN"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0965">Cell junction</keyword>
    <keyword evidence="5 6" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Angiomotin C-terminal" evidence="8">
        <location>
            <begin position="604"/>
            <end position="809"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="130"/>
            <end position="174"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="225"/>
            <end position="247"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="273"/>
            <end position="311"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="801"/>
            <end position="821"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="864"/>
            <end position="947"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="437"/>
            <end position="631"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="726"/>
            <end position="760"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="137"/>
            <end position="150"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="152"/>
            <end position="174"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="804"/>
            <end position="813"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="897"/>
            <end position="907"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004282"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF12240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033727.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="954" mass="106665" checksum="C93B8DA4A5D0B4C9" modified="2019-05-08" version="1">MAARLFTDWFIGIDIPFLFLIGPLSACYNPSSPVQILEDIDFLYQDHQLFASRLDVSQLTAEAVSGAKNKAVEDPLCNFHPPNFLGISEVEMRGSEDVAAGTVLQRLIQEQLRYGNPNENMNLLAIQHQATGSAGPSNSTTSTLSSTENLTQEDPQMLQQSARQEPQGQEHQVDNTVMEKQSRATQPHQNNEELPTYEEAKAQSQFFRGQQPGAVGPSFYIAGVSSQKSRTEGRPTVNRANSGQAHKDEALKELKQGHVRSLSERIMQLSLERNGAKQHPPTSGSNKGFKTGGPPPTQSSGKGMDPRGPPPEYPFKAKQMVSPVSKTQEHGLFYNDQHSGMMQMLKPSYPAPLPARTEGAVVRYQPPPEYGVTSRQCQPAFPSVPVQQHSPMSSQVSSITGPLHSATLPLLPSAVTAQPSSATPPSQQLTPDAYAIVERAQQMVEMLSEENHVLRQELEGYYEKADKLQKFEIEIQRISEAYEGLVKTTTKRESLDKAMRNKLEGEIRRLHDFNRDLRDRLETANRQLASREFDGHEDKATEGFYASQNKEHLKERERLEMELAAVRSASEDQRRHIEILDQALNNAQAKVIKLEEELRKKQAYVEKVEKLQQALTQLQAACEKREQMERRLRTRLERELDSLRMQQRQGNYQAESLPEHNAPALMELVREKEERILALEADMTKWEQKYLEESTIRHFAMNAAATAATERDTSVINHSRNGSYNENSLEVRIWQEEEEIVQANRRCQDMEYTIKNLHAKIIEKDAMIKVLQQRLRKDAGKTDNTSLRPARSVPSIAAAMGAHSRQTSLTGNQIAEEKKEEKTWKGSIGLLLGKDHHDHSSIPALPPLPPSLSCAASSLPVSTFHAKTGSKDSSTQTDKSSELFWPNTASFPGRGRLSTTPSNSPSLRHTGAKGARDKLENSPSHGKLSDSKGRVSSLVHKPEFPDADMMEVLI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J047</accession>
    <name>A0A452J047_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">ATP-dependent RNA helicase</fullName>
            <ecNumber evidence="9">3.6.4.13</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">RNA helicase.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="7 9">
            <text>ATP + H2O = ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:13065"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="3.6.4.13"/>
        </reaction>
    </comment>
    <comment type="domain">
        <text evidence="9">The Q motif is unique to and characteristic of the DEAD box family of RNA helicases and controls ATP binding and hydrolysis.</text>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the DEAD box helicase family. DDX10/DBP4 subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.6.4.13" evidence="9"/>
    <dbReference type="AlphaFoldDB" id="A0A452J047"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038160.1">
        <property type="protein sequence ID" value="ENSGAGP00000033692.1"/>
        <property type="gene ID" value="ENSGAGG00000023967.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="RHEA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003724">
        <property type="term" value="F:RNA helicase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="CDD" id="cd17941">
        <property type="entry name" value="DEADc_DDX10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18787">
        <property type="entry name" value="SF2_C_DEAD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000874">
        <property type="entry name" value="RNA helicase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:001089">
        <property type="entry name" value="RNA helicase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011545">
        <property type="entry name" value="DEAD/DEAH_box_helicase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014001">
        <property type="entry name" value="Helicase_ATP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001650">
        <property type="entry name" value="Helicase_C-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000629">
        <property type="entry name" value="RNA-helicase_DEAD-box_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014014">
        <property type="entry name" value="RNA_helicase_DEAD_Q_motif"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025313">
        <property type="entry name" value="SPB4-like_CTE"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24031">
        <property type="entry name" value="RNA HELICASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13959">
        <property type="entry name" value="CTE_SPB4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00270">
        <property type="entry name" value="DEAD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00271">
        <property type="entry name" value="Helicase_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00487">
        <property type="entry name" value="DEXDc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01178">
        <property type="entry name" value="DUF4217"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00490">
        <property type="entry name" value="HELICc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00039">
        <property type="entry name" value="DEAD_ATP_HELICASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51192">
        <property type="entry name" value="HELICASE_ATP_BIND_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51194">
        <property type="entry name" value="HELICASE_CTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51195">
        <property type="entry name" value="Q_MOTIF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 9" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="3 9" id="KW-0347">Helicase</keyword>
    <keyword evidence="2 9" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="1 9" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 9" id="KW-0694">RNA-binding</keyword>
    <feature type="domain" description="DEAD-box RNA helicase Q" evidence="13">
        <location>
            <begin position="70"/>
            <end position="98"/>
        </location>
    </feature>
    <feature type="domain" description="Helicase ATP-binding" evidence="11">
        <location>
            <begin position="101"/>
            <end position="275"/>
        </location>
    </feature>
    <feature type="domain" description="Helicase C-terminal" evidence="12">
        <location>
            <begin position="301"/>
            <end position="450"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="564"/>
            <end position="609"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="678"/>
            <end position="703"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="725"/>
            <end position="782"/>
        </location>
    </feature>
    <feature type="short sequence motif" description="Q motif" evidence="8">
        <location>
            <begin position="70"/>
            <end position="98"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="564"/>
            <end position="583"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="10">
        <location>
            <begin position="591"/>
            <end position="604"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="728"/>
            <end position="747"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022806"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00038084"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00047984"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00552"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="RuleBase" id="RU365068"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS51192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS51194"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS51195"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033692.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="803" mass="92049" checksum="E44A8B27D7B3BC09" modified="2019-05-08" version="1">MGGIGRHVGSGDPDPDPVLNFLRWKKNYNRRKNKLRLQRKEQQRPAGVVEREGISLLEKRYPEINPNEIKRFSDFPLSKKTLKGLQEAQYRMVTEIQRQTIGLALQGKDVLGAAKTGSGKTLAFIVPALERLYRQQWTSIDGLGVLIISPTRELAYQTFEVLRKVGKNHDFSAGLIIGGKDLKKEVTKIHYVNMLICTPGRLLQHMDETSYFHASDLQMLILDEADRILDMGFTDTMNAIIENLPKKRQTLLFSATQTKSVKDLARLSLKDPEYVWVHEKAKFSTPATLDQNYVVCGLQQKINMLYSFLRSHLNKKSIVFYASCKEVQYLFRVFCRLRPGIPILALHGKQPQMKRMEVYNNFVRKKSAVLFATDVAARGLDFPAVNWVIQFDCPEDANTYIHRVGRTARYKEGGEALLVLLPSEEGGMVHQLSQRKVPVNKIKINPEKLVDIQKKMQSFLAQDKELKERAQMCFVSYLRSVYLMKNKEVFDVFKLPLAEYAHSLGLAVEPRVRFLQKAQKQLSVNESLEGKSHPEGTGQCEDACSLVSNEEVEERGTHFKEKISTVQRMKGEGRKEEPEEKELPSQVIDKEGDESEELSEEEEVPSINKIPKAESINFLADDDDDFRDLDLLTNTSNSELKKKATKTKEAKKILKRKFKVNTKIVFTEDGELVQQWPPVQKSCVDSEEKEDESGGINLDKAKERLQEEDKFDKEVYRKKIKEKHREKRLREKAERRKASRTSEKEGVEAFLDYSGNEEEFDPSTLPDPDKYRGSDQEADLDTEDRDGFSFIFVIAKLTSSVFS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J048</accession>
    <name>A0A452J048_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Angiomotin C-terminal domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell junction</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the angiomotin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J048"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038201.1">
        <property type="protein sequence ID" value="ENSGAGP00000033723.1"/>
        <property type="gene ID" value="ENSGAGG00000024003.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000038202.1">
        <property type="protein sequence ID" value="ENSGAGP00000033724.1"/>
        <property type="gene ID" value="ENSGAGG00000024003.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005923">
        <property type="term" value="C:bicellular tight junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031410">
        <property type="term" value="C:cytoplasmic vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030036">
        <property type="term" value="P:actin cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001525">
        <property type="term" value="P:angiogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003365">
        <property type="term" value="P:establishment of cell polarity involved in ameboidal cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035329">
        <property type="term" value="P:hippo signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030334">
        <property type="term" value="P:regulation of cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009114">
        <property type="entry name" value="Angiomotin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051747">
        <property type="entry name" value="Angiomotin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024646">
        <property type="entry name" value="Angiomotin_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14826">
        <property type="entry name" value="ANGIOMOTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14826:SF12">
        <property type="entry name" value="ANGIOMOTIN-LIKE PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12240">
        <property type="entry name" value="Angiomotin_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01807">
        <property type="entry name" value="ANGIOMOTIN"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0965">Cell junction</keyword>
    <keyword evidence="5 6" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Angiomotin C-terminal" evidence="8">
        <location>
            <begin position="513"/>
            <end position="718"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="39"/>
            <end position="83"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="134"/>
            <end position="156"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="182"/>
            <end position="225"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="710"/>
            <end position="730"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="773"/>
            <end position="863"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="346"/>
            <end position="540"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="635"/>
            <end position="669"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="46"/>
            <end position="59"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="61"/>
            <end position="83"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="713"/>
            <end position="722"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="806"/>
            <end position="816"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="7">
        <location>
            <begin position="854"/>
            <end position="863"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004282"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF12240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033724.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="863" mass="96553" checksum="E5F76650C3779240" modified="2019-05-08" version="1">MRGSEDVAAGTVLQRLIQEQLRYGNPNENMNLLAIQHQATGSAGPSNSTTSTLSSTENLTQEDPQMLQQSARQEPQGQEHQVDNTVMEKQSRATQPHQNNEELPTYEEAKAQSQFFRGQQPGAVGPSFYIAGVSSQKSRTEGRPTVNRANSGQAHKDEALKELKQGHVRSLSERIMQLSLERNGAKQHPPTSGSNKGFKTGGPPPTQSSGKGMDPRGPPPEYPFKAKQMVSPVSKTQEHGLFYNDQHSGMMQMLKPSYPAPLPARTEGAVVRYQPPPEYGVTSRQCQPAFPSVPVQQHSPMSSQVSSITGPLHSATLPLLPSAVTAQPSSATPPSQQLTPDAYAIVERAQQMVEMLSEENHVLRQELEGYYEKADKLQKFEIEIQRISEAYEGLVKTTTKRESLDKAMRNKLEGEIRRLHDFNRDLRDRLETANRQLASREFDGHEDKATEGFYASQNKEHLKERERLEMELAAVRSASEDQRRHIEILDQALNNAQAKVIKLEEELRKKQAYVEKVEKLQQALTQLQAACEKREQMERRLRTRLERELDSLRMQQRQGNYQAESLPEHNAPALMELVREKEERILALEADMTKWEQKYLEESTIRHFAMNAAATAATERDTSVINHSRNGSYNENSLEVRIWQEEEEIVQANRRCQDMEYTIKNLHAKIIEKDAMIKVLQQRLRKDAGKTDNTSLRPARSVPSIAAAMGAHSRQTSLTGNQIAEEKKEEKTWKGSIGLLLGKDHHDHSSIPALPPLPPSLSCAASSLPVSTFHAKTGSKDSSTQTDKSSELFWPNTASFPGRGRLSTTPSNSPSLRHTGAKGARDKLENSPSHGKLSDSKGRVSSLVHKPEFPDADMMEVLI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J049</accession>
    <name>A0A452J049_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Clathrin heavy chain</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="6">Clathrin is the major protein of the polyhedral coat of coated pits and vesicles.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="6">Cytoplasmic vesicle membrane</location>
            <topology evidence="6">Peripheral membrane protein</topology>
            <orientation evidence="6">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="6">Membrane</location>
            <location evidence="6">Coated pit</location>
            <topology evidence="6">Peripheral membrane protein</topology>
            <orientation evidence="6">Cytoplasmic side</orientation>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="1 6">Belongs to the clathrin heavy chain family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038210.1">
        <property type="protein sequence ID" value="ENSGAGP00000033731.1"/>
        <property type="gene ID" value="ENSGAGG00000024009.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030132">
        <property type="term" value="C:clathrin coat of coated pit"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030130">
        <property type="term" value="C:clathrin coat of trans-Golgi network vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071439">
        <property type="term" value="C:clathrin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045334">
        <property type="term" value="C:clathrin-coated endocytic vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070062">
        <property type="term" value="C:extracellular exosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005819">
        <property type="term" value="C:spindle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032051">
        <property type="term" value="F:clathrin light chain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005198">
        <property type="term" value="F:structural molecule activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048268">
        <property type="term" value="P:clathrin coat assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006886">
        <property type="term" value="P:intracellular protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000278">
        <property type="term" value="P:mitotic cell cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006898">
        <property type="term" value="P:receptor-mediated endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000001">
        <property type="entry name" value="Clathrin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000002">
        <property type="entry name" value="Clathrin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000007">
        <property type="entry name" value="Clathrin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000095">
        <property type="entry name" value="Clathrin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.730:FF:000001">
        <property type="entry name" value="Clathrin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.110:FF:000001">
        <property type="entry name" value="Clathrin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.730">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.110">
        <property type="entry name" value="Clathrin heavy-chain terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.10">
        <property type="entry name" value="Tetratricopeptide repeat domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055358">
        <property type="entry name" value="CHCR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000547">
        <property type="entry name" value="Clathrin_H-chain/VPS_repeat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015348">
        <property type="entry name" value="Clathrin_H-chain_linker_core"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016025">
        <property type="entry name" value="Clathrin_H-chain_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022365">
        <property type="entry name" value="Clathrin_H-chain_propeller_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016341">
        <property type="entry name" value="Clathrin_heavy_chain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011990">
        <property type="entry name" value="TPR-like_helical_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10292:SF7">
        <property type="entry name" value="CLATHRIN HEAVY CHAIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10292">
        <property type="entry name" value="CLATHRIN HEAVY CHAIN RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00637">
        <property type="entry name" value="Clathrin"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09268">
        <property type="entry name" value="Clathrin-link"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13838">
        <property type="entry name" value="Clathrin_H_link"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01394">
        <property type="entry name" value="Clathrin_propel"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF002290">
        <property type="entry name" value="Clathrin_H_chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00299">
        <property type="entry name" value="CLH"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50989">
        <property type="entry name" value="Clathrin heavy-chain terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50236">
        <property type="entry name" value="CHCR"/>
        <property type="match status" value="7"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 6" id="KW-0168">Coated pit</keyword>
    <keyword evidence="5 6" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="3 6" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="Clathrin heavy chain linker core motif" evidence="8">
        <location>
            <begin position="331"/>
            <end position="354"/>
        </location>
    </feature>
    <feature type="repeat" description="CHCR" evidence="7">
        <location>
            <begin position="537"/>
            <end position="683"/>
        </location>
    </feature>
    <feature type="repeat" description="CHCR" evidence="7">
        <location>
            <begin position="686"/>
            <end position="828"/>
        </location>
    </feature>
    <feature type="repeat" description="CHCR" evidence="7">
        <location>
            <begin position="833"/>
            <end position="972"/>
        </location>
    </feature>
    <feature type="repeat" description="CHCR" evidence="7">
        <location>
            <begin position="979"/>
            <end position="1124"/>
        </location>
    </feature>
    <feature type="repeat" description="CHCR" evidence="7">
        <location>
            <begin position="1128"/>
            <end position="1269"/>
        </location>
    </feature>
    <feature type="repeat" description="CHCR" evidence="7">
        <location>
            <begin position="1274"/>
            <end position="1420"/>
        </location>
    </feature>
    <feature type="repeat" description="CHCR" evidence="7">
        <location>
            <begin position="1423"/>
            <end position="1566"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00009535"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023176"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PIRNR" id="PIRNR002290"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01006"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF09268"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033731.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1675" mass="191655" checksum="90B6A7C8AD28C18E" modified="2019-05-08" version="1">MAQILPIRFQEHLQLQNLGINPANIGFSTLTMESDKFICIREKVGEQAQVVIIDMNDPSNPIRRPISADSAIMNPASKVIALKAGKTLQIFNIEMKSKMKAHTMTDDVTFWKWISLNTVALVTDNAVYHWSMEGESQPVKMFDRHSSLAGCQIINYRTDAKQKWLLLTGISAQQNRVVGAMQLYSVDRKVSQPIEGHAASFAQFKMEGNAEESTLFCFAVRGQAGGKLHIIEVGTPPTGNQPFPKKAVDVFFPPEAQNDFPVAMQISEKHDVVFLITKYGYIHLYDLETGTCIYMNRISGETIFVTAPHEATAGIIGVNRKGQVLSVCVEEENIIPYITNVLQNPDLALRMAVRNNLAGAEELFARKFNALFAQGNYSEAAKVAANAPKGILRTPDTIRRFQSVQAQPGQTSPLLQYFGILLDQGQLNKYESLELCRPVLQQGRKQLLEKWLKEDKLECSEELGDLVKSVDPTLALSVYLRANVPNKVIQCFAETGQVQKIVLYAKKVGYTPDWIFLLRNVMRISPDQGQQFAQMLVQDEEPLADITQIVDVFMEYNLIQQCTAFLLDALKNNRPSEGPLQTRLLEMNLMHAPQVADAILGNQMFTHYDRAHIAQLCEKAGLLQRALEHFTDLYDIKRAVVHTHLLNPEWLVNYFGSLSVEDSLECLRAMLSANIRQNLQICVQVASKYHEQLSTQSLIELFESFKSFEGLFYFLGSIVNFSQDPDVHFKYIQAACKTGQIKEVERICRESNCYDPERVKNFLKEAKLTDQLPLIIVCDRFDFVHDLVLYLYRNNLQKYIEIYVQKVNPSRLPVVIGGLLDVDCSEDVIKNLILVVRGQFSTDELVAEVEKRNRLKLLLPWLEARIHEGCEEPATHNALAKIYIDSNNNPERFLRENPYYDSRVVGKYCEKRDPHLACVAYERGQCDLELINVCNENSLFKSLSRYLVRRKDPELWGSVLLESNPYRRPLIDQVVQTALSETQDPEEVSVTVKAFMTADLPNELIELLEKIVLDNSVFSEHRNLQNLLILTAIKADRTRVMEYINRLDNYDAPDIANIAISNELFEEAFAIFRKFDVNTSAVQVLIEHIGNLDRAYEFAERCNEPAVWSQLAKAQLQKGMVKEAIDSYIKADDPSSYMEVVQAANASGNWEELVKYLQMARKKARESYVETELIFALAKTNRLAELEEFINGPNNAHIQQVGDRCYDEKMYDAAKLLYNNVSNFGRLASTLVHLGEYQAAVDGARKANSTRTWKEVCFACVDGKEFRLAQMCGLHIVVHADELEELINYYQDRGYFEELITMLEAALGLERAHMGMFTELAILYSKFKPQKMREHLELFWSRVNIPKVLRAAEQAHLWAELVFLYDKYEEYDNAIITMMNHPTDAWKEGQFKDIITKVANVELYYKAIQFYLEFKPLLLNDLLMVLSPRLDHTRAVNFFSKTKQLPLVKPYLRSVQNHNNKSVNESLNNLFIIEEDYQALRTSIDAYDNFDNISLAQRLEKHELIEFRRIAAYLFKGNNRWKQSVELCKKDRLYKDAMQYASESKDTELAEELLQWFLQEEKRECFGACLFTCYDLLRPDVVLETAWRHNIMDFAMPYFIQVMKEYLSKVDKLDASESLRKEEEQATETQPIVYGQPQLMLTAGPSVAVPPQAPFGYGYTAPPYGQPQPGFGYTM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J079</accession>
    <name>A0A452J079_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Glucose-6-phosphate exchanger SLC37A2</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="13">Solute carrier family 37 member 2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>D-glucose 6-phosphate(in) + phosphate(out) = D-glucose 6-phosphate(out) + phosphate(in)</text>
            <dbReference type="Rhea" id="RHEA:71535"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:61548"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the major facilitator superfamily. Organophosphate:Pi antiporter (OPA) (TC 2.A.1.4) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J079"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038242.1">
        <property type="protein sequence ID" value="ENSGAGP00000033761.1"/>
        <property type="gene ID" value="ENSGAGG00000024013.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061513">
        <property type="term" value="F:glucose 6-phosphate:phosphate antiporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035435">
        <property type="term" value="P:phosphate ion transmembrane transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd17344">
        <property type="entry name" value="MFS_SLC37A1_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1250.20:FF:000050">
        <property type="entry name" value="glucose-6-phosphate exchanger SLC37A2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1250.20:FF:000028">
        <property type="entry name" value="Sugar phosphate exchanger 3 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1250.20">
        <property type="entry name" value="MFS general substrate transporter like domains"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011701">
        <property type="entry name" value="MFS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020846">
        <property type="entry name" value="MFS_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036259">
        <property type="entry name" value="MFS_trans_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044740">
        <property type="entry name" value="SLC37A1_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000849">
        <property type="entry name" value="Sugar_P_transporter"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43184:SF9">
        <property type="entry name" value="GLUCOSE-6-PHOSPHATE EXCHANGER SLC37A2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43184">
        <property type="entry name" value="MAJOR FACILITATOR SUPERFAMILY TRANSPORTER 16, ISOFORM B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07690">
        <property type="entry name" value="MFS_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF002808">
        <property type="entry name" value="Hexose_phosphate_transp"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103473">
        <property type="entry name" value="MFS general substrate transporter"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50850">
        <property type="entry name" value="MFS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0050">Antiport</keyword>
    <keyword evidence="7" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9 14" id="KW-0472">Membrane</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0762">Sugar transport</keyword>
    <keyword evidence="6 14" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 14" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="12"/>
            <end position="30"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="82"/>
            <end position="99"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="106"/>
            <end position="128"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="134"/>
            <end position="162"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="174"/>
            <end position="193"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="199"/>
            <end position="220"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="269"/>
            <end position="290"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="310"/>
            <end position="327"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="334"/>
            <end position="351"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="363"/>
            <end position="388"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="400"/>
            <end position="422"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="434"/>
            <end position="454"/>
        </location>
    </feature>
    <feature type="domain" description="Major facilitator superfamily (MFS) profile" evidence="15">
        <location>
            <begin position="19"/>
            <end position="457"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004477"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009598"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022449"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022597"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00034251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00039218"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00042041"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50850"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033761.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="482" mass="52208" checksum="7BE1C73C0A5139D6" modified="2019-05-08" version="1">MVSRKTFSLGGWYRGLTLVLTFVFYASFHLSRKPISIVKSELHTNCSSLEKIPPNDTNSTTWCDWAPFDQANYNELFGALDNAFLVAYAIGMFISGIFGERLPLRYYLSGGMVLSGLFTSLFGLAYFWEIHVLWYYIIIQICNGLVQTTGWPSVVTCVGNWFGKGKRGLIMGIWNSHTSVGNILGSLIAGVWVSSAWGLSFIVPGIIIAIMGIVCFLFLVECECELVTLSADDGSCSRENTIDSSEYPKEPAEQPGAISFLGALRIPGVVEFSLCLLFAKLVSYTFLYWLPLYIVNVAHFSAKEAGDLSTLFDVGGILGGILAGLISDYTDGRATTCCVMLIVAAPMLFLYNHVGQNGIGSSIAMLIVCGALVNGPYALITTAVSADLGTHECLKGNAKALSTVTAIIDGTGSVGAALGPLLAGLISPTGWNNVFYMLITADILACLFLSRVVYKEIKGWCGCYTRKRGYVLLKSQVCRVFL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J080</accession>
    <name>A0A452J080_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="6">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the CLEC16A/gop-1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J080"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038248.1">
        <property type="protein sequence ID" value="ENSGAGP00000033767.1"/>
        <property type="gene ID" value="ENSGAGG00000024031.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005770">
        <property type="term" value="C:late endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006914">
        <property type="term" value="P:autophagy"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016197">
        <property type="term" value="P:endosomal transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1901096">
        <property type="term" value="P:regulation of autophagosome maturation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007034">
        <property type="term" value="P:vacuolar transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039272">
        <property type="entry name" value="CLEC16A/TT9"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045820">
        <property type="entry name" value="CLEC16A/TT9_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019155">
        <property type="entry name" value="CLEC16A/TT9_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21481">
        <property type="entry name" value="PROTEIN CLEC16A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21481:SF0">
        <property type="entry name" value="PROTEIN CLEC16A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19439">
        <property type="entry name" value="CLEC16A_C"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09758">
        <property type="entry name" value="FPL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0072">Autophagy</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="FPL" evidence="4">
        <location>
            <begin position="51"/>
            <end position="198"/>
        </location>
    </feature>
    <feature type="domain" description="CLEC16A/TT9 C-terminal" evidence="5">
        <location>
            <begin position="243"/>
            <end position="618"/>
        </location>
    </feature>
    <feature type="domain" description="CLEC16A/TT9 C-terminal" evidence="5">
        <location>
            <begin position="619"/>
            <end position="971"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="379"/>
            <end position="429"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="843"/>
            <end position="893"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="905"/>
            <end position="932"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="980"/>
            <end position="1003"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="3">
        <location>
            <begin position="379"/>
            <end position="390"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="3">
        <location>
            <begin position="867"/>
            <end position="884"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="905"/>
            <end position="915"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="992"/>
            <end position="1003"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006441"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023006"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="Pfam" id="PF09758"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="Pfam" id="PF19439"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033767.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1003" mass="112771" checksum="C1B93C57A2D42F4E" modified="2019-05-08" version="1">MFGRSRSWVGGGHGKAARSIHSLDHLKYMYHVLTKNTTVTDHNRNLLVETIRSITEILIWGDQNDSSVFDFFLEKNMFVFFLNILRQKSGRYVCVQLLQTLNILFENISHETSLYYLLSNNYVNSIIVHKFDFSDEEIMAYYISFLKTLSLKLNNHTVHFFYNEHTNDFALYTEAIKFFNHPESMVRIAVRTITLNVYKVDNQPMLHYIRDKTAVPYFSNLVWFIGSHVIELDNCVQTDEEHRNRGKLSDLVAEHLDHLHYLNDILIINCEFLNDVLTDHLLNRLFLPLYVYSLVNQDKGGERPKISLQVSLYLLSQVFLIIHYAPLVNSLAEVILNGDLSVFSSKVEQDIQRSSAKSSIRCFIKPTETLERSLEINKHKGKKRLQKRPNYKNVGEEEEEERGLENTQEDPDKAKGTEGSSKGIKASGENEEIEMVIMERCKMSELSISVVTEQNTTDEEKSAAASGSEMTNWSRPFLDMVYNAQDCAEDDYYALFVLCLLYAMSQNKGIDPVKLERIQLPAQHAEEKSSYNHVLAERLIRIMNYAAQPDGKVRLATLELGCLLLKQLVLSKSGSIIKDVHLACLEGAREESVHLLRRFYKGEEIFLDMFEDEYRSMTAIRVFFMLRSLSLHLRDEPETQLPLTREEDLIKTDDVLDLNNSDLIACTVITKDGGQVQRFLAVDIYQMSLVEPDVARLGWGVVKFAGLLQDMQVTGVEDDSRALNIIIHKPASSPHSKPFPILQATFIFSDHIRCIIAKQRLAKGRIQARRMKMQRIAALLDLPVQPSTEVMGFGHSSASAAQHLPFRFYDQSRRGSCDPAVQRSVFASVDKVPGFAVAQYINQPNPSPLSSPSPSSSGSGSTGHCDSVTASSTSTPSAAHSPSVDDSSAPEQPPIIFSSQMMMVDETSSAASKSGKNPVKNPEVETANLSPSLIPAQQPTISLLSDDNADALSVESLTLVPPIDPHSIHTFSCIPQSSLPSEVCGVTEGEGSDQSQQADYDEQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J084</accession>
    <name>A0A452J084_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">NACHT domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452J084"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038283.1">
        <property type="protein sequence ID" value="ENSGAGP00000033797.1"/>
        <property type="gene ID" value="ENSGAGG00000024047.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000038289.1">
        <property type="protein sequence ID" value="ENSGAGP00000033800.1"/>
        <property type="gene ID" value="ENSGAGG00000024047.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045345">
        <property type="term" value="P:positive regulation of MHC class I biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045348">
        <property type="term" value="P:positive regulation of MHC class II biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045944">
        <property type="term" value="P:positive regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:001028">
        <property type="entry name" value="Class II major histocompatibility complex transactivator"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001611">
        <property type="entry name" value="Leu-rich_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025875">
        <property type="entry name" value="Leu-rich_rpt_4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008095">
        <property type="entry name" value="MHC_II_transact"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007111">
        <property type="entry name" value="NACHT_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041267">
        <property type="entry name" value="NLRP_HD2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47189">
        <property type="entry name" value="MHC CLASS II TRANSACTIVATOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47189:SF1">
        <property type="entry name" value="MHC CLASS II TRANSACTIVATOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12799">
        <property type="entry name" value="LRR_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13516">
        <property type="entry name" value="LRR_6"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05729">
        <property type="entry name" value="NACHT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17776">
        <property type="entry name" value="NLRC4_HD2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01719">
        <property type="entry name" value="MHCIIACTVATR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00368">
        <property type="entry name" value="LRR_RI"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52047">
        <property type="entry name" value="RNI-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51450">
        <property type="entry name" value="LRR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50837">
        <property type="entry name" value="NACHT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="1" id="KW-0433">Leucine-rich repeat</keyword>
    <keyword evidence="3" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="NACHT" evidence="5">
        <location>
            <begin position="443"/>
            <end position="578"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022614"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033800.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1172" mass="131858" checksum="B52455A5D0F92A5C" modified="2019-05-08" version="1">MDSAVLTENSFLDLLHSNIDPLHLYSLLDTRLSDEESELSTDPEIDTINHDQFSNLDVFYTMESNESVEESIFSNAGEAYSRIAGLAEYVLKDQQERQMEDIFGKLILDEMANETTEGFTEMKKQKCQKRSFVSTAESYSDASKPKCRRPDAARRMTEQVCTEGSRAVGDVRASCQKRVESPGSTVNGSFVAVPLNSSPANSTSLSHLHMQFSVTSIHPTGKNVVMPVSSGNSNPDCLLVGAKGIQVISGFAVLPQQIINLPVGNGASNIIIYPVSSSPTETLQISPAFPTTLAAQVDLAVQSILGSPDKNTSIGPFAYMPETAAVTDLTDEEEISNEISQIPVVPSEQILKRPKPVEAFCTSLKDYFQDVCKFMAMDREVTLDHLYIDGALMQGQTETKSGKNVIKVMEKELVIYNLEEKEKAAIERSQIFQVLGGKELETKVIVVLGKAGMGKSILVQKICQDWSNGKFSQFEFVFWFECRRLSLPEKQYSLKDLLLELFVKPREGRDQVFEYILQNPNKVLLIFDGFEELQDHDGFIHYPDCQSYKELYSIRELFAGLFQKKILSGCTLLLTARPKDKFNQYLSKVDKIIEMIGFSPQQVELYIARYFEGLPYSDDAVKLITNCQYLFSHCYSPVICRFICFLCEMMFEMGDKDLPPTLTSLFVKYLQQKLIPTQTDVTAIGNQQHVTRLAQLAWYLGEKNQNALKSSLFPSKEVKEFALKYGLVLPFTFPKHSGNGEEEFGSAFSDFVIQNFLSALHLLLAEDIKDKSLTKYLSLPARKKKPHNWLDLVPRFLPGLLYLQHDPFLCSLSDEHVMKIITKKQKTLLKYIKKIQISDLYPEKLLELLHCVYETQDSYLLQHVALRIKAELSFLGTLLTPPDVYVLHFILKRSPKEFSLDLRNSSINLQGLKHLVGLKNVTSFRASLSDTVRLWESLERTKEYELLKISVEKFDIDPFKAETMKDIHDLSDLVQIQEKMIHCMEEAFGSCYQIPAIKNLRKLEFALGPACGLQGFLKLVEILAAFPSLQHLDLDALSENEIGDEGTKSLCEVLSELTSLETLNLSQNKITDLGAEQLATALPSLSSLKTLSLYNNYICDAGAENIAKILPAMASLRVLHVQCNKITDAGAQKLTDSLRKCPHIKSVAMWNPTIPYGVLEHLQLLDSRISLL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J094</accession>
    <name>A0A452J094_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Eukaryotic translation initiation factor 4E binding protein 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the eIF4E-binding protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J094"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038258.1">
        <property type="protein sequence ID" value="ENSGAGP00000033777.1"/>
        <property type="gene ID" value="ENSGAGG00000024040.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008190">
        <property type="term" value="F:eukaryotic initiation factor 4E binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045947">
        <property type="term" value="P:negative regulation of translational initiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008606">
        <property type="entry name" value="EIF4EBP"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12669">
        <property type="entry name" value="EUKARYOTIC TRANSLATION INITIATION FACTOR 4E-BINDING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12669:SF14">
        <property type="entry name" value="EUKARYOTIC TRANSLATION INITIATION FACTOR 4E-BINDING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05456">
        <property type="entry name" value="eIF_4EBP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0652">Protein synthesis inhibitor</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0810">Translation regulation</keyword>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="1"/>
            <end position="46"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="1"/>
            <end position="10"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="35"/>
            <end position="46"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00005480"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022845"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023193"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033777.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="7">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="111" mass="11855" checksum="EC7C095F1054C964" modified="2019-05-08" version="1">MSGSCRQGHTLSRAIPAKRLSLPDGAPLPPGDYSTTPGGTVFGTTPGGTRIIYDRKFLMECRNSPVAKTSPCDLPDIPGVTSPNVEELKIENNHVHNYLEKVGIVPRALAT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J095</accession>
    <name>A0A452J095_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Integrin alpha-V</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="19">Vitronectin receptor subunit alpha</fullName>
        </alternativeName>
    </protein>
    <organism evidence="26 27">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="27" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="26" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="26" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell junction</location>
            <location evidence="1">Focal adhesion</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2 21">Membrane</location>
            <topology evidence="2 21">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 21">Belongs to the integrin alpha chain family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J095"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038304.1">
        <property type="protein sequence ID" value="ENSGAGP00000033815.1"/>
        <property type="gene ID" value="ENSGAGG00000023825.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009897">
        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
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        <property type="term" value="C:integrin complex"/>
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        <property type="project" value="InterPro"/>
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        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001525">
        <property type="term" value="P:angiogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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    <dbReference type="GO" id="GO:0033627">
        <property type="term" value="P:cell adhesion mediated by integrin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098609">
        <property type="term" value="P:cell-cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007160">
        <property type="term" value="P:cell-matrix adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007229">
        <property type="term" value="P:integrin-mediated signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
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        <property type="entry name" value="Integrin alpha V"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Integrin alpha V"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="integrin alpha-V isoform X2"/>
        <property type="match status" value="1"/>
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    <dbReference type="FunFam" id="1.20.5.930:FF:000001">
        <property type="entry name" value="Integrin subunit alpha V"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1460:FF:000001">
        <property type="entry name" value="Integrin, alpha V"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.930">
        <property type="entry name" value="Bicelle-embedded integrin alpha(iib) transmembrane segment"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.130">
        <property type="entry name" value="Integrin alpha, N-terminal"/>
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    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1460">
        <property type="entry name" value="Integrin domains. Chain A, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1510">
        <property type="entry name" value="ntegrin, alpha v. Chain A, domain 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1530">
        <property type="entry name" value="ntegrin, alpha v. Chain A, domain 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013517">
        <property type="entry name" value="FG-GAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013519">
        <property type="entry name" value="Int_alpha_beta-p"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000413">
        <property type="entry name" value="Integrin_alpha"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018184">
        <property type="entry name" value="Integrin_alpha_C_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013649">
        <property type="entry name" value="Integrin_alpha_Ig-like_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048285">
        <property type="entry name" value="Integrin_alpha_Ig-like_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048286">
        <property type="entry name" value="Integrin_alpha_Ig-like_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028994">
        <property type="entry name" value="Integrin_alpha_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032695">
        <property type="entry name" value="Integrin_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23220">
        <property type="entry name" value="INTEGRIN ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23220:SF4">
        <property type="entry name" value="INTEGRIN ALPHA-V"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="FG-GAP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08441">
        <property type="entry name" value="Integrin_A_Ig_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20805">
        <property type="entry name" value="Integrin_A_Ig_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20806">
        <property type="entry name" value="Integrin_A_Ig_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00357">
        <property type="entry name" value="Integrin_alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01185">
        <property type="entry name" value="INTEGRINA"/>
    </dbReference>
    <dbReference type="SMART" id="SM00191">
        <property type="entry name" value="Int_alpha"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69318">
        <property type="entry name" value="Integrin alpha N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="3"/>
    </dbReference>
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        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00242">
        <property type="entry name" value="INTEGRIN_ALPHA"/>
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    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0106">Calcium</keyword>
    <keyword evidence="10 21" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="11" id="KW-0965">Cell junction</keyword>
    <keyword evidence="4" id="KW-0165">Cleavage on pair of basic residues</keyword>
    <keyword evidence="15" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="17" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13 21" id="KW-0401">Integrin</keyword>
    <keyword evidence="14 21" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="16 21" id="KW-0675">Receptor</keyword>
    <keyword evidence="27" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="7 21" id="KW-0732">Signal</keyword>
    <keyword evidence="5 21" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="12 21" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="21">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5018809939" description="Integrin alpha-V" evidence="21">
        <location>
            <begin position="21"/>
            <end position="997"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="943"/>
            <end position="965"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="20">
        <location>
            <begin position="22"/>
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        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="20">
        <location>
            <begin position="98"/>
            <end position="159"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="20">
        <location>
            <begin position="190"/>
            <end position="244"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="20">
        <location>
            <begin position="245"/>
            <end position="310"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="20">
        <location>
            <begin position="311"/>
            <end position="368"/>
        </location>
    </feature>
    <feature type="repeat" description="FG-GAP" evidence="20">
        <location>
            <begin position="372"/>
            <end position="435"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha first immunoglubulin-like" evidence="23">
        <location>
            <begin position="420"/>
            <end position="578"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha second immunoglobulin-like" evidence="24">
        <location>
            <begin position="579"/>
            <end position="707"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin alpha third immunoglobulin-like" evidence="25">
        <location>
            <begin position="725"/>
            <end position="932"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="976"/>
            <end position="997"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="22">
        <location>
            <begin position="976"/>
            <end position="991"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00008054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022685"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
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    </evidence>
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        </source>
    </evidence>
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    <sequence length="997" mass="110567" checksum="BB4122ABDF3380AE" modified="2019-05-08" version="1">MARLRLLLLLLGRWLALGGAFNLDADGPAVYSGSEGSYFGFAVDFFAPDSSSIFLLVGAPKANTTQPDIVEGGQVLKCNWNSNKNCQPVIFDSTGDRNFAPNDPLEFKSHQWFGASVRSKNDKILACAPLYHWRTEVKQEREPVGTCYLHDGSKAVEYAPCRSSQLISDRVAEIVSKYNSKVYSIKYDDQLATRSASAAFDDSYLGYSVAVGEFSGDGIEDFVSGVPRAARTLGMVSIYNGKNMSSLYNFTGEQMAAYFGYSVAATDINGDNYTDLFIGAPLFMERGSDGKLQEVGQVSICLQQASGGFQIIKLNGFEIFARFSSSIAPLGDLDQDGFNDIAVAAPYGGEDKRGVVYIYNGRATGLNAVPSQILEGQWAARTMPPSFGYSLKGATDVDKNGYPDLIVGAFGVDTAVLYRARPVISVNAGLEVNPTILNPEEKICELPDTRVKVSCFNVKFCLTADGKGKLPNTLTFQVELLLDKLKQKGALRRALFLHNKQPGHSKNMTVVKGGTMRCEELQAFLRDESEFRDKLTPITVFMEYRLDYKTAKDSAGLQPILNQFTPANISRQTHILLDCGEDNICKPKLQVAVHSDQKQIYIGDDNPLTLIVDAQNQGEGAYEAELFVIIPPQADFIGVVRNNEALARLSCAFKTENQTRLVVCDLGNPMKAGTKLLAGLRFSVNQQSEMDTSVKFDLQIRSSNLFDNLSPTTSYQADLAILASVEIRGVSSPDHIFLPIANWQPQENPETEEDIGPLVQHIYELRNNGPSAFSKVMLNLQWPYKYNNNTLLYIMQYEIDGPMNCTSDMEINPLNIKVTDEKNETIRDEDNRDHHINRRDITAIEGDVHTLGCGTAECLKIVCQVGRLERGKSAILYVKSRLWTQTFMNKENQNHSYSLKSSASFNVIEFPYKNLSFEDIHNSTVVTTNITWGIQPQPMPVPVWVIILAVLAGLLLLAVLVFVMYRMGFFKRVRPPQEEQEREQLQPHENGEGTSEA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHH1</accession>
    <name>A0A452GHH1_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000001192.1">
        <property type="protein sequence ID" value="ENSGAGP00000001058.1"/>
        <property type="gene ID" value="ENSGAGG00000000775.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060271">
        <property type="term" value="P:cilium assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR33487">
        <property type="entry name" value="CILIA- AND FLAGELLA-ASSOCIATED PROTEIN 54"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR33487:SF1">
        <property type="entry name" value="CILIA- AND FLAGELLA-ASSOCIATED PROTEIN 54"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="116"/>
            <end position="138"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="438"/>
            <end position="468"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="1">
        <location>
            <begin position="124"/>
            <end position="134"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="440"/>
            <end position="459"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001058.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2081" mass="236549" checksum="87FB76DC1EBE37E6" modified="2019-05-08" version="1">MVVAIELSNWLNDANYALQSVVQCYGLLAPIIYHKIASVPVVQILIKCLTVLQEVPSSTLQRKQTGCYESIQHMIACSSFYTAQVLRSWKEYELALIIINYGKKLLDSSQTSHGQSKFSSIEEMQIDPETEDESPYSKKKSQVAAVEKAAENLAALESNLLKLTRPGTGPEITGQEDPFFLYPIISCWTTKNAYREVMKFRKKPRFLEFFVQVLHKALNEEKFQRIVEWTGSVKDFLRRRNNHLLGIKGTLAKEVSELSVYEDAKRHTTAVVEFQKSVETSALRKPRKEVMSPFRGRESLRQYFLRHSSLGKSPEAQRKQRDEIEKVAFHTLVVLLNPIVSNYLKRKRFHQVCLEEMPWRSQMNIYLAITHFNLFKKRLEEQYNIGVRSLQHLNSYDVLDPEIFSLNNSGIIVVRGGTDDNIGTLSHLLIKRIQSGSSRKITDPDKSSERSTKFSEADTPRSQMTNDTELSITLSAREHERPMSNAILLEHFTKTFLHLRRAMVLAHRGGHWTLLQNTCRELWNHSQEALSIAKHSHSLQGLFPITRNFLISTIWLPFYLASDAILDMIIELQTSNSLKIFDPKGDFCVPSCVGGIADEDGGSNLTFCCPLDDISVVDLRWICGLVLRTIELMYQMKKWEALVHIAMHFNTVTHERYAEQVTPLLVYAQRQLLERIQQFRGPDSPQPHFIKYQGDNANKINCRNFIGKQLRIASTFNEVTVSGHHFVPEMPDTYTDGSQAQALVCVPLDVTDTLKCFRETLEKSKYHSRALRHSRKLLSLFLAHTQETVGRVPNQIFTNGKVEFSVGAELTFLPKPPDLSQEKYNFPSMVESRPLPESQLSVVVSSYEKTIEILQINNQRDLKVQALHELGNLHFYAGKKRSAFKYWCQALDETLKIADVLDNWQELEGFSKNVPDDFTDGSSKHFSEKCIYQAGIWGCLHAAVLAAKIAQYILTSSVRLRTKCCIFSAILFKTLFKVSIPHPTADCDFAQYEIHRLIPGIDLFSDRYRADVSTVVASLSFLIFELHCAKQNLIVLPLFTLYQYFVSEICRDPVKCIEGRILKIKVLTDLGFFAEAFYELCLLNHGEKIPWKVSAGYRLTGKMMSFQKFNASKPLLSNANLQVLEDIFNWSLPPAVVSMCDQQIMNKLTLAKMHFIICLSATINSIPEKVMKTIYSMDNTSLKKPSRIMESSIKINAEVTRHQEQSSIIVQLVRCKDELNMAMLKGILLTETEERLNSLVANIQNEYDGKISRCSAVELETVIEAKLQLAVISQQRHQTAFSAALAFSAIRLLQDANIFRKNVPHIQLNIQRTERRDSYVDSSQLPYNVIAREHMNIRLWLRCRMALVTALTAQIRGIGDMKENDMTDCSSLINEVIIEAEAFSDIETQAEVLVQAVILDLQERRPVTGIKLLLQDIINLLQEKPFISPMASLTLVKSMLLLADILTTQTDEDTEHHSSAIEQLNLLILAHKFTIKQIFTLGAPIKQCIEDPALTSPVVPLKNNYLPVINLLAKVKMRIAHALAQEVASTSKRENPLEWLPALSHLETALDLCRTSATRELDLEAEILFQKGKVERQIIKAGNNKSPRAAETLLDAIKVSQRNDQNFGLIRKSYLEIALLYFHLATYKEETSQTVKIEASKTKTTPKEHLASTERIPHITEVYRTQAWIAVRAATQVSEAMLASQLLIGKKGVKGHQVKAITEQNIPEFASMDLLASYKDFLSDGYEVRYKNLMASSIKNEEIIQSEKDQIESEGTECNISKDIQRKQTITWVHLIRYHNYLIRHNMSPLFAVSKSTPGSFSMEDALFTSVFDTGVVLRLAEMHFFLKKHLYIYSTCCLEDFPKQLQFLEKPGASLTAVPVKMSLEILGAPHKMSTVRIASTVSFCTDTDIQLDNRATIAPNKELCIQWYIPSLEKSTKDMETMVLLLYAYNTKYVKIVSIKTFNSANVFSGHLWIPLTRIISLHEKLSNVKQQIEILVQSSTSFPLQEQSERQRRMPSTKSKINLAKVELDEKIEEMLKQCLCEIKALLCVVPDLSSPLTEIPFDVTLQSIQNLEKMFDPANGCVIAGGSLFSWVISLLP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHH2</accession>
    <name>A0A452GHH2_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GHH2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001204.1">
        <property type="protein sequence ID" value="ENSGAGP00000001070.1"/>
        <property type="gene ID" value="ENSGAGG00000000862.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:1990589">
        <property type="term" value="C:ATF4-CREB1 transcription factor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001630">
        <property type="entry name" value="Leuzip_CREB"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45879:SF4">
        <property type="entry name" value="CAMP-RESPONSIVE ELEMENT MODULATOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45879">
        <property type="entry name" value="CYCLIC AMP RESPONSE ELEMENT-BINDING PROTEIN B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="1" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="1">
        <location>
            <begin position="20"/>
            <end position="38"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001070.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="147" mass="15726" checksum="9BD1DAA15B48FCF6" modified="2019-05-08" version="1">MLSFNLSVLLLFEVSVEIHRALYLNMIIIILAGFYSWIKCWKRIFSCNSSTVAFWSNCTRPGSDSSHTAISHSVTTNAGCSVAIAQGGAIQISNPASDGVQGLQTLTMTNSGAPQPGATIVQYAAQTPEGTQQFFVPGSQVVVQVRL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHH6</accession>
    <name>A0A452GHH6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">protein-serine/threonine phosphatase</fullName>
            <ecNumber evidence="3">3.1.3.16</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29035"/>
        </cofactor>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the PP2C family.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.16" evidence="3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001197.1">
        <property type="protein sequence ID" value="ENSGAGP00000001063.1"/>
        <property type="gene ID" value="ENSGAGG00000000310.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005654">
        <property type="term" value="C:nucleoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004722">
        <property type="term" value="F:protein serine/threonine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="CDD" id="cd00143">
        <property type="entry name" value="PP2Cc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.60.40.10:FF:000023">
        <property type="entry name" value="Protein phosphatase, Mg2+/Mn2+-dependent, 1G"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.60.40.10">
        <property type="entry name" value="PPM-type phosphatase domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015655">
        <property type="entry name" value="PP2C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036457">
        <property type="entry name" value="PPM-type-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001932">
        <property type="entry name" value="PPM-type_phosphatase-like_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13832:SF803">
        <property type="entry name" value="PROTEIN PHOSPHATASE 1G"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13832">
        <property type="entry name" value="PROTEIN PHOSPHATASE 2C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00481">
        <property type="entry name" value="PP2C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00332">
        <property type="entry name" value="PP2Cc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81606">
        <property type="entry name" value="PP2C-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51746">
        <property type="entry name" value="PPM_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="6" id="KW-0460">Magnesium</keyword>
    <keyword evidence="8" id="KW-0464">Manganese</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="PPM-type phosphatase" evidence="10">
        <location>
            <begin position="125"/>
            <end position="376"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="126"/>
            <end position="198"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="386"/>
            <end position="409"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="9">
        <location>
            <begin position="149"/>
            <end position="177"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="178"/>
            <end position="193"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="388"/>
            <end position="397"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001936"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006702"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00013081"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023211"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS51746"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001063.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="409" mass="44446" checksum="2CC0BC0CED2BB40A" modified="2019-05-08" version="1">GPAFPSGLSAQLALCPGPLGGARFSSNLCSLSGAEVALYCAKYLPEIIKEQKAYKEGKLQKALEDAFLAIDVKLTTEEVIKELAQIAGRPQDFREEKEKVADEDDDDLLGHRLPLAELILSILPEGLVPSEPGTPECSEDEDGHSSETAEIEEDEGDTEEIEEDEGDTEEIEEDEDKEEKIMLPRMERKKEPGSDSGTTAVVALIRGKQLIVANAGDSRCVVSEXEGGKAVDMSHDHKPEDELELARINNAGGKVTMDGRVNGRLNLSRAIGDHFYKQNKNLPPEEQMISALPDIKVLTINEEHDFMVIACDGIWNVMSSQEVVDFIQAKISQKDENGDLRPLSSIVEELLDQCLAPQASGDGIGCDNMTCIIICFNPRSTGKRKLAAAEAQAAGEENGSDSSKKAKRD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHH7</accession>
    <name>A0A452GHH7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Tubby-like protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 6">Belongs to the TUB family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHH7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001209.1">
        <property type="protein sequence ID" value="ENSGAGP00000001075.1"/>
        <property type="gene ID" value="ENSGAGG00000000861.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005929">
        <property type="term" value="C:cilium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061512">
        <property type="term" value="P:protein localization to cilium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.20.90.10:FF:000001">
        <property type="entry name" value="Tubby-like protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.20.90.10">
        <property type="entry name" value="Tubby Protein, Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025659">
        <property type="entry name" value="Tubby-like_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000007">
        <property type="entry name" value="Tubby_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018066">
        <property type="entry name" value="Tubby_C_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005398">
        <property type="entry name" value="Tubby_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16517">
        <property type="entry name" value="TUBBY-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16517:SF138">
        <property type="entry name" value="TUBBY-RELATED PROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01167">
        <property type="entry name" value="Tub"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16322">
        <property type="entry name" value="Tub_N"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01573">
        <property type="entry name" value="SUPERTUBBY"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54518">
        <property type="entry name" value="Tubby C-terminal domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01200">
        <property type="entry name" value="TUB_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01201">
        <property type="entry name" value="TUB_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0964">Secreted</keyword>
    <feature type="domain" description="Tubby N-terminal" evidence="9">
        <location>
            <begin position="1"/>
            <end position="38"/>
        </location>
    </feature>
    <feature type="domain" description="Tubby N-terminal" evidence="9">
        <location>
            <begin position="39"/>
            <end position="148"/>
        </location>
    </feature>
    <feature type="domain" description="Tubby C-terminal" evidence="8">
        <location>
            <begin position="169"/>
            <end position="412"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1"/>
            <end position="29"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="97"/>
            <end position="152"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="7">
        <location>
            <begin position="105"/>
            <end position="114"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="122"/>
            <end position="149"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007129"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="RuleBase" id="RU361125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF01167"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF16322"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001075.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="417" mass="46674" checksum="6176E56E2F9F51FF" modified="2019-05-08" version="1">MVQPNPEAKLRRSKPKGCEEQTPLVESHTPFHSDVVLHGIDGPAAFLKPEVQDFGTKLQILSVGSSTTENDADQDNDEDTLIDTAAKPDLQEILQKRGISGSLNFDEDTEEEEEAGKRSRSHSPYSESERPSSATSQKSATDTGASCTASPEADAQLGEVENLEEFALRPAPRGITVKCRITRDKKGMDRGLFPTYYMHLERDDNRKTFLLAGRKRKKSKTSNYLISIDPTDLSREGESFIGKLRSNLMGTKFTVYDHGVSPGKAQGLIEKAHTRQELAAICYETNVLGFKGPRKMSVIIPGMNMNHKRIPIRPRNEHESLLSKWQNKNLENLIELHNKAPVWNDDTQSYVLNFHGRVTQASVKNFQIVHENDPDYIVMQFGRVAEDVFTLDYNYPLCALQAFAIGLSSFDSKLACE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHI2</accession>
    <name>A0A452GHI2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">SAM domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="5">May regulate the disassembly of focal adhesions. Did not bind receptor-like tyrosine phosphatases type 2A.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the liprin family. Liprin-beta subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHI2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001199.1">
        <property type="protein sequence ID" value="ENSGAGP00000001064.1"/>
        <property type="gene ID" value="ENSGAGG00000000781.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048786">
        <property type="term" value="C:presynaptic active zone"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007528">
        <property type="term" value="P:neuromuscular junction development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd09563">
        <property type="entry name" value="SAM_liprin-beta1_2_repeat1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd09566">
        <property type="entry name" value="SAM_liprin-beta1_2_repeat2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd09569">
        <property type="entry name" value="SAM_liprin-beta1_2_repeat3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.50:FF:000005">
        <property type="entry name" value="Liprin-beta-1 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.50:FF:000007">
        <property type="entry name" value="Liprin-beta-1 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.50:FF:000017">
        <property type="entry name" value="Liprin-beta-1 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.50">
        <property type="entry name" value="Transcription Factor, Ets-1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058914">
        <property type="entry name" value="LIPB1/2_CC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037617">
        <property type="entry name" value="LIPB1/2_SAM_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037618">
        <property type="entry name" value="LIPB1/2_SAM_2nd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037619">
        <property type="entry name" value="LIPB1/2_SAM_3rd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029515">
        <property type="entry name" value="Liprin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001660">
        <property type="entry name" value="SAM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013761">
        <property type="entry name" value="SAM/pointed_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12587">
        <property type="entry name" value="LAR INTERACTING PROTEIN LIP -RELATED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12587:SF16">
        <property type="entry name" value="LIPRIN-BETA-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26022">
        <property type="entry name" value="CC_Liprin_beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00536">
        <property type="entry name" value="SAM_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07647">
        <property type="entry name" value="SAM_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00454">
        <property type="entry name" value="SAM"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47769">
        <property type="entry name" value="SAM/Pointed domain"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50105">
        <property type="entry name" value="SAM_DOMAIN"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 6" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="SAM" evidence="8">
        <location>
            <begin position="674"/>
            <end position="738"/>
        </location>
    </feature>
    <feature type="domain" description="SAM" evidence="8">
        <location>
            <begin position="746"/>
            <end position="809"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="430"/>
            <end position="613"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="626"/>
            <end position="661"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="99"/>
            <end position="350"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="458"/>
            <end position="472"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="489"/>
            <end position="501"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="576"/>
            <end position="586"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007547"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00060046"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50105"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001064.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1038" mass="116310" checksum="105B816EB18D950D" modified="2019-05-08" version="1">MMSDASDMLAAALEQMDGIIAGSKALEYSNGIFDCQSPTSPFMGSLRALHLVEDLRGLLEMMEADEKEGLRCQVPDSTAEALIEWLQSQMTNGHISGNGDIYQERLARLENDKESLVLQVSVLTDQVEAQGEKIRDLEFCLEEHREKLNATEEMLQQELLSRTSLETQKLNLMAEISNLKLKLTAVEKDRLDYEDRYGNTEGLMQEISELRLRVGEMDNERLQYEKKLKSTKSLMAKLSSMKIKVGQMQYEKQRMEQKCQILKDELASLKEKLEQKEAEVRKLQEKLVCKMKGEGVEITDRDENCKKKLKDKNIEVQKMKKAVESLMAANEEKDRKIEELRQSLNRYKKVQDMVILAQGKKGKEGECEEFLNSGSVSAVLLDMQSLTDQEKSPSPVPVISFPSHEEFNTTVDEENSLQIHTSILQVSIPAFPPAPESSEAATEKVNAQPKPEVASEVSEGTTPCSSVETQPCDSPLMSALQKSSSLGNLKKEASEVDKEPVQKPAETKAPMEGNKCGTLPPKSPSHSGTGDEDSFGTRKARSSFGRGFFKIKSNKRTASAPNLDRSRSASAPTLAETEKGSADHLDLAGLPPRPKETDSGQMSPSSPDSKKKARGIKKLFGRLKRSQSTTFNPDDMSEAEFKRGGTRATAGPRLGWSRDLGQSNSDLDMPFAKWTKEQVCNWLQGQGLGSYMNHGKHWILSGQTLLQASQQDLEKELGIKHPLHRKKLQLALQALGSEEENNHGKLDYNWVTRWLDDIGLPQYKTQFDEGRVDGRMLHYMSVDDLLSLKVVSVLHHLSIKRAIQVLRINNFEPNCLRRRPSDENNITPAEVTQWTNHRVMEWLRSVDLAEYAPNLRGSGVHGGLMVLEPRFNVETMAQLLNIPPNKTLLRRHLATHFNLLVGQDAQHQKREAMESPDYVLLTATAKVKPKKLAFSNFGSLRKKKQDDVEEYVCPMELGQASGSGSKKGFKAGLDIRVYDDDDLDRLEQMEDSEGTVRQIGAFSEGINNLTHMLKEDEMFKDFATCSPSASITDEDSNV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHI4</accession>
    <name>A0A452GHI4_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000001167.1">
        <property type="protein sequence ID" value="ENSGAGP00000001035.1"/>
        <property type="gene ID" value="ENSGAGG00000000775.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060271">
        <property type="term" value="P:cilium assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR33487">
        <property type="entry name" value="CILIA- AND FLAGELLA-ASSOCIATED PROTEIN 54"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR33487:SF1">
        <property type="entry name" value="CILIA- AND FLAGELLA-ASSOCIATED PROTEIN 54"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="378"/>
            <end position="400"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="700"/>
            <end position="730"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="1">
        <location>
            <begin position="386"/>
            <end position="396"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="702"/>
            <end position="721"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001035.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2343" mass="265301" checksum="8F874CE9B36B2720" modified="2019-05-08" version="1">MNKIKKNKLSKAIFLMQKAILMFPKGSASSSQKHLLEEALALIQKVEAEQNALYSSLKQAMTSKKNRIPPPPLLLSRSHCSMTFKPAPFTSDVQIPADGKSLLEIKGLDPNEKYIFAVAAYSSDGKLIGDAIGETTKPILAYPPLSATTARAYLTQIAYQTDNYTLAKKAFSPLWDYFVLPPSTPSADPAVISASNNLTISQNRLCLDAVSQTSPILLHLFLRSIFIINEINVKEGALFCDSICSKEILYNGQVARLAECERMVVAIELSNWLNDANYALQSVVQCYGLLAPIIYHKIASVPVVQILIKCLTVLQEVPSSTLQRKQTGCYESIQHMIACSSFYTAQVLRSWKEYELALIIINYGKKLLDSSQTSHGQSKFSSIEEMQIDPETEDESPYSKKKSQVAAVEKAAENLAALESNLLKLTRPGTGPEITGQEDPFFLYPIISCWTTKNAYREVMKFRKKPRFLEFFVQVLHKALNEEKFQRIVEWTGSVKDFLRRRNNHLLGIKGTLAKEVSELSVYEDAKRHTTAVVEFQKSVETSALRKPRKEVMSPFRGRESLRQYFLRHSSLGKSPEAQRKQRDEIEKVAFHTLVVLLNPIVSNYLKRKRFHQVCLEEMPWRSQMNIYLAITHFNLFKKRLEEQYNIGVRSLQHLNSYDVLDPEIFSLNNSGIIVVRGGTDDNIGTLSHLLIKRIQSGSSRKITDPDKSSERSTKFSEADTPRSQMTNDTELSITLSAREHERPMSNAILLEHFTKTFLHLRRAMVLAHRGGHWTLLQNTCRELWNHSQEALSIAKHSHSLQGLFPITRNFLISTIWLPFYLASDAILDMIIELQTSNSLKIFDPKGDFCVPSCVGGIADEDGGSNLTFCCPLDDISVVDLRWICGLVLRTIELMYQMKKWEALVHIAMHFNTVTHERYAEQVTPLLVYAQRQLLERIQQFRGPDSPQPHFIKYQGDNANKINCRNFIGKQLRIASTFNEVTVSGHHFVPEMPDTYTDGSQAQALVCVPLDVTDTLKCFRETLEKSKYHSRALRHSRKLLSLFLAHTQETVGRVPNQIFTNGKVEFSVGAELTFLPKPPDLSQEKYNFPSMVESRPLPESQLSVVVSSYEKTIEILQINNQRDLKVQALHELGNLHFYAGKKRSAFKYWCQALDETLKIADVLDNWQELEGFSKNVPDDFTDGSSKHFSEKCIYQAGIWGCLHAAVLAAKIAQYILTSSVRLRTKCCIFSAILFKTLFKVSIPHPTADCDFAQYEIHRLIPGIDLFSDRYRADVSTVVASLSFLIFELHCAKQNLIVLPLFTLYQYFVSEICRDPVKCIEGRILKIKVLTDLGFFAEAFYELCLLNHGEKIPWKVSAGYRLTGKMMSFQKFNASKPLLSNANLQVLEDIFNWSLPPAVVSMCDQQIMNKLTLAKMHFIICLSATINSIPEKVMKTIYSMDNTSLKKPSRIMESSIKINAEVTRHQEQSSIIVQLVRCKDELNMAMLKGILLTETEERLNSLVANIQNEYDGKISRCSAVELETVIEAKLQLAVISQQRHQTAFSAALAFSAIRLLQDANIFRKNVPHIQLNIQRTERRDSYVDSSQLPYNVIAREHMNIRLWLRCRMALVTALTAQIRGIGDMKENDMTDCSSLINEVIIEAEAFSDIETQAEVLVQAVILDLQERRPVTGIKLLLQDIINLLQEKPFISPMASLTLVKSMLLLADILTTQTDEDTEHHSSAIEQLNLLILAHKFTIKQIFTLGAPIKQCIEDPALTSPVVPLKNNYLPVINLLAKVKMRIAHALAQEVASTSKRENPLEWLPALSHLETALDLCRTSATRELDLEAEILFQKGKVERQIIKAGNNKSPRAAETLLDAIKVSQRNDQNFGLIRKSYLEIALLYFHLATYKEETSQTVKIEASKTKTTPKEHLASTERIPHITEVYRTQAWIAVRAATQVSEAMLASQLLIGKKGVKGHQVKAITEQNIPEFASMDLLASYKDFLSDGYEVRYKNLMASSIKNEEIIQSEKDQIESEGTECNISKDIQRKQTITWVHLIRYHNYLIRHNMSPLFAVSKSTPGSFSMEDALFTSVFDTGVVLRLAEMHFFLKKHLYIYSTCCLEDFPKQLQFLEKPGASLTAVPVKMSLEILGAPHKMSTVRIASTVSFCTDTDIQLDNRATIAPNKELCIQWYIPSLEKSTKDMETMVLLLYAYNTKYVKIVSIKTFNSANVFSGHLWIPLTRIISLHEKLSNVKQQIEILVQSSTSFPLQEQSERQRRMPSTKSKINLAKVELDEKIEEMLKQCLCEIKALLCVVPDLSSPLTEIPFDVTLQSIQNLEKMFDPANGCVIAGGSLFSWVISLLP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHI5</accession>
    <name>A0A452GHI5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">U3 small nucleolar RNA-associated protein NOL7 C-terminal domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GHI5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001207.1">
        <property type="protein sequence ID" value="ENSGAGP00000001073.1"/>
        <property type="gene ID" value="ENSGAGG00000000863.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005730">
        <property type="term" value="C:nucleolus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012579">
        <property type="entry name" value="NOL7_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR32337">
        <property type="entry name" value="NUCLEOLAR PROTEIN 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR32337:SF2">
        <property type="entry name" value="NUCLEOLAR PROTEIN 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08157">
        <property type="entry name" value="NUC129"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="U3 small nucleolar RNA-associated protein NOL7 C-terminal" evidence="2">
        <location>
            <begin position="142"/>
            <end position="204"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="46"/>
            <end position="68"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="98"/>
            <end position="132"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="100"/>
            <end position="113"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="1">
        <location>
            <begin position="114"/>
            <end position="132"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF08157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001073.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="213" mass="24583" checksum="D3AE8CCAD207D39F" modified="2019-05-08" version="1">MAMTRGRLAMLAKEAAALEPPMAMAAAASEDEEEDEVPEEVTFERARAAAETERRQAGETARRDRELLKEKRRRREELFKEQKKRKLLPENVLKELTSRPRINNNNLSATTKQGKNRKKNEVRNRRRNNKGKKRVVRLKGRYMAVRLKDQDLTSQHQKVAKDFIQSHLYGPGTNRTTANQFFSFANKKSPVKKAAVQFVNKSWGKCNESEIVC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHI9</accession>
    <name>A0A452GHI9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">TEA domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 6">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHI9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001171.1">
        <property type="protein sequence ID" value="ENSGAGP00000001040.1"/>
        <property type="gene ID" value="ENSGAGG00000000816.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005667">
        <property type="term" value="C:transcription regulator complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048568">
        <property type="term" value="P:embryonic organ development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035329">
        <property type="term" value="P:hippo signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="2.70.50.80:FF:000001">
        <property type="entry name" value="Transcriptional enhancer factor TEF-1, putative"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.70.50.80">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.20.40">
        <property type="entry name" value="TEA/ATTS domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000818">
        <property type="entry name" value="TEA/ATTS_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038096">
        <property type="entry name" value="TEA/ATTS_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050937">
        <property type="entry name" value="TEC1_TEAD_TF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016361">
        <property type="entry name" value="TEF_metazoa"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041086">
        <property type="entry name" value="YBD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11834">
        <property type="entry name" value="TRANSCRIPTIONAL ENHANCER FACTOR TEF RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11834:SF2">
        <property type="entry name" value="TRANSCRIPTIONAL ENHANCER FACTOR TEF-3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01285">
        <property type="entry name" value="TEA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17725">
        <property type="entry name" value="YBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF002603">
        <property type="entry name" value="TEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00065">
        <property type="entry name" value="TEADOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00426">
        <property type="entry name" value="TEA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51088">
        <property type="entry name" value="TEA_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3 6" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="5 6" id="KW-0539">Nucleus</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 6" id="KW-0804">Transcription</keyword>
    <keyword evidence="2 6" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="TEA" evidence="8">
        <location>
            <begin position="1"/>
            <end position="39"/>
        </location>
    </feature>
    <feature type="DNA-binding region" description="TEA" evidence="7">
        <location>
            <begin position="1"/>
            <end position="39"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PIRNR" id="PIRNR002603"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00505"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51088"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001040.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="360" mass="40999" checksum="F6165468426587AE" modified="2019-05-08" version="1">MYCRNELIARYIKLRTGKTRTRKQVSSHIQVLARRKAREIQAKLKDQAAKDKAMQSMATMSSAQIISATAFHSKMVLPGLPRPAYPAVSGFWQGALPGQAGSSHDVKPFSQQTYAVQPPLPLPGFESPAGLSPSPSAPPWQGRRVASSKLWMLEFSAFLEQQQDQDTYNKHLFVHIGQSSPSYSDPYLEAVDIRQIYDKFPEKKGGLKELFEKGPPNAFFLVKFWADLNTNIEDESRSFYGVSSQYESPENMIITCSTKVCSFGKQVVEKVETEYAHYENGHYSYRIHRSPLCEYMINFIHKLKHLPEKYMMNSVLENFTILQVVTNRDTQETLLCIAYVFEVSASEHGAQHHIYRLVKD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHJ6</accession>
    <name>A0A452GHJ6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">TEA domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 6">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHJ6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001176.1">
        <property type="protein sequence ID" value="ENSGAGP00000001044.1"/>
        <property type="gene ID" value="ENSGAGG00000000816.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005667">
        <property type="term" value="C:transcription regulator complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048568">
        <property type="term" value="P:embryonic organ development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035329">
        <property type="term" value="P:hippo signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="2.70.50.80:FF:000003">
        <property type="entry name" value="Scalloped, isoform D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.70.50.80">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.20.40">
        <property type="entry name" value="TEA/ATTS domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000818">
        <property type="entry name" value="TEA/ATTS_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038096">
        <property type="entry name" value="TEA/ATTS_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050937">
        <property type="entry name" value="TEC1_TEAD_TF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016361">
        <property type="entry name" value="TEF_metazoa"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041086">
        <property type="entry name" value="YBD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11834">
        <property type="entry name" value="TRANSCRIPTIONAL ENHANCER FACTOR TEF RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11834:SF2">
        <property type="entry name" value="TRANSCRIPTIONAL ENHANCER FACTOR TEF-3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01285">
        <property type="entry name" value="TEA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17725">
        <property type="entry name" value="YBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF002603">
        <property type="entry name" value="TEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00065">
        <property type="entry name" value="TEADOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00426">
        <property type="entry name" value="TEA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00554">
        <property type="entry name" value="TEA_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51088">
        <property type="entry name" value="TEA_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3 6" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="5 6" id="KW-0539">Nucleus</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 6" id="KW-0804">Transcription</keyword>
    <keyword evidence="2 6" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="TEA" evidence="8">
        <location>
            <begin position="1"/>
            <end position="39"/>
        </location>
    </feature>
    <feature type="DNA-binding region" description="TEA" evidence="7">
        <location>
            <begin position="1"/>
            <end position="39"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PIRNR" id="PIRNR002603"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00505"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51088"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001044.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="388" mass="43387" checksum="E211E30E80BE4E6B" modified="2019-05-08" version="1">MYGRNELIARYIKLRTGKTRTRKQVSSHIQVLARRKAREIQAKLKDQAAKDKAMQSMATMSSAQIISATAFHSKMVLPGLPRPAYPAVSGFWQGALPGQAGSSHDVKPFSQQTYAVQPPLPLPGFESPAGLSPSPSAPPWQGRRVASSKLWMLEFSAFLEQQQDQDTYNKHLFVHIGQSSPSYSDPYLEAVDIRQIYDKFPEKKGGLKELFEKGPPNAFFLVKFWADLNTNIEDESRSFYGVSSQYESPENMIITCSTKVCSFGKQVVEKVETEYAHYENGHYSYRIHRSPLCEYMINFIHKLKHLPEKYMMNSVLENFTILQGFTVVGTPPPPITDSCISPRPIKPFTSCTQGTRGVGANGAICPWPRRGPTSPSLVTVRVFSGILA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHJ8</accession>
    <name>A0A452GHJ8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Olduvai domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Golgi apparatus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000001230.1">
        <property type="protein sequence ID" value="ENSGAGP00000001095.1"/>
        <property type="gene ID" value="ENSGAGG00000000835.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060090">
        <property type="term" value="F:molecular adaptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007098">
        <property type="term" value="P:centrosome cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090063">
        <property type="term" value="P:positive regulation of microtubule nucleation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903358">
        <property type="term" value="P:regulation of Golgi organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056273">
        <property type="entry name" value="CDK5RAP2_MYOME_CC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012943">
        <property type="entry name" value="Cnn_1N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052593">
        <property type="entry name" value="MT-associated_AKAP9-binding"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010630">
        <property type="entry name" value="Olduvai_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46501">
        <property type="entry name" value="MYOMEGALIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46501:SF2">
        <property type="entry name" value="MYOMEGALIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23246">
        <property type="entry name" value="CC_CDK5RAP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07989">
        <property type="entry name" value="Cnn_1N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01148">
        <property type="entry name" value="DUF1220"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51316">
        <property type="entry name" value="ODV"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="5" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Olduvai" evidence="9">
        <location>
            <begin position="1643"/>
            <end position="1734"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="23"/>
            <end position="46"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1084"/>
            <end position="1131"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1403"/>
            <end position="1424"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1484"/>
            <end position="1510"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1678"/>
            <end position="1784"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="2023"/>
            <end position="2064"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="2249"/>
            <end position="2297"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="165"/>
            <end position="228"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="254"/>
            <end position="310"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="396"/>
            <end position="451"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="510"/>
            <end position="676"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="705"/>
            <end position="790"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="892"/>
            <end position="1034"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="1152"/>
            <end position="1179"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="1345"/>
            <end position="1372"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="1453"/>
            <end position="1480"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="1842"/>
            <end position="1922"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="2190"/>
            <end position="2224"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="1114"/>
            <end position="1123"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="1405"/>
            <end position="1416"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="1686"/>
            <end position="1703"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="1727"/>
            <end position="1738"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="1742"/>
            <end position="1762"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="1769"/>
            <end position="1784"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="2023"/>
            <end position="2032"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="2043"/>
            <end position="2061"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="2274"/>
            <end position="2286"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004555"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51316"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001095.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2316" mass="259954" checksum="174BAE65C8AA22B1" modified="2019-05-08" version="1">MLADAGLEYAGLSRGRWLRSRLPRRRTSGARLPEQPMRSPVGAGTVQAPASHGPGCCCCCSREAPGLDPAEMCTPARGDYCDGDGKEMLPGASHVTAVLCDAPEQRNLDISDEDSLRQTPTSGLFETAHHPVPAQTLPGLEQSPLMQTHVLSDLEMGPVVQTQTLRDFEKHLNDLKKENFSLKLRIYFLEERMQQKYEASREDVYKRNIELKVEVESLKQELQERQQTLDKAWAAAENLTSCNEAEFQRRYEERQQEKEHVSELLENKIQLLQEEARLAKSEAERTAALVEAEKGRCLELTKKLKELTKEEEKRGAQVLRDDHCAALAQKDKHIEELIQELSAREQLIELLATEKQNLLQRLEEPREIEVQNLPEDHQQLTQCDPVASGRVSESHAAELQDKIQQYDATNKLLQEKLNEMNFELKSIQEASQRQDRTIQSLNEALKSKESETEELYHVIEGQNETMGKLRDMLHRSQLGHLQTSEGPSPSQQQQHMALLDLQNTLFCTQLEMQKLKRAQWQKEHQLAEARRTTQLLEATVQEEQQLKEVSWKHNQELRGVVQQVQAELQGKVQELWALEGEKCCEIRAQEQRVQRLSQRLAHKEQLLQESWELLQRRQSLEKNPVVTDTMLEKLQQRIKHRDAALERAIDEKFCAVEEKEQELQQLHISVREREHDLERLRSVLSSNEATIHSMESLLKAKTLELEQVTATCQNLQWLKEEVEAKSNCWQKEQEGIIQQLQASLHDRNKEVEELTATLLCKLGPGQSEIAEELCLRLQRKERMLQDLLSDRNRQAGEHETELRELLQAMSIREQRSRVTAEKMVQALAEKSDEVQFLRQRLVGNEPGKNPVADGRLLLQKDKQPFQELLPGGCGNATATRISKGEDSKYRMERGLLETAAELEKELINAREELELMARKERESRLELSSLQSVVTAQEEELQVQASDIESLTRNIQSKEDLIKDLQMQLVDPEEIPAVERLTQEVLVLREKVATAESRGQEATGNRQQQLLLMLEGLVAEKSRLNEALQAERQLYSSLVKFHAHPDSPERGQTLQAELEGAQALRGRLEESLGRSMEHLRRLETLGAFGGQPGRDDAKDGSTDSIEGEAPPRTVNHQTSQGLPANSPEAKAAQPDLLTLAPLERDSSLQAELLHVRAKNQQLLEQKAKVEGELWELKALMEEAGFSSLSSIRKVLLSLCLETAELKEQLGGSPLSEGWEDKAEARAALCTQNSKLQSSETVVTLLKEQLVLNCPEGESKFPPQLVTGMATEIDRLWTEMGVASGKRPALEGQLQEDPAKRPCPRSQAVELGPAQAPRGMSQQMEACPAGSWQRLVEPRVGPSGQLVQCRQRNQELQDKLAVSEATVRAQAEQLEQYRTLLSEPLVQQDNKQVQVDLQDLGYETCGRSENEADREETPSPECEEPDVFSKAGLMEELSRPGPWGGLCWGQSEDVAVLQQHVRELQLQLQNSNKALQRLQRRTRSFSSTSDYASGAERLHKPVPSATDEDEGWQSDGVGAFCPPALQASRDLEWLVHRVSLLEAQLHGRWAGGVLPEELRSAVWPGKYDSLIQAQARELSHLRQKMREGRSVCHLLTQHLRDTVKSFEELLRGTDIDYYMGQSFREHLAQGGQLAERLSSKLSSRDRSDVEDKTGHEFVALRLSKELREKERMIETLQAKLQERCETPSSSRALSESPRSNSSASFLSDGPEACSDGDATSEYSQLQGDRSEQPSRRLTDVTHPAALASNPSPAAAPHQASAESIRGYPTGPSSQHPPMDSGQTRTGADLLEGHLAEIRSLRQRLEESICINDRLREQLEKRLATTTKGNGYPTNIYIQGLESASSLSSETQALREENQSLQLQFAHLSRELEQLRETTRCRLQKAETENRTLAQELAEQRQSGRQLREERLALQENNHRLECTVPLLQQQCQENQQLCQALCAELRIHETLSSSSQAALSACSGDVHRRPPSSSELGALLAEVRTLRRKLLRGIQVNSALRQQLARQQPEGSAAPLFAAHQATPNWQPFQDSSPSPPVRDVGMSSPAPLSPPALLSGPAPLAHQLKEPPVDDALVRRSAPAPAGDAPSSSSASKGGCQVIGHVDDYYTLKQLILEGKTLVHQMASLLRPALAMPSLEPRGTKALDRGSVQQLLSTASALHQLLEQCSSLLTTFWRGALPAAPSPAQPQTAEQAMKDELLALRARLAEQENLLQRAAERLQDTCRLKDNMEHFIVSHLTQTHAVLRKARTNLEKSKPQGSSVKSPPFIGAGEISVGLSQHPSHVIPTNQEHRRKKSCPDTLKWQEDQSWPCFVHLPSS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHK1</accession>
    <name>A0A452GHK1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="19">E3 ubiquitin-protein ligase UHRF</fullName>
            <ecNumber evidence="19">2.3.2.27</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="19">RING-type E3 ubiquitin transferase UHRF</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="19">Ubiquitin-like PHD and RING finger domain-containing protein</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="19">Ubiquitin-like-containing PHD and RING finger domains protein</fullName>
        </alternativeName>
    </protein>
    <organism evidence="25 26">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="26" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="25" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="19">Multi domain E3 ubiquitin ligase that also plays a role in DNA methylation and histone modifications.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="1 27">
            <text>S-ubiquitinyl-[E2 ubiquitin-conjugating enzyme]-L-cysteine + [acceptor protein]-L-lysine = [E2 ubiquitin-conjugating enzyme]-L-cysteine + N(6)-ubiquitinyl-[acceptor protein]-L-lysine.</text>
            <dbReference type="EC" id="2.3.2.27"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="2 19">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="18 19">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="domain">
        <text evidence="19">The YDG domain mediates the interaction with histone H3.</text>
    </comment>
    <comment type="domain">
        <text evidence="19">The tudor-like regions specifically recognize and bind histone H3 unmethylated at 'Arg-2' (H3R2me0), while the PHD-type zinc finger specifically recognizes and binds histone H3 trimethylated at 'Lys-9' (H3K9me3).</text>
    </comment>
    <dbReference type="EC" id="2.3.2.27" evidence="19"/>
    <dbReference type="AlphaFoldDB" id="A0A452GHK1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001182.1">
        <property type="protein sequence ID" value="ENSGAGP00000001050.1"/>
        <property type="gene ID" value="ENSGAGG00000000852.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000001214.1">
        <property type="protein sequence ID" value="ENSGAGP00000001079.1"/>
        <property type="gene ID" value="ENSGAGG00000000852.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00143"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042393">
        <property type="term" value="F:histone binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061630">
        <property type="term" value="F:ubiquitin protein ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0044027">
        <property type="term" value="P:negative regulation of gene expression via chromosomal CpG island methylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016567">
        <property type="term" value="P:protein ubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006950">
        <property type="term" value="P:response to stress"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd15616">
        <property type="entry name" value="PHD_UHRF1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd16769">
        <property type="entry name" value="RING-HC_UHRF1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd20455">
        <property type="entry name" value="Tudor_UHRF1_rpt1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd20457">
        <property type="entry name" value="Tudor_UHRF1_rpt2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd17122">
        <property type="entry name" value="Ubl_UHRF1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.280.10:FF:000001">
        <property type="entry name" value="E3 ubiquitin-protein ligase UHRF1 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.140:FF:000068">
        <property type="entry name" value="E3 ubiquitin-protein ligase UHRF1 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.20.90:FF:000143">
        <property type="entry name" value="E3 ubiquitin-protein ligase UHRF1 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.1150:FF:000001">
        <property type="entry name" value="E3 ubiquitin-protein ligase UHRF2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.40.10:FF:000066">
        <property type="entry name" value="E3 ubiquitin-protein ligase UHRF2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.1150">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.20.90">
        <property type="entry name" value="Phosphatidylinositol 3-kinase Catalytic Subunit, Chain A, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.280.10">
        <property type="entry name" value="SRA-YDG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015947">
        <property type="entry name" value="PUA-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036987">
        <property type="entry name" value="SRA-YDG_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003105">
        <property type="entry name" value="SRA_YDG"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021991">
        <property type="entry name" value="TTD_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000626">
        <property type="entry name" value="Ubiquitin-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029071">
        <property type="entry name" value="Ubiquitin-like_domsf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019956">
        <property type="entry name" value="Ubiquitin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047406">
        <property type="entry name" value="Ubl_UHRF1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045134">
        <property type="entry name" value="UHRF1/2-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011011">
        <property type="entry name" value="Znf_FYVE_PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001965">
        <property type="entry name" value="Znf_PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019787">
        <property type="entry name" value="Znf_PHD-finger"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001841">
        <property type="entry name" value="Znf_RING"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017907">
        <property type="entry name" value="Znf_RING_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14140">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE UHRF-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14140:SF2">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE UHRF1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00628">
        <property type="entry name" value="PHD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02182">
        <property type="entry name" value="SAD_SRA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12148">
        <property type="entry name" value="TTD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00240">
        <property type="entry name" value="ubiquitin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00348">
        <property type="entry name" value="UBIQUITIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00249">
        <property type="entry name" value="PHD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00184">
        <property type="entry name" value="RING"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00466">
        <property type="entry name" value="SRA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00213">
        <property type="entry name" value="UBQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57903">
        <property type="entry name" value="FYVE/PHD zinc finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF88697">
        <property type="entry name" value="PUA domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57850">
        <property type="entry name" value="RING/U-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54236">
        <property type="entry name" value="Ubiquitin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50053">
        <property type="entry name" value="UBIQUITIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51015">
        <property type="entry name" value="YDG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50016">
        <property type="entry name" value="ZF_PHD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00518">
        <property type="entry name" value="ZF_RING_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50089">
        <property type="entry name" value="ZF_RING_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="16" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="11" id="KW-0156">Chromatin regulator</keyword>
    <keyword evidence="13 19" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="6 19" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="15 18 19" id="KW-0539">Nucleus</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="26" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0678">Repressor</keyword>
    <keyword evidence="14" id="KW-0804">Transcription</keyword>
    <keyword evidence="12" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="5 19" id="KW-0808">Transferase</keyword>
    <keyword evidence="9 19" id="KW-0833">Ubl conjugation pathway</keyword>
    <keyword evidence="10 19" id="KW-0862">Zinc</keyword>
    <keyword evidence="8 17 19" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="Ubiquitin-like" evidence="22">
        <location>
            <begin position="1"/>
            <end position="80"/>
        </location>
    </feature>
    <feature type="domain" description="PHD-type" evidence="21">
        <location>
            <begin position="327"/>
            <end position="378"/>
        </location>
    </feature>
    <feature type="domain" description="RING-type" evidence="23">
        <location>
            <begin position="330"/>
            <end position="376"/>
        </location>
    </feature>
    <feature type="domain" description="YDG" evidence="24">
        <location>
            <begin position="431"/>
            <end position="594"/>
        </location>
    </feature>
    <feature type="domain" description="RING-type" evidence="23">
        <location>
            <begin position="721"/>
            <end position="760"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="86"/>
            <end position="116"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="629"/>
            <end position="675"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="20">
        <location>
            <begin position="96"/>
            <end position="114"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="20">
        <location>
            <begin position="629"/>
            <end position="640"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022491"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00358"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="RuleBase" id="RU369101"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50016"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS50053"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS50089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS51015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001050.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="790" mass="88955" checksum="38F4ACF7D4A63D14" modified="2019-05-08" version="1">MWIQVRTMDGKETHRVDSLSKLTKVEELRLRIHEVFGDGVEPDRQRLFYRGKQMEDGHSLFDYNVGLNDIVQLLVKQSPAMLPTVNKEKDLELSDTDSGCGSGQSESDKNSNNGEGAIELECQPSTVAQADWVDPGFGLYKINEFVDARDMNMGAWFEAQIVNVTRKEKAANESSDSDTESELQSTAAEEDILYHVKYEDYPENGVVELSSKDVRTRARTVLKWHQLEVGEVVMVNYNPDEPKERGFWYDAEILRKRETRMNKEMYAKILLGDAGDSLNDCRIIFVDEIYKIEEPGSVSPASTSPLKRQSGPVCKHCKDNPNKTCRMCACHVCGGKQDPDKQLMCDECDMAFHIYCLNPPLSNIPDDEDWYCPECRNDASEVVLAGEKLKESKKKAKMASANSSSQRDWGKGMACVGRTRECTIVPSNHYGPIPGVPVGTMWKFRVQVSESGVHRPHVAGIHGRSNDGAYSLVLAGGYEDDIDHGNSFTYTGSGGRDLSGNKRTAEQSCDQKLTNMNRALALNCSAPINDKDGAEAKDWRAGKPVRVVRNVKGGKHSKYAPREGNRYDGIYKVVKYWPETGKSGFLVWRYLLRRDDEEPAPWTKEGKDRMKKLGLTMQYPEGYLEAVANKDKEKENKRGDEECETPGKGKRKRKSAGDEIKLPASPRGTPKKTKVEPYKLTSQQKALIKDDESNEKLWNEVLEALKDGPKFLNKVEETFLCICCQEVVFRPVTTVCQHNVCKDCLDRSFRADVYSCPACRYDLGKNYTMQVNETLQALLTQLFPGYGNGR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHK2</accession>
    <name>A0A452GHK2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">ELK domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
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                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
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</entry>
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                <person name="Henen B.T."/>
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                <person name="Murphy R.W."/>
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        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
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<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
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        <citation type="submission" date="2025-08" db="UniProtKB">
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        <scope>IDENTIFICATION</scope>
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        <property type="entry name" value="Polyadenylate-binding protein"/>
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    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Nucleotide-bd_a/b_plait_sf"/>
    </dbReference>
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        <property type="entry name" value="PABP-dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002004">
        <property type="entry name" value="PABP_HYD_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034364">
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    </dbReference>
    <dbReference type="InterPro" id="IPR035979">
        <property type="entry name" value="RBD_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045305">
        <property type="entry name" value="RRM2_I_PABPs"/>
    </dbReference>
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        <property type="entry name" value="RRM_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003954">
        <property type="entry name" value="RRM_euk-type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24012">
        <property type="entry name" value="RNA BINDING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00658">
        <property type="entry name" value="MLLE"/>
        <property type="match status" value="1"/>
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    <dbReference type="Pfam" id="PF00076">
        <property type="entry name" value="RRM_1"/>
        <property type="match status" value="4"/>
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        <property type="entry name" value="PolyA"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00361">
        <property type="entry name" value="RRM_1"/>
        <property type="match status" value="2"/>
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    <dbReference type="SUPFAM" id="SSF63570">
        <property type="entry name" value="PABC (PABP) domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54928">
        <property type="entry name" value="RNA-binding domain, RBD"/>
        <property type="match status" value="2"/>
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        <property type="match status" value="1"/>
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        <property type="match status" value="4"/>
    </dbReference>
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    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="5 6" id="KW-0694">RNA-binding</keyword>
    <feature type="domain" description="RRM" evidence="7">
        <location>
            <begin position="11"/>
            <end position="89"/>
        </location>
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    <feature type="domain" description="RRM" evidence="7">
        <location>
            <begin position="99"/>
            <end position="175"/>
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            <begin position="191"/>
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            <begin position="259"/>
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    </feature>
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        <location>
            <begin position="510"/>
            <end position="587"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
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    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008557"/>
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    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
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    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00176"/>
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    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50102"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51309"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001087.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="604" mass="67116" checksum="1BC624C3AF8F7169" modified="2019-05-08" version="1">MNPSAPSYPMASLYVGDLHPDVTEAMLYEKFSPAGPILSIRVCRDMITRRSLGYAYVNFQQPADAERALDTMNFDVIKGKPVRIMWSQRDPSLRKSGVGNIFIKNLDKSIDNKALYDTFSAFGNILSCKVVCDENGSKGYGFVHFETQEAAERAIEKMNGMLLNDRKVFVGRFKSRKEREAELGARAKEFTNVYIKNFGEDMDDERLKELFGKFGPALSVKVMTDESGKSKGFGFVSFERHEDAQKAVDEMNGKELNGKQIYVGRAQKKVERQTELKRKFEQMKQDRITRYQVMMEGGRSKGFGFVCFSSPEEATKAVTEMNGRIVATKPLYVALAQRKEERQAHLTNQYMQRMASVRAVPNPVINPYQPAPPSGYFMAALPPTQNRAAYYPTSQLAQLRPSPRWTAQGARPHPFQNMPGAIRPAAPRPPFSTMRPASSQVPRVMSTQRVANTSTQTMGPRPAAAAAAATPAVRTVPQYKYAAGVRNPQQHLNTQPQVAMQQPAVHVQGQEPLTASMLASAPPQEQKQMLGERLFPLIQAMHPTLAGKITGMLLEIDNSELLHMLESPESLRSKVDEAVAVLQAHQAKEAAQKAVNNPTGVPSV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GI02</accession>
    <name>A0A452GI02_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Protein FAM196B</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GI02"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001410.1">
        <property type="protein sequence ID" value="ENSGAGP00000001257.1"/>
        <property type="gene ID" value="ENSGAGG00000001004.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029337">
        <property type="entry name" value="INSYN2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR28682">
        <property type="entry name" value="INHIBITORY SYNAPTIC FACTOR 2A-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR28682:SF6">
        <property type="entry name" value="MUCIN-5AC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15265">
        <property type="entry name" value="FAM196"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1"/>
            <end position="86"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="202"/>
            <end position="222"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="272"/>
            <end position="360"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="430"/>
            <end position="462"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="513"/>
            <end position="547"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="51"/>
            <end position="62"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="70"/>
            <end position="81"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="2">
        <location>
            <begin position="284"/>
            <end position="304"/>
        </location>
    </feature>
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        <location>
            <begin position="307"/>
            <end position="321"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="336"/>
            <end position="351"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="430"/>
            <end position="449"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001257.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="5">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="590" mass="62783" checksum="B7F046251D535A3C" modified="2019-05-08" version="1">MVSKETSPCLARSNEGSQEKAMTVRSVLLNRDSPNIETRLKHRRNRPQQVRFKDLLDGKPGEEVSPPQEPTATPDTNTPCASTPARESPELVASCASDAQWGWHEAKPCSLTLPVPSKVCLSTAIQTSPSLQQQFPASRVRSKSACDFAQQEVLAAAWEPSKSQTGCWGPARAPASTRASGDLPHCILTSLSAHNHLPASPLAQHTSREGQPGPTPHATLCHGLTPPPRAPCPSYPGTIRCTAANYTSQQKGCPPAPSPGLCSSQPPAYTRNCGSPKDSHDAPAPCPLRPPSVPCGQPGPPAEPRPQSRSQSEQDLPSQPAHNHMAGSHGLGQPAQCPSQRGPTAALQQQHSSSWASPSCLPSNHIPSTVLPPPSLPSTTATMLKKTPATLSHQAPSSGWAQSVAARPSVCSQLKATQTECGLMNLEEPQQLTSTSSQGHPPRTQQFTRTPGAPWSNEPLGEPCLTAKQLETLHQVQDLLQLVVAAKGQVGLSKGDEHFLTSRGHVGVGPGQMGDLQSQLQSLEGVLETSQQTIKMLLDVIQDLEKKEAQRDGRHSYRTGQDIANCGTCRDCACIIYRSTSVHGKEATKP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GI04</accession>
    <name>A0A452GI04_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">TFIIS central domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000001355.1">
        <property type="protein sequence ID" value="ENSGAGP00000001207.1"/>
        <property type="gene ID" value="ENSGAGG00000000965.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006351">
        <property type="term" value="P:DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd21540">
        <property type="entry name" value="SPOC_SPOCD1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.472.30">
        <property type="entry name" value="Transcription elongation factor S-II, central domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012921">
        <property type="entry name" value="SPOC_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003618">
        <property type="entry name" value="TFIIS_cen_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036575">
        <property type="entry name" value="TFIIS_cen_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11477:SF18">
        <property type="entry name" value="SPOC DOMAIN-CONTAINING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11477">
        <property type="entry name" value="TRANSCRIPTION FACTOR S-II ZINC FINGER DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07744">
        <property type="entry name" value="SPOC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07500">
        <property type="entry name" value="TFIIS_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00510">
        <property type="entry name" value="TFS2M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46942">
        <property type="entry name" value="Elongation factor TFIIS domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51321">
        <property type="entry name" value="TFIIS_CENTRAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="TFIIS central" evidence="2">
        <location>
            <begin position="945"/>
            <end position="1061"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="17"/>
            <end position="42"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="386"/>
            <end position="420"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="651"/>
            <end position="676"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="908"/>
            <end position="931"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1137"/>
            <end position="1160"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1631"/>
            <end position="1658"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="17"/>
            <end position="29"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="400"/>
            <end position="415"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="1631"/>
            <end position="1656"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="PROSITE" id="PS51321"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001207.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1670" mass="185694" checksum="1F43D52FB85CAAE8" modified="2019-05-08" version="1">MEHQKFHGEQSKFCLNVNSKNMETSNTEAGSLGSGGPDRDINPEEEIFKKRAVVAGEAFEIPKINCVDNDDNDKGSRTVQSETWFGVGDDLDENCIKELSSTLKIPALEALGENEKSHSMKSYLKVQNQEESVTSQGARLTGVDAVKAEFPLNNEGTKLKELLSQMDFSGNLSNSWDQGTHSKSPIIPDLKEEKELFSFHLPACSEQVTGKRVGETYASDHVSNEEQSEQSTVTLNVVNQNKVSSESEVRVSGDGSSYSRSPYVKEIKAKETDFREKMIVSGGPSESPKQNCDNDERGSRTILIKPCNIGLKDLKDELKEKCKDSLISLDIETQSLKMKNESKASFSDVFQVPLHKNNVKKQNKIKIQNIKAPLENEKLNNTEKKEIKSHGEAVTSRGSDSMDTKIHHNSEETQGKRSFTQMDSPDLSDMIGDGIFSRLSLVEKEYKRSTINLSNVEEEGSRTKRLQSDSRPLSSMIPEVEKEKSREVITPVAKFRKFWNNPVTQLGKEEISYSYLVMNPNRKGFNSRLSERLFTSNISQSGRIILAPENTLQTKDLQGIPVEGTSTYLSASKTENVFGEGKSHNRSPVLPDQSDGIEDIHVTVCEPSCISPPAVEKFGTNQKGELSEMKNQMPRVEGLVQYFLEESAEEVSQGQLDVEEQKSLNQGNFKSNDESDDSSWFLMEIAEDLDNVDVANVPDGQNKAVEQETRGEHLNPSKVTLQSESMEVPVSGRHKAECYSCSNDINKIKHNDIKKSTAFCCLESDRDGQSSCVGKVAAPDSAKETCLIGKMSGPVVELLGVPSQACKPRRLQRLVELEDSSDDEMCVCCQPPKKMSRLVLGVSKSKACKPCKMRRVEEMEDLSEDVYVDFQVTQEIKDSDYHPTLYKREMKQLLKPYAAHVRWQKPLNSDYSNEKKRRGRTPASKEASASVHGLQLSEEQSRIKVIDSLRGMLQKRLEESPDLDVHEDTILRLANNVEKEIFNLFLCVDQRYKNKYRSLLFNLKSPKNKLLFHQVVLGEVSPQSLVQMNSLELAPKELAEWRAKESKHVLEVIEKQQREPPRCCPTKLTHKGEIEIHREVDEDFTLEDLYGSALCMDKKSISQPAALSKRDTTEQHRSHLLDLDCLICTGRMAPDGESGFNPLQSKPTSKKKAEDSSRRLHSTGVYSYKERKHLEKDRPASVDVLKKSPRPKKQVRDTVLWEGFIQMFSIKQFAAKAYPVSGYGTCLIQALPELIQSRGCILPEDVWDYLESIWPAEAKEMSMIRFCPAMTKDFNTYNMLYSYLNNKQRYGIVNSNQMEMFMVPLPAFQPVPAKFHPLGGPGLDASHSCLLLGLILPKKVSRDPSTSSMGPSLLRETKRKTVTFKNNLVTKCFSPPLDSSGSTKLAEPFQQVPSRAFTGKRLSRNEYPPAVFHSQEPCNNMFALNTAVGEVIHGGTCHAASSSHCQNRKEETNQSMQDCAVHSLSSFSWMGGQSYSTAQESLGRQQNMIGGHTEFQADAGPAMTQNWNHGPLSSCSLCSHECGDNVALDVNMLDIQQLSSMLYPGVASSGGVSQVPSVAHLVSNEPPTASDLQNHAATGDSNPTIPVGEALSLIQHLEALVQLNSQIQNQTLFPLPSLDLSAILQATTPGESQIVGGSTSPPAQQESGVDPQSQLTLPPFCGSIYPPSGC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GI05</accession>
    <name>A0A452GI05_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="1">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="1 2">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="2" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="1" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="1" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GI05"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001415.1">
        <property type="protein sequence ID" value="ENSGAGP00000001262.1"/>
        <property type="gene ID" value="ENSGAGG00000001002.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036282">
        <property type="entry name" value="Glutathione-S-Trfase_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029063">
        <property type="entry name" value="SAM-dependent_MTases_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13369">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13369:SF0">
        <property type="entry name" value="GLUTATHIONE S-TRANSFERASE C-TERMINAL DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47616">
        <property type="entry name" value="GST C-terminal domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53335">
        <property type="entry name" value="S-adenosyl-L-methionine-dependent methyltransferases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000313" key="1">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001262.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="588" mass="65817" checksum="663A702272BEDDD8" modified="2019-05-08" version="1">MKGGVTEEHLYLAYSHQFEDCIFPLHTSVSLFLLSYCDCKLFKVFLVPASDSTDDQLFIKAQPGDLEVHFISRCQLPLVVQSCCLPAIVEENGKFCRAGLSVVLRHIIQKTYEVDPSKKLVLELLGFKKTCLKACAEVSQWTRLCEISIPQAVESLLNASPDQCQVIPPEVLLLEKKLGEPVRVHNDDKIRRQKLQQQKAGYKTAAPALTKRATKNLSPEEGTIVEIKDQEPASLELKVAFSKLVVQEIPAATNREPFHIRKTKTSDLPPLEHVFAEGLYFTLADIVLLPCIHQFLVFIKKHNKLLDLPLISSWYHRVQEVPGVKSAAAKCNMLFLQFPDLLSSPDEQLRDFYTIPDDLEEEHEDYQFIGGPRPTMTKLMESGIEAKFSLHPCPRWTLDWNSLPAAVSPGEGKMSSDRALRKQQQLNNLVSMVTKLSKPGDVIVDFCSGGGHVGIVLAHMLPSCQVALHACGVATDMVIEHCIKAHAAFVICPCCYGFIQNTVKFTFPRSHQFKEVLSYKEHMILCRFADQTAVQLPPKRRLIGKHCMGLVDLDRAWAAEGCNYSVQVMSMEPESCSPKNNLVVGVPI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GI08</accession>
    <name>A0A452GI08_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Peptidase S1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GI08"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001416.1">
        <property type="protein sequence ID" value="ENSGAGP00000001263.1"/>
        <property type="gene ID" value="ENSGAGG00000000827.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00190">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.10:FF:000005">
        <property type="entry name" value="Serine protease 37"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.10.10">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009003">
        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043504">
        <property type="entry name" value="Peptidase_S1_PA_chymotrypsin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001314">
        <property type="entry name" value="Peptidase_S1A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001254">
        <property type="entry name" value="Trypsin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018114">
        <property type="entry name" value="TRYPSIN_HIS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271:SF81">
        <property type="entry name" value="GRANZYME B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271">
        <property type="entry name" value="KALLIKREIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00089">
        <property type="entry name" value="Trypsin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00722">
        <property type="entry name" value="CHYMOTRYPSIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00020">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50494">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50240">
        <property type="entry name" value="TRYPSIN_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00134">
        <property type="entry name" value="TRYPSIN_HIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0732">Signal</keyword>
    <keyword evidence="2" id="KW-0865">Zymogen</keyword>
    <feature type="domain" description="Peptidase S1" evidence="4">
        <location>
            <begin position="14"/>
            <end position="245"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023145"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001263.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="246" mass="27089" checksum="4DFEF6EEA4006D63" modified="2019-05-08" version="1">MQSLSCLVSSEGDIIGGREARPHSRPYMAFVETKMGPESTTECGGFLVREDFVLTAAHCDEEGSGDAPTVVILGAHDLNRKERTQQEIQVRRKIPHEDFDRTTLENDIMLLQLELRAEWNNAVKPIRLPPANFLLKPEATCSVAGWGRTGVHMVRPATKLQEVTLKMADDKQCELFFDSVHVSQHGPSSKHGTGYFKIGPGDTGGPLVCNGFAQSIVSYGKDSGLPSAVYTRMSAFIPWLRKKMGT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYJ7</accession>
    <name>A0A452GYJ7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Protein TOPAZ1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="7">Testis- and ovary-specific PAZ domain-containing protein 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">Important for normal spermatogenesis and male fertility. Specifically required for progression to the post-meiotic stages of spermatocyte development. Seems to be necessary for normal expression levels of a number of testis-expressed gene transcripts, although its role in this process is unclear.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytosol</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000008316.1">
        <property type="protein sequence ID" value="ENSGAGP00000007190.1"/>
        <property type="gene ID" value="ENSGAGG00000005743.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048137">
        <property type="term" value="P:spermatocyte division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038952">
        <property type="entry name" value="TOPAZ1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029435">
        <property type="entry name" value="TOPAZ1_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR35671">
        <property type="entry name" value="PROTEIN TOPAZ1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR35671:SF1">
        <property type="entry name" value="PROTEIN TOPAZ1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14669">
        <property type="entry name" value="Asp_Glu_race_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="5" id="KW-0221">Differentiation</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0744">Spermatogenesis</keyword>
    <feature type="domain" description="Protein TOPAZ1" evidence="9">
        <location>
            <begin position="1113"/>
            <end position="1288"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="27"/>
            <end position="53"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00002132"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004514"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00016464"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022871"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00031943"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF14669"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007190.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1518" mass="172801" checksum="86DDF77685EBD1D6" modified="2019-05-08" version="1">KQLETDLNISKDYNIKKPLVAYSESLQEAKSDGTRSKAPVCPGLKRGSAVQGNRRRNRITKIRCLQNLDVKVTENKVTKSRLVKSKDEKCREIPNYHNNLPSTVERNPVVKLYDCRYLNTFVKSSVDATNCSYKIGGGFLHVAQGKRETSCPDSVMGQSDMNSKLTVMQMSTQKSIPFDKEGICQNRKNCQNENSKCLRERKWNIDRKPRKRMKLFEKSSVQNVLPVMDNKYCEYELHTEDKTATTDSLAILKLNSNDTTSSSSFDQKSDPHLEPNIQEAQNGCTASFNRKMHLKPCATEGSFEKRLELSEIAKGDKSSFDSESIEQQDRSNTLNSPEVVTLVSDIISDYYNDASCQKKERIVSNEKFKSNVKKLHLYSCQRIVPMSGKNVWPRESCARTSEWFHKNHVSVLEGKTSTSGSQECSYENNTEAVREMVLTDNSRQLDLSMLEAESVKESVCKITMDSNIECLTSVRTPESSLLDICGTLRVTCNENLESSVDSSPIASKSEAVHEAQVILNLAVKQKQDDKNKGALTKKNRMMALQDGISEDDKTSKMIVNHKTSMIKKTVTIPNLMKMLNPGNLSNFKIPLRKNKTESRSVDYIRSLDKETYSPLECLHRSAATSARQNRAEENSSVVVSKQQPISEEADISVIPLKENADRMISKDFGSGGSENLSNEINILSERVSTYQCASLERELESSALEDVIDSSVTASPSRTLDYSSKTVDCSISLEMHDDDDKKKSKMDLAQSKIKNFPDVLKAYEDDILVIDVIQDDPDLFGTTDEGELALTMNHVTEKHCNISTTSEEKQDVKPKCLVLPETKDTRNYFSEYLSFALILEKHLYYFVKRVFLLLFSNDSKYIEFVSSELQLGLQLPSLQINAFQEDTVLKPWINGILVCNSVTCVSHLIQQNLGMIELPRKYCRFYFSTLRGCERVECWFWHVPEQGDEKICMAILKTYININESSLLQRSVQIFMNYYREATPGVHFDSQVLNDLLISLIKCCLLKEVFQLLNVAVMIKMLPAIDVFLKVFEHVASMNIRDAVASLIDIFCKLFDAGMIFELEHFNCIIKFLHQLQVSSQEINVILNMKSRFQAKQFKKNWLRDLNLAIAEIQHCKEKSDWAKLGTLYVNMRIGCENFGDLQKLTLYIADILTRDSKEERPGVPFCEFAEAVIKNPQHNEADRDFLGRFGISVMYSYHTVQQWSKGRKVLDKLHELQIHFTSLKGIIGPERLASRCQIVNAAAEIFLKSGSLDGATWVLRESEWIINTPLWPCDKMDVLNRHNLLCAIACEYLTKNLYRQAFEILQNLPGFQYCCDTLDVSQYSLLFNKLLDACIESKGLGISSCVTDFMLSKNIPIDFTLIRGLITALGRSCLWLKARIYYKSALSMGCYPPLEGNLYRKLLLIPSYMSEIEMLLAIEIFLVSNASNIQSPCATNQTLQVVLKRSEEQKVRSKDDYQAAVERLIHAARISNPKLFLKHMTMNVNMEQVYSLEHNSALKWLKENMKWAGKVWLFSSR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYK0</accession>
    <name>A0A452GYK0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">VLRF1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="domain">
        <text evidence="14">The VLRF1 domain mediates binding to the 60S ribosomal subunit.</text>
    </comment>
    <comment type="similarity">
        <text evidence="2 14">Belongs to the ANKZF1/VMS1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYK0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008369.1">
        <property type="protein sequence ID" value="ENSGAGP00000007238.1"/>
        <property type="gene ID" value="ENSGAGG00000005817.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004519">
        <property type="term" value="F:endonuclease activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016787">
        <property type="term" value="F:hydrolase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036503">
        <property type="term" value="P:ERAD pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047139">
        <property type="entry name" value="ANKZ1/VMS1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041540">
        <property type="entry name" value="VATC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041175">
        <property type="entry name" value="VLRF1/Vms1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16036">
        <property type="entry name" value="ANKYRIN REPEAT AND ZINC FINGER DOMAIN-CONTAINING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16036:SF2">
        <property type="entry name" value="TRNA ENDONUCLEASE ANKZF1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00023">
        <property type="entry name" value="Ank"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18826">
        <property type="entry name" value="bVLRF1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18716">
        <property type="entry name" value="VATC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS52044">
        <property type="entry name" value="VLRF1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="11 13" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="12 15" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3 14" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7 14" id="KW-0255">Endonuclease</keyword>
    <keyword evidence="9 14" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="5" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="4 14" id="KW-0540">Nuclease</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="10" id="KW-0862">Zinc</keyword>
    <keyword evidence="8" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="VLRF1" evidence="17">
        <location>
            <begin position="199"/>
            <end position="342"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="13">
        <location>
            <begin position="549"/>
            <end position="581"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="107"/>
            <end position="155"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="373"/>
            <end position="474"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="501"/>
            <end position="536"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="15">
        <location>
            <begin position="627"/>
            <end position="667"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="16">
        <location>
            <begin position="414"/>
            <end position="425"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="16">
        <location>
            <begin position="426"/>
            <end position="439"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="443"/>
            <end position="457"/>
        </location>
    </feature>
    <feature type="active site" evidence="14">
        <location>
            <position position="242"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009262"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022759"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01389"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS52044"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007238.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="735" mass="80668" checksum="F470574305201EC5" modified="2019-05-08" version="1">MQPLEVRSVFEAAQDPTFLHGLSLVNCSSEDLPSTDCNAITQEKPVVADGEERAQGVPEISDRMCCSSCSRVFDSREEQTEHYRLDWHRFNLKQRLLGRQALTAEEFEEKTRAGDVSSISGSDSDDSDSGSESDLLPPREHTDPGSSQQSLRPGTRSAKVLFRNSQGQLLSAYRCVLGSKKGGGEQQAELVASLQSMGTGTCWVILMTGGGHFAGAVFRGPEVLDHKTFHRYTVRARRGTVQGVRDAQGQASMPKSAGASLRRYNEAALLKDIQDLLASWVQHLQGAQRIFLRVPCANRALLFSGRNPPLHKDDPRICSIPFSTRRATFREVLRVHAALATLQVYGKDTVVADLAGFPRKGWRKVPCARVEEPQQGDASFLSEEENEESQAGELEMVDVTLSTLHLRESEVTPKRSRKKRRKKDKVGRGARAEGPRGQEEPEGSSSQPGAETVTDPQEGSGAEPAPWRKGETPLGQLRDALFTACKTGDVGTLRRLLGVAESGAPPAGSEDGAMGQSQGHPGRVQSEGSEPNHAVPGLLSLLSEPIDGTGFTLLHVAAAAGKGAAVRLLLEAGADPALRDRQERPPYCVSADKSTRNVFRKFMVDHPAKYDYGRAKVPGPLTAEMEARRLEKQRAQKTQRKQRAQELREQQQRLAQEQEEKRRFAALPDREKRALAAERRLAAQLQDTGAALANTRRCWQCGESLLGRIPFHYHDFSFCSTACLQTHRRGRAGPP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYK1</accession>
    <name>A0A452GYK1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">CFA20 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GYK1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008438.1">
        <property type="protein sequence ID" value="ENSGAGP00000007297.1"/>
        <property type="gene ID" value="ENSGAGG00000005860.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040441">
        <property type="entry name" value="CFA20/CFAP20DC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007714">
        <property type="entry name" value="CFA20_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12458">
        <property type="entry name" value="ORF PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05018">
        <property type="entry name" value="CFA20_dom"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="CFA20" evidence="2">
        <location>
            <begin position="1"/>
            <end position="175"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="219"/>
            <end position="247"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="334"/>
            <end position="449"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="225"/>
            <end position="247"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="334"/>
            <end position="372"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="378"/>
            <end position="393"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="1">
        <location>
            <begin position="435"/>
            <end position="449"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF05018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007297.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="449" mass="50168" checksum="AB698D336E370BD9" modified="2019-05-08" version="1">MFRNEYQGGPFVEIFSAQGKNPGAKWKIFGSPSAIWKEFDKEVKGFVFVLEGSSQTNKIQLPKETRQTLGLIQRFLVLQIYVPLGQDFSTELLVTDLGNIKRRLYLSTVHKELSITPLHAKLPLFMIKRRIWCNLCIDLVAFTSEIFRGAVFQSLDGIIISANCKLRKIFTLKSKPQDTAEEDADDPTDTIPRTCQLNTDVPQVTQLLNMTKLRQPEIRFGGRPLTSSESDQLINRGQGSTHNSKNQDVSHIAFGSKVLGPPPPSGRKVNTRVSGEITKSLGSRSNRSCQQLTLEKGSESMRDTKRSGLCLSPFYKAFDQGDKENIHQTNQITSQNDDWIFPENSSDSFQIDSNGQSVATHNTSCSNLSLPPNTEVESESKQTGEHQISHKEIFTFSSRPRSAPHGKSPNMSPEGCMSPLDLNEDSDGFCRGAQMEDDFYGSDSNEEFL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYK2</accession>
    <name>A0A452GYK2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Actin gamma 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 8">Belongs to the actin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYK2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008256.1">
        <property type="protein sequence ID" value="ENSGAGP00000007133.1"/>
        <property type="gene ID" value="ENSGAGG00000005729.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.420.40:FF:000131">
        <property type="entry name" value="Actin, alpha skeletal muscle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.420.40:FF:000205">
        <property type="entry name" value="Actin, alpha skeletal muscle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.640.10:FF:000047">
        <property type="entry name" value="Actin, alpha skeletal muscle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.420.40:FF:000058">
        <property type="entry name" value="Putative actin-related protein 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.420.40">
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.640.10">
        <property type="entry name" value="Actin, Chain A, domain 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004000">
        <property type="entry name" value="Actin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004001">
        <property type="entry name" value="Actin_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043129">
        <property type="entry name" value="ATPase_NBD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11937">
        <property type="entry name" value="ACTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00022">
        <property type="entry name" value="Actin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00268">
        <property type="entry name" value="ACTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53067">
        <property type="entry name" value="Actin-like ATPase domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00432">
        <property type="entry name" value="ACTINS_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="4" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="6" id="KW-0558">Oxidation</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006752"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023097"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="RuleBase" id="RU000487"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007133.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="11">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="295" mass="32620" checksum="729D69E703FA1629" modified="2019-05-08" version="1">MEEEIAALVIDNGSGMCKAGFAGDDAPRAVFPSIVGRPRHQIMFETFNTPAMYVAIQAVLSLYASGRTTGIVMDSGDGVTHTVPIYEGYALPHAILRLDLAGRDLTDYLMKILTERGYSFTTTAEREIVRDIKEKLCYVALDFEQEMATAASSSSLEKSYELPDGQVITIGNERFRCPEALFQPSFLGMESCGIHETTFNSIMKCDVDIRKDLYANTVLSGGTTMYPGIADRMQKEITALAPSTMKIKIIAPPERKYSVWIGGSILASLSTFQQMWISKQEYDESGPSIVHRKCF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYK6</accession>
    <name>A0A452GYK6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">CFA20 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GYK6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008443.1">
        <property type="protein sequence ID" value="ENSGAGP00000007302.1"/>
        <property type="gene ID" value="ENSGAGG00000005860.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040441">
        <property type="entry name" value="CFA20/CFAP20DC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007714">
        <property type="entry name" value="CFA20_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12458">
        <property type="entry name" value="ORF PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05018">
        <property type="entry name" value="CFA20_dom"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="CFA20" evidence="2">
        <location>
            <begin position="1"/>
            <end position="175"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="338"/>
            <end position="424"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="338"/>
            <end position="373"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="379"/>
            <end position="394"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF05018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007302.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="485" mass="54261" checksum="8F419F59DC17EA52" modified="2019-05-08" version="1">MFRNEYQGGPFVEIFSAQGKNPGAKWKIFGSPSAIWKEFDKEVKGFVFVLEGSSQTNKIQLPKETRQTLGLIQRFLVLQIYVPLGQDFSTELLVTDLGNIKRRLYLSTVHKELSITPLHAKLPLFMIKRRIWCNLCIDLVAFTSEIFRGAVFQSLDGIIISANCKLRKIFTLKSKPQDTAEEDADDPTDTIPRTCQLNTDVPQVTQLLNMTKLRQPEIRFGGRPLTSSESVDQLINRGQGSTHNSKNQDVSHIAFGSKVLGPPPPSGRKVNTRVSGEITKSLGSRSNRSCQQLTLEKGSESMRDTKRSGLCLSPFYKAFDQGDKENIHQTNQITSQNDDWIFPENSSDSFQIDSNGQSVATHNTSCSNLSLPPNTEVESESKQTGEHQISHKEIFTFSSRPRSAPHGKSPNMSPEGCMSPLDLNEDSDGFCRGAQMEDDFYGSDSNEETGKEVQQSYVTARQMKLCKNTVPFLLGNYFTDANYNQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYK7</accession>
    <name>A0A452GYK7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Fibronectin type-II domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="3">Belongs to the sel-1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYK7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008448.1">
        <property type="protein sequence ID" value="ENSGAGP00000007307.1"/>
        <property type="gene ID" value="ENSGAGG00000004194.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031012">
        <property type="term" value="C:extracellular matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036503">
        <property type="term" value="P:ERAD pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00062">
        <property type="entry name" value="FN2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.10.10:FF:000001">
        <property type="entry name" value="Fibronectin 1a isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000208">
        <property type="entry name" value="Protein sel-1 homolog 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000200">
        <property type="entry name" value="protein sel-1 homolog 1 precursor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.10.10">
        <property type="entry name" value="Fibronectin, type II, collagen-binding"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.10">
        <property type="entry name" value="Tetratricopeptide repeat domain"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000562">
        <property type="entry name" value="FN_type2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036943">
        <property type="entry name" value="FN_type2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013806">
        <property type="entry name" value="Kringle-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006597">
        <property type="entry name" value="Sel1-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050767">
        <property type="entry name" value="Sel1_AlgK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011990">
        <property type="entry name" value="TPR-like_helical_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11102">
        <property type="entry name" value="SEL-1-LIKE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11102:SF147">
        <property type="entry name" value="SEL1L ADAPTOR SUBUNIT OF ERAD E3 UBIQUITIN LIGASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00040">
        <property type="entry name" value="fn2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08238">
        <property type="entry name" value="Sel1"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00013">
        <property type="entry name" value="FNTYPEII"/>
    </dbReference>
    <dbReference type="SMART" id="SM00059">
        <property type="entry name" value="FN2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00671">
        <property type="entry name" value="SEL1"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81901">
        <property type="entry name" value="HCP-like"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57440">
        <property type="entry name" value="Kringle-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00023">
        <property type="entry name" value="FN2_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51092">
        <property type="entry name" value="FN2_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 4" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="665"/>
            <end position="683"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-II" evidence="7">
        <location>
            <begin position="78"/>
            <end position="126"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="1"/>
            <end position="41"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="1"/>
            <end position="10"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="19"/>
            <end position="41"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="4">
        <location>
            <begin position="83"/>
            <end position="109"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="4">
        <location>
            <begin position="97"/>
            <end position="124"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00038101"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS51092"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007307.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="714" mass="80329" checksum="1E8EC58F06F39580" modified="2019-05-08" version="1">EQVGDRSTGTRVVAGQIFHESEESDSHTEDDESFRSPEEEKESMLEELNSLEMSNLLNKNLEDVEIQKPVLTAIGGTADSEPCHFPFLFLEKEYAECTADGREDGRLWCATTYDYKRDQKWGFCETEEQSNKRRQMQEAEDVYQTGMKIFNESSKKAQKKEAYQYLLKAADMNHTKAMEKVSYALLFGDYLKQNIQSAKELFEKLTEEGSPKGQTRVAAGYRYWGGIGVLQSCESALTHYRLVANHVASDISLTGGTVVQRIRLADEVENPGMASGMLEEDLIQYYQFLAEKGDVQAQVGLGQLHLHGGRGVEQNHQRAFEYFNQAANAGNSHAMAFLGKLYSEGSDIIPQSNETALQYFKKAADMGNPVGQSGLGMAYLYGRGVPVNYELALKYFQKAAEQGWVDGQLQLGSMYYNGIGVKRDYKQALKYFNLASQGGHILAFYNLAQMHATGTGVMRSCHTAVELFKNVCERGRWSERLMTAYNSYKDGDSNAAVVQYLLLAEQGYEVAQSNAAFILDQKEASIVGENETYPQALLHWNRAASQGYTVARIKLGDYHFYGFGTDVDYETAFIHYRLASEQQHSAQAMFNLGYMHEKGLGIKQDIHLAKRFYDMAAEASPDAQVPVFLALCKLGIVYSLQYVRETNIREIVSHLDMDQLLGPEWDLYLMTIIALLLGTVIAYRQRQHQAILRPVGLRPAVPQQEPPPEHQPPQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYK9</accession>
    <name>A0A452GYK9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="5">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the OXR1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYK9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008332.1">
        <property type="protein sequence ID" value="ENSGAGP00000007203.1"/>
        <property type="gene ID" value="ENSGAGG00000005780.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006979">
        <property type="term" value="P:response to oxidative stress"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00118">
        <property type="entry name" value="LysM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.350.10">
        <property type="entry name" value="LysM domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018392">
        <property type="entry name" value="LysM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036779">
        <property type="entry name" value="LysM_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006571">
        <property type="entry name" value="TLDc_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23354:SF68">
        <property type="entry name" value="NUCLEAR RECEPTOR COACTIVATOR 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23354">
        <property type="entry name" value="NUCLEOLAR PROTEIN 7/ESTROGEN RECEPTOR COACTIVATOR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01476">
        <property type="entry name" value="LysM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07534">
        <property type="entry name" value="TLD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00257">
        <property type="entry name" value="LysM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00584">
        <property type="entry name" value="TLDc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54106">
        <property type="entry name" value="LysM domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51782">
        <property type="entry name" value="LYSM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51886">
        <property type="entry name" value="TLDC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="LysM" evidence="3">
        <location>
            <begin position="125"/>
            <end position="168"/>
        </location>
    </feature>
    <feature type="domain" description="TLDc" evidence="4">
        <location>
            <begin position="751"/>
            <end position="912"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1"/>
            <end position="50"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="293"/>
            <end position="317"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="339"/>
            <end position="380"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="448"/>
            <end position="475"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="1"/>
            <end position="12"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="25"/>
            <end position="42"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="297"/>
            <end position="317"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="360"/>
            <end position="380"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="448"/>
            <end position="464"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00009540"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS51782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS51886"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007203.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="912" mass="102753" checksum="C0AC91B6E94B8927" modified="2019-05-08" version="1">METKEEKKERRQGYFARLKKKKQAKQNAETVTSNSRGTTASRTHVGKDDGSKVILEQISSEDNCKLAEEKEECVTEEEKKKKKYNQLKDIRRTELKRYYNTDDNQNKTNEKKEKKMVTQKPLGTIEYTAGSQDTLNSIALKFNITPNRLVELNRLFTHTIVPGQVLYVPDTDHYSSASRLASSSPGAPVSPSSSDAEYDKLPGVVGGVMIVTPNNIMFDPHKSDPLVIENGCEEYGLICPMEEVVSIALYNDISHMKIKDALPSDIPKDLCPLYRPGEWEDLSSEKDINPFSKFKSLSKEKRQQNGDDSKSMKPLDKEKSADFELLKPFDSTISMEAKSLEPPNGMASAEASGRSAVGAHTKEPSGEKAASDSKTKERSLIEGGDDDFIELQETASLTNTRLYRTDAVTESFSFDPKELGKARSQVTKSVSEVQDLLAFDSLGTKNDVEPKADLDSESCEKQDVMPEVNKCASSPTGKVETTLDIQKELDKNKLIEFYLSKDKDGSQSSENLHEAGIGEGENNKAAGVDLTLSCSESQVNKLLKVKSGDSSCIERKEPSLEVCLDAVEEKDLKESVKSSSVPSLDNVCQETQLDNNSEIRLWLLQRIQVPIEDMLPSKEEKSKTPPMFLCIKVGKPMRKSFVTQSTTIAQQYGKRRKQPEYWFAVPRERVDHLYTFFVQWSPEVYGKDAKEQGFVVVEKEELDMIDNFFSEPSTKSWEIITVEEAKRRKSICSYYEEDDDDSLPVLKHHSALLENMHIEQLARCLPARVQGYPWRLAYSTLEHGTSLKTLYRKSASLDSPVLLVIKDMDNQIFGAYATHPFRFSDHYYGTGETFLYTFSPNFKVFKWSGENTYFINGDIGSLELGGGGGRFGLWLDVDLYHGRSNSCSTFNNDILSKKEDFIIQDVEVWTFE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYL1</accession>
    <name>A0A452GYL1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Homeobox domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 6 7">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the NK-2 homeobox family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYL1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008453.1">
        <property type="protein sequence ID" value="ENSGAGP00000007312.1"/>
        <property type="gene ID" value="ENSGAGG00000005873.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00086">
        <property type="entry name" value="homeodomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.60:FF:000078">
        <property type="entry name" value="NK2 homeobox 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.60">
        <property type="entry name" value="Homeodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001356">
        <property type="entry name" value="HD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020479">
        <property type="entry name" value="HD_metazoa"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017970">
        <property type="entry name" value="Homeobox_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050394">
        <property type="entry name" value="Homeobox_NK-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009057">
        <property type="entry name" value="Homeodomain-like_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24340">
        <property type="entry name" value="HOMEOBOX PROTEIN NKX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24340:SF32">
        <property type="entry name" value="HOMEOBOX PROTEIN NKX-2.3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00046">
        <property type="entry name" value="Homeodomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00024">
        <property type="entry name" value="HOMEOBOX"/>
    </dbReference>
    <dbReference type="SMART" id="SM00389">
        <property type="entry name" value="HOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46689">
        <property type="entry name" value="Homeodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00027">
        <property type="entry name" value="HOMEOBOX_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50071">
        <property type="entry name" value="HOMEOBOX_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 6 7" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="4 6 7" id="KW-0371">Homeobox</keyword>
    <keyword evidence="5 6 7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Homeobox" evidence="9">
        <location>
            <begin position="69"/>
            <end position="129"/>
        </location>
    </feature>
    <feature type="DNA-binding region" description="Homeobox" evidence="6">
        <location>
            <begin position="71"/>
            <end position="130"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="24"/>
            <end position="75"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="58"/>
            <end position="67"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005661"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023155"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00108"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU000682"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50071"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007312.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="196" mass="21802" checksum="AF70EDB5D8D8EAE8" modified="2019-05-08" version="1">MLPSPVTSTPFSVKDILSLEQQDCPVGQAPLPRPGPALTLPLPHAERCALKQQPGAAEKPEEAERPKPRSRRKPRVLFSQAQVFELERRFKQQRYLSAPEREHLASSLKLTSTQVKIWFQNRRYKCKRQRQDKSLEMGPAPPPPRRVAVPVLVRDGKPCLGGSPGYGSPYNAPYSYNAPYNARPTGTGLTPVLGAE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYL2</accession>
    <name>A0A452GYL2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">tRNA (guanine(10)-N(2))-methyltransferase TRMT11</fullName>
            <ecNumber evidence="12">2.1.1.214</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="14">tRNA methyltransferase 11 homolog</fullName>
        </alternativeName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">Catalytic subunit of the TRMT11-TRM112 methyltransferase complex, that specifically mediates the S-adenosyl-L-methionine-dependent N(2)-methylation of guanosine nucleotide at position 10 (m2G10) in tRNAs. This is one of the major tRNA (guanine-N(2))-methyltransferases.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="9">
            <text>guanosine(10) in tRNA + S-adenosyl-L-methionine = N(2)-methylguanosine(10) in tRNA + S-adenosyl-L-homocysteine + H(+)</text>
            <dbReference type="Rhea" id="RHEA:43128"/>
            <dbReference type="Rhea" id="RHEA-COMP:10355"/>
            <dbReference type="Rhea" id="RHEA-COMP:10357"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57856"/>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
            <dbReference type="ChEBI" id="CHEBI:74269"/>
            <dbReference type="ChEBI" id="CHEBI:74481"/>
            <dbReference type="EC" id="2.1.1.214"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="9">
            <dbReference type="Rhea" id="RHEA:43129"/>
        </physiologicalReaction>
    </comment>
    <comment type="subunit">
        <text evidence="11">Part of the heterodimeric TRMT11-TRM112 methyltransferase complex; this complex forms an active tRNA methyltransferase, where TRMT112 acts as an activator of the catalytic subunit TRMT11.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="15">Belongs to the class I-like SAM-binding methyltransferase superfamily. TRM11 methyltransferase family.</text>
    </comment>
    <dbReference type="EC" id="2.1.1.214" evidence="12"/>
    <dbReference type="AlphaFoldDB" id="A0A452GYL2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008380.1">
        <property type="protein sequence ID" value="ENSGAGP00000007248.1"/>
        <property type="gene ID" value="ENSGAGG00000005813.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043527">
        <property type="term" value="C:tRNA methyltransferase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0160102">
        <property type="term" value="F:tRNA (guanine(10)-N2)-methyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000049">
        <property type="term" value="F:tRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032259">
        <property type="term" value="P:methylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008033">
        <property type="term" value="P:tRNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.150:FF:000069">
        <property type="entry name" value="tRNA (Guanine(10)-N2)-methyltransferase homolog isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.150">
        <property type="entry name" value="Vaccinia Virus protein VP39"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002052">
        <property type="entry name" value="DNA_methylase_N6_adenine_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000241">
        <property type="entry name" value="RlmKL-like_Mtase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029063">
        <property type="entry name" value="SAM-dependent_MTases_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016691">
        <property type="entry name" value="TRMT11"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR059073">
        <property type="entry name" value="TRMT11_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13370">
        <property type="entry name" value="RNA METHYLASE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13370:SF3">
        <property type="entry name" value="TRNA (GUANINE(10)-N2)-METHYLTRANSFERASE HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25904">
        <property type="entry name" value="Tmrp11_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01170">
        <property type="entry name" value="UPF0020"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF017259">
        <property type="entry name" value="tRNA_mtfrase_TRM11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00507">
        <property type="entry name" value="N12N6MTFRASE"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53335">
        <property type="entry name" value="S-adenosyl-L-methionine-dependent methyltransferases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00092">
        <property type="entry name" value="N6_MTASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51627">
        <property type="entry name" value="SAM_MT_TRM11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4 15" id="KW-0489">Methyltransferase</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8 15" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="6 15" id="KW-0949">S-adenosyl-L-methionine</keyword>
    <keyword evidence="16" id="KW-0732">Signal</keyword>
    <keyword evidence="5 15" id="KW-0808">Transferase</keyword>
    <keyword evidence="7 15" id="KW-0819">tRNA processing</keyword>
    <keyword evidence="3 15" id="KW-0820">tRNA-binding</keyword>
    <feature type="signal peptide" evidence="16">
        <location>
            <begin position="1"/>
            <end position="16"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5018989257" description="tRNA (guanine(10)-N(2))-methyltransferase TRMT11" evidence="16">
        <location>
            <begin position="17"/>
            <end position="446"/>
        </location>
    </feature>
    <feature type="domain" description="tRNA (guanine(10)-N(2))-methyltransferase TRMT11 N-terminal" evidence="18">
        <location>
            <begin position="22"/>
            <end position="193"/>
        </location>
    </feature>
    <feature type="domain" description="Ribosomal RNA large subunit methyltransferase K/L-like methyltransferase" evidence="17">
        <location>
            <begin position="203"/>
            <end position="334"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022555"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022603"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022691"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022694"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00050985"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00056270"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00065434"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00066937"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00067484"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00075308"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00959"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF01170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF25904"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007248.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="446" mass="51247" checksum="9E4DCEAFF911F100" modified="2019-05-08" version="1">MRGWAERFMASGGAMARGLLPSYLLLLAQENLEFRLPEIKSLLSLCGGQFSSQQEIRINSPFWILNIPSEEMARGLMKRTVCAKSVFELWGQGKSAEELYTSLKNYPMEKMVPYLQSNSTYKIHIHAFNKTLTQDKKIRRIDALEFLPFEGKVNLKNPEHIFWILEDYGMDPNNVPEDPLDLYFGRWIANGQRELIESYSVKRRHFIGNTSMDACLSFIMANHGRVKANDVVFDPFVGTGGLLISSARFGAYVCGTDIDYNTVHGLGKASRKNQKWRGPDENIRANLRQYGLEKYYLDALVSDASKPIWRKEMLFDAIITDPPYGIREATRRTGSQKEILKATERGTENHISISSNYHLSDIFFDLLNSAAEYLVMGGRLVYWLPVYKPEYTEEIIPRHPCLKLISNCEQMLSSHTSRRLITMEKVKEFEVKCSWVCCAKWRLSNI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYL5</accession>
    <name>A0A452GYL5_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="5">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the OXR1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYL5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008326.1">
        <property type="protein sequence ID" value="ENSGAGP00000007198.1"/>
        <property type="gene ID" value="ENSGAGG00000005780.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000008337.1">
        <property type="protein sequence ID" value="ENSGAGP00000007208.1"/>
        <property type="gene ID" value="ENSGAGG00000005780.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006979">
        <property type="term" value="P:response to oxidative stress"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00118">
        <property type="entry name" value="LysM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.350.10">
        <property type="entry name" value="LysM domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018392">
        <property type="entry name" value="LysM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036779">
        <property type="entry name" value="LysM_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006571">
        <property type="entry name" value="TLDc_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23354:SF68">
        <property type="entry name" value="NUCLEAR RECEPTOR COACTIVATOR 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23354">
        <property type="entry name" value="NUCLEOLAR PROTEIN 7/ESTROGEN RECEPTOR COACTIVATOR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01476">
        <property type="entry name" value="LysM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07534">
        <property type="entry name" value="TLD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00257">
        <property type="entry name" value="LysM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00584">
        <property type="entry name" value="TLDc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54106">
        <property type="entry name" value="LysM domain"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="LYSM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51886">
        <property type="entry name" value="TLDC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="LysM" evidence="3">
        <location>
            <begin position="125"/>
            <end position="168"/>
        </location>
    </feature>
    <feature type="domain" description="TLDc" evidence="4">
        <location>
            <begin position="793"/>
            <end position="954"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1"/>
            <end position="50"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="177"/>
            <end position="199"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="335"/>
            <end position="359"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="381"/>
            <end position="422"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="490"/>
            <end position="517"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="1"/>
            <end position="12"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="25"/>
            <end position="42"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="177"/>
            <end position="195"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="339"/>
            <end position="359"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="402"/>
            <end position="422"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="490"/>
            <end position="506"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00009540"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS51782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS51886"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007208.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="954" mass="107584" checksum="E3AB99BE838D8582" modified="2019-05-08" version="1">METKEEKKERRQGYFARLKKKKQAKQNAETVTSNSRGTTASRTHVGKDDGSKVILEQISSEDNCKLAEEKEECVTEEEKKKKKYNQLKDIRRTELKRYYNTDDNQNKTNEKKEKKMVTQKPLGTIEYTAGSQDTLNSIALKFNITPNRLVELNRLFTHTIVPGQVLYVPDTDHYSSASRLASSSPGAPVSPSSSDAEYDKLPDADLARKAFKPIERVLSSTSEEDEPIVVKFLKMNCRYFTDGKGVVGGVMIVTPNNIMFDPHKSDPLVIENGCEEYGLICPMEEVVSIALYNDISHMKIKDALPSDIPKDLCPLYRPGEWEDLSSEKDINPFSKFKSLSKEKRQQNGDDSKSMKPLDKEKSADFELLKPFDSTISMEAKSLEPPNGMASAEASGRSAVGAHTKEPSGEKAASDSKTKERSLIEGGDDDFIELQETASLTNTRLYRTDAVTESFSFDPKELGKARSQVTKSVSEVQDLLAFDSLGTKNDVEPKADLDSESCEKQDVMPEVNKCASSPTGKVETTLDIQKELDKNKLIEFYLSKDKDGSQSSENLHEAGIGEGENNKAAGVDLTLSCSESQVNKLLKVKSGDSSCIERKEPSLEVCLDAVEEKDLKESVKSSSVPSLDNVCQETQLDNNSEIRLWLLQRIQVPIEDMLPSKEEKSKTPPMFLCIKVGKPMRKSFVTQSTTIAQQYGKRRKQPEYWFAVPRERVDHLYTFFVQWSPEVYGKDAKEQGFVVVEKEELDMIDNFFSEPSTKSWEIITVEEAKRRKSICSYYEEDDDDSLPVLKHHSALLENMHIEQLARCLPARVQGYPWRLAYSTLEHGTSLKTLYRKSASLDSPVLLVIKDMDNQIFGAYATHPFRFSDHYYGTGETFLYTFSPNFKVFKWSGENTYFINGDIGSLELGGGGGRFGLWLDVDLYHGRSNSCSTFNNDILSKKEDFIIQDVEVWTFE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYL6</accession>
    <name>A0A452GYL6_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="6">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1 5">Belongs to the GDA1/CD39 NTPase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYL6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008458.1">
        <property type="protein sequence ID" value="ENSGAGP00000007317.1"/>
        <property type="gene ID" value="ENSGAGG00000005862.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004382">
        <property type="term" value="F:GDP phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017111">
        <property type="term" value="F:ribonucleoside triphosphate phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045134">
        <property type="term" value="F:UDP phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009134">
        <property type="term" value="P:nucleoside diphosphate catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.420.40:FF:000068">
        <property type="entry name" value="Ectonucleoside triphosphate diphosphohydrolase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.420.40">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.420.150">
        <property type="entry name" value="Exopolyphosphatase. Domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000407">
        <property type="entry name" value="GDA1_CD39_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11782">
        <property type="entry name" value="ADENOSINE/GUANOSINE DIPHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11782:SF33">
        <property type="entry name" value="ECTONUCLEOSIDE TRIPHOSPHATE DIPHOSPHOHYDROLASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01150">
        <property type="entry name" value="GDA1_CD39"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01238">
        <property type="entry name" value="GDA1_CD39_NTPASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="2 5" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="4" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="active site" description="Proton acceptor" evidence="3">
        <location>
            <position position="178"/>
        </location>
    </feature>
    <feature type="binding site" evidence="4">
        <location>
            <begin position="224"/>
            <end position="228"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00009283"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PIRSR" id="PIRSR600407-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PIRSR" id="PIRSR600407-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="RuleBase" id="RU003833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007317.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="288" mass="31560" checksum="59375EFF45624DBE" modified="2019-05-08" version="1">MQLSGVGKPKAVWFCTGSHLVGLGGSRIHLLLLLRAGLCASLSFSLSRPVPQYGIVLDAGSSHTAMFVYKWPADKENDTGIVSQHSMCDVEGGGISSYAENPPAAGKSLKQCLDQAVSDVPKERHPLTPLYLGATAGMRLLNLTDPQASDRVLSAVTSTLKSFPFDFRGAKILSGEDEGVFGWVTANYLLENFIKALLQSWIFLQPSRNWSHSKPTLGAMDLGGASTQITFETQKPIEDPRNEVTLRLYGQTYKVYTHSFLCYGRDQILRRMLSKVMKVRRSGSVPHP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYM8</accession>
    <name>A0A452GYM8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">TLDc domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GYM8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008347.1">
        <property type="protein sequence ID" value="ENSGAGP00000007218.1"/>
        <property type="gene ID" value="ENSGAGG00000005780.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006979">
        <property type="term" value="P:response to oxidative stress"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006571">
        <property type="entry name" value="TLDc_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23354:SF68">
        <property type="entry name" value="NUCLEAR RECEPTOR COACTIVATOR 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23354">
        <property type="entry name" value="NUCLEOLAR PROTEIN 7/ESTROGEN RECEPTOR COACTIVATOR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07534">
        <property type="entry name" value="TLD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00584">
        <property type="entry name" value="TLDc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51886">
        <property type="entry name" value="TLDC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="TLDc" evidence="1">
        <location>
            <begin position="58"/>
            <end position="219"/>
        </location>
    </feature>
    <evidence type="ECO:0000259" key="1">
        <source>
            <dbReference type="PROSITE" id="PS51886"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007218.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="219" mass="25284" checksum="06D11003593F7717" modified="2019-05-08" version="1">MRVRKIPLDIQIVYCVRPDQEPFVQIITVEEAKRRKSICSYYEEDDDDSLPVLKHHSALLENMHIEQLARCLPARVQGYPWRLAYSTLEHGTSLKTLYRKSASLDSPVLLVIKDMDNQIFGAYATHPFRFSDHYYGTGETFLYTFSPNFKVFKWSGENTYFINGDIGSLELGGGGGRFGLWLDVDLYHGRSNSCSTFNNDILSKKEDFIIQDVEVWTFE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYP1</accession>
    <name>A0A452GYP1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Rho guanine nucleotide exchange factor 11</fullName>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">May play a role in the regulation of RhoA GTPase by guanine nucleotide-binding alpha-12 (GNA12) and alpha-13 (GNA13). Acts as guanine nucleotide exchange factor (GEF) for RhoA GTPase and may act as GTPase-activating protein (GAP) for GNA12 and GNA13. Involved in neurotrophin-induced neurite outgrowth.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Membrane</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000008502.1">
        <property type="protein sequence ID" value="ENSGAGP00000007357.1"/>
        <property type="gene ID" value="ENSGAGG00000005896.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001664">
        <property type="term" value="F:G protein-coupled receptor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007186">
        <property type="term" value="P:G protein-coupled receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035556">
        <property type="term" value="P:intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd23069">
        <property type="entry name" value="PDZ_ARHGEF11-12-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd08753">
        <property type="entry name" value="RGS_PDZRhoGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00160">
        <property type="entry name" value="RhoGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.167.10:FF:000010">
        <property type="entry name" value="Rho guanine nucleotide exchange factor (GEF) 11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.900.10:FF:000006">
        <property type="entry name" value="Rho guanine nucleotide exchange factor (GEF) 11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000033">
        <property type="entry name" value="Rho guanine nucleotide exchange factor (GEF) 11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000072">
        <property type="entry name" value="Rho guanine nucleotide exchange factor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.42.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.900.10">
        <property type="entry name" value="Dbl homology (DH) domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.167.10">
        <property type="entry name" value="Regulator of G-protein Signalling 4, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035899">
        <property type="entry name" value="DBL_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000219">
        <property type="entry name" value="DH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001331">
        <property type="entry name" value="GDS_CDC24_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001478">
        <property type="entry name" value="PDZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036034">
        <property type="entry name" value="PDZ_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037889">
        <property type="entry name" value="PDZRhoGEF_RGS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041020">
        <property type="entry name" value="PH_16"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016137">
        <property type="entry name" value="RGS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015212">
        <property type="entry name" value="RGS-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036305">
        <property type="entry name" value="RGS_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044926">
        <property type="entry name" value="RGS_subdomain_2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45872:SF1">
        <property type="entry name" value="RHO GUANINE NUCLEOTIDE EXCHANGE FACTOR 11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45872">
        <property type="entry name" value="RHO GUANINE NUCLEOTIDE EXCHANGE FACTOR 2, ISOFORM D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00595">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17838">
        <property type="entry name" value="PH_16"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09128">
        <property type="entry name" value="RGS-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00621">
        <property type="entry name" value="RhoGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00228">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00233">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00315">
        <property type="entry name" value="RGS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00325">
        <property type="entry name" value="RhoGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48065">
        <property type="entry name" value="DBL homology domain (DH-domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50156">
        <property type="entry name" value="PDZ domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48097">
        <property type="entry name" value="Regulator of G-protein signaling, RGS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00741">
        <property type="entry name" value="DH_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50010">
        <property type="entry name" value="DH_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50106">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50132">
        <property type="entry name" value="RGS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="3" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="6" id="KW-0344">Guanine-nucleotide releasing factor</keyword>
    <keyword evidence="9" id="KW-0472">Membrane</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="PDZ" evidence="15">
        <location>
            <begin position="48"/>
            <end position="112"/>
        </location>
    </feature>
    <feature type="domain" description="RGS" evidence="16">
        <location>
            <begin position="358"/>
            <end position="451"/>
        </location>
    </feature>
    <feature type="domain" description="DH" evidence="14">
        <location>
            <begin position="779"/>
            <end position="968"/>
        </location>
    </feature>
    <feature type="domain" description="PH" evidence="13">
        <location>
            <begin position="1010"/>
            <end position="1123"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1"/>
            <end position="39"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="131"/>
            <end position="176"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="231"/>
            <end position="278"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="558"/>
            <end position="603"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="618"/>
            <end position="729"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1163"/>
            <end position="1242"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1274"/>
            <end position="1454"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1523"/>
            <end position="1590"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="10"/>
            <end position="26"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="131"/>
            <end position="140"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="12">
        <location>
            <begin position="148"/>
            <end position="162"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="254"/>
            <end position="269"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="566"/>
            <end position="578"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="626"/>
            <end position="637"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="646"/>
            <end position="682"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="696"/>
            <end position="709"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="12">
        <location>
            <begin position="1195"/>
            <end position="1212"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="1213"/>
            <end position="1224"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="1300"/>
            <end position="1311"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="1361"/>
            <end position="1373"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="1395"/>
            <end position="1404"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="12">
        <location>
            <begin position="1405"/>
            <end position="1424"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="1427"/>
            <end position="1438"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="1537"/>
            <end position="1554"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="1581"/>
            <end position="1590"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004370"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022658"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00057618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00074303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50010"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50106"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50132"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007357.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1590" mass="175099" checksum="67A45ADB4EDF5D38" modified="2019-05-08" version="1">MSVRPPQTALDRLSSLSSLGDSTTGRRSPGHRRQPSDCTETTGLVQRCVIIQKDQDGFGFTVSGDRIVLVQSVRPGGAAMKAGVQEGDRIVKVNGTMVTNSSHLEVVKLIKSGAYVALTLLGSPPPSVVLSSPQQDVSVSGAPGIAPLCPPPPPPPPLPPPQRITGPKPLQDPEVQKHATQILRNMLRQEEAELQRFCEVYSRNPATTLEERIEGARRRVSQLQLKIQQETGSSVDLGRPCCNSSSASLRGTEGRLSLDSQDGDSGLDSGTERFPSVSEISLNRNSVLSDPGLDSPRTSPVITSKMFQHHRRQGSDTPFLPIAEQGVDRTPRPLIIGPEEDYDPGYFNNEQCDSLFQDLGKLKSRPAHLGVFLRYIFSQADPSPLLFYLCAEVCQQMNSKDSRALGKEIWNIFLEKNAPLRIKVSEQLLADIESRLRNGEDVRSAFSEAQDAAMPEIQEQIQDYRTKRTMGLGSLYGENDLLDLDGDPQKERQVAEKQLAQLGDILSKYEEDRSSPMAFALSTYMSHTGIRMREARPASTSEKSQAFPDKDKWLPFFPKAKKSSSTKKDKDVMEDKKRNPILKYIGKPKSSSQSTFHVPLSPVEVKPGNVRNIIQHFENNQHSESQEPGTQRLSTGSFPEELLDSDSSRSEVKLGRSESLKGREEMKKSRKAENVPRSRSDVDMDAAAEATRLHQSASSSASSLSTRSLENPTPPYTPKMGRRSIESPSLGFGTDSFLPHLLEDDLGQLSDLEPEPDSQNWQHTVGREVMADLTHREIDRQEVINELFATEASHLRILRVLDVLFYQRMKRENLLSREELGLLFPNLPDLIDIHNSLSESMKKLREEGPIIKEIGDLMLSRFDGPAREEIQQVAADFCSYQSIALELIKTKQRKETRFQIFMQEAESNPQCRRLQLKDLIISEMQRLTKYPLLLENVLKHTDGGTPEHKKLCRARDQCREILRYVNEAVKRAENRHRLEAYQKRLDATSLERTSNPLAAEFKNLDLTSRCMIHEGALSWRISKEKTLDLHVLLLEDLLVLLQKQDEKLVLKFHSKTAPGSSDTKQTFSPVLKLNSMLIRSVATDKRALFIICTSELGPQIYELVALTPSEKNTWMELLEEAVQSSMRNATSPPKRRARELPIHRPAPSSLVLQDPDVSPILSQADSSRTEVDDVSSADDNPPDFLGKGKHPVLPEEPEAEGSEEEEEEEGEEAPAAPVPAQSSAELDEAGVSEEPGPSMHLPLPGLVLSKGLAESALEDVENLRHLILRILLPCRDPEPEDDLTPTPSVAGGAAQPWDSVLSSQDSGCQEQSVEHSGRGGPTTDRAGLPGHSPSGRPEPELESESRPSAGDVEETGWSSDGGREASQEVERPPAAEGQGGYKVVRKGNYFYVSMPAGPAESSTEPPAPKPPASPVEQPPSPPCPALGSQQHNQDSTLGPGSLDGAPPPEPPAPSLVIQDVDMIFRTIEQLTLKLTRLKDMELAHRELLRSLGRGSSAETTPVGVTALEMGRWLECPPCPASDHCPSPARRGCRPQQPASLAAEPPRRSPSAPVPCQGDLPKDGWVAGVAGHPLDPPNLGKDPWEVGKPEC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYP2</accession>
    <name>A0A452GYP2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Copper homeostasis protein cutC homolog</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">May play a role in copper homeostasis. Can bind one Cu(1+) per subunit.</text>
    </comment>
    <comment type="subunit">
        <text evidence="4">Homotetramer.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the CutC family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYP2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008418.1">
        <property type="protein sequence ID" value="ENSGAGP00000007279.1"/>
        <property type="gene ID" value="ENSGAGG00000005848.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005507">
        <property type="term" value="F:copper ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.20.20.380:FF:000002">
        <property type="entry name" value="copper homeostasis protein cutC homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.20.20.380">
        <property type="entry name" value="Copper homeostasis (CutC) domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="HAMAP" id="MF_00795">
        <property type="entry name" value="CutC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005627">
        <property type="entry name" value="CutC-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036822">
        <property type="entry name" value="CutC-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12598">
        <property type="entry name" value="COPPER HOMEOSTASIS PROTEIN CUTC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12598:SF0">
        <property type="entry name" value="COPPER HOMEOSTASIS PROTEIN CUTC HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03932">
        <property type="entry name" value="CutC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF110395">
        <property type="entry name" value="CutC-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0186">Copper</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9" id="KW-0539">Nucleus</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007768"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00011881"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00019014"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023008"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00055012"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007279.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="267" mass="28352" checksum="21737AB2355A1481" modified="2019-05-08" version="1">MGRRDRKPGSGPGNAGMGDGFLMEVCVDSVESAVNAERGGAGRIELCAGLMEGGTTPSMGLLQVVKQCVRVPVFVMIRPRGGDFLYSDRELEVMKADIRLAKLHGADGLVFGALTEDGHINTELCTALLAVCRPLPVTFHRAFDMVHDPLVALETLVSLGFERVLTSGCDSSALEGLPLIKRLTEQAKGRIVVVPGGGITERNLQRILEGSGAPEFHCSARSARDSGMKFRNSSVAMGASVSVSEYSLKVADVAKVRTLNAIAKNIL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYP4</accession>
    <name>A0A452GYP4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">RING-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYP4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008507.1">
        <property type="protein sequence ID" value="ENSGAGP00000007362.1"/>
        <property type="gene ID" value="ENSGAGG00000005909.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000139">
        <property type="term" value="C:Golgi membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061630">
        <property type="term" value="F:ubiquitin protein ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036503">
        <property type="term" value="P:ERAD pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd16475">
        <property type="entry name" value="RING-H2_RNF121-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.40.10:FF:000074">
        <property type="entry name" value="Ring finger protein 121"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040176">
        <property type="entry name" value="RNF121/RNF175"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001841">
        <property type="entry name" value="Znf_RING"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13407:SF1">
        <property type="entry name" value="E3 UBIQUITIN LIGASE RNF121"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13407">
        <property type="entry name" value="RNF121 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00184">
        <property type="entry name" value="RING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57850">
        <property type="entry name" value="RING/U-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50089">
        <property type="entry name" value="ZF_RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7 9" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 9" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 9" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <keyword evidence="4 8" id="KW-0863">Zinc-finger</keyword>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="37"/>
            <end position="55"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="84"/>
            <end position="103"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="124"/>
            <end position="146"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="158"/>
            <end position="175"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="285"/>
            <end position="307"/>
        </location>
    </feature>
    <feature type="domain" description="RING-type" evidence="10">
        <location>
            <begin position="209"/>
            <end position="259"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007362.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="310" mass="36376" checksum="BA2733C631384A97" modified="2019-05-08" version="1">MRKQVDLSRLSPEERWRVEHARMHAKHRGHEAMHAEMVLILIATLVVAQLLLVQWKQRHPRSYNMVTLFQMWVVPLYFTAKLHWWRFLVIWVLFSAVTAFVTFRATRKPLVQTTPRLVYKWFLLIYKISYATGIVGYMAVMFTLFGLNLLFRIKPEDAMDFGVSLLFYGLYYGVLERDFAEMCADYMASTIGFYSASGMPTKHLSDSVCAVCGQQIFVDVNEEGIIENTYRLSCNHVFHEFCIRGWCIVGKKQTCPYCKEKVDLKRMFSNPWERPHVMYGQLLDWLRYLVAWQPVIIGLVQAINYILGLE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYQ3</accession>
    <name>A0A452GYQ3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">tRNA (guanine(10)-N(2))-methyltransferase TRMT11</fullName>
            <ecNumber evidence="12">2.1.1.214</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="14">tRNA methyltransferase 11 homolog</fullName>
        </alternativeName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">Catalytic subunit of the TRMT11-TRM112 methyltransferase complex, that specifically mediates the S-adenosyl-L-methionine-dependent N(2)-methylation of guanosine nucleotide at position 10 (m2G10) in tRNAs. This is one of the major tRNA (guanine-N(2))-methyltransferases.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="9">
            <text>guanosine(10) in tRNA + S-adenosyl-L-methionine = N(2)-methylguanosine(10) in tRNA + S-adenosyl-L-homocysteine + H(+)</text>
            <dbReference type="Rhea" id="RHEA:43128"/>
            <dbReference type="Rhea" id="RHEA-COMP:10355"/>
            <dbReference type="Rhea" id="RHEA-COMP:10357"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57856"/>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
            <dbReference type="ChEBI" id="CHEBI:74269"/>
            <dbReference type="ChEBI" id="CHEBI:74481"/>
            <dbReference type="EC" id="2.1.1.214"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="9">
            <dbReference type="Rhea" id="RHEA:43129"/>
        </physiologicalReaction>
    </comment>
    <comment type="subunit">
        <text evidence="11">Part of the heterodimeric TRMT11-TRM112 methyltransferase complex; this complex forms an active tRNA methyltransferase, where TRMT112 acts as an activator of the catalytic subunit TRMT11.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="15">Belongs to the class I-like SAM-binding methyltransferase superfamily. TRM11 methyltransferase family.</text>
    </comment>
    <dbReference type="EC" id="2.1.1.214" evidence="12"/>
    <dbReference type="AlphaFoldDB" id="A0A452GYQ3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008371.1">
        <property type="protein sequence ID" value="ENSGAGP00000007240.1"/>
        <property type="gene ID" value="ENSGAGG00000005813.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043527">
        <property type="term" value="C:tRNA methyltransferase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0160102">
        <property type="term" value="F:tRNA (guanine(10)-N2)-methyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000049">
        <property type="term" value="F:tRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032259">
        <property type="term" value="P:methylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008033">
        <property type="term" value="P:tRNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.150:FF:000069">
        <property type="entry name" value="tRNA (Guanine(10)-N2)-methyltransferase homolog isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.150">
        <property type="entry name" value="Vaccinia Virus protein VP39"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002052">
        <property type="entry name" value="DNA_methylase_N6_adenine_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000241">
        <property type="entry name" value="RlmKL-like_Mtase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029063">
        <property type="entry name" value="SAM-dependent_MTases_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016691">
        <property type="entry name" value="TRMT11"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR059073">
        <property type="entry name" value="TRMT11_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13370">
        <property type="entry name" value="RNA METHYLASE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13370:SF3">
        <property type="entry name" value="TRNA (GUANINE(10)-N2)-METHYLTRANSFERASE HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25904">
        <property type="entry name" value="Tmrp11_N"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01170">
        <property type="entry name" value="UPF0020"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF017259">
        <property type="entry name" value="tRNA_mtfrase_TRM11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00507">
        <property type="entry name" value="N12N6MTFRASE"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53335">
        <property type="entry name" value="S-adenosyl-L-methionine-dependent methyltransferases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00092">
        <property type="entry name" value="N6_MTASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51627">
        <property type="entry name" value="SAM_MT_TRM11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4 15" id="KW-0489">Methyltransferase</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8 15" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="6 15" id="KW-0949">S-adenosyl-L-methionine</keyword>
    <keyword evidence="16" id="KW-0732">Signal</keyword>
    <keyword evidence="5 15" id="KW-0808">Transferase</keyword>
    <keyword evidence="7 15" id="KW-0819">tRNA processing</keyword>
    <keyword evidence="3 15" id="KW-0820">tRNA-binding</keyword>
    <feature type="signal peptide" evidence="16">
        <location>
            <begin position="1"/>
            <end position="16"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019436784" description="tRNA (guanine(10)-N(2))-methyltransferase TRMT11" evidence="16">
        <location>
            <begin position="17"/>
            <end position="442"/>
        </location>
    </feature>
    <feature type="domain" description="tRNA (guanine(10)-N(2))-methyltransferase TRMT11 N-terminal" evidence="18">
        <location>
            <begin position="22"/>
            <end position="106"/>
        </location>
    </feature>
    <feature type="domain" description="tRNA (guanine(10)-N(2))-methyltransferase TRMT11 N-terminal" evidence="18">
        <location>
            <begin position="113"/>
            <end position="162"/>
        </location>
    </feature>
    <feature type="domain" description="Ribosomal RNA large subunit methyltransferase K/L-like methyltransferase" evidence="17">
        <location>
            <begin position="172"/>
            <end position="303"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022555"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022603"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022691"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022694"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00050985"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00056270"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00065434"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00066937"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00067484"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00075308"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00959"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF01170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF25904"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007240.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="442" mass="50849" checksum="33F9F9847A59C2D5" modified="2019-05-08" version="1">MRGWAERFMASGGAMARGLLPSYLLLLAQENLEFRLPEIKSLLSLCGGQFSSQQEIRINSPFWILNIPSEEMARGLMKRTVCAKSVFELWGQGKSAEELYTSLKNYPMEKMALEFLPFEGKVNLKNPEHIFWILEDYGMDPNNVPEDPLDLYFGRWIANGQRELIESYSVKRRHFIGNTSMDACLSFIMANHGRVKANDVVFDPFVGTGGLLISSARFGAYVCGTDIDYNTVHGLGKASRKNQKWRGPDENIRANLRQYGLEKYYLDALVSDASKPIWRKEMLFDAIITDPPYGIREATRRTGSQKEILKATERGTENHISISSNYHLSDIFFDLLNSAAEYLVMGGRLVYWLPVYKPEYTEEIIPRHPCLKLISNCEQMLSSHTSRRLITMEKVKEFEDEDQYSHLLDYQSLLYKGHNSFREKYFSGVTKRIAKEEKDNQK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYQ4</accession>
    <name>A0A452GYQ4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Gamma-tubulin complex component</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="6">Component of the gamma-tubulin ring complex (gTuRC) which mediates microtubule nucleation. The gTuRC regulates the minus-end nucleation of alpha-beta tubulin heterodimers that grow into microtubule protafilaments, a critical step in centrosome duplication and spindle formation.</text>
    </comment>
    <comment type="subunit">
        <text evidence="7">Component of the gamma-tubulin ring complex (gTuRC) consisting of TUBGCP2, TUBGCP3, TUBGCP4, TUBGCP5 and TUBGCP6 and gamma-tubulin TUBG1 or TUBG2. TUBGCP2, TUBGCP3, TUBGCP4, TUBGCP5 and TUBGCP6 assemble in a 5:5:2:1:1 stoichiometry; each is associated with a gamma-tubulin, thereby arranging 14 gamma-tubulins in a helical manner. Gamma-tubulin at the first position is blocked by TUBGCP3 at the last position, allowing 13 protafilaments to grow into a microtubule. The gTuRC (via TUBGCP3 and TUBGCP6) interacts with ACTB and MZT1; the interactions form a luminal bridge that stabilizes the initial structure during complex assembly. The gTuRC (via TUBGCP2) interacts with MZT2A/MZT2B and CDK5RAP2 (via CM1 motif); the interactions play a role in gTuRC activation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Microtubule organizing center</location>
            <location evidence="1">Centrosome</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 8">Belongs to the TUBGCP family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYQ4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008315.1">
        <property type="protein sequence ID" value="ENSGAGP00000007189.1"/>
        <property type="gene ID" value="ENSGAGG00000004413.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000930">
        <property type="term" value="C:gamma-tubulin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005874">
        <property type="term" value="C:microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000922">
        <property type="term" value="C:spindle pole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043015">
        <property type="term" value="F:gamma-tubulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051011">
        <property type="term" value="F:microtubule minus-end binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031122">
        <property type="term" value="P:cytoplasmic microtubule organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051321">
        <property type="term" value="P:meiotic cell cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007020">
        <property type="term" value="P:microtubule nucleation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000278">
        <property type="term" value="P:mitotic cell cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051225">
        <property type="term" value="P:spindle assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd22572">
        <property type="entry name" value="GCP5_NTD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.1900:FF:000005">
        <property type="entry name" value="Gamma-tubulin complex component"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.1900">
        <property type="entry name" value="Gamma-tubulin complex, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007259">
        <property type="entry name" value="GCP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR059169">
        <property type="entry name" value="GCP5_N_ext"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040457">
        <property type="entry name" value="GCP_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042241">
        <property type="entry name" value="GCP_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041470">
        <property type="entry name" value="GCP_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19302">
        <property type="entry name" value="GAMMA TUBULIN COMPLEX PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19302:SF33">
        <property type="entry name" value="GAMMA-TUBULIN COMPLEX COMPONENT 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04130">
        <property type="entry name" value="GCP_C_terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17681">
        <property type="entry name" value="GCP_N_terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 8" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="5 8" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="4 8" id="KW-0493">Microtubule</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Gamma tubulin complex component protein N-terminal" evidence="11">
        <location>
            <begin position="250"/>
            <end position="599"/>
        </location>
    </feature>
    <feature type="domain" description="Gamma tubulin complex component C-terminal" evidence="10">
        <location>
            <begin position="698"/>
            <end position="983"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="129"/>
            <end position="160"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="507"/>
            <end position="526"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="9">
        <location>
            <begin position="140"/>
            <end position="151"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="513"/>
            <end position="525"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010337"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00093416"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00093551"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="RuleBase" id="RU363050"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="Pfam" id="PF04130"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF17681"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007189.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1006" mass="116386" checksum="2FA6410372BCD9CE" modified="2019-05-08" version="1">MAGQPPRWSPLEQAQDRDVRALIRRLTGLREAQGGEEPGGRFQTALIFAWSNFRFHRFLDVSSHKVERTIEGIYEKLIVHSDLSKAASWKRLIEKFLNSPLPSTEETKTDAHYGILSLLLCLSDSPSNTNYVEKPRDKEDWSEESEGEEDAQQPLSREDSGIQVDRTPQEEQDQNKKTVSQVTWKVGEPDARSWLEQHVVPQYWTGRAPRFSHSFHLHSNLAAVWDQHLYSSDPLYMPEERTLVTETQIIRETLWLLSGVKKLFIFQLNEGKVTVRNDIIVTHLTHNCLRSVLEQIAAYGQVVFRLQKFIDEVMGHSSESVIPGTISTPKKTTEPPFRTYQAFMWALYKYFISFKEELTEIEKCVINKDKTVTLSIVVDKLSPRLAQLKVLHKVFSTGIAEVPQVPPDTRNVVRASHLLNTLYKAILEYDNVGEASEQTVSLLFALWVETVRPYLQTVDEWIVHGNLFDPAKEFIIQRNKNVPVNHRDFWYATYTLYSVSEKMENEEKMSDNASASSGSDQAPSSRQHTMVSFLKPVLKQIIMAGKSMQLLKNLQSKDDSPRQAASRDAERKSLYTLFLESVQSRLRHGEESVPDIITEQQATKQSLIKMQSIAERHLELDDVHDPLLAINFARLYLEQSDFHEKFTGGDVCVDRSSESVTCQTFELTLRSCLYPHIDKQYLECCGNLMQTLKKDYRLVEYLQAMRNFFLLEAGDTMYDFYTSIFDKIREKETWQNVSFLNVQLQEAVGQRYPEDSSRLSISFESIDTAKKKLPVHTLDGLTLSYKVPWPVDIVISLECQKIYNQVFLLLLQIKWAKYSLDVLRFDELVNATEKPQLKEGPQLEQETVPQFGSQTELIKQQIHRMFLVRVKLMHFVNSLHNYIMTRILHSTGLEFQHQVEEAKDLDQLIKIHYRYLSTIHDRCLLREKVSFVKEAIMKVLNLVLIFADRWQAGLGAWKMESIEKMESDFKNCHMFLVTVLNKAVCRGSFPHLLVMAVFFFLHVSML</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYQ7</accession>
    <name>A0A452GYQ7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">DUF7789 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GYQ7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008435.1">
        <property type="protein sequence ID" value="ENSGAGP00000007294.1"/>
        <property type="gene ID" value="ENSGAGG00000005863.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056691">
        <property type="entry name" value="DUF7789"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR39299">
        <property type="entry name" value="TRANSMEMBRANE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR39299:SF1">
        <property type="entry name" value="TRANSMEMBRANE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25044">
        <property type="entry name" value="DUF7789"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0472">Membrane</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="2" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="50"/>
            <end position="68"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="80"/>
            <end position="97"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="109"/>
            <end position="132"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="138"/>
            <end position="160"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="218"/>
            <end position="239"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="246"/>
            <end position="267"/>
        </location>
    </feature>
    <feature type="domain" description="DUF7789" evidence="3">
        <location>
            <begin position="37"/>
            <end position="161"/>
        </location>
    </feature>
    <feature type="domain" description="DUF7789" evidence="3">
        <location>
            <begin position="172"/>
            <end position="270"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="Pfam" id="PF25044"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007294.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="276" mass="31044" checksum="E43EC92CF8A6D84C" modified="2019-05-08" version="1">MEPLDNRDPLFSPPNETADLPLPNLTPRHQELIPTPCGPIKPWSELSSPVKIYFCVTILSLLSVIGLTTETLFQQTHEDFTISLIQLIGVVFCVYYVTRGILQENRQELIVFILSILLVMFRSIVNFTVVSAEQKPNVLARFILILLVGSFDVICAIILIQSQSMMAFRVGGALESLQSQYFMLNLCFSMLTFDLQAQLCLCALLLTSLHEISLLHSSILAIGVFWACLKVTVGIIAILKELKPLVWIFLSQNIPEVANLVYLLYMVTHQVFVDLK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYQ8</accession>
    <name>A0A452GYQ8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">tRNA (guanine(10)-N(2))-methyltransferase TRMT11</fullName>
            <ecNumber evidence="12">2.1.1.214</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="14">tRNA methyltransferase 11 homolog</fullName>
        </alternativeName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">Catalytic subunit of the TRMT11-TRM112 methyltransferase complex, that specifically mediates the S-adenosyl-L-methionine-dependent N(2)-methylation of guanosine nucleotide at position 10 (m2G10) in tRNAs. This is one of the major tRNA (guanine-N(2))-methyltransferases.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="9">
            <text>guanosine(10) in tRNA + S-adenosyl-L-methionine = N(2)-methylguanosine(10) in tRNA + S-adenosyl-L-homocysteine + H(+)</text>
            <dbReference type="Rhea" id="RHEA:43128"/>
            <dbReference type="Rhea" id="RHEA-COMP:10355"/>
            <dbReference type="Rhea" id="RHEA-COMP:10357"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57856"/>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
            <dbReference type="ChEBI" id="CHEBI:74269"/>
            <dbReference type="ChEBI" id="CHEBI:74481"/>
            <dbReference type="EC" id="2.1.1.214"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="9">
            <dbReference type="Rhea" id="RHEA:43129"/>
        </physiologicalReaction>
    </comment>
    <comment type="subunit">
        <text evidence="11">Part of the heterodimeric TRMT11-TRM112 methyltransferase complex; this complex forms an active tRNA methyltransferase, where TRMT112 acts as an activator of the catalytic subunit TRMT11.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="15">Belongs to the class I-like SAM-binding methyltransferase superfamily. TRM11 methyltransferase family.</text>
    </comment>
    <dbReference type="EC" id="2.1.1.214" evidence="12"/>
    <dbReference type="AlphaFoldDB" id="A0A452GYQ8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008376.1">
        <property type="protein sequence ID" value="ENSGAGP00000007245.1"/>
        <property type="gene ID" value="ENSGAGG00000005813.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043527">
        <property type="term" value="C:tRNA methyltransferase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0160102">
        <property type="term" value="F:tRNA (guanine(10)-N2)-methyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000049">
        <property type="term" value="F:tRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032259">
        <property type="term" value="P:methylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008033">
        <property type="term" value="P:tRNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.150">
        <property type="entry name" value="Vaccinia Virus protein VP39"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002052">
        <property type="entry name" value="DNA_methylase_N6_adenine_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000241">
        <property type="entry name" value="RlmKL-like_Mtase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029063">
        <property type="entry name" value="SAM-dependent_MTases_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016691">
        <property type="entry name" value="TRMT11"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR059073">
        <property type="entry name" value="TRMT11_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13370">
        <property type="entry name" value="RNA METHYLASE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13370:SF3">
        <property type="entry name" value="TRNA (GUANINE(10)-N2)-METHYLTRANSFERASE HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25904">
        <property type="entry name" value="Tmrp11_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01170">
        <property type="entry name" value="UPF0020"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF017259">
        <property type="entry name" value="tRNA_mtfrase_TRM11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53335">
        <property type="entry name" value="S-adenosyl-L-methionine-dependent methyltransferases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00092">
        <property type="entry name" value="N6_MTASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51627">
        <property type="entry name" value="SAM_MT_TRM11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4 15" id="KW-0489">Methyltransferase</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8 15" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="6 15" id="KW-0949">S-adenosyl-L-methionine</keyword>
    <keyword evidence="16" id="KW-0732">Signal</keyword>
    <keyword evidence="5 15" id="KW-0808">Transferase</keyword>
    <keyword evidence="7 15" id="KW-0819">tRNA processing</keyword>
    <keyword evidence="3 15" id="KW-0820">tRNA-binding</keyword>
    <feature type="signal peptide" evidence="16">
        <location>
            <begin position="1"/>
            <end position="16"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019233070" description="tRNA (guanine(10)-N(2))-methyltransferase TRMT11" evidence="16">
        <location>
            <begin position="17"/>
            <end position="429"/>
        </location>
    </feature>
    <feature type="domain" description="tRNA (guanine(10)-N(2))-methyltransferase TRMT11 N-terminal" evidence="18">
        <location>
            <begin position="22"/>
            <end position="193"/>
        </location>
    </feature>
    <feature type="domain" description="Ribosomal RNA large subunit methyltransferase K/L-like methyltransferase" evidence="17">
        <location>
            <begin position="203"/>
            <end position="334"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022555"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022603"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022691"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022694"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00050985"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00056270"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00065434"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00066937"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00067484"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00075308"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00959"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF01170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF25904"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007245.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="429" mass="49475" checksum="ADF7463BBB238C35" modified="2019-05-08" version="1">MRGWAERFMASGGAMARGLLPSYLLLLAQENLEFRLPEIKSLLSLCGGQFSSQQEIRINSPFWILNIPSEEMARGLMKRTVCAKSVFELWGQGKSAEELYTSLKNYPMEKMVPYLQSNSTYKIHIHAFNKTLTQDKKIRRIDALEFLPFEGKVNLKNPEHIFWILEDYGMDPNNVPEDPLDLYFGRWIANGQRELIESYSVKRRHFIGNTSMDACLSFIMANHGRVKANDVVFDPFVGTGGLLISSARFGAYVCGTDIDYNTVHGLGKASRKNQKWRGPDENIRANLRQYGLEKYYLDALVSDASKPIWRKEMLFDAIITDPPYGIREATRRTGSQKEILKATERGYTEEIIPRHPCLKLISNCEQMLSSHTSRRLITMEKVKEFEDEDQYSHLLDYQSLLYKGHNSFREKYFSGVTKRIAKEEKDNQK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ13</accession>
    <name>A0A452GZ13_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="10">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Mitochondrion outer membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the Tom40 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZ13"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008668.1">
        <property type="protein sequence ID" value="ENSGAGP00000007505.1"/>
        <property type="gene ID" value="ENSGAGG00000006008.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005741">
        <property type="term" value="C:mitochondrial outer membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008320">
        <property type="term" value="F:protein transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030150">
        <property type="term" value="P:protein import into mitochondrial matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd07305">
        <property type="entry name" value="Porin3_Tom40"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.160.10:FF:000005">
        <property type="entry name" value="mitochondrial import receptor subunit TOM40 homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.160.10">
        <property type="entry name" value="Porin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023614">
        <property type="entry name" value="Porin_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027246">
        <property type="entry name" value="Porin_Euk/Tom40"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037930">
        <property type="entry name" value="Tom40"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10802">
        <property type="entry name" value="MITOCHONDRIAL IMPORT RECEPTOR SUBUNIT TOM40"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01459">
        <property type="entry name" value="Porin_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="6" id="KW-1000">Mitochondrion outer membrane</keyword>
    <keyword evidence="7" id="KW-0653">Protein transport</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4" id="KW-1134">Transmembrane beta strand</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004374"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010510"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022452"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022787"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007505.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="305" mass="33937" checksum="830C0FAD221C8365" modified="2019-05-08" version="1">MGNVLAPVLDLHRPLRREEPLSNPGSFDELHRQCKEVFPQQMEGVKLIINKALSSHFQVTHTIHMSTVGQSSYHFNTTYVGDRQLSPTETFPMLVGDIDSCGSLNAQVLHLVAERIRTKAVFQTQQARFVTWQFDGELRGDDYTATLTLGNPDMLSQSVIVVAHFLQSVTSRLVLGGEMVYHRRPGEEGAIVTLAGKYTALKWVATLNIGYGGAHASYYHRANEQVQVGVELEANTRLQDTTFAFGYQLNLPQANMVFRGLLDSNWCVGGVLEKKLPPLPVTLALGAFLNHWKNRFHCGFSVIVG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ16</accession>
    <name>A0A452GZ16_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Tyrosine-protein kinase receptor</fullName>
            <ecNumber evidence="20">2.7.10.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="17">Receptor with tyrosine-protein kinase activity. Functions as a pH sensing receptor which is activated by increased extracellular pH. Activates an intracellular signaling pathway that involves IRS1 and AKT1/PKB.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="16">
            <text>L-tyrosyl-[protein] + ATP = O-phospho-L-tyrosyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10596"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.10.1"/>
        </reaction>
    </comment>
    <comment type="subunit">
        <text evidence="18">Probable tetramer of 2 alpha and 2 beta chains linked by disulfide bonds. The alpha chains contribute to the formation of the ligand-binding domain, while the beta chains carry the kinase domain.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="20">Belongs to the protein kinase superfamily. Tyr protein kinase family. Insulin receptor subfamily.</text>
    </comment>
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    <dbReference type="AlphaFoldDB" id="A0A452GZ16"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008569.1">
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        <property type="gene ID" value="ENSGAGG00000005944.1"/>
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        <property type="component" value="Unassembled WGS sequence"/>
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        <property type="term" value="C:axon"/>
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        <property type="project" value="TreeGrafter"/>
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        <property type="project" value="UniProtKB-UniRule"/>
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        <property type="term" value="F:insulin receptor activity"/>
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        <property type="term" value="F:insulin receptor substrate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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        <property type="match status" value="3"/>
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        <property type="entry name" value="FU"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="PTKc_InsR_like"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Tyrosine-protein kinase receptor"/>
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        <property type="entry name" value="Hormone Receptor, Insulin-like Growth Factor Receptor 1, Chain A, domain 2"/>
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        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="4"/>
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        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
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        <property type="entry name" value="Receptor L-domain"/>
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        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
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        <property type="entry name" value="FN3_dom"/>
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        <property type="entry name" value="FN3_sf"/>
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        <property type="entry name" value="Furin-like_Cys-rich_dom"/>
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        <property type="entry name" value="Furin_repeat"/>
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        <property type="entry name" value="Growth_fac_rcpt_cys_sf"/>
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        <property type="entry name" value="Ig-like_fold"/>
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        <property type="entry name" value="Kinase-like_dom_sf"/>
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        <property type="entry name" value="Prot_kinase_dom"/>
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        <property type="entry name" value="Protein_kinase_ATP_BS"/>
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        <property type="entry name" value="Rcpt_L-dom"/>
    </dbReference>
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        <property type="entry name" value="Rcpt_L-dom_sf"/>
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        <property type="entry name" value="RTK"/>
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    <dbReference type="InterPro" id="IPR001245">
        <property type="entry name" value="Ser-Thr/Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008266">
        <property type="entry name" value="Tyr_kinase_AS"/>
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        <property type="entry name" value="Tyr_kinase_cat_dom"/>
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        <property type="entry name" value="Tyr_kinase_rcpt_2_CS"/>
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        <property type="entry name" value="INSULIN RECEPTOR-RELATED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416">
        <property type="entry name" value="TYROSINE-PROTEIN KINASE RECEPTOR"/>
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    </dbReference>
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    </dbReference>
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        <property type="entry name" value="PK_Tyr_Ser-Thr"/>
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    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00109">
        <property type="entry name" value="TYRKINASE"/>
    </dbReference>
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        <property type="entry name" value="FN3"/>
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    </dbReference>
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        <property type="entry name" value="FU"/>
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    </dbReference>
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        <property type="entry name" value="TyrKc"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="3"/>
    </dbReference>
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        <property type="entry name" value="Growth factor receptor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52058">
        <property type="entry name" value="L domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
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    </dbReference>
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        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
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    </dbReference>
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        <property type="entry name" value="PROTEIN_KINASE_TYR"/>
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    </dbReference>
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        <property type="entry name" value="RECEPTOR_TYR_KIN_II"/>
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    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9 19" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="13" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="15" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="8" id="KW-0418">Kinase</keyword>
    <keyword evidence="11" id="KW-0472">Membrane</keyword>
    <keyword evidence="7 19" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="2 20" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="14 20" id="KW-0675">Receptor</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <keyword evidence="3" id="KW-0808">Transferase</keyword>
    <keyword evidence="4 20" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="10" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="12" id="KW-0829">Tyrosine-protein kinase</keyword>
    <feature type="domain" description="Fibronectin type-III" evidence="23">
        <location>
            <begin position="308"/>
            <end position="428"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="23">
        <location>
            <begin position="654"/>
            <end position="749"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="22">
        <location>
            <begin position="803"/>
            <end position="1078"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="21">
        <location>
            <begin position="562"/>
            <end position="590"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="21">
        <location>
            <begin position="562"/>
            <end position="573"/>
        </location>
    </feature>
    <feature type="binding site" evidence="19">
        <location>
            <position position="837"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
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    </evidence>
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            <dbReference type="ARBA" id="ARBA00022840"/>
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    </evidence>
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        <source>
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    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00065424"/>
        </source>
    </evidence>
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        <source>
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    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
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        <source>
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    </evidence>
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    </evidence>
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    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1152" mass="129939" checksum="F6815622F20A874E" modified="2019-05-08" version="1">MGDVVCPGSLAKQGKWELLPAVLREKPYNERNSSPRNGCYGIACTAAGECCHPECLGGCSRPWDSRACVACRHFHFNGHCLTSCPTRTYRYEGWRCVTAEYCASLRKVSENPRDASEFVIHHERCLSECPSGYTRNDSSIFCHKCEGLCPKECKVGTKTIDSTRAAQELAGCTYVEGNLIVNIRRGYNLASELQNSLGLIETITGFLKIKHSFALVSLSFFKNLKLIRGDSMVDGNYTLYVLDNQNLQQLWDWSHHTLSIPVGKVYFAFNPKLCLSEIYRMEEVTGTKGRQNKAEINPRTNGDRASCESRTLRFVSNITESDRIFLKWERYRPPEYRDLLSFIVYYKESTFQNVTEYVGQDACGANSWNVVDVELPLNNDQDPGVALLNLKPWTQYAIFVRAITLTTAEEGRNYGAQSEVVYVRTMPAAPTVPRDVLSMSNSSSHIIVRWKPPSQRNGNITYYLVLWQQLAEDLELYVNDYCHKGLKLPTSSADTRFNVDDSGGPELEQDTEERCCPCHRDSGQLQMEGEAVSFQKKFENFLHNSIIIPRTRKRRRDILAMTSSGHPLSNASSLEPPGKPPSLNLTGAGPKAQTDFRIFEDKVVRDRTVISQLRHFTEYRIDIHACNHAAHTVGCSAATFVFARTMPELEADNIPGNVTWEPASKNSVLLRWEEPKNPNGLILKYEIKYSRENEVTTVVCVSRHRYAKYRGVHLALLQPGNYSAKVRATSLAGNGSWTGLIKFYILGPGMYLGEALTFVLLFPPRRNNDGYPSGTLYASVNPEYFSTSDMYIPDEWEVSRDKITVIRELGQGSFGMVYEGIAQGIEKEDEETKVALKTVNELATMRERIEFLNEASVMKAFKCHHVVRLLGVVSQGQPALVIMELMTRGDLKSYLRSLRPDAENNPGLLPPSLKNMIQMAGEIADGMAYLNAKKFVHRDLAARNCMVSEDFTVKIGDFGMTRDIYETDYYRKGGKGLLPVRWMSPEALKDGIFNTHSDVWSFGVVLWEIATLAEQPYQGLSNEQVLRFVMDNGILKRPENCPEKLHELMSWCWQQNPRQRPSFTQLLESIKDDMDPTFRNISFFYSAENKRRGSCEPSDVETDNLLEEEPSTALAALRLPLATNGSPGPAVAPQKDRSCSRNCCHLNGAPYL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ20</accession>
    <name>A0A452GZ20_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">CFA20 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GZ20"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008436.1">
        <property type="protein sequence ID" value="ENSGAGP00000007295.1"/>
        <property type="gene ID" value="ENSGAGG00000005860.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040441">
        <property type="entry name" value="CFA20/CFAP20DC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007714">
        <property type="entry name" value="CFA20_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12458">
        <property type="entry name" value="ORF PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05018">
        <property type="entry name" value="CFA20_dom"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="CFA20" evidence="2">
        <location>
            <begin position="1"/>
            <end position="175"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="219"/>
            <end position="247"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="336"/>
            <end position="451"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="225"/>
            <end position="247"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="336"/>
            <end position="372"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="378"/>
            <end position="393"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="1">
        <location>
            <begin position="435"/>
            <end position="451"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF05018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007295.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="451" mass="50323" checksum="6BA64B71ED336E37" modified="2019-05-08" version="1">MFRNEYQGGPFVEIFSAQGKNPGAKWKIFGSPSAIWKEFDKEVKGFVFVLEGSSQTNKIQLPKETRQTLGLIQRFLVLQIYVPLGQDFSTELLVTDLGNIKRRLYLSTVHKELSITPLHAKLPLFMIKRRIWCNLCIDLVAFTSEIFRGAVFQSLDGIIISANCKLRKIFTLKSKPQDTAEEDADDPTDTIPRTCQLNTDVPQVTQLLNMTKLRQPEIRFGGRPLTSSESDQLINRGQGSTHNSKNQDVSHIAFGSKVLGPPPPSGRKVNTRVSGEITKSLGSRSNRSCQQLTLEKGSESMRDTKRSGLCLSPFYKAFDQGDKENIHQTNQITSQNDDWIFPENSSDSFQIDSNGQSVATHNTSCSNLSLPPNTEVESESKQTGEHQISHKEIFTFSSRPRSAPHGKSPNMSPEGCMSPLDLNEDSDGFCRGAQMEDDFYGSDSNEETLSN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ22</accession>
    <name>A0A452GZ22_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">ATP-dependent DNA helicase</fullName>
            <ecNumber evidence="17">5.6.2.4</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="23">
            <text>ATP + H2O = ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:13065"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="16">
            <text>Couples ATP hydrolysis with the unwinding of duplex DNA by translocating in the 3'-5' direction.</text>
            <dbReference type="EC" id="5.6.2.4"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2 17">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 17">Belongs to the helicase family. RecQ subfamily.</text>
    </comment>
    <dbReference type="EC" id="5.6.2.4" evidence="17"/>
    <dbReference type="AlphaFoldDB" id="A0A452GZ22"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008574.1">
        <property type="protein sequence ID" value="ENSGAGP00000007424.1"/>
        <property type="gene ID" value="ENSGAGG00000005383.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005694">
        <property type="term" value="C:chromosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043138">
        <property type="term" value="F:3'-5' DNA helicase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="RHEA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009378">
        <property type="term" value="F:four-way junction helicase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006260">
        <property type="term" value="P:DNA replication"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000724">
        <property type="term" value="P:double-strand break repair via homologous recombination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000723">
        <property type="term" value="P:telomere maintenance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd18016">
        <property type="entry name" value="DEXHc_RecQ2_BLM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18794">
        <property type="entry name" value="SF2_C_RecQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000537">
        <property type="entry name" value="Bloom syndrome RecQ-like helicase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000340">
        <property type="entry name" value="Bloom syndrome, RecQ helicase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.80">
        <property type="entry name" value="HRDC domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.10">
        <property type="entry name" value="Winged helix-like DNA-binding domain superfamily/Winged helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012532">
        <property type="entry name" value="BDHCT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032439">
        <property type="entry name" value="BDHCT_assoc"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011545">
        <property type="entry name" value="DEAD/DEAH_box_helicase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002464">
        <property type="entry name" value="DNA/RNA_helicase_DEAH_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004589">
        <property type="entry name" value="DNA_helicase_ATP-dep_RecQ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014001">
        <property type="entry name" value="Helicase_ATP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001650">
        <property type="entry name" value="Helicase_C-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044876">
        <property type="entry name" value="HRDC_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032284">
        <property type="entry name" value="RecQ_Zn-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018982">
        <property type="entry name" value="RQC_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036388">
        <property type="entry name" value="WH-like_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036390">
        <property type="entry name" value="WH_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00614">
        <property type="entry name" value="recQ_fam"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13710">
        <property type="entry name" value="DNA HELICASE RECQ FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13710:SF153">
        <property type="entry name" value="RECQ-LIKE DNA HELICASE BLM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08072">
        <property type="entry name" value="BDHCT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16204">
        <property type="entry name" value="BDHCT_assoc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00270">
        <property type="entry name" value="DEAD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00271">
        <property type="entry name" value="Helicase_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16124">
        <property type="entry name" value="RecQ_Zn_bind"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09382">
        <property type="entry name" value="RQC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00487">
        <property type="entry name" value="DEXDc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00490">
        <property type="entry name" value="HELICc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00956">
        <property type="entry name" value="RQC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46785">
        <property type="entry name" value="Winged helix' DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00690">
        <property type="entry name" value="DEAH_ATP_HELICASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51192">
        <property type="entry name" value="HELICASE_ATP_BIND_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51194">
        <property type="entry name" value="HELICASE_CTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="11 17" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="7" id="KW-0227">DNA damage</keyword>
    <keyword evidence="13" id="KW-0234">DNA repair</keyword>
    <keyword evidence="4" id="KW-0235">DNA replication</keyword>
    <keyword evidence="12" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="9 17" id="KW-0347">Helicase</keyword>
    <keyword evidence="8 17" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="14" id="KW-0413">Isomerase</keyword>
    <keyword evidence="5" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="6 17" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="15 17" id="KW-0539">Nucleus</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="Helicase ATP-binding" evidence="19">
        <location>
            <begin position="382"/>
            <end position="557"/>
        </location>
    </feature>
    <feature type="domain" description="Helicase C-terminal" evidence="20">
        <location>
            <begin position="583"/>
            <end position="730"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1"/>
            <end position="30"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="10"/>
            <end position="22"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005446"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022705"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022806"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023235"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00034617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="RuleBase" id="RU364117"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS51192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS51194"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007424.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="970" mass="109416" checksum="735241DAB6A730E9" modified="2019-05-08" version="1">SKKQYNGKQKITEGEKNSHSPKELAIGWDETPDYNGIEEDDYLDFIPPSPEEELLSSSSSVKNHLFSVMEDICKLVDSIPIHELKSLSCGNDLLQQRNLRRELLANHIDSNANKSDLGLVFASNWKSHPGHVTSMDDSIPVQSGNLNMSTFSTTKLQTLDTSGASDFSQYTALDTNITKNAGLSFSKMDVKERTSLSFHSEISSFNSSLCEKPRMEKDGSCFFSEMPVTSTTLKVENKSSSSLNNVGGDDSLGARDIDFDLDNFVLDDDFDERWDNPRCVSADEKTSTPLYQPIREGEPAKSLLKSFSTVKETSPIYNPVPKPNILLSAKSCSDPLLKKSPSKNPALARFRGCTFSHSNDMMRIFHKKFGLHHFRTNQLEAINASLLGEDCFILMPTGGGKSLCYQLPACLSSGVTIVISPLRSLILDQVEKLMTLDISATYLTGDRTDTEASKIYIQLSKKDPQIKLLYVTPEKVCSSGRLISALKNLYERDLLARFVIDEAHCVSQWGHDFRQDYKRLNMLRQKFPSVPMMALTATANPRVQKDILNQLEMLKPQVFTMSFNRHNLKYDVLPKKPKRVALDCLEWIKKYHPHDSGIIYCLSRQDCNKTADILQEKGLAALAYHAGLSDTVRDLVQRKWINQDGCQVICATIAFGMGIDKPDVRYVIHASLPKSVEGYYQESGRAGRDGEVSHCLLFYSYSDVTRLRRLIQMEKDGNSQTRKTHFDNLYSMVHYCENVHECRRIQLLAYFGETGFNLNFCKEHPEVICDNCCRKKDYKSRNVTDEVKSIIRFVQEHCGQMGRRNSGSGRYTLNMMIDIFLGTKSARIQSGIFGKGAAYSRHNVERLFRKLVLDKILEEELYITAAEQAVAYVTVGEKAQAVLNGYLQVEFYETENASSIRKQRASVTKMSQREEMVKKCLVELTDVCKTLGKVFDVHYFNIFNTATLKRIAGNSRIHPGVGLHWGGGTL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ24</accession>
    <name>A0A452GZ24_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">POF1B helix-loop-helix domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GZ24"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008554.1">
        <property type="protein sequence ID" value="ENSGAGP00000007405.1"/>
        <property type="gene ID" value="ENSGAGG00000005876.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005884">
        <property type="term" value="C:actin filament"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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    <sequence length="390" mass="45681" checksum="6AB9CA4106A625D5" modified="2019-05-08" version="1">QWYRTEGGWQHQDGNVVYERVRTYSCPGGKRVQVLELPPCSPRPVSPLENAPGHQHVIWSRDPYCPGNNVIYEKTIRKYELLNPEQVHSLQETNGQLDSRYFGELLAEVNRKSNDLYSCLLQHVEKIGRRTQDNESTSQVEDIEDLIPKGLSELTKQQIRYLLQMRVTSDKSLRLVLSTFGSLREELCHLQDDLGKLETDKMLLEKDLAFKESQVKEYETLLASVRENNRQQQQGLRDSASKCRSLEEQIISLRYAEGEKDCRLKELEYCKRALEQEIQNLRMQTCSNPVIQTTTDELSSRYVEMINNLREDKDREIRNLRSQINQFQQDISRRDGSNSDFQMRLHELTSMLEEKETFIKQQQEELFRLKQERLSCNQSTGVTKSIITKK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
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    <name>A0A452GZ26_9SAUR</name>
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        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
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            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
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        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
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    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
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            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
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        <citation type="submission" date="2025-09" db="UniProtKB">
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                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
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        <text evidence="8">Interacts with CPAP, LIP8 and ZNF521.</text>
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        <subcellularLocation>
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        <property type="gene ID" value="ENSGAGG00000005878.1"/>
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    </dbReference>
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            <end position="3339"/>
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            <end position="3563"/>
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    </feature>
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        <source>
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    </evidence>
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    </evidence>
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        <source>
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        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
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        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="3718" mass="421442" checksum="0F0B81790FBF768D" modified="2019-05-08" version="1">MSSASNSLAREFLIDVNRLCNAVVQRAEAREEEEEETHMAALGQYLVQGRGFTLLTTLNSNMDQELTCREELLTLLLSLLPLVWKIPVQEEKAINFNLPFTVDVCLTKETNSNSLRSAHEKQNLEGIVHPSFQTPVKLNSSFRKIRRQRKITHRYSVRDARKSQLSTSDSEGNSDNKTTIMIKHRRSQILQPFVTHLPRDDYHAEVVRSFNPSPISLENSRQIIPTQESKSEVLNESAAAAVVSSMDNSPFDLCHVLLSLLEKVCKFDITLNHNSALAATIVPSLTEFLSGFGDCCNVSGDAESEVVSAGWTEEPVALVQRMLFRTVLHLLSIDITNAETMPDNLKKNLTDLLRAALKIRTCLEKQSDPFVPRHKKTLQEVQDNFVFSKYRHRVLLLPELLEGVLQILICCLQSAASNPFYFSQAVDLVHEFIQHQGFKLLEITVLQMEWLVSKDEEITPEASEHLKALISSIMKIISTVKKVKSEQLHQSMCTRKRHRRCEYSHFMHHHRDLSGLPVSAFKNQVSRNPFEEVVDGEVHYPERCCCIAVCAHQCFHLLQQVSLSSTCAQILSGVHNVGICCCMDPKSVITPLLQASKLPVLKNFQQHILSILNKFILDQLGGAEVSQKIIQAACNICTIDCDQLAELEDTLQSNSTDIGLASNFSCPSYRFQGILPSSGSEDMLLKWDALEAYQDFAFEEDKLRSVQIASHICHLIQKGNVVVQWKLYNYIYNPVLQRGVELAHRSQQLGITAVCNHVCSCHSLCLPQEVLQIYLQTLPALFKSRIIQNLFLSCNGVNQMTELNYLDSVRSHSLKVFETLIFCLGDQQKDQAMPELEGMDNEQKESILDLVTSFRSQQMISDVPQSLSKFYAGLRKAYPKKKKCVSQEVHVNLIHLFLCVAFLCVSKEEECDRDSANDSEDTSGYDSTASEPLSHKLPCLSLESLTLPSPEHIHRAADIWSMCRWIYLCSSVFQRQFHRLGGFEVCHRLIIMIIQKLAKNKEKEQEKKERPLSENNKCLNASPLNQPVFKDDSYYLTVNKQLTQSELNSLDENPDASGQPVSECISCDHSLKVEQISVAMVELEELKKSVKQENEWSLQSIRLLEALLTICLHSANSTQQKAEVELLHQSVCLEDVLLEMRDQLAQSRAIETCLAKPLLDSLLRVALGNYSADLDHSEEKTEKILHTEKASVPQPEDLSEEAEESQCCSFKLFAEEEGYEADSESNPDDSESKDEGTDTKREAECMVPSSYSSDLTENISQGELIYPEICMLELKLLSAGNPSLDVLAHVFQSFLKVIRQKKKNATFLMQQGAVKSLLGGFLNILIQKDSSFHACQIVLVELLVSLMSSRTCAEELTLLLRIFLEETPCTEILLKGILKIVETNIYMSPLQYLSFPLLKSPNLNSSSSQKSTSALNSKTVGLLKRAKFSQNKKEADGENLTHQLLSSWHVAPIHLPLVGQNCWPHMSEGFSVSLWFHVECVQETESPTEKGKKMKKRSKPLIVRESSFDGTEGSKSVRMEYLNLGDKLVEDGCIHIISLGSKALMIQVWADLNTGAFIFRMCMDPNDEMKAGLLAQAESPENIFLVGKWEHLAITYLQQPEGKKNIHGELLLWVSGQRKCEISLDYTLPRKTTLSDSNKTFCMIGHCTSSQEEPLQLAGKWDLGNLLLFNGAKIGPEEAFYLYACGPDFTAIMPCKYGKTMTDYSKYITKDILQCERIKEFFMKKKEADVGPLIESLAVVYTTCCPAQYTIYEPVIRLKGQVKTVPSQRPFSSKEVQSNFLDPYHLKMLQPVEYQTLQGILHEIGGTGAFVFLFARVIDITNCEETQALALQVILSLTKYNQQRIQEMENCNGYSMIHQVLIKPKCIVGFHILKTLLEGCFSDEILNINESGQFMLDVDSTAVIQDIKLLEDLLLDWKIWSKAQQGVWETLLAALEILIRANHHQQMFNIKQLLKARVVHHFLLTCQVLQEQKEGHLTSLPREVCRSFVKIIEEVLGSPPDLELLTLIFNFLLAVHPPTNTYVCHNPTNFYFSLHIDGKIFQEKVQSIKYLRNSSIGGKSVDSPSLTTSPAGFTTSPLGGSIISKIIIPQMMISQALQRRCKSLPASPTSSPQVHPKRLPGSLGRSIDNLQEVWESDSVPQEKNIFVESSETLKRIQEELFVSSCESAKTVCDSELTSAEALVDISKEELELSENKELNTELKSNENGAAVESLLRRPDNLKGLPSFQKSQNNFAGLGLAFSAQSGSSSVACWPSLADRSALPEDWESLTFSSVSEQTFKRSERPDYRSTEDCLVLICCGLYDLLRGVLLILPDIMLEDVIDKLIQPDTLIVLVNHPSPLIQERVIKLLDAYFNRATREQKEQFLKNRGFSLLANQLYLHQGTQELVECFLEMLFGRSVGLDEDIPLNDYKLLVCDIQQLLVAVTIHACSSLGSQYFRVIEDLIVLLGHLQNNKNEKTQNMAIALQFRVLQAAIELIKATANQDSQDLANSLRLPATHHHAMFQKRKSIADSRKFSRAQSDTLVMKMRLVASDELNMMMQRRMSQENPIRATENEFIQRLQRLTVLAVNRFIYQEPTEDCFDVLGTPETTIQCRLSTSQLEASEEEDQRELPSNTNSFLKEMFKMMIEGIKVSIATSRISTPRQQWRRILWSCKDTFRMQLGRLLVHILSPAQPLQERKQALEIVCETNHQEIIRDCLSVPLQHGPKLALYLYELMHDHKDELTREEQVSSGVFMNALKFCGYKCISPSAPPKPDLLNAMKEDRKKFEAEECVNKTVWEKTVINNQQNLSQRLDAKSKDISKIAADITQAVSLSQGMERKKVIQHIRGMYKTDLSASRHWQELVQQLTHDRTLWYDPVYYPTSWQLDPTEGPNRERRRLRRCYLTIPNKYLLRDRQKQEEVIKPPLSYLFEDKTHSSHSSTVKDKAASENIRVNRRCINVAPSRETAGELLLGKYGMYFVEDNASDTIESSSFHGETEPASFSWTYEEIKEVHKRWWQLRDNAVEIFLTNGRTLLLAFDNTKVRDDVYHNILMNNLPNLLEYGNITALTHLWYTGQITNFEYLTHLNKHAGRSFNDLMQYPVFPFILSDYTSETLDLNDPSVYRNLAKPIAVQSKEKEDRYVDTYKYLEEEYRKGAREDDPMPPVQPYHYGSHYSNSGTVLHFLVRMPPFTKMFLAYQDQSFDIPDRTFHSTHTTWRLSSYESMTDVKELIPEFFYLPEFLVNREGFDFGLRQNGERVNHVNLPPWARNDPRLFILIHRQALESDHVSQTICNWIDLVFGYKQKGKASVQAINVFHPATYFGMDVSAVEDPVQRRALETMIKTYGQTPRQLFHTAHVSRPGSRLTIEGELPAAMGLLVQFAFRETREHAKEIVYPSPLPWIKGLKWGEYVGSPSAPDPVVCFSQPHGERFGSLQALPTRAICGLSQKFCLLMVYSKEQGVRSMHSTDIQWSAILSWGYADNILRLKSKQSEPPINFIQSSQLHQVTSCAWVPDSCQLFTGSKSGVITAYMNRFSSNMHSEIEMESQVHLYGHTAEITSLFVCKPYSIMISVSKDGTCIIWDLNRLCYVQSLAGHKSPVTAVSASEMTGDIATVCDSVGGGSDLRLWTVNGDLVGHVHCREIICSVAFSNQPEGVSINVIAGGLENGVVRLWSTWDLKPVREITFPKSNKPIVSLTFSCDGHHLYTANSEGTVIAWCRKDQQRLKLPMFYSFLSSYAAG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ28</accession>
    <name>A0A452GZ28_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">TRPM SLOG domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GZ28"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008691.1">
        <property type="protein sequence ID" value="ENSGAGP00000007527.1"/>
        <property type="gene ID" value="ENSGAGG00000006018.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005227">
        <property type="term" value="F:calcium-activated cation channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0099604">
        <property type="term" value="F:ligand-gated calcium channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050927">
        <property type="entry name" value="TRPM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041491">
        <property type="entry name" value="TRPM_SLOG"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13800:SF6">
        <property type="entry name" value="TRANSIENT RECEPTOR POTENTIAL CATION CHANNEL SUBFAMILY M MEMBER 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13800">
        <property type="entry name" value="TRANSIENT RECEPTOR POTENTIAL CATION CHANNEL, SUBFAMILY M, MEMBER 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18139">
        <property type="entry name" value="LSDAT_euk"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="TRPM SLOG" evidence="2">
        <location>
            <begin position="41"/>
            <end position="156"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="152"/>
            <end position="183"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="152"/>
            <end position="168"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF18139"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007527.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="183" mass="18962" checksum="159A7DC745600F7D" modified="2019-05-08" version="1">VGDAFGTAIVSKWDSDQHTTEGPTDAFGELEFVGAGGKPSKFIRLSDSSDPALAYNLVTEHWKISPPNLVVSVVGGDGEAPVRGRVTFSACATVRAPTGAWIMTGGLQAGIGQYVGEAVRDHGTASTSPCARVVAMGIAPWGVVANREGTLREELGVPEGQREGDPKGRSGRTLGEGDPRESS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GZ29</accession>
    <name>A0A452GZ29_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">ATP synthase peripheral stalk subunit OSCP, mitochondrial</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="14">ATP synthase subunit O</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="11">Oligomycin sensitivity conferral protein</fullName>
        </alternativeName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="12">Component of the ATP synthase complex composed at least of ATP5F1A/subunit alpha, ATP5F1B/subunit beta, ATP5MC1/subunit c (homooctomer), MT-ATP6/subunit a, MT-ATP8/subunit 8, ATP5ME/subunit e, ATP5MF/subunit f, ATP5MG/subunit g, ATP5MK/subunit k, ATP5MJ/subunit j, ATP5F1C/subunit gamma, ATP5F1D/subunit delta, ATP5F1E/subunit epsilon, ATP5PF/subunit F6, ATP5PB/subunit b, ATP5PD/subunit d, ATP5PO/subunit OSCP. ATP synthase complex consists of a soluble F(1) head domain (subunits alpha(3) and beta(3)) - the catalytic core - and a membrane F(0) domain - the membrane proton channel (subunits c, a, 8, e, f, g, k and j). These two domains are linked by a central stalk (subunits gamma, delta, and epsilon) rotating inside the F1 region and a stationary peripheral stalk (subunits F6, b, d, and OSCP).</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Mitochondrion inner membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the ATPase delta chain family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GZ29"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008562.1">
        <property type="protein sequence ID" value="ENSGAGP00000007413.1"/>
        <property type="gene ID" value="ENSGAGG00000005893.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005743">
        <property type="term" value="C:mitochondrial inner membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045259">
        <property type="term" value="C:proton-transporting ATP synthase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046933">
        <property type="term" value="F:proton-transporting ATP synthase activity, rotational mechanism"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.520.20:FF:000002">
        <property type="entry name" value="ATP synthase subunit O, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.520.20">
        <property type="entry name" value="N-terminal domain of the delta subunit of the F1F0-ATP synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026015">
        <property type="entry name" value="ATP_synth_OSCP/delta_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000711">
        <property type="entry name" value="ATPase_OSCP/dsu"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01145">
        <property type="entry name" value="ATP_synt_delta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11910">
        <property type="entry name" value="ATP SYNTHASE DELTA CHAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00213">
        <property type="entry name" value="OSCP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00125">
        <property type="entry name" value="ATPASEDELTA"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47928">
        <property type="entry name" value="N-terminal domain of the delta subunit of the F1F0-ATP synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0066">ATP synthesis</keyword>
    <keyword evidence="4" id="KW-0375">Hydrogen ion transport</keyword>
    <keyword evidence="7" id="KW-0406">Ion transport</keyword>
    <keyword evidence="9" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="5" id="KW-0999">Mitochondrion inner membrane</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0809">Transit peptide</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007046"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022781"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022792"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023310"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00033369"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00064647"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00073432"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00078525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007413.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="171" mass="19373" checksum="C520DC3BFA1CCB4C" modified="2019-05-08" version="1">MLQTVTNLTSFFFKPPIQVYGLEGRYATALYSAASKQKKLDQVEKELSRVSTLLKDPKLSSVVVNPHVKSVIKQKTINDALVKEKLSPITINFMKLLAENGRLQQTPDVISAFGKIMSAYRGEVLCSVTTANKMSSQHKVLLLHMEVRWLSRGKTLVRLFELRCELLAFFR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKK0</accession>
    <name>A0A452HKK0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Type II GnRH receptor</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKK0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017659.1">
        <property type="protein sequence ID" value="ENSGAGP00000015459.1"/>
        <property type="gene ID" value="ENSGAGG00000011590.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004930">
        <property type="term" value="F:G protein-coupled receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042277">
        <property type="term" value="F:peptide binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016500">
        <property type="term" value="F:protein-hormone receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032870">
        <property type="term" value="P:cellular response to hormone stimulus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1070.10:FF:000199">
        <property type="entry name" value="Gonadotropin-releasing hormone II receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1070.10">
        <property type="entry name" value="Rhodopsin 7-helix transmembrane proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000276">
        <property type="entry name" value="GPCR_Rhodpsn"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017452">
        <property type="entry name" value="GPCR_Rhodpsn_7TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001658">
        <property type="entry name" value="GphnRH_fam_rcpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24241:SF69">
        <property type="entry name" value="GONADOTROPIN-RELEASING HORMONE II RECEPTOR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24241">
        <property type="entry name" value="NEUROPEPTIDE RECEPTOR-RELATED G-PROTEIN COUPLED RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00001">
        <property type="entry name" value="7tm_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00529">
        <property type="entry name" value="GNADOTRPHINR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00237">
        <property type="entry name" value="GPCRRHODOPSN"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81321">
        <property type="entry name" value="Family A G protein-coupled receptor-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50262">
        <property type="entry name" value="G_PROTEIN_RECEP_F1_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6" id="KW-0297">G-protein coupled receptor</keyword>
    <keyword evidence="7 12" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="9" id="KW-0675">Receptor</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0807">Transducer</keyword>
    <keyword evidence="4 12" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 12" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="38"/>
            <end position="60"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="72"/>
            <end position="91"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="111"/>
            <end position="131"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="152"/>
            <end position="171"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="202"/>
            <end position="225"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="259"/>
            <end position="283"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="295"/>
            <end position="318"/>
        </location>
    </feature>
    <feature type="domain" description="G-protein coupled receptors family 1 profile" evidence="13">
        <location>
            <begin position="51"/>
            <end position="315"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023224"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00082552"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50262"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015459.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="388" mass="42379" checksum="2DC822F6F2ACE6C1" modified="2019-05-08" version="1">MPKNCFCASSPEPQRAQQTSNSGALQLPTFSPAAKARVILTFVLLTLSAAGNLAVLWAGAGGRRGKLSHVRVLLLHLAAADLLVTFVVMPLDAGWNLTVQWQAGDLACRLLMYLKLLAMYASAFITVVISLDRQAAILRPLAIAHAHRRNRLMLGPALLSFAPLCFQLFLFHTVTIRSPQNFTQCTTQGSFPQRWHETAYNMLGFACLFLLPLLIMVCCYARILLEISPLLPVALVSKEVPLRRSTTNIPRARLRVLKLSLVIVSSFVVCWTPYYLLGLWYWFCPRAMEETVSPSLAHVLFIFGLLNACLDPLTYGLFTIPFHRGLGHCCRGGHGAGSEPCSPPTGSLRCSATSLHAKRGATATPVHETPAAQSYLGNGTSSVDSSYL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKK7</accession>
    <name>A0A452HKK7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">C2H2-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="10">Belongs to the Ikaros C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKK7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017663.1">
        <property type="protein sequence ID" value="ENSGAGP00000015463.1"/>
        <property type="gene ID" value="ENSGAGG00000011670.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003700">
        <property type="term" value="F:DNA-binding transcription factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000073">
        <property type="entry name" value="IKAROS family zinc finger 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000265">
        <property type="entry name" value="IKAROS family zinc finger 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000525">
        <property type="entry name" value="IKAROS family zinc finger 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000080">
        <property type="entry name" value="IKAROS family zinc finger 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050589">
        <property type="entry name" value="Ikaros_C2H2-ZF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24404:SF36">
        <property type="entry name" value="DNA-BINDING PROTEIN IKAROS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24404">
        <property type="entry name" value="ZINC FINGER PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="2" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="8" id="KW-0804">Transcription</keyword>
    <keyword evidence="6" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <keyword evidence="4 11" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="56"/>
            <end position="83"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="84"/>
            <end position="111"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="112"/>
            <end position="140"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1"/>
            <end position="49"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="293"/>
            <end position="316"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00038390"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015463.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="429" mass="48242" checksum="AE6DCB972A496755" modified="2019-05-08" version="1">METDEAQDMSQVSGKESPPISDVPDDADEPMPVPEDLSTSTGGQQKNDRVLGERPFHCTQCGASFTQKGNLLRHIKLHSGEKPFKCHLCNYACRRRDALTGHLRTHSVGKPHKCGYCGRSYKQRSSLEEHKERCHNYLQTMSLSGSLYPVIKEETNQSEMAEDLCKIGSERSLVLDRLASNVAKRKSSMPQKFVGEKCLSDVPYDATTNYEKESEMMQTHVIDQAINNAISYLGAESLRPLVQTPPGSSEAAPIISSMYQLHKPLGDNHSRSNHTAQDTAVENLLLLSKAKSISSERDASPSNSCQDSTDTESNNEERSGLIYLTNHIVPHARNGLSIKEEQRQYEVLRAGTDNSHDAFKVITSNGEQVKVYKCEHCRVLFLDHVMYTIHMGCHGFRDPFECNMCGYHSQDRYEFSSHITRGEHRFHMS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKK9</accession>
    <name>A0A452HKK9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="9">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="caution">
        <text evidence="6">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKK9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017675.1">
        <property type="protein sequence ID" value="ENSGAGP00000015473.1"/>
        <property type="gene ID" value="ENSGAGG00000011220.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005080">
        <property type="term" value="F:protein kinase C binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045778">
        <property type="term" value="P:positive regulation of ossification"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045667">
        <property type="term" value="P:regulation of osteoblast differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00054">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000120">
        <property type="entry name" value="Protein kinase C-binding protein NELL1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000102">
        <property type="entry name" value="Protein kinase C-binding protein NELL2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000111">
        <property type="entry name" value="Protein kinase C-binding protein NELL2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.200">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.20.200.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001881">
        <property type="entry name" value="EGF-like_Ca-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000152">
        <property type="entry name" value="EGF-type_Asp/Asn_hydroxyl_site"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018097">
        <property type="entry name" value="EGF_Ca-bd_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009030">
        <property type="entry name" value="Growth_fac_rcpt_cys_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024731">
        <property type="entry name" value="NELL2-like_EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049883">
        <property type="entry name" value="NOTCH1_EGF-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051586">
        <property type="entry name" value="PKC-binding_NELL"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048287">
        <property type="entry name" value="TSPN-like_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001007">
        <property type="entry name" value="VWF_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24042">
        <property type="entry name" value="NEL HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24042:SF2">
        <property type="entry name" value="PROTEIN KINASE C-BINDING PROTEIN NELL1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12947">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07645">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00093">
        <property type="entry name" value="VWC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00179">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00210">
        <property type="entry name" value="TSPN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00214">
        <property type="entry name" value="VWC"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57603">
        <property type="entry name" value="FnI-like domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57184">
        <property type="entry name" value="Growth factor receptor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00010">
        <property type="entry name" value="ASX_HYDROXYL"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01186">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01187">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01208">
        <property type="entry name" value="VWFC_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50184">
        <property type="entry name" value="VWFC_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="1 6" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="5" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0732">Signal</keyword>
    <feature type="domain" description="VWFC" evidence="8">
        <location>
            <begin position="180"/>
            <end position="241"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="7">
        <location>
            <begin position="343"/>
            <end position="384"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="7">
        <location>
            <begin position="385"/>
            <end position="425"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50184"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015473.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="437" mass="48807" checksum="D6CC9ED90D4CBD4F" modified="2019-05-08" version="1">TELDLVNTTIGVTQVSGLHASKAFLFQDVEREIHAAPHVSEKLIQLFRNKSEFTFLATLQQRASTSGVVLSIRELEHRIYERVIDPPEMNLTPGSSLWLGQRNRKHGFFKGIIQDVKVIFMPNGYITQCPNLNRTCPTCSDFLSLVQGIMDLQELLAKMTAKLNYAETRLSQLENCHCEKTCQVNGVIYRDKDSWVEDDHCRNCICKSGVVECRRMSCPPLNCAPDTLPVHMGGQCCKVCRSKCIFGGRVLAEGQRVLTKNCRECRNGVLVKVTETCPPLNCSEKDHILPENQCCSVCRGHNFCAEGHRCGENSECKNWNTKATCECKTGYISNQGDSAYCKDIDECAAKMHYCHANTVCVNLPGSYRCDCVPGYIRVDDFSCTEHDECGSGQHNCDANAICTNTVRGYSCTCKPGYVGNGTICRGRSTWTITLRLR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKL1</accession>
    <name>A0A452HKL1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Type II GnRH receptor</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKL1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017666.1">
        <property type="protein sequence ID" value="ENSGAGP00000015465.1"/>
        <property type="gene ID" value="ENSGAGG00000011590.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004930">
        <property type="term" value="F:G protein-coupled receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042277">
        <property type="term" value="F:peptide binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016500">
        <property type="term" value="F:protein-hormone receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032870">
        <property type="term" value="P:cellular response to hormone stimulus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd15383">
        <property type="entry name" value="7tmA_GnRHR_vertebrate"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1070.10:FF:000199">
        <property type="entry name" value="Gonadotropin-releasing hormone II receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1070.10">
        <property type="entry name" value="Rhodopsin 7-helix transmembrane proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000276">
        <property type="entry name" value="GPCR_Rhodpsn"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017452">
        <property type="entry name" value="GPCR_Rhodpsn_7TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001658">
        <property type="entry name" value="GphnRH_fam_rcpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24241:SF69">
        <property type="entry name" value="GONADOTROPIN-RELEASING HORMONE II RECEPTOR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24241">
        <property type="entry name" value="NEUROPEPTIDE RECEPTOR-RELATED G-PROTEIN COUPLED RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00001">
        <property type="entry name" value="7tm_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00529">
        <property type="entry name" value="GNADOTRPHINR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00237">
        <property type="entry name" value="GPCRRHODOPSN"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81321">
        <property type="entry name" value="Family A G protein-coupled receptor-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50262">
        <property type="entry name" value="G_PROTEIN_RECEP_F1_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6" id="KW-0297">G-protein coupled receptor</keyword>
    <keyword evidence="7 12" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="9" id="KW-0675">Receptor</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0807">Transducer</keyword>
    <keyword evidence="4 12" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 12" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="55"/>
            <end position="77"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="89"/>
            <end position="108"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="128"/>
            <end position="148"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="169"/>
            <end position="189"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="220"/>
            <end position="243"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="277"/>
            <end position="301"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="313"/>
            <end position="336"/>
        </location>
    </feature>
    <feature type="domain" description="G-protein coupled receptors family 1 profile" evidence="13">
        <location>
            <begin position="68"/>
            <end position="333"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023224"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00082552"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50262"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015465.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="406" mass="44205" checksum="BB2C6D62777BBD62" modified="2019-05-08" version="1">GVGVGPGLQLPLLLRPLHPAAQGLPVLPCGWGPWPNGTEGALQLPTFSPAAKARVILTFVLLTLSAAGNLAVLWAGAGGRRGKLSHVRVLLLHLAAADLLVTFVVMPLDAGWNLTVQWQAGDLACRLLMYLKLLAMYASAFITVVISLDRQAAILRPLAIAHAHRRNRLMLYAAWLLSAGLSLPQLFLFHTVTIRSPQNFTQCTTQGSFPQRWHETAYNMLGFACLFLLPLLIMVCCYARILLEISRRMGTSLFSKEVPLRRSTTNIPRARLRVLKLSLVIVSSFVVCWTPYYLLGLWYWFCPRAMEETVSPSLAHVLFIFGLLNACLDPLTYGLFTIPFHRGLGHCCRGGHGAGSEPCSPPTGSLRCSATSLHAKRGATATPVHETPAAQSYLGNGTSSVDSSYL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKL3</accession>
    <name>A0A452HKL3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Slit homolog 2 protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="15">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000017677.1">
        <property type="protein sequence ID" value="ENSGAGP00000015475.1"/>
        <property type="gene ID" value="ENSGAGG00000011388.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048495">
        <property type="term" value="F:Roundabout binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072359">
        <property type="term" value="P:circulatory system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060560">
        <property type="term" value="P:developmental growth involved in morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060562">
        <property type="term" value="P:epithelial tube morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001822">
        <property type="term" value="P:kidney development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050919">
        <property type="term" value="P:negative chemotaxis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030308">
        <property type="term" value="P:negative regulation of cell growth"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010605">
        <property type="term" value="P:negative regulation of macromolecule metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010977">
        <property type="term" value="P:negative regulation of neuron projection development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0021772">
        <property type="term" value="P:olfactory bulb development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050770">
        <property type="term" value="P:regulation of axonogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032535">
        <property type="term" value="P:regulation of cellular component size"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050767">
        <property type="term" value="P:regulation of neurogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031290">
        <property type="term" value="P:retinal ganglion cell axon guidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0022029">
        <property type="term" value="P:telencephalon cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd00054">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="CDD" id="cd00110">
        <property type="entry name" value="LamG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000045">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000053">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000054">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000062">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000063">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000099">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000289">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.200:FF:000013">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000002">
        <property type="entry name" value="Slit guidance ligand 2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000419">
        <property type="entry name" value="Slit homolog 2 (Drosophila)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.200">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000483">
        <property type="entry name" value="Cys-rich_flank_reg_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006207">
        <property type="entry name" value="Cys_knot_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001881">
        <property type="entry name" value="EGF-like_Ca-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000152">
        <property type="entry name" value="EGF-type_Asp/Asn_hydroxyl_site"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018097">
        <property type="entry name" value="EGF_Ca-bd_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003645">
        <property type="entry name" value="Fol_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001791">
        <property type="entry name" value="Laminin_G"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001611">
        <property type="entry name" value="Leu-rich_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003591">
        <property type="entry name" value="Leu-rich_rpt_typical-subtyp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000372">
        <property type="entry name" value="LRRNT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051355">
        <property type="entry name" value="Notch/Slit_guidance"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45836">
        <property type="entry name" value="SLIT HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45836:SF2">
        <property type="entry name" value="SLIT HOMOLOG 2 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00008">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00054">
        <property type="entry name" value="Laminin_G_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00560">
        <property type="entry name" value="LRR_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13855">
        <property type="entry name" value="LRR_8"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01463">
        <property type="entry name" value="LRRCT"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01462">
        <property type="entry name" value="LRRNT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00041">
        <property type="entry name" value="CT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="SMART" id="SM00179">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SMART" id="SM00274">
        <property type="entry name" value="FOLN"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00282">
        <property type="entry name" value="LamG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00368">
        <property type="entry name" value="LRR_RI"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00365">
        <property type="entry name" value="LRR_SD22"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SMART" id="SM00369">
        <property type="entry name" value="LRR_TYP"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="SMART" id="SM00082">
        <property type="entry name" value="LRRCT"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00013">
        <property type="entry name" value="LRRNT"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52058">
        <property type="entry name" value="L domain-like"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00010">
        <property type="entry name" value="ASX_HYDROXYL"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01185">
        <property type="entry name" value="CTCK_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01225">
        <property type="entry name" value="CTCK_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01186">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01187">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50025">
        <property type="entry name" value="LAM_G_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51450">
        <property type="entry name" value="LRR"/>
        <property type="match status" value="5"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0145">Chemotaxis</keyword>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="10" id="KW-0221">Differentiation</keyword>
    <keyword evidence="12 15" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5 15" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7" id="KW-0358">Heparin-binding</keyword>
    <keyword evidence="6" id="KW-0433">Leucine-rich repeat</keyword>
    <keyword evidence="11" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="8" id="KW-0732">Signal</keyword>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="810"/>
            <end position="845"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="847"/>
            <end position="886"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="888"/>
            <end position="924"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="926"/>
            <end position="964"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="966"/>
            <end position="1002"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="1011"/>
            <end position="1047"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin G" evidence="17">
        <location>
            <begin position="1050"/>
            <end position="1223"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="1227"/>
            <end position="1258"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="1261"/>
            <end position="1297"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="18">
        <location>
            <begin position="1302"/>
            <end position="1338"/>
        </location>
    </feature>
    <feature type="domain" description="CTCK" evidence="16">
        <location>
            <begin position="1343"/>
            <end position="1418"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="835"/>
            <end position="844"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="876"/>
            <end position="885"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="914"/>
            <end position="923"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="954"/>
            <end position="963"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="992"/>
            <end position="1001"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="1037"/>
            <end position="1046"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="1248"/>
            <end position="1257"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="1265"/>
            <end position="1275"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="1287"/>
            <end position="1296"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="1306"/>
            <end position="1316"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="1328"/>
            <end position="1337"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022500"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022614"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022674"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00074955"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS01225"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50025"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015475.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1419" mass="157698" checksum="455156AFD63E92F4" modified="2019-05-08" version="1">RVIGACFVGLARPATPRPLLTLLRSPAWRSAHFHLIEIHCEILFLHDILSQKFFHTDKICTTTFKFPFIFFFCSDLSENQIQAIPRKAFRGAVDIKNLQLDYNQISCIEDGAFRALRDLEVLTLNNNNITRLSVASFNHMPKLRTFRLHSNNLYCDCHLAWLSDWLRQRPRVGLYTQCMGPSHLRGHNVAEVQKREFVCSGHQSFMAPSCSVLHCPTACTCSNNIVDCRGKGLTEIPTNLPETITEIRLEQNSIKVIPPGAFSPYKKLRRIDLSNNQISETAPDAFQGLRSLNSLVLYGNKITELPKGLFEGLFSLQLLLLNANKINCLRVDAFQDLHNLNLLSLYDNKLQTIAKGTFSPLRSIQTLHLAQNPFICDCHLKWLADYLHANPIETSGARCTSPRRLANKRIGQIKSKKFRCSAKEQYFIPGTEDYRSKLSGDCFADLACPEKCRCEGTTVDCSNQKLNKIPEHVPQYTAELRLNNNEFTVLEATGIFKKLPQLRKINLSNNKITDIEEGAFEGASGVNELLLTSNRLETIRHKMFKGLESLKTLMLRSNRISCIGNDSFTGLSSVRLLSLYDNQITTVAPGAFDTLHSLSTLNLLANPFNCNCHLAWLGDWLRKKRIVTGNPRCQKPYFLKEIPIQDVAIQDFTCDDGNDDNSCSPLSRCPAECTCLDTVVRCSNKGLKVLPKGIPKDVTELDLSNNRISTLSNQSFSNMTQLLTLFVFVIQSKHIKLLCFFPPSAIGANPLYCDCNMQWLSDWVKSEYKEPGIARCAGPGEMADKLLLTTPSKKFTCQGPVDVNILAKCSPCLSNPCKNDGTCNNDPVDFYTCTCPYGFKGQDCDIPIHACISNPCQHGGTCHLKEGEKDGFWCTCADGFEGENCEVNVDDCEDNDCENNSTCVDGINNYTCLCPPEYTGELCEEKLDFCAQDLNPCQHDSKCILTLKGYKCDCTPGYVGEHCDMDYDDCQDNKCKNGAQCTDAVNGYTCICPEGYSGLFCEFSPPVVLPRTSPCDNYECQNGAQCIVRVNEPICQCLSGYQGDKCEKLVSVNFVSKESYLQIPSSKVRPQTNITLQIATDEDSGILLYKGDRDHIAMELYRGRVRVSYDTGSYPASAIYSVETINDGNFHIVELLAMDQILSLSVDGGSPKIITNLSKQSTLNVDSPLYVGGMPVKNNVAALRQSPGQNGSSFHGCIRNLYINSELQDFRNVPLQVGILPGCEPCHKKVCVHGTCHAMSQSGFTCECEGGWTGPLCDQQTNDPCLGNKCVHGTCLPINAFSYSCKCLQGHGGVLCDEEEELFNPCQSIKCKHGKCRLSGLGKPYCECSSGYTGDSCDKEISCRGERIRDYYQKQQGYAACQTTKKVSRLECKGGCSTGQCCGPLRSKRRKYSFECTDGSSFVDEVEKVVKCGCTKCPS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKM5</accession>
    <name>A0A452HKM5_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="10">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="caution">
        <text evidence="7">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKM5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017685.1">
        <property type="protein sequence ID" value="ENSGAGP00000015483.1"/>
        <property type="gene ID" value="ENSGAGG00000011220.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005080">
        <property type="term" value="F:protein kinase C binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045778">
        <property type="term" value="P:positive regulation of ossification"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045667">
        <property type="term" value="P:regulation of osteoblast differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00054">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="CDD" id="cd00110">
        <property type="entry name" value="LamG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000120">
        <property type="entry name" value="Protein kinase C-binding protein NELL1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.200:FF:000015">
        <property type="entry name" value="protein kinase C-binding protein NELL1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000102">
        <property type="entry name" value="Protein kinase C-binding protein NELL2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000111">
        <property type="entry name" value="Protein kinase C-binding protein NELL2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.200">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.20.200.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001881">
        <property type="entry name" value="EGF-like_Ca-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000152">
        <property type="entry name" value="EGF-type_Asp/Asn_hydroxyl_site"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018097">
        <property type="entry name" value="EGF_Ca-bd_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009030">
        <property type="entry name" value="Growth_fac_rcpt_cys_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001791">
        <property type="entry name" value="Laminin_G"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024731">
        <property type="entry name" value="NELL2-like_EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049883">
        <property type="entry name" value="NOTCH1_EGF-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051586">
        <property type="entry name" value="PKC-binding_NELL"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048287">
        <property type="entry name" value="TSPN-like_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001007">
        <property type="entry name" value="VWF_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24042">
        <property type="entry name" value="NEL HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24042:SF2">
        <property type="entry name" value="PROTEIN KINASE C-BINDING PROTEIN NELL1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12947">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07645">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02210">
        <property type="entry name" value="Laminin_G_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00093">
        <property type="entry name" value="VWC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00179">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00282">
        <property type="entry name" value="LamG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00210">
        <property type="entry name" value="TSPN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00214">
        <property type="entry name" value="VWC"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57603">
        <property type="entry name" value="FnI-like domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57184">
        <property type="entry name" value="Growth factor receptor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00010">
        <property type="entry name" value="ASX_HYDROXYL"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01186">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01187">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01208">
        <property type="entry name" value="VWFC_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50184">
        <property type="entry name" value="VWFC_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0106">Calcium</keyword>
    <keyword evidence="5" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="1 7" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="6" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0732">Signal</keyword>
    <feature type="domain" description="VWFC" evidence="9">
        <location>
            <begin position="237"/>
            <end position="298"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="8">
        <location>
            <begin position="400"/>
            <end position="441"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="8">
        <location>
            <begin position="442"/>
            <end position="482"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50184"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015483.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="518" mass="57878" checksum="43E703A5B541CA52" modified="2019-05-08" version="1">TELDLVNTTIGVTQVSGLHASKAFLFQDVEREIHAAPHVSEKLIQLFRNKSEFTFLATLQQRASTSGVVLSIRELEHSYFELESSGLRDEIRYHYRFNGKARTEVFPYRMADGQWHKIALSLSASHLLLHVDCNRIYERVIDPPEMNLTPGSSLWLGQRNRKHGFFKGIIQDVKVIFMPNGYITQCPNLNRTCPTCSDFLSLVQGIMDLQELLAKMTAKLNYAETRLSQLENCHCEKTCQVNGVIYRDKDSWVEDDHCRNCICKSGVVECRRMSCPPLNCAPDTLPVHMGGQCCKVCRSKCIFGGRVLAEGQRVLTKNCRECRNGVLVKVTETCPPLNCSEKDHILPENQCCSVCRGHNFCAEGHRCGENSECKNWNTKATCECKTGYISNQGDSAYCKDIDECAAKMHYCHANTVCVNLPGSYRCDCVPGYIRVDDFSCTEHDECGSGQHNCDANAICTNTVRGYSCTCKPGYVGNGTICRGSACCYCVLGITLSHKMLLIAPSLILQFLASKTLLP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKM9</accession>
    <name>A0A452HKM9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Cytochrome P450 family 4 subfamily B member 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="9">
            <name>heme</name>
            <dbReference type="ChEBI" id="CHEBI:30413"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 10">Belongs to the cytochrome P450 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKM9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017692.1">
        <property type="protein sequence ID" value="ENSGAGP00000015488.1"/>
        <property type="gene ID" value="ENSGAGG00000011439.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0020037">
        <property type="term" value="F:heme binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005506">
        <property type="term" value="F:iron ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004497">
        <property type="term" value="F:monooxygenase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016705">
        <property type="term" value="F:oxidoreductase activity, acting on paired donors, with incorporation or reduction of molecular oxygen"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.630.10:FF:000005">
        <property type="entry name" value="cytochrome P450 4F22 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.630.10">
        <property type="entry name" value="Cytochrome P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001128">
        <property type="entry name" value="Cyt_P450"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017972">
        <property type="entry name" value="Cyt_P450_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002401">
        <property type="entry name" value="Cyt_P450_E_grp-I"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036396">
        <property type="entry name" value="Cyt_P450_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050196">
        <property type="entry name" value="Cytochrome_P450_Monoox"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24291">
        <property type="entry name" value="CYTOCHROME P450 FAMILY 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24291:SF201">
        <property type="entry name" value="CYTOCHROME P450, FAMILY 4, SUBFAMILY B, POLYPEPTIDE 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00067">
        <property type="entry name" value="p450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00463">
        <property type="entry name" value="EP450I"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00385">
        <property type="entry name" value="P450"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48264">
        <property type="entry name" value="Cytochrome P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00086">
        <property type="entry name" value="CYTOCHROME_P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="3 9 10" id="KW-0349">Heme</keyword>
    <keyword evidence="6 9 10" id="KW-0408">Iron</keyword>
    <keyword evidence="8 11" id="KW-0472">Membrane</keyword>
    <keyword evidence="4 9 10" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7 10" id="KW-0503">Monooxygenase</keyword>
    <keyword evidence="10" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="11" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="20"/>
            <end position="40"/>
        </location>
    </feature>
    <feature type="binding site" description="axial binding residue" evidence="9">
        <location>
            <position position="447"/>
        </location>
        <ligand>
            <name>heme</name>
            <dbReference type="ChEBI" id="CHEBI:30413"/>
        </ligand>
        <ligandPart>
            <name>Fe</name>
            <dbReference type="ChEBI" id="CHEBI:18248"/>
        </ligandPart>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004586"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023033"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PIRSR" id="PIRSR602401-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="RuleBase" id="RU000461"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015488.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="14">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="502" mass="58928" checksum="9E8B69F5DED923EB" modified="2019-05-08" version="1">MTSVVVRLITAIWCWLKKDVFRIVYLLTVLCLTYMVLKAIKLYQWRRKLNKALQSFPGPPVHWLFGNVHEDLDKLLEWSKKYARAFPVWFGKFVVFVSLNDADYAKAVCARGGKRWWKGLLTLQGPKWFQHRKLLTPGFHYGVLKPYVSLITKSTQVMLDKWEHLITQEKSVELFGHVSLMTLDSILKCAFSHESNCQMDRSNSYIKSVYEITYQINYRIWYLPYQIDFIYWLSRDGRRFQKACEVVHLQTDKVIKERKKSLKDERELEKIQKKRHLDFLDILLCAKDEKGDGLSDEDIRAEVDTFMFGGHDTTASGICWTLYCLAMYPEHQQKCREEIRELLGDRETIQWDDLGKMPNTTMCIKESMRIFPPVPFVGRQLSKPITFHDGRYLPEGTLVGINIYLIHRDPSLWKDPEVFDPMRFSPENSSKRHSHAFMPFSAGARNCIGQQFAMIEMKVAIALSLLRFEFSLNPAKIPRKIPQMVLQSKNGIHVHLKKIHWH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKN3</accession>
    <name>A0A452HKN3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Cytochrome P450 family 4 subfamily B member 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="5">
            <name>heme</name>
            <dbReference type="ChEBI" id="CHEBI:30413"/>
        </cofactor>
    </comment>
    <comment type="similarity">
        <text evidence="1 6">Belongs to the cytochrome P450 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKN3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017701.1">
        <property type="protein sequence ID" value="ENSGAGP00000015497.1"/>
        <property type="gene ID" value="ENSGAGG00000011439.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0020037">
        <property type="term" value="F:heme binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005506">
        <property type="term" value="F:iron ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004497">
        <property type="term" value="F:monooxygenase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016705">
        <property type="term" value="F:oxidoreductase activity, acting on paired donors, with incorporation or reduction of molecular oxygen"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.630.10">
        <property type="entry name" value="Cytochrome P450"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001128">
        <property type="entry name" value="Cyt_P450"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017972">
        <property type="entry name" value="Cyt_P450_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002403">
        <property type="entry name" value="Cyt_P450_E_grp-IV"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036396">
        <property type="entry name" value="Cyt_P450_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050196">
        <property type="entry name" value="Cytochrome_P450_Monoox"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24291">
        <property type="entry name" value="CYTOCHROME P450 FAMILY 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24291:SF201">
        <property type="entry name" value="CYTOCHROME P450, FAMILY 4, SUBFAMILY B, POLYPEPTIDE 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00067">
        <property type="entry name" value="p450"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00465">
        <property type="entry name" value="EP450IV"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00385">
        <property type="entry name" value="P450"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48264">
        <property type="entry name" value="Cytochrome P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00086">
        <property type="entry name" value="CYTOCHROME_P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 5 6" id="KW-0349">Heme</keyword>
    <keyword evidence="4 5 6" id="KW-0408">Iron</keyword>
    <keyword evidence="7" id="KW-0472">Membrane</keyword>
    <keyword evidence="3 5 6" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="6" id="KW-0503">Monooxygenase</keyword>
    <keyword evidence="6" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="20"/>
            <end position="40"/>
        </location>
    </feature>
    <feature type="binding site" description="axial binding residue" evidence="5">
        <location>
            <position position="332"/>
        </location>
        <ligand>
            <name>heme</name>
            <dbReference type="ChEBI" id="CHEBI:30413"/>
        </ligand>
        <ligandPart>
            <name>Fe</name>
            <dbReference type="ChEBI" id="CHEBI:18248"/>
        </ligandPart>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00010617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PIRSR" id="PIRSR602403-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="RuleBase" id="RU000461"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015497.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="10">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="387" mass="44718" checksum="18A2FE179955D697" modified="2019-05-08" version="1">MTSVVVRLITAIWCWLKKDVFRIVYLLTVLCLTYMVLKAIKLYQWRRKLNKALQSFPGPPVHWLFGNVHEDLDKLLEWSKKYARAFPVWFGKFVVFVSLNDADYAKAVCARGGKRWWKGLLTLQGPKWFQHRKLLTPGFHYGVLKPYVSLITKSTQVMLDKWEHLITQEKSVELFGHVSLMTLDSILKCAFSHESNCQMDRSVFYATCSMDSQHGYCLPVLLHREGGCHSISLACRDDLGKMPNTTMCIKESMRIFPPVPFVGRQLSKPITFHDGRYLPEGTLVGINIYLIHRDPSLWKDPEVFDPMRFSPENSSKRHSHAFMPFSAGARNCIGQQFAMIEMKVAIALSLLRFEFSLNPAKIPRKIPQMVLQSKNGIHVHLKKIHWH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKN4</accession>
    <name>A0A452HKN4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Latent-transforming growth factor beta-binding protein 1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="19">Latent-transforming growth factor beta-binding protein 2</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="20">Transforming growth factor beta-1-binding protein 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="25 26">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="26" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="25" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="25" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">Key regulator of transforming growth factor beta (TGFB1, TGFB2 and TGFB3) that controls TGF-beta activation by maintaining it in a latent state during storage in extracellular space. Associates specifically via disulfide bonds with the Latency-associated peptide (LAP), which is the regulatory chain of TGF-beta, and regulates integrin-dependent activation of TGF-beta. Outcompeted by LRRC32/GARP for binding to LAP regulatory chain of TGF-beta.</text>
    </comment>
    <comment type="function">
        <text evidence="14">May play an integral structural role in elastic-fiber architectural organization and/or assembly.</text>
    </comment>
    <comment type="subunit">
        <text evidence="16">Forms part of the large latent transforming growth factor beta precursor complex; removal is essential for activation of complex. Interacts with SDC4. Interacts (via C-terminal domain) with FBN1 (via N-terminal domain) in a Ca(+2)-dependent manner.</text>
    </comment>
    <comment type="subunit">
        <text evidence="17">Interacts with TGFB1; associates via disulfide bonds with the Latency-associated peptide chain (LAP) regulatory chain of TGFB1, leading to regulate activation of TGF-beta-1. LTBP1 does not bind directly to TGF-beta-1, the active chain of TGFB1. Interacts (via C-terminal domain) with FBN1 (via N-terminal domain). Interacts with FBN2. Interacts with ADAMTSL2. Interacts with EFEMP2.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="13">Belongs to the LTBP family.</text>
    </comment>
    <comment type="caution">
        <text evidence="21">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000017697.1">
        <property type="protein sequence ID" value="ENSGAGP00000015493.1"/>
        <property type="gene ID" value="ENSGAGG00000008780.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019838">
        <property type="term" value="F:growth factor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00054">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="12"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000005">
        <property type="entry name" value="Fibrillin 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000194">
        <property type="entry name" value="Latent transforming growth factor beta binding protein 2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000469">
        <property type="entry name" value="Latent transforming growth factor beta binding protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.290.10:FF:000004">
        <property type="entry name" value="latent-transforming growth factor beta-binding protein 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000019">
        <property type="entry name" value="latent-transforming growth factor beta-binding protein 1 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000046">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 1 isoform x2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000014">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.290.10:FF:000001">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 3 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.290.10:FF:000002">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 3 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000056">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 3 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000115">
        <property type="entry name" value="latent-transforming growth factor beta-binding protein 4 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000024">
        <property type="entry name" value="Putative latent-transforming growth factor beta-binding protein 2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000273">
        <property type="entry name" value="Putative latent-transforming growth factor beta-binding protein 2"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Laminin"/>
        <property type="match status" value="17"/>
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        <property type="entry name" value="TGF-beta binding (TB) domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026823">
        <property type="entry name" value="cEGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001881">
        <property type="entry name" value="EGF-like_Ca-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013032">
        <property type="entry name" value="EGF-like_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000152">
        <property type="entry name" value="EGF-type_Asp/Asn_hydroxyl_site"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018097">
        <property type="entry name" value="EGF_Ca-bd_CS"/>
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    <dbReference type="InterPro" id="IPR009030">
        <property type="entry name" value="Growth_fac_rcpt_cys_sf"/>
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    <dbReference type="InterPro" id="IPR049883">
        <property type="entry name" value="NOTCH1_EGF-like"/>
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    <dbReference type="InterPro" id="IPR017878">
        <property type="entry name" value="TB_dom"/>
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    <dbReference type="InterPro" id="IPR036773">
        <property type="entry name" value="TB_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24039:SF48">
        <property type="entry name" value="FIBRILLIN-2 ISOFORM X1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24039">
        <property type="entry name" value="FIBRILLIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12662">
        <property type="entry name" value="cEGF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07645">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="12"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12661">
        <property type="entry name" value="hEGF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00683">
        <property type="entry name" value="TB"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="18"/>
    </dbReference>
    <dbReference type="SMART" id="SM00179">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="16"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57184">
        <property type="entry name" value="Growth factor receptor domain"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57581">
        <property type="entry name" value="TB module/8-cys domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00010">
        <property type="entry name" value="ASX_HYDROXYL"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01186">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01187">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51364">
        <property type="entry name" value="TB"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9 21" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="4 21" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="3" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11" id="KW-0340">Growth factor binding</keyword>
    <keyword evidence="6" id="KW-0358">Heparin-binding</keyword>
    <keyword evidence="12" id="KW-0379">Hydroxylation</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="26" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="89"/>
            <end position="121"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="278"/>
            <end position="310"/>
        </location>
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    <feature type="domain" description="TB" evidence="24">
        <location>
            <begin position="425"/>
            <end position="468"/>
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    <feature type="domain" description="TB" evidence="24">
        <location>
            <begin position="545"/>
            <end position="597"/>
        </location>
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    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="701"/>
            <end position="742"/>
        </location>
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    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="743"/>
            <end position="779"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="786"/>
            <end position="825"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="866"/>
            <end position="906"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="991"/>
            <end position="1032"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="1079"/>
            <end position="1119"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="1120"/>
            <end position="1161"/>
        </location>
    </feature>
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        <location>
            <begin position="1224"/>
            <end position="1278"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="1300"/>
            <end position="1338"/>
        </location>
    </feature>
    <feature type="domain" description="TB" evidence="24">
        <location>
            <begin position="1398"/>
            <end position="1450"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="1587"/>
            <end position="1631"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="395"/>
            <end position="420"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="618"/>
            <end position="641"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="22">
        <location>
            <begin position="395"/>
            <end position="419"/>
        </location>
    </feature>
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            <begin position="93"/>
            <end position="103"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="21">
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            <begin position="111"/>
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        </location>
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        </location>
    </feature>
    <feature type="disulfide bond" evidence="21">
        <location>
            <begin position="300"/>
            <end position="309"/>
        </location>
    </feature>
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        <source>
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        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
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            <dbReference type="ARBA" id="ARBA00022553"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022674"/>
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    </evidence>
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    </evidence>
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        </source>
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        <source>
            <dbReference type="ARBA" id="ARBA00023278"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00038081"/>
        </source>
    </evidence>
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        <source>
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    </evidence>
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    </evidence>
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    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
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    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS51364"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015493.1"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1643" mass="177975" checksum="AB9C3D01C64E81D3" modified="2019-05-08" version="1">TWNVPLFLPRQISKPSCLPLSIGARVGQAPSPAPQLERGKPETPLPRGPWVVVCPPRARPTIILINTVRATLACCTGSHVSLFTPFHVPPAVCDPPCQNKGSCSRPQLCICRSGFQGAQCEEVVPERVYHPPGSVAALQPPAGSLPKRQQLCAMFLNPLPVPPSCRRIVFPVSRHTGSSRTVRRYPAGNTQLMSNALPNGNGHEQSSSGPHAAYQEHTLSLWGANLTEKIRKIKIVFTPTICKQTCRSGHCYNSCQKGDTTTLYSQGGQNHDPKSGFRIYFCQIPCLNGGRCVGRDECWCPSNSTGKFCHLPVPKLEKNQVGRVPKTLASGSIKQSMYTLPLSNQLASLNPSLVNVHIKHPPEATVQIHQVARVRGDLGQSEENSVEAVLLSQLPTGPQHSYSNGNSNSITTESGQQQRPPELMGRCFRETLHGQCANPLEGLAKFEDCCGSVGLFWGVNRCIPCPPRPAHPVIENRQVECPQGYKRLNRSHCQDINECLMLGLCKDTECVNTRGSFLCTCKPGTMLDPSRSRCVSDKAVSMEQGLCYRSAAEGVCALPLAQSITKQICCCSRVGKGWGRNCEKCPMLGSETFKEICPAGHGYTYSSSDIRLSMRKAEAEELPFSSEEQGESSDQTPTQAAKRLQLQEALSGGVTGTLARAGTSAAEGVLGCDATLGNPCTICWGTGEQWPVTSPCLSLSEMDRCASVPDPCGPGTCVNLPDRYSCICGPGYQLHPGLPQCIDDDECMRDPCAGKGHCINSVGSYSCLCYPGYTLLASQGTQTCQDLNECEQPSMCQGGQCTNTPGSYHCECARGYIMGHTGQCEDIDECADPSTCPRGRCVNALGSYECLSCAAGYQPMNGRCVDDDECFTVGICAHGLCINLAGSFRCSCYHGYEVTADEKSCQDINECATHNACPMGLCINSEGSFSCAACGAGYTVSRDGSMCEDIDECSSATACPLGICTNTDGSFACRACDAGYMLSSNRFTCEDMNECLASEICSPNGECLNSHGSYFCICAPGFSNAAGGVSCQGEQLGQMAAAKCALDPGYRPGLSLAPTGPVYTHRTASVLIFILSLPDVDECADKSRCLRGQCFNTEGSYRCLCENGFRHSQETDDCVDIDECANETLCGSHAFCDNTDGSFRCLCDSGYENLPPGQDCVDVNECELMVAVCGTALCENVEGSFLCLCLSDHEEYNTEAGECRPRAGDPERPIAPRPAGSERKECYYHISDVQLCDSVLAKNTTKEECCCTEGAAWGDNCETHPCPILGTVEYSEICPSGKGYIPVEGTLLFGQTSYTDADECEMFGPEICRNGHCVNTVPGYKCFCRSGYLYDSSRLEYQDECENEMSCVNGECLNTAGSFHCFCGLPLVLDVTRNRCINVSSRAEDPEEHEIHLDICWQRVSNYICNQPLRGRRTTYTECCCRYGEAWSQDCALCPHRASEDFAHLCNVARGETEREPGLQERPGYEYGPALEDSHYGLYSPDLGPYYNYLSPEYGAPDASFPQRELSSEVGGSTFLRTPQLPPAQSQPRYLPRHFDSFEGLQAEECGILNGCENGRCVRVREGYTCDCFDGFQLDMTHMACVDINECEEVSSSGPLCEGGTCENTEGSYRCNCSPGFVVQAEPHRCAPAPAQRPEAAQL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKN6</accession>
    <name>A0A452HKN6_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="4">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the cytochrome P450 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKN6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017705.1">
        <property type="protein sequence ID" value="ENSGAGP00000015501.1"/>
        <property type="gene ID" value="ENSGAGG00000011439.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0020037">
        <property type="term" value="F:heme binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005506">
        <property type="term" value="F:iron ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004497">
        <property type="term" value="F:monooxygenase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016705">
        <property type="term" value="F:oxidoreductase activity, acting on paired donors, with incorporation or reduction of molecular oxygen"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.630.10">
        <property type="entry name" value="Cytochrome P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001128">
        <property type="entry name" value="Cyt_P450"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002401">
        <property type="entry name" value="Cyt_P450_E_grp-I"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036396">
        <property type="entry name" value="Cyt_P450_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050196">
        <property type="entry name" value="Cytochrome_P450_Monoox"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24291">
        <property type="entry name" value="CYTOCHROME P450 FAMILY 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24291:SF201">
        <property type="entry name" value="CYTOCHROME P450, FAMILY 4, SUBFAMILY B, POLYPEPTIDE 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00067">
        <property type="entry name" value="p450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00463">
        <property type="entry name" value="EP450I"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00385">
        <property type="entry name" value="P450"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48264">
        <property type="entry name" value="Cytochrome P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0408">Iron</keyword>
    <keyword evidence="3" id="KW-0472">Membrane</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="3" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="3">
        <location>
            <begin position="20"/>
            <end position="40"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00010617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015501.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="437" mass="51485" checksum="53A1FA889149F2F7" modified="2019-05-08" version="1">MTSVVVRLITAIWCWLKKDVFRIVYLLTVLCLTYMVLKAIKLYQWRRKLNKALQSFPGPPVHWLFGNVHEVRLKNQKQLSCIERLPDNIFYRYVLPWIGKGLLTLQGPKWFQHRKLLTPGFHYGVLKPYVSLITKSTQVMLDKWEHLITQEKSVELFGHVSLMTLDSILKCAFSHESNCQMDRSNSYIKSVYEITYQINYRIWYLPYQIDFIYWLSRDGRRFQKACEVVHLQTDKVIKERKKSLKDERELEKIQKKRHLDFLDILLCAKDEKGDGLSDEDIRAEVDTFMFGGHDTTASGICWTLYCLAMYPEHQQKCREEIRELLGDRETIQWDDLGKMPNTTMCIKESMRIFPPVPFVGRQLSKPITFHDGRYLPEGTLVGINIYLIHRDPSLWKDPEVFDPMRFSPENSSKRHSHAFMPFSAGARWGIVLVFMSS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKN7</accession>
    <name>A0A452HKN7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">CDK5 regulatory subunit-associated protein 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Golgi apparatus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000017695.1">
        <property type="protein sequence ID" value="ENSGAGP00000015491.1"/>
        <property type="gene ID" value="ENSGAGG00000011684.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035371">
        <property type="term" value="C:microtubule plus-end"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097431">
        <property type="term" value="C:mitotic spindle pole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000242">
        <property type="term" value="C:pericentriolar material"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043015">
        <property type="term" value="F:gamma-tubulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008017">
        <property type="term" value="F:microtubule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007099">
        <property type="term" value="P:centriole replication"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007059">
        <property type="term" value="P:chromosome segregation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000132">
        <property type="term" value="P:establishment of mitotic spindle orientation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001578">
        <property type="term" value="P:microtubule bundle formation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046600">
        <property type="term" value="P:negative regulation of centriole replication"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090266">
        <property type="term" value="P:regulation of mitotic cell cycle spindle assembly checkpoint"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042791">
        <property type="entry name" value="CDK5RAP2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056273">
        <property type="entry name" value="CDK5RAP2_MYOME_CC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012943">
        <property type="entry name" value="Cnn_1N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46930">
        <property type="entry name" value="CDK5 REGULATORY SUBUNIT-ASSOCIATED PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46930:SF1">
        <property type="entry name" value="CDK5 REGULATORY SUBUNIT-ASSOCIATED PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23246">
        <property type="entry name" value="CC_CDK5RAP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07989">
        <property type="entry name" value="Cnn_1N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="7" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Centrosomin N-terminal motif 1" evidence="11">
        <location>
            <begin position="57"/>
            <end position="130"/>
        </location>
    </feature>
    <feature type="domain" description="CDK5 regulatory subunit-associated protein 2/Myomegalin coiled coil" evidence="12">
        <location>
            <begin position="461"/>
            <end position="580"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="888"/>
            <end position="925"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="58"/>
            <end position="128"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="155"/>
            <end position="182"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="276"/>
            <end position="414"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="451"/>
            <end position="578"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="619"/>
            <end position="656"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="1414"/>
            <end position="1445"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="1512"/>
            <end position="1546"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="1616"/>
            <end position="1654"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="891"/>
            <end position="900"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004555"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00020479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF07989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF23246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015491.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1929" mass="222816" checksum="F68C9C14FA5499F0" modified="2019-05-08" version="1">MDSLPGEDLTLPVEFSGSVSISRLSDVSRDEPRIGLGISGALLDVAEDAISPTRARTMKDYENQITDLKKENFNLKLRIYFLEERMQQKFDGPTEDIYKINIELKVEVESLKRDLQEREKLLIKASKAVESLAQGGDAEIQRVKEEARKKVQQVEEILTSRINLLEENLKAAQEEVEKAFAVTEREKVLRMAVEQKLSGVTNTHLKELDSITELPEKDRLIEQLTLSLKSKEAIIQHLEEEKSQSESCDGNLPAEKIRELTVALRHEKDSEIEAIKMELRNERNCFEKKVQNEELNSEIENLNVSLAKAREATHKAQIQKFKGAEDYQALLMEKETLLAELRSENLTKDTENRKLQRRIKRTDQELNDLNLEREKMERELDEAQLQKSRGDKTINDLRNQLEKLHGEMTEREKAVEHHYNILLSESNQKLQSQELVIKRLTDGVSQKDLLLQKFDAAIKEKDVELQELIKKCKNLLKAKEDLELKNEDLIKEKCASESQQSIIKIVKELEEKKESEYEELILVLRKEQHIYSTLVKTLKDSDSINSLQTELNNIFLLRKQLEEDILANRNLRKVLEDQIKNTKNREDETFSFCGDQTSYMSICLGEHDHLNLQVDHLSLEELKKKVAELILMVKDLHSVNQELKKKQLRLSRSETQGKETRKIINQNEFLERTEETMTQAGDFDNKNAESCQCSERTSEACPQLPLDHPLDLDMQIPPSDYSDGVCKDRHYRNPNKPWFEHDIAGKHFGRIEMGEHEVEEMSLLTSLLNENGTSVLESRQEQMKIVIALLDHLSTTEQDCSSEDIENKDEKDLRQMIIQLRAELRHFKQVNKILKQQVELKSTSDGKEKLYPDTAPQVNKDIELLKVELKDTATQTMTIEGNVMRLKHEGKRGASEEKTKYSRLNAEPQQENAYKKGEQHESLSVTSRTNKSDFIYPFKKSRLPIPLKSSRSLGNIPLTSCTQKPDMHLQHRLMALDEGLKERKIENKQLHKELSLCGSVIETVQEKAEEERVLVRAESLVRMDCKEIPVDSQAAGFDTHDKNENKMVTDQRTSQETETPHNISDMSILHSLKELNKEFYSMENQDIDSATTNSGSKSLLSLKLSVPSTDAFDDYEVIDDLEELRQRIKDMKSELEKYKMFIFHLQPTNPFLSSDIFNIVLNDAELPIERHVIQMQDTDGQETQTFSTVHQQVDYEIRIENLNSQSAETPDEQTSQNLKQLLLENEAELEREQIANMHLLDEVYRLQNKLRETSPSKYDSLVHSQARELSFQRQQIKEIHSICVTYHQHLTSLIKAFEELLQASDVDYYVAEGFREQLNQSVQLFEKLEKKFLYGESLDAEVAKLYELAQRDNETTTTMHQQLKDESPIPSAFHSLSDFEMSEKSSIGSPEPKQDLVEEDKLALVPSKFPPALLMEHLQEIRSLRQRLEDSIKTNERLRLQLERQVTDMELNQGSANIFIHGSEQHNSLTSEIHFLRKQNKVLNVMLAKGSRDKQKENEKLRESLSKKNAIIEHLHQDYECFKKENEKLQKQIGKKEDENRHLTHEIYSSHNELNRLQTEMNVKQHQLSENDKLLQSLRVELKVYEKLDEARRRKTDILDPGHEEFQKDQNNPLDLHELLTEIQSLRVQLERSIETNKTLHEKLEEQLSKEKREEAASVSAVNINYLLKRDSQHYTRMNEVGHKFSDPSSNVCELQNCGHGTSPGKAGTPKDKNIINKQMFCIPETQLAAKDDLDSSSRCSSSSSTGTSCTPRLVPGHRMWADKNGRHILGLIEDYNALRKQISEGQKLVSEMEAPLREITGMKNLEPGVKVPDQESLNNFSANIHTVQQNLEEAARLLKLLWRVSLPMKVVHSTAHCSQDEGMKAEILRLHKKLSEQEKKLHSTVKRLHSTNQLKENMEKVIIDQLALTHDVLKKARGNLEVSKICLH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKN8</accession>
    <name>A0A452HKN8_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="11">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000017710.1">
        <property type="protein sequence ID" value="ENSGAGP00000015504.1"/>
        <property type="gene ID" value="ENSGAGG00000011700.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003779">
        <property type="term" value="F:actin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000083">
        <property type="entry name" value="Uveal autoantigen with coiled-coil domains and ankyrin repeats"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.1490">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1170.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042420">
        <property type="entry name" value="RAI14/UACA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24129">
        <property type="entry name" value="ANKYCORBIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24129:SF1">
        <property type="entry name" value="UVEAL AUTOANTIGEN WITH COILED-COIL DOMAINS AND ANKYRIN REPEATS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00023">
        <property type="entry name" value="Ank"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13637">
        <property type="entry name" value="Ank_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01415">
        <property type="entry name" value="ANKYRIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="5"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5 9" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="6 10" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <feature type="repeat" description="ANK" evidence="9">
        <location>
            <begin position="46"/>
            <end position="78"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="9">
        <location>
            <begin position="79"/>
            <end position="111"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="9">
        <location>
            <begin position="112"/>
            <end position="144"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="9">
        <location>
            <begin position="145"/>
            <end position="177"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="9">
        <location>
            <begin position="178"/>
            <end position="210"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="10">
        <location>
            <begin position="274"/>
            <end position="346"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="10">
        <location>
            <begin position="411"/>
            <end position="484"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="10">
        <location>
            <begin position="511"/>
            <end position="580"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="10">
        <location>
            <begin position="608"/>
            <end position="703"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="10">
        <location>
            <begin position="732"/>
            <end position="1070"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="10">
        <location>
            <begin position="1103"/>
            <end position="1133"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="10">
        <location>
            <begin position="1158"/>
            <end position="1192"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="10">
        <location>
            <begin position="1274"/>
            <end position="1350"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015504.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1369" mass="157675" checksum="AA54D86FBA7AF37A" modified="2019-05-08" version="1">MASLWLNTDWSKYDDRLMKAAERGDVEKLSSVLAKKGVNPTKLDVEGRSAFHVVASKGNLDCLNTILVHGVDIVASDAAGRNALHLAAKYGHALCLQKLLQYNCPTENLDLHGRTALHDAAMSDCSSSIQLLCDHGASVNAKDGDGRTPLVLATQMCRPTVCQLLIDRGADLNARDKQNRTALMLGCEYGCKDAVEVLLKNGADISLVDALGHDCSYYARIGDNIDILALIKTAIEDSSKAKELMKKGQPEYKVTNVSQKRNQLYAQEVNFKQYQREQLSMQELEAENEDLKNRLREIQQEHRILLNRVSVLELQLNEEQMVADDLESEKDELRRLLTAKEKQQEESLRSIETLKSKLRHYERDYLGSGSDSGKEDMLLKQGQVFATESQTKSMLRPLELSLSCQSSSSENEILKRELDNMRISYGTAKEEISKLQKELSHKVAECKALASECERTKEESDRQIKQLEDALKDVQKRMFDSEGKVKQMQTHFLVLKDHLTNEAASGSNKLTEDLKDQLKDMKAKYEGVSAEVGKLRNQIKQNELLVVEFKRDEGRLVEENKRLQKELSMLEMKREKSGRNVLEVQGQLKEMAAKLVLSVPVEKFENMKSLLSTEVNEKAKKLVEMEREYEKLQVELRLLKREFESQKAKFSQYVRPEEHEQMRGRFERKTEELEKTISELSKNQTLQKEVERVHMDNKLLKQQIHTLTTEIKSQYVPLKLNEEMKKTSDLTIGDLTKKLAEVTQRYNENKAEAEKLMAEKTSLSETVSHFQAVYLSPEKHEKEVAALKSTVVELKRQLAELHQKYDAEQARVSKLVSENTTLRDSLNDQYVPAITHEEIKTVLNSTLEKTNRELSDLKEKTEDIKREFMRVNEENGTLKKKLRLLQTHIQTEYICVKDHESKVSTLNKSMQELQDNNTETLAKYKRGQEEILQLHAEIEAQKKELDTIQECIKSKYAPVASFEEREQNFEAVVKELKKQLLEQAQRCREWEEETQKYKQENEKLKNRILSIENDLQQNYILAEKSHEMEKMFTIRMEELNKQLRELLQKYTEIKHEKDKLQEESAKQISEALAVQTRMQGHYVPVQQFEALKRTLGHTIEDLKEDLNSKKACYDRELEKVRELQQELADLHNSSISLAEHAQMKEAFEKEIVAVKTSLKKEVEENQAKSEEISKLQSEIQSTKQALTELESKEVVDMLEYKSMKSALETQVASIAGNLASLNRKYEEMCEEALQAKKEELSAKDEKELLQSRSFCIEQEIKDQKERCDKSLTTITDLQKRIQESAKQVEAKDNKITELLNDVERLKQALNGLSQLTYTCGVPSKRQNQQVEMLQHQVKTLQQQLADADRQHQEVISIYRTHLLSAVQVC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKN9</accession>
    <name>A0A452HKN9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">CDK5 regulatory subunit-associated protein 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Golgi apparatus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000017686.1">
        <property type="protein sequence ID" value="ENSGAGP00000015484.1"/>
        <property type="gene ID" value="ENSGAGG00000011684.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035371">
        <property type="term" value="C:microtubule plus-end"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097431">
        <property type="term" value="C:mitotic spindle pole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000242">
        <property type="term" value="C:pericentriolar material"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043015">
        <property type="term" value="F:gamma-tubulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008017">
        <property type="term" value="F:microtubule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007099">
        <property type="term" value="P:centriole replication"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007059">
        <property type="term" value="P:chromosome segregation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000132">
        <property type="term" value="P:establishment of mitotic spindle orientation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001578">
        <property type="term" value="P:microtubule bundle formation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046600">
        <property type="term" value="P:negative regulation of centriole replication"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090266">
        <property type="term" value="P:regulation of mitotic cell cycle spindle assembly checkpoint"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042791">
        <property type="entry name" value="CDK5RAP2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056273">
        <property type="entry name" value="CDK5RAP2_MYOME_CC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012943">
        <property type="entry name" value="Cnn_1N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46930">
        <property type="entry name" value="CDK5 REGULATORY SUBUNIT-ASSOCIATED PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46930:SF1">
        <property type="entry name" value="CDK5 REGULATORY SUBUNIT-ASSOCIATED PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23246">
        <property type="entry name" value="CC_CDK5RAP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07989">
        <property type="entry name" value="Cnn_1N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="7" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Centrosomin N-terminal motif 1" evidence="11">
        <location>
            <begin position="57"/>
            <end position="130"/>
        </location>
    </feature>
    <feature type="domain" description="CDK5 regulatory subunit-associated protein 2/Myomegalin coiled coil" evidence="12">
        <location>
            <begin position="501"/>
            <end position="620"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="928"/>
            <end position="965"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="58"/>
            <end position="128"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="155"/>
            <end position="182"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="276"/>
            <end position="454"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="491"/>
            <end position="618"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="659"/>
            <end position="696"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="1454"/>
            <end position="1547"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="1617"/>
            <end position="1655"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="931"/>
            <end position="940"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004555"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00020479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF07989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF23246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015484.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1930" mass="223151" checksum="DDE5EDCDA0EE60AA" modified="2019-05-08" version="1">MDSLPGEDLTLPVEFSGSVSISRLSDVSRDEPRIGLGISGALLDVAEDAISPTRARTMKDYENQITDLKKENFNLKLRIYFLEERMQQKFDGPTEDIYKINIELKVEVESLKRDLQEREKLLIKASKAVESLAQGGDAEIQRVKEEARKKVQQVEEILTSRINLLEENLKAAQEEVEKAFAVTEREKVLRMAVEQKLSGVTNTHLKELDSITELPEKDRLIEQLTLSLKSKEAIIQHLEEEKSQSESCDGNLPAEKIRELTVALRHEKDSEIEAIKMELRNERNCFEKKVQSLQEELREREDELSIEKKNGLKRDKTIQGLTLALKTKEKENEELNSEIENLNVSLAKAREATHKAQIQKFKGAEDYQALLMEKETLLAELRSENLTKDTENRKLQRRIKRTDQELNDLNLEREKMERELDEAQLQKSRGDKTINDLRNQLEKLHGEMTEREKAVEHHYNILLSESNQKLQSQELVIKRLTDGVSQKDLLLQKFDAAIKEKDVELQELIKKCKNLLKAKEDLELKNEDLIKEKCASESQQSIIKIVKELEEKKESEYEELILVLRKEQHIYSTLVKTLKDSDSINSLQTELNNIFLLRKQLEEDILANRNLRKVLEDQIKNTKNREDETFSFCGDQTSYMSICLGEHDHLNLQVDHLSLEELKKKVAELILMVKDLHSVNQELKKKQLRLSRSETQGKETRKIINQNEFLERTEETMTQAGDFDNKNAESCQCSERTSEACPQLPLDHPLDLDMQIPPSDYSDGVCKDRHYRNPNKPWFEHDIAGKHFGRIEMGEHEVEEMSLLTSLLNENGTSVLESRQEQMKIVIALLDHLSTTEQDCSSEDIENKDEKDLRQMIIQLRAELRHFKQVNKILKQQVELKSTSDGKEKLYPDTAPQVNKDIELLKVELKDTATQTMTIEGNVMRLKHEGKRGASEEKTKYSRLNAEPQQENAYKKGEQHESLSVTSRTNKSDFIYPFKKSRLPIPLKSSRSLGNIPLTSCTQKPDMHLQHRLMALDEGLKERKIENKQLHKELSLCGSVIETVQEKAEEERVLVRAESLVRMDCKEIPVDSQAAGFDTHDKNENKMVTDQRTSQETETPHNISDMSILHSLKELNKEFYSMENQDIDSATTNSGSKSLLSLKLSVPSTDAFDDYEVIDDLEELRQRIKDMKSELEKYKMFIFHLQPTNPFLSSDIFNIVLNDAELPIERHVIQMQDTDGQETQTFSTVHQQVDYEIRIENLNSQSAETPDEQTSQNLKQLLLENEAELEREQIANMHLLDEVYRLQNKLRETSPSKYDSLVHSQARELSFQRQQIKEIHSICVTYHQHLTSLIKAFEELLQASDVDYYVAEGFREQLNQSVQLFEKLEKKFLYGESLDAEVAKLYELAQRDNETTTTMHQQLKDESPIPSAFHSLSDFEMSEKSSIGSPEPKQDLVEEDKLALVPSKFPPALLMEHLQEIRSLRQRLEDSIKTNERLRLQLERQVTDMELNQDKQKENEKLRESLSKKNAIIEHLHQDYECFKKENEKLQKQIGKKEDENRHLTHEIYSSHNELNRLQTEMNVKQHQLSENDKLLQSLRVELKVYEKLDEARRRKTDILDPGHEEFQKDQNNPLDLHELLTEIQSLRVQLERSIETNKTLHEKLEEQLSKEKREEAASVSAVNINYLLKRDSQHYTRMNEVGHKFSDPSSNVCELQNCGHGTSPGKAGTPKDKNIINKQMFCIPETQLAAKDDLDSSSRCSSSSSTGTSCTPRLVPGHRMWADKNGRHILGLIEDYNALRKQISEGQKLVSEMEAPLREITGMKNLEPGVKVPDQESLNNFSANIHTVQQNLEEAARLLKLLWRVSLPMKVVHSTAHCSQDEGMKAEILRLHKKLSEQEKKLHSTVKRLHSTNQLKENMEKVIIDQLALTHDVLKKARGNLEVSKICLH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKQ0</accession>
    <name>A0A452HKQ0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Groucho/TLE N-terminal Q-rich domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="6">Transcriptional corepressor that binds to a number of transcription factors. Inhibits the transcriptional activation mediated by CTNNB1 and TCF family members in Wnt signaling. The effects of full-length TLE family members may be modulated by association with dominant-negative AES.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the WD repeat Groucho/TLE family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKQ0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017698.1">
        <property type="protein sequence ID" value="ENSGAGP00000015494.1"/>
        <property type="gene ID" value="ENSGAGG00000011690.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005667">
        <property type="term" value="C:transcription regulator complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003714">
        <property type="term" value="F:transcription corepressor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090090">
        <property type="term" value="P:negative regulation of canonical Wnt signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00200">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:000001">
        <property type="entry name" value="transducin-like enhancer protein 3 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005617">
        <property type="entry name" value="Groucho/TLE_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009146">
        <property type="entry name" value="Groucho_enhance"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019775">
        <property type="entry name" value="WD40_repeat_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036322">
        <property type="entry name" value="WD40_repeat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001680">
        <property type="entry name" value="WD40_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10814">
        <property type="entry name" value="TRANSDUCIN-LIKE ENHANCER PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10814:SF24">
        <property type="entry name" value="TRANSDUCIN-LIKE ENHANCER PROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03920">
        <property type="entry name" value="TLE_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00400">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01850">
        <property type="entry name" value="GROUCHOFAMLY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00320">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50978">
        <property type="entry name" value="WD40 repeat-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00678">
        <property type="entry name" value="WD_REPEATS_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50082">
        <property type="entry name" value="WD_REPEATS_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50294">
        <property type="entry name" value="WD_REPEATS_REGION"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0539">Nucleus</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="3 7" id="KW-0853">WD repeat</keyword>
    <feature type="domain" description="Groucho/TLE N-terminal Q-rich" evidence="10">
        <location>
            <begin position="18"/>
            <end position="125"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="7">
        <location>
            <begin position="502"/>
            <end position="543"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="7">
        <location>
            <begin position="544"/>
            <end position="585"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="130"/>
            <end position="283"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="8">
        <location>
            <begin position="26"/>
            <end position="60"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="132"/>
            <end position="144"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="145"/>
            <end position="183"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="208"/>
            <end position="219"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="220"/>
            <end position="235"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="236"/>
            <end position="245"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="246"/>
            <end position="258"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005969"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022574"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00045617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="Pfam" id="PF03920"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015494.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="702" mass="76121" checksum="0D117C5263D3A22F" modified="2019-05-08" version="1">MYPQGRHPAPHQPGQPGFKFTVAESCDRIKDEFQFLQAQYHSLKVEYDKLANEKTEMQRHYVMYYEMSYGLNIEMHKQTEIAKRLNTILAQIMPFLSQEHQQQVAQAVERAKQVTMTELNAIIGVRGLPNLPLTNNSVSPSDSLRASEKHRSSTDYSIDAKKRKAEEKDSMSRYDSDGDKSDDLVVDVSNEDPATPRVSPAHSPPENGIDKTRGLKKDAPNSPASVASSSSTPSSKTKDLGHNDKSSTPGLKSNTPTPRNEAPTPGTSTTPGLRSVPGKAAGMDPLASALRTPISITGPYAAPFAMMGHHEMNGSLTSPGAYAGLHNIPPQMSTAAAAAYGRSPMVSFGAVGFDPHPPMRGPGLPTSLASIPGGKPAYSFHVSADGQMQPVPFPHDALAGPGIPRHARQINTLSHGEVVCAVTISNPTRHVYTGGKGCVKIWDISQPGSKSPISQLDCLNRDNYIRSCKLLPDGRTLIVGGEASTLTIWDLASPTPRIKAELTSSAPACYALAISPDAKVCFSCCSDGNIAVWDLHNQTLVRQFQGHTDGASCIDISHDGTKLWTGGLDNTVRSWDLREGRQLQQHDFTSQIFSLGYCPTGEWLAVGMESSNVEVLHHTKPDKYQLHLHESCVLSLKFAYCGKWFVSTGKDNLLNAWRTPYGASIFQSKESSSVLSCDISADDKYIVTGSGDKKATVYEVIY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKQ1</accession>
    <name>A0A452HKQ1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Semaphorin-6A</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="18">Semaphorin VIA</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="17">Semaphorin-6A-1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Cell surface receptor for PLXNA2 that plays an important role in cell-cell signaling. Required for normal granule cell migration in the developing cerebellum. Promotes reorganization of the actin cytoskeleton and plays an important role in axon guidance in the developing central nervous system. Can act as repulsive axon guidance cue. Has repulsive action towards migrating granular neurons. May play a role in channeling sympathetic axons into the sympathetic chains and controlling the temporal sequence of sympathetic target innervation.</text>
    </comment>
    <comment type="subunit">
        <text evidence="15">Active as a homodimer or oligomer. The SEMA6A homodimer interacts with a PLXNA2 homodimer, giving rise to a heterotetramer. Interacts with EVL.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the semaphorin family.</text>
    </comment>
    <comment type="caution">
        <text evidence="19">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKQ1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017727.1">
        <property type="protein sequence ID" value="ENSGAGP00000015517.1"/>
        <property type="gene ID" value="ENSGAGG00000011703.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045499">
        <property type="term" value="F:chemorepellent activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030215">
        <property type="term" value="F:semaphorin receptor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007411">
        <property type="term" value="P:axon guidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001755">
        <property type="term" value="P:neural crest cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:2001224">
        <property type="term" value="P:positive regulation of neuron migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071526">
        <property type="term" value="P:semaphorin-plexin signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd11266">
        <property type="entry name" value="Sema_6A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:000028">
        <property type="entry name" value="semaphorin-6A isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1680.10:FF:000007">
        <property type="entry name" value="semaphorin-6A isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1680.10">
        <property type="entry name" value="ligand-binding face of the semaphorins, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002165">
        <property type="entry name" value="Plexin_repeat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001627">
        <property type="entry name" value="Semap_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036352">
        <property type="entry name" value="Semap_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027231">
        <property type="entry name" value="Semaphorin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11036">
        <property type="entry name" value="SEMAPHORIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11036:SF12">
        <property type="entry name" value="SEMAPHORIN-6A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01437">
        <property type="entry name" value="PSI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01403">
        <property type="entry name" value="Sema"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00630">
        <property type="entry name" value="Sema"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103575">
        <property type="entry name" value="Plexin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF101912">
        <property type="entry name" value="Sema domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51004">
        <property type="entry name" value="SEMA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="3" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="8" id="KW-0221">Differentiation</keyword>
    <keyword evidence="12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11 21" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7 22" id="KW-0732">Signal</keyword>
    <keyword evidence="6 21" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="10 21" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="22">
        <location>
            <begin position="1"/>
            <end position="18"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019146260" description="Semaphorin-6A" evidence="22">
        <location>
            <begin position="19"/>
            <end position="972"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="590"/>
            <end position="618"/>
        </location>
    </feature>
    <feature type="domain" description="Sema" evidence="23">
        <location>
            <begin position="24"/>
            <end position="512"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="701"/>
            <end position="725"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="804"/>
            <end position="844"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="862"/>
            <end position="909"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="935"/>
            <end position="972"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="20">
        <location>
            <begin position="864"/>
            <end position="875"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="20">
        <location>
            <begin position="876"/>
            <end position="896"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="20">
        <location>
            <begin position="959"/>
            <end position="972"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009492"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00053495"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00063062"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00074111"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00079635"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00080740"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00352"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS51004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015517.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="972" mass="108490" checksum="8A44CC2CB1C5663C" modified="2019-05-08" version="1">MRSEALLLYFTLLHFAGAGFPEDSEPISISHGNYTKQYPVFVGHKPGRNATQRHRLDIQLIMIMNRTLYIAARDHIYTVDMDTSHTEEIYFIKKLTWKSRQADVDTCRMKGKHKDECHNFIKVLLKRNDDTLFICGTNAFNPSCRNYKMETLEFLGDEFSGMARCPYDAKHANVALFAEGKLYSATVTDFLAIDAVIYRSLGDSPTLRTVKHDSKWLKEPYFVQAVDYGDYIYFFFREIAVEYNSMGKVVFPRVAQVCKNDMGGSQRVLEKQWTSFLKARLNCSVPGDSHFYFNILQAVTDVIHINGHDIVLATFSTPYNSIPGSAVCAYDMLDIANVFTGRFKEQKSPDSTWTPVPDERVPKPRPGCCAGSVSLEKYITSNEFPDDTLNFIKTHPLMDEAVPSIVSRPWFLRTMVRYRLTKIAVDTAAGPHQNHTVVFLGSEKGIILKFLARTGNSGFLNDSLFLEEMNVYNPEKCSYDGVEDKRIMGMQLDKHSRSLYVAFSTCVIKIPLGRCERHGKCKKACIASRDPYCGWVKEGGVCIQLLSGVKLAFEQDIEHGNTDGLGDCQNSFVALNGVIRESYLKGHDQLVPVTLLAIAVILAFVMGAVFSGIVVYCICDHRRRDVAVVQRKEKELAHSRRGSMSSVTKLSGLFGDTQSKEPKPEAILTPLMHNGKLATPGSTAKMLIKADQHHLDLAALPTPESTPTLQQKRKPSRGSRDWERNQNLINACTKDIPAMSSPVIPTDLPLRASPSHIPSVVVLPICQQGYQHEYVDQPKMSDMAQMAMEDQNATLEYKTIKEHLSSKSPNHGVNLVENLDSMPPKVPQREASLGPPGSSLAQSGLGKRLEMHSSAYNVDYKRNYPTNSLTRNHQTSTLKRNNTNSSNSSHLSRNQSFGRGDNPPPAPQRVDSIQVHAAQAQAVTVSRQPSLTAYNSLTRSGLKRTPSLKPDVPPKPSFTPLSTSMKPNDACT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKQ4</accession>
    <name>A0A452HKQ4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Semaphorin-6A</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="18">Semaphorin VIA</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="17">Semaphorin-6A-1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Cell surface receptor for PLXNA2 that plays an important role in cell-cell signaling. Required for normal granule cell migration in the developing cerebellum. Promotes reorganization of the actin cytoskeleton and plays an important role in axon guidance in the developing central nervous system. Can act as repulsive axon guidance cue. Has repulsive action towards migrating granular neurons. May play a role in channeling sympathetic axons into the sympathetic chains and controlling the temporal sequence of sympathetic target innervation.</text>
    </comment>
    <comment type="subunit">
        <text evidence="15">Active as a homodimer or oligomer. The SEMA6A homodimer interacts with a PLXNA2 homodimer, giving rise to a heterotetramer. Interacts with EVL.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the semaphorin family.</text>
    </comment>
    <comment type="caution">
        <text evidence="19">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKQ4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017715.1">
        <property type="protein sequence ID" value="ENSGAGP00000015508.1"/>
        <property type="gene ID" value="ENSGAGG00000011703.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045499">
        <property type="term" value="F:chemorepellent activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030215">
        <property type="term" value="F:semaphorin receptor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007411">
        <property type="term" value="P:axon guidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001755">
        <property type="term" value="P:neural crest cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:2001224">
        <property type="term" value="P:positive regulation of neuron migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071526">
        <property type="term" value="P:semaphorin-plexin signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd11266">
        <property type="entry name" value="Sema_6A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:000028">
        <property type="entry name" value="semaphorin-6A isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1680.10:FF:000007">
        <property type="entry name" value="semaphorin-6A isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1680.10">
        <property type="entry name" value="ligand-binding face of the semaphorins, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002165">
        <property type="entry name" value="Plexin_repeat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001627">
        <property type="entry name" value="Semap_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036352">
        <property type="entry name" value="Semap_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027231">
        <property type="entry name" value="Semaphorin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11036">
        <property type="entry name" value="SEMAPHORIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11036:SF12">
        <property type="entry name" value="SEMAPHORIN-6A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01437">
        <property type="entry name" value="PSI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01403">
        <property type="entry name" value="Sema"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00630">
        <property type="entry name" value="Sema"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103575">
        <property type="entry name" value="Plexin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF101912">
        <property type="entry name" value="Sema domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51004">
        <property type="entry name" value="SEMA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="3" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="8" id="KW-0221">Differentiation</keyword>
    <keyword evidence="12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11 21" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7 22" id="KW-0732">Signal</keyword>
    <keyword evidence="6 21" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="10 21" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="22">
        <location>
            <begin position="1"/>
            <end position="18"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019154623" description="Semaphorin-6A" evidence="22">
        <location>
            <begin position="19"/>
            <end position="1027"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="645"/>
            <end position="673"/>
        </location>
    </feature>
    <feature type="domain" description="Sema" evidence="23">
        <location>
            <begin position="24"/>
            <end position="512"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="583"/>
            <end position="604"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="755"/>
            <end position="778"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="859"/>
            <end position="899"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="917"/>
            <end position="964"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="990"/>
            <end position="1027"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="20">
        <location>
            <begin position="583"/>
            <end position="592"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="20">
        <location>
            <begin position="919"/>
            <end position="930"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="20">
        <location>
            <begin position="931"/>
            <end position="951"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="20">
        <location>
            <begin position="1014"/>
            <end position="1027"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009492"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00053495"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00063062"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00074111"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00079635"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00080740"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00352"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS51004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015508.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1027" mass="114405" checksum="6D84F60D62F5299D" modified="2019-05-08" version="1">MRSEALLLYFTLLHFAGAGFPEDSEPISISHGNYTKQYPVFVGHKPGRNATQRHRLDIQLIMIMNRTLYIAARDHIYTVDMDTSHTEEIYFIKKLTWKSRQADVDTCRMKGKHKDECHNFIKVLLKRNDDTLFICGTNAFNPSCRNYKMETLEFLGDEFSGMARCPYDAKHANVALFAEGKLYSATVTDFLAIDAVIYRSLGDSPTLRTVKHDSKWLKEPYFVQAVDYGDYIYFFFREIAVEYNSMGKVVFPRVAQVCKNDMGGSQRVLEKQWTSFLKARLNCSVPGDSHFYFNILQAVTDVIHINGHDIVLATFSTPYNSIPGSAVCAYDMLDIANVFTGRFKEQKSPDSTWTPVPDERVPKPRPGCCAGSVSLEKYITSNEFPDDTLNFIKTHPLMDEAVPSIVSRPWFLRTMVRYRLTKIAVDTAAGPHQNHTVVFLGSEKGIILKFLARTGNSGFLNDSLFLEEMNVYNPEKCSYDGVEDKRIMGMQLDKHSRSLYVAFSTCVIKIPLGRCERHGKCKKACIASRDPYCGWVKEGGVCIQLLSGVKLAFEQDIEHGNTDGLGDCQNSFVALNGHSSSLVPSTTTSDSPAQEGYDTRGRMLDWKDPVGSSENTDPYAAVSSHNHQDKKGVIRESYLKGHDQLVPVTLLAIAVILAFVMGAVFSGIVVYCICDHRRRDVAVVQRKEKELAHSRRGSMSSVTKLSGLFGDTQSKEPKPEAILTPLMHNGKLATPGSTAKMLIKADQHHLDLAALPTPESTPTLQQKRKPSRGSRDWERNQNLINACTKDIPAMSSPVIPTDLPLRASPSHIPSVVVLPICQQGYQHEYVDQPKMSDMAQMAMEDQNATLEYKTIKEHLSSKSPNHGVNLVENLDSMPPKVPQREASLGPPGSSLAQSGLGKRLEMHSSAYNVDYKRNYPTNSLTRNHQTSTLKRNNTNSSNSSHLSRNQSFGRGDNPPPAPQRVDSIQVHAAQAQAVTVSRQPSLTAYNSLTRSGLKRTPSLKPDVPPKPSFTPLSTSMKPNDACT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="22" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKQ5</accession>
    <name>A0A452HKQ5_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="1">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="1 2">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="2" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="1" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="1" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HKQ5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017722.1">
        <property type="protein sequence ID" value="ENSGAGP00000015515.1"/>
        <property type="gene ID" value="ENSGAGG00000011439.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0020037">
        <property type="term" value="F:heme binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005506">
        <property type="term" value="F:iron ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004497">
        <property type="term" value="F:monooxygenase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016705">
        <property type="term" value="F:oxidoreductase activity, acting on paired donors, with incorporation or reduction of molecular oxygen"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.630.10">
        <property type="entry name" value="Cytochrome P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036396">
        <property type="entry name" value="Cyt_P450_sf"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48264">
        <property type="entry name" value="Cytochrome P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000313" key="1">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015515.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="163" mass="18047" checksum="1F07099AE1FFC218" modified="2019-05-08" version="1">ICAYLVSASASDLGAYATGCQQPSCFVKSLHLAQTLNKALQSFPGPPVHWLFGNVHEFSRRGKDLDKLLEWSKKYARAFPVWFGKFVVFVSLNDADYAKAVCARGGKRVEVKGLEGISQEEFSSVPSWVQGGFCMWVVAAFTKPRSEKRCNLGSLIQAWSGQY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKQ7</accession>
    <name>A0A452HKQ7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Latent-transforming growth factor beta-binding protein 1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="19">Latent-transforming growth factor beta-binding protein 2</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="20">Transforming growth factor beta-1-binding protein 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="25 26">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="26" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="25" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="25" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">Key regulator of transforming growth factor beta (TGFB1, TGFB2 and TGFB3) that controls TGF-beta activation by maintaining it in a latent state during storage in extracellular space. Associates specifically via disulfide bonds with the Latency-associated peptide (LAP), which is the regulatory chain of TGF-beta, and regulates integrin-dependent activation of TGF-beta. Outcompeted by LRRC32/GARP for binding to LAP regulatory chain of TGF-beta.</text>
    </comment>
    <comment type="function">
        <text evidence="14">May play an integral structural role in elastic-fiber architectural organization and/or assembly.</text>
    </comment>
    <comment type="subunit">
        <text evidence="16">Forms part of the large latent transforming growth factor beta precursor complex; removal is essential for activation of complex. Interacts with SDC4. Interacts (via C-terminal domain) with FBN1 (via N-terminal domain) in a Ca(+2)-dependent manner.</text>
    </comment>
    <comment type="subunit">
        <text evidence="17">Interacts with TGFB1; associates via disulfide bonds with the Latency-associated peptide chain (LAP) regulatory chain of TGFB1, leading to regulate activation of TGF-beta-1. LTBP1 does not bind directly to TGF-beta-1, the active chain of TGFB1. Interacts (via C-terminal domain) with FBN1 (via N-terminal domain). Interacts with FBN2. Interacts with ADAMTSL2. Interacts with EFEMP2.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="13">Belongs to the LTBP family.</text>
    </comment>
    <comment type="caution">
        <text evidence="21">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000017731.1">
        <property type="protein sequence ID" value="ENSGAGP00000015520.1"/>
        <property type="gene ID" value="ENSGAGG00000008780.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019838">
        <property type="term" value="F:growth factor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00054">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="12"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000005">
        <property type="entry name" value="Fibrillin 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000194">
        <property type="entry name" value="Latent transforming growth factor beta binding protein 2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000469">
        <property type="entry name" value="Latent transforming growth factor beta binding protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.290.10:FF:000004">
        <property type="entry name" value="latent-transforming growth factor beta-binding protein 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000019">
        <property type="entry name" value="latent-transforming growth factor beta-binding protein 1 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000046">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 1 isoform x2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000014">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000077">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 3 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.290.10:FF:000001">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 3 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.290.10:FF:000002">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 3 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000056">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 3 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000115">
        <property type="entry name" value="latent-transforming growth factor beta-binding protein 4 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000024">
        <property type="entry name" value="Putative latent-transforming growth factor beta-binding protein 2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000273">
        <property type="entry name" value="Putative latent-transforming growth factor beta-binding protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="18"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.290.10">
        <property type="entry name" value="TGF-beta binding (TB) domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026823">
        <property type="entry name" value="cEGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001881">
        <property type="entry name" value="EGF-like_Ca-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013032">
        <property type="entry name" value="EGF-like_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000152">
        <property type="entry name" value="EGF-type_Asp/Asn_hydroxyl_site"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018097">
        <property type="entry name" value="EGF_Ca-bd_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009030">
        <property type="entry name" value="Growth_fac_rcpt_cys_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049883">
        <property type="entry name" value="NOTCH1_EGF-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017878">
        <property type="entry name" value="TB_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036773">
        <property type="entry name" value="TB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052080">
        <property type="entry name" value="vWF_C/EGF_Fibrillin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47333:SF4">
        <property type="entry name" value="EGF-LIKE DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47333">
        <property type="entry name" value="VON WILLEBRAND FACTOR C AND EGF DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12662">
        <property type="entry name" value="cEGF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07645">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12661">
        <property type="entry name" value="hEGF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00683">
        <property type="entry name" value="TB"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="19"/>
    </dbReference>
    <dbReference type="SMART" id="SM00179">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="17"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57184">
        <property type="entry name" value="Growth factor receptor domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57581">
        <property type="entry name" value="TB module/8-cys domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00010">
        <property type="entry name" value="ASX_HYDROXYL"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01186">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01187">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51364">
        <property type="entry name" value="TB"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9 21" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="4 21" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="3" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11" id="KW-0340">Growth factor binding</keyword>
    <keyword evidence="6" id="KW-0358">Heparin-binding</keyword>
    <keyword evidence="12" id="KW-0379">Hydroxylation</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="26" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="54"/>
            <end position="86"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="249"/>
            <end position="281"/>
        </location>
    </feature>
    <feature type="domain" description="TB" evidence="24">
        <location>
            <begin position="396"/>
            <end position="439"/>
        </location>
    </feature>
    <feature type="domain" description="TB" evidence="24">
        <location>
            <begin position="516"/>
            <end position="568"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="693"/>
            <end position="734"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="735"/>
            <end position="771"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="778"/>
            <end position="817"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="858"/>
            <end position="898"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="983"/>
            <end position="1025"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="1056"/>
            <end position="1096"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="1141"/>
            <end position="1182"/>
        </location>
    </feature>
    <feature type="domain" description="TB" evidence="24">
        <location>
            <begin position="1247"/>
            <end position="1301"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="1323"/>
            <end position="1361"/>
        </location>
    </feature>
    <feature type="domain" description="TB" evidence="24">
        <location>
            <begin position="1425"/>
            <end position="1477"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="1599"/>
            <end position="1643"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="366"/>
            <end position="391"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="589"/>
            <end position="612"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="22">
        <location>
            <begin position="366"/>
            <end position="390"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="21">
        <location>
            <begin position="58"/>
            <end position="68"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="21">
        <location>
            <begin position="76"/>
            <end position="85"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="21">
        <location>
            <begin position="253"/>
            <end position="263"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="21">
        <location>
            <begin position="271"/>
            <end position="280"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022674"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023183"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023278"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00038081"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00058734"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00059743"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00062144"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00062844"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00072992"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00072997"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00075443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS51364"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015520.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1655" mass="178252" checksum="104A95F46568A446" modified="2019-05-08" version="1">FLPRQISKPSCLPLSIGARVGQAPSPAPQLERGKATLACCTGSHVSLFTPFHVPPAVCDPPCQNKGSCSRPQLCICRSGFQGAQCEEVVPERVYHPPGSVAALQPPAGSLPKRAGSAEREGLVCAPRGSPPHLLAAGRIVFPVSRHTGSSRTVRRYPAGNTQLMSNALPNGNGHEQSSSGPHAAYQEHTLSLWGANLTEKIRKIKIVFTPTICKQTCRSGHCYNSCQKGDTTTLYSQGGQNHDPKSGFRIYFCQIPCLNGGRCVGRDECWCPSNSTGKFCHLPVPKLEKNQVGRVPKTLASGSIKQSMYTLPLSNQLASLNPSLVNVHIKHPPEATVQIHQVARVRGDLGQSEENSVEAVLLSQLPTGPQHSYSNGNSNSITTESGQQQRPPELMGRCFRETLHGQCANPLEGLAKFEDCCGSVGLFWGVNRCIPCPPRPAHPVIENRQVECPQGYKRLNRSHCQDINECLMLGLCKDTECVNTRGSFLCTCKPGTMLDPSRSRCVSDKAVSMEQGLCYRSAAEGVCALPLAQSITKQICCCSRVGKGWGRNCEKCPMLGSETFKEICPAGHGYTYSSSDIRLSMRKAEAEELPFSSEEQGESSDQTPTQAAKRLQLQEALSGGVTGTLARAGTSAAEGVLGCDATLVAALHTGGTADATQEPAPHPGGNPCTICWGTGEQWPVTSPCLSLSEMDRCASVPDPCGPGTCVNLPDRYSCICGPGYQLHPGLPQCIDDDECMRDPCAGKGHCINSVGSYSCLCYPGYTLLASQGTQTCQDLNECEQPSMCQGGQCTNTPGSYHCECARGYIMGHTGQCEDIDECADPSTCPRGRCVNALGSYECLSCAAGYQPMNGRCVDDDECFTVGICAHGLCINLAGSFRCSCYHGYEVTADEKSCQDINECATHNACPMGLCINSEGSFSCAACGAGYTVSRDGSMCEDIDECSSATACPLGICTNTDGSFACRACDAGYMLSSNRFTCEDMNECLASEICSPNGECLNSHGSYFCICAPGFSNAAGGVTKCALDPGYRPGLSLGPVYTHRTASVLIFILSLPDVDECADKSRCLRGQCFNTEGSYRCLCENGFRHSQETDDCVDVDECVDFGDTVCGTWRCQNTPGSYRCVMGCQPGFHRTPLGDCIDIDECANETLCGSHAFCDNTDGSFRCLCDSGYENLPPGQDCVDVNECELMVAVCGTALCENVEGSFLCLCLSDHEEYNTEAGECRPRAGMGDPERPIAPRPAGSERKECYYHISDVQLCDSVLAKNTTKEECCCTEGAAWGDNCETHPCPILGTVEYSEICPSGKGYIPVEGTLLFGQTSYTDADECEMFGPEICRNGHCVNTVPGYKCFCRSGYLYDSSRLECVDQDECENEMSCVNGECLNTAGSFHCFCGLPLVLDVTRNRCINVSSRAGEWDPEEHEIHLDICWQRVSNYICNQPLRGRRTTYTECCCRYGEAWSQDCALCPHRASEDFAHLCNVARGETEREPGLQERPGYEYGPALEDSHYGLYSPDLGPYYNYLSPEYGAPDPSPAPTVCSLQADGPSVSTGHFDSFEGLQAEECGILNGCENGRCVRVREGYTCDCFDGFQLDMTHMACVDINECEEVSSSGPLCEGGTCENTEGSYRCNCSPGFVVQAEPHRCAPAPAQRPEAAQL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKR0</accession>
    <name>A0A452HKR0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">SRCR domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="9">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKR0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017736.1">
        <property type="protein sequence ID" value="ENSGAGP00000015524.1"/>
        <property type="gene ID" value="ENSGAGG00000011707.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005507">
        <property type="term" value="F:copper ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004720">
        <property type="term" value="F:protein-lysine 6-oxidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030199">
        <property type="term" value="P:collagen fibril organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000001">
        <property type="entry name" value="Lysyl oxidase 4 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000016">
        <property type="entry name" value="Scavenger receptor cysteine-rich protein type 12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.250.10">
        <property type="entry name" value="SRCR-like domain"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050912">
        <property type="entry name" value="LOX-like_protein"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001695">
        <property type="entry name" value="Lysyl_oxidase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001190">
        <property type="entry name" value="SRCR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036772">
        <property type="entry name" value="SRCR-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45817:SF5">
        <property type="entry name" value="LYSYL OXIDASE HOMOLOG 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45817">
        <property type="entry name" value="LYSYL OXIDASE-LIKE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01186">
        <property type="entry name" value="Lysyl_oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00530">
        <property type="entry name" value="SRCR"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00258">
        <property type="entry name" value="SPERACTRCPTR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00202">
        <property type="entry name" value="SR"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56487">
        <property type="entry name" value="SRCR-like"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00420">
        <property type="entry name" value="SRCR_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50287">
        <property type="entry name" value="SRCR_2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7 9" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="8" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6" id="KW-0472">Membrane</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="3 10" id="KW-0732">Signal</keyword>
    <keyword evidence="2" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="10">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5018974066" description="SRCR domain-containing protein" evidence="10">
        <location>
            <begin position="23"/>
            <end position="447"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="11">
        <location>
            <begin position="29"/>
            <end position="130"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="11">
        <location>
            <begin position="151"/>
            <end position="263"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="11">
        <location>
            <begin position="282"/>
            <end position="330"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="9">
        <location>
            <begin position="55"/>
            <end position="119"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="9">
        <location>
            <begin position="68"/>
            <end position="129"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="9">
        <location>
            <begin position="99"/>
            <end position="109"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="9">
        <location>
            <begin position="228"/>
            <end position="238"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004167"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00196"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50287"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015524.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="447" mass="48907" checksum="CAAB53EEB296CE68" modified="2019-05-08" version="1">MLAGPIPTLLPLLLCLATPSWQQPLQIQVRLAGSRNQAGEGRLEVLYDGQWGTVCDDDFDIHVASVACRELGFESAVTWAHSAKYGQGEGPIWLDNLRCAGTERSLAECDSNGWGVSDCNHGEDAGIVCSAQHPPGPRTLGPAPGLQLEEVRLKPVLAWAKLSVPVSEGVVEVKHEGRWRQVCAAGWTRNNSRVVCGMLGFPREKWLNPRFYRYLSQKNTFWIHRVSCLGTEPHLSHCPVQVSPAARRRPACAHGMHAVISCVPGPEFQKAKAKPFRKGLRVRLAGGRSPEEGVVEVLVPVGGALKWGAVCGERWGLNEAMVVCRQLGLGFARLQRRFACANFGEQGVSAGCWDTYRHDIDCQWVDISDVGPGSYVLQVVVNPKHEVAESDFSNNVMRCQCKYDGHRIWLHGFSHADAGQASNQSLPNSLPLGDVFDVCTPELGRAP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKR3</accession>
    <name>A0A452HKR3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Mortality factor 4-like protein 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKR3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017737.1">
        <property type="protein sequence ID" value="ENSGAGP00000015525.1"/>
        <property type="gene ID" value="ENSGAGG00000011716.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035267">
        <property type="term" value="C:NuA4 histone acetyltransferase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006325">
        <property type="term" value="P:chromatin organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006310">
        <property type="term" value="P:DNA recombination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006281">
        <property type="term" value="P:DNA repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006355">
        <property type="term" value="P:regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd18983">
        <property type="entry name" value="CBD_MSL3_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.274.30:FF:000001">
        <property type="entry name" value="Mortality factor 4-like protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.140:FF:000024">
        <property type="entry name" value="Mortality factor 4-like protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.274.30">
        <property type="entry name" value="MRG domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016197">
        <property type="entry name" value="Chromo-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000953">
        <property type="entry name" value="Chromo/chromo_shadow_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008676">
        <property type="entry name" value="MRG"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038217">
        <property type="entry name" value="MRG_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026541">
        <property type="entry name" value="MRG_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025995">
        <property type="entry name" value="Tudor-knot"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10880:SF48">
        <property type="entry name" value="MORTALITY FACTOR 4 LIKE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10880">
        <property type="entry name" value="MORTALITY FACTOR 4-LIKE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05712">
        <property type="entry name" value="MRG"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11717">
        <property type="entry name" value="Tudor-knot"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF038133">
        <property type="entry name" value="HAT_Nua4_EAF3/MRG15"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00298">
        <property type="entry name" value="CHROMO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54160">
        <property type="entry name" value="Chromo domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51640">
        <property type="entry name" value="MRG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0007">Acetylation</keyword>
    <keyword evidence="4" id="KW-0156">Chromatin regulator</keyword>
    <keyword evidence="3" id="KW-0227">DNA damage</keyword>
    <keyword evidence="8" id="KW-0233">DNA recombination</keyword>
    <keyword evidence="9" id="KW-0234">DNA repair</keyword>
    <keyword evidence="2" id="KW-0341">Growth regulation</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0804">Transcription</keyword>
    <keyword evidence="6" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="Chromo" evidence="13">
        <location>
            <begin position="10"/>
            <end position="78"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="77"/>
            <end position="99"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="129"/>
            <end position="166"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022604"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023172"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00071326"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="SMART" id="SM00298"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015525.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="366" mass="42092" checksum="837F883842A0CF66" modified="2019-05-08" version="1">MAPKQDPKPKFQEGERVLCFHGPLLYEAKCVKVAIKDKQVKYFIHYSGWNKNWDEWVPESRVLKYVDTNLQKQKELQKANQEQYAEGKMRGAAPGKKTSGLQQKNVEVFFRRDGAHTVCCLETPTISTRKTKKNKQKTPGIGEGSSTSETPQPPRKKRARVDPTVESEETFMNRVEVKVKIPEELKPWLVDDWDLITRQKQIIIYFFPGIFTQAVLPIPHLKPLFYLPAKKNVDSILEDYASYKKSRGNTDNKEYAVNEVVAGIKEYFNVMLGTQLLYKFERPQYAEILADHPDAPMSQVYGAPHLLRLFVRIGAMLAYTPLDEKSLALLLNYLHDFLKYLAKNASTLFSASDYEVAPPEYHRKAV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKR5</accession>
    <name>A0A452HKR5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Latent-transforming growth factor beta-binding protein 1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="19">Latent-transforming growth factor beta-binding protein 2</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="20">Transforming growth factor beta-1-binding protein 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="25 26">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="26" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="25" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="25" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">Key regulator of transforming growth factor beta (TGFB1, TGFB2 and TGFB3) that controls TGF-beta activation by maintaining it in a latent state during storage in extracellular space. Associates specifically via disulfide bonds with the Latency-associated peptide (LAP), which is the regulatory chain of TGF-beta, and regulates integrin-dependent activation of TGF-beta. Outcompeted by LRRC32/GARP for binding to LAP regulatory chain of TGF-beta.</text>
    </comment>
    <comment type="function">
        <text evidence="14">May play an integral structural role in elastic-fiber architectural organization and/or assembly.</text>
    </comment>
    <comment type="subunit">
        <text evidence="16">Forms part of the large latent transforming growth factor beta precursor complex; removal is essential for activation of complex. Interacts with SDC4. Interacts (via C-terminal domain) with FBN1 (via N-terminal domain) in a Ca(+2)-dependent manner.</text>
    </comment>
    <comment type="subunit">
        <text evidence="17">Interacts with TGFB1; associates via disulfide bonds with the Latency-associated peptide chain (LAP) regulatory chain of TGFB1, leading to regulate activation of TGF-beta-1. LTBP1 does not bind directly to TGF-beta-1, the active chain of TGFB1. Interacts (via C-terminal domain) with FBN1 (via N-terminal domain). Interacts with FBN2. Interacts with ADAMTSL2. Interacts with EFEMP2.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="13">Belongs to the LTBP family.</text>
    </comment>
    <comment type="caution">
        <text evidence="21">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000017741.1">
        <property type="protein sequence ID" value="ENSGAGP00000015529.1"/>
        <property type="gene ID" value="ENSGAGG00000008780.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019838">
        <property type="term" value="F:growth factor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00054">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000194">
        <property type="entry name" value="Latent transforming growth factor beta binding protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000469">
        <property type="entry name" value="Latent transforming growth factor beta binding protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.290.10:FF:000004">
        <property type="entry name" value="latent-transforming growth factor beta-binding protein 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000019">
        <property type="entry name" value="latent-transforming growth factor beta-binding protein 1 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000046">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 1 isoform x2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000014">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.290.10:FF:000001">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 3 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.290.10:FF:000002">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 3 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000056">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 3 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000115">
        <property type="entry name" value="latent-transforming growth factor beta-binding protein 4 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000024">
        <property type="entry name" value="Putative latent-transforming growth factor beta-binding protein 2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000273">
        <property type="entry name" value="Putative latent-transforming growth factor beta-binding protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="18"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.290.10">
        <property type="entry name" value="TGF-beta binding (TB) domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026823">
        <property type="entry name" value="cEGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001881">
        <property type="entry name" value="EGF-like_Ca-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013032">
        <property type="entry name" value="EGF-like_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000152">
        <property type="entry name" value="EGF-type_Asp/Asn_hydroxyl_site"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018097">
        <property type="entry name" value="EGF_Ca-bd_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009030">
        <property type="entry name" value="Growth_fac_rcpt_cys_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049883">
        <property type="entry name" value="NOTCH1_EGF-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017878">
        <property type="entry name" value="TB_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036773">
        <property type="entry name" value="TB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052080">
        <property type="entry name" value="vWF_C/EGF_Fibrillin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47333:SF4">
        <property type="entry name" value="EGF-LIKE DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47333">
        <property type="entry name" value="VON WILLEBRAND FACTOR C AND EGF DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12662">
        <property type="entry name" value="cEGF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07645">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12661">
        <property type="entry name" value="hEGF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00683">
        <property type="entry name" value="TB"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="18"/>
    </dbReference>
    <dbReference type="SMART" id="SM00179">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="17"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57184">
        <property type="entry name" value="Growth factor receptor domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57581">
        <property type="entry name" value="TB module/8-cys domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00010">
        <property type="entry name" value="ASX_HYDROXYL"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01186">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01187">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51364">
        <property type="entry name" value="TB"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9 21" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="4 21" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="3" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11" id="KW-0340">Growth factor binding</keyword>
    <keyword evidence="6" id="KW-0358">Heparin-binding</keyword>
    <keyword evidence="12" id="KW-0379">Hydroxylation</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="26" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="60"/>
            <end position="92"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="254"/>
            <end position="286"/>
        </location>
    </feature>
    <feature type="domain" description="TB" evidence="24">
        <location>
            <begin position="401"/>
            <end position="444"/>
        </location>
    </feature>
    <feature type="domain" description="TB" evidence="24">
        <location>
            <begin position="521"/>
            <end position="573"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="672"/>
            <end position="713"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="714"/>
            <end position="750"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="757"/>
            <end position="796"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="837"/>
            <end position="877"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="950"/>
            <end position="990"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="1028"/>
            <end position="1068"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="1113"/>
            <end position="1154"/>
        </location>
    </feature>
    <feature type="domain" description="TB" evidence="24">
        <location>
            <begin position="1212"/>
            <end position="1266"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="1288"/>
            <end position="1326"/>
        </location>
    </feature>
    <feature type="domain" description="TB" evidence="24">
        <location>
            <begin position="1386"/>
            <end position="1438"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="1565"/>
            <end position="1609"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="1"/>
            <end position="31"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="371"/>
            <end position="396"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="594"/>
            <end position="617"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="22">
        <location>
            <begin position="8"/>
            <end position="18"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="22">
        <location>
            <begin position="371"/>
            <end position="395"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="21">
        <location>
            <begin position="64"/>
            <end position="74"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="21">
        <location>
            <begin position="82"/>
            <end position="91"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="21">
        <location>
            <begin position="258"/>
            <end position="268"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="21">
        <location>
            <begin position="276"/>
            <end position="285"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022674"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023183"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023278"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00038081"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00058734"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00059743"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00062144"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00062844"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00072992"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00072997"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00075443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS51364"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015529.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1621" mass="175767" checksum="6909E5294E853B59" modified="2019-05-08" version="1">KLSAPIHRSQSGASSQPCSPAGAGQARDPTPQRARLKCLEDQIQPLGWSHVSLFTPFHVPPAVCDPPCQNKGSCSRPQLCICRSGFQGAQCEEVVPERVYHPPGSVAALQPPAGSLPKRAGRLVGPEPPFPKFRLGKSWTQGRIVFPVSRHTGSSRTVRRYPAGNTQLMSNALPNGNGHEQSSSGPHAAYQEHTLSLWGANLTEKIRKIKIVFTPTICKQTCRSGHCYNSCQKGDTTTLYSQGGQNHDPKSGFRIYFCQIPCLNGGRCVGRDECWCPSNSTGKFCHLPVPKLEKNQVGRVPKTLASGSIKQSMYTLPLSNQLASLNPSLVNVHIKHPPEATVQIHQVARVRGDLGQSEENSVEAVLLSQLPTGPQHSYSNGNSNSITTESGQQQRPPELMGRCFRETLHGQCANPLEGLAKFEDCCGSVGLFWGVNRCIPCPPRPAHPVIENRQVECPQGYKRLNRSHCQDINECLMLGLCKDTECVNTRGSFLCTCKPGTMLDPSRSRCVSDKAVSMEQGLCYRSAAEGVCALPLAQSITKQICCCSRVGKGWGRNCEKCPMLGSETFKEICPAGHGYTYSSSDIRLSMRKAEAEELPFSSEEQGESSDQTPTQAAKRLQLQEALSGGVTGTLARAGTSAAEGVLGCDATLVAALHTGQWPVTSPCLSLSEMDRCASVPDPCGPGTCVNLPDRYSCICGPGYQLHPGLPQCIDDDECMRDPCAGKGHCINSVGSYSCLCYPGYTLLASQGTQTCQDLNECEQPSMCQGGQCTNTPGSYHCECARGYIMGHTGQCEDIDECADPSTCPRGRCVNALGSYECLSCAAGYQPMNGRCVDDDECFTVGICAHGLCINLAGSFRCSCYHGYEVTADEKSCQDINECATHNACPMGLCINSEGSFSCAACGAGYTVSRDGSMCEDIDECSSATACPLGICTNTDGSFACRACDADIDECEDPTVSCLGGECLNTPGSYVCHCRPGFELLNGTMCEAGGVTKCALDPGYLFIVQGPVYTHRTASVLIFILSLPDVDECADKSRCLRGQCFNTEGSYRCLCENGFRHSQETDDCVDVDECVDFGDTVCGTWRCQNTPGSYRCVMGCQPGFHRTPLGDCIDIDECANETLCGSHAFCDNTDGSFRCLCDSGYENLPPGQDCVDVNECELMVAVCGTALCENVEGSFLCLCLSDHEEYNTEAGECRPRAGMAPRPAGSERKECYYHISDVQLCDSVLAKNTTKEECCCTEGAAWGDNCETHPCPILGTVEYSEICPSGKGYIPVEGTLLFGQTSYTDADECEMFGPEICRNGHCVNTVPGYKCFCRSGYLYDSSRLEYQDECENEMSCVNGECLNTAGSFHCFCGLPLVLDVTRNRCINVSSRAEDPEEHEIHLDICWQRVSNYICNQPLRGRRTTYTECCCRYGEAWSQDCALCPHRASEDFAHLCNVARGETEREPGLQERPGYEYGPALEDSHYGLYSPDLGPYYNYLSPEYGAPDSTWTGPHLCYPDQYPYLLPLSPLHHPPASFEGLQAEECGILNGCENGRCVRVREGYTCDCFDGFQLDMTHMACVDINECEEVSSSGPLCEGGTCENTEGSYRCNCSPGFVVQAEPHRCAPAPAQRPEAAQL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HL00</accession>
    <name>A0A452HL00_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Ninein-like protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Microtubule organizing center</location>
            <location evidence="1">Centrosome</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HL00"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017830.1">
        <property type="protein sequence ID" value="ENSGAGP00000015610.1"/>
        <property type="gene ID" value="ENSGAGG00000011767.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005874">
        <property type="term" value="C:microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034454">
        <property type="term" value="P:microtubule anchoring at centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00051">
        <property type="entry name" value="EFh"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.238.10:FF:000094">
        <property type="entry name" value="ninein isoform X7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.238.10:FF:000209">
        <property type="entry name" value="Ninein like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.238.10">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011992">
        <property type="entry name" value="EF-hand-dom_pair"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018247">
        <property type="entry name" value="EF_Hand_1_Ca_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002048">
        <property type="entry name" value="EF_hand_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18905">
        <property type="entry name" value="NINEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18905:SF12">
        <property type="entry name" value="NINEIN-LIKE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13499">
        <property type="entry name" value="EF-hand_7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00054">
        <property type="entry name" value="EFh"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47473">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00018">
        <property type="entry name" value="EF_HAND_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50222">
        <property type="entry name" value="EF_HAND_2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0106">Calcium</keyword>
    <keyword evidence="9 12" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="10" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="5" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="4" id="KW-0493">Microtubule</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="8" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="EF-hand" evidence="14">
        <location>
            <begin position="8"/>
            <end position="43"/>
        </location>
    </feature>
    <feature type="domain" description="EF-hand" evidence="14">
        <location>
            <begin position="198"/>
            <end position="233"/>
        </location>
    </feature>
    <feature type="domain" description="EF-hand" evidence="14">
        <location>
            <begin position="235"/>
            <end position="270"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="149"/>
            <end position="168"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="12">
        <location>
            <begin position="291"/>
            <end position="325"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="12">
        <location>
            <begin position="355"/>
            <end position="382"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="12">
        <location>
            <begin position="413"/>
            <end position="615"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="12">
        <location>
            <begin position="648"/>
            <end position="909"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="13">
        <location>
            <begin position="152"/>
            <end position="162"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00071185"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50222"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015610.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="913" mass="106967" checksum="1D71BCF7E986C62D" modified="2019-05-08" version="1">MGEEAENKYVSQLKDVYNSCDTTGTGYLDKEELTELCRKLHLERQLPVLLQTLLGRNCFARVNFEEFKEGFVAVLSSNIDLGISDEESSYLEPAVPDDVKPKYVNGAKWYGRWTKPELQETEVETTKYLQDQQVKASVKSQLRRSASLESVESLKSDEEAQSAKESQNEMLAAQGQMGTWNPDLFDSPRRTASPCFDMMEGQIRGIWEELDVGNTGYLSKCELATVCKNIGLKELEQEELEDLFRRLDRDGDGRVSFQEFELGLSSHGPVSFPVSSTPFKQLRQPPDHQENRRLQSEVTEVVEKLSESEKRVSKLQKELDFVLKDKLDMVDPHSTELLHQEERFAEIITEYELQSRELWDKNDELQIELEKLRSQLQESQYRRTWRKMKENKPGGRLLPAGVLFVEAGACAVSIETELMIEQLKEQLQDLKIQLETKVNYYEREIELMKRNFEKERKDIEQGFKMEISELEEQKGDLEELKVKYQEVIGGLKDQLQKSTPSQELEKRFEKERSEMEQYYAKEICNLGQRLAQERDQLEEEMKMQHKNELQLTRMELHRVSEDNALLRNKLRRFQEDVEEASNKQRKQIDELQREKEMALSETEELNQLNKTYKQEICQLNARTLQLSSQLSDLSLQSETSHSTIQLLNQRLAETVSQKEEEAAVAKQLQETSSKLELEFGQHQSAWQQERELLHQQLRESREKVTQVQELEVELERVTQECQTLRLMAAQFREELEESQDQLLEANTRLSLAQSQHRQELQQLKDHLESTVSRDCLAQLQSRWAEEQRKVQQLQESLSFQAEQASRQLATFQEDHERLLRTMEERTEELEWDVRNMQRMLQEMVTQLQEQRDRNMKCDLLLKDLYVENAQLVKALQATEQRQKNAEKKNLALEEKLSALTELIRRMAQASLSV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HL36</accession>
    <name>A0A452HL36_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Ubiquinol-cytochrome c chaperone domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the CBP3 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HL36"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017871.1">
        <property type="protein sequence ID" value="ENSGAGP00000015646.1"/>
        <property type="gene ID" value="ENSGAGG00000011771.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034551">
        <property type="term" value="P:mitochondrial respiratory chain complex III assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021150">
        <property type="entry name" value="Ubiq_cyt_c_chap"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007129">
        <property type="entry name" value="Ubiqinol_cyt_c_chaperone_CPB3"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12184">
        <property type="entry name" value="UBIQUINOL-CYTOCHROME C REDUCTASE COMPLEX ASSEMBLY FACTOR 1 FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12184:SF1">
        <property type="entry name" value="UBIQUINOL-CYTOCHROME-C REDUCTASE COMPLEX ASSEMBLY FACTOR 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03981">
        <property type="entry name" value="Ubiq_cyt_C_chap"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Ubiquinol-cytochrome c chaperone" evidence="2">
        <location>
            <begin position="50"/>
            <end position="185"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006407"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF03981"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015646.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="213" mass="24577" checksum="59E31A83C6535F89" modified="2019-05-08" version="1">EEGPVVRIKKVFGGGHGEVAQGSVTKIKIAALRMYTCCVEKTDYEEFFHRCHMPDTLNSWFLIAQLHVWMCLVRMKQEGRAGKYMCRYIVHSMWEDVEQRGKVMGIDSFTLKQSMRSMTENFYAAIFGYDEGILSDDHVLAAALWRNLFNKHCEDPRQLELLVEYVRKQMQYLDAINGEDLLLTGEVNWRPLVESNAQSILKPTSPAYNDEGL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HL37</accession>
    <name>A0A452HL37_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">ubiquitinyl hydrolase 1</fullName>
            <ecNumber evidence="3">3.4.19.12</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>Thiol-dependent hydrolysis of ester, thioester, amide, peptide and isopeptide bonds formed by the C-terminal Gly of ubiquitin (a 76-residue protein attached to proteins as an intracellular targeting signal).</text>
            <dbReference type="EC" id="3.4.19.12"/>
        </reaction>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the peptidase C19 family.</text>
    </comment>
    <dbReference type="EC" id="3.4.19.12" evidence="3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017886.1">
        <property type="protein sequence ID" value="ENSGAGP00000015658.1"/>
        <property type="gene ID" value="ENSGAGG00000011442.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004843">
        <property type="term" value="F:cysteine-type deubiquitinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016579">
        <property type="term" value="P:protein deubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009966">
        <property type="term" value="P:regulation of signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd02659">
        <property type="entry name" value="peptidase_C19C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.70.10">
        <property type="entry name" value="Cysteine proteinases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056850">
        <property type="entry name" value="ARM_UBP34_24_USP9X_Y"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021905">
        <property type="entry name" value="DUF3517"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038765">
        <property type="entry name" value="Papain-like_cys_pep_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050164">
        <property type="entry name" value="Peptidase_C19"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001394">
        <property type="entry name" value="Peptidase_C19_UCH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028889">
        <property type="entry name" value="USP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018200">
        <property type="entry name" value="USP_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24006">
        <property type="entry name" value="UBIQUITIN CARBOXYL-TERMINAL HYDROLASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24006:SF827">
        <property type="entry name" value="UBIQUITIN CARBOXYL-TERMINAL HYDROLASE 34"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25010">
        <property type="entry name" value="ARM_UBP24_USP9X-Y"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12030">
        <property type="entry name" value="DUF3517"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00443">
        <property type="entry name" value="UCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54001">
        <property type="entry name" value="Cysteine proteinases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00972">
        <property type="entry name" value="USP_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00973">
        <property type="entry name" value="USP_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50235">
        <property type="entry name" value="USP_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="4" id="KW-0645">Protease</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0788">Thiol protease</keyword>
    <keyword evidence="5" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="USP" evidence="9">
        <location>
            <begin position="1616"/>
            <end position="1961"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="227"/>
            <end position="251"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="267"/>
            <end position="359"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="374"/>
            <end position="400"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="3055"/>
            <end position="3194"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="235"/>
            <end position="250"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="276"/>
            <end position="286"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="287"/>
            <end position="306"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="307"/>
            <end position="325"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="8">
        <location>
            <begin position="326"/>
            <end position="346"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="3059"/>
            <end position="3070"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="3157"/>
            <end position="3179"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000707"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009085"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012759"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022807"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50235"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015658.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="3270" mass="372784" checksum="3BA239997729AA4D" modified="2025-10-08" version="2">MVFLLISRIFSKICVEIMVRMVIDLRSLMFQYLRQYAQDSFVPETYILGKSYCNSYKSDFHKFKIVNQLHTFNDVCNNESLVSDTETSIAKELADWLISNNVVEHIFGPNLHIEIIKQCQVILNFLAAEGRLSTQHIDCIWAAAQLKHCSRYIHDLFPSLIKNLDPVPLRHLLNLVSTLHPSAHTEQTLYLASMLIKALWNNALAAKAQLSKQSSFASLLSTNLPMGNKKGSPAASPQSSDNSDTHQSGGSDIEMEEQIINRNKHVQQRLSDTEESMQGSSDETANSGEDGSSGPGSSSGHSDGSSNEANSSHASQSAGSPGSEVQSEDIADIEALKEDDEEEEEDRGPNPQKNSRSTELLSRKIESQAGICLGDSQGAADRSGASNGSGKDHVFNTESMPPVDSRIRMLDACSHTEDAEHDMTGEMNPAHISQGSQDSCITHTGDFLGETIGNELFNCRQFIGPQHHHHHHHHHHHHHHHDGHMVDDMLSADDVSCSSSQVSAKSEKNMADFDGEESGCEEELVQINSHAELTSHLQQHLPNLASIYHEHLSQGPAVHKHQYNSNAVTDINLDNVCKKGNTLLWDLVQDEDAIHLSEGLINEAEKLLCSLVCWFTDRQIRMRFIEGCLENLANNRSVVVSLRLLPKLFGTFQQFGSSYDTHWITMWAEKELNMMKLFFDNLIHYIQAVREGRHKHALYSHSAEVQVRLQFLTCVFSTLGSPDHFRLSLEQVDILWHCLVEDSECYDDALHWFLNQVRSKDQHAMGMETYKHLFLEKMPQLKPETISMTGLNLFQHLCNLARLATSAYDGGSNSELCGMDQFWGIALRAQSGDVSRAAIQYINSYYINGKTGLEKEQEFISKCMESLMIASSNLEQDSHSSLTIIERGLLMLKTHLEAFRRRFAYHLRQWQIEGTGISSHLKALSDKQSLPLRIVCQPAGLPDKMTIEMYPSDQVADLRAEVTHWYENLQKEQINQQAQLQEFGQSSRKGDFPGGLMGPVRMISSGHELTTDYDEKTLHELGFKDMQMVFVSLGAPRRERKGEGVQLPASCLPPPQKDNIPMLLLLQEPHLTTLFDLLEMLASFKPPSGEMAMEDSESARCEELHLHAENLSRRVWELLMLLPTCPNMLQAFQNISDEQGNDGFSWKDLLRIKSAHKLLYALEIIEALGKPNRRIRRESTGSYSDLYPDSDDSSEDQIENSKNTWSCKFVAAGGLQQLLEIFNSGILEPKEQEPWTVWQLDCLACLLKLICQFAVDPSDLDLAYHDVFAWSGVAESHRKRTWPGKSRKTAGDHVKGLHIPRLTEVFLVLVQGTSLIQRLMSVAYTYDNLAPRVLKAQSDHRSRHEVTHYSMWLLVSWAHCCSLVKSSLADSDNLQDWLKKLTLLIPETAVRHESCNGLYKLSLCGLDGGDTINRSFLLLAASTLLKFLPDAQALKPIRIEDYEEEPVLRSGCKEYFWLLCKLIDNIHVKDASQTTLLDLDALARHLADCIRSREILDHQDGNIEDDGLTGLLRLATSVIKHKPPFKFSREGQEFLRDIFNLLFLLPSLKDRQQPKCKSHSSRAAAYDLLVEMVKGSVENYRLLHNWVMAQHMQSSHAPYKWDYWPHDDVRAECRFVGLTNLGATCYLASTIQQLYMIPEARQAIFTAKYSEDMKHKTTLLELQKMFTYLMESECKAYNPRPFCKTYTMDKQPLNTGEQKDMTEFFTDLITKIEEMSPELKNTVKSLFGGVITNNVVSLDCEHVSQTAEEFYTVRCQVADMKNIYESLDEVTIKDTLEGDNMYTCSHCGKKVRAEKRACFKKLPRILSFNTMRYTFNMVTMMKEKVNTHFSFPLRLDMTPYTEDFLMGKNDRKEGFKDNSEYLKETESYEYDLIGVTVHTGTADGGHYYSFIRDIVNPHAYKNNKWYLFNDAEVKPFDSAQLASECFGGEMTTKTYDSVTDKFMDFSFEKTHSAYMLFYKRMEPEEENGKDYKFDVSSELLEWIWHDNMQFLQDKNIFEHTYFGFMWQLCSSIPSTLPDPKAVSLMTAKLSTSFVLETFIHSKEKPTMLQWIELLTKQFNNSQAACEWFLDRMADDDWWPMQILIKCPNQIVRQMFQRLCIHVIQRLRPVHAHLYLQPGMEDCSDDIDGPVEDIGSRSCVTRFVKTLLSIMEHGVKPHSKHLTEYFAFLYEFAKMGEEESQFLLSLQAISTMVHFYMGTKGPENPQVEVLSEEEGEEEEEEEDILSLAEEKYRPAALEKMIALIALLVEQSRSERHLTLSQNDMAALTGGKGFPFLFQHIRDGINIRQTCNLIFSLCRYNNRLAEHIVSMLFTSIAKLTPEAANPFFKLLTMLMEFAGGPPGMPPFASYILQRIWEVIEYNPSQCLDWLAVQTPRNKLAHSWVLQNMENWVERFLLAHNYPRVRTSAAYLLVSLIPSNSFRQMFRSTRSLHIHTRDLPLSPDTTVVLHQVYNVLLGLLSRAKLYVDAAVHGTTKLVPYFSFMTYCLISKTEKLMFSTYFMDLWNLFQPKLSEPAIATNHNKQALLSFWYNGCVDCPENVRLIVQNPVVTKNIAFNYILADHDDQDVVLFNRGMLPAYYGILRLCCEQSPAFTRQLASHQNIQWAFKNLTPHASQYPGAVEELFNLMQLFVAQRPDMREEEIEDIKQFKKTTISCYLRCLDGRSCWTTLISAFRILLESDEDRLLVVFNRGLILMTESFNTLHMMYHEATACHVTGDLVELLSIFLSVLKSTRPYLQRKDVKQALIQWQERIEFAHKLLTLLNSYSPSELRNACIDVLKELVLLSPHDFLHTLVPFLQHNHCTYHHSNIPMSLGPYFPCRENLKLIGGKSNIRPPRPELNMCLLPTMMEASKGKDDVYDRMLLDYFFSYHQFIHLLCRVAINCEKFTETLVKMSVLVAYEGLPLHLALFPKLWTELCQTQSAMSKNCIKLLCEDPVFAEYIKCILMDERTFLNNNIVYTFLTHFLLKVQGQVFSEANCANLINTLVTNLINQYQSLESDFSSQRVEISKASSTLNGDLRALALLLSVHTPKQLNSALIPTLQELLNKCRACQQQRNSLQEQEAKERKTKDDEGATPVKRRRVSSDEEHTVDSCISDMKTEPRDTLTPTSTSDNETRDSSIIDPGTEQDPPSPENSSVKEYRMEVPSSFSEEMMLTTRSQHTEEQSSNGKFEECKEFKDLQSSKDPSAVEDDSEFPSTSISAVLSDLADLRGCDGQALPSQDPENSLSISCGHSRGPFSHMQQHDILDILCRTIESTIHVVTRISSKGNQAAS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HL40</accession>
    <name>A0A452HL40_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Glycerol-3-phosphate dehydrogenase [NAD(+)]</fullName>
            <ecNumber evidence="11">1.1.1.8</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="6">
            <text>sn-glycerol 3-phosphate + NAD(+) = dihydroxyacetone phosphate + NADH + H(+)</text>
            <dbReference type="Rhea" id="RHEA:11092"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57540"/>
            <dbReference type="ChEBI" id="CHEBI:57597"/>
            <dbReference type="ChEBI" id="CHEBI:57642"/>
            <dbReference type="ChEBI" id="CHEBI:57945"/>
            <dbReference type="EC" id="1.1.1.8"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="6">
            <dbReference type="Rhea" id="RHEA:11093"/>
        </physiologicalReaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 10">Belongs to the NAD-dependent glycerol-3-phosphate dehydrogenase family.</text>
    </comment>
    <dbReference type="EC" id="1.1.1.8" evidence="11"/>
    <dbReference type="AlphaFoldDB" id="A0A452HL40"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017878.1">
        <property type="protein sequence ID" value="ENSGAGP00000015651.1"/>
        <property type="gene ID" value="ENSGAGG00000011806.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0141152">
        <property type="term" value="F:glycerol-3-phosphate dehydrogenase (NAD+) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051287">
        <property type="term" value="F:NAD binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042803">
        <property type="term" value="F:protein homodimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005975">
        <property type="term" value="P:carbohydrate metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046168">
        <property type="term" value="P:glycerol-3-phosphate catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.720:FF:000088">
        <property type="entry name" value="Glycerol-3-phosphate dehydrogenase [NAD(+)]"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.1040.10:FF:000084">
        <property type="entry name" value="Glycerol-3-phosphate dehydrogenase [NAD(+)], cytoplasmic"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1040.10">
        <property type="entry name" value="N-(1-d-carboxylethyl)-l-norvaline Dehydrogenase, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.720">
        <property type="entry name" value="NAD(P)-binding Rossmann-like Domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008927">
        <property type="entry name" value="6-PGluconate_DH-like_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013328">
        <property type="entry name" value="6PGD_dom2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006168">
        <property type="entry name" value="G3P_DH_NAD-dep"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006109">
        <property type="entry name" value="G3P_DH_NAD-dep_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017751">
        <property type="entry name" value="G3P_DH_NAD-dep_euk"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011128">
        <property type="entry name" value="G3P_DH_NAD-dep_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036291">
        <property type="entry name" value="NAD(P)-bd_dom_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR03376">
        <property type="entry name" value="glycerol3P_DH"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="GLYCEROL-3-PHOSPHATE DEHYDROGENASE"/>
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        <property type="entry name" value="GLYCEROL-3-PHOSPHATE DEHYDROGENASE 1-LIKE PROTEIN"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="NAD_Gly3P_dh_C"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="NAD_Gly3P_dh_N"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Glycerol-3-P_dh"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="GPDHDRGNASE"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48179">
        <property type="entry name" value="6-phosphogluconate dehydrogenase C-terminal domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="NAD(P)-binding Rossmann-fold domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="5 9 10" id="KW-0520">NAD</keyword>
    <keyword evidence="4 10" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Glycerol-3-phosphate dehydrogenase NAD-dependent N-terminal" evidence="12">
        <location>
            <begin position="15"/>
            <end position="169"/>
        </location>
    </feature>
    <feature type="domain" description="Glycerol-3-phosphate dehydrogenase NAD-dependent C-terminal" evidence="13">
        <location>
            <begin position="193"/>
            <end position="339"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="7">
        <location>
            <position position="204"/>
        </location>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="41"/>
        </location>
        <ligand>
            <name>NAD(+)</name>
            <dbReference type="ChEBI" id="CHEBI:57540"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="97"/>
        </location>
        <ligand>
            <name>NAD(+)</name>
            <dbReference type="ChEBI" id="CHEBI:57540"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="8">
        <location>
            <position position="120"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
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    <feature type="binding site" evidence="9">
        <location>
            <position position="153"/>
        </location>
        <ligand>
            <name>NAD(+)</name>
            <dbReference type="ChEBI" id="CHEBI:57540"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="8">
        <location>
            <begin position="269"/>
            <end position="270"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
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    <feature type="binding site" evidence="9">
        <location>
            <position position="269"/>
        </location>
        <ligand>
            <name>NAD(+)</name>
            <dbReference type="ChEBI" id="CHEBI:57540"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="296"/>
        </location>
        <ligand>
            <name>NAD(+)</name>
            <dbReference type="ChEBI" id="CHEBI:57540"/>
        </ligand>
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    <feature type="binding site" evidence="9">
        <location>
            <position position="298"/>
        </location>
        <ligand>
            <name>NAD(+)</name>
            <dbReference type="ChEBI" id="CHEBI:57540"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00011009"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023027"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00048723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PIRSR" id="PIRSR000114-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PIRSR" id="PIRSR000114-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PIRSR" id="PIRSR000114-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="RuleBase" id="RU000437"/>
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    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="RuleBase" id="RU361243"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF01210"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="Pfam" id="PF07479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015651.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="349" mass="38413" checksum="D946684432E2B1EC" modified="2019-05-08" version="1">MLHLCKVGLSINHRGSAVAKIIGNNVKNMQKFASTVKMWVFEENINGRKLTDIINNEHENVKYLPGYKLPENVVAIPNLNEAVQDADLLVFVIPHQFIHKICDEITGRVPKKAVGITLIKGIDEGPEGLKLISDIIREKMGIDISVLMGANIANEVAAEKFCETTIGSKILENGLLFKELLQTPNFRITVVDDADTVELCGALKNIVAVGAGFCDGLRCGDNTKAAVIRLGLMEMIAFAKIFCKGQVSIATFLESCGVADLITTCYGGRNRRVAEAFVKTGKSIEELEQEMLNGQKLQGPQTSAEVYRILKQKGMVDKFPLFTAVYQICYEGKPVQEMISCLQSHPEHM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INN0</accession>
    <name>A0A452INN0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Macro domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
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            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Chromosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452INN0"/>
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        <property type="protein sequence ID" value="ENSGAGP00000029694.1"/>
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    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
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        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
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        <property type="project" value="UniProtKB-KW"/>
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        <property type="term" value="F:protein heterodimerization activity"/>
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        <property type="project" value="InterPro"/>
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        <property type="term" value="F:structural constituent of chromatin"/>
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        <property type="project" value="InterPro"/>
    </dbReference>
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        <property type="term" value="P:chromatin organization"/>
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        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
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        <property type="entry name" value="HFD_H2A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.20.10:FF:000013">
        <property type="entry name" value="Core histone macro-H2A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.220.10:FF:000002">
        <property type="entry name" value="Core histone macro-H2A"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Histone, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.220.10">
        <property type="entry name" value="Leucine Aminopeptidase, subunit E, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="H2A/H2B/H3"/>
    </dbReference>
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        <property type="entry name" value="Histone-fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002119">
        <property type="entry name" value="Histone_H2A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032454">
        <property type="entry name" value="Histone_H2A_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002589">
        <property type="entry name" value="Macro_dom"/>
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        <property type="entry name" value="Macro_dom-like"/>
    </dbReference>
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        <property type="entry name" value="HISTONE H2A"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Macro"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00620">
        <property type="entry name" value="HISTONEH2A"/>
    </dbReference>
    <dbReference type="SMART" id="SM00506">
        <property type="entry name" value="A1pp"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00414">
        <property type="entry name" value="H2A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47113">
        <property type="entry name" value="Histone-fold"/>
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    </dbReference>
    <dbReference type="SUPFAM" id="SSF52949">
        <property type="entry name" value="Macro domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="MACRO"/>
        <property type="match status" value="1"/>
    </dbReference>
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    <keyword evidence="6" id="KW-0156">Chromatin regulator</keyword>
    <keyword evidence="3" id="KW-0158">Chromosome</keyword>
    <keyword evidence="7" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="4" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="9" id="KW-0544">Nucleosome core</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="Macro" evidence="11">
        <location>
            <begin position="183"/>
            <end position="314"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="128"/>
            <end position="179"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="10">
        <location>
            <begin position="144"/>
            <end position="159"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="164"/>
            <end position="176"/>
        </location>
    </feature>
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        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00004286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022454"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023269"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS51154"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029694.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="314" mass="33132" checksum="7ED1C978BC00E51C" modified="2019-05-08" version="1">MSSRGGKKKSTKTSRSAKAGVIFPVGRMLRYIKKGHPKYRIGVGAPVYMAAVLEYLTAEILELAGNAARDNKKGRVTPRHILLAVANDEELNQLLKGVTIASGGVLPNIHPELLAKKRGSKGKLEAIITPPPAKKAKSPSQKKTVSKKAGGKKGSRKSKKQGEVSKSASADSTTEGTPADGFTVLSTKSLFLGQKLQVVQADIATIDSDAVVHPTNSDFYTGGEVGSTLEKKGGKEFVEAVIELRKKNGPLDIAGAVVSAGHGLPAKFVIHCNSPGWGSDKCEELLEKTVKNCLALADEKKLKSIAFPSIGSGR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INP4</accession>
    <name>A0A452INP4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Acrosin-binding protein</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="6">Acrosin-binding protein, 60 kDa form</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="7">Proacrosin-binding protein sp32</fullName>
        </alternativeName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="8">Acrosomal protein that maintains proacrosin (pro-ACR) as an enzymatically inactive zymogen in the acrosome. Involved also in the acrosome formation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasmic vesicle</location>
            <location evidence="1">Secretory vesicle</location>
            <location evidence="1">Acrosome</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452INP4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033735.1">
        <property type="protein sequence ID" value="ENSGAGP00000029712.1"/>
        <property type="gene ID" value="ENSGAGG00000021441.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001669">
        <property type="term" value="C:acrosomal vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002350">
        <property type="entry name" value="Kazal_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009865">
        <property type="entry name" value="Proacrosin-bd"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21362">
        <property type="entry name" value="ACROSIN-BINDING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21362:SF1">
        <property type="entry name" value="ACROSIN-BINDING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07648">
        <property type="entry name" value="Kazal_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07222">
        <property type="entry name" value="PBP_sp32"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <feature type="domain" description="Kazal-like" evidence="10">
        <location>
            <begin position="372"/>
            <end position="407"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="115"/>
            <end position="158"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="135"/>
            <end position="146"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004218"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00018940"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00032734"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00033453"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00045517"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="Pfam" id="PF07648"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029712.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="438" mass="49038" checksum="DA461814C4C4D155" modified="2025-10-08" version="2">MPKPRLLLPMVQPVWSMSSLVPGSPLSDSEYQVFFSALMPSWKAGVACQIRRTQGCHDPKVIRLDQFENHGWIPEGPICSDLPDASWFQTFCQFAHYRCSNRKYYTKVSHIPDVPTQRSLKPGGDPELAGPLQVQPVTATPETSPSAMKPSVKTGRATEQHMQKSIQQLINSALSLEGSPEIETTQVQCSTLCSSLKEPAGNYRSQKLTCCSFLCFLLCISLLALERDEALVILCYAMLEGICLSSAVTQAWKRMEARTLGFGDSVCDSLGRHHMELCPDCAFCSLKREQCRGVSDLRRVHCDGHSPVSNAPLCLQVGSLEAMGYYGMEVYGGLRGDYWCGRLATHGCDDPRVILWLQVEYAFFQGGDFPNKICDSDRVQHPNYCAFKSHQCLQHSLSSQKVLRYGCRRNETYQVLSEEKGEEEVLLWSQKFFSLTEG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INP7</accession>
    <name>A0A452INP7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="24">Dual specificity mitogen-activated protein kinase kinase 7</fullName>
            <ecNumber evidence="20">2.7.12.2</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="26">JNK-activating kinase 2</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="25">MAPK/ERK kinase 7</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="27">c-Jun N-terminal kinase kinase 2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="31 32">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="32" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="31" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="31" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="21">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.12.2"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="22">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.12.2"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="23">
            <text>L-tyrosyl-[protein] + ATP = O-phospho-L-tyrosyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10596"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.12.2"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="19">Belongs to the protein kinase superfamily. STE Ser/Thr protein kinase family. MAP kinase kinase subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.12.2" evidence="20"/>
    <dbReference type="AlphaFoldDB" id="A0A452INP7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033522.1">
        <property type="protein sequence ID" value="ENSGAGP00000029522.1"/>
        <property type="gene ID" value="ENSGAGG00000021324.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019899">
        <property type="term" value="F:enzyme binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000287">
        <property type="term" value="F:magnesium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004708">
        <property type="term" value="F:MAP kinase kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004713">
        <property type="term" value="F:protein tyrosine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006915">
        <property type="term" value="P:apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043410">
        <property type="term" value="P:positive regulation of MAPK cascade"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006950">
        <property type="term" value="P:response to stress"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd06618">
        <property type="entry name" value="PKc_MKK7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000040">
        <property type="entry name" value="Dual specificity mitogen-activated protein kinase kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000214">
        <property type="entry name" value="Dual specificity mitogen-activated protein kinase kinase 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052468">
        <property type="entry name" value="Dual_spec_MAPK_kinase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47238:SF2">
        <property type="entry name" value="DUAL SPECIFICITY MITOGEN-ACTIVATED PROTEIN KINASE KINASE HEMIPTEROUS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47238">
        <property type="entry name" value="MITOGEN-ACTIVATED PROTEIN KINASE KINASE 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="14" id="KW-0007">Acetylation</keyword>
    <keyword evidence="8" id="KW-0053">Apoptosis</keyword>
    <keyword evidence="12" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="16 28" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11" id="KW-0418">Kinase</keyword>
    <keyword evidence="13" id="KW-0460">Magnesium</keyword>
    <keyword evidence="9" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="18" id="KW-0539">Nucleus</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="32" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="15" id="KW-0346">Stress response</keyword>
    <keyword evidence="7" id="KW-0808">Transferase</keyword>
    <keyword evidence="17" id="KW-0829">Tyrosine-protein kinase</keyword>
    <feature type="domain" description="Protein kinase" evidence="30">
        <location>
            <begin position="136"/>
            <end position="396"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="29">
        <location>
            <begin position="63"/>
            <end position="96"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="28">
        <location>
            <begin position="3"/>
            <end position="30"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="29">
        <location>
            <begin position="68"/>
            <end position="77"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022703"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023016"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023137"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00038035"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00038999"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00049014"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00049299"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00051693"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00073834"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00077430"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00081151"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="ARBA" id="ARBA00083185"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="30">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="31">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029522.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="32">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="436" mass="49368" checksum="13E5363E177C84EF" modified="2019-05-08" version="1">MAASSLEQKLSRLEAKLKQENREARRKIDLNLEISPARARPIIVITLSPAPAPSQRAALQLPLANDGSSRSSSSENSPQHQTHQPRRPLLGLPQPSFPIPRSMESIEIDQKLQEIMKQTGYLTIGGQRYQAEINDLENLGEIGSGTCGQVWKMRFKKTGHIIAVKQMRRSGNKEENKRILMDLDVVLKSHDCPYIVQCFGTFITNTDVFIAMELMGTCAEKLKKRIQGPIPERILGKMTVAIVKALYYLKEKHGVIHRDVKPSNILLDERGQIKLCDFGISGRLVDSKAKTRSAGCAAYMAPERIDPPDPTKPDYDIRADVWSLGISLVELATGQFPYKNCKTDFEVLTKVLQEDPPLLPNNMSFSVDFQSFVKDCLTKDHRKRPKYNKLLEHNFIKRYETLEVDVASWFKDVMAKTESPRTSSILSQHHLPFFTR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INQ5</accession>
    <name>A0A452INQ5_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
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            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
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        <property type="protein sequence ID" value="ENSGAGP00000029495.1"/>
        <property type="gene ID" value="ENSGAGG00000021299.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
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        <property type="term" value="P:mRNA 3'-end processing"/>
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        <property type="project" value="InterPro"/>
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        <property type="entry name" value="WD repeat domain 33"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
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        <property type="entry name" value="Pfs2-like"/>
    </dbReference>
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        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
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    <dbReference type="PANTHER" id="PTHR22836:SF0">
        <property type="entry name" value="PRE-MRNA 3' END PROCESSING PROTEIN WDR33"/>
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    </dbReference>
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        <property type="match status" value="2"/>
    </dbReference>
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        <property type="match status" value="3"/>
    </dbReference>
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    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
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    <feature type="repeat" description="WD" evidence="1">
        <location>
            <begin position="157"/>
            <end position="189"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="1">
        <location>
            <begin position="198"/>
            <end position="230"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029495.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="270" mass="31930" checksum="68BF236CDF67E1F1" modified="2019-05-08" version="1">MATEIGSPPRFFHMPRFQHQAPRQLFYKRPDFAQQQAMQQLTFDGKRMRKAVNRKTIDYNPSVIKYLENRVWQRDQRDMRAIQPDAGYYNDLVPPIGMLNNPMNAVTTKFVRTSTNKVKCPVFVVRWTPEGRRLVTGASSGEFTLWNGLTFNFETILQAHDSPVRAMTWSHNDMWMLTADHGGYVKYWQSNMNNVKMFQAHKEAIREASFSPTDNKFATCSDDGTVRIWDFLRCHEERILRDLWPTWISTHDNMDEEREYLNYWTTVLRS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INT1</accession>
    <name>A0A452INT1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Phospholipase A2</fullName>
            <ecNumber evidence="8">3.1.1.4</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
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        <lineage>
            <taxon>Eukaryota</taxon>
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            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="8">
            <text>a 1,2-diacyl-sn-glycero-3-phosphocholine + H2O = a 1-acyl-sn-glycero-3-phosphocholine + a fatty acid + H(+)</text>
            <dbReference type="Rhea" id="RHEA:15801"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:28868"/>
            <dbReference type="ChEBI" id="CHEBI:57643"/>
            <dbReference type="ChEBI" id="CHEBI:58168"/>
            <dbReference type="EC" id="3.1.1.4"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="5">
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </cofactor>
        <text evidence="5">Binds 1 Ca(2+) ion per subunit.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 8">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="7">Belongs to the phospholipase A2 family.</text>
    </comment>
    <dbReference type="EC" id="3.1.1.4" evidence="8"/>
    <dbReference type="AlphaFoldDB" id="A0A452INT1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033782.1">
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        <property type="gene ID" value="ENSGAGG00000021477.1"/>
    </dbReference>
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        <property type="project" value="UniProtKB-SubCell"/>
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        <property type="project" value="InterPro"/>
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        <property type="term" value="F:calcium-dependent phospholipase A2 activity"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="F:phospholipid binding"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="P:arachidonate secretion"/>
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        <property type="project" value="InterPro"/>
    </dbReference>
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        <property type="term" value="P:lipid catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006644">
        <property type="term" value="P:phospholipid metabolic process"/>
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        <property type="project" value="InterPro"/>
    </dbReference>
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        <property type="entry name" value="PLA2c"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.90.10:FF:000001">
        <property type="entry name" value="Basic phospholipase A2 homolog"/>
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    </dbReference>
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        <property type="entry name" value="Phospholipase A2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="PLA2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016090">
        <property type="entry name" value="PLA2-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033112">
        <property type="entry name" value="PLA2_Asp_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033113">
        <property type="entry name" value="PLA2_histidine"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036444">
        <property type="entry name" value="PLipase_A2_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11716:SF8">
        <property type="entry name" value="GROUP IIF SECRETORY PHOSPHOLIPASE A2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11716">
        <property type="entry name" value="PHOSPHOLIPASE A2 FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00068">
        <property type="entry name" value="Phospholip_A2_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00389">
        <property type="entry name" value="PHPHLIPASEA2"/>
    </dbReference>
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        <property type="entry name" value="PA2c"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48619">
        <property type="entry name" value="Phospholipase A2, PLA2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00119">
        <property type="entry name" value="PA2_ASP"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="PA2_HIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 8" id="KW-0106">Calcium</keyword>
    <keyword evidence="3 6" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="8" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="5" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 8" id="KW-0964">Secreted</keyword>
    <feature type="domain" description="Phospholipase A2-like central" evidence="9">
        <location>
            <begin position="8"/>
            <end position="127"/>
        </location>
    </feature>
    <feature type="active site" evidence="4">
        <location>
            <position position="54"/>
        </location>
    </feature>
    <feature type="active site" evidence="4">
        <location>
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        </location>
    </feature>
    <feature type="binding site" evidence="5">
        <location>
            <position position="36"/>
        </location>
        <ligand>
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="5">
        <location>
            <position position="38"/>
        </location>
        <ligand>
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="5">
        <location>
            <position position="55"/>
        </location>
        <ligand>
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </ligand>
    </feature>
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        <location>
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            <end position="126"/>
        </location>
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            <end position="51"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="6">
        <location>
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        </location>
    </feature>
    <feature type="disulfide bond" evidence="6">
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        </location>
    </feature>
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            <end position="97"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
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    </evidence>
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        <source>
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    </evidence>
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        <source>
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        </source>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        <source>
            <dbReference type="RuleBase" id="RU003654"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="RuleBase" id="RU361236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="SMART" id="SM00085"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029753.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="133" mass="15325" checksum="6C1B32FE40027BA0" modified="2019-05-08" version="1">SRDIVVILLLNFKFMIRRITGKSALISYNGYGCHCGLGGRGQPLDKTDWCCHAHDCCYKELTNQSCHPFFDYYQYSIIDEDVMCWNTNNSTCAKQACDCDRTAVLCFRKEASSYNKGYRYYFNFFCKGSTPSC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INT5</accession>
    <name>A0A452INT5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">Ras-GAP domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
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        <lineage>
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            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
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                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000033567.1">
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        <property type="gene ID" value="ENSGAGG00000019812.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903479">
        <property type="term" value="P:mitotic actomyosin contractile ring assembly actin filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05131">
        <property type="entry name" value="RasGAP_IQGAP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00201">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.506.10:FF:000004">
        <property type="entry name" value="IQ motif containing GTPase activating protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.506.10">
        <property type="entry name" value="GTPase Activation - p120gap, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000048">
        <property type="entry name" value="IQ_motif_EF-hand-BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000593">
        <property type="entry name" value="RasGAP_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023152">
        <property type="entry name" value="RasGAP_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001936">
        <property type="entry name" value="RasGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001202">
        <property type="entry name" value="WW_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149">
        <property type="entry name" value="RAS GTPASE-ACTIVATING PROTEIN WITH IQ MOTIF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149:SF12">
        <property type="entry name" value="RAS GTPASE-ACTIVATING-LIKE PROTEIN IQGAP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00612">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00616">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03836">
        <property type="entry name" value="RasGAP_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00015">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00323">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00456">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF143885">
        <property type="entry name" value="RGC domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00509">
        <property type="entry name" value="RAS_GTPASE_ACTIV_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50018">
        <property type="entry name" value="RAS_GTPASE_ACTIV_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01159">
        <property type="entry name" value="WW_DOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Ras-GAP" evidence="1">
        <location>
            <begin position="622"/>
            <end position="855"/>
        </location>
    </feature>
    <evidence type="ECO:0000259" key="1">
        <source>
            <dbReference type="PROSITE" id="PS50018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029559.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1282" mass="146971" checksum="BD169FF486CFA5CC" modified="2019-05-08" version="1">LTHKTIRSHNWCNGKNPINFLYFPPELVAVDHINTAIRNCDANKTLVALMKPEAQLPVVHPFAAAVYQTELFNLQQQNAVRYLAHDELSIAVEMLSAVVLLNQALGSKDILAIKNHLRNPSVGFNNVEDENFQRYADTLLSIKSEASSQGQDYLSWNDVQNCIDMVNMQVQEENDRIVAVSYINEAIDQGNPGKTLETLLLPTVKLHDVNPTNARHYQDVLHDAKVQKCKNCICVYVKYKMSDVEMQFCCLVALGTSMINNCLEKHDSQPVLDILQSPKFGLRVVPECAETYYKNLLEAKNLKTKEDSSEGPWLKLIMKNRYDYYYNVETGESTFVPPEGFIPKTSWLTSEEIQTIVGQVTADYNREQLWFANENLILQLQALARGFLVRKSYEERKGFLQKQEPSVIKIQVLLCNIKAQSLLLGIQSWVKMWLARKAYKKRLQYFKDHVRHFSTPSPFEKRTNVVVMHVFLEITDTVVGAENPPLNVLRKFAYLLDQSDLDFQEELEVTRLREEVVTKIRSNQQLEKDLNLMDIKIGLLVKNRISLQDVVLHSKKLNKKSKNQVEEMVTGDKQGIKGLSKERRKQLEAYQHLFYLLQTNPMYLAKLIFQMPQNKSTKFMDTVIFTLYNYASNQREEYLLLKLFKTALEEEINSKVDQIQDIVTGNPTVIKMVVSFNRGARGQNSLRQLLAPVVKEIMDDKSLLINTSPVEVYKAWVNQLETQTGEASKLPYDVTTEQALEHPEVVNRLESSIQSLRTVTDKVLVSIFSSLSMMPYGMRYIAKVLKNSLHEKFPDATEDELLKIVGNLLYYRYMNPAIVAPDGFDIIDMTAGGQIHSDQRRNLGCVAKVLQHAASNKLFEGESAHLSSMNTYLSQTYQKFRNFFQIACDVPEPEEKFNIDEYSDMVTLSKPVIYISIEEIINTHSLLLEHQDAIAPEQNDLLNELLKGLGAVPDVESFLGEGAVDPNDPNKENTLSQLAKTEISLSLTSKYELQEGEDQDLKGLMIKTKRLVVDVIRTQPGDTLLEILEIPASAQQESEHLKVVEKRAILDSKTPEKMKHSQSMLEDGQLPIEQKKRKIQRNLRTLEQAGLVSSANKYQEIINEIAKDIRNQRRYRHHRKAELVKLQQTLSALQSKTAFYEDQVNYYNTYIKTCLDNLTRKNSRRSIKLDGKVEEKGSKKLKHTSLKYTAARLHEKGVILEIEDLQTNQFKNVMFDITPGEEMGDFEIKAKFLGVQMEKVELHFQDLLQMQYEGVAVMKMFDKAKVNVNLLIFLLNKKFYGK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INT7</accession>
    <name>A0A452INT7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Annexin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="domain">
        <text evidence="6">A pair of annexin repeats may form one binding site for calcium and phospholipid.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1 6">Belongs to the annexin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452INT7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033788.1">
        <property type="protein sequence ID" value="ENSGAGP00000029758.1"/>
        <property type="gene ID" value="ENSGAGG00000021484.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034704">
        <property type="term" value="C:calcium channel complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012506">
        <property type="term" value="C:vesicle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005262">
        <property type="term" value="F:calcium channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005544">
        <property type="term" value="F:calcium-dependent phospholipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001786">
        <property type="term" value="F:phosphatidylserine binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048513">
        <property type="term" value="P:animal organ development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055074">
        <property type="term" value="P:calcium ion homeostasis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032506">
        <property type="term" value="P:cytokinetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006909">
        <property type="term" value="P:phagocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009888">
        <property type="term" value="P:tissue development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.220.10:FF:000001">
        <property type="entry name" value="Annexin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.220.10:FF:000002">
        <property type="entry name" value="Annexin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.220.10:FF:000003">
        <property type="entry name" value="Annexin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.220.10:FF:000004">
        <property type="entry name" value="Annexin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.220.10">
        <property type="entry name" value="Annexin"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001464">
        <property type="entry name" value="Annexin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018502">
        <property type="entry name" value="Annexin_repeat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018252">
        <property type="entry name" value="Annexin_repeat_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037104">
        <property type="entry name" value="Annexin_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008157">
        <property type="entry name" value="ANX11"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10502">
        <property type="entry name" value="ANNEXIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10502:SF29">
        <property type="entry name" value="ANNEXIN A11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00191">
        <property type="entry name" value="Annexin"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00196">
        <property type="entry name" value="ANNEXIN"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01810">
        <property type="entry name" value="ANNEXINXI"/>
    </dbReference>
    <dbReference type="SMART" id="SM00335">
        <property type="entry name" value="ANX"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47874">
        <property type="entry name" value="Annexin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00223">
        <property type="entry name" value="ANNEXIN_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51897">
        <property type="entry name" value="ANNEXIN_2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 6" id="KW-0041">Annexin</keyword>
    <keyword evidence="3 6" id="KW-0106">Calcium</keyword>
    <keyword evidence="5 6" id="KW-0111">Calcium/phospholipid-binding</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 6" id="KW-0677">Repeat</keyword>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="74"/>
            <end position="147"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="7">
        <location>
            <begin position="74"/>
            <end position="85"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="7">
        <location>
            <begin position="102"/>
            <end position="112"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="7">
        <location>
            <begin position="138"/>
            <end position="147"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007831"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023216"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023302"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="RuleBase" id="RU003540"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029758.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="502" mass="54313" checksum="CCAF2EFD444571C9" modified="2019-05-08" version="1">MSYPGYPPAGGYPPAAPGSNPWGGSGYPPINPPPIGLENVANYANQFNPDYMSGMASNLSGTFGGANVPQSIYPPAPGGYPPVPPGGFGQPPSGQQASYGTYPPPGGNPPAGMPAYPGYPGSHMPPPGQQPMTYPGQQPMPLPGQQPVPLPGQQPMPSYPSGPAVNPSMPPYSGSVMPTITPGTSGNRGTITDAPGFDPLRDAEVLRKAMKGFGTDEQAIIDCLGSRSNKQRQKIILSFKTAYGKDLIKDLKSELSGNFEKTILAMLKTPVMYDVCEIKEAIKGAGTDEACLIEIMSSRSNEHIREISRTYKTEFKKTLEEAIRSDTSGHFQRLLISLSQGNRDESTNVDMSLVQRDVQELYAAGENRLGTDESKFNAILCTRSRAHLRAVFSEYQRMCNRDIEKSISREMSGDLESGMVAVVKCMKNTPAFFAERLHKAMKGAGTKDRTLIRIMVSRSEVDLLDIRTEYKRMYGKSLYADITGDTSGDYRKILLKLCGGND</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INT9</accession>
    <name>A0A452INT9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="5">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452INT9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033572.1">
        <property type="protein sequence ID" value="ENSGAGP00000029564.1"/>
        <property type="gene ID" value="ENSGAGG00000021354.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031410">
        <property type="term" value="C:cytoplasmic vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019905">
        <property type="term" value="F:syntaxin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008286">
        <property type="term" value="P:insulin receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061178">
        <property type="term" value="P:regulation of insulin secretion involved in cellular response to glucose stimulus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd06698">
        <property type="entry name" value="PDZ1_hSTXBP4-PDZ2_GgSTXBP4-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00201">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.70.10:FF:000034">
        <property type="entry name" value="syntaxin-binding protein 4 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.20.70.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.42.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001478">
        <property type="entry name" value="PDZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051342">
        <property type="entry name" value="PDZ_scaffold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036034">
        <property type="entry name" value="PDZ_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001202">
        <property type="entry name" value="WW_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036020">
        <property type="entry name" value="WW_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19964">
        <property type="entry name" value="MULTIPLE PDZ DOMAIN PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19964:SF16">
        <property type="entry name" value="SYNTAXIN-BINDING PROTEIN 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00595">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00397">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00228">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00456">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50156">
        <property type="entry name" value="PDZ domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51045">
        <property type="entry name" value="WW domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50106">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01159">
        <property type="entry name" value="WW_DOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50020">
        <property type="entry name" value="WW_DOMAIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="2">
        <location>
            <begin position="1"/>
            <end position="16"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019207238" evidence="2">
        <location>
            <begin position="17"/>
            <end position="486"/>
        </location>
    </feature>
    <feature type="domain" description="PDZ" evidence="4">
        <location>
            <begin position="14"/>
            <end position="88"/>
        </location>
    </feature>
    <feature type="domain" description="WW" evidence="3">
        <location>
            <begin position="429"/>
            <end position="462"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="269"/>
            <end position="359"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS50020"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50106"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029564.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="486" mass="54348" checksum="28E643A068AB1092" modified="2019-05-08" version="1">MLIFFSFFSLSDSAFQIITVSKGTGLGLNIVGGINRNEGPLVYIQEVIPGGDCHKDGRLKPGDQLVSINKESMIGVSYEEAKSIINRTKLRSESAWEIAFIRQRVFPSYFENPQSPGSLLTSSAGCGQQQTLLAFNLLSPPNENPASKVPTPYTVTVASSWTEDYGPQERKISLKPTVRLKAEKLEMALNYLGIQPTDEQQRVLRHQLQKDAKGTVSFGDFVQVARNLFCLQLDEPGGGQRPITFGASEIANLLDSQLVSCDSLEMDDVERLKRERNEALKEISKLKEELCESVGLRKQLTEELQTVKQEAKAAVEEARALRSRIHLAEAAQRQARGMEMDYEEIIRLLEAEITELKAQLTGRSGQNKDSVQDLRKRITVLDCQLRKSESARKTSEVATGKPLKWPLQNSDNSIPLSNLRVILFLYFTLGLPYGWEEAYTADGIKYFINHVTQTTSWIHPVTSALNLSCSGENEEDHIRELPDPKS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INU0</accession>
    <name>A0A452INU0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Calponin-homology (CH) domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the hook family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452INU0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033652.1">
        <property type="protein sequence ID" value="ENSGAGP00000029637.1"/>
        <property type="gene ID" value="ENSGAGG00000021396.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005874">
        <property type="term" value="C:microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051959">
        <property type="term" value="F:dynein light intermediate chain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008017">
        <property type="term" value="F:microtubule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031122">
        <property type="term" value="P:cytoplasmic microtubule organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030705">
        <property type="term" value="P:cytoskeleton-dependent intracellular transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.418.10:FF:000197">
        <property type="entry name" value="Protein Hook homolog 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.418.10">
        <property type="entry name" value="Calponin-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001715">
        <property type="entry name" value="CH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036872">
        <property type="entry name" value="CH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008636">
        <property type="entry name" value="Hook_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043936">
        <property type="entry name" value="HOOK_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18947">
        <property type="entry name" value="HOOK PROTEINS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18947:SF38">
        <property type="entry name" value="PROTEIN HOOK HOMOLOG 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05622">
        <property type="entry name" value="HOOK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19047">
        <property type="entry name" value="HOOK_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF116907">
        <property type="entry name" value="Hook domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50021">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="4" id="KW-0493">Microtubule</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Calponin-homology (CH)" evidence="9">
        <location>
            <begin position="1"/>
            <end position="83"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="639"/>
            <end position="679"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="128"/>
            <end position="387"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="446"/>
            <end position="607"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="644"/>
            <end position="663"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029637.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="679" mass="78865" checksum="5B070E406F64839E" modified="2019-05-08" version="1">MTGDRDPAYFDENWLNRIKTEVGDNWRLKISNLKKILKGILDYNHEILGQQINDFTLPDVNLIGEHSDVAELGRMLQLILGCAVNCEQKQEYIQTIMMMEESVQHVVMTAIQELMSKESPVSVGNDAYVDLDRQLKKTTEELNEALATKEEIAQRCHELDMQVAALQEEKSSLLAENQILMERLNQSDSIEDPNSPAGRRHLQLQTQLEQLQEETFRLEAAKDDYRIRCEELEKEIAELRQQNEELTTFAEEAQSLKDEMDVLRHSSDKVAKLESQVESYKKKLEDLGDLRRQVKLLEEKNTMYMQNTVSLEEELRKAHAARSQLETYKRQVVELQNRLSEESKKADKLEFEYKLLKEKVDGLQKEKDRLRTERDSLKETIEELRCVQAQEGQLTTSGLMPLGSQESSDSLAAEIVTPEIKEKLIRLQHENKMLKLNQEGSDNEKIALLQSLLDDANLRKNELETENRLVNQRLLEVQSQVEELQKSLQEQGSKAEDSVLLKKKLEEHLEKLHEANNEIQRKRAIIEDLEPRYNNSSLKIEELQEALRKKEEEMKQMEERYKKYLEKAKSVIRTLDPKQNQGTAPEIQALKNQLQERDRMFHSLEKEYEKTKTQREMEEKFIVSAWYNMGMTLHKKAAEDRLASTGSGQSFLARQRQATNTRRSYPGHVQPATASDVIA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INU1</accession>
    <name>A0A452INU1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">General transcription factor 3C polypeptide 5</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="12">TF3C-epsilon</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="10">Transcription factor IIIC 63 kDa subunit</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="11">Transcription factor IIIC subunit epsilon</fullName>
        </alternativeName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="6">Involved in RNA polymerase III-mediated transcription. Integral, tightly associated component of the DNA-binding TFIIIC2 subcomplex that directly binds tRNA and virus-associated RNA promoters.</text>
    </comment>
    <comment type="subunit">
        <text evidence="8">Part of the TFIIIC subcomplex TFIIIC2, consisting of six subunits, GTF3C1, GTF3C2, GTF3C3, GTF3C4, GTF3C5 and GTF3C6. Interacts with BRF1, GTF3C6 and TBP.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="7">Belongs to the TFIIIC subunit 5 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452INU1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033525.1">
        <property type="protein sequence ID" value="ENSGAGP00000029525.1"/>
        <property type="gene ID" value="ENSGAGG00000021325.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000127">
        <property type="term" value="C:transcription factor TFIIIC complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001002">
        <property type="term" value="F:RNA polymerase III type 1 promoter sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001003">
        <property type="term" value="F:RNA polymerase III type 2 promoter sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006384">
        <property type="term" value="P:transcription initiation at RNA polymerase III promoter"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.160:FF:000001">
        <property type="entry name" value="General transcription factor IIIC subunit 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.160">
        <property type="entry name" value="TFIIIC, subcomplex tauA, subunit Sfc1, barrel domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019136">
        <property type="entry name" value="TF_IIIC_su-5_HTH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040454">
        <property type="entry name" value="TF_IIIC_Tfc1/Sfc1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041499">
        <property type="entry name" value="Tfc1/Sfc1_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042536">
        <property type="entry name" value="TFIIIC_tauA_Sfc1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13230:SF5">
        <property type="entry name" value="GENERAL TRANSCRIPTION FACTOR 3C POLYPEPTIDE 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13230">
        <property type="entry name" value="GENERAL TRANSCRIPTION FACTOR IIIC, POLYPEPTIDE 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09734">
        <property type="entry name" value="Tau95"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17682">
        <property type="entry name" value="Tau95_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0007">Acetylation</keyword>
    <keyword evidence="3" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="5" id="KW-0539">Nucleus</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0804">Transcription</keyword>
    <feature type="domain" description="Transcription factor IIIC subunit Tfc1/Sfc1 triple barrel" evidence="15">
        <location>
            <begin position="23"/>
            <end position="129"/>
        </location>
    </feature>
    <feature type="domain" description="Transcription factor IIIC subunit 5 HTH" evidence="14">
        <location>
            <begin position="166"/>
            <end position="320"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="472"/>
            <end position="514"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="13">
        <location>
            <begin position="482"/>
            <end position="495"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="13">
        <location>
            <begin position="502"/>
            <end position="514"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00057927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00061645"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00064690"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00069548"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00077529"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00078452"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00081022"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="Pfam" id="PF09734"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="Pfam" id="PF17682"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029525.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="514" mass="59359" checksum="235988FF5194613B" modified="2019-05-08" version="1">MAAGGGGEHVSAVLELPRGPRLVCVEYPGLVRDVGKMLQTLGGEEGVSRIYADPGKRLELYFRPKDPYCHPVCANRFPTSSMLFKVKRKIRRKQLEAEGETQEEVKFEMEVLGVVSTVYKFQGMSDFQFLAMHSGPENKQISMYDKILMLKPEKEEFFNRELPLYIPPPIFSRLDTPIDYFYRPDVQHREGYNNPQVSCENLIGLSRARRPHNAIFVNFDDDDVPTKPLEPAVQTWKKACTNPVDKKVEEELRKLFEIRPVWSRNAVKANISVHPEKLKLLLPYLAYYMLTGPWRSLWVRYGYDPRKHPEAKIYQVLDFRIRCGMKYGYAPSDIPVKAKRSTYNYSLPITIKKPVSHTVNVQDLKHGLDPASTSSAKKPASSKYKLKDSIYIFQEGDLPPYRQMFYQLCDLNVDSLQKIIHRNDGMETECTERDGWCLQKTSDDLRDTMSLMIKHIIRSTRPALFSSTLNAEDGKERLTYDSGEEEEEEEDEEEDFKPSDGSENEMETEILDYV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INU2</accession>
    <name>A0A452INU2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Phospholipase A2</fullName>
            <ecNumber evidence="8">3.1.1.4</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="8">
            <text>a 1,2-diacyl-sn-glycero-3-phosphocholine + H2O = a 1-acyl-sn-glycero-3-phosphocholine + a fatty acid + H(+)</text>
            <dbReference type="Rhea" id="RHEA:15801"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:28868"/>
            <dbReference type="ChEBI" id="CHEBI:57643"/>
            <dbReference type="ChEBI" id="CHEBI:58168"/>
            <dbReference type="EC" id="3.1.1.4"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="5">
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </cofactor>
        <text evidence="5">Binds 1 Ca(2+) ion per subunit.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 8">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="7">Belongs to the phospholipase A2 family.</text>
    </comment>
    <dbReference type="EC" id="3.1.1.4" evidence="8"/>
    <dbReference type="AlphaFoldDB" id="A0A452INU2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033796.1">
        <property type="protein sequence ID" value="ENSGAGP00000029763.1"/>
        <property type="gene ID" value="ENSGAGG00000021486.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0047498">
        <property type="term" value="F:calcium-dependent phospholipase A2 activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005543">
        <property type="term" value="F:phospholipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050482">
        <property type="term" value="P:arachidonate secretion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016042">
        <property type="term" value="P:lipid catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042130">
        <property type="term" value="P:negative regulation of T cell proliferation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006644">
        <property type="term" value="P:phospholipid metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00125">
        <property type="entry name" value="PLA2c"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.90.10:FF:000001">
        <property type="entry name" value="Basic phospholipase A2 homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.90.10">
        <property type="entry name" value="Phospholipase A2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001211">
        <property type="entry name" value="PLA2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016090">
        <property type="entry name" value="PLA2-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033112">
        <property type="entry name" value="PLA2_Asp_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033113">
        <property type="entry name" value="PLA2_histidine"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036444">
        <property type="entry name" value="PLipase_A2_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11716">
        <property type="entry name" value="PHOSPHOLIPASE A2 FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11716:SF9">
        <property type="entry name" value="PHOSPHOLIPASE A2, MEMBRANE ASSOCIATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00068">
        <property type="entry name" value="Phospholip_A2_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00389">
        <property type="entry name" value="PHPHLIPASEA2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00085">
        <property type="entry name" value="PA2c"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48619">
        <property type="entry name" value="Phospholipase A2, PLA2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00119">
        <property type="entry name" value="PA2_ASP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00118">
        <property type="entry name" value="PA2_HIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 8" id="KW-0106">Calcium</keyword>
    <keyword evidence="3 6" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="8" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="5" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 8" id="KW-0964">Secreted</keyword>
    <feature type="domain" description="Phospholipase A2-like central" evidence="9">
        <location>
            <begin position="6"/>
            <end position="123"/>
        </location>
    </feature>
    <feature type="active site" evidence="4">
        <location>
            <position position="52"/>
        </location>
    </feature>
    <feature type="active site" evidence="4">
        <location>
            <position position="96"/>
        </location>
    </feature>
    <feature type="binding site" evidence="5">
        <location>
            <position position="32"/>
        </location>
        <ligand>
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="5">
        <location>
            <position position="34"/>
        </location>
        <ligand>
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </ligand>
    </feature>
    <feature type="disulfide bond" evidence="6">
        <location>
            <begin position="31"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="6">
        <location>
            <begin position="33"/>
            <end position="49"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="6">
        <location>
            <begin position="48"/>
            <end position="102"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="6">
        <location>
            <begin position="54"/>
            <end position="129"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="6">
        <location>
            <begin position="55"/>
            <end position="95"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="6">
        <location>
            <begin position="64"/>
            <end position="88"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="6">
        <location>
            <begin position="82"/>
            <end position="93"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PIRSR" id="PIRSR601211-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PIRSR" id="PIRSR601211-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PIRSR" id="PIRSR601211-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU003654"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="RuleBase" id="RU361236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="SMART" id="SM00085"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029763.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="129" mass="14696" checksum="8EAF54C452C89852" modified="2019-05-08" version="1">SFLVLFLLEFRKMINQATGKSAILSYYGYGCYCGSDGKGEPKDATDWCCRAHHCCYKRLKASGCYGNTERYSYTYKDGDILCALGSWCEEQVCDCDKSTAFCLERNLKTFNSLYVRYRDSKCTGFTPNC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INU3</accession>
    <name>A0A452INU3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="22">Cell division cycle 14B</fullName>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>O-phospho-L-seryl-[protein] + H2O = L-seryl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:20629"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="EC" id="3.1.3.16"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="16 23">
            <text>O-phospho-L-threonyl-[protein] + H2O = L-threonyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:47004"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="EC" id="3.1.3.16"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="14">Cell projection</location>
            <location evidence="14">Kinocilium</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
            <location evidence="2">Microtubule organizing center</location>
            <location evidence="2">Centrosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Cytoskeleton</location>
            <location evidence="3">Spindle pole</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the protein-tyrosine phosphatase family. Non-receptor class CDC14 subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452INU3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033578.1">
        <property type="protein sequence ID" value="ENSGAGP00000029570.1"/>
        <property type="gene ID" value="ENSGAGG00000021351.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060091">
        <property type="term" value="C:kinocilium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000922">
        <property type="term" value="C:spindle pole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004722">
        <property type="term" value="F:protein serine/threonine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007605">
        <property type="term" value="P:sensory perception of sound"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd14499">
        <property type="entry name" value="CDC14_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd17657">
        <property type="entry name" value="CDC14_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000032">
        <property type="entry name" value="dual specificity protein phosphatase CDC14A isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000006">
        <property type="entry name" value="Dual specificity protein phosphatase CDC14B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044506">
        <property type="entry name" value="CDC14_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029260">
        <property type="entry name" value="DSPn"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000340">
        <property type="entry name" value="Dual-sp_phosphatase_cat-dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050561">
        <property type="entry name" value="PTP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016130">
        <property type="entry name" value="Tyr_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020422">
        <property type="entry name" value="TYR_PHOSPHATASE_DUAL_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23339">
        <property type="entry name" value="TYROSINE SPECIFIC PROTEIN PHOSPHATASE AND DUAL SPECIFICITY PROTEIN PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00782">
        <property type="entry name" value="DSPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14671">
        <property type="entry name" value="DSPn"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00195">
        <property type="entry name" value="DSPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00383">
        <property type="entry name" value="TYR_PHOSPHATASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50054">
        <property type="entry name" value="TYR_PHOSPHATASE_DUAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="13" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="7" id="KW-0132">Cell division</keyword>
    <keyword evidence="12" id="KW-0966">Cell projection</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="10" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="17" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="9" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="17" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="17" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="403"/>
            <end position="429"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="18">
        <location>
            <begin position="178"/>
            <end position="337"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="19">
        <location>
            <begin position="260"/>
            <end position="322"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004647"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00007315"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00037822"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00047761"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00048336"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029570.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="22">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="446" mass="50981" checksum="6F887EF289F41415" modified="2019-05-08" version="1">MSVRKESRPSVEFVKGRLYFAILCQKPKSGVANTHYFCIDDELVYENFYADFGPLNLAMVYRYCCKLNKKLKSFTLIRKKIIHYTGFDQKKQANAAFLIGSYAIIYLKESPEDVYRLLLAGNASYLPFRDASFGTCSFHLTLLDCFHAINKALRYGFLDFSAFDVNEYEHYEKAENGDFNWIIPNKFIAFSGPHSRSKTENGYPHHAPEAYFPYFRRHKVTTIIRLNRKMYDAKRFADAGFEHYDLFFADGSIPSDTIVKTFLNICENAEGVIAVHCKAGLGRTGTLIACYIMKHYRMTAAETIAWIRINRPGSVIGPQQHFLVEKQTELWREGDVFRAKLKENRKIAVTKILTGVDDISINDAKNRRANRKDTELYSDEDDANCVTQGDKLRALKSKRQAKAAATVPLTWLLAMLVSTLCSIVIWWIVCGSLLPSLLFSLDGLRT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INU4</accession>
    <name>A0A452INU4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">Ras-GAP domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000033555.1">
        <property type="protein sequence ID" value="ENSGAGP00000029548.1"/>
        <property type="gene ID" value="ENSGAGG00000019812.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005938">
        <property type="term" value="C:cell cortex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903479">
        <property type="term" value="P:mitotic actomyosin contractile ring assembly actin filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05131">
        <property type="entry name" value="RasGAP_IQGAP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00201">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.506.10:FF:000004">
        <property type="entry name" value="IQ motif containing GTPase activating protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.506.10">
        <property type="entry name" value="GTPase Activation - p120gap, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000048">
        <property type="entry name" value="IQ_motif_EF-hand-BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000593">
        <property type="entry name" value="RasGAP_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023152">
        <property type="entry name" value="RasGAP_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001936">
        <property type="entry name" value="RasGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001202">
        <property type="entry name" value="WW_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149">
        <property type="entry name" value="RAS GTPASE-ACTIVATING PROTEIN WITH IQ MOTIF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149:SF12">
        <property type="entry name" value="RAS GTPASE-ACTIVATING-LIKE PROTEIN IQGAP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00612">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00616">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03836">
        <property type="entry name" value="RasGAP_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00015">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00323">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00456">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF143885">
        <property type="entry name" value="RGC domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00509">
        <property type="entry name" value="RAS_GTPASE_ACTIV_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50018">
        <property type="entry name" value="RAS_GTPASE_ACTIV_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01159">
        <property type="entry name" value="WW_DOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Ras-GAP" evidence="1">
        <location>
            <begin position="622"/>
            <end position="855"/>
        </location>
    </feature>
    <evidence type="ECO:0000259" key="1">
        <source>
            <dbReference type="PROSITE" id="PS50018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029548.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1282" mass="146913" checksum="0FDCEFD2D47C9A2D" modified="2019-05-08" version="1">LTHKTIRSHNWCNGKNPINFLYFPPELVAVDHINTAIRNCDANKTLVALMKPEAQLPVVHPFAAAVYQTELFNLQQQNAVRYLAHDELSIAVEMLSAVVLLNQALGSKDILAIKNHLRNPSVGFNNVEDENFQRYADTLLSIKSEASSQGQDYLSWNDVQNCIDMVNMQVQEENDRIVAVSYINEAIDQGNPGKTLETLLLPTVKLHDVNPTNARHYQDVLHDAKVQKCKVEPQLNCICVYVKYKMSDVEMQFCCLVALGTSMINNCLEKHDSQPVLDILQSPKFGLRVVPECAETYYKNLLEAKNLKTKEDSSEGPWLKLIMKNRYDYYYNVETGESTFVPPEGFIPKTSWLTSEEIQTIVGQVTADYNREQLWFANENLILQLQALARGFLVRKSYEERKGFLQKQEPSVIKIQVLLCNIKYSSLSLFIEHDLLFEVSCSMLSKVFSLWSFFQNEEIVKIQAFLRANKARDDYRTLIGAENPPLNVLRKFAYLLDQSDLDFQEELEVTRLREEVVTKIRSNQQLEKDLNLMDIKIGLLVKNRISLQDVVLHSKKLNKKSKNQVEEMVTGDKQGIKGLSKERRKQLEAYQHLFYLLQTNPMYLAKLIFQMPQNKSTKFMDTVIFTLYNYASNQREEYLLLKLFKTALEEEINSKVDQIQDIVTGNPTVIKMVVSFNRGARGQNSLRQLLAPVVKEIMDDKSLLINTSPVEVYKAWVNQLETQTGEASKLPYDVTTEQALEHPEVVNRLESSIQSLRTVTDKVLVSIFSSLSMMPYGMRYIAKVLKNSLHEKFPDATEDELLKIVGNLLYYRYMNPAIVAPDGFDIIDMTAGGQIHSDQRRNLGCVAKVLQHAASNKLFEGESAHLSSMNTYLSQTYQKFRNFFQIACDVPEPEEKFNIDEYSDMVTLSKPVIYISIEEIINTHSLLLEHQDAIAPEQNDLLNELLKGLGAVPDVESFLGEGAVDPNDPNKENTLSQLAKTEISLSLTSKYELQEGEDQDLKGLMIKTKRLVVDVIRTQPGDTLLEILEIPASAQQESEHLKVVEKRAILDSKTPEKMKHSQSMLEDGQLPIEQKKRKIQRNLRTLEQAGLVSSANKYQEIINEIAKDIRNQRRYRHHRKAELVKLQQTLSALQSKTAFYEDQVNYYNTYIKTCLDNLTRKNSRRSIKLDGKVEEKGSKKLKHTSLKYTAARLHEKGVILEIEDLQTNQFKNVMFDITPGEEMGDFEIKAKFLGVQMEKVELHFQDLLQMQYEGVAVMKMFDKAKVNVNLLIFLLNKKFYGK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J000</accession>
    <name>A0A452J000_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Rho GTPase-activating protein 17</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="14">Rho-type GTPase-activating protein 17</fullName>
        </alternativeName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="11">Rho GTPase-activating protein involved in the maintenance of tight junction by regulating the activity of CDC42, thereby playing a central role in apical polarity of epithelial cells. Specifically acts as a GTPase activator for the CDC42 GTPase by converting it to an inactive GDP-bound state. The complex formed with AMOT acts by regulating the uptake of polarity proteins at tight junctions, possibly by deciding whether tight junction transmembrane proteins are recycled back to the plasma membrane or sent elsewhere. Participates in the Ca(2+)-dependent regulation of exocytosis, possibly by catalyzing GTPase activity of Rho family proteins and by inducing the reorganization of the cortical actin filaments. Acts as a GTPase activator in vitro for RAC1.</text>
    </comment>
    <comment type="subunit">
        <text evidence="12">Component of a complex whose core is composed of ARHGAP17, AMOT, PALS1, PATJ and PARD3/PAR3. Interacts with NHERF1, FNBP1, TRIP10, CAPZA (CAPZA1, CAPZA2 or CAPZA3), CAPZB, CD2AP and SH3KBP1/CIN85.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell junction</location>
            <location evidence="2">Tight junction</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Peripheral membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J000"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038137.1">
        <property type="protein sequence ID" value="ENSGAGP00000033671.1"/>
        <property type="gene ID" value="ENSGAGG00000023966.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005923">
        <property type="term" value="C:bicellular tight junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017124">
        <property type="term" value="F:SH3 domain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032956">
        <property type="term" value="P:regulation of actin cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035020">
        <property type="term" value="P:regulation of Rac protein signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd04386">
        <property type="entry name" value="RhoGAP_nadrin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.555.10:FF:000001">
        <property type="entry name" value="Rho GTPase activating protein 44"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1270.60:FF:000019">
        <property type="entry name" value="rho GTPase-activating protein 17 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.60">
        <property type="entry name" value="Arfaptin homology (AH) domain/BAR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.555.10">
        <property type="entry name" value="Rho GTPase activation protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027267">
        <property type="entry name" value="AH/BAR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004148">
        <property type="entry name" value="BAR_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047165">
        <property type="entry name" value="RHG17/44/SH3BP1-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000198">
        <property type="entry name" value="RhoGAP_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14130">
        <property type="entry name" value="3BP-1 RELATED RHOGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14130:SF3">
        <property type="entry name" value="RHO GTPASE-ACTIVATING PROTEIN 17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03114">
        <property type="entry name" value="BAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00620">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00721">
        <property type="entry name" value="BAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00324">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103657">
        <property type="entry name" value="BAR/IMD domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51021">
        <property type="entry name" value="BAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50238">
        <property type="entry name" value="RHOGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8" id="KW-0965">Cell junction</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="5" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="10" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0729">SH3-binding</keyword>
    <keyword evidence="4" id="KW-0796">Tight junction</keyword>
    <feature type="domain" description="BAR" evidence="17">
        <location>
            <begin position="1"/>
            <end position="230"/>
        </location>
    </feature>
    <feature type="domain" description="Rho-GAP" evidence="16">
        <location>
            <begin position="236"/>
            <end position="426"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="496"/>
            <end position="517"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="552"/>
            <end position="867"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="15">
        <location>
            <begin position="552"/>
            <end position="564"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="15">
        <location>
            <begin position="585"/>
            <end position="605"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="15">
        <location>
            <begin position="624"/>
            <end position="634"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="15">
        <location>
            <begin position="662"/>
            <end position="689"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="15">
        <location>
            <begin position="695"/>
            <end position="710"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="15">
        <location>
            <begin position="735"/>
            <end position="747"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="15">
        <location>
            <begin position="760"/>
            <end position="777"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="15">
        <location>
            <begin position="792"/>
            <end position="802"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="15">
        <location>
            <begin position="804"/>
            <end position="831"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004435"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022427"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023036"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00055904"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00065623"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00070237"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00083392"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50238"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS51021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033671.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="867" mass="94051" checksum="F77F4728367E6E09" modified="2019-05-08" version="1">MAEKTEVLSEDLLQIERRLDTVRSVCHISQKRLIACFQGQYGTDADKRHKKLPLTAFAQNMQEGSVQLSDDTLLGKMLDTCGDAENKLAVELSHHEVQIEKEILDPLSLLTEAEIPNIQKQRKQLAKLVLDWDSAKARYSQACKTSGTNFQMHPSKIDSLKEEMDEAGNKVEQCKDQLAADMYNCVSKEGEYARYFVMLLEAQADYHRKSLAVLEKALPEIQAHQDKWTEKPAFGTPLEEHLKRSGREIALPIEACVMMLLETGMKEEGLFRIAAGASKLKKLKAALDCSTSQLDEFYSDPHAVAGALKSYLRELPEPLMTYSLYEEWTQAANIQDQDKKLQELWKICKRLPKANLANFRYLIKFLATLAQNSDINKMTPSNIAIVLGPNLLWAKTEGSLAEMAAATSVHVVAIIEPIIQHADWFFPEDVDFNVSGAFVALPATNSNHSSQLGSEYESGTLERKRPVSMAVMEGDLLKKEGFGVKVVDFQVNPRRCGTINRKHTSPAFQPPLPPTEAGMLAQPVAEQHSQGSVAETSPVSATFAPFAGTAEHLQSQGNEDVSTSKPRDTAASTTPPPVRNGGHLSTVQNQPQSSSSSNQLCVSQPPNAAGPSPHAVRRAVKKPAPAPPKPPPGQPGNHGSTAAAQLPSVSVKPPTRSPSPPDQLTNPGAVQTSTSSQVSAPRRYSSSLSPIQAPNHPPPQPPTQATPPLHPKLNSQVSSPPAVPSGEYGPEQPSCTPPQTPTPPDTPPLGKHSASSALFPLQSSTQETSQSHSPPQTGTLPRPKPVPKPRNRPNVPPPPHPPVSQSSGDSVLTNPTQTASKIITDSNSNVQEPLKRLSSELPTESASKELHNHLVLNVDNDTESTAL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J001</accession>
    <name>A0A452J001_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">ATP-dependent RNA helicase</fullName>
            <ecNumber evidence="9">3.6.4.13</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">RNA helicase.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="7">
            <text>ATP + H2O = ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:13065"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="3.6.4.13"/>
        </reaction>
    </comment>
    <comment type="domain">
        <text evidence="9">The Q motif is unique to and characteristic of the DEAD box family of RNA helicases and controls ATP binding and hydrolysis.</text>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the DEAD box helicase family. DDX10/DBP4 subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.6.4.13" evidence="9"/>
    <dbReference type="AlphaFoldDB" id="A0A452J001"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038151.1">
        <property type="protein sequence ID" value="ENSGAGP00000033685.1"/>
        <property type="gene ID" value="ENSGAGG00000023967.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="RHEA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003724">
        <property type="term" value="F:RNA helicase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="CDD" id="cd17941">
        <property type="entry name" value="DEADc_DDX10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18787">
        <property type="entry name" value="SF2_C_DEAD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:001089">
        <property type="entry name" value="RNA helicase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011545">
        <property type="entry name" value="DEAD/DEAH_box_helicase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014001">
        <property type="entry name" value="Helicase_ATP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001650">
        <property type="entry name" value="Helicase_C-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000629">
        <property type="entry name" value="RNA-helicase_DEAD-box_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025313">
        <property type="entry name" value="SPB4-like_CTE"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24031">
        <property type="entry name" value="RNA HELICASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13959">
        <property type="entry name" value="CTE_SPB4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00270">
        <property type="entry name" value="DEAD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00271">
        <property type="entry name" value="Helicase_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00487">
        <property type="entry name" value="DEXDc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01178">
        <property type="entry name" value="DUF4217"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00490">
        <property type="entry name" value="HELICc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00039">
        <property type="entry name" value="DEAD_ATP_HELICASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51192">
        <property type="entry name" value="HELICASE_ATP_BIND_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51194">
        <property type="entry name" value="HELICASE_CTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 8" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="3 8" id="KW-0347">Helicase</keyword>
    <keyword evidence="2 8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="1 8" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 9" id="KW-0694">RNA-binding</keyword>
    <feature type="domain" description="Helicase ATP-binding" evidence="11">
        <location>
            <begin position="10"/>
            <end position="184"/>
        </location>
    </feature>
    <feature type="domain" description="Helicase C-terminal" evidence="12">
        <location>
            <begin position="210"/>
            <end position="359"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="460"/>
            <end position="515"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="604"/>
            <end position="630"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="648"/>
            <end position="749"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="463"/>
            <end position="492"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="10">
        <location>
            <begin position="500"/>
            <end position="513"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="654"/>
            <end position="673"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="10">
        <location>
            <begin position="702"/>
            <end position="715"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022806"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00038084"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00047984"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="RuleBase" id="RU000492"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="RuleBase" id="RU365068"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS51192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS51194"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033685.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="774" mass="88343" checksum="D40B64327695FAB3" modified="2019-05-08" version="1">MVTEIQRQTIGLALQGKDVLGAAKTGSGKTLAFIVPALERLYRQQWTSIDGLGVLIISPTRELAYQTFEVLRKVGKNHDFSAGLIIGGKDLKKEVTKIHYVNMLICTPGRLLQHMDETSYFHASDLQMLILDEADRILDMGFTDTMNAIIENLPKKRQTLLFSATQTKSVKDLARLSLKDPEYVWVHEKAKFSTPATLDQNYVVCGLQQKINMLYSFLRSHLNKKSIVFYASCKEVQYLFRVFCRLRPGIPILALHGKQPQMKRMEVYNNFVRKKSAVLFATDVAARGLDFPAVNWVIQFDCPEDANTYIHRVGRTARYKEGGEALLVLLPSEEGGMVHQLSQRKVPVNKIKINPEKLVDIQKKMQSFLAQDKELKERAQMCFVSYLRSVYLMKNKEVFDVFKLPLAEYAHSLGLAVEPRVRFLQKAQKQLSVNESLEGKSHPEGTGQCEDACSLVSNEEVEERGTHFKEKISTVQRMKGEGRKEEPEEKELPSQVIDKEGDESEELSEEEEVPSINKIPKAESINFLADDDDDFRDLDLLTVKRQNVFGLEAKDLAVLNTSNSELKKKATKTKEAKKILKRKFKVNTKIVFTEDGELVQQWPPVQKSCVDSEEKEDESGGINLDKAKERLQEEDKFDKEVYRKKIKEKHREKRLREKAERRKASRTSEKEGVEAFLDYSGNEEEFDPSTLPDPDKYRGSDQEADLDTEDRDGELEEKNGTRKRSKSSNNAAEESPLRRKKAKLSDEHSFVPLDTGLSLAEDEELVLHLLNSQK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J002</accession>
    <name>A0A452J002_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">ATP-dependent RNA helicase</fullName>
            <ecNumber evidence="9">3.6.4.13</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">RNA helicase.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="7">
            <text>ATP + H2O = ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:13065"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="3.6.4.13"/>
        </reaction>
    </comment>
    <comment type="domain">
        <text evidence="9">The Q motif is unique to and characteristic of the DEAD box family of RNA helicases and controls ATP binding and hydrolysis.</text>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the DEAD box helicase family. DDX10/DBP4 subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.6.4.13" evidence="9"/>
    <dbReference type="AlphaFoldDB" id="A0A452J002"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038159.1">
        <property type="protein sequence ID" value="ENSGAGP00000033691.1"/>
        <property type="gene ID" value="ENSGAGG00000023967.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="RHEA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003724">
        <property type="term" value="F:RNA helicase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="CDD" id="cd17941">
        <property type="entry name" value="DEADc_DDX10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18787">
        <property type="entry name" value="SF2_C_DEAD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000874">
        <property type="entry name" value="RNA helicase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:001089">
        <property type="entry name" value="RNA helicase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011545">
        <property type="entry name" value="DEAD/DEAH_box_helicase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014001">
        <property type="entry name" value="Helicase_ATP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001650">
        <property type="entry name" value="Helicase_C-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000629">
        <property type="entry name" value="RNA-helicase_DEAD-box_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014014">
        <property type="entry name" value="RNA_helicase_DEAD_Q_motif"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025313">
        <property type="entry name" value="SPB4-like_CTE"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24031">
        <property type="entry name" value="RNA HELICASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13959">
        <property type="entry name" value="CTE_SPB4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00270">
        <property type="entry name" value="DEAD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00271">
        <property type="entry name" value="Helicase_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00487">
        <property type="entry name" value="DEXDc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01178">
        <property type="entry name" value="DUF4217"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00490">
        <property type="entry name" value="HELICc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00039">
        <property type="entry name" value="DEAD_ATP_HELICASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51192">
        <property type="entry name" value="HELICASE_ATP_BIND_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51194">
        <property type="entry name" value="HELICASE_CTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50890">
        <property type="entry name" value="PUA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51195">
        <property type="entry name" value="Q_MOTIF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 9" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="3 9" id="KW-0347">Helicase</keyword>
    <keyword evidence="2 9" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="1 9" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 9" id="KW-0694">RNA-binding</keyword>
    <feature type="domain" description="DEAD-box RNA helicase Q" evidence="13">
        <location>
            <begin position="70"/>
            <end position="98"/>
        </location>
    </feature>
    <feature type="domain" description="Helicase ATP-binding" evidence="11">
        <location>
            <begin position="101"/>
            <end position="275"/>
        </location>
    </feature>
    <feature type="domain" description="Helicase C-terminal" evidence="12">
        <location>
            <begin position="301"/>
            <end position="450"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="563"/>
            <end position="606"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="701"/>
            <end position="720"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="770"/>
            <end position="832"/>
        </location>
    </feature>
    <feature type="short sequence motif" description="Q motif" evidence="8">
        <location>
            <begin position="70"/>
            <end position="98"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="563"/>
            <end position="583"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="10">
        <location>
            <begin position="591"/>
            <end position="604"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022806"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00038084"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00047984"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00552"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="RuleBase" id="RU365068"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS51192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS51194"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS51195"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033691.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="860" mass="98234" checksum="59FB132A368B9D07" modified="2019-05-08" version="1">MGGIGRHVGSGDPDPDPVLNFLRWKKNYNRRKNKLRLQRKEQQRPAGVVEREGISLLEKRYPEINPNEIKRFSDFPLSKKTLKGLQEAQYRMVTEIQRQTIGLALQGKDVLGAAKTGSGKTLAFIVPALERLYRQQWTSIDGLGVLIISPTRELAYQTFEVLRKVGKNHDFSAGLIIGGKDLKKEVTKIHYVNMLICTPGRLLQHMDETSYFHASDLQMLILDEADRILDMGFTDTMNAIIENLPKKRQTLLFSATQTKSVKDLARLSLKDPEYVWVHEKAKFSTPATLDQNYVVCGLQQKINMLYSFLRSHLNKKSIVFYASCKEVQYLFRVFCRLRPGIPILALHGKQPQMKRMEVYNNFVRKKSAVLFATDVAARGLDFPAVNWVIQFDCPEDANTYIHRVGRTARYKEGGEALLVLLPSEEGGMVHQLSQRKVPVNKIKINPEKLVDIQKKMQSFLAQDKELKERAQMCFVSYLRSVYLMKNKEVFDVFKLPLAEYAHSLGLAVEPRVRFLQKAQKQLSVNESLEGKSHPEGTGQCEDACSLVSNEEVEERGTHFKEKISTVQRMKGEGRKEEPEEKELPSQVIDKEGDESEELSEEEEVPSINKIPKAESINFLADDDDDFRDLDLLTVKRQNVFGLEAKDLAVLVSNTSNSELKKKATKTKEAKKILKRKFKVNTKIVFTEDGELVQQWPPVQKSCVDSEEKEDESGGINLDKAKERLQEEDKFDKEVYRKKIKEKHRVIVCLSLKLFLLLQEGVEAFLDYSGNEEEFDPSTLPDPDKYRGSDQEADLDTEDRDGYEKNGTRKRSKSSNNAAEESPLRRKKAKLSDEHSFVPLDTGLSLAEDEELVLHLLNSQK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J006</accession>
    <name>A0A452J006_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Phosphatase and actin regulator</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="5">Binds PPP1CA and actin.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1 5">Belongs to the phosphatase and actin regulator family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J006"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038167.1">
        <property type="protein sequence ID" value="ENSGAGP00000033697.1"/>
        <property type="gene ID" value="ENSGAGG00000023986.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003779">
        <property type="term" value="F:actin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004864">
        <property type="term" value="F:protein phosphatase inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030036">
        <property type="term" value="P:actin cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.1750">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.2130">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004018">
        <property type="entry name" value="RPEL_repeat"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12751:SF5">
        <property type="entry name" value="PHOSPHATASE AND ACTIN REGULATOR 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12751">
        <property type="entry name" value="PHOSPHATASE AND ACTIN REGULATOR PHACTR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02755">
        <property type="entry name" value="RPEL"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00707">
        <property type="entry name" value="RPEL"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51073">
        <property type="entry name" value="RPEL"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 5" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 5" id="KW-0677">Repeat</keyword>
    <feature type="repeat" description="RPEL" evidence="4">
        <location>
            <begin position="60"/>
            <end position="85"/>
        </location>
    </feature>
    <feature type="repeat" description="RPEL" evidence="4">
        <location>
            <begin position="401"/>
            <end position="426"/>
        </location>
    </feature>
    <feature type="repeat" description="RPEL" evidence="4">
        <location>
            <begin position="439"/>
            <end position="464"/>
        </location>
    </feature>
    <feature type="repeat" description="RPEL" evidence="4">
        <location>
            <begin position="477"/>
            <end position="502"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="1"/>
            <end position="33"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="104"/>
            <end position="433"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="460"/>
            <end position="479"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="13"/>
            <end position="26"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="152"/>
            <end position="165"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="173"/>
            <end position="183"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="204"/>
            <end position="231"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="6">
        <location>
            <begin position="245"/>
            <end position="264"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="265"/>
            <end position="275"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="358"/>
            <end position="367"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="6">
        <location>
            <begin position="377"/>
            <end position="387"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="412"/>
            <end position="433"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="460"/>
            <end position="477"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00009795"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00401"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="RuleBase" id="RU301113"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033697.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="588" mass="64778" checksum="455BCA5EA4AFB5E1" modified="2019-05-08" version="1">MDNAVDGLDKASIANSDGPTPGSQTPPFKRKGKLSTIGKIFKPWKWRKKKTSDKFKETSAVLERKISTRQSREELIRKGVLKEMPEQVGDVTVNFETSNGHTITISEETIQEENVVKTTEDNGSLSAKASAVEGKKEDKKASSSHVKKLPISKSSSSQSLSSTSSHPKASKETAPSKTGTSGTARVKKKPGKQSAPRTRADGDVSSTSNAAANSLETKVESSQPEQPSVIISESEDTDQQSKLIVPPPPTAPPPPPPPLLPPSSPSSASQSTGSSETPDVPISTKAGLSVPGSDSNPLLQTEHETDNEDLSSIALDNSLDRSEENTTCSVTKDDQEEETPNLASSQPTEEPPEPMESESSKESHTSDSDSDGPILYTDDDDDDDDNTSAESSLASKIRRRDTLAIKLGNRPSKKELEDKNILQRTSEEERQEIRHQIGTKLVRRLSQRPTTEELEQRNILKQKNAEEEQEARRELKRSLSRKLSLRPTVAELQARRILRFNEYVEVTDSPDYDRRADKPWARLTPADKAAIRKELNEFKSTEMEVHEESRQFTSMSQMKFQQSAGMGGEPCFHRITENYMKVSSSIIV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J009</accession>
    <name>A0A452J009_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="8">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the autoinducer-2 exporter (AI-2E) (TC 2.A.86) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J009"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038149.1">
        <property type="protein sequence ID" value="ENSGAGP00000033683.1"/>
        <property type="gene ID" value="ENSGAGG00000023964.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002549">
        <property type="entry name" value="AI-2E-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21716">
        <property type="entry name" value="TRANSMEMBRANE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21716:SF4">
        <property type="entry name" value="TRANSMEMBRANE PROTEIN 245"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01594">
        <property type="entry name" value="AI-2E_transport"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 7" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 7" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="46"/>
            <end position="70"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="110"/>
            <end position="128"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="140"/>
            <end position="166"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="172"/>
            <end position="196"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="208"/>
            <end position="226"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="232"/>
            <end position="249"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="338"/>
            <end position="359"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="436"/>
            <end position="456"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="607"/>
            <end position="623"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="629"/>
            <end position="647"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="694"/>
            <end position="714"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="720"/>
            <end position="740"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="752"/>
            <end position="771"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="791"/>
            <end position="812"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="285"/>
            <end position="322"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="285"/>
            <end position="297"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="307"/>
            <end position="318"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009773"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033683.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="841" mass="92929" checksum="C42CD822B850DE3A" modified="2019-05-08" version="1">MYSSESPERSAAGPPGAAEGAAPRLLLLPPAAVSSLPRLEKPIKQAFYNTGALLFAGLCCGAAVLVYFILEAFLRPLLWAVLCGTFLHPFKSSLTALGRRWLGRLHRSRTPVLLAALLLPICFANYGAEALGEQVLRRRRLLLLLGAGGPLLYGLYCLGSSLGVQVLLAQTGLLICQGLDFFSSLWIWTLVVGYLLTVSFKWNASTECYLRAISIPVWIILLFHLASVAGSWRIPVFLVIVFLMTVGMLHEQQSGKQSSGTELPGQMISIAASTIAMAISVTGYDSSNEQHSPQSSEHAPPEAWPNSSSSFSSSSSTSGGRHRPEIGAFLRKKKTSDIYFVTLVWAIVVVQIWLNLWIVQLLPVPVAVWVVKKLVIHFGVVNFMAKRCSSWWHVLENFVKERQEALAPRPIRGLGRVMLKLDSKMIVWLEKMLDKIISIFIIFLLVIGTLLLALLLTAKVHQESVHLIQVTSSLINETVANHPEWANWLPEAQVIQKALNSAANNVYQYGREWITHKLHKILGEKVNNTAVIEKQVLELWDRLYHSWFVKNVTHSGRHKGHKLQINRQNSWLGDILDWQDIASFVHENIETFLSILESLWIVMSRNVSLLFTTVTTLVTILFYSGTALLNFVLSLVIFLTTLFYLLSSSDEYYKPVKWVISLTPLSQPGPSSNIVGQSVEEAIRGVFDASLKMAGFYGLYTWLTHTIFGINIVFIPSALAAILGAVPFLGTYWAAIPAILDLWLTQGEGCKAILLLVFHLLPTYFVDTAIYSDISGGGHPYLTGLAVAGGAYYLGLEGAIIGPILLCILVVASNIYSAMLVSPTNSVPTPSQTPWPLQIYQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J016</accession>
    <name>A0A452J016_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Nucleoprotein TPR</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="29">Megator</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="27">NPC-associated intranuclear protein</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="28">Translocated promoter region protein</fullName>
        </alternativeName>
    </protein>
    <organism evidence="35 36">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="36" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="35" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="35" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="5">Chromosome</location>
            <location evidence="5">Centromere</location>
            <location evidence="5">Kinetochore</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Spindle</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Nucleus membrane</location>
            <topology evidence="2">Peripheral membrane protein</topology>
            <orientation evidence="2">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Nucleus membrane</location>
            <topology evidence="4">Peripheral membrane protein</topology>
            <orientation evidence="4">Nucleoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Nucleus</location>
            <location evidence="3">Nuclear pore complex</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the TPR family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038180.1">
        <property type="protein sequence ID" value="ENSGAGP00000033707.1"/>
        <property type="gene ID" value="ENSGAGG00000023969.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000776">
        <property type="term" value="C:kinetochore"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031965">
        <property type="term" value="C:nuclear membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034399">
        <property type="term" value="C:nuclear periphery"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0044615">
        <property type="term" value="C:nuclear pore nuclear basket"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005819">
        <property type="term" value="C:spindle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017056">
        <property type="term" value="F:structural constituent of nuclear pore"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006406">
        <property type="term" value="P:mRNA export from nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090316">
        <property type="term" value="P:positive regulation of intracellular protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046824">
        <property type="term" value="P:positive regulation of nucleocytoplasmic transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006606">
        <property type="term" value="P:protein import into nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:1901673">
        <property type="term" value="P:regulation of mitotic spindle assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.1490:FF:000004">
        <property type="entry name" value="nucleoprotein TPR isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.1490:FF:000005">
        <property type="entry name" value="nucleoprotein TPR isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.1490">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057974">
        <property type="entry name" value="NUA/TPR/MLP1-2-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012929">
        <property type="entry name" value="Nucleoprot-TPR/MLP1-2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057577">
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    <keyword evidence="25" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="13" id="KW-0132">Cell division</keyword>
    <keyword evidence="26" id="KW-0137">Centromere</keyword>
    <keyword evidence="9" id="KW-0158">Chromosome</keyword>
    <keyword evidence="20 30" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="11" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="23" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="16" id="KW-0995">Kinetochore</keyword>
    <keyword evidence="22" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0488">Methylation</keyword>
    <keyword evidence="14" id="KW-0498">Mitosis</keyword>
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    <keyword evidence="21" id="KW-0906">Nuclear pore complex</keyword>
    <keyword evidence="24" id="KW-0539">Nucleus</keyword>
    <keyword evidence="12" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="17" id="KW-0653">Protein transport</keyword>
    <keyword evidence="36" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="19" id="KW-0811">Translocation</keyword>
    <keyword evidence="8" id="KW-0813">Transport</keyword>
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            <begin position="209"/>
            <end position="288"/>
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            <begin position="1079"/>
            <end position="1206"/>
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            <begin position="1"/>
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    <sequence length="2403" mass="271225" checksum="C578FACB8FC4593E" modified="2019-05-08" version="1">PTANLGFHPPTPQAASPIQPAAGLGSQPCPHRRLPPNMAAVLQQVLERAELAKLPKPVQGKLERFLADQQSEIDGLRARHERFKVDSEQQYFEVEKRLAQSQERLVNETQECQTLRQELNKLNEQLKSLHEKNKELEAAHDRNATIQSQLTRAKEELEGEKRDLVRTSERRSQEIEHLNEDVKRLNEKLTEANSAKVELQLKLDELQTSDVSVKYREKRLEQEKELLQNQNTWLHTELKAKTDELLLISREKGNEILELKCSLENKKEEVSRMEEQVNSLKLSNENLQKHVEDLLIKLKEAKEQQAGMEERFHNELNAHIKLSNLYKSAADDSEAKSNELTRAVEELHKLLQEASEANKATEDHLAEMEESKTAMEKELGEKISKLEKELENANDLLSATKRKGAILSEEELTAMSPTAAAVAKVVKPGMKLTELYNAYVETQDQLLLEKRENKRINKYLDEIVQEVEAKAPILKRQREEYERSQKAVASLSAKLEQAMKEIQRLQEDTDKANKHASVLERENQRLEIQVKDLSQQVRVLLMELEEARGNHVIRDEEVSSADISSSSEVISQHLVSYRNIEELQQQNQRLLVALRELGEAREKEEQETTSSRISELQSQLEEAIHELEKLREARHVQLQLVDSIVRQRDMYRVLLAQTTGVVIPVQASSMLPEEISFTSTPKRSTIPQAMSTPAPVSMTESAEAVEAKAALKQLREVFENYRKEKAENDKLLNEQNEKLQEQVTDLRSQNAKMSTQLEFASKRYEMLQDNVEGYRREITSLQERTQKLTATTQKQEQIINTMTQDLRGANEKLAMAEVRAENLKKEKDILKMAEARLTQQRESLLAEQRGQNLLLTNLRTIQVILERSETETKQRLNNQVEKLEREISQLKKKLENEVEQRHVLTKNQDVHLLDTKRQLETEMNRHLNTKELLKNAQKETATLKQQLNNTEAQLASQTSQQAPGKGQSSANEDVDDLVSRLRQAEEQVNDLKERLKTSSSNVEQYRAMVLSLEESLNKEKQVTEEVRATVEARLKDSSEYQAQLEKKLMEAEKEKQELQDEKRKAIESMEQQLSELKKSLSNVQSEVQEALQRAGTAINNEQQARRDCQEQAKIASEAQNKYERELMLHAADVEALQAAKSQVAKNSAIRQQLEEATQKAESALLECKASWEERERMLKDEASKLGSRCEDLEKQNRLLHEQLENLSNKMVTSVKESVPSALNVSLNEEGKSQEQILEILRFIRREKEIAETRFEVAQVESLRYRQRVDHLERELQDLHDSLNAEREKVQITAKTIAQHEELMKKTETMNVLIETNKMLREEKERLEQELQQMQAKVRKLEADILPLQESNAELSEKSGMLQAEKKLLEEDVKRWKARTQHLLSQQKDTDLEEYRKLLSEKEANTKRIQQMSEETGRLKAEIARSNASLTTSQNLVQTLRDELTKTRTEKDTLQKELENKVADIQEKVKTITQVKKIGRRYKTQYEELKAQYDKVVAETSTQSYAEEQEQQASVQEIQELKESLSQAEVKTKALENQIENLQKTIAEKESEARSLQEQISQLQSELSRLHQDLQERTTQEEQLRQQITEKEEKTRKTLLAAKQKIAQLAGTKEQLTKENDEWKQKSSALEEQKSELEVRMSALKSQYEGRICRLERELREQQERHHEQRDEPPESTNKVPEQQRQITLKSTPASGERGIASTSDPPTANIKPTPVVSTPSKVTATAIPGSKSTPRASIRPMVTPATVTNPTTTPTATVMPTTQVETQEAMQSEGPVEHVPVFGSTSGSVRSTSPNVQTSLSQPILTVQQQTQATAFVQPTQQSHPQIEPATQEPPPTIVEVVQSSQIERPSTSTAVFGTVSATPSSSLPKRPREEEEDSTVENAEQISEETVDIPLPKKLRSVQRVGPEEEITAEESTDGEVETQIYNQDSQDSIGEGVTQGEYAPMEDSEETSQSLPIDLGPLQSDQQNTSSSQDGQSKRDDVIVIDSDDEEDDEEENEGEQEVYEEEEDEDDDEDTGMGDEGDDSNEGTGSADGNDGYEADDAEGADGTDPCTETEESMGGGESNQRAADSQNSGEGSTSAAESTFSHESSREQQPTSASERQASRLPQSPRRPPHPLPPRLTIHAPPQELGPPVQRIQMTRRQSVGRGLQLTPGIGGMQQHFFDDEDRTVPSTPTLVVPHRTDGFAEAIHSPQVAGVPRFRFGPPEDMPQTSSSHSDLGQLASQGGLGMYETPLFLAHEEESGGRSVPTTPLQVAAPVTVFTESTTADPSEHASQSVPMVTTSTGSLSTTTESGVGDDGDEVFVEAESEGITSEAGLEIDSQQEEESVQAPDESDLPSTSQDPPSSSSADTSSNQPKPFRRVRLQPPTLRTGVRGRQFNRQRGVTHAMGGRGGLNRGNIS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J018</accession>
    <name>A0A452J018_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Phosphatase and actin regulator</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="5">Binds PPP1CA and actin.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1 5">Belongs to the phosphatase and actin regulator family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J018"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038173.1">
        <property type="protein sequence ID" value="ENSGAGP00000033701.1"/>
        <property type="gene ID" value="ENSGAGG00000023986.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003779">
        <property type="term" value="F:actin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004864">
        <property type="term" value="F:protein phosphatase inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030036">
        <property type="term" value="P:actin cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.1750">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.2130">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004018">
        <property type="entry name" value="RPEL_repeat"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12751:SF5">
        <property type="entry name" value="PHOSPHATASE AND ACTIN REGULATOR 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12751">
        <property type="entry name" value="PHOSPHATASE AND ACTIN REGULATOR PHACTR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02755">
        <property type="entry name" value="RPEL"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00707">
        <property type="entry name" value="RPEL"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51073">
        <property type="entry name" value="RPEL"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 5" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 5" id="KW-0677">Repeat</keyword>
    <feature type="repeat" description="RPEL" evidence="4">
        <location>
            <begin position="70"/>
            <end position="95"/>
        </location>
    </feature>
    <feature type="repeat" description="RPEL" evidence="4">
        <location>
            <begin position="411"/>
            <end position="436"/>
        </location>
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        <location>
            <begin position="449"/>
            <end position="474"/>
        </location>
    </feature>
    <feature type="repeat" description="RPEL" evidence="4">
        <location>
            <begin position="487"/>
            <end position="512"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="1"/>
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        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
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        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="470"/>
            <end position="489"/>
        </location>
    </feature>
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        <location>
            <begin position="1"/>
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        <location>
            <begin position="255"/>
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            <begin position="368"/>
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        <location>
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        <location>
            <begin position="470"/>
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        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00009795"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00401"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="RuleBase" id="RU301113"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033701.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="568" mass="62542" checksum="28B79E683B762DD1" modified="2019-05-08" version="1">MGQTSVSSLSEQASVDGLDKASIANSDGPTPGSQTPPFKRKGKLSTIGKIFKPWKWRKKKTSDKFKETSAVLERKISTRQSREELIRKGVLKEMPEQVGDVTVNFETSNGHTITISEETIQEENVVKTTEDNGSLSAKASAVEGKKEDKKASSSHVKKLPISKSSSSQSLSSTSSHPKASKETAPSKTGTSGTARVKKKPGKQSAPRTRADGDVSSTSNAAANSLETKVESSQPEQPSVIISESEDTDQQSKLIVPPPPTAPPPPPPPLLPPSSPSSASQSTGSSETPDVPISTKAGLSVPGSDSNPLLQTEHETDNEDLSSIALDNSLDRSEENTTCSVTKDDQEEETPNLASSQPTEEPPEPMESESSKESHTSDSDSDGPILYTDDDDDDDDNTSAESSLASKIRRRDTLAIKLGNRPSKKELEDKNILQRTSEEERQEIRHQIGTKLVRRLSQRPTTEELEQRNILKQKNAEEEQEARRELKRSLSRKLSLRPTVAELQARRILRFNEYVEVTDSPDYDRRADKPWARLTPADKAAIRKELNEFKSTEMEVHEESRQFTRFHRP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J023</accession>
    <name>A0A452J023_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000038214.1">
        <property type="protein sequence ID" value="ENSGAGP00000033735.1"/>
        <property type="gene ID" value="ENSGAGG00000024011.1"/>
    </dbReference>
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    </dbReference>
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        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051018">
        <property type="term" value="F:protein kinase A binding"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008382">
        <property type="entry name" value="SPHK1-interactor_AKAP_110"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10226">
        <property type="entry name" value="A KINASE ANCHOR PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10226:SF7">
        <property type="entry name" value="A-KINASE ANCHOR PROTEIN SPHKAP"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <location>
            <begin position="226"/>
            <end position="248"/>
        </location>
    </feature>
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</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J031</accession>
    <name>A0A452J031_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Protein odr-4 homolog</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">May play a role in the trafficking of a subset of G-protein coupled receptors.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the ODR-4 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J031"/>
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        <property type="gene ID" value="ENSGAGG00000024018.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008104">
        <property type="term" value="P:intracellular protein localization"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029454">
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    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR33966:SF1">
        <property type="entry name" value="PROTEIN ODR-4 HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14778">
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    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6" id="KW-1133">Transmembrane helix</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00003891"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004370"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00010131"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00020550"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033745.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="436" mass="48769" checksum="B3D2F170C253F736" modified="2019-05-08" version="1">MGRSYFVEEAVGQYLSDFSTKPKACVTGLLIGQCSPQRDYVILAIRTPPRGEQNESNKNYHPKLADIDEEWVTGHASQLSRMLPGGLLVLGVFIVAAPELAKDIQNALRKLVFSVEKSIGKKRLWNFAEEEVSDRVALHLCSATKKILCRTYDMQDPKSSAKPADWKYQSGLSAFWLSLECTINVNIHIPLSATSVNYELEKNTRKKSSRGNIQHSIQPFDVKVLTQLSQSSGHRSTATVQVCSGSINLKGAMKCRAYIHNNKPKVKDAIQVLKRDIINTLSDRCEILFEDLLINEAPEKKNSEREYHALPHRAFASIAGSNVMLCDYKFGDETAGEIQERFFEMLDQVVHGEAIHIAEEINIVDINPPVDNTDDTQQMQLTKATLLLKFQQNIGKLPPISPLRHCRTDMYSLSLLFAPTIVIVISGCEDIVNGII</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J050</accession>
    <name>A0A452J050_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000038216.1">
        <property type="protein sequence ID" value="ENSGAGP00000033737.1"/>
        <property type="gene ID" value="ENSGAGG00000024011.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051018">
        <property type="term" value="F:protein kinase A binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008382">
        <property type="entry name" value="SPHK1-interactor_AKAP_110"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10226">
        <property type="entry name" value="A KINASE ANCHOR PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10226:SF7">
        <property type="entry name" value="A-KINASE ANCHOR PROTEIN SPHKAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="226"/>
            <end position="248"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="362"/>
            <end position="392"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="534"/>
            <end position="553"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="761"/>
            <end position="791"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1234"/>
            <end position="1264"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1396"/>
            <end position="1433"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1516"/>
            <end position="1551"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="228"/>
            <end position="238"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="371"/>
            <end position="380"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="534"/>
            <end position="551"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="1">
        <location>
            <begin position="764"/>
            <end position="781"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="1410"/>
            <end position="1424"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="1525"/>
            <end position="1540"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033737.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1645" mass="182774" checksum="48DD906E4B2A7642" modified="2019-05-08" version="1">MAGHSLLTVPSNFESPLMYEVSEHQVSGTDSSESSLGSSVTACKKVLCSNSLLESTDYWLQNQRTPCRIGFVEDKSENKCASVCFVNLDANKNDCSEEHLKQRLINVSPNLPKLISSMNVQPPKENEIVLLSGLTSGNLQADFEVSQCPWLADVCLVQCARGNRKNSIGCIIFEINKFLIGLELVQERQLQIETSILKPEDDTNCSVSSIEEDFLTASEHLEDENEADEYKNGHDNIHGTEPASDIKKHKGKGLENLHYRKTTLPFALDDNCFNKENLASAKNSTDSEEALNIVSEDSILQTVDKSIQQIQWKNGLAGRVRSSPNTNSLERSLTDKAENSYPICSSEGVSLTRMAKEELASQCDTTAKQNNKLDTEERTSERKHHPPLQNEQATTGQYATNLAESVLQDAFIRLSQSQPTFTKEAAISISVGNSFSSATCTTEDITSRSWNELPKIVIVQSPDSCENTSEWPGSGFPNLCHWSESESSAEISDYLQEENSKGHTQNVLEVALACAAKVIGTISSPHAAERLKRELATDSKSKTVDNEEEQIKSSQVLDNGADIEYSFPSALCGMTQVASAVAVCGLGETKEDKYSATSRGLLSAAETSAAITLHCSIAIGSSMEKLNRNIAETLLKEASIILTKPNTYRNVGDFMESINGRIIETATRPRISPLDEVICDELAQNVSNVILRHSVEEVRKKRQLHPGSDNNLDFSTHDMFMETANELLFNVIYFTCKKMRDIAQVGECSPLFSEDKDRWRVTEAESQATQTTATQPSQQAPDHTNEPFSTFYSANAGKDLVNMTNTKKDNEEGVPNTMESATLHSDLLCSASGHYSLVAKTSPKKRYLKRATRDCYRSPYQSKKDLRSFSDRENTLTVNECRHGVQEQLSSDAIVNTENQSKHKCDALLNNEVQVSVPLLGNHILLPSQPVLQVKHPRDTYCVTDFAEELAETVVSMATEIAAICLENSNGKQPWFCAWKRGNEYLVTQGLSCRTIKRKKESQPNGSVVRKHRPPRLSEIKRKTDEHPELKERLMNRVVDESINLEDTPESVSIFANEVAAKIMNLTELSMVDNVWQGPNHPRNRLHCERWGRAKGSSCESIPEEDLDSKGSVNTLGPMNMLGQPGSQTSSVSKQSSCESITDEFSRFMVNQMENEGRGFDLLLDYYAGKNASSILTSALQQVARKNCHLNVRPSCPSKQSSTESITEEFYKYMLREIEKENKDDASSIRSSREWSSSLLSPSPRSPFCFRQSSMPDSRSSGSRLTVNAPVKAKSLDGFAHNSHRDSLSIQQVSTVSSLGLCKSDSCLYQRCRTDQVTDMLIHETWASSIESLMRKNKIIGDEAEVTDADQFPSDSPLHVEQYANRLAANIVESGKTLIVVHQDSFDYTNREHVAEGRSLPSVSQSQSKPTRDRINLDGKKEPLTTHGSLQGSHIGSSAFLREVPLIQIETDQREELNKDLEPLTSSIIPAGETKEHLNKEKPPAAYFGKHIVSSSVLNTSVDTRSKPDAELVVETKTAEDFPNPLNSSDESTGSWSQLVNDEDNPDDTSSYLQLSERSMSELAKEKAFLKEQPENIEESTSELSVETANCQKDLLVINFDLEPECPDAELRATLQWIAASELGIRTIYFQRSQENRIEKVLFTM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J053</accession>
    <name>A0A452J053_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="22">Solute carrier family 12 member 7</fullName>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>K(+)(in) + chloride(in) = K(+)(out) + chloride(out)</text>
            <dbReference type="Rhea" id="RHEA:72427"/>
            <dbReference type="ChEBI" id="CHEBI:17996"/>
            <dbReference type="ChEBI" id="CHEBI:29103"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="14">Belongs to the SLC12A transporter family. K/Cl co-transporter subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J053"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038168.1">
        <property type="protein sequence ID" value="ENSGAGP00000033698.1"/>
        <property type="gene ID" value="ENSGAGG00000023984.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045202">
        <property type="term" value="C:synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="GOC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015379">
        <property type="term" value="F:potassium:chloride symporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006884">
        <property type="term" value="P:cell volume homeostasis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007268">
        <property type="term" value="P:chemical synaptic transmission"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055064">
        <property type="term" value="P:chloride ion homeostasis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055075">
        <property type="term" value="P:potassium ion homeostasis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1990573">
        <property type="term" value="P:potassium ion import across plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1740.10:FF:000049">
        <property type="entry name" value="Solute carrier family 12 (potassium/chloride transporter), member 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1740.10:FF:000040">
        <property type="entry name" value="Solute carrier family 12 member 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1740.10">
        <property type="entry name" value="Amino acid/polyamine transporter I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004841">
        <property type="entry name" value="AA-permease/SLC12A_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000076">
        <property type="entry name" value="KCL_cotranspt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018491">
        <property type="entry name" value="SLC12_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004842">
        <property type="entry name" value="SLC12A_fam"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00930">
        <property type="entry name" value="2a30"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11827:SF47">
        <property type="entry name" value="SOLUTE CARRIER FAMILY 12 MEMBER 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11827">
        <property type="entry name" value="SOLUTE CARRIER FAMILY 12, CATION COTRANSPORTERS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00324">
        <property type="entry name" value="AA_permease"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03522">
        <property type="entry name" value="SLC12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01081">
        <property type="entry name" value="KCLTRNSPORT"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="13" id="KW-0868">Chloride</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10" id="KW-0406">Ion transport</keyword>
    <keyword evidence="11 17" id="KW-0472">Membrane</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="8" id="KW-0630">Potassium</keyword>
    <keyword evidence="4" id="KW-0633">Potassium transport</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0769">Symport</keyword>
    <keyword evidence="6 17" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9 17" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="2" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="113"/>
            <end position="136"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="148"/>
            <end position="171"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="192"/>
            <end position="213"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="246"/>
            <end position="265"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="272"/>
            <end position="291"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="411"/>
            <end position="431"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="451"/>
            <end position="477"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="497"/>
            <end position="515"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="551"/>
            <end position="569"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="575"/>
            <end position="597"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="609"/>
            <end position="633"/>
        </location>
    </feature>
    <feature type="domain" description="Amino acid permease/ SLC12A" evidence="18">
        <location>
            <begin position="119"/>
            <end position="291"/>
        </location>
    </feature>
    <feature type="domain" description="Amino acid permease/ SLC12A" evidence="18">
        <location>
            <begin position="412"/>
            <end position="690"/>
        </location>
    </feature>
    <feature type="domain" description="SLC12A transporter C-terminal" evidence="19">
        <location>
            <begin position="820"/>
            <end position="1043"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1"/>
            <end position="47"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022538"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022847"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022958"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023214"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00046331"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00047825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF00324"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="Pfam" id="PF03522"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033698.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="22">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1066" mass="118407" checksum="1E3DAC2B869441CA" modified="2019-05-08" version="1">MPTSFTVVPVEGQEPQPSRPLEDSNQQLAECSSGEPGDGIPKESSPFINSAEMDRGNIYEGKNMALFEEEMDSNPMVSSLLNKLANYTNLTQGVVEHEADEDSKRKEIKTPCMGTFIGVYLPCLQNILGVILFLRLTWIVGTSGVLESFIIVFMCCACTMLTAISMSAIATNGVVPAGGSYYMISRSLGPEFGGAVGLCFYLGTTFAGAMYILGTIEILLTYISPSAAIFKAEDADGETEAMLNNMRVYGTCIVALMAIVVFVGVKYVNKLALVFLSCVILSIFAIYAGVIKTAFDPPDFPICLLGNRTLSRRNFDICAKFIEVNNETVTTKLWSLFCDNRSLSATCDEYFNLNNVTEIQGIPGIASGVLIDNLWSAYTDKGSIIEKKGHPSITGTEEIKVGGLPYVFTDIMTYFTMLVGIYFPSVTGIMAGSNRSGDLRDAQKSIPTGTILAISTTSFIYLSCIVLFGACIEGVILRDKFGEAVNRNLVIGTLAWPSPWVIVIGSFFSTCGAGLQSLTGAPRLLQAIARDGIIPFLQVFGHGKANGEPTWALLLTACICEIGILIASLDSVAPILSMFFLMCYMFVNLACAVQTLLCTPNWRPRFKYYHWALSFLGMSLCFALMFICSWYYALIAMLIAGCIYKYIEYRGAEKEWGDGIRGLSLNAARYALLRVEHGAPHTKNWRPQVLVLLNLDSEQMVKHPRLLSFTSQLKAGKGLTIVGSVLQGIYLDKYIEAQKAEENIKSLMGVEKTKGFCQIVVSPSFRDGMSHLIQSAGLGGMTHNTVLMAWPQSWKQTDNPFSWKNFVDTVRETTAAQQALLVAKNIDLFPTNQERFSEGNVDVWWIVHDGGMLMLLPFLLRQHKVWRKCKMRIFTVAQMDDNSIQMKKDLQMFLYHLRLNAEVEVVEMFENDISAFTYEKTLMMEQRSQMLKQMQLSKNEREREAQLIHDRNTASHSAAVVKCDAAAATPEKVQMTWTKEKFIAEKHKNRETNITGFKDIFNMKPNQSNVRRMHTAVKLNGVILNKSQHAQLVLLNMPGPPKNKKGDENCILCCIVLMVTYTMLQK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J057</accession>
    <name>A0A452J057_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Nucleoprotein TPR</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="29">Megator</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="27">NPC-associated intranuclear protein</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="28">Translocated promoter region protein</fullName>
        </alternativeName>
    </protein>
    <organism evidence="35 36">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="36" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="35" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="35" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="5">Chromosome</location>
            <location evidence="5">Centromere</location>
            <location evidence="5">Kinetochore</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Spindle</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Nucleus membrane</location>
            <topology evidence="2">Peripheral membrane protein</topology>
            <orientation evidence="2">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Nucleus membrane</location>
            <topology evidence="4">Peripheral membrane protein</topology>
            <orientation evidence="4">Nucleoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Nucleus</location>
            <location evidence="3">Nuclear pore complex</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the TPR family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038213.1">
        <property type="protein sequence ID" value="ENSGAGP00000033734.1"/>
        <property type="gene ID" value="ENSGAGG00000023969.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000776">
        <property type="term" value="C:kinetochore"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031965">
        <property type="term" value="C:nuclear membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034399">
        <property type="term" value="C:nuclear periphery"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0044615">
        <property type="term" value="C:nuclear pore nuclear basket"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005819">
        <property type="term" value="C:spindle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017056">
        <property type="term" value="F:structural constituent of nuclear pore"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006406">
        <property type="term" value="P:mRNA export from nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090316">
        <property type="term" value="P:positive regulation of intracellular protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046824">
        <property type="term" value="P:positive regulation of nucleocytoplasmic transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006606">
        <property type="term" value="P:protein import into nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:1901673">
        <property type="term" value="P:regulation of mitotic spindle assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.1490:FF:000004">
        <property type="entry name" value="nucleoprotein TPR isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.1490:FF:000005">
        <property type="entry name" value="nucleoprotein TPR isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.1490">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057974">
        <property type="entry name" value="NUA/TPR/MLP1-2-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012929">
        <property type="entry name" value="Nucleoprot-TPR/MLP1-2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057577">
        <property type="entry name" value="Nucleoprot-TPR/MLP1_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18898:SF2">
        <property type="entry name" value="NUCLEOPROTEIN TPR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18898">
        <property type="entry name" value="NUCLEOPROTEIN TPR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25481">
        <property type="entry name" value="Nucleoprot-TPR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25785">
        <property type="entry name" value="TPR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07926">
        <property type="entry name" value="TPR_MLP1_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="18" id="KW-0007">Acetylation</keyword>
    <keyword evidence="25" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="13" id="KW-0132">Cell division</keyword>
    <keyword evidence="26" id="KW-0137">Centromere</keyword>
    <keyword evidence="9" id="KW-0158">Chromosome</keyword>
    <keyword evidence="20 30" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="11" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="23" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="16" id="KW-0995">Kinetochore</keyword>
    <keyword evidence="22" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0488">Methylation</keyword>
    <keyword evidence="14" id="KW-0498">Mitosis</keyword>
    <keyword evidence="15" id="KW-0509">mRNA transport</keyword>
    <keyword evidence="21" id="KW-0906">Nuclear pore complex</keyword>
    <keyword evidence="24" id="KW-0539">Nucleus</keyword>
    <keyword evidence="12" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="17" id="KW-0653">Protein transport</keyword>
    <keyword evidence="36" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="19" id="KW-0811">Translocation</keyword>
    <keyword evidence="8" id="KW-0813">Transport</keyword>
    <feature type="domain" description="Nucleoprotein TPR/MPL1" evidence="33">
        <location>
            <begin position="172"/>
            <end position="251"/>
        </location>
    </feature>
    <feature type="domain" description="NUA/TPR/MLP1-2-like" evidence="34">
        <location>
            <begin position="470"/>
            <end position="567"/>
        </location>
    </feature>
    <feature type="domain" description="Nucleoprotein TPR/MLP1-2" evidence="32">
        <location>
            <begin position="1028"/>
            <end position="1155"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="31">
        <location>
            <begin position="99"/>
            <end position="135"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="31">
        <location>
            <begin position="1561"/>
            <end position="1583"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="31">
        <location>
            <begin position="1609"/>
            <end position="1686"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="31">
        <location>
            <begin position="1713"/>
            <end position="1749"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="31">
        <location>
            <begin position="1766"/>
            <end position="2064"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="31">
        <location>
            <begin position="2176"/>
            <end position="2301"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="30">
        <location>
            <begin position="219"/>
            <end position="364"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="30">
        <location>
            <begin position="425"/>
            <end position="511"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="30">
        <location>
            <begin position="541"/>
            <end position="594"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="30">
        <location>
            <begin position="659"/>
            <end position="957"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="30">
        <location>
            <begin position="983"/>
            <end position="1158"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="30">
        <location>
            <begin position="1210"/>
            <end position="1423"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="31">
        <location>
            <begin position="115"/>
            <end position="135"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="31">
        <location>
            <begin position="1563"/>
            <end position="1583"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="31">
        <location>
            <begin position="1609"/>
            <end position="1621"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="31">
        <location>
            <begin position="1623"/>
            <end position="1642"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="31">
        <location>
            <begin position="1732"/>
            <end position="1749"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="31">
        <location>
            <begin position="1791"/>
            <end position="1817"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="31">
        <location>
            <begin position="1882"/>
            <end position="1894"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="31">
        <location>
            <begin position="1905"/>
            <end position="1940"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="31">
        <location>
            <begin position="1950"/>
            <end position="1971"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="31">
        <location>
            <begin position="1979"/>
            <end position="2013"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="31">
        <location>
            <begin position="2176"/>
            <end position="2192"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="31">
        <location>
            <begin position="2193"/>
            <end position="2205"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="31">
        <location>
            <begin position="2207"/>
            <end position="2220"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="31">
        <location>
            <begin position="2233"/>
            <end position="2247"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="31">
        <location>
            <begin position="2248"/>
            <end position="2265"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004186"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004335"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004567"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004620"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004629"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00005274"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00019789"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022454"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022776"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022816"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022838"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023010"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00023132"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00023328"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="ARBA" id="ARBA00077074"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="ARBA" id="ARBA00079318"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="ARBA" id="ARBA00083105"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="31">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="32">
        <source>
            <dbReference type="Pfam" id="PF07926"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="33">
        <source>
            <dbReference type="Pfam" id="PF25481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="34">
        <source>
            <dbReference type="Pfam" id="PF25785"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="35">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033734.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="36">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2301" mass="260953" checksum="5F272F2D9B078EE7" modified="2019-05-08" version="1">MAAVLQQVLERAELAKLPKPVQGKLERFLADQQSEIDGLRARHERFKVDSEQQYFEVEKRLAQSQERLVNETQECQTLRQELNKLNEQLKSLHEKNKELEAAHDRNATIQSQLTRAKEELEGEKRDLVRTSERRSQEIEHLNEDVKRLNEKLTEANSAKVELQLKLDELQTSDVSVKYREKRLEQEKELLQNQNTWLHTELKAKTDELLLISREKGNEILELKCSLENKKEEVSRMEEQVNSLKLSNENLQKHVEDLLIKLKEAKEQQAGMEERFHNELNAHIKLSNLYKSAADDSEAKSNELTRAVEELHKLLQEATNKATEDHLAEMEESKTAMEKELGEKISKLEKELENANDLLSATKRKGAILSEEELTAMSPTAAAVAKVVKPGMKLTELYNAYVETQDQLLLEKRENKRINKYLDEIVQEVEAKAPILKRQREEYERSQKAVASLSAKLEQAMKEIQRLQEDTDKANKHASVLERENQRLEIQVKDLSQQVRVLLMELEEARGNHVIRDEEVSSADISSSSEVISQHLVSYRNIEELQQQNQRLLVALRELGEAREKEEQETTSSRISELQSQLEEAIHELEKLREARHVQLQLVDSIVRQRDMYRVLLAQTTGVVIPVQEEISFTSTPKRSTIPQAMSTPAPVSMTESAEAVEAKAALKQLREVFENYRKEKAENDKLLNEQNEKLQEQVTDLRSQNAKMSTQLEFASKRYEMLQDNVEGYRREITSLQERTQKLTATTQKQEQIINTMTQDLRGANEKLAMAEVRAENLKKEKDILKMAEARLTQQRESLLAEQRGQNLLLTNLRTIQVILERSETETKQRLNNQVEKLEREISQLKKKLENEVEQRHVLTKNQDVHLLDTKRQLETEMNRHLNTKELLKNAQKETATLKQQLNNTEAQLASQTSQQSSANEDVDDLVSRLRQAEEQVNDLKERLKTSSSNVEQYRAMVLSLEESLNKEKQVTEEVRATVEARLKDSSEYQAQLEKKLMEAEKEKQELQDEKRKAIESMEQQLSELKKSLSNVQSEVQEALQRAGTAINNEQQARRDCQEQAKIASEAQNKYERELMLHAADVEALQAAKSQVAKNSAIRQQLEEATQKAESALLECKASWEERERMLKDEASKLGSRCEDLEKQNRLLHEQLENLSNKMVTSVKESVPSALNVSLNEEGKSQEQILEILRFIRREKEIAETRFEVAQVESLRYRQRVDHLERELQDLHDSLNAEREKVQITAKTIAQHEELMKKTETMNVLIETNKMLREEKERLEQELQQMQAKVRKLEADILPLQESNAELSEKSGMLQAEKKLLEEDVKRWKARTQHLLSQQKDTDLEEYRKLLSEKEANTKRIQQMSEETGRLKAEIARSNASLTTSQNLVQTLRDELTKTRTEKDTLQKELENKVADIQEKVKTITQVKKIGRRYKTQYEELKAQYDKVVAETSTQSYAEEQEQQASVQEIQELKESLSQAEVKTKALENQIENLQKTIAEKESEARSLQEQISQLQSELSRLHQDLQERTTQEEQLRQQITEKEEKTRKTLLAAKQKIAQLAGTKEQLTKENDEWKQKSSALEEQKSELEVRMSALKSQYEGRICRLERELREQQERHHEQRDEPPESTNKVPEQQRQITLKSTPASGERGIASTSDPPTANIKPTPVVSTPSKVTATAIPGSKSTPRASIRPMVTPATVTNPTTTPTATVMPTTQVETQEAMQSEGPVEHVPVFGSTSGSVRSTSPNVQTSLSQPILTVQQQTQATAFVQPTQQSHPQIEPATQEPPPTIVEVVQSSQIERPSTSTAVFGTVSATPSSSLPKRPREEEEDSTVENAEQISEETVDIPLPKKLRSVQRVGPEVCKNGRYFHEETSQSLPIDLGPLQSDQQNTSSSQDGQSKRDDVIVIDSDDEEDDEEENEDEEEEDEDDDEDTGMGDEGDDSNEGTGSADGNDGYEADDAEGADGTDPCTETEESMGGGESNQRAADSQNSGSTSAAESTFSHESSREQQPTSASERQASRLPQSPRRPPHPLPPRLTIHAPPQELGPPVQRIQMTRRQSVGRGLQLTPGIGGMQHFFDDEDRTVPSTPTLVVPHRTDGFAEAIHSPQVAGVPRFRFGPPEDMPQTSSSHSDLGQLASQGGLGMYETPLFLAHEEESGGRSVPTTPLQVAAPVTVFTESTTADPSEHASQSVPMVTTSTGSLSTTTESGVGDDGDEVFVEAESEGITSEAGLEIDSQQEEESVQAPDESDLPSTSQDPPSSSSADTSSNQPKPFRRVRLQPPTLRTGVRGRQFNRQRGKWLSML</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J058</accession>
    <name>A0A452J058_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Phosphatase and actin regulator</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="5">Binds PPP1CA and actin.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1 5">Belongs to the phosphatase and actin regulator family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J058"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038175.1">
        <property type="protein sequence ID" value="ENSGAGP00000033703.1"/>
        <property type="gene ID" value="ENSGAGG00000023986.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003779">
        <property type="term" value="F:actin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004864">
        <property type="term" value="F:protein phosphatase inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030036">
        <property type="term" value="P:actin cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004018">
        <property type="entry name" value="RPEL_repeat"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12751:SF5">
        <property type="entry name" value="PHOSPHATASE AND ACTIN REGULATOR 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12751">
        <property type="entry name" value="PHOSPHATASE AND ACTIN REGULATOR PHACTR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02755">
        <property type="entry name" value="RPEL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00707">
        <property type="entry name" value="RPEL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51073">
        <property type="entry name" value="RPEL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 5" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 5" id="KW-0677">Repeat</keyword>
    <feature type="repeat" description="RPEL" evidence="4">
        <location>
            <begin position="70"/>
            <end position="95"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="1"/>
            <end position="44"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="127"/>
            <end position="146"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="1"/>
            <end position="13"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="23"/>
            <end position="36"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00009795"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00401"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="RuleBase" id="RU301113"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033703.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="172" mass="18965" checksum="614ECF89F44026C6" modified="2019-05-08" version="1">MGQTSVSSLSEQASVDGLDKASIANSDGPTPGSQTPPFKRKGKLSTIGKIFKPWKWRKKKTSDKFKETSAVLERKISTRQSREELIRKGVLKEMPEQVGDVTVNFETSNGHTITISEETIQEENVVKTTEDNGSLSAKASAVEGKKEDKKVCLIQEVEAREQRDSPRGSVSR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J059</accession>
    <name>A0A452J059_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Protein odr-4 homolog</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">May play a role in the trafficking of a subset of G-protein coupled receptors.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the ODR-4 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J059"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038221.1">
        <property type="protein sequence ID" value="ENSGAGP00000033741.1"/>
        <property type="gene ID" value="ENSGAGG00000024018.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012505">
        <property type="term" value="C:endomembrane system"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008104">
        <property type="term" value="P:intracellular protein localization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029454">
        <property type="entry name" value="ODR-4-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR33966">
        <property type="entry name" value="PROTEIN ODR-4 HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR33966:SF1">
        <property type="entry name" value="PROTEIN ODR-4 HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14778">
        <property type="entry name" value="ODR4-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7 8" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 8" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 8" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="425"/>
            <end position="446"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00003891"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004370"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00010131"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00020550"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033741.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="448" mass="50036" checksum="6041B1F5B2DFA738" modified="2019-05-08" version="1">MGRSYFVEEAVGQYLSDFSTKPKACVTGLLIGQCSPQRDYVILAIRTPPRGEQNESNKNYHPKLADIDEEWVTGHASQLSRMLPGGLLVLGVFIVAAPELAKDIQNALRKLVFSVEKSIGKKRLWNFAEEEVSDRVALHLCSATKKILCRTYDMQDPKSSAKPADWKYQSGLSAFWLSLECTINVNIHIPLSATSVNYELEKNTRNGLTRWAKQIEEGVFLINGQAKDDDAELLEGQKKSSRGNIQHSIQPFDVKVLTQLSQSSGHRSTATVQVCSGSINLKGAMKCRAYIHNNKPKVKDAIQVLKRDIINTLSDRCEILFEDLLINEAPEKKNSEREYHALPHRAFASIAGSNVMLCDYKFGDETAGEIQERFFEMLDQVVHGEAIHIAEEINIVDINPPVDNTDDTQQMQLTKATLLLKFQQNIGVVIAAAVAIFAVAFSFNYFSD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J062</accession>
    <name>A0A452J062_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="20">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Muscle contraction.</text>
    </comment>
    <comment type="subunit">
        <text evidence="15">Muscle myosin is a hexameric protein that consists of 2 heavy chain subunits (MHC), 2 alkali light chain subunits (MLC) and 2 regulatory light chain subunits (MLC-2).</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Myofibril</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 16">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000038219.1">
        <property type="protein sequence ID" value="ENSGAGP00000033739.1"/>
        <property type="gene ID" value="ENSGAGG00000020256.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030016">
        <property type="term" value="C:myofibril"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032982">
        <property type="term" value="C:myosin filament"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016460">
        <property type="term" value="C:myosin II complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000146">
        <property type="term" value="F:microfilament motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd01377">
        <property type="entry name" value="MYSc_class_II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000003">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000004">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000006">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000003">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000007">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000008">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.4820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.530:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.360:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.720:FF:000001">
        <property type="entry name" value="Myosin heavy chain, muscle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000050">
        <property type="entry name" value="Myosin heavy chain, muscle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000101">
        <property type="entry name" value="Slow myosin heavy chain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.340">
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.370">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.4820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.2420">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.360">
        <property type="entry name" value="Myosin S1 fragment, N-terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008989">
        <property type="entry name" value="Myosin_S1_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002928">
        <property type="entry name" value="Myosin_tail"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004009">
        <property type="entry name" value="SH3_Myosin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014751">
        <property type="entry name" value="XRCC4-like_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615">
        <property type="entry name" value="MYOSIN HEAVY CHAIN, NON-MUSCLE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615:SF69">
        <property type="entry name" value="MYOSIN-6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02736">
        <property type="entry name" value="Myosin_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01576">
        <property type="entry name" value="Myosin_tail_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90257">
        <property type="entry name" value="Myosin rod fragments"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57997">
        <property type="entry name" value="Tropomyosin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51844">
        <property type="entry name" value="SH3_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="13 16" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="7 16" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="8" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0488">Methylation</keyword>
    <keyword evidence="11 16" id="KW-0505">Motor protein</keyword>
    <keyword evidence="12" id="KW-0514">Muscle protein</keyword>
    <keyword evidence="10 16" id="KW-0518">Myosin</keyword>
    <keyword evidence="6 16" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0787">Thick filament</keyword>
    <feature type="domain" description="Myosin N-terminal SH3-like" evidence="19">
        <location>
            <begin position="29"/>
            <end position="78"/>
        </location>
    </feature>
    <feature type="domain" description="Myosin motor" evidence="18">
        <location>
            <begin position="82"/>
            <end position="745"/>
        </location>
    </feature>
    <feature type="region of interest" description="Actin-binding" evidence="16">
        <location>
            <begin position="622"/>
            <end position="644"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1220"/>
            <end position="1257"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1851"/>
            <end position="1899"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="17">
        <location>
            <begin position="1220"/>
            <end position="1231"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="1232"/>
            <end position="1248"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="17">
        <location>
            <begin position="1878"/>
            <end position="1899"/>
        </location>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <begin position="175"/>
            <end position="182"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004657"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022433"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023179"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00037488"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00038612"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS51844"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033739.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1899" mass="218627" checksum="04D5185A4D8767CE" modified="2019-05-08" version="1">VDGGVQGGSAPYLRKSDKERLEAQTRIFDLRSDCFVSDEQEEFVKGKIVSRDGERLTVETENGKTVTVKESDVHQQNPPKFDKIEDMAMLTFLHEPAVLYNLKERYAAWMIYTYSGLFCVTVNPYKWLPVYNAEVVAAYRGKKRSEAPPHIFSISDNAYQNMLTNRENQSILITGESGAGKTVNTKRVIQYFASIGTLEDQIIQANPALEAFGNAKTVRNDNSSRFGKFIRIHFGATGKLASADIETYLLEKSRVIFQLKAERNYHIFYQILSNQKPELLEMMLVTNNPYDYSFISQGEVTVASINDSEELIATDSAFDVLGFTPEEKVGVYKLTGAIMHFGNMKFKQKQREEQAEPDGTEDADKSAYLMGLNSADLLKGLCHPRVKVGNEYVTKGQNVQQVYYSIGALAKSVYEKMFSWMVTRINSTLETKQPRQYFIGVLDIAGFEIFDFNSFEQLCINFTNEKLQQFFNHHMFVLEQEEYKKEGIEWVFIDFGMDLQACIDLIEKPMGIMSILEEECMFPKATDMTFKAKLFDNHLGKSANFSKPRNIKGKPEAHFSLIHYAGTVDYNILGWLQKNKDPLNETVVGLYQKSSLKLLALLFKGEEFPLGSYYTPTPSRFLTLLMANLKTTHPHFVRCIIPNERKAPGVMDNPLVMHQLRCNGVLEGIRICRKGFPNRILYGDFRQRYRILNPAAIPEGQFIDSRKGAEKLLSSLDIDPNQYKFGHTKVFFKAGLLGLLEEMRDERLSRIITRIQAQSRGQLMRIEFKKILERRDALLVIQWNIRAFMGVKNWPWMKLYFKIKPLLKSAETEKEMQVRSKEFGRLKEALEKSEARRKELEEKMVSLLQEKNDLQLQVQAEQDNLNDAEERCDQLIKNKIQLEAKVKELTERMEDEEEMNAELTAKKRKLEDECSELKKDIDDLELTLAKVEKEKHATENKVKNLTEEMAGLDEIIAKLTKEKKSLQEAHQQALDDLQVEEDKVNSLTKAKLKLEQQADDLEGSLEQEKKIRMDLERAKRKLEGDLKLTQENIMDLENDKQQLEEKKEFDINQQNSKIEDEQALGLQLQKKLKELQARIEELEEELEAERTARAKVEKLRSDLSRELEEISERLEEAGGATSVQIELNKKREAEFQKMRRDLEEATLQHEATAATLRKKHADSVAELGEQIDNLQRVKQKLEKEKSELKLELDDISSNMEQLLKAKVSLEKLSRTLEDQTAEHRAKLEETQRAMNDTSTQRAKLQTENGEAPPSTALGPLRGTSRALCPPPQAKNALAHALQSARHDCDLLREQYEEETEAKAELQRSLSKANSEVAQWRTKYETDAIQRTEELEEAKKKLAQRLQEAEEAVEAVNAKCSSLEKTKHRLQNEIEDLMVDVERSNAAAAALDKKQRNFDKILSEWKQKFEESQTELESSQKEARSLSTELFKLKNAYEESLEHLETFKRENKNLQEEISDLTEQLGEGGKNLHELEKVRKQLEAEKMELQAALEEAEASLEHEEGKILRAQLEFNQFKADFERKLAEKDEEMEQAKRNHLRVVDSLQTSLDAETRSRNEALRVKKKMEGDLNEMEIQLSHANRMAAEAQSHLKATQTQLKDSQLQLDDFIRANEDLKENIAIVERRNSLLQAELEELRAVVEQTERTRRLAEQELIEASERVHSISLLINQKKKMEADLSQLQTEVEEAVQECRNAEEKAKKAITDAAMMAEELKKEQDTSAHLERMKKNMEQTIKDLQLRLDEAEQLALKGGKKQLQKLEARVRELENELESEQKRNAESIKGLRKSERRIKELTYQTEEDKKNLMRLQDLVDKLQMKVKAYKRQAEEAEEQANTNLAKFRKVQHELDEAEERADIAESQVNKLRAKSRDAGGKWGTSVQSQRSGNSQSQNASGHNREV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J063</accession>
    <name>A0A452J063_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="22">Solute carrier family 12 member 7</fullName>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>K(+)(in) + chloride(in) = K(+)(out) + chloride(out)</text>
            <dbReference type="Rhea" id="RHEA:72427"/>
            <dbReference type="ChEBI" id="CHEBI:17996"/>
            <dbReference type="ChEBI" id="CHEBI:29103"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="14">Belongs to the SLC12A transporter family. K/Cl co-transporter subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J063"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038182.1">
        <property type="protein sequence ID" value="ENSGAGP00000033708.1"/>
        <property type="gene ID" value="ENSGAGG00000023984.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045202">
        <property type="term" value="C:synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="GOC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015379">
        <property type="term" value="F:potassium:chloride symporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006884">
        <property type="term" value="P:cell volume homeostasis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007268">
        <property type="term" value="P:chemical synaptic transmission"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055064">
        <property type="term" value="P:chloride ion homeostasis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055075">
        <property type="term" value="P:potassium ion homeostasis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1990573">
        <property type="term" value="P:potassium ion import across plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1740.10:FF:000049">
        <property type="entry name" value="Solute carrier family 12 (potassium/chloride transporter), member 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1740.10">
        <property type="entry name" value="Amino acid/polyamine transporter I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004841">
        <property type="entry name" value="AA-permease/SLC12A_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000076">
        <property type="entry name" value="KCL_cotranspt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018491">
        <property type="entry name" value="SLC12_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004842">
        <property type="entry name" value="SLC12A_fam"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11827:SF47">
        <property type="entry name" value="SOLUTE CARRIER FAMILY 12 MEMBER 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11827">
        <property type="entry name" value="SOLUTE CARRIER FAMILY 12, CATION COTRANSPORTERS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00324">
        <property type="entry name" value="AA_permease"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03522">
        <property type="entry name" value="SLC12"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01081">
        <property type="entry name" value="KCLTRNSPORT"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="13" id="KW-0868">Chloride</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10" id="KW-0406">Ion transport</keyword>
    <keyword evidence="11 17" id="KW-0472">Membrane</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="8" id="KW-0630">Potassium</keyword>
    <keyword evidence="4" id="KW-0633">Potassium transport</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0769">Symport</keyword>
    <keyword evidence="6 17" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9 17" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="2" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="113"/>
            <end position="136"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="148"/>
            <end position="171"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="192"/>
            <end position="213"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="246"/>
            <end position="265"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="272"/>
            <end position="291"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="411"/>
            <end position="431"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="451"/>
            <end position="477"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="551"/>
            <end position="576"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="582"/>
            <end position="611"/>
        </location>
    </feature>
    <feature type="domain" description="Amino acid permease/ SLC12A" evidence="18">
        <location>
            <begin position="119"/>
            <end position="291"/>
        </location>
    </feature>
    <feature type="domain" description="Amino acid permease/ SLC12A" evidence="18">
        <location>
            <begin position="413"/>
            <end position="584"/>
        </location>
    </feature>
    <feature type="domain" description="SLC12A transporter C-terminal" evidence="19">
        <location>
            <begin position="787"/>
            <end position="928"/>
        </location>
    </feature>
    <feature type="domain" description="SLC12A transporter C-terminal" evidence="19">
        <location>
            <begin position="1018"/>
            <end position="1087"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1"/>
            <end position="47"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022538"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022847"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022958"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023214"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00046331"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00047825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF00324"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="Pfam" id="PF03522"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033708.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="22">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1087" mass="120325" checksum="6BEE2D01CA639873" modified="2019-05-08" version="1">MPTSFTVVPVEGQEPQPSRPLEDSNQQLAECSSGEPGDGIPKESSPFINSAEMDRGNIYEGKNMALFEEEMDSNPMVSSLLNKLANYTNLTQGVVEHEADEDSKRKEIKTPCMGTFIGVYLPCLQNILGVILFLRLTWIVGTSGVLESFIIVFMCCACTMLTAISMSAIATNGVVPAGGSYYMISRSLGPEFGGAVGLCFYLGTTFAGAMYILGTIEILLTYISPSAAIFKAEDADGETEAMLNNMRVYGTCIVALMAIVVFVGVKYVNKLALVFLSCVILSIFAIYAGVIKTAFDPPDFPICLLGNRTLSRRNFDICAKFIEVNNETVTTKLWSLFCDNRSLSATCDEYFNLNNVTEIQGIPGIASGVLIDNLWSAYTDKGSIIEKKGHPSITGTEEIKVGGLPYVFTDIMTYFTMLVGIYFPSVTGIMAGSNRSGDLRDAQKSIPTGTILAISTTSFIYLSCIVLFGACIEGVILRDKFGEAVNRNLVIGTLAWPSPWVIVIGSFFSTCGAGLQSLTGAPRLLQAIARDGIIPFLQVFGHGKANGEPTWALLLTACICEIGILIASLDSVAPILSMALSFLGMSLCFALMFICSWYYALIAMLIAGCIYKYIEYRGAEKEWGDGIRGLSLNAARYALLRVEHGAPHTKNWRPQVLVLLNLDSEQMVKHPRLLSFTSQLKAGKGLTIVGSVLQGIYLDKYIEAQKAEENIKSLMGVEKTKGFCQIVVSPSFRDGMSHLIQSAGLGGMTHNTVLMAWPQSWKQTDNPFSWKNFVDTVRETTAAQQALLVAKNIDLFPTNQERFSEGNVDVWWIVHDGGMLMLLPFLLRQHKVWRKCKMRIFTVAQMDDNSIQMKKDLQMFLYHLRLNAEVEVVEMFENDISAFTYEKTLMMEQRSQMLKQMQLSKNEREREIQSITDESRGSVRRKIYSSPQSIGQDIQAPLPDEEEAQLIHDRNTASHSAAVVKCDAAAATPEKVQMTWTKEKFIAEKHKNRETNITGFKDIFNMKPEWGNLNQSNVRRMHTAVKLNGVILNKSQHAQLVLLNMPGPPKNKKGDENYMEFLEVLTEGLNRVLLVRGGGREVITIYS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J064</accession>
    <name>A0A452J064_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">60S ribosomal protein L3-like</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="4">Heart- and skeletal muscle-specific component of the ribosome, which regulates muscle function. Component of the large ribosomal subunit in striated muscle cells: replaces the RPL3 paralog in the ribosome in these cells. The ribosome is a large ribonucleoprotein complex responsible for the synthesis of proteins in the cell. Inhibits myotube growth and muscle function.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1 6">Belongs to the universal ribosomal protein uL3 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J064"/>
    <dbReference type="Ensembl" id="ENSGAGT00000038226.1">
        <property type="protein sequence ID" value="ENSGAGP00000033746.1"/>
        <property type="gene ID" value="ENSGAGG00000024019.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0022625">
        <property type="term" value="C:cytosolic large ribosomal subunit"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003735">
        <property type="term" value="F:structural constituent of ribosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006412">
        <property type="term" value="P:translation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.30.10:FF:000079">
        <property type="entry name" value="60S ribosomal protein L3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1430.10:FF:000001">
        <property type="entry name" value="60S ribosomal protein L3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.960.10:FF:000001">
        <property type="entry name" value="60S ribosomal protein L3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.960.10:FF:000002">
        <property type="entry name" value="60S ribosomal protein L3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.30.10:FF:000351">
        <property type="entry name" value="Ribosomal protein L3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1430.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.960.10">
        <property type="entry name" value="Ribosomal protein L3, domain 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.30.10">
        <property type="entry name" value="Translation factors"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045077">
        <property type="entry name" value="L3_arc_euk"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044892">
        <property type="entry name" value="Ribosomal_L3_dom_3_arc_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000597">
        <property type="entry name" value="Ribosomal_uL3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019926">
        <property type="entry name" value="Ribosomal_uL3_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009000">
        <property type="entry name" value="Transl_B-barrel_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11363">
        <property type="entry name" value="60S RIBOSOMAL PROTEIN L3-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11363:SF7">
        <property type="entry name" value="RIBOSOMAL PROTEIN UL3-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00297">
        <property type="entry name" value="Ribosomal_L3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50447">
        <property type="entry name" value="Translation proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00474">
        <property type="entry name" value="RIBOSOMAL_L3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 6" id="KW-0687">Ribonucleoprotein</keyword>
    <keyword evidence="2 6" id="KW-0689">Ribosomal protein</keyword>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1"/>
            <end position="36"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="7">
        <location>
            <begin position="1"/>
            <end position="31"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006540"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022980"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023274"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00055008"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00077439"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="RuleBase" id="RU003905"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000033746.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="407" mass="46737" checksum="34245AE6A778E163" modified="2019-05-08" version="1">MSHRKFSAPRHGHLGFLPHKRSRRHRGKVKTWPKDDPSKPVHLTAFLGYKAGMTHILREVHRPGMKLSKREEVEAVTIVETPPLVVVGLVGYIDTPRGLRNFRTIFAEHISDECRRRFYKNWHKSKKKAFTKYCKKWQDKDGKKQLEKDFAALKKYCKVIRVIVHTQMKLLPLRQKKAHVMEIQLNGGTVADKVDWAHEKLEKQVSVHAVFSQNEMIDVIGVTKGHGMKGVTSRWHAKKLPRKTHKGLRKVACIGAWHPARVGYSIARAGQKGYHHRTELNKKIYRIGRGIHVQDGKIVKNNATTNYDVTEKTITPLGGFPHYGAVNNDFMMLKGCVMGTKKRVLTLRKSLLVHSSRRALEPIELKFIDTTSKFGHGCFQTAQEKRAFMGPQKKHLVKEKPESQEEL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHA2</accession>
    <name>A0A452GHA2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">DMAP1-binding domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell projection</location>
            <location evidence="1">Neuron projection</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the DIP2 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000001068.1">
        <property type="protein sequence ID" value="ENSGAGP00000000943.1"/>
        <property type="gene ID" value="ENSGAGG00000000729.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009986">
        <property type="term" value="C:cell surface"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043005">
        <property type="term" value="C:neuron projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd05905">
        <property type="entry name" value="Dip2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.300.30:FF:000003">
        <property type="entry name" value="DIP2 disco-interacting protein 2 homolog A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.300.30:FF:000001">
        <property type="entry name" value="DIP2 disco-interacting protein 2 homolog C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.12780:FF:000004">
        <property type="entry name" value="Disco interacting protein 2 homolog A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.12780:FF:000002">
        <property type="entry name" value="Disco interacting protein 2 homolog B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.300.30">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.12780">
        <property type="entry name" value="N-terminal domain of ligase-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025110">
        <property type="entry name" value="AMP-bd_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045851">
        <property type="entry name" value="AMP-bd_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000873">
        <property type="entry name" value="AMP-dep_synth/lig_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042099">
        <property type="entry name" value="ANL_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037337">
        <property type="entry name" value="Dip2-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010506">
        <property type="entry name" value="DMAP1-bd"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22754">
        <property type="entry name" value="DISCO-INTERACTING PROTEIN 2 DIP2 -RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22754:SF34">
        <property type="entry name" value="DISCO-INTERACTING PROTEIN 2 HOMOLOG A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00501">
        <property type="entry name" value="AMP-binding"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23024">
        <property type="entry name" value="AMP-dom_DIP2-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06464">
        <property type="entry name" value="DMAP_binding"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01137">
        <property type="entry name" value="DMAP_binding"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56801">
        <property type="entry name" value="Acetyl-CoA synthetase-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51912">
        <property type="entry name" value="DMAP1_BIND"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0966">Cell projection</keyword>
    <keyword evidence="4" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="DMAP1-binding" evidence="7">
        <location>
            <begin position="13"/>
            <end position="131"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="67"/>
            <end position="97"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="71"/>
            <end position="89"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004487"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007735"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS51912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000943.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1491" mass="162980" checksum="5117910158D75EBD" modified="2019-05-08" version="1">MAAERGGCSLEAVPVPLPPEVRESLAELELELSEGDITQKGYEKKRAKLLARYIPLIQGVDPSLQAENRIPGSSQATAAVSKQQKSRSTNSRDERFRSDVHTEAVQAALAKYKERKMPMPSKRRSVLVHSSVETYTPPDNHSAPPDVTTGLVEHLHHERPQVASVRGIPRGYNSSILETADGVPVNSRVSSKIQQLLNTLKRPKRPPLKEFFVDDFEELLEVQQPDPNQPKPEGGEMTALKGEPLGVVTNWPPSLLAALQRWGTTQPKTPCLTALDTTGKAVYTLTYGKLWSRSLKLAYTLLNKLTSKNEPLLKPGHRVALVFPNSDPVMFMVAFYGCLLADLVPVPIEVPLTRKDAGSQQIGFLLGSCGVTLALTTDVCQKGLPKAQTGEVVTFKGWPRLVWFVIDGKHLAKPAKDWHPQVRDASADIAYIEYKTSKEGSTMGITISHASMLAHCHALTQACGYSEAETLTNVLDFKRDAGLWHGVLTSVMNRMHVVSIPYALMKVNPLSWIQKVCSYKACVAVVKSRDMHWSLLAQRDQRDVNLSSLRMLIVADGANPWSISSCDAFLNVFQSRGLRPEVICPCASSSEALTVAIRRPPDLGGPPPGKAVLSMNCLSYNVIRVDTEEKLSVLTVQDVGQVMPGANVCVVKADGTPYLCKTDEVGEICVNSGATGTAYYGLLGITKNVFETIPVTAAGVPVSDQPFTRTGLLGFIGPDSLVFVVGKLDGLMVVNIRKHNADDVVATALAVEPMKFVYRGRIAVFSVTVLHDDRIVLVAEQRPDASEEDSFQWMSRVLQAIDSIHQVGVYCLALVPANTLPKAPLGGIHISETKQRFLEGALHPCNVLMCPHTCVTNLPKPRQKQPDVGPASMIVGNLVAGKRIAQASGRDVTQLEDNDQARKFLYLADVLQWRAQTTPDHPLFLLLNSKGTVASTASCIQLHKKAERVAAALMEKGRLNVGDHVALVYPPGVDLIATFYGCLYAGCIPITVRPPHPQNLATTLPTVKMIVEVSKSACILTTQGIMKLLKSKEAATAVDIKTWPTILDTDDIPKKKVANIFRPPSPDMLAYLDFSVSTTGILAGVKMSHAATSALCRSVKLQCELYPSRQIAICLDPYCGLGFALWCLCSVYSGHQSILVPPLELESNVSLWLSAVSQYKVRVTFCSYSVMEMCTKGLGTQTDVLRMKGVNLSCVRTCMVVAEERPRIALTQSFSKLFKDLGLSARSVSTTFGCRVNVAICLQGTAGPDPTTVYVDMRALRHDRVRLVERGSPHSLPLMESGKILPGVKVIIAHTETKGPLGDSHLGEIWVSSPHNATGYYTVYGEEALHADHFSARLSFGDTQTVWARTGYLGFLRRTELTDANGERHDALYVVGSLDETLELRGMRYHPIDIETSVIRAHKSIAECAVFTWTNLLVVVVELEGSEQEALDLVALVTNVVLEEHYLIVGVVVIVDPGVIPINSRGEKQRMHLRDGFLADQLDPIYVAYNM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHA4</accession>
    <name>A0A452GHA4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Zinc finger protein 883-like</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHA4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001092.1">
        <property type="protein sequence ID" value="ENSGAGP00000000967.1"/>
        <property type="gene ID" value="ENSGAGG00000000708.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005654">
        <property type="term" value="C:nucleoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001227">
        <property type="term" value="F:DNA-binding transcription repressor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001817">
        <property type="term" value="P:regulation of cytokine production"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002682">
        <property type="term" value="P:regulation of immune system process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07765">
        <property type="entry name" value="KRAB_A-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001909">
        <property type="entry name" value="KRAB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036051">
        <property type="entry name" value="KRAB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050758">
        <property type="entry name" value="Znf_C2H2-type"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23234:SF8">
        <property type="entry name" value="C2H2-TYPE DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23234">
        <property type="entry name" value="ZNF44 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01352">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="SMART" id="SM00349">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109640">
        <property type="entry name" value="KRAB domain (Kruppel-associated box)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50805">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="13"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="6" id="KW-0539">Nucleus</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <keyword evidence="4 7" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="KRAB" evidence="9">
        <location>
            <begin position="5"/>
            <end position="80"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="8">
        <location>
            <begin position="81"/>
            <end position="108"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="8">
        <location>
            <begin position="109"/>
            <end position="136"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="8">
        <location>
            <begin position="137"/>
            <end position="164"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="8">
        <location>
            <begin position="165"/>
            <end position="192"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="8">
        <location>
            <begin position="193"/>
            <end position="220"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="8">
        <location>
            <begin position="221"/>
            <end position="248"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="8">
        <location>
            <begin position="249"/>
            <end position="276"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="8">
        <location>
            <begin position="277"/>
            <end position="304"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="8">
        <location>
            <begin position="305"/>
            <end position="332"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="8">
        <location>
            <begin position="333"/>
            <end position="360"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="8">
        <location>
            <begin position="361"/>
            <end position="388"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="8">
        <location>
            <begin position="395"/>
            <end position="422"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="8">
        <location>
            <begin position="423"/>
            <end position="450"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000967.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="12">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="455" mass="52347" checksum="157E9DEB08B1CBF0" modified="2025-10-08" version="2">MADPVTFEEVAVYFTREEWALLDPVQRALYRDVMQENYETVTSLGKESWINNKKVKYTLGTHQVEATKEEKDLGGNVKTPYTCTECGKSFSQSSNLITHQRIHTGEKPYTCTECGKSFNWSSTLNTHRRIHTGETPYTCTECGKSFSWRSDLVRHLRIHTGETPYTCTECGRRFSWRSHLIGHQRIHTGETPYTCTECGKSFSQCSNLITHQRIHTGEKPYSCSECGKSFSRSSHLIRHQIIHTGETPYTCTECGKSFNRSSTLNTHRRIHTGKLPYTCTECEKSFSRSSDLIGHQRIHTREKPYTCSECGKSFSCSSDLIRHQRIHTGETPYTCTECGRSFNRSSHLIGHQRIHTGEKPYTCCECEKSFSQSSHLIGHQRIHTGETIHTGETPYTCTECGKSFNRSSTLNTHRRIHTREMPYTCSECGQSFSQRSRLIRHQKMHRRENCNKSLD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHA6</accession>
    <name>A0A452GHA6_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="9">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Lipid-anchor</topology>
            <orientation evidence="1">Cytoplasmic side</orientation>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHA6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001073.1">
        <property type="protein sequence ID" value="ENSGAGP00000000948.1"/>
        <property type="gene ID" value="ENSGAGG00000000769.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008360">
        <property type="term" value="P:regulation of cell shape"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004965">
        <property type="entry name" value="Paralemmin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10498:SF10">
        <property type="entry name" value="PALM2 AND AKAP2 FUSION-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10498">
        <property type="entry name" value="PARALEMMIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03285">
        <property type="entry name" value="Paralemmin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="4 7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="6" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="5" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="358"/>
            <end position="397"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="467"/>
            <end position="515"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="615"/>
            <end position="647"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="802"/>
            <end position="885"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="956"/>
            <end position="1020"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="1"/>
            <end position="34"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="70"/>
            <end position="107"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="379"/>
            <end position="389"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="487"/>
            <end position="500"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="635"/>
            <end position="645"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="802"/>
            <end position="823"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="824"/>
            <end position="848"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="849"/>
            <end position="870"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="871"/>
            <end position="883"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="996"/>
            <end position="1005"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004342"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000948.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1086" mass="121570" checksum="372A08883A6C3975" modified="2019-05-08" version="1">MAEAELHKERLQAIAEKRKRQTEIEGKRQQLEDQILQLQHFKSKALREKWLLQGIPAGSAEEEEARRRQSEEDELKVKKLEGNIHRLEQEIEKLESEESQISAKEQIILEKLKETEKSFEDFQKSFSHQDGDAVNYIYSQIPELPTLYSRTAEPVPGWDRSSRVTGAKGMAVLPKDGRGQCSCESPETAVTGMKSSRIPENDIRLRKDKDQNCANIPEPVSVLAAKEEKMEIEVPVSEHKNITVVDSTHPVDNQTHFFSPVASHNGLKDRHESLDSEVAKEIRYLDEVLEANCCDSAADNTFNGTCSPEPSAILVDASGPPVPVNSESVPNEKDAIVAGRQASPITEQNETSVMAEKLKPNGHSSDGQKEELCGNLKMPSSPSSSTSSRRSSKDGELTLTTIKKEAKFELRAFQEEKKPSKLFDNDNEKEKYSVRKVRPSEEIIELEKERRELIKSQAVKKNPGIAAKWWNPPQEKTLEEQLDEEHLESHKKYKERKERQQQQNTTPTSPKQVAYSFVPPEPGCVKKEDIVTEQIDFSAARKQFQLMENSGQSNSQIAPKRYGAPKMFSIKPFYKPIGPSQIDRPTVSMTRPASVPLENNDITIVKAQKVSCVSEGHPRNIQTSPTDPVRELSYGDTSKTGQTTKLWPDDSEFMSARAVFMVVKDDDPGILDQFSRSVNVSSPPEELDSGLDELSVRSQDTTVLETLSNDFSMDNISDSGASNETMNPLQENSLADFSLPQTPQMDTPSECRVGVSKSFSDHGFYSPSSTLLDSMLVDDQLDYHAGLLVQNAIQQAIAEQADKTGSKQATDKAERKNSVKGQEEVSTAPCSVKEQSTMFEPPQVSSPVQEKKDTIPKTSRDEDSGLREGKTSQQSSVSPTSQPFIVEENRQEVSYFSKYSEAAELRSTASILATQEPEVTVGPFKLRSRKQRTLSMIEEEIRAAQEREEELKRQRQGLQMAQSPITKSAPPLPTRTVAYKTAPGKIEKIRPPPSPTTEGPLSQSDLLPDEAAGSQRPKNLLQTLMEDYETHKTKRRERMDDSSYTCQLLSSKVTSEVLEATRVNRRKSALALRWEAGIYANQEENE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHB3</accession>
    <name>A0A452GHB3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">SAM domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="5">May regulate the disassembly of focal adhesions. Did not bind receptor-like tyrosine phosphatases type 2A.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the liprin family. Liprin-beta subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHB3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001123.1">
        <property type="protein sequence ID" value="ENSGAGP00000000995.1"/>
        <property type="gene ID" value="ENSGAGG00000000781.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048786">
        <property type="term" value="C:presynaptic active zone"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007528">
        <property type="term" value="P:neuromuscular junction development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd09563">
        <property type="entry name" value="SAM_liprin-beta1_2_repeat1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd09566">
        <property type="entry name" value="SAM_liprin-beta1_2_repeat2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd09569">
        <property type="entry name" value="SAM_liprin-beta1_2_repeat3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.50:FF:000005">
        <property type="entry name" value="Liprin-beta-1 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.50:FF:000007">
        <property type="entry name" value="Liprin-beta-1 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.50:FF:000017">
        <property type="entry name" value="Liprin-beta-1 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.50">
        <property type="entry name" value="Transcription Factor, Ets-1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058914">
        <property type="entry name" value="LIPB1/2_CC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037617">
        <property type="entry name" value="LIPB1/2_SAM_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037618">
        <property type="entry name" value="LIPB1/2_SAM_2nd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037619">
        <property type="entry name" value="LIPB1/2_SAM_3rd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029515">
        <property type="entry name" value="Liprin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001660">
        <property type="entry name" value="SAM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013761">
        <property type="entry name" value="SAM/pointed_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12587">
        <property type="entry name" value="LAR INTERACTING PROTEIN LIP -RELATED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12587:SF16">
        <property type="entry name" value="LIPRIN-BETA-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26022">
        <property type="entry name" value="CC_Liprin_beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00536">
        <property type="entry name" value="SAM_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07647">
        <property type="entry name" value="SAM_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00454">
        <property type="entry name" value="SAM"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47769">
        <property type="entry name" value="SAM/Pointed domain"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50105">
        <property type="entry name" value="SAM_DOMAIN"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 6" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="SAM" evidence="8">
        <location>
            <begin position="677"/>
            <end position="741"/>
        </location>
    </feature>
    <feature type="domain" description="SAM" evidence="8">
        <location>
            <begin position="749"/>
            <end position="812"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="433"/>
            <end position="616"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="629"/>
            <end position="664"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="99"/>
            <end position="353"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="461"/>
            <end position="475"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="492"/>
            <end position="504"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="579"/>
            <end position="589"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007547"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00060046"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50105"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000995.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1041" mass="116653" checksum="D362ECA264500B16" modified="2019-05-08" version="1">MMSDASDMLAAALEQMDGIIAGSKALEYSNGIFDCQSPTSPFMGSLRALHLVEDLRGLLEMMEADEKEGLRCQVPDSTAEALIEWLQSQMTNGHISGNGDIYQERLARLENDKESLVLQVSVLTDQVEAQGEKIRDLEFCLEEHREKLNATEEMLQQELLSRTSLETQKLNLMAEISNLKLKLTAVEKDRLDYEDRYGNTESLHNVIADLKQQMEKVEVELALFQEKRLQYGDQEGLMQEISELRLRVGEMDNERLQYEKKLKSTKDELASLKEKLEQKEAEVRKLQEKLVCKMKGEGVEITDRDENCKKKLKDKNIEVQKMKKAVESLMAANEEKDRKIEELRQSLNRYKKVQDMVILAQGKKGKEGECEEFLNSGSVSAVLLDMQSLTDQEKSPSPVPVISFPSHEEFNTTVDEENSLQIHTSILQVSIPAFPPAPESSEAATEKVNAQPKPEVASEVSEGTTPCSSVETQPCDSPLMSALQKSSSLGNLKKEASEVDKEPVQKPAETKAPMEGNKCGTLPPKSPSHSGTGDEDSFGTRKARSSFGRGFFKIKSNKRTASAPNLDRSRSASAPTLAETEKGSADHLDLAGLPPRPKETDSGQMSPSSPDSKKKARGIKKLFGRLKRSQSTTFNPDDMSEAEFKRGGTRATAGPRLGWSRDLGQSNSDLDMPFAKWTKEQVCNWLQGQGLGSYMNHGKHWILSGQTLLQASQQDLEKELGIKHPLHRKKLQLALQALGSEEENNHGKLDYNWVTRWLDDIGLPQYKTQFDEGRVDGRMLHYMSVDDLLSLKVVSVLHHLSIKRAIQVLRINNFEPNCLRRRPSDENNITPAEVTQWTNHRVMEWLRSVDLAEYAPNLRGSGVHGGLMVLEPRFNVETMAQLLNIPPNKTLLRRHLATHFNLLVGQDAQHQKREAMESPDYVLLTATAKVKPKKLAFSNFGSLRKKKQDDVEEYVCPMELGQASGSGSKKGFKAGLDIRVYDDDDLDRLEQMEDSEGTVRQIGAFSEGINNLTHMLKEDEMFKDFATCSPSASITDEDSNV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHB4</accession>
    <name>A0A452GHB4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Cullin family profile domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="pathway">
        <text evidence="1">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="similarity">
        <text evidence="2 5 6">Belongs to the cullin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHB4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001107.1">
        <property type="protein sequence ID" value="ENSGAGP00000000980.1"/>
        <property type="gene ID" value="ENSGAGG00000000219.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031461">
        <property type="term" value="C:cullin-RING ubiquitin ligase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031625">
        <property type="term" value="F:ubiquitin protein ligase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006511">
        <property type="term" value="P:ubiquitin-dependent protein catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1310.10:FF:000012">
        <property type="entry name" value="Cullin 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1310.10:FF:000016">
        <property type="entry name" value="Cullin 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1310.10:FF:000018">
        <property type="entry name" value="Cullin 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.230.130:FF:000003">
        <property type="entry name" value="Cullin 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1310.10:FF:000022">
        <property type="entry name" value="Cullin-2 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1310.10">
        <property type="entry name" value="Cullin Repeats"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.230.130">
        <property type="entry name" value="Cullin, Chain C, Domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045093">
        <property type="entry name" value="Cullin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR059120">
        <property type="entry name" value="Cullin-like_AB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016158">
        <property type="entry name" value="Cullin_homology"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036317">
        <property type="entry name" value="Cullin_homology_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001373">
        <property type="entry name" value="Cullin_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016159">
        <property type="entry name" value="Cullin_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11932">
        <property type="entry name" value="CULLIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00888">
        <property type="entry name" value="Cullin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26557">
        <property type="entry name" value="Cullin_AB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00182">
        <property type="entry name" value="CULLIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF75632">
        <property type="entry name" value="Cullin homology domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF74788">
        <property type="entry name" value="Cullin repeat-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50069">
        <property type="entry name" value="CULLIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0832">Ubl conjugation</keyword>
    <keyword evidence="3" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="Cullin family profile" evidence="7">
        <location>
            <begin position="386"/>
            <end position="618"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006019"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00330"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="RuleBase" id="RU003829"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50069"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000980.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="670" mass="78343" checksum="2E8F81943D8D8479" modified="2019-05-08" version="1">MSLKPRVVDFDETWNKLLTTIKAVVMLDYVERATWNDRFSDIYALCVAYPEPLGERLYTETKFFLENHVRHLHKRVLESEEQVLVMYHRYWEEYSKGADYMDCLYRYLNTQFIKKNKLTEADLQYGYGGVDISEPLMEIGELALDMWRKLMIEPLQGILIRMLLREIKNDRCGEDPNQKVIHGVINSFVHVEQYKKKFPLKFYQEIFECPFLNETGEYYKQEASNLLQESNCSQYMEKVLGRLKDEEMRCRKYLHPSSYGKVIHECQQRMVADHLQFLHAECHNIIRQEKRSDMANMYTLLRAVSSGLPHMIQELQNHIHDEGLRATSNLSQENMPTQFVESVLEVHSKFVQLINTVLNGDQHFMSALDKALTSVVNYREPKSICKAPELLAKYCDNLLKKSAKGMTENEVEDKLTSFITVFKYIDDKDVFQKFYARMLAKRLIHGLSMSMDSEEAMINKLKQACGYEFTSKLHRMYTDMSVSADLNNKFNNFIKNQDAIIDLGISFQIYVLQAGAWPLTQAPSSTFAIPQELEKSVQMFELFYSQHFSGRKLTWLHYLCTGEVKMNYLCKPYVAMVTTYQMAVLLAFNNSETVSYKELQDSTQMNEKELTKTIKSLLDVKMINHDLDKEDIEAESTFSLNLNFSSKRTKFKITTSMQKDTPQASIFQYL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHC0</accession>
    <name>A0A452GHC0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">protein-serine/threonine phosphatase</fullName>
            <ecNumber evidence="3">3.1.3.16</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29035"/>
        </cofactor>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the PP2C family.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.16" evidence="3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001141.1">
        <property type="protein sequence ID" value="ENSGAGP00000001010.1"/>
        <property type="gene ID" value="ENSGAGG00000000310.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005654">
        <property type="term" value="C:nucleoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004722">
        <property type="term" value="F:protein serine/threonine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="CDD" id="cd00143">
        <property type="entry name" value="PP2Cc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.60.40.10:FF:000023">
        <property type="entry name" value="Protein phosphatase, Mg2+/Mn2+-dependent, 1G"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.60.40.10">
        <property type="entry name" value="PPM-type phosphatase domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015655">
        <property type="entry name" value="PP2C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036457">
        <property type="entry name" value="PPM-type-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001932">
        <property type="entry name" value="PPM-type_phosphatase-like_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13832:SF803">
        <property type="entry name" value="PROTEIN PHOSPHATASE 1G"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13832">
        <property type="entry name" value="PROTEIN PHOSPHATASE 2C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00481">
        <property type="entry name" value="PP2C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00332">
        <property type="entry name" value="PP2Cc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81606">
        <property type="entry name" value="PP2C-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51746">
        <property type="entry name" value="PPM_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="6" id="KW-0460">Magnesium</keyword>
    <keyword evidence="8" id="KW-0464">Manganese</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="PPM-type phosphatase" evidence="10">
        <location>
            <begin position="182"/>
            <end position="436"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="141"/>
            <end position="256"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="445"/>
            <end position="467"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="9">
        <location>
            <begin position="209"/>
            <end position="237"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="238"/>
            <end position="253"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="446"/>
            <end position="455"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001936"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006702"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00013081"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023211"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS51746"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001010.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="467" mass="50561" checksum="C002B4BCEE0840B0" modified="2019-05-08" version="1">GPAFPSGLSAQLALCPGPLGGARFSSNLCSLSGAEVALYCAKYLPEIIKEQKAYKEGKLQKALEDAFLAIDVKLTTEEVIKELAQIAGRPQDFREEKEKVADEDDVDNEEAALRHEEATMTIEEPLTRYGQNCIKNLRNKPLAPAGEMGQEHGGEVSVNGESSPGELAKHRERKNGKPDDEATGISSTTDGPGTPECSEDEDGHSSETAEIEEDEGDTEEIEEDEGDTEEIEEDEDKEEKIMLPRMERKKEPGSDSGTTAVVALIRGKQLIVANAGDSRCVVSEXEGGKAVDMSHDHKPEDELELARINNAGGKVTMDGRVNGRLNLSRAIGDHFYKQNKNLPPEEQMISALPDIKVLTINEEHDFMVIACDGIWNVMSSQEVVDFIQAKISQKDENGDLRPLSSIVEELLDQCLAPQASGDGIGCDNMTCIIILASDSGKRKLAAAEAQAAGEENGSDSSKKAKRD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHC1</accession>
    <name>A0A452GHC1_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000001119.1">
        <property type="protein sequence ID" value="ENSGAGP00000000991.1"/>
        <property type="gene ID" value="ENSGAGG00000000775.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060271">
        <property type="term" value="P:cilium assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR33487">
        <property type="entry name" value="CILIA- AND FLAGELLA-ASSOCIATED PROTEIN 54"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR33487:SF1">
        <property type="entry name" value="CILIA- AND FLAGELLA-ASSOCIATED PROTEIN 54"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="409"/>
            <end position="431"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="731"/>
            <end position="761"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="1">
        <location>
            <begin position="417"/>
            <end position="427"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="733"/>
            <end position="752"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000991.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2348" mass="265774" checksum="042DC77028A5714D" modified="2019-05-08" version="1">MNKIKKNKLSKAIFLMQKAILMFPKGSASSSQKHLLEEALALIQKVEAEQNALYSSLKQAMTSKKNRIPPPPLLLSRSHCSMTFKPAPFTSDVQVSWYSILGCVAAGSNLKVRLNNHHLPNAGEEIPADGKSLLEIKGLDPNEKYIFAVAAYSSDGKLIGDAIGETTKPILAYPPLSATTARAYLTQIAYQTDNYTLAKKAFSPLWDYFVLPPSTPSADPAVISASNNLTISQNRLCLDAVSQTSPILLHLFLRSIFIINEINVKEGALFCDSICSKEILYNGQVARLAECERMVVAIELSNWLNDANYALQSVVQCYGLLAPIIYHKIASVPVVQILIKCLTVLQEVPSSTLQRKQTGCYESIQHMIACSSFYTAQVLRSWKEYELALIIINYGKKLLDSSQTSHGQSKFSSIEEMQIDPETEDESPYSKKKSQVAAVEKAAENLAALESNLLKLTRPGTGPEITGQEDPFFLYPIISCWTTKNAYREVMKFRKKPRFLEFFVQVLHKALNEEKFQRIVEWTGSVKDFLRRRNNHLLGIKGTLAKEVSELSVYEDAKRHTTAVVEFQKSVETSALRKPRKEVMSPFRGRESLRQYFLRHSSLGKSPEAQRKQRDEIEKVAFHTLVVLLNPIVSNYLKRKRFHQVCLEEMPWRSQMNIYLAITHFNLFKKRLEEQYNIGVRSLQHLNSYDVLDPEIFSLNNSGIIVVRGGTDDNIGTLSHLLIKRIQSGSSRKITDPDKSSERSTKFSEADTPRSQMTNDTELSITLSAREHERPMSNAILLEHFTKTFLHLRRAMVLAHRGGHWTLLQNTCRELWNHSQEALSIAKHSHSLQGLFPITRNFLISTIWLPFYLASDAILDMIIELQTSNSLKIFDPKGDFCVPSCVGGIADEDGGSNLTFCCPLDDISVVDLRWICGLVLRTIELMYQMKKWEALVHIAMHFNTVTHERYAEQVTPLLVYAQRQLLERIQQFRGPDSPQPHFIKYQGDNANKINCRNFIGKQLRIASTFNEVTVSGHHFVPEMPDTYTDGSQAQALVCVPLDVTDTLKCFRETLEKSKYHSRALRHSRKLLSLFLAHTQETVGRVPNQIFTNGKVEFSVGAELTFLPKPPDLSQEKYNFPSMVESRPLPESQLSVVVSSYEKTIEILQINNQRDLKVQALHELGNLHFYAGKKRSAFKYWCQALDETLKIADVLDNWQELEGFSKNVPDDFTDGSSKHFSEKCIYQAGIWGCLHAAVLAAKIAQYILTSSVRLRTKCCIFSAILFKTLFKVSIPHPTADCDFAQYEIHRLIPGIDLFSDRYRADVSTVVASLSFLIFELHCAKQNLIVLPLFTLYQYFVSEICRDPVKCIEGRILKIKVLTDLGFFAEAFYELCLLNHGEKIPWKVSAGYRLTGKMMSFQKFNASKPLLSNANLQVLEDIFNWSLPPAVVSMCDQQIMNKLTLAKMHFIICLSATINSIPEKVMKTIYSMDNTSLKKPSRIMESSIKINAEVTRHQEQSSIIVQLVRCKDELNMAMLKGILLTETEERLNSLVANIQNEYDGKISRCSAVELETVIEAKLQLAVISQQRHQTAFSAALAFSAIRLLQDANIFRKNVPHIQLNIQRTERRDSYVDSSQLPYNVIAREHMNIRLWLRCRMALVTALTAQIRGIGDMKENDMTDCSSLINEVIIEAEAFSDIETQAEVLVQAVILDLQERRPVTGIKLLLQDIINLLQEKPFISPMASLTLVKSMLLLADILTTQTDEDTEHHSSAIEQLNLLILAHKFTIKQIFTLGAPIKQCIEDPALTSPVVPLKNNYLPVINLLAKVKMRIAHALAQEVASTSKRENPLEWLPALSHLETALDLCRTSATRELDLEAEILFQKGKVERQIIKAGNNKSPRAAETLLDAIKVSQRNDQNFGLIRKSYLEIALLYFHLATYKEETSQTVKIEASKTKTTPKEHLASTERIPHITEVYRTQAWIAVRAATQVSEAMLASQLLIGKKGVKGHQVKAITEQNIPEFASMDLLASYKDFLSDGYEVRYKNLMASSIKNEEIIQSEKDQIESEGTECNISKDIQRKQTITWVHLIRYHNYLIRHNMSPLFAVSKSTPGSFSMEDALFTSVFDTGVVLRLAEMHFFLKKHLYIYSTCCLEDFPKQLQFLEKPGASLTAVPVKMSLEILGAPHKMSTVRIASTVSFCTDTDIQLDNRATIAPNKELCIQWYIPSLEKSTKDMETMVLLLYAYNTKYVKIVSIKTFNSANVFSGHLWIPLTRIISLHEKLSNVKQQIEILVQSSTSFPLQEQSERQRRMPSTKSKINLAKVELDEKIEIPFDVTLQSIQNLEKMFDPANGCVIAGGSLFSWVISLLP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHC4</accession>
    <name>A0A452GHC4_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="7">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GHC4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001147.1">
        <property type="protein sequence ID" value="ENSGAGP00000001015.1"/>
        <property type="gene ID" value="ENSGAGG00000000810.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034605">
        <property type="term" value="P:cellular response to heat"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd19499">
        <property type="entry name" value="RecA-like_ClpB_Hsp104-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.8.60:FF:000040">
        <property type="entry name" value="caseinolytic peptidase B protein homolog isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000641">
        <property type="entry name" value="caseinolytic peptidase B protein homolog isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.8.60">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003593">
        <property type="entry name" value="AAA+_ATPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003959">
        <property type="entry name" value="ATPase_AAA_core"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019489">
        <property type="entry name" value="Clp_ATPase_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001270">
        <property type="entry name" value="ClpA/B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050130">
        <property type="entry name" value="ClpA_ClpB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11638">
        <property type="entry name" value="ATP-DEPENDENT CLP PROTEASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11638:SF93">
        <property type="entry name" value="MITOCHONDRIAL DISAGGREGASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07724">
        <property type="entry name" value="AAA_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10431">
        <property type="entry name" value="ClpB_D2-small"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00300">
        <property type="entry name" value="CLPPROTEASEA"/>
    </dbReference>
    <dbReference type="SMART" id="SM00382">
        <property type="entry name" value="AAA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM01086">
        <property type="entry name" value="ClpB_D2-small"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="2" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="1" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <feature type="repeat" description="ANK" evidence="3">
        <location>
            <begin position="120"/>
            <end position="156"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="3">
        <location>
            <begin position="193"/>
            <end position="225"/>
        </location>
    </feature>
    <feature type="domain" description="AAA+ ATPase" evidence="5">
        <location>
            <begin position="301"/>
            <end position="445"/>
        </location>
    </feature>
    <feature type="domain" description="Clp ATPase C-terminal" evidence="6">
        <location>
            <begin position="499"/>
            <end position="588"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="588"/>
            <end position="609"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="SMART" id="SM00382"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="SMART" id="SM01086"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001015.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="636" mass="71829" checksum="C295CCADD50BDC9B" modified="2019-05-08" version="1">MGTGTGGGRVPGLQELGMDMGEFRTPGLQELGMDMGEFRVPWLQELGPCSTLARAPELPFLVAPGPELELQYPGPAVEWCVRLRPDPTGHQKKRESWTRRAVLPAFAAALVYCYSKSHASKEDALLEAARINNVSEVDSVVKILLAAGADPNLGDEFNSVYETAKEKGLHSLEVLVTREDDFNNRLNNRASFKGCTALHYAVLADDYVTVKLLLDGGASPLQRNEMGHTPLDYARDGEVMKLLKASETKFQEEQRKRELEERRRFPLEQRLKEHIIGQENAIATVGAAIRRKENGWYDEEHPLVFLFLGSSGIGKTELAKQTAKYMHKDAKKGFIRLDMSEFQERHEVAKFIGSPPGYIGHEEGGQLTKKLKQCPNAVVLFDEVDKAHPDVLTIMLQLFDEGRLTDGKGKTIDCKDAIFIMTSNVASDEIAQHALQLRQEALEMSRNRIAENLDDVQLTDKITISKQFKENVIRPILKAHFRRDEFLGRINEIVYFLPFCHSELIQLVSKELSFWAKRAKVRHNITLVWDREVMDVLADGYNLHYGARSIKHEVERRVVNQLAAAYEQDLLPRGCTLRITVDDSDKQLLKTKDSSSPEEEEKKAPTLRLEIVEKDSKSRKLDIQAPVSPDEISFML</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHC5</accession>
    <name>A0A452GHC5_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="9">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the cathelicidin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000001139.1">
        <property type="protein sequence ID" value="ENSGAGP00000001008.1"/>
        <property type="gene ID" value="ENSGAGG00000000799.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061844">
        <property type="term" value="P:antimicrobial humoral immune response mediated by antimicrobial peptide"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042742">
        <property type="term" value="P:defense response to bacterium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.450.10:FF:000003">
        <property type="entry name" value="Cathelicidin antimicrobial peptide"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.450.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001894">
        <property type="entry name" value="Cathelicidin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018216">
        <property type="entry name" value="Cathelicidin_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046350">
        <property type="entry name" value="Cystatin_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10206">
        <property type="entry name" value="CATHELICIDIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10206:SF0">
        <property type="entry name" value="CATHELICIDIN B1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00666">
        <property type="entry name" value="Cathelicidins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54403">
        <property type="entry name" value="Cystatin/monellin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00947">
        <property type="entry name" value="CATHELICIDINS_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0044">Antibiotic</keyword>
    <keyword evidence="4" id="KW-0929">Antimicrobial</keyword>
    <keyword evidence="6" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0964">Secreted</keyword>
    <keyword evidence="8" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="8">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019379755" evidence="8">
        <location>
            <begin position="21"/>
            <end position="203"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="181"/>
            <end position="203"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005320"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022529"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023022"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001008.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="203" mass="21807" checksum="E3B44EC80A54DBA2" modified="2019-05-08" version="1">METCCRCLLLLGIVAAAATAVPXAVPSQHKTLSYEEAVALAVDFYNQESGIDHAFRLLKAGPQPEWDVFSKTIQSLKFTVKETVCPVSGNRDPNQCEFKEDGLVKDCSGFFSTEQDPPSIIIKCEEASEEVSGLDSLLSAHGWADAGLRVSLAEILARYFTLHKCPQNKCPAPPTGSELAKASCPTSLEVEPTAASRTAGSKF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHC9</accession>
    <name>A0A452GHC9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="30">ATP-dependent DNA helicase DDX11</fullName>
            <ecNumber evidence="28">5.6.2.3</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="32">DEAD/H-box protein 11</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="31">DNA 5'-3' helicase DDX11</fullName>
        </alternativeName>
    </protein>
    <organism evidence="34 35">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="35" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="34" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="34" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="29">
            <text>ATP + H2O = ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:13065"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="5.6.2.3"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>[4Fe-4S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:49883"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Cytoskeleton</location>
            <location evidence="3">Microtubule organizing center</location>
            <location evidence="3">Centrosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="5">Cytoplasm</location>
            <location evidence="5">Cytoskeleton</location>
            <location evidence="5">Spindle pole</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Midbody</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Nucleus</location>
            <location evidence="4">Nucleolus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the DEAD box helicase family. DEAH subfamily. DDX11/CHL1 sub-subfamily.</text>
    </comment>
    <dbReference type="EC" id="5.6.2.3" evidence="28"/>
    <dbReference type="AlphaFoldDB" id="A0A452GHC9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001096.1">
        <property type="protein sequence ID" value="ENSGAGP00000000970.1"/>
        <property type="gene ID" value="ENSGAGG00000000211.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030496">
        <property type="term" value="C:midbody"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005730">
        <property type="term" value="C:nucleolus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000922">
        <property type="term" value="C:spindle pole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051539">
        <property type="term" value="F:4 iron, 4 sulfur cluster binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043139">
        <property type="term" value="F:5'-3' DNA helicase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016818">
        <property type="term" value="F:hydrolase activity, acting on acid anhydrides, in phosphorus-containing anhydrides"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006281">
        <property type="term" value="P:DNA repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006260">
        <property type="term" value="P:DNA replication"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034085">
        <property type="term" value="P:establishment of sister chromatid cohesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd18788">
        <property type="entry name" value="SF2_C_XPD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:001050">
        <property type="entry name" value="ATP-dependent DNA helicase DDX11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000910">
        <property type="entry name" value="probable ATP-dependent DNA helicase DDX11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000909">
        <property type="entry name" value="Putative ATP-dependent RNA helicase DDX11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006555">
        <property type="entry name" value="ATP-dep_Helicase_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045028">
        <property type="entry name" value="DinG/Rad3-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014013">
        <property type="entry name" value="Helic_SF1/SF2_ATP-bd_DinG/Rad3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006554">
        <property type="entry name" value="Helicase-like_DEXD_c2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010614">
        <property type="entry name" value="RAD3-like_helicase_DEAD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013020">
        <property type="entry name" value="Rad3/Chl1-like"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00604">
        <property type="entry name" value="rad3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11472:SF41">
        <property type="entry name" value="ATP-DEPENDENT DNA HELICASE DDX11-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11472">
        <property type="entry name" value="DNA REPAIR DEAD HELICASE RAD3/XP-D SUBFAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06733">
        <property type="entry name" value="DEAD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13307">
        <property type="entry name" value="Helicase_C_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00488">
        <property type="entry name" value="DEXDc2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00491">
        <property type="entry name" value="HELICc2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51193">
        <property type="entry name" value="HELICASE_ATP_BIND_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0004">4Fe-4S</keyword>
    <keyword evidence="22" id="KW-0010">Activator</keyword>
    <keyword evidence="16" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="9" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="25" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="7" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="13" id="KW-0227">DNA damage</keyword>
    <keyword evidence="24" id="KW-0234">DNA repair</keyword>
    <keyword evidence="10" id="KW-0235">DNA replication</keyword>
    <keyword evidence="21" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="15" id="KW-0347">Helicase</keyword>
    <keyword evidence="14" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="18" id="KW-0408">Iron</keyword>
    <keyword evidence="19" id="KW-0411">Iron-sulfur</keyword>
    <keyword evidence="26" id="KW-0413">Isomerase</keyword>
    <keyword evidence="11" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="12" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="27" id="KW-0539">Nucleus</keyword>
    <keyword evidence="35" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="17" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="23" id="KW-0804">Transcription</keyword>
    <keyword evidence="20" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="Helicase ATP-binding" evidence="33">
        <location>
            <begin position="16"/>
            <end position="425"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001966"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004214"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004604"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004647"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00008435"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022485"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022705"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022806"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023014"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00023204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00023235"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="ARBA" id="ARBA00044969"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="ARBA" id="ARBA00048954"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="ARBA" id="ARBA00074489"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="31">
        <source>
            <dbReference type="ARBA" id="ARBA00079245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="32">
        <source>
            <dbReference type="ARBA" id="ARBA00080369"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="33">
        <source>
            <dbReference type="PROSITE" id="PS51193"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="34">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000970.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="35">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="893" mass="101324" checksum="DB76FDE4C2B34726" modified="2019-05-08" version="1">MCFLCQKAAGAAQGKGRVNFPFPYTPYSIQEDFMANLYDVLEAGKVGIFESPTGTGKSLSLICGALSWLRDFEEKKKQEEARLLADEHTGELVWITEFMQKKEEQAMVDRLKEEQIRRKKREDRLEKIRHNVQLKYASKRKRLEDDETQRLLQLSKEILSTELGSDVLEQLDHEEEELILAEYESDEEKKGRQTALLEFLQSSFSFSSQIYYCSRTHSQLAQFVHEVQKSPFGEEIRLVSLGSRQNLCVNEEVRHLGAVQLINDRCVEMQKNKHGKKSEGEESEGKKRRVSRAVCPFYSYEQMQFLRDEVLVKVKDIEQLVALGKETKACPYYGSRYAIPAAQLVVLPYQMLLHESTRCASGIKLKDQVVIIDEAHNLIDTITCIYSAEVRGSQLCCAHSQLLQYMERYRKRLKAKNLMYIKQILYLLERFVSILGGNVNQNPNAQSISQAGTELKSINDFLFQSQIDNINLFKCVCILCKSYFLFMWQLFGFVERYGATSVVKINKEKQKMAGLQNFLLTLHQGPDKAAIQCHPVEVEDGQPKMASPLMQIEGFLSALTSANQDGRVILNRQGTVAKSSLKFLLLNPAVHFAKVLKECRAAIVAGGTMQPVSDFREQLLSCAIGADRVVEFSCGHVIPPDNILPIILCSGPSNQQLEFTYQKRDLPHMVSTMDEMGRILGNLCNVVPGGVVCFFPSYEYEKQVYTHWEGTGLLTRLATKKKIFQEPKKANQVEQVLAEYTKCIKRCSQVGGHVTGALLFSVVGGKMSEGINFSDDLGRCVVMVGMPYPNIKSPELQEKMAYLDKTMPRTAGQPPSKMLIENLCMKAVNQSIGRAIRHQKDFASILLVDHRYARPSVFNKLPQWIKERTEIKYTFGLAFAALRKVSTISCTYQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHD3</accession>
    <name>A0A452GHD3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">SAM domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="5">May regulate the disassembly of focal adhesions. Did not bind receptor-like tyrosine phosphatases type 2A.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the liprin family. Liprin-beta subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHD3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001160.1">
        <property type="protein sequence ID" value="ENSGAGP00000001028.1"/>
        <property type="gene ID" value="ENSGAGG00000000781.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048786">
        <property type="term" value="C:presynaptic active zone"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007528">
        <property type="term" value="P:neuromuscular junction development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd09563">
        <property type="entry name" value="SAM_liprin-beta1_2_repeat1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd09566">
        <property type="entry name" value="SAM_liprin-beta1_2_repeat2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd09569">
        <property type="entry name" value="SAM_liprin-beta1_2_repeat3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.50:FF:000005">
        <property type="entry name" value="Liprin-beta-1 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.50:FF:000007">
        <property type="entry name" value="Liprin-beta-1 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.50:FF:000017">
        <property type="entry name" value="Liprin-beta-1 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.50">
        <property type="entry name" value="Transcription Factor, Ets-1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058914">
        <property type="entry name" value="LIPB1/2_CC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037617">
        <property type="entry name" value="LIPB1/2_SAM_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037618">
        <property type="entry name" value="LIPB1/2_SAM_2nd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037619">
        <property type="entry name" value="LIPB1/2_SAM_3rd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029515">
        <property type="entry name" value="Liprin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001660">
        <property type="entry name" value="SAM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013761">
        <property type="entry name" value="SAM/pointed_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12587">
        <property type="entry name" value="LAR INTERACTING PROTEIN LIP -RELATED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12587:SF16">
        <property type="entry name" value="LIPRIN-BETA-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26022">
        <property type="entry name" value="CC_Liprin_beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00536">
        <property type="entry name" value="SAM_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07647">
        <property type="entry name" value="SAM_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00454">
        <property type="entry name" value="SAM"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47769">
        <property type="entry name" value="SAM/Pointed domain"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50105">
        <property type="entry name" value="SAM_DOMAIN"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 6" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="SAM" evidence="8">
        <location>
            <begin position="697"/>
            <end position="761"/>
        </location>
    </feature>
    <feature type="domain" description="SAM" evidence="8">
        <location>
            <begin position="769"/>
            <end position="832"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="453"/>
            <end position="636"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="649"/>
            <end position="684"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="99"/>
            <end position="373"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="481"/>
            <end position="495"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="512"/>
            <end position="524"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="599"/>
            <end position="609"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007547"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00060046"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50105"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001028.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1061" mass="118993" checksum="C965FB57FE5D0110" modified="2019-05-08" version="1">MMSDASDMLAAALEQMDGIIAGSKALEYSNGIFDCQSPTSPFMGSLRALHLVEDLRGLLEMMEADEKEGLRCQVPDSTAEALIEWLQSQMTNGHISGNGDIYQERLARLENDKESLVLQVSVLTDQVEAQGEKIRDLEFCLEEHREKLNATEEMLQQELLSRTSLETQKLNLMAEISNLKLKLTAVEKDRLDYEDRYGNTESLHNVIADLKQQMEKVEVELALFQEKRLQYGDQEGLMQEISELRLRVGEMDNERLQYEKKLKSTKSLMAKLSSMKIKVGQMQYEKQRMEQKCQILKDELASLKEKLEQKEAEVRKLQEKLVCKMKGEGVEITDRDIEVQKMKKAVESLMAANEEKDRKIEELRQSLNRYKKVQDMVILAQGKKGKEGECEEFLNSGSVSAVLLDMQSLTDQEKSPSPVPVISFPSHEEFNTTVDEENSLQIHTSILQVSIPAFPPAPESSEAATEKVNAQPKPEVASEVSEGTTPCSSVETQPCDSPLMSALQKSSSLGNLKKEASEVDKEPVQKPAETKAPMEGNKCGTLPPKSPSHSGTGDEDSFGTRKARSSFGRGFFKIKSNKRTASAPNLDRSRSASAPTLAETEKGSADHLDLAGLPPRPKETDSGQMSPSSPDSKKKARGIKKLFGRLKRSQSTTFNPDDMSEAEFKRGGTRATAGPRLGWSRDLGQSNSDLDMPFAKWTKEQVCNWLQGQGLGSYMNHGKHWILSGQTLLQASQQDLEKELGIKHPLHRKKLQLALQALGSEEENNHGKLDYNWVTRWLDDIGLPQYKTQFDEGRVDGRMLHYMSVDDLLSLKVVSVLHHLSIKRAIQVLRINNFEPNCLRRRPSDENNITPAEVTQWTNHRVMEWLRSVDLAEYAPNLRGSGVHGGLMVLEPRFNVETMAQLLNIPPNKTLLRRHLATHFNLLVGQDAQHQKREAMESPDYVLLTATAKVKPKKLAFSNFGSLRKKKQDDVEEYVCPMELGQASGSGSKKGFKAGLDIRVYDDDDLDRLEQMEDSEGTVRQIGAFSEGINNLTHMLKEDEMFKDFATCSPSASITDEDSNV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHD6</accession>
    <name>A0A452GHD6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">SAM domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="5">May regulate the disassembly of focal adhesions. Did not bind receptor-like tyrosine phosphatases type 2A.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the liprin family. Liprin-beta subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHD6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001140.1">
        <property type="protein sequence ID" value="ENSGAGP00000001009.1"/>
        <property type="gene ID" value="ENSGAGG00000000781.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048786">
        <property type="term" value="C:presynaptic active zone"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007528">
        <property type="term" value="P:neuromuscular junction development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd09563">
        <property type="entry name" value="SAM_liprin-beta1_2_repeat1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd09566">
        <property type="entry name" value="SAM_liprin-beta1_2_repeat2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd09569">
        <property type="entry name" value="SAM_liprin-beta1_2_repeat3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.50:FF:000005">
        <property type="entry name" value="Liprin-beta-1 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.50:FF:000007">
        <property type="entry name" value="Liprin-beta-1 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.50:FF:000017">
        <property type="entry name" value="Liprin-beta-1 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.50">
        <property type="entry name" value="Transcription Factor, Ets-1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058914">
        <property type="entry name" value="LIPB1/2_CC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037617">
        <property type="entry name" value="LIPB1/2_SAM_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037618">
        <property type="entry name" value="LIPB1/2_SAM_2nd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037619">
        <property type="entry name" value="LIPB1/2_SAM_3rd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029515">
        <property type="entry name" value="Liprin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001660">
        <property type="entry name" value="SAM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013761">
        <property type="entry name" value="SAM/pointed_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12587">
        <property type="entry name" value="LAR INTERACTING PROTEIN LIP -RELATED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12587:SF16">
        <property type="entry name" value="LIPRIN-BETA-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26022">
        <property type="entry name" value="CC_Liprin_beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00536">
        <property type="entry name" value="SAM_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07647">
        <property type="entry name" value="SAM_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00454">
        <property type="entry name" value="SAM"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47769">
        <property type="entry name" value="SAM/Pointed domain"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50105">
        <property type="entry name" value="SAM_DOMAIN"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 6" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="SAM" evidence="8">
        <location>
            <begin position="632"/>
            <end position="696"/>
        </location>
    </feature>
    <feature type="domain" description="SAM" evidence="8">
        <location>
            <begin position="704"/>
            <end position="767"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="399"/>
            <end position="510"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="526"/>
            <end position="571"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="584"/>
            <end position="619"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="99"/>
            <end position="319"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="427"/>
            <end position="441"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="458"/>
            <end position="470"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="534"/>
            <end position="544"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007547"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00060046"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50105"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001009.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="996" mass="111498" checksum="86801EA52E7CADB4" modified="2019-05-08" version="1">MMSDASDMLAAALEQMDGIIAGSKALEYSNGIFDCQSPTSPFMGSLRALHLVEDLRGLLEMMEADEKEGLRCQVPDSTAEALIEWLQSQMTNGHISGNGDIYQERLARLENDKESLVLQVSVLTDQVEAQGEKIRDLEFCLEEHREKLNATEEMLQQELLSRTSLETQKLNLMAEISNLKLKLTAVEKDRLDYEDRYGNTEGLMQEISELRLRVGEMDNERLQYEKKLKSTKDELASLKEKLEQKEAEVRKLQEKLVCKMKGEGVEITDRDENCKKKLKDKNIEVQKMKKAVESLMAANEEKDRKIEELRQSLNRYKKVQDMVILAQGKKGKEGECEEFLNSGSVSAVLLDMQSLTDQEKSPSPVPVISFPSHEEFNTTVDEENSLQIHTSILQVSIPAFPPAPESSEAATEKVNAQPKPEVASEVSEGTTPCSSVETQPCDSPLMSALQKSSSLGNLKKEASEVDKEPVQKPAETKAPMEGNKCGTLPPKSPSHSGTGDEDSFGTRKARSSFGRGFFKIKSNKRTASAPNLAETEKGSADHLDLAGLPPRPKETDSGQMSPSSPDSKKKARGIKKLFGRLKRSQSTTFNPDDMSEAEFKRGGTRATAGPRLGWSRDLGQSNSDLDMPFAKWTKEQVCNWLQGQGLGSYMNHGKHWILSGQTLLQASQQDLEKELGIKHPLHRKKLQLALQALGSEEENNHGKLDYNWVTRWLDDIGLPQYKTQFDEGRVDGRMLHYMSVDDLLSLKVVSVLHHLSIKRAIQVLRINNFEPNCLRRRPSDENNITPAEVTQWTNHRVMEWLRSVDLAEYAPNLRGSGVHGGLMVLEPRFNVETMAQLLNIPPNKTLLRRHLATHFNLLVGQDAQHQKREAMESPDYVLLTATAKVKPKKLAFSNFGSLRKKKQDDVEEYVCPMELGQASGSGSKKGFKAGLDIRVYDDDDLDRLEQMEDSEGTVRQIGAFSEGINNLTHMLKEDEMFKDFATCSPSASITDEDSNV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHD7</accession>
    <name>A0A452GHD7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">TEA domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 6">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHD7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001165.1">
        <property type="protein sequence ID" value="ENSGAGP00000001033.1"/>
        <property type="gene ID" value="ENSGAGG00000000816.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005667">
        <property type="term" value="C:transcription regulator complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048568">
        <property type="term" value="P:embryonic organ development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035329">
        <property type="term" value="P:hippo signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="2.70.50.80:FF:000001">
        <property type="entry name" value="Transcriptional enhancer factor TEF-1, putative"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.70.50.80">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.20.40">
        <property type="entry name" value="TEA/ATTS domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000818">
        <property type="entry name" value="TEA/ATTS_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038096">
        <property type="entry name" value="TEA/ATTS_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050937">
        <property type="entry name" value="TEC1_TEAD_TF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016361">
        <property type="entry name" value="TEF_metazoa"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041086">
        <property type="entry name" value="YBD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11834">
        <property type="entry name" value="TRANSCRIPTIONAL ENHANCER FACTOR TEF RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11834:SF2">
        <property type="entry name" value="TRANSCRIPTIONAL ENHANCER FACTOR TEF-3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01285">
        <property type="entry name" value="TEA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17725">
        <property type="entry name" value="YBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF002603">
        <property type="entry name" value="TEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00065">
        <property type="entry name" value="TEADOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00426">
        <property type="entry name" value="TEA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00554">
        <property type="entry name" value="TEA_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51088">
        <property type="entry name" value="TEA_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3 6" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="5 6" id="KW-0539">Nucleus</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 6" id="KW-0804">Transcription</keyword>
    <keyword evidence="2 6" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="TEA" evidence="8">
        <location>
            <begin position="1"/>
            <end position="39"/>
        </location>
    </feature>
    <feature type="DNA-binding region" description="TEA" evidence="7">
        <location>
            <begin position="1"/>
            <end position="39"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PIRNR" id="PIRNR002603"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00505"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51088"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001033.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="360" mass="40953" checksum="82C4590B8819D682" modified="2019-05-08" version="1">MYGRNELIARYIKLRTGKTRTRKQVSSHIQVLARRKAREIQAKLKDQAAKDKAMQSMATMSSAQIISATAFHSKMVLPGLPRPAYPAVSGFWQGALPGQAGSSHDVKPFSQQTYAVQPPLPLPGFESPAGLSPSPSAPPWQGRRVASSKLWMLEFSAFLEQQQDQDTYNKHLFVHIGQSSPSYSDPYLEAVDIRQIYDKFPEKKGGLKELFEKGPPNAFFLVKFWADLNTNIEDESRSFYGVSSQYESPENMIITCSTKVCSFGKQVVEKVETEYAHYENGHYSYRIHRSPLCEYMINFIHKLKHLPEKYMMNSVLENFTILQVVTNRDTQETLLCIAYVFEVSASEHGAQHHIYRLVKD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHE8</accession>
    <name>A0A452GHE8_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="11">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the tollip family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHE8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001121.1">
        <property type="protein sequence ID" value="ENSGAGP00000000993.1"/>
        <property type="gene ID" value="ENSGAGG00000000803.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043130">
        <property type="term" value="F:ubiquitin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031624">
        <property type="term" value="F:ubiquitin conjugating enzyme binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006914">
        <property type="term" value="P:autophagy"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006954">
        <property type="term" value="P:inflammatory response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045087">
        <property type="term" value="P:innate immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006511">
        <property type="term" value="P:ubiquitin-dependent protein catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd04016">
        <property type="entry name" value="C2_Tollip"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd14363">
        <property type="entry name" value="CUE_TOLIP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.8.10:FF:000036">
        <property type="entry name" value="Toll-interacting protein-like Protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000055">
        <property type="entry name" value="Toll-interacting protein-like Protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.8.10">
        <property type="entry name" value="DNA helicase RuvA subunit, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000008">
        <property type="entry name" value="C2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003892">
        <property type="entry name" value="CUE"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041799">
        <property type="entry name" value="TOLIP_CUE"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037301">
        <property type="entry name" value="Tollip_C2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009060">
        <property type="entry name" value="UBA-like_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16461">
        <property type="entry name" value="TOLL-INTERACTING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16461:SF5">
        <property type="entry name" value="TOLL-INTERACTING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00168">
        <property type="entry name" value="C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02845">
        <property type="entry name" value="CUE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00239">
        <property type="entry name" value="C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00546">
        <property type="entry name" value="CUE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49562">
        <property type="entry name" value="C2 domain (Calcium/lipid-binding domain, CaLB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46934">
        <property type="entry name" value="UBA-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50004">
        <property type="entry name" value="C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51140">
        <property type="entry name" value="CUE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0072">Autophagy</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0391">Immunity</keyword>
    <keyword evidence="8" id="KW-0395">Inflammatory response</keyword>
    <keyword evidence="4" id="KW-0399">Innate immunity</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="C2" evidence="9">
        <location>
            <begin position="35"/>
            <end position="152"/>
        </location>
    </feature>
    <feature type="domain" description="CUE" evidence="10">
        <location>
            <begin position="229"/>
            <end position="272"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009278"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022588"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022859"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023006"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023198"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS51140"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000993.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="274" mass="30399" checksum="CE74433DB494EC1C" modified="2019-05-08" version="1">MATTVSTQRGPVYIGELPQDFLRITPTQQQQQIQLDAQAAQQLQYGGPMGTVGRLSITVVQAKLAKNYGMTRMDPYCRVRLGYAVYETPTAHNGAKNPRWNKVIQCTVPPGVDSFYLEIFDERAFSMDDRIAWTHITIPESLKQGKVEDEWYSLSGRQGDDKEGMINLVMSYASIPAAMMMQPQPVVLMPTVYQQGVGYVPIAGMPAVCNPGMVPVAVPPPPVNPQHLCNEEDLKSIQDMFPNMDKEVIRSVLEAQRGNKDAAINSLLQIAEEL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHF0</accession>
    <name>A0A452GHF0_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="8">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell projection</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHF0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001159.1">
        <property type="protein sequence ID" value="ENSGAGP00000001027.1"/>
        <property type="gene ID" value="ENSGAGG00000000830.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030424">
        <property type="term" value="C:axon"/>
        <property type="evidence" value="ECO:0007669"/>
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        <property type="term" value="C:endocytic vesicle"/>
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        <property type="project" value="TreeGrafter"/>
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        <property type="term" value="C:plasma membrane"/>
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    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
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        <property type="project" value="InterPro"/>
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    <dbReference type="GO" id="GO:0043542">
        <property type="term" value="P:endothelial cell migration"/>
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        <property type="project" value="TreeGrafter"/>
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    <dbReference type="GO" id="GO:0007266">
        <property type="term" value="P:Rho protein signal transduction"/>
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        <property type="project" value="TreeGrafter"/>
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    <dbReference type="CDD" id="cd13244">
        <property type="entry name" value="PH_PLEKHG5_G6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd17068">
        <property type="entry name" value="RBD_PLEKHG5"/>
        <property type="match status" value="1"/>
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        <property type="match status" value="1"/>
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    <dbReference type="FunFam" id="2.30.29.30:FF:000141">
        <property type="entry name" value="Pleckstrin homology domain-containing family G member 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.900.10:FF:000017">
        <property type="entry name" value="pleckstrin homology domain-containing family G member 5 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.900.10">
        <property type="entry name" value="Dbl homology (DH) domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035899">
        <property type="entry name" value="DBL_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000219">
        <property type="entry name" value="DH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040181">
        <property type="entry name" value="PKHG5/7"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055251">
        <property type="entry name" value="SOS1_NGEF_PH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029071">
        <property type="entry name" value="Ubiquitin-like_domsf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13217:SF11">
        <property type="entry name" value="PLECKSTRIN HOMOLOGY DOMAIN-CONTAINING FAMILY G MEMBER 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13217">
        <property type="entry name" value="PLECKSTRIN HOMOLOGY DOMAIN-CONTAINING FAMILY G MEMBER 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00621">
        <property type="entry name" value="RhoGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22697">
        <property type="entry name" value="SOS1_NGEF_PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00233">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00325">
        <property type="entry name" value="RhoGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48065">
        <property type="entry name" value="DBL homology domain (DH-domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54236">
        <property type="entry name" value="Ubiquitin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50010">
        <property type="entry name" value="DH_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0966">Cell projection</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="DH" evidence="7">
        <location>
            <begin position="327"/>
            <end position="519"/>
        </location>
    </feature>
    <feature type="domain" description="PH" evidence="6">
        <location>
            <begin position="575"/>
            <end position="675"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="18"/>
            <end position="68"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="691"/>
            <end position="733"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="751"/>
            <end position="776"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="831"/>
            <end position="853"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="975"/>
            <end position="1000"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="721"/>
            <end position="733"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="5">
        <location>
            <begin position="761"/>
            <end position="776"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="5">
        <location>
            <begin position="979"/>
            <end position="994"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004316"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50010"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001027.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1053" mass="117135" checksum="F3AE1878A4E90F46" modified="2019-05-08" version="1">MHFDGHIRFDLPPQGSILARNVSTRSCPPRTSPASDVEEEEEGLVDGKGDKKSSALKLPKKKARRRHTDDPSKECFTLKFDFNVDIETEIVPAMKKKSLGEVLLPVFERKGIELAKVDIYLDQSNTPLSLSFEAYRFGGHYLRVKAKPGDELKVEQAVKDFKSLSLPIMRSSGSASTFLFTPAGERLDQSPFRRESMDILAPGRKRKNMNEFLGDSSIPGQDPLQHFSCSLPSNSIDTWKNRAASRFSGFFGSGANTGPFGREMDKMEQLESKLHSYSIFGLPKLPQQLRFDQDSWEEEEDDTSLCLEDSWQEITEGTEALTRRQCHQQEAVWELLHTEATYIRKLKVITDLFLCCLLNLQESGLLCEVEAERLFSNIQEIIQLHRALWSGVMAPILEKARKTKALLDPVDFLKGFKMFGSLFKPYIHYCMEEEGCMEYMRTLLRDNDLFRLYVTWAEKHKQCNRLKLSDMLVKPHQRLTKYPLLLKSVLKKTDDPCTRDAIITMISSVEQFINHVNWRMRQRQEQQKLVAILSRIDAYEVVDSSTEEVDKLLKEFLRLDLTAPIPGTSPEDTRQLLLEGSLKMKEGKDSKMDVYCFLFTDLFLITKPVKKAERTKVIRQPLLVDKIICRELKDPGSFLLIYLNELRSAVGAYTFQASGQSLCRSWIEALYNAQNLLQRLRLQERQRSQRQRLQGLEEGEDGESGASAASSPTILRRSRNSLDSQRCPSDGSTETISVVVVDASEELSFPDLEVGPFSSQSDETSISTTASSITPTRELLEAGGELAETRSSPSSNCLTLDSSGCRSASIDSAYGTLSPTSIQEFAEAQQLEPGAEDGGPLHTQRASSPKLRRRTPVQLLPCKVKALKSKSEASLLQLIPASPPLTQSKSLCNLFTALGQATFLAGPSQGLARREVPAGFQRTSRAGAGIGEARLDCSSLGLSICSSSSSSSSSELSEPEELMCAKGFAYPAESNVTSPAAPGQEGPQAGPEGASAPCRAPSTTVTECCQELPLAHRTLSDPQAAQHRKLTLAQLYRIRTTLLLNSTLTASEV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHF3</accession>
    <name>A0A452GHF3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">non-specific serine/threonine protein kinase</fullName>
            <ecNumber evidence="3">2.7.11.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="12">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452GHF3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001127.1">
        <property type="protein sequence ID" value="ENSGAGP00000000998.1"/>
        <property type="gene ID" value="ENSGAGG00000000802.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048471">
        <property type="term" value="C:perinuclear region of cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007059">
        <property type="term" value="P:chromosome segregation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035556">
        <property type="term" value="P:intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd14041">
        <property type="entry name" value="STKc_TLK2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000037">
        <property type="entry name" value="Serine/threonine-protein kinase tousled-like 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22974">
        <property type="entry name" value="MIXED LINEAGE PROTEIN KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22974:SF20">
        <property type="entry name" value="SERINE_THREONINE-PROTEIN KINASE TOUSLED-LIKE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8 13" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="9 14" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-0418">Kinase</keyword>
    <keyword evidence="6 13" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="16" id="KW-0732">Signal</keyword>
    <keyword evidence="5" id="KW-0808">Transferase</keyword>
    <feature type="signal peptide" evidence="16">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019300641" description="non-specific serine/threonine protein kinase" evidence="16">
        <location>
            <begin position="21"/>
            <end position="755"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="17">
        <location>
            <begin position="445"/>
            <end position="724"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="88"/>
            <end position="215"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="347"/>
            <end position="374"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="14">
        <location>
            <begin position="396"/>
            <end position="430"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="15">
        <location>
            <begin position="92"/>
            <end position="105"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="15">
        <location>
            <begin position="107"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="15">
        <location>
            <begin position="171"/>
            <end position="215"/>
        </location>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <position position="474"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012513"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00047899"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00048679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000998.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="755" mass="86606" checksum="FBAA7ACD49E401E1" modified="2019-05-08" version="1">MWQVAFYCITILNLYCMTEIICIQSLDTTTIFEHHQCHRKYVLSLFFALRNMVYFSPIAAEMMEELHSLDPRRQELLEARFTGVGVAKGPLNSESSNQSLCSVGSLSDKELETPEKKQNDQRNRKRKAEAYETSQGKGTPRGHKISDYFEFAGGSGPGTSPGRSVPPVARSSPQHSLSNPLPAQQGSPSSTGSGNTEHSCTSQKQISVQHKQTQSDLTMDKISALENNKNSDLEKKEGRIDDLLRANCDLRRQIDEQQKMLEKYKERLNRCVTMSKKLLIEKSKQEKMACRDKSMQDRLRLGHFTTVRHGASFTEQWTDGYAFQNLIKQQERINSQREEIERQRKMLAKRKPPAMGQTPPASNEQKQRKNKTNGAENETLTLAEYHEQEEIFKLRLGHLKKEEAEIQAELERLERVRNLHIRELKRIHNEDNSQFKDHPTLNDRYLLLHLLGRGGFSEVYKAFDLTEQRYVAVKIHQLNKNWRDEKKENYHKHACREYRIHKELDHPRIVKLYDYFSLDTDSFCTVLEYCEGNDLDFYLKQHKLMSEKEARSIIMQIVNALKYLNEIKPPIIHYDLKPGNILLVNGTACGEIKITDFGLSKIMDDDSYNSVDGMELTSQGAGTYWYLPPECFVVGKEPPKISNKVDVWSVGVIFYQCLYGRKPFGHNQSQQDILQENTILKATEVQFPPKPVVTPEAKAFIRRCLAYRKEDRIDVQQLACDPYLLPHIRKSVSTSSPAGAAIASTSGSSNNSSSN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHF4</accession>
    <name>A0A452GHF4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Transmembrane protein 107</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHF4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001164.1">
        <property type="protein sequence ID" value="ENSGAGP00000001032.1"/>
        <property type="gene ID" value="ENSGAGG00000000838.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036038">
        <property type="term" value="C:MKS complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1905515">
        <property type="term" value="P:non-motile cilium assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1904491">
        <property type="term" value="P:protein localization to ciliary transition zone"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029248">
        <property type="entry name" value="TMEM107"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR34341">
        <property type="entry name" value="TRANSMEMBRANE PROTEIN 107"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR34341:SF1">
        <property type="entry name" value="TRANSMEMBRANE PROTEIN 107"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14995">
        <property type="entry name" value="TMEM107"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0970">Cilium biogenesis/degradation</keyword>
    <keyword evidence="6 7" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 7" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="12"/>
            <end position="29"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="53"/>
            <end position="72"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="84"/>
            <end position="102"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="108"/>
            <end position="135"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00015652"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022794"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001032.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="140" mass="15546" checksum="6A3D6525E3154489" modified="2019-05-08" version="1">MVLVSGLVPARFLTLTAHLVIVITLFWSRDNNVLASLPLQYSQQQYQRQDVELVVALSVTLGLFAVELAGFLSGVSMFNNTQSLLSIGAHAAATVALLFFLFDQWDSSLYWWIFAFCSALPAAFEILLFVSVFGLKRKPL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHF5</accession>
    <name>A0A452GHF5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">C2H2-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">May be involved in transcriptional regulation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHF5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001128.1">
        <property type="protein sequence ID" value="ENSGAGP00000000999.1"/>
        <property type="gene ID" value="ENSGAGG00000000708.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000557">
        <property type="entry name" value="zinc finger and SCAN domain-containing protein 29"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000002">
        <property type="entry name" value="Zinc finger protein 1 homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000759">
        <property type="entry name" value="zinc finger protein 16"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000016">
        <property type="entry name" value="zinc finger protein 37 homolog"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000238">
        <property type="entry name" value="Zinc finger protein 485"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000367">
        <property type="entry name" value="Zinc finger protein 572"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002134">
        <property type="entry name" value="Zinc finger protein 616"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000431">
        <property type="entry name" value="zinc finger protein 629 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000912">
        <property type="entry name" value="Zinc finger protein 660"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000176">
        <property type="entry name" value="zinc finger protein 70"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001954">
        <property type="entry name" value="Zinc finger protein 787"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="12"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23235:SF178">
        <property type="entry name" value="C2H2-TYPE DOMAIN-CONTAINING PROTEIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23235">
        <property type="entry name" value="KRUEPPEL-LIKE TRANSCRIPTION FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="12"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="12"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="12"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="10" id="KW-0804">Transcription</keyword>
    <keyword evidence="8" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="7" id="KW-0862">Zinc</keyword>
    <keyword evidence="6 12" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="41"/>
            <end position="68"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="69"/>
            <end position="96"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="97"/>
            <end position="124"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="125"/>
            <end position="151"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="152"/>
            <end position="179"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="180"/>
            <end position="207"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="208"/>
            <end position="235"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="236"/>
            <end position="263"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="264"/>
            <end position="291"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="292"/>
            <end position="319"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="320"/>
            <end position="347"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="348"/>
            <end position="375"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00003767"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000999.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="380" mass="43715" checksum="2A865F725C8CA471" modified="2019-05-08" version="1">YFSVTGLGCEDVTLYDERREAHGILSDLITSGRINLEETRYRCHECGKSFSWSSTLVRHRRIHTGETPYTCTECGKSFSQSSNLITHQRIHTGEKPYTCTECGKSFNWSSTLNTHRRIHTGETPYTCTECGKSFSCSTLNTHRRIHTGKLPYTCTECEKSFSRSSDLIGHQRIHTREKPYTCSECGKSFSCSSDLIRHQRIHTGETPYTCTECGRSFNRSSHLIGHQRIHTGEKPYTCCECEKSFSQSSHLIGHQRIHTGETPYTCTECGRRFSWRSHLIGHQRIHTGETPYTCTECGKSFSQCSNLITHQRIHTGETPYTCTECGKSFNRSSTLNTHRRIHTREMPYTCSECGQSFSQRSRLIRHQKMHRRENCNKSLD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHF7</accession>
    <name>A0A452GHF7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Mitochondrial disaggregase</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="15">Suppressor of potassium transport defect 3</fullName>
        </alternativeName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="12">Functions as a regulatory ATPase and participates in secretion/protein trafficking process. Has ATP-dependent protein disaggregase activity and is required to maintain the solubility of key mitochondrial proteins. Involved in mitochondrial-mediated antiviral innate immunity, activates RIG-I-mediated signal transduction and production of IFNB1 and pro-inflammatory cytokine IL6. Plays a role in granulocyte differentiation.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>ATP + H2O = ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:13065"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
    </comment>
    <comment type="subunit">
        <text evidence="13">Homododecamer when substrate-bound; the homododecamer consists of 2 homohexamers stacked head-to-head via ANK repeat-mediated interactions. The active substrate-bound form is likely to exist in a dynamic equilibrium between homohexamers and homododecamers. Homotetradecamer in the unbound state which is remodeled upon substrate binding into the homododecamer. Interacts with PHB and PHB2. Interacts with MAVS; the interaction is enhanced by Sendai virus infection.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Mitochondrion intermembrane space</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the ClpA/ClpB family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHF7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001134.1">
        <property type="protein sequence ID" value="ENSGAGP00000001005.1"/>
        <property type="gene ID" value="ENSGAGG00000000810.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005758">
        <property type="term" value="C:mitochondrial intermembrane space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034605">
        <property type="term" value="P:cellular response to heat"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd19499">
        <property type="entry name" value="RecA-like_ClpB_Hsp104-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.8.60:FF:000040">
        <property type="entry name" value="caseinolytic peptidase B protein homolog isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000203">
        <property type="entry name" value="caseinolytic peptidase B protein homolog isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000641">
        <property type="entry name" value="caseinolytic peptidase B protein homolog isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.8.60">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003593">
        <property type="entry name" value="AAA+_ATPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003959">
        <property type="entry name" value="ATPase_AAA_core"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019489">
        <property type="entry name" value="Clp_ATPase_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001270">
        <property type="entry name" value="ClpA/B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050130">
        <property type="entry name" value="ClpA_ClpB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11638">
        <property type="entry name" value="ATP-DEPENDENT CLP PROTEASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11638:SF93">
        <property type="entry name" value="MITOCHONDRIAL DISAGGREGASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07724">
        <property type="entry name" value="AAA_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10431">
        <property type="entry name" value="ClpB_D2-small"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00300">
        <property type="entry name" value="CLPPROTEASEA"/>
    </dbReference>
    <dbReference type="SMART" id="SM00382">
        <property type="entry name" value="AAA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM01086">
        <property type="entry name" value="ClpB_D2-small"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0007">Acetylation</keyword>
    <keyword evidence="9 16" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="6" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="5" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="10" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="4" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="7" id="KW-0809">Transit peptide</keyword>
    <feature type="repeat" description="ANK" evidence="16">
        <location>
            <begin position="153"/>
            <end position="185"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="16">
        <location>
            <begin position="222"/>
            <end position="254"/>
        </location>
    </feature>
    <feature type="domain" description="AAA+ ATPase" evidence="18">
        <location>
            <begin position="330"/>
            <end position="459"/>
        </location>
    </feature>
    <feature type="domain" description="Clp ATPase C-terminal" evidence="19">
        <location>
            <begin position="513"/>
            <end position="602"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="602"/>
            <end position="623"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004569"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008675"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00049360"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00060073"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00066129"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00069063"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00083127"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="SMART" id="SM00382"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="SMART" id="SM01086"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001005.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="650" mass="73156" checksum="3DB2CF395ECE76AB" modified="2019-05-08" version="1">MGTGTGGGRVPGLQELGMDMGEFRTPGLQELGMDMGEFRVPWLQELGPCSTLARAPELPFLVAPGPELELQYPGPAVEWCVRLRPDPTGHQKKRESWTRRAVLPAFAAALVYCYSKSHASKEDALLEAARINNVSEVDRLLLEGTDVNARHKLGWTALMVAAINRNSSVVKILLAAGADPNLGDEFNSVYETAKEKGLHSLEVLVTREDDFNNRLNNRASFKGCTALHYAVLADDYVTVKLLLDGGASPLQRNEMGHTPLDYARDGEVMKLLKASETKFQEEQRKRELEERRRFPLEQRLKEHIIGQENAIATVGAAIRRKENGWYDEEHPLVFLFLGSSGIGKTELAKQTAKYMHKDAKKVAKFIGSPPGYIGHEEGGQLTKKLKQCPNAVVLFDEVDKAHPDVLTIMLQLFDEGRLTDGKGKTIDCKDAIFIMTSNVASDEIAQHALQLRQEALEMSRNRIAENLDDVQLTDKITISKQFKENVIRPILKAHFRRDEFLGRINEIVYFLPFCHSELIQLVSKELSFWAKRAKVRHNITLVWDREVMDVLADGYNLHYGARSIKHEVERRVVNQLAAAYEQDLLPRGCTLRITVDDSDKQLLKTKDSSSPEEEEKKAPTLRLEIVEKDSKSRKLDIQAPVSPDEISFML</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHG0</accession>
    <name>A0A452GHG0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Cullin family profile domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="pathway">
        <text evidence="1">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="similarity">
        <text evidence="2 5 6">Belongs to the cullin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GHG0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001169.1">
        <property type="protein sequence ID" value="ENSGAGP00000001037.1"/>
        <property type="gene ID" value="ENSGAGG00000000219.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031461">
        <property type="term" value="C:cullin-RING ubiquitin ligase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031625">
        <property type="term" value="F:ubiquitin protein ligase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006511">
        <property type="term" value="P:ubiquitin-dependent protein catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1310.10:FF:000012">
        <property type="entry name" value="Cullin 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1310.10:FF:000016">
        <property type="entry name" value="Cullin 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1310.10:FF:000018">
        <property type="entry name" value="Cullin 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.230.130:FF:000003">
        <property type="entry name" value="Cullin 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.1030.10:FF:000002">
        <property type="entry name" value="cullin homolog 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1310.10:FF:000022">
        <property type="entry name" value="Cullin-2 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1310.10">
        <property type="entry name" value="Cullin Repeats"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.1030.10">
        <property type="entry name" value="Ring Box Chain A, domain 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.10">
        <property type="entry name" value="Winged helix-like DNA-binding domain superfamily/Winged helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045093">
        <property type="entry name" value="Cullin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR059120">
        <property type="entry name" value="Cullin-like_AB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016158">
        <property type="entry name" value="Cullin_homology"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036317">
        <property type="entry name" value="Cullin_homology_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001373">
        <property type="entry name" value="Cullin_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016159">
        <property type="entry name" value="Cullin_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036388">
        <property type="entry name" value="WH-like_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11932">
        <property type="entry name" value="CULLIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00888">
        <property type="entry name" value="Cullin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26557">
        <property type="entry name" value="Cullin_AB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00182">
        <property type="entry name" value="CULLIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF75632">
        <property type="entry name" value="Cullin homology domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF74788">
        <property type="entry name" value="Cullin repeat-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50069">
        <property type="entry name" value="CULLIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0832">Ubl conjugation</keyword>
    <keyword evidence="3" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="Cullin family profile" evidence="7">
        <location>
            <begin position="415"/>
            <end position="647"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006019"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00330"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="RuleBase" id="RU003829"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50069"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001037.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="681" mass="79740" checksum="1687E719C8A35288" modified="2019-05-08" version="1">MSLKPRVVDFDETWNKLLTTIKAVVMLDYVERATWNDRFSDIYALCVAYPEPLGERLYTETKFFLENHVRHLHKRVLESEEQVLVMYHRYWEEYSKGADYMDCLYRYLNTQFIKKNKLTEADLQYGYGGVDISEPLMEIGELALDMWRKLMIEPLQGILIRMLLREIKNRSYTSTRQAECQLLHLCLNTAGSRQDHEFDRCGEDPNQKVIHGVINSFVHVEQYKKKFPLKFYQEIFECPFLNETGEYYKQEASNLLQESNCSQYMEKVLGRLKDEEMRCRKYLHPSSYGKVIHECQQRMVADHLQFLHAECHNIIRQEKRSDMANMYTLLRAVSSGLPHMIQELQNHIHDEGLRATSNLSQENMPTQFVESVLEVHSKFVQLINTVLNGDQHFMSALDKALTSVVNYREPKSICKAPELLAKYCDNLLKKSAKGMTENEVEDKLTSFITVFKYIDDKDVFQKFYARMLAKRLIHGLSMSMDSEEAMINKLKQACGYEFTSKLHRMYTDMSVSADLNNKFNNFIKNQDAIIDLGISFQIYVLQAGAWPLTQAPSSTFAIPQELEKSVQMFELFYSQHFSGRKLTWLHYLCTGEVKMNYLCKPYVAMVTTYQMAVLLAFNNSETVSYKELQDSTQMNEKELTKTIKSLLDVKMINHDLDKVCCNVTKRMCNISSHVFFLFIWF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHG3</accession>
    <name>A0A452GHG3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">non-specific serine/threonine protein kinase</fullName>
            <ecNumber evidence="3">2.7.11.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="12">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452GHG3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001142.1">
        <property type="protein sequence ID" value="ENSGAGP00000001011.1"/>
        <property type="gene ID" value="ENSGAGG00000000802.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048471">
        <property type="term" value="C:perinuclear region of cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007059">
        <property type="term" value="P:chromosome segregation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035556">
        <property type="term" value="P:intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd14041">
        <property type="entry name" value="STKc_TLK2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000037">
        <property type="entry name" value="Serine/threonine-protein kinase tousled-like 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22974">
        <property type="entry name" value="MIXED LINEAGE PROTEIN KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22974:SF20">
        <property type="entry name" value="SERINE_THREONINE-PROTEIN KINASE TOUSLED-LIKE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8 13" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="9 14" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-0418">Kinase</keyword>
    <keyword evidence="6 13" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="16" id="KW-0732">Signal</keyword>
    <keyword evidence="5" id="KW-0808">Transferase</keyword>
    <feature type="signal peptide" evidence="16">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019338396" description="non-specific serine/threonine protein kinase" evidence="16">
        <location>
            <begin position="21"/>
            <end position="723"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="17">
        <location>
            <begin position="413"/>
            <end position="692"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="88"/>
            <end position="183"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="315"/>
            <end position="342"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="14">
        <location>
            <begin position="364"/>
            <end position="398"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="15">
        <location>
            <begin position="92"/>
            <end position="105"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="15">
        <location>
            <begin position="107"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="15">
        <location>
            <begin position="152"/>
            <end position="183"/>
        </location>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <position position="442"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012513"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00047899"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00048679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001011.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="723" mass="83525" checksum="06388FA89B99B83B" modified="2019-05-08" version="1">MWQVAFYCITILNLYCMTEIICIQSLDTTTIFEHHQCHRKYVLSLFFALRNMVYFSPIAAEMMEELHSLDPRRQELLEARFTGVGVAKGPLNSESSNQSLCSVGSLSDKELETPEKKQNDQRNRKRKAEAYETSQGKGTPRGHKISDYFEAQQGSPSSTGSGNTEHSCTSQKQISVQHKQTQSDLTMDKISALENNKNSDLEKKEGRIDDLLRANCDLRRQIDEQQKMLEKYKERLNRCVTMSKKLLIEKSKQEKMACRDKSMQDRLRLGHFTTVRHGASFTEQWTDGYAFQNLIKQQERINSQREEIERQRKMLAKRKPPAMGQTPPASNEQKQRKNKTNGAENETLTLAEYHEQEEIFKLRLGHLKKEEAEIQAELERLERVRNLHIRELKRIHNEDNSQFKDHPTLNDRYLLLHLLGRGGFSEVYKAFDLTEQRYVAVKIHQLNKNWRDEKKENYHKHACREYRIHKELDHPRIVKLYDYFSLDTDSFCTVLEYCEGNDLDFYLKQHKLMSEKEARSIIMQIVNALKYLNEIKPPIIHYDLKPGNILLVNGTACGEIKITDFGLSKIMDDDSYNSVDGMELTSQGAGTYWYLPPECFVVGKEPPKISNKVDVWSVGVIFYQCLYGRKPFGHNQSQQDILQENTILKATEVQFPPKPVVTPEAKAFIRRCLAYRKEDRIDVQQLACDPYLLPHIRKSVSTSSPAGAAIASTSGSSNNSSSN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GHG4</accession>
    <name>A0A452GHG4_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000001143.1">
        <property type="protein sequence ID" value="ENSGAGP00000001012.1"/>
        <property type="gene ID" value="ENSGAGG00000000775.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060271">
        <property type="term" value="P:cilium assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR33487">
        <property type="entry name" value="CILIA- AND FLAGELLA-ASSOCIATED PROTEIN 54"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR33487:SF1">
        <property type="entry name" value="CILIA- AND FLAGELLA-ASSOCIATED PROTEIN 54"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="409"/>
            <end position="432"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="701"/>
            <end position="731"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="1">
        <location>
            <begin position="417"/>
            <end position="427"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="703"/>
            <end position="722"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000001012.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2344" mass="265216" checksum="10F0C1505D91A7B3" modified="2019-05-08" version="1">MNKIKKNKLSKAIFLMQKAILMFPKGSASSSQKHLLEEALALIQKVEAEQNALYSSLKQAMTSKKNRIPPPPLLLSRSHCSMTFKPAPFTSDVQVSWYSILGCVAAGSNLKVRLNNHHLPNAGEEIPADGKSLLEIKGLDPNEKYIFAVAAYSSDGKLIGDAIGETTKPILAYPPLSATTARAYLTQIAYQTDNYTLAKKAFSPLWDYFVLPPSTPSADPAVISASNNLTISQNRLCLDAVSQTSPILLHLFLRSIFIINEINVKEGALFCDSICSKEILYNGQVARLAECERMVVAIELSNWLNDANYALQSVVQCYGLLAPIIYHKIASVPVVQILIKCLTVLQEVPSSTLQRKQTGCYESIQHMIACSSFYTAQVLRSWKEYELALIIINYGKKLLDSSQTSHGQSKFSSIEEMQIDPETEDESPYSKKKSQVAAVEKAAENLAALESNLLKLTRPVMKFRKKPRFLEFFVQVLHKALNEEKFQRIVEWTGSVKDFLRRRNNHLLGIKGTLAKEVSELSVYEDAKRHTTAVVEFQKSVETSALRKPRKEVMSPFRGRESLRQYFLRHSSLGKSPEAQRKQRDEIEKVAFHTLVVLLNPIVSNYLKRKRFHQVCLEEMPWRSQMNIYLAITHFNLFKKRLEEQYNIGVRSLQHLNSYDVLDPEIFSLNNSGIIVVRGGTDDNIGTLSHLLIKRIQSGSSRKITDPDKSSERSTKFSEADTPRSQMTNDTELSITLSAREHERPMSNAILLEHFTKTFLHLRRAMVLAHRGGHWTLLQNTCRELWNHSQEALSIAKHSHSLQGLFPITRNFLISTIWLPFYLASDAILDMIIELQTSNSLKIFDPKGDFCVPSCVGGIADEDGGSNLTFCCPLDDISVVDLRWICGLVLRTIELMYQMKKWEALVHIAMHFNTVTHERYAEQVTPLLVYAQRQLLERIQQFRGPDSPQPHFIKYQGDNANKINCRNFIGKQLRIASTFNEVTVSGHHFVPEMPDTYTDGSQAQALVCVPLDVTDTLKCFRETLEKSKYHSRALRHSRKLLSLFLAHTQETVGRVPNQIFTNGKVEFSVGAELTFLPKPPDLSQEKYNFPSMVESRPLPESQLSVVVSSYEKTIEILQINNQRDLKVQALHELGNLHFYAGKKRSAFKYWCQALDETLKIADVLDNWQELEGFSKNVPDDFTDGSSKHFSEKCIYQAGIWGCLHAAVLAAKIAQYILTSSVRLRTKCCIFSAILFKTLFKVSIPHPTADCDFAQYEIHRLIPGIDLFSDRYRADVSTVVASLSFLIFELHCAKQNLIVLPLFTLYQYFVSEICRDPVKCIEGRILKIKVLTDLGFFAEAFYELCLLNHGEKIPWKVSAGYRLTGKMMSFQKFNASKPLLSNANLQVLEDIFNWSLPPAVVSMCDQQIMNKLTLAKMHFIICLSATINSIPEKVMKTIYSMDNTSLKKPSRIMESSIKINAEVTRHQEQSSIIVQLVRCKDELNMAMLKGILLTETEERLNSLVANIQNEYDGKISRCSAVELETVIEAKLQLAVISQQRHQTAFSAALAFSAIRLLQDANIFRKNVPHIQLNIQRTERRDSYVDSSQLPYNVIAREHMNIRLWLRCRMALVTALTAQIRGIGDMKENDMTDCSSLINEVIIEAEAFSDIETQAEVLVQAVILDLQERRPVTGIKLLLQDIINLLQEKPFISPMASLTLVKSMLLLADILTTQTDEDTEHHSSAIEQLNLLILAHKFTIKQIFTLGAPIKQCIEDPALTSPVVPLKNNYLPVINLLAKVKMRIAHALAQEVASTSKRENPLEWLPALSHLETALDLCRTSATRELDLEAEILFQKGKVERQIIKAGNNKSPRAAETLLDAIKVSQRNDQNFGLIRKSYLEIALLYFHLATYKEETSQTVKIEASKTKTTPKEHLASTERIPHITEVYRTQAWIAVRAATQVSEAMLASQLLIGKKGVKGHQVKAITEQNIPEFASMDLLASYKDFLSDGYEVRYKNLMASSIKNEEIIQSEKDQIESEGTECNISKDIQRKQTITWVHLIRYHNYLIRHNMSPLFAVSKSTPGSFSMEDALFTSVFDTGVVLRLAEMHFFLKKHLYIYSTCCLEDFPKQLQFLEKPGASLTAVPVKMSLEILGAPHKMSTVRIASTVSFCTDTDIQLDNRATIAPNKELCIQWYIPSLEKSTKDMETMVLLLYAYNTKYVKIVSIKTFNSANVFSGHLWIPLTRIISLHEKLSNVKQQIEILVQSSTSFPLQEQSERQRRMPSTKSKINLAKVELDEKIEEMLKQCLCEIKALLCVVPDLSSPLTEIPFDVTLQSIQNLEKMFDPANGCVIAGGSLFSWVISLLP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYA1</accession>
    <name>A0A452GYA1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Cellular communication network factor 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the CCN family.</text>
    </comment>
    <comment type="caution">
        <text evidence="6">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYA1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008288.1">
        <property type="protein sequence ID" value="ENSGAGP00000007163.1"/>
        <property type="gene ID" value="ENSGAGG00000005703.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031012">
        <property type="term" value="C:extracellular matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005921">
        <property type="term" value="C:gap junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008083">
        <property type="term" value="F:growth factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045597">
        <property type="term" value="P:positive regulation of cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051239">
        <property type="term" value="P:regulation of multicellular organismal process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.70.10">
        <property type="entry name" value="Complement Module, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.20.100.10">
        <property type="entry name" value="Thrombospondin type-1 (TSP1) repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050941">
        <property type="entry name" value="CCN"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006207">
        <property type="entry name" value="Cys_knot_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006208">
        <property type="entry name" value="Glyco_hormone_CN"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009030">
        <property type="entry name" value="Growth_fac_rcpt_cys_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000867">
        <property type="entry name" value="IGFBP-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012395">
        <property type="entry name" value="IGFBP_CNN"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043973">
        <property type="entry name" value="TSP1_CCN"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000884">
        <property type="entry name" value="TSP1_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036383">
        <property type="entry name" value="TSP1_rpt_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001007">
        <property type="entry name" value="VWF_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11348:SF37">
        <property type="entry name" value="CONNECTIVE TISSUE GROWTH FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11348">
        <property type="entry name" value="CONNECTIVE TISSUE GROWTH FACTOR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00007">
        <property type="entry name" value="Cys_knot"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00219">
        <property type="entry name" value="IGFBP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19035">
        <property type="entry name" value="TSP1_CCN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00093">
        <property type="entry name" value="VWC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF036495">
        <property type="entry name" value="IGFBP_rP_CNN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00041">
        <property type="entry name" value="CT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00121">
        <property type="entry name" value="IB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00209">
        <property type="entry name" value="TSP1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00214">
        <property type="entry name" value="VWC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57184">
        <property type="entry name" value="Growth factor receptor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF82895">
        <property type="entry name" value="TSP-1 type 1 repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01185">
        <property type="entry name" value="CTCK_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01225">
        <property type="entry name" value="CTCK_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51323">
        <property type="entry name" value="IGFBP_N_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50092">
        <property type="entry name" value="TSP1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01208">
        <property type="entry name" value="VWFC_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50184">
        <property type="entry name" value="VWFC_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0964">Secreted</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <feature type="domain" description="IGFBP N-terminal" evidence="9">
        <location>
            <begin position="8"/>
            <end position="79"/>
        </location>
    </feature>
    <feature type="domain" description="VWFC" evidence="8">
        <location>
            <begin position="82"/>
            <end position="148"/>
        </location>
    </feature>
    <feature type="domain" description="CTCK" evidence="7">
        <location>
            <begin position="232"/>
            <end position="306"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00039"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS01225"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50184"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51323"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007163.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="12">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="330" mass="36939" checksum="32E86D6BE28A8C15" modified="2025-10-08" version="2">MGQQIHAESQLCSYPCQCPSQPLQCPVGTSHVWDACGCCKVCARQLGELCSLQTPCDHHKGLHCDFSKIHRDVGICLVHEGATCDLLGKIYHNGESFQPTCKLQCICMDGAIGCIPLCADDLQLPSPACPNPLRVRLPAQCCEEWICQGGSQETALAVYRENPAPRSELNDLQENCLVQTTKWSACSRSCGMGISTRITNDNPQCRLEKESRLCMVRPCPVQPWHRQKGKKCTRTPKSRRGIRFEFSGCTSVRSYRPKFCGSCTDGRCCTPHTTSTAGVEFQCPEGDFFHRKLMFIKTCSCHHDCPRDNDIFLATYRRRMAGDHVKIERQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYA3</accession>
    <name>A0A452GYA3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="19">Neuropilin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="29 30">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="30" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="29" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="29" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="17">High affinity receptor for semaphorins 3C, 3F, VEGF-165 and VEGF-145 isoforms of VEGF, and the PLGF-2 isoform of PGF.</text>
    </comment>
    <comment type="subunit">
        <text evidence="18">Heterodimer with NRP1. Binds PLXNB1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 19">Belongs to the neuropilin family.</text>
    </comment>
    <comment type="caution">
        <text evidence="22">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYA3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008216.1">
        <property type="protein sequence ID" value="ENSGAGP00000007096.1"/>
        <property type="gene ID" value="ENSGAGG00000005700.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030424">
        <property type="term" value="C:axon"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098978">
        <property type="term" value="C:glutamatergic synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017154">
        <property type="term" value="F:semaphorin receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005021">
        <property type="term" value="F:vascular endothelial growth factor receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001525">
        <property type="term" value="P:angiogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007411">
        <property type="term" value="P:axon guidance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
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    </dbReference>
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        <property type="entry name" value="FA58C"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="CDD" id="cd06263">
        <property type="entry name" value="MAM"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Coagulation factor VIII"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Neuropilin"/>
        <property type="match status" value="1"/>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.260">
        <property type="entry name" value="Galactose-binding domain-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.290">
        <property type="entry name" value="Spermadhesin, CUB domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
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    <dbReference type="InterPro" id="IPR000859">
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    <dbReference type="InterPro" id="IPR000421">
        <property type="entry name" value="FA58C"/>
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    <dbReference type="InterPro" id="IPR008979">
        <property type="entry name" value="Galactose-bd-like_sf"/>
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        <property type="entry name" value="MAM_dom"/>
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        <property type="entry name" value="Neuropilin"/>
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    <dbReference type="InterPro" id="IPR050633">
        <property type="entry name" value="Neuropilin_MCO_CoagFactor"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035914">
        <property type="entry name" value="Sperma_CUB_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46806">
        <property type="entry name" value="F5/8 TYPE C DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46806:SF2">
        <property type="entry name" value="NEUROPILIN-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00431">
        <property type="entry name" value="CUB"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00754">
        <property type="entry name" value="F5_F8_type_C"/>
        <property type="match status" value="2"/>
    </dbReference>
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        <property type="entry name" value="MAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF036960">
        <property type="entry name" value="Neuropilin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00020">
        <property type="entry name" value="MAMDOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00042">
        <property type="entry name" value="CUB"/>
        <property type="match status" value="2"/>
    </dbReference>
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        <property type="entry name" value="FA58C"/>
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    </dbReference>
    <dbReference type="SMART" id="SM00137">
        <property type="entry name" value="MAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49785">
        <property type="entry name" value="Galactose-binding domain-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49854">
        <property type="entry name" value="Spermadhesin, CUB domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01180">
        <property type="entry name" value="CUB"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01285">
        <property type="entry name" value="FA58C_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01286">
        <property type="entry name" value="FA58C_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50022">
        <property type="entry name" value="FA58C_3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50060">
        <property type="entry name" value="MAM_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10 19 20" id="KW-0106">Calcium</keyword>
    <keyword evidence="3 19" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="9 19" id="KW-0221">Differentiation</keyword>
    <keyword evidence="14 21 22" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="16" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="4" id="KW-0358">Heparin-binding</keyword>
    <keyword evidence="13 19 24" id="KW-0472">Membrane</keyword>
    <keyword evidence="6 20" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11 19" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="15 19" id="KW-0675">Receptor</keyword>
    <keyword evidence="30" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="7 25" id="KW-0732">Signal</keyword>
    <keyword evidence="5 24" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="12 24" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="25">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019195100" description="Neuropilin" evidence="25">
        <location>
            <begin position="23"/>
            <end position="912"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="24">
        <location>
            <begin position="842"/>
            <end position="867"/>
        </location>
    </feature>
    <feature type="domain" description="CUB" evidence="26">
        <location>
            <begin position="28"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="domain" description="CUB" evidence="26">
        <location>
            <begin position="149"/>
            <end position="267"/>
        </location>
    </feature>
    <feature type="domain" description="F5/8 type C" evidence="27">
        <location>
            <begin position="277"/>
            <end position="427"/>
        </location>
    </feature>
    <feature type="domain" description="F5/8 type C" evidence="27">
        <location>
            <begin position="434"/>
            <end position="592"/>
        </location>
    </feature>
    <feature type="domain" description="MAM" evidence="28">
        <location>
            <begin position="642"/>
            <end position="806"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="23">
        <location>
            <begin position="297"/>
            <end position="318"/>
        </location>
    </feature>
    <feature type="binding site" evidence="20">
        <location>
            <position position="197"/>
        </location>
        <ligand>
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="20">
        <location>
            <position position="211"/>
        </location>
        <ligand>
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="20">
        <location>
            <position position="252"/>
        </location>
        <ligand>
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </ligand>
    </feature>
    <feature type="disulfide bond" evidence="21 22">
        <location>
            <begin position="28"/>
            <end position="55"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="21">
        <location>
            <begin position="83"/>
            <end position="105"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="21">
        <location>
            <begin position="149"/>
            <end position="175"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="21">
        <location>
            <begin position="208"/>
            <end position="230"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="21">
        <location>
            <begin position="277"/>
            <end position="427"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="21">
        <location>
            <begin position="434"/>
            <end position="592"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006078"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022674"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00053517"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00063815"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="PIRNR" id="PIRNR036960"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="PIRSR" id="PIRSR036960-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="PIRSR" id="PIRSR036960-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00059"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="26">
        <source>
            <dbReference type="PROSITE" id="PS01180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="27">
        <source>
            <dbReference type="PROSITE" id="PS50022"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="28">
        <source>
            <dbReference type="PROSITE" id="PS50060"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="29">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007096.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="30">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="912" mass="102090" checksum="880C764CFF4D86CE" modified="2019-05-08" version="1">MDTLPLVWGFVALYLAGSGVLGEAVQSCGGRLNSKDAGYITSPEYPQDYPSHQNCEWVIYAPEANQKIILNFNPHFEIEKHDCKYDFIEIRDGDSESADLLGKHCGNIAPPTIISSGSSLYIKFTSDYARQGAGFSLRYEIYKTGSEDCSRNFTSSNGTIESPGFPDKYPHNLDCTFTIMAKPKMEIILQFMTFDLEHDPLQVGEGDCKYDWLDIWDGISQVGPLIGRYCGTRTPSEIRSATGILSLTFHTDLAVAKDGFSARYYLVQQEIPENFQCNVPLGMESGRISNEQISASSTYSDGRWTPQQSRLNSDDNGWTPNLDSNKEFLQVDLRFLTMLTAIATQGAISKETQNGYYVRSYKLEISTNGEDWMMYRHGKNHKVFQANTDVTEVVLNKIHAPVLTRFVRIRPQVWHNGIALRLELYGCRITDSPCSNMLGMLSGLISDSQISASSIRGYDWSPSMARLVSSRSGWFPRIPQAQPGEEWLQVDLGVPKNVKGVVIQGARGGDSITMVEARSFVKKFRVAYSMNGKDWEYIQDTKTMQPKLFEGNMHYDIPEVRRFDPVPAQYIRVHPERWSPAGIGMRLEVLGCDWTDVKPTSETLKPTLKSEETTSPYPTDEEITECGDGCGEGDDAQVPVGFNCNFDLAEDLCGWSNDPATDYTWSFHSRSTWFSNSGPSTGSLPESKSYLQVQSSGRREDQRVRLVSPMIYLPRSAVCMVFRYQASGGSGIMLQVWREASQEKKLLWIISEDQGDEWKEGRIILPSYDMDYQIVFDGVIRKGHSGEIAIDDIRIGSDISLENCMEPITAFPGENFYIPGGTLLPGSEPTVDTVSVQPIPAYWYYVMAAGGAVLVLVSVALALVLHYHRFRYAAKKSDHAITYKTSHYTNGAPLAVEPTLTIKLEQDHSSRC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYB0</accession>
    <name>A0A452GYB0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">CCN family member 3</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="14">Cellular communication network factor 3</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="15">Protein NOV homolog</fullName>
        </alternativeName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell junction</location>
            <location evidence="2">Gap junction</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the CCN family.</text>
    </comment>
    <comment type="caution">
        <text evidence="16">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYB0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008223.1">
        <property type="protein sequence ID" value="ENSGAGP00000007101.1"/>
        <property type="gene ID" value="ENSGAGG00000005703.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031012">
        <property type="term" value="C:extracellular matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005921">
        <property type="term" value="C:gap junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008083">
        <property type="term" value="F:growth factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045597">
        <property type="term" value="P:positive regulation of cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051239">
        <property type="term" value="P:regulation of multicellular organismal process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000046">
        <property type="entry name" value="Cellular communication network factor 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.70.10">
        <property type="entry name" value="Complement Module, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.20.100.10">
        <property type="entry name" value="Thrombospondin type-1 (TSP1) repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050941">
        <property type="entry name" value="CCN"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006207">
        <property type="entry name" value="Cys_knot_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006208">
        <property type="entry name" value="Glyco_hormone_CN"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043973">
        <property type="entry name" value="TSP1_CCN"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000884">
        <property type="entry name" value="TSP1_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036383">
        <property type="entry name" value="TSP1_rpt_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001007">
        <property type="entry name" value="VWF_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11348:SF37">
        <property type="entry name" value="CONNECTIVE TISSUE GROWTH FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11348">
        <property type="entry name" value="CONNECTIVE TISSUE GROWTH FACTOR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00007">
        <property type="entry name" value="Cys_knot"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19035">
        <property type="entry name" value="TSP1_CCN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00093">
        <property type="entry name" value="VWC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00041">
        <property type="entry name" value="CT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00209">
        <property type="entry name" value="TSP1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00214">
        <property type="entry name" value="VWC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57603">
        <property type="entry name" value="FnI-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF82895">
        <property type="entry name" value="TSP-1 type 1 repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01185">
        <property type="entry name" value="CTCK_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01225">
        <property type="entry name" value="CTCK_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50092">
        <property type="entry name" value="TSP1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01208">
        <property type="entry name" value="VWFC_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50184">
        <property type="entry name" value="VWFC_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0965">Cell junction</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="8" id="KW-0303">Gap junction</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10" id="KW-0339">Growth factor</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0964">Secreted</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
    <feature type="domain" description="VWFC" evidence="18">
        <location>
            <begin position="34"/>
            <end position="100"/>
        </location>
    </feature>
    <feature type="domain" description="CTCK" evidence="17">
        <location>
            <begin position="222"/>
            <end position="296"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004610"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00008125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022868"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00039944"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00042352"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00077787"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00039"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS01225"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50184"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007101.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="313" mass="34753" checksum="33B7E7C828FE40F1" modified="2019-05-08" version="1">VGPGFQPRICEHLLEKISPPLPPPSSLPTVHEGATCDLLGKIYHNGESFQPTCKLQCICMDGAIGCIPLCADDLQLPSPACPNPLRVRLPAQCCEEWICQGGTKARWQGLVEGTGAAADSAAPVLWLQSVCGAFYDHKKVGKGSDIIQLPNQTPRSELNDLQENCLVQTTKWSACSRSCGMGISTRITNDNPQCRLEKESRLCMVRPCDFPREKIFKKGKKCTRTPKSRRGIRFEFSGCTSVRSYRPKFCGSCTDGRCCTPHTTSTAGVEFQCPEGDFFHRKLMFIKTCSCHHDCPRDNDIFLATYRIFNDHE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYB3</accession>
    <name>A0A452GYB3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">SMC hinge domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Chromosome</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000008165.1">
        <property type="protein sequence ID" value="ENSGAGP00000007049.1"/>
        <property type="gene ID" value="ENSGAGG00000005287.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005694">
        <property type="term" value="C:chromosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051276">
        <property type="term" value="P:chromosome organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006302">
        <property type="term" value="P:double-strand break repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1060.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.1620">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058611">
        <property type="entry name" value="Ig_SMCHD1_1st"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058612">
        <property type="entry name" value="Ig_SMCHD1_2nd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058613">
        <property type="entry name" value="Ig_SMCHD1_4th"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058614">
        <property type="entry name" value="Ig_SMCHD1_5th"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058615">
        <property type="entry name" value="Ig_SMCHD1_6th"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058617">
        <property type="entry name" value="Ig_SMCHD1_7th"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058616">
        <property type="entry name" value="Ig_SMCHD1_8th"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010935">
        <property type="entry name" value="SMC_hinge"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036277">
        <property type="entry name" value="SMC_hinge_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038892">
        <property type="entry name" value="SMCHD1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055109">
        <property type="entry name" value="SMCHD1_S5"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22640">
        <property type="entry name" value="STRUCTURAL MAINTENANCE OF CHROMOSOMES FLEXIBLE HINGE DOMAIN-CONTAINING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22640:SF2">
        <property type="entry name" value="STRUCTURAL MAINTENANCE OF CHROMOSOMES FLEXIBLE HINGE DOMAIN-CONTAINING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26194">
        <property type="entry name" value="Ig_SMCHD1_1st"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26195">
        <property type="entry name" value="Ig_SMCHD1_2nd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26196">
        <property type="entry name" value="Ig_SMCHD1_4th"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26197">
        <property type="entry name" value="Ig_SMCHD1_5th"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26198">
        <property type="entry name" value="Ig_SMCHD1_6th"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26201">
        <property type="entry name" value="Ig_SMCHD1_7th"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26199">
        <property type="entry name" value="Ig_SMCHD1_8th"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06470">
        <property type="entry name" value="SMC_hinge"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22899">
        <property type="entry name" value="SMCHD1_S5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00968">
        <property type="entry name" value="SMC_hinge"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF75553">
        <property type="entry name" value="Smc hinge domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0158">Chromosome</keyword>
    <keyword evidence="3" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="SMC hinge" evidence="5">
        <location>
            <begin position="1464"/>
            <end position="1591"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="1700"/>
            <end position="1735"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="3">
        <location>
            <begin position="1367"/>
            <end position="1422"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022454"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="SMART" id="SM00968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007049.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1735" mass="195168" checksum="353F1FA2EAAEFB3A" modified="2019-05-08" version="1">ATSSLPPPPPWQSAGLRHSGAGEGGGTCTLCSRSSLLLLECCKPADLLQAEGAGGRHLSAGSAGGQEVLGGIALVFPVCVFGLVKNIFLMNFRHIYHYYIHGPKGNETNATVKEAGPFNNIDIEILMFEKGKVPRMVNLREIKDDMQTLYINSAADSFEFKSHVEGEGIVEGIIRYHPFLYDKETYPDDPCFPSRLKDDDDDDDCFVVEKGARGKRPIFECFWNGRLIPYTSVEDFDWCAPPKKRGLAPIECYNRISGTLFTNDKFQVSTNKLTFMDLELKLKDKNTLFTRVFNGQEQRMKIDREFALWLKDCHEKYDKQIKFTLFKEISTRTDLASKRMQSPWATYSAIEWDGKTYRTGQLVKTIKTLPIFYGSILKFFLYGDHDGDVYATGGEVQLALEPQALYDEIKIIPISKLDRAMSEKVVKKYIEDEMARLPDRLSVTWPEGDELLPNEIKLAGTPIGALRIEILNKKGEAMQKLPGTSHGGSKKLLVELKVILHSPVGNKEIISHISQHGGKWPYWFKKMENIHKLGSYTLKLQVVLNESNADTYAGRPLPSKMFKFNIVEGKPEKFSVSLLDPPFRVGVPFNIPLDLQDEFGHPTQLASDIEPILEASGLTLQYEEINRGTNCVIKGVTAKGHVNNYLGKNFNLKVILPGLKEDSQILKIRLLPGPPRQLKVKPDSEMLKIENGTSFPFQVEVLDEAGNITAQPKLIVHCKFVGAPNLPVYAVDCSNAGTNILTGPVVHVQNIKKDQMLKARIEIPSCKDVPPVEKTIKLLPSSHVARLQIFSVEGQKAIQIKHQDEINWIAGDVMHNLIFQMYDEGEREIIITADLAEKIKVNWTPRINRDQMIQGLLPDVKVPTSVKDVRYCLITFHDDHVSLESAFTVRPLPDEPKHIKCKLKGPNTLQMGEELQNVMVTDQYGNQIQTLSASCINSLGVSGNGLDKSNLKTTWQESTQTMAVKGIRFYPGPPGSKELCFAWREFSDFLRVNLTSGPPVKLHLMDWPDLEEPIAVINGRELQKPLIVQLYDQWENPSPEPNVKISLVKASNLKIIPSSQQHKTDENGRANLGIISIHAPRGEHTLQLKASYNRNTLDCPIIKFNVLPDPEKPVCLNVKYDKNASFAAGSTFPDFMVSVISEDSNIIKNINPARVSMKIWEGLSSGNRIPANVTTLGCSKVKDGKDEGFFYFRDKMVPERVGTYSIQFAFAIDKTNILSSEQIIIEVVANDPVQLLPESLPATPVVSNVRTVASRTLIKDLCLHIKDEYNNRTGYDLVGKIIAAIKSSSEEDTEIPVFQGKVKTIEFPFNRGSAEIVNLVLAENSPGRDSTEYILEFEADLPTLAKPLEPYRLSFMFYNDFKKQQRMATLTRERDELSKSIAVYRNMFDTTEQLITEMKCQVQEAETKQSHLKNELKKHQIDIPQTSILQCVDSLIKQKMLEQEGVKKSPRRACTLSNYPKDNQDILGKIAHLAQIEDDRAAKVISWHMASDMDCVVTLTTAAARRIFDETQGRQQVLPLDSVYKKTLPDWNRTLPHLRNGRNHFRPIGNPVFARDLLTFPENVEHCQIVFGMLLGDTIIIDNLDAANHYRKEVVKITHCPTLLTREGDRIRSNGKFGGLQNKAPPMDKLRGMVFGAPMPKLYSVLSGQIDLLQQYRTAVVKLHSVNQDLHLQLQSLNTPEMQKRKQELAEQEKSLKLIEEKLGMTPSGKGTESLLPLEALDMSDSPTPPKRMRR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYB7</accession>
    <name>A0A452GYB7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Peptidase S1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the peptidase S1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYB7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008312.1">
        <property type="protein sequence ID" value="ENSGAGP00000007186.1"/>
        <property type="gene ID" value="ENSGAGG00000005766.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00190">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.10:FF:000077">
        <property type="entry name" value="Predicted protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.10.10">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009003">
        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043504">
        <property type="entry name" value="Peptidase_S1_PA_chymotrypsin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001314">
        <property type="entry name" value="Peptidase_S1A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001254">
        <property type="entry name" value="Trypsin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033116">
        <property type="entry name" value="TRYPSIN_SER"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24252">
        <property type="entry name" value="ACROSIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24252:SF26">
        <property type="entry name" value="TRANSMEMBRANE SERINE PROTEASE 9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00089">
        <property type="entry name" value="Trypsin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00722">
        <property type="entry name" value="CHYMOTRYPSIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00020">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50494">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50240">
        <property type="entry name" value="TRYPSIN_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00135">
        <property type="entry name" value="TRYPSIN_SER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="2" id="KW-0645">Protease</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0720">Serine protease</keyword>
    <feature type="domain" description="Peptidase S1" evidence="7">
        <location>
            <begin position="62"/>
            <end position="275"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="1"/>
            <end position="34"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="279"/>
            <end position="356"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="281"/>
            <end position="341"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007664"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007186.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="356" mass="37021" checksum="51B33D86F6F988C8" modified="2019-05-08" version="1">SSKPTSAPKDSPATPHQPGSRTTGVRAPSLPIPRGSESTNGLCFGFCLECGGRPGLSKPSKIVGGIEASRGEIPWQVSLKEGSRHFCGATVIGERWLLSAAHSGADGNAVKVSIKKVVQHPSYNPIILDFDVAVLELASALHFNKFIQPVCLPLAVQKFPVGKKCMISGWGNVKEGNITKPESLQKASVSIIDQKTCNVLYNFSLTDRMICAGFLEGRVDSCQGDSGGPLACEETPGLFYLAGVVSWGIGCAQAKRPGVYSRITKLKDWILDAISSKPNLSSGAGTSPSTAAARTSSTSSSPAKPSSEAATSPASQITTKPTATPRKTTTAAPKTTGAARPSQLSGNRLGLSLGCL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYB9</accession>
    <name>A0A452GYB9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">SMC hinge domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Chromosome</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYB9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008170.1">
        <property type="protein sequence ID" value="ENSGAGP00000007054.1"/>
        <property type="gene ID" value="ENSGAGG00000005287.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005694">
        <property type="term" value="C:chromosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051276">
        <property type="term" value="P:chromosome organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006302">
        <property type="term" value="P:double-strand break repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1060.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.1620">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058617">
        <property type="entry name" value="Ig_SMCHD1_7th"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058616">
        <property type="entry name" value="Ig_SMCHD1_8th"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010935">
        <property type="entry name" value="SMC_hinge"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036277">
        <property type="entry name" value="SMC_hinge_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038892">
        <property type="entry name" value="SMCHD1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22640">
        <property type="entry name" value="STRUCTURAL MAINTENANCE OF CHROMOSOMES FLEXIBLE HINGE DOMAIN-CONTAINING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22640:SF2">
        <property type="entry name" value="STRUCTURAL MAINTENANCE OF CHROMOSOMES FLEXIBLE HINGE DOMAIN-CONTAINING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26201">
        <property type="entry name" value="Ig_SMCHD1_7th"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26199">
        <property type="entry name" value="Ig_SMCHD1_8th"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06470">
        <property type="entry name" value="SMC_hinge"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00968">
        <property type="entry name" value="SMC_hinge"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF75553">
        <property type="entry name" value="Smc hinge domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0158">Chromosome</keyword>
    <keyword evidence="3" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="SMC hinge" evidence="5">
        <location>
            <begin position="373"/>
            <end position="500"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="609"/>
            <end position="644"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="3">
        <location>
            <begin position="276"/>
            <end position="331"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022454"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="SMART" id="SM00968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007054.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="644" mass="72813" checksum="6BF591D4D718C7FF" modified="2019-05-08" version="1">MKIHLNNCPIIKFNVLPDPEKPVCLNVKYDKNASFAAGSTFPDFMVSVISEDSNIIKNINPARVSMKIWEGLSSGNRIPANVTTLGCSKVKDGKDEGFFYFRDKMVPERVGTYSIQFAFAIDKTNILSSEQIIIEVVANDPVQLLPESLPATPVVSNVRTVASRTLIKDLCLHIKDEYNNRTGYDLVGKIIAAIKSSSEEDTEIPVFQGKVKTIEFPFNRGSAEIVNLVLAENSPGRDSTEYILEFEADLPTLAKPLEPYRLSFMFYNDFKKQQRMATLTRERDELSKSIAVYRNMFDTTEQLITEMKCQVQEAETKQSHLKNELKKHQIDIPQTSILQCVDSLIKQKMLEQEGVKKSPRRACTLSNYPKDNQDILGKIAHLAQIEDDRAAKVISWHMASDMDCVVTLTTAAARRIFDETQGRQQVLPLDSVYKKTLPDWNRTLPHLRNGRNHFRPIGNPVFARDLLTFPENVEHCQIVFGMLLGDTIIIDNLDAANHYRKEVVKITHCPTLLTREGDRIRSNGKFGGLQNKAPPMDKLRGMVFGAPMPKLYSVLSGQIDLLQQYRTAVVKLHSVNQDLHLQLQSLNTPEMQKRKQELAEQEKSLKLIEEKLGMTPSGKGTESLLPLEALDMSDSPTPPKRMRR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYC4</accession>
    <name>A0A452GYC4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">SMC hinge domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Chromosome</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYC4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008177.1">
        <property type="protein sequence ID" value="ENSGAGP00000007060.1"/>
        <property type="gene ID" value="ENSGAGG00000005287.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005694">
        <property type="term" value="C:chromosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051276">
        <property type="term" value="P:chromosome organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006302">
        <property type="term" value="P:double-strand break repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1060.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.1620">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058616">
        <property type="entry name" value="Ig_SMCHD1_8th"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010935">
        <property type="entry name" value="SMC_hinge"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036277">
        <property type="entry name" value="SMC_hinge_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038892">
        <property type="entry name" value="SMCHD1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22640">
        <property type="entry name" value="STRUCTURAL MAINTENANCE OF CHROMOSOMES FLEXIBLE HINGE DOMAIN-CONTAINING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22640:SF2">
        <property type="entry name" value="STRUCTURAL MAINTENANCE OF CHROMOSOMES FLEXIBLE HINGE DOMAIN-CONTAINING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26199">
        <property type="entry name" value="Ig_SMCHD1_8th"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06470">
        <property type="entry name" value="SMC_hinge"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00968">
        <property type="entry name" value="SMC_hinge"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF75553">
        <property type="entry name" value="Smc hinge domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0158">Chromosome</keyword>
    <keyword evidence="3" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="SMC hinge" evidence="5">
        <location>
            <begin position="140"/>
            <end position="267"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="376"/>
            <end position="411"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="3">
        <location>
            <begin position="43"/>
            <end position="98"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022454"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="SMART" id="SM00968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007060.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="411" mass="47073" checksum="3CA3C7177D432F44" modified="2019-05-08" version="1">IPGRDSTEYILEFEADLPTLAKPLEPYRLSFMFYNDFKKQQRMATLTRERDELSKSIAVYRNMFDTTEQLITEMKCQVQEAETKQSHLKNELKKHQIDIPQTSILQCVDSLIKQKMLEQEGVKKSPRRACTLSNYPKDNQDILGKIAHLAQIEDDRAAKVISWHMASDMDCVVTLTTAAARRIFDETQGRQQVLPLDSVYKKTLPDWNRTLPHLRNGRNHFRPIGNPVFARDLLTFPENVEHCQIVFGMLLGDTIIIDNLDAANHYRKEVVKITHCPTLLTREGDRIRSNGKFGGLQNKAPPMDKLRGMVFGAPMPKLYSVLSGQIDLLQQYRTAVVKLHSVNQDLHLQLQSLNTPEMQKRKQELAEQEKSLKLIEEKLGMTPSGKGTESLLPLEALDMSDSPTPPKRMRR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYC9</accession>
    <name>A0A452GYC9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Translation initiation factor eIF2B subunit delta</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="8">eIF2B GDP-GTP exchange factor subunit delta</fullName>
        </alternativeName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="6">Acts as a component of the translation initiation factor 2B (eIF2B) complex, which catalyzes the exchange of GDP for GTP on eukaryotic initiation factor 2 (eIF2) gamma subunit. Its guanine nucleotide exchange factor activity is repressed when bound to eIF2 complex phosphorylated on the alpha subunit, thereby limiting the amount of methionyl-initiator methionine tRNA available to the ribosome and consequently global translation is repressed.</text>
    </comment>
    <comment type="subunit">
        <text evidence="9">Component of the translation initiation factor 2B (eIF2B) complex which is a heterodecamer of two sets of five different subunits: alpha, beta, gamma, delta and epsilon. Subunits alpha, beta and delta comprise a regulatory subcomplex and subunits epsilon and gamma comprise a catalytic subcomplex. Within the complex, the hexameric regulatory complex resides at the center, with the two heterodimeric catalytic subcomplexes bound on opposite sides.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytosol</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 10">Belongs to the eIF-2B alpha/beta/delta subunits family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYC9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008290.1">
        <property type="protein sequence ID" value="ENSGAGP00000007165.1"/>
        <property type="gene ID" value="ENSGAGG00000005695.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005851">
        <property type="term" value="C:eukaryotic translation initiation factor 2B complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003743">
        <property type="term" value="F:translation initiation factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048513">
        <property type="term" value="P:animal organ development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007417">
        <property type="term" value="P:central nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10470:FF:000002">
        <property type="entry name" value="Probable translation initiation factor eIF-2B subunit delta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.10470">
        <property type="entry name" value="Translation initiation factor eif-2b, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000649">
        <property type="entry name" value="IF-2B-related"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042529">
        <property type="entry name" value="IF_2B-like_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037171">
        <property type="entry name" value="NagB/RpiA_transferase-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10233">
        <property type="entry name" value="TRANSLATION INITIATION FACTOR EIF-2B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10233:SF14">
        <property type="entry name" value="TRANSLATION INITIATION FACTOR EIF-2B SUBUNIT DELTA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01008">
        <property type="entry name" value="IF-2B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF100950">
        <property type="entry name" value="NagB/RpiA/CoA transferase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0396">Initiation factor</keyword>
    <keyword evidence="5" id="KW-0648">Protein biosynthesis</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="28"/>
            <end position="51"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004514"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022540"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022917"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00043898"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00044147"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00044356"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00046432"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="RuleBase" id="RU003814"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007165.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="362" mass="39164" checksum="00B2087540DB160C" modified="2019-05-08" version="1">MRHIAGRAGSRAQRAKRVLGAACRRGRSAGCQEGGRQLRWTGGPREPWDQRTLHRDACGRSTGAAGPASGRVPPAACRRAWGGEMARAVRAPWVGGMGVHTPTRSVEHGQLWGEPEGKMSVLSGPRVTGTASPQAKGKLQGSIDGYVREKIQLAAEAISKYAFEKISDGDVILVYGCSSLVSRTLCDAHANGRAFRVIVVDSRPRLEGRDTLRRLVRKGIRCTYVMITAISYMLPEVSKVLLGAHALLANGSVMSRVGTSQIALVSKAYNVPVLVCCETYKFCERVQTDSFVSNELDDPDDLILPGKGGAHLRGWKENSSLRLLNLVYDVTPPELVDLVITDLGMIPCTSVPVVLRVKNVEQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYE5</accession>
    <name>A0A452GYE5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">UBR-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000008265.1">
        <property type="protein sequence ID" value="ENSGAGP00000007142.1"/>
        <property type="gene ID" value="ENSGAGG00000005721.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043130">
        <property type="term" value="F:ubiquitin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034450">
        <property type="term" value="F:ubiquitin-ubiquitin ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090263">
        <property type="term" value="P:positive regulation of canonical Wnt signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000209">
        <property type="term" value="P:protein polyubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd14423">
        <property type="entry name" value="CUE_UBR5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd19675">
        <property type="entry name" value="UBR-box_UBR5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.30:FF:000007">
        <property type="entry name" value="E3 ubiquitin-protein ligase UBR5 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1750.10:FF:000016">
        <property type="entry name" value="E3 ubiquitin-protein ligase UBR5 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.8.10:FF:000009">
        <property type="entry name" value="Putative E3 ubiquitin-protein ligase UBR5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.8.10">
        <property type="entry name" value="DNA helicase RuvA subunit, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1750.10">
        <property type="entry name" value="Hect, E3 ligase catalytic domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.30">
        <property type="entry name" value="Regulator of chromosome condensation 1/beta-lactamase-inhibitor protein II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009091">
        <property type="entry name" value="RCC1/BLIP-II"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047503">
        <property type="entry name" value="UBR-box_UBR5"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024725">
        <property type="entry name" value="UBR5_UBA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003126">
        <property type="entry name" value="Znf_UBR"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46276">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE UBR5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46276:SF1">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE UBR5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11547">
        <property type="entry name" value="E3_UbLigase_EDD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00396">
        <property type="entry name" value="ZnF_UBR1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50985">
        <property type="entry name" value="RCC1/BLIP-II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51157">
        <property type="entry name" value="ZF_UBR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0862">Zinc</keyword>
    <keyword evidence="2" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="UBR-type" evidence="6">
        <location>
            <begin position="1121"/>
            <end position="1189"/>
        </location>
    </feature>
    <feature type="zinc finger region" description="UBR-type" evidence="4">
        <location>
            <begin position="1121"/>
            <end position="1189"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="77"/>
            <end position="175"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="225"/>
            <end position="249"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="271"/>
            <end position="296"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="523"/>
            <end position="591"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="943"/>
            <end position="976"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="996"/>
            <end position="1019"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="1243"/>
            <end position="1262"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="1459"/>
            <end position="1681"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="1803"/>
            <end position="1834"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="1928"/>
            <end position="1965"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="2059"/>
            <end position="2086"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="77"/>
            <end position="88"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="5">
        <location>
            <begin position="135"/>
            <end position="144"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="5">
        <location>
            <begin position="227"/>
            <end position="238"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="271"/>
            <end position="291"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="527"/>
            <end position="548"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="5">
        <location>
            <begin position="558"/>
            <end position="572"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="5">
        <location>
            <begin position="961"/>
            <end position="976"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="5">
        <location>
            <begin position="996"/>
            <end position="1018"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="5">
        <location>
            <begin position="1468"/>
            <end position="1481"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="1482"/>
            <end position="1497"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="5">
        <location>
            <begin position="1503"/>
            <end position="1518"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="5">
        <location>
            <begin position="1549"/>
            <end position="1558"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="1573"/>
            <end position="1582"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="5">
        <location>
            <begin position="1585"/>
            <end position="1601"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="5">
        <location>
            <begin position="1612"/>
            <end position="1625"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="5">
        <location>
            <begin position="1670"/>
            <end position="1681"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="1823"/>
            <end position="1834"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="5">
        <location>
            <begin position="1929"/>
            <end position="1942"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00508"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS51157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007142.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2293" mass="252079" checksum="C20FDAE05894544F" modified="2019-05-08" version="1">MTSIHFVVHPLPGTEDQLNDRLREVSEKLNKYNLNSHPPLNVLEQATIKQCVVGPNHAAFLLEDGRVCRIGFSVQPDRLELGKPDNNDGSKLSSGSGAGRTSRPGRTSDSPWFLSGSETLGRLAGNTLGSRWSSGVGGSGGGSSGRSSAGARDSRRQTRVIRTGRDRGSGLLGSQPQPVIPASVIPEELISQAQVVLQGKSRSVIIRELQRTNLDVNLAVNNLLSRDDEDGDDGDDTASESYLPGERDSELLRERESVLRLRERRWLDGASFDNERGSTSKEGEPNLDKKNTPVQSPVSLGEDLQWWPDKDGTKFISIGALYSELVAVSSKGELYQWKWSESEPYRNAQNPSLHHPRAMFLGLTNEKIVLLSANSIRATVATENNKVATWVDETLSSVASKLEHTAQTYSELQGERIVSLHCCALYTCAQLENNLYWWGVVPFSQRKKMLEKARAKNKKPKSSAGISSMPNITVGTQVCLRNNPLYHAGAVAFSISAGIPKVGVLMESVWNMNDSCRFQLRSPESLKNMEKASKTTEAKPESKQEPVKTEMGPPPSPASTCSDASSIASSASMPYKRRRSTPAPKEEEKVNEEQWSLREVVFVEDVKNVPVGKVLKVDGAYVAVKFPGTSSNANCQSSSSSDPDPSSLLQDCRLLRIDELQVVKTGGTPKVPDCFQRTPKKLCIPEKTEILAVNVDSKGVHAVLKTGNWVRYCIFDLATGKAEQENNFPTSSIAFLGQNERSVAIFTAGQESPIILRDGNGTIYPMAKDCMGGIRDPDWLDLPPISSLGMGVHSLTNLPANSTIKKKAAIIIMAVEKQTLMQHILRCDYEACRQYLMNLEQAVVLDQNLQMLQTFLSHRCDGNRNILHACVSVCFPTSNKETKEEEEAERSERNTFAERLSAVEAIANAISVVSSNGPGNRAGSSSSRSLRLREMMRRSLRAAGLGRHEAGASSSDHQDPVSPPIAPPSWVPDPPAMDPDGDIDFILAPAVGSLTTAATGTGPGPSTSTIPGPSTEPSVVESKDRKANAHFILKLLCDSVVLQPYLRELLSAKDARGMTPFMSAVSGRAYPAAITILETAQKIAKAEATSSEKEEDVFMGMVCPSGTNPDDSPLYVLCCNDTCSFTWTGAEHINQDIFECRTCGLLESLCCCTECARVCHKGHDCKLKRTSPTAYCDCWEKCKCKTLIAGQKSARLDLLYRLLTTTNLVTMPNSRGEHLLLFLVQTVARQTVEHCQYRPPRIREDRNRKTANPEDSDMPDHDLEPPRFAQLALERVLQDWNALKSMIMFGSQENKDPLSASSRIGHLLPEEQVYLNQQSGTIRLDCFTHCLIVKCTADILLLDTLLGTLVKELQNKYTPGRREEAIAVTMRFLRSVARVFVILSVEMASSKKKNNFIPQPIGKCKRVFQALLPYAVEELCNVAESLIIPVRMGIARPTAPFTLASTSIDAMQGSEELFSVEPLPPRPSSDQSSSSSQSQSSYIIRNPQQRRISQSQPVRGRDEEQDDIVSADVEEVEVVEGVAGEEDHHDEQEEHGEENAEAEGQHDEHDEDGSDMELDLLAAAETESDSESNHSNQDNASGRRSVVTAATAGSEAGASSVPAFFSEDDSQSNDSSDSDSSSSQSDDIEQETFMLDEPLERTTNSSHANGAAQAPRSMQWAVRNTQNQRTTSTAPSSTSAPAASSAGLIYIDPSNLRRSGTISTSAAAAAAALEASNASSYLTSASSLARAYSIVIRQISDLMGLIPKYNHLVYSQIPAAVKLTYQDAVNLQNYVEEKLIPTWNWMVSIMDSTEAQLRYGSALASAGDPGHPNHPLHASQNSTRRERMTAREEASLRTLEGRRRATLLSARQGMMSARGDFLNYALSLMRSHNDEHSDVLPVLDVCSLKHVAYVFQALIYWIKAMNQQTTLDTPQLERKRTRELLELGLDNEDSEHENDDDTNQSATLNDKDDDSLPAETGQNHPFFRRSDSMTFLGCIPPNPFEVPLAEAIPLADQPHLLQPNARKEDLFGRPSQGLYSSSANSGKCLMEVTVDRNCLEVLPTKMSYAANLKNVMSMQNRQKKEGEEQNVPTEESETSKPGPSAHDLAAQLKSSLLAEIGLTESEGPPLTSFRPQCSFMGMVISHDMLLGRWRLSLELFGRVFMEDVGAEPGSILTELGGFEVKESKFRREMEKLRNQQSRDLTLEVDRDRDLLIQQTMRQLNNHFGRRCATTPMAVHRVKVTFKDEPGEGSGVARSFYTAIAQAFLSNEKLPNLDCIQNANKGTHTSKYHREGIPWLPLIITVDTLFTLEY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYE7</accession>
    <name>A0A452GYE7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Protein FAM221A</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the FAM221 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GYE7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008356.1">
        <property type="protein sequence ID" value="ENSGAGP00000007226.1"/>
        <property type="gene ID" value="ENSGAGG00000005803.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026755">
        <property type="entry name" value="Fam221a/b"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31214:SF2">
        <property type="entry name" value="PROTEIN FAM221A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31214">
        <property type="entry name" value="PROTEIN FAM221A-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14753">
        <property type="entry name" value="FAM221"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="230"/>
            <end position="269"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="3">
        <location>
            <begin position="234"/>
            <end position="255"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00011026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00039630"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007226.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="299" mass="33262" checksum="C668DE14A4B578B0" modified="2019-05-08" version="1">MERLRVQAEGAAAVDEYVEYRRIVGDDDGGKLFTPEQYEEYKRKMLPIRLQNRLYVSWRSPTGMDCKLVGPETQCFCTHRYKQHKTDYEEIPKDRPICVPCRVSRCPCKSYHYVPLNGTQPIRCRCKHFADQHSAAPGFSCNSCSKCSGFHSCFTCGCGQPTYAHETVVETKQERLAQGKPVGQDVPYAAMGGLTGFSSLAEGYMRLDDSGVGAPSAEFLESPSSSMDHPFLKAFQGPSSSAQASSQLTGGSSAAKQVSNLRSSEKDDMAYFEKQYQERLKKEKAAKQKVKGLLLSKNP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYF8</accession>
    <name>A0A452GYF8_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="6">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GYF8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008279.1">
        <property type="protein sequence ID" value="ENSGAGP00000007154.1"/>
        <property type="gene ID" value="ENSGAGG00000005748.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="CDD" id="cd23767">
        <property type="entry name" value="IQCD"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000078">
        <property type="entry name" value="Inversin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000092">
        <property type="entry name" value="inversin isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000134">
        <property type="entry name" value="inversin isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000048">
        <property type="entry name" value="IQ_motif_EF-hand-BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051165">
        <property type="entry name" value="Multifunctional_ANK_Repeat"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24123">
        <property type="entry name" value="ANKYRIN REPEAT-CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24123:SF33">
        <property type="entry name" value="PROTEIN HOS4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00023">
        <property type="entry name" value="Ank"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13637">
        <property type="entry name" value="Ank_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00612">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01415">
        <property type="entry name" value="ANKYRIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="SMART" id="SM00015">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2 3" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="4" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0677">Repeat</keyword>
    <feature type="repeat" description="ANK" evidence="3">
        <location>
            <begin position="47"/>
            <end position="79"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="3">
        <location>
            <begin position="90"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="3">
        <location>
            <begin position="162"/>
            <end position="182"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="3">
        <location>
            <begin position="196"/>
            <end position="220"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="3">
        <location>
            <begin position="230"/>
            <end position="262"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="3">
        <location>
            <begin position="298"/>
            <end position="330"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="3">
        <location>
            <begin position="331"/>
            <end position="363"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="3">
        <location>
            <begin position="364"/>
            <end position="396"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="3">
        <location>
            <begin position="397"/>
            <end position="429"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="3">
        <location>
            <begin position="430"/>
            <end position="462"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="566"/>
            <end position="589"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="602"/>
            <end position="624"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="4">
        <location>
            <begin position="520"/>
            <end position="551"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="576"/>
            <end position="585"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="602"/>
            <end position="620"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007154.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1050" mass="116996" checksum="79D43E15D28E93A4" modified="2019-05-08" version="1">MNVSENLLVNGSSLASQVHAAAINGNKGALQKLIAGNPELKDKDDQFGRTPLMYCVLADRLDCAEALLKAGVGINKADHSRRTALHLAAQKQTALHWSAYYNNPEHVKLLIKHDSNIGIPDIEGKIPLHWAANHKDPSAVLTVKCILDAAPTESLLNWQDYEGRTPLHFAVADGNVAVVDVLTSYEGCNVTSYDNLFRTPLHWAALLGHAQIVHLLLERNKFGTIPSDSQGATPLHYAAQSNFAETVEVFLKHPSVKDDSDLEGRTSFMWAAGKGSDDVIRTMLNLKLNIDINMADKYAGTALHAAALSGHVSTVKLLLEHNAQVDATDVMKHTPLFRACEMGHKEVIQTLIKGGARVDLVDQDGHSPLHWAALGGNADVCQILIENKINPNMQDYAGRTPLQCAAYGGYINCMVVLLENNADPNIQDKEGRTALHWLCNNGYLDAIKLLLGFDAFPNHMENNEERYTPLDYALLGEHHEVIQFMLEHGALSIAAIQDIAAFKIQAIYKGYKVRKAFQDRKNLLMKHEQLRKDAVAKKREEENKRKEAELLKGMQSLVLNRVQVQPHPANREKSNRTPSTPQLPSKQIDVHSNRLFYSNISQNQAGKNNRGSPATSQNKTTPKEKCLSAELLGEDHKNKKELSRKHTKSKTACVHFHCGKENDGAKVEANHQVVAATEWYGEKHKEQTASAKGLAHGNEKHIPCCNTARAEEQLADQSQSIAGNRSYCEGTTGHLQKINCTDGNSRNSRQCETLPKGKPRMLKARNKELNSERCSPAGSSRPGSAKMRFVNIRNDSSHAVVQTNNVTNNELAKKTCPLLSIKAGTTRPVPSDPAAHAIFDDSLNLEKIDITGSKNADHQLCSHLPCQSMNIELIPLEIRMQIIEKERKRKELFRKKNYAATVIQRTWRSYQLRQKLVQLFSAERQCKEDEDKWRQETASFFIQVAWKKQLNHGPVKSVSSCKNLKSINKSNSAIKTSKQSILKQIYGRSQEGKVCHPARPSKHKLSDVHVISVSNLQYVHLLENEGKLKQFSYNMRPTTAAKSKTKRPEH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYG0</accession>
    <name>A0A452GYG0_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="5">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GYG0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008372.1">
        <property type="protein sequence ID" value="ENSGAGP00000007241.1"/>
        <property type="gene ID" value="ENSGAGG00000005814.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004672">
        <property type="term" value="F:protein kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd20430">
        <property type="entry name" value="Tudor_TDRD8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.140:FF:000018">
        <property type="entry name" value="Serine/threonine-protein kinase 31"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000518">
        <property type="entry name" value="serine/threonine-protein kinase 31 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.50.90">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052451">
        <property type="entry name" value="Ser/Thr_kinase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035437">
        <property type="entry name" value="SNase_OB-fold_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002999">
        <property type="entry name" value="Tudor"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047383">
        <property type="entry name" value="Tudor_TDRD8"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48008">
        <property type="entry name" value="LEUCINE-RICH REPEAT RECEPTOR-LIKE PROTEIN KINASE IMK3-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48008:SF6">
        <property type="entry name" value="LEUCINE-RICH REPEAT RECEPTOR-LIKE PROTEIN KINASE IMK3-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00567">
        <property type="entry name" value="TUDOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00333">
        <property type="entry name" value="TUDOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50199">
        <property type="entry name" value="Staphylococcal nuclease"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF63748">
        <property type="entry name" value="Tudor/PWWP/MBT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50304">
        <property type="entry name" value="TUDOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Tudor" evidence="4">
        <location>
            <begin position="55"/>
            <end position="114"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="3">
        <location>
            <begin position="650"/>
            <end position="913"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="937"/>
            <end position="959"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="280"/>
            <end position="328"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="937"/>
            <end position="950"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50304"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007241.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="959" mass="109336" checksum="DFC514057C6D1F55" modified="2019-05-08" version="1">MDENTGYSKVEDVVGCHVEDAVTFWAQNMSRNNDILKLSHALAEVCPQANSVFGNPDFTKIYGGCFSEDKCWYRCKVQQVISDEKCQVLYIDYGNSEMLNRSEIVEIPKNLQFPSIAKKYRLWGLQIPPDQDLNQFDQGRMFLCSLIFEKEIKMRHKATYQDGTIVAQAEYGKVDIGEEVAKKGFAEKCKSPINANNCDEKKIDVAQYQPWNAKVSAPAWENRPKPSVSNRLKGRFGDQIGTNMRNENHTGNHMPFVKEKLMARDFNIGSNISLAKIKQDQKLIEENEKLRNEKEVLQKENQILLQNYKELESTIAYLNHQVQEEKETSKETLKHLGKTLQTYVGTKMRSLAAQIEVLKNVRCSHMNVHFGDDLFRAVKEVTERCLIAPSSLEKLEKIWAEYNLAQEMIQLCRNVNESDILILKRNEVRQNLYSAVEEFILEVDELPLSERSTILQELSGSLEAIYGQVSEANDSEEAFQQFFEWKQTKLEEFNRVRSVTNASLQILATWFSGMKKLFDLTSEVSLKSEEVVGNVDEVLEKAESDICKELEISLVEQDEADKEIILNAYSKVIRKIRQEQHLITVVQSKYLASVESDDYSEYEITKIKEDITGLRNNLIQEIYNEQEEYETLTYLVQKWFPELPLLHPEAGILKYMNSGGLLAVSLERDLLDAEPLKELSTKHPLVCSEVQEQKVLLKGYAVDVSTEAKVIERTARYHRAWSEFKEKSGLLQLMFLFLCKSDPLVYLMVPYYPGASLGALQASTPLTSEEILKVMEGVARGLHTLHKANIVHGSLHTNNVFAVNRKQGIVGDFDFTKSVDQRASADSMVVGNLNLISPELRMGHLASPSSDMYAYGCLLFWLFAGDQEFKIKENGTPEVDELNMDDKVKSLLLNLICSNRMTAEQVLNDDCFLLPDVIPAPPQNEPGEKLKCEKAEEKGEVIINSDKKIDDEEEMDPSQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYG4</accession>
    <name>A0A452GYG4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Protein kinase domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GYG4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008378.1">
        <property type="protein sequence ID" value="ENSGAGP00000007246.1"/>
        <property type="gene ID" value="ENSGAGG00000005814.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004672">
        <property type="term" value="F:protein kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000518">
        <property type="entry name" value="serine/threonine-protein kinase 31 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.50.90">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052451">
        <property type="entry name" value="Ser/Thr_kinase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035437">
        <property type="entry name" value="SNase_OB-fold_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002999">
        <property type="entry name" value="Tudor"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48008">
        <property type="entry name" value="LEUCINE-RICH REPEAT RECEPTOR-LIKE PROTEIN KINASE IMK3-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48008:SF6">
        <property type="entry name" value="LEUCINE-RICH REPEAT RECEPTOR-LIKE PROTEIN KINASE IMK3-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00567">
        <property type="entry name" value="TUDOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Protein kinase" evidence="2">
        <location>
            <begin position="273"/>
            <end position="536"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="560"/>
            <end position="582"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="560"/>
            <end position="573"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007246.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="582" mass="65974" checksum="53336B4EF2E3FB8E" modified="2019-05-08" version="1">MDENTGYSKVEDVVGCHVEDAVTFWAQNMSRNNDILKLSHALAEVCPQANSVFGNPDFTKIYGGCFSEDKCWYRCKVQQVISDEKEEKETSKETLKHLGKTLQTYVGTKMRSLAAQIEVLKNVRCSHMNVHFGDDLFRAVKEVTERCLIAPSSLEKLEKIWAEYNLAQEMIQLCRNVNESDILILKRNEVRQNLYSAVEEFILEVDELPLSERSTILQSDDYSEYEITKIKEDITGLRNNLIQEIYNEQEEYETLTYLVQKWFPELPLLHPEAGILKYMNSGGLLAVSLERDLLDAEPLKELSTKHPLVCSEVQEQKVLLKGYAVDVSTEAKVIERTARYHRAWSEFKEKSGLLQLMFLFLCKSDPLVYLMVPYYPGASLGALQASTPLTSEEILKVMEGVARGLHTLHKANIVHGSLHTNNVFAVNRKQGIVGDFDFTKSVDQRASADSMVVGNLNLISPELRMGHLASPSSDMYAYGCLLFWLFAGDQEFKIKENGTPEVDELNMDDKVKSLLLNLICSNRMTAEQVLNDDCFLLPDVIPAPPQNEPGEKLKCEKAEEKGEVIINSDKKIDDEEEMDPSQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYG5</accession>
    <name>A0A452GYG5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Peroxisomal trans-2-enoyl-CoA reductase</fullName>
            <ecNumber evidence="13">1.3.1.38</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="11">Participates in chain elongation of fatty acids. Catalyzes the reduction of trans-2-enoyl-CoAs of varying chain lengths from 6:1 to 16:1, having maximum activity with 10:1 CoA. Has no 2,4-dienoyl-CoA reductase activity.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>(2E)-decenoyl-CoA + NADPH + H(+) = decanoyl-CoA + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:44960"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:61406"/>
            <dbReference type="ChEBI" id="CHEBI:61430"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="19">
            <dbReference type="Rhea" id="RHEA:44961"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>(2E)-dodecenoyl-CoA + NADPH + H(+) = dodecanoyl-CoA + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:44964"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57330"/>
            <dbReference type="ChEBI" id="CHEBI:57375"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="15">
            <dbReference type="Rhea" id="RHEA:44965"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>(2E)-hexenoyl-CoA + NADPH + H(+) = hexanoyl-CoA + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:44956"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:62077"/>
            <dbReference type="ChEBI" id="CHEBI:62620"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="17">
            <dbReference type="Rhea" id="RHEA:44957"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="20">
            <text>(2E)-octenoyl-CoA + NADPH + H(+) = octanoyl-CoA + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:44952"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57386"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:62242"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="20">
            <dbReference type="Rhea" id="RHEA:44953"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="16">
            <text>(2E)-tetradecenoyl-CoA + NADPH + H(+) = tetradecanoyl-CoA + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:44968"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57385"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:61405"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="16">
            <dbReference type="Rhea" id="RHEA:44969"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>a (2E)-enoyl-CoA + NADPH + H(+) = a 2,3-saturated acyl-CoA + NADP(+)</text>
            <dbReference type="Rhea" id="RHEA:33763"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:58856"/>
            <dbReference type="ChEBI" id="CHEBI:65111"/>
            <dbReference type="EC" id="1.3.1.38"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="18">
            <dbReference type="Rhea" id="RHEA:33764"/>
        </physiologicalReaction>
    </comment>
    <comment type="pathway">
        <text evidence="2">Lipid metabolism; fatty acid biosynthesis.</text>
    </comment>
    <comment type="subunit">
        <text evidence="12">Interacts with PEX5, probably required to target it into peroxisomes.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Peroxisome</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="1.3.1.38" evidence="13"/>
    <dbReference type="AlphaFoldDB" id="A0A452GYG5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008335.1">
        <property type="protein sequence ID" value="ENSGAGP00000007206.1"/>
        <property type="gene ID" value="ENSGAGG00000005788.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005777">
        <property type="term" value="C:peroxisome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019166">
        <property type="term" value="F:trans-2-enoyl-CoA reductase (NADPH) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006633">
        <property type="term" value="P:fatty acid biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033306">
        <property type="term" value="P:phytol metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05369">
        <property type="entry name" value="TER_DECR_SDR_a"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.720:FF:000335">
        <property type="entry name" value="Peroxisomal trans-2-enoyl-CoA reductase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.720">
        <property type="entry name" value="NAD(P)-binding Rossmann-like Domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036291">
        <property type="entry name" value="NAD(P)-bd_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052388">
        <property type="entry name" value="Peroxisomal_t2-enoyl-CoA_red"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002347">
        <property type="entry name" value="SDR_fam"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24317">
        <property type="entry name" value="PEROXISOMAL TRANS-2-ENOYL-COA REDUCTASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24317:SF7">
        <property type="entry name" value="PEROXISOMAL TRANS-2-ENOYL-COA REDUCTASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13561">
        <property type="entry name" value="adh_short_C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00081">
        <property type="entry name" value="GDHRDH"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00080">
        <property type="entry name" value="SDRFAMILY"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51735">
        <property type="entry name" value="NAD(P)-binding Rossmann-fold domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="10" id="KW-0275">Fatty acid biosynthesis</keyword>
    <keyword evidence="5" id="KW-0276">Fatty acid metabolism</keyword>
    <keyword evidence="3" id="KW-0444">Lipid biosynthesis</keyword>
    <keyword evidence="8" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="6" id="KW-0521">NADP</keyword>
    <keyword evidence="7" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="9" id="KW-0576">Peroxisome</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004275"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005194"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022516"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022832"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022857"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023140"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023160"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00037124"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00038622"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00038849"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00041063"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00047570"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00048686"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00049108"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00049251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00049386"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00049559"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007206.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="277" mass="29635" checksum="DAD06AEDBDBD2D9C" modified="2019-05-08" version="1">VWCLSLSPPPVFKTAAAPLTTGQDSAAAPPRPGNFKGCNVVIASRKLDRLQTAAKELAAKISSSNPVKVTPIQCNIRREEEVEALVKSTLDLHGRIDFLVNNGGGQFPSPAEAISSKGWNAVIETNLSGTFYCCKAVYNAWMREHGGAIVNIVADMWKGVPGMSHTGAARAAVDNLTKSLAIEWAHSGVRINSVAPGVIFSETAVANYKELGEEMFKNYILKIPAKRLGVPEEVSPTVCFLLSHAASFITGETVKVDAGQSLYSSPWEIPGKNRLMG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYG8</accession>
    <name>A0A452GYG8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">acetyl-CoA carboxylase</fullName>
            <ecNumber evidence="3">6.4.1.2</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>biotin</name>
            <dbReference type="ChEBI" id="CHEBI:57586"/>
        </cofactor>
    </comment>
    <comment type="pathway">
        <text evidence="2">Lipid metabolism; malonyl-CoA biosynthesis; malonyl-CoA from acetyl-CoA: step 1/1.</text>
    </comment>
    <dbReference type="EC" id="6.4.1.2" evidence="3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008287.1">
        <property type="protein sequence ID" value="ENSGAGP00000007162.1"/>
        <property type="gene ID" value="ENSGAGG00000004854.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00655">
        <property type="reaction ID" value="UER00711"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003989">
        <property type="term" value="F:acetyl-CoA carboxylase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006633">
        <property type="term" value="P:fatty acid biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:2001295">
        <property type="term" value="P:malonyl-CoA biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="CDD" id="cd06850">
        <property type="entry name" value="biotinyl_domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.460.10:FF:000001">
        <property type="entry name" value="Acetyl-CoA carboxylase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.50.100:FF:000005">
        <property type="entry name" value="Acetyl-CoA carboxylase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.470.20:FF:000005">
        <property type="entry name" value="Acetyl-CoA carboxylase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1770.10:FF:000001">
        <property type="entry name" value="acetyl-CoA carboxylase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1490.20:FF:000003">
        <property type="entry name" value="acetyl-CoA carboxylase isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.20:FF:000005">
        <property type="entry name" value="acetyl-CoA carboxylase isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.226.10:FF:000010">
        <property type="entry name" value="acetyl-CoA carboxylase isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.50.100">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.226.10">
        <property type="entry name" value="2-enoyl-CoA Hydratase, Chain A, domain 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1490.20">
        <property type="entry name" value="ATP-grasp fold, A domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.470.20">
        <property type="entry name" value="ATP-grasp fold, B domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.460.10">
        <property type="entry name" value="Biotin dependent carboxylase carboxyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1770.10">
        <property type="entry name" value="PreATP-grasp domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049076">
        <property type="entry name" value="ACCA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049074">
        <property type="entry name" value="ACCA_BT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034733">
        <property type="entry name" value="AcCoA_carboxyl_beta"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013537">
        <property type="entry name" value="AcCoA_COase_cen"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011761">
        <property type="entry name" value="ATP-grasp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013815">
        <property type="entry name" value="ATP_grasp_subdomain_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005481">
        <property type="entry name" value="BC-like_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011764">
        <property type="entry name" value="Biotin_carboxylation_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005482">
        <property type="entry name" value="Biotin_COase_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000089">
        <property type="entry name" value="Biotin_lipoyl"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029045">
        <property type="entry name" value="ClpP/crotonase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011763">
        <property type="entry name" value="COA_CT_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011762">
        <property type="entry name" value="COA_CT_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005479">
        <property type="entry name" value="CPAse_ATP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016185">
        <property type="entry name" value="PreATP-grasp_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011054">
        <property type="entry name" value="Rudment_hybrid_motif"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011053">
        <property type="entry name" value="Single_hybrid_motif"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45728:SF1">
        <property type="entry name" value="ACETYL-COA CARBOXYLASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45728">
        <property type="entry name" value="ACETYL-COA CARBOXYLASE, ISOFORM A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08326">
        <property type="entry name" value="ACC_central"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21385">
        <property type="entry name" value="ACCA_BT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02785">
        <property type="entry name" value="Biotin_carb_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00289">
        <property type="entry name" value="Biotin_carb_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00364">
        <property type="entry name" value="Biotin_lipoyl"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01039">
        <property type="entry name" value="Carboxyl_trans"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02786">
        <property type="entry name" value="CPSase_L_D2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00878">
        <property type="entry name" value="Biotin_carb_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52096">
        <property type="entry name" value="ClpP/crotonase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56059">
        <property type="entry name" value="Glutathione synthetase ATP-binding domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52440">
        <property type="entry name" value="PreATP-grasp domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51246">
        <property type="entry name" value="Rudiment single hybrid motif"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51230">
        <property type="entry name" value="Single hybrid motif"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50975">
        <property type="entry name" value="ATP_GRASP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50979">
        <property type="entry name" value="BC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50968">
        <property type="entry name" value="BIOTINYL_LIPOYL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50989">
        <property type="entry name" value="COA_CT_CTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50980">
        <property type="entry name" value="COA_CT_NTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00866">
        <property type="entry name" value="CPSASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00867">
        <property type="entry name" value="CPSASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8 13" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="11" id="KW-0092">Biotin</keyword>
    <keyword evidence="10" id="KW-0275">Fatty acid biosynthesis</keyword>
    <keyword evidence="7" id="KW-0276">Fatty acid metabolism</keyword>
    <keyword evidence="5" id="KW-0436">Ligase</keyword>
    <keyword evidence="4" id="KW-0444">Lipid biosynthesis</keyword>
    <keyword evidence="9" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="12" id="KW-0511">Multifunctional enzyme</keyword>
    <keyword evidence="6 13" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Biotin carboxylation" evidence="16">
        <location>
            <begin position="48"/>
            <end position="549"/>
        </location>
    </feature>
    <feature type="domain" description="ATP-grasp" evidence="15">
        <location>
            <begin position="206"/>
            <end position="397"/>
        </location>
    </feature>
    <feature type="domain" description="Lipoyl-binding" evidence="14">
        <location>
            <begin position="676"/>
            <end position="750"/>
        </location>
    </feature>
    <feature type="domain" description="CoA carboxyltransferase N-terminal" evidence="17">
        <location>
            <begin position="1469"/>
            <end position="1799"/>
        </location>
    </feature>
    <feature type="domain" description="CoA carboxyltransferase C-terminal" evidence="18">
        <location>
            <begin position="1803"/>
            <end position="2119"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001953"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004956"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00013058"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022516"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022598"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022832"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023160"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023267"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023268"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00409"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50975"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50979"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50980"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007162.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2165" mass="245224" checksum="9A72F41301C8C67D" modified="2019-05-08" version="1">GTPEGFHRRLTWLHLVKRGREQKKLELQRDFTVASPAEFVTRFGGNRIIEKVLIANNGIAAVKCMRSIRRWAYEMFRNERAIRFVVMVTPEDLKANAEYIKMADHYVPVPGGANNNNYANVELIVDISKRIPVQAVWAGWGHASENPRLPELLQKHGVAFLGPPSEAMWALGDKVASTIVAQTVQIPTLPWSGSGLMAEWTEDCEQRRTISVSLETYARGCVKDVDESLEAAERIGYPVMIKASEGGGGKGIRRAETAEEFTGCFRQVQSEAPGSPIFVMKLAQHARHLEVQVLADQYGNAVSLFGRDCSLQRRHQKIIEEAPVTIAAPSVFDFMEQCAVRLAKMVGYVSAGTVEYLYSEDGGFHFLELNPRLQVEHPCTEMIADVNLPAAQLQIAMGVPLHRIKDIRVLYGESPWADTPICFESPANIPVPRGHVIAARITSENPDEEFKPSSGTVQELNFRSSKNVWGYFSVAAAGGLHEFADSQFGHCFSWGENREEAISNMVVALKELSIRGDFRTTVEYLIKLLETESFQNNEIDTGWLDHLIAEKVQAEKPDTMLGVVCCALNVADALFRTCMTDFLHSLERGQVLPAASLLNIVEVELIYEGVKYALKVARQSLTTYVIIMNNTHIEIDVHRLNDGGVLLSYDGNSYTTYMKEEVDCYRITVGNKTCYFEKEKDPTMLRSPSAGKLLKYTVEDGGHIFIGHSFAEIEVMKIIFTLEVKESGHIRYIKRPGALLEAGCVIARLELDDPTKVHPAQLYTGGLPTQQMLPIGGLKLHQVFHSVLENLINVMNGYCLPEPYFSAKMKEWVCELMKTLRDPSLPLLELQEIMTNVSGRIPLSVEKSIRKVMAQYASNITSVLCQFPSQQIASILDSYAASLQRKADREVFFMNTQSIVQLVQRYRSGIRGYMKSVVLDLLRRYLQVETQFQQAHYDKCVINLREQYKLDMNPVLECIFSHAQVAKKNLLVIMLIDQLCGRDSTMTEELMAILTELTQLSKTEHSKVALRARQVLIASHLPSYELRHNQVESIFLSAIDMYGHQFCPEDLKKLILSETTIFDVLPTFFYHSNQLVRMAALEVYVRRGYIAYELNSLQHRQLSDGTCVVEFQFMLPSSHPNRMSIPISVSNPDLARHSTELFMDSGFSPLCQRMGAMVAFDRFEDFTRESPVQRGAXSCFADPPSESPLFSEAQSTLYEEGDNKTIQEEPIHILNTAIRWAGHLEDEELVPVFRTFAQSKVLFNFVNFFTFRARDEFAEDRIYRHLEPALAFQLELNRMRNFDLTAVPCANHRMHLYLGAAKVEEGNEVTDYRFFIRAIIRHSDLITKEASFEYLQNEGERLLLEAMDELEVAFSNTSVRTDCNHIFLNFVPTVIMDPSKIEEAVRSMVMRYGSRLWKLRVLQAEVKINIRLTPVGKAIPIRLFLTNESGYYLDISLYKEVNDPRTGNIMFHSYGDKQGLQHGMLINTPYVTKDLLQSKRFQAQSLGTTYVYDFPEMFKQALFKLWGSTEMYPKDVLTYTELVLDSQGQLVEMNRIPGGNEVGMVAFKMKLKTPEYPKGRDMIVISNDITCKIGSFGPEEDLLFLRASQLARSEGIPRIYIAANSGARIGFADEKKHMFQVAWVDPMDPYKGFKYLYLTPQDYTRINSVNSVHCEHVEEGGESRYVLTDIIGKDEGFGVENLRASGTIAGESSIAYDEIVTISMVTCRAIGIGAYLVRLGQRVIQVENSHIILTGASALNKVLGREVYTSNNQLGGVQIMHNNGVSHVTVPDDFEGIYTILQWLSYMPKDNRSPVPVTATSDPIEREIDFVPSKGPYDPRWMLAGRPHPIVKGTWQSGFFDHGTFMEIMQPWAQTVVVGRARLGGIPIGVVAVETRTVEMAIPADPANLDSEAKIVQQAGQVWFPDSAFKTAQAIRDFDRERLPLLIFANWRGFSGGMKDMYDQVLKFGAYIVDSLRQFKQPVLVYIPPHAELRGGSWVVIDPTINPVYMELYADKESRGGILEAEGTVEIKFRRKDLIKTMRRIDSVCARLVEQLGTPELLEEDHRELKKQLKAREELLLPIFHQVAVQFADLHDTPGRMHEKGVIADILEWKNARTFLYWRLRRLLLEEVVKTEILNADSELSNIHIQSMLRRWFMETEGAVKVSLHLENHWRASLFPCYGSV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYH0</accession>
    <name>A0A452GYH0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8 9">RNA cytidine acetyltransferase</fullName>
            <ecNumber evidence="9">2.3.1.-</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="9">18S rRNA cytosine acetyltransferase</fullName>
        </alternativeName>
    </protein>
    <gene>
        <name evidence="9" type="primary">NAT10</name>
    </gene>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">RNA cytidine acetyltransferase with specificity toward both 18S rRNA and tRNAs. Catalyzes the formation of N(4)-acetylcytidine (ac4C) in 18S rRNA. Required for early nucleolar cleavages of precursor rRNA at sites A0, A1 and A2 during 18S rRNA synthesis. Catalyzes the formation of ac4C in serine and leucine tRNAs. Requires the tRNA-binding adapter protein THUMPD1 for full tRNA acetyltransferase activity but not for 18S rRNA acetylation.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="9">
            <text>a cytidine in 18S rRNA + acetyl-CoA + ATP + H2O = an N(4)-acetylcytidine in 18S rRNA + ADP + phosphate + CoA + H(+)</text>
            <dbReference type="Rhea" id="RHEA:51424"/>
            <dbReference type="Rhea" id="RHEA-COMP:13575"/>
            <dbReference type="Rhea" id="RHEA-COMP:13576"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57288"/>
            <dbReference type="ChEBI" id="CHEBI:74900"/>
            <dbReference type="ChEBI" id="CHEBI:82748"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="9">
            <text>a cytidine in tRNA + acetyl-CoA + ATP + H2O = an N(4)-acetylcytidine in tRNA + ADP + phosphate + CoA + H(+)</text>
            <dbReference type="Rhea" id="RHEA:53876"/>
            <dbReference type="Rhea" id="RHEA-COMP:13670"/>
            <dbReference type="Rhea" id="RHEA-COMP:13671"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57288"/>
            <dbReference type="ChEBI" id="CHEBI:74900"/>
            <dbReference type="ChEBI" id="CHEBI:82748"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
    </comment>
    <comment type="subunit">
        <text evidence="9">Interacts with THUMPD1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 9">Nucleus</location>
            <location evidence="1 9">Nucleolus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="9">Belongs to the RNA cytidine acetyltransferase family. NAT10 subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.3.1.-" evidence="9"/>
    <dbReference type="AlphaFoldDB" id="A0A452GYH0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008383.1">
        <property type="protein sequence ID" value="ENSGAGP00000007251.1"/>
        <property type="gene ID" value="ENSGAGG00000002160.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030686">
        <property type="term" value="C:90S preribosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005730">
        <property type="term" value="C:nucleolus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:1990883">
        <property type="term" value="F:18S rRNA cytidine N-acetyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000049">
        <property type="term" value="F:tRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051392">
        <property type="term" value="F:tRNA cytidine N4-acetyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="RHEA"/>
    </dbReference>
    <dbReference type="GO" id="GO:1904812">
        <property type="term" value="P:rRNA acetylation involved in maturation of SSU-rRNA"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051391">
        <property type="term" value="P:tRNA acetylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="CDD" id="cd04301">
        <property type="entry name" value="NAT_SF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.630.30:FF:000019">
        <property type="entry name" value="RNA cytidine acetyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:002218">
        <property type="entry name" value="tRNA(Met) cytidine acetyltransferase TmcA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.630.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="HAMAP" id="MF_03211">
        <property type="entry name" value="RNA_acetyltr_Nat10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016181">
        <property type="entry name" value="Acyl_CoA_acyltransferase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000182">
        <property type="entry name" value="GNAT_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033688">
        <property type="entry name" value="NAT10"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007807">
        <property type="entry name" value="TcmA/NAT10_helicase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032672">
        <property type="entry name" value="TmcA/NAT10/Kre33"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027992">
        <property type="entry name" value="tRNA_bind_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10925">
        <property type="entry name" value="N-ACETYLTRANSFERASE 10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10925:SF5">
        <property type="entry name" value="RNA CYTIDINE ACETYLTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13718">
        <property type="entry name" value="GNAT_acetyltr_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05127">
        <property type="entry name" value="NAT10_TcmA_helicase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13725">
        <property type="entry name" value="tRNA_bind_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55729">
        <property type="entry name" value="Acyl-CoA N-acyltransferases (Nat)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51186">
        <property type="entry name" value="GNAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7 9" id="KW-0012">Acyltransferase</keyword>
    <keyword evidence="6 9" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="5 9" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="9" id="KW-0539">Nucleus</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 9" id="KW-0698">rRNA processing</keyword>
    <keyword evidence="3 9" id="KW-0808">Transferase</keyword>
    <keyword evidence="4 9" id="KW-0819">tRNA processing</keyword>
    <feature type="domain" description="N-acetyltransferase" evidence="10">
        <location>
            <begin position="400"/>
            <end position="595"/>
        </location>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <begin position="129"/>
            <end position="138"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="312"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <begin position="471"/>
            <end position="473"/>
        </location>
        <ligand>
            <name>acetyl-CoA</name>
            <dbReference type="ChEBI" id="CHEBI:57288"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <begin position="478"/>
            <end position="484"/>
        </location>
        <ligand>
            <name>acetyl-CoA</name>
            <dbReference type="ChEBI" id="CHEBI:57288"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="567"/>
        </location>
        <ligand>
            <name>acetyl-CoA</name>
            <dbReference type="ChEBI" id="CHEBI:57288"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004604"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022552"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022694"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023315"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00068357"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="HAMAP-Rule" id="MF_03211"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS51186"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007251.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="843" mass="94753" checksum="510ED826BF799B9D" modified="2019-05-08" version="1">MNSLKQLYTITMDVHSRYRTEAHQDIVGRFNERFILSLAACKNCIVIDDQLNILPISSHIANITPVLPQSLVSDLELKELKESLQDTQPVGVLVDSCKTLDQAKALLKFIEAISEKTLRSTVALTAARGRGKSAALGLAIAGAVAFGYSNIFVTSPSPDNLHTLFEFVFKGFDALQYEEHLDYEIIQSLNPEFNKAVVRVNVFKEHRQTIQYIHPADAVKLGQAELVVIDEAAAIPLPLVKNLLGPYLVFMASTINGYEGTGRSLSLKLIQQLRQQSAQAQASITAENKSTATAKLTSARTLHEVSLHESIRYAPGDPVEKWLNDLLCLDCLNITRIISGCPLPETCNLYYVNRDTLFCYHKASEIFLQRLMALYVASHYKNSPNDLQMLSDAPAHHLFCLLPPVPPTQNSLPEVLAVVQVCLEGEISRQSIMNSLSRGKKASGDLIPWTVSEQFQDPDFGSLSGGRVVRIAVHPDYQGMGYGSRALTLLQMYYEGKFPCLEEEVTPKPKGIATVSSEAVSLLEEVVTPRKDLPPLLLKLSERQAENLDYLGVSYGLTPRLLKFWKRAGFVPVYLRQTPNDLTGEHSCIMLKMLNEEESEQEHWLAAFWKDFRRRFLSLLSYQFSTFPPSLALNILQNKNIKQELQPPISRPELEAMFLPYDLKRLEMYSQNMVDYHLIMDTIPAVSRMYFLSQLGDTSLSAAQSALLLGIGLQHKSLDVLEKEIELPSSQLMGLFNRIIRKVVQLFSRIQEKAVEEQMAVTKEVVMEPTLKSLNEDLEEAAKEFQEKQKQEMGKLKDMDFTQYIIRGDDEEWNEVLNKAGQNTSIVNAAHGDFYTVCLESLV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYH4</accession>
    <name>A0A452GYH4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12 13">RNA cytidine acetyltransferase</fullName>
            <ecNumber evidence="13">2.3.1.-</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="13">18S rRNA cytosine acetyltransferase</fullName>
        </alternativeName>
    </protein>
    <gene>
        <name evidence="13" type="primary">NAT10</name>
    </gene>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">RNA cytidine acetyltransferase with specificity toward both 18S rRNA and tRNAs. Catalyzes the formation of N(4)-acetylcytidine (ac4C) in 18S rRNA. Required for early nucleolar cleavages of precursor rRNA at sites A0, A1 and A2 during 18S rRNA synthesis. Catalyzes the formation of ac4C in serine and leucine tRNAs. Requires the tRNA-binding adapter protein THUMPD1 for full tRNA acetyltransferase activity but not for 18S rRNA acetylation.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>a cytidine in 18S rRNA + acetyl-CoA + ATP + H2O = an N(4)-acetylcytidine in 18S rRNA + ADP + phosphate + CoA + H(+)</text>
            <dbReference type="Rhea" id="RHEA:51424"/>
            <dbReference type="Rhea" id="RHEA-COMP:13575"/>
            <dbReference type="Rhea" id="RHEA-COMP:13576"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57288"/>
            <dbReference type="ChEBI" id="CHEBI:74900"/>
            <dbReference type="ChEBI" id="CHEBI:82748"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>a cytidine in tRNA + acetyl-CoA + ATP + H2O = an N(4)-acetylcytidine in tRNA + ADP + phosphate + CoA + H(+)</text>
            <dbReference type="Rhea" id="RHEA:53876"/>
            <dbReference type="Rhea" id="RHEA-COMP:13670"/>
            <dbReference type="Rhea" id="RHEA-COMP:13671"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57288"/>
            <dbReference type="ChEBI" id="CHEBI:74900"/>
            <dbReference type="ChEBI" id="CHEBI:82748"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
    </comment>
    <comment type="subunit">
        <text evidence="13">Interacts with THUMPD1.</text>
    </comment>
    <comment type="subunit">
        <text evidence="11">Part of the small subunit (SSU) processome, composed of more than 70 proteins and the RNA chaperone small nucleolar RNA (snoRNA) U3. Interacts with THUMPD1. Interacts with SUN1 (via N-terminus). Interacts with TERT.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 13">Nucleus</location>
            <location evidence="1 13">Nucleolus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="13">Belongs to the RNA cytidine acetyltransferase family. NAT10 subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.3.1.-" evidence="13"/>
    <dbReference type="AlphaFoldDB" id="A0A452GYH4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008294.1">
        <property type="protein sequence ID" value="ENSGAGP00000007168.1"/>
        <property type="gene ID" value="ENSGAGG00000002160.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030686">
        <property type="term" value="C:90S preribosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005730">
        <property type="term" value="C:nucleolus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:1990883">
        <property type="term" value="F:18S rRNA cytidine N-acetyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000049">
        <property type="term" value="F:tRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051392">
        <property type="term" value="F:tRNA cytidine N4-acetyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="RHEA"/>
    </dbReference>
    <dbReference type="GO" id="GO:1904812">
        <property type="term" value="P:rRNA acetylation involved in maturation of SSU-rRNA"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051391">
        <property type="term" value="P:tRNA acetylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="CDD" id="cd04301">
        <property type="entry name" value="NAT_SF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.11040:FF:000006">
        <property type="entry name" value="RNA cytidine acetyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.630.30:FF:000019">
        <property type="entry name" value="RNA cytidine acetyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:002218">
        <property type="entry name" value="tRNA(Met) cytidine acetyltransferase TmcA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.11040">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.630.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="HAMAP" id="MF_03211">
        <property type="entry name" value="RNA_acetyltr_Nat10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016181">
        <property type="entry name" value="Acyl_CoA_acyltransferase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000182">
        <property type="entry name" value="GNAT_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033688">
        <property type="entry name" value="NAT10"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007807">
        <property type="entry name" value="TcmA/NAT10_helicase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032672">
        <property type="entry name" value="TmcA/NAT10/Kre33"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013562">
        <property type="entry name" value="TmcA/NAT10_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027992">
        <property type="entry name" value="tRNA_bind_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10925">
        <property type="entry name" value="N-ACETYLTRANSFERASE 10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10925:SF5">
        <property type="entry name" value="RNA CYTIDINE ACETYLTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13718">
        <property type="entry name" value="GNAT_acetyltr_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05127">
        <property type="entry name" value="NAT10_TcmA_helicase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08351">
        <property type="entry name" value="TmcA_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13725">
        <property type="entry name" value="tRNA_bind_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55729">
        <property type="entry name" value="Acyl-CoA N-acyltransferases (Nat)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51186">
        <property type="entry name" value="GNAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0007">Acetylation</keyword>
    <keyword evidence="10 13" id="KW-0012">Acyltransferase</keyword>
    <keyword evidence="7 13" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="6 13" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="9 13" id="KW-0539">Nucleus</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 13" id="KW-0698">rRNA processing</keyword>
    <keyword evidence="4 13" id="KW-0808">Transferase</keyword>
    <keyword evidence="5 13" id="KW-0819">tRNA processing</keyword>
    <feature type="domain" description="N-acetyltransferase" evidence="15">
        <location>
            <begin position="515"/>
            <end position="710"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="869"/>
            <end position="905"/>
        </location>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <begin position="244"/>
            <end position="253"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <position position="427"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <begin position="586"/>
            <end position="588"/>
        </location>
        <ligand>
            <name>acetyl-CoA</name>
            <dbReference type="ChEBI" id="CHEBI:57288"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <begin position="593"/>
            <end position="599"/>
        </location>
        <ligand>
            <name>acetyl-CoA</name>
            <dbReference type="ChEBI" id="CHEBI:57288"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <position position="682"/>
        </location>
        <ligand>
            <name>acetyl-CoA</name>
            <dbReference type="ChEBI" id="CHEBI:57288"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004604"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022552"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022694"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023315"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00065380"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00068357"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="HAMAP-Rule" id="MF_03211"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS51186"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007168.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="905" mass="102380" checksum="ABA1D9FAFD0BAA7F" modified="2019-05-08" version="1">MLSKAAVKARPSVLWCYKKELGFSSHRKKRMRQLQKKIKSGTLNLNQDDPFELFVAATNIRYCYYKETHKILGNTFGMCVLQDFEALTPNLLARTVETVEGGGIVVILLRTMNSLKQLYTITMDVHSRYRTEAHQDIVGRFNERFILSLAACKNCIVIDDQLNILPISSHIANITPVLPQSLDNIRPQDLELKELKESLQDTQPVGVLVDSCKTLDQAKALLKFIEAISEKTLRSTVALTAARGRGKSAALGLAIAGAVAFGYSNIFVTSPSPDNLHTLFEFVFKGFDALQYEEHLDYEIIQSLNPEFNKAVVRVNVFKEHRQTIQYIHPADAVKLGQAELVVIDEAAAIPLPLVKNLLGPYLVFMASTINGYEGTGRSLSLKLIQQLRQQSAQAQASITAENKSTATAKLTSARTLHEVSLHESIRYAPGDPVEKWLNDLLCLDCLNITRIISGCPLPETCNLYYVNRDTLFCYHKASEIFLQRLMALYVASHYKNSPNDLQMLSDAPAHHLFCLLPPVPPTQNSLPEVLAVVQVCLEGEISRQSIMNSLSRGKKASGDLIPWTVSEQFQDPDFGSLSGGRVVRIAVHPDYQGMGYGSRALTLLQMYYEGKFPCLEEEVTPKPKGIATVSSEAVSLLEEVVTPRKDLPPLLLKLSERQAENLDYLGVSYGLTPRLLKFWKRAGFVPVYLRQTPNDLTGEHSCIMLKMLNEEESEQEHWLAAFWKDFRRRFLSLLSYQFSTFPPSLALNILQNKNIKQELQPPISRPELEAMFLPYDLKRLEMYSQNMVDYHLIMDTIPAVSRMYFLSQLGDTSLSAAQSEEAAKEFQEKQKQEMGKLKDMDFTQYIIRGDDEEWNEVLNKAGQNTSIVSLKSEKKRKLDMEKGPKQHKKFKNKKDPKQKQKWKK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2025-10-08" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYH5</accession>
    <name>A0A452GYH5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Protein TOPAZ1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="7">Testis- and ovary-specific PAZ domain-containing protein 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">Important for normal spermatogenesis and male fertility. Specifically required for progression to the post-meiotic stages of spermatocyte development. Seems to be necessary for normal expression levels of a number of testis-expressed gene transcripts, although its role in this process is unclear.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytosol</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000008296.1">
        <property type="protein sequence ID" value="ENSGAGP00000007170.1"/>
        <property type="gene ID" value="ENSGAGG00000005743.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000008301.1">
        <property type="protein sequence ID" value="ENSGAGP00000007175.1"/>
        <property type="gene ID" value="ENSGAGG00000005743.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048137">
        <property type="term" value="P:spermatocyte division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038952">
        <property type="entry name" value="TOPAZ1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029435">
        <property type="entry name" value="TOPAZ1_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR35671">
        <property type="entry name" value="PROTEIN TOPAZ1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR35671:SF1">
        <property type="entry name" value="PROTEIN TOPAZ1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14669">
        <property type="entry name" value="Asp_Glu_race_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="5" id="KW-0221">Differentiation</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0744">Spermatogenesis</keyword>
    <feature type="domain" description="Protein TOPAZ1" evidence="8">
        <location>
            <begin position="1040"/>
            <end position="1215"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00002132"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004514"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00016464"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022871"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00031943"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF14669"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007175.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1445" mass="164335" checksum="EE7CFDE781FA365F" modified="2019-05-08" version="1">MGQSDMNSKLTVMQMSTQKSIPFDKEGICQNRKNCQNENSKCLRERKWNIDRKPRKRMKLFEKSSVQNVLPVMDNKYCEYELHTEDKTATTDSLAILKLNSNDTTSSSSFDQKSDPHLEPNIQEAQNGCTASFNRKMHLKPCATEGSFEKRLELSEIAKGDKSSFDSESIEQQDRSNTLNSPEVVTLVSDIISDYYNDASCQKKERIVSNEKFKSNVKKLHLYSCQRIVPMSGKNVWPRESCARTSEWFHKNHVSVLEGKTSTSGSQECSYENNTEAVREMVLTDNSRQLDLSMLEAESVKESVCKITMDSNIECLTSVRTPESSLLDICGTLRVTCNENLESSVDSSPIASKSEAVHEAQVILNLAVKQKQDDKNKGALTKKNRMMALQDGISEDDKTSKMIVNHKTSMIKKTVTIPNLMKMLNPGNLSNFKIPLRKNKTESRSVDYIRSLDKETYSPLECLHRSAATSARQNRAEENSSVVVSKQQPISEEADISVIPLKENADRMISKDFGSGGSENLSNEINILSERVSTYQCASLERELESSALEDVIDSSVTASPSRTLDYSSKTVDCSISLEMHDDDDKKKSKMDLAQSKIKNFPDVLKAYEDDILVIDVIQDDPDLFGTTDEGELALTMNHVTEKHCNISTTSEEKQDVKPKCLVLPETKDTRNYFRESPIQESGTLKSITDSYDALLMANKIQKHDSSSGSCPLGGVIEESFEDGQLVESDELLKDSHMDEKYSFSGKLAAIKEEKVNIHEAEKSNDSKYIEFVSSELQLGLQLPSLQINAFQEDTVLKPWINDFRYSGKSSLLPLPNSDCYEPWKMDKNVMVSHLIQQNLGMIELPRKYCRFYFSTLRGCERVECWFWHVPEQGDEKICMAILKTYININESSLLQRSVQIFMNYYREATPGVHFDSQVLNDLLISLIKCCLLKEVFQLLNVAVMIKMLPAIDVFLKVFEHVASMNIRDAVASLIDIFCKLFDAGMIFELEHFNCIIKFLHQLQVSSQEINVILNMKSRFQAKQFKKNWLRDLNLAIAEIQHCKEKSDWAKLGTLYVNMRIGCENFGDLQKLTLYIADILTRDSKEERPGVPFCEFAEAVIKNPQHNEADRDFLGRFGISVMYSYHTVQQWSKGRKVLDKLHELQIHFTSLKGIIGPERLASRCQIVNAAAEIFLKSGSLDGATWVLRESEWIINTPLWPCDKMDVLNRHNLLCAIACEYLTKNLYRQAFEILQNLPGFQYCCDTLDVSQYSLLFNKLLDACIESKGLGISSCVTDFMLSKNIPIDFTLIRGLITALGRSCLWLKARIYYKSALSMGCYPPLEGNLYRKLLLIPSYMSEIEMLLAIEIFLVSNASNIQSPCATNQTLQVVLKRSEEQKVRSKDDYQAAVERLIHAARISNPKLFLKHMTMNVNMEQVYSLEHNSALKWLKENMKWAGKVWLFSSR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYH8</accession>
    <name>A0A452GYH8_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="3">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GYH8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008235.1">
        <property type="protein sequence ID" value="ENSGAGP00000007113.1"/>
        <property type="gene ID" value="ENSGAGG00000005718.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028031">
        <property type="entry name" value="DUF4460"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027989">
        <property type="entry name" value="DUF4461"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027986">
        <property type="entry name" value="TCAIM"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31596">
        <property type="entry name" value="T-CELL ACTIVATION INHIBITOR, MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31596:SF1">
        <property type="entry name" value="T-CELL ACTIVATION INHIBITOR, MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14687">
        <property type="entry name" value="DUF4460"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14688">
        <property type="entry name" value="DUF4461"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="DUF4460" evidence="1">
        <location>
            <begin position="25"/>
            <end position="133"/>
        </location>
    </feature>
    <feature type="domain" description="DUF4461" evidence="2">
        <location>
            <begin position="185"/>
            <end position="486"/>
        </location>
    </feature>
    <evidence type="ECO:0000259" key="1">
        <source>
            <dbReference type="Pfam" id="PF14687"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF14688"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007113.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="487" mass="56770" checksum="B4078E8933447F17" modified="2019-05-08" version="1">MFCHLGAVRRLCLEKILLQWVLQARPLSGAEAVNALRPFYFAVHPDFFGQHPREREINENSLKRLNGYLENLQKPGFKSFKPTQLTFYVREREQNSSDVQESFSAAGFRAVSFTLHTRDLLSTVLDILNSCSLSTEHVQSVNSQPHKEVKGTFYRPIKWDKTYYSFTGFKDPEEELEQAQRVEITLISWLDNSEKSAIKKLRKSLPLREELNRLKDELTHQLQLSDIRWQRSWGIAHRCSQLHSLGRLAQQSPEVLRNVKGHTVVFTDCSGMSATGHVMLGTMDVHHHWIKIFERLPSYYKLQKKMLLLEDRISHLLGGIQITYIEELQPLLTLEEYYTSLDAFYNRLLNSKLSFHPRSLRGLQMILESDRYAPNLHELGHFNIPTMCDLATLQWFIITKAQQARENLKRKEELTVMEKELISASTEKFSLKQLYKEPSISHAQMIDCCKRLLEESLPYLQGMHLCISHFYSVLQDGDLCIPWNWKS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYH9</accession>
    <name>A0A452GYH9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">t-SNARE coiled-coil homology domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GYH9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008299.1">
        <property type="protein sequence ID" value="ENSGAGP00000007173.1"/>
        <property type="gene ID" value="ENSGAGG00000005759.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016192">
        <property type="term" value="P:vesicle-mediated transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.110">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010989">
        <property type="entry name" value="SNARE"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR059001">
        <property type="entry name" value="STX17_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000727">
        <property type="entry name" value="T_SNARE_dom"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05739">
        <property type="entry name" value="SNARE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26585">
        <property type="entry name" value="STX17_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47661">
        <property type="entry name" value="t-snare proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50192">
        <property type="entry name" value="T_SNARE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="2" id="KW-0472">Membrane</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="2" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="215"/>
            <end position="235"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="241"/>
            <end position="261"/>
        </location>
    </feature>
    <feature type="domain" description="T-SNARE coiled-coil homology" evidence="3">
        <location>
            <begin position="180"/>
            <end position="210"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS50192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007173.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="288" mass="32027" checksum="033188B7D4365F61" modified="2019-05-08" version="1">MSEDEEKVNIRRLEPAIQKFIKVAIPTDLERLRKHQINIKKYQKCRLWDRLHEEHINAGRTVQQLRANIREMEKLCLRVRLEDKQILQRMINPVKEEASFAIKEFLQLHSESAEELKSQFNEQEDNSLKSPLTRSATVGGEPLYNTGAVDTSQSFIQMYSPLPEIPQDENAAESWETLEESQQEKIDRIEDHVNSAAVNVEEGTKNLGKAAKYKLAALPVAGALIGGVVGGPIGLLAGFKVAGIAAALGGGVLGFTGGKLIQRRKQKMIEELSSSCPDLPSQSDKKSS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GYI6</accession>
    <name>A0A452GYI6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Protein TOPAZ1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="7">Testis- and ovary-specific PAZ domain-containing protein 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">Important for normal spermatogenesis and male fertility. Specifically required for progression to the post-meiotic stages of spermatocyte development. Seems to be necessary for normal expression levels of a number of testis-expressed gene transcripts, although its role in this process is unclear.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytosol</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000008313.1">
        <property type="protein sequence ID" value="ENSGAGP00000007187.1"/>
        <property type="gene ID" value="ENSGAGG00000005743.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048137">
        <property type="term" value="P:spermatocyte division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038952">
        <property type="entry name" value="TOPAZ1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029435">
        <property type="entry name" value="TOPAZ1_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR35671">
        <property type="entry name" value="PROTEIN TOPAZ1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR35671:SF1">
        <property type="entry name" value="PROTEIN TOPAZ1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14669">
        <property type="entry name" value="Asp_Glu_race_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="5" id="KW-0221">Differentiation</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0744">Spermatogenesis</keyword>
    <feature type="domain" description="Protein TOPAZ1" evidence="9">
        <location>
            <begin position="1064"/>
            <end position="1239"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="37"/>
            <end position="64"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00002132"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004514"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00016464"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022871"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00031943"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF14669"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007187.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1469" mass="167303" checksum="187A7729E4BA1F94" modified="2019-05-08" version="1">EVKEDSDAQDGKQLETDLNISKDYNIKKPLVAYSESLQEAKSDGTRSKAPVCPGLKRGSAVQGNRRRNRITKIRCLQNLDVKVTENKVTKSRLVKSKDEKCREIPNYHNNLPSTVERNPVVKLYDCRYLNTFVKSSVDATNCSYKIGGGFLHVAQGKRETSCPDSVMGQSDMNSKLTVMQMSTQKSIPFDKEGICQNRKNCQNENSKCLRERKWNIDRKPRKRMKLFEKSSVQNVLPVMDNKYCEYELHTEDKTATTDSLAILKLNSNDTTSSSSFDQKSDPHLEPNIQEAQNGCTASFNRKMHLKPCATEGSFEKRLELSEIAKGDKSSFDSESIEQQDRSNTLNSPEVVTLVSDIISDYYNDASCQKKERIVSNEKFKSNVKKLHLYSCQRIVPMSGKNVWPRESCARTSEWFHKNHVSVLEGKTSTSGSQECSYENNTEAVREMVLTDNSRQLDLSMLEAESVKESVCKITMDSNIECLTSVRTPESSLLDICGTLRVTCNENLESSVDSSPIASKSEAVHEAQVILNLAVKQKQDDKNKGALTKKNRMMALQDGISEDDKTSKMIVNHKTSMIKKTVTIPNLMKMLNPGNLSNFKIPLRKNKTESRSVDYIRSLDKETYSPLECLHRSAATSARQNRAEENSSVVVSKQQPISEEADISVIPLKENADRMISKDFGSGGSENLSNEINILSERDVIDSSVTASPSRTLDYSSKTVDCSISLEMHDDDDKKKSKMDLAQSKIKNFPDVLKAYEDDILVIDVIQDDPDLFGTTDEGELALTMNHVTEKHCNISTTSEEKQDVKPKCLVLPETKDTRNYFRYAYHCFTFIKIFVYSGKSSLLPLPNSDCYEPWKMDKNVMQNLGMIELPRKYCRFYFSTLRGCERVECWFWHVPEQGDEKICMAILKTYININESSLLQRSVQIFMNYYREATPGVHFDSQVLNDLLISLIKCCLLKEVFQLLNVAVMIKMLPAIDVFLKVFEHVASMNIRDAVASLIDIFCKLFDAGMIFELEHFNCIIKFLHQLQVSSQEINVILNMKSRFQAKQFKKNWLRDLNLAIAEIQHCKEKSDWAKLGTLYVNMRIGCENFGDLQKLTLYIADILTRDSKEERPGVPFCEFAEAVIKNPQHNEADRDFLGRFGISVMYSYHTVQQWSKGRKVLDKLHELQIHFTSLKGIIGPERLASRCQIVNAAAEIFLKSGSLDGATWVLRESEWIINTPLWPCDKMDVLNRHNLLCAIACEYLTKNLYRQAFEILQNLPGFQYCCDTLDVSQYSLLFNKLLDACIESKGLGISSCVTDFMLSKNIPIDFTLIRGLITALGRSCLWLKARIYYKSALSMGCYPPLEGNLYRKLLLIPSYMSEIEMLLAIEIFLVSNASNIQSPCATNQTLQVVLKRSEEQKVRSKDDYQAAVERLIHAARISNPKLFLKHMTMNVNMEQVYSLEHNSALKWLKENMKWAGKVWLFSSR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKC7</accession>
    <name>A0A452HKC7_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="9">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 7">Membrane</location>
            <topology evidence="1 7">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 7">Belongs to the reduced folate carrier (RFC) transporter (TC 2.A.48) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKC7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017557.1">
        <property type="protein sequence ID" value="ENSGAGP00000015373.1"/>
        <property type="gene ID" value="ENSGAGG00000011594.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015234">
        <property type="term" value="F:thiamine transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1250.20:FF:000225">
        <property type="entry name" value="Solute carrier family 19 member 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1250.20">
        <property type="entry name" value="MFS general substrate transporter like domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002666">
        <property type="entry name" value="Folate_carrier"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036259">
        <property type="entry name" value="MFS_trans_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00806">
        <property type="entry name" value="rfc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10686">
        <property type="entry name" value="FOLATE TRANSPORTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10686:SF37">
        <property type="entry name" value="THIAMINE TRANSPORTER 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01770">
        <property type="entry name" value="Folate_carrier"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF028739">
        <property type="entry name" value="Folate_carrier"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103473">
        <property type="entry name" value="MFS general substrate transporter"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6 7 8" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 8" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 8" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="3 7" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="12"/>
            <end position="31"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="51"/>
            <end position="73"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="85"/>
            <end position="105"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="111"/>
            <end position="131"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="143"/>
            <end position="162"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="168"/>
            <end position="190"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="369"/>
            <end position="387"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005773"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PIRNR" id="PIRNR028739"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015373.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="400" mass="45480" checksum="87D259AA2CB28565" modified="2019-05-08" version="1">MACCKEIRSSSWIYPTLVLCIYGFFSMMRPSEPFLTPYLTGPDKNLTINEVTNQVFPVWTYSYLVLLFPVFLITDYLRYKPIISLQGLSFIITWLMLLFFQGVLAMQWMEFFYGMVTATEVAYYAYIYSVVSADHYQKVTSYCRSITLVSTTVAAVLGQLLVSLAELSYFYLNAISLASLSLAFLASFFLPMPKKSMFFHKNHIPEIPQGTMGQDTMSNAPSINEQPSCQEDKDVVKSSVISRTLPECQSDKHQCHMLVQLCKDLKECYCTKKLLYWSLWWALATAGYNQIVNYIQVLWDDRAPSQNYEIYNGAVEAAATFLSSVTSMAVGYVKINWDLSGELALGIFSAVDAGSLFLMHFTTNIWACYASYLVFKSCYMLLITISTKSPLMDFNPFLIT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKD1</accession>
    <name>A0A452HKD1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">protein-arginine deiminase</fullName>
            <ecNumber evidence="3">3.5.3.15</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="7">
            <text>L-arginyl-[protein] + H2O = L-citrullyl-[protein] + NH4(+)</text>
            <dbReference type="Rhea" id="RHEA:18089"/>
            <dbReference type="Rhea" id="RHEA-COMP:10532"/>
            <dbReference type="Rhea" id="RHEA-COMP:10588"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:28938"/>
            <dbReference type="ChEBI" id="CHEBI:29965"/>
            <dbReference type="ChEBI" id="CHEBI:83397"/>
            <dbReference type="EC" id="3.5.3.15"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the protein arginine deiminase family.</text>
    </comment>
    <dbReference type="EC" id="3.5.3.15" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452HKD1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017536.1">
        <property type="protein sequence ID" value="ENSGAGP00000015354.1"/>
        <property type="gene ID" value="ENSGAGG00000011443.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004668">
        <property type="term" value="F:protein-arginine deiminase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1700:FF:000001">
        <property type="entry name" value="Protein-arginine deiminase type-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.75.10.10:FF:000003">
        <property type="entry name" value="Protein-arginine deiminase type-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.75.10.10">
        <property type="entry name" value="L-arginine/glycine Amidinotransferase, Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1700">
        <property type="entry name" value="Protein-arginine deiminase, central domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1860">
        <property type="entry name" value="Protein-arginine deiminase, N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008972">
        <property type="entry name" value="Cupredoxin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004303">
        <property type="entry name" value="PAD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013530">
        <property type="entry name" value="PAD_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036556">
        <property type="entry name" value="PAD_central_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013732">
        <property type="entry name" value="PAD_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038685">
        <property type="entry name" value="PAD_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013733">
        <property type="entry name" value="Prot_Arg_deaminase_cen_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10837">
        <property type="entry name" value="PEPTIDYLARGININE DEIMINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10837:SF11">
        <property type="entry name" value="PROTEIN-ARGININE DEIMINASE TYPE-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03068">
        <property type="entry name" value="PAD"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08527">
        <property type="entry name" value="PAD_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08526">
        <property type="entry name" value="PAD_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF001247">
        <property type="entry name" value="Protein-arginine_deiminase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49503">
        <property type="entry name" value="Cupredoxins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55909">
        <property type="entry name" value="Pentein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF110083">
        <property type="entry name" value="Peptidylarginine deiminase Pad4, middle domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0106">Calcium</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="5" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Protein-arginine deiminase (PAD) N-terminal" evidence="10">
        <location>
            <begin position="11"/>
            <end position="92"/>
        </location>
    </feature>
    <feature type="domain" description="Protein-arginine deiminase (PAD) central" evidence="11">
        <location>
            <begin position="109"/>
            <end position="246"/>
        </location>
    </feature>
    <feature type="domain" description="Protein-arginine deiminase C-terminal" evidence="9">
        <location>
            <begin position="257"/>
            <end position="494"/>
        </location>
    </feature>
    <feature type="domain" description="Protein-arginine deiminase C-terminal" evidence="9">
        <location>
            <begin position="507"/>
            <end position="622"/>
        </location>
    </feature>
    <feature type="active site" evidence="8">
        <location>
            <position position="327"/>
        </location>
    </feature>
    <feature type="active site" evidence="8">
        <location>
            <position position="447"/>
        </location>
    </feature>
    <feature type="active site" evidence="8">
        <location>
            <position position="449"/>
        </location>
    </feature>
    <feature type="active site" evidence="8">
        <location>
            <position position="607"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008166"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012200"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00048487"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PIRSR" id="PIRSR001247-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF03068"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="Pfam" id="PF08526"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF08527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015354.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="625" mass="69879" checksum="ACA6A59CD37DA9BF" modified="2019-05-08" version="1">MSLGKGVRVLSIPSGATSFDARGTSGVKIYVVYNPNAVKKAAGASRWTLNSQVEVIVSMDIPSSTVNDSKVRISYYGQKENAPIEGAWLYLTCVTWSVFVPDPFVFPLQTKWTWGPEGQGAVLLVNCDRDSPGAKDMDSAYQYVQSYADLEDMSLMILRTQGPSAIFADYKLVLHISVADVNKVRVFHASGSGSLAQYKHVLGSDKRSYVVDRSSGQEEDTFYVEGLAFPDADFSGLVSFHVTLLEVSDTNAPDTPIFTDTVVFRVAPWIMTPNTLQPLEVFVCSMERGSNPNGDFLEAMRALVRKANCKLTVCPEVENRRDRWIQDEMEFGYVEAPHKMFPVVFDSPRDRGLRDFAFRRILGPDFGYVTRVPPHGHVSSLDSFGNLDVSPPVTVQGKTYPLGRILIGSSLPRSDGRRMTKAVRDFLYAQKVQPPVELYSDWLSVGHVDEFLSFVPASDRKGFRLLLASPDACYKLFKEKQREGHGKAAQFAGEWLSQDAFGTHRGCCIDWNRHTLKEELGLTEQDIIDIPQLFMLKGSRADAFFPDVVNMIVLGKHLGIPKPFGPIVDGRCCLEEKVRSLLEPLGLVCIFINDFFSYHTLLGEIHCGTNVLRKPFSFKWWNMIL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKD3</accession>
    <name>A0A452HKD3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Latent-transforming growth factor beta-binding protein 1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="19">Latent-transforming growth factor beta-binding protein 2</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="20">Transforming growth factor beta-1-binding protein 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="25 26">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="26" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="25" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="25" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">Key regulator of transforming growth factor beta (TGFB1, TGFB2 and TGFB3) that controls TGF-beta activation by maintaining it in a latent state during storage in extracellular space. Associates specifically via disulfide bonds with the Latency-associated peptide (LAP), which is the regulatory chain of TGF-beta, and regulates integrin-dependent activation of TGF-beta. Outcompeted by LRRC32/GARP for binding to LAP regulatory chain of TGF-beta.</text>
    </comment>
    <comment type="function">
        <text evidence="14">May play an integral structural role in elastic-fiber architectural organization and/or assembly.</text>
    </comment>
    <comment type="subunit">
        <text evidence="16">Forms part of the large latent transforming growth factor beta precursor complex; removal is essential for activation of complex. Interacts with SDC4. Interacts (via C-terminal domain) with FBN1 (via N-terminal domain) in a Ca(+2)-dependent manner.</text>
    </comment>
    <comment type="subunit">
        <text evidence="17">Interacts with TGFB1; associates via disulfide bonds with the Latency-associated peptide chain (LAP) regulatory chain of TGFB1, leading to regulate activation of TGF-beta-1. LTBP1 does not bind directly to TGF-beta-1, the active chain of TGFB1. Interacts (via C-terminal domain) with FBN1 (via N-terminal domain). Interacts with FBN2. Interacts with ADAMTSL2. Interacts with EFEMP2.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="13">Belongs to the LTBP family.</text>
    </comment>
    <comment type="caution">
        <text evidence="21">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000017564.1">
        <property type="protein sequence ID" value="ENSGAGP00000015379.1"/>
        <property type="gene ID" value="ENSGAGG00000008780.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019838">
        <property type="term" value="F:growth factor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00054">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000005">
        <property type="entry name" value="Fibrillin 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000194">
        <property type="entry name" value="Latent transforming growth factor beta binding protein 2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000469">
        <property type="entry name" value="Latent transforming growth factor beta binding protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.290.10:FF:000004">
        <property type="entry name" value="latent-transforming growth factor beta-binding protein 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000019">
        <property type="entry name" value="latent-transforming growth factor beta-binding protein 1 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000046">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 1 isoform x2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000014">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000077">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 3 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.290.10:FF:000001">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 3 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.290.10:FF:000002">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 3 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000056">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 3 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000115">
        <property type="entry name" value="latent-transforming growth factor beta-binding protein 4 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000024">
        <property type="entry name" value="Putative latent-transforming growth factor beta-binding protein 2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000273">
        <property type="entry name" value="Putative latent-transforming growth factor beta-binding protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="18"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.290.10">
        <property type="entry name" value="TGF-beta binding (TB) domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026823">
        <property type="entry name" value="cEGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001881">
        <property type="entry name" value="EGF-like_Ca-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013032">
        <property type="entry name" value="EGF-like_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000152">
        <property type="entry name" value="EGF-type_Asp/Asn_hydroxyl_site"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018097">
        <property type="entry name" value="EGF_Ca-bd_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009030">
        <property type="entry name" value="Growth_fac_rcpt_cys_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049883">
        <property type="entry name" value="NOTCH1_EGF-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017878">
        <property type="entry name" value="TB_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036773">
        <property type="entry name" value="TB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052080">
        <property type="entry name" value="vWF_C/EGF_Fibrillin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47333:SF4">
        <property type="entry name" value="EGF-LIKE DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47333">
        <property type="entry name" value="VON WILLEBRAND FACTOR C AND EGF DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12662">
        <property type="entry name" value="cEGF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07645">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12661">
        <property type="entry name" value="hEGF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00683">
        <property type="entry name" value="TB"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="19"/>
    </dbReference>
    <dbReference type="SMART" id="SM00179">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="18"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57184">
        <property type="entry name" value="Growth factor receptor domain"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57581">
        <property type="entry name" value="TB module/8-cys domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00010">
        <property type="entry name" value="ASX_HYDROXYL"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01186">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01187">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51364">
        <property type="entry name" value="TB"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9 21" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="4 21" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="3" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11" id="KW-0340">Growth factor binding</keyword>
    <keyword evidence="6" id="KW-0358">Heparin-binding</keyword>
    <keyword evidence="12" id="KW-0379">Hydroxylation</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="26" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="86"/>
            <end position="118"/>
        </location>
    </feature>
    <feature type="domain" description="TB" evidence="24">
        <location>
            <begin position="233"/>
            <end position="276"/>
        </location>
    </feature>
    <feature type="domain" description="TB" evidence="24">
        <location>
            <begin position="353"/>
            <end position="405"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="514"/>
            <end position="555"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="556"/>
            <end position="592"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="599"/>
            <end position="638"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="679"/>
            <end position="719"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="804"/>
            <end position="844"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="845"/>
            <end position="886"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="887"/>
            <end position="927"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="972"/>
            <end position="1013"/>
        </location>
    </feature>
    <feature type="domain" description="TB" evidence="24">
        <location>
            <begin position="1078"/>
            <end position="1132"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="1154"/>
            <end position="1192"/>
        </location>
    </feature>
    <feature type="domain" description="TB" evidence="24">
        <location>
            <begin position="1254"/>
            <end position="1306"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="1446"/>
            <end position="1490"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="203"/>
            <end position="228"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="426"/>
            <end position="507"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="1373"/>
            <end position="1396"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="22">
        <location>
            <begin position="203"/>
            <end position="227"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="22">
        <location>
            <begin position="1383"/>
            <end position="1396"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="21">
        <location>
            <begin position="90"/>
            <end position="100"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="21">
        <location>
            <begin position="108"/>
            <end position="117"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022674"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023183"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023278"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00038081"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00058734"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00059743"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00062144"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00062844"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00072992"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00072997"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00075443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS51364"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015379.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1502" mass="162844" checksum="76992DEFD1D5BCBE" modified="2019-05-08" version="1">MSNALPNGNGHEQSSSGPHAAYQEHTLSLWGANLTEKIRKIKIVFTPTICKQTCRSGHCYNSCQKGDTTTLYSQGGQNHDPKSGFRIYFCQIPCLNGGRCVGRDECWCPSNSTGKFCHLPVPKLEKNQVGRVPKTLASGSIKQSMYTLPLSNQLASLNPSLVNVHIKHPPEATVQIHQVARVRGDLGQSEENSVEAVLLSQLPTGPQHSYSNGNSNSITTESGQQQRPPELMGRCFRETLHGQCANPLEGLAKFEDCCGSVGLFWGVNRCIPCPPRPAHPVIENRQVECPQGYKRLNRSHCQDINECLMLGLCKDTECVNTRGSFLCTCKPGTMLDPSRSRCVSDKAVSMEQGLCYRSAAEGVCALPLAQSITKQICCCSRVGKGWGRNCEKCPMLGSETFKEICPAGHGYTYSSSDIRLSMRKAEAEELPFSSEEQGESSDQTPTQAAKRLQLQEALSGGVTGHSPDSEGTADATQEPAPHPGDVVEETFPHPPSQPGPEAAGERTAAAQITEMDRCASVPDPCGPGTCVNLPDRYSCICGPGYQLHPGLPQCIDDDECMRDPCAGKGHCINSVGSYSCLCYPGYTLLASQGTQTCQDLNECEQPSMCQGGQCTNTPGSYHCECARGYIMGHTGQCEDIDECADPSTCPRGRCVNALGSYECLSCAAGYQPMNGRCVDDDECFTVGICAHGLCINLAGSFRCSCYHGYEVTADEKSCQDINECATHNACPMGLCINSEGSFSCAACGAGYTVSRDGSMCEDIDECSSATACPLGICTNTDGSFACRACDAGYMLSSNRFTCEDIDECEDPTVSCLGGECLNTPGSYVCHCRPGFELLNGTMCEDMNECLASEICSPNGECLNSHGSYFCICAPGFSNAAGGVSCQDVDECADKSRCLRGQCFNTEGSYRCLCENGFRHSQETDDCVDVDECVDFGDTVCGTWRCQNTPGSYRCVMGCQPGFHRTPLGDCIDIDECANETLCGSHAFCDNTDGSFRCLCDSGYENLPPGQDCVDVNECELMVAVCGTALCENVEGSFLCLCLSDHEEYNTEAGECRPRAGMGDPERPIAPRPAGSERKECYYHISDVQLCDSVLAKNTTKEECCCTEGAAWGDNCETHPCPILGTVEYSEICPSGKGYIPVEGTLLFGQTSYTDADECEMFGPEICRNGHCVNTVPGYKCFCRSGYLYDSSRLECVDQDECENEMSCVNGECLNTAGSFHCFCGLPLVLDVTRNRCINVSSRAEDPEEHEIHLDICWQRVSNYICNQPLRGRRTTYTECCCRYGEAWSQDCALCPHRASEDFAHLCNVARGETEREPGLQERPGYEYGPALEDSHYGLYSPDLGPYYNYLSPEYGAPDASFPQRELSSEVGGSTFLRTPQLPPAQSQPRYLPSQTGHFDSFEGLQAEECGILNGCENGRCVRVREGYTCDCFDGFQLDMTHMACVDINECEEVSSSGPLCEGGTCENTEGSYRCNCSPGFVVQAEPHRCAPAPAQRPEAAQL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKD6</accession>
    <name>A0A452HKD6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">ubiquitinyl hydrolase 1</fullName>
            <ecNumber evidence="3">3.4.19.12</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="8">Deubiquitinating enzyme A</fullName>
        </alternativeName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>Thiol-dependent hydrolysis of ester, thioester, amide, peptide and isopeptide bonds formed by the C-terminal Gly of ubiquitin (a 76-residue protein attached to proteins as an intracellular targeting signal).</text>
            <dbReference type="EC" id="3.4.19.12"/>
        </reaction>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the peptidase C85 family.</text>
    </comment>
    <dbReference type="EC" id="3.4.19.12" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452HKD6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017541.1">
        <property type="protein sequence ID" value="ENSGAGP00000015359.1"/>
        <property type="gene ID" value="ENSGAGG00000011581.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004843">
        <property type="term" value="F:cysteine-type deubiquitinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061578">
        <property type="term" value="F:K63-linked deubiquitinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010629">
        <property type="term" value="P:negative regulation of gene expression"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051241">
        <property type="term" value="P:negative regulation of multicellular organismal process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016579">
        <property type="term" value="P:protein deubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001817">
        <property type="term" value="P:regulation of cytokine production"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd22752">
        <property type="entry name" value="OTU_OTUD5-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.70.80:FF:000002">
        <property type="entry name" value="OTU domain-containing protein 5 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.70.80">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003323">
        <property type="entry name" value="OTU_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038765">
        <property type="entry name" value="Papain-like_cys_pep_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050704">
        <property type="entry name" value="Peptidase_C85-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12419">
        <property type="entry name" value="OTU DOMAIN CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12419:SF4">
        <property type="entry name" value="OTU DOMAIN-CONTAINING PROTEIN 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02338">
        <property type="entry name" value="OTU"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54001">
        <property type="entry name" value="Cysteine proteinases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50802">
        <property type="entry name" value="OTU"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="4" id="KW-0645">Protease</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0788">Thiol protease</keyword>
    <keyword evidence="5" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="OTU" evidence="10">
        <location>
            <begin position="1"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="199"/>
            <end position="283"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="205"/>
            <end position="218"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="225"/>
            <end position="236"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000707"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010407"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012759"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022807"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00033460"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50802"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015359.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="348" mass="38817" checksum="4BD961674F621D89" modified="2019-05-08" version="1">IKQMKEDGACLFRAVADQVYGDQDMHEVVRKHCMDYLMKNADYFSNYVTEDFTTYINRKRKNNCHGNHIEMQAMAEMYNRPVEVYQYGTEPINTFHGIQQNEDEPIRVSYHRNIHYNSVVNPNKATIGVGLGLPSFKPGYAEQSLMKSAIKTSEESWIEQQMLEDKKRATDWEATNEAIEEQVARESYLQWLRDQEKQARQPRKASATCSSATAAATSGLEDWSSRSPRQRSSAPSPEHPDLHGEITALKPPSPGTAATLALPKPPSPCAPGTSSQLSAGGGRATPPLVSLYPALECRAIMQQMSPTAFGLTDWDEDEILASVLAVSQQEYLESMKGSLHRDSPADKS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKD7</accession>
    <name>A0A452HKD7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Tripartite motif containing 39</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HKD7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017569.1">
        <property type="protein sequence ID" value="ENSGAGP00000015383.1"/>
        <property type="gene ID" value="ENSGAGG00000011603.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd19762">
        <property type="entry name" value="Bbox2_TRIM7-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12888">
        <property type="entry name" value="SPRY_PRY_TRIM7_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.920:FF:000004">
        <property type="entry name" value="Butyrophilin subfamily 1 member A1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.920">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001870">
        <property type="entry name" value="B30.2/SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043136">
        <property type="entry name" value="B30.2/SPRY_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003879">
        <property type="entry name" value="Butyrophylin_SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006574">
        <property type="entry name" value="PRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003877">
        <property type="entry name" value="SPRY_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050143">
        <property type="entry name" value="TRIM/RBCC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000315">
        <property type="entry name" value="Znf_B-box"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24103">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE TRIM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13765">
        <property type="entry name" value="PRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00622">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00643">
        <property type="entry name" value="zf-B_box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01407">
        <property type="entry name" value="BUTYPHLNCDUF"/>
    </dbReference>
    <dbReference type="SMART" id="SM00336">
        <property type="entry name" value="BBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00589">
        <property type="entry name" value="PRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00449">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57845">
        <property type="entry name" value="B-box zinc-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50188">
        <property type="entry name" value="B302_SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50119">
        <property type="entry name" value="ZF_BBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="1 3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0862">Zinc</keyword>
    <keyword evidence="1 3" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="B box-type" evidence="5">
        <location>
            <begin position="9"/>
            <end position="50"/>
        </location>
    </feature>
    <feature type="domain" description="B30.2/SPRY" evidence="6">
        <location>
            <begin position="180"/>
            <end position="376"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="4">
        <location>
            <begin position="51"/>
            <end position="144"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00024"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50119"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50188"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015383.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="9">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="376" mass="42928" checksum="7387DE84658C6AA7" modified="2019-05-08" version="1">NGPAAKGSGGERVCEKHQEILKLFCEEDQAPICVICRESQAHRAHMVVPIEEAAQEYKEKIQIQLQILKEEREKLMGLKLTKERLKILSEFEQLRQFLEEQERLLLARLEELDKEIEKRRDENIAKLLEEISCLNDMISELEGKCQQPASEFLQVRYWILDTQEPWSPGEVSPEPEKTPSSISHKYIALKETLRKFQVDVTLDPDTAHPRLVLSEDGKSVRWEDTRQHVPDNPKRFDSSRCVLGLEGFSSGRHYWEVEVEDGGAWAVGVARESVRRKGRVNVNPEEGIWAVGQCGTQYLALTSPKIPLSLAERPRTIGIYLDCEEGRVAFFDADNEGPIFTFPPASVPGEKILPLLCLGRWKKKSCVKLQCCYNFG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKF1</accession>
    <name>A0A452HKF1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">PH domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the GAB family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKF1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017559.1">
        <property type="protein sequence ID" value="ENSGAGP00000015375.1"/>
        <property type="gene ID" value="ENSGAGG00000011596.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035591">
        <property type="term" value="F:signaling adaptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046355">
        <property type="entry name" value="Gab1-4-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45960">
        <property type="entry name" value="GRB2-ASSOCIATED-BINDING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45960:SF3">
        <property type="entry name" value="GRB2-ASSOCIATED-BINDING PROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00169">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="PH" evidence="3">
        <location>
            <begin position="1"/>
            <end position="80"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="92"/>
            <end position="136"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="399"/>
            <end position="431"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="92"/>
            <end position="122"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00029462"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015375.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="561" mass="62139" checksum="A869602561A92669" modified="2019-05-08" version="1">MSGNPDVLEYYRNNHSKKPIRIIDLNECEVLKHSGSNFIKKEFQNNFVFIVKTTYRSFYLVAKTEEEMQIWVHNISQICNFGHLEDGTDSVESLSHTTSSLQPSPAVSTHTSRITDPSSSVGSATTDASASEETRSESESVFLPDYLFLSNCETGKFNHIRCDSWSNSDRSLEQTSSDDVFVDALQSQPSLHVQSSSNGIVHQDGGQPANHGIIPRSIDFSGPFSDVPSSSPLLGPPLTPTIQIDKSQTALPYGVTELDILSNTPPPRPPKPTHFSDRSGEELSNGALQNGHGGICGTQIALVPRRISLSSLDNMRHWKASMEGSSLRSRDKRLSLNLPCRFTPLYSSASDNTEDSYVPMHPSTSPPIPGSDCSPDGYTPMSPSSATFTFPVITNDKLSSPLPELRTDLEPPPVNRDLKPRRKSRPPPLDLRNLSTIREHATLTRMWTVPYNRTSFISPDKDGINSARIFGNSVSGEEEESYIQMEHSQATSPCSGAFPWSKKSNLDYLALDFNSASPSPVQKKPFLSEEQRVDYVQVDEQKTQALQNTKQEWTDVRQSKV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKF5</accession>
    <name>A0A452HKF5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Interleukin 34</fullName>
        </recommendedName>
    </protein>
    <organism evidence="1 2">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="2" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="1" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="1" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HKF5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017566.1">
        <property type="protein sequence ID" value="ENSGAGP00000015380.1"/>
        <property type="gene ID" value="ENSGAGG00000011602.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005157">
        <property type="term" value="F:macrophage colony-stimulating factor receptor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008284">
        <property type="term" value="P:positive regulation of cell population proliferation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045651">
        <property type="term" value="P:positive regulation of macrophage differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045657">
        <property type="term" value="P:positive regulation of monocyte differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1250.80">
        <property type="entry name" value="Interleukin-34"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020415">
        <property type="entry name" value="IL-34"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038328">
        <property type="entry name" value="IL-34_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR28606">
        <property type="entry name" value="INTERLEUKIN-34"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR28606:SF1">
        <property type="entry name" value="INTERLEUKIN-34"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15036">
        <property type="entry name" value="IL34"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01938">
        <property type="entry name" value="INTRLEUKIN34"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000313" key="1">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015380.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="3">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="156" mass="18291" checksum="60EC0A6EAC358620" modified="2019-05-08" version="1">MQKECRIIEALQNKLTYRQRLQYMKHYFPINYTVNVQFEEVLRASNITRLRDQNVSELSLRCLWLSVNSQVLLKIGAVLLEKHLSWEYIQDLCLLFEHLAEEYEDCNQGNVDTNVWDVVEQVLTGDTGSGQKAVRPKALLDNCVKVMWMLYSESCK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKG0</accession>
    <name>A0A452HKG0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">FAD-binding PCMH-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="1 14">
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="15">
            <name>Mo-molybdopterin</name>
            <dbReference type="ChEBI" id="CHEBI:71302"/>
        </cofactor>
        <text evidence="15">Binds 1 Mo-molybdopterin (Mo-MPT) cofactor per subunit.</text>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="12">
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="15">
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:49601"/>
        </cofactor>
        <text evidence="15">Binds 2 [2Fe-2S] clusters.</text>
    </comment>
    <comment type="subunit">
        <text evidence="3">Homodimer.</text>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the xanthine dehydrogenase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKG0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017610.1">
        <property type="protein sequence ID" value="ENSGAGP00000015416.1"/>
        <property type="gene ID" value="ENSGAGG00000010622.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051537">
        <property type="term" value="F:2 iron, 2 sulfur cluster binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071949">
        <property type="term" value="F:FAD binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005506">
        <property type="term" value="F:iron ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016491">
        <property type="term" value="F:oxidoreductase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.365.10:FF:000003">
        <property type="entry name" value="Aldehyde oxidase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1170.50:FF:000001">
        <property type="entry name" value="Aldehyde oxidase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.365.10:FF:000025">
        <property type="entry name" value="Aldehyde oxidase 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.365.10:FF:000001">
        <property type="entry name" value="Xanthine dehydrogenase oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.365.10:FF:000004">
        <property type="entry name" value="Xanthine dehydrogenase oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.390.50:FF:000001">
        <property type="entry name" value="Xanthine dehydrogenase oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.43.10:FF:000001">
        <property type="entry name" value="Xanthine dehydrogenase/oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.465.10:FF:000004">
        <property type="entry name" value="Xanthine dehydrogenase/oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.20.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.465.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.120">
        <property type="entry name" value="[2Fe-2S]-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1170.50">
        <property type="entry name" value="Aldehyde oxidase/xanthine dehydrogenase, a/b hammerhead"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.365.10">
        <property type="entry name" value="Aldehyde oxidase/xanthine dehydrogenase, molybdopterin binding domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.390.50">
        <property type="entry name" value="CO dehydrogenase flavoprotein, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.43.10">
        <property type="entry name" value="Uridine Diphospho-n-acetylenolpyruvylglucosamine Reductase, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002888">
        <property type="entry name" value="2Fe-2S-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036884">
        <property type="entry name" value="2Fe-2S-bd_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036010">
        <property type="entry name" value="2Fe-2S_ferredoxin-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001041">
        <property type="entry name" value="2Fe-2S_ferredoxin-type"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006058">
        <property type="entry name" value="2Fe2S_fd_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000674">
        <property type="entry name" value="Ald_Oxase/Xan_DH_a/b"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036856">
        <property type="entry name" value="Ald_Oxase/Xan_DH_a/b_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016208">
        <property type="entry name" value="Ald_Oxase/xanthine_DH-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008274">
        <property type="entry name" value="AldOxase/xan_DH_MoCoBD1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046867">
        <property type="entry name" value="AldOxase/xan_DH_MoCoBD2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037165">
        <property type="entry name" value="AldOxase/xan_DH_Mopterin-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012675">
        <property type="entry name" value="Beta-grasp_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005107">
        <property type="entry name" value="CO_DH_flav_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036683">
        <property type="entry name" value="CO_DH_flav_C_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016166">
        <property type="entry name" value="FAD-bd_PCMH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036318">
        <property type="entry name" value="FAD-bd_PCMH-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016167">
        <property type="entry name" value="FAD-bd_PCMH_sub1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016169">
        <property type="entry name" value="FAD-bd_PCMH_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002346">
        <property type="entry name" value="Mopterin_DH_FAD-bd"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45444">
        <property type="entry name" value="XANTHINE DEHYDROGENASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45444:SF3">
        <property type="entry name" value="XANTHINE DEHYDROGENASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01315">
        <property type="entry name" value="Ald_Xan_dh_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03450">
        <property type="entry name" value="CO_deh_flav_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00941">
        <property type="entry name" value="FAD_binding_5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00111">
        <property type="entry name" value="Fer2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01799">
        <property type="entry name" value="Fer2_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02738">
        <property type="entry name" value="MoCoBD_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20256">
        <property type="entry name" value="MoCoBD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000127">
        <property type="entry name" value="Xanthine_DH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01008">
        <property type="entry name" value="Ald_Xan_dh_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01092">
        <property type="entry name" value="CO_deh_flav_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54292">
        <property type="entry name" value="2Fe-2S ferredoxin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55447">
        <property type="entry name" value="CO dehydrogenase flavoprotein C-terminal domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47741">
        <property type="entry name" value="CO dehydrogenase ISP C-domain like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54665">
        <property type="entry name" value="CO dehydrogenase molybdoprotein N-domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56176">
        <property type="entry name" value="FAD-binding/transporter-associated domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56003">
        <property type="entry name" value="Molybdenum cofactor-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00197">
        <property type="entry name" value="2FE2S_FER_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51387">
        <property type="entry name" value="FAD_PCMH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6 15" id="KW-0001">2Fe-2S</keyword>
    <keyword evidence="8 14" id="KW-0274">FAD</keyword>
    <keyword evidence="5" id="KW-0285">Flavoprotein</keyword>
    <keyword evidence="10 15" id="KW-0408">Iron</keyword>
    <keyword evidence="11 15" id="KW-0411">Iron-sulfur</keyword>
    <keyword evidence="7 15" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="4 15" id="KW-0500">Molybdenum</keyword>
    <keyword evidence="9" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="FAD-binding PCMH-type" evidence="16">
        <location>
            <begin position="173"/>
            <end position="358"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="13">
        <location>
            <position position="1202"/>
        </location>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="20"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="25"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="28"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="50"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="90"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>2</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="93"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>2</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <begin position="201"/>
            <end position="208"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="304"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="366"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="708"/>
        </location>
        <ligand>
            <name>Mo-molybdopterin</name>
            <dbReference type="ChEBI" id="CHEBI:71302"/>
        </ligand>
        <ligandPart>
            <name>Mo</name>
            <dbReference type="ChEBI" id="CHEBI:28685"/>
        </ligandPart>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="739"/>
        </location>
        <ligand>
            <name>Mo-molybdopterin</name>
            <dbReference type="ChEBI" id="CHEBI:71302"/>
        </ligand>
        <ligandPart>
            <name>Mo</name>
            <dbReference type="ChEBI" id="CHEBI:28685"/>
        </ligandPart>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="853"/>
        </location>
        <ligand>
            <name>Mo-molybdopterin</name>
            <dbReference type="ChEBI" id="CHEBI:71302"/>
        </ligand>
        <ligandPart>
            <name>Mo</name>
            <dbReference type="ChEBI" id="CHEBI:28685"/>
        </ligandPart>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="855"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="1020"/>
        </location>
        <ligand>
            <name>Mo-molybdopterin</name>
            <dbReference type="ChEBI" id="CHEBI:71302"/>
        </ligand>
        <ligandPart>
            <name>Mo</name>
            <dbReference type="ChEBI" id="CHEBI:28685"/>
        </ligandPart>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001974"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006849"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00011738"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022505"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022630"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022714"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022827"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023014"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00034078"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PIRSR" id="PIRSR000127-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PIRSR" id="PIRSR000127-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PIRSR" id="PIRSR000127-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS51387"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015416.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1273" mass="140783" checksum="7E3C16A0D5102A18" modified="2019-05-08" version="1">QLFSLTAAFAVRLSGTKYGCGIGGCGACTVMISTYEPTSKKIMHYPANSCLLPICSLHSSAVTTVEGVGSTKTRVHPIQERLAKCHGSQCGFCTPGMVMSMYALLRNHPEPSMEQITAALDGRFTFTSSKLKEMHSGLCNPEEFQPVDPTQEFIFPPELMRMAQEQQKRTLVFRGERTTWISPTNLRELLELKAMYPKAPLIVGNTSVGLEMKRDGVHHPVFIHPTRIPDLHVVSNTNNGLVIGAACSLAQLRDILMKVVPELPEEKTKIFRALLQQLKTLAGEQIRSLASLGGHIVSRGSTWDLNPVLAVGNSVLSLASKDGTRQIPLNDQFLAGYENADLKPKEIIVSVFIPYSKKDDFVSAFRQAERRRNAVSVVNSGMRVLFKPNTDIIMDMNILYGGTGPITVSARKSCQRLIGRQWNEQMLDEACKLVLGEISLSPSAPGGKVEYRRTLLVSFLFKFYLEVLYGLNKMYPAKFPDLSKKNLSALGAFHSEVPQGMQTYQDVDPVQPPQDPVGRPIMHQSGVKHATGEAVYCNDIRAVDEELFLAVVTSSRAHAKIVTIDVSEALKVPGVVDVVTAEDVPGKNGTDDEQAYAKDEVICVGQIVCAVVAESAVQAKQGVEKMKIEYEDLEPILTIEDAIKHNSYFGKEKKIEQGNIEEGFKSADEIIEGEIHIGGQEHFYLETNSVLVVPRGEDNEMDVHVSTQDAARVQEFVALALDVQSSKIMCHTKRVGGAFGGKITKPAIFAVVAAVAANKTGRPIRFTLERDEDMLITGGRHPFFGKYKVGFTNDGRITAADIEPYINAGCTLDDSEMVIEYVVLKTDNAYKIPNLRIRGRACKTNLPSNTAFRGFGFPQSGLFTESWITTVATQTGLPQEKIREINMYKEVNLTPFKEELDATNLVKCWKECMEKSEYYSRKTAVEEFNKQNYWKKKGIAIIPMKFSVGFNATQYHQAAALVHIYTDGSVLVTHGGIELGQGIHTKMLQIASRELKVPLSYVHFCETSTITVPNAIVTAGSIGSDVNGKAVQNACQILRKRLEPIINKNPEGKWEDWITEAYDQSVSLSATGYFRGYMVNMNWDTGEGQAFPYFVFGAACSEVEIDCLTGDHKNIRTDIVMDACFSINPAVDIGQIEGAFVQGVGLYTMEEIKFSPEGEQYTLGPDTYKIPAVCDIPEQFRVYLLPNSRNPIAIYSAKGIGEAGLFLGCSVFFAIWDAVAAVRKERGLTGTFTLNSPLTIERIRMACADNFTEMVMLHLFYVLHYTHWSLHSI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKG3</accession>
    <name>A0A452HKG3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Programmed cell death protein 5</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="5">May function in the process of apoptosis.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the PDCD5 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKG3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017577.1">
        <property type="protein sequence ID" value="ENSGAGP00000015390.1"/>
        <property type="gene ID" value="ENSGAGG00000011611.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006915">
        <property type="term" value="P:apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.8.140:FF:000001">
        <property type="entry name" value="Programmed cell death protein 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.8.140">
        <property type="entry name" value="PDCD5-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002836">
        <property type="entry name" value="PDCD5-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036883">
        <property type="entry name" value="PDCD5-like_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10840">
        <property type="entry name" value="PROGRAMMED CELL DEATH PROTEIN 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10840:SF0">
        <property type="entry name" value="PROGRAMMED CELL DEATH PROTEIN 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01984">
        <property type="entry name" value="dsDNA_bind"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF015730">
        <property type="entry name" value="TFAR19"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46950">
        <property type="entry name" value="Double-stranded DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0007">Acetylation</keyword>
    <keyword evidence="3" id="KW-0053">Apoptosis</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="13"/>
            <end position="41"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="13"/>
            <end position="24"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00010490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022703"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00056824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00071574"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015390.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="127" mass="14304" checksum="5AF54B51ECB8AA33" modified="2019-05-08" version="1">MADEELEALRKQRLAELQAKHGDPSGDPAQQEAKQRESEMRNSILAQVLDQAARARLSNLALVKPEKAKAVENYLIQMARFGQLGGKVSEQGLIEILEKVSQQTEKKTTVKFNRRKVLDSDEEDDDD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKG5</accession>
    <name>A0A452HKG5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Methionine--tRNA ligase, cytoplasmic</fullName>
            <ecNumber evidence="3">6.1.1.10</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="12">Methionyl-tRNA synthetase</fullName>
        </alternativeName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>tRNA(Met) + L-methionine + ATP = L-methionyl-tRNA(Met) + AMP + diphosphate</text>
            <dbReference type="Rhea" id="RHEA:13481"/>
            <dbReference type="Rhea" id="RHEA-COMP:9667"/>
            <dbReference type="Rhea" id="RHEA-COMP:9698"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:33019"/>
            <dbReference type="ChEBI" id="CHEBI:57844"/>
            <dbReference type="ChEBI" id="CHEBI:78442"/>
            <dbReference type="ChEBI" id="CHEBI:78530"/>
            <dbReference type="ChEBI" id="CHEBI:456215"/>
            <dbReference type="EC" id="6.1.1.10"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 14">Belongs to the class-I aminoacyl-tRNA synthetase family.</text>
    </comment>
    <dbReference type="EC" id="6.1.1.10" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452HKG5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017618.1">
        <property type="protein sequence ID" value="ENSGAGP00000015421.1"/>
        <property type="gene ID" value="ENSGAGG00000011640.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017101">
        <property type="term" value="C:aminoacyl-tRNA synthetase multienzyme complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004825">
        <property type="term" value="F:methionine-tRNA ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006431">
        <property type="term" value="P:methionyl-tRNA aminoacylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd07957">
        <property type="entry name" value="Anticodon_Ia_Met"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00814">
        <property type="entry name" value="MetRS_core"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00939">
        <property type="entry name" value="MetRS_RNA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.28.20:FF:000001">
        <property type="entry name" value="Methionine--tRNA ligase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.730.10:FF:000010">
        <property type="entry name" value="methionine--tRNA ligase, cytoplasmic"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1050.10:FF:000026">
        <property type="entry name" value="Methionine--tRNA ligase, cytoplasmic"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.620:FF:000640">
        <property type="entry name" value="Uncharacterized protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1050.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.30.10">
        <property type="entry name" value="Glutaredoxin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.620">
        <property type="entry name" value="HUPs"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.730.10">
        <property type="entry name" value="Isoleucyl-tRNA Synthetase, Domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.20.28.20">
        <property type="entry name" value="Methionyl-tRNA synthetase, Zn-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.10">
        <property type="entry name" value="S15/NS1, RNA-binding"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="HAMAP" id="MF_00098">
        <property type="entry name" value="Met_tRNA_synth_type1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001412">
        <property type="entry name" value="aa-tRNA-synth_I_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041872">
        <property type="entry name" value="Anticodon_Met"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010987">
        <property type="entry name" value="Glutathione-S-Trfase_C-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036282">
        <property type="entry name" value="Glutathione-S-Trfase_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041598">
        <property type="entry name" value="MARS_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023458">
        <property type="entry name" value="Met-tRNA_ligase_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014758">
        <property type="entry name" value="Met-tRNA_synth"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015413">
        <property type="entry name" value="Methionyl/Leucyl_tRNA_Synth"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033911">
        <property type="entry name" value="MetRS_core"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029038">
        <property type="entry name" value="MetRS_Zn"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014729">
        <property type="entry name" value="Rossmann-like_a/b/a_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009080">
        <property type="entry name" value="tRNAsynth_Ia_anticodon-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009068">
        <property type="entry name" value="uS15_NS1_RNA-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000738">
        <property type="entry name" value="WHEP-TRS_dom"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00398">
        <property type="entry name" value="metG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF001100">
        <property type="entry name" value="PRK00133.1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45765">
        <property type="entry name" value="METHIONINE--TRNA LIGASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45765:SF1">
        <property type="entry name" value="METHIONINE--TRNA LIGASE, CYTOPLASMIC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19303">
        <property type="entry name" value="Anticodon_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18485">
        <property type="entry name" value="GST_N_5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09334">
        <property type="entry name" value="tRNA-synt_1g"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00458">
        <property type="entry name" value="WHEP-TRS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01041">
        <property type="entry name" value="TRNASYNTHMET"/>
    </dbReference>
    <dbReference type="SMART" id="SM00991">
        <property type="entry name" value="WHEP-TRS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47323">
        <property type="entry name" value="Anticodon-binding domain of a subclass of class I aminoacyl-tRNA synthetases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47616">
        <property type="entry name" value="GST C-terminal domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57770">
        <property type="entry name" value="Methionyl-tRNA synthetase (MetRS), Zn-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52374">
        <property type="entry name" value="Nucleotidylyl transferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47060">
        <property type="entry name" value="S15/NS1 RNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00178">
        <property type="entry name" value="AA_TRNA_LIGASE_I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50405">
        <property type="entry name" value="GST_CTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51185">
        <property type="entry name" value="WHEP_TRS_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="11 14" id="KW-0030">Aminoacyl-tRNA synthetase</keyword>
    <keyword evidence="8 14" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6 14" id="KW-0436">Ligase</keyword>
    <keyword evidence="7 14" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="10 14" id="KW-0648">Protein biosynthesis</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0694">RNA-binding</keyword>
    <feature type="domain" description="GST C-terminal" evidence="15">
        <location>
            <begin position="67"/>
            <end position="194"/>
        </location>
    </feature>
    <feature type="domain" description="WHEP-TRS" evidence="16">
        <location>
            <begin position="843"/>
            <end position="899"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005594"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012838"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00018335"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022598"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022917"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023146"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00030904"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00047364"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="RuleBase" id="RU363039"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50405"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS51185"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015421.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="903" mass="101019" checksum="91D14D1F693F21C3" modified="2019-05-08" version="1">MPLNPPLSLGPGLLGLGSSALMPHAFPSAGHVVPFLTHPRVPALHLDSGTVLFSPNAICQYFFLLAGEEPNDLTNQWLEWEATELQPAVSRALYCHLVQGKKGGEVLGAIRKTLTYIDQSLTSKATPYLAGEAESVADIVLWATLFPVLRDESCLPDELQALRSWFQTMNLSEPCRSAAVSVLTPKGAQAFKVYLQKQPGPILHWEKPAANEPEEEELAERSLSEEEIAAAAEAWARGPAALPERWQPPSPVLPVEGRRNVLITSALPYVNNVPHLGNIIGCVLSADVFARYCRLRNWNTLYVCGTDEYGTATETKALEEGLTPQQICDKYNAIHVQIYRWFDISFDYFGRTTTTQQTTIAQDIFQRLLARNMLLTDTVDQLRCERCQRFLADRFVEGTCPFCNYEEARGDQCDKCGKLINAVELKKPQCKVCKDSPVVKSSQHLFLDLPQLEERLEKWLEQSWATGNWTANSRFITRSWIRDGLKPRCITRDLKWGTPVPLDGFRDKVFYVWFDAPIGYLSITANYTDQWERWWKNPQQVQLYNFMAKDNVPFHSVIFPCTLLGAEDNYTLVNHLVATEYLNYEDGKFSKSRGVGVFGDMAQDTGIPADIWRFYLLFLRPEGQDSAFSWTDLMLKNNSELLNNLGNFVNRAGMFVCKFFGGRVPPMELGADDKRLLAHVTLELRQYNQLLEKIRIRDALRCILSISRYGNQYIQVNEPWKRIKGHDADRTRAGTVTGVAVNIAALISVMLQPYMPGISATIQRQLCVPQACNVLTDTFLCLLPPGHQIGTVTPLFQKLENEQIEALRKQFGGGQVQAARVASADVTPPGPGGGAVLAGDPQKVKELAEEVAKQADHVRQLKASKAEKGQIDGEVAKLLELKKQLALAEGKTPELPAPKGKRK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKG6</accession>
    <name>A0A452HKG6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Type II GnRH receptor</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKG6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017613.1">
        <property type="protein sequence ID" value="ENSGAGP00000015418.1"/>
        <property type="gene ID" value="ENSGAGG00000011590.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004930">
        <property type="term" value="F:G protein-coupled receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042277">
        <property type="term" value="F:peptide binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016500">
        <property type="term" value="F:protein-hormone receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032870">
        <property type="term" value="P:cellular response to hormone stimulus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1070.10:FF:000199">
        <property type="entry name" value="Gonadotropin-releasing hormone II receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1070.10">
        <property type="entry name" value="Rhodopsin 7-helix transmembrane proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000276">
        <property type="entry name" value="GPCR_Rhodpsn"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017452">
        <property type="entry name" value="GPCR_Rhodpsn_7TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001658">
        <property type="entry name" value="GphnRH_fam_rcpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24241:SF69">
        <property type="entry name" value="GONADOTROPIN-RELEASING HORMONE II RECEPTOR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24241">
        <property type="entry name" value="NEUROPEPTIDE RECEPTOR-RELATED G-PROTEIN COUPLED RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00001">
        <property type="entry name" value="7tm_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00529">
        <property type="entry name" value="GNADOTRPHINR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00237">
        <property type="entry name" value="GPCRRHODOPSN"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81321">
        <property type="entry name" value="Family A G protein-coupled receptor-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50262">
        <property type="entry name" value="G_PROTEIN_RECEP_F1_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6" id="KW-0297">G-protein coupled receptor</keyword>
    <keyword evidence="7 12" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="9" id="KW-0675">Receptor</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0807">Transducer</keyword>
    <keyword evidence="4 12" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 12" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="30"/>
            <end position="49"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="69"/>
            <end position="89"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="110"/>
            <end position="130"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="161"/>
            <end position="184"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="218"/>
            <end position="242"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="254"/>
            <end position="277"/>
        </location>
    </feature>
    <feature type="domain" description="G-protein coupled receptors family 1 profile" evidence="13">
        <location>
            <begin position="9"/>
            <end position="274"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023224"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00082552"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50262"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015418.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="347" mass="38343" checksum="EF7A34AF85D689AC" modified="2019-05-08" version="1">MARLGYGMGGLWVLWAGAGGRRGKLSHVRVLLLHLAAADLLVTFVVMPLDAGWNLTVQWQAGDLACRLLMYLKLLAMYASAFITVVISLDRQAAILRPLAIAHAHRRNRLMLYAAWLLSAGLSLPQLFLFHTVTIRSPQNFTQCTTQGSFPQRWHETAYNMLGFACLFLLPLLIMVCCYARILLEISRRMGTSLFSKEVPLRRSTTNIPRARLRVLKLSLVIVSSFVVCWTPYYLLGLWYWFCPRAMEETVSPSLAHVLFIFGLLNACLDPLTYGLFTIPFHRGLGHCCRGGHGAGSEPCSPPTGSLRCSATSLHAKRGATATPVHETPAAQSYLGNGTSSVDSSYL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKG8</accession>
    <name>A0A452HKG8_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="8">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the GAS2 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000017620.1">
        <property type="protein sequence ID" value="ENSGAGP00000015423.1"/>
        <property type="gene ID" value="ENSGAGG00000011642.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005884">
        <property type="term" value="C:actin filament"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035371">
        <property type="term" value="C:microtubule plus-end"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001725">
        <property type="term" value="C:stress fiber"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008093">
        <property type="term" value="F:cytoskeletal anchor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008017">
        <property type="term" value="F:microtubule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051764">
        <property type="term" value="P:actin crosslink formation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001578">
        <property type="term" value="P:microtubule bundle formation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1904825">
        <property type="term" value="P:protein localization to microtubule plus-end"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031110">
        <property type="term" value="P:regulation of microtubule polymerization or depolymerization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd21268">
        <property type="entry name" value="CH_GAS2L1_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.418.10">
        <property type="entry name" value="Calponin-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.920.20">
        <property type="entry name" value="Gas2-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001715">
        <property type="entry name" value="CH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036872">
        <property type="entry name" value="CH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003108">
        <property type="entry name" value="GAR_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036534">
        <property type="entry name" value="GAR_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46756:SF18">
        <property type="entry name" value="GAS2-LIKE PROTEIN PICKLED EGGS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46756">
        <property type="entry name" value="TRANSGELIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00307">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02187">
        <property type="entry name" value="GAS2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00033">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00243">
        <property type="entry name" value="GAS2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47576">
        <property type="entry name" value="Calponin-homology domain, CH-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF143575">
        <property type="entry name" value="GAS2 domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50021">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51460">
        <property type="entry name" value="GAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="3" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Calponin-homology (CH)" evidence="6">
        <location>
            <begin position="26"/>
            <end position="154"/>
        </location>
    </feature>
    <feature type="domain" description="GAR" evidence="7">
        <location>
            <begin position="158"/>
            <end position="245"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="260"/>
            <end position="301"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="313"/>
            <end position="609"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="671"/>
            <end position="694"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="714"/>
            <end position="751"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="810"/>
            <end position="835"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="847"/>
            <end position="868"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="882"/>
            <end position="975"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="5">
        <location>
            <begin position="267"/>
            <end position="287"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="332"/>
            <end position="347"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="363"/>
            <end position="379"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="5">
        <location>
            <begin position="410"/>
            <end position="422"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="426"/>
            <end position="446"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="5">
        <location>
            <begin position="450"/>
            <end position="461"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="476"/>
            <end position="485"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="534"/>
            <end position="544"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="5">
        <location>
            <begin position="552"/>
            <end position="570"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="591"/>
            <end position="602"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="715"/>
            <end position="725"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="727"/>
            <end position="742"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="906"/>
            <end position="916"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00038441"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS51460"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015423.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="975" mass="107054" checksum="17C42CEEB12157F1" modified="2019-05-08" version="1">MADLSNIQSAASKSIRPFRSSEEYLYAMKEDLAEWLNILYGWDVRVENFMETLETGCDLCQHANNVNHIALEFQQQHPEVAAHMRVPQNEVVYQAKNVVPGSFIARDNVSNFIQWCRQDLGIQDVLMFETNDLVLKKNEKNFVLCLLEVARRGSKFGMLAPMLIQMEQEIEEEMRDQRAYGGLETQEASQDLGPESPVYPSRAQRISLCDLKNLDELVLRSHVMVRVGGGWDTLEHYLDKHDPCRCASLSHRLMQTRALGFSPQKTASPGSFSSSPRAASPSSQRPSEGMNPYRPTETLRMGDKRLSAGGNLTQAKMDRSGHPASGAAGDLSVQSDSSRQIPNVTSHVRQEKLEGLPSRAGTVPSSGQSSCSPLRSSTPLAHRDVSEPRASTTLRLQDQLPPRVRRLSGDSDSSASSTQSGTLGRRRGEDGALRTAHRKEVGRRMPEGAPRSQPSSRSQSQERGAFLGPRPSVGARRAEPEDRSRSRMVNGPGRPSPRARSQGRGSGEPVLLISRGKDGQHSWARANEVKENGQDSGRATSRTKSPARGHLARVSSRSPAPTSRRASLPANKLAEPSVRAPQQMLGSVRQRQVDETEPRVPVREQLSGETLQARELSESFSQELEELAQSFRTPLRLDPSQEQQLYRCLEEEFLANSQMIELAEDNGESLGCAHDPGRLQQTTPDPGAVDSAYCSSSSSSSSLNFFSKHSFQADFGERPKDDPKRNPASQRPWASNCSTGPTELQDGSGWDSHVALEPATCWRKPALSSSSDESNYYPALNDLQEVAEGVEVSGPGMDAAWSILELAGSDTDLGASAPDEPKAEEDSAEGPLGETSLDAMLDVLQTQEVXKTEEDSAEGPLGETSLDAMLDVLQTQEVVLRQKKSLKKPGRVPSIYKLKLRPKAKPRVDTQPDKKPSKIPTPLSYKTSGASPKGSPPKRPAEHKPWRAFHSVFSSFMEPPQGQMEPRGPDEDAWA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKI0</accession>
    <name>A0A452HKI0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">SLIT-ROBO Rho GTPase-activating protein 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="28 29">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="29" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="28" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="28" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="5">Cell projection</location>
            <location evidence="5">Dendritic spine</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cell projection</location>
            <location evidence="3">Lamellipodium</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cytoplasm</location>
            <location evidence="4">Cytosol</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasmic vesicle</location>
            <location evidence="2">Phagosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="18">Postsynaptic cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="19">Postsynaptic density</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKI0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017635.1">
        <property type="protein sequence ID" value="ENSGAGP00000015436.1"/>
        <property type="gene ID" value="ENSGAGG00000010110.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043197">
        <property type="term" value="C:dendritic spine"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030027">
        <property type="term" value="C:lamellipodium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045335">
        <property type="term" value="C:phagocytic vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0014069">
        <property type="term" value="C:postsynaptic density"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd04383">
        <property type="entry name" value="RhoGAP_srGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd11955">
        <property type="entry name" value="SH3_srGAP1-3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.555.10:FF:000010">
        <property type="entry name" value="SLIT-ROBO Rho GTPase-activating protein 1 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1270.60:FF:000006">
        <property type="entry name" value="SLIT-ROBO Rho GTPase-activating protein 1 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.40:FF:000266">
        <property type="entry name" value="SLIT-ROBO Rho GTPase-activating protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.60">
        <property type="entry name" value="Arfaptin homology (AH) domain/BAR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.555.10">
        <property type="entry name" value="Rho GTPase activation protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027267">
        <property type="entry name" value="AH/BAR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031160">
        <property type="entry name" value="F_BAR_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001060">
        <property type="entry name" value="FCH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000198">
        <property type="entry name" value="RhoGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051627">
        <property type="entry name" value="SLIT-ROBO_RhoGAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035648">
        <property type="entry name" value="srGAP1/2/3_SH3"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14166">
        <property type="entry name" value="SLIT-ROBO RHO GTPASE ACTIVATING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00611">
        <property type="entry name" value="FCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00620">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00018">
        <property type="entry name" value="SH3_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00055">
        <property type="entry name" value="FCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00324">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00326">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103657">
        <property type="entry name" value="BAR/IMD domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51741">
        <property type="entry name" value="F_BAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50238">
        <property type="entry name" value="RHOGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="16" id="KW-0966">Cell projection</keyword>
    <keyword evidence="12 22 23" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="9" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="17" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="7" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="13" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="14" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-0628">Postsynaptic cell membrane</keyword>
    <keyword evidence="29" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6 21" id="KW-0728">SH3 domain</keyword>
    <keyword evidence="11" id="KW-0770">Synapse</keyword>
    <feature type="domain" description="F-BAR" evidence="27">
        <location>
            <begin position="19"/>
            <end position="277"/>
        </location>
    </feature>
    <feature type="domain" description="Rho-GAP" evidence="26">
        <location>
            <begin position="460"/>
            <end position="643"/>
        </location>
    </feature>
    <feature type="domain" description="SH3" evidence="25">
        <location>
            <begin position="692"/>
            <end position="751"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="24">
        <location>
            <begin position="188"/>
            <end position="221"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="24">
        <location>
            <begin position="657"/>
            <end position="677"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="24">
        <location>
            <begin position="751"/>
            <end position="782"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="23">
        <location>
            <begin position="337"/>
            <end position="371"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="24">
        <location>
            <begin position="188"/>
            <end position="203"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="24">
        <location>
            <begin position="204"/>
            <end position="213"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="24">
        <location>
            <begin position="755"/>
            <end position="765"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004262"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004510"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004514"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004552"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023257"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00034100"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00034105"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00073507"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01077"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="26">
        <source>
            <dbReference type="PROSITE" id="PS50238"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="27">
        <source>
            <dbReference type="PROSITE" id="PS51741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="28">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015436.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="29">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="824" mass="93953" checksum="5ABE6801A8BA697F" modified="2019-05-08" version="1">MTSPAKFKKDKEIIAEYDTQVKEIRTQLIEQLKCLDQQCELRVQLLQDLQDFFRKKAEIEMDYSRNLEKLAERFLAKTRSTKDQQFKKDQNVLSPVNCWNLLLNQVKRESRDHTTLSDIYLNNIIPRFVQVSEDSGRLFKKSKEVGLQLQEDLMKVLNELYTVMKTYHMYNADSISAQSKLKEAEKQEEKQIGKSVKQEDKQTPRSPDSTSNVRFEEKHVRRSSVKKIEKMKEKRQAKYTENKLKAIKARNEYLLALEATNASVFKYYIHDLSDLIDVSVLLLGLMSFPLTPLCASPSLTPFVLTADVANSIMMYHNDFTGASFLQVSQLCAQQPVQSELVQRCQQLQSRLSTLKIENEEVKKTMEATLQTIQDIVTIEDFDVSDCFQYSNSMESVKSTVSETFMSKPSIAKRRANQQETEQFYFTKMKEYLEGRNLITKLQAKHDLLQKTLGESQRTDCSLDSSQAIPLVVESCIRFISRHGLQHEGIFRVSGSQVEVNDIKNAFERGEDPLAGDQNDHDMDSIAGVLKLYFRGLEHPLFPKEIFHDLIACVTMDNLQERALHIRKVLLSLPKTTLIVMRYLFAFLSHLSQFSEENMMDPYNLAICFGPTLMSVPEGHDQVSCQAHVNELIKTIIIQHENIFPGLRELEGPVYSRGGSTEDYCESPHGETASAEDSVQDIAAEHHTSDDECEPIEAIAKFDYIGRTARELSFKKGASLLLYQRASDDWWEGRHNGIDGLIPHQYIVVQDTEDGSSERSSPKSEIEINSEPPEEKVTARASANCPSGGHVADIYLANINNSSFLPLSPVPPPQFHIIPCPKMKI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKI2</accession>
    <name>A0A452HKI2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">aldehyde oxidase</fullName>
            <ecNumber evidence="5">1.2.3.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="16">
            <text>an aldehyde + O2 + H2O = a carboxylate + H2O2 + H(+)</text>
            <dbReference type="Rhea" id="RHEA:16829"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:15379"/>
            <dbReference type="ChEBI" id="CHEBI:16240"/>
            <dbReference type="ChEBI" id="CHEBI:17478"/>
            <dbReference type="ChEBI" id="CHEBI:29067"/>
            <dbReference type="EC" id="1.2.3.1"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 18">
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="19">
            <name>Mo-molybdopterin</name>
            <dbReference type="ChEBI" id="CHEBI:71302"/>
        </cofactor>
        <text evidence="19">Binds 1 Mo-molybdopterin (Mo-MPT) cofactor per subunit.</text>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="15">
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="19">
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:49601"/>
        </cofactor>
        <text evidence="19">Binds 2 [2Fe-2S] clusters.</text>
    </comment>
    <comment type="subunit">
        <text evidence="4">Homodimer.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the xanthine dehydrogenase family.</text>
    </comment>
    <dbReference type="EC" id="1.2.3.1" evidence="5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017631.1">
        <property type="protein sequence ID" value="ENSGAGP00000015432.1"/>
        <property type="gene ID" value="ENSGAGG00000010622.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051537">
        <property type="term" value="F:2 iron, 2 sulfur cluster binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004031">
        <property type="term" value="F:aldehyde oxidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071949">
        <property type="term" value="F:FAD binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005506">
        <property type="term" value="F:iron ion binding"/>
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        <property type="entry name" value="Uridine Diphospho-n-acetylenolpyruvylglucosamine Reductase, domain 2"/>
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        <property type="entry name" value="Fer2"/>
        <property type="match status" value="1"/>
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    </dbReference>
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    </dbReference>
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    </dbReference>
    <proteinExistence type="inferred from homology"/>
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    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11 18" id="KW-0274">FAD</keyword>
    <keyword evidence="8" id="KW-0285">Flavoprotein</keyword>
    <keyword evidence="13 19" id="KW-0408">Iron</keyword>
    <keyword evidence="14 19" id="KW-0411">Iron-sulfur</keyword>
    <keyword evidence="10 19" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7 19" id="KW-0500">Molybdenum</keyword>
    <keyword evidence="12" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
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        <location>
            <begin position="231"/>
            <end position="416"/>
        </location>
    </feature>
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        <location>
            <position position="1260"/>
        </location>
    </feature>
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        <location>
            <position position="40"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
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            <label>1</label>
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        <location>
            <position position="45"/>
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            <name>[2Fe-2S] cluster</name>
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            <label>1</label>
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        <location>
            <position position="48"/>
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        <location>
            <position position="70"/>
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            <name>[2Fe-2S] cluster</name>
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            <label>1</label>
        </ligand>
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        <location>
            <position position="110"/>
        </location>
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            <name>[2Fe-2S] cluster</name>
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            <label>2</label>
        </ligand>
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        <location>
            <position position="113"/>
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            <name>[2Fe-2S] cluster</name>
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            <label>2</label>
        </ligand>
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        <location>
            <position position="145"/>
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            <name>[2Fe-2S] cluster</name>
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            <label>2</label>
        </ligand>
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        <location>
            <position position="147"/>
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            <name>[2Fe-2S] cluster</name>
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            <label>2</label>
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        <location>
            <begin position="259"/>
            <end position="266"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
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        <location>
            <position position="362"/>
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        </ligand>
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        <location>
            <position position="424"/>
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            <name>FAD</name>
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        </ligand>
    </feature>
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        <location>
            <position position="766"/>
        </location>
        <ligand>
            <name>Mo-molybdopterin</name>
            <dbReference type="ChEBI" id="CHEBI:71302"/>
        </ligand>
        <ligandPart>
            <name>Mo</name>
            <dbReference type="ChEBI" id="CHEBI:28685"/>
        </ligandPart>
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        <location>
            <position position="797"/>
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            <name>Mo-molybdopterin</name>
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            <name>Mo</name>
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        <location>
            <position position="911"/>
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        </ligandPart>
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        <ligand>
            <name>substrate</name>
        </ligand>
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            <position position="1078"/>
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            <name>Mo</name>
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        </ligandPart>
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        <source>
            <dbReference type="ARBA" id="ARBA00001974"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00006849"/>
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            <dbReference type="ARBA" id="ARBA00013041"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00034078"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00047679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="PIRSR" id="PIRSR000127-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="PIRSR" id="PIRSR000127-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS51387"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015432.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1316" mass="145105" checksum="643F713C797342B4" modified="2019-05-08" version="1">MACSTQSDALVFFVNGRKNQQLFSLTAAFAVRLSGTKYGCGIGGCGACTVMISTYEPTSKKIMHYPANSCLLPICSLHSSAVTTVEGVGSTKTRVHPIQERLAKCHGSQCGFCTPGMVMSMYALLRNHPEPSMEQITAALDGNLCRCTGYRPIIDTYKSFTGETACCQLRGTGQCCLDKEDVCSSSDKGNQMHSGLCNPEEFQPVDPTQEFIFPPELMRMAQEQQKRTLVFRGERTTWISPTNLRELLELKAMYPKAPLIVGNTSVGLEMKRDGVHHPVFIHPTRIPDLHVVSNTNNGLVIGAACSLAQLRDILMKVVPELPEEKTKIFRALLQQLKTLAGEQIRSLASLGGHIVSRGSTWDLNPVLAVGNSVLSLASKDGTRQIPLNDQFLAGYENADLKPKEIIVSVFIPYSKKDDFVSAFRQAERRRNAVSVVNSGMRVLFKPNTDIIMDMNILYGGTGPITVSARKSCQRLIGRQWNEQMLDEACKLVLGEISLSPSAPGGKVEYRRTLLVSFLFKFYLEVLYGLNKMYPAKFPDLSKKNLSALGAFHSEVPQGMQTYQDVDPVQPPQDPVGRPIMHQSGVKHATGEAVYCNDIRAVDEELFLAVVTSSRAHAKIVTIDVSEALKVPGVVDVVTAEDVPGKNGTDDEQAYAKDEVICVGQIVCAVVAESAVQAKQGVEKMKIEYEDLEPILTIEDAIKHNSYFGKEKKIEQGNIEEGFKSADEIIEGEIHIGGQEHFYLETNSVLVVPRGEDNEMDVHVSTQDAARVQEFVALALDVQSSKIMCHTKRVGGAFGGKITKPAIFAVVAAVAANKTGRPIRFTLERDEDMLITGGRHPFFGKYKVGFTNDGRITAADIEPYINAGCTLDDSEMVIEYVVLKTDNAYKIPNLRIRGRACKTNLPSNTAFRGFGFPQSGLFTESWITTVATQTGLPQEKIREINMYKEVNLTPFKEELDATNLVKCWKECMEKSEYYSRKTAVEEFNKQNYWKKKGIAIIPMKFSVGFNATQYHQAAALVHIYTDGSVLVTHGGIELGQGIHTKMLQIASRELKVPLSYVHFCETSTITVPNAIVTAGSIGSDVNGKAVQNACQILRKRLEPIINKNPEGKWEDWITEAYDQSVSLSATGYFRGYMVNMNWDTGEGQAFPYFVFGAACSEVEIDCLTGDHKNIRTDIVMDACFSINPAVDIGQIEGAFVQGVGLYTMEEIKFSPEGEQYTLGPDTYKIPAVCDIPEQFRVYLLPNSRNPIAIYSAKGIGEAGLFLGCSVFFAIWDAVAAVRKERGLTGTFTLNSPLTIERIRMACADNFTEMVMVF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKI6</accession>
    <name>A0A452HKI6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Protein FAM13B</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the FAM13 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKI6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017633.1">
        <property type="protein sequence ID" value="ENSGAGP00000015434.1"/>
        <property type="gene ID" value="ENSGAGG00000011649.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd04393">
        <property type="entry name" value="RhoGAP_FAM13A1a"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.555.10:FF:000020">
        <property type="entry name" value="protein FAM13B isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.555.10">
        <property type="entry name" value="Rho GTPase activation protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039102">
        <property type="entry name" value="FAM13"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR059029">
        <property type="entry name" value="FAM13A_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000198">
        <property type="entry name" value="RhoGAP_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15904">
        <property type="entry name" value="FAM13"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15904:SF16">
        <property type="entry name" value="PROTEIN FAM13B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26116">
        <property type="entry name" value="FAM13A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00620">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00324">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50238">
        <property type="entry name" value="RHOGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Rho-GAP" evidence="5">
        <location>
            <begin position="23"/>
            <end position="212"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="243"/>
            <end position="263"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="336"/>
            <end position="361"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="407"/>
            <end position="428"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="492"/>
            <end position="516"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="628"/>
            <end position="674"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="811"/>
            <end position="830"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="4">
        <location>
            <begin position="243"/>
            <end position="253"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="254"/>
            <end position="263"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="338"/>
            <end position="361"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="414"/>
            <end position="428"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="640"/>
            <end position="652"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="664"/>
            <end position="674"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="811"/>
            <end position="821"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007549"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00074176"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50238"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015434.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="921" mass="105874" checksum="C1F6DE757BE95F13" modified="2019-05-08" version="1">MRKSSSPSLSNCNSVLANKIFGIPLDELQQEGQPDNEVPFIVRHVVDYIEEHGGLEQEGLFQVNGNAETVEWLRKRYDNGEDVDLVKEADVPSAISLLRFFLQELPEPVIPGSLHIHLMQFSQDYNNEDEFGRKLRFLLQQLPPVNYSLLKFLCRFLANVASHHEEIWSASSLAAVFGPDVFHIYTDVEDLKEQEIVSRIMAGLLENYYEFFENEEEDFSSTNDLSSITEQINDLLEEEEDIKLEQTEELPEDSTEKPEERPDMVHGMTESILDSSSVTASTSAHLSPISILPASADILERTIRAAVEQHLFDLQSSIDDDLKNLQQHRLVCNNEAKNINRDEEGSNNKDDVTKDNDINSSENTRDCFETLACTDLDSAVLQPDSVYEEEENDQTVGILLKPCNEDAEVDEENNSERQDCESVDDNDKMSSQLVLDSSSKICDLNANTESEISGDGSVNVPEESTCIQVPHLDLKNVSDGNKWEEPFPAFKSWQEDSESGEAQLSPQAGRMNSHPLEDCHPVLSHRSLDFGQSQRFLHDPETLDFSSKALSFVRTRRASFSSKDDRREDRTPYQLVKKLQKKIRQFEEQFEKDKNSKPSYSDIAANPKVLKWMTELTKLRKQIKDEKHKCSDGEFIPQTRPRSNTLPKSFGSSLEHEDEENEDEPRVMQKEKKPTKEATLELILKRLKEKRVERCLPEDIKKMTKDHLAEEKTSLQKSLLYYESQHGRPVTREERHIVKPLYDRYRLVKQMLTRASITPILGSPSTKRRSQMLQPIIEGETAHFFEEIKEEEEDVDSLSSEFNDILKTAVETQSISSPAENSESDLEDSQEKLTRDLRLSSIRAASMPELLEQLWKARAEKKKLRKTLREFEEGFYQQNGRNVQKEDRAPMFDEYREYKKIKAKLRLLEVLISKQDFSKSI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKJ7</accession>
    <name>A0A452HKJ7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Agouti-related protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="8">Plays a role in weight homeostasis. Involved in the control of feeding behavior through the central melanocortin system. Acts as alpha melanocyte-stimulating hormone antagonist by inhibiting cAMP production mediated by stimulation of melanocortin receptors within the hypothalamus and adrenal gland. Has very low activity with MC5R. Is an inverse agonist for MC3R and MC4R being able to suppress their constitutive activity. It promotes MC3R and MC4R endocytosis in an arrestin-dependent manner.</text>
    </comment>
    <comment type="subunit">
        <text evidence="9">Interacts with melanocortin receptors MC3R, MC4R and MC5R.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="7">Golgi apparatus lumen</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="11">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKJ7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017652.1">
        <property type="protein sequence ID" value="ENSGAGP00000015452.1"/>
        <property type="gene ID" value="ENSGAGG00000011650.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005796">
        <property type="term" value="C:Golgi lumen"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005184">
        <property type="term" value="F:neuropeptide hormone activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070996">
        <property type="term" value="F:type 1 melanocortin receptor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008343">
        <property type="term" value="P:adult feeding behavior"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009755">
        <property type="term" value="P:hormone-mediated signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007218">
        <property type="term" value="P:neuropeptide signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:2000253">
        <property type="term" value="P:positive regulation of feeding behavior"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.760.10:FF:000003">
        <property type="entry name" value="Agouti-related peptide 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.760.10">
        <property type="entry name" value="Agouti domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007733">
        <property type="entry name" value="Agouti"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027300">
        <property type="entry name" value="Agouti_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036836">
        <property type="entry name" value="Agouti_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16551">
        <property type="entry name" value="AGOUTI RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16551:SF4">
        <property type="entry name" value="AGOUTI-RELATED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05039">
        <property type="entry name" value="Agouti"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00792">
        <property type="entry name" value="Agouti"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57055">
        <property type="entry name" value="Agouti-related protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS60024">
        <property type="entry name" value="AGOUTI_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51150">
        <property type="entry name" value="AGOUTI_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6 11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="4" id="KW-0960">Knottin</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="3 12" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="12">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019076094" description="Agouti-related protein" evidence="12">
        <location>
            <begin position="21"/>
            <end position="153"/>
        </location>
    </feature>
    <feature type="domain" description="Agouti" evidence="13">
        <location>
            <begin position="109"/>
            <end position="150"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="11">
        <location>
            <begin position="109"/>
            <end position="124"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="11">
        <location>
            <begin position="116"/>
            <end position="130"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="11">
        <location>
            <begin position="123"/>
            <end position="141"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="11">
        <location>
            <begin position="132"/>
            <end position="139"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022854"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023769"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00056588"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00065157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00068128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00494"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS51150"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015452.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="153" mass="17069" checksum="F61A17EFF96355E1" modified="2019-05-08" version="1">MLNMLLLSWGVLQGIQTILTSDLSSNHLSEMHRSLDGADRSSYPSLLRKVKEAPMGPTGTLPRPGFDRMAMELRVADEDLIKEASVLEPEVLSAALEAQGREERSPRRCVRLLESCLGHQLPCCDPCATCYCRFFNAFCYCRKINATFPCGKN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKM1</accession>
    <name>A0A452HKM1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">C2H2-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="10">Belongs to the Ikaros C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKM1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017661.1">
        <property type="protein sequence ID" value="ENSGAGP00000015461.1"/>
        <property type="gene ID" value="ENSGAGG00000011670.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003700">
        <property type="term" value="F:DNA-binding transcription factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002376">
        <property type="term" value="P:immune system process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000073">
        <property type="entry name" value="IKAROS family zinc finger 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000265">
        <property type="entry name" value="IKAROS family zinc finger 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000525">
        <property type="entry name" value="IKAROS family zinc finger 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000080">
        <property type="entry name" value="IKAROS family zinc finger 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000168">
        <property type="entry name" value="zinc finger protein Eos isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050589">
        <property type="entry name" value="Ikaros_C2H2-ZF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24404:SF36">
        <property type="entry name" value="DNA-BINDING PROTEIN IKAROS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24404">
        <property type="entry name" value="ZINC FINGER PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="2" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="8" id="KW-0804">Transcription</keyword>
    <keyword evidence="6" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <keyword evidence="4 11" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="115"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="143"/>
            <end position="170"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="171"/>
            <end position="198"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="199"/>
            <end position="227"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1"/>
            <end position="49"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="335"/>
            <end position="358"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00038390"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015461.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="471" mass="52388" checksum="EA5691734E21BD73" modified="2019-05-08" version="1">METDEAQDMSQVSGKESPPISDVPDDADEPMPVPEDLSTSTGGQQKNDRVLASNIKIETQSDEENGRACEMNGEECAEDLRMLDASGEKMNGSHNGPGSKAMSGVGGIRLPNGKLKCDICGIICIGPNVLMVHKRSHTGERPFHCTQCGASFTQKGNLLRHIKLHSGEKPFKCHLCNYACRRRDALTGHLRTHSVGKPHKCGYCGRSYKQRSSLEEHKERCHNYLQTMSLSGSLYPGEKCLSDVPYDATTNYEKESEMMQTHVIDQAINNAISYLGAESLRPLVQTPPGSSEAAPIISSMYQLHKPLGDNHSRSNHTAQDTAVENLLLLSKAKSISSERDASPSNSCQDSTDTESNNEERSGLIYLTNHIVPHARNGLSIKEEQRQYEVLRAGTDNSHDAFKVITSNGEQVKVYKCEHCRVLFLDHVMYTIHMGCHGFRDPFECNMCGYHSQDRYEFSSHITRGEHRFHMS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INF1</accession>
    <name>A0A452INF1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">glutamate carboxypeptidase II</fullName>
            <ecNumber evidence="18">3.4.17.21</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>Release of an unsubstituted, C-terminal glutamyl residue, typically from Ac-Asp-Glu or folylpoly-gamma-glutamates.</text>
            <dbReference type="EC" id="3.4.17.21"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Membrane</location>
            <topology evidence="2">Single-pass type II membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the peptidase M28 family. M28B subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.4.17.21" evidence="18"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033484.1">
        <property type="protein sequence ID" value="ENSGAGP00000029488.1"/>
        <property type="gene ID" value="ENSGAGG00000020963.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016805">
        <property type="term" value="F:dipeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004181">
        <property type="term" value="F:metallocarboxypeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd08022">
        <property type="entry name" value="M28_PSMA_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.930.40:FF:000001">
        <property type="entry name" value="N-acetylated-alpha-linked acidic dipeptidase 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.630.10:FF:000009">
        <property type="entry name" value="N-acetylated-alpha-linked acidic dipeptidase 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.50.30.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.930.40">
        <property type="entry name" value="Transferrin receptor-like, dimerisation domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.630.10">
        <property type="entry name" value="Zn peptidases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046450">
        <property type="entry name" value="PA_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007484">
        <property type="entry name" value="Peptidase_M28"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039373">
        <property type="entry name" value="Peptidase_M28B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007365">
        <property type="entry name" value="TFR-like_dimer_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036757">
        <property type="entry name" value="TFR-like_dimer_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10404:SF36">
        <property type="entry name" value="GLUTAMATE CARBOXYPEPTIDASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10404">
        <property type="entry name" value="N-ACETYLATED-ALPHA-LINKED ACIDIC DIPEPTIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04389">
        <property type="entry name" value="Peptidase_M28"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04253">
        <property type="entry name" value="TFR_dimer"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52025">
        <property type="entry name" value="PA domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47672">
        <property type="entry name" value="Transferrin receptor-like dimerisation domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53187">
        <property type="entry name" value="Zn-dependent exopeptidases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0121">Carboxypeptidase</keyword>
    <keyword evidence="12" id="KW-0224">Dipeptidase</keyword>
    <keyword evidence="15" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="14" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13" id="KW-0482">Metalloprotease</keyword>
    <keyword evidence="16" id="KW-0511">Multifunctional enzyme</keyword>
    <keyword evidence="5" id="KW-0645">Protease</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0735">Signal-anchor</keyword>
    <keyword evidence="6" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="9" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="Peptidase M28" evidence="20">
        <location>
            <begin position="325"/>
            <end position="436"/>
        </location>
    </feature>
    <feature type="domain" description="Transferrin receptor-like dimerisation" evidence="19">
        <location>
            <begin position="592"/>
            <end position="712"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004606"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005634"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022645"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022997"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023049"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023268"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00052003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00066561"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="Pfam" id="PF04253"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="Pfam" id="PF04389"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029488.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="715" mass="80336" checksum="A74537A0C06E8C5A" modified="2019-05-08" version="1">GIIQCLAHCGLPKTFPGGPPDSDALSXKGNNPRPLFPCYFLPGSPPWTTLGDSLLPKLTSNYKIQFLVLTANTCPFLFPSNFTQRPHLAGTAENLRLAQQIQAQWKEFGLDSVELAHYDVLLSYPNETNPNYISIIDEHGTEIFNTSLFEPPPPGYETVTDVVPPYSAFSAPGVPEVKSNRLLKSRKLKVSCSQLGLIFRGSWKIRPHKLYSPGNFQLMRFCRTAAGHLEHGSAKINTSLIPPEYTYRYEEARAVGLPKLPVHPIGYNDAEKLLRDMGGPAPPDSSWKGSLNVSYNIGPGFMSRYSTRKVKMHIHSSNKVTRIYNVIGTIRGATEPDRYVILGGHRDSWVFGGIDPQSGAAVVHEIVRSYGKLKKEGWRPRRTIIFASWDAEEFGLLGSTEWAEENTKLLQARGVAYINADSSVEGNYTLRIDCTPLMYSLVYSLTREIPSPDEGFKGKSLYESWHENNPSTEYKDVPRINKLGSGNDFEVFFQRLGIASGRARYSKNWVSLSYPVYHSVYETYEIVERFYDPTFKNHLTVAQVRGGLVFELADSVVLPFDCRDYAEALRNYVQVIYNLSKNHQMGLTTYSVSFDSLFSAANNFTEAATDFHRRLQQVDINNPIAVRALNDQLMFLERAFIDPLGLPGRPFYRHIIFAPSSHNKYAGESFPGIYDAMFDIENKADQRKAWEEVKRQISIAAFTVQAAAGTLKDLA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INF2</accession>
    <name>A0A452INF2_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="11">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the RIN (Ras interaction/interference) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452INF2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033380.1">
        <property type="protein sequence ID" value="ENSGAGP00000029394.1"/>
        <property type="gene ID" value="ENSGAGG00000021249.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030139">
        <property type="term" value="C:endocytic vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031267">
        <property type="term" value="F:small GTPase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016192">
        <property type="term" value="P:vesicle-mediated transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd16131">
        <property type="entry name" value="RA_Rin2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd10394">
        <property type="entry name" value="SH2_RIN2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1050.80:FF:000002">
        <property type="entry name" value="Ras and Rab interactor 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.505.10:FF:000052">
        <property type="entry name" value="Ras and Rab interactor 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.505.10">
        <property type="entry name" value="SH2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1050.80">
        <property type="entry name" value="VPS9 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000159">
        <property type="entry name" value="RA_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035868">
        <property type="entry name" value="RIN2_SH2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000980">
        <property type="entry name" value="SH2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036860">
        <property type="entry name" value="SH2_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029071">
        <property type="entry name" value="Ubiquitin-like_domsf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003123">
        <property type="entry name" value="VPS9"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045046">
        <property type="entry name" value="Vps9-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037191">
        <property type="entry name" value="VPS9_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23101">
        <property type="entry name" value="RAB GDP/GTP EXCHANGE FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23101:SF51">
        <property type="entry name" value="RAS AND RAB INTERACTOR 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00788">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23268">
        <property type="entry name" value="RIN1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02204">
        <property type="entry name" value="VPS9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00314">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00252">
        <property type="entry name" value="SH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00167">
        <property type="entry name" value="VPS9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55550">
        <property type="entry name" value="SH2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54236">
        <property type="entry name" value="Ubiquitin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109993">
        <property type="entry name" value="VPS9 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50200">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50001">
        <property type="entry name" value="SH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51205">
        <property type="entry name" value="VPS9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="3" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 6" id="KW-0727">SH2 domain</keyword>
    <feature type="domain" description="SH2" evidence="8">
        <location>
            <begin position="61"/>
            <end position="154"/>
        </location>
    </feature>
    <feature type="domain" description="VPS9" evidence="10">
        <location>
            <begin position="581"/>
            <end position="720"/>
        </location>
    </feature>
    <feature type="domain" description="Ras-associating" evidence="9">
        <location>
            <begin position="750"/>
            <end position="841"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="176"/>
            <end position="197"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="247"/>
            <end position="325"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="421"/>
            <end position="445"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="7">
        <location>
            <begin position="182"/>
            <end position="193"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="247"/>
            <end position="262"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="7">
        <location>
            <begin position="269"/>
            <end position="278"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="279"/>
            <end position="302"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006919"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022999"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00191"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50001"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50200"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS51205"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029394.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="857" mass="96950" checksum="13E424F98FD0C502" modified="2019-05-08" version="1">MGPRKKEQMHQQAPKRGLVKDMDNVCPEKKDLKVCPRDSGYDSLSNKLSILDKLLHTHPVWLQLGLNDAEAVEILQSQPPGIFLVRKSTRMQKKVLSLRLPNEFGSRLKEFAIKESTYTFSLEGSGISFADLFRLIAFYCISRDVLPFTLKLPHAIAAAKTEAELEDIAQLGLNFWNSPANNNPPNPSPPRRPLPSDNACKDSRQLCLINGVHSIRTRTPSELECSQTNGALCFINPLFLKVHSQDVSGSLKRQSPRTQDVNGTEKPRSPPPRPPPPSINSLLTSPQLSRTIKQGSMPGTVNRNKHRNLDLLQKKPTPVPPPRLKKQAVCSEGEDNAKTLAVIRSSHNSVHGSEAASVPGEALCEQNLAACRKTLATNSESQVQWNGGRQRLSDMSISTSSSDSLDFDRSMPLFGYEGDTNSSLEDFEGESDQESMAPPLKPKKKRNSSFVLPKIVKSQLRKVSGVFSSFMTPEKRMIKKIAELSQDKRTYFGCLVQDYVSFLQENKECHVSSTDMLQTIRQFMTQVKNYLSQSSELDPPIESLIPEDQIDVVLEKAMHKCILKPLKGHIEAMLKEFHTVDGSWKQLKENLQLVRQRNPQELGVYVPTPDFVDVEKIKVKFMTMQKMYSPEKKVMLLLRVCKLIYTVMENNSGRMYGADDFLPVLTYVVAQCDMLELDTEIEYMMELLDPSLLHGEGGYYLTSAYGALSLIKNFQEEQAARLLSSEARDTLRQWHKRRTANRTIPSVDDFQNYLRVAFQEVNSGCTGKTLLVRPYITTEDVCQLCAEKFKVDSPEDYSLFLFIDDTWQQLTEDTYPQKIKAELHSRPQPQVFHFVYKRINSDPYGAIFQNNDDNSTL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INF3</accession>
    <name>A0A452INF3_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="3">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000033381.1">
        <property type="protein sequence ID" value="ENSGAGP00000029395.1"/>
        <property type="gene ID" value="ENSGAGG00000020783.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005938">
        <property type="term" value="C:cell cortex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903479">
        <property type="term" value="P:mitotic actomyosin contractile ring assembly actin filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd23767">
        <property type="entry name" value="IQCD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12207">
        <property type="entry name" value="RasGAP_IQGAP3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.506.10:FF:000004">
        <property type="entry name" value="IQ motif containing GTPase activating protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.190">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.506.10">
        <property type="entry name" value="GTPase Activation - p120gap, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000048">
        <property type="entry name" value="IQ_motif_EF-hand-BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000593">
        <property type="entry name" value="RasGAP_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023152">
        <property type="entry name" value="RasGAP_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001936">
        <property type="entry name" value="RasGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001202">
        <property type="entry name" value="WW_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149">
        <property type="entry name" value="RAS GTPASE-ACTIVATING PROTEIN WITH IQ MOTIF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149:SF10">
        <property type="entry name" value="RAS GTPASE-ACTIVATING-LIKE PROTEIN IQGAP3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00612">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00616">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03836">
        <property type="entry name" value="RasGAP_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00015">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00323">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF143885">
        <property type="entry name" value="RGC domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00509">
        <property type="entry name" value="RAS_GTPASE_ACTIV_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50018">
        <property type="entry name" value="RAS_GTPASE_ACTIV_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01159">
        <property type="entry name" value="WW_DOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50020">
        <property type="entry name" value="WW_DOMAIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="WW" evidence="2">
        <location>
            <begin position="416"/>
            <end position="449"/>
        </location>
    </feature>
    <feature type="domain" description="Ras-GAP" evidence="1">
        <location>
            <begin position="740"/>
            <end position="973"/>
        </location>
    </feature>
    <evidence type="ECO:0000259" key="1">
        <source>
            <dbReference type="PROSITE" id="PS50018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="PROSITE" id="PS50020"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029395.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1390" mass="158706" checksum="507544F3FEB6E30E" modified="2019-05-08" version="1">MCNGGWLFPLQYLQESPESEDIYDRCLTQAEIQGNINKVNVHGALEYVDDALERQDPLALCQALQDPVLALRGVQRENAAWYLDQLSTDREQKALELGYVDLLEQEEVQAGVYAANERGDQEHAMRRAVSRINASIRRGEAMETVRELMSPEAQLPKAFPFAAQLYQHELSLLQQHHPQGELGQEELYVAVEMLSAVALIDRALEAGDVSGFWSSLVSPALGLSDIEDANAQRYFDDLLKLKRRSGKAGVAFLSWNDIQACVNSANLAVQDENDKILAIRLINEALAQDDPEKTLTALLLPSARFSAVTLPGARRYHDVLTAALKQKAQVTGDDGAALWWEEIQQGVSKANQDTEAARRLALGIAAINQAIKEGKPAQTARVLRNPDVSLRGIVAECAGAYQAQLAALLAAKRQAGNANHNWVRHQLKDGSVYYFSLQNFEGSWERPHDCVLNSTHLSREEIQSTITGVTAAHDRERLWESNAGFVVRLQARMRGYLVRQEFVARRHLLQEQLPAVIKIQSCWRGYRQRRDYLQRLRYLQKTRDAVIKIQTYVRMWRARKRYRERLRYFRKNINAVIKIQAFVRANKARGDYRMLVHARDPPLSIVRRFVHLLEQSQHDFWEESELLKLREEVVKRIRSNQQLESDLNLMDIKIGLLVKNRITLQEVVSHCKKLTRKNKEQLSNLMAIDKQKGLKSLSKEKRQKLEAYQHLFYLLQTHPVYLAKLIFQMPQNKSTKFMESVVFTLYNYASNPREAYLLLQLFKTALQEEIRSKVDQLRDILTGNPTVIRLVVSFYRNARGQNALRQILGSPVQEVLQDKTLSIRTNPVDIYKAWINQLESQSGQKSKFPYDVSPEQALSHPEVQRRLDISIRNLLAVTDKFLSAITSSVDKIPYGMRYVAKVLKTSLAEKFPAAGEEDICKIVGNLLYYRFMNPAVVAPDGFDIVDISAGVALHPEHRRNLGSIAKILQHAAANKMFEGENGHLRVVNQYLEETHSKFRKFICAACSVPEPEERFNVDEYSEMVAVAKPVIYITVAELINTHKLLLEHQDSISPDHKDPLHELLEDLDELPTVQSLIGESAPNPGGDSPEQALAQLGKQEVSLTLTNKFDAVEASSEEADTRSLLLSTKQMLVDVIQSQPGDSLLEILRTAASEDQEAFHHHQMHRRALREAQTPDRLKRNRSLAGNSQLSMEEKKRKVIRNLRRLENLGVVDSANQYQEIINEMAKDIRNQRRYRQHRKGELVKLRQTLCGLNSKTSFYEEQIDYYNQYIRTCLDNLAASSKLNGKSKKQQSLHYTAARLFEKGVLLEIEDLPVSQFRNVIFDIIPCGEAGKFQVKAKFMGIDMEKFQLHYQDLLQLQYEGVAVMKLFDKAKVNVNLLIFLLNKKFFRK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INF5</accession>
    <name>A0A452INF5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Folate hydrolase 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the peptidase M28 family. M28B subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452INF5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033416.1">
        <property type="protein sequence ID" value="ENSGAGP00000029427.1"/>
        <property type="gene ID" value="ENSGAGG00000020963.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016805">
        <property type="term" value="F:dipeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004181">
        <property type="term" value="F:metallocarboxypeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd08022">
        <property type="entry name" value="M28_PSMA_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.50.30.30">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.930.40">
        <property type="entry name" value="Transferrin receptor-like, dimerisation domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.630.10">
        <property type="entry name" value="Zn peptidases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046450">
        <property type="entry name" value="PA_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007484">
        <property type="entry name" value="Peptidase_M28"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039373">
        <property type="entry name" value="Peptidase_M28B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007365">
        <property type="entry name" value="TFR-like_dimer_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036757">
        <property type="entry name" value="TFR-like_dimer_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10404:SF36">
        <property type="entry name" value="GLUTAMATE CARBOXYPEPTIDASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10404">
        <property type="entry name" value="N-ACETYLATED-ALPHA-LINKED ACIDIC DIPEPTIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04389">
        <property type="entry name" value="Peptidase_M28"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04253">
        <property type="entry name" value="TFR_dimer"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52025">
        <property type="entry name" value="PA domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47672">
        <property type="entry name" value="Transferrin receptor-like dimerisation domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53187">
        <property type="entry name" value="Zn-dependent exopeptidases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0224">Dipeptidase</keyword>
    <keyword evidence="9" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="5" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8" id="KW-0482">Metalloprotease</keyword>
    <keyword evidence="10" id="KW-0511">Multifunctional enzyme</keyword>
    <keyword evidence="3" id="KW-0645">Protease</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="Peptidase M28" evidence="12">
        <location>
            <begin position="266"/>
            <end position="468"/>
        </location>
    </feature>
    <feature type="domain" description="Transferrin receptor-like dimerisation" evidence="11">
        <location>
            <begin position="536"/>
            <end position="656"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005634"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022997"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023049"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023268"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF04253"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF04389"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029427.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="15">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="659" mass="74510" checksum="84FF8066053C15C4" modified="2025-10-08" version="2">MDFKSCDENTAVLSLRSVLSPVGCSCSSWFTKPTDEVTATSFHEKVKKAFMAEMKAENIKQFLYNFTQRPHLAGTAENLRLAQQIQAQWKEFGLDSVELAHYDVLLSYPNETNPNYISIIDEHGTEIFNTSLFEPPPPGYETVTDVVPPYSAFSAPGVPEVKRLELLNSRLPALSKINTSLIPPEYTYRYEEARAVGLPKLPVHPIGYNDAEKLLRDMGGPAPPDSSWKGSLNVSYNIGPGFMSRYSTRKVKMHIHSSNKVTRIYNVIGTIRGATEPDRYVILGGHRDSWVFGGIDPQSGAAVVHEIVRSYGKLKKEGWRPRRTIIFASWDAEEFGLLGSTEWAEENTKLLQARGVAYINADSSVEGNYTLRIDCTPLMYSLVYSLTREIPSPDEGFKGKSLYESWHENNPSTEYKDVPRINKLGSGNDFEVFFQRLGIASGRARYSKNWRVEKYSSYPVYHSVYETYEIVERFYDPTFKNHLTVAQVRGGLVFELADSVVLPFDCRDYAEALRNYVQVIYNLSKNHQMGLTTYSVSFDSLFSAANNFTEAATDFHRRLQQVDINNPIAVRALNDQLMFLERAFIDPLGLPGRPFYRHIIFAPSSHNKYAGESFPGIYDAMFDIENKADQRKAWEEVKRQISIAAFTVQAAAGTLKDLA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INF6</accession>
    <name>A0A452INF6_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452INF6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033489.1">
        <property type="protein sequence ID" value="ENSGAGP00000029493.1"/>
        <property type="gene ID" value="ENSGAGG00000021299.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005847">
        <property type="term" value="C:mRNA cleavage and polyadenylation specificity factor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031124">
        <property type="term" value="P:mRNA 3'-end processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:000778">
        <property type="entry name" value="WD repeat domain 33"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045245">
        <property type="entry name" value="Pfs2-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036322">
        <property type="entry name" value="WD40_repeat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001680">
        <property type="entry name" value="WD40_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22836:SF0">
        <property type="entry name" value="PRE-MRNA 3' END PROCESSING PROTEIN WDR33"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22836">
        <property type="entry name" value="WD40 REPEAT PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00400">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00320">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50978">
        <property type="entry name" value="WD40 repeat-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50082">
        <property type="entry name" value="WD_REPEATS_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50294">
        <property type="entry name" value="WD_REPEATS_REGION"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0853">WD repeat</keyword>
    <feature type="repeat" description="WD" evidence="1">
        <location>
            <begin position="157"/>
            <end position="189"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="1">
        <location>
            <begin position="198"/>
            <end position="230"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029493.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="264" mass="31009" checksum="4FC37E224945ECA7" modified="2019-05-08" version="1">MATEIGSPPRFFHMPRFQHQAPRQLFYKRPDFAQQQAMQQLTFDGKRMRKAVNRKTIDYNPSVIKYLENRVWQRDQRDMRAIQPDAGYYNDLVPPIGMLNNPMNAVTTKFVRTSTNKVKCPVFVVRWTPEGRRLVTGASSGEFTLWNGLTFNFETILQAHDSPVRAMTWSHNDMWMLTADHGGYVKYWQSNMNNVKMFQAHKEAIREASFSPTDNKFATCSDDGTVRIWDFLRCHEERILRAWLLYFSGISLELELSFLFVCWL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INF7</accession>
    <name>A0A452INF7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Ras-GAP domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000033632.1">
        <property type="protein sequence ID" value="ENSGAGP00000029618.1"/>
        <property type="gene ID" value="ENSGAGG00000019812.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005938">
        <property type="term" value="C:cell cortex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903479">
        <property type="term" value="P:mitotic actomyosin contractile ring assembly actin filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05131">
        <property type="entry name" value="RasGAP_IQGAP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00201">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.506.10:FF:000004">
        <property type="entry name" value="IQ motif containing GTPase activating protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.506.10">
        <property type="entry name" value="GTPase Activation - p120gap, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000048">
        <property type="entry name" value="IQ_motif_EF-hand-BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000593">
        <property type="entry name" value="RasGAP_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023152">
        <property type="entry name" value="RasGAP_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001936">
        <property type="entry name" value="RasGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001202">
        <property type="entry name" value="WW_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149">
        <property type="entry name" value="RAS GTPASE-ACTIVATING PROTEIN WITH IQ MOTIF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149:SF12">
        <property type="entry name" value="RAS GTPASE-ACTIVATING-LIKE PROTEIN IQGAP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00612">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00616">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03836">
        <property type="entry name" value="RasGAP_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00015">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00323">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00456">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF143885">
        <property type="entry name" value="RGC domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00509">
        <property type="entry name" value="RAS_GTPASE_ACTIV_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50018">
        <property type="entry name" value="RAS_GTPASE_ACTIV_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01159">
        <property type="entry name" value="WW_DOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Ras-GAP" evidence="2">
        <location>
            <begin position="657"/>
            <end position="885"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="PROSITE" id="PS50018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029618.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1289" mass="147474" checksum="2511843BFF3B6B9B" modified="2019-05-08" version="1">LEGEKWDDNGDRRNKTEKEKDMAIPGLVKSSFRIRWSNALIIDLSDGKNPINFLYFPPELVAVDHINTAIRNCDANKTLVALMKPEAQLPVVHPFAAAVYQTELFNLQQQNAVRYLAHDELSIAVEMLSAVVLLNQALGSKDILAIKNHLRNPSVGFNNVEDENFQRYADTLLSIKSEASSQGQDYLSWNDVQNCIDMVNMQVQEENDRIVAVSYINEAIDQGNPGKTLETLLLPTVKLHDVNPTNARHYQDVLHDAKVQKCKESHDESAVLWLDEIQKGINDANSNIEKAANLALGTSMINNCLEKHDSQPVLDILQSPKFGLRVVPECAETYYKNLLEAKNLKTKEDSSEGPWLKLIMKNRYDYYYNVETGESTFVPPEGFIPKTSWLTSEEIQTIVGQVTADYNREQLWFANENLILQLQALARGFLVRKSYEERKGFLQKQEPSVIKIQVLLCNIKAQSLLLEHDLLFEVSCSMLSKVFSLWSFFQNEEIVKIQAFLRANKARDDYRTLIGAENPPLNVLRKFAYLLDQSDLDFQEELEVTRLREEVVTKIRSNQQLEKDLNLMDIKIGLLVKNRISLQDVVLHSKKLNKKSKNQVEEMVTGDKQGIKGLSKERRKQLEAYQHLFYLLQTNPMYLAKLIFQMPQNKSTKFMDTVIFTLYNYASNQREEYLLLKLFKTALEEEINSKVDQIQDIVTGNPTVIKMVVSFNRGARGQNSLRQLLAPVVKEIMDDKSLLINTSPVEVYKAWVNQLETQTGEASKLPYDVTTEQALEHPEVVNRLESSIQSLRTVTDKVLVSIFSSYGMRYIAKVLKNSLHEKFPDATEDELLKIVGNLLYYRYMNPAIVAPDGFDIIDMTAGGQIHSDQRRNLGCVAKVLQHAASNKLFEGESAHLSSMNTYLSQTYQKFRNFFQIACDVPEPEEKFNIDEYSDMVTLSKPVIYISIEEIINTHSLLLEHQDAIAPEQNDLLNELLKGLGAVPDVESFLGAVDPNDPNKENTLSQLAKTEISLSLTSKYELQEGEDQDLKGTKRLVVDVIRTQPGDTLLEILEIPASAQQESEHLKVVEKRAILDSKTPEKMKHSQSMLEDGQLPIEQKKRKIQRNLRTLEQAGLVSSANKYQEIINEIAKDIRNQRRYRHHRKAELVKLQQTLSALQSKTAFYEDQVNYYNTYIKTCLDNLTRNKKLKHTSLKYTAARLHEKGVILEIEDLQTNQFKNVMFDITPGEEMGDFEIKAKFLGVQMEKVELHFQDLLQMQYEGVAVMKMFDKAKVNVNLLIFLLNKKFYGK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452ING0</accession>
    <name>A0A452ING0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">SMB domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452ING0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033421.1">
        <property type="protein sequence ID" value="ENSGAGP00000029432.1"/>
        <property type="gene ID" value="ENSGAGG00000020186.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009986">
        <property type="term" value="C:cell surface"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004551">
        <property type="term" value="F:dinucleotide phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003676">
        <property type="term" value="F:nucleic acid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004528">
        <property type="term" value="F:phosphodiesterase I activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046034">
        <property type="term" value="P:ATP metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030505">
        <property type="term" value="P:inorganic diphosphate transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045599">
        <property type="term" value="P:negative regulation of fat cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009143">
        <property type="term" value="P:nucleoside triphosphate catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030500">
        <property type="term" value="P:regulation of bone mineralization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd16018">
        <property type="entry name" value="Enpp"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00091">
        <property type="entry name" value="NUC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.410.20:FF:000003">
        <property type="entry name" value="Ectonucleotide pyrophosphatase/phosphodiesterase family member 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.410.20:FF:000001">
        <property type="entry name" value="Ectonucleotide pyrophosphatase/phosphodiesterase family member 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.720.10:FF:000145">
        <property type="entry name" value="Uncharacterized protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.410.20">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.720.10">
        <property type="entry name" value="Alkaline Phosphatase, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.570.10">
        <property type="entry name" value="Extracellular Endonuclease, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017850">
        <property type="entry name" value="Alkaline_phosphatase_core_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044929">
        <property type="entry name" value="DNA/RNA_non-sp_Endonuclease_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001604">
        <property type="entry name" value="Endo_G_ENPP1-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020821">
        <property type="entry name" value="ENPP1-3/EXOG-like_nuc-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044925">
        <property type="entry name" value="His-Me_finger_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002591">
        <property type="entry name" value="Phosphodiest/P_Trfase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036024">
        <property type="entry name" value="Somatomedin_B-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001212">
        <property type="entry name" value="Somatomedin_B_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10151">
        <property type="entry name" value="ECTONUCLEOTIDE PYROPHOSPHATASE/PHOSPHODIESTERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10151:SF77">
        <property type="entry name" value="ECTONUCLEOTIDE PYROPHOSPHATASE_PHOSPHODIESTERASE FAMILY MEMBER 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01223">
        <property type="entry name" value="Endonuclease_NS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01663">
        <property type="entry name" value="Phosphodiest"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01033">
        <property type="entry name" value="Somatomedin_B"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00892">
        <property type="entry name" value="Endonuclease_NS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00477">
        <property type="entry name" value="NUC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00201">
        <property type="entry name" value="SO"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53649">
        <property type="entry name" value="Alkaline phosphatase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54060">
        <property type="entry name" value="His-Me finger endonucleases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90188">
        <property type="entry name" value="Somatomedin B domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00524">
        <property type="entry name" value="SMB_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50958">
        <property type="entry name" value="SMB_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="4" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <feature type="domain" description="SMB" evidence="7">
        <location>
            <begin position="7"/>
            <end position="47"/>
        </location>
    </feature>
    <feature type="domain" description="SMB" evidence="7">
        <location>
            <begin position="50"/>
            <end position="94"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50958"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029432.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="803" mass="92332" checksum="B0146527EA294AE7" modified="2019-05-08" version="1">MTLSNISVKTCKGRCFERTFGSCRCDRDCVELGNCCLDFQETCIQPGKTHIWTCNKFRCGEKRWPENRCSCSDDCVENKDCCVNYYIVCQGAKSWVEEKCEDINEPQCPPGFAKPPVLLFSLDGFRAEYLHTWSGLLPVISKLQKCGTYTPNMRPVYPTKTFPNHYTIVTGLYPESHGIIDNKIYDPHRNVSFTLKNAEKFNPQWYQGQPIWLTAMHQGLKAGTFFWPGSDVRINGTFPDLYRKYNGYVKHRVCVCVCVCACVRAPHFYTLYLEEPDSSGHNFGPVSSGVILALQRVDNIVGMLMDGLKQMNLHKCLNIILLSDHGMETASCSKTAYLSTYMENVQDITVIPGPAARLRPQNVPDEYFSYPDQHFKAFMKELLPKRFHYVNNDRIERVHFYLDPQWQLARKPSEIKYCNGGFHGSDNRFMSMQAIFIGFGPGFKFKTQVDPFENIEIYNLICDLLGITPAYNNGTHGSLNHLLKNPVYIPRHPEEESHPSVYPLMKTTSSLDIGCSCDLYIELVINSVSALVKKIEKYSLPYGRPRVLQKRSTYYLLYHHQYVSGYSQDLNMTLWSAYTVTKYDNRTASVENLSNCLHVDIRIPSSSSKTCSFYKNHHQLSYGFLFPPNTKKSSYEVLLNSNIVPMYHEFKVLWNYFYDVLLPKYTLARNGVNVVSGPVFDYDSNGLYDTPEKVKRLTHDPEVLVPTHYFIVLTSCKNISQTPLQCEGALDALSFIVPHRPDNSESCARHKRDSLWVEERMRFHTARVRDVELLTGLSFYQDRKQPVSDILQLKTYLPTFPME</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452ING3</accession>
    <name>A0A452ING3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">RING-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
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            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
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            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
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        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
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</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452ING7</accession>
    <name>A0A452ING7_9SAUR</name>
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        <recommendedName>
            <fullName evidence="2">Transforming growth factor-beta-induced protein ig-h3</fullName>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
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        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
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    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
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                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
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    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="11">Plays a role in cell adhesion. May play a role in cell-collagen interactions.</text>
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        <text evidence="12">Binds to type I, II, and IV collagens.</text>
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        <subcellularLocation>
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            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
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    </dbReference>
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        <property type="entry name" value="TRANSFORMING GROWTH FACTOR-BETA-INDUCED PROTEIN IG-H3"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Fasciclin"/>
        <property type="match status" value="4"/>
    </dbReference>
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        <property type="entry name" value="FAS1"/>
        <property type="match status" value="4"/>
    </dbReference>
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    </dbReference>
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        <property type="match status" value="1"/>
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    <keyword evidence="10" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="3" id="KW-0301">Gamma-carboxyglutamic acid</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="7 13" id="KW-0732">Signal</keyword>
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        <location>
            <begin position="1"/>
            <end position="21"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019527157" description="Transforming growth factor-beta-induced protein ig-h3" evidence="13">
        <location>
            <begin position="22"/>
            <end position="667"/>
        </location>
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        <location>
            <begin position="41"/>
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        </location>
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        </location>
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        </location>
    </feature>
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            <begin position="497"/>
            <end position="627"/>
        </location>
    </feature>
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    </evidence>
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    </evidence>
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        <source>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00045945"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00047016"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50213"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="PROSITE" id="PS51041"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029503.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="667" mass="72808" checksum="D4149F550D74E5FD" modified="2019-05-08" version="1">MERGVLLLALAWALGLTLAGAARSPYQQVLHHSRLRGRQHGPNVCAVQKLLGTNRKYYTNCKQWYQRKVCGKSTVISYECCPGYEKVPGEKGCPAALPLSNIYDTLGIVGASTTQLYSLSSNLKPDIEAPGSFTIFAPNNEAWAALPAEMLDSLVSNVNIELLNALRYHMVNRRVLTDDLKHGTSLPSMYQNIDLQIHHYPNGIVTVNCARLLKADHHATNGVVHLIDKVIPTTTNTIQQIVETDENLETLRAAVAASGLNTLLESDGQFTLLAPTNEAFEKIPPETLNRILGDPEALKDLLNHHILKSAMCAEAIIAGLSMETLEGTMLEVGCTGDELTLNGRPIIAEKDILATNGVVHFVNELLIPDSAKTLFELGKESDVSTSIDLFRQADLSSHLTGNERVTLLAPINTVFKDGTPPMNQNTKSLLLDHILEEQLSSKYLYHGQMLKTLGGKELRVFVYRNNLCIENVCIAAHDRRGRFGTLFTVDKMLTPPVGTLMDVLKADQRFSTLVAAIQSAGLMETLNRPGAFTVFAPTNEAFRAMPQGELNKLMGNAKELATILKYHIGDEILVSGAVGALVRLKSLQGDKIEVSSKNNVVNVNKEPVAETDIMATNGVIYAVNAVLQPPAFRPQERGDEPADPALEIFKQASALSKVSQRSARLGK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452ING8</accession>
    <name>A0A452ING8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">glutamate carboxypeptidase II</fullName>
            <ecNumber evidence="18">3.4.17.21</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>Release of an unsubstituted, C-terminal glutamyl residue, typically from Ac-Asp-Glu or folylpoly-gamma-glutamates.</text>
            <dbReference type="EC" id="3.4.17.21"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Membrane</location>
            <topology evidence="2">Single-pass type II membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the peptidase M28 family. M28B subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.4.17.21" evidence="18"/>
    <dbReference type="AlphaFoldDB" id="A0A452ING8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033432.1">
        <property type="protein sequence ID" value="ENSGAGP00000029442.1"/>
        <property type="gene ID" value="ENSGAGG00000020963.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016805">
        <property type="term" value="F:dipeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004181">
        <property type="term" value="F:metallocarboxypeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd08022">
        <property type="entry name" value="M28_PSMA_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.930.40:FF:000001">
        <property type="entry name" value="N-acetylated-alpha-linked acidic dipeptidase 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.630.10:FF:000009">
        <property type="entry name" value="N-acetylated-alpha-linked acidic dipeptidase 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.930.40">
        <property type="entry name" value="Transferrin receptor-like, dimerisation domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.630.10">
        <property type="entry name" value="Zn peptidases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046450">
        <property type="entry name" value="PA_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007484">
        <property type="entry name" value="Peptidase_M28"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039373">
        <property type="entry name" value="Peptidase_M28B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007365">
        <property type="entry name" value="TFR-like_dimer_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036757">
        <property type="entry name" value="TFR-like_dimer_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10404:SF36">
        <property type="entry name" value="GLUTAMATE CARBOXYPEPTIDASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10404">
        <property type="entry name" value="N-ACETYLATED-ALPHA-LINKED ACIDIC DIPEPTIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04389">
        <property type="entry name" value="Peptidase_M28"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04253">
        <property type="entry name" value="TFR_dimer"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52025">
        <property type="entry name" value="PA domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47672">
        <property type="entry name" value="Transferrin receptor-like dimerisation domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53187">
        <property type="entry name" value="Zn-dependent exopeptidases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0121">Carboxypeptidase</keyword>
    <keyword evidence="12" id="KW-0224">Dipeptidase</keyword>
    <keyword evidence="15" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="14 19" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13" id="KW-0482">Metalloprotease</keyword>
    <keyword evidence="16" id="KW-0511">Multifunctional enzyme</keyword>
    <keyword evidence="5" id="KW-0645">Protease</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0735">Signal-anchor</keyword>
    <keyword evidence="6 19" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 19" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="9" id="KW-0862">Zinc</keyword>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="21"/>
            <end position="42"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase M28" evidence="21">
        <location>
            <begin position="297"/>
            <end position="499"/>
        </location>
    </feature>
    <feature type="domain" description="Transferrin receptor-like dimerisation" evidence="20">
        <location>
            <begin position="567"/>
            <end position="687"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004606"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005634"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022645"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022997"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023049"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023268"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00052003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00066561"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="Pfam" id="PF04253"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="Pfam" id="PF04389"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029442.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="690" mass="78278" checksum="B63D3687109CDEC9" modified="2019-05-08" version="1">MWTALRSGAEQSSPWGSRGRVCALSLAVAAALLLLGFLIGWFTKPTDEVTATSFHEKVKKAFMAEMKAENIKQFLYNFTQRPHLAGTAENLRLAQQIQAQWKEFGLDSVELAHYDVLLSYPNETNPNYISIIDEHGTEIFNTSLFEPPPPGYETVTDVVPPYSAFSAPGVPEVKSNRLLKSRKLKVSCSQLGLIFRGSWKIRPHKLYSPGNFQLMRFCRTAAGHLEHGVWEHSWIMTSSKINTSLIPPEYTYRYEEARAVGLPKLPVHPIGYNDAEKLLRKVKMHIHSSNKVTRIYNVIGTIRGATEPDRYVILGGHRDSWVFGGIDPQSGAAVVHEIVRSYGKLKKEGWRPRRTIIFASWDAEEFGLLGSTEWAEENTKLLQARGVAYINADSSVEGNYTLRIDCTPLMYSLVYSLTREIPSPDEGFKGKSLYESWHENNPSTEYKDVPRINKLGSGNDFEVFFQRLGIASGRARYSKNWRVEKYSSYPVYHSVYETYEIVERFYDPTFKNHLTVAQVRGGLVFELADSVVLPFDCRDYAEALRNYVQVIYNLSKNHQMGLTTYSVSFDSLFSAANNFTEAATDFHRRLQQVDINNPIAVRALNDQLMFLERAFIDPLGLPGRPFYRHIIFAPSSHNKYAGESFPGIYDAMFDIENKADQRKAWEEVKRQISIAAFTVQAAAGTLKDLA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452ING9</accession>
    <name>A0A452ING9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">non-specific serine/threonine protein kinase</fullName>
            <ecNumber evidence="3">2.7.11.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="12">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033396.1">
        <property type="protein sequence ID" value="ENSGAGP00000029410.1"/>
        <property type="gene ID" value="ENSGAGG00000021255.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.20.90:FF:000007">
        <property type="entry name" value="Serine/threonine-protein kinase WNK1 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000006">
        <property type="entry name" value="Serine/threonine-protein kinase WNK1 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.20.90:FF:000012">
        <property type="entry name" value="Serine/threonine-protein kinase WNK1 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000494">
        <property type="entry name" value="serine/threonine-protein kinase WNK2 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.20.90">
        <property type="entry name" value="Phosphatidylinositol 3-kinase Catalytic Subunit, Chain A, domain 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056865">
        <property type="entry name" value="CCTL2_WNK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024678">
        <property type="entry name" value="Kinase_OSR1/WNK_CCT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050588">
        <property type="entry name" value="WNK_Ser-Thr_kinase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13902">
        <property type="entry name" value="SERINE/THREONINE-PROTEIN KINASE WNK WITH NO LYSINE -RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24889">
        <property type="entry name" value="CCTL2_WNK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12202">
        <property type="entry name" value="OSR1_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="10" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="13" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9" id="KW-0418">Kinase</keyword>
    <keyword evidence="8" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="7" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Protein kinase" evidence="15">
        <location>
            <begin position="192"/>
            <end position="450"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1"/>
            <end position="184"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="606"/>
            <end position="625"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1094"/>
            <end position="1135"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1209"/>
            <end position="1248"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1273"/>
            <end position="1302"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1384"/>
            <end position="1404"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1862"/>
            <end position="1926"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="2204"/>
            <end position="2233"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="2245"/>
            <end position="2277"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="13">
        <location>
            <begin position="1953"/>
            <end position="1980"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="1"/>
            <end position="12"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="110"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="125"/>
            <end position="144"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="147"/>
            <end position="173"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="606"/>
            <end position="615"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="616"/>
            <end position="625"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="14">
        <location>
            <begin position="1116"/>
            <end position="1126"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="1220"/>
            <end position="1229"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="1230"/>
            <end position="1239"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="1882"/>
            <end position="1891"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="1903"/>
            <end position="1926"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="2209"/>
            <end position="2218"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012513"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00047899"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00048679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029410.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2277" mass="246802" checksum="40AEA1AD1D929E29" modified="2019-05-08" version="1">METGCAASSAPPRGGGGASPAAARPPPADGSAAVGGLGRRSGAGMETAAVPERGPAPGLPPRGPGAREPRVGHLRFLRRSVVDSDQEEPPLELSEAEPRRIVLLSKTRRIIAERAKERHGEADLEPPAVEPAAEPKGAVPGVPATAEEQKELGRDAERKEPSDKDQSKSKKQEPEEEADMKAVSTSPDGRFLKFDIELGRGSFKTVYKGLDTETWVEVAWCELQDRKLTKVERQRFKEEAEMLKGLQHPNIVRFYDFWESCVKGKKCIVLVTELMTSGTLKTYLKRFKVMKPKVLRSWCRQILKGLLFLHTRTPPIIHRDLKCDNIFITGPTGSVKIGDLGLATLKRASFAKSVIGTPEFMAPEMYEEHYDESVDVYAFGMCMLEMATSEYPYSECQNAAQIYRKVTCGVKPASFEKVTDPEIKEIIGECICKNKEERYEIKDLLSHAFFAEDTGVRVELAEEDHGRKSSIALRLWVEDPKKLKGKPKDNGAIEFTFDLEKETPDDVAQEMVDSGFFHESDVKIVAKSIRDRVALIQWRRERIWPMIQSEDHQVSECLEKLKVPQAQQVQVTSLSHAGQQTLSALEEPEADQHLFQHNLPTSVTSVASDSTFDSGQGSTVYSDSQSSQQSVIYSSLPDSVPPAIQRVYSPPLSESQALPQSLQQLGHYQQPSVPGLTVAQAPPAVLSQRPQHYQEPAMTFPVIQPVTVASSQMGQSQPVLASQQQPIPQQAPLQQVLASQPVCPLQPAAHLLPQYQTQTSQVAATSTQLTPLQISTVRQLPHVPPSQIPQLPVIPAVTPLAGLDNLPPSLSDLPAANVPSVPAPSQYFTPTVIIPSLPMNPALPLASNPPALSMQAVNLPHTTVASLVLPCQTIVPNMSAATIPLLAVAPPGVSALPTHHAISHLSQQQMYQPAFQQIIQSDAPSPHHTQNMQAVSQQQLPPLPQSLQPSIIHPSEQTMSTSGVGNQIIGHPQYGLEIGAQMPVIPMQPSIVMPPPLQLQPEILPPCLGPESLSQLHGNLAAASATVPIDHCLPLQPTGLLQLQPDLSQPLGQQAPCSAIQSVTESTQEEHAIQEKQQTLSLCCESYMSPDVASGKEMSDSFEGVVGSGKQEGKPSKKHKKSTRVRSRQEKSSRPKLTILNVCNTGDKMVECQLETHNHKMVTFKFDLDGDAPEEIATYMVEKEFILQTEKETFIEQMKDIIDKAEDMLSEDTEGERSSDQGTSPQQDASRLEMNEENRQSQVKTPMYQQNVLHTGKRWFIICPVVENPTEEVPEFSPAVPQSKQAEGPDPEQSDDQQMGSAVTDTPSVFFCHQLPSQTSVFDSPCGTPAPLAQVPAAASSAVLQTKAEILAATLPGTAPHVPEQFVTNVGQPEYPPLHLVMPSQPDASAHNAASLEGPSDAQLAPKSSLQTLQEMDEDSCIPDVEISCCTVRPPSSILTVPAQGAEFCPVPALSDVVVLELQPAPQIPQVSEASQPGVMRHSFVNQLLPGGVVSDPLSLTGSQLQSPPQLPVAVSQQHTMLTQSQPVTCTGLGVGPLQPPCAVESDGEGPPRVDFADDTIKSLDEKLRNLLYQECVPTSAASAGTPDSFVPLEQGESELASLPPLNEELIPTLVLDSREPGLNVPDTFQEANAAQDQCVSLVPSAMPPYPVLASQALTVPFIPINPAAMETSAVLNNATATSHSRTEFRIQEPMLEKFETTSIDVSSSKVRVYETGSAANISSTSAVTTQGLQVAPASSTIPEQMETSRRNKEAKIKISSALTENLIQTSTTDREINNLQRNDFVDPGSCGKQSEVHGNSTPAKTIGRFSVISTHDELTLTSPHCLRYSAPPDVYLDELPSSPDLKTSVRRVQTASFLDVLSDHGSSDSGDESLRRKSVAAQLSPQSSIAANDITKKPASLLQRSGKTSSPGPDSPNGQGTKIPTINITSFHSQSSYMSSDNDSEFEDADMKKELQNLREKHMKEIAELQTQQKNEIEALFVRLGKPLPPNVGFLHSVPPTGRRRRTSKNKLKAGKLLNPLVQQLKTAPSNTSNLSVCKESLPKETTKTHAGSTEATAVTSSTSATFGNSVQTQQPCSVKASLSSDICSGLATEGGNAHGSSGQGWTVFHQTSERVTYKSSSKPRTRFLSGPMSLSIWSTLKRLCLGKDHSSRSSTNSLVTCPEPLPPPTLQVQVQANNSNNKKGTFTDDLHKLVDQWTSKTVGAAQSKPSLNQLKQNQKRQDMEPKANPMQAQNETCGILKMRMPLPPIDAGPAQLQYQAQGPMSAKEDEEEDVK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INH0</accession>
    <name>A0A452INH0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Eyes absent homolog</fullName>
            <ecNumber evidence="15">3.1.3.48</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="12">
            <text>O-phospho-L-tyrosyl-[protein] + H2O = L-tyrosyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:10684"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="EC" id="3.1.3.48"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="14 15">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
        <text evidence="14 15">Binds 1 Mg(2+) ion per subunit.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 15">Belongs to the HAD-like hydrolase superfamily. EYA family.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.48" evidence="15"/>
    <dbReference type="AlphaFoldDB" id="A0A452INH0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033395.1">
        <property type="protein sequence ID" value="ENSGAGP00000029409.1"/>
        <property type="gene ID" value="ENSGAGG00000021258.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:2001240">
        <property type="term" value="P:negative regulation of extrinsic apoptotic signaling pathway in absence of ligand"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045739">
        <property type="term" value="P:positive regulation of DNA repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd02601">
        <property type="entry name" value="HAD_Eya"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.12350:FF:000001">
        <property type="entry name" value="Eyes absent homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.12350">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028472">
        <property type="entry name" value="EYA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006545">
        <property type="entry name" value="EYA_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042577">
        <property type="entry name" value="EYA_dom_metazoan"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038102">
        <property type="entry name" value="EYA_dom_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01658">
        <property type="entry name" value="EYA-cons_domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10190">
        <property type="entry name" value="EYES ABSENT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10190:SF17">
        <property type="entry name" value="EYES ABSENT HOMOLOG 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00702">
        <property type="entry name" value="Hydrolase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDG01129">
        <property type="entry name" value="C1.5:_HAD__Beta-PGM__Phosphata"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDS00003">
        <property type="entry name" value="Haloacid_Dehalogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0010">Activator</keyword>
    <keyword evidence="3" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="5 15" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="6 14 15" id="KW-0460">Magnesium</keyword>
    <keyword evidence="4 14 15" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="7 15" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0804">Transcription</keyword>
    <keyword evidence="8 15" id="KW-0805">Transcription regulation</keyword>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1"/>
            <end position="42"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="69"/>
            <end position="93"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="180"/>
            <end position="207"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="272"/>
            <end position="347"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1"/>
            <end position="14"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="16">
        <location>
            <begin position="15"/>
            <end position="37"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="16">
        <location>
            <begin position="76"/>
            <end position="93"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="16">
        <location>
            <begin position="276"/>
            <end position="290"/>
        </location>
    </feature>
    <feature type="active site" description="Nucleophile" evidence="13">
        <location>
            <position position="354"/>
        </location>
    </feature>
    <feature type="active site" description="Proton donor" evidence="13">
        <location>
            <position position="356"/>
        </location>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="354"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="356"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="582"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010501"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00051722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PIRSR" id="PIRSR628472-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PIRSR" id="PIRSR628472-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="RuleBase" id="RU362036"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029409.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="618" mass="67235" checksum="16E6666A06A18DEB" modified="2019-05-08" version="1">MEMQDLTSPHNLVGSSDNPSSSKLDKSNLSSTSITTNGTGGDNMTVLNTADWLLSCSTPSSATMSLLAVKTEPMNSSETTTTTTTGDGSLDTFTGSVITSSGYSPRSAHQYSPQIYPSKPYPHILSTPAAQSMSAYSGQTQYSGMQQPAVYTAYSQTGQPYSLPTYDLGVMLPGIKTESGLSQTQSPLQSGCLSYSPGFSTPQPGQTPYSYQMPGSSFTPSSTIYANNSVSNSTNFSSSQQDYPSYTAFGQNQYAQYYSASTYGAYMTSNNTADGTSSSSSTYQLQESLSGLNSQPGTDLHPGEFDTVQSPSTPIKDLDERTCRSSGSKSRGRGRKNNPSPPPDSDLERVFVWDLDETIIVFHSLLTGSYAQKYGKDPPMAVTLGLRMEEMIFNLADTHLFFNDLEECDQVHIDDVSSDDNGQDLSTYSFATDGFHAAASSANLCLPTGVRGGVDWMRKLAFRYRRVKELYNTYKNNIGGLLGPAKRDAWLQLRAEIEALTDSWLTNALKSLSIISTRSNCVNVLVTTTQLIPALAKVLLYSLGGAFPIENIYSATKIGKESCFERIMQRFGRKVVYVVIGDGVEEEQAAKKHNMPFWRISSHSDLLALHQALELEYL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INH1</accession>
    <name>A0A452INH1_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="15">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the peptidase S1C family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452INH1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033647.1">
        <property type="protein sequence ID" value="ENSGAGP00000029633.1"/>
        <property type="gene ID" value="ENSGAGG00000021395.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019838">
        <property type="term" value="F:growth factor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043065">
        <property type="term" value="P:positive regulation of apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012501">
        <property type="term" value="P:programmed cell death"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00104">
        <property type="entry name" value="KAZAL_FS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.120:FF:000002">
        <property type="entry name" value="HtrA serine peptidase 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.60.30:FF:000026">
        <property type="entry name" value="Insulin-like growth factor-binding protein 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.42.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.10.120">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.60.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.40.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009030">
        <property type="entry name" value="Growth_fac_rcpt_cys_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000867">
        <property type="entry name" value="IGFBP-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002350">
        <property type="entry name" value="Kazal_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036058">
        <property type="entry name" value="Kazal_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036034">
        <property type="entry name" value="PDZ_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009003">
        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001940">
        <property type="entry name" value="Peptidase_S1C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22939">
        <property type="entry name" value="SERINE PROTEASE FAMILY S1C HTRA-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22939:SF13">
        <property type="entry name" value="SERINE PROTEASE HTRA1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00219">
        <property type="entry name" value="IGFBP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07648">
        <property type="entry name" value="Kazal_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13365">
        <property type="entry name" value="Trypsin_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00834">
        <property type="entry name" value="PROTEASES2C"/>
    </dbReference>
    <dbReference type="SMART" id="SM00121">
        <property type="entry name" value="IB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00280">
        <property type="entry name" value="KAZAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57184">
        <property type="entry name" value="Growth factor receptor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF100895">
        <property type="entry name" value="Kazal-type serine protease inhibitors"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50494">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51323">
        <property type="entry name" value="IGFBP_N_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51465">
        <property type="entry name" value="KAZAL_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="10" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="11" id="KW-0340">Growth factor binding</keyword>
    <keyword evidence="8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="6" id="KW-0645">Protease</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0964">Secreted</keyword>
    <keyword evidence="9" id="KW-0720">Serine protease</keyword>
    <keyword evidence="7 12" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="12">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019279250" evidence="12">
        <location>
            <begin position="23"/>
            <end position="421"/>
        </location>
    </feature>
    <feature type="domain" description="IGFBP N-terminal" evidence="13">
        <location>
            <begin position="31"/>
            <end position="101"/>
        </location>
    </feature>
    <feature type="domain" description="Kazal-like" evidence="14">
        <location>
            <begin position="84"/>
            <end position="145"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00010541"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023183"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS51323"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS51465"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029633.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="421" mass="44928" checksum="51F4729EE8A5778C" modified="2019-05-08" version="1">MLLWLSWAGGCCLLLLLPGSAAEPARRYDERAAGCSERCDRARCPAVPADCAGGEALDACGCCPVCAAQEGEPCGGPAGEPPCGEGLQCRKRAKAGLCVCTSSEPVCGSDARTYGNLCQLRAASRRSERLQQPPVIAIQRGACGQGQEDPNSLRHKYNFIADVVEKIAPAVVHIELFRKLPFTKREVPVASGSGFIVSEDGLIVTNAHVVTNKHRVKVELKNGATYEAKIKDVDEKADIALIKIDTQGKLPVLLLGRSADLRPGEFVVAIGSPFSLQNTVTTGIVSTTQRGGKELGLRNSDMDYIQTDAIINYGNSGGPLVNLDGEVIGINTLKVTAGISFAIPSDKIKKFLTESHDRQAKGKAITKKKYIGIRMMSLSPSKAKELKEHHKDFPDVVSGAYIIEVIPDTPAEASAVISETF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INH3</accession>
    <name>A0A452INH3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">UDENN FNIP1/2-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Lysosome membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the FNIP family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452INH3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033437.1">
        <property type="protein sequence ID" value="ENSGAGP00000029447.1"/>
        <property type="gene ID" value="ENSGAGG00000021275.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005765">
        <property type="term" value="C:lysosomal membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042030">
        <property type="term" value="F:ATPase inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051087">
        <property type="term" value="F:protein-folding chaperone binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037545">
        <property type="entry name" value="DENN_FNIP1/2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028086">
        <property type="entry name" value="FNIP_C_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026156">
        <property type="entry name" value="FNIP_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028085">
        <property type="entry name" value="FNIP_mid_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028084">
        <property type="entry name" value="FNIP_N_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21634:SF12">
        <property type="entry name" value="FOLLICULIN-INTERACTING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21634">
        <property type="entry name" value="RE13835P"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14638">
        <property type="entry name" value="FNIP_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14637">
        <property type="entry name" value="FNIP_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14636">
        <property type="entry name" value="FNIP_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR02073">
        <property type="entry name" value="FOLLICULNIP1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51836">
        <property type="entry name" value="DENN_FNIP12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0458">Lysosome</keyword>
    <keyword evidence="5" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="UDENN FNIP1/2-type" evidence="8">
        <location>
            <begin position="10"/>
            <end position="1139"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="65"/>
            <end position="97"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="602"/>
            <end position="626"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="894"/>
            <end position="940"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
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            <begin position="68"/>
            <end position="84"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
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            <begin position="894"/>
            <end position="907"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004656"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007541"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
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    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023228"/>
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    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51836"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029447.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1157" mass="128908" checksum="92819CCC3D0B2760" modified="2019-05-08" version="1">MLPSWPLPEFDPSQIRLIVYQDCERRGRNVLFDSSSKRKTEDVSVSKLCSDAQVRVFGKCCQLKPGGDSSSSLDSSINSSSLSSDTKEQCPKYQGSRCSSDANMLGEMMFGSVAMSYKGSTLKIHQIRSPPQLMLSKVFTARTGSSIYGSLNTLQDSLEFINQDSNTLKPDHSTIMNGLLGNIGLSQLCSPRRAFSEQGPLRLIRSASFFAVHSNPMDMPGREQNEDRDSGIARSASLSSLLITPFPSPGSSLTRSCASSYQRRWRRSQTTSLENGVFPRWSVEESFNLSDDSSGLSPGIVRKKKIAIGVIFSLSKDEDENNKFHEFFFSHFPLFESHMNKLKSAIEQAMKMSRRSADASQRSLAYNRIVDALNEFRTTICNLYTMPRIGEPVWLTMMSGIPEKNQLCHRFMKEFTFLMENASKNQFLPALLTAILTNHLAWVPTVMPNGQPPIKIFLEKHSSQSVDMLAKTHPYNPLWAQLGDLYGAIGSPVRLAKTVVVGKRQDLVQRLLYFLTYFIRCSELQETHLLENGEDEAIVMPGTVITTTLEKGEVEESEYVLITIHRNRSNLLSSESEEMRTPNCSCKYCKCPLLLGQNTEGVSHQEREYAPNASKVEQETSSDESRTTCAADCQENAADIKQCKDKLISCLDTKLETVICTGSASVDKCAMADLGLEPTAGTWKTEESLESGNQASRVTRSTGIAVEKKPPDKLLSDAFPCSTAEAQTKVTFLIGDSMSPDSDIDLRSQAVVEQIARHHNQAAIEEEASAAQNCEAKQTVEDQNRDCGTAEPFPRVASEHQNWNPNPYNAESMSLFDEYFTDDGSIETRTIDDIPVQAAVEVLDHGSSLELSKKLGTMNSKLPSEFCKFMESVRQETCKNCFAEQDQRDKISICVPHGDKESSEKKVAPGSDWDIPRNESSDSALGDSESEDTGHDLTRQNSNYYVGEHEDWTEEDEIPFPGSKLVEVNSIQPSIANFGRSLLGGYCSSYVPDFVLQGVGNDEKLRHCLVSDLSHAVQHPVLDEPIAEAVCIIADTDKWTVQVASSQRRMIDNKLGKEVLVSSLVSNLLHSTLQLYKHNLSPNFCVMHLEDRLQELYFKSKMLSEYLKGQMRVHVKELGVVLGYDHNIVQYNFHLCFSHHKSSSHLLVGLSLPQGLH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INH5</accession>
    <name>A0A452INH5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">non-specific serine/threonine protein kinase</fullName>
            <ecNumber evidence="3">2.7.11.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="12">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033403.1">
        <property type="protein sequence ID" value="ENSGAGP00000029415.1"/>
        <property type="gene ID" value="ENSGAGG00000021255.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.20.90:FF:000007">
        <property type="entry name" value="Serine/threonine-protein kinase WNK1 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000006">
        <property type="entry name" value="Serine/threonine-protein kinase WNK1 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.20.90:FF:000012">
        <property type="entry name" value="Serine/threonine-protein kinase WNK1 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000494">
        <property type="entry name" value="serine/threonine-protein kinase WNK2 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.20.90">
        <property type="entry name" value="Phosphatidylinositol 3-kinase Catalytic Subunit, Chain A, domain 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056865">
        <property type="entry name" value="CCTL2_WNK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024678">
        <property type="entry name" value="Kinase_OSR1/WNK_CCT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050588">
        <property type="entry name" value="WNK_Ser-Thr_kinase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13902">
        <property type="entry name" value="SERINE/THREONINE-PROTEIN KINASE WNK WITH NO LYSINE -RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24889">
        <property type="entry name" value="CCTL2_WNK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12202">
        <property type="entry name" value="OSR1_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="10" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="13" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9" id="KW-0418">Kinase</keyword>
    <keyword evidence="8" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="7" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Protein kinase" evidence="15">
        <location>
            <begin position="192"/>
            <end position="450"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1"/>
            <end position="184"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="606"/>
            <end position="625"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="923"/>
            <end position="950"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="992"/>
            <end position="1032"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1107"/>
            <end position="1146"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1171"/>
            <end position="1200"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1282"/>
            <end position="1302"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1760"/>
            <end position="1824"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="2101"/>
            <end position="2131"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="2143"/>
            <end position="2175"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="13">
        <location>
            <begin position="1851"/>
            <end position="1878"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="1"/>
            <end position="12"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="110"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="125"/>
            <end position="144"/>
        </location>
    </feature>
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        <location>
            <begin position="147"/>
            <end position="173"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="606"/>
            <end position="615"/>
        </location>
    </feature>
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        <location>
            <begin position="616"/>
            <end position="625"/>
        </location>
    </feature>
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        <location>
            <begin position="923"/>
            <end position="935"/>
        </location>
    </feature>
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        <location>
            <begin position="936"/>
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        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="14">
        <location>
            <begin position="1014"/>
            <end position="1024"/>
        </location>
    </feature>
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        <location>
            <begin position="1118"/>
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        </location>
    </feature>
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        <location>
            <begin position="1128"/>
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    </feature>
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        <location>
            <begin position="1780"/>
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        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="1801"/>
            <end position="1824"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="2107"/>
            <end position="2116"/>
        </location>
    </feature>
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        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012513"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
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    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00047899"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00048679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029415.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2175" mass="236092" checksum="91873E9EB6FDA0DA" modified="2019-05-08" version="1">METGCAASSAPPRGGGGASPAAARPPPADGSAAVGGLGRRSGAGMETAAVPERGPAPGLPPRGPGAREPRVGHLRFLRRSVVDSDQEEPPLELSEAEPRRIVLLSKTRRIIAERAKERHGEADLEPPAVEPAAEPKGAVPGVPATAEEQKELGRDAERKEPSDKDQSKSKKQEPEEEADMKAVSTSPDGRFLKFDIELGRGSFKTVYKGLDTETWVEVAWCELQDRKLTKVERQRFKEEAEMLKGLQHPNIVRFYDFWESCVKGKKCIVLVTELMTSGTLKTYLKRFKVMKPKVLRSWCRQILKGLLFLHTRTPPIIHRDLKCDNIFITGPTGSVKIGDLGLATLKRASFAKSVIGTPEFMAPEMYEEHYDESVDVYAFGMCMLEMATSEYPYSECQNAAQIYRKVTCGVKPASFEKVTDPEIKEIIGECICKNKEERYEIKDLLSHAFFAEDTGVRVELAEEDHGRKSSIALRLWVEDPKKLKGKPKDNGAIEFTFDLEKETPDDVAQEMVDSGFFHESDVKIVAKSIRDRVALIQWRRERIWPMIQSEDHQVSECLEKLKVPQAQQVQVTSLSHAGQQTLSALEEPEADQHLFQHNLPTSVTSVASDSTFDSGQGSTVYSDSQSSQQSVIYSSLPDSVPPAIQRVYSPPLSESQALPQSLQQLGHYQQPSVPGLTVAQAPPAVLSQRPQHYQEPAMTFPVIQPVTVASSQMGQSQPVLASQQQPIPQQAPLQQVLASQPVCPLQPAAHLLPQYQTQTSQVAATSTQLTPLQISTVRQLPHVPPSQIPQLPVIPAVTPLAGLDNLPPSLSDLPAANVPSVPAPSQYFTPTVIIPSLPMNPALPLASNPPALSMQAVNLPHTTVASLVLPCQTIVPNMSAATIPLLAVAPPGVSALPTHHAISHLSQQQMYQPAFQQIIQSDAPSPHHTQNMQAVSQQQLPPLPQSLQPSIIHPSEQTMSTSGVGNQEHAIQEKQQTLSLCCESYMSPDVASGKEMSDSFEGVVGSGKQEGKPSKKHKKSTRVRSRQEKSSRPKLTILNVCNTGDKMVECQLETHNHKMVTFKFDLDGDAPEEIATYMVEKEFILQTEKETFIEQMKDIIDKAEDMLSEDTEGERSSDQGTSPQQDASRLEMNEENRQSQVKTPMYQQNVLHTGKRWFIICPVVENPTEEVPEFSPAVPQSKQAEGPDPEQSDDQQMGSAVTDTPSVFFCHQLPSQTSVFDSPCGTPAPLAQVPAAASSAVLQTKAEILAATLPGTAPHVPEQFVTNVGQPEYPPLHLVMPSQPDASAHNAASLEGPSDAQLAPKSSLQTLQEMDEDSCIPDVEISCCTVRPPSSILTVPAQGAEFCPVPALSDVVVLELQPAPQIPQVSEASQPGVMRHSFVNQLLPGGVVSDPLSLTGSQLQSPPQLPVAVSQQHTMLTQSQPVTCTGLGVGPLQPPCAVESDGEGPPRVDFADDTIKSLDEKLRNLLYQECVPTSAASAGTPDSFVPLEQGESELASLPPLNEELIPTLVLDSREPGLNVPDTFQEANAAQDQCVSLVPSAMPPYPVLASQALTVPFIPINPAAMETSAVLNNATATSHSRTEFRIQEPMLEKFETTSIDVSSSKVRVYETGSAANISSTSAVTTQGLQVAPASSTIPEQMETSRRNKEAKIKISSALTENLIQTSTTDREINNLQRNDFVDPGSCGKQSEVHGNSTPAKTIGRFSVISTHDELTLTSPHCLRYSAPPDVYLDELPSSPDLKTSVRRVQTASFLDVLSDHGSSDSGDESLRRKSVAAQLSPQSSIAANDITKKPASLLQRSGKTSSPGPDSPNGQGTKIPTINITSFHSQSSYMSSDNDSEFEDADMKKELQNLREKHMKEIAELQTQQKNEIEALFVRLGKPLPPNVGFLHSVPPTGRRRRTSKNKLKAGKLLNPLVQQLKTAPSNTSNLSVCKESLPKETTKTHAGSTEATAVTSSTSATFGNSVQTQQPCSVKASLSSDICSGLATEGGNAHGSSGQGWTVFHQTSERVTYKSSSKPRTRFLSGPMSLSIWSTLKRLCLGKDHSSRSSTNSLVTCPEPLPPPTLQVQVQANNSNNKKGTFTDDLHKLVDQWTSKTVGAAQSKPSLNQLKQNQKRQDMEPKANPMQAQNETCGILKMRMPLPPIDAGPAQLQYQAQGPMSAKEDEEEDVK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INH7</accession>
    <name>A0A452INH7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Tuberin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Catalytic component of the TSC-TBC complex, a multiprotein complex that acts as a negative regulator of the canonical mTORC1 complex, an evolutionarily conserved central nutrient sensor that stimulates anabolic reactions and macromolecule biosynthesis to promote cellular biomass generation and growth. Within the TSC-TBC complex, TSC2 acts as a GTPase-activating protein (GAP) for the small GTPase RHEB, a direct activator of the protein kinase activity of mTORC1. In absence of nutrients, the TSC-TBC complex inhibits mTORC1, thereby preventing phosphorylation of ribosomal protein S6 kinase (RPS6KB1 and RPS6KB2) and EIF4EBP1 (4E-BP1) by the mTORC1 signaling. The TSC-TBC complex is inactivated in response to nutrients, relieving inhibition of mTORC1. Involved in microtubule-mediated protein transport via its ability to regulate mTORC1 signaling. Also stimulates the intrinsic GTPase activity of the Ras-related proteins RAP1A and RAB5.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytosol</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="8">Lysosome membrane</location>
            <topology evidence="8">Peripheral membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000033653.1">
        <property type="protein sequence ID" value="ENSGAGP00000029638.1"/>
        <property type="gene ID" value="ENSGAGG00000021398.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005765">
        <property type="term" value="C:lysosomal membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045202">
        <property type="term" value="C:synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033596">
        <property type="term" value="C:TSC1-TSC2 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046627">
        <property type="term" value="P:negative regulation of insulin receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051898">
        <property type="term" value="P:negative regulation of phosphatidylinositol 3-kinase/protein kinase B signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032007">
        <property type="term" value="P:negative regulation of TOR signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030178">
        <property type="term" value="P:negative regulation of Wnt signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051726">
        <property type="term" value="P:regulation of cell cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016241">
        <property type="term" value="P:regulation of macroautophagy"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051056">
        <property type="term" value="P:regulation of small GTPase mediated signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.11210:FF:000004">
        <property type="entry name" value="Tuberin isoform X3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.11210">
        <property type="entry name" value="Rap/Ran-GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035974">
        <property type="entry name" value="Rap/Ran-GAP_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000331">
        <property type="entry name" value="Rap/Ran_GAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003913">
        <property type="entry name" value="Tuberin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018515">
        <property type="entry name" value="Tuberin-type_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027107">
        <property type="entry name" value="Tuberin/Ral-act_asu"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024584">
        <property type="entry name" value="Tuberin_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10063">
        <property type="entry name" value="TUBERIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10063:SF0">
        <property type="entry name" value="TUBERIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11864">
        <property type="entry name" value="DUF3384"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02145">
        <property type="entry name" value="Rap_GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03542">
        <property type="entry name" value="Tuberin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01431">
        <property type="entry name" value="TUBERIN"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF111347">
        <property type="entry name" value="Rap/Ran-GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50085">
        <property type="entry name" value="RAPGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="2" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="7" id="KW-0458">Lysosome</keyword>
    <keyword evidence="6" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="Rap-GAP" evidence="12">
        <location>
            <begin position="1524"/>
            <end position="1751"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1"/>
            <end position="34"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1088"/>
            <end position="1167"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1315"/>
            <end position="1342"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1390"/>
            <end position="1482"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1767"/>
            <end position="1786"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="1"/>
            <end position="16"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="17"/>
            <end position="30"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="1390"/>
            <end position="1406"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004514"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023228"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023765"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00054764"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00070662"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50085"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029638.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1802" mass="201370" checksum="F9EA1630C1528221" modified="2019-05-08" version="1">MAKQQNKDPGLKEKFKSLLGLGPSRPSSKSSEGKQTEFIITAEILKELSIECGLNNRIRAISQICEVAKTKKFEEHAVEAIWKVVADMLQPELPAEARHAVLHLLKAIIQGQGERLGILRAHFFKVIKDYPSNEDLHERLAVFKALTDNGRSITHLEEELAEFVLQWMDVGLSSEFLLVLVNLVKFNSCYLEDYVADMVHMICLLCIQTSSSVDIEVSLQVLDAVVCYNCLPSESLPVFIITLCRTINVKELCEPCWKLMRNLLGTHLGHSAIYNMCRIMEDRAYMTDAALLRGAVFFVGMALWGAHRLYSLKNSPTSVLPSFLKAMTCPNAVVSYEIVLSITRLIKKYGKELQAVTWDILLDIIERLLQQLQSIENQDLKSIVHALLTTVEELCDQNEFHGSEERFFELVERCADQRPESSLLNLITYKAQSIHPAKDGWIHNLKVLMERFFRNECRSAVRIKVLDVLSFVLSINRQFYEEELINVVVISQLAHIPEDKDHQVRKLATQLLVDLAEGCNTHHFNSLLDIIEKVAARSLWAPVDLEERDFLSHSAALEDVKTAVLGLLVILQTKLYSLPASHATRVYEMLINHIELHYKNVYCLPIASSIRLQAFDFLLMLRADSLHRLGLTNKDGAVRFSPYCLCDLVEPEKRVSEKKPAGTLSPPSGSPSVSSQNATLRMGYLPYSLLFRVLLQCLKQETDWKVLKLVLNKLPESLRYKVLFLTSPCKLDQLCSALCSMLTDKKTAERLHGTPDGFSRNDLHLAVVPVLTALISYHNYLDKAKQREMVYCLEHGLIYRCANQCVVALSVCSVEMPDIIIKALPVLIVKLTHISATANMAIPLLEFLSTLARLPHLYRNFAAEQYASVFAISLPYTNPSKFNQYIVCLAHHVIAMWFIRCRLPFRKDFVPYITKGLRSNVLLSFDDTPEKDSFRARSTSLNERPKSSLRIAKNARQGLNNSPPVKELKEASAVDAFRSRSISVSEHVVRSRIQTSITSSSLGSADENSMAQADDNLKNLHLELTETCLDMMARYVFSNFTAVPKRSPVGEFLLAGGRTKTWLVGNKLVTITTSVGTGTRSLLGLDSGELQSSTESSPDPVLQVRQTKEAPAKLESQAGQQVCRGSRNRVRSMSGGHALRVGALDSSAPQGLRSPAASAPRTEKATPGAHALLQKEKTNLAAYVPLLTQGWAEILVRRPTGNTSWLMSLENPLSPFSSDINNMPLQELSNALMAAERFKEHRETALYKSLSVPSSSLATGTSKPSLLQRSNTESAVVLEEGSQLEVDLKETDSLVSQEFEDFKVVCSEQAMAEEKFSKPHNAYSRSSSTSSQEEKSLKSNELVAGGIPIGRVVTSEDGRTLEELSFQPSQPLSKSSSSPELQTLQEVLKDTNAKEATGRLSTELKSKSQSGNLEGEGASSWLSRGDDAGITGTDRLDCNVPSSSPRSPSGHRPRGYTISDSAPSRRGKRIERDAFKSRTAASNAEKVPGINPSFVFLQLYHSPFFGDESNKPLLLPNETFESSVQLLDQIPSYDTHKIAVLYVGEGQSNNEIAILSNEHGSYRYTEFLTGLGKLIELKDCQPDKIYLGGLDVCGEDGQFTYCWHDDIMQAIFHIATLMPTKDLDKYRCDKKRHLGNDFVSIVYNDSGEDFKLGTIKGQFNFVHVIITPLDYDCNLVTLQCRKDMEGLVDTSVAKIVSDKNLPFVARQMALHANMASQVHHSRSNPTDTYPSKWIARLRHIKRLRHRIREETQYQTPGFPLMQMHPSALTKPPPQVPQDPASTYETGQRKRLISSVDDFTEFV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INH9</accession>
    <name>A0A452INH9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Unc-13 homolog A</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000033442.1">
        <property type="protein sequence ID" value="ENSGAGP00000029452.1"/>
        <property type="gene ID" value="ENSGAGG00000020811.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031594">
        <property type="term" value="C:neuromuscular junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098831">
        <property type="term" value="C:presynaptic active zone cytoplasmic component"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042734">
        <property type="term" value="C:presynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030672">
        <property type="term" value="C:synaptic vesicle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043195">
        <property type="term" value="C:terminal bouton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019992">
        <property type="term" value="F:diacylglycerol binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005543">
        <property type="term" value="F:phospholipid binding"/>
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        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017075">
        <property type="term" value="F:syntaxin-1 binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061789">
        <property type="term" value="P:dense core granule priming"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0099525">
        <property type="term" value="P:presynaptic dense core vesicle exocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035249">
        <property type="term" value="P:synaptic transmission, glutamatergic"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016081">
        <property type="term" value="P:synaptic vesicle docking"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016082">
        <property type="term" value="P:synaptic vesicle priming"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd20859">
        <property type="entry name" value="C1_Munc13-2-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd08394">
        <property type="entry name" value="C2A_Munc13"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04027">
        <property type="entry name" value="C2B_Munc13"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.357.50:FF:000001">
        <property type="entry name" value="Protein unc-13 homolog B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000002">
        <property type="entry name" value="Protein unc-13 homolog B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000031">
        <property type="entry name" value="Protein unc-13 homolog B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.60.20:FF:000001">
        <property type="entry name" value="Protein unc-13 homolog B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.357.50">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.1100">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.60.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046349">
        <property type="entry name" value="C1-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000008">
        <property type="entry name" value="C2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010439">
        <property type="entry name" value="MUN_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014770">
        <property type="entry name" value="Munc13_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002219">
        <property type="entry name" value="PKC_DAG/PE"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027080">
        <property type="entry name" value="Unc-13"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037302">
        <property type="entry name" value="Unc-13_C2B"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10480">
        <property type="entry name" value="PROTEIN UNC-13 HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10480:SF1">
        <property type="entry name" value="PROTEIN UNC-13 HOMOLOG A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00130">
        <property type="entry name" value="C1_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00168">
        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06292">
        <property type="entry name" value="MUN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00360">
        <property type="entry name" value="C2DOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00109">
        <property type="entry name" value="C1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00239">
        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM01145">
        <property type="entry name" value="DUF1041"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49562">
        <property type="entry name" value="C2 domain (Calcium/lipid-binding domain, CaLB)"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57889">
        <property type="entry name" value="Cysteine-rich domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50004">
        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51258">
        <property type="entry name" value="MHD1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00479">
        <property type="entry name" value="ZF_DAG_PE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50081">
        <property type="entry name" value="ZF_DAG_PE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0268">Exocytosis</keyword>
    <keyword evidence="2" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <keyword evidence="4" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2" evidence="7">
        <location>
            <begin position="1"/>
            <end position="97"/>
        </location>
    </feature>
    <feature type="domain" description="Phorbol-ester/DAG-type" evidence="8">
        <location>
            <begin position="516"/>
            <end position="566"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="7">
        <location>
            <begin position="622"/>
            <end position="746"/>
        </location>
    </feature>
    <feature type="domain" description="MHD1" evidence="9">
        <location>
            <begin position="1056"/>
            <end position="1199"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="155"/>
            <end position="408"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="191"/>
            <end position="200"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="201"/>
            <end position="220"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="227"/>
            <end position="242"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="251"/>
            <end position="262"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="263"/>
            <end position="275"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="283"/>
            <end position="299"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="307"/>
            <end position="316"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="6">
        <location>
            <begin position="317"/>
            <end position="332"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="333"/>
            <end position="368"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="379"/>
            <end position="396"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50081"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51258"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029452.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="12">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1321" mass="150948" checksum="62D32240E005F6F7" modified="2019-05-08" version="1">TFYSLFAVKKAKFDGAQEKFNTYVTLKVQNVKSTTIAVRGNLPCWEQDFMFEINRLDLGLTVEVWNKGLIWDTMVGTVWIPLQTIRQSNEEGPGEWLTLDSQVIMTDNEISGTKDPTFHRILLDTHFELPLDIPEEEARYWAKKLEQLNAMKDQDDYEFQEEQDKPLPVPGSQCCKYRSPPASNDDPDSTMDDRDSDYRSETSNSIPPPYYTTSQPNASVHQYPMRHQQQSSRDSYMDSVQNYKMDYSDQRPISPTGSSRYASSGELSQGSSQLSDDFDPDNESLRGSELDERDGDSYHSCHSSVSYHDDSPRWEEDFYPEEEDEDYSENEQDFPKVEAEVLRDREADKAKPPEREPYPQPPRKEEKAAYVPEETPEIPQEKPAAKEPPVPERAPEPPKPAEGYREQEDVEVQDAKSRAKANWLKAFNKVCMQLQEGPVQRGLRRAHRVGGGAGCDSSPLIQLFGSVSLQQSLVQSRKAGITSALASSTLNNEELKNHVYKKTLQALIYPISSTTPHNFEVWTATTPTYCYECEGLLWGIARQGMRCTECGVKCHEKCQDLLNADCLQRAAEKSSKHGAEDRTQNIIMVLKDRMKIRERNKPEIFELIQEIFAVSKATHTQQMKAVKQSVLDGTSKWSAKISITVVCAQGLQAKDKTGSSDPYVTVQVGKTKKRTKTIYGNLNPVWEENFHFECHNSSDRIKVRVWDEDDDIKSRVKQRFKRESDDFLGQTIIEVRTLSGEMDVWYNLDKRTDKSAVSGAIRLHISVEIKGEEKVAPYHVQYTCLHENLFHFVTDIQNNGVVKIPEAKGDDAWKVYFDETAQEIVDEFAMRYGVESIYQAMTHFACLSSKYMCPGVPAVMSTLLANINAYYAHTTASTNVSASDRFAASNFGKERFVKLLDQLHNSLRIDLSMYRNNFPASSPERLQDLKSTVDLLTSITFFRMKVQELQSPPRASQVVKDCVKACLNSTYEYIFNNCHELYSREYQTDPTKKGEVPPEEQGPSIKNLDFWSKLITLIVSIIEEDKNSYTPCLNQFPQELNVGKISAEVMWNLFAQDMKYAMEEHDKHRLCKSADYMNLHFKVKWLYNEYVTELPAFKSRVPEYPAWFEPFVIQWLDENEEVSRDFLHGALERDKKDGFQQTSEHALFSCSVVDVFSQLNQSFEIIKKLECPDPQIVGHYMRRFAKTISNVLLQYAEIISKDFALYCSKEKEKVPCILMNNIQQLRVQLEKMFEAMGGKELDAEASDILKELQVKLNNVLDELSRVFATSFQPHIEECVKQMGDILSQVKGTGNVPASACSSVAQDADNVLQPIMDLLDSK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INI0</accession>
    <name>A0A452INI0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Long-chain-fatty-acid--CoA ligase</fullName>
            <ecNumber evidence="20">6.2.1.3</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="20">Catalyzes the conversion of long-chain fatty acids to their active form acyl-CoAs for both synthesis of cellular lipids, and degradation via beta-oxidation.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>(5Z,8Z,11Z,14Z)-eicosatetraenoate + ATP + CoA = (5Z,8Z,11Z,14Z)-eicosatetraenoyl-CoA + AMP + diphosphate</text>
            <dbReference type="Rhea" id="RHEA:19713"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:32395"/>
            <dbReference type="ChEBI" id="CHEBI:33019"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57368"/>
            <dbReference type="ChEBI" id="CHEBI:456215"/>
            <dbReference type="EC" id="6.2.1.15"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="17">
            <dbReference type="Rhea" id="RHEA:19714"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>(E)-hexadec-2-enoate + ATP + CoA = (2E)-hexadecenoyl-CoA + AMP + diphosphate</text>
            <dbReference type="Rhea" id="RHEA:36139"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:33019"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:61526"/>
            <dbReference type="ChEBI" id="CHEBI:72745"/>
            <dbReference type="ChEBI" id="CHEBI:456215"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="18">
            <dbReference type="Rhea" id="RHEA:36140"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>12-hydroxy-(5Z,8Z,10E,14Z)-eicosatetraenoate + ATP + CoA = 12-hydroxy-(5Z,8Z,10E,14Z)-eicosatetraenoyl-CoA + AMP + diphosphate</text>
            <dbReference type="Rhea" id="RHEA:52112"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:33019"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:90718"/>
            <dbReference type="ChEBI" id="CHEBI:136408"/>
            <dbReference type="ChEBI" id="CHEBI:456215"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="15">
            <dbReference type="Rhea" id="RHEA:52113"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="16">
            <text>15-hydroxy-(5Z,8Z,11Z,13E)-eicosatetraenoate + ATP + CoA = 15-hydroxy-(5Z,8Z,11Z,13E)-eicosatetraenoyl-CoA + AMP + diphosphate</text>
            <dbReference type="Rhea" id="RHEA:52116"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:33019"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:78832"/>
            <dbReference type="ChEBI" id="CHEBI:136409"/>
            <dbReference type="ChEBI" id="CHEBI:456215"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="16">
            <dbReference type="Rhea" id="RHEA:52117"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>5-hydroxy-(6E,8Z,11Z,14Z)-eicosatetraenoate + ATP + CoA = 5-hydroxy-(6E,8Z,11Z,14Z)-eicosatetraenoyl-CoA + AMP + diphosphate</text>
            <dbReference type="Rhea" id="RHEA:52108"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:33019"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:65341"/>
            <dbReference type="ChEBI" id="CHEBI:136407"/>
            <dbReference type="ChEBI" id="CHEBI:456215"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="13">
            <dbReference type="Rhea" id="RHEA:52109"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="14">
            <text>a long-chain fatty acid + ATP + CoA = a long-chain fatty acyl-CoA + AMP + diphosphate</text>
            <dbReference type="Rhea" id="RHEA:15421"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:33019"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57560"/>
            <dbReference type="ChEBI" id="CHEBI:83139"/>
            <dbReference type="ChEBI" id="CHEBI:456215"/>
            <dbReference type="EC" id="6.2.1.3"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="14">
            <dbReference type="Rhea" id="RHEA:15422"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>hexadecanoate + ATP + CoA = hexadecanoyl-CoA + AMP + diphosphate</text>
            <dbReference type="Rhea" id="RHEA:30751"/>
            <dbReference type="ChEBI" id="CHEBI:7896"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:33019"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57379"/>
            <dbReference type="ChEBI" id="CHEBI:456215"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="19">
            <dbReference type="Rhea" id="RHEA:30752"/>
        </physiologicalReaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Endoplasmic reticulum membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Mitochondrion membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 20">Belongs to the ATP-dependent AMP-binding enzyme family.</text>
    </comment>
    <dbReference type="EC" id="6.2.1.3" evidence="20"/>
    <dbReference type="AlphaFoldDB" id="A0A452INI0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033409.1">
        <property type="protein sequence ID" value="ENSGAGP00000029420.1"/>
        <property type="gene ID" value="ENSGAGG00000021268.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031966">
        <property type="term" value="C:mitochondrial membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0047676">
        <property type="term" value="F:arachidonate-CoA ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd05927">
        <property type="entry name" value="LC-FACS_euk"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.12780:FF:000006">
        <property type="entry name" value="long-chain-fatty-acid--CoA ligase 6 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.12780">
        <property type="entry name" value="N-terminal domain of ligase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020845">
        <property type="entry name" value="AMP-binding_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000873">
        <property type="entry name" value="AMP-dep_synth/lig_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042099">
        <property type="entry name" value="ANL_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045311">
        <property type="entry name" value="LC-FACS_euk"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43272">
        <property type="entry name" value="LONG-CHAIN-FATTY-ACID--COA LIGASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43272:SF54">
        <property type="entry name" value="LONG-CHAIN-FATTY-ACID--COA LIGASE 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00501">
        <property type="entry name" value="AMP-binding"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56801">
        <property type="entry name" value="Acetyl-CoA synthetase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00455">
        <property type="entry name" value="AMP_BINDING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 20" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="6" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="7 20" id="KW-0276">Fatty acid metabolism</keyword>
    <keyword evidence="4 20" id="KW-0436">Ligase</keyword>
    <keyword evidence="10 20" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="9" id="KW-0460">Magnesium</keyword>
    <keyword evidence="12 21" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="5 20" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="21" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="21" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="21"/>
            <end position="45"/>
        </location>
    </feature>
    <feature type="domain" description="AMP-dependent synthetase/ligase" evidence="22">
        <location>
            <begin position="117"/>
            <end position="520"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004325"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004586"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00006432"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022598"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022832"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00024469"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00024484"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00024495"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00024532"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00024548"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00024565"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00049139"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="RuleBase" id="RU369030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="Pfam" id="PF00501"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029420.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="698" mass="78360" checksum="06686A822376A1C1" modified="2019-05-08" version="1">MQAQEILRSLRLPEFDNVSQFFHSLPATTLVGIGAFAAVIAYWFASRPKVLKPPCDLLMQSEEAEGWEGARRSVIGDSTELLTHYYDDARTMYEVFRRGFNISENGPCLGFRKPKQPYQWLSYKEVSERAEALGSGLLQQGCKPSASQFIGVFSQNRPEWIISELACYTYSMVVVPLYDTLGHGAIRYIINTADISTVICDKPEKARILLDHMEKRETPGLKSIILMDPFEKDLKERGKRCGVHVQSMQEVEASGRESWHVPVPPQPEDLSIVCFTSGTTGNPKGAMLTHRNVVADFSGFLKVTEKVIFPRQDDVLISFLPLAHMFERVIQSVVYCHGGRIGFFQGDIRLLSDDMKALHPTIFPVVPRLLNRMYDKIFSQADTPLKRWLLEFAAKRKKAEVQSGIIRNDSLWDKLFFNKIQASLGGCVRMIVTGAAPASPTVLGFLRAALGCQVYEGYGQTECTAGCTFTTPGDWTSGHVGAPLPCNLIKLMDVEEMNYFASKGEGEICVKGPNVFKGYLKDGEKTAEALDKEGWLHTGDIGKWLPTGTLKIIDRKKHIFKLAQGEYIAPEKIENIYIRSEPIAQIYVHGDSLQAFLVGIVVPDSEVMPGWARKRGFEGTYAELCKNTELTQAIMEDMVRLGKEGGLHSFEQVKAIYIHSDMFSIENGLLTPTLKARRRELRDYFKKQIEELYSSVSV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INI1</accession>
    <name>A0A452INI1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Cytoplasmic FMR1 interacting protein 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="1 2">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="2" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="1" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="1" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452INI1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033518.1">
        <property type="protein sequence ID" value="ENSGAGP00000029518.1"/>
        <property type="gene ID" value="ENSGAGG00000020525.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031267">
        <property type="term" value="F:small GTPase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030833">
        <property type="term" value="P:regulation of actin filament polymerization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008081">
        <property type="entry name" value="Cytoplasmic_FMR1-int"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12195">
        <property type="entry name" value="CYTOPLASMIC FMR1-INTERACTING PROTEIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05994">
        <property type="entry name" value="FragX_IP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF008153">
        <property type="entry name" value="FMR1_interacting"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000313" key="1">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029518.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="3">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="944" mass="109100" checksum="B990BB0B55110B9B" modified="2019-05-08" version="1">MTAQVTLEDALSNVDLLEELPLPDQQPCIEPPPSSLLYQPNFNTNFEDRNAFVTGIARYIEQATVHSNMVMGFGLYLMDGSVSNIYKLDAKKRINLTKIDKFFKQLQVVPLFGDMQIELARYIKTSAHYEENKSRWTCTSSSSSPQYNICEQMIQIREDHMRFISELARYSNSEVVTGSGRQEAQKTDAEYRKLFDLSLQGLQLLSQWSAHVMEVYSWKLVHPTDKYSNKDCPDNAEEYERATRYNYTSEEKFALVEVIAMIKGLQVLMGRMESVFNHAIRHTIYAALQDFSQVTLREPLRQAIKKKKNVIQSVLQAIRKTVCDWEAGHEPFNDPALRGEKDPKSGFDIKVPRYVLYSLDLYNDSAHYALTKFKKQFLYDEIEAEVNLCFDQFVYKLADQIFAYYKIMAGSLLLDKRLRSECKNQGATIHLPPSNRYETLLKQRHVQLLGRSIDLNRLITQRISAAMYKSMELAIGRFESEDLTSIVELDGLVEINKMTHKLLSRYMTLDSFDAMFREANHNVSAPYGRITLHVFWELNYDFLPNYCYNGSTNRFVRTVLPFSQEFQRDKQPNAQPQYLHGSKALNLTYSSIYSNYRNFVGPPHFKVICRLLGYQGIAVVMEELLKVVKSLLQGTILQYVKTLMEVMPKICRLPRHEYGSPGILEFFHHQLKDIVEYAELKTVCFQNLREVGNAVLFCLLIEQSLSLEEVCDLLHAAPFQNILPRVHVKEGERLDAKMKRLESKYAPLHLVPLIERLGTPQQIAIAREGDLLTKERLCCGLSMFEVILTRIRTFLDDPIWRGPLPSNGVMHVDECVEFHRLWSAMQFVYCIPVGTHEFTVEQCFGDGLHWAGCMIIVLLGQQRRFDVLDFCYHLLKVQKHDGKDEVIKNVPLKKMVERIRKFQILNNEIITILDKYMKSGDGEGTPVEHVRCFQPPIHQSLASN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INI3</accession>
    <name>A0A452INI3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Centrosomal protein of 170 kDa</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Plays a role in microtubule organization. Required for centriole subdistal appendage assembly.</text>
    </comment>
    <comment type="subunit">
        <text evidence="10">Interacts with CCDC68 and CCDC120; leading to recruitment to centrosomes. Interacts with PLK1. Interacts with NIN. Interacts with FHDC1. Interacts with CCDC61. Interacts with TBK1; efficient complex formation may be dependent on the presence of CCDC61.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Microtubule organizing center</location>
            <location evidence="1">Centrosome</location>
            <location evidence="1">Centriole</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
            <location evidence="2">Spindle</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the CEP170 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000033415.1">
        <property type="protein sequence ID" value="ENSGAGP00000029426.1"/>
        <property type="gene ID" value="ENSGAGG00000021269.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005814">
        <property type="term" value="C:centriole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005874">
        <property type="term" value="C:microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005819">
        <property type="term" value="C:spindle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="CDD" id="cd22724">
        <property type="entry name" value="FHA_Cep170A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.200.20:FF:000018">
        <property type="entry name" value="Centrosomal protein of 170 kDa"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.200.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051176">
        <property type="entry name" value="Cent_Immune-Sig_Mod"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029300">
        <property type="entry name" value="CEP170_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000253">
        <property type="entry name" value="FHA_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008984">
        <property type="entry name" value="SMAD_FHA_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15715">
        <property type="entry name" value="CENTROSOMAL PROTEIN OF 170 KDA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15715:SF17">
        <property type="entry name" value="CENTROSOMAL PROTEIN OF 170 KDA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15308">
        <property type="entry name" value="CEP170_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00498">
        <property type="entry name" value="FHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00240">
        <property type="entry name" value="FHA"/>
        <property type="match status" value="1"/>
    </dbReference>
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</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INI5</accession>
    <name>A0A452INI5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Long-chain-fatty-acid--CoA ligase</fullName>
            <ecNumber evidence="14">6.2.1.3</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Catalyzes the conversion of long-chain fatty acids to their active form acyl-CoAs for both synthesis of cellular lipids, and degradation via beta-oxidation.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>(5Z,8Z,11Z,14Z)-eicosatetraenoate + ATP + CoA = (5Z,8Z,11Z,14Z)-eicosatetraenoyl-CoA + AMP + diphosphate</text>
            <dbReference type="Rhea" id="RHEA:19713"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:32395"/>
            <dbReference type="ChEBI" id="CHEBI:33019"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57368"/>
            <dbReference type="ChEBI" id="CHEBI:456215"/>
            <dbReference type="EC" id="6.2.1.15"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="11">
            <dbReference type="Rhea" id="RHEA:19714"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="12">
            <text>(E)-hexadec-2-enoate + ATP + CoA = (2E)-hexadecenoyl-CoA + AMP + diphosphate</text>
            <dbReference type="Rhea" id="RHEA:36139"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
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            <dbReference type="ChEBI" id="CHEBI:61526"/>
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        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="12">
            <dbReference type="Rhea" id="RHEA:36140"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="9">
            <text>12-hydroxy-(5Z,8Z,10E,14Z)-eicosatetraenoate + ATP + CoA = 12-hydroxy-(5Z,8Z,10E,14Z)-eicosatetraenoyl-CoA + AMP + diphosphate</text>
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            <dbReference type="ChEBI" id="CHEBI:90718"/>
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        </reaction>
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            <dbReference type="Rhea" id="RHEA:52113"/>
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    </comment>
    <comment type="catalytic activity">
        <reaction evidence="10">
            <text>15-hydroxy-(5Z,8Z,11Z,13E)-eicosatetraenoate + ATP + CoA = 15-hydroxy-(5Z,8Z,11Z,13E)-eicosatetraenoyl-CoA + AMP + diphosphate</text>
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    <comment type="catalytic activity">
        <reaction evidence="7">
            <text>5-hydroxy-(6E,8Z,11Z,14Z)-eicosatetraenoate + ATP + CoA = 5-hydroxy-(6E,8Z,11Z,14Z)-eicosatetraenoyl-CoA + AMP + diphosphate</text>
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            <dbReference type="ChEBI" id="CHEBI:30616"/>
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            <dbReference type="ChEBI" id="CHEBI:65341"/>
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    <comment type="catalytic activity">
        <reaction evidence="8">
            <text>a long-chain fatty acid + ATP + CoA = a long-chain fatty acyl-CoA + AMP + diphosphate</text>
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            <dbReference type="ChEBI" id="CHEBI:30616"/>
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            <dbReference type="EC" id="6.2.1.3"/>
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            <dbReference type="Rhea" id="RHEA:15422"/>
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    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>hexadecanoate + ATP + CoA = hexadecanoyl-CoA + AMP + diphosphate</text>
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            <dbReference type="ChEBI" id="CHEBI:7896"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
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            <dbReference type="ChEBI" id="CHEBI:57379"/>
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        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="13">
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        </physiologicalReaction>
    </comment>
    <comment type="similarity">
        <text evidence="1 14">Belongs to the ATP-dependent AMP-binding enzyme family.</text>
    </comment>
    <dbReference type="EC" id="6.2.1.3" evidence="14"/>
    <dbReference type="AlphaFoldDB" id="A0A452INI5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033414.1">
        <property type="protein sequence ID" value="ENSGAGP00000029425.1"/>
        <property type="gene ID" value="ENSGAGG00000021268.1"/>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005783">
        <property type="term" value="C:endoplasmic reticulum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0047676">
        <property type="term" value="F:arachidonate-CoA ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd05927">
        <property type="entry name" value="LC-FACS_euk"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.12780">
        <property type="entry name" value="N-terminal domain of ligase-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020845">
        <property type="entry name" value="AMP-binding_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000873">
        <property type="entry name" value="AMP-dep_synth/lig_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042099">
        <property type="entry name" value="ANL_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045311">
        <property type="entry name" value="LC-FACS_euk"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43272">
        <property type="entry name" value="LONG-CHAIN-FATTY-ACID--COA LIGASE"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="LONG-CHAIN-FATTY-ACID--COA LIGASE 6"/>
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    <sequence length="698" mass="78267" checksum="FEF0F84277495805" modified="2019-05-08" version="1">MDQTALFLGQHSSMFIKEFALSIFEKMQAQEILRSLRLPEFDNVSQFFHSLPATTLVGIGAFAAVIAYWFASRPKVLKPPCDLLMQSEEAEGWEGARRSVIGDSTELLTHYYDDARTMYEVFRRGFNISENGPCLGFRKPKQPYQWLSYKEVSERAEALGSGLLQQGCKPSASQFIGVFSQNRPEWIISELACYTYSMVVVPLYDTLGHGAIRYIINTADISTVICDKPEKARILLDHMEKRETPGLKSIILMDPFEKDLKERGKRCGVHVQSMQEVEASGRESWHVPVPPQPEDLSIVCFTSGTTGNPKGAMLTHRNVVADFSGFLKVTESVVYCHGGRIGFFQGDIRLLSDDMKALHPTIFPVVPRLLNRMYDKIFSQADTPLKRWLLEFAAKRKKAEVQSGIIRNDSLWDKLFFNKIQASLGGCVRMIVTGAAPASPTVLGFLRAALGCQVYEGYGQTECTAGCTFTTPGDWTSGHVGAPLPCNLIKLMDVEEMNYFASKGEGEICVKGPNVFKGYLKDGEKTAEALDKEGWLHTGDIGKWLPTGTLKIIDRKKHIFKLAQGEYIAPEKIENIYIRSEPIAQIYVHGDSLQAFLVGIVVPDSEVMPGWARKRGFEGTYAELCKNTELTQAIMEDMVRLGKEGGLHSFEQVKAIYIHSDMFSIENGLLTPTLKARRRELRDYFKKQIEELYSSVSV</sequence>
</entry>
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    <accession>A0A452INI8</accession>
    <name>A0A452INI8_9SAUR</name>
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        <recommendedName>
            <fullName evidence="14">PDZ domain-containing protein</fullName>
        </recommendedName>
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        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
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        <lineage>
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            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
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            <taxon>Testudines</taxon>
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            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
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            <taxon>Gopherus</taxon>
        </lineage>
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        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
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                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
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        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
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        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
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        <subcellularLocation>
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    </comment>
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    <keyword evidence="10" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
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    <keyword evidence="6" id="KW-0524">Neurogenesis</keyword>
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        </location>
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    </feature>
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        </location>
    </feature>
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        </location>
    </feature>
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        </location>
    </feature>
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        </location>
    </feature>
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        </location>
    </feature>
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    </evidence>
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    </evidence>
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        </source>
    </evidence>
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    </evidence>
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        </source>
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        </source>
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        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
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        </source>
    </evidence>
    <sequence length="1199" mass="133867" checksum="859B6ABD672EE18D" modified="2019-05-08" version="1">MMKTESSGERSTLRSASPHRNAYRTEFQALKSTFDKPKSDGDQKTKDEGEVSQQSRGRKYGSNVNRIKNLFMQMGMEPSESTGVTPKTRGKSGPLSPQRIRPKEFVEKADGSVVKLESSVSERISRFDTVHGGPSYSKFTETRKMFERNVHEMGHSNRYSPKKERVVSNELQDEWCSSKSNRGSTDSLDSLSSRTETISPTVSQLSAVFENTDSHTVIVLEKSENNEEYSVTGHYPLNLSSTVENLSSTVANLDGFSPLKDASAWSSPAKQTTSSVSAETFQQNSTSSTIREASNSTLPVSKTNEEIRKNKEASFDPVDVCAATKQELASINSEKSEESYTRTKEKIDTEVSLQQKEHSQNAENATESPYDDASGKEVQKDLNNFQSSHVYMHSDYNVYRVRSRYNSDWGETGTEQDDQDDSDENNYYEPDMEYAEVIGLPAEDEIPANRRIKFSSAPIKVFNTYSNEDYDRRNDEVDPVAASAEYELEKRVEKLELFPVELGKDEDGLGISIIGMGVGADAGLEKLGIFVKTVTEGGAAERDGRIQVNDQIVEVDGISLVGVTQNFAATVLRNTKGNVRFVIGREKPGQVSEVAQLISQTLEQERRQRELLEQHYAQYDADDDETGEYATDEEDEDMGPIVPGADMAIEVFDLPENDDMFSPTELDTTKLAHKFKELQIKHAVTEAEIQKLKTKLQAAENEKVRWEREKTQLQQNIEENKERMMKLESYWIEAQTLCHTVNEHLKETQSQYQALEKKYNKAKKLIKDFQQKELDFIKRQEAERKKIEELEKAHLAEVQGLQARIRDLEAEVFKLLKQNGSQVNNNNNIFERQTSFGEVSRGDPVESLDVEQVSCLDDLSQDFNEAVPETERLDSKALKTRAQLSVRNKRQRPTRTRLYDSISSTDGEDSLERKPSDSYSSPMHISKSLLPICVRASSPASDSGASSLSPVASSAAFSFDNVAENQEGGQQHKGGVLSHYARGDTPLSSPSKSSHSSETSPLHHPTNRNVLQDKDPGSFLLNRTTQEMLESSASNASMSERSSSLPYSIPFTWYGGSGRGSNSSSNLPSSSSSTEPSSENLSPAKKGSKNFTFTDDFSPSSTSSADLSGLGAEPKPPGFSHSLALSSDETLGMTSSQDRAIIKKKLKEMKVSIERARKAQEKMEKQREKLRKKEQERLQRKSKKLDKSSSDATEGANEQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INI9</accession>
    <name>A0A452INI9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
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            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
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                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
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            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
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        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
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                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
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        <text evidence="9">Plays a role in microtubule organization. Required for centriole subdistal appendage assembly.</text>
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        <text evidence="10">Interacts with CCDC68 and CCDC120; leading to recruitment to centrosomes. Interacts with PLK1. Interacts with NIN. Interacts with FHDC1. Interacts with CCDC61. Interacts with TBK1; efficient complex formation may be dependent on the presence of CCDC61.</text>
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        </source>
    </evidence>
    <sequence length="1633" mass="180479" checksum="66553DFE44DB2B1F" modified="2019-05-08" version="1">MSLTSWFLVSSGGTRHRLPREMIFVGRDDCELMLQSRSVDKQHAVVNYDASTDEHLVKDLGSLNGTFVNDVRIPEQTYITLKLEDKLRFGYDTNLFTVVRGEMRVPEEALKHEKFTSQLQLSQKSSEAEGSKPVSSTGTEPNVPDATGEVHHKATEALKSEEKSMDLSAMPRGTPLYGQPSWWGEDEADEKNGCKRDIKHEEKIHETGASGCSTDAKQVVEKSSAANEEELYPFCREPSYFEIPTKEFHQPSQIAESTIHEIPTKDTQTSHAAGAGHASFTIEFDDTTPGKVTIKDHVTKFTSDQRHKSKKSPLGGQDLPGLQTVMMAAESKVADWLAQNNPPRMIWEATEEDSKSIKSDVPVYLKRLKGNKHDDGTQSDSENAGARRHYNKRATVDEHLSHQHLEQKKSYQKAHTTEKHEEQAAISQTAFMIEFFDEDHPRKRRSYSFSQNVGALCSETPSSTPHVRAEKVKGTSGDIKGVSLPLQTSRVAHGVHAKLLKQKSEDPPVPLPFLQTALLRSSGSLGHRPSQNQEMDKKLKSQLVSAATEKDNDDDQSDKGTYTIELENPNAEEMEARKMIDKEGSSASGSKRWVSQWASLAANHTRHDQDDMRLESSAPAPRENDTDISESGVSVRSTSSATSVTSQGERKRRTLPQLPKEEKATESSRAKVTPHQRSEIGEKQDTELQEKETPTHASQKEMPQDAGSRSMAKANRTMNGLSSKPGGDNKVSSHPISCSSFSKEKSEIVRETSVVKQALAKIQQQEQKEQAHRTSTKVSSAKLLANQIEKGREETSTPHKMLDNHQEKAPGHSTSKAEMKIVQSEGKRKKTEEIIKGQTPKGFGGEKKESSKPLVRQGSFTIDKPSTNIPIELIPHINKQTGSTPPSFSLTSTNRIRERSDSVDTDSSLDTTLILKDTEAVMAFLEAKLREENKTDEGPETPSYNRDNSISPESDVDTASTISLVTGDTERKTTQKRKSFTTLYKDRCSTSSPSKDVLKSSATNAREKIDKKTKSRSSDAGSRAEGRKLVQPSGRMRQPSVDLTDDDQTSSVPHSAISDILSSDQETYSGKSHGRGHFTSADELLHSKMEGSKSAKSKISPVTTGQSSKSITLPRPRPTRTSLLRRARLGEASDSELADADKASVASEVSTTSSTSKPPSGRRNISRIDLLAQPRRIRLGSLSARSDSEATITRSAASSRTAEAIIRSSGRLTPSADSSKVSARTRANSISRLSDSKSKSLTSAHNSPSEKSQLLPDPDPAVETYSVNSRWRRFPTDYASTSEDEFGSNRNSPKHTRLRTSPALKTPRLQSSGTAMQTSNTFKHRIKEQEDYIRDWTAHREEIARISQDLALIAREINDVAGEIDSVTSSGTAPSTTVSTAATTPGSAIDTREEVGDLHGEMHKLVDRVFDESLNFRKIPPLVHAKPPEGNSRSNDSRPQLTDALDPPTITRRRTWSRDEVMGDSLLLSSVFQFSRKIRQSIDKTAGKIRILFKDKDRNWEEIESKLRTESEVPIVKTSSMEISSILQELKRVEKQLQVINAMIDPDGTLDALSNLGFTSPVLPAQPKHKSSPVSHSTRAQIQPNCQPEARALRPAVTSVSTAFDNVESKPDFSIHFNRFNPDGEEEDPSMRE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INJ8</accession>
    <name>A0A452INJ8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">HECT-type E3 ubiquitin transferase</fullName>
            <ecNumber evidence="4">2.3.2.26</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>S-ubiquitinyl-[E2 ubiquitin-conjugating enzyme]-L-cysteine + [acceptor protein]-L-lysine = [E2 ubiquitin-conjugating enzyme]-L-cysteine + N(6)-ubiquitinyl-[acceptor protein]-L-lysine.</text>
            <dbReference type="EC" id="2.3.2.26"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="3">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="2.3.2.26" evidence="4"/>
    <dbReference type="AlphaFoldDB" id="A0A452INJ8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033533.1">
        <property type="protein sequence ID" value="ENSGAGP00000029533.1"/>
        <property type="gene ID" value="ENSGAGG00000021328.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00143"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019871">
        <property type="term" value="F:sodium channel inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061630">
        <property type="term" value="F:ubiquitin protein ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016567">
        <property type="term" value="P:protein ubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048814">
        <property type="term" value="P:regulation of dendrite morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051049">
        <property type="term" value="P:regulation of transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006511">
        <property type="term" value="P:ubiquitin-dependent protein catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd04033">
        <property type="entry name" value="C2_NEDD4_NEDD4L"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00078">
        <property type="entry name" value="HECTc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00201">
        <property type="entry name" value="WW"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1750.10:FF:000026">
        <property type="entry name" value="E3 ubiquitin-protein ligase HACE1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.2160.10:FF:000001">
        <property type="entry name" value="E3 ubiquitin-protein ligase NEDD4-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.2410.10:FF:000001">
        <property type="entry name" value="E3 ubiquitin-protein ligase NEDD4-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1750.10:FF:000001">
        <property type="entry name" value="E3 ubiquitin-protein ligase NEDD4-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.70.10:FF:000006">
        <property type="entry name" value="E3 ubiquitin-protein ligase NEDD4-like protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.70.10:FF:000008">
        <property type="entry name" value="E3 ubiquitin-protein ligase NEDD4-like protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.70.10:FF:000025">
        <property type="entry name" value="E3 ubiquitin-protein ligase NEDD4-like protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000047">
        <property type="entry name" value="Putative E3 ubiquitin-protein ligase NEDD4-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.20.70.10">
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.2160.10">
        <property type="entry name" value="Hect, E3 ligase catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.2410.10">
        <property type="entry name" value="Hect, E3 ligase catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1750.10">
        <property type="entry name" value="Hect, E3 ligase catalytic domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000008">
        <property type="entry name" value="C2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024928">
        <property type="entry name" value="E3_ub_ligase_SMURF1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050409">
        <property type="entry name" value="E3_ubiq-protein_ligase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000569">
        <property type="entry name" value="HECT_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035983">
        <property type="entry name" value="Hect_E3_ubiquitin_ligase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001202">
        <property type="entry name" value="WW_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036020">
        <property type="entry name" value="WW_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11254:SF440">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE NEDD-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11254">
        <property type="entry name" value="HECT DOMAIN UBIQUITIN-PROTEIN LIGASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00168">
        <property type="entry name" value="C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00632">
        <property type="entry name" value="HECT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00397">
        <property type="entry name" value="WW"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF001569">
        <property type="entry name" value="E3_ub_ligase_SMURF1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00360">
        <property type="entry name" value="C2DOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00239">
        <property type="entry name" value="C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00119">
        <property type="entry name" value="HECTc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00456">
        <property type="entry name" value="WW"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49562">
        <property type="entry name" value="C2 domain (Calcium/lipid-binding domain, CaLB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56204">
        <property type="entry name" value="Hect, E3 ligase catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51045">
        <property type="entry name" value="WW domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50004">
        <property type="entry name" value="C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50237">
        <property type="entry name" value="HECT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01159">
        <property type="entry name" value="WW_DOMAIN_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50020">
        <property type="entry name" value="WW_DOMAIN_2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="7" id="KW-0808">Transferase</keyword>
    <keyword evidence="10" id="KW-0832">Ubl conjugation</keyword>
    <keyword evidence="9 12" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="C2" evidence="14">
        <location>
            <begin position="4"/>
            <end position="124"/>
        </location>
    </feature>
    <feature type="domain" description="WW" evidence="15">
        <location>
            <begin position="176"/>
            <end position="209"/>
        </location>
    </feature>
    <feature type="domain" description="WW" evidence="15">
        <location>
            <begin position="348"/>
            <end position="381"/>
        </location>
    </feature>
    <feature type="domain" description="WW" evidence="15">
        <location>
            <begin position="460"/>
            <end position="493"/>
        </location>
    </feature>
    <feature type="domain" description="WW" evidence="15">
        <location>
            <begin position="511"/>
            <end position="535"/>
        </location>
    </feature>
    <feature type="domain" description="HECT" evidence="16">
        <location>
            <begin position="622"/>
            <end position="956"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="137"/>
            <end position="184"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="227"/>
            <end position="293"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="389"/>
            <end position="458"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="13">
        <location>
            <begin position="147"/>
            <end position="158"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="13">
        <location>
            <begin position="264"/>
            <end position="273"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="13">
        <location>
            <begin position="423"/>
            <end position="434"/>
        </location>
    </feature>
    <feature type="active site" description="Glycyl thioester intermediate" evidence="11 12">
        <location>
            <position position="924"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000885"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00012485"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PIRSR" id="PIRSR001569-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00104"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50020"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50237"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029533.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="957" mass="110407" checksum="ECEAB320E3A0680C" modified="2019-05-08" version="1">MASGAVEPVYGLSEDEDESRLLRVKVVSGIDLAKKDIFGASDPYVKLSLYVADENRELALVQTKTIKKTLNPKWNEEFYFRVNPTNHRLLFEVFDENRLTEDPTMERPYTFKDFLLRPRSHKSRVKGFLRLKMAYMPKNGGQEEETTEQREESERAWEVVDSNDSVSHHQEELPPPPLPPGWEEKVDNLGRTYYVNHNNRTTQWHRPSLIDVASDSDNNIRQINQEAAHRRFRSRRHISEDLEPEPSESGDASEPWETISEEASISGDSLSLSLPPPPASPVSRTSPQELSDELNRRLQVTPDSNGEQFGSLIQRDPSSRLRSCSVTDAVAEQSHLSLPSVAYVHTTPGLPSGWEERKDAKGRTYYVNHNNRTTTWTRPIMQLAEDGAVGSAANSNNHLSEPQIRRPRSLSSPTVTLSAPLEFAKDSPVRRAVKDTLSNPQSPQPSPYNSPKPQHKVTQSFLPPGWEMRIAPNGRPFFIDHNTKTTTWEDPRLKFPVHLRSKASLNPNDLGPLPPGWEERIHLDGRTFYIDHRSQVLICGDIDTRDLVPSYDSKITQWEDPRLQNPAITGPAVPYSREFKQKYDYFRKKLKKPADIPNRFEMKLHRNNIFEESYRRIMSVKRSDVLKARLWIEFESEKGLDYGGVAREWFFLLSKEMFNPYYGLFEYSATDNYTLQINPNSGLCNEDHLSYFTFIGRVAGLAVFHGKLLDGFFIRPFYKMMLGKQITLKDMESVDSEYYNSLKWILENDPTELDLMFCIDEENFGQTYQVDLKPNGSEIMVTNENKREYIDLVIQWRFVNRVQKQMNAFLEGFTELLPIDLIKIFDENELELLMCGLGDVDVNDWRQHTIYKNGYCPNHPVIQWYWKAVLLMDAEKRIRLLQFVTGTSRVPMNGFAELYGSNGPQLFTIEQWGSPEKLPRAHTCFNRLDLPPYESFEDLREKLLMAVENAQGFEGVD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INK0</accession>
    <name>A0A452INK0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Unc-13 homolog A</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
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            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
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    <dbReference type="PANTHER" id="PTHR10480">
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="DUF1041"/>
        <property type="match status" value="1"/>
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        <property type="match status" value="1"/>
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        <property type="match status" value="2"/>
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        <property type="entry name" value="ZF_DAG_PE_2"/>
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    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0268">Exocytosis</keyword>
    <keyword evidence="2" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <keyword evidence="4" id="KW-0863">Zinc-finger</keyword>
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            <begin position="1"/>
            <end position="97"/>
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    <feature type="domain" description="Phorbol-ester/DAG-type" evidence="8">
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        <location>
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    </feature>
    <feature type="domain" description="MHD1" evidence="9">
        <location>
            <begin position="1069"/>
            <end position="1212"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="155"/>
            <end position="408"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="191"/>
            <end position="200"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
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            <begin position="201"/>
            <end position="220"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
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            <end position="242"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
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            <end position="262"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
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    </feature>
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        </source>
    </evidence>
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        </source>
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        <source>
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        </source>
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        </source>
    </evidence>
    <evidence type="ECO:0008006" key="12">
        <source>
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        </source>
    </evidence>
    <sequence length="1334" mass="152506" checksum="9E06919D9147145E" modified="2019-05-08" version="1">TFYSLFAVKKAKFDGAQEKFNTYVTLKVQNVKSTTIAVRGNLPCWEQDFMFEINRLDLGLTVEVWNKGLIWDTMVGTVWIPLQTIRQSNEEGPGEWLTLDSQVIMTDNEISGTKDPTFHRILLDTHFELPLDIPEEEARYWAKKLEQLNAMKDQDDYEFQEEQDKPLPVPGSQCCKYRSPPASNDDPDSTMDDRDSDYRSETSNSIPPPYYTTSQPNASVHQYPMRHQQQSSRDSYMDSVQNYKMDYSDQRPISPTGSSRYASSGELSQGSSQLSDDFDPDNESLRGSELDERDGDSYHSCHSSVSYHDDSPRWEEDFYPEEEDEDYSENEQDFPKVEAEVLRDREADKAKPPEREPYPQPPRKEEKAAYVPEETPEIPQEKPAAKEPPVPERAPEPPKPAEGYREQEDVEVQDAKSRAKANWLKAFNKVCMQLQEGPVQRGLRRGRLTTFPSHCSPGPGGGLIIIDSMPDIRKRKPIPLVSDLSLVQSRKAGITSALASSTLNNEELKNHVYKKTLQALIYPISSTTPHNFEVWTATTPTYCYECEGLLWGIARQGMRCTECGVKCHEKCQDLLNADCLQRAAEKSSKHGAEDRTQNIIMVLKDRMKIRERNKPEIFELIQEIFAVSKATHTQQMKAVKQSVLDGTSKWSAKISITVVCAQGLQAKDKTGSSDPYVTVQVGKTKKRTKTIYGNLNPVWEENFHFECHNSSDRIKVRVWDEDDDIKSRVKQRFKRESDDFLGQTIIEVRTLSGEMDVWYNLDKRTDKSAVSGAIRLHISVEIKGEEKVAPYHVQYTCLHENLFHFVTDIQNNGVVKIPEAKGDDAWKVYFDETAQEIVDEFAMRYGVESIYQAMTHFACLSSKYMCPGVPAVMSTLLANINAYYAHTTASTNVSASDRFAASNFGKERFVKLLDQLHNSLRIDLSMYRNNFPASSPERLQDLKSTVDLLTSITFFRMKVQELQSPPRASQVVKDCVKACLNSTYEYIFNNCHELYSREYQTDPTKKGEVPPEEQGPSIKNLDFWSKLITLIVSIIEEDKNSYTPCLNQFPQELNVGKISAEVMWNLFAQDMKYAMEEHDKHRLCKSADYMNLHFKVKWLYNEYVTELPAFKSRVPEYPAWFEPFVIQWLDENEEVSRDFLHGALERDKKDGFQQTSEHALFSCSVVDVFSQLNQSFEIIKKLECPDPQIVGHYMRRFAKTISNVLLQYAEIISKDFALYCSKEKEKVPCILMNNIQQLRVQLEKMFEAMGGKELDAEASDILKELQVKLNNVLDELSRVFATSFQPHIEECVKQMGDILSQVKGTGNVPASACSSVAQDADNVLQPIMDLLDSK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INK1</accession>
    <name>A0A452INK1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1 10">Catechol O-methyltransferase</fullName>
            <ecNumber evidence="1 10">2.1.1.6</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">Catalyzes the O-methylation, and thereby the inactivation, of catecholamine neurotransmitters and catechol hormones.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="10">
            <text>a catechol + S-adenosyl-L-methionine = a guaiacol + S-adenosyl-L-homocysteine + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17877"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:33566"/>
            <dbReference type="ChEBI" id="CHEBI:57856"/>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
            <dbReference type="ChEBI" id="CHEBI:134251"/>
            <dbReference type="EC" id="2.1.1.6"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="10 13">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
        <text evidence="10 13">Binds 1 Mg(2+) ion per subunit.</text>
    </comment>
    <comment type="similarity">
        <text evidence="9 10">Belongs to the class I-like SAM-binding methyltransferase superfamily. Cation-dependent O-methyltransferase family.</text>
    </comment>
    <dbReference type="EC" id="2.1.1.6" evidence="1 10"/>
    <dbReference type="AlphaFoldDB" id="A0A452INK1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033680.1">
        <property type="protein sequence ID" value="ENSGAGP00000029663.1"/>
        <property type="gene ID" value="ENSGAGG00000021411.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030424">
        <property type="term" value="C:axon"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030425">
        <property type="term" value="C:dendrite"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016206">
        <property type="term" value="F:catechol O-methyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000287">
        <property type="term" value="F:magnesium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042424">
        <property type="term" value="P:catecholamine catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032502">
        <property type="term" value="P:developmental process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042417">
        <property type="term" value="P:dopamine metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032259">
        <property type="term" value="P:methylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="CDD" id="cd02440">
        <property type="entry name" value="AdoMet_MTases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.150:FF:000054">
        <property type="entry name" value="Catechol O-methyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.150">
        <property type="entry name" value="Vaccinia Virus protein VP39"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017128">
        <property type="entry name" value="Catechol_O-MeTrfase_euk"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029063">
        <property type="entry name" value="SAM-dependent_MTases_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002935">
        <property type="entry name" value="SAM_O-MeTrfase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43836:SF3">
        <property type="entry name" value="CATECHOL O-METHYLTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43836">
        <property type="entry name" value="CATECHOL O-METHYLTRANSFERASE 1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01596">
        <property type="entry name" value="Methyltransf_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF037177">
        <property type="entry name" value="Catechol_O-mtfrase_euk"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53335">
        <property type="entry name" value="S-adenosyl-L-methionine-dependent methyltransferases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51682">
        <property type="entry name" value="SAM_OMT_I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 10" id="KW-0128">Catecholamine metabolism</keyword>
    <keyword evidence="6 10 13" id="KW-0460">Magnesium</keyword>
    <keyword evidence="5 10 13" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="2 10" id="KW-0489">Methyltransferase</keyword>
    <keyword evidence="7 10" id="KW-0531">Neurotransmitter degradation</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 10 11" id="KW-0949">S-adenosyl-L-methionine</keyword>
    <keyword evidence="3 10" id="KW-0808">Transferase</keyword>
    <feature type="binding site" evidence="11">
        <location>
            <position position="92"/>
        </location>
        <ligand>
            <name>S-adenosyl-L-methionine</name>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="11">
        <location>
            <position position="122"/>
        </location>
        <ligand>
            <name>S-adenosyl-L-methionine</name>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="11">
        <location>
            <position position="140"/>
        </location>
        <ligand>
            <name>S-adenosyl-L-methionine</name>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <position position="191"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="11">
        <location>
            <position position="191"/>
        </location>
        <ligand>
            <name>S-adenosyl-L-methionine</name>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="12">
        <location>
            <position position="194"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <position position="219"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <position position="220"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="12">
        <location>
            <position position="220"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="12">
        <location>
            <position position="249"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00012880"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022603"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022691"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022867"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022939"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023453"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PIRNR" id="PIRNR037177"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PIRSR" id="PIRSR037177-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PIRSR" id="PIRSR037177-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PIRSR" id="PIRSR037177-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029663.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="264" mass="30107" checksum="FA62B9CD8FC947A0" modified="2019-05-08" version="1">MKMLESYPILLSIVTSLLFIVLLLAVLIRNNGWAAILWNEIIMQKINNFIMAQSKEERILNFVLQNAVRGDPQSVVDTIDKYCSQKEWAMNVGAEKGLILDKILEETKPSVLLELGTYCGYSAVRIARLLKPSARLLTIEFNPDFAAIARQMIEFAGVQDKVKIVEGSSETIIPQLKKKHEVDTLDFVFLDHWKDRYLPDTILLLDQGLLRKGSVILADNVIFPGAPEFIKYIRNNNRFECTNYPSYVEYMKVKDAMEKAVFVG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INK5</accession>
    <name>A0A452INK5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Unc-13 homolog A</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452INK5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033478.1">
        <property type="protein sequence ID" value="ENSGAGP00000029482.1"/>
        <property type="gene ID" value="ENSGAGG00000020811.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031594">
        <property type="term" value="C:neuromuscular junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098831">
        <property type="term" value="C:presynaptic active zone cytoplasmic component"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042734">
        <property type="term" value="C:presynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030672">
        <property type="term" value="C:synaptic vesicle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043195">
        <property type="term" value="C:terminal bouton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019992">
        <property type="term" value="F:diacylglycerol binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005543">
        <property type="term" value="F:phospholipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017075">
        <property type="term" value="F:syntaxin-1 binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061789">
        <property type="term" value="P:dense core granule priming"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0099525">
        <property type="term" value="P:presynaptic dense core vesicle exocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035249">
        <property type="term" value="P:synaptic transmission, glutamatergic"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016081">
        <property type="term" value="P:synaptic vesicle docking"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016082">
        <property type="term" value="P:synaptic vesicle priming"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd20859">
        <property type="entry name" value="C1_Munc13-2-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd08394">
        <property type="entry name" value="C2A_Munc13"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04027">
        <property type="entry name" value="C2B_Munc13"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.357.50:FF:000001">
        <property type="entry name" value="Protein unc-13 homolog B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000002">
        <property type="entry name" value="Protein unc-13 homolog B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000031">
        <property type="entry name" value="Protein unc-13 homolog B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.60.20:FF:000001">
        <property type="entry name" value="Protein unc-13 homolog B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.357.50">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.1100">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.60.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046349">
        <property type="entry name" value="C1-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000008">
        <property type="entry name" value="C2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010439">
        <property type="entry name" value="MUN_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014770">
        <property type="entry name" value="Munc13_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002219">
        <property type="entry name" value="PKC_DAG/PE"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027080">
        <property type="entry name" value="Unc-13"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037302">
        <property type="entry name" value="Unc-13_C2B"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10480">
        <property type="entry name" value="PROTEIN UNC-13 HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10480:SF1">
        <property type="entry name" value="PROTEIN UNC-13 HOMOLOG A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00130">
        <property type="entry name" value="C1_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00168">
        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06292">
        <property type="entry name" value="MUN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00360">
        <property type="entry name" value="C2DOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00109">
        <property type="entry name" value="C1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00239">
        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM01145">
        <property type="entry name" value="DUF1041"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49562">
        <property type="entry name" value="C2 domain (Calcium/lipid-binding domain, CaLB)"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57889">
        <property type="entry name" value="Cysteine-rich domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50004">
        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51258">
        <property type="entry name" value="MHD1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00479">
        <property type="entry name" value="ZF_DAG_PE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50081">
        <property type="entry name" value="ZF_DAG_PE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0268">Exocytosis</keyword>
    <keyword evidence="2" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <keyword evidence="4" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2" evidence="7">
        <location>
            <begin position="1"/>
            <end position="97"/>
        </location>
    </feature>
    <feature type="domain" description="Phorbol-ester/DAG-type" evidence="8">
        <location>
            <begin position="450"/>
            <end position="500"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="7">
        <location>
            <begin position="556"/>
            <end position="680"/>
        </location>
    </feature>
    <feature type="domain" description="MHD1" evidence="9">
        <location>
            <begin position="964"/>
            <end position="1086"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="155"/>
            <end position="242"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="277"/>
            <end position="342"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="191"/>
            <end position="200"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="201"/>
            <end position="220"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="227"/>
            <end position="241"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="281"/>
            <end position="303"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="333"/>
            <end position="342"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50081"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51258"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029482.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="12">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1208" mass="136720" checksum="5CACDCAC46579128" modified="2019-05-08" version="1">TFYSLFAVKKAKFDGAQEKFNTYVTLKVQNVKSTTIAVRGNLPCWEQDFMFEINRLDLGLTVEVWNKGLIWDTMVGTVWIPLQTIRQSNEEGPGEWLTLDSQVIMTDNEISGTKDPTFHRILLDTHFELPLDIPEEEARYWAKKLEQLNAMKDQDDYEFQEEQDKPLPVPGSQCCKYRSPPASNDDPDSTMDDRDSDYRSETSNSIPPPYYTTSQPNASVHQYPMRHQQQSSRDSYMDSQTGTGGCTWVRTAPWHATSELQAVSGLPSCPCSTRLVLSDSPGRTPSAGQPPQGPTPSGVSPSASLFLGPNLQAFSPPGLRRELPSASLPGLHPQREHYREQEDVEVQDAKSRAKANWLKAFNKVCMQLQEGPVQRGLRRAHRVGGGAGCDSSPLIQLFGSVSLQQSLVQSRKAGITSALASSTLNNEELKNHVYKKTLQALIYPISSTTPHNFEVWTATTPTYCYECEGLLWGIARQGMRCTECGVKCHEKCQDLLNADCLQRAAEKSSKHGAEDRTQNIIMVLKDRMKIRERNKPEIFELIQEIFAVSKATHTQQMKAVKQSVLDGTSKWSAKISITVVCAQGLQAKDKTGSSDPYVTVQVGKTKKRTKTIYGNLNPVWEENFHFECHNSSDRIKVRVWDEDDDIKSRVKQRFKRESDDFLGQTIIEVRTLSGEMDVWYNLDKRTDKSAVSGAIRLHISVEIKGEEKVAPYHVQYTCLHENLFHFVTDIQNNGVVKIPEAKGDDAWKVYFDETAQEIVDEFAMRYGVESIYQAMTHFACLSSKYMCPGVPAVMSTLLANINAYYAHTTASTNVSASDRFAASNFGKERFVKLLDQLHNSLRIDLSMYRNNFPASSPERLQDLKSTVDLLTSITFFRMKVQELQSPPRASQVVKDCVKACLNSTYEYIFNNCHELYSREYQTDPTKKGEVPPEEQGPSIKNLDFWSKLITLIVSIIEEDKNSYTPCLNHADYMNLHFKVKWLYNEYVTELPAFKSRVPEYPWLDENEEVSRDFLHGALERDKKDGFQQTSEHALFSCSVVDVFSQLNQSFEIIKKLECPDPQIVGHYMRRFAKTISNVLLQYAEIISKDFALYCSKEKEKVPCILMNNIQQLRVQLEKMFEAMGGKELDAEASDILKELQVKLNNVLDELSRVFATSFQPHIEECVKQMGDILSQVKGTGNVPASACSSVAQDADNVLQPIMDLLDSK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INK8</accession>
    <name>A0A452INK8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">E3 ubiquitin-protein ligase NEDD4-like</fullName>
            <ecNumber evidence="6">2.3.2.26</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="17">HECT-type E3 ubiquitin transferase NED4L</fullName>
        </alternativeName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>S-ubiquitinyl-[E2 ubiquitin-conjugating enzyme]-L-cysteine + [acceptor protein]-L-lysine = [E2 ubiquitin-conjugating enzyme]-L-cysteine + N(6)-ubiquitinyl-[acceptor protein]-L-lysine.</text>
            <dbReference type="EC" id="2.3.2.26"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="5">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Endosome</location>
            <location evidence="4">Multivesicular body</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Golgi apparatus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="2.3.2.26" evidence="6"/>
    <dbReference type="AlphaFoldDB" id="A0A452INK8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033550.1">
        <property type="protein sequence ID" value="ENSGAGP00000029545.1"/>
        <property type="gene ID" value="ENSGAGG00000021328.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00143"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005771">
        <property type="term" value="C:multivesicular body"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019871">
        <property type="term" value="F:sodium channel inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061630">
        <property type="term" value="F:ubiquitin protein ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006811">
        <property type="term" value="P:monoatomic ion transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010766">
        <property type="term" value="P:negative regulation of sodium ion transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016567">
        <property type="term" value="P:protein ubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048814">
        <property type="term" value="P:regulation of dendrite morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006511">
        <property type="term" value="P:ubiquitin-dependent protein catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd00078">
        <property type="entry name" value="HECTc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00201">
        <property type="entry name" value="WW"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.70.10:FF:000017">
        <property type="entry name" value="E3 ubiquitin-protein ligase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1750.10:FF:000026">
        <property type="entry name" value="E3 ubiquitin-protein ligase HACE1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.2160.10:FF:000001">
        <property type="entry name" value="E3 ubiquitin-protein ligase NEDD4-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.2410.10:FF:000001">
        <property type="entry name" value="E3 ubiquitin-protein ligase NEDD4-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1750.10:FF:000001">
        <property type="entry name" value="E3 ubiquitin-protein ligase NEDD4-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.70.10:FF:000006">
        <property type="entry name" value="E3 ubiquitin-protein ligase NEDD4-like protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.70.10:FF:000008">
        <property type="entry name" value="E3 ubiquitin-protein ligase NEDD4-like protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.70.10:FF:000025">
        <property type="entry name" value="E3 ubiquitin-protein ligase NEDD4-like protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.20.70.10">
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.2160.10">
        <property type="entry name" value="Hect, E3 ligase catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.2410.10">
        <property type="entry name" value="Hect, E3 ligase catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1750.10">
        <property type="entry name" value="Hect, E3 ligase catalytic domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050409">
        <property type="entry name" value="E3_ubiq-protein_ligase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000569">
        <property type="entry name" value="HECT_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035983">
        <property type="entry name" value="Hect_E3_ubiquitin_ligase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001202">
        <property type="entry name" value="WW_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036020">
        <property type="entry name" value="WW_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11254:SF440">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE NEDD-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11254">
        <property type="entry name" value="HECT DOMAIN UBIQUITIN-PROTEIN LIGASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00632">
        <property type="entry name" value="HECT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00397">
        <property type="entry name" value="WW"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00119">
        <property type="entry name" value="HECTc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00456">
        <property type="entry name" value="WW"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56204">
        <property type="entry name" value="Hect, E3 ligase catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51045">
        <property type="entry name" value="WW domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50237">
        <property type="entry name" value="HECT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01159">
        <property type="entry name" value="WW_DOMAIN_1"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50020">
        <property type="entry name" value="WW_DOMAIN_2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="12" id="KW-0221">Differentiation</keyword>
    <keyword evidence="11" id="KW-0967">Endosome</keyword>
    <keyword evidence="15" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="8" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0677">Repeat</keyword>
    <keyword evidence="9" id="KW-0808">Transferase</keyword>
    <keyword evidence="14" id="KW-0832">Ubl conjugation</keyword>
    <keyword evidence="13 18" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="WW" evidence="20">
        <location>
            <begin position="483"/>
            <end position="516"/>
        </location>
    </feature>
    <feature type="domain" description="WW" evidence="20">
        <location>
            <begin position="675"/>
            <end position="708"/>
        </location>
    </feature>
    <feature type="domain" description="WW" evidence="20">
        <location>
            <begin position="787"/>
            <end position="820"/>
        </location>
    </feature>
    <feature type="domain" description="WW" evidence="20">
        <location>
            <begin position="838"/>
            <end position="871"/>
        </location>
    </feature>
    <feature type="domain" description="HECT" evidence="21">
        <location>
            <begin position="930"/>
            <end position="1264"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="130"/>
            <end position="159"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="177"/>
            <end position="199"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="312"/>
            <end position="335"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="362"/>
            <end position="390"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="445"/>
            <end position="491"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="534"/>
            <end position="668"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="716"/>
            <end position="785"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="19">
        <location>
            <begin position="133"/>
            <end position="148"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="318"/>
            <end position="334"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="379"/>
            <end position="389"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="454"/>
            <end position="465"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="19">
        <location>
            <begin position="571"/>
            <end position="580"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="641"/>
            <end position="659"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="750"/>
            <end position="761"/>
        </location>
    </feature>
    <feature type="active site" description="Glycyl thioester intermediate" evidence="18">
        <location>
            <position position="1232"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000885"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004555"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004559"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00012485"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00068697"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00082247"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00104"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50020"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50237"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029545.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1265" mass="143913" checksum="C24A5B7375D57A46" modified="2019-05-08" version="1">MAHRLRFHFGSGRSNTAPESEILDQEREDDFFMAFHTLPRRNSPHPFSQHGADYGGGDLQEVGSLKRSTSMFIPQLLNNIDVRPTRSSSMQISLQRKAVNGCADECGAPESPEETLSCKFSDFREHYTCPVDSHSSSPSSPQVTSENSPPRMPEETGQRLNGAVSCNHRIFCTIPSNNQNDDVATTAEEDEASEAASGLNISGVRIQHRASSADMHQVMLSPFGSEVQDNALASEPRRWSLQHLPDTSANSGKKLFVFQLQQSQASGTGSDYNFGFTGTKGNRLVRYPRIRLERSTSYPVQPRPEQISLVEDEGNSELHGNSRNGSEISRSNSVERIPQGQECLFKIRPDQNTRQQHFRILVTRGTEEQNQVSDEDSPDTSGPTVTSATTRDDFLGQVDVPLNHLPTEDPTMERPYTFKDFLLRPRSHKSRVKGFLRLKMAYMPKNGGQEEETTEQREESERAWEVVDSNDSVSHHQEELPPPPLPPGWEEKVDNLGRTYYVNHNNRTTQWHRPSLIDVASDSDNNIRQINQEAAHRRFRSRRHISEDLEPEPSESGDASEPWETISEEASISGDSLSLSLPPPPASPVSRTSPQELSDELNRRLQVTPDSNGEQFGSLIQRDPSSRLRSCSVTDAVAEQSHLSLQSGPSRRARSSTVTGGEEPTPSVAYVHTTPGLPSGWEERKDAKGRTYYVNHNNRTTTWTRPIMQLAEDGAVGSAANSNNHLSEPQIRRPRSLSSPTVTLSAPLEFAKDSPVRRAVKDTLSNPQSPQPSPYNSPKPQHKVTQSFLPPGWEMRIAPNGRPFFIDHNTKTTTWEDPRLKFPVHLRSKASLNPNDLGPLPPGWEERIHLDGRTFYIDHNSKITQWEDPRLQNPAITGPAVPYSREFKQKYDYFRKKLKKPADIPNRFEMKLHRNNIFEESYRRIMSVKRSDVLKARLWIEFESEKGLDYGGVAREWFFLLSKEMFNPYYGLFEYSATDNYTLQINPNSGLCNEDHLSYFTFIGRVAGLAVFHGKLLDGFFIRPFYKMMLGKQITLKDMESVDSEYYNSLKWILENDPTELDLMFCIDEENFGQTYQVDLKPNGSEIMVTNENKREYIDLVIQWRFVNRVQKQMNAFLEGFTELLPIDLIKIFDENELELLMCGLGDVDVNDWRQHTIYKNGYCPNHPVIQWYWKAVLLMDAEKRIRLLQFVTGTSRVPMNGFAELYGSNGPQLFTIEQWGSPEKLPRAHTCFNRLDLPPYESFEDLREKLLMAVENAQGFEGVD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INL2</accession>
    <name>A0A452INL2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Interleukin 1 receptor type 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the interleukin-1 receptor family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452INL2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033695.1">
        <property type="protein sequence ID" value="ENSGAGP00000029674.1"/>
        <property type="gene ID" value="ENSGAGG00000021421.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016787">
        <property type="term" value="F:hydrolase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004908">
        <property type="term" value="F:interleukin-1 receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050727">
        <property type="term" value="P:regulation of inflammatory response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10140:FF:000002">
        <property type="entry name" value="Interleukin 1 receptor accessory protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:001302">
        <property type="entry name" value="Interleukin 1 receptor like 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000188">
        <property type="entry name" value="Interleukin-1 receptor accessory protein-like 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.10140">
        <property type="entry name" value="Toll/interleukin-1 receptor homology (TIR) domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015621">
        <property type="entry name" value="IL-1_rcpt_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004074">
        <property type="entry name" value="IL-1_rcpt_I/II-typ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000157">
        <property type="entry name" value="TIR_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035897">
        <property type="entry name" value="Toll_tir_struct_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11890">
        <property type="entry name" value="INTERLEUKIN-1 RECEPTOR FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11890:SF26">
        <property type="entry name" value="INTERLEUKIN-1 RECEPTOR TYPE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01582">
        <property type="entry name" value="TIR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01536">
        <property type="entry name" value="INTRLKN1R12F"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01537">
        <property type="entry name" value="INTRLKN1R1F"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00255">
        <property type="entry name" value="TIR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52200">
        <property type="entry name" value="Toll/Interleukin receptor TIR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50104">
        <property type="entry name" value="TIR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="13" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="9 14" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-0520">NAD</keyword>
    <keyword evidence="11" id="KW-0675">Receptor</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="4 15" id="KW-0732">Signal</keyword>
    <keyword evidence="3 14" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 14" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="15">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019178521" description="Interleukin 1 receptor type 1" evidence="15">
        <location>
            <begin position="21"/>
            <end position="542"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="313"/>
            <end position="338"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="17">
        <location>
            <begin position="40"/>
            <end position="119"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="17">
        <location>
            <begin position="147"/>
            <end position="214"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="17">
        <location>
            <begin position="223"/>
            <end position="305"/>
        </location>
    </feature>
    <feature type="domain" description="TIR" evidence="16">
        <location>
            <begin position="362"/>
            <end position="517"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009752"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023027"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50104"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029674.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="20">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="542" mass="62266" checksum="CCA39EA633896DFD" modified="2019-05-08" version="1">MAAAVFTYSVILVFLSSVKAEECQVHDVLHSQNLISVLVGEPLTIDCLLHKILALQHMNYSLTWYKSGKKMPVTQVKLSRIHQHENLLWFMPAILEDSGSYECVMWNLTSCNKIYFSITVFKNTAGLCFNGDFLYAQEISASSNEKIVCPHLDYFRDEKNTLPIHWYKECNLIDNERFLSWDDDLIINSANVNDSGNYSCKMTYTYMEKEYNISRNIYVTVIGSNVIVDCNVSSSKDNSVGISWRVNNTLVNFLFMGRVQEGNQEDSFPDGYLFSTVTLNITEVKQEDYGRQFVCHAGEVAAYIALRQPVRNFLGYLIGGLIAPVFVIIVAILIYKYFKVDIVLWYRKSCRPFLSKEVSDGKIYDAYVLYPKINKVDCIYTPDNFVLKLLPQVLERQCGYNLFILGRDDLPGKAVVNVVDETIKQSRRLIIILVPDSSSCSLLEDIFEQQLAVYNALVRDGIKVILIELDKIKDYANMPESIKYIKQKHGAIRWKGDFTEKSHSANTKFWKNVRYRMPPRCPTFSELHLLPTALNTFQTTER</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INL5</accession>
    <name>A0A452INL5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Neurabin-1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="14">Neurabin-I</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="13">Neural tissue-specific F-actin-binding protein I</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="15">Protein phosphatase 1 regulatory subunit 9A</fullName>
        </alternativeName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="11">Synapse</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000033458.1">
        <property type="protein sequence ID" value="ENSGAGP00000029464.1"/>
        <property type="gene ID" value="ENSGAGG00000020904.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015629">
        <property type="term" value="C:actin cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030425">
        <property type="term" value="C:dendrite"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0014069">
        <property type="term" value="C:postsynaptic density"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007015">
        <property type="term" value="P:actin filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019722">
        <property type="term" value="P:calcium-mediated signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031175">
        <property type="term" value="P:neuron projection development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd06790">
        <property type="entry name" value="PDZ_neurabin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd09512">
        <property type="entry name" value="SAM_Neurabin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000010">
        <property type="entry name" value="Neurabin-1 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.50:FF:000008">
        <property type="entry name" value="Neurabin-1 isoform 1-like protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.42.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.50">
        <property type="entry name" value="Transcription Factor, Ets-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040645">
        <property type="entry name" value="Neurabin-1/2_PDZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043446">
        <property type="entry name" value="Neurabin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001478">
        <property type="entry name" value="PDZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036034">
        <property type="entry name" value="PDZ_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001660">
        <property type="entry name" value="SAM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013761">
        <property type="entry name" value="SAM/pointed_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16154">
        <property type="entry name" value="NEURABIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16154:SF22">
        <property type="entry name" value="NEURABIN-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00595">
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        <property type="entry name" value="PDZ_5"/>
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        <property type="entry name" value="SAM_2"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="PDZ"/>
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    </dbReference>
    <dbReference type="SUPFAM" id="SSF50156">
        <property type="entry name" value="PDZ domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="SAM/Pointed domain"/>
        <property type="match status" value="1"/>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50105">
        <property type="entry name" value="SAM_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="9" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="8 16" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="10" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="5" id="KW-0221">Differentiation</keyword>
    <keyword evidence="6" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0770">Synapse</keyword>
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        <location>
            <begin position="507"/>
            <end position="595"/>
        </location>
    </feature>
    <feature type="domain" description="SAM" evidence="18">
        <location>
            <begin position="1092"/>
            <end position="1155"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
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        </location>
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    <feature type="region of interest" description="Disordered" evidence="17">
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        </location>
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        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="890"/>
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        </location>
    </feature>
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            <begin position="1158"/>
            <end position="1202"/>
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            <begin position="603"/>
            <end position="630"/>
        </location>
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        <location>
            <begin position="683"/>
            <end position="826"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="17">
        <location>
            <begin position="1"/>
            <end position="12"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="17">
        <location>
            <begin position="33"/>
            <end position="49"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="17">
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            <begin position="182"/>
            <end position="195"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="17">
        <location>
            <begin position="926"/>
            <end position="939"/>
        </location>
    </feature>
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        <location>
            <begin position="975"/>
            <end position="994"/>
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    </feature>
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        <location>
            <begin position="1158"/>
            <end position="1192"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
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            <dbReference type="ARBA" id="ARBA00022553"/>
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            <dbReference type="ARBA" id="ARBA00023018"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00077125"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00082439"/>
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    </evidence>
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    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
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    </evidence>
    <evidence type="ECO:0000259" key="18">
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    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50106"/>
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    </evidence>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029464.1"/>
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    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
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    </evidence>
    <sequence length="1202" mass="134573" checksum="D60E72FA8453D1D6" modified="2019-05-08" version="1">MMKTESSGERSTLRSASPHRNAYRTEFQALKSTFDKPKSDGDQKTKDEGEVSQQSRGRKYGSNVNRIKNLFMQMGMEPSESTGVTPKTRGKSGPLSPQRIRPKEFVEKADGSVVKLESSVSERISRFDTVHGGPSYSKFTETRKMFERNVHEMGHSNRYSPKKERVVSNELQDEWCSSKSNRGSTDSLDSLSSRTETISPTVSQLSAVFENTDSHTVIVLEKSENNEEYSVTGHYPLNLSSTVENLSSTVANLDGFSPLKDASAWSSPAKQTTSSVSAETFQQNSTSSTIREASNSTLPVSKTNEEIRKNKEASFDPVDVCAATKQELASINSEKMPXHSQNAESACVGTGAADIPRNLVAGGDFAADVASDATESPYDDASGKEVQKDLNNFQSSHVYMHSDYNVYRVRSRYNSDWGETGTEQDDQDDSDENNYYEPDMEYAEVIGLPAEDEIPANRRIKFSSAPIKVFNTYSNEDYDRRNDEVDPVAASAEYELEKRVEKLELFPVELGKDEDGLGISIIGMGVGADAGLEKLGIFVKTVTEGGAAERDGRIQVNDQIVEVDGISLVGVTQNFAATVLRNTKGNVRFVIGREKPGQVSEVAQLISQTLEQERRQRELLEQHYAQYDADDDETGEYATDEEDEDMGPIVPGADMAIEVFDLPENDDMFSPTELDTTKLAHKFKELQIKHAVTEAEIQKLKTKLQAAENEKVRWEREKTQLQQNIEENKERMMKLESYWIEAQTLCHTVNEHLKETQSQYQALEKKYNKAKKLIKDFQQKELDFIKRQEAERKKIEELEKAHLAEVQGLQARIRDLEAEVFKLLKQNGSQVNNNNNIFERQTSFGEVSRGDPVESLDVEQVSCLDDLSQDFNEAVPETERLDSKALKTRAQLSVRNKRQRPTRTRLYDSISSTDGEDSLERKVRRASSPASDSGASSLSPVASSAAFSFDNVAENQEGGQQHKGGVLSHYARGDTPLSSPSKSSHSSETSPLHHPTNRNVLQDKAWKEKASLLQDVCIRSYVDFDPSFCLQNFTFTDDFSPSSTSSADLSGLGAEPKPPGFSHSLALSSDEILDDGQSPKHSQCQNRAVQEWSIQQVSHWLMSLNFEQYVSEFTAQNINGEHLLQLDGSKLKTLGMTSSQDRAIIKKKLKEMKVSIERARKAQEKMEKQREKLRKKEQERLQRKSKKLDKSSSDATEGANEQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INM0</accession>
    <name>A0A452INM0_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452INM0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033493.1">
        <property type="protein sequence ID" value="ENSGAGP00000029497.1"/>
        <property type="gene ID" value="ENSGAGG00000021299.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="term" value="C:mRNA cleavage and polyadenylation specificity factor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="P:mRNA 3'-end processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
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        <property type="entry name" value="WD repeat domain 33"/>
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        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Pfs2-like"/>
    </dbReference>
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        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
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    </dbReference>
    <dbReference type="PANTHER" id="PTHR22836:SF0">
        <property type="entry name" value="PRE-MRNA 3' END PROCESSING PROTEIN WDR33"/>
        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
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        <property type="entry name" value="WD_REPEATS_REGION"/>
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    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
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        <location>
            <begin position="157"/>
            <end position="189"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="1">
        <location>
            <begin position="198"/>
            <end position="230"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029497.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="260" mass="30476" checksum="9742B3A87F461FA1" modified="2019-05-08" version="1">MATEIGSPPRFFHMPRFQHQAPRQLFYKRPDFAQQQAMQQLTFDGKRMRKAVNRKTIDYNPSVIKYLENRVWQRDQRDMRAIQPDAGYYNDLVPPIGMLNNPMNAVTTKFVRTSTNKVKCPVFVVRWTPEGRRLVTGASSGEFTLWNGLTFNFETILQAHDSPVRAMTWSHNDMWMLTADHGGYVKYWQSNMNNVKMFQAHKEAIREASFSPTDNKFATCSDDGTVRIWDFLRCHEERILREIFLFLLLATHTRETQLGD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INM1</accession>
    <name>A0A452INM1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">PDZ domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="11">Synapse</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000033463.1">
        <property type="protein sequence ID" value="ENSGAGP00000029469.1"/>
        <property type="gene ID" value="ENSGAGG00000020904.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015629">
        <property type="term" value="C:actin cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030425">
        <property type="term" value="C:dendrite"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0014069">
        <property type="term" value="C:postsynaptic density"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007015">
        <property type="term" value="P:actin filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019722">
        <property type="term" value="P:calcium-mediated signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031175">
        <property type="term" value="P:neuron projection development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd06790">
        <property type="entry name" value="PDZ_neurabin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000010">
        <property type="entry name" value="Neurabin-1 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.42.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040645">
        <property type="entry name" value="Neurabin-1/2_PDZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043446">
        <property type="entry name" value="Neurabin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001478">
        <property type="entry name" value="PDZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036034">
        <property type="entry name" value="PDZ_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16154">
        <property type="entry name" value="NEURABIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16154:SF22">
        <property type="entry name" value="NEURABIN-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00595">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
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    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57997">
        <property type="entry name" value="Tropomyosin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50106">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="9" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="8 12" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="10" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="5" id="KW-0221">Differentiation</keyword>
    <keyword evidence="6" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0770">Synapse</keyword>
    <feature type="domain" description="PDZ" evidence="14">
        <location>
            <begin position="504"/>
            <end position="592"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="1"/>
            <end position="104"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="176"/>
            <end position="196"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="275"/>
            <end position="302"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="880"/>
            <end position="928"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="970"/>
            <end position="1013"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="12">
        <location>
            <begin position="600"/>
            <end position="627"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="12">
        <location>
            <begin position="680"/>
            <end position="823"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="13">
        <location>
            <begin position="1"/>
            <end position="12"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="13">
        <location>
            <begin position="33"/>
            <end position="49"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="13">
        <location>
            <begin position="182"/>
            <end position="195"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="13">
        <location>
            <begin position="990"/>
            <end position="1009"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00034103"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50106"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029469.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1058" mass="118198" checksum="05B3FF0D450D3184" modified="2019-05-08" version="1">MMKTESSGERSTLRSASPHRNAYRTEFQALKSTFDKPKSDGDQKTKDEGEVSQQSRGRKYGSNVNRIKNLFMQMGMEPSESTGVTPKTRGKSGPLSPQRIRPKEFVEKADGSVVKLESSVSERISRFDTVHGGPSYSKFTETRKMFERNVHEMGHSNRYSPKKERVVSNELQDEWCSSKSNRGSTDSLDSLSSRTETISPTVSQLSAVFENTDSHTVIVLEKSENNEEYSVTGHYPLNLSSTVENLSSTVANLDGFSPLKDASAWSSPAKQTTSSVSAETFQQNSTSSTIREASNSTLPVSKTNEEIRKNKEASFDPVDVCAATKQELASINMPXHSQNAESACVGTGAADIPRNLVAGGDFAADVASDATESPYDDASGKEVQKDLNNFQSSHVYMHSDYNVYRVRSRYNSDWGETGTEQDDQDDSDENNYYEPDMEYAEVIGLPAEDEIPANRRIKFSSAPIKVFNTYSNEDYDRRNDEVDPVAASAEYELEKRVEKLELFPVELGKDEDGLGISIIGMGVGADAGLEKLGIFVKTVTEGGAAERDGRIQVNDQIVEVDGISLVGVTQNFAATVLRNTKGNVRFVIGREKPGQVSEVAQLISQTLEQERRQRELLEQHYAQYDADDDETGEYATDEEDEDMGPIVPGADMAIEVFDLPENDDMFSPTELDTTKLAHKFKELQIKHAVTEAEIQKLKTKLQAAENEKVRWEREKTQLQQNIEENKERMMKLESYWIEAQTLCHTVNEHLKETQSQYQALEKKYNKAKKLIKDFQQKELDFIKRQEAERKKIEELEKAHLAEVQGLQARIRDLEAEVFKLLKQNGSQVNNNNNIFERQTSFGEVSRGDPVESLDVEQVSCLDDLSQDFNEAVPETERLDSKALKTRAQLSVRNKRQRPTRTRLYDSISSTDGEDSLERKPSDSYSSPMHISKSLLPICVRASSPASDSGASSLSPVASSAAFSFDNVAENQEGGQQHKGGVLSHYARGDTPLSSPSKSSHSSETSPLHHPTNRNVLQDKVGLKQKEVFLHTTHSQPVELVARQCCEGQKYNWVQKSVR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INM2</accession>
    <name>A0A452INM2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">non-specific serine/threonine protein kinase</fullName>
            <ecNumber evidence="2">2.7.11.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the protein kinase superfamily. AGC Ser/Thr protein kinase family. RAC subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452INM2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033457.1">
        <property type="protein sequence ID" value="ENSGAGP00000029463.1"/>
        <property type="gene ID" value="ENSGAGG00000021286.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd01241">
        <property type="entry name" value="PH_PKB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05593">
        <property type="entry name" value="STKc_PKB_gamma"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000033">
        <property type="entry name" value="Non-specific serine/threonine protein kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000027">
        <property type="entry name" value="Non-specific serine/threonine protein kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000838">
        <property type="entry name" value="Non-specific serine/threonine protein kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000961">
        <property type="entry name" value="AGC-kinase_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034675">
        <property type="entry name" value="Akt3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039026">
        <property type="entry name" value="PH_PKB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017892">
        <property type="entry name" value="Pkinase_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24351">
        <property type="entry name" value="RIBOSOMAL PROTEIN S6 KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00169">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00433">
        <property type="entry name" value="Pkinase_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00233">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00133">
        <property type="entry name" value="S_TK_X"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51285">
        <property type="entry name" value="AGC_KINASE_CTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 9" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="7" id="KW-0418">Kinase</keyword>
    <keyword evidence="6 9" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="5" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="PH" evidence="11">
        <location>
            <begin position="30"/>
            <end position="132"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="12">
        <location>
            <begin position="173"/>
            <end position="430"/>
        </location>
    </feature>
    <feature type="domain" description="AGC-kinase C-terminal" evidence="13">
        <location>
            <begin position="431"/>
            <end position="504"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="483"/>
            <end position="504"/>
        </location>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="212"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006935"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00012513"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS51285"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029463.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="504" mass="58547" checksum="5B96C683AD2CA610" modified="2019-05-08" version="1">MKKIVQANSDTVNVGHGYSTVTSHIKVFFFFSINAYFSHIGEYIKNWRPRYFLLKTDGSFIGYKEKPQDVDLPYPLNNFSVAKCQLMKTERPKPNTFIIRCLQWTTVIERTFHVDTPEEREEWTEAIQAVADRLQRQEEERMNCSPTSQIDNIGEEEMDASTTHHKRKTMNDFDYLKLLGKGTFGKVILVREKASGKYYAMKILKKEVIIAKDEVAHTLTESRVLKNTRHPFLTSLKYSFQTKDRLCFVMEYVNGGELFFHLSRERVFSEDRTRFYGAEIVSALDYLHSGKIVYRDLKLENLMLDKDGHIKITDFGLCKEGITDAATMKTFCGTPEYLAPEVLEDNDYGRAVDWWGLGVVMYEMMCGRLPFYNQDHEKLFELILMEDIKFPRTLSSDAKSLLSGLLIKDPNKRLGGGPDDAKEIMRHSFFVGVNWQDVYDKKLVPPFKPQVTSETDTRYFDEEFTAQTITITPPEKYDEDGMDCMDNERRPHFPQFSYSASGRE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INM3</accession>
    <name>A0A452INM3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="22">Cell division cycle 14B</fullName>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>O-phospho-L-seryl-[protein] + H2O = L-seryl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:20629"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="EC" id="3.1.3.16"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="16">
            <text>O-phospho-L-threonyl-[protein] + H2O = L-threonyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:47004"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="EC" id="3.1.3.16"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="14">Cell projection</location>
            <location evidence="14">Kinocilium</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
            <location evidence="2">Microtubule organizing center</location>
            <location evidence="2">Centrosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Cytoskeleton</location>
            <location evidence="3">Spindle pole</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the protein-tyrosine phosphatase family. Non-receptor class CDC14 subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452INM3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033568.1">
        <property type="protein sequence ID" value="ENSGAGP00000029560.1"/>
        <property type="gene ID" value="ENSGAGG00000021351.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000033569.1">
        <property type="protein sequence ID" value="ENSGAGP00000029561.1"/>
        <property type="gene ID" value="ENSGAGG00000021351.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060091">
        <property type="term" value="C:kinocilium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000922">
        <property type="term" value="C:spindle pole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004722">
        <property type="term" value="F:protein serine/threonine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007605">
        <property type="term" value="P:sensory perception of sound"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd14499">
        <property type="entry name" value="CDC14_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd17657">
        <property type="entry name" value="CDC14_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000032">
        <property type="entry name" value="dual specificity protein phosphatase CDC14A isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000006">
        <property type="entry name" value="Dual specificity protein phosphatase CDC14B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044506">
        <property type="entry name" value="CDC14_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029260">
        <property type="entry name" value="DSPn"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000340">
        <property type="entry name" value="Dual-sp_phosphatase_cat-dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050561">
        <property type="entry name" value="PTP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016130">
        <property type="entry name" value="Tyr_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020422">
        <property type="entry name" value="TYR_PHOSPHATASE_DUAL_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23339">
        <property type="entry name" value="TYROSINE SPECIFIC PROTEIN PHOSPHATASE AND DUAL SPECIFICITY PROTEIN PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00782">
        <property type="entry name" value="DSPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14671">
        <property type="entry name" value="DSPn"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00195">
        <property type="entry name" value="DSPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00383">
        <property type="entry name" value="TYR_PHOSPHATASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50054">
        <property type="entry name" value="TYR_PHOSPHATASE_DUAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="13" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="7" id="KW-0132">Cell division</keyword>
    <keyword evidence="12" id="KW-0966">Cell projection</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="10" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="9" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="18">
        <location>
            <begin position="212"/>
            <end position="371"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="19">
        <location>
            <begin position="294"/>
            <end position="356"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1"/>
            <end position="44"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="17">
        <location>
            <begin position="1"/>
            <end position="13"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004647"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00007315"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00037822"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00047761"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00048336"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029560.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="22">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="478" mass="54381" checksum="FB4F65F0159C8A2B" modified="2019-05-08" version="1">MKRKNWAGARRHSAPAPPAVKKTRSAAAAAAGVEEEPPPSGPESCMEITGRLYFAILCQKPKSGVANTHYFCIDDELVYENFYADFGPLNLAMVYRYCCKLNKKLKSFTLIRKKIIHYTGFDQKKQANAAFLIGSYAIIYLKESPEDVYRLLLAGNASYLPFRDASFGTCSFHLTLLDCFHAINKALRYGFLDFSAFDVNEYEHYEKAENGDFNWIIPNKFIAFSGPHSRSKTENGYPHHAPEAYFPYFRRHKVTTIIRLNRKMYDAKRFADAGFEHYDLFFADGSIPSDTIVKTFLNICENAEGVIAVHCKAGLGRTGTLIACYIMKHYRMTAAETIAWIRINRPGSVIGPQQHFLVEKQTELWREGDVFRAKLKENRKIAVTKILTGVDDISINDAKNRRANRKDTELYSDEDDANCVTQGDKLRALKSKRQAKAAATVPLTLIPQKEKRKRNALQQMRLMKLCGHSIPRTRTVLR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INM7</accession>
    <name>A0A452INM7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">DUF4455 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452INM7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033462.1">
        <property type="protein sequence ID" value="ENSGAGP00000029468.1"/>
        <property type="gene ID" value="ENSGAGG00000021287.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028089">
        <property type="entry name" value="DUF4455"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21444">
        <property type="entry name" value="COILED-COIL DOMAIN-CONTAINING PROTEIN 180"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21444:SF14">
        <property type="entry name" value="COILED-COIL DOMAIN-CONTAINING PROTEIN 180"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14643">
        <property type="entry name" value="DUF4455"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="DUF4455" evidence="2">
        <location>
            <begin position="129"/>
            <end position="594"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="620"/>
            <end position="706"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="635"/>
            <end position="649"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="1">
        <location>
            <begin position="660"/>
            <end position="671"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="1">
        <location>
            <begin position="684"/>
            <end position="696"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF14643"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029468.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1136" mass="132058" checksum="89197A2DBF5E5812" modified="2019-05-08" version="1">MVLVGAVRVVPSGKVYRQIFDDEVHLVRSLEEVRNKAEQHKFPMNSGRIPVVRDLGMCAGKGLLSHRQQKWVEGMPNDDFTENPVLYREMTFRKAQESEDVIAAREVRGLSDFIIPEKTNSNIIERLSERRRGRHNEAVAVLEQELACIGREMEPFILESGELLLTKLSESDKAIELLFKKIESDTDLEAFSFTSFGELWDAISQESLYRRQRIKEMDETLIKLEVARAEKIKDLLRKYTKILKDISYFSPSDVHRFIHTEAMMINQALLANQRAIAKLFVNLMEADLKRELSQWLKWQERLKDWKIMQKDYVVRSFREFMARKEVQEPTAVKRDLENMIKDQISLSRKRMELLQVLCDLLPPTHSKAEINEWYESLVALNKYIDTHNVQHMMIIRIQYERVCQECLANVQLCKNKLLSLKVCTEKEAEKVVNPDFFKMVGRLQSRFEQELEQMDRDFEDLAKHIEQDCKDLYKYFQQAIVLWDEHQLKLSQQENELQVKLNECRRKHENLNQIREANLDIILDKLRTQSSERKLKNLLGKAFASLDVIRAGYEAFHQDLVKIVASYPEAILKELISYSSSIGQYFGVKEVYKQVGSAFEEQVFVEDSEVQNMAETTGDSLLQESEETEQQSESFPQGSTEAEQQTESLTNKTEETNTVENEESSVQETEEAEKPGDGESTAPESEETEHLEDSAESEANRFENGDSPAQAYEEMDYLKDILPESEVLFHDSGEAESCEVIAEVFTTSSGNTYSVLGHKETGKTKTPEIYYAEFAEKTLPVYLENVLVPDSLFTELKKRIRLCFFEHLEKWFIQSLSNSWIIVTAKKEELDSELQLRLHMHQPRRERIEKDIHNVRAAELLLHQERLECHCVGVVEALNKEKNEFMKLQEDQNDISKNFRRRIQDMEAIFLSSTRAEKLVALSNSLHSELLNHVEVIQISLRSYRNYLEEALGKLRDSNVDFLKACRLFSEGGNFSPEEVEFFCKRLEKETGRIDFIEGLIMIDMEKMESNYLEQATEVINKFETKFHNLAMDRLFMEKIQRFLTNLQVKIKSEVANSNLQAQTLNSSLEKFLKKIDACAHPTVDKEAVTSEELYDFAKTVMKELKKRSKYLNCLLLQKGEQSIGNVFKGMSTLRN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INP1</accession>
    <name>A0A452INP1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Regulator of G-protein signaling 12</fullName>
        </recommendedName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">Regulates G protein-coupled receptor signaling cascades. Inhibits signal transduction by increasing the GTPase activity of G protein alpha subunits, thereby driving them into their inactive GDP-bound form.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell projection</location>
            <location evidence="2">Dendrite</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="14">Synapse</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000033592.1">
        <property type="protein sequence ID" value="ENSGAGP00000029582.1"/>
        <property type="gene ID" value="ENSGAGG00000021288.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030425">
        <property type="term" value="C:dendrite"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045202">
        <property type="term" value="C:synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008277">
        <property type="term" value="P:regulation of G protein-coupled receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd06710">
        <property type="entry name" value="PDZ_RGS12-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd13162">
        <property type="entry name" value="PTB_RGS12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd17136">
        <property type="entry name" value="RBD1_RGS12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd17138">
        <property type="entry name" value="RBD2_RGS12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd08742">
        <property type="entry name" value="RGS_RGS12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.167.10:FF:000001">
        <property type="entry name" value="Putative regulator of g-protein signaling 12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000296">
        <property type="entry name" value="Regulator of G protein signaling 12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000115">
        <property type="entry name" value="Regulator of G-protein signaling 12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.196.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.42.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.20.90">
        <property type="entry name" value="Phosphatidylinositol 3-kinase Catalytic Subunit, Chain A, domain 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.167.10">
        <property type="entry name" value="Regulator of G-protein Signalling 4, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003109">
        <property type="entry name" value="GoLoco_motif"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001478">
        <property type="entry name" value="PDZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036034">
        <property type="entry name" value="PDZ_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006020">
        <property type="entry name" value="PTB/PI_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003116">
        <property type="entry name" value="RBD_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016137">
        <property type="entry name" value="RGS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046995">
        <property type="entry name" value="RGS10/12/14-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037880">
        <property type="entry name" value="RGS12_RGS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036305">
        <property type="entry name" value="RGS_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024066">
        <property type="entry name" value="RGS_subdom1/3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044926">
        <property type="entry name" value="RGS_subdomain_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029071">
        <property type="entry name" value="Ubiquitin-like_domsf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45945:SF1">
        <property type="entry name" value="REGULATOR OF G-PROTEIN SIGNALING 12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45945">
        <property type="entry name" value="REGULATOR OF G-PROTEIN SIGNALING LOCO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02188">
        <property type="entry name" value="GoLoco"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00595">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00640">
        <property type="entry name" value="PID"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02196">
        <property type="entry name" value="RBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00615">
        <property type="entry name" value="RGS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16613">
        <property type="entry name" value="RGS12_us1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16611">
        <property type="entry name" value="RGS12_us2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16612">
        <property type="entry name" value="RGS12_usC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01301">
        <property type="entry name" value="RGSPROTEIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00390">
        <property type="entry name" value="GoLoco"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00228">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00462">
        <property type="entry name" value="PTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00455">
        <property type="entry name" value="RBD"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00315">
        <property type="entry name" value="RGS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50156">
        <property type="entry name" value="PDZ domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48097">
        <property type="entry name" value="Regulator of G-protein signaling, RGS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54236">
        <property type="entry name" value="Ubiquitin-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50877">
        <property type="entry name" value="GOLOCO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50106">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="PID"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="RBD"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50132">
        <property type="entry name" value="RGS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="13" id="KW-0966">Cell projection</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="7" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="5" id="KW-0488">Methylation</keyword>
    <keyword evidence="12" id="KW-0539">Nucleus</keyword>
    <keyword evidence="8" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0677">Repeat</keyword>
    <keyword evidence="11" id="KW-0770">Synapse</keyword>
    <keyword evidence="10" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="PDZ" evidence="19">
        <location>
            <begin position="22"/>
            <end position="99"/>
        </location>
    </feature>
    <feature type="domain" description="PID" evidence="18">
        <location>
            <begin position="227"/>
            <end position="339"/>
        </location>
    </feature>
    <feature type="domain" description="RGS" evidence="20">
        <location>
            <begin position="708"/>
            <end position="825"/>
        </location>
    </feature>
    <feature type="domain" description="RBD" evidence="21">
        <location>
            <begin position="901"/>
            <end position="971"/>
        </location>
    </feature>
    <feature type="domain" description="RBD" evidence="21">
        <location>
            <begin position="973"/>
            <end position="1043"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="835"/>
            <end position="854"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1054"/>
            <end position="1103"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1159"/>
            <end position="1219"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="840"/>
            <end position="849"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="1061"/>
            <end position="1088"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="17">
        <location>
            <begin position="1090"/>
            <end position="1103"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="1193"/>
            <end position="1208"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004279"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00034103"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00059154"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00069205"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS01179"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50106"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50132"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50898"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029582.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1313" mass="145491" checksum="C7F4ABED4C140D51" modified="2019-05-08" version="1">MYRPGETVKCRLNMHAPPRIRSVEVARGRAGYGFTLSGQAPCVLSCVMKGSPADYVGLKAGDEIFAINEINVKQASHEDVVKLIGKCSGVLHMVIAEGNSHIDSCSSDEEVGFYDGKGWLKPKPDSKALGINRAEKVVEEMQSGGIFNMIFENPSFCAGNAENSVPKQRSLSESSAITFETGHESINNNPNLLSKEEISKVLHDDSVFQIGLENPEDFGLDASILNVAMVVGYLGSIELPSTSSNLENDSLQAIRGCMRRLRAEQKIHSLVMMKIMHDCIQLCSDKSGVVAEYPAEKLAFSAVCPDDRRFFGLVTMQTNDDASLAQEDEGVLRTSCHVFMVDPELFHHKIHQGIARRFGLECTADPDTNGCLEFPTSSLPVLQFISVLYRDMGELIEGMRARAFLDGDADAHQNNSTSSNSDSGIGNFNQEEKSNRVLVVDLGGNPSKHIPNSIWENPVGRGQNQHASHWNGFCQEQEGNAHLEIIQNDKSQNLSKHLSPSARIEVPLVSSRSSVPPSKKSIVGIGNQRWLPVHVLQEWQHGNASDQESYTDSTDGWSSINCGTLPPPMNKIPADRYRVDGSFGQRQLTSYINEWSKNVFCTHKKFGPQHNIRKSKEAKKGAKFGHPVGLALAPQRVSVRRSFGRSKRFSITRSLDDLESATVSDGELNSSDLKDCISDNSLSSNASLPSVQSCRRLRERRVASWAVSFERLLQDPVGVRFFSEFLRKEFSEENILFWQACEYFNHVPAHDKKELSYRAREIFSKFLCSKATTPVNIDSQAQLADDILSAPHPDMFKEQQLQIFNLMKFDSYTRFLKSPLYQECILAEVEGRPLPDPQRVPSSPTSKHSINSERSHISTPKKFDLLDCNTVCLFLTFIFVSDKSDASRLSASVPDKEKSSKYCCINLPDGSSCKMVVKSGFSVKEMLSGLCEKHGINIAAVDLFLVGGDKPLVLHQDSSILDSRDLRLEKRTLFRLDLVPINRSVGLKAKPTKPVTEVLRPVVAKYGLNLNELVAKLNGEQEPLDLGVPISNLDGQRVVLYEKEPTKGRVFTDRQKGAPVKQSTVVTSTSRNQASTGEGRTLGKSNSIKIKGENGKNARDARLSKREESLAKIGEKKYQKINLDEAEEFFELISKAQSNRADDQRGLLRKEDLVLPDFLRLPLSGPEPPSSTPVSSKGLNKRIAKNDGKDKCTQPSGKLETTQNYNENSIKKTGHSENKHKSALTTLNSQKTYNVPFSAPLSPIPHVQENNMTIWKRQSGELQAEGIQTIDDENVADLTLVAEGDISSPNSTLLPPPFFPQFVNIVTYSCWSL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INP2</accession>
    <name>A0A452INP2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">glutamate carboxypeptidase II</fullName>
            <ecNumber evidence="18">3.4.17.21</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>Release of an unsubstituted, C-terminal glutamyl residue, typically from Ac-Asp-Glu or folylpoly-gamma-glutamates.</text>
            <dbReference type="EC" id="3.4.17.21"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Membrane</location>
            <topology evidence="2">Single-pass type II membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the peptidase M28 family. M28B subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.4.17.21" evidence="18"/>
    <dbReference type="AlphaFoldDB" id="A0A452INP2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033492.1">
        <property type="protein sequence ID" value="ENSGAGP00000029496.1"/>
        <property type="gene ID" value="ENSGAGG00000020963.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016805">
        <property type="term" value="F:dipeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
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        <property type="term" value="F:metallocarboxypeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd08022">
        <property type="entry name" value="M28_PSMA_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.930.40:FF:000001">
        <property type="entry name" value="N-acetylated-alpha-linked acidic dipeptidase 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.630.10:FF:000009">
        <property type="entry name" value="N-acetylated-alpha-linked acidic dipeptidase 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.50.30.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.930.40">
        <property type="entry name" value="Transferrin receptor-like, dimerisation domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.630.10">
        <property type="entry name" value="Zn peptidases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046450">
        <property type="entry name" value="PA_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007484">
        <property type="entry name" value="Peptidase_M28"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039373">
        <property type="entry name" value="Peptidase_M28B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007365">
        <property type="entry name" value="TFR-like_dimer_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036757">
        <property type="entry name" value="TFR-like_dimer_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10404:SF36">
        <property type="entry name" value="GLUTAMATE CARBOXYPEPTIDASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10404">
        <property type="entry name" value="N-ACETYLATED-ALPHA-LINKED ACIDIC DIPEPTIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04389">
        <property type="entry name" value="Peptidase_M28"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04253">
        <property type="entry name" value="TFR_dimer"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52025">
        <property type="entry name" value="PA domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47672">
        <property type="entry name" value="Transferrin receptor-like dimerisation domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53187">
        <property type="entry name" value="Zn-dependent exopeptidases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0121">Carboxypeptidase</keyword>
    <keyword evidence="12" id="KW-0224">Dipeptidase</keyword>
    <keyword evidence="15" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="14" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13" id="KW-0482">Metalloprotease</keyword>
    <keyword evidence="16" id="KW-0511">Multifunctional enzyme</keyword>
    <keyword evidence="5" id="KW-0645">Protease</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0735">Signal-anchor</keyword>
    <keyword evidence="6" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="9" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="Peptidase M28" evidence="20">
        <location>
            <begin position="231"/>
            <end position="433"/>
        </location>
    </feature>
    <feature type="domain" description="Transferrin receptor-like dimerisation" evidence="19">
        <location>
            <begin position="501"/>
            <end position="621"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004606"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005634"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022645"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022997"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023049"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023268"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00052003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00066561"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="Pfam" id="PF04253"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="Pfam" id="PF04389"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029496.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="624" mass="70601" checksum="AE87A2728D6E0F29" modified="2019-05-08" version="1">FEDLVHNVLNKLVLFLVLTANTCPFLFPSNFTQRPHLAGTAENLRLAQQIQAQWKEFGLDSVELAHYDVLLSYPNETNPNYISIIDEHGTEIFNTSLFEPPPPGYETVTDVVPPYSAFSAPGVPEVKRLELLNSRLPALSKINTSLIPPEYTYRYEEARAVGLPKLPVHPIGYNDAEKLLRDMGGPAPPDSSWKGSLNVSYNIGPGFMSRYSTRKVKMHIHSSNKVTRIYNVIGTIRGATEPDRYVILGGHRDSWVFGGIDPQSGAAVVHEIVRSYGKLKKEGWRPRRTIIFASWDAEEFGLLGSTEWAEENTKLLQARGVAYINADSSVEGNYTLRIDCTPLMYSLVYSLTREIPSPDEGFKGKSLYESWHENNPSTEYKDVPRINKLGSGNDFEVFFQRLGIASGRARYSKNWRVEKYSSYPVYHSVYETYEIVERFYDPTFKNHLTVAQVRGGLVFELADSVVLPFDCRDYAEALRNYVQVIYNLSKNHQMGLTTYSVSFDSLFSAANNFTEAATDFHRRLQQVDINNPIAVRALNDQLMFLERAFIDPLGLPGRPFYRHIIFAPSSHNKYAGESFPGIYDAMFDIENKADQRKAWEEVKRQISIAAFTVQAAAGTLKDLA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INP3</accession>
    <name>A0A452INP3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Ral guanine nucleotide dissociation stimulator</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452INP3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033517.1">
        <property type="protein sequence ID" value="ENSGAGP00000029517.1"/>
        <property type="gene ID" value="ENSGAGG00000021309.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007265">
        <property type="term" value="P:Ras protein signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd17209">
        <property type="entry name" value="RA_RalGDS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00155">
        <property type="entry name" value="RasGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd06224">
        <property type="entry name" value="REM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.840.10:FF:000005">
        <property type="entry name" value="Ral guanine nucleotide dissociation stimulator isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.20.90:FF:000042">
        <property type="entry name" value="Ral guanine nucleotide dissociation stimulator isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.20.90">
        <property type="entry name" value="Phosphatidylinositol 3-kinase Catalytic Subunit, Chain A, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.840.10">
        <property type="entry name" value="Ras guanine-nucleotide exchange factors catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.870.10">
        <property type="entry name" value="Son of sevenless (SoS) protein Chain: S domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000159">
        <property type="entry name" value="RA_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015758">
        <property type="entry name" value="RalGDS_RA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008937">
        <property type="entry name" value="Ras-like_GEF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000651">
        <property type="entry name" value="Ras-like_Gua-exchang_fac_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019804">
        <property type="entry name" value="Ras_G-nucl-exch_fac_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023578">
        <property type="entry name" value="Ras_GEF_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001895">
        <property type="entry name" value="RASGEF_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036964">
        <property type="entry name" value="RASGEF_cat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029071">
        <property type="entry name" value="Ubiquitin-like_domsf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23113">
        <property type="entry name" value="GUANINE NUCLEOTIDE EXCHANGE FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23113:SF35">
        <property type="entry name" value="RAL GUANINE NUCLEOTIDE DISSOCIATION STIMULATOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00788">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00617">
        <property type="entry name" value="RasGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00618">
        <property type="entry name" value="RasGEF_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00314">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00147">
        <property type="entry name" value="RasGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00229">
        <property type="entry name" value="RasGEFN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48366">
        <property type="entry name" value="Ras GEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54236">
        <property type="entry name" value="Ubiquitin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50200">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00720">
        <property type="entry name" value="RASGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50009">
        <property type="entry name" value="RASGEF_CAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50212">
        <property type="entry name" value="RASGEF_NTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1 2" id="KW-0344">Guanine-nucleotide releasing factor</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="N-terminal Ras-GEF" evidence="6">
        <location>
            <begin position="1"/>
            <end position="124"/>
        </location>
    </feature>
    <feature type="domain" description="Ras-GEF" evidence="4">
        <location>
            <begin position="195"/>
            <end position="460"/>
        </location>
    </feature>
    <feature type="domain" description="Ras-associating" evidence="5">
        <location>
            <begin position="611"/>
            <end position="698"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="564"/>
            <end position="585"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="3">
        <location>
            <begin position="564"/>
            <end position="584"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022658"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00168"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50009"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50200"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029517.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="9">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="728" mass="81079" checksum="651DAB8ADDA53856" modified="2019-05-08" version="1">KAGTLEKLVEYLVSAFKGNDSTYVTIFLCTYRAFATTKQVLDLLLNRSAYQGACVLPGLSSQSVGLLLPSTISSILGAWLDQYSEDFRKPPDFACLKQLISYVRQNIPGSDLERRARILLAQFQQQEHSEAEPDGEQRFLPVPVSLSALCGVSWVSPVTPVRRTVYEAVDQTSCTFRIVEENGLGEEKPDFLAFSQEMVAEQFTLMDAELFKKVVPYHCLGCIWSQRDKKGKEHLAPTIRATVSQFNSVTNCVIATCLGDRSLKPQQRAKVVERWIEVARECRILKNFSSLRAILSALQCNAVHRLKKTWDEVSRESFRTFHELSEIFSDENNHSLSRELLIKEGTSKFATLEINPKRAQKRQQQQREMGVMQGTIPYLGTFLTDLVMLDTAMKDFLDGGLINFEKRRKEFEVIAQIKLLQSACNNYSFTQEDGFVEWFHGVERLSEAESYSLSCEIEPLSESASNTLKAKKNTGIIKRWSDRQVPSAEPCTSGSSHSKSFDQLKCGQYLCSGDTADSVSVTSAGSSSSDVEEINISFIPESPDCQEKKFWECTSLSSLDTSGIGSGSSSASSSSVSSTPVTASRTHKRSVSGISSYSSLSLPLYNQQIDDCCIIRVSLAVDNGNMYKSILVTSQDKTPVVIRKAMAKHNLDGDRPEDYELVQIISEERELKIPDNANVFYAMNSAANYDFVLKKRGFSKGGVKIKHGSSSTLPRMKQKGLKIAKGIF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INP5</accession>
    <name>A0A452INP5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">glutamate carboxypeptidase II</fullName>
            <ecNumber evidence="18">3.4.17.21</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>Release of an unsubstituted, C-terminal glutamyl residue, typically from Ac-Asp-Glu or folylpoly-gamma-glutamates.</text>
            <dbReference type="EC" id="3.4.17.21"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Membrane</location>
            <topology evidence="2">Single-pass type II membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the peptidase M28 family. M28B subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.4.17.21" evidence="18"/>
    <dbReference type="AlphaFoldDB" id="A0A452INP5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033481.1">
        <property type="protein sequence ID" value="ENSGAGP00000029485.1"/>
        <property type="gene ID" value="ENSGAGG00000020963.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016805">
        <property type="term" value="F:dipeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004181">
        <property type="term" value="F:metallocarboxypeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd08022">
        <property type="entry name" value="M28_PSMA_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.930.40:FF:000001">
        <property type="entry name" value="N-acetylated-alpha-linked acidic dipeptidase 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.630.10:FF:000009">
        <property type="entry name" value="N-acetylated-alpha-linked acidic dipeptidase 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.50.30.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.930.40">
        <property type="entry name" value="Transferrin receptor-like, dimerisation domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.630.10">
        <property type="entry name" value="Zn peptidases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046450">
        <property type="entry name" value="PA_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007484">
        <property type="entry name" value="Peptidase_M28"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039373">
        <property type="entry name" value="Peptidase_M28B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007365">
        <property type="entry name" value="TFR-like_dimer_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036757">
        <property type="entry name" value="TFR-like_dimer_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10404:SF36">
        <property type="entry name" value="GLUTAMATE CARBOXYPEPTIDASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10404">
        <property type="entry name" value="N-ACETYLATED-ALPHA-LINKED ACIDIC DIPEPTIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04389">
        <property type="entry name" value="Peptidase_M28"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04253">
        <property type="entry name" value="TFR_dimer"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52025">
        <property type="entry name" value="PA domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47672">
        <property type="entry name" value="Transferrin receptor-like dimerisation domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53187">
        <property type="entry name" value="Zn-dependent exopeptidases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0121">Carboxypeptidase</keyword>
    <keyword evidence="12" id="KW-0224">Dipeptidase</keyword>
    <keyword evidence="15" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="14" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13" id="KW-0482">Metalloprotease</keyword>
    <keyword evidence="16" id="KW-0511">Multifunctional enzyme</keyword>
    <keyword evidence="5" id="KW-0645">Protease</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0735">Signal-anchor</keyword>
    <keyword evidence="6" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="9" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="Peptidase M28" evidence="20">
        <location>
            <begin position="249"/>
            <end position="451"/>
        </location>
    </feature>
    <feature type="domain" description="Transferrin receptor-like dimerisation" evidence="19">
        <location>
            <begin position="519"/>
            <end position="639"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004606"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005634"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022645"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022997"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023049"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023268"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00052003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00066561"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="Pfam" id="PF04253"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="Pfam" id="PF04389"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029485.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="642" mass="72630" checksum="42F96797DC65E6AD" modified="2019-05-08" version="1">FAMSKSHVLFIFVCHSYSGDSFHKLTSNYKIQFLVLTANTCPFLFPSNFTQRPHLAGTAENLRLAQQIQAQWKEFGLDSVELAHYDVLLSYPNETNPNYISIIDEHGTEIFNTSLFEPPPPGYETVTDVVPPYSAFSAPGVPEVKRLELLNSRLPALSKINTSLIPPEYTYRYEEARAVGLPKLPVHPIGYNDAEKLLRDMGGPAPPDSSWKGSLNVSYNIGPGFMSRYSTRSVKMHIHSSNKVTRIYNVIGTIRGATEPDRYVILGGHRDSWVFGGIDPQSGAAVVHEIVRSYGKLKKEGWRPRRTIIFASWDAEEFGLLGSTEWAEENTKLLQARGVAYINADSSVEGNYTLRIDCTPLMYSLVYSLTREIPSPDEGFKGKSLYESWHENNPSTEYKDVPRINKLGSGNDFEVFFQRLGIASGRARYSKNWRVEKYSSYPVYHSVYETYEIVERFYDPTFKNHLTVAQVRGGLVFELADSVVLPFDCRDYAEALRNYVQVIYNLSKNHQMGLTTYSVSFDSLFSAANNFTEAATDFHRRLQQVDINNPIAVRALNDQLMFLERAFIDPLGLPGRPFYRHIIFAPSSHNKYAGESFPGIYDAMFDIENKADQRKAWEEVKRQISIAAFTVQAAAGTLKDLA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGS9</accession>
    <name>A0A452GGS9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">BOD1/SHG1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Chromosome</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the BOD1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GGS9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000894.1">
        <property type="protein sequence ID" value="ENSGAGP00000000793.1"/>
        <property type="gene ID" value="ENSGAGG00000000628.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005694">
        <property type="term" value="C:chromosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055264">
        <property type="entry name" value="BOD1/SHG1_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043244">
        <property type="entry name" value="BOD1L1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47391">
        <property type="entry name" value="BIORIENTATION OF CHROMOSOMES IN CELL DIVISION 1 LIKE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47391:SF1">
        <property type="entry name" value="BIORIENTATION OF CHROMOSOMES IN CELL DIVISION 1 LIKE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05205">
        <property type="entry name" value="COMPASS-Shg1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0158">Chromosome</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="BOD1/SHG1" evidence="5">
        <location>
            <begin position="41"/>
            <end position="136"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="146"/>
            <end position="172"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="4">
        <location>
            <begin position="154"/>
            <end position="172"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008463"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022454"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="Pfam" id="PF05205"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000793.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="172" mass="19898" checksum="DEBDA16F35223414" modified="2019-05-08" version="1">MQWNGSTQNVARRRRRQLCRWRWERRLRGPGCAGYQLIALIVEQLKSRGLFDGFRRDCLADVDTKPAYQNLRQKVDNFVSTHLDKQEWNPTMNKNQLRNGLRQSVIQSGMLEAGVDRIISQVVDPKLNHIFRPQIEKAIHEFLAAQKKEETMPALPPEPENQDPPAPSQDAS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGT3</accession>
    <name>A0A452GGT3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">K Homology domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000000900.1">
        <property type="protein sequence ID" value="ENSGAGP00000000799.1"/>
        <property type="gene ID" value="ENSGAGG00000000470.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045087">
        <property type="term" value="P:innate immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd22503">
        <property type="entry name" value="KH-I_ANKHD1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000046">
        <property type="entry name" value="Ankyrin repeat and KH domain-containing protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1370.10:FF:000029">
        <property type="entry name" value="ankyrin repeat and KH domain-containing protein 1 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000012">
        <property type="entry name" value="ankyrin repeat domain-containing protein 17 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000028">
        <property type="entry name" value="ankyrin repeat domain-containing protein 17 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000131">
        <property type="entry name" value="ankyrin repeat domain-containing protein 17 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000014">
        <property type="entry name" value="ankyrin repeat domain-containing protein 17 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000068">
        <property type="entry name" value="ankyrin repeat domain-containing protein 17 isoform X5"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="7"/>
    </dbReference>
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        <property type="entry name" value="K Homology domain, type 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051631">
        <property type="entry name" value="Ankyrin-KH/SAM_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047374">
        <property type="entry name" value="KH-I_ANKHD1"/>
    </dbReference>
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        <property type="entry name" value="KH_dom"/>
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        <property type="entry name" value="KH_dom_type_1"/>
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        <property type="entry name" value="KH_dom_type_1_sf"/>
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        <property type="entry name" value="ANKYRIN REPEAT AND KH DOMAIN-CONTAINING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="MASK PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00013">
        <property type="entry name" value="KH_1"/>
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    </dbReference>
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        <property type="entry name" value="ANKYRIN"/>
    </dbReference>
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        <property type="entry name" value="ANK"/>
        <property type="match status" value="21"/>
    </dbReference>
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        <property type="entry name" value="KH"/>
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        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54791">
        <property type="entry name" value="Eukaryotic type KH-domain (KH-domain type I)"/>
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        <property type="match status" value="16"/>
    </dbReference>
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    <keyword evidence="3 5" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="4 7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0694">RNA-binding</keyword>
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    <feature type="repeat" description="ANK" evidence="5">
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    <feature type="repeat" description="ANK" evidence="5">
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    <feature type="repeat" description="ANK" evidence="5">
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    <feature type="repeat" description="ANK" evidence="5">
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            <begin position="1599"/>
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            <dbReference type="SAM" id="MobiDB-lite"/>
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    <sequence length="2467" mass="262136" checksum="9E9D759606A9ED77" modified="2019-05-08" version="1">MLTDGTAAPVGDEEIDSVTPRAPPTAEAPVAAGPAPLGLRAVRLFGEAGPGGPGGPGAPDAGEAALDFKLAAAVLRSGGGGGGSGSDEDEVSEVESFILDQEDLDNPVLKTTSELFLSSAAEGTDLRTVDPETQARLEALLEAAGIGKLSTADGKAFADPEVLRRLTSSVSCALDEAAAALTRMRAENNHNAGQVDNRSLAEACSDGDVNAVRKLLDEGRSVNEHTEEGESLLCLACSAGYYELAQVLLAMHANVEDRGNKGDITPLMAAASGGYVDIVKLLLVHCADVNAQSSTGNTALTYACAGGFVDIVKVLLKAGANIEDHNENGHTPLMEAASAGHVEVARVLLEYGAGINTHSNEFKESALTLACYKGHLDMVRFLLEAGADQEHKTDEMHTALMEACMDGHVEVARLLLDSGAQVNMPADSFESPLTLAACGGHVELAALLIERGANLEEVNDEGYTPLMEAAREGHEEMVALLLAQGANINAQTEETQETALTLACCGGFSEVADFLIKAGADIELGCSTPLMEAAQEGHLELVKYLLAAGANVHATTATGDTALTYACENGHTDVADVLLQAGVFNLNYSLLDNFKLNFTEKSPFLYSTEARRTPLTVAFSVSKLSLQDGSTMLIEAAKGGHTNVVSYLLDYPNNVLSVPTADMSQLTPPSQDQSQVPRVPMHALAMVVPPQEPDRAPQENSPPLLGVQKGASKQKSSSLQVADQDLLPPFHPYQPLECIVEETEGKLNELGQRISAIEKAQLKSLELIQGEPLNKDKIEELKKNREEQVQKKKKILKELQKVERQLQMKTQQQFTKEYLETKGQTDTPLPLQQQCPHAGVFPEVEGDEGLPEENFSELPQVDTFLSKDDEQQQSSPPAEQIEFVPIQALSTPQCNFSSNLGYNGTDSLDLQKVLGNQQNVGQQQQIAGHGQGLLVQEPDGLMVATPAQTLTDTLDDLIAAVNSRVPTGSTSSSLTTESPAPEPCAQTPSNVGSQSVLPMYPSVDIDAHVSNLATDLSSSCNTESNHDTALTLACAGGHEELVVDLLLARGANKEHRNVSDYTPLSLAASGGYVNIIKILLNAGAEINSRTGSKLGISPLMLAAMNGHVPAVKLLLDMGSDINAQIETNRNTALTLACFQGRAEVVSLLLDRKANVEHRAKTGLTPLMEAASGGYAEVGRVLLDKGADVNAPPVPSSRDTALTIAADKGHYKFCELLINRGAHIDVRNKKGNTPLWLAANGGHFDVVQLLVQAGADVDAADNRKITPLMSAFRKGHVKVVQYLVKEVNQFPSDIECMRYIATITDKELLKKCHQCVETIVKAKDQQAAEANKNASILLKELDLEKSREESRKQALAAKREKRKEKRKKKKEEQKRKQEEDEENKPKETLELQEDDDEEENDDEEVPIEPPSATTTTTIGISATSATFTNVFGKKRANVVTTPSTNRKNKKNKTKDTPQNVQIILPDQHISLAQQKADKNKINGEPRGGGAGGNSDSDNLDSTDCNSESSSGSKSQELNFTMDTNSSGDRGYASLLLPSQEEKTSTSASKIQTRLEGDANPNSLSASYKSVSLPLTSPNIKLNLTSPKRGQKREEGWKEVVRRSKKLSVPASVVSRIMGRGGCNITAIQDVTGAHIDVDKQKDKNGERMITIRGGTESTRYAVQLISALIQDPAKELEDLIPKTHIRTPAPNTKSIHANFSSGVNTAAASNKNSFPLGAPPLVTSQSSTLSTFQTTNKLNKNVPANVRSSFPVSLPLAYSHPHFALLAAQTMQQIRHPRLPMAQFGGTFSPSPNTWGPFPVRPVNPSSTNSSPKHNSSSRVGNQNGNILQTESTGLATASCPITVSSVVASTQPLCATSSRTPSSVRKQLFACVPKTSAVATAISTVTSTCSTLPSPSSAPANNGQVPTAFLPSNVPQTQHSALKTDSFSAISAPKEKVSTLDQPTGNTCAPSSVASSCNISASSNPGVLETRPSSSPAPLNNVQDEILPTSMSDISSSVSMPFSSSSEPTPLSLASPRSIAADNQDNSNLPQLAVPAPRVSHRMQPRGSFYSVVPNANLHQDHQSIFVTNPVPLTPSQGPPAAVQLSSAMNVMNGSQMHINPANKSLPPTFGPATLFNHFSSLFDSNQVPANQGWGDCPLSTRAAADPSFTVQSTFLSNSVLGHIENVHPDNSKAPGFRPPSQRVSTSPVGLPSLDPSNGSSAASSAPLTGFSANIQRVYLQGPAPVGTPSFNRQHFSPHPWTSATNSCESPIPSVSSGSSSPLSAASAPSNLGQPKAGSTSQDRKVPPPIGTERLARIRQGGSVTPTPLGTNFTAPVGHSGIWSFGVNSVSEGMSGWSQPIMGNHPMHQQLSDPGTFSQHQPMERDDSGIVAPSNIFHQPMPNSFGLPISMYGGALIPSHPQLADGPGGPLFNGLHTPDPAWNPMIKVVQNSTECTDAQQVNLFDPPLLPSSHPWHHLSPPRRRGPR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGW5</accession>
    <name>A0A452GGW5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">protein-tyrosine-phosphatase</fullName>
            <ecNumber evidence="2">3.1.3.48</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="12">
            <text>O-phospho-L-tyrosyl-[protein] + H2O = L-tyrosyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:10684"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="EC" id="3.1.3.48"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="11">Belongs to the protein-tyrosine phosphatase family. Receptor class 3 subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.48" evidence="2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000930.1">
        <property type="protein sequence ID" value="ENSGAGP00000000823.1"/>
        <property type="gene ID" value="ENSGAGG00000000682.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043235">
        <property type="term" value="C:receptor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045296">
        <property type="term" value="F:cadherin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001525">
        <property type="term" value="P:angiogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="CDD" id="cd14617">
        <property type="entry name" value="R-PTPc-B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000369">
        <property type="entry name" value="Protein tyrosine phosphatase, receptor type B"/>
        <property type="match status" value="12"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:001376">
        <property type="entry name" value="Protein tyrosine phosphatase, receptor type B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000009">
        <property type="entry name" value="Receptor-type tyrosine-protein phosphatase beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000242">
        <property type="entry name" value="PTP_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041201">
        <property type="entry name" value="PTPRJ_TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050713">
        <property type="entry name" value="RTP_Phos/Ushers"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016130">
        <property type="entry name" value="Tyr_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003595">
        <property type="entry name" value="Tyr_Pase_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46957">
        <property type="entry name" value="CYTOKINE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46957:SF2">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE BETA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18861">
        <property type="entry name" value="PTP_tm"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00102">
        <property type="entry name" value="Y_phosphatase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00700">
        <property type="entry name" value="PRTYPHPHTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="17"/>
    </dbReference>
    <dbReference type="SMART" id="SM00194">
        <property type="entry name" value="PTPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00404">
        <property type="entry name" value="PTPc_motif"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="16"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00383">
        <property type="entry name" value="TYR_PHOSPHATASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50055">
        <property type="entry name" value="TYR_PHOSPHATASE_PTP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="9 13" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="4 14" id="KW-0732">Signal</keyword>
    <keyword evidence="3 13" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 13" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="14">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019518794" description="protein-tyrosine-phosphatase" evidence="14">
        <location>
            <begin position="23"/>
            <end position="1999"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="13">
        <location>
            <begin position="1618"/>
            <end position="1644"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="17">
        <location>
            <begin position="114"/>
            <end position="207"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="17">
        <location>
            <begin position="293"/>
            <end position="380"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="17">
        <location>
            <begin position="471"/>
            <end position="558"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="17">
        <location>
            <begin position="644"/>
            <end position="731"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="17">
        <location>
            <begin position="732"/>
            <end position="822"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="17">
        <location>
            <begin position="911"/>
            <end position="998"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="17">
        <location>
            <begin position="1177"/>
            <end position="1264"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="17">
        <location>
            <begin position="1265"/>
            <end position="1358"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="17">
        <location>
            <begin position="1359"/>
            <end position="1450"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="15">
        <location>
            <begin position="1705"/>
            <end position="1965"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="16">
        <location>
            <begin position="1883"/>
            <end position="1956"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00013064"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00025789"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00051722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000823.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1999" mass="224481" checksum="57742A768D2D03F8" modified="2019-05-08" version="1">MLRSAAGVSLWLALALIKKGLGEPVKCAINLTEWRVSSQSVYLRWSSLGNLCNFTLTCSSDNVSHTECHPVQKTNQSYECSYNGLEAGTLYHLRIFSLSDTETSNISLQTDPLPPARFEINKEKTTSTSFQVSWSPSSGKVDWYQLQLSDHNNKKIQNVSVQGSISRREETFSNLTPGSKYNVIITAVAGNKSSPAIHISGSTVPSPVQNIQISVKTDTIHVSWLPGSGYVDRYRLVLLDKEGPVREVTHEERLLSYTFLGLTTGHLYNLTIIAEAAGLESYNFKLVRTAPAQVSDLMVTNDGSFDILKVTWKRPLGNLDFYNITLSHLGSIKEIKTLQPQVTEVQFDKLIPGSLYQVTVSTISGELFTDKMATGRTVPQKVSELKASSGGWLRSLTVNWLPPAGDWDRYHIVVFNLSTVLLNTTLKKDRREYVIQDIGLIPGRQYEVAVIVESGGLQNTARCKGRTAPQPVLQLRVKHANESSLSVMWMTPVAEWDSYVVSLGDRDLTIINKALVKEAKEFTFSDLVPGRKYTATVTSISGDLSNWTSVEGRTVPAQVTSLHVTNQGATNSLFTNWTRALGDVESYQVLLIHENVVIKNESVSSETSRYHFHSLKAGGLYSVVVTTVSGGIWSRQTIAEGRTVPSSVSGVTVNNSGRSDYLSVSWLPATGDVDSYLVTLSHDSQTIQTLTISKALSECSFSSLIAGRLYNVMITTKSGKYENHSFSQERTVPSSVQGLTVSNSARSDYLKVSWLHATGDFDNYKVIIKNNNDFNQTKSVPKDENECVFTKLVPGRQYSVTVSTQSGKYETSKRINGRTMPESVKELTLSNRSTEDLQVTWSKAEGDVDKYEIQLLFNDMKVFPPIFLGNTIEEYWFTALTPGRLYKILVLTISGDTQRATFIEGLTAPSAVKNIHVSANGMTNSLKVNWTPGGGDVDSYTVTLFHQNYQLDSQTVSKPVYEHTFSKLEAGEQYRVVVQSNSGTLHNSLTALGRTIPASVQGLFADHAYSSHSLLVTWHSAPGVAERYDILLLTEQGILLSNKSEPATAKQHKFEDLIPGKKYKVQMLTVSGGLFSRRAETEGRTVPAAVTNLKVTENINGRLSFSWTTSQGELDSYNIFLYNPDKSLQDRISGDQNLQQCSFQNLRQGRIYKMVIVTHSGELTNESSVFGRTVPAPVINLSGSNRNMTNSLWFTWGPASGDVDYYELNLYNPNGTQKETKHGKDLREWHFLGLVPGRKYTLVVVTHSGDLTNTAKTEGRTAPSPPSAVSFADVANTSLSITWLGPPDWTDYDDFELQWFPRDPLTVFNPYSNSRSKGRIMYGLRPGRLYNFSVRSVSGNTWKTYSQSLSGTVRTRPDRIQNLHCRPQNSTAIACSWTPPDSDFDGYSIECKKMDTGEVEFSKRIEKERSLQNIMMLVPHKRYLVSIKVQSADMTSEVAEDSTITMIDRPPPPPLHIRVNKKDALITKSSINFTFNCSWFSDTNGAVKYFTVVVREADGSEELKPDQQHPLPSYMEYKHNNSIRVYQTNYFASKCAENLDSNYKSFDIKLGGEMESLGGKCDPNQQKFCDGPLKPRTAYRISIRAFTQLFSEDMREFTEPLFSDTFFSLPITTQSEPMFGVIEGVSAGLFLILMVVAVTALFVYRQKINNGHERPTARLSIRRDRPLSVHLNLGQRGSRRISSPIKVNQFEAHFTKLQADSNYLLSKEYEDLKDVGRNQSCDIALLPENRGKNRYNNILPYDTSRVKLSNVDDDPCSDYINASYIPGNNFRREYIATQGPLPGTKDDFWKMAWEQNVHNIVMVTQCVEKGRVKCDHYWPLDQDSLYYGDLIVEMLSESVLPEWTIREFKICSEEQLDSKRLIRHFHYTVWPDHGVPETTQSLIQFVRTVRDYINRTPDTGPTILHCSAGVGRTGTFIALDRILQQLDSKDSVDIYGAVHDLRLHRVHMVQTECQYVYLHQCVRDVLRARKLRSEQENPLFPIYENVNPECHRDAVYSRH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGW8</accession>
    <name>A0A452GGW8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Zinc finger protein 883-like</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GGW8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000952.1">
        <property type="protein sequence ID" value="ENSGAGP00000000837.1"/>
        <property type="gene ID" value="ENSGAGG00000000691.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031981">
        <property type="term" value="C:nuclear lumen"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001228">
        <property type="term" value="F:DNA-binding transcription activator activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd07765">
        <property type="entry name" value="KRAB_A-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002063">
        <property type="entry name" value="RB associated KRAB zinc finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000056">
        <property type="entry name" value="Zinc finger and SCAN domain-containing 20"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002343">
        <property type="entry name" value="Zinc finger protein 33A"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000016">
        <property type="entry name" value="zinc finger protein 37 homolog"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000212">
        <property type="entry name" value="zinc finger protein 382 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002090">
        <property type="entry name" value="Zinc finger protein 473"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001270">
        <property type="entry name" value="zinc finger protein 583 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001437">
        <property type="entry name" value="Zinc finger protein 594"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000176">
        <property type="entry name" value="zinc finger protein 70"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001909">
        <property type="entry name" value="KRAB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036051">
        <property type="entry name" value="KRAB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24393">
        <property type="entry name" value="ZINC FINGER PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24393:SF100">
        <property type="entry name" value="ZINC FINGER PROTEIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01352">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="SMART" id="SM00349">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109640">
        <property type="entry name" value="KRAB domain (Kruppel-associated box)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50805">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="15"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="9" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 11" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="KRAB" evidence="13">
        <location>
            <begin position="11"/>
            <end position="88"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="130"/>
            <end position="157"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="158"/>
            <end position="185"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="186"/>
            <end position="213"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="214"/>
            <end position="241"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="242"/>
            <end position="269"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="270"/>
            <end position="297"/>
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        <location>
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            <end position="325"/>
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    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="326"/>
            <end position="353"/>
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    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="354"/>
            <end position="381"/>
        </location>
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        <location>
            <begin position="382"/>
            <end position="409"/>
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        <location>
            <begin position="410"/>
            <end position="437"/>
        </location>
    </feature>
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        <location>
            <begin position="438"/>
            <end position="465"/>
        </location>
    </feature>
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        <location>
            <begin position="465"/>
            <end position="492"/>
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        <location>
            <begin position="493"/>
            <end position="520"/>
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    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="521"/>
            <end position="548"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
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            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000837.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="16">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="583" mass="67036" checksum="53F1FEAD1F7AFD4A" modified="2019-05-08" version="1">MGDNCYFQEPVTFEEVAVYFTEEEWALLDPGQRALYRDVMQENYENHEEVNPQLGGLEQVEQHGTLLGRSKGNICWSVGQGKTCESQCSTERLLGNQPVKKVDKSTNCGRGLKHLKKTRVQRIHLGERPSTCSECGKSFSSHSHLLIHQRIHTGEKPYKCPQCGKSFNQSPNLVTHLRIHTGERPYKCQECGKSFTANSTFIKHQRIHTGEKPYKCPDCGRSFNQSSDLIRHQRLHTGEKKTTCTKCGTNFKQRSDLIRHERIHTGEKLYNCPNCGKSFNNSSALTLHQKTHTGERPYKCLNFGKNFTRSSRLTEHQRFHTGEKPYKCLDCGKSFSVNSALIRHWRVHTGEKPYKCPDCDKRFNVSSSLKTHRRIHTGERLYRCPDCGKSFKERSSLINHERIHTGERPFKCLDCGKNFNQISALLLHQRLHTEERPYKCPDCGKNFRVSSALIRHQRIHTEKPYKCPDCGKSFSISSSLITHQRIHTEERPYKCSSCRKSFNISSSLIKHQRMHTEERLYKWTNCGESFMQSSYLIKHQRIHIAETPINAPVSSHSKNKPFSDFPPTSVTFGSQICGCPSQG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGX1</accession>
    <name>A0A452GGX1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Serine/threonine-protein kinase</fullName>
            <ecNumber evidence="20">2.7.11.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="26 27">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="27" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="26" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="26" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="20">Proto-oncogene with serine/threonine kinase activity involved in cell survival and cell proliferation.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subunit">
        <text evidence="17">Interacts with RP9. Interacts with HSP90AA1, this interaction stabilizes PIM1 protein levels. Interacts (ubiquitinated form) with HSP70 and promotes its proteasomal degradation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4 20">Belongs to the protein kinase superfamily. CAMK Ser/Thr protein kinase family. PIM subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="20"/>
    <dbReference type="AlphaFoldDB" id="A0A452GGX1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000954.1">
        <property type="protein sequence ID" value="ENSGAGP00000000839.1"/>
        <property type="gene ID" value="ENSGAGG00000000395.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0106310">
        <property type="term" value="F:protein serine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006915">
        <property type="term" value="P:apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043066">
        <property type="term" value="P:negative regulation of apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000209">
        <property type="entry name" value="Serine/threonine-protein kinase pim-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000232">
        <property type="entry name" value="Serine/threonine-protein kinase pim-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017348">
        <property type="entry name" value="PIM1/2/3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051138">
        <property type="entry name" value="PIM_Ser/Thr_kinase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22984">
        <property type="entry name" value="SERINE/THREONINE-PROTEIN KINASE PIM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22984:SF29">
        <property type="entry name" value="SERINE_THREONINE-PROTEIN KINASE PIM-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF037993">
        <property type="entry name" value="STPK_Pim-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0053">Apoptosis</keyword>
    <keyword evidence="12 20 22" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="16" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11 20" id="KW-0418">Kinase</keyword>
    <keyword evidence="13" id="KW-0460">Magnesium</keyword>
    <keyword evidence="10 20 23" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="15" id="KW-0539">Nucleus</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="27" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6 20 24" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="8 20" id="KW-0808">Transferase</keyword>
    <keyword evidence="14" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="Protein kinase" evidence="25">
        <location>
            <begin position="38"/>
            <end position="290"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="21">
        <location>
            <position position="167"/>
        </location>
    </feature>
    <feature type="binding site" evidence="22">
        <location>
            <begin position="44"/>
            <end position="52"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="22 23">
        <location>
            <position position="67"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="22">
        <location>
            <position position="121"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="22">
        <location>
            <position position="128"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00005505"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022703"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00047040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00047899"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00048679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="PIRNR" id="PIRNR037993"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="PIRSR" id="PIRSR037993-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="PIRSR" id="PIRSR037993-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="RuleBase" id="RU000304"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000839.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="304" mass="34245" checksum="DAC5F558DA161594" modified="2019-05-08" version="1">MLLSKINSLAHLCSGSGGELHPAKLQPGKEKEPLEKLYQVGPLLGSGGFGSVYSGTRLSDGAPVAIKHVARDRISDWGELPGGSRVPMEILLLKKVGSGFRGVIRLLDWFERPDSFVLVLERPDPVQDLFDFITERGALPEELARSFFRQVLEAVRHCHSCGVLHRDIKDENILIDLNSGELKLIDFGSGALLKDTVYTDFDGTRVYSPPEWIRYHRYHGRSAAVWSLGVLLYDMVCGDIPFEHDEEIIRGQVYFRQRVSPECQHLIKWCLALRPSDRPSFEDLLNHSWMQDVLFVPDLLSPGI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGX3</accession>
    <name>A0A452GGX3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase</fullName>
            <ecNumber evidence="15">2.7.4.24</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Bifunctional inositol kinase that acts in concert with the IP6K kinases IP6K1, IP6K2 and IP6K3 to synthesize the diphosphate group-containing inositol pyrophosphates diphosphoinositol pentakisphosphate, PP-InsP5, and bis-diphosphoinositol tetrakisphosphate, (PP)2-InsP4. PP-InsP5 and (PP)2-InsP4, also respectively called InsP7 and InsP8, regulate a variety of cellular processes, including apoptosis, vesicle trafficking, cytoskeletal dynamics, exocytosis, insulin signaling and neutrophil activation. Phosphorylates inositol hexakisphosphate (InsP6) at position 1 to produce PP-InsP5 which is in turn phosphorylated by IP6Ks to produce (PP)2-InsP4. Alternatively, phosphorylates PP-InsP5 at position 1, produced by IP6Ks from InsP6, to produce (PP)2-InsP4. Activated when cells are exposed to hyperosmotic stress.</text>
    </comment>
    <comment type="function">
        <text evidence="15">Bifunctional inositol kinase that acts in concert with the IP6K kinases to synthesize the diphosphate group-containing inositol pyrophosphates diphosphoinositol pentakisphosphate, PP-InsP5, and bis-diphosphoinositol tetrakisphosphate, (PP)2-InsP4. PP-InsP5 and (PP)2-InsP4, also respectively called InsP7 and InsP8, may regulate a variety of cellular processes, including apoptosis, vesicle trafficking, cytoskeletal dynamics, and exocytosis. Phosphorylates inositol hexakisphosphate (InsP6).</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>1D-myo-inositol hexakisphosphate + ATP = 1-diphospho-1D-myo-inositol 2,3,4,5,6-pentakisphosphate + ADP</text>
            <dbReference type="Rhea" id="RHEA:37459"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:58130"/>
            <dbReference type="ChEBI" id="CHEBI:74946"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.4.24"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="13">
            <dbReference type="Rhea" id="RHEA:37460"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="12">
            <text>5-diphospho-1D-myo-inositol 1,2,3,4,6-pentakisphosphate + ATP + H(+) = 1,5-bis(diphospho)-1D-myo-inositol 2,3,4,6-tetrakisphosphate + ADP</text>
            <dbReference type="Rhea" id="RHEA:10276"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:58628"/>
            <dbReference type="ChEBI" id="CHEBI:77983"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.4.24"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="12">
            <dbReference type="Rhea" id="RHEA:10277"/>
        </physiologicalReaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2 15">Cytoplasm</location>
            <location evidence="2 15">Cytosol</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 15">Belongs to the histidine acid phosphatase family. VIP1 subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.4.24" evidence="15"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000923.1">
        <property type="protein sequence ID" value="ENSGAGP00000000819.1"/>
        <property type="gene ID" value="ENSGAGG00000000614.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033857">
        <property type="term" value="F:5-diphosphoinositol pentakisphosphate 1-kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0052723">
        <property type="term" value="F:inositol hexakisphosphate 1-kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="RHEA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006020">
        <property type="term" value="P:inositol metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032958">
        <property type="term" value="P:inositol phosphate biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07061">
        <property type="entry name" value="HP_HAP_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.470.20:FF:000003">
        <property type="entry name" value="Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.11950:FF:000001">
        <property type="entry name" value="Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.11950:FF:000003">
        <property type="entry name" value="Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1240:FF:000013">
        <property type="entry name" value="Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.11950">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.470.20">
        <property type="entry name" value="ATP-grasp fold, B domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1240">
        <property type="entry name" value="Phosphoglycerate mutase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033379">
        <property type="entry name" value="Acid_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000560">
        <property type="entry name" value="His_Pase_clade-2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037446">
        <property type="entry name" value="His_Pase_VIP1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029033">
        <property type="entry name" value="His_PPase_superfam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040557">
        <property type="entry name" value="VIP1_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12750">
        <property type="entry name" value="DIPHOSPHOINOSITOL PENTAKISPHOSPHATE KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12750:SF11">
        <property type="entry name" value="INOSITOL HEXAKISPHOSPHATE AND DIPHOSPHOINOSITOL-PENTAKISPHOSPHATE KINASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00328">
        <property type="entry name" value="His_Phos_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18086">
        <property type="entry name" value="PPIP5K2_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53254">
        <property type="entry name" value="Phosphoglycerate mutase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00616">
        <property type="entry name" value="HIS_ACID_PHOSPHAT_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10 15" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="5 15" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9 15" id="KW-0418">Kinase</keyword>
    <keyword evidence="11" id="KW-0472">Membrane</keyword>
    <keyword evidence="8 15" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7 15" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="VIP1 N-terminal" evidence="17">
        <location>
            <begin position="55"/>
            <end position="144"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="910"/>
            <end position="974"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1087"/>
            <end position="1160"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1193"/>
            <end position="1361"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1"/>
            <end position="15"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="16">
        <location>
            <begin position="1101"/>
            <end position="1118"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1131"/>
            <end position="1149"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1323"/>
            <end position="1345"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004514"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005609"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00033696"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00034629"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00037056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="RuleBase" id="RU365032"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF18086"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000819.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1361" mass="152431" checksum="15C0B3FEC7738963" modified="2019-05-08" version="1">MSSLTTSHEGASSTPRFFVGTRDDETDFLGSNMKTDEIDFFEDDEESESPPERQIVVGICAMTKKSKSKPMTQILERLCKFEYITVVIMGEDVILNEPVENWPPCDCLISFHSKGFPLDKAVAYAKLCNPFLINDLDMQYYIQDRREVYRILQEEGIDLPRYAVLNRDPDKPDECSLVEGEDQVEVNGAVFPKPFVEKPVSAEDHNVYIYYPTSAGGGSQRLFRKIGSRSSVYSPESCVRKTGSYIYEEFMPTDGTDVKVYTVGPDYAHAEARKSPALDGKVERDSEGKEIRYPVMLTAMEKLVARKVCVAFKQTVCGFDLLRANGHSFVCDVNGFSFVKNSMKYYDDCTKILGNIIMRELAPQFQIPWSIPTEAEDIPIVPTTSGTMMELRCVIAVIRHGDRTPKQKMKMEVKHSRFFELFEKYDGYKTGKLKLKKPEQLQEVLDIARLLVVELGTHNDCEIEERKSKLEQLKTVLEMYGHFSGINRKVQLTYLPRGHPKASSEDEDARKEPSPSLLLVLKWGGELTPAGRVQAEELGRAFRCMYPGGQGDYAGFPGCGLLRLHSTYRHDLKIYASDEGRVQMTAAAFAKGLLALEGELTPILVQMVKSANMNGLLDSDSDSLSSCQHRVKARLHEIMQKDAKFCEEDYEKLAPTGSISLVSSMTFIQNPVEICDQVFALIENLTSQICKRLEDPKSADLQLYHSETLELMLQRWSKLERDFRMKNGRYDISKIPDIYDCIKYDVQHNGTLKLEGTAELFKLSKALADVIIPQEYGINKEEKLEIAIGFCLPLIKKIQLDLQRTHEDESVNKLHPLYSRGVLSPGRHVRTRLYFTSESHVHSLLSIFRYGGLLDESKDQQWKRAMDYLSAISELNYMTQIVIMLYEDNNKDPSSEERFHVELHFSPGVKGCEEDGSAPTGFGFRPASAENDDKKPNQGSMEDLSKVKSIDETDQAQERSPQPSEPICIQRRSPLIRNRKTGSMEVLSESSSKTGGYRLFSSYSRQSTEMKQSGLGFEGCSMVPTIYPLETLHNSLSLRQVNEFLTSVCRCYSSLHSRASTALFDSVIASQTPGDPFMPQRILSSSSLPLRQRSDKPPWYSSGPSSTVSSAGPSSPTSMENYARFGFTEKPSISPQISEERLNCQSPEQSEMEPSGEALELDLGGSGSLVENPLGEMPEQGLMRMPGLLEVPEPSKEELSEEVPEQGTMRVLGLSKTRQSGKTSEKDGSVVLEQGGEKPAGGTWELGETGKLEAGEIGLSGKISDSDEEGPAGEMLEPDVVRPVAPGETSELDEEEPVMQTLEPSEKGQPVEILEKEDEELSTDNALSDQQPFSHLYQDQQLSSEKSVEDEQPSSQQELQN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGX4</accession>
    <name>A0A452GGX4_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000000924.1">
        <property type="protein sequence ID" value="ENSGAGP00000000820.1"/>
        <property type="gene ID" value="ENSGAGG00000000675.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051726">
        <property type="term" value="P:regulation of cell cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048024">
        <property type="term" value="P:regulation of mRNA splicing, via spliceosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032922">
        <property type="entry name" value="SON"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46528">
        <property type="entry name" value="PROTEIN SON"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46528:SF1">
        <property type="entry name" value="PROTEIN SON"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17069">
        <property type="entry name" value="RSRP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1"/>
            <end position="63"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="88"/>
            <end position="172"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1163"/>
            <end position="1184"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="2377"/>
            <end position="4117"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="39"/>
            <end position="63"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="1">
        <location>
            <begin position="118"/>
            <end position="148"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="2377"/>
            <end position="2395"/>
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        <name type="scientific">Gopherus agassizii</name>
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            <taxon>Durocryptodira</taxon>
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        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
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                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
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        <text evidence="13">The conversion to 3-oxoalanine (also known as C-formylglycine, FGly), of a serine or cysteine residue in prokaryotes and of a cysteine residue in eukaryotes, is critical for catalytic activity.</text>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF00884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF12548"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000845.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="797" mass="92816" checksum="FDD251B4512ADB3E" modified="2019-05-08" version="1">MQVMNKTRRIMEQGGAHFINAFVTTPMCCPSRSSILTGKYVHNHNTYTNNENCSSPSWQAQHEIRTFAIYLNNTGYRTAFFGKYLNEYNGSYVPPGWKEWVGLLKNSRFYNYTLCRNGVKEKHGFDYSRDYLTDLITNDSVSFFRISKKMYPHRPVLMVLSHAAPHGPEDSAPQYSQLFPNASQHITPSYNYAPNPDKHWIMRYTGPMKPIHMEFTNMLQRKRLQTLMSVDDSMEMIYNTLVETGELDNTYIIYTADHGYHIGQFGLVKGKSMPYEFDIRVPFYVRGPNVEAGSLNPHIVLNIDLAPTILDIAGLDIPPDMDGKSILKLLDSERPVNRFHLKKKMKVWRDSFLVERGKLLHKRENEKVDAQEENFLPKYQRVKDLCQRAEYQTACEQLGQKWQCIEDASGKLKLHKCKGMAHLAAASSSKGLSNLMPKYYSRKSEDCNCEENEYKLSPIGQRTKLFKKQYKPSYVRNRSIRSVSIELDGEVYNLGLEDGYQPVLPRNITKRHKAEGDAAGEEDKDIGEYSGTGGIAEYTAPNLIKVTHRCYILENDTVQCDTDLYKSLQAWKDHKLHIDHEIETLQNKIKNLREVRGHLKKKRPEECDCNKIRKVLQEKDKLWLLREQRRKKKLRKLLKRIKNNDTCSMPGLTCFTHDNQHWQTAPLWTLGPFCACTSANNNTYWCMRTINETHNFLFCEFATGFLEYFDLNTDPYQLINAVNTLDRDILNQLHIQLMELRSCKGYKQCNPRTRNMDLGLKDGSYEQFRQFQRRRWPEMKRPSSSKSLGQLWEGWEG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGX8</accession>
    <name>A0A452GGX8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Folate receptor-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the folate receptor family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GGX8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000953.1">
        <property type="protein sequence ID" value="ENSGAGP00000000838.1"/>
        <property type="gene ID" value="ENSGAGG00000000517.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009897">
        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0038023">
        <property type="term" value="F:signaling receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004269">
        <property type="entry name" value="Folate_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018143">
        <property type="entry name" value="Folate_rcpt-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10517">
        <property type="entry name" value="FOLATE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10517:SF14">
        <property type="entry name" value="FOLATE RECEPTOR 1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03024">
        <property type="entry name" value="Folate_rec"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 4" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="4">
        <location>
            <begin position="1"/>
            <end position="33"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019347687" description="Folate receptor-like domain-containing protein" evidence="4">
        <location>
            <begin position="34"/>
            <end position="239"/>
        </location>
    </feature>
    <feature type="domain" description="Folate receptor-like" evidence="5">
        <location>
            <begin position="41"/>
            <end position="195"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007932"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="Pfam" id="PF03024"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000838.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="239" mass="26909" checksum="4B8B4786EC1D0628" modified="2019-05-08" version="1">GSVSATENQLMCRLQHTLWWAVLGLLAACVASAAKESVLNVCMDAKHHKTKPGPEGALHGQCALWKDNACCTAETSTGAHQDQSYLYNFNWNHCGVMPEMCKRHFIQDTCLYECSPNLGPWIDQADTSWRRERILNVPLCKEDCELWWEACKDAVTCKENWHKGWNWTSADLCEKIWSNSYKYTTEHWGSGRCIQMWFDPANGNPNVAVAKYYAQNGRDASPAPQAVLLLLPAALLSLL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGY3</accession>
    <name>A0A452GGY3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Alpha-protein kinase 3</fullName>
            <ecNumber evidence="3">2.7.11.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">Involved in cardiomyocyte differentiation.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="14">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the protein kinase superfamily. Alpha-type protein kinase family. ALPK subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000972.1">
        <property type="protein sequence ID" value="ENSGAGP00000000855.1"/>
        <property type="gene ID" value="ENSGAGG00000000711.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055013">
        <property type="term" value="P:cardiac muscle cell development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd16973">
        <property type="entry name" value="Alpha_kinase_ALPK3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00096">
        <property type="entry name" value="Ig"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000069">
        <property type="entry name" value="Alpha-protein kinase 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000543">
        <property type="entry name" value="Alpha-protein kinase 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.20.200.10:FF:000003">
        <property type="entry name" value="alpha-protein kinase 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.20.200.10">
        <property type="entry name" value="MHCK/EF2 kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004166">
        <property type="entry name" value="a-kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013098">
        <property type="entry name" value="Ig_I-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47091">
        <property type="entry name" value="ALPHA-PROTEIN KINASE 2-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47091:SF1">
        <property type="entry name" value="ALPHA-PROTEIN KINASE 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02816">
        <property type="entry name" value="Alpha_kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07679">
        <property type="entry name" value="I-set"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00811">
        <property type="entry name" value="Alpha_kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51158">
        <property type="entry name" value="ALPHA_KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="10" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="12" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="9" id="KW-0418">Kinase</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="7" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Ig-like" evidence="18">
        <location>
            <begin position="75"/>
            <end position="166"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="18">
        <location>
            <begin position="1275"/>
            <end position="1363"/>
        </location>
    </feature>
    <feature type="domain" description="Alpha-type protein kinase" evidence="19">
        <location>
            <begin position="1391"/>
            <end position="1622"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1"/>
            <end position="44"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="196"/>
            <end position="219"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="257"/>
            <end position="299"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="319"/>
            <end position="370"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="391"/>
            <end position="418"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="623"/>
            <end position="750"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="768"/>
            <end position="809"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="846"/>
            <end position="896"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="915"/>
            <end position="971"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1000"/>
            <end position="1024"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1181"/>
            <end position="1274"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1626"/>
            <end position="1700"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="17">
        <location>
            <begin position="22"/>
            <end position="31"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="17">
        <location>
            <begin position="196"/>
            <end position="218"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="266"/>
            <end position="278"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="17">
        <location>
            <begin position="288"/>
            <end position="299"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="344"/>
            <end position="370"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="636"/>
            <end position="645"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="17">
        <location>
            <begin position="646"/>
            <end position="660"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="17">
        <location>
            <begin position="698"/>
            <end position="709"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="17">
        <location>
            <begin position="873"/>
            <end position="884"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="942"/>
            <end position="951"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="17">
        <location>
            <begin position="961"/>
            <end position="971"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="17">
        <location>
            <begin position="1689"/>
            <end position="1700"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012513"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00047899"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00048679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00053245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00073276"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS51158"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000855.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1700" mass="186277" checksum="F2E22986E1AF43A0" modified="2019-05-08" version="1">MGSRRRPLSRSYSVNGRYSLYSPSSAPGPGAESEEWEGPGWSTRPDSRSYLLSIRPENRSTFCTIIAQLTEETQPQFETTLKSRAVSEDSDAKFTCIVTGYPQPEVTWYKDDEEMDRYCGLPKYEIFRHGNRHTLQLYKCREDDAAIYQASARNTKGIVSCSSVLEVGTMTEYKIHQRWFAKLKQKAEAKLHEIEQSRKRGKENMEVEKLRRTSPERFQRKRRLTGNLNLRSGVSLWDKEDVAKVHVVDPKSRLREDVAEPKEPSLNATAGFSNNFLTRPQDAEVTTNGDSSMESGEENGNSFFTYIYETVEEITTKPTTRESFAKKKKKKGEEPAAVMAAKQDVSSQENGLIESQNLSQSQRNAPSVTLPGSMQSVAAISQLVGDVPEVKELGTPSSQDAETGKPQPCSPPSRGDTYFPLAEMYLESGAKHVMEEKMPQTPEEAENRTQLQSLAVSINRQREILPPASEVFLVGAVAGEEQKGQPVERELEGNEAIGTKATKKESKLHGNLNKKRVDPPSGTLKLSSEMRHSLLMGNASGTSCISSCKNVQETDLLDSVQLNTKQPLHHATPSLNQVAPCLDESQLRQEAGGSLLQGSLEEQVVFKVPCEAQEAVRQVVSSTSDLKHSALEPTEDTNSQASQLKELTRRKPICMERETEQSEQPSSPRLRGTGMLKISQESSQRPLAGSPGLTQPSWEEHPPWKEEGAGHSLNFPGKETQQRPTPSEENEELATAGSALKKASRMPGGKALLKPGACWSGMGEGEKIAQPLTAETEPAGSWLGDSLSCTEVEESPIEPSHETSQQADGVVGGVSWVPKQAAPVSIQVSKEELHPIAIAEISESVEALGSASPSPVKEAPAGGKQSAAQPPGDVKREMETRKTNESAPRLAAMPHRERLPVSPAMELSAKLAPVSLTPESWEESEQRAVTPGKEPAAAAGRNSMTSTGTSELKTEAPLAPRAEERTPDGEVSLEHKVQHRNLVSSLKNYLLLLLKMSSSEPDKCKGPTKKDTETEEKESQPAGEEKALLEVGIAGLSPGTSRKILERVQNNQLFQSAESLHLSPRTSRRLTGMINQELVTSKEALGISPLACSTLPAREASEPEGSALLSVPSIVVGNVSLEGTRQADDGLSDLHLESSTKENSHDLVAALPCATPEELASGARRKIYLAKSKQLEEAEAAASESQVHLKRDSPTVSPRQSRKNPALLQAPVPTLSPPMDRHSPTLPRKMATLEVPKAYEEPSEEISGTPEGPEVTKQEGQAGESRKTNNPFKAPQLIRKIRAEQFSDASGNLKLWCQFFNILSDSKLTWYKDEFPVAEARRSSGDEGQAALAIVQASRKDCGVYKCTIENEYGTDSTDFLLSAEVLSGFVSREEIEVGEEIEMTPMVFAKGLADSGYWGDKFFGRVVSEDMEVGAGFLRKAWRARIIYGLEPVFESGRTCIIKVHNLIVFGTKNENTLIEKNYDITIQECKIQNLSREYCKIFAAEARAIPDFGAVPEIIPLYLIYRPANTVPYATIEEDLAGPFERYCVKERDGSLAVQSSSEIAQKCCAFQHWLYQWTNGNILVTDMEGVGWRITNMRIATKLKGYQGLKESCFPALLDQFAAAHRCNRYCEILGLKSLESLQQPAKPKGSKSPSLGRKAPSAQSSPQLQKKGLASPQGTRKGAVSPKASRKSTEAGEAQPVLKHKALESDRSSQLQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGY5</accession>
    <name>A0A452GGY5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">T-box domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 6">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="6">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GGY5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000939.1">
        <property type="protein sequence ID" value="ENSGAGP00000000830.1"/>
        <property type="gene ID" value="ENSGAGG00000000667.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000785">
        <property type="term" value="C:chromatin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009653">
        <property type="term" value="P:anatomical structure morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001708">
        <property type="term" value="P:cell fate specification"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045893">
        <property type="term" value="P:positive regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd20187">
        <property type="entry name" value="T-box_TBX1_10-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.820">
        <property type="entry name" value="Transcription factor, T-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008967">
        <property type="entry name" value="p53-like_TF_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046360">
        <property type="entry name" value="T-box_DNA-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036960">
        <property type="entry name" value="T-box_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001699">
        <property type="entry name" value="TF_T-box"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018186">
        <property type="entry name" value="TF_T-box_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11267">
        <property type="entry name" value="T-BOX PROTEIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11267:SF102">
        <property type="entry name" value="T-BOX TRANSCRIPTION FACTOR TBX10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00907">
        <property type="entry name" value="T-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00937">
        <property type="entry name" value="TBOX"/>
    </dbReference>
    <dbReference type="SMART" id="SM00425">
        <property type="entry name" value="TBOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49417">
        <property type="entry name" value="p53-like transcription factors"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01283">
        <property type="entry name" value="TBOX_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01264">
        <property type="entry name" value="TBOX_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50252">
        <property type="entry name" value="TBOX_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3 6" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="5 6" id="KW-0539">Nucleus</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0804">Transcription</keyword>
    <keyword evidence="2" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="T-box" evidence="7">
        <location>
            <begin position="75"/>
            <end position="244"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00201"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50252"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000830.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="332" mass="36353" checksum="85225C6D93EB5346" modified="2025-10-08" version="2">MPAFLTASLSVLTSDGGPFASGPEWGNPVGAPCLLGPPPACGRFGVPAGEGPAGLGDGVQPARKNQHVSNVSVQLEMRSLWEEFNSLGTEMIVTKAGRRMFPTFQVKLSGLDPLADYVLLMDFVPLDDKRYRYAFHSSSWLVAGRADPAAPGRVHFHPDSPAKGGQWMRQIVSFDKLKLTNNLLDDNGHIILNSMHRYQPRFHVVYVDPRRDSERFAHENFKSFTFVETQFMAVTAYQNHRVSSPPGAPRLGIIVPLPVSPALRSPPPAHHLRATAPPAAEGRLGPPLYKPLSYHSVYVRAKPRVAPYPLPASLHPFNIYGASPPALNYGQQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGY8</accession>
    <name>A0A452GGY8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Ig-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="9">Belongs to the immunoglobulin superfamily. TIM family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GGY8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000967.1">
        <property type="protein sequence ID" value="ENSGAGP00000000851.1"/>
        <property type="gene ID" value="ENSGAGG00000000714.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001786">
        <property type="term" value="F:phosphatidylserine binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043277">
        <property type="term" value="P:apoptotic cell clearance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060097">
        <property type="term" value="P:cytoskeletal rearrangement involved in phagocytosis, engulfment"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000774">
        <property type="entry name" value="Hepatitis A virus cellular receptor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013106">
        <property type="entry name" value="Ig_V-set"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46608">
        <property type="entry name" value="T-CELL IMMUNOGLOBULIN AND MUCIN DOMAIN-CONTAINING PROTEIN 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46608:SF3">
        <property type="entry name" value="T-CELL IMMUNOGLOBULIN AND MUCIN DOMAIN-CONTAINING PROTEIN 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07686">
        <property type="entry name" value="V-set"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="7" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="8" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="5 10" id="KW-0472">Membrane</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 11" id="KW-0732">Signal</keyword>
    <keyword evidence="2 10" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 10" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="11">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019500008" description="Ig-like domain-containing protein" evidence="11">
        <location>
            <begin position="23"/>
            <end position="246"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="178"/>
            <end position="200"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="12">
        <location>
            <begin position="3"/>
            <end position="130"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00038203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000851.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="246" mass="26846" checksum="99FE442A027EDF66" modified="2019-05-08" version="1">MAPCFILKRILMVLFTGLSVSGSPVRGVVGQNVTLPCKYRVNHQNDVTTMCWGRGSCPSSQCSQTILWTDGRRVTERQSSRYRLEGNLAQGDVSLTIVNAAEADGGMYCCRVEIPGWFNDQLNNLEVVIEKAAANASDTSSAITPQWPLVLSSEAPEDVSVTMLTSIHQQMEPESTRAGVHVGISIFVVLLVILVLALIISKRYFPNRQKLKTLASSVSFSNPEHGGLQSTLEAGDHAEENIYTMS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGY9</accession>
    <name>A0A452GGY9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Glutaryl-CoA dehydrogenase, mitochondrial</fullName>
            <ecNumber evidence="11">1.3.8.6</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>glutaryl-CoA + oxidized [electron-transfer flavoprotein] + 2 H(+) = (2E)-butenoyl-CoA + reduced [electron-transfer flavoprotein] + CO2</text>
            <dbReference type="Rhea" id="RHEA:13389"/>
            <dbReference type="Rhea" id="RHEA-COMP:10685"/>
            <dbReference type="Rhea" id="RHEA-COMP:10686"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:57332"/>
            <dbReference type="ChEBI" id="CHEBI:57378"/>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
            <dbReference type="ChEBI" id="CHEBI:58307"/>
            <dbReference type="EC" id="1.3.8.6"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 14">
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </cofactor>
    </comment>
    <comment type="pathway">
        <text evidence="9">Amino-acid metabolism; lysine degradation.</text>
    </comment>
    <comment type="pathway">
        <text evidence="10">Amino-acid metabolism; tryptophan metabolism.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Mitochondrion matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 14">Belongs to the acyl-CoA dehydrogenase family.</text>
    </comment>
    <dbReference type="EC" id="1.3.8.6" evidence="11"/>
    <dbReference type="AlphaFoldDB" id="A0A452GGY9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000979.1">
        <property type="protein sequence ID" value="ENSGAGP00000000862.1"/>
        <property type="gene ID" value="ENSGAGG00000000718.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005743">
        <property type="term" value="C:mitochondrial inner membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005759">
        <property type="term" value="C:mitochondrial matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000062">
        <property type="term" value="F:fatty-acyl-CoA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050660">
        <property type="term" value="F:flavin adenine dinucleotide binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004361">
        <property type="term" value="F:glutaryl-CoA dehydrogenase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033539">
        <property type="term" value="P:fatty acid beta-oxidation using acyl-CoA dehydrogenase"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046949">
        <property type="term" value="P:fatty-acyl-CoA biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd01151">
        <property type="entry name" value="GCD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.540.10:FF:000003">
        <property type="entry name" value="glutaryl-CoA dehydrogenase, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.140.10:FF:000006">
        <property type="entry name" value="Glutaryl-CoA dehydrogenase, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.110.10:FF:000008">
        <property type="entry name" value="Glutaryl-CoA dehydrogenase, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.540.10">
        <property type="entry name" value="Acyl-CoA dehydrogenase/oxidase, N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.110.10">
        <property type="entry name" value="Butyryl-CoA Dehydrogenase, subunit A, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.140.10">
        <property type="entry name" value="Butyryl-CoA Dehydrogenase, subunit A, domain 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006089">
        <property type="entry name" value="Acyl-CoA_DH_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006091">
        <property type="entry name" value="Acyl-CoA_Oxase/DH_mid-dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046373">
        <property type="entry name" value="Acyl-CoA_Oxase/DH_mid-dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036250">
        <property type="entry name" value="AcylCo_DH-like_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009075">
        <property type="entry name" value="AcylCo_DH/oxidase_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013786">
        <property type="entry name" value="AcylCoA_DH/ox_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037069">
        <property type="entry name" value="AcylCoA_DH/ox_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009100">
        <property type="entry name" value="AcylCoA_DH/oxidase_NM_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052033">
        <property type="entry name" value="Glutaryl-CoA_DH_mitochondrial"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR42807">
        <property type="entry name" value="GLUTARYL-COA DEHYDROGENASE, MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR42807:SF1">
        <property type="entry name" value="GLUTARYL-COA DEHYDROGENASE, MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00441">
        <property type="entry name" value="Acyl-CoA_dh_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02770">
        <property type="entry name" value="Acyl-CoA_dh_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02771">
        <property type="entry name" value="Acyl-CoA_dh_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47203">
        <property type="entry name" value="Acyl-CoA dehydrogenase C-terminal domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56645">
        <property type="entry name" value="Acyl-CoA dehydrogenase NM domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00073">
        <property type="entry name" value="ACYL_COA_DH_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 14" id="KW-0274">FAD</keyword>
    <keyword evidence="4 14" id="KW-0285">Flavoprotein</keyword>
    <keyword evidence="8" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="7 14" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0809">Transit peptide</keyword>
    <feature type="domain" description="Acyl-CoA dehydrogenase/oxidase N-terminal" evidence="17">
        <location>
            <begin position="53"/>
            <end position="164"/>
        </location>
    </feature>
    <feature type="domain" description="Acyl-CoA oxidase/dehydrogenase middle" evidence="16">
        <location>
            <begin position="168"/>
            <end position="261"/>
        </location>
    </feature>
    <feature type="domain" description="Acyl-CoA dehydrogenase/oxidase C-terminal" evidence="15">
        <location>
            <begin position="280"/>
            <end position="420"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001974"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004305"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00009347"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022630"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022827"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00037899"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00037927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00039033"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00039507"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00049493"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="RuleBase" id="RU362125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="Pfam" id="PF00441"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF02770"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF02771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000862.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="430" mass="47216" checksum="F3A2341322F62F94" modified="2019-05-08" version="1">MALRLAVTRLLRCSSCRVVGVGAIRPQGTIAAQKAPGAQFDWRDPLQLERQLTQEELMIRDSFRTYCQGKLMPRIVMANRNEVFHRDIVSELGELGVLGSTIKGYGCAGTTYVAYGLLAREIERVDSSYRSVLSVQSSLVMHPINAYGTEQQKQKYLPRLATGELLGCFGLTEPNHGSDPGGMETRARYHPASKTYTLNGSKTWITNSPLADLCVVWAVCEDGRIRGFLLERGMKGLSTPKIEGKFSLRASTTGMILMEDVEVPEENLLPHVSGLAGPFGCLNNARYGIAWGALGAAEFCLETARQYALDRIQFGVPLARNQLVQKKLADMLTEITIGLQACLQLGRLKDEDKAAPEMISMLKRNSCGKALDIARQARDMLGGNGIADEYHVIRHVMNLEAVNTYEGTHDIHALILGRAITGLQSFTVGK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGZ2</accession>
    <name>A0A452GGZ2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">SRCR domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="12">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GGZ2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000983.1">
        <property type="protein sequence ID" value="ENSGAGP00000000865.1"/>
        <property type="gene ID" value="ENSGAGG00000000435.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000012">
        <property type="entry name" value="CD163 molecule like 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000003">
        <property type="entry name" value="Deleted in malignant brain tumors 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000002">
        <property type="entry name" value="Scavenger receptor cysteine-rich type 1 protein M130"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000004">
        <property type="entry name" value="Scavenger receptor cysteine-rich type 1 protein M130"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.250.10">
        <property type="entry name" value="SRCR-like domain"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001190">
        <property type="entry name" value="SRCR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036772">
        <property type="entry name" value="SRCR-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19331">
        <property type="entry name" value="SCAVENGER RECEPTOR DOMAIN-CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19331:SF484">
        <property type="entry name" value="SRCR DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00530">
        <property type="entry name" value="SRCR"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00258">
        <property type="entry name" value="SPERACTRCPTR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00202">
        <property type="entry name" value="SR"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56487">
        <property type="entry name" value="SRCR-like"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00420">
        <property type="entry name" value="SRCR_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50287">
        <property type="entry name" value="SRCR_2"/>
        <property type="match status" value="6"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="10 12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9 13" id="KW-0472">Membrane</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="6 14" id="KW-0732">Signal</keyword>
    <keyword evidence="5 13" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 13" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="14">
        <location>
            <begin position="1"/>
            <end position="21"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019144370" description="SRCR domain-containing protein" evidence="14">
        <location>
            <begin position="22"/>
            <end position="714"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="13">
        <location>
            <begin position="678"/>
            <end position="701"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="15">
        <location>
            <begin position="22"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="15">
        <location>
            <begin position="127"/>
            <end position="224"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="15">
        <location>
            <begin position="224"/>
            <end position="324"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="15">
        <location>
            <begin position="343"/>
            <end position="444"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="15">
        <location>
            <begin position="449"/>
            <end position="549"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="15">
        <location>
            <begin position="565"/>
            <end position="665"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="47"/>
            <end position="111"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="60"/>
            <end position="121"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="91"/>
            <end position="101"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="196"/>
            <end position="206"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="249"/>
            <end position="313"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="262"/>
            <end position="323"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="293"/>
            <end position="303"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="369"/>
            <end position="433"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="382"/>
            <end position="443"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="413"/>
            <end position="423"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="473"/>
            <end position="537"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="517"/>
            <end position="527"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="590"/>
            <end position="654"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="603"/>
            <end position="664"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="634"/>
            <end position="644"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00196"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50287"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000865.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="714" mass="76763" checksum="7F200CD5318E5B15" modified="2019-05-08" version="1">WGPGSLLALTCSVSLFEGAEELRLADGGSPCAGRVEVKHQDQWGTVCSKGWDKADAGVVCKQLGCGAAVSAPRWGHFGPGSGPIWLDEVACVGTESALWHCRNRGWNVHNCYHEWDAGVTCSGHIELHLVGGDNACSGRVEVRHKDNWATICDAHFDLKAASVICNALQCGIAVSVPEGAHFRKGHGPIWTEEFQCVGNESHHVYCPRVSHGSQIVAYSRIGCFRLVNGSTACSGRVEIQVLGAWGTLCDSHWDLPDANVLCHQLYCGFAVSAPGEAHFGKGNGSVWTDTFHCKGNEPHLRHCPVTALGASQCSHDNDASVVCSGNVIQCIFYLLSPGLSESLRLLVNGTGRCAGRVEIYYNGSWGTVCDDFWDLLDSNVVCKQLGCGRAINATVSGHYGQGSGQIWLDDVNCSGNESDLWACPSRSWGQHNCRHKEDAGVLCSEFTDLRLVSNSYCAGRLEVFYNGTWGSVCANQMSGVTPEIVCKQLNCGDGGQIARDFEYGAGSGPTWLDHVTCSEQHSSLWQCPSEPWNRKSCDNRAEETHISCTGNSETTFSAFPDQEKLRVMGGEDGCSGRVEVWYQGSWGTVCDDSWDMADANVVCKLLGCGSAISALDKAAFGEGTGPIWVEKLNCRGTESSLWDCPAKPWGESNCDHKEDVAVNCSAPSRRLPTDSGRVTVPVIICIILGALLCLVLIILGAQVQSARAQRRGGS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGZ3</accession>
    <name>A0A452GGZ3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">protein-tyrosine-phosphatase</fullName>
            <ecNumber evidence="2">3.1.3.48</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endomembrane system</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="3.1.3.48" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452GGZ3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000973.1">
        <property type="protein sequence ID" value="ENSGAGP00000000856.1"/>
        <property type="gene ID" value="ENSGAGG00000000715.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030054">
        <property type="term" value="C:cell junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012505">
        <property type="term" value="C:endomembrane system"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019901">
        <property type="term" value="F:protein kinase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd14611">
        <property type="entry name" value="R-PTPc-R"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000020">
        <property type="entry name" value="Tyrosine-protein phosphatase non-receptor type 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000242">
        <property type="entry name" value="PTP_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR059011">
        <property type="entry name" value="PTPRR_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016130">
        <property type="entry name" value="Tyr_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003595">
        <property type="entry name" value="Tyr_Pase_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008356">
        <property type="entry name" value="Tyr_Pase_KIM-con"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016334">
        <property type="entry name" value="Tyr_Pase_rcpt_R/non-rcpt_5"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46198">
        <property type="entry name" value="PROTEIN-TYROSINE-PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46198:SF2">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE R"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26155">
        <property type="entry name" value="PTPRR_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00102">
        <property type="entry name" value="Y_phosphatase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF001997">
        <property type="entry name" value="PTPRR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01778">
        <property type="entry name" value="KIMPTPASE"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00700">
        <property type="entry name" value="PRTYPHPHTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00194">
        <property type="entry name" value="PTPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00404">
        <property type="entry name" value="PTPc_motif"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00383">
        <property type="entry name" value="TYR_PHOSPHATASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50055">
        <property type="entry name" value="TYR_PHOSPHATASE_PTP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="8" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="6" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7" id="KW-1133">Transmembrane helix</keyword>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="11">
        <location>
            <begin position="344"/>
            <end position="598"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="12">
        <location>
            <begin position="513"/>
            <end position="589"/>
        </location>
    </feature>
    <feature type="active site" description="Phosphocysteine intermediate" evidence="9">
        <location>
            <position position="539"/>
        </location>
    </feature>
    <feature type="binding site" evidence="10">
        <location>
            <position position="505"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="10">
        <location>
            <begin position="539"/>
            <end position="545"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="10">
        <location>
            <position position="583"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004308"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00013064"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PIRSR" id="PIRSR001997-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PIRSR" id="PIRSR608356-51"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000856.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="608" mass="68282" checksum="53336B819E57E13A" modified="2019-05-08" version="1">MFSLLSTGCFSGDSDHFLAIHQRKSGKPVFIYHHVQSVEKSLDSDSQKIYKHNFHSSSHAKISNLHPAIIVKSAFPRRAYDPSLNLLAMTAQEPEVENLPIPATNVIVVALQMDVNKLNITLLRIFRQGVAAALGLLPQQIHINRLIGKKNCVEMFVSPTNRKPGISDALPSEEVLRSLNINVLHQSLSQFGITEVSPEVLYRLKEKIQFSLQQDKEKKQEIHLSPIPLQTTRSEYKTANRMVQPEQAPKVVNVIVDPQGQCVPEIKPPLSASPSPFKMKPVGLQERRGSNVSLTLDMSSLGNVEPFVAVPTPREKVAMEYLQSASRVLTRQQLRDTVACSHLLQTEFMEIPMNFVDPKEIAIPSHGTKNRYKTILPNPLSRVCLKPKNPADSLSTYINANYIRGYGGKEKSFIATQGPMINTVNDFWQMVWQEDSPVIVMITKLKEKNEKCVLYWPEKRGIYGKVEVLIKDVQECENYTVRNLTLKQGSQSQNVKHYWYTSWPDHKTPYSAQPLLQLMLDVEEDRMASAGRGPVIVHCSAGIGRTGCFIATAIGCHQLKEEGVVDALSIVCQLRVDRGGMIQTSEQYEFVHHALSLYESRLSAEAVQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGZ8</accession>
    <name>A0A452GGZ8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Ig-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="9">Belongs to the immunoglobulin superfamily. TIM family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GGZ8"/>
    <dbReference type="STRING" id="38772.ENSGAGP00000000849"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000964.1">
        <property type="protein sequence ID" value="ENSGAGP00000000849.1"/>
        <property type="gene ID" value="ENSGAGG00000000714.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001786">
        <property type="term" value="F:phosphatidylserine binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043277">
        <property type="term" value="P:apoptotic cell clearance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060097">
        <property type="term" value="P:cytoskeletal rearrangement involved in phagocytosis, engulfment"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000774">
        <property type="entry name" value="Hepatitis A virus cellular receptor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013106">
        <property type="entry name" value="Ig_V-set"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46608">
        <property type="entry name" value="T-CELL IMMUNOGLOBULIN AND MUCIN DOMAIN-CONTAINING PROTEIN 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46608:SF3">
        <property type="entry name" value="T-CELL IMMUNOGLOBULIN AND MUCIN DOMAIN-CONTAINING PROTEIN 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07686">
        <property type="entry name" value="V-set"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="7" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="8" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="5 10" id="KW-0472">Membrane</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 11" id="KW-0732">Signal</keyword>
    <keyword evidence="2 10" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 10" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="11">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019032931" description="Ig-like domain-containing protein" evidence="11">
        <location>
            <begin position="23"/>
            <end position="263"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="195"/>
            <end position="217"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="12">
        <location>
            <begin position="3"/>
            <end position="130"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00038203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000849.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="263" mass="28695" checksum="D64497778E1E715B" modified="2019-05-08" version="1">MAPCFILKRILMVLFTGLSVSGSPVRGVVGQNVTLPCKYRVNHQNDVTTMCWGRGSCPSSQCSQTILWTDGRRVTERQSSRYRLEGNLAQGDVSLTIVNAAEADGGMYCCRVEIPGWFNDQLNNLEVVIEKARTSTADPHPYTPKHTSAAANASDTSSAITPQWPLVLSSEAPEDVSVTMLTSIHQQMEPESTRAGVHVGISIFVVLLVILVLALIISKRYFPNRQKLKTLASSVSFSNPEHGGLQSTLEAGDHAEENIYTMS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GH52</accession>
    <name>A0A452GH52_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Folate receptor-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the folate receptor family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GH52"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001019.1">
        <property type="protein sequence ID" value="ENSGAGP00000000899.1"/>
        <property type="gene ID" value="ENSGAGG00000000517.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009897">
        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0038023">
        <property type="term" value="F:signaling receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004269">
        <property type="entry name" value="Folate_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018143">
        <property type="entry name" value="Folate_rcpt-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10517">
        <property type="entry name" value="FOLATE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10517:SF14">
        <property type="entry name" value="FOLATE RECEPTOR 1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03024">
        <property type="entry name" value="Folate_rec"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0732">Signal</keyword>
    <feature type="domain" description="Folate receptor-like" evidence="4">
        <location>
            <begin position="11"/>
            <end position="159"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007932"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="Pfam" id="PF03024"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000899.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="203" mass="23380" checksum="361904A4DF86D15A" modified="2019-05-08" version="1">MLLHPQHPLYNACCTAETSTGAHQDQSYLYNFNWNHCGVMPEMCKRHFIQDTCLYECSPNLGPWIDQADTSWRRERILNVPLCKEDCELWWEACKDAVTCKENWHKGWNWTSGTNQCPHSSTCQLFKYIFPRPADLCEKIWSNSYKYTTEHWGSGRCIQMWFDPANGNPNVAVAKYYAQNGRDASPAPQAVLLLLPAALLSLL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GH53</accession>
    <name>A0A452GH53_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="14">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GH53"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001057.1">
        <property type="protein sequence ID" value="ENSGAGP00000000932.1"/>
        <property type="gene ID" value="ENSGAGG00000000708.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd07765">
        <property type="entry name" value="KRAB_A-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000557">
        <property type="entry name" value="zinc finger and SCAN domain-containing protein 29"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000759">
        <property type="entry name" value="zinc finger protein 16"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002343">
        <property type="entry name" value="Zinc finger protein 33A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000016">
        <property type="entry name" value="zinc finger protein 37 homolog"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000238">
        <property type="entry name" value="Zinc finger protein 485"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000367">
        <property type="entry name" value="Zinc finger protein 572"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000038">
        <property type="entry name" value="Zinc finger protein 624"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000953">
        <property type="entry name" value="Zinc finger protein 691"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000176">
        <property type="entry name" value="zinc finger protein 70"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001954">
        <property type="entry name" value="Zinc finger protein 787"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001909">
        <property type="entry name" value="KRAB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036051">
        <property type="entry name" value="KRAB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050527">
        <property type="entry name" value="Snail/Krueppel_Znf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24388">
        <property type="entry name" value="ZINC FINGER PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24388:SF51">
        <property type="entry name" value="ZINC FINGER PROTEIN 281-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01352">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="SMART" id="SM00349">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109640">
        <property type="entry name" value="KRAB domain (Kruppel-associated box)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50805">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="11"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="9" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 11" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="KRAB" evidence="13">
        <location>
            <begin position="8"/>
            <end position="83"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="115"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="143"/>
            <end position="170"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="171"/>
            <end position="198"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="199"/>
            <end position="226"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="227"/>
            <end position="254"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="257"/>
            <end position="284"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="285"/>
            <end position="312"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="315"/>
            <end position="337"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="338"/>
            <end position="365"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="366"/>
            <end position="393"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="394"/>
            <end position="421"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000932.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="426" mass="49225" checksum="7D309D5E94FAD573" modified="2019-05-08" version="1">MLLQQDPVTFEEVAVYFTREEWALLDPVQRALYRDVMQENYETVTSLGKESCPLDDERREAHGMLSGRSKENVSRSCALPEKPKACETLHRPEGNVSSLSDLITSGRINLEETRYRCHECGKSFSWSSTLVRHRRIHTGETPYTCTECGKSFSQSSNLITHQRIHTGEKPYTCTECGKSFNWSSTLNTHRRIHTGETPYTCTECGKSFSWRSDLVRHLRIHTGETPYTCTECGRRFSWRSHLIGHQRIHTGETEKPYTCSECGKSFSCSSDLIRHQRIHTGETPYTCTECGRSFNRSSHLIGHQRIHTGEKPYTCCERFSWRSHLIGHQRIHTGETPYTCTECGKSFSQCSNLITHQRIHTGETPYTCTECGKSFNRSSTLNTHRRIHTREMPYTCSECGQSFSQRSRLIRHQKMHRRENCNKSLD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GH79</accession>
    <name>A0A452GH79_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Phenylalanine--tRNA ligase alpha subunit</fullName>
            <ecNumber evidence="3">6.1.1.20</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="13">Phenylalanyl-tRNA synthetase alpha subunit</fullName>
        </alternativeName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the class-II aminoacyl-tRNA synthetase family. Phe-tRNA synthetase alpha subunit type 2 subfamily.</text>
    </comment>
    <dbReference type="EC" id="6.1.1.20" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452GH79"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001047.1">
        <property type="protein sequence ID" value="ENSGAGP00000000924.1"/>
        <property type="gene ID" value="ENSGAGG00000000753.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009328">
        <property type="term" value="C:phenylalanine-tRNA ligase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004826">
        <property type="term" value="F:phenylalanine-tRNA ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000049">
        <property type="term" value="F:tRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006432">
        <property type="term" value="P:phenylalanyl-tRNA aminoacylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00496">
        <property type="entry name" value="PheRS_alpha_core"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.2320:FF:000001">
        <property type="entry name" value="phenylalanine--tRNA ligase alpha subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.2330:FF:000001">
        <property type="entry name" value="phenylalanine--tRNA ligase alpha subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.930.10:FF:000036">
        <property type="entry name" value="phenylalanine--tRNA ligase alpha subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.2320">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.2330">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1370.240">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.930.10">
        <property type="entry name" value="Bira Bifunctional Protein, Domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006195">
        <property type="entry name" value="aa-tRNA-synth_II"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045864">
        <property type="entry name" value="aa-tRNA-synth_II/BPL/LPL"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004529">
        <property type="entry name" value="Phe-tRNA-synth_IIc_asu"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002319">
        <property type="entry name" value="Phenylalanyl-tRNA_Synthase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040724">
        <property type="entry name" value="PheRS_DBD1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040586">
        <property type="entry name" value="PheRS_DBD2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040725">
        <property type="entry name" value="PheRS_DBD3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036390">
        <property type="entry name" value="WH_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00468">
        <property type="entry name" value="pheS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF003210">
        <property type="entry name" value="PRK04172.1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11538:SF40">
        <property type="entry name" value="PHENYLALANINE--TRNA LIGASE ALPHA SUBUNIT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11538">
        <property type="entry name" value="PHENYLALANYL-TRNA SYNTHETASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18552">
        <property type="entry name" value="PheRS_DBD1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18554">
        <property type="entry name" value="PheRS_DBD2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18553">
        <property type="entry name" value="PheRS_DBD3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01409">
        <property type="entry name" value="tRNA-synt_2d"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55681">
        <property type="entry name" value="Class II aaRS and biotin synthetases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46785">
        <property type="entry name" value="Winged helix' DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50862">
        <property type="entry name" value="AA_TRNA_LIGASE_II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12" id="KW-0030">Aminoacyl-tRNA synthetase</keyword>
    <keyword evidence="9" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0436">Ligase</keyword>
    <keyword evidence="10" id="KW-0460">Magnesium</keyword>
    <keyword evidence="7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="11" id="KW-0648">Protein biosynthesis</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Aminoacyl-transfer RNA synthetases class-II family profile" evidence="14">
        <location>
            <begin position="236"/>
            <end position="498"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006703"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012814"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00015409"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022598"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022917"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023146"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00030612"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50862"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000924.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="504" mass="57037" checksum="8F8F2936FF960DF3" modified="2019-05-08" version="1">MSPAMATDGPALAELLLQRLERAGGGGLGSLEAAAALGLEHQQLVGAVKSLQALGEVIEAEQRSSKRWELTPEGKEIVQEGSHEARVFDSIPAEGLPQSELMKLPSGKVGFSKAMSNKWVRLDKNADGGPRVFRAVESVQDLVREKLQLVQRGDAESLEEKDKNELKKRKLLMEVNIKTYWIRKGSAFSTTVTKQETDLTPEMLASGSWRDLKFKAYNFEALGIMPESGHLHPLLKVRTQFRQIFLEMGFTEMPTNNFIESSFWNFDALFQPQQHPARDQHDTFFLQDPAEATDFPTDYLKRVKKIHSQGGYGSQGYKYDWKLEEARKNILRTHTTAVSARMLYRLAQQEKFTPAKYFSIDRVFRNETLDATHLAEFHQIEGVVADYGLTLGDLMGTLREFFTKLGISQLRFKPAYNPYTEPSMEVFSYHRGLQKWVEVGNSGIFRPEMLLPMGLPEGVSVIAWGLSLERPTMIKYGINNIRELVGHKVNLQMVYDSPLCRLDA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GH81</accession>
    <name>A0A452GH81_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">2',3'-cyclic-nucleotide 3'-phosphodiesterase</fullName>
            <ecNumber evidence="6">3.1.4.37</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">Catalyzes the formation of 2'-nucleotide products from 2',3'-cyclic substrates. May participate in RNA metabolism in the myelinating cell, CNP is the third most abundant protein in central nervous system myelin.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>a nucleoside 2',3'-cyclic phosphate + H2O = a nucleoside 2'-phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:14489"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:66954"/>
            <dbReference type="ChEBI" id="CHEBI:78552"/>
            <dbReference type="EC" id="3.1.4.37"/>
        </reaction>
    </comment>
    <comment type="subunit">
        <text evidence="5">Exists as monomers and homodimers.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Melanosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Membrane</location>
            <topology evidence="3">Lipid-anchor</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the 2H phosphoesterase superfamily. CNPase family.</text>
    </comment>
    <dbReference type="EC" id="3.1.4.37" evidence="6"/>
    <dbReference type="AlphaFoldDB" id="A0A452GH81"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001046.1">
        <property type="protein sequence ID" value="ENSGAGP00000000923.1"/>
        <property type="gene ID" value="ENSGAGG00000000759.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042470">
        <property type="term" value="C:melanosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004113">
        <property type="term" value="F:2',3'-cyclic-nucleotide 3'-phosphodiesterase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009214">
        <property type="term" value="P:cyclic nucleotide catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000795">
        <property type="entry name" value="Tetratricopeptide repeat protein 25"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1740.10:FF:000001">
        <property type="entry name" value="Tetratricopeptide repeat protein 25"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1740.10">
        <property type="entry name" value="2',3'-cyclic nucleotide 3'-phosphodiesterase superfamily"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008431">
        <property type="entry name" value="CNPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047325">
        <property type="entry name" value="CNPase_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009097">
        <property type="entry name" value="Cyclic_Pdiesterase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10156">
        <property type="entry name" value="2',3'-CYCLIC-NUCLEOTIDE 3'-PHOSPHODIESTERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10156:SF0">
        <property type="entry name" value="2',3'-CYCLIC-NUCLEOTIDE 3'-PHOSPHODIESTERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13671">
        <property type="entry name" value="AAA_33"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05881">
        <property type="entry name" value="CNPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000970">
        <property type="entry name" value="CNPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55144">
        <property type="entry name" value="LigT-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="13" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="12" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-0488">Methylation</keyword>
    <keyword evidence="9" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="14" id="KW-0636">Prenylation</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="11" id="KW-0694">RNA-binding</keyword>
    <feature type="domain" description="Cyclic nucleotide phosphodiesterase catalytic" evidence="19">
        <location>
            <begin position="213"/>
            <end position="448"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1"/>
            <end position="21"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="18">
        <location>
            <begin position="12"/>
            <end position="21"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="16">
        <location>
            <position position="278"/>
        </location>
    </feature>
    <feature type="active site" description="Proton donor" evidence="16">
        <location>
            <position position="357"/>
        </location>
    </feature>
    <feature type="binding site" evidence="17">
        <location>
            <position position="280"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="17">
        <location>
            <position position="359"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000610"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004223"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004635"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00008662"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00011781"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00012317"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00014478"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023289"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00045937"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PIRSR" id="PIRSR000970-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="PIRSR" id="PIRSR000970-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="Pfam" id="PF05881"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000923.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="449" mass="50049" checksum="D76F75BE107C7159" modified="2019-05-08" version="1">MKIAAGGPVTQPRPLRSAPAAPAGAALAMNRGFTRKSHTFLPKIFRKMSTQSSKERLENLQFPFLDDDTTISTVKESKTLFIIRGLPGSGKSTLAQAIQDQYKDACKVISADTYKITPAISSSIPEDYSKLDEDLVDYCKRDISVVVVDDTHHERERLDQLFDIADKYQFKVLFVEPKTPWKMDCLQLKDKNQWKLSVEDLKKMKPSLEKDFLPLYYGWFLSKKSSENLRKTGQAFLDELGSLKVFKKESKYFLPAEDPKVKLDLTSYFAKRPPSVLHCTTKFTDFGKAMGADEYAQQEVVKASYGKAFTLTISALFLTTKTAGARVELSEQELLLWPGDVDKILPTDNLPKGSRAHITLGCASGIEPVQTGIDLLEFVKLEKAGTKGDEIEEIGGGKLQYFGNGMWMLLLSKKIEVKAIFSGYYGKGKLVPTQGSNKQGSAFNPCTII</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GH82</accession>
    <name>A0A452GH82_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Protein JTB</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">Required for normal cytokinesis during mitosis. Plays a role in the regulation of cell proliferation. May be a component of the chromosomal passenger complex (CPC), a complex that acts as a key regulator of mitosis. The CPC complex has essential functions at the centromere in ensuring correct chromosome alignment and segregation and is required for chromatin-induced microtubule stabilization and spindle assembly. Increases AURKB activity. Inhibits apoptosis induced by TGFB1. Overexpression induces swelling of mitochondria and reduces mitochondrial membrane potential.</text>
    </comment>
    <comment type="subunit">
        <text evidence="17">Interacts with AURKA, AURKB, BIRC5 and INCENP. May be a component of the CPC at least composed of BIRC5/survivin, CDCA8/borealin, INCENP and AURKB/Aurora-B.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Cytoskeleton</location>
            <location evidence="3">Microtubule organizing center</location>
            <location evidence="3">Centrosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
            <location evidence="2">Spindle</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Membrane</location>
            <topology evidence="4">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Mitochondrion</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="16">Belongs to the JTB family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GH82"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001066.1">
        <property type="protein sequence ID" value="ENSGAGP00000000941.1"/>
        <property type="gene ID" value="ENSGAGG00000000752.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030496">
        <property type="term" value="C:midbody"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005819">
        <property type="term" value="C:spindle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000281">
        <property type="term" value="P:mitotic cytokinesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.720.220:FF:000001">
        <property type="entry name" value="Jumping translocation breakpoint"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.720.220">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008657">
        <property type="entry name" value="JTB"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13041">
        <property type="entry name" value="JTB PROTEIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13041:SF3">
        <property type="entry name" value="PROTEIN JTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05439">
        <property type="entry name" value="JTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="14" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="6" id="KW-0132">Cell division</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="13" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="12" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="9" id="KW-0498">Mitosis</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0732">Signal</keyword>
    <keyword evidence="7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="10" id="KW-1133">Transmembrane helix</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004173"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004186"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022776"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00058368"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00060886"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00063184"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00068227"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000941.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="136" mass="15190" checksum="4FAC3FF3E2FF4322" modified="2019-05-08" version="1">QCPWAGRRGLSQDLGLRLDRALNLSPLFSVSPLVATPCWQAEDFVVVQDCAPCTIFQAKTMPECSTTGFIEQVSCAVSKKEEYKSCRSVVTEAHSFWKFEGAMICVAVVFALLVMCRQRVLDRKALEKVLKQIESI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GH85</accession>
    <name>A0A452GH85_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">SRCR domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="caution">
        <text evidence="5">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GH85"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001054.1">
        <property type="protein sequence ID" value="ENSGAGP00000000929.1"/>
        <property type="gene ID" value="ENSGAGG00000000435.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000012">
        <property type="entry name" value="CD163 molecule like 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000006">
        <property type="entry name" value="neurotrypsin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000002">
        <property type="entry name" value="Scavenger receptor cysteine-rich type 1 protein M130"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.250.10">
        <property type="entry name" value="SRCR-like domain"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001190">
        <property type="entry name" value="SRCR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036772">
        <property type="entry name" value="SRCR-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19331:SF465">
        <property type="entry name" value="EGG PEPTIDE SPERACT RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19331">
        <property type="entry name" value="SCAVENGER RECEPTOR DOMAIN-CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00530">
        <property type="entry name" value="SRCR"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00258">
        <property type="entry name" value="SPERACTRCPTR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00202">
        <property type="entry name" value="SR"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56487">
        <property type="entry name" value="SRCR-like"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00420">
        <property type="entry name" value="SRCR_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50287">
        <property type="entry name" value="SRCR_2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3 5" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="4" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <keyword evidence="1" id="KW-0732">Signal</keyword>
    <feature type="domain" description="SRCR" evidence="6">
        <location>
            <begin position="30"/>
            <end position="129"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="6">
        <location>
            <begin position="179"/>
            <end position="279"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="6">
        <location>
            <begin position="281"/>
            <end position="381"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="6">
        <location>
            <begin position="387"/>
            <end position="487"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="6">
        <location>
            <begin position="492"/>
            <end position="592"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="6">
        <location>
            <begin position="597"/>
            <end position="697"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="6">
        <location>
            <begin position="711"/>
            <end position="811"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="54"/>
            <end position="118"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="67"/>
            <end position="128"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="98"/>
            <end position="108"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="204"/>
            <end position="268"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="217"/>
            <end position="278"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="248"/>
            <end position="258"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="306"/>
            <end position="370"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="319"/>
            <end position="380"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="350"/>
            <end position="360"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="425"/>
            <end position="486"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="456"/>
            <end position="466"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
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            <end position="581"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="530"/>
            <end position="591"/>
        </location>
    </feature>
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        <location>
            <begin position="561"/>
            <end position="571"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="621"/>
            <end position="685"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="665"/>
            <end position="675"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="736"/>
            <end position="800"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="749"/>
            <end position="810"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="780"/>
            <end position="790"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00196"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50287"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000929.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="813" mass="87557" checksum="F23C3B657595CF12" modified="2019-05-08" version="1">ISFTFLSISPSLLPPPNTSFCPPAPETFPYLLSTGSPCAGRVEVKHQDQWGTVCSKGWDKADAGVVCKQLGCGAAVSAPRWGHFGPGSGPIWLDEVACVGTESALWHCRNRGWNVHNCYHEWDAGVTCSGKNQSPLGMGGARDNGERKGEVMGHGAVGRGMGWEREGDTGMGAGRHIELHLVGGDNACSGRVEVRHKDNWATICDAHFDLKAASVICNALQCGIAVSVPEGAHFRKGHGPIWTEEFQCVGNESHHVYCPRVSHGSQMCSHANDASVICSRFRLVNGSTACSGRVEIQVLGAWGTLCDSHWDLPDANVLCHQLYCGFAVSAPGEAHFGKGNGSVWTDTFHCKGNEPHLRHCPVTALGASQCSHDNDASVVCSGLSESLRLLNGESWCDGRVEISLRGVWSRVLDDQWDMNDARVVCRQLQCGVAEKAYNPPKSERGTGPVGLRRVQCAGNETRLTLCDNSTSETSQAGITEDVGVVCSGSKQIRLVNGTGRCAGRVEIYYNGSWGTVCDDFWDLLDSNVVCKQLGCGRAINATVSGHYGQGSGQIWLDDVNCSGNESDLWACPSRSWGQHNCRHKEDAGVLCSEFTDLRLVSNSYCAGRLEVFYNGTWGSVCANQMSGVTPEIVCKQLNCGDGGQIARDFEYGAGSGPTWLDHVTCSEQHSSLWQCPSEPWNRKSCDNRAEETHISCTVTCEVSAFPDQEKLRVMGGEDGCSGRVEVWYQGSWGTVCDDSWDMADANVVCKLLGCGSAISALDKAAFGEGTGPIWVEKLNCRGTESSLWDCPAKPWGESNCDHKEDVAVNCSGE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GH88</accession>
    <name>A0A452GH88_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="5">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
            <location evidence="1">Nucleolus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the SLX9 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GH88"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001091.1">
        <property type="protein sequence ID" value="ENSGAGP00000000966.1"/>
        <property type="gene ID" value="ENSGAGG00000000641.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030686">
        <property type="term" value="C:90S preribosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005730">
        <property type="term" value="C:nucleolus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030688">
        <property type="term" value="C:preribosome, small subunit precursor"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000462">
        <property type="term" value="P:maturation of SSU-rRNA from tricistronic rRNA transcript (SSU-rRNA, 5.8S rRNA, LSU-rRNA)"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
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        <property type="entry name" value="Slx9-like"/>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31109:SF2">
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    </dbReference>
    <dbReference type="Pfam" id="PF15341">
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        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0539">Nucleus</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="coiled-coil region" evidence="4">
        <location>
            <begin position="71"/>
            <end position="98"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004604"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00011022"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000966.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="185" mass="20602" checksum="EBF2CBB1C3CFFFEF" modified="2019-05-08" version="1">GHVCLILSLAQDWAFLSSDVFAGMKIDPKTLVKKLDLDSRSVVSAQTGKGCFTAVCSYFSGVEEKTLLPKREKMKLRKERWLQKIEGLKLARQKQKAEAKRKATPVVGDMQPLLEALPELSELTTASKARKQPKSLIQGSLAFLREEEVARFHEVIANPVYKANPLAAISEHLCKRLKEEAGNTI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GH92</accession>
    <name>A0A452GH92_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">SRCR domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="12">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GH92"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001058.1">
        <property type="protein sequence ID" value="ENSGAGP00000000933.1"/>
        <property type="gene ID" value="ENSGAGG00000000435.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000012">
        <property type="entry name" value="CD163 molecule like 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000006">
        <property type="entry name" value="neurotrypsin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000002">
        <property type="entry name" value="Scavenger receptor cysteine-rich type 1 protein M130"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000004">
        <property type="entry name" value="Scavenger receptor cysteine-rich type 1 protein M130"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.250.10">
        <property type="entry name" value="SRCR-like domain"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001190">
        <property type="entry name" value="SRCR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036772">
        <property type="entry name" value="SRCR-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19331:SF468">
        <property type="entry name" value="SCAVENGER RECEPTOR CYSTEINE-RICH TYPE 1 PROTEIN M160"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19331">
        <property type="entry name" value="SCAVENGER RECEPTOR DOMAIN-CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00530">
        <property type="entry name" value="SRCR"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00258">
        <property type="entry name" value="SPERACTRCPTR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00202">
        <property type="entry name" value="SR"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56487">
        <property type="entry name" value="SRCR-like"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00420">
        <property type="entry name" value="SRCR_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50287">
        <property type="entry name" value="SRCR_2"/>
        <property type="match status" value="6"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="10 12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9 14" id="KW-0472">Membrane</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="5 14" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 14" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="603"/>
            <end position="624"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="15">
        <location>
            <begin position="1"/>
            <end position="86"/>
        </location>
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            <begin position="88"/>
            <end position="188"/>
        </location>
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            <begin position="194"/>
            <end position="238"/>
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            <begin position="282"/>
            <end position="382"/>
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            <begin position="384"/>
            <end position="484"/>
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    <feature type="domain" description="SRCR" evidence="15">
        <location>
            <begin position="500"/>
            <end position="600"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="633"/>
            <end position="707"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="11"/>
            <end position="75"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="24"/>
            <end position="85"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="55"/>
            <end position="65"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="113"/>
            <end position="177"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="126"/>
            <end position="187"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="157"/>
            <end position="167"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="307"/>
            <end position="371"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="320"/>
            <end position="381"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="351"/>
            <end position="361"/>
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    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="408"/>
            <end position="472"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="452"/>
            <end position="462"/>
        </location>
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    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="525"/>
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    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="538"/>
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    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="569"/>
            <end position="579"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
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    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
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    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
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    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00196"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50287"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000933.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="759" mass="81709" checksum="CEBF0560BBC2C01D" modified="2019-05-08" version="1">VRHKDNWATICDAHFDLKAASVICNALQCGIAVSVPEGAHFRKGHGPIWTEEFQCVGNESHHVYCPRVSHGSQMCSHANDASVICSRFRLVNGSTACSGRVEIQVLGAWGTLCDSHWDLPDANVLCHQLYCGFAVSAPGEAHFGKGNGSVWTDTFHCKGNEPHLRHCPVTALGASQCSHDNDASVVCSGLSESLRLLNGESWCDGRVEISLRGVWSRVLDDQWDMNDARVVCRQLQCGVAEKAYNPPKSERGTGNETRLTLCDNSTSETSQLGSIPAGSKQIRLVNGTGRCAGRVEIYYNGSWGTVCDDFWDLLDSNVVCKQLGCGRAINATVSGHYGQGSGQIWLDDVNCSGNESDLWACPSRSWGQHNCRHKEDAGVLCSDLRLVSNSYCAGRLEVFYNGTWGSVCANQMSGVTPEIVCKQLNCGDGGQIARDFEYGAGSGPTWLDHVTCSEQHSSLWQCPSEPWNRKSCDNRAEETHISCTGNSETTFSAFPDQEKLRVMGGEDGCSGRVEVWYQGSWGTVCDDSWDMADANVVCKLLGCGSAISALDKAAFGEGTGPIWVEKLNCRGTESSLWDCPAKPWGESNCDHKEDVAVNCSGEVTVPVIICIILGALLCLVLIILGAQVQSARAQPSSAERAPRNGDTLAPDLGDHSKVGGQPPRNPGPSLVGVTRTLQTGESSRAGRHLRKRNGSKDSDISPIPLHQGSAEVRKVPHDRGTIPPLTCIPTVIHSKNAQSGLKPPAITYLWAGSICSTSF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GH98</accession>
    <name>A0A452GH98_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="14">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">May be involved in transcriptional regulation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GH98"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001106.1">
        <property type="protein sequence ID" value="ENSGAGP00000000979.1"/>
        <property type="gene ID" value="ENSGAGG00000000708.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031981">
        <property type="term" value="C:nuclear lumen"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd07765">
        <property type="entry name" value="KRAB_A-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000557">
        <property type="entry name" value="zinc finger and SCAN domain-containing protein 29"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000002">
        <property type="entry name" value="Zinc finger protein 1 homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000759">
        <property type="entry name" value="zinc finger protein 16"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000295">
        <property type="entry name" value="zinc finger protein 19"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000016">
        <property type="entry name" value="zinc finger protein 37 homolog"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002090">
        <property type="entry name" value="Zinc finger protein 473"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000362">
        <property type="entry name" value="Zinc finger protein 606"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002134">
        <property type="entry name" value="Zinc finger protein 616"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000912">
        <property type="entry name" value="Zinc finger protein 660"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000176">
        <property type="entry name" value="zinc finger protein 70"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001909">
        <property type="entry name" value="KRAB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036051">
        <property type="entry name" value="KRAB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23226">
        <property type="entry name" value="ZINC FINGER AND SCAN DOMAIN-CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23226:SF377">
        <property type="entry name" value="ZINC FINGER AND SCAN DOMAIN-CONTAINING PROTEIN 20"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01352">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="SMART" id="SM00349">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109640">
        <property type="entry name" value="KRAB domain (Kruppel-associated box)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50805">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="14"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="9" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0862">Zinc</keyword>
    <keyword evidence="6 11" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="KRAB" evidence="13">
        <location>
            <begin position="13"/>
            <end position="87"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="79"/>
            <end position="106"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="107"/>
            <end position="134"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="135"/>
            <end position="162"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="163"/>
            <end position="190"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="191"/>
            <end position="218"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="219"/>
            <end position="246"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="247"/>
            <end position="274"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="275"/>
            <end position="302"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="303"/>
            <end position="330"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="331"/>
            <end position="358"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="359"/>
            <end position="386"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="387"/>
            <end position="414"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="415"/>
            <end position="442"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="443"/>
            <end position="470"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00003767"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000979.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="475" mass="54454" checksum="C6FFCEB2B2219AA3" modified="2019-05-08" version="1">MNSGVVSGAAGPEVWTLVAVYFTREEWALLDPVQRALYRDVMQENYETVTSLGKEQENDCHPGEAHGMLSGRSKENKPYTCTECGKSFNWSSTLNTHRRIHTGETPYTCTECGKSFSWRSDLVRHLRIHTGETPYTCTECGRRFSWRSHLIGHQRIHTGETPYTCTECGKSFSQCSNLITHQRIHTGEKPYSCSECGKSFSRSSHLIRHQIIHTGETPYTCTECGKSFNRSSTLNTHRRIHTGKLPYTCTECEKSFSRSSDLIGHQRIHTREKPYTCSECGKSFSCSSDLIRHQRIHTGETPYTCTECGRSFNRSSHLIGHQRIHTGEKPYTCCECEKSFSQSSHLIGHQRIHTGETPYTCTECGRRFSWRSHLIGHQRIHTGETPYTCTECGKSFSQCSNLITHQRIHTGETPYTCTECGKSFNRSSTLNTHRRIHTREMPYTCSECGQSFSQRSRLIRHQKMHRRENCNKSLD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXU2</accession>
    <name>A0A452GXU2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">UBA domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum membrane</location>
            <topology evidence="1">Single-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000008078.1">
        <property type="protein sequence ID" value="ENSGAGP00000006971.1"/>
        <property type="gene ID" value="ENSGAGG00000005610.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019894">
        <property type="term" value="F:kinesin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007018">
        <property type="term" value="P:microtubule-based movement"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024854">
        <property type="entry name" value="Kinectin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007794">
        <property type="entry name" value="Rib_rcpt_KP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015940">
        <property type="entry name" value="UBA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18864">
        <property type="entry name" value="KINECTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18864:SF1">
        <property type="entry name" value="KINECTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05104">
        <property type="entry name" value="Rib_recp_KP_reg"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50030">
        <property type="entry name" value="UBA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="5 8" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 8" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 8" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="7"/>
            <end position="29"/>
        </location>
    </feature>
    <feature type="domain" description="UBA" evidence="9">
        <location>
            <begin position="744"/>
            <end position="790"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="48"/>
            <end position="194"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="307"/>
            <end position="370"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="407"/>
            <end position="498"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="566"/>
            <end position="1050"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="1116"/>
            <end position="1252"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="73"/>
            <end position="86"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="7">
        <location>
            <begin position="113"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="123"/>
            <end position="135"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="7">
        <location>
            <begin position="161"/>
            <end position="171"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="172"/>
            <end position="181"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004389"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006971.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1345" mass="153706" checksum="DCA9BBCCB8EEC6A1" modified="2019-05-08" version="1">MEFYESTYFIILIPSVVITVIFLFFWLFMKETLYDEVLAKQKRDQKFLPSKMDKKKTEKKKNKRKDAQNGNLHESDSESTPREFKLSDAMIADEEPILPVPLGLTEASSSVRERKKKEKKHKTVQEEHVSKESEGSKPAGKKIEPVPVTKQPTPPSEAAGSKKKPGQKKQKNGIDDQDSKSESFTSPTKKQDPLLFQHEIKQESGSGKKKVSTKKQKIDNVAALVDEPLIHATTYIPLMDNSDLNPLAEKREVADVGKQDTTEMIQKSGAKKLKNETDKENAEVKFKDFLLAMKSMIFTEDEALCVVELLKEKSSVIQDALQKASKEDSAATMHQLQEKDKMLAALKEEAAITKDQCKQLTQELLAEKERSNLVTTKMRERINVLEKEHGTFQSKIHVSYQENQGMKMKFQQLREQMEAEIGHLKQENGILRDAVSTSTNQMESKQSSELNKLRQDCARLVNELTEKNNKLQQEELQKKNAEQTIGQLKVQQQEAERRWEEIQAYLRKRIAEHEAAQQDVQNKLVAKDSEVQSLHSKLTDTIVSKQQLEQRMLQLMDAEQKRVTAEDSVQLQVQELMEQNEALKAQLQKFHSQMAAQSSASVLAEELQKVIAEKDKQIKQVEDSLVNEHANLAGKEEELKVIQNMNLSLKAEVQKLQALTNEQAAAAHELERMQKSIHIKDDKIRALEEQLQGQLAQVSNTREEFKVLKDQNKALQSEVQKLQALLSEQANKELLEQMERSITEKDEKIKTVEELLEAGLIQVANREEELKALRAENSSLKKEVQNLQIQQSEQVSFESVVEELQKVIREKDGKIKLVEELLEAEVLKVTNKEKTVQAFTQEIEALKEEVGHSQLEMKKQVSIASQVNDLQNLLKGKEEQMKTMEALLEEKAKDIAKKGESLQSQQDTITHLTSKVQELEQQNSEQLQQVSTVSQTQDLEILLKEKEEEVKKMEAVLEERGREITNKGKQLQEAQKENELFKMQIQELKQETHKQASLAVQSDELLQVIAGKETEITSLQNELGSLKNAVEQQRKKNNDLREKNWKAMEALASTEKLLQDKVNKTAKERQQYLETVEMETRELLQKLFPKVSLPSNLSHKEWIDGFEKMASEYLQEASGSEEVKAMEQKLKEAEEMHIMLQLECEKYKAVLAETEGILQRLQRSVEEEESKWKIKIEESQKELRQMHSSVGSLEHEIERLKEEIKEAETLKDLLTELQKKLDDSYSEAVRQNEELNLLKTQLNDTLAKLKIDQSERQKVAGDLPKAQESLASLEKEIGKAAGDANVIENSDGCSEPELTEKRLNLAVTLSQDVGHLKKLVVSISQMLSKGKEHYQLNEVTGESLI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXU4</accession>
    <name>A0A452GXU4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">UBA domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum membrane</location>
            <topology evidence="1">Single-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000008086.1">
        <property type="protein sequence ID" value="ENSGAGP00000006979.1"/>
        <property type="gene ID" value="ENSGAGG00000005610.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019894">
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007018">
        <property type="term" value="P:microtubule-based movement"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024854">
        <property type="entry name" value="Kinectin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007794">
        <property type="entry name" value="Rib_rcpt_KP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015940">
        <property type="entry name" value="UBA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18864">
        <property type="entry name" value="KINECTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18864:SF1">
        <property type="entry name" value="KINECTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05104">
        <property type="entry name" value="Rib_recp_KP_reg"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50030">
        <property type="entry name" value="UBA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="5 8" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 8" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 8" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="7"/>
            <end position="29"/>
        </location>
    </feature>
    <feature type="domain" description="UBA" evidence="9">
        <location>
            <begin position="744"/>
            <end position="790"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="48"/>
            <end position="194"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="307"/>
            <end position="370"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="407"/>
            <end position="498"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="566"/>
            <end position="1046"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="1087"/>
            <end position="1251"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="73"/>
            <end position="86"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="7">
        <location>
            <begin position="113"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="123"/>
            <end position="135"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="7">
        <location>
            <begin position="161"/>
            <end position="171"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="172"/>
            <end position="181"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004389"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006979.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1344" mass="153792" checksum="31BA5B20CCA085DC" modified="2019-05-08" version="1">MEFYESTYFIILIPSVVITVIFLFFWLFMKETLYDEVLAKQKRDQKFLPSKMDKKKTEKKKNKRKDAQNGNLHESDSESTPREFKLSDAMIADEEPILPVPLGLTEASSSVRERKKKEKKHKTVQEEHVSKESEGSKPAGKKIEPVPVTKQPTPPSEAAGSKKKPGQKKQKNGIDDQDSKSESFTSPTKKQDPLLFQHEIKQESGSGKKKVSTKKQKIDNVAALVDEPLIHATTYIPLMDNSDLNPLAEKREVADVGKQDTTEMIQKSGAKKLKNETDKENAEVKFKDFLLAMKSMIFTEDEALCVVELLKEKSSVIQDALQKASKEDSAATMHQLQEKDKMLAALKEEAAITKDQCKQLTQELLAEKERSNLVTTKMRERINVLEKEHGTFQSKIHVSYQENQGMKMKFQQLREQMEAEIGHLKQENGILRDAVSTSTNQMESKQSSELNKLRQDCARLVNELTEKNNKLQQEELQKKNAEQTIGQLKVQQQEAERRWEEIQAYLRKRIAEHEAAQQDVQNKLVAKDSEVQSLHSKLTDTIVSKQQLEQRMLQLMDAEQKRVTAEDSVQLQVQELMEQNEALKAQLQKFHSQMAAQSSASVLAEELQKVIAEKDKQIKQVEDSLVNEHANLAGKEEELKVIQNMNLSLKAEVQKLQALTNEQAAAAHELERMQKSIHIKDDKIRALEEQLQGQLAQVSNTREEFKVLKDQNKALQSEVQKLQALLSEQANKELLEQMERSITEKDEKIKTVEELLEAGLIQVANREEELKALRAENSSLKKEVQNLQIQQSEQVSFESVVEELQKVIREKDGKIKLVEELLEAEVLKVTNKEKTVQAFTQEIEALKEEVGHSQLEMKKQVSIASQVNDLQNLLKGKEEQMKTMEALLEEKAKDIAKKGESLQSQQDTITHLTSKVQELEQQNSEQLQQVSTVSQTQDLEILLKEKEEEVKKMEAVLEERGREITNKGKQLQEAQKENELFKMQIQELKQETHKQASLAVQSDELLQVIAGKETEITSLQNELGSLKNAVEQQRKKNNERQQYLETVEMETRELLQKLFPKVSLPSNLSHKEWIDGFEKMASEYLQEASGSEEVKAMEQKLKEAEEMHIMLQLECEKYKAVLAETEGILQRLQRSVEEEESKWKIKIEESQKELRQMHSSVGSLEHEIERLKEEIKEAETLKREREHLESELEKAEIERSTYVSEVRELKDLLTELQKKLDDSYSEAVRQNEELNLLKTQLNDTLAKLKIDQSERQKVAGDLPKAQESLASLEKEIGKAAGDANVIENSDGCSEPELTEKRLNLAVTLSQDVGHLKKLVVSISQMLSKGKEHYQLNEVTGESLI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXU9</accession>
    <name>A0A452GXU9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Dynein axonemal light chain 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Cilium axoneme</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="10">Belongs to the dynein light chain LC1-type family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GXU9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007967.1">
        <property type="protein sequence ID" value="ENSGAGP00000006867.1"/>
        <property type="gene ID" value="ENSGAGG00000005536.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005930">
        <property type="term" value="C:axoneme"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030286">
        <property type="term" value="C:dynein complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005874">
        <property type="term" value="C:microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043014">
        <property type="term" value="F:alpha-tubulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045504">
        <property type="term" value="F:dynein heavy chain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036158">
        <property type="term" value="P:outer dynein arm assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000049">
        <property type="entry name" value="Dynein light chain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001611">
        <property type="entry name" value="Leu-rich_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025875">
        <property type="entry name" value="Leu-rich_rpt_4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15454:SF33">
        <property type="entry name" value="DYNEIN AXONEMAL LIGHT CHAIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15454">
        <property type="entry name" value="NISCHARIN RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12799">
        <property type="entry name" value="LRR_4"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00365">
        <property type="entry name" value="LRR_SD22"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52058">
        <property type="entry name" value="L domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51450">
        <property type="entry name" value="LRR"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0966">Cell projection</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="6" id="KW-0243">Dynein</keyword>
    <keyword evidence="3" id="KW-0433">Leucine-rich repeat</keyword>
    <keyword evidence="4" id="KW-0493">Microtubule</keyword>
    <keyword evidence="7" id="KW-0505">Motor protein</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004430"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022614"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023017"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00049659"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00049760"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006867.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="181" mass="20450" checksum="CAAB37E40DAE9451" modified="2019-05-08" version="1">MEERNGQKASEAKEVKLYAQIPPVEKMDASLSTLGNCEKLSLSTNCIEKIANLNGLKNLRILSLGRNNIKNLNGLEAVGDTLEELWISYNLIEKLKGIHVMKKLKILYMSNNLVKDWAEFVRLADLPLLEDLVFVGNPLEEKYSADQQSNWVEEATKRVPRLKKLDGVPVIKQEEGEEGEN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXX0</accession>
    <name>A0A452GXX0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">G-protein coupled receptors family 1 profile domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GXX0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008070.1">
        <property type="protein sequence ID" value="ENSGAGP00000006963.1"/>
        <property type="gene ID" value="ENSGAGG00000005601.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004930">
        <property type="term" value="F:G protein-coupled receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1070.10">
        <property type="entry name" value="Rhodopsin 7-helix transmembrane proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000276">
        <property type="entry name" value="GPCR_Rhodpsn"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017452">
        <property type="entry name" value="GPCR_Rhodpsn_7TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026234">
        <property type="entry name" value="MRGPCRFAMILY"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11334">
        <property type="entry name" value="MAS-RELATED G-PROTEIN COUPLED RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11334:SF29">
        <property type="entry name" value="MAS-RELATED G-PROTEIN COUPLED RECEPTOR MEMBER X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00237">
        <property type="entry name" value="GPCRRHODOPSN"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81321">
        <property type="entry name" value="Family A G protein-coupled receptor-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50262">
        <property type="entry name" value="G_PROTEIN_RECEP_F1_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="5" id="KW-0297">G-protein coupled receptor</keyword>
    <keyword evidence="6 9" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0675">Receptor</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0807">Transducer</keyword>
    <keyword evidence="3 9" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 9" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="29"/>
            <end position="53"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="65"/>
            <end position="90"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="96"/>
            <end position="120"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="140"/>
            <end position="159"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="179"/>
            <end position="205"/>
        </location>
    </feature>
    <feature type="domain" description="G-protein coupled receptors family 1 profile" evidence="10">
        <location>
            <begin position="45"/>
            <end position="130"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023224"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50262"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006963.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="241" mass="27609" checksum="681C3FDC5F392AC3" modified="2019-05-08" version="1">SKEAGKLGLLSVLRRICPITAHLVLHQEFVISSISLFICLVGLVGNGMVLWFLGFPVKKEPFTIYILNLAIADFGFLLCMVAFLIIIILYLHMDFLVGFVVIEAIQWVALFIYNTGLYLLTDISIHRCLSCYFCINDYDCSKMTIFSCVLSFLIFTPLMVRCSSHRLIVLMAFLNHSMIHLFVMSLVILLPCMNSSINSFLYFLIGRHGKQQLREPLREVFQRIFNEKAGPQEKRPTKQVK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXX1</accession>
    <name>A0A452GXX1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="22">Serine/threonine-protein kinase VRK2</fullName>
            <ecNumber evidence="5">2.7.11.1</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="23">Vaccinia-related kinase 2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="26 27">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="27" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="26" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="26" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="20">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="20">
            <dbReference type="Rhea" id="RHEA:17990"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="19">
            <dbReference type="Rhea" id="RHEA:46609"/>
        </physiologicalReaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="4">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum membrane</location>
            <topology evidence="1">Single-pass type IV membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Mitochondrion membrane</location>
            <topology evidence="3">Single-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Nucleus envelope</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="21">Belongs to the protein kinase superfamily. CK1 Ser/Thr protein kinase family. VRK subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="5"/>
    <dbReference type="AlphaFoldDB" id="A0A452GXX1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007990.1">
        <property type="protein sequence ID" value="ENSGAGP00000006888.1"/>
        <property type="gene ID" value="ENSGAGG00000005556.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031966">
        <property type="term" value="C:mitochondrial membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005635">
        <property type="term" value="C:nuclear envelope"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000532">
        <property type="entry name" value="VRK serine/threonine kinase 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050235">
        <property type="entry name" value="CK1_Ser-Thr_kinase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11909">
        <property type="entry name" value="CASEIN KINASE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="14" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="13" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="12" id="KW-0418">Kinase</keyword>
    <keyword evidence="17" id="KW-0472">Membrane</keyword>
    <keyword evidence="16" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="11" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="18" id="KW-0539">Nucleus</keyword>
    <keyword evidence="8" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="27" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="9" id="KW-0808">Transferase</keyword>
    <keyword evidence="10" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="15" id="KW-1133">Transmembrane helix</keyword>
    <feature type="domain" description="Protein kinase" evidence="25">
        <location>
            <begin position="29"/>
            <end position="327"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="24">
        <location>
            <begin position="406"/>
            <end position="427"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004259"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004304"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012513"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00048659"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00048977"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00061699"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00073146"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00081231"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006888.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="540" mass="61531" checksum="91AF6171F24DA82F" modified="2019-05-08" version="1">MPPKGRGKSKLPVPLPKGMIMKDTEGKEWRLGNIIGQGGFGLIYLASSHVHIPVEDDAVHVIKVEYLENGPLFSELKFYQRAAKHEHIKKWMTLRQIGFLGIPMFWGTGLAEYNGKSYRFMVMERLGVDLQRILDNDGHRFKKETVLQLGARMLDALEYMHENEYVHGDIKAANLLLGYTNPHEVYLADYGLSYRYCPNGNHKQYKENPKKGHNGTIEFTSLDAHKGVAPSRRGDLEILGYCMLQWLCGKLPWEQNLKDPVSVQTAKTKLMDELPDSVMKWGPAGSSCCEIAKFLACVYGLTYEEKPKYQVLKKILVDGLKSSGTFYDGPLEFSTAGCALNLCASKDLKADLPKPTPKQCKQKENKLNDKEHNRLRKAGIQVTQRQLQEQEKPAGLNKMHQHNEPLLADLPKPTPKQCKQKENKLDDKEHNRLRKAGIQVTQRQLQEQEKPVGLNKMCQPNEPLLENTVRQLHRHYLQTVFCEVKKKPHENSITYSPVSLQEPSRGFTSPSVSNIFKLAFTEERSPYLGSMNWSQVDLLH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXX5</accession>
    <name>A0A452GXX5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Calpain-2 catalytic subunit</fullName>
            <ecNumber evidence="6">3.4.22.53</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="18">Calcium-activated neutral proteinase 2</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="19">Calpain M-type</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="20">Calpain-2 large subunit</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="17">Millimolar-calpain</fullName>
        </alternativeName>
    </protein>
    <organism evidence="26 27">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="27" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="26" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="26" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="21">Calcium-regulated non-lysosomal thiol-protease which catalyze limited proteolysis of substrates involved in cytoskeletal remodeling and signal transduction.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>Broad endopeptidase specificity.</text>
            <dbReference type="EC" id="3.4.22.53"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="2">
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the peptidase C2 family.</text>
    </comment>
    <dbReference type="EC" id="3.4.22.53" evidence="6"/>
    <dbReference type="AlphaFoldDB" id="A0A452GXX5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008075.1">
        <property type="protein sequence ID" value="ENSGAGP00000006968.1"/>
        <property type="gene ID" value="ENSGAGG00000004685.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004198">
        <property type="term" value="F:calcium-dependent cysteine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00214">
        <property type="entry name" value="Calpain_III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00044">
        <property type="entry name" value="CysPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.380:FF:000001">
        <property type="entry name" value="Calpain-1 catalytic subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.70.10:FF:000001">
        <property type="entry name" value="Calpain-1 catalytic subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.380">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.70.10">
        <property type="entry name" value="Cysteine proteinases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.238.10">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033883">
        <property type="entry name" value="C2_III"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022684">
        <property type="entry name" value="Calpain_cysteine_protease"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022682">
        <property type="entry name" value="Calpain_domain_III"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022683">
        <property type="entry name" value="Calpain_III"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036213">
        <property type="entry name" value="Calpain_III_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011992">
        <property type="entry name" value="EF-hand-dom_pair"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018247">
        <property type="entry name" value="EF_Hand_1_Ca_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002048">
        <property type="entry name" value="EF_hand_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038765">
        <property type="entry name" value="Papain-like_cys_pep_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000169">
        <property type="entry name" value="Pept_cys_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001300">
        <property type="entry name" value="Peptidase_C2_calpain_cat"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10183">
        <property type="entry name" value="CALPAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10183:SF268">
        <property type="entry name" value="CALPAIN-2 CATALYTIC SUBUNIT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01067">
        <property type="entry name" value="Calpain_III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00648">
        <property type="entry name" value="Peptidase_C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00704">
        <property type="entry name" value="CALPAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00720">
        <property type="entry name" value="calpain_III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00230">
        <property type="entry name" value="CysPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49758">
        <property type="entry name" value="Calpain large subunit, middle domain (domain III)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54001">
        <property type="entry name" value="Cysteine proteinases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47473">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50203">
        <property type="entry name" value="CALPAIN_CAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00018">
        <property type="entry name" value="EF_HAND_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50222">
        <property type="entry name" value="EF_HAND_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00139">
        <property type="entry name" value="THIOL_PROTEASE_CYS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="15" id="KW-0106">Calcium</keyword>
    <keyword evidence="8" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="9" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="13 23" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="16" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10 23" id="KW-0645">Protease</keyword>
    <keyword evidence="27" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="12" id="KW-0677">Repeat</keyword>
    <keyword evidence="14 23" id="KW-0788">Thiol protease</keyword>
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        <location>
            <begin position="45"/>
            <end position="344"/>
        </location>
    </feature>
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        <location>
            <begin position="528"/>
            <end position="563"/>
        </location>
    </feature>
    <feature type="active site" evidence="22 23">
        <location>
            <position position="105"/>
        </location>
    </feature>
    <feature type="active site" evidence="22 23">
        <location>
            <position position="262"/>
        </location>
    </feature>
    <feature type="active site" evidence="22 23">
        <location>
            <position position="286"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001223"/>
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        <source>
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    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
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    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00007623"/>
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    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00012481"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00014052"/>
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    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
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    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
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    </evidence>
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        <source>
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    </evidence>
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    </evidence>
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        <source>
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    </evidence>
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        <source>
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    </evidence>
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        <source>
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    </evidence>
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        <source>
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        <source>
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            <dbReference type="ARBA" id="ARBA00032544"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00054695"/>
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    </evidence>
    <evidence type="ECO:0000256" key="22">
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    </evidence>
    <evidence type="ECO:0000256" key="23">
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        <source>
            <dbReference type="PROSITE" id="PS50222"/>
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    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
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    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="621" mass="70778" checksum="8B550AFA6E73F33A" modified="2019-05-08" version="1">MSGIAVKLAKERETARGLGSNAKAFKYLSQDFETLRNECLQAGKLFQDPSFPAIASSLGYKELGPNSYKTQGITWKRPTELCSNPQFIVGGATRTDICQGALGDCWLLAAIASLTLNEEILARVVPKDQSFQDKYAGIFHFQFWQYGEWVDVVVDDKLPTKNGELLFVHSAEGSEFWSALLEKAYAKVNGSYEALSGGSTTEGFEDFTGGIAEWYELQKAPPNLFKIIQKALLKGSLLGCSIDITSAAETEAVTSQKLVKGHAYSVTGAEEVNCRGSMQKLIRIRNPWGEVEWTGKWNDNCSNWNSVAPEVRERLTRRHEDGEFWMSFSDFLRHYSRLEICNLTPDTLANEKYKKWSLTKLDGNWRRGSTAGGCRNYPNTFWMNPQYLIKLEEEDEDPDDPEKGCTFLVGLIQKHRRRQRKMGEDMHTIGFGIYEASRFSGQTNVHLSKNFFLTNRAREKSNTFINLREVLNRFKLPPGEYVIVPSTFEPNKDGDFCLRVFSEKNADSQYVSDGSGKLGLKEFYMLWTKIQKYQKIYREIDVDRSGTMNSYEMRRALEEAGFKLTCQLHQVIVARFADEQLIIDFDNFVRCLIRLETLFKIFKKLDVEKSGTVEMNLNTVS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXX8</accession>
    <name>A0A452GXX8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Calpain-2 catalytic subunit</fullName>
            <ecNumber evidence="6">3.4.22.53</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="18">Calcium-activated neutral proteinase 2</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="19">Calpain M-type</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="20">Calpain-2 large subunit</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="17">Millimolar-calpain</fullName>
        </alternativeName>
    </protein>
    <organism evidence="26 27">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="27" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="26" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="26" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="21">Calcium-regulated non-lysosomal thiol-protease which catalyze limited proteolysis of substrates involved in cytoskeletal remodeling and signal transduction.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>Broad endopeptidase specificity.</text>
            <dbReference type="EC" id="3.4.22.53"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="2">
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the peptidase C2 family.</text>
    </comment>
    <dbReference type="EC" id="3.4.22.53" evidence="6"/>
    <dbReference type="AlphaFoldDB" id="A0A452GXX8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008072.1">
        <property type="protein sequence ID" value="ENSGAGP00000006965.1"/>
        <property type="gene ID" value="ENSGAGG00000004685.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004198">
        <property type="term" value="F:calcium-dependent cysteine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00214">
        <property type="entry name" value="Calpain_III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00044">
        <property type="entry name" value="CysPc"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Calpain-1 catalytic subunit"/>
        <property type="match status" value="1"/>
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        <property type="match status" value="1"/>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Cysteine proteinases"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="C2_III"/>
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        <property type="entry name" value="Calpain_cysteine_protease"/>
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        <property type="entry name" value="Calpain_domain_III"/>
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        <property type="entry name" value="Calpain_III"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036213">
        <property type="entry name" value="Calpain_III_sf"/>
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    <dbReference type="InterPro" id="IPR011992">
        <property type="entry name" value="EF-hand-dom_pair"/>
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    <dbReference type="InterPro" id="IPR018247">
        <property type="entry name" value="EF_Hand_1_Ca_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002048">
        <property type="entry name" value="EF_hand_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038765">
        <property type="entry name" value="Papain-like_cys_pep_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000169">
        <property type="entry name" value="Pept_cys_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001300">
        <property type="entry name" value="Peptidase_C2_calpain_cat"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10183">
        <property type="entry name" value="CALPAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10183:SF268">
        <property type="entry name" value="CALPAIN-2 CATALYTIC SUBUNIT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01067">
        <property type="entry name" value="Calpain_III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00648">
        <property type="entry name" value="Peptidase_C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00704">
        <property type="entry name" value="CALPAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00720">
        <property type="entry name" value="calpain_III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00230">
        <property type="entry name" value="CysPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49758">
        <property type="entry name" value="Calpain large subunit, middle domain (domain III)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54001">
        <property type="entry name" value="Cysteine proteinases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47473">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50203">
        <property type="entry name" value="CALPAIN_CAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00018">
        <property type="entry name" value="EF_HAND_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50222">
        <property type="entry name" value="EF_HAND_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00139">
        <property type="entry name" value="THIOL_PROTEASE_CYS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="15" id="KW-0106">Calcium</keyword>
    <keyword evidence="8" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="9" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="13 23" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="16" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10 23" id="KW-0645">Protease</keyword>
    <keyword evidence="27" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="12" id="KW-0677">Repeat</keyword>
    <keyword evidence="14 23" id="KW-0788">Thiol protease</keyword>
    <feature type="domain" description="Calpain catalytic" evidence="24">
        <location>
            <begin position="45"/>
            <end position="344"/>
        </location>
    </feature>
    <feature type="domain" description="EF-hand" evidence="25">
        <location>
            <begin position="569"/>
            <end position="604"/>
        </location>
    </feature>
    <feature type="active site" evidence="22 23">
        <location>
            <position position="105"/>
        </location>
    </feature>
    <feature type="active site" evidence="22 23">
        <location>
            <position position="262"/>
        </location>
    </feature>
    <feature type="active site" evidence="22 23">
        <location>
            <position position="286"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001223"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00001913"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00007623"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00012481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00014052"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
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        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
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        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00030109"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00031308"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00032449"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00032544"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00054695"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="PIRSR" id="PIRSR622684-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS50203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS50222"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006965.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="662" mass="75417" checksum="D3CC650A91476314" modified="2019-05-08" version="1">MSGIAVKLAKERETARGLGSNAKAFKYLSQDFETLRNECLQAGKLFQDPSFPAIASSLGYKELGPNSYKTQGITWKRPTELCSNPQFIVGGATRTDICQGALGDCWLLAAIASLTLNEEILARVVPKDQSFQDKYAGIFHFQFWQYGEWVDVVVDDKLPTKNGELLFVHSAEGSEFWSALLEKAYAKVNGSYEALSGGSTTEGFEDFTGGIAEWYELQKAPPNLFKIIQKALLKGSLLGCSIDITSAAETEAVTSQKLVKGHAYSVTGAEEVNCRGSMQKLIRIRNPWGEVEWTGKWNDNCSNWNSVAPEVRERLTRRHEDGEFWMSFSDFLRHYSRLEICNLTPDTLANEKYKKWSLTKLDGNWRRGSTAGGCRNYPNTFWMNPQYLIKLEEEDEDPDDPEKGCTFLVGLIQKHRRRQRKMGEDMHTIGFGIYEASRFSGQTNVHLSKNFFLTNRAREKSNTFINLREVLNRFKLPPGEYVIVPSTFEPNKDGDFCLRVFSEKNADSQYVDAEISAFELCNILRKIMAKRQDIKSDGFSIETCKIMVDLLDSDGSGKLGLKEFYILWTKIQKYQKIYREIDVDRSGTMNSYEMRRALEEAGFKLTCQLHQVIVARFADEQLIIDFDNFVRCLIRLETLFKIFKKLDVEKSGTVEMNLNTVS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXY0</accession>
    <name>A0A452GXY0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">G-protein coupled receptors family 1 profile domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="9">Belongs to the G-protein coupled receptor 1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GXY0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008124.1">
        <property type="protein sequence ID" value="ENSGAGP00000007014.1"/>
        <property type="gene ID" value="ENSGAGG00000005601.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004930">
        <property type="term" value="F:G protein-coupled receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1070.10">
        <property type="entry name" value="Rhodopsin 7-helix transmembrane proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000276">
        <property type="entry name" value="GPCR_Rhodpsn"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017452">
        <property type="entry name" value="GPCR_Rhodpsn_7TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026234">
        <property type="entry name" value="MRGPCRFAMILY"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11334">
        <property type="entry name" value="MAS-RELATED G-PROTEIN COUPLED RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11334:SF29">
        <property type="entry name" value="MAS-RELATED G-PROTEIN COUPLED RECEPTOR MEMBER X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00001">
        <property type="entry name" value="7tm_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00237">
        <property type="entry name" value="GPCRRHODOPSN"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR02108">
        <property type="entry name" value="MRGPCRFAMILY"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81321">
        <property type="entry name" value="Family A G protein-coupled receptor-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00237">
        <property type="entry name" value="G_PROTEIN_RECEP_F1_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50262">
        <property type="entry name" value="G_PROTEIN_RECEP_F1_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="5 9" id="KW-0297">G-protein coupled receptor</keyword>
    <keyword evidence="6 10" id="KW-0472">Membrane</keyword>
    <keyword evidence="7 9" id="KW-0675">Receptor</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8 9" id="KW-0807">Transducer</keyword>
    <keyword evidence="3 9 10" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 10" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="14"/>
            <end position="41"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="53"/>
            <end position="78"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="84"/>
            <end position="108"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="151"/>
            <end position="177"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="183"/>
            <end position="209"/>
        </location>
    </feature>
    <feature type="domain" description="G-protein coupled receptors family 1 profile" evidence="11">
        <location>
            <begin position="33"/>
            <end position="126"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023224"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="RuleBase" id="RU000688"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50262"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007014.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="245" mass="28047" checksum="0A10C2A94BEDCDEA" modified="2019-05-08" version="1">DLPTIYTHSSATSLLIFLGLSISLFICLVGLVGNGMVLWFLGFPVKKEPFTIYILNLAIADFGFLLCMVAFLIIIILYLHMDFLVGFVVIEAIQWVALFIYNTGLYLLTDISIHRCLSVLYPIWYRCHIQKTCLPLCVLCSGAISTLFIKLYIVIMVTILFFLMFALPPRVIVLMAFLNHSMIHLFVMSLVILLPCMNSSINSFLYFLIGRHGKQQLREPLREVFQRIFNEKAGPQEKRPTKQVK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXY1</accession>
    <name>A0A452GXY1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Calpain-2 catalytic subunit</fullName>
            <ecNumber evidence="6">3.4.22.53</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="18">Calcium-activated neutral proteinase 2</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="19">Calpain M-type</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="20">Calpain-2 large subunit</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="17">Millimolar-calpain</fullName>
        </alternativeName>
    </protein>
    <organism evidence="26 27">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="27" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="26" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="26" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="21">Calcium-regulated non-lysosomal thiol-protease which catalyze limited proteolysis of substrates involved in cytoskeletal remodeling and signal transduction.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>Broad endopeptidase specificity.</text>
            <dbReference type="EC" id="3.4.22.53"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="2">
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the peptidase C2 family.</text>
    </comment>
    <dbReference type="EC" id="3.4.22.53" evidence="6"/>
    <dbReference type="AlphaFoldDB" id="A0A452GXY1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008080.1">
        <property type="protein sequence ID" value="ENSGAGP00000006973.1"/>
        <property type="gene ID" value="ENSGAGG00000004685.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004198">
        <property type="term" value="F:calcium-dependent cysteine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00214">
        <property type="entry name" value="Calpain_III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00044">
        <property type="entry name" value="CysPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.380:FF:000001">
        <property type="entry name" value="Calpain-1 catalytic subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.70.10:FF:000001">
        <property type="entry name" value="Calpain-1 catalytic subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.380">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.70.10">
        <property type="entry name" value="Cysteine proteinases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.238.10">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033883">
        <property type="entry name" value="C2_III"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022684">
        <property type="entry name" value="Calpain_cysteine_protease"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022682">
        <property type="entry name" value="Calpain_domain_III"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022683">
        <property type="entry name" value="Calpain_III"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036213">
        <property type="entry name" value="Calpain_III_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011992">
        <property type="entry name" value="EF-hand-dom_pair"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018247">
        <property type="entry name" value="EF_Hand_1_Ca_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002048">
        <property type="entry name" value="EF_hand_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038765">
        <property type="entry name" value="Papain-like_cys_pep_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000169">
        <property type="entry name" value="Pept_cys_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001300">
        <property type="entry name" value="Peptidase_C2_calpain_cat"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10183">
        <property type="entry name" value="CALPAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10183:SF268">
        <property type="entry name" value="CALPAIN-2 CATALYTIC SUBUNIT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01067">
        <property type="entry name" value="Calpain_III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00648">
        <property type="entry name" value="Peptidase_C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00704">
        <property type="entry name" value="CALPAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00720">
        <property type="entry name" value="calpain_III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00230">
        <property type="entry name" value="CysPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49758">
        <property type="entry name" value="Calpain large subunit, middle domain (domain III)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54001">
        <property type="entry name" value="Cysteine proteinases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47473">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50203">
        <property type="entry name" value="CALPAIN_CAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00018">
        <property type="entry name" value="EF_HAND_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50222">
        <property type="entry name" value="EF_HAND_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00139">
        <property type="entry name" value="THIOL_PROTEASE_CYS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="15" id="KW-0106">Calcium</keyword>
    <keyword evidence="8" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="9" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="13 23" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="16" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10 23" id="KW-0645">Protease</keyword>
    <keyword evidence="27" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="12" id="KW-0677">Repeat</keyword>
    <keyword evidence="14 23" id="KW-0788">Thiol protease</keyword>
    <feature type="domain" description="Calpain catalytic" evidence="24">
        <location>
            <begin position="45"/>
            <end position="344"/>
        </location>
    </feature>
    <feature type="domain" description="EF-hand" evidence="25">
        <location>
            <begin position="528"/>
            <end position="563"/>
        </location>
    </feature>
    <feature type="active site" evidence="22 23">
        <location>
            <position position="105"/>
        </location>
    </feature>
    <feature type="active site" evidence="22 23">
        <location>
            <position position="262"/>
        </location>
    </feature>
    <feature type="active site" evidence="22 23">
        <location>
            <position position="286"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001223"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00001913"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00007623"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00012481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00014052"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022807"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00030109"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00031308"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00032449"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00032544"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00054695"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="PIRSR" id="PIRSR622684-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS50203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS50222"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006973.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="621" mass="70760" checksum="8DFFCC5604B559FA" modified="2019-05-08" version="1">MSGIAVKLAKERETARGLGSNAKAFKYLSQDFETLRNECLQAGKLFQDPSFPAIASSLGYKELGPNSYKTQGITWKRPTELCSNPQFIVGGATRTDICQGALGDCWLLAAIASLTLNEEILARVVPKDQSFQDKYAGIFHFQFWQYGEWVDVVVDDKLPTKNGELLFVHSAEGSEFWSALLEKAYAKVNGSYEALSGGSTTEGFEDFTGGIAEWYELQKAPPNLFKIIQKALLKGSLLGCSIDITSAAETEAVTSQKLVKGHAYSVTGAEEVNCRGSMQKLIRIRNPWGEVEWTGKWNDNCSNWNSVAPEVRERLTRRHEDGEFWMSFSDFLRHYSRLEICNLTPDTLANEKYKKWSLTKLDGNWRRGSTAGGCRNYPNTFWMNPQYLIKLEEEDEDPDDPEKGCTFLVGLIQKHRRRQRKMGEDMHTIGFGIYEASRFSGQTNVHLSKNFFLTNRAREKSNTFINLREVLNRFKLPPGEYVIVPSTFEPNKDGDFCLRVFSEKNADSQYVSDGSGKLGLKEFYILWTKIQKYQKIYREIDVDRSGTMNSYEMRRALEEAGFKLTCQLHQVIVARFADEQLIIDFDNFVRCLIRLETLFKIFKKLDVEKSGTVEMNLNTVS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXY2</accession>
    <name>A0A452GXY2_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="9">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 5">Belongs to the D-isomer specific 2-hydroxyacid dehydrogenase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GXY2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008139.1">
        <property type="protein sequence ID" value="ENSGAGP00000007025.1"/>
        <property type="gene ID" value="ENSGAGG00000005657.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140297">
        <property type="term" value="F:DNA-binding transcription factor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051287">
        <property type="term" value="F:NAD binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016616">
        <property type="term" value="F:oxidoreductase activity, acting on the CH-OH group of donors, NAD or NADP as acceptor"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003713">
        <property type="term" value="F:transcription coactivator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001221">
        <property type="term" value="F:transcription coregulator binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003714">
        <property type="term" value="F:transcription corepressor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05299">
        <property type="entry name" value="CtBP_dh"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.720:FF:000012">
        <property type="entry name" value="C-terminal-binding protein 2 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.720">
        <property type="entry name" value="NAD(P)-binding Rossmann-like Domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043322">
        <property type="entry name" value="CtBP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051638">
        <property type="entry name" value="CTBP_dehydrogenase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006139">
        <property type="entry name" value="D-isomer_2_OHA_DH_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029753">
        <property type="entry name" value="D-isomer_DH_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029752">
        <property type="entry name" value="D-isomer_DH_CS1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006140">
        <property type="entry name" value="D-isomer_DH_NAD-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036291">
        <property type="entry name" value="NAD(P)-bd_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46029">
        <property type="entry name" value="C-TERMINAL-BINDING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46029:SF2">
        <property type="entry name" value="C-TERMINAL-BINDING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00389">
        <property type="entry name" value="2-Hacid_dh"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02826">
        <property type="entry name" value="2-Hacid_dh_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52283">
        <property type="entry name" value="Formate/glycerate dehydrogenase catalytic domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51735">
        <property type="entry name" value="NAD(P)-binding Rossmann-fold domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00065">
        <property type="entry name" value="D_2_HYDROXYACID_DH_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00671">
        <property type="entry name" value="D_2_HYDROXYACID_DH_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0539">Nucleus</keyword>
    <keyword evidence="3 5" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="D-isomer specific 2-hydroxyacid dehydrogenase catalytic" evidence="7">
        <location>
            <begin position="38"/>
            <end position="352"/>
        </location>
    </feature>
    <feature type="domain" description="D-isomer specific 2-hydroxyacid dehydrogenase NAD-binding" evidence="8">
        <location>
            <begin position="135"/>
            <end position="317"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="409"/>
            <end position="441"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="430"/>
            <end position="441"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005854"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="RuleBase" id="RU003719"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF00389"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF02826"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007025.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="441" mass="47822" checksum="1DCCA345A6BDCD80" modified="2019-05-08" version="1">MGSSHLLNKGLPLGIRPPIMNGPMHPRPLVALLDGRDCTVEMPILKDVATVAFCDAQSTQEIHEKVLNEAVGALMYHTITLTREDLEKFKALRIIVRIGSGFDNIDIKSAGDLGIAVCNVPAASVEETADSTMCHILNLYRRTTWLHQALREGTRVQSVEQIREVASGAARIRGETLGIIGLGRVGQAVALRAKAFGFSVLFYDPYLSDGIERALGLQRVSTLQDLLFHSDCVTLHCNLNEHNHHLINDFTIKQMRQGAFLVNTARGGLVDEKALAQALKEGRIRGAALDVHESEPFSFSQGPLKDAPNLICTPHAAWYSEQASIEMREEAAREIRRAITGRIPDSLKNCVNKDHLTAATPWASMDPGVVHPELNGAAYSRYPPGVVSMAPAGIPTAVEGIVPNPMSLSHGLPAVAHPPHAPSPGQTVKPEADRDHPSDQL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXY8</accession>
    <name>A0A452GXY8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">ATP binding cassette subfamily C member 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>17beta-estradiol 17-O-(beta-D-glucuronate)(in) + ATP + H2O = 17beta-estradiol 17-O-(beta-D-glucuronate)(out) + ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:60128"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:82961"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="15">
            <dbReference type="Rhea" id="RHEA:60129"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>ATP + H2O + xenobioticSide 1 = ADP + phosphate + xenobioticSide 2.</text>
            <dbReference type="EC" id="7.6.2.2"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="14">
            <text>leukotriene C4(in) + ATP + H2O = leukotriene C4(out) + ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:38963"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:57973"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="14">
            <dbReference type="Rhea" id="RHEA:38964"/>
        </physiologicalReaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the ABC transporter superfamily. ABCC family. Conjugate transporter (TC 3.A.1.208) subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GXY8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008133.1">
        <property type="protein sequence ID" value="ENSGAGP00000007021.1"/>
        <property type="gene ID" value="ENSGAGG00000004641.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016324">
        <property type="term" value="C:apical plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008559">
        <property type="term" value="F:ABC-type xenobiotic transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006869">
        <property type="term" value="P:lipid transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd18595">
        <property type="entry name" value="ABC_6TM_MRP1_2_3_6_D1_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18603">
        <property type="entry name" value="ABC_6TM_MRP1_2_3_6_D2_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd03250">
        <property type="entry name" value="ABCC_MRP_domain1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd03244">
        <property type="entry name" value="ABCC_MRP_domain2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000293">
        <property type="entry name" value="ATP binding cassette subfamily C member 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1560.10:FF:000001">
        <property type="entry name" value="ATP-binding cassette subfamily C member 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1560.10:FF:000007">
        <property type="entry name" value="ATP-binding cassette subfamily C member 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000074">
        <property type="entry name" value="Multidrug resistance-associated protein 5 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1560.10">
        <property type="entry name" value="ABC transporter type 1, transmembrane domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003593">
        <property type="entry name" value="AAA+_ATPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011527">
        <property type="entry name" value="ABC1_TM_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036640">
        <property type="entry name" value="ABC1_TM_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003439">
        <property type="entry name" value="ABC_transporter-like_ATP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017871">
        <property type="entry name" value="ABC_transporter-like_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050173">
        <property type="entry name" value="ABC_transporter_C-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24223">
        <property type="entry name" value="ATP-BINDING CASSETTE SUB-FAMILY C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24223:SF176">
        <property type="entry name" value="ATP-BINDING CASSETTE SUB-FAMILY C MEMBER 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00664">
        <property type="entry name" value="ABC_membrane"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00005">
        <property type="entry name" value="ABC_tran"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00382">
        <property type="entry name" value="AAA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90123">
        <property type="entry name" value="ABC transporter transmembrane region"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50929">
        <property type="entry name" value="ABC_TM1F"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00211">
        <property type="entry name" value="ABC_TRANSPORTER_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50893">
        <property type="entry name" value="ABC_TRANSPORTER_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11" id="KW-0445">Lipid transport</keyword>
    <keyword evidence="12 16" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="9" id="KW-1278">Translocase</keyword>
    <keyword evidence="5 16" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="10 16" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="40"/>
            <end position="65"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="111"/>
            <end position="134"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="140"/>
            <end position="159"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="221"/>
            <end position="245"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="265"/>
            <end position="285"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="623"/>
            <end position="646"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="677"/>
            <end position="696"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="745"/>
            <end position="762"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="768"/>
            <end position="788"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="853"/>
            <end position="874"/>
        </location>
    </feature>
    <feature type="domain" description="ABC transmembrane type-1" evidence="18">
        <location>
            <begin position="2"/>
            <end position="282"/>
        </location>
    </feature>
    <feature type="domain" description="ABC transporter" evidence="17">
        <location>
            <begin position="304"/>
            <end position="530"/>
        </location>
    </feature>
    <feature type="domain" description="ABC transmembrane type-1" evidence="18">
        <location>
            <begin position="679"/>
            <end position="895"/>
        </location>
    </feature>
    <feature type="domain" description="ABC transporter" evidence="17">
        <location>
            <begin position="927"/>
            <end position="1161"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009726"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022967"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00034018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00047523"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00047576"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50893"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50929"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007021.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="21">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1172" mass="128533" checksum="C5797979448CFBB0" modified="2019-05-08" version="1">KSVAFKVVHDLLVFVSPQLLKLLISFVADPTAYAWQGYLYALLLFVTALVQSLCLQQYFQLCFVLGMSLRTALMAAVYKKALTVSNATRKEYTQGETVNLMSADTQRFMDLATFIHQLWSAPLQIALSIIFLWWELGPSVLAGIGVMVLLIPINAVLATKARRIQMKNMKNKDQRMKIMNEILSGIKILKLFAWEPSFERQVGEIRVRELKGLLHFTNLQAFSIFIFSCAPFLVSVASFAVYVMVDEKNVLDAQKAFTSISLFNILRFPMATLPLVISSLVQTAVSTERLERYLGSGNLDPSAIKCDPSVAAQRCSESQLSRPSRVTLDIKPGHLVAVVGAVGSGKSSLVSAVLGEMENLKGHINIQGSVAYVPQQAWIQNATLKDNILFGSDLDEARYQQVVEACALLPDLKLLPGGDLTEIGEKGINLSGGQKQRVSLARAVYSDADIYLLDDPLSAVDAHVGKHLFDQVLGPDGLLQGKTRILVTHSVSFLPRTDEIVVLVAGAVSEQGSYSTLLANGGAFAQLLSTYGSQEGSTPEGEATGMWAGAEPRVSGWGARVAPSPIANYEQSPYCSLSTSSTKSLRKALQAGEVGGACGGWTPWDAATECAALQVKFSLYLRYLHAVGWGFSAWVFITYVAQYAAFMGSNLWLSDWTDDSLRYWNQTYPTSQRDMRIGVFGALGTVQALLATRGAIRASRVLHQQLLSNILRVPMSFFDTTPTGRIVNRFAKDIFTVDETIPMSFRSWLTCFLGIISTLLMICLATPYFALIIIPLSILYFFLLRYYVSTSRQLRRLDSVTRSPIYSHFGETVSGLSVIRAYGHQERFLQHNDRIVDMNQKSVYSWIVSNRWLAVRLEFVGNLVVFSAALLAVITRDNLGSGIVGLSISSALNVSPVTQGNEASLLSGKCCCTFGHQRGSPPGGCQLGGGHWCRVLGWMEPRKRCSHSEGGVLSPQVGVVGRTGAGKSSLTNCLFRILEAAGGKIVIDGVDISTIGLHDLRQSLTIIPQDPVLFSGTLRMNLDPFDQHSDEEIWKALELAHLKTYVLDLPERLSHQVSEAGENLSVGQRQLLCLARALLRKSKILILDEATAAVDLETDHLIQTTIRSEFANCTVLTIAHRLHTIMDSNRVMVLQAGRIVEYDSPERLLQQQGLFSTMAKDAGIVSAQATVL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXY9</accession>
    <name>A0A452GXY9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Protein TANC1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">May be a scaffold component in the postsynaptic density.</text>
    </comment>
    <comment type="subunit">
        <text evidence="10">Interacts probably directly with DLG1, DLG4, HOMER1. Interacts with DLGAP1, INA, CAMK2A, GRIN2B and GRIA1. Interacts with TNIK and MINK1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="7">Postsynaptic density</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="8">Belongs to the TANC family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000008082.1">
        <property type="protein sequence ID" value="ENSGAGP00000006975.1"/>
        <property type="gene ID" value="ENSGAGG00000005019.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043197">
        <property type="term" value="C:dendritic spine"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0014069">
        <property type="term" value="C:postsynaptic density"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061001">
        <property type="term" value="P:regulation of dendritic spine morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd22541">
        <property type="entry name" value="SP5_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000229">
        <property type="entry name" value="protein TANC1 isoform X3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000036">
        <property type="entry name" value="protein TANC2 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000044">
        <property type="entry name" value="Tetratricopeptide repeat, ankyrin repeat and coiled-coil containing 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.10">
        <property type="entry name" value="Tetratricopeptide repeat domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058018">
        <property type="entry name" value="AAA_lid_TANC1/2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050889">
        <property type="entry name" value="Dendritic_Spine_Reg/Scaffold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011990">
        <property type="entry name" value="TPR-like_helical_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019734">
        <property type="entry name" value="TPR_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058056">
        <property type="entry name" value="WH_TANC1/2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24166:SF21">
        <property type="entry name" value="PROTEIN TANC2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24166">
        <property type="entry name" value="ROLLING PEBBLES, ISOFORM B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25520">
        <property type="entry name" value="AAA_lid_TANC1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00023">
        <property type="entry name" value="Ank"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13637">
        <property type="entry name" value="Ank_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13181">
        <property type="entry name" value="TPR_8"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25521">
        <property type="entry name" value="WHD_TANC1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01415">
        <property type="entry name" value="ANKYRIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="SMART" id="SM00028">
        <property type="entry name" value="TPR"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48452">
        <property type="entry name" value="TPR-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50005">
        <property type="entry name" value="TPR"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0007">Acetylation</keyword>
    <keyword evidence="6 12" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="1" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0770">Synapse</keyword>
    <keyword evidence="3 13" id="KW-0802">TPR repeat</keyword>
    <feature type="domain" description="TANC1/2-like AAA+ ATPase lid" evidence="15">
        <location>
            <begin position="390"/>
            <end position="484"/>
        </location>
    </feature>
    <feature type="domain" description="TANC1/2-like winged helix" evidence="16">
        <location>
            <begin position="486"/>
            <end position="638"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="12">
        <location>
            <begin position="709"/>
            <end position="741"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="12">
        <location>
            <begin position="810"/>
            <end position="842"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="12">
        <location>
            <begin position="843"/>
            <end position="875"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="12">
        <location>
            <begin position="876"/>
            <end position="908"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="12">
        <location>
            <begin position="909"/>
            <end position="941"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="12">
        <location>
            <begin position="942"/>
            <end position="974"/>
        </location>
    </feature>
    <feature type="repeat" description="TPR" evidence="13">
        <location>
            <begin position="1021"/>
            <end position="1054"/>
        </location>
    </feature>
    <feature type="repeat" description="TPR" evidence="13">
        <location>
            <begin position="1068"/>
            <end position="1101"/>
        </location>
    </feature>
    <feature type="repeat" description="TPR" evidence="13">
        <location>
            <begin position="1102"/>
            <end position="1135"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="188"/>
            <end position="234"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1152"/>
            <end position="1171"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1200"/>
            <end position="1386"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1405"/>
            <end position="1429"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1555"/>
            <end position="1577"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="211"/>
            <end position="223"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="1152"/>
            <end position="1161"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="1239"/>
            <end position="1268"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="1370"/>
            <end position="1385"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022803"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00034105"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00038259"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00054404"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00065031"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00070020"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00339"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="Pfam" id="PF25520"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF25521"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006975.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1577" mass="176477" checksum="A90CE920461E2B53" modified="2019-05-08" version="1">MWQLTKKNPGEVREEWVMRSERKCLRGLMPERNLRRPYPQWIREIGQWECEKEYQDLEPGLIAVCFPLAATSSAHLEDLAYLDEQRHTPLRTSLRMPRQSMAGALRFAPYRPPDISLKPLLFEVPSITTESVFVGRDWVFHEIDAQLQSANASVNRGVVIVGNIGYGKTAIISRLVALSCHGTRMRQIASDSPHASPKHVDTNRELPLTQPPSAHSSITSGSCPGTPEMRRRQEEAMRRLAAQVVAYHYCQADNAYTCLVPEFVHNVAALLCRSPQFVAYREQLLREPHLQSMLSLRSCVQDPMASFRRGVLEPLENLHNERKIPDEDYIILIDGLNEAEFHKPDYGDTIVSFLSKMIGKFPSWLKLIVTVRTSLQEIIKLLPFHRIFLDRLEENEAIDQDLQAYILHRIHSSSEIQNNISLNGKMDNTTFGKLSSHLKTLSQGSYLYLKLTFDLIEKGYLVLKSSSYKVVPVSLSEVYLLQCNMKFPTQSSFDRVMPLLNVAVASLHPLTDEHIFQAINAGSIEGTLEWDDFQQRMENLSMFLIKRRDMTRMFVHPSFREWLIWREEGEKTKFLCDPRSGHTLLAFWFSRQEGKLNRQQTIELGHHILKAHIFKGLSKKVGVSSSILQGLWISYSTEGLSMALASLRNLYTPNIKVSRLLILGGANVNYRTEVLNNAPILCVQSHLGYTEMVALLLEFGANVDASSESGLTPLGYAAAAGYLSIVMLLCKKRAQVDHLDKNGQCALVHAALRGHLEVVKFLIQCDWTMAGQQQGVFKKSHAIQQALIAAASMGYTEVERAQINSFDSLWGETALTAAAGRGKLEVCRLLLEQGAAVAQPNRRGVVPLFSAVRQGHWQIVDLLLTHGADVNMADKQGRTPLMMAASEGHLGTMEFLLAQGGSISLMDKEGLTALSWACLKGHLSVVRSLVESGAATDHADKNGRTPLDLAAFYGDAEVVSFLVDHGAMIEHVDYSGMRPLDRAVGCRNTSVVVTLLKKGAKIGPATWAMATSKPDIMIILLSKLMEEGDMFYKKGKVKEAAQRYQYALKKFPREGFGEDLKTFRELKVSLLLNLSRCRRKMNDFGMAEEFATKALELKPKSYEAYYARARAKRSSRQFSAALEDLNEAIKLCPNNREIQRLLMRVEEECRQMQQQQQQQQPPVEPEPEAQHEELYSVQDLFEEEYLEQDVENVSIGLQTESRPSQGLPIIQSPPPSPAHRDSAYISGSPLGSHQVFDFRSNSSVGSPTRQGYQSTSPALSPTHQNNHYRPSPPHTSPAHQGSSYRFSPPPVAGQGKEYQSPPPSPLRRGPQYRASPPAESISVYRSQSGSPVRYQQEPSVSQLPGRPKSPLSKMTQRPLLHSQSVGLRFSPSSNSISSTSNLTPTFRPSASIQQMEIPLKPAYDRSCDELSPVSPPQGSYPSEPARSRTTPFMGIIDKTARTQQYPHLHQQTRTWAVSSVDTVISPTSPGNLPQPESFSPPSSISNIAFYNKTNNAQNGHLLEEDYYSPHGMLANGSRGDLLERVSQASSYPDVKVARTLPVAQAYQDNLYRQLSRDSRQGQTSPIKPKRPFVESNV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXZ0</accession>
    <name>A0A452GXZ0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">acetyl-CoA carboxylase</fullName>
            <ecNumber evidence="3">6.4.1.2</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>biotin</name>
            <dbReference type="ChEBI" id="CHEBI:57586"/>
        </cofactor>
    </comment>
    <comment type="pathway">
        <text evidence="2">Lipid metabolism; malonyl-CoA biosynthesis; malonyl-CoA from acetyl-CoA: step 1/1.</text>
    </comment>
    <dbReference type="EC" id="6.4.1.2" evidence="3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008152.1">
        <property type="protein sequence ID" value="ENSGAGP00000007036.1"/>
        <property type="gene ID" value="ENSGAGG00000004854.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00655">
        <property type="reaction ID" value="UER00711"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003989">
        <property type="term" value="F:acetyl-CoA carboxylase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006633">
        <property type="term" value="P:fatty acid biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:2001295">
        <property type="term" value="P:malonyl-CoA biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="CDD" id="cd06850">
        <property type="entry name" value="biotinyl_domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.460.10:FF:000001">
        <property type="entry name" value="Acetyl-CoA carboxylase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.50.100:FF:000005">
        <property type="entry name" value="Acetyl-CoA carboxylase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.470.20:FF:000005">
        <property type="entry name" value="Acetyl-CoA carboxylase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1770.10:FF:000001">
        <property type="entry name" value="acetyl-CoA carboxylase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1490.20:FF:000003">
        <property type="entry name" value="acetyl-CoA carboxylase isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.20:FF:000005">
        <property type="entry name" value="acetyl-CoA carboxylase isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.226.10:FF:000010">
        <property type="entry name" value="acetyl-CoA carboxylase isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.50.100">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.226.10">
        <property type="entry name" value="2-enoyl-CoA Hydratase, Chain A, domain 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1490.20">
        <property type="entry name" value="ATP-grasp fold, A domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.470.20">
        <property type="entry name" value="ATP-grasp fold, B domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.460.10">
        <property type="entry name" value="Biotin dependent carboxylase carboxyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1770.10">
        <property type="entry name" value="PreATP-grasp domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049076">
        <property type="entry name" value="ACCA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049074">
        <property type="entry name" value="ACCA_BT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034733">
        <property type="entry name" value="AcCoA_carboxyl_beta"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013537">
        <property type="entry name" value="AcCoA_COase_cen"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011761">
        <property type="entry name" value="ATP-grasp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013815">
        <property type="entry name" value="ATP_grasp_subdomain_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005481">
        <property type="entry name" value="BC-like_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011764">
        <property type="entry name" value="Biotin_carboxylation_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005482">
        <property type="entry name" value="Biotin_COase_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000089">
        <property type="entry name" value="Biotin_lipoyl"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029045">
        <property type="entry name" value="ClpP/crotonase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011763">
        <property type="entry name" value="COA_CT_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011762">
        <property type="entry name" value="COA_CT_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005479">
        <property type="entry name" value="CPAse_ATP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016185">
        <property type="entry name" value="PreATP-grasp_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011054">
        <property type="entry name" value="Rudment_hybrid_motif"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011053">
        <property type="entry name" value="Single_hybrid_motif"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45728:SF1">
        <property type="entry name" value="ACETYL-COA CARBOXYLASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45728">
        <property type="entry name" value="ACETYL-COA CARBOXYLASE, ISOFORM A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08326">
        <property type="entry name" value="ACC_central"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21385">
        <property type="entry name" value="ACCA_BT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02785">
        <property type="entry name" value="Biotin_carb_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00289">
        <property type="entry name" value="Biotin_carb_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00364">
        <property type="entry name" value="Biotin_lipoyl"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01039">
        <property type="entry name" value="Carboxyl_trans"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02786">
        <property type="entry name" value="CPSase_L_D2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00878">
        <property type="entry name" value="Biotin_carb_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52096">
        <property type="entry name" value="ClpP/crotonase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56059">
        <property type="entry name" value="Glutathione synthetase ATP-binding domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52440">
        <property type="entry name" value="PreATP-grasp domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51246">
        <property type="entry name" value="Rudiment single hybrid motif"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51230">
        <property type="entry name" value="Single hybrid motif"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50975">
        <property type="entry name" value="ATP_GRASP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50979">
        <property type="entry name" value="BC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50968">
        <property type="entry name" value="BIOTINYL_LIPOYL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50989">
        <property type="entry name" value="COA_CT_CTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50980">
        <property type="entry name" value="COA_CT_NTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00866">
        <property type="entry name" value="CPSASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00867">
        <property type="entry name" value="CPSASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8 13" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="11" id="KW-0092">Biotin</keyword>
    <keyword evidence="10" id="KW-0275">Fatty acid biosynthesis</keyword>
    <keyword evidence="7" id="KW-0276">Fatty acid metabolism</keyword>
    <keyword evidence="5" id="KW-0436">Ligase</keyword>
    <keyword evidence="4" id="KW-0444">Lipid biosynthesis</keyword>
    <keyword evidence="9" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="12" id="KW-0511">Multifunctional enzyme</keyword>
    <keyword evidence="6 13" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Biotin carboxylation" evidence="16">
        <location>
            <begin position="47"/>
            <end position="548"/>
        </location>
    </feature>
    <feature type="domain" description="ATP-grasp" evidence="15">
        <location>
            <begin position="205"/>
            <end position="396"/>
        </location>
    </feature>
    <feature type="domain" description="Lipoyl-binding" evidence="14">
        <location>
            <begin position="675"/>
            <end position="749"/>
        </location>
    </feature>
    <feature type="domain" description="CoA carboxyltransferase N-terminal" evidence="17">
        <location>
            <begin position="1479"/>
            <end position="1809"/>
        </location>
    </feature>
    <feature type="domain" description="CoA carboxyltransferase C-terminal" evidence="18">
        <location>
            <begin position="1813"/>
            <end position="2129"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001953"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004956"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00013058"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022516"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022598"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022832"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023160"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023267"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023268"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00409"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50975"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50979"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50980"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007036.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2175" mass="246088" checksum="57D3CEAA45992479" modified="2019-05-08" version="1">DINLTLPSMSGLHLVKRGREQKKLELQRDFTVASPAEFVTRFGGNRIIEKVLIANNGIAAVKCMRSIRRWAYEMFRNERAIRFVVMVTPEDLKANAEYIKMADHYVPVPGGANNNNYANVELIVDISKRIPVQAVWAGWGHASENPRLPELLQKHGVAFLGPPSEAMWALGDKVASTIVAQTVQIPTLPWSGSGLMAEWTEDCEQRRTISVSLETYARGCVKDVDESLEAAERIGYPVMIKASEGGGGKGIRRAETAEEFTGCFRQVQSEAPGSPIFVMKLAQHARHLEVQVLADQYGNAVSLFGRDCSLQRRHQKIIEEAPVTIAAPSVFDFMEQCAVRLAKMVGYVSAGTVEYLYSEDGGFHFLELNPRLQVEHPCTEMIADVNLPAAQLQIAMGVPLHRIKDIRVLYGESPWADTPICFESPANIPVPRGHVIAARITSENPDEEFKPSSGTVQELNFRSSKNVWGYFSVAAAGGLHEFADSQFGHCFSWGENREEAISNMVVALKELSIRGDFRTTVEYLIKLLETESFQNNEIDTGWLDHLIAEKVQAEKPDTMLGVVCCALNVADALFRTCMTDFLHSLERGQVLPAASLLNIVEVELIYEGVKYALKVARQSLTTYVIIMNNTHIEIDVHRLNDGGVLLSYDGNSYTTYMKEEVDCYRITVGNKTCYFEKEKDPTMLRSPSAGKLLKYTVEDGGHIFIGHSFAEIEVMKIIFTLEVKESGHIRYIKRPGALLEAGCVIARLELDDPTKVHPAQLYTGGLPTQQMLPIGGLKLHQVFHSVLENLINVMNGYCLPEPYFSAKMKEWVCELMKTLRDPSLPLLELQEIMTNVSGRIPLSVEKSIRKVMAQYASNITSVLCQFPSQQIASILDSYAASLQRKADREVFFMNTQSIVQLVQRYRSGIRGYMKSVVLDLLRRYLQVETQFQQAHYDKCVINLREQYKLDMNPVLECIFSHAQVAKKNLLVIMLIDQLCGRDSTMTEELMAILTELTQLSKTEHSKVALRARQVLIASHLPSYELRHNQVESIFLSAIDMYGHQFCPEDLKKLILSETTIFDVLPTFFYHSNQLVRMAALEVYVRRGYIAYELNSLQHRQLSDGTCVVEFQFMLPSSHPNRMSIPISVSNPDLARHSTELFMDSGFSPLCQRMGAMVAFDRFEDFTRNFDEVISCFADPPSESPLFSEAQSTLYEEGDNKTIQEEPIHILNTAIRWAGHLEDEELVPVFRTFAQSKKNILVDCGLRRITFLIVQKVSLLSSLVRTQFAEDRIYRHLEPALAFQLELNRMRNFDLTAVPCANHRMHLYLGAAKVEEGNEVTDYRFFIRAIIRHSDLITKEASFEYLQNEGERLLLEAMDELEVAFSNTSVRTDCNHIFLNFVPTVIMDPSKIEEAVRSMVMRYGSRLWKLRVLQAEVKINIRLTPVGKAIPIRLFLTNESGYYLDISLYKEVNDPRTGNIMFHSYGDKQGLQHGMLINTPYVTKDLLQSKRFQAQSLGTTYVYDFPEMFKQALFKLWGSTEMYPKDVLTYTELVLDSQGQLVEMNRIPGGNEVGMVAFKMKLKTPEYPKGRDMIVISNDITCKIGSFGPEEDLLFLRASQLARSEGIPRIYIAANSGARIGFADEKKHMFQVAWVDPMDPYKGFKYLYLTPQDYTRINSVNSVHCEHVEEGGESRYVLTDIIGKDEGFGVENLRASGTIAGESSIAYDEIVTISMVTCRAIGIGAYLVRLGQRVIQVENSHIILTGASALNKVLGREVYTSNNQLGGVQIMHNNGVSHVTVPDDFEGIYTILQWLSYMPKDNRSPVPVTATSDPIEREIDFVPSKGPYDPRWMLAGRPHPIVKGTWQSGFFDHGTFMEIMQPWAQTVVVGRARLGGIPIGVVAVETRTVEMAIPADPANLDSEAKIVQQAGQVWFPDSAFKTAQAIRDFDRERLPLLIFANWRGFSGGMKDMYDQVLKFGAYIVDSLRQFKQPVLVYIPPHAELRGGSWVVIDPTINPVYMELYADKESRGGILEAEGTVEIKFRRKDLIKTMRRIDSVCARLVEQLGTPELLEEDHRELKKQLKAREELLLPIFHQVAVQFADLHDTPGRMHEKGVIADILEWKNARTFLYWRLRRLLLEEVVKTEILNADSELSNIHIQSMLRRWFMETEGAVKVSLHLENHWRASLFPCYGSV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXZ4</accession>
    <name>A0A452GXZ4_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="12">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GXZ4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008087.1">
        <property type="protein sequence ID" value="ENSGAGP00000006980.1"/>
        <property type="gene ID" value="ENSGAGG00000002943.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd08849">
        <property type="entry name" value="ArfGap_ASAP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd13251">
        <property type="entry name" value="PH_ASAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.220.150:FF:000002">
        <property type="entry name" value="arf-GAP with SH3 domain, ANK repeat and PH domain-containing protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.950:FF:000001">
        <property type="entry name" value="Arf-GAP with SH3 domain, ANK repeat and PH domain-containing protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000006">
        <property type="entry name" value="Arf-GAP with SH3 domain, ANK repeat and PH domain-containing protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000012">
        <property type="entry name" value="Arf-GAP with SH3 domain, ANK repeat and PH domain-containing protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.950">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.220.150">
        <property type="entry name" value="Arf GTPase activating protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.60">
        <property type="entry name" value="Arfaptin homology (AH) domain/BAR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027267">
        <property type="entry name" value="AH/BAR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037278">
        <property type="entry name" value="ARFGAP/RecO"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001164">
        <property type="entry name" value="ArfGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038508">
        <property type="entry name" value="ArfGAP_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043593">
        <property type="entry name" value="ASAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004148">
        <property type="entry name" value="BAR_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037844">
        <property type="entry name" value="PH_ASAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45854:SF4">
        <property type="entry name" value="ARF-GAP WITH SH3 DOMAIN, ANK REPEAT AND PH DOMAIN-CONTAINING PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45854">
        <property type="entry name" value="ASAP FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01412">
        <property type="entry name" value="ArfGap"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16746">
        <property type="entry name" value="BAR_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00169">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00405">
        <property type="entry name" value="REVINTRACTNG"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00105">
        <property type="entry name" value="ArfGap"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00233">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57863">
        <property type="entry name" value="ArfGap/RecO-like zinc finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103657">
        <property type="entry name" value="BAR/IMD domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50115">
        <property type="entry name" value="ARFGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6 7" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <keyword evidence="8" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="PH" evidence="10">
        <location>
            <begin position="148"/>
            <end position="240"/>
        </location>
    </feature>
    <feature type="domain" description="Arf-GAP" evidence="11">
        <location>
            <begin position="264"/>
            <end position="386"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="7">
        <location>
            <begin position="427"/>
            <end position="462"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="7">
        <location>
            <begin position="463"/>
            <end position="495"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="131"/>
            <end position="155"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="543"/>
            <end position="564"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="658"/>
            <end position="688"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="131"/>
            <end position="143"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="9">
        <location>
            <begin position="667"/>
            <end position="688"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50115"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006980.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="688" mass="77165" checksum="75A30933B845E07A" modified="2019-05-08" version="1">MLKLSHTHNAAGQGNAKIRRWMKNRVFPHPPFHDLYIYLLELEGTLEVNEIKIKKGVDLLQNLIKYFHAQCNFFQDGLKAVESLKPSIETLSTDLHTIKQAQDEERKQLIHLRDILKSALQVEQKEDSQIRQSTAYSLHQPQGNKEHGTERSGSLYKKSDGIRKVWQKRKCSVKNGFLTISHGTANRPPAKLNLLTCQVKTNPEEKKCFDLISHDRTYHFQAEDEQECQIWTSVLQNSKEEALNNAFKGDENTGENNIVQELTKEIISEVQRMTGNDVCCDCGAQDPSWLSTNLGILTCIECSGIHRELGVHYSRMQSLTLDVLGTSELLLAKNIGNAGFNEIMEFCLPADETIKPNPSSDMNARKDYITAKYFERKYARKKHVDNAAKLYSLCEAVKARDIFGLVQVYADGVDLTEKIPLANGHEQDETALHLAVRSVDRTSLHIVDFLVQNSGNLDKQTYKGSTALHYCCLTDNVECLKLLLRGKASIEIANEAGETPLDIAKRLKHTHCEELLTQALSGRFNSHVHVEYEWRLLHEDLDESDDDVDEKLQQPSPNRREDRPVSFYQLGSNQLQPNVASLARDAANIAKDKQRSFMPSILQNETYGAILSGSPPPIQPAVPVTTSAPPLPPRNIGKDILFPAPPPPVAKTASVIEALNQQSKQPPAVPQSKPIPPPLPPQPPIFNS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXZ7</accession>
    <name>A0A452GXZ7_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="13">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="6">Postsynapse</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="7">Belongs to the TANC family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000008096.1">
        <property type="protein sequence ID" value="ENSGAGP00000006988.1"/>
        <property type="gene ID" value="ENSGAGG00000005019.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043197">
        <property type="term" value="C:dendritic spine"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061001">
        <property type="term" value="P:regulation of dendritic spine morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000022">
        <property type="entry name" value="protein TANC2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000036">
        <property type="entry name" value="protein TANC2 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000044">
        <property type="entry name" value="Tetratricopeptide repeat, ankyrin repeat and coiled-coil containing 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.10">
        <property type="entry name" value="Tetratricopeptide repeat domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058018">
        <property type="entry name" value="AAA_lid_TANC1/2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050889">
        <property type="entry name" value="Dendritic_Spine_Reg/Scaffold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011990">
        <property type="entry name" value="TPR-like_helical_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019734">
        <property type="entry name" value="TPR_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058056">
        <property type="entry name" value="WH_TANC1/2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24166:SF21">
        <property type="entry name" value="PROTEIN TANC2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24166">
        <property type="entry name" value="ROLLING PEBBLES, ISOFORM B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25520">
        <property type="entry name" value="AAA_lid_TANC1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00023">
        <property type="entry name" value="Ank"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13181">
        <property type="entry name" value="TPR_8"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25521">
        <property type="entry name" value="WHD_TANC1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01415">
        <property type="entry name" value="ANKYRIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="SMART" id="SM00028">
        <property type="entry name" value="TPR"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48452">
        <property type="entry name" value="TPR-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50005">
        <property type="entry name" value="TPR"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 8" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="1" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0770">Synapse</keyword>
    <keyword evidence="3 9" id="KW-0802">TPR repeat</keyword>
    <feature type="domain" description="TANC1/2-like AAA+ ATPase lid" evidence="11">
        <location>
            <begin position="394"/>
            <end position="488"/>
        </location>
    </feature>
    <feature type="domain" description="TANC1/2-like winged helix" evidence="12">
        <location>
            <begin position="490"/>
            <end position="642"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="8">
        <location>
            <begin position="713"/>
            <end position="745"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="8">
        <location>
            <begin position="829"/>
            <end position="861"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="8">
        <location>
            <begin position="862"/>
            <end position="894"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="8">
        <location>
            <begin position="895"/>
            <end position="927"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="8">
        <location>
            <begin position="928"/>
            <end position="960"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="8">
        <location>
            <begin position="961"/>
            <end position="993"/>
        </location>
    </feature>
    <feature type="repeat" description="TPR" evidence="9">
        <location>
            <begin position="1040"/>
            <end position="1073"/>
        </location>
    </feature>
    <feature type="repeat" description="TPR" evidence="9">
        <location>
            <begin position="1087"/>
            <end position="1120"/>
        </location>
    </feature>
    <feature type="repeat" description="TPR" evidence="9">
        <location>
            <begin position="1121"/>
            <end position="1154"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="192"/>
            <end position="239"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1171"/>
            <end position="1190"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1219"/>
            <end position="1371"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1481"/>
            <end position="1507"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1610"/>
            <end position="1634"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1760"/>
            <end position="1782"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="215"/>
            <end position="227"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="1171"/>
            <end position="1180"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="1258"/>
            <end position="1287"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022803"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00034110"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00038259"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00339"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF25520"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF25521"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006988.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1782" mass="197724" checksum="B8298C688B9F8B5F" modified="2019-05-08" version="1">MGNALRKEMSAIPTMDQGDWPVGGPATFRMVHQVKRGNAGQAWSIPTYPQGILCEKEYQDLEPGLIAVCFPLAATSSAHLEDLAYLDEQRHTPLRTSLRMPRQSMAGALRFAPYRPPDISLKPLLFEVPSITTESVFVGRDWVFHEIDAQLQSANASVNRGVVIVGNIGYGKTAIISRLVALSCHGTRMRQIASDSPHASPKHVDTNRELPLTQPPSAHSSITSGSCPGTPEMRRRQEEAMRRLAAQVVAYHYCQADNAYTCLVPEFVHNVAALLCRSPQFVAYREQLLREPHLQSMLSLRSCVQDPMASFRRGVLEPLENLHNERKIPDEDYIILIDGLNEAEFHKPDYGDTIVSFLSKMIGKFPSWLKLIVTVRTSLQEIIKLLPFHRIFLDRLEENEAIDQDLQAYILHRIHSSSEIQNNISLNGKMDNTTFGKLSSHLKTLSQGSYLYLKLTFDLIEKGYLVLKSSSYKVVPVSLSEVYLLQCNMKFPTQSSFDRVMPLLNVAVASLHPLTDEHIFQAINAGSIEGTLEWDDFQQRMENLSMFLIKRRDMTRMFVHPSFREWLIWREEGEKTKFLCDPRSGHTLLAFWFSRQEGKLNRQQTIELGHHILKAHIFKGLSKKVGVSSSILQGLWISYSTEGLSMALASLRNLYTPNIKVSRLLILGGANVNYRTEVLNNAPILCVQSHLGYTEMVALLLEFGANVDASSESGLTPLGYAAAAGYLSIVMLLCKKRAQVDHLDKNGQCALVHAALRGHLEVVKFLIQCDWTMAGQQQGVFKKSHAIQQALIAAASMGYTEIVSYLLDLPEKDEEEVERAQINSFDSLWGETALTAAAGRGKLEVCRLLLEQGAAVAQPNRRGVVPLFSAVRQGHWQIVDLLLTHGADVNMADKQGRTPLMMAASEGHLGTMEFLLAQGGSISLMDKEGLTALSWACLKGHLSVVRSLVESGAATDHADKNGRTPLDLAAFYGDAEVVQFLVDHGAMIEHVDYSGMRPLDRAVGCRNTSVVVTLLKKGAKIGPATWAMATSKPDIMIILLSKLMEEGDMFYKKGKVKEAAQRYQYALKKFPREGFGEDLKTFRELKVSLLLNLSRCRRKMNDFGMAEEFATKALELKPKSYEAYYARARAKRSSRQFSAALEDLNEAIKLCPNNREIQRLLMRVEEECRQMQQQQQQQQPPVEPEPEAQHEELYSVQDLFEEEYLEQDVENVSIGLQTESRPSQGLPIIQSPPPSPAHRDSAYISGSPLGSHQVFDFRSNSSVGSPTRQGYQSTSPALSPTHQNNHYRPSPPHTSPAHQGSSYRFSPPPVAGQGKEYQSPPPSPLRRGPQYRASPPAESISVYRSQSGSPVRYQQEPSVSQLPGRPKSPLSKMTQRSFQIPQLPVAVPQQGHRLQPAKAQIVRSNQPSSAVHSSTVIPTGAYSQVAHSMASKYQSASGEMGVGQSRLVYQGSISGIVGDGRPLQHVQASLSAGAICQHGGLTKEDLPQRPSSAYRGGMRYSQTPQIGRSQSASYYPVCHSKLDLERSSLPSSQLGSPDVSHLIRRPITINPNEIKPHPPTPRPLLHSQSVGLRFSPSSNSISSTSNLTPTFRPSASIQQMEIPLKPAYDRSCDELSPVSPPQGSYPSEPARSRTTPFMGIIDKTARTQQYPHLHQQTRTWAVSSVDTVISPTSPGNLPQPESFSPPSSISNIAFYNKTNNAQNGHLLEEDYYSPHGMLANGSRGDLLERVSQASSYPDVKVARTLPVAQAYQDNLYRQLSRDSRQGQTSPIKPKRPFVESNV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXZ8</accession>
    <name>A0A452GXZ8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Glycogen debranching enzyme</fullName>
            <ecNumber evidence="5">2.4.1.25</ecNumber>
            <ecNumber evidence="6">3.2.1.33</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="18">Glycogen debrancher</fullName>
        </alternativeName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="3">Multifunctional enzyme acting as 1,4-alpha-D-glucan:1,4-alpha-D-glucan 4-alpha-D-glycosyltransferase and amylo-1,6-glucosidase in glycogen degradation.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="2">
            <text>Hydrolysis of (1-&gt;6)-alpha-D-glucosidic branch linkages in glycogen phosphorylase limit dextrin.</text>
            <dbReference type="EC" id="3.2.1.33"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>Transfers a segment of a (1-&gt;4)-alpha-D-glucan to a new position in an acceptor, which may be glucose or a (1-&gt;4)-alpha-D-glucan.</text>
            <dbReference type="EC" id="2.4.1.25"/>
        </reaction>
    </comment>
    <comment type="subunit">
        <text evidence="19">Monomer. Interacts with NHLRC1/malin.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="4">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="17">Belongs to the glycogen debranching enzyme family.</text>
    </comment>
    <dbReference type="EC" id="2.4.1.25" evidence="5"/>
    <dbReference type="EC" id="3.2.1.33" evidence="6"/>
    <dbReference type="STRING" id="38772.ENSGAGP00000006892"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007994.1">
        <property type="protein sequence ID" value="ENSGAGP00000006892.1"/>
        <property type="gene ID" value="ENSGAGG00000005555.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000008002.1">
        <property type="protein sequence ID" value="ENSGAGP00000006900.1"/>
        <property type="gene ID" value="ENSGAGG00000005555.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000008015.1">
        <property type="protein sequence ID" value="ENSGAGP00000006912.1"/>
        <property type="gene ID" value="ENSGAGG00000005555.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000008028.1">
        <property type="protein sequence ID" value="ENSGAGP00000006925.1"/>
        <property type="gene ID" value="ENSGAGG00000005555.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000008046.1">
        <property type="protein sequence ID" value="ENSGAGP00000006943.1"/>
        <property type="gene ID" value="ENSGAGG00000005555.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004134">
        <property type="term" value="F:4-alpha-glucanotransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004135">
        <property type="term" value="F:amylo-alpha-1,6-glucosidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005978">
        <property type="term" value="P:glycogen biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005980">
        <property type="term" value="P:glycogen catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd11327">
        <property type="entry name" value="AmyAc_Glg_debranch_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.20.20.80:FF:000070">
        <property type="entry name" value="GDB1p Glycogen debranching enzyme"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.50.10.10:FF:000039">
        <property type="entry name" value="Glycogen debranching enzyme Gdb1, putative"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.20.20.80:FF:000051">
        <property type="entry name" value="glycogen debranching enzyme isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.20.20.80">
        <property type="entry name" value="Glycosidases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008928">
        <property type="entry name" value="6-hairpin_glycosidase_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010401">
        <property type="entry name" value="AGL/Gdb1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032788">
        <property type="entry name" value="AGL_central"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029436">
        <property type="entry name" value="AGL_euk_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032792">
        <property type="entry name" value="AGL_glucanoTrfase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032790">
        <property type="entry name" value="GDE_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017853">
        <property type="entry name" value="GH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006421">
        <property type="entry name" value="Glycogen_debranch_met"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01531">
        <property type="entry name" value="glyc_debranch"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10569">
        <property type="entry name" value="GLYCOGEN DEBRANCHING ENZYME"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10569:SF2">
        <property type="entry name" value="GLYCOGEN DEBRANCHING ENZYME"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06202">
        <property type="entry name" value="GDE_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14701">
        <property type="entry name" value="hDGE_amylase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14702">
        <property type="entry name" value="hGDE_central"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14699">
        <property type="entry name" value="hGDE_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51445">
        <property type="entry name" value="(Trans)glycosidases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48208">
        <property type="entry name" value="Six-hairpin glycosidases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="14" id="KW-0320">Glycogen biosynthesis</keyword>
    <keyword evidence="16" id="KW-0326">Glycosidase</keyword>
    <keyword evidence="10" id="KW-0328">Glycosyltransferase</keyword>
    <keyword evidence="12" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="15" id="KW-0511">Multifunctional enzyme</keyword>
    <keyword evidence="9" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="11" id="KW-0808">Transferase</keyword>
    <keyword evidence="13" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="Eukaryotic glycogen debranching enzyme N-terminal" evidence="21">
        <location>
            <begin position="31"/>
            <end position="115"/>
        </location>
    </feature>
    <feature type="domain" description="Glycogen debranching enzyme glucanotransferase" evidence="22">
        <location>
            <begin position="121"/>
            <end position="550"/>
        </location>
    </feature>
    <feature type="domain" description="Glycogen debranching enzyme central" evidence="23">
        <location>
            <begin position="696"/>
            <end position="973"/>
        </location>
    </feature>
    <feature type="domain" description="Glycogen debranching enzyme C-terminal" evidence="20">
        <location>
            <begin position="1072"/>
            <end position="1526"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000439"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00000927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00003530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012560"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00012778"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00020723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022676"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023268"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023295"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00025780"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00031477"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00063438"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="Pfam" id="PF06202"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="Pfam" id="PF14699"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="Pfam" id="PF14701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="Pfam" id="PF14702"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006925.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1531" mass="174272" checksum="DEF0292372A4B9D4" modified="2019-05-08" version="1">MPPSNQTRVLLLSDMEKQEKTLFRLEQGFELQFRLGPTLQGKHVTVYTNYPASGELFCRRKFRALSWHNPTGREDDSDKYCKLDLQISGSYQYYFSHENEKSGGGYIVVDPILRVGADNQVLHLDCVTLQTFLAKCLGPFEEWEDRLKVAKESGYNMIHFTPLQKLGLSRSCYSLADQLEVNPDFSSPNKKCTWSNIGNLVEKMKNEWNMLCITDVVYNHTAANSEWIRVHPECGYNLVNSPHLKPAWILDRALCHLSCSVAEGKYRDKGLPSSIENDHHLNCIRKIIWEDIYPRIKLWEFFQVDVNKAVQQFKTLLTQGNRKTKSDPNQHLQRIQDPEYKRLGCTVDMNIALATFIPHSNGPAAIEECCNWFQKRIEELNAEQFQQTTYHQEQAVTCLLETVVYERLADHGPKQGPVSRKYPLVTRYFTYPFKEMTLEEEELLMHQPDKACHFLAHNGWVMGDDPLRNFAEPGSNVYLRRELICWGDSVKLRYGNKPEDCPYLWAHMKKYTEITAKYFHGVRLDNCHSTPLHVAEDMLAAARSVRPNLYVIAELFTGSERIDNVFVNRLGITSLIREAMSAYNSHEEGRLVYRFGGEPVGSFVQPRLRPLMPAIAHALFMDVTHDNECPIQHRSEYDALPSSVIVSMACCATGSTKGYDELVPHQISVVSEERFYAKWNPAATPSTPGEVNFQTGIIAGRQAMNRLHQELGAKGFIQVYVDQVDEDIVAVTRHCPNTHQSVVAVSRTAFRDPKTSSYSKEVPQMCIPGKIEEVVLEARTIERNVRPYKKDEHFINGLPNYTVEIREHIQIRESKIIKQAGTAIKGPNEFVQEIEFENLTPGSVIIFRVSLDPKAQEAVGVLRNHLIQFSPHFKAGSLPDDHSAPILKNLFSSIASKLTLADLNQVLYRCDAEEQEDGGGCYNIPDWSALKYAGLQGLMSVMADIRPKNDLGHPFCGNLRSGDWMIDYVSDRLISRAGACAEVGKWLKAMFVYLKQIPRYLIPCYFDAILVGAYTTLLDLAWSQMSSFVQNGSTFVKHLSLGSVQMCGIGKYPSLPNLSPTLHDVPYRMNEITKEREQCCVSLAAGLPHFSSGIFRCWGRDTFIALRGLMLITGRYLEARNIILAFASTMRHGLIPNLLGQGTHARFNCRDAVWWWLQCIQDYCKIVPNGTDILKCPVSRMYPTDDSPAQPAGTLDQPLYEVIQEVMQQHMQGIEFRERNAGPQLDRNMRDEGFNVTVGVDHETGFVFGGNRFNCGTWMDKMGESDRARNKGIPATPRDGSAVEIVGLCKSTIRWLLELSKKKVFPYQGVTIKRHGKKETITYDEWNGKIQGHFEKLFFVSENPADPNEKHPNLVHKRGIYKDSYGASSPWCDYQLRPNFTIAMVVAPEIFSPERAWKALEIAEEKLLGPLGMKTLDPDDMVYCGVYDNALDSDNYNVAKGFNYHQGPEWLWPIGYFLRAKLYFSRLIGPEMYAKTVVLIKNVLSRHYVHLERSSWKGLPELTNENGQYCPFSCESQAWSIAVILEVLYDL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GY60</accession>
    <name>A0A452GY60_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">receptor protein-tyrosine kinase</fullName>
            <ecNumber evidence="3">2.7.10.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="25 26">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="26" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="25" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="25" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="20 27">
            <text>L-tyrosyl-[protein] + ATP = O-phospho-L-tyrosyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10596"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.10.1"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="2.7.10.1" evidence="3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008237.1">
        <property type="protein sequence ID" value="ENSGAGP00000007115.1"/>
        <property type="gene ID" value="ENSGAGG00000005698.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009925">
        <property type="term" value="C:basal plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043235">
        <property type="term" value="C:receptor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005154">
        <property type="term" value="F:epidermal growth factor receptor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004714">
        <property type="term" value="F:transmembrane receptor protein tyrosine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0038130">
        <property type="term" value="P:ERBB4 signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043066">
        <property type="term" value="P:negative regulation of apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0022008">
        <property type="term" value="P:neurogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008284">
        <property type="term" value="P:positive regulation of cell population proliferation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009966">
        <property type="term" value="P:regulation of signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd00064">
        <property type="entry name" value="FU"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="CDD" id="cd12092">
        <property type="entry name" value="TM_ErbB4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.220.10:FF:000001">
        <property type="entry name" value="Receptor protein-tyrosine kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.220.10:FF:000004">
        <property type="entry name" value="Receptor protein-tyrosine kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000044">
        <property type="entry name" value="Receptor protein-tyrosine kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.20.20:FF:000003">
        <property type="entry name" value="Receptor protein-tyrosine kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.880">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.220.10">
        <property type="entry name" value="Hormone Receptor, Insulin-like Growth Factor Receptor 1, Chain A, domain 2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.20.20">
        <property type="entry name" value="Receptor L-domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006211">
        <property type="entry name" value="Furin-like_Cys-rich_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006212">
        <property type="entry name" value="Furin_repeat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032778">
        <property type="entry name" value="GF_recep_IV"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009030">
        <property type="entry name" value="Growth_fac_rcpt_cys_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000494">
        <property type="entry name" value="Rcpt_L-dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036941">
        <property type="entry name" value="Rcpt_L-dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050122">
        <property type="entry name" value="RTK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001245">
        <property type="entry name" value="Ser-Thr/Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049328">
        <property type="entry name" value="TM_ErbB1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416:SF90">
        <property type="entry name" value="RECEPTOR TYROSINE-PROTEIN KINASE ERBB-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416">
        <property type="entry name" value="TYROSINE-PROTEIN KINASE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00757">
        <property type="entry name" value="Furin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14843">
        <property type="entry name" value="GF_recep_IV"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07714">
        <property type="entry name" value="PK_Tyr_Ser-Thr"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01030">
        <property type="entry name" value="Recep_L_domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21314">
        <property type="entry name" value="TM_ErbB1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00261">
        <property type="entry name" value="FU"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57184">
        <property type="entry name" value="Growth factor receptor domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52058">
        <property type="entry name" value="L domain-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="11 21" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="16" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="18" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10" id="KW-0418">Kinase</keyword>
    <keyword evidence="14 23" id="KW-0472">Membrane</keyword>
    <keyword evidence="9 21" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="19" id="KW-0539">Nucleus</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="17" id="KW-0675">Receptor</keyword>
    <keyword evidence="26" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0808">Transferase</keyword>
    <keyword evidence="7 23" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="13 23" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="15" id="KW-0829">Tyrosine-protein kinase</keyword>
    <keyword evidence="12" id="KW-0832">Ubl conjugation</keyword>
    <feature type="transmembrane region" description="Helical" evidence="23">
        <location>
            <begin position="497"/>
            <end position="520"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="24">
        <location>
            <begin position="563"/>
            <end position="801"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="739"/>
            <end position="801"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="22">
        <location>
            <begin position="775"/>
            <end position="801"/>
        </location>
    </feature>
    <feature type="binding site" evidence="21">
        <location>
            <position position="596"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00011902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023137"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00051243"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007115.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="801" mass="89324" checksum="5C992E09DBE93267" modified="2019-05-08" version="1">CYADTIHWQDIVRNPRASNFTLVPTNGSSGCGRCHKSCTGRCWGPTENHCQSLTKTVCAEQCDGRCYGPYVSDCCHRECAGGCSGPKDTDCFACMNFNDSGACVTQCPQTFVYNPTTFQLEHNHNAKYTYGAFCVKKCPHNFVVDSSSCVRACPSSKMEVEENGIKMCKPCTDICPKACDGIGTGSLTSAQTVDSSNIDKFINCTKINGNLIFLVTGIHGDPYHTIAAMDPEKLNIFQTVREITGFLNIQSWPQNMTDFRVFSNLVTIGGRALYSGLSLLILKQQAITSLQFQSLKQISAGNIYITDNTNLCYYHTINWTSLFSTPNQKTVIHHNKKPENCTAEGKVCNNLCSSDGCWGPGPDQCLSCKRFSRGRTCIESCNLYDGEFREFANGSVCVECDPQCEKMEDNMITCYGXGPDHCTKCFHFKDGPNCVEKCPDGLQGANSFIFKYANEDRECHPCHPNCTQGCRGPTSHDCIYYPWTRQSTSPQHARTPLIAAGVIGGLFIIIIVGLTFAVYVRRKSIKKKRALRRFLETELVEPLTPSGTAPNQAQLRILKETELKRVKVLGSGAFGTVYKGIWVPEGETVKIPVAIKILNETTGPKANVEFMDEALIMASMDHPHLVRLLGVCLSPTIQLVTQLMPHGCLLDYVHEHKDNIGSQLLLNWCVQIAKVRSYSFLDVSRTKQMNRVISESQQYPFLCRKQERKIYSPKAIWVRIPTHRNPNPSSKCLIHWNPKPREGIPTHKNPKPNSKCPIHRNPNPRAGIRIDRNPKFSSKCPTHWNPNPNSKCPTHTNPNPS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GY65</accession>
    <name>A0A452GY65_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">EF-hand domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GY65"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008242.1">
        <property type="protein sequence ID" value="ENSGAGP00000007120.1"/>
        <property type="gene ID" value="ENSGAGG00000005720.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005743">
        <property type="term" value="C:mitochondrial inner membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061891">
        <property type="term" value="F:calcium ion sensor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00051">
        <property type="entry name" value="EFh"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.238.10">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011992">
        <property type="entry name" value="EF-hand-dom_pair"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002048">
        <property type="entry name" value="EF_hand_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040365">
        <property type="entry name" value="EFHD1/2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13025">
        <property type="entry name" value="EF-HAND DOMAIN-CONTAINING PROTEIN D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13025:SF5">
        <property type="entry name" value="EF-HAND DOMAIN-CONTAINING PROTEIN D1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13499">
        <property type="entry name" value="EF-hand_7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47473">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50222">
        <property type="entry name" value="EF_HAND_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0106">Calcium</keyword>
    <keyword evidence="4" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="1" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="EF-hand" evidence="5">
        <location>
            <begin position="5"/>
            <end position="29"/>
        </location>
    </feature>
    <feature type="domain" description="EF-hand" evidence="5">
        <location>
            <begin position="30"/>
            <end position="65"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="4">
        <location>
            <begin position="106"/>
            <end position="133"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50222"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007120.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="143" mass="16316" checksum="9015DA7534BE420D" modified="2025-10-08" version="2">MFRVMYDSGKDGYIDLMELKLMMEKLGAPQTHLGLKNMIKEVDEDFDGKLSFREFLLIFHKAAAGELEEDSGLMTLAKLSEIDVSTEGVKGAKNFFEAKVQALSSASKFEAEIKAEQDERKREEEEKKHRRAAFRELKSAFSQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GY67</accession>
    <name>A0A452GY67_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Arpin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the Arpin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GY67"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008224.1">
        <property type="protein sequence ID" value="ENSGAGP00000007102.1"/>
        <property type="gene ID" value="ENSGAGG00000005693.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051126">
        <property type="term" value="P:negative regulation of actin nucleation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018889">
        <property type="entry name" value="Arpin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31199">
        <property type="entry name" value="ARPIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31199:SF1">
        <property type="entry name" value="ARPIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10574">
        <property type="entry name" value="UPF0552"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="1"/>
            <end position="26"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="196"/>
            <end position="226"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00008453"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00019314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007102.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="226" mass="25280" checksum="967CAD044039C839" modified="2019-05-08" version="1">MSRPPEERGLRARPVHSERLPGRWDPPALQGGNGVILEGELLDVSRHLILDINGRKERYYVLYIRPSRVHRRNFDAKGNEREPNFSDTRKVNTGFLMSSFKVEAKGDTDRLSPEELKGLVNKPELLSVTERHTPSQAVAFWLLETELEKMELELGTEIRLKTQGDSPFIFSLAKLHAGTVTKCNFVGDGQAGASWTDNILANRPPGGPAPEPRGQGDGAEEDEWDD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GY92</accession>
    <name>A0A452GY92_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="19">Nitric oxide synthase 3</fullName>
            <ecNumber evidence="6">1.14.13.39</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="16">Constitutive NOS</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="22">EC-NOS</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="20">NOS type III</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="21">Nitric oxide synthase, endothelial</fullName>
        </alternativeName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>2 L-arginine + 3 NADPH + 4 O2 + H(+) = 2 L-citrulline + 2 nitric oxide + 3 NADP(+) + 4 H2O</text>
            <dbReference type="Rhea" id="RHEA:19897"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:15379"/>
            <dbReference type="ChEBI" id="CHEBI:16480"/>
            <dbReference type="ChEBI" id="CHEBI:32682"/>
            <dbReference type="ChEBI" id="CHEBI:57743"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="EC" id="1.14.13.39"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="17">
            <dbReference type="Rhea" id="RHEA:19898"/>
        </physiologicalReaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="2">
            <name>(6R)-L-erythro-5,6,7,8-tetrahydrobiopterin</name>
            <dbReference type="ChEBI" id="CHEBI:59560"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="4">
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>FMN</name>
            <dbReference type="ChEBI" id="CHEBI:58210"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="3">
            <name>heme b</name>
            <dbReference type="ChEBI" id="CHEBI:60344"/>
        </cofactor>
    </comment>
    <comment type="subunit">
        <text evidence="18">Homodimer. Interacts with NOSIP and NOSTRIN. Interacts with HSP90AB1. Forms a complex with ASL, ASS1 and SLC7A1; the complex regulates cell-autonomous L-arginine synthesis and citrulline recycling while channeling extracellular L-arginine to nitric oxide synthesis pathway.</text>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the NOS family.</text>
    </comment>
    <dbReference type="EC" id="1.14.13.39" evidence="6"/>
    <dbReference type="AlphaFoldDB" id="A0A452GY92"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008249.1">
        <property type="protein sequence ID" value="ENSGAGP00000007127.1"/>
        <property type="gene ID" value="ENSGAGG00000000423.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010181">
        <property type="term" value="F:FMN binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004517">
        <property type="term" value="F:nitric-oxide synthase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006809">
        <property type="term" value="P:nitric oxide biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.440.10:FF:000001">
        <property type="entry name" value="Endothelial nitric oxide synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1230.10:FF:000001">
        <property type="entry name" value="Nitric oxide synthase, brain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.360">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.340.10">
        <property type="entry name" value="Nitric Oxide Synthase, Chain A, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1230.10">
        <property type="entry name" value="Nitric Oxide Synthase, Chain A, domain 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.440.10">
        <property type="entry name" value="Nitric Oxide Synthase,Heme Domain,Chain A domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001094">
        <property type="entry name" value="Flavdoxin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008254">
        <property type="entry name" value="Flavodoxin/NO_synth"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029039">
        <property type="entry name" value="Flavoprotein-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050607">
        <property type="entry name" value="NOS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044943">
        <property type="entry name" value="NOS_dom_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044940">
        <property type="entry name" value="NOS_dom_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044944">
        <property type="entry name" value="NOS_dom_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004030">
        <property type="entry name" value="NOS_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036119">
        <property type="entry name" value="NOS_N_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43410:SF1">
        <property type="entry name" value="NITRIC OXIDE SYNTHASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43410">
        <property type="entry name" value="NITRIC OXIDE SYNTHASE OXYGENASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00258">
        <property type="entry name" value="Flavodoxin_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02898">
        <property type="entry name" value="NO_synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00369">
        <property type="entry name" value="FLAVODOXIN"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52218">
        <property type="entry name" value="Flavoproteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56512">
        <property type="entry name" value="Nitric oxide (NO) synthase oxygenase domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50902">
        <property type="entry name" value="FLAVODOXIN_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="13" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="11" id="KW-0274">FAD</keyword>
    <keyword evidence="8" id="KW-0285">Flavoprotein</keyword>
    <keyword evidence="9" id="KW-0288">FMN</keyword>
    <keyword evidence="7" id="KW-0349">Heme</keyword>
    <keyword evidence="15" id="KW-0408">Iron</keyword>
    <keyword evidence="10" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="12" id="KW-0521">NADP</keyword>
    <keyword evidence="14" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Flavodoxin-like" evidence="23">
        <location>
            <begin position="314"/>
            <end position="464"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001917"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00001950"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00001970"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00001974"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00006267"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00012989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022630"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022643"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022827"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022857"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00029794"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00047419"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00049679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00049764"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00049791"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00049814"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00049818"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS50902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007127.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="464" mass="51774" checksum="5416E4B072AF17B3" modified="2019-05-08" version="1">MFGFLCTHLQYATNRGNLRSAITIFPQRMPGRGDFRIWNSQLIRYAGYPQPDGSVLGDPANVDITELCIQHGWTPGHGRFDVLPLLLQAPDEAPELFLLPPELVLEVPIQHPTLAWFGELGLRWHALPAVANMLLEIGGLQFPAAPFNGWYVGSEIGMRNFCDSNRYNLLQEVALHMGLDTGKTSSLWKDKAAVEINVAVLHSFQVAKVTIVDHHVATESFVKHMDNELQTRGGCPADWVWIVPPISGSLTPVFHQEMVNYLLSPCFRYQPDAWKVYTSKGTSITRKKTFKEVANAVKFSAKLMVHVMARRVKATILYATETGKSRTYAWNLGELFRRAFDPKVLCMDEYDIVSLEHETLVLVVTSTFGNGDPPESGESFAKALMEMSCPYVTWAHCASPPRSYKVRFNSVSQSDGLVASWKKKRKESSNTDSAGALGTLRFSVFGLGSRAYPHFCAFAPVKVL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GY99</accession>
    <name>A0A452GY99_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">CCN family member 3</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="14">Cellular communication network factor 3</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="15">Protein NOV homolog</fullName>
        </alternativeName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell junction</location>
            <location evidence="2">Gap junction</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the CCN family.</text>
    </comment>
    <comment type="caution">
        <text evidence="16">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GY99"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008283.1">
        <property type="protein sequence ID" value="ENSGAGP00000007158.1"/>
        <property type="gene ID" value="ENSGAGG00000005703.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031012">
        <property type="term" value="C:extracellular matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005921">
        <property type="term" value="C:gap junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008083">
        <property type="term" value="F:growth factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045597">
        <property type="term" value="P:positive regulation of cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051239">
        <property type="term" value="P:regulation of multicellular organismal process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000046">
        <property type="entry name" value="Cellular communication network factor 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.40.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.70.10">
        <property type="entry name" value="Complement Module, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.20.100.10">
        <property type="entry name" value="Thrombospondin type-1 (TSP1) repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050941">
        <property type="entry name" value="CCN"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006207">
        <property type="entry name" value="Cys_knot_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006208">
        <property type="entry name" value="Glyco_hormone_CN"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009030">
        <property type="entry name" value="Growth_fac_rcpt_cys_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000867">
        <property type="entry name" value="IGFBP-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012395">
        <property type="entry name" value="IGFBP_CNN"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043973">
        <property type="entry name" value="TSP1_CCN"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000884">
        <property type="entry name" value="TSP1_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036383">
        <property type="entry name" value="TSP1_rpt_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001007">
        <property type="entry name" value="VWF_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11348:SF37">
        <property type="entry name" value="CONNECTIVE TISSUE GROWTH FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11348">
        <property type="entry name" value="CONNECTIVE TISSUE GROWTH FACTOR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00007">
        <property type="entry name" value="Cys_knot"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00219">
        <property type="entry name" value="IGFBP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19035">
        <property type="entry name" value="TSP1_CCN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00093">
        <property type="entry name" value="VWC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF036495">
        <property type="entry name" value="IGFBP_rP_CNN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00041">
        <property type="entry name" value="CT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00121">
        <property type="entry name" value="IB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00209">
        <property type="entry name" value="TSP1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00214">
        <property type="entry name" value="VWC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57184">
        <property type="entry name" value="Growth factor receptor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF82895">
        <property type="entry name" value="TSP-1 type 1 repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01185">
        <property type="entry name" value="CTCK_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01225">
        <property type="entry name" value="CTCK_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51323">
        <property type="entry name" value="IGFBP_N_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50092">
        <property type="entry name" value="TSP1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01208">
        <property type="entry name" value="VWFC_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50184">
        <property type="entry name" value="VWFC_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0965">Cell junction</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="8" id="KW-0303">Gap junction</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10" id="KW-0339">Growth factor</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0964">Secreted</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
    <feature type="domain" description="IGFBP N-terminal" evidence="19">
        <location>
            <begin position="27"/>
            <end position="98"/>
        </location>
    </feature>
    <feature type="domain" description="VWFC" evidence="18">
        <location>
            <begin position="101"/>
            <end position="167"/>
        </location>
    </feature>
    <feature type="domain" description="CTCK" evidence="17">
        <location>
            <begin position="246"/>
            <end position="320"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004610"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00008125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022868"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00039944"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00042352"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00077787"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00039"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS01225"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50184"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS51323"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007158.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="344" mass="38262" checksum="2D3A3E2318BF02ED" modified="2019-05-08" version="1">LLVVSIGSVRLAGEGFYFHGLSVLQAESQLCSYPCQCPSQPLQCPVGTSHVWDACGCCKVCARQLGELCSLQTPCDHHKGLHCDFSKIHRDVGICLVHEGATCDLLGKIYHNGESFQPTCKLQCICMDGAIGCIPLCADDLQLPSPACPNPLRVRLPAQCCEEWICQGGSQETALAAPRSELNDLQENCLVQTTKWSACSRSCGMGISTRITNDNPQCRLEKESRLCMVRPCDFPREKIFKKGKKCTRTPKSRRGIRFEFSGCTSVRSYRPKFCGSCTDGRCCTPHTTSTAGVEFQCPEGDFFHRKLMFIKTCSCHHDCPRDNDIFLATYRRRMAGDHVKIERQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9U0</accession>
    <name>A0A452H9U0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Cyclin-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the cyclin family. Cyclin Y subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9U0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013208.1">
        <property type="protein sequence ID" value="ENSGAGP00000011535.1"/>
        <property type="gene ID" value="ENSGAGG00000008908.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019901">
        <property type="term" value="F:protein kinase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd20540">
        <property type="entry name" value="CYCLIN_CCNY_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.472.10:FF:000011">
        <property type="entry name" value="Cyclin-Y isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.472.10">
        <property type="entry name" value="Cyclin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013763">
        <property type="entry name" value="Cyclin-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036915">
        <property type="entry name" value="Cyclin-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006671">
        <property type="entry name" value="Cyclin_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012399">
        <property type="entry name" value="Cyclin_Y"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14248">
        <property type="entry name" value="CYCLIN Y, ISOFORM A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00134">
        <property type="entry name" value="Cyclin_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF028934">
        <property type="entry name" value="Cyclin_CG14939"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00385">
        <property type="entry name" value="CYCLIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47954">
        <property type="entry name" value="Cyclin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 3" id="KW-0195">Cyclin</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Cyclin-like" evidence="4">
        <location>
            <begin position="33"/>
            <end position="118"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00005463"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023127"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="RuleBase" id="RU000383"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="SMART" id="SM00385"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011535.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="201" mass="23451" checksum="334F7B5F994E3F25" modified="2019-05-08" version="1">LRTVTNSRHFLFQREEVPNDYGKHDPDHKHIYRFVRTLFSAAQLTAECAIVTLVYLERLLTYAEIDICPSNWKRIVLGAILLASKVWDDQAVWNVDYCQILKDITVEDMNEMERHFLELLQFNINVPASVYAKYYFDLRSLADDNNLNFLLEPLSKERAQKLEAISRLCEDKYKDLSKAAMRRSFSADNLVGIRRSNAILS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9U1</accession>
    <name>A0A452H9U1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Cathepsin O</fullName>
            <ecNumber evidence="13">3.4.22.42</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="12">Proteolytic enzyme possibly involved in normal cellular protein degradation and turnover.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>The recombinant human enzyme hydrolyzes synthetic endopeptidase substrates including Z-Phe-Arg-NHMec and Z-Arg-Arg-NHMec.</text>
            <dbReference type="EC" id="3.4.22.42"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Lysosome</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the peptidase C1 family.</text>
    </comment>
    <dbReference type="EC" id="3.4.22.42" evidence="13"/>
    <dbReference type="AlphaFoldDB" id="A0A452H9U1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013016.1">
        <property type="protein sequence ID" value="ENSGAGP00000011370.1"/>
        <property type="gene ID" value="ENSGAGG00000008781.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005764">
        <property type="term" value="C:lysosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008234">
        <property type="term" value="F:cysteine-type peptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd02248">
        <property type="entry name" value="Peptidase_C1A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.70.10:FF:000079">
        <property type="entry name" value="Cathepsin O"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.70.10">
        <property type="entry name" value="Cysteine proteinases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038765">
        <property type="entry name" value="Papain-like_cys_pep_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000169">
        <property type="entry name" value="Pept_cys_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025660">
        <property type="entry name" value="Pept_his_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013128">
        <property type="entry name" value="Peptidase_C1A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000668">
        <property type="entry name" value="Peptidase_C1A_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039417">
        <property type="entry name" value="Peptidase_C1A_papain-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12411">
        <property type="entry name" value="CYSTEINE PROTEASE FAMILY C1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00112">
        <property type="entry name" value="Peptidase_C1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00705">
        <property type="entry name" value="PAPAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00645">
        <property type="entry name" value="Pept_C1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54001">
        <property type="entry name" value="Cysteine proteinases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00139">
        <property type="entry name" value="THIOL_PROTEASE_CYS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00639">
        <property type="entry name" value="THIOL_PROTEASE_HIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="9" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="5" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="10" id="KW-0458">Lysosome</keyword>
    <keyword evidence="3" id="KW-0645">Protease</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="6" id="KW-0788">Thiol protease</keyword>
    <keyword evidence="7" id="KW-0865">Zymogen</keyword>
    <feature type="domain" description="Peptidase C1A papain C-terminal" evidence="15">
        <location>
            <begin position="69"/>
            <end position="281"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004371"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008455"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022807"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023145"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023228"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00051025"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00053492"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00066464"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00072046"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="SMART" id="SM00645"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011370.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="282" mass="31598" checksum="3EF8377C31ADFB78" modified="2019-05-08" version="1">MEEVPKMRESIKRIRSLNSLSSDDNTTAFYGINQFSHFFPEDFKAIYLKSRPYSVPRYTEVLNRKETPLPAKFDWRDKNVVTQVRNQQTCGGCWAFSVVGGIESAYAIKMKILEELSVQQVIDCSYNNYGCHGGSTVSALSWLNQTQVKLVKDSEYAFKAQTGLCHYFQSSDIGVSITDYTAYDFSGQEDKMMKKLVNCGPLAVTVDAVSWQDYLGGIIQYHCSSGKANHAVLITGFDRTGSIPYWIVRNSWGRTWGINGYAHIKIGSNVCGIADTVSSVFV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9U4</accession>
    <name>A0A452H9U4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">C2H2-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">May be involved in transcriptional regulation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9U4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013022.1">
        <property type="protein sequence ID" value="ENSGAGP00000011375.1"/>
        <property type="gene ID" value="ENSGAGG00000008773.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003700">
        <property type="term" value="F:DNA-binding transcription factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07765">
        <property type="entry name" value="KRAB_A-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000295">
        <property type="entry name" value="zinc finger protein 19"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002343">
        <property type="entry name" value="Zinc finger protein 33A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001498">
        <property type="entry name" value="Zinc finger protein 404"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002090">
        <property type="entry name" value="Zinc finger protein 473"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001270">
        <property type="entry name" value="zinc finger protein 583 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001437">
        <property type="entry name" value="Zinc finger protein 594"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000028">
        <property type="entry name" value="zinc finger protein 90 homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050589">
        <property type="entry name" value="Ikaros_C2H2-ZF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001909">
        <property type="entry name" value="KRAB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036051">
        <property type="entry name" value="KRAB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24404">
        <property type="entry name" value="ZINC FINGER PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24404:SF97">
        <property type="entry name" value="ZINC FINGER PROTEIN 350"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01352">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="SMART" id="SM00349">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109640">
        <property type="entry name" value="KRAB domain (Kruppel-associated box)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="13"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="10" id="KW-0804">Transcription</keyword>
    <keyword evidence="8" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="7" id="KW-0862">Zinc</keyword>
    <keyword evidence="6 12" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="102"/>
            <end position="129"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="130"/>
            <end position="157"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="158"/>
            <end position="185"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="186"/>
            <end position="213"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="214"/>
            <end position="241"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="242"/>
            <end position="269"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="270"/>
            <end position="297"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="298"/>
            <end position="325"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="326"/>
            <end position="353"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="354"/>
            <end position="381"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="382"/>
            <end position="409"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="410"/>
            <end position="437"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="438"/>
            <end position="465"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00003767"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011375.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="486" mass="56111" checksum="1DFBC95F8CA7D543" modified="2019-05-08" version="1">APFPHLRTFQLPQPPLCPFKAAPQRQVLSFGLLLFQEPVTFEEVAVYFTTLYRDVMQENYENVTSLGKGYCPLSKCISCRGTQKDLKKTTTQEEILMGKRKNTCTECGKNVCDYSALIKHQQLHTGERPYECSECGKNFTRRSHLINHQRIHTGERPYKCSECGKSFTQRSHLTNHQKIHIGERPYECSECGKSFTQRSHLTRHQTIHTGERPYECSECGKSFNHSSALSEHQRIHIGERAYECKECGKNFTQRSGLFLHQRIHTGERPYECCECGQTFSHSSHLSNHQTIHTGERPYECRECGKSFTSSSALSEHLRIHTGERPYECHECGKSFRFSSALSEHQRIHTGEKRYECSECGKTFGRSSCLFQHQRIHTGERPYKCIECGKRFTTSSDLSKHQRIHTGERPYKCSECGKSFNQRSTLIKHQRIHTGERPYECHECAKTFNSSSSLIRHQRIHTGERPYECSDITIISASALWFSRGKT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9U5</accession>
    <name>A0A452H9U5_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000013214.1">
        <property type="protein sequence ID" value="ENSGAGP00000011540.1"/>
        <property type="gene ID" value="ENSGAGG00000008909.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051018">
        <property type="term" value="F:protein kinase A binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008104">
        <property type="term" value="P:intracellular protein localization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008382">
        <property type="entry name" value="SPHK1-interactor_AKAP_110"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10226">
        <property type="entry name" value="A KINASE ANCHOR PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10226:SF3">
        <property type="entry name" value="A-KINASE ANCHOR PROTEIN 11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="408"/>
            <end position="458"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1189"/>
            <end position="1209"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1797"/>
            <end position="1831"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011540.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1935" mass="215793" checksum="EA745D29C20667A0" modified="2019-05-08" version="1">MDTYTRARSSQTKPKVSMKKDFTDGVLRSVKSLLHSRKELCNVSAEECLRQEEQDNFIEVTFLGFAGETNAAHMQELAAVSVELPDVLKSLHFCSLNEKEVIFLKNINKPMEISDVCKHQNHLSGVLCVMRLSPSFPRIKVDSVFTLLSKYATGVRYTMEVYSLQKHETEMSHAEDDDTNQSVSSIEDDFVTAFEHLDEDEPSKIQNAGTTYTTQNHRDAASQTIPAHCLEAIDSKIIVRSGHRKSSAKSSASLINILGLKELSSSVKNSVTTSISDPWIQRSFYKPCNSSDQRVNVLPKTLFSSSPAESSESDCSSPSPVIFLDEEGYQKSLKAKLQLPKIPVVKDGIEDSDSEVSEFFDSFDQFDELEHTLESTCKLIRDPILGNPPQKREISLEQLHSKTTTMNPQKFKFDRPTLPANVRKPTPRKPESPYSSLFDVPDSPRPMKTGEDNGGLFSPIRSSAFSPLGSCASTECLCRMDINGDGLNQNHDTLYSEYSDYANNVSFEILGSVFPSQSTSLQKHAENISSLNRIVSKEEKGQNTELQRKTSDKEPAKKVKYKHKSQMIKDSIQKFAAELVEKSFGTAFKDLQKGVSSCTNALCHLAARLTSSVFQMAFYEIGRRRALSLKERAINGLASFLVSEAITSALKEMRYVKQQIFTNTVARFAADLTEELIFEGIMEVCQFSHPSTPTTARDWAFDYEDKVVRSYAKDLSESVIQEAFIELSQVDVTFTTQAAISVSMDNIKYVSAESPLQSTQTTTAFPNFHDNALLALTPVQETGKEYTVQKALFCTSGVVSSILVPLAGTALYQHQASSDVYKAKVWTCSTAGGSLKMYKNATQPYFTTKKKEEEVASLRNIYLTADHSESTESSARCFFNQNDFKQRNNPEVNNTSDLTTGPTGINSFSGTMVDMIVNEAYEAVTSSGVTKTVEEYTDCVTRKTPRWQCVGGDIPKNTFADHLAKCIVKQSVDETKSMFSHTNETVECNGGLQMNTDNCKKELHSAVKKQESEKHRFVPIIVEPQQLPLNNPPNFLVTSVDSGQCLLSVPKDCVQERKGNEVCRFSSSTPPPCPTVTLASCNVEEFTDSGSSSIKSQSKLLKKYDMQKTTLAPSALGQESCFLHASNLSLVMFGCEDSLQMEDKSSIRDGNVCAVPNTPPPTPLVPSQTSSERNLRKLTKKLKGELAKEFAPATPPSTPYNSSAAALSATEHDSLEKEEFMLKLMRSLSEEVESSEDEEHSEMLVEKTEHSEKTVQYADHLAGHIISVATEMAASHLDDKTIKRKADRHCQLNAQNKICGYTAFINIPEETLHSLWNYAGDMAGKVINEAKKMIKSRHCKLLRLKRVNCHVDCLSLRKDDRDCRSKEKWCPVVNQWSGEVDSSVLSLPQSLGTTGLTYRYPSCESVTDEYADHIIRVLKREGGNSELIMDQYASRLAYRSIKSGLQQAARKIKLTYNRRMFPVQNTQVNGRLELFKILNKDMDTKWQMKSGVQRCGGQACEINNLHRTDGTELLHFSESLAHRITCDVRRKLKMSAVCLPKSLTDSCLYRKSKFDETTGDCLKTTLSKTVLPFSQKHKLYHSTGNLNEYGYREDIIQAIEEYARKVVDDTLEMSLESAVLQAAENRKNGDRFTYTEKLSPFSGTACRYCSMKEYQYCTGSSSPHLPVQEPLPRIRQIPNSGLGGICQKSRMFHLDIPKIHIDMEQKTVFSEKVVGAAIEKAERELSNTSLTADSGIGHGGVSFAESLTTEIMTSAMTNIGQVVNISSMGREGFHSAESVVSQQISLSIGDDSIGSWSNLSFEDEHPDESSSFLHLSDSDGTEDKDEDPKNAVEGVGQVGKALLIVNVDMEPHMVDPQLRTTLQWLAASETEVSELRFHDTATKEFILLSKRLRERGWKVGDLLQAVLKYCEMMEKMSDGDRLLNKPLFGWLLENV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9U9</accession>
    <name>A0A452H9U9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">RRM domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9U9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013220.1">
        <property type="protein sequence ID" value="ENSGAGP00000011545.1"/>
        <property type="gene ID" value="ENSGAGG00000008873.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005730">
        <property type="term" value="C:nucleolus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003729">
        <property type="term" value="F:mRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd12413">
        <property type="entry name" value="RRM1_RBM28_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12414">
        <property type="entry name" value="RRM2_RBM28_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12415">
        <property type="entry name" value="RRM3_RBM28_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.330:FF:000315">
        <property type="entry name" value="RNA-binding motif protein 28"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.330:FF:000340">
        <property type="entry name" value="RNA-binding motif protein 28"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.330">
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012677">
        <property type="entry name" value="Nucleotide-bd_a/b_plait_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035979">
        <property type="entry name" value="RBD_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000504">
        <property type="entry name" value="RRM_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051945">
        <property type="entry name" value="RRM_MRD1_RNA_proc_ribogen"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48039">
        <property type="entry name" value="RNA-BINDING MOTIF PROTEIN 14B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48039:SF5">
        <property type="entry name" value="RNA-BINDING PROTEIN 28"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00076">
        <property type="entry name" value="RRM_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00360">
        <property type="entry name" value="RRM"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54928">
        <property type="entry name" value="RNA-binding domain, RBD"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50102">
        <property type="entry name" value="RRM"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0539">Nucleus</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <keyword evidence="3 5" id="KW-0694">RNA-binding</keyword>
    <feature type="domain" description="RRM" evidence="7">
        <location>
            <begin position="4"/>
            <end position="80"/>
        </location>
    </feature>
    <feature type="domain" description="RRM" evidence="7">
        <location>
            <begin position="86"/>
            <end position="163"/>
        </location>
    </feature>
    <feature type="domain" description="RRM" evidence="7">
        <location>
            <begin position="241"/>
            <end position="319"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="428"/>
            <end position="562"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="456"/>
            <end position="466"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="6">
        <location>
            <begin position="524"/>
            <end position="541"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00176"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50102"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011545.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="594" mass="65229" checksum="B243C23A75C741F3" modified="2019-05-08" version="1">MAGLTLLVRGLPASARGPGLEQLFGRLGPLRRCFVVTRKGSETCRGFGYVTFSLLEDAQRALREVTTFEGHEIKVAVAKKKLREKKQRQLGELGTVCSEDDLKALFSPFGAVLEVNIPRKTDGKMRGFAFVQFKNILQAGKALKGMNMKEIKGRTVAVDWAVAKDKYRAMQSVGSGALTPLPGEPLVEGRAWPGHAGQLAATISSEAAGATWAAGSVAGSRSCWGSCRQWHGLTFGAPGRNLAFDTEEEELGEMLEQFGDLKYVRIVLHPDTEHSRGCAFAQFTTQEAAQKCLEAAQDDSEGGGLRLGGRRLHLDLAVSRDEAQKLRAQKVKKPTGTRNLYLAREGLIRAGTQAAEGVSTSDMAKRARFEELKRQKLKDQNIFVSRTRLCIHNLPKAVDDKQLRQLLLRAAGGRPGLRIKEVNPCWGEQLNRQKDPRGPRTQGTKGQKDALGPKGQGYAAGAVGQQNPRKRSPEGQGASTPWAGFQTKAEVEQVELPDGKKRRKVLTLPTHRGPKIRQRDRGKPAAPKKPKAQTSQRRHEKQRVAPRQAPGKRRKLVGGRAEARFDQLVEQYKRKILGSAPSAPGAKRSKWFES</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9X0</accession>
    <name>A0A452H9X0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Elongation factor 1-alpha</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="8">This protein promotes the GTP-dependent binding of aminoacyl-tRNA to the A-site of ribosomes during protein biosynthesis.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1 8">Belongs to the TRAFAC class translation factor GTPase superfamily. Classic translation factor GTPase family. EF-Tu/EF-1A subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9X0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013072.1">
        <property type="protein sequence ID" value="ENSGAGP00000011417.1"/>
        <property type="gene ID" value="ENSGAGG00000008805.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005525">
        <property type="term" value="F:GTP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003924">
        <property type="term" value="F:GTPase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003746">
        <property type="term" value="F:translation elongation factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="CDD" id="cd01883">
        <property type="entry name" value="EF1_alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd03693">
        <property type="entry name" value="EF1_alpha_II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd03705">
        <property type="entry name" value="EF1_alpha_III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.30.10:FF:000005">
        <property type="entry name" value="Elongation factor 1-alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000090">
        <property type="entry name" value="Elongation factor 1-alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.30.10:FF:000168">
        <property type="entry name" value="Elongation factor 1-alpha 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.30.10">
        <property type="entry name" value="Translation factors"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004161">
        <property type="entry name" value="EFTu-like_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031157">
        <property type="entry name" value="G_TR_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR054696">
        <property type="entry name" value="GTP-eEF1A_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000795">
        <property type="entry name" value="T_Tr_GTP-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050100">
        <property type="entry name" value="TRAFAC_GTPase_members"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009000">
        <property type="entry name" value="Transl_B-barrel_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004539">
        <property type="entry name" value="Transl_elong_EF1A_euk/arc"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00483">
        <property type="entry name" value="EF-1_alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF008969">
        <property type="entry name" value="PRK12317.1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23115">
        <property type="entry name" value="TRANSLATION FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22594">
        <property type="entry name" value="GTP-eEF1A_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00009">
        <property type="entry name" value="GTP_EFTU"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03144">
        <property type="entry name" value="GTP_EFTU_D2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00315">
        <property type="entry name" value="ELONGATNFCT"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50447">
        <property type="entry name" value="Translation proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00301">
        <property type="entry name" value="G_TR_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51722">
        <property type="entry name" value="G_TR_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 8" id="KW-0251">Elongation factor</keyword>
    <keyword evidence="7 8" id="KW-0342">GTP-binding</keyword>
    <keyword evidence="2" id="KW-0488">Methylation</keyword>
    <keyword evidence="4 8" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="6" id="KW-0648">Protein biosynthesis</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Tr-type G" evidence="9">
        <location>
            <begin position="5"/>
            <end position="242"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007249"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022768"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022917"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="RuleBase" id="RU000325"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011417.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="426" mass="46710" checksum="DE9B44116AC1C557" modified="2019-05-08" version="1">MGKEKTHINIVVIGHVDSGKSTTTGHLIYKCGGIDKRTIEKFEKEAAEMGKGSFKYAWVLDKLKAERERGITIDISLWKFETTKYYITIIDAPGHRDFIKNMITGTSQADCAVLIVAAGVGEFEAGISKNGQTREHALLAYTLGVKQLIVGINKMDSTEPPYSEKRYDEIVKEVSAYIKKIGYNPATVPFVPISGWHGDNMLEPSPNMPWFKGWKVERKEGNASGVSLLEALDTILPPTRPTDKPLRLPLQDVYKIGGIGTVPVGRVETGILRPGMVVTFAPVNITTEVKSVEMHHEALSEALPGDNVGFNVKNVSVKDIRRGNVCGDSKSDPPQEAAQFTSQVIILNHPGQISAGYSPVIDCHTAHIACKFAELKEKIDRRSGKKLEDNPKSLKSGDAAIVEMIPGKPMCVESFSQYPPLESSLH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9X1</accession>
    <name>A0A452H9X1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Peptidyl-prolyl cis-trans isomerase</fullName>
            <shortName evidence="6">PPIase</shortName>
            <ecNumber evidence="6">5.2.1.8</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="6">PPIases accelerate the folding of proteins. It catalyzes the cis-trans isomerization of proline imidic peptide bonds in oligopeptides.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="1 10">
            <text>[protein]-peptidylproline (omega=180) = [protein]-peptidylproline (omega=0)</text>
            <dbReference type="Rhea" id="RHEA:16237"/>
            <dbReference type="Rhea" id="RHEA-COMP:10747"/>
            <dbReference type="Rhea" id="RHEA-COMP:10748"/>
            <dbReference type="ChEBI" id="CHEBI:83833"/>
            <dbReference type="ChEBI" id="CHEBI:83834"/>
            <dbReference type="EC" id="5.2.1.8"/>
        </reaction>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the cyclophilin-type PPIase family.</text>
    </comment>
    <dbReference type="EC" id="5.2.1.8" evidence="6"/>
    <dbReference type="AlphaFoldDB" id="A0A452H9X1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013248.1">
        <property type="protein sequence ID" value="ENSGAGP00000011572.1"/>
        <property type="gene ID" value="ENSGAGG00000008887.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016018">
        <property type="term" value="F:cyclosporin A binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003755">
        <property type="term" value="F:peptidyl-prolyl cis-trans isomerase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006457">
        <property type="term" value="P:protein folding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.100.10:FF:000020">
        <property type="entry name" value="E3 SUMO-protein ligase RanBP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.100.10">
        <property type="entry name" value="Cyclophilin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029000">
        <property type="entry name" value="Cyclophilin-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024936">
        <property type="entry name" value="Cyclophilin-type_PPIase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020892">
        <property type="entry name" value="Cyclophilin-type_PPIase_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002130">
        <property type="entry name" value="Cyclophilin-type_PPIase_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11071">
        <property type="entry name" value="PEPTIDYL-PROLYL CIS-TRANS ISOMERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11071:SF561">
        <property type="entry name" value="PEPTIDYL-PROLYL CIS-TRANS ISOMERASE D-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00160">
        <property type="entry name" value="Pro_isomerase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF001467">
        <property type="entry name" value="Peptidylpro_ismrse"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00153">
        <property type="entry name" value="CSAPPISMRASE"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50891">
        <property type="entry name" value="Cyclophilin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00170">
        <property type="entry name" value="CSA_PPIASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50072">
        <property type="entry name" value="CSA_PPIASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 6" id="KW-0413">Isomerase</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="4 6" id="KW-0697">Rotamase</keyword>
    <feature type="domain" description="PPIase cyclophilin-type" evidence="7">
        <location>
            <begin position="8"/>
            <end position="136"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000971"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023110"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023235"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="RuleBase" id="RU363019"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50072"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011572.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="166" mass="18736" checksum="14532707F86E85C8" modified="2019-05-08" version="1">METNPVVYFEVSADDEPLGHITMELFSNIVPRTAENFRALCTGEKGFGFKNSIFHRIVPDFVCQGGDITRHDGTGGRSIYGDAFEDENFEVKHTGPGLLSMANRGRDTNNSQFFITLKKAEHLDFKHVVFGFVKDGWGPLQYRNKNEGEHVRTKLTLSTCNYCIHS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9X2</accession>
    <name>A0A452H9X2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">C2H2-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9X2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013060.1">
        <property type="protein sequence ID" value="ENSGAGP00000011409.1"/>
        <property type="gene ID" value="ENSGAGG00000008773.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006355">
        <property type="term" value="P:regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd07765">
        <property type="entry name" value="KRAB_A-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000295">
        <property type="entry name" value="zinc finger protein 19"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002343">
        <property type="entry name" value="Zinc finger protein 33A"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000135">
        <property type="entry name" value="Zinc finger protein 358"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002090">
        <property type="entry name" value="Zinc finger protein 473"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001270">
        <property type="entry name" value="zinc finger protein 583 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000431">
        <property type="entry name" value="zinc finger protein 629 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001909">
        <property type="entry name" value="KRAB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036051">
        <property type="entry name" value="KRAB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050331">
        <property type="entry name" value="Zinc_finger"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16515">
        <property type="entry name" value="PR DOMAIN ZINC FINGER PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16515:SF62">
        <property type="entry name" value="ZINC FINGER PROTEIN 883-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01352">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="SMART" id="SM00349">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109640">
        <property type="entry name" value="KRAB domain (Kruppel-associated box)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="11"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 8" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="9">
        <location>
            <begin position="134"/>
            <end position="161"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="9">
        <location>
            <begin position="162"/>
            <end position="189"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="9">
        <location>
            <begin position="190"/>
            <end position="217"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="9">
        <location>
            <begin position="218"/>
            <end position="245"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="9">
        <location>
            <begin position="246"/>
            <end position="273"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="9">
        <location>
            <begin position="274"/>
            <end position="301"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="9">
        <location>
            <begin position="302"/>
            <end position="329"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="9">
        <location>
            <begin position="330"/>
            <end position="357"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="9">
        <location>
            <begin position="358"/>
            <end position="385"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="9">
        <location>
            <begin position="386"/>
            <end position="413"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="9">
        <location>
            <begin position="414"/>
            <end position="441"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011409.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="441" mass="50616" checksum="3D1C954E0A5B6D04" modified="2019-05-08" version="1">LLLFQEPVTFEEVAVYFTTLYRDVMQENYENVTSLGKGYCPLSSWKRKRRDKVHTSTTMSPLLCPVSASPQYASDTLTARDPPLLQNTLGTEHRSSIGNHPGEKVGKCISCRGTQKDLKKTTTQEEILMGKRKNTCTECGKNVCDYSALIKHQQLHTGERPYECSECGKNFTRRSHLINHQRIHTGERPYKCSECGKSFTQRSHLTNHQKIHIGERPYECSECGKSFTQRSHLTRHQTIHTGERPYECHECGKSFRFSSALSEHQRIHTGEKRYECSECGKTFGRSSCLFQHQRIHTGERPYKCIECGKRFTTSSDLSKHQRIHTGERPYKCSECGKSFNQRSTLIKHQRIHTGERPYECHECAKTFNSSSSLIRHQRIHTGERPYECSECGKRFTTDSALFQHQSNHTGKRTHKCSECGKSFTTSLALSEHQRIHAGEGL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9X6</accession>
    <name>A0A452H9X6_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452H9X6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013089.1">
        <property type="protein sequence ID" value="ENSGAGP00000011433.1"/>
        <property type="gene ID" value="ENSGAGG00000008816.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026718">
        <property type="entry name" value="Luzp2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22414">
        <property type="entry name" value="LEUCINE ZIPPER PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22414:SF0">
        <property type="entry name" value="LEUCINE ZIPPER PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="12"/>
            <end position="134"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="166"/>
            <end position="197"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011433.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="263" mass="30330" checksum="2F6278B9B51C757A" modified="2019-05-08" version="1">MVKQGYYLQFGYEELERQLKEVFKERSNILRQLSKTSKELEGIKVNLQSLKNDEASAKSDVQKLLELGQKQREEMKYLQEAFQKQLNEVAEKAEKQQATINFLKTEMERKSKMIRDLQNENKSLKNKLLSGNKLCGIHAEESKKIQAQLKEFRYGKKDLIFKAQQLTDLEQKLAIAKDKLEKAALDKESQLKALKETVQLCLSSVLHNQPPATNIIPSKPTEKLTSQVTNDSKVPFQVTNTKILGEPLPEILYLQDSLFLLYQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9X9</accession>
    <name>A0A452H9X9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="19">Guanylate cyclase</fullName>
            <ecNumber evidence="19">4.6.1.2</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="17">Receptor for the C-type natriuretic peptide NPPC/CNP hormone. Has guanylate cyclase activity upon binding of its ligand. May play a role in the regulation of skeletal growth.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="1 19">
            <text>GTP = 3',5'-cyclic GMP + diphosphate</text>
            <dbReference type="Rhea" id="RHEA:13665"/>
            <dbReference type="ChEBI" id="CHEBI:33019"/>
            <dbReference type="ChEBI" id="CHEBI:37565"/>
            <dbReference type="ChEBI" id="CHEBI:57746"/>
            <dbReference type="EC" id="4.6.1.2"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="18">Belongs to the adenylyl cyclase class-4/guanylyl cyclase family.</text>
    </comment>
    <dbReference type="EC" id="4.6.1.2" evidence="19"/>
    <dbReference type="AlphaFoldDB" id="A0A452H9X9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013083.1">
        <property type="protein sequence ID" value="ENSGAGP00000011427.1"/>
        <property type="gene ID" value="ENSGAGG00000002083.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004016">
        <property type="term" value="F:adenylate cyclase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005525">
        <property type="term" value="F:GTP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004383">
        <property type="term" value="F:guanylate cyclase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016941">
        <property type="term" value="F:natriuretic peptide receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017046">
        <property type="term" value="F:peptide hormone binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004672">
        <property type="term" value="F:protein kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035556">
        <property type="term" value="P:intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001503">
        <property type="term" value="P:ossification"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007168">
        <property type="term" value="P:receptor guanylyl cyclase signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07302">
        <property type="entry name" value="CHD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd14042">
        <property type="entry name" value="PK_GC-A_B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000270">
        <property type="entry name" value="Guanylate cyclase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:001106">
        <property type="entry name" value="Guanylate cyclase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.1230:FF:000004">
        <property type="entry name" value="Guanylate cyclase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.2300:FF:000101">
        <property type="entry name" value="Guanylate cyclase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.2300">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.1230">
        <property type="entry name" value="Nucleotide cyclase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001054">
        <property type="entry name" value="A/G_cyclase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018297">
        <property type="entry name" value="A/G_cyclase_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001828">
        <property type="entry name" value="ANF_lig-bd_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001170">
        <property type="entry name" value="ANPR/GUC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050401">
        <property type="entry name" value="Cyclic_nucleotide_synthase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029787">
        <property type="entry name" value="Nucleotide_cyclase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028082">
        <property type="entry name" value="Peripla_BP_I"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001245">
        <property type="entry name" value="Ser-Thr/Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11920:SF494">
        <property type="entry name" value="ATRIAL NATRIURETIC PEPTIDE RECEPTOR 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11920">
        <property type="entry name" value="GUANYLYL CYCLASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01094">
        <property type="entry name" value="ANF_receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00211">
        <property type="entry name" value="Guanylate_cyc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07714">
        <property type="entry name" value="PK_Tyr_Ser-Thr"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00255">
        <property type="entry name" value="NATPEPTIDER"/>
    </dbReference>
    <dbReference type="SMART" id="SM00044">
        <property type="entry name" value="CYCc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55073">
        <property type="entry name" value="Nucleotide cyclase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53822">
        <property type="entry name" value="Periplasmic binding protein-like I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00458">
        <property type="entry name" value="ANF_RECEPTORS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00452">
        <property type="entry name" value="GUANYLATE_CYCLASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50125">
        <property type="entry name" value="GUANYLATE_CYCLASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="16 19" id="KW-0141">cGMP biosynthesis</keyword>
    <keyword evidence="20" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="14" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10" id="KW-0342">GTP-binding</keyword>
    <keyword evidence="15 18" id="KW-0456">Lyase</keyword>
    <keyword evidence="11" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="8" id="KW-0892">Osteogenesis</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="13" id="KW-0675">Receptor</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="5" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9" id="KW-1133">Transmembrane helix</keyword>
    <feature type="domain" description="Protein kinase" evidence="21">
        <location>
            <begin position="499"/>
            <end position="772"/>
        </location>
    </feature>
    <feature type="domain" description="Guanylate cyclase" evidence="22">
        <location>
            <begin position="847"/>
            <end position="977"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="20">
        <location>
            <begin position="784"/>
            <end position="811"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001436"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022855"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023293"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00043880"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="RuleBase" id="RU000405"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="RuleBase" id="RU003431"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS50125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011427.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1032" mass="116398" checksum="7A3D3D21CB03068C" modified="2019-05-08" version="1">MTGRWGRKIWRGPEGANRSLSLAVVLPERNVSYPWAWPRVGPAIRLALEALERGDPPLLPGPLAVRLHFRSSELEGACSEYVAPLNAVDLKLYHDPDVLFGPGCVYPAASVGRFASHWRLPLITAGAVATGFSHKQEHYGTTVRTGPSAPKLGAFVSQLHAHFNWSARAALLYADHKTDDRPYYFTIEGVYQELQEALNLSVRYHIYAPEEGGADAALHFLRASARGEFIYVCGPLEMLHQIMRLAQRESLTNGDYVFFYVDVFGESLRASATRDALKPWQNKQSQDVGLRQAFQTVLVITYHEPQNPEYQHFQTQLILRAKEEFKVELGYSLMNLVAGCFYDGVLLYAMALNETLAEGGSKKDGTRIVQKMRDRKFQGVTGLVSMDSNNDRDTDFSLWLCSLSMQVAGHYAGAEKKLNWLGRPIQWVKGAPPPDNPPCVFDVDDPSCDKRGPASLCPSSVWTGRKLMLEKELASMLWRIRWDELQFGNAERYYKGAGSRLTLSLRGSSYGSLMTAQGKYQIFANTGHFKGNVVAVKHVNKKRIELTRQVLFELKHMRDIQFNHLTRFIGACIDPPNVCIVTEYCPRGSLQDILENESINLDWMFRYSLINDIVKGMAFLHNSIIGHHGSLKSSNCVVDSRFVLKITDYGLASFRSSCDAEDSHALYAKKLWTAPELLQQGRLPLPGMQKADVYSFSIILQEIALRNGPFYIEGMDLSPKEIVQKVRNSQKPFFRPSIDMGLHSEELAVLMERCWAQEPAERPDFSQIKIFIRRFNKEGSTSILDNLLSRMEQYANNLEKLVEERTQAYLEEKRKAENLLYQILPHSVAEQLKRGETVRAEAFDSVTIYFSDIVGFTALSAESTPMQVVTLLNDLYTCFDAIIDNFDVYKVETIGDAYMVVSGLPVRNGKLHANEIVRMALALLEAVKTFKIRHRPNDQLRLRIGIHTGPVCAGVVGLKMPRYCLFGDTVNTASRMESNGQALKIHVSSTTKEVLDEFGCFELELRGDVEMKGKGKMRTYWLLGERTDDSII</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9Y1</accession>
    <name>A0A452H9Y1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="23">1-phosphatidylinositol 3-phosphate 5-kinase</fullName>
            <ecNumber evidence="3">2.7.1.150</ecNumber>
            <ecNumber evidence="4">2.7.11.1</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="26">FYVE finger-containing phosphoinositide kinase</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="27">PIKfyve</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="25">Phosphatidylinositol 3-phosphate 5-kinase type III</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="24">Serine-protein kinase PIKFYVE</fullName>
        </alternativeName>
    </protein>
    <organism evidence="34 35">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="35" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="34" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="34" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="19">
            <dbReference type="Rhea" id="RHEA:17990"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="20">
            <text>a 1,2-diacyl-sn-glycero-3-phospho-(1D-myo-inositol) + ATP = a 1,2-diacyl-sn-glycero-3-phospho-(1D-myo-inositol-5-phosphate) + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:44680"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:57795"/>
            <dbReference type="ChEBI" id="CHEBI:57880"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="20">
            <dbReference type="Rhea" id="RHEA:44681"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="21">
            <text>a 1,2-diacyl-sn-glycero-3-phospho-(1D-myo-inositol-3-phosphate) + ATP = a 1,2-diacyl-sn-glycero-3-phospho-(1D-myo-inositol-3,5-bisphosphate) + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:13609"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:57923"/>
            <dbReference type="ChEBI" id="CHEBI:58088"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.1.150"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="21">
            <dbReference type="Rhea" id="RHEA:13610"/>
        </physiologicalReaction>
    </comment>
    <comment type="subunit">
        <text evidence="22">Component of the PI(3,5)P2 regulatory complex/PAS complex, at least composed of PIKFYVE, FIG4 and VAC14. VAC14 nucleates the assembly of the complex and serves as a scaffold by pentamerizing into a star-shaped structure, which can bind a single copy each of PIKFYVE and FIG4 and coordinates their activities. Interacts (via chaperonin-like domain) with RABEPK; the interaction recruits RABEPK to the endosomal membrane. Interacts with SPAG9. Interacts with EGFR.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="18">Cytoplasmic vesicle</location>
            <location evidence="18">Phagosome membrane</location>
            <topology evidence="18">Peripheral membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Early endosome membrane</location>
            <topology evidence="1">Peripheral membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Late endosome membrane</location>
            <topology evidence="2">Peripheral membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="2.7.1.150" evidence="3"/>
    <dbReference type="EC" id="2.7.11.1" evidence="4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013088.1">
        <property type="protein sequence ID" value="ENSGAGP00000011432.1"/>
        <property type="gene ID" value="ENSGAGG00000008802.1"/>
    </dbReference>
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        </source>
    </evidence>
    <sequence length="2089" mass="237005" checksum="48F5EB82790CFDE3" modified="2019-05-08" version="1">MLLLLYHIQVCKACVYNTLKFFSRTHEMATDDRQSPTLDSVSDLPRSPSSPSHLTHFKPLTPDQDEPPFKSAYSSFVNLFRFGKDDGRLASPAESGQGEQQALSGTWSSPQHTSRAQSLRSPVPYKKQLTGELQRRASATLDTRRKVEPSLSGHDPRTAVQLRSLSTVLKRLKEIMEGKSQDSDLKQYWMPDSQCKECYDCSEKFTTFRRRHHCRLCGQIFCSRCCNQEIPGKFMGYTGDLRACTYCRKIALSYAHSTDSNSIGEDLNALSDSTCCVSVLEPGEPRTPVGSRKASRNIFLEEDLTWQSLIHPDSSTTTLSTRLVSVQEDPGKSPARNRSASITNLSLDRSASPMVPAYETSVSPQPNRTYLKAETNEDERKILLDSVQLKDLWKKICHHNSGMEFQDHRYWLRTHPNCIVGKELVNWLIRNGHITTRAQAIAIGQALVDGRWLDCVSHHDQIFRDEYALYRPLQSTEFSETPSPDSDSVNSVEGHSEPSWFKDIKFDDSDTEQIADEGEDNLAKYLNPDNGGQQMISISDAFIKESLFNRRVEEKSKELFFTPLGWHHSNLDLLREENGEKQAMERLLSANHNHMMALLQQLLNNESLSLSWRDIIVPVVCQVVQTVRPDVKNRDDDMDIRQFVHIKKIPGGKKFDSVVVNGFVCTKNVAHKKMNTYIKNPKILLLKCSIEYLYREETKFTCIDPIVLQEREFLKNYVQRIVDVRPTLVLVEKTVSRIAQDMLLEHGITLVINVKPQVLDRISRMTQGDLVMSMDQLLTKPHLGTCHKFYMQVFQLLNDQTKTLMFFEGCPPHLGCTIKLRGAADYELTRVKEILIFMVCVAYHSQLEISFLMDEFAMPPSLTKNFHTLIESQGYENGHQDLFNGEEFSTVVRDVELSPEKVPMVSESVSSDEASTLEQRTLFEEDQDDDDDVLREGSSLRHQELLRMEYSLPVSSNITTPVPESRLPFHNTDQHPGTLYSQHLETLKQPGDLQDSKSQMRVFRDPLQDDTGLYVMEEVTSSEDRLKAYSSAFKQELKDVILCISPVITFHEPFLLTEMGMRCPARDYFPEQVYLSPFLNKEYKELESRRKKQLLRDLSGLQGINGSIQAKAVQVLPTHELVNTRIAEHLGDSQSLARMLADYRARGGRIQQKNADPFAQCKDASGVPGGKAGGRNEEDERGLVRGESVWSHKVDCLSPVNHQRLCVLFSSSSAQSSNAPSACVSPWIVTMEFYGKNDLTLGVFLERYCFRPSYQCPSMFCETPMVHHVRRFVHGQGCVQIILKELDSPVPGYQHTILTYSWCRLCKQVTPVVPLSNDSWSMSFAKYLELRFYGHQYTRRANAEPCGHSIHHDYHQYFSYNQMVASFSYSPIRLLEVCVPHPKIYIKRQAPLKVTILQDLKDFSQKVSQVYLAVDDRLTSLKTDTFSKTREEKMEDLFAQKEMEEVEFRNWIEKIQARMLSPSLDTSQQLQSVFESLIAKKQSLCEMLQAWNNRLQDLFQQEKGRKRPSVPPSPGRLRQGEENKISAMDASPRNTSPALQNGEKDDRFLTTLSSQSSTSSTHLQLPTPPEIVSEQLTGGPSFANSLSEPDTTSSSEDIFDGHSLGSTDSQVKEKSTMKAILANLLPGNSYNPIPFPFDPDKHYLMYEHERVPIAVCEKEPSSIIAFALSCKEYRNGLDELSKASLKSSSEEGLPPNSMSDNKPKSSSPVRLPETNVGQANRTAEAEQPKKVSGVLSFFRGTGGKSPDLSAQKKETLRGADSAYYQVGQMGKEGPESQGAETQDEADGGDAQKKQLANPHVELQFSDANAKFYCRIYYAGEFHKMRDVILGSREEDFIRSLSHSLPWQARGGKSGAAFYVTEDDRFILKQMPRLEVQSFLDFAPHYFTYITNAVQQKKPTALAKILGVYRIGYKNSQNNTEKKLDLLVMENLFYGRKMAQVFDLKGSLRNRNVKTDTGKESCDVVLLDENLLKMVRDNPLYIRSHCKAVLRASIHSDSHFLSSHLIIDYSLLVGRDDTSNELVVGIIDYIRTFTWDKKLEMVVKSTGILGGQGKMPTVVSPELYRTRFCEAMDKYFLMVPDHWTGLGLNC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9Y5</accession>
    <name>A0A452H9Y5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">TROVE domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452H9Y5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013100.1">
        <property type="protein sequence ID" value="ENSGAGP00000011442.1"/>
        <property type="gene ID" value="ENSGAGG00000008821.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005697">
        <property type="term" value="C:telomerase holoenzyme complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003720">
        <property type="term" value="F:telomerase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070034">
        <property type="term" value="F:telomerase RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000722">
        <property type="term" value="P:telomere maintenance via recombination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052652">
        <property type="entry name" value="Telomerase_Complex_Comp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008858">
        <property type="entry name" value="TROVE_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037214">
        <property type="entry name" value="TROVE_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR44791:SF1">
        <property type="entry name" value="TELOMERASE PROTEIN COMPONENT 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR44791">
        <property type="entry name" value="TELOMERASE PROTEIN COMPONENT 1 TEP1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05731">
        <property type="entry name" value="TROVE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF140864">
        <property type="entry name" value="TROVE domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50988">
        <property type="entry name" value="TROVE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="TROVE" evidence="2">
        <location>
            <begin position="25"/>
            <end position="238"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="177"/>
            <end position="238"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="1">
        <location>
            <begin position="177"/>
            <end position="187"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="PROSITE" id="PS50988"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011442.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="238" mass="26006" checksum="B4B1FB3BD67DA026" modified="2019-05-08" version="1">MGSSAEVGKLRPPGPHPPVGPFYLALGLPAGEASPWPLPHSLSSPCCQRSGRHGWLWPGGKAASPAALSELCEELAQLEPEFILKVSLYTRQELNIRGTANFLLALAARLPPCRPHLRRYLCATLQLPSDWIQVVALFQSLAGAGRPLAPLPRCLRAGLADKFQQFDAYQLAKYNSRKSRGKARGRPWPRQPRRELRSPALGADHMEPGKSPTQLPWLEPRSGARPGPCSLTRARGPD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9Y8</accession>
    <name>A0A452H9Y8_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="15">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the plexin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9Y8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013125.1">
        <property type="protein sequence ID" value="ENSGAGP00000011464.1"/>
        <property type="gene ID" value="ENSGAGG00000008707.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002116">
        <property type="term" value="C:semaphorin receptor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017154">
        <property type="term" value="F:semaphorin receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007162">
        <property type="term" value="P:negative regulation of cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050772">
        <property type="term" value="P:positive regulation of axonogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030334">
        <property type="term" value="P:regulation of cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008360">
        <property type="term" value="P:regulation of cell shape"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd12792">
        <property type="entry name" value="RasGAP_plexin_B2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000203">
        <property type="entry name" value="Plexin B2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000798">
        <property type="entry name" value="Plexin B2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.506.10:FF:000035">
        <property type="entry name" value="Plexin b2a"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.506.10">
        <property type="entry name" value="GTPase Activation - p120gap, domain 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014756">
        <property type="entry name" value="Ig_E-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002909">
        <property type="entry name" value="IPT_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031148">
        <property type="entry name" value="Plexin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013548">
        <property type="entry name" value="Plexin_cytoplasmic_RasGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046800">
        <property type="entry name" value="Plexin_RBD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016201">
        <property type="entry name" value="PSI"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057533">
        <property type="entry name" value="PSI_Plexin-B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041019">
        <property type="entry name" value="TIG1_plexin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22625">
        <property type="entry name" value="PLEXIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22625:SF9">
        <property type="entry name" value="PLEXIN-B2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08337">
        <property type="entry name" value="Plexin_cytopl"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20170">
        <property type="entry name" value="Plexin_RBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24317">
        <property type="entry name" value="PSI_Plexin-B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01833">
        <property type="entry name" value="TIG"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17960">
        <property type="entry name" value="TIG_plexin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00429">
        <property type="entry name" value="IPT"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00423">
        <property type="entry name" value="PSI"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81296">
        <property type="entry name" value="E set domains"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="12" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="9" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="8" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0675">Receptor</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <keyword evidence="4" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7" id="KW-1133">Transmembrane helix</keyword>
    <feature type="domain" description="PSI" evidence="13">
        <location>
            <begin position="108"/>
            <end position="158"/>
        </location>
    </feature>
    <feature type="domain" description="PSI" evidence="13">
        <location>
            <begin position="204"/>
            <end position="250"/>
        </location>
    </feature>
    <feature type="domain" description="IPT/TIG" evidence="14">
        <location>
            <begin position="251"/>
            <end position="344"/>
        </location>
    </feature>
    <feature type="domain" description="IPT/TIG" evidence="14">
        <location>
            <begin position="345"/>
            <end position="431"/>
        </location>
    </feature>
    <feature type="domain" description="IPT/TIG" evidence="14">
        <location>
            <begin position="433"/>
            <end position="541"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="12">
        <location>
            <begin position="666"/>
            <end position="693"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010297"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="SMART" id="SM00423"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="SMART" id="SM00429"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011464.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1276" mass="144719" checksum="4A1FA7EAC0E58E52" modified="2019-05-08" version="1">MPNHWLWSPDGVCMAITSAHPQNMSRKAPGKVELTVSPLPELTESDMLWCQFGETTNIAELRDGTIICDPPTTIPHTPKGQDHVSVSLILTFGEEKMFFASATYPFYDCEEAMNLSENLPCSSCASSRWNCQWDWKKHFCEDSSLEEDVITQNMEDCPQFLNPNPQIIPMEFPTIVHFEGKNLENYQVNELIPSSHLTYMTLYNCSYGRSDCSLCLAADPGYDCVWCEEEGKPNKCIYKNLCKTSTNTCPHPVITRIEPKTGPLSGGILLTIMGSNLGIKAEDVKSITVAGTECLFKEDFYSVSIRYLIVCQAGNVSVPSQGKIEVDINGKRGSSTSEVQFTYQQPHPTGISPTEGPRAGGTILTITGNNLETGSQKDVRVLLGDVPCNITDFGEQIVCITGQHESGGKVQVTVDHGDTENQLNDPWFTYSENPTVTKFFPKNSFSSGGRNITVTGTGFDIIQSFSIVVTADLHGPSRRKKRQIRSLDSKKVVCMYICITIVFSSPPILEDPENYRITTLIKMDGCSLVLNAENNDPFKYVADPTFENFTDGIKKQVNKLIHAKGSNLNKAMTIEEAQAFVGAEPCNIKTLTEKDLYCEPPEVQPQPKKRQKRDTINNLPEFIVKFGLREWILGRVEYDTRVSDVPLDVILPSVIFPMVAIIIISIYCYRRKSQQAEREYEKIKSQLEGLEESVRDRCKKEFTDLMIEMEDQTNDVSEAGIPVLDYKMYTDRVFFLPSKDGEKDVMITGKLDIPEARKQTVEQALNQFSNLLNSKSFLINFIHTLEKQREFSAREKVYFASLLTVALHGKLEYYTDIMRTLFLELMEQYVVAKNPKLMLRRSETVVERMLSNWMAICLYQYLKDNAGEPLYKLFKAIKHQVEKGPVDAVLMKAKYTLNDTGLLGDDVEYTQLTVNVYVQDGGTDSVPVKVLNCDTISQVKEKIIDQVYRNLPCSQWPKADSVVLVVFKPASNSHVLWDDFSIFVFLKVRDGATLILSKIGISQQPEDNQQDVPGERHALLEDENKVWHLVRPADEVDEGKSKRGSMKEKERTKAITEIYLTRLLSVKGTLQQFVDNFFHSVLNSNHVVPPAVKYFFDFLDEQAEKHELKDEDTIHIWKTNSLPLRFWVNILKNPHFIFDVHVHEVVDASLSVIAQTFMDACTRTEHKLSRDSPSNKLLYAKEISNYKKMVEDYYKGIRQMVQVSDQDMNTHLAEISRAHTDSLNTLVALHQLYQYTNKYYDEIINALEEDPAAQKMQLAFRLQQIAAALENKVTDL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9Z9</accession>
    <name>A0A452H9Z9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Beta/gamma crystallin 'Greek key' domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">Crystallins are the dominant structural components of the vertebrate eye lens.</text>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the beta/gamma-crystallin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9Z9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013140.1">
        <property type="protein sequence ID" value="ENSGAGP00000011478.1"/>
        <property type="gene ID" value="ENSGAGG00000008850.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005212">
        <property type="term" value="F:structural constituent of eye lens"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002088">
        <property type="term" value="P:lens development in camera-type eye"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007601">
        <property type="term" value="P:visual perception"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.20.10:FF:000001">
        <property type="entry name" value="Crystallin gamma S"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.20.10">
        <property type="entry name" value="Crystallins"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050252">
        <property type="entry name" value="Beta/Gamma-Crystallin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001064">
        <property type="entry name" value="Beta/gamma_crystallin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011024">
        <property type="entry name" value="G_crystallin-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11818">
        <property type="entry name" value="BETA/GAMMA CRYSTALLIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11818:SF119">
        <property type="entry name" value="GAMMA-CRYSTALLIN D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00030">
        <property type="entry name" value="Crystall"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01367">
        <property type="entry name" value="BGCRYSTALLIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00247">
        <property type="entry name" value="XTALbg"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49695">
        <property type="entry name" value="gamma-Crystallin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50915">
        <property type="entry name" value="CRYSTALLIN_BETA_GAMMA"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0273">Eye lens protein</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="Beta/gamma crystallin 'Greek key'" evidence="5">
        <location>
            <begin position="5"/>
            <end position="43"/>
        </location>
    </feature>
    <feature type="domain" description="Beta/gamma crystallin 'Greek key'" evidence="5">
        <location>
            <begin position="44"/>
            <end position="85"/>
        </location>
    </feature>
    <feature type="domain" description="Beta/gamma crystallin 'Greek key'" evidence="5">
        <location>
            <begin position="118"/>
            <end position="160"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00003689"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009646"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50915"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011478.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="163" mass="18829" checksum="AD219484080679DD" modified="2019-05-08" version="1">MGRPMNIILFEDRNFQGRSVECSSDRPDLQSHLSRCNSIRVESGCFMLYERPNFQGQQFFVKRGDYPDMQSEGFSTSIKSCRMIPPVSSILLLKHGNVAQATIIRHLLDHFQCSLVLGHGILYEMPNYRGRQYLLRPGQYRRFSEWGAMSGRVGSLRRATDLY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJW3</accession>
    <name>A0A452HJW3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">5'-nucleotidase domain containing 3</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="6">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
        <text evidence="6">Binds 1 Mg(2+) ion per subunit.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the 5'(3')-deoxyribonucleotidase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJW3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017371.1">
        <property type="protein sequence ID" value="ENSGAGP00000015210.1"/>
        <property type="gene ID" value="ENSGAGG00000011446.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008253">
        <property type="term" value="F:5'-nucleotidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1000">
        <property type="entry name" value="HAD superfamily/HAD-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036412">
        <property type="entry name" value="HAD-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008380">
        <property type="entry name" value="HAD-SF_hydro_IG_5-nucl"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023214">
        <property type="entry name" value="HAD_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016695">
        <property type="entry name" value="Pur_nucleotidase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12103">
        <property type="entry name" value="5'-NUCLEOTIDASE DOMAIN-CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12103:SF11">
        <property type="entry name" value="5'-NUCLEOTIDASE DOMAIN-CONTAINING PROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05761">
        <property type="entry name" value="5_nucleotid"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF017434">
        <property type="entry name" value="Purine_5'-nucleotidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56784">
        <property type="entry name" value="HAD-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="4 6" id="KW-0460">Magnesium</keyword>
    <keyword evidence="2 6" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <feature type="active site" description="Nucleophile" evidence="5">
        <location>
            <position position="31"/>
        </location>
    </feature>
    <feature type="active site" description="Proton donor" evidence="5">
        <location>
            <position position="33"/>
        </location>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="31"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="33"/>
        </location>
        <ligand>
            <name>GMP</name>
            <dbReference type="ChEBI" id="CHEBI:58115"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="272"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00009589"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PIRSR" id="PIRSR017434-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PIRSR" id="PIRSR017434-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015210.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="9">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="434" mass="51391" checksum="BE7A10FD64496412" modified="2025-10-08" version="2">MLLVLPNLLNPDAIFTNNEMSLSDIEIYGFDYDYTLVFYSKHLHTLIYNAARDLLINEHRYPAEIRKYDYDPNFAIRGLHYDVHRALLMKIDAFHYIQLGTVYRGFSVVPDEEVIAMYDGSHVPLEQMSDFYGKDSIRDVHIKGIMYRAIEADIEKYVCYAEQTRAVLRKLADHGKKIFLITNSPSSFVDKGMKFLVGKDWRDLFDVVIVQADKPNFFNDKRRPFRKVNERGVLLWDKIHKLQKGQIYKQGNLYEFLKLTGWRGSKVLYFGDHIYSDLADLTLKHGWRTGAIIPELRSEIKIMNTEQYIQTMTWLQTLTGLLEHMQIYQDPDSQMILQEWKKERKEMREMTRNFFNSQFGSLFRTDQNPTYFLRRLSRFADIYMASLSCLLNYDLNYTFYPRRTPLQHELPTWSDQLCSGAFRIPLLQEMAQIK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJW5</accession>
    <name>A0A452HJW5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">5'-nucleotidase domain containing 3</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="6">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
        <text evidence="6">Binds 1 Mg(2+) ion per subunit.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the 5'(3')-deoxyribonucleotidase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJW5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017369.1">
        <property type="protein sequence ID" value="ENSGAGP00000015208.1"/>
        <property type="gene ID" value="ENSGAGG00000011446.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008253">
        <property type="term" value="F:5'-nucleotidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1000">
        <property type="entry name" value="HAD superfamily/HAD-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036412">
        <property type="entry name" value="HAD-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008380">
        <property type="entry name" value="HAD-SF_hydro_IG_5-nucl"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023214">
        <property type="entry name" value="HAD_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016695">
        <property type="entry name" value="Pur_nucleotidase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12103">
        <property type="entry name" value="5'-NUCLEOTIDASE DOMAIN-CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12103:SF11">
        <property type="entry name" value="5'-NUCLEOTIDASE DOMAIN-CONTAINING PROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05761">
        <property type="entry name" value="5_nucleotid"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF017434">
        <property type="entry name" value="Purine_5'-nucleotidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56784">
        <property type="entry name" value="HAD-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="4 6" id="KW-0460">Magnesium</keyword>
    <keyword evidence="2 6" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <feature type="active site" description="Nucleophile" evidence="5">
        <location>
            <position position="45"/>
        </location>
    </feature>
    <feature type="active site" description="Proton donor" evidence="5">
        <location>
            <position position="47"/>
        </location>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="45"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="47"/>
        </location>
        <ligand>
            <name>GMP</name>
            <dbReference type="ChEBI" id="CHEBI:58115"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="286"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00009589"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PIRSR" id="PIRSR017434-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PIRSR" id="PIRSR017434-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015208.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="9">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="448" mass="53028" checksum="24D386AE4A932002" modified="2025-10-08" version="2">MKKLKYEIAELLDLVPSIMSNLLNPDAIFTNNEMSLSDIEIYGFDYDYTLVFYSKHLHTLIYNAARDLLINEHRYPAEIRKYDYDPNFAIRGLHYDVHRALLMKIDAFHYIQLGTVYRGFSVVPDEEVIAMYDGSHVPLEQMSDFYGKDSIRDVHIKGIMYRAIEADIEKYVCYAEQTRAVLRKLADHGKKIFLITNSPSSFVDKGMKFLVGKDWRDLFDVVIVQADKPNFFNDKRRPFRKVNERGVLLWDKIHKLQKGQIYKQGNLYEFLKLTGWRGSKVLYFGDHIYSDLADLTLKHGWRTGAIIPELRSEIKIMNTEQYIQTMTWLQTLTGLLEHMQIYQDPDSQMILQEWKKERKEMREMTRNFFNSQFGSLFRTDQNPTYFLRRLSRFADIYMASLSCLLNYDLNYTFYPRRTPLQHELPTWSDQLCSGAFRIPLLQEMAQIK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJZ2</accession>
    <name>A0A452HJZ2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">Calcium-transporting ATPase</fullName>
            <ecNumber evidence="21">7.2.2.10</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="21">Catalyzes the hydrolysis of ATP coupled with the transport of calcium.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>Ca(2+)(in) + ATP + H2O = Ca(2+)(out) + ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:18105"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="7.2.2.10"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="19">
            <dbReference type="Rhea" id="RHEA:18106"/>
        </physiologicalReaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subunit">
        <text evidence="20">Interacts with sarcolipin (SLN). Interacts with phospholamban (PLN). Interacts with myoregulin (MRLN). Interacts with DWORF.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Endoplasmic reticulum membrane</location>
            <topology evidence="3">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="21">Membrane</location>
            <topology evidence="21">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Sarcoplasmic reticulum membrane</location>
            <topology evidence="2">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4 21">Belongs to the cation transport ATPase (P-type) (TC 3.A.3) family. Type IIA subfamily.</text>
    </comment>
    <comment type="caution">
        <text evidence="21">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="EC" id="7.2.2.10" evidence="21"/>
    <dbReference type="AlphaFoldDB" id="A0A452HJZ2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017408.1">
        <property type="protein sequence ID" value="ENSGAGP00000015237.1"/>
        <property type="gene ID" value="ENSGAGG00000011401.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033017">
        <property type="term" value="C:sarcoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005388">
        <property type="term" value="F:P-type calcium transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="CDD" id="cd02083">
        <property type="entry name" value="P-type_ATPase_SERCA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.1110.10:FF:000003">
        <property type="entry name" value="Calcium-transporting ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1000:FF:000005">
        <property type="entry name" value="Calcium-transporting ATPase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1110.10:FF:000065">
        <property type="entry name" value="Sarcoplasmic/endoplasmic reticulum calcium ATPase 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.70.150.10:FF:000160">
        <property type="entry name" value="Sarcoplasmic/endoplasmic reticulum calcium ATPase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.1110.10">
        <property type="entry name" value="Calcium-transporting ATPase, cytoplasmic domain N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.70.150.10">
        <property type="entry name" value="Calcium-transporting ATPase, cytoplasmic transduction domain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1110.10">
        <property type="entry name" value="Calcium-transporting ATPase, transmembrane domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1000">
        <property type="entry name" value="HAD superfamily/HAD-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004014">
        <property type="entry name" value="ATPase_P-typ_cation-transptr_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023299">
        <property type="entry name" value="ATPase_P-typ_cyto_dom_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018303">
        <property type="entry name" value="ATPase_P-typ_P_site"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023298">
        <property type="entry name" value="ATPase_P-typ_TM_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008250">
        <property type="entry name" value="ATPase_P-typ_transduc_dom_A_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR059000">
        <property type="entry name" value="ATPase_P-type_domA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036412">
        <property type="entry name" value="HAD-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023214">
        <property type="entry name" value="HAD_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005782">
        <property type="entry name" value="P-type_ATPase_IIA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001757">
        <property type="entry name" value="P_typ_ATPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044492">
        <property type="entry name" value="P_typ_ATPase_HD_dom"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01116">
        <property type="entry name" value="ATPase-IIA1_Ca"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01494">
        <property type="entry name" value="ATPase_P-type"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR42861">
        <property type="entry name" value="CALCIUM-TRANSPORTING ATPASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13246">
        <property type="entry name" value="Cation_ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00690">
        <property type="entry name" value="Cation_ATPase_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00122">
        <property type="entry name" value="E1-E2_ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08282">
        <property type="entry name" value="Hydrolase_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00119">
        <property type="entry name" value="CATATPASE"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00120">
        <property type="entry name" value="HATPASE"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDG00002">
        <property type="entry name" value="C1.7:_P-type_atpase_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDS00003">
        <property type="entry name" value="Haloacid_Dehalogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDF00027">
        <property type="entry name" value="p-type_atpase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00831">
        <property type="entry name" value="Cation_ATPase_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81653">
        <property type="entry name" value="Calcium ATPase, transduction domain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81665">
        <property type="entry name" value="Calcium ATPase, transmembrane domain M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56784">
        <property type="entry name" value="HAD-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81660">
        <property type="entry name" value="Metal cation-transporting ATPase, ATP-binding domain N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00154">
        <property type="entry name" value="ATPASE_E1_E2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12 21" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="11 21" id="KW-0106">Calcium</keyword>
    <keyword evidence="7 21" id="KW-0109">Calcium transport</keyword>
    <keyword evidence="10" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="17 21" id="KW-0406">Ion transport</keyword>
    <keyword evidence="13" id="KW-0460">Magnesium</keyword>
    <keyword evidence="18 21" id="KW-0472">Membrane</keyword>
    <keyword evidence="9 21" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="14" id="KW-0703">Sarcoplasmic reticulum</keyword>
    <keyword evidence="15" id="KW-1278">Translocase</keyword>
    <keyword evidence="8 21" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="16 21" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="5 21" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="54"/>
            <end position="72"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="78"/>
            <end position="101"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="254"/>
            <end position="274"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="286"/>
            <end position="315"/>
        </location>
    </feature>
    <feature type="domain" description="Cation-transporting P-type ATPase N-terminal" evidence="23">
        <location>
            <begin position="3"/>
            <end position="71"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="766"/>
            <end position="789"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="22">
        <location>
            <begin position="769"/>
            <end position="789"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004326"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004477"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00005675"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022568"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022951"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022967"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00047282"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00061760"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="RuleBase" id="RU361146"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="SMART" id="SM00831"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015237.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="789" mass="86085" checksum="EE894F364437A450" modified="2019-05-08" version="1">MENSHAKTVEECLAYFGVSENSGLTPEQVKKNLDKYGPNGKSIWELVLEQFEDLLVRILLLAACISFVLAWFEEGEETVTAFVEPFVILLILIANAIVGVWQERNAENAIEALKEYEPEMGKVYRADRKSVQRIKARDMVPGDIAEVAVGDKVPADIRIISIKSTTLRVDQSILTGESVSVIKHTDPVPDPRAVNQDKKNMLFSGTNVAAGKAVGIVIATGVGTEIGKIRDEMAATEQEKTPLQQKLDEFGEQLSKVISLICVAVWLINIGHFNDPVHGGSWIRGAIYYFKIAVALAVAAIPEGLPAVITTCLALGTRRMAKKNAIVRSLPSVETLGCTSVICSDKTGTLTTNQMSVCKMFVIEKVEGDVCSLNEFSITGSTYAPEGEVLKNDKHVKAGQYDGLVELATICALCNDSSLDYNEAKGAYEKVGEATETALTCLVEKMNVFNTDVRTLSKVERANACNSVIKQLMKKEFTLEFSRDRKSMSVYCSPAKASRAAVSNKMFIKGAPEGVIDRCNYVRVGTTRVPLTSLVKDKILTVIKEWGTGRDTLRCLALATRDTPPRREDMVLEDSTKFVEYEVDLTFVGCVGMLDPPRKEVMGSIQLCRDAGIRVIMITGDNKGTAIAICRRIGIFTEDEEVMGRAYTGREFDDLPPAEQREACKRACCFARVEPAHKSKIVEFLQSFDEITAMTGDGVNDAPALKKAEIGIAMGSGTAVAKTASEMVLADDNFSTIVSAVEEGRAIYNNMKQFIRYLISSNVGEVPRQPSPPLPQPCHPSPTPATPAG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJZ3</accession>
    <name>A0A452HJZ3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Septin-type G domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="8">Belongs to the TRAFAC class TrmE-Era-EngA-EngB-Septin-like GTPase superfamily. Septin GTPase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJZ3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017409.1">
        <property type="protein sequence ID" value="ENSGAGP00000015238.1"/>
        <property type="gene ID" value="ENSGAGG00000011502.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005525">
        <property type="term" value="F:GTP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd01850">
        <property type="entry name" value="CDC_Septin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000143">
        <property type="entry name" value="septin-9 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR030379">
        <property type="entry name" value="G_SEPTIN_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016491">
        <property type="entry name" value="Septin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18884">
        <property type="entry name" value="SEPTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00735">
        <property type="entry name" value="Septin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51719">
        <property type="entry name" value="G_SEPTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="3" id="KW-0132">Cell division</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="5 8" id="KW-0342">GTP-binding</keyword>
    <keyword evidence="4 8" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Septin-type G" evidence="10">
        <location>
            <begin position="282"/>
            <end position="554"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="14"/>
            <end position="172"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="222"/>
            <end position="261"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="20"/>
            <end position="47"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="97"/>
            <end position="109"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="116"/>
            <end position="134"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="RuleBase" id="RU004560"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS51719"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015238.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="571" mass="64578" checksum="188292F16D14BBEA" modified="2019-05-08" version="1">MERDRISALKRSFEVEEVDQSLNASTPQRRTPNPLLRSTVSSSTQKFQDLGARNSETPARHADSPGQRSPKSSLRRVELSGPKLPEPVSRRTEISIDISSKQVENSTPGLSRFGLKRAEGLGHKPPEPTPRRTEITIGKPQEPAVRRMETPTSKIPELPQRKAETASAPVPDVATKRVEIQVAKSPEVPVPPVRQVENLEPSLASQHLQPESKPQPVMVEIPPKSPEGPEAVPSYTPSMTDAARDTGLKQAAPPRPEKPAPDFSYVGIDAILEQMRRKAMKQGFEFNIMVVGQSGLGKSTLINTLFKSKVSRKSVQPASEERIPKTIEIKSITHDIEEKGVRMKLTVIDTPGFGDHINNENCWQPIMKFINDQYEKYLQEELNVNRKRRIPDTRVHCCIYFIPATGHALRPLDIEFMRRLSKVVNIVPVIAKADTLTLEERDYFKKRITADLLSNDIDVYPQKEFDEDSEDRLVNEKFREMIPFAVVGSDQEYQVNGRRILGRKTKWGTIEVENTTHCEFAYLRDLLIRTHMQNIKDITSSFHFEAYRVKRLNEGRSALSNGMADQEANEM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJZ4</accession>
    <name>A0A452HJZ4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Septin-type G domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="8">Belongs to the TRAFAC class TrmE-Era-EngA-EngB-Septin-like GTPase superfamily. Septin GTPase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJZ4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017405.1">
        <property type="protein sequence ID" value="ENSGAGP00000015234.1"/>
        <property type="gene ID" value="ENSGAGG00000011502.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005525">
        <property type="term" value="F:GTP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd01850">
        <property type="entry name" value="CDC_Septin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000143">
        <property type="entry name" value="septin-9 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR030379">
        <property type="entry name" value="G_SEPTIN_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016491">
        <property type="entry name" value="Septin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18884">
        <property type="entry name" value="SEPTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00735">
        <property type="entry name" value="Septin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51719">
        <property type="entry name" value="G_SEPTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="3" id="KW-0132">Cell division</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="5 8" id="KW-0342">GTP-binding</keyword>
    <keyword evidence="4 8" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Septin-type G" evidence="10">
        <location>
            <begin position="282"/>
            <end position="554"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="1"/>
            <end position="172"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="222"/>
            <end position="261"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="20"/>
            <end position="47"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="97"/>
            <end position="109"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="116"/>
            <end position="134"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="RuleBase" id="RU004560"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS51719"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015234.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="571" mass="64487" checksum="B48DF8DA568E14F1" modified="2019-05-08" version="1">MKKSVHSALKRSFEVEEVDQSLNASTPQRRTPNPLLRSTVSSSTQKFQDLGARNSETPARHADSPGQRSPKSSLRRVELSGPKLPEPVSRRTEISIDISSKQVENSTPGLSRFGLKRAEGLGHKPPEPTPRRTEITIGKPQEPAVRRMETPTSKIPELPQRKAETASAPVPDVATKRVEIQVAKSPEVPVPPVRQVENLEPSLASQHLQPESKPQPVMVEIPPKSPEGPEAVPSYTPSMTDAARDTGLKQAAPPRPEKPAPDFSYVGIDAILEQMRRKAMKQGFEFNIMVVGQSGLGKSTLINTLFKSKVSRKSVQPASEERIPKTIEIKSITHDIEEKGVRMKLTVIDTPGFGDHINNENCWQPIMKFINDQYEKYLQEELNVNRKRRIPDTRVHCCIYFIPATGHALRPLDIEFMRRLSKVVNIVPVIAKADTLTLEERDYFKKRITADLLSNDIDVYPQKEFDEDSEDRLVNEKFREMIPFAVVGSDQEYQVNGRRILGRKTKWGTIEVENTTHCEFAYLRDLLIRTHMQNIKDITSSFHFEAYRVKRLNEGRSALSNGMADQEANEM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJZ5</accession>
    <name>A0A452HJZ5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Ubiquitin carboxyl-terminal hydrolase 47</fullName>
            <ecNumber evidence="2">3.4.19.12</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="9">Deubiquitinating enzyme 47</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="8">Ubiquitin thioesterase 47</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="10">Ubiquitin-specific-processing protease 47</fullName>
        </alternativeName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>Thiol-dependent hydrolysis of ester, thioester, amide, peptide and isopeptide bonds formed by the C-terminal Gly of ubiquitin (a 76-residue protein attached to proteins as an intracellular targeting signal).</text>
            <dbReference type="EC" id="3.4.19.12"/>
        </reaction>
    </comment>
    <dbReference type="EC" id="3.4.19.12" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452HJZ5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017411.1">
        <property type="protein sequence ID" value="ENSGAGP00000015240.1"/>
        <property type="gene ID" value="ENSGAGG00000011497.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004843">
        <property type="term" value="F:cysteine-type deubiquitinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016579">
        <property type="term" value="P:protein deubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd02659">
        <property type="entry name" value="peptidase_C19C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd17039">
        <property type="entry name" value="Ubl_ubiquitin_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.70.10">
        <property type="entry name" value="Cysteine proteinases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038765">
        <property type="entry name" value="Papain-like_cys_pep_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050164">
        <property type="entry name" value="Peptidase_C19"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001394">
        <property type="entry name" value="Peptidase_C19_UCH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028889">
        <property type="entry name" value="USP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045578">
        <property type="entry name" value="USP47_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018200">
        <property type="entry name" value="USP_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24006">
        <property type="entry name" value="UBIQUITIN CARBOXYL-TERMINAL HYDROLASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24006:SF702">
        <property type="entry name" value="UBIQUITIN CARBOXYL-TERMINAL HYDROLASE 47"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00443">
        <property type="entry name" value="UCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19718">
        <property type="entry name" value="USP47_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54001">
        <property type="entry name" value="Cysteine proteinases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00972">
        <property type="entry name" value="USP_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00973">
        <property type="entry name" value="USP_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50235">
        <property type="entry name" value="USP_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="11" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="3" id="KW-0645">Protease</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0788">Thiol protease</keyword>
    <keyword evidence="4" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="USP" evidence="13">
        <location>
            <begin position="88"/>
            <end position="464"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="30"/>
            <end position="61"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="325"/>
            <end position="352"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="734"/>
            <end position="755"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="781"/>
            <end position="813"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="828"/>
            <end position="925"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="11">
        <location>
            <begin position="478"/>
            <end position="505"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="330"/>
            <end position="352"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="734"/>
            <end position="750"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="784"/>
            <end position="803"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="829"/>
            <end position="839"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="841"/>
            <end position="851"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="854"/>
            <end position="870"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="898"/>
            <end position="907"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="12">
        <location>
            <begin position="908"/>
            <end position="921"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000707"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00012759"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022807"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00026136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00029910"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00030277"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00032453"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50235"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015240.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1276" mass="146052" checksum="B4DA183C177A2005" modified="2019-05-08" version="1">MTPLDQTSDKSILEARFEVGKKNFLRLTDKDGEQPHIMSEESGTTDDGNQERFIGPLPTEPSLGSTNDYVCQNYSYSSILSKSETGYVGLVNQAMTCYLNSLLQTLFMTPEFRNALYKWEFEESEEDAVMSIPYQLQRLFVLLQTSKKRAIETTDVTRSFGWDSSEAWQQHDVQELCRVMFDALEQKWKQTEQADLINQLYQGKLKDYVRCLECGYEGWRIDTYLDIPLVIRPYGSNQAFASVEEALHAFIQPEILDGPNQYFCERCKKKCDARKGLRFLHFPYLLTLQLKRFDFDYTTMHRIKLNDRMTFPEELDMSTFIDVEDEKSPQTESCTDSGAENEGSCHSDQMSNDFSNDDGVDEGICLESNSGAERISKVGIEKNSLLYELFSVMVHSGSAAGGHYYACIKSFNDEQWYSFNDQHVSKITQEDIKKTYGGSSGSRGYYSSAFASSTNAYMLIYRLKDPTRNAKFLDVNEYPEHIKQLVQKEKELEEQEKKQREIERNTCKIKLFCMHPVKQIMMENKLEVHKDKTLKEAVKIAYKIMDIEEAVPLDCCRLVKYDEFHDYLERSFEEEEEDIPMGLLLGGVKSTYMFDLLLETRTPDQVFQCYKPGEVMVKVHVVDLKTESVAPSISVRAYLNQTVTEFKQLISKAVCLPAETMRVVLERCYNDLRLLNVGNKTLKAEGFFRSNKVFVESSESLDRQMAYTDSHLWKLLDRHANTIRLYVSLPEQSPGSQFKRTVNQKASGDSGNLDEACARVKGPLGNMRSVEAVLEESTEKLKSLSLQQEQEGDNADSSKSTEASDFESIESPLNEVDSLASAENRELENQIQISDPENFQSEERSDSDVNNDRSTSSVDSDILSSSHSSDTLCNVDSAPMPLANGLDSHSITSSRRSKANEGKKETWDTAEDDSGTDSEYDESGKSTGEAQYMYFKAEPYASDEGSGEGQKWLLVHVDKRITLSSFKQHLEPFVGVLSSHFKVFRVYASNQEFESVRLNETLSSFSDDNKITIRLGRALKKGEYRVKVYQLLINEPEPCKFLLDAVFAKGMTVRQSKEELLPQLREQCGLDLTIDRFRLRKKTWKNPGTVFLDYHVYEEDINISSNWEVFLEILDGAEKMKSMSQLAVLSRRWRPSEMKLDSFQEVVLESSSVDELKEKLSEISGIPLEYIEFAKGRGTFPCDISVLEIHQDLDWNPKVSALNVWPLYICDDGAVIFYRDKTEELIELTDEQRNELMKKESSRLQKTGHRVTYSPRKEKALKIYLDGAPNRESTQD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJZ7</accession>
    <name>A0A452HJZ7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Septin-type G domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="8">Belongs to the TRAFAC class TrmE-Era-EngA-EngB-Septin-like GTPase superfamily. Septin GTPase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJZ7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017413.1">
        <property type="protein sequence ID" value="ENSGAGP00000015242.1"/>
        <property type="gene ID" value="ENSGAGG00000011502.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005525">
        <property type="term" value="F:GTP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd01850">
        <property type="entry name" value="CDC_Septin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000143">
        <property type="entry name" value="septin-9 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR030379">
        <property type="entry name" value="G_SEPTIN_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016491">
        <property type="entry name" value="Septin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18884">
        <property type="entry name" value="SEPTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00735">
        <property type="entry name" value="Septin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51719">
        <property type="entry name" value="G_SEPTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="3" id="KW-0132">Cell division</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="5 8" id="KW-0342">GTP-binding</keyword>
    <keyword evidence="4 8" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Septin-type G" evidence="10">
        <location>
            <begin position="293"/>
            <end position="565"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="29"/>
            <end position="183"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="232"/>
            <end position="272"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="31"/>
            <end position="58"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="108"/>
            <end position="120"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="127"/>
            <end position="145"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="RuleBase" id="RU004560"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS51719"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015242.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="582" mass="65617" checksum="6F286ECAF0976E2C" modified="2019-05-08" version="1">MSDPAVNAHLEGIISDFEALKRSFEVEEVDQSLNASTPQRRTPNPLLRSTVSSSTQKFQDLGARNSETPARHADSPGQRSPKSSLRRVELSGPKLPEPVSRRTEISIDISSKQVENSTPGLSRFGLKRAEGLGHKPPEPTPRRTEITIGKPQEPAVRRMETPTSKIPELPQRKAETASAPVPDVATKRVEIQVAKSPEVPVPPVRQVENLEPSLASQHLQPESKPQPVMVEIPPKSPEGPEAVPSYTPSMTDAARDTGLKQAAPPRPEKPAPDFSYVGIDAILEQMRRKAMKQGFEFNIMVVGQSGLGKSTLINTLFKSKVSRKSVQPASEERIPKTIEIKSITHDIEEKGVRMKLTVIDTPGFGDHINNENCWQPIMKFINDQYEKYLQEELNVNRKRRIPDTRVHCCIYFIPATGHALRPLDIEFMRRLSKVVNIVPVIAKADTLTLEERDYFKKRITADLLSNDIDVYPQKEFDEDSEDRLVNEKFREMIPFAVVGSDQEYQVNGRRILGRKTKWGTIEVENTTHCEFAYLRDLLIRTHMQNIKDITSSFHFEAYRVKRLNEGRSALSNGMADQEANEM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJZ8</accession>
    <name>A0A452HJZ8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Ubiquitin carboxyl-terminal hydrolase 47</fullName>
            <ecNumber evidence="2">3.4.19.12</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="9">Deubiquitinating enzyme 47</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="8">Ubiquitin thioesterase 47</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="10">Ubiquitin-specific-processing protease 47</fullName>
        </alternativeName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>Thiol-dependent hydrolysis of ester, thioester, amide, peptide and isopeptide bonds formed by the C-terminal Gly of ubiquitin (a 76-residue protein attached to proteins as an intracellular targeting signal).</text>
            <dbReference type="EC" id="3.4.19.12"/>
        </reaction>
    </comment>
    <dbReference type="EC" id="3.4.19.12" evidence="2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017415.1">
        <property type="protein sequence ID" value="ENSGAGP00000015243.1"/>
        <property type="gene ID" value="ENSGAGG00000011497.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004843">
        <property type="term" value="F:cysteine-type deubiquitinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016579">
        <property type="term" value="P:protein deubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd02659">
        <property type="entry name" value="peptidase_C19C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd17039">
        <property type="entry name" value="Ubl_ubiquitin_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.70.10">
        <property type="entry name" value="Cysteine proteinases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038765">
        <property type="entry name" value="Papain-like_cys_pep_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050164">
        <property type="entry name" value="Peptidase_C19"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001394">
        <property type="entry name" value="Peptidase_C19_UCH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028889">
        <property type="entry name" value="USP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045578">
        <property type="entry name" value="USP47_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018200">
        <property type="entry name" value="USP_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24006">
        <property type="entry name" value="UBIQUITIN CARBOXYL-TERMINAL HYDROLASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24006:SF702">
        <property type="entry name" value="UBIQUITIN CARBOXYL-TERMINAL HYDROLASE 47"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25985">
        <property type="entry name" value="Ubiquitin_USP47_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00443">
        <property type="entry name" value="UCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19718">
        <property type="entry name" value="USP47_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54001">
        <property type="entry name" value="Cysteine proteinases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00972">
        <property type="entry name" value="USP_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00973">
        <property type="entry name" value="USP_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50235">
        <property type="entry name" value="USP_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="11" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="3" id="KW-0645">Protease</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0788">Thiol protease</keyword>
    <keyword evidence="4" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="USP" evidence="13">
        <location>
            <begin position="168"/>
            <end position="544"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="112"/>
            <end position="141"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="405"/>
            <end position="432"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="814"/>
            <end position="835"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="861"/>
            <end position="893"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="908"/>
            <end position="1005"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="11">
        <location>
            <begin position="558"/>
            <end position="585"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="410"/>
            <end position="432"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="814"/>
            <end position="830"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="864"/>
            <end position="883"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="909"/>
            <end position="919"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="921"/>
            <end position="931"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="934"/>
            <end position="950"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="978"/>
            <end position="987"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="12">
        <location>
            <begin position="988"/>
            <end position="1001"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000707"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00012759"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022807"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00026136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00029910"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00030277"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00032453"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50235"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015243.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1356" mass="154862" checksum="5E619D5E05622832" modified="2019-05-08" version="1">MVPNEENQLVPKEIESAADEPRVLCIIQDTTNSKTVNERITLNLPASTPLRRLFEDVASKVGYVNGTFDLVWGNGINVADMTPLDQTSDKSILEARFEVGKKNFLRLTDKDGEQPHIMSEESGTTDDGNQERFIGPLPTEPSLGSTNDYVCQNYSYSSILSKSETGYVGLVNQAMTCYLNSLLQTLFMTPEFRNALYKWEFEESEEDAVMSIPYQLQRLFVLLQTSKKRAIETTDVTRSFGWDSSEAWQQHDVQELCRVMFDALEQKWKQTEQADLINQLYQGKLKDYVRCLECGYEGWRIDTYLDIPLVIRPYGSNQAFASVEEALHAFIQPEILDGPNQYFCERCKKKCDARKGLRFLHFPYLLTLQLKRFDFDYTTMHRIKLNDRMTFPEELDMSTFIDVEDEKSPQTESCTDSGAENEGSCHSDQMSNDFSNDDGVDEGICLESNSGAERISKVGIEKNSLLYELFSVMVHSGSAAGGHYYACIKSFNDEQWYSFNDQHVSKITQEDIKKTYGGSSGSRGYYSSAFASSTNAYMLIYRLKDPTRNAKFLDVNEYPEHIKQLVQKEKELEEQEKKQREIERNTCKIKLFCMHPVKQIMMENKLEVHKDKTLKEAVKIAYKIMDIEEAVPLDCCRLVKYDEFHDYLERSFEEEEEDIPMGLLLGGVKSTYMFDLLLETRTPDQVFQCYKPGEVMVKVHVVDLKTESVAPSISVRAYLNQTVTEFKQLISKAVCLPAETMRVVLERCYNDLRLLNVGNKTLKAEGFFRSNKVFVESSESLDRQMAYTDSHLWKLLDRHANTIRLYVSLPEQSPGSQFKRTVNQKASGDSGNLDEACARVKGPLGNMRSVEAVLEESTEKLKSLSLQQEQEGDNADSSKSTEASDFESIESPLNEVDSLASAENRELENQIQISDPENFQSEERSDSDVNNDRSTSSVDSDILSSSHSSDTLCNVDSAPMPLANGLDSHSITSSRRSKANEGKKETWDTAEDDSGTDSEYDESGKSTGEAQYMYFKAEPYASDEGSGEGQKWLLVHVDKRITLSSFKQHLEPFVGVLSSHFKVFRVYASNQEFESVRLNETLSSFSDDNKITIRLGRALKKGEYRVKVYQLLINEPEPCKFLLDAVFAKGMTVRQSKEELLPQLREQCGLDLTIDRFRLRKKTWKNPGTVFLDYHVYEEDINISSNWEVFLEILDGAEKMKSMSQLAVLSRRWRPSEMKLDSFQEVVLESSSVDELKEKLSEISGIPLEYIEFAKGRGTFPCDISVLEIHQDLDWNPKVSALNVWPLYICDDGAVIFYRDKTEELIELTDEQRNELMKKESSRLQKTGHRVTYSPRKEKALKIYLDGAPNRESTQD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJZ9</accession>
    <name>A0A452HJZ9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="19">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="14">Interacts with SHTN1; the interaction occurs in axonal growth cones. Interacts with isoform 2 of BSG.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Membrane</location>
            <topology evidence="2">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the immunoglobulin superfamily. L1/neurofascin/NgCAM family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000017410.1">
        <property type="protein sequence ID" value="ENSGAGP00000015239.1"/>
        <property type="gene ID" value="ENSGAGG00000006819.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030424">
        <property type="term" value="C:axon"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009986">
        <property type="term" value="C:cell surface"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061564">
        <property type="term" value="P:axon development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098609">
        <property type="term" value="P:cell-cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="CDD" id="cd05845">
        <property type="entry name" value="IgI_2_L1-CAM_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000658">
        <property type="entry name" value="Neural cell adhesion molecule L1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000367">
        <property type="entry name" value="Neural cell adhesion molecule L1-like protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000005">
        <property type="entry name" value="Neuronal cell adhesion molecule"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000028">
        <property type="entry name" value="Neuronal cell adhesion molecule"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000038">
        <property type="entry name" value="Neuronal cell adhesion molecule"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000078">
        <property type="entry name" value="Neuronal cell adhesion molecule"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000347">
        <property type="entry name" value="Neuronal cell adhesion molecule"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013098">
        <property type="entry name" value="Ig_I-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013151">
        <property type="entry name" value="Immunoglobulin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026966">
        <property type="entry name" value="Neurofascin/L1/NrCAM_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR44170:SF44">
        <property type="entry name" value="L1 CELL ADHESION MOLECULE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR44170">
        <property type="entry name" value="PROTEIN SIDEKICK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13882">
        <property type="entry name" value="Bravo_FIGEY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07679">
        <property type="entry name" value="I-set"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00047">
        <property type="entry name" value="ig"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13927">
        <property type="entry name" value="Ig_3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="6"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="10 16" id="KW-0472">Membrane</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="5 16" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9 16" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="1093"/>
            <end position="1114"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="17">
        <location>
            <begin position="1"/>
            <end position="127"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="17">
        <location>
            <begin position="134"/>
            <end position="221"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="17">
        <location>
            <begin position="235"/>
            <end position="323"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="17">
        <location>
            <begin position="328"/>
            <end position="415"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="17">
        <location>
            <begin position="420"/>
            <end position="504"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="17">
        <location>
            <begin position="513"/>
            <end position="603"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="757"/>
            <end position="853"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="858"/>
            <end position="960"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="18">
        <location>
            <begin position="964"/>
            <end position="1059"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="620"/>
            <end position="653"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="740"/>
            <end position="768"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="1040"/>
            <end position="1075"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="1124"/>
            <end position="1163"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="1193"/>
            <end position="1223"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="15">
        <location>
            <begin position="1124"/>
            <end position="1143"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="15">
        <location>
            <begin position="1209"/>
            <end position="1223"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00008588"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00063896"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015239.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1223" mass="134784" checksum="F9208CE57A409D70" modified="2019-05-08" version="1">MGEXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXPPPRVSTDLLCLPTPHSSPFRWTKDGRWFDPRQEAGVTVVEGSGTLYVNNSNAGRHQGVYRCYASNDLGTAMGPEIRVIAESTPQWPKEKVLPLEVEEGESVILPCNPPASAMPPKIYWLNSRIVHIAQDERVSMGQDGYLYFANVRRSDSHPDYICHAHYLGPRTIIQKEPIELRVRPTNSVKFRQPRLMVPPGESSSHVALRGSSLALECIAEGFPTPTVEWIRLNGRLPAERVALENSQKTLRIRDVVEEDDGEYQCLARNTQGTARHTYTVTVEAAPYWAQQPHSRVYGPGENVRLDCSVYGKPNPLVSWRINGVPWGELAPDPTRTLRSGALILSQVQPNDSLVTQCEAVNHHGRLLANAYIYVIQLPARILTPDKQRYAVVENQTASLDCQTFGAPMPSVEWFTGRMEPALQDERAFVHTNGSLRLPRAQRQDTGTYVCLAQNDQTNVTISAWLEVKGATRIELGPQSAVVKKSSNVTFQCLVRFDETLQERGVEWWRDGRPIRESDDSDKYTIEPGWLRISGVDYTDQGVYSCRAHTELDAVEESAELRVVGTCLERWVKLSWTPGDDHNSPIESKGTPRPSDPRASPPQPHCPCCSSRHPLPQAGQSWGASGQFQVGTGGFGGQVVGGVSEEVLEFVVEAEDGLRDPGTWRDVSTVPGHQPWAQLHLVPYGDFRFRVRAANRFGRGEPSAASAPVRTSPAAPERYPEGVAGEGNETNNMLITWKPLNRSDWNAPDLSYRVQWRRLGKGGGWSEETVSGPPLLVTRTPTFVPYEIKVQAVNPIGKGPEPPTVVGYSGEDVPQVNPENVGVEIFNSTSVLVSWTLPNRDQLRGHLRGFRVFYWRLGSVGERSRRXPPPLVLTVEGEASWARLGGLRPWSWYRLYVVVFNGRSDGPHSEDIDFSTPEGVPSPPSSLLLERVSDTVLALEWTAPQFPNGALTEYLLQYQQGESRGRWAGGHKGWNRGSYGWGGLAIGQVQDGYWLDPTPSGEQCSRRGKLRHTQASSPRVLGPGAGSLGNAARRSAPPVLTTDQRSFATEGWFIGFICAVVLLLLLLLILCFVKRSKGGKYSVKDKEDIQVDSEARPMKDETFGEYSDNEEKPFGSSQPSLNGALPALGSEDSLADYGGSVDVQFNEDGSFIGQYSGHTGKEPAGNNSSGATSPTTATAALE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK83</accession>
    <name>A0A452HK83_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">BTB domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HK83"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017507.1">
        <property type="protein sequence ID" value="ENSGAGP00000015328.1"/>
        <property type="gene ID" value="ENSGAGG00000011556.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="CDD" id="cd18481">
        <property type="entry name" value="BACK_KBTBD4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18272">
        <property type="entry name" value="BTB_POZ_KBTBD4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.420">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.120.10.80">
        <property type="entry name" value="Kelch-type beta propeller"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.710.10">
        <property type="entry name" value="Potassium Channel Kv1.1, Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011705">
        <property type="entry name" value="BACK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017096">
        <property type="entry name" value="BTB-kelch_protein"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000210">
        <property type="entry name" value="BTB/POZ_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042884">
        <property type="entry name" value="KBTBD4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042950">
        <property type="entry name" value="KBTBD4_BACK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042949">
        <property type="entry name" value="KBTBD4_BTB_POZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015915">
        <property type="entry name" value="Kelch-typ_b-propeller"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011498">
        <property type="entry name" value="Kelch_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011333">
        <property type="entry name" value="SKP1/BTB/POZ_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47195">
        <property type="entry name" value="KELCH REPEAT AND BTB DOMAIN-CONTAINING PROTEIN 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47195:SF1">
        <property type="entry name" value="KELCH REPEAT AND BTB DOMAIN-CONTAINING PROTEIN 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07707">
        <property type="entry name" value="BACK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00651">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07646">
        <property type="entry name" value="Kelch_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF037037">
        <property type="entry name" value="Kelch-like_protein_gigaxonin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00875">
        <property type="entry name" value="BACK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00225">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF117281">
        <property type="entry name" value="Kelch motif"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54695">
        <property type="entry name" value="POZ domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50097">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0880">Kelch repeat</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="BTB" evidence="3">
        <location>
            <begin position="46"/>
            <end position="113"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022441"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS50097"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015328.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="519" mass="58261" checksum="E62CF0D00AB86236" modified="2019-05-08" version="1">MDSSEETGGSSTEENYFVNYTFTDRSHSGRVAQGIMKLCLEDELFADVTISVEGKEFQLHRLVLSAQSCFFRSMFTSNLKEAHNRVIELQDVSESVFQLLVDYIYHGTVKLRAEELQETYEVADMYQLTALFEECSRFLARTVQVRNCLQVMWLADQHSDMELYTAAKHCAKSHLSQLQDTEEFLHLPLRLLTDILTDGVPCSQNPTVAIETWINFNKEERAGFLEVLRSSLKEIGENVHIYLIGKESSRTHSLAVSLHCADDDSISVSGQNSLCHQITAACKHGSDLYVVGGSIPRRMWKCNNATVDWEWCAPLPRDRLQHTLVSVPSKDAIYSLGGKTLQDTLSNAVIYYRVRDNVWTETSQLEVAVSGAAGANLNGIIYLLGGEENDLDFFTKPSRLIQCYDTNTEKCHVKPYVLPFAGHMHAAVHKDLVFIVAEGDSLLCYNPLLDSFTRLCLPDAWSSVPSLWKIASCNGSIYVFRDRYKKGDANTFKLNPATSVVTVTSGIKVLLTNLQFVLA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK85</accession>
    <name>A0A452HK85_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">protein-serine/threonine phosphatase</fullName>
            <ecNumber evidence="3">3.1.3.16</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="8 15">
            <text>O-phospho-L-threonyl-[protein] + H2O = L-threonyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:47004"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="EC" id="3.1.3.16"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the protein-tyrosine phosphatase family.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.16" evidence="3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017487.1">
        <property type="protein sequence ID" value="ENSGAGP00000015309.1"/>
        <property type="gene ID" value="ENSGAGG00000011542.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003779">
        <property type="term" value="F:actin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004722">
        <property type="term" value="F:protein serine/threonine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030837">
        <property type="term" value="P:negative regulation of actin filament polymerization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd14569">
        <property type="entry name" value="DSP_slingshot_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd11652">
        <property type="entry name" value="SSH-N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000004">
        <property type="entry name" value="Protein phosphatase Slingshot homolog 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014876">
        <property type="entry name" value="DEK_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000340">
        <property type="entry name" value="Dual-sp_phosphatase_cat-dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043587">
        <property type="entry name" value="Phosphatase_SSH-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043588">
        <property type="entry name" value="SSH-N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016130">
        <property type="entry name" value="Tyr_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020422">
        <property type="entry name" value="TYR_PHOSPHATASE_DUAL_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45864:SF3">
        <property type="entry name" value="PROTEIN PHOSPHATASE SLINGSHOT HOMOLOG 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45864">
        <property type="entry name" value="SLINGSHOT PROTEIN PHOSPHATASE HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08766">
        <property type="entry name" value="DEK_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00782">
        <property type="entry name" value="DSPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23040">
        <property type="entry name" value="PH_SSH1-like_1st"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00195">
        <property type="entry name" value="DSPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51998">
        <property type="entry name" value="DEK_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00383">
        <property type="entry name" value="TYR_PHOSPHATASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50054">
        <property type="entry name" value="TYR_PHOSPHATASE_DUAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="5" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="6" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="DEK-C" evidence="12">
        <location>
            <begin position="169"/>
            <end position="224"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="10">
        <location>
            <begin position="228"/>
            <end position="369"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="11">
        <location>
            <begin position="293"/>
            <end position="347"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="549"/>
            <end position="568"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="587"/>
            <end position="608"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="621"/>
            <end position="664"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="718"/>
            <end position="789"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="849"/>
            <end position="899"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="1103"/>
            <end position="1130"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="1174"/>
            <end position="1218"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="1291"/>
            <end position="1329"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="554"/>
            <end position="567"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="587"/>
            <end position="603"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="630"/>
            <end position="641"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="738"/>
            <end position="748"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="9">
        <location>
            <begin position="762"/>
            <end position="772"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="849"/>
            <end position="858"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="1110"/>
            <end position="1128"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="1199"/>
            <end position="1209"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="9">
        <location>
            <begin position="1300"/>
            <end position="1313"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009580"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00013081"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
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    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
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    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00048336"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS51998"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015309.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1329" mass="147533" checksum="7D485E8D63A7A8FE" modified="2019-05-08" version="1">MFILLRPEDNIRLAVRLESTYQNRTRYMVVVSTNGRQDTEENIVLGMDFSSNDSSTCTMGLVLPLWSDTLIHLDGDGGFSVSTDNRVHVFKPVSVQAMWSALQSLHKACEVARSHNYYPGSLFLTWVSYYESHIDSDQSSVNEWNAMQDVQSHRPDSPALFTDVPTERERTERLIKTKLREIMMQKDLENITSKEIRTELEMQMVCNLREFKEFIDNEMIVILGQMDSPTQIFDHVFLGSEWNASNLEDLQNKGVRYILNVTREIDNFFPGIFEYHNIRVYDEEATDLLAYWNDTYKFISKAKKNGSKCLVHCKMGVSRSASTVIAYAMKEYGWDLDRAYEHVKERRAVTKPNPSFMRQLEEYQGILLASKQRHNKLWRSHSDSDLSEHHQPICKPGMELNKKEITTSAEQLSEPTANDNHQPALPVFSVGPHVGRLLQPDTDVIEELGAKERRIHLEFAARDFRAEQIEDELNLNNINRCPAGCCLEESEFPLDNCHSSEALVQLKEPLEMATAFPDLTVDDLETDALKTDLNVHLVPMEEFTACLKDLPRSPDQNSPSPQLSSHLETTDRIDFLSALEKFEELSQESRSHACSHTRMDEQGGGRNGVSKMSVLEVLPAGTAVDTQRNSSASSSPAAPEESSTDEEQPKDISELTTLGPLTRSHSENAISVKEIITEIESINQGIGPCQQKMEVSANHAQAPKRSTVHELPVEVVWTSENKPGKAEQSERACTAEQETAKDLPKPDQEGVPTFQLPTCASDVEESDPAEDQQEPRVQGLNPGSKWCPGSVKRATLEFEERLRQEQEHQHVPLICLLPTRKNSKNDSAAAELAPKGRTEELTWELDPAHGENESDQARAKSPAAEGALPHGPELPGGTHLSPPLAPCFVQPAQEPPLAEPGAEQEVLRTVVSLNGSGEPALPCHLPKRIEIIEYPHMGTPPDLRERDSSREQGTLENAAALLPVDENLNPSVCAETAPVHPDALPLRNPSVLEHKAIGQATFTLGSPEEEYGAAPINLDTSSFTGQVRRLPSAGLDDLWHQRVVHLEGVTEQSTSADDEPVAPSSLSRNGHVLCEASDSAFVELSNEDLDFLSDSTEAVDVSFLPGGLPHSSNSPSIKDSSEAPSSLKQRAKELEARVRQAGLTTPSQMKRSASLAKLGCLELSRDDLTKRELASSHASLTGPDLLPASPRTGQGKCDALPEHGLEGPPEKTCASSPLIRQDSKPTKHFVEQLKTTECIVQSQPVERPPVQYAKEFGSAPQNLYPSTDPRLTSSEEGLPLLQMQVLDSRLPAQSLAGAPRHQRSRTHPLRRLKMATDKRRTSNPFYNTM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKA1</accession>
    <name>A0A452HKA1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">SH2B adapter protein 2</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="11">Adapter protein with pleckstrin homology and Src homology 2 domains</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="10">SH2 and PH domain-containing adapter protein APS</fullName>
        </alternativeName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="8">Homodimer. Interacts with KIT/c-KIT, SHC1, EPOR, PDGFR, VAV1 and VAV3. Interacts (via N-terminal region) with SHC1. Interacts (via the phosphorylated C-terminus) with GRB2. Interacts (via its SH2 domain) with EPOR, INSR and KIT. Interacts with GRB2 after B-cell antigen receptor stimulation. Interacts (via PH domain) with VAV3. Interacts with NTRK1, NTRK2 and NTRK3 (phosphorylated); after stimulation of the receptor by its extracellular ligand and subsequent autophosphorylation of the receptor. Binds INSR, GRB2, ASB6 and CAP. Insulin stimulation leads to dissociation of CAP. Binds CBS only when SH2B2/APS has become phosphorylated. INSR binding does not depend on the phosphorylation of SH2B2/APS.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the SH2B adapter family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKA1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017503.1">
        <property type="protein sequence ID" value="ENSGAGP00000015324.1"/>
        <property type="gene ID" value="ENSGAGG00000011554.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005068">
        <property type="term" value="F:transmembrane receptor protein tyrosine kinase adaptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035556">
        <property type="term" value="P:intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd01231">
        <property type="entry name" value="PH_SH2B_family"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd10346">
        <property type="entry name" value="SH2_SH2B_family"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.505.10:FF:000008">
        <property type="entry name" value="SH2B adapter protein 1 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000187">
        <property type="entry name" value="SH2B adapter protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.110">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.505.10">
        <property type="entry name" value="SH2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015012">
        <property type="entry name" value="Phe_ZIP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036290">
        <property type="entry name" value="Phe_ZIP_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000980">
        <property type="entry name" value="SH2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036860">
        <property type="entry name" value="SH2_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR030523">
        <property type="entry name" value="SH2B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035057">
        <property type="entry name" value="SH2B1_SH2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10872">
        <property type="entry name" value="SH2B ADAPTER PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10872:SF3">
        <property type="entry name" value="SH2B ADAPTER PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00169">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08916">
        <property type="entry name" value="Phe_ZIP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00017">
        <property type="entry name" value="SH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00401">
        <property type="entry name" value="SH2DOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00233">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00252">
        <property type="entry name" value="SH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109805">
        <property type="entry name" value="Phenylalanine zipper"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55550">
        <property type="entry name" value="SH2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50001">
        <property type="entry name" value="SH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-0472">Membrane</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6 12" id="KW-0727">SH2 domain</keyword>
    <feature type="domain" description="PH" evidence="15">
        <location>
            <begin position="237"/>
            <end position="352"/>
        </location>
    </feature>
    <feature type="domain" description="SH2" evidence="14">
        <location>
            <begin position="483"/>
            <end position="581"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="100"/>
            <end position="165"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="178"/>
            <end position="212"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="592"/>
            <end position="612"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="13">
        <location>
            <begin position="103"/>
            <end position="121"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="13">
        <location>
            <begin position="131"/>
            <end position="154"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="13">
        <location>
            <begin position="194"/>
            <end position="208"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004370"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00010220"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022999"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00065784"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00073702"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00075397"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00080463"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00191"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50001"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015324.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="626" mass="67473" checksum="ED73B5ECA4296567" modified="2019-05-08" version="1">MNGSVAPPGVVEAAAPLPSPGWRDFCEAHARAAALDFARRFRAFLSENPQFAAPGAEAAFSRRFAEHFVEHFEAEVSRACSGDSPPRCDIAPFTGTHHCPGGASSSSTRDLSETCSDSSVASPVEPQPGLASGLSSSQSRSSEDVSASAATASTKPKLKKRFSLRNVSRSVRGSMRGILQWKSSAESPTEEEAGGANTNSNSSGSGAADSLPTERWTHKFERLRLSKAPAPKVELSSVRREGLLNYIVADDSGGGGGSRARWQKCRLLLRKAGKGEGEGYLLEFYIPPKSTKARVSILCSAVTDVRTTTPLEMPDKENTFVLKMENSLEYILETVDSLQMRSWLADIQDCMSPGENVDSMDLPCLNHSESMPSRELPLVPSESNEQLSQGAYGGLSERPSASISPSSISIAASHFNSMELLPPELPPRIPIDEAAERLQGPYPPGLLHTPFPDTPDTTGSFLFQGEPDPGGGDPEHPLSEYPWFHGTLSRLKAAQLVLAGGPSSHGVFLVRQSETRRGEYVLTFNFQGKAKHLRLSLNEDGQCRVQHLWFQTIFDMLEHFRVHPIPLESGGSSDVTLVSYVVASQRLHGRDRAGSRAAACEPTQRSPDSPGLVSIGANDCLAEHLP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKB2</accession>
    <name>A0A452HKB2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Phosphatidylinositol-binding clathrin assembly protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="18">Cytoplasmic adapter protein that plays a critical role in clathrin-mediated endocytosis which is important in processes such as internalization of cell receptors, synaptic transmission or removal of apoptotic cells. Recruits AP-2 and attaches clathrin triskelions to the cytoplasmic side of plasma membrane leading to clathrin-coated vesicles (CCVs) assembly. Furthermore, regulates clathrin-coated vesicle size and maturation by directly sensing and driving membrane curvature. In addition to binding to clathrin, mediates the endocytosis of small R-SNARES (Soluble NSF Attachment Protein REceptors) between plasma membranes and endosomes including VAMP2, VAMP3, VAMP4, VAMP7 or VAMP8. In turn, PICALM-dependent SNARE endocytosis is required for the formation and maturation of autophagic precursors. Modulates thereby autophagy and the turnover of autophagy substrates such as MAPT/TAU or amyloid precursor protein cleaved C-terminal fragment (APP-CTF).</text>
    </comment>
    <comment type="subunit">
        <text evidence="19">Binds to clathrin; involves primarily the C-terminal sequences, but the full-length protein is required for full binding capacity. Binds phosphatidylinositol 4,5- bisphosphate. Interacts with PIMREG; this interaction may change the subcellular location into the nucleus. Interacts with AP2A1 (via its alpha-appendage domain). Interacts (via N-terminus) with VAMP2; VAMP3; VAMP7 and VAMP8 (Via N-terminus). Interacts with LC3/MAP1LC3A.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasmic vesicle</location>
            <location evidence="2">Clathrin-coated vesicle</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Golgi apparatus</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="5">Membrane</location>
            <location evidence="5">Clathrin-coated pit</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the PICALM/SNAP91 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKB2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017540.1">
        <property type="protein sequence ID" value="ENSGAGP00000015358.1"/>
        <property type="gene ID" value="ENSGAGG00000011583.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005905">
        <property type="term" value="C:clathrin-coated pit"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030136">
        <property type="term" value="C:clathrin-coated vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098894">
        <property type="term" value="C:extrinsic component of presynaptic endocytic zone membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008021">
        <property type="term" value="C:synaptic vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005545">
        <property type="term" value="F:1-phosphatidylinositol binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032050">
        <property type="term" value="F:clathrin heavy chain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005546">
        <property type="term" value="F:phosphatidylinositol-4,5-bisphosphate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000149">
        <property type="term" value="F:SNARE binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048268">
        <property type="term" value="P:clathrin coat assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072583">
        <property type="term" value="P:clathrin-dependent endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016185">
        <property type="term" value="P:synaptic vesicle budding from presynaptic endocytic zone membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd16985">
        <property type="entry name" value="ANTH_N_AP180"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.150:FF:000001">
        <property type="entry name" value="phosphatidylinositol-binding clathrin assembly protein-like isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.90:FF:000001">
        <property type="entry name" value="phosphatidylinositol-binding clathrin assembly protein-like isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.90">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.150">
        <property type="entry name" value="ANTH domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011417">
        <property type="entry name" value="ANTH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014712">
        <property type="entry name" value="ANTH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045192">
        <property type="entry name" value="AP180-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013809">
        <property type="entry name" value="ENTH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008942">
        <property type="entry name" value="ENTH_VHS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22951">
        <property type="entry name" value="CLATHRIN ASSEMBLY PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22951:SF11">
        <property type="entry name" value="ENTH DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07651">
        <property type="entry name" value="ANTH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00273">
        <property type="entry name" value="ENTH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48464">
        <property type="entry name" value="ENTH/VHS domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF89009">
        <property type="entry name" value="GAT-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50942">
        <property type="entry name" value="ENTH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12" id="KW-0007">Acetylation</keyword>
    <keyword evidence="7" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="15" id="KW-0168">Coated pit</keyword>
    <keyword evidence="21" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="17" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="10" id="KW-0254">Endocytosis</keyword>
    <keyword evidence="13" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="8" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="14" id="KW-0472">Membrane</keyword>
    <keyword evidence="16" id="KW-0539">Nucleus</keyword>
    <keyword evidence="9" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="11" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="ENTH" evidence="23">
        <location>
            <begin position="14"/>
            <end position="145"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="565"/>
            <end position="589"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="21">
        <location>
            <begin position="318"/>
            <end position="348"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="22">
        <location>
            <begin position="571"/>
            <end position="587"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004132"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004555"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004600"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00008011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022583"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023176"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00055144"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00061829"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00068054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS50942"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015358.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="655" mass="71358" checksum="AD9C534026B40DA8" modified="2019-05-08" version="1">MSGQSITDRIAAAQHSVTGSAISKAVCKATTHEVMGPKKKHLDYLIQCTNEMNVNIPQLADTLFERTVNSSWVVVFKALITTHHLMMYGNERFIQYLASRNTLFNLNNYLDKSAMQGYDMSTFIRRYSKYLNEKALSYRLVAVDFTKMKRGIDGVMRTMNTEKLLKTLPIIQNQLDALLDFDANPNELTNGVINAAFMLLFKDSIRLFAAYNEGVINLLEKYFDMKKTQCKEGLDIYKKFLARMTKLSEFLKVAEQVGIDQGDIPDLTQAPSSLLEALEQHLASLEGKKTKEVSAASRASTLSSAVSTLANTGMSFSKMDEREKQQALEEEQARLQALKEQRLKEISVVSNSTSTSASPSTLSGKSVNTTTAVDLFAAPAPTTSSMPNLSSDLFDLQPAFVPTVQSTPAIATSASSAWGGGPFSSSNGCVGSPSHLDIFDMKPVEEAVKSTVPFVSSTFTSKQTVEQFSGFPLHSAPQSTTSSIINVDFDAVFGGKSTAHEYRTTSEDVLQPTVPTQCQRTVLINQQSGKILANDLDSSLANLVGNLGFGGTPSKKSDMQWNQPTEKKLTGGTNWQAKTSTSTTWNTAPIPPAPHMVPAPMTYPVTTPQVPVYGMVPPQIGAAPLTAPQPMIYTQPGLRPTNPFASVPGPQIQFM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKB5</accession>
    <name>A0A452HKB5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Phosphatidylinositol-binding clathrin assembly protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="18">Cytoplasmic adapter protein that plays a critical role in clathrin-mediated endocytosis which is important in processes such as internalization of cell receptors, synaptic transmission or removal of apoptotic cells. Recruits AP-2 and attaches clathrin triskelions to the cytoplasmic side of plasma membrane leading to clathrin-coated vesicles (CCVs) assembly. Furthermore, regulates clathrin-coated vesicle size and maturation by directly sensing and driving membrane curvature. In addition to binding to clathrin, mediates the endocytosis of small R-SNARES (Soluble NSF Attachment Protein REceptors) between plasma membranes and endosomes including VAMP2, VAMP3, VAMP4, VAMP7 or VAMP8. In turn, PICALM-dependent SNARE endocytosis is required for the formation and maturation of autophagic precursors. Modulates thereby autophagy and the turnover of autophagy substrates such as MAPT/TAU or amyloid precursor protein cleaved C-terminal fragment (APP-CTF).</text>
    </comment>
    <comment type="subunit">
        <text evidence="19">Binds to clathrin; involves primarily the C-terminal sequences, but the full-length protein is required for full binding capacity. Binds phosphatidylinositol 4,5- bisphosphate. Interacts with PIMREG; this interaction may change the subcellular location into the nucleus. Interacts with AP2A1 (via its alpha-appendage domain). Interacts (via N-terminus) with VAMP2; VAMP3; VAMP7 and VAMP8 (Via N-terminus). Interacts with LC3/MAP1LC3A.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasmic vesicle</location>
            <location evidence="2">Clathrin-coated vesicle</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Golgi apparatus</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="5">Membrane</location>
            <location evidence="5">Clathrin-coated pit</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the PICALM/SNAP91 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKB5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017549.1">
        <property type="protein sequence ID" value="ENSGAGP00000015366.1"/>
        <property type="gene ID" value="ENSGAGG00000011583.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005905">
        <property type="term" value="C:clathrin-coated pit"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030136">
        <property type="term" value="C:clathrin-coated vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098894">
        <property type="term" value="C:extrinsic component of presynaptic endocytic zone membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008021">
        <property type="term" value="C:synaptic vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005545">
        <property type="term" value="F:1-phosphatidylinositol binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032050">
        <property type="term" value="F:clathrin heavy chain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005546">
        <property type="term" value="F:phosphatidylinositol-4,5-bisphosphate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000149">
        <property type="term" value="F:SNARE binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048268">
        <property type="term" value="P:clathrin coat assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072583">
        <property type="term" value="P:clathrin-dependent endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016185">
        <property type="term" value="P:synaptic vesicle budding from presynaptic endocytic zone membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd16985">
        <property type="entry name" value="ANTH_N_AP180"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.150:FF:000001">
        <property type="entry name" value="phosphatidylinositol-binding clathrin assembly protein-like isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.90:FF:000001">
        <property type="entry name" value="phosphatidylinositol-binding clathrin assembly protein-like isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.90">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.150">
        <property type="entry name" value="ANTH domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011417">
        <property type="entry name" value="ANTH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014712">
        <property type="entry name" value="ANTH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045192">
        <property type="entry name" value="AP180-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013809">
        <property type="entry name" value="ENTH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008942">
        <property type="entry name" value="ENTH_VHS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22951">
        <property type="entry name" value="CLATHRIN ASSEMBLY PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22951:SF11">
        <property type="entry name" value="ENTH DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07651">
        <property type="entry name" value="ANTH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00273">
        <property type="entry name" value="ENTH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48464">
        <property type="entry name" value="ENTH/VHS domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF89009">
        <property type="entry name" value="GAT-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50942">
        <property type="entry name" value="ENTH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12" id="KW-0007">Acetylation</keyword>
    <keyword evidence="7" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="15" id="KW-0168">Coated pit</keyword>
    <keyword evidence="21" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="17" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="10" id="KW-0254">Endocytosis</keyword>
    <keyword evidence="13" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="8" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="14" id="KW-0472">Membrane</keyword>
    <keyword evidence="16" id="KW-0539">Nucleus</keyword>
    <keyword evidence="9" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="11" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="ENTH" evidence="22">
        <location>
            <begin position="14"/>
            <end position="145"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="21">
        <location>
            <begin position="318"/>
            <end position="348"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004132"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004555"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004600"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00008011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022583"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023176"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00055144"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00061829"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00068054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS50942"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015366.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="614" mass="67039" checksum="B9E7F8069AC338AA" modified="2019-05-08" version="1">MSGQSITDRIAAAQHSVTGSAISKAVCKATTHEVMGPKKKHLDYLIQCTNEMNVNIPQLADTLFERTVNSSWVVVFKALITTHHLMMYGNERFIQYLASRNTLFNLNNYLDKSAMQGYDMSTFIRRYSKYLNEKALSYRLVAVDFTKMKRGIDGVMRTMNTEKLLKTLPIIQNQLDALLDFDANPNELTNGVINAAFMLLFKDSIRLFAAYNEGVINLLEKYFDMKKTQCKEGLDIYKKFLARMTKLSEFLKVAEQVGIDQGDIPDLTQAPSSLLEALEQHLASLEGKKTKEVSAASRASTLSSAVSTLANTGMSFSKMDEREKQQALEEEQARLQALKEQRLKEISVVSNSTSTSASPSTLSGKSVNTTTAVDLFAAPAPTTSSMPNLSSDLFDLQPAFVPTVQSTPAIATSASSAWGGGFPLHSAPQSTTSSIINVDFDAVFGGKSTAHEYRTTSEDVLQPTVPTQCQRTVLINQQSGKILANDLDSSLANLVGNLGFGGTPSKKSDMQWNQPTEKKLTGGTNWQAKTSTSTTWNTAPIPPAPHMNGVHYPGYVPAPMTYPVTTPQVPVYGMVPPQIGAAPLTAPQPMIYTQPGLRPTNPFASVPGPQIQFM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKB7</accession>
    <name>A0A452HKB7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">SLIT-ROBO Rho GTPase-activating protein 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="28 29">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="29" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="28" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="28" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="5">Cell projection</location>
            <location evidence="5">Dendritic spine</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cell projection</location>
            <location evidence="3">Lamellipodium</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cytoplasm</location>
            <location evidence="4">Cytosol</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasmic vesicle</location>
            <location evidence="2">Phagosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="18">Postsynaptic cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="19">Postsynaptic density</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKB7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017546.1">
        <property type="protein sequence ID" value="ENSGAGP00000015363.1"/>
        <property type="gene ID" value="ENSGAGG00000010110.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043197">
        <property type="term" value="C:dendritic spine"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030027">
        <property type="term" value="C:lamellipodium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045335">
        <property type="term" value="C:phagocytic vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0014069">
        <property type="term" value="C:postsynaptic density"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd04383">
        <property type="entry name" value="RhoGAP_srGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd11955">
        <property type="entry name" value="SH3_srGAP1-3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.555.10:FF:000010">
        <property type="entry name" value="SLIT-ROBO Rho GTPase-activating protein 1 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1270.60:FF:000006">
        <property type="entry name" value="SLIT-ROBO Rho GTPase-activating protein 1 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.40:FF:000266">
        <property type="entry name" value="SLIT-ROBO Rho GTPase-activating protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.60">
        <property type="entry name" value="Arfaptin homology (AH) domain/BAR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.555.10">
        <property type="entry name" value="Rho GTPase activation protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027267">
        <property type="entry name" value="AH/BAR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031160">
        <property type="entry name" value="F_BAR_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001060">
        <property type="entry name" value="FCH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000198">
        <property type="entry name" value="RhoGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051627">
        <property type="entry name" value="SLIT-ROBO_RhoGAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035648">
        <property type="entry name" value="srGAP1/2/3_SH3"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14166">
        <property type="entry name" value="SLIT-ROBO RHO GTPASE ACTIVATING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00611">
        <property type="entry name" value="FCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00620">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00018">
        <property type="entry name" value="SH3_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00055">
        <property type="entry name" value="FCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00324">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00326">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103657">
        <property type="entry name" value="BAR/IMD domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51741">
        <property type="entry name" value="F_BAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50238">
        <property type="entry name" value="RHOGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="16" id="KW-0966">Cell projection</keyword>
    <keyword evidence="12 22 23" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="9" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="17" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="7" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="13" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="14" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-0628">Postsynaptic cell membrane</keyword>
    <keyword evidence="29" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6 21" id="KW-0728">SH3 domain</keyword>
    <keyword evidence="11" id="KW-0770">Synapse</keyword>
    <feature type="domain" description="F-BAR" evidence="27">
        <location>
            <begin position="7"/>
            <end position="251"/>
        </location>
    </feature>
    <feature type="domain" description="Rho-GAP" evidence="26">
        <location>
            <begin position="413"/>
            <end position="602"/>
        </location>
    </feature>
    <feature type="domain" description="SH3" evidence="25">
        <location>
            <begin position="651"/>
            <end position="710"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="24">
        <location>
            <begin position="616"/>
            <end position="636"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="24">
        <location>
            <begin position="710"/>
            <end position="735"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="24">
        <location>
            <begin position="760"/>
            <end position="854"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="24">
        <location>
            <begin position="951"/>
            <end position="989"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="23">
        <location>
            <begin position="300"/>
            <end position="334"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="24">
        <location>
            <begin position="714"/>
            <end position="724"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="24">
        <location>
            <begin position="787"/>
            <end position="799"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="24">
        <location>
            <begin position="812"/>
            <end position="823"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="24">
        <location>
            <begin position="837"/>
            <end position="851"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="24">
        <location>
            <begin position="967"/>
            <end position="989"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004262"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004510"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004514"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004552"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023257"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00034100"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00034105"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00073507"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01077"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="26">
        <source>
            <dbReference type="PROSITE" id="PS50238"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="27">
        <source>
            <dbReference type="PROSITE" id="PS51741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="28">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015363.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="29">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="989" mass="111412" checksum="A8D935DF46A00DC4" modified="2019-05-08" version="1">MSSPTRKKLQLTRFIRTQLIEQLKCLDQQCELRVQLLQDLQDFFRKKAEIEMDYSRNLEKLAERFLAKTRSTKDQQFKKDQNVLSPVNCWNLLLNQVKRESRDHTTLSDIYLNNIIPRFVQVSEDSGRLFKKSKEVGLQLQEDLMKVLNELYTVMKTYHMYNADSISAQSKLKEAEKQEEKQIGKSVKFEEKHVRRSSVKKIEKMKEKRQAKYTENKLKAIKARNEYLLALEATNASVFKYYIHDLSDLIDVSVLLLGLMSFPLTPLCASPSLTPFPPQKGHLFDSHLGVSQLCAQQPVQSELVQRCQQLQSRLSTLKIENEEVKKTMEATLQTIQDIVTIEDFDVSDCFQYSNSMESVKSTVSETFMSKPSIAKRRANQQETEQFYFTKMKEYLEGRNLITKLQAKHDLLQKTLGESKSGTRFQAIPLVVESCIRFISRHGLQHEGIFRVSGSQVEVNDIKNAFERGEDPLAGDQNDHDMDSIAGVLKLYFRGLEHPLFPKEIFHDLIACVTMDNLQERALHIRKVLLSLPKTTLIVMRYLFAFLSHLSQFSEENMMDPYNLAICFGPTLMSVPEGHDQVSCQAHVNELIKTIIIQHENIFPGLRELEGPVYSRGGSTEDYCESPHGETASAEDSVQDIAAEHHTSDDECEPIEAIAKFDYIGRTARELSFKKGASLLLYQRASDDWWEGRHNGIDGLIPHQYIVVQDTEDGSSERSSPKSEIEINSETSILRYGPSPTPACTDLFCFCVGRQRKRPESGSIRRAFRQSDSHGLASSLVEPASPGATASSRPSSQPIMSPSLPKEVPDKCSSSGHGSLNSISRHSSLKNRIDSPQIRKTVTAGRSKSFNNHRPMDPEVIVQDIEATMNTALNELRELERQSSVKHTPDVVLDTLEPLKTSPVVAPTSEPSSPLHTQILKDSEPAFQRSASTAGDIPCAFRPVKSVKMAAQVKPPATRPKPAVFPKTNATSPGVASSASQQPTDKSCTV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HKB8</accession>
    <name>A0A452HKB8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">ENTH domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the PICALM/SNAP91 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HKB8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017544.1">
        <property type="protein sequence ID" value="ENSGAGP00000015362.1"/>
        <property type="gene ID" value="ENSGAGG00000011583.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005905">
        <property type="term" value="C:clathrin-coated pit"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030136">
        <property type="term" value="C:clathrin-coated vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098894">
        <property type="term" value="C:extrinsic component of presynaptic endocytic zone membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008021">
        <property type="term" value="C:synaptic vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005545">
        <property type="term" value="F:1-phosphatidylinositol binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032050">
        <property type="term" value="F:clathrin heavy chain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005546">
        <property type="term" value="F:phosphatidylinositol-4,5-bisphosphate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000149">
        <property type="term" value="F:SNARE binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048268">
        <property type="term" value="P:clathrin coat assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072583">
        <property type="term" value="P:clathrin-dependent endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016185">
        <property type="term" value="P:synaptic vesicle budding from presynaptic endocytic zone membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd16985">
        <property type="entry name" value="ANTH_N_AP180"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.90:FF:000001">
        <property type="entry name" value="phosphatidylinositol-binding clathrin assembly protein-like isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.90">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.150">
        <property type="entry name" value="ANTH domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011417">
        <property type="entry name" value="ANTH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014712">
        <property type="entry name" value="ANTH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045192">
        <property type="entry name" value="AP180-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013809">
        <property type="entry name" value="ENTH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008942">
        <property type="entry name" value="ENTH_VHS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22951">
        <property type="entry name" value="CLATHRIN ASSEMBLY PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22951:SF11">
        <property type="entry name" value="ENTH DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07651">
        <property type="entry name" value="ANTH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00273">
        <property type="entry name" value="ENTH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48464">
        <property type="entry name" value="ENTH/VHS domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF89009">
        <property type="entry name" value="GAT-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50942">
        <property type="entry name" value="ENTH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="6" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="ENTH" evidence="7">
        <location>
            <begin position="14"/>
            <end position="145"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="304"/>
            <end position="334"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50942"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015362.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="649" mass="70766" checksum="06D9983C239245B8" modified="2019-05-08" version="1">MSGQSITDRIAAAQHSVTGSAISKAVCKATTHEVMGPKKKHLDYLIQCTNEMNVNIPQLADTLFERTVNSSWVVVFKALITTHHLMMYGNERFIQYLASRNTLFNLNNYLDKSAMQGYDMSTFIRRYSKYLNEKALSYRLVAVDFTKMKRGIDGVMRTMNTEKLLKTLPIIQNQLDALLDFDANPNELTNGVINAAFMLLFKDSIRLFAAYNEGVINLLEKYFDMKKTQCKEGLDIYKKFLARMTKLSEFLKVAEAPSSLLEALEQHLASLEGKKTKEVSAASRASTLSSAVSTLANTGMSFSKMDEREKQQALEEEQARLQALKEQRLKEISVVSNSTSTSASPSTLSGKSVNTTTAVDLFAAPAPTTSSMPNLSSDLFDLQPAFVPTVQSTPAIATSASSAWGGGPFSSSNGCVGSPSHLDIFDMKPVEEAVKSTVPFVSSTFTSKQTVEQFSGFPLHSAPQSTTSSIINVDFDAVFGGKSTAHEYRTTSEDVLQPTVPTQCQRTVLINQQSGKILANDLDSSLANLVGNLGFGGTPSKKSDMQWNQPTEKKLTGGTNWQAKTSTSTTWNTAPIPPAPHMNGVHYPGYVPAPMTYPVTTPQVPVYGMVPPQIGAAPLTAPQPMIYTQPGLRPTNPFASVPGPQIQFM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMZ2</accession>
    <name>A0A452IMZ2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Kinesin-like protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
            <location evidence="2">Microtubule organizing center</location>
            <location evidence="2">Centrosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Spindle</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="13 14">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Kinesin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMZ2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033188.1">
        <property type="protein sequence ID" value="ENSGAGP00000029224.1"/>
        <property type="gene ID" value="ENSGAGG00000021132.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005874">
        <property type="term" value="C:microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005819">
        <property type="term" value="C:spindle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008017">
        <property type="term" value="F:microtubule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003777">
        <property type="term" value="F:microtubule motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007019">
        <property type="term" value="P:microtubule depolymerization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007018">
        <property type="term" value="P:microtubule-based movement"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd01367">
        <property type="entry name" value="KISc_KIF2_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000006">
        <property type="entry name" value="Kinesin-like protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR054473">
        <property type="entry name" value="KIF2A-like_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027640">
        <property type="entry name" value="Kinesin-like_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019821">
        <property type="entry name" value="Kinesin_motor_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001752">
        <property type="entry name" value="Kinesin_motor_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47971:SF24">
        <property type="entry name" value="KINESIN-LIKE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47971">
        <property type="entry name" value="KINESIN-RELATED PROTEIN 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22923">
        <property type="entry name" value="KIF2A-like_1st"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00225">
        <property type="entry name" value="Kinesin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00380">
        <property type="entry name" value="KINESINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00129">
        <property type="entry name" value="KISc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00411">
        <property type="entry name" value="KINESIN_MOTOR_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50067">
        <property type="entry name" value="KINESIN_MOTOR_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 13 14" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="12" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="4" id="KW-0132">Cell division</keyword>
    <keyword evidence="9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="5 14" id="KW-0493">Microtubule</keyword>
    <keyword evidence="7" id="KW-0498">Mitosis</keyword>
    <keyword evidence="10 13 14" id="KW-0505">Motor protein</keyword>
    <keyword evidence="6 13 14" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Kinesin motor" evidence="16">
        <location>
            <begin position="223"/>
            <end position="553"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="64"/>
            <end position="141"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="165"/>
            <end position="186"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="15">
        <location>
            <begin position="123"/>
            <end position="140"/>
        </location>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <begin position="313"/>
            <end position="320"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004186"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022776"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00283"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="RuleBase" id="RU000394"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50067"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029224.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="687" mass="77666" checksum="13F20584A2A97322" modified="2019-05-08" version="1">MAAANFGKIQIGIYVEIKRSDGRIHQAMVTSLNEDNESVTVEWIENGDTKGKEIDLESIFSLNPDLAPDEDIEPSPETPPPPTPSAKVNKIVKNRRTVAPIKNETPARDNRAVGSARARPTQLPDQSSSTQQNGSVSDISPVQAAKKEFGPPSRRKSNCVKEVEKLQEKREKRRLQQQELREKRAQDVDATNPNYEIMCMIRDFRGSLDYRPLTTTDPIDEHRICVCVRKRPLNKKETVMKDLDVITIPSKDVVMVHEPKQKVDLTRYLENQTFRFDYAFDDTAPNEMVYRFTARPLVETIFERGMATCFAYGQTGSGKTHTMGGDFSGKNQDCSKGIYALAARDVFLMLKKPNYKKLELQVYATFFEIYSGKVFDLLNRKTKLRVLEDGKQQVQVVGLQEREVKCVEDVLKLIEIGNSCRTSGQTSANAHSSRSHAVFQIILRRKGKLHGKFSLIDLAGNERGADTSSADRQTRLEGAEINKSLLALKECIRALGRNKPHTPFRASKLTQVLRDSFIGENSRTCMIATISPGMASCENTLNTLRYANRVKELTVDPSAAGDIRPIIHHAPNQIDDLETQWGVGSSPQRDDLKLLCEQNEEEVSPQLFTFHEAVSQMVEMEEQVVEDHRAVFQESIRWLEDEKALLEMTEEVDYDVDSYATQLEAILEQKIDILTELRGKAVFFVHL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMZ3</accession>
    <name>A0A452IMZ3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Ubiquitin-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the DDI1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMZ3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033200.1">
        <property type="protein sequence ID" value="ENSGAGP00000029234.1"/>
        <property type="gene ID" value="ENSGAGG00000021142.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004190">
        <property type="term" value="F:aspartic-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072711">
        <property type="term" value="P:cellular response to hydroxyurea"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051603">
        <property type="term" value="P:proteolysis involved in protein catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097752">
        <property type="term" value="P:regulation of DNA stability"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031647">
        <property type="term" value="P:regulation of protein stability"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd05479">
        <property type="entry name" value="RP_DDI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd01796">
        <property type="entry name" value="Ubl_Ddi1_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.70.10:FF:000005">
        <property type="entry name" value="DNA damage inducible 1 homolog 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.20.90:FF:000107">
        <property type="entry name" value="protein DDI1 homolog 2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.70.10">
        <property type="entry name" value="Acid Proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.20.90">
        <property type="entry name" value="Phosphatidylinositol 3-kinase Catalytic Subunit, Chain A, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057273">
        <property type="entry name" value="Ddi1/2_HDD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033882">
        <property type="entry name" value="DDI1_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019103">
        <property type="entry name" value="Peptidase_aspartic_DDI1-type"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021109">
        <property type="entry name" value="Peptidase_aspartic_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000626">
        <property type="entry name" value="Ubiquitin-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029071">
        <property type="entry name" value="Ubiquitin-like_domsf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15397:SF3">
        <property type="entry name" value="DNA DAMAGE INDUCIBLE 1 HOMOLOG 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15397">
        <property type="entry name" value="SODIUM-GLUCOSE COTRANSPORTER REGULATORY PROTEIN -RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09668">
        <property type="entry name" value="Asp_protease"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24669">
        <property type="entry name" value="Ddi2_HDD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00240">
        <property type="entry name" value="ubiquitin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50630">
        <property type="entry name" value="Acid proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54236">
        <property type="entry name" value="Ubiquitin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50053">
        <property type="entry name" value="UBIQUITIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0064">Aspartyl protease</keyword>
    <keyword evidence="4" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="2" id="KW-0645">Protease</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Ubiquitin-like" evidence="6">
        <location>
            <begin position="1"/>
            <end position="81"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="102"/>
            <end position="132"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00009136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022750"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50053"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029234.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="397" mass="44460" checksum="D27B15AC22C0C366" modified="2019-05-08" version="1">MLLTVFCLRRDRSELTFSLQVDADFELQNFRALCELESGIPAAESQIVYAERPLTDNHRSLASYGLKDGDVVILRQKENAEPRPPVQFPGLPRIDFSSIAVPGTSAQQQQQPAQHLRPSPPDTSSSPQGLDNPALLRDMLLANPHELSLLKERNPPLAEALLSGDLEKFTRVLVEQQQDRARREQERIRLFSADPFDLEAQAKIEEDIRQQNIEENMTIAMEEAPESFGQVVMLYINCKVNGHPVKAFVDSGAQMTIMSQACAERCNIMRLVDRRWAGIAKGVGTQKIIGRVHLAQVQIEGDFLPCSFSILEEQPMDMLLGLDMLKRHQCSIDLKKNVLVIGTTGSQTTFLPEGELPECARLAYGPGREDMRPEEIADQELAEALQKSVEEAEHQKP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMZ5</accession>
    <name>A0A452IMZ5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Serologically defined colon cancer antigen 8</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IMZ5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033439.1">
        <property type="protein sequence ID" value="ENSGAGP00000029449.1"/>
        <property type="gene ID" value="ENSGAGG00000021276.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="term" value="C:centriole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007098">
        <property type="term" value="P:centrosome cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030010">
        <property type="term" value="P:establishment of cell polarity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001764">
        <property type="term" value="P:neuron migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="P:tube formation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031887">
        <property type="entry name" value="SDCCAG8"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR34343">
        <property type="entry name" value="SEROLOGICALLY DEFINED COLON CANCER ANTIGEN 8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR34343:SF1">
        <property type="entry name" value="SEROLOGICALLY DEFINED COLON CANCER ANTIGEN 8"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="CCCAP"/>
        <property type="match status" value="1"/>
    </dbReference>
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    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
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        <location>
            <begin position="34"/>
            <end position="60"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="81"/>
            <end position="106"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="125"/>
            <end position="166"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="229"/>
            <end position="270"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="305"/>
            <end position="591"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029449.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="5">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="666" mass="77115" checksum="CBA2D617439312DD" modified="2019-05-08" version="1">MREAREESDLERYQRSLRDRASKSIHQLKYALVEGDPTAGESEVAFDQSSTGNDESGETAWQELQHSHAVNQLKALLREQEGLESEVSPSRRRKMSPSRTSEAADQSLPTLCDLVPIINDQSQYIHHLEAEVKFCKEELSGMKDRIQVVVLENEKLNEELKSLTVEQTLREQTLLDASANVRNSWTVGGDDSYLCQPAKSSPSHNKEQVLATRDTAETEKWQLELEKLKRLYQERTDTLEAQVRSLRKDLSESEKNCEDLKGRLKHQESLIATSNYNRVGGLCLKCAQHEAVLSQTHSNVHMHTIERLTKERDDLMGALVSVRRDLSEMQQRESSAYEQVKQAVQMTEEANLEKMKALVQCEQLKSEMERQKDRLERELIAQLDKRAGEKEALRGEMKKEREDLASTVMVLSQNVATLEARLERIIREKNSIANQLEEAQNQLTSQEMEVNKVCGEMRFQLNQTKMKKDEAEKEHREYRTKTIRELEIKDQEIEKLGLELNENKLRLEQAQQDAARAKDECLKLTELLGKSEHQLHLTRLKKESIQHSFSNEAKTRALQAEQREQELTQKMQQMEAQHDKTVNELDSLLTSQNTFIAKLKEECCMLARKLEHVTEKNRLHQLDKHCQTTAQQLVELLNKQNQLFRERQMLTEEVQFLRTQLPCVPV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMZ6</accession>
    <name>A0A452IMZ6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">AKNA domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IMZ6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033305.1">
        <property type="protein sequence ID" value="ENSGAGP00000029326.1"/>
        <property type="gene ID" value="ENSGAGG00000021212.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052655">
        <property type="entry name" value="AKNA_Centrosome-Trans_reg"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022150">
        <property type="entry name" value="AKNA_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21510">
        <property type="entry name" value="AKNA DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21510:SF16">
        <property type="entry name" value="PROTEIN AKNAD1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12443">
        <property type="entry name" value="AKNA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="AKNA" evidence="3">
        <location>
            <begin position="440"/>
            <end position="527"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="320"/>
            <end position="340"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="629"/>
            <end position="656"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="489"/>
            <end position="551"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="645"/>
            <end position="656"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="Pfam" id="PF12443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029326.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="988" mass="112687" checksum="1E95E8641608C981" modified="2019-05-08" version="1">MEYNKDIQINTSKSRQPDHLKKQVNNILLDNFDYPEDATDEEQEDLPYDGDLERTYHHNNESNNLKACVSVENISNNFLNLTCSEINRSLKEEINYETQQLSQEKVFSHRMPATRTNGIMTEMAIEAPVSGMPLETELSVGAFSSLDRKEHFTNSNISDVLLRHFSKEELSNTGQLIDCETIPETSFTESVDETVLTKTRISECTKPTLRTEQWAKHFEDYHLNRQEEKSEDDYEDKHLIDENKFVIDDRATSYGDEKNCIQENSQLIAENEDTNNFKNIRNNHSHQKGLFERTGSSHELKYGQGQVNYCLPDFSKVPPKVKIPKRNNSDKSAPTMKRTKFSPNLVGKSAIVNDVLETINYFDSVEVKNQEEEMRIPELLQQLELLTKHAEAQNRIDHLRFDPKIYTHSASPNPNLSIHSRCMGAASEESVFHPLTVPIQPLLGLSQASFSGLQSGKIISSLPSASSAGEVHCLNPNLMQNTTKGEKMSQMLKEQAEQLKAKVEEFSKCVIQEAFPLQDRCLVLKELKGYLDTLERKYLATKEEHRGLQLQNYNHKSISVGEFDPDRKVEGEIFRLGMLLEDVKEKIDDSICSPCPSSFTTSPMSPSLTQSESVCSSCSCLRESPVVSSISDPPERSATEVNFHNNERNKGENRSHAIDVIPKKTSQLSLQNDNCDPFPEIQLGLQKRESCAYVKGMEPLEKAGLLANKHSSDVMQCFLPHEADNGPGGLESHRQLILSQKRNIDQENPKGSLNFWQMKSPPKTKPYNMCNLEHKIINLNVRQEQGSDSEDISFCDSFLESHSDEFLEHRTKNYKSRRAELKEQSKALQYRDWNINFCTSGQRNKCSRESDKNNIASSQKKRIVANSVCTSRQPDQFVYRISDKDNLETPQTCYSGMDDTIILSPQHLASRKDYGNKSIINIRNRHTSYTDTKILNSTLDHAIQTATSLQETTERMVQVVAEDLAKVKTHRRRRLKLNILSRNTIKLR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INA0</accession>
    <name>A0A452INA0_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="9">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000033319.1">
        <property type="protein sequence ID" value="ENSGAGP00000029339.1"/>
        <property type="gene ID" value="ENSGAGG00000020783.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005938">
        <property type="term" value="C:cell cortex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0120025">
        <property type="term" value="C:plasma membrane bounded cell projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903479">
        <property type="term" value="P:mitotic actomyosin contractile ring assembly actin filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd21276">
        <property type="entry name" value="CH_IQGAP3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.418.10:FF:000013">
        <property type="entry name" value="IQ motif containing GTPase activating protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.506.10:FF:000004">
        <property type="entry name" value="IQ motif containing GTPase activating protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.418.10">
        <property type="entry name" value="Calponin-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.506.10">
        <property type="entry name" value="GTPase Activation - p120gap, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001715">
        <property type="entry name" value="CH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036872">
        <property type="entry name" value="CH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000048">
        <property type="entry name" value="IQ_motif_EF-hand-BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000593">
        <property type="entry name" value="RasGAP_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023152">
        <property type="entry name" value="RasGAP_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001936">
        <property type="entry name" value="RasGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001202">
        <property type="entry name" value="WW_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149">
        <property type="entry name" value="RAS GTPASE-ACTIVATING PROTEIN WITH IQ MOTIF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149:SF10">
        <property type="entry name" value="RAS GTPASE-ACTIVATING-LIKE PROTEIN IQGAP3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00307">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00612">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00616">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03836">
        <property type="entry name" value="RasGAP_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00033">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00015">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00323">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47576">
        <property type="entry name" value="Calponin-homology domain, CH-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF143885">
        <property type="entry name" value="RGC domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50021">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00509">
        <property type="entry name" value="RAS_GTPASE_ACTIV_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50018">
        <property type="entry name" value="RAS_GTPASE_ACTIV_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01159">
        <property type="entry name" value="WW_DOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50020">
        <property type="entry name" value="WW_DOMAIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="4" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="1" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="Calponin-homology (CH)" evidence="8">
        <location>
            <begin position="35"/>
            <end position="150"/>
        </location>
    </feature>
    <feature type="domain" description="WW" evidence="7">
        <location>
            <begin position="668"/>
            <end position="701"/>
        </location>
    </feature>
    <feature type="domain" description="Ras-GAP" evidence="6">
        <location>
            <begin position="964"/>
            <end position="1197"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="1356"/>
            <end position="1379"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="4">
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            <begin position="159"/>
            <end position="186"/>
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            <begin position="1357"/>
            <end position="1366"/>
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            <dbReference type="ARBA" id="ARBA00022553"/>
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            <dbReference type="ARBA" id="ARBA00022737"/>
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    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
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    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
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    <evidence type="ECO:0000259" key="6">
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            <dbReference type="PROSITE" id="PS50018"/>
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            <dbReference type="PROSITE" id="PS50020"/>
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            <dbReference type="PROSITE" id="PS50021"/>
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    <evidence type="ECO:0000313" key="9">
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            <dbReference type="Ensembl" id="ENSGAGP00000029339.1"/>
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    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1576" mass="179897" checksum="E9A48CB4C46CE08D" modified="2019-05-08" version="1">MEPAAESGPARTGYERLTADEMDEQRRQNVAYQYLCRLEEAKRWMEACLNEELASPTELEESLRNGVILAKLGHCFAPDIIPLKKIYDREQMRYEATGLHFRHTDNINHWREAMAHLGLPTIFHPETTDIYDKKNMPRVVYCIHALSLYLFKLGLAPQIQDLYGKVNFTEEEINNMKRELEKYGLQMPAFSKIGGILASELSVDEAAVHAAVLAINEAVERGVVEQTMVALHNPSAMLLNLRENLAAVYQEMLFQVKMEKAGNARNRVRADEDIYDRCLTQAEIQGNINKVNVHGALEYVDDALERQDPLALCQALQDPVLALRGVQRENAAWYLDQLSTDREQKALELGYVDLLEQEEVQAGVYAANERGDQEHAMRRAVSRINASIRRGEAMETVRELMSPEAQLPKAFPFAAQLYQHELSLLQQHHPQGELGQEELYVAVEMLSAVALIDRALEAGDVSGFWSSLVSPALGLSDIEDANAQRYFDDLLKLKRRSGKAGVAFLSWNDIQACVNSANLAVQDENDKILAIRLINEALAQDDPEKTLTALLLPSARFSAVTLPGARRYHDVLTAALKQKAQVTGDDGAALWWEEIQQGVSKANQDTEAARRLALGIAAINQAIKEGKPAQTARVLRNPDVSLRGIVAECAGAYQAQLAALLAAKRQAGNANHNWVRHQLKDGSVYYFSLQNFEGSWERPHDCVLNSTHLSREEIQSTITGVTAAHDRERLWESNAGFVVRLQARMRGYLVRQEFVARRHLLQEQLPAVIKIQVRSAARLPSRSHADGHDDIYIIQTYVRMWRARKRYRERLRYFRKNVSVHARDPPLSIVRRFVHLLEQSQHDFWEESELLKLREEVVKRIRSNQQLESDLNLMDIKIGLLVKNRITLQEVVSHCKKLTRKNKEQLSNLMAIDKQKGLKSLSKEKRQKLEAYQHLFYLLQTHPVYLAKLIFQMPQNKSTKFMESVVFTLYNYASNPREAYLLLQLFKTALQEEIRSKVDQLRDILTGNPTVIRLVVSFYRNARGQNALRQILGSPVQEVLQDKTLSIRTNPVDIYKAWINQLESQSGQKSKFPYDVSPEQALSHPEVQRRLDISIRNLLAVTDKFLSAITSSVDKIPYGMRYVAKVLKTSLAEKFPAAGEEDICKIVGNLLYYRFMNPAVVAPDGFDIVDISAGVALHPEHRRNLGSIAKILQHAAANKMFEGENGHLRVVNQYLEETHSKFRKFICAACSVPEPEERFNVDEYSEMVAVAKPVIYITVAELINTHKLLLEHQDSISPDHKDPLHELLEDLDELPTVQSLIGTCRLLVSFKPAPPLVSSFNTKQMLVDVIQSQPGDSLLEILRTEAFHHHQMHRRALREAQTPDRLKRNRSLAGNSQLSMEEKKRKVIRNLRRLENLGVVDSANQYQEIINEMAKDIRNQRRYRQHRKGELVKLRQTLCGLNSKTSFYEEQIDYYNQYIRTCLDNLAASSKYVGKKQQSLHYTAARLFEKGVLLEIEDLPVSQFRNVIFDIIPCGEAGKFQVKAKFMGIDMEKFQLHYQDLLQLQYEGVAVMKLFDKAKVNVNLLIFLLNKKFFRK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INA1</accession>
    <name>A0A452INA1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Centrosomal protein of 170 kDa</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Plays a role in microtubule organization. Required for centriole subdistal appendage assembly.</text>
    </comment>
    <comment type="subunit">
        <text evidence="10">Interacts with CCDC68 and CCDC120; leading to recruitment to centrosomes. Interacts with PLK1. Interacts with NIN. Interacts with FHDC1. Interacts with CCDC61. Interacts with TBK1; efficient complex formation may be dependent on the presence of CCDC61.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Microtubule organizing center</location>
            <location evidence="1">Centrosome</location>
            <location evidence="1">Centriole</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
            <location evidence="2">Spindle</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the CEP170 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000033422.1">
        <property type="protein sequence ID" value="ENSGAGP00000029433.1"/>
        <property type="gene ID" value="ENSGAGG00000021269.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005814">
        <property type="term" value="C:centriole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005874">
        <property type="term" value="C:microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005819">
        <property type="term" value="C:spindle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="CDD" id="cd22724">
        <property type="entry name" value="FHA_Cep170A"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Centrosomal protein of 170 kDa"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.200.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051176">
        <property type="entry name" value="Cent_Immune-Sig_Mod"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029300">
        <property type="entry name" value="CEP170_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000253">
        <property type="entry name" value="FHA_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008984">
        <property type="entry name" value="SMAD_FHA_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15715">
        <property type="entry name" value="CENTROSOMAL PROTEIN OF 170 KDA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15715:SF17">
        <property type="entry name" value="CENTROSOMAL PROTEIN OF 170 KDA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15308">
        <property type="entry name" value="CEP170_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00498">
        <property type="entry name" value="FHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00240">
        <property type="entry name" value="FHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49879">
        <property type="entry name" value="SMAD/FHA domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50006">
        <property type="entry name" value="FHA_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="6" id="KW-0493">Microtubule</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
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            <begin position="23"/>
            <end position="73"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
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            <end position="149"/>
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    </feature>
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    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1181"/>
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        </location>
    </feature>
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        <location>
            <begin position="1349"/>
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        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1404"/>
            <end position="1435"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1546"/>
            <end position="1573"/>
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        <location>
            <begin position="1597"/>
            <end position="1616"/>
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            <begin position="116"/>
            <end position="125"/>
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    </feature>
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            <end position="533"/>
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            <end position="1111"/>
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            <begin position="1606"/>
            <end position="1616"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00004186"/>
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            <dbReference type="ARBA" id="ARBA00010436"/>
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            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
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    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
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    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00053332"/>
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    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00065884"/>
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    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00070079"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
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    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50006"/>
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    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029433.1"/>
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    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1616" mass="178608" checksum="4E68E0976CD770ED" modified="2019-05-08" version="1">MSLTSWFLVSSGGTRHRLPREMIFVGRDDCELMLQSRSVDKQHAVVNYDASTDEHLVKDLGSLNGTFVNDVRIPEQTYITLKLEDKLRFGYDTNLFTVVRGEMRVPEEALKHEKFTSQLQLSQKSSEAEGSKPVSSTGTEPNVPDATGEVHHKATEALKSEEKSMDLSAMPRGTPLYGQPSWWGEDEADEKNGCKRDIKHEEKIHETGASGCSTDAKQVVEKSSAANEEELYPFCREPSYFEIPTKEFHQPSQIAESTIHEIPTKDTQTSHAAGAGHASFTIEFDDTTPGKVTIKDHVTKFTSDQRHKSKKSPLGGQDLPGLQTVMMAAESKVADWLAQNNPPRMIWEATEEDSKSIKSDVPVYLKRLKGNKHDDGTQSDSENAGARRHYNKRATVDEHLSHQHLEQKKSYQKAHTTEKHEEQAAISQTAFMIEFFDEDHPRKRRSYSFSQNVGALCSETPSSTPHVRAEKVKGTSGDIKGVSLPLQTSRVAHGVHAKLLKQKSEDPPVPLPFLQTALLRSSGSLGHRPSQNQEMDKKLKSQLVSAATEKDNDDDQSDKGTYTIELENPNAEEMEARKMIDKEGSSASGSKRWVSQWASLAANHTRHDQDDMRLESSAPAPRENDTDISESGVSVRSTSSATSVTSQGERKRRTLPQLPKEEKATESSRAKVTPHQRSEIGEKQDTELQEKETPTHASQKEMPQDAGSRSMAKANRTMNGLSSKPGGDNKVSSHPISCSSFSKEKSEIVRETSVVKQALAKIQQQEQKEQAHRTSTKVSSAKLLANQIEKGREETSTPHKMLDNHQEKAPGHSTSKAEMKIVQSEGKRKKTEEIIKGQTPKGFGGEKKESSKPLVRQGSFTIDKPSTNIPIELIPHINKQTGSTPPSFSLTSTNRIRERSDSVDTDSSLDTTLILKDTEAVMAFLEAKLREENKTDEGPETPSYNRDNSISPESDVDTASTISLVTGDTERKTTQKRKSFTTLYKDRCSTSSPSKDVLKSSATNAREKIDKKTKSRSSDAGSRAEGRKLVQPSGRMRQPSVDLTDDDQTSSVPHSAISDILSSDQETYSGKSHGRGHFTSADELLHSKMEGSKSAKSKISPVTTGQSSKSITLPRPRPTRTSLLRRARLGEASDSELADADKASVASEVSTTSSTSKPPSGRRNISRIDLLAQPRRIRLGSLSARSDSEATITRSAASSRTAEAIIRSSGRLTPSADSSKVSARTRANSISRLSDSKSKSLTSAHNSPSVNSRWRRFPTDYASTSEDEFGSNRNSPKHTRLRTSPALKTPRLQSSGTAMQTSNTFKHRIKEQEDYIRDWTAHREEIARISQDLALIAREINDVAGEIDSVTSSGTAPSTTVSTAATTPGSAIDTREEVGDLHGEMHKLVDRVFDESLNFRKIPPLVHAKPPEGNSRSNDSRPQLTDALDPPTITRRRTWSRDEVMGDSLLLSSVFQFSRKIRQSIDKTAGKIRILFKDKDRNWEEIESKLRTESEVPIVKTSSMEISSILQELKRVEKQLQVINAMIDPDGTLDALSNLGFTSPVLPAQPKHKSSPVSHSTRAQIQPNCQPEARALRPAVTSVSTAFDNVESKPDFSIHFNRFNPDGEEEDPSMRE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INA3</accession>
    <name>A0A452INA3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">SMB domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452INA3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033328.1">
        <property type="protein sequence ID" value="ENSGAGP00000029348.1"/>
        <property type="gene ID" value="ENSGAGG00000020186.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009986">
        <property type="term" value="C:cell surface"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004551">
        <property type="term" value="F:dinucleotide phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003676">
        <property type="term" value="F:nucleic acid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004528">
        <property type="term" value="F:phosphodiesterase I activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046034">
        <property type="term" value="P:ATP metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030505">
        <property type="term" value="P:inorganic diphosphate transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045599">
        <property type="term" value="P:negative regulation of fat cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009143">
        <property type="term" value="P:nucleoside triphosphate catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030500">
        <property type="term" value="P:regulation of bone mineralization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd16018">
        <property type="entry name" value="Enpp"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00091">
        <property type="entry name" value="NUC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.410.20">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.720.10">
        <property type="entry name" value="Alkaline Phosphatase, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.570.10">
        <property type="entry name" value="Extracellular Endonuclease, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017850">
        <property type="entry name" value="Alkaline_phosphatase_core_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044929">
        <property type="entry name" value="DNA/RNA_non-sp_Endonuclease_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001604">
        <property type="entry name" value="Endo_G_ENPP1-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020821">
        <property type="entry name" value="ENPP1-3/EXOG-like_nuc-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044925">
        <property type="entry name" value="His-Me_finger_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002591">
        <property type="entry name" value="Phosphodiest/P_Trfase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036024">
        <property type="entry name" value="Somatomedin_B-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001212">
        <property type="entry name" value="Somatomedin_B_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10151">
        <property type="entry name" value="ECTONUCLEOTIDE PYROPHOSPHATASE/PHOSPHODIESTERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10151:SF77">
        <property type="entry name" value="ECTONUCLEOTIDE PYROPHOSPHATASE_PHOSPHODIESTERASE FAMILY MEMBER 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01223">
        <property type="entry name" value="Endonuclease_NS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01663">
        <property type="entry name" value="Phosphodiest"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01033">
        <property type="entry name" value="Somatomedin_B"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00892">
        <property type="entry name" value="Endonuclease_NS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00477">
        <property type="entry name" value="NUC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00201">
        <property type="entry name" value="SO"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53649">
        <property type="entry name" value="Alkaline phosphatase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54060">
        <property type="entry name" value="His-Me finger endonucleases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90188">
        <property type="entry name" value="Somatomedin B domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51257">
        <property type="entry name" value="PROKAR_LIPOPROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00524">
        <property type="entry name" value="SMB_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50958">
        <property type="entry name" value="SMB_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="4" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="7" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="12"/>
            <end position="40"/>
        </location>
    </feature>
    <feature type="domain" description="SMB" evidence="8">
        <location>
            <begin position="47"/>
            <end position="87"/>
        </location>
    </feature>
    <feature type="domain" description="SMB" evidence="8">
        <location>
            <begin position="88"/>
            <end position="132"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50958"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029348.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="847" mass="97176" checksum="881C39E9068F4B74" modified="2025-10-08" version="2">MKISSRAAGCLPLTYSVFSLSIPVLSACVLTIILGCIFGLKPSCSRDVKTCKGRCFERTFGSCRCDRDCVELGNCCLDFQETCIQPAHIWTCNKFRCGEKRWPENRCSCSDDCVENKDCCVNYYIVCQGAKSWVEEKCEDINEPQCPPGFAKPPVLLFSLDGFRAEYLHTWSGLLPVISKLQKCGTYTPNMRPVYPTKTFPNHYTIVTGLYPESHGIIDNKIYDPHRNVSFTLKNAEKFNPQWYQGQPIWLTAMHQGLKAGTFFWPGSDVRINGTFPDLYRKYNGSIPFEERVIAILHWLQLPKDERPHFYTLYLEEPDSSGHNFGPVSSGVILALQRVDNIVGMLMDGLKQMNLHKCLNIILLSDHGMETASCSKTAYLSTYMENVQDITVIPGPAARLRPQNVPDEYFSYPDQHFKAFMKELLPKRFHYVNNDRIERVHFYLDPQWQLARKPSEIKYCNGGFHGSDNRFMSMQAIFIGFGPGFKFKTQVDPFENIEIYNLICDLLGITPAYNNGTHGSLNHLLKNPVYIPRHPEEESHPSVYPLMKTTSSLDIGCSCDLAGLPIRDFQQQLNHTASEVKKIEKYSLPYGRPRVLQKRSTYYLLYHHQYVSGYSQDLNMTLWSAYTDNRTASVENLSNCLHVDIRIPSSSSKTCSFYKNHHQLSYGFLFPPNTKKSSYEVLLNSNIVPMYHEFKVLWNYFYDVLLPKYTLARNGVNVVSGPVFDYDSNGLYDTPEKVKRLTHDPEVLVPTHYFIVLTSCKNISQTPLQCEGALDALSFIVPHRPDNSESCARHKRDSLWVEERMRFHTARVRDVELLTGLSFYQDRKQPVSDILQLKTYLPTFPME</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INA5</accession>
    <name>A0A452INA5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">glutamate carboxypeptidase II</fullName>
            <ecNumber evidence="18">3.4.17.21</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>Release of an unsubstituted, C-terminal glutamyl residue, typically from Ac-Asp-Glu or folylpoly-gamma-glutamates.</text>
            <dbReference type="EC" id="3.4.17.21"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Membrane</location>
            <topology evidence="2">Single-pass type II membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the peptidase M28 family. M28B subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.4.17.21" evidence="18"/>
    <dbReference type="AlphaFoldDB" id="A0A452INA5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033428.1">
        <property type="protein sequence ID" value="ENSGAGP00000029438.1"/>
        <property type="gene ID" value="ENSGAGG00000020963.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016805">
        <property type="term" value="F:dipeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004181">
        <property type="term" value="F:metallocarboxypeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd08022">
        <property type="entry name" value="M28_PSMA_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.930.40:FF:000001">
        <property type="entry name" value="N-acetylated-alpha-linked acidic dipeptidase 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.630.10:FF:000009">
        <property type="entry name" value="N-acetylated-alpha-linked acidic dipeptidase 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.50.30.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.930.40">
        <property type="entry name" value="Transferrin receptor-like, dimerisation domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.630.10">
        <property type="entry name" value="Zn peptidases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046450">
        <property type="entry name" value="PA_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007484">
        <property type="entry name" value="Peptidase_M28"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039373">
        <property type="entry name" value="Peptidase_M28B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007365">
        <property type="entry name" value="TFR-like_dimer_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036757">
        <property type="entry name" value="TFR-like_dimer_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10404:SF36">
        <property type="entry name" value="GLUTAMATE CARBOXYPEPTIDASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10404">
        <property type="entry name" value="N-ACETYLATED-ALPHA-LINKED ACIDIC DIPEPTIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04389">
        <property type="entry name" value="Peptidase_M28"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04253">
        <property type="entry name" value="TFR_dimer"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52025">
        <property type="entry name" value="PA domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47672">
        <property type="entry name" value="Transferrin receptor-like dimerisation domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53187">
        <property type="entry name" value="Zn-dependent exopeptidases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0121">Carboxypeptidase</keyword>
    <keyword evidence="12" id="KW-0224">Dipeptidase</keyword>
    <keyword evidence="15" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="14 19" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13" id="KW-0482">Metalloprotease</keyword>
    <keyword evidence="16" id="KW-0511">Multifunctional enzyme</keyword>
    <keyword evidence="5" id="KW-0645">Protease</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0735">Signal-anchor</keyword>
    <keyword evidence="6 19" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11 19" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="9" id="KW-0862">Zinc</keyword>
    <feature type="transmembrane region" description="Helical" evidence="19">
        <location>
            <begin position="21"/>
            <end position="42"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase M28" evidence="21">
        <location>
            <begin position="330"/>
            <end position="532"/>
        </location>
    </feature>
    <feature type="domain" description="Transferrin receptor-like dimerisation" evidence="20">
        <location>
            <begin position="600"/>
            <end position="720"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004606"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005634"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022645"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022997"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023049"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023268"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00052003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00066561"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="Pfam" id="PF04253"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="Pfam" id="PF04389"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029438.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="723" mass="81793" checksum="169DA229C6F10C0D" modified="2019-05-08" version="1">MWTALRSGAEQSSPWGSRGRVCALSLAVAAALLLLGFLIGWFTKPTDEVTATSFHEKVKKAFMAEMKAENIKQFLYNFTQRPHLAGTAENLRLAQQIQAQWKEFGLDSVELAHYDVLLSYPNETNPNYISIIDEHGTEIFNTSLFEPPPPGYETVTDVVPPYSAFSAPGVPEVKSNRLLKSRKLKVSCSQLGLIFRGSWKIRPHKLYSPGNFQLMRFCRTAAGHLEHGVWEHSWIMTSSKINTSLIPPEYTYRYEEARAVGLPKLPVHPIGYNDAEKLLRDMGGPAPPDSSWKGSLNVSYNIGPGFMSRYSTRKVKMHIHSSNKVTRIYNVIGTIRGATEPDRYVILGGHRDSWVFGGIDPQSGAAVVHEIVRSYGKLKKEGWRPRRTIIFASWDAEEFGLLGSTEWAEENTKLLQARGVAYINADSSVEGNYTLRIDCTPLMYSLVYSLTREIPSPDEGFKGKSLYESWHENNPSTEYKDVPRINKLGSGNDFEVFFQRLGIASGRARYSKNWRVEKYSSYPVYHSVYETYEIVERFYDPTFKNHLTVAQVRGGLVFELADSVVLPFDCRDYAEALRNYVQVIYNLSKNHQMGLTTYSVSFDSLFSAANNFTEAATDFHRRLQQVDINNPIAVRALNDQLMFLERAFIDPLGLPGRPFYRHIIFAPSSHNKYAGESFPGIYDAMFDIENKADQRKAWEEVKRQISIAAFTVQAAAGTLKDLA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INA8</accession>
    <name>A0A452INA8_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="8">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000033333.1">
        <property type="protein sequence ID" value="ENSGAGP00000029353.1"/>
        <property type="gene ID" value="ENSGAGG00000020783.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005938">
        <property type="term" value="C:cell cortex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0120025">
        <property type="term" value="C:plasma membrane bounded cell projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903479">
        <property type="term" value="P:mitotic actomyosin contractile ring assembly actin filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd21276">
        <property type="entry name" value="CH_IQGAP3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.418.10:FF:000013">
        <property type="entry name" value="IQ motif containing GTPase activating protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.506.10:FF:000004">
        <property type="entry name" value="IQ motif containing GTPase activating protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.190">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.418.10">
        <property type="entry name" value="Calponin-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.506.10">
        <property type="entry name" value="GTPase Activation - p120gap, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001715">
        <property type="entry name" value="CH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036872">
        <property type="entry name" value="CH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000048">
        <property type="entry name" value="IQ_motif_EF-hand-BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000593">
        <property type="entry name" value="RasGAP_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023152">
        <property type="entry name" value="RasGAP_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001936">
        <property type="entry name" value="RasGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001202">
        <property type="entry name" value="WW_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149">
        <property type="entry name" value="RAS GTPASE-ACTIVATING PROTEIN WITH IQ MOTIF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149:SF10">
        <property type="entry name" value="RAS GTPASE-ACTIVATING-LIKE PROTEIN IQGAP3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00307">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00612">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00616">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03836">
        <property type="entry name" value="RasGAP_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00033">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00015">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00323">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47576">
        <property type="entry name" value="Calponin-homology domain, CH-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF143885">
        <property type="entry name" value="RGC domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50021">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00509">
        <property type="entry name" value="RAS_GTPASE_ACTIV_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50018">
        <property type="entry name" value="RAS_GTPASE_ACTIV_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01159">
        <property type="entry name" value="WW_DOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50020">
        <property type="entry name" value="WW_DOMAIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="4" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="1" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="Calponin-homology (CH)" evidence="7">
        <location>
            <begin position="42"/>
            <end position="157"/>
        </location>
    </feature>
    <feature type="domain" description="WW" evidence="6">
        <location>
            <begin position="655"/>
            <end position="688"/>
        </location>
    </feature>
    <feature type="domain" description="Ras-GAP" evidence="5">
        <location>
            <begin position="954"/>
            <end position="1187"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="4">
        <location>
            <begin position="166"/>
            <end position="193"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50020"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029353.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1578" mass="179108" checksum="CA8AE304A0E58274" modified="2019-05-08" version="1">SSATDPLCDLGQVTLAALSLHERLTADEMDEQRRQNVAYQYLCRLEEAKRWMEACLNEELASPTELEESLRNGVILAKLGHCFAPDIIPLKKIYDREQMRYEATGLHFRHTDNINHWREAMAHLGLPTIFHPETTDIYDKKNMPRVVYCIHALSLYLFKLGLAPQIQDLYGKVNFTEEEINNMKRELEKYGLQMPAFSKIGGILASELSVDEAAVHAAVLAINEAVERGVVEQTMVALHNPSAMLLNLRENLAAVYQEMLFQVKMEKAGNARNRVRADPESEDIYDRCLTQAEIQGNINKVNVHGALEYVDDALERQDPLALCQALQDPVLALRGVQRENAAWYLDQLSTDREQKALQEEVQAGVYAANERGDQEHAMRRAVSRINASIRRGEAMETVRELMSPEAQLPKAFPFAAQLYQHELSLLQQHHPQLGQEELYVAVEMLSAVALIDRALEAGDVSGFWSSLVSPALGLSDIEDANAQRYFDDLLKLKRRSGKAGVAFLSWNDIQACVNSANLAVQDENDKILAIRLINEALAQDDPEKTLTALLLPSARFSAVTLPGARRYHDVLTAALKQKAQVGPLALRRLEQLAGEGVEMALGIAAINQAIKEGKPAQTARVLRNPDVSLRGIVAECAGAYQAQLAALLAAKRQAGNANHNWVRHQLKDGSVYYFSLQNFEGSWERPHDCVLNSTHLSREEIQSTITGVTAAHDRERLWESNAGFVVRLQARMRGYLVRQEFVARRHLLQEQLPAVIKIQVRSAARLPSRSHADGHDDIYIALGLQGINAVIKIQAFVRANKARGDYRMLVHARDPPLSIVRRFVHLLEQSQHDFWEESELLKLREEVVKRIRSNQQLESDLNLMDIKIGLLVKNRITLQEVVSHCKKLTRKNKEQLSNLMAIDKQKGLKSLSKEKRQKLEAYQHLFYLLQTHPVYLAKLIFQMPQNKSTKFMESVVFTLYNYASNPREAYLLLQLFKTALQEEIRSKVDQLRDILTGNPTVIRLVVSFYRNARGQNALRQILGSPVQEVLQDKTLSIRTNPVDIYKAWINQLESQSGQKSKFPYDVSPEQALSHPEVQRRLDISIRNLLAVTDKFLSAITSSVDKIPYGMRYVAKVLKTSLAEKFPAAGEEDICKIVGNLLYYRFMNPAVVAPDGFDIVDISAGVALHPEHRRNLGSIAKILQHAAANKMFEGENGHLRVVNQYLEETHSKFRLFICAACSVPEPEERFNVDEYSEMVAVAKPVIYITVAELINTHKLLLEHQDSISPDHKDPLHELLEDLDELPTVQSLIGTCRLLVSFKPAPPLVSSFNPQNGTKQMLVDVIQSQPGDSLLEILRTAASEDQEAFHHHQMHRRALREAQTPDRLKRNRSLAGNSQLSMEEKKRKVIRNLRRLENLGVVDSANQYQEIINEMAKDIRNQRRYRQHRKGELVKLRQTLCGLNSKTSFYEEQIDYYNQYIRTCLDNLAASSKLNGKSKKQQSLHYTAARLFEKGVLLEIEDLPVSQFRNVIFDIIPCGEAGKFQVKAKFMGIDMEKFQLHYQDLLQLQYEGVAVMKLFDKAKVNVNLLIFLLNKKFFRK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INB2</accession>
    <name>A0A452INB2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Neurabin-1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="14">Neurabin-I</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="13">Neural tissue-specific F-actin-binding protein I</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="15">Protein phosphatase 1 regulatory subunit 9A</fullName>
        </alternativeName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
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        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
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<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
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    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149">
        <property type="entry name" value="RAS GTPASE-ACTIVATING PROTEIN WITH IQ MOTIF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149:SF10">
        <property type="entry name" value="RAS GTPASE-ACTIVATING-LIKE PROTEIN IQGAP3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00307">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00612">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00616">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03836">
        <property type="entry name" value="RasGAP_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00033">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00015">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00323">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47576">
        <property type="entry name" value="Calponin-homology domain, CH-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF143885">
        <property type="entry name" value="RGC domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50021">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00509">
        <property type="entry name" value="RAS_GTPASE_ACTIV_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50018">
        <property type="entry name" value="RAS_GTPASE_ACTIV_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01159">
        <property type="entry name" value="WW_DOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50020">
        <property type="entry name" value="WW_DOMAIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="4" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="1" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="Calponin-homology (CH)" evidence="7">
        <location>
            <begin position="33"/>
            <end position="148"/>
        </location>
    </feature>
    <feature type="domain" description="WW" evidence="6">
        <location>
            <begin position="669"/>
            <end position="702"/>
        </location>
    </feature>
    <feature type="domain" description="Ras-GAP" evidence="5">
        <location>
            <begin position="960"/>
            <end position="1193"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="4">
        <location>
            <begin position="157"/>
            <end position="184"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50020"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029358.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1598" mass="181898" checksum="26B8654B326EF82E" modified="2019-05-08" version="1">SSQYAAGDGREGLRLTADEMDEQRRQNVAYQYLCRLEEAKRWMEACLNEELASPTELEESLRNGVILAKLGHCFAPDIIPLKKIYDREQMRYEATGLHFRHTDNINHWREAMAHLGLPTIFHPETTDIYDKKNMPRVVYCIHALSLYLFKLGLAPQIQDLYGKVNFTEEEINNMKRELEKYGLQMPAFSKIGGILASELSVDEAAVHAAVLAINEAVERGVVEQTMVALHNPSAMLLNLRENLAAVYQEMLFQVKMEKAGNARNRVRADPESEDIYDRCLTQAEIQGNINKVNVHGALEYVDDALERQDPLALCQALQDPVLALRGVQRENAAWYLDQLSTDREQKALELGYVDLLEQEEVQAGVYAANERGDQEHAMRRAVSRINASIRRGEAMETVRELMSPEAQLPKAFPFAAQLYQHELSLLQQHHPQGELGQEELYVAVEMLSAVALIDRALEAGDVSGFWSSLVSPALGLSDIEDANAQRYFDDLLKLKRRSGKAGVAFLSWNDIQACVNSANLAVQDENDKILAIRLINEALAQDDPEKTLTALLLPSARFSAVTLPGARRYHDVLTAALKQKAQVTGDDGAALWWEEIQQGVSKANQDTEAARRLALGIAAINQAIKEGKPAQTARVLRNPDVSLRGIVAECAGAYQAQLAALLAAKRQAGNANHNWVRHQLKDGSVYYFSLQNFEGSWERPHDCVLNSTHLSREEIQSTITGVTAAHDRERLWESNAGFVVRLQARMRGYLVRQEFVARRHLLQEQLPAVIKIQSCWRGYRQRRDYLQRLRYLQKTRDAVIKVLKPCLEISLCPLPVHARDPPLSIVRRFVHLLEQSQHDFWEESELLKLREEVVKRIRSNQQLESDLNLMDIKIGLLVKNRITLQEVVSHCKKLTRKNKEQLSNLMAIDKQKGLKSLSKEKRQKLEAYQHLFYLLQTHPVYLAKLIFQMPQNKSTKFMESVVFTLYNYASNPREAYLLLQLFKTALQEEIRSKVDQLRDILTGNPTVIRLVVSFYRNARGQNALRQILGSPVQEVLQDKTLSIRTNPVDIYKAWINQLESQSGQKSKFPYDVSPEQALSHPEVQRRLDISIRNLLAVTDKFLSAITSSVDKIPYGMRYVAKVLKTSLAEKFPAAGEEDICKIVGNLLYYRFMNPAVVAPDGFDIVDISAGVALHPEHRRNLGSIAKILQHAAANKMFEGENGHLRVVNQYLEETHSKFRKFICAACSVPEPEERFNVDEYSEMVAVAKPVIYITVAELINTHKLLLEHQDSISPDHKDPLHELLEDLDELPTVQSLIGTSLAQLGKQEVSLTLTNKFDAVEASSEEADTRSLLLSTKQMLVDVIQSQPGDSLLEILRTAASEDQEAFHHHQMHRRALREAQTPDRLKRNRSLAGNSQLSMEEKKRKVIRNLRRLENLGVVDSANQYQEIINEMAKDIRNQRRYRQHRKGELVKLRQTLCGLNSKTSFYEEQIDYYNQYIRTCLDNLAASSKLNGKSKKQQSLHYTAARLFEKGVLLEIEDLPVSQFRNVIFDIIPCGEAGKFQVKAKFMGIDMEKFQLHYQDLLQLQYEGVAVMKLFDKAKVNVNLLIFLLNKKFFRK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INC2</accession>
    <name>A0A452INC2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Serologically defined colon cancer antigen 8</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452INC2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033443.1">
        <property type="protein sequence ID" value="ENSGAGP00000029453.1"/>
        <property type="gene ID" value="ENSGAGG00000021276.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005814">
        <property type="term" value="C:centriole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007098">
        <property type="term" value="P:centrosome cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030010">
        <property type="term" value="P:establishment of cell polarity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001764">
        <property type="term" value="P:neuron migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035148">
        <property type="term" value="P:tube formation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031887">
        <property type="entry name" value="SDCCAG8"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR34343">
        <property type="entry name" value="SEROLOGICALLY DEFINED COLON CANCER ANTIGEN 8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR34343:SF1">
        <property type="entry name" value="SEROLOGICALLY DEFINED COLON CANCER ANTIGEN 8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15964">
        <property type="entry name" value="CCCAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="34"/>
            <end position="60"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="81"/>
            <end position="106"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="125"/>
            <end position="166"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="229"/>
            <end position="270"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="305"/>
            <end position="527"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="562"/>
            <end position="603"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="640"/>
            <end position="709"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029453.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="5">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="719" mass="83877" checksum="1418F52EDE5F4E6B" modified="2019-05-08" version="1">MREAREESDLERYQRSLRDRASKSIHQLKYALVEGDPTAGESEVAFDQSSTGNDESGETAWQELQHSHAVNQLKALLREQEGLESEVSPSRRRKMSPSRTSEAADQSLPTLCDLVPIINDQSQYIHHLEAEVKFCKEELSGMKDRIQVVVLENEKLNEELKSLTVEQTLREQTLLDASANVRNSWTVGGDDSYLCQPAKSSPSHNKEQVLATRDTAETEKWQLELEKLKRLYQERTDTLEAQVRSLRKDLSESEKNCEDLKGRLKHQESLIATSNYNRVGGLCLKCAQHEAVLSQTHSNVHMHTIERLTKERDDLMGALVSVRRDLSEMQQRESSAYEQVKQAVQMTEEANLEKMKALVQCEQLKSEMERQKDRLERELIAQLDKRAGEKEALRGEMKKEREDLASTVMVLSQNVATLEARLERIIREKNSIANQLEEAQNQLTSQEMEVNKVCGEMRFQLNQTKMKKDEAEKEHREYRTKTIRELEIKDQEIEKLGLELNENKLRLEQAQQDAARAKDECLKLTELLGKSEHQLHLTRKLNKKLVHYDYRLKKESIQHSFSNEAKTRALQAEQREQELTQKMQQMEAQHDKTVNELDSLLTSQNTFIAKLKEECCMLARKLEHVTEKNRSEVSQLSQENEYFHDRLEKMQKRNDELEEQCIQHGRMHERMKLRLHQLDKHCQTTAQQLVELLNKQNQLFRERQMLTEEVQFLRTQIEA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INC4</accession>
    <name>A0A452INC4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="22">Sodium/calcium exchanger membrane region domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>Ca(2+)(out) + K(+)(out) + 4 Na(+)(in) = Ca(2+)(in) + K(+)(in) + 4 Na(+)(out)</text>
            <dbReference type="Rhea" id="RHEA:69967"/>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
            <dbReference type="ChEBI" id="CHEBI:29103"/>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the Ca(2+):cation antiporter (CaCA) (TC 2.A.19) family. SLC24A subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452INC4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033382.1">
        <property type="protein sequence ID" value="ENSGAGP00000029396.1"/>
        <property type="gene ID" value="ENSGAGG00000021253.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005262">
        <property type="term" value="F:calcium channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008273">
        <property type="term" value="F:calcium, potassium:sodium antiporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006874">
        <property type="term" value="P:intracellular calcium ion homeostasis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1420.30:FF:000005">
        <property type="entry name" value="sodium/potassium/calcium exchanger 3 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1420.30:FF:000006">
        <property type="entry name" value="sodium/potassium/calcium exchanger 4 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1420.30">
        <property type="entry name" value="NCX, central ion-binding region"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004481">
        <property type="entry name" value="K/Na/Ca-exchanger"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004837">
        <property type="entry name" value="NaCa_Exmemb"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044880">
        <property type="entry name" value="NCX_ion-bd_dom_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00367">
        <property type="entry name" value="calcium/sodium antiporter"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10846">
        <property type="entry name" value="SODIUM/POTASSIUM/CALCIUM EXCHANGER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10846:SF42">
        <property type="entry name" value="SODIUM_POTASSIUM_CALCIUM EXCHANGER 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01699">
        <property type="entry name" value="Na_Ca_ex"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0050">Antiport</keyword>
    <keyword evidence="11" id="KW-0106">Calcium</keyword>
    <keyword evidence="7" id="KW-0109">Calcium transport</keyword>
    <keyword evidence="5" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="17" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="15" id="KW-0406">Ion transport</keyword>
    <keyword evidence="16 21" id="KW-0472">Membrane</keyword>
    <keyword evidence="12" id="KW-0630">Potassium</keyword>
    <keyword evidence="6" id="KW-0633">Potassium transport</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0677">Repeat</keyword>
    <keyword evidence="9" id="KW-0732">Signal</keyword>
    <keyword evidence="14" id="KW-0915">Sodium</keyword>
    <keyword evidence="18" id="KW-0739">Sodium transport</keyword>
    <keyword evidence="8 21" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="13 21" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="45"/>
            <end position="67"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="136"/>
            <end position="154"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="175"/>
            <end position="199"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="205"/>
            <end position="227"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="239"/>
            <end position="258"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="264"/>
            <end position="284"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="515"/>
            <end position="534"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="541"/>
            <end position="564"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="584"/>
            <end position="604"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="611"/>
            <end position="629"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="641"/>
            <end position="667"/>
        </location>
    </feature>
    <feature type="domain" description="Sodium/calcium exchanger membrane region" evidence="22">
        <location>
            <begin position="141"/>
            <end position="281"/>
        </location>
    </feature>
    <feature type="domain" description="Sodium/calcium exchanger membrane region" evidence="22">
        <location>
            <begin position="507"/>
            <end position="658"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="1"/>
            <end position="38"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="414"/>
            <end position="468"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="20">
        <location>
            <begin position="1"/>
            <end position="10"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="20">
        <location>
            <begin position="433"/>
            <end position="463"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005364"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022449"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022538"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022568"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022958"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023053"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023201"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00033627"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="Pfam" id="PF01699"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029396.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="675" mass="75407" checksum="D8AFE39832D7E1AA" modified="2019-05-08" version="1">MEAAKEKRRQPVPAAVAGARFSRPEMLPLQPGKRHRSQARRRRELLLAQLCLGGSLALLLASLSSWAGQSGYDVLNYPVETYDKWSGRRLLQATEDNATEIPEETENRYVKNCTEPALHEFPSDIFTNEDRRHGAVVLHVLCAMYMFYALAIVCDDFFVPSLEKISERLHLSEDVAGATFMAAGSSAPELFTSVIGVFITKGDVGVGTIVGSAVFNILCIIGVCGLFAGQVIALSSWSLLRDSTYYTLSVVALIVFIYDEIVSWWESLVLVLMYFIYILIMKYNSSIHHCFERKTKNAPNMVNGLASNTEMDDNSNCDATVVLLKKGNCSRNFHRKASVIMVDELLSAYPHQLSFSEAGLRIMITSHFPPKTRLSMASRMLINERQRLINSRAYTNGESEVAIKIPIKHALENGTVPSNSAERGVNGTRRDEDMAEAGNETENEDNENNENDEEEEEEEDDNDGPYTPFDLPSGKMEILKWLFTWPLSFLLHFTVPNCNKPYWEKWFMVTFASSTLWIAALSYVMVWMVTIIGYTLGIPDVIMGITFLAAGTSVPDCMASLIVARQGMGDMAVSNSIGSNVFDILIGLGLPWALQTLAVDYGSYIQLNSRGLIYSVGLLLASVFVTVFGVHMNKWKLDKKLGIVCLFLYGIFLCFSIMTEFNVFTFVNLPMCRDH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INC5</accession>
    <name>A0A452INC5_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="10">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452INC5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033349.1">
        <property type="protein sequence ID" value="ENSGAGP00000029368.1"/>
        <property type="gene ID" value="ENSGAGG00000021101.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004222">
        <property type="term" value="F:metalloendopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016485">
        <property type="term" value="P:protein processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd08662">
        <property type="entry name" value="M13"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.390.10">
        <property type="entry name" value="Collagenase (Catalytic Domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1380.10">
        <property type="entry name" value="Neutral endopeptidase , domain2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024079">
        <property type="entry name" value="MetalloPept_cat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000718">
        <property type="entry name" value="Peptidase_M13"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018497">
        <property type="entry name" value="Peptidase_M13_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042089">
        <property type="entry name" value="Peptidase_M13_dom_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008753">
        <property type="entry name" value="Peptidase_M13_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11733:SF141">
        <property type="entry name" value="MEMBRANE METALLO-ENDOPEPTIDASE-LIKE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11733">
        <property type="entry name" value="ZINC METALLOPROTEASE FAMILY M13 NEPRILYSIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01431">
        <property type="entry name" value="Peptidase_M13"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05649">
        <property type="entry name" value="Peptidase_M13_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00786">
        <property type="entry name" value="NEPRILYSIN"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55486">
        <property type="entry name" value="Metalloproteases ('zincins'), catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51885">
        <property type="entry name" value="NEPRILYSIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="6" id="KW-0482">Metalloprotease</keyword>
    <keyword evidence="2" id="KW-0645">Protease</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="Peptidase M13 N-terminal" evidence="9">
        <location>
            <begin position="44"/>
            <end position="460"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase M13 C-terminal" evidence="8">
        <location>
            <begin position="521"/>
            <end position="708"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="464"/>
            <end position="507"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023049"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF01431"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF05649"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029368.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="710" mass="81818" checksum="24EA2D33CBCC9A4F" modified="2019-05-08" version="1">MALCGFIPPTDMQYGTNSGSICTTPGCVTAAARILQNMDPTTEPCTDFYQYACGGWLNWHVIPETSSRYSIFDILRDELEIILKGVLESPEQGDREAFRKAKMLYTSCMNESFIEQRDSLPLLEVLAMVGDWPVASADWNITKGNKPNWSMEEKLSTLNSRFNKRVFIDMFVWNDDQDSSRHIIYIDQPSLGMPSRDYYFNGGNYQRVREAYLQFMITIAKMIREDKNMAKDDTFVQQEMAKVMKLETEIANATAPAEERHDVTLLYNKMTLRELQEKFALNVSLGIGVCLSLPSQEFNWTLFIQGIMSSVNVQIHLDEEVVVYGAPYLQELKAIISNYSASTIQNYLVWRLVTDRVSSLSRRFKDARAHYRKALYGTTLEEARWRECVSYVNNNMENAVGALYVRETFAGESKRMVRDLIDKIREVFIETLDELQWMDETSKEKAREKAMAIKEQIGYPDYILEDQNEKLDQEYANLNFSEHKYFENSLENLRAGAQKSLKKLRERVDQDIWIIGAAVVNAFYSPNRNQIVFPAGILQPPFFSKHQPQALNFGGIGMVIGHEITHGFDDNGRNFDKDGNMFDWWSNFSAMHFKDQSRCMVYQYGNYTWELAGGQNVSGISTLGENIADNGGVRQAYKAYLKWLEREGKEPKLPGLDLTHKQLFFLNFAQVWCGSYRPEYAIVRTWATFSEAFHCKKGTAMHPVEKCRVW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INC6</accession>
    <name>A0A452INC6_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="11">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452INC6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033345.1">
        <property type="protein sequence ID" value="ENSGAGP00000029364.1"/>
        <property type="gene ID" value="ENSGAGG00000021101.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004222">
        <property type="term" value="F:metalloendopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016485">
        <property type="term" value="P:protein processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd08662">
        <property type="entry name" value="M13"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.390.10">
        <property type="entry name" value="Collagenase (Catalytic Domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1380.10">
        <property type="entry name" value="Neutral endopeptidase , domain2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024079">
        <property type="entry name" value="MetalloPept_cat_dom_sf"/>
    </dbReference>
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        <property type="entry name" value="MEMBRANE METALLO-ENDOPEPTIDASE-LIKE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="ZINC METALLOPROTEASE FAMILY M13 NEPRILYSIN-RELATED"/>
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        <property type="entry name" value="Peptidase_M13_N"/>
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        <property type="entry name" value="NEPRILYSIN"/>
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        <property type="entry name" value="Metalloproteases ('zincins'), catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="NEPRILYSIN"/>
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    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="8" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="6" id="KW-0482">Metalloprotease</keyword>
    <keyword evidence="2" id="KW-0645">Protease</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="21"/>
            <end position="48"/>
        </location>
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    <feature type="domain" description="Peptidase M13 N-terminal" evidence="10">
        <location>
            <begin position="65"/>
            <end position="465"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase M13 C-terminal" evidence="9">
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            <begin position="526"/>
            <end position="732"/>
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        <location>
            <begin position="469"/>
            <end position="512"/>
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            <dbReference type="ARBA" id="ARBA00022670"/>
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        <source>
            <dbReference type="SAM" id="Coils"/>
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        <source>
            <dbReference type="SAM" id="Phobius"/>
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    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF01431"/>
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    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="Pfam" id="PF05649"/>
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    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029364.1"/>
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    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="733" mass="84780" checksum="7B991DF924C16741" modified="2019-05-08" version="1">MGKSESQMDIVEKSTKSGKKSWSFVTIGLSVLLLLMTCAVVALVILYANSRAARILQNMDPTTEPCTDFYQYACGGWLNWHVIPETSSRYSIFDILRDELEIILKGVLESPEQGDREAFRKAKMLYTSCMNESFIEQRDSLPLLEVLAMVGDWPVASADWNITKEPNWSMEEKLSTLNSRFNKRVFIDMFVWNDDQDSSRHIIYIDQPSLGMPSRDYYFNGGNYQRVREAYLQFMITIAKMIREDKNMAKDDTFVQQEMAKVMKLETEIANATAPAEERHDVTLLYNKMTLRELQEKFALNEFNWTLFIQGIMSSVNVQIHLDEEVVVYGAPYLQELKAIISNYSASTIQNYLVWRLVTDRVSSLSRRFKDARAHYRKALYGTTLEEARWRECVSYVNNNMENAVGALYVRETFAGESKRMVRDLIDKIREVFIETLDELQWMDETSKEKAREKAMAIKEQIGYPDYILEDQNEKLDQEYANLNFSEHKYFENSLENLRAGAQKSLKKLRERVDQDIWIIGAAVVNAFYSPNRNQIVFPAGILQPPFFSKHQPQALNFGGIGMVIGHEITHGFDDNGRNFDKDGNMFDWWSNFSAMHFKDQSRCMVYQYGNYTWELAGGQNVSGISTLGENIADNGGVRQAYKAYLKWLEREGKEPKLPGLDLTHKQLFFLNFAQVWCGSYRPEYASQSIKTDVHSPLKYRVMGSLQNFEAFSEAFHCKKGTAMHPVEKCRVW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INC8</accession>
    <name>A0A452INC8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Regulator of G-protein signaling 12</fullName>
        </recommendedName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">Regulates G protein-coupled receptor signaling cascades. Inhibits signal transduction by increasing the GTPase activity of G protein alpha subunits, thereby driving them into their inactive GDP-bound form.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell projection</location>
            <location evidence="2">Dendrite</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="14">Synapse</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000033598.1">
        <property type="protein sequence ID" value="ENSGAGP00000029588.1"/>
        <property type="gene ID" value="ENSGAGG00000021288.1"/>
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        <property type="component" value="Unassembled WGS sequence"/>
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        <property type="term" value="C:cytoplasm"/>
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        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030425">
        <property type="term" value="C:dendrite"/>
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        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
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        <property type="project" value="UniProtKB-SubCell"/>
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    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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        <property type="term" value="C:synapse"/>
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        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
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        <property type="term" value="F:GTPase activator activity"/>
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        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008277">
        <property type="term" value="P:regulation of G protein-coupled receptor signaling pathway"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
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    </dbReference>
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        <property type="entry name" value="Phosphatidylinositol 3-kinase Catalytic Subunit, Chain A, domain 1"/>
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        <property type="entry name" value="PDZ_sf"/>
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        <property type="entry name" value="RGS_subdomain_2"/>
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    <dbReference type="InterPro" id="IPR029071">
        <property type="entry name" value="Ubiquitin-like_domsf"/>
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    <dbReference type="PANTHER" id="PTHR45945:SF1">
        <property type="entry name" value="REGULATOR OF G-PROTEIN SIGNALING 12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45945">
        <property type="entry name" value="REGULATOR OF G-PROTEIN SIGNALING LOCO"/>
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    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="PID"/>
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        <property type="entry name" value="RGS12_us1"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="RGS12_us2"/>
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    <dbReference type="Pfam" id="PF16612">
        <property type="entry name" value="RGS12_usC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01301">
        <property type="entry name" value="RGSPROTEIN"/>
    </dbReference>
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        <property type="entry name" value="GoLoco"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="2"/>
    </dbReference>
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    </dbReference>
    <dbReference type="SUPFAM" id="SSF48097">
        <property type="entry name" value="Regulator of G-protein signaling, RGS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54236">
        <property type="entry name" value="Ubiquitin-like"/>
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    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
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    </dbReference>
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    </dbReference>
    <dbReference type="PROSITE" id="PS50132">
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    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="13" id="KW-0966">Cell projection</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="7" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="5" id="KW-0488">Methylation</keyword>
    <keyword evidence="12" id="KW-0539">Nucleus</keyword>
    <keyword evidence="8" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0677">Repeat</keyword>
    <keyword evidence="11" id="KW-0770">Synapse</keyword>
    <keyword evidence="10" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="PDZ" evidence="19">
        <location>
            <begin position="22"/>
            <end position="99"/>
        </location>
    </feature>
    <feature type="domain" description="PID" evidence="18">
        <location>
            <begin position="227"/>
            <end position="339"/>
        </location>
    </feature>
    <feature type="domain" description="RGS" evidence="20">
        <location>
            <begin position="708"/>
            <end position="825"/>
        </location>
    </feature>
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        <location>
            <begin position="901"/>
            <end position="971"/>
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            <begin position="973"/>
            <end position="1043"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
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        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
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        </location>
    </feature>
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        <location>
            <begin position="1159"/>
            <end position="1220"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1286"/>
            <end position="1338"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="840"/>
            <end position="849"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="1061"/>
            <end position="1088"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="17">
        <location>
            <begin position="1090"/>
            <end position="1103"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="1193"/>
            <end position="1208"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="1320"/>
            <end position="1338"/>
        </location>
    </feature>
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        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00004279"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00034103"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00059154"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00069205"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS01179"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50106"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50132"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50898"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029588.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1338" mass="147858" checksum="19E7205CBF270523" modified="2019-05-08" version="1">MYRPGETVKCRLNMHAPPRIRSVEVARGRAGYGFTLSGQAPCVLSCVMKGSPADYVGLKAGDEIFAINEINVKQASHEDVVKLIGKCSGVLHMVIAEGNSHIDSCSSDEEVGFYDGKGWLKPKPDSKALGINRAEKVVEEMQSGGIFNMIFENPSFCAGNAENSVPKQRSLSESSAITFETGHESINNNPNLLSKEEISKVLHDDSVFQIGLENPEDFGLDASILNVAMVVGYLGSIELPSTSSNLENDSLQAIRGCMRRLRAEQKIHSLVMMKIMHDCIQLCSDKSGVVAEYPAEKLAFSAVCPDDRRFFGLVTMQTNDDASLAQEDEGVLRTSCHVFMVDPELFHHKIHQGIARRFGLECTADPDTNGCLEFPTSSLPVLQFISVLYRDMGELIEGMRARAFLDGDADAHQNNSTSSNSDSGIGNFNQEEKSNRVLVVDLGGNPSKHIPNSIWENPVGRGQNQHASHWNGFCQEQEGNAHLEIIQNDKSQNLSKHLSPSARIEVPLVSSRSSVPPSKKSIVGIGNQRWLPVHVLQEWQHGNASDQESYTDSTDGWSSINCGTLPPPMNKIPADRYRVDGSFGQRQLTSYINEWSKNVFCTHKKFGPQHNIRKSKEAKKGAKFGHPVGLALAPQRVSVRRSFGRSKRFSITRSLDDLESATVSDGELNSSDLKDCISDNSLSSNASLPSVQSCRRLRERRVASWAVSFERLLQDPVGVRFFSEFLRKEFSEENILFWQACEYFNHVPAHDKKELSYRAREIFSKFLCSKATTPVNIDSQAQLADDILSAPHPDMFKEQQLQIFNLMKFDSYTRFLKSPLYQECILAEVEGRPLPDPQRVPSSPTSKHSINSERSHISTPKKFDLLDCNTVCLFLTFIFVSDKSDASRLSASVPDKEKSSKYCCINLPDGSSCKMVVKSGFSVKEMLSGLCEKHGINIAAVDLFLVGGDKPLVLHQDSSILDSRDLRLEKRTLFRLDLVPINRSVGLKAKPTKPVTEVLRPVVAKYGLNLNELVAKLNGEQEPLDLGVPISNLDGQRVVLYEKEPTKGRVFTDRQKGAPVKQSTVVTSTSRNQASTGEGRTLGKSNSIKIKGENGKNARDARLSKREESLAKIGEKKYQKINLDEAEEFFELISKAQSNRADDQRGLLRKEDLVLPDFLRLPLSGPEPPSSTPVSSKGLNKRIAKNDGKDKCTQPSGKLETTQNYNENSIKKTGHSENKHKSALTTLNSQKTYNVPFSAPLSPIPHVQENNMTIWKRQSGELQAEGIQTIDDENVADLTLVAEGDISSPNSTLLPPPPPTPLSDISKLTEANYPPPTPLAGNQENSGYQRSNSGIASF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IND2</accession>
    <name>A0A452IND2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">non-specific serine/threonine protein kinase</fullName>
            <ecNumber evidence="2">2.7.11.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the protein kinase superfamily. AGC Ser/Thr protein kinase family. RAC subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452IND2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033460.1">
        <property type="protein sequence ID" value="ENSGAGP00000029466.1"/>
        <property type="gene ID" value="ENSGAGG00000021286.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd01241">
        <property type="entry name" value="PH_PKB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000033">
        <property type="entry name" value="Non-specific serine/threonine protein kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000027">
        <property type="entry name" value="Non-specific serine/threonine protein kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000961">
        <property type="entry name" value="AGC-kinase_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039026">
        <property type="entry name" value="PH_PKB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017892">
        <property type="entry name" value="Pkinase_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24351">
        <property type="entry name" value="RIBOSOMAL PROTEIN S6 KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00169">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00433">
        <property type="entry name" value="Pkinase_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00233">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00133">
        <property type="entry name" value="S_TK_X"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51285">
        <property type="entry name" value="AGC_KINASE_CTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 9 10" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="7" id="KW-0418">Kinase</keyword>
    <keyword evidence="6 9 10" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 10" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="5" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="PH" evidence="12">
        <location>
            <begin position="30"/>
            <end position="132"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="13">
        <location>
            <begin position="112"/>
            <end position="385"/>
        </location>
    </feature>
    <feature type="domain" description="AGC-kinase C-terminal" evidence="14">
        <location>
            <begin position="386"/>
            <end position="459"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="438"/>
            <end position="459"/>
        </location>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="212"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006935"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00012513"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="RuleBase" id="RU000304"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS51285"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029466.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="459" mass="53303" checksum="23026EDE5A1DDB62" modified="2019-05-08" version="1">MKKIVQANSDTVNVGHGYSTVTSHIKVFFFFSINAYFSHIGEYIKNWRPRYFLLKTDGSFIGYKEKPQDVDLPYPLNNFSVAKCQLMKTERPKPNTFIIRCLQWTTVIERTFHVDTPEEREEWTEAIQAVADRLQRQEEERMNCSPTSQIDNIGEEEMDASTTHHKRKTMNDFDYLKLLGKGTFGKVILVREKASGKYYAMKILKKEVIIAKLFFHLSRERVFSEDRTRFYGAEIVSALDYLHSGKIVYRDLKLENLMLDKDGHIKITDFGLCKEGITDAATMKTFCGTPEYLAPEVLEDNDYGRAVDWWGLGVVMYEMMCGRLPFYNQDHEKLFELILMEDIKFPRTLSSDAKSLLSGLLIKDPNKRLGGGPDDAKEIMRHSFFVGVNWQDVYDKKLVPPFKPQVTSETDTRYFDEEFTAQTITITPPEKYDEDGMDCMDNERRPHFPQFSYSASGRE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IND3</accession>
    <name>A0A452IND3_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IND3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033605.1">
        <property type="protein sequence ID" value="ENSGAGP00000029593.1"/>
        <property type="gene ID" value="ENSGAGG00000021375.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000033614.1">
        <property type="protein sequence ID" value="ENSGAGP00000029602.1"/>
        <property type="gene ID" value="ENSGAGG00000021375.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007288">
        <property type="term" value="P:sperm axoneme assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000225">
        <property type="entry name" value="Armadillo"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038905">
        <property type="entry name" value="ARMC2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21356">
        <property type="entry name" value="ARMADILLO REPEAT CONTAINING 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21356:SF1">
        <property type="entry name" value="ARMADILLO REPEAT-CONTAINING PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00185">
        <property type="entry name" value="ARM"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1"/>
            <end position="125"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="147"/>
            <end position="173"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="221"/>
            <end position="260"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="58"/>
            <end position="81"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="150"/>
            <end position="159"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="221"/>
            <end position="250"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029593.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="864" mass="97744" checksum="514E45010B90F97D" modified="2019-05-08" version="1">MHSSKNKNVEKPEPFYRLSAPKQKTSAEIINEARNALRTLRTQRPFTPREDQRKLFGSASSRTLENRPPSTFSLHASSFESGESRPVSGTRLSPLDHKPKLLSSPIKDEDSSISFPKPPVDPMEVRKVSSARARLCRTASQGNLLPDKIFLPDENKEGVNSEEPATMDDLSRRSNKSCLAELSPHSNFKDDGIPLIYNEHNKAEEQDFLKEAVATSPLQFRSGSDQIKNGDRASSNLSRISWNTSGSKAGSENDSRVNETEEEENYWNVKIVPIVYELENEDNIENLCRACTSLHQALEEGNMFGKRFKRRSALLKLLYKLVDIGSDPLGLKLARMILALKVSGKNLLNVCKLVFKISRNEKNDSLIQNDSLLDSLLEILRFEDLQTNNEAFLYCLGAIKFISGNAVLLNELVNKGTVEILFQLMKQINNIKENDTRFSNLGHLLVQLTATLRNLVDLPLSRCQLLCSGAVSELWVVLEQRMNDKDICANIVRIFSKLSSYNDCCAALAACSRCYVLFLALLNKHQKKQDLVVRVVFILGNLTAKNNQAREQFFKKKGSADTLISLFQTYNEFDLNTKKEKHEGEENKSKQKHPSEAEDVLIKLIRVLANLSIHPNVGAVLAANHHIVALLITVLEYKSVDDCEELVINAAATINNLSYYKVRNSVIEDKKLHIAELLLKLLMSSNMDGILEAVRVFGNLSQYLEICDFIVQKKVYKYMIALLDAKHQDVCFSACGVLLNLTVDKNKRAILKEEGGIKKLVDCLRDFGPTDWQLACLICKTLWNYSENITNAASCFEEDDINTLLVLLALFLDEELALDYSFDRDLRDYHKLYWETEFKPVAQKLLNRIQSHHSFLQPITITSF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IND4</accession>
    <name>A0A452IND4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">non-specific serine/threonine protein kinase</fullName>
            <ecNumber evidence="3">2.7.11.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="12">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033392.1">
        <property type="protein sequence ID" value="ENSGAGP00000029406.1"/>
        <property type="gene ID" value="ENSGAGG00000021255.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.20.90:FF:000007">
        <property type="entry name" value="Serine/threonine-protein kinase WNK1 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000006">
        <property type="entry name" value="Serine/threonine-protein kinase WNK1 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.20.90:FF:000012">
        <property type="entry name" value="Serine/threonine-protein kinase WNK1 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000494">
        <property type="entry name" value="serine/threonine-protein kinase WNK2 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.20.90">
        <property type="entry name" value="Phosphatidylinositol 3-kinase Catalytic Subunit, Chain A, domain 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056865">
        <property type="entry name" value="CCTL2_WNK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024678">
        <property type="entry name" value="Kinase_OSR1/WNK_CCT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050588">
        <property type="entry name" value="WNK_Ser-Thr_kinase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13902">
        <property type="entry name" value="SERINE/THREONINE-PROTEIN KINASE WNK WITH NO LYSINE -RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24889">
        <property type="entry name" value="CCTL2_WNK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12202">
        <property type="entry name" value="OSR1_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="10" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="13" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9" id="KW-0418">Kinase</keyword>
    <keyword evidence="8" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="7" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Protein kinase" evidence="15">
        <location>
            <begin position="192"/>
            <end position="450"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1"/>
            <end position="184"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="606"/>
            <end position="625"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1094"/>
            <end position="1135"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1209"/>
            <end position="1248"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1273"/>
            <end position="1302"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1384"/>
            <end position="1404"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1862"/>
            <end position="1926"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="2171"/>
            <end position="2244"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="13">
        <location>
            <begin position="1953"/>
            <end position="1980"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="1"/>
            <end position="12"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="110"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="125"/>
            <end position="144"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="147"/>
            <end position="173"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="606"/>
            <end position="615"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="616"/>
            <end position="625"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="14">
        <location>
            <begin position="1116"/>
            <end position="1126"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="1220"/>
            <end position="1229"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="1230"/>
            <end position="1239"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="1882"/>
            <end position="1891"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="1903"/>
            <end position="1926"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="2176"/>
            <end position="2185"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012513"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00047899"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00048679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029406.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2244" mass="242918" checksum="1B5CE48DADC452C4" modified="2019-05-08" version="1">METGCAASSAPPRGGGGASPAAARPPPADGSAAVGGLGRRSGAGMETAAVPERGPAPGLPPRGPGAREPRVGHLRFLRRSVVDSDQEEPPLELSEAEPRRIVLLSKTRRIIAERAKERHGEADLEPPAVEPAAEPKGAVPGVPATAEEQKELGRDAERKEPSDKDQSKSKKQEPEEEADMKAVSTSPDGRFLKFDIELGRGSFKTVYKGLDTETWVEVAWCELQDRKLTKVERQRFKEEAEMLKGLQHPNIVRFYDFWESCVKGKKCIVLVTELMTSGTLKTYLKRFKVMKPKVLRSWCRQILKGLLFLHTRTPPIIHRDLKCDNIFITGPTGSVKIGDLGLATLKRASFAKSVIGTPEFMAPEMYEEHYDESVDVYAFGMCMLEMATSEYPYSECQNAAQIYRKVTCGVKPASFEKVTDPEIKEIIGECICKNKEERYEIKDLLSHAFFAEDTGVRVELAEEDHGRKSSIALRLWVEDPKKLKGKPKDNGAIEFTFDLEKETPDDVAQEMVDSGFFHESDVKIVAKSIRDRVALIQWRRERIWPMIQSEDHQVSECLEKLKVPQAQQVQVTSLSHAGQQTLSALEEPEADQHLFQHNLPTSVTSVASDSTFDSGQGSTVYSDSQSSQQSVIYSSLPDSVPPAIQRVYSPPLSESQALPQSLQQLGHYQQPSVPGLTVAQAPPAVLSQRPQHYQEPAMTFPVIQPVTVASSQMGQSQPVLASQQQPIPQQAPLQQVLASQPVCPLQPAAHLLPQYQTQTSQVAATSTQLTPLQISTVRQLPHVPPSQIPQLPVIPAVTPLAGLDNLPPSLSDLPAANVPSVPAPSQYFTPTVIIPSLPMNPALPLASNPPALSMQAVNLPHTTVASLVLPCQTIVPNMSAATIPLLAVAPPGVSALPTHHAISHLSQQQMYQPAFQQIIQSDAPSPHHTQNMQAVSQQQLPPLPQSLQPSIIHPSEQTMSTSGVGNQIIGHPQYGLEIGAQMPVIPMQPSIVMPPPLQLQPEILPPCLGPESLSQLHGNLAAASATVPIDHCLPLQPTGLLQLQPDLSQPLGQQAPCSAIQSVTESTQEEHAIQEKQQTLSLCCESYMSPDVASGKEMSDSFEGVVGSGKQEGKPSKKHKKSTRVRSRQEKSSRPKLTILNVCNTGDKMVECQLETHNHKMVTFKFDLDGDAPEEIATYMVEKEFILQTEKETFIEQMKDIIDKAEDMLSEDTEGERSSDQGTSPQQDASRLEMNEENRQSQVKTPMYQQNVLHTGKRWFIICPVVENPTEEVPEFSPAVPQSKQAEGPDPEQSDDQQMGSAVTDTPSVFFCHQLPSQTSVFDSPCGTPAPLAQVPAAASSAVLQTKAEILAATLPGTAPHVPEQFVTNVGQPEYPPLHLVMPSQPDASAHNAASLEGPSDAQLAPKSSLQTLQEMDEDSCIPDVEISCCTVRPPSSILTVPAQGAEFCPVPALSDVVVLELQPAPQIPQVSEASQPGVMRHSFVNQLLPGGVVSDPLSLTGSQLQSPPQLPVAVSQQHTMLTQSQPVTCTGLGVGPLQPPCAVESDGEGPPRVDFADDTIKSLDEKLRNLLYQECVPTSAASAGTPDSFVPLEQGESELASLPPLNEELIPTLVLDSREPGLNVPDTFQEANAAQDQCVSLVPSAMPPYPVLASQALTVPFIPINPAAMETSAVLNNATATSHSRTEFRIQEPMLEKFETTSIDVSSSKVRVYETGSAANISSTSAVTTQGLQVAPASSTIPEQMETSRRNKEAKIKISSALTENLIQTSTTDREINNLQRNDFVDPGSCGKQSEVHGNSTPAKTIGRFSVISTHDELTLTSPHCLRYSAPPDVYLDELPSSPDLKTSVRRVQTASFLDVLSDHGSSDSGDESLRRKSVAAQLSPQSSIAANDITKKPASLLQRSGKTSSPGPDSPNGQGTKIPTINITSFHSQSSYMSSDNDSEFEDADMKKELQNLREKHMKEIAELQTQQKNEIEALFVRLGKPLPPNVGFLHSVPPTGRRRRTSKNKLKAGKLLNPLVQQLKTAPSNTSNLSVCKESLPKETTKTHAGSTEATAVTSSTSATFGNSVQTQQPCSVKASLSSDICSGLATEGGNAHGSSGQGSTLKRLCLGKDHSSRSSTNSLVTCPEPLPPPTLQVQVQANNSNNKKGTFTDDLHKLVDQWTSKTVGAAQSKPSLNQLKQNQKRQDMEPKANPMQAQNETCGILKMRMPLPPIDAGPAQLQYQAQGPMSAKEDEEEDVK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IND7</accession>
    <name>A0A452IND7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">glutamate carboxypeptidase II</fullName>
            <ecNumber evidence="18">3.4.17.21</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>Release of an unsubstituted, C-terminal glutamyl residue, typically from Ac-Asp-Glu or folylpoly-gamma-glutamates.</text>
            <dbReference type="EC" id="3.4.17.21"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Membrane</location>
            <topology evidence="2">Single-pass type II membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the peptidase M28 family. M28B subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.4.17.21" evidence="18"/>
    <dbReference type="AlphaFoldDB" id="A0A452IND7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033465.1">
        <property type="protein sequence ID" value="ENSGAGP00000029471.1"/>
        <property type="gene ID" value="ENSGAGG00000020963.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016805">
        <property type="term" value="F:dipeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004181">
        <property type="term" value="F:metallocarboxypeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd08022">
        <property type="entry name" value="M28_PSMA_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.930.40:FF:000001">
        <property type="entry name" value="N-acetylated-alpha-linked acidic dipeptidase 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.630.10:FF:000009">
        <property type="entry name" value="N-acetylated-alpha-linked acidic dipeptidase 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.50.30.30">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.930.40">
        <property type="entry name" value="Transferrin receptor-like, dimerisation domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.630.10">
        <property type="entry name" value="Zn peptidases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046450">
        <property type="entry name" value="PA_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007484">
        <property type="entry name" value="Peptidase_M28"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039373">
        <property type="entry name" value="Peptidase_M28B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007365">
        <property type="entry name" value="TFR-like_dimer_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036757">
        <property type="entry name" value="TFR-like_dimer_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10404:SF36">
        <property type="entry name" value="GLUTAMATE CARBOXYPEPTIDASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10404">
        <property type="entry name" value="N-ACETYLATED-ALPHA-LINKED ACIDIC DIPEPTIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04389">
        <property type="entry name" value="Peptidase_M28"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04253">
        <property type="entry name" value="TFR_dimer"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52025">
        <property type="entry name" value="PA domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47672">
        <property type="entry name" value="Transferrin receptor-like dimerisation domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53187">
        <property type="entry name" value="Zn-dependent exopeptidases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0121">Carboxypeptidase</keyword>
    <keyword evidence="12" id="KW-0224">Dipeptidase</keyword>
    <keyword evidence="15" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="14" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13" id="KW-0482">Metalloprotease</keyword>
    <keyword evidence="16" id="KW-0511">Multifunctional enzyme</keyword>
    <keyword evidence="5" id="KW-0645">Protease</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0735">Signal-anchor</keyword>
    <keyword evidence="6" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="9" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="Peptidase M28" evidence="20">
        <location>
            <begin position="298"/>
            <end position="409"/>
        </location>
    </feature>
    <feature type="domain" description="Transferrin receptor-like dimerisation" evidence="19">
        <location>
            <begin position="565"/>
            <end position="685"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004606"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005634"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022645"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022997"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023049"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023268"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00052003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00066561"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="Pfam" id="PF04253"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="Pfam" id="PF04389"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029471.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="688" mass="77546" checksum="EF7689B586E397E0" modified="2019-05-08" version="1">FRGQGCHPITAQPSPLQPDSICRGSVYPAKLTSNYKIQFLVLTANTCPFLFPSNFTQRPHLAGTAENLRLAQQIQAQWKEFGLDSVELAHYDVLLSYPNETNPNYISIIDEHGTEIFNTSLFEPPPPGYETVTDVVPPYSAFSAPGVPEVKSNRLLKSRKLKVSCSQLGLIFRGSWKIRPHKLYSPGNFQLMRFCRTAAGHLEHGSAKINTSLIPPEYTYRYEEARAVGLPKLPVHPIGYNDAEKLLRDMGGPAPPDSSWKGSLNVSYNIGPGFMSRYSTRKVKMHIHSSNKVTRIYNVIGTIRGATEPDRYVILGGHRDSWVFGGIDPQSGAAVVHEIVRSYGKLKKEGWRPRRTIIFASWDAEEFGLLGSTEWAEENTKLLQARGVAYINADSSVEGNYTLRIDCTPLMYSLVYSLTREIPSPDEGFKGKSLYESWHENNPSTEYKDVPRINKLGSGNDFEVFFQRLGIASGRARYSKNWVSLSYPVYHSVYETYEIVERFYDPTFKNHLTVAQVRGGLVFELADSVVLPFDCRDYAEALRNYVQVIYNLSKNHQMGLTTYSVSFDSLFSAANNFTEAATDFHRRLQQVDINNPIAVRALNDQLMFLERAFIDPLGLPGRPFYRHIIFAPSSHNKYAGESFPGIYDAMFDIENKADQRKAWEEVKRQISIAAFTVQAAAGTLKDLA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IND8</accession>
    <name>A0A452IND8_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="3">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IND8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033610.1">
        <property type="protein sequence ID" value="ENSGAGP00000029598.1"/>
        <property type="gene ID" value="ENSGAGG00000021375.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007288">
        <property type="term" value="P:sperm axoneme assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000225">
        <property type="entry name" value="Armadillo"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038905">
        <property type="entry name" value="ARMC2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21356">
        <property type="entry name" value="ARMADILLO REPEAT CONTAINING 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21356:SF1">
        <property type="entry name" value="ARMADILLO REPEAT-CONTAINING PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00185">
        <property type="entry name" value="ARM"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="2" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="49"/>
            <end position="66"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="120"/>
            <end position="201"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="227"/>
            <end position="253"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="138"/>
            <end position="161"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="230"/>
            <end position="239"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029598.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="913" mass="103553" checksum="91864D2C2C82FABC" modified="2019-05-08" version="1">MKRKEQYYLTSNGKSRFLEVSENIFPTLTLVSEFVLFKSYINCMWITELFINVLIINHLQVVLYFVPPIHSASCKVFCRRMHSSKNKNVEKPEPFYRLSAPKQKTSAEIINEARNALRTLRTQRPFTPREDQRKLFGSASSRTLENRPPSTFSLHASSFESGESRPVSGTRLSPLDHKPKLLSSPIKDEDSSISFPKPPVDPMEVRKVSSARARLCRTASQGNLLPDKIFLPDENKEGVNSEEPATMDDLSRRSNKSCLAELSPHSNFKDDGIPLIYNEHNKAEEQDFLKEAVATSPLQFRSGSDQIKNGDRASSNLSRISWNTSGSKAEDNIENLCRACTSLHQALEEGNMFGKRFKRRSALLKLLYKLVDIGSDPLGLKLARMILALKVSGKNLLNVCKLVFKISRNEKNDSLIQNDSLLDSLLEILRFEDLQTNNEAFLYCLGAIKFISGNAVLLNELVNKGTVEILFQLMKQINNIKENDTRFSNLGHLLVQLTATLRNLVDLPLSRCQLLCSGAVSELWVVLEQRMNDKDICANIVRIFSKLSSYNDCCAALAACSRCYVLFLALLNKHQKKQDLVVRVVFILGNLTAKNNQAREQFFKKKGSADTLISLFQTYNEFDLNTKKEKHEGEENKSKQKHPSEAEDVLIKLIRVLANLSIHPNVGAVLAANHHIVALLITVLEYKSVDDCEELVINAAATINNLSYYKVRNSVIEDKKLHIAELLLKLLMSSNMDGILEAVRVFGNLSQYLEICDFIVQKKVYKYMIALLDAKHQDVCFSACGVLLNLTVDKNKRAILKEEGGIKKLVDCLRDFGPTDWQLACLICKTLWNYSENITNAASCFEEDDINTLLVLLALFLDEELALDYSFDRDLRDYHKLYWETEFKPVAQKLLNRIQSHHSFLQPITITSF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IND9</accession>
    <name>A0A452IND9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">SMB domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IND9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033397.1">
        <property type="protein sequence ID" value="ENSGAGP00000029411.1"/>
        <property type="gene ID" value="ENSGAGG00000020186.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009986">
        <property type="term" value="C:cell surface"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004551">
        <property type="term" value="F:dinucleotide phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003676">
        <property type="term" value="F:nucleic acid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004528">
        <property type="term" value="F:phosphodiesterase I activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046034">
        <property type="term" value="P:ATP metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030505">
        <property type="term" value="P:inorganic diphosphate transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045599">
        <property type="term" value="P:negative regulation of fat cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009143">
        <property type="term" value="P:nucleoside triphosphate catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030500">
        <property type="term" value="P:regulation of bone mineralization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd16018">
        <property type="entry name" value="Enpp"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00091">
        <property type="entry name" value="NUC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.410.20:FF:000003">
        <property type="entry name" value="Ectonucleotide pyrophosphatase/phosphodiesterase family member 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.410.20:FF:000001">
        <property type="entry name" value="Ectonucleotide pyrophosphatase/phosphodiesterase family member 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.720.10:FF:000145">
        <property type="entry name" value="Uncharacterized protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.410.20">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.720.10">
        <property type="entry name" value="Alkaline Phosphatase, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.570.10">
        <property type="entry name" value="Extracellular Endonuclease, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017850">
        <property type="entry name" value="Alkaline_phosphatase_core_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044929">
        <property type="entry name" value="DNA/RNA_non-sp_Endonuclease_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001604">
        <property type="entry name" value="Endo_G_ENPP1-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020821">
        <property type="entry name" value="ENPP1-3/EXOG-like_nuc-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044925">
        <property type="entry name" value="His-Me_finger_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002591">
        <property type="entry name" value="Phosphodiest/P_Trfase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036024">
        <property type="entry name" value="Somatomedin_B-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001212">
        <property type="entry name" value="Somatomedin_B_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10151">
        <property type="entry name" value="ECTONUCLEOTIDE PYROPHOSPHATASE/PHOSPHODIESTERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10151:SF77">
        <property type="entry name" value="ECTONUCLEOTIDE PYROPHOSPHATASE_PHOSPHODIESTERASE FAMILY MEMBER 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01223">
        <property type="entry name" value="Endonuclease_NS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01663">
        <property type="entry name" value="Phosphodiest"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01033">
        <property type="entry name" value="Somatomedin_B"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00892">
        <property type="entry name" value="Endonuclease_NS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00477">
        <property type="entry name" value="NUC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00201">
        <property type="entry name" value="SO"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53649">
        <property type="entry name" value="Alkaline phosphatase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54060">
        <property type="entry name" value="His-Me finger endonucleases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90188">
        <property type="entry name" value="Somatomedin B domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00524">
        <property type="entry name" value="SMB_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50958">
        <property type="entry name" value="SMB_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="4" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <feature type="domain" description="SMB" evidence="7">
        <location>
            <begin position="7"/>
            <end position="47"/>
        </location>
    </feature>
    <feature type="domain" description="SMB" evidence="7">
        <location>
            <begin position="48"/>
            <end position="92"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50958"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029411.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="830" mass="95846" checksum="50F6D53F632B8045" modified="2019-05-08" version="1">MTLSNISVKTCKGRCFERTFGSCRCDRDCVELGNCCLDFQETCIQPAHIWTCNKFRCGEKRWPENRCSCSDDCVENKDCCVNYYIVCQGAKSWVEEKCEDINEPQCPPGFAKPPVLLFSLDGFRAEYLHTWSGLLPVISKLQKCGTYTPNMRPVYPTKTFPNHYTIVTGLYPESHGIIDNKIYDPHRNVSFTLKNAEKFNPQWYQGQPIWLTAMHQGLKAGTFFWPGSDVRINGTFPDLYRKYNGYVKHRVCVCVSTETYGNTYVNRVCRKRSFICIWAVLCLTFYIHQIQIYEFVSKCLTSILHHHISILQVILALQRVDNIVGMLMDGLKQMNLHKCLNIILLSDHGMETASCSKTAYLSTYMENVQDITVIPGPAARLRPQNCIDPDQHFKAFMKELLPKRFHYVNNDRIERVHFYLDPQWQLARKPSEIKYCNGGFHGSDNRFMSMQAIFIGFGPGFKFKTQVDPFENIEIYNLICDLLGITPAYNNGTHGSLNHLLKNPVYIPRHPEEESHPSVYPLMKTTSSLDIGCSCDLAVKNLNKYAKYIELVINSVSALVKKIEKYSLPYGRPRVLQKRSTYYLLYHHQYVSGYSQDLNMTLWSAYTVTKYDNRTASVENLSNCLHVDIRIPSSSSKTCSFYKNHHQLSYGFLFPPSKFTSSYEVLLNSNIVPMYHEFKVLWNYFYDVLLPKYTLARNGVNVVSGPVFDYDSNGLYDTPEKVKRLTHDPEVLVPTHYFIVLTSCKNISQTPLQCEGALDALSFIVPHRPDNSESCRHKRDSLWVEERMRFHTARVRDVELLTGLSFYQDRKQPVSDILQLKTYLPTFPME</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INE1</accession>
    <name>A0A452INE1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Homeobox domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 6">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the TALE/IRO homeobox family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452INE1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033367.1">
        <property type="protein sequence ID" value="ENSGAGP00000029384.1"/>
        <property type="gene ID" value="ENSGAGG00000021244.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048468">
        <property type="term" value="P:cell development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030182">
        <property type="term" value="P:neuron differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00086">
        <property type="entry name" value="homeodomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.60:FF:000003">
        <property type="entry name" value="Iroquois-class homeobox protein IRX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.60">
        <property type="entry name" value="Homeodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001356">
        <property type="entry name" value="HD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017970">
        <property type="entry name" value="Homeobox_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009057">
        <property type="entry name" value="Homeodomain-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003893">
        <property type="entry name" value="Iroquois_homeo"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008422">
        <property type="entry name" value="KN_HD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11211">
        <property type="entry name" value="IROQUOIS-CLASS HOMEODOMAIN PROTEIN IRX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11211:SF14">
        <property type="entry name" value="IROQUOIS-CLASS HOMEODOMAIN PROTEIN IRX-3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05920">
        <property type="entry name" value="Homeobox_KN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00389">
        <property type="entry name" value="HOX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00548">
        <property type="entry name" value="IRO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46689">
        <property type="entry name" value="Homeodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00027">
        <property type="entry name" value="HOMEOBOX_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50071">
        <property type="entry name" value="HOMEOBOX_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 6" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="4 6" id="KW-0371">Homeobox</keyword>
    <keyword evidence="5 6" id="KW-0539">Nucleus</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Homeobox" evidence="8">
        <location>
            <begin position="116"/>
            <end position="169"/>
        </location>
    </feature>
    <feature type="DNA-binding region" description="Homeobox" evidence="6">
        <location>
            <begin position="118"/>
            <end position="170"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="171"/>
            <end position="344"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="7">
        <location>
            <begin position="197"/>
            <end position="209"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="210"/>
            <end position="219"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="226"/>
            <end position="236"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="7">
        <location>
            <begin position="237"/>
            <end position="247"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="248"/>
            <end position="260"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="326"/>
            <end position="337"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008446"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023155"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00108"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50071"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029384.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="434" mass="47161" checksum="54D025C35006B866" modified="2019-05-08" version="1">MSFPQLGYQYIRPIYPSERQGSGGARSGTELTPSGTLSNVLSSMYGAPYAAAAAAQGYGAFLPYTAELPIFPQLGAQYELKDSPGVQHPAFPHHHPAFYPYGQYQFGDPSRPKNATRESTSTLKAWLNEHRKNPYPTKGEKIMLAIITKMTLTQVSTWFANARRRLKKENKMTWAPRSRTDEEGNSYGSDHEGEEDKREDEEEIDLENIDTEHMESQKDELEEDPDLLHSDSKTDSEGSEGFEDLPGSEERFLKAMEGGRPHPPAGGLEPLKLPPQQHRLSPPAPASSPTEGSLASAGQKPKIWSLAETATSPDNPRKSPGGGSGCSPPAAAPQSLPLAPPPPHRLVSSCPLGKFPNWTSRAFSAHHHHPHPLTLLSTPHLLGLGATPGAASSAAAIAAFPRPADQAQSTESAGTGPRTNLKQLWFYQRCHHHS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452INE9</accession>
    <name>A0A452INE9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="6">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="1 3">Belongs to the ETS family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452INE9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033621.1">
        <property type="protein sequence ID" value="ENSGAGP00000029608.1"/>
        <property type="gene ID" value="ENSGAGG00000021383.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043565">
        <property type="term" value="F:sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd08542">
        <property type="entry name" value="SAM_PNT-ETS-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.50:FF:000014">
        <property type="entry name" value="Protein c-ets-1 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.50">
        <property type="entry name" value="Transcription Factor, Ets-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.10">
        <property type="entry name" value="Winged helix-like DNA-binding domain superfamily/Winged helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045688">
        <property type="entry name" value="Ets1_N_flank"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000418">
        <property type="entry name" value="Ets_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046328">
        <property type="entry name" value="ETS_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003118">
        <property type="entry name" value="Pointed_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013761">
        <property type="entry name" value="SAM/pointed_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041886">
        <property type="entry name" value="SAM_PNT-ETS-1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036388">
        <property type="entry name" value="WH-like_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036390">
        <property type="entry name" value="WH_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11849">
        <property type="entry name" value="ETS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11849:SF209">
        <property type="entry name" value="ETS TRANSLOCATION VARIANT 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00178">
        <property type="entry name" value="Ets"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19525">
        <property type="entry name" value="Ets1_N_flank"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02198">
        <property type="entry name" value="SAM_PNT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00413">
        <property type="entry name" value="ETS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00251">
        <property type="entry name" value="SAM_PNT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47769">
        <property type="entry name" value="SAM/Pointed domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46785">
        <property type="entry name" value="Winged helix' DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00345">
        <property type="entry name" value="ETS_DOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50061">
        <property type="entry name" value="ETS_DOMAIN_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51433">
        <property type="entry name" value="PNT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 3" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0539">Nucleus</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="PNT" evidence="5">
        <location>
            <begin position="60"/>
            <end position="145"/>
        </location>
    </feature>
    <feature type="domain" description="ETS" evidence="4">
        <location>
            <begin position="344"/>
            <end position="380"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00005562"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="RuleBase" id="RU004019"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50061"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS51433"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029608.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="399" mass="45582" checksum="24D11C32E225068D" modified="2019-05-08" version="1">MSCGFQSTCYQQRICYPVWDESTIQEVPTGIERYNSDMECADVPLLTPSSKEMMSQALKATFSGFTKEQQRLGIPKDPQQWTETHVRDWVMWAVNEFSLKDVDFQKFCMNGAALCALGKDCFLELAPDFVGDILWEHLEILQKDEIKPYSANGVNPAYPESRYTSDYFISYGIEHAQCVPPSEFSEPSFITESYQTLHPISSEELFSLKYENDYPSVILRDPVQTDSLQTDYFTIKQEVVTPDNMCMGRTSRGKLGGQDSFESIESYDSCDRLTQSWNSQSSFQSLQRVPSYDSFDSEDYPAALPNHKPKGTFKDYVRDRANMNKDKPVIPAAALAGYTGSGPIQLWQFLLELLTDKSCQSFISWTGDGWEFKLSDPDEVWPIVTDQRPLKMLRRIPKS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGR3</accession>
    <name>A0A452GGR3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase</fullName>
            <ecNumber evidence="15">2.7.4.24</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Bifunctional inositol kinase that acts in concert with the IP6K kinases IP6K1, IP6K2 and IP6K3 to synthesize the diphosphate group-containing inositol pyrophosphates diphosphoinositol pentakisphosphate, PP-InsP5, and bis-diphosphoinositol tetrakisphosphate, (PP)2-InsP4. PP-InsP5 and (PP)2-InsP4, also respectively called InsP7 and InsP8, regulate a variety of cellular processes, including apoptosis, vesicle trafficking, cytoskeletal dynamics, exocytosis, insulin signaling and neutrophil activation. Phosphorylates inositol hexakisphosphate (InsP6) at position 1 to produce PP-InsP5 which is in turn phosphorylated by IP6Ks to produce (PP)2-InsP4. Alternatively, phosphorylates PP-InsP5 at position 1, produced by IP6Ks from InsP6, to produce (PP)2-InsP4. Activated when cells are exposed to hyperosmotic stress.</text>
    </comment>
    <comment type="function">
        <text evidence="15">Bifunctional inositol kinase that acts in concert with the IP6K kinases to synthesize the diphosphate group-containing inositol pyrophosphates diphosphoinositol pentakisphosphate, PP-InsP5, and bis-diphosphoinositol tetrakisphosphate, (PP)2-InsP4. PP-InsP5 and (PP)2-InsP4, also respectively called InsP7 and InsP8, may regulate a variety of cellular processes, including apoptosis, vesicle trafficking, cytoskeletal dynamics, and exocytosis. Phosphorylates inositol hexakisphosphate (InsP6).</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>1D-myo-inositol hexakisphosphate + ATP = 1-diphospho-1D-myo-inositol 2,3,4,5,6-pentakisphosphate + ADP</text>
            <dbReference type="Rhea" id="RHEA:37459"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:58130"/>
            <dbReference type="ChEBI" id="CHEBI:74946"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.4.24"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="13">
            <dbReference type="Rhea" id="RHEA:37460"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="12">
            <text>5-diphospho-1D-myo-inositol 1,2,3,4,6-pentakisphosphate + ATP + H(+) = 1,5-bis(diphospho)-1D-myo-inositol 2,3,4,6-tetrakisphosphate + ADP</text>
            <dbReference type="Rhea" id="RHEA:10276"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:58628"/>
            <dbReference type="ChEBI" id="CHEBI:77983"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.4.24"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="12">
            <dbReference type="Rhea" id="RHEA:10277"/>
        </physiologicalReaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2 15">Cytoplasm</location>
            <location evidence="2 15">Cytosol</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 15">Belongs to the histidine acid phosphatase family. VIP1 subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.4.24" evidence="15"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000864.1">
        <property type="protein sequence ID" value="ENSGAGP00000000767.1"/>
        <property type="gene ID" value="ENSGAGG00000000614.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033857">
        <property type="term" value="F:5-diphosphoinositol pentakisphosphate 1-kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0052723">
        <property type="term" value="F:inositol hexakisphosphate 1-kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="RHEA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006020">
        <property type="term" value="P:inositol metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032958">
        <property type="term" value="P:inositol phosphate biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07061">
        <property type="entry name" value="HP_HAP_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.470.20:FF:000003">
        <property type="entry name" value="Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.11950:FF:000001">
        <property type="entry name" value="Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.11950:FF:000003">
        <property type="entry name" value="Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.11950">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.470.20">
        <property type="entry name" value="ATP-grasp fold, B domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1240">
        <property type="entry name" value="Phosphoglycerate mutase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033379">
        <property type="entry name" value="Acid_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000560">
        <property type="entry name" value="His_Pase_clade-2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037446">
        <property type="entry name" value="His_Pase_VIP1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029033">
        <property type="entry name" value="His_PPase_superfam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040557">
        <property type="entry name" value="VIP1_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12750">
        <property type="entry name" value="DIPHOSPHOINOSITOL PENTAKISPHOSPHATE KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12750:SF11">
        <property type="entry name" value="INOSITOL HEXAKISPHOSPHATE AND DIPHOSPHOINOSITOL-PENTAKISPHOSPHATE KINASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00328">
        <property type="entry name" value="His_Phos_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18086">
        <property type="entry name" value="PPIP5K2_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53254">
        <property type="entry name" value="Phosphoglycerate mutase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00616">
        <property type="entry name" value="HIS_ACID_PHOSPHAT_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10 15" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="5 15" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9 15" id="KW-0418">Kinase</keyword>
    <keyword evidence="11" id="KW-0472">Membrane</keyword>
    <keyword evidence="8 15" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7 15" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="VIP1 N-terminal" evidence="17">
        <location>
            <begin position="55"/>
            <end position="144"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="910"/>
            <end position="974"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1147"/>
            <end position="1226"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1253"/>
            <end position="1421"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1"/>
            <end position="15"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="16">
        <location>
            <begin position="1161"/>
            <end position="1178"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1191"/>
            <end position="1209"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1383"/>
            <end position="1405"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004514"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005609"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00033696"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00034629"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00037056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="RuleBase" id="RU365032"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF18086"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000767.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1421" mass="159513" checksum="AEEA2700321481EB" modified="2019-05-08" version="1">MSSLTTSHEGASSTPRFFVGTRDDETDFLGSNMKTDEIDFFEDDEESESPPERQIVVGICAMTKKSKSKPMTQILERLCKFEYITVVIMGEDVILNEPVENWPPCDCLISFHSKGFPLDKAVAYAKLCNPFLINDLDMQYYIQDRREVYRILQEEGIDLPRYAVLNRDPDKPDECSLVEGEDQVEVNGAVFPKPFVEKPVSAEDHNVYIYYPTSAGGGSQRLFRKIGSRSSVYSPESCVRKTGSYIYEEFMPTDGTDVKVYTVGPDYAHAEARKSPALDGKVERDSEGKEIRYPVMLTAMEKLVARKVCVAFKQTVCGFDLLRANGHSFVCDVNGFSFVKNSMKYYDDCTKILGNIIMRELAPQFQIPWSIPTEAEDIPIVPTTSGTMMELRCVIAVIRHGDRTPKQKMKMEVKHSRFFELFEKYDGYKTGKLKLKKPEQLQEVLDIARLLVVELGTHNDCEIEERKSKLEQLKTVLEMYGHFSGINRKVQLTYLPRGHPKASSEDEDARKEPSPSLLLVLKWGGELTPAGRVQAEELGRAFRCMYPGGQGDYAGFPGCGLLRLHSTYRHDLKIYASDEGRVQMTAAAFAKGLLALEGELTPILVQMVKSANMNGLLDSDSDSLSSCQHRVKARLHEIMQKDAKFCEEDYEKLAPTGSISLVSSMTFIQNPVEICDQVFALIENLTSQICKRLEDPKSADLQLYHSETLELMLQRWSKLERDFRMKNGRYDISKIPDIYDCIKYDVQHNGTLKLEGTAELFKLSKALADVIIPQEYGINKEEKLEIAIGFCLPLIKKIQLDLQRTHEDESVNKLHPLYSRGVLSPGRHVRTRLYFTSESHVHSLLSIFRYGGLLDESKDQQWKRAMDYLSAISELNYMTQIVIMLYEDNNKDPSSEERFHVELHFSPGVKGCEEDGSAPTGFGFRPASAENDDKKPNQGSMEDLSKVKSIDETDQAQERSPQPSEPICIQRRSPLIRNRKTGSMEVLSESSSKTGGYRLFSSYSRQSTEMKQSGLDVLEHCRVVPLLCRFPSDYAMRQNISLDATLAHHLHQCSYHLRLFRSWLISGQDDLECLYGFEGCSMVPTIYPLETLHNSLSLRQVNEFLTSVCRCYSSLHSRASTALFDSVIASQTPGDPFMPQRILSSSSLPLRQRSDKPPWYSSGPSSTVSSAGPSSPTSMENYARFGFTEKPSISPQISEERLNCQSPEQSEMEPSGEALELDLGGSGSLVENPLGEMPEQGLMRMPGLLEVPEPSKEELSEEVPEQGTMRVLGLSKTRQSGKTSEKDGSVVLEQGGEKPAGGTWELGETGKLEAGEIGLSGKISDSDEEGPAGEMLEPDVVRPVAPGETSELDEEEPVMQTLEPSEKGQPVEILEKEDEELSTDNALSDQQPFSHLYQDQQLSSEKSVEDEQPSSQQELQN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGR9</accession>
    <name>A0A452GGR9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Cullin family profile domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="pathway">
        <text evidence="1">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="similarity">
        <text evidence="2 4 5">Belongs to the cullin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GGR9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000858.1">
        <property type="protein sequence ID" value="ENSGAGP00000000760.1"/>
        <property type="gene ID" value="ENSGAGG00000000219.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031461">
        <property type="term" value="C:cullin-RING ubiquitin ligase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031625">
        <property type="term" value="F:ubiquitin protein ligase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006511">
        <property type="term" value="P:ubiquitin-dependent protein catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1310.10:FF:000012">
        <property type="entry name" value="Cullin 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1310.10:FF:000016">
        <property type="entry name" value="Cullin 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1310.10:FF:000018">
        <property type="entry name" value="Cullin 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.230.130:FF:000003">
        <property type="entry name" value="Cullin 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.280.240">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1310.10">
        <property type="entry name" value="Cullin Repeats"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.230.130">
        <property type="entry name" value="Cullin, Chain C, Domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045093">
        <property type="entry name" value="Cullin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR059120">
        <property type="entry name" value="Cullin-like_AB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016158">
        <property type="entry name" value="Cullin_homology"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036317">
        <property type="entry name" value="Cullin_homology_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001373">
        <property type="entry name" value="Cullin_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016159">
        <property type="entry name" value="Cullin_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11932">
        <property type="entry name" value="CULLIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00888">
        <property type="entry name" value="Cullin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26557">
        <property type="entry name" value="Cullin_AB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00182">
        <property type="entry name" value="CULLIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF75632">
        <property type="entry name" value="Cullin homology domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF74788">
        <property type="entry name" value="Cullin repeat-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50069">
        <property type="entry name" value="CULLIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="Cullin family profile" evidence="6">
        <location>
            <begin position="366"/>
            <end position="598"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006019"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00330"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="RuleBase" id="RU003829"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50069"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000760.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="665" mass="77884" checksum="837FFEF4161B0F56" modified="2019-05-08" version="1">MSLKPRVVDFDETWNKLLTTIKAVVMLDYVERATWNDRFSDIYALCVAYPEPLGERLYTETKFFLENHVRHLHKRVLESEEQVLVMYHRYWEEYSKGADYMDCLYRYLNTQFIKKNKLTEADLQYGYGGVDISEPLMEIGELALDMWRKLMIEPLQGILIRMLLREIKNDRCGEDPNQKVIHGVINSFVHVEQYKKKFPLKFYQEIFECPFLNETGEYYKQEASNLLQESNCSQYMEKVLGRLKDEEMRCRKYLHPSSYGKVIHECQQRMVADHLQFLHAECHNIIRQEKRSDMANMYTLLRAVSSGLPHMIQELQNHIHDEGLRATSNLSQENMPTQFVESVLEVHSKFVQLINTVLNGDQHFMSALDKLAKYCDNLLKKSAKGMTENEVEDKLTSFITVFKYIDDKDVFQKFYARMLAKRLIHGLSMSMDSEEAMINKLKQACGYEFTSKLHRMYTDMSVSADLNNKFNNFIKNQDAIIDLGISFQIYVLQAGAWPLTQAPSSTFAIPQELEKSVQMFELFYSQHFSGRKLTWLHYLCTGEVKMNYLCKPYVAMVTTYQMAVLLAFNNSETVSYKELQDSTQMNEKELTKTIKSLLDVKMINHDLDKEDIEAESTFSLNLNFSSKRTKFKITTSMQKDTPQARVAVLVWICKSDRVLCFRIHP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGS0</accession>
    <name>A0A452GGS0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Alpha-endosulfine</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">Protein phosphatase inhibitor that specifically inhibits protein phosphatase 2A (PP2A) during mitosis.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 10">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 10">Belongs to the endosulfine family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GGS0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000880.1">
        <property type="protein sequence ID" value="ENSGAGP00000000781.1"/>
        <property type="gene ID" value="ENSGAGG00000000656.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004864">
        <property type="term" value="F:protein phosphatase inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006760">
        <property type="entry name" value="Endosulphine"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10358:SF21">
        <property type="entry name" value="ALPHA-ENDOSULFINE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10358">
        <property type="entry name" value="ENDOSULFINE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04667">
        <property type="entry name" value="Endosulfine"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 10" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="5 10" id="KW-0132">Cell division</keyword>
    <keyword evidence="3 10" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6 10" id="KW-0498">Mitosis</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="7 10" id="KW-0650">Protein phosphatase inhibitor</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1"/>
            <end position="52"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="79"/>
            <end position="134"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="1"/>
            <end position="37"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="81"/>
            <end position="92"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010520"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022776"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023272"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00040822"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="RuleBase" id="RU363120"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000781.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="134" mass="14785" checksum="6FD71625047DD092" modified="2019-05-08" version="1">MSEKLSADSRCEQRGDEKQDTQEKEAVVPERAEEAKLKAKYPNLGQKPGGSDFLMKRLQKGQKYFDSGDYNMAKAKMKNKQLPTTGPDNNLVTGDHIPTPQDLPPRKSSLVTSKLAGPENHDSVNAACGMQDHN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGS5</accession>
    <name>A0A452GGS5_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="6">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the DONSON family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GGS5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000888.1">
        <property type="protein sequence ID" value="ENSGAGP00000000788.1"/>
        <property type="gene ID" value="ENSGAGG00000000167.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="project" value="UniProtKB-SubCell"/>
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        <property type="term" value="P:nuclear DNA replication"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="entry name" value="Donson"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12972:SF0">
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    </dbReference>
    <dbReference type="PRINTS" id="PR02064">
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    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="3" id="KW-0539">Nucleus</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="205"/>
            <end position="225"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="205"/>
            <end position="216"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00025806"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000788.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="431" mass="48367" checksum="38C4BCD4B8C4D41B" modified="2019-05-08" version="1">ERADFDLHLPVSIPDVSSSPSTEFPADWSIKTRLLFMSSQPFTWAEHLKAQEEAQGFAQHCRATSVNFPQSIQEPKLSTELRCAFQQSLIYWLHPSLPWLQLFPRIGADRKTAGKTSLWSCDETLQQVLMSEWSVSFTSLYNLLKAKLCPYFYVCTYQFTVLFRTAGLAGSDVITAVVSPTTRGLREAMRNEGIEFSLPLVEETRINSPETDKNTEDGEEQGVSDDESFSWLEEMGVQDEIKKPDAISVKLCKEKNEVQVDHKPESVALVKGTNTFMLLNFLINCKSLVAAAGPQAGLPPTLLSPVAFRGATMQTLKARSINVKTPVHSCYNDIFSLEITGPVMPHCLHTLTMLLKSAQRGAFSAVLYTHEPTAVFNTKIERILNKETMCKDLTKCGLHLKTLDQLIQLPTLGKSSIRLLEMRDYAYTWKS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGS6</accession>
    <name>A0A452GGS6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Phosphatase and actin regulator 4</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">Regulator of protein phosphatase 1 (PP1) required for neural tube and optic fissure closure, and enteric neural crest cell (ENCCs) migration during development. Acts as an activator of PP1. During neural tube closure, localizes to the ventral neural tube and activates PP1, leading to down-regulate cell proliferation within cranial neural tissue and the neural retina. Also acts as a regulator of migration of enteric neural crest cells (ENCCs) by activating PP1, leading to repression of the integrin signaling through the rho/rock pathway.</text>
    </comment>
    <comment type="subunit">
        <text evidence="2 10">Binds PPP1CA and actin.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="10">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="10">Cell projection</location>
            <location evidence="10">Lamellipodium</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="1 10">Belongs to the phosphatase and actin regulator family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GGS6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000885.1">
        <property type="protein sequence ID" value="ENSGAGP00000000786.1"/>
        <property type="gene ID" value="ENSGAGG00000000652.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030027">
        <property type="term" value="C:lamellipodium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003779">
        <property type="term" value="F:actin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008157">
        <property type="term" value="F:protein phosphatase 1 binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072542">
        <property type="term" value="F:protein phosphatase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030036">
        <property type="term" value="P:actin cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061386">
        <property type="term" value="P:closure of optic fissure"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048484">
        <property type="term" value="P:enteric nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:2001045">
        <property type="term" value="P:negative regulation of integrin-mediated signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001755">
        <property type="term" value="P:neural crest cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001843">
        <property type="term" value="P:neural tube closure"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051726">
        <property type="term" value="P:regulation of cell cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007266">
        <property type="term" value="P:Rho protein signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.1750">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.2130">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004018">
        <property type="entry name" value="RPEL_repeat"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12751:SF4">
        <property type="entry name" value="PHOSPHATASE AND ACTIN REGULATOR 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12751">
        <property type="entry name" value="PHOSPHATASE AND ACTIN REGULATOR PHACTR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02755">
        <property type="entry name" value="RPEL"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00707">
        <property type="entry name" value="RPEL"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51073">
        <property type="entry name" value="RPEL"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7 10" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="8 10" id="KW-0966">Cell projection</keyword>
    <keyword evidence="4 10" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="3 10" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="6 10" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <feature type="repeat" description="RPEL" evidence="9">
        <location>
            <begin position="49"/>
            <end position="74"/>
        </location>
    </feature>
    <feature type="repeat" description="RPEL" evidence="9">
        <location>
            <begin position="574"/>
            <end position="599"/>
        </location>
    </feature>
    <feature type="repeat" description="RPEL" evidence="9">
        <location>
            <begin position="612"/>
            <end position="637"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1"/>
            <end position="24"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="66"/>
            <end position="365"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="449"/>
            <end position="564"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="132"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="179"/>
            <end position="201"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="217"/>
            <end position="235"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="11">
        <location>
            <begin position="236"/>
            <end position="245"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="11">
        <location>
            <begin position="318"/>
            <end position="339"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="11">
        <location>
            <begin position="451"/>
            <end position="461"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="11">
        <location>
            <begin position="491"/>
            <end position="506"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="551"/>
            <end position="564"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00009795"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00011844"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00401"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="RuleBase" id="RU367131"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000786.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="693" mass="76490" checksum="BC8481DE2329D48C" modified="2019-05-08" version="1">MGMGADVLESGDTTPPTKRKSKFSGFGKIFKPWKWRKKKSSDKFKETSEVLERKISMRKPREELVKRGVLLEDPEQDGEEPDKFSHAALKNGHTVPIGGSGMSNLAHLEEEVGKKSSLRKSVPVEEPKKRPGSSSSNPSSEMEPPHEPHAPKQPLLPPKRPSSSSQEASDVQARDPVPASRTTRTASSSTVTAATSIAKTVNSTVAPCPAPRTVPPAFASTNTTATTSTTSVVPAKQPPIPPPKPVNRNSNPLIAELSQAVNSGMGSSKPSPPLPPKRGIPSNVAPSLESAAASKPPSDRIASATRPASIPVHITVSHPPPSPSPPLPSHVPPEPPRTPLPASSPAMEPQISLELPKETPPQEDLRSLEVPKRAAEQGFGEPHVPTWLPQIPLHIRIQQALTSPLPVTPPAEGSHRAHSLLFENDSLYEDNSTLGRARSLPVTIEMLKVPDDDEEEEEADQEGERSSGPHVYIGDAPSVTVIPRLMPQPLQEEEEEEEEGMSDSDSEGPILYKDEEDEEEDESHNSALANKVKRKDTLAMKLGTTATQQELEEKNAFPRKSKEEWNEIRQQIGTALIRRLSQRPTAEELEQRNILQQKNEADRQAEKREIKRRLTRKLSQRPTVAELQARKILRFHEYVEVTDAQDYDRRADKPWTKLTPADKAAIRKELNEFKSSEMQVHEDSKHFTRFHRP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGS7</accession>
    <name>A0A452GGS7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Phosphatase and actin regulator 4</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">Regulator of protein phosphatase 1 (PP1) required for neural tube and optic fissure closure, and enteric neural crest cell (ENCCs) migration during development. Acts as an activator of PP1. During neural tube closure, localizes to the ventral neural tube and activates PP1, leading to down-regulate cell proliferation within cranial neural tissue and the neural retina. Also acts as a regulator of migration of enteric neural crest cells (ENCCs) by activating PP1, leading to repression of the integrin signaling through the rho/rock pathway.</text>
    </comment>
    <comment type="subunit">
        <text evidence="2 10">Binds PPP1CA and actin.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="10">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="10">Cell projection</location>
            <location evidence="10">Lamellipodium</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="1 10">Belongs to the phosphatase and actin regulator family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GGS7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000881.1">
        <property type="protein sequence ID" value="ENSGAGP00000000782.1"/>
        <property type="gene ID" value="ENSGAGG00000000652.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030027">
        <property type="term" value="C:lamellipodium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003779">
        <property type="term" value="F:actin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008157">
        <property type="term" value="F:protein phosphatase 1 binding"/>
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        <property type="project" value="UniProtKB-UniRule"/>
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    <dbReference type="GO" id="GO:0072542">
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        <property type="project" value="UniProtKB-UniRule"/>
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        <property type="term" value="P:actin cytoskeleton organization"/>
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        <property type="project" value="UniProtKB-UniRule"/>
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    <dbReference type="GO" id="GO:0061386">
        <property type="term" value="P:closure of optic fissure"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048484">
        <property type="term" value="P:enteric nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:2001045">
        <property type="term" value="P:negative regulation of integrin-mediated signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001755">
        <property type="term" value="P:neural crest cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001843">
        <property type="term" value="P:neural tube closure"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051726">
        <property type="term" value="P:regulation of cell cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007266">
        <property type="term" value="P:Rho protein signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.1750">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.2130">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004018">
        <property type="entry name" value="RPEL_repeat"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12751:SF4">
        <property type="entry name" value="PHOSPHATASE AND ACTIN REGULATOR 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12751">
        <property type="entry name" value="PHOSPHATASE AND ACTIN REGULATOR PHACTR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02755">
        <property type="entry name" value="RPEL"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00707">
        <property type="entry name" value="RPEL"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51073">
        <property type="entry name" value="RPEL"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7 10" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="8 10" id="KW-0966">Cell projection</keyword>
    <keyword evidence="4 10" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="3 10" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="6 10" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <feature type="repeat" description="RPEL" evidence="9">
        <location>
            <begin position="562"/>
            <end position="587"/>
        </location>
    </feature>
    <feature type="repeat" description="RPEL" evidence="9">
        <location>
            <begin position="600"/>
            <end position="625"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1"/>
            <end position="353"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="437"/>
            <end position="552"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="580"/>
            <end position="609"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="11">
        <location>
            <begin position="46"/>
            <end position="55"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="56"/>
            <end position="73"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="120"/>
            <end position="130"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="167"/>
            <end position="189"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="205"/>
            <end position="223"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="11">
        <location>
            <begin position="224"/>
            <end position="233"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="11">
        <location>
            <begin position="306"/>
            <end position="327"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="11">
        <location>
            <begin position="439"/>
            <end position="449"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="11">
        <location>
            <begin position="479"/>
            <end position="494"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="539"/>
            <end position="552"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="587"/>
            <end position="597"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="11">
        <location>
            <begin position="598"/>
            <end position="607"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00009795"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00011844"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00401"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="RuleBase" id="RU367131"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000782.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="681" mass="74941" checksum="26ED526D2A120B49" modified="2019-05-08" version="1">MEENTAEEVDHPPSDMGMGADVLESGDTTPPTKRKSKFSGFGKIFKPWKWRKKKSSDKFKETSEDGEEPDKFSHAALKNGHTVPIGGSGMSNLAHLEEEVGKKSSLRKSVPVEEPKKRPGSSSSNPSSEMEPPHEPHAPKQPLLPPKRPSSSSQEASDVQARDPVPASRTTRTASSSTVTAATSIAKTVNSTVAPCPAPRTVPPAFASTNTTATTSTTSVVPAKQPPIPPPKPVNRNSNPLIAELSQAVNSGMGSSKPSPPLPPKRGIPSNVAPSLESAAASKPPSDRIASATRPASIPVHITVSHPPPSPSPPLPSHVPPEPPRTPLPASSPAMEPQISLELPKETPPQEDLRSLEVPKRAAEQGFGEPHVPTWLPQIPLHIRIQQALTSPLPVTPPAEGSHRAHSLLFENDSLYEDNSTLGRARSLPVTIEMLKVPDDDEEEEEADQEGERSSGPHVYIGDAPSVTVIPRLMPQPLQEEEEEEEEGMSDSDSEGPILYKDEEDEEEDESHNSALANKVKRKDTLAMKLGTTATQQELEEKNAFPRKSKEEWNEIRQQIGTALIRRLSQRPTAEELEQRNILQQKNEADRQAEKREIKRRLTRKLSQRPTVAELQARKILRFHEYVEVTDAQDYDRRADKPWTKLTPADKAAIRKELNEFKSSEMQVHEDSKHFTRFHRP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGS8</accession>
    <name>A0A452GGS8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Serine/threonine-protein kinase</fullName>
            <ecNumber evidence="20">2.7.11.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="26 27">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="27" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="26" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="26" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="20">Proto-oncogene with serine/threonine kinase activity involved in cell survival and cell proliferation.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subunit">
        <text evidence="17">Interacts with RP9. Interacts with HSP90AA1, this interaction stabilizes PIM1 protein levels. Interacts (ubiquitinated form) with HSP70 and promotes its proteasomal degradation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4 20">Belongs to the protein kinase superfamily. CAMK Ser/Thr protein kinase family. PIM subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="20"/>
    <dbReference type="AlphaFoldDB" id="A0A452GGS8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000892.1">
        <property type="protein sequence ID" value="ENSGAGP00000000791.1"/>
        <property type="gene ID" value="ENSGAGG00000000395.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0106310">
        <property type="term" value="F:protein serine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006915">
        <property type="term" value="P:apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043066">
        <property type="term" value="P:negative regulation of apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000209">
        <property type="entry name" value="Serine/threonine-protein kinase pim-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000232">
        <property type="entry name" value="Serine/threonine-protein kinase pim-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017348">
        <property type="entry name" value="PIM1/2/3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051138">
        <property type="entry name" value="PIM_Ser/Thr_kinase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22984">
        <property type="entry name" value="SERINE/THREONINE-PROTEIN KINASE PIM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22984:SF29">
        <property type="entry name" value="SERINE_THREONINE-PROTEIN KINASE PIM-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF037993">
        <property type="entry name" value="STPK_Pim-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0053">Apoptosis</keyword>
    <keyword evidence="12 20 22" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="16" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11 20" id="KW-0418">Kinase</keyword>
    <keyword evidence="13" id="KW-0460">Magnesium</keyword>
    <keyword evidence="10 20 23" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="15" id="KW-0539">Nucleus</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="27" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6 20 24" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="8 20" id="KW-0808">Transferase</keyword>
    <keyword evidence="14" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="Protein kinase" evidence="25">
        <location>
            <begin position="38"/>
            <end position="290"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="21">
        <location>
            <position position="167"/>
        </location>
    </feature>
    <feature type="binding site" evidence="22">
        <location>
            <begin position="44"/>
            <end position="52"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="22 23">
        <location>
            <position position="67"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="22">
        <location>
            <position position="121"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="22">
        <location>
            <position position="128"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00005505"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022703"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00047040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00047899"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00048679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="PIRNR" id="PIRNR037993"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="PIRSR" id="PIRSR037993-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="PIRSR" id="PIRSR037993-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="RuleBase" id="RU000304"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000791.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="313" mass="35317" checksum="FF3ECDF44231CEFD" modified="2019-05-08" version="1">MLLSKINSLAHLCSGSGGELHPAKLQPGKEKEPLEKLYQVGPLLGSGGFGSVYSGTRLSDGAPVAIKHVARDRISDWGELPGGSRVPMEILLLKKVGSGFRGVIRLLDWFERPDSFVLVLERPDPVQDLFDFITERGALPEELARSFFRQVLEAVRHCHSCGVLHRDIKDENILIDLNSGELKLIDFGSGALLKDTVYTDFDGTRVYSPPEWIRYHRYHGRSAAVWSLGVLLYDMVCGDIPFEHDEEIIRGQVYFRQRVSPECQHLIKWCLALRPSDRPSFEDLLNHSWMQDVLLPQKTAEIHLHSLIQESGK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGT6</accession>
    <name>A0A452GGT6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Alpha-endosulfine</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">Protein phosphatase inhibitor that specifically inhibits protein phosphatase 2A (PP2A) during mitosis.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 10">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 10">Belongs to the endosulfine family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GGT6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000872.1">
        <property type="protein sequence ID" value="ENSGAGP00000000774.1"/>
        <property type="gene ID" value="ENSGAGG00000000656.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004864">
        <property type="term" value="F:protein phosphatase inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006760">
        <property type="entry name" value="Endosulphine"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10358:SF21">
        <property type="entry name" value="ALPHA-ENDOSULFINE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10358">
        <property type="entry name" value="ENDOSULFINE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04667">
        <property type="entry name" value="Endosulfine"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 10" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="5 10" id="KW-0132">Cell division</keyword>
    <keyword evidence="3 10" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6 10" id="KW-0498">Mitosis</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="7 10" id="KW-0650">Protein phosphatase inhibitor</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1"/>
            <end position="52"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="79"/>
            <end position="108"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="1"/>
            <end position="37"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="81"/>
            <end position="92"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010520"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022776"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023272"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00040822"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="RuleBase" id="RU363120"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000774.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="136" mass="15180" checksum="FB75A4064A95D2CD" modified="2019-05-08" version="1">MSEKLSADSRCEQRGDEKQDTQEKEAVVPERAEEAKLKAKYPNLGQKPGGSDFLMKRLQKGQKYFDSGDYNMAKAKMKNKQLPTTGPDNNLVTGDHIPTPQDLPPRKSSLVTSKLAGCIWPALFPPIDLPCNQEDY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGT8</accession>
    <name>A0A452GGT8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase</fullName>
            <ecNumber evidence="15">2.7.4.24</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Bifunctional inositol kinase that acts in concert with the IP6K kinases IP6K1, IP6K2 and IP6K3 to synthesize the diphosphate group-containing inositol pyrophosphates diphosphoinositol pentakisphosphate, PP-InsP5, and bis-diphosphoinositol tetrakisphosphate, (PP)2-InsP4. PP-InsP5 and (PP)2-InsP4, also respectively called InsP7 and InsP8, regulate a variety of cellular processes, including apoptosis, vesicle trafficking, cytoskeletal dynamics, exocytosis, insulin signaling and neutrophil activation. Phosphorylates inositol hexakisphosphate (InsP6) at position 1 to produce PP-InsP5 which is in turn phosphorylated by IP6Ks to produce (PP)2-InsP4. Alternatively, phosphorylates PP-InsP5 at position 1, produced by IP6Ks from InsP6, to produce (PP)2-InsP4. Activated when cells are exposed to hyperosmotic stress.</text>
    </comment>
    <comment type="function">
        <text evidence="15">Bifunctional inositol kinase that acts in concert with the IP6K kinases to synthesize the diphosphate group-containing inositol pyrophosphates diphosphoinositol pentakisphosphate, PP-InsP5, and bis-diphosphoinositol tetrakisphosphate, (PP)2-InsP4. PP-InsP5 and (PP)2-InsP4, also respectively called InsP7 and InsP8, may regulate a variety of cellular processes, including apoptosis, vesicle trafficking, cytoskeletal dynamics, and exocytosis. Phosphorylates inositol hexakisphosphate (InsP6).</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>1D-myo-inositol hexakisphosphate + ATP = 1-diphospho-1D-myo-inositol 2,3,4,5,6-pentakisphosphate + ADP</text>
            <dbReference type="Rhea" id="RHEA:37459"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:58130"/>
            <dbReference type="ChEBI" id="CHEBI:74946"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.4.24"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="13">
            <dbReference type="Rhea" id="RHEA:37460"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="12">
            <text>5-diphospho-1D-myo-inositol 1,2,3,4,6-pentakisphosphate + ATP + H(+) = 1,5-bis(diphospho)-1D-myo-inositol 2,3,4,6-tetrakisphosphate + ADP</text>
            <dbReference type="Rhea" id="RHEA:10276"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:58628"/>
            <dbReference type="ChEBI" id="CHEBI:77983"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.4.24"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="12">
            <dbReference type="Rhea" id="RHEA:10277"/>
        </physiologicalReaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2 15">Cytoplasm</location>
            <location evidence="2 15">Cytosol</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 15">Belongs to the histidine acid phosphatase family. VIP1 subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.4.24" evidence="15"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000907.1">
        <property type="protein sequence ID" value="ENSGAGP00000000805.1"/>
        <property type="gene ID" value="ENSGAGG00000000614.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033857">
        <property type="term" value="F:5-diphosphoinositol pentakisphosphate 1-kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0052723">
        <property type="term" value="F:inositol hexakisphosphate 1-kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="RHEA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006020">
        <property type="term" value="P:inositol metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032958">
        <property type="term" value="P:inositol phosphate biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07061">
        <property type="entry name" value="HP_HAP_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.470.20:FF:000003">
        <property type="entry name" value="Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.11950:FF:000001">
        <property type="entry name" value="Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.11950:FF:000003">
        <property type="entry name" value="Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1240:FF:000013">
        <property type="entry name" value="Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.11950">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.470.20">
        <property type="entry name" value="ATP-grasp fold, B domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1240">
        <property type="entry name" value="Phosphoglycerate mutase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033379">
        <property type="entry name" value="Acid_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000560">
        <property type="entry name" value="His_Pase_clade-2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037446">
        <property type="entry name" value="His_Pase_VIP1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029033">
        <property type="entry name" value="His_PPase_superfam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040557">
        <property type="entry name" value="VIP1_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12750">
        <property type="entry name" value="DIPHOSPHOINOSITOL PENTAKISPHOSPHATE KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12750:SF11">
        <property type="entry name" value="INOSITOL HEXAKISPHOSPHATE AND DIPHOSPHOINOSITOL-PENTAKISPHOSPHATE KINASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00328">
        <property type="entry name" value="His_Phos_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18086">
        <property type="entry name" value="PPIP5K2_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53254">
        <property type="entry name" value="Phosphoglycerate mutase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00616">
        <property type="entry name" value="HIS_ACID_PHOSPHAT_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10 15" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="5 15" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9 15" id="KW-0418">Kinase</keyword>
    <keyword evidence="11" id="KW-0472">Membrane</keyword>
    <keyword evidence="8 15" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7 15" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="VIP1 N-terminal" evidence="17">
        <location>
            <begin position="55"/>
            <end position="144"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="910"/>
            <end position="974"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1108"/>
            <end position="1187"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1214"/>
            <end position="1382"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1"/>
            <end position="15"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="16">
        <location>
            <begin position="1122"/>
            <end position="1139"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1152"/>
            <end position="1170"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1344"/>
            <end position="1366"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004514"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005609"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00033696"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00034629"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00037056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="RuleBase" id="RU365032"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF18086"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000805.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1382" mass="154799" checksum="9C02E9AD3CE6E939" modified="2019-05-08" version="1">MSSLTTSHEGASSTPRFFVGTRDDETDFLGSNMKTDEIDFFEDDEESESPPERQIVVGICAMTKKSKSKPMTQILERLCKFEYITVVIMGEDVILNEPVENWPPCDCLISFHSKGFPLDKAVAYAKLCNPFLINDLDMQYYIQDRREVYRILQEEGIDLPRYAVLNRDPDKPDECSLVEGEDQVEVNGAVFPKPFVEKPVSAEDHNVYIYYPTSAGGGSQRLFRKIGSRSSVYSPESCVRKTGSYIYEEFMPTDGTDVKVYTVGPDYAHAEARKSPALDGKVERDSEGKEIRYPVMLTAMEKLVARKVCVAFKQTVCGFDLLRANGHSFVCDVNGFSFVKNSMKYYDDCTKILGNIIMRELAPQFQIPWSIPTEAEDIPIVPTTSGTMMELRCVIAVIRHGDRTPKQKMKMEVKHSRFFELFEKYDGYKTGKLKLKKPEQLQEVLDIARLLVVELGTHNDCEIEERKSKLEQLKTVLEMYGHFSGINRKVQLTYLPRGHPKASSEDEDARKEPSPSLLLVLKWGGELTPAGRVQAEELGRAFRCMYPGGQGDYAGFPGCGLLRLHSTYRHDLKIYASDEGRVQMTAAAFAKGLLALEGELTPILVQMVKSANMNGLLDSDSDSLSSCQHRVKARLHEIMQKDAKFCEEDYEKLAPTGSISLVSSMTFIQNPVEICDQVFALIENLTSQICKRLEDPKSADLQLYHSETLELMLQRWSKLERDFRMKNGRYDISKIPDIYDCIKYDVQHNGTLKLEGTAELFKLSKALADVIIPQEYGINKEEKLEIAIGFCLPLIKKIQLDLQRTHEDESVNKLHPLYSRGVLSPGRHVRTRLYFTSESHVHSLLSIFRYGGLLDESKDQQWKRAMDYLSAISELNYMTQIVIMLYEDNNKDPSSEERFHVELHFSPGVKGCEEDGSAPTGFGFRPASAENDDKKPNQGSMEDLSKVKSIDETDQAQERSPQPSEPICIQRRSPLIRNRKTGSMEVLSESSSKTGGYRLFSSYSRQSTEMKQSGLELLSMPAVKRFSVSFAKYPTNGFEGCSMVPTIYPLETLHNSLSLRQVNEFLTSVCRCYSSLHSRASTALFDSVIASQTPGDPFMPQRILSSSSLPLRQRSDKPPWYSSGPSSTVSSAGPSSPTSMENYARFGFTEKPSISPQISEERLNCQSPEQSEMEPSGEALELDLGGSGSLVENPLGEMPEQGLMRMPGLLEVPEPSKEELSEEVPEQGTMRVLGLSKTRQSGKTSEKDGSVVLEQGGEKPAGGTWELGETGKLEAGEIGLSGKISDSDEEGPAGEMLEPDVVRPVAPGETSELDEEEPVMQTLEPSEKGQPVEILEKEDEELSTDNALSDQQPFSHLYQDQQLSSEKSVEDEQPSSQQELQN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGT9</accession>
    <name>A0A452GGT9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Cullin family profile domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="pathway">
        <text evidence="1">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="similarity">
        <text evidence="2 5 6">Belongs to the cullin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GGT9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000879.1">
        <property type="protein sequence ID" value="ENSGAGP00000000780.1"/>
        <property type="gene ID" value="ENSGAGG00000000219.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031461">
        <property type="term" value="C:cullin-RING ubiquitin ligase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031625">
        <property type="term" value="F:ubiquitin protein ligase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006511">
        <property type="term" value="P:ubiquitin-dependent protein catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1310.10:FF:000012">
        <property type="entry name" value="Cullin 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1310.10:FF:000016">
        <property type="entry name" value="Cullin 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1310.10:FF:000018">
        <property type="entry name" value="Cullin 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.230.130:FF:000003">
        <property type="entry name" value="Cullin 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1310.10:FF:000022">
        <property type="entry name" value="Cullin-2 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1310.10">
        <property type="entry name" value="Cullin Repeats"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.230.130">
        <property type="entry name" value="Cullin, Chain C, Domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045093">
        <property type="entry name" value="Cullin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR059120">
        <property type="entry name" value="Cullin-like_AB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016158">
        <property type="entry name" value="Cullin_homology"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036317">
        <property type="entry name" value="Cullin_homology_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001373">
        <property type="entry name" value="Cullin_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016159">
        <property type="entry name" value="Cullin_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11932">
        <property type="entry name" value="CULLIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00888">
        <property type="entry name" value="Cullin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26557">
        <property type="entry name" value="Cullin_AB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00182">
        <property type="entry name" value="CULLIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF75632">
        <property type="entry name" value="Cullin homology domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF74788">
        <property type="entry name" value="Cullin repeat-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50069">
        <property type="entry name" value="CULLIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0832">Ubl conjugation</keyword>
    <keyword evidence="3" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="Cullin family profile" evidence="7">
        <location>
            <begin position="386"/>
            <end position="618"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006019"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00330"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="RuleBase" id="RU003829"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50069"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000780.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="677" mass="79158" checksum="F8AF0B2196E1850D" modified="2019-05-08" version="1">MSLKPRVVDFDETWNKLLTTIKAVVMLDYVERATWNDRFSDIYALCVAYPEPLGERLYTETKFFLENHVRHLHKRVLESEEQVLVMYHRYWEEYSKGADYMDCLYRYLNTQFIKKNKLTEADLQYGYGGVDISEPLMEIGELALDMWRKLMIEPLQGILIRMLLREIKNDRCGEDPNQKVIHGVINSFVHVEQYKKKFPLKFYQEIFECPFLNETGEYYKQEASNLLQESNCSQYMEKVLGRLKDEEMRCRKYLHPSSYGKVIHECQQRMVADHLQFLHAECHNIIRQEKRSDMANMYTLLRAVSSGLPHMIQELQNHIHDEGLRATSNLSQENMPTQFVESVLEVHSKFVQLINTVLNGDQHFMSALDKALTSVVNYREPKSICKAPELLAKYCDNLLKKSAKGMTENEVEDKLTSFITVFKYIDDKDVFQKFYARMLAKRLIHGLSMSMDSEEAMINKLKQACGYEFTSKLHRMYTDMSVSADLNNKFNNFIKNQDAIIDLGISFQIYVLQAGAWPLTQAPSSTFAIPQELEKSVQMFELFYSQHFSGRKLTWLHYLCTGEVKMNYLCKPYVAMVTTYQMAVLLAFNNSETVSYKELQDSTQMNEKELTKTIKSLLDVKMINHDLDKEDIEAESTFSLNLNFSSKRTKFKITTSMQKDTPQATGFMDNGRREVRF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGU0</accession>
    <name>A0A452GGU0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">ATP-citrate synthase</fullName>
            <ecNumber evidence="6">2.3.3.8</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="20">ATP-citrate (pro-S-)-lyase</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="21">Citrate cleavage enzyme</fullName>
        </alternativeName>
    </protein>
    <organism evidence="27 28">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="28" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="27" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="27" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="23">Catalyzes the cleavage of citrate into oxaloacetate and acetyl-CoA, the latter serving as common substrate in multiple biochemical reactions in protein, carbohydrate and lipid metabolism.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="22">
            <text>oxaloacetate + acetyl-CoA + ADP + phosphate = citrate + ATP + CoA</text>
            <dbReference type="Rhea" id="RHEA:21160"/>
            <dbReference type="ChEBI" id="CHEBI:16452"/>
            <dbReference type="ChEBI" id="CHEBI:16947"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57288"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.3.3.8"/>
        </reaction>
        <physiologicalReaction direction="right-to-left" evidence="22">
            <dbReference type="Rhea" id="RHEA:21162"/>
        </physiologicalReaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subunit">
        <text evidence="5">Homotetramer.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytosol</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">In the C-terminal section; belongs to the succinate/malate CoA ligase alpha subunit family.</text>
    </comment>
    <comment type="similarity">
        <text evidence="4">In the N-terminal section; belongs to the succinate/malate CoA ligase beta subunit family.</text>
    </comment>
    <dbReference type="EC" id="2.3.3.8" evidence="6"/>
    <dbReference type="AlphaFoldDB" id="A0A452GGU0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000878.1">
        <property type="protein sequence ID" value="ENSGAGP00000000779.1"/>
        <property type="gene ID" value="ENSGAGG00000000607.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003878">
        <property type="term" value="F:ATP citrate synthase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006085">
        <property type="term" value="P:acetyl-CoA biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006101">
        <property type="term" value="P:citrate metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006633">
        <property type="term" value="P:fatty acid biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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    <dbReference type="CDD" id="cd06100">
        <property type="entry name" value="CCL_ACL-C"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="ATP-citrate synthase"/>
        <property type="match status" value="1"/>
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        <property type="match status" value="1"/>
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        <property type="entry name" value="ATP-citrate synthase"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="ATP-citrate synthase subunit"/>
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        <property type="entry name" value="ATP-citrate synthase subunit"/>
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        <property type="entry name" value="Probable ATP-citrate synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.580.10">
        <property type="entry name" value="Citrate Synthase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.230.10">
        <property type="entry name" value="Cytochrome P450-Terp, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="NAD(P)-binding Rossmann-like Domain"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Succinyl-CoA synthetase domains"/>
        <property type="match status" value="2"/>
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        <property type="entry name" value="Citrate-bd"/>
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        <property type="entry name" value="Citrate_synth-like_lrg_a-sub"/>
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        <property type="entry name" value="Citrate_synth-like_sm_a-sub"/>
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        <property type="entry name" value="Citrate_synth_N"/>
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    <dbReference type="InterPro" id="IPR002020">
        <property type="entry name" value="Citrate_synthase"/>
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        <property type="entry name" value="Citrate_synthase_sf"/>
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        <property type="entry name" value="Citrt_syn/SCS-alpha_CS"/>
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        <property type="entry name" value="CoA-bd"/>
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        <property type="entry name" value="ATP-CITRATE SYNTHASE"/>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Glutathione synthetase ATP-binding domain-like"/>
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    </dbReference>
    <dbReference type="SUPFAM" id="SSF52210">
        <property type="entry name" value="Succinyl-CoA synthetase domains"/>
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    </dbReference>
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        <property type="entry name" value="SUCCINYL_COA_LIG_1"/>
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    </dbReference>
    <dbReference type="PROSITE" id="PS00399">
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    <dbReference type="PROSITE" id="PS01217">
        <property type="entry name" value="SUCCINYL_COA_LIG_3"/>
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    </dbReference>
    <proteinExistence type="inferred from homology"/>
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    <keyword evidence="15" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="8" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="10" id="KW-0444">Lipid biosynthesis</keyword>
    <keyword evidence="19" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="16" id="KW-0460">Magnesium</keyword>
    <keyword evidence="13" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="14" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="11" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="28" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="12" id="KW-0808">Transferase</keyword>
    <keyword evidence="17" id="KW-0832">Ubl conjugation</keyword>
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        <location>
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            <end position="601"/>
        </location>
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            <begin position="464"/>
            <end position="485"/>
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            <position position="760"/>
        </location>
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    <evidence type="ECO:0000313" key="28">
        <source>
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    </evidence>
    <sequence length="1045" mass="114294" checksum="CACFC4AF2C5BFFDA" modified="2019-05-08" version="1">MSAKAISEQTGKEFLYKYICTSSAIQNRFKYARVTSDTDWDRLTQDHPWLLTERLVVKPDQLIKRRGKLGLVGVNLDLGQVKAWLKLRLGKETTITKATGILKNFLIEPFVPHKQEEEFYVCIYAAREGDYVLFHHQGGVDVGDVDAKAQKLLVRVDEKLSESDVKKHLLVHALEEKKDVLASFISGLFNLYEDLYFTYLEINPLVVTKDGVYVLDLAAKIDATADYICKVKWGDVEFPPPFGREAYPEEAYIADLDAKSGASLKLTLLNPKGRIWTMVAGGGASVVYSDTICDLGGVNELANYGEYSGAPSEQQTYDYAKTILSLMTREKHPEGKILIIGGSIANFTNVAATFKGIVRAIKDYQGPLQEHGVRIFVRRGGPNYQEGLRVMGEVGKTTGIPIHVFGTETHMTAIVGMALGQRPIPNQPPAAAHTANFLLNASGSCSTPAPSRTASFSECKPDGIAPAKKAKPMTPPDSAPASRPVQGKATTLFSCHTKAIVWGMQTRAVQGMLDFDYICSRDEPSVTAMVYPFTGDHKQKFYWGHKEILIPVFKNMSDAMRKHPEVDVLINFASLRSAYDSTIETMSFPQIRTIAIIAEGIPEALTRKLIKTADKKGVTIIGPATVGGIKPGCFKIGNTGGMLDNILASKLYRPGSVAYVSRSGGMSNELNNIISRATDGVYEGVAIGGDRYPGSTFMDHVLRYQDTPGVKMIVVLGEIGGTEEYKISRGIKEGRITKPVVCWCIGTCATMFSSEVQFGHAGACANQASETAVAKNQALKEAGVFVPQSFDELGDVIQSVYEDLVSKGVIEPAQEVPPPTVPMDYSWARELGLIRKPASFMTSICDERGQELIYAGMPITEVFKEEMGIGGVLGLLWFQRRLPKYACHFIEMCLMVTADHGPAVSGAHNTIVCARAGKDLVSSLTSGLLTIGDRFGGALDAAAKMFSKAFDSGIIPMEFVNKMKKEGKLIMGIGHRVKSINNPDMRVQILKDYVKQHFPATPLLDYALEVEKITTSKKPNLILNVDGFIGVAFVDVLRNCGSFTR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGU1</accession>
    <name>A0A452GGU1_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="5">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000000905.1">
        <property type="protein sequence ID" value="ENSGAGP00000000803.1"/>
        <property type="gene ID" value="ENSGAGG00000000675.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051726">
        <property type="term" value="P:regulation of cell cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048024">
        <property type="term" value="P:regulation of mRNA splicing, via spliceosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd19870">
        <property type="entry name" value="DSRM_SON-like"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="protein SON isoform X1"/>
        <property type="match status" value="1"/>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="dsRBD_dom"/>
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        <property type="entry name" value="G_patch_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032922">
        <property type="entry name" value="SON"/>
    </dbReference>
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        <property type="entry name" value="PROTEIN SON"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46528:SF1">
        <property type="entry name" value="PROTEIN SON"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14709">
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="RSRP"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50137">
        <property type="entry name" value="DS_RBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50174">
        <property type="entry name" value="G_PATCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0694">RNA-binding</keyword>
    <feature type="domain" description="G-patch" evidence="4">
        <location>
            <begin position="4368"/>
            <end position="4414"/>
        </location>
    </feature>
    <feature type="domain" description="DRBM" evidence="3">
        <location>
            <begin position="4433"/>
            <end position="4503"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1"/>
            <end position="63"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="88"/>
            <end position="172"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1163"/>
            <end position="1184"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="2377"/>
            <end position="4117"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="39"/>
            <end position="63"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="2">
        <location>
            <begin position="118"/>
            <end position="148"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="2377"/>
            <end position="2395"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="2">
        <location>
            <begin position="2396"/>
            <end position="2433"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="2448"/>
            <end position="2465"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="2473"/>
            <end position="2495"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="2503"/>
            <end position="2522"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="2">
        <location>
            <begin position="2523"/>
            <end position="2532"/>
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                <person name="Henen B.T."/>
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        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
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                <person name="Kusumi K."/>
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        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
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    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
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        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
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        <scope>IDENTIFICATION</scope>
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    <comment type="function">
        <text evidence="14">Bifunctional inositol kinase that acts in concert with the IP6K kinases IP6K1, IP6K2 and IP6K3 to synthesize the diphosphate group-containing inositol pyrophosphates diphosphoinositol pentakisphosphate, PP-InsP5, and bis-diphosphoinositol tetrakisphosphate, (PP)2-InsP4. PP-InsP5 and (PP)2-InsP4, also respectively called InsP7 and InsP8, regulate a variety of cellular processes, including apoptosis, vesicle trafficking, cytoskeletal dynamics, exocytosis, insulin signaling and neutrophil activation. Phosphorylates inositol hexakisphosphate (InsP6) at position 1 to produce PP-InsP5 which is in turn phosphorylated by IP6Ks to produce (PP)2-InsP4. Alternatively, phosphorylates PP-InsP5 at position 1, produced by IP6Ks from InsP6, to produce (PP)2-InsP4. Activated when cells are exposed to hyperosmotic stress.</text>
    </comment>
    <comment type="function">
        <text evidence="15">Bifunctional inositol kinase that acts in concert with the IP6K kinases to synthesize the diphosphate group-containing inositol pyrophosphates diphosphoinositol pentakisphosphate, PP-InsP5, and bis-diphosphoinositol tetrakisphosphate, (PP)2-InsP4. PP-InsP5 and (PP)2-InsP4, also respectively called InsP7 and InsP8, may regulate a variety of cellular processes, including apoptosis, vesicle trafficking, cytoskeletal dynamics, and exocytosis. Phosphorylates inositol hexakisphosphate (InsP6).</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>1D-myo-inositol hexakisphosphate + ATP = 1-diphospho-1D-myo-inositol 2,3,4,5,6-pentakisphosphate + ADP</text>
            <dbReference type="Rhea" id="RHEA:37459"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:58130"/>
            <dbReference type="ChEBI" id="CHEBI:74946"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.4.24"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="13">
            <dbReference type="Rhea" id="RHEA:37460"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="12">
            <text>5-diphospho-1D-myo-inositol 1,2,3,4,6-pentakisphosphate + ATP + H(+) = 1,5-bis(diphospho)-1D-myo-inositol 2,3,4,6-tetrakisphosphate + ADP</text>
            <dbReference type="Rhea" id="RHEA:10276"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:58628"/>
            <dbReference type="ChEBI" id="CHEBI:77983"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
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        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="12">
            <dbReference type="Rhea" id="RHEA:10277"/>
        </physiologicalReaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2 15">Cytoplasm</location>
            <location evidence="2 15">Cytosol</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 15">Belongs to the histidine acid phosphatase family. VIP1 subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.4.24" evidence="15"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000887.1">
        <property type="protein sequence ID" value="ENSGAGP00000000789.1"/>
        <property type="gene ID" value="ENSGAGG00000000614.1"/>
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</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGV1</accession>
    <name>A0A452GGV1_9SAUR</name>
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        <recommendedName>
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        </recommendedName>
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        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
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        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
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        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
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    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
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        <scope>IDENTIFICATION</scope>
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    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
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</entry>
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        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
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    <accession>A0A452GH33</accession>
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                <person name="Henen B.T."/>
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                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
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        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
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            <authorList>
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            <authorList>
                <consortium name="Ensembl"/>
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        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GH33"/>
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        <property type="gene ID" value="ENSGAGG00000000744.1"/>
    </dbReference>
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    <dbReference type="InterPro" id="IPR029681">
        <property type="entry name" value="CCDC157"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43696">
        <property type="entry name" value="COILED-COIL DOMAIN-CONTAINING PROTEIN 157"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43696:SF9">
        <property type="entry name" value="COILED-COIL DOMAIN-CONTAINING PROTEIN 157"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="194"/>
            <end position="232"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="255"/>
            <end position="277"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="21"/>
            <end position="118"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="143"/>
            <end position="180"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="196"/>
            <end position="216"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="217"/>
            <end position="232"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000905.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="5">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="277" mass="30592" checksum="229EE4C048C451F0" modified="2019-05-08" version="1">PTSLPATWVTSVHIICLPLSVKSLQAKQRALLQQLDCLDQECEELRASLGEAEDDKAKLEHQLKETQAEKQQVQGQLEAQQQLLENLQQEKLSFEQSTSELLRNISELDELLQELKERERLLVSFPDLHIPAETQFECTGNITDDMEKQLQANNIRISVLEEENSRLRTALAKMKEAAQQGVLRVSLTPAIGGEGTSQCKGGQLLNSNRQSAMQAESGQPASSQPKPSSAEAMWKAMSCVSLPVETSAISTYARLKGKGSAARVQARSAHSPRNHQK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GH36</accession>
    <name>A0A452GH36_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">DMAP1-binding domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell projection</location>
            <location evidence="1">Neuron projection</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the DIP2 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000001016.1">
        <property type="protein sequence ID" value="ENSGAGP00000000896.1"/>
        <property type="gene ID" value="ENSGAGG00000000729.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009986">
        <property type="term" value="C:cell surface"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043005">
        <property type="term" value="C:neuron projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd05905">
        <property type="entry name" value="Dip2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.300.30:FF:000003">
        <property type="entry name" value="DIP2 disco-interacting protein 2 homolog A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.300.30:FF:000001">
        <property type="entry name" value="DIP2 disco-interacting protein 2 homolog C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.12780:FF:000004">
        <property type="entry name" value="Disco interacting protein 2 homolog A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.12780:FF:000002">
        <property type="entry name" value="Disco interacting protein 2 homolog B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.300.30">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.12780">
        <property type="entry name" value="N-terminal domain of ligase-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025110">
        <property type="entry name" value="AMP-bd_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045851">
        <property type="entry name" value="AMP-bd_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000873">
        <property type="entry name" value="AMP-dep_synth/lig_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042099">
        <property type="entry name" value="ANL_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037337">
        <property type="entry name" value="Dip2-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010506">
        <property type="entry name" value="DMAP1-bd"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22754">
        <property type="entry name" value="DISCO-INTERACTING PROTEIN 2 DIP2 -RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22754:SF34">
        <property type="entry name" value="DISCO-INTERACTING PROTEIN 2 HOMOLOG A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00501">
        <property type="entry name" value="AMP-binding"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23024">
        <property type="entry name" value="AMP-dom_DIP2-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06464">
        <property type="entry name" value="DMAP_binding"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01137">
        <property type="entry name" value="DMAP_binding"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56801">
        <property type="entry name" value="Acetyl-CoA synthetase-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51912">
        <property type="entry name" value="DMAP1_BIND"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0966">Cell projection</keyword>
    <keyword evidence="4" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="DMAP1-binding" evidence="7">
        <location>
            <begin position="13"/>
            <end position="131"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="67"/>
            <end position="201"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="71"/>
            <end position="89"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="90"/>
            <end position="102"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="131"/>
            <end position="143"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="178"/>
            <end position="189"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004487"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007735"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS51912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000896.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1573" mass="171270" checksum="26A62B3A41AC2E58" modified="2019-05-08" version="1">MAAERGGCSLEAVPVPLPPEVRESLAELELELSEGDITQKGYEKKRAKLLARYIPLIQGVDPSLQAENRIPGSSQATAAVSKQQKSRSTNSRDERFRSDVHTEAVQAALAKYKERKMPMPSKRRSVLVHSSVETYTPPDTSSASEDEGSLRRQGRIPSTPFQSHSNVEPWLNRVIQGSSTSSSASSTSSHPGGKPAAASNTATATAAATVLADLMAHAQLDNHSAPPDVTTGLVEHLHHERPQVASVRGIPRGYNSSILETADGVPVNSRVSSKIQQLLNTLKRPKRPPLKEFFVDDFEELLEVQQPDPNQPKPEGGEMTALKGEPLGVVTNWPPSLLAALQRWGTTQPKTPCLTALDTTGKAVYTLTYGKLWSRSLKLAYTLLNKLTSKNEPLLKPGHRVALVFPNSDPVMFMVAFYGCLLADLVPVPIEVPLTRKDAGSQQIGFLLGSCGVTLALTTDVCQKGLPKAQTGEVVTFKGWPRLVWFVIDGKHLAKPAKDWHPQVRDASADIAYIEYKTSKEGSTMGITISHASMLAHCHALTQACGYSEAETLTNVLDFKRDAGLWHGVLTSVMNRMHVVSIPYALMKVNPLSWIQKVCSYKACVAVVKSRDMHWSLLAQRDQRDVNLSSLRMLIVADGANPWSISSCDAFLNVFQSRGLRPEVICPCASSSEALTVAIRRPPDLGGPPPGKAVLSMNCLSYNVIRVDTEEKLSVLTVQDVGQVMPGANVCVVKADGTPYLCKTDEVGEICVNSGATGTAYYGLLGITKNVFETIPVTAAGVPVSDQPFTRTGLLGFIGPDSLVFVVGKLDGLMVVNIRKHNADDVVATALAVEPMKFVYRGRIAVFSVTVLHDDRIVLVAEQRPDASEEDSFQWMSRVLQAIDSIHQVGVYCLALVPANTLPKAPLGGIHISETKQRFLEGALHPCNVLMCPHTCVTNLPKPRQKQPDVGPASMIVGNLVAGKRIAQASGRDVTQLEDNDQARKFLYLADVLQWRAQTTPDHPLFLLLNSKGTVASTASCIQLHKKAERVAAALMEKGRLNVGDHVALVYPPGVDLIATFYGCLYAGCIPITVRPPHPQNLATTLPTVKMIVEVSKSACILTTQGIMKLLKSKEAATAVDIKTWPTILDTDDIPKKKVANIFRPPSPDMLAYLDFSVSTTGILAGVKMSHAATSALCRSVKLQCELYPSRQIAICLDPYCGLGFALWCLCSVYSGHQSILVPPLELESNVSLWLSAVSQYKVRVTFCSYSVMEMCTKGLGTQTDVLRMKGVNLSCVRTCMVVAEERPRIALTQSFSKLFKDLGLSARSVSTTFGCRVNVAICLQGTAGPDPTTVYVDMRALRHDRVRLVERGSPHSLPLMESGKILPGVKVIIAHTETKGPLGDSHLGEIWVSSPHNATGYYTVYGEEALHADHFSARLSFGDTQTVWARTGYLGFLRRTELTDANGERHDALYVVGSLDETLELRGMRYHPIDIETSVIRAHKSIAECAVFTWTNLLVVVVELEGSEQEALDLVALVTNVVLEEHYLIVGVVVIVDPGVIPINSRGEKQRMHLRDGFLADQLDPIYVAYNM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GH39</accession>
    <name>A0A452GH39_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">VWFA domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GH39"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001039.1">
        <property type="protein sequence ID" value="ENSGAGP00000000917.1"/>
        <property type="gene ID" value="ENSGAGG00000000183.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.410">
        <property type="entry name" value="von Willebrand factor, type A domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011704">
        <property type="entry name" value="ATPase_dyneun-rel_AAA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039891">
        <property type="entry name" value="VWA8"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002035">
        <property type="entry name" value="VWF_A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036465">
        <property type="entry name" value="vWFA_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21610">
        <property type="entry name" value="VON WILLEBRAND FACTOR A DOMAIN-CONTAINING PROTEIN 8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21610:SF9">
        <property type="entry name" value="VON WILLEBRAND FACTOR A DOMAIN-CONTAINING PROTEIN 8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07728">
        <property type="entry name" value="AAA_5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00327">
        <property type="entry name" value="VWA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53300">
        <property type="entry name" value="vWA-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50234">
        <property type="entry name" value="VWFA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="VWFA" evidence="1">
        <location>
            <begin position="933"/>
            <end position="1115"/>
        </location>
    </feature>
    <evidence type="ECO:0000259" key="1">
        <source>
            <dbReference type="PROSITE" id="PS50234"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000917.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1124" mass="125372" checksum="E0FD5A6D9DBFAD6A" modified="2019-05-08" version="1">MCKLQKKINLATRRKTCLGKTSIIVGWSKEKNINKSFADGFFLLDTTVQSLTLQPSVTDGHITYEDSPLVKSVKLGHILVIDEADKAPTNVTCILKTLVESGEMVLSDGRRLVANPANVDGRENVIVIHPDFRMIVLANRPGFPFLGNDFFGTLGDIFSCHAVDNPKPQSELAMLRQYGPDVPEPILQKLVAAFGELRSLADQGIINYPYSTREVVNIVKHLQKFPNEGLANVVRNVFDFDSYNNEMRDTLISTLHKHGIPIGAKPTNIQLAKELPLPDHKFMGYWRINQSGNAQQKLLCPTETHQLNIKGPVYLNVQGFPLERHEARSLSFTEELAFWTLPLNEVNLISDVAVAEGTTENMLYVATCNPVSLYFMNVSSKNGYFVDCYDIFPRTVGSIWQPFVTVAPLGNPLKGQVILHEEQSNVVLLLDTKTGAIHRLLFSSDTQEVPKRASWWSNKEETVRNKFSHKNWLVFYKEEGNHLIVLDVLEGRTHTISLPINLRSVFLVAEDRWLLVESKTNQKFLLTKPAHMVSEESGVCQLHALNEDPVDKGFGLTTGTQCKPVSLIFLSTLWALVATPLFRLPQVTLSLSQSPSEVYSWKRGSSLGSRHASPSDPFYYGRRSKTGAAKQINCVTLLATNQIVRILAPGDVPLKELYPKDITPPQTAGYLEVTDLNSKKLKYIPIPRSVSLSPYTSWLSKISDTDVLLAPLGKVGLVTVDMGGSVRLWETGLDSLQRSLQDWRNMIGQEDGRPVQITIERDSGLEVSSPKHGKTDPDNMPHVGGNTWAGGTGGRDTAGLGGKGGPYRLDAGHKVYQISQAEKDAVPEEVKRAAREMGEKAFKQRLKEIQMSEYDASTYERFSGAVRRQVQSLRIILDNLQAKGKERQWLRHQAIGELDDAKIIDGLTGEKAIYKRRGELEPELGSPQQKPKRLRLVVDVSGSMYRFNGVDGRLERSMEAVCMVMEAFENYEHKFKYDIVGHSGDGFNIALVGSDKVPKNNKQRLEIIKIMHAHAQFCMSGDHTLEGTEHAVREIAKEEADEYFVIVLSDANLERYGIPPARFAQTLTINPQVNAFAIFIGSLGDQADRLQKMLPAGRSFIAMDTKQIPQILQQIFTSTMLSSA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GH41</accession>
    <name>A0A452GH41_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">SRCR domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="12">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GH41"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001037.1">
        <property type="protein sequence ID" value="ENSGAGP00000000915.1"/>
        <property type="gene ID" value="ENSGAGG00000000435.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000012">
        <property type="entry name" value="CD163 molecule like 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000006">
        <property type="entry name" value="neurotrypsin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000002">
        <property type="entry name" value="Scavenger receptor cysteine-rich type 1 protein M130"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000004">
        <property type="entry name" value="Scavenger receptor cysteine-rich type 1 protein M130"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.250.10">
        <property type="entry name" value="SRCR-like domain"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001190">
        <property type="entry name" value="SRCR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036772">
        <property type="entry name" value="SRCR-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19331:SF468">
        <property type="entry name" value="SCAVENGER RECEPTOR CYSTEINE-RICH TYPE 1 PROTEIN M160"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19331">
        <property type="entry name" value="SCAVENGER RECEPTOR DOMAIN-CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00530">
        <property type="entry name" value="SRCR"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00258">
        <property type="entry name" value="SPERACTRCPTR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00202">
        <property type="entry name" value="SR"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56487">
        <property type="entry name" value="SRCR-like"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00420">
        <property type="entry name" value="SRCR_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50287">
        <property type="entry name" value="SRCR_2"/>
        <property type="match status" value="6"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="10 12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9 13" id="KW-0472">Membrane</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="6 14" id="KW-0732">Signal</keyword>
    <keyword evidence="5 13" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 13" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="14">
        <location>
            <begin position="1"/>
            <end position="28"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019527557" description="SRCR domain-containing protein" evidence="14">
        <location>
            <begin position="29"/>
            <end position="741"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="13">
        <location>
            <begin position="705"/>
            <end position="728"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="15">
        <location>
            <begin position="32"/>
            <end position="132"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="15">
        <location>
            <begin position="137"/>
            <end position="217"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="15">
        <location>
            <begin position="217"/>
            <end position="336"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="15">
        <location>
            <begin position="371"/>
            <end position="471"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="15">
        <location>
            <begin position="476"/>
            <end position="576"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="15">
        <location>
            <begin position="592"/>
            <end position="692"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="57"/>
            <end position="121"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="70"/>
            <end position="131"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="101"/>
            <end position="111"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="206"/>
            <end position="216"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="261"/>
            <end position="325"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="274"/>
            <end position="335"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="305"/>
            <end position="315"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="396"/>
            <end position="460"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="409"/>
            <end position="470"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="440"/>
            <end position="450"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="500"/>
            <end position="564"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="544"/>
            <end position="554"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="617"/>
            <end position="681"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="630"/>
            <end position="691"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="661"/>
            <end position="671"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00196"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50287"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000915.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="741" mass="79561" checksum="FD4430AB24FD2016" modified="2019-05-08" version="1">RRGGWETSGRGKCGSLLALTCSVSLFEGAEELRLADGGSPCAGRVEVKHQDQWGTVCSKGWDKADAGVVCKQLGCGAAVSAPRWGHFGPGSGPIWLDEVACVGTESALWHCRNRGWNVHNCYHEWDAGVTCSGHIELHLVGGDNACSGRVEVRHKDNWATICDAHFDLKAASVICNALQCGIAVSVPEGAHFRKGHGPIWTEEFQCVGNESHHVYCPRVSHGSQMCSHANDASVICSRKNFTQSCSGRVEIQVLGAWGTLCDSHWDLPDANVLCHQLYCGFAVSAPGEAHFGKGNGSVWTDTFHCKGNEPHLRHCPVTALGASQCSHDNDASVVCSGNETRLTLCDNSTSETSQAGITEDVGVVCSGSKQIRLVNGTGRCAGRVEIYYNGSWGTVCDDFWDLLDSNVVCKQLGCGRAINATVSGHYGQGSGQIWLDDVNCSGNESDLWACPSRSWGQHNCRHKEDAGVLCSEFTDLRLVSNSYCAGRLEVFYNGTWGSVCANQMSGVTPEIVCKQLNCGDGGQIARDFEYGAGSGPTWLDHVTCSEQHSSLWQCPSEPWNRKSCDNRAEETHISCTGNSETTFSAFPDQEKLRVMGGEDGCSGRVEVWYQGSWGTVCDDSWDMADANVVCKLLGCGSAISALDKAAFGEGTGPIWVEKLNCRGTESSLWDCPAKPWGESNCDHKEDVAVNCSAPSRRLPTDSGRVTVPVIICIILGALLCLVLIILGAQVQSARAQRRGGS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GH45</accession>
    <name>A0A452GH45_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="6">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
            <location evidence="1">Nucleolus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the SLX9 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GH45"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001043.1">
        <property type="protein sequence ID" value="ENSGAGP00000000920.1"/>
        <property type="gene ID" value="ENSGAGG00000000641.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030686">
        <property type="term" value="C:90S preribosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005730">
        <property type="term" value="C:nucleolus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030688">
        <property type="term" value="C:preribosome, small subunit precursor"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000462">
        <property type="term" value="P:maturation of SSU-rRNA from tricistronic rRNA transcript (SSU-rRNA, 5.8S rRNA, LSU-rRNA)"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028160">
        <property type="entry name" value="Slx9-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31109">
        <property type="entry name" value="PROTEIN FAM207A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31109:SF2">
        <property type="entry name" value="RIBOSOME BIOGENESIS PROTEIN SLX9 HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15341">
        <property type="entry name" value="SLX9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0539">Nucleus</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="4">
        <location>
            <begin position="85"/>
            <end position="112"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004604"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00011022"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000920.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="199" mass="22238" checksum="DA802E49648C855A" modified="2019-05-08" version="1">NQEQRRKRAEQHSSSVPPSGGHLGQLRCLSALEPALMADWAFLSSDVFAGMKIDPKTLVKKLDLDSRSVVSAQTGVEEKTLLPKREKMKLRKERWLQKIEGLKLARQKQKAEAKRKATPVVGDMQPLLEALPELSELTTASKARKQPKSLIQGSLAFLREEEVARFHEVIANPVYKANPLAAISEHLCKRLKEEAGNTI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="20" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GH48</accession>
    <name>A0A452GH48_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="1">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="1 2">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="2" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="1" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GH48"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001038.1">
        <property type="protein sequence ID" value="ENSGAGP00000000916.1"/>
        <property type="gene ID" value="ENSGAGG00000000756.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000001048.1">
        <property type="protein sequence ID" value="ENSGAGP00000000925.1"/>
        <property type="gene ID" value="ENSGAGG00000000756.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10285">
        <property type="entry name" value="URIDINE KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000313" key="1">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000925.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="139" mass="16479" checksum="2290B8FADAC74D5D" modified="2019-05-08" version="1">MDAMVSTVHAWIEEPRKFARSHGVNVRSNAKEPVPEEIHILVIEGFLLYNYKPLTDVFNKRYYLTVPYDECKRRRSTRHYPIPDPPGLFDGHVWPMYLKYRKEMETHGIDAVYLDGLKSRDDLYNQVFEEIQNMLLNCS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GH58</accession>
    <name>A0A452GH58_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">SRCR domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="caution">
        <text evidence="5">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GH58"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001062.1">
        <property type="protein sequence ID" value="ENSGAGP00000000937.1"/>
        <property type="gene ID" value="ENSGAGG00000000435.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009897">
        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000012">
        <property type="entry name" value="CD163 molecule like 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000006">
        <property type="entry name" value="neurotrypsin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000002">
        <property type="entry name" value="Scavenger receptor cysteine-rich type 1 protein M130"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.250.10">
        <property type="entry name" value="SRCR-like domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001190">
        <property type="entry name" value="SRCR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036772">
        <property type="entry name" value="SRCR-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19331:SF468">
        <property type="entry name" value="SCAVENGER RECEPTOR CYSTEINE-RICH TYPE 1 PROTEIN M160"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19331">
        <property type="entry name" value="SCAVENGER RECEPTOR DOMAIN-CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00530">
        <property type="entry name" value="SRCR"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00258">
        <property type="entry name" value="SPERACTRCPTR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00202">
        <property type="entry name" value="SR"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56487">
        <property type="entry name" value="SRCR-like"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00420">
        <property type="entry name" value="SRCR_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50287">
        <property type="entry name" value="SRCR_2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3 5" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="4" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <keyword evidence="1" id="KW-0732">Signal</keyword>
    <keyword evidence="7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="470"/>
            <end position="491"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="8">
        <location>
            <begin position="33"/>
            <end position="133"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="8">
        <location>
            <begin position="149"/>
            <end position="249"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="8">
        <location>
            <begin position="251"/>
            <end position="351"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="8">
        <location>
            <begin position="367"/>
            <end position="467"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="500"/>
            <end position="574"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="71"/>
            <end position="132"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="102"/>
            <end position="112"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="174"/>
            <end position="238"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="187"/>
            <end position="248"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="218"/>
            <end position="228"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="275"/>
            <end position="339"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="319"/>
            <end position="329"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="392"/>
            <end position="456"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="405"/>
            <end position="466"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="436"/>
            <end position="446"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00196"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50287"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000937.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="626" mass="67578" checksum="687EE9E11C91E725" modified="2019-05-08" version="1">MCLCKAVKQGQNLSLDVIQCIFYLLSPGLSESLRLLNGESWCDGRVEISLRGVWSRVLDDQWDMNDARVVCRQLQCGVAEKAYNPPKSERGTGPVGLRRVQCAGNETRLTLCDNSTSETSQAGITEDVGVVCSGDSFPNLIDIFWSKQIRLVNGTGRCAGRVEIYYNGSWGTVCDDFWDLLDSNVVCKQLGCGRAINATVSGHYGQGSGQIWLDDVNCSGNESDLWACPSRSWGQHNCRHKEDAGVLCSDLRLVSNSYCAGRLEVFYNGTWGSVCANQMSGVTPEIVCKQLNCGDGGQIARDFEYGAGSGPTWLDHVTCSEQHSSLWQCPSEPWNRKSCDNRAEETHISCTGNSETTFSAFPDQEKLRVMGGEDGCSGRVEVWYQGSWGTVCDDSWDMADANVVCKLLGCGSAISALDKAAFGEGTGPIWVEKLNCRGTESSLWDCPAKPWGESNCDHKEDVAVNCSGEVTVPVIICIILGALLCLVLIILGAQVQSARAQPSSAERAPRNGDTLAPDLGDHSKVGGQPPRNPGPSLVGVTRTLQTGESSRAGRHLRKRNGSKDSDISPIPLHQGSAEVRKVPHDRGTIPPLTCIPTVIHSKNAQSGLKPPAITYLWAGSICSTSF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GH59</accession>
    <name>A0A452GH59_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="9">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Lipid-anchor</topology>
            <orientation evidence="1">Cytoplasmic side</orientation>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000001060.1">
        <property type="protein sequence ID" value="ENSGAGP00000000935.1"/>
        <property type="gene ID" value="ENSGAGG00000000769.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008360">
        <property type="term" value="P:regulation of cell shape"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004965">
        <property type="entry name" value="Paralemmin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10498:SF10">
        <property type="entry name" value="PALM2 AND AKAP2 FUSION-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10498">
        <property type="entry name" value="PARALEMMIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03285">
        <property type="entry name" value="Paralemmin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="4 7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="6" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="5" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="373"/>
            <end position="400"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="592"/>
            <end position="631"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="701"/>
            <end position="749"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="849"/>
            <end position="881"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1036"/>
            <end position="1119"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1190"/>
            <end position="1254"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="1"/>
            <end position="34"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="70"/>
            <end position="107"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="613"/>
            <end position="623"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="721"/>
            <end position="734"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="869"/>
            <end position="879"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="1036"/>
            <end position="1057"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="1058"/>
            <end position="1082"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="1083"/>
            <end position="1104"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="1105"/>
            <end position="1117"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="1230"/>
            <end position="1239"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004342"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000935.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1320" mass="147092" checksum="402F860DD85AB582" modified="2019-05-08" version="1">MAEAELHKERLQAIAEKRKRQTEIEGKRQQLEDQILQLQHFKSKALREKWLLQGIPAGSAEEEEARRRQSEEDELKVKKLEGNIHRLEQEIEKLESEESQISAKEQIILEKLKETEKSFEDFQKSFSHQDGDAVNYIYSQIPELPTLYSRTAEPVPGWDRSSRVTALYAMEINVEKDKLTGETKILSASPIGPEGTHQRGFKVYDDATKVVYEVHSGSTVVENGVHKLSSKDVDELIQKAGQSSVRGGYEMTVSARNVIADGNLSHVKEQMLFKEAKLEMVHKPSKGNSGNPQHQAKISGDEVPEASMDQPVTMIFMGYQNIDDEEETKKVLGYDETIKAELVLIDEDDEKSLREKTVTDVSTMDGNAAELVSGRPVSDTTEPSSSEGKEEILPMEPVPGAKGMAVLPKDGRGQCSCESPETAVTGMKSSRIPENDIRLRKDKDQNCANIPEPVSVLAAKEEKMEIEVPVSEHKNITVVDSTHPVDNQTHFFSPVASHNGLKDRHESLDSEVAKEIRYLDEVLEANCCDSAADNTFNGTCSPEPSAILVDASGPPVPVNSESVPNEKDAIVAGRQASPITEQNETSVMAEKLKPNGHSSDGQKEELCGNLKMPSSPSSSTSSRRSSKDGELTLTTIKKEAKFELRAFQEEKKPSKLFDNDNEKEKYSVRKVRPSEEIIELEKERRELIKSQAVKKNPGIAAKWWNPPQEKTLEEQLDEEHLESHKKYKERKERQQQQNTTPTSPKQVAYSFVPPEPGCVKKEDIVTEQIDFSAARKQFQLMENSGQSNSQIAPKRYGAPKMFSIKPFYKPIGPSQIDRPTVSMTRPASVPLENNDITIVKAQKVSCVSEGHPRNIQTSPTDPVRELSYGDTSKTGQTTKLWPDDSEFMSARAVFMVVKDDDPGILDQFSRSVNVSSPPEELDSGLDELSVRSQDTTVLETLSNDFSMDNISDSGASNETMNPLQENSLADFSLPQTPQMDTPSECRVGVSKSFSDHGFYSPSSTLLDSMLVDDQLDYHAGLLVQNAIQQAIAEQADKTGSKQATDKAERKNSVKGQEEVSTAPCSVKEQSTMFEPPQVSSPVQEKKDTIPKTSRDEDSGLREGKTSQQSSVSPTSQPFIVEENRQEVSYFSKYSEAAELRSTASILATQEPEVTVGPFKLRSRKQRTLSMIEEEIRAAQEREEELKRQRQGLQMAQSPITKSAPPLPTRTVAYKTAPGKIEKIRPPPSPTTEGPLSQSDLLPDEAAGSQRPKNLLQTLMEDYETHKTKRRERMDDSSYTCQLLSSKVTSEVLEATRVNRRKSALALRWEAGIYANQEENE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GH60</accession>
    <name>A0A452GH60_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">SRCR domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="caution">
        <text evidence="5">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GH60"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001023.1">
        <property type="protein sequence ID" value="ENSGAGP00000000903.1"/>
        <property type="gene ID" value="ENSGAGG00000000435.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000012">
        <property type="entry name" value="CD163 molecule like 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000006">
        <property type="entry name" value="neurotrypsin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000002">
        <property type="entry name" value="Scavenger receptor cysteine-rich type 1 protein M130"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.250.10">
        <property type="entry name" value="SRCR-like domain"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001190">
        <property type="entry name" value="SRCR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036772">
        <property type="entry name" value="SRCR-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19331">
        <property type="entry name" value="SCAVENGER RECEPTOR DOMAIN-CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19331:SF484">
        <property type="entry name" value="SRCR DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00530">
        <property type="entry name" value="SRCR"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00258">
        <property type="entry name" value="SPERACTRCPTR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00202">
        <property type="entry name" value="SR"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56487">
        <property type="entry name" value="SRCR-like"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00420">
        <property type="entry name" value="SRCR_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50287">
        <property type="entry name" value="SRCR_2"/>
        <property type="match status" value="6"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3 5" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="4" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <keyword evidence="1" id="KW-0732">Signal</keyword>
    <feature type="domain" description="SRCR" evidence="6">
        <location>
            <begin position="40"/>
            <end position="140"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="6">
        <location>
            <begin position="145"/>
            <end position="245"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="6">
        <location>
            <begin position="247"/>
            <end position="347"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="6">
        <location>
            <begin position="384"/>
            <end position="484"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="6">
        <location>
            <begin position="489"/>
            <end position="589"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="6">
        <location>
            <begin position="603"/>
            <end position="703"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="65"/>
            <end position="129"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="78"/>
            <end position="139"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="109"/>
            <end position="119"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="170"/>
            <end position="234"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="183"/>
            <end position="244"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="214"/>
            <end position="224"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="272"/>
            <end position="336"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="285"/>
            <end position="346"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="316"/>
            <end position="326"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="409"/>
            <end position="473"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="422"/>
            <end position="483"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="453"/>
            <end position="463"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="513"/>
            <end position="577"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="557"/>
            <end position="567"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="628"/>
            <end position="692"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="641"/>
            <end position="702"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="5">
        <location>
            <begin position="672"/>
            <end position="682"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00196"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50287"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000903.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="747" mass="80627" checksum="E8FBC3E880CCC8FA" modified="2019-05-08" version="1">MQTAVETTGGREWNLRATGRRGSLLALTCSVSLFEGAEELRLADGGSPCAGRVEVKHQDQWGTVCSKGWDKADAGVVCKQLGCGAAVSAPRWGHFGPGSGPIWLDEVACVGTESALWHCRNRGWNVHNCYHEWDAGVTCSGHIELHLVGGDNACSGRVEVRHKDNWATICDAHFDLKAASVICNALQCGIAVSVPEGAHFRKGHGPIWTEEFQCVGNESHHVYCPRVSHGSQMCSHANDASVICSRFRLVNGSTACSGRVEIQVLGAWGTLCDSHWDLPDANVLCHQLYCGFAVSAPGEAHFGKGNGSVWTDTFHCKGNEPHLRHCPVTALGASQCSHDNDASVVCSGKCRRMAGLMTPAPQCVILPHTPVSPLGSIPAGSKQIRLVNGTGRCAGRVEIYYNGSWGTVCDDFWDLLDSNVVCKQLGCGRAINATVSGHYGQGSGQIWLDDVNCSGNESDLWACPSRSWGQHNCRHKEDAGVLCSEFTDLRLVSNSYCAGRLEVFYNGTWGSVCANQMSGVTPEIVCKQLNCGDGGQIARDFEYGAGSGPTWLDHVTCSEQHSSLWQCPSEPWNRKSCDNRAEETHISCTVTCEVSAFPDQEKLRVMGGEDGCSGRVEVWYQGSWGTVCDDSWDMADANVVCKLLGCGSAISALDKAAFGEGTGPIWVEKLNCRGTESSLWDCPAKPWGESNCDHKEDVAVNCSGERPLDPFSEAVYEEIDYNLMREKQEVFSHSGLCVSLLSNALSI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GH62</accession>
    <name>A0A452GH62_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="9">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Lipid-anchor</topology>
            <orientation evidence="1">Cytoplasmic side</orientation>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000001067.1">
        <property type="protein sequence ID" value="ENSGAGP00000000942.1"/>
        <property type="gene ID" value="ENSGAGG00000000769.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008360">
        <property type="term" value="P:regulation of cell shape"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004965">
        <property type="entry name" value="Paralemmin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10498:SF10">
        <property type="entry name" value="PALM2 AND AKAP2 FUSION-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10498">
        <property type="entry name" value="PARALEMMIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03285">
        <property type="entry name" value="Paralemmin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="4 7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="6" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="5" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="373"/>
            <end position="400"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="592"/>
            <end position="631"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="701"/>
            <end position="749"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="849"/>
            <end position="881"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1036"/>
            <end position="1120"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1179"/>
            <end position="1287"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="1"/>
            <end position="34"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="70"/>
            <end position="107"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="613"/>
            <end position="623"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="721"/>
            <end position="734"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="869"/>
            <end position="879"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="1036"/>
            <end position="1057"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="1058"/>
            <end position="1082"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="1083"/>
            <end position="1104"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="1105"/>
            <end position="1117"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="1230"/>
            <end position="1239"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004342"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000942.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1307" mass="145651" checksum="DF02B2B1BE082B7C" modified="2019-05-08" version="1">MAEAELHKERLQAIAEKRKRQTEIEGKRQQLEDQILQLQHFKSKALREKWLLQGIPAGSAEEEEARRRQSEEDELKVKKLEGNIHRLEQEIEKLESEESQISAKEQIILEKLKETEKSFEDFQKSFSHQDGDAVNYIYSQIPELPTLYSRTAEPVPGWDRSSRVTALYAMEINVEKDKLTGETKILSASPIGPEGTHQRGFKVYDDATKVVYEVHSGSTVVENGVHKLSSKDVDELIQKAGQSSVRGGYEMTVSARNVIADGNLSHVKEQMLFKEAKLEMVHKPSKGNSGNPQHQAKISGDEVPEASMDQPVTMIFMGYQNIDDEEETKKVLGYDETIKAELVLIDEDDEKSLREKTVTDVSTMDGNAAELVSGRPVSDTTEPSSSEGKEEILPMEPVPGAKGMAVLPKDGRGQCSCESPETAVTGMKSSRIPENDIRLRKDKDQNCANIPEPVSVLAAKEEKMEIEVPVSEHKNITVVDSTHPVDNQTHFFSPVASHNGLKDRHESLDSEVAKEIRYLDEVLEANCCDSAADNTFNGTCSPEPSAILVDASGPPVPVNSESVPNEKDAIVAGRQASPITEQNETSVMAEKLKPNGHSSDGQKEELCGNLKMPSSPSSSTSSRRSSKDGELTLTTIKKEAKFELRAFQEEKKPSKLFDNDNEKEKYSVRKVRPSEEIIELEKERRELIKSQAVKKNPGIAAKWWNPPQEKTLEEQLDEEHLESHKKYKERKERQQQQNTTPTSPKQVAYSFVPPEPGCVKKEDIVTEQIDFSAARKQFQLMENSGQSNSQIAPKRYGAPKMFSIKPFYKPIGPSQIDRPTVSMTRPASVPLENNDITIVKAQKVSCVSEGHPRNIQTSPTDPVRELSYGDTSKTGQTTKLWPDDSEFMSARAVFMVVKDDDPGILDQFSRSVNVSSPPEELDSGLDELSVRSQDTTVLETLSNDFSMDNISDSGASNETMNPLQENSLADFSLPQTPQMDTPSECRVGVSKSFSDHGFYSPSSTLLDSMLVDDQLDYHAGLLVQNAIQQAIAEQADKTGSKQATDKAERKNSVKGQEEVSTAPCSVKEQSTMFEPPQVSSPVQEKKDTIPKTSRDEDSGLREGKTSQQSSVSPTSQPFIVEENRQEVSYFSKYSEAAELRSTASILATQEPEVTVGPFKLRSRKQRTLSMIEEEIRAAQEREEELKRQRQGLQMAQSPITKSAPPLPTRTVAYKTAPGKIEKIRPPPSPTTEGPLSQSDLLPDEAAGSQRPKNLLQTLMEDYETHKTKRRERMDDSSVLEATRVNRRKSALALRWEAGIYANQEENE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GH63</accession>
    <name>A0A452GH63_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Outer dynein arm-docking complex subunit 4</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="8">Tetratricopeptide repeat protein 25</fullName>
        </alternativeName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Cilium axoneme</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GH63"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001029.1">
        <property type="protein sequence ID" value="ENSGAGP00000000909.1"/>
        <property type="gene ID" value="ENSGAGG00000000687.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005930">
        <property type="term" value="C:axoneme"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000274">
        <property type="entry name" value="Tetratricopeptide repeat domain 25"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.10">
        <property type="entry name" value="Tetratricopeptide repeat domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040111">
        <property type="entry name" value="ODAD4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011990">
        <property type="entry name" value="TPR-like_helical_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019734">
        <property type="entry name" value="TPR_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23040">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23040:SF1">
        <property type="entry name" value="OUTER DYNEIN ARM-DOCKING COMPLEX SUBUNIT 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13424">
        <property type="entry name" value="TPR_12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13176">
        <property type="entry name" value="TPR_7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00028">
        <property type="entry name" value="TPR"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48452">
        <property type="entry name" value="TPR-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50005">
        <property type="entry name" value="TPR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0966">Cell projection</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="5" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="4 9" id="KW-0802">TPR repeat</keyword>
    <feature type="repeat" description="TPR" evidence="9">
        <location>
            <begin position="305"/>
            <end position="338"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="456"/>
            <end position="517"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="10">
        <location>
            <begin position="456"/>
            <end position="466"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="467"/>
            <end position="517"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004430"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022803"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00034139"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00034143"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00339"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000909.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="517" mass="59623" checksum="27B3D9965FF6B1F4" modified="2019-05-08" version="1">MCYLKLGDTENSLRDAEASLQNDKTFSKGLYQKAETLYTMGDFEFALVYYHRGHRLRPELQKFRLGIQKAQEAIDNSVGSPSSVKLENKGDLCFLSRQAESKKVKQKLQVKAPKKDQKQQSKVGPIRNEKTERQLLGELYADKAYLEKLLKDADLIKSSTKYGMTVEDLILGGINYLDTRTEFWQQQKPIYARVRDRKLMQQRWLPERKRKPSEVARYILKSMEEIDMLLTSSCPEASCKKAEHVLKTIQSWSEDEIPNKNELIGNLYSCIGNAQIEMGQMGAALQSHKMDLEIARENDLPDAVSRALDNMGRVYARIGKFQQAIDTWEEKIPLAKTSLEKTWLFHEIGRCYLELDEAAEAQDYGEKSLQSAEEEGDVEWQLNASVLVAQAQVKLQDFQSAVMNFEKALEKAKLIHNDAAQQAIIIALDDANKDFIEELRAEQRRERLESFRDYDYDDDYKEGEEAENQKEREENKQKETEKEENERNGRGEEAEGKEETEGEKGEDYAEPHQDLPE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GH69</accession>
    <name>A0A452GH69_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">tRNA N(3)-cytidine methyltransferase</fullName>
            <ecNumber evidence="10">2.1.1.-</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">S-adenosyl-L-methionine-dependent methyltransferase.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="8">
            <text>cytidine(32) in tRNA(Arg)(CCU) + S-adenosyl-L-methionine = N(3)-methylcytidine(32) in tRNA(Arg)(CCU) + S-adenosyl-L-homocysteine + H(+)</text>
            <dbReference type="Rhea" id="RHEA:60912"/>
            <dbReference type="Rhea" id="RHEA-COMP:15710"/>
            <dbReference type="Rhea" id="RHEA-COMP:15712"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57856"/>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
            <dbReference type="ChEBI" id="CHEBI:74894"/>
            <dbReference type="ChEBI" id="CHEBI:82748"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="8">
            <dbReference type="Rhea" id="RHEA:60913"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="9">
            <text>cytidine(32) in tRNA(Thr) + S-adenosyl-L-methionine = N(3)-methylcytidine(32) in tRNA(Thr) + S-adenosyl-L-homocysteine + H(+)</text>
            <dbReference type="Rhea" id="RHEA:50960"/>
            <dbReference type="Rhea" id="RHEA-COMP:12850"/>
            <dbReference type="Rhea" id="RHEA-COMP:12852"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57856"/>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
            <dbReference type="ChEBI" id="CHEBI:74894"/>
            <dbReference type="ChEBI" id="CHEBI:82748"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="9">
            <dbReference type="Rhea" id="RHEA:50961"/>
        </physiologicalReaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 10">Belongs to the methyltransferase superfamily. METL family.</text>
    </comment>
    <dbReference type="EC" id="2.1.1.-" evidence="10"/>
    <dbReference type="AlphaFoldDB" id="A0A452GH69"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001071.1">
        <property type="protein sequence ID" value="ENSGAGP00000000945.1"/>
        <property type="gene ID" value="ENSGAGG00000000748.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0052735">
        <property type="term" value="F:tRNA (cytidine-3-)-methyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032259">
        <property type="term" value="P:methylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008033">
        <property type="term" value="P:tRNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd02440">
        <property type="entry name" value="AdoMet_MTases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.150">
        <property type="entry name" value="Vaccinia Virus protein VP39"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013217">
        <property type="entry name" value="Methyltransf_12"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026113">
        <property type="entry name" value="METTL2/6/8-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029063">
        <property type="entry name" value="SAM-dependent_MTases_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22809">
        <property type="entry name" value="METHYLTRANSFERASE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22809:SF4">
        <property type="entry name" value="TRNA N(3)-METHYLCYTIDINE METHYLTRANSFERASE METTL2A-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08242">
        <property type="entry name" value="Methyltransf_12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF037755">
        <property type="entry name" value="Mettl2_prd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53335">
        <property type="entry name" value="S-adenosyl-L-methionine-dependent methyltransferases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4 10" id="KW-0489">Methyltransferase</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0949">S-adenosyl-L-methionine</keyword>
    <keyword evidence="5 10" id="KW-0808">Transferase</keyword>
    <keyword evidence="7" id="KW-0819">tRNA processing</keyword>
    <feature type="domain" description="Methyltransferase type 12" evidence="12">
        <location>
            <begin position="139"/>
            <end position="242"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="52"/>
            <end position="76"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="61"/>
            <end position="75"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009725"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022603"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022691"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022694"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00048054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00049374"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PIRNR" id="PIRNR037755"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF08242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000945.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="334" mass="38488" checksum="4E47175C829311C3" modified="2019-05-08" version="1">QENSRQLVPQDKQDEYELNANEYWNDFYKIHENGFFKDRHWLFTEFPELAPDQNLSQPEASEYKDGSTRKPRETRLGSCENGLCSLETRTQEHHQLDLMKDSNDIINRLDTDREAVPKLGDLKLSDGDFPGSSATYRILEVGCGAGNTVFPILQTNNDPGLFVYCCDFSNTAVDLVQTNAEYDSSRCFAFVHDLCKDQSPYPMPEESLDIIILIFVLSALLPEKMQCVINRLSRLLKPGGMILLRDYGRYDLAQLRFKKGRCLSDNFYVRGDGTRVYFFTQDELDALFTTAGLEKVQNLVDRRLQVNRGKQVTMHRVWIQCKYRKPFPSSTGQK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GH71</accession>
    <name>A0A452GH71_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="30">ATP-dependent DNA helicase DDX11</fullName>
            <ecNumber evidence="28">5.6.2.3</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="32">DEAD/H-box protein 11</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="31">DNA 5'-3' helicase DDX11</fullName>
        </alternativeName>
    </protein>
    <organism evidence="35 36">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="36" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="35" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="35" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="29">
            <text>ATP + H2O = ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:13065"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="5.6.2.3"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>[4Fe-4S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:49883"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Cytoskeleton</location>
            <location evidence="3">Microtubule organizing center</location>
            <location evidence="3">Centrosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="5">Cytoplasm</location>
            <location evidence="5">Cytoskeleton</location>
            <location evidence="5">Spindle pole</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Midbody</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Nucleus</location>
            <location evidence="4">Nucleolus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the DEAD box helicase family. DEAH subfamily. DDX11/CHL1 sub-subfamily.</text>
    </comment>
    <dbReference type="EC" id="5.6.2.3" evidence="28"/>
    <dbReference type="AlphaFoldDB" id="A0A452GH71"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001077.1">
        <property type="protein sequence ID" value="ENSGAGP00000000952.1"/>
        <property type="gene ID" value="ENSGAGG00000000211.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030496">
        <property type="term" value="C:midbody"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005730">
        <property type="term" value="C:nucleolus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000922">
        <property type="term" value="C:spindle pole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051539">
        <property type="term" value="F:4 iron, 4 sulfur cluster binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043139">
        <property type="term" value="F:5'-3' DNA helicase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016818">
        <property type="term" value="F:hydrolase activity, acting on acid anhydrides, in phosphorus-containing anhydrides"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006281">
        <property type="term" value="P:DNA repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006260">
        <property type="term" value="P:DNA replication"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034085">
        <property type="term" value="P:establishment of sister chromatid cohesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd18788">
        <property type="entry name" value="SF2_C_XPD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:001050">
        <property type="entry name" value="ATP-dependent DNA helicase DDX11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000910">
        <property type="entry name" value="probable ATP-dependent DNA helicase DDX11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000909">
        <property type="entry name" value="Putative ATP-dependent RNA helicase DDX11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006555">
        <property type="entry name" value="ATP-dep_Helicase_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045028">
        <property type="entry name" value="DinG/Rad3-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014013">
        <property type="entry name" value="Helic_SF1/SF2_ATP-bd_DinG/Rad3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006554">
        <property type="entry name" value="Helicase-like_DEXD_c2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010614">
        <property type="entry name" value="RAD3-like_helicase_DEAD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013020">
        <property type="entry name" value="Rad3/Chl1-like"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00604">
        <property type="entry name" value="rad3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11472:SF41">
        <property type="entry name" value="ATP-DEPENDENT DNA HELICASE DDX11-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11472">
        <property type="entry name" value="DNA REPAIR DEAD HELICASE RAD3/XP-D SUBFAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06733">
        <property type="entry name" value="DEAD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13307">
        <property type="entry name" value="Helicase_C_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00488">
        <property type="entry name" value="DEXDc2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00491">
        <property type="entry name" value="HELICc2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51193">
        <property type="entry name" value="HELICASE_ATP_BIND_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0004">4Fe-4S</keyword>
    <keyword evidence="22" id="KW-0010">Activator</keyword>
    <keyword evidence="16" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="9" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="25" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="7" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="13" id="KW-0227">DNA damage</keyword>
    <keyword evidence="24" id="KW-0234">DNA repair</keyword>
    <keyword evidence="10" id="KW-0235">DNA replication</keyword>
    <keyword evidence="21" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="15" id="KW-0347">Helicase</keyword>
    <keyword evidence="14" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="18" id="KW-0408">Iron</keyword>
    <keyword evidence="19" id="KW-0411">Iron-sulfur</keyword>
    <keyword evidence="26" id="KW-0413">Isomerase</keyword>
    <keyword evidence="11" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="12" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="27" id="KW-0539">Nucleus</keyword>
    <keyword evidence="36" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="17" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="23" id="KW-0804">Transcription</keyword>
    <keyword evidence="20" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="Helicase ATP-binding" evidence="34">
        <location>
            <begin position="8"/>
            <end position="440"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="33">
        <location>
            <begin position="77"/>
            <end position="97"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="33">
        <location>
            <begin position="84"/>
            <end position="97"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001966"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004214"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004604"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004647"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00008435"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022485"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022705"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022806"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023014"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00023204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00023235"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="ARBA" id="ARBA00044969"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="ARBA" id="ARBA00048954"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="ARBA" id="ARBA00074489"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="31">
        <source>
            <dbReference type="ARBA" id="ARBA00079245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="32">
        <source>
            <dbReference type="ARBA" id="ARBA00080369"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="33">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="34">
        <source>
            <dbReference type="PROSITE" id="PS51193"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="35">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000952.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="36">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="853" mass="96743" checksum="BFFFCEC366E4647B" modified="2019-05-08" version="1">MQNSQGKGRVNFPFPYTPYSIQEDFMANLYDVLEAGKVGIFESPTGTGKSLSLICGALSWLRDFEEKKKQEEARLLADEHSGQDKQQQPTSCSAEASNCQGSAGELVWITEFMQKKEEQAMVDRLKEEQIRRKKREDRLEKIRHNVQLKYASKRKRLEDDETQRLLQLSKEILSTELGSDVLEQLDHEEEELILAEYESDEEKKVELGLDDDDDDDLEEEHVTKIYYCSRTHSQLAQFVHEVQKSPFGEEIRLVSLGSRQNLCVNEEVRHLGAVQLINDRCVEMQKNKHGKKSEGEESEGKKRRVSRAVCPFYSYEQMQFLRDEVLVKVKDIEQLVALGKETKACPYYGSRYAIPAAQLVVLPYQMLLHESTRCASGIKLKDQVVIIDEAHNLIDTITCIYSAEVRGSQLCCAHSQLLQYMERYRKRLKAKNLMYIKQILYLLERFVSILGGNVNQNPNAQSISQAGTELKSINDFLFQSQIDNINLFKVCHPVEVEDGQPKMASPLMQIEGFLSALTSANQDGRVILNRQGTVAKSSLKFLLLNPAVHFAKVLKECRAAIVAGGTMQPVSDFREQLLSCAIGADRVVEFSCGHVIPPDNILPIILCSGPSNQQLEFTYQKRDLPHMVSTMGRILGNLCNVVPGGVVCFFPSYEYEKQVYTHWEGTGLLTRLATKKKIFQEPKKANQVEQVLAEYTKCIKRCSQVGGHVTGALLFSVVGGKMSEGINFSDDLGRCVVMVGMPYPNIKSPELQEKMAYLDKTMPRTAGQPPSKMLIENLCMKAVNQSIGRAIRHQKDFASILLVDHRYARPSVFNKLPQWIKERTEIKYTFGLAFAALRKFHHEKKSCYRSEAS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GH74</accession>
    <name>A0A452GH74_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">DMAP1-binding domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell projection</location>
            <location evidence="1">Neuron projection</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the DIP2 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000001042.1">
        <property type="protein sequence ID" value="ENSGAGP00000000919.1"/>
        <property type="gene ID" value="ENSGAGG00000000729.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009986">
        <property type="term" value="C:cell surface"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043005">
        <property type="term" value="C:neuron projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd05905">
        <property type="entry name" value="Dip2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.300.30:FF:000003">
        <property type="entry name" value="DIP2 disco-interacting protein 2 homolog A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.300.30:FF:000001">
        <property type="entry name" value="DIP2 disco-interacting protein 2 homolog C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.12780:FF:000004">
        <property type="entry name" value="Disco interacting protein 2 homolog A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.12780:FF:000002">
        <property type="entry name" value="Disco interacting protein 2 homolog B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.300.30">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.12780">
        <property type="entry name" value="N-terminal domain of ligase-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025110">
        <property type="entry name" value="AMP-bd_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045851">
        <property type="entry name" value="AMP-bd_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000873">
        <property type="entry name" value="AMP-dep_synth/lig_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042099">
        <property type="entry name" value="ANL_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037337">
        <property type="entry name" value="Dip2-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010506">
        <property type="entry name" value="DMAP1-bd"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22754">
        <property type="entry name" value="DISCO-INTERACTING PROTEIN 2 DIP2 -RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22754:SF34">
        <property type="entry name" value="DISCO-INTERACTING PROTEIN 2 HOMOLOG A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00501">
        <property type="entry name" value="AMP-binding"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23024">
        <property type="entry name" value="AMP-dom_DIP2-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06464">
        <property type="entry name" value="DMAP_binding"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01137">
        <property type="entry name" value="DMAP_binding"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56801">
        <property type="entry name" value="Acetyl-CoA synthetase-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51912">
        <property type="entry name" value="DMAP1_BIND"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0966">Cell projection</keyword>
    <keyword evidence="4" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="DMAP1-binding" evidence="7">
        <location>
            <begin position="13"/>
            <end position="109"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="78"/>
            <end position="161"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="91"/>
            <end position="103"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="138"/>
            <end position="149"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004487"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007735"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS51912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000919.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1543" mass="168076" checksum="664CE405AD3839C9" modified="2019-05-08" version="1">MAAERGGCSLEAVPVPLPPEVRESLAELELELSEGDITQKGYEKKRAKLLARYIPLIQDVHTEAVQAALAKYKERKMPMPSKRRSVLVHSSVETYTPPDTSSASEDEGSLRRQGRIPSTPFQSHSNVEPWLNRVIQGSSTSSSASSTSSHPGGKPAAASNTATATAAATVLADLMAHAQLDNHSAPPDVTTGLVEHLHHERPQVASVRGIPRGYNSSILETADGVPVNSRVSSKIQQLLNTLKRPKRPPLKEFFVDDFEELLEVQQPDPNQPKPEGGEMTALKGEPLGVVTNWPPSLLAALQRWGTTQPKTPCLTALDTTGKAVYTLTYGKLWSRSLKLAYTLLNKLTSKNEPLLKPGHRVALVFPNSDPVMFMVAFYGCLLADLVPVPIEVPLTRKDAGSQQIGFLLGSCGVTLALTTDVCQKGLPKAQTGEVVTFKGWPRLVWFVIDGKHLAKPAKDWHPQVRDASADIAYIEYKTSKEGSTMGITISHASMLAHCHALTQACGYSEAETLTNVLDFKRDAGLWHGVLTSVMNRMHVVSIPYALMKVNPLSWIQKVCSYKACVAVVKSRDMHWSLLAQRDQRDVNLSSLRMLIVADGANPWSISSCDAFLNVFQSRGLRPEVICPCASSSEALTVAIRRPPDLGGPPPGKAVLSMNCLSYNVIRVDTEEKLSVLTVQDVGQVMPGANVCVVKADGTPYLCKTDEVGEICVNSGATGTAYYGLLGITKNVFETIPVTAAGVPVSDQPFTRTGLLGFIGPDSLVFVVGKLDGLMVVNIRKHNADDVVATALAVEPMKFVYRGRIAVFSVTVLHDDRIVLVAEQRPDASEEDSFQWMSRVLQAIDSIHQVGVYCLALVPANTLPKAPLGGIHISETKQRFLEGALHPCNVLMCPHTCVTNLPKPRQKQPDVGPASMIVGNLVAGKRIAQASGRDVTQLEDNDQARKFLYLADVLQWRAQTTPDHPLFLLLNSKGTVASTASCIQLHKKAERVAAALMEKGRLNVGDHVALVYPPGVDLIATFYGCLYAGCIPITVRPPHPQNLATTLPTVKMIVEVSKSACILTTQGIMKLLKSKEAATAVDIKTWPTILDTDDIPKKKVANIFRPPSPDMLAYLDFSVSTTGILAGVKMSHAATSALCRSVKLQCELYPSRQIAICLDPYCGLGFALWCLCSVYSGHQSILVPPLELESNVSLWLSAVSQYKVRVTFCSYSVMEMCTKGLGTQTDVLRMKGVNLSCVRTCMVVAEERPRIALTQSFSKLFKDLGLSARSVSTTFGCRVNVAICLQPNRLGRLAEQGTAGPDPTTVYVDMRALRHDRVRLVERGSPHSLPLMESGKILPGVKVIIAHTETKGPLGDSHLGEIWVSSPHNATGYYTVYGEEALHADHFSARLSFGDTQTVWARTGYLGFLRRTELTDANGERHDALYVVGSLDETLELRGMRYHPIDIETSVIRAHKSIAECAVFTWTNLLVVVVELEGSEQEALDLVALVTNVVLEEHYLIVGVVVIVDPGVIPINSRGEKQRMHLRDGFLADQLDPIYVAYNM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXQ8</accession>
    <name>A0A452GXQ8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Zinc finger protein 777</fullName>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">May be involved in transcriptional regulation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GXQ8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007999.1">
        <property type="protein sequence ID" value="ENSGAGP00000006897.1"/>
        <property type="gene ID" value="ENSGAGG00000005537.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005654">
        <property type="term" value="C:nucleoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001227">
        <property type="term" value="F:DNA-binding transcription repressor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001817">
        <property type="term" value="P:regulation of cytokine production"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002682">
        <property type="term" value="P:regulation of immune system process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07765">
        <property type="entry name" value="KRAB_A-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002343">
        <property type="entry name" value="Zinc finger protein 33A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001498">
        <property type="entry name" value="Zinc finger protein 404"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001119">
        <property type="entry name" value="zinc finger protein 408"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001602">
        <property type="entry name" value="Zinc finger protein 490"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000180">
        <property type="entry name" value="Zinc finger protein 689"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001909">
        <property type="entry name" value="KRAB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036051">
        <property type="entry name" value="KRAB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24399:SF54">
        <property type="entry name" value="GASTRULA ZINC FINGER PROTEIN XLCGF26.1-LIKE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24399">
        <property type="entry name" value="ZINC FINGER AND BTB DOMAIN-CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01352">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109640">
        <property type="entry name" value="KRAB domain (Kruppel-associated box)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50805">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="5"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="10" id="KW-0804">Transcription</keyword>
    <keyword evidence="8" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="7" id="KW-0862">Zinc</keyword>
    <keyword evidence="6 12" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="KRAB" evidence="15">
        <location>
            <begin position="128"/>
            <end position="214"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="14">
        <location>
            <begin position="272"/>
            <end position="299"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="14">
        <location>
            <begin position="359"/>
            <end position="386"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="14">
        <location>
            <begin position="389"/>
            <end position="416"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="14">
        <location>
            <begin position="417"/>
            <end position="444"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="14">
        <location>
            <begin position="445"/>
            <end position="472"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="1"/>
            <end position="21"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="223"/>
            <end position="255"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="299"/>
            <end position="343"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="469"/>
            <end position="497"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="13">
        <location>
            <begin position="475"/>
            <end position="490"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00003767"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006897.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="18">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="497" mass="55998" checksum="1FB47B3C7207DE00" modified="2019-05-08" version="1">GAGTLPAQSRRSARPRSQAGSADLQTAEISLTVVAEIQAVDRKVEVQAAQLMNLEGRMRMAETKLVGCEKTAVEFGNQLESKWTALGTLIQEYGQLQRRLENMENLLKNRNFWILRLPPGANGEVPKVRRSQLVVRGCKQEWKSLDEWQKELYRHIMKGNYEAVISLGKDCLQNEAFVMPLDDVNVSVLSVLTDEQQCGEEGPANLDLPSLLPGDWEELFSSPEEELTKESQSSSAVLQRNPPGSGLRRSARRGRDLMKTVSEEAHAAEGPYICCECGESFIDKQLLATHQRIHASGGACTSSLEHGENIRQKPKVTTPPRTQAPVRSKPPKRPEPEKSSNLKYGFVKHQANHMVERPYTCTQCRESFSLEVSLILHQKLHTGKGDGPLTCTYCGKDFRDLSKAIRHQRIHTGERPYQCTECGKSFIRRDHLLKHWRVHTGETPYQCPVCGKNFRYKESLNCHQKIHTRNPQPLESSQQLETGTQTTLLGVKKENKQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXT4</accession>
    <name>A0A452GXT4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="19">Nitric oxide synthase 3</fullName>
            <ecNumber evidence="6">1.14.13.39</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="16">Constitutive NOS</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="22">EC-NOS</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="20">NOS type III</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="21">Nitric oxide synthase, endothelial</fullName>
        </alternativeName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>2 L-arginine + 3 NADPH + 4 O2 + H(+) = 2 L-citrulline + 2 nitric oxide + 3 NADP(+) + 4 H2O</text>
            <dbReference type="Rhea" id="RHEA:19897"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:15379"/>
            <dbReference type="ChEBI" id="CHEBI:16480"/>
            <dbReference type="ChEBI" id="CHEBI:32682"/>
            <dbReference type="ChEBI" id="CHEBI:57743"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="EC" id="1.14.13.39"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="17">
            <dbReference type="Rhea" id="RHEA:19898"/>
        </physiologicalReaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="2">
            <name>(6R)-L-erythro-5,6,7,8-tetrahydrobiopterin</name>
            <dbReference type="ChEBI" id="CHEBI:59560"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="4">
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>FMN</name>
            <dbReference type="ChEBI" id="CHEBI:58210"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="3">
            <name>heme b</name>
            <dbReference type="ChEBI" id="CHEBI:60344"/>
        </cofactor>
    </comment>
    <comment type="subunit">
        <text evidence="18">Homodimer. Interacts with NOSIP and NOSTRIN. Interacts with HSP90AB1. Forms a complex with ASL, ASS1 and SLC7A1; the complex regulates cell-autonomous L-arginine synthesis and citrulline recycling while channeling extracellular L-arginine to nitric oxide synthesis pathway.</text>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the NOS family.</text>
    </comment>
    <dbReference type="EC" id="1.14.13.39" evidence="6"/>
    <dbReference type="AlphaFoldDB" id="A0A452GXT4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007950.1">
        <property type="protein sequence ID" value="ENSGAGP00000006851.1"/>
        <property type="gene ID" value="ENSGAGG00000000423.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010181">
        <property type="term" value="F:FMN binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004517">
        <property type="term" value="F:nitric-oxide synthase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006809">
        <property type="term" value="P:nitric oxide biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00795">
        <property type="entry name" value="NOS_oxygenase_euk"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.440.10:FF:000001">
        <property type="entry name" value="Endothelial nitric oxide synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1230.10:FF:000001">
        <property type="entry name" value="Nitric oxide synthase, brain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.360">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.340.10">
        <property type="entry name" value="Nitric Oxide Synthase, Chain A, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1230.10">
        <property type="entry name" value="Nitric Oxide Synthase, Chain A, domain 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.440.10">
        <property type="entry name" value="Nitric Oxide Synthase,Heme Domain,Chain A domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001094">
        <property type="entry name" value="Flavdoxin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008254">
        <property type="entry name" value="Flavodoxin/NO_synth"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029039">
        <property type="entry name" value="Flavoprotein-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050607">
        <property type="entry name" value="NOS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044943">
        <property type="entry name" value="NOS_dom_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044940">
        <property type="entry name" value="NOS_dom_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044944">
        <property type="entry name" value="NOS_dom_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004030">
        <property type="entry name" value="NOS_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036119">
        <property type="entry name" value="NOS_N_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43410:SF1">
        <property type="entry name" value="NITRIC OXIDE SYNTHASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43410">
        <property type="entry name" value="NITRIC OXIDE SYNTHASE OXYGENASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00258">
        <property type="entry name" value="Flavodoxin_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02898">
        <property type="entry name" value="NO_synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00369">
        <property type="entry name" value="FLAVODOXIN"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52218">
        <property type="entry name" value="Flavoproteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56512">
        <property type="entry name" value="Nitric oxide (NO) synthase oxygenase domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50902">
        <property type="entry name" value="FLAVODOXIN_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="13" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="11" id="KW-0274">FAD</keyword>
    <keyword evidence="8" id="KW-0285">Flavoprotein</keyword>
    <keyword evidence="9" id="KW-0288">FMN</keyword>
    <keyword evidence="7" id="KW-0349">Heme</keyword>
    <keyword evidence="15" id="KW-0408">Iron</keyword>
    <keyword evidence="10" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="12" id="KW-0521">NADP</keyword>
    <keyword evidence="14" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Flavodoxin-like" evidence="23">
        <location>
            <begin position="417"/>
            <end position="526"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001917"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00001950"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00001970"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00001974"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00006267"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00012989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022630"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022643"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022827"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022857"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00029794"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00047419"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00049679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00049764"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00049791"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00049814"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00049818"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS50902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006851.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="526" mass="58854" checksum="FC61BD4C84510A3D" modified="2019-05-08" version="1">MFPKQMVCQADEGPPAKEELLPLARDFIHQYYSSIKRADSRVHMQRVQEVEQEIRATGTYQLLEPELIFGAKQAWRNAARCLGRIQWNKLQVFDARDCASVQEMFGFLCTHLQYATNRGNLRSAITIFPQRMPGRGDFRIWNSQLIRYAGYPQPDGSVLGDPANVDITELCIQHGWTPGHGRFDVLPLLLQAPDEAPELFLLPPELVLEVPIQHPTLAWFGELGLRWHALPAVANMLLEIGGLQFPAAPFNGWYVGSEIGMRNFCDSNRYNLLQEVALHMGLDTGKTSSLWKDKAAVEINVAVLHSFQVAKVTIVDHHVATESFVKHMDNELQTRGGCPADWVWIVPPISGSLTPVFHQEMVNYLLSPCFRYQPDAWKVYTSKGTSITRKKTFKEVANAVKFSAKLMVHVMARRVKATILYATETGKSRTYAWNLGELFRRAFDPKVLCMDEYDIVSLEHETLVLVVTSTFGNGDPPESGEPGALGPNVASLATATAAHVPLVPPSWSVGLMPCVPLGREFSIVMQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXV7</accession>
    <name>A0A452GXV7_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="16">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>heme b</name>
            <dbReference type="ChEBI" id="CHEBI:60344"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Membrane</location>
            <topology evidence="2">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the FRRS1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GXV7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008060.1">
        <property type="protein sequence ID" value="ENSGAGP00000006953.1"/>
        <property type="gene ID" value="ENSGAGG00000005607.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016722">
        <property type="term" value="F:oxidoreductase activity, acting on metal ions"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006879">
        <property type="term" value="P:intracellular iron ion homeostasis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd08760">
        <property type="entry name" value="Cyt_b561_FRRS1_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd09628">
        <property type="entry name" value="DOMON_SDR_2_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd08544">
        <property type="entry name" value="Reeler"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.4060:FF:000003">
        <property type="entry name" value="Ferric chelate reductase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.1770">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.4060">
        <property type="entry name" value="Reeler domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006593">
        <property type="entry name" value="Cyt_b561/ferric_Rdtase_TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005018">
        <property type="entry name" value="DOMON_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051237">
        <property type="entry name" value="Ferric-chelate_Red/DefProt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002861">
        <property type="entry name" value="Reeler_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042307">
        <property type="entry name" value="Reeler_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45828">
        <property type="entry name" value="CYTOCHROME B561/FERRIC REDUCTASE TRANSMEMBRANE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45828:SF3">
        <property type="entry name" value="FERRIC-CHELATE REDUCTASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03351">
        <property type="entry name" value="DOMON"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02014">
        <property type="entry name" value="Reeler"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00665">
        <property type="entry name" value="B561"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00664">
        <property type="entry name" value="DoH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50939">
        <property type="entry name" value="CYTOCHROME_B561"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50836">
        <property type="entry name" value="DOMON"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51019">
        <property type="entry name" value="REELIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0249">Electron transport</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="8" id="KW-0408">Iron</keyword>
    <keyword evidence="9 11" id="KW-0472">Membrane</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="12" id="KW-0732">Signal</keyword>
    <keyword evidence="5 11" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 11" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="4" id="KW-0813">Transport</keyword>
    <feature type="signal peptide" evidence="12">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019470524" evidence="12">
        <location>
            <begin position="23"/>
            <end position="592"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="375"/>
            <end position="396"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="416"/>
            <end position="435"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="447"/>
            <end position="466"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="478"/>
            <end position="500"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="512"/>
            <end position="531"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="571"/>
            <end position="589"/>
        </location>
    </feature>
    <feature type="domain" description="Reelin" evidence="15">
        <location>
            <begin position="13"/>
            <end position="178"/>
        </location>
    </feature>
    <feature type="domain" description="DOMON" evidence="13">
        <location>
            <begin position="216"/>
            <end position="331"/>
        </location>
    </feature>
    <feature type="domain" description="Cytochrome b561" evidence="14">
        <location>
            <begin position="336"/>
            <end position="534"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001970"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00009195"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022982"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50836"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50939"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS51019"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006953.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="592" mass="66064" checksum="593408900E168CAB" modified="2019-05-08" version="1">MEPPVLAFALWIFAFLSVPAAGYPNGKVREACSSMMPYHGHSPQPSPEHTISVNETKFRPGDHIKVSLSGPVFKGFFIQARNAENLTGAAVGSFMLAEERISQLLTCGLVKNSAVSHTSKSKKTNVEAYWVAPTDAPEHVQFLATVVEKYKIFWVKIPGPIVSQPNAPPLTTPTSITPDTLPTSQPVFRLTTSFNASSCGITKFCIRNPPKCDPGAAYCFFLSFKQNNHLVLIEMSGPSEGYIAFALSHDQWMGDDDAYLCINEDHRINIKTAYLSGRTPPVLDSENGLEDISWRVADGLLQCSFRRNLHLPAYKGRFNLDANYYIFLADGEASEGGLIHKHHRQPLITNGMYNFTDSPKDVGGSRSPILLKLHASLMFAAWMTTVSIGVIVARFFKPVWPHSLFGKEIWFQVHRMLMLTTVLLTSIAFVLPFIYRGGWSEWAGFHPYLGCTVMALAIFQPLMAGFRPPPHAPRRQVFNWVHWSTGTTARILAVVTMFLGMDLPALNLPDPWDTYTMIGFVAWHVSIDILLEIHSYCLIRKVEVIDEDRIQILQSLTSAEAEGHTFKEMVLTIYICGNIAFLITFLAAIHQL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXV9</accession>
    <name>A0A452GXV9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Prohibitin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="6">Protein with pleiotropic attributes mediated in a cell-compartment- and tissue-specific manner, which include the plasma membrane-associated cell signaling functions, mitochondrial chaperone, and transcriptional co-regulator of transcription factors and sex steroid hormones in the nucleus.</text>
    </comment>
    <comment type="subunit">
        <text evidence="7">The mitochondrial prohibitin complex consists of two subunits (PHB1 and PHB2), assembled into a membrane-associated ring-shaped supercomplex of approximately 1 mDa.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 8">Mitochondrion inner membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 8">Belongs to the prohibitin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GXV9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008100.1">
        <property type="protein sequence ID" value="ENSGAGP00000006992.1"/>
        <property type="gene ID" value="ENSGAGG00000005631.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005743">
        <property type="term" value="C:mitochondrial inner membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007005">
        <property type="term" value="P:mitochondrion organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd03401">
        <property type="entry name" value="SPFH_prohibitin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.479.30:FF:000001">
        <property type="entry name" value="Prohibitin 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.479.30">
        <property type="entry name" value="Band 7 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001107">
        <property type="entry name" value="Band_7"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036013">
        <property type="entry name" value="Band_7/SPFH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000163">
        <property type="entry name" value="Prohibitin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23222">
        <property type="entry name" value="PROHIBITIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23222:SF1">
        <property type="entry name" value="PROHIBITIN-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01145">
        <property type="entry name" value="Band_7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00679">
        <property type="entry name" value="PROHIBITIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00244">
        <property type="entry name" value="PHB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF117892">
        <property type="entry name" value="Band 7/SPFH domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="3 8" id="KW-0999">Mitochondrion inner membrane</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Band 7" evidence="9">
        <location>
            <begin position="39"/>
            <end position="201"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009658"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022792"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00037479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00064091"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="RuleBase" id="RU366048"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="SMART" id="SM00244"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006992.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="289" mass="32139" checksum="8A2F7DC7C5656DC4" modified="2019-05-08" version="1">MAQNLKDLAGRLPAGPRGMGTALKLLLGAGAVAYGVRESVFTVEGGQRAIFFNRIGGIQPDTILAEGLHFRIPWFQYPIIYDIRARPRKISSPTGSKDLQMVNISLRVLSRPNAAELPKMYQRLGLDYEERVLPSIVNEVLKSVVAKFNASQLITQRAQVSLLIRRELTERAKDFSLILDDVAITELSFSREYTAAVEAKQVAQQEAQRAQFLVEKAKQEQKQKIVQAEGEATAAKMIGEALSKNPGYIKLRKIRAAQNISKTIAASQNRVYLTADNLQLNLQDEGFTR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXW3</accession>
    <name>A0A452GXW3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">G-protein coupled receptors family 1 profile domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="9">Belongs to the G-protein coupled receptor 1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GXW3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008110.1">
        <property type="protein sequence ID" value="ENSGAGP00000007000.1"/>
        <property type="gene ID" value="ENSGAGG00000005601.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004930">
        <property type="term" value="F:G protein-coupled receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1070.10">
        <property type="entry name" value="Rhodopsin 7-helix transmembrane proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000276">
        <property type="entry name" value="GPCR_Rhodpsn"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017452">
        <property type="entry name" value="GPCR_Rhodpsn_7TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026234">
        <property type="entry name" value="MRGPCRFAMILY"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11334">
        <property type="entry name" value="MAS-RELATED G-PROTEIN COUPLED RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11334:SF29">
        <property type="entry name" value="MAS-RELATED G-PROTEIN COUPLED RECEPTOR MEMBER X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00237">
        <property type="entry name" value="GPCRRHODOPSN"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR02108">
        <property type="entry name" value="MRGPCRFAMILY"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81321">
        <property type="entry name" value="Family A G protein-coupled receptor-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00237">
        <property type="entry name" value="G_PROTEIN_RECEP_F1_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50262">
        <property type="entry name" value="G_PROTEIN_RECEP_F1_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="5 9" id="KW-0297">G-protein coupled receptor</keyword>
    <keyword evidence="6 10" id="KW-0472">Membrane</keyword>
    <keyword evidence="7 9" id="KW-0675">Receptor</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8 9" id="KW-0807">Transducer</keyword>
    <keyword evidence="3 9 10" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 10" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="20"/>
            <end position="46"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="58"/>
            <end position="83"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="89"/>
            <end position="113"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="134"/>
            <end position="155"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="175"/>
            <end position="202"/>
        </location>
    </feature>
    <feature type="domain" description="G-protein coupled receptors family 1 profile" evidence="11">
        <location>
            <begin position="38"/>
            <end position="131"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023224"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="RuleBase" id="RU000688"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50262"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007000.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="238" mass="26947" checksum="2674910B73D4D245" modified="2019-05-08" version="1">MPLNPTPTHPAQPSAIPDLGFAPVLSISLFICLVGLVGNGMVLWFLGFPVKKEPFTIYILNLAIADFGFLLCMVAFLIIIILYLHMDFLVGFVVIEAIQWVALFIYNTGLYLLTDISIHRCLSVLYPIWYRCHIQKTCLPLCVLCSGAISASMTMTVRCSSHRLIVLMAFLNHSMIHLFVMSLVILLPCMNSSINSFLYFLIGRHGKQQLREPLREVFQRIFNEKAGPQEKRPTKQVK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXW9</accession>
    <name>A0A452GXW9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">18S rRNA (pseudouridine(1248)-N1)-methyltransferase</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="17">18S rRNA Psi1248 methyltransferase</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="13">Nucleolar protein EMG1 homolog</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="16">Protein C2f</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="14">Ribosome biogenesis protein NEP1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>pseudouridine(1248) in human 18S rRNA + S-adenosyl-L-methionine = N(1)-methylpseudouridine(1248) in human 18S rRNA + S-adenosyl-L-homocysteine + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46712"/>
            <dbReference type="Rhea" id="RHEA-COMP:11638"/>
            <dbReference type="Rhea" id="RHEA-COMP:11639"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57856"/>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
            <dbReference type="ChEBI" id="CHEBI:65314"/>
            <dbReference type="ChEBI" id="CHEBI:74890"/>
        </reaction>
    </comment>
    <comment type="subunit">
        <text evidence="12">Homodimer. Part of the small subunit (SSU) processome, composed of more than 70 proteins and the RNA chaperone small nucleolar RNA (snoRNA) U3.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
            <location evidence="1">Nucleolus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the class IV-like SAM-binding methyltransferase superfamily. RNA methyltransferase NEP1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GXW9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008113.1">
        <property type="protein sequence ID" value="ENSGAGP00000007003.1"/>
        <property type="gene ID" value="ENSGAGG00000005641.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032040">
        <property type="term" value="C:small-subunit processome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070037">
        <property type="term" value="F:rRNA (pseudouridine) methyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019843">
        <property type="term" value="F:rRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070475">
        <property type="term" value="P:rRNA base methylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd18088">
        <property type="entry name" value="Nep1-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.1280.10:FF:000003">
        <property type="entry name" value="Ribosomal RNA small subunit methyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.1280.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029028">
        <property type="entry name" value="Alpha/beta_knot_MTases"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005304">
        <property type="entry name" value="Rbsml_bgen_MeTrfase_EMG1/NEP1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029026">
        <property type="entry name" value="tRNA_m1G_MTases_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12636">
        <property type="entry name" value="NEP1/MRA1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12636:SF5">
        <property type="entry name" value="RIBOSOMAL RNA SMALL SUBUNIT METHYLTRANSFERASE NEP1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03587">
        <property type="entry name" value="EMG1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF75217">
        <property type="entry name" value="alpha/beta knot"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0489">Methyltransferase</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0690">Ribosome biogenesis</keyword>
    <keyword evidence="9" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="4" id="KW-0698">rRNA processing</keyword>
    <keyword evidence="8" id="KW-0699">rRNA-binding</keyword>
    <keyword evidence="7" id="KW-0949">S-adenosyl-L-methionine</keyword>
    <keyword evidence="6" id="KW-0808">Transferase</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004604"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008115"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022517"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022552"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022603"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022691"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022730"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00051029"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00063756"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00075957"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00077532"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00078441"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00079565"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00079971"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007003.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="151" mass="16542" checksum="2CBDAB6489384D00" modified="2019-05-08" version="1">MLMDSPLNRAGLLQVYIHTQKNVLIEVNPQTRIPRTFDRFCGLMVQLLHKLSVRAADGPQKLLKVIKNPVTDHLPVGCMKIGTSFSVPNVTDVRELVPTTEPVAIVVGAFAHGSLNVNYTEKMISISNYPLSAALTCAKITTAFEEAWGVV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GY25</accession>
    <name>A0A452GY25_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="5">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GY25"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008135.1">
        <property type="protein sequence ID" value="ENSGAGP00000007023.1"/>
        <property type="gene ID" value="ENSGAGG00000005653.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000008163.1">
        <property type="protein sequence ID" value="ENSGAGP00000007047.1"/>
        <property type="gene ID" value="ENSGAGG00000005653.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd13242">
        <property type="entry name" value="PH_puratrophin-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00160">
        <property type="entry name" value="RhoGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.60">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.900.10">
        <property type="entry name" value="Dbl homology (DH) domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035899">
        <property type="entry name" value="DBL_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000219">
        <property type="entry name" value="DH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052231">
        <property type="entry name" value="Rho_GEF_signaling-related"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055251">
        <property type="entry name" value="SOS1_NGEF_PH"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45845:SF1">
        <property type="entry name" value="PLECKSTRIN HOMOLOGY AND RHOGEF DOMAIN CONTAINING G4B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45845">
        <property type="entry name" value="RHO GUANINE NUCLEOTIDE EXCHANGE FACTOR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00621">
        <property type="entry name" value="RhoGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22697">
        <property type="entry name" value="SOS1_NGEF_PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00233">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00325">
        <property type="entry name" value="RhoGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48065">
        <property type="entry name" value="DBL homology domain (DH-domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46966">
        <property type="entry name" value="Spectrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50010">
        <property type="entry name" value="DH_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="2">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5044604469" evidence="2">
        <location>
            <begin position="24"/>
            <end position="1203"/>
        </location>
    </feature>
    <feature type="domain" description="DH" evidence="4">
        <location>
            <begin position="685"/>
            <end position="862"/>
        </location>
    </feature>
    <feature type="domain" description="PH" evidence="3">
        <location>
            <begin position="874"/>
            <end position="982"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="485"/>
            <end position="524"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="563"/>
            <end position="601"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="662"/>
            <end position="684"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1042"/>
            <end position="1066"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1108"/>
            <end position="1191"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="1">
        <location>
            <begin position="493"/>
            <end position="504"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="505"/>
            <end position="515"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="575"/>
            <end position="593"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="672"/>
            <end position="681"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="1">
        <location>
            <begin position="1052"/>
            <end position="1066"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="1111"/>
            <end position="1120"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="1">
        <location>
            <begin position="1121"/>
            <end position="1140"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="1160"/>
            <end position="1171"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="1181"/>
            <end position="1191"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50010"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007023.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1203" mass="138394" checksum="0E68C922A325A61E" modified="2019-05-08" version="1">MLSQRLFFLCFIWLGSRDRGGRAVVQVCTRSTIWSSEHSTTTELARLLMYFYRIPRKEVRDLGLVILVDARKCQPSPGLFKAFTVLQSAVPHSIHSVLILTEKESAFRPDKDTAIQYEVLTSLKALQKHIDCTQLTAEFDGTFLYNHHDWMRFRKKLEPFITNCKEAIVFLQNSVCSLNSNRTLSTVQEVRDLIDKHKTMMKLVLEDALLVTLRLEGGTILARLRKEEFCNTEDYRDAMEAITRLYNQVDEEVHRLVLISNKCLQQLENLQEIRKFEEHCDQIKLWFENERERYLGPLDQELLSLESVKKQQEEFRVFSDKVTQYCEKGLQLIQKNCSLNLEEKVQTFKSYLSNVSTQTEERKSELEKLIHLCEFYETAHRWMLHCNEYLEHLTVENKYSQSVIQSLQSYQQEATKVSAENFQKVNEIVLGLASKKELKQWNILWLKCQQTRHHLEEVLSEALKGSNKETSRKVPQEADVKVNCEFDTEGARRPSQSLSRLSSSTDNDIWDNNTKPLHKPQHMSLPSGVPPFLDHENSNQNMDVYVQLNASPPQTFTPLLGGRQRASPISEDMDSVSTSGSASCYSEPVQSPTTRHRKHPLKKIMKKTQSFEFTQLESSPSELHRHGYTGVYIKGLEVASNVAAEKKYMQRLNIKSPLISRNRSLSSPSRIHHTEEEEKKRAGSKVQHIMDEMITTEREYVRSLGYIIDNYFPEMERIDLPQDLRGKRNIIFGNLEKLYDFHCQYFLKELEHCRDFPLRVSHCFLRHEEQFGMYALYSKNKPQSDALLTSHGNAFFKNKQQDLGDKMDLASYLLKPIQRMSKYALLLKDMIKECTKTQEQELNDLRTAEEMVKFQLRHGNDLLAMDAIRGCDVNLKEQGQLRCQDEFIVCCGRKKYLRHVFLFEDLILFSKTKKIDGGYDIYMYKQSFKTAEIGMTENVGDSGLRFEIWFRRRRKSQDTYILQANSAEVKIAWTSIIGRILWRQALRNRELRMQEMVSMGIGNKPFMDIKPSEAAISDRAIDYIMKGTESRSRASIAVSSFDHSTPFKRPHSTISNSSTSSSSSQSSSSILGSLNLHVYSSPPHPGLLGPPFYHWSYDIRTCIEEDELEQETGSQPSMITESSESSQCTSGESVSGFSSSAHPGLSQLSTETFLDEDASNPGSKPPSQCTSPALPEKNPTSKHTNQYVTANKTSHVLGLSTMV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GY26</accession>
    <name>A0A452GY26_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Ribonuclease A-domain domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the pancreatic ribonuclease family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GY26"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008123.1">
        <property type="protein sequence ID" value="ENSGAGP00000007013.1"/>
        <property type="gene ID" value="ENSGAGG00000005645.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003676">
        <property type="term" value="F:nucleic acid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004540">
        <property type="term" value="F:RNA nuclease activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050830">
        <property type="term" value="P:defense response to Gram-positive bacterium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd06265">
        <property type="entry name" value="RNase_A_canonical"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.130.10">
        <property type="entry name" value="Ribonuclease A-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001427">
        <property type="entry name" value="RNaseA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036816">
        <property type="entry name" value="RNaseA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023412">
        <property type="entry name" value="RNaseA_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11437:SF10">
        <property type="entry name" value="ANGIOGENIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11437">
        <property type="entry name" value="RIBONUCLEASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00074">
        <property type="entry name" value="RnaseA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00794">
        <property type="entry name" value="RIBONUCLEASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00092">
        <property type="entry name" value="RNAse_Pc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54076">
        <property type="entry name" value="RNase A-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0964">Secreted</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="5">
        <location>
            <begin position="1"/>
            <end position="34"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019330107" description="Ribonuclease A-domain domain-containing protein" evidence="5">
        <location>
            <begin position="35"/>
            <end position="160"/>
        </location>
    </feature>
    <feature type="domain" description="Ribonuclease A-domain" evidence="6">
        <location>
            <begin position="36"/>
            <end position="155"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005600"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="SMART" id="SM00092"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007013.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="160" mass="17676" checksum="820A122A4575BE48" modified="2019-05-08" version="1">MSPEIATVGPAMKTRLGPGLLLLIFLLPACLTAAGRETPYEKFQRQHVDTSGSWEPDPNRYCNLMMPRRNMTVSFCKDLNSFVHGALAMITAVCGSGGTWHHEDFYYSNSPFQVTDCQTTGASRWPRCIYRGDICSKRICVACKNGQPVHFARPSVCGGP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GY27</accession>
    <name>A0A452GY27_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">PIPK domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GY27"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008176.1">
        <property type="protein sequence ID" value="ENSGAGP00000007059.1"/>
        <property type="gene ID" value="ENSGAGG00000005661.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016308">
        <property type="term" value="F:1-phosphatidylinositol-4-phosphate 5-kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046854">
        <property type="term" value="P:phosphatidylinositol phosphate biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd17306">
        <property type="entry name" value="PIPKc_PIP5K1A_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.800.10:FF:000001">
        <property type="entry name" value="phosphatidylinositol 4-phosphate 5-kinase type-1 gamma"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.810.10">
        <property type="entry name" value="2-Layer Sandwich"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.800.10">
        <property type="entry name" value="Phosphatidylinositol Phosphate Kinase II Beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027483">
        <property type="entry name" value="PInositol-4-P-4/5-kinase_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002498">
        <property type="entry name" value="PInositol-4-P-4/5-kinase_core"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027484">
        <property type="entry name" value="PInositol-4-P-5-kinase_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023610">
        <property type="entry name" value="PInositol-4/5-P-5/4-kinase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23086:SF54">
        <property type="entry name" value="PHOSPHATIDYLINOSITOL 4-PHOSPHATE 5-KINASE TYPE-1 ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23086">
        <property type="entry name" value="PHOSPHATIDYLINOSITOL-4-PHOSPHATE 5-KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01504">
        <property type="entry name" value="PIP5K"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00330">
        <property type="entry name" value="PIPKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56104">
        <property type="entry name" value="SAICAR synthase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51455">
        <property type="entry name" value="PIPK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="3" id="KW-0418">Kinase</keyword>
    <keyword evidence="3" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="PIPK" evidence="5">
        <location>
            <begin position="63"/>
            <end position="431"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="27"/>
            <end position="59"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="439"/>
            <end position="554"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="27"/>
            <end position="44"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="463"/>
            <end position="473"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="503"/>
            <end position="515"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="523"/>
            <end position="535"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="536"/>
            <end position="546"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00781"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS51455"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007059.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="554" mass="61161" checksum="A42AEA0DEBD9AF06" modified="2019-05-08" version="1">MASAGPESGGSSSSSAGVAGSATFKKSYTSEMSGSSGQPGSQTIKKGHRGVDSTGETTYKKTTSSALKGAIQLGITHTVGSLSTKPERDVLMQDFYVVESIFFPSEGSNLTPAHHYNDFRFKTYAPVAFRYFRELFGIRPDDYLYSLCSEPLIELSNSGASGSLFYVSSDDEFIIKTVQHKEAEFLQKLLPGYYMNLNQNPRTLLPKFYGLYCVQAGGKNIRIVVMNNLLPRSVKMHLKYDLKGSTYKRRASQKEREKVFPTYKDLDFMQDIPDGLFLDSDMYNALCKTLQRDCLVLQSFKIMDYSLLVAVHNIDQALRERAIMDGTSQSDTRRPATQKALYSTAMESIRGEARRGGTIETDDQMGGIPARNSKGERLLLYIGIIDVLQSYRFVKKLEHSWKALVHDGDTVSVHRPSFYAERFQRFMCNTAFKKIPLKLSPSKKSRSGTTVPRRPGQGGAPYQSHSSCETKAQVTAEADVEQGSHLGRPDLLPRTPPVDEANSDSMATTLSTSSMGSGGLNSPANSGSAGPSVTERSSELTEHFEDVQETEISF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GY28</accession>
    <name>A0A452GY28_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Cytochrome b5 heme-binding domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="4">Belongs to the cytochrome b5 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GY28"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008064.1">
        <property type="protein sequence ID" value="ENSGAGP00000006957.1"/>
        <property type="gene ID" value="ENSGAGG00000005604.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005783">
        <property type="term" value="C:endoplasmic reticulum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004128">
        <property type="term" value="F:cytochrome-b5 reductase activity, acting on NAD(P)H"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0020037">
        <property type="term" value="F:heme binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006801">
        <property type="term" value="P:superoxide metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.120.10">
        <property type="entry name" value="Cytochrome b5-like heme/steroid binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001199">
        <property type="entry name" value="Cyt_B5-like_heme/steroid-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036400">
        <property type="entry name" value="Cyt_B5-like_heme/steroid_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018506">
        <property type="entry name" value="Cyt_B5_heme-BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051872">
        <property type="entry name" value="Cytochrome_b5/Flavoprotein_Rdt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46237:SF1">
        <property type="entry name" value="CYTOCHROME B5 REDUCTASE 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46237">
        <property type="entry name" value="CYTOCHROME B5 REDUCTASE 4 FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00173">
        <property type="entry name" value="Cyt-b5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00363">
        <property type="entry name" value="CYTOCHROMEB5"/>
    </dbReference>
    <dbReference type="SMART" id="SM01117">
        <property type="entry name" value="Cyt-b5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55856">
        <property type="entry name" value="Cytochrome b5-like heme/steroid binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00191">
        <property type="entry name" value="CYTOCHROME_B5_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50255">
        <property type="entry name" value="CYTOCHROME_B5_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="1 4" id="KW-0349">Heme</keyword>
    <keyword evidence="3 4" id="KW-0408">Iron</keyword>
    <keyword evidence="2 4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Cytochrome b5 heme-binding" evidence="6">
        <location>
            <begin position="54"/>
            <end position="121"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="1"/>
            <end position="28"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="5">
        <location>
            <begin position="11"/>
            <end position="24"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="RuleBase" id="RU362121"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50255"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006957.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="121" mass="13399" checksum="A25401D295F02401" modified="2019-05-08" version="1">MLNVPAQAFPASGSQQRAAAGGRSKVPLKPGRSLMDWIRLTKSGKDLTGLKGRLIEVTEQELAKHSKKDDCWICIRGLVYNVTPYMEYHPGGDDELMRAAGTDGTDLFDQVRILELHFLTV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GY51</accession>
    <name>A0A452GY51_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">CCN family member 3</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="14">Cellular communication network factor 3</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="15">Protein NOV homolog</fullName>
        </alternativeName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell junction</location>
            <location evidence="2">Gap junction</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the CCN family.</text>
    </comment>
    <comment type="caution">
        <text evidence="16">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GY51"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008227.1">
        <property type="protein sequence ID" value="ENSGAGP00000007105.1"/>
        <property type="gene ID" value="ENSGAGG00000005703.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031012">
        <property type="term" value="C:extracellular matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005921">
        <property type="term" value="C:gap junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008083">
        <property type="term" value="F:growth factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045597">
        <property type="term" value="P:positive regulation of cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051239">
        <property type="term" value="P:regulation of multicellular organismal process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000046">
        <property type="entry name" value="Cellular communication network factor 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.70.10">
        <property type="entry name" value="Complement Module, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.20.100.10">
        <property type="entry name" value="Thrombospondin type-1 (TSP1) repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050941">
        <property type="entry name" value="CCN"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006207">
        <property type="entry name" value="Cys_knot_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006208">
        <property type="entry name" value="Glyco_hormone_CN"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009030">
        <property type="entry name" value="Growth_fac_rcpt_cys_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000867">
        <property type="entry name" value="IGFBP-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012395">
        <property type="entry name" value="IGFBP_CNN"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043973">
        <property type="entry name" value="TSP1_CCN"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000884">
        <property type="entry name" value="TSP1_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036383">
        <property type="entry name" value="TSP1_rpt_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001007">
        <property type="entry name" value="VWF_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11348:SF37">
        <property type="entry name" value="CONNECTIVE TISSUE GROWTH FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11348">
        <property type="entry name" value="CONNECTIVE TISSUE GROWTH FACTOR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00007">
        <property type="entry name" value="Cys_knot"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00219">
        <property type="entry name" value="IGFBP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19035">
        <property type="entry name" value="TSP1_CCN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00093">
        <property type="entry name" value="VWC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF036495">
        <property type="entry name" value="IGFBP_rP_CNN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00041">
        <property type="entry name" value="CT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00121">
        <property type="entry name" value="IB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00209">
        <property type="entry name" value="TSP1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00214">
        <property type="entry name" value="VWC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57184">
        <property type="entry name" value="Growth factor receptor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF82895">
        <property type="entry name" value="TSP-1 type 1 repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01185">
        <property type="entry name" value="CTCK_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01225">
        <property type="entry name" value="CTCK_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51323">
        <property type="entry name" value="IGFBP_N_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50092">
        <property type="entry name" value="TSP1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01208">
        <property type="entry name" value="VWFC_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50184">
        <property type="entry name" value="VWFC_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0965">Cell junction</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="8" id="KW-0303">Gap junction</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10" id="KW-0339">Growth factor</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0964">Secreted</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
    <feature type="domain" description="IGFBP N-terminal" evidence="19">
        <location>
            <begin position="5"/>
            <end position="76"/>
        </location>
    </feature>
    <feature type="domain" description="VWFC" evidence="18">
        <location>
            <begin position="79"/>
            <end position="145"/>
        </location>
    </feature>
    <feature type="domain" description="CTCK" evidence="17">
        <location>
            <begin position="224"/>
            <end position="298"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004610"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00008125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022868"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00039944"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00042352"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00077787"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00039"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS01225"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50184"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS51323"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007105.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="322" mass="36045" checksum="00C643116CE31614" modified="2019-05-08" version="1">TQKIRSQLCSYPCQCPSQPLQCPVGTSHVWDACGCCKVCARQLGELCSLQTPCDHHKGLHCDFSKIHRDVGICLVHEGATCDLLGKIYHNGESFQPTCKLQCICMDGAIGCIPLCADDLQLPSPACPNPLRVRLPAQCCEEWICQGGSQETALAAPRSELNDLQENCLVQTTKWSACSRSCGMGISTRITNDNPQCRLEKESRLCMVRPCDFPREKIFKKGKKCTRTPKSRRGIRFEFSGCTSVRSYRPKFCGSCTDGRCCTPHTTSTAGVEFQCPEGDFFHRKLMFIKTCSCHHDCPRDNDIFLATYRRRMAGDHVKIERQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GY58</accession>
    <name>A0A452GY58_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">RING-CH-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endomembrane system</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Endosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Lysosome membrane</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GY58"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008173.1">
        <property type="protein sequence ID" value="ENSGAGP00000007056.1"/>
        <property type="gene ID" value="ENSGAGG00000005628.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005768">
        <property type="term" value="C:endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005765">
        <property type="term" value="C:lysosomal membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002376">
        <property type="term" value="P:immune system process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011016">
        <property type="entry name" value="Znf_RING-CH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45981">
        <property type="entry name" value="LD02310P"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12906">
        <property type="entry name" value="RINGv"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00744">
        <property type="entry name" value="RINGv"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57850">
        <property type="entry name" value="RING/U-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51292">
        <property type="entry name" value="ZF_RING_CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0391">Immunity</keyword>
    <keyword evidence="8" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5" id="KW-0863">Zinc-finger</keyword>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="70"/>
            <end position="92"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="112"/>
            <end position="132"/>
        </location>
    </feature>
    <feature type="domain" description="RING-CH-type" evidence="9">
        <location>
            <begin position="1"/>
            <end position="54"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004127"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004177"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004656"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022859"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51292"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007056.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="206" mass="23874" checksum="FA11993AAC972923" modified="2019-05-08" version="1">RLESSLVDRDEESPLITPCHCTGSHFVHQACLQQWIKSSDTRYCELCKYEFIMETKLKKLQMTASERRKIMCSVTFHIIAITCVVWSLYVLIDRTAEEIKQGQTTGILEWPFWTKLVVVAIGFTGGLLFMYVQCKVYVQLWKRLKAYNRVIYLQNCPETSKKSIFEKPALMEPNTENKEMLGVHHSDTNSSHYTEPEDCGAEILHV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GY59</accession>
    <name>A0A452GY59_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">UBA domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum membrane</location>
            <topology evidence="1">Single-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000008095.1">
        <property type="protein sequence ID" value="ENSGAGP00000006987.1"/>
        <property type="gene ID" value="ENSGAGG00000005610.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019894">
        <property type="term" value="F:kinesin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007018">
        <property type="term" value="P:microtubule-based movement"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024854">
        <property type="entry name" value="Kinectin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007794">
        <property type="entry name" value="Rib_rcpt_KP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015940">
        <property type="entry name" value="UBA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18864">
        <property type="entry name" value="KINECTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18864:SF1">
        <property type="entry name" value="KINECTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05104">
        <property type="entry name" value="Rib_recp_KP_reg"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50030">
        <property type="entry name" value="UBA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="5 8" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 8" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 8" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="7"/>
            <end position="29"/>
        </location>
    </feature>
    <feature type="domain" description="UBA" evidence="9">
        <location>
            <begin position="744"/>
            <end position="790"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="48"/>
            <end position="194"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="307"/>
            <end position="370"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="407"/>
            <end position="498"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="566"/>
            <end position="1050"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="1116"/>
            <end position="1252"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="73"/>
            <end position="86"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="7">
        <location>
            <begin position="113"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="123"/>
            <end position="135"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="7">
        <location>
            <begin position="161"/>
            <end position="171"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="172"/>
            <end position="181"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004389"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50030"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006987.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1338" mass="153131" checksum="3631284C3E337806" modified="2019-05-08" version="1">MEFYESTYFIILIPSVVITVIFLFFWLFMKETLYDEVLAKQKRDQKFLPSKMDKKKTEKKKNKRKDAQNGNLHESDSESTPREFKLSDAMIADEEPILPVPLGLTEASSSVRERKKKEKKHKTVQEEHVSKESEGSKPAGKKIEPVPVTKQPTPPSEAAGSKKKPGQKKQKNGIDDQDSKSESFTSPTKKQDPLLFQHEIKQESGSGKKKVSTKKQKIDNVAALVDEPLIHATTYIPLMDNSDLNPLAEKREVADVGKQDTTEMIQKSGAKKLKNETDKENAEVKFKDFLLAMKSMIFTEDEALCVVELLKEKSSVIQDALQKASKEDSAATMHQLQEKDKMLAALKEEAAITKDQCKQLTQELLAEKERSNLVTTKMRERINVLEKEHGTFQSKIHVSYQENQGMKMKFQQLREQMEAEIGHLKQENGILRDAVSTSTNQMESKQSSELNKLRQDCARLVNELTEKNNKLQQEELQKKNAEQTIGQLKVQQQEAERRWEEIQAYLRKRIAEHEAAQQDVQNKLVAKDSEVQSLHSKLTDTIVSKQQLEQRMLQLMDAEQKRVTAEDSVQLQVQELMEQNEALKAQLQKFHSQMAAQSSASVLAEELQKVIAEKDKQIKQVEDSLVNEHANLAGKEEELKVIQNMNLSLKAEVQKLQALTNEQAAAAHELERMQKSIHIKDDKIRALEEQLQGQLAQVSNTREEFKVLKDQNKALQSEVQKLQALLSEQANKELLEQMERSITEKDEKIKTVEELLEAGLIQVANREEELKALRAENSSLKKEVQNLQIQQSEQVSFESVVEELQKVIREKDGKIKLVEELLEAEVLKVTNKEKTVQAFTQEIEALKEEVGHSQLEMKKQVSIASQVNDLQNLLKGKEEQMKTMEALLEEKAKDIAKKGESLQSQQDTITHLTSKVQELEQQNSEQLQQVSTVSQTQDLEILLKEKEEEVKKMEAVLEERGREITNKGKQLQEAQKENELFKMQIQELKQETHKQASLAVQSDELLQVIAGKETEITSLQNELGSLKNAVEQQRKKNNDLREKNWKAMEALASTEKLLQDKVNKTAKERQQYLETVEMETRELLQKLFPKVSLPSNLSHKEWIDGFEKMASEYLQEASGSEEVKAMEQKLKEAEEMHIMLQLECEKYKAVLAETEGILQRLQRSVEEEESKWKIKIEESQKELRQMHSSVGSLEHEIERLKEEIKEAETLKREREHLESELEKAEIERSTYVSEVRELKTQLNDTLAKLKIDQSERQKVAGDLPKAQESLASLEKEIGKAAGDANVIENSDGCSEPELTEKRLNLAVTLSQDVGHLKKLVVSISQMLSKGKEHYQLVE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GY72</accession>
    <name>A0A452GY72_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">small monomeric GTPase</fullName>
            <ecNumber evidence="2">3.6.5.2</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="6">
            <text>GTP + H2O = GDP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:19669"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:37565"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:58189"/>
            <dbReference type="EC" id="3.6.5.2"/>
        </reaction>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the small GTPase superfamily. Ras family.</text>
    </comment>
    <dbReference type="EC" id="3.6.5.2" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452GY72"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008180.1">
        <property type="protein sequence ID" value="ENSGAGP00000007063.1"/>
        <property type="gene ID" value="ENSGAGG00000005672.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003925">
        <property type="term" value="F:G protein activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005525">
        <property type="term" value="F:GTP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051065">
        <property type="entry name" value="Ras-related_GTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001806">
        <property type="entry name" value="Small_GTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45704">
        <property type="entry name" value="RAS-LIKE FAMILY MEMBER 11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00071">
        <property type="entry name" value="Ras"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00175">
        <property type="entry name" value="RAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00173">
        <property type="entry name" value="RAS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00174">
        <property type="entry name" value="RHO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51419">
        <property type="entry name" value="RAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51421">
        <property type="entry name" value="RAS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0342">GTP-binding</keyword>
    <keyword evidence="4" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="3" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00008344"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00011984"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00048098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007063.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="255" mass="28572" checksum="BA6CC2EE25DE9A78" modified="2019-05-08" version="1">MSSMFGKPRATPSTTDRHLPSLLPECNVAILGCRGSGKSEDTYMSEETVDQQPVLMKVMDTADQDVPVNCERYLNWANAFMVVYSIDNKRSFEGCHQYLDTISLHAKGIQHDYPILLLGNKLDLEQYRQVTKAEGVSLAAKYGCLFYEVSACLEFEAVQHVFHEAAREVRREMERNLPVRPLFITEERPFLHLAAPTAVASKHGLASCTLNTLSTVNYKEIPSVAQAKLVTVKSSRAQSKRKAPTLTLLKGFKIF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GY73</accession>
    <name>A0A452GY73_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">GRB10 interacting GYF protein 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GY73"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008229.1">
        <property type="protein sequence ID" value="ENSGAGP00000007107.1"/>
        <property type="gene ID" value="ENSGAGG00000001575.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043204">
        <property type="term" value="C:perikaryon"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1990635">
        <property type="term" value="C:proximal dendrite"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031982">
        <property type="term" value="C:vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048009">
        <property type="term" value="P:insulin-like growth factor receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051640">
        <property type="entry name" value="GRB10-interact_GYF"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14445">
        <property type="entry name" value="GRB10 INTERACTING GYF PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14445:SF38">
        <property type="entry name" value="GRB10-INTERACTING GYF PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="97"/>
            <end position="234"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="299"/>
            <end position="473"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="563"/>
            <end position="583"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="139"/>
            <end position="170"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="180"/>
            <end position="199"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="213"/>
            <end position="226"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="1">
        <location>
            <begin position="306"/>
            <end position="323"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="325"/>
            <end position="349"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="361"/>
            <end position="379"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="384"/>
            <end position="402"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="442"/>
            <end position="459"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007107.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="4">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="583" mass="64441" checksum="E3B3EC66771576C6" modified="2025-10-08" version="2">MAALRALSSGGSVTSPPLSPALPKYKLADYRYGREEMLALFLKDNKIPSDLLDKEFQPILQEEPLPPLALVPFTEEEQRNFSMSVNSAAVLRLTGRGGGTVVGAPRGRSSSRGRGRGRGESGFYQRSFDEVEGGFGRGGGREMHRSQSWEERGDRRFEKPGRKDPVRPNFEEGGTTTTGRKHEFIRSESENWRIFREEQNGEDDDGGWRLAGPRRDGERWRPHSPDGPRSAGWREPMERRRRFEFDFRDRDDERGYRRVRCGSGSMEDDRDSLPEWCLEDAEEEMGTFDSSGAFLSLKKVQKESIPEEQEMDFQPVEGEEPSDPEGGHGEEAKELEKTTKREGDKRDTTTAEAAEEAVRASSPSIRSGTPPKLQSQEPLQASLFERKEECVPERTEKMEAKESQTVNTLPSKVPSRGEDLTPAVQPLPQISPATVSPVHLSSPLSNTNPALKPLQTSVSAAPGMGNISADPDDEEGLKHLEQQAEKMVAYLQDSALDDERLAAKLQEHRAKGSSMPLLHEAMQKWYYKDPQGEIQAPPVPAVFDTTTVCTGVGTAAKSSPLIPAAAATGSPLTAVPGPEDENI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GY86</accession>
    <name>A0A452GY86_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">EF-hand domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GY86"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008244.1">
        <property type="protein sequence ID" value="ENSGAGP00000007122.1"/>
        <property type="gene ID" value="ENSGAGG00000005720.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005743">
        <property type="term" value="C:mitochondrial inner membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061891">
        <property type="term" value="F:calcium ion sensor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00051">
        <property type="entry name" value="EFh"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.238.10:FF:000112">
        <property type="entry name" value="EF-hand domain family, member D2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.238.10">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011992">
        <property type="entry name" value="EF-hand-dom_pair"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002048">
        <property type="entry name" value="EF_hand_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040365">
        <property type="entry name" value="EFHD1/2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13025">
        <property type="entry name" value="EF-HAND DOMAIN-CONTAINING PROTEIN D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13025:SF5">
        <property type="entry name" value="EF-HAND DOMAIN-CONTAINING PROTEIN D1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13499">
        <property type="entry name" value="EF-hand_7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47473">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50222">
        <property type="entry name" value="EF_HAND_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0106">Calcium</keyword>
    <keyword evidence="4" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="1" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="EF-hand" evidence="5">
        <location>
            <begin position="3"/>
            <end position="27"/>
        </location>
    </feature>
    <feature type="domain" description="EF-hand" evidence="5">
        <location>
            <begin position="28"/>
            <end position="63"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="4">
        <location>
            <begin position="104"/>
            <end position="131"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50222"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007122.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="141" mass="16082" checksum="60427E4B8D81DAF3" modified="2019-05-08" version="1">RFKYDSGKDGYIDLMELKLMMEKLGAPQTHLGLKNMIKEVDEDFDGKLSFREFLLIFHKAAAGELEEDSGLMTLAKLSEIDVSTEGVKGAKNFFEAKVQALSSASKFEAEIKAEQDERKREEEEKKHRRAAFRELKSAFSQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GY89</accession>
    <name>A0A452GY89_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="6">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the tropomodulin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GY89"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008205.1">
        <property type="protein sequence ID" value="ENSGAGP00000007086.1"/>
        <property type="gene ID" value="ENSGAGG00000005690.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000008208.1">
        <property type="protein sequence ID" value="ENSGAGP00000007088.1"/>
        <property type="gene ID" value="ENSGAGG00000005690.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005865">
        <property type="term" value="C:striated muscle thin filament"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003779">
        <property type="term" value="F:actin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005523">
        <property type="term" value="F:tropomyosin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007015">
        <property type="term" value="P:actin filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006936">
        <property type="term" value="P:muscle contraction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030239">
        <property type="term" value="P:myofibril assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051694">
        <property type="term" value="P:pointed-end actin filament capping"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000006">
        <property type="entry name" value="Tropomodulin 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004934">
        <property type="entry name" value="TMOD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10901">
        <property type="entry name" value="TROPOMODULIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10901:SF9">
        <property type="entry name" value="TROPOMODULIN-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03250">
        <property type="entry name" value="Tropomodulin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52047">
        <property type="entry name" value="RNI-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="5" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009345"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007088.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="342" mass="38804" checksum="71A517C0D7CB8364" modified="2019-05-08" version="1">MTSYRMELEKYRDIDEEKLLRELSPEELDQLDLELQEMDPENMMLPAGLRQRDQTKKSPTGPLDREALMQHLEKQALEVKEREDLVPFTGEKRGKPFIPKNPQREIPKEEQITLEPELEEALANATEAEMCDIAAILGMYTLMSNKQYYEALCSGNITSTEGINSVVKPDRYKPVPDEPPNPTNVEETLKKIQSNDKDLEEVNLNNIKDIPIPKLKEICEAMKTNTHVKKLSLVATRSNDPVAKAVAEMLKENKTLQSLNIESNFITSAGMMAIIKGMQQNTSLAELKVDNQSQRLGDSVEMEMAAMLEKCPSIIRFGYHFTQQGPRARASIAISKNNELRK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9Q0</accession>
    <name>A0A452H9Q0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Dynein regulatory complex protein 1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="8">Coiled-coil domain-containing protein 164</fullName>
        </alternativeName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Component of the nexin-dynein regulatory complex (N-DRC) a key regulator of ciliary/flagellar motility which maintains the alignment and integrity of the distal axoneme and regulates microtubule sliding in motile axonemes. Plays a critical role in the assembly of N-DRC and also stabilizes the assembly of multiple inner dynein arms and radial spokes. Coassembles with CCDC65/DRC2 to form a central scaffold needed for assembly of the N-DRC and its attachment to the outer doublet microtubules.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Flagellum axoneme</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the DRC1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9Q0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013221.1">
        <property type="protein sequence ID" value="ENSGAGP00000011546.1"/>
        <property type="gene ID" value="ENSGAGG00000008764.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005858">
        <property type="term" value="C:axonemal dynein complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070286">
        <property type="term" value="P:axonemal dynein complex assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060285">
        <property type="term" value="P:cilium-dependent cell motility"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003352">
        <property type="term" value="P:regulation of cilium movement"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039505">
        <property type="entry name" value="DRC1/2_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039750">
        <property type="entry name" value="DRC1/DRC2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029440">
        <property type="entry name" value="DRC1_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21625:SF1">
        <property type="entry name" value="DYNEIN REGULATORY COMPLEX PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21625">
        <property type="entry name" value="NYD-SP28 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14772">
        <property type="entry name" value="NYD-SP28"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14775">
        <property type="entry name" value="NYD-SP28_assoc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0966">Cell projection</keyword>
    <keyword evidence="6" id="KW-0969">Cilium</keyword>
    <keyword evidence="5" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0282">Flagellum</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Dynein regulatory complex protein 1/2 N-terminal" evidence="11">
        <location>
            <begin position="60"/>
            <end position="161"/>
        </location>
    </feature>
    <feature type="domain" description="Dynein regulatory complex protein 1 C-terminal" evidence="12">
        <location>
            <begin position="579"/>
            <end position="638"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1"/>
            <end position="38"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="11"/>
            <end position="38"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004611"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009688"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00013815"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022846"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023069"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00031554"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00046115"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF14772"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF14775"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011546.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="659" mass="77058" checksum="FA6D9E37F9D3A791" modified="2019-05-08" version="1">VTKVFLLSTSPHREALGEDPDAKKAEAEELSRSHKQIEESRQRLAKLLNDGTQLVTNIQVAADARETQRRAEEDELKRQRLEKLENESKTNQDKFEEITSKWGSAKEKTIPQDLWDVLNQQQQHCALLIEEKNKLISELQQELKSKDDQYVKDLKKQSDDINLLVDRMEEQIRNLMKTYRRELTQIEKAFELERRELLSSNKKKWEQAMQAHNAQELENLITRMQKVEEYEKQLNQLRVQDAEEYNTIKIQLENDVQILEQQLQQMKAIYQLNQEKLEYNFQVLKKRDEENTIIKSQQKRKLNRLHDVLNALRMKLDKQIKQFQEENQTLTADYKRIVEQYKDLQRTMRHFAVVDAQKFLEVWLMNEEEAKGLMWKALKADRIIHTQQLGLPWVEPDYWFLDNVGPIVHHMAIRSASKLAREVMDAVEEGRKEAAAEAGEEVKPPRHISVKSVKHILELLCDESGFLIESKLLGLLQPLERHERSLMRLDAIFAALGIDNEDDVYTLVDFFLKFNAQQVVSSQGKQWAEDQCPEELAVDAADSDEGSAALKEEGTAHQSWAMSRVKNMMEERDDSKDSEYWKALAHVIPEPKLKVWDALEAALEKYYSVLSQRSKLLTDLDGLQQQNTELHMLLHQYLTSKVNEELLIPPTHVIELMPP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9Q2</accession>
    <name>A0A452H9Q2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Palmitoyltransferase</fullName>
            <ecNumber evidence="7">2.3.1.225</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="7">
            <text>L-cysteinyl-[protein] + hexadecanoyl-CoA = S-hexadecanoyl-L-cysteinyl-[protein] + CoA</text>
            <dbReference type="Rhea" id="RHEA:36683"/>
            <dbReference type="Rhea" id="RHEA-COMP:10131"/>
            <dbReference type="Rhea" id="RHEA-COMP:11032"/>
            <dbReference type="ChEBI" id="CHEBI:29950"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57379"/>
            <dbReference type="ChEBI" id="CHEBI:74151"/>
            <dbReference type="EC" id="2.3.1.225"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="domain">
        <text evidence="7">The DHHC domain is required for palmitoyltransferase activity.</text>
    </comment>
    <comment type="similarity">
        <text evidence="7">Belongs to the DHHC palmitoyltransferase family.</text>
    </comment>
    <dbReference type="EC" id="2.3.1.225" evidence="7"/>
    <dbReference type="AlphaFoldDB" id="A0A452H9Q2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000012971.1">
        <property type="protein sequence ID" value="ENSGAGP00000011327.1"/>
        <property type="gene ID" value="ENSGAGG00000008762.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019706">
        <property type="term" value="F:protein-cysteine S-palmitoyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001594">
        <property type="entry name" value="Palmitoyltrfase_DHHC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039859">
        <property type="entry name" value="PFA4/ZDH16/20/ERF2-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12246">
        <property type="entry name" value="PALMITOYLTRANSFERASE ZDHHC16"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01529">
        <property type="entry name" value="DHHC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50216">
        <property type="entry name" value="DHHC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6 7" id="KW-0012">Acyltransferase</keyword>
    <keyword evidence="5 7" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 7" id="KW-0808">Transferase</keyword>
    <keyword evidence="3 7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 7" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="43"/>
            <end position="67"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="73"/>
            <end position="92"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="171"/>
            <end position="194"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="214"/>
            <end position="235"/>
        </location>
    </feature>
    <feature type="domain" description="Palmitoyltransferase DHHC" evidence="8">
        <location>
            <begin position="128"/>
            <end position="254"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023315"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU079119"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF01529"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011327.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="299" mass="34312" checksum="83E66474AA6FB70A" modified="2019-05-08" version="1">MMITPIHRFRDIERRPEYLQPDKCVPPPSQSSLGTMWFIRDGCGIACAVVTWMLVFYADFVVLLVMLVPSRDYIYSVINGILFNMLAFLALASHFRAMLSDPGAVPKGNATKEFIESLQLKPGQVVYKCPKCCSIKPDRAHHCSVCKRCIRKMDHHCPWVNNCVGENNQKYFVLFTMYIALISMHALIMVGFHFLYCFEEDWTKCSSFSPPTTVILLILLCFEALLFLIFTSVMFGTQVHSICTDETGIEQLKKEERRWAKKTKWMNMKAVFGHPFSIAWLSPFATPDQGKADPYQYVV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9Q4</accession>
    <name>A0A452H9Q4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">non-specific serine/threonine protein kinase</fullName>
            <ecNumber evidence="2">2.7.11.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="10">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="12">Belongs to the protein kinase superfamily. CK1 Ser/Thr protein kinase family.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452H9Q4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013159.1">
        <property type="protein sequence ID" value="ENSGAGP00000011492.1"/>
        <property type="gene ID" value="ENSGAGG00000008859.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015630">
        <property type="term" value="C:microtubule cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000167">
        <property type="entry name" value="Tau tubulin kinase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000358">
        <property type="entry name" value="Tau tubulin kinase 2b"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050235">
        <property type="entry name" value="CK1_Ser-Thr_kinase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11909">
        <property type="entry name" value="CASEIN KINASE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9 13" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0418">Kinase</keyword>
    <keyword evidence="7 13" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="6" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Protein kinase" evidence="15">
        <location>
            <begin position="34"/>
            <end position="297"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="546"/>
            <end position="672"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="569"/>
            <end position="583"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="617"/>
            <end position="636"/>
        </location>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <position position="63"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00012513"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00047899"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00048679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00061588"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011492.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="751" mass="85187" checksum="11EC4222252DF8AB" modified="2019-05-08" version="1">MQCLVAAIREEPNMSGGGEQVDILPANYVVKDRWKVLKKIGGGGFGEIYEAMDLLTRENVALKVESAQQPKQVLKMEVAVLKKLQGKDHVCRFIGCGRNEKFNYVVMQLQGRNLADLRRSQPRGTFTLSTTLRLGKQILESIEAIHSVGFLHRDIKPSNFSMGRLPSTYRKCYMLDFGLARQYTNTNGEVRPPRNVAGFRGTVRYASVNAHKNREMGRHDDLWSLFYMLVEFAVGQLPWRKIKDKEQVGMIKEKYEHRMLLKHMPSEFHLFLDHIANLDYFTKPDYQLIMSVFENSMKERGITENEAFDWEKAGTDILLSTSTSTPPQQNTRQTAAMFGVVNVTPVPGDMLRENTEDVLQGEHLSDQENAPPILPGRPVEGLGQAPNMAFNETEVWEETDVNRNKLRISISKTQCMMEEEQRSAVCPSSPIRAPPDSPTTQVRSLRYRRVNSPESERLSIADGKADLHERRSRVDLPCSPSRLVCSSQPAQMLSIDTGQVDRQASGRMDVSASVEHEALSNAFRSVPLAEEEDFDSKEWVIIDKETELKDFHPGAEPSTSGTTDEEPEELRPIEEGEERRRLGAEAAVRPKVHDGRTWGMQPVTEEDSSHRHEGYCQTVSDTKPEQQPGSPAQLSLHSAPALRQHRRESEPTGPQRQLEEDRHSQHSLPRYSPLRRLASSVFSSSALETEHYPQLGGTFIQRSRSAESSPVRMPHRRHMPLMAGNHRLMPSVLRISRSQLQQVWARFTHKT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9Q6</accession>
    <name>A0A452H9Q6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">mitogen-activated protein kinase</fullName>
            <ecNumber evidence="3">2.7.11.24</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the protein kinase superfamily. CMGC Ser/Thr protein kinase family. MAP kinase subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.24" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452H9Q6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013170.1">
        <property type="protein sequence ID" value="ENSGAGP00000011501.1"/>
        <property type="gene ID" value="ENSGAGG00000008872.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004707">
        <property type="term" value="F:MAP kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000063">
        <property type="entry name" value="Mitogen-activated protein kinase 14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000769">
        <property type="entry name" value="Mitogen-activated protein kinase 14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050117">
        <property type="entry name" value="MAPK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008352">
        <property type="entry name" value="MAPK_HOG-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24055">
        <property type="entry name" value="MITOGEN-ACTIVATED PROTEIN KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01773">
        <property type="entry name" value="P38MAPKINASE"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9 11" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="8" id="KW-0418">Kinase</keyword>
    <keyword evidence="7 11" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="10" id="KW-0346">Stress response</keyword>
    <keyword evidence="6" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Protein kinase" evidence="12">
        <location>
            <begin position="23"/>
            <end position="291"/>
        </location>
    </feature>
    <feature type="binding site" evidence="11">
        <location>
            <position position="53"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008832"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012411"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023016"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011501.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="345" mass="39982" checksum="EE8C46ED499EB542" modified="2019-05-08" version="1">MSERGGFYKQELNKTVWEVPQRYQNLTPVGSGAYGSVCSTYDTKTRQKVAVKKLSRPFQSLIHARRTYRELRLLKHMKHENVIGLLDVFTPATSIENFNEVYLVTNLMGADLNNIVKCQKLTDDHIQFLIYQLLRGLKYIHSAGIIHRILDFGLARQTDDEMTGYVATRWYRAPEIMLNWMHYNQTVDIWSVGCIMAELLKGKALFPGNDYIDQLKRIMEVVGTPSSELLKKISSEHARKYIESLPHMPQQDLKAVFRGANPLAVDLLEKMLILDSDKRITASEALAHAYFVQYHDPDDEPEAELYDESIENKERTIDEWKEITYEEMISFKPPDLKMDSLEIEQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9Q9</accession>
    <name>A0A452H9Q9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">Fibronectin type-III domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452H9Q9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013166.1">
        <property type="protein sequence ID" value="ENSGAGP00000011497.1"/>
        <property type="gene ID" value="ENSGAGG00000008869.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47135">
        <property type="entry name" value="FIBRONECTIN TYPE III DOMAIN-CONTAINING PROTEIN 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47135:SF1">
        <property type="entry name" value="FIBRONECTIN TYPE III DOMAIN-CONTAINING PROTEIN 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="5"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Fibronectin type-III" evidence="1">
        <location>
            <begin position="7"/>
            <end position="94"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="1">
        <location>
            <begin position="95"/>
            <end position="181"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="1">
        <location>
            <begin position="182"/>
            <end position="267"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="1">
        <location>
            <begin position="268"/>
            <end position="352"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="1">
        <location>
            <begin position="436"/>
            <end position="523"/>
        </location>
    </feature>
    <evidence type="ECO:0000259" key="1">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011497.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="711" mass="75504" checksum="F98EFCE3E6D9A8BA" modified="2019-05-08" version="1">MHLAPNTPEIPVIDQAYSKLSNSITVEWVAVPGATSYLLTAQDGDSSIETIVINSPGTVTGLKAATLYKITIRSINSGGRSLPSPSKTAKTVLAAPVLSVSSPSYDSIVVSWGTVYMAAGFSVSIMRSDGLGSMMKENTTHTSLVFTSLDPGTLYTIKAHAWNVNGIPGDDFTYNQITSPRAPSDVQVAFDSGTLRATVSWIPTEGTLNYSVTACSEASKLNCSTSSTSCPLSSLQCGSEYSVYVVANNSAGSSKPTDAVSLKTVPCAPRSITIEEDNPGKLSVSWSNVDLGDYYVVFVKSDDGLEVHCNTSHTQCHFQSDCGFTYFISVFAYNKAGQSPLGDIFNYTTSPCCPSDFYPVFVSSDTVEIVWSPVRGAEMYETKAVDGVGVVLCHDTATICILSALQCNTQYNITVYSYSEIRGTNTSCVSKYVATAPCSPEIKNISQDDLSIIRVHWQANNDDAVYTVTARGEVGLWHCTSSGNSCSITGLPCGSVFSVSAVAITTAGQSLPSYSVPLETAPCCPNDLTVTQVMQSITNISWSVGAGARTYITILESPKGQAKCHTLQNHCLLGCITCGTNYTVSLKVITETGLASYCTYQGYSSSACCPSGVKLYRLGNNGIRIYWQASAGLISTDLYGSKGNFTCTPTSGLSYCDVTEIPCGDVYTVVVSPVPVKGSKLTFCPKKIYSVTCSGSSVGMGTRMLSVFHLQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9V0</accession>
    <name>A0A452H9V0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Anoctamin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="8">Membrane</location>
            <topology evidence="8">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 8">Belongs to the anoctamin family.</text>
    </comment>
    <comment type="caution">
        <text evidence="8">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9V0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013027.1">
        <property type="protein sequence ID" value="ENSGAGP00000011379.1"/>
        <property type="gene ID" value="ENSGAGG00000007328.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005229">
        <property type="term" value="F:intracellularly calcium-gated chloride channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046983">
        <property type="term" value="F:protein dimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032394">
        <property type="entry name" value="Anoct_dimer"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007632">
        <property type="entry name" value="Anoctamin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049452">
        <property type="entry name" value="Anoctamin_TM"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12308">
        <property type="entry name" value="ANOCTAMIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12308:SF28">
        <property type="entry name" value="ANOCTAMIN-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16178">
        <property type="entry name" value="Anoct_dimer"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04547">
        <property type="entry name" value="Anoctamin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6 8" id="KW-0472">Membrane</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 8" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 8" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="187"/>
            <end position="213"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="263"/>
            <end position="282"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="339"/>
            <end position="365"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="385"/>
            <end position="406"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="434"/>
            <end position="455"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="554"/>
            <end position="576"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="604"/>
            <end position="628"/>
        </location>
    </feature>
    <feature type="domain" description="Anoctamin dimerisation" evidence="11">
        <location>
            <begin position="1"/>
            <end position="173"/>
        </location>
    </feature>
    <feature type="domain" description="Anoctamin transmembrane" evidence="10">
        <location>
            <begin position="176"/>
            <end position="757"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="756"/>
            <end position="783"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009671"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="RuleBase" id="RU280814"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="Pfam" id="PF04547"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF16178"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011379.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="792" mass="92754" checksum="DC7E7072C5972BF0" modified="2019-05-08" version="1">MEKESSLTNSDIIFVKLHAPWEVLGRYAEVMNVRMPFRIERQISRFQGWLPRKPMRLDKETLPDLEENDCYTAPFSQQRIHHFIIHNKDTFFNNATRSRIVHHILQRVKYEEGKNKIGLNRLLTNGSYEAAFPLHEGSYRSKNSIRTHGAENHRHLLYECWASWGMWYKYQPLDLVRRYFGEKIGLYFAWLGWYTGMLFPAAFIGLFVFLYGVTTLNHCQVSKEVCQATDIIMCPICDKFCPFMRLSDSCVYAKVTHLFDNGATVFFAVFMAVWATVFLEFWKRRRAVIAYDWDLIDWEEEEEEIRPQFEAKYSKKERMNPISGKPEPYQAFADKCSRLIVSASGIFFMICVVIAAVFGIVIYRVVTVSTFAAFKWALIRNNSQVATTGTAVCINFCIIMLLNVLYEKVALLLTNLEQPRTESEWENSFTLKMFLFQFVNLNSSTFYIAFFLGRFTGHPGAYLRLINRWRLEECHPSGCLIDLCMQMGIIMVLKQTWNNFMELGYPLIQNWWTRRKLYQEHGSQRKTSFPQWEKDYNLQPMNAYGLFDEYLEMILQFGFTTIFVAAFPLAPLLALLNNIIEIRLDAYKFVTQWRRPLASRAKDIGIWYGILEGIGILSVITNAFVVAVTSDFIPRLVYAYKYGPCAGQGEAGQKCMVGYVNASLSVFLVSDFENRSEPTSNGSEFSGSPLKYCRYRDYRDPPHSPVPYGYTLQFWHVLAARLAFIIVFEHLVFCIKHLISYLIPDLPKDLRDRMRREKYLIQEMMYEAELERLQKERKERKKNGKSHHNEWP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9V1</accession>
    <name>A0A452H9V1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">DUF4200 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452H9V1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013229.1">
        <property type="protein sequence ID" value="ENSGAGP00000011554.1"/>
        <property type="gene ID" value="ENSGAGG00000008918.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051147">
        <property type="entry name" value="CFAP_domain-containing"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025252">
        <property type="entry name" value="DUF4200"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21683:SF3">
        <property type="entry name" value="CILIA AND FLAGELLA ASSOCIATED PROTEIN 100"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21683">
        <property type="entry name" value="COILED-COIL DOMAIN-CONTAINING PROTEIN 42 LIKE-2-LIKE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13863">
        <property type="entry name" value="DUF4200"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1 2" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="DUF4200" evidence="4">
        <location>
            <begin position="160"/>
            <end position="278"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="1"/>
            <end position="57"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="295"/>
            <end position="322"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="340"/>
            <end position="378"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="2">
        <location>
            <begin position="74"/>
            <end position="136"/>
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    </feature>
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        <location>
            <begin position="174"/>
            <end position="204"/>
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        <location>
            <begin position="417"/>
            <end position="483"/>
        </location>
    </feature>
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        <location>
            <begin position="549"/>
            <end position="599"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="27"/>
            <end position="49"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="365"/>
            <end position="378"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="Pfam" id="PF13863"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011554.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="633" mass="73348" checksum="95D073F4106EFBFB" modified="2019-05-08" version="1">MSVLSCTRSTGSQETSSSGAISKARTLPQQTVPTSGIPFTNSTVETKSPLSDDDEFQLRKNPFKIPSDIDIFLLRDKERERAKAERERKKTLKVHEKMTYSTMMNAKQSGFKKELQKEEEAEDKELAAEAQRLKALQESISWKIAVTKDQQVEKETLHEYIDKKRQMFLLQYAVAIKRDEIQKLENLAAEEEAKLEKAEQYLEKDAAMFDEFLKENDKNSAQALKTAEKETKAKIEKIIEIRELTAQMMSIQSEISRFEDTLKEYKKYKNFLYQLSPKEWREEYEKKQMKTKAMKMSSEVKEEKISVSPITPKGQDSTPRTRCGKLPYLGYTGCQSVDFSSGGPDLRAPSQITQRRESLRPGKSPSKQNLKSQQTRRQSIAALAMEEKASTTFSDNEDEDEEPELYFTDPQQLLHIFTELEEQNLSLIQNSQETEETLDELQHTLTTTRSKMDREIKQLKQLAATLKASIIKEEETAADLELKARVFSFGEYKADVQDKMLVSLNKKVTEVYRRCIGENEANLGTLQMLTVIEHQLDDLLECLERVPQAKIEQAEKAKEKERRMRMRDEKVRQQRQLQEERLQRALARAQADIKKKTGRKLMFRSEPAPIKEKEDEDQGLIDQEKEEALYYFT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9V4</accession>
    <name>A0A452H9V4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">IF rod domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="3 4">Belongs to the intermediate filament family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9V4"/>
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        <property type="protein sequence ID" value="ENSGAGP00000011390.1"/>
        <property type="gene ID" value="ENSGAGG00000008332.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="C:keratin filament"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
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        <property type="term" value="F:structural constituent of skin epidermis"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045109">
        <property type="term" value="P:intermediate filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="P:keratinization"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="entry name" value="Keratin type II"/>
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        <property type="entry name" value="Type II keratin 23"/>
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        <property type="entry name" value="Vasodilator-stimulated phosphoprotein"/>
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        <property type="entry name" value="GATA-TYPE DOMAIN-CONTAINING PROTEIN"/>
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    </dbReference>
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    </dbReference>
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    </dbReference>
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    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
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        <location>
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            <end position="264"/>
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            <end position="242"/>
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        <location>
            <begin position="311"/>
            <end position="329"/>
        </location>
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            <dbReference type="ARBA" id="ARBA00061646"/>
        </source>
    </evidence>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
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        </source>
    </evidence>
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        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS51842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011390.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="329" mass="36424" checksum="5D3C920F03627C3A" modified="2019-05-08" version="1">MFEAYINNLRRQLESLGSERLRLDGELKNMQDTVEDFRIRYEDEVNKRTAAENEFVVLKKDVDTAYMNKVELEAKVDALTDEMNFLRALYEAELSHFQSQISDTSVVLSMDNSRNLDLDGIIAEVKAQYEDIANRSRAEAESWYQTKYEELQLTAGRHGDDLRNTKIEISEMNRMIQRLHSEIDNVKKQCASLQTAIADAEQRGELALKDAQAKLAELEEALQKTKADMARQLREYQELMNVKLALDIEIATYRKLLEGEENSSYGTGGGLGLGVGLGNAAGLGYGIGGSSFSASSGRGLVGGFSAGGGSSSSMKIVSKTTSTKKSIRS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9V7</accession>
    <name>A0A452H9V7_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="8">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
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            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452H9V7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013046.1">
        <property type="protein sequence ID" value="ENSGAGP00000011395.1"/>
        <property type="gene ID" value="ENSGAGG00000008776.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004386">
        <property type="term" value="F:helicase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016787">
        <property type="term" value="F:hydrolase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd17979">
        <property type="entry name" value="DEXHc_DHX34"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18791">
        <property type="entry name" value="SF2_C_RHA"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="ATP-dependent RNA helicase DHX34"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000540">
        <property type="entry name" value="probable ATP-dependent RNA helicase DHX34"/>
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    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000725">
        <property type="entry name" value="probable ATP-dependent RNA helicase DHX34"/>
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    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
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        <property type="entry name" value="DEAD/DEAH_box_helicase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048333">
        <property type="entry name" value="HA2_WH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007502">
        <property type="entry name" value="Helicase-assoc_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014001">
        <property type="entry name" value="Helicase_ATP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001650">
        <property type="entry name" value="Helicase_C-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18934">
        <property type="entry name" value="ATP-DEPENDENT RNA HELICASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18934:SF221">
        <property type="entry name" value="ATP-DEPENDENT RNA HELICASE DHX34-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00270">
        <property type="entry name" value="DEAD"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00271">
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04408">
        <property type="entry name" value="WHD_HA2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00487">
        <property type="entry name" value="DEXDc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00847">
        <property type="entry name" value="HA2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00490">
        <property type="entry name" value="HELICc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51192">
        <property type="entry name" value="HELICASE_ATP_BIND_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51194">
        <property type="entry name" value="HELICASE_CTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="3" id="KW-0347">Helicase</keyword>
    <keyword evidence="2" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="1" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Helicase ATP-binding" evidence="6">
        <location>
            <begin position="188"/>
            <end position="348"/>
        </location>
    </feature>
    <feature type="domain" description="Helicase C-terminal" evidence="7">
        <location>
            <begin position="386"/>
            <end position="554"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="1"/>
            <end position="35"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="720"/>
            <end position="820"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="734"/>
            <end position="751"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="803"/>
            <end position="820"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022806"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS51192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS51194"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011395.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="820" mass="91157" checksum="7D50B9C5B0A68062" modified="2019-05-08" version="1">PTPCLCREGGGRRLGAQRRSRDPSPSRGHQGGRWDWSCPEMRRRLEELYFREQDYIRAGSEDSRQFWAFFERLQRFQTLKAEGELAPRAPPARADPSAALGLAGLPQEYDPRYRINLSVLSGDVEGLAQGRSGGPRRGPGVPAERLSQFRAALLHYLDFAQKQSFAKLAKLQRERAALPIAQYRDRLLRAVAQNQVVVVAGDTGCGKSTQVPQYLLAAGYSHVACTQPRRIACISLAKRVGFESLHQYGAEVGYQIRFESTRSLATKIVFLTEGLLLRQVQRELALPGYQVLIADEVHERHLHSDFLLGVLRRLLPSRPDLKLVLMSATINIRLFAGYFGGAPVVQVPGRLFPITVVYQPIPQEEAAASGKPERLDPRPYLRVLQAIDHKYPPEERGDLLVFLSGVAEIGAVLEAAQAYATHTQRWVVLPLHSTLSLAEQDKVFDMPPSGVRKCIVSTNIAETSVTIDGVRFVLDSGKVKELSYDPQAKLQRLQEFWISRASAEQRKGRAGRTGPGVCYRLYAESDYDAFAPYPVPEIQRVALDALVLQMKSMGLGDPRTFPFLEPPPQSSLEMAVRYLREQGALDEAESLTPIGSLLAQLPVDVVIGKMLVLGALFGLAGPVLTVAAALSVQSPFLRPVRANPDCATARRPLESPHGDPLTLLNTFNEWVQVKSEWAGGSRKWCRRRGLEEHRLYEAANLRRQFQELLRDHGLLEAVQGPRDSYARQSRHRVRQELHRLQRRHEETEGRQRKVLKLQGGEAGSSSDEGEGARSAGPPGVDIQVRGRGPQCGAEGPPRLTAHWGQSQHQELSTAMSSPRQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9W1</accession>
    <name>A0A452H9W1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Cullin family profile domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="pathway">
        <text evidence="1">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="similarity">
        <text evidence="2 6 7">Belongs to the cullin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9W1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013051.1">
        <property type="protein sequence ID" value="ENSGAGP00000011400.1"/>
        <property type="gene ID" value="ENSGAGG00000008767.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00143"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031461">
        <property type="term" value="C:cullin-RING ubiquitin ligase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031625">
        <property type="term" value="F:ubiquitin protein ligase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016567">
        <property type="term" value="P:protein ubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006511">
        <property type="term" value="P:ubiquitin-dependent protein catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.230.130:FF:000001">
        <property type="entry name" value="Cullin 4A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.10:FF:000050">
        <property type="entry name" value="Cullin 4B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1310.10:FF:000004">
        <property type="entry name" value="Cullin 4B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1310.10:FF:000008">
        <property type="entry name" value="Cullin 4B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1310.10:FF:000059">
        <property type="entry name" value="Cullin-4B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1310.10">
        <property type="entry name" value="Cullin Repeats"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.230.130">
        <property type="entry name" value="Cullin, Chain C, Domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.10">
        <property type="entry name" value="Winged helix-like DNA-binding domain superfamily/Winged helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045093">
        <property type="entry name" value="Cullin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR059120">
        <property type="entry name" value="Cullin-like_AB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016157">
        <property type="entry name" value="Cullin_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016158">
        <property type="entry name" value="Cullin_homology"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036317">
        <property type="entry name" value="Cullin_homology_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001373">
        <property type="entry name" value="Cullin_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019559">
        <property type="entry name" value="Cullin_neddylation_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016159">
        <property type="entry name" value="Cullin_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036388">
        <property type="entry name" value="WH-like_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036390">
        <property type="entry name" value="WH_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11932">
        <property type="entry name" value="CULLIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00888">
        <property type="entry name" value="Cullin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26557">
        <property type="entry name" value="Cullin_AB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10557">
        <property type="entry name" value="Cullin_Nedd8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00182">
        <property type="entry name" value="CULLIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00884">
        <property type="entry name" value="Cullin_Nedd8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF75632">
        <property type="entry name" value="Cullin homology domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF74788">
        <property type="entry name" value="Cullin repeat-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46785">
        <property type="entry name" value="Winged helix' DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01256">
        <property type="entry name" value="CULLIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50069">
        <property type="entry name" value="CULLIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0832">Ubl conjugation</keyword>
    <keyword evidence="4" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="Cullin family profile" evidence="9">
        <location>
            <begin position="520"/>
            <end position="719"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1"/>
            <end position="85"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="105"/>
            <end position="126"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="52"/>
            <end position="81"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="108"/>
            <end position="125"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006019"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00330"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU003829"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50069"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011400.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="846" mass="97371" checksum="4FB386ADAE1F562E" modified="2019-05-08" version="1">MFPTGFSSPNPPAAAQEVRPATDGNSSSSSSSCKKRKLNNSSNNSEREEFDSISSCSSSPSKNNSSSSSSIITTSSGGPSSNHHLQKKLRFEDSLDFIGLDVKMAEESSSSSSPAASSQQQHQQQLKNKSLLISSVAVGHHANGLTKAASSTVSSFANSKPGSAKKLVIKNFKDKPKLPENYTDETWQKLKEAVEAIQNSTSIKYNLEELYQAVENLCSYKISANLYKQLRQICEDHIKAQIHHVLFLKKIDKCWQDHCRQMIMIRSIFLFLDRTYVLQNSMLPSIWDMGLELFRTHIISDQKVQNKTIDGILLLIERERNGEAIDRSLLRSLLSMLSDLQIYQDSFEHRFLEETNRLYAAEGQRLMQEREVPEYLHHVNKRLEEEADRIITYLDQSTQKPLIATVEKQLLGEHLTAILQKGLNHLLDENRIQDLSLLYQLFSRVRGGVQVLLQHWIEYIKAFGSTIVINPEKDKTMVQELLDFKDKVDHIIDVCFLKNEKFVNAMKEAFETFINKRPNKPAELIAKYVDSKLRAGNKEATDEELEKMLDKIMIIFRFIYGKGYSFVYPWEGMIFFKDKMQLENSTTFSICFCFSLQYMQNQNVPGNIELTVNILTMGYWPTYVPMEVHLPPEMVKLQEIFKTFYLGKHSGRKLQWQSTLGHCVLKAEFKEGRKELQVSLFQTLVLLMFNEGEEFSLEEIKQATGIEDGELRRTLQSLACGKARVLTKSPKGKDVEDGDKFTCNDDFRHKLFRIKINQIQMKETVEEQASTTERVFQDRQYQIDAAIVRIMKMRKTLSHNLLVSEVYNQLKFPVKPADLKKRIESLIDRDYMERDKENPNQYNYIA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9W3</accession>
    <name>A0A452H9W3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Dynein regulatory complex protein 1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="8">Coiled-coil domain-containing protein 164</fullName>
        </alternativeName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Component of the nexin-dynein regulatory complex (N-DRC) a key regulator of ciliary/flagellar motility which maintains the alignment and integrity of the distal axoneme and regulates microtubule sliding in motile axonemes. Plays a critical role in the assembly of N-DRC and also stabilizes the assembly of multiple inner dynein arms and radial spokes. Coassembles with CCDC65/DRC2 to form a central scaffold needed for assembly of the N-DRC and its attachment to the outer doublet microtubules.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Flagellum axoneme</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the DRC1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9W3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013246.1">
        <property type="protein sequence ID" value="ENSGAGP00000011570.1"/>
        <property type="gene ID" value="ENSGAGG00000008764.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005858">
        <property type="term" value="C:axonemal dynein complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070286">
        <property type="term" value="P:axonemal dynein complex assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060285">
        <property type="term" value="P:cilium-dependent cell motility"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003352">
        <property type="term" value="P:regulation of cilium movement"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039505">
        <property type="entry name" value="DRC1/2_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039750">
        <property type="entry name" value="DRC1/DRC2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029440">
        <property type="entry name" value="DRC1_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21625:SF1">
        <property type="entry name" value="DYNEIN REGULATORY COMPLEX PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21625">
        <property type="entry name" value="NYD-SP28 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14772">
        <property type="entry name" value="NYD-SP28"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14775">
        <property type="entry name" value="NYD-SP28_assoc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0966">Cell projection</keyword>
    <keyword evidence="6" id="KW-0969">Cilium</keyword>
    <keyword evidence="5" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0282">Flagellum</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Dynein regulatory complex protein 1/2 N-terminal" evidence="11">
        <location>
            <begin position="45"/>
            <end position="146"/>
        </location>
    </feature>
    <feature type="domain" description="Dynein regulatory complex protein 1 C-terminal" evidence="12">
        <location>
            <begin position="553"/>
            <end position="612"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004611"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009688"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00013815"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022846"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023069"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00031554"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00046115"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF14772"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF14775"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011570.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="633" mass="74322" checksum="585C206A0B224A4E" modified="2019-05-08" version="1">LGEDPDAKKAEAEELSRSHKQIEESRQRLAKLLNDGTQLVTNIQVAADARETQRRAEEDELKRQRLEKLENESKTNQDKFEEITSKWGSAKEKTIPQDLWDVLNQQQQHCALLIEEKNKLISELQQELKSKDDQYVKDLKKQSDDINLLVDRMEEQIRNLMKTYRRELTQIEKAFELERRELLSSNKKKWEQAMQAHNAQELENLITRMQKVEEYEKQLNQLRVQDAEEYNTIKIQLENDVQILEQQLQQMKAIYQLNQEKLEYNFQVLKKRDEENTIIKSQQKRKLNRLHDVLNALRMKLDKQIKQFQEENQTLTADYKRIVEQYKDLQRTMRHFAVVDAQKFLEVWLMNEEEAKGLMWKALKADRIIHTQQLGLPWVEPDYWFLDNVGPIVHHMAIRSASKLAREVMDAVEEGRKEAAAEAGEEVKPPRHISVKSVKHILELLCDESGFLIESKLLGLLQPLERHERSLMRLDAIFAALGIDNEDDVYTLVDFFLKFNAQQVVSSQTQQPREEMGLVNGALSPGLLSLPFRDKRPQVKNMMEERDDSKDSEYWKALAHVIPEPKLKVWDALEAALEKYYSVLSQRSKLLTDLDGLQQQNTELHMLLHQYLTSKVNEELLIPPTHVIELMPP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9W9</accession>
    <name>A0A452H9W9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Anoctamin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="8">Membrane</location>
            <topology evidence="8">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 8">Belongs to the anoctamin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9W9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013055.1">
        <property type="protein sequence ID" value="ENSGAGP00000011404.1"/>
        <property type="gene ID" value="ENSGAGG00000007328.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005229">
        <property type="term" value="F:intracellularly calcium-gated chloride channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046983">
        <property type="term" value="F:protein dimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032394">
        <property type="entry name" value="Anoct_dimer"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007632">
        <property type="entry name" value="Anoctamin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049452">
        <property type="entry name" value="Anoctamin_TM"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12308">
        <property type="entry name" value="ANOCTAMIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12308:SF28">
        <property type="entry name" value="ANOCTAMIN-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16178">
        <property type="entry name" value="Anoct_dimer"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04547">
        <property type="entry name" value="Anoctamin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6 8" id="KW-0472">Membrane</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 8" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 8" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="289"/>
            <end position="315"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="365"/>
            <end position="384"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="441"/>
            <end position="467"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="487"/>
            <end position="507"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="535"/>
            <end position="556"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="655"/>
            <end position="677"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="705"/>
            <end position="729"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="790"/>
            <end position="811"/>
        </location>
    </feature>
    <feature type="domain" description="Anoctamin dimerisation" evidence="11">
        <location>
            <begin position="56"/>
            <end position="275"/>
        </location>
    </feature>
    <feature type="domain" description="Anoctamin transmembrane" evidence="10">
        <location>
            <begin position="278"/>
            <end position="833"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="832"/>
            <end position="859"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009671"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="RuleBase" id="RU280814"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="Pfam" id="PF04547"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF16178"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011404.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="868" mass="101863" checksum="CFBB982F96157046" modified="2019-05-08" version="1">IFQNGSTKFRPLNSCLGTKKEKESSLGSNGDSQLCFGNCFCSSQGVPDKNKSNGLYFRDGKCRIDYILVYRKSSPQTEKREIFERNIRAEGLQMEKEVRDDIIFVKLHAPWEVLGRYAEVMNVRMPFRRKIYYLHRRHKFMNRQISRFQGWLPRKPMRLDKETLPDLEENDCYTAPFSQQRIHHFIIHNKDTFFNNATRSRIVHHILQRVKYEEGKNKIGLNRLLTNGSYEAAFPLHEGSYRSKNSIRTHGAENHRHLLYECWASWGMWYKYQPLDLVRRYFGEKIGLYFAWLGWYTGMLFPAAFIGLFVFLYGVTTLNHCQVSKEVCQATDIIMCPICDKFCPFMRLSDSCVYAKVTHLFDNGATVFFAVFMAVWATVFLEFWKRRRAVIAYDWDLIDWEEEEEEIRPQFEAKYSKKERMNPISGKPEPYQAFADKCSRLIVSASGIFFMICVVIAAVFGIVIYRVVTVSTFAAFKWALIRNNSQVATTGTAVCINFCIIMLLNLYEKVALLLTNLEQPRTESEWENSFTLKMFLFQFVNLNSSTFYIAFFLGRFTGHPGAYLRLINRWRLEECHPSGCLIDLCMQMGIIMVLKQTWNNFMELGYPLIQNWWTRRKLYQEHGSQRKTSFPQWEKDYNLQPMNAYGLFDEYLEMILQFGFTTIFVAAFPLAPLLALLNNIIEIRLDAYKFVTQWRRPLASRAKDIGIWYGILEGIGILSVITNAFVVAVTSDFIPRLVYAYKYGPCAGQGEAGQKCMVGYVNASLSVFLVYRDYRDPPHSPVPYGYTLQFWHVLAARLAFIIVFEHLVFCIKHLISYLIPDLPKDLRDRMRREKYLIQEMMYEAELERLQKERKERKKNGKSHHNEWP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJR8</accession>
    <name>A0A452HJR8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Beta-defensin-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HJR8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017314.1">
        <property type="protein sequence ID" value="ENSGAGP00000015157.1"/>
        <property type="gene ID" value="ENSGAGG00000011441.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006952">
        <property type="term" value="P:defense response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.360.10">
        <property type="entry name" value="Antimicrobial Peptide, Beta-defensin 2, Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001855">
        <property type="entry name" value="Defensin_beta-like"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00711">
        <property type="entry name" value="Defensin_beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57392">
        <property type="entry name" value="Defensin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="1">
        <location>
            <begin position="1"/>
            <end position="18"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019425879" description="Beta-defensin-like domain-containing protein" evidence="1">
        <location>
            <begin position="19"/>
            <end position="69"/>
        </location>
    </feature>
    <feature type="domain" description="Beta-defensin-like" evidence="2">
        <location>
            <begin position="21"/>
            <end position="54"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF00711"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015157.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="69" mass="7666" checksum="A88C8510B74BD790" modified="2019-05-08" version="1">MKAAAFLTLFLCAGFTQFINNPFACRRAGGICRRSCYPNLRPIGRCGFAQSCCRRSWVSSGCHKEDITV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJS1</accession>
    <name>A0A452HJS1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Pecanex-like protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 6">Membrane</location>
            <topology evidence="1 6">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 6">Belongs to the pecanex family.</text>
    </comment>
    <comment type="caution">
        <text evidence="6">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000017301.1">
        <property type="protein sequence ID" value="ENSGAGP00000015144.1"/>
        <property type="gene ID" value="ENSGAGG00000011428.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039797">
        <property type="entry name" value="Pecanex"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007735">
        <property type="entry name" value="Pecanex_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12372">
        <property type="entry name" value="PECANEX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12372:SF2">
        <property type="entry name" value="PECANEX-LIKE PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05041">
        <property type="entry name" value="Pecanex_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 6" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 6" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 6" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="798"/>
            <end position="816"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="823"/>
            <end position="840"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="860"/>
            <end position="876"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="910"/>
            <end position="935"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="947"/>
            <end position="971"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="983"/>
            <end position="1003"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="1057"/>
            <end position="1079"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="1085"/>
            <end position="1102"/>
        </location>
    </feature>
    <feature type="domain" description="Pecanex C-terminal" evidence="8">
        <location>
            <begin position="1573"/>
            <end position="1799"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1"/>
            <end position="48"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="168"/>
            <end position="189"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="246"/>
            <end position="585"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="642"/>
            <end position="667"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1841"/>
            <end position="1899"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="8"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="7">
        <location>
            <begin position="170"/>
            <end position="180"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="268"/>
            <end position="279"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="292"/>
            <end position="308"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="312"/>
            <end position="332"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="343"/>
            <end position="353"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="364"/>
            <end position="376"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="426"/>
            <end position="442"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="7">
        <location>
            <begin position="452"/>
            <end position="467"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="503"/>
            <end position="533"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="1851"/>
            <end position="1866"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="RuleBase" id="RU367089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF05041"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015144.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2131" mass="236014" checksum="278DE2F050185E58" modified="2019-05-08" version="1">MDRGAKTEQGNTSTRRTDSNGPSDPGGGIEMSEFIREATPPVGCSSRNSYAALDANNQIGSGLARLGTAATIKGDTDTAKTSDDISLSLGRSCSLCKEGSEEQDLAADQKILHLVSNDSFVSIQPWLSSGQDLPRDNSDKVCLPNNHHVDQSLSSACDTEMASLVPLHSHSYRKDHRPRGVPRTSSSAVAFPDASLNDFPLYQQRRGLDPVSELEASKPHCGSKESLAGNSCISGVFQLDDILKSSSPQPLAKSGKSKSLKAEKSVDSLRSLSGRSSGSTESYCSGIDRDTNSTISSYKSEQTSSTHVENVLSEHEESPREESETSKRKDCCIDSEDDSSSATDKRSSSERTAVEMSTNSGVQETKDSNTSDEIHSQRGLSASASEEANKNPHANELTTQVDRPPGKAAETRDDQSEKLAALVDSRTCKETGGKQKEGDVRPKSSSLIHRTASVHKSSRRRTGKKRASSFDSSRHREYISFRGVSGTKPQSAVFCHDEDSSDQSDLSRTSSMQSAHQFSSDSSSSTTSHSCQSPEGKYNALKTKYIKERGGADSNTHKAHLGSEGISKKRSTRRTSSTHSVKSRARVLSLDSGTVACLNNPNSLMAPGNIKQLTTSKSDLEAKEGEVLDELSLLGRASHLESVTRSRNSLPSQAMFPEGEEQESASGGKFSSTLFEMGGCDMSLVNFEPAARRSSNICDTDSHISSSTSVRFYPHDLLSLPQIRLNRLLTIDTDLLEHQDVDLSPDLQDRMQSQEEAAQKVKQYYRFWILPKLWIGINFDRLTLLALFDRNREILENVLAVILAILVAFLGSVLLIEGFFKDIWVFQFCLVIASCQYSLLKSVQPDSSSPRHGHNRIIAYSRPVYFCLCCGLIWLLDYGSRNTSTTRFRLYGVVFTNPLLLLSARDLVIVFTLCFPIVFFIGLLPQVNTFVMYLCEQLDIHVFGGNATTSLLAALYSFICSIVAVALLYGLCYGALKDSWDGQHIPVLFSIFCGLLVAISYHLSRQSSDPSVLFSLVQSKFFPNSEDKNPEDPLSEVKDPLPEKLRNSVNERLQSDLIVCLVIGVLYFAIHVSTVFTVLQPVLSYILYALVGSVGFLTHYVLPQLRKQLPWHCFSHSLLKTKEYYQFEVRNAAHVMWFEKLHVWLLFVEKNVIYPLIVLNELSSSAKNIASPKKLDTEIGALMITIAGLKLLRSSFSNPTCQYVTVIFTVLFFTFDYKSFSETMLLDLFFMSILFSKLWELFYKLRFVYTYIAPWQITWGSAFHAFAQPFAVPHSAMLFVQAAVSALFSTPLNPFLGSAIFITSYVRPVKFWERDYNTKRVDHSNTRLASQLDRNPGSDDNNLNSIFYEHLTRSLQHSLCGDLLLGRWGNYSTGDCFILASDYLNALVHLIEIGNGLVTFQLRGLEFRGTYCQQREVEAITEGVEEDEGFCCCEPGHIPHMLSFNATFGQRWLAWEVLVTKYVLEGYSITDNSAASMLQVFDLRKILTTYYVKGIIYYVTTSPKLEEWLANETMQEGLQPCADCNYVDVDPTFNPNIDEDYDHHLAGVSRESFCATYLNWIEYCCSRREKPLDADKDSSLVTLCYGLCVLGRRALGTASHHMSSNLESFLYGLHALFKGDFRISSVRDEWIFADMELLRKVVVPGIRMSLKLHQDHFTSPDEYDDPTVLYEAIVSHEQNLVIAHEGDPAWRSAVLSNSPSLLALRHVMDDGTNEYKIIMLNKRYLSFRVIKVNKECVRGLWAGQQQELVFLRNRNPERGSIQNAKQALRNMINSSCDQPIGYPIYVSPLTTSYSDSHDQLKEFLGGAISLGNIKNFIVSTWHRLRKGCGAGCNSGGNIEDSDAGGGASCTSNSAAANESQSSMSQAGGNLITGQGPGVGLHPPVTSHPSTLGTSHSSHSVQSSLVRHSPARASVASHSSYCYSSRHSSLRTSTTGFVPCRRSSTSQISLRNLPSSIQSRLSMVNQIEPISQTGIVIMQHGLPSSSSSSQSIPACKQHALVGFLGTEGGNNVTDAQSGGNASPANHMQAKRDEVIYRIQIMDPSQVLEGINMSKRKELQWPDEVIRLKAGRNSWKDWSPQEGMEGHVIHRWVPCSRDPGSRSHIDKTILLVQIEDKYVAVFETGVLELGAEV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJU6</accession>
    <name>A0A452HJU6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Calponin-homology (CH) domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HJU6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017347.1">
        <property type="protein sequence ID" value="ENSGAGP00000015189.1"/>
        <property type="gene ID" value="ENSGAGG00000011467.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003779">
        <property type="term" value="F:actin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.418.10:FF:000003">
        <property type="entry name" value="Spectrin beta chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.418.10:FF:000089">
        <property type="entry name" value="Spectrin beta chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.418.10">
        <property type="entry name" value="Calponin-like domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001589">
        <property type="entry name" value="Actinin_actin-bd_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001715">
        <property type="entry name" value="CH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036872">
        <property type="entry name" value="CH_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11915">
        <property type="entry name" value="SPECTRIN/FILAMIN RELATED CYTOSKELETAL PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00307">
        <property type="entry name" value="CH"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00033">
        <property type="entry name" value="CH"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47576">
        <property type="entry name" value="Calponin-homology domain, CH-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00020">
        <property type="entry name" value="ACTININ_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50021">
        <property type="entry name" value="CH"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="Calponin-homology (CH)" evidence="3">
        <location>
            <begin position="1"/>
            <end position="72"/>
        </location>
    </feature>
    <feature type="domain" description="Calponin-homology (CH)" evidence="3">
        <location>
            <begin position="87"/>
            <end position="192"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS50021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015189.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="233" mass="26646" checksum="66FD7B2113495E23" modified="2019-05-08" version="1">MLIKLLEVLSGESLPKPTKGRMRIHCLENVDKALQFLKEKHVHLENMGSHDIVDGNHRLVLGLIWTIILRFQIEDIIVQTPEGRETRSAKEALLLWCQMKTAGYPHVNVTNFTSSWKDGLAFNALIHKHRPDLIDFKNLEHSNAQHNLEHAFSVAERQLGITQLLDPEDVFTENPDEKSIITYVVAFYHYFSKMKVRAVKGKRVAKVSAVPLAPDSAHLGLLHELPFLLFRLH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJU7</accession>
    <name>A0A452HJU7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">SAM domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HJU7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017350.1">
        <property type="protein sequence ID" value="ENSGAGP00000015192.1"/>
        <property type="gene ID" value="ENSGAGG00000011466.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="CDD" id="cd09579">
        <property type="entry name" value="SAM_Samd7_11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.50">
        <property type="entry name" value="Transcription Factor, Ets-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001660">
        <property type="entry name" value="SAM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013761">
        <property type="entry name" value="SAM/pointed_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10417">
        <property type="entry name" value="GLUCOCORTICOID MODULATORY ELEMENT-BINDING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10417:SF15">
        <property type="entry name" value="STERILE ALPHA MOTIF DOMAIN-CONTAINING 11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00536">
        <property type="entry name" value="SAM_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00454">
        <property type="entry name" value="SAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47769">
        <property type="entry name" value="SAM/Pointed domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50105">
        <property type="entry name" value="SAM_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0539">Nucleus</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0804">Transcription</keyword>
    <keyword evidence="1" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="SAM" evidence="5">
        <location>
            <begin position="486"/>
            <end position="532"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="139"/>
            <end position="173"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="303"/>
            <end position="432"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="564"/>
            <end position="625"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="157"/>
            <end position="169"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="330"/>
            <end position="352"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="4">
        <location>
            <begin position="355"/>
            <end position="369"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="387"/>
            <end position="405"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="409"/>
            <end position="424"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="580"/>
            <end position="600"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50105"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015192.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="625" mass="68912" checksum="8E3F87E12DAEA7D2" modified="2019-05-08" version="1">MKRRIHTHWDLNISFRETSCSRDNDLSTIISNLHQSRQLVMPETQSRCEFKRNSIDVGLGAAENSHTPVLEDSVTQMTPEEHYRRMMSALNEHSTFEEQQQQRLYQLANSMAVPSHGDLLRARQEAAAAARNPAAMEAHLPAASNSSSQRRKQGLPQHRDTHFPERDMSHPPPLLSPQNAPHIALGPHLRPPFLGVPSALCQTPGYGFLQPAQAEMFARQQEMLRKQNLARLEMSAEIIRQKELENLHRQRLLAGDPLSLHPALPQDHPALRNMHDIPEGHPLRDELSRRNAMLVLRHNNTPLLSLNHGVPSATPPKEQGRRGSRKSMHHRAEPPGLHEAKELAEPRARDGVPDCNDEEMKDSESDAEVSDEKPESLKAESSSSVSELKECKETSKVYEGAKELSEVASGQSAPCSSSSTESPSHLLGPGISKNEVKYLPPASIPPPQPLPFGFPYTMSPYFHAGAMGGLFLDGEESPALEDISKWTVEDVCNFVGSLSGCAEYTQVFREQAIDGETLPLLTEEHLLTNMGLKLGPALKIRSQVAKRVGRVLYMASFPMAFPLQPPGLRPPDRDLAPAELRPSSTGSVSSPYSSSLLPASRISPKQENGNPSLMAGINDTPKPPS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJU9</accession>
    <name>A0A452HJU9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">protein-tyrosine-phosphatase</fullName>
            <ecNumber evidence="2">3.1.3.48</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="10">
            <text>O-phospho-L-tyrosyl-[protein] + H2O = L-tyrosyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:10684"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="EC" id="3.1.3.48"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="3.1.3.48" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452HJU9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017346.1">
        <property type="protein sequence ID" value="ENSGAGP00000015188.1"/>
        <property type="gene ID" value="ENSGAGG00000004356.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd14636">
        <property type="entry name" value="R-PTPc-K-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000003">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase kappa isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000005">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase kappa isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000019">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase kappa isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000048">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase U isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057598">
        <property type="entry name" value="Fn3_PTPRU"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050348">
        <property type="entry name" value="Protein-Tyr_Phosphatase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000242">
        <property type="entry name" value="PTP_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016130">
        <property type="entry name" value="Tyr_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003595">
        <property type="entry name" value="Tyr_Pase_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19134">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19134:SF209">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE KAPPA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23144">
        <property type="entry name" value="Fn3_PTPRU"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00102">
        <property type="entry name" value="Y_phosphatase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00700">
        <property type="entry name" value="PRTYPHPHTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00194">
        <property type="entry name" value="PTPc"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00404">
        <property type="entry name" value="PTPc_motif"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00383">
        <property type="entry name" value="TYR_PHOSPHATASE_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50055">
        <property type="entry name" value="TYR_PHOSPHATASE_PTP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="9" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="5" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="8 11" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="3 11" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 11" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="404"/>
            <end position="424"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="14">
        <location>
            <begin position="55"/>
            <end position="151"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="14">
        <location>
            <begin position="152"/>
            <end position="256"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="12">
        <location>
            <begin position="494"/>
            <end position="738"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="13">
        <location>
            <begin position="658"/>
            <end position="729"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="12">
        <location>
            <begin position="770"/>
            <end position="1032"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="13">
        <location>
            <begin position="946"/>
            <end position="1023"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00013064"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00051722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015188.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1036" mass="117465" checksum="531726EA9925CD50" modified="2019-05-08" version="1">KSDMAIPISALLLSSGRSTETGLKSPLYLYKGPRCVDRCRCYLHYILCLLEPMRTPKTLKIAEIRARHIAVDWESLGYNITRCHTFNVTICYHYFHGHNENKADCLDMDPKAPQHVVDHLPPYTNVSLKMILTNPEGRKESEETIIQTDEDVPGPVPAKSLKGTPFEDKIFLNWKEPVDPNGIITQYEVIERTSRKLRVILGALSYIQHRPKKNTHHVFSHLHPGTTYQFFIRASTVKGFGPATAINVTTNISAPTLPDYEGVDASLNETATTITVLLRPAQAKGAPISAYQIVVEELHPHRTKRETGTMECYQVPVTYQNALSGSSPYYFAAELPPGNLPEPAPFTVGDNRTYQGFWNPPLAPRKGYNIYFQAMSSVEKVTAATEEPEVIPDPAKQTDRVVKIAGISAGILVFILLLLVVILINKCYLLLWAFVTTDENHSATAESSRLLDVPRYLGEGTESPYQTGQLHPAIRVADLLQHINLMKTSDSYGFKEEYESFFEGQSASWDVAKKDQNRTKNRYGNIIAYDHSRVILQPVEDDPSSDYINANYIDVSGPVHETVYDFWRMIWQEQSACVVMVTNLVEVGRVKCYKYWPDDTEVYGDFKVTCVEMEPLAEYVVRTFTLERRGYNEIRELKQFHFTGWPDHGVPYNATGLLSFIRRVKLSNPPSAGPIVVHCSAGAGRTGCYIVIDIMLDMAEREGVVDIYNCVKALRSRRINMVQTEEQYIFIHDAILEACLCGETAIPVCEFKAAYFDMIRIDSQTNSSHLKDEFQTLNSVTPRLQAEDCSIACLPRNHDKNRFMDMLPPDRCLPFLITIDGESSNYINAALMDSYRQPAAFIVTQHPLPNTVKDFWRLVYDYGCTSVVMLNEVDLAQGCPQYWPEEGMLRYGPIQVECMSCSMDCDVINRIFRICNLTRPQEGYLMVQQFQYLGWASHRDVPGSKRSFLKLILQVEKWQEECEEGEGRTIIHCLNGGGRSGMFCAIGIVAEMVKRQNVVDVFHAVKTLRNSKPNMVETPEQYRFCYDVALEYLESS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJV5</accession>
    <name>A0A452HJV5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">SAM domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HJV5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017355.1">
        <property type="protein sequence ID" value="ENSGAGP00000015196.1"/>
        <property type="gene ID" value="ENSGAGG00000011466.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003682">
        <property type="term" value="F:chromatin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042393">
        <property type="term" value="F:histone binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045892">
        <property type="term" value="P:negative regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd09579">
        <property type="entry name" value="SAM_Samd7_11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.50">
        <property type="entry name" value="Transcription Factor, Ets-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050548">
        <property type="entry name" value="PcG_chromatin_remod_factors"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001660">
        <property type="entry name" value="SAM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013761">
        <property type="entry name" value="SAM/pointed_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12247">
        <property type="entry name" value="POLYCOMB GROUP PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12247:SF67">
        <property type="entry name" value="STERILE ALPHA MOTIF DOMAIN-CONTAINING PROTEIN 11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00536">
        <property type="entry name" value="SAM_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00454">
        <property type="entry name" value="SAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47769">
        <property type="entry name" value="SAM/Pointed domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50105">
        <property type="entry name" value="SAM_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="SAM" evidence="2">
        <location>
            <begin position="474"/>
            <end position="520"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1"/>
            <end position="60"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="127"/>
            <end position="161"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="291"/>
            <end position="420"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="552"/>
            <end position="613"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="39"/>
            <end position="53"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="145"/>
            <end position="157"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="318"/>
            <end position="340"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="1">
        <location>
            <begin position="343"/>
            <end position="357"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="375"/>
            <end position="393"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="1">
        <location>
            <begin position="397"/>
            <end position="412"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="1">
        <location>
            <begin position="568"/>
            <end position="588"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="PROSITE" id="PS50105"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015196.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="613" mass="67306" checksum="B9106095E9A8118E" modified="2019-05-08" version="1">MKGVETQEGQKAAEQMMKLNEILGRRRVCSPNSSSECPLESKKTRSESPKENSHTPVLEDSVTQMTPEEHYRRMMSALNEHSTFEEQQQQRLYQLANSMAVPSHGDLLRARQEAAAAARNPAAMEAHLPAASNSSSQRRKQGLPQHRDTHFPERDMSHPPPLLSPQNAPHIALGPHLRPPFLGVPSALCQTPGYGFLQPAQAEMFARQQEMLRKQNLARLEMSAEIIRQKELENLHRQRLLAGDPLSLHPALPQDHPALRNMHDIPEGHPLRDELSRRNAMLVLRHNNTPLLSLNHGVPSATPPKEQGRRGSRKSMHHRAEPPGLHEAKELAEPRARDGVPDCNDEEMKDSESDAEVSDEKPESLKAESSSSVSELKECKETSKVYEGAKELSEVASGQSAPCSSSSTESPSHLLGPGISKNEVKYLPPASIPPPQPLPFGFPYTMSPYFHAGAMGGLFLDGEESPALEDISKWTVEDVCNFVGSLSGCAEYTQVFREQAIDGETLPLLTEEHLLTNMGLKLGPALKIRSQVAKRVGRVLYMASFPMAFPLQPPGLRPPDRDLAPAELRPSSTGSVSSPYSSSLLPASRISPKQENGNPSLMAGINDTPKPPS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJW7</accession>
    <name>A0A452HJW7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">SLIT-ROBO Rho GTPase-activating protein 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="28 29">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="29" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="28" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="28" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="5">Cell projection</location>
            <location evidence="5">Dendritic spine</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cell projection</location>
            <location evidence="3">Lamellipodium</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cytoplasm</location>
            <location evidence="4">Cytosol</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasmic vesicle</location>
            <location evidence="2">Phagosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="18">Postsynaptic cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="19">Postsynaptic density</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJW7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017373.1">
        <property type="protein sequence ID" value="ENSGAGP00000015212.1"/>
        <property type="gene ID" value="ENSGAGG00000010110.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043197">
        <property type="term" value="C:dendritic spine"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030027">
        <property type="term" value="C:lamellipodium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045335">
        <property type="term" value="C:phagocytic vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0014069">
        <property type="term" value="C:postsynaptic density"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd04383">
        <property type="entry name" value="RhoGAP_srGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd11955">
        <property type="entry name" value="SH3_srGAP1-3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.555.10:FF:000010">
        <property type="entry name" value="SLIT-ROBO Rho GTPase-activating protein 1 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1270.60:FF:000006">
        <property type="entry name" value="SLIT-ROBO Rho GTPase-activating protein 1 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.40:FF:000266">
        <property type="entry name" value="SLIT-ROBO Rho GTPase-activating protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.60">
        <property type="entry name" value="Arfaptin homology (AH) domain/BAR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.555.10">
        <property type="entry name" value="Rho GTPase activation protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027267">
        <property type="entry name" value="AH/BAR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031160">
        <property type="entry name" value="F_BAR_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001060">
        <property type="entry name" value="FCH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000198">
        <property type="entry name" value="RhoGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051627">
        <property type="entry name" value="SLIT-ROBO_RhoGAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035648">
        <property type="entry name" value="srGAP1/2/3_SH3"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14166">
        <property type="entry name" value="SLIT-ROBO RHO GTPASE ACTIVATING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00611">
        <property type="entry name" value="FCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00620">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00018">
        <property type="entry name" value="SH3_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00055">
        <property type="entry name" value="FCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00324">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00326">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103657">
        <property type="entry name" value="BAR/IMD domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51741">
        <property type="entry name" value="F_BAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50238">
        <property type="entry name" value="RHOGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="16" id="KW-0966">Cell projection</keyword>
    <keyword evidence="12 22 23" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="9" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="17" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="7" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="13" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="14" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-0628">Postsynaptic cell membrane</keyword>
    <keyword evidence="29" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6 21" id="KW-0728">SH3 domain</keyword>
    <keyword evidence="11" id="KW-0770">Synapse</keyword>
    <feature type="domain" description="F-BAR" evidence="27">
        <location>
            <begin position="7"/>
            <end position="251"/>
        </location>
    </feature>
    <feature type="domain" description="Rho-GAP" evidence="26">
        <location>
            <begin position="437"/>
            <end position="626"/>
        </location>
    </feature>
    <feature type="domain" description="SH3" evidence="25">
        <location>
            <begin position="675"/>
            <end position="734"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="24">
        <location>
            <begin position="640"/>
            <end position="660"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="24">
        <location>
            <begin position="734"/>
            <end position="767"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="24">
        <location>
            <begin position="783"/>
            <end position="884"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="24">
        <location>
            <begin position="981"/>
            <end position="1019"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="23">
        <location>
            <begin position="324"/>
            <end position="358"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="24">
        <location>
            <begin position="738"/>
            <end position="748"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="24">
        <location>
            <begin position="817"/>
            <end position="829"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="24">
        <location>
            <begin position="842"/>
            <end position="853"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="24">
        <location>
            <begin position="867"/>
            <end position="881"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="24">
        <location>
            <begin position="997"/>
            <end position="1019"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004262"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004510"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004514"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004552"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023257"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00034100"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00034105"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00073507"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01077"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="26">
        <source>
            <dbReference type="PROSITE" id="PS50238"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="27">
        <source>
            <dbReference type="PROSITE" id="PS51741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="28">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015212.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="29">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1019" mass="114611" checksum="3E73D389CCADAF16" modified="2019-05-08" version="1">MSSPTRKKLQLTRFIRTQLIEQLKCLDQQCELRVQLLQDLQDFFRKKAEIEMDYSRNLEKLAERFLAKTRSTKDQQFKKDQNVLSPVNCWNLLLNQVKRESRDHTTLSDIYLNNIIPRFVQVSEDSGRLFKKSKEVGLQLQEDLMKVLNELYTVMKTYHMYNADSISAQSKLKEAEKQEEKQIGKSVKFEEKHVRRSSVKKIEKMKEKRQAKYTENKLKAIKARNEYLLALEATNASVFKYYIHDLSDLIDVSVLLLGLMSFPLTPLCASPSLTPFLNSCDPKDGETPLFPSYTCIFCLSRLGKIYQGASFLQVSQLCAQQPVQSELVQRCQQLQSRLSTLKIENEEVKKTMEATLQTIQDIVTIEDFDVSDCFQYSNSMESVKSTVSETFMSKPSIAKRRANQQETEQFYFTKMKEYLEGRNLITKLQAKHDLLQKTLGESKSGTRFQAIPLVVESCIRFISRHGLQHEGIFRVSGSQVEVNDIKNAFERGEDPLAGDQNDHDMDSIAGVLKLYFRGLEHPLFPKEIFHDLIACVTMDNLQERALHIRKVLLSLPKTTLIVMRYLFAFLSHLSQFSEENMMDPYNLAICFGPTLMSVPEGHDQVSCQAHVNELIKTIIIQHENIFPGLRELEGPVYSRGGSTEDYCESPHGETASAEDSVQDIAAEHHTSDDECEPIEAIAKFDYIGRTARELSFKKGASLLLYQRASDDWWEGRHNGIDGLIPHQYIVVQDTEDGSSERSSPKSEIEINSEPPEEKVTARASANCPSGGHVADIYLANINKQRKRPESGSIRRAFRQSDSHGLASSLVEPASPGATASSRPSSQPIMSPSLPKEVPDKCSSSGHGSLNSISRHSSLKNRIDSPQIRKTVTAGRSKSFNNHRPMDPEVIVQDIEATMNTALNELRELERQSSVKHTPDVVLDTLEPLKTSPVVAPTSEPSSPLHTQILKDSEPAFQRSASTAGDIPCAFRPVKSVKMAAQVKPPATRPKPAVFPKTNATSPGVASSASQQPTDKSCTV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJW8</accession>
    <name>A0A452HJW8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Sacsin</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="8">DnaJ homolog subfamily C member 29</fullName>
        </alternativeName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="6">Co-chaperone which acts as a regulator of the Hsp70 chaperone machinery and may be involved in the processing of other ataxia-linked proteins.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000017370.1">
        <property type="protein sequence ID" value="ENSGAGP00000015209.1"/>
        <property type="gene ID" value="ENSGAGG00000011482.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030544">
        <property type="term" value="F:Hsp70 protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="entry name" value="DnaJ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.330:FF:000009">
        <property type="entry name" value="Sacsin molecular chaperone"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.110">
        <property type="entry name" value="DnaJ domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.330">
        <property type="entry name" value="Nucleotidyltransferases domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001623">
        <property type="entry name" value="DnaJ_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036890">
        <property type="entry name" value="HATPase_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007842">
        <property type="entry name" value="HEPN_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036869">
        <property type="entry name" value="J_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058210">
        <property type="entry name" value="SACS/Nov_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052972">
        <property type="entry name" value="Sacsin_chaperone_reg"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF047352">
        <property type="entry name" value="P_loop_sacsin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15600">
        <property type="entry name" value="SACSIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15600:SF42">
        <property type="entry name" value="SACSIN"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25794">
        <property type="entry name" value="SACS"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00748">
        <property type="entry name" value="HEPN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55874">
        <property type="entry name" value="ATPase domain of HSP90 chaperone/DNA topoisomerase II/histidine kinase"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46565">
        <property type="entry name" value="Chaperone J-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81593">
        <property type="entry name" value="Nucleotidyltransferase substrate binding subunit/domain"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="DNAJ_2"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0007">Acetylation</keyword>
    <keyword evidence="5" id="KW-0143">Chaperone</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="J" evidence="10">
        <location>
            <begin position="4163"/>
            <end position="4247"/>
        </location>
    </feature>
    <feature type="domain" description="HEPN" evidence="11">
        <location>
            <begin position="4304"/>
            <end position="4420"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="4101"/>
            <end position="4152"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="4107"/>
            <end position="4119"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="9">
        <location>
            <begin position="4142"/>
            <end position="4152"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023186"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00054721"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00073920"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00076379"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50910"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015209.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="4432" mass="503964" checksum="2D194B165BF73086" modified="2019-05-08" version="1">[encoded data, 4432 characters]</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJX0</accession>
    <name>A0A452HJX0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Gamma-sarcoglycan</fullName>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
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        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">Component of the sarcoglycan complex, a subcomplex of the dystrophin-glycoprotein complex which forms a link between the F-actin cytoskeleton and the extracellular matrix.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cell membrane</location>
            <location evidence="3">Sarcolemma</location>
            <topology evidence="3">Single-pass type II membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the sarcoglycan beta/delta/gamma/zeta family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJX0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017376.1">
        <property type="protein sequence ID" value="ENSGAGP00000015215.1"/>
        <property type="gene ID" value="ENSGAGG00000011484.1"/>
    </dbReference>
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    </dbReference>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016012">
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        <property type="project" value="InterPro"/>
    </dbReference>
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        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
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        <property type="term" value="P:cardiac muscle tissue development"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="P:heart contraction"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="entry name" value="Sarcoglycan"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039972">
        <property type="entry name" value="Sarcoglycan_gamma/delta/zeta"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
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    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
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    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
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        <location>
            <begin position="36"/>
            <end position="58"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00002860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004274"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00007574"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
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        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015215.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="291" mass="32557" checksum="43B3D51E077DF213" modified="2019-05-08" version="1">MVREQYTSTTQSTSLERPENQYVYNIGIYGWRKRCLYLFVLLLLIVLVVNFALTIWILKVMWFSPTGMGHLHVTKEGLRLEGESEFLFPLYAKEIHSREDSSLLLQSTHNVTVNARNSDGEVTGRLNVGPKMVEVHGQQFQINSVDGKPLFTVDENEVVVGTDKLRITGPEGAIFEHSVETPLVRAEAFKPLRLESPTRSLSMDAPKGINIKAQAGNIEALSQMDIKLHSSEGVLLLDAETVRLPKLPEGTRGGSDRSQELYEICVCPDGKLYLSAAGRYSTCQEYSHVCQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJX1</accession>
    <name>A0A452HJX1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">SAM domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
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        <lineage>
            <taxon>Eukaryota</taxon>
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            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HJX1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017348.1">
        <property type="protein sequence ID" value="ENSGAGP00000015190.1"/>
        <property type="gene ID" value="ENSGAGG00000011466.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="CDD" id="cd09579">
        <property type="entry name" value="SAM_Samd7_11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.50">
        <property type="entry name" value="Transcription Factor, Ets-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001660">
        <property type="entry name" value="SAM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013761">
        <property type="entry name" value="SAM/pointed_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10417">
        <property type="entry name" value="GLUCOCORTICOID MODULATORY ELEMENT-BINDING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10417:SF15">
        <property type="entry name" value="STERILE ALPHA MOTIF DOMAIN-CONTAINING 11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00536">
        <property type="entry name" value="SAM_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00454">
        <property type="entry name" value="SAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47769">
        <property type="entry name" value="SAM/Pointed domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50105">
        <property type="entry name" value="SAM_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0539">Nucleus</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0804">Transcription</keyword>
    <keyword evidence="1" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="SAM" evidence="5">
        <location>
            <begin position="517"/>
            <end position="563"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="72"/>
            <end position="101"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="170"/>
            <end position="204"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="334"/>
            <end position="463"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="595"/>
            <end position="656"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="82"/>
            <end position="96"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="188"/>
            <end position="200"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="361"/>
            <end position="383"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="4">
        <location>
            <begin position="386"/>
            <end position="400"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="418"/>
            <end position="436"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="440"/>
            <end position="455"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="611"/>
            <end position="631"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50105"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015190.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="656" mass="72370" checksum="EF05098F6DC9062B" modified="2019-05-08" version="1">MKRRIHTHWDLNISFRETSCSRDNDLSTIISNLHQSRQLVMPETQSRCEFKRNSIDVGLGAADEILGRRRVCSPNSSSECPLESKKTRSESPKENSHTPVLEDSVTQMTPEEHYRRMMSALNEHSTFEEQQQQRLYQLANSMAVPSHGDLLRARQEAAAAARNPAAMEAHLPAASNSSSQRRKQGLPQHRDTHFPERDMSHPPPLLSPQNAPHIALGPHLRPPFLGVPSALCQTPGYGFLQPAQAEMFARQQEMLRKQNLARLEMSAEIIRQKELENLHRQRLLAGDPLSLHPALPQDHPALRNMHDIPEGHPLRDELSRRNAMLVLRHNNTPLLSLNHGVPSATPPKEQGRRGSRKSMHHRAEPPGLHEAKELAEPRARDGVPDCNDEEMKDSESDAEVSDEKPESLKAESSSSVSELKECKETSKVYEGAKELSEVASGQSAPCSSSSTESPSHLLGPGISKNEVKYLPPASIPPPQPLPFGFPYTMSPYFHAGAMGGLFLDGEESPALEDISKWTVEDVCNFVGSLSGCAEYTQVFREQAIDGETLPLLTEEHLLTNMGLKLGPALKIRSQVAKRVGRVLYMASFPMAFPLQPPGLRPPDRDLAPAELRPSSTGSVSSPYSSSLLPASRISPKQENGNPSLMAGINDTPKPPS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJX5</accession>
    <name>A0A452HJX5_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="7">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="6">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
        <text evidence="6">Binds 1 Mg(2+) ion per subunit.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the 5'(3')-deoxyribonucleotidase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJX5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017375.1">
        <property type="protein sequence ID" value="ENSGAGP00000015214.1"/>
        <property type="gene ID" value="ENSGAGG00000011446.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008253">
        <property type="term" value="F:5'-nucleotidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1000:FF:000026">
        <property type="entry name" value="NT5DC3 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1000">
        <property type="entry name" value="HAD superfamily/HAD-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036412">
        <property type="entry name" value="HAD-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008380">
        <property type="entry name" value="HAD-SF_hydro_IG_5-nucl"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023214">
        <property type="entry name" value="HAD_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016695">
        <property type="entry name" value="Pur_nucleotidase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12103">
        <property type="entry name" value="5'-NUCLEOTIDASE DOMAIN-CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12103:SF11">
        <property type="entry name" value="5'-NUCLEOTIDASE DOMAIN-CONTAINING PROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05761">
        <property type="entry name" value="5_nucleotid"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF017434">
        <property type="entry name" value="Purine_5'-nucleotidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56784">
        <property type="entry name" value="HAD-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="4 6" id="KW-0460">Magnesium</keyword>
    <keyword evidence="2 6" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <feature type="active site" description="Nucleophile" evidence="5">
        <location>
            <position position="27"/>
        </location>
    </feature>
    <feature type="active site" description="Proton donor" evidence="5">
        <location>
            <position position="29"/>
        </location>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="27"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="29"/>
        </location>
        <ligand>
            <name>GMP</name>
            <dbReference type="ChEBI" id="CHEBI:58115"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="271"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00009589"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PIRSR" id="PIRSR017434-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PIRSR" id="PIRSR017434-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015214.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="431" mass="51185" checksum="2A529B2BEA35B10A" modified="2019-05-08" version="1">MSNLLNPDAIFTNNEMSLSDIEIYGFDYDYTLVFYSKHLHTLIYNAARDLLINEHRYPAEIRKYDYDPNFAIRGLHYDVHRALLMKIDAFHYIQLGTVYRGFSVVPDEEVIAMYDGSHVPLEQMSDFYGKKIIQKSILFTLMFNMILKLIFFLEKYVCYAEQTRAVLRKLADHGKKIFLITNSPSSFVDKGMKFLVGKDWRDLFDVVIVQADKPNFFNDKRRPFRKVNERGVLLWDKIHKLQKGQIYKQGNLYEFLKLTGWRGSKVLYFGDHIYSDLADLTLKHGWRTGAIIPELRSEIKIMNTEQYIQTMTWLQTLTGLLEHMQVYPDSQMILQEWKKERKEMREMTRNFFNSQFGSLFRTDQNPTYFLRRLSRFADIYMASLSCLLNYDLNYTFYPRRTPLQHELPTWSDQLCSGAFRIPLLQEMAQIK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJX9</accession>
    <name>A0A452HJX9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">C-type lectin domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type II membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJX9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017390.1">
        <property type="protein sequence ID" value="ENSGAGP00000015225.1"/>
        <property type="gene ID" value="ENSGAGG00000011489.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030246">
        <property type="term" value="F:carbohydrate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd03593">
        <property type="entry name" value="CLECT_NK_receptors_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.100.10">
        <property type="entry name" value="Mannose-Binding Protein A, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001304">
        <property type="entry name" value="C-type_lectin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016186">
        <property type="entry name" value="C-type_lectin-like/link_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050828">
        <property type="entry name" value="C-type_lectin/matrix_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016187">
        <property type="entry name" value="CTDL_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033992">
        <property type="entry name" value="NKR-like_CTLD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45710:SF35">
        <property type="entry name" value="C-TYPE LECTIN DOMAIN FAMILY 2 MEMBER D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45710">
        <property type="entry name" value="C-TYPE LECTIN DOMAIN-CONTAINING PROTEIN 180"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00059">
        <property type="entry name" value="Lectin_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00034">
        <property type="entry name" value="CLECT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56436">
        <property type="entry name" value="C-type lectin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50041">
        <property type="entry name" value="C_TYPE_LECTIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0430">Lectin</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="3">
        <location>
            <begin position="1"/>
            <end position="19"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019571910" description="C-type lectin domain-containing protein" evidence="3">
        <location>
            <begin position="20"/>
            <end position="167"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="4">
        <location>
            <begin position="50"/>
            <end position="153"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004401"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022734"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50041"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015225.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="167" mass="18984" checksum="D4A9D6EBD9DB7CE2" modified="2019-05-08" version="1">RKQRDTFLIAAILLTLVKSPPESNLPRVSTPVRLNVRFPAPCCLDGWVGYRGKCYYFSEAEGTWDSSQSFCSSLNASLAEIDTEKDLKFIMRYKGVSEFWIGLKRESAQLWGWVNGEQFNNLFTVRGEGDCAYLSDGFATSSWCSTKRYWICSKPDGMQREDTMPGD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJY5</accession>
    <name>A0A452HJY5_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="15">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">May be involved in transcriptional regulation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJY5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017386.1">
        <property type="protein sequence ID" value="ENSGAGP00000015224.1"/>
        <property type="gene ID" value="ENSGAGG00000011487.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003700">
        <property type="term" value="F:DNA-binding transcription factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07765">
        <property type="entry name" value="KRAB_A-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000739">
        <property type="entry name" value="Zgc:171418 protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001024">
        <property type="entry name" value="Zinc finger and SCAN domain-containing protein 20"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000295">
        <property type="entry name" value="zinc finger protein 19"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000352">
        <property type="entry name" value="zinc finger protein 3 homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002343">
        <property type="entry name" value="Zinc finger protein 33A"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001270">
        <property type="entry name" value="zinc finger protein 583 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000990">
        <property type="entry name" value="zinc finger protein 629 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000912">
        <property type="entry name" value="Zinc finger protein 660"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002074">
        <property type="entry name" value="Zinc finger protein 841"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050589">
        <property type="entry name" value="Ikaros_C2H2-ZF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001909">
        <property type="entry name" value="KRAB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036051">
        <property type="entry name" value="KRAB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24404">
        <property type="entry name" value="ZINC FINGER PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24404:SF100">
        <property type="entry name" value="ZINC FINGER PROTEIN 501"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01352">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="SMART" id="SM00349">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109640">
        <property type="entry name" value="KRAB domain (Kruppel-associated box)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50805">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="10"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="10" id="KW-0804">Transcription</keyword>
    <keyword evidence="8" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="7" id="KW-0862">Zinc</keyword>
    <keyword evidence="6 12" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="KRAB" evidence="14">
        <location>
            <begin position="13"/>
            <end position="96"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="115"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="143"/>
            <end position="170"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="171"/>
            <end position="198"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="199"/>
            <end position="226"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="227"/>
            <end position="254"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="255"/>
            <end position="282"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="283"/>
            <end position="310"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="311"/>
            <end position="338"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="339"/>
            <end position="366"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="367"/>
            <end position="398"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00003767"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015224.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="398" mass="45699" checksum="8B4DBEF9BB1E7B1C" modified="2019-05-08" version="1">VLRVGLLLFQGRLTFEEVAVYFTREEWSLLDPAQRALYGDVMQENYETVVLLGKESSPLGGWTRYLLRSKGKVSRSCARREKAKAYETQQMPEENFSSHSDLITHKRINLEWRRYTCHECGKSFNWACLLVRHRRTHTGEKPYMCSECGKSFGQHSTLATHRRIHTGEMPYVCSECGKRFRDRSAHAKHVRLHTGEMPYTCPECGKSFIDSTCLISHQRTHTGETPYTCSECGQSFTLSGTLSRHRRIHTGEKPYGCSECGKSFSQRSDLITHQRTHTGEKPYTCSECGKSFYNSSSLKKHGRIHTGEKPYRCSECGKSFNRSSALISHERIHTGETPYTCAECGKSFAYHSGLITHKRIHTGEKPYTCSECGKRFHQSSHLHWELTKHQRRHTGETP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK35</accession>
    <name>A0A452HK35_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">protein-tyrosine-phosphatase</fullName>
            <ecNumber evidence="2">3.1.3.48</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="10">
            <text>O-phospho-L-tyrosyl-[protein] + H2O = L-tyrosyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:10684"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="EC" id="3.1.3.48"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="3.1.3.48" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452HK35"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017453.1">
        <property type="protein sequence ID" value="ENSGAGP00000015276.1"/>
        <property type="gene ID" value="ENSGAGG00000004356.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd14636">
        <property type="entry name" value="R-PTPc-K-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000003">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase kappa isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000005">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase kappa isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000048">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase U isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057598">
        <property type="entry name" value="Fn3_PTPRU"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050348">
        <property type="entry name" value="Protein-Tyr_Phosphatase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000242">
        <property type="entry name" value="PTP_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016130">
        <property type="entry name" value="Tyr_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003595">
        <property type="entry name" value="Tyr_Pase_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19134">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19134:SF209">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE KAPPA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23144">
        <property type="entry name" value="Fn3_PTPRU"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00102">
        <property type="entry name" value="Y_phosphatase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00700">
        <property type="entry name" value="PRTYPHPHTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00194">
        <property type="entry name" value="PTPc"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00404">
        <property type="entry name" value="PTPc_motif"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00383">
        <property type="entry name" value="TYR_PHOSPHATASE_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50055">
        <property type="entry name" value="TYR_PHOSPHATASE_PTP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="9" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="5" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="8 11" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="3 11" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 11" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="319"/>
            <end position="339"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="14">
        <location>
            <begin position="67"/>
            <end position="171"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="12">
        <location>
            <begin position="409"/>
            <end position="653"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="13">
        <location>
            <begin position="573"/>
            <end position="644"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="12">
        <location>
            <begin position="685"/>
            <end position="947"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="13">
        <location>
            <begin position="861"/>
            <end position="938"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00013064"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00051722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015276.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="951" mass="107672" checksum="E39C8ED0634347A4" modified="2019-05-08" version="1">QTTWEPYPHWYDTISISLDIDKPWIALTAVIDGVYHHPMFSLLLSPSLSLSLLLEVLLLKNSNLFAVPGPVPAKSLKGTPFEDKIFLNWKEPVDPNGIITQYEVIERTSRKLRVILGALSYIQHRPKKNTHHVFSHLHPGTTYQFFIRASTVKGFGPATAINVTTNISAPTLPDYEGVDASLNETATTITVLLRPAQAKGAPISAYQIVVEELHPHRTKRETGTMECYQVPVTYQNALSGSSPYYFAAELPPGNLPEPAPFTVGDNRTYQGFWNPPLAPRKGYNIYFQAMSSVEKVTAATEEPEVIPDPAKQTDRVVKIAGISAGILVFILLLLVVILINKCYLLLWAFVTTDENHSATAESSRLLDVPRYLGEGTESPYQTGQLHPAIRVADLLQHINLMKTSDSYGFKEEYESFFEGQSASWDVAKKDQNRTKNRYGNIIAYDHSRVILQPVEDDPSSDYINANYIDVSGPVHETVYDFWRMIWQEQSACVVMVTNLVEVGRVKCYKYWPDDTEVYGDFKVTCVEMEPLAEYVVRTFTLERRGYNEIRELKQFHFTGWPDHGVPYNATGLLSFIRRVKLSNPPSAGPIVVHCSAGAGRTGCYIVIDIMLDMAEREGVVDIYNCVKALRSRRINMVQTEEQYIFIHDAILEACLCGETAIPVCEFKAAYFDMIRIDSQTNSSHLKDEFQTLNSVTPRLQAEDCSIACLPRNHDKNRFMDMLPPDRCLPFLITIDGESSNYINAALMDSYRQPAAFIVTQHPLPNTVKDFWRLVYDYGCTSVVMLNEVDLAQGCPQYWPEEGMLRYGPIQVECMSCSMDCDVINRIFRICNLTRPQEGYLMVQQFQYLGWASHRDVPGSKRSFLKLILQVEKWQEECEEGEGRTIIHCLNGGGRSGMFCAIGIVAEMVKRQNVVDVFHAVKTLRNSKPNMVETPEQYRFCYDVALEYLESS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK37</accession>
    <name>A0A452HK37_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">protein-tyrosine-phosphatase</fullName>
            <ecNumber evidence="2">3.1.3.48</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="10">
            <text>O-phospho-L-tyrosyl-[protein] + H2O = L-tyrosyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:10684"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="EC" id="3.1.3.48"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="3.1.3.48" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452HK37"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017434.1">
        <property type="protein sequence ID" value="ENSGAGP00000015261.1"/>
        <property type="gene ID" value="ENSGAGG00000004356.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd14636">
        <property type="entry name" value="R-PTPc-K-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000003">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase kappa isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000005">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase kappa isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000019">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase kappa isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000048">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase U isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057598">
        <property type="entry name" value="Fn3_PTPRU"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050348">
        <property type="entry name" value="Protein-Tyr_Phosphatase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000242">
        <property type="entry name" value="PTP_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016130">
        <property type="entry name" value="Tyr_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003595">
        <property type="entry name" value="Tyr_Pase_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19134">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19134:SF209">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE KAPPA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23144">
        <property type="entry name" value="Fn3_PTPRU"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00102">
        <property type="entry name" value="Y_phosphatase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00700">
        <property type="entry name" value="PRTYPHPHTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00194">
        <property type="entry name" value="PTPc"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00404">
        <property type="entry name" value="PTPc_motif"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00383">
        <property type="entry name" value="TYR_PHOSPHATASE_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50055">
        <property type="entry name" value="TYR_PHOSPHATASE_PTP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="9" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="5" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="8 11" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="3 11" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 11" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="363"/>
            <end position="383"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="14">
        <location>
            <begin position="3"/>
            <end position="99"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="14">
        <location>
            <begin position="100"/>
            <end position="204"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="12">
        <location>
            <begin position="453"/>
            <end position="697"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="13">
        <location>
            <begin position="617"/>
            <end position="688"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="12">
        <location>
            <begin position="729"/>
            <end position="991"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="13">
        <location>
            <begin position="905"/>
            <end position="982"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00013064"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00051722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015261.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="995" mass="112830" checksum="B1B4F1CE99EECE11" modified="2019-05-08" version="1">MRTPKTLKIAEIRARHIAVDWESLGYNITRCHTFNVTICYHYFHGHNENKADCLDMDPKAPQHVVDHLPPYTNVSLKMILTNPEGRKESEETIIQTDEDVPGPVPAKSLKGTPFEDKIFLNWKEPVDPNGIITQYEVIERTSRKLRVILGALSYIQHRPKKNTHHVFSHLHPGTTYQFFIRASTVKGFGPATAINVTTNISAPTLPDYEGVDASLNETATTITVLLRPAQAKGAPISAYQIVVEELHPHRTKRETGTMECYQVPVTYQNALSGSSPYYFAAELPPGNLPEPAPFTVGDNRTYQGFWNPPLAPRKGYNIYFQAMSSVEKVKLLAHDNFPVSAAATEEPEVIPDPAKQTDRVVKIAGISAGILVFILLLLVVILINKCYLLLWAFVTTDENHSATAESSRLLDVPRYLGEGTESPYQTGQLHPAIRVADLLQHINLMKTSDSYGFKEEYESFFEGQSASWDVAKKDQNRTKNRYGNIIAYDHSRVILQPVEDDPSSDYINANYIDVSGPVHETVYDFWRMIWQEQSACVVMVTNLVEVGRVKCYKYWPDDTEVYGDFKVTCVEMEPLAEYVVRTFTLERRGYNEIRELKQFHFTGWPDHGVPYNATGLLSFIRRVKLSNPPSAGPIVVHCSAGAGRTGCYIVIDIMLDMAEREGVVDIYNCVKALRSRRINMVQTEEQYIFIHDAILEACLCGETAIPVCEFKAAYFDMIRIDSQTNSSHLKDEFQTLNSVTPRLQAEDCSIACLPRNHDKNRFMDMLPPDRCLPFLITIDGESSNYINAALMDSYRQPAAFIVTQHPLPNTVKDFWRLVYDYGCTSVVMLNEVDLAQGCPQYWPEEGMLRYGPIQVECMSCSMDCDVINRIFRICNLTRPQEGYLMVQQFQYLGWASHRDVPGSKRSFLKLILQVEKWQEECEEGEGRTIIHCLNGGGRSGMFCAIGIVAEMVKRQNVVDVFHAVKTLRNSKPNMVETPEQYRFCYDVALEYLESS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK41</accession>
    <name>A0A452HK41_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">protein-tyrosine-phosphatase</fullName>
            <ecNumber evidence="2">3.1.3.48</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="10">
            <text>O-phospho-L-tyrosyl-[protein] + H2O = L-tyrosyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:10684"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="EC" id="3.1.3.48"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="3.1.3.48" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452HK41"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017466.1">
        <property type="protein sequence ID" value="ENSGAGP00000015289.1"/>
        <property type="gene ID" value="ENSGAGG00000004356.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd14636">
        <property type="entry name" value="R-PTPc-K-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000003">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase kappa isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000005">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase kappa isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000048">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase U isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057598">
        <property type="entry name" value="Fn3_PTPRU"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050348">
        <property type="entry name" value="Protein-Tyr_Phosphatase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000242">
        <property type="entry name" value="PTP_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016130">
        <property type="entry name" value="Tyr_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003595">
        <property type="entry name" value="Tyr_Pase_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19134">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19134:SF209">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE KAPPA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23144">
        <property type="entry name" value="Fn3_PTPRU"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00102">
        <property type="entry name" value="Y_phosphatase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00700">
        <property type="entry name" value="PRTYPHPHTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00194">
        <property type="entry name" value="PTPc"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00404">
        <property type="entry name" value="PTPc_motif"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00383">
        <property type="entry name" value="TYR_PHOSPHATASE_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50055">
        <property type="entry name" value="TYR_PHOSPHATASE_PTP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="9" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="5" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="8 11" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="3 11" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 11" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="315"/>
            <end position="336"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="14">
        <location>
            <begin position="50"/>
            <end position="156"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="12">
        <location>
            <begin position="449"/>
            <end position="703"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="13">
        <location>
            <begin position="623"/>
            <end position="694"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="12">
        <location>
            <begin position="735"/>
            <end position="997"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="13">
        <location>
            <begin position="911"/>
            <end position="988"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00013064"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00051722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015289.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1001" mass="112896" checksum="8C673EEF76E0B855" modified="2019-05-08" version="1">AHTPSRRIALTAVIDGVYHHPMFSLLLSPSLSLSLLLEVLLLKNSNLFAVPGPVPAKSLKGTPFEDKIFLNWKEPVDPNGIITQYEVSYSSIRSFDPAVLVAGPPQTVSKVWNSTHHVFSHLHPGTTYQFFIRASTVKGFGPATAINVTTNISAPTLPDYEGVDASLNETATTITVLLRPAQAKGAPISAYQIVVEELHPHRTKRETGTMECYQVPVTYQNALSGSSPYYFAAELPPGNLPEPAPFTVGDNRTYQGFWNPPLAPRKGYNIYFQAMSSVEKETKTQCVRIATKAAATEEPEVIPDPAKQTDRVVKIAGISAGILVFILLLLVVILIVKKSKLAKKRKDAMGNTRQEMTHMVNAMDRSYADQSTLHAEDPLSITFMDSHNFSPRYENHSATAESSRLLDVPRYLGEGTESPYQTGQLHPAIRVADLLQHINLMKTSDSYGFKEEYESFFEGQSASWDVAKKDQNRTKNRYGNIIAYDHSRVILQPVEDDPSSDYINANYIDGYQRPSHYIATQGPVHETVYDFWRMIWQEQSACVVMVTNLVEVGRVKCYKYWPDDTEVYGDFKVTCVEMEPLAEYVVRTFTLERRGYNEIRELKQFHFTGWPDHGVPYNATGLLSFIRRVKLSNPPSAGPIVVHCSAGAGRTGCYIVIDIMLDMAEREGVVDIYNCVKALRSRRINMVQTEEQYIFIHDAILEACLCGETAIPVCEFKAAYFDMIRIDSQTNSSHLKDEFQTLNSVTPRLQAEDCSIACLPRNHDKNRFMDMLPPDRCLPFLITIDGESSNYINAALMDSYRQPAAFIVTQHPLPNTVKDFWRLVYDYGCTSVVMLNEVDLAQGCPQYWPEEGMLRYGPIQVECMSCSMDCDVINRIFRICNLTRPQEGYLMVQQFQYLGWASHRDVPGSKRSFLKLILQVEKWQEECEEGEGRTIIHCLNGGGRSGMFCAIGIVAEMVKRQNVVDVFHAVKTLRNSKPNMVETPEQYRFCYDVALEYLESS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK68</accession>
    <name>A0A452HK68_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">protein-serine/threonine phosphatase</fullName>
            <ecNumber evidence="3">3.1.3.16</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="8">
            <text>O-phospho-L-threonyl-[protein] + H2O = L-threonyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:47004"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="EC" id="3.1.3.16"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the protein-tyrosine phosphatase family.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.16" evidence="3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017490.1">
        <property type="protein sequence ID" value="ENSGAGP00000015312.1"/>
        <property type="gene ID" value="ENSGAGG00000011542.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003779">
        <property type="term" value="F:actin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004722">
        <property type="term" value="F:protein serine/threonine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030837">
        <property type="term" value="P:negative regulation of actin filament polymerization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd14569">
        <property type="entry name" value="DSP_slingshot_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000004">
        <property type="entry name" value="Protein phosphatase Slingshot homolog 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014876">
        <property type="entry name" value="DEK_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000340">
        <property type="entry name" value="Dual-sp_phosphatase_cat-dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043587">
        <property type="entry name" value="Phosphatase_SSH-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043588">
        <property type="entry name" value="SSH-N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016130">
        <property type="entry name" value="Tyr_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020422">
        <property type="entry name" value="TYR_PHOSPHATASE_DUAL_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45864:SF3">
        <property type="entry name" value="PROTEIN PHOSPHATASE SLINGSHOT HOMOLOG 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45864">
        <property type="entry name" value="SLINGSHOT PROTEIN PHOSPHATASE HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08766">
        <property type="entry name" value="DEK_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00782">
        <property type="entry name" value="DSPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23040">
        <property type="entry name" value="PH_SSH1-like_1st"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00195">
        <property type="entry name" value="DSPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51998">
        <property type="entry name" value="DEK_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00383">
        <property type="entry name" value="TYR_PHOSPHATASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50054">
        <property type="entry name" value="TYR_PHOSPHATASE_DUAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="5" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="6" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="DEK-C" evidence="12">
        <location>
            <begin position="169"/>
            <end position="224"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="10">
        <location>
            <begin position="228"/>
            <end position="369"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="11">
        <location>
            <begin position="293"/>
            <end position="347"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="549"/>
            <end position="568"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="587"/>
            <end position="608"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="621"/>
            <end position="664"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="718"/>
            <end position="789"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="849"/>
            <end position="899"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="1103"/>
            <end position="1130"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="1174"/>
            <end position="1218"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="1291"/>
            <end position="1329"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="554"/>
            <end position="567"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="587"/>
            <end position="603"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="630"/>
            <end position="641"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="738"/>
            <end position="748"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="9">
        <location>
            <begin position="762"/>
            <end position="772"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="849"/>
            <end position="858"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="1110"/>
            <end position="1128"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="1199"/>
            <end position="1209"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="9">
        <location>
            <begin position="1300"/>
            <end position="1313"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009580"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00013081"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00048336"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS51998"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015312.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1329" mass="147388" checksum="58258BBFBBECAD47" modified="2019-05-08" version="1">MPTCPEDCDSYPQAVRLESTYQNRTRYMVVVSTNGRQDTEENIVLGMDFSSNDSSTCTMGLVLPLWSDTLIHLDGDGGFSVSTDNRVHVFKPVSVQAMWSALQSLHKACEVARSHNYYPGSLFLTWVSYYESHIDSDQSSVNEWNAMQDVQSHRPDSPALFTDVPTERERTERLIKTKLREIMMQKDLENITSKEIRTELEMQMVCNLREFKEFIDNEMIVILGQMDSPTQIFDHVFLGSEWNASNLEDLQNKGVRYILNVTREIDNFFPGIFEYHNIRVYDEEATDLLAYWNDTYKFISKAKKNGSKCLVHCKMGVSRSASTVIAYAMKEYGWDLDRAYEHVKERRAVTKPNPSFMRQLEEYQGILLASKQRHNKLWRSHSDSDLSEHHQPICKPGMELNKKEITTSAEQLSEPTANDNHQPALPVFSVGPHVGRLLQPDTDVIEELGAKERRIHLEFAARDFRAEQIEDELNLNNINRCPAGCCLEESEFPLDNCHSSEALVQLKEPLEMATAFPDLTVDDLETDALKTDLNVHLVPMEEFTACLKDLPRSPDQNSPSPQLSSHLETTDRIDFLSALEKFEELSQESRSHACSHTRMDEQGGGRNGVSKMSVLEVLPAGTAVDTQRNSSASSSPAAPEESSTDEEQPKDISELTTLGPLTRSHSENAISVKEIITEIESINQGIGPCQQKMEVSANHAQAPKRSTVHELPVEVVWTSENKPGKAEQSERACTAEQETAKDLPKPDQEGVPTFQLPTCASDVEESDPAEDQQEPRVQGLNPGSKWCPGSVKRATLEFEERLRQEQEHQHVPLICLLPTRKNSKNDSAAAELAPKGRTEELTWELDPAHGENESDQARAKSPAAEGALPHGPELPGGTHLSPPLAPCFVQPAQEPPLAEPGAEQEVLRTVVSLNGSGEPALPCHLPKRIEIIEYPHMGTPPDLRERDSSREQGTLENAAALLPVDENLNPSVCAETAPVHPDALPLRNPSVLEHKAIGQATFTLGSPEEEYGAAPINLDTSSFTGQVRRLPSAGLDDLWHQRVVHLEGVTEQSTSADDEPVAPSSLSRNGHVLCEASDSAFVELSNEDLDFLSDSTEAVDVSFLPGGLPHSSNSPSIKDSSEAPSSLKQRAKELEARVRQAGLTTPSQMKRSASLAKLGCLELSRDDLTKRELASSHASLTGPDLLPASPRTGQGKCDALPEHGLEGPPEKTCASSPLIRQDSKPTKHFVEQLKTTECIVQSQPVERPPVQYAKEFGSAPQNLYPSTDPRLTSSEEGLPLLQMQVLDSRLPAQSLAGAPRHQRSRTHPLRRLKMATDKRRTSNPFYNTM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK69</accession>
    <name>A0A452HK69_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="9">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the prickle / espinas / testin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HK69"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017492.1">
        <property type="protein sequence ID" value="ENSGAGP00000015313.1"/>
        <property type="gene ID" value="ENSGAGG00000011537.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd09415">
        <property type="entry name" value="LIM1_Prickle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd09418">
        <property type="entry name" value="LIM2_Prickle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd09827">
        <property type="entry name" value="PET_Prickle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.110.10:FF:000035">
        <property type="entry name" value="prickle-like protein 2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.110.10:FF:000005">
        <property type="entry name" value="Testin isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.110.10">
        <property type="entry name" value="Cysteine Rich Protein"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033725">
        <property type="entry name" value="LIM1_prickle"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033726">
        <property type="entry name" value="LIM2_prickle"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010442">
        <property type="entry name" value="PET_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033723">
        <property type="entry name" value="PET_prickle"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047120">
        <property type="entry name" value="Pk/Esn/Tes"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001781">
        <property type="entry name" value="Znf_LIM"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24211">
        <property type="entry name" value="LIM DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24211:SF19">
        <property type="entry name" value="PRICKLE PLANAR CELL POLARITY PROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00412">
        <property type="entry name" value="LIM"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06297">
        <property type="entry name" value="PET"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00132">
        <property type="entry name" value="LIM"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57716">
        <property type="entry name" value="Glucocorticoid receptor-like (DNA-binding domain)"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00478">
        <property type="entry name" value="LIM_DOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50023">
        <property type="entry name" value="LIM_DOMAIN_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51303">
        <property type="entry name" value="PET"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 6" id="KW-0440">LIM domain</keyword>
    <keyword evidence="2 6" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="4 6" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="PET" evidence="8">
        <location>
            <begin position="14"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="domain" description="LIM zinc-binding" evidence="7">
        <location>
            <begin position="124"/>
            <end position="188"/>
        </location>
    </feature>
    <feature type="domain" description="LIM zinc-binding" evidence="7">
        <location>
            <begin position="189"/>
            <end position="249"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00008268"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023038"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015313.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="354" mass="40157" checksum="90A96B314303472F" modified="2019-05-08" version="1">VPVDLERMMCRLISDFQRHSISDDDSGCASEDYAWVPPGLKPEQVYQFFSCLPEDKVPYVNSPGERYRIRQLLHQLPPHDSEAQYCSPLEEEEKKELKLFSQQRKRENLGRGTVRLFPVTITGAICEQCGKQIRGGDIAVFASRAGHGACWHPQCFQCTTCQELLVDLIYFYQEGKIYCGRHHAEHLKPRCQACDEIIFADECTEAEGRHWHMRHFCCFECEEALGGQRYVMRQSRPYCCRCYQALYAEYCDACGEPIGINQGQMTYEGQHWHATDACFCCTCCRLPLLGKPFLPKGGQIFCSRGCSLGGDEASASDSCDSALHGLRPQDGRRAPPRHPAVARSISFADRPLFL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK70</accession>
    <name>A0A452HK70_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Nuclear speckle splicing regulatory protein 1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="4">Coiled-coil domain-containing protein 55</fullName>
        </alternativeName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the NSRP1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HK70"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017470.1">
        <property type="protein sequence ID" value="ENSGAGP00000015293.1"/>
        <property type="gene ID" value="ENSGAGG00000011536.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000381">
        <property type="term" value="P:regulation of alternative mRNA splicing, via spliceosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046850">
        <property type="entry name" value="NRP1_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042816">
        <property type="entry name" value="Nsrp1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018612">
        <property type="entry name" value="NSRP1_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31938">
        <property type="entry name" value="NUCLEAR SPECKLE SPLICING REGULATORY PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31938:SF4">
        <property type="entry name" value="NUCLEAR SPECKLE SPLICING REGULATORY PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20427">
        <property type="entry name" value="NRP1_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09745">
        <property type="entry name" value="NSRP1_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Nuclear speckle splicing regulatory protein 1 N-terminal" evidence="6">
        <location>
            <begin position="36"/>
            <end position="155"/>
        </location>
    </feature>
    <feature type="domain" description="Nuclear speckle splicing regulatory protein 1 RS-like" evidence="7">
        <location>
            <begin position="273"/>
            <end position="447"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="93"/>
            <end position="117"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="167"/>
            <end position="477"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="93"/>
            <end position="111"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="179"/>
            <end position="190"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="197"/>
            <end position="217"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="5">
        <location>
            <begin position="218"/>
            <end position="232"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="233"/>
            <end position="259"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="289"/>
            <end position="477"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00010126"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00020556"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00030718"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="Pfam" id="PF09745"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF20427"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015293.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="520" mass="61766" checksum="8770B2BA9913EC14" modified="2019-05-08" version="1">MLKKHSVFADDSDEETSVGESLQREALKKQVMKQTKLEIQKALAEDSTVYEYDSIYDEMQQQKKESNAKVLSGKDDKKPKYIQNILKAAEVRKQEQEKRMEKKIQKEREMEGGEFDDKEAFVTSAYKKKLQERAEEEEREKKEAALEACLDVTKQKDLSGFYRHLLNQTMGEEEVPECSLREARIKEEKSITNSDESIQRNKSPDERPRLRTPAKEEDNPDADSDLESDSSDDDHKNSRASSKKKENRDSSESNEEDSKHHRSRKHSNSSESSSEEGSHHTKPHINRPRRGESGTGRREKDEHHKERDRNQEKDHRREKEDRHRYGDHTSKDNYRKRNEQEDKQRGKDRKEREGYDKEGRKEKEREDKYSEREREKERIRNGKDRYSDRERGEKCREKEEHLKEKRERDDKDDKKYRDKKQSSPLSLEKDRQCDQEKETKAKDRKSNDERVSCSERFPEPKGKDEEEKQKGAEQMEKKSLCVSKFAKRSSEETVISARDRYLARQMARVSTKSYIEKEED</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK71</accession>
    <name>A0A452HK71_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">protein-arginine deiminase</fullName>
            <ecNumber evidence="3">3.5.3.15</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="7">
            <text>L-arginyl-[protein] + H2O = L-citrullyl-[protein] + NH4(+)</text>
            <dbReference type="Rhea" id="RHEA:18089"/>
            <dbReference type="Rhea" id="RHEA-COMP:10532"/>
            <dbReference type="Rhea" id="RHEA-COMP:10588"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:28938"/>
            <dbReference type="ChEBI" id="CHEBI:29965"/>
            <dbReference type="ChEBI" id="CHEBI:83397"/>
            <dbReference type="EC" id="3.5.3.15"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the protein arginine deiminase family.</text>
    </comment>
    <dbReference type="EC" id="3.5.3.15" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452HK71"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017499.1">
        <property type="protein sequence ID" value="ENSGAGP00000015320.1"/>
        <property type="gene ID" value="ENSGAGG00000011443.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004668">
        <property type="term" value="F:protein-arginine deiminase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1700:FF:000001">
        <property type="entry name" value="Protein-arginine deiminase type-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.75.10.10:FF:000003">
        <property type="entry name" value="Protein-arginine deiminase type-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.75.10.10">
        <property type="entry name" value="L-arginine/glycine Amidinotransferase, Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1700">
        <property type="entry name" value="Protein-arginine deiminase, central domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1860">
        <property type="entry name" value="Protein-arginine deiminase, N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008972">
        <property type="entry name" value="Cupredoxin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004303">
        <property type="entry name" value="PAD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013530">
        <property type="entry name" value="PAD_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036556">
        <property type="entry name" value="PAD_central_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013732">
        <property type="entry name" value="PAD_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038685">
        <property type="entry name" value="PAD_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013733">
        <property type="entry name" value="Prot_Arg_deaminase_cen_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10837">
        <property type="entry name" value="PEPTIDYLARGININE DEIMINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10837:SF11">
        <property type="entry name" value="PROTEIN-ARGININE DEIMINASE TYPE-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03068">
        <property type="entry name" value="PAD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08527">
        <property type="entry name" value="PAD_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08526">
        <property type="entry name" value="PAD_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF001247">
        <property type="entry name" value="Protein-arginine_deiminase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49503">
        <property type="entry name" value="Cupredoxins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55909">
        <property type="entry name" value="Pentein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF110083">
        <property type="entry name" value="Peptidylarginine deiminase Pad4, middle domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0106">Calcium</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="5" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Protein-arginine deiminase (PAD) N-terminal" evidence="10">
        <location>
            <begin position="14"/>
            <end position="95"/>
        </location>
    </feature>
    <feature type="domain" description="Protein-arginine deiminase (PAD) central" evidence="11">
        <location>
            <begin position="112"/>
            <end position="259"/>
        </location>
    </feature>
    <feature type="domain" description="Protein-arginine deiminase C-terminal" evidence="9">
        <location>
            <begin position="266"/>
            <end position="639"/>
        </location>
    </feature>
    <feature type="active site" evidence="8">
        <location>
            <position position="331"/>
        </location>
    </feature>
    <feature type="active site" evidence="8">
        <location>
            <position position="451"/>
        </location>
    </feature>
    <feature type="active site" evidence="8">
        <location>
            <position position="453"/>
        </location>
    </feature>
    <feature type="active site" evidence="8">
        <location>
            <position position="624"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008166"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012200"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00048487"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PIRSR" id="PIRSR001247-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF03068"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="Pfam" id="PF08526"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF08527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015320.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="642" mass="72131" checksum="5DF9DF5913CA0DA6" modified="2019-05-08" version="1">QSKQRRQEICQELSIPSGATSFDARGTSGVKIYVVYNPNAVKKAAGASRWTLNSQVEVIVSMDIPSSTVNDSKVRISYYGQKENAPIEGAWLYLTCVSKYLFVPDPFVFPLQTKWTWGPEGQGAVLLVNCDRDSPGAKDMDSAYQYVQSYAGTHREQTSALDLEDMSLMILRTQGPSAIFADYKLVLHISVADVNKVRVFHASGDGSLAQYKHVLGSDKRSYVVDRSSGQEEDTFYVEGLAFPDADFSGLVSFHVTLLENAPDTPIFTDTVVFRVAPWIMTPNTLQPLEVFVCSNPNGDFLEAMRALVRKANCKLTVCPEVENRRDRWIQDEMEFGYVEAPHKMFPVVFDSPRDRGLRDFAFRRILGPDFGYVTRVPPHGHVSSLDSFGNLDVSPPVTVQGKTYPLGRILIGSSLPRSDGRRMTKAVRDFLYAQKVQPPVELYSDWLSVGHVDEFLSFVPASDRKGFRLLLASPDACYKLFKEKQREGHGKAAQFAGLENVHTITIDEMLADKTLRSDSEYVQSCIDWNRHTLKEELGLTEQDIIDIPQLFMLKGSRADAFFPDVVNMIVLGKHLGIPKPFGPIVDGRCCLEEKVRSLLEPLGLVCIFINDFFSYHTLLGEIHCGTNVLRKPFSFKWWNMIL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK73</accession>
    <name>A0A452HK73_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">Calcium-transporting ATPase</fullName>
            <ecNumber evidence="21">7.2.2.10</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="21">Catalyzes the hydrolysis of ATP coupled with the transport of calcium.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>Ca(2+)(in) + ATP + H2O = Ca(2+)(out) + ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:18105"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="7.2.2.10"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="19">
            <dbReference type="Rhea" id="RHEA:18106"/>
        </physiologicalReaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subunit">
        <text evidence="20">Interacts with sarcolipin (SLN). Interacts with phospholamban (PLN). Interacts with myoregulin (MRLN). Interacts with DWORF.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Endoplasmic reticulum membrane</location>
            <topology evidence="3">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="21">Membrane</location>
            <topology evidence="21">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Sarcoplasmic reticulum membrane</location>
            <topology evidence="2">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4 21">Belongs to the cation transport ATPase (P-type) (TC 3.A.3) family. Type IIA subfamily.</text>
    </comment>
    <comment type="caution">
        <text evidence="21">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="EC" id="7.2.2.10" evidence="21"/>
    <dbReference type="AlphaFoldDB" id="A0A452HK73"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017496.1">
        <property type="protein sequence ID" value="ENSGAGP00000015317.1"/>
        <property type="gene ID" value="ENSGAGG00000011401.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033017">
        <property type="term" value="C:sarcoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005388">
        <property type="term" value="F:P-type calcium transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="CDD" id="cd02083">
        <property type="entry name" value="P-type_ATPase_SERCA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.1110.10:FF:000003">
        <property type="entry name" value="Calcium-transporting ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1000:FF:000005">
        <property type="entry name" value="Calcium-transporting ATPase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1110.10:FF:000065">
        <property type="entry name" value="Sarcoplasmic/endoplasmic reticulum calcium ATPase 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.70.150.10:FF:000160">
        <property type="entry name" value="Sarcoplasmic/endoplasmic reticulum calcium ATPase 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.1110.10">
        <property type="entry name" value="Calcium-transporting ATPase, cytoplasmic domain N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.70.150.10">
        <property type="entry name" value="Calcium-transporting ATPase, cytoplasmic transduction domain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1110.10">
        <property type="entry name" value="Calcium-transporting ATPase, transmembrane domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1000">
        <property type="entry name" value="HAD superfamily/HAD-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004014">
        <property type="entry name" value="ATPase_P-typ_cation-transptr_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023299">
        <property type="entry name" value="ATPase_P-typ_cyto_dom_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018303">
        <property type="entry name" value="ATPase_P-typ_P_site"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023298">
        <property type="entry name" value="ATPase_P-typ_TM_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008250">
        <property type="entry name" value="ATPase_P-typ_transduc_dom_A_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR059000">
        <property type="entry name" value="ATPase_P-type_domA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036412">
        <property type="entry name" value="HAD-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023214">
        <property type="entry name" value="HAD_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005782">
        <property type="entry name" value="P-type_ATPase_IIA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001757">
        <property type="entry name" value="P_typ_ATPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044492">
        <property type="entry name" value="P_typ_ATPase_HD_dom"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01116">
        <property type="entry name" value="ATPase-IIA1_Ca"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01494">
        <property type="entry name" value="ATPase_P-type"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR42861">
        <property type="entry name" value="CALCIUM-TRANSPORTING ATPASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13246">
        <property type="entry name" value="Cation_ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00690">
        <property type="entry name" value="Cation_ATPase_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00122">
        <property type="entry name" value="E1-E2_ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08282">
        <property type="entry name" value="Hydrolase_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00119">
        <property type="entry name" value="CATATPASE"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00120">
        <property type="entry name" value="HATPASE"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDG00002">
        <property type="entry name" value="C1.7:_P-type_atpase_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDS00003">
        <property type="entry name" value="Haloacid_Dehalogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDF00027">
        <property type="entry name" value="p-type_atpase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00831">
        <property type="entry name" value="Cation_ATPase_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81653">
        <property type="entry name" value="Calcium ATPase, transduction domain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81665">
        <property type="entry name" value="Calcium ATPase, transmembrane domain M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56784">
        <property type="entry name" value="HAD-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81660">
        <property type="entry name" value="Metal cation-transporting ATPase, ATP-binding domain N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00154">
        <property type="entry name" value="ATPASE_E1_E2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12 21" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="11 21" id="KW-0106">Calcium</keyword>
    <keyword evidence="7 21" id="KW-0109">Calcium transport</keyword>
    <keyword evidence="10" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="17 21" id="KW-0406">Ion transport</keyword>
    <keyword evidence="13" id="KW-0460">Magnesium</keyword>
    <keyword evidence="18 21" id="KW-0472">Membrane</keyword>
    <keyword evidence="9 21" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="14" id="KW-0703">Sarcoplasmic reticulum</keyword>
    <keyword evidence="15" id="KW-1278">Translocase</keyword>
    <keyword evidence="8 21" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="16 21" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="5 21" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="60"/>
            <end position="78"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="84"/>
            <end position="107"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="260"/>
            <end position="280"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="292"/>
            <end position="321"/>
        </location>
    </feature>
    <feature type="domain" description="Cation-transporting P-type ATPase N-terminal" evidence="22">
        <location>
            <begin position="3"/>
            <end position="77"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004326"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004477"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00005675"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022568"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022951"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022967"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00047282"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00061760"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="RuleBase" id="RU361146"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="SMART" id="SM00831"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015317.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="787" mass="86061" checksum="4D25320DEBFF1897" modified="2019-05-08" version="1">MENSHAKTVEECLAYFGVSENSGLTPEQVKKNLDKYGPNELPAEEGKSIWELVLEQFEDLLVRILLLAACISFVLAWFEEGEETVTAFVEPFVILLILIANAIVGVWQERNAENAIEALKEYEPEMGKVYRADRKSVQRIKARDMVPGDIAEVAVGDKVPADIRIISIKSTTLRVDQSILTGESVSVIKHTDPVPDPRAVNQDKKNMLFSGTNVAAGKAVGIVIATGVGTEIGKIRDEMAATEQEKTPLQQKLDEFGEQLSKVISLICVAVWLINIGHFNDPVHGGSWIRGAIYYFKIAVALAVAAIPEGLPAVITTCLALGTRRMAKKNAIVRSLPSVETLGCTSVICSDKTGTLTTNQMSVCKMFVIEKVEGDVCSLNEFSITGSTYAPEGEVLKNDKHVKAGQYDGLVELATICALCNDSSLDYNEAKGAYEKVGEATETALTCLVEKMNVFNTDVRTLSKVERANACNSVIKQLMKKEFTLEFSRDRKSMSVYCSPAKASRAAVSNKMFIKGAPEGVIDRCNYVRVGTTRVPLTSLVKDKILTVIKEWGTGRDTLRCLALATRDTPPRREDMVLEDSTKFVEYEVDLTFVGCVGMLDPPRKEVMGSIQLCRDAGIRVIMITGDNKGTAIAICRRIGIFTEDEEVMGRAYTGREFDDLPPAEQREACKRACCFARVEPAHKSKIVEFLQSFDEITAMTGDGVNDAPALKKAEIGIAMGSGTAVAKTASEMVLADDNFSTIVSAVEEGRAIYNNMKQFIRYLISSNVGEVPRQPSPPLPQPCHPS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK75</accession>
    <name>A0A452HK75_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Coronin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="4">Belongs to the WD repeat coronin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HK75"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017504.1">
        <property type="protein sequence ID" value="ENSGAGP00000015325.1"/>
        <property type="gene ID" value="ENSGAGG00000011551.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007015">
        <property type="term" value="P:actin filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:000871">
        <property type="entry name" value="Coronin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:003604">
        <property type="entry name" value="Coronin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015505">
        <property type="entry name" value="Coronin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015048">
        <property type="entry name" value="DUF1899"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019775">
        <property type="entry name" value="WD40_repeat_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036322">
        <property type="entry name" value="WD40_repeat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001680">
        <property type="entry name" value="WD40_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10856">
        <property type="entry name" value="CORONIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10856:SF23">
        <property type="entry name" value="CORONIN-6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08953">
        <property type="entry name" value="DUF1899"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00400">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16300">
        <property type="entry name" value="WD40_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01166">
        <property type="entry name" value="DUF1899"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01167">
        <property type="entry name" value="DUF1900"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00320">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50978">
        <property type="entry name" value="WD40 repeat-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00678">
        <property type="entry name" value="WD_REPEATS_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50082">
        <property type="entry name" value="WD_REPEATS_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50294">
        <property type="entry name" value="WD_REPEATS_REGION"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 4" id="KW-0677">Repeat</keyword>
    <keyword evidence="1 3 4" id="KW-0853">WD repeat</keyword>
    <feature type="domain" description="DUF1899" evidence="7">
        <location>
            <begin position="4"/>
            <end position="68"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="3">
        <location>
            <begin position="77"/>
            <end position="111"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="3">
        <location>
            <begin position="127"/>
            <end position="169"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="370"/>
            <end position="389"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="5">
        <location>
            <begin position="395"/>
            <end position="422"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022574"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="RuleBase" id="RU280818"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="SMART" id="SM01166"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015325.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="431" mass="48480" checksum="C29CA95FC5B986DF" modified="2019-05-08" version="1">MSRRVVRQSKFRHVFGQPVKADQMYEDIRVSKVTWDSSFCAVNPKFVAIIVESGGGGAFMVLPLAKTGRVDKNYPLVTGHTAPVLDIDWCPHNDNVIASASEDTTVMVWQIPDYVPMRNITEPIVTLEGHSKRVGIITWHPTARNVLLSAGGDNLLILWNVGTGEMLLTLDDMHGDVIYNVCWNRTGSLLVTTCKDKHVRVIDPRKQQVVAENFEEPIALQEMDTSNGVLLPFYDPDSSIVYLCGKGDSSIRYFEITAEAPYVHYLSTYSSKEPQRGMGFMPKRGLDVSKCEIARFFKLHERKCEPIVMTVPRKSDLFQDDLYPDTPGPEPALEAAEWLAGKDAEPVLVSLRDGYVPLKNRELKVTRKNILDNKPPMGPRRSHSTCDANFSRPSLEDLLEEIRSLRQTVQAQEKRISDLENKLCKFTNGTA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK78</accession>
    <name>A0A452HK78_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Coronin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="5">Belongs to the WD repeat coronin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HK78"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017501.1">
        <property type="protein sequence ID" value="ENSGAGP00000015322.1"/>
        <property type="gene ID" value="ENSGAGG00000011551.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007015">
        <property type="term" value="P:actin filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:000003">
        <property type="entry name" value="Coronin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015505">
        <property type="entry name" value="Coronin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015048">
        <property type="entry name" value="DUF1899"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019775">
        <property type="entry name" value="WD40_repeat_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036322">
        <property type="entry name" value="WD40_repeat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001680">
        <property type="entry name" value="WD40_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10856">
        <property type="entry name" value="CORONIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10856:SF23">
        <property type="entry name" value="CORONIN-6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08953">
        <property type="entry name" value="DUF1899"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00400">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16300">
        <property type="entry name" value="WD40_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01166">
        <property type="entry name" value="DUF1899"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01167">
        <property type="entry name" value="DUF1900"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00320">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50978">
        <property type="entry name" value="WD40 repeat-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00678">
        <property type="entry name" value="WD_REPEATS_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50082">
        <property type="entry name" value="WD_REPEATS_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50294">
        <property type="entry name" value="WD_REPEATS_REGION"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 6" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 5" id="KW-0677">Repeat</keyword>
    <keyword evidence="1 4 5" id="KW-0853">WD repeat</keyword>
    <feature type="domain" description="DUF1899" evidence="8">
        <location>
            <begin position="4"/>
            <end position="68"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="4">
        <location>
            <begin position="77"/>
            <end position="111"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="4">
        <location>
            <begin position="127"/>
            <end position="169"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="410"/>
            <end position="429"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="435"/>
            <end position="462"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022574"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="RuleBase" id="RU280818"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="SMART" id="SM01166"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015322.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="471" mass="53066" checksum="B53E99B90BEBD524" modified="2019-05-08" version="1">MSRRVVRQSKFRHVFGQPVKADQMYEDIRVSKVTWDSSFCAVNPKFVAIIVESGGGGAFMVLPLAKTGRVDKNYPLVTGHTAPVLDIDWCPHNDNVIASASEDTTVMVWQIPDYVPMRNITEPIVTLEGHSKRVGIITWHPTARNVLLSAGGDNLLILWNVGTGEMLLTLDDMHGDVIYNVCWNRTGSLLVTTCKDKHVRVIDPRKQQVVAEKVKPHEGARPIRAIFMADGKIFTTGFSRMSERQLALWDLENFEEPIALQEMDTSNGVLLPFYDPDSSIVYLCGKGDSSIRYFEITAEAPYVHYLSTYSSKEPQRGMGFMPKRGLDVSKCEIARFFKLHERKCEPIVMTVPRKSDLFQDDLYPDTPGPEPALEAAEWLAGKDAEPVLVSLRDGYVPLKNRELKVTRKNILDNKPPMGPRRSHSTCDANFSRPSLEDLLEEIRSLRQTVQAQEKRISDLENKLCKFTNGTA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK86</accession>
    <name>A0A452HK86_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">RRM domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the CELF/BRUNOL family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HK86"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017514.1">
        <property type="protein sequence ID" value="ENSGAGP00000015335.1"/>
        <property type="gene ID" value="ENSGAGG00000011562.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006397">
        <property type="term" value="P:mRNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd12631">
        <property type="entry name" value="RRM1_CELF1_2_Bruno"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12634">
        <property type="entry name" value="RRM2_CELF1_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12638">
        <property type="entry name" value="RRM3_CELF1_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.330:FF:000013">
        <property type="entry name" value="CUGBP Elav-like family member 1 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.330:FF:000015">
        <property type="entry name" value="CUGBP Elav-like family member 1 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.330:FF:000016">
        <property type="entry name" value="CUGBP Elav-like family member 1 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.330">
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034196">
        <property type="entry name" value="CELF1/2_RRM1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034198">
        <property type="entry name" value="CELF1/2_RRM2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034199">
        <property type="entry name" value="CELF1/2_RRM3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012677">
        <property type="entry name" value="Nucleotide-bd_a/b_plait_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035979">
        <property type="entry name" value="RBD_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000504">
        <property type="entry name" value="RRM_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24012">
        <property type="entry name" value="RNA BINDING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00076">
        <property type="entry name" value="RRM_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00360">
        <property type="entry name" value="RRM"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54928">
        <property type="entry name" value="RNA-binding domain, RBD"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50102">
        <property type="entry name" value="RRM"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="5" id="KW-0507">mRNA processing</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="7 9" id="KW-0694">RNA-binding</keyword>
    <feature type="domain" description="RRM" evidence="11">
        <location>
            <begin position="16"/>
            <end position="99"/>
        </location>
    </feature>
    <feature type="domain" description="RRM" evidence="11">
        <location>
            <begin position="108"/>
            <end position="188"/>
        </location>
    </feature>
    <feature type="domain" description="RRM" evidence="11">
        <location>
            <begin position="399"/>
            <end position="477"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="276"/>
            <end position="307"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00009621"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022664"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00176"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50102"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015335.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="484" mass="51561" checksum="935B9890029B87C9" modified="2019-05-08" version="1">MNGTLDHPDQPDLDAIKMFVGQVPRSWCEKDLRELFEQYGAVYEINVLRDRSQNPPQSKGCCFVTFYTRKAALEAQNALHNMKILPGMHHPIQMKPADSEKNNAVEDRKLFIGMISKKCNENDIRVMFSSFGQIEECRILRGPDGLSRGCAFVTFTTRAMAQTAIKAMHQAQTMEGCSSPIVVKFADTQKDKEQKRIAQQLQQQMQQISAASVWGNLAGLNTLGPQYLALLQQTAAASSGNLNTLSSLHPMGGLNAMQLQNLAALAAAASAAQNTPSGTAALTSSSSPLSVLTSSGSSPSSSSSSSVNPMASLGALQTLAGATAGLNVSSLAGMAALNGGLGSGGLSNGTGSTMEALTQAYSGIQQYAAAALPTLYSQSLLTQQSIGAAGSQKEGPEGANLFIYHLPQEFGDQDLLQMFMPFGNVVSAKVFIDKQTNLSKCFGFVSYDNPVSAQAAIQSMNGFQIGMKRLKVQLKRSKNDSKPY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK89</accession>
    <name>A0A452HK89_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">XLR/SYCP3/FAM9 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the XLR/SYCP3 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HK89"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017511.1">
        <property type="protein sequence ID" value="ENSGAGP00000015332.1"/>
        <property type="gene ID" value="ENSGAGG00000011560.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000795">
        <property type="term" value="C:synaptonemal complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051321">
        <property type="term" value="P:meiotic cell cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007286">
        <property type="term" value="P:spermatid development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051443">
        <property type="entry name" value="XLR/SYCP3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006888">
        <property type="entry name" value="XLR/SYCP3/FAM9_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19368">
        <property type="entry name" value="XLR/SCP3/FAM9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19368:SF15">
        <property type="entry name" value="XLR_SYCP3_FAM9 DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04803">
        <property type="entry name" value="Cor1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="XLR/SYCP3/FAM9" evidence="4">
        <location>
            <begin position="87"/>
            <end position="218"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="27"/>
            <end position="61"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="2">
        <location>
            <begin position="144"/>
            <end position="210"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="3">
        <location>
            <begin position="1"/>
            <end position="11"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00010283"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="Pfam" id="PF04803"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015332.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="240" mass="28094" checksum="821B3BA7D809D39C" modified="2019-05-08" version="1">MAPSGRKHSGKAGKPIQDDQALCAYDFQEEEKKNLSGSEEDIREGDTPVIEKHGKKRPSVTPHAVVEEDVGGEVQTMLERFGADINKALLAKRKRLEMYTKASLKTSNQKIEHVWKTQQEQRQKLNQEYSQQFLTLFQQWDMDMQKAEEQEEKLANMFRQQQKVFQQARIVQSQRLKTIKQLYEQFLKSMEELEKNHENLLAGAQNELRKEMAMLQKKIMMDTQQQEMATVRKSLQSMLF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK95</accession>
    <name>A0A452HK95_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="3">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HK95"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017526.1">
        <property type="protein sequence ID" value="ENSGAGP00000015345.1"/>
        <property type="gene ID" value="ENSGAGG00000011572.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006355">
        <property type="term" value="P:regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd07765">
        <property type="entry name" value="KRAB_A-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003655">
        <property type="entry name" value="aKRAB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001909">
        <property type="entry name" value="KRAB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036051">
        <property type="entry name" value="KRAB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050169">
        <property type="entry name" value="Krueppel_C2H2_ZnF"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23232:SF142">
        <property type="entry name" value="GASTRULA ZINC FINGER PROTEIN XLCGF57.1-LIKE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23232">
        <property type="entry name" value="KRAB DOMAIN C2H2 ZINC FINGER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01352">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00349">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109640">
        <property type="entry name" value="KRAB domain (Kruppel-associated box)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50805">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50806">
        <property type="entry name" value="KRAB_RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="KRAB-related" evidence="2">
        <location>
            <begin position="2"/>
            <end position="72"/>
        </location>
    </feature>
    <feature type="domain" description="KRAB" evidence="1">
        <location>
            <begin position="5"/>
            <end position="76"/>
        </location>
    </feature>
    <evidence type="ECO:0000259" key="1">
        <source>
            <dbReference type="PROSITE" id="PS50805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="PROSITE" id="PS50806"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015345.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="86" mass="9883" checksum="7024FC573619B291" modified="2019-05-08" version="1">LEGPVTFEDVAVYFTREEWALLDPTQRALYRAIMQENYENVTSLGFPVSKPDVISQLEQGEEPWVPDLQGSEEREILRVPLTFKAV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK99</accession>
    <name>A0A452HK99_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Latent-transforming growth factor beta-binding protein 1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="19">Latent-transforming growth factor beta-binding protein 2</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="20">Transforming growth factor beta-1-binding protein 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="25 26">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="26" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="25" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="25" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">Key regulator of transforming growth factor beta (TGFB1, TGFB2 and TGFB3) that controls TGF-beta activation by maintaining it in a latent state during storage in extracellular space. Associates specifically via disulfide bonds with the Latency-associated peptide (LAP), which is the regulatory chain of TGF-beta, and regulates integrin-dependent activation of TGF-beta. Outcompeted by LRRC32/GARP for binding to LAP regulatory chain of TGF-beta.</text>
    </comment>
    <comment type="function">
        <text evidence="14">May play an integral structural role in elastic-fiber architectural organization and/or assembly.</text>
    </comment>
    <comment type="subunit">
        <text evidence="16">Forms part of the large latent transforming growth factor beta precursor complex; removal is essential for activation of complex. Interacts with SDC4. Interacts (via C-terminal domain) with FBN1 (via N-terminal domain) in a Ca(+2)-dependent manner.</text>
    </comment>
    <comment type="subunit">
        <text evidence="17">Interacts with TGFB1; associates via disulfide bonds with the Latency-associated peptide chain (LAP) regulatory chain of TGFB1, leading to regulate activation of TGF-beta-1. LTBP1 does not bind directly to TGF-beta-1, the active chain of TGFB1. Interacts (via C-terminal domain) with FBN1 (via N-terminal domain). Interacts with FBN2. Interacts with ADAMTSL2. Interacts with EFEMP2.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="13">Belongs to the LTBP family.</text>
    </comment>
    <comment type="caution">
        <text evidence="21">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000017527.1">
        <property type="protein sequence ID" value="ENSGAGP00000015346.1"/>
        <property type="gene ID" value="ENSGAGG00000008780.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019838">
        <property type="term" value="F:growth factor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00054">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="12"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000005">
        <property type="entry name" value="Fibrillin 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000194">
        <property type="entry name" value="Latent transforming growth factor beta binding protein 2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000469">
        <property type="entry name" value="Latent transforming growth factor beta binding protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.290.10:FF:000004">
        <property type="entry name" value="latent-transforming growth factor beta-binding protein 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000019">
        <property type="entry name" value="latent-transforming growth factor beta-binding protein 1 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000046">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 1 isoform x2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000014">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000077">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 3 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.290.10:FF:000001">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 3 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.290.10:FF:000002">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 3 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000056">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 3 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000115">
        <property type="entry name" value="latent-transforming growth factor beta-binding protein 4 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000024">
        <property type="entry name" value="Putative latent-transforming growth factor beta-binding protein 2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000273">
        <property type="entry name" value="Putative latent-transforming growth factor beta-binding protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="18"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.290.10">
        <property type="entry name" value="TGF-beta binding (TB) domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026823">
        <property type="entry name" value="cEGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001881">
        <property type="entry name" value="EGF-like_Ca-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013032">
        <property type="entry name" value="EGF-like_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000152">
        <property type="entry name" value="EGF-type_Asp/Asn_hydroxyl_site"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018097">
        <property type="entry name" value="EGF_Ca-bd_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009030">
        <property type="entry name" value="Growth_fac_rcpt_cys_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049883">
        <property type="entry name" value="NOTCH1_EGF-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017878">
        <property type="entry name" value="TB_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036773">
        <property type="entry name" value="TB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052080">
        <property type="entry name" value="vWF_C/EGF_Fibrillin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47333:SF4">
        <property type="entry name" value="EGF-LIKE DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47333">
        <property type="entry name" value="VON WILLEBRAND FACTOR C AND EGF DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12662">
        <property type="entry name" value="cEGF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07645">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12661">
        <property type="entry name" value="hEGF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00683">
        <property type="entry name" value="TB"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="19"/>
    </dbReference>
    <dbReference type="SMART" id="SM00179">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="17"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57184">
        <property type="entry name" value="Growth factor receptor domain"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57581">
        <property type="entry name" value="TB module/8-cys domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00010">
        <property type="entry name" value="ASX_HYDROXYL"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01186">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01187">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51364">
        <property type="entry name" value="TB"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9 21" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="4 21" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="3" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11" id="KW-0340">Growth factor binding</keyword>
    <keyword evidence="6" id="KW-0358">Heparin-binding</keyword>
    <keyword evidence="12" id="KW-0379">Hydroxylation</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="26" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="60"/>
            <end position="92"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="258"/>
            <end position="290"/>
        </location>
    </feature>
    <feature type="domain" description="TB" evidence="24">
        <location>
            <begin position="405"/>
            <end position="448"/>
        </location>
    </feature>
    <feature type="domain" description="TB" evidence="24">
        <location>
            <begin position="525"/>
            <end position="577"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="685"/>
            <end position="726"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="727"/>
            <end position="763"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="770"/>
            <end position="809"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="850"/>
            <end position="890"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="975"/>
            <end position="1017"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="1048"/>
            <end position="1088"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="1133"/>
            <end position="1174"/>
        </location>
    </feature>
    <feature type="domain" description="TB" evidence="24">
        <location>
            <begin position="1236"/>
            <end position="1288"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="1310"/>
            <end position="1348"/>
        </location>
    </feature>
    <feature type="domain" description="TB" evidence="24">
        <location>
            <begin position="1410"/>
            <end position="1462"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="23">
        <location>
            <begin position="1584"/>
            <end position="1626"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="1"/>
            <end position="31"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="105"/>
            <end position="132"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="375"/>
            <end position="400"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="598"/>
            <end position="621"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="1516"/>
            <end position="1535"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="22">
        <location>
            <begin position="8"/>
            <end position="18"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="22">
        <location>
            <begin position="375"/>
            <end position="399"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="21">
        <location>
            <begin position="64"/>
            <end position="74"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="21">
        <location>
            <begin position="82"/>
            <end position="91"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="21">
        <location>
            <begin position="262"/>
            <end position="272"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="21">
        <location>
            <begin position="280"/>
            <end position="289"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022674"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023183"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023278"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00038081"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00058734"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00059743"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00062144"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00062844"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00072992"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00072997"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00075443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS51364"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015346.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1640" mass="177183" checksum="CC9F8E1E2AD5FFA3" modified="2019-05-08" version="1">KLSAPIHRSQSGASSQPCSPAGAGQARDPTPQRARLKCLEDQIQPLGWSHVSLFTPFHVPPAVCDPPCQNKGSCSRPQLCICRSGFQGAQCEEVVPERVYHPPGSVAALQPPAGSLPKRAGSAEREGSPRGAQTPILKQALAGTQGRIVFPVSRHTGSSRTVRRYPAGNTQLMSNALPNGNGHEQSSSGPHAAYQEHTLSLWGANLTEKIRKIKIVFTPTICKQTCRSGHCYNSCQKGDTTTLYSQGGQNHDPKSGFRIYFCQIPCLNGGRCVGRDECWCPSNSTGKFCHLPVPKLEKNQVGRVPKTLASGSIKQSMYTLPLSNQLASLNPSLVNVHIKHPPEATVQIHQVARVRGDLGQSEENSVEAVLLSQLPTGPQHSYSNGNSNSITTESGQQQRPPELMGRCFRETLHGQCANPLEGLAKFEDCCGSVGLFWGVNRCIPCPPRPAHPVIENRQVECPQGYKRLNRSHCQDINECLMLGLCKDTECVNTRGSFLCTCKPGTMLDPSRSRCVSDKAVSMEQGLCYRSAAEGVCALPLAQSITKQICCCSRVGKGWGRNCEKCPMLGSETFKEICPAGHGYTYSSSDIRLSMRKAEAEELPFSSEEQGESSDQTPTQAAKRLQLQEALSGGVTGTLARAGTSAAEGVLGCDATLVAGTADATQEPAPHPGQWPVTSPCLSLSEMDRCASVPDPCGPGTCVNLPDRYSCICGPGYQLHPGLPQCIDDDECMRDPCAGKGHCINSVGSYSCLCYPGYTLLASQGTQTCQDLNECEQPSMCQGGQCTNTPGSYHCECARGYIMGHTGQCEDIDECADPSTCPRGRCVNALGSYECLSCAAGYQPMNGRCVDDDECFTVGICAHGLCINLAGSFRCSCYHGYEVTADEKSCQDINECATHNACPMGLCINSEGSFSCAACGAGYTVSRDGSMCEDIDECSSATACPLGICTNTDGSFACRACDAGYMLSSNRFTCEDMNECLASEICSPNGECLNSHGSYFCICAPGFSNAAGGVTKCALDPGYRPGLSLGPVYTHRTASVLIFILSLPDVDECADKSRCLRGQCFNTEGSYRCLCENGFRHSQETDDCVDVDECVDFGDTVCGTWRCQNTPGSYRCVMGCQPGFHRTPLGDCIDIDECANETLCGSHAFCDNTDGSFRCLCDSGYENLPPGQDCVDVNECELMVAVCGTALCENVEGSFLCLCLSDHEEYNTEAGECRPRAGMGTGMAPRPAGSERKECYYHISDVQLCDSVLAKNTTKEECCCTEGAAWGDNCETHPCPILVEYSEICPSGKGYIPVEGTLLFGQTSYTDADECEMFGPEICRNGHCVNTVPGYKCFCRSGYLYDSSRLECVDQDECENEMSCVNGECLNTAGSFHCFCGLPLVLDVTRNRCINVSSRAEDPEEHEIHLDICWQRVSNYICNQPLRGRRTTYTECCCRYGEAWSQDCALCPHRASEDFAHLCNVARGETEREPGLQERPGYEYGPALEDSHYGLYSPDLGPYYNYLSPEYGAPDASFPQRELSSEADGPSVSTGHFDSFEGLQAEECGILNGCENGRCVRVREGYTCDCFDGFQLDMTHMACVDINECEEVSSSGPLCEGGTCENTEGSYRCNCSPGFVVVAGHCRAKETPVLMEDKWRC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMU8</accession>
    <name>A0A452IMU8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">ATP synthase lipid-binding protein</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="15">ATPase protein 9</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="13">ATPase subunit c</fullName>
        </alternativeName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Mitochondrion membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 16">Belongs to the ATPase C chain family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMU8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033246.1">
        <property type="protein sequence ID" value="ENSGAGP00000029272.1"/>
        <property type="gene ID" value="ENSGAGG00000021173.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031966">
        <property type="term" value="C:mitochondrial membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045259">
        <property type="term" value="C:proton-transporting ATP synthase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033177">
        <property type="term" value="C:proton-transporting two-sector ATPase complex, proton-transporting domain"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008289">
        <property type="term" value="F:lipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015078">
        <property type="term" value="F:proton transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015986">
        <property type="term" value="P:proton motive force-driven ATP synthesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd18182">
        <property type="entry name" value="ATP-synt_Fo_c_ATP5G3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.20.10:FF:000003">
        <property type="entry name" value="Atp synthase f complex subunit mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.20.10">
        <property type="entry name" value="F1F0 ATP synthase subunit C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="HAMAP" id="MF_01396">
        <property type="entry name" value="ATP_synth_c_bact"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000454">
        <property type="entry name" value="ATP_synth_F0_csu"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020537">
        <property type="entry name" value="ATP_synth_F0_csu_DDCD_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038662">
        <property type="entry name" value="ATP_synth_F0_csu_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002379">
        <property type="entry name" value="ATPase_proteolipid_c-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035921">
        <property type="entry name" value="F/V-ATP_Csub_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10031:SF54">
        <property type="entry name" value="ATP SYNTHASE F(0) COMPLEX SUBUNIT C2, MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10031">
        <property type="entry name" value="ATP SYNTHASE LIPID-BINDING PROTEIN, MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00137">
        <property type="entry name" value="ATP-synt_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00124">
        <property type="entry name" value="ATPASEC"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81333">
        <property type="entry name" value="F1F0 ATP synthase subunit C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00605">
        <property type="entry name" value="ATPASE_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0138">CF(0)</keyword>
    <keyword evidence="7 16" id="KW-0375">Hydrogen ion transport</keyword>
    <keyword evidence="9 16" id="KW-0406">Ion transport</keyword>
    <keyword evidence="10 16" id="KW-0446">Lipid-binding</keyword>
    <keyword evidence="12 16" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-0488">Methylation</keyword>
    <keyword evidence="11" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6 16" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 16" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="3 16" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="75"/>
            <end position="101"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="113"/>
            <end position="141"/>
        </location>
    </feature>
    <feature type="domain" description="V-ATPase proteolipid subunit C-like" evidence="17">
        <location>
            <begin position="76"/>
            <end position="138"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004225"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006704"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022547"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022781"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023121"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00029852"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00032304"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00033111"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="RuleBase" id="RU004221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF00137"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029272.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="142" mass="15286" checksum="84F37D4C444D9628" modified="2019-05-08" version="1">MRRTLLFLPFLLEYRVRNNSRLLCRPVSASLLSRPEARTDELSSTPAPQNALLQAVRREVHTSSSSWDIDTAAKFIGAGAATVGVAGSGAGIGTVFGSLIIGYARNPSLKQQLFSYAILGFALSEAMGLFCLMVAFLILFAM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMV1</accession>
    <name>A0A452IMV1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Nck-associated protein 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="6">Cell projection</location>
            <location evidence="6">Lamellipodium membrane</location>
            <topology evidence="6">Single-pass membrane protein</topology>
            <orientation evidence="6">Cytoplasmic side</orientation>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="7">Belongs to the HEM-1/HEM-2 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMV1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033174.1">
        <property type="protein sequence ID" value="ENSGAGP00000029210.1"/>
        <property type="gene ID" value="ENSGAGG00000019065.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031258">
        <property type="term" value="C:lamellipodium membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031209">
        <property type="term" value="C:SCAR complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030031">
        <property type="term" value="P:cell projection assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030866">
        <property type="term" value="P:cortical actin cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048812">
        <property type="term" value="P:neuron projection morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019137">
        <property type="entry name" value="Nck-associated_protein-1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12093">
        <property type="entry name" value="NCK-ASSOCIATED PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12093:SF11">
        <property type="entry name" value="NCK-ASSOCIATED PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09735">
        <property type="entry name" value="Nckap1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="1" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="5" id="KW-0966">Cell projection</keyword>
    <keyword evidence="4" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="3" id="KW-1133">Transmembrane helix</keyword>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="524"/>
            <end position="549"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="535"/>
            <end position="549"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00037839"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00037947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00039689"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029210.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="976" mass="111247" checksum="7E02F8E2E15204F5" modified="2019-05-08" version="1">VNNMEKRTVNFDLTKNYLDLIITYTTLMILLSRIEERKAIIGLYNYAHEMTHGASDREYPRLGQMIVDYENPLKKMMEEFVPHSKSLSDALISLQMVYPRRNLSADQWRNAQLLSLISAPSTMLNPAQSDTMPCEYLSLDAMEKWIIFGFILCHGILNSDATALNLWKLALQSSSCLSLFRDEVFHIHKAAEDLFVNIRGYNKRINDIRECKEAAVSHAGSMHRERRKFLRSALKELATVLSDQPGLLGPKALFVFMALSFARDEIIWLLRHADNMPKKSADDFIDKHIAELIFYMEELRAHVRKYGPVMQRYYVQYLSGFDAVVLNELVQNLSVCPEDESIIMSSFVNTMTSLSVKQVEDGEVFDFRGMRLDWFRLQAYTSVSKASLSLADHRELGKMMNTIIFHTKMVDSLVEMLVETSDLSIFCFYSRAFEKMFQQCLELPSQSRYSIAFPLLCTHFMSCTHELCPEERHHIGDRSLSLCNMFLDEMAKQARNLITDICTEQCTLSDQLLPKHCAKTISQAVNKKSKKQTGKKGEPEREKPGVESMRKNRLVVTNLDKLHTALSELCFSINYVPNMVVWEHTFTPREYLTSHLEIRFTKSIVGMTMYNQATQEIAKPSELLTSVRAYMTVLQSIENYVQIDITRVFNNVLLQQTQHLDSHGEPTITSLYTNWYLETLLRQVSNGHIAYFPAMKAFVNLPTENELTFNAEEYSDISEMRALSELLGPYGMKFLSESLMWHISSQVAELKKLVVENVEVLTQMRTSFDKPDQMAALFKRLSCGVLKRMTIIGVILSFRSLAQEALRDVAMNVYELSSAAGLPCEIDPALVVALSSQKSENISPEEEYKIACLLMVFVAVSLPTLASNVMSQYSPAIEGHCNNIHCLAKAINQIAAALFTIHKGSIEDQLTATKIRLKNQTKTLNICCNSDLPFRCTVIVQESPFLTMDLLESCFPYVLLRNAYHAVYKQSVTSSA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMX7</accession>
    <name>A0A452IMX7_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="7">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="3">Belongs to the RecA family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMX7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033285.1">
        <property type="protein sequence ID" value="ENSGAGP00000029306.1"/>
        <property type="gene ID" value="ENSGAGG00000021203.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000794">
        <property type="term" value="C:condensed nuclear chromosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140664">
        <property type="term" value="F:ATP-dependent DNA damage sensor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000150">
        <property type="term" value="F:DNA strand exchange activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003690">
        <property type="term" value="F:double-stranded DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003697">
        <property type="term" value="F:single-stranded DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070192">
        <property type="term" value="P:chromosome organization involved in meiotic cell cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000730">
        <property type="term" value="P:DNA recombinase assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042148">
        <property type="term" value="P:DNA strand invasion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006312">
        <property type="term" value="P:mitotic recombination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007131">
        <property type="term" value="P:reciprocal meiotic recombination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013632">
        <property type="entry name" value="Rad51_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020588">
        <property type="entry name" value="RecA_ATP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020587">
        <property type="entry name" value="RecA_monomer-monomer_interface"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22942:SF30">
        <property type="entry name" value="MEIOTIC RECOMBINATION PROTEIN DMC1_LIM15 HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22942">
        <property type="entry name" value="RECA/RAD51/RADA DNA STRAND-PAIRING FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08423">
        <property type="entry name" value="Rad51"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50162">
        <property type="entry name" value="RECA_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50163">
        <property type="entry name" value="RECA_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 3" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="4" id="KW-0472">Membrane</keyword>
    <keyword evidence="1 3" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="4">
        <location>
            <begin position="28"/>
            <end position="52"/>
        </location>
    </feature>
    <feature type="domain" description="RecA family profile 1" evidence="5">
        <location>
            <begin position="1"/>
            <end position="100"/>
        </location>
    </feature>
    <feature type="domain" description="RecA family profile 2" evidence="6">
        <location>
            <begin position="107"/>
            <end position="148"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="RuleBase" id="RU003422"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50162"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029306.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="151" mass="17295" checksum="280D92B6D29A569D" modified="2019-05-08" version="1">MRKLESSSYWYEPFGNFTLLHQLNAFRIIFSFAPSFSFTTFILFCIFTMQIIDSIMALFRVDFSGRGELAERQQKLAQMLSRLQKISEEYNVAVFVTNQMTADPGATMTFQADPKKPIGGHILAHASTTRISLRKGRGELRIAKIYDSCYL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMX9</accession>
    <name>A0A452IMX9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="5">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the STRIP family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMX9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033173.1">
        <property type="protein sequence ID" value="ENSGAGP00000029209.1"/>
        <property type="gene ID" value="ENSGAGG00000020767.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007010">
        <property type="term" value="P:cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040185">
        <property type="entry name" value="Far11/STRP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021819">
        <property type="entry name" value="Far11/STRP_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012486">
        <property type="entry name" value="Far11/STRP_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13239">
        <property type="entry name" value="PROTEIN REQUIRED FOR HYPHAL ANASTOMOSIS HAM-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13239:SF6">
        <property type="entry name" value="STRIATIN-INTERACTING PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11882">
        <property type="entry name" value="DUF3402"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07923">
        <property type="entry name" value="N1221"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01293">
        <property type="entry name" value="DUF3402"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01292">
        <property type="entry name" value="N1221"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Far11/STRP N-terminal" evidence="3">
        <location>
            <begin position="1"/>
            <end position="299"/>
        </location>
    </feature>
    <feature type="domain" description="Far11/STRP C-terminal" evidence="4">
        <location>
            <begin position="291"/>
            <end position="706"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="304"/>
            <end position="323"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="2">
        <location>
            <begin position="312"/>
            <end position="323"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007062"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="SMART" id="SM01292"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="SMART" id="SM01293"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029209.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="728" mass="84384" checksum="8221EAC4444E7473" modified="2019-05-08" version="1">EPEGPFFLCVSSPIELYSYTEEPEFTINRKCFEEDFQTQVQGKRWPELDLAKQKAYVMRLLDGLEVVNRDKRLKAARAVLYLAQGVFGECDVEAEVLHWSRHNNFLLFQMGTFSAFLELLNMEIDNSQACSSALRKPAISLADSTELRVLLSVMYLMVENIRVEAEADPPEWKSSRETFRTELSFPVHSEEPFALLLFTMVAKFCSGHAPHFPMKKVLLLLWKVLLFTLGGFEALQAMKTKKREELGLPPLPEDSIQVVRRMRAASPPTYTIELLDQQSSKRGHRSRRVRPLVKQDSLDIYNERDPFKNDEAGVDEEENDDVDSGIEGELDLMERDAIIQAGPAQRPPIERVSFPKGLPWAPKVRQKDIEHFLELCFFLPYLSACQGEEEVQETPCEMLYRAMLYNLPQYMIALLKILLAAAPTSKAKTDSINILADVLPEEMPITVLQSMKLGVDVNRHKEIIVKSISASLLLLLKHFKLNHIYQFEYVSQHLVFANCIPLILKFFNQNIMSYITAKNSISVLDYPHCTVHELPELTAESLEAGDNNQFCWRNLFSCINLLRLLNKLTKWKHSRTMMLVVFKSAPILKRALKVKQAMMQLYVLKLLKIQTKYLGRQWRKSNMKTMSAIYQKVRHRMNDDWAYGNDIDARPWDFQAEECTLRANIEAFNSRRYDKSQDSEFAPVDNCLQSVLGQPLELPEDFHYSYELWLEREVFSQPIRWEELLHYQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMY2</accession>
    <name>A0A452IMY2_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="7">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IMY2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033290.1">
        <property type="protein sequence ID" value="ENSGAGP00000029311.1"/>
        <property type="gene ID" value="ENSGAGG00000021205.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd16594">
        <property type="entry name" value="RING-HC_TRIM7-like_C-IV"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12888">
        <property type="entry name" value="SPRY_PRY_TRIM7_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.920:FF:000004">
        <property type="entry name" value="Butyrophilin subfamily 1 member A1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.920">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001870">
        <property type="entry name" value="B30.2/SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043136">
        <property type="entry name" value="B30.2/SPRY_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003879">
        <property type="entry name" value="Butyrophylin_SPRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006574">
        <property type="entry name" value="PRY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003877">
        <property type="entry name" value="SPRY_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050143">
        <property type="entry name" value="TRIM/RBCC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001841">
        <property type="entry name" value="Znf_RING"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017907">
        <property type="entry name" value="Znf_RING_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24103">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE TRIM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13765">
        <property type="entry name" value="PRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00622">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15227">
        <property type="entry name" value="zf-C3HC4_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01407">
        <property type="entry name" value="BUTYPHLNCDUF"/>
    </dbReference>
    <dbReference type="SMART" id="SM00589">
        <property type="entry name" value="PRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00184">
        <property type="entry name" value="RING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00449">
        <property type="entry name" value="SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57850">
        <property type="entry name" value="RING/U-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50188">
        <property type="entry name" value="B302_SPRY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00518">
        <property type="entry name" value="ZF_RING_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50089">
        <property type="entry name" value="ZF_RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0862">Zinc</keyword>
    <keyword evidence="2 4" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="RING-type" evidence="5">
        <location>
            <begin position="16"/>
            <end position="57"/>
        </location>
    </feature>
    <feature type="domain" description="B30.2/SPRY" evidence="6">
        <location>
            <begin position="88"/>
            <end position="266"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50188"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029311.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="266" mass="29668" checksum="D6B7D03B714B6CC7" modified="2019-05-08" version="1">MAAANPEKTLQDELTCPICLEYFKDPVSLDCDHNFCQACITQFWEGLATDVSCPQCRATFPQRNLRLNRQLRNVVEAVRELRLSSGREPETEGLCEKHKEPLKLFLNVTLDPDTAHPELVLSEDRKSVRYGDTWQDLPNNPERFDCWEYLLGCEGFTSGRHCWEVEVRGGGFWSVGVARESVGRKGWINLWGGQVQALTSPVTPLPLSRVPSRIRVCLDCDRGQVTFIGAGDEAPIFTFPPGSVPGGRIRPWLWAGPGGGTGNQPL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMY3</accession>
    <name>A0A452IMY3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Pre-mRNA-splicing factor ISY1 homolog</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="4">Identified in the spliceosome C complex. Component of the XAB2 complex, a multimeric protein complex composed of XAB2, PRPF19, AQR, ZNF830, ISY1, and PPIE. Identified in a pentameric intron-binding (IB) complex composed of AQR, XAB2, ISY1, ZNF830 and PPIE that is incorporated into the spliceosome as a preassembled complex. The IB complex does not contain PRPF19. Interacts with CPSF3; this interaction is in an RNA independent manner. Interacts with the microprocessor complex subunits DGCR8 and DROSHA; this interaction is in an RNA dependent manner.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the ISY1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMY3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033429.1">
        <property type="protein sequence ID" value="ENSGAGP00000029439.1"/>
        <property type="gene ID" value="ENSGAGG00000021274.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000350">
        <property type="term" value="P:generation of catalytic spliceosome for second transesterification step"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.660:FF:000001">
        <property type="entry name" value="pre-mRNA-splicing factor ISY1 homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.660">
        <property type="entry name" value="Helix hairpin bin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029012">
        <property type="entry name" value="Helix_hairpin_bin_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009360">
        <property type="entry name" value="Isy1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037200">
        <property type="entry name" value="Isy1_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13021">
        <property type="entry name" value="PRE-MRNA-SPLICING FACTOR ISY1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06246">
        <property type="entry name" value="Isy1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF140102">
        <property type="entry name" value="ISY1 domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0539">Nucleus</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="176"/>
            <end position="216"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="6">
        <location>
            <begin position="182"/>
            <end position="201"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="202"/>
            <end position="213"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00065310"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00070851"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029439.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="262" mass="30237" checksum="91F88C301BAC7AB5" modified="2019-05-08" version="1">MRKSFLFGERRPFLASECNELPKAEKWRRQIIGEISKKVAQIQNAGLGEFRIRDLNDEINKLLREKGHWEVRIKELGGPDYARVGPKMLDHEGKEVPGNRGYKYFGAAKDLPGVRELFEKEPLPPPRKTRAELMKAIDAEYYGYRDEDDGVLEPLEQEHEKKVIAEAVEKWKVEREARVAGGDEEEEEEINIYAVDDESDEESAKEKESEEGQQKFIAHVPVPSQQEIEEALVRRKKMELLQKYASETLMAQSEDAKRLLGL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMY4</accession>
    <name>A0A452IMY4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">protein kinase C</fullName>
            <ecNumber evidence="2">2.7.11.13</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the protein kinase superfamily. AGC Ser/Thr protein kinase family. PKC subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.13" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452IMY4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033178.1">
        <property type="protein sequence ID" value="ENSGAGP00000029214.1"/>
        <property type="gene ID" value="ENSGAGG00000021128.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004697">
        <property type="term" value="F:diacylglycerol-dependent serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000023">
        <property type="entry name" value="Protein kinase C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000080">
        <property type="entry name" value="Protein kinase C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000961">
        <property type="entry name" value="AGC-kinase_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017892">
        <property type="entry name" value="Pkinase_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24351">
        <property type="entry name" value="RIBOSOMAL PROTEIN S6 KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00433">
        <property type="entry name" value="Pkinase_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00133">
        <property type="entry name" value="S_TK_X"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51285">
        <property type="entry name" value="AGC_KINASE_CTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12 13" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="10" id="KW-0418">Kinase</keyword>
    <keyword evidence="6" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8 13" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="5" id="KW-0808">Transferase</keyword>
    <keyword evidence="11" id="KW-0862">Zinc</keyword>
    <keyword evidence="9" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="Protein kinase" evidence="15">
        <location>
            <begin position="175"/>
            <end position="433"/>
        </location>
    </feature>
    <feature type="domain" description="AGC-kinase C-terminal" evidence="16">
        <location>
            <begin position="434"/>
            <end position="504"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1"/>
            <end position="48"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="447"/>
            <end position="469"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="8"/>
            <end position="17"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="27"/>
            <end position="42"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="456"/>
            <end position="468"/>
        </location>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <position position="204"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00005490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00012429"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS51285"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029214.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="504" mass="57738" checksum="6B05677074EF1E36" modified="2019-05-08" version="1">MIQNPLKSTERPNHRITDYQGWKGPQEVSSPTPCSKQDQSLTDFCPRSLDGPLKDSTHRCRRKREWSRTVVFNSQAKNRLEPSVLDSLDTTNPACLSYVLFLLCPSRFKLLSQEEGEYFNVPVPPEGEEGNEELRQKFERARIGPGAKAAEERMTHAISKFDNNGNRDRMKLSDFNFLMVLGKGSFGKVMLSERKGTDELYAVKILKKDVVIQDDDVECTMVEKRVLALAGKPPFLTQLHSCFQTMDRLYFVMEYVNGGDLMYQIQQVGRFKEPHAVFYAAEIAIGLFFLQSKGIIYRDLKLDNVMLDNEGHIKIADFGMCKENIWDGATTKTFCGTPDYIAPEIIAYQPYGKSVDWWAFGVLLYEMLAGQAPFEGEDEDELFQSIMEHNVAYPKSMSKEAVAICKGLMTKHPGKRLGCGPEGERDIKEHAFFRYIDWEKLERKEVQPPFKPKAKDKRDTSNFDKEFTRQPVELTPTDKLFIMNLDQTEFAGFSYTNPEFVINV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMY8</accession>
    <name>A0A452IMY8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">UDENN FNIP1/2-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Lysosome membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the FNIP family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMY8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033434.1">
        <property type="protein sequence ID" value="ENSGAGP00000029444.1"/>
        <property type="gene ID" value="ENSGAGG00000021275.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005765">
        <property type="term" value="C:lysosomal membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042030">
        <property type="term" value="F:ATPase inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051087">
        <property type="term" value="F:protein-folding chaperone binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037545">
        <property type="entry name" value="DENN_FNIP1/2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028086">
        <property type="entry name" value="FNIP_C_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026156">
        <property type="entry name" value="FNIP_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028085">
        <property type="entry name" value="FNIP_mid_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028084">
        <property type="entry name" value="FNIP_N_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21634:SF12">
        <property type="entry name" value="FOLLICULIN-INTERACTING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21634">
        <property type="entry name" value="RE13835P"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14638">
        <property type="entry name" value="FNIP_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14637">
        <property type="entry name" value="FNIP_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14636">
        <property type="entry name" value="FNIP_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR02073">
        <property type="entry name" value="FOLLICULNIP1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51836">
        <property type="entry name" value="DENN_FNIP12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0458">Lysosome</keyword>
    <keyword evidence="5" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="UDENN FNIP1/2-type" evidence="8">
        <location>
            <begin position="36"/>
            <end position="1137"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="91"/>
            <end position="123"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="600"/>
            <end position="624"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="892"/>
            <end position="938"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="94"/>
            <end position="110"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="892"/>
            <end position="905"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004656"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007541"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023228"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51836"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029444.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1155" mass="128887" checksum="4B0FFA19DCB6FE6A" modified="2019-05-08" version="1">MPPTLFQKLFNKKYGLISPARDSREECVFSWPLPEFDPSQIRLIVYQDCERRGRNVLFDSSSKRKTEDVSVSKLCSDAQVRVFGKCCQLKPGGDSSSSLDSSINSSSLSSDTKEQCPKYQGSRCSSDANMLGEMMFGSVAMSYKGSTLKIHQIRSPPQLMLSKVFTARTGSSIYGSLNTLQDSLEFINQDSNTLKPDHSTIMNGLLGNIVHSNPMDMPGREQNEDRDSGIARSASLSSLLITPFPSPGSSLTRSCASSYQRRWRRSQTTSLENGVFPRWSVEESFNLSDDSSGLSPGIVRKKKIAIGVIFSLSKDEDENNKFHEFFFSHFPLFESHMNKLKSAIEQAMKMSRRSADASQRSLAYNRIVDALNEFRTTICNLYTMPRIGEPVWLTMMSGIPEKNQLCHRFMKEFTFLMENASKNQFLPALLTAILTNHLAWVPTVMPNGQPPIKIFLEKHSSQSVDMLAKTHPYNPLWAQLGDLYGAIGSPVRLAKTVVVGKRQDLVQRLLYFLTYFIRCSELQETHLLENGEDEAIVMPGTVITTTLEKGEVEESEYVLITIHRNRSNLLSSESEEMRTPNCSCKYCKCPLLLGQNTEGVSHQEREYAPNASKVEQETSSDESRTTCAADCQENAADIKQCKDKLISCLDTKLETVICTGSASVDKCAMADLGLEPTAGTWKTEESLESGNQASRVTRSTGIAVEKKPPDKLLSDAFPCSTAEAQTKVTFLIGDSMSPDSDIDLRSQAVVEQIARHHNQAAIEEEASAAQNCEAKQTVEDQNRDCGTAEPFPRVASEHQNWNPNPYNAESMSLFDEYFTDDGSIETRTIDDIPVQAAVEVLDHGSSLELSKKLGTMNSKLPSEFCKFMESVRQETCKNCFAEQDQRDKISICVPHGDKESSEKKVAPGSDWDIPRNESSDSALGDSESEDTGHDLTRQNSNYYVGEHEDWTEEDEIPFPGSKLVEVNSIQPSIANFGRSLLGGYCSSYVPDFVLQGVGNDEKLRHCLVSDLSHAVQHPVLDEPIAEAVCIIADTDKWTVQVASSQRRMIDNKLGKEVLVSSLVSNLLHSTLQLYKHNLSPNFCVMHLEDRLQELYFKSKMLSEYLKGQMRVHVKELGVVLGYDHNIVQYNFHLCFSHHKSSSHLLVGLSLPQGLH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN68</accession>
    <name>A0A452IN68_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="24">Dual specificity mitogen-activated protein kinase kinase 7</fullName>
            <ecNumber evidence="20">2.7.12.2</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="26">JNK-activating kinase 2</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="25">MAPK/ERK kinase 7</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="27">c-Jun N-terminal kinase kinase 2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="30 31">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="31" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="30" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="30" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="21">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.12.2"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="22">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.12.2"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="23">
            <text>L-tyrosyl-[protein] + ATP = O-phospho-L-tyrosyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10596"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.12.2"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="19">Belongs to the protein kinase superfamily. STE Ser/Thr protein kinase family. MAP kinase kinase subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.12.2" evidence="20"/>
    <dbReference type="AlphaFoldDB" id="A0A452IN68"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033524.1">
        <property type="protein sequence ID" value="ENSGAGP00000029524.1"/>
        <property type="gene ID" value="ENSGAGG00000021324.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019899">
        <property type="term" value="F:enzyme binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000287">
        <property type="term" value="F:magnesium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004708">
        <property type="term" value="F:MAP kinase kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004713">
        <property type="term" value="F:protein tyrosine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006915">
        <property type="term" value="P:apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043410">
        <property type="term" value="P:positive regulation of MAPK cascade"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006950">
        <property type="term" value="P:response to stress"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd06618">
        <property type="entry name" value="PKc_MKK7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000040">
        <property type="entry name" value="Dual specificity mitogen-activated protein kinase kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000214">
        <property type="entry name" value="Dual specificity mitogen-activated protein kinase kinase 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052468">
        <property type="entry name" value="Dual_spec_MAPK_kinase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47238:SF2">
        <property type="entry name" value="DUAL SPECIFICITY MITOGEN-ACTIVATED PROTEIN KINASE KINASE HEMIPTEROUS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47238">
        <property type="entry name" value="MITOGEN-ACTIVATED PROTEIN KINASE KINASE 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="14" id="KW-0007">Acetylation</keyword>
    <keyword evidence="8" id="KW-0053">Apoptosis</keyword>
    <keyword evidence="12" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="16" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11" id="KW-0418">Kinase</keyword>
    <keyword evidence="13" id="KW-0460">Magnesium</keyword>
    <keyword evidence="9" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="18" id="KW-0539">Nucleus</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="31" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="15" id="KW-0346">Stress response</keyword>
    <keyword evidence="7" id="KW-0808">Transferase</keyword>
    <keyword evidence="17" id="KW-0829">Tyrosine-protein kinase</keyword>
    <feature type="domain" description="Protein kinase" evidence="29">
        <location>
            <begin position="120"/>
            <end position="380"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="28">
        <location>
            <begin position="19"/>
            <end position="80"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="28">
        <location>
            <begin position="19"/>
            <end position="30"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="28">
        <location>
            <begin position="52"/>
            <end position="61"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022703"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023016"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023137"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00038035"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00038999"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00049014"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00049299"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00051693"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00073834"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00077430"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00081151"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="ARBA" id="ARBA00083185"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="29">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="30">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029524.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="31">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="420" mass="47782" checksum="2E4763ADCA1B5ABC" modified="2019-05-08" version="1">MAASSLEQKLSRLEAKLKQENREARRKIDLNLEISPARARPTLQLPLANDGSSRSSSSENSPQHQTHQPRRPLLGLPQPSFPIPRSMESIEIDQKLQEIMKQTGYLTIGGQRYQAEINDLENLGEIGSGTCGQVWKMRFKKTGHIIAVKQMRRSGNKEENKRILMDLDVVLKSHDCPYIVQCFGTFITNTDVFIAMELMGTCAEKLKKRIQGPIPERILGKMTVAIVKALYYLKEKHGVIHRDVKPSNILLDERGQIKLCDFGISGRLVDSKAKTRSAGCAAYMAPERIDPPDPTKPDYDIRADVWSLGISLVELATGQFPYKNCKTDFEVLTKVLQEDPPLLPNNMSFSVDFQSFVKDCLTKDHRKRPKYNKLLEHNFIKRYETLEVDVASWFKDVMAKTESPRTSSILSQHHLPFFTR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN70</accession>
    <name>A0A452IN70_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">non-specific serine/threonine protein kinase</fullName>
            <ecNumber evidence="3">2.7.11.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="12">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033388.1">
        <property type="protein sequence ID" value="ENSGAGP00000029402.1"/>
        <property type="gene ID" value="ENSGAGG00000021255.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.20.90:FF:000007">
        <property type="entry name" value="Serine/threonine-protein kinase WNK1 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000006">
        <property type="entry name" value="Serine/threonine-protein kinase WNK1 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.20.90:FF:000012">
        <property type="entry name" value="Serine/threonine-protein kinase WNK1 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000494">
        <property type="entry name" value="serine/threonine-protein kinase WNK2 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.20.90">
        <property type="entry name" value="Phosphatidylinositol 3-kinase Catalytic Subunit, Chain A, domain 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056865">
        <property type="entry name" value="CCTL2_WNK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024678">
        <property type="entry name" value="Kinase_OSR1/WNK_CCT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050588">
        <property type="entry name" value="WNK_Ser-Thr_kinase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13902">
        <property type="entry name" value="SERINE/THREONINE-PROTEIN KINASE WNK WITH NO LYSINE -RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24889">
        <property type="entry name" value="CCTL2_WNK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12202">
        <property type="entry name" value="OSR1_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="10" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="13" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9" id="KW-0418">Kinase</keyword>
    <keyword evidence="8" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="7" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Protein kinase" evidence="15">
        <location>
            <begin position="192"/>
            <end position="450"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1"/>
            <end position="184"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="606"/>
            <end position="625"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1031"/>
            <end position="1072"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1146"/>
            <end position="1185"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1210"/>
            <end position="1239"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1321"/>
            <end position="1341"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1799"/>
            <end position="1863"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="2141"/>
            <end position="2170"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="2182"/>
            <end position="2214"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="13">
        <location>
            <begin position="1890"/>
            <end position="1917"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="1"/>
            <end position="12"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="110"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="125"/>
            <end position="144"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="147"/>
            <end position="173"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="606"/>
            <end position="615"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="616"/>
            <end position="625"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="14">
        <location>
            <begin position="1053"/>
            <end position="1063"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="1157"/>
            <end position="1166"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="1167"/>
            <end position="1176"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="1819"/>
            <end position="1828"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="1840"/>
            <end position="1863"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="2146"/>
            <end position="2155"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012513"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00047899"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00048679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029402.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2214" mass="240019" checksum="615FF1A54890DA1B" modified="2019-05-08" version="1">METGCAASSAPPRGGGGASPAAARPPPADGSAAVGGLGRRSGAGMETAAVPERGPAPGLPPRGPGAREPRVGHLRFLRRSVVDSDQEEPPLELSEAEPRRIVLLSKTRRIIAERAKERHGEADLEPPAVEPAAEPKGAVPGVPATAEEQKELGRDAERKEPSDKDQSKSKKQEPEEEADMKAVSTSPDGRFLKFDIELGRGSFKTVYKGLDTETWVEVAWCELQDRKLTKVERQRFKEEAEMLKGLQHPNIVRFYDFWESCVKGKKCIVLVTELMTSGTLKTYLKRFKVMKPKVLRSWCRQILKGLLFLHTRTPPIIHRDLKCDNIFITGPTGSVKIGDLGLATLKRASFAKSVIGTPEFMAPEMYEEHYDESVDVYAFGMCMLEMATSEYPYSECQNAAQIYRKVTCGVKPASFEKVTDPEIKEIIGECICKNKEERYEIKDLLSHAFFAEDTGVRVELAEEDHGRKSSIALRLWVEDPKKLKGKPKDNGAIEFTFDLEKETPDDVAQEMVDSGFFHESDVKIVAKSIRDRVALIQWRRERIWPMIQSEDHQVSECLEKLKVPQAQQVQVTSLSHAGQQTLSALEEPEADQHLFQHNLPTSVTSVASDSTFDSGQGSTVYSDSQSSQQSVIYSSLPDSVPPAIQRVYSPPLSESQALPQSLQQLGHYQQPSVPGLTVAQAPPAVLSQRPQHYQEPAMTFPVIQPVTVASSQMGQSQPVLASQQIPQLPVIPAVTPLAGLDNLPPSLSDLPAANVPSVPAPSQYFTPTVIIPSLPMNPALPLASNPPALSMQAVNLPHTTVASLVLPCQTIVPNMSAATIPLLAVAPPGVSALPTHHAISHLSQQQMYQPAFQQIIQSDAPSPHHTQNMQAVSQQQLPPLPQSLQPSIIHPSEQTMSTSGVGNQIIGHPQYGLEIGAQMPVIPMQPSIVMPPPLQLQPEILPPCLGPESLSQLHGNLAAASATVPIDHCLPLQPTGLLQLQPDLSQPLGQQAPCSAIQSVTESTQEEHAIQEKQQTLSLCCESYMSPDVASGKEMSDSFEGVVGSGKQEGKPSKKHKKSTRVRSRQEKSSRPKLTILNVCNTGDKMVECQLETHNHKMVTFKFDLDGDAPEEIATYMVEKEFILQTEKETFIEQMKDIIDKAEDMLSEDTEGERSSDQGTSPQQDASRLEMNEENRQSQVKTPMYQQNVLHTGKRWFIICPVVENPTEEVPEFSPAVPQSKQAEGPDPEQSDDQQMGSAVTDTPSVFFCHQLPSQTSVFDSPCGTPAPLAQVPAAASSAVLQTKAEILAATLPGTAPHVPEQFVTNVGQPEYPPLHLVMPSQPDASAHNAASLEGPSDAQLAPKSSLQTLQEMDEDSCIPDVEISCCTVRPPSSILTVPAQGAEFCPVPALSDVVVLELQPAPQIPQVSEASQPGVMRHSFVNQLLPGGVVSDPLSLTGSQLQSPPQLPVAVSQQHTMLTQSQPVTCTGLGVGPLQPPCAVESDGEGPPRVDFADDTIKSLDEKLRNLLYQECVPTSAASAGTPDSFVPLEQGESELASLPPLNEELIPTLVLDSREPGLNVPDTFQEANAAQDQCVSLVPSAMPPYPVLASQALTVPFIPINPAAMETSAVLNNATATSHSRTEFRIQEPMLEKFETTSIDVSSSKVRVYETGSAANISSTSAVTTQGLQVAPASSTIPEQMETSRRNKEAKIKISSALTENLIQTSTTDREINNLQRNDFVDPGSCGKQSEVHGNSTPAKTIGRFSVISTHDELTLTSPHCLRYSAPPDVYLDELPSSPDLKTSVRRVQTASFLDVLSDHGSSDSGDESLRRKSVAAQLSPQSSIAANDITKKPASLLQRSGKTSSPGPDSPNGQGTKIPTINITSFHSQSSYMSSDNDSEFEDADMKKELQNLREKHMKEIAELQTQQKNEIEALFVRLGKPLPPNVGFLHSVPPTGRRRRTSKNKLKAGKLLNPLVQQLKTAPSNTSNLSVCKESLPKETTKTHAGSTEATAVTSSTSATFGNSVQTQQPCSVKASLSSDICSGLATEGGNAHGSSGQGWTVFHQTSERVTYKSSSKPRTRFLSGPMSLSIWSTLKRLCLGKDHSSRSSTNSLVTCPEPLPPPTLQVQVQANNSNNKKGTFTDDLHKLVDQWTSKTVGAAQSKPSLNQLKQNQKRQDMEPKANPMQAQNETCGILKMRMPLPPIDAGPAQLQYQAQGPMSAKEDEEEDVK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN72</accession>
    <name>A0A452IN72_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Carbamoyl-phosphate synthase [ammonia], mitochondrial</fullName>
            <ecNumber evidence="10">6.3.4.16</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="14">Carbamoyl-phosphate synthetase I</fullName>
        </alternativeName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="12">Involved in the urea cycle of ureotelic animals where the enzyme plays an important role in removing excess ammonia from the cell.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>hydrogencarbonate + NH4(+) + 2 ATP = carbamoyl phosphate + 2 ADP + phosphate + 2 H(+)</text>
            <dbReference type="Rhea" id="RHEA:18029"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:17544"/>
            <dbReference type="ChEBI" id="CHEBI:28938"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:58228"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="6.3.4.16"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Mitochondrion</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="6.3.4.16" evidence="10"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033320.1">
        <property type="protein sequence ID" value="ENSGAGP00000029340.1"/>
        <property type="gene ID" value="ENSGAGG00000021129.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004087">
        <property type="term" value="F:carbamoyl-phosphate synthase (ammonia) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004088">
        <property type="term" value="F:carbamoyl-phosphate synthase (glutamine-hydrolyzing) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006541">
        <property type="term" value="P:glutamine metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000050">
        <property type="term" value="P:urea cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd01744">
        <property type="entry name" value="GATase1_CPSase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd01423">
        <property type="entry name" value="MGS_CPS_I_III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.470.20:FF:000030">
        <property type="entry name" value="Carbamoyl-phosphate synthase 1, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.20:FF:000012">
        <property type="entry name" value="Carbamoyl-phosphate synthase 1, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.880:FF:000006">
        <property type="entry name" value="Carbamoyl-phosphate synthase 1, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1380:FF:000010">
        <property type="entry name" value="carbamoyl-phosphate synthase [ammonia], mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.1030.10:FF:000001">
        <property type="entry name" value="Carbamoyl-phosphate synthase large chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1490.20:FF:000001">
        <property type="entry name" value="Carbamoyl-phosphate synthase large chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.470.20:FF:000001">
        <property type="entry name" value="Carbamoyl-phosphate synthase large chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.20:FF:000002">
        <property type="entry name" value="Carbamoyl-phosphate synthase large chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.20">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.880">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1490.20">
        <property type="entry name" value="ATP-grasp fold, A domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.470.20">
        <property type="entry name" value="ATP-grasp fold, B domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1030.10">
        <property type="entry name" value="Carbamoyl-phosphate synthetase, large subunit oligomerisation domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1380">
        <property type="entry name" value="Methylglyoxal synthase-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011761">
        <property type="entry name" value="ATP-grasp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013815">
        <property type="entry name" value="ATP_grasp_subdomain_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036897">
        <property type="entry name" value="CarbamoylP_synth_lsu_oligo_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002474">
        <property type="entry name" value="CarbamoylP_synth_ssu_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036480">
        <property type="entry name" value="CarbP_synth_ssu_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029062">
        <property type="entry name" value="Class_I_gatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005479">
        <property type="entry name" value="CPAse_ATP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005483">
        <property type="entry name" value="CPSase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035686">
        <property type="entry name" value="CPSase_GATase1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006275">
        <property type="entry name" value="CPSase_lsu"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005480">
        <property type="entry name" value="CPSase_lsu_oligo"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058047">
        <property type="entry name" value="CPSase_preATP-grasp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017926">
        <property type="entry name" value="GATASE"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011607">
        <property type="entry name" value="MGS-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036914">
        <property type="entry name" value="MGS-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016185">
        <property type="entry name" value="PreATP-grasp_dom_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01369">
        <property type="entry name" value="CPSaseII_lrg"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF003671">
        <property type="entry name" value="PRK05294.1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF009455">
        <property type="entry name" value="PRK12815.1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11405:SF53">
        <property type="entry name" value="CARBAMOYL-PHOSPHATE SYNTHASE [AMMONIA], MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11405">
        <property type="entry name" value="CARBAMOYLTRANSFERASE FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25596">
        <property type="entry name" value="CPSase_L_D1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02786">
        <property type="entry name" value="CPSase_L_D2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02787">
        <property type="entry name" value="CPSase_L_D3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00988">
        <property type="entry name" value="CPSase_sm_chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00117">
        <property type="entry name" value="GATase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02142">
        <property type="entry name" value="MGS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00098">
        <property type="entry name" value="CPSASE"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00099">
        <property type="entry name" value="CPSGATASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM01096">
        <property type="entry name" value="CPSase_L_D3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01097">
        <property type="entry name" value="CPSase_sm_chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00851">
        <property type="entry name" value="MGS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48108">
        <property type="entry name" value="Carbamoyl phosphate synthetase, large subunit connection domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52021">
        <property type="entry name" value="Carbamoyl phosphate synthetase, small subunit N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52317">
        <property type="entry name" value="Class I glutamine amidotransferase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56059">
        <property type="entry name" value="Glutathione synthetase ATP-binding domain-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52335">
        <property type="entry name" value="Methylglyoxal synthase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52440">
        <property type="entry name" value="PreATP-grasp domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50975">
        <property type="entry name" value="ATP_GRASP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00866">
        <property type="entry name" value="CPSASE_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00867">
        <property type="entry name" value="CPSASE_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51273">
        <property type="entry name" value="GATASE_TYPE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51855">
        <property type="entry name" value="MGS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0021">Allosteric enzyme</keyword>
    <keyword evidence="7 15" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="4" id="KW-0436">Ligase</keyword>
    <keyword evidence="9" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="6 15" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="8" id="KW-0809">Transit peptide</keyword>
    <keyword evidence="2" id="KW-0835">Urea cycle</keyword>
    <feature type="domain" description="ATP-grasp" evidence="16">
        <location>
            <begin position="518"/>
            <end position="710"/>
        </location>
    </feature>
    <feature type="domain" description="ATP-grasp" evidence="16">
        <location>
            <begin position="1060"/>
            <end position="1251"/>
        </location>
    </feature>
    <feature type="domain" description="MGS-like" evidence="17">
        <location>
            <begin position="1322"/>
            <end position="1467"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004173"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022436"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022533"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022598"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00044063"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00047359"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00059545"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00074238"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00082675"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00409"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50975"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS51855"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029340.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1467" mass="161236" checksum="AC07DD17B5828E1A" modified="2019-05-08" version="1">MGDEGICALHHKWHGSRVPWNFLQTHHCLVFGSSLLAGLCWLFAPPAPCQQLSVCFSPRYTETLTDPSYRGQILTMVNPIVGNGGVPDTAALDEMGLSKFLESDGIKVSGLLVLDYSNDYSHWRAVKTLGQWLQEEKVRDLWATRPDTRHGISILRKLLSDSLSNGNQEREILEVKVYGKGNPIKVVAVDCGLKNNAIRLLVKRGAEVHLVPWNHDFTGMEYDGLFLSGGPGDPMQAQELIQNLQKVLKSDRKEPLFGVSMGNLITGIAAGATSYKMPMANRGQNQPVMNLSNGQAFITAQNHGYALQSNTLPAGWKPLFVNVNDQTNEGIMHETKPIFTAQFNPEASAGPTDTEFLFDSFISLIKKGKGTTVASVLPKARTVASRLEVSKVLILGSGGLSIGQAGEFDYSGSQAVKAMKEENVKTVLMNPNIASVQTNEVGLKQADTVYFLPITPQFVTEVIKAERPDGLILGMGGQTALNCGVELFRRGVLKEYGVKVLGTSVEAIMATEDRQLFSDKLTEINEKIAPSFAVESIEDALNAAEKIGYPVMIRSAYALGGLGSGICPDKERLLDLGTKAFAVTNQILVERSVVGWKEIEYEVVRDASDNCITVCNMENVDAMGVHTGDSIVVAPSQTLSNEEFQMLRESAIKVVRHLGIVGECNIQYALHPTSLEYCIIEVNARLSRSSALASKATGYPLAFIAAKIALGIPLPEIKNVVSGKTSACFEPSLDYMVAKIPRWDLDRFQGTSSQIGSSMKSVGEVMAIGRTFEECFQKALRMCHPSVDGFTSRLPMNKEWPANLDLQKELSDPSSIRVYAIAKALHDGVPVDDIHKLTAIDRWFIYKMRDIVNMEKSLKELSSESIPEETMRRAKQIGFSDKQIGKCLGVTEAQSRELRLKKNIKPSVKQIDTLAAEYPAITNYLYVTYNGQENDVKFDDHGMMVLGCGPYHIGSSVEFDWCAVSSIRTLRQLGHKTVVVNCNPETVSTDFDECDRLYFEELSLERILDIYQHEKCAGCIISVGGQIPNNLAVPLYKNGVKIMGTNPMQIDRAEDRSVFSAVLDELNVAQAPWKAVSTLKDAVEFANSVGYPCLLRPSYVLSGSAMNVVFTEEEMKKFLAEATRVSQEHPVVLTKFIEGAREVEMDAVAKAGKVVSHAISEHVEDAGVHSGDATLMLPTQNISQGALEKVKIITKKIARAFSISGPFNVQFLVRGNDVLVIECNLRASRSFPFVSKTIGVDFIDVATKVMIGEKIDESSLPTLERPIIPSEYVGIKAPMFSWPRLRDADPILRCEMASTGEVACFGESIHSAFLKAMLSTGFKFPQKGILVGIQHSFRPKFLGVAEQLHEEGFKLYATEATSNWLNANSVPTTPVAWPSQESHTTSLPAIRRLIKDGNIDLVINLPNSNTKFVHDNYVIRRTAVDSGIALLTNFQVTKLFAEAIKYSGKVDAKSLFHYRKLNTSNPP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN97</accession>
    <name>A0A452IN97_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Tubby-like protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 6">Belongs to the TUB family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IN97"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033558.1">
        <property type="protein sequence ID" value="ENSGAGP00000029551.1"/>
        <property type="gene ID" value="ENSGAGG00000021348.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005929">
        <property type="term" value="C:cilium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061512">
        <property type="term" value="P:protein localization to cilium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.20.90.10:FF:000001">
        <property type="entry name" value="Tubby-like protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.20.90.10">
        <property type="entry name" value="Tubby Protein, Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025659">
        <property type="entry name" value="Tubby-like_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000007">
        <property type="entry name" value="Tubby_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018066">
        <property type="entry name" value="Tubby_C_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005398">
        <property type="entry name" value="Tubby_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16517:SF20">
        <property type="entry name" value="TUBBY PROTEIN HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16517">
        <property type="entry name" value="TUBBY-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01167">
        <property type="entry name" value="Tub"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16322">
        <property type="entry name" value="Tub_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01573">
        <property type="entry name" value="SUPERTUBBY"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01574">
        <property type="entry name" value="TUBBYPROTEIN"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54518">
        <property type="entry name" value="Tubby C-terminal domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01200">
        <property type="entry name" value="TUB_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01201">
        <property type="entry name" value="TUB_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0964">Secreted</keyword>
    <feature type="domain" description="Tubby N-terminal" evidence="9">
        <location>
            <begin position="29"/>
            <end position="232"/>
        </location>
    </feature>
    <feature type="domain" description="Tubby C-terminal" evidence="8">
        <location>
            <begin position="250"/>
            <end position="493"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="36"/>
            <end position="67"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="102"/>
            <end position="135"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="157"/>
            <end position="238"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="7">
        <location>
            <begin position="192"/>
            <end position="202"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="216"/>
            <end position="237"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007129"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="RuleBase" id="RU361125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF01167"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF16322"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029551.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="498" mass="55921" checksum="8C09998552C9F0F5" modified="2019-05-08" version="1">MTSKHQADWIPYSVLDDEGSNLRQQKLDRQRALLEQKQKKKRQEPLMVQSNVDSRTRTRRMKHSEEQAPLVESYLNSNSSTIFHVQEAEQEDVKVVADAQVPKSTKKTKAAAASCQTGSTRKEKKGKHKGIDGPAAFQDDVQEVGSQIQILTVGQSSHNEDDGEMMASGQQQSRQDLRVTMQKKGISSSMNFDEEDEDDEDSSSSSQLNSNTRPGSATSKKSNKEAASAPSPSANEPVIDVDDLEEFAVRPAPQGVTVKCRITRDKKGMDRGMYPTYYLHLEREDGKKVFLLAGRKRKKSKTSNYLISIDPTDLSRGGESFIGKLRSNLMGTKFTVYDNGVNPMKTTSSLEASNLRQELAAICYETNVLGFKGPRKMSVVIPGMNMDHERVSIRPRNEHETLLARWQNKNTESVIELHNKTPVWNDDTQSYVLNFHGRVTQASVKNFQIIHDNDPDYIVMQFGRVAEDVFTMDYNYPMCALQAFAIALSSFDSKLACE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN98</accession>
    <name>A0A452IN98_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Carbamoyl-phosphate synthase [ammonia], mitochondrial</fullName>
            <ecNumber evidence="10">6.3.4.16</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="14">Carbamoyl-phosphate synthetase I</fullName>
        </alternativeName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="12">Involved in the urea cycle of ureotelic animals where the enzyme plays an important role in removing excess ammonia from the cell.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>hydrogencarbonate + NH4(+) + 2 ATP = carbamoyl phosphate + 2 ADP + phosphate + 2 H(+)</text>
            <dbReference type="Rhea" id="RHEA:18029"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:17544"/>
            <dbReference type="ChEBI" id="CHEBI:28938"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:58228"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="6.3.4.16"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Mitochondrion</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="6.3.4.16" evidence="10"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033323.1">
        <property type="protein sequence ID" value="ENSGAGP00000029343.1"/>
        <property type="gene ID" value="ENSGAGG00000021129.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004087">
        <property type="term" value="F:carbamoyl-phosphate synthase (ammonia) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004088">
        <property type="term" value="F:carbamoyl-phosphate synthase (glutamine-hydrolyzing) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006541">
        <property type="term" value="P:glutamine metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000050">
        <property type="term" value="P:urea cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd01744">
        <property type="entry name" value="GATase1_CPSase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd01423">
        <property type="entry name" value="MGS_CPS_I_III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.470.20:FF:000030">
        <property type="entry name" value="Carbamoyl-phosphate synthase 1, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.20:FF:000012">
        <property type="entry name" value="Carbamoyl-phosphate synthase 1, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.880:FF:000006">
        <property type="entry name" value="Carbamoyl-phosphate synthase 1, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1380:FF:000010">
        <property type="entry name" value="carbamoyl-phosphate synthase [ammonia], mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.1030.10:FF:000001">
        <property type="entry name" value="Carbamoyl-phosphate synthase large chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1490.20:FF:000001">
        <property type="entry name" value="Carbamoyl-phosphate synthase large chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.470.20:FF:000001">
        <property type="entry name" value="Carbamoyl-phosphate synthase large chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.20:FF:000002">
        <property type="entry name" value="Carbamoyl-phosphate synthase large chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.20">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.880">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1490.20">
        <property type="entry name" value="ATP-grasp fold, A domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.470.20">
        <property type="entry name" value="ATP-grasp fold, B domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1030.10">
        <property type="entry name" value="Carbamoyl-phosphate synthetase, large subunit oligomerisation domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1380">
        <property type="entry name" value="Methylglyoxal synthase-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011761">
        <property type="entry name" value="ATP-grasp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013815">
        <property type="entry name" value="ATP_grasp_subdomain_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036897">
        <property type="entry name" value="CarbamoylP_synth_lsu_oligo_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002474">
        <property type="entry name" value="CarbamoylP_synth_ssu_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036480">
        <property type="entry name" value="CarbP_synth_ssu_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029062">
        <property type="entry name" value="Class_I_gatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005479">
        <property type="entry name" value="CPAse_ATP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005483">
        <property type="entry name" value="CPSase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035686">
        <property type="entry name" value="CPSase_GATase1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006275">
        <property type="entry name" value="CPSase_lsu"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005480">
        <property type="entry name" value="CPSase_lsu_oligo"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058047">
        <property type="entry name" value="CPSase_preATP-grasp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017926">
        <property type="entry name" value="GATASE"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011607">
        <property type="entry name" value="MGS-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036914">
        <property type="entry name" value="MGS-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016185">
        <property type="entry name" value="PreATP-grasp_dom_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01369">
        <property type="entry name" value="CPSaseII_lrg"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF003671">
        <property type="entry name" value="PRK05294.1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF009455">
        <property type="entry name" value="PRK12815.1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11405:SF53">
        <property type="entry name" value="CARBAMOYL-PHOSPHATE SYNTHASE [AMMONIA], MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11405">
        <property type="entry name" value="CARBAMOYLTRANSFERASE FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25596">
        <property type="entry name" value="CPSase_L_D1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02786">
        <property type="entry name" value="CPSase_L_D2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02787">
        <property type="entry name" value="CPSase_L_D3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00988">
        <property type="entry name" value="CPSase_sm_chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00117">
        <property type="entry name" value="GATase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02142">
        <property type="entry name" value="MGS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00098">
        <property type="entry name" value="CPSASE"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00099">
        <property type="entry name" value="CPSGATASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM01096">
        <property type="entry name" value="CPSase_L_D3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01097">
        <property type="entry name" value="CPSase_sm_chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00851">
        <property type="entry name" value="MGS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48108">
        <property type="entry name" value="Carbamoyl phosphate synthetase, large subunit connection domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52021">
        <property type="entry name" value="Carbamoyl phosphate synthetase, small subunit N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52317">
        <property type="entry name" value="Class I glutamine amidotransferase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56059">
        <property type="entry name" value="Glutathione synthetase ATP-binding domain-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52335">
        <property type="entry name" value="Methylglyoxal synthase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52440">
        <property type="entry name" value="PreATP-grasp domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50975">
        <property type="entry name" value="ATP_GRASP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00866">
        <property type="entry name" value="CPSASE_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00867">
        <property type="entry name" value="CPSASE_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51273">
        <property type="entry name" value="GATASE_TYPE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51855">
        <property type="entry name" value="MGS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0021">Allosteric enzyme</keyword>
    <keyword evidence="7 15" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="4" id="KW-0436">Ligase</keyword>
    <keyword evidence="9" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="6 15" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="16" id="KW-0732">Signal</keyword>
    <keyword evidence="8" id="KW-0809">Transit peptide</keyword>
    <keyword evidence="2" id="KW-0835">Urea cycle</keyword>
    <feature type="signal peptide" evidence="16">
        <location>
            <begin position="1"/>
            <end position="40"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019506162" description="Carbamoyl-phosphate synthase [ammonia], mitochondrial" evidence="16">
        <location>
            <begin position="41"/>
            <end position="1455"/>
        </location>
    </feature>
    <feature type="domain" description="ATP-grasp" evidence="17">
        <location>
            <begin position="507"/>
            <end position="699"/>
        </location>
    </feature>
    <feature type="domain" description="ATP-grasp" evidence="17">
        <location>
            <begin position="1049"/>
            <end position="1240"/>
        </location>
    </feature>
    <feature type="domain" description="MGS-like" evidence="18">
        <location>
            <begin position="1311"/>
            <end position="1455"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004173"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022436"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022533"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022598"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00044063"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00047359"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00059545"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00074238"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00082675"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00409"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50975"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS51855"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029343.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1455" mass="159801" checksum="EA8B5EC48BA66453" modified="2019-05-08" version="1">MCLNLSLLGSFLQTHHCLVFGSSLLAGLCWLFAPPAPCQQLSVCFSPRYTETLTDPSYRGQILTMVNPIVGNGGVPDTAALDEMGLSKFLESDGIKVSGLLVLDYSNDYSHWRAVKTLGQWLQEEKVRDLWATRPDTRHGISILRKLLSDSLSNGNQEREILEVKVYGKGNPIKVVAVDCGLKNNAIRLLVKRGAEVHLVPWNHDFTGMEYDGLFLSGGPGDPMQAQELIQNLQKVLKSDRKEPLFGVSMGNLITGIAAGATSYKMPMANRGQNQPVMNLSNGQAFITAQNHGYALQSNTLPAGWKPLFVNVNDQTNEGIMHETKPIFTAQFNPEASAGPTDTEFLFDSFISLIKKGKGTTVASVLPKARTVASRLEVSKVLILGSGGLSIGQAGEFDYSGSQAVKAMKEENVKTVLMNPNIASVQTNEVGLKQADTVYFLPITPQFVTEVIKAERPDGLILGMGGQTALNCGVELFRRGVLKEYGVKVLGTSVEAIMATEDRQLFSDKLTEINEKIAPSFAVESIEDALNAAEKIGYPVMIRSAYALGGLGSGICPDKERLLDLGTKAFAVTNQILVERSVVGWKEIEYEVVRDASDNCITVCNMENVDAMGVHTGDSIVVAPSQTLSNEEFQMLRESAIKVVRHLGIVGECNIQYALHPTSLEYCIIEVNARLSRSSALASKATGYPLAFIAAKIALGIPLPEIKNVVSGKTSACFEPSLDYMVAKIPRWDLDRFQGTSSQIGSSMKSVGEVMAIGRTFEECFQKALRMCHPSVDGFTSRLPMNKEWPANLDLQKELSDPSSIRVYAIAKALHDGVPVDDIHKLTAIDRWFIYKMRDIVNMEKSLKELSSESIPEETMRRAKQIGFSDKQIGKCLGVTEAQSRELRLKKNIKPSVKQIDTLAAEYPAITNYLYVTYNGQENDVKFDDHGMMVLGCGPYHIGSSVEFDWCAVSSIRTLRQLGHKTVVVNCNPETVSTDFDECDRLYFEELSLERILDIYQHEKCAGCIISVGGQIPNNLAVPLYKNGVKIMGTNPMQIDRAEDRSVFSAVLDELNVAQAPWKAVSTLKDAVEFANSVGYPCLLRPSYVLSGSAMNVVFTEEEMKKFLAEATRVSQEHPVVLTKFIEGAREVEMDAVAKAGKVVSHAISEHVEDAGVHSGDATLMLPTQNISQGALEKVKIITKKIARAFSISGPFNVQFLVRGNDVLVIECNLRASRSFPFVSKTIGVDFIDVATKVMIGEKIDESSLPTLERPIIPSEYVGIKAPMFSWPRLRDADPILRCEMASTGEVACFGESIHSAFLKAMLSTGFKFPQKGILVGIQHSFRPKFLGVAEQLHEEGFKLYATEATSNWLNANSVPTTPVAWPSQESHTTSLPAIRRLIKDGNIDLVINLPNSNTKFVHDNYVIRRTAVDSGIALLTNFQVTKLFAEAIKYSGKVDAKSLFHYRKLNTRAL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGN3</accession>
    <name>A0A452GGN3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7 24">ATP-citrate synthase</fullName>
            <ecNumber evidence="6 24">2.3.3.8</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="20 24">ATP-citrate (pro-S-)-lyase</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="21 24">Citrate cleavage enzyme</fullName>
        </alternativeName>
    </protein>
    <organism evidence="27 28">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="28" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="27" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="27" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="23">Catalyzes the cleavage of citrate into oxaloacetate and acetyl-CoA, the latter serving as common substrate in multiple biochemical reactions in protein, carbohydrate and lipid metabolism.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="22">
            <text>oxaloacetate + acetyl-CoA + ADP + phosphate = citrate + ATP + CoA</text>
            <dbReference type="Rhea" id="RHEA:21160"/>
            <dbReference type="ChEBI" id="CHEBI:16452"/>
            <dbReference type="ChEBI" id="CHEBI:16947"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57288"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.3.3.8"/>
        </reaction>
        <physiologicalReaction direction="right-to-left" evidence="22">
            <dbReference type="Rhea" id="RHEA:21162"/>
        </physiologicalReaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subunit">
        <text evidence="5 24">Homotetramer.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytosol</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 24">In the C-terminal section; belongs to the succinate/malate CoA ligase alpha subunit family.</text>
    </comment>
    <comment type="similarity">
        <text evidence="4 24">In the N-terminal section; belongs to the succinate/malate CoA ligase beta subunit family.</text>
    </comment>
    <dbReference type="EC" id="2.3.3.8" evidence="6 24"/>
    <dbReference type="AlphaFoldDB" id="A0A452GGN3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000832.1">
        <property type="protein sequence ID" value="ENSGAGP00000000741.1"/>
        <property type="gene ID" value="ENSGAGG00000000607.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003878">
        <property type="term" value="F:ATP citrate synthase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006085">
        <property type="term" value="P:acetyl-CoA biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006101">
        <property type="term" value="P:citrate metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006633">
        <property type="term" value="P:fatty acid biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd06100">
        <property type="entry name" value="CCL_ACL-C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.230.10:FF:000004">
        <property type="entry name" value="ATP-citrate synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.470.110:FF:000001">
        <property type="entry name" value="ATP-citrate synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.261:FF:000003">
        <property type="entry name" value="ATP-citrate synthase subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.261:FF:000004">
        <property type="entry name" value="ATP-citrate synthase subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.720:FF:000024">
        <property type="entry name" value="Probable ATP-citrate synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.470.110">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.230.10">
        <property type="entry name" value="Cytochrome P450-Terp, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.720">
        <property type="entry name" value="NAD(P)-binding Rossmann-like Domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.261">
        <property type="entry name" value="Succinyl-CoA synthetase domains"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014608">
        <property type="entry name" value="ATP-citrate_synthase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017440">
        <property type="entry name" value="Cit_synth/succinyl-CoA_lig_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032263">
        <property type="entry name" value="Citrate-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016143">
        <property type="entry name" value="Citrate_synth-like_sm_a-sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056749">
        <property type="entry name" value="Citrate_synth_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002020">
        <property type="entry name" value="Citrate_synthase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036969">
        <property type="entry name" value="Citrate_synthase_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033847">
        <property type="entry name" value="Citrt_syn/SCS-alpha_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003781">
        <property type="entry name" value="CoA-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036291">
        <property type="entry name" value="NAD(P)-bd_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017866">
        <property type="entry name" value="Succ-CoA_synthase_bsu_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005811">
        <property type="entry name" value="SUCC_ACL_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016102">
        <property type="entry name" value="Succinyl-CoA_synth-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23118">
        <property type="entry name" value="ATP-CITRATE SYNTHASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23118:SF42">
        <property type="entry name" value="ATP-CITRATE SYNTHASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16114">
        <property type="entry name" value="Citrate_bind"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00285">
        <property type="entry name" value="Citrate_synt"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24948">
        <property type="entry name" value="Citrate_synth_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02629">
        <property type="entry name" value="CoA_binding"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00549">
        <property type="entry name" value="Ligase_CoA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF036511">
        <property type="entry name" value="ATP_citrt_syn"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00881">
        <property type="entry name" value="CoA_binding"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48256">
        <property type="entry name" value="Citrate synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56059">
        <property type="entry name" value="Glutathione synthetase ATP-binding domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51735">
        <property type="entry name" value="NAD(P)-binding Rossmann-fold domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52210">
        <property type="entry name" value="Succinyl-CoA synthetase domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01216">
        <property type="entry name" value="SUCCINYL_COA_LIG_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00399">
        <property type="entry name" value="SUCCINYL_COA_LIG_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01217">
        <property type="entry name" value="SUCCINYL_COA_LIG_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="18" id="KW-0007">Acetylation</keyword>
    <keyword evidence="15 24" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="8 24" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="10 24" id="KW-0444">Lipid biosynthesis</keyword>
    <keyword evidence="19 24" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="16 24" id="KW-0460">Magnesium</keyword>
    <keyword evidence="13 24" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="14 24" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="11" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="28" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="12 24" id="KW-0808">Transferase</keyword>
    <keyword evidence="17" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="CoA-binding" evidence="26">
        <location>
            <begin position="482"/>
            <end position="591"/>
        </location>
    </feature>
    <feature type="active site" description="Tele-phosphohistidine intermediate" evidence="25">
        <location>
            <position position="750"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004514"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005899"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00010719"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00011881"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00012639"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00015259"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022516"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00030151"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00030982"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00048669"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00093367"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="PIRNR" id="PIRNR036511"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="PIRSR" id="PIRSR036511-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="26">
        <source>
            <dbReference type="SMART" id="SM00881"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000741.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="28">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1091" mass="119807" checksum="813F4FB02D2CF129" modified="2019-05-08" version="1">MSAKAISEQTGKEFLYKYICTSSAIQNRFKYARVTSDTDWDRLTQDHPWLLTERLVVKPDQLIKRRGKLGLVGVNLDLGQVKAWLKLRLGKETTITKATGILKNFLIEPFVPHKQEEEFYVCIYAAREGDYVLFHHQGGVDVGDVDAKAQKLLVRVDEKLSESDVKKHLLVHALEEKKDVLASFISGLFNLYEDLYFTYLEINPLVVTKDGVYVLDLAAKIDATADYICKVKWGDVEFPPPFGREAYPEEAYIADLDAKSGASLKLTLLNPKGRIWTMVAGGGASVVYSDTICDLGGVNELANYGEYSGAPSEQQTYDYAKTILSLMTREKHPEGKILIIGGSIANFTNVAATFKGIVRAIKDYQGPLQEHGVRIFVRRGGPNYQEGLRVMGEVGKTTGIPIHVFGTETHMTAIVGMALGQRPIPNQPPAAAHTANFLLNASGSCSTPAPSRTASFSECKPDGIAPAKKAKPMTPPGKATTLFSCHTKAIVWGMQTRAVQGMLDFDYICSRDEPSVTAMVYPFTGDHKQKFYWGHKEILIPVFKNMSDAMRKHPEVDVLINFASLRSAYDSTIETMSFPQIRTIAIIAEGIPEALTRKLIKTADKKGVTIIGPATVGGIKPGCFKIGNTGGMLDNILASKLYRPGSVAYVSRSGGMSNELNNIISRATDGVYEGVAIGGDRYPGSTFMDHVLRYQDTPGVKMIVVLGEIGGTEEYKISRGIKEGRITKPVVCWCIGTCATMFSSEVQFGHAGACANQASETAVAKNQALKEAGVFVPQSFDELGDVIQSVYEDLVSKGVIEPAQEVPPPTVPMDYSWARELGLIRKPASFMTSICDERGQELIYAGMPITEVFKEEMGIGGVLGLLWFQRRLPKYACHFIEMCLMVTADHGPAVSGAHNTIVCARAGKDLVSSLTSGLLTIGDRFGGALDAAAKMFSKAFDSGIIPMEFVNKMKKEGKLIMGIGHRVKSINNPDMRVQILKDYVKQHFPATPLLDYALEVEKITTSKKPNLILNVDGFIGVAFVDVLRNCGSFTREEADEYIDIGALNGIFVLGRSMGFIGHYLDQKRLKQGLYRHPWDDISYVLPEHMTM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGP1</accession>
    <name>A0A452GGP1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase</fullName>
            <ecNumber evidence="15">2.7.4.24</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Bifunctional inositol kinase that acts in concert with the IP6K kinases IP6K1, IP6K2 and IP6K3 to synthesize the diphosphate group-containing inositol pyrophosphates diphosphoinositol pentakisphosphate, PP-InsP5, and bis-diphosphoinositol tetrakisphosphate, (PP)2-InsP4. PP-InsP5 and (PP)2-InsP4, also respectively called InsP7 and InsP8, regulate a variety of cellular processes, including apoptosis, vesicle trafficking, cytoskeletal dynamics, exocytosis, insulin signaling and neutrophil activation. Phosphorylates inositol hexakisphosphate (InsP6) at position 1 to produce PP-InsP5 which is in turn phosphorylated by IP6Ks to produce (PP)2-InsP4. Alternatively, phosphorylates PP-InsP5 at position 1, produced by IP6Ks from InsP6, to produce (PP)2-InsP4. Activated when cells are exposed to hyperosmotic stress.</text>
    </comment>
    <comment type="function">
        <text evidence="15">Bifunctional inositol kinase that acts in concert with the IP6K kinases to synthesize the diphosphate group-containing inositol pyrophosphates diphosphoinositol pentakisphosphate, PP-InsP5, and bis-diphosphoinositol tetrakisphosphate, (PP)2-InsP4. PP-InsP5 and (PP)2-InsP4, also respectively called InsP7 and InsP8, may regulate a variety of cellular processes, including apoptosis, vesicle trafficking, cytoskeletal dynamics, and exocytosis. Phosphorylates inositol hexakisphosphate (InsP6).</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>1D-myo-inositol hexakisphosphate + ATP = 1-diphospho-1D-myo-inositol 2,3,4,5,6-pentakisphosphate + ADP</text>
            <dbReference type="Rhea" id="RHEA:37459"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:58130"/>
            <dbReference type="ChEBI" id="CHEBI:74946"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.4.24"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="13">
            <dbReference type="Rhea" id="RHEA:37460"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="12">
            <text>5-diphospho-1D-myo-inositol 1,2,3,4,6-pentakisphosphate + ATP + H(+) = 1,5-bis(diphospho)-1D-myo-inositol 2,3,4,6-tetrakisphosphate + ADP</text>
            <dbReference type="Rhea" id="RHEA:10276"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:58628"/>
            <dbReference type="ChEBI" id="CHEBI:77983"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.4.24"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="12">
            <dbReference type="Rhea" id="RHEA:10277"/>
        </physiologicalReaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2 15">Cytoplasm</location>
            <location evidence="2 15">Cytosol</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 15">Belongs to the histidine acid phosphatase family. VIP1 subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.4.24" evidence="15"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000846.1">
        <property type="protein sequence ID" value="ENSGAGP00000000750.1"/>
        <property type="gene ID" value="ENSGAGG00000000614.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033857">
        <property type="term" value="F:5-diphosphoinositol pentakisphosphate 1-kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0052723">
        <property type="term" value="F:inositol hexakisphosphate 1-kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="RHEA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006020">
        <property type="term" value="P:inositol metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032958">
        <property type="term" value="P:inositol phosphate biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07061">
        <property type="entry name" value="HP_HAP_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.470.20:FF:000003">
        <property type="entry name" value="Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.11950:FF:000001">
        <property type="entry name" value="Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.11950:FF:000003">
        <property type="entry name" value="Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.11950">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.470.20">
        <property type="entry name" value="ATP-grasp fold, B domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1240">
        <property type="entry name" value="Phosphoglycerate mutase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033379">
        <property type="entry name" value="Acid_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000560">
        <property type="entry name" value="His_Pase_clade-2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037446">
        <property type="entry name" value="His_Pase_VIP1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029033">
        <property type="entry name" value="His_PPase_superfam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040557">
        <property type="entry name" value="VIP1_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12750">
        <property type="entry name" value="DIPHOSPHOINOSITOL PENTAKISPHOSPHATE KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12750:SF11">
        <property type="entry name" value="INOSITOL HEXAKISPHOSPHATE AND DIPHOSPHOINOSITOL-PENTAKISPHOSPHATE KINASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00328">
        <property type="entry name" value="His_Phos_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18086">
        <property type="entry name" value="PPIP5K2_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53254">
        <property type="entry name" value="Phosphoglycerate mutase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00616">
        <property type="entry name" value="HIS_ACID_PHOSPHAT_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10 15" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="5 15" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9 15" id="KW-0418">Kinase</keyword>
    <keyword evidence="11" id="KW-0472">Membrane</keyword>
    <keyword evidence="8 15" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7 15" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="VIP1 N-terminal" evidence="17">
        <location>
            <begin position="55"/>
            <end position="144"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="910"/>
            <end position="988"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1149"/>
            <end position="1228"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1255"/>
            <end position="1423"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1"/>
            <end position="15"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="16">
        <location>
            <begin position="1163"/>
            <end position="1180"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1193"/>
            <end position="1211"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1385"/>
            <end position="1407"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004514"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005609"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00033696"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00034629"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00037056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="RuleBase" id="RU365032"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF18086"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000750.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1423" mass="159092" checksum="998829B3AEACAA85" modified="2019-05-08" version="1">MSSLTTSHEGASSTPRFFVGTRDDETDFLGSNMKTDEIDFFEDDEESESPPERQIVVGICAMTKKSKSKPMTQILERLCKFEYITVVIMGEDVILNEPVENWPPCDCLISFHSKGFPLDKAVAYAKLCNPFLINDLDMQYYIQDRREVYRILQEEGIDLPRYAVLNRDPDKPDECSLVEGEDQVEVNGAVFPKPFVEKPVSAEDHNVYIYYPTSAGGGSQRLFRKIGSRSSVYSPESCVRKTGSYIYEEFMPTDGTDVKVYTVGPDYAHAEARKSPALDGKVERDSEGKEIRYPVMLTAMEKLVARKVCVAFKQTVCGFDLLRANGHSFVCDVNGFSFVKNSMKYYDDCTKILGNIIMRELAPQFQIPWSIPTEAEDIPIVPTTSGTMMELRCVIAVIRHGDRTPKQKMKMEVKHSRFFELFEKYDGYKTGKLKLKKPEQLQEVLDIARLLVVELGTHNDCEIEERKSKLEQLKTVLEMYGHFSGINRKVQLTYLPRGHPKASSEDEDARKEPSPSLLLVLKWGGELTPAGRVQAEELGRAFRCMYPGGQGDYAGFPGCGLLRLHSTYRHDLKIYASDEGRVQMTAAAFAKGLLALEGELTPILVQMVKSANMNGLLDSDSDSLSSCQHRVKARLHEIMQKDAKFCEEDYEKLAPTGSISLVSSMTFIQNPVEICDQVFALIENLTSQICKRLEDPKSADLQLYHSETLELMLQRWSKLERDFRMKNGRYDISKIPDIYDCIKYDVQHNGTLKLEGTAELFKLSKALADVIIPQEYGINKEEKLEIAIGFCLPLIKKIQLDLQRTHEDESVNKLHPLYSRGVLSPGRHVRTRLYFTSESHVHSLLSIFRYGGLLDESKDQQWKRAMDYLSAISELNYMTQIVIMLYEDNNKDPSSEERFHVELHFSPGVKGCEEDGSAPTGFGFRPASAENDDKKPNQGSMEDLSKVKSIDETDQAQERSPQPSEPICIQRRSPLIRNRKTGSMEVLSESSSKTGGYRLFSSYSRQSTEMKQSGLGSQCTGLFSTTVLGGSSSAPNLQDYVRSHGKKFTASLTYKDELLSMPAVKRFSVSFAKYPTNGFEGCSMVPTIYPLETLHNSLSLRQVNEFLTSVCRCYSSLHSRASTALFDSVIASQTPGDPFMPQRILSSSSLPLRQRSDKPPWYSSGPSSTVSSAGPSSPTSMENYARFGFTEKPSISPQISEERLNCQSPEQSEMEPSGEALELDLGGSGSLVENPLGEMPEQGLMRMPGLLEVPEPSKEELSEEVPEQGTMRVLGLSKTRQSGKTSEKDGSVVLEQGGEKPAGGTWELGETGKLEAGEIGLSGKISDSDEEGPAGEMLEPDVVRPVAPGETSELDEEEPVMQTLEPSEKGQPVEILEKEDEELSTDNALSDQQPFSHLYQDQQLSSEKSVEDEQPSSQQELQN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGQ0</accession>
    <name>A0A452GGQ0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Protein aurora borealis</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the BORA family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GGQ0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000833.1">
        <property type="protein sequence ID" value="ENSGAGP00000000742.1"/>
        <property type="gene ID" value="ENSGAGG00000000621.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019901">
        <property type="term" value="F:protein kinase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007088">
        <property type="term" value="P:regulation of mitotic nuclear division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060236">
        <property type="term" value="P:regulation of mitotic spindle organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023252">
        <property type="entry name" value="Aurora_borealis_protein"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14728">
        <property type="entry name" value="PROTEIN AURORA BOREALIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14728:SF2">
        <property type="entry name" value="PROTEIN AURORA BOREALIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15280">
        <property type="entry name" value="BORA_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR02038">
        <property type="entry name" value="AURORABORA"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="3" id="KW-0132">Cell division</keyword>
    <keyword evidence="4" id="KW-0498">Mitosis</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="304"/>
            <end position="328"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="314"/>
            <end position="326"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00010963"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00020055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022776"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000742.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="612" mass="67497" checksum="3008EA7CC7E95067" modified="2019-05-08" version="1">MGDAKETKMQITPKTPGRATVLNPFESPSDYYSLQEQIVSSPSVFKSTKSSSTPGKFRWSIDQLALINPVEIDSEDIRRQAMYLSHARIDKEMEDKRQKAIEEFFTKSLIVPSPWTDHEGKQVSQFHSSTKCIDLNNISPIGRQLTVQPGKSNAACQTVLSLPVDFNLEKILGEYFRTGEFADQSQESLSSSSLRRKLFLDENGSVSECLSLSPQSPCNNPPASLGVLCSIDISPARCRSPVETSSSGQFSSSPIQGGTRAYSLGSITSPTFPERSPANIASPTFSPIAFQIRKTPLSEQRKLTFRSPDVPSAPVSNRMTPPSTRSPYIDGCSPIKNCSPMRLGACRGTAQYQTSLIRIPFTLENHNEEAEDKENALPAEVLLPQMDTGVNLQQQDGDTFAHGTHLVVATVSISPDHSEACEQGLASLQDIEGLKENNTVDMVDSVEVSDENTWVKERVGNSSMPMTSFMTGITFSIESSHMCISPLAESSVIPCDNSSIQVDSGYNTQTCGSSIMDTVGAENNCRDNNVHSYEAQNKSQLSKTKEFPVLNYKDNQLLRTKSPESQPCFHKVKTYNTLFSQNATCNFSTWKHTNEHQIQGFHKNDEDPEMFS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGQ3</accession>
    <name>A0A452GGQ3_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="5">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the DONSON family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GGQ3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000855.1">
        <property type="protein sequence ID" value="ENSGAGP00000000757.1"/>
        <property type="gene ID" value="ENSGAGG00000000167.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033260">
        <property type="term" value="P:nuclear DNA replication"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024861">
        <property type="entry name" value="Donson"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12972">
        <property type="entry name" value="DOWNSTREAM NEIGHBOR OF SON"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12972:SF0">
        <property type="entry name" value="PROTEIN DOWNSTREAM NEIGHBOR OF SON"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR02064">
        <property type="entry name" value="DONSON"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="3" id="KW-0539">Nucleus</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00025806"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000757.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="440" mass="49585" checksum="CF4B0E5F703F0854" modified="2019-05-08" version="1">QENKSTWKEECSERDLHLPVSIPDVSSSPSTEFPADWSIKTRLLFMSSQPFTWAEHLKAQEEAQGFAQHCRATSVNFPQSIQEPKLSTELRCAFQQSLIYWLHPSLPWLQLFPRIGADRKTAGKTSLWSCDETLQQVLMSEWSVSFTSLYNLLKAKLCPYFYVCTYQFTVLFRTAGLAGSDVITAVVSPTTRGLREAMRNEGIEFSLPLVEETRSKTQKNSETNLDTEVNRVSDDESFSWLEEMGVQDEIKKPDAISVKLCKEKNEVQVDHKPESVALVKGTNTFMLLNFLINCKSLVAAAGPQAGLPPTLLSPVAFRGATMQTLKARSINVKTPVHSCYNDIFSLEITGPVMPHCLHTLTMLLKSAQRGAFSAVLYTHEPTAVFNTKIERILNKETMCKDLTKCGLHLKTLDQLIQLPTLGKSSIRLLEMRDYAYTWKS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGQ4</accession>
    <name>A0A452GGQ4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Endonuclease 8-like 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="1 2">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="2" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="1" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="1" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GGQ4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000834.1">
        <property type="protein sequence ID" value="ENSGAGP00000000743.1"/>
        <property type="gene ID" value="ENSGAGG00000000596.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019104">
        <property type="term" value="F:DNA N-glycosylase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003906">
        <property type="term" value="F:DNA-(apurinic or apyrimidinic site) endonuclease activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003676">
        <property type="term" value="F:nucleic acid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006284">
        <property type="term" value="P:base-excision repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.8.50">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035937">
        <property type="entry name" value="FPG_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010979">
        <property type="entry name" value="Ribosomal_uS13-like_H2TH"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22993:SF27">
        <property type="entry name" value="ENDONUCLEASE 8-LIKE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22993">
        <property type="entry name" value="FORMAMIDOPYRIMIDINE-DNA GLYCOSYLASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81624">
        <property type="entry name" value="N-terminal domain of MutM-like DNA repair proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46946">
        <property type="entry name" value="S13-like H2TH domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000313" key="1">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000743.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="3">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="388" mass="42806" checksum="535506114F748B77" modified="2019-05-08" version="1">MPEGPELHLAICYINTVCAGLIFSGKVEKSEVSRNPEVLFESNAYLISATSRGKEIKLTLTPLKEEKNTLPGPCVMLEYEKFRENVLRNLSDKAFNKPICEALLNQKFFNGIGDYLRAEILYRLKIPPFEKARTVLEALQHRKQSPDLSLSKKVKLKRENPDLLELCHAVPVEVIHMGKEAAQGAQPRVPALQQAMEPAGGGDSWLAKSTRLTQLGVVYMLEAVSKPGMSALTVKQCKRQPMLREDSCPTVQIVPWLGESEPVQRTSQEPGLTHMQPTLKLYGQSFNIPVPFAFPVSSPARKVHTVAPPACAPPAGNLKGPGAPGHCYCSSGLGPFKSSPEPWVVGVRQHRWAGWGRLPLPAPPLPPEALPLLGLPPVLVRIYYYFYP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGR1</accession>
    <name>A0A452GGR1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">Serine/threonine-protein kinase SMG1</fullName>
            <ecNumber evidence="5">2.7.11.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="16">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29035"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the PI3/PI4-kinase family.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000871.1">
        <property type="protein sequence ID" value="ENSGAGP00000000773.1"/>
        <property type="gene ID" value="ENSGAGG00000000611.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005694">
        <property type="term" value="C:chromosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000077">
        <property type="term" value="P:DNA damage checkpoint signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006281">
        <property type="term" value="P:DNA repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000184">
        <property type="term" value="P:nuclear-transcribed mRNA catabolic process, nonsense-mediated decay"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000723">
        <property type="term" value="P:telomere maintenance"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05170">
        <property type="entry name" value="PIKKc_SMG1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000136">
        <property type="entry name" value="serine/threonine-protein kinase SMG1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1010.10:FF:000010">
        <property type="entry name" value="serine/threonine-protein kinase SMG1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.1070.11:FF:000008">
        <property type="entry name" value="serine/threonine-protein kinase SMG1 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1070.11">
        <property type="entry name" value="Phosphatidylinositol 3-/4-kinase, catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1010.10">
        <property type="entry name" value="Phosphatidylinositol 3-kinase Catalytic Subunit, Chain A, domain 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050517">
        <property type="entry name" value="DDR_Repair_Kinase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003152">
        <property type="entry name" value="FATC_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000403">
        <property type="entry name" value="PI3/4_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036940">
        <property type="entry name" value="PI3/4_kinase_cat_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018936">
        <property type="entry name" value="PI3/4_kinase_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014009">
        <property type="entry name" value="PIK_FAT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031559">
        <property type="entry name" value="SMG1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035175">
        <property type="entry name" value="SMG1_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039414">
        <property type="entry name" value="SMG1_PIKKc"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11139">
        <property type="entry name" value="ATAXIA TELANGIECTASIA MUTATED ATM -RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11139:SF71">
        <property type="entry name" value="SERINE_THREONINE-PROTEIN KINASE SMG1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02260">
        <property type="entry name" value="FATC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00454">
        <property type="entry name" value="PI3_PI4_kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15785">
        <property type="entry name" value="SMG1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17229">
        <property type="entry name" value="SMG1_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01343">
        <property type="entry name" value="FATC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00146">
        <property type="entry name" value="PI3Kc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01345">
        <property type="entry name" value="Rapamycin_bind"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51189">
        <property type="entry name" value="FAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51190">
        <property type="entry name" value="FATC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00916">
        <property type="entry name" value="PI3_4_KINASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50290">
        <property type="entry name" value="PI3_4_KINASE_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="11" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="18" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="10" id="KW-0418">Kinase</keyword>
    <keyword evidence="13" id="KW-0464">Manganese</keyword>
    <keyword evidence="12" id="KW-0866">Nonsense-mediated mRNA decay</keyword>
    <keyword evidence="9" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="14" id="KW-0539">Nucleus</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="8" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="FAT" evidence="21">
        <location>
            <begin position="1236"/>
            <end position="1818"/>
        </location>
    </feature>
    <feature type="domain" description="PI3K/PI4K catalytic" evidence="20">
        <location>
            <begin position="2076"/>
            <end position="2415"/>
        </location>
    </feature>
    <feature type="domain" description="FATC" evidence="22">
        <location>
            <begin position="3581"/>
            <end position="3613"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1"/>
            <end position="69"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1104"/>
            <end position="1128"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1849"/>
            <end position="1872"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="3519"/>
            <end position="3544"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="18">
        <location>
            <begin position="1926"/>
            <end position="1953"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="25"/>
            <end position="38"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="51"/>
            <end position="65"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="1104"/>
            <end position="1114"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="3519"/>
            <end position="3531"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001936"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00011031"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012513"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023161"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023211"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00047899"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00048679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00068299"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50290"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS51189"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS51190"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000773.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="3613" mass="404862" checksum="3F2D4E24B8537DC3" modified="2019-05-08" version="1">MSRRAPGSRLSGIGTRQHYHPRSWNDWQPRTDSASADQDNLKYPASRDRGSSASYGLQPSNSAVVSRQRHDDIRVHTDIQNEEKGGYNVNGGSGENTYGRKSVGQELRVNNVTNPDFTSVQHGNRALATKDMRKSQERSLSYSDESRLSNLLRRITREDDRDRRLATVKQLKEFIQQPENKLVLVKQLDNILTAIHDVLNESSKLLQELRQEGACCLGLLCASLSYEAEKIFKWIFSKFSSSTKDEVKLLYLRATYKALETVGEKKAFSSVMQLVMTSLQSILENVDTPELLCKCVKCILLVSRCYPHIFSTNFRDTVDILVGWHIDHTQKPSLTQQVSGWLQSLEPFWVADLAFSTTLLGQFLEDMEAYAEDLSHVASGESVDEDIPPPSVSLPKLAALLRVFSTVVRSIGERFSPIRGQPITEAYVTDVLYRVMRCVTTANQVFFSEAVLTAANECAGVLLGSLDPSMTIRCDMVITYGLDQLENCQTCGTDYIISVLNLLTLIVEQINTKLPSTFVEKLFIPESKLLVLRYHKEKEVVAAAHAVYQAVLSLKNIPVLETAYRLILGEMTCALNSLLCSLQLPEACSEIQHDAFKNHVFNVANAKFVVIFDLSALTTIGNAKNSLIGMWALSPTVFALLSKNLMIVHSDIAVHFPAVQYAVLYTLYSHCTRHDHFISSSLSSSSPSLFDGAVISTVTTATKKHFSIILNLLGLLLKKDNLIQDTRKLLMTWALEVAVLMKKSETYAPLLSLPSFHKFCKGLLGNTLLEDVNICLQACSSLHALSSSLPDDLLQRCVDVCRVQLVHSGARIRQAFGKLLKSIPLDVVLRKDNWLERLFYSCQRLDKRDQSTIPRNLLKTDAVLWQWAVWEAAQFTVLSKLRTPLGRAQDTFQTIEGIIRSLAAHTLNPDQDVSQWTTADNDEGHSSNQLRLVLLLQYLENLEKLMYNAYEGCANALTSPPKVIRTFFYTNRQTCQDWLTRIRLSIMRVGLLAGQPAVTVRHGFDLLTEMKTNNISQGSELEVTIMTLVEALCELHCPEAIQGIAVWSSAVVGKSLSWINSVAQQAEGRFEKATVEYQEHLSSMTGVDCSVAGFDKSVLKLANSSSGNNASPKHSLNGEARKTVLTKPSDSSPEVINYLGNKACECYIAIADWAAVQEWQNAVHDLKKSTSNTSVNLKADFNYIKSLSSFECGELTDCTGQLELLPGENINLLTGGSKEKIDMKKLLPNMLSPDPREFQKAIEVQLLRSSVCLATAINHTEQEQKWQLITENIVKYLKQTSRIAVGPLRLSTLTVCQSLPVLSTLQLYCSSALENTVSSRLSSEDCLIPLFSDALRSCKQHDVRPWMHALRYTVYQNQLLEKLKEQTVSVRSHLMELGLTAAKFARKRGNVALATRLLAQCSEVQLGKTTTAQDLVQHFKKLSAPGQIDEKWSPELDIEKTKLLYTAGQSTHAMEMLSSCAISFCKSAKAEYAVAKSILTLAKWIQAEWKEISGQLKQVYRARQQQNLTGLSTLSKNIYTLIDLPAVNTIEEEYPRIESESTVNIGVGEPDFILGQLYHLSSVQAPEVAKSWAALASWAYRWGRKVVDNASQGGVRLLPKEKSEVQSLLPDNISEEDREKIYGILGQAVCRPAGIQDEDMTLQITENEDNEEDDMVDVIWRQLLTSCPWLSDLDENATEGLIKVWRKVVDRIFSLYKLSCSAYFTFLKLNAGQVPLDEDDPRLLLRNTSEQSTDDVIVMATLRLLRLLVKHAGELRQYLEQGLETTPTAPWRGIIPQLFSRLNHPEVYVRQSICNLLCRVAQDSPHLILYPAIVGTISLSSESQSSGNKLPSAIPTLLGNIQGEELLGSECEGGSPPASQESNKDEPKNGLNEDQAMMQDCYSKIVEKLSTANPTMVLQVQMLVAELRRVTVLWDELWLGVLLQQHMYVLRRIQQLEDEVKRVQNNNTLRKEEKIAIMREKHTALMKPIVFALEHVQSITAAPAETPHEKWFQDNYGDAIENALEKLKNPSNPAKPGSSWIPFKEVMLSLQQRAQKRASYLLHLEEISPWLAATMNTEIALPGEVSARDTVTIQSVGSTITILPTKTKPKKLLFLGSDGKNYPYLFKGLEDLHLDERIMQFLSIVNTMFATINRQETPRFHARHYSVTPLGTRSGLIQWVDGATPLFGLYKRWQQREAALQAQKAQDSYQTPQNPGIVPRPSELYYSKIGPALKAVGLSLDVSRRDWPLNVMKAVLEELMEATPPNLLAKELWSSCATPDEWWRVTQSYARSTAVMSMVGYIIGLGDRHLDNVLIDMTTGEVVHIDYNVCFEKGKSLRVPEKVPFRMTHNIETALGVTGVEGVFRLSCEQVLHIMRRGRETLLTLLEAFVYDPLVDWTAGGEAGFAGAVYGGGGQQAENKQSKREMERDITRSLFSSRVAEIKVNWFKNRDEMLVVLPKLDSSLEEYLSLQEQLTDIEKLQGKLLEEIEFLEGAEGVDHPSHTLQHRYSEHTQLQSQQRAVQEAIQVKLNEFEQWITHYQAAFNNLEATQLASLLQEISTQMDLGPPSYVPATAFLQNAGQAHLISQCEQLEGEVGALLQQRRSVLRGCLEQLHNYATVALQYPKAVFQKHRIEQWKTWMEELICNMTIERCQDLYGKYEMQYAPQPPPTVCQFITATEMTLQRYAADINTRLIRQVERLKQEAVTVPVCEEQLKEIERCIKVFLHENGEEGSLSLASVVISALCTLTRRNLMMEGAASSAGEQLVDLTSRDGAWFLEELCSMSGNVTCLVQLLKQCHLVPQDLDIPNPIEASEAVHLANGVYISLQELNSNFRQIIFPEALRCLMKGEYTLESMLHELDNLIEQTTDGVPLQTLVESLQAYLRNAAMGLEEETHTHYIDVARMLHAQYGELIQPRNGSVDETPKMSAGQMLLVAFDGMFAQVETAFGLLIEKLNKMEVPVAWRKIDIIREARSTQVNFFDDDDNRQVLEEIFFLKRLQTIKEFFRLCGTFSKTLSGSSSLEDQNTVNGPVQIVNVKALYRNSCFSEDQMAKPIKAFTADFVRQLLIGLPNQALGLTLCSFISALGVDIIAQVEAKDFGAESKVSVDDLCKKAVEHNIQIGKFSQLVMNRATVLASSYDTAWKKHDLVRRLETSISSCKTSLQRVQLHIAMFQWQHEDLLISRPQAMSITPPPRSAILASMKKKLHTLSQIEASIGTVQEKLAALEASIEQRLKWAGGANPALAPVLQDFDATIAERRNLVLKESQRASQVTFLCSNIIHFESLRTRTAEALSLDTALFELIKRCQQMCSFASQFNSSVSDLELRLLQRVGTNLEHPIGSSEWLLSAHKQLTQDMSTQRAVQAEKEQQIETVCETIQNLVDSIKTVLTGHNRQLGDVKHLLKAMAKDEEAALADGEDVPYENSVRQFLGEYKSWQDNIQTVLFTLVQAMGQVRNQEHVEMLQEITPTLKELKTQSQSVYNNLVSFASPLVTDTSNECSSPTSTATYQPTFAAAVRSNTGQKTQPEVMSQNARKPIQKNLATSADTPPSTVPGTGKGVACSPKKAVRDPKTGKAVQERNSYAVSVWKRVKAKLEGRDVDPNRRMSVAEQVDFVIKEATNLDNLAQLYEGWTAWV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGR2</accession>
    <name>A0A452GGR2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Phosphatase and actin regulator 4</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">Regulator of protein phosphatase 1 (PP1) required for neural tube and optic fissure closure, and enteric neural crest cell (ENCCs) migration during development. Acts as an activator of PP1. During neural tube closure, localizes to the ventral neural tube and activates PP1, leading to down-regulate cell proliferation within cranial neural tissue and the neural retina. Also acts as a regulator of migration of enteric neural crest cells (ENCCs) by activating PP1, leading to repression of the integrin signaling through the rho/rock pathway.</text>
    </comment>
    <comment type="subunit">
        <text evidence="2 10">Binds PPP1CA and actin.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="10">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="10">Cell projection</location>
            <location evidence="10">Lamellipodium</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="1 10">Belongs to the phosphatase and actin regulator family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GGR2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000869.1">
        <property type="protein sequence ID" value="ENSGAGP00000000771.1"/>
        <property type="gene ID" value="ENSGAGG00000000652.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030027">
        <property type="term" value="C:lamellipodium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003779">
        <property type="term" value="F:actin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008157">
        <property type="term" value="F:protein phosphatase 1 binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072542">
        <property type="term" value="F:protein phosphatase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030036">
        <property type="term" value="P:actin cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061386">
        <property type="term" value="P:closure of optic fissure"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048484">
        <property type="term" value="P:enteric nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:2001045">
        <property type="term" value="P:negative regulation of integrin-mediated signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001755">
        <property type="term" value="P:neural crest cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001843">
        <property type="term" value="P:neural tube closure"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051726">
        <property type="term" value="P:regulation of cell cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007266">
        <property type="term" value="P:Rho protein signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.1750">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.2130">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004018">
        <property type="entry name" value="RPEL_repeat"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12751:SF4">
        <property type="entry name" value="PHOSPHATASE AND ACTIN REGULATOR 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12751">
        <property type="entry name" value="PHOSPHATASE AND ACTIN REGULATOR PHACTR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02755">
        <property type="entry name" value="RPEL"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00707">
        <property type="entry name" value="RPEL"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51073">
        <property type="entry name" value="RPEL"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7 10" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="8 10" id="KW-0966">Cell projection</keyword>
    <keyword evidence="4 10" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="3 10" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="6 10" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <feature type="repeat" description="RPEL" evidence="9">
        <location>
            <begin position="64"/>
            <end position="89"/>
        </location>
    </feature>
    <feature type="repeat" description="RPEL" evidence="9">
        <location>
            <begin position="589"/>
            <end position="614"/>
        </location>
    </feature>
    <feature type="repeat" description="RPEL" evidence="9">
        <location>
            <begin position="627"/>
            <end position="652"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1"/>
            <end position="38"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="81"/>
            <end position="380"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="464"/>
            <end position="579"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="598"/>
            <end position="623"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="147"/>
            <end position="157"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="194"/>
            <end position="216"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="232"/>
            <end position="250"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="11">
        <location>
            <begin position="251"/>
            <end position="260"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="11">
        <location>
            <begin position="333"/>
            <end position="354"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="11">
        <location>
            <begin position="466"/>
            <end position="476"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="11">
        <location>
            <begin position="506"/>
            <end position="521"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="566"/>
            <end position="579"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="614"/>
            <end position="623"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00009795"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00011844"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00401"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="RuleBase" id="RU367131"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000771.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="708" mass="78171" checksum="67D43A5AA009E64D" modified="2019-05-08" version="1">MEENTAEEVDHPPSDMGMGADVLESGDTTPPTKRKSKFSGFGKIFKPWKWRKKKSSDKFKETSEVLERKISMRKPREELVKRGVLLEDPEQDGEEPDKFSHAALKNGHTVPIGGSGMSNLAHLEEEVGKKSSLRKSVPVEEPKKRPGSSSSNPSSEMEPPHEPHAPKQPLLPPKRPSSSSQEASDVQARDPVPASRTTRTASSSTVTAATSIAKTVNSTVAPCPAPRTVPPAFASTNTTATTSTTSVVPAKQPPIPPPKPVNRNSNPLIAELSQAVNSGMGSSKPSPPLPPKRGIPSNVAPSLESAAASKPPSDRIASATRPASIPVHITVSHPPPSPSPPLPSHVPPEPPRTPLPASSPAMEPQISLELPKETPPQEDLRSLEVPKRAAEQGFGEPHVPTWLPQIPLHIRIQQALTSPLPVTPPAEGSHRAHSLLFENDSLYEDNSTLGRARSLPVTIEMLKVPDDDEEEEEADQEGERSSGPHVYIGDAPSVTVIPRLMPQPLQEEEEEEEEGMSDSDSEGPILYKDEEDEEEDESHNSALANKVKRKDTLAMKLGTTATQQELEEKNAFPRKSKEEWNEIRQQIGTALIRRLSQRPTAEELEQRNILQQKNEADRQAEKREIKRRLTRKLSQRPTVAELQARKILRFHEYVEVTDAQDYDRRADKPWTKLTPADKAAIRKELNEFKSSEMQVHEDSKHFTRFHRP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GH00</accession>
    <name>A0A452GH00_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="30">ATP-dependent DNA helicase DDX11</fullName>
            <ecNumber evidence="28">5.6.2.3</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="32">DEAD/H-box protein 11</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="31">DNA 5'-3' helicase DDX11</fullName>
        </alternativeName>
    </protein>
    <organism evidence="35 36">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="36" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="35" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="35" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="29">
            <text>ATP + H2O = ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:13065"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="5.6.2.3"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>[4Fe-4S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:49883"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
            <location evidence="3">Cytoskeleton</location>
            <location evidence="3">Microtubule organizing center</location>
            <location evidence="3">Centrosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="5">Cytoplasm</location>
            <location evidence="5">Cytoskeleton</location>
            <location evidence="5">Spindle pole</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Midbody</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Nucleus</location>
            <location evidence="4">Nucleolus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the DEAD box helicase family. DEAH subfamily. DDX11/CHL1 sub-subfamily.</text>
    </comment>
    <dbReference type="EC" id="5.6.2.3" evidence="28"/>
    <dbReference type="AlphaFoldDB" id="A0A452GH00"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000978.1">
        <property type="protein sequence ID" value="ENSGAGP00000000861.1"/>
        <property type="gene ID" value="ENSGAGG00000000211.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030496">
        <property type="term" value="C:midbody"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005730">
        <property type="term" value="C:nucleolus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000922">
        <property type="term" value="C:spindle pole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051539">
        <property type="term" value="F:4 iron, 4 sulfur cluster binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043139">
        <property type="term" value="F:5'-3' DNA helicase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016818">
        <property type="term" value="F:hydrolase activity, acting on acid anhydrides, in phosphorus-containing anhydrides"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006281">
        <property type="term" value="P:DNA repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006260">
        <property type="term" value="P:DNA replication"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034085">
        <property type="term" value="P:establishment of sister chromatid cohesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd18788">
        <property type="entry name" value="SF2_C_XPD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:001050">
        <property type="entry name" value="ATP-dependent DNA helicase DDX11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000910">
        <property type="entry name" value="probable ATP-dependent DNA helicase DDX11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000909">
        <property type="entry name" value="Putative ATP-dependent RNA helicase DDX11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006555">
        <property type="entry name" value="ATP-dep_Helicase_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045028">
        <property type="entry name" value="DinG/Rad3-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014013">
        <property type="entry name" value="Helic_SF1/SF2_ATP-bd_DinG/Rad3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006554">
        <property type="entry name" value="Helicase-like_DEXD_c2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010614">
        <property type="entry name" value="RAD3-like_helicase_DEAD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013020">
        <property type="entry name" value="Rad3/Chl1-like"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00604">
        <property type="entry name" value="rad3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11472:SF41">
        <property type="entry name" value="ATP-DEPENDENT DNA HELICASE DDX11-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11472">
        <property type="entry name" value="DNA REPAIR DEAD HELICASE RAD3/XP-D SUBFAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06733">
        <property type="entry name" value="DEAD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13307">
        <property type="entry name" value="Helicase_C_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00488">
        <property type="entry name" value="DEXDc2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00491">
        <property type="entry name" value="HELICc2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51193">
        <property type="entry name" value="HELICASE_ATP_BIND_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0004">4Fe-4S</keyword>
    <keyword evidence="22" id="KW-0010">Activator</keyword>
    <keyword evidence="16" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="9" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="25" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="7" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="13" id="KW-0227">DNA damage</keyword>
    <keyword evidence="24" id="KW-0234">DNA repair</keyword>
    <keyword evidence="10" id="KW-0235">DNA replication</keyword>
    <keyword evidence="21" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="15" id="KW-0347">Helicase</keyword>
    <keyword evidence="14" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="18" id="KW-0408">Iron</keyword>
    <keyword evidence="19" id="KW-0411">Iron-sulfur</keyword>
    <keyword evidence="26" id="KW-0413">Isomerase</keyword>
    <keyword evidence="11" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="12" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="27" id="KW-0539">Nucleus</keyword>
    <keyword evidence="36" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="17" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="23" id="KW-0804">Transcription</keyword>
    <keyword evidence="20" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="Helicase ATP-binding" evidence="34">
        <location>
            <begin position="8"/>
            <end position="433"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="33">
        <location>
            <begin position="77"/>
            <end position="97"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="33">
        <location>
            <begin position="84"/>
            <end position="97"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001966"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004214"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004604"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004647"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00008435"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022485"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022705"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022806"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023014"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00023204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00023235"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="ARBA" id="ARBA00044969"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="ARBA" id="ARBA00048954"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="ARBA" id="ARBA00074489"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="31">
        <source>
            <dbReference type="ARBA" id="ARBA00079245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="32">
        <source>
            <dbReference type="ARBA" id="ARBA00080369"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="33">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="34">
        <source>
            <dbReference type="PROSITE" id="PS51193"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="35">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000861.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="36">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="885" mass="100193" checksum="569DA345F672CB32" modified="2019-05-08" version="1">RSAAVGKGRVNFPFPYTPYSIQEDFMANLYDVLEAGKVGIFESPTGTGKSLSLICGALSWLRDFEEKKKQEEARLLADEHSGQDKQQQPTSCSAEASNCQGSAGELVWITEFMQKKEEQAMVDRLKEEQIRRKKREDRLEKIRHNVQLKYASKRKRLEDDETQRLLQLSKEILSTELGSDVLEQLDHEEEELILAEYESDEEKKGRQTALLEFLQSSFSFSSQIYYCSRTHSQLAQFVHEVQKSPFGEEIRLVSLGSRQNLCVNEEVRHLGAVQLINDRCVEMQKNKHEESEGKKRRVSRAVCPFYSYEQMQFLRDEVLVKVKDIEQLVALGKETKACPYYGSRYAIPAAQLVVLPYQMLLHESTRCASGIKLKDQVVIIDEAHNLIDTITCIYSAEVRGSQLCCAHSQLLQYMERYRKRLKAKNLMYIKQILYLLERFVSILGGNVNQNPNAQSISQAGTELKSINDFLFQSQIDNINLFKLFGFVERYGATSVVKINKEKQKMAGLQNFLLTLHQGPDKGEFRCRVEDGQPKMASPLMQIEGFLSALTSANQDGRVILNRQGTVAKSSLKFLLLNPAVHFAKVLKECRAAIVAGGTMQPVSDFREQLLSCAIGADRVVEFSCGHVIPPDNILPIILCSGPSNQQLEFTYQKRDLPHMMDEMGRILGNLCNVVPGGVVCFFPSYEYEKQVYTHWEGTGLLTRLATKKKIFQEPKKANQVEQVLAEYTKCIKRCSQVGGHVTGALLFSVVGGKMSEGINFSDDLGRCVVMVGMPYPNIKSPELQEKMAYLDKTMPRTAGQPPSKMLIENLCMKAVNQSIGRAIRHQKDFASILLVDHRYARPSVFNKLPQWIKERTEIKYTFGLAFAALRKVSQKPPEGDGRALC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GH15</accession>
    <name>A0A452GH15_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">protein-tyrosine-phosphatase</fullName>
            <ecNumber evidence="2">3.1.3.48</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endomembrane system</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="3.1.3.48" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452GH15"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000980.1">
        <property type="protein sequence ID" value="ENSGAGP00000000863.1"/>
        <property type="gene ID" value="ENSGAGG00000000715.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030054">
        <property type="term" value="C:cell junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012505">
        <property type="term" value="C:endomembrane system"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019901">
        <property type="term" value="F:protein kinase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000020">
        <property type="entry name" value="Tyrosine-protein phosphatase non-receptor type 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000242">
        <property type="entry name" value="PTP_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR059011">
        <property type="entry name" value="PTPRR_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003595">
        <property type="entry name" value="Tyr_Pase_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008356">
        <property type="entry name" value="Tyr_Pase_KIM-con"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016334">
        <property type="entry name" value="Tyr_Pase_rcpt_R/non-rcpt_5"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46198">
        <property type="entry name" value="PROTEIN-TYROSINE-PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46198:SF2">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE R"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26155">
        <property type="entry name" value="PTPRR_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00102">
        <property type="entry name" value="Y_phosphatase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF001997">
        <property type="entry name" value="PTPRR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01778">
        <property type="entry name" value="KIMPTPASE"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00700">
        <property type="entry name" value="PRTYPHPHTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00194">
        <property type="entry name" value="PTPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00404">
        <property type="entry name" value="PTPc_motif"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50055">
        <property type="entry name" value="TYR_PHOSPHATASE_PTP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="8 10" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="6" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 10" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 10" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="217"/>
            <end position="239"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="11">
        <location>
            <begin position="381"/>
            <end position="583"/>
        </location>
    </feature>
    <feature type="active site" description="Phosphocysteine intermediate" evidence="9">
        <location>
            <position position="576"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004308"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00013064"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PIRSR" id="PIRSR001997-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000863.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="607" mass="68549" checksum="BAC2BC6A1173AF1F" modified="2019-05-08" version="1">MFSLLSTGCFSGDSDHFLAIHQRKSGKPVFIYHHVQSVEKSLDSDSQKIYKHNFHSSSHAKISNLHPAIIVKSAFPRRAYDPSLNLLAMTAQEPEVENLPIPATNVIVVALQMDVNKLNITLLRIFRQGVAAALGLLPQQIHINRLIGKKNCVEMFVSPTNRKPGISDALPSEEVLRSLNINVLHQSLSQFGITEVSPEKNVLQGQHEADKIWSKEGFYAVVIFLSIFVIVATCLMVLYRLKEKIQFSLQQDKEKKQEIHLSPIPLQTTRSEYKTANRMVQPEQAPKVVNVIVDPQGQCVPEIKPPLSASPSPFKMKPVGLQERRGSNVSLTLDMSSLGNVEPFVAVPTPREKVAMEYLQSASRVLTRQQLRDTVACSHLLQTEFMEIPMNFVDPKEIAIPSHGTKNRYKTILPNPLSRVCLKPKNPADSLSTYINANYIRGYGGKEKSFIATQGPMINTVNDFWQMVWQEDSPVIVMITKLKEKNEKCVLYWPEKRGIYGKVEVLIKDVQECENYTVRNLTLKQGSQSQNVKHYWYTSWPDHKTPYSAQPLLQLMLDVEEDRMASAGRGPVIVHCRGGMIQTSEQYEFVHHALSLYESRLSAEAVQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GH18</accession>
    <name>A0A452GH18_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Outer dynein arm-docking complex subunit 4</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="8">Tetratricopeptide repeat protein 25</fullName>
        </alternativeName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Cilium axoneme</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GH18"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001009.1">
        <property type="protein sequence ID" value="ENSGAGP00000000890.1"/>
        <property type="gene ID" value="ENSGAGG00000000687.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005930">
        <property type="term" value="C:axoneme"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003341">
        <property type="term" value="P:cilium movement"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036158">
        <property type="term" value="P:outer dynein arm assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000189">
        <property type="entry name" value="Tetratricopeptide repeat domain 25"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000274">
        <property type="entry name" value="Tetratricopeptide repeat domain 25"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.10">
        <property type="entry name" value="Tetratricopeptide repeat domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040111">
        <property type="entry name" value="ODAD4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011990">
        <property type="entry name" value="TPR-like_helical_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019734">
        <property type="entry name" value="TPR_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23040">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23040:SF1">
        <property type="entry name" value="OUTER DYNEIN ARM-DOCKING COMPLEX SUBUNIT 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13424">
        <property type="entry name" value="TPR_12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13176">
        <property type="entry name" value="TPR_7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00028">
        <property type="entry name" value="TPR"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48452">
        <property type="entry name" value="TPR-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50005">
        <property type="entry name" value="TPR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0966">Cell projection</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="5" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="4 9" id="KW-0802">TPR repeat</keyword>
    <feature type="repeat" description="TPR" evidence="9">
        <location>
            <begin position="354"/>
            <end position="387"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004430"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022803"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00034139"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00034143"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00339"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000890.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="530" mass="60544" checksum="7C668FF7FB728EF9" modified="2019-05-08" version="1">MLDLKLSRVLVCARCADKPTTSLTECFSVFAFQALQLQAGNKHCLVARSMCYLKLGDTENSLRDAEASLQNDKTFSKGLYQKAETLYTMGDFEFALVYYHRGHRLRPELQKFRLGIQKAQEAIDNSVGSPSSVKLENKGDLCFLSRQAESKKVKQKLQVKAPKKDQKQQSKVGPIRNEKTERQLLGELYADKAYLEKLLKDADLIKSSTKYGMTVEDLILGGINYLDTRTEFWQQQKPIYARVRDRKLMQQRWLPERKRKPSEVARYILKSMEEIDMLLTSSCPEASCKKAEHVLKTIQSWSEDEIPNKNELIGNLYSCIGNAQIEMGQMGAALQSHKMDLEIARENDLPDAVSRALDNMGRVYARIGKFQQAIDTWEEKIPLAKTSLEKTWLFHEIGRCYLELDEAAEAQDYGEKSLQSAEEEGDVEWQLNASVLVAQAQVKLQDFQSAVMNFEKALEKAKLIHNDAAQQAIIIALDDANKDFIEELRAEQRRERLESFRGTNSPGNWCVKQHRKFCFKSKIVKSSSPQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GH19</accession>
    <name>A0A452GH19_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">RING-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GH19"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000987.1">
        <property type="protein sequence ID" value="ENSGAGP00000000868.1"/>
        <property type="gene ID" value="ENSGAGG00000000730.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd16670">
        <property type="entry name" value="RING-H2_RNF215"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.40.10:FF:000009">
        <property type="entry name" value="E3 ubiquitin-protein ligase RNF130"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.50.30.30:FF:000031">
        <property type="entry name" value="RING finger protein 215"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.50.30.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001841">
        <property type="entry name" value="Znf_RING"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051073">
        <property type="entry name" value="ZNRF3_Arkadia_E3_ligases"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16200">
        <property type="entry name" value="RING ZINC FINGER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13639">
        <property type="entry name" value="zf-RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00184">
        <property type="entry name" value="RING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57850">
        <property type="entry name" value="RING/U-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50089">
        <property type="entry name" value="ZF_RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <keyword evidence="4 8" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="RING-type" evidence="9">
        <location>
            <begin position="223"/>
            <end position="264"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004370"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000868.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="275" mass="30738" checksum="BB91EA115B81A0C2" modified="2019-05-08" version="1">MGDAEPELEGEDSWIAVVPVGEEQAEIPRGSKEESFTVAVVNKMKRALVLGASALLILALNQNTIRELDVSQVLSKPVIVIQTSDNVTRLLGALLRGLRVTAKITYQAMLLENLGVTLTLWSTCGLSRGGLYGEWQGVICTGENSSRVQKYLQQLWNTILFIALILCTGVIVQAQRQSRQGQPDQDTELDLKQHILRRLSALKTRRYHPGKHSWNQARDIDSCVVCLDQFHKNQCLRVLPCSHEFHRDCVDPWLLLQQTCPLCKHNILGNCCGES</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GH21</accession>
    <name>A0A452GH21_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">C2H2-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GH21"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000988.1">
        <property type="protein sequence ID" value="ENSGAGP00000000869.1"/>
        <property type="gene ID" value="ENSGAGG00000000708.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006355">
        <property type="term" value="P:regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000557">
        <property type="entry name" value="zinc finger and SCAN domain-containing protein 29"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000002">
        <property type="entry name" value="Zinc finger protein 1 homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000759">
        <property type="entry name" value="zinc finger protein 16"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000295">
        <property type="entry name" value="zinc finger protein 19"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002343">
        <property type="entry name" value="Zinc finger protein 33A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000016">
        <property type="entry name" value="zinc finger protein 37 homolog"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000238">
        <property type="entry name" value="Zinc finger protein 485"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000367">
        <property type="entry name" value="Zinc finger protein 572"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000362">
        <property type="entry name" value="Zinc finger protein 606"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002134">
        <property type="entry name" value="Zinc finger protein 616"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000176">
        <property type="entry name" value="zinc finger protein 70"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001909">
        <property type="entry name" value="KRAB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050758">
        <property type="entry name" value="Znf_C2H2-type"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23234:SF8">
        <property type="entry name" value="C2H2-TYPE DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23234">
        <property type="entry name" value="ZNF44 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01352">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="14"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="9" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 11" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="90"/>
            <end position="117"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="135"/>
            <end position="162"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="163"/>
            <end position="190"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="191"/>
            <end position="218"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="219"/>
            <end position="246"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="247"/>
            <end position="274"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="275"/>
            <end position="302"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="303"/>
            <end position="330"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="331"/>
            <end position="358"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="359"/>
            <end position="386"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="387"/>
            <end position="414"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="415"/>
            <end position="442"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="443"/>
            <end position="470"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="471"/>
            <end position="498"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000869.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="503" mass="58450" checksum="EFB36C037CF6BB5F" modified="2019-05-08" version="1">METEWALLDPVQRALYRDVIPHLEYCVQFYSPMFKKDELKLEQVQRRATRMIQGMENLSYERRLKELEGNVSSLSDLITSGRINLEETRYRCHECGKSFSWSSTLVRHRRIHTGETPYTSTLNTHRRIHTGETPYTCTECGKSFSWRSDLVRHLRIHTGETPYTCTECGRRFSWRSHLIGHQRIHTGETPYTCTECGKSFSQCSNLITHQRIHTGEKPYSCSECGKSFSRSSHLIRHQIIHTGETPYTCTECGKSFNRSSTLNTHRRIHTGKLPYTCTECEKSFSRSSDLIGHQRIHTREKPYTCSECGKSFSCSSDLIRHQRIHTGETPYTCTECGRSFNRSSHLIGHQRIHTGEKPYTCCECEKSFSQSSHLIGHQRIHTGETPYTCTECGRRFSWRSHLIGHQRIHTGETPYTCTECGKSFSQCSNLITHQRIHTGETPYTCTECGKSFNRSSTLNTHRRIHTREMPYTCSECGQSFSQRSRLIRHQKMHRRENCNKSLD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GH22</accession>
    <name>A0A452GH22_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">POPDC1-3 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the popeye family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GH22"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001017.1">
        <property type="protein sequence ID" value="ENSGAGP00000000897.1"/>
        <property type="gene ID" value="ENSGAGG00000000745.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042383">
        <property type="term" value="C:sarcolemma"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030552">
        <property type="term" value="F:cAMP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007507">
        <property type="term" value="P:heart development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042391">
        <property type="term" value="P:regulation of membrane potential"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007519">
        <property type="term" value="P:skeletal muscle tissue development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051146">
        <property type="term" value="P:striated muscle cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018490">
        <property type="entry name" value="cNMP-bd_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006916">
        <property type="entry name" value="POPDC1-3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055272">
        <property type="entry name" value="POPDC1-3_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12101">
        <property type="entry name" value="POPEYE DOMAIN CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12101:SF15">
        <property type="entry name" value="POPEYE DOMAIN-CONTAINING PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04831">
        <property type="entry name" value="POPDC1-3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51206">
        <property type="entry name" value="cAMP-binding domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 7" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 7" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="47"/>
            <end position="70"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="77"/>
            <end position="97"/>
        </location>
    </feature>
    <feature type="domain" description="POPDC1-3" evidence="8">
        <location>
            <begin position="26"/>
            <end position="251"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="301"/>
            <end position="325"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007146"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF04831"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000897.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="354" mass="40260" checksum="9E1221C248F8C285" modified="2019-05-08" version="1">MSRNSSPWDQLLFQPPLCEGWKQNDEGAAYHLGNCILLLGYMGGSGIFGSIYIFSLLATGYICYALWGWLNACGLDIFIWNMLLVMACVLQLAHLVYRLHKDTFLEEFDLLYKTLCQPLQVPFEVYKEIVKCCEEKVLSLARDQNYAVEGKTPINCLSLLLSGRVRVSQDGQFLHYVFPYQFLDSPEWESLRPSEEGTFQVTLTAETDCSYITWPRRKLYLLLAKDRYIARLFSVFLGYDISEKLYALNDKLFAKFGLRFDIRLPSLHHVLGPASSQVEAEKELDEWEERAMPHRPMSTFVQTATSAPPPPPSARPKMPRPDSALLASGSLTRSYPCTLFKGRAPLAPTQTPEL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GH24</accession>
    <name>A0A452GH24_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Outer dynein arm-docking complex subunit 4</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="8">Tetratricopeptide repeat protein 25</fullName>
        </alternativeName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Cilium axoneme</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GH24"/>
    <dbReference type="Ensembl" id="ENSGAGT00000001022.1">
        <property type="protein sequence ID" value="ENSGAGP00000000902.1"/>
        <property type="gene ID" value="ENSGAGG00000000687.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005930">
        <property type="term" value="C:axoneme"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003341">
        <property type="term" value="P:cilium movement"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036158">
        <property type="term" value="P:outer dynein arm assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000189">
        <property type="entry name" value="Tetratricopeptide repeat domain 25"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000274">
        <property type="entry name" value="Tetratricopeptide repeat domain 25"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.10">
        <property type="entry name" value="Tetratricopeptide repeat domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040111">
        <property type="entry name" value="ODAD4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011990">
        <property type="entry name" value="TPR-like_helical_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019734">
        <property type="entry name" value="TPR_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23040">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23040:SF1">
        <property type="entry name" value="OUTER DYNEIN ARM-DOCKING COMPLEX SUBUNIT 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13424">
        <property type="entry name" value="TPR_12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13176">
        <property type="entry name" value="TPR_7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00028">
        <property type="entry name" value="TPR"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48452">
        <property type="entry name" value="TPR-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50005">
        <property type="entry name" value="TPR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0966">Cell projection</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="5" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="4 9" id="KW-0802">TPR repeat</keyword>
    <feature type="repeat" description="TPR" evidence="9">
        <location>
            <begin position="329"/>
            <end position="362"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004430"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022803"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00034139"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00034143"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00339"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000902.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="505" mass="57870" checksum="E6863E82C3569204" modified="2019-05-08" version="1">RRETESQDALQLQAGNKHCLVARSMCYLKLGDTENSLRDAEASLQNDKTFSKGLYQKAETLYTMGDFEFALVYYHRGHRLRPELQKFRLGIQKAQEAIDNSVGSPSSVKLENKGDLCFLSRQAESKKVKQKLQVKAPKKDQKQQSKVGPIRNEKTERQLLGELYADKAYLEKLLKDADLIKSSTKYGMTVEDLILGGINYLDTRTEFWQQQKPIYARVRDRKLMQQRWLPERKRKPSEVARYILKSMEEIDMLLTSSCPEASCKKAEHVLKTIQSWSEDEIPNKNELIGNLYSCIGNAQIEMGQMGAALQSHKMDLEIARENDLPDAVSRALDNMGRVYARIGKFQQAIDTWEEKIPLAKTSLEKTWLFHEIGRCYLELDEAAEAQDYGEKSLQSAEEEGDVEWQLNASVLVAQAQVKLQDFQSAVMNFEKALEKAKLIHNDAAQQAIIIALDDANKDFIEELRAEQRRERLESFRGTNSPGNWCVKQHRKFCFKSKIVKSSSPQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXP4</accession>
    <name>A0A452GXP4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">PDZ domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the peptidase S1C family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GXP4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007902.1">
        <property type="protein sequence ID" value="ENSGAGP00000006807.1"/>
        <property type="gene ID" value="ENSGAGG00000005493.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043065">
        <property type="term" value="P:positive regulation of apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012501">
        <property type="term" value="P:programmed cell death"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd06785">
        <property type="entry name" value="cpPDZ_HtrA-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.42.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.10.120">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001478">
        <property type="entry name" value="PDZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041489">
        <property type="entry name" value="PDZ_6"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036034">
        <property type="entry name" value="PDZ_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009003">
        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001940">
        <property type="entry name" value="Peptidase_S1C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001254">
        <property type="entry name" value="Trypsin_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22939">
        <property type="entry name" value="SERINE PROTEASE FAMILY S1C HTRA-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22939:SF127">
        <property type="entry name" value="SERINE PROTEASE HTRA2, MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17820">
        <property type="entry name" value="PDZ_6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00089">
        <property type="entry name" value="Trypsin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00834">
        <property type="entry name" value="PROTEASES2C"/>
    </dbReference>
    <dbReference type="SMART" id="SM00228">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50156">
        <property type="entry name" value="PDZ domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50494">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50106">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="2" id="KW-0645">Protease</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="PDZ" evidence="4">
        <location>
            <begin position="141"/>
            <end position="196"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00010541"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50106"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006807.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="209" mass="22879" checksum="B0B03FC4B396AA40" modified="2019-05-08" version="1">MRCEAGDLGYFSQGLCSLPLLLTTPGSLGFVSAPQWECRCWGLTLPLSLLLQFGNSGGPLVNLDGEVIGVNTMKVTSGISFAIPSDRLREFLQKEDQRKSSWFGSTEMKRRYIGVMMLTLTPSILAELKLRDSSFPDVSYGVLIHKVIIGSPAHQAGLKAGDVVQEINGQAVRRAEDIYEAVRTQNSLALLVRRGHDMLLVNVSPEVTE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXP6</accession>
    <name>A0A452GXP6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="19">tRNA-dihydrouridine(47) synthase [NAD(P)(+)]</fullName>
            <ecNumber evidence="19">1.3.1.-</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="19">tRNA-dihydrouridine synthase 3</fullName>
        </alternativeName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">Catalyzes the synthesis of dihydrouridine, a modified base, in various RNAs, such as tRNAs, mRNAs and some long non-coding RNAs (lncRNAs). Mainly modifies the uridine in position 47 (U47) in the D-loop of most cytoplasmic tRNAs. Also able to mediate the formation of dihydrouridine in some mRNAs, thereby regulating their translation.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="14">
            <text>5,6-dihydrouridine(47) in tRNA + NAD(+) = uridine(47) in tRNA + NADH + H(+)</text>
            <dbReference type="Rhea" id="RHEA:53364"/>
            <dbReference type="Rhea" id="RHEA-COMP:13539"/>
            <dbReference type="Rhea" id="RHEA-COMP:13540"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57540"/>
            <dbReference type="ChEBI" id="CHEBI:57945"/>
            <dbReference type="ChEBI" id="CHEBI:65315"/>
            <dbReference type="ChEBI" id="CHEBI:74443"/>
            <dbReference type="EC" id="1.3.1.89"/>
        </reaction>
        <physiologicalReaction direction="right-to-left" evidence="14">
            <dbReference type="Rhea" id="RHEA:53366"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>5,6-dihydrouridine(47) in tRNA + NADP(+) = uridine(47) in tRNA + NADPH + H(+)</text>
            <dbReference type="Rhea" id="RHEA:53360"/>
            <dbReference type="Rhea" id="RHEA-COMP:13539"/>
            <dbReference type="Rhea" id="RHEA-COMP:13540"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:65315"/>
            <dbReference type="ChEBI" id="CHEBI:74443"/>
            <dbReference type="EC" id="1.3.1.89"/>
        </reaction>
        <physiologicalReaction direction="right-to-left" evidence="17">
            <dbReference type="Rhea" id="RHEA:53362"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>a 5,6-dihydrouridine in mRNA + NAD(+) = a uridine in mRNA + NADH + H(+)</text>
            <dbReference type="Rhea" id="RHEA:69851"/>
            <dbReference type="Rhea" id="RHEA-COMP:14658"/>
            <dbReference type="Rhea" id="RHEA-COMP:17789"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57540"/>
            <dbReference type="ChEBI" id="CHEBI:57945"/>
            <dbReference type="ChEBI" id="CHEBI:65315"/>
            <dbReference type="ChEBI" id="CHEBI:74443"/>
        </reaction>
        <physiologicalReaction direction="right-to-left" evidence="15">
            <dbReference type="Rhea" id="RHEA:69853"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="16">
            <text>a 5,6-dihydrouridine in mRNA + NADP(+) = a uridine in mRNA + NADPH + H(+)</text>
            <dbReference type="Rhea" id="RHEA:69855"/>
            <dbReference type="Rhea" id="RHEA-COMP:14658"/>
            <dbReference type="Rhea" id="RHEA-COMP:17789"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="ChEBI" id="CHEBI:65315"/>
            <dbReference type="ChEBI" id="CHEBI:74443"/>
        </reaction>
        <physiologicalReaction direction="right-to-left" evidence="16">
            <dbReference type="Rhea" id="RHEA:69857"/>
        </physiologicalReaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 19">
            <name>FMN</name>
            <dbReference type="ChEBI" id="CHEBI:58210"/>
        </cofactor>
    </comment>
    <comment type="similarity">
        <text evidence="19">Belongs to the dus family. Dus3 subfamily.</text>
    </comment>
    <dbReference type="EC" id="1.3.1.-" evidence="19"/>
    <dbReference type="AlphaFoldDB" id="A0A452GXP6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007991.1">
        <property type="protein sequence ID" value="ENSGAGP00000006889.1"/>
        <property type="gene ID" value="ENSGAGG00000005527.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000007997.1">
        <property type="protein sequence ID" value="ENSGAGP00000006894.1"/>
        <property type="gene ID" value="ENSGAGG00000005527.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050660">
        <property type="term" value="F:flavin adenine dinucleotide binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0102265">
        <property type="term" value="F:tRNA-dihydrouridine47 synthase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006397">
        <property type="term" value="P:mRNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd02801">
        <property type="entry name" value="DUS_like_FMN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.20.20.70:FF:000067">
        <property type="entry name" value="tRNA-dihydrouridine(47) synthase [NAD(P)(+)]"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.1000.10:FF:000029">
        <property type="entry name" value="tRNA-dihydrouridine(47) synthase [NAD(P)(+)]"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.20.20.70">
        <property type="entry name" value="Aldolase class I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013785">
        <property type="entry name" value="Aldolase_TIM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035587">
        <property type="entry name" value="DUS-like_FMN-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018517">
        <property type="entry name" value="tRNA_hU_synthase_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000571">
        <property type="entry name" value="Znf_CCCH"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45846">
        <property type="entry name" value="TRNA-DIHYDROURIDINE(47) SYNTHASE [NAD(P)(+)]-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45846:SF1">
        <property type="entry name" value="TRNA-DIHYDROURIDINE(47) SYNTHASE [NAD(P)(+)]-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01207">
        <property type="entry name" value="Dus"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25585">
        <property type="entry name" value="zf-CCCH_DUS3L"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51395">
        <property type="entry name" value="FMN-linked oxidoreductases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01136">
        <property type="entry name" value="UPF0034"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50103">
        <property type="entry name" value="ZF_C3H1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 19" id="KW-0285">Flavoprotein</keyword>
    <keyword evidence="3 19" id="KW-0288">FMN</keyword>
    <keyword evidence="6 18 19" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="4" id="KW-0507">mRNA processing</keyword>
    <keyword evidence="12" id="KW-0520">NAD</keyword>
    <keyword evidence="10" id="KW-0521">NADP</keyword>
    <keyword evidence="11 19" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="5 19" id="KW-0819">tRNA processing</keyword>
    <keyword evidence="9 18 19" id="KW-0862">Zinc</keyword>
    <keyword evidence="8 18 19" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C3H1-type" evidence="21">
        <location>
            <begin position="47"/>
            <end position="77"/>
        </location>
    </feature>
    <feature type="zinc finger region" description="C3H1-type" evidence="18">
        <location>
            <begin position="47"/>
            <end position="77"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="1"/>
            <end position="43"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="165"/>
            <end position="190"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="20">
        <location>
            <begin position="1"/>
            <end position="26"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="20">
        <location>
            <begin position="27"/>
            <end position="43"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="20">
        <location>
            <begin position="169"/>
            <end position="181"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001917"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022630"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022643"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022664"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022694"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022857"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023027"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00045365"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00048266"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00048342"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00049447"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00049513"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="RuleBase" id="RU291113"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50103"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006889.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="589" mass="66504" checksum="F385EE12874937D4" modified="2019-05-08" version="1">MKLDGGDQVRKQGSQEEVLKEEEKKPQERKRARGQNKSRPCMKPKHYEKDRLCQSVIQECAEKCYFGSQCRFLHDVGEYMAVKPSDLGDSCILFKTFGKCIYGVTCRFARAHLGEDYRNIINGDLVKQWEGKSLVRNSLSKELQQRLRKKKVCFDKSSEYLSHLGKPSSAGNAGNKATGNSTEEKGMTNCTASKGNAEGLDVLPPGEEAPECLTVSEKGEASESMVLQSSCLVNASKDSTLKTIGPVTDEDVIKLRLCEKKRLDIQGKLYLAPLTTCGNLPFRRICKRFGADVTCGEMAVCTNLLQGQSSEWALLKRHYTEDLFGVQLEGAFPDTMTRCAELLNQTTEMDFVDINVGCPIDLVYKKGGGCALMNRSNKFEQIVRGMNSVLDVPLTVKIRTGVQEKVNLAHKVIPNIRKWGASMVTLHGRSREQRYTKVADWEYIAECAKLASPMPLFGNGDILSYEDANRAMQTGVSGIMIARGALIKPWIFTEVKEQRHWDISSNERFDILKDFTNYGLEHWGSDTQGVEKTRKFMLEWLSFLCRYIPAGLLEYLPQKINERPPYYMGRDYLETLMASQNVDDWIKIR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXP7</accession>
    <name>A0A452GXP7_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GXP7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008026.1">
        <property type="protein sequence ID" value="ENSGAGP00000006924.1"/>
        <property type="gene ID" value="ENSGAGG00000005577.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090730">
        <property type="term" value="C:Las1 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030687">
        <property type="term" value="C:preribosome, large subunit precursor"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004519">
        <property type="term" value="F:endonuclease activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000460">
        <property type="term" value="P:maturation of 5.8S rRNA"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000470">
        <property type="term" value="P:maturation of LSU-rRNA"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007174">
        <property type="entry name" value="Las1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15002">
        <property type="entry name" value="RIBOSOMAL BIOGENESIS PROTEIN LAS1L"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15002:SF0">
        <property type="entry name" value="RIBOSOMAL BIOGENESIS PROTEIN LAS1L"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04031">
        <property type="entry name" value="Las1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="55"/>
            <end position="86"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="345"/>
            <end position="388"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="1">
        <location>
            <begin position="55"/>
            <end position="76"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="1">
        <location>
            <begin position="369"/>
            <end position="386"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006924.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="469" mass="53749" checksum="789BB93C6CF3F342" modified="2019-05-08" version="1">MNIPIWIVDLRHDLTHGKLPQLVACRKGCDVVLEWLRRTYWSRQLGNSLARECEEEQNEEMETAATETEVDSDSEEAQNPQPPGYQKHKELHEKVRELLVSYKNQQFGVLQELQSLRKARKAWCNSSSEVEWILAQIKDLLQESREVVAEALLDDGSLIPTVEELETLNIKCEENKQLLDFKIPQTFYYFWQPLLKGLQSRNFTQTLLEKMFADLKQCSESSELRPQYLINWVAEILTTNTKAKARRKMRCPSQSQISEGKLFHHRVSLQWIKLIDECLEAPCWATPHLLQRILSDMEPPLPSVAQENLLYLSSIYTQEGGSLSSPGSSSDSSWQPIYTTESLQWKARQESQTKGHRVERRETVLGGVGEEEDNGEEEAEEEEMEAEPTPFEELLHVDNMMVIAEKRAALHGSIWQISADGVQWKDFPLGKLPGQTDDPDSLLVDSYSMMSMLDQPVTRGDGRNSPSTK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXP9</accession>
    <name>A0A452GXP9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GXP9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008030.1">
        <property type="protein sequence ID" value="ENSGAGP00000006927.1"/>
        <property type="gene ID" value="ENSGAGG00000005577.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090730">
        <property type="term" value="C:Las1 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030687">
        <property type="term" value="C:preribosome, large subunit precursor"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004519">
        <property type="term" value="F:endonuclease activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000460">
        <property type="term" value="P:maturation of 5.8S rRNA"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000470">
        <property type="term" value="P:maturation of LSU-rRNA"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007174">
        <property type="entry name" value="Las1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15002">
        <property type="entry name" value="RIBOSOMAL BIOGENESIS PROTEIN LAS1L"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15002:SF0">
        <property type="entry name" value="RIBOSOMAL BIOGENESIS PROTEIN LAS1L"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="155"/>
            <end position="199"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="1">
        <location>
            <begin position="179"/>
            <end position="196"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006927.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="337" mass="38558" checksum="D5416977B2B34AC0" modified="2019-05-08" version="1">FWQPLLKGLQSRNFTQTLLEKMFADLKQCSESSELRPQYLINWVAEILTTNTKAKARRKMRCPSQSQISEGKLFHHRVSLQWIKLIDECLEAPCWATPHLLQILSDMEPPLPSVAQENLLYLSSIYTQEGGSLSSPGSSSDSSWQPIYTTESLQWKARQESQTKGHRVERRETVLGGVGEEEDNGEEEAEEEEMEAEPTPFEELLHVDNMMVIAEKRAALHGSIWQISADGVQWKDFPLGKLPGQTDDPDSLLVDSYSMMSMLDQPVTREICFLLSCWKETQEAAILSSIVWKEGRRGRTNTHHHKVPPSDIKSMLVVLLWEPLWLSSQLLSEQQWH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXR1</accession>
    <name>A0A452GXR1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4 12">NADH dehydrogenase [ubiquinone] iron-sulfur protein 4, mitochondrial</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1 12">Accessory subunit of the mitochondrial membrane respiratory chain NADH dehydrogenase (Complex I), that is believed not to be involved in catalysis. Complex I functions in the transfer of electrons from NADH to the respiratory chain. The immediate electron acceptor for the enzyme is believed to be ubiquinone.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2 12">Mitochondrion inner membrane</location>
            <topology evidence="2 12">Peripheral membrane protein</topology>
            <orientation evidence="2 12">Matrix side</orientation>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 12">Belongs to the complex I NDUFS4 subunit family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GXR1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007921.1">
        <property type="protein sequence ID" value="ENSGAGP00000006825.1"/>
        <property type="gene ID" value="ENSGAGG00000005502.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005743">
        <property type="term" value="C:mitochondrial inner membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0022900">
        <property type="term" value="P:electron transport chain"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.190:FF:000001">
        <property type="entry name" value="NADH-ubiquinone oxidoreductase 21 kDa subunit mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.190">
        <property type="entry name" value="atu1810 like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006885">
        <property type="entry name" value="NADH_UbQ_FeS_4_mit-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038532">
        <property type="entry name" value="NDUFS4-like_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12219:SF8">
        <property type="entry name" value="NADH DEHYDROGENASE [UBIQUINONE] IRON-SULFUR PROTEIN 4, MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12219">
        <property type="entry name" value="NADH-UBIQUINONE OXIDOREDUCTASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04800">
        <property type="entry name" value="NDUS4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9 12" id="KW-0249">Electron transport</keyword>
    <keyword evidence="11 12" id="KW-0472">Membrane</keyword>
    <keyword evidence="10 12" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="7 12" id="KW-0999">Mitochondrion inner membrane</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6 12" id="KW-0679">Respiratory chain</keyword>
    <keyword evidence="8 12" id="KW-0809">Transit peptide</keyword>
    <keyword evidence="5 12" id="KW-0813">Transport</keyword>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="151"/>
            <end position="171"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00003195"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005882"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00015796"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022660"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022792"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022982"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="RuleBase" id="RU367010"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006825.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="171" mass="19240" checksum="F8CDD072A1D051E9" modified="2019-05-08" version="1">MAAALSLSAARRQLARGRGAFAACLTPARSLSTSTWRLAQDQAQDTQLITVDEKLDITPLTGVPEEHIKTRKVQIFVPARNAMQAGVNNTKKWKLEFDTRERWENPLMGWASTADPLSNVVLAFSTKEDAVAFAEKNGWSYDIQERRIPKPKSKSYGENFSWNKRTRVSTK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXR4</accession>
    <name>A0A452GXR4_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="15">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GXR4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008053.1">
        <property type="protein sequence ID" value="ENSGAGP00000006946.1"/>
        <property type="gene ID" value="ENSGAGG00000002943.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd08849">
        <property type="entry name" value="ArfGap_ASAP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd13251">
        <property type="entry name" value="PH_ASAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.220.150:FF:000002">
        <property type="entry name" value="arf-GAP with SH3 domain, ANK repeat and PH domain-containing protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.950:FF:000001">
        <property type="entry name" value="Arf-GAP with SH3 domain, ANK repeat and PH domain-containing protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000006">
        <property type="entry name" value="Arf-GAP with SH3 domain, ANK repeat and PH domain-containing protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000012">
        <property type="entry name" value="Arf-GAP with SH3 domain, ANK repeat and PH domain-containing protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.950">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.220.150">
        <property type="entry name" value="Arf GTPase activating protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.60">
        <property type="entry name" value="Arfaptin homology (AH) domain/BAR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027267">
        <property type="entry name" value="AH/BAR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037278">
        <property type="entry name" value="ARFGAP/RecO"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001164">
        <property type="entry name" value="ArfGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038508">
        <property type="entry name" value="ArfGAP_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043593">
        <property type="entry name" value="ASAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004148">
        <property type="entry name" value="BAR_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037844">
        <property type="entry name" value="PH_ASAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45854:SF4">
        <property type="entry name" value="ARF-GAP WITH SH3 DOMAIN, ANK REPEAT AND PH DOMAIN-CONTAINING PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45854">
        <property type="entry name" value="ASAP FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="ArfGap"/>
        <property type="match status" value="1"/>
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    <dbReference type="Pfam" id="PF16746">
        <property type="entry name" value="BAR_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00169">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="SH3_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00405">
        <property type="entry name" value="REVINTRACTNG"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="2"/>
    </dbReference>
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        <property type="entry name" value="ArfGap"/>
        <property type="match status" value="1"/>
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    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57863">
        <property type="entry name" value="ArfGap/RecO-like zinc finger"/>
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    </dbReference>
    <dbReference type="SUPFAM" id="SSF103657">
        <property type="entry name" value="BAR/IMD domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
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        <property type="match status" value="2"/>
    </dbReference>
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        <property type="entry name" value="ARFGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7 8" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="2 9" id="KW-0728">SH3 domain</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="10" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="PH" evidence="13">
        <location>
            <begin position="148"/>
            <end position="240"/>
        </location>
    </feature>
    <feature type="domain" description="Arf-GAP" evidence="14">
        <location>
            <begin position="264"/>
            <end position="386"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="8">
        <location>
            <begin position="427"/>
            <end position="462"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="8">
        <location>
            <begin position="463"/>
            <end position="495"/>
        </location>
    </feature>
    <feature type="domain" description="SH3" evidence="12">
        <location>
            <begin position="777"/>
            <end position="840"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="131"/>
            <end position="155"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="543"/>
            <end position="564"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="696"/>
            <end position="727"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="131"/>
            <end position="143"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="11">
        <location>
            <begin position="702"/>
            <end position="719"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50115"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006946.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="840" mass="94928" checksum="EC7F628D1E5E0EAE" modified="2019-05-08" version="1">MLKLSHTHNAAGQGNAKIRRWMKNRVFPHPPFHDLYIYLLELEGTLEVNEIKIKKGVDLLQNLIKYFHAQCNFFQDGLKAVESLKPSIETLSTDLHTIKQAQDEERKQLIHLRDILKSALQVEQKEDSQIRQSTAYSLHQPQGNKEHGTERSGSLYKKSDGIRKVWQKRKCSVKNGFLTISHGTANRPPAKLNLLTCQVKTNPEEKKCFDLISHDRTYHFQAEDEQECQIWTSVLQNSKEEALNNAFKGDENTGENNIVQELTKEIISEVQRMTGNDVCCDCGAQDPSWLSTNLGILTCIECSGIHRELGVHYSRMQSLTLDVLGTSELLLAKNIGNAGFNEIMEFCLPADETIKPNPSSDMNARKDYITAKYFERKYARKKHVDNAAKLYSLCEAVKARDIFGLVQVYADGVDLTEKIPLANGHEQDETALHLAVRSVDRTSLHIVDFLVQNSGNLDKQTYKGSTALHYCCLTDNVECLKLLLRGKASIEIANEAGETPLDIAKRLKHTHCEELLTQALSGRFNSHVHVEYEWRLLHEDLDESDDDVDEKLQQPSPNRREDRPVSFYQLGSNQLQPNVASLARDAANIAKDKQRSFMPSILQNETYGAILSGSPPPIQPAVPVTTSAPPLPPRNIGKGTNAGKYNKLVDEILRIMHSIKVQFHCTLYFAFFIDILFPAPPPPVAKTASVIEALNQQSKQPPAVPQSKPIPPPLPPQPPSRISQKKPATGWSNKIYYLTHFVSLISGLGPTRLQLHCLSFVMFHYLYIHIFVLQSKTKPKRVKAIYNCVADNPDELTFSEGDIIVVDGEEDQEWWVSIFTCAFISLVFHSKYAHTFRVHW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXS0</accession>
    <name>A0A452GXS0_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="12">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GXS0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007931.1">
        <property type="protein sequence ID" value="ENSGAGP00000006835.1"/>
        <property type="gene ID" value="ENSGAGG00000005509.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035102">
        <property type="term" value="C:PRC1 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003682">
        <property type="term" value="F:chromatin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042393">
        <property type="term" value="F:histone binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045892">
        <property type="term" value="P:negative regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd09577">
        <property type="entry name" value="SAM_Ph1_2_3"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Polyhomeotic-like protein 2 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.60.160:FF:000002">
        <property type="entry name" value="Polyhomeotic-like protein 2 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.60.160">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.50">
        <property type="entry name" value="Transcription Factor, Ets-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050548">
        <property type="entry name" value="PcG_chromatin_remod_factors"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001660">
        <property type="entry name" value="SAM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013761">
        <property type="entry name" value="SAM/pointed_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012313">
        <property type="entry name" value="Znf_FCS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038603">
        <property type="entry name" value="Znf_FCS_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12247">
        <property type="entry name" value="POLYCOMB GROUP PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12247:SF86">
        <property type="entry name" value="POLYHOMEOTIC-LIKE PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16616">
        <property type="entry name" value="PHC2_SAM_assoc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00536">
        <property type="entry name" value="SAM_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21319">
        <property type="entry name" value="zf-FCS_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00454">
        <property type="entry name" value="SAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47769">
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51024">
        <property type="entry name" value="ZF_FCS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="6" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <keyword evidence="4 8" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="FCS-type" evidence="11">
        <location>
            <begin position="98"/>
            <end position="132"/>
        </location>
    </feature>
    <feature type="domain" description="SAM" evidence="10">
        <location>
            <begin position="264"/>
            <end position="328"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="1"/>
            <end position="26"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="75"/>
            <end position="99"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="149"/>
            <end position="188"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="203"/>
            <end position="241"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="1"/>
            <end position="11"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="203"/>
            <end position="213"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="214"/>
            <end position="229"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00367"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50105"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS51024"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006835.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="328" mass="36067" checksum="2ED15CE255A01BCA" modified="2019-05-08" version="1">MTSGNGNSASTVAGAAPQNGENKPPQAIVKPQVLTHVIEGFVIQEGAEPFPVGRSSLLVGNLKKKYAQGLLAEKLPQQDNTTTTDSEMEEPYLQESKEEGNPPKLKCELCGRVDFAYKFKRSKRFCSMACAKRYNVGCTKRVGLFHPDRSKLQKPGAATHGRRRTCKGALPVLSKDTKKQTPVSLSTGSVPLSVTASLQLSRSQEDSSCCSDNSSYEEPLSPLSASSSTSRRRQGERDLELPDMHMRDLVGIGHHFLPSEPTKWNVEDVYEFIRSLPGCQEIAEEFRAQEIDGQALLLLKEDHLMSAMNIKLGPALKIYARISMLKDS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXS5</accession>
    <name>A0A452GXS5_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="7">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GXS5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007937.1">
        <property type="protein sequence ID" value="ENSGAGP00000006840.1"/>
        <property type="gene ID" value="ENSGAGG00000005511.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:1990904">
        <property type="term" value="C:ribonucleoprotein complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003729">
        <property type="term" value="F:mRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006396">
        <property type="term" value="P:RNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd08033">
        <property type="entry name" value="LARP_6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.10:FF:000158">
        <property type="entry name" value="La ribonucleoprotein domain family member 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.330">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.10">
        <property type="entry name" value="Winged helix-like DNA-binding domain superfamily/Winged helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045180">
        <property type="entry name" value="La_dom_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006630">
        <property type="entry name" value="La_HTH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002344">
        <property type="entry name" value="Lupus_La"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012677">
        <property type="entry name" value="Nucleotide-bd_a/b_plait_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035979">
        <property type="entry name" value="RBD_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024642">
        <property type="entry name" value="SUZ-C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036388">
        <property type="entry name" value="WH-like_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036390">
        <property type="entry name" value="WH_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22792:SF61">
        <property type="entry name" value="LA RIBONUCLEOPROTEIN DOMAIN FAMILY MEMBER 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22792">
        <property type="entry name" value="LUPUS LA PROTEIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05383">
        <property type="entry name" value="La"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00302">
        <property type="entry name" value="LUPUSLA"/>
    </dbReference>
    <dbReference type="SMART" id="SM00715">
        <property type="entry name" value="LA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54928">
        <property type="entry name" value="RNA-binding domain, RBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46785">
        <property type="entry name" value="Winged helix' DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50961">
        <property type="entry name" value="HTH_LA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51938">
        <property type="entry name" value="SUZ_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0539">Nucleus</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 4" id="KW-0694">RNA-binding</keyword>
    <feature type="domain" description="HTH La-type RNA-binding" evidence="5">
        <location>
            <begin position="30"/>
            <end position="121"/>
        </location>
    </feature>
    <feature type="domain" description="SUZ-C" evidence="6">
        <location>
            <begin position="365"/>
            <end position="425"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00332"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50961"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS51938"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006840.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="434" mass="48069" checksum="78DCD6408618CD52" modified="2019-05-08" version="1">MQRQPPSSAVWSFYDLNDIFGADSFECDSSVPDAQLIQRIVSQVEFYLSDENLAKDAFLLKHVQKNKLGFVSIKLLTSFKKVKYLTRDWRLTLYALHFSELLEVNEEGTKVRRKIPIPESLLSIPPSKLLLAWDLLPQEQRLSPPLQKSFIETITKMFSPFGAIASIRILRPGKKLPSDVKKYSSRYPELLTKCCALVEYENLESARKAYEELNRGQGSPSRESIKVVRLSGKGSKKKSGDDVEEVELVDQLGRKQQASETLAYAAGDSFFCSSSESDSILASPPIPIQKYLSTPVWSACSLGVSFKPGFFSSPQVSPLLSNKSLAQLHCPSPLAAELGNGGSSSPDTSPEFRRLSDSCWDSGIGSGSSWVQRRRVAAHSQSLETRVLPCSSLALRKMPDSFGLRSGVMRLPHGPDGTKGFYNSIGRGKLVLRH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GY10</accession>
    <name>A0A452GY10_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">Calpain catalytic domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Calcium-regulated non-lysosomal thiol-protease which catalyze limited proteolysis of substrates involved in cytoskeletal remodeling and signal transduction.</text>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the peptidase C2 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GY10"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008106.1">
        <property type="protein sequence ID" value="ENSGAGP00000006996.1"/>
        <property type="gene ID" value="ENSGAGG00000004685.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004198">
        <property type="term" value="F:calcium-dependent cysteine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00214">
        <property type="entry name" value="Calpain_III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00044">
        <property type="entry name" value="CysPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.380:FF:000001">
        <property type="entry name" value="Calpain-1 catalytic subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.70.10:FF:000001">
        <property type="entry name" value="Calpain-1 catalytic subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.380">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.70.10">
        <property type="entry name" value="Cysteine proteinases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.238.10">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033883">
        <property type="entry name" value="C2_III"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022684">
        <property type="entry name" value="Calpain_cysteine_protease"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022682">
        <property type="entry name" value="Calpain_domain_III"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022683">
        <property type="entry name" value="Calpain_III"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036213">
        <property type="entry name" value="Calpain_III_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011992">
        <property type="entry name" value="EF-hand-dom_pair"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038765">
        <property type="entry name" value="Papain-like_cys_pep_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000169">
        <property type="entry name" value="Pept_cys_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001300">
        <property type="entry name" value="Peptidase_C2_calpain_cat"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10183">
        <property type="entry name" value="CALPAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10183:SF268">
        <property type="entry name" value="CALPAIN-2 CATALYTIC SUBUNIT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01067">
        <property type="entry name" value="Calpain_III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00648">
        <property type="entry name" value="Peptidase_C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00704">
        <property type="entry name" value="CALPAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00720">
        <property type="entry name" value="calpain_III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00230">
        <property type="entry name" value="CysPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49758">
        <property type="entry name" value="Calpain large subunit, middle domain (domain III)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54001">
        <property type="entry name" value="Cysteine proteinases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47473">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50203">
        <property type="entry name" value="CALPAIN_CAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00139">
        <property type="entry name" value="THIOL_PROTEASE_CYS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12" id="KW-0106">Calcium</keyword>
    <keyword evidence="5" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="10 16" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="13" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7 16" id="KW-0645">Protease</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0677">Repeat</keyword>
    <keyword evidence="11 16" id="KW-0788">Thiol protease</keyword>
    <feature type="domain" description="Calpain catalytic" evidence="17">
        <location>
            <begin position="45"/>
            <end position="344"/>
        </location>
    </feature>
    <feature type="active site" evidence="15 16">
        <location>
            <position position="105"/>
        </location>
    </feature>
    <feature type="active site" evidence="15 16">
        <location>
            <position position="262"/>
        </location>
    </feature>
    <feature type="active site" evidence="15 16">
        <location>
            <position position="286"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001913"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00007623"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022807"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00054695"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PIRSR" id="PIRSR622684-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006996.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="584" mass="66332" checksum="1618EF8534EE1130" modified="2019-05-08" version="1">MSGIAVKLAKERETARGLGSNAKAFKYLSQDFETLRNECLQAGKLFQDPSFPAIASSLGYKELGPNSYKTQGITWKRPTELCSNPQFIVGGATRTDICQGALGDCWLLAAIASLTLNEEILARVVPKDQSFQDKYAGIFHFQFWQYGEWVDVVVDDKLPTKNGELLFVHSAEGSEFWSALLEKAYAKVNGSYEALSGGSTTEGFEDFTGGIAEWYELQKAPPNLFKIIQKALLKGSLLGCSIDITSAAETEAVTSQKLVKGHAYSVTGAEEVNCRGSMQKLIRIRNPWGEVEWTGKWNDNCSNWNSVAPEVRERLTRRHEDGEFWMSFSDFLRHYSRLEICNLTPDTLANEKYKKWSLTKLDGNWRRGSTAGGCRNYPNTFWMNPQYLIKLEEEDEDPDDPEKGCTFLVGLIQKHRRRQRKMGEDMHTIGFGIYEASRFSGQTNVHLSKNFFLTNRAREKSNTFINLREVLNRFKLPPGEYVIVPSTFEPNKDGDFCLRVFSEKNADSQYVLAKCFHLLFCPVGFKLTCQLHQVIVARFADEQLIIDFDNFVRCLIRLETLFKIFKKLDVEKSGTVEMNLNTVS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GY11</accession>
    <name>A0A452GY11_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12 13">RNA cytidine acetyltransferase</fullName>
            <ecNumber evidence="13">2.3.1.-</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="13">18S rRNA cytosine acetyltransferase</fullName>
        </alternativeName>
    </protein>
    <gene>
        <name evidence="13" type="primary">NAT10</name>
    </gene>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">RNA cytidine acetyltransferase with specificity toward both 18S rRNA and tRNAs. Catalyzes the formation of N(4)-acetylcytidine (ac4C) in 18S rRNA. Required for early nucleolar cleavages of precursor rRNA at sites A0, A1 and A2 during 18S rRNA synthesis. Catalyzes the formation of ac4C in serine and leucine tRNAs. Requires the tRNA-binding adapter protein THUMPD1 for full tRNA acetyltransferase activity but not for 18S rRNA acetylation.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>a cytidine in 18S rRNA + acetyl-CoA + ATP + H2O = an N(4)-acetylcytidine in 18S rRNA + ADP + phosphate + CoA + H(+)</text>
            <dbReference type="Rhea" id="RHEA:51424"/>
            <dbReference type="Rhea" id="RHEA-COMP:13575"/>
            <dbReference type="Rhea" id="RHEA-COMP:13576"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57288"/>
            <dbReference type="ChEBI" id="CHEBI:74900"/>
            <dbReference type="ChEBI" id="CHEBI:82748"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>a cytidine in tRNA + acetyl-CoA + ATP + H2O = an N(4)-acetylcytidine in tRNA + ADP + phosphate + CoA + H(+)</text>
            <dbReference type="Rhea" id="RHEA:53876"/>
            <dbReference type="Rhea" id="RHEA-COMP:13670"/>
            <dbReference type="Rhea" id="RHEA-COMP:13671"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57288"/>
            <dbReference type="ChEBI" id="CHEBI:74900"/>
            <dbReference type="ChEBI" id="CHEBI:82748"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
    </comment>
    <comment type="subunit">
        <text evidence="13">Interacts with THUMPD1.</text>
    </comment>
    <comment type="subunit">
        <text evidence="11">Part of the small subunit (SSU) processome, composed of more than 70 proteins and the RNA chaperone small nucleolar RNA (snoRNA) U3. Interacts with THUMPD1. Interacts with SUN1 (via N-terminus). Interacts with TERT.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 13">Nucleus</location>
            <location evidence="1 13">Nucleolus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="13">Belongs to the RNA cytidine acetyltransferase family. NAT10 subfamily.</text>
    </comment>
    <comment type="caution">
        <text evidence="13">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="EC" id="2.3.1.-" evidence="13"/>
    <dbReference type="AlphaFoldDB" id="A0A452GY11"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008174.1">
        <property type="protein sequence ID" value="ENSGAGP00000007057.1"/>
        <property type="gene ID" value="ENSGAGG00000002160.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030686">
        <property type="term" value="C:90S preribosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005730">
        <property type="term" value="C:nucleolus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:1990883">
        <property type="term" value="F:18S rRNA cytidine N-acetyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000049">
        <property type="term" value="F:tRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051392">
        <property type="term" value="F:tRNA cytidine N4-acetyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="RHEA"/>
    </dbReference>
    <dbReference type="GO" id="GO:1904812">
        <property type="term" value="P:rRNA acetylation involved in maturation of SSU-rRNA"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051391">
        <property type="term" value="P:tRNA acetylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="CDD" id="cd04301">
        <property type="entry name" value="NAT_SF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.11040:FF:000006">
        <property type="entry name" value="RNA cytidine acetyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.630.30:FF:000019">
        <property type="entry name" value="RNA cytidine acetyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:002218">
        <property type="entry name" value="tRNA(Met) cytidine acetyltransferase TmcA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.11040">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.630.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="HAMAP" id="MF_03211">
        <property type="entry name" value="RNA_acetyltr_Nat10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016181">
        <property type="entry name" value="Acyl_CoA_acyltransferase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000182">
        <property type="entry name" value="GNAT_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033688">
        <property type="entry name" value="NAT10"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007807">
        <property type="entry name" value="TcmA/NAT10_helicase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032672">
        <property type="entry name" value="TmcA/NAT10/Kre33"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013562">
        <property type="entry name" value="TmcA/NAT10_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027992">
        <property type="entry name" value="tRNA_bind_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10925">
        <property type="entry name" value="N-ACETYLTRANSFERASE 10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10925:SF5">
        <property type="entry name" value="RNA CYTIDINE ACETYLTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13718">
        <property type="entry name" value="GNAT_acetyltr_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05127">
        <property type="entry name" value="NAT10_TcmA_helicase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08351">
        <property type="entry name" value="TmcA_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13725">
        <property type="entry name" value="tRNA_bind_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55729">
        <property type="entry name" value="Acyl-CoA N-acyltransferases (Nat)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51186">
        <property type="entry name" value="GNAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0007">Acetylation</keyword>
    <keyword evidence="10 13" id="KW-0012">Acyltransferase</keyword>
    <keyword evidence="7 13" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="6 13" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="9 13" id="KW-0539">Nucleus</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 13" id="KW-0698">rRNA processing</keyword>
    <keyword evidence="4 13" id="KW-0808">Transferase</keyword>
    <keyword evidence="5 13" id="KW-0819">tRNA processing</keyword>
    <feature type="domain" description="N-acetyltransferase" evidence="14">
        <location>
            <begin position="553"/>
            <end position="732"/>
        </location>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <begin position="282"/>
            <end position="291"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <position position="465"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <begin position="624"/>
            <end position="626"/>
        </location>
        <ligand>
            <name>acetyl-CoA</name>
            <dbReference type="ChEBI" id="CHEBI:57288"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <begin position="631"/>
            <end position="637"/>
        </location>
        <ligand>
            <name>acetyl-CoA</name>
            <dbReference type="ChEBI" id="CHEBI:57288"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004604"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022552"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022694"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023315"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00065380"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00068357"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="HAMAP-Rule" id="MF_03211"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS51186"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007057.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1009" mass="113758" checksum="6EC23CD254736B23" modified="2019-05-08" version="1">MQRKKVDNRIRVLIENGMAERHRTLFVLVGDHGKDQVVILHHMLSKAAVKARPSVLWCYKKELGFSSHRKKRMRQLQKKIKSGTLNLNQDDPFELFVAATNIRYCYYKETHKILGNTFGMCVLQDFEALTPNLLARTVETVEGGGIVVILLRTMNSLKQLYTITMDVHSRYRTEAHQDIVGRFNERFILSLAACKNCIVIDDQLNILPISSHIANITPVLPQSLVSDLELKELKESLQDTQPVGVLVDSCKTLDQAKALLKFIEAISEKTLRSTVALTAARGRGKSAALGLAIAGAVAFGYSNIFVTSPSPDNLHTLFEFVFKGFDALQYEEHLDYEIIQSLNPEFNKAVVRVNVFKEHRQTIQYIHPADAVKLGQAELVVIDEAAAIPLPLVKNLLGPYLVFMASTINGYEGTGRSLSLKLIQQLRQQSAQAQASITAENKSTATAKLTSARTLHEVSLHESIRYAPGDPVEKWLNDLLCLDCLNITRIISGCPLPETCNLYYVNRDTLFCYHKASEIFLQRLMALYVASHYKNSPNDLQMLSDAPAHHLFCLLPPVPPTQNSLPEVLAVVQVCLEGEISRQSIMNSLSRGKKASGDLIPWTVSEQFQDPDFGSLSGGRVVRIAVHPDYQGMGYGSRALTLLQMYYEGKFPCLEEEVTPKPKGIATVSSEAVSLLEEVVTPRKDLPPLLLKLSERQAENLDYLGRFSNYGPQTPRNDLTGEHSCIMLKMLNEEESEQEHWLAAFWKDFRRRFLSLLSYQFSTFPPSLALNILQNKNIKQELQPPISRPELEAMFLPYDLKRLEMYSQNMVDYHLIMDTIPAVSRMYFLSQLGDTSLSAAQSALLLGIGLQHKSLDVLEKEIELPSSQLMGLFNRIIRKVVQLFSRIQEKAVEEQMAVTKEVVMEPTLKSLNEDLEEAAKEFQEKQKQEMGKLKDMDFTQYIIRGDDEEWNEVLNKAGQNTSIVSLKRLGKCVCVCVCVLFCFVLFVFLKIRAHLFSGSVLPSLLYCNV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GY12</accession>
    <name>A0A452GY12_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Histone-binding protein RBBP4-like N-terminal domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the WD repeat RBAP46/RBAP48/MSI1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GY12"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008116.1">
        <property type="protein sequence ID" value="ENSGAGP00000007006.1"/>
        <property type="gene ID" value="ENSGAGG00000005642.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006325">
        <property type="term" value="P:chromatin organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006260">
        <property type="term" value="P:DNA replication"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022052">
        <property type="entry name" value="Histone-bd_RBBP4-like_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020472">
        <property type="entry name" value="WD40_PAC1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019775">
        <property type="entry name" value="WD40_repeat_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036322">
        <property type="entry name" value="WD40_repeat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001680">
        <property type="entry name" value="WD40_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050459">
        <property type="entry name" value="WD_repeat_RBAP46/RBAP48/MSI1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22850">
        <property type="entry name" value="WD40 REPEAT FAMILY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12265">
        <property type="entry name" value="CAF1C_H4-bd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00400">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00320">
        <property type="entry name" value="GPROTEINBRPT"/>
    </dbReference>
    <dbReference type="SMART" id="SM00320">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50978">
        <property type="entry name" value="WD40 repeat-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00678">
        <property type="entry name" value="WD_REPEATS_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50082">
        <property type="entry name" value="WD_REPEATS_2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50294">
        <property type="entry name" value="WD_REPEATS_REGION"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0156">Chromatin regulator</keyword>
    <keyword evidence="4" id="KW-0235">DNA replication</keyword>
    <keyword evidence="7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="3 8" id="KW-0853">WD repeat</keyword>
    <feature type="domain" description="Histone-binding protein RBBP4-like N-terminal" evidence="9">
        <location>
            <begin position="18"/>
            <end position="87"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="8">
        <location>
            <begin position="172"/>
            <end position="214"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="8">
        <location>
            <begin position="222"/>
            <end position="264"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="8">
        <location>
            <begin position="343"/>
            <end position="377"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009341"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022574"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022705"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF12265"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007006.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="399" mass="44653" checksum="52C608DF63EBF4C5" modified="2019-05-08" version="1">MANKEMFEDTVEERVINEEYKIWKKNTPFLYDLVMTHALEWPSLTVQWLPDVTRPEGKDYALHWLVLGTHTSDEQNHLVVARVQIPNDDAQFDASHYDSEKGEFGGFGSVTGKIETEIKINHEGEVNRARYMPQNPCIIATKTPSSDVLVFDYTKHPSKPDPSGECNPDLRLRGHQKEGYGLSWNSNLSGHLLSASDDHTVCLWDISAGPKEGKVVDAKAIFTGHSAVVEDVAWHLLHESLFGSVADDQKLMIWDTRSNTTSKPSHSVDAHTAEVNCLSFNPYSEFILATGSADKVHWSPHNETILASSGTDRRLNVWDLSKIGEEQSAEDAEDGPPELLFIHGGHTAKISDFSWNPNEPWVICSVSEDNIMQIWQMAENIYNDEEPDIAASELEGQGT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GY31</accession>
    <name>A0A452GY31_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="5">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GY31"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008146.1">
        <property type="protein sequence ID" value="ENSGAGP00000007030.1"/>
        <property type="gene ID" value="ENSGAGG00000005653.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd13242">
        <property type="entry name" value="PH_puratrophin-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00160">
        <property type="entry name" value="RhoGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.60">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.900.10">
        <property type="entry name" value="Dbl homology (DH) domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035899">
        <property type="entry name" value="DBL_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000219">
        <property type="entry name" value="DH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052231">
        <property type="entry name" value="Rho_GEF_signaling-related"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055251">
        <property type="entry name" value="SOS1_NGEF_PH"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45845:SF1">
        <property type="entry name" value="PLECKSTRIN HOMOLOGY AND RHOGEF DOMAIN CONTAINING G4B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45845">
        <property type="entry name" value="RHO GUANINE NUCLEOTIDE EXCHANGE FACTOR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00621">
        <property type="entry name" value="RhoGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22697">
        <property type="entry name" value="SOS1_NGEF_PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00233">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00325">
        <property type="entry name" value="RhoGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48065">
        <property type="entry name" value="DBL homology domain (DH-domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46966">
        <property type="entry name" value="Spectrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50010">
        <property type="entry name" value="DH_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="2">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019216654" evidence="2">
        <location>
            <begin position="24"/>
            <end position="1192"/>
        </location>
    </feature>
    <feature type="domain" description="DH" evidence="4">
        <location>
            <begin position="686"/>
            <end position="863"/>
        </location>
    </feature>
    <feature type="domain" description="PH" evidence="3">
        <location>
            <begin position="875"/>
            <end position="983"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="485"/>
            <end position="524"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="563"/>
            <end position="601"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="662"/>
            <end position="687"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1043"/>
            <end position="1067"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1109"/>
            <end position="1192"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="1">
        <location>
            <begin position="493"/>
            <end position="504"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="505"/>
            <end position="515"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="575"/>
            <end position="593"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="672"/>
            <end position="681"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="1">
        <location>
            <begin position="1053"/>
            <end position="1067"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="1112"/>
            <end position="1121"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="1">
        <location>
            <begin position="1122"/>
            <end position="1141"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="1161"/>
            <end position="1172"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="1182"/>
            <end position="1192"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50010"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007030.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1192" mass="137226" checksum="F719516948FA85B7" modified="2019-05-08" version="1">MLSQRLFFLCFIWLGSRDRGGRAVVQVCTRSTIWSSEHSTTTELARLLMYFYRIPRKEVRDLGLVILVDARKCQPSPGLFKAFTVLQSAVPHSIHSVLILTEKESAFRPDKDTAIQYEVLTSLKALQKHIDCTQLTAEFDGTFLYNHHDWMRFRKKLEPFITNCKEAIVFLQNSVCSLNSNRTLSTVQEVRDLIDKHKTMMKLVLEDALLVTLRLEGGTILARLRKEEFCNTEDYRDAMEAITRLYNQVDEEVHRLVLISNKCLQQLENLQEIRKFEEHCDQIKLWFENERERYLGPLDQELLSLESVKKQQEEFRVFSDKVTQYCEKGLQLIQKNCSLNLEEKVQTFKSYLSNVSTQTEERKSELEKLIHLCEFYETAHRWMLHCNEYLEHLTVENKYSQSVIQSLQSYQQEATKVSAENFQKVNEIVLGLASKKELKQWNILWLKCQQTRHHLEEVLSEALKGSNKETSRKVPQEADVKVNCEFDTEGARRPSQSLSRLSSSTDNDIWDNNTKPLHKPQHMSLPSGVPPFLDHENSNQNMDVYVQLNASPPQTFTPLLGGRQRASPISEDMDSVSTSGSASCYSEPVQSPTTRHRKHPLKKIMKKTQSFEFTQLESSPSELHRHGYTGVYIKGLEVASNVAAEKKYMQRLNIKSPLISRNRSLSSPSRIHHTEEEEKKRAGSSKVQHIMDEMITTEREYVRSLGYIIDNYFPEMERIDLPQDLRGKRNIIFGNLEKLYDFHCQYFLKELEHCRDFPLRVSHCFLRHEEQFGMYALYSKNKPQSDALLTSHGNAFFKNKQQDLGDKMDLASYLLKPIQRMSKYALLLKDMIKECTKTQEQELNDLRTAEEMVKFQLRHGNDLLAMDAIRGCDVNLKEQGQLRCQDEFIVCCGRKKYLRHVFLFEDLILFSKTKKIDGGYDIYMYKQSFKTAEIGMTENVGDSGLRFEIWFRRRRKSQDTYILQANSAEVKIAWTSIIGRILWRQALRNRELRMQEMVSMGIGNKPFMDIKPSEAAISDRAIDYIMKGTESRSRASIAVSSFDHSTPFKRPHSTISNSSTSSSSSQSSSSILGSLNLHVYSSPPHPGLLGPPFYHWSYDIRTCIEEDELEQETGSQPSMITESSESSQCTSGESVSGFSSSAHPGLSQLSTETFLDEDASNPGSKPPSQCTSPALPEKNPTSKHTNQYVTAL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GY34</accession>
    <name>A0A452GY34_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">SMC hinge domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Chromosome</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000008202.1">
        <property type="protein sequence ID" value="ENSGAGP00000007083.1"/>
        <property type="gene ID" value="ENSGAGG00000005287.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005694">
        <property type="term" value="C:chromosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051276">
        <property type="term" value="P:chromosome organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006302">
        <property type="term" value="P:double-strand break repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1060.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.1620">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058611">
        <property type="entry name" value="Ig_SMCHD1_1st"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058612">
        <property type="entry name" value="Ig_SMCHD1_2nd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058613">
        <property type="entry name" value="Ig_SMCHD1_4th"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058614">
        <property type="entry name" value="Ig_SMCHD1_5th"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058615">
        <property type="entry name" value="Ig_SMCHD1_6th"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058617">
        <property type="entry name" value="Ig_SMCHD1_7th"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058616">
        <property type="entry name" value="Ig_SMCHD1_8th"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010935">
        <property type="entry name" value="SMC_hinge"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036277">
        <property type="entry name" value="SMC_hinge_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038892">
        <property type="entry name" value="SMCHD1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055109">
        <property type="entry name" value="SMCHD1_S5"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22640">
        <property type="entry name" value="STRUCTURAL MAINTENANCE OF CHROMOSOMES FLEXIBLE HINGE DOMAIN-CONTAINING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22640:SF2">
        <property type="entry name" value="STRUCTURAL MAINTENANCE OF CHROMOSOMES FLEXIBLE HINGE DOMAIN-CONTAINING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26194">
        <property type="entry name" value="Ig_SMCHD1_1st"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26195">
        <property type="entry name" value="Ig_SMCHD1_2nd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26196">
        <property type="entry name" value="Ig_SMCHD1_4th"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26197">
        <property type="entry name" value="Ig_SMCHD1_5th"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26198">
        <property type="entry name" value="Ig_SMCHD1_6th"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26201">
        <property type="entry name" value="Ig_SMCHD1_7th"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26199">
        <property type="entry name" value="Ig_SMCHD1_8th"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06470">
        <property type="entry name" value="SMC_hinge"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22899">
        <property type="entry name" value="SMCHD1_S5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00968">
        <property type="entry name" value="SMC_hinge"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF75553">
        <property type="entry name" value="Smc hinge domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0158">Chromosome</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="SMC hinge" evidence="3">
        <location>
            <begin position="1472"/>
            <end position="1599"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022454"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="SMART" id="SM00968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007083.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1723" mass="193922" checksum="47B8F399292D3AD1" modified="2019-05-08" version="1">MAKLLWISQIFMGAGPDSVHVYAASLPCYSEAGAGTLYSCNMLWAGCLPLCSYLPPAKGVQGEQIKTFVYSVAKQQGRASAVHHPQLLSGLLGHKATVNCGRRGIVCTCCVFTNILYVQCYNWDVPQPYSTQAQPTGTAGQEASRLGEAHCTAIALVFPVCVFGLVKNIFLMNFRHIYHYYIHGPKGNETNATVKEAGPFNNIDIEILMFEKGKVPRMVNLREIKDDMQTLYINSAADSFEFKSHVEGEGIVEGIIRYHPFLYDKETYPDDPCFPSRLKDDDDDDDCFVVEKGARGKRPIFECFWNGRLIPYTSVEDFDWCAPPKKRGLAPIECYNRISGTLFTNDKFQVSTNKLTFMDLELKLKDKNTLFTRVFNGQEQRMKIDREFALWLKDCHEKYDKQIKFTLFKEISTRTDLASKRMQSPWATYSAIEWDGKTYRTGQLVKTIKTLPIFYGSILKFFLYGDHDGDVYATGGEVQLALEPQALYDEIKIIPISKLDRAMSEKVVKKYIEDEMARLPDRLSVTWPEGDELLPNEIKLAGTPIGALRIEILNKKGEAMQKLPGTSHGGSKKLLVELKVILHSPVGNKEIISHISQHGGKWPYWFKKMENIHKLGSYTLKLQVVLNESNADTYAGRPLPSKMFKFNIVEGKPEKFSVSLLDPPFRVGVPFNIPLDLQDEFGHPTQLASDIEPILEASGLTLQYEEINRGTNCVIKGVTAKGHVNNYLGKVGPPRQLKVKPDSEMLKIENGTSFPFQVEVLDEAGNITAQPKLIVHCKFVGAPNLPVYAVDCSNAGTNILTGPVVHVQNIKKDQMLKARIEIPSCKDVPPVEKTIKLLPSSHVARLQIFSVEGQKAIQIKHQDEINWIAGDVMHNLIFQMYDEGEREIIITADLAEKIKVNWTPRINRDQMIQGLLPDVKVPTSVKDVRYCLITFHDDHVSLESAFTVRPLPDEPKHIKCKLKGPNTLQMGEELQSEIDVMVTDQYGNQIQTLSASCINSLGVSGNGLDKSNLKTTWQESTQTMAVKGIRFYPGPPGSKELCFAWREFSDFLRVNLTSGPPVKLHLMDWPDLEEPIAVINGRELQKPLIVQLYDQWENPSPEPNVKISLVKASNLKIIPSSQQHKTDENGRANLGIISIHAPRGEHTLQLKASYNRNTLDCPIIKFNVLPDPEKPVCLNVKYDKNASFAAGSTFPDFMVSVISEDSNIIKNINPARVSMKIWEGLSSGNRIPANVSNWKIIVGQPTYSNDPVQLLPESLPATPVVSNVRTVASRTLIKDLCLHIKDEYNNRTGYDLVGKIIAAIKSSSEEDTEIPVFQGKVKTIEFPFNRGSAEIVVSNLVLAENSPGRDSTEYILEFEADLPTLAKPLEPYRLSFMFYNDFKKQQRMATLTRERDELSKSIAVYRNMFDTTEQLITEMKCKSYDVQLLTFSFNLQLQCVDSLIKQKMLEQEGVKKSPRRACTLSNYPKDNQDILGKIAHLAQIEDDRAAKVISWHMASDMDCVVTLTTAAARRIFDETQGRQQVLPLDSVYKKTLPDWNRTLPHLRNGRNHFRPIGNPVFARDLLTFPENVEHCQIVFGMLLGDTIIIDNLDAANHYRKEVVKITHCPTLLTREGDRIRSNGKFGGLQNKAPPMDKLRGMVFGAPMPKLYSGLVCNPWPSKHLFSCCNSPLSQETHPNSTTTFSTKDRFHRVFSACSNRHAVTSLHAVINLPTDRFLTLKQE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GY35</accession>
    <name>A0A452GY35_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="9">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 5">Belongs to the D-isomer specific 2-hydroxyacid dehydrogenase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GY35"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008151.1">
        <property type="protein sequence ID" value="ENSGAGP00000007035.1"/>
        <property type="gene ID" value="ENSGAGG00000005657.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140297">
        <property type="term" value="F:DNA-binding transcription factor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051287">
        <property type="term" value="F:NAD binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016616">
        <property type="term" value="F:oxidoreductase activity, acting on the CH-OH group of donors, NAD or NADP as acceptor"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003713">
        <property type="term" value="F:transcription coactivator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001221">
        <property type="term" value="F:transcription coregulator binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003714">
        <property type="term" value="F:transcription corepressor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05299">
        <property type="entry name" value="CtBP_dh"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.720:FF:000012">
        <property type="entry name" value="C-terminal-binding protein 2 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.720">
        <property type="entry name" value="NAD(P)-binding Rossmann-like Domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043322">
        <property type="entry name" value="CtBP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051638">
        <property type="entry name" value="CTBP_dehydrogenase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006139">
        <property type="entry name" value="D-isomer_2_OHA_DH_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029753">
        <property type="entry name" value="D-isomer_DH_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029752">
        <property type="entry name" value="D-isomer_DH_CS1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006140">
        <property type="entry name" value="D-isomer_DH_NAD-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036291">
        <property type="entry name" value="NAD(P)-bd_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46029">
        <property type="entry name" value="C-TERMINAL-BINDING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46029:SF2">
        <property type="entry name" value="C-TERMINAL-BINDING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00389">
        <property type="entry name" value="2-Hacid_dh"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02826">
        <property type="entry name" value="2-Hacid_dh_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52283">
        <property type="entry name" value="Formate/glycerate dehydrogenase catalytic domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51735">
        <property type="entry name" value="NAD(P)-binding Rossmann-fold domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00065">
        <property type="entry name" value="D_2_HYDROXYACID_DH_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00671">
        <property type="entry name" value="D_2_HYDROXYACID_DH_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0539">Nucleus</keyword>
    <keyword evidence="3 5" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="D-isomer specific 2-hydroxyacid dehydrogenase catalytic" evidence="7">
        <location>
            <begin position="27"/>
            <end position="341"/>
        </location>
    </feature>
    <feature type="domain" description="D-isomer specific 2-hydroxyacid dehydrogenase NAD-binding" evidence="8">
        <location>
            <begin position="124"/>
            <end position="306"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="398"/>
            <end position="430"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="419"/>
            <end position="430"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005854"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="RuleBase" id="RU003719"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF00389"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF02826"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007035.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="430" mass="46691" checksum="36FFB43EA10E5E71" modified="2019-05-08" version="1">MSGIRPPIMNGPMHPRPLVALLDGRDCTVEMPILKDVATVAFCDAQSTQEIHEKVLNEAVGALMYHTITLTREDLEKFKALRIIVRIGSGFDNIDIKSAGDLGIAVCNVPAASVEETADSTMCHILNLYRRTTWLHQALREGTRVQSVEQIREVASGAARIRGETLGIIGLGRVGQAVALRAKAFGFSVLFYDPYLSDGIERALGLQRVSTLQDLLFHSDCVTLHCNLNEHNHHLINDFTIKQMRQGAFLVNTARGGLVDEKALAQALKEGRIRGAALDVHESEPFSFSQGPLKDAPNLICTPHAAWYSEQASIEMREEAAREIRRAITGRIPDSLKNCVNKDHLTAATPWASMDPGVVHPELNGAAYSRYPPGVVSMAPAGIPTAVEGIVPNPMSLSHGLPAVAHPPHAPSPGQTVKPEADRDHPSDQL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GY41</accession>
    <name>A0A452GY41_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">acetyl-CoA carboxylase</fullName>
            <ecNumber evidence="3">6.4.1.2</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>biotin</name>
            <dbReference type="ChEBI" id="CHEBI:57586"/>
        </cofactor>
    </comment>
    <comment type="pathway">
        <text evidence="2">Lipid metabolism; malonyl-CoA biosynthesis; malonyl-CoA from acetyl-CoA: step 1/1.</text>
    </comment>
    <dbReference type="EC" id="6.4.1.2" evidence="3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008148.1">
        <property type="protein sequence ID" value="ENSGAGP00000007032.1"/>
        <property type="gene ID" value="ENSGAGG00000004854.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00655">
        <property type="reaction ID" value="UER00711"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003989">
        <property type="term" value="F:acetyl-CoA carboxylase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006633">
        <property type="term" value="P:fatty acid biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:2001295">
        <property type="term" value="P:malonyl-CoA biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="CDD" id="cd06850">
        <property type="entry name" value="biotinyl_domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.460.10:FF:000001">
        <property type="entry name" value="Acetyl-CoA carboxylase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.50.100:FF:000005">
        <property type="entry name" value="Acetyl-CoA carboxylase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.470.20:FF:000005">
        <property type="entry name" value="Acetyl-CoA carboxylase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1770.10:FF:000001">
        <property type="entry name" value="acetyl-CoA carboxylase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1490.20:FF:000003">
        <property type="entry name" value="acetyl-CoA carboxylase isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.20:FF:000005">
        <property type="entry name" value="acetyl-CoA carboxylase isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.226.10:FF:000010">
        <property type="entry name" value="acetyl-CoA carboxylase isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.50.100">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.226.10">
        <property type="entry name" value="2-enoyl-CoA Hydratase, Chain A, domain 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1490.20">
        <property type="entry name" value="ATP-grasp fold, A domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.470.20">
        <property type="entry name" value="ATP-grasp fold, B domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.460.10">
        <property type="entry name" value="Biotin dependent carboxylase carboxyltransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1770.10">
        <property type="entry name" value="PreATP-grasp domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049076">
        <property type="entry name" value="ACCA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049074">
        <property type="entry name" value="ACCA_BT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034733">
        <property type="entry name" value="AcCoA_carboxyl_beta"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013537">
        <property type="entry name" value="AcCoA_COase_cen"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011761">
        <property type="entry name" value="ATP-grasp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013815">
        <property type="entry name" value="ATP_grasp_subdomain_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005481">
        <property type="entry name" value="BC-like_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011764">
        <property type="entry name" value="Biotin_carboxylation_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005482">
        <property type="entry name" value="Biotin_COase_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000089">
        <property type="entry name" value="Biotin_lipoyl"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029045">
        <property type="entry name" value="ClpP/crotonase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011763">
        <property type="entry name" value="COA_CT_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011762">
        <property type="entry name" value="COA_CT_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005479">
        <property type="entry name" value="CPAse_ATP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016185">
        <property type="entry name" value="PreATP-grasp_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011054">
        <property type="entry name" value="Rudment_hybrid_motif"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011053">
        <property type="entry name" value="Single_hybrid_motif"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45728:SF1">
        <property type="entry name" value="ACETYL-COA CARBOXYLASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45728">
        <property type="entry name" value="ACETYL-COA CARBOXYLASE, ISOFORM A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08326">
        <property type="entry name" value="ACC_central"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21385">
        <property type="entry name" value="ACCA_BT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02785">
        <property type="entry name" value="Biotin_carb_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00289">
        <property type="entry name" value="Biotin_carb_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00364">
        <property type="entry name" value="Biotin_lipoyl"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01039">
        <property type="entry name" value="Carboxyl_trans"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02786">
        <property type="entry name" value="CPSase_L_D2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00878">
        <property type="entry name" value="Biotin_carb_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52096">
        <property type="entry name" value="ClpP/crotonase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56059">
        <property type="entry name" value="Glutathione synthetase ATP-binding domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52440">
        <property type="entry name" value="PreATP-grasp domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51246">
        <property type="entry name" value="Rudiment single hybrid motif"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51230">
        <property type="entry name" value="Single hybrid motif"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50975">
        <property type="entry name" value="ATP_GRASP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50979">
        <property type="entry name" value="BC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50968">
        <property type="entry name" value="BIOTINYL_LIPOYL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50989">
        <property type="entry name" value="COA_CT_CTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50980">
        <property type="entry name" value="COA_CT_NTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00866">
        <property type="entry name" value="CPSASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00867">
        <property type="entry name" value="CPSASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8 13" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="11" id="KW-0092">Biotin</keyword>
    <keyword evidence="10" id="KW-0275">Fatty acid biosynthesis</keyword>
    <keyword evidence="7" id="KW-0276">Fatty acid metabolism</keyword>
    <keyword evidence="5" id="KW-0436">Ligase</keyword>
    <keyword evidence="4" id="KW-0444">Lipid biosynthesis</keyword>
    <keyword evidence="9" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="12" id="KW-0511">Multifunctional enzyme</keyword>
    <keyword evidence="6 13" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="15" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="15">
        <location>
            <begin position="1"/>
            <end position="21"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019143682" description="acetyl-CoA carboxylase" evidence="15">
        <location>
            <begin position="22"/>
            <end position="2372"/>
        </location>
    </feature>
    <feature type="domain" description="Biotin carboxylation" evidence="18">
        <location>
            <begin position="255"/>
            <end position="756"/>
        </location>
    </feature>
    <feature type="domain" description="ATP-grasp" evidence="17">
        <location>
            <begin position="413"/>
            <end position="604"/>
        </location>
    </feature>
    <feature type="domain" description="Lipoyl-binding" evidence="16">
        <location>
            <begin position="883"/>
            <end position="957"/>
        </location>
    </feature>
    <feature type="domain" description="CoA carboxyltransferase N-terminal" evidence="19">
        <location>
            <begin position="1676"/>
            <end position="2006"/>
        </location>
    </feature>
    <feature type="domain" description="CoA carboxyltransferase C-terminal" evidence="20">
        <location>
            <begin position="2010"/>
            <end position="2326"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="24"/>
            <end position="50"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001953"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004956"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00013058"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022516"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022598"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022832"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023160"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023267"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023268"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00409"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50975"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50979"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50980"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007032.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2372" mass="267455" checksum="6613A98BAE3F4123" modified="2019-05-08" version="1">MVLLALLSCLILLWFIWTEMANSPPVNQQTERPSPEPPAASENKEKPSKGSTCLLQERAVTPPQSTGPAQTSPTSGHVSVDALQQAASMLWQHKLPNIPSSSESAQDVSANSLAEPLCQVLPPSSKQQMKLKKSISREKQEQFIEHTQVQLKRQAMAKFVLGSFEDNSSDDELIPGSFKVTSRSSMMSVGSSSSFESFLLTSLSESGSSSTTMRPSMSGLHLVKRGREQKKLELQRDFTVASPAEFVTRFGGNRIIEKVLIANNGIAAVKCMRSIRRWAYEMFRNERAIRFVVMVTPEDLKANAEYIKMADHYVPVPGGANNNNYANVELIVDISKRIPVQAVWAGWGHASENPRLPELLQKHGVAFLGPPSEAMWALGDKVASTIVAQTVQIPTLPWSGSGLMAEWTEDCEQRRTISVSLETYARGCVKDVDESLEAAERIGYPVMIKASEGGGGKGIRRAETAEEFTGCFRQVQSEAPGSPIFVMKLAQHARHLEVQVLADQYGNAVSLFGRDCSLQRRHQKIIEEAPVTIAAPSVFDFMEQCAVRLAKMVGYVSAGTVEYLYSEDGGFHFLELNPRLQVEHPCTEMIADVNLPAAQLQIAMGVPLHRIKDIRVLYGESPWADTPICFESPANIPVPRGHVIAARITSENPDEEFKPSSGTVQELNFRSSKNVWGYFSVAAAGGLHEFADSQFGHCFSWGENREEAISNMVVALKELSIRGDFRTTVEYLIKLLETESFQNNEIDTGWLDHLIAEKVQAEKPDTMLGVVCCALNVADALFRTCMTDFLHSLERGQVLPAASLLNIVEVELIYEGVKYALKVARQSLTTYVIIMNNTHIEIDVHRLNDGGVLLSYDGNSYTTYMKEEVDCYRITVGNKTCYFEKEKDPTMLRSPSAGKLLKYTVEDGGHIFIGHSFAEIEVMKIIFTLEVKESGHIRYIKRPGALLEAGCVIARLELDDPTKVHPAQLYTGGLPTQQMLPIGGLKLHQVFHSVLENLINVMNGYCLPEPYFSAKMKEWVCELMKTLRDPSLPLLELQEIMTNVSGRIPLSVEKSIRKVMAQYASNITSVLCQFPSQQIASILDSYAASLQRKADREVFFMNTQSIVQLVQRYRSGIRGYMKSVVLDLLRRYLQVETQFQQAHYDKCVINLREQYKLDMNPVLECIFSHAQVAKKNLLVIMLIDQLCGRDSTMTEELMAILTELTQLSKTEHSKVALRARQVLIASHLPSYELRHNQVESIFLSAIDMYGHQFCPEDLKKLILSETTIFDVLPTFFYHSNQLVRMAALEVYVRRGYIAYELNSLQHRQLSDGTCVVEFQFMLPSSHPNRMSIPISVSNPDLARHSTELFMDSGFSPLCQRMGAMVAFDRFEDFTRESPVQRGAXSCFADPPSESPLFSEAQSTLYEEGDNKTIQEEPIHILNTAIRWAGHLEDEELVPVFRTFAQSKVLFNFVNFFTFRARDEFAEDRIYRHLEPALAFQLELNRMRNFDLTAVPCANHRMHLYLGAAKVEEGNEVTDYRFFIRAIIRHSDLITKEASFEYLQNEGERLLLEAMDELEVAFSNTSVRTDCNHIFLNFVPTVIMDPSKIEEAVRSMVMRYGSRLWKLRVLQAEVKINIRLTPVGKAIPIRLFLTNESGYYLDISLYKEVNDPRTGNIMFHSYGDKQGLQHGMLINTPYVTKDLLQSKRFQAQSLGTTYVYDFPEMFKQALFKLWGSTEMYPKDVLTYTELVLDSQGQLVEMNRIPGGNEVGMVAFKMKLKTPEYPKGRDMIVISNDITCKIGSFGPEEDLLFLRASQLARSEGIPRIYIAANSGARIGFADEKKHMFQVAWVDPMDPYKGFKYLYLTPQDYTRINSVNSVHCEHVEEGGESRYVLTDIIGKDEGFGVENLRASGTIAGESSIAYDEIVTISMVTCRAIGIGAYLVRLGQRVIQVENSHIILTGASALNKVLGREVYTSNNQLGGVQIMHNNGVSHVTVPDDFEGIYTILQWLSYMPKDNRSPVPVTATSDPIEREIDFVPSKGPYDPRWMLAGRPHPIVKGTWQSGFFDHGTFMEIMQPWAQTVVVGRARLGGIPIGVVAVETRTVEMAIPADPANLDSEAKIVQQAGQVWFPDSAFKTAQAIRDFDRERLPLLIFANWRGFSGGMKDMYDQVLKFGAYIVDSLRQFKQPVLVYIPPHAELRGGSWVVIDPTINPVYMELYADKESRGGILEAEGTVEIKFRRKDLIKTMRRIDSVCARLVEQLGTPELLEEDHRELKKQLKAREELLLPIFHQVAVQFADLHDTPGRMHEKGVIADILEWKNARTFLYWRLRRLLLEEVVKTEILNADSELSNIHIQSMLRRWFMETEGAVKVSLHLENHWRASLFPCYGSV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GY49</accession>
    <name>A0A452GY49_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Tyrosine-protein phosphatase non-receptor type</fullName>
            <ecNumber evidence="10">3.1.3.48</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="9 20">
            <text>O-phospho-L-tyrosyl-[protein] + H2O = L-tyrosyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:10684"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="EC" id="3.1.3.48"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 10">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 10">Belongs to the protein-tyrosine phosphatase family. Non-receptor class 2 subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.48" evidence="10"/>
    <dbReference type="AlphaFoldDB" id="A0A452GY49"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008084.1">
        <property type="protein sequence ID" value="ENSGAGP00000006977.1"/>
        <property type="gene ID" value="ENSGAGG00000005618.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004726">
        <property type="term" value="F:non-membrane spanning protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001784">
        <property type="term" value="F:phosphotyrosine residue binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035556">
        <property type="term" value="P:intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000278">
        <property type="term" value="P:mitotic cell cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd14606">
        <property type="entry name" value="PTPc-N6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd09931">
        <property type="entry name" value="SH2_C-SH2_SHP_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd10340">
        <property type="entry name" value="SH2_N-SH2_SHP_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.505.10:FF:000012">
        <property type="entry name" value="Tyrosine-protein phosphatase non-receptor type"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.505.10:FF:000018">
        <property type="entry name" value="Tyrosine-protein phosphatase non-receptor type"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000018">
        <property type="entry name" value="Tyrosine-protein phosphatase non-receptor type"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.505.10">
        <property type="entry name" value="SH2 domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052123">
        <property type="entry name" value="Non-rcpt_Tyr_Phosphatase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000242">
        <property type="entry name" value="PTP_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000980">
        <property type="entry name" value="SH2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036860">
        <property type="entry name" value="SH2_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016130">
        <property type="entry name" value="Tyr_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003595">
        <property type="entry name" value="Tyr_Pase_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012152">
        <property type="entry name" value="Tyr_Pase_non-rcpt_typ-6/11"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46257">
        <property type="entry name" value="TYROSINE-PROTEIN PHOSPHATASE CORKSCREW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46257:SF4">
        <property type="entry name" value="TYROSINE-PROTEIN PHOSPHATASE NON-RECEPTOR TYPE 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00017">
        <property type="entry name" value="SH2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00102">
        <property type="entry name" value="Y_phosphatase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000929">
        <property type="entry name" value="Tyr-Ptase_nr_6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00700">
        <property type="entry name" value="PRTYPHPHTASE"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00401">
        <property type="entry name" value="SH2DOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00194">
        <property type="entry name" value="PTPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00404">
        <property type="entry name" value="PTPc_motif"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00252">
        <property type="entry name" value="SH2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55550">
        <property type="entry name" value="SH2 domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50001">
        <property type="entry name" value="SH2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00383">
        <property type="entry name" value="TYR_PHOSPHATASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50055">
        <property type="entry name" value="TYR_PHOSPHATASE_PTP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 10" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6 10" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="7 10" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="8 13" id="KW-0727">SH2 domain</keyword>
    <feature type="domain" description="SH2" evidence="15">
        <location>
            <begin position="4"/>
            <end position="100"/>
        </location>
    </feature>
    <feature type="domain" description="SH2" evidence="15">
        <location>
            <begin position="110"/>
            <end position="224"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="16">
        <location>
            <begin position="255"/>
            <end position="526"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="17">
        <location>
            <begin position="441"/>
            <end position="517"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="557"/>
            <end position="611"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="14">
        <location>
            <begin position="561"/>
            <end position="570"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="587"/>
            <end position="605"/>
        </location>
    </feature>
    <feature type="active site" description="Phosphocysteine intermediate" evidence="11">
        <location>
            <position position="464"/>
        </location>
    </feature>
    <feature type="binding site" evidence="12">
        <location>
            <position position="430"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="12">
        <location>
            <position position="511"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010750"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022999"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00051722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PIRNR" id="PIRNR000929"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PIRSR" id="PIRSR000929-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PIRSR" id="PIRSR000929-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00191"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50001"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006977.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="611" mass="69115" checksum="1225B0782AF88896" modified="2019-05-08" version="1">MVRWFHRDLSGLQAEALLKGRGVHGSFLARPSRKNLGDFSLSVRVGDQVTHIRIQNTGDYYDLYGGEKFATLSELVEYYTQQQGSLQDKDGTVIDLRYPLNCSDPTTERWYHGHLSGPAAESLLQSKAIPWTFLVRESLSKPGDFVLSVLTDQPKAGPEAPPTGAAGTPGARLKVTHVKILCENGRYTVGGAEMFDSLADLVEHFKKTGIEELSGSYVYLRQPYYATRVNAADIENRVHELNKQSPAEETAKAGFWEEFDSLQKQEAKHLYDRHEGQRPENKGKNRYKNILPFDHSRVILQGRDPNIPGADYINANYIKNQLISPDEFTKTYIASQGCLDATVNDFWQMVWQENSRIVVMTTREVEKGRNKCVPYWPEEGSTKEYGPYIVENLGERDALEYKLRQLCLSPIDNCKAVREIWQYQYLGWPDHGVPSEPGGVLSFLDQINQKQESIPEAGPIVVHCSAGIGRTGTIIVIDMIVETISTKGLDCDIDIQKTIQMVRAQRSGMVQTEAQYKFIYMAICQFIETTKKKLDVMQSQKGRAKESEYGNIAYPPALKNQHVKASRKSSKQKEEPTVYENLGNAKGKKDERVRKQRSSEKEKLKSSLKKK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9P0</accession>
    <name>A0A452H9P0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Cleavage and polyadenylation specificity factor subunit 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
            <location evidence="1">Nucleoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the CPSF1 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000013209.1">
        <property type="protein sequence ID" value="ENSGAGP00000011536.1"/>
        <property type="gene ID" value="ENSGAGG00000004276.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005654">
        <property type="term" value="C:nucleoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003676">
        <property type="term" value="F:nucleic acid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031123">
        <property type="term" value="P:RNA 3'-end processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.910:FF:000005">
        <property type="entry name" value="Cleavage and polyadenylation specific factor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:002223">
        <property type="entry name" value="Cleavage and polyadenylation specific factor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:000100">
        <property type="entry name" value="Cleavage and polyadenylation specificity factor subunit 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:000118">
        <property type="entry name" value="Cleavage and polyadenylation specificity factor subunit 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:000544">
        <property type="entry name" value="cleavage and polyadenylation specificity factor subunit 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.910">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018846">
        <property type="entry name" value="Beta-prop_RSE1/DDB1/CPSF1_1st"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058543">
        <property type="entry name" value="Beta-prop_RSE1/DDB1/CPSF1_2nd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050358">
        <property type="entry name" value="RSE1/DDB1/CFT1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004871">
        <property type="entry name" value="RSE1/DDB1/CPSF1_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10644">
        <property type="entry name" value="DNA REPAIR/RNA PROCESSING CPSF FAMILY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10433">
        <property type="entry name" value="Beta-prop_RSE1_1st"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23726">
        <property type="entry name" value="Beta-prop_RSE1_2nd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03178">
        <property type="entry name" value="CPSF_A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0539">Nucleus</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="RSE1/DDB1/CPSF1 first beta-propeller" evidence="7">
        <location>
            <begin position="14"/>
            <end position="403"/>
        </location>
    </feature>
    <feature type="domain" description="RSE1/DDB1/CPSF1 second beta-propeller" evidence="8">
        <location>
            <begin position="520"/>
            <end position="991"/>
        </location>
    </feature>
    <feature type="domain" description="RSE1/DDB1/CPSF1 C-terminal" evidence="6">
        <location>
            <begin position="1064"/>
            <end position="1384"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="407"/>
            <end position="437"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="542"/>
            <end position="569"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="742"/>
            <end position="764"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="542"/>
            <end position="559"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="749"/>
            <end position="764"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004642"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00038446"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00068483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="Pfam" id="PF03178"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF10433"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF23726"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011536.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1419" mass="158107" checksum="A0A81602AD6F8634" modified="2019-05-08" version="1">MYAVYRQAHPPTGVEFSMYCNFFSNAERNLVVAGTSQLYVYRLNHDSEVPCPLTKQSTHAEGKGHKEKLELVASFSFFGNVMSMASVQLAGAKRDALLLSFKDAKLSVVEYDPGTHDLKTLSLHYFEEPELKDGFVQNVHIPKVRVDPDGRCAVMLIYGTRLVVLPFRRETLTDEHEGVMGEGQKSSFLPSYIIDVRELDEKLLNIIDMQFLYGYYEPTLLILFEPNQTWPGRVAVRQDTCSIVAISLNIMQKVHPVIWSLSNLPFDCTQALAVPKPIGGVVIFAVNSLLYLNQSVPPYGVSLNSLTTGTTVFPLRIQDGVKITLDCAQAAFISYDKMVISLKGGEIYVLTLITDGMRSVRSFHFDKAAASVLTTCMITMEPGYLFLGSRLGNSLLLKYTEKLQEPPVNVAKDPEEPPVKKKRSESSGNWAGGKSAPQDEVDEIEVYGSEAQSGTQLATYSFEVCDSILNIGPCANAAMGEPAFLSEEFQNSPEPDLEIVVCSGYGKNGALSVLQKSIRPQVVTTFELPGCYDMWTVISPQKKEEDESAKGESAEKESSPPEPEDDGKRHGFLILSREDSTMILQTGQEIMELDTSGFATQGPTVFAGNIGENRYIVQVSPLGIRLLEGVNQLHFIPVDLGSPIVHCAVADPYVVIMSAEGQVTMFVLKSDTYGGRTHRLTLQKPQLHHQSKVIALCVYRDVRGMFTTESKTASSRDEWVGRGARGRVNTVDDEEEMLYGDSSTLFSPSKEEPRRSSLPLADRDPHQYKVEPTHWCVLVRENGAMEIYQLPDWRLVFLVKNFPMGQRVLVDSSFGQPAAQGDAKKEEVTRQGEMPLVKEVLLVALGNRQSRPYLLVHVDQELLIYEAFSHDSQLGQSNLKVRFKKVPHNINFREKKLKLSKKKSESAGGEESGGARGRVARFRYFEDIYGYSGVFICGPSPHWLLVTARGALRLHPMTIDGLIESFAPFHNVNCPKGFLYFNRQGELRISVLPAYLSYDAPWPVRKIPLRCTTHYVAYHVESKVYAVATSVVNPCTRIPRMTGEEKEFESIERDERYIHPQQEAFSIQLISPVSWETIPNTRIELEEWEHVTCMKTVSLKSEETVSGLKGYIAAGTCLMQGEEVTCRGRILIMDIIEVVPEPGQPLTKNKFKVLYEKEQKGPVTALCHCHGYLVSAIGQKIFLWSLKDNDLTGMAFIDTQLYIHQMISVKNFILAADVMKSISLLRYQEESKTLSLVSRVAQPGTIVYSVDFMVDTDQLGFLGRFCSFVPCTLAPTASGGGSGQARGARARTPWPCCGDPAACTLLPAPGQSCLATLDGGIGLLLPMLEKTYRRLLMLQNALTTMLSHHAGLNPRAFRMLHVDRRILQNAVRNILDGELLSRYLYLSAMERSELAKKIGTTPDIILEDLLEIDRVTAHF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9R0</accession>
    <name>A0A452H9R0_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="13">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9R0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013231.1">
        <property type="protein sequence ID" value="ENSGAGP00000011556.1"/>
        <property type="gene ID" value="ENSGAGG00000008921.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd07765">
        <property type="entry name" value="KRAB_A-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000352">
        <property type="entry name" value="zinc finger protein 3 homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002343">
        <property type="entry name" value="Zinc finger protein 33A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000127">
        <property type="entry name" value="Zinc finger protein 354C"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000135">
        <property type="entry name" value="Zinc finger protein 358"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000710">
        <property type="entry name" value="Zinc finger protein 768"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001954">
        <property type="entry name" value="Zinc finger protein 787"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001909">
        <property type="entry name" value="KRAB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036051">
        <property type="entry name" value="KRAB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23235:SF178">
        <property type="entry name" value="C2H2-TYPE DOMAIN-CONTAINING PROTEIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23235">
        <property type="entry name" value="KRUEPPEL-LIKE TRANSCRIPTION FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01352">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SMART" id="SM00349">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109640">
        <property type="entry name" value="KRAB domain (Kruppel-associated box)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50805">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="9"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 9" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="KRAB" evidence="12">
        <location>
            <begin position="3"/>
            <end position="74"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="11">
        <location>
            <begin position="113"/>
            <end position="143"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="11">
        <location>
            <begin position="144"/>
            <end position="171"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="11">
        <location>
            <begin position="172"/>
            <end position="199"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="11">
        <location>
            <begin position="200"/>
            <end position="227"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="11">
        <location>
            <begin position="228"/>
            <end position="255"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="11">
        <location>
            <begin position="256"/>
            <end position="283"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="11">
        <location>
            <begin position="284"/>
            <end position="311"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="11">
        <location>
            <begin position="312"/>
            <end position="334"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="11">
        <location>
            <begin position="337"/>
            <end position="364"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="74"/>
            <end position="93"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011556.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="370" mass="41737" checksum="43F27FBBD2302DE9" modified="2019-05-08" version="1">LSQECKWFSETAGTGEWALLNPAQRALYREVTQENYATVASLGFPVSKPDVISQLERGEEPWVPDLQGSEERELQKGAQIGEASIKPTQKLEPRRTLAGRCKEKVSGHCGEAKACQNQHRPEKSFSSGSDLIQHERINMGKRPYTCSECGKSFSQSSTLITHYRVHTGERPYTCHECGKSFSHGSALITHQRIHTGEMPYTCPECGKSFNHSSNLIKHRRIHTGETLYTCSECGRNFCRRSALITHQRIHTGETPYSCTECKKTFGESSDLIRHCIIHTGEKPYTCSECGKSFNQSSNLITHLRIHTGEKPYTCSECGKSFCRSSNLITHQRIHRPYTCSECGKSFSRNSNLITHQRLHTGERPFTCTEV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9R4</accession>
    <name>A0A452H9R4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">Integrin beta</fullName>
        </recommendedName>
    </protein>
    <organism evidence="26 27">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="27" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="26" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="26" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="20">Heterodimer of an alpha and a beta subunit. Interacts with FLNB. Interacts with HAX1. ITGAV:ITGB6 interacts with FBN1. ITGAV:ITGB6 interacts with TGFB1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell junction</location>
            <location evidence="1">Focal adhesion</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2 21">Cell membrane</location>
            <topology evidence="2 21">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 21">Belongs to the integrin beta chain family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9R4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013171.1">
        <property type="protein sequence ID" value="ENSGAGP00000011502.1"/>
        <property type="gene ID" value="ENSGAGG00000007913.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009986">
        <property type="term" value="C:cell surface"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005925">
        <property type="term" value="C:focal adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034685">
        <property type="term" value="C:integrin alphav-beta6 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033627">
        <property type="term" value="P:cell adhesion mediated by integrin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098609">
        <property type="term" value="P:cell-cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007160">
        <property type="term" value="P:cell-matrix adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007229">
        <property type="term" value="P:integrin-mediated signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.100:FF:000004">
        <property type="entry name" value="Integrin beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000043">
        <property type="entry name" value="Integrin beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000075">
        <property type="entry name" value="Integrin beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000328">
        <property type="entry name" value="Integrin beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.410:FF:000002">
        <property type="entry name" value="Integrin beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.1240.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.100">
        <property type="entry name" value="Cytochrome c1, transmembrane anchor, C-terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1510">
        <property type="entry name" value="ntegrin, alpha v. Chain A, domain 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.410">
        <property type="entry name" value="von Willebrand factor, type A domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040622">
        <property type="entry name" value="EGF_integrin_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057073">
        <property type="entry name" value="EGF_integrin_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015812">
        <property type="entry name" value="Integrin_bsu"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014836">
        <property type="entry name" value="Integrin_bsu_cyt_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012896">
        <property type="entry name" value="Integrin_bsu_tail"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036349">
        <property type="entry name" value="Integrin_bsu_tail_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002369">
        <property type="entry name" value="Integrin_bsu_VWA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032695">
        <property type="entry name" value="Integrin_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057243">
        <property type="entry name" value="Integrin_I-EGF_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036465">
        <property type="entry name" value="vWFA_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10082">
        <property type="entry name" value="INTEGRIN BETA SUBUNIT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10082:SF11">
        <property type="entry name" value="INTEGRIN BETA-6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23105">
        <property type="entry name" value="EGF_integrin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18372">
        <property type="entry name" value="I-EGF_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08725">
        <property type="entry name" value="Integrin_b_cyt"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07965">
        <property type="entry name" value="Integrin_B_tail"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00362">
        <property type="entry name" value="Integrin_beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01186">
        <property type="entry name" value="INTEGRINB"/>
    </dbReference>
    <dbReference type="SMART" id="SM00187">
        <property type="entry name" value="INB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01241">
        <property type="entry name" value="Integrin_b_cyt"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01242">
        <property type="entry name" value="Integrin_B_tail"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69687">
        <property type="entry name" value="Integrin beta tail domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69179">
        <property type="entry name" value="Integrin domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53300">
        <property type="entry name" value="vWA-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00243">
        <property type="entry name" value="I_EGF_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS52047">
        <property type="entry name" value="I_EGF_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0106">Calcium</keyword>
    <keyword evidence="12 21" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="13" id="KW-0965">Cell junction</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="17" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="19" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="15 21" id="KW-0401">Integrin</keyword>
    <keyword evidence="11" id="KW-0460">Magnesium</keyword>
    <keyword evidence="16 22" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="18" id="KW-0675">Receptor</keyword>
    <keyword evidence="27" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0677">Repeat</keyword>
    <keyword evidence="8" id="KW-0732">Signal</keyword>
    <keyword evidence="6 21 22" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="14 22" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="22">
        <location>
            <begin position="613"/>
            <end position="635"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin beta subunit VWA" evidence="23">
        <location>
            <begin position="6"/>
            <end position="359"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin beta subunit tail" evidence="25">
        <location>
            <begin position="529"/>
            <end position="612"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin beta subunit cytoplasmic" evidence="24">
        <location>
            <begin position="636"/>
            <end position="682"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007449"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023037"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00046864"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="RuleBase" id="RU000633"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="SMART" id="SM00187"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="SMART" id="SM01241"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="SMART" id="SM01242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011502.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="694" mass="76584" checksum="D8D7DCA29CF1D436" modified="2019-05-08" version="1">DHSVLVKCVNLFSISPMALLGNEETIEIKVRQTEDYPVDLYYLMDLSASMDDDLNTIKELGSTLSKEMSKLTSNFRLGFGSFVEKPVSPFIKTTAEEINNPCRSVPRDCLPTFGYKHVLPLTNDAERFNEIVKEQKISANIDTPEGGFDAIMQAAVCKEKIGWRNDSLHLLVFVSDADSHFGMDSKLAGIVIPNDGHCHLDQNNEYSMSTVLEYPTIGQLTDKLVQNNVLLIFAVTKEQVYFYENYAKLIPGATVGQLQKDSGNILQLIIDAYKELRSEVELEVLGDTEGLNLSFTAICNNGSIFPHQRKCSHIKVGETVSFNMTVSLPTCEKRRRQIVIKPVGLSDALEIDIQLECSCNCQKEAEMNSSKCSKGKGSFECGVCACNPGYMGPYCECNENSLSTNSCKRSPEQSSCSGRGDCYCGQCICHFSVYGNIYGLYCECDNFSCVRFKGLQCGGNGDCDCGECVCHRGWTGEYCNCTTDTNSCVSEDGILCSGRGECICGTCVCTNPGASGSTCERCPTCGDPCNSKRSCIECHLASDVQSQAECNEKCRLVDATVNSAEDFSKDKSIPCSLQGENECVTTFLMATDEQGRTIIHSIKQKDCPQPPNIPMIMLGVSLAILLIGIVLLCIWKLLVSFQDRKEVAKFEAERSKAKWQTGTNPLYRGSTTTFKNVTYKHQEKQKVDITADFY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9R5</accession>
    <name>A0A452H9R5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Peptidyl-prolyl cis-trans isomerase</fullName>
            <shortName evidence="6">PPIase</shortName>
            <ecNumber evidence="6">5.2.1.8</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="6">PPIases accelerate the folding of proteins. It catalyzes the cis-trans isomerization of proline imidic peptide bonds in oligopeptides.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="1 11">
            <text>[protein]-peptidylproline (omega=180) = [protein]-peptidylproline (omega=0)</text>
            <dbReference type="Rhea" id="RHEA:16237"/>
            <dbReference type="Rhea" id="RHEA-COMP:10747"/>
            <dbReference type="Rhea" id="RHEA-COMP:10748"/>
            <dbReference type="ChEBI" id="CHEBI:83833"/>
            <dbReference type="ChEBI" id="CHEBI:83834"/>
            <dbReference type="EC" id="5.2.1.8"/>
        </reaction>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the cyclophilin-type PPIase family.</text>
    </comment>
    <dbReference type="EC" id="5.2.1.8" evidence="6"/>
    <dbReference type="AlphaFoldDB" id="A0A452H9R5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013236.1">
        <property type="protein sequence ID" value="ENSGAGP00000011561.1"/>
        <property type="gene ID" value="ENSGAGG00000008887.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016018">
        <property type="term" value="F:cyclosporin A binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003755">
        <property type="term" value="F:peptidyl-prolyl cis-trans isomerase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006457">
        <property type="term" value="P:protein folding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.100.10:FF:000020">
        <property type="entry name" value="E3 SUMO-protein ligase RanBP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.100.10">
        <property type="entry name" value="Cyclophilin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029000">
        <property type="entry name" value="Cyclophilin-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024936">
        <property type="entry name" value="Cyclophilin-type_PPIase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020892">
        <property type="entry name" value="Cyclophilin-type_PPIase_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002130">
        <property type="entry name" value="Cyclophilin-type_PPIase_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11071">
        <property type="entry name" value="PEPTIDYL-PROLYL CIS-TRANS ISOMERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11071:SF561">
        <property type="entry name" value="PEPTIDYL-PROLYL CIS-TRANS ISOMERASE D-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00160">
        <property type="entry name" value="Pro_isomerase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF001467">
        <property type="entry name" value="Peptidylpro_ismrse"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00153">
        <property type="entry name" value="CSAPPISMRASE"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50891">
        <property type="entry name" value="Cyclophilin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00170">
        <property type="entry name" value="CSA_PPIASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50072">
        <property type="entry name" value="CSA_PPIASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5 6" id="KW-0413">Isomerase</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="4 6" id="KW-0697">Rotamase</keyword>
    <feature type="domain" description="PPIase cyclophilin-type" evidence="8">
        <location>
            <begin position="39"/>
            <end position="194"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="2"/>
            <end position="29"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000971"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023110"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023235"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="RuleBase" id="RU363019"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50072"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011561.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="195" mass="21859" checksum="2275D399AB40EEA2" modified="2019-05-08" version="1">MADSFKRRFEECQQNLSELQRAHVSLAAELSKETNPVVYFEVSADDEPLGHITMELFSNIVPRTAENFRALCTGEKGFGFKNSIFHRIVPDFVCQGGDITRHDGTGGRSIYGDAFEDENFEVKHTGPGLLSMANRGRDTNNSQFFITLKKAEHLDFKHVVFGFVKDGMDVVKKMESFGSPKGLMNRIVITDCGQI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9R6</accession>
    <name>A0A452H9R6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Guanylate cyclase soluble subunit alpha-1</fullName>
            <ecNumber evidence="3">4.6.1.2</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="10">Soluble guanylate cyclase large subunit</fullName>
        </alternativeName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>GTP = 3',5'-cyclic GMP + diphosphate</text>
            <dbReference type="Rhea" id="RHEA:13665"/>
            <dbReference type="ChEBI" id="CHEBI:33019"/>
            <dbReference type="ChEBI" id="CHEBI:37565"/>
            <dbReference type="ChEBI" id="CHEBI:57746"/>
            <dbReference type="EC" id="4.6.1.2"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="11">Belongs to the adenylyl cyclase class-4/guanylyl cyclase family.</text>
    </comment>
    <dbReference type="EC" id="4.6.1.2" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452H9R6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000012978.1">
        <property type="protein sequence ID" value="ENSGAGP00000011334.1"/>
        <property type="gene ID" value="ENSGAGG00000008765.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000012981.1">
        <property type="protein sequence ID" value="ENSGAGP00000011337.1"/>
        <property type="gene ID" value="ENSGAGG00000008765.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008074">
        <property type="term" value="C:guanylate cyclase complex, soluble"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005525">
        <property type="term" value="F:GTP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004383">
        <property type="term" value="F:guanylate cyclase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0020037">
        <property type="term" value="F:heme binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019934">
        <property type="term" value="P:cGMP-mediated signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070482">
        <property type="term" value="P:response to oxygen levels"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07302">
        <property type="entry name" value="CHD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.450.260:FF:000002">
        <property type="entry name" value="guanylate cyclase soluble subunit alpha-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.1230:FF:000007">
        <property type="entry name" value="Guanylate cyclase soluble subunit alpha-3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1520.10:FF:000002">
        <property type="entry name" value="Guanylate cyclase soluble subunit alpha-3 isoform A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1520.10">
        <property type="entry name" value="H-NOX domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.450.260">
        <property type="entry name" value="Haem NO binding associated domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.1230">
        <property type="entry name" value="Nucleotide cyclase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001054">
        <property type="entry name" value="A/G_cyclase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018297">
        <property type="entry name" value="A/G_cyclase_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038158">
        <property type="entry name" value="H-NOX_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042463">
        <property type="entry name" value="HNOB_dom_associated_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024096">
        <property type="entry name" value="NO_sig/Golgi_transp_ligand-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029787">
        <property type="entry name" value="Nucleotide_cyclase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45655:SF4">
        <property type="entry name" value="GUANYLATE CYCLASE SOLUBLE SUBUNIT ALPHA-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45655">
        <property type="entry name" value="GUANYLATE CYCLASE SOLUBLE SUBUNIT BETA-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00211">
        <property type="entry name" value="Guanylate_cyc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00044">
        <property type="entry name" value="CYCc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF111126">
        <property type="entry name" value="Ligand-binding domain in the NO signalling and Golgi transport"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55073">
        <property type="entry name" value="Nucleotide cyclase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00452">
        <property type="entry name" value="GUANYLATE_CYCLASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50125">
        <property type="entry name" value="GUANYLATE_CYCLASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0141">cGMP biosynthesis</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0342">GTP-binding</keyword>
    <keyword evidence="7 11" id="KW-0456">Lyase</keyword>
    <keyword evidence="5" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Guanylate cyclase" evidence="12">
        <location>
            <begin position="452"/>
            <end position="579"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001436"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012202"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023293"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00040514"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00042184"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="RuleBase" id="RU000405"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011337.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="661" mass="74741" checksum="DC5111BD8F9234A6" modified="2019-05-08" version="1">MFCTKLKDLKITGECPFSLLTQSHISDEHEEECAEVPNSSRAALPICKEVSEKDTHGTLPQRKTSRSRVYLHTLAESICKLLFPEFERLHLALQRTLAKNKIKENRKYGEREDFEKIVTDHANAAGVPVETVKEYLGEELFKICYEEDEHILGVVGGTLKDFLNSFNTLLKQSGHCQEADKKSKLEEASILCLEKDQDFLNVYCFFPKKITGLILPGIIKAAAHILYETEVEVMLMPPCFRNDCTEFINQPYLLYSVLVKSTRPSLSPCKPQSSLVISASLFCKTFPFHFMFDKDMAVLQVGNGIRRLLNRREFLANPNFEECFEILTPRINCTFSGIMTMLNMQFTVRVKRWDNAVKKSSRVMDLKGQMIYIFESSAILFLGSPCVDRLEDFTGRGLYLSDIPIHNALRDVVLIGEQARAQDGLKKRLGKLKATLEMAHQALEEEKKKTVDLLCSIFPGFTAICAQCSPMQVITMLNELYTRFDYQCGELDVYKVETIGDAYCVAGGLHKESETHAVQIALMALKMMELSDEVMSPHGEPIKMRIGLHSGSVFAGVVGVKMPRYCLFGNNVTLANKFESCSIPGKINVSPTTYRLLKEYPDFVFTPRSREELPPNFPSDIPGICYFLDAYLQGTNSKPWFQKRDFEDGNANFLGKATGVD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9S2</accession>
    <name>A0A452H9S2_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="14">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Lipid-anchor</topology>
            <topology evidence="1">GPI-anchor</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9S2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000012989.1">
        <property type="protein sequence ID" value="ENSGAGP00000011344.1"/>
        <property type="gene ID" value="ENSGAGG00000008630.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030424">
        <property type="term" value="C:axon"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043025">
        <property type="term" value="C:neuronal cell body"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098552">
        <property type="term" value="C:side of membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008046">
        <property type="term" value="F:axon guidance receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007156">
        <property type="term" value="P:homophilic cell adhesion via plasma membrane adhesion molecules"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050808">
        <property type="term" value="P:synapse organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00096">
        <property type="entry name" value="Ig"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="CDD" id="cd06263">
        <property type="entry name" value="MAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000240">
        <property type="entry name" value="MAM domain containing glycosylphosphatidylinositol anchor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000262">
        <property type="entry name" value="MAM domain containing glycosylphosphatidylinositol anchor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000303">
        <property type="entry name" value="MAM domain containing glycosylphosphatidylinositol anchor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.200:FF:000019">
        <property type="entry name" value="MAM domain containing glycosylphosphatidylinositol anchor 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000165">
        <property type="entry name" value="MAM domain containing glycosylphosphatidylinositol anchor 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000243">
        <property type="entry name" value="MAM domain-containing glycosylphosphatidylinositol anchor protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.200">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050958">
        <property type="entry name" value="Cell_Adh-Cytoskel_Orgn"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000998">
        <property type="entry name" value="MAM_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45080">
        <property type="entry name" value="CONTACTIN 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45080:SF32">
        <property type="entry name" value="MAM DOMAIN CONTAINING GLYCOSYLPHOSPHATIDYLINOSITOL ANCHOR 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13927">
        <property type="entry name" value="Ig_3"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00629">
        <property type="entry name" value="MAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00020">
        <property type="entry name" value="MAMDOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SMART" id="SM00137">
        <property type="entry name" value="MAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50060">
        <property type="entry name" value="MAM_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="7" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="8" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="3" id="KW-0336">GPI-anchor</keyword>
    <keyword evidence="10" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="9" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="6" id="KW-0472">Membrane</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <feature type="domain" description="Ig-like" evidence="12">
        <location>
            <begin position="6"/>
            <end position="103"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="12">
        <location>
            <begin position="112"/>
            <end position="210"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="12">
        <location>
            <begin position="220"/>
            <end position="303"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="12">
        <location>
            <begin position="318"/>
            <end position="412"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="12">
        <location>
            <begin position="420"/>
            <end position="511"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="12">
        <location>
            <begin position="517"/>
            <end position="605"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="13">
        <location>
            <begin position="617"/>
            <end position="717"/>
        </location>
    </feature>
    <feature type="domain" description="MAM" evidence="11">
        <location>
            <begin position="725"/>
            <end position="892"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004609"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022622"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50060"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011344.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="929" mass="104087" checksum="AECA36078975E24D" modified="2019-05-08" version="1">MLLAFPAQIIHAGQACVVKEDNISERVYTIREGNTLVLQCLVTGHPRPQVRWTKTAGSASDKFQETSVFNETLRIEKIQRLQGGRYYCKAENGVGVPAIKSIRVDVQYLDEPILTVHQTISDVRGSFYQEKTVFLRCTVNSNPPARFIWKRGAETLSHSQDNGVDIYEPLYTQGETKVLKLKNLRPQDYASYTCQVSVRNVCSILDKTITFQLTNTTAPPALKLSVNETLVVNPGDNVTMQCSLIGGDPLPEVVWSHSPSPMPHNSLIQKGNLTIWGICMEDSGYYNCTAINNVGNPAKKTVNLLVRSMKNVTFQITPDVIKESESIQLGQDLKLSCHVDAVPQEKVIYFWYKNGKPARVSDRLLISRNDPELPPVTSSLEIIDLRFSDYGTYLCIASFQGAPIPDLSVEVNISSETVPPTISVPKGQSTITVREGSRAELQCEVRGKPKPPIIWSRVDKEAPMPSGAMTVETYDGKLCLESVSREMSGTYKCQTARYNGFNIRPREALVQLNVQFPPVVEPTSQDVRQGMGRSVTLRCTMLKGSPMKVATAVWRFNGNLLTLLPTEQQDYSELMLDSVNRETSGSYECSISNDVGVSTCLFHVSAKAYNPEFYYDTPNPTLSQKQSKNYSYVLQWTQKEPDAVDPILKYRLDVRQLNQRNTVQTLIPVKKMEKGLLLEHILRNLKVPQSYEVRLTPITNFGAGDMAARIIRYMEPINYPNLADNTCHFEDEKICGFTQDKMDNFDWTRQNALTQNPKRTVNTGPSTDISGTEEGYYMFIEASRPRVQGDKARLISPLYNITGKYFCVSFFYHMYGKHIGSLNLLVRVQNKRAIDTQVWSLSENKGNEWKQAHVPISPPGPFQIIFEGVRGTSYEGDIAIDDVTLKRGDCPRKSIGPNKAVALPGSGVPTLHKSSLGGPLTFFLYVLLR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9S3</accession>
    <name>A0A452H9S3_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="18">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>K(+)(in) = K(+)(out)</text>
            <dbReference type="Rhea" id="RHEA:29463"/>
            <dbReference type="ChEBI" id="CHEBI:29103"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>Li(+)(in) = Li(+)(out)</text>
            <dbReference type="Rhea" id="RHEA:78551"/>
            <dbReference type="ChEBI" id="CHEBI:49713"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="14">
            <text>Na(+)(in) = Na(+)(out)</text>
            <dbReference type="Rhea" id="RHEA:34963"/>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="16">Belongs to the amiloride-sensitive sodium channel (TC 1.A.6) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9S3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000012995.1">
        <property type="protein sequence ID" value="ENSGAGP00000011350.1"/>
        <property type="gene ID" value="ENSGAGG00000008774.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015280">
        <property type="term" value="F:ligand-gated sodium channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.470.10:FF:000006">
        <property type="entry name" value="Acid-sensing ion channel 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.820">
        <property type="entry name" value="Acid-sensing ion channel domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.470.10">
        <property type="entry name" value="Acid-sensing ion channels like domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001873">
        <property type="entry name" value="ENaC"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11690:SF286">
        <property type="entry name" value="ACID-SENSING ION CHANNEL 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11690">
        <property type="entry name" value="AMILORIDE-SENSITIVE SODIUM CHANNEL-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00858">
        <property type="entry name" value="ASC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01078">
        <property type="entry name" value="AMINACHANNEL"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="10" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="12 16" id="KW-0407">Ion channel</keyword>
    <keyword evidence="8 16" id="KW-0406">Ion transport</keyword>
    <keyword evidence="9 17" id="KW-0472">Membrane</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0915">Sodium</keyword>
    <keyword evidence="3 16" id="KW-0894">Sodium channel</keyword>
    <keyword evidence="11 16" id="KW-0739">Sodium transport</keyword>
    <keyword evidence="5 16 17" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 17" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="2 16" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="61"/>
            <end position="79"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="119"/>
            <end position="140"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="458"/>
            <end position="475"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022461"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023053"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023201"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00034430"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00036239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00044635"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="RuleBase" id="RU000679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011350.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="492" mass="56454" checksum="9FAC55A405D077A6" modified="2019-05-08" version="1">IIHTKHSPSQCRLLEKIRLYLTRKPLPSLEERKKFDHEFAMSTSFHGLHNIVQTRSKIRKILWMLVVLGSITIVTWQIWNRFSNYFSWPTTTSIIVQYVENIEFPAVTFCNMNRFQVHAVANLSIIFFLWSIVSGVLHAFDTDNKFSQELKDFLQGNQNFSIEEFTRNNGFYLNSSTLLECDFFGQPCYPEDFEHVFTEYGNCFTFNHMDLPARRRVSVSGRGLSLLFDIKQAQFTDDPTFGFIDAGIIYVIHSPKEPPRFDGLGLSTPVGMHAHAAISQLKTVIQEYPWGECNPNIKLQHHKIYSTYGCLQECKARHIQDKCGCLPFLLPGNGTECDLQKFYSCVSPALYNIELMGLCTVGTHNSTCPVPCEETDYPSTVTYSSFASEKALKFFSTKLKRSPEYIRENLVYIDIKYHDLNYKMTRQQKARSVSELLADVGGQLGLFCGASMITIIEIVEYVFTNFCWLCIFLLLKAPEMPHLTVSCNLPSF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9S5</accession>
    <name>A0A452H9S5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="25">Tyrosine-protein kinase receptor</fullName>
            <ecNumber evidence="25">2.7.10.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="29 30">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="30" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="29" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="29" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="21">
            <text>L-tyrosyl-[protein] + ATP = O-phospho-L-tyrosyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10596"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.10.1"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="25">Belongs to the protein kinase superfamily. Tyr protein kinase family. Insulin receptor subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.10.1" evidence="25"/>
    <dbReference type="AlphaFoldDB" id="A0A452H9S5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013249.1">
        <property type="protein sequence ID" value="ENSGAGP00000011573.1"/>
        <property type="gene ID" value="ENSGAGG00000008928.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030424">
        <property type="term" value="C:axon"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043235">
        <property type="term" value="C:receptor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043121">
        <property type="term" value="F:neurotrophin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005030">
        <property type="term" value="F:neurotrophin receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004714">
        <property type="term" value="F:transmembrane receptor protein tyrosine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048513">
        <property type="term" value="P:animal organ development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007169">
        <property type="term" value="P:cell surface receptor protein tyrosine kinase signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:1990090">
        <property type="term" value="P:cellular response to nerve growth factor stimulus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010976">
        <property type="term" value="P:positive regulation of neuron projection development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051897">
        <property type="term" value="P:positive regulation of phosphatidylinositol 3-kinase/protein kinase B signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd04971">
        <property type="entry name" value="IgI_TrKABC_d5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05094">
        <property type="entry name" value="PTKc_TrkC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000701">
        <property type="entry name" value="Tyrosine-protein kinase receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000251">
        <property type="entry name" value="Tyrosine-protein kinase receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000265">
        <property type="entry name" value="Tyrosine-protein kinase receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000033">
        <property type="entry name" value="Tyrosine-protein kinase receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000035">
        <property type="entry name" value="Tyrosine-protein kinase receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000483">
        <property type="entry name" value="Cys-rich_flank_reg_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013098">
        <property type="entry name" value="Ig_I-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013151">
        <property type="entry name" value="Immunoglobulin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001611">
        <property type="entry name" value="Leu-rich_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020777">
        <property type="entry name" value="NTRK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020446">
        <property type="entry name" value="NTRK3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031635">
        <property type="entry name" value="NTRK_LRRCT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050122">
        <property type="entry name" value="RTK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001245">
        <property type="entry name" value="Ser-Thr/Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008266">
        <property type="entry name" value="Tyr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020635">
        <property type="entry name" value="Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002011">
        <property type="entry name" value="Tyr_kinase_rcpt_2_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416:SF66">
        <property type="entry name" value="NT-3 GROWTH FACTOR RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416">
        <property type="entry name" value="TYROSINE-PROTEIN KINASE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07679">
        <property type="entry name" value="I-set"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00047">
        <property type="entry name" value="ig"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13855">
        <property type="entry name" value="LRR_8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16920">
        <property type="entry name" value="LRRCT_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07714">
        <property type="entry name" value="PK_Tyr_Ser-Thr"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01939">
        <property type="entry name" value="NTKRECEPTOR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01942">
        <property type="entry name" value="NTKRECEPTOR3"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00109">
        <property type="entry name" value="TYRKINASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00082">
        <property type="entry name" value="LRRCT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00219">
        <property type="entry name" value="TyrKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52058">
        <property type="entry name" value="L domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51450">
        <property type="entry name" value="LRR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00109">
        <property type="entry name" value="PROTEIN_KINASE_TYR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00239">
        <property type="entry name" value="RECEPTOR_TYR_KIN_II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12 23 24" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="11" id="KW-0221">Differentiation</keyword>
    <keyword evidence="17" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="19" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="20" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="10" id="KW-0418">Kinase</keyword>
    <keyword evidence="4" id="KW-0433">Leucine-rich repeat</keyword>
    <keyword evidence="15 26" id="KW-0472">Membrane</keyword>
    <keyword evidence="13" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="9 23 24" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="3 25" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="18 25" id="KW-0675">Receptor</keyword>
    <keyword evidence="30" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
    <keyword evidence="5" id="KW-0808">Transferase</keyword>
    <keyword evidence="6 25 26" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="14 26" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="16" id="KW-0829">Tyrosine-protein kinase</keyword>
    <feature type="transmembrane region" description="Helical" evidence="26">
        <location>
            <begin position="12"/>
            <end position="31"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="26">
        <location>
            <begin position="432"/>
            <end position="455"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="28">
        <location>
            <begin position="210"/>
            <end position="300"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="27">
        <location>
            <begin position="477"/>
            <end position="778"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="22">
        <location>
            <position position="618"/>
        </location>
    </feature>
    <feature type="binding site" evidence="23">
        <location>
            <begin position="483"/>
            <end position="491"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="23 24">
        <location>
            <position position="511"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022614"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023137"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00051243"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="PIRSR" id="PIRSR620777-50"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="PIRSR" id="PIRSR620777-51"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="RuleBase" id="RU000312"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="27">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="28">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="29">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011573.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="30">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="789" mass="89322" checksum="11EA416E14789F3E" modified="2019-05-08" version="1">MDVSLCPTKCSFWRVFLLWSIWGDYLLSVLACPANCLCSKIDINCKKPDDGNLFPLLEGQDSGTSNGNTSINITDISRNITSIHIENWKNLQTLNAVDMELYTGLQRLTIRNSGLRSIQPRAFAKNPHLRYINLSDNRLTTLSWQLFQTLRLSELTLERNLFNCSCDIRWIQLWQEKGEANLQKQELYCMNMEAAVISLQEMNITQCDLPEISVSHINLTVREGENAVITCNGSGAPLPDVDWTVADLHSITTHQTNLNWTNVHAINLTLVNVTSEDNGFLLTCIAENVVGMTNASVLLTVYYPPRIQSLEEPELRLEHCIEFVVHGNPAPSLHWLHNGQVLRETEIIRMEFYQQGEVSEGCLLFNKPTHYNNGNYTLVATNQLGTANQTIKGHFLEKPFPESTDNFVSIGDYEVSPTPPITVTHKPEEDTFGVSIAVGLAAFACVLLVVLFIMINKYGRRSKFGMKDVQHIKRRDIVLKRELGEGAFGKVFLAECYNLSPTKDKMLVAVKALKDPTLAARKDFQREAELLTNLQHEHIVKFYGVCGDGDPLIMVFEYMKHGDLNKFLRAHGPDAMILVDGQPRQAKGELGLSQMLHIASQIASGMVYLASQHFVHRDLATRNCLVGANLLVKIGDFGMSRDVYSTDYYREGPHPKGQCSAAWQRQRLARPAAKTVGGHTMLPIRWMPPESIMYRKFTTESDVWSFGVILWEIFTYGKQPWFQLSNTEVIECITQGRVLERPRVCPKEVYDIMLGCWQREPQQRLNIKEIYKILHALGKATPIYLDILG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9S7</accession>
    <name>A0A452H9S7_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="18">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>K(+)(in) = K(+)(out)</text>
            <dbReference type="Rhea" id="RHEA:29463"/>
            <dbReference type="ChEBI" id="CHEBI:29103"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>Li(+)(in) = Li(+)(out)</text>
            <dbReference type="Rhea" id="RHEA:78551"/>
            <dbReference type="ChEBI" id="CHEBI:49713"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="14">
            <text>Na(+)(in) = Na(+)(out)</text>
            <dbReference type="Rhea" id="RHEA:34963"/>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="16">Belongs to the amiloride-sensitive sodium channel (TC 1.A.6) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9S7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000012994.1">
        <property type="protein sequence ID" value="ENSGAGP00000011349.1"/>
        <property type="gene ID" value="ENSGAGG00000008774.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015280">
        <property type="term" value="F:ligand-gated sodium channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.470.10:FF:000006">
        <property type="entry name" value="Acid-sensing ion channel 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.820">
        <property type="entry name" value="Acid-sensing ion channel domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.470.10">
        <property type="entry name" value="Acid-sensing ion channels like domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001873">
        <property type="entry name" value="ENaC"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11690:SF286">
        <property type="entry name" value="ACID-SENSING ION CHANNEL 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11690">
        <property type="entry name" value="AMILORIDE-SENSITIVE SODIUM CHANNEL-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00858">
        <property type="entry name" value="ASC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01078">
        <property type="entry name" value="AMINACHANNEL"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="10" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="12 16" id="KW-0407">Ion channel</keyword>
    <keyword evidence="8 16" id="KW-0406">Ion transport</keyword>
    <keyword evidence="9 17" id="KW-0472">Membrane</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0915">Sodium</keyword>
    <keyword evidence="3 16" id="KW-0894">Sodium channel</keyword>
    <keyword evidence="11 16" id="KW-0739">Sodium transport</keyword>
    <keyword evidence="5 16 17" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 17" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="2 16" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="55"/>
            <end position="73"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="113"/>
            <end position="134"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="452"/>
            <end position="469"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022461"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023053"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023201"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00034430"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00036239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00044635"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="RuleBase" id="RU000679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011349.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="487" mass="55943" checksum="9DC3B33815FA2B1E" modified="2019-05-08" version="1">MVRIHKEKEGNKLYLTRKPLPSLEERKKFDHEFAMSTSFHGLHNIVQTRSKIRKILWMLVVLGSITIVTWQIWNRFSNYFSWPTTTSIIVQYVENIEFPAVTFCNMNRFQVHAVANLSIIFFLWSIVSGVLHAFDTDNKFSQELKDFLQGNQNFSIEEFTRNNGFYLNSSTLLECDFFGQPCYPEDFEHVFTEYGNCFTFNHMDLPARRRVSVSGRGLSLLFDIKQAQFTDDPTFGFIDAGIIYVIHSPKEPPRFDGLGLSTPVGMHAHAAISQLKTVIQEYPWGECNPNIKLQHHKIYSTYGCLQECKARHIQDKCGCLPFLLPGNGTECDLQKFYSCVSPALYNIELMGLCTVGTHNSTCPVPCEETDYPSTVTYSSFASEKALKFFSTKLKRSPEYIRENLVYIDIKYHDLNYKMTRQQKARSVSELLADVGGQLGLFCGASMITIIEIVEYVFTNFCWLCIFLLLKAPEMPQWNIPTPAVWVS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9S9</accession>
    <name>A0A452H9S9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Cullin family profile domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="pathway">
        <text evidence="1">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="similarity">
        <text evidence="2 6 7">Belongs to the cullin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9S9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013000.1">
        <property type="protein sequence ID" value="ENSGAGP00000011355.1"/>
        <property type="gene ID" value="ENSGAGG00000008767.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00143"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031461">
        <property type="term" value="C:cullin-RING ubiquitin ligase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031625">
        <property type="term" value="F:ubiquitin protein ligase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016567">
        <property type="term" value="P:protein ubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006511">
        <property type="term" value="P:ubiquitin-dependent protein catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.230.130:FF:000001">
        <property type="entry name" value="Cullin 4A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.10:FF:000050">
        <property type="entry name" value="Cullin 4B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1310.10:FF:000004">
        <property type="entry name" value="Cullin 4B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1310.10:FF:000008">
        <property type="entry name" value="Cullin 4B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1310.10:FF:000059">
        <property type="entry name" value="Cullin-4B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1310.10">
        <property type="entry name" value="Cullin Repeats"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.230.130">
        <property type="entry name" value="Cullin, Chain C, Domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.10">
        <property type="entry name" value="Winged helix-like DNA-binding domain superfamily/Winged helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045093">
        <property type="entry name" value="Cullin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR059120">
        <property type="entry name" value="Cullin-like_AB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016157">
        <property type="entry name" value="Cullin_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016158">
        <property type="entry name" value="Cullin_homology"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036317">
        <property type="entry name" value="Cullin_homology_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001373">
        <property type="entry name" value="Cullin_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019559">
        <property type="entry name" value="Cullin_neddylation_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016159">
        <property type="entry name" value="Cullin_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036388">
        <property type="entry name" value="WH-like_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036390">
        <property type="entry name" value="WH_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11932">
        <property type="entry name" value="CULLIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00888">
        <property type="entry name" value="Cullin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26557">
        <property type="entry name" value="Cullin_AB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10557">
        <property type="entry name" value="Cullin_Nedd8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00182">
        <property type="entry name" value="CULLIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00884">
        <property type="entry name" value="Cullin_Nedd8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF75632">
        <property type="entry name" value="Cullin homology domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF74788">
        <property type="entry name" value="Cullin repeat-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46785">
        <property type="entry name" value="Winged helix' DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01256">
        <property type="entry name" value="CULLIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50069">
        <property type="entry name" value="CULLIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0832">Ubl conjugation</keyword>
    <keyword evidence="4" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="Cullin family profile" evidence="9">
        <location>
            <begin position="530"/>
            <end position="771"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1"/>
            <end position="95"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="115"/>
            <end position="136"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="62"/>
            <end position="91"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="118"/>
            <end position="135"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006019"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00330"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU003829"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50069"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011355.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="898" mass="102871" checksum="D7D03D8D8F9B087B" modified="2019-05-08" version="1">GPNYILSGDGCDNGCFSSPNPPAAAQEVRPATDGNSSSSSSSCKKRKLNNSSNNSEREEFDSISSCSSSPSKNNSSSSSSIITTSSGGPSSNHHLQKKLRFEDSLDFIGLDVKMAEESSSSSSPAASSQQQHQQQLKNKSLLISSVAVGHHANGLTKAASSTVSSFANSKPGSAKKLVIKNFKDKPKLPENYTDETWQKLKEAVEAIQNSTSIKYNLEELYQAVENLCSYKISANLYKQLRQICEDHIKAQIHHVLFLKKIDKCWQDHCRQMIMIRSIFLFLDRTYVLQNSMLPSIWDMGLELFRTHIISDQKVQNKTIDGILLLIERERNGEAIDRSLLRSLLSMLSDLQIYQDSFEHRFLEETNRLYAAEGQRLMQEREVPEYLHHVNKRLEEEADRIITYLDQSTQKPLIATVEKQLLGEHLTAILQKGLNHLLDENRIQDLSLLYQLFSRVRGGVQVLLQHWIEYIKAFGSTIVINPEKDKTMVQELLDFKDKVDHIIDVCFLKNEKFVNAMKEAFETFINKRPNKPAELIAKYVDSKLRAGNKEATDEELEKMLDKIMIIFRFIYGKDVFEAFYKKDLAKRLLVGKSASVDAEKSMLSKLKHGEYLALPPCFSLLEIIGMIFFKDKMQLENSTTFSICFCFSLQYMQNQNVPGNIELTVNILTMGYWPTYVPMEVHLPPEMVKLQEIFKTFYLGKHSGRKLQWQSTLGHCVLKAEFKEGRKELQVSLFQTLVLLMFNEGEEFSLEEIKQATGIEDGELRRTLQSLACGKARVLTKSPKGKDVEDGDKFTCNDDFRHKLFRIKINQIQMKETVEEQASTTERVFQDRQYQIDAAIVRIMKMRKTLSHNLLVSEVYNQLKFPVKPADLKKRIESLIDRDYMERDKENPNQYNYIA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJQ2</accession>
    <name>A0A452HJQ2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Cytochrome P450 family 4 subfamily B member 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>heme</name>
            <dbReference type="ChEBI" id="CHEBI:30413"/>
        </cofactor>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the cytochrome P450 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJQ2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017276.1">
        <property type="protein sequence ID" value="ENSGAGP00000015123.1"/>
        <property type="gene ID" value="ENSGAGG00000011253.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0020037">
        <property type="term" value="F:heme binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005506">
        <property type="term" value="F:iron ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004497">
        <property type="term" value="F:monooxygenase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016705">
        <property type="term" value="F:oxidoreductase activity, acting on paired donors, with incorporation or reduction of molecular oxygen"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.630.10">
        <property type="entry name" value="Cytochrome P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001128">
        <property type="entry name" value="Cyt_P450"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002402">
        <property type="entry name" value="Cyt_P450_E_grp-II"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036396">
        <property type="entry name" value="Cyt_P450_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050196">
        <property type="entry name" value="Cytochrome_P450_Monoox"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24291">
        <property type="entry name" value="CYTOCHROME P450 FAMILY 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24291:SF201">
        <property type="entry name" value="CYTOCHROME P450, FAMILY 4, SUBFAMILY B, POLYPEPTIDE 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00067">
        <property type="entry name" value="p450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00464">
        <property type="entry name" value="EP450II"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00385">
        <property type="entry name" value="P450"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48264">
        <property type="entry name" value="Cytochrome P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0349">Heme</keyword>
    <keyword evidence="6" id="KW-0408">Iron</keyword>
    <keyword evidence="8" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7" id="KW-0503">Monooxygenase</keyword>
    <keyword evidence="5" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="20"/>
            <end position="40"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="72"/>
            <end position="91"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001971"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023033"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015123.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="11">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="432" mass="49818" checksum="EEB92AF1E34D6003" modified="2019-05-08" version="1">TSELSPVPSTRLFATPSSLLLYPTAGFCLIYVLLKAIQLYRKRQQLLKAFESFPGPPSHWLYGHSSQVGKDIFVFMVCQLLTLYLSSLLNFTDPKGLTIYKFLIPWIGKGLLILHGPKWFQHRRLLTPGFHYDVLKPYVSLMAESTKVMLDKWEWLITVDKSVELFEHVSLMTLDSIMKCAFSCHSNCQTDRDSDYYIKAVYDLTYLVFHRIRFFPFHNDFIYRISSQGRRFQEACRLAHQHTDKVIKERKTSLQDQQELEKIQKKKYLDFLDILLCAKDENGAGLSDEDLRAEVDTFMFEGHDTTASGVSWLLYCMALHPEHQQRCREEIKEILGDRETIQWDDLGKMTYSTMCIKESLRLYPPVPGVSRLLSKPITFHDGRTVPEGLFSLFLLKGPILPSAAYMPLPLKSHFGPECSTGGGRERISLCFP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJQ5</accession>
    <name>A0A452HJQ5_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="14">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Golgi apparatus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 13">Belongs to the small GTPase superfamily. Arf family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJQ5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017302.1">
        <property type="protein sequence ID" value="ENSGAGP00000015145.1"/>
        <property type="gene ID" value="ENSGAGG00000011435.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005525">
        <property type="term" value="F:GTP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003924">
        <property type="term" value="F:GTPase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016192">
        <property type="term" value="P:vesicle-mediated transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:003500">
        <property type="entry name" value="ADP-ribosylation factor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005225">
        <property type="entry name" value="Small_GTP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024156">
        <property type="entry name" value="Small_GTPase_ARF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006689">
        <property type="entry name" value="Small_GTPase_ARF/SAR"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00231">
        <property type="entry name" value="small_GTP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11711">
        <property type="entry name" value="ADP RIBOSYLATION FACTOR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00025">
        <property type="entry name" value="Arf"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00328">
        <property type="entry name" value="SAR1GTPBP"/>
    </dbReference>
    <dbReference type="SMART" id="SM00177">
        <property type="entry name" value="ARF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51417">
        <property type="entry name" value="ARF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0931">ER-Golgi transport</keyword>
    <keyword evidence="8" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="9 11 13" id="KW-0342">GTP-binding</keyword>
    <keyword evidence="10" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="12" id="KW-0460">Magnesium</keyword>
    <keyword evidence="12" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="4" id="KW-0519">Myristate</keyword>
    <keyword evidence="5 11 13" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="7" id="KW-0653">Protein transport</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <feature type="binding site" evidence="11">
        <location>
            <begin position="113"/>
            <end position="120"/>
        </location>
        <ligand>
            <name>GTP</name>
            <dbReference type="ChEBI" id="CHEBI:37565"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="12">
        <location>
            <position position="120"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="12">
        <location>
            <position position="137"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="11">
        <location>
            <position position="159"/>
        </location>
        <ligand>
            <name>GTP</name>
            <dbReference type="ChEBI" id="CHEBI:37565"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004555"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010290"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022707"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022892"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PIRSR" id="PIRSR606689-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PIRSR" id="PIRSR606689-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="RuleBase" id="RU003925"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015145.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="207" mass="22353" checksum="A76995AFE04D4BA2" modified="2019-05-08" version="1">MRNDVVVGPRSDARRSRKALVAAAGWGGGVEGGSAYLPFWALSFGSGPAVAAAFKSPAPAGGVRALLCRGDARSVTSSGRGTGPSQPSLAMGLIFAKLWSFFCDQEHKVIIVGLDNAGKTTILYQFLMNEVVHTSPTIGSNVEEIVVKNTHFLMWDIGGQESLRSSWNTYYSNTEFIILVVDSIDRERLSITKEELYKMLAHEIVGV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJQ9</accession>
    <name>A0A452HJQ9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">KRAB domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HJQ9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017304.1">
        <property type="protein sequence ID" value="ENSGAGP00000015147.1"/>
        <property type="gene ID" value="ENSGAGG00000011437.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006355">
        <property type="term" value="P:regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd07765">
        <property type="entry name" value="KRAB_A-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001909">
        <property type="entry name" value="KRAB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036051">
        <property type="entry name" value="KRAB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050169">
        <property type="entry name" value="Krueppel_C2H2_ZnF"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23232">
        <property type="entry name" value="KRAB DOMAIN C2H2 ZINC FINGER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23232:SF163">
        <property type="entry name" value="ZINC FINGER PROTEIN 589"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01352">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00349">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109640">
        <property type="entry name" value="KRAB domain (Kruppel-associated box)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50805">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="KRAB" evidence="1">
        <location>
            <begin position="1"/>
            <end position="79"/>
        </location>
    </feature>
    <evidence type="ECO:0000259" key="1">
        <source>
            <dbReference type="PROSITE" id="PS50805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015147.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="105" mass="12083" checksum="6D23A9081ABC6A3D" modified="2019-05-08" version="1">MESLEVAVYFTREEWALLDPAQRALYRAIMQENYENVTSLGKDSFKVHATSTRPPLHCPLSASPQYANLSPLPPSSLPKALPYPYSFFCPFLNLFQFHYIFFEMG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJR0</accession>
    <name>A0A452HJR0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Argininosuccinate lyase</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="2">Homotetramer.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the lyase 1 family. Argininosuccinate lyase subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJR0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017308.1">
        <property type="protein sequence ID" value="ENSGAGP00000015151.1"/>
        <property type="gene ID" value="ENSGAGG00000011438.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004056">
        <property type="term" value="F:argininosuccinate lyase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005212">
        <property type="term" value="F:structural constituent of eye lens"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042450">
        <property type="term" value="P:L-arginine biosynthetic process via ornithine"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd01359">
        <property type="entry name" value="Argininosuccinate_lyase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.40.30:FF:000001">
        <property type="entry name" value="Argininosuccinate lyase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.200.10:FF:000015">
        <property type="entry name" value="argininosuccinate lyase isoform X2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.40.30">
        <property type="entry name" value="Fumarase/aspartase (C-terminal domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.200.10">
        <property type="entry name" value="Fumarase/aspartase (Central domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.275.10">
        <property type="entry name" value="Fumarase/aspartase (N-terminal domain)"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="HAMAP" id="MF_00006">
        <property type="entry name" value="Arg_succ_lyase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029419">
        <property type="entry name" value="Arg_succ_lyase_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009049">
        <property type="entry name" value="Argininosuccinate_lyase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024083">
        <property type="entry name" value="Fumarase/histidase_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020557">
        <property type="entry name" value="Fumarate_lyase_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000362">
        <property type="entry name" value="Fumarate_lyase_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022761">
        <property type="entry name" value="Fumarate_lyase_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008948">
        <property type="entry name" value="L-Aspartase-like"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00838">
        <property type="entry name" value="argH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43814">
        <property type="entry name" value="ARGININOSUCCINATE LYASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43814:SF1">
        <property type="entry name" value="ARGININOSUCCINATE LYASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14698">
        <property type="entry name" value="ASL_C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00206">
        <property type="entry name" value="Lyase_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00145">
        <property type="entry name" value="ARGSUCLYASE"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00149">
        <property type="entry name" value="FUMRATELYASE"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48557">
        <property type="entry name" value="L-aspartase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00163">
        <property type="entry name" value="FUMARATE_LYASES"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0273">Eye lens protein</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Fumarate lyase N-terminal" evidence="4">
        <location>
            <begin position="70"/>
            <end position="277"/>
        </location>
    </feature>
    <feature type="domain" description="Argininosuccinate lyase C-terminal" evidence="5">
        <location>
            <begin position="340"/>
            <end position="407"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00010755"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00011881"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="Pfam" id="PF00206"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="Pfam" id="PF14698"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015151.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="8">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="438" mass="48501" checksum="A9255F8A53BB9A0A" modified="2019-05-08" version="1">MASEGDKLWGGRFVGGTDPIMEALNSSMAYDQRLSEVDIRGSMAYAKALEKSGILTKGDLEKILSGLEKELIGDVAGKLHTGRSRNDQVVTDLRLYMRNSCSVLSAHLLQFIKTLVERAAAETDVIFPGYTHLQRAQPIRWSHFLLSHAVAFTRDSERLGEVRRRINVLPLGSGAVAGNPLGIDREFLRKELDFASISLNSMDATSERDFVVEFLFFASLLMIHLSKMAEDLIIYSTKEFGFVTLSDAYSTGSSLMPQKKNPDSLELIRSKAGRVFGRCSGMMMILKGLPSTYNKDLQEDKEAVFDVIDTLSSVLQVATGVISTLQIHKENMEKALSPDMLATDLALYLVRKGMPFRQAHAASGKAVHLAETKGIAINQLTVEDMKSISPLFGTDVSQVWDYANSVEQYTTPGGTSKSSVMVQIEQLRELLRRQKDLF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJR1</accession>
    <name>A0A452HJR1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Pecanex-like protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 6">Membrane</location>
            <topology evidence="1 6">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 6">Belongs to the pecanex family.</text>
    </comment>
    <comment type="caution">
        <text evidence="6">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000017307.1">
        <property type="protein sequence ID" value="ENSGAGP00000015150.1"/>
        <property type="gene ID" value="ENSGAGG00000011428.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039797">
        <property type="entry name" value="Pecanex"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007735">
        <property type="entry name" value="Pecanex_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12372">
        <property type="entry name" value="PECANEX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12372:SF2">
        <property type="entry name" value="PECANEX-LIKE PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05041">
        <property type="entry name" value="Pecanex_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 6" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 6" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 6" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="812"/>
            <end position="830"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="837"/>
            <end position="854"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="874"/>
            <end position="890"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="924"/>
            <end position="949"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="961"/>
            <end position="985"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="997"/>
            <end position="1017"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="1071"/>
            <end position="1093"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="1099"/>
            <end position="1116"/>
        </location>
    </feature>
    <feature type="domain" description="Pecanex C-terminal" evidence="8">
        <location>
            <begin position="1587"/>
            <end position="1813"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1"/>
            <end position="48"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="168"/>
            <end position="189"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="246"/>
            <end position="585"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="640"/>
            <end position="675"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="690"/>
            <end position="712"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="1855"/>
            <end position="1913"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="8"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="7">
        <location>
            <begin position="170"/>
            <end position="180"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="268"/>
            <end position="279"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="292"/>
            <end position="308"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="312"/>
            <end position="332"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="343"/>
            <end position="353"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="364"/>
            <end position="376"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="7">
        <location>
            <begin position="426"/>
            <end position="442"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="7">
        <location>
            <begin position="452"/>
            <end position="467"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="503"/>
            <end position="533"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="664"/>
            <end position="674"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="1865"/>
            <end position="1880"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="RuleBase" id="RU367089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF05041"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015150.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2145" mass="237690" checksum="C52EA3960A9A4CA6" modified="2019-05-08" version="1">MDRGAKTEQGNTSTRRTDSNGPSDPGGGIEMSEFIREATPPVGCSSRNSYAALDANNQIGSGLARLGTAATIKGDTDTAKTSDDISLSLGRSCSLCKEGSEEQDLAADQKILHLVSNDSFVSIQPWLSSGQDLPRDNSDKVCLPNNHHVDQSLSSACDTEMASLVPLHSHSYRKDHRPRGVPRTSSSAVAFPDASLNDFPLYQQRRGLDPVSELEASKPHCGSKESLAGNSCISGVFQLDDILKSSSPQPLAKSGKSKSLKAEKSVDSLRSLSGRSSGSTESYCSGIDRDTNSTISSYKSEQTSSTHVENVLSEHEESPREESETSKRKDCCIDSEDDSSSATDKRSSSERTAVEMSTNSGVQETKDSNTSDEIHSQRGLSASASEEANKNPHANELTTQVDRPPGKAAETRDDQSEKLAALVDSRTCKETGGKQKEGDVRPKSSSLIHRTASVHKSSRRRTGKKRASSFDSSRHREYISFRGVSGTKPQSAVFCHDEDSSDQSDLSRTSSMQSAHQFSSDSSSSTTSHSCQSPEGKYNALKTKYIKERGGADSNTHKAHLGSEGISKKRSTRRTSSTHSVKSRARVLSLDSGTVACLNNPNSLMAPGNIKQLTTSKSDLEAKEGEVLDELSLLGRASHLESVTRSRNSLPSQAMFPEGEEQESASGAAQASEETVMFHRERSTFRRQAVRRRHNAGSNPTPPTLLIGSPLSDTDSHISSSTSVRFYPHDLLSLPQIRLNRLLTIDTDLLEHQDVDLSPDLQDRMQSQEEAAQKVKQYYRFWILPKLWIGINFDRLTLLALFDRNREILENVLAVILAILVAFLGSVLLIEGFFKDIWVFQFCLVIASCQYSLLKSVQPDSSSPRHGHNRIIAYSRPVYFCLCCGLIWLLDYGSRNTSTTRFRLYGVVFTNPLLLLSARDLVIVFTLCFPIVFFIGLLPQVNTFVMYLCEQLDIHVFGGNATTSLLAALYSFICSIVAVALLYGLCYGALKDSWDGQHIPVLFSIFCGLLVAISYHLSRQSSDPSVLFSLVQSKFFPNSEDKNPEDPLSEVKDPLPEKLRNSVNERLQSDLIVCLVIGVLYFAIHVSTVFTVLQPVLSYILYALVGSVGFLTHYVLPQLRKQLPWHCFSHSLLKTKEYYQFEVRNAAHVMWFEKLHVWLLFVEKNVIYPLIVLNELSSSAKNIASPKKLDTEIGALMITIAGLKLLRSSFSNPTCQYVTVIFTVLFFTFDYKSFSETMLLDLFFMSILFSKLWELFYKLRFVYTYIAPWQITWGSAFHAFAQPFAVPHSAMLFVQAAVSALFSTPLNPFLGSAIFITSYVRPVKFWERDYNTKRVDHSNTRLASQLDRNPGSDDNNLNSIFYEHLTRSLQHSLCGDLLLGRWGNYSTGDCFILASDYLNALVHLIEIGNGLVTFQLRGLEFRGTYCQQREVEAITEGVEEDEGFCCCEPGHIPHMLSFNATFGQRWLAWEVLVTKYVLEGYSITDNSAASMLQVFDLRKILTTYYVKGIIYYVTTSPKLEEWLANETMQEGLQPCADCNYVDVDPTFNPNIDEDYDHHLAGVSRESFCATYLNWIEYCCSRREKPLDADKDSSLVTLCYGLCVLGRRALGTASHHMSSNLESFLYGLHALFKGDFRISSVRDEWIFADMELLRKVVVPGIRMSLKLHQDHFTSPDEYDDPTVLYEAIVSHEQNLVIAHEGDPAWRSAVLSNSPSLLALRHVMDDGTNEYKIIMLNKRYLSFRVIKVNKECVRGLWAGQQQELVFLRNRNPERGSIQNAKQALRNMINSSCDQPIGYPIYVSPLTTSYSDSHDQLKEFLGGAISLGNIKNFIVSTWHRLRKGCGAGCNSGGNIEDSDAGGGASCTSNSAAANESQSSMSQAGGNLITGQGPGVGLHPPVTSHPSTLGTSHSSHSVQSSLVRHSPARASVASHSSYCYSSRHSSLRTSTTGFVPCRRSSTSQISLRNLPSSIQSRLSMVNQIEPISQTGIVIMQHGLPSSSSSSQSIPACKQHALVGFLGTEGGNNVTDAQSGGNASPANHMQAKRDEVIYRIQIMDPSQVLEGINMSKRKELQWPDEVIRLKAGRNSWKDWSPQEGMEGHVIHRWVPCSRDPGSRSHIDKTILLVQIEDKYVAVFETGVLELGAEV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJS2</accession>
    <name>A0A452HJS2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Interleukin-2 receptor subunit alpha</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">Receptor for interleukin-2. The receptor is involved in the regulation of immune tolerance by controlling regulatory T cells (TREGs) activity. TREGs suppress the activation and expansion of autoreactive T-cells.</text>
    </comment>
    <comment type="subunit">
        <text evidence="13">Non-covalent dimer of an alpha and a beta subunit. IL2R exists in 3 different forms: a high affinity dimer, an intermediate affinity monomer (beta subunit), and a low affinity monomer (alpha subunit). The high and intermediate affinity forms also associate with a gamma subunit.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Membrane</location>
            <topology evidence="2">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="14">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJS2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017317.1">
        <property type="protein sequence ID" value="ENSGAGP00000015160.1"/>
        <property type="gene ID" value="ENSGAGG00000011448.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019976">
        <property type="term" value="F:interleukin-2 binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004911">
        <property type="term" value="F:interleukin-2 receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002376">
        <property type="term" value="P:immune system process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006954">
        <property type="term" value="P:inflammatory response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00033">
        <property type="entry name" value="CCP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.70.10">
        <property type="entry name" value="Complement Module, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015486">
        <property type="entry name" value="IL-2_rcpt_alpha"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035976">
        <property type="entry name" value="Sushi/SCR/CCP_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000436">
        <property type="entry name" value="Sushi_SCR_CCP_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10573">
        <property type="entry name" value="INTERLEUKIN-2 RECEPTOR ALPHA CHAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10573:SF0">
        <property type="entry name" value="INTERLEUKIN-2 RECEPTOR SUBUNIT ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00084">
        <property type="entry name" value="Sushi"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00032">
        <property type="entry name" value="CCP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57535">
        <property type="entry name" value="Complement control module/SCR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50923">
        <property type="entry name" value="SUSHI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="10" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7" id="KW-0391">Immunity</keyword>
    <keyword evidence="9 16" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-0675">Receptor</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5 17" id="KW-0732">Signal</keyword>
    <keyword evidence="14" id="KW-0768">Sushi</keyword>
    <keyword evidence="4 16" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 16" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="17">
        <location>
            <begin position="1"/>
            <end position="17"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019172820" description="Interleukin-2 receptor subunit alpha" evidence="17">
        <location>
            <begin position="18"/>
            <end position="165"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="137"/>
            <end position="161"/>
        </location>
    </feature>
    <feature type="domain" description="Sushi" evidence="18">
        <location>
            <begin position="49"/>
            <end position="111"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="18"/>
            <end position="48"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00002381"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00013445"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022859"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00025938"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00302"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50923"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015160.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="165" mass="18227" checksum="AD4A7DD4B750CFF1" modified="2019-05-08" version="1">MELIFLFLWGLLRSGEAAGHGNGDSRATSQQSERKHENPVTIVPSDTAGYCGIPSPVEHTTPRVTEYAVGQELHYKCLHGYDARPPTSDISTCKEEHGKTFWTRLSLLCTNDSKSGEEMTQPIPITDTSEPSHVPSVMFPVTVAVSTVFPIIFIILGWIIVSKLW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJS3</accession>
    <name>A0A452HJS3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Leptin receptor overlapping transcript-like 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="6">Negatively regulates growth hormone (GH) receptor cell surface expression in liver. May play a role in liver resistance to GH during periods of reduced nutrient availability.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the OB-RGRP/VPS55 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJS3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017319.1">
        <property type="protein sequence ID" value="ENSGAGP00000015162.1"/>
        <property type="gene ID" value="ENSGAGG00000011449.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005768">
        <property type="term" value="C:endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032511">
        <property type="term" value="P:late endosome to vacuole transport via multivesicular body sorting pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060400">
        <property type="term" value="P:negative regulation of growth hormone receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007262">
        <property type="entry name" value="Vps55/LEPROT"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12050:SF4">
        <property type="entry name" value="LEPTIN RECEPTOR OVERLAPPING TRANSCRIPT-LIKE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12050">
        <property type="entry name" value="LEPTIN RECEPTOR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04133">
        <property type="entry name" value="Vps55"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51257">
        <property type="entry name" value="PROKAR_LIPOPROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 8" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 8" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 8" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="7"/>
            <end position="29"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="35"/>
            <end position="57"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="69"/>
            <end position="89"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="95"/>
            <end position="121"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005645"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00037298"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00040108"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015162.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="132" mass="14489" checksum="B5181A9576B4AA13" modified="2019-05-08" version="1">MAGIKALISLSFGGAIGLMFLMLGCALPQYNSQYWPLFVLFFYILSPIPYCIARRLVDETDATSNACKELAIFLTTGIVVSAFGLPIVFARAQLILWGACALVLTGNTVIFATILGFFLVFGSNDDFSWQQW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJS5</accession>
    <name>A0A452HJS5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Latent-transforming growth factor beta-binding protein 1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="19">Latent-transforming growth factor beta-binding protein 2</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="20">Transforming growth factor beta-1-binding protein 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="25 26">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="26" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="25" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="25" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">Key regulator of transforming growth factor beta (TGFB1, TGFB2 and TGFB3) that controls TGF-beta activation by maintaining it in a latent state during storage in extracellular space. Associates specifically via disulfide bonds with the Latency-associated peptide (LAP), which is the regulatory chain of TGF-beta, and regulates integrin-dependent activation of TGF-beta. Outcompeted by LRRC32/GARP for binding to LAP regulatory chain of TGF-beta.</text>
    </comment>
    <comment type="function">
        <text evidence="14">May play an integral structural role in elastic-fiber architectural organization and/or assembly.</text>
    </comment>
    <comment type="subunit">
        <text evidence="16">Forms part of the large latent transforming growth factor beta precursor complex; removal is essential for activation of complex. Interacts with SDC4. Interacts (via C-terminal domain) with FBN1 (via N-terminal domain) in a Ca(+2)-dependent manner.</text>
    </comment>
    <comment type="subunit">
        <text evidence="17">Interacts with TGFB1; associates via disulfide bonds with the Latency-associated peptide chain (LAP) regulatory chain of TGFB1, leading to regulate activation of TGF-beta-1. LTBP1 does not bind directly to TGF-beta-1, the active chain of TGFB1. Interacts (via C-terminal domain) with FBN1 (via N-terminal domain). Interacts with FBN2. Interacts with ADAMTSL2. Interacts with EFEMP2.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="13">Belongs to the LTBP family.</text>
    </comment>
    <comment type="caution">
        <text evidence="21">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000017325.1">
        <property type="protein sequence ID" value="ENSGAGP00000015168.1"/>
        <property type="gene ID" value="ENSGAGG00000008780.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019838">
        <property type="term" value="F:growth factor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00054">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="12"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000005">
        <property type="entry name" value="Fibrillin 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000194">
        <property type="entry name" value="Latent transforming growth factor beta binding protein 2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000469">
        <property type="entry name" value="Latent transforming growth factor beta binding protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.290.10:FF:000004">
        <property type="entry name" value="latent-transforming growth factor beta-binding protein 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000019">
        <property type="entry name" value="latent-transforming growth factor beta-binding protein 1 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000046">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 1 isoform x2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000014">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000077">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 3 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.290.10:FF:000001">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 3 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.290.10:FF:000002">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 3 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000056">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 3 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000115">
        <property type="entry name" value="latent-transforming growth factor beta-binding protein 4 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000024">
        <property type="entry name" value="Putative latent-transforming growth factor beta-binding protein 2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000273">
        <property type="entry name" value="Putative latent-transforming growth factor beta-binding protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="18"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.290.10">
        <property type="entry name" value="TGF-beta binding (TB) domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026823">
        <property type="entry name" value="cEGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001881">
        <property type="entry name" value="EGF-like_Ca-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013032">
        <property type="entry name" value="EGF-like_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000152">
        <property type="entry name" value="EGF-type_Asp/Asn_hydroxyl_site"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018097">
        <property type="entry name" value="EGF_Ca-bd_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051145">
        <property type="entry name" value="GAS-SHBG-PROS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009030">
        <property type="entry name" value="Growth_fac_rcpt_cys_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049883">
        <property type="entry name" value="NOTCH1_EGF-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017878">
        <property type="entry name" value="TB_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036773">
        <property type="entry name" value="TB_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24040:SF16">
        <property type="entry name" value="FIBRILLIN-2-LIKE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24040">
        <property type="entry name" value="LAMININ G-LIKE DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12662">
        <property type="entry name" value="cEGF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07645">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12661">
        <property type="entry name" value="hEGF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00683">
        <property type="entry name" value="TB"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="19"/>
    </dbReference>
    <dbReference type="SMART" id="SM00179">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="17"/>
    </dbReference>
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        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="7"/>
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        <property type="entry name" value="Growth factor receptor domain"/>
        <property type="match status" value="4"/>
    </dbReference>
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        <property type="entry name" value="TB module/8-cys domain"/>
        <property type="match status" value="4"/>
    </dbReference>
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        <property type="entry name" value="ASX_HYDROXYL"/>
        <property type="match status" value="10"/>
    </dbReference>
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        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="2"/>
    </dbReference>
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        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="8"/>
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    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9 21" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="4 21" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="3" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
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    <keyword evidence="6" id="KW-0358">Heparin-binding</keyword>
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    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="26" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
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            <begin position="1156"/>
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            <begin position="1255"/>
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            <dbReference type="SAM" id="MobiDB-lite"/>
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    <sequence length="1677" mass="181196" checksum="29D166A0BAEFBE34" modified="2019-05-08" version="1">KLSAPIHRSQSGASSQPCSPAGAGQARDPTPQRARLKCLEDQIQPLGWSHVSLFTPFHVPPAVCDPPCQNKGSCSRPQLCICRSGFQGAQCEEVVPERVYHPPGSVAALQPPAGSLPKRAGSAEREGSPRGAQTPILKQALAGTQGRIVFPVSRHTGSSRTVRRYPAGNTQLMSNALPNGNGHEQSSSGPHAAYQEHTLSLWGANLTEKIRKIKIVFTPTICKQTCRSGHCYNSCQKGDTTTLYSQGGQNHDPKSGFRIYFCQIPCLNGGRCVGRDECWCPSNSTGKFCHLPVPKLEKNQVGRVPKTLASGSIKQSMYTLPLSNQLASLNPSLVNVHIKHPPEATVQIHQVARVRGDLGQSEENSVEAVLLSQLPTGPQHSYSNGNSNSITTESGQQQRPPELMGRCFRETLHGQCANPLEGLAKFEDCCGSVGLFWGVNRCIPCPPRPAHPVIENRQVECPQGYKRLNRSHCQDINECLMLGLCKDTECVNTRGSFLCTCKPGTMLDPSRSRCVSDKAVSMEQGLCYRSAAEGVCALPLAQSITKQICCCSRVGKGWGRNCEKCPMLGSETFKEICPAGHGYTYSSSDIRLSMRKAEAEELPFSSEEQGESSDQTPTQAAKRLQLQEALSGGVTGTLARAGTSAAEGVLGCDATLVAGTADATQEPAPHPGGNPCTICWGTGEQWPVTSPCLSLSEMDRCASVPDPCGPGTCVNLPDRYSCICGPGYQLHPGLPQCIDDDECMRDPCAGKGHCINSVGSYSCLCYPGYTLLASQGTQTCQDLNECEQPSMCQGGQCTNTPGSYHCECARGYIMGHTGQCEDIDECADPSTCPRGRCVNALGSYECLSCAAGYQPMNGRCVDDDECFTVGICAHGLCINLAGSFRCSCYHGYEVTADEKSCQDINECATHNACPMGLCINSEGSFSCAACGAGYTVSRDGSMCEDIDECSSATACPLGICTNTDGSFACRACDAGYMLSSNRFTCEDMNECLASEICSPNGECLNSHGSYFCICAPGFSNAAGGVSCQGEQLGQMAAAKCALDPGYLFIVQGPVYTHRTASVLIFILSLPDVDECADKSRCLRGQCFNTEGSYRCLCENGFRHSQETDDCVDVDECVDFGDTVCGTWRCQNTPGSYRCVMGCQPGFHRTPLGDCIDIDECANETLCGSHAFCDNTDGSFRCLCDSGYENLPPGQDCVDVNECELMVAVCGTALCENVEGSFLCLCLSDHEEYNTEAGECRPRAGMAPRPAGSERKECYYHISDVQLCDSVLAKNTTKEECCCTEGAAWGDNCETHPCPILVEYSEICPSGKGYIPVEGTLLFGQTSYTDADECEMFGPEICRNGHCVNTVPGYKCFCRSGYLYDSSRLECVDQDECENEMSCVNGECLNTAGSFHCFCGLPLVLDVTRNRCINVSSRAEDPEEHEIHLDICWQRVSNYICNQPLRGRRTTYTECCCRYGEAWSQDCALCPHRASEDFAHLCNVARGETEREPGLQERPGYEYGPALEDSHYGLYSPDLGPYYNYLSPEYGAPDASFPQRELSSEVGGSTFLRTPQLPPAQSQPRYLPKSNPPPPSPKGLQAEECGILNGCENGRCVRVREGYTCDCFDGFQLDMTHMACVDINECEEVSSSGPLCEGGTCENTEGSYRCNCSPGFVVVAGHCRAKETPVLMEDKWRC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJS6</accession>
    <name>A0A452HJS6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">SAM-dependent MTase RsmB/NOP-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1 6">Belongs to the class I-like SAM-binding methyltransferase superfamily. RsmB/NOP family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJS6"/>
    <dbReference type="SMR" id="A0A452HJS6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017322.1">
        <property type="protein sequence ID" value="ENSGAGP00000015165.1"/>
        <property type="gene ID" value="ENSGAGG00000011440.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008173">
        <property type="term" value="F:RNA methyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001510">
        <property type="term" value="P:RNA methylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd02440">
        <property type="entry name" value="AdoMet_MTases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd21150">
        <property type="entry name" value="PUA_NSun6-like"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="PUA domain"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Vaccinia Virus protein VP39"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="MeTrfase_RsmB-F_NOP2_cat"/>
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    <dbReference type="InterPro" id="IPR001678">
        <property type="entry name" value="MeTrfase_RsmB-F_NOP2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002478">
        <property type="entry name" value="PUA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015947">
        <property type="entry name" value="PUA-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036974">
        <property type="entry name" value="PUA_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023267">
        <property type="entry name" value="RCMT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018314">
        <property type="entry name" value="RsmB/NOL1/NOP2-like_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029063">
        <property type="entry name" value="SAM-dependent_MTases_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22807">
        <property type="entry name" value="NOP2 YEAST -RELATED NOL1/NOP2/FMU SUN DOMAIN-CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22807:SF34">
        <property type="entry name" value="TRNA (CYTOSINE(72)-C(5))-METHYLTRANSFERASE NSUN6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01189">
        <property type="entry name" value="Methyltr_RsmB-F"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR02008">
        <property type="entry name" value="RCMTFAMILY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00359">
        <property type="entry name" value="PUA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF88697">
        <property type="entry name" value="PUA domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53335">
        <property type="entry name" value="S-adenosyl-L-methionine-dependent methyltransferases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01153">
        <property type="entry name" value="NOL1_NOP2_SUN"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="PUA"/>
        <property type="match status" value="1"/>
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    <dbReference type="PROSITE" id="PS51686">
        <property type="entry name" value="SAM_MT_RSMB_NOP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 6" id="KW-0489">Methyltransferase</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 6" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="4 6" id="KW-0949">S-adenosyl-L-methionine</keyword>
    <keyword evidence="3 6" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="SAM-dependent MTase RsmB/NOP-type" evidence="7">
        <location>
            <begin position="140"/>
            <end position="465"/>
        </location>
    </feature>
    <feature type="active site" description="Nucleophile" evidence="6">
        <location>
            <position position="372"/>
        </location>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <begin position="241"/>
            <end position="247"/>
        </location>
        <ligand>
            <name>S-adenosyl-L-methionine</name>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="265"/>
        </location>
        <ligand>
            <name>S-adenosyl-L-methionine</name>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="292"/>
        </location>
        <ligand>
            <name>S-adenosyl-L-methionine</name>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="322"/>
        </location>
        <ligand>
            <name>S-adenosyl-L-methionine</name>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007494"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022603"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022691"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS51686"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015165.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="468" mass="51244" checksum="A16A93D8EB88D61B" modified="2019-05-08" version="1">MYSVHTVILLSGDLSSTSNVFRFDLNCALGTLEAERKYESLLNCLSHPPAFTTVRVNTHLASVKQVKNLLFEEIHKQFKDLSVPVLQHPELQDVLLIPVIGPRKDLQKQSSEVIVGAQCGNSVLRGAHVFVPGIISASKFMKAGDVVSVYSDIEGKCKRGAKVFLGTKVFIGNGISELSRSEIFSSSGSLKGIGIRMTDPVYLSPSLDNVLSSYLFLQNLPSAVVSHVLNPQPGERILDMCAAPGGKTTHLATLMHDQGEVIAMDKIANKVKKIKQNAALLQLNCVKAFCFDGTKVLATGKKEDGQEGPPFSAESFDRILLDVPCSGMGQRPNMAYFWTLKEVMSYQPLQRKLFSTAVKLLKPGGTLVYSTCTITLSENEEQVAWALETFPCLQLQSQEPHIGGEGMMGAGLSLDQLKLLQRFDPSSVTSREMDINSLQDSREDDLILLANKDCIGFFIAKFIKLNSK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJS8</accession>
    <name>A0A452HJS8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Protein max</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="10">Myc-associated factor X</fullName>
        </alternativeName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the MAX family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJS8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017324.1">
        <property type="protein sequence ID" value="ENSGAGP00000015167.1"/>
        <property type="gene ID" value="ENSGAGG00000011450.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090575">
        <property type="term" value="C:RNA polymerase II transcription regulator complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003700">
        <property type="term" value="F:DNA-binding transcription factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046983">
        <property type="term" value="F:protein dimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045944">
        <property type="term" value="P:positive regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd11406">
        <property type="entry name" value="bHLHzip_Max"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.280.10:FF:000023">
        <property type="entry name" value="MAX isoform 13"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.280.10">
        <property type="entry name" value="Helix-loop-helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011598">
        <property type="entry name" value="bHLH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036638">
        <property type="entry name" value="HLH_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10328:SF3">
        <property type="entry name" value="PROTEIN MAX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10328">
        <property type="entry name" value="PROTEIN MAX MYC-ASSOCIATED FACTOR X"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00010">
        <property type="entry name" value="HLH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00353">
        <property type="entry name" value="HLH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47459">
        <property type="entry name" value="HLH, helix-loop-helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50888">
        <property type="entry name" value="BHLH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0010">Activator</keyword>
    <keyword evidence="6" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="9" id="KW-0539">Nucleus</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0678">Repressor</keyword>
    <keyword evidence="8" id="KW-0804">Transcription</keyword>
    <keyword evidence="5" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="BHLH" evidence="12">
        <location>
            <begin position="20"/>
            <end position="71"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="89"/>
            <end position="116"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007628"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00017633"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022491"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00029944"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50888"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015167.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="116" mass="13643" checksum="DB717CDBDD880416" modified="2019-05-08" version="1">MLAWYLASHSEARHDDLTADKRAHHNALERKRRDHIKDSFHSLRDSVPSLQGEKASRAQILDKATEYIQYMRRKNHTHQQDIDDLKRQNALLEQQGDSESEPDEPQSRKKLRMEAS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJT2</accession>
    <name>A0A452HJT2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">DNA-directed RNA polymerase III subunit RPC9</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="13">DNA-directed RNA polymerase III subunit rpc9</fullName>
        </alternativeName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">Accessory protein for the calcitonin gene-related peptide (CGRP) receptor. It modulates CGRP responsiveness in a variety of tissues.</text>
    </comment>
    <comment type="function">
        <text evidence="12">DNA-dependent RNA polymerase catalyzes the transcription of DNA into RNA using the four ribonucleoside triphosphates as substrates. Specific peripheric component of RNA polymerase III (Pol III) which synthesizes small non-coding RNAs including 5S rRNA, snRNAs, tRNAs and miRNAs from at least 500 distinct genomic loci. With POLR3H/RPC8 forms a mobile stalk that protrudes from Pol III core and functions primarily in transcription initiation. Pol III plays a key role in sensing and limiting infection by intracellular bacteria and DNA viruses. Acts as nuclear and cytosolic DNA sensor involved in innate immune response. Can sense non-self dsDNA that serves as template for transcription into dsRNA. The non-self RNA polymerase III transcripts, such as Epstein-Barr virus-encoded RNAs (EBERs) induce type I interferon and NF-kappa-B through the RIG-I pathway.</text>
    </comment>
    <comment type="subunit">
        <text evidence="11">Component of the RNA polymerase III complex consisting of 17 subunits: a ten-subunit horseshoe-shaped catalytic core composed of POLR3A/RPC1, POLR3B/RPC2, POLR1C/RPAC1, POLR1D/RPAC2, POLR3K/RPC10, POLR2E/RPABC1, POLR2F/RPABC2, POLR2H/RPABC3, POLR2K/RPABC4 and POLR2L/RPABC5; a mobile stalk composed of two subunits POLR3H/RPC8 and CRCP/RPC9, protruding from the core and functioning primarily in transcription initiation; and additional subunits homologous to general transcription factors of the RNA polymerase II machinery, POLR3C/RPC3-POLR3F/RPC6-POLR3G/RPC7 heterotrimer required for transcription initiation and POLR3D/RPC4-POLR3E/RPC5 heterodimer involved in both transcription initiation and termination.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Peripheral membrane protein</topology>
            <orientation evidence="2">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the eukaryotic RPC9 RNA polymerase subunit family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJT2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017311.1">
        <property type="protein sequence ID" value="ENSGAGP00000015154.1"/>
        <property type="gene ID" value="ENSGAGG00000011445.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005666">
        <property type="term" value="C:RNA polymerase III complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000166">
        <property type="term" value="F:nucleotide binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006384">
        <property type="term" value="P:transcription initiation at RNA polymerase III promoter"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1250.40:FF:000002">
        <property type="entry name" value="DNA-directed RNA polymerase III subunit RPC9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1250.40">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010997">
        <property type="entry name" value="HRDC-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006590">
        <property type="entry name" value="RNA_pol_Rpb4/RPC9_core"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005574">
        <property type="entry name" value="Rpb4/RPC9"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038324">
        <property type="entry name" value="Rpb4/RPC9_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038846">
        <property type="entry name" value="RPC9"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15561">
        <property type="entry name" value="CALCITONIN GENE-RELATED PEPTIDE-RECEPTOR COMPONENT PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15561:SF0">
        <property type="entry name" value="DNA-DIRECTED RNA POLYMERASE III SUBUNIT RPC9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03874">
        <property type="entry name" value="RNA_pol_Rpb4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00657">
        <property type="entry name" value="RPOL4c"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47819">
        <property type="entry name" value="HRDC-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="6" id="KW-0240">DNA-directed RNA polymerase</keyword>
    <keyword evidence="7" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-0539">Nucleus</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0804">Transcription</keyword>
    <feature type="domain" description="RNA polymerase Rpb4/RPC9 core" evidence="15">
        <location>
            <begin position="6"/>
            <end position="132"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="129"/>
            <end position="152"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004413"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00006898"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00016672"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022478"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00043924"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00044007"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00045808"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00073026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="SMART" id="SM00657"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015154.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="152" mass="17406" checksum="EA4812045167A95A" modified="2019-05-08" version="1">MEQTSTERKDANAALLSNFEVFQLLTDLKQQRKDSGKHKQSSGQQNLNTIMYETLKYISKTPCRFQSPEIVRDFLIAMKGHKLTKAEKLQLLNHRPMTAVEIQLMVEESEERLTEEQIESLLQTVSSILPGNPEEELQRQATMAAEEEGDPE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJT3</accession>
    <name>A0A452HJT3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Voltage-dependent L-type calcium channel subunit beta-2</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="15">Calcium channel voltage-dependent subunit beta 2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="16">Beta subunit of voltage-dependent calcium channels which contributes to the function of the calcium channel by increasing peak calcium current. Plays a role in shifting voltage dependencies of activation and inactivation of the channel. May modulate G protein inhibition. May contribute to beta-adrenergic augmentation of Ca(2+) influx in cardiomyocytes, thereby regulating increases in heart rate and contractile force. Involved in membrane targeting of the alpha-1 subunit CACNA1C.</text>
    </comment>
    <comment type="subunit">
        <text evidence="17">Component of a calcium channel complex consisting of a pore-forming alpha subunit (CACNA1S) and the ancillary subunits CACNB1 or CACNB2, CACNG1 and CACNA2D1. The channel complex contains alpha, beta, gamma and delta subunits in a 1:1:1:1 ratio, i.e. it contains either CACNB1 or CACNB2. Interacts with CACNA1C. Interacts with RRAD; interaction may be involved in beta-adrenergic regulation of heart rate and contractile force. Interaction with RRAD regulates the trafficking of CACNA1C to the cell membrane. Interacts with TMIGD2. Interacts with CAMK2D. Interacts with CBARP. Interacts with CAMK2A.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <location evidence="1">Sarcolemma</location>
            <topology evidence="1">Peripheral membrane protein</topology>
            <orientation evidence="1">Cytoplasmic side</orientation>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the calcium channel beta subunit family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJT3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017334.1">
        <property type="protein sequence ID" value="ENSGAGP00000015177.1"/>
        <property type="gene ID" value="ENSGAGG00000011455.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042383">
        <property type="term" value="C:sarcolemma"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005891">
        <property type="term" value="C:voltage-gated calcium channel complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005245">
        <property type="term" value="F:voltage-gated calcium channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.40:FF:000015">
        <property type="entry name" value="Voltage-dependent L-type calcium channel subunit beta-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000023">
        <property type="entry name" value="Voltage-dependent L-type calcium channel subunit beta-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046937">
        <property type="entry name" value="CAB1-4_N_A-dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008145">
        <property type="entry name" value="GK/Ca_channel_bsu"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005444">
        <property type="entry name" value="VDCC_L_b2su"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000584">
        <property type="entry name" value="VDCC_L_bsu"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11824">
        <property type="entry name" value="VOLTAGE-DEPENDENT CALCIUM CHANNEL BETA SUBUNIT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00625">
        <property type="entry name" value="Guanylate_kin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12052">
        <property type="entry name" value="VGCC_beta4Aa_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01626">
        <property type="entry name" value="LCACHANNELB"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01628">
        <property type="entry name" value="LCACHANNELB2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00072">
        <property type="entry name" value="GuKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0106">Calcium</keyword>
    <keyword evidence="9" id="KW-0107">Calcium channel</keyword>
    <keyword evidence="8" id="KW-0109">Calcium transport</keyword>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="14" id="KW-0407">Ion channel</keyword>
    <keyword evidence="12" id="KW-0406">Ion transport</keyword>
    <keyword evidence="13" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 18" id="KW-0728">SH3 domain</keyword>
    <keyword evidence="5" id="KW-0813">Transport</keyword>
    <keyword evidence="11" id="KW-0851">Voltage-gated channel</keyword>
    <feature type="domain" description="SH3" evidence="20">
        <location>
            <begin position="54"/>
            <end position="123"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1"/>
            <end position="39"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="130"/>
            <end position="202"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="427"/>
            <end position="599"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="15"/>
            <end position="25"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="19">
        <location>
            <begin position="137"/>
            <end position="149"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="431"/>
            <end position="441"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="500"/>
            <end position="542"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="550"/>
            <end position="563"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="571"/>
            <end position="599"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004278"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010836"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00019005"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022568"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022673"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022882"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00030521"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00046042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00046619"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015177.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="599" mass="67767" checksum="C92FB614833B102D" modified="2019-05-08" version="1">MYLYGSEDYEVGSADSYTSRPSDSDVSLEEDREAVRREAERQAQAQLEKAKTKPVAFAVRTNVGYSAGHDDDVPVPGMAISFEAKDFLHVKEKFNNDWWIGRLVKEGCEIGFIPSPVKLENMRIQHEQRVKQGKFYSSKSGGNSSSSLGDIVPSSRKSTPPSSAIDIDATGLDAEENDIPANHRSPKPSANSVTSPHSKEKRMPFFKKTEHIPPYDVVPSMRPVVLVGPSLKGYEVTDMMQKALFDFLKHRFEGRISITRVTADISLAKRSVLNNPSKHAIIERSNTRSSLAEVQSEIERIFELARTLQLVVLDADTINHPAQLSKTSLAPIIVYVKISSPKVLQRLIKSRGKSQAKHLNVQMVAADKLAQCPPEMFDVILDENQLEDACEHLADYLEAYWKATHPPCSNPPNQLLTRTLATATLPISPGPNINLQGSQGDQRNDHSAPERTGSQVEEEARIEPMKKFPHRSSSSAHHHNHRSGVSRGLSRQETFDSETQDSRDSAYIEPKEEYSHEHGDHNDSHRDHSHRDESHGSSDHRHRESRHRSKERDREQDHNECNKQRSRHKSRDQYCDKDGEMMSRKRNDTGDWNRDVYIR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJT7</accession>
    <name>A0A452HJT7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">SAM-dependent MTase RsmB/NOP-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1 6">Belongs to the class I-like SAM-binding methyltransferase superfamily. RsmB/NOP family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJT7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017316.1">
        <property type="protein sequence ID" value="ENSGAGP00000015159.1"/>
        <property type="gene ID" value="ENSGAGG00000011440.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008173">
        <property type="term" value="F:RNA methyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001510">
        <property type="term" value="P:RNA methylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd02440">
        <property type="entry name" value="AdoMet_MTases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd21150">
        <property type="entry name" value="PUA_NSun6-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.130.10">
        <property type="entry name" value="PUA domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.150">
        <property type="entry name" value="Vaccinia Virus protein VP39"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049560">
        <property type="entry name" value="MeTrfase_RsmB-F_NOP2_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001678">
        <property type="entry name" value="MeTrfase_RsmB-F_NOP2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002478">
        <property type="entry name" value="PUA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015947">
        <property type="entry name" value="PUA-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036974">
        <property type="entry name" value="PUA_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023267">
        <property type="entry name" value="RCMT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018314">
        <property type="entry name" value="RsmB/NOL1/NOP2-like_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029063">
        <property type="entry name" value="SAM-dependent_MTases_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22807">
        <property type="entry name" value="NOP2 YEAST -RELATED NOL1/NOP2/FMU SUN DOMAIN-CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22807:SF34">
        <property type="entry name" value="TRNA (CYTOSINE(72)-C(5))-METHYLTRANSFERASE NSUN6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01189">
        <property type="entry name" value="Methyltr_RsmB-F"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR02008">
        <property type="entry name" value="RCMTFAMILY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00359">
        <property type="entry name" value="PUA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF88697">
        <property type="entry name" value="PUA domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53335">
        <property type="entry name" value="S-adenosyl-L-methionine-dependent methyltransferases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01153">
        <property type="entry name" value="NOL1_NOP2_SUN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50890">
        <property type="entry name" value="PUA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51686">
        <property type="entry name" value="SAM_MT_RSMB_NOP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 6" id="KW-0489">Methyltransferase</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 6" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="4 6" id="KW-0949">S-adenosyl-L-methionine</keyword>
    <keyword evidence="3 6" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="SAM-dependent MTase RsmB/NOP-type" evidence="7">
        <location>
            <begin position="143"/>
            <end position="468"/>
        </location>
    </feature>
    <feature type="active site" description="Nucleophile" evidence="6">
        <location>
            <position position="375"/>
        </location>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <begin position="244"/>
            <end position="250"/>
        </location>
        <ligand>
            <name>S-adenosyl-L-methionine</name>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="268"/>
        </location>
        <ligand>
            <name>S-adenosyl-L-methionine</name>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="295"/>
        </location>
        <ligand>
            <name>S-adenosyl-L-methionine</name>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="325"/>
        </location>
        <ligand>
            <name>S-adenosyl-L-methionine</name>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007494"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022603"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022691"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS51686"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015159.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="471" mass="51633" checksum="1BE753F8445ED318" modified="2019-05-08" version="1">MTRRASPTTIKIQLRPQPFPGVCARLRLISALGTLEAERKYESLLNCLSHPPAFTTVRVNTHLASVKQVKNLLFEEIHKQFKDLSVPVLQHPELQDVLLIPVIGPRKDLQKQSSEVIVGAQCGNSVLRGAHVFVPGIISASKFMKAGDVVSVYSDIEGKCKRGAKVFLGTKVFIGNGISELSRSEIFSSSGSLKGIGIRMTDPVYLSPSLDNVLSSYLFLQNLPSAVVSHVLNPQPGERILDMCAAPGGKTTHLATLMHDQGEVIAMDKIANKVKKIKQNAALLQLNCVKAFCFDGTKVLATGKKEDGQEGPPFSAESFDRILLDVPCSGMGQRPNMAYFWTLKEVMSYQPLQRKLFSTAVKLLKPGGTLVYSTCTITLSENEEQVAWALETFPCLQLQSQEPHIGGEGMMGAGLSLDQLKLLQRFDPSSVTSREMDINSLQDSREDDLILLANKDCIGFFIAKFIKLNSK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJU1</accession>
    <name>A0A452HJU1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Voltage-dependent L-type calcium channel subunit beta-2</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="15">Calcium channel voltage-dependent subunit beta 2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="16">Beta subunit of voltage-dependent calcium channels which contributes to the function of the calcium channel by increasing peak calcium current. Plays a role in shifting voltage dependencies of activation and inactivation of the channel. May modulate G protein inhibition. May contribute to beta-adrenergic augmentation of Ca(2+) influx in cardiomyocytes, thereby regulating increases in heart rate and contractile force. Involved in membrane targeting of the alpha-1 subunit CACNA1C.</text>
    </comment>
    <comment type="subunit">
        <text evidence="17">Component of a calcium channel complex consisting of a pore-forming alpha subunit (CACNA1S) and the ancillary subunits CACNB1 or CACNB2, CACNG1 and CACNA2D1. The channel complex contains alpha, beta, gamma and delta subunits in a 1:1:1:1 ratio, i.e. it contains either CACNB1 or CACNB2. Interacts with CACNA1C. Interacts with RRAD; interaction may be involved in beta-adrenergic regulation of heart rate and contractile force. Interaction with RRAD regulates the trafficking of CACNA1C to the cell membrane. Interacts with TMIGD2. Interacts with CAMK2D. Interacts with CBARP. Interacts with CAMK2A.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <location evidence="1">Sarcolemma</location>
            <topology evidence="1">Peripheral membrane protein</topology>
            <orientation evidence="1">Cytoplasmic side</orientation>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the calcium channel beta subunit family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJU1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017339.1">
        <property type="protein sequence ID" value="ENSGAGP00000015181.1"/>
        <property type="gene ID" value="ENSGAGG00000011455.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042383">
        <property type="term" value="C:sarcolemma"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005891">
        <property type="term" value="C:voltage-gated calcium channel complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005245">
        <property type="term" value="F:voltage-gated calcium channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.40:FF:000015">
        <property type="entry name" value="Voltage-dependent L-type calcium channel subunit beta-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000023">
        <property type="entry name" value="Voltage-dependent L-type calcium channel subunit beta-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046937">
        <property type="entry name" value="CAB1-4_N_A-dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008145">
        <property type="entry name" value="GK/Ca_channel_bsu"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005444">
        <property type="entry name" value="VDCC_L_b2su"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000584">
        <property type="entry name" value="VDCC_L_bsu"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11824">
        <property type="entry name" value="VOLTAGE-DEPENDENT CALCIUM CHANNEL BETA SUBUNIT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00625">
        <property type="entry name" value="Guanylate_kin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12052">
        <property type="entry name" value="VGCC_beta4Aa_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01626">
        <property type="entry name" value="LCACHANNELB"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01628">
        <property type="entry name" value="LCACHANNELB2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00072">
        <property type="entry name" value="GuKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0106">Calcium</keyword>
    <keyword evidence="9" id="KW-0107">Calcium channel</keyword>
    <keyword evidence="8" id="KW-0109">Calcium transport</keyword>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="14" id="KW-0407">Ion channel</keyword>
    <keyword evidence="12" id="KW-0406">Ion transport</keyword>
    <keyword evidence="13" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 18" id="KW-0728">SH3 domain</keyword>
    <keyword evidence="5" id="KW-0813">Transport</keyword>
    <keyword evidence="11" id="KW-0851">Voltage-gated channel</keyword>
    <feature type="domain" description="SH3" evidence="20">
        <location>
            <begin position="66"/>
            <end position="135"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="24"/>
            <end position="47"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="142"/>
            <end position="214"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="439"/>
            <end position="611"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="27"/>
            <end position="37"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="19">
        <location>
            <begin position="149"/>
            <end position="161"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="443"/>
            <end position="453"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="512"/>
            <end position="554"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="562"/>
            <end position="575"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="583"/>
            <end position="611"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004278"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010836"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00019005"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022568"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022673"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022882"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00030521"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00046042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00046619"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015181.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="611" mass="69058" checksum="6B5A513BE4C2B282" modified="2019-05-08" version="1">MKATWIRLLKRAKGGRLKNSDICGSADSYTSRPSDSDVSLEEDREAVRREAERQAQAQLEKAKTKPVAFAVRTNVGYSAGHDDDVPVPGMAISFEAKDFLHVKEKFNNDWWIGRLVKEGCEIGFIPSPVKLENMRIQHEQRVKQGKFYSSKSGGNSSSSLGDIVPSSRKSTPPSSAIDIDATGLDAEENDIPANHRSPKPSANSVTSPHSKEKRMPFFKKTEHIPPYDVVPSMRPVVLVGPSLKGYEVTDMMQKALFDFLKHRFEGRISITRVTADISLAKRSVLNNPSKHAIIERSNTRSSLAEVQSEIERIFELARTLQLVVLDADTINHPAQLSKTSLAPIIVYVKISSPKVLQRLIKSRGKSQAKHLNVQMVAADKLAQCPPEMFDVILDENQLEDACEHLADYLEAYWKATHPPCSNPPNQLLTRTLATATLPISPGPNINLQGSQGDQRNDHSAPERTGSQVEEEARIEPMKKFPHRSSSSAHHHNHRSGVSRGLSRQETFDSETQDSRDSAYIEPKEEYSHEHGDHNDSHRDHSHRDESHGSSDHRHRESRHRSKERDREQDHNECNKQRSRHKSRDQYCDKDGEMMSRKRNDTGDWNRDVYIR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJU2</accession>
    <name>A0A452HJU2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Interleukin-2 receptor subunit alpha</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">Receptor for interleukin-2. The receptor is involved in the regulation of immune tolerance by controlling regulatory T cells (TREGs) activity. TREGs suppress the activation and expansion of autoreactive T-cells.</text>
    </comment>
    <comment type="subunit">
        <text evidence="13">Non-covalent dimer of an alpha and a beta subunit. IL2R exists in 3 different forms: a high affinity dimer, an intermediate affinity monomer (beta subunit), and a low affinity monomer (alpha subunit). The high and intermediate affinity forms also associate with a gamma subunit.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Membrane</location>
            <topology evidence="2">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJU2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017321.1">
        <property type="protein sequence ID" value="ENSGAGP00000015164.1"/>
        <property type="gene ID" value="ENSGAGG00000011448.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019976">
        <property type="term" value="F:interleukin-2 binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004911">
        <property type="term" value="F:interleukin-2 receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002376">
        <property type="term" value="P:immune system process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006954">
        <property type="term" value="P:inflammatory response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.20.28.230">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.70.10">
        <property type="entry name" value="Complement Module, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015486">
        <property type="entry name" value="IL-2_rcpt_alpha"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035976">
        <property type="entry name" value="Sushi/SCR/CCP_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000436">
        <property type="entry name" value="Sushi_SCR_CCP_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10573">
        <property type="entry name" value="INTERLEUKIN-2 RECEPTOR ALPHA CHAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10573:SF0">
        <property type="entry name" value="INTERLEUKIN-2 RECEPTOR SUBUNIT ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00084">
        <property type="entry name" value="Sushi"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00032">
        <property type="entry name" value="CCP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57535">
        <property type="entry name" value="Complement control module/SCR domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="10" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7" id="KW-0391">Immunity</keyword>
    <keyword evidence="9" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-0675">Receptor</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <keyword evidence="4" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8" id="KW-1133">Transmembrane helix</keyword>
    <feature type="domain" description="Sushi" evidence="14">
        <location>
            <begin position="12"/>
            <end position="74"/>
        </location>
    </feature>
    <feature type="domain" description="Sushi" evidence="14">
        <location>
            <begin position="94"/>
            <end position="150"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00002381"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00013445"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022859"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00025938"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="SMART" id="SM00032"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015164.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="173" mass="19878" checksum="ABE1F27A2DE3D952" modified="2019-05-08" version="1">SASKQTMRWQNLGTMHGKRGGIQTCDPPFAVMELNNKVRYKCEAGYRRIPGESNFIVCINDTGLIHWTTKASVCERNRTNPQLNPARSKHIFYMHSPVEHTTPRVTEYAVGQELHYKCLHGYDARPPTSDISTCKEEHGKTFWTRLSLLCTNDSKSGEEMTTFRYVLHSNQRL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK20</accession>
    <name>A0A452HK20_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">FAD-binding PCMH-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="1 14">
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="15">
            <name>Mo-molybdopterin</name>
            <dbReference type="ChEBI" id="CHEBI:71302"/>
        </cofactor>
        <text evidence="15">Binds 1 Mo-molybdopterin (Mo-MPT) cofactor per subunit.</text>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="12">
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="15">
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:49601"/>
        </cofactor>
        <text evidence="15">Binds 2 [2Fe-2S] clusters.</text>
    </comment>
    <comment type="subunit">
        <text evidence="3">Homodimer.</text>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the xanthine dehydrogenase family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000017436.1">
        <property type="protein sequence ID" value="ENSGAGP00000015263.1"/>
        <property type="gene ID" value="ENSGAGG00000010622.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051537">
        <property type="term" value="F:2 iron, 2 sulfur cluster binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071949">
        <property type="term" value="F:FAD binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005506">
        <property type="term" value="F:iron ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051287">
        <property type="term" value="F:NAD binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016491">
        <property type="term" value="F:oxidoreductase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.120:FF:000001">
        <property type="entry name" value="Aldehyde oxidase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.365.10:FF:000003">
        <property type="entry name" value="Aldehyde oxidase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1170.50:FF:000001">
        <property type="entry name" value="Aldehyde oxidase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.365.10:FF:000001">
        <property type="entry name" value="Xanthine dehydrogenase oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.365.10:FF:000004">
        <property type="entry name" value="Xanthine dehydrogenase oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.390.50:FF:000001">
        <property type="entry name" value="Xanthine dehydrogenase oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.43.10:FF:000001">
        <property type="entry name" value="Xanthine dehydrogenase/oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.465.10:FF:000004">
        <property type="entry name" value="Xanthine dehydrogenase/oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.20.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.465.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.120">
        <property type="entry name" value="[2Fe-2S]-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1170.50">
        <property type="entry name" value="Aldehyde oxidase/xanthine dehydrogenase, a/b hammerhead"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.365.10">
        <property type="entry name" value="Aldehyde oxidase/xanthine dehydrogenase, molybdopterin binding domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.390.50">
        <property type="entry name" value="CO dehydrogenase flavoprotein, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.43.10">
        <property type="entry name" value="Uridine Diphospho-n-acetylenolpyruvylglucosamine Reductase, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002888">
        <property type="entry name" value="2Fe-2S-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036884">
        <property type="entry name" value="2Fe-2S-bd_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036010">
        <property type="entry name" value="2Fe-2S_ferredoxin-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006058">
        <property type="entry name" value="2Fe2S_fd_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000674">
        <property type="entry name" value="Ald_Oxase/Xan_DH_a/b"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036856">
        <property type="entry name" value="Ald_Oxase/Xan_DH_a/b_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016208">
        <property type="entry name" value="Ald_Oxase/xanthine_DH-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014313">
        <property type="entry name" value="Aldehyde_oxidase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008274">
        <property type="entry name" value="AldOxase/xan_DH_MoCoBD1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046867">
        <property type="entry name" value="AldOxase/xan_DH_MoCoBD2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037165">
        <property type="entry name" value="AldOxase/xan_DH_Mopterin-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012675">
        <property type="entry name" value="Beta-grasp_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005107">
        <property type="entry name" value="CO_DH_flav_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036683">
        <property type="entry name" value="CO_DH_flav_C_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016166">
        <property type="entry name" value="FAD-bd_PCMH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036318">
        <property type="entry name" value="FAD-bd_PCMH-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016167">
        <property type="entry name" value="FAD-bd_PCMH_sub1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016169">
        <property type="entry name" value="FAD-bd_PCMH_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002346">
        <property type="entry name" value="Mopterin_DH_FAD-bd"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR02969">
        <property type="entry name" value="mam_aldehyde_ox"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45444">
        <property type="entry name" value="XANTHINE DEHYDROGENASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45444:SF3">
        <property type="entry name" value="XANTHINE DEHYDROGENASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01315">
        <property type="entry name" value="Ald_Xan_dh_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03450">
        <property type="entry name" value="CO_deh_flav_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00941">
        <property type="entry name" value="FAD_binding_5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01799">
        <property type="entry name" value="Fer2_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02738">
        <property type="entry name" value="MoCoBD_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20256">
        <property type="entry name" value="MoCoBD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000127">
        <property type="entry name" value="Xanthine_DH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01008">
        <property type="entry name" value="Ald_Xan_dh_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01092">
        <property type="entry name" value="CO_deh_flav_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54292">
        <property type="entry name" value="2Fe-2S ferredoxin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55447">
        <property type="entry name" value="CO dehydrogenase flavoprotein C-terminal domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47741">
        <property type="entry name" value="CO dehydrogenase ISP C-domain like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54665">
        <property type="entry name" value="CO dehydrogenase molybdoprotein N-domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56176">
        <property type="entry name" value="FAD-binding/transporter-associated domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56003">
        <property type="entry name" value="Molybdenum cofactor-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00197">
        <property type="entry name" value="2FE2S_FER_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51387">
        <property type="entry name" value="FAD_PCMH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6 15" id="KW-0001">2Fe-2S</keyword>
    <keyword evidence="8 14" id="KW-0274">FAD</keyword>
    <keyword evidence="5" id="KW-0285">Flavoprotein</keyword>
    <keyword evidence="10 15" id="KW-0408">Iron</keyword>
    <keyword evidence="11 15" id="KW-0411">Iron-sulfur</keyword>
    <keyword evidence="7 15" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="4 15" id="KW-0500">Molybdenum</keyword>
    <keyword evidence="9" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="FAD-binding PCMH-type" evidence="16">
        <location>
            <begin position="214"/>
            <end position="399"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="13">
        <location>
            <position position="1239"/>
        </location>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="15"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="20"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="23"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="45"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="85"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>2</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="88"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>2</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="120"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>2</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="122"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>2</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <begin position="242"/>
            <end position="249"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="345"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="407"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="745"/>
        </location>
        <ligand>
            <name>Mo-molybdopterin</name>
            <dbReference type="ChEBI" id="CHEBI:71302"/>
        </ligand>
        <ligandPart>
            <name>Mo</name>
            <dbReference type="ChEBI" id="CHEBI:28685"/>
        </ligandPart>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="776"/>
        </location>
        <ligand>
            <name>Mo-molybdopterin</name>
            <dbReference type="ChEBI" id="CHEBI:71302"/>
        </ligand>
        <ligandPart>
            <name>Mo</name>
            <dbReference type="ChEBI" id="CHEBI:28685"/>
        </ligandPart>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="890"/>
        </location>
        <ligand>
            <name>Mo-molybdopterin</name>
            <dbReference type="ChEBI" id="CHEBI:71302"/>
        </ligand>
        <ligandPart>
            <name>Mo</name>
            <dbReference type="ChEBI" id="CHEBI:28685"/>
        </ligandPart>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="892"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="1057"/>
        </location>
        <ligand>
            <name>Mo-molybdopterin</name>
            <dbReference type="ChEBI" id="CHEBI:71302"/>
        </ligand>
        <ligandPart>
            <name>Mo</name>
            <dbReference type="ChEBI" id="CHEBI:28685"/>
        </ligandPart>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001974"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006849"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00011738"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022505"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022630"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022714"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022827"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023014"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00034078"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PIRSR" id="PIRSR000127-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PIRSR" id="PIRSR000127-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PIRSR" id="PIRSR000127-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS51387"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015263.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1306" mass="143985" checksum="0F2763C8AFE92C39" modified="2019-05-08" version="1">TAAFAVRLSGTKYGCGIGGCGACTVMISTYEPTSKKIMHYPANSCLLPICSLHSSAVTTVEGVGSTKTRVHPIQERLAKCHGSQCGFCTPGMVMSMYALLRNHPEPSMEQITAALDGNLCRCTGYRPIIDTYKSFTGVGRKTNRSPFSKVSTNFECSVSGSLTKIRFSELLCSHLSTKNLMHSGLCNPEEFQPVDPTQEFIFPPELMQKRTLVFRGERTTWISPTNLRELLELKAMYPKAPLIVGNTSVGLEMKRDGVHHPVFIHPTRIPDLHVVSNTNNGLVIGAACSLAQLRDILMKVVPELPEEKTKIFRALLQQLKTLAGEQIRSLASLGGHIVSRGSTWDLNPVLAVGNSVLSLASKDGTRQIPLNDQFLAGYENADLKPKEIIVSVFIPYSKKDDFVSAFRQAERRRNAVSVVNSGMRVLFKPNTDIIMDMNILYGGTGPITVSARKSCQRLIGRQWNEQMLDEACKLVLGEISLSPSAPGGKVEYRRTLLVSFLFKFYLEVLYGLNKMVSGFPLNGLRSPTGPHCIAATGYHILIKQDVDPVQPPQDPVGRPIMHQSGVKHATGEAVYCNDIRAVDEELFLAVVTSSRAHAKIVTIDVSEALKVPGVVDVVTAEDVPGKNGTDDEQAYAKDEVICVGQIVCAVVAESAVQAKQGVEKMKIEYEDLEPILTIEVISQLSFFCLKKIEQGNIEEGFKSADEIIEGEIHIGGQEHFYLETNSVLVVPRGEDNEMDVHVSTQDAARVQEFVALALDVQSSKIMCHTKRVGGAFGGKITKPAIFAVVAAVAANKTGRPIRFTLERDEDMLITGGRHPFFGKYKVGFTNDGRITAADIEPYINAGCTLDDSEMVIEYVVLKTDNAYKIPNLRIRGRACKTNLPSNTAFRGFGFPQSGLFTESWITTVATQTGLPQEKIREINMYKEVNLTPFKEELDATNLVKCWKECMEKSEYYSRKTAVEEFNKQNYWKKKGIAIIPMKFSVGFNATQYHQAAALVHIYTDGSVLVTHGGIELGQGIHTKMLQIASRELKVPLSYVHFCETSTITVPNAIVTAGSIGSDVNGKAVQNACQILRKRLEPIINKNPEGKWEDWITEAYDQSVSLSATGYFRGYMVNMNWDTGEGQAFPYFVFGAACSEVEIDCLTGDHKNIRTDIVMDACFSINPAVDIGQIEGAFVQGVGLYTMEEIKFSPEGEQYTLGPDTYKIPAVCDIPEQFRVYLLPNSRNPIAIYSAKGIGEAGLFLGCSVFFAIWDAVAAVRKERGLTGTFTLNSPLTIERIRMACADNFTEMVRLDDHNGLFWPWNL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK23</accession>
    <name>A0A452HK23_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="11">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Endomembrane system</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="8">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HK23"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017443.1">
        <property type="protein sequence ID" value="ENSGAGP00000015269.1"/>
        <property type="gene ID" value="ENSGAGG00000011517.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012505">
        <property type="term" value="C:endomembrane system"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006898">
        <property type="term" value="P:receptor-mediated endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00112">
        <property type="entry name" value="LDLa"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.400.10:FF:000087">
        <property type="entry name" value="low-density lipoprotein receptor class A domain-containing protein 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.400.10">
        <property type="entry name" value="Low-density Lipoprotein Receptor"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036055">
        <property type="entry name" value="LDL_receptor-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050685">
        <property type="entry name" value="LDLR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023415">
        <property type="entry name" value="LDLR_class-A_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002172">
        <property type="entry name" value="LDrepeatLR_classA_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24270:SF24">
        <property type="entry name" value="LOW-DENSITY LIPOPROTEIN RECEPTOR CLASS A DOMAIN-CONTAINING PROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24270">
        <property type="entry name" value="LOW-DENSITY LIPOPROTEIN RECEPTOR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00057">
        <property type="entry name" value="Ldl_recept_a"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00261">
        <property type="entry name" value="LDLRECEPTOR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00192">
        <property type="entry name" value="LDLa"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57424">
        <property type="entry name" value="LDL receptor-like module"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01209">
        <property type="entry name" value="LDLRA_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50068">
        <property type="entry name" value="LDLRA_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7 8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6 10" id="KW-0472">Membrane</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="3 10" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 10" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="144"/>
            <end position="166"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="246"/>
            <end position="324"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="268"/>
            <end position="297"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="298"/>
            <end position="310"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="8">
        <location>
            <begin position="63"/>
            <end position="78"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="8">
        <location>
            <begin position="85"/>
            <end position="97"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="8">
        <location>
            <begin position="92"/>
            <end position="110"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="8">
        <location>
            <begin position="104"/>
            <end position="119"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004167"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004308"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00124"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015269.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="324" mass="36055" checksum="BDDB51ED373B314A" modified="2019-05-08" version="1">GATLFLLRNSLHHLLSRNVIYLSPGNEGRRLTSFLLAKAKSKCGSTFFPCASGIHCIIGRFRCNGFEDCPDGSDEENCTANPLLCSTARFHCKNGLCIDKSFVCDSQNNCQDNSDEESCESPQETGSGQDYVTSENQLLYYPSITYAIIGSSVIFVLVVAFLALVLHHQRKRNNLMSLPVHRLQHPVLLSRLVVLDHPHHCSVTYNVNNGIQYMSNQTCHRPLDLDSPPSYSEAVLDQRPPWFDLPPPPYCSDSESPSQTDLPPYRSRTGSLVSTDTPMAGSPLSTGSSRTRSVSNSMDHRRMLCERTPEQNDSLINPVTEREM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK24</accession>
    <name>A0A452HK24_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Low-density lipoprotein receptor class A domain-containing protein 3</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">May influence APP processing, resulting in a decrease in sAPP-alpha production and increased amyloidogenic P3 peptide production. May regulate ITCH and NEDD4 E3 ligase activity and degradation.</text>
    </comment>
    <comment type="subunit">
        <text evidence="14">Interacts with APP precursor C-terminus. Interacts directly with ITCH; this interaction promotes ITCH auto-ubiquitination leading to its degradation. Interacts directly with NEDD4; this interaction promotes NEDD4 auto-ubiquitination. Interacts directly with NEDD4L.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the LDLR family.</text>
    </comment>
    <comment type="caution">
        <text evidence="16">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HK24"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017438.1">
        <property type="protein sequence ID" value="ENSGAGP00000015264.1"/>
        <property type="gene ID" value="ENSGAGG00000011517.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006898">
        <property type="term" value="P:receptor-mediated endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00112">
        <property type="entry name" value="LDLa"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.400.10:FF:000087">
        <property type="entry name" value="low-density lipoprotein receptor class A domain-containing protein 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.400.10:FF:000104">
        <property type="entry name" value="Low-density lipoprotein receptor class A domain-containing protein 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.400.10">
        <property type="entry name" value="Low-density Lipoprotein Receptor"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036055">
        <property type="entry name" value="LDL_receptor-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050685">
        <property type="entry name" value="LDLR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023415">
        <property type="entry name" value="LDLR_class-A_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002172">
        <property type="entry name" value="LDrepeatLR_classA_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24270:SF24">
        <property type="entry name" value="LOW-DENSITY LIPOPROTEIN RECEPTOR CLASS A DOMAIN-CONTAINING PROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24270">
        <property type="entry name" value="LOW-DENSITY LIPOPROTEIN RECEPTOR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00057">
        <property type="entry name" value="Ldl_recept_a"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00261">
        <property type="entry name" value="LDLRECEPTOR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00192">
        <property type="entry name" value="LDLa"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57424">
        <property type="entry name" value="LDL receptor-like module"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01209">
        <property type="entry name" value="LDLRA_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50068">
        <property type="entry name" value="LDLRA_2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="9 16" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="12" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="8 18" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0675">Receptor</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <keyword evidence="4 18" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 18" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="18">
        <location>
            <begin position="168"/>
            <end position="190"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="271"/>
            <end position="349"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="293"/>
            <end position="322"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="17">
        <location>
            <begin position="323"/>
            <end position="335"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="33"/>
            <end position="51"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="45"/>
            <end position="60"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="87"/>
            <end position="102"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="109"/>
            <end position="121"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="116"/>
            <end position="134"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="128"/>
            <end position="143"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009939"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00059533"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00064132"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00074720"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00124"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015264.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="349" mass="38772" checksum="D377F5EE89091594" modified="2019-05-08" version="1">MWLLWLLLGSAESQLLPGNNFTNECNIPGNFMCSNGRCIPGAWQCDGLPDCFDESDEKECPKAKSKCGSTFFPCASGIHCIIGRFRCNGFEDCPDGSDEENCTANPLLCSTARFHCKNGLCIDKSFVCDSQNNCQDNSDEESCESPQETGSGQDYVTSENQLLYYPSITYAIIGSSVIFVLVVAFLALVLHHQRKRNNLMSLPVHRLQHPVLLSRLVVLDHPHHCSVTYNVNNGIQYMSNQTCHRPLDLDSPPSYSEAVLDQSRPPWFDLPPPPYCSDSESPSQTDLPPYRSRTGSLVSTDTPMAGSPLSTGSSRTRSVSNSMDHRRMLCERTPEQNDSLINPVTEREM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK25</accession>
    <name>A0A452HK25_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Ubiquitin carboxyl-terminal hydrolase 47</fullName>
            <ecNumber evidence="2">3.4.19.12</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="9">Deubiquitinating enzyme 47</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="8">Ubiquitin thioesterase 47</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="10">Ubiquitin-specific-processing protease 47</fullName>
        </alternativeName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1 16">
            <text>Thiol-dependent hydrolysis of ester, thioester, amide, peptide and isopeptide bonds formed by the C-terminal Gly of ubiquitin (a 76-residue protein attached to proteins as an intracellular targeting signal).</text>
            <dbReference type="EC" id="3.4.19.12"/>
        </reaction>
    </comment>
    <dbReference type="EC" id="3.4.19.12" evidence="2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017402.1">
        <property type="protein sequence ID" value="ENSGAGP00000015231.1"/>
        <property type="gene ID" value="ENSGAGG00000011497.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000017419.1">
        <property type="protein sequence ID" value="ENSGAGP00000015246.1"/>
        <property type="gene ID" value="ENSGAGG00000011497.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004843">
        <property type="term" value="F:cysteine-type deubiquitinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016579">
        <property type="term" value="P:protein deubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd02659">
        <property type="entry name" value="peptidase_C19C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd17039">
        <property type="entry name" value="Ubl_ubiquitin_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.70.10">
        <property type="entry name" value="Cysteine proteinases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038765">
        <property type="entry name" value="Papain-like_cys_pep_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050164">
        <property type="entry name" value="Peptidase_C19"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001394">
        <property type="entry name" value="Peptidase_C19_UCH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028889">
        <property type="entry name" value="USP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045578">
        <property type="entry name" value="USP47_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018200">
        <property type="entry name" value="USP_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24006">
        <property type="entry name" value="UBIQUITIN CARBOXYL-TERMINAL HYDROLASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24006:SF702">
        <property type="entry name" value="UBIQUITIN CARBOXYL-TERMINAL HYDROLASE 47"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25985">
        <property type="entry name" value="Ubiquitin_USP47_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00443">
        <property type="entry name" value="UCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19718">
        <property type="entry name" value="USP47_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54001">
        <property type="entry name" value="Cysteine proteinases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00972">
        <property type="entry name" value="USP_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00973">
        <property type="entry name" value="USP_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50235">
        <property type="entry name" value="USP_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="11" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="3" id="KW-0645">Protease</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0788">Thiol protease</keyword>
    <keyword evidence="4" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="USP" evidence="13">
        <location>
            <begin position="174"/>
            <end position="550"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="118"/>
            <end position="147"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="411"/>
            <end position="438"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="820"/>
            <end position="841"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="867"/>
            <end position="899"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="914"/>
            <end position="1011"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="11">
        <location>
            <begin position="564"/>
            <end position="591"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="416"/>
            <end position="438"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="820"/>
            <end position="836"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="870"/>
            <end position="889"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="915"/>
            <end position="925"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="927"/>
            <end position="937"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="940"/>
            <end position="956"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="984"/>
            <end position="993"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="12">
        <location>
            <begin position="994"/>
            <end position="1007"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000707"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00012759"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022807"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00026136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00029910"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00030277"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00032453"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50235"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015246.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1362" mass="155762" checksum="01283188D9BE0A59" modified="2019-05-08" version="1">MFWSCRQSLFDEMKKKFSQIESAADEPRVLCIIQDTTNSKTVNERITLNLPASTPLRRLFEDVASKVGYVNGTFDLVWGNGINVADMTPLDQTSDKSILEARFEVGKKNFLRLTDKDGEQPHIMSEESGTTDDGNQERFIGPLPTEPSLGSTNDYVCQNYSYSSILSKSETGYVGLVNQAMTCYLNSLLQTLFMTPEFRNALYKWEFEESEEDAVMSIPYQLQRLFVLLQTSKKRAIETTDVTRSFGWDSSEAWQQHDVQELCRVMFDALEQKWKQTEQADLINQLYQGKLKDYVRCLECGYEGWRIDTYLDIPLVIRPYGSNQAFASVEEALHAFIQPEILDGPNQYFCERCKKKCDARKGLRFLHFPYLLTLQLKRFDFDYTTMHRIKLNDRMTFPEELDMSTFIDVEDEKSPQTESCTDSGAENEGSCHSDQMSNDFSNDDGVDEGICLESNSGAERISKVGIEKNSLLYELFSVMVHSGSAAGGHYYACIKSFNDEQWYSFNDQHVSKITQEDIKKTYGGSSGSRGYYSSAFASSTNAYMLIYRLKDPTRNAKFLDVNEYPEHIKQLVQKEKELEEQEKKQREIERNTCKIKLFCMHPVKQIMMENKLEVHKDKTLKEAVKIAYKIMDIEEAVPLDCCRLVKYDEFHDYLERSFEEEEEDIPMGLLLGGVKSTYMFDLLLETRTPDQVFQCYKPGEVMVKVHVVDLKTESVAPSISVRAYLNQTVTEFKQLISKAVCLPAETMRVVLERCYNDLRLLNVGNKTLKAEGFFRSNKVFVESSESLDRQMAYTDSHLWKLLDRHANTIRLYVSLPEQSPGSQFKRTVNQKASGDSGNLDEACARVKGPLGNMRSVEAVLEESTEKLKSLSLQQEQEGDNADSSKSTEASDFESIESPLNEVDSLASAENRELENQIQISDPENFQSEERSDSDVNNDRSTSSVDSDILSSSHSSDTLCNVDSAPMPLANGLDSHSITSSRRSKANEGKKETWDTAEDDSGTDSEYDESGKSTGEAQYMYFKAEPYASDEGSGEGQKWLLVHVDKRITLSSFKQHLEPFVGVLSSHFKVFRVYASNQEFESVRLNETLSSFSDDNKITIRLGRALKKGEYRVKVYQLLINEPEPCKFLLDAVFAKGMTVRQSKEELLPQLREQCGLDLTIDRFRLRKKTWKNPGTVFLDYHVYEEDINISSNWEVFLEILDGAEKMKSMSQLAVLSRRWRPSEMKLDSFQEVVLESSSVDELKEKLSEISGIPLEYIEFAKGRGTFPCDISVLEIHQDLDWNPKVSALNVWPLYICDDGAVIFYRDKTEELIELTDEQRNELMKKESSRLQKTGHRVTYSPRKEKALKIYLDGAPNRESTQD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK42</accession>
    <name>A0A452HK42_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">Calcium-transporting ATPase</fullName>
            <ecNumber evidence="21">7.2.2.10</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="25 26">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="26" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="25" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="25" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="21">Catalyzes the hydrolysis of ATP coupled with the transport of calcium.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>Ca(2+)(in) + ATP + H2O = Ca(2+)(out) + ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:18105"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="7.2.2.10"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="19">
            <dbReference type="Rhea" id="RHEA:18106"/>
        </physiologicalReaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subunit">
        <text evidence="20">Interacts with sarcolipin (SLN). Interacts with phospholamban (PLN). Interacts with myoregulin (MRLN). Interacts with DWORF.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Endoplasmic reticulum membrane</location>
            <topology evidence="3">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="21">Membrane</location>
            <topology evidence="21">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Sarcoplasmic reticulum membrane</location>
            <topology evidence="2">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4 21">Belongs to the cation transport ATPase (P-type) (TC 3.A.3) family. Type IIA subfamily.</text>
    </comment>
    <comment type="caution">
        <text evidence="21">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="EC" id="7.2.2.10" evidence="21"/>
    <dbReference type="AlphaFoldDB" id="A0A452HK42"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017467.1">
        <property type="protein sequence ID" value="ENSGAGP00000015290.1"/>
        <property type="gene ID" value="ENSGAGG00000011401.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033017">
        <property type="term" value="C:sarcoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005388">
        <property type="term" value="F:P-type calcium transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.1110.10:FF:000003">
        <property type="entry name" value="Calcium-transporting ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1000:FF:000005">
        <property type="entry name" value="Calcium-transporting ATPase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1110.10:FF:000065">
        <property type="entry name" value="Sarcoplasmic/endoplasmic reticulum calcium ATPase 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.70.150.10:FF:000160">
        <property type="entry name" value="Sarcoplasmic/endoplasmic reticulum calcium ATPase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.1110.10">
        <property type="entry name" value="Calcium-transporting ATPase, cytoplasmic domain N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.70.150.10">
        <property type="entry name" value="Calcium-transporting ATPase, cytoplasmic transduction domain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1110.10">
        <property type="entry name" value="Calcium-transporting ATPase, transmembrane domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1000">
        <property type="entry name" value="HAD superfamily/HAD-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004014">
        <property type="entry name" value="ATPase_P-typ_cation-transptr_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023299">
        <property type="entry name" value="ATPase_P-typ_cyto_dom_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018303">
        <property type="entry name" value="ATPase_P-typ_P_site"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023298">
        <property type="entry name" value="ATPase_P-typ_TM_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008250">
        <property type="entry name" value="ATPase_P-typ_transduc_dom_A_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR059000">
        <property type="entry name" value="ATPase_P-type_domA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036412">
        <property type="entry name" value="HAD-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023214">
        <property type="entry name" value="HAD_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005782">
        <property type="entry name" value="P-type_ATPase_IIA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001757">
        <property type="entry name" value="P_typ_ATPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044492">
        <property type="entry name" value="P_typ_ATPase_HD_dom"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01116">
        <property type="entry name" value="ATPase-IIA1_Ca"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01494">
        <property type="entry name" value="ATPase_P-type"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR42861">
        <property type="entry name" value="CALCIUM-TRANSPORTING ATPASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13246">
        <property type="entry name" value="Cation_ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00690">
        <property type="entry name" value="Cation_ATPase_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00122">
        <property type="entry name" value="E1-E2_ATPase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08282">
        <property type="entry name" value="Hydrolase_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00119">
        <property type="entry name" value="CATATPASE"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00120">
        <property type="entry name" value="HATPASE"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDG00002">
        <property type="entry name" value="C1.7:_P-type_atpase_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDS00003">
        <property type="entry name" value="Haloacid_Dehalogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDF00027">
        <property type="entry name" value="p-type_atpase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81653">
        <property type="entry name" value="Calcium ATPase, transduction domain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81665">
        <property type="entry name" value="Calcium ATPase, transmembrane domain M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56784">
        <property type="entry name" value="HAD-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81660">
        <property type="entry name" value="Metal cation-transporting ATPase, ATP-binding domain N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00154">
        <property type="entry name" value="ATPASE_E1_E2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12 21" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="11 21" id="KW-0106">Calcium</keyword>
    <keyword evidence="7 21" id="KW-0109">Calcium transport</keyword>
    <keyword evidence="10" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="17 21" id="KW-0406">Ion transport</keyword>
    <keyword evidence="13" id="KW-0460">Magnesium</keyword>
    <keyword evidence="18 21" id="KW-0472">Membrane</keyword>
    <keyword evidence="9 21" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="26" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="14" id="KW-0703">Sarcoplasmic reticulum</keyword>
    <keyword evidence="15" id="KW-1278">Translocase</keyword>
    <keyword evidence="8 21" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="16 21" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="5 21" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="53"/>
            <end position="71"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="21">
        <location>
            <begin position="77"/>
            <end position="100"/>
        </location>
    </feature>
    <feature type="domain" description="Cation-transporting P-type ATPase N-terminal" evidence="24">
        <location>
            <begin position="34"/>
            <end position="65"/>
        </location>
    </feature>
    <feature type="domain" description="P-type ATPase A" evidence="23">
        <location>
            <begin position="116"/>
            <end position="232"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="765"/>
            <end position="788"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="22">
        <location>
            <begin position="768"/>
            <end position="788"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004326"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004477"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00005675"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022568"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022951"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022967"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00047282"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00061760"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="RuleBase" id="RU361146"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="Pfam" id="PF00122"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="Pfam" id="PF00690"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015290.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="788" mass="85718" checksum="D1C78F156561ACB0" modified="2019-05-08" version="1">MYLSVSCAHTLPLACQRKIGVGSLGGYPCCSLTLPHSPGKSIWELVLEQFEDLLVRILLLAACISFVLAWFEEGEETVTAFVEPFVILLILIANAIVGVWQERNAENAIEALKEYEPEMGKVYRADRKSVQRIKARDMVPGDIAEVAVGDKVPADIRIISIKSTTLRVDQSILTGESVSVIKHTDPVPDPRAVNQDKKNMLFSGTNVAAGKAVGIVIATGVGTEIGKIRDEMAATEQEKTPLQQKLDEFGEQLSKVISLICVAVWLINIGHFNDPVHGGSWIRGAIYYFKIAVALAVAAIPEGLPAVITTCLALGTRRMAKKNAIVRSLPSVETLGCTSVICSDKTGTLTTNQMSVCKMFVIEKVEGDVCSLNEFSITGSTYAPEGEVLKNDKHVKAGQYDGLVELATICALCNDSSLDYNEAKGAYEKVGEATETALTCLVEKMNVFNTDVRTLSKVERANACNSVIKQLMKKEFTLEFSRDRKSMSVYCSPAKASRAAVSNKMFIKGAPEGVIDRCNYVRVGTTRVPLTSLVKDKILTVIKEWGTGRDTLRCLALATRDTPPRREDMVLEDSTKFVEYEVDLTFVGCVGMLDPPRKEVMGSIQLCRDAGIRVIMITGDNKGTAIAICRRIGIFTEDEEVMGRAYTGREFDDLPPAEQREACKRACCFARVEPAHKSKIVEFLQSFDEITAMTGDGVNDAPALKKAEIGIAMGSGTAVAKTASEMVLADDNFSTIVSAVEEGRAIYNNMKQFIRYLISSNVGEVPRQPSPPLPQPCHPSPTPATPAG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK45</accession>
    <name>A0A452HK45_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">5-aminolevulinate synthase</fullName>
            <ecNumber evidence="15">2.3.1.37</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="15">5-aminolevulinic acid synthase</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="15">Delta-ALA synthase</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="15">Delta-aminolevulinate synthase</fullName>
        </alternativeName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="12">Catalyzes the pyridoxal 5'-phosphate (PLP)-dependent condensation of succinyl-CoA and glycine to form aminolevulinic acid (ALA), with CoA and CO2 as by-products. Contributes significantly to heme formation during erythropoiesis.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>succinyl-CoA + glycine + H(+) = 5-aminolevulinate + CO2 + CoA</text>
            <dbReference type="Rhea" id="RHEA:12921"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57292"/>
            <dbReference type="ChEBI" id="CHEBI:57305"/>
            <dbReference type="ChEBI" id="CHEBI:356416"/>
            <dbReference type="EC" id="2.3.1.37"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="13">
            <dbReference type="Rhea" id="RHEA:12922"/>
        </physiologicalReaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 14">
            <name>pyridoxal 5'-phosphate</name>
            <dbReference type="ChEBI" id="CHEBI:597326"/>
        </cofactor>
    </comment>
    <comment type="pathway">
        <text evidence="2 15">Porphyrin-containing compound metabolism; protoporphyrin-IX biosynthesis; 5-aminolevulinate from glycine: step 1/1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="15">Mitochondrion inner membrane</location>
            <topology evidence="15">Peripheral membrane protein</topology>
        </subcellularLocation>
        <text evidence="15">Localizes to the matrix side of the mitochondrion inner membrane.</text>
    </comment>
    <comment type="similarity">
        <text evidence="3 14">Belongs to the class-II pyridoxal-phosphate-dependent aminotransferase family.</text>
    </comment>
    <dbReference type="EC" id="2.3.1.37" evidence="15"/>
    <dbReference type="AlphaFoldDB" id="A0A452HK45"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017458.1">
        <property type="protein sequence ID" value="ENSGAGP00000015281.1"/>
        <property type="gene ID" value="ENSGAGG00000011529.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000017464.1">
        <property type="protein sequence ID" value="ENSGAGP00000015287.1"/>
        <property type="gene ID" value="ENSGAGG00000011529.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00251">
        <property type="reaction ID" value="UER00375"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005743">
        <property type="term" value="C:mitochondrial inner membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005759">
        <property type="term" value="C:mitochondrial matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003870">
        <property type="term" value="F:5-aminolevulinate synthase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030170">
        <property type="term" value="F:pyridoxal phosphate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048821">
        <property type="term" value="P:erythrocyte development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042541">
        <property type="term" value="P:hemoglobin biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006782">
        <property type="term" value="P:protoporphyrinogen IX biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="CDD" id="cd06454">
        <property type="entry name" value="KBL_like"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="5-aminolevulinate synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="5-aminolevulinate synthase, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1150.10">
        <property type="entry name" value="Aspartate Aminotransferase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.640.10">
        <property type="entry name" value="Type I PLP-dependent aspartate aminotransferase-like (Major domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010961">
        <property type="entry name" value="4pyrrol_synth_NH2levulA_synth"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015118">
        <property type="entry name" value="5aminolev_synth_preseq"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001917">
        <property type="entry name" value="Aminotrans_II_pyridoxalP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004839">
        <property type="entry name" value="Aminotransferase_I/II_large"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050087">
        <property type="entry name" value="AON_synthase_class-II"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015424">
        <property type="entry name" value="PyrdxlP-dep_Trfase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015421">
        <property type="entry name" value="PyrdxlP-dep_Trfase_major"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015422">
        <property type="entry name" value="PyrdxlP-dep_Trfase_small"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01821">
        <property type="entry name" value="5aminolev_synth"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13693:SF58">
        <property type="entry name" value="5-AMINOLEVULINATE SYNTHASE, ERYTHROID-SPECIFIC, MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13693">
        <property type="entry name" value="CLASS II AMINOTRANSFERASE/8-AMINO-7-OXONONANOATE SYNTHASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00155">
        <property type="entry name" value="Aminotran_1_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09029">
        <property type="entry name" value="Preseq_ALAS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53383">
        <property type="entry name" value="PLP-dependent transferases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00599">
        <property type="entry name" value="AA_TRANSFER_CLASS_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="11 15" id="KW-0012">Acyltransferase</keyword>
    <keyword evidence="9 15" id="KW-0350">Heme biosynthesis</keyword>
    <keyword evidence="10" id="KW-0472">Membrane</keyword>
    <keyword evidence="8 15" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="5" id="KW-0999">Mitochondrion inner membrane</keyword>
    <keyword evidence="6 14" id="KW-0663">Pyridoxal phosphate</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 15" id="KW-0808">Transferase</keyword>
    <keyword evidence="7" id="KW-0809">Transit peptide</keyword>
    <feature type="domain" description="5-aminolevulinate synthase presequence" evidence="18">
        <location>
            <begin position="3"/>
            <end position="108"/>
        </location>
    </feature>
    <feature type="domain" description="Aminotransferase class I/classII large" evidence="17">
        <location>
            <begin position="202"/>
            <end position="546"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="41"/>
            <end position="79"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001933"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005029"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00008392"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022792"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022898"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023133"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023315"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00045913"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00049013"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="RuleBase" id="RU003693"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="RuleBase" id="RU910713"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF00155"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF09029"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015287.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="597" mass="65058" checksum="6DCBFE4F73DA74F7" modified="2019-05-08" version="1">MASFLQRCPFLTRDPSIFLLKARPLLVSSAQRCPIMVARTLMNSSPDPQRGTDNPIPLAAGSVPQELETPETASSPSQTHCPFMELEVRDRRSRIVQRAGPEVQEDVKVYKADPLSSLLLEVQSSLRKKFLGRDGPGPDARLLPTHLLQDNLPGSGAFSYDSFLEGRLEAKRRDHTYRVFKTVTRRADAFPFAREAAGPEVSVWCSNDYLAMSRHPRVLQAAMDALQHHGLGAGGTRNISGTSQYHVDLERELAQLHRKDAALLFSSCYVANDSTLFTLARMMPGCEIFSDAGNHASMIQGIRNSGVPKHVFRHNDPQHLAQLLGRSPPGIPKIVAFETVHSMDGGICPLEELCDVAHEHGALTFVDEVHAVGLYGARGAGIAERDGVLGKVDIVSGTLGKAFGCVGGYIASTAALVDTVRSYAAGFIFTTSLPPPVLAGALASLRVLAGPEGRGLRRAHQRNVKHMRQLLMDAGLPLVRCPSHIIPIRVGDAALNSQLCDLLLSQHNIYVQAINYPTVPRGEELLRLAPSPHHTPPMMDYFVERLVQGWQDVGLPLQPPASSACHFCQRPLHFALMSEWERDSFGSLGAQCVTSSA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK47</accession>
    <name>A0A452HK47_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="17">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the tenascin family.</text>
    </comment>
    <comment type="caution">
        <text evidence="12">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000017472.1">
        <property type="protein sequence ID" value="ENSGAGP00000015295.1"/>
        <property type="gene ID" value="ENSGAGG00000011532.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030155">
        <property type="term" value="P:regulation of cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00054">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="CDD" id="cd00087">
        <property type="entry name" value="FReD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000099">
        <property type="entry name" value="Fibronectin 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000001">
        <property type="entry name" value="Tenascin C"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000162">
        <property type="entry name" value="Tenascin C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000201">
        <property type="entry name" value="Tenascin C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000207">
        <property type="entry name" value="Tenascin C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000274">
        <property type="entry name" value="Tenascin C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000293">
        <property type="entry name" value="Tenascin C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000398">
        <property type="entry name" value="Tenascin C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000529">
        <property type="entry name" value="Tenascin C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000611">
        <property type="entry name" value="Tenascin C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.215.10:FF:000001">
        <property type="entry name" value="Tenascin isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.20.25.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.215.10">
        <property type="entry name" value="Gamma Fibrinogen, chain A, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050991">
        <property type="entry name" value="ECM_Regulatory_Proteins"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013111">
        <property type="entry name" value="EGF_extracell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041161">
        <property type="entry name" value="EGF_Tenascin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036056">
        <property type="entry name" value="Fibrinogen-like_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014716">
        <property type="entry name" value="Fibrinogen_a/b/g_C_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002181">
        <property type="entry name" value="Fibrinogen_a/b/g_C_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF040941">
        <property type="entry name" value="GGGWT_bact"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46708">
        <property type="entry name" value="TENASCIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46708:SF1">
        <property type="entry name" value="TENASCIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07974">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25024">
        <property type="entry name" value="EGF_TEN"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18720">
        <property type="entry name" value="EGF_Tenascin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00147">
        <property type="entry name" value="Fibrinogen_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="SMART" id="SM00186">
        <property type="entry name" value="FBG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56496">
        <property type="entry name" value="Fibrinogen C-terminal domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01186">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51406">
        <property type="entry name" value="FIBRINOGEN_C_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="8"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="10 12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5 12" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="4" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0964">Secreted</keyword>
    <keyword evidence="6 13" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="13">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019539652" evidence="13">
        <location>
            <begin position="23"/>
            <end position="1687"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="14">
        <location>
            <begin position="188"/>
            <end position="219"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="15">
        <location>
            <begin position="657"/>
            <end position="747"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="15">
        <location>
            <begin position="837"/>
            <end position="927"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="15">
        <location>
            <begin position="928"/>
            <end position="1018"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="15">
        <location>
            <begin position="1019"/>
            <end position="1108"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="15">
        <location>
            <begin position="1109"/>
            <end position="1197"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="15">
        <location>
            <begin position="1198"/>
            <end position="1287"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="15">
        <location>
            <begin position="1288"/>
            <end position="1374"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="15">
        <location>
            <begin position="1375"/>
            <end position="1463"/>
        </location>
    </feature>
    <feature type="domain" description="Fibrinogen C-terminal" evidence="16">
        <location>
            <begin position="1461"/>
            <end position="1676"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="192"/>
            <end position="202"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="209"/>
            <end position="218"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008673"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS51406"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015295.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1687" mass="186177" checksum="BA9C0CE7A7661DD3" modified="2019-05-08" version="1">MGFPMQVLTCAIIALLHQHVNGGLIKRIIRQKRETGLNVTLPEDNQSVVFNHVYNINVPMGSLCSVDLDSANEDTDLKSQVEPSKTYQEHTVNEGNQIVFTHRINIPRRACGCAAAPDIKDLLSRLEELEGLVSSLREQCSSSAGCCPSGLTAEGQVDTTPYCSGRGNYSSEHCGCICEPGWKGPNCSEPECLNNCNNKGRCVNGKCICDEGFTGDDCSQLTCPNDCSDQGKCINGVCLCFEEYTGEDCSEERCPVACNEHGKCVKGRCVCDEGFTGEDCSEPLCLNNCNNRGRCVEKECVCNEGYTGEDCGELICPNDCYDRGRCVNGTCYCEEGFTGEDCGELTCPNNCNNNGRCDNGLCICNEGFVGDDCSERRCPKDCNKRGRCVNGKCVCNEGFAGAACGQVKCPKDCNKQGRCVNGQCVCQEGFMGEDCSELRCPGDCSNWGRCVNGRCECDEGFTGEDCSQLKCPNDCHNRGRCVNRQCVCNEGFIGEDCGELRCLNDCNNRGRCVNGQCVCEEGFVGEDCGELRCPNNCNEHGRCINGQCVCDEGYMGEDCAELRCPNDCNNRGLCVNGQCECDNGFTAMDCSQQRCPNDCNNRGRCINGQCLCEEGFGGVDCSERSCPNNCNNMGHCVDGQCICDDGYEGDDCSNVSPPSDLTVTDVTDKTVNLEWKNENLVNEYLITYVPTSVGGLELQFRVPGNETSATIRELEPGVEYFIRVFAILKNKKSIPISARVATYLPAPEGLKFKSIRETSVQVEWDPLNISFDGWELIFRNMKKEDNGDITNSLKRPDTSYMQPGLAPGQQYNVSLHVVKNKTRGPGLFRVFTTKLDGPSQIEARDVTDTTALITWFKPLAEIDDMELTYGPKDVPGDRTTIDLSEDESQYSVGNLKPHTEYEVTLISRRGDMASDPVKEIFVTDLDAPKNLKRVSQTDNSITLEWKNSQASIDTYRIKFAPISGGDHAEITVPRSNQTITKATLTGLRPGTEYGIGVTAMKQDRESTPATINAATDLDNPKDLVVSDNTETTLSLRWRRPLAKFDRYRLTYMIPSGRKNEIEIPADSTSYILRELDPGTEYTITLVAEKGRHKSKPTTVKESTEAEPEVDNLLVSDATPSGFRLSWTADDGVFDSFVLKIRDTDRQSDPRELIVPGHERTQDITGLKEGTKYDIELYGLISGQRSQPISAVASTAVGSPKAISFSDITENSATVNWSPPRTRVESFRISYVPITGGAPNVVTVDGTKTRTKLVKLIPGVDYTVSIISIKGFEESEPISGILTTALDSPSGLVAVNITDSEALAVWQPAIAAVDNYVISYIAEDEPEVTQTVSGNTVEFDLNGLRPATEYMLRIYAVKGPQKSETVFTKFTTGMDAPRDLSAIEVQSETAVLSWRPPRASVTGYLLIYESIDGKVKEVILGPEATSYSLSELNPSTQYTVKLQALNRALKSKIIQTIFTTIGLLYPYPKDCSQALLNGESASGLYTIYMNGNKSQPMEVYCDMISDGGGWIVFLRRHNGKEDFYRNWRTYAAGFGDPTDEFWIGLENLHKITSQGQYELRVDLRDKGETAYALYDRFSVGDSKTRYRLKVDGYSGTAGDSMIYHNGRSFSTFDKDNDAAITNCALSYKGAFWYKNCHRVNLMGRYGDNSHSQGVNWFHWKGHEYSIQFAEMKLRPISFRNLEGRRKRA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK48</accession>
    <name>A0A452HK48_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="17">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the tenascin family.</text>
    </comment>
    <comment type="caution">
        <text evidence="12">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000017468.1">
        <property type="protein sequence ID" value="ENSGAGP00000015291.1"/>
        <property type="gene ID" value="ENSGAGG00000011532.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030155">
        <property type="term" value="P:regulation of cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00054">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="CDD" id="cd00087">
        <property type="entry name" value="FReD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000099">
        <property type="entry name" value="Fibronectin 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000001">
        <property type="entry name" value="Tenascin C"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000162">
        <property type="entry name" value="Tenascin C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000201">
        <property type="entry name" value="Tenascin C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000207">
        <property type="entry name" value="Tenascin C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000293">
        <property type="entry name" value="Tenascin C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000398">
        <property type="entry name" value="Tenascin C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000529">
        <property type="entry name" value="Tenascin C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000611">
        <property type="entry name" value="Tenascin C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.215.10:FF:000001">
        <property type="entry name" value="Tenascin isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.20.25.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.215.10">
        <property type="entry name" value="Gamma Fibrinogen, chain A, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050991">
        <property type="entry name" value="ECM_Regulatory_Proteins"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013111">
        <property type="entry name" value="EGF_extracell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041161">
        <property type="entry name" value="EGF_Tenascin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036056">
        <property type="entry name" value="Fibrinogen-like_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014716">
        <property type="entry name" value="Fibrinogen_a/b/g_C_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002181">
        <property type="entry name" value="Fibrinogen_a/b/g_C_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF040941">
        <property type="entry name" value="GGGWT_bact"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46708">
        <property type="entry name" value="TENASCIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46708:SF1">
        <property type="entry name" value="TENASCIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07974">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25024">
        <property type="entry name" value="EGF_TEN"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18720">
        <property type="entry name" value="EGF_Tenascin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00147">
        <property type="entry name" value="Fibrinogen_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="SMART" id="SM00186">
        <property type="entry name" value="FBG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56496">
        <property type="entry name" value="Fibrinogen C-terminal domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01186">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51406">
        <property type="entry name" value="FIBRINOGEN_C_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="9"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="10 12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5 12" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="4" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0964">Secreted</keyword>
    <keyword evidence="6 13" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="13">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019261999" evidence="13">
        <location>
            <begin position="23"/>
            <end position="1778"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="14">
        <location>
            <begin position="188"/>
            <end position="219"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="15">
        <location>
            <begin position="657"/>
            <end position="747"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="15">
        <location>
            <begin position="837"/>
            <end position="927"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="15">
        <location>
            <begin position="928"/>
            <end position="1018"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="15">
        <location>
            <begin position="1019"/>
            <end position="1107"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="15">
        <location>
            <begin position="1108"/>
            <end position="1198"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="15">
        <location>
            <begin position="1199"/>
            <end position="1288"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="15">
        <location>
            <begin position="1289"/>
            <end position="1378"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="15">
        <location>
            <begin position="1379"/>
            <end position="1465"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="15">
        <location>
            <begin position="1466"/>
            <end position="1554"/>
        </location>
    </feature>
    <feature type="domain" description="Fibrinogen C-terminal" evidence="16">
        <location>
            <begin position="1552"/>
            <end position="1767"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="192"/>
            <end position="202"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="209"/>
            <end position="218"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008673"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS51406"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015291.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1778" mass="196158" checksum="67F8C8E6333EEFE7" modified="2019-05-08" version="1">MGFPMQVLTCAIIALLHQHVNGGLIKRIIRQKRETGLNVTLPEDNQSVVFNHVYNINVPMGSLCSVDLDSANEDTDLKSQVEPSKTYQEHTVNEGNQIVFTHRINIPRRACGCAAAPDIKDLLSRLEELEGLVSSLREQCSSSAGCCPSGLTAEGQVDTTPYCSGRGNYSSEHCGCICEPGWKGPNCSEPECLNNCNNKGRCVNGKCICDEGFTGDDCSQLTCPNDCSDQGKCINGVCLCFEEYTGEDCSEERCPVACNEHGKCVKGRCVCDEGFTGEDCSEPLCLNNCNNRGRCVEKECVCNEGYTGEDCGELICPNDCYDRGRCVNGTCYCEEGFTGEDCGELTCPNNCNNNGRCDNGLCICNEGFVGDDCSERRCPKDCNKRGRCVNGKCVCNEGFAGAACGQVKCPKDCNKQGRCVNGQCVCQEGFMGEDCSELRCPGDCSNWGRCVNGRCECDEGFTGEDCSQLKCPNDCHNRGRCVNRQCVCNEGFIGEDCGELRCLNDCNNRGRCVNGQCVCEEGFVGEDCGELRCPNNCNEHGRCINGQCVCDEGYMGEDCAELRCPNDCNNRGLCVNGQCECDNGFTAMDCSQQRCPNDCNNRGRCINGQCLCEEGFGGVDCSERSCPNNCNNMGHCVDGQCICDDGYEGDDCSNVSPPSDLTVTDVTDKTVNLEWKNENLVNEYLITYVPTSVGGLELQFRVPGNETSATIRELEPGVEYFIRVFAILKNKKSIPISARVATYLPAPEGLKFKSIRETSVQVEWDPLNISFDGWELIFRNMKKEDNGDITNSLKRPDTSYMQPGLAPGQQYNVSLHVVKNKTRGPGLFRVFTTKLDGPSQIEARDVTDTTALITWFKPLAEIDDMELTYGPKDVPGDRTTIDLSEDESQYSVGNLKPHTEYEVTLISRRGDMASDPVKEIFVTDLDAPKNLKRVSQTDNSITLEWKNSQASIDTYRIKFAPISGGDHAEITVPRSNQTITKATLTGLRPGTEYGIGVTAMKQDRESTPATINAATDLDNPKDLVVSDNTETTLSLRWRRPLAKFDRYRLTYMIPSGRKNEIEIPADSTSYILRELDPGTEYTITLVAEKGRHKSKPTTVKESTEEEPELGKLTVSKVSWDGFQLNWTAAEGAYETFVIQVQEPNNPEDTRNLTVPGSLRFVNVTGLKANMPYTVILYGVIQGYRTKPLSVNTTTAAEPEVGELMVSSITPESFNLSWTAPDEAFKTFTIEIIDSSRLLEPMEYNISGNLRTAHISGLSPKTDFIVYLSGSTRGFRTKTISAAATTAVGSPKAISFSDITENSATVNWSPPRTRVESFRISYVPITGGAPNVVTVDGTKTRTKLVKLIPGVDYTVSIISIKGFEESEPISGILTTALDSPSGLVAVNITDSEALAVWQPAIAAVDNYVISYIAEDEPEVTQTVSGNTVEFDLNGLRPATEYMLRIYAVKGPQKSETVFTKFTTGMDAPRDLSAIEVQSETAVLSWRPPRASVTGYLLIYESIDGKVKEVILGPEATSYSLSELNPSTQYTVKLQALNRALKSKIIQTIFTTIGLLYPYPKDCSQALLNGESASGLYTIYMNGNKSQPMEVYCDMISDGGGWIVFLRRHNGKEDFYRNWRTYAAGFGDPTDEFWIGLENLHKITSQGQYELRVDLRDKGETAYALYDRFSVGDSKTRYRLKVDGYSGTAGDSMIYHNGRSFSTFDKDNDAAITNCALSYKGAFWYKNCHRVNLMGRYGDNSHSQGVNWFHWKGHEYSIQFAEMKLRPISFRNLEGRRKRA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK50</accession>
    <name>A0A452HK50_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="17">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the tenascin family.</text>
    </comment>
    <comment type="caution">
        <text evidence="12">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000017477.1">
        <property type="protein sequence ID" value="ENSGAGP00000015300.1"/>
        <property type="gene ID" value="ENSGAGG00000011532.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030155">
        <property type="term" value="P:regulation of cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="CDD" id="cd00087">
        <property type="entry name" value="FReD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000099">
        <property type="entry name" value="Fibronectin 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000001">
        <property type="entry name" value="Tenascin C"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000162">
        <property type="entry name" value="Tenascin C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000201">
        <property type="entry name" value="Tenascin C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000207">
        <property type="entry name" value="Tenascin C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000293">
        <property type="entry name" value="Tenascin C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000398">
        <property type="entry name" value="Tenascin C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000529">
        <property type="entry name" value="Tenascin C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000611">
        <property type="entry name" value="Tenascin C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.215.10:FF:000001">
        <property type="entry name" value="Tenascin isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.20.25.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.215.10">
        <property type="entry name" value="Gamma Fibrinogen, chain A, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050991">
        <property type="entry name" value="ECM_Regulatory_Proteins"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013111">
        <property type="entry name" value="EGF_extracell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041161">
        <property type="entry name" value="EGF_Tenascin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036056">
        <property type="entry name" value="Fibrinogen-like_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014716">
        <property type="entry name" value="Fibrinogen_a/b/g_C_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002181">
        <property type="entry name" value="Fibrinogen_a/b/g_C_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF040941">
        <property type="entry name" value="GGGWT_bact"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46708">
        <property type="entry name" value="TENASCIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46708:SF1">
        <property type="entry name" value="TENASCIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07974">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25024">
        <property type="entry name" value="EGF_TEN"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18720">
        <property type="entry name" value="EGF_Tenascin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00147">
        <property type="entry name" value="Fibrinogen_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="SMART" id="SM00186">
        <property type="entry name" value="FBG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56496">
        <property type="entry name" value="Fibrinogen C-terminal domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01186">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51406">
        <property type="entry name" value="FIBRINOGEN_C_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="7"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="10 12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5 12" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="4" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0964">Secreted</keyword>
    <keyword evidence="6 13" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="13">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019136251" evidence="13">
        <location>
            <begin position="23"/>
            <end position="1596"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="14">
        <location>
            <begin position="188"/>
            <end position="219"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="15">
        <location>
            <begin position="657"/>
            <end position="747"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="15">
        <location>
            <begin position="837"/>
            <end position="927"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="15">
        <location>
            <begin position="928"/>
            <end position="1018"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="15">
        <location>
            <begin position="1019"/>
            <end position="1108"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="15">
        <location>
            <begin position="1109"/>
            <end position="1195"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="15">
        <location>
            <begin position="1196"/>
            <end position="1283"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="15">
        <location>
            <begin position="1284"/>
            <end position="1372"/>
        </location>
    </feature>
    <feature type="domain" description="Fibrinogen C-terminal" evidence="16">
        <location>
            <begin position="1370"/>
            <end position="1585"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="192"/>
            <end position="202"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="12">
        <location>
            <begin position="209"/>
            <end position="218"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008673"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS51406"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015300.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1596" mass="176172" checksum="20B75919196D19C1" modified="2019-05-08" version="1">MGFPMQVLTCAIIALLHQHVNGGLIKRIIRQKRETGLNVTLPEDNQSVVFNHVYNINVPMGSLCSVDLDSANEDTDLKSQVEPSKTYQEHTVNEGNQIVFTHRINIPRRACGCAAAPDIKDLLSRLEELEGLVSSLREQCSSSAGCCPSGLTAEGQVDTTPYCSGRGNYSSEHCGCICEPGWKGPNCSEPECLNNCNNKGRCVNGKCICDEGFTGDDCSQLTCPNDCSDQGKCINGVCLCFEEYTGEDCSEERCPVACNEHGKCVKGRCVCDEGFTGEDCSEPLCLNNCNNRGRCVEKECVCNEGYTGEDCGELICPNDCYDRGRCVNGTCYCEEGFTGEDCGELTCPNNCNNNGRCDNGLCICNEGFVGDDCSERRCPKDCNKRGRCVNGKCVCNEGFAGAACGQVKCPKDCNKQGRCVNGQCVCQEGFMGEDCSELRCPGDCSNWGRCVNGRCECDEGFTGEDCSQLKCPNDCHNRGRCVNRQCVCNEGFIGEDCGELRCLNDCNNRGRCVNGQCVCEEGFVGEDCGELRCPNNCNEHGRCINGQCVCDEGYMGEDCAELRCPNDCNNRGLCVNGQCECDNGFTAMDCSQQRCPNDCNNRGRCINGQCLCEEGFGGVDCSERSCPNNCNNMGHCVDGQCICDDGYEGDDCSNVSPPSDLTVTDVTDKTVNLEWKNENLVNEYLITYVPTSVGGLELQFRVPGNETSATIRELEPGVEYFIRVFAILKNKKSIPISARVATYLPAPEGLKFKSIRETSVQVEWDPLNISFDGWELIFRNMKKEDNGDITNSLKRPDTSYMQPGLAPGQQYNVSLHVVKNKTRGPGLFRVFTTKLDGPSQIEARDVTDTTALITWFKPLAEIDDMELTYGPKDVPGDRTTIDLSEDESQYSVGNLKPHTEYEVTLISRRGDMASDPVKEIFVTDLDAPKNLKRVSQTDNSITLEWKNSQASIDTYRIKFAPISGGDHAEITVPRSNQTITKATLTGLRPGTEYGIGVTAMKQDRESTPATINAATDLDNPKDLVVSDNTETTLSLRWRRPLAKFDRYRLTYMIPSGRKNEIEIPADSTSYILRELDPGTEYTITLVAEKGRHKSKPTTVKESTAVGSPKAISFSDITENSATVNWSPPRTRVESFRISYVPITGGAPNVVTVDGTKTRTKLVKLIPGVDYTVSIISIKGFEESEPISGILTTALDSPSGLVAVNITDSEALAVWQPAIAAVDNYVISYIAEDEPEVTQTVSGNTVEFDLNGLRPATEYMLRIYAVKGPQKSETVFTKFTTGMDAPRDLSAIEVQSETAVLSWRPPRASVTGYLLIYESIDGKVKEVILGPEATSYSLSELNPSTQYTVKLQALNRALKSKIIQTIFTTIGLLYPYPKDCSQALLNGESASGLYTIYMNGNKSQPMEVYCDMISDGGGWIVFLRRHNGKEDFYRNWRTYAAGFGDPTDEFWIGLENLHKITSQGQYELRVDLRDKGETAYALYDRFSVGDSKTRYRLKVDGYSGTAGDSMIYHNGRSFSTFDKDNDAAITNCALSYKGAFWYKNCHRVNLMGRYGDNSHSQGVNWFHWKGHEYSIQFAEMKLRPISFRNLEGRRKRA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK51</accession>
    <name>A0A452HK51_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">protein-arginine deiminase</fullName>
            <ecNumber evidence="3">3.5.3.15</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="7">
            <text>L-arginyl-[protein] + H2O = L-citrullyl-[protein] + NH4(+)</text>
            <dbReference type="Rhea" id="RHEA:18089"/>
            <dbReference type="Rhea" id="RHEA-COMP:10532"/>
            <dbReference type="Rhea" id="RHEA-COMP:10588"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:28938"/>
            <dbReference type="ChEBI" id="CHEBI:29965"/>
            <dbReference type="ChEBI" id="CHEBI:83397"/>
            <dbReference type="EC" id="3.5.3.15"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the protein arginine deiminase family.</text>
    </comment>
    <dbReference type="EC" id="3.5.3.15" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452HK51"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017450.1">
        <property type="protein sequence ID" value="ENSGAGP00000015273.1"/>
        <property type="gene ID" value="ENSGAGG00000011443.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004668">
        <property type="term" value="F:protein-arginine deiminase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1700:FF:000001">
        <property type="entry name" value="Protein-arginine deiminase type-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.75.10.10:FF:000003">
        <property type="entry name" value="Protein-arginine deiminase type-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.75.10.10">
        <property type="entry name" value="L-arginine/glycine Amidinotransferase, Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1700">
        <property type="entry name" value="Protein-arginine deiminase, central domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1860">
        <property type="entry name" value="Protein-arginine deiminase, N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008972">
        <property type="entry name" value="Cupredoxin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004303">
        <property type="entry name" value="PAD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013530">
        <property type="entry name" value="PAD_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036556">
        <property type="entry name" value="PAD_central_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013732">
        <property type="entry name" value="PAD_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038685">
        <property type="entry name" value="PAD_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013733">
        <property type="entry name" value="Prot_Arg_deaminase_cen_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10837">
        <property type="entry name" value="PEPTIDYLARGININE DEIMINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10837:SF11">
        <property type="entry name" value="PROTEIN-ARGININE DEIMINASE TYPE-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03068">
        <property type="entry name" value="PAD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08527">
        <property type="entry name" value="PAD_M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08526">
        <property type="entry name" value="PAD_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF001247">
        <property type="entry name" value="Protein-arginine_deiminase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49503">
        <property type="entry name" value="Cupredoxins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55909">
        <property type="entry name" value="Pentein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF110083">
        <property type="entry name" value="Peptidylarginine deiminase Pad4, middle domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0106">Calcium</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="5" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Protein-arginine deiminase (PAD) N-terminal" evidence="10">
        <location>
            <begin position="22"/>
            <end position="99"/>
        </location>
    </feature>
    <feature type="domain" description="Protein-arginine deiminase (PAD) central" evidence="11">
        <location>
            <begin position="126"/>
            <end position="263"/>
        </location>
    </feature>
    <feature type="domain" description="Protein-arginine deiminase C-terminal" evidence="9">
        <location>
            <begin position="270"/>
            <end position="645"/>
        </location>
    </feature>
    <feature type="active site" evidence="8">
        <location>
            <position position="337"/>
        </location>
    </feature>
    <feature type="active site" evidence="8">
        <location>
            <position position="457"/>
        </location>
    </feature>
    <feature type="active site" evidence="8">
        <location>
            <position position="459"/>
        </location>
    </feature>
    <feature type="active site" evidence="8">
        <location>
            <position position="630"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008166"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012200"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00048487"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PIRSR" id="PIRSR001247-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF03068"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="Pfam" id="PF08526"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF08527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015273.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="648" mass="72682" checksum="F2B1F7AF978217FD" modified="2019-05-08" version="1">KVPCAPWHPILFIPALPGLDCGATSFDARGTSGVKIYVVYNPNAVKKAAGASRWTLNSQVEVIVSMDIPSSTVNDSKVRISYYGQKENAPIEGAWLYLTCVSKYCKNNRLNTLETCNLDPFVFPLQTKWTWGPEGQGAVLLVNCDRDSPGAKDMDSAYQYVQSYADLEDMSLMILRTQGPSAIFADYKLVLHISVADVNKVRVFHASGDGSLAQYKHVLGSDKRSYVVDRSSGQEEDTFYVEGLAFPDADFSGLVSFHVTLLENAPDTPIFTDTVVFRVAPWIMTPNTLQPLEVFVCRGSNPNGDFLEAMRALVRKANCKLTVCPEVENRRDRWIQDEMEFGYVEAPHKMFPVVFDSPRDRGLRDFAFRRILGPDFGYVTRVPPHGHVSSLDSFGNLDVSPPVTVQGKTYPLGRILIGSSLPRSDGRRMTKAVRDFLYAQKVQPPVELYSDWLSVGHVDEFLSFVPASDRKGFRLLLASPDACYKLFKEKQREGHGKAAQFAGLENVHTITIDEMLADKTLRSDSEYVQSCIDWNRHTLKEELGLTEQDIIDIPQLFMLKGSRADAFFPDVVNMIVLGKHLGIPKPFGPIVDGRCCLEEKVRSLLEPLGLVCIFINDFFSYHTLLGEIHCGTNVLRKPFSFKWWNMIL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK52</accession>
    <name>A0A452HK52_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="26">NACHT, LRR and PYD domains-containing protein 3</fullName>
        </recommendedName>
    </protein>
    <organism evidence="31 32">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="32" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="31" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="31" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="27">Independently of inflammasome activation, regulates the differentiation of T helper 2 (Th2) cells and has a role in Th2 cell-dependent asthma and tumor growth. During Th2 differentiation, required for optimal IRF4 binding to IL4 promoter and for IRF4-dependent IL4 transcription. Binds to the consensus DNA sequence 5'-GRRGGNRGAG-3'. May also participate in the transcription of IL5, IL13, GATA3, CCR3, CCR4 and MAF.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="28">
            <text>ATP + H2O = ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:13065"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="28">
            <dbReference type="Rhea" id="RHEA:13066"/>
        </physiologicalReaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="4">Cytoplasm</location>
            <location evidence="4">Cytoskeleton</location>
            <location evidence="4">Microtubule organizing center</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Endoplasmic reticulum</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="5">Golgi apparatus membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Inflammasome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Mitochondrion</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="6">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="7">Belongs to the NLRP family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HK52"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017478.1">
        <property type="protein sequence ID" value="ENSGAGP00000015301.1"/>
        <property type="gene ID" value="ENSGAGG00000011540.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061702">
        <property type="term" value="C:canonical inflammasome complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000139">
        <property type="term" value="C:Golgi membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005815">
        <property type="term" value="C:microtubule organizing center"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006954">
        <property type="term" value="P:inflammatory response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045087">
        <property type="term" value="P:innate immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd08321">
        <property type="entry name" value="Pyrin_ASC-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000442">
        <property type="entry name" value="NACHT, LRR and PYD domains-containing protein 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.533.10">
        <property type="entry name" value="Death Domain, Fas"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004020">
        <property type="entry name" value="DAPIN"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011029">
        <property type="entry name" value="DEATH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001611">
        <property type="entry name" value="Leu-rich_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007111">
        <property type="entry name" value="NACHT_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041267">
        <property type="entry name" value="NLRP_HD2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050637">
        <property type="entry name" value="NLRP_innate_immun_reg"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041075">
        <property type="entry name" value="NOD1/2_WH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45690">
        <property type="entry name" value="NACHT, LRR AND PYD DOMAINS-CONTAINING PROTEIN 12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45690:SF19">
        <property type="entry name" value="NACHT, LRR AND PYD DOMAINS-CONTAINING PROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13516">
        <property type="entry name" value="LRR_6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05729">
        <property type="entry name" value="NACHT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17776">
        <property type="entry name" value="NLRC4_HD2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02758">
        <property type="entry name" value="PYRIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17779">
        <property type="entry name" value="WHD_NOD2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00368">
        <property type="entry name" value="LRR_RI"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM01289">
        <property type="entry name" value="PYRIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47986">
        <property type="entry name" value="DEATH domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52047">
        <property type="entry name" value="RNI-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50824">
        <property type="entry name" value="DAPIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50837">
        <property type="entry name" value="NACHT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="21" id="KW-0010">Activator</keyword>
    <keyword evidence="16" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="8" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="24" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="15" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="19" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="14" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="25" id="KW-1271">Inflammasome</keyword>
    <keyword evidence="23" id="KW-0395">Inflammatory response</keyword>
    <keyword evidence="9" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="20" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="13" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="11" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="32" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="12" id="KW-0677">Repeat</keyword>
    <keyword evidence="10" id="KW-0964">Secreted</keyword>
    <keyword evidence="22" id="KW-0804">Transcription</keyword>
    <keyword evidence="18" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="17" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="Pyrin" evidence="29">
        <location>
            <begin position="1"/>
            <end position="94"/>
        </location>
    </feature>
    <feature type="domain" description="NACHT" evidence="30">
        <location>
            <begin position="154"/>
            <end position="359"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004110"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004173"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004267"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004394"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00008665"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00023198"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00023233"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00040040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="ARBA" id="ARBA00045987"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="ARBA" id="ARBA00048778"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="29">
        <source>
            <dbReference type="PROSITE" id="PS50824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="30">
        <source>
            <dbReference type="PROSITE" id="PS50837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="31">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015301.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="32">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="721" mass="82647" checksum="1AC2139DA839DCC8" modified="2019-05-08" version="1">VSGEEGGINDLLIHALNNLSQEEFKGFKDKLSHSDFEGKGTIPRGRLENADAIDTKNLLMAFYGGDAAVDVAIEVFNQISLREAAANLREERENDYRVKYREHIQNKYRVIKDLNSRLGENVTLNSRYTKLIIRTSFITLGNLFKCDEDGQTPQIVVLLGVAGIGKTMTARKIMFDWASGELYKEKFDYAFYINCREVNFGSEQGRIADLILKNCPNKNAPVEEILINPEKLLFIIDGFDELRFSLDQPEDSLCSDPCEKQPMEITLSSLVRKTVLLESYVIITTRPTALEKLRQCLECARYAEILGFSAAEREEYFHNFLGNEEKARKAFNFVKEHEMLFTLCFVPIACWIICTVLKQQMEAGEDLAQTSKTITGVYMLYLFSLLKGHSSNSKKHTPANLKGLCSLAADGIWRQKILFGEEEIKKHGLDISDSLFLNENMFQKDIDCECVYSFIHLSFQEFFATLFYVLEKEETTTVEGSETAMNNVKKLLENYGKSRNYLMLTVRFLFGLLNEERMEDMEKKLSCKISPKIKPELLEWELFLEPDEEFSSFKRNAEMIYQLEEFHCLYEIQEESFVQSALNHFTELYLAEYNFTKMDQMALSFCLKNCCKLETLYLNLCTFGSEDPNEDSLREQHQDEPTCSHIYLLCQALKEPNCKLKKLKLSNCHLTAVCGGDLSSLLSTKPTLMELDIGENNLGDSGVQQLCEGLKHPHSKLQKLR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK53</accession>
    <name>A0A452HK53_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">FAD-binding PCMH-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="1 13">
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="14">
            <name>Mo-molybdopterin</name>
            <dbReference type="ChEBI" id="CHEBI:71302"/>
        </cofactor>
        <text evidence="14">Binds 1 Mo-molybdopterin (Mo-MPT) cofactor per subunit.</text>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="11">
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="14">
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:49601"/>
        </cofactor>
        <text evidence="14">Binds 2 [2Fe-2S] clusters.</text>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the xanthine dehydrogenase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HK53"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017474.1">
        <property type="protein sequence ID" value="ENSGAGP00000015297.1"/>
        <property type="gene ID" value="ENSGAGG00000010622.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051537">
        <property type="term" value="F:2 iron, 2 sulfur cluster binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071949">
        <property type="term" value="F:FAD binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005506">
        <property type="term" value="F:iron ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016491">
        <property type="term" value="F:oxidoreductase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.365.10:FF:000003">
        <property type="entry name" value="Aldehyde oxidase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.365.10:FF:000001">
        <property type="entry name" value="Xanthine dehydrogenase oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.365.10:FF:000004">
        <property type="entry name" value="Xanthine dehydrogenase oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.390.50:FF:000001">
        <property type="entry name" value="Xanthine dehydrogenase oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.43.10:FF:000001">
        <property type="entry name" value="Xanthine dehydrogenase/oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.465.10:FF:000004">
        <property type="entry name" value="Xanthine dehydrogenase/oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.20.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.465.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.120">
        <property type="entry name" value="[2Fe-2S]-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1170.50">
        <property type="entry name" value="Aldehyde oxidase/xanthine dehydrogenase, a/b hammerhead"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.365.10">
        <property type="entry name" value="Aldehyde oxidase/xanthine dehydrogenase, molybdopterin binding domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.390.50">
        <property type="entry name" value="CO dehydrogenase flavoprotein, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.43.10">
        <property type="entry name" value="Uridine Diphospho-n-acetylenolpyruvylglucosamine Reductase, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002888">
        <property type="entry name" value="2Fe-2S-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036884">
        <property type="entry name" value="2Fe-2S-bd_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036010">
        <property type="entry name" value="2Fe-2S_ferredoxin-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006058">
        <property type="entry name" value="2Fe2S_fd_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000674">
        <property type="entry name" value="Ald_Oxase/Xan_DH_a/b"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036856">
        <property type="entry name" value="Ald_Oxase/Xan_DH_a/b_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016208">
        <property type="entry name" value="Ald_Oxase/xanthine_DH-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008274">
        <property type="entry name" value="AldOxase/xan_DH_MoCoBD1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046867">
        <property type="entry name" value="AldOxase/xan_DH_MoCoBD2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037165">
        <property type="entry name" value="AldOxase/xan_DH_Mopterin-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012675">
        <property type="entry name" value="Beta-grasp_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005107">
        <property type="entry name" value="CO_DH_flav_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036683">
        <property type="entry name" value="CO_DH_flav_C_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016166">
        <property type="entry name" value="FAD-bd_PCMH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036318">
        <property type="entry name" value="FAD-bd_PCMH-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016167">
        <property type="entry name" value="FAD-bd_PCMH_sub1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016169">
        <property type="entry name" value="FAD-bd_PCMH_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002346">
        <property type="entry name" value="Mopterin_DH_FAD-bd"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45444">
        <property type="entry name" value="XANTHINE DEHYDROGENASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45444:SF3">
        <property type="entry name" value="XANTHINE DEHYDROGENASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03450">
        <property type="entry name" value="CO_deh_flav_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00941">
        <property type="entry name" value="FAD_binding_5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01799">
        <property type="entry name" value="Fer2_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02738">
        <property type="entry name" value="MoCoBD_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20256">
        <property type="entry name" value="MoCoBD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000127">
        <property type="entry name" value="Xanthine_DH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01008">
        <property type="entry name" value="Ald_Xan_dh_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01092">
        <property type="entry name" value="CO_deh_flav_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54292">
        <property type="entry name" value="2Fe-2S ferredoxin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55447">
        <property type="entry name" value="CO dehydrogenase flavoprotein C-terminal domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47741">
        <property type="entry name" value="CO dehydrogenase ISP C-domain like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54665">
        <property type="entry name" value="CO dehydrogenase molybdoprotein N-domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56176">
        <property type="entry name" value="FAD-binding/transporter-associated domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56003">
        <property type="entry name" value="Molybdenum cofactor-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00197">
        <property type="entry name" value="2FE2S_FER_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51387">
        <property type="entry name" value="FAD_PCMH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 14" id="KW-0001">2Fe-2S</keyword>
    <keyword evidence="7 13" id="KW-0274">FAD</keyword>
    <keyword evidence="4" id="KW-0285">Flavoprotein</keyword>
    <keyword evidence="9 14" id="KW-0408">Iron</keyword>
    <keyword evidence="10 14" id="KW-0411">Iron-sulfur</keyword>
    <keyword evidence="6 14" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="3 14" id="KW-0500">Molybdenum</keyword>
    <keyword evidence="8" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="FAD-binding PCMH-type" evidence="15">
        <location>
            <begin position="197"/>
            <end position="382"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="12">
        <location>
            <position position="1207"/>
        </location>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="15"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="20"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="23"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="45"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="85"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>2</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="88"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>2</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="120"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>2</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="122"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>2</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <begin position="225"/>
            <end position="232"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <position position="328"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <position position="390"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="713"/>
        </location>
        <ligand>
            <name>Mo-molybdopterin</name>
            <dbReference type="ChEBI" id="CHEBI:71302"/>
        </ligand>
        <ligandPart>
            <name>Mo</name>
            <dbReference type="ChEBI" id="CHEBI:28685"/>
        </ligandPart>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="744"/>
        </location>
        <ligand>
            <name>Mo-molybdopterin</name>
            <dbReference type="ChEBI" id="CHEBI:71302"/>
        </ligand>
        <ligandPart>
            <name>Mo</name>
            <dbReference type="ChEBI" id="CHEBI:28685"/>
        </ligandPart>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="858"/>
        </location>
        <ligand>
            <name>Mo-molybdopterin</name>
            <dbReference type="ChEBI" id="CHEBI:71302"/>
        </ligand>
        <ligandPart>
            <name>Mo</name>
            <dbReference type="ChEBI" id="CHEBI:28685"/>
        </ligandPart>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <position position="860"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="1025"/>
        </location>
        <ligand>
            <name>Mo-molybdopterin</name>
            <dbReference type="ChEBI" id="CHEBI:71302"/>
        </ligand>
        <ligandPart>
            <name>Mo</name>
            <dbReference type="ChEBI" id="CHEBI:28685"/>
        </ligandPart>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001974"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006849"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022505"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022630"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022714"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022827"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023014"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00034078"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PIRSR" id="PIRSR000127-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PIRSR" id="PIRSR000127-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PIRSR" id="PIRSR000127-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS51387"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015297.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1274" mass="141209" checksum="9439D60808647017" modified="2019-05-08" version="1">TAAFAVRLSGTKYGCGIGGCGACTVMISTYEPTSKKIMHYPANSCLLPICSLHSSAVTTVEGVGSTKTRVHPIQERLAKCHGSQCGFCTPGMVMSMYALLRNHPEPSMEQITAALDGNLCRCTGYRPIIDTYKSFTGVGRKTNRSPFSKVSTNFECSVSGSLTKIRFSELLYPTQEFIFPPELMRMAQEQQKRTLVFRGERTTWISPTNLRELLELKAMYPKAPLIVGNTSVGLEMKRDGVHHPVFIHPTRIPDLHVVSNTNNGLVIGAACSLAQLRDILMKVVPELPEEKTKIFRALLQQLKTLAGEQIRSLASLGGHIVSRGSTWDLNPVLAVGNSVLSLASKDGTRQIPLNDQFLAGYENADLKPKEIIVSVFIPYSKKDDFVSAFRQAERRRNAVSVVNSGMRVLFKPNTDIIMDMNILYGGTGPITVSARKSCQRLIGRQWNEQMLDEACKLVLGEISLSPSAPGGKVEYRRTLLVSFLFKFYLEVLYGLNKMAWESPNWNGYEHSVVSVLSLGYHILIKQDVDPVQPPQDPVGRPIMHQSGVKHATGEAVYCNDIRAVDEELFLAVVTSSRAHAKIVTYYLNITPLPYTVTVHLYVLGVLQVICVGQIVCAVVAESAVQAKQGVEKMKIEYEDLEPILTIEVISQLSFFCLKKIEQGNIEEGFKSADEIIEGEIHIGGQEHFYLETNSVLVVPRGEDNEMDVHVSTQDAARVQEFVALALDVQSSKIMCHTKRVGGAFGGKITKPAIFAVVAAVAANKTGRPIRFTLERDEDMLITGGRHPFFGKYKVGFTNDGRITAADIEPYINAGCTLDDSEMVIEYVVLKTDNAYKIPNLRIRGRACKTNLPSNTAFRGFGFPQSGLFTESWITTVATQTGLPQEKIREINMYKEVNLTPFKEELDATNLVKCWKECMEKSEYYSRKTAVEEFNKQNYWKKKGIAIIPMKFSVGFNATQYHQAAALVHIYTDGSVLVTHGGIELGQGIHTKMLQIASRELKVPLSYVHFCETSTITVPNAIVTAGSIGSDVNGKAVQNACQILRKRLEPIINKNPEGKWEDWITEAYDQSVSLSATGYFRGYMVNMNWDTGEGQAFPYFVFGAACSEVEIDCLTGDHKNIRTDIVMDACFSINPAVDIGQIEGAFVQGVGLYTMEEIKFSPEGEQYTLGPDTYKIPAVCDIPEQFRVYLLPNSRNPIAIYSAKGIGEAGLFLGCSVFFAIWDAVAAVRKERGLTGTFTLNSPLTIERIRMACADNFTEMVRLDDHNGLFWPWNL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK54</accession>
    <name>A0A452HK54_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">NACHT domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Inflammasome</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the NLRP family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HK54"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017475.1">
        <property type="protein sequence ID" value="ENSGAGP00000015298.1"/>
        <property type="gene ID" value="ENSGAGG00000011535.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006954">
        <property type="term" value="P:inflammatory response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045087">
        <property type="term" value="P:innate immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001611">
        <property type="entry name" value="Leu-rich_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007111">
        <property type="entry name" value="NACHT_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041267">
        <property type="entry name" value="NLRP_HD2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050637">
        <property type="entry name" value="NLRP_innate_immun_reg"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041075">
        <property type="entry name" value="NOD1/2_WH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45690">
        <property type="entry name" value="NACHT, LRR AND PYD DOMAINS-CONTAINING PROTEIN 12"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45690:SF19">
        <property type="entry name" value="NACHT, LRR AND PYD DOMAINS-CONTAINING PROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13516">
        <property type="entry name" value="LRR_6"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05729">
        <property type="entry name" value="NACHT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17776">
        <property type="entry name" value="NLRC4_HD2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17779">
        <property type="entry name" value="WHD_NOD2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00368">
        <property type="entry name" value="LRR_RI"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52047">
        <property type="entry name" value="RNI-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50837">
        <property type="entry name" value="NACHT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9" id="KW-1271">Inflammasome</keyword>
    <keyword evidence="8" id="KW-0395">Inflammatory response</keyword>
    <keyword evidence="5" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="7" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="NACHT" evidence="10">
        <location>
            <begin position="83"/>
            <end position="217"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004110"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008665"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023198"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023233"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015298.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="813" mass="93216" checksum="210ACA9782DF433C" modified="2019-05-08" version="1">TDAKEFYQTVQKHSPLCSCSTPTGQKYRYTKLIIFNKYRHEKEREHEIMAVRRRHADIMSEQAHSSITMDTLFKPDEDGQMSQIIVLLGAAGTGKIMTARKIMLDWAGGRLYQEFDYVFYINGREMNLLTKQGNMADMILKHCPNNSGRIRHILENTEKLLFIIDGFDELRFSFDQPKDNLCSDPWEKKPVEIVLSSLFRKTVLSECYLIITTRPTALEELRQFLEYPHEKQATAAFNFVCENEILLTMCFVPIVCWIICTVLKQQMERGKDLAQTSKTLTEVYLLYLFSLVQSPSSSSKPHMQASLRRLWSLAVDGIWKQKILFEEVEIKKYGLDQVDSLQRIMVNIFQKDIDSECAYSFIPLTFQEFFAALFYVLEEDEETMKDSVIPNRDVKELLIHYGQSTSCLITVHFLFGLLNKNRMKDKDEKLGCKISHEIKPVLKTWVISSLPSNINQHYYLEKFYYLYEIQEENFVKNALEHLTNLETWEKIFTQMDQMVLSFCVKHYHQLESLLIHYCVFRFEDHEEEEFPSLPNCWHFAAVLRTDQSLTELDLGYNQYIGDSRVQLLCEGLKHSNCKLQRLGLWNCDLKGASCGYLAAVLRTSQSLTELELRDNVSLGDSGVQLLCEGLKHPNCKLQRLGLPSCSLTTAFRGNLSSVLSTSQTLTELDLKNNTIRDSGIQLLCEGLKHPNCKLQKLGLEKCDFTAACCGELSSALSTSQTLTELDLRENKVEDSRVKVLCEGLTQPACNLQKLSPSSSSQSALDLMILNNFILSGKFATSLFIPFSRSLKNMVNSTGPSTDPWRTLLSMSPL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK55</accession>
    <name>A0A452HK55_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">protein-serine/threonine phosphatase</fullName>
            <ecNumber evidence="3">3.1.3.16</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="8">
            <text>O-phospho-L-threonyl-[protein] + H2O = L-threonyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:47004"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="EC" id="3.1.3.16"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the protein-tyrosine phosphatase family.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.16" evidence="3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017483.1">
        <property type="protein sequence ID" value="ENSGAGP00000015305.1"/>
        <property type="gene ID" value="ENSGAGG00000011542.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003779">
        <property type="term" value="F:actin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004722">
        <property type="term" value="F:protein serine/threonine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030837">
        <property type="term" value="P:negative regulation of actin filament polymerization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
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        <property type="entry name" value="DSP_slingshot_2"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Protein phosphatase Slingshot homolog 2"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
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        <property type="entry name" value="DEK_C"/>
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        <property type="entry name" value="Dual-sp_phosphatase_cat-dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043587">
        <property type="entry name" value="Phosphatase_SSH-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
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        <property type="entry name" value="Tyr_Pase_AS"/>
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    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020422">
        <property type="entry name" value="TYR_PHOSPHATASE_DUAL_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45864:SF3">
        <property type="entry name" value="PROTEIN PHOSPHATASE SLINGSHOT HOMOLOG 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45864">
        <property type="entry name" value="SLINGSHOT PROTEIN PHOSPHATASE HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="DEK_C"/>
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        <property type="entry name" value="DSPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23040">
        <property type="entry name" value="PH_SSH1-like_1st"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00195">
        <property type="entry name" value="DSPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51998">
        <property type="entry name" value="DEK_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00383">
        <property type="entry name" value="TYR_PHOSPHATASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
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    </dbReference>
    <dbReference type="PROSITE" id="PS50054">
        <property type="entry name" value="TYR_PHOSPHATASE_DUAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="5" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="6" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="DEK-C" evidence="12">
        <location>
            <begin position="229"/>
            <end position="284"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="10">
        <location>
            <begin position="288"/>
            <end position="429"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="11">
        <location>
            <begin position="353"/>
            <end position="407"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="609"/>
            <end position="628"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="647"/>
            <end position="668"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="681"/>
            <end position="724"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="778"/>
            <end position="849"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="909"/>
            <end position="959"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="1163"/>
            <end position="1190"/>
        </location>
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    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="1234"/>
            <end position="1278"/>
        </location>
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    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="1351"/>
            <end position="1389"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="614"/>
            <end position="627"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="647"/>
            <end position="663"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="690"/>
            <end position="701"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="798"/>
            <end position="808"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="9">
        <location>
            <begin position="822"/>
            <end position="832"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
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            <begin position="909"/>
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            <begin position="1170"/>
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        <location>
            <begin position="1259"/>
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    </feature>
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        <location>
            <begin position="1360"/>
            <end position="1373"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00009580"/>
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            <dbReference type="ARBA" id="ARBA00023212"/>
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            <dbReference type="ARBA" id="ARBA00048336"/>
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    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
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        <source>
            <dbReference type="PROSITE" id="PS50054"/>
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        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
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        </source>
    </evidence>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015305.1"/>
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    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
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        </source>
    </evidence>
    <sequence length="1389" mass="154254" checksum="4773CC00144DDF5C" modified="2019-05-08" version="1">MEGVSAAWCIQWRYLRSISESFLTVKGAALFLPRGNGSSTPRINNRRNKHAGDLQQHLQAMFILLRPEDNIRLAVRLESTYQNRTRYMVVVSTNGRQDTEENIVLGMDFSSNDSSTCTMGLVLPLWSDTLIHLDGDGGFSVSTDNRVHVFKPVSVQAMWSALQSLHKACEVARSHNYYPGSLFLTWVSYYESHIDSDQSSVNEWNAMQDVQSHRPDSPALFTDVPTERERTERLIKTKLREIMMQKDLENITSKEIRTELEMQMVCNLREFKEFIDNEMIVILGQMDSPTQIFDHVFLGSEWNASNLEDLQNKGVRYILNVTREIDNFFPGIFEYHNIRVYDEEATDLLAYWNDTYKFISKAKKNGSKCLVHCKMGVSRSASTVIAYAMKEYGWDLDRAYEHVKERRAVTKPNPSFMRQLEEYQGILLASKQRHNKLWRSHSDSDLSEHHQPICKPGMELNKKEITTSAEQLSEPTANDNHQPALPVFSVGPHVGRLLQPDTDVIEELGAKERRIHLEFAARDFRAEQIEDELNLNNINRCPAGCCLEESEFPLDNCHSSEALVQLKEPLEMATAFPDLTVDDLETDALKTDLNVHLVPMEEFTACLKDLPRSPDQNSPSPQLSSHLETTDRIDFLSALEKFEELSQESRSHACSHTRMDEQGGGRNGVSKMSVLEVLPAGTAVDTQRNSSASSSPAAPEESSTDEEQPKDISELTTLGPLTRSHSENAISVKEIITEIESINQGIGPCQQKMEVSANHAQAPKRSTVHELPVEVVWTSENKPGKAEQSERACTAEQETAKDLPKPDQEGVPTFQLPTCASDVEESDPAEDQQEPRVQGLNPGSKWCPGSVKRATLEFEERLRQEQEHQHVPLICLLPTRKNSKNDSAAAELAPKGRTEELTWELDPAHGENESDQARAKSPAAEGALPHGPELPGGTHLSPPLAPCFVQPAQEPPLAEPGAEQEVLRTVVSLNGSGEPALPCHLPKRIEIIEYPHMGTPPDLRERDSSREQGTLENAAALLPVDENLNPSVCAETAPVHPDALPLRNPSVLEHKAIGQATFTLGSPEEEYGAAPINLDTSSFTGQVRRLPSAGLDDLWHQRVVHLEGVTEQSTSADDEPVAPSSLSRNGHVLCEASDSAFVELSNEDLDFLSDSTEAVDVSFLPGGLPHSSNSPSIKDSSEAPSSLKQRAKELEARVRQAGLTTPSQMKRSASLAKLGCLELSRDDLTKRELASSHASLTGPDLLPASPRTGQGKCDALPEHGLEGPPEKTCASSPLIRQDSKPTKHFVEQLKTTECIVQSQPVERPPVQYAKEFGSAPQNLYPSTDPRLTSSEEGLPLLQMQVLDSRLPAQSLAGAPRHQRSRTHPLRRLKMATDKRRTSNPFYNTM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK60</accession>
    <name>A0A452HK60_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">5-aminolevulinate synthase</fullName>
            <ecNumber evidence="15">2.3.1.37</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="15">5-aminolevulinic acid synthase</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="15">Delta-ALA synthase</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="15">Delta-aminolevulinate synthase</fullName>
        </alternativeName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="12">Catalyzes the pyridoxal 5'-phosphate (PLP)-dependent condensation of succinyl-CoA and glycine to form aminolevulinic acid (ALA), with CoA and CO2 as by-products. Contributes significantly to heme formation during erythropoiesis.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>succinyl-CoA + glycine + H(+) = 5-aminolevulinate + CO2 + CoA</text>
            <dbReference type="Rhea" id="RHEA:12921"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57292"/>
            <dbReference type="ChEBI" id="CHEBI:57305"/>
            <dbReference type="ChEBI" id="CHEBI:356416"/>
            <dbReference type="EC" id="2.3.1.37"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="13">
            <dbReference type="Rhea" id="RHEA:12922"/>
        </physiologicalReaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 14">
            <name>pyridoxal 5'-phosphate</name>
            <dbReference type="ChEBI" id="CHEBI:597326"/>
        </cofactor>
    </comment>
    <comment type="pathway">
        <text evidence="2 15">Porphyrin-containing compound metabolism; protoporphyrin-IX biosynthesis; 5-aminolevulinate from glycine: step 1/1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="15">Mitochondrion inner membrane</location>
            <topology evidence="15">Peripheral membrane protein</topology>
        </subcellularLocation>
        <text evidence="15">Localizes to the matrix side of the mitochondrion inner membrane.</text>
    </comment>
    <comment type="similarity">
        <text evidence="3 14">Belongs to the class-II pyridoxal-phosphate-dependent aminotransferase family.</text>
    </comment>
    <dbReference type="EC" id="2.3.1.37" evidence="15"/>
    <dbReference type="AlphaFoldDB" id="A0A452HK60"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017460.1">
        <property type="protein sequence ID" value="ENSGAGP00000015283.1"/>
        <property type="gene ID" value="ENSGAGG00000011529.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00251">
        <property type="reaction ID" value="UER00375"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005743">
        <property type="term" value="C:mitochondrial inner membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005759">
        <property type="term" value="C:mitochondrial matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003870">
        <property type="term" value="F:5-aminolevulinate synthase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030170">
        <property type="term" value="F:pyridoxal phosphate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048821">
        <property type="term" value="P:erythrocyte development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042541">
        <property type="term" value="P:hemoglobin biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006782">
        <property type="term" value="P:protoporphyrinogen IX biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="CDD" id="cd06454">
        <property type="entry name" value="KBL_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1150.10:FF:000029">
        <property type="entry name" value="5-aminolevulinate synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.640.10:FF:000006">
        <property type="entry name" value="5-aminolevulinate synthase, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1150.10">
        <property type="entry name" value="Aspartate Aminotransferase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.640.10">
        <property type="entry name" value="Type I PLP-dependent aspartate aminotransferase-like (Major domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010961">
        <property type="entry name" value="4pyrrol_synth_NH2levulA_synth"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015118">
        <property type="entry name" value="5aminolev_synth_preseq"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001917">
        <property type="entry name" value="Aminotrans_II_pyridoxalP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004839">
        <property type="entry name" value="Aminotransferase_I/II_large"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050087">
        <property type="entry name" value="AON_synthase_class-II"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015424">
        <property type="entry name" value="PyrdxlP-dep_Trfase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015421">
        <property type="entry name" value="PyrdxlP-dep_Trfase_major"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015422">
        <property type="entry name" value="PyrdxlP-dep_Trfase_small"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01821">
        <property type="entry name" value="5aminolev_synth"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13693:SF58">
        <property type="entry name" value="5-AMINOLEVULINATE SYNTHASE, ERYTHROID-SPECIFIC, MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13693">
        <property type="entry name" value="CLASS II AMINOTRANSFERASE/8-AMINO-7-OXONONANOATE SYNTHASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00155">
        <property type="entry name" value="Aminotran_1_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09029">
        <property type="entry name" value="Preseq_ALAS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53383">
        <property type="entry name" value="PLP-dependent transferases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00599">
        <property type="entry name" value="AA_TRANSFER_CLASS_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="11 15" id="KW-0012">Acyltransferase</keyword>
    <keyword evidence="9 15" id="KW-0350">Heme biosynthesis</keyword>
    <keyword evidence="10" id="KW-0472">Membrane</keyword>
    <keyword evidence="8 15" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="5" id="KW-0999">Mitochondrion inner membrane</keyword>
    <keyword evidence="6 14" id="KW-0663">Pyridoxal phosphate</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 15" id="KW-0808">Transferase</keyword>
    <keyword evidence="7" id="KW-0809">Transit peptide</keyword>
    <feature type="domain" description="5-aminolevulinate synthase presequence" evidence="18">
        <location>
            <begin position="86"/>
            <end position="188"/>
        </location>
    </feature>
    <feature type="domain" description="Aminotransferase class I/classII large" evidence="17">
        <location>
            <begin position="241"/>
            <end position="585"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="122"/>
            <end position="159"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001933"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005029"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00008392"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022792"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022898"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023133"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023315"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00045913"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00049013"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="RuleBase" id="RU003693"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="RuleBase" id="RU910713"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF00155"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF09029"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015283.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="636" mass="68640" checksum="F137B7F2AB1F0F80" modified="2019-05-08" version="1">MRATLGAGGRDRVLGYKTQASGAQPLWGLSLPPSCNMVPSLGLIEPVLICGPGRLRYGVGLARGGPGSLSGGVSQCWLFVLSAGQQRCPFLTRDPSIFLLKARPLLVSSAQRCPIMVARTLMNSSPDPQRGTDNPIPLAAGSVPQELETPETASSPSQTHCPFMELEVRDRRSRIVQRAGPEVQEDVKVYKAGSGAFSYDSFLEGRLEAKRRDHTYRVFKTVTRRADAFPFAREAAGPEVSVWCSNDYLAMSRHPRVLQAAMDALQHHGLGAGGTRNISGTSQYHVDLERELAQLHRKDAALLFSSCYVANDSTLFTLARMMPGCEIFSDAGNHASMIQGIRNSGVPKHVFRHNDPQHLAQLLGRSPPGIPKIVAFETVHSMDGGICPLEELCDVAHEHGALTFVDEVHAVGLYGARGAGIAERDGVLGKVDIVSGTLGKAFGCVGGYIASTAALVDTVRSYAAGFIFTTSLPPPVLAGALASLRVLAGPEGRGLRRAHQRNVKHMRQLLMDAGLPLVRCPSHIIPIRVGDAALNSQLCDLLLSQHNIYVQAINYPTVPRGEELLRLAPSPHHTPPMMDYFVERLVQGWQDVGLPLQPPASSACHFCQRPLHFALMSEWERDSFGSLGAQCVTSSA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK62</accession>
    <name>A0A452HK62_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">protein-tyrosine-phosphatase</fullName>
            <ecNumber evidence="2">3.1.3.48</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="10 17">
            <text>O-phospho-L-tyrosyl-[protein] + H2O = L-tyrosyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:10684"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="EC" id="3.1.3.48"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="3.1.3.48" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452HK62"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017489.1">
        <property type="protein sequence ID" value="ENSGAGP00000015311.1"/>
        <property type="gene ID" value="ENSGAGG00000004356.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000003">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase kappa isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000005">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase kappa isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000019">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase kappa isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000048">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase U isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057598">
        <property type="entry name" value="Fn3_PTPRU"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050348">
        <property type="entry name" value="Protein-Tyr_Phosphatase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000242">
        <property type="entry name" value="PTP_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016130">
        <property type="entry name" value="Tyr_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003595">
        <property type="entry name" value="Tyr_Pase_cat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19134">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR19134:SF209">
        <property type="entry name" value="RECEPTOR-TYPE TYROSINE-PROTEIN PHOSPHATASE KAPPA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23144">
        <property type="entry name" value="Fn3_PTPRU"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00102">
        <property type="entry name" value="Y_phosphatase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00700">
        <property type="entry name" value="PRTYPHPHTASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00194">
        <property type="entry name" value="PTPc"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00404">
        <property type="entry name" value="PTPc_motif"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00383">
        <property type="entry name" value="TYR_PHOSPHATASE_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50055">
        <property type="entry name" value="TYR_PHOSPHATASE_PTP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="9" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="5" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="8 11" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="3 11" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 11" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="420"/>
            <end position="441"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="14">
        <location>
            <begin position="58"/>
            <end position="154"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="14">
        <location>
            <begin position="155"/>
            <end position="261"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="12">
        <location>
            <begin position="566"/>
            <end position="820"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="13">
        <location>
            <begin position="740"/>
            <end position="811"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="12">
        <location>
            <begin position="852"/>
            <end position="1100"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="13">
        <location>
            <begin position="1028"/>
            <end position="1105"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00013064"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00051722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015311.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1116" mass="125876" checksum="67B460DF8232A75B" modified="2019-05-08" version="1">GLFLSTWPVHLSSGRFPERSQWCTVGYRPEASTVDLWSTETGLKSPLYLYKGPQPMRTPKTLKIAEIRARHIAVDWESLGYNITRCHTFNVTICYHYFHGHNENKADCLDMDPKAPQHVVDHLPPYTNVSLKMILTNPEGRKESEETIIQTDEDVPGPVPAKSLKGTPFEDKIFLNWKEPVDPNGIITQYEVSYSSIRSFDPAVLVAGPPQTVSKVWNSTHHVFSHLHPGTTYQFFIRASTVKGFGPATAINVTTNISAPTLPDYEGVDASLNETATTITVLLRPAQAKGAPISAYQIVVEELHPHRTKRETGTMECYQVPVTYQNALSGSSPYYFAAELPPGNLPEPAPFTVGDNRTYQGFWNPPLAPRKGYNIYFQAMSSVEKETKTQCVRIATKAAATEEPEVIPDPAKQTDRVVKIAGISAGILVFILLLLVVILIVKKSKLAKKRKDAMGNTRQEMTHMVNAMDRSYADQSTLHAEDPLSITFMDSHNFSPRLPNDPLVPTAVLDENHSATAESSRLLDVPRYLGEGTESPYQTGQLHPAIRVADLLQHINLMKTSDSYGFKEEYESFFEGQSASWDVAKKDQNRTKNRYGNIIAYDHSRVILQPVEDDPSSDYINANYIDGYQRPSHYIATQGPVHETVYDFWRMIWQEQSACVVMVTNLVEVGRVKCYKYWPDDTEVYGDFKVTCVEMEPLAEYVVRTFTLERRGYNEIRELKQFHFTGWPDHGVPYNATGLLSFIRRVKLSNPPSAGPIVVHCSAGAGRTGCYIVIDIMLDMAEREGVVDIYNCVKALRSRRINMVQTEEQYIFIHDAILEACLCGETAIPVCEFKAAYFDMIRIDSQTNSSHLKDEFQTLNSVTPRLQAEDCSIACLPRNHDKNRFMDMLPPDRCLPFLITIDGESSNYINAALMDSYRQPAAFIVTQHPLPNTVKDFWRLVYDYGCTSVVMLNEVDLAQGCPQYWPEEGMLRYGPIQVECMSCSMDCDVINRIFRICNLTRPQEGYLMVQQFQYLGWASHRDVPGSKRSFLKLILQVEKWQEECEEGEGRTIIHCLNGGGRSGMFCAIGIVAEMVKRQNVVDVFHAVKTLRNSKPNMVETPVNIPPLPASGFCNLK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK63</accession>
    <name>A0A452HK63_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">protein-serine/threonine phosphatase</fullName>
            <ecNumber evidence="3">3.1.3.16</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="8">
            <text>O-phospho-L-threonyl-[protein] + H2O = L-threonyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:47004"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="EC" id="3.1.3.16"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the protein-tyrosine phosphatase family.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.16" evidence="3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017485.1">
        <property type="protein sequence ID" value="ENSGAGP00000015307.1"/>
        <property type="gene ID" value="ENSGAGG00000011542.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003779">
        <property type="term" value="F:actin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004722">
        <property type="term" value="F:protein serine/threonine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030837">
        <property type="term" value="P:negative regulation of actin filament polymerization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd14569">
        <property type="entry name" value="DSP_slingshot_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd11652">
        <property type="entry name" value="SSH-N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000004">
        <property type="entry name" value="Protein phosphatase Slingshot homolog 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014876">
        <property type="entry name" value="DEK_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000340">
        <property type="entry name" value="Dual-sp_phosphatase_cat-dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043587">
        <property type="entry name" value="Phosphatase_SSH-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043588">
        <property type="entry name" value="SSH-N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016130">
        <property type="entry name" value="Tyr_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000387">
        <property type="entry name" value="Tyr_Pase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020422">
        <property type="entry name" value="TYR_PHOSPHATASE_DUAL_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45864:SF3">
        <property type="entry name" value="PROTEIN PHOSPHATASE SLINGSHOT HOMOLOG 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45864">
        <property type="entry name" value="SLINGSHOT PROTEIN PHOSPHATASE HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08766">
        <property type="entry name" value="DEK_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00782">
        <property type="entry name" value="DSPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23040">
        <property type="entry name" value="PH_SSH1-like_1st"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00195">
        <property type="entry name" value="DSPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51998">
        <property type="entry name" value="DEK_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00383">
        <property type="entry name" value="TYR_PHOSPHATASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50056">
        <property type="entry name" value="TYR_PHOSPHATASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50054">
        <property type="entry name" value="TYR_PHOSPHATASE_DUAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="5" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="6" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="DEK-C" evidence="12">
        <location>
            <begin position="231"/>
            <end position="286"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine-protein phosphatase" evidence="10">
        <location>
            <begin position="290"/>
            <end position="431"/>
        </location>
    </feature>
    <feature type="domain" description="Tyrosine specific protein phosphatases" evidence="11">
        <location>
            <begin position="355"/>
            <end position="409"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="611"/>
            <end position="630"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="649"/>
            <end position="670"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="683"/>
            <end position="726"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="780"/>
            <end position="851"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="911"/>
            <end position="961"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="1165"/>
            <end position="1192"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="1236"/>
            <end position="1280"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="1353"/>
            <end position="1391"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="616"/>
            <end position="629"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="649"/>
            <end position="665"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="692"/>
            <end position="703"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="800"/>
            <end position="810"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="9">
        <location>
            <begin position="824"/>
            <end position="834"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="911"/>
            <end position="920"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="1172"/>
            <end position="1190"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="1261"/>
            <end position="1271"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="9">
        <location>
            <begin position="1362"/>
            <end position="1375"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009580"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00013081"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00048336"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS51998"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015307.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1391" mass="154336" checksum="5423ABD66318BEAF" modified="2019-05-08" version="1">MIPYFSDNAVISQNAINQLISESFLTVKGAALFLPRGNGSSTPRINNRRNKHAGDLQQHLQAMFILLRPEDNIRLAVRLESTYQNRTRYMVVVSTNGRQDTEENIVLGMDFSSNDSSTCTMGLVLPLWSDTLIHLDGDGGFSVSTDNRVHVFKPVSVQAMWSALQSLHKACEVARSHNYYPGSLFLTWVSYYESHIDSDQSSVNEWNAMQDVQSHRPDSPALFTDVPTERERTERLIKTKLREIMMQKDLENITSKEIRTELEMQMVCNLREFKEFIDNEMIVILGQMDSPTQIFDHVFLGSEWNASNLEDLQNKGVRYILNVTREIDNFFPGIFEYHNIRVYDEEATDLLAYWNDTYKFISKAKKNGSKCLVHCKMGVSRSASTVIAYAMKEYGWDLDRAYEHVKERRAVTKPNPSFMRQLEEYQGILLASKQRHNKLWRSHSDSDLSEHHQPICKPGMELNKKEITTSAEQLSEPTANDNHQPALPVFSVGPHVGRLLQPDTDVIEELGAKERRIHLEFAARDFRAEQIEDELNLNNINRCPAGCCLEESEFPLDNCHSSEALVQLKEPLEMATAFPDLTVDDLETDALKTDLNVHLVPMEEFTACLKDLPRSPDQNSPSPQLSSHLETTDRIDFLSALEKFEELSQESRSHACSHTRMDEQGGGRNGVSKMSVLEVLPAGTAVDTQRNSSASSSPAAPEESSTDEEQPKDISELTTLGPLTRSHSENAISVKEIITEIESINQGIGPCQQKMEVSANHAQAPKRSTVHELPVEVVWTSENKPGKAEQSERACTAEQETAKDLPKPDQEGVPTFQLPTCASDVEESDPAEDQQEPRVQGLNPGSKWCPGSVKRATLEFEERLRQEQEHQHVPLICLLPTRKNSKNDSAAAELAPKGRTEELTWELDPAHGENESDQARAKSPAAEGALPHGPELPGGTHLSPPLAPCFVQPAQEPPLAEPGAEQEVLRTVVSLNGSGEPALPCHLPKRIEIIEYPHMGTPPDLRERDSSREQGTLENAAALLPVDENLNPSVCAETAPVHPDALPLRNPSVLEHKAIGQATFTLGSPEEEYGAAPINLDTSSFTGQVRRLPSAGLDDLWHQRVVHLEGVTEQSTSADDEPVAPSSLSRNGHVLCEASDSAFVELSNEDLDFLSDSTEAVDVSFLPGGLPHSSNSPSIKDSSEAPSSLKQRAKELEARVRQAGLTTPSQMKRSASLAKLGCLELSRDDLTKRELASSHASLTGPDLLPASPRTGQGKCDALPEHGLEGPPEKTCASSPLIRQDSKPTKHFVEQLKTTECIVQSQPVERPPVQYAKEFGSAPQNLYPSTDPRLTSSEEGLPLLQMQVLDSRLPAQSLAGAPRHQRSRTHPLRRLKMATDKRRTSNPFYNTM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMT3</accession>
    <name>A0A452IMT3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Mannose receptor C-type 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="14">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000033118.1">
        <property type="protein sequence ID" value="ENSGAGP00000029156.1"/>
        <property type="gene ID" value="ENSGAGG00000020766.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030246">
        <property type="term" value="F:carbohydrate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006897">
        <property type="term" value="P:endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd23407">
        <property type="entry name" value="beta-trefoil_Ricin_MRC1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00037">
        <property type="entry name" value="CLECT"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="CDD" id="cd00062">
        <property type="entry name" value="FN2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.80.10.50:FF:000032">
        <property type="entry name" value="macrophage mannose receptor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000014">
        <property type="entry name" value="Macrophage mannose receptor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000016">
        <property type="entry name" value="macrophage mannose receptor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000023">
        <property type="entry name" value="Macrophage mannose receptor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000031">
        <property type="entry name" value="macrophage mannose receptor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000025">
        <property type="entry name" value="Mannose receptor C-type 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000030">
        <property type="entry name" value="Mannose receptor C-type 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000027">
        <property type="entry name" value="Mannose receptor, C type 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.80.10.50">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.10.10">
        <property type="entry name" value="Fibronectin, type II, collagen-binding"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.100.10">
        <property type="entry name" value="Mannose-Binding Protein A, subunit A"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001304">
        <property type="entry name" value="C-type_lectin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016186">
        <property type="entry name" value="C-type_lectin-like/link_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050111">
        <property type="entry name" value="C-type_lectin/snaclec_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018378">
        <property type="entry name" value="C-type_lectin_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016187">
        <property type="entry name" value="CTDL_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000562">
        <property type="entry name" value="FN_type2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036943">
        <property type="entry name" value="FN_type2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013806">
        <property type="entry name" value="Kringle-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035992">
        <property type="entry name" value="Ricin_B-like_lectins"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000772">
        <property type="entry name" value="Ricin_B_lectin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22803">
        <property type="entry name" value="MANNOSE, PHOSPHOLIPASE, LECTIN RECEPTOR RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24562">
        <property type="entry name" value="CysR_MRC2_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00040">
        <property type="entry name" value="fn2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00059">
        <property type="entry name" value="Lectin_C"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00013">
        <property type="entry name" value="FNTYPEII"/>
    </dbReference>
    <dbReference type="SMART" id="SM00034">
        <property type="entry name" value="CLECT"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SMART" id="SM00059">
        <property type="entry name" value="FN2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00458">
        <property type="entry name" value="RICIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56436">
        <property type="entry name" value="C-type lectin-like"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57440">
        <property type="entry name" value="Kringle-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50370">
        <property type="entry name" value="Ricin B-like lectins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00615">
        <property type="entry name" value="C_TYPE_LECTIN_1"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50041">
        <property type="entry name" value="C_TYPE_LECTIN_2"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51092">
        <property type="entry name" value="FN2_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50231">
        <property type="entry name" value="RICIN_B_LECTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8" id="KW-0106">Calcium</keyword>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0254">Endocytosis</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6" id="KW-0430">Lectin</keyword>
    <keyword evidence="10 15" id="KW-0472">Membrane</keyword>
    <keyword evidence="12" id="KW-0675">Receptor</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <keyword evidence="4 15" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9 15" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="1362"/>
            <end position="1385"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-II" evidence="17">
        <location>
            <begin position="161"/>
            <end position="212"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="16">
        <location>
            <begin position="226"/>
            <end position="341"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="16">
        <location>
            <begin position="369"/>
            <end position="486"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="16">
        <location>
            <begin position="511"/>
            <end position="626"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="16">
        <location>
            <begin position="655"/>
            <end position="774"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="16">
        <location>
            <begin position="791"/>
            <end position="907"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="16">
        <location>
            <begin position="935"/>
            <end position="1057"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="16">
        <location>
            <begin position="1079"/>
            <end position="1190"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="16">
        <location>
            <begin position="1219"/>
            <end position="1336"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022583"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022734"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50041"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS51092"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029156.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="20">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1431" mass="162896" checksum="C22F8C1D30B1D61A" modified="2019-05-08" version="1">ILPSITTFTIYFFSFCFTGSGIFLIHNEDHKLCVKAQSSNSVIIVTCNQDDDSQKFRWVSDHQILSMALKLCLGVPSMKDQAAITLYPCNKTSELQQWECRNKTLLAIQGEDLFFNSAHGEERKVMLSKGSGSWNRWKIHGTTENLCSQGYEDMFTLLGNAFGAPCVFPFKFNSTWYAECTKDRKAGNKQGFPWCATSTDFDEDSLFGYCPLKDNIDKDFWITNPFTGTHYQINSNSALTWHQARKSCQQQNAELLSITELHEQTYLTGLFGRANSELWIGLNRLNSNGGWQWIGGSPFRYLNWAPGSPSLESEKICGLLQSGNGKWKNQQCDQKLGYICKKGNSSLDSFIIPSGDFRPIKCPSGWIPYSGHCYMLYRAPKIWKDALSSCRKEGGDLASIHNIEEYSFTVSQLGYKPTDELWIGLNDLKIHMYFDWSDGTPVTYTKWLHEEPSHANNRKEDCVSMKGEDGYWADDVCEKKLGYICKMKPLAEESEKVEVIIPGCQKGWIRHGFYCYSVGQTSVTFSEAKKICEGNKGYLVTVKDRYEQAFLTVLIGFSPVKYFWIALSNVEEQGTFKWADGEAVLFTHWNSGMPGREPGCVAMRTGTTAGLWDVVNCEEKSMFLCKQLVEGATPPPVPTTTPAPSCPDGWHSRAHSSSCFKHFQNGRKHMKTWFDARDFCRAIGGDLASIHSEEDQNVINSLAHYYQSYWMGMSAFDPDGGFTWSDGSPVNYENWADGEPNNYDGNEKCAMFNEYTDMQWNDMFCEKFGDWTCQIKKEYTYKVSEDGWIIYEDKAYYFSHETLPMEKAREYCKKNSGDLVVIEGESKRKFLWKYSFYNDFRESLYIGLSVSLDKKFSWIDGTPLNYVAWSPNEPNFANNDENCVVMYTQTGMWNDLNCGAPNSFICERLNNITHSTVAPTSPAPLGGCPQNWFLFNNKCFKIFGSSENESLTWHAARTACINLEGNLASIPSEEVQAFLITHLKDASTDTWIGLNDVNSEYTFLWTDGSGVSYTSWAKGFPNYRSEADCVMMIKDPIEEAGKWKDEDCQMNKSYICQRNADPKLYNFQTTVPVSGFTLYGNSSYSLISSRMNWEEAQKNCKAEYSELASILDPYSQSFLWLQVLKIGQPVWIGLNSNVTSGFYKWTDNWRIKYSKWASEEPKQNLACVYLDVDGTWKTGSCDENYCSVCKRSDVLAPTELPQLPGKCPTSDIRRSWIPFHGHCYYIESSEMESWPQSSMECVRLGASLVSIENMAEMNFLLHHLDAFASDSRQFWIGMYKNVDGEWTWADNSAVEFVNWKEREPTDGYHEYCVDMDASDGGWRSYYCSADKRFICKIPKITEVEPSEKLSDNKDEAVPAHHLSGMAVILVLLILVGAGLAAYIFFKKKRHNQLPIDVTFDNTLYCNRDAVPAASDSKYLVANIEQNEQAML</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMU1</accession>
    <name>A0A452IMU1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Atos homolog protein A</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="4">Transcription regulator that syncronizes transcriptional and translational programs to promote macrophage invasion of tissues.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the ATOS family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMU1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033163.1">
        <property type="protein sequence ID" value="ENSGAGP00000029200.1"/>
        <property type="gene ID" value="ENSGAGG00000021121.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033473">
        <property type="entry name" value="Atos-like_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025261">
        <property type="entry name" value="Atos-like_cons_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051506">
        <property type="entry name" value="ATOS_Transcription_Regulators"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13199:SF13">
        <property type="entry name" value="ATOS HOMOLOG PROTEIN A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13199">
        <property type="entry name" value="GH03947P"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13889">
        <property type="entry name" value="Chromosome_seg"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13915">
        <property type="entry name" value="DUF4210"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01177">
        <property type="entry name" value="DUF4210"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0539">Nucleus</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Atos-like conserved" evidence="6">
        <location>
            <begin position="889"/>
            <end position="947"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00034497"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00037191"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00040290"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="SMART" id="SM01177"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029200.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1087" mass="122321" checksum="2F0998C107A9FCE3" modified="2019-05-08" version="1">MKPERDPLDEYFEYEAEEFLVSLALLITEGRTPEFSVKGRTEGFHCPPAQSSQPLTSKHECTDKLAQCRQARRTRSEVMLLWKNNIPIMIEVMLLPDCCYSDEGPTTEGSDLNDPAIKQDALLLERWILEPVPRQSGDRFIEEKTLLLAVRSFVFFSQLSAWLSVSHGAVPRNILYRVSAADVDLQWTFSQTPTEHVFPVPNVSHNVALKVSVQSLPRQSNYPVLTCSIHTSLRFYEKGIQEHNVHQHCDSIEREQCSVSSSQRSCGKQTWTLIPEGLLNTKISCEFSTSVRNIKLYPATGLGSAFGTSQSKVQCYNAMGNDKTQPHETSVRTFKSFSLIDPQVPNSHCSHQRIGETTPLIGSLLQERQEVIARIAQHLIHCDPSTSLVTGCPFSMHETSSVTSKVFWNTYEDENLLKKGKETSSVSTANTDITLSEDCSEDKTRAIPENPLVDCHVPMNHCSRQSLGETNSLIGSLLLERQEAIARIAQHLIHCDPATSHITRHPVNVHESNSITSKVFQSSYEDENLLKKGKETSSISFSKSDVTIEDSSKVKTRTPDTPLTSYRLDGELKASPKPEARRKLILAKSSEVVQNTFQQTPNKTTSSFISISISCSKENKSELPGKWEAVLSGCPHKDQMTKKDIGKQCSNFNTPDEQICKNKLKERTITSENSNGDCLINPQLDKSKIFEGTKKATLMQASNILHKNELKCLDKDSKKPNICEQNTQLVSIENYLNKDYDSFKCKNKQDKLKTAHDENEDPTDYEFQSNSQKKSVEDYLARCERLKNADVLRMPPLKHSSIWRKNNFHSVDGTVTKAFHPRTGLPLLSSPVPQRKTQSGCFDLDSSLLRLKCLSTRSPQQCINTDSDPEGHGKPFLSSSAPPITSLSLLGNFEESVLNYRLDPLGIVDGFTAEVGASGVFCPTHMTLPVEVSFYSVSDDNAPSPYMGVITLESLGKRGYRVPPSGTIQVTLFNPNKTVVKMFVVIYDLREMPANHQTFLRQRTFSIPVRREVKRSINKENIRQTEERLLRYLIHLRFQSSKSGKIYLHRDVRLLFSRKSMEVDSGAAYELKSYTESPTNPQFSPRC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2025-10-08" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMU2</accession>
    <name>A0A452IMU2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Nucleoporin NUP188</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
            <location evidence="1">Nuclear pore complex</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="8">Belongs to the Nup188 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000033126.1">
        <property type="protein sequence ID" value="ENSGAGP00000029164.1"/>
        <property type="gene ID" value="ENSGAGG00000021096.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000033128.1">
        <property type="protein sequence ID" value="ENSGAGP00000029166.1"/>
        <property type="gene ID" value="ENSGAGG00000021096.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0044611">
        <property type="term" value="C:nuclear pore inner ring"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017056">
        <property type="term" value="F:structural constituent of nuclear pore"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051028">
        <property type="term" value="P:mRNA transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006606">
        <property type="term" value="P:protein import into nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006405">
        <property type="term" value="P:RNA export from nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018864">
        <property type="entry name" value="Nucleoporin_Nup188_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044840">
        <property type="entry name" value="Nup188"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048883">
        <property type="entry name" value="Nup188_N-subdom_III"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31431:SF1">
        <property type="entry name" value="NUCLEOPORIN NUP188"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31431">
        <property type="entry name" value="NUCLEOPORIN NUP188 HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10487">
        <property type="entry name" value="Nup188_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21093">
        <property type="entry name" value="Nup188_N-subdom_III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21094">
        <property type="entry name" value="Nup188_SH3-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0509">mRNA transport</keyword>
    <keyword evidence="6" id="KW-0906">Nuclear pore complex</keyword>
    <keyword evidence="7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="4" id="KW-0653">Protein transport</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0811">Translocation</keyword>
    <keyword evidence="2" id="KW-0813">Transport</keyword>
    <feature type="domain" description="Nucleoporin Nup188 N-terminal" evidence="11">
        <location>
            <begin position="4"/>
            <end position="360"/>
        </location>
    </feature>
    <feature type="domain" description="Nucleoporin Nup188 N-terminal subdomain III" evidence="12">
        <location>
            <begin position="409"/>
            <end position="817"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1131"/>
            <end position="1152"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1387"/>
            <end position="1414"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1576"/>
            <end position="1600"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="1137"/>
            <end position="1152"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="1396"/>
            <end position="1406"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="1580"/>
            <end position="1593"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004567"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022816"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023010"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023132"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00038387"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00040174"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF10487"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF21093"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029166.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1618" mass="181604" checksum="A919A83DAA76152B" modified="2019-05-08" version="1">MLKLADYYYEERMCILRCVLNLLTYFQDERHPYMAQYSQCVEKLDKELVPNYRKQFEDLYKAEAPTWETHGNLMTERQVSRWFVQCLREQSMLLEVIFLYYAYFEMAPDELLSFTRLFKEQGFGSRQTNRHLVDESMDHLVDRIGYFSALILVEGMEIDSLHERVLDDRTEEHKFASNGRMHQEMDHLLLTLGDISHHAPVLLAWALLRHTVTPEESTNAIRRMGSTAIQLNIFQYLTKLLRSLSSGGKNCTASTACMCIYGLLSFVLTSLELHTLGNQQDIIDAACEVLAASSIPQLFWRTEPTAGLGIILDSVCGMFPHLLSPLLQLLKALVSDKSTAKKVYSFLDKMSFYIELYKHKPPDVISHEDGTLWRRQVPKLLYPLGLGQTNLRIPQGTVGQVMLDDRAYLVRWEYSYSSWTLFTCEIEMLLHVVSTADVIQHCQRVKPVIDLVHKVISTDVSIADCLLPITSRIYMLLQRLTSVISPPVDVIASCVNCLTVLAARMPAKVWTDLHHTGFLPFVANPVSSMSHMISAEGMNAGGYGNLLMSVEQPQGEYGVTISFLHLVTTLVRGQLGSTQSQGLMPCIVFVLKEMLPNYHKWRYNSHGVRERIGCLILQLIHAILNLCPEMDSHSSNAPSLQSLCIFSLANTEAGQAVINIMGIGVDTIDMVMASQSCSDESLGQGQLLIQTVKLAFSVTNNVIRLKSPSSVVSPLEQALTQHGAHGNNLIAVLAKYIYHIHDPALPRLAIQLLKRLATVAPMSVYACLGSDAAAIRDAFLTRLQSKIEDMRIKVMILEFLTVAVETQPGLIELFLNLEVKDGSDGSKEFSLGEWSCLQVVLELIDSKQQERYWCPPLLHRAAIAFLHALWQDRRDSAMLVLRTKPKFWENLTNPLFGTLSPPSETSELSVLDTCALIMKIICLEIYYVVKGSLDQSLKDTLKKFSTEKRFAYWSGYVKSLALHMAETEGSSCTSLTEYQMLISAWRMLLIISTSHADIMHLTETAVRQQLFLEVLSGTKALLLVPMSVLCLQLGSMLCTLLLILLRQWKSELGSVDKIINSLTQILEGVLQADQQMMEKTKAKVFSALITVLQMKEMKVSEIPQYSQLVMSVCETLQDEVIVLFDRTRHSLTSGDSADDKDSMETDDACRGKHKDQRDGVCVLGLHLAKELCEVDEDGDYWLQVTRKVPMLPTIFTALEISLRVKQNLHFTEASLHLLLTLARTQQGAAAVASAGVTQSICLSLLSVYQLSTNGTIQTPASSRKSLDAPSWPGVYRLSISLMERLLKTLRYNFLTEALDFVGVHQERILQCLNAVRTVQSLACLEEADHTVGFILQLSNFMKEWHFHLPQLLRDIQVNLCYLCQACTSLLHSRKMLQHYLQTKNGDSLPSAVTPRVQRTPQTSSKHPSPESEAAEQKALRTVQYGLLKILSKSLAALRHFTPDVCQILLDQSLDLAEYNLLFVLSFTTPAFDSDVAPSFGTLLATVNVALNMLGELDKKKEPFAQAAGLNTQEETKTLKSLLMFTMENCFYLLISQAVRYLRDPAVHPRDKQRMKQELSSELSTLLSSLSRYFRRGGPSSPASGVLPSSQGKSASKIGPEGQEPLVQLVQAFVRHVQR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMU4</accession>
    <name>A0A452IMU4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">RING-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000033389.1">
        <property type="protein sequence ID" value="ENSGAGP00000029403.1"/>
        <property type="gene ID" value="ENSGAGG00000021257.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000245">
        <property type="term" value="P:spliceosomal complex assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057031">
        <property type="entry name" value="SFR19-like_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001841">
        <property type="entry name" value="Znf_RING"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017907">
        <property type="entry name" value="Znf_RING_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47048">
        <property type="entry name" value="PROTEIN SCAF11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47048:SF1">
        <property type="entry name" value="PROTEIN SCAF11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23030">
        <property type="entry name" value="SCAF11-like_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57850">
        <property type="entry name" value="RING/U-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00518">
        <property type="entry name" value="ZF_RING_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50089">
        <property type="entry name" value="ZF_RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="1" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0862">Zinc</keyword>
    <keyword evidence="2 4" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="RING-type" evidence="7">
        <location>
            <begin position="41"/>
            <end position="89"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="257"/>
            <end position="440"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="624"/>
            <end position="1034"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="1049"/>
            <end position="1092"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="1157"/>
            <end position="1177"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="5">
        <location>
            <begin position="445"/>
            <end position="485"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="295"/>
            <end position="306"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="312"/>
            <end position="323"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="408"/>
            <end position="425"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="426"/>
            <end position="439"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="624"/>
            <end position="633"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="645"/>
            <end position="660"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="6">
        <location>
            <begin position="682"/>
            <end position="692"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="714"/>
            <end position="750"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="760"/>
            <end position="782"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="789"/>
            <end position="808"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="6">
        <location>
            <begin position="809"/>
            <end position="830"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="837"/>
            <end position="862"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="870"/>
            <end position="909"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="923"/>
            <end position="944"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="992"/>
            <end position="1001"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="1004"/>
            <end position="1017"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="1049"/>
            <end position="1059"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="6">
        <location>
            <begin position="1163"/>
            <end position="1173"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029403.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1346" mass="152496" checksum="9D6D69BA08B97A84" modified="2019-05-08" version="1">MKNKKKCIPNIEDQKHEDMEGEENEESCSCPAVLYDEADRCPICLNCLLEQEVGFPENCRHFFCMPCILKWAEVQEKIQLREKGEKENCCCNKEKCYHMNTKIFMRRDTCADKGPLIEKLCKDLKNYSGISDERNKETASVKINKPRRQTCSNECFRNHFCNRSPSTGESFSSYRNDCTELTEIHTVIRQKRRELELSSSSVLARVQRNSFMFCEAEAETFLLASSVMPETVFPTHTTPLEKFDSLGKRYAVACTQEGEEKKQASGTSGTRGTRKKTANTTPRRSARNRKTETMSQSRSSPRSSNSGCDAPNDSNSTINVSSADSEKPVSKRKSKRGVKQQVPLVKKKLRSSLRSEKSSSDSVEDEVAESDTPPVLDKDHLSDIESSNSCLMHKNNAETESANGLESCSEHAESEDLTRDCDKEQNTLANSDSGSSIQESPVLALEAQKCEIKDVNKNVTDFENKDVCENILEQIEQSVDNHRDQLLEDSITPTKVDQTLDELPEYPESVAKVVEPLDSPRNELTEHPESIEIDDSEDKEQAANCNTSNIANIKTVQDTETAVLINNISLDINELDQSLEEEPIPVCEEARTTELLRVNVEQSNKHFNEDNNEMIPMECDSFCSDQAESQIEQPSPVDSVKQEEANSLQQDSENCVSYSGLSDKKDEASSQETECSTEFKKDKKTRTRRSRFHSPSTTWSPSRREAGKSQSPSPKRETVREGRRSRSPDRETAREGRRSPSQSPKRETVRGGRKTPSRSPKRETDREGRRSPSRSRVKDSSPRQKSRSQSKDRDSDRDGQRRDHDRERRNRRWSRSRSRSRSRSRSRTRSKGPSFIRNERDGHSPRWKERDGHSPRWKERWTNDSWRSPRGNDRYKRSEQEKQNENIQKDNTEKSTDEQCSSEKHKNDCPDWVMERINSVPETRSRERDKFKDPHWEENRHDNSGHSWNKNFGSGWISNRSRGGRNRGGRGRGGFMYGDQNEDHWQNRKPLSGNSNSSGNETARFPEHQPYRRKSEQDFSFDTPADRSGWTSASSWAVRKTLPADVQSYYSRRGRNSSSPQSGWMRQEDETSEQDPNLKDQTNQPGDGSQLPINMMQQQMNVMPQVNAQHQPMNIFPYPVGVHAPIMNIQHNPFNIHPPLPMHLHTGVPLIQVAAPTSVSQGLPPPPPPPPPSQQVSYIASQPDGKQLQGIPSAPHVSNNMSAPLLPAPTAVLGNVGTVQGPSSGNATSSSHIKTSNAAVKFAEGKVSVTVEASADSSKKDQKLLIQEKAAQEVKLAIKPFYQNKDITKEEYKEIVRKAVDKVCHSKSGEVNSAKVANLVKAYVDKYKHSRKKNPEEVVSAERKRE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMV6</accession>
    <name>A0A452IMV6_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="7">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the major facilitator superfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMV6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033145.1">
        <property type="protein sequence ID" value="ENSGAGP00000029183.1"/>
        <property type="gene ID" value="ENSGAGG00000021090.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1250.20:FF:000508">
        <property type="entry name" value="Sodium-dependent glucose transporter 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1250.20">
        <property type="entry name" value="MFS general substrate transporter like domains"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036259">
        <property type="entry name" value="MFS_trans_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23121">
        <property type="entry name" value="SODIUM-DEPENDENT GLUCOSE TRANSPORTER 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23121:SF9">
        <property type="entry name" value="SODIUM-DEPENDENT GLUCOSE TRANSPORTER 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103473">
        <property type="entry name" value="MFS general substrate transporter"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 6" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 6" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 6" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="36"/>
            <end position="58"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="65"/>
            <end position="81"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="87"/>
            <end position="114"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="126"/>
            <end position="146"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="187"/>
            <end position="209"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="230"/>
            <end position="257"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="302"/>
            <end position="320"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="326"/>
            <end position="348"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="360"/>
            <end position="380"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="392"/>
            <end position="412"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008335"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029183.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="521" mass="56056" checksum="8A88C2AD0D3D9B94" modified="2019-05-08" version="1">MNKQNIPTVFQGMSIAVLGPTFQDLAANVNKNVSDIYYIFMGRSLGYLGGSVIGGIIFDCMNPHLLLGLTMLGTAVGLYAIPWCKRALLLTALMSVIGTSVGVLDTGGNVLALNIWGAEAGPHMQALHFSFALGAVVAPILTKMALSSFISPNFQTEDEKTNHSVPSDIPNALFGSALKLHSNVSFMWSYIVIGTYILIVSLLFVVLYSRTGLARDKAKSSLQKCQIAKYHYALLILLFLFFFCYVGAEVTYGSYIFNYAEVYADMKESEAASLNSLFWGAFAACRGLAICFATCLYPGTMILLSVIGSTVSSLFLALLSKHPISLWIGTAVYGASMATVFPSGISWIEQYTTIQGKSASLFVVGAALGEMCIPAVVGFLEGKLHNVPVMMYAALGTSVMTAVLFPVMYKLATSPSESKLKDGGGSEDQKALLSNFELNEDEEDEEDAGEWNEADFEVIEMNDTVRNSVVETCQKIPGESPAKASTQPPSNIFSSSPGTLLLVTLHIPKLNFKGNFTVLVG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMV9</accession>
    <name>A0A452IMV9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Kinesin-like protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
            <location evidence="2">Microtubule organizing center</location>
            <location evidence="2">Centrosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Spindle</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="13 14">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Kinesin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMV9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033184.1">
        <property type="protein sequence ID" value="ENSGAGP00000029220.1"/>
        <property type="gene ID" value="ENSGAGG00000021132.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000033186.1">
        <property type="protein sequence ID" value="ENSGAGP00000029222.1"/>
        <property type="gene ID" value="ENSGAGG00000021132.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005874">
        <property type="term" value="C:microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005819">
        <property type="term" value="C:spindle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008017">
        <property type="term" value="F:microtubule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003777">
        <property type="term" value="F:microtubule motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007019">
        <property type="term" value="P:microtubule depolymerization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007018">
        <property type="term" value="P:microtubule-based movement"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd01367">
        <property type="entry name" value="KISc_KIF2_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000006">
        <property type="entry name" value="Kinesin-like protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR054473">
        <property type="entry name" value="KIF2A-like_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027640">
        <property type="entry name" value="Kinesin-like_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019821">
        <property type="entry name" value="Kinesin_motor_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001752">
        <property type="entry name" value="Kinesin_motor_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47971:SF24">
        <property type="entry name" value="KINESIN-LIKE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47971">
        <property type="entry name" value="KINESIN-RELATED PROTEIN 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22923">
        <property type="entry name" value="KIF2A-like_1st"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00225">
        <property type="entry name" value="Kinesin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00380">
        <property type="entry name" value="KINESINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00129">
        <property type="entry name" value="KISc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00411">
        <property type="entry name" value="KINESIN_MOTOR_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50067">
        <property type="entry name" value="KINESIN_MOTOR_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 13 14" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="12" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="4" id="KW-0132">Cell division</keyword>
    <keyword evidence="9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="5 14" id="KW-0493">Microtubule</keyword>
    <keyword evidence="7" id="KW-0498">Mitosis</keyword>
    <keyword evidence="10 13 14" id="KW-0505">Motor protein</keyword>
    <keyword evidence="6 13 14" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Kinesin motor" evidence="16">
        <location>
            <begin position="223"/>
            <end position="553"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="64"/>
            <end position="141"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="165"/>
            <end position="186"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="15">
        <location>
            <begin position="123"/>
            <end position="140"/>
        </location>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <begin position="313"/>
            <end position="320"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004186"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022776"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00283"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="RuleBase" id="RU000394"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50067"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029220.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="725" mass="81835" checksum="29AD1AC1DD8C644F" modified="2019-05-08" version="1">MAAANFGKIQIGIYVEIKRSDGRIHQAMVTSLNEDNESVTVEWIENGDTKGKEIDLESIFSLNPDLAPDEDIEPSPETPPPPTPSAKVNKIVKNRRTVAPIKNETPARDNRAVGSARARPTQLPDQSSSTQQNGSVSDISPVQAAKKEFGPPSRRKSNCVKEVEKLQEKREKRRLQQQELREKRAQDVDATNPNYEIMCMIRDFRGSLDYRPLTTTDPIDEHRICVCVRKRPLNKKETVMKDLDVITIPSKDVVMVHEPKQKVDLTRYLENQTFRFDYAFDDTAPNEMVYRFTARPLVETIFERGMATCFAYGQTGSGKTHTMGGDFSGKNQDCSKGIYALAARDVFLMLKKPNYKKLELQVYATFFEIYSGKVFDLLNRKTKLRVLEDGKQQVQVVGLQEREVKCVEDVLKLIEIGNSCRTSGQTSANAHSSRSHAVFQIILRRKGKLHGKFSLIDLAGNERGADTSSADRQTRLEGAEINKSLLALKECIRALGRNKPHTPFRASKLTQVLRDSFIGENSRTCMIATISPGMASCENTLNTLRYANRVKEFGISPSDIPFSQSSGSRSDLSPSYEYDDFSPSITRVKELTVDPSAAGDIRPIIHHAPNQIDDLETQWGVGSSPQRDDLKLLCEQNEEEVSPQLFTFHEAVSQMVEMEEQVVEDHRAVFQESIRWLEDEKALLEMTEEVDYDVDSYATQLEAILEQKIDILTELRGKAVFFVHL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMW0</accession>
    <name>A0A452IMW0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">SMB domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMW0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033408.1">
        <property type="protein sequence ID" value="ENSGAGP00000029419.1"/>
        <property type="gene ID" value="ENSGAGG00000020186.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009986">
        <property type="term" value="C:cell surface"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004551">
        <property type="term" value="F:dinucleotide phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003676">
        <property type="term" value="F:nucleic acid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004528">
        <property type="term" value="F:phosphodiesterase I activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046034">
        <property type="term" value="P:ATP metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030505">
        <property type="term" value="P:inorganic diphosphate transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045599">
        <property type="term" value="P:negative regulation of fat cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009143">
        <property type="term" value="P:nucleoside triphosphate catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030500">
        <property type="term" value="P:regulation of bone mineralization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd16018">
        <property type="entry name" value="Enpp"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00091">
        <property type="entry name" value="NUC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.410.20:FF:000003">
        <property type="entry name" value="Ectonucleotide pyrophosphatase/phosphodiesterase family member 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.410.20:FF:000001">
        <property type="entry name" value="Ectonucleotide pyrophosphatase/phosphodiesterase family member 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.720.10:FF:000145">
        <property type="entry name" value="Uncharacterized protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.410.20">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.720.10">
        <property type="entry name" value="Alkaline Phosphatase, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.570.10">
        <property type="entry name" value="Extracellular Endonuclease, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017850">
        <property type="entry name" value="Alkaline_phosphatase_core_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044929">
        <property type="entry name" value="DNA/RNA_non-sp_Endonuclease_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001604">
        <property type="entry name" value="Endo_G_ENPP1-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020821">
        <property type="entry name" value="ENPP1-3/EXOG-like_nuc-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044925">
        <property type="entry name" value="His-Me_finger_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002591">
        <property type="entry name" value="Phosphodiest/P_Trfase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036024">
        <property type="entry name" value="Somatomedin_B-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001212">
        <property type="entry name" value="Somatomedin_B_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10151">
        <property type="entry name" value="ECTONUCLEOTIDE PYROPHOSPHATASE/PHOSPHODIESTERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10151:SF77">
        <property type="entry name" value="ECTONUCLEOTIDE PYROPHOSPHATASE_PHOSPHODIESTERASE FAMILY MEMBER 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01223">
        <property type="entry name" value="Endonuclease_NS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01663">
        <property type="entry name" value="Phosphodiest"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01033">
        <property type="entry name" value="Somatomedin_B"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00892">
        <property type="entry name" value="Endonuclease_NS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00477">
        <property type="entry name" value="NUC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00201">
        <property type="entry name" value="SO"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53649">
        <property type="entry name" value="Alkaline phosphatase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54060">
        <property type="entry name" value="His-Me finger endonucleases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90188">
        <property type="entry name" value="Somatomedin B domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00524">
        <property type="entry name" value="SMB_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50958">
        <property type="entry name" value="SMB_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="4" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <feature type="domain" description="SMB" evidence="7">
        <location>
            <begin position="7"/>
            <end position="47"/>
        </location>
    </feature>
    <feature type="domain" description="SMB" evidence="7">
        <location>
            <begin position="50"/>
            <end position="94"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50958"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029419.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="818" mass="94578" checksum="379B361C7538511A" modified="2019-05-08" version="1">MTLSNISVKTCKGRCFERTFGSCRCDRDCVELGNCCLDFQETCIQPGKTHIWTCNKFRCGEKRWPENRCSCSDDCVENKDCCVNYYIVCQGAKSWVEEKCEDINEPQCPPGFAKPPVLLFSLDGFRAEYLHTWSGLLPVISKLQKCGTYTPNMRPVYPTKTFPNHYTIVTGLYPESHGIIDNKIYDPHRNVSFTLKNAEKFNPQWYQGQPIWLTAMHQGLKAGTFFWPGSDVRINGTFPDLYRKYNGSIPFEERVIAILHWLQLPKDERPHFYTLYLEEPDSSGHNFGPVSSGVILALQRVDNIVGMLMDGLKQMNLHKCLNIILLSDHGKLMCTHCSKTAYLSTYMENVQDITVIPGPAARLRPQNVPDEYFSYPDQHFKAFMKELLPKRFHYVNNDRIERVHFYLDPQWQLARKPSEIKYCNGGFHGSDNRFMSMQAIFIGFGPGFKFKTQVDPFENIEIYNLICDLLGITPAYNNGTHGSLNHLLKNPVYIPRHPEEESHPSVYPLMKTTSSLDIDIFCPFMNNWILLQICYIELVINSVSALVKKIEKYSLPYGRPRVLQKRSTYYLLYHHQYVSGYSQDLNMTLWSAYTVTKYDNRTASVENLSNCLHVDIRIPSSSSKTCSFYKNHHQLSYGFLFPPNTKKSSYEVLLNSNIVPMYHEFKVLWNYFYDVLLPKYTLARNGVNVVSGPVFDYDSNGLYDTPEKVKRLTHDPEVLVPTHYFIVLTSCKNISQTPLQCEGALDALSFIVPHRPDNSESCARHKRDSLWVEERMRFHTARVRDVELLTGLSFYQDRKQPVSDILQLKTYLPTFPME</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMW6</accession>
    <name>A0A452IMW6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">DNA polymerase epsilon catalytic subunit</fullName>
            <ecNumber evidence="16">2.7.7.7</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="16">DNA polymerase II participates in chromosomal DNA replication.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="16">
            <text>DNA(n) + a 2'-deoxyribonucleoside 5'-triphosphate = DNA(n+1) + diphosphate</text>
            <dbReference type="Rhea" id="RHEA:22508"/>
            <dbReference type="Rhea" id="RHEA-COMP:17339"/>
            <dbReference type="Rhea" id="RHEA-COMP:17340"/>
            <dbReference type="ChEBI" id="CHEBI:33019"/>
            <dbReference type="ChEBI" id="CHEBI:61560"/>
            <dbReference type="ChEBI" id="CHEBI:173112"/>
            <dbReference type="EC" id="2.7.7.7"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="16">
            <name>[4Fe-4S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:49883"/>
        </cofactor>
    </comment>
    <comment type="subunit">
        <text evidence="15">Component of the DNA polymerase epsilon complex consisting of four subunits: the catalytic subunit POLE and the accessory subunits POLE2, POLE3 and POLE4. Interacts with RAD17 and TOPBP1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 16">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 16">Belongs to the DNA polymerase type-B family.</text>
    </comment>
    <dbReference type="EC" id="2.7.7.7" evidence="16"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033156.1">
        <property type="protein sequence ID" value="ENSGAGP00000029193.1"/>
        <property type="gene ID" value="ENSGAGG00000020939.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008622">
        <property type="term" value="C:epsilon DNA polymerase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051539">
        <property type="term" value="F:4 iron, 4 sulfur cluster binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003887">
        <property type="term" value="F:DNA-directed DNA polymerase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000166">
        <property type="term" value="F:nucleotide binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008310">
        <property type="term" value="F:single-stranded DNA 3'-5' DNA exonuclease activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006287">
        <property type="term" value="P:base-excision repair, gap-filling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045004">
        <property type="term" value="P:DNA replication proofreading"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006272">
        <property type="term" value="P:leading strand elongation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000278">
        <property type="term" value="P:mitotic cell cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006297">
        <property type="term" value="P:nucleotide-excision repair, DNA gap filling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05779">
        <property type="entry name" value="DNA_polB_epsilon_exo"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05535">
        <property type="entry name" value="POLBc_epsilon"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.132.60:FF:000002">
        <property type="entry name" value="DNA polymerase epsilon catalytic subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.420.10:FF:000010">
        <property type="entry name" value="DNA polymerase epsilon catalytic subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1600.10:FF:000006">
        <property type="entry name" value="DNA polymerase epsilon catalytic subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.132.60">
        <property type="entry name" value="DNA polymerase family B, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.342.10">
        <property type="entry name" value="DNA Polymerase, chain B, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1600.10">
        <property type="entry name" value="Palm domain of DNA polymerase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.420.10">
        <property type="entry name" value="Ribonuclease H-like superfamily/Ribonuclease H"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006172">
        <property type="entry name" value="DNA-dir_DNA_pol_B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006133">
        <property type="entry name" value="DNA-dir_DNA_pol_B_exonuc"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043502">
        <property type="entry name" value="DNA/RNA_pol_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042087">
        <property type="entry name" value="DNA_pol_B_thumb"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013697">
        <property type="entry name" value="DNA_pol_e_suA_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023211">
        <property type="entry name" value="DNA_pol_palm_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029703">
        <property type="entry name" value="POL2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055191">
        <property type="entry name" value="POL2_thumb"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012337">
        <property type="entry name" value="RNaseH-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036397">
        <property type="entry name" value="RNaseH_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR054475">
        <property type="entry name" value="Znf-DPOE"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10670">
        <property type="entry name" value="DNA POLYMERASE EPSILON CATALYTIC SUBUNIT A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10670:SF0">
        <property type="entry name" value="DNA POLYMERASE EPSILON CATALYTIC SUBUNIT A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03104">
        <property type="entry name" value="DNA_pol_B_exo1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08490">
        <property type="entry name" value="DUF1744"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22634">
        <property type="entry name" value="POL2_thumb"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22912">
        <property type="entry name" value="zf-DPOE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23250">
        <property type="entry name" value="zf_DPOE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01159">
        <property type="entry name" value="DUF1744"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00486">
        <property type="entry name" value="POLBc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56672">
        <property type="entry name" value="DNA/RNA polymerases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53098">
        <property type="entry name" value="Ribonuclease H-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 16" id="KW-0004">4Fe-4S</keyword>
    <keyword evidence="6 16" id="KW-0235">DNA replication</keyword>
    <keyword evidence="13 16" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="10 16" id="KW-0239">DNA-directed DNA polymerase</keyword>
    <keyword evidence="11 16" id="KW-0408">Iron</keyword>
    <keyword evidence="12 16" id="KW-0411">Iron-sulfur</keyword>
    <keyword evidence="7 16" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="5 16" id="KW-0548">Nucleotidyltransferase</keyword>
    <keyword evidence="14 16" id="KW-0539">Nucleus</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 16" id="KW-0808">Transferase</keyword>
    <keyword evidence="9 16" id="KW-0862">Zinc</keyword>
    <keyword evidence="8 16" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="DNA polymerase epsilon catalytic subunit A C-terminal" evidence="18">
        <location>
            <begin position="1531"/>
            <end position="1920"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1"/>
            <end position="42"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1922"/>
            <end position="1957"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="2005"/>
            <end position="2026"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="17">
        <location>
            <begin position="32"/>
            <end position="42"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="17">
        <location>
            <begin position="1940"/>
            <end position="1957"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005755"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022485"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022695"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022705"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022932"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023014"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00062298"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="RuleBase" id="RU365029"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="SMART" id="SM01159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029193.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2273" mass="260262" checksum="55CFD96E6171CD9D" modified="2019-05-08" version="1">MVLRNSGRRRAEPAGGDPDTPRDDGSGLSALKRLERSQRTDRLDAQFGFERPKEPGEKTGWLINMHPTEILDDDKRLVSSVDYYFIQEDGSRFKVALPYKPYFYIATRKGCEREVSSFLSKRYQGKIAKLETVPKEDLDLPNHLVGLKRNYVKLSFNTVDDLVKVRREISPAVKKNRERDQASDTYTSMLSSALIGGSLTMDDEGPSKKTVNQMDNIVDMREYDVPYHIRLSIDLKIHVAHWYNVRYRGSIYPPEITRRDDLVERPDPVVLAFDIETTKLPLKFPDAETDQIMMISYMIDGQGYLITNREIVSEDIEDFEFTPKPEYEGPFCIFNEPDEAHLIQRWFEHVQETKPTIMVTYNGDFFDWPFVEARAAVHGMNMYQEIGFQKDSQGEYKASQCIHMDCLRWVKRDSYLPVGSHNLKAAAKAKLGYDPVELDPEEMCRMATEEPQTLATYSVSDAVATYYMYMKYVHPFIFALCTIIPMEPDEVLRKGSGTLCEALLMVQAYHANIIFPNKQEQEFNKLTEDGHVLDSETYVGGHVEALESGVFRSDIPCRFKMNPVAFDFLLQRVEKTLRHAIEEEEGLPLDQVTNFQEVCDEIKVKLNSLKDIPNRIECPLIYHLDVGAMYPNIILTNRLQPSAMVDEATCAACDFNKPGANCQRRMTWQWRGEFMPASRSEYHRIQQQLESEKLPPLYPDGPPRAFHELSREEQAKFEKKRLADYCRKAYKKLHVTKVEERITTICQRENSFYVDTVRAFRDRRYEFKGLHKVWKKKLSAATEMGDATEVKRCKNMEILYDSLQLAHKCILNSFYGYVMRKGARWYSMEMAGIVCFTGANIITQARELIEQIGRPLELDTDGIWCVLPNSFPENFVIKSTNAKKPKVTISYPGAMLNILVKEGFTNDQYQELVDPASLTYVTRSENSIFFEVDGPYLAMILPASKEEGKKLKKRYAVFNEDGSLAELKGFEVKRRGELQLVKIFQSSVFESFLKGTTLEEVYASVAKVADYWLDVLYSKAANMPDSELFELISENRSMSRKLEDYGEQKSTSISTAKRLAEFLGDQMVKDAGLSCRFIISKKPEGSPVTERAIPLAIFQAELTVRKHYLRKWLKSPSLQDFNIRTILDWDYYIERLGSTIQKIITIPAALQQVKNPVPRVRHPDWLHKKLLEKNDVYKQKKINELFTSEGKRQVTANQLQEGTPCTQVADMEDFGASKPMQPSVPIASKRKRVLVTGESQEMSQNLELTQSWREILGPPPPTGTTKEERLAWLRYHKKKWELQARQRQARRKKRRLEDGEVVLGGGVIRDGSSKGLSSYLRRTARSILDLPWQIVQVAETSQPGLFRLWAVIGSDLHCMKLSVPRVFYVNQRVAKPEEGATYRKVNRVLPRSNLLYNLYEYSVPEDMYQEHINEINAGLSAPDIEGVYETQVPLLLRALIHLGCVCMVSRQLVRHLTGREAETFALEHLEMRSLAQFSYLEPGSIRHIYLYHNSQGSKALFGLFVPSQRKASVFVLDTVRGGLQAAGPWIAMLEKVGEELLPPDKHTFEVRAETDLKTVSRAIQRLLLGYKDERRGPTLIAVQSNWDVKRLVSGIPVFEEFPLVPMRVTDDISYAVLDWQRHAARRMIRHYLNLDTCLSQAFEMSRYYHIPIGNLPDDISTFGSDLFFSRHLRRHNHLLWLSPTARPDLGGKEADDNRLVMEFDDRASVEINNPGCYSTVCLELDIQSLAVNTILQSHHINDMEGASSMSISFDVIQQASLEDMVTGNQAASIPASYDETALCSNTFRILKSMVVGWVKEITQYHNVYADNQVIHFYRWLRSPASLLHDPALHRTLHNMMKKLFLQLVAEFKRLGSSVVYANFNRIILCTKKRRIEDAIAYVEYITNSIHSKEIFHSLTISFSRCWEFLLWMDPANYGGIKGRVPSGEVSTQGRQAGAEEGSEDEEEEAEEEEEPGEASVEDLLENNWNIVQYLPQAASCQNYFLMIVSAYIVAVYHSMKEEMRRSAPGSTPVRRRPSSQASQEAVGEVGTMPGVITFSQDYVSNELTQSFFTITQKIQKKVTGSRHATLPSDMFPALPGSYLPLNNPALEFIKYVCKVLSLDANITNQVNKLKRDLLRLVDVGEFSEEAQFRDPCRSYVLSEVICRNCNFCRDLDLCKDRSPCQDAAVPPSWVCSNCQAEYERGSIEAALVEALQKKLMAFMLQDLVCVKCKGVKETHMPIYCTCAGDFTLLIQTKAFVEQINVFRSIARHYSMAHLLETIEWLLQLNPQLR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMW8</accession>
    <name>A0A452IMW8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Unc-13 homolog A</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IMW8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033172.1">
        <property type="protein sequence ID" value="ENSGAGP00000029208.1"/>
        <property type="gene ID" value="ENSGAGG00000020811.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031594">
        <property type="term" value="C:neuromuscular junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098831">
        <property type="term" value="C:presynaptic active zone cytoplasmic component"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042734">
        <property type="term" value="C:presynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030672">
        <property type="term" value="C:synaptic vesicle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043195">
        <property type="term" value="C:terminal bouton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019992">
        <property type="term" value="F:diacylglycerol binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005543">
        <property type="term" value="F:phospholipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017075">
        <property type="term" value="F:syntaxin-1 binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061789">
        <property type="term" value="P:dense core granule priming"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0099525">
        <property type="term" value="P:presynaptic dense core vesicle exocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035249">
        <property type="term" value="P:synaptic transmission, glutamatergic"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016081">
        <property type="term" value="P:synaptic vesicle docking"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016082">
        <property type="term" value="P:synaptic vesicle priming"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd20859">
        <property type="entry name" value="C1_Munc13-2-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd08394">
        <property type="entry name" value="C2A_Munc13"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04027">
        <property type="entry name" value="C2B_Munc13"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.357.50:FF:000001">
        <property type="entry name" value="Protein unc-13 homolog B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000002">
        <property type="entry name" value="Protein unc-13 homolog B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000031">
        <property type="entry name" value="Protein unc-13 homolog B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.60.20:FF:000001">
        <property type="entry name" value="Protein unc-13 homolog B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.357.50">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.1100">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.60.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046349">
        <property type="entry name" value="C1-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000008">
        <property type="entry name" value="C2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010439">
        <property type="entry name" value="MUN_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014770">
        <property type="entry name" value="Munc13_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002219">
        <property type="entry name" value="PKC_DAG/PE"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027080">
        <property type="entry name" value="Unc-13"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037302">
        <property type="entry name" value="Unc-13_C2B"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10480">
        <property type="entry name" value="PROTEIN UNC-13 HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10480:SF1">
        <property type="entry name" value="PROTEIN UNC-13 HOMOLOG A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00130">
        <property type="entry name" value="C1_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00168">
        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06292">
        <property type="entry name" value="MUN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00360">
        <property type="entry name" value="C2DOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00109">
        <property type="entry name" value="C1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00239">
        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM01145">
        <property type="entry name" value="DUF1041"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49562">
        <property type="entry name" value="C2 domain (Calcium/lipid-binding domain, CaLB)"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57889">
        <property type="entry name" value="Cysteine-rich domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50004">
        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51258">
        <property type="entry name" value="MHD1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00479">
        <property type="entry name" value="ZF_DAG_PE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50081">
        <property type="entry name" value="ZF_DAG_PE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0268">Exocytosis</keyword>
    <keyword evidence="2" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <keyword evidence="4" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2" evidence="7">
        <location>
            <begin position="1"/>
            <end position="97"/>
        </location>
    </feature>
    <feature type="domain" description="Phorbol-ester/DAG-type" evidence="8">
        <location>
            <begin position="465"/>
            <end position="515"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="7">
        <location>
            <begin position="571"/>
            <end position="695"/>
        </location>
    </feature>
    <feature type="domain" description="MHD1" evidence="9">
        <location>
            <begin position="1003"/>
            <end position="1146"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="176"/>
            <end position="318"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="195"/>
            <end position="204"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="205"/>
            <end position="224"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="231"/>
            <end position="246"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="255"/>
            <end position="266"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="267"/>
            <end position="279"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="287"/>
            <end position="303"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50081"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51258"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029208.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="12">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1275" mass="144865" checksum="18BB7DA868BFCBDF" modified="2019-05-08" version="1">TFYSLFAVKKAKFDGAQEKFNTYVTLKVQNVKSTTIAVRGNLPCWEQDFMFEINRLDLGLTVEVWNKGLIWDTMVGTVWIPLQTIRQSNEEGPGEWLTLDSQVIMTDNEISGTKDPTFHRILLDTHFELPLDIPEEEARYWAKKLEQLNAMKDQDDVNTGYLIPLRHVVASGPLPCETLTGDNRPFADDDPDSTMDDRDSDYRSETSNSIPPPYYTTSQPNASVHQYPMRHQQQSSRDSYMDSVQNYKMDYSDQRPISPTGSSRYASSGELSQGSSQLSDDFDPDNESLRGSELDERDGDSYHSCHSSVSYHDDSPRWEEDFYPEEEDEDYSENEQDFPKVEAEVLRDREQEDVEVQDAKSRAKANWLKAFNKVCMQLQEVSAPSLPSLPAGRGPGGGLIIIDSMPDIRKRKPIPLVSDLSLVQSRKAGITSALASSTLNNEELKNHVYKKTLQALIYPISSTTPHNFEVWTATTPTYCYECEGLLWGIARQGMRCTECGVKCHEKCQDLLNADCLQRAAEKSSKHGAEDRTQNIIMVLKDRMKIRERNKPEIFELIQEIFAVSKATHTQQMKAVKQSVLDGTSKWSAKISITVVCAQGLQAKDKTGSSDPYVTVQVGKTKKRTKTIYGNLNPVWEENFHFECHNSSDRIKVRVWDEDDDIKSRVKQRFKRESDDFLGQTIIEVRTLSGEMDVWYNLDKRTDKSAVSGAIRLHISVEIKGEEKVAPYHVQYTCLHENLFHFVTDIQNNGVVKIPEAKGDDAWKVYFDETAQEIVDEFAMRYGVESIYQAMTHFACLSSKYMCPGVPAVMSTLLANINAYYAHTTASTNVSASDRFAASNFGKERFVKLLDQLHNSLRIDLSMYRNNFPASSPERLQDLKSTVDLLTSITFFRMKVQELQSPPRASQVVKDCVKACLNSTYEYIFNNCHELYSREYQTDPKGEVPPEEQGPSIKNLDFWSKLITLIVSIIEEDKNSYTPCLNQFPQELNVGKISAEVMWNLFAQDMKYAMEEHDKHRLCKSADYMNLHFKVKWLYNEYVTELPAFKSRVPEYPAWFEPFVIQWLDENEEVSRDFLHGALERDKKDGFQQTSEHALFSCSVVDVFSQLNQSFEIIKKLECPDPQIVGHYMRRFAKTISNVLLQYAEIISKDFALYCSKEKEKPCILMNNIQQLRVQLEKMFEAMGGKELDAEASDILKELQVKLNNVLDELSRVFATSFQPHIEECVKQMGDILSQVKGTGNVPASACSSVAQDADNQVSAGGLGGPVGRGTGGTAG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMW9</accession>
    <name>A0A452IMW9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="5">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the STRIP family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMW9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033177.1">
        <property type="protein sequence ID" value="ENSGAGP00000029213.1"/>
        <property type="gene ID" value="ENSGAGG00000020767.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007010">
        <property type="term" value="P:cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040185">
        <property type="entry name" value="Far11/STRP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021819">
        <property type="entry name" value="Far11/STRP_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012486">
        <property type="entry name" value="Far11/STRP_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13239">
        <property type="entry name" value="PROTEIN REQUIRED FOR HYPHAL ANASTOMOSIS HAM-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13239:SF6">
        <property type="entry name" value="STRIATIN-INTERACTING PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11882">
        <property type="entry name" value="DUF3402"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07923">
        <property type="entry name" value="N1221"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01293">
        <property type="entry name" value="DUF3402"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01292">
        <property type="entry name" value="N1221"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Far11/STRP N-terminal" evidence="3">
        <location>
            <begin position="16"/>
            <end position="312"/>
        </location>
    </feature>
    <feature type="domain" description="Far11/STRP C-terminal" evidence="4">
        <location>
            <begin position="409"/>
            <end position="749"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="317"/>
            <end position="336"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="2">
        <location>
            <begin position="325"/>
            <end position="336"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007062"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="SMART" id="SM01292"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="SMART" id="SM01293"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029213.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="771" mass="88872" checksum="6318B9E7BF8D1D77" modified="2019-05-08" version="1">MSHPHPGCIFFLHCDPAFKPFFLCVSSPIELYSYTEEPEFTINRKCFEEDFQTQGIPVRTIRWPELDLAKQKAYVMRLLDGLEVVNRDKRLKAARAVLYLAQGVFGECDVEAEVLHWSRHNNFLLFQMGTFSAFLELLNMEIDNSQACSSALRKPAISLADSTELRVLLSVMYLMVENIRVEAEADPPEWKSSRETFRTELSEYEEPFALLLFTMVAKFCSGHAPHFPMKKVLLLLWKVLLFTLGGFEALQAMKTKKREELGLPPLPEDSIQVVRRMRAASPPTYTIELLDQQSSKRGHRSRRPLVKQDSLDIYNERDPFKNDEAGVDEEENDDVDSGIEGELDLMERDAIIQAGPAQRPPIERVSFPKGLPWAPKVRQKDIEHFLEASRNKFIGFTLGQDTDTLVGLPRPIHESVKTLKQVMQWLCALNICSCSGLWDLDIGDGLSDILNCENIALLKILLAAAPTSKAKTDSINILADVLPEEMPITVLQSMKLGVDVNRHKEIIVKSISASLLLLLKHFKLNHIYQFEYVSQHLVFANCIPLILKFFNQNIMSYITAKNSISVLDYPHCTVHELPELTAESLEAGDNNQFCWRNLFSCINLLRLLNKLTKWKHSRTMMLVVFKSAPILKRALKVKQAMMQLYVLKLLKIQTKYLGRQWRKSNMKTMSAIYQKVRHRMNDDWAYGNDIDARPWDFQAEECTLRANIEAFNSRRYDKSQDSEFAPVDNCLQSVLGQPLELPEDFHYSYELWLEREVFSQPIRWEELLHYQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMX2</accession>
    <name>A0A452IMX2_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="15">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">May be involved in transcriptional regulation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMX2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033280.1">
        <property type="protein sequence ID" value="ENSGAGP00000029301.1"/>
        <property type="gene ID" value="ENSGAGG00000021197.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003700">
        <property type="term" value="F:DNA-binding transcription factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07765">
        <property type="entry name" value="KRAB_A-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002063">
        <property type="entry name" value="RB associated KRAB zinc finger"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001430">
        <property type="entry name" value="Uncharacterized protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001772">
        <property type="entry name" value="Uncharacterized protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000478">
        <property type="entry name" value="Zinc finger protein 133"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000688">
        <property type="entry name" value="zinc finger protein 197 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002343">
        <property type="entry name" value="Zinc finger protein 33A"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001119">
        <property type="entry name" value="zinc finger protein 408"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001343">
        <property type="entry name" value="Zinc finger protein 568"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001270">
        <property type="entry name" value="zinc finger protein 583 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000624">
        <property type="entry name" value="zinc finger protein 697"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="22"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050589">
        <property type="entry name" value="Ikaros_C2H2-ZF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001909">
        <property type="entry name" value="KRAB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036051">
        <property type="entry name" value="KRAB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24404">
        <property type="entry name" value="ZINC FINGER PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24404:SF100">
        <property type="entry name" value="ZINC FINGER PROTEIN 501"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01352">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="SMART" id="SM00349">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="22"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="12"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109640">
        <property type="entry name" value="KRAB domain (Kruppel-associated box)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50805">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="18"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="22"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="10" id="KW-0804">Transcription</keyword>
    <keyword evidence="8" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="7" id="KW-0862">Zinc</keyword>
    <keyword evidence="6 12" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="KRAB" evidence="14">
        <location>
            <begin position="7"/>
            <end position="76"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="82"/>
            <end position="109"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="110"/>
            <end position="137"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="138"/>
            <end position="165"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="166"/>
            <end position="193"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="194"/>
            <end position="221"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="222"/>
            <end position="249"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="250"/>
            <end position="277"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="278"/>
            <end position="305"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="306"/>
            <end position="333"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="334"/>
            <end position="361"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="362"/>
            <end position="389"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="390"/>
            <end position="417"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="418"/>
            <end position="445"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="446"/>
            <end position="473"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="474"/>
            <end position="501"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="502"/>
            <end position="529"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="530"/>
            <end position="559"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="560"/>
            <end position="587"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="588"/>
            <end position="615"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="616"/>
            <end position="643"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="644"/>
            <end position="671"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="672"/>
            <end position="699"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00003767"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029301.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="700" mass="81267" checksum="C96CE06193A248BE" modified="2019-05-08" version="1">QPHLVPVTFEEVAVYFTQRQGALLDPAQTALYRDVMLENYETVTSLGKGFPSPRLLESVGPAMGGPNDSTVQQTNPNEENPYQCLKCGKGLILRSQLVTHQTIHTREKPLQCLDHGKTFNNRSDLKIHGRSHTREKPSRCLEHKKSFYCKSALITHQRIHTQERPHKCLHCGKSFTRRSHLIQHGRIHTGERPHKCLDCGKSFRYRSVFLSHLRTHTGEKPHKCLDCGKSFRYRSVLLSHLRTHTGEKPHKCLDCGKSFRYRSVLLSHLRTHTGEKPHKCLDCGKSFRYRSVLLSHLRTHTGEKPHKCLDCGKSFRYRSVLLSHLRTHTGEKPHKCLECGKSFMWRSTLVLHQRIHTGERPHKCLDCGKSFTRRSHLIQHGRIHTGERPHKCLDCGKSFRYRSVFLSHLRTHTGEKPHKCLDCGKSFRYRSVLLSHLRTHTGERPHKCLECGKSFMWRSTLVLHQRIHTGERPHKCLECGKSFMWRSALVLHQRIHTGERPHKCLECGKSFMWRSTLVLHQRIHTGERPHKCLECGKSFIFMRRSDLDLHQRIHTGERPHRCLDCGKSFTWRSALVKHGRTHTGERPHKCLDCGKSFTWRSVLVKHGRIHTGERPHKCLDCGKSFTWRSALVSHQRIHTGERPHKCLDCGKSFIWRSALVSHKRIHTEERPHKGLDCGKSFTQRSKLIKHGRIHTASKIL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN54</accession>
    <name>A0A452IN54_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Protein kinase domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the protein kinase superfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IN54"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033277.1">
        <property type="protein sequence ID" value="ENSGAGP00000029298.1"/>
        <property type="gene ID" value="ENSGAGG00000021195.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004672">
        <property type="term" value="F:protein kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd14011">
        <property type="entry name" value="PK_SCY1_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000179">
        <property type="entry name" value="SCY1 like pseudokinase 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000189">
        <property type="entry name" value="SCY1-like pseudokinase 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051177">
        <property type="entry name" value="CIK-Related_Protein"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12984:SF6">
        <property type="entry name" value="SCY1-LIKE PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12984">
        <property type="entry name" value="SCY1-RELATED S/T PROTEIN KINASE-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Protein kinase" evidence="3">
        <location>
            <begin position="32"/>
            <end position="327"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="631"/>
            <end position="710"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="805"/>
            <end position="848"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="879"/>
            <end position="921"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="637"/>
            <end position="669"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="678"/>
            <end position="692"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="693"/>
            <end position="703"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="805"/>
            <end position="824"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="892"/>
            <end position="902"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="903"/>
            <end position="914"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00038349"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029298.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="921" mass="102363" checksum="A3BAB26F6B83C22F" modified="2019-05-08" version="1">MESMLNKLKSTVTKVTADVTSAVMGNPVTREFDVGRHIASGGNGLAWKIFNGTKKSTKQEVAVFVFDKKLIDKYQKFEKDQIIDSLKRGVQQLTRLRHPRLVTVQHPLEESRDCLAFCTEPVFASLANVLGNWDNLPSPLPSDIKEYKLYDVETKYGLLQVSEGLSFLHSSVKMVHGNITPENIILNKSGAWKIMGFDFCIQSTNPSEQELKFLCKEWDPNLPSLCLPNPEYLAPEYILSVSCETASDMYSLGAVTYAVFNKGKPIFEVNKKDIYKSFSRQLDQLSRLGSTTLQNIPEEVREHVKLLLNVAPAVRPDADQMTKIPFFDDVGAMTLQYFDSLFQRDNLQKSQFFKGLPKVLPKLPKRVIVQRILPYLTSEFVNPDMVPFVLPNVLLIAEECTKEEYVKLILPDLGPVFKQQEPVQILLIFLQKMDLLLTKTPPDEIKNSVLPMVYRALEAPSIQIQELCLNIIPTFANLIDYPSMKNALIPRIKNACLQTSSLAVRVNSLVCLGKILEYLDKWFVLDDILPFLQQIPSKEPAVLMGILGIYKCTFTHKKLGITKEQLAGKVLPHLIPLSIENNLNLNQFNSFISVIKEMLSRLELEHKTKLEQLHIMQEQQKSLDITNQMNVPEESKSNSTGNQIDKVFNNTGNDVPTSGSDNKENGLSSKQKKVSLTLEEKQKLAKEQEQAQKLKNQQPLKPQSHTITAPVKQTKDLTDTLIDNMSSLTSHSISKPKVASSSSFSSVSSMGVGMGFLSPVDSTKRNIANGLNTNMGFQTSGFSLGAGTTNQNFFSSPTATGMANVPNYSPVSATSGISQPTQHNRSPDMSALDSLFGPQKPKVSMNQLSQQKPNQWLTQFVPPQGSPGASSSVLGPQMNALGQPGFGMQGNPFFNPQNFAQPTNTMTNSSSASNDLKDLFG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN57</accession>
    <name>A0A452IN57_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="11">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the RIN (Ras interaction/interference) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IN57"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033374.1">
        <property type="protein sequence ID" value="ENSGAGP00000029390.1"/>
        <property type="gene ID" value="ENSGAGG00000021249.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000033377.1">
        <property type="protein sequence ID" value="ENSGAGP00000029392.1"/>
        <property type="gene ID" value="ENSGAGG00000021249.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030139">
        <property type="term" value="C:endocytic vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031267">
        <property type="term" value="F:small GTPase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016192">
        <property type="term" value="P:vesicle-mediated transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd16131">
        <property type="entry name" value="RA_Rin2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd10394">
        <property type="entry name" value="SH2_RIN2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1050.80:FF:000002">
        <property type="entry name" value="Ras and Rab interactor 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.505.10:FF:000052">
        <property type="entry name" value="Ras and Rab interactor 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.505.10">
        <property type="entry name" value="SH2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1050.80">
        <property type="entry name" value="VPS9 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000159">
        <property type="entry name" value="RA_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035868">
        <property type="entry name" value="RIN2_SH2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000980">
        <property type="entry name" value="SH2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036860">
        <property type="entry name" value="SH2_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029071">
        <property type="entry name" value="Ubiquitin-like_domsf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003123">
        <property type="entry name" value="VPS9"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045046">
        <property type="entry name" value="Vps9-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037191">
        <property type="entry name" value="VPS9_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23101">
        <property type="entry name" value="RAB GDP/GTP EXCHANGE FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23101:SF51">
        <property type="entry name" value="RAS AND RAB INTERACTOR 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00788">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23268">
        <property type="entry name" value="RIN1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02204">
        <property type="entry name" value="VPS9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00314">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00252">
        <property type="entry name" value="SH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00167">
        <property type="entry name" value="VPS9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55550">
        <property type="entry name" value="SH2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54236">
        <property type="entry name" value="Ubiquitin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109993">
        <property type="entry name" value="VPS9 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50200">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50001">
        <property type="entry name" value="SH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51205">
        <property type="entry name" value="VPS9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="3" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 6" id="KW-0727">SH2 domain</keyword>
    <feature type="domain" description="SH2" evidence="8">
        <location>
            <begin position="95"/>
            <end position="188"/>
        </location>
    </feature>
    <feature type="domain" description="VPS9" evidence="10">
        <location>
            <begin position="615"/>
            <end position="754"/>
        </location>
    </feature>
    <feature type="domain" description="Ras-associating" evidence="9">
        <location>
            <begin position="784"/>
            <end position="875"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="210"/>
            <end position="231"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="281"/>
            <end position="359"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="455"/>
            <end position="479"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="7">
        <location>
            <begin position="216"/>
            <end position="227"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="281"/>
            <end position="296"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="7">
        <location>
            <begin position="303"/>
            <end position="312"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="313"/>
            <end position="336"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006919"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022999"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00191"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50001"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50200"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS51205"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029392.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="891" mass="100684" checksum="E4186A78C3DE0677" modified="2019-05-08" version="1">MSSVTMEVRCLEKRGSFFKLIDTIASEIGELKQEMVQTDLMLEDGSTDLQSLVKDMDNVCPEKKDLKVCPRDSGYDSLSNKLSILDKLLHTHPVWLQLGLNDAEAVEILQSQPPGIFLVRKSTRMQKKVLSLRLPNEFGSRLKEFAIKESTYTFSLEGSGISFADLFRLIAFYCISRDVLPFTLKLPHAIAAAKTEAELEDIAQLGLNFWNSPANNNPPNPSPPRRPLPSDNACKDSRQLCLINGVHSIRTRTPSELECSQTNGALCFINPLFLKVHSQDVSGSLKRQSPRTQDVNGTEKPRSPPPRPPPPSINSLLTSPQLSRTIKQGSMPGTVNRNKHRNLDLLQKKPTPVPPPRLKKQAVCSEGEDNAKTLAVIRSSHNSVHGSEAASVPGEALCEQNLAACRKTLATNSESQVQWNGGRQRLSDMSISTSSSDSLDFDRSMPLFGYEGDTNSSLEDFEGESDQESMAPPLKPKKKRNSSFVLPKIVKSQLRKVSGVFSSFMTPEKRMIKKIAELSQDKRTYFGCLVQDYVSFLQENKECHVSSTDMLQTIRQFMTQVKNYLSQSSELDPPIESLIPEDQIDVVLEKAMHKCILKPLKGHIEAMLKEFHTVDGSWKQLKENLQLVRQRNPQELGVYVPTPDFVDVEKIKVKFMTMQKMYSPEKKVMLLLRVCKLIYTVMENNSGRMYGADDFLPVLTYVVAQCDMLELDTEIEYMMELLDPSLLHGEGGYYLTSAYGALSLIKNFQEEQAARLLSSEARDTLRQWHKRRTANRTIPSVDDFQNYLRVAFQEVNSGCTGKTLLVRPYITTEDVCQLCAEKFKVDSPEDYSLFLFIDDTWQQLTEDTYPQKIKAELHSRPQPQVFHFVYKRINSDPYGAIFQNNDDNSTL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN61</accession>
    <name>A0A452IN61_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Carbamoyl-phosphate synthase [ammonia], mitochondrial</fullName>
            <ecNumber evidence="10">6.3.4.16</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="14">Carbamoyl-phosphate synthetase I</fullName>
        </alternativeName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="12">Involved in the urea cycle of ureotelic animals where the enzyme plays an important role in removing excess ammonia from the cell.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>hydrogencarbonate + NH4(+) + 2 ATP = carbamoyl phosphate + 2 ADP + phosphate + 2 H(+)</text>
            <dbReference type="Rhea" id="RHEA:18029"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:17544"/>
            <dbReference type="ChEBI" id="CHEBI:28938"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:58228"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="6.3.4.16"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Mitochondrion</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="6.3.4.16" evidence="10"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033309.1">
        <property type="protein sequence ID" value="ENSGAGP00000029330.1"/>
        <property type="gene ID" value="ENSGAGG00000021129.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004087">
        <property type="term" value="F:carbamoyl-phosphate synthase (ammonia) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004088">
        <property type="term" value="F:carbamoyl-phosphate synthase (glutamine-hydrolyzing) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006541">
        <property type="term" value="P:glutamine metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000050">
        <property type="term" value="P:urea cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd01744">
        <property type="entry name" value="GATase1_CPSase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd01423">
        <property type="entry name" value="MGS_CPS_I_III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.470.20:FF:000030">
        <property type="entry name" value="Carbamoyl-phosphate synthase 1, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.20:FF:000012">
        <property type="entry name" value="Carbamoyl-phosphate synthase 1, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.880:FF:000006">
        <property type="entry name" value="Carbamoyl-phosphate synthase 1, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1380:FF:000010">
        <property type="entry name" value="carbamoyl-phosphate synthase [ammonia], mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.1030.10:FF:000001">
        <property type="entry name" value="Carbamoyl-phosphate synthase large chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1490.20:FF:000001">
        <property type="entry name" value="Carbamoyl-phosphate synthase large chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.470.20:FF:000001">
        <property type="entry name" value="Carbamoyl-phosphate synthase large chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.20:FF:000002">
        <property type="entry name" value="Carbamoyl-phosphate synthase large chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.20">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.880">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1490.20">
        <property type="entry name" value="ATP-grasp fold, A domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.470.20">
        <property type="entry name" value="ATP-grasp fold, B domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.50.30.20">
        <property type="entry name" value="Carbamoyl-phosphate synthase small subunit, N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1030.10">
        <property type="entry name" value="Carbamoyl-phosphate synthetase, large subunit oligomerisation domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1380">
        <property type="entry name" value="Methylglyoxal synthase-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011761">
        <property type="entry name" value="ATP-grasp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013815">
        <property type="entry name" value="ATP_grasp_subdomain_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036897">
        <property type="entry name" value="CarbamoylP_synth_lsu_oligo_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002474">
        <property type="entry name" value="CarbamoylP_synth_ssu_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036480">
        <property type="entry name" value="CarbP_synth_ssu_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029062">
        <property type="entry name" value="Class_I_gatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005479">
        <property type="entry name" value="CPAse_ATP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005483">
        <property type="entry name" value="CPSase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035686">
        <property type="entry name" value="CPSase_GATase1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006275">
        <property type="entry name" value="CPSase_lsu"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005480">
        <property type="entry name" value="CPSase_lsu_oligo"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058047">
        <property type="entry name" value="CPSase_preATP-grasp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017926">
        <property type="entry name" value="GATASE"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011607">
        <property type="entry name" value="MGS-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036914">
        <property type="entry name" value="MGS-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016185">
        <property type="entry name" value="PreATP-grasp_dom_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01369">
        <property type="entry name" value="CPSaseII_lrg"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF003671">
        <property type="entry name" value="PRK05294.1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF009455">
        <property type="entry name" value="PRK12815.1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF009475">
        <property type="entry name" value="PRK12838.1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11405:SF53">
        <property type="entry name" value="CARBAMOYL-PHOSPHATE SYNTHASE [AMMONIA], MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11405">
        <property type="entry name" value="CARBAMOYLTRANSFERASE FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25596">
        <property type="entry name" value="CPSase_L_D1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02786">
        <property type="entry name" value="CPSase_L_D2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02787">
        <property type="entry name" value="CPSase_L_D3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00988">
        <property type="entry name" value="CPSase_sm_chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00117">
        <property type="entry name" value="GATase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02142">
        <property type="entry name" value="MGS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00098">
        <property type="entry name" value="CPSASE"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00099">
        <property type="entry name" value="CPSGATASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM01096">
        <property type="entry name" value="CPSase_L_D3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01097">
        <property type="entry name" value="CPSase_sm_chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00851">
        <property type="entry name" value="MGS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48108">
        <property type="entry name" value="Carbamoyl phosphate synthetase, large subunit connection domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52021">
        <property type="entry name" value="Carbamoyl phosphate synthetase, small subunit N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52317">
        <property type="entry name" value="Class I glutamine amidotransferase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56059">
        <property type="entry name" value="Glutathione synthetase ATP-binding domain-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52335">
        <property type="entry name" value="Methylglyoxal synthase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52440">
        <property type="entry name" value="PreATP-grasp domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50975">
        <property type="entry name" value="ATP_GRASP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00866">
        <property type="entry name" value="CPSASE_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00867">
        <property type="entry name" value="CPSASE_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51273">
        <property type="entry name" value="GATASE_TYPE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51855">
        <property type="entry name" value="MGS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0021">Allosteric enzyme</keyword>
    <keyword evidence="7 15" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="4" id="KW-0436">Ligase</keyword>
    <keyword evidence="9" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="6 15" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="8" id="KW-0809">Transit peptide</keyword>
    <keyword evidence="2" id="KW-0835">Urea cycle</keyword>
    <feature type="domain" description="ATP-grasp" evidence="16">
        <location>
            <begin position="498"/>
            <end position="690"/>
        </location>
    </feature>
    <feature type="domain" description="ATP-grasp" evidence="16">
        <location>
            <begin position="1040"/>
            <end position="1231"/>
        </location>
    </feature>
    <feature type="domain" description="MGS-like" evidence="17">
        <location>
            <begin position="1302"/>
            <end position="1447"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004173"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022436"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022533"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022598"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00044063"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00047359"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00059545"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00074238"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00082675"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00409"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50975"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS51855"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029330.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1447" mass="158594" checksum="F37133EBA4E74337" modified="2019-05-08" version="1">AYAQTANLVLDDGTKMKGYSFGHPSSAAGEVVFNTGLAGYTETLTDPSYRGQILTMVNPIVGNGGVPDTAALDEMGLSKFLESDGIKVSGLLVLDYSNDYSHWRAVKTLGQWLQEEKVRDLWATRPDTRHGISILRKLLSDSLSNGNQEREILEVKVYGKGNPIKVVAVDCGLKNNAIRLLVKRGAEVHLVPWNHDFTGMEYDGLFLSGGPGDPMQAQELIQNLQKVLKSDRKEPLFGVSMGNLITGIAAGATSYKMPMANRGQNQPVMNLSNGQAFITAQNHGYALQSNTLPAGWKPLFVNVNDQTNEGIMHETKPIFTAQFNPEASAGPTDTEFLFDSFISLIKKGKGTTVASVLPKARTVASRLEVSKVLILGSGGLSIGQAGEFDYSGSQAVKAMKEENVKTVLMNPNIASVQTNEVGLKQADTVYFLPITPQFVTEVIKAERPDGLILGMGGQTALNCGVELFRRGVLKEYGVKVLGTSVEAIMATEDRQLFSDKLTEINEKIAPSFAVESIEDALNAAEKIGYPVMIRSAYALGGLGSGICPDKERLLDLGTKAFAVTNQILVERSVVGWKEIEYEVVRDASDNCITVCNMENVDAMGVHTGDSIVVAPSQTLSNEEFQMLRESAIKVVRHLGIVGECNIQYALHPTSLEYCIIEVNARLSRSSALASKATGYPLAFIAAKIALGIPLPEIKNVVSGKTSACFEPSLDYMVAKIPRWDLDRFQGTSSQIGSSMKSVGEVMAIGRTFEECFQKALRMCHPSVDGFTSRLPMNKEWPANLDLQKELSDPSSIRVYAIAKALHDGVPVDDIHKLTAIDRWFIYKMRDIVNMEKSLKELSSESIPEETMRRAKQIGFSDKQIGKCLGVTEAQSRELRLKKNIKPSVKQIDTLAAEYPAITNYLYVTYNGQENDVKFDDHGMMVLGCGPYHIGSSVEFDWCAVSSIRTLRQLGHKTVVVNCNPETVSTDFDECDRLYFEELSLERILDIYQHEKCAGCIISVGGQIPNNLAVPLYKNGVKIMGTNPMQIDRAEDRSVFSAVLDELNVAQAPWKAVSTLKDAVEFANSVGYPCLLRPSYVLSGSAMNVVFTEEEMKKFLAEATRVSQEHPVVLTKFIEGAREVEMDAVAKAGKVVSHAISEHVEDAGVHSGDATLMLPTQNISQGALEKVKIITKKIARAFSISGPFNVQFLVRGNDVLVIECNLRASRSFPFVSKTIGVDFIDVATKVMIGEKIDESSLPTLERPIIPSEYVGIKAPMFSWPRLRDADPILRCEMASTGEVACFGESIHSAFLKAMLSTGFKFPQKGILVGIQHSFRPKFLGVAEQLHEEGFKLYATEATSNWLNANSVPTTPVAWPSQESHTTSLPAIRRLIKDGNIDLVINLPNSNTKFVHDNYVIRRTAVDSGIALLTNFQVTKLFAEAIKYSGKVDAKSLFHYRKLNTSNPP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN62</accession>
    <name>A0A452IN62_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">C2H2-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IN62"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033278.1">
        <property type="protein sequence ID" value="ENSGAGP00000029299.1"/>
        <property type="gene ID" value="ENSGAGG00000021197.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001430">
        <property type="entry name" value="Uncharacterized protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000355">
        <property type="entry name" value="zinc finger and SCAN domain-containing protein 20 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000478">
        <property type="entry name" value="Zinc finger protein 133"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002343">
        <property type="entry name" value="Zinc finger protein 33A"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001343">
        <property type="entry name" value="Zinc finger protein 568"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001270">
        <property type="entry name" value="zinc finger protein 583 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000624">
        <property type="entry name" value="zinc finger protein 697"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050758">
        <property type="entry name" value="Znf_C2H2-type"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23234:SF8">
        <property type="entry name" value="C2H2-TYPE DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23234">
        <property type="entry name" value="ZNF44 PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="12"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="13"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="9" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 11" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="37"/>
            <end position="64"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="87"/>
            <end position="114"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="115"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="143"/>
            <end position="170"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="171"/>
            <end position="198"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="199"/>
            <end position="226"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="227"/>
            <end position="254"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="255"/>
            <end position="282"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="283"/>
            <end position="310"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="311"/>
            <end position="338"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="339"/>
            <end position="366"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="367"/>
            <end position="394"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="395"/>
            <end position="421"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029299.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="422" mass="49356" checksum="87802B6A1511CBF7" modified="2019-05-08" version="1">DGEERERGREKKPLQCLDHERNPLDVLSTRNLSTRPHKCLHCGKSFTRRSHLIQHGRIHTGERPHKCLDCGKILLSHLRTHTGEKPHKCLDCGKSFRYRSVLLSHLRTHTGEKPHKCLDCGKSFRYRSVLLSHLRTHTGEKPHKCLDCGKSFRYRSVLLSHLRTHTGEKPHKCLECGKSFMWRSTLVLHQRIHTGERPHKCLDCGKSFTRRSHLIQHGRIHTGERPHKCLDCGKSFRYRSVFLSHLRTHTGEKPHKCLDCGKSFRYRSVLLSHLRTHTGERPHKCLECGKSFMWRSTLVLHQRIHTGERPHKCLECGKSFMWRSALVLHQRIHTGERPHKCLECGKSFMWRSTLVLHQRIHTGERPHKCLECGKSFMWRSALVLHQRIHTGERPHKCLDCGKSFTRRSLIKHGRIHTASKIL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN64</accession>
    <name>A0A452IN64_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Ral guanine nucleotide dissociation stimulator</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IN64"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033519.1">
        <property type="protein sequence ID" value="ENSGAGP00000029519.1"/>
        <property type="gene ID" value="ENSGAGG00000021309.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007265">
        <property type="term" value="P:Ras protein signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd17209">
        <property type="entry name" value="RA_RalGDS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00155">
        <property type="entry name" value="RasGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd06224">
        <property type="entry name" value="REM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.840.10:FF:000005">
        <property type="entry name" value="Ral guanine nucleotide dissociation stimulator isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.20.90:FF:000042">
        <property type="entry name" value="Ral guanine nucleotide dissociation stimulator isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.20.90">
        <property type="entry name" value="Phosphatidylinositol 3-kinase Catalytic Subunit, Chain A, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.840.10">
        <property type="entry name" value="Ras guanine-nucleotide exchange factors catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.870.10">
        <property type="entry name" value="Son of sevenless (SoS) protein Chain: S domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000159">
        <property type="entry name" value="RA_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015758">
        <property type="entry name" value="RalGDS_RA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008937">
        <property type="entry name" value="Ras-like_GEF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000651">
        <property type="entry name" value="Ras-like_Gua-exchang_fac_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019804">
        <property type="entry name" value="Ras_G-nucl-exch_fac_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023578">
        <property type="entry name" value="Ras_GEF_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001895">
        <property type="entry name" value="RASGEF_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036964">
        <property type="entry name" value="RASGEF_cat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029071">
        <property type="entry name" value="Ubiquitin-like_domsf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23113">
        <property type="entry name" value="GUANINE NUCLEOTIDE EXCHANGE FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23113:SF35">
        <property type="entry name" value="RAL GUANINE NUCLEOTIDE DISSOCIATION STIMULATOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00788">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00617">
        <property type="entry name" value="RasGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00618">
        <property type="entry name" value="RasGEF_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00314">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00147">
        <property type="entry name" value="RasGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00229">
        <property type="entry name" value="RasGEFN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48366">
        <property type="entry name" value="Ras GEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54236">
        <property type="entry name" value="Ubiquitin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50200">
        <property type="entry name" value="RA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00720">
        <property type="entry name" value="RASGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50009">
        <property type="entry name" value="RASGEF_CAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50212">
        <property type="entry name" value="RASGEF_NTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1 2" id="KW-0344">Guanine-nucleotide releasing factor</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="N-terminal Ras-GEF" evidence="6">
        <location>
            <begin position="1"/>
            <end position="110"/>
        </location>
    </feature>
    <feature type="domain" description="Ras-GEF" evidence="4">
        <location>
            <begin position="181"/>
            <end position="446"/>
        </location>
    </feature>
    <feature type="domain" description="Ras-associating" evidence="5">
        <location>
            <begin position="597"/>
            <end position="684"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="550"/>
            <end position="571"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="3">
        <location>
            <begin position="550"/>
            <end position="570"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022658"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00168"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50009"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50200"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029519.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="9">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="714" mass="79709" checksum="FA2C888CA0CA8A8F" modified="2019-05-08" version="1">KAGTLEKLVEYLVSAFKGNDSTYVTIFLCTYRAFATTKQVLDLLFNSQSVGLLLPSTISSILGAWLDQYSEDFRKPPDFACLKQLISYVRQNIPGSDLERRARILLAQFQQQEHSEAEPDGEQRFLPVPVSLSALCGVSWVSPVTPVRRTVYEAVDQTSCTFRIVEENGLGEEKPDFLAFSQEMVAEQFTLMDAELFKKVVPYHCLGCIWSQRDKKGKEHLAPTIRATVSQFNSVTNCVIATCLGDRSLKPQQRAKVVERWIEVARECRILKNFSSLRAILSALQCNAVHRLKKTWDEVSRESFRTFHELSEIFSDENNHSLSRELLIKEGTSKFATLEINPKRAQKRQQQQREMGVMQGTIPYLGTFLTDLVMLDTAMKDFLDGGLINFEKRRKEFEVIAQIKLLQSACNNYSFTQEDGFVEWFHGVERLSEAESYSLSCEIEPLSESASNTLKAKKNTGIIKRWSDRQVPSAEPCTSGSSHSKSFDQLKCGQYLCSGDTADSVSVTSAGSSSSDVEEINISFIPESPDCQEKKFWECTSLSSLDTSGIGSGSSSASSSSVSSTPVTASRTHKRSVSGISSYSSLSLPLYNQQIDDCCIIRVSLAVDNGNMYKSILVTSQDKTPVVIRKAMAKHNLDGDRPEDYELVQIISEERELKIPDNANVFYAMNSAANYDFVLKKRGFSKGGVKIKHGSSSTLPRMKQKGLKIAKGIF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN65</accession>
    <name>A0A452IN65_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">C2H2-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">May be involved in transcriptional regulation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000033287.1">
        <property type="protein sequence ID" value="ENSGAGP00000029308.1"/>
        <property type="gene ID" value="ENSGAGG00000021197.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002063">
        <property type="entry name" value="RB associated KRAB zinc finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001772">
        <property type="entry name" value="Uncharacterized protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000247">
        <property type="entry name" value="Zinc finger protein 236"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002343">
        <property type="entry name" value="Zinc finger protein 33A"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001343">
        <property type="entry name" value="Zinc finger protein 568"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000624">
        <property type="entry name" value="zinc finger protein 697"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001005">
        <property type="entry name" value="Zinc finger protein 75A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050752">
        <property type="entry name" value="C2H2-ZF_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24384">
        <property type="entry name" value="FINGER PUTATIVE TRANSCRIPTION FACTOR FAMILY-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24384:SF218">
        <property type="entry name" value="ZINC FINGER PROTEIN 502"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="11"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="11" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="4" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="5" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13" id="KW-0539">Nucleus</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="12" id="KW-0804">Transcription</keyword>
    <keyword evidence="10" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="9" id="KW-0832">Ubl conjugation</keyword>
    <keyword evidence="8" id="KW-0862">Zinc</keyword>
    <keyword evidence="7 14" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="49"/>
            <end position="76"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="77"/>
            <end position="104"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="105"/>
            <end position="132"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="135"/>
            <end position="153"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="154"/>
            <end position="181"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="182"/>
            <end position="209"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="210"/>
            <end position="237"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="238"/>
            <end position="265"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="266"/>
            <end position="293"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="294"/>
            <end position="321"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="322"/>
            <end position="349"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00003767"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029308.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="361" mass="42206" checksum="98CD75EDF1AD7C88" modified="2019-05-08" version="1">MLENYETVTSLGKGFPSPRLLESVGPAMLMLVTFFPGQLGLWRXVEKPSRCLEHKKSFYCKSALITHQRIHTQERPHKCLHCGKSFTRRSHLIQHGRIHTGERPHKCLDCGKSFRYRSVFLSHLRTHTGEKPHNFRYRSVLLSHLRTHTGERPHKCLECGKSFMWRSTLVLHQRIHTGERPHKCLECGKSFMWRSALVLHQRIHTGERPHKCLECGKSFMWRSTLVLHQRIHTGERPHKCLECGKSFMWRSALVLHQRIHTGERPHKCLDCGKSFTRRSVLVSHQRIHIVQRPHKCLDCGKSFIRRSDLIKHGRIHTEERPHKCLDCGKSFTRRPALVSHQTIHTGERSLKCLDCGNFNCQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN66</accession>
    <name>A0A452IN66_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="17">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">May be involved in transcriptional regulation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IN66"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033283.1">
        <property type="protein sequence ID" value="ENSGAGP00000029304.1"/>
        <property type="gene ID" value="ENSGAGG00000021197.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd07765">
        <property type="entry name" value="KRAB_A-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002063">
        <property type="entry name" value="RB associated KRAB zinc finger"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001772">
        <property type="entry name" value="Uncharacterized protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000056">
        <property type="entry name" value="Zinc finger and SCAN domain-containing 20"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000478">
        <property type="entry name" value="Zinc finger protein 133"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000688">
        <property type="entry name" value="zinc finger protein 197 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000247">
        <property type="entry name" value="Zinc finger protein 236"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002343">
        <property type="entry name" value="Zinc finger protein 33A"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001119">
        <property type="entry name" value="zinc finger protein 408"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001343">
        <property type="entry name" value="Zinc finger protein 568"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000624">
        <property type="entry name" value="zinc finger protein 697"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000931">
        <property type="entry name" value="zinc finger protein 697"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001005">
        <property type="entry name" value="Zinc finger protein 75A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="27"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001909">
        <property type="entry name" value="KRAB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036051">
        <property type="entry name" value="KRAB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23235:SF178">
        <property type="entry name" value="C2H2-TYPE DOMAIN-CONTAINING PROTEIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23235">
        <property type="entry name" value="KRUEPPEL-LIKE TRANSCRIPTION FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01352">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="18"/>
    </dbReference>
    <dbReference type="SMART" id="SM00349">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="27"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="14"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109640">
        <property type="entry name" value="KRAB domain (Kruppel-associated box)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50805">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="24"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="27"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="11" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="4" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="5" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13" id="KW-0539">Nucleus</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="12" id="KW-0804">Transcription</keyword>
    <keyword evidence="10" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="9" id="KW-0832">Ubl conjugation</keyword>
    <keyword evidence="8" id="KW-0862">Zinc</keyword>
    <keyword evidence="7 14" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="KRAB" evidence="16">
        <location>
            <begin position="9"/>
            <end position="75"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="78"/>
            <end position="105"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="106"/>
            <end position="133"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="134"/>
            <end position="161"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="162"/>
            <end position="189"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="190"/>
            <end position="217"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="218"/>
            <end position="245"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="246"/>
            <end position="273"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="274"/>
            <end position="301"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="302"/>
            <end position="329"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="330"/>
            <end position="357"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="358"/>
            <end position="385"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="386"/>
            <end position="413"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="414"/>
            <end position="441"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="442"/>
            <end position="469"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="470"/>
            <end position="497"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="498"/>
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        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="526"/>
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        </location>
    </feature>
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        <location>
            <begin position="554"/>
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        </location>
    </feature>
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        <location>
            <begin position="582"/>
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        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="610"/>
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        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="638"/>
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        </location>
    </feature>
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        <location>
            <begin position="666"/>
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        </location>
    </feature>
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        <location>
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        </location>
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        <location>
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        <location>
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            <end position="777"/>
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    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
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        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="15">
        <location>
            <begin position="806"/>
            <end position="833"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00003767"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
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    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
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    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029304.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="834" mass="97320" checksum="8717BB637F1A1C7E" modified="2019-05-08" version="1">HPVFQEVPVTFEEVAVYFTQRQGALLDPAQTALYRDVMLENYETVTSLGKGFPSPRTDTETIWLMKEKQTNPNEENPYQCLKCGKGLILRSQLVTHQTIHTREKPLQCLDHGKTFNNRSDLKIHGRSHTREKPSRCLEHKKSFYCKSALITHQRIHTQERPHKCLHCGKSFTRRSHLIQHGRIHTGERPHKCLDCGKSFRYRSVFLSHLRTHTGEKPHKCLDCGKSFRYRSVLLSHLRTHTGEKPHKCLDCGKSFRYRSVLLSHLRTHTGEKPHKCLDCGKSFRYRSVLLSHLRTHTGEKPHKCLDCGKSFRYRSVLLSHLRTHTGEKPHKCLECGKSFMWRSTLVLHQRIHTGERPHKCLDCGKSFTRRSHLIQHGRIHTGERPHKCLDCGKSFRYRSVFLSHLRTHTGEKPHKCLDCGKSFRYRSVLLSHLRTHTGERPHKCLECGKSFMWRSTLVLHQRIHTGERPHKCLECGKSFMWRSALVLHQRIHTGERPHKCLECGKSFMWRSTLVLHQRIHTGERPHKCLECGKSFMWRSALVLHQRIHTGERPHKCLDCGKSFTRRSVLVSHQRIHIVQRPHKCLDCGKSFIRRSDLIKHGRIHTEERPHKCLDCGKSFTRRPALVSHQTIHTGERSLKCLDCGNCFRWKSNLIRHQRIHTVESPHKCLACGKSFMRRSDLDLHQRIHTGERPHRCLDCGKSFTWRSALVKHGRTHTGERPHKCLDCGKSFTWRSVLVKHGRIHTGERPHKCLDCGKSFTWRSALVSHQRIHTGERPHKCLDCGKSFIWRSALVSHKRIHTEERPHKGLDCGKSFTQRSKLIKHGRIHTASKIL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN67</accession>
    <name>A0A452IN67_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">AKNA domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IN67"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033314.1">
        <property type="protein sequence ID" value="ENSGAGP00000029335.1"/>
        <property type="gene ID" value="ENSGAGG00000021212.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052655">
        <property type="entry name" value="AKNA_Centrosome-Trans_reg"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022150">
        <property type="entry name" value="AKNA_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21510">
        <property type="entry name" value="AKNA DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21510:SF16">
        <property type="entry name" value="PROTEIN AKNAD1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12443">
        <property type="entry name" value="AKNA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="AKNA" evidence="3">
        <location>
            <begin position="440"/>
            <end position="527"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="320"/>
            <end position="340"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="629"/>
            <end position="656"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="489"/>
            <end position="551"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="645"/>
            <end position="656"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="Pfam" id="PF12443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029335.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1059" mass="120560" checksum="31AC387A59BCD7AB" modified="2019-05-08" version="1">MEYNKDIQINTSKSRQPDHLKKQVNNILLDNFDYPEDATDEEQEDLPYDGDLERTYHHNNESNNLKACVSVENISNNFLNLTCSEINRSLKEEINYETQQLSQEKVFSHRMPATRTNGIMTEMAIEAPVSGMPLETELSVGAFSSLDRKEHFTNSNISDVLLRHFSKEELSNTGQLIDCETIPETSFTESVDETVLTKTRISECTKPTLRTEQWAKHFEDYHLNRQEEKSEDDYEDKHLIDENKFVIDDRATSYGDEKNCIQENSQLIAENEDTNNFKNIRNNHSHQKGLFERTGSSHELKYGQGQVNYCLPDFSKVPPKVKIPKRNNSDKSAPTMKRTKFSPNLVGKSAIVNDVLETINYFDSVEVKNQEEEMRIPELLQQLELLTKHAEAQNRIDHLRFDPKIYTHSASPNPNLSIHSRCMGAASEESVFHPLTVPIQPLLGLSQASFSGLQSGKIISSLPSASSAGEVHCLNPNLMQNTTKGEKMSQMLKEQAEQLKAKVEEFSKCVIQEAFPLQDRCLVLKELKGYLDTLERKYLATKEEHRGLQLQNYNHKSISVGEFDPDRKVEGEIFRLGMLLEDVKEKIDDSICSPCPSSFTTSPMSPSLTQSESVCSSCSCLRESPVVSSISDPPERSATEVNFHNNERNKGENRSHAIDVIPKKTSQLSLQNDNCDPFPEIQLGLQKRESCAYVKGMEPLEKAGLLANKHSSDVMQCFLPHEADNGPGGLESHRQLILSQKRNIDQENPKGSLNFWQMKSPPKTKPYNMCNLEHKIINLNVRQEQGNLAHPSDHCFCGRVEKCSNAHDAALYPKWKIHWSKNANKELCSCSVNRNKKTEDRKTDSERSKCGRYSVFIQEKAVELDLSGSDSEDISFCDSFLESHSDEFLEHRTKNYKSRRAELKEQSKALQYRDWNINFCTSGQRNKCSRESDKNNIASSQKKRIVANSVCTSRQPDQFVYRISDKDNLETPQTCYSGMDDTIILSPQHLASRKDYGNKSIINIRNRHTSYTDTKILNSTLDHAIQTATSLQETTERMVQVVAEDLAKVKTHRSVHLES</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN75</accession>
    <name>A0A452IN75_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Eyes absent homolog</fullName>
            <ecNumber evidence="15">3.1.3.48</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="12 15">
            <text>O-phospho-L-tyrosyl-[protein] + H2O = L-tyrosyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:10684"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="EC" id="3.1.3.48"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="14 15">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
        <text evidence="14 15">Binds 1 Mg(2+) ion per subunit.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 15">Belongs to the HAD-like hydrolase superfamily. EYA family.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.48" evidence="15"/>
    <dbReference type="AlphaFoldDB" id="A0A452IN75"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033393.1">
        <property type="protein sequence ID" value="ENSGAGP00000029407.1"/>
        <property type="gene ID" value="ENSGAGG00000021258.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
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        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
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        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:2001240">
        <property type="term" value="P:negative regulation of extrinsic apoptotic signaling pathway in absence of ligand"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="P:positive regulation of DNA repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="entry name" value="HAD_Eya"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.12350:FF:000001">
        <property type="entry name" value="Eyes absent homolog"/>
        <property type="match status" value="1"/>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028472">
        <property type="entry name" value="EYA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006545">
        <property type="entry name" value="EYA_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042577">
        <property type="entry name" value="EYA_dom_metazoan"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038102">
        <property type="entry name" value="EYA_dom_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01658">
        <property type="entry name" value="EYA-cons_domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10190">
        <property type="entry name" value="EYES ABSENT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10190:SF17">
        <property type="entry name" value="EYES ABSENT HOMOLOG 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00702">
        <property type="entry name" value="Hydrolase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDG01129">
        <property type="entry name" value="C1.5:_HAD__Beta-PGM__Phosphata"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDS00003">
        <property type="entry name" value="Haloacid_Dehalogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0010">Activator</keyword>
    <keyword evidence="3" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="5 15" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="6 14 15" id="KW-0460">Magnesium</keyword>
    <keyword evidence="4 14 15" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="7 15" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0804">Transcription</keyword>
    <keyword evidence="8 15" id="KW-0805">Transcription regulation</keyword>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1"/>
            <end position="88"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="154"/>
            <end position="180"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="245"/>
            <end position="314"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="18"/>
            <end position="43"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="16">
        <location>
            <begin position="44"/>
            <end position="62"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="63"/>
            <end position="88"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="16">
        <location>
            <begin position="249"/>
            <end position="263"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="264"/>
            <end position="280"/>
        </location>
    </feature>
    <feature type="active site" description="Nucleophile" evidence="13">
        <location>
            <position position="321"/>
        </location>
    </feature>
    <feature type="active site" description="Proton donor" evidence="13">
        <location>
            <position position="323"/>
        </location>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="321"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="323"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="549"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
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            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
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    </evidence>
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        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
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            <dbReference type="RuleBase" id="RU362036"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="585" mass="64112" checksum="7922EB07E7633309" modified="2019-05-08" version="1">MEDSQDLNEQSVKKTCTESDVSEAQNSRSMEMQDLTSPHNLVGSSDNPSSSKLDKSNLSSTSITTNGTGVITSSGYSPRSAHQYSPQIYPSKPYPHILSTPAAQSMSAYSGQTQYSGMQQPAVYTAYSQTGQPYSLPTYDLGVMLPGIKTESGLSQTQSPLQSGCLSYSPGFSTPQPGQTPYSYQMPGSSFTPSSTIYANNSVSNSTNFSSSQQDYPSYTAFGQNQYAQYYSASTYGAYMTSNNTADGTSSSSSTYQLQESLSGLNSQPGEFDTVQSPSTPIKDLDERTCRSSGSKSRGRGRKNNPSPPPDSDLERVFVWDLDETIIVFHSLLTGSYAQKYGKDPPMAVTLGLRMEEMIFNLADTHLFFNDLEECDQVHIDDVSSDDNGQDLSTYSFATDGFHAAASSANLCLPTGVRGGVDWMRKLAFRYRRVKELYNTYKNNIGGLLGPAKRDAWLQLRAEIEALTDSWLTNALKSLSIISTRSNCVNVLVTTTQLIPALAKVLLYSLGGAFPIENIYSATKIGKESCFERIMQRFGRKVVYVVIGDGVEEEQAAKKHNMPFWRISSHSDLLALHQALELEYL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN78</accession>
    <name>A0A452IN78_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="6">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
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    <dbReference type="AlphaFoldDB" id="A0A452IN78"/>
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    </dbReference>
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        <property type="project" value="UniProtKB-SubCell"/>
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    </dbReference>
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        <property type="entry name" value="Solute_trans_a"/>
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    </dbReference>
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            <end position="63"/>
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    </evidence>
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    </evidence>
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        </source>
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        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="410" mass="46252" checksum="8FA05F5D011DC4F8" modified="2019-05-08" version="1">MIVVTGIMSAPELGSTATTPVAAPNFSWMSDDGSPTAMEQPIFLMTTAAQAISGFFVWTALLITCHQIYMHLRCYSCPNEQRYIVRILFIVPIYAFDSWLSLLFFTNDQYYVYFGTVRDCYEAFVIYNFLSLCYEYLGGESSIMSEIRGKSIESSCMYGTCCLWGKTYSIGFLRFCKQATLQFCVVKPLMAISTVILQAFGKYRDGDFDVTRGYLYVTIIYNISVSLALYALFLFYFATRELLSPYSPVLKFFMVKSVIFLSFWQGMLLAILEKCGAIPKIHSANVSVGEGTVAAGYQDFIICVEMFFAALALRHAFTYKVYADKHLDAQGRCAPMKSISSSLKETMNPHDIVQDAIHNFSPAYQQYTQQSTLEHGPPWRNGSHIISRSHSCSGARDNEKTLLLSSDDEF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN80</accession>
    <name>A0A452IN80_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Chloride channel CLIC-like protein 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the chloride channel MCLC family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IN80"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033330.1">
        <property type="protein sequence ID" value="ENSGAGP00000029350.1"/>
        <property type="gene ID" value="ENSGAGG00000021220.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005783">
        <property type="term" value="C:endoplasmic reticulum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005254">
        <property type="term" value="F:chloride channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009231">
        <property type="entry name" value="Chloride_chnl_CLIC-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR34093">
        <property type="entry name" value="CHLORIDE CHANNEL CLIC-LIKE PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR34093:SF1">
        <property type="entry name" value="CHLORIDE CHANNEL CLIC-LIKE PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05934">
        <property type="entry name" value="MCLC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="6 9" id="KW-0472">Membrane</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0732">Signal</keyword>
    <keyword evidence="4 9" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 9" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="10">
        <location>
            <begin position="1"/>
            <end position="16"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019081732" description="Chloride channel CLIC-like protein 1" evidence="10">
        <location>
            <begin position="17"/>
            <end position="548"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="188"/>
            <end position="209"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="221"/>
            <end position="238"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="337"/>
            <end position="358"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="393"/>
            <end position="433"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="493"/>
            <end position="548"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="68"/>
            <end position="95"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="412"/>
            <end position="421"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="515"/>
            <end position="526"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005944"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00015571"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029350.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="548" mass="62381" checksum="DA9452C70188875A" modified="2019-05-08" version="1">MLVPLVLCATLMIGSGEIQDDEWIDPTDMLNYDAASGTMRRPAKVNYDDSENREKPDAVVSLPSTAAVSECHRKLESLIQKIEEYEKKEKTKLSESHSIYVFRRYLNKILIEAGRLGLPDENIGDVHYDAEIILTSQALLEINRFLNEEDWKSGALDDAVNNILINFKHHNFEVWKWRFEDTFGIDPYNLLMVLLCLLCITVTVATELWTHIGWYTQVKRILFISFLISCGWNWIYLYKIAFAQHQAEVARMGNYDTVCDKKIDWSESLFEWLRSSWTFQDDPCQKYYETILVNPIWLVPPTKVLAVTFTNFVTEPLKHIGKGIGEFIKALMKEIPMMLQIPVLIILAVAVLGFCYGAGRSVATLKHLTGPDRKPSPSLPSSDRQHWEQIHYSAGDSDSNYRRNVGAISGGPHDRGDDHMRLQNGNRSSEVLLPGGTPNAQREEHYKCGLHPVSQDQTIFRVKTNVDQNCDREIALEEQKLCTLGVEQESEKNCEPQASCSLKKEKTNKQNSVEKPGDLEKTEKGSLSRLMEGTEETKEPLNSTVSFV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN81</accession>
    <name>A0A452IN81_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="9">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000033298.1">
        <property type="protein sequence ID" value="ENSGAGP00000029319.1"/>
        <property type="gene ID" value="ENSGAGG00000020783.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005938">
        <property type="term" value="C:cell cortex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0120025">
        <property type="term" value="C:plasma membrane bounded cell projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903479">
        <property type="term" value="P:mitotic actomyosin contractile ring assembly actin filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd21276">
        <property type="entry name" value="CH_IQGAP3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.418.10:FF:000013">
        <property type="entry name" value="IQ motif containing GTPase activating protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.506.10:FF:000004">
        <property type="entry name" value="IQ motif containing GTPase activating protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.190">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.418.10">
        <property type="entry name" value="Calponin-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.506.10">
        <property type="entry name" value="GTPase Activation - p120gap, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001715">
        <property type="entry name" value="CH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036872">
        <property type="entry name" value="CH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000048">
        <property type="entry name" value="IQ_motif_EF-hand-BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000593">
        <property type="entry name" value="RasGAP_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023152">
        <property type="entry name" value="RasGAP_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001936">
        <property type="entry name" value="RasGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001202">
        <property type="entry name" value="WW_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149">
        <property type="entry name" value="RAS GTPASE-ACTIVATING PROTEIN WITH IQ MOTIF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149:SF10">
        <property type="entry name" value="RAS GTPASE-ACTIVATING-LIKE PROTEIN IQGAP3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00307">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00612">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00616">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03836">
        <property type="entry name" value="RasGAP_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00033">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00015">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00323">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47576">
        <property type="entry name" value="Calponin-homology domain, CH-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF143885">
        <property type="entry name" value="RGC domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50021">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00509">
        <property type="entry name" value="RAS_GTPASE_ACTIV_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50018">
        <property type="entry name" value="RAS_GTPASE_ACTIV_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01159">
        <property type="entry name" value="WW_DOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50020">
        <property type="entry name" value="WW_DOMAIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="4" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="1" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="Calponin-homology (CH)" evidence="8">
        <location>
            <begin position="44"/>
            <end position="159"/>
        </location>
    </feature>
    <feature type="domain" description="WW" evidence="7">
        <location>
            <begin position="675"/>
            <end position="708"/>
        </location>
    </feature>
    <feature type="domain" description="Ras-GAP" evidence="6">
        <location>
            <begin position="974"/>
            <end position="1207"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="1367"/>
            <end position="1388"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="4">
        <location>
            <begin position="168"/>
            <end position="195"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50020"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029319.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1587" mass="180164" checksum="EB0F625433038A90" modified="2019-05-08" version="1">DSLVVEVAGICVLGTEGAHFSSVPRLTADEMDEQRRQNVAYQYLCRLEEAKRWMEACLNEELASPTELEESLRNGVILAKLGHCFAPDIIPLKKIYDREQMRYEATGLHFRHTDNINHWREAMAHLGLPTIFHPETTDIYDKKNMPRVVYCIHALSLYLFKLGLAPQIQDLYGKVNFTEEEINNMKRELEKYGLQMPAFSKIGGILASELSVDEAAVHAAVLAINEAVERGVVEQTMVALHNPSAMLLNLRENLAAVYQEMLFQVKMEKAGNARNRESPESEDIYDRCLTQAEIQGNINKVNVHGALEYVDDALERQDPLALCQALQDPVLALRGVQRENAAWYLDQLSTDREQKALLGYVDLLEQEEVQAGVYAANERGDQEHAMRRAVSRINASIRRGEAMETVRELMSPEAQLPKAFPFAAQLYQHELSLLQQHHPQLGQEELYVAVEMLSAVALIDRALEAGDVSGFWSSLVSPALGLSDIEDANAQRYFDDLLKLKRRSGKAGVAFLSWNDIQACVNSANLAVQDENDKILAIRLINEALAQDDPEKTLTALLLPSARFSAVTLPGARRYHDVLTAALKQKAQVTGDDGAALWWEEIQQGVSKANQDTEAARRLALGIAAINQAIKEGKPAQTARVLRNPDVSLRGIVAECAGAYQAQLAALLAAKRQAGNANHNWVRHQLKDGSVYYFSLQNFEGSWERPHDCVLNSTHLSREEIQSTITGVTAAHDRERLWESNAGFVVRLQARMRGYLVRQEFVARRHLLQEQLPAVIKIQVRSAARLPSRSHADGHDDIYIALGLQGINAVIKIQAFVRANKARGDYRMLVHARDPPLSIVRRFVHLLEQSQHDFWEESELLKLREEVVKRIRSNQQLESDLNLMDIKIGLLVKNRITLQEVVSHCKKLTRKNKEQLSNLMAIDKQKGLKSLSKEKRQKLEAYQHLFYLLQTHPVYLAKLIFQMPQNKSTKFMESVVFTLYNYASNPREAYLLLQLFKTALQEEIRSKVDQLRDILTGNPTVIRLVVSFYRNARGQNALRQILGSPVQEVLQDKTLSIRTNPVDIYKAWINQLESQSGQKSKFPYDVSPEQALSHPEVQRRLDISIRNLLAVTDKFLSAITSSVDKIPYGMRYVAKVLKTSLAEKFPAAGEEDICKIVGNLLYYRFMNPAVVAPDGFDIVDISAGVALHPEHRRNLGSIAKILQHAAANKMFEGENGHLRVVNQYLEETHSKFRLFICAACSVPEPEERFNVDEYSEMVAVAKPVIYITVAELINTHKLLLEHQDSISPDHKDPLHELLEDLDELPTVQSLIGTCRLLVSFKPQALAQLGKQEVSLTLTNKFDAVEASSEEADTRTFHHHQMHRRALREAQTPDRLKRNRSLAGNSQLSMEEKKRKVIRNLRRLENLGVVDSANQYQEIINEMAKDIRNQRRYRQHRKGELVKLRQTLCGLNSKTSFYEEQIDYYNQYIRTCLDNLAASSKLNGKSKKQQSLHYTAARLFEKGVLLEIEDLPVSQFRNVIFDIIPCGEAGKFQVKAKFMGIDMEKFQLHYQDLLQLQYEGVAVMKLFDKAKVNVNLLIFLLNKKFFRK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN83</accession>
    <name>A0A452IN83_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="8">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000033307.1">
        <property type="protein sequence ID" value="ENSGAGP00000029328.1"/>
        <property type="gene ID" value="ENSGAGG00000020783.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005938">
        <property type="term" value="C:cell cortex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0120025">
        <property type="term" value="C:plasma membrane bounded cell projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903479">
        <property type="term" value="P:mitotic actomyosin contractile ring assembly actin filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd21276">
        <property type="entry name" value="CH_IQGAP3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.418.10:FF:000013">
        <property type="entry name" value="IQ motif containing GTPase activating protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.506.10:FF:000004">
        <property type="entry name" value="IQ motif containing GTPase activating protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.190">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.418.10">
        <property type="entry name" value="Calponin-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.506.10">
        <property type="entry name" value="GTPase Activation - p120gap, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001715">
        <property type="entry name" value="CH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036872">
        <property type="entry name" value="CH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000048">
        <property type="entry name" value="IQ_motif_EF-hand-BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000593">
        <property type="entry name" value="RasGAP_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023152">
        <property type="entry name" value="RasGAP_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001936">
        <property type="entry name" value="RasGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001202">
        <property type="entry name" value="WW_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149">
        <property type="entry name" value="RAS GTPASE-ACTIVATING PROTEIN WITH IQ MOTIF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149:SF10">
        <property type="entry name" value="RAS GTPASE-ACTIVATING-LIKE PROTEIN IQGAP3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00307">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00612">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00616">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03836">
        <property type="entry name" value="RasGAP_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00033">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00015">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00323">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47576">
        <property type="entry name" value="Calponin-homology domain, CH-domain"/>
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    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF143885">
        <property type="entry name" value="RGC domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50021">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00509">
        <property type="entry name" value="RAS_GTPASE_ACTIV_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50018">
        <property type="entry name" value="RAS_GTPASE_ACTIV_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01159">
        <property type="entry name" value="WW_DOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50020">
        <property type="entry name" value="WW_DOMAIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="4" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="1" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="Calponin-homology (CH)" evidence="7">
        <location>
            <begin position="44"/>
            <end position="159"/>
        </location>
    </feature>
    <feature type="domain" description="WW" evidence="6">
        <location>
            <begin position="654"/>
            <end position="687"/>
        </location>
    </feature>
    <feature type="domain" description="Ras-GAP" evidence="5">
        <location>
            <begin position="953"/>
            <end position="1186"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="4">
        <location>
            <begin position="168"/>
            <end position="195"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50020"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029328.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1577" mass="178944" checksum="E35724FED6E275DA" modified="2019-05-08" version="1">DSLVVEVAGICVLGTEGAHFSSVPRLTADEMDEQRRQNVAYQYLCRLEEAKRWMEACLNEELASPTELEESLRNGVILAKLGHCFAPDIIPLKKIYDREQMRYEATGLHFRHTDNINHWREAMAHLGLPTIFHPETTDIYDKKNMPRVVYCIHALSLYLFKLGLAPQIQDLYGKVNFTEEEINNMKRELEKYGLQMPAFSKIGGILASELSVDEAAVHAAVLAINEAVERGVVEQTMVALHNPSAMLLNLRENLAAVYQEMLFQVKMEKAGNARNRVRAEHLVKGGTPLPALEYVDDALERQDPLALCQALQDPVLALRGVQRENAAWYLDQLSTDREQKRCLKTSPEFRPPQLICLVLSRGRAVRRAVSRINASIRRGEAMETVRELMSPEAQLPKAFPFAAQLYQHELSLLQQHHPQLGQEELYVAVEMLSAVALIDRALEAGDVSGFWSSLVSPALGLSDIEDANAQRYFDDLLKLKRRSGKAGVAFLSWNDIQACVNSANLAVQDENDKILAIRLINEALAQDDPEKTLTALLLPSARFSAVTLPGARRYHDVLTAALKQKAQVTGDDGAALWWEEIQQGVSKANQDTEAARRLALGIAAINQAIKEGKPAQTARVLRNPDVSLRGIVAECAGAYQAQLAALLAAKRQAGNANHNWVRHQLKDGSVYYFSLQNFEGSWERPHDCVLNSTHLSREEIQSTITGVTAAHDRERLWESNAGFVVRLQARMRGYLVRQEFVARRHLLQEQLPAVIKIQVRSAARLPSRSHADGHDDIYIALGLQGINAVIKIQAFVRANKARGDYRMLVHARDPPLSIVRRFVHLLEQSQHDFWEESELLKLREEVVKRIRSNQQLESDLNLMDIKIGLLVKNRITLQEVVSHCKKLTRKNKEQLSNLMAIDKQKGLKSLSKEKRQKLEAYQHLFYLLQTHPVYLAKLIFQMPQNKSTKFMESVVFTLYNYASNPREAYLLLQLFKTALQEEIRSKVDQLRDILTGNPTVIRLVVSFYRNARGQNALRQILGSPVQEVLQDKTLSIRTNPVDIYKAWINQLESQSGQKSKFPYDVSPEQALSHPEVQRRLDISIRNLLAVTDKFLSAITSSVDKIPYGMRYVAKVLKTSLAEKFPAAGEEDICKIVGNLLYYRFMNPAVVAPDGFDIVDISAGVALHPEHRRNLGSIAKILQHAAANKMFEGENGHLRVVNQYLEETHSKFRLFICAACSVPEPEERFNVDEYSEMVAVAKPVIYITVAELINTHKLLLEHQDSISPDHKDPLHELLEDLDELPTVQSLIGTCRLLVSFKPAPPLVSSFNPQNGTKQMLVDVIQSQPGDSLLEILRTAASEDQEAFHHHQMHRRALREAQTPDRLKRNRSLAGNSQLSMEEKKRKVIRNLRRLENLGVVDSANQYQEIINEMAKDIRNQRRYRQHRKGELVKLRQTLCGLNSKTSFYEEQIDYYNQYIRTCLDNLAASSKLNGKSKKQQSLHYTAARLFEKGVLLEIEDLPVSQFRNVIFDIIPCGEAGKFQVKAKFMGIDMEKFQLHYQDLLQLQYEGVAVMKLFDKAKVNVNLLIFLLNKKFFRK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN86</accession>
    <name>A0A452IN86_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">SMB domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IN86"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033335.1">
        <property type="protein sequence ID" value="ENSGAGP00000029355.1"/>
        <property type="gene ID" value="ENSGAGG00000020186.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009986">
        <property type="term" value="C:cell surface"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004551">
        <property type="term" value="F:dinucleotide phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003676">
        <property type="term" value="F:nucleic acid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004528">
        <property type="term" value="F:phosphodiesterase I activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046034">
        <property type="term" value="P:ATP metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030505">
        <property type="term" value="P:inorganic diphosphate transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045599">
        <property type="term" value="P:negative regulation of fat cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009143">
        <property type="term" value="P:nucleoside triphosphate catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030500">
        <property type="term" value="P:regulation of bone mineralization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd16018">
        <property type="entry name" value="Enpp"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00091">
        <property type="entry name" value="NUC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.410.20:FF:000003">
        <property type="entry name" value="Ectonucleotide pyrophosphatase/phosphodiesterase family member 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.410.20:FF:000001">
        <property type="entry name" value="Ectonucleotide pyrophosphatase/phosphodiesterase family member 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.720.10:FF:000145">
        <property type="entry name" value="Uncharacterized protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.410.20">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.720.10">
        <property type="entry name" value="Alkaline Phosphatase, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.570.10">
        <property type="entry name" value="Extracellular Endonuclease, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017850">
        <property type="entry name" value="Alkaline_phosphatase_core_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044929">
        <property type="entry name" value="DNA/RNA_non-sp_Endonuclease_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001604">
        <property type="entry name" value="Endo_G_ENPP1-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020821">
        <property type="entry name" value="ENPP1-3/EXOG-like_nuc-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044925">
        <property type="entry name" value="His-Me_finger_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002591">
        <property type="entry name" value="Phosphodiest/P_Trfase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036024">
        <property type="entry name" value="Somatomedin_B-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001212">
        <property type="entry name" value="Somatomedin_B_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10151">
        <property type="entry name" value="ECTONUCLEOTIDE PYROPHOSPHATASE/PHOSPHODIESTERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10151:SF77">
        <property type="entry name" value="ECTONUCLEOTIDE PYROPHOSPHATASE_PHOSPHODIESTERASE FAMILY MEMBER 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01223">
        <property type="entry name" value="Endonuclease_NS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01663">
        <property type="entry name" value="Phosphodiest"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01033">
        <property type="entry name" value="Somatomedin_B"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00892">
        <property type="entry name" value="Endonuclease_NS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00477">
        <property type="entry name" value="NUC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00201">
        <property type="entry name" value="SO"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53649">
        <property type="entry name" value="Alkaline phosphatase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54060">
        <property type="entry name" value="His-Me finger endonucleases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90188">
        <property type="entry name" value="Somatomedin B domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00524">
        <property type="entry name" value="SMB_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50958">
        <property type="entry name" value="SMB_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="4" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <feature type="domain" description="SMB" evidence="7">
        <location>
            <begin position="36"/>
            <end position="76"/>
        </location>
    </feature>
    <feature type="domain" description="SMB" evidence="7">
        <location>
            <begin position="77"/>
            <end position="121"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50958"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029355.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="844" mass="96761" checksum="F3D48F4FEBF9530D" modified="2019-05-08" version="1">FKGLGALAAATVVLSACVLTIILGCIFGLKPSCSRDVKTCKGRCFERTFGSCRCDRDCVELGNCCLDFQETCIQPAHIWTCNKFRCGEKRWPENRCSCSDDCVENKDCCVNYYIVCQGAKSWVEEKCEDINEPQCPPGFAKPPVLLFSLDGFRAEYLHTWSGLLPVISKLQKCGTYTPNMRPVYPTKTFPNHYTIVTGLYPESHGIIDNKIYDPHRNVSFTLKNAEKFNPQWYQGQPIWLTAMHQGLKAGTFFWPGSDVRINGTFPDLYRKYNGSIPFEERVIAILHWLQLPKDERPHFYTLYLEEPDSSGHNFGPVSSGVILALQRVDNIVGMLMDGLKQMNLHKCLNIILLSDHGMETASCSKTAYLSTYMENVQDITVIPGPAARLRPQNVPDEYFSCLENLTCEEDPDQHFKAFMKELLPKRFHYVNNDRIERVHFYLDPQWQLARKPSEIKYCNGGFHGSDNRFMSMQAIFIGFGPGFKFKTQVDPFENIEIYNLICDLLGITPAYNNGTHGSLNHLLKNPVYIPRHPEEESHPSVYPLMKTTSSLDIGCSCDLAGLPIRDFQQQLNHTASEVKKIEKYSLPYGRPRVLQKRSTYYLLYHHQYVSGYSQDLNMTLWSAYTDNRTASVENLSNCLHVDIRIPSSSSKTCSFYKNHHQLSYGFLFPPSKFTSSYEVLLNSNIVPMYHEFKVLWNYFYDVLLPKYTLARNGVNVVSGPVFDYDSNGLYDTPEKVKRLTHDPEVLVPTHYFIVLTSCKNISQTPLQCEGALDALSFIVPHRPDNSESCRHKRDSLWVEERMRFHTARVRDVELLTGLSFYQDRKQPVSDILQLKTYLPTFPME</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN87</accession>
    <name>A0A452IN87_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">AKNA domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IN87"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033303.1">
        <property type="protein sequence ID" value="ENSGAGP00000029324.1"/>
        <property type="gene ID" value="ENSGAGG00000021212.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052655">
        <property type="entry name" value="AKNA_Centrosome-Trans_reg"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022150">
        <property type="entry name" value="AKNA_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21510">
        <property type="entry name" value="AKNA DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21510:SF16">
        <property type="entry name" value="PROTEIN AKNAD1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12443">
        <property type="entry name" value="AKNA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="AKNA" evidence="3">
        <location>
            <begin position="440"/>
            <end position="527"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="320"/>
            <end position="340"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="629"/>
            <end position="656"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="489"/>
            <end position="551"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="645"/>
            <end position="656"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="Pfam" id="PF12443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029324.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1029" mass="117220" checksum="A09B3830D6C572ED" modified="2019-05-08" version="1">MEYNKDIQINTSKSRQPDHLKKQVNNILLDNFDYPEDATDEEQEDLPYDGDLERTYHHNNESNNLKACVSVENISNNFLNLTCSEINRSLKEEINYETQQLSQEKVFSHRMPATRTNGIMTEMAIEAPVSGMPLETELSVGAFSSLDRKEHFTNSNISDVLLRHFSKEELSNTGQLIDCETIPETSFTESVDETVLTKTRISECTKPTLRTEQWAKHFEDYHLNRQEEKSEDDYEDKHLIDENKFVIDDRATSYGDEKNCIQENSQLIAENEDTNNFKNIRNNHSHQKGLFERTGSSHELKYGQGQVNYCLPDFSKVPPKVKIPKRNNSDKSAPTMKRTKFSPNLVGKSAIVNDVLETINYFDSVEVKNQEEEMRIPELLQQLELLTKHAEAQNRIDHLRFDPKIYTHSASPNPNLSIHSRCMGAASEESVFHPLTVPIQPLLGLSQASFSGLQSGKIISSLPSASSAGEVHCLNPNLMQNTTKGEKMSQMLKEQAEQLKAKVEEFSKCVIQEAFPLQDRCLVLKELKGYLDTLERKYLATKEEHRGLQLQNYNHKSISVGEFDPDRKVEGEIFRLGMLLEDVKEKIDDSICSPCPSSFTTSPMSPSLTQSESVCSSCSCLRESPVVSSISDPPERSATEVNFHNNERNKGENRSHAIDVIPKKTSQLSLQNDNCDPFPEIQLGLQKRESCAYVKGMEPLEKAGLLANKHSSDVMQCFLPHEADNGPGGLESHRQLILSQKRNIDQENPKGSLNFWQMKSPPKTKPYNMCNLEHKIINLNVRQEQGNLAHPSDHCFCGRVEKCSNAHDAALYPKWKIHWSKNANKELCSCSVNRNKKTEDRKTDSERSKCGRYSVFIQEKAVELDLSALQYRDWNINFCTSGQRNKCSRESDKNNIASSQKKRIVANSVCTSRQPDQFVYRISDKDNLETPQTCYSGMDDTIILSPQHLASRKDYGNKSIINIRNRHTSYTDTKILNSTLDHAIQTATSLQETTERMVQVVAEDLAKVKTHRRRRLKLNILSRNTIKLR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN91</accession>
    <name>A0A452IN91_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="8">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000033308.1">
        <property type="protein sequence ID" value="ENSGAGP00000029329.1"/>
        <property type="gene ID" value="ENSGAGG00000020783.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005938">
        <property type="term" value="C:cell cortex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0120025">
        <property type="term" value="C:plasma membrane bounded cell projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903479">
        <property type="term" value="P:mitotic actomyosin contractile ring assembly actin filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd21276">
        <property type="entry name" value="CH_IQGAP3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.418.10:FF:000013">
        <property type="entry name" value="IQ motif containing GTPase activating protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.506.10:FF:000004">
        <property type="entry name" value="IQ motif containing GTPase activating protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.190">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.418.10">
        <property type="entry name" value="Calponin-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.506.10">
        <property type="entry name" value="GTPase Activation - p120gap, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001715">
        <property type="entry name" value="CH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036872">
        <property type="entry name" value="CH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000048">
        <property type="entry name" value="IQ_motif_EF-hand-BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000593">
        <property type="entry name" value="RasGAP_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023152">
        <property type="entry name" value="RasGAP_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001936">
        <property type="entry name" value="RasGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001202">
        <property type="entry name" value="WW_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149">
        <property type="entry name" value="RAS GTPASE-ACTIVATING PROTEIN WITH IQ MOTIF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149:SF10">
        <property type="entry name" value="RAS GTPASE-ACTIVATING-LIKE PROTEIN IQGAP3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00307">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00612">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00616">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03836">
        <property type="entry name" value="RasGAP_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00033">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00015">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00323">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47576">
        <property type="entry name" value="Calponin-homology domain, CH-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF143885">
        <property type="entry name" value="RGC domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50021">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00509">
        <property type="entry name" value="RAS_GTPASE_ACTIV_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50018">
        <property type="entry name" value="RAS_GTPASE_ACTIV_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01159">
        <property type="entry name" value="WW_DOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50020">
        <property type="entry name" value="WW_DOMAIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="4" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="1" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="Calponin-homology (CH)" evidence="7">
        <location>
            <begin position="43"/>
            <end position="158"/>
        </location>
    </feature>
    <feature type="domain" description="WW" evidence="6">
        <location>
            <begin position="673"/>
            <end position="706"/>
        </location>
    </feature>
    <feature type="domain" description="Ras-GAP" evidence="5">
        <location>
            <begin position="972"/>
            <end position="1205"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="4">
        <location>
            <begin position="167"/>
            <end position="194"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50020"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029329.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1596" mass="181439" checksum="2DFBAE73E98F5FF5" modified="2019-05-08" version="1">MNPWGVGVQNYPHPQAVPHAQDERLTADEMDEQRRQNVAYQYLCRLEEAKRWMEACLNEELASPTELEESLRNGVILAKLGHCFAPDIIPLKKIYDREQMRYEATGLHFRHTDNINHWREAMAHLGLPTIFHPETTDIYDKKNMPRVVYCIHALSLYLFKLGLAPQIQDLYGKVNFTEEEINNMKRELEKYGLQMPAFSKIGGILASELSVDEAAVHAAVLAINEAVERGVVEQTMVALHNPSAMLLNLRENLAAVYQEMLFQVKMEKAGNARNRESPESEDIYDRCLTQAEIQGNINKVNVHGALEYVDDALERQDPLALCQALQDPVLALRGVQRENAAWYLDQLSTDREQKELGYVDLLEQEEVQAGVYAANERGDQEHAMRRAVSRINASIRRGEAMETVRELMSPEAQLPKAFPFAAQLYQHELSLLQQHHPQLGQEELYVAVEMLSAVALIDRALEAGDVSGFWSSLVSPALGLSDIEDANAQRYFDDLLKLKRRSGKAGVAFLSWNDIQACVNSANLAVQDENDKILAIRLINEALAQDDPEKTLTALLLPSARFSAVTLPGARRYHDVLTAALKQKAQVTGDDGAALWWEEIQQGVSKANQDTEAARRLALGIAAINQAIKEGKPAQTARVLRNPDVSLRGIVAECAGAYQAQLAALLAAKRQAGNANHNWVRHQLKDGSVYYFSLQNFEGSWERPHDCVLNSTHLSREEIQSTITGVTAAHDRERLWESNAGFVVRLQARMRGYLVRQEFVARRHLLQEQLPAVIKIQVRSAARLPSRSHADGHDDIYIALGLQGINAVIKIQAFVRANKARGDYRMLVHARDPPLSIVRRFVHLLEQSQHDFWEESELLKLREEVVKRIRSNQQLESDLNLMDIKIGLLVKNRITLQEVVSHCKKLTRKNKEQLSNLMAIDKQKGLKSLSKEKRQKLEAYQHLFYLLQTHPVYLAKLIFQMPQNKSTKFMESVVFTLYNYASNPREAYLLLQLFKTALQEEIRSKVDQLRDILTGNPTVIRLVVSFYRNARGQNALRQILGSPVQEVLQDKTLSIRTNPVDIYKAWINQLESQSGQKSKFPYDVSPEQALSHPEVQRRLDISIRNLLAVTDKFLSAITSSVDKIPYGMRYVAKVLKTSLAEKFPAAGEEDICKIVGNLLYYRFMNPAVVAPDGFDIVDISAGVALHPEHRRNLGSIAKILQHAAANKMFEGENGHLRVVNQYLEETHSKFRLFICAACSVPEPEERFNVDEYSEMVAVAKPVIYITVAELINTHKLLLEHQDSISPDHKDPLHELLEDLDELPTVQSLIGTCRLLVSFKPAPPLVSSFNPQNGTKQMLVDVIQSQPGDSLLEILRTAASEDQEAFHHHQMHRRALREAQTPDRLKRNRSLAGNSQLSMEEKKRKVIRNLRRLENLGVVDSANQYQEIINEMAKDIRNQRRYRQHRKGELVKLRQTLCGLNSKTSFYEEQIDYYNQYIRTCLDNLAASSKLNGKSKKQQSLHYTAARLFEKGVLLEIEDLPVSQFRNVIFDIIPCGEAGKFQVKAKFMGIDMEKFQLHYQDLLQLQYEGVAVMKLFDKAKVNVNLLIFLLNKKFFRK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN92</accession>
    <name>A0A452IN92_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">SMB domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IN92"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033341.1">
        <property type="protein sequence ID" value="ENSGAGP00000029360.1"/>
        <property type="gene ID" value="ENSGAGG00000020186.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009986">
        <property type="term" value="C:cell surface"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004551">
        <property type="term" value="F:dinucleotide phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003676">
        <property type="term" value="F:nucleic acid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004528">
        <property type="term" value="F:phosphodiesterase I activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046034">
        <property type="term" value="P:ATP metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030505">
        <property type="term" value="P:inorganic diphosphate transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045599">
        <property type="term" value="P:negative regulation of fat cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009143">
        <property type="term" value="P:nucleoside triphosphate catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030500">
        <property type="term" value="P:regulation of bone mineralization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd16018">
        <property type="entry name" value="Enpp"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00091">
        <property type="entry name" value="NUC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.410.20:FF:000003">
        <property type="entry name" value="Ectonucleotide pyrophosphatase/phosphodiesterase family member 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.410.20:FF:000001">
        <property type="entry name" value="Ectonucleotide pyrophosphatase/phosphodiesterase family member 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.720.10:FF:000145">
        <property type="entry name" value="Uncharacterized protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.410.20">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.720.10">
        <property type="entry name" value="Alkaline Phosphatase, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.570.10">
        <property type="entry name" value="Extracellular Endonuclease, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017850">
        <property type="entry name" value="Alkaline_phosphatase_core_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044929">
        <property type="entry name" value="DNA/RNA_non-sp_Endonuclease_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001604">
        <property type="entry name" value="Endo_G_ENPP1-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020821">
        <property type="entry name" value="ENPP1-3/EXOG-like_nuc-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044925">
        <property type="entry name" value="His-Me_finger_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002591">
        <property type="entry name" value="Phosphodiest/P_Trfase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036024">
        <property type="entry name" value="Somatomedin_B-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001212">
        <property type="entry name" value="Somatomedin_B_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10151">
        <property type="entry name" value="ECTONUCLEOTIDE PYROPHOSPHATASE/PHOSPHODIESTERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10151:SF77">
        <property type="entry name" value="ECTONUCLEOTIDE PYROPHOSPHATASE_PHOSPHODIESTERASE FAMILY MEMBER 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01663">
        <property type="entry name" value="Phosphodiest"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01033">
        <property type="entry name" value="Somatomedin_B"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00892">
        <property type="entry name" value="Endonuclease_NS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00477">
        <property type="entry name" value="NUC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00201">
        <property type="entry name" value="SO"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53649">
        <property type="entry name" value="Alkaline phosphatase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54060">
        <property type="entry name" value="His-Me finger endonucleases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90188">
        <property type="entry name" value="Somatomedin B domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00524">
        <property type="entry name" value="SMB_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50958">
        <property type="entry name" value="SMB_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="4" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="7">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019107429" description="SMB domain-containing protein" evidence="7">
        <location>
            <begin position="24"/>
            <end position="806"/>
        </location>
    </feature>
    <feature type="domain" description="SMB" evidence="8">
        <location>
            <begin position="31"/>
            <end position="71"/>
        </location>
    </feature>
    <feature type="domain" description="SMB" evidence="8">
        <location>
            <begin position="72"/>
            <end position="116"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50958"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029360.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="806" mass="91793" checksum="5D608052F91208B5" modified="2019-05-08" version="1">PDPFKLLVLSACVLTIILGCIFGLKPSCSRDVKTCKGRCFERTFGSCRCDRDCVELGNCCLDFQETCIQPAHIWTCNKFRCGEKRWPENRCSCSDDCVENKDCCVNYYIVCQGAKSWVEEKCEDINEPQCPPGFAKPPVLLFSLDGFRAEYLHTWSGLLPVISKLQKCGTYTPNMRPVYPTKTFPNHYTIVTGLYPESHGIIDNKIYDPHRNVSFTLKNAEKFNPQWYQGQPIWLTAMHQGLKAGTFFWPGSDVRINGTFPDLYRKYNGSIPFEERVIAILHWLQLPKDERPHFYTLYLEEPDSSGHNFGPVSSGVILALQRVDNIVGMLMDGLKQMNLHKCLNIILLSDHGMETASCSKTAYLSTYMENVQDITVIPGPAARLRPQNVPDEYFSYPDQHFKAFMKELLPKRFHYVNNDRIERVHFYLDPQWQLARKPSEIKYCNGGFHGSDNRFMSMQAIFIGFGPGFKFKTQVDPFENIEIYNLICDLLGITPAYNNGTHGSLNHLLKNPVYIPRHPEEESHPSVYPLMKTTSSLDIGCSCDLAGLPIRDFQQQLNHTASEVKKIEKYSLPYGRPRVLQKRSTYYLLYHHQYVSGYSQDLNMTLWSAYTDNRTASVENLSNCLHVDIRIPSSSSKTCSFYKNHHQLSYGFLFPPNTKKSSYEVLLNSNIVPMYHEFKVLWNYFYDVLLPKYTLARNGVNVVSGPVFDYDSNGLYDTPEKVKRLTHDPEVLVPTHYFIVLTSCKNISQTPLQCEGALDALSFIVPHRPDNSESCAVSFGFSPGWWGPEPQGAVPSSGASAFCPGP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGF4</accession>
    <name>A0A452GGF4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Unconventional myosin-XVIIIa</fullName>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">May link Golgi membranes to the cytoskeleton and participate in the tensile force required for vesicle budding from the Golgi. Thereby, may play a role in Golgi membrane trafficking and could indirectly give its flattened shape to the Golgi apparatus. Alternatively, in concert with LURAP1 and CDC42BPA/CDC42BPB, has been involved in modulating lamellar actomyosin retrograde flow that is crucial to cell protrusion and migration. May be involved in the maintenance of the stromal cell architectures required for cell to cell contact. Regulates trafficking, expression, and activation of innate immune receptors on macrophages. Plays a role to suppress inflammatory responsiveness of macrophages via a mechanism that modulates CD14 trafficking. Acts as a receptor of surfactant-associated protein A (SFTPA1/SP-A) and plays an important role in internalization and clearance of SFTPA1-opsonized S.aureus by alveolar macrophages. Strongly enhances natural killer cell cytotoxicity.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Microtubule organizing center</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Golgi apparatus</location>
            <location evidence="2">trans-Golgi network</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="13">Golgi outpost</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 16">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <comment type="caution">
        <text evidence="16">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000000750.1">
        <property type="protein sequence ID" value="ENSGAGP00000000666.1"/>
        <property type="gene ID" value="ENSGAGG00000000355.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005815">
        <property type="term" value="C:microtubule organizing center"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032982">
        <property type="term" value="C:myosin filament"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016460">
        <property type="term" value="C:myosin II complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008094">
        <property type="term" value="F:ATP-dependent activity, acting on DNA"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003774">
        <property type="term" value="F:cytoskeletal motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031032">
        <property type="term" value="P:actomyosin structure organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006259">
        <property type="term" value="P:DNA metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903028">
        <property type="term" value="P:positive regulation of opsonization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043030">
        <property type="term" value="P:regulation of macrophage activation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd01386">
        <property type="entry name" value="MYSc_Myo18"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.1160:FF:000006">
        <property type="entry name" value="Myosin-XVIIIa isoform a"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.820:FF:000004">
        <property type="entry name" value="unconventional myosin-XVIIIa isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000020">
        <property type="entry name" value="unconventional myosin-XVIIIa isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.270.10:FF:000002">
        <property type="entry name" value="unconventional myosin-XVIIIa isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.720:FF:000007">
        <property type="entry name" value="unconventional myosin-XVIIIa isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.530:FF:000011">
        <property type="entry name" value="unconventional myosin-XVIIIa isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.340">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.1590">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.270.10">
        <property type="entry name" value="Myosin, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.1160">
        <property type="entry name" value="Vasodilator-stimulated phosphoprotein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000048">
        <property type="entry name" value="IQ_motif_EF-hand-BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002928">
        <property type="entry name" value="Myosin_tail"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036064">
        <property type="entry name" value="MYSc_Myo18"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020587">
        <property type="entry name" value="RecA_monomer-monomer_interface"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057772">
        <property type="entry name" value="SH3_Myo18a"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615">
        <property type="entry name" value="MYOSIN HEAVY CHAIN, NON-MUSCLE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615:SF13">
        <property type="entry name" value="UNCONVENTIONAL MYOSIN-XVIIIA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01576">
        <property type="entry name" value="Myosin_tail_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24556">
        <property type="entry name" value="SH3_Myosin-XVIIIa"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00015">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90257">
        <property type="entry name" value="Myosin rod fragments"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50163">
        <property type="entry name" value="RECA_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="16" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="7 16" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="9 17" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="12" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="8" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="11 16" id="KW-0505">Motor protein</keyword>
    <keyword evidence="10 16" id="KW-0518">Myosin</keyword>
    <keyword evidence="6 16" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Myosin motor" evidence="20">
        <location>
            <begin position="268"/>
            <end position="1044"/>
        </location>
    </feature>
    <feature type="domain" description="RecA family profile 2" evidence="19">
        <location>
            <begin position="404"/>
            <end position="470"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1"/>
            <end position="180"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="583"/>
            <end position="602"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1711"/>
            <end position="1761"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1821"/>
            <end position="1986"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1999"/>
            <end position="2052"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="2106"/>
            <end position="2137"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="2163"/>
            <end position="2330"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="17">
        <location>
            <begin position="1108"/>
            <end position="1212"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="17">
        <location>
            <begin position="1248"/>
            <end position="1537"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="18">
        <location>
            <begin position="26"/>
            <end position="39"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="48"/>
            <end position="59"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="89"/>
            <end position="100"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="18">
        <location>
            <begin position="111"/>
            <end position="120"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="121"/>
            <end position="145"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="1846"/>
            <end position="1855"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="1874"/>
            <end position="1889"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="1913"/>
            <end position="1922"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="1944"/>
            <end position="1954"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="1968"/>
            <end position="1977"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="18">
        <location>
            <begin position="1999"/>
            <end position="2011"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="2013"/>
            <end position="2022"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="2024"/>
            <end position="2043"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="2127"/>
            <end position="2136"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="2184"/>
            <end position="2203"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="18">
        <location>
            <begin position="2216"/>
            <end position="2225"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="18">
        <location>
            <begin position="2237"/>
            <end position="2255"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="18">
        <location>
            <begin position="2257"/>
            <end position="2290"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="2293"/>
            <end position="2316"/>
        </location>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <begin position="361"/>
            <end position="368"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004267"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004601"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00008314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00024182"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00058021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00073449"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000666.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2330" mass="262178" checksum="7B6BD6A79AA85CC8" modified="2019-05-08" version="1">MNGLQGSRAGSLAQPEPLGEESQGLSRKRVVRVVRKVVRKVLPGEDSGCTKEQGRDGKSPEPASPAKKEEKAPPRVMSSPPLPASSPKPEPKETGPKDEISVGLRSLMSRGKTKQHRPRLRLAEKQEEEKPPELKTPPVRKEPEKPGSPSPAQEEGPVATPGPRAKQEQTVQPSGAKVEPVQVAPQNAAALGQAKAKTEEQITAEEAWYETDKVWLVHKDGFSLASQLRADETSLLPEGKVKVKLDHDGAVLEVDEDDVEKANPPSCDRVEDLANLVYLNESSVLHTVRQRYGGNLIHTYAGPTMLVVNPLSSPSMYSEKVMHMFKGCRREDTSPHIYAVAQAAYRDMLMSRQDQSIVLLGSSGSGKTTCCQHLIQYLATIAGSTGKVFSVEKWQALYTILEAFGNSSTSMNGNATRFSQIVSLDFDQAGQVASASIQTMLLEKLRVTKRPTNEATFNVFYYLLAGTDNALRTELHFNHLAETSVFGIVPLSKPEEKQKAAQQFSKLQAAMKVLGISSDEQKAFWLVLGSIYHLGAAGATKDTDKAGRKQFARHEWAQKAAYLLGCSLEELSSSVFKHQPKGSLQRSTSFRQGPEEAGLGDGSGSKLTALECLEGLASGLYSELFTLLISLLNRALKSSQHSLCSMMLVDTPGFQNPELANQSRGASFEELCHNYAQERLQMLFHERTFTQELERYKEESIELALDDIEPSTSGSVATVDQASHQALVRSLARTDEARGLLWLLEEEALQPAGNEDTLLERLFAYYGPQEGDKKGHSPLLRSDKPRHFFLGHSYGTNWVEYDATGWLQYVRQNPASQNAVALLQDSQKKIISSLFMGRAGTAMVLSGSIAGLEGGSQLALRRATSMRKTFTTGVAAVKKKSLCIQIKLQVDALVDTIKKSKIHFVHCLLPKAEGWSGDPKGLAARRLSSTELDLHGEHCEAGLMQLDVPLLRAQVRGSRLLDALRMYRQGYPDHMVFSEFRRRFDVLAPHLTKKHGRNYIVTDEKRAVEELLESLDLEKSSSHMGLSRVFFRAGTLAKLEEQRDKQTSKNIAHFQAACRGFLARQQFKKRKIQELAIRCVQKNIQKNRGVKDWPWWKLFTTVRPLIEVQLTEDQIRGKDEEIQQLKGRLEKVERERNELRLGSDRLESRVAELTAELTDERHTGESASQLLDAETAERLRAEKEMKDLQIKYEALKKQMDSMEMEVMEARLIRAAELNGDLDDDDSGGEWRLKYERAIREVDFTKKRMQQEFEDKLELEQQSKRQQERRLADLHADNEECQRALQQLKKKCQRLTAELQDTKLHLEGQQGRNHELEKKQRRFDSELAQAHEEAQREKLLREKLSREKDMLIAEVFSLKQQLEDKDSDIMSFTQKVEALEAELQDVSSQESKDEASLAKVKKQLRDLEAKAKDQEEELDEQAGTIQLLEQAKLRLEMEMERLRQTHSKEVESRDEEVEEIRQSCQKKLKQMEVQLEEEYEDKQKVLREKRELESKLAAASDQVSQRDLETEKRLRKDLKRTKALLADAQIMLDHLKNNAPSKREIAQLRNQLEESEFTCAAAVKARKSMEVEIEDLHLQIDDISKAKAALEEQLSRLQREKNEVQSRLEEDQEDMNELMKKHKASVAQASRDLAQINDLQSQLEDANKEKQELQEKLQGLQSQLEFLEQSMVDKSLVSRQEAKIRELETRLEFERTQVKRLENLASRLKENMEKLTEERDQRAAAENREKEQNKRLQRQLRDSKEEMGELAKKETEASRKKHELEMDLESLEAANQSLQSDLKLAFKRIGDLQAAIEDEMESDDNEDLINSLQDVVTKYQKRKNKLEGDSDMDSELEDRVDGVKSWLSRNKGSSKAVSDDGSWKSSRPALNSAPKEGKEGREPKEVEERPVSVTSSLSYRKRLNLKDSIGGQGDSETLFSTLSERPASPERSFRKAKRSSAASSQEETGSVSSWRKSLGSEDPDDRGSVISQAFSEASSRARKGLEKRWSVTTADFDQASALSAPASRAASSRHGLEADEADARSTLSFTLSSPGSLRRSTSRLSEPLAGVSGALSPQLSRRAELGLETSTLGSRVDSRLCRSRSRLEELDDVSSVALSDTRSLYSQHSLGRSFSVPPRPRTSASDKMQLEDSDIRPVSHRSYLDPDLEAAISEVLNYKPIKFKRSSLDPDPDLDSEEDKRSVRSARSATRPESSERSFSNLRRSASALDCSRPRSSRGPRSSSSSSEEDSSEEERKKSARKHAKKKAKKKSKKKQLPLDSDSSSDSSSSSYHSSSSVKKGPKKSPASEGEGAAEGRDAEKSSKRLKKEEKQRKKQVDSLMMKYLYRPESD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGG5</accession>
    <name>A0A452GGG5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Protein zer-1 homolog-like C-terminal domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the zyg-11 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GGG5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000737.1">
        <property type="protein sequence ID" value="ENSGAGP00000000654.1"/>
        <property type="gene ID" value="ENSGAGG00000000551.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031462">
        <property type="term" value="C:Cul2-RING ubiquitin ligase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:001025">
        <property type="entry name" value="Protein zyg-11 homolog A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.10.10:FF:000086">
        <property type="entry name" value="protein zyg-11 homolog B isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.10.10">
        <property type="entry name" value="Leucine-rich Repeat Variant"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011989">
        <property type="entry name" value="ARM-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000225">
        <property type="entry name" value="Armadillo"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055142">
        <property type="entry name" value="ZER1-like_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051341">
        <property type="entry name" value="Zyg-11_UBL_adapter"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12904">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12904:SF22">
        <property type="entry name" value="ZYG-11 FAMILY MEMBER B, CELL CYCLE REGULATOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22964">
        <property type="entry name" value="ZER1-like_2nd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00185">
        <property type="entry name" value="ARM"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52047">
        <property type="entry name" value="RNI-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0433">Leucine-rich repeat</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="Protein zer-1 homolog-like C-terminal" evidence="5">
        <location>
            <begin position="360"/>
            <end position="714"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00009420"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022614"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="Pfam" id="PF22964"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000654.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="735" mass="83105" checksum="F28D503370D97AAE" modified="2019-05-08" version="1">MEEASPYTLLDICLNFLTADLEKFCTERQDGTLCLQEPGIFPQEVADRLLQTMAFHGLLNDGTVGIFRGNQMRLKRACIRKAKISAVAFRKAFCHHKLVELDATGVNADITITDIISGLGSNKWIKQNLQCLVLNSLTLSLEDPYERCFSQLSGLRALSITNVLFYNEDLADVASLPRLESLDISNTSVTDITALLTCKEQLKSLTMHHLKCLKMTTTQILDVLRELKYLNHLDISDDKQFTSDIALRLLEQKDILPNLVSLDISGRKHVTDEAVEAFIQQRPTMQFVGLLATDAGYSEFLTGEGNLKVSGEANETQISEALKRYSERAFFVREALFHLFSLTHIMEKTKPEILKLVVIGMRNHPLNLPVQLAASACVFNLTKQDLAAGMPVRLLADVTHLLLKAMEHFPNHQQLQKNCLLSLCSDRILQDVPFNRFEAAKLVMQWLCNHEDQNMQRMAVAIISILAAKLSTEQTAQLGAELFIVRQLLQIVKQKTNQNLVDTTLKFTLSALWNLTDESPTTCRHFIENQGLELFMRVLESFPSESSIQQKVLGLLNNIAEVKELHSELMWKDFIDHISKLLHSVEVEVSYFAAGIIAHLISRGEQAWTLSHAQRTSLLEELHSAILNWPTPECEMVAYRSFNPFFPLLGCFMTPGVQLWAVWAMQHVCSKNPARYCNMLIEEGGLQHLYNIKENGQTDPDVQRIAIAILDSLEKHIVRHGRPPPCKKQPQTKRN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGG6</accession>
    <name>A0A452GGG6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">TP53-binding protein 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Chromosome</location>
            <location evidence="2">Centromere</location>
            <location evidence="2">Kinetochore</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000000675.1">
        <property type="protein sequence ID" value="ENSGAGP00000000601.1"/>
        <property type="gene ID" value="ENSGAGG00000000496.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000776">
        <property type="term" value="C:kinetochore"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016604">
        <property type="term" value="C:nuclear body"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035861">
        <property type="term" value="C:site of double-strand break"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042393">
        <property type="term" value="F:histone binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140005">
        <property type="term" value="F:histone H4K20me2 reader activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000077">
        <property type="term" value="P:DNA damage checkpoint signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006303">
        <property type="term" value="P:double-strand break repair via nonhomologous end joining"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:2000042">
        <property type="term" value="P:negative regulation of double-strand break repair via homologous recombination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045830">
        <property type="term" value="P:positive regulation of isotype switching"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045944">
        <property type="term" value="P:positive regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd17745">
        <property type="entry name" value="BRCT_p53bp1_rpt1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd17724">
        <property type="entry name" value="BRCT_p53bp1_rpt2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd20383">
        <property type="entry name" value="Tudor_53BP1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.140:FF:000021">
        <property type="entry name" value="Tumor suppressor p53-binding protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.30:FF:000019">
        <property type="entry name" value="Tumor suppressor p53-binding protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10190:FF:000003">
        <property type="entry name" value="Tumor suppressor p53-binding protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10190:FF:000005">
        <property type="entry name" value="Tumor suppressor p53-binding protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.10190">
        <property type="entry name" value="BRCT domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015125">
        <property type="entry name" value="53-BP1_Tudor"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001357">
        <property type="entry name" value="BRCT_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036420">
        <property type="entry name" value="BRCT_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047249">
        <property type="entry name" value="BRCT_p53bp1-like_rpt1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047250">
        <property type="entry name" value="BRCT_p53bp1-like_rpt2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014722">
        <property type="entry name" value="Rib_uL2_dom2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047252">
        <property type="entry name" value="TP53BP1-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15321:SF3">
        <property type="entry name" value="TP53-BINDING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15321">
        <property type="entry name" value="TUMOR SUPPRESSOR P53-BINDING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09038">
        <property type="entry name" value="53-BP1_Tudor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18428">
        <property type="entry name" value="BRCT_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00292">
        <property type="entry name" value="BRCT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52113">
        <property type="entry name" value="BRCT domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF63748">
        <property type="entry name" value="Tudor/PWWP/MBT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50172">
        <property type="entry name" value="BRCT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="13" id="KW-0010">Activator</keyword>
    <keyword evidence="17" id="KW-0137">Centromere</keyword>
    <keyword evidence="3" id="KW-0158">Chromosome</keyword>
    <keyword evidence="8" id="KW-0227">DNA damage</keyword>
    <keyword evidence="15" id="KW-0234">DNA repair</keyword>
    <keyword evidence="12" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="5" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="9" id="KW-0995">Kinetochore</keyword>
    <keyword evidence="4" id="KW-0488">Methylation</keyword>
    <keyword evidence="16" id="KW-0539">Nucleus</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="14" id="KW-0804">Transcription</keyword>
    <keyword evidence="11" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="10" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="BRCT" evidence="20">
        <location>
            <begin position="1662"/>
            <end position="1730"/>
        </location>
    </feature>
    <feature type="domain" description="BRCT" evidence="20">
        <location>
            <begin position="1746"/>
            <end position="1846"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="62"/>
            <end position="112"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="149"/>
            <end position="300"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="524"/>
            <end position="553"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="627"/>
            <end position="719"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="914"/>
            <end position="1023"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1051"/>
            <end position="1105"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1148"/>
            <end position="1218"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1232"/>
            <end position="1337"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1515"/>
            <end position="1602"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="72"/>
            <end position="82"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="86"/>
            <end position="100"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="154"/>
            <end position="167"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="191"/>
            <end position="200"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="203"/>
            <end position="234"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="532"/>
            <end position="547"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="643"/>
            <end position="653"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="686"/>
            <end position="696"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="971"/>
            <end position="1013"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="19">
        <location>
            <begin position="1161"/>
            <end position="1206"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="19">
        <location>
            <begin position="1522"/>
            <end position="1534"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="1589"/>
            <end position="1602"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004629"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022454"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022838"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023328"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00073180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50172"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000601.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1854" mass="198500" checksum="256AF5C7D22CEC2C" modified="2019-05-08" version="1">MDPSQSQLDLGFSQQDTPCLIVEDSQPESAVTEEDTAHGCLSLLALPLPNCQSPVLEIMTSPQGSKLAAGGGEDRGLEEKPLTEPMDSSTHLDTTGSLSQVIDRLPQPTKTNRVPGMVAAAAEEAGEDASHCATPLLCAEDSGTSQLGFGALELSQSQDVEGDSTLSEEGRGHQLQPADTVPTCCNPMKNDPNEDLHIVRSEAVSTGESYTDVQCQQEKSGSYSSGLITPSSGAFGSHPLISGSPTEQEEAKDEQMNISAPAEGGDLLQEKQRGDEPVEVGNPPVPPGPHVLPQVSTPVSQNAPAFVPGSFPIPSQPEFSHDIFISTQSLEEKPSGEEKGGALLGSCLPADVSETTDALRKISTGDPELSCKLVLSASECSQPTRTEGNTSSLKTNQGSENIQVEMDSTLQASELKPCSTESYSMQLRLSEDSQALLSENCEKPKREAVWATGEPTICPVAEGVAAEGSVAQMACIDHSCDSGSQTAATLSVQSEISSHTLGQEALTEIKEKLSAEASEFEAVPETLCEEQGGERQAEEKQLDEKGSSDPLTLSQGLILLRETKEHEGQEVMEVESSEAFGKNSEKLSDLRHGLVQMDLETRKKETPKDSNVVAEVKSMKDLSLKAAPEDVAKLSEVPPQPSVGESLQQQDDSWPQVKVDVAPGLASCAKMCPGSQGNKSQAEVDLELKQVQKDRQPPPAPLERGDEVPATPQESQQPHLMTLEKEVDMPEKPFKTSNLEQEEERAAEQSPCPSTGTLFHFALPKEGDVIQPLTSVTPSLIGQLKMGPRRHSTPIVDGSCPDRTIVTSDVTAEGSIGTIDVTVESAMASADVLEECEKGDSGAAPEADGKLCLRMNLATPVSEESEGSLPFSLEKPAASERKNGSSAVAGAVASSQKTLSVFSLVCEVQREDEARGHDLPASPFRGNLFSFPSTQEEEEEQEQQHDNPEAWQQHQKHIPCGEIKVSQPAEQDSKHQPSHAKEGEAMEADIEKEGKSAQEKLSKALAAHERDKGQQLWAGTSNKEVQTTGGSVLAPAAVSAATQTMARVSSQVEVGTSMAGQRPGQQDAKVQTEESGEKLANAPGDDTESLHSQGEEEFNLPQPPRGRIQRRHMRTIREVRTVVTRIITDVYYVDGAEVERKVIEETEEPIVDCHESETDISPSRTTGGSSLTSGDLGDISSLSSKVSSLHRTSSGASSGLSATHGGSSSGRGAGSFKGKVSGMEAGEFALPSGRGVLGKLSPRRGAGQPGSPLRAGQTGATACEEDEDAALGSRQGGKAPLTPRGRGRRGRPPSRATGTRDAAGSGLLGVEDISTTVSPEEKPFVRIVGHPPDGGDRFDTAGPCTLRRSDSPEIPLQVVAGPSDSADLSSGSSFVGLRVVAKWSSNAYFYSGTITQDVGAGKYKLLFDDGYECDVLGKDILLCDPIPLETEVTALSEDEYFSAGVVKGHRKESGELYYCIEKEGQRKWYKRMAVILSLEQGNKLREQFGLSPYEPITPLTKAADISLDNLVEGKRKRRSNFGSPGTSSSSTTPTRKGLESPRVAPGLLSGKRKLIASEDERSPAKRGRKSAAMKPGAVRAGEFVSPCESGDHTGEPSTLEEHHGPLPHNKTLFLGYAFLLTMATSSDKLAIHQKPSDGPAGSSEEEEEFSIPYNKEYTALQLRTGAGYILEDFNETQCNAAYQCLLIADQHCRTRKYFLCLASGIPCVSHVWVRDSCHANQLQNYRNYLLPAGYSLQEQRLLEWHPRENPFGNLKVLLVSNQQQNFLELWSEILMTGGAASVKQHYSNAQNKDIALGVFDVVVSDSSCPAAVLKCAEALRLPVVSQEWVIQSLIAGERVGYNKHPKYKHDYGPC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGJ0</accession>
    <name>A0A452GGJ0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Clathrin light chain</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="6">Clathrin is the major protein of the polyhedral coat of coated pits and vesicles.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 6">Cytoplasmic vesicle membrane</location>
            <topology evidence="1 6">Peripheral membrane protein</topology>
            <orientation evidence="1 6">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="6">Membrane</location>
            <location evidence="6">Coated pit</location>
            <topology evidence="6">Peripheral membrane protein</topology>
            <orientation evidence="6">Cytoplasmic side</orientation>
        </subcellularLocation>
        <text evidence="6">Cytoplasmic face of coated pits and vesicles.</text>
    </comment>
    <comment type="similarity">
        <text evidence="2 6">Belongs to the clathrin light chain family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GGJ0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000761.1">
        <property type="protein sequence ID" value="ENSGAGP00000000677.1"/>
        <property type="gene ID" value="ENSGAGG00000000556.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030132">
        <property type="term" value="C:clathrin coat of coated pit"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030130">
        <property type="term" value="C:clathrin coat of trans-Golgi network vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0099631">
        <property type="term" value="C:postsynaptic endocytic zone cytoplasmic component"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030672">
        <property type="term" value="C:synaptic vesicle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032050">
        <property type="term" value="F:clathrin heavy chain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005198">
        <property type="term" value="F:structural molecule activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072583">
        <property type="term" value="P:clathrin-dependent endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006886">
        <property type="term" value="P:intracellular protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000996">
        <property type="entry name" value="Clathrin_L-chain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10639">
        <property type="entry name" value="CLATHRIN LIGHT CHAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10639:SF28">
        <property type="entry name" value="CLATHRIN LIGHT CHAIN B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01086">
        <property type="entry name" value="Clathrin_lg_ch"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00224">
        <property type="entry name" value="CLATHRIN_LIGHT_CHN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00581">
        <property type="entry name" value="CLATHRIN_LIGHT_CHN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 6" id="KW-0168">Coated pit</keyword>
    <keyword evidence="7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5 6" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="3 6" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="37"/>
            <end position="58"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="100"/>
            <end position="142"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="1"/>
            <end position="14"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005263"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023176"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="RuleBase" id="RU363137"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000677.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="204" mass="22789" checksum="C6B9100C5708DF5E" modified="2019-05-08" version="1">MADDFGFFSSSESGAPDEDPAAAFLAQQESEIAGIENDEGFGAADGGPGQAPDLGADFEDVGATVNGEVFQESNGPTDGYAAIARADRLTQEPESIRKWREEQKKRLQELDAASKVMEQEWREKAKKDLEEWNVRQSEQMEKNRVNNRASEEAFLKESKEENPGTAWEKVAQLCDFNPKSSKQWKDVSRMRLVLISLKQTPLSR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGJ2</accession>
    <name>A0A452GGJ2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Alpha-1,3-glucosyltransferase</fullName>
            <ecNumber evidence="12">2.4.1.-</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">Dolichyl pyrophosphate Man9GlcNAc2 alpha-1,3-glucosyltransferase that operates in the biosynthetic pathway of dolichol-linked oligosaccharides, the glycan precursors employed in protein asparagine (N)-glycosylation. The assembly of dolichol-linked oligosaccharides begins on the cytosolic side of the endoplasmic reticulum membrane and finishes in its lumen. The sequential addition of sugars to dolichol pyrophosphate produces dolichol-linked oligosaccharides containing fourteen sugars, including two GlcNAcs, nine mannoses and three glucoses. Once assembled, the oligosaccharide is transferred from the lipid to nascent proteins by oligosaccharyltransferases. In the lumen of the endoplasmic reticulum, adds the first glucose residue from dolichyl phosphate glucose (Dol-P-Glc) onto the lipid-linked oligosaccharide intermediate Man(9)GlcNAc(2)-PP-Dol to produce Glc(1)Man(9)GlcNAc(2)-PP-Dol. Glc(1)Man(9)GlcNAc(2)-PP-Dol is a substrate for ALG8, the following enzyme in the biosynthetic pathway.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>an alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;6)]-alpha-D-Man-(1-&gt;6)]-beta-D-Man-(1-&gt;4)-beta-D-GlcNAc-(1-&gt;4)-alpha-D-GlcNAc-diphospho-di-trans,poly-cis-dolichol + a di-trans,poly-cis-dolichyl beta-D-glucosyl phosphate = an alpha-D-Glc-(1-&gt;3)-alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;3)-[alpha-D-Man-(1-&gt;2)-alpha-D-Man-(1-&gt;6)]-alpha-D-Man-(1-&gt;6)]-beta-D-Man-(1-&gt;4)-beta-D-GlcNAc-(1-&gt;4)-alpha-D-GlcNAc-diphospho-di-trans,poly-cis-dolichol + a di-trans,poly-cis-dolichyl phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:30635"/>
            <dbReference type="Rhea" id="RHEA-COMP:19498"/>
            <dbReference type="Rhea" id="RHEA-COMP:19502"/>
            <dbReference type="Rhea" id="RHEA-COMP:19520"/>
            <dbReference type="Rhea" id="RHEA-COMP:19521"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57525"/>
            <dbReference type="ChEBI" id="CHEBI:57683"/>
            <dbReference type="ChEBI" id="CHEBI:132520"/>
            <dbReference type="ChEBI" id="CHEBI:132521"/>
            <dbReference type="EC" id="2.4.1.267"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="11">
            <dbReference type="Rhea" id="RHEA:30636"/>
        </physiologicalReaction>
    </comment>
    <comment type="pathway">
        <text evidence="2 12">Protein modification; protein glycosylation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 12">Endoplasmic reticulum membrane</location>
            <topology evidence="1 12">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 12">Belongs to the ALG6/ALG8 glucosyltransferase family.</text>
    </comment>
    <dbReference type="EC" id="2.4.1.-" evidence="12"/>
    <dbReference type="AlphaFoldDB" id="A0A452GGJ2"/>
    <dbReference type="STRING" id="38772.ENSGAGP00000000649"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000730.1">
        <property type="protein sequence ID" value="ENSGAGP00000000649.1"/>
        <property type="gene ID" value="ENSGAGG00000000549.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000000765.1">
        <property type="protein sequence ID" value="ENSGAGP00000000680.1"/>
        <property type="gene ID" value="ENSGAGG00000000549.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000000775.1">
        <property type="protein sequence ID" value="ENSGAGP00000000689.1"/>
        <property type="gene ID" value="ENSGAGG00000000549.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00378"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042281">
        <property type="term" value="F:dolichyl pyrophosphate Man9GlcNAc2 alpha-1,3-glucosyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004856">
        <property type="entry name" value="Glyco_trans_ALG6/ALG8"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12413">
        <property type="entry name" value="DOLICHYL GLYCOSYLTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12413:SF1">
        <property type="entry name" value="DOLICHYL PYROPHOSPHATE MAN9GLCNAC2 ALPHA-1,3-GLUCOSYLTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03155">
        <property type="entry name" value="Alg6_Alg8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7 12" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="4 12" id="KW-0328">Glycosyltransferase</keyword>
    <keyword evidence="9 12" id="KW-0472">Membrane</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="13" id="KW-0732">Signal</keyword>
    <keyword evidence="5 12" id="KW-0808">Transferase</keyword>
    <keyword evidence="6 12" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 12" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="13">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5044604440" description="Alpha-1,3-glucosyltransferase" evidence="13">
        <location>
            <begin position="23"/>
            <end position="507"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="110"/>
            <end position="132"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="144"/>
            <end position="161"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="173"/>
            <end position="197"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="230"/>
            <end position="249"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="300"/>
            <end position="318"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="387"/>
            <end position="408"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="442"/>
            <end position="460"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="472"/>
            <end position="495"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004477"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004922"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00008715"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022676"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
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    <sequence length="507" mass="57804" checksum="B96F6D17DAA36F0B" modified="2019-05-08" version="1">MEKWSLMTITVLLGLIVRWTVSFGSYSGAGKPPMYGDYEAQRHWQEVTYNLPIKQWYFNTSDNNLQYWGLDYPPLTAYHSLLCAYVAKLINPDWIALHTSRGYESQPHKLFMRATVLIADLLVYIPAVILYCCYLKETSTKKKIASALCILLYPGLILIDYGHFQYNSVSLGFALWGVLFLSYNWDLLGSLAFCLAINYKQMELYHSLPFFCYLLGKCCQKGLTGKGLVLLIKIAFTVVASFAICWLPFCTEVEQILQVFRRLFPIDRGLFEDKVANIWCSLSVLIKIKNVMSPQTQLKLSFAFTFLTLLPACIKLTLQPSLRRFKFALVSCALSFFLFSFQVHEKSILLVSIPVCLMINEIPLMCTWFLLVSTFSLLPLLLKDGLLLAYVVTTLAFLTVCATFFSIFEKTSEEDLQLKSLSLSIRAYIPSFKAFPKIIKSLFIVSLTLMGILTSMSATLTPPQKLPDLFPVSVSVVSCLHFLFFLMYFNAIVLWDSRNGRNQKKIN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
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            <taxon>Vertebrata</taxon>
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            <taxon>Archelosauria</taxon>
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            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
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        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
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        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
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        <scope>IDENTIFICATION</scope>
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        </source>
    </evidence>
    <sequence length="3139" mass="349094" checksum="545EF5377BFC01FE" modified="2019-05-08" version="1">MGCSSQDPMVAQLLSNTDPDNKASLTVSCLGEGDWSNLGHPQFQQLINLKLNPDPVLPEMDGVSEFAEYVSETVDVPTPFDLLEPPTSGGFLKLSKPCCYIFPGGRGDSALFAVNGFNILVDGGSERKSCFWKLVRHLDRIDSILLTHIGADNLPGINGLLQRKIAEQDEEQSQGSTNYSDWMKNLISPELGVVFFNVPEKLKMPESSMKVKRSIEEACLTLQYLNKLGIKSEPLYRVLSNTIEPITLFHKMGVGKLDMYILNPVKDSKEMQFLMQKWAGNSKAKTGIILANGKEGEISVPYLTSITALVVWLPASPTEKIVRVLFPGNAPQNKILEGLEKLKHLDLLRYPVATQKDMSSGLPPPSLKQTKIKQRTDSKESLKSSPKPVATKQARKEEASDDGAKEIKPEIVKEARTEKKVKEHTEKIPEKPAKPEKIKTESSDALKAEKRKLTKDKIGKKHLKEKVSKLDEKKDKEKKEIKKERKDFKKDDGKKEEKKDSKKEAKPELKKMSKPDLKPFTPEVRKTLYKAKVPGRIKTEKTRVKAEKEAPVEQKMSPAQKRPVQQEPGMADMAEQSLVLSSPEDLTKDFEELNYEEKVELQVAQEMTDIEASDAVMRGPETGGKRVPEMTAQGHVDEALILEENIMMKVTPLDSPDEGIITFDRERELPCEEKPVQLEVRAGQAETFEDERAALEEPLEIGEFAEKAERKEEAKLLPEVHKDERGQAPTHTEAAAREAQTPELKEKEEEQKISLDRKQREGEMTCEKYIEGMDTGEEGEKEKREDVIEKAELEETEEQYTEEKDMKTKEFYQLHQDEEKGTKFTAKEKKREFSEEISEKSKLLEKEVTKEEYSLRSAPVPPGATAEHISYIQDETIPGYSETEQTISDEEIHDEQEERIPHLKYDVSAYDISVPDHPGSFEAVHGIQATSITDIAAKGYISQESEILVYPTSIVAAPLAEEEHVSSATSVTECDKLSSFATSVAEDQSVASITAPQTEETGKSSLLLDTVNSIPSSRTEATQGLDYVPSAGTISPTSSLEEDKCFKSPPSEDFQALAEGERKFKAQKKGLQEETEEEEEEEEEDETPNIEMPEKLRGHYGAQLFTQQGHLADVLIESRTEASMLLLSTEETQLSQVDSCSGEAEERCISPDDSTVKMASPTQSGPTSAGHTPFHQSPVEEKLETVEAELFENDRDASVKKGEGQTSGVEKGAESESEPLKKAEAPAANDIPEEELPSVIQEKPGLANKVLLPMESMFPRNEKQELPVGKEHISYHKQKTIEISEKDDSKLYIPSTDIISKPEKQAEFKETEMMQDVSLEFAKYTAKEPVSTKGISPKNAEVPQQSQLESDKSSQIFIKEISPDDSKSLSFGEESLCKEKSSDISIREASPEEEKPLYFTGHTLEKEKSTHIFIKDISPEDTKSISLTEESPGKETSPDIFMKGTSPEDTKSLTFTEKSPAKETFLATSTNEFSPEAVIPLHFMQQSPATSEKSQETAAKDIIKEKPKSPLCPEHSPIKDMLPTEISSAEISPEDTRVFSLTEMSLARGKALEMCSQETFPDMKFLHFTESRRIQDKESSDVCSEDVKSCDFMEETQTKEKSPEMEDIYMKNLNYEKKVWFPEEMEEIRSKESQQKYDKERESTFLDEVQEYEKKSSLTVTEEEKIIHSMAYEGYALETVEKKTSDSEYGYFQEEDAPRQAGYLPKKEDYLCYGEDKPVPQETEAGHFKVDSAEAQEFARWEDKGLSLETNKMEISPTEITSSEKSEVVRATPSSQHVCIEEKQVKPSSAAKGREESLFPSKQDYSYLDHKEKAVLDLYAKSLDRALTVSPLPASSSALEKEVDTSTQELEKHFMTDQSKPVLQAEVLQGATVEISDADALNQETSDSKYMLDIEQNAESKSTEAPDDPRGSKEQSAAAFGLLYSTSPAEYEYVAASQKDSDAASKNSMHSFSGPYYPKSSSDIIGHSEDMGSFSQVEYHSESCTRTSSSPSEKEVCASKGDDHAKQLEQEEREPTPYPDERTFQYADIYDQVVVSGVGLPSLLLKDVDSLGHPDKEKLKPTFVASRTEKVSVSQKQLLYHISAKEEESCLYTSAEKELSSPTSPKDKVDSSFAYTGIQERYSPLAESDKHGMPSDLRFKEMQEGQNGLSSSTAAPKEPYLTLPPSSPGSRECTPPTYTIDAFYQEERTPYIEAAHPDENIISISQEQRSLSPMEPQPKSFYYEKAGRGEESTSDSELEKGAKEKSEKESKLPSPPQAPEKVSEEEVYADISVMEMAQVSELLHQDKHQESHLSTSIASSPCDPVQKEELGVEKTETATVIETGTAGYSTLTDTCRDVWASEMSYSSHISLQQGKDAEKSPKEAHFLVTDYHHAEIASKGYDKDKVIYLKSTAKYVKEEHQEGLALQSKEMDLSSSGFEFSSLEKHEPSIVFRHETSSSGHLGDKSLILKFSDALLSDQGKKDEYLEVSKKDTSVDSSSLSFSPLSPYKEDKLFSGAVEREVASQQRAKEDCPRLSEDASPEATTPVIHTTAESFYSHMLSAYTGAEPDTRSLWDVSPLIPADSVKTYPAETEGFVSAENCGSAFTCDVDDTLPCRIECQSTLMSPLSTEKQKQSCAFSPEQHLSVTETLMAMTSLPDVLTSFPPHQQEAAATANGPTEVSTSPQAFRSVQSAAEVKDEKALPDSDWTVSPSLGGEDMARQSRPSSLRSPEHIFQPFFPEAQRGGRTEDHGDASHEGEPCLGASSDNRQKFSSCEYKQRKGELSPSFINPSPHDFSEDSDLSQDEGHPSVKGRPHHTTGSDVQATMIEETPPTSVSDSGTSQSDSDVPPETEECPSITADAAMDSDEDADFLPVDKAVGTSHHGSTRSSHDPQPVPMIDPHPHPPHPDVCMVDPEMLVNEQNVSRTDKISKKDLKEKSKGVRKPLSKPKSSSPARKLDVKGKRSPTPVKQPLDRSPKSTTAKKERDGGQKQSKPRTQLVQPSHIEEKDELSRSSHSNPGKTLVNGIKTNPVSSSPKSSLSVLPGPPVYVDLAYIPNHCSGKNSDLEFFKRVRASYYVVSGNDPVNGEPSRVVLDALLEGKSQWGENLQVTLIPTHDTEVTREWYQQTHEKQQELNIMVLASSSTVVMQDESFPACKIEF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGK0</accession>
    <name>A0A452GGK0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Chloride intracellular channel protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>chloride(in) = chloride(out)</text>
            <dbReference type="Rhea" id="RHEA:29823"/>
            <dbReference type="ChEBI" id="CHEBI:17996"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="16">Membrane</location>
            <topology evidence="16">Single-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="16">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="domain">
        <text evidence="16">Members of this family may change from a globular, soluble state to a state where the N-terminal domain is inserted into the membrane and functions as chloride channel. A conformation change of the N-terminal domain is thought to expose hydrophobic surfaces that trigger membrane insertion.</text>
    </comment>
    <comment type="similarity">
        <text evidence="2 16">Belongs to the chloride channel CLIC family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GGK0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000787.1">
        <property type="protein sequence ID" value="ENSGAGP00000000698.1"/>
        <property type="gene ID" value="ENSGAGG00000000580.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034707">
        <property type="term" value="C:chloride channel complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005254">
        <property type="term" value="F:chloride channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016491">
        <property type="term" value="F:oxidoreductase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd10297">
        <property type="entry name" value="GST_C_CLIC5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd03061">
        <property type="entry name" value="GST_N_CLIC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1050.10:FF:000001">
        <property type="entry name" value="Chloride intracellular channel 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.30.10:FF:000021">
        <property type="entry name" value="Chloride intracellular channel 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1050.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.30.10">
        <property type="entry name" value="Glutaredoxin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002946">
        <property type="entry name" value="CLIC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042069">
        <property type="entry name" value="CLIC5_C_GST"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR053823">
        <property type="entry name" value="CLIC_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010987">
        <property type="entry name" value="Glutathione-S-Trfase_C-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036282">
        <property type="entry name" value="Glutathione-S-Trfase_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040079">
        <property type="entry name" value="Glutathione_S-Trfase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036249">
        <property type="entry name" value="Thioredoxin-like_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00862">
        <property type="entry name" value="O-ClC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45476:SF4">
        <property type="entry name" value="CHLORIDE INTRACELLULAR CHANNEL PROTEIN 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45476">
        <property type="entry name" value="CHLORIDE INTRACELLULAR CHANNEL PROTEIN 6-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22441">
        <property type="entry name" value="CLIC-like_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01263">
        <property type="entry name" value="INTCLCHANNEL"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDS00019">
        <property type="entry name" value="Glutathione_Transferase_(cytos"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SFLD" id="SFLDG00358">
        <property type="entry name" value="Main_(cytGST)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47616">
        <property type="entry name" value="GST C-terminal domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52833">
        <property type="entry name" value="Thioredoxin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50405">
        <property type="entry name" value="GST_CTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="13 16" id="KW-0868">Chloride</keyword>
    <keyword evidence="12 16" id="KW-0869">Chloride channel</keyword>
    <keyword evidence="5 16" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="14 16" id="KW-0407">Ion channel</keyword>
    <keyword evidence="10 16" id="KW-0406">Ion transport</keyword>
    <keyword evidence="11" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="3 16" id="KW-0813">Transport</keyword>
    <keyword evidence="7 16" id="KW-0851">Voltage-gated channel</keyword>
    <feature type="domain" description="GST C-terminal" evidence="17">
        <location>
            <begin position="78"/>
            <end position="244"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004162"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007655"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022882"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023173"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023214"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00024167"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="RuleBase" id="RU362009"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50405"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000698.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="251" mass="28306" checksum="76A52AF1E989C9B2" modified="2019-05-08" version="1">MTDTVPANEEGKDPDIELFVKAGVDGESIGNCPFSQRLFMILWLKGVVFNVTTVDLKRKPADLHNLAPGTHPPFLTFNGEVKTDINKIEEFLEEMLAPPKYPKLAAVHRESNTAGIDIFSKFSAYIKNTNQQDNAGLERGLTKALKKLDDYLNTPLPEEIDADSTEEEKVSKRKFLDGDELTLADCNLLPKLHVVKIVAKKYRNYEFPTEMTGLWRYLKNAYARDEFTNTCAADKEIEQAYADVAKRLSNS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGK3</accession>
    <name>A0A452GGK3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Dynein axonemal intermediate chain 4</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="11">WD repeat-containing protein 78</fullName>
        </alternativeName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Flagellum axoneme</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="9">Dynein axonemal particle</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GGK3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000778.1">
        <property type="protein sequence ID" value="ENSGAGP00000000692.1"/>
        <property type="gene ID" value="ENSGAGG00000000553.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005858">
        <property type="term" value="C:axonemal dynein complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0120293">
        <property type="term" value="C:dynein axonemal particle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045504">
        <property type="term" value="F:dynein heavy chain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045503">
        <property type="term" value="F:dynein light chain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003341">
        <property type="term" value="P:cilium movement"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:001248">
        <property type="entry name" value="WD repeat domain 78"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050687">
        <property type="entry name" value="Dynein_IC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036322">
        <property type="entry name" value="WD40_repeat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001680">
        <property type="entry name" value="WD40_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12442:SF12">
        <property type="entry name" value="DYNEIN AXONEMAL INTERMEDIATE CHAIN 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12442">
        <property type="entry name" value="DYNEIN INTERMEDIATE CHAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00400">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00320">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50978">
        <property type="entry name" value="WD40 repeat-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50082">
        <property type="entry name" value="WD_REPEATS_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8" id="KW-0966">Cell projection</keyword>
    <keyword evidence="6" id="KW-0969">Cilium</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="5" id="KW-0282">Flagellum</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="3 12" id="KW-0853">WD repeat</keyword>
    <feature type="repeat" description="WD" evidence="12">
        <location>
            <begin position="654"/>
            <end position="696"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="12">
        <location>
            <begin position="697"/>
            <end position="731"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="1"/>
            <end position="43"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="150"/>
            <end position="175"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="302"/>
            <end position="339"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="410"/>
            <end position="434"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="13">
        <location>
            <begin position="150"/>
            <end position="162"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="13">
        <location>
            <begin position="302"/>
            <end position="320"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="13">
        <location>
            <begin position="321"/>
            <end position="338"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="13">
        <location>
            <begin position="416"/>
            <end position="429"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004611"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022574"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022846"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023069"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00024190"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00040002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00041557"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000692.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="799" mass="89430" checksum="BF7E6F7AF92D4968" modified="2019-05-08" version="1">MSSSTQAGKGRRQSVSLTSKMELNTKGTSISGTSSQRRMSYVSGVNPNRVESLSRRSILIASDGKTMDKGSLIGSRHTIQVFDEQGKDVTPRPLYRPEPSASLHRQSKILSTQDFSGATGSDFLSSFSMHQTSGVNASLVGPFSRTYGSSTISKSTTSTTESMADEIVEPGSRRDTAISLSDVQVRRDEIKEQLTKEDIDKRVDIYLTETETIWLLDMPTVMMSVESEDAEKVQERNKDYIELCKNKVGNDRFVERMMQTLNGAPKTKEVQCEKILMEDKGIMATSWDLYDSFNALEPMLTSSVTERSSRPTSVSSSKSNVTKDRERTMSTTSTDRESITSSSIIDLESVILARIHEEEKDHSEAILKSEKFQQDLFFMERILMENVFQPKLAAYRQLPVLLEPILTSDAGKKEAETEEAEEGEEEEEEKKETVVSSSLLLDLSEAPEEILPPNLERLWSYTCEWTNGHNVSSMAWNKLNPWPERIYQCDHGVTALDFSLANPNLLAVGMYNGTIAIYNVQTHNTTALLDSSESFDKHIGPVWQLKWIEQDRGITGDDKGEILISISADGRITKWLIRKGLGCTDLMKLKRTASERKKLTGEKEKKSEALISRQAPGMCFDFHPRNTNIYLAGTEEGHIHKCSCSYNEQFLETYRAHKGPVYKIAWNPFSADMFLSCSADWSIILWRQDSVRPILTFSSTTNVVHDIMWSPKSAFVFAAVNEGRVEIWDLSVSTLDPLIVTFANPGVKFTTVLFARNTDCILIGDNNGIVSVYELRNMTASDSNKVDALHDIIGSALAS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGK8</accession>
    <name>A0A452GGK8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Serine/threonine-protein kinase</fullName>
            <ecNumber evidence="20">2.7.11.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="26 27">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="27" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="26" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="26" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="20">Proto-oncogene with serine/threonine kinase activity involved in cell survival and cell proliferation.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subunit">
        <text evidence="17">Interacts with RP9. Interacts with HSP90AA1, this interaction stabilizes PIM1 protein levels. Interacts (ubiquitinated form) with HSP70 and promotes its proteasomal degradation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4 20">Belongs to the protein kinase superfamily. CAMK Ser/Thr protein kinase family. PIM subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="20"/>
    <dbReference type="AlphaFoldDB" id="A0A452GGK8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000785.1">
        <property type="protein sequence ID" value="ENSGAGP00000000696.1"/>
        <property type="gene ID" value="ENSGAGG00000000395.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0106310">
        <property type="term" value="F:protein serine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006915">
        <property type="term" value="P:apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043066">
        <property type="term" value="P:negative regulation of apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000209">
        <property type="entry name" value="Serine/threonine-protein kinase pim-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000232">
        <property type="entry name" value="Serine/threonine-protein kinase pim-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017348">
        <property type="entry name" value="PIM1/2/3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051138">
        <property type="entry name" value="PIM_Ser/Thr_kinase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22984">
        <property type="entry name" value="SERINE/THREONINE-PROTEIN KINASE PIM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22984:SF29">
        <property type="entry name" value="SERINE_THREONINE-PROTEIN KINASE PIM-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF037993">
        <property type="entry name" value="STPK_Pim-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0053">Apoptosis</keyword>
    <keyword evidence="12 20 22" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="16" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11 20" id="KW-0418">Kinase</keyword>
    <keyword evidence="13" id="KW-0460">Magnesium</keyword>
    <keyword evidence="10 20 23" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="15" id="KW-0539">Nucleus</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="27" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6 20 24" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="8 20" id="KW-0808">Transferase</keyword>
    <keyword evidence="14" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="Protein kinase" evidence="25">
        <location>
            <begin position="38"/>
            <end position="290"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="21">
        <location>
            <position position="167"/>
        </location>
    </feature>
    <feature type="binding site" evidence="22">
        <location>
            <begin position="44"/>
            <end position="52"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="22 23">
        <location>
            <position position="67"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="22">
        <location>
            <position position="121"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="22">
        <location>
            <position position="128"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00005505"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022703"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00047040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00047899"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00048679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="PIRNR" id="PIRNR037993"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="PIRSR" id="PIRSR037993-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="PIRSR" id="PIRSR037993-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="RuleBase" id="RU000304"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000696.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="313" mass="35258" checksum="26409634C92D9B0F" modified="2019-05-08" version="1">MLLSKINSLAHLCSGSGGELHPAKLQPGKEKEPLEKLYQVGPLLGSGGFGSVYSGTRLSDGAPVAIKHVARDRISDWGELPGGSRVPMEILLLKKVGSGFRGVIRLLDWFERPDSFVLVLERPDPVQDLFDFITERGALPEELARSFFRQVLEAVRHCHSCGVLHRDIKDENILIDLNSGELKLIDFGSGALLKDTVYTDFDGTRVYSPPEWIRYHRYHGRSAAVWSLGVLLYDMVCGDIPFEHDEEIIRGQVYFRQRVSPECQHLIKWCLALRPSDRPSFEDLLNHSWMQDVLLPQKTANFKTPLVELGSIL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGL0</accession>
    <name>A0A452GGL0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">Bromodomain adjacent to zinc finger domain 2A</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the WAL family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000000799.1">
        <property type="protein sequence ID" value="ENSGAGP00000000709.1"/>
        <property type="gene ID" value="ENSGAGG00000000428.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033553">
        <property type="term" value="C:rDNA heterochromatin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd05503">
        <property type="entry name" value="Bromo_BAZ2A_B_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd01397">
        <property type="entry name" value="HAT_MBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd15629">
        <property type="entry name" value="PHD_BAZ2A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.40.10:FF:000199">
        <property type="entry name" value="Bromodomain adjacent to zinc finger domain 2B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.890.10:FF:000002">
        <property type="entry name" value="Bromodomain adjacent to zinc finger domain protein 2B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.920.10">
        <property type="entry name" value="Bromodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.890.10">
        <property type="entry name" value="Methyl-cpg-binding Protein 2, Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017956">
        <property type="entry name" value="AT_hook_DNA-bd_motif"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037374">
        <property type="entry name" value="BAZ2A/B_Bromo"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001487">
        <property type="entry name" value="Bromodomain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036427">
        <property type="entry name" value="Bromodomain-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018359">
        <property type="entry name" value="Bromodomain_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018501">
        <property type="entry name" value="DDT_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016177">
        <property type="entry name" value="DNA-bd_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001739">
        <property type="entry name" value="Methyl_CpG_DNA-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028940">
        <property type="entry name" value="PHD_BAZ2A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028942">
        <property type="entry name" value="WHIM1_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028941">
        <property type="entry name" value="WHIM2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011011">
        <property type="entry name" value="Znf_FYVE_PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001965">
        <property type="entry name" value="Znf_PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019787">
        <property type="entry name" value="Znf_PHD-finger"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45915:SF5">
        <property type="entry name" value="BROMODOMAIN ADJACENT TO ZINC FINGER DOMAIN PROTEIN 2A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45915">
        <property type="entry name" value="TRANSCRIPTION INTERMEDIARY FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00439">
        <property type="entry name" value="Bromodomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02791">
        <property type="entry name" value="DDT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01429">
        <property type="entry name" value="MBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00628">
        <property type="entry name" value="PHD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15612">
        <property type="entry name" value="WHIM1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15613">
        <property type="entry name" value="WSD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00929">
        <property type="entry name" value="ATHOOK"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00503">
        <property type="entry name" value="BROMODOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00384">
        <property type="entry name" value="AT_hook"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00297">
        <property type="entry name" value="BROMO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00571">
        <property type="entry name" value="DDT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00391">
        <property type="entry name" value="MBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00249">
        <property type="entry name" value="PHD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47370">
        <property type="entry name" value="Bromodomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54171">
        <property type="entry name" value="DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57903">
        <property type="entry name" value="FYVE/PHD zinc finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00633">
        <property type="entry name" value="BROMODOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50014">
        <property type="entry name" value="BROMODOMAIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50827">
        <property type="entry name" value="DDT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50982">
        <property type="entry name" value="MBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50016">
        <property type="entry name" value="ZF_PHD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 12" id="KW-0103">Bromodomain</keyword>
    <keyword evidence="7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="9" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0804">Transcription</keyword>
    <keyword evidence="6" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <keyword evidence="4 13" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="MBD" evidence="18">
        <location>
            <begin position="110"/>
            <end position="181"/>
        </location>
    </feature>
    <feature type="domain" description="DDT" evidence="17">
        <location>
            <begin position="386"/>
            <end position="451"/>
        </location>
    </feature>
    <feature type="domain" description="PHD-type" evidence="16">
        <location>
            <begin position="1149"/>
            <end position="1199"/>
        </location>
    </feature>
    <feature type="domain" description="Bromo" evidence="15">
        <location>
            <begin position="1263"/>
            <end position="1333"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1"/>
            <end position="38"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="215"/>
            <end position="246"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="291"/>
            <end position="334"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="565"/>
            <end position="616"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="670"/>
            <end position="732"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="752"/>
            <end position="892"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="1"/>
            <end position="10"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="22"/>
            <end position="38"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="220"/>
            <end position="234"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="14">
        <location>
            <begin position="235"/>
            <end position="244"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="303"/>
            <end position="334"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="574"/>
            <end position="585"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="597"/>
            <end position="607"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="752"/>
            <end position="767"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="833"/>
            <end position="847"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007444"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023117"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00035"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00146"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50014"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50016"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50827"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50982"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000709.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="21">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1359" mass="152874" checksum="98B83E9CF49B7327" modified="2019-05-08" version="1">MDNSQLVNDKSSLEAAPDISTLECSESPSLNNSSSFSLLPDDSQAPASLFVSPDSPPVLGEAVLQGRGCLWSGTGLRVALGPARAALQSAQLVGTICSSPHPGEVPRRRIATAEEVRFPLQHGWRREVRIKKGSHRWQGETWYYGPCGKRMKQFPEVIKYLSRNMVQSVRREHFSFSPRMPVGDFYEERDTPQGLQWVKLTQEEIPSRIQAITGKRGRPRNAEKETAKAKEVPKVKRGRGRPPKAKMADLLSKTDARLLKKLEAQEVLSDEDRLKMSKIKKKMRRKVRAALGEGSLNLPPPGAKKEKGWPEKAKEKGRPKEKGRPKEKAKAVRKVDKNLLAQRRLEERRRQQIILEEMKKPTEDMCLGDHQPLPPFSRIPGLVLPSGAFSDCLTIVEFLHSYGRVLGFEVPRDVPSLCTLQEGLFGMDDSLGEVQDLLVRLLRAALYDPGLPSYCQSLKILGEKVSEISLNRDTVSEILRCFLMAYGADADLCHGLRTKPFQALPPDRKAAILAFLVNELNGSALIINEIDKTLENMSNYRRNKWIIEGKLRRLKIALAKRTGRPESEITGLDEGPRRRSSRITEENSGLEAEEEESRGRKSRKDEEVTSIPASSVPELERQIEKLAKRQMFFRKKLLHASQTLRAAALGQDRYRRRYWVLPHLGGIFEEEEEEKEAAPEISPVKKEPVELPIPNRLKCTASRDLSRPGRPKPPPLNGLLEDSMPPCQSQHDLSQSAFLSWLSQTQSSLLNSSVLTPDSSPGKGDPSPRAPEAPALEESFPETMEKQGPWFNLLPRTPCHASPTLTTSSEEPPARLALPKGPSPLARVGVALTGTSHPRSLSSQPNGPTSPPLASTPIHASPRVPSACPRGRGGPEKAQEPPGQPKRRGRPPTKFFKQIEQKYLTQLMAQPVPPEMQSGWWWLKDAEELEAVVRALHPRGIREKALHKHLTKHKEYLREVCTRPQATGHPVSQEALARWSVTERAYEMDLALLQWVEELEQRVLMADLQIRGWTCPSPDSARDDLRYCEHKVGALEEITLKNRREGLPPSREATNPLDLAVLRLAALEQNMERRYLKEPLWAPHEVVVEKAMLSSPEALDLGSTEIAYEITPRMRTWRQTLERCRSAAQVSLCIYQLEKSIAWEKSVNKVTCLVCRKGDDDEHLLLCDGCDRGCHLYCHRPRMTEVPDGDWFCSVCISQVRSCGSPAGLRPLHGLSRAVPLTEIFPSPDNTLSPPPLLGARAFGTYPCAMPLPFCRIILMEMESQEDAWPFLEPVNPRLVPGYRKIIKNPMDFATMRTRLLRGEYVSCEEFATDATLVFDNCQTFNEDDSEVGKAGRAMRRFFESRWEEFYQGKHAPNP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGL6</accession>
    <name>A0A452GGL6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Bromodomain adjacent to zinc finger domain 2A</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GGL6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000811.1">
        <property type="protein sequence ID" value="ENSGAGP00000000720.1"/>
        <property type="gene ID" value="ENSGAGG00000000428.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033553">
        <property type="term" value="C:rDNA heterochromatin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd05503">
        <property type="entry name" value="Bromo_BAZ2A_B_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd15629">
        <property type="entry name" value="PHD_BAZ2A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.920.10">
        <property type="entry name" value="Bromodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037374">
        <property type="entry name" value="BAZ2A/B_Bromo"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001487">
        <property type="entry name" value="Bromodomain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036427">
        <property type="entry name" value="Bromodomain-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018359">
        <property type="entry name" value="Bromodomain_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028940">
        <property type="entry name" value="PHD_BAZ2A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028941">
        <property type="entry name" value="WHIM2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011011">
        <property type="entry name" value="Znf_FYVE_PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001965">
        <property type="entry name" value="Znf_PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019787">
        <property type="entry name" value="Znf_PHD-finger"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45915:SF5">
        <property type="entry name" value="BROMODOMAIN ADJACENT TO ZINC FINGER DOMAIN PROTEIN 2A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45915">
        <property type="entry name" value="TRANSCRIPTION INTERMEDIARY FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00439">
        <property type="entry name" value="Bromodomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00628">
        <property type="entry name" value="PHD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15613">
        <property type="entry name" value="WSD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00503">
        <property type="entry name" value="BROMODOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00297">
        <property type="entry name" value="BROMO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00249">
        <property type="entry name" value="PHD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47370">
        <property type="entry name" value="Bromodomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57903">
        <property type="entry name" value="FYVE/PHD zinc finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00633">
        <property type="entry name" value="BROMODOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50014">
        <property type="entry name" value="BROMODOMAIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50016">
        <property type="entry name" value="ZF_PHD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6 9" id="KW-0103">Bromodomain</keyword>
    <keyword evidence="2" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0804">Transcription</keyword>
    <keyword evidence="5" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="4" id="KW-0862">Zinc</keyword>
    <keyword evidence="3 10" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="PHD-type" evidence="13">
        <location>
            <begin position="315"/>
            <end position="365"/>
        </location>
    </feature>
    <feature type="domain" description="Bromo" evidence="12">
        <location>
            <begin position="449"/>
            <end position="519"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1"/>
            <end position="50"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="372"/>
            <end position="434"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="11">
        <location>
            <begin position="1"/>
            <end position="12"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="14"/>
            <end position="24"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023117"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00035"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00146"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50014"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50016"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000720.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="16">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="545" mass="62544" checksum="F2EDD90FC541B208" modified="2025-10-08" version="2">MLPHGLRGRCRPVPRAADQALPGAAPGPGRGGPEKAQEPPGQPKRRGRPPTKFFKQIEQKYLTQLMAQPVPPEMQSGWWWLKDAEELEAVVRALHPRGIREKALHKHLTKHKEYLREVCTRATADPIFQPQATGHPVSQEALARWSVTERAYEMDLALLQWVEELEQRVLMADLQIRGWTCPSPDSARDDLRYCEHKVGALEEITLKNRREGLPPSREATNPLDLAVLRLAALEQNMERRYLKEPLWAPHEVVVEKAMLSSPEALDLGSTEIAYEITPRMRTWRQTLERCRSAAQVSLCIYQLEKSIAWEKSVNKVTCLVCRKGDDDEHLLLCDGCDRGCHLYCHRPRMTEVPDGDWFCSVCISQVREYYEDPSSPRRGKKRKGGRLFGGGFPEEEESPGRRVLPRRRDTLSVPRYSGEGLSPSKRRRLSPRGQSSDLTFCEIILMEMESQEDAWPFLEPVNPRLVPGYRKIIKNPMDFATMRTRLLRGEYVSCEEFATDATLVFDNCQTFNEDDSEVGKAGRAMRRFFESRWEEFYQGKHAPNP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGM5</accession>
    <name>A0A452GGM5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Phosphoglucomutase-1</fullName>
            <ecNumber evidence="6">5.4.2.2</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="16">Glucose phosphomutase 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="17">Catalyzes the reversible isomerization of alpha-D-glucose 1-phosphate to alpha-D-glucose 6-phosphate. The mechanism proceeds via the intermediate compound alpha-D-glucose 1,6-bisphosphate. This enzyme participates in both the breakdown and synthesis of glucose.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>O-phospho-L-seryl-[protein] + alpha-D-glucose 1-phosphate = alpha-D-glucose 1,6-bisphosphate + L-seryl-[protein]</text>
            <dbReference type="Rhea" id="RHEA:68748"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:58392"/>
            <dbReference type="ChEBI" id="CHEBI:58601"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>alpha-D-glucose 1,6-bisphosphate + L-seryl-[protein] = O-phospho-L-seryl-[protein] + alpha-D-glucose 6-phosphate</text>
            <dbReference type="Rhea" id="RHEA:68752"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:58225"/>
            <dbReference type="ChEBI" id="CHEBI:58392"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>alpha-D-glucose 1-phosphate = alpha-D-glucose 6-phosphate</text>
            <dbReference type="Rhea" id="RHEA:23536"/>
            <dbReference type="ChEBI" id="CHEBI:58225"/>
            <dbReference type="ChEBI" id="CHEBI:58601"/>
            <dbReference type="EC" id="5.4.2.2"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="2">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subunit">
        <text evidence="5">Monomer.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4 20">Belongs to the phosphohexose mutase family.</text>
    </comment>
    <dbReference type="EC" id="5.4.2.2" evidence="6"/>
    <dbReference type="AlphaFoldDB" id="A0A452GGM5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000820.1">
        <property type="protein sequence ID" value="ENSGAGP00000000729.1"/>
        <property type="gene ID" value="ENSGAGG00000000620.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000287">
        <property type="term" value="F:magnesium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004614">
        <property type="term" value="F:phosphoglucomutase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006006">
        <property type="term" value="P:glucose metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd03085">
        <property type="entry name" value="PGM1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.310.50:FF:000002">
        <property type="entry name" value="Phosphoglucomutase 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.120.10:FF:000004">
        <property type="entry name" value="Phosphoglucomutase 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.120.10:FF:000005">
        <property type="entry name" value="Phosphoglucomutase 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.120.10:FF:000007">
        <property type="entry name" value="Phosphoglucomutase 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.120.10">
        <property type="entry name" value="Alpha-D-Glucose-1,6-Bisphosphate, subunit A, domain 3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.310.50">
        <property type="entry name" value="Alpha-D-phosphohexomutase, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005844">
        <property type="entry name" value="A-D-PHexomutase_a/b/a-I"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016055">
        <property type="entry name" value="A-D-PHexomutase_a/b/a-I/II/III"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005845">
        <property type="entry name" value="A-D-PHexomutase_a/b/a-II"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005846">
        <property type="entry name" value="A-D-PHexomutase_a/b/a-III"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036900">
        <property type="entry name" value="A-D-PHexomutase_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016066">
        <property type="entry name" value="A-D-PHexomutase_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005841">
        <property type="entry name" value="Alpha-D-phosphohexomutase_SF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045244">
        <property type="entry name" value="PGM"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF005737">
        <property type="entry name" value="PRK07564.1-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22573:SF37">
        <property type="entry name" value="PHOSPHOGLUCOMUTASE-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22573">
        <property type="entry name" value="PHOSPHOHEXOMUTASE FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24947">
        <property type="entry name" value="PGM1_C_vert_fung"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02878">
        <property type="entry name" value="PGM_PMM_I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02879">
        <property type="entry name" value="PGM_PMM_II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02880">
        <property type="entry name" value="PGM_PMM_III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00509">
        <property type="entry name" value="PGMPMM"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55957">
        <property type="entry name" value="Phosphoglucomutase, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53738">
        <property type="entry name" value="Phosphoglucomutase, first 3 domains"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00710">
        <property type="entry name" value="PGM_PMM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12" id="KW-0007">Acetylation</keyword>
    <keyword evidence="14" id="KW-0119">Carbohydrate metabolism</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0313">Glucose metabolism</keyword>
    <keyword evidence="13" id="KW-0413">Isomerase</keyword>
    <keyword evidence="11 20" id="KW-0460">Magnesium</keyword>
    <keyword evidence="10 20" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Alpha-D-phosphohexomutase alpha/beta/alpha" evidence="21">
        <location>
            <begin position="15"/>
            <end position="157"/>
        </location>
    </feature>
    <feature type="domain" description="Alpha-D-phosphohexomutase alpha/beta/alpha" evidence="22">
        <location>
            <begin position="194"/>
            <end position="299"/>
        </location>
    </feature>
    <feature type="domain" description="Alpha-D-phosphohexomutase alpha/beta/alpha" evidence="23">
        <location>
            <begin position="307"/>
            <end position="420"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00010231"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00011245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00012728"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022526"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023235"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023277"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00040178"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00043051"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00045679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00049318"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00049409"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="RuleBase" id="RU004326"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="Pfam" id="PF02878"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="Pfam" id="PF02879"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="Pfam" id="PF02880"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000729.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="562" mass="61619" checksum="F808D8F2D555DD7B" modified="2019-05-08" version="1">MVKIVTVRTQAYADQKPGTSGLRKRVRVFQDNAHYAENFIQSILATVPAAERQEAALVVGGDGRFYMTEAIQLIIRIAAANGIGRLVIGQNGILSTPAVSCIIRKIKAIGGIILTASHNPGGPNGDFGIKFNISNGGPAPEGITDKIFQISKTIEEYAICPDLKVDLGTIGKQQFDLENKFKPFTVEIVDSVEAYANMLRNIFDFSALKELLSGQNHLKIRIDAMHGVMGPYVKKILCEELGAPANSAVNCTPLEDFGGHHPDPNLTYAADLVQSMKTGEYDFGAAFDGDGDRNMILGKHGFFVNPSDSVAVIAANIFSIPYFQHTGVRGFARSMPTSGAIDRVAKATKIALFETPTGWKFFGNLMDANKLSLCGEESFGTGSDHIREKDGLWAVLAWLSIIAVRKQSVEDILKDHWQKYGRNFFTRYDYEEVDADRANKMMKDLETVMFDRSFVGKQLSAGDKVYTVEKADNFEYSDPVDGSISRNQGLRLIFSDGSRIIFRLSGTGSAGATVRLYIDSYEKDAPKIYEDPQVMLAPLIGIALQLSQLHERTGRTGPTVIT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGM8</accession>
    <name>A0A452GGM8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase</fullName>
            <ecNumber evidence="15">2.7.4.24</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Bifunctional inositol kinase that acts in concert with the IP6K kinases IP6K1, IP6K2 and IP6K3 to synthesize the diphosphate group-containing inositol pyrophosphates diphosphoinositol pentakisphosphate, PP-InsP5, and bis-diphosphoinositol tetrakisphosphate, (PP)2-InsP4. PP-InsP5 and (PP)2-InsP4, also respectively called InsP7 and InsP8, regulate a variety of cellular processes, including apoptosis, vesicle trafficking, cytoskeletal dynamics, exocytosis, insulin signaling and neutrophil activation. Phosphorylates inositol hexakisphosphate (InsP6) at position 1 to produce PP-InsP5 which is in turn phosphorylated by IP6Ks to produce (PP)2-InsP4. Alternatively, phosphorylates PP-InsP5 at position 1, produced by IP6Ks from InsP6, to produce (PP)2-InsP4. Activated when cells are exposed to hyperosmotic stress.</text>
    </comment>
    <comment type="function">
        <text evidence="15">Bifunctional inositol kinase that acts in concert with the IP6K kinases to synthesize the diphosphate group-containing inositol pyrophosphates diphosphoinositol pentakisphosphate, PP-InsP5, and bis-diphosphoinositol tetrakisphosphate, (PP)2-InsP4. PP-InsP5 and (PP)2-InsP4, also respectively called InsP7 and InsP8, may regulate a variety of cellular processes, including apoptosis, vesicle trafficking, cytoskeletal dynamics, and exocytosis. Phosphorylates inositol hexakisphosphate (InsP6).</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>1D-myo-inositol hexakisphosphate + ATP = 1-diphospho-1D-myo-inositol 2,3,4,5,6-pentakisphosphate + ADP</text>
            <dbReference type="Rhea" id="RHEA:37459"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:58130"/>
            <dbReference type="ChEBI" id="CHEBI:74946"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.4.24"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="13">
            <dbReference type="Rhea" id="RHEA:37460"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="12">
            <text>5-diphospho-1D-myo-inositol 1,2,3,4,6-pentakisphosphate + ATP + H(+) = 1,5-bis(diphospho)-1D-myo-inositol 2,3,4,6-tetrakisphosphate + ADP</text>
            <dbReference type="Rhea" id="RHEA:10276"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:58628"/>
            <dbReference type="ChEBI" id="CHEBI:77983"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.4.24"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="12">
            <dbReference type="Rhea" id="RHEA:10277"/>
        </physiologicalReaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2 15">Cytoplasm</location>
            <location evidence="2 15">Cytosol</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 15">Belongs to the histidine acid phosphatase family. VIP1 subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.4.24" evidence="15"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000825.1">
        <property type="protein sequence ID" value="ENSGAGP00000000734.1"/>
        <property type="gene ID" value="ENSGAGG00000000614.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033857">
        <property type="term" value="F:5-diphosphoinositol pentakisphosphate 1-kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0052723">
        <property type="term" value="F:inositol hexakisphosphate 1-kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="RHEA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006020">
        <property type="term" value="P:inositol metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032958">
        <property type="term" value="P:inositol phosphate biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07061">
        <property type="entry name" value="HP_HAP_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.470.20:FF:000003">
        <property type="entry name" value="Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.11950:FF:000001">
        <property type="entry name" value="Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.11950:FF:000003">
        <property type="entry name" value="Inositol hexakisphosphate and diphosphoinositol-pentakisphosphate kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.11950">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.470.20">
        <property type="entry name" value="ATP-grasp fold, B domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1240">
        <property type="entry name" value="Phosphoglycerate mutase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033379">
        <property type="entry name" value="Acid_Pase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000560">
        <property type="entry name" value="His_Pase_clade-2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037446">
        <property type="entry name" value="His_Pase_VIP1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029033">
        <property type="entry name" value="His_PPase_superfam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040557">
        <property type="entry name" value="VIP1_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12750">
        <property type="entry name" value="DIPHOSPHOINOSITOL PENTAKISPHOSPHATE KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12750:SF11">
        <property type="entry name" value="INOSITOL HEXAKISPHOSPHATE AND DIPHOSPHOINOSITOL-PENTAKISPHOSPHATE KINASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00328">
        <property type="entry name" value="His_Phos_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18086">
        <property type="entry name" value="PPIP5K2_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53254">
        <property type="entry name" value="Phosphoglycerate mutase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00616">
        <property type="entry name" value="HIS_ACID_PHOSPHAT_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10 15" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="5 15" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9 15" id="KW-0418">Kinase</keyword>
    <keyword evidence="11" id="KW-0472">Membrane</keyword>
    <keyword evidence="8 15" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7 15" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="VIP1 N-terminal" evidence="17">
        <location>
            <begin position="55"/>
            <end position="144"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="910"/>
            <end position="988"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1188"/>
            <end position="1267"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1294"/>
            <end position="1462"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1"/>
            <end position="15"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="16">
        <location>
            <begin position="1202"/>
            <end position="1219"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1232"/>
            <end position="1250"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1424"/>
            <end position="1446"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004514"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005609"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00033696"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00034629"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00037056"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="RuleBase" id="RU365032"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF18086"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000734.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1462" mass="163806" checksum="B82AB6CB1D526726" modified="2019-05-08" version="1">MSSLTTSHEGASSTPRFFVGTRDDETDFLGSNMKTDEIDFFEDDEESESPPERQIVVGICAMTKKSKSKPMTQILERLCKFEYITVVIMGEDVILNEPVENWPPCDCLISFHSKGFPLDKAVAYAKLCNPFLINDLDMQYYIQDRREVYRILQEEGIDLPRYAVLNRDPDKPDECSLVEGEDQVEVNGAVFPKPFVEKPVSAEDHNVYIYYPTSAGGGSQRLFRKIGSRSSVYSPESCVRKTGSYIYEEFMPTDGTDVKVYTVGPDYAHAEARKSPALDGKVERDSEGKEIRYPVMLTAMEKLVARKVCVAFKQTVCGFDLLRANGHSFVCDVNGFSFVKNSMKYYDDCTKILGNIIMRELAPQFQIPWSIPTEAEDIPIVPTTSGTMMELRCVIAVIRHGDRTPKQKMKMEVKHSRFFELFEKYDGYKTGKLKLKKPEQLQEVLDIARLLVVELGTHNDCEIEERKSKLEQLKTVLEMYGHFSGINRKVQLTYLPRGHPKASSEDEDARKEPSPSLLLVLKWGGELTPAGRVQAEELGRAFRCMYPGGQGDYAGFPGCGLLRLHSTYRHDLKIYASDEGRVQMTAAAFAKGLLALEGELTPILVQMVKSANMNGLLDSDSDSLSSCQHRVKARLHEIMQKDAKFCEEDYEKLAPTGSISLVSSMTFIQNPVEICDQVFALIENLTSQICKRLEDPKSADLQLYHSETLELMLQRWSKLERDFRMKNGRYDISKIPDIYDCIKYDVQHNGTLKLEGTAELFKLSKALADVIIPQEYGINKEEKLEIAIGFCLPLIKKIQLDLQRTHEDESVNKLHPLYSRGVLSPGRHVRTRLYFTSESHVHSLLSIFRYGGLLDESKDQQWKRAMDYLSAISELNYMTQIVIMLYEDNNKDPSSEERFHVELHFSPGVKGCEEDGSAPTGFGFRPASAENDDKKPNQGSMEDLSKVKSIDETDQAQERSPQPSEPICIQRRSPLIRNRKTGSMEVLSESSSKTGGYRLFSSYSRQSTEMKQSGLGSQCTGLFSTTVLGGSSSAPNLQDYVRSHGKKFTASLTYKDDVLEHCRVVPLLCRFPSDYAMRQNISLDATLAHHLHQCSYHLRLFRSWLISGQDDLECLYGFEGCSMVPTIYPLETLHNSLSLRQVNEFLTSVCRCYSSLHSRASTALFDSVIASQTPGDPFMPQRILSSSSLPLRQRSDKPPWYSSGPSSTVSSAGPSSPTSMENYARFGFTEKPSISPQISEERLNCQSPEQSEMEPSGEALELDLGGSGSLVENPLGEMPEQGLMRMPGLLEVPEPSKEELSEEVPEQGTMRVLGLSKTRQSGKTSEKDGSVVLEQGGEKPAGGTWELGETGKLEAGEIGLSGKISDSDEEGPAGEMLEPDVVRPVAPGETSELDEEEPVMQTLEPSEKGQPVEILEKEDEELSTDNALSDQQPFSHLYQDQQLSSEKSVEDEQPSSQQELQN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXH1</accession>
    <name>A0A452GXH1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">C2H2-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GXH1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007910.1">
        <property type="protein sequence ID" value="ENSGAGP00000006815.1"/>
        <property type="gene ID" value="ENSGAGG00000005496.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010468">
        <property type="term" value="P:regulation of gene expression"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000233">
        <property type="entry name" value="Putative zinc finger protein 362"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000158">
        <property type="entry name" value="Zinc finger protein 362"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000377">
        <property type="entry name" value="zinc finger protein 362 isoform X4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000216">
        <property type="entry name" value="Zinc finger protein 384 like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050331">
        <property type="entry name" value="Zinc_finger"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16515:SF49">
        <property type="entry name" value="GASTRULA ZINC FINGER PROTEIN XLCGF49.1-LIKE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16515">
        <property type="entry name" value="PR DOMAIN ZINC FINGER PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="6"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="9" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 11" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="275"/>
            <end position="302"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="303"/>
            <end position="330"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="331"/>
            <end position="358"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="359"/>
            <end position="388"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="389"/>
            <end position="416"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="13">
        <location>
            <begin position="419"/>
            <end position="442"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="110"/>
            <end position="135"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="170"/>
            <end position="204"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="231"/>
            <end position="250"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="182"/>
            <end position="202"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006815.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="468" mass="51144" checksum="A9B99393473323FE" modified="2019-05-08" version="1">MSGCWSYSAAPSSPWGCSGNLLAGGACTKPREPIAGDSSACFPPLPAPQCWIRWALTSLCFSAERSWMAEPRFNNPYFWPPPPTMPSQLDNLVLINKIKEQLMAEKIRPPHLPPTSVSSQQPLLVPPSPAESSQSIMSLPKLQQVPGLHPQAVPQPDVALHARPATSTVTGLGLASRATAVSTSESSTGTSTPSTPTSTSQSRLIASSPTLISGITSPPLLDSIKTIQGHSLLGAPKPERGRKKIKAENPSGPPVLVVPYPILASGETAKEGKTYRCKVCPLTFFTKSEMQIHSKSHTEAKPHKCPHCSKSFANASYLAQHLRIHLGVKPYHCSYCEKSFRQLSHLQQHTRIHTGDRPYKCPHPGCEKAFTQLSNLQSHQRQHNKDKPYKCPNCYRAYSDSASLQIHLSAHAIKHAKAYCCSMCGRAYTSETYLMKHMSKHTVVEHLVSQHSPQRTESPGIPVRISLI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXI0</accession>
    <name>A0A452GXI0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="23">Nitric oxide synthase 3</fullName>
            <ecNumber evidence="7">1.14.13.39</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="20">Constitutive NOS</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="26">EC-NOS</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="24">NOS type III</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="25">Nitric oxide synthase, endothelial</fullName>
        </alternativeName>
    </protein>
    <organism evidence="30 31">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="31" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="30" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="30" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="21">
            <text>2 L-arginine + 3 NADPH + 4 O2 + H(+) = 2 L-citrulline + 2 nitric oxide + 3 NADP(+) + 4 H2O</text>
            <dbReference type="Rhea" id="RHEA:19897"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:15379"/>
            <dbReference type="ChEBI" id="CHEBI:16480"/>
            <dbReference type="ChEBI" id="CHEBI:32682"/>
            <dbReference type="ChEBI" id="CHEBI:57743"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="EC" id="1.14.13.39"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="21">
            <dbReference type="Rhea" id="RHEA:19898"/>
        </physiologicalReaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="2">
            <name>(6R)-L-erythro-5,6,7,8-tetrahydrobiopterin</name>
            <dbReference type="ChEBI" id="CHEBI:59560"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="4">
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>FMN</name>
            <dbReference type="ChEBI" id="CHEBI:58210"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="3">
            <name>heme b</name>
            <dbReference type="ChEBI" id="CHEBI:60344"/>
        </cofactor>
    </comment>
    <comment type="subunit">
        <text evidence="22">Homodimer. Interacts with NOSIP and NOSTRIN. Interacts with HSP90AB1. Forms a complex with ASL, ASS1 and SLC7A1; the complex regulates cell-autonomous L-arginine synthesis and citrulline recycling while channeling extracellular L-arginine to nitric oxide synthesis pathway.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="5">Cell membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the NOS family.</text>
    </comment>
    <dbReference type="EC" id="1.14.13.39" evidence="7"/>
    <dbReference type="AlphaFoldDB" id="A0A452GXI0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007912.1">
        <property type="protein sequence ID" value="ENSGAGP00000006817.1"/>
        <property type="gene ID" value="ENSGAGG00000000423.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050660">
        <property type="term" value="F:flavin adenine dinucleotide binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010181">
        <property type="term" value="F:FMN binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0020037">
        <property type="term" value="F:heme binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050661">
        <property type="term" value="F:NADP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004517">
        <property type="term" value="F:nitric-oxide synthase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006809">
        <property type="term" value="P:nitric oxide biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00795">
        <property type="entry name" value="NOS_oxygenase_euk"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.440.10:FF:000001">
        <property type="entry name" value="Endothelial nitric oxide synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.360:FF:000003">
        <property type="entry name" value="Nitric oxide synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1230.10:FF:000001">
        <property type="entry name" value="Nitric oxide synthase, brain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.360">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.990.10">
        <property type="entry name" value="NADPH-cytochrome p450 Reductase, Chain A, domain 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.340.10">
        <property type="entry name" value="Nitric Oxide Synthase, Chain A, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1230.10">
        <property type="entry name" value="Nitric Oxide Synthase, Chain A, domain 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.440.10">
        <property type="entry name" value="Nitric Oxide Synthase,Heme Domain,Chain A domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.30.10">
        <property type="entry name" value="Translation factors"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003097">
        <property type="entry name" value="CysJ-like_FAD-binding"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001094">
        <property type="entry name" value="Flavdoxin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008254">
        <property type="entry name" value="Flavodoxin/NO_synth"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029039">
        <property type="entry name" value="Flavoprotein-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023173">
        <property type="entry name" value="NADPH_Cyt_P450_Rdtase_alpha"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050607">
        <property type="entry name" value="NOS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044943">
        <property type="entry name" value="NOS_dom_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044940">
        <property type="entry name" value="NOS_dom_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044944">
        <property type="entry name" value="NOS_dom_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012144">
        <property type="entry name" value="NOS_euk"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004030">
        <property type="entry name" value="NOS_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036119">
        <property type="entry name" value="NOS_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017938">
        <property type="entry name" value="Riboflavin_synthase-like_b-brl"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43410:SF1">
        <property type="entry name" value="NITRIC OXIDE SYNTHASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43410">
        <property type="entry name" value="NITRIC OXIDE SYNTHASE OXYGENASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00667">
        <property type="entry name" value="FAD_binding_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00258">
        <property type="entry name" value="Flavodoxin_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02898">
        <property type="entry name" value="NO_synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000333">
        <property type="entry name" value="NOS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00369">
        <property type="entry name" value="FLAVODOXIN"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52218">
        <property type="entry name" value="Flavoproteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56512">
        <property type="entry name" value="Nitric oxide (NO) synthase oxygenase domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF63380">
        <property type="entry name" value="Riboflavin synthase domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50902">
        <property type="entry name" value="FLAVODOXIN_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="16" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="8" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="14" id="KW-0274">FAD</keyword>
    <keyword evidence="11" id="KW-0285">Flavoprotein</keyword>
    <keyword evidence="12" id="KW-0288">FMN</keyword>
    <keyword evidence="10 27" id="KW-0349">Heme</keyword>
    <keyword evidence="18 27" id="KW-0408">Iron</keyword>
    <keyword evidence="19" id="KW-0472">Membrane</keyword>
    <keyword evidence="13 27" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="15" id="KW-0521">NADP</keyword>
    <keyword evidence="17" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="9" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="31" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Flavodoxin-like" evidence="29">
        <location>
            <begin position="495"/>
            <end position="678"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="28">
        <location>
            <begin position="1"/>
            <end position="30"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="28">
        <location>
            <begin position="1"/>
            <end position="10"/>
        </location>
    </feature>
    <feature type="binding site" description="axial binding residue" evidence="27">
        <location>
            <position position="159"/>
        </location>
        <ligand>
            <name>heme b</name>
            <dbReference type="ChEBI" id="CHEBI:60344"/>
        </ligand>
        <ligandPart>
            <name>Fe</name>
            <dbReference type="ChEBI" id="CHEBI:18248"/>
        </ligandPart>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001917"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00001950"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00001970"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00001974"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00006267"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00012989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022630"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022643"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022827"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022857"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00029794"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00047419"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00049679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00049764"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00049791"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00049814"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00049818"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="PIRSR" id="PIRSR000333-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="29">
        <source>
            <dbReference type="PROSITE" id="PS50902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="30">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006817.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="31">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="863" mass="95834" checksum="1EADFC0D79BECA87" modified="2019-05-08" version="1">AISCQKTEGQPGTEPPPVFRGNIKTSPASSDLPFAEPVEWLTGAGVHAGSDPNSPQPQLCPSVLQELPCTGKRCLGSLMFPKQMVCQADEGPPAKEELLPLARDFIHQYYSSIKRADSRVHMQRVQEVEQEIRATGTYQLLEPELIFGAKQAWRNAARCLGRIQWNKLQVFDARDCASVQEMFGFLCTHLQYATNRGNLRSAITIFPQRMPGRGDFRIWNSQLIRYAGYPQPDGSVLGDPANVDITELCIQHGWTPGHGRFDVLPLLLQAPDEAPELFLLPPELVLEVPIQHPTLAWFGELGLRWHALPAVANMLLEIGGLQFPAAPFNGWYVGSEIGMRNFCDSNRYNLLQEVALHMGLDTGKTSSLWKDKAAVEINVAVLHSFQVAKVTIVDHHVATESFVKHMDNELQTRGGCPADWVWIVPPISGSLTPVFHQEMVNYLLSPCFRYQPDAWKVYTSKGTSITRKKTFKEVANAVKFSAKLMVHVMARRVKATILYATETGKSRTYAWNLGELFRRAFDPKVLCMDEYDIVSLEHETLVLVVTSTFGNGDPPESGESFAKALMEMSCPYVSSTQPEQQKSYKVRFNSVSQSDGLVASWKKKRKESSNTDSAGALGTLRFSVFGLGSRAYPHFCAFAHAVDTRLEELGGERILAMGEGDELCAQEETFRTWAKRVFQAACDTFCVGDDAEEAAREIFTTRRSWKRQKYQLSVHANPPDMLTGLSQVHKRRVFPCRVLSVQNLQSKESSRSTLLVRLDTAGQPELQYLPGDHVGIFPANRLELVQELLERVEDPPPDEEPVAVEFLEKATSALGAGNLVLVPGGSQGGFWGGQCQCLGGPMRVWGDPGASAWRIPWGVLRGG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXI1</accession>
    <name>A0A452GXI1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">SAM domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GXI1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007953.1">
        <property type="protein sequence ID" value="ENSGAGP00000006854.1"/>
        <property type="gene ID" value="ENSGAGG00000005509.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035102">
        <property type="term" value="C:PRC1 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003682">
        <property type="term" value="F:chromatin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042393">
        <property type="term" value="F:histone binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045892">
        <property type="term" value="P:negative regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd09577">
        <property type="entry name" value="SAM_Ph1_2_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.50:FF:000011">
        <property type="entry name" value="Polyhomeotic-like protein 2 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.50">
        <property type="entry name" value="Transcription Factor, Ets-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050548">
        <property type="entry name" value="PcG_chromatin_remod_factors"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001660">
        <property type="entry name" value="SAM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013761">
        <property type="entry name" value="SAM/pointed_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12247">
        <property type="entry name" value="POLYCOMB GROUP PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12247:SF86">
        <property type="entry name" value="POLYHOMEOTIC-LIKE PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16616">
        <property type="entry name" value="PHC2_SAM_assoc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00536">
        <property type="entry name" value="SAM_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00454">
        <property type="entry name" value="SAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47769">
        <property type="entry name" value="SAM/Pointed domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50105">
        <property type="entry name" value="SAM_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="6" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <keyword evidence="4" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="SAM" evidence="9">
        <location>
            <begin position="102"/>
            <end position="166"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1"/>
            <end position="80"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="1"/>
            <end position="10"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="21"/>
            <end position="51"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="52"/>
            <end position="67"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="71"/>
            <end position="80"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50105"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006854.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="166" mass="18520" checksum="FF94AD3EC1CBA829" modified="2019-05-08" version="1">VHEQAPETRGGHARAPPHLQGSLTCPQQGHQETASLQLSRSQEDSSCCSDNSSYEEPLSPLSASSSTSRRRQGERDLELPDMHMRDLVGIGHHFLPSEPTKWNVEDVYEFIRSLPGCQEIAEEFRAQEIDGQALLLLKEDHLMSAMNIKLGPALKIYARISMLKDS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXL2</accession>
    <name>A0A452GXL2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Acyl-coenzyme A thioesterase THEM4</fullName>
            <ecNumber evidence="19">3.1.2.2</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="21">Thioesterase superfamily member 4</fullName>
        </alternativeName>
    </protein>
    <organism evidence="29 30">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="30" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="29" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="29" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="16">
            <text>(5Z,8Z,11Z,14Z)-eicosatetraenoyl-CoA + H2O = (5Z,8Z,11Z,14Z)-eicosatetraenoate + CoA + H(+)</text>
            <dbReference type="Rhea" id="RHEA:40151"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:32395"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57368"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="16">
            <dbReference type="Rhea" id="RHEA:40152"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>(9Z)-octadecenoyl-CoA + H2O = (9Z)-octadecenoate + CoA + H(+)</text>
            <dbReference type="Rhea" id="RHEA:40139"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30823"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57387"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="17">
            <dbReference type="Rhea" id="RHEA:40140"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="24">
            <text>decanoyl-CoA + H2O = decanoate + CoA + H(+)</text>
            <dbReference type="Rhea" id="RHEA:40059"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:27689"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:61430"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="24">
            <dbReference type="Rhea" id="RHEA:40060"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="25">
            <text>dodecanoyl-CoA + H2O = dodecanoate + CoA + H(+)</text>
            <dbReference type="Rhea" id="RHEA:30135"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:18262"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57375"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="25">
            <dbReference type="Rhea" id="RHEA:30136"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="23">
            <text>hexadecanoyl-CoA + H2O = hexadecanoate + CoA + H(+)</text>
            <dbReference type="Rhea" id="RHEA:16645"/>
            <dbReference type="ChEBI" id="CHEBI:7896"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57379"/>
            <dbReference type="EC" id="3.1.2.2"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="23">
            <dbReference type="Rhea" id="RHEA:16646"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="22">
            <text>octanoyl-CoA + H2O = octanoate + CoA + H(+)</text>
            <dbReference type="Rhea" id="RHEA:30143"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:25646"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57386"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="22">
            <dbReference type="Rhea" id="RHEA:30144"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="26">
            <text>tetradecanoyl-CoA + H2O = tetradecanoate + CoA + H(+)</text>
            <dbReference type="Rhea" id="RHEA:40119"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30807"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57385"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="26">
            <dbReference type="Rhea" id="RHEA:40120"/>
        </physiologicalReaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cell projection</location>
            <location evidence="3">Ruffle membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Mitochondrion inner membrane</location>
            <topology evidence="4">Peripheral membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Mitochondrion intermembrane space</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="18">Belongs to the THEM4/THEM5 thioesterase family.</text>
    </comment>
    <dbReference type="EC" id="3.1.2.2" evidence="19"/>
    <dbReference type="AlphaFoldDB" id="A0A452GXL2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007958.1">
        <property type="protein sequence ID" value="ENSGAGP00000006858.1"/>
        <property type="gene ID" value="ENSGAGG00000005485.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005743">
        <property type="term" value="C:mitochondrial inner membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005758">
        <property type="term" value="C:mitochondrial intermembrane space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032587">
        <property type="term" value="C:ruffle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016787">
        <property type="term" value="F:hydrolase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006915">
        <property type="term" value="P:apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006631">
        <property type="term" value="P:fatty acid metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd03443">
        <property type="entry name" value="PaaI_thioesterase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.129.10">
        <property type="entry name" value="Hotdog Thioesterase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029069">
        <property type="entry name" value="HotDog_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052365">
        <property type="entry name" value="THEM4/THEM5_acyl-CoA_thioest"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006683">
        <property type="entry name" value="Thioestr_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12418">
        <property type="entry name" value="ACYL-COENZYME A THIOESTERASE THEM4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12418:SF19">
        <property type="entry name" value="ACYL-COENZYME A THIOESTERASE THEM4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03061">
        <property type="entry name" value="4HBT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54637">
        <property type="entry name" value="Thioesterase/thiol ester dehydrase-isomerase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0053">Apoptosis</keyword>
    <keyword evidence="5" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="15" id="KW-0966">Cell projection</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="10" id="KW-0276">Fatty acid metabolism</keyword>
    <keyword evidence="9" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="12" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="14" id="KW-0472">Membrane</keyword>
    <keyword evidence="13" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="8" id="KW-0999">Mitochondrion inner membrane</keyword>
    <keyword evidence="30" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="11" id="KW-0809">Transit peptide</keyword>
    <feature type="domain" description="Thioesterase" evidence="28">
        <location>
            <begin position="121"/>
            <end position="193"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="27">
        <location>
            <begin position="1"/>
            <end position="32"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004569"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004632"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004637"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022703"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022792"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022832"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00035852"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00037002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00038456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00038848"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00040123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00043210"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00047588"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00047734"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00047969"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00048074"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00048180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="28">
        <source>
            <dbReference type="Pfam" id="PF03061"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="29">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006858.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="30">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="218" mass="24099" checksum="2570A68334D851CB" modified="2019-05-08" version="1">AARAEALRVRPQRAAEPMEPGGGKRQSKDYALPNSSWSQGMMDLYSRFLEMCKDGTWKRIPSYSNSVDHIPGKKERKDTRLFPRNMDTEGVGFEYAMFLNPSEKRMVCLFQPGPYLEGPSGFAHGGSTATIIDSTIGSCATFVAGRVMTANLSINYRNPIPLGSVVLVDSKVDQVEGRKVFLSCHVRSIDGQTLHAEATALFIRLDSTKSLQQQMSCQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXL5</accession>
    <name>A0A452GXL5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Integrin beta</fullName>
        </recommendedName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 18">Cell membrane</location>
            <topology evidence="1 18">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 18">Belongs to the integrin beta chain family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GXL5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007954.1">
        <property type="protein sequence ID" value="ENSGAGP00000006855.1"/>
        <property type="gene ID" value="ENSGAGG00000001361.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009986">
        <property type="term" value="C:cell surface"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005925">
        <property type="term" value="C:focal adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008305">
        <property type="term" value="C:integrin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001540">
        <property type="term" value="F:amyloid-beta binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019901">
        <property type="term" value="F:protein kinase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033627">
        <property type="term" value="P:cell adhesion mediated by integrin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007160">
        <property type="term" value="P:cell-matrix adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007229">
        <property type="term" value="P:integrin-mediated signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007159">
        <property type="term" value="P:leukocyte cell-cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030593">
        <property type="term" value="P:neutrophil chemotaxis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000076">
        <property type="entry name" value="Integrin beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000098">
        <property type="entry name" value="Integrin beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1680.10:FF:000012">
        <property type="entry name" value="Integrin beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.410:FF:000002">
        <property type="entry name" value="Integrin beta"/>
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        <property type="entry name" value="Laminin"/>
        <property type="match status" value="3"/>
    </dbReference>
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        <property type="entry name" value="ligand-binding face of the semaphorins, domain 2"/>
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    </dbReference>
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        <property type="entry name" value="ntegrin, alpha v. Chain A, domain 3"/>
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    </dbReference>
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        <property type="entry name" value="von Willebrand factor, type A domain"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="EGF_integrin_1"/>
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        <property type="entry name" value="Integrin_beta_N"/>
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        <property type="entry name" value="Integrin_bsu"/>
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        <property type="entry name" value="Integrin_bsu_tail"/>
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        <property type="entry name" value="Integrin_bsu_tail_dom_sf"/>
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        <property type="entry name" value="Integrin_bsu_VWA"/>
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        <property type="entry name" value="Integrin_dom_sf"/>
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        <property type="entry name" value="PSI"/>
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        <property type="entry name" value="vWFA_dom_sf"/>
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        <property type="entry name" value="INTEGRIN BETA-2"/>
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    </dbReference>
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    </dbReference>
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        <property type="entry name" value="INTEGRINB"/>
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        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69687">
        <property type="entry name" value="Integrin beta tail domain"/>
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    </dbReference>
    <dbReference type="SUPFAM" id="SSF69179">
        <property type="entry name" value="Integrin domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103575">
        <property type="entry name" value="Plexin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53300">
        <property type="entry name" value="vWA-like"/>
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        <property type="entry name" value="I_EGF_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0106">Calcium</keyword>
    <keyword evidence="11 18" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="15 17" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="4" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="16" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13 18" id="KW-0401">Integrin</keyword>
    <keyword evidence="10" id="KW-0460">Magnesium</keyword>
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    <keyword evidence="6" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="7 19" id="KW-0732">Signal</keyword>
    <keyword evidence="5 18" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="12" id="KW-1133">Transmembrane helix</keyword>
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        <location>
            <begin position="1"/>
            <end position="25"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019227302" description="Integrin beta" evidence="19">
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            <begin position="26"/>
            <end position="777"/>
        </location>
    </feature>
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        <location>
            <begin position="38"/>
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        </location>
    </feature>
    <feature type="domain" description="Integrin beta subunit VWA" evidence="20">
        <location>
            <begin position="46"/>
            <end position="463"/>
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    </feature>
    <feature type="domain" description="Integrin beta subunit tail" evidence="22">
        <location>
            <begin position="637"/>
            <end position="713"/>
        </location>
    </feature>
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            <begin position="47"/>
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    <feature type="disulfide bond" evidence="17">
        <location>
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        <source>
            <dbReference type="ARBA" id="ARBA00007449"/>
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            <dbReference type="ARBA" id="ARBA00022475"/>
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    </evidence>
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    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
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        </source>
    </evidence>
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    </evidence>
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        </source>
    </evidence>
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        <source>
            <dbReference type="SMART" id="SM00423"/>
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    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="SMART" id="SM01242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006855.1"/>
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    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="777" mass="84818" checksum="31BC40472F456BB4" modified="2019-05-08" version="1">MPGVAVGCPWGLGLSLMGWPSSSHASSLPLCVTALALECPKIKVSTCQDCIRSGPGCAWCKKLNFTKAGEPDSIRCDTKEQLQQRGCDAKDIVFPESAFIQLKNALLDDKTQLAPQQVRLMLRLGQPAVFNVTFRRAEGYPIDLYYLMDLSYSMHDDLEKKVKKLGGDLLAALNSITGSGQIGFGSFVDKTVLPFVNTHPEKLKNPCTNKETTCQPPFAFRHVLSLTENARTFEREVQKQSISGNLDAPEGGLDAMMQVAVCGETIGWRNVTRLLVYATDDGFHFAGDGKLGAILTPNDGRCHLEDNLYKKSNEFDYPSVGQLVQKLAENNIQPIFAVTSRMVGIYQKLRDMIPKSAVGVLHDDSSNVIQLIKEAYNNLSSRVILDHGTLPDSLKITYDSFCHGKKATIDGVRGECGDVKIHDKITFQVKVTAKECIPNQSFTIRPLGFTDTLTVELVSRCDCDCDERQSPTACNGNGKIVCGTCSCNPGHTGKNCECETKGKTSKELEGSCRKDNASVICSGLGDCVCGQCTCHTSDVPNKEIYGTFCECDNMNCEVHNGQLCGGEERGRCDCGECKCSDKFEGSACQCKKSTDGCVNPRGKVCSDRGHCRCNTCQCKAGYQPPFCEECPGCPSPCARYISCVECQAFKSGPFQKNCTDACLNILKLDSLASGRQCREKDSENCWISYLMKQEDGEDKYSFQVIDKKGTGVSSHTPLCATPEAPIGQEPWPMGAVGAVPADGAAHRAPWPHLHILCRSRRGRRESNLQNLPVLPES</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXL9</accession>
    <name>A0A452GXL9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">SHC adaptor protein 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GXL9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007995.1">
        <property type="protein sequence ID" value="ENSGAGP00000006893.1"/>
        <property type="gene ID" value="ENSGAGG00000005552.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030971">
        <property type="term" value="F:receptor tyrosine kinase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007169">
        <property type="term" value="P:cell surface receptor protein tyrosine kinase signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035556">
        <property type="term" value="P:intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd01209">
        <property type="entry name" value="PTB_Shc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd09925">
        <property type="entry name" value="SH2_SHC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000036">
        <property type="entry name" value="SHC-transforming protein 1 isoform 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.505.10:FF:000005">
        <property type="entry name" value="SHC-transforming protein 1 isoform 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.505.10">
        <property type="entry name" value="SH2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051235">
        <property type="entry name" value="CEP152/SHC-Transforming"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006019">
        <property type="entry name" value="PID_Shc-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006020">
        <property type="entry name" value="PTB/PI_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000980">
        <property type="entry name" value="SH2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036860">
        <property type="entry name" value="SH2_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035676">
        <property type="entry name" value="SHC_SH2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10337">
        <property type="entry name" value="SHC TRANSFORMING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10337:SF5">
        <property type="entry name" value="SHC-TRANSFORMING PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00640">
        <property type="entry name" value="PID"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="SH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00401">
        <property type="entry name" value="SH2DOMAIN"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00629">
        <property type="entry name" value="SHCPIDOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00462">
        <property type="entry name" value="PTB"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55550">
        <property type="entry name" value="SH2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01179">
        <property type="entry name" value="PID"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50001">
        <property type="entry name" value="SH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1 2" id="KW-0727">SH2 domain</keyword>
    <feature type="domain" description="PID" evidence="4">
        <location>
            <begin position="133"/>
            <end position="315"/>
        </location>
    </feature>
    <feature type="domain" description="SH2" evidence="5">
        <location>
            <begin position="458"/>
            <end position="549"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="41"/>
            <end position="63"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="425"/>
            <end position="451"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="3">
        <location>
            <begin position="41"/>
            <end position="53"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022999"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00191"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS01179"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50001"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006893.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="8">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="553" mass="60678" checksum="CBC7B06DEE45910D" modified="2019-05-08" version="1">SLPGPGGASVEQEAVTTFCMLIPKIPQWKFSSASGFLSRSPSSASKDLSSPAKPGGHGEAEGSSSGLAAVLSACEPVCHSPCSLQVMNRFRGGGGRKAARVEGVKLAGEEWNRKGSFINKPAQGWLHPDERVLGPGVSYIVRYMGCIEVLRSMRSLDFNTRTQVTREAINRLYEAVPGVKGAWKKKAPNKALFSILGKNNLHFAGMSITVNISIDGLNLMIPTTRQIIANHHMQSISFASGGDTDMTDYVAYVAKDPINQRACHILECCDGLAQSIINTVGQAFELRFKQYLHSPPKVVVPPDRVMGTEESAWGNEEETAEHDYYNSIPGKEPPLGGLIDSRLRHTTLVPGAGPGLEARLCLPGQSCDGYLQADSQPLGCRDYEEHMYVNTQCLDGWEHEASPPKDLFDMRPFEDALKLHESTVAGGASPPLEDQWPSPPTRKAPIAPTEEQLKREPWYHGKMSRRDAEKLLQMDGDFLVRDSITNPGQYVLTGMNSGQPKHLLLVDPEGVVRTKDVLFESISHLINYHLQNEQPIVAAESELHLRQVVKRKQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXM1</accession>
    <name>A0A452GXM1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Acyl-coenzyme A thioesterase THEM4</fullName>
            <ecNumber evidence="19">3.1.2.2</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="21">Thioesterase superfamily member 4</fullName>
        </alternativeName>
    </protein>
    <organism evidence="28 29">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="29" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="28" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="28" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="16">
            <text>(5Z,8Z,11Z,14Z)-eicosatetraenoyl-CoA + H2O = (5Z,8Z,11Z,14Z)-eicosatetraenoate + CoA + H(+)</text>
            <dbReference type="Rhea" id="RHEA:40151"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:32395"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57368"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="16">
            <dbReference type="Rhea" id="RHEA:40152"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>(9Z)-octadecenoyl-CoA + H2O = (9Z)-octadecenoate + CoA + H(+)</text>
            <dbReference type="Rhea" id="RHEA:40139"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30823"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57387"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="17">
            <dbReference type="Rhea" id="RHEA:40140"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="24">
            <text>decanoyl-CoA + H2O = decanoate + CoA + H(+)</text>
            <dbReference type="Rhea" id="RHEA:40059"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:27689"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:61430"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="24">
            <dbReference type="Rhea" id="RHEA:40060"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="25">
            <text>dodecanoyl-CoA + H2O = dodecanoate + CoA + H(+)</text>
            <dbReference type="Rhea" id="RHEA:30135"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:18262"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57375"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="25">
            <dbReference type="Rhea" id="RHEA:30136"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="23">
            <text>hexadecanoyl-CoA + H2O = hexadecanoate + CoA + H(+)</text>
            <dbReference type="Rhea" id="RHEA:16645"/>
            <dbReference type="ChEBI" id="CHEBI:7896"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57379"/>
            <dbReference type="EC" id="3.1.2.2"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="23">
            <dbReference type="Rhea" id="RHEA:16646"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="22">
            <text>octanoyl-CoA + H2O = octanoate + CoA + H(+)</text>
            <dbReference type="Rhea" id="RHEA:30143"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:25646"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57386"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="22">
            <dbReference type="Rhea" id="RHEA:30144"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="26">
            <text>tetradecanoyl-CoA + H2O = tetradecanoate + CoA + H(+)</text>
            <dbReference type="Rhea" id="RHEA:40119"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30807"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57385"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="26">
            <dbReference type="Rhea" id="RHEA:40120"/>
        </physiologicalReaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cell projection</location>
            <location evidence="3">Ruffle membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Mitochondrion inner membrane</location>
            <topology evidence="4">Peripheral membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Mitochondrion intermembrane space</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="18">Belongs to the THEM4/THEM5 thioesterase family.</text>
    </comment>
    <dbReference type="EC" id="3.1.2.2" evidence="19"/>
    <dbReference type="AlphaFoldDB" id="A0A452GXM1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007960.1">
        <property type="protein sequence ID" value="ENSGAGP00000006860.1"/>
        <property type="gene ID" value="ENSGAGG00000005485.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005743">
        <property type="term" value="C:mitochondrial inner membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005758">
        <property type="term" value="C:mitochondrial intermembrane space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032587">
        <property type="term" value="C:ruffle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016787">
        <property type="term" value="F:hydrolase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006915">
        <property type="term" value="P:apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006631">
        <property type="term" value="P:fatty acid metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd03443">
        <property type="entry name" value="PaaI_thioesterase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.129.10">
        <property type="entry name" value="Hotdog Thioesterase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029069">
        <property type="entry name" value="HotDog_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052365">
        <property type="entry name" value="THEM4/THEM5_acyl-CoA_thioest"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006683">
        <property type="entry name" value="Thioestr_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12418">
        <property type="entry name" value="ACYL-COENZYME A THIOESTERASE THEM4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12418:SF19">
        <property type="entry name" value="ACYL-COENZYME A THIOESTERASE THEM4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03061">
        <property type="entry name" value="4HBT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54637">
        <property type="entry name" value="Thioesterase/thiol ester dehydrase-isomerase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0053">Apoptosis</keyword>
    <keyword evidence="5" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="15" id="KW-0966">Cell projection</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="10" id="KW-0276">Fatty acid metabolism</keyword>
    <keyword evidence="9" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="12" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="14" id="KW-0472">Membrane</keyword>
    <keyword evidence="13" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="8" id="KW-0999">Mitochondrion inner membrane</keyword>
    <keyword evidence="29" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="11" id="KW-0809">Transit peptide</keyword>
    <feature type="domain" description="Thioesterase" evidence="27">
        <location>
            <begin position="131"/>
            <end position="203"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004569"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004632"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004637"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022703"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022792"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022832"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00035852"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00037002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00038456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00038848"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00040123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00043210"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00047588"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00047734"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00047969"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00048074"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00048180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="27">
        <source>
            <dbReference type="Pfam" id="PF03061"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="28">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006860.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="29">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="228" mass="25293" checksum="C202AB49F878A333" modified="2019-05-08" version="1">MWRNCVRLARRGLGRGAVLRPDPTVSRCGVSKDYALPNSSWSQGMMDLYSRFLEMCKDGTWKRIPSYSNSVDHIPGKAFWGVHSKYQPQLWQTINMDTEGVGFEYAMFLNPSEKRMVCLFQPGPYLEGPSGFAHGGSTATIIDSTIGSCATFVAGRVMTANLSINYRNPIPLGSVVLVDSKVDQVEGRKVFLSCHVRSIDGQTLHAEATALFIRLDSTKSLQQQMSCQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXN3</accession>
    <name>A0A452GXN3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">ATP binding cassette subfamily C member 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>17beta-estradiol 17-O-(beta-D-glucuronate)(in) + ATP + H2O = 17beta-estradiol 17-O-(beta-D-glucuronate)(out) + ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:60128"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:82961"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="15">
            <dbReference type="Rhea" id="RHEA:60129"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>ATP + H2O + xenobioticSide 1 = ADP + phosphate + xenobioticSide 2.</text>
            <dbReference type="EC" id="7.6.2.2"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="14">
            <text>leukotriene C4(in) + ATP + H2O = leukotriene C4(out) + ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:38963"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:57973"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="14">
            <dbReference type="Rhea" id="RHEA:38964"/>
        </physiologicalReaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the ABC transporter superfamily. ABCC family. Conjugate transporter (TC 3.A.1.208) subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GXN3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008012.1">
        <property type="protein sequence ID" value="ENSGAGP00000006909.1"/>
        <property type="gene ID" value="ENSGAGG00000004641.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016324">
        <property type="term" value="C:apical plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008559">
        <property type="term" value="F:ABC-type xenobiotic transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006869">
        <property type="term" value="P:lipid transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd18595">
        <property type="entry name" value="ABC_6TM_MRP1_2_3_6_D1_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18603">
        <property type="entry name" value="ABC_6TM_MRP1_2_3_6_D2_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd03250">
        <property type="entry name" value="ABCC_MRP_domain1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd03244">
        <property type="entry name" value="ABCC_MRP_domain2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000293">
        <property type="entry name" value="ATP binding cassette subfamily C member 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1560.10:FF:000001">
        <property type="entry name" value="ATP-binding cassette subfamily C member 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1560.10:FF:000007">
        <property type="entry name" value="ATP-binding cassette subfamily C member 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000074">
        <property type="entry name" value="Multidrug resistance-associated protein 5 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1560.10">
        <property type="entry name" value="ABC transporter type 1, transmembrane domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003593">
        <property type="entry name" value="AAA+_ATPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011527">
        <property type="entry name" value="ABC1_TM_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036640">
        <property type="entry name" value="ABC1_TM_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003439">
        <property type="entry name" value="ABC_transporter-like_ATP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017871">
        <property type="entry name" value="ABC_transporter-like_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050173">
        <property type="entry name" value="ABC_transporter_C-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24223">
        <property type="entry name" value="ATP-BINDING CASSETTE SUB-FAMILY C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24223:SF176">
        <property type="entry name" value="ATP-BINDING CASSETTE SUB-FAMILY C MEMBER 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00664">
        <property type="entry name" value="ABC_membrane"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00005">
        <property type="entry name" value="ABC_tran"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00382">
        <property type="entry name" value="AAA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90123">
        <property type="entry name" value="ABC transporter transmembrane region"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50929">
        <property type="entry name" value="ABC_TM1F"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00211">
        <property type="entry name" value="ABC_TRANSPORTER_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50893">
        <property type="entry name" value="ABC_TRANSPORTER_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11" id="KW-0445">Lipid transport</keyword>
    <keyword evidence="12 16" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="9" id="KW-1278">Translocase</keyword>
    <keyword evidence="5 16" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="10 16" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="46"/>
            <end position="67"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="117"/>
            <end position="140"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="146"/>
            <end position="165"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="227"/>
            <end position="251"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="271"/>
            <end position="289"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="581"/>
            <end position="604"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="635"/>
            <end position="654"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="703"/>
            <end position="720"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="726"/>
            <end position="746"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="811"/>
            <end position="832"/>
        </location>
    </feature>
    <feature type="domain" description="ABC transmembrane type-1" evidence="18">
        <location>
            <begin position="10"/>
            <end position="288"/>
        </location>
    </feature>
    <feature type="domain" description="ABC transporter" evidence="17">
        <location>
            <begin position="287"/>
            <end position="518"/>
        </location>
    </feature>
    <feature type="domain" description="ABC transmembrane type-1" evidence="18">
        <location>
            <begin position="637"/>
            <end position="868"/>
        </location>
    </feature>
    <feature type="domain" description="ABC transporter" evidence="17">
        <location>
            <begin position="904"/>
            <end position="1138"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009726"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022967"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00034018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00047523"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00047576"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50893"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50929"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006909.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="21">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1149" mass="127113" checksum="0235C8F27CEB906F" modified="2019-05-08" version="1">RTFVGNLLKSVAFKVVHDLLVFVSPQLLKLLISFVADPTAYAWQGYLYALLLFVTALVQSLCLQQYFQLCFVLGMSLRTALMAAVYKKVSVGTPVSSYTQGETVNLMSADTQRFMDLATFIHQLWSAPLQIALSIIFLWWELGPSVLAGIGVMVLLIPINAVLATKARRIQMKNMKNKDQRMKIMNEILSGIKILKLFAWEPSFERQVGEIRVRELKGLLHFTNLQAFSIFIFSCAPFLVSVASFAVYVMVDEKNVLDAQKAFTSISLFNILRFPMATLPLVISSLVQVSSRGRAERRRVESWALLTHSGALVVTLDIKPGHLVAVVGAVGSGKSSLVSAVLGEMENLKGHINIQGSVAYVPQQAWIQNATLKDNILFGSDLDEARYQQVVEACALLPDLKLLPGGDLTEIGEKGINLSGGQKQRVSLARAVYSDADIYLLDDPLSAVDAHVGKHLFDQVLGPDGLLQGKTRILVTHSVSFLPRTDEIVVLVAGAVSEQGSYSTLLANGGAFAQLLSTYGSQEGSTPEGEATGMWAGADPIANYEQSPYCSLSTSSTKSLRKALQAECAALQVKFSLYLRYLHAVGWGFSAWVFITYVAQYAAFMGSNLWLSDWTDDSLRYWNQTYPTSQRDMRIGVFGALGTVQALLATRGAIRASRVLHQQLLSNILRVPMSFFDTTPTGRIVNRFAKDIFTVDETIPMSFRSWLTCFLGIISTLLMICLATPYFALIIIPLSILYFFLLRYYVSTSRQLRRLDSVTRSPIYSHFGETVSGLSVIRAYGHQERFLQHNDRIVDMNQKSVYSWIVSNRWLAVRLEFVGNLVVFSAALLAVITRDNLGSGIVGLSISSALNITQTLNWLVRMTSELETNIVAVERVREYTEVENEAQWVTAQRPPPGWPSKGEIQITDYQVRYRPGLELVLHGLTCDIGSTEKVGVVGRTGAGKSSLTNCLFRILEAAGGKIVIDGVDISTIGLHDLRQSLTIIPQDPVLFSGTLRMNLDPFDQHSDEEIWKALELAHLKTYVLDLPERLSHQVSEAGENLSVGQRQLLCLARALLRKSKILILDEATAAVDLETDHLIQTTIRSEFANCTVLTIAHRLHTIMDSNRVMVLQAGRIVEYDSPERLLQQQGLFSTMAKDAGIVSAQATVL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXN4</accession>
    <name>A0A452GXN4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Centromere protein J</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the TCP10 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GXN4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007884.1">
        <property type="protein sequence ID" value="ENSGAGP00000006792.1"/>
        <property type="gene ID" value="ENSGAGG00000005474.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005814">
        <property type="term" value="C:centriole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015631">
        <property type="term" value="F:tubulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061511">
        <property type="term" value="P:centriole elongation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060271">
        <property type="term" value="P:cilium assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.450.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009852">
        <property type="entry name" value="CENPJ_C_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047002">
        <property type="entry name" value="Tcp10_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026581">
        <property type="entry name" value="TCP10L/CENPJ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058029">
        <property type="entry name" value="Tubulin-bd_CENPJ"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10331">
        <property type="entry name" value="T COMPLEX PROTEIN 10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10331:SF25">
        <property type="entry name" value="T-COMPLEX PROTEIN 10A-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07202">
        <property type="entry name" value="Tcp10_C"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25779">
        <property type="entry name" value="Tubulin-bind_CPAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="CENPJ tubulin-binding region" evidence="5">
        <location>
            <begin position="173"/>
            <end position="235"/>
        </location>
    </feature>
    <feature type="domain" description="Centromere protein J C-terminal" evidence="4">
        <location>
            <begin position="1064"/>
            <end position="1097"/>
        </location>
    </feature>
    <feature type="domain" description="Centromere protein J C-terminal" evidence="4">
        <location>
            <begin position="1137"/>
            <end position="1168"/>
        </location>
    </feature>
    <feature type="domain" description="Centromere protein J C-terminal" evidence="4">
        <location>
            <begin position="1175"/>
            <end position="1207"/>
        </location>
    </feature>
    <feature type="domain" description="Centromere protein J C-terminal" evidence="4">
        <location>
            <begin position="1211"/>
            <end position="1241"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="81"/>
            <end position="139"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="305"/>
            <end position="334"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="366"/>
            <end position="394"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="483"/>
            <end position="557"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="575"/>
            <end position="659"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="876"/>
            <end position="1015"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="1028"/>
            <end position="1063"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="2">
        <location>
            <begin position="784"/>
            <end position="850"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="3">
        <location>
            <begin position="83"/>
            <end position="94"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="116"/>
            <end position="138"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="306"/>
            <end position="334"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="375"/>
            <end position="394"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="495"/>
            <end position="509"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="3">
        <location>
            <begin position="598"/>
            <end position="608"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="637"/>
            <end position="654"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="884"/>
            <end position="895"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="900"/>
            <end position="911"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="917"/>
            <end position="927"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="928"/>
            <end position="937"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="3">
        <location>
            <begin position="944"/>
            <end position="961"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="991"/>
            <end position="1015"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="1028"/>
            <end position="1052"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="1053"/>
            <end position="1063"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00005627"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="Pfam" id="PF07202"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="Pfam" id="PF25779"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006792.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="8">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1253" mass="141475" checksum="11178D7726CD9106" modified="2019-05-08" version="1">MMEQMEQLQRLVTEQQKIITYYNPGFSVYPGTPSQLIAMVPTLPRFPATLFPVQLPLENSSPVQNHGHLQTSPSIMQTSLQRCSPASSPNNNCSETLGKAIEPLSRSAGSPPKKLSRQDAQEEEITASKDESPLKEETYPNTLPAIKEEEGEQTDEQIPLSPFGIRMDARTRNLEDRPIRPGIGVRQKTFEEFVEEQLKVDSQITENHQQNFCEAKATPRESFLKRGEGIARLEKNKENLAKEQGKHPRRVSFCQNIFSFPHQQDAGKLQLGKHLNRLHQRSSSPTVLASNEKNTNCTISMSEIKAQVDSKTRDPEQCPEERREGGGGEECAKEDPAVPLQMNWAELLQQCQEQITEMKLQIGEKNKAQSTDSLGHSDRADRSSMESTVQKDLDIVDQRVKGNLTQKAEKPLEILQEGSKLQNLEGSQTKQMDWSTGLKFTSGMKGISSCHNEDYTVSKSQESQSGTTTGFKMVNDRIMKVTHKSTQGMDKKKNITSVSHQQEWQSNRSAAYRWKAPPLSSGSGCTSTDSEDDPKSYCTRSPTMHVPQKVGHTDKNLDLSDADYATDELSGAEDLALKKHTKPPPKKLSAQEIKAKQDLSLSTSSSDSGNRVARLRGNKACSSLRKSPFRPPRSARGGREPEPRSERNVEDGKNVELQSSPSTCDLVASLFPAFKSKETLAEERAQRKSVLGTLEAVQRRLTDKLEELEKEVGVYHGETPLLAMMKEEQEKARHFLRTQMDQFETSKAQELNCLEEYKRDCIHTPKKEKVEFKKYATAARVTREDVKSEEIQMLKQQIAGLQEEFRRNESHWHAAHGKLKSQIEMLTKQNLELRDELRVSEHQRLEAEKKSGAVDFISRKSETPVSEAILRGTSSLANLEERSLQSSPKSRNTTPVGRKTPVEKHPPRDVNMKAMKSTVQRTESLKSVTRESREKTPSNHFHSRSTTPTGRRTPHPGRLTPFEPEKVMHQSSLNEQRTYGRKSPVIPVSHLTVSKETNSSSYIKGRFSSTSGSSEDTVLFRNQNEDVCSSASCSNNEETQTSPPKSTLSRRSSLYEESKKRDEEVREKIEYADGKVEEVLTDGRRIITFRNGTKKEISADKRMTVVTFFNGDVKKIMPDQRVIYYYADAETTHTTYPNGLEVLQFPNNQIEKHHPDGTKEIVFPDQTVKRFYDGGLEETVFPDGTVVKVEKNGDKMVMFSNGQKEIHTSQFKRREYPDGTIKTVYSSGQQETKYSSGRVRIKDEEGNIILNKK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GY00</accession>
    <name>A0A452GY00_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">GYF domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the GIGYF family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GY00"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008092.1">
        <property type="protein sequence ID" value="ENSGAGP00000006985.1"/>
        <property type="gene ID" value="ENSGAGG00000001575.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043204">
        <property type="term" value="C:perikaryon"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1990635">
        <property type="term" value="C:proximal dendrite"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031982">
        <property type="term" value="C:vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048009">
        <property type="term" value="P:insulin-like growth factor receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00072">
        <property type="entry name" value="GYF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1490.40">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051640">
        <property type="entry name" value="GRB10-interact_GYF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003169">
        <property type="entry name" value="GYF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035445">
        <property type="entry name" value="GYF-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14445">
        <property type="entry name" value="GRB10 INTERACTING GYF PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14445:SF38">
        <property type="entry name" value="GRB10-INTERACTING GYF PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02213">
        <property type="entry name" value="GYF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00444">
        <property type="entry name" value="GYF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55277">
        <property type="entry name" value="GYF domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50829">
        <property type="entry name" value="GYF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="GYF" evidence="4">
        <location>
            <begin position="522"/>
            <end position="570"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="97"/>
            <end position="234"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="299"/>
            <end position="430"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="901"/>
            <end position="944"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="956"/>
            <end position="989"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="1028"/>
            <end position="1054"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="2">
        <location>
            <begin position="674"/>
            <end position="855"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="139"/>
            <end position="170"/>
        </location>
    </feature>
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        <location>
            <begin position="180"/>
            <end position="199"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="213"/>
            <end position="226"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="3">
        <location>
            <begin position="306"/>
            <end position="323"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="325"/>
            <end position="349"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="361"/>
            <end position="379"/>
        </location>
    </feature>
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        <location>
            <begin position="384"/>
            <end position="402"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="901"/>
            <end position="918"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="3">
        <location>
            <begin position="922"/>
            <end position="932"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="970"/>
            <end position="989"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="3">
        <location>
            <begin position="1034"/>
            <end position="1048"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00038015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50829"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006985.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1241" mass="143177" checksum="E51EA0BA68539F89" modified="2025-10-08" version="2">MAALRALSSGGSVTSPPLSPALPKYKLADYRYGREEMLALFLKDNKIPSDLLDKEFQPILQEEPLPPLALVPFTEEEQRNFSMSVNSAAVLRLTGRGGGTVVGAPRGRSSSRGRGRGRGESGFYQRSFDEVEGGFGRGGGREMHRSQSWEERGDRRFEKPGRKDPVRPNFEEGGTTTTGRKHEFIRSESENWRIFREEQNGEDDDGGWRLAGPRRDGERWRPHSPDGPRSAGWREPMERRRRFEFDFRDRDDERGYRRVRCGSGSMEDDRDSLPEWCLEDAEEEMGTFDSSGAFLSLKKVQKESIPEEQEMDFQPVEGEEPSDPEGGHGEEAKELEKTTKREGDKRDTTTAEAAEEAVRASSPSIRSGTPPKLQSQEPLQASLFERKEECVPERTEKMEAKESQTVNTLPSKVPSRGEDLTPAVQPLPQISPATVSPVHLSSPLSNTNPALKPLQTSVSAAPGMGNISADPDDEEGLKHLEQQAEKMVAYLQDSALDDERLAAKLQEHRAKGSSMPLLHEAMQKWYYKDPQGEIQGPFSNQEMAEWFQAGYFTMSLLVKRACDENFQPLGDIMKMWGRVPFTPGPAPPPHLGEMDQERLTRQQELTALYQMQHLQYQQFLIQQQYAQVLAQQQKAALSSQQQQQLALLLQQFQALKMSSVWDLPIDAAAQEPALEQLQQLEKAKAVKLEQERREMEMRAKREEEERKRQEELRRQQEELLRRQQEEERKRREEEELARRKQEEALRRQREQEIALRRQREEEERQQQEEALRRLEERRREEEERRQREELLRKQEEEAAKWAREEEEAQRRLEENRLHMEEEAARLRLEEEERKRKELELQRQKELLRQRQQQQEALQFVIIISCFSQLPSSSTWGQQSSSGACQSQTTLSLAEIQKLEEERERQLREEQRRQQRELMKALQQQQQQQQQQQKMSGWGNVTKPTGATKSLLEIQQEEARQMQKQQQQQQPNRVRSTTHSSLHSSIGNSVWGSLNTNPTSQWASDLVTSIWSNSDTKNSNMGFWDDAVKEVGPRNSTNKNKSNASLSKSIGITSRQNKKVEQEEKLLKLFQGVNKAQDGFTQWCEQMLHALNTANNLDVPTFVSFLKEVESPYEVHDYIRAYLGDTPEAKEFAKQFLERRAKQKASQQQQQMSQYLGICPASQPEFEFLFQDSVWGMNHSALHSVFQTNQSNNQQSNFEAVQSSKRKKKQKMVRADPSLLGFSVNASSERLNMGEIETLEDY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GY04</accession>
    <name>A0A452GY04_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">SH3 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GY04"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008154.1">
        <property type="protein sequence ID" value="ENSGAGP00000007038.1"/>
        <property type="gene ID" value="ENSGAGG00000005659.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002039">
        <property type="term" value="F:p53 binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006915">
        <property type="term" value="P:apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042981">
        <property type="term" value="P:regulation of apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd11953">
        <property type="entry name" value="SH3_ASPP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000008">
        <property type="entry name" value="Apoptosis-stimulating of p53 protein 2 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
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    </dbReference>
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        <property type="entry name" value="SH3_domain"/>
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        <property type="entry name" value="APOPTOSIS-STIMULATING OF P53 PROTEIN"/>
        <property type="match status" value="1"/>
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    <dbReference type="PANTHER" id="PTHR24131:SF8">
        <property type="entry name" value="APOPTOSIS-STIMULATING OF P53 PROTEIN 2"/>
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        <property type="entry name" value="Ank_2"/>
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        <property type="entry name" value="SPECTRNALPHA"/>
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    </dbReference>
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    </dbReference>
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    </dbReference>
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    </dbReference>
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    <keyword evidence="3" id="KW-0053">Apoptosis</keyword>
    <keyword evidence="9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="6" id="KW-0539">Nucleus</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="2 8" id="KW-0728">SH3 domain</keyword>
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        <location>
            <begin position="913"/>
            <end position="945"/>
        </location>
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            <end position="978"/>
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    </feature>
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            <begin position="1012"/>
            <end position="1074"/>
        </location>
    </feature>
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            <begin position="19"/>
            <end position="58"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
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            <begin position="344"/>
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    </feature>
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        </location>
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            <begin position="490"/>
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        </location>
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    </feature>
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            <begin position="722"/>
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            <begin position="66"/>
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    </feature>
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            <begin position="722"/>
            <end position="749"/>
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    </feature>
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        <source>
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    </evidence>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022703"/>
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    </evidence>
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            <dbReference type="ARBA" id="ARBA00022737"/>
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    </evidence>
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            <dbReference type="ARBA" id="ARBA00023043"/>
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    </evidence>
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    </evidence>
    <evidence type="ECO:0000256" key="8">
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    </evidence>
    <evidence type="ECO:0000256" key="9">
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    </evidence>
    <evidence type="ECO:0000256" key="10">
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    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007038.1"/>
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    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1083" mass="120301" checksum="2FD9EB0D6EEF51E4" modified="2019-05-08" version="1">MLDILQHFGVQRNEVRFFLRHERPPSRETGSGQRPQDPSLKRNGVKVPGDRRLENGISVPRMDMTLAELQEMASRQQQQIEAQQQMLANKEQRLKFLKQQDQRQQQQAAEKEKLKRLKEIAENQEAKLKKVRALKGHVEQKRLSNGKLVEEIEQMNSLFQQKQRELVLAVSKVEELTRQLEMLKNGRIDGYHDNQSAVAELDRLYKELQLRNKLNQEQNAKLQQQRECLNKRNSEVAVMDKRVNELRDRLWKKKAALQQKENVPISSDGNPPQQVVSAPSRVAAVGPYIQSSTMPRVASRPELLVKPAFPEGTPTLQAPDGPLKTQTLPNMRAVGVSQAKAPAGLTIPGSKPPPAVSEWCNSNADNFISQGPVSSSAKGNVTNEGQAGDGETPLREKEKKVRPFSMFDSVDQPTGGPALGSLRKNQSSEDLLRDGQIANKNVTKVPPPVPTKPKQINLPYFGQASQLPSSETKPDGNPQKLPLAIVTMGNKQKSAAQQPPHTSQQIQQRISVPPSGPSTIQDQILSSSKQESPPAAAVRPFTPQPSKEIPLPSFRKPQTVAASSIYSMYTQQQTPGKNFQQAVQSALTRAQTRGPHFSSMYGKPVIAGIGAQNQSQQTENVYSNSHGSPEPEMETIASVHENHETERIPRPLSPTKLLPFLSNPYRNQSDADLEALRKKLSNAPRPLKKRSSITEPEGPNGPNIQKLLYQRTTLAAMETISTPSYPSKQTSMSASPESPTEIQNPYLSTESEKEVVPSTPEPAITEEAENTHTDQNEVPLPSAGLDSVPEVTSDSDALVQSEMEESSLESSEPLEVYMEEYPPYPPPPYPSGEPGGLGEDSFSMRPPEVTGQFSLPPGKRTNLRKTGSERIAHGMRVKFNPLALLLDSSLEGEFDLVQRIIYEVEDPSLPNDEGITALHNAVCAGHTEIVKFLVQFGVNVNAADSDGWTPLHCAASCNNVQVCKFLVESGAAVFAMTYSDMQTAADKCEEMEEGYTQCSQFLYGVQEKMGIMNKGVIYALWDYDAQNDDELSMKEGDCMTVLRREDEDEIEWWWARLNDKEGYIPRNLLGLYPRIKPRQRSLA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GY05</accession>
    <name>A0A452GY05_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">GYF domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the GIGYF family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GY05"/>
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        <property type="protein sequence ID" value="ENSGAGP00000006991.1"/>
        <property type="gene ID" value="ENSGAGG00000001575.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="project" value="TreeGrafter"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1990635">
        <property type="term" value="C:proximal dendrite"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="C:vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="P:insulin-like growth factor receptor signaling pathway"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="match status" value="1"/>
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        <property type="match status" value="1"/>
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        <property type="entry name" value="GYF"/>
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        <property type="entry name" value="GYF-like_dom_sf"/>
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    <dbReference type="PANTHER" id="PTHR14445">
        <property type="entry name" value="GRB10 INTERACTING GYF PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14445:SF38">
        <property type="entry name" value="GRB10-INTERACTING GYF PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
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    </dbReference>
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    </dbReference>
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    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
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            <end position="570"/>
        </location>
    </feature>
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            <begin position="1"/>
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    </feature>
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        <location>
            <begin position="922"/>
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        </location>
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        </location>
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        <location>
            <begin position="1033"/>
            <end position="1045"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00038015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="PROSITE" id="PS50829"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006991.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1054" mass="121627" checksum="DA172B28000BD6BF" modified="2025-10-08" version="2">MAALRALSSGGSVTSPPLSPALPKYKLADYRYGREEMLALFLKDNKIPSDLLDKEFQPILQEEPLPPLALVPFTEEEQRNFSMSVNSAAVLRLTGRGGGTVVGAPRGRSSSRGRGRGRGESGFYQRSFDEVEGGFGRGGGREMHRSQSWEERGDRRFEKPGRKDPVRPNFEEGGTTTTGRKHEFIRSESENWRIFREEQNGEDDDGGWRLAGPRRDGERWRPHSPDGPRSAGWREPMERRRRFEFDFRDRDDERGYRRVRCGSGSMEDDRDSLPEWCLEDAEEEMGTFDSSGAFLSLKKVQKESIPEEQEMDFQPVEGEEPSDPEGGHGEEAKELEKTTKREGDKRDTTTAEAAEEAVRASSPSIRSGTPPKLQSQEPLQASLFERKEECVPERTEKMEAKESQTVNTLPSKVPSRGEDLTPAVQPLPQISPATVSPVHLSSPLSNTNPALKPLQTSVSAAPGMGNISADPDDEEGLKHLEQQAEKMVAYLQDSALDDERLAAKLQEHRAKGSSMPLLHEAMQKWYYKDPQGEIQGPFSNQEMAEWFQAGYFTMSLLVKRACDENFQPLGDIMKMWGRVPFTPGPAPPPHLGEMDQERLTRQQELTALYQMQHLQYQQFLIQQQYAQVLAQQQKAALSSQQQQQLALLLQQFQALKMSSVWDLPIDAAAQEPALEQLQQLEKAKAVKLEQERREMEMRAKREEEERKRQEELRRQQEELLRRQQEEERKRREEEELARRKQEEALRRQREQEIALRRQREEEERQQQEEALRRLEERRREEEERRQREELLRKQEEEAAKWAREEEEAQRRLEENRLHMEEEAARLRLEEEERKRKELELQRQKELLRQRQQQQEALQFVIIISCFSQLPSSSTWGQQSSSGACQSQTTLSLAEIQKLEEERERQLREEQRRQQRELMKALQQQQQQQQQQQKMSGWGNVTKPTGATKSLLEIQQEEARQMQKQQQQQQPNRVRSTTHSSLHSSIGNSVWGSLNTNPTSQWASDLVTSIWSNSDTKNSNMGFWDDAVKEVGPRNSTNKNKSNASLSPPGFLRNA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GY09</accession>
    <name>A0A452GY09_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="7">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GY09"/>
    <dbReference type="Ensembl" id="ENSGAGT00000008159.1">
        <property type="protein sequence ID" value="ENSGAGP00000007043.1"/>
        <property type="gene ID" value="ENSGAGG00000005663.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015179">
        <property type="term" value="F:L-amino acid transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1740.10:FF:000056">
        <property type="entry name" value="Y+L amino acid transporter 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1740.10">
        <property type="entry name" value="Amino acid/polyamine transporter I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002293">
        <property type="entry name" value="AA/rel_permease1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050598">
        <property type="entry name" value="AminoAcid_Transporter"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11785">
        <property type="entry name" value="AMINO ACID TRANSPORTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11785:SF323">
        <property type="entry name" value="CYSTINE_GLUTAMATE TRANSPORTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13520">
        <property type="entry name" value="AA_permease_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4 6" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 6" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="3 6" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="46"/>
            <end position="64"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="76"/>
            <end position="98"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="133"/>
            <end position="151"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="163"/>
            <end position="186"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="192"/>
            <end position="214"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="226"/>
            <end position="246"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="1"/>
            <end position="29"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="9"/>
            <end position="20"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007043.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="249" mass="26957" checksum="D732CA993EF44C2E" modified="2019-05-08" version="1">MFRKPVLSTIPNGSSFQGQSIGRLPSLDSKEPTQEEKVVLKKKVTFLRGVSIIIGTIIGAGIFISPKGILKNTGSVGMSLILWTVCGILSLFGALCYAELGTCIKKSGGHYAYILEAFGPLPAFVRVWVELLIIRPAATAVISLAFGRYILEPFFIQCEIPDLAIKLITAVGITVVMVLNSMSVSWSARIQIFLTFCKLVAILIIIVPGVMQLIKGETQHFKDAFAGNDASVMGLPLAFYSGMYAYSGW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9H0</accession>
    <name>A0A452H9H0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Anoctamin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="8">Membrane</location>
            <topology evidence="8">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 8">Belongs to the anoctamin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9H0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013058.1">
        <property type="protein sequence ID" value="ENSGAGP00000011407.1"/>
        <property type="gene ID" value="ENSGAGG00000007328.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005229">
        <property type="term" value="F:intracellularly calcium-gated chloride channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046983">
        <property type="term" value="F:protein dimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032394">
        <property type="entry name" value="Anoct_dimer"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007632">
        <property type="entry name" value="Anoctamin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049452">
        <property type="entry name" value="Anoctamin_TM"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12308">
        <property type="entry name" value="ANOCTAMIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12308:SF28">
        <property type="entry name" value="ANOCTAMIN-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16178">
        <property type="entry name" value="Anoct_dimer"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04547">
        <property type="entry name" value="Anoctamin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6 8" id="KW-0472">Membrane</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 8" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 8" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="326"/>
            <end position="352"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="402"/>
            <end position="421"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="478"/>
            <end position="504"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="524"/>
            <end position="544"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="572"/>
            <end position="593"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="692"/>
            <end position="714"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="742"/>
            <end position="766"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="827"/>
            <end position="848"/>
        </location>
    </feature>
    <feature type="domain" description="Anoctamin dimerisation" evidence="11">
        <location>
            <begin position="93"/>
            <end position="312"/>
        </location>
    </feature>
    <feature type="domain" description="Anoctamin transmembrane" evidence="10">
        <location>
            <begin position="315"/>
            <end position="870"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="869"/>
            <end position="896"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009671"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="RuleBase" id="RU280814"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="Pfam" id="PF04547"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF16178"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011407.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="905" mass="105533" checksum="0659A9F34C1514A8" modified="2019-05-08" version="1">PAASQELHYAPQVPASTALSKVLASFHLQGTQCFLLELQGVTAPALPCGGSYMGLLSPDWLSLGSNGDSQLCFGNCFCSSQGVPDKNKSNGLYFRDGKCRIDYILVYRKSSPQTEKREIFERNIRAEGLQMEKEVRDDIIFVKLHAPWEVLGRYAEVMNVRMPFRRKIYYLHRRHKFMNRQISRFQGWLPRKPMRLDKETLPDLEENDCYTAPFSQQRIHHFIIHNKDTFFNNATRSRIVHHILQRVKYEEGKNKIGLNRLLTNGSYEAAFPLHEGSYRSKNSIRTHGAENHRHLLYECWASWGMWYKYQPLDLVRRYFGEKIGLYFAWLGWYTGMLFPAAFIGLFVFLYGVTTLNHCQVSKEVCQATDIIMCPICDKFCPFMRLSDSCVYAKVTHLFDNGATVFFAVFMAVWATVFLEFWKRRRAVIAYDWDLIDWEEEEEEIRPQFEAKYSKKERMNPISGKPEPYQAFADKCSRLIVSASGIFFMICVVIAAVFGIVIYRVVTVSTFAAFKWALIRNNSQVATTGTAVCINFCIIMLLNLYEKVALLLTNLEQPRTESEWENSFTLKMFLFQFVNLNSSTFYIAFFLGRFTGHPGAYLRLINRWRLEECHPSGCLIDLCMQMGIIMVLKQTWNNFMELGYPLIQNWWTRRKLYQEHGSQRKTSFPQWEKDYNLQPMNAYGLFDEYLEMILQFGFTTIFVAAFPLAPLLALLNNIIEIRLDAYKFVTQWRRPLASRAKDIGIWYGILEGIGILSVITNAFVVAVTSDFIPRLVYAYKYGPCAGQGEAGQKCMVGYVNASLSVFLVYRDYRDPPHSPVPYGYTLQFWHVLAARLAFIIVFEHLVFCIKHLISYLIPDLPKDLRDRMRREKYLIQEMMYEAELERLQKERKERKKNGKSHHNEWP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9H1</accession>
    <name>A0A452H9H1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">C3H1-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9H1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013064.1">
        <property type="protein sequence ID" value="ENSGAGP00000011411.1"/>
        <property type="gene ID" value="ENSGAGG00000008803.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035198">
        <property type="term" value="F:miRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035196">
        <property type="term" value="P:miRNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010608">
        <property type="term" value="P:post-transcriptional regulation of gene expression"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000070">
        <property type="entry name" value="zinc finger CCCH domain-containing protein 7B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.10">
        <property type="entry name" value="Tetratricopeptide repeat domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011990">
        <property type="entry name" value="TPR-like_helical_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019734">
        <property type="entry name" value="TPR_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039691">
        <property type="entry name" value="ZC3H7A/B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000571">
        <property type="entry name" value="Znf_CCCH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036855">
        <property type="entry name" value="Znf_CCCH_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14928">
        <property type="entry name" value="MICRO-RNA BINDING ZINC FINGER CCCH DOMAIN-CONTAINING PROTEIN 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14928:SF6">
        <property type="entry name" value="ZINC FINGER CCCH DOMAIN-CONTAINING PROTEIN 7B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00642">
        <property type="entry name" value="zf-CCCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00028">
        <property type="entry name" value="TPR"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00356">
        <property type="entry name" value="ZnF_C3H1"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90229">
        <property type="entry name" value="CCCH zinc finger"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48452">
        <property type="entry name" value="TPR-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50005">
        <property type="entry name" value="TPR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50103">
        <property type="entry name" value="ZF_C3H1"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3 9" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="8" id="KW-0802">TPR repeat</keyword>
    <keyword evidence="6 9" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 9" id="KW-0863">Zinc-finger</keyword>
    <feature type="repeat" description="TPR" evidence="8">
        <location>
            <begin position="82"/>
            <end position="115"/>
        </location>
    </feature>
    <feature type="domain" description="C3H1-type" evidence="10">
        <location>
            <begin position="517"/>
            <end position="541"/>
        </location>
    </feature>
    <feature type="domain" description="C3H1-type" evidence="10">
        <location>
            <begin position="649"/>
            <end position="671"/>
        </location>
    </feature>
    <feature type="domain" description="C3H1-type" evidence="10">
        <location>
            <begin position="787"/>
            <end position="815"/>
        </location>
    </feature>
    <feature type="domain" description="C3H1-type" evidence="10">
        <location>
            <begin position="919"/>
            <end position="947"/>
        </location>
    </feature>
    <feature type="zinc finger region" description="C3H1-type" evidence="9">
        <location>
            <begin position="517"/>
            <end position="541"/>
        </location>
    </feature>
    <feature type="zinc finger region" description="C3H1-type" evidence="9">
        <location>
            <begin position="649"/>
            <end position="671"/>
        </location>
    </feature>
    <feature type="zinc finger region" description="C3H1-type" evidence="9">
        <location>
            <begin position="787"/>
            <end position="815"/>
        </location>
    </feature>
    <feature type="zinc finger region" description="C3H1-type" evidence="9">
        <location>
            <begin position="919"/>
            <end position="947"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00339"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50103"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011411.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="990" mass="111794" checksum="F9FD565848F49251" modified="2019-05-08" version="1">MERQKRKEEIEKGLQFIQSTLPLSQEDYEAFIQKLVQNLFAEGNDLFREKDFKLSLVQYVEGLNVADYAASDEVTIPIELLCKLHVNRAACYFAMGLYEKALEDSEKALMLDKENIRALFRKARSLNELGRHKEAYECNSRCLLSLPHDESVTQLGQELAQKLGLRVRKAYKRPQMTSNGLGSIDDIETDFSMDLQCIPAPLATSIPVSEGAVLSASDMDAKGLPTSLPAATLLHSPDCVSLPVPESVEDFTDGEVIGEELDSLLDSLADGSQYPLSLVQGTIPTNLPTEMPQLIPVFPGGTPLLPPVVTANIPINQHPLPPASFGLVMDPTKQLSSSVLDAFEPTGGSGGSPLDSLDSLDLLPYTEPRLDALDAFGTSRGSLDALDSFAVEEAGPQELRHTAGSQKPSPAISLSGVSHPTIPKGAELLQSQPEAVIPNTALLVKNPLASTHVFKQACHLCYPKTGPKAGDYTYREGLEHKCKRDILLGRLRNSEDKTWKRIRPRPTKTNFVGSYYLCKDMINKQDCKYGDNCTFAYHQEEIDVWTEERKGTLNRDLLFDPLGGIKRGNLTIAKLLQEHQGIFTFLCEICFDSKPRIISKGTKDSPAVCSNLAAKHSFYDNKCLVHIVRSTSLKYSKIRQFQEHFQFDVCRHEVRYGCLREDSCHFAHSFIELKVWLLQQYSGMTHEDIVQESKKYWQLMEVHASKATSGTAAPRAPTSSSFDLQMKFVCGQCWRNGQVVEPDKDLKYCSAKARHCWTKERRVLLVMSKAKRKWVSVRPLPSIRNFPQQYDLCIHAQNGRKCQYVGNCSFAHSPEERDMWTFMKENKILDMQQTYDMWLKKHNPGKPGEGTPLTSREGEKQIQMPTDYADIMMGYHCWLCGKNSNSKKQWQQHIQSEKHKEKVFTSDNDSSCWNYRFPMGEFRLCDRFQKTKACPEGDKCCCAHGQDELTEWLDRREVLKQKLAKARKDMLLCPKDDDFGKYNFLLRDGV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9H2</accession>
    <name>A0A452H9H2_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452H9H2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000012869.1">
        <property type="protein sequence ID" value="ENSGAGP00000011236.1"/>
        <property type="gene ID" value="ENSGAGG00000008690.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043123">
        <property type="term" value="P:positive regulation of canonical NF-kappaB signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001819">
        <property type="term" value="P:positive regulation of cytokine production"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039499">
        <property type="entry name" value="LURA1/LRA25"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037443">
        <property type="entry name" value="LURAP1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR33767:SF2">
        <property type="entry name" value="LEUCINE RICH ADAPTOR PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR33767">
        <property type="entry name" value="LEUCINE RICH ADAPTOR PROTEIN 1-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14854">
        <property type="entry name" value="LURAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="148"/>
            <end position="191"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="1"/>
            <end position="10"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011236.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="218" mass="24185" checksum="F38A4A9F9FEF5618" modified="2019-05-08" version="1">NRGSVSSQKSLPHLESKVGRRPPEGLVRWLREDPAAALLRESPVRGQRRGLAGKIKALKLELAYLRAIDVKILQQLVVVNEGIEAVKWLLEEKGTLTSHCSSLASSQYSLVESQETSRRGSWTSLQDPGEKLDSISVGSYLDTLADEMDEPGIPPARLEQDWSRSSPDRGLPSKPASKEAQGARLSKGSPTRMLNGKLHLEYDAHWHWVQSQDDVTFL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9H5</accession>
    <name>A0A452H9H5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">BZIP domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the bZIP family. PAR subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9H5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013073.1">
        <property type="protein sequence ID" value="ENSGAGP00000011418.1"/>
        <property type="gene ID" value="ENSGAGG00000008808.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd14695">
        <property type="entry name" value="bZIP_HLF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.170:FF:000007">
        <property type="entry name" value="hepatic leukemia factor isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.170">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004827">
        <property type="entry name" value="bZIP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046347">
        <property type="entry name" value="bZIP_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040223">
        <property type="entry name" value="PAR_bZIP"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11988:SF24">
        <property type="entry name" value="THYROTROPH EMBRYONIC FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11988">
        <property type="entry name" value="THYROTROPH EMBRYONIC FACTOR RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07716">
        <property type="entry name" value="bZIP_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00338">
        <property type="entry name" value="BRLZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57959">
        <property type="entry name" value="Leucine zipper domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50217">
        <property type="entry name" value="BZIP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0010">Activator</keyword>
    <keyword evidence="4" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0804">Transcription</keyword>
    <keyword evidence="3" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="BZIP" evidence="9">
        <location>
            <begin position="218"/>
            <end position="281"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="30"/>
            <end position="50"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="118"/>
            <end position="160"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="32"/>
            <end position="43"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="118"/>
            <end position="130"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="131"/>
            <end position="142"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009208"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50217"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011418.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="294" mass="32687" checksum="C9DC477A1E2728B4" modified="2019-05-08" version="1">MSGCDNLGVPPAMDFPEVLKSLLEYSLPWADTRSDKEKEKEKLEEDEAAAASTMAVSASLMPPIWDKTIPYDGESFHLEYMDLDEFLLENGIPSSPTHLGLNQNPLLPVAELEGKELASASAASPPSSSSTVVFQPSEAASGTESSPESERETPSPVDPDCIEIDVNFNPDPADLVLSSVPGGELFNPRKHKFAEEDLKPQPMIKKAKKVFVPDEQKDEKYWTRRKKNNVAAKRSRDARRLKENQITIRAAFLEKENTALRTEVAELRKEVGKCKNIVSKYETRYGPFDLSDSE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9H7</accession>
    <name>A0A452H9H7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">ADP-ribosylation factor-like protein 15</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the small GTPase superfamily. Arf family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9H7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000012874.1">
        <property type="protein sequence ID" value="ENSGAGP00000011241.1"/>
        <property type="gene ID" value="ENSGAGG00000008693.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005525">
        <property type="term" value="F:GTP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003924">
        <property type="term" value="F:GTPase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000934">
        <property type="entry name" value="ADP-ribosylation factor-like 15 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042292">
        <property type="entry name" value="ARL15"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006689">
        <property type="entry name" value="Small_GTPase_ARF/SAR"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46693">
        <property type="entry name" value="ADP-RIBOSYLATION FACTOR-LIKE PROTEIN 15"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46693:SF1">
        <property type="entry name" value="ADP-RIBOSYLATION FACTOR-LIKE PROTEIN 15"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00025">
        <property type="entry name" value="Arf"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00328">
        <property type="entry name" value="SAR1GTPBP"/>
    </dbReference>
    <dbReference type="SMART" id="SM00177">
        <property type="entry name" value="ARF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00178">
        <property type="entry name" value="SAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51417">
        <property type="entry name" value="ARF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 5" id="KW-0342">GTP-binding</keyword>
    <keyword evidence="6" id="KW-0460">Magnesium</keyword>
    <keyword evidence="6" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="2 5" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <feature type="binding site" evidence="5">
        <location>
            <begin position="25"/>
            <end position="32"/>
        </location>
        <ligand>
            <name>GTP</name>
            <dbReference type="ChEBI" id="CHEBI:37565"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="32"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="49"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="5">
        <location>
            <position position="71"/>
        </location>
        <ligand>
            <name>GTP</name>
            <dbReference type="ChEBI" id="CHEBI:37565"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="5">
        <location>
            <begin position="128"/>
            <end position="131"/>
        </location>
        <ligand>
            <name>GTP</name>
            <dbReference type="ChEBI" id="CHEBI:37565"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00010290"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00072404"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PIRSR" id="PIRSR606689-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PIRSR" id="PIRSR606689-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011241.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="190" mass="21239" checksum="21E5B6301E9AFA7E" modified="2019-05-08" version="1">LLCFKALCCKGPPPPRPEYDLVCIGLTGSGKTSLLSQLCSESQENIVSTTGFSIKAVPFQNAVLNVKELGGADTIRKYWSRYYQGSQGVIFVLDSASSEDDLETARNELHSALQHPQLCTLPFLILANHQDKPAARSVQEIKKYFELEPLARGKRWILKPCSLDDMDAVKDSFTLLIHLLEERELEPSRM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9L1</accession>
    <name>A0A452H9L1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Lysosome-associated membrane glycoprotein 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasmic vesicle</location>
            <location evidence="3">Autophagosome membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Endosome membrane</location>
            <topology evidence="2">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="16">Lysosome membrane</location>
            <topology evidence="16">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="16">Belongs to the LAMP family.</text>
    </comment>
    <comment type="caution">
        <text evidence="16">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9L1"/>
    <dbReference type="STRING" id="38772.ENSGAGP00000011280"/>
    <dbReference type="Ensembl" id="ENSGAGT00000012920.1">
        <property type="protein sequence ID" value="ENSGAGP00000011280.1"/>
        <property type="gene ID" value="ENSGAGG00000008724.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031902">
        <property type="term" value="C:late endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005765">
        <property type="term" value="C:lysosomal membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072594">
        <property type="term" value="P:establishment of protein localization to organelle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.160.110:FF:000001">
        <property type="entry name" value="lysosome-associated membrane glycoprotein 2 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.160.110">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048528">
        <property type="entry name" value="Lamp2-like_luminal"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048524">
        <property type="entry name" value="Lamp2-like_TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018134">
        <property type="entry name" value="LAMP_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002000">
        <property type="entry name" value="Lysosome-assoc_membr_glycop"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11506">
        <property type="entry name" value="LYSOSOME-ASSOCIATED MEMBRANE GLYCOPROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11506:SF6">
        <property type="entry name" value="LYSOSOME-ASSOCIATED MEMBRANE GLYCOPROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01299">
        <property type="entry name" value="Lamp2-like_luminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21222">
        <property type="entry name" value="Lamp2_2nd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00336">
        <property type="entry name" value="LYSASSOCTDMP"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00310">
        <property type="entry name" value="LAMP_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51407">
        <property type="entry name" value="LAMP_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0072">Autophagy</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="14" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="11 16" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="7" id="KW-0967">Endosome</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13 16" id="KW-0458">Lysosome</keyword>
    <keyword evidence="10 16 17" id="KW-0472">Membrane</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6 18" id="KW-0732">Signal</keyword>
    <keyword evidence="5 16 17" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 17" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="18">
        <location>
            <begin position="1"/>
            <end position="33"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019408432" description="Lysosome-associated membrane glycoprotein 2" evidence="18">
        <location>
            <begin position="34"/>
            <end position="430"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="395"/>
            <end position="418"/>
        </location>
    </feature>
    <feature type="domain" description="Lysosome-associated membrane glycoprotein 2-like luminal" evidence="19">
        <location>
            <begin position="233"/>
            <end position="377"/>
        </location>
    </feature>
    <feature type="domain" description="Lysosome-associated membrane glycoprotein 2-like transmembrane" evidence="20">
        <location>
            <begin position="397"/>
            <end position="428"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="158"/>
            <end position="194"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="350"/>
            <end position="387"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004652"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023006"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023228"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00074380"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00740"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="Pfam" id="PF01299"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="Pfam" id="PF21222"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011280.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="430" mass="47278" checksum="5A99C98FC9F913AA" modified="2019-05-08" version="1">MATRSPWPRPLSLPCCCLLLLALGGPGFFQSHAVEVDVKDLSNNTCIYAKWMMSFLIKYETNNSEYKNATLNPSTNVTHDGSTCGNNTHAPLLAIQFGAGHSWSINFTKTNETYQGNIISFTYNTSDAVFKDAKTKGPITIVVKDTMPPVQLNTVYKCHHLDSIEVANVSLFFWDVILQAFVQNGTVSKNDSTCEADKSTVAPTTVANLTTAPITTASSPLPTTTSKPVEKPLIGNYSLKDGDKICLLATMGLQLNVSQEKPVLININPKTTNVTGSCGNTSATLRLNDSNRRLIDFTFAVKNTSANTQRFYLKEVNITLISSINGSEPLNAGNNNLSIWDTFLGSSYMCQKEQTVLVTEDLQVHTFDLRIQPFRVQNNKYSKAEECFADSDLNFLIPIAVGMALGFLIILVFISYVIGRRKSRTGYQSV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9L4</accession>
    <name>A0A452H9L4_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="1">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="1 2">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="2" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="1" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="1" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452H9L4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000012913.1">
        <property type="protein sequence ID" value="ENSGAGP00000011273.1"/>
        <property type="gene ID" value="ENSGAGG00000008720.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014848">
        <property type="entry name" value="Rgp1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12507">
        <property type="entry name" value="REDUCED GROWTH PHENOTYPE 1 RGP1, YEAST -RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08737">
        <property type="entry name" value="Rgp1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000313" key="1">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011273.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="387" mass="43055" checksum="412E7D72220971E2" modified="2019-05-08" version="1">MIEVLAQLSRGPVFLAGEMLECGIVFTNPLSATSTSASSEMLAWASAQIHCQFHSSESRVALPPSDDSKHDVQAANETVFVPNRGERGQCILSTPPKILFCDLRLDPGESKSYSYCETLPVDGPPSFRGQSVKYVYKLTIGCQRVNSPIKLLRVPFRVLVLHGLKDYQFPQDEVVAPSNPFLEEEEGVKKDSRLADLATELLMAATSRRSLHVYNISNTRGRVGKFGIFKTVYKIGEDILGTFNFSEGDIPCLQYSVSLQTEESIQEEFQRRHGQPVSYSTHARHQESCLHTARSSFSLPVPLSSTPGFTTNIVSLKWRLHFEFVTSRKSGEAHTVPENQSETITWTGVEQIEVDTFSWDLPIKVLPTNPVLASYVTQFSSTNSITI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9L6</accession>
    <name>A0A452H9L6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Lysosome-associated membrane glycoprotein 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cytoplasmic vesicle</location>
            <location evidence="3">Autophagosome membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Endosome membrane</location>
            <topology evidence="2">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="16">Lysosome membrane</location>
            <topology evidence="16">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="16">Belongs to the LAMP family.</text>
    </comment>
    <comment type="caution">
        <text evidence="16">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9L6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000012925.1">
        <property type="protein sequence ID" value="ENSGAGP00000011285.1"/>
        <property type="gene ID" value="ENSGAGG00000008724.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031902">
        <property type="term" value="C:late endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005765">
        <property type="term" value="C:lysosomal membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072594">
        <property type="term" value="P:establishment of protein localization to organelle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.160.110:FF:000001">
        <property type="entry name" value="lysosome-associated membrane glycoprotein 2 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.160.110">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048528">
        <property type="entry name" value="Lamp2-like_luminal"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048524">
        <property type="entry name" value="Lamp2-like_TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018134">
        <property type="entry name" value="LAMP_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002000">
        <property type="entry name" value="Lysosome-assoc_membr_glycop"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11506">
        <property type="entry name" value="LYSOSOME-ASSOCIATED MEMBRANE GLYCOPROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11506:SF6">
        <property type="entry name" value="LYSOSOME-ASSOCIATED MEMBRANE GLYCOPROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01299">
        <property type="entry name" value="Lamp2-like_luminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21222">
        <property type="entry name" value="Lamp2_2nd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00336">
        <property type="entry name" value="LYSASSOCTDMP"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00310">
        <property type="entry name" value="LAMP_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51407">
        <property type="entry name" value="LAMP_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0072">Autophagy</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="14" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="11 16" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="7" id="KW-0967">Endosome</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13 16" id="KW-0458">Lysosome</keyword>
    <keyword evidence="10 16 17" id="KW-0472">Membrane</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6 18" id="KW-0732">Signal</keyword>
    <keyword evidence="5 16 17" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 17" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="18">
        <location>
            <begin position="1"/>
            <end position="33"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019270101" description="Lysosome-associated membrane glycoprotein 2" evidence="18">
        <location>
            <begin position="34"/>
            <end position="429"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="394"/>
            <end position="417"/>
        </location>
    </feature>
    <feature type="domain" description="Lysosome-associated membrane glycoprotein 2-like luminal" evidence="19">
        <location>
            <begin position="233"/>
            <end position="377"/>
        </location>
    </feature>
    <feature type="domain" description="Lysosome-associated membrane glycoprotein 2-like transmembrane" evidence="20">
        <location>
            <begin position="396"/>
            <end position="427"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="158"/>
            <end position="194"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="16">
        <location>
            <begin position="350"/>
            <end position="387"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004652"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023006"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023228"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00074380"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00740"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="Pfam" id="PF01299"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="Pfam" id="PF21222"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011285.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="429" mass="47002" checksum="60701158A37B02AA" modified="2019-05-08" version="1">MATRSPWPRPLSLPCCCLLLLALGGPGFFQSHAVEVDVKDLSNNTCIYAKWMMSFLIKYETNNSEYKNATLNPSTNVTHDGSTCGNNTHAPLLAIQFGAGHSWSINFTKTNETYQGNIISFTYNTSDAVFKDAKTKGPITIVVKDTMPPVQLNTVYKCHHLDSIEVANVSLFFWDVILQAFVQNGTVSKNDSTCEADKSTVAPTTVANLTTAPITTASSPLPTTTSKPVEKPLIGNYSLKDGDKICLLATMGLQLNVSQEKPVLININPKTTNVTGSCGNTSATLRLNDSNRRLIDFTFAVKNTSANTQRFYLKEVNITLISSINGSEPLNAGNNNLSIWDTFLGSSYMCQKEQTVLVTEDLQVHTFDLRIQPFRVQNNKYSKAQECSLDDDTILIPIIVGAALAGLIVIIVIAYIIGRRKTYAGYQTL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9M0</accession>
    <name>A0A452H9M0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="22">Integrin beta</fullName>
        </recommendedName>
    </protein>
    <organism evidence="27 28">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="28" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="27" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="27" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="20">Heterodimer of an alpha and a beta subunit. Interacts with FLNB. Interacts with HAX1. ITGAV:ITGB6 interacts with FBN1. ITGAV:ITGB6 interacts with TGFB1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell junction</location>
            <location evidence="1">Focal adhesion</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2 22">Cell membrane</location>
            <topology evidence="2 22">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 22">Belongs to the integrin beta chain family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9M0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013114.1">
        <property type="protein sequence ID" value="ENSGAGP00000011455.1"/>
        <property type="gene ID" value="ENSGAGG00000007913.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009986">
        <property type="term" value="C:cell surface"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005925">
        <property type="term" value="C:focal adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034685">
        <property type="term" value="C:integrin alphav-beta6 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033627">
        <property type="term" value="P:cell adhesion mediated by integrin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098609">
        <property type="term" value="P:cell-cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007160">
        <property type="term" value="P:cell-matrix adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007229">
        <property type="term" value="P:integrin-mediated signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000043">
        <property type="entry name" value="Integrin beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000075">
        <property type="entry name" value="Integrin beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000328">
        <property type="entry name" value="Integrin beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1680.10:FF:000002">
        <property type="entry name" value="Integrin beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.410:FF:000002">
        <property type="entry name" value="Integrin beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.1240.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.100">
        <property type="entry name" value="Cytochrome c1, transmembrane anchor, C-terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1680.10">
        <property type="entry name" value="ligand-binding face of the semaphorins, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1510">
        <property type="entry name" value="ntegrin, alpha v. Chain A, domain 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.410">
        <property type="entry name" value="von Willebrand factor, type A domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040622">
        <property type="entry name" value="EGF_integrin_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057073">
        <property type="entry name" value="EGF_integrin_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033760">
        <property type="entry name" value="Integrin_beta_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015812">
        <property type="entry name" value="Integrin_bsu"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014836">
        <property type="entry name" value="Integrin_bsu_cyt_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012896">
        <property type="entry name" value="Integrin_bsu_tail"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036349">
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    <keyword evidence="10" id="KW-0106">Calcium</keyword>
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        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
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    </evidence>
    <sequence length="755" mass="83207" checksum="F040816EEE3E8EB8" modified="2019-05-08" version="1">VTCEDCLLSGPQCAWCSQENYTDSSGVHGRCDAPENLLSKGCQLKFIEFPISKVEIHKNKPLSIGIQNNSDVTQISPQKLTFRLRPGNEETIEIKVRQTEDYPVDLYYLMDLSASMDDDLNTIKELGSTLSKEMSKLTSNFRLGFGSFVEKPVSPFIKTTAEEINNPCRSVPRDCLPTFGYKHVLPLTNDAERFNEIVKEQKISANIDTPEGGFDAIMQAAVCKEKIGWRNDSLHLLVFVSDADSHFGMDSKLAGIVIPNDGHCHLDQNNEYSMSTVLEYPTIGQLTDKLVQNNVLLIFAVTKEQVYFYENYAKLIPGATVGQLQKDSGNILQLIIDAYKELRSEVELEVLGDTEGLNLSFTAICNNGSIFPHQRKCSHIKVGETVSFNMTVSLPTCEKRRRQIVIKPVGLSDALEIDIQLECSCNCQKEAEMNSSKCSKGKGSFECGVCACNPGYMGPYCECNENSLSTNSCKRSPEQSSCSGRGDCYCGQCICHFSVYGNIYGLYCECDNFSCVRFKGLQCGGNGDCDCGECVCHRGWTGEYCNCTTDTNSCVSEDGILCSGRGECICGTCVCTNPGASGSTCERCPTCGDPCNSKRSCIECHLASDVQSQAECNEKCRLVDATVNSAEDFSKDKSIPCSLQGENECVTTFLMATDEQGRTIIHSIKQKDCPQPPNIPMIMLGVSLAILLIGIVLLCIWKLLVSFQDRKEVAKFEAERSKAKWQTVCICINFIQVSHISVSMLKAEMPSEGNF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9M1</accession>
    <name>A0A452H9M1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Beta/gamma crystallin 'Greek key' domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">Crystallins are the dominant structural components of the vertebrate eye lens.</text>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the beta/gamma-crystallin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9M1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013126.1">
        <property type="protein sequence ID" value="ENSGAGP00000011465.1"/>
        <property type="gene ID" value="ENSGAGG00000008842.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005212">
        <property type="term" value="F:structural constituent of eye lens"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002088">
        <property type="term" value="P:lens development in camera-type eye"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007601">
        <property type="term" value="P:visual perception"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="entry name" value="Crystallin gamma S"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.20.10:FF:000003">
        <property type="entry name" value="Crystallin gamma S"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Crystallins"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050252">
        <property type="entry name" value="Beta/Gamma-Crystallin"/>
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    <dbReference type="InterPro" id="IPR001064">
        <property type="entry name" value="Beta/gamma_crystallin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038776">
        <property type="entry name" value="C2orf80"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011024">
        <property type="entry name" value="G_crystallin-like"/>
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    <dbReference type="PANTHER" id="PTHR11818">
        <property type="entry name" value="BETA/GAMMA CRYSTALLIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11818:SF119">
        <property type="entry name" value="GAMMA-CRYSTALLIN D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00030">
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        <property type="match status" value="2"/>
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        <property type="match status" value="1"/>
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        <property type="entry name" value="BGCRYSTALLIN"/>
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        <property type="match status" value="2"/>
    </dbReference>
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        <property type="entry name" value="CRYSTALLIN_BETA_GAMMA"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
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    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="Beta/gamma crystallin 'Greek key'" evidence="6">
        <location>
            <begin position="80"/>
            <end position="118"/>
        </location>
    </feature>
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            <begin position="119"/>
            <end position="160"/>
        </location>
    </feature>
    <feature type="domain" description="Beta/gamma crystallin 'Greek key'" evidence="6">
        <location>
            <begin position="207"/>
            <end position="249"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="50"/>
            <end position="79"/>
        </location>
    </feature>
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        <source>
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        <source>
            <dbReference type="ARBA" id="ARBA00009646"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="PROSITE" id="PS50915"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011465.1"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="252" mass="29081" checksum="65C62593791D437C" modified="2019-05-08" version="1">MGPSATRWYRSAKGHRIHPFNLSLHPFAMLTAPKAAEHAWKQSVKFQKAAANQHATTNSTRTNKDCKRSESPSTAKQPDNKIILFEDRNFQGRSVECSSDRPDLQSHLSRCNSVRVESGCFMLYEHPNFQGQQFSLKWGDYPDMQSEGFSTSIKSCRMIPSHRGTYRIKIYDKEDHRGNMVELTEDSPRVMDQLRSPEMLSCSVLEGYWILYELPNYRGRQYLLRPGQYRIFSEWGAMSGKVGSLRRAPDLY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9M2</accession>
    <name>A0A452H9M2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Cleavage and polyadenylation specificity factor subunit 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
            <location evidence="1">Nucleoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the CPSF1 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000013193.1">
        <property type="protein sequence ID" value="ENSGAGP00000011521.1"/>
        <property type="gene ID" value="ENSGAGG00000004276.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005654">
        <property type="term" value="C:nucleoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003676">
        <property type="term" value="F:nucleic acid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031123">
        <property type="term" value="P:RNA 3'-end processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.910:FF:000005">
        <property type="entry name" value="Cleavage and polyadenylation specific factor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:002223">
        <property type="entry name" value="Cleavage and polyadenylation specific factor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:000100">
        <property type="entry name" value="Cleavage and polyadenylation specificity factor subunit 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:000118">
        <property type="entry name" value="Cleavage and polyadenylation specificity factor subunit 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:000544">
        <property type="entry name" value="cleavage and polyadenylation specificity factor subunit 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.910">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018846">
        <property type="entry name" value="Beta-prop_RSE1/DDB1/CPSF1_1st"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058543">
        <property type="entry name" value="Beta-prop_RSE1/DDB1/CPSF1_2nd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050358">
        <property type="entry name" value="RSE1/DDB1/CFT1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004871">
        <property type="entry name" value="RSE1/DDB1/CPSF1_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10644">
        <property type="entry name" value="DNA REPAIR/RNA PROCESSING CPSF FAMILY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10433">
        <property type="entry name" value="Beta-prop_RSE1_1st"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23726">
        <property type="entry name" value="Beta-prop_RSE1_2nd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03178">
        <property type="entry name" value="CPSF_A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0539">Nucleus</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="RSE1/DDB1/CPSF1 first beta-propeller" evidence="7">
        <location>
            <begin position="14"/>
            <end position="400"/>
        </location>
    </feature>
    <feature type="domain" description="RSE1/DDB1/CPSF1 second beta-propeller" evidence="8">
        <location>
            <begin position="521"/>
            <end position="1003"/>
        </location>
    </feature>
    <feature type="domain" description="RSE1/DDB1/CPSF1 C-terminal" evidence="6">
        <location>
            <begin position="1076"/>
            <end position="1396"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="404"/>
            <end position="438"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="543"/>
            <end position="570"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="714"/>
            <end position="776"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="410"/>
            <end position="420"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="543"/>
            <end position="560"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="761"/>
            <end position="776"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004642"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00038446"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00068483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="Pfam" id="PF03178"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF10433"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF23726"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011521.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1431" mass="159303" checksum="77A67D159DD5A1D8" modified="2019-05-08" version="1">MYAVYRQAHPPTGVEFSMYCNFFSNAERNLVVAGTSQLYVYRLNHDSEPSTKGDRNTEGKGHKEKLELVASFSFFGNVMSMASVQLAGAKRDALLLSFKDAKLSVVEYDPGTHDLKTLSLHYFEEPELKDGFVQNVHIPKVRVDPDGRCAVMLIYGTRLVVLPFRRETLTDEHEGVMGEGQKSSFLPSYIIDVRELDEKLLNIIDMQFLYGYYEPTLLILFEPNQTWPGRVAVRQDTCSIVAISLNIMQKVHPVIWSLSNLPFDCTQALAVPKPIGGVVIFAVNSLLYLNQSVPPYGVSLNSLTTGTTVFPLRIQDGVKITLDCAQAAFISYDKMVISLKGGEIYVLTLITDGMRSVRSFHFDKAAASVLTTCMITMEPGYLFLGSRLGNSLLLKYTEKLQEPPVNVAKDPADKQEEPPVKKKRSESSGNWAGGKSAPQDEVDEIEVYGSEAQSGTQLATYSFEVCDSILNIGPCANAAMGEPAFLSEEFQNSPEPDLEIVVCSGYGKNGALSVLQKSIRPQVVTTFELPGCYDMWTVISPQKKEEDESAKGESAEKESSPPEPEDDGKRHGFLILSREDSTMILQTGQEIMELDTSGFATQGPTVFAGNIGENRYIVQVSPLGIRLLEGVNQLHFIPVDLGSPIVHCAVADPYVVIMSAEGQVTMFVLKSDTYGGRTHRLTLQKPQLHHQSKVIALCVYRDVRGMFTTESKTASSRDEFSLRRQGGPLPASPAPAARHSNTVDDEEEMLYGDSSTLFSPSKEEPRRSSLPLADRDPHQYKVEPTHWCVLVRENGAMEIYQLPDWRLVFLVKNFPMGQRVLVDSSFGQPAAQGDAKKEEVTRQGEMPLVKEVLLVALGNRQSRPYLLVHVDQELLIYEAFSHDSQLGQSNLKVRFKKVPHNINFREKKLKLSKKKSESAGGEESGGARGRVARFRYFEDIYGYSGVFICGPSPHWLLVTARGALRLHPMTIDGLIESFAPFHNVNCPKGFLYFNRQGELRISVLPAYLSYDAPWPVRKIPLRCTTHYVAYHVESKVYAVATSVVNPCTRIPRMTGEEKEFESIERDERYIHPQQEAFSIQLISPVSWETIPNTRIELEEWEHVTCMKTVSLKSEETVSGLKGYIAAGTCLMQGEEVTCRGRILIMDIIEVVPEPGQPLTKNKFKVLYEKEQKGPVTALCHCHGYLVSAIGQKIFLWSLKDNDLTGMAFIDTQLYIHQMISVKNFILAADVMKSISLLRYQEESKTLSLVSRVAQPGTIVYSVDFMVDTDQLGFLGRFCSFVPCTLAPTASGGGSGQARGARARTPWPCCGDPAACTLLPAPGQSCLATLDGGIGLLLPMLEKTYRRLLMLQNALTTMLSHHAGLNPRAFRMLHVDRRILQNAVRNILDGELLSRYLYLSAMERSELAKKIGTTPDIILEDLLEIDRVTAHF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9M7</accession>
    <name>A0A452H9M7_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="15">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the plexin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9M7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013131.1">
        <property type="protein sequence ID" value="ENSGAGP00000011470.1"/>
        <property type="gene ID" value="ENSGAGG00000008707.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002116">
        <property type="term" value="C:semaphorin receptor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017154">
        <property type="term" value="F:semaphorin receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007162">
        <property type="term" value="P:negative regulation of cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050772">
        <property type="term" value="P:positive regulation of axonogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030334">
        <property type="term" value="P:regulation of cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008360">
        <property type="term" value="P:regulation of cell shape"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd12792">
        <property type="entry name" value="RasGAP_plexin_B2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000203">
        <property type="entry name" value="Plexin B2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000798">
        <property type="entry name" value="Plexin B2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.506.10:FF:000035">
        <property type="entry name" value="Plexin b2a"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.506.10">
        <property type="entry name" value="GTPase Activation - p120gap, domain 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014756">
        <property type="entry name" value="Ig_E-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002909">
        <property type="entry name" value="IPT_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031148">
        <property type="entry name" value="Plexin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013548">
        <property type="entry name" value="Plexin_cytoplasmic_RasGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046800">
        <property type="entry name" value="Plexin_RBD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016201">
        <property type="entry name" value="PSI"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057533">
        <property type="entry name" value="PSI_Plexin-B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041019">
        <property type="entry name" value="TIG1_plexin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22625">
        <property type="entry name" value="PLEXIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22625:SF9">
        <property type="entry name" value="PLEXIN-B2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08337">
        <property type="entry name" value="Plexin_cytopl"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20170">
        <property type="entry name" value="Plexin_RBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24317">
        <property type="entry name" value="PSI_Plexin-B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01833">
        <property type="entry name" value="TIG"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17960">
        <property type="entry name" value="TIG_plexin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00429">
        <property type="entry name" value="IPT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00423">
        <property type="entry name" value="PSI"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81296">
        <property type="entry name" value="E set domains"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="12" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="9" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="8" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0675">Receptor</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <keyword evidence="4" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7" id="KW-1133">Transmembrane helix</keyword>
    <feature type="domain" description="PSI" evidence="13">
        <location>
            <begin position="108"/>
            <end position="158"/>
        </location>
    </feature>
    <feature type="domain" description="PSI" evidence="13">
        <location>
            <begin position="204"/>
            <end position="250"/>
        </location>
    </feature>
    <feature type="domain" description="IPT/TIG" evidence="14">
        <location>
            <begin position="251"/>
            <end position="344"/>
        </location>
    </feature>
    <feature type="domain" description="IPT/TIG" evidence="14">
        <location>
            <begin position="345"/>
            <end position="431"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="12">
        <location>
            <begin position="620"/>
            <end position="647"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010297"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="SMART" id="SM00423"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="SMART" id="SM00429"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011470.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1230" mass="139696" checksum="EC8C3508E1FE0890" modified="2019-05-08" version="1">MPNHWLWSPDGVCMAITSAHPQNMSRKAPGKVELTVSPLPELTESDMLWCQFGETTNIAELRDGTIICDPPTTIPHTPKGQDHVSVSLILTFGEEKMFFASATYPFYDCEEAMNLSENLPCSSCASSRWNCQWDWKKHFCEDSSLEEDVITQNMEDCPQFLNPNPQIIPMEFPTIVHFEGKNLENYQVNELIPSSHLTYMTLYNCSYGRSDCSLCLAADPGYDCVWCEEEGKPNKCIYKNLCKTSTNTCPHPVITRIEPKTGPLSGGILLTIMGSNLGIKAEDVKSITVAGTECLFKEDFYSVSIRYLIVCQAGNVSVPSQGKIEVDINGKRGSSTSEVQFTYQQPHPTGISPTEGPRAGGTILTITGNNLETGSQKDVRVLLGDVPCNITDFGEQIVCITGQHESGGKVQVTVDHGDTENQLNDPWFTYSENPTVTKFFPKNSFNITKINESTIVFSSPPILEDPENYRITTLIKMDGCSLVLNAENNDPFKYVADPTFENFTDGIKKQVNKLIHAKGSNLNKAMTIEEAQAFVGAEPCNIKTLTEKDLYCEPPEVQPQPKKRQKRDTINNLPEFIVKFGLREWILGRVEYDTRVSDVPLDVILPSVIFPMVAIIIISIYCYRRKSQQAEREYEKIKSQLEGLEESVRDRCKKEFTDLMIEMEDQTNDVSEAGIPVLDYKMYTDRVFFLPSKDGEKDVMITGKLDIPEARKQTVEQALNQFSNLLNSKSFLINFIHTLEKQREFSAREKVYFASLLTVALHGKLEYYTDIMRTLFLELMEQYVVAKNPKLMLRRSETVVERMLSNWMAICLYQYLKDNAGEPLYKLFKAIKHQVEKGPVDAVLMKAKYTLNDTGLLGDDVEYTQLTVNVYVQDGGTDSVPVKVLNCDTISQVKEKIIDQVYRNLPCSQWPKADSVVLVVFKPASNSHVLWDDFSIFVFLKVRDGATLILSKIGISQQPEDNQQDVPGERHALLEDENKVWHLVRPADEVDEGKSKRGSMKEKERTKAITEIYLTRLLSVKGTLQQFVDNFFHSVLNSNHVVPPAVKYFFDFLDEQAEKHELKDEDTIHIWKTNSLPLRFWVNILKNPHFIFDVHVHEVVDASLSVIAQTFMDACTRTEHKLSRDSPSNKLLYAKEISNYKKMVEDYYKGIRQMVQVSDQDMNTHLAEISRAHTDSLNTLVALHQLYQYTNKYYDEIINALEEDPAAQKMQLAFRLQQIAAALENKVTDL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9N2</accession>
    <name>A0A452H9N2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">BTB domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452H9N2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013137.1">
        <property type="protein sequence ID" value="ENSGAGP00000011475.1"/>
        <property type="gene ID" value="ENSGAGG00000008844.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="CDD" id="cd18501">
        <property type="entry name" value="BACK_ANKFY1_Rank5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18303">
        <property type="entry name" value="BTB_POZ_Rank-5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000116">
        <property type="entry name" value="Ankyrin repeat and FYVE domain containing 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000142">
        <property type="entry name" value="Ankyrin repeat and FYVE domain containing 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000210">
        <property type="entry name" value="Ankyrin repeat and FYVE domain containing 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.710.10">
        <property type="entry name" value="Potassium Channel Kv1.1, Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049765">
        <property type="entry name" value="ANFY1_BTB_POZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049763">
        <property type="entry name" value="ANKFY1_BACK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000210">
        <property type="entry name" value="BTB/POZ_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011333">
        <property type="entry name" value="SKP1/BTB/POZ_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24198">
        <property type="entry name" value="ANKYRIN REPEAT AND PROTEIN KINASE DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24198:SF191">
        <property type="entry name" value="RABANKYRIN-5-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00023">
        <property type="entry name" value="Ank"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00651">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01415">
        <property type="entry name" value="ANKYRIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="21"/>
    </dbReference>
    <dbReference type="SMART" id="SM00225">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54695">
        <property type="entry name" value="POZ domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50097">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2 3" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="BTB" evidence="4">
        <location>
            <begin position="55"/>
            <end position="117"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="3">
        <location>
            <begin position="275"/>
            <end position="307"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="3">
        <location>
            <begin position="477"/>
            <end position="509"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="3">
        <location>
            <begin position="528"/>
            <end position="560"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="3">
        <location>
            <begin position="607"/>
            <end position="639"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="3">
        <location>
            <begin position="640"/>
            <end position="672"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="3">
        <location>
            <begin position="755"/>
            <end position="787"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="3">
        <location>
            <begin position="856"/>
            <end position="888"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="3">
        <location>
            <begin position="891"/>
            <end position="923"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="3">
        <location>
            <begin position="924"/>
            <end position="956"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="3">
        <location>
            <begin position="957"/>
            <end position="978"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="3">
        <location>
            <begin position="1029"/>
            <end position="1061"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50097"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011475.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1112" mass="122938" checksum="987228253E9EB7E5" modified="2019-05-08" version="1">MLLRQEYVKLQKKLADTERRCTLLAAQSNKENASDSFISRLLTIVAELYQQEQYSDLKIKVGDKHIRAHKFVLAARSETWSLANLVSTEELDLSDAEPEVTMAMLRWIYTDELELREDDVFLTELMKLANRFQLQLLRERCEKGVMSLVNVRNCIRFYQTAEELNASTLLNYCAEIIASHWDDLRKEDFSSMSAQLLYKMFKSKTEYPLHKAIKVEREDVVFLYLIEMDSQLPGKLNELDHNGDLALDLALSRRLESIATTLVNYKADVDMVDKRGWSLLHKAIRRGDKFSANFLIKNGARVNAATLGDQETPLHLVASYSPKKHLPDVMSEMAQIAESLLQAGANPNMQDCKGRTPLHASIVVRNDPVFSQLLQCKQLDLELKDHEGSTALWLAVQYIKVSSDLSINPFDDVPVVNGTSFDENSFAARLIQRGSNTDAPDTVTGNCLLQRAAGAGNEAAALFLATHGAKVNHRNKWGETPLHTSCRHGLSNLTAELLQQGANPNIQTEEAVPSQKDASSPPASGNVYLQTPLHMAIAYNHPDVVSVILEQKANALHATNNLQIIPDFSLKDSRDQTVLGLALWTGMHTIAAQLLGSGASINDTMSDGQTLLHMAIQRQDSKSALFLLEHQADINVRTQDGETALQLAIKNQLPLVVDAICTRGADMSVPDEKGNPPLWLALENNLEDIASTLVRHGCDATCWGPGPGGCLQTLLHRSIDENSEQIACFLIRSGCDVNSPRQPGANGEGEEEARDGQTPLHLAACWGLEEVVQCLLEFGANVNAQDAEGRTPIHVAISNQHNVIIQLMISHPEIRLNMRDRQGMTPFACAMTYKNNKAAEAILKREAGAAEQVDNKGRNFLHIAVQNSDIESVLFLISVQANVNSRVQDASKLTPLHLAVQAGSEIIVRNLLLAGAKVNELTKHRQTALHLAAQQDLPIICSVLLENGVDFAAVDENGNNALHLAVMHGRLNNIRILLTECNVDAEAFNLRGQSPMHILGHYGKDNAAAICELFLECMPEYPLDKPDAEGNTVLLLAYMKGNANLCRAIVRAGARLGVNNNQGVNIFNYQVATKQLLFRLLDMLSKEPPWCDGSNCYECTAKFGVTTRKHHW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9N3</accession>
    <name>A0A452H9N3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">G-protein coupled receptors family 1 profile domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9N3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000012918.1">
        <property type="protein sequence ID" value="ENSGAGP00000011278.1"/>
        <property type="gene ID" value="ENSGAGG00000008716.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000012928.1">
        <property type="protein sequence ID" value="ENSGAGP00000011288.1"/>
        <property type="gene ID" value="ENSGAGG00000008716.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000012935.1">
        <property type="protein sequence ID" value="ENSGAGP00000011294.1"/>
        <property type="gene ID" value="ENSGAGG00000008716.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008528">
        <property type="term" value="F:G protein-coupled peptide receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016500">
        <property type="term" value="F:protein-hormone receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007189">
        <property type="term" value="P:adenylate cyclase-activating G protein-coupled receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090263">
        <property type="term" value="P:positive regulation of canonical Wnt signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd15363">
        <property type="entry name" value="7tmA_LGR5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:001164">
        <property type="entry name" value="GH01279p"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1070.10:FF:000028">
        <property type="entry name" value="leucine-rich repeat-containing G-protein coupled receptor 4 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000770">
        <property type="entry name" value="Uncharacterized protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1070.10">
        <property type="entry name" value="Rhodopsin 7-helix transmembrane proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000276">
        <property type="entry name" value="GPCR_Rhodpsn"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017452">
        <property type="entry name" value="GPCR_Rhodpsn_7TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002131">
        <property type="entry name" value="Gphrmn_rcpt_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001611">
        <property type="entry name" value="Leu-rich_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003591">
        <property type="entry name" value="Leu-rich_rpt_typical-subtyp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026906">
        <property type="entry name" value="LRR_5"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24372">
        <property type="entry name" value="GLYCOPROTEIN HORMONE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24372:SF71">
        <property type="entry name" value="LEUCINE-RICH REPEAT-CONTAINING G-PROTEIN COUPLED RECEPTOR 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00001">
        <property type="entry name" value="7tm_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13306">
        <property type="entry name" value="LRR_5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13855">
        <property type="entry name" value="LRR_8"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00373">
        <property type="entry name" value="GLYCHORMONER"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00237">
        <property type="entry name" value="GPCRRHODOPSN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00364">
        <property type="entry name" value="LRR_BAC"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00365">
        <property type="entry name" value="LRR_SD22"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00369">
        <property type="entry name" value="LRR_TYP"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81321">
        <property type="entry name" value="Family A G protein-coupled receptor-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52058">
        <property type="entry name" value="L domain-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50262">
        <property type="entry name" value="G_PROTEIN_RECEP_F1_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51450">
        <property type="entry name" value="LRR"/>
        <property type="match status" value="5"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="10" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="8" id="KW-0297">G-protein coupled receptor</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="3" id="KW-0433">Leucine-rich repeat</keyword>
    <keyword evidence="9 14" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-0675">Receptor</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <keyword evidence="13" id="KW-0807">Transducer</keyword>
    <keyword evidence="4 14" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 14" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="484"/>
            <end position="509"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="521"/>
            <end position="545"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="565"/>
            <end position="586"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="607"/>
            <end position="628"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="648"/>
            <end position="673"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="694"/>
            <end position="717"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="729"/>
            <end position="749"/>
        </location>
    </feature>
    <feature type="domain" description="G-protein coupled receptors family 1 profile" evidence="15">
        <location>
            <begin position="500"/>
            <end position="746"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022614"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023224"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50262"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011294.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="834" mass="92496" checksum="EEE1BD3C67A00550" modified="2019-05-08" version="1">MNNITKLPSNPLHNLRFLEELRLAGNGLTYIPRGAFAGLFSLKVLMLQNNQLKQVPTEALQNLRSLQSLRLDANHINYVPPNCFSGLVSLRHLWLDDNSLTEIPVQAFRSLSALQAMTLALNKIHHIPDYAFGNLSSLVVLHLHNNRIYSLGKKCFDGLHNLETLDLNYNSLDEFPIAIRTLTNLKELGFHSNNIKSIPEHAFVGNPSLITIHFYDNPIQLVGQSAFQHLPELRTLTLNSASQITEFPDLTGTTSLESLTLTGAQVTSLPKVACDQLPNLQVLDLSYNLLQDLPCFTACKKLQKIDLHHNEIYEIKANTFQQLVALRSLDLAWNKIAVIHPSAFSSLPSLIKLDLSSNLLSSFPVTGLHGLTHLKLTGNHDLQSLISSENFPELKVIEMPYAYQCCAFGTCESHYKISSQWNKEENSSADDFHKKDSGMLHIQDEHDFEDFLLDFEEDLKAHHSVQCTPSPGPFKPCDHLFGSWLIRIGVWTIVGLAFICNALLTATVFRSPLYISSIKLLLGLIAIVNMLMGLSSGVLASMDAFTFGSFAQYGAWWESGVGCQVTGLLSIFASETSIFLLTLAALERGFSVKYATKCKTKSSFASVKIAIFFCFMLALIIAVVPLLSESKYGVSPLCLPLPFGEPSSMGFMVALVLLNSLCFLVMTVAYTKLYCSLEKGELDNIWDCSMIKHIALLLFTNCILYCPVAFLSFSSLLNLTFISPEVIKSILLVIVPLPACLNPLLYILFNPHFKEDLGSLRKQTLLWSRSKHTSLISVNSDDVEKQSCDSTQALVTFASTSISYDVPASNSLMPSSYQTTESCNLSPVAFVPCR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9N4</accession>
    <name>A0A452H9N4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">FYVE-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452H9N4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013132.1">
        <property type="protein sequence ID" value="ENSGAGP00000011471.1"/>
        <property type="gene ID" value="ENSGAGG00000008844.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd18501">
        <property type="entry name" value="BACK_ANKFY1_Rank5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd15728">
        <property type="entry name" value="FYVE_ANFY1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000116">
        <property type="entry name" value="Ankyrin repeat and FYVE domain containing 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000142">
        <property type="entry name" value="Ankyrin repeat and FYVE domain containing 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000210">
        <property type="entry name" value="Ankyrin repeat and FYVE domain containing 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.40.10:FF:000104">
        <property type="entry name" value="Ankyrin repeat and FYVE domain-containing 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.710.10">
        <property type="entry name" value="Potassium Channel Kv1.1, Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049764">
        <property type="entry name" value="ANFY1_FYVE"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049763">
        <property type="entry name" value="ANKFY1_BACK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011333">
        <property type="entry name" value="SKP1/BTB/POZ_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000306">
        <property type="entry name" value="Znf_FYVE"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017455">
        <property type="entry name" value="Znf_FYVE-rel"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011011">
        <property type="entry name" value="Znf_FYVE_PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24198">
        <property type="entry name" value="ANKYRIN REPEAT AND PROTEIN KINASE DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24198:SF191">
        <property type="entry name" value="RABANKYRIN-5-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00023">
        <property type="entry name" value="Ank"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01363">
        <property type="entry name" value="FYVE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01415">
        <property type="entry name" value="ANKYRIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="21"/>
    </dbReference>
    <dbReference type="SMART" id="SM00064">
        <property type="entry name" value="FYVE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57903">
        <property type="entry name" value="FYVE/PHD zinc finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50178">
        <property type="entry name" value="ZF_FYVE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5 6" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="1" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0862">Zinc</keyword>
    <keyword evidence="3 7" id="KW-0863">Zinc-finger</keyword>
    <feature type="repeat" description="ANK" evidence="6">
        <location>
            <begin position="174"/>
            <end position="206"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="6">
        <location>
            <begin position="376"/>
            <end position="408"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="6">
        <location>
            <begin position="427"/>
            <end position="459"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="6">
        <location>
            <begin position="506"/>
            <end position="538"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="6">
        <location>
            <begin position="539"/>
            <end position="571"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="6">
        <location>
            <begin position="654"/>
            <end position="686"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="6">
        <location>
            <begin position="755"/>
            <end position="787"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="6">
        <location>
            <begin position="790"/>
            <end position="822"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="6">
        <location>
            <begin position="823"/>
            <end position="855"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="6">
        <location>
            <begin position="856"/>
            <end position="877"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="6">
        <location>
            <begin position="928"/>
            <end position="960"/>
        </location>
    </feature>
    <feature type="domain" description="FYVE-type" evidence="8">
        <location>
            <begin position="989"/>
            <end position="1049"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00091"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50178"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011471.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1054" mass="116131" checksum="F46CB921088FE274" modified="2019-05-08" version="1">MAMLRWIYTDELELREDDVFLTELMKLANRFQLQLLRERCEKGVMSLVNVRNCIRFYQTAEELNASTLLNYCAEIIASHWDDLRKEDFSSMSAQLLYKMFKSKTEYPLHKAIKVEREDVVFLYLIEMDSQLPGKLNELDHNGDLALDLALSRRLESIATTLVNYKADVDMVDKRGWSLLHKAIRRGDKFSANFLIKNGARVNAATLGDQETPLHLVASYSPKKHLPDVMSEMAQIAESLLQAGANPNMQDCKGRTPLHASIVVRNDPVFSQLLQCKQLDLELKDHEGSTALWLAVQYIKVSSDLSINPFDDVPVVNGTSFDENSFAARLIQRGSNTDAPDTVTGNCLLQRAAGAGNEAAALFLATHGAKVNHRNKWGETPLHTSCRHGLSNLTAELLQQGANPNIQTEEAVPSQKDASSPPASGNVYLQTPLHMAIAYNHPDVVSVILEQKANALHATNNLQIIPDFSLKDSRDQTVLGLALWTGMHTIAAQLLGSGASINDTMSDGQTLLHMAIQRQDSKSALFLLEHQADINVRTQDGETALQLAIKNQLPLVVDAICTRGADMSVPDEKGNPPLWLALENNLEDIASTLVRHGCDATCWGPGPGGCLQTLLHRSIDENSEQIACFLIRSGCDVNSPRQPGANGEGEEEARDGQTPLHLAACWGLEEVVQCLLEFGANVNAQDAEGRTPIHVAISNQHNVIIQLMISHPEIRLNMRDRQGMTPFACAMTYKNNKAAEAILKREAGAAEQVDNKGRNFLHIAVQNSDIESVLFLISVQANVNSRVQDASKLTPLHLAVQAGSEIIVRNLLLAGAKVNELTKHRQTALHLAAQQDLPIICSVLLENGVDFAAVDENGNNALHLAVMHGRLNNIRILLTECNVDAEAFNLRGQSPMHILGHYGKDNAAAICELFLECMPEYPLDKPDAEGNTVLLLAYMKGNANLCRAIVRAGARLGVNNNQGVNIFNYQVATKQLLFRLLDMLSKEPPWCDGSNCYECTAKFGVTTRKHHCRHCGRLLCHKCSTKEIPIIKFDLNKPVRVCNICFDVLTLGGVS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9N8</accession>
    <name>A0A452H9N8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">Ig-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="12">Synapse</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="13">Belongs to the LRFN family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9N8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000012941.1">
        <property type="protein sequence ID" value="ENSGAGP00000011300.1"/>
        <property type="gene ID" value="ENSGAGG00000008741.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045202">
        <property type="term" value="C:synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000235">
        <property type="entry name" value="Leucine-rich repeat and fibronectin type III domain-containing 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000091">
        <property type="entry name" value="Leucine-rich repeat and fibronectin type III domain-containing protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000016">
        <property type="entry name" value="Leucine-rich repeat and fibronectin type III domain-containing protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000019">
        <property type="entry name" value="leucine-rich repeat and fibronectin type III domain-containing protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000483">
        <property type="entry name" value="Cys-rich_flank_reg_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013098">
        <property type="entry name" value="Ig_I-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001611">
        <property type="entry name" value="Leu-rich_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003591">
        <property type="entry name" value="Leu-rich_rpt_typical-subtyp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050467">
        <property type="entry name" value="LRFN"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45842:SF7">
        <property type="entry name" value="LEUCINE-RICH REPEAT AND FIBRONECTIN TYPE III DOMAIN-CONTAINING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45842">
        <property type="entry name" value="SYNAPTIC ADHESION-LIKE MOLECULE SALM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07679">
        <property type="entry name" value="I-set"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00560">
        <property type="entry name" value="LRR_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13855">
        <property type="entry name" value="LRR_8"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00369">
        <property type="entry name" value="LRR_TYP"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SMART" id="SM00082">
        <property type="entry name" value="LRRCT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52058">
        <property type="entry name" value="L domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51450">
        <property type="entry name" value="LRR"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="2" id="KW-0433">Leucine-rich repeat</keyword>
    <keyword evidence="8 15" id="KW-0472">Membrane</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="4 16" id="KW-0732">Signal</keyword>
    <keyword evidence="7" id="KW-0770">Synapse</keyword>
    <keyword evidence="3 15" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 15" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="16">
        <location>
            <begin position="1"/>
            <end position="17"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019056041" description="Ig-like domain-containing protein" evidence="16">
        <location>
            <begin position="18"/>
            <end position="597"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="513"/>
            <end position="539"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="17">
        <location>
            <begin position="287"/>
            <end position="375"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="548"/>
            <end position="597"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="557"/>
            <end position="570"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022614"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00034103"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00038433"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011300.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="597" mass="65630" checksum="FDD8729DAD4EE0E3" modified="2019-05-08" version="1">MEKLLLPLLMLGTMAKGQHCPGRCICQNISPTLTMLCAKTGLLFVPPSIDRRTVELRLTDNFITIIRRKDFSNMTNLVHLTLSRNTISQVMPYAFADLRALRALHMNSNRLASLRNEHFKGLGNLRHLILGNNQISNIQPASFDEFLATVEDLDLSYNNLETLPWEAIGQMVSLNTLTLDHNLIDHIAEGTFSQLHKLVRLDMTSNRLQKLPPDNLFLRAQVLANVRGSHPSTLTISFGGNPLHCNCELLWLRRLTREDDLETCASPEHLMDKYFWSVSEEEFICEPPLITRQYSTKAFIMEGQGVSLKCKAVGDPDPSIHWIAPDGKLIHNTTRATVYDNGTLEVHITTLKDNGVFTCIASNAAGEATAPVEVCIVPLPLLVNNTGHIKEPDPGLSDITTSTKSGANDTKNHLEKKIVAAELTSTSVVIHWPSERHIPGIRMYQIQYNSSLDDSLVYRMIPPSSRAFLVNDLAAGRAYDLCVLAVYDDGVTALTATRVVGCTTQCHALHTQFLGGTMIIIIGGIIVASVLVFIIILMIRYKVHSGHEGPKKAKVSNVYSQTNGDGSGSPTRPRRGSCADPTDAGSGALRSGEGGQK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9N9</accession>
    <name>A0A452H9N9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Beta/gamma crystallin 'Greek key' domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">Crystallins are the dominant structural components of the vertebrate eye lens.</text>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the beta/gamma-crystallin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9N9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013138.1">
        <property type="protein sequence ID" value="ENSGAGP00000011476.1"/>
        <property type="gene ID" value="ENSGAGG00000008850.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005212">
        <property type="term" value="F:structural constituent of eye lens"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002088">
        <property type="term" value="P:lens development in camera-type eye"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007601">
        <property type="term" value="P:visual perception"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.20.10:FF:000001">
        <property type="entry name" value="Crystallin gamma S"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.20.10:FF:000003">
        <property type="entry name" value="Crystallin gamma S"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.20.10">
        <property type="entry name" value="Crystallins"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050252">
        <property type="entry name" value="Beta/Gamma-Crystallin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001064">
        <property type="entry name" value="Beta/gamma_crystallin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011024">
        <property type="entry name" value="G_crystallin-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11818">
        <property type="entry name" value="BETA/GAMMA CRYSTALLIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11818:SF119">
        <property type="entry name" value="GAMMA-CRYSTALLIN D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00030">
        <property type="entry name" value="Crystall"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01367">
        <property type="entry name" value="BGCRYSTALLIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00247">
        <property type="entry name" value="XTALbg"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49695">
        <property type="entry name" value="gamma-Crystallin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50915">
        <property type="entry name" value="CRYSTALLIN_BETA_GAMMA"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0273">Eye lens protein</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="Beta/gamma crystallin 'Greek key'" evidence="5">
        <location>
            <begin position="5"/>
            <end position="43"/>
        </location>
    </feature>
    <feature type="domain" description="Beta/gamma crystallin 'Greek key'" evidence="5">
        <location>
            <begin position="44"/>
            <end position="85"/>
        </location>
    </feature>
    <feature type="domain" description="Beta/gamma crystallin 'Greek key'" evidence="5">
        <location>
            <begin position="132"/>
            <end position="169"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00003689"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009646"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50915"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011476.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="169" mass="19958" checksum="9E4E3FB495EFC0E4" modified="2019-05-08" version="1">MGRPMNIILFEDRNFQGRSVECSSDRPDLQSHLSRCNSIRVESGCFMLYERPNFQGQQFFVKRGDYPDMQSEGFSTSIKSCRMIPPHRGTYRIKIYDKEDHRGNMVELTEDSPRVMDQLRSLEMLSCCVLEGYWILYELPNYRARQYLLMTGQYRRFSEWGTMSGRVSS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJI6</accession>
    <name>A0A452HJI6_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="11">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="8">Presynapse</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJI6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017194.1">
        <property type="protein sequence ID" value="ENSGAGP00000015048.1"/>
        <property type="gene ID" value="ENSGAGG00000009356.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030424">
        <property type="term" value="C:axon"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048788">
        <property type="term" value="C:cytoskeleton of presynaptic active zone"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098882">
        <property type="term" value="F:structural constituent of presynaptic active zone"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007274">
        <property type="term" value="P:neuromuscular synaptic transmission"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048167">
        <property type="term" value="P:regulation of synaptic plasticity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.1490">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019323">
        <property type="entry name" value="ELKS/CAST"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18861">
        <property type="entry name" value="ELKS/RAB6-INTERACTING/CAST PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18861:SF3">
        <property type="entry name" value="ERC PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10174">
        <property type="entry name" value="Cast"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90257">
        <property type="entry name" value="Myosin rod fragments"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0966">Cell projection</keyword>
    <keyword evidence="5 9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0770">Synapse</keyword>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1"/>
            <end position="44"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="147"/>
            <end position="174"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="211"/>
            <end position="276"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="332"/>
            <end position="846"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="1"/>
            <end position="13"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="14"/>
            <end position="25"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00034106"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015048.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="895" mass="103620" checksum="B969E5FBF5906AD5" modified="2019-05-08" version="1">MYGSARTISNLEGSPSRSPRLPRSPRLGHRRTSSGGGGGTGKTLSMENIQSLNAAYATSGPMYLSDHEGVASTTFPKGTMTLGRATNRAVYGGRVTAMGSSPNIASAGLSHTDVLSYTDQHGGLTTSSHHHHHQVPSMLRQVRDSTMLDLQAQLKELQRENDLLRKELDIKDSKLGSSMNSIKTFWSPELKKERVLRKEEAARMSVLKEQMRVSHEENQHLQLTIQALQDELRTQRDLNHLLQQESGNRGTEHFTIELTEENFRRLQAEHDRQAKELFLLRKTLEEMELRIETQKQTLNARDESIKKLLEMLQSKGLPSKGMEDDNERARRMAEAESQVNHLEVILDQKEKENIHLREELHRRTQLQPEPAKTKALQTVIEMKDTKIASLERNIRDLEDEIQMLKTNGVLNIEDREEEIKQIEVYKSHSKFMKNKIDQLKQELSKKESELLALQTKLETLSNQNSDCKQHIEVLKESLTAKEQRAAILQTEVDALRLRLEEKETFLNKKTKQLQDLTEEKGTLAGEIRDMKDMLEVKERKINVLQKKIENLQEQLRDKDKQLSNLKDRVKSLQTDSSNTDTALATLEEALSEKERIIERLKEQRERDDRERLEEIESFKKENKDLKEKVNALQAELTEKESNLIDLKEHASSLASTGLKRDSKLKSLEIAIEQKKEECSKLEAQLKKAHEAEEEARLNPEFADRMKQLDKEASYYREECGKAQAEVDRLLEILKEVENEKNDKDKKIAELERHVKDQNKKVANLKHNQQMEKKKNVQLLEEVRRREDNMTDNSQHLQLEELMNALEKTRQELDATKARLASTQQSLAEKEAHLANLRIERRKQLEEILEMKMTLLCLDQSGNYVVMIMDRKFKEHEIYCKIKSKINRPFLTFSFP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJI8</accession>
    <name>A0A452HJI8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">cAMP-dependent protein kinase</fullName>
            <ecNumber evidence="1">2.7.11.11</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="8">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.11"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="7">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.11"/>
        </reaction>
    </comment>
    <dbReference type="EC" id="2.7.11.11" evidence="1"/>
    <dbReference type="AlphaFoldDB" id="A0A452HJI8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017224.1">
        <property type="protein sequence ID" value="ENSGAGP00000015076.1"/>
        <property type="gene ID" value="ENSGAGG00000010963.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005952">
        <property type="term" value="C:cAMP-dependent protein kinase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004691">
        <property type="term" value="F:cAMP-dependent protein kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034237">
        <property type="term" value="F:protein kinase A regulatory subunit binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24353:SF153">
        <property type="entry name" value="CAMP-DEPENDENT PROTEIN KINASE CATALYTIC SUBUNIT 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24353">
        <property type="entry name" value="CYCLIC NUCLEOTIDE-DEPENDENT PROTEIN KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6 9" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="5" id="KW-0418">Kinase</keyword>
    <keyword evidence="4 9" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="3" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Protein kinase" evidence="11">
        <location>
            <begin position="41"/>
            <end position="308"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="243"/>
            <end position="308"/>
        </location>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="70"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00012444"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00047292"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00047454"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015076.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="308" mass="34543" checksum="907C969DAAE45162" modified="2019-05-08" version="1">GQGARLAKEGSPVKEFLAKGKEDFLKKWENPSQNTACLDQFERVKTLGMGSFGRVMLVKHKETGNYYAMKVLEKQKVRLVKLKQIEHTLNEKRILQAINFPLLIKLEYSFKDNSNLESHAWFYDLKPENLLLDQQGYIKVMDFGFAKRVKGRTWSLCGTPEYLAPEIILSKDYNKAVDWWALGVLIYEMAPIQIYEKIVSGKVFGLGEGTLVRFGGGFANQGSATCGTSSKHDRRADFEQHPATLSAPAHPSPLREQEAEAWFCGSQAPPPPRLPPTWRFPAPPPLLRQSVDGPSKGEGAERQPQAAP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJI9</accession>
    <name>A0A452HJI9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Auxilin</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="11">DnaJ homolog subfamily C member 6</fullName>
        </alternativeName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="9">Forms a complex composed of HSPA8, CLTC and DNAJC6. Interacts with HSPA8/HSC70 in an ATP-dependent manner; this interaction stimulates the HSPA8's ATPase activity. Interacts with CLTC; this interaction produces a local change in heavy-chain contacts, creating a detectable global distortion of the clathrin coat. Interacts with AP2A2. Interacts with DNM1(GTP-bound form); this interaction allows clathrin-coated vesicle (CCV) formation at the plasma membrane.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasmic vesicle</location>
            <location evidence="1">Clathrin-coated vesicle</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJI9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017226.1">
        <property type="protein sequence ID" value="ENSGAGP00000015078.1"/>
        <property type="gene ID" value="ENSGAGG00000011384.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030136">
        <property type="term" value="C:clathrin-coated vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0014069">
        <property type="term" value="C:postsynaptic density"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098793">
        <property type="term" value="C:presynapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="GOC"/>
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    <dbReference type="GO" id="GO:0030276">
        <property type="term" value="F:clathrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004721">
        <property type="term" value="F:phosphoprotein phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017124">
        <property type="term" value="F:SH3 domain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072583">
        <property type="term" value="P:clathrin-dependent endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016191">
        <property type="term" value="P:synaptic vesicle uncoating"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd06257">
        <property type="entry name" value="DnaJ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd14563">
        <property type="entry name" value="PTP_auxilin_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1110:FF:000001">
        <property type="entry name" value="cyclin-G-associated kinase isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000255">
        <property type="entry name" value="putative tyrosine-protein phosphatase auxilin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.110:FF:000002">
        <property type="entry name" value="putative tyrosine-protein phosphatase auxilin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1110">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.110">
        <property type="entry name" value="DnaJ domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001623">
        <property type="entry name" value="DnaJ_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036869">
        <property type="entry name" value="J_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014020">
        <property type="entry name" value="Tensin_C2-dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029023">
        <property type="entry name" value="Tensin_phosphatase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23172">
        <property type="entry name" value="AUXILIN/CYCLIN G-ASSOCIATED KINASE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23172:SF4">
        <property type="entry name" value="TYROSINE-PROTEIN PHOSPHATASE AUXILIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10409">
        <property type="entry name" value="PTEN_C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00271">
        <property type="entry name" value="DnaJ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01326">
        <property type="entry name" value="PTEN_C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49562">
        <property type="entry name" value="C2 domain (Calcium/lipid-binding domain, CaLB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46565">
        <property type="entry name" value="Chaperone J-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51182">
        <property type="entry name" value="C2_TENSIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50076">
        <property type="entry name" value="DNAJ_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51181">
        <property type="entry name" value="PPASE_TENSIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0143">Chaperone</keyword>
    <keyword evidence="8" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="4" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="5" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0729">SH3-binding</keyword>
    <feature type="domain" description="Phosphatase tensin-type" evidence="14">
        <location>
            <begin position="55"/>
            <end position="222"/>
        </location>
    </feature>
    <feature type="domain" description="C2 tensin-type" evidence="15">
        <location>
            <begin position="228"/>
            <end position="366"/>
        </location>
    </feature>
    <feature type="domain" description="J" evidence="13">
        <location>
            <begin position="866"/>
            <end position="930"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="431"/>
            <end position="522"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="537"/>
            <end position="794"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="434"/>
            <end position="456"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="464"/>
            <end position="475"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="510"/>
            <end position="522"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="664"/>
            <end position="684"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="692"/>
            <end position="707"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="745"/>
            <end position="755"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="756"/>
            <end position="772"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004132"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023036"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023186"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00064305"/>
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    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00069335"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00075670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS51181"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS51182"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015078.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="930" mass="101451" checksum="A86187A4D9CCF6A8" modified="2019-05-08" version="1">MKESENKGASSPDMESTYGGGLLDMVKGGAGRLFSNLKDNLKDTLKDTSSKVMQSVASYTKGELDISYVTSRIIVMSFPAEGVELGFRNHIEDVRTFLDSRHPDHYTVFNLSQKSYRSAKFHNRVSECSWPMRQAPSLHNLYAVCKNMHNWLQQNPKNVCVIHCMDGRAASSVLVSAMFCFCHLFSSPVPAIQLLNSKRPGIGLWPSHRRYLGYICDLIAEKPIDPHCKPLAIKSVCLSPVPCFNKQRNGCRPFCDILIGETRIFTTSQEYERMKEFRVQEGKVLIPLGVTVHGDVVLSVYHMRSTIGGRLQAKMTNTQIFQIQFHTGFIALGTTMLKFTKPELDACDSPEKYPQLFHVTLEIEIESADRQTDLTPPWENFTTKDINASILFSSHQEHQDALALAGRGPVDAHQDNIKNVGQGTFLSSLSWQDQKSDKSSSHPTSEDRAALVHEESEQSDDELLSLSSQHSNASSDKTHGTPKHSKKQQEPPAPLPPEDVDLLGLDGSPMSKNFPTQSTALPSNSDLLSDLFGVSDSRAHSQGVQAAAEDVFHVGGPGSAHSTPRRSAASASPSPAPRVGEATAFDPFGTRPKQSGPDLLGSFLGSSNATGDPFLQATRSPSPTVHSDPFHMASSTPTVAVQPDVSGGWEWSNKSGGLGMGSKSAATSPTGSLHSTPTHQSKPQTLDPFADLGALGANLAGGSSFASKPTTPTGISGGFPPMGSPQKPSPQPMGGSGWQQGAGYSWQQSQSKSQQNIPHASPQNRPNYNVSFSAMPGGQNDRGKGPANADSKPKVSADFEDLLSGQGFNAHKDKKGPRTIAEMRKEEMAKDMDPEKLKILEWIEGKERNIRALLSTMHTVLWAGETKWKPIGMADLVTPEQVKKVYRRAVLVVHPDKATGQPYEQYAKMIFMELNDAWSEFENQGQKPLY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJJ5</accession>
    <name>A0A452HJJ5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Auxilin</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="11">DnaJ homolog subfamily C member 6</fullName>
        </alternativeName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="9">Forms a complex composed of HSPA8, CLTC and DNAJC6. Interacts with HSPA8/HSC70 in an ATP-dependent manner; this interaction stimulates the HSPA8's ATPase activity. Interacts with CLTC; this interaction produces a local change in heavy-chain contacts, creating a detectable global distortion of the clathrin coat. Interacts with AP2A2. Interacts with DNM1(GTP-bound form); this interaction allows clathrin-coated vesicle (CCV) formation at the plasma membrane.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasmic vesicle</location>
            <location evidence="1">Clathrin-coated vesicle</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJJ5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017232.1">
        <property type="protein sequence ID" value="ENSGAGP00000015083.1"/>
        <property type="gene ID" value="ENSGAGG00000011384.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030136">
        <property type="term" value="C:clathrin-coated vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0014069">
        <property type="term" value="C:postsynaptic density"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098793">
        <property type="term" value="C:presynapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="GOC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030276">
        <property type="term" value="F:clathrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004721">
        <property type="term" value="F:phosphoprotein phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017124">
        <property type="term" value="F:SH3 domain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072583">
        <property type="term" value="P:clathrin-dependent endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016191">
        <property type="term" value="P:synaptic vesicle uncoating"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd06257">
        <property type="entry name" value="DnaJ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd14563">
        <property type="entry name" value="PTP_auxilin_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1110:FF:000001">
        <property type="entry name" value="cyclin-G-associated kinase isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000255">
        <property type="entry name" value="putative tyrosine-protein phosphatase auxilin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.110:FF:000002">
        <property type="entry name" value="putative tyrosine-protein phosphatase auxilin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1110">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.110">
        <property type="entry name" value="DnaJ domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001623">
        <property type="entry name" value="DnaJ_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036869">
        <property type="entry name" value="J_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014020">
        <property type="entry name" value="Tensin_C2-dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029023">
        <property type="entry name" value="Tensin_phosphatase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23172">
        <property type="entry name" value="AUXILIN/CYCLIN G-ASSOCIATED KINASE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23172:SF4">
        <property type="entry name" value="TYROSINE-PROTEIN PHOSPHATASE AUXILIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10409">
        <property type="entry name" value="PTEN_C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00271">
        <property type="entry name" value="DnaJ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01326">
        <property type="entry name" value="PTEN_C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49562">
        <property type="entry name" value="C2 domain (Calcium/lipid-binding domain, CaLB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46565">
        <property type="entry name" value="Chaperone J-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51182">
        <property type="entry name" value="C2_TENSIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50076">
        <property type="entry name" value="DNAJ_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51181">
        <property type="entry name" value="PPASE_TENSIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0143">Chaperone</keyword>
    <keyword evidence="8" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="4" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="5" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0729">SH3-binding</keyword>
    <feature type="domain" description="Phosphatase tensin-type" evidence="14">
        <location>
            <begin position="55"/>
            <end position="222"/>
        </location>
    </feature>
    <feature type="domain" description="C2 tensin-type" evidence="15">
        <location>
            <begin position="228"/>
            <end position="366"/>
        </location>
    </feature>
    <feature type="domain" description="J" evidence="13">
        <location>
            <begin position="843"/>
            <end position="907"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="431"/>
            <end position="522"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="537"/>
            <end position="771"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="434"/>
            <end position="456"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="464"/>
            <end position="475"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="510"/>
            <end position="522"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="641"/>
            <end position="661"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="669"/>
            <end position="684"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="722"/>
            <end position="732"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="733"/>
            <end position="749"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004132"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023036"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023186"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00064305"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00069335"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00075670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS51181"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS51182"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015083.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="907" mass="99062" checksum="27299B5408840233" modified="2019-05-08" version="1">MKESENKGASSPDMESTYGGGLLDMVKGGAGRLFSNLKDNLKDTLKDTSSKVMQSVASYTKGELDISYVTSRIIVMSFPAEGVELGFRNHIEDVRTFLDSRHPDHYTVFNLSQKSYRSAKFHNRVSECSWPMRQAPSLHNLYAVCKNMHNWLQQNPKNVCVIHCMDGRAASAVLVSAMFCFCHLFSSPVPAIQLLNSKRPGIGLWPSHRRYLGYICDLIAEKPIDPHCKPLAIKSVCLSPVPCFNKQRNGCRPFCDILIGETRIFTTSQEYERMKEFRVQEGKVLIPLGVTVHGDVVLSVYHMRSTIGGRLQAKMTNTQIFQIQFHTGFIALGTTMLKFTKPELDACDSPEKYPQLFHVTLEIEIESADRQTDLTPPWENFTTKDINASILFSSHQEHQDALALAGRGPVDAHQDNIKNVGQGTFLSSLSWQDQKSDKSSSHPTSEDRAALVHEESEQSDDELLSLSSQHSNASSDKTHGTPKHSKKQQEPPAPLPPEDVDLLGLDGSPMSKNFPTQSTALPSNSDLLSDLFGVSDSRAHSQGVQAAAEDVFHVGGPGSAHSTPRRSAASASPSPAPRVGEATAFDPFGTRPKQSGPDLLGSFLGSSNATGDPFLQATRSPSPTVHSDPFHMGGLGMGSKSAATSPTGSLHSTPTHQSKPQTLDPFADLGALGANLAGGSSFASKPTTPTGISGGFPPMGSPQKPSPQPMGGSGWQQGAGYSWQQSQSKSQQNIPHASPQNRPNYNVSFSAMPGGQNDRGKGPANADSKPKVSADFEDLLSGQGFNAHKDKKGPRTIAEMRKEEMAKDMDPEKLKILEWIEGKERNIRALLSTMHTVLWAGETKWKPIGMADLVTPEQVKKVYRRAVLVVHPDKATGQPYEQYAKMIFMELNDAWSEFENQGQKPLY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJJ6</accession>
    <name>A0A452HJJ6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Auxilin</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="11">DnaJ homolog subfamily C member 6</fullName>
        </alternativeName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="9">Forms a complex composed of HSPA8, CLTC and DNAJC6. Interacts with HSPA8/HSC70 in an ATP-dependent manner; this interaction stimulates the HSPA8's ATPase activity. Interacts with CLTC; this interaction produces a local change in heavy-chain contacts, creating a detectable global distortion of the clathrin coat. Interacts with AP2A2. Interacts with DNM1(GTP-bound form); this interaction allows clathrin-coated vesicle (CCV) formation at the plasma membrane.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasmic vesicle</location>
            <location evidence="1">Clathrin-coated vesicle</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJJ6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017239.1">
        <property type="protein sequence ID" value="ENSGAGP00000015089.1"/>
        <property type="gene ID" value="ENSGAGG00000011384.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030136">
        <property type="term" value="C:clathrin-coated vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0014069">
        <property type="term" value="C:postsynaptic density"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098793">
        <property type="term" value="C:presynapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="GOC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030276">
        <property type="term" value="F:clathrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004721">
        <property type="term" value="F:phosphoprotein phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017124">
        <property type="term" value="F:SH3 domain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072583">
        <property type="term" value="P:clathrin-dependent endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016191">
        <property type="term" value="P:synaptic vesicle uncoating"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd06257">
        <property type="entry name" value="DnaJ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd14563">
        <property type="entry name" value="PTP_auxilin_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1110:FF:000001">
        <property type="entry name" value="cyclin-G-associated kinase isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000255">
        <property type="entry name" value="putative tyrosine-protein phosphatase auxilin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.110:FF:000002">
        <property type="entry name" value="putative tyrosine-protein phosphatase auxilin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1110">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.110">
        <property type="entry name" value="DnaJ domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001623">
        <property type="entry name" value="DnaJ_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036869">
        <property type="entry name" value="J_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014020">
        <property type="entry name" value="Tensin_C2-dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029023">
        <property type="entry name" value="Tensin_phosphatase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23172">
        <property type="entry name" value="AUXILIN/CYCLIN G-ASSOCIATED KINASE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23172:SF4">
        <property type="entry name" value="TYROSINE-PROTEIN PHOSPHATASE AUXILIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10409">
        <property type="entry name" value="PTEN_C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00271">
        <property type="entry name" value="DnaJ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01326">
        <property type="entry name" value="PTEN_C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49562">
        <property type="entry name" value="C2 domain (Calcium/lipid-binding domain, CaLB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46565">
        <property type="entry name" value="Chaperone J-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51182">
        <property type="entry name" value="C2_TENSIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50076">
        <property type="entry name" value="DNAJ_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51181">
        <property type="entry name" value="PPASE_TENSIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0143">Chaperone</keyword>
    <keyword evidence="8" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="4" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="5" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0729">SH3-binding</keyword>
    <feature type="domain" description="Phosphatase tensin-type" evidence="14">
        <location>
            <begin position="55"/>
            <end position="222"/>
        </location>
    </feature>
    <feature type="domain" description="C2 tensin-type" evidence="15">
        <location>
            <begin position="228"/>
            <end position="366"/>
        </location>
    </feature>
    <feature type="domain" description="J" evidence="13">
        <location>
            <begin position="798"/>
            <end position="862"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="431"/>
            <end position="522"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="537"/>
            <end position="726"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="434"/>
            <end position="456"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="464"/>
            <end position="475"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="510"/>
            <end position="522"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="636"/>
            <end position="646"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="677"/>
            <end position="687"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="688"/>
            <end position="704"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004132"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023036"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023186"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00064305"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00069335"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00075670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS51181"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS51182"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015089.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="862" mass="94732" checksum="FC953A3A831602AF" modified="2019-05-08" version="1">MKESENKGASSPDMESTYGGGLLDMVKGGAGRLFSNLKDNLKDTLKDTSSKVMQSVASYTKGELDISYVTSRIIVMSFPAEGVELGFRNHIEDVRTFLDSRHPDHYTVFNLSQKSYRSAKFHNRVSECSWPMRQAPSLHNLYAVCKNMHNWLQQNPKNVCVIHCMDGRAASAVLVSAMFCFCHLFSSPVPAIQLLNSKRPGIGLWPSHRRYLGYICDLIAEKPIDPHCKPLAIKSVCLSPVPCFNKQRNGCRPFCDILIGETRIFTTSQEYERMKEFRVQEGKVLIPLGVTVHGDVVLSVYHMRSTIGGRLQAKMTNTQIFQIQFHTGFIALGTTMLKFTKPELDACDSPEKYPQLFHVTLEIEIESADRQTDLTPPWENFTTKDINASILFSSHQEHQDALALAGRGPVDAHQDNIKNVGQGTFLSSLSWQDQKSDKSSSHPTSEDRAALVHEESEQSDDELLSLSSQHSNASSDKTHGTPKHSKKQQEPPAPLPPEDVDLLGLDGSPMSKNFPTQSTALPSNSDLLSDLFGVSDSRAHSQGVQAAAEDVFHVGGPGSAHSTPRRSAASASPSPAPRVGEATAFDPFGTRPKQSGPDLLGSFLGSSNATGDPFLQATRSPSPTVHSDPFHMGGSSFASKPTTPTGISGGFPPMGSPQKPSPQPMGGSGWQQGAGYSWQQSQSKSQQNIPHASPQNRPNYNVSFSAMPGGQNDRGKGPANADSKPKVSADFEDLLSGQGFNAHKDKKGPRTIAEMRKEEMAKDMDPEKLKILEWIEGKERNIRALLSTMHTVLWAGETKWKPIGMADLVTPEQVKKVYRRAVLVVHPDKATGQPYEQYAKMIFMELNDAWSEFENQGQKPLY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJM0</accession>
    <name>A0A452HJM0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Retinol dehydrogenase 16</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1 2">Belongs to the short-chain dehydrogenases/reductases (SDR) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJM0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017261.1">
        <property type="protein sequence ID" value="ENSGAGP00000015109.1"/>
        <property type="gene ID" value="ENSGAGG00000011406.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016491">
        <property type="term" value="F:oxidoreductase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008202">
        <property type="term" value="P:steroid metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.720">
        <property type="entry name" value="NAD(P)-binding Rossmann-like Domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036291">
        <property type="entry name" value="NAD(P)-bd_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002347">
        <property type="entry name" value="SDR_fam"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43313:SF48">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43313">
        <property type="entry name" value="SHORT-CHAIN DEHYDROGENASE/REDUCTASE FAMILY 9C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00106">
        <property type="entry name" value="adh_short"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00081">
        <property type="entry name" value="GDHRDH"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00080">
        <property type="entry name" value="SDRFAMILY"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51735">
        <property type="entry name" value="NAD(P)-binding Rossmann-fold domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006484"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="RuleBase" id="RU000363"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015109.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="5">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="224" mass="24988" checksum="8C8E9CC34AE7231E" modified="2019-05-08" version="1">HHIKEKGTGRQATPKPFCPLCPGLWGLVNNAGIAVPTAPNEWLTKDDFVKVLDVNLIGLIEVTLSFLPLVKRARGRVVNVASTMGRVAMLGGGYSLSKYGVEAFSDSLRRELLPFGVKVCIIEPGAFRTQILNTQLLQDMVKEVWGRVPSEIKESYGQRYFDSCDPNLFLVTDCMEHALTARHPRTRYSGGWDAKLYFIPVSYLPAWCQDFLLTLQQPKPAHGI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJM1</accession>
    <name>A0A452HJM1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Aryl hydrocarbon receptor nuclear translocator</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Required for activity of the AHR. Upon ligand binding, AHR translocates into the nucleus, where it heterodimerizes with ARNT and induces transcription by binding to xenobiotic response elements (XRE). Not required for the ligand-binding subunit to translocate from the cytosol to the nucleus after ligand binding. The complex initiates transcription of genes involved in the regulation of a variety of biological processes, including angiogenesis, hematopoiesis, drug and lipid metabolism, cell motility and immune modulation. The heterodimer binds to core DNA sequence 5'-TACGTG-3' within the hypoxia response element (HRE) of target gene promoters and functions as a transcriptional regulator of the adaptive response to hypoxia. The heterodimer ARNT:AHR binds to core DNA sequence 5'-TGCGTG-3' within the dioxin response element (DRE) of target gene promoters and activates their transcription.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJM1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017263.1">
        <property type="protein sequence ID" value="ENSGAGP00000015111.1"/>
        <property type="gene ID" value="ENSGAGG00000011404.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005667">
        <property type="term" value="C:transcription regulator complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003700">
        <property type="term" value="F:DNA-binding transcription factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046983">
        <property type="term" value="F:protein dimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd18947">
        <property type="entry name" value="bHLH-PAS_ARNT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00130">
        <property type="entry name" value="PAS"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.450.20:FF:000028">
        <property type="entry name" value="Aryl hydrocarbon receptor nuclear translocator 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.450.20:FF:000003">
        <property type="entry name" value="Aryl hydrocarbon receptor nuclear translocator 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.280.10:FF:000011">
        <property type="entry name" value="Aryl hydrocarbon receptor nuclear translocator 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.280.10">
        <property type="entry name" value="Helix-loop-helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.450.20">
        <property type="entry name" value="PAS domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011598">
        <property type="entry name" value="bHLH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050933">
        <property type="entry name" value="Circadian_TF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036638">
        <property type="entry name" value="HLH_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001067">
        <property type="entry name" value="Nuc_translocat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001610">
        <property type="entry name" value="PAC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000014">
        <property type="entry name" value="PAS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035965">
        <property type="entry name" value="PAS-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013767">
        <property type="entry name" value="PAS_fold"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00229">
        <property type="entry name" value="sensory_box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23042">
        <property type="entry name" value="CIRCADIAN PROTEIN CLOCK/ARNT/BMAL/PAS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00010">
        <property type="entry name" value="HLH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00989">
        <property type="entry name" value="PAS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14598">
        <property type="entry name" value="PAS_11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00785">
        <property type="entry name" value="NCTRNSLOCATR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00353">
        <property type="entry name" value="HLH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00086">
        <property type="entry name" value="PAC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00091">
        <property type="entry name" value="PAS"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47459">
        <property type="entry name" value="HLH, helix-loop-helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55785">
        <property type="entry name" value="PYP-like sensor domain (PAS domain)"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50888">
        <property type="entry name" value="BHLH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50112">
        <property type="entry name" value="PAS"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0010">Activator</keyword>
    <keyword evidence="4" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0804">Transcription</keyword>
    <keyword evidence="3" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="BHLH" evidence="12">
        <location>
            <begin position="76"/>
            <end position="129"/>
        </location>
    </feature>
    <feature type="domain" description="PAS" evidence="11">
        <location>
            <begin position="148"/>
            <end position="222"/>
        </location>
    </feature>
    <feature type="domain" description="PAS" evidence="11">
        <location>
            <begin position="354"/>
            <end position="405"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1"/>
            <end position="70"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="453"/>
            <end position="475"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="639"/>
            <end position="697"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="715"/>
            <end position="758"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="10">
        <location>
            <begin position="47"/>
            <end position="57"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="58"/>
            <end position="70"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="453"/>
            <end position="473"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="639"/>
            <end position="650"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="661"/>
            <end position="683"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="725"/>
            <end position="736"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00039652"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00045949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50112"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50888"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015111.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="770" mass="84556" checksum="34C9B92906F0061B" modified="2019-05-08" version="1">MAAAGNPEMASDVPSLSAAVSSGNPGPGTQAGVAIVQRTNKRRPGLDFDDDDEEEEEGSKFLRCDDDQIPNDKERFARENHSEIERRRRNKMTAYITELSDMVPTCSALARKPDKLTILRMAVSHMKSLRGTGNTSTDGTYKPSFLTDQELKHLILEAADGFLFIVSCETGRVVYVSDSVTPVLNQPQSEWFSSTLYDQVHPDDVDKLREQLSTSENALTGRILDLKTGTVKKEGQQSMRMCMGSRRSFICRMRCGNSSVDPASMNRLSFVRSRCRSGLSAVKDGEPHYVVVHCTGYIKAWPPAGVSLPDDDPDAGQGSKFCLVAIGRLQVTSSPNCTDLNSICQPTEFISRHNTESIFTFVDHRCVATVGYQPQELLGKDIVDFCHPEDQQLLRDSFQQVVKLKGQVLSVMFRFRSKNREWLWMRTSSFSFQNPYSDEIEYIICTNTNVKNSSQESRPAVSNSMQRPQQGQNVGLPLDVGTAQLASRQQQPQQTALEVVPGRESLSSYDHSQVAVQPVTAAGPERSKPLEKTEALFSQERDPRFSEIYSGINTDQSKVLPASTVPANQPLFPQGNTFTPARPTESFRSSSMVPPVINIQHQPASAGQILAQISRHSNTAPVTATNWVSATRPSFTAQQVASQTAKTRSPSFGIGSFQGTPSSFSPMSAPSSTASPGGAAYPPLANRGTGFATETGQTPAQFQTRTAEGVGMWPQWPGQHHGRSSGEQHVQQQPSQPEVFPDMLSMLGDQGTNYSNEEFPDLNIFPAFSE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJM3</accession>
    <name>A0A452HJM3_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="11">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="8">Presynapse</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJM3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017264.1">
        <property type="protein sequence ID" value="ENSGAGP00000015112.1"/>
        <property type="gene ID" value="ENSGAGG00000009356.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030424">
        <property type="term" value="C:axon"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048788">
        <property type="term" value="C:cytoskeleton of presynaptic active zone"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098882">
        <property type="term" value="F:structural constituent of presynaptic active zone"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007274">
        <property type="term" value="P:neuromuscular synaptic transmission"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048167">
        <property type="term" value="P:regulation of synaptic plasticity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.1490">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019323">
        <property type="entry name" value="ELKS/CAST"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18861">
        <property type="entry name" value="ELKS/RAB6-INTERACTING/CAST PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18861:SF3">
        <property type="entry name" value="ERC PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10174">
        <property type="entry name" value="Cast"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90257">
        <property type="entry name" value="Myosin rod fragments"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0966">Cell projection</keyword>
    <keyword evidence="5 9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0770">Synapse</keyword>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1"/>
            <end position="44"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="147"/>
            <end position="174"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="211"/>
            <end position="276"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="332"/>
            <end position="862"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="1"/>
            <end position="13"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="14"/>
            <end position="25"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00034106"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015112.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="867" mass="100287" checksum="FEF307FABBAD356C" modified="2019-05-08" version="1">MYGSARTISNLEGSPSRSPRLPRSPRLGHRRTSSGGGGGTGKTLSMENIQSLNAAYATSGPMYLSDHEGVASTTFPKGTMTLGRATNRAVYGGRVTAMGSSPNIASAGLSHTDVLSYTDQHGGLTTSSHHHHHQVPSMLRQVRDSTMLDLQAQLKELQRENDLLRKELDIKDSKLGSSMNSIKTFWSPELKKERVLRKEEAARMSVLKEQMRVSHEENQHLQLTIQALQDELRTQRDLNHLLQQESGNRGTEHFTIELTEENFRRLQAEHDRQAKELFLLRKTLEEMELRIETQKQTLNARDESIKKLLEMLQSKGLPSKGMEDDNERARRMAEAESQVNHLEVILDQKEKENIHLREELHRRTQLQPEPAKTKALQTVIEMKDTKIASLERNIRDLEDEIQMLKTNGVLNIEDREEEIKQIEVYKSHSKFMKNKIDQLKQELSKKESELLALQTKLETLSNQNSDCKQHIEVLKESLTAKEQRAAILQTEVDALRLRLEEKETFLNKKTKQLQDLTEEKGTLAGEIRDMKDMLEVKERKINVLQKKIENLQEQLRDKDKQLSNLKDRVKSLQTDSSNTDTALATLEEALSEKERIIERLKEQRERDDRERLEEIESFKKENKDLKEKVNALQAELTEKESNLIDLKEHASSLASTGLKRDSKLKSLEIAIEQKKEECSKLEAQLKKAHEAEEEARLNPEFADRMKQLDKEASYYREECGKAQAEVDRLLEILKEVENEKNDKDKKIAELERHVKDQNKKVANLKHNQQMEKKKNVQLLEEVRRREDNMTDNSQHLQVKKLFSLAYEDRTRNNLEELMNALEKTRQELDATKARLASTQQSLAEKEAHLANLRIERRKQLEEILEMK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJM4</accession>
    <name>A0A452HJM4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Protein S100-B</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="15">S-100 protein beta chain</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="17">S-100 protein subunit beta</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="16">S100 calcium-binding protein B</fullName>
        </alternativeName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="18">Dimer of either two alpha chains, or two beta chains, or one alpha and one beta chain. The S100B dimer binds two molecules of STK38. Interacts with CACYBP in a calcium-dependent manner. Interacts with ATAD3A; this interaction probably occurs in the cytosol prior to ATAD3A mitochondrial targeting. Interacts with S100A6. The S100B dimer interacts with two molecules of CAPZA1. Interacts with AGER. Interacts with PPP5C (via TPR repeats); the interaction is calcium-dependent and modulates PPP5C activity. Interacts with TPPP; this interaction inhibits TPPP dimerization. Interacts with isoform CLSTN3beta of CLSTN3; interaction promotes secretion.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the S-100 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJM4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017243.1">
        <property type="protein sequence ID" value="ENSGAGP00000015093.1"/>
        <property type="gene ID" value="ENSGAGG00000011396.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048306">
        <property type="term" value="F:calcium-dependent protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050786">
        <property type="term" value="F:RAGE receptor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0044548">
        <property type="term" value="F:S100 protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043123">
        <property type="term" value="P:positive regulation of canonical NF-kappaB signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008284">
        <property type="term" value="P:positive regulation of cell population proliferation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05027">
        <property type="entry name" value="S-100B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.238.10:FF:000044">
        <property type="entry name" value="Protein S100"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.238.10">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011992">
        <property type="entry name" value="EF-hand-dom_pair"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002048">
        <property type="entry name" value="EF_hand_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028481">
        <property type="entry name" value="S100-B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001751">
        <property type="entry name" value="S100/CaBP7/8-like_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013787">
        <property type="entry name" value="S100_Ca-bd_sub"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11639:SF134">
        <property type="entry name" value="PROTEIN S100-A1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11639">
        <property type="entry name" value="S100 CALCIUM-BINDING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01023">
        <property type="entry name" value="S_100"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00054">
        <property type="entry name" value="EFh"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01394">
        <property type="entry name" value="S_100"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47473">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50222">
        <property type="entry name" value="EF_HAND_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00303">
        <property type="entry name" value="S100_CABP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="13" id="KW-0007">Acetylation</keyword>
    <keyword evidence="11" id="KW-0106">Calcium</keyword>
    <keyword evidence="12" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="14" id="KW-0539">Nucleus</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0677">Repeat</keyword>
    <keyword evidence="7" id="KW-0964">Secreted</keyword>
    <keyword evidence="10" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="EF-hand" evidence="19">
        <location>
            <begin position="49"/>
            <end position="84"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00007323"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00021222"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00030603"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00031938"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00033127"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00047135"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50222"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015093.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="92" mass="10492" checksum="96F11CA570350197" modified="2019-05-08" version="1">MSALEKAMIAIIEAFHQYSGKEGDKHKLKKSELKELINNELPHFLGEIKDQETVDKVMETLDADGDAECDFQEFVAFIAMVTSACHEFFEHE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJM5</accession>
    <name>A0A452HJM5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Retinol dehydrogenase 16</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the short-chain dehydrogenases/reductases (SDR) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJM5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017266.1">
        <property type="protein sequence ID" value="ENSGAGP00000015114.1"/>
        <property type="gene ID" value="ENSGAGG00000011406.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016491">
        <property type="term" value="F:oxidoreductase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008202">
        <property type="term" value="P:steroid metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.720">
        <property type="entry name" value="NAD(P)-binding Rossmann-like Domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036291">
        <property type="entry name" value="NAD(P)-bd_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002347">
        <property type="entry name" value="SDR_fam"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43313:SF48">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43313">
        <property type="entry name" value="SHORT-CHAIN DEHYDROGENASE/REDUCTASE FAMILY 9C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00106">
        <property type="entry name" value="adh_short"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51735">
        <property type="entry name" value="NAD(P)-binding Rossmann-fold domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="2">
        <location>
            <begin position="1"/>
            <end position="19"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019291741" description="Retinol dehydrogenase 16" evidence="2">
        <location>
            <begin position="20"/>
            <end position="170"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006484"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015114.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="5">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="170" mass="19158" checksum="F3D4CE2141F250BC" modified="2019-05-08" version="1">MWLYLAALLGLYFLRRWYRERQSVEGLPEKYVLITGCDSGFGNQLAKQLDTRGLRVLAACLTQQGAERLRKATSERLQTVILDVTRTDSIAAATAWVKEQVGNKGLEATASPNLFLVTDCMEHALTARHPRTRYSGGWDAKLYFIPVSYLPAWCQDFLLTLQQPKPAHGI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJM9</accession>
    <name>A0A452HJM9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="13">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="7">
            <text>Ca(2+)(in) = Ca(2+)(out)</text>
            <dbReference type="Rhea" id="RHEA:29671"/>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the CSC1 (TC 1.A.17) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJM9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017269.1">
        <property type="protein sequence ID" value="ENSGAGP00000015117.1"/>
        <property type="gene ID" value="ENSGAGG00000011403.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005227">
        <property type="term" value="F:calcium-activated cation channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045122">
        <property type="entry name" value="Csc1-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003864">
        <property type="entry name" value="CSC1/OSCA1-like_7TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027815">
        <property type="entry name" value="CSC1/OSCA1-like_cyt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032880">
        <property type="entry name" value="CSC1/OSCA1-like_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13018:SF38">
        <property type="entry name" value="CSC1-LIKE PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13018">
        <property type="entry name" value="PROBABLE MEMBRANE PROTEIN DUF221-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14703">
        <property type="entry name" value="PHM7_cyt"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02714">
        <property type="entry name" value="RSN1_7TM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13967">
        <property type="entry name" value="RSN1_TM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6 9" id="KW-0472">Membrane</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 9" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 9" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="40"/>
            <end position="67"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="150"/>
            <end position="170"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="196"/>
            <end position="215"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="422"/>
            <end position="446"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="466"/>
            <end position="489"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="510"/>
            <end position="533"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="564"/>
            <end position="586"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="607"/>
            <end position="629"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="635"/>
            <end position="650"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="671"/>
            <end position="691"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="703"/>
            <end position="725"/>
        </location>
    </feature>
    <feature type="domain" description="CSC1/OSCA1-like N-terminal transmembrane" evidence="11">
        <location>
            <begin position="112"/>
            <end position="216"/>
        </location>
    </feature>
    <feature type="domain" description="CSC1/OSCA1-like cytosolic" evidence="12">
        <location>
            <begin position="232"/>
            <end position="414"/>
        </location>
    </feature>
    <feature type="domain" description="CSC1/OSCA1-like 7TM region" evidence="10">
        <location>
            <begin position="425"/>
            <end position="697"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="770"/>
            <end position="816"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="8">
        <location>
            <begin position="777"/>
            <end position="792"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007779"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00036634"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="Pfam" id="PF02714"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF13967"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF14703"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015117.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="816" mass="92803" checksum="DAF2BCA31255B771" modified="2019-05-08" version="1">MLPFLFATLGSAGATCKVPDCNSTKDYCYSARIRSTVLQGLPFGGVPTVLALDFMCFLALLFVFSILRKVAWDYGRLALVTDADSVASALHSDSHDRYERLTSVSSSVDFDQRDNGFCSWLTAIFRIKDDEIRDKCGADAVHYLSFQRHIIGLLVAVGVLSVGIVLPVNFSGDLLENNPYSFGRTTIANLNSGNNLLWLHTSFAFLYLLLTVYSMRRHTSKMRYKEDDLVKRTLFINGISKYAESEKIKKHFEEAYSNCTVLEARPCYNVARLMFLDTERKKAERGKIYFTNLKTKENASSMINPKPCGHLCCCVIKGCEEVEAIEYYTKLEEKLKDDYKREKEKVNEKPLGMAFVTFHNETITAIILKDFNVCKCQGCVCHGEPRASSCSESLHISNWTVTYAPDPQNIYWEHLSIRGFIWWIRCLVINVILFLLLFFLTTPAIIITTMDKFNVTKPVEYLNNPIITQFFPTLLLWCFSALLPTIVYYSAFFEAHWTRSGENRTTMHKCYTFLIFMVLLLPSLGLSSLDVFFRWLFDKKFLAEAAVRFECVFLPDNGAFFVNYVIASAFIGNAMDLLRIPGLLMYMIRLCLARSAAERRNVKRHQAYEFQFGAAYAWMMCVFTVVMTYSITCPIIVPFGLMYMLLKHLVDRYNLYYAYLPAKLDKKIHSGAVNQVVAAPILCLFWLLFFSTMRTGFLAPTSMFTFVVLVITIVICLCHVCFGHFKYLSAHNYKIDHTEVDAVDGRVNGQPAAVPPAPKSAQKYIAQVLQDSSPEGDAAESEEQGSQDEELINPDGMNDTDFQSCEDSLIENEIHQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJN2</accession>
    <name>A0A452HJN2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">cAMP-dependent protein kinase</fullName>
            <ecNumber evidence="1">2.7.11.11</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="8">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.11"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="7">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.11"/>
        </reaction>
    </comment>
    <dbReference type="EC" id="2.7.11.11" evidence="1"/>
    <dbReference type="AlphaFoldDB" id="A0A452HJN2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017277.1">
        <property type="protein sequence ID" value="ENSGAGP00000015125.1"/>
        <property type="gene ID" value="ENSGAGG00000010963.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005952">
        <property type="term" value="C:cAMP-dependent protein kinase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004691">
        <property type="term" value="F:cAMP-dependent protein kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034237">
        <property type="term" value="F:protein kinase A regulatory subunit binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24353:SF153">
        <property type="entry name" value="CAMP-DEPENDENT PROTEIN KINASE CATALYTIC SUBUNIT 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24353">
        <property type="entry name" value="CYCLIC NUCLEOTIDE-DEPENDENT PROTEIN KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6 9" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="5" id="KW-0418">Kinase</keyword>
    <keyword evidence="4 9" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="3" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Protein kinase" evidence="10">
        <location>
            <begin position="1"/>
            <end position="228"/>
        </location>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="78"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00012444"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00047292"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00047454"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015125.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="228" mass="25947" checksum="38B02F2A984D1B50" modified="2019-05-08" version="1">MTLRAASSVDMVSHSSSSESLGEFLAKGKEDFLKKWENPSQNTACLDQFERVKTLGMGSFGRVMLVKHKETGNYYAMKVLEKQKVRLVKLKQIEHTLNEKRILQAINFPLLIKLEYSFKDNSNLESHAWFYDLKPENLLLDQQGYIKVMDFGFAKRVKGRTWSLCGTPEYLAPEIILSKDYNKAVDWWALGVLIYEMAPIQIYEKIVSGKVFGLGEGTLVRFGGGFAS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJP0</accession>
    <name>A0A452HJP0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">cAMP-dependent protein kinase</fullName>
            <ecNumber evidence="1">2.7.11.11</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="8">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.11"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="7">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.11"/>
        </reaction>
    </comment>
    <dbReference type="EC" id="2.7.11.11" evidence="1"/>
    <dbReference type="AlphaFoldDB" id="A0A452HJP0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017280.1">
        <property type="protein sequence ID" value="ENSGAGP00000015126.1"/>
        <property type="gene ID" value="ENSGAGG00000010963.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005952">
        <property type="term" value="C:cAMP-dependent protein kinase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004691">
        <property type="term" value="F:cAMP-dependent protein kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034237">
        <property type="term" value="F:protein kinase A regulatory subunit binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24353:SF153">
        <property type="entry name" value="CAMP-DEPENDENT PROTEIN KINASE CATALYTIC SUBUNIT 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24353">
        <property type="entry name" value="CYCLIC NUCLEOTIDE-DEPENDENT PROTEIN KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6 9" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="5" id="KW-0418">Kinase</keyword>
    <keyword evidence="4 9" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="3" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Protein kinase" evidence="10">
        <location>
            <begin position="1"/>
            <end position="211"/>
        </location>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="61"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00012444"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00047292"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00047454"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015126.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="211" mass="24237" checksum="FCAED9B6DF229957" modified="2025-10-08" version="2">MGLPLEFLAKGKEDFLKKWENPSQNTACLDQFERVKTLGMGSFGRVMLVKHKETGNYYAMKVLEKQKVRLVKLKQIEHTLNEKRILQAINFPLLIKLEYSFKDNSNLESHAWFYDLKPENLLLDQQGYIKVMDFGFAKRVKGRTWSLCGTPEYLAPEIILSKDYNKAVDWWALGVLIYEMAPIQIYEKIVSGKVFGLGEGTLVRFGGGFAS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK01</accession>
    <name>A0A452HK01_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="14">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HK01"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017383.1">
        <property type="protein sequence ID" value="ENSGAGP00000015221.1"/>
        <property type="gene ID" value="ENSGAGG00000011487.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003700">
        <property type="term" value="F:DNA-binding transcription factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07765">
        <property type="entry name" value="KRAB_A-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000739">
        <property type="entry name" value="Zgc:171418 protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001024">
        <property type="entry name" value="Zinc finger and SCAN domain-containing protein 20"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002343">
        <property type="entry name" value="Zinc finger protein 33A"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000200">
        <property type="entry name" value="zinc finger protein 510 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000912">
        <property type="entry name" value="Zinc finger protein 660"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001954">
        <property type="entry name" value="Zinc finger protein 787"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002074">
        <property type="entry name" value="Zinc finger protein 841"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002196">
        <property type="entry name" value="zinc finger protein 850-like isoform X3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001933">
        <property type="entry name" value="Zinc finger protein 870"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001909">
        <property type="entry name" value="KRAB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036051">
        <property type="entry name" value="KRAB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24390">
        <property type="entry name" value="ZINC FINGER PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24390:SF260">
        <property type="entry name" value="ZINC FINGER PROTEIN 383-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01352">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="SMART" id="SM00349">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109640">
        <property type="entry name" value="KRAB domain (Kruppel-associated box)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50805">
        <property type="entry name" value="KRAB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="10"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="9" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 11" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="KRAB" evidence="13">
        <location>
            <begin position="24"/>
            <end position="95"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="136"/>
            <end position="163"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="164"/>
            <end position="191"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="192"/>
            <end position="219"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="220"/>
            <end position="247"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="248"/>
            <end position="275"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="276"/>
            <end position="303"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="304"/>
            <end position="331"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="332"/>
            <end position="359"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="360"/>
            <end position="387"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="12">
        <location>
            <begin position="388"/>
            <end position="415"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015221.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="424" mass="48372" checksum="D10429BC6AB5A346" modified="2019-05-08" version="1">MEVKSINVGQQVLRVGLLLFQGRLTFEEVAVYFTREEWSLLDPAQRALYGDVMQENYETVVLLGFPISKPDVILQLERGEEPWVPDLQASEEEVLLRAACTEKAKAYETQQMPEENFSSHSDLITHKRINLEWRRYTCHECGKSFNWACLLVRHRRTHTGEKPYMCSECGKSFGQHSTLATHRRIHTGEMPYVCSECGKRFRDRSAHAKHVRLHTGEMPYTCPECGKSFIDSTCLISHQRTHTGETPYTCSECGQSFTLSGTLSRHRRIHTGEKPYGCSECGKSFSQRSDLITHQRTHTGEKPYTCSECGKSFYNSSSLKKHGRIHTGEKPYRCSECGKSFNRSSALISHERIHTGETPYTCAECGKSFAYHSGLITHKRIHTGEKPYTCSECGKRFHQSSHLVAHQRIHTDEKPYGCSDSDLT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK02</accession>
    <name>A0A452HK02_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">FAD-binding PCMH-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="1 14">
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="15">
            <name>Mo-molybdopterin</name>
            <dbReference type="ChEBI" id="CHEBI:71302"/>
        </cofactor>
        <text evidence="15">Binds 1 Mo-molybdopterin (Mo-MPT) cofactor per subunit.</text>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="12">
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="15">
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:49601"/>
        </cofactor>
        <text evidence="15">Binds 2 [2Fe-2S] clusters.</text>
    </comment>
    <comment type="subunit">
        <text evidence="3">Homodimer.</text>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the xanthine dehydrogenase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HK02"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017420.1">
        <property type="protein sequence ID" value="ENSGAGP00000015247.1"/>
        <property type="gene ID" value="ENSGAGG00000010622.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051537">
        <property type="term" value="F:2 iron, 2 sulfur cluster binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071949">
        <property type="term" value="F:FAD binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005506">
        <property type="term" value="F:iron ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016491">
        <property type="term" value="F:oxidoreductase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.365.10:FF:000003">
        <property type="entry name" value="Aldehyde oxidase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1170.50:FF:000001">
        <property type="entry name" value="Aldehyde oxidase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.365.10:FF:000025">
        <property type="entry name" value="Aldehyde oxidase 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.365.10:FF:000001">
        <property type="entry name" value="Xanthine dehydrogenase oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.365.10:FF:000004">
        <property type="entry name" value="Xanthine dehydrogenase oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.390.50:FF:000001">
        <property type="entry name" value="Xanthine dehydrogenase oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.43.10:FF:000001">
        <property type="entry name" value="Xanthine dehydrogenase/oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.465.10:FF:000004">
        <property type="entry name" value="Xanthine dehydrogenase/oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.20.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.465.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.120">
        <property type="entry name" value="[2Fe-2S]-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1170.50">
        <property type="entry name" value="Aldehyde oxidase/xanthine dehydrogenase, a/b hammerhead"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.365.10">
        <property type="entry name" value="Aldehyde oxidase/xanthine dehydrogenase, molybdopterin binding domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.390.50">
        <property type="entry name" value="CO dehydrogenase flavoprotein, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.43.10">
        <property type="entry name" value="Uridine Diphospho-n-acetylenolpyruvylglucosamine Reductase, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002888">
        <property type="entry name" value="2Fe-2S-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036884">
        <property type="entry name" value="2Fe-2S-bd_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036010">
        <property type="entry name" value="2Fe-2S_ferredoxin-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001041">
        <property type="entry name" value="2Fe-2S_ferredoxin-type"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006058">
        <property type="entry name" value="2Fe2S_fd_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000674">
        <property type="entry name" value="Ald_Oxase/Xan_DH_a/b"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036856">
        <property type="entry name" value="Ald_Oxase/Xan_DH_a/b_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016208">
        <property type="entry name" value="Ald_Oxase/xanthine_DH-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008274">
        <property type="entry name" value="AldOxase/xan_DH_MoCoBD1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046867">
        <property type="entry name" value="AldOxase/xan_DH_MoCoBD2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037165">
        <property type="entry name" value="AldOxase/xan_DH_Mopterin-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012675">
        <property type="entry name" value="Beta-grasp_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005107">
        <property type="entry name" value="CO_DH_flav_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036683">
        <property type="entry name" value="CO_DH_flav_C_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016166">
        <property type="entry name" value="FAD-bd_PCMH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036318">
        <property type="entry name" value="FAD-bd_PCMH-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016167">
        <property type="entry name" value="FAD-bd_PCMH_sub1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016169">
        <property type="entry name" value="FAD-bd_PCMH_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002346">
        <property type="entry name" value="Mopterin_DH_FAD-bd"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45444">
        <property type="entry name" value="XANTHINE DEHYDROGENASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45444:SF3">
        <property type="entry name" value="XANTHINE DEHYDROGENASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01315">
        <property type="entry name" value="Ald_Xan_dh_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03450">
        <property type="entry name" value="CO_deh_flav_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00941">
        <property type="entry name" value="FAD_binding_5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00111">
        <property type="entry name" value="Fer2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01799">
        <property type="entry name" value="Fer2_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02738">
        <property type="entry name" value="MoCoBD_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20256">
        <property type="entry name" value="MoCoBD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000127">
        <property type="entry name" value="Xanthine_DH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01008">
        <property type="entry name" value="Ald_Xan_dh_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01092">
        <property type="entry name" value="CO_deh_flav_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54292">
        <property type="entry name" value="2Fe-2S ferredoxin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55447">
        <property type="entry name" value="CO dehydrogenase flavoprotein C-terminal domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47741">
        <property type="entry name" value="CO dehydrogenase ISP C-domain like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54665">
        <property type="entry name" value="CO dehydrogenase molybdoprotein N-domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56176">
        <property type="entry name" value="FAD-binding/transporter-associated domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56003">
        <property type="entry name" value="Molybdenum cofactor-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00197">
        <property type="entry name" value="2FE2S_FER_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51387">
        <property type="entry name" value="FAD_PCMH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6 15" id="KW-0001">2Fe-2S</keyword>
    <keyword evidence="8 14" id="KW-0274">FAD</keyword>
    <keyword evidence="5" id="KW-0285">Flavoprotein</keyword>
    <keyword evidence="10 15" id="KW-0408">Iron</keyword>
    <keyword evidence="11 15" id="KW-0411">Iron-sulfur</keyword>
    <keyword evidence="7 15" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="4 15" id="KW-0500">Molybdenum</keyword>
    <keyword evidence="9" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="FAD-binding PCMH-type" evidence="16">
        <location>
            <begin position="177"/>
            <end position="360"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="13">
        <location>
            <position position="1207"/>
        </location>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="24"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="29"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="32"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="54"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="94"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>2</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="97"/>
        </location>
        <ligand>
            <name>[2Fe-2S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:190135"/>
            <label>2</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="306"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="368"/>
        </location>
        <ligand>
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="713"/>
        </location>
        <ligand>
            <name>Mo-molybdopterin</name>
            <dbReference type="ChEBI" id="CHEBI:71302"/>
        </ligand>
        <ligandPart>
            <name>Mo</name>
            <dbReference type="ChEBI" id="CHEBI:28685"/>
        </ligandPart>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="744"/>
        </location>
        <ligand>
            <name>Mo-molybdopterin</name>
            <dbReference type="ChEBI" id="CHEBI:71302"/>
        </ligand>
        <ligandPart>
            <name>Mo</name>
            <dbReference type="ChEBI" id="CHEBI:28685"/>
        </ligandPart>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="858"/>
        </location>
        <ligand>
            <name>Mo-molybdopterin</name>
            <dbReference type="ChEBI" id="CHEBI:71302"/>
        </ligand>
        <ligandPart>
            <name>Mo</name>
            <dbReference type="ChEBI" id="CHEBI:28685"/>
        </ligandPart>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <position position="860"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <feature type="binding site" evidence="15">
        <location>
            <position position="1025"/>
        </location>
        <ligand>
            <name>Mo-molybdopterin</name>
            <dbReference type="ChEBI" id="CHEBI:71302"/>
        </ligand>
        <ligandPart>
            <name>Mo</name>
            <dbReference type="ChEBI" id="CHEBI:28685"/>
        </ligandPart>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001974"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006849"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00011738"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022505"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022630"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022714"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022827"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023014"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00034078"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PIRSR" id="PIRSR000127-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PIRSR" id="PIRSR000127-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PIRSR" id="PIRSR000127-3"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS51387"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015247.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1274" mass="140973" checksum="943C70355CE04C27" modified="2019-05-08" version="1">TENQQLFSLTAAFAVRLSGTKYGCGIGGCGACTVMISTYEPTSKKIMHYPANSCLLPICSLHSSAVTTVEGVGSTKTRVHPIQERLAKCHGSQCGFCTPGMVMSMYALLRNHPEPSMEQITAALDGRFTFTIVSRILQNGGNQVCYEFQPVDPTQEFIFPPELMRMAQEQQKRTLVFRGERTTWISPTNLRELLELKAMYPKAPLIVGNTRLEMKRDGVHHPVFIHPTRIPDLHVVSNTNNGLVIGAACSLAQLRDILMKVVPELPEEKTKIFRALLQQLKTLAGEQIRSLASLGGHIVSRGSTWDLNPVLAVGNSVLSLASKDGTRQIPLNDQFLAGYENADLKPKEIIVSVFIPYSKKDDFVSAFRQAERRRNAVSVVNSGMRVLFKPNTDIIMDMNILYGGTGPITVSARKSCQRLIGRQWNEQMLDEACKLVLGEISLSPSAPGGKVEYRRTLLVSFLFKFYLEVLYGLNKMYPAKFPDLSKKNLSALGAFHSEVPQGMQTYQVSEDVDPVQPPQDPVGRPIMHQSGVKHATGEAVYCNDIRAVDEELFLAVVTSSRAHAKIVTIDVSEALKVPGVVDVVTAEDVPGKNGTDDEQAYAKDEVICVGQIVCAVVAESAVQAKQGVEKMKIEYEDLEPILTIEDAIKHNSYFGKEKKIEQGNIEEGFKSADEIIEGEIHIGGQEHFYLETNSVLVVPRGEDNEMDVHVSTQDAARVQEFVALALDVQSSKIMCHTKRVGGAFGGKITKPAIFAVVAAVAANKTGRPIRFTLERDEDMLITGGRHPFFGKYKVGFTNDGRITAADIEPYINAGCTLDDSEMVIEYVVLKTDNAYKIPNLRIRGRACKTNLPSNTAFRGFGFPQSGLFTESWITTVATQTGLPQEKIREINMYKEVNLTPFKEELDATNLVKCWKECMEKSEYYSRKTAVEEFNKQNYWKKKGIAIIPMKFSVGFNATQYHQAAALVHIYTDGSVLVTHGGIELGQGIHTKMLQIASRELKVPLSYVHFCETSTITVPNAIVTAGSIGSDVNGKAVQNACQILRKRLEPIINKNPEGKWEDWITEAYDQSVSLSATGYFRGYMVNMNWDTGEGQAFPYFVFGAACSEVEIDCLTGDHKNIRTDIVMDACFSINPAVDIGQIEGAFVQGVGLYTMEEIKFSPEGEQYTLGPDTYKIPAVCDIPEQFRVYLLPNSRNPIAIYSAKGIGEAGLFLGCSVFFAIWDAVAAVRKERGLTGTFTLNSPLTIERIRMACADNFTEMVRLDDHNGLFWPWNL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK05</accession>
    <name>A0A452HK05_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Cortactin-binding protein-2 N-terminal domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HK05"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017417.1">
        <property type="protein sequence ID" value="ENSGAGP00000015244.1"/>
        <property type="gene ID" value="ENSGAGG00000011507.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015629">
        <property type="term" value="C:actin cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.4090">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050719">
        <property type="entry name" value="Cortactin-Actin_Reg"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019131">
        <property type="entry name" value="Cortactin-binding_p2_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23166:SF3">
        <property type="entry name" value="FILAMIN-A-INTERACTING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23166">
        <property type="entry name" value="FILAMIN/GPBP-INTERACTING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09727">
        <property type="entry name" value="CortBP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1 2" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Cortactin-binding protein-2 N-terminal" evidence="4">
        <location>
            <begin position="73"/>
            <end position="254"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="1"/>
            <end position="70"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="1110"/>
            <end position="1158"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="2">
        <location>
            <begin position="196"/>
            <end position="223"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="2">
        <location>
            <begin position="257"/>
            <end position="287"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="2">
        <location>
            <begin position="327"/>
            <end position="598"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="2">
        <location>
            <begin position="628"/>
            <end position="760"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="1"/>
            <end position="13"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="3">
        <location>
            <begin position="1123"/>
            <end position="1137"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="1138"/>
            <end position="1154"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="Pfam" id="PF09727"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015244.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1175" mass="134701" checksum="58E8E2408257F896" modified="2019-05-08" version="1">MRSRNQGGESSPNGHFPSSKPIVSNAENENLQEDAKKKNKSHRKEEEVIVSGTVKRQLKSPAPGERKNKKSIELSKEDLIRLLSIMEGELQAREDVIHMLKTEKTKPEVLEAHYGSAAPENVLRVLHRDAILAQEKSIGEDVYEKPISELDRLEEKQKETYRRMLEQLLLAEKCHRRTVYELENEKHKHTDYMNKSDDFTNLLEQERERLKKLLEQEKIYQARKEKEHTKRLNKLRDELVKLKSFALMLVDERQMHIEQLGQQSQKIQDLAQKLKEEEEKLKVISSKTKEDGQKLMKLEAELEHKTLSFSQEHEEMTAKLGNQESHNRQLRLKLVGLTRRIEELEETNKNLHKAEEELQELRDKIAKGECGNSSLMAEVENLRKRVLEMEGKDEEITKTESQCRELKKKLQEEEHHSKELKFEVEKLQKRMSELEKLEEAFSKSKTECTQLHINLEKENNLTKDLISELDMVKTRVKDLESSESKLEKAELSLKEDLTKLKSFTVMLVDERKNMMEKIKQEERKVEGLNKNVKVEQGKVMDVTEKLIDESKKLLKLKTEMEEKVSSLTKERDELIGKLKSEEEKSSELSCRVDLLKKRIDGIEEVEREIARGRARKGQELAYHEDNKIKELTIETERLKKRLKQLEVVEGDLMKTEDEYDQLEQKFRTEQDKANFLSQQLEEMKLQIAKNKAIEKGEVVSQEAELRHRFRLEEAKSRDLKAEVQALKEKIHELMNKEDQLSQLQVDYSVLQQRFMEEEDKNKNMGQEVLNLTKELELSKRYSRVLRPSMNGRRMVDVPVTSTGVQTDAVSSEVAEEETPAVFIRKSFQEENHIMSNLRQVGLKKPIERSSVLNRYPPAANDLAMRKSWIPWMRKRENVSQGAHDKGARTHGSPGHPGEVVLSPKQGQPLHIRVTPDHENSTATLEITSPTSEEFFSSTTVIPTLGNQKPRITIIPSPNVSQKVKGGESSVGPERAMSPVTITTFSREKSADGGRAPFMERPISPIQIMTVSTSAAPSEISVSPELQDMTMGRAVFKVTPEKQTVPTPIRKYNANANIITTEDNKIHIHLGSQFKRSPGAASEGASPVITVRPMNVAAEKEVMTGTVLRSPRNHFSSRPGASKVTSTITITPVTTSSTRGTQSMTGQDGSSQRPTPTRIPVSKESVIIHQLRMNSR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK07</accession>
    <name>A0A452HK07_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">PPM-type phosphatase domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HK07"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017423.1">
        <property type="protein sequence ID" value="ENSGAGP00000015250.1"/>
        <property type="gene ID" value="ENSGAGG00000011331.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00143">
        <property type="entry name" value="PP2Cc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.60.40.10:FF:000003">
        <property type="entry name" value="PH domain and leucine-rich repeat protein phosphatase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.60.40.10">
        <property type="entry name" value="PPM-type phosphatase domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001611">
        <property type="entry name" value="Leu-rich_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003591">
        <property type="entry name" value="Leu-rich_rpt_typical-subtyp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050216">
        <property type="entry name" value="LRR_domain-containing"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036457">
        <property type="entry name" value="PPM-type-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001932">
        <property type="entry name" value="PPM-type_phosphatase-like_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48051">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48051:SF59">
        <property type="entry name" value="PH DOMAIN AND LEUCINE RICH REPEAT PROTEIN PHOSPHATASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13855">
        <property type="entry name" value="LRR_8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00481">
        <property type="entry name" value="PP2C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00364">
        <property type="entry name" value="LRR_BAC"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SMART" id="SM00369">
        <property type="entry name" value="LRR_TYP"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SMART" id="SM00332">
        <property type="entry name" value="PP2Cc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52058">
        <property type="entry name" value="L domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81606">
        <property type="entry name" value="PP2C-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51450">
        <property type="entry name" value="LRR"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51746">
        <property type="entry name" value="PPM_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0433">Leucine-rich repeat</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="PPM-type phosphatase" evidence="4">
        <location>
            <begin position="253"/>
            <end position="501"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="521"/>
            <end position="617"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="639"/>
            <end position="658"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="752"/>
            <end position="785"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="3">
        <location>
            <begin position="525"/>
            <end position="559"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="589"/>
            <end position="608"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="754"/>
            <end position="773"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022614"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS51746"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015250.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="785" mass="85335" checksum="0ACEFF9789074538" modified="2019-05-08" version="1">TDKGNSLCPPRTLSILSLRKLMVGHNHVQSLPPLLEHIPLEVLDLQHNLITKLPEMFFFKALNLRCLNVSANSLESLPSVCPGEESLSVLQLLYLTNNNLTDLCIPDLVGHPNLRVLHLANNQLQTFPASKLSKLEQLEELNLSGNKLKTIPTTIANCKQLHTLIAHSNEISIFPEILHLPRIQFVDLSCNELTEIMIPEVLPATLQELDLTGNTNLVLEHKTLDIFSHISTLKIDAKPSLTADSALSSTLWNHGLAEMAGQRNKLCVSALALGSFADSVEAVYGMFDGDKNEELPRLLQCTMADVLLEEVQQSNIDSVFMCNTFLVSHRKLGMAGQKLGSSAVLCCIRHDATDLAGSFTLTLANVGTCEAILCRSGKPLPLSKVFSLEQYPEEARRVKEQKAIITEDNKVNGVTCCTRMLGCTYLHPWILPKPHVYSIPLTVEDEMLLLGNKALWEHLSYTEAVMAVRHLHDPLAAAKKVCTLAQSYGCQDNVGAMVVCLNIGEDSCTCEVHGLPLPGPRGGISTAKPATPSSSSGTASEFSSELSASEVSSEVGSTASDEHSGAGLDAGLVPQPERRCSLHPVPPSSIFQRQPSCATFSSNQSDNGLDSDDEQPVEGVMINGSKVEVEVDIHCCQGKSLEPVPPEAEEGSAASSEDDSGGLFFRIWRQNSMTSNILLAGTKERCELQKSPSTCCLYGKKLSNGSIVPLEESLNLIEVATEVPKKKTGYFAAPAQLEPEDRFVVPPDLEEEMKEQIKQHHESGAEQKPREKAAVAPPDEFDTAL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK10</accession>
    <name>A0A452HK10_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="15">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="8">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HK10"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017422.1">
        <property type="protein sequence ID" value="ENSGAGP00000015249.1"/>
        <property type="gene ID" value="ENSGAGG00000011506.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004222">
        <property type="term" value="F:metalloendopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033631">
        <property type="term" value="P:cell-cell adhesion mediated by integrin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006509">
        <property type="term" value="P:membrane protein ectodomain proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007179">
        <property type="term" value="P:transforming growth factor beta receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd04269">
        <property type="entry name" value="ZnMc_adamalysin_II_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.390.10:FF:000002">
        <property type="entry name" value="Disintegrin and metalloproteinase domain-containing protein 22"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.70.10:FF:000001">
        <property type="entry name" value="Disintegrin and metalloproteinase domain-containing protein 22"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.390.10">
        <property type="entry name" value="Collagenase (Catalytic Domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.70.10">
        <property type="entry name" value="Disintegrin domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006586">
        <property type="entry name" value="ADAM_Cys-rich"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018358">
        <property type="entry name" value="Disintegrin_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001762">
        <property type="entry name" value="Disintegrin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036436">
        <property type="entry name" value="Disintegrin_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013111">
        <property type="entry name" value="EGF_extracell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024079">
        <property type="entry name" value="MetalloPept_cat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001590">
        <property type="entry name" value="Peptidase_M12B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002870">
        <property type="entry name" value="Peptidase_M12B_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034027">
        <property type="entry name" value="Reprolysin_adamalysin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11905">
        <property type="entry name" value="ADAM A DISINTEGRIN AND METALLOPROTEASE DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11905:SF136">
        <property type="entry name" value="DISINTEGRIN AND METALLOPROTEINASE DOMAIN-CONTAINING PROTEIN 9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08516">
        <property type="entry name" value="ADAM_CR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00200">
        <property type="entry name" value="Disintegrin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07974">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01562">
        <property type="entry name" value="Pep_M12B_propep"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01421">
        <property type="entry name" value="Reprolysin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00289">
        <property type="entry name" value="DISINTEGRIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00608">
        <property type="entry name" value="ACR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00050">
        <property type="entry name" value="DISIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57552">
        <property type="entry name" value="Blood coagulation inhibitor (disintegrin)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55486">
        <property type="entry name" value="Metalloproteases ('zincins'), catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50215">
        <property type="entry name" value="ADAM_MEPRO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00427">
        <property type="entry name" value="DISINTEGRIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50214">
        <property type="entry name" value="DISINTEGRIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01186">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6 8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="2 8" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="5 11" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 11" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 11" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="9" id="KW-0862">Zinc</keyword>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="668"/>
            <end position="687"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase M12B" evidence="14">
        <location>
            <begin position="181"/>
            <end position="375"/>
        </location>
    </feature>
    <feature type="domain" description="Disintegrin" evidence="13">
        <location>
            <begin position="383"/>
            <end position="470"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="12">
        <location>
            <begin position="609"/>
            <end position="643"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="704"/>
            <end position="738"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="761"/>
            <end position="805"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="723"/>
            <end position="735"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="761"/>
            <end position="772"/>
        </location>
    </feature>
    <feature type="active site" evidence="9">
        <location>
            <position position="317"/>
        </location>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="316"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <note>catalytic</note>
        </ligand>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="320"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <note>catalytic</note>
        </ligand>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="326"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <note>catalytic</note>
        </ligand>
    </feature>
    <feature type="disulfide bond" evidence="9">
        <location>
            <begin position="334"/>
            <end position="339"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="7">
        <location>
            <begin position="442"/>
            <end position="462"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="8">
        <location>
            <begin position="633"/>
            <end position="642"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00068"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00276"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50214"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50215"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015249.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="805" mass="89988" checksum="C4E01C9FAB38FA50" modified="2019-05-08" version="1">MGFQQTSHLSSYEIIHPQKLTRERRAASNSSSVQDKMSYVIHVEGKKHILHLEKNKEFLPKDFTIYTYSEEGKLQSENPDMQDHCHYQGYVEGIPDSIVAVSTCIGLRGLLQVENVTYGIEPMDSSSGLKHIIYRMENVKKEPMMCGVNNTAPEREHTEDEHPPSMTQLLRRRRAILPQTRYVELFIVVDKERYELLGRSEVAVREEMVHLANYLDSIYIMLNIRIVLVGLEIWKNENLISTDGSAGDVLGNFVQWREKNLVMRRRHDSAQFVLKKGFGGTAGMAYVGTVCSKSHAGGINVFGQISVQMFASIMAHELGHNLGMNHDDERVCHCGAESCIMNSGASGSRNFSSCSAEDFEKLTLNKGGSCLLNVPNPDETYSIPFCGNKLVDAGEECDCGSPKECESDPCCESDTCKLRAGAECAYGDCCKKCRFLPGGTVCRASTNECDLPEYCNGTSQFCQQDVTVQNGYPCQNDEAYCYNGVCQYYDAQCQAIFGSRAKAAPNSCFVEVNSKGDRFGNCGYQGHDYKKCSSWNAICGKLQCENVKTMPVFGIEPAIIKTPIRNSTCWGVDFQLGSDVPDPGMVNEGTKCNTGKICRHFQCVSVSVLNYDCDVEKKCHGHGVCNNNKNCHCDSGWAPPYCDTKGYGGSIDSGPPYNDKDTSLRDGLLVFFFLVLPLLLAAAVAFAKRRVLKQWFRRRISHCHQSHTSVPTPPPPRTETDPGNFQQNMPYTSRSVPMDMPANRFTVPSYAVNHHQPAYHQPYNYPQPYQEHQQYKIPTRPPPPQQKNIPQGHYFPSRPAPLPPM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK13</accession>
    <name>A0A452HK13_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Carboxypeptidase D</fullName>
            <ecNumber evidence="5">3.4.17.22</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="22">Metallocarboxypeptidase D</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="23">gp180</fullName>
        </alternativeName>
    </protein>
    <organism evidence="28 29">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="29" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="28" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="28" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>Releases C-terminal Arg and Lys from polypeptides.</text>
            <dbReference type="EC" id="3.4.17.22"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="2">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cell membrane</location>
            <topology evidence="3">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4 24">Belongs to the peptidase M14 family.</text>
    </comment>
    <comment type="caution">
        <text evidence="24">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="EC" id="3.4.17.22" evidence="5"/>
    <dbReference type="AlphaFoldDB" id="A0A452HK13"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017431.1">
        <property type="protein sequence ID" value="ENSGAGP00000015258.1"/>
        <property type="gene ID" value="ENSGAGG00000011436.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004181">
        <property type="term" value="F:metallocarboxypeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006518">
        <property type="term" value="P:peptide metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016485">
        <property type="term" value="P:protein processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd06245">
        <property type="entry name" value="M14_CPD_III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd11308">
        <property type="entry name" value="Peptidase_M14NE-CP-C_like"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1120:FF:000005">
        <property type="entry name" value="Carboxypeptidase D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1120:FF:000006">
        <property type="entry name" value="Carboxypeptidase D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1120:FF:000008">
        <property type="entry name" value="Carboxypeptidase D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.630.10:FF:000020">
        <property type="entry name" value="Carboxypeptidase D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.630.10:FF:000026">
        <property type="entry name" value="Carboxypeptidase D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.630.10:FF:000079">
        <property type="entry name" value="Carboxypeptidase D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1120">
        <property type="entry name" value="Carboxypeptidase-like, regulatory domain"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.630.10">
        <property type="entry name" value="Zn peptidases"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008969">
        <property type="entry name" value="CarboxyPept-like_regulatory"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057246">
        <property type="entry name" value="CARBOXYPEPT_ZN_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057247">
        <property type="entry name" value="CARBOXYPEPT_ZN_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033848">
        <property type="entry name" value="M14_CPD_III"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000834">
        <property type="entry name" value="Peptidase_M14"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050753">
        <property type="entry name" value="Peptidase_M14_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11532:SF73">
        <property type="entry name" value="CARBOXYPEPTIDASE D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11532">
        <property type="entry name" value="PROTEASE M14 CARBOXYPEPTIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13620">
        <property type="entry name" value="CarboxypepD_reg"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00246">
        <property type="entry name" value="Peptidase_M14"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00765">
        <property type="entry name" value="CRBOXYPTASEA"/>
    </dbReference>
    <dbReference type="SMART" id="SM00631">
        <property type="entry name" value="Zn_pept"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49464">
        <property type="entry name" value="Carboxypeptidase regulatory domain-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53187">
        <property type="entry name" value="Zn-dependent exopeptidases"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00132">
        <property type="entry name" value="CARBOXYPEPT_ZN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00133">
        <property type="entry name" value="CARBOXYPEPT_ZN_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS52035">
        <property type="entry name" value="PEPTIDASE_M14"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0121">Carboxypeptidase</keyword>
    <keyword evidence="7" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="20" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="14" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="21" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="18 26" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="17" id="KW-0482">Metalloprotease</keyword>
    <keyword evidence="19" id="KW-0564">Palmitate</keyword>
    <keyword evidence="9" id="KW-0645">Protease</keyword>
    <keyword evidence="29" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="13" id="KW-0677">Repeat</keyword>
    <keyword evidence="12" id="KW-0732">Signal</keyword>
    <keyword evidence="10 26" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="16 26" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="15" id="KW-0862">Zinc</keyword>
    <feature type="transmembrane region" description="Helical" evidence="26">
        <location>
            <begin position="1060"/>
            <end position="1083"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase M14" evidence="27">
        <location>
            <begin position="1"/>
            <end position="136"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase M14" evidence="27">
        <location>
            <begin position="263"/>
            <end position="553"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase M14" evidence="27">
        <location>
            <begin position="687"/>
            <end position="971"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="25">
        <location>
            <begin position="367"/>
            <end position="395"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="25">
        <location>
            <begin position="796"/>
            <end position="819"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="25">
        <location>
            <begin position="1118"/>
            <end position="1140"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="25">
        <location>
            <begin position="371"/>
            <end position="380"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="25">
        <location>
            <begin position="796"/>
            <end position="805"/>
        </location>
    </feature>
    <feature type="active site" description="Proton donor/acceptor" evidence="24">
        <location>
            <position position="106"/>
        </location>
    </feature>
    <feature type="active site" description="Proton donor/acceptor" evidence="24">
        <location>
            <position position="523"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000614"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00005988"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012811"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00014107"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022645"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023049"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023139"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00023288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00030819"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00082537"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01379"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="27">
        <source>
            <dbReference type="PROSITE" id="PS52035"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="28">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015258.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="29">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1140" mass="127050" checksum="36411EF2494726AF" modified="2019-05-08" version="1">MQRNKFVLSGNLHGGSVVASYPYDDSPMHKPNGIYSKSADDEVFKYLAKAYASNHPIMKTGTPNCPGEQFETFKDGITNGAHWYDVEGGMQDYNYVWANCFEITLELSCCKYPMASQLQQEWENNRESLLTFIEKVHMGVKGFVRDLVTGSGLANATIAVAGIDHNITAGRFGDYHRLLVPGTYNITAAVMGYVPMTVENIEVKEGAATGVDFSMQPAVLVPVPDPTPAAATPAPGSAVNATATARAASPTSLHQPLQPEDFRHHHFPDMEIFLRRYASEYTNITRLYSVGKSVEQRELYVMEISDNPGIHEPGEPEFKYVGNMHGNEVIGRELLLNLIDYLCKNFGTDPEVTELVLNTRIHIMPSMNPDGYEKSQEGDRSSTVGRNNSNNYDLNRNFPDQFLPMTDPLQPETLAVMNWLRAFPFVLSANLHGGSLVVNYPFDDDEKGIAIYSKSPDDGVFQQLALSYSKENTQMYQGSPCKDMYPNEYFPHGITNGAHWYNVQGGMQDWNYLNTNCFEVTIELGCVKYPNAEDLPKYWEENHRSLLQFIKQVHQGVKGFVLDATSRRGISNATISVADIDHPVTSYKAGDYWRLLVPGSYKITASARGYNPLTRTVTVGDRGAVQVNFTLSRTDSKVEEGKIPVLTNEDTSDPSTKEFETLIKDLSAENGLERLVLTPSVDVSPYRYRPYKDLSDFLRALHLNYPQITNLTNLGHSVEYRQMWSLEISNKPNQSEPEEPKIRFVAGIHGNAPVGTELLLALAEFLCVNYKKNAAITKLIDRTRIVIVPSLNPDGRETAQERDCTSKTGQTNAHGKDLDTDFTSNFSSYSGIGEPETKAIIENLILKRDFSLSVALDGGSLLVTYPYDKPVQTVENKETLRHLATLYANNHPSMHLGQPGCKNESDNIPGGVIRGAEWHSHLGSMKDFSVTFGHCPEITVYTSCCYFPSAGQLPNLWTENKKSLLSMVVEVHKGVHGFVQDKSGKPVSKAVIVLNEGVRVYTKEGGYFHVLLAPGLHNISATAEGYQQKHVLVQVRHDAASSVVIVFDMDNRIFGLSRELVVTVAGATMSAMVLTACIIWCVCSIKSNRHKDGFHRLRQHNDDYEDEIRMMSTGSKKSLLSHEFQDETDTEEETLYSSKH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK15</accession>
    <name>A0A452HK15_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="15">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="8">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HK15"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017427.1">
        <property type="protein sequence ID" value="ENSGAGP00000015254.1"/>
        <property type="gene ID" value="ENSGAGG00000011506.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004222">
        <property type="term" value="F:metalloendopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033631">
        <property type="term" value="P:cell-cell adhesion mediated by integrin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006509">
        <property type="term" value="P:membrane protein ectodomain proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007179">
        <property type="term" value="P:transforming growth factor beta receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd04269">
        <property type="entry name" value="ZnMc_adamalysin_II_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.390.10:FF:000002">
        <property type="entry name" value="Disintegrin and metalloproteinase domain-containing protein 22"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.70.10:FF:000001">
        <property type="entry name" value="Disintegrin and metalloproteinase domain-containing protein 22"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.390.10">
        <property type="entry name" value="Collagenase (Catalytic Domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.70.10">
        <property type="entry name" value="Disintegrin domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006586">
        <property type="entry name" value="ADAM_Cys-rich"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018358">
        <property type="entry name" value="Disintegrin_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001762">
        <property type="entry name" value="Disintegrin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036436">
        <property type="entry name" value="Disintegrin_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013111">
        <property type="entry name" value="EGF_extracell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024079">
        <property type="entry name" value="MetalloPept_cat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001590">
        <property type="entry name" value="Peptidase_M12B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002870">
        <property type="entry name" value="Peptidase_M12B_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034027">
        <property type="entry name" value="Reprolysin_adamalysin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11905">
        <property type="entry name" value="ADAM A DISINTEGRIN AND METALLOPROTEASE DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11905:SF136">
        <property type="entry name" value="DISINTEGRIN AND METALLOPROTEINASE DOMAIN-CONTAINING PROTEIN 9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08516">
        <property type="entry name" value="ADAM_CR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00200">
        <property type="entry name" value="Disintegrin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07974">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01562">
        <property type="entry name" value="Pep_M12B_propep"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01421">
        <property type="entry name" value="Reprolysin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00289">
        <property type="entry name" value="DISINTEGRIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00608">
        <property type="entry name" value="ACR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00050">
        <property type="entry name" value="DISIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57552">
        <property type="entry name" value="Blood coagulation inhibitor (disintegrin)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55486">
        <property type="entry name" value="Metalloproteases ('zincins'), catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50215">
        <property type="entry name" value="ADAM_MEPRO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00427">
        <property type="entry name" value="DISINTEGRIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50214">
        <property type="entry name" value="DISINTEGRIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01186">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6 8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="2 8" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="5 11" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 11" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 11" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="9" id="KW-0862">Zinc</keyword>
    <feature type="transmembrane region" description="Helical" evidence="11">
        <location>
            <begin position="679"/>
            <end position="698"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase M12B" evidence="14">
        <location>
            <begin position="192"/>
            <end position="386"/>
        </location>
    </feature>
    <feature type="domain" description="Disintegrin" evidence="13">
        <location>
            <begin position="394"/>
            <end position="481"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="12">
        <location>
            <begin position="620"/>
            <end position="654"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="715"/>
            <end position="749"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="772"/>
            <end position="816"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="734"/>
            <end position="746"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="772"/>
            <end position="783"/>
        </location>
    </feature>
    <feature type="active site" evidence="9">
        <location>
            <position position="328"/>
        </location>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="327"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <note>catalytic</note>
        </ligand>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="331"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <note>catalytic</note>
        </ligand>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="337"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <note>catalytic</note>
        </ligand>
    </feature>
    <feature type="disulfide bond" evidence="9">
        <location>
            <begin position="345"/>
            <end position="350"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="7">
        <location>
            <begin position="453"/>
            <end position="473"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="8">
        <location>
            <begin position="644"/>
            <end position="653"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00068"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00276"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50214"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50215"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015254.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="816" mass="91339" checksum="4FF7DDD21A36A986" modified="2019-05-08" version="1">MELFFKTELLLSSFQQTSHLSSYEIIHPQKLTRERRAASNSSSVQDKMSYVIHVEGKKHILHLEKNKEFLPKDFTIYTYSEEGKLQSENPDMQDHCHYQGYVEGIPDSIVAVSTCIGLRGLLQVENVTYGIEPMDSSSGLKHIIYRMENVKKEPMMCGVNNTAPEREHTEDEHPPSMTQLLRRRRAILPQTRYVELFIVVDKERYELLGRSEVAVREEMVHLANYLDSIYIMLNIRIVLVGLEIWKNENLISTDGSAGDVLGNFVQWREKNLVMRRRHDSAQFVLKKGFGGTAGMAYVGTVCSKSHAGGINVFGQISVQMFASIMAHELGHNLGMNHDDERVCHCGAESCIMNSGASGSRNFSSCSAEDFEKLTLNKGGSCLLNVPNPDETYSIPFCGNKLVDAGEECDCGSPKECESDPCCESDTCKLRAGAECAYGDCCKKCRFLPGGTVCRASTNECDLPEYCNGTSQFCQQDVTVQNGYPCQNDEAYCYNGVCQYYDAQCQAIFGSRAKAAPNSCFVEVNSKGDRFGNCGYQGHDYKKCSSWNAICGKLQCENVKTMPVFGIEPAIIKTPIRNSTCWGVDFQLGSDVPDPGMVNEGTKCNTGKICRHFQCVSVSVLNYDCDVEKKCHGHGVCNNNKNCHCDSGWAPPYCDTKGYGGSIDSGPPYNDKDTSLRDGLLVFFFLVLPLLLAAAVAFAKRRVLKQWFRRRISHCHQSHTSVPTPPPPRTETDPGNFQQNMPYTSRSVPMDMPANRFTVPSYAVNHHQPAYHQPYNYPQPYQEHQQYKIPTRPPPPQQKNIPQGHYFPSRPAPLPPM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HK19</accession>
    <name>A0A452HK19_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">PPM-type phosphatase domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HK19"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017432.1">
        <property type="protein sequence ID" value="ENSGAGP00000015259.1"/>
        <property type="gene ID" value="ENSGAGG00000011331.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00143">
        <property type="entry name" value="PP2Cc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.60.40.10:FF:000003">
        <property type="entry name" value="PH domain and leucine-rich repeat protein phosphatase 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.60.40.10">
        <property type="entry name" value="PPM-type phosphatase domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001611">
        <property type="entry name" value="Leu-rich_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003591">
        <property type="entry name" value="Leu-rich_rpt_typical-subtyp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050216">
        <property type="entry name" value="LRR_domain-containing"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036457">
        <property type="entry name" value="PPM-type-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001932">
        <property type="entry name" value="PPM-type_phosphatase-like_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48051">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48051:SF59">
        <property type="entry name" value="PH DOMAIN AND LEUCINE RICH REPEAT PROTEIN PHOSPHATASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13855">
        <property type="entry name" value="LRR_8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00481">
        <property type="entry name" value="PP2C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00364">
        <property type="entry name" value="LRR_BAC"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SMART" id="SM00369">
        <property type="entry name" value="LRR_TYP"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SMART" id="SM00332">
        <property type="entry name" value="PP2Cc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52058">
        <property type="entry name" value="L domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81606">
        <property type="entry name" value="PP2C-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51450">
        <property type="entry name" value="LRR"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51746">
        <property type="entry name" value="PPM_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0433">Leucine-rich repeat</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="PPM-type phosphatase" evidence="4">
        <location>
            <begin position="260"/>
            <end position="508"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="528"/>
            <end position="624"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="646"/>
            <end position="665"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="759"/>
            <end position="792"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="3">
        <location>
            <begin position="532"/>
            <end position="566"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="596"/>
            <end position="615"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="761"/>
            <end position="780"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022614"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS51746"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015259.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="792" mass="86471" checksum="139DAF14B97407EA" modified="2019-05-08" version="1">YEENWMLHLHPWTLCPPRTLSILSLRKLMVGHNHVQSLPPLLEHIPLEVLDLQHNLITKLPEMFFFKALNLRCLNVSANSLESLPSVCPGEESLSVLQLLYLTNNNLTDLCIPDLVGHPNLRVLHLANNQLQTFPASKLSKLEQLEELNLSGNKLKTIPTTIANCKQLHTLIAHSNEISIFPEILHLPRIQFVDLSCNELTEIMIPEVLPATLQELDLTGNTNLVLEHKTLDIFSHISTLKIDAKPSLTADSALSSTLWNHGLAEMAGQRNKLCVSALALGSFADSVEAVYGMFDGDKNEELPRLLQCTMADVLLEEVQQSNIDSVFMCNTFLVSHRKLGMAGQKLGSSAVLCCIRHDATDLAGSFTLTLANVGTCEAILCRSGKPLPLSKVFSLEQYPEEARRVKEQKAIITEDNKVNGVTCCTRMLGCTYLHPWILPKPHVYSIPLTVEDEMLLLGNKALWEHLSYTEAVMAVRHLHDPLAAAKKVCTLAQSYGCQDNVGAMVVCLNIGEDSCTCEVHGLPLPGPRGGISTAKPATPSSSSGTASEFSSELSASEVSSEVGSTASDEHSGAGLDAGLVPQPERRCSLHPVPPSSIFQRQPSCATFSSNQSDNGLDSDDEQPVEGVMINGSKVEVEVDIHCCQGKSLEPVPPEAEEGSAASSEDDSGGLFFRIWRQNSMTSNILLAGTKERCELQKSPSTCCLYGKKLSNGSIVPLEESLNLIEVATEVPKKKTGYFAAPAQLEPEDRFVVPPDLEEEMKEQIKQHHESGAEQKPREKAAVAPPDEFDTAL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMM2</accession>
    <name>A0A452IMM2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">DNA polymerase epsilon catalytic subunit</fullName>
            <ecNumber evidence="15">2.7.7.7</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">DNA polymerase II participates in chromosomal DNA replication.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>DNA(n) + a 2'-deoxyribonucleoside 5'-triphosphate = DNA(n+1) + diphosphate</text>
            <dbReference type="Rhea" id="RHEA:22508"/>
            <dbReference type="Rhea" id="RHEA-COMP:17339"/>
            <dbReference type="Rhea" id="RHEA-COMP:17340"/>
            <dbReference type="ChEBI" id="CHEBI:33019"/>
            <dbReference type="ChEBI" id="CHEBI:61560"/>
            <dbReference type="ChEBI" id="CHEBI:173112"/>
            <dbReference type="EC" id="2.7.7.7"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="15">
            <name>[4Fe-4S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:49883"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 15">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 15">Belongs to the DNA polymerase type-B family.</text>
    </comment>
    <dbReference type="EC" id="2.7.7.7" evidence="15"/>
    <dbReference type="AlphaFoldDB" id="A0A452IMM2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033158.1">
        <property type="protein sequence ID" value="ENSGAGP00000029195.1"/>
        <property type="gene ID" value="ENSGAGG00000020939.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008622">
        <property type="term" value="C:epsilon DNA polymerase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051539">
        <property type="term" value="F:4 iron, 4 sulfur cluster binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003887">
        <property type="term" value="F:DNA-directed DNA polymerase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000166">
        <property type="term" value="F:nucleotide binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008310">
        <property type="term" value="F:single-stranded DNA 3'-5' DNA exonuclease activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006287">
        <property type="term" value="P:base-excision repair, gap-filling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045004">
        <property type="term" value="P:DNA replication proofreading"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006272">
        <property type="term" value="P:leading strand elongation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000278">
        <property type="term" value="P:mitotic cell cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006297">
        <property type="term" value="P:nucleotide-excision repair, DNA gap filling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05535">
        <property type="entry name" value="POLBc_epsilon"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.690:FF:000005">
        <property type="entry name" value="DNA polymerase epsilon catalytic subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.420.10:FF:000217">
        <property type="entry name" value="DNA polymerase epsilon catalytic subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1600.10:FF:000006">
        <property type="entry name" value="DNA polymerase epsilon catalytic subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.132.60">
        <property type="entry name" value="DNA polymerase family B, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1600.10">
        <property type="entry name" value="Palm domain of DNA polymerase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.420.10">
        <property type="entry name" value="Ribonuclease H-like superfamily/Ribonuclease H"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006172">
        <property type="entry name" value="DNA-dir_DNA_pol_B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006133">
        <property type="entry name" value="DNA-dir_DNA_pol_B_exonuc"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006134">
        <property type="entry name" value="DNA-dir_DNA_pol_B_multi_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043502">
        <property type="entry name" value="DNA/RNA_pol_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042087">
        <property type="entry name" value="DNA_pol_B_thumb"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023211">
        <property type="entry name" value="DNA_pol_palm_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029703">
        <property type="entry name" value="POL2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055191">
        <property type="entry name" value="POL2_thumb"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012337">
        <property type="entry name" value="RNaseH-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036397">
        <property type="entry name" value="RNaseH_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10670">
        <property type="entry name" value="DNA POLYMERASE EPSILON CATALYTIC SUBUNIT A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10670:SF0">
        <property type="entry name" value="DNA POLYMERASE EPSILON CATALYTIC SUBUNIT A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00136">
        <property type="entry name" value="DNA_pol_B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03104">
        <property type="entry name" value="DNA_pol_B_exo1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22634">
        <property type="entry name" value="POL2_thumb"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00486">
        <property type="entry name" value="POLBc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56672">
        <property type="entry name" value="DNA/RNA polymerases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53098">
        <property type="entry name" value="Ribonuclease H-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 15" id="KW-0004">4Fe-4S</keyword>
    <keyword evidence="6 15" id="KW-0235">DNA replication</keyword>
    <keyword evidence="13 15" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="10 15" id="KW-0239">DNA-directed DNA polymerase</keyword>
    <keyword evidence="11 15" id="KW-0408">Iron</keyword>
    <keyword evidence="12 15" id="KW-0411">Iron-sulfur</keyword>
    <keyword evidence="7 15" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="5 15" id="KW-0548">Nucleotidyltransferase</keyword>
    <keyword evidence="14 15" id="KW-0539">Nucleus</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 15" id="KW-0808">Transferase</keyword>
    <keyword evidence="9 15" id="KW-0862">Zinc</keyword>
    <keyword evidence="8 15" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="DNA-directed DNA polymerase family B exonuclease" evidence="17">
        <location>
            <begin position="16"/>
            <end position="64"/>
        </location>
    </feature>
    <feature type="domain" description="DNA-directed DNA polymerase family B multifunctional" evidence="16">
        <location>
            <begin position="433"/>
            <end position="491"/>
        </location>
    </feature>
    <feature type="domain" description="DNA polymerase epsilon ,catalytic subunit A thumb" evidence="18">
        <location>
            <begin position="610"/>
            <end position="657"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005755"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022485"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022695"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022705"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022932"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023014"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="RuleBase" id="RU365029"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF00136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF03104"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF22634"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029195.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="687" mass="78546" checksum="C72B94682FE78175" modified="2019-05-08" version="1">MLRLLPRPFVEARAAVHGMNMYQEIGFQKDSQGEYKASQCIHMDCLRWVKRDSYLPVGSHNLKAAAKAKLGYDPVELDPEEMCRMATEEPQTLATYSVSDAVATYYMYMKYVHPFIFALCTIIPMEPDEVLRKGSGTLCEALLMVQAYHANIIFPNKQEQEFNKLTEDGHVLDSETYVGGHVEALESGVFRSDIPCRFKMNPVAFDFLLQRVEKTLRHAIEEEEGLPLDQVTNFQEVCDEIKVKLNSLKDIPNRIECPLIYHLDVGAMYPNIILTNRLQPSAMVDEATCAACDFNKPGANCQRRMTWQWRGEFMPASRSEYHRIQQQLESEKLPPLYPDGPPRAFHELSREEQAKFEKKRLADYCRKAYKKLHVTKVEERITTICQRENSFYVDTVRAFRDRRYEFKGLHKVWKKKLSAATEMGDATEVKRCKNMEILYDSLQLAHKCILNSFYGYVMRKGARWYSMEMAGIVCFTGANIITQARELIEQIGRPLELDTDGIWCVLPNSFPENFVIKSTNAKKPKVTISYPGAMLNILVKEGFTNDQYQELVDPASLTYVTRSENSIFFEVDGPYLPWLQDRPAGAVSPRYAVFNEDGSLAELKGFEVKRRGELQLVKIFQSSVFESFLKGTTLEEVYASVAKVADYWLDVLYSKVSAQPLASLCCSVGPGVEMGATPWMGGQDMHW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMM5</accession>
    <name>A0A452IMM5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Carbamoyl-phosphate synthase [ammonia], mitochondrial</fullName>
            <ecNumber evidence="10">6.3.4.16</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="14">Carbamoyl-phosphate synthetase I</fullName>
        </alternativeName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="12">Involved in the urea cycle of ureotelic animals where the enzyme plays an important role in removing excess ammonia from the cell.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>hydrogencarbonate + NH4(+) + 2 ATP = carbamoyl phosphate + 2 ADP + phosphate + 2 H(+)</text>
            <dbReference type="Rhea" id="RHEA:18029"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:17544"/>
            <dbReference type="ChEBI" id="CHEBI:28938"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:58228"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="6.3.4.16"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Mitochondrion</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="6.3.4.16" evidence="10"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033317.1">
        <property type="protein sequence ID" value="ENSGAGP00000029337.1"/>
        <property type="gene ID" value="ENSGAGG00000021129.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004087">
        <property type="term" value="F:carbamoyl-phosphate synthase (ammonia) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004088">
        <property type="term" value="F:carbamoyl-phosphate synthase (glutamine-hydrolyzing) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006207">
        <property type="term" value="P:'de novo' pyrimidine nucleobase biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006541">
        <property type="term" value="P:glutamine metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000050">
        <property type="term" value="P:urea cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd01744">
        <property type="entry name" value="GATase1_CPSase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd01423">
        <property type="entry name" value="MGS_CPS_I_III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.470.20:FF:000030">
        <property type="entry name" value="Carbamoyl-phosphate synthase 1, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.20:FF:000012">
        <property type="entry name" value="Carbamoyl-phosphate synthase 1, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.880:FF:000006">
        <property type="entry name" value="Carbamoyl-phosphate synthase 1, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1380:FF:000010">
        <property type="entry name" value="carbamoyl-phosphate synthase [ammonia], mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.1030.10:FF:000001">
        <property type="entry name" value="Carbamoyl-phosphate synthase large chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1490.20:FF:000001">
        <property type="entry name" value="Carbamoyl-phosphate synthase large chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.470.20:FF:000001">
        <property type="entry name" value="Carbamoyl-phosphate synthase large chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.20:FF:000002">
        <property type="entry name" value="Carbamoyl-phosphate synthase large chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.20">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.880">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1490.20">
        <property type="entry name" value="ATP-grasp fold, A domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.470.20">
        <property type="entry name" value="ATP-grasp fold, B domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.50.30.20">
        <property type="entry name" value="Carbamoyl-phosphate synthase small subunit, N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1030.10">
        <property type="entry name" value="Carbamoyl-phosphate synthetase, large subunit oligomerisation domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1380">
        <property type="entry name" value="Methylglyoxal synthase-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011761">
        <property type="entry name" value="ATP-grasp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013815">
        <property type="entry name" value="ATP_grasp_subdomain_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036897">
        <property type="entry name" value="CarbamoylP_synth_lsu_oligo_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006274">
        <property type="entry name" value="CarbamoylP_synth_ssu"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002474">
        <property type="entry name" value="CarbamoylP_synth_ssu_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036480">
        <property type="entry name" value="CarbP_synth_ssu_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029062">
        <property type="entry name" value="Class_I_gatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005479">
        <property type="entry name" value="CPAse_ATP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005483">
        <property type="entry name" value="CPSase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035686">
        <property type="entry name" value="CPSase_GATase1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006275">
        <property type="entry name" value="CPSase_lsu"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005480">
        <property type="entry name" value="CPSase_lsu_oligo"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058047">
        <property type="entry name" value="CPSase_preATP-grasp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017926">
        <property type="entry name" value="GATASE"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011607">
        <property type="entry name" value="MGS-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036914">
        <property type="entry name" value="MGS-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016185">
        <property type="entry name" value="PreATP-grasp_dom_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01369">
        <property type="entry name" value="CPSaseII_lrg"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01368">
        <property type="entry name" value="CPSaseIIsmall"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF003671">
        <property type="entry name" value="PRK05294.1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF009455">
        <property type="entry name" value="PRK12815.1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF009475">
        <property type="entry name" value="PRK12838.1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11405:SF53">
        <property type="entry name" value="CARBAMOYL-PHOSPHATE SYNTHASE [AMMONIA], MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11405">
        <property type="entry name" value="CARBAMOYLTRANSFERASE FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25596">
        <property type="entry name" value="CPSase_L_D1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02786">
        <property type="entry name" value="CPSase_L_D2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02787">
        <property type="entry name" value="CPSase_L_D3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00988">
        <property type="entry name" value="CPSase_sm_chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00117">
        <property type="entry name" value="GATase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02142">
        <property type="entry name" value="MGS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00098">
        <property type="entry name" value="CPSASE"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00099">
        <property type="entry name" value="CPSGATASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM01096">
        <property type="entry name" value="CPSase_L_D3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01097">
        <property type="entry name" value="CPSase_sm_chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00851">
        <property type="entry name" value="MGS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48108">
        <property type="entry name" value="Carbamoyl phosphate synthetase, large subunit connection domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52021">
        <property type="entry name" value="Carbamoyl phosphate synthetase, small subunit N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52317">
        <property type="entry name" value="Class I glutamine amidotransferase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56059">
        <property type="entry name" value="Glutathione synthetase ATP-binding domain-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52335">
        <property type="entry name" value="Methylglyoxal synthase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52440">
        <property type="entry name" value="PreATP-grasp domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50975">
        <property type="entry name" value="ATP_GRASP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00866">
        <property type="entry name" value="CPSASE_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00867">
        <property type="entry name" value="CPSASE_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51273">
        <property type="entry name" value="GATASE_TYPE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51855">
        <property type="entry name" value="MGS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0021">Allosteric enzyme</keyword>
    <keyword evidence="7 15" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="4" id="KW-0436">Ligase</keyword>
    <keyword evidence="9" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="6 15" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="8" id="KW-0809">Transit peptide</keyword>
    <keyword evidence="2" id="KW-0835">Urea cycle</keyword>
    <feature type="domain" description="ATP-grasp" evidence="16">
        <location>
            <begin position="506"/>
            <end position="698"/>
        </location>
    </feature>
    <feature type="domain" description="ATP-grasp" evidence="16">
        <location>
            <begin position="1048"/>
            <end position="1239"/>
        </location>
    </feature>
    <feature type="domain" description="MGS-like" evidence="17">
        <location>
            <begin position="1310"/>
            <end position="1455"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004173"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022436"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022533"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022598"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00044063"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00047359"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00059545"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00074238"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00082675"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00409"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50975"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS51855"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029337.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1455" mass="159630" checksum="B28B427FA4E7E7E0" modified="2019-05-08" version="1">TAQTANLVLDDGTKMKGYSFGHPSSAAGEVVFNTGLAGYTETLTDPSYRGQILTMVNPIVGNGGVPDTAALDEMGLSKFLESDGIKVSGLLVLDYSNDYSHWRAVKTLGQWLQEEKVRDLWATRPDTRHGISILRKLLSDSLSNGNQEREILVYSELELLSSDIWFEVKVYGKGNPIKVVAVDCGLKNNAIRLLVKRGAEVHLVPWNHDFTGMEYDGLFLSGGPGDPMQAQELIQNLQKVQRSDRKEPLFGVSMGNLITGIAAGATSYKMPMANRGQNQPVMNLSNGQAFITAQNHGYALQSNTLPAGWKPLFVNVNDQTNEGIMHETKPIFTAQFNPEASAGPTDTEFLFDSFISLIKKGKGTTVASVLPKARTVVSKVLILGSGGLSIGQAGEFDYSGSQAVKAMKEENVKTVLMNPNIASVQTNEVGLKQADTVYFLPITPQFVTEVIKAERPDGLILGMGGQTALNCGVELFRRGVLKEYGVKVLGTSVEAIMATEDRQLFSDKLTEINEKIAPSFAVESIEDALNAAEKIGYPVMIRSAYALGGLGSGICPDKERLLDLGTKAFAVTNQILVERSVVGWKEIEYEVVRDASDNCITVCNMENVDAMGVHTGDSIVVAPSQTLSNEEFQMLRESAIKVVRHLGIVGECNIQYALHPTSLEYCIIEVNARLSRSSALASKATGYPLAFIAAKIALGIPLPEIKNVVSGKTSACFEPSLDYMVAKIPRWDLDRFQGTSSQIGSSMKSVGEVMAIGRTFEECFQKALRMCHPSVDGFTSRLPMNKEWPANLDLQKELSDPSSIRVYAIAKALHDGVPVDDIHKLTAIDRWFIYKMRDIVNMEKSLKELSSESIPEETMRRAKQIGFSDKQIGKCLGVTEAQSRELRLKKNIKPSVKQIDTLAAEYPAITNYLYVTYNGQENDVKFDDHGMMVLGCGPYHIGSSVEFDWCAVSSIRTLRQLGHKTVVVNCNPETVSTDFDECDRLYFEELSLERILDIYQHEKCAGCIISVGGQIPNNLAVPLYKNGVKIMGTNPMQIDRAEDRSVFSAVLDELNVAQAPWKAVSTLKDAVEFANSVGYPCLLRPSYVLSGSAMNVVFTEEEMKKFLAEATRVSQEHPVVLTKFIEGAREVEMDAVAKAGKVVSHAISEHVEDAGVHSGDATLMLPTQNISQGALEKVKIITKKIARAFSISGPFNVQFLVRGNDVLVIECNLRASRSFPFVSKTIGVDFIDVATKVMIGEKIDESSLPTLERPIIPSEYVGIKAPMFSWPRLRDADPILRCEMASTGEVACFGESIHSAFLKAMLSTGFKFPQKGILVGIQHSFRPKFLGVAEQLHEEGFKLYATEATSNWLNANSVPTTPVAWPSQESHTTSLPAIRRLIKDGNIDLVINLPNSNTKFVHDNYVIRRTAVDSGIALLTNFQVTKLFAEAIKYSGKVDAKSLFHYRKLNTSNPP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMM7</accession>
    <name>A0A452IMM7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">DNA polymerase zeta catalytic subunit</fullName>
            <ecNumber evidence="4">2.7.7.7</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="19">Protein reversionless 3-like</fullName>
        </alternativeName>
    </protein>
    <organism evidence="26 27">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="27" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="26" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="26" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="17">Catalytic subunit of the DNA polymerase zeta complex, an error-prone polymerase specialized in translesion DNA synthesis (TLS). Lacks an intrinsic 3'-5' exonuclease activity and thus has no proofreading function.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="28">
            <text>DNA(n) + a 2'-deoxyribonucleoside 5'-triphosphate = DNA(n+1) + diphosphate</text>
            <dbReference type="Rhea" id="RHEA:22508"/>
            <dbReference type="Rhea" id="RHEA-COMP:17339"/>
            <dbReference type="Rhea" id="RHEA-COMP:17340"/>
            <dbReference type="ChEBI" id="CHEBI:33019"/>
            <dbReference type="ChEBI" id="CHEBI:61560"/>
            <dbReference type="ChEBI" id="CHEBI:173112"/>
            <dbReference type="EC" id="2.7.7.7"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>[4Fe-4S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:49883"/>
        </cofactor>
    </comment>
    <comment type="subunit">
        <text evidence="18">Heterodimer with MAD2L2. This dimer forms the minimal DNA polymerase zeta complex (Pol-zeta2), with REV3L bearing DNA polymerase catalytic activity, although its activity is very low in this context. Component of the tetrameric Pol-zeta complex (Pol-zeta4), which consists of REV3L, MAD2L2, POLD2 and POLD3; Pol-zeta4 is the fully active form of DNA polymerase zeta.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the DNA polymerase type-B family.</text>
    </comment>
    <dbReference type="EC" id="2.7.7.7" evidence="4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033089.1">
        <property type="protein sequence ID" value="ENSGAGP00000029132.1"/>
        <property type="gene ID" value="ENSGAGG00000021078.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031981">
        <property type="term" value="C:nuclear lumen"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016035">
        <property type="term" value="C:zeta DNA polymerase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003887">
        <property type="term" value="F:DNA-directed DNA polymerase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051536">
        <property type="term" value="F:iron-sulfur cluster binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000166">
        <property type="term" value="F:nucleotide binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000724">
        <property type="term" value="P:double-strand break repair via homologous recombination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042276">
        <property type="term" value="P:error-prone translesion synthesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05778">
        <property type="entry name" value="DNA_polB_zeta_exo"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05534">
        <property type="entry name" value="POLBc_zeta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd22287">
        <property type="entry name" value="REV3L_RBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.690:FF:000002">
        <property type="entry name" value="DNA polymerase zeta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.420.10:FF:000024">
        <property type="entry name" value="DNA polymerase zeta catalytic subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.342.10:FF:000002">
        <property type="entry name" value="DNA polymerase zeta catalytic subunit isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.132.60:FF:000005">
        <property type="entry name" value="Putative DNA polymerase zeta catalytic subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.132.60">
        <property type="entry name" value="DNA polymerase family B, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.342.10">
        <property type="entry name" value="DNA Polymerase, chain B, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.690">
        <property type="entry name" value="Helix hairpin bin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1600.10">
        <property type="entry name" value="Palm domain of DNA polymerase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.420.10">
        <property type="entry name" value="Ribonuclease H-like superfamily/Ribonuclease H"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006172">
        <property type="entry name" value="DNA-dir_DNA_pol_B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017964">
        <property type="entry name" value="DNA-dir_DNA_pol_B_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006133">
        <property type="entry name" value="DNA-dir_DNA_pol_B_exonuc"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006134">
        <property type="entry name" value="DNA-dir_DNA_pol_B_multi_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043502">
        <property type="entry name" value="DNA/RNA_pol_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042087">
        <property type="entry name" value="DNA_pol_B_thumb"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023211">
        <property type="entry name" value="DNA_pol_palm_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056435">
        <property type="entry name" value="DPOD/Z_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032757">
        <property type="entry name" value="DUF4683"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR030559">
        <property type="entry name" value="PolZ_Rev3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012337">
        <property type="entry name" value="RNaseH-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036397">
        <property type="entry name" value="RNaseH_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025687">
        <property type="entry name" value="Znf-C4pol"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45812">
        <property type="entry name" value="DNA POLYMERASE ZETA CATALYTIC SUBUNIT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45812:SF1">
        <property type="entry name" value="DNA POLYMERASE ZETA CATALYTIC SUBUNIT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00136">
        <property type="entry name" value="DNA_pol_B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03104">
        <property type="entry name" value="DNA_pol_B_exo1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15735">
        <property type="entry name" value="DUF4683"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24055">
        <property type="entry name" value="POL3_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14260">
        <property type="entry name" value="zf-C4pol"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00106">
        <property type="entry name" value="DNAPOLB"/>
    </dbReference>
    <dbReference type="SMART" id="SM00486">
        <property type="entry name" value="POLBc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56672">
        <property type="entry name" value="DNA/RNA polymerases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53098">
        <property type="entry name" value="Ribonuclease H-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00116">
        <property type="entry name" value="DNA_POLYMERASE_B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0227">DNA damage</keyword>
    <keyword evidence="14" id="KW-0234">DNA repair</keyword>
    <keyword evidence="11" id="KW-0239">DNA-directed DNA polymerase</keyword>
    <keyword evidence="12" id="KW-0408">Iron</keyword>
    <keyword evidence="13" id="KW-0411">Iron-sulfur</keyword>
    <keyword evidence="8" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7" id="KW-0548">Nucleotidyltransferase</keyword>
    <keyword evidence="15" id="KW-0539">Nucleus</keyword>
    <keyword evidence="27" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0808">Transferase</keyword>
    <keyword evidence="10" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="DNA polymerase delta/zeta catalytic subunit N-terminal" evidence="25">
        <location>
            <begin position="44"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="domain" description="DUF4683" evidence="24">
        <location>
            <begin position="733"/>
            <end position="1124"/>
        </location>
    </feature>
    <feature type="domain" description="DNA-directed DNA polymerase family B exonuclease" evidence="22">
        <location>
            <begin position="2257"/>
            <end position="2507"/>
        </location>
    </feature>
    <feature type="domain" description="DNA-directed DNA polymerase family B multifunctional" evidence="21">
        <location>
            <begin position="2573"/>
            <end position="3023"/>
        </location>
    </feature>
    <feature type="domain" description="C4-type zinc-finger of DNA polymerase delta" evidence="23">
        <location>
            <begin position="3065"/>
            <end position="3132"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="254"/>
            <end position="283"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="418"/>
            <end position="446"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="461"/>
            <end position="499"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="1027"/>
            <end position="1064"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="1182"/>
            <end position="1204"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="1556"/>
            <end position="1607"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="1698"/>
            <end position="1718"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="1734"/>
            <end position="1753"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="1846"/>
            <end position="1875"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="1962"/>
            <end position="1989"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="2136"/>
            <end position="2199"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="20">
        <location>
            <begin position="254"/>
            <end position="265"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="20">
        <location>
            <begin position="422"/>
            <end position="437"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="20">
        <location>
            <begin position="464"/>
            <end position="474"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="20">
        <location>
            <begin position="486"/>
            <end position="497"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="20">
        <location>
            <begin position="1035"/>
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        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="20">
        <location>
            <begin position="1700"/>
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        <location>
            <begin position="1742"/>
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        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="20">
        <location>
            <begin position="1851"/>
            <end position="1867"/>
        </location>
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            <begin position="2153"/>
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        </location>
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    <evidence type="ECO:0000256" key="1">
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            <dbReference type="ARBA" id="ARBA00001966"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
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        </source>
    </evidence>
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    </evidence>
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    </evidence>
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    </evidence>
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            <dbReference type="ARBA" id="ARBA00022695"/>
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    </evidence>
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    </evidence>
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    </evidence>
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    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
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    </evidence>
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    </evidence>
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    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00075683"/>
        </source>
    </evidence>
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            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
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            <dbReference type="Pfam" id="PF00136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
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            <dbReference type="Pfam" id="PF03104"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
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            <dbReference type="Pfam" id="PF14260"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="Pfam" id="PF15735"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="Pfam" id="PF24055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029132.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="3153" mass="354734" checksum="3CE678C9DF3EA3A1" modified="2019-05-08" version="1">MILGVSLGGLAAPCLFPTEEGCSLGLLAGRKEVSGQKTCLHLHGIFPYLYVPYDGYGKQSERYLRQVAFSIDRALNVALGNPSSTVQHVFKVSLVSGMPFYGYHEKEREFMKIYLYNPSMVKRVCELLQGGAIMNKCYQPHEAHIPYLLQLFIDYNLYGMNLINLAAVKFRKAKRRSDTLGGIGCYKDHLPENSITGTFFRWEEDEIPSSLILDGVEPQSTCELEVDAVAADILNRLDVEAQIGRNPGLQAIWEDEKQRRREKNESSQISPSDSQDRGFVPTTESEKIFQKRLKEILKQNDFSVTLSGSVDYSNGSEEFSAELTLHSEVLSPKVVPCSPANMVEVHKDKLNTEPNRDTNKEEDVLINEEAILNIMENSQCFHPLSQRLNQTAVFMDSSSDQAMIHLLAGLEDDGYQMGGHRTLSQNHSLGSNRNLQNSDDEENEPQIEKEEMELSLLMSQRWDSSNEEHGEKRRSLSKNTHRSSTEEEDSSSEEEMEWSGNSLLLANLSIPQLDGTAEENGDNPLNNESSRTHSSIIAASKMSVKTSMFHRDATTLEPPSSAKITFQCKHTSALSSHVLNNEDLVEELSQPNTEVRPDPSVHSFTKESTYAAKCPTSFSNSAHSENSHKESNKDILPVSSCENNIFDYEEDISSVSRQIPSRKYTSIRKVEKDDASFMHMSCHISDSVLSKNSFSFSELSLSKNKIPSEANEKSSSTGINSFFPSTVTESCELVSCSGGNRTAGHCVERSTDEEGSLNKLKIRFEEFQEHKTEKPSLSQQAAHYMFFPSVVLSNCLSRPQKLAPVTYKLQQGNRQSRLKLNKKKLGLGNHQEPAGVSDIASTKESCNTQDDACSIIPGKDSALPSDLVQVPHGAFENKMPTYNANCIDCQFRDGSLETEQSFGLCGNKYTLRTKRKVNYETEDNESSFFMHNSKISLPQPIESVESLDGSQKSRKRRKLSKKLPPVIIKYIIINRFRGRKNMLVKIGKIDSSEEQVILTEEKMNLYKKLAPLKDFWPNVPDSPATKYPIYPLTPKKSHRRKAKHKSAKKKVGKPQKASSKNIKRTLSFRKRTHAILSPPSPSYNAEAEDCDYHYSDVMSKLGFLSERSISPVNASPPRCWSPTDPKAEEIINGLDKEALLIKGPNMYISKTVNSIVGKNSRAKTQVKTCKKKFATVSTKKRNKINQTNKTANDGKKKPRTKPRQKIAEKISRKHMTVKDEKGNICSAAEVKFVLKHQDLPEITHNFGSSQPFLTQKDVPLTGYSTGHLPSTQLPSTADVQGNSSCYFHPLLETDKSVGLQGLSTPRENLHSSVISAPVIPGREITKISSQRSKSQNAVFRMKESSLIQNNIFDPSNHSSQVTHNVCISKDKTATKTKEIINSQNRYLSSVGKLNETHSFLDTMKTDQLHTTNFLHCKDSNQQQIVCLPEASKHADPYSSILKSSEEGHGSLQLPKNCFVTSLRSPVKQLSWEQKQRGFIFDMSHFKPERIKHRSLSETISQTKTISQCENRTVMTPSAFSEGQSGLAVLKELLQKRQQKAQHAGILQEPLSVSPQINKSISCSPEQHKAGKRSQSVTSPRKSRTPRNTKPKEKTPKPLKVASSNQHNTNQIDHFISDDSPIFFSDPGFESCYSLEDSLSPDHNYNFDINTIGQTGFCSLYSASQFVPADQSLPQKFLSDAVQDLFPGQTAENELFSHNNQKSEEEKHHSSDPSTWITSGALSPELFERTSLDNNENHRHGQWRNNVHPSASRTSSSVDIFCTQQAGDYVNEKFKLNRNPVNKEVFLSLPQSSCSDFIQSCTRKETTFEPCQPLGSINTSFTSVLSSPDGELMDAASEDLELYVSRNNEGLTPTPDSSPRSTSSPSQSKNGSFTPRTAHILKPLMSPPSREEIMATLLDHDLAETIYQEPFCSNPSDAPEKPREIGGRLLTVETRLPNDLPEFEGDFSLEGLRLWKTAFSAMTQSPRPGSPPRNCHSIAGKGEINSHKTSEDKKIVVMPCKCAPSRQQVQMWLHAKEEYEHIKKLPKAQPVDTEKATENFRSIALPIDQPTEVVKAAKNFSLPSLQDSMPIVPTKDAPVSQADPTPTQTKETCDKYINTVQISTNTIEPSKTPPESRTLLLAPSAIELDKEEEDDDYYVNYSSPDSPVLPPWQQAASPDSKPSSLEDADWQNQNIISSPVEENDTAHSENVQKHVKDEGRASFSTSPLLVREGAGNSESVCLHSTPIVQRKCQERIAEALDFTPLSTEPKMQKLNHRRGSNSDTLRRVLLTTQVKNQFAALNAPKKETSQIEGPSLNNSYGFKVSVQNLQEAKALHEVQNLTLISMELHARTRRDLEPDPEFDPICALFFCISSDAALPNTDKREITGAIVIDKDRTISSQGSRDQAPLLTRSGITGLEVIYAADERALFQEVVNIVKRYDPDMMLGYEVQMHSWGYLLQRAAALSVDLCQMISRVPDDKKENRFAAEQDEYGSDTMSEIHIVGRIVLNLWRMMRNEVTLTNYTFENVGFHVLHQRFPLFTSRVLSDWFDNKADIYRWKMVDHYISRVRGNLQMLEQLDLIGRTSEMARLFGIQFLHVLTRGSQYRVESMMLRIAKPMNYIPVTPSVQQRAQMKAPQCIPLIMEPESRFYSNTVLVLDFQSLYPSIVIAYNYCFSTCLGHIENLGKFDEFKFGCTSLRVPPDLLYQIRHDITVSPNGVAFVKPSVRKGVLPRMLEEILRTRIMVKQSIKAYKHDKAVTRMLEARQMGLKLIANVTFGYTSANFSGRMPCMEIGDSIVHKARETLERAIKLVNDTKKWGARVVYGDTDSMFVLLKGATKEQSFKIGQEIAEAVTATNPKPVKLKFEKVYLPCVLQTKKRYVGYMYEALDQKDPVFDAKGIETVRRDSCPAVSKILERSIKLLFETRDISQIKQYVQNQCMKLLEGKASMQDFIFAKEYRGSSAYRPGACVPALEITRRMLAYDRRSEPRVGERVPYVIVYGMPGLPLIQLVRRPIEVLQDTNLRLNATYYITKQILPPLARIFSLIGIDVFSWYHELPRIQRAASAARTELEGRKGTISQYFTTLHCPVCDELTQHGICSKCRSQPQHVAVILSQEIRELEHKQEQLVKICKNCTGCFDRQIQCVSLNCPVLFKLSRVSRELSKAPYLRQLLDQF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMP7</accession>
    <name>A0A452IMP7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Methylcrotonoyl-CoA carboxylase beta chain, mitochondrial</fullName>
            <ecNumber evidence="3">6.4.1.4</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="6">3-methylcrotonyl-CoA carboxylase 2</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="4">3-methylcrotonyl-CoA carboxylase non-biotin-containing subunit</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="5">3-methylcrotonyl-CoA:carbon dioxide ligase subunit beta</fullName>
        </alternativeName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="8">Carboxyltransferase subunit of the 3-methylcrotonyl-CoA carboxylase, an enzyme that catalyzes the conversion of 3-methylcrotonyl-CoA to 3-methylglutaconyl-CoA, a critical step for leucine and isovaleric acid catabolism.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="7">
            <text>3-methylbut-2-enoyl-CoA + hydrogencarbonate + ATP = 3-methyl-(2E)-glutaconyl-CoA + ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:13589"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:17544"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:57344"/>
            <dbReference type="ChEBI" id="CHEBI:57346"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="6.4.1.4"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="2">Amino-acid degradation; L-leucine degradation; (S)-3-hydroxy-3-methylglutaryl-CoA from 3-isovaleryl-CoA: step 2/3.</text>
    </comment>
    <comment type="subunit">
        <text evidence="9">Probably a dodecamer composed of six biotin-containing alpha subunits (MCCC1) and six beta (MCCC2) subunits.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the AccD/PCCB family.</text>
    </comment>
    <dbReference type="EC" id="6.4.1.4" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452IMP7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033069.1">
        <property type="protein sequence ID" value="ENSGAGP00000029113.1"/>
        <property type="gene ID" value="ENSGAGG00000021062.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00363">
        <property type="reaction ID" value="UER00861"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:1905202">
        <property type="term" value="C:methylcrotonoyl-CoA carboxylase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004485">
        <property type="term" value="F:methylcrotonoyl-CoA carboxylase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006552">
        <property type="term" value="P:L-leucine catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.226.10:FF:000004">
        <property type="entry name" value="Methylcrotonoyl-CoA carboxylase beta chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.226.10:FF:000007">
        <property type="entry name" value="Methylcrotonoyl-CoA carboxylase subunit beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.226.10">
        <property type="entry name" value="2-enoyl-CoA Hydratase, Chain A, domain 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034733">
        <property type="entry name" value="AcCoA_carboxyl_beta"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029045">
        <property type="entry name" value="ClpP/crotonase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011763">
        <property type="entry name" value="COA_CT_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011762">
        <property type="entry name" value="COA_CT_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045190">
        <property type="entry name" value="MCCB/AccD1-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22855">
        <property type="entry name" value="ACETYL, PROPIONYL, PYRUVATE, AND GLUTACONYL CARBOXYLASE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22855:SF13">
        <property type="entry name" value="METHYLCROTONOYL-COA CARBOXYLASE BETA CHAIN, MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01039">
        <property type="entry name" value="Carboxyl_trans"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52096">
        <property type="entry name" value="ClpP/crotonase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50989">
        <property type="entry name" value="COA_CT_CTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50980">
        <property type="entry name" value="COA_CT_NTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="CoA carboxyltransferase N-terminal" evidence="11">
        <location>
            <begin position="50"/>
            <end position="307"/>
        </location>
    </feature>
    <feature type="domain" description="CoA carboxyltransferase C-terminal" evidence="12">
        <location>
            <begin position="307"/>
            <end position="556"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006102"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00025711"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00026116"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00031109"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00031237"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00031404"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00052347"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00053988"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00062256"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00073673"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50980"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029113.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="564" mass="61267" checksum="BEE5DA414317CB26" modified="2019-05-08" version="1">MLAGLKWRLGLRLCRPAATLARAYHGDRVAAVGSQPDAGSPLYQENYERMKALVTELQELAEKIRLGGGEKARKLHTSRGKLLPRERIDRLIDPGSPFLEFSQFAGYQLYGNEEVPAGGIITGIGRVSGVECLIVANDATVKGGTYYPVTVKKHLRAQEIAMQNHLPCLYLVDSGGANLPRQAEVFPDRDHFGRIFYNQALMSSQRIPQIAVVMGSCTAGGAYVPAMADESIIVGKQGTIFLGGPPLVKAATGEEVSAEDLGGADLHCRKSGVTDHYALDDNHALHLARKVVRSLNYQKKVDVTVEPPEEPLYPADELYGIVGDRLKRNFDVKEVIARIVDGSKFDEFKALYGDTLITGFARLFGYPVGIIGNNGVLFSESAKKGTHFIQLCCQRNIPIVFLQNITGFMVGRDYEAGGIAKDGAKMVTAVACAGVPKITVIIGGSYGAGNYGMCGRAYSPRFLYMWPNARISVMGGEQAATVLATVAKDQKAREGKQLSEADEAALKEPIIRRFEEEGNPYYSSARLWDDGIIDPADTRLVLGLSLSAALNAPTKRTEFGVFRM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMP9</accession>
    <name>A0A452IMP9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="11">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="domain">
        <text evidence="6">The CKK domain binds microtubules.</text>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the CAMSAP1 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000033337.1">
        <property type="protein sequence ID" value="ENSGAGP00000029357.1"/>
        <property type="gene ID" value="ENSGAGG00000021226.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036449">
        <property type="term" value="C:microtubule minus-end"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051011">
        <property type="term" value="F:microtubule minus-end binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030507">
        <property type="term" value="F:spectrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031122">
        <property type="term" value="P:cytoplasmic microtubule organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007026">
        <property type="term" value="P:negative regulation of microtubule depolymerization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031175">
        <property type="term" value="P:neuron projection development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.20.360:FF:000001">
        <property type="entry name" value="Calmodulin-regulated spectrin-associated protein 3 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.20.360">
        <property type="entry name" value="CKK domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032940">
        <property type="entry name" value="CAMSAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031372">
        <property type="entry name" value="CAMSAP_CC1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058042">
        <property type="entry name" value="CAMSAP_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022613">
        <property type="entry name" value="CH_CAMSAP_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001715">
        <property type="entry name" value="CH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036872">
        <property type="entry name" value="CH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014797">
        <property type="entry name" value="CKK_CAMSAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038209">
        <property type="entry name" value="CKK_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011033">
        <property type="entry name" value="PRC_barrel-like_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21595:SF1">
        <property type="entry name" value="CALMODULIN-REGULATED SPECTRIN-ASSOCIATED PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21595">
        <property type="entry name" value="PATRONIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17095">
        <property type="entry name" value="CAMSAP_CC1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11971">
        <property type="entry name" value="CAMSAP_CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08683">
        <property type="entry name" value="CAMSAP_CKK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25532">
        <property type="entry name" value="CH_CAMSAP2_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01051">
        <property type="entry name" value="CAMSAP_CKK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47576">
        <property type="entry name" value="Calponin-homology domain, CH-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50346">
        <property type="entry name" value="PRC-barrel domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50021">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51508">
        <property type="entry name" value="CKK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="5" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="3 6" id="KW-0493">Microtubule</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Calponin-homology (CH)" evidence="9">
        <location>
            <begin position="211"/>
            <end position="324"/>
        </location>
    </feature>
    <feature type="domain" description="CKK" evidence="10">
        <location>
            <begin position="1353"/>
            <end position="1487"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="440"/>
            <end position="463"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="610"/>
            <end position="629"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="663"/>
            <end position="721"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="948"/>
            <end position="981"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1200"/>
            <end position="1229"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1244"/>
            <end position="1283"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1297"/>
            <end position="1347"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="752"/>
            <end position="779"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="881"/>
            <end position="918"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="1040"/>
            <end position="1076"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="440"/>
            <end position="456"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="668"/>
            <end position="690"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="694"/>
            <end position="704"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="961"/>
            <end position="972"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="8">
        <location>
            <begin position="1249"/>
            <end position="1263"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="1299"/>
            <end position="1310"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00841"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS51508"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029357.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1493" mass="169150" checksum="6F802A5C010CB91B" modified="2019-05-08" version="1">MGDAADLKEMRKTFIVPAVKPFDHYDFTRAKIACNVAWLVAKAFGTENVPEELREPFYTDQYDQEHIKPPVVNLLLSAELYCRAGSLILKSDAAKPLLGHDAVIQALAQKGLYVTDQEKLVTERDLHKKPIQMSAHLAMIDTLMMAYTVEMVSVEKVIACAQQYSVFFQATDLPYDIEDAVMYWINKVNEHLKDIMEQEQKLKEHHSVETPGGQKARYRKEQTLLKQLPCIPLVENLLKDGTDGCALAALIHFYCPDIVKLEDVCLKETMSLADSLYNLQLIQEFCQEYLNQCCHFTLEDMLYAAASVKSNYLVFMAELFWWFEVVKPSFVQPRVVINPQAEPVKDASSVPILNANRRNYMDSSCGSDFIASVEGPTFTPSHQLLPVRHTHHQPYSAAPGVIRRSTSMSYVDGYVGTWPKEKRSSMHGVSFDISFDNENSIQTSTPNRGITRSVSNEGIMPNINRMPKHIKKNLSFKPVNGEEETQDIEEEKEIVSHIDPKASASLNTNERNANENSQYKLPNGALQNRVVMDDFGNEIETPSIEEALQIIHDTEKSPSVIQPDQITNGFFLHNQEISILNSNIKLNQPNPDIITDTKVALSPVTDNTEVDTGIHVPSEDIPETMDEDSSLRDYTVSMDSDMEDPPKCLQDYDIRASNHRELLSPCPSSISTKSQAGSSASSSSGVKMTSFAEQKFRKLNHTDSRSSGSSSQKTTPEGSELNIPHVVAWTHIPEETPLPQGRDTTQLLASEMVHLRMKLEEKRRAIEAQKKKVEAAFTKQRQKMGRTAFLNVVKKKGDGISPLREEAAGAEDEKVFTDSNRLNEKDTQKPIEQINKTAEVIKEGAENSQAKWLKSPNTPMDAEKQWNLMSPAEETINEGDILEYTKSIEKLNASLHFLQQEMQRLSLQQEMLMQMREQQAWVISPPQPSPQKQIRELKSSMRQTGLSSPIAPFALEPPRPTHQSPQTSNRKNASFPVKMQRTPRPNELKIIPFNRTLTAPRSVDSLPRLRRFSPSQVPIQTRSFVCFGDDGERISEPQLKRNLLKEVKTTEDEAKEQQQKATLEECEQKVKEKEIKPFESKVSEVLSQPITETVCLTPNEDQLSQPVLPTLIPKTANLIEVSLSDLKSPEKAEVSVEKYEGESDREQFEDDQKVCCGFFFKDDQKAENDMAVKRAALLEKRLRRERETLLRKQQLEAELEHKKEETRRKTEEERQKKEDERARREFIKQEYMRRKQLKLMEEMDTVIKPRPHSSKQKKPRPKSIHRDHIESPKTPLKGPPGSQLYRVFSVSSLSLASLNTGDNDSVQSGKRTPRSESVEGFLSPSRCGSRNGEKDWENASTTSSVASATEYTGPKLYKEPSAKSNKHIIQNALAHCCLAGKVNEGQKKKILEEMEKSDANNFLILFRDSGCQFRSLYTYCPDTEEISKLTGIGPKSITKKMIEGLYKYNSDRKQFSHIPAKTMSASVDAITIHSHLWQSKRPVTPKKLLPTKA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMQ1</accession>
    <name>A0A452IMQ1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Endophilin-B2</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="8">SH3 domain-containing GRB2-like protein B2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="6">Homodimer, and heterodimer with SH3GLB1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the endophilin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMQ1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033097.1">
        <property type="protein sequence ID" value="ENSGAGP00000029140.1"/>
        <property type="gene ID" value="ENSGAGG00000021083.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061024">
        <property type="term" value="P:membrane organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07617">
        <property type="entry name" value="BAR_Endophilin_B2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd11944">
        <property type="entry name" value="SH3_Endophilin_B2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1270.60:FF:000017">
        <property type="entry name" value="endophilin-B2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.40:FF:000028">
        <property type="entry name" value="endophilin-B2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.60">
        <property type="entry name" value="Arfaptin homology (AH) domain/BAR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027267">
        <property type="entry name" value="AH/BAR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004148">
        <property type="entry name" value="BAR_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035640">
        <property type="entry name" value="Endophilin_B2_SH3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050384">
        <property type="entry name" value="Endophilin_SH3RF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14167:SF68">
        <property type="entry name" value="DREBRIN-LIKE PROTEIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14167">
        <property type="entry name" value="SH3 DOMAIN-CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03114">
        <property type="entry name" value="BAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14604">
        <property type="entry name" value="SH3_9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00721">
        <property type="entry name" value="BAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00326">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103657">
        <property type="entry name" value="BAR/IMD domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51021">
        <property type="entry name" value="BAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 10" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 9" id="KW-0728">SH3 domain</keyword>
    <feature type="domain" description="BAR" evidence="12">
        <location>
            <begin position="24"/>
            <end position="287"/>
        </location>
    </feature>
    <feature type="domain" description="SH3" evidence="11">
        <location>
            <begin position="343"/>
            <end position="403"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="10">
        <location>
            <begin position="207"/>
            <end position="234"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006697"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00038771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00040331"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00042180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS51021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029140.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="403" mass="44728" checksum="8254D715F451EA53" modified="2019-05-08" version="1">MDFNMKKLASDAGVFFSRAVQFTEEKLGQAEKTELDAHFESLLAKADCTKNWTEKILHQTEVLLQPNPSVRVEEFLYEKLDRKVPSRVTNGELLAQYMVEAANDFGPGTPYGKTLLKVGETQKRLGGAERDFMHSASINFLNPLRNFLEGDWRTITKERRILENRRLDLDACKARLKKAKAAEAKASAVPDFQETRPRNYILSASASALWSDEVEKAEHELRVAQTEFDRQAELTRLLLEGISSTHVNHLRCLHQFVESQATYYAQGYQYMLDLQKQLGSSKGEIFPGTFVGNAESTSPPPATTSPTTVAATIPVVATVGRAPNPATAVEGALNLNDVKPPASGTRKARVLYDYEAADSSELALLADEMITVYSLPGMDPDWLIGERGNQKGKVPVTYLELLS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMQ3</accession>
    <name>A0A452IMQ3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Kinesin-like protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
            <location evidence="2">Microtubule organizing center</location>
            <location evidence="2">Centrosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Spindle</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="13 14">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Kinesin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMQ3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033190.1">
        <property type="protein sequence ID" value="ENSGAGP00000029226.1"/>
        <property type="gene ID" value="ENSGAGG00000021132.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005874">
        <property type="term" value="C:microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005819">
        <property type="term" value="C:spindle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008017">
        <property type="term" value="F:microtubule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003777">
        <property type="term" value="F:microtubule motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007019">
        <property type="term" value="P:microtubule depolymerization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007018">
        <property type="term" value="P:microtubule-based movement"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd01367">
        <property type="entry name" value="KISc_KIF2_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000006">
        <property type="entry name" value="Kinesin-like protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR054473">
        <property type="entry name" value="KIF2A-like_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027640">
        <property type="entry name" value="Kinesin-like_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019821">
        <property type="entry name" value="Kinesin_motor_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001752">
        <property type="entry name" value="Kinesin_motor_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47971:SF24">
        <property type="entry name" value="KINESIN-LIKE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47971">
        <property type="entry name" value="KINESIN-RELATED PROTEIN 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22923">
        <property type="entry name" value="KIF2A-like_1st"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00225">
        <property type="entry name" value="Kinesin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00380">
        <property type="entry name" value="KINESINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00129">
        <property type="entry name" value="KISc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00411">
        <property type="entry name" value="KINESIN_MOTOR_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50067">
        <property type="entry name" value="KINESIN_MOTOR_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 13 14" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="12" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="4" id="KW-0132">Cell division</keyword>
    <keyword evidence="9 15" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="5 14" id="KW-0493">Microtubule</keyword>
    <keyword evidence="7" id="KW-0498">Mitosis</keyword>
    <keyword evidence="10 13 14" id="KW-0505">Motor protein</keyword>
    <keyword evidence="6 13 14" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Kinesin motor" evidence="17">
        <location>
            <begin position="177"/>
            <end position="507"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1"/>
            <end position="114"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="122"/>
            <end position="141"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="15">
        <location>
            <begin position="614"/>
            <end position="648"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="96"/>
            <end position="106"/>
        </location>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <begin position="267"/>
            <end position="274"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004186"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022776"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00283"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="RuleBase" id="RU000394"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50067"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029226.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="660" mass="75006" checksum="EE3DB2A61DC52C65" modified="2019-05-08" version="1">MVTSLNEDNESVTVEWIENGDTKGKEIDLESIFSLNPDLAPDEDIEPSPETPPPPTPSAKVNKIVKNRRTVAPIKNETPARDNRAVGSARARPTQLPDQSSSTQQNARRKSNCVKEVEKLQEKREKRRLQQQELREKRAQDVDATNPNYEIMCMIRDFRGSLDYRPLTTTDPIDEHRICVCVRKRPLNKKETVMKDLDVITIPSKDVVMVHEPKQKVDLTRYLENQTFRFDYAFDDTAPNEMVYRFTARPLVETIFERGMATCFAYGQTGSGKTHTMGGDFSGKNQDCSKGIYALAARDVFLMLKKPNYKKLELQVYATFFEIYSGKVFDLLNRKTKLRVLEDGKQQVQVVGLQEREVKCVEDVLKLIEIGNSCRTSGQTSANAHSSRSHAVFQIILRRKGKLHGKFSLIDLAGNERGADTSSADRQTRLEGAEINKSLLALKECIRALGRNKPHTPFRASKLTQVLRDSFIGENSRTCMIATISPGMASCENTLNTLRYANRVKELTVDPSAAGDIRPIIHHAPNQIDDLETQWGVGSSPQRDDLKLLCEQNEEEVSPQLFTFHEAVSQMVEMEEQVVEDHRAVFQESIRWLEDEKALLEMTEEVDYDVDSYATQLEAILEQKIDILTELRDKVKSFRAALQEEEQASKQINPKRPRAL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMQ4</accession>
    <name>A0A452IMQ4_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="8">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
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        <property type="gene ID" value="ENSGAGG00000020783.1"/>
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        <property type="component" value="Unassembled WGS sequence"/>
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        <property type="evidence" value="ECO:0007669"/>
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        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
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        <property type="project" value="TreeGrafter"/>
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        <property type="project" value="UniProtKB-KW"/>
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        <property type="term" value="P:mitotic actomyosin contractile ring assembly actin filament organization"/>
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        <property type="project" value="TreeGrafter"/>
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        <property type="entry name" value="Calponin-like domain"/>
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        <property type="entry name" value="GTPase Activation - p120gap, domain 1"/>
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    <dbReference type="PANTHER" id="PTHR14149:SF10">
        <property type="entry name" value="RAS GTPASE-ACTIVATING-LIKE PROTEIN IQGAP3"/>
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    </dbReference>
    <dbReference type="PROSITE" id="PS00509">
        <property type="entry name" value="RAS_GTPASE_ACTIV_1"/>
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    </dbReference>
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    <proteinExistence type="predicted"/>
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    <keyword evidence="4" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="1" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="Calponin-homology (CH)" evidence="7">
        <location>
            <begin position="34"/>
            <end position="149"/>
        </location>
    </feature>
    <feature type="domain" description="WW" evidence="6">
        <location>
            <begin position="673"/>
            <end position="706"/>
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    </feature>
    <feature type="domain" description="Ras-GAP" evidence="5">
        <location>
            <begin position="972"/>
            <end position="1205"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="4">
        <location>
            <begin position="158"/>
            <end position="185"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
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    </evidence>
    <evidence type="ECO:0000256" key="4">
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        </source>
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        </source>
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        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1596" mass="181188" checksum="96DED33AC59238FB" modified="2019-05-08" version="1">MIDLVSSLSNCPDERLTADEMDEQRRQNVAYQYLCRLEEAKRWMEACLNEELASPTELEESLRNGVILAKLGHCFAPDIIPLKKIYDREQMRYEATGLHFRHTDNINHWREAMAHLGLPTIFHPETTDIYDKKNMPRVVYCIHALSLYLFKLGLAPQIQDLYGKVNFTEEEINNMKRELEKYGLQMPAFSKIGGILASELSVDEAAVHAAVLAINEAVERGVVEQTMVALHNPSAMLLNLRENLAAVYQEMLFQVKMEKAGNARNRESPESEDIYDRCLTQAEIQGNINKVNVHGALEYVDDALERQDPLALCQALQDPVLALRGVQRENAAWYLDQLSTDREQKALLGYVDLLEQEEVQAGVYAANERGDQEHAMRRAVSRINASIRRGEAMETVRELMSPEAQLPKAFPFAAQLYQHELSLLQQHHPQLGQEELYVAVEMLSAVALIDRALEAGDVSGFWSSLVSPALGLSDIEDANAQRYFDDLLKLKRRSGKAGVAFLSWNDIQACVNSANLAVQDENDKILAIRLINEALAQDDPEKTLTALLLPSARFSAVTLPGARRYHDVLTAALKQKAQVTGDDGAALWWEEIQQGVSKANQDTEAARRLALGIAAINQAIKEGKPAQTARVLRNPDVSLRGIVAECAGAYQAQLAALLAAKRQAGKSLASCPGNANHNWVRHQLKDGSVYYFSLQNFEGSWERPHDCVLNSTHLSREEIQSTITGVTAAHDRERLWESNAGFVVRLQARMRGYLVRQEFVARRHLLQEQLPAVIKIQVRSAARLPSRSHADGHDDIYIALGLQGINAVIKIQAFVRANKARGDYRMLVHARDPPLSIVRRFVHLLEQSQHDFWEESELLKLREEVVKRIRSNQQLESDLNLMDIKIGLLVKNRITLQEVVSHCKKLTRKNKEQLSNLMAIDKQKGLKSLSKEKRQKLEAYQHLFYLLQTHPVYLAKLIFQMPQNKSTKFMESVVFTLYNYASNPREAYLLLQLFKTALQEEIRSKVDQLRDILTGNPTVIRLVVSFYRNARGQNALRQILGSPVQEVLQDKTLSIRTNPVDIYKAWINQLESQSGQKSKFPYDVSPEQALSHPEVQRRLDISIRNLLAVTDKFLSAITSSVDKIPYGMRYVAKVLKTSLAEKFPAAGEEDICKIVGNLLYYRFMNPAVVAPDGFDIVDISAGVALHPEHRRNLGSIAKILQHAAANKMFEGENGHLRVVNQYLEETHSKFRLFICAACSVPEPEERFNVDEYSEMVAVAKPVIYITVAELINTHKLLLEHQDSISPDHKDPLHELLEDLDELPTVQSLIGTCRLLVSFKPAPPLVSSFNPQNGTKQMLVDVIQSQPGDSLLEILRTAASEDQEAFHHHQMHRRALREAQTPDRLKRNRSLAGNSQLSMEEKKRKVIRNLRRLENLGVVDSANQYQEIINEMAKDIRNQRRYRQHRKGELVKLRQTLCGLNSKTSFYEEQIDYYNQYIRTCLDNLAASSKLNGKSKKQQSLHYTAARLFEKGVLLEIEDLPVSQFRNVIFDIIPCGEAGKFQVKAKFMGIDMEKFQLHYQDLLQLQYEGVAVMKLFDKAKVNVNLLIFLLNKKFFRK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMQ5</accession>
    <name>A0A452IMQ5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Endophilin-B2</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="8">SH3 domain-containing GRB2-like protein B2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="6">Homodimer, and heterodimer with SH3GLB1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the endophilin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMQ5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033105.1">
        <property type="protein sequence ID" value="ENSGAGP00000029145.1"/>
        <property type="gene ID" value="ENSGAGG00000021083.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061024">
        <property type="term" value="P:membrane organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07617">
        <property type="entry name" value="BAR_Endophilin_B2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd11944">
        <property type="entry name" value="SH3_Endophilin_B2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1270.60:FF:000017">
        <property type="entry name" value="endophilin-B2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.40:FF:000028">
        <property type="entry name" value="endophilin-B2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.60">
        <property type="entry name" value="Arfaptin homology (AH) domain/BAR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027267">
        <property type="entry name" value="AH/BAR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004148">
        <property type="entry name" value="BAR_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035640">
        <property type="entry name" value="Endophilin_B2_SH3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050384">
        <property type="entry name" value="Endophilin_SH3RF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14167:SF68">
        <property type="entry name" value="DREBRIN-LIKE PROTEIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14167">
        <property type="entry name" value="SH3 DOMAIN-CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03114">
        <property type="entry name" value="BAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14604">
        <property type="entry name" value="SH3_9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00721">
        <property type="entry name" value="BAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00326">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103657">
        <property type="entry name" value="BAR/IMD domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51021">
        <property type="entry name" value="BAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 10" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 9" id="KW-0728">SH3 domain</keyword>
    <feature type="domain" description="BAR" evidence="12">
        <location>
            <begin position="24"/>
            <end position="266"/>
        </location>
    </feature>
    <feature type="domain" description="SH3" evidence="11">
        <location>
            <begin position="322"/>
            <end position="382"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="10">
        <location>
            <begin position="179"/>
            <end position="213"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006697"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00038771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00040331"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00042180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS51021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029145.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="382" mass="42452" checksum="4C113742160E53B2" modified="2019-05-08" version="1">MDFNMKKLASDAGVFFSRAVQFTEEKLGQAEKTELDAHFESLLAKADCTKNWTEKILHQTEVLLQPNPSVRVEEFLYEKLDRKVPSRVTNGELLAQYMVEAANDFGPGTPYGKTLLKVGETQKRLGGAERDFMHSASINFLNPLRNFLEGDWRTITKERRILENRRLDLDACKARLKKAKAAEAKASLWSDEVEKAEHELRVAQTEFDRQAELTRLLLEGISSTHVNHLRCLHQFVESQATYYAQGYQYMLDLQKQLGSSKGEIFPGTFVGNAESTSPPPATTSPTTVAATIPVVATVGRAPNPATAVEGALNLNDVKPPASGTRKARVLYDYEAADSSELALLADEMITVYSLPGMDPDWLIGERGNQKGKVPVTYLELLS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMQ9</accession>
    <name>A0A452IMQ9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">DNA polymerase epsilon catalytic subunit</fullName>
            <ecNumber evidence="15">2.7.7.7</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">DNA polymerase II participates in chromosomal DNA replication.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>DNA(n) + a 2'-deoxyribonucleoside 5'-triphosphate = DNA(n+1) + diphosphate</text>
            <dbReference type="Rhea" id="RHEA:22508"/>
            <dbReference type="Rhea" id="RHEA-COMP:17339"/>
            <dbReference type="Rhea" id="RHEA-COMP:17340"/>
            <dbReference type="ChEBI" id="CHEBI:33019"/>
            <dbReference type="ChEBI" id="CHEBI:61560"/>
            <dbReference type="ChEBI" id="CHEBI:173112"/>
            <dbReference type="EC" id="2.7.7.7"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="15">
            <name>[4Fe-4S] cluster</name>
            <dbReference type="ChEBI" id="CHEBI:49883"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 15">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 15">Belongs to the DNA polymerase type-B family.</text>
    </comment>
    <dbReference type="EC" id="2.7.7.7" evidence="15"/>
    <dbReference type="AlphaFoldDB" id="A0A452IMQ9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033112.1">
        <property type="protein sequence ID" value="ENSGAGP00000029150.1"/>
        <property type="gene ID" value="ENSGAGG00000020939.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008622">
        <property type="term" value="C:epsilon DNA polymerase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051539">
        <property type="term" value="F:4 iron, 4 sulfur cluster binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003887">
        <property type="term" value="F:DNA-directed DNA polymerase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000166">
        <property type="term" value="F:nucleotide binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008310">
        <property type="term" value="F:single-stranded DNA 3'-5' DNA exonuclease activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006287">
        <property type="term" value="P:base-excision repair, gap-filling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045004">
        <property type="term" value="P:DNA replication proofreading"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006272">
        <property type="term" value="P:leading strand elongation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000278">
        <property type="term" value="P:mitotic cell cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006297">
        <property type="term" value="P:nucleotide-excision repair, DNA gap filling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05779">
        <property type="entry name" value="DNA_polB_epsilon_exo"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05535">
        <property type="entry name" value="POLBc_epsilon"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.132.60:FF:000003">
        <property type="entry name" value="DNA polymerase epsilon catalytic subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.690:FF:000005">
        <property type="entry name" value="DNA polymerase epsilon catalytic subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.420.10:FF:000010">
        <property type="entry name" value="DNA polymerase epsilon catalytic subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1600.10:FF:000006">
        <property type="entry name" value="DNA polymerase epsilon catalytic subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.132.60">
        <property type="entry name" value="DNA polymerase family B, C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.342.10">
        <property type="entry name" value="DNA Polymerase, chain B, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.690">
        <property type="entry name" value="Helix hairpin bin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1600.10">
        <property type="entry name" value="Palm domain of DNA polymerase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.420.10">
        <property type="entry name" value="Ribonuclease H-like superfamily/Ribonuclease H"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006172">
        <property type="entry name" value="DNA-dir_DNA_pol_B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006133">
        <property type="entry name" value="DNA-dir_DNA_pol_B_exonuc"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043502">
        <property type="entry name" value="DNA/RNA_pol_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042087">
        <property type="entry name" value="DNA_pol_B_thumb"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023211">
        <property type="entry name" value="DNA_pol_palm_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029703">
        <property type="entry name" value="POL2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055191">
        <property type="entry name" value="POL2_thumb"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012337">
        <property type="entry name" value="RNaseH-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036397">
        <property type="entry name" value="RNaseH_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10670">
        <property type="entry name" value="DNA POLYMERASE EPSILON CATALYTIC SUBUNIT A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10670:SF0">
        <property type="entry name" value="DNA POLYMERASE EPSILON CATALYTIC SUBUNIT A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03104">
        <property type="entry name" value="DNA_pol_B_exo1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22634">
        <property type="entry name" value="POL2_thumb"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00486">
        <property type="entry name" value="POLBc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56672">
        <property type="entry name" value="DNA/RNA polymerases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53098">
        <property type="entry name" value="Ribonuclease H-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3 15" id="KW-0004">4Fe-4S</keyword>
    <keyword evidence="6 15" id="KW-0235">DNA replication</keyword>
    <keyword evidence="13 15" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="10 15" id="KW-0239">DNA-directed DNA polymerase</keyword>
    <keyword evidence="11 15" id="KW-0408">Iron</keyword>
    <keyword evidence="12 15" id="KW-0411">Iron-sulfur</keyword>
    <keyword evidence="7 15" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="5 15" id="KW-0548">Nucleotidyltransferase</keyword>
    <keyword evidence="14 15" id="KW-0539">Nucleus</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 15" id="KW-0808">Transferase</keyword>
    <keyword evidence="9 15" id="KW-0862">Zinc</keyword>
    <keyword evidence="8 15" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="DNA-directed DNA polymerase family B exonuclease" evidence="16">
        <location>
            <begin position="208"/>
            <end position="414"/>
        </location>
    </feature>
    <feature type="domain" description="DNA polymerase epsilon ,catalytic subunit A thumb" evidence="17">
        <location>
            <begin position="963"/>
            <end position="1090"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005755"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022485"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022695"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022705"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022932"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023014"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="RuleBase" id="RU365029"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF03104"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF22634"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029150.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1104" mass="126474" checksum="7AECA733C952CBEB" modified="2019-05-08" version="1">VPWFGGGGGPRLSRGLVDDGSGLSALKRLERSQRTDRLDAQFGFERPKEPGEKTGWLINMHPTEILDDDKRLVSSVDYYFIQEDGSRFKVSPFHFSVQGCEREVSSFLSKRYQGKIAKLETVPKEDLDLPNHLVGLKRNYVKLSFNTVDDLVKVRREISPAVKKNRERDQASDTYTSMLSSALIGGSLTMDDEGPSKKTVNQMDNIVDMREYDVPYHIRLSIDLKIHVAHWYNVRYRGSIYPPEITRRDDLVERPDPVVLAFDIETTKLPLKFPDAETDQIMMISYMIDGQGYLITNREIVSEDIEDFEFTPKPEYEGPFCIFNEPDEAHLIQRWFEHVQETKPTIMVTYNGDFFDWPFVEARAAVHGMNMYQEIGFQKDSQGEYKASQCIHMDCLRWVKRDSYLPVGSHNLKAAAKAKLGYDPVELDPEEMCRMATEEPQTLATYSVSDAVATYYMYMKYVHPFIFALCTIIPMEPDEVLRKGSGTLCEALLMVQAYHANIIFPNKQEQEFNKLTEDGHVLDSETYVGGHVEALESGVFRSDIPCRFKMNPVAFDFLLQRVEKTLRHAIEEEEGLPLDQVTNFQEVCDEIKVKLNSLKDIPNRIECPLIYHLDVGAMYPNIILTNRLQPSAMVDEATCAACDFNKPGANCQRRMTWQWRGEFMPASRSEYHRIQQQLESEKLPPLYPDGPPRAFHELSREEQAKFEKKRLADYCRKAYKKLHVTKVEERITTICQRENSFYVDTVRAFRDRRYEFKGLHKVWKKKLSAATEMGDATEVKRCKNMEILYDSLQLAHKCILNSFYGYVMRKGARWYSMEMAGIVCFTGANIITQARELIEQIGRPLELDTDGIWCVLPNSFPENFVIKSTNAKKPKVTISYPGAMLNILVKEGFTNDQYQELVDPASLTYVTRSENSIFFEVDGPYLAMILPASKEEGKKLKKRYAVFNEDGSLAELKGFEVKRRGELQLVKIFQSSVFESFLKGTTLEEVYASVAKVADYWLDVLYSKAANMPDSELFELISENRSMSRKLEDYGEQKSTSISTAKRLAEFLGDQMVKDAGLSCRFIISKKPEGSPVTERAIPLAIFQAELTPIPAGLQHPDGK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMR7</accession>
    <name>A0A452IMR7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Gastricsin</fullName>
            <ecNumber evidence="12">3.4.23.3</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="11">Hydrolyzes a variety of proteins.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="10">
            <text>More restricted specificity than pepsin A, but shows preferential cleavage at Tyr-|-Xaa bonds. High activity on hemoglobin.</text>
            <dbReference type="EC" id="3.4.23.3"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 16">Belongs to the peptidase A1 family.</text>
    </comment>
    <dbReference type="EC" id="3.4.23.3" evidence="12"/>
    <dbReference type="AlphaFoldDB" id="A0A452IMR7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033215.1">
        <property type="protein sequence ID" value="ENSGAGP00000029242.1"/>
        <property type="gene ID" value="ENSGAGG00000021150.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004190">
        <property type="term" value="F:aspartic-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007586">
        <property type="term" value="P:digestion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.70.10:FF:000006">
        <property type="entry name" value="Cathepsin E"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.70.10:FF:000004">
        <property type="entry name" value="Pepsin A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.60">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.70.10">
        <property type="entry name" value="Acid Proteases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001461">
        <property type="entry name" value="Aspartic_peptidase_A1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001969">
        <property type="entry name" value="Aspartic_peptidase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012848">
        <property type="entry name" value="Aspartic_peptidase_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033121">
        <property type="entry name" value="PEPTIDASE_A1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021109">
        <property type="entry name" value="Peptidase_aspartic_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47966">
        <property type="entry name" value="BETA-SITE APP-CLEAVING ENZYME, ISOFORM A-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47966:SF72">
        <property type="entry name" value="GASTRICSIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07966">
        <property type="entry name" value="A1_Propeptide"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00026">
        <property type="entry name" value="Asp"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00792">
        <property type="entry name" value="PEPSIN"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50630">
        <property type="entry name" value="Acid proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00141">
        <property type="entry name" value="ASP_PROTEASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51767">
        <property type="entry name" value="PEPTIDASE_A1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="16" id="KW-0064">Aspartyl protease</keyword>
    <keyword evidence="6" id="KW-0222">Digestion</keyword>
    <keyword evidence="9 15" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="7 16" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="4 16" id="KW-0645">Protease</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0964">Secreted</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <keyword evidence="8" id="KW-0865">Zymogen</keyword>
    <feature type="domain" description="Peptidase A1" evidence="17">
        <location>
            <begin position="41"/>
            <end position="358"/>
        </location>
    </feature>
    <feature type="active site" evidence="14">
        <location>
            <position position="59"/>
        </location>
    </feature>
    <feature type="active site" evidence="14">
        <location>
            <position position="249"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="72"/>
            <end position="77"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="15">
        <location>
            <begin position="283"/>
            <end position="316"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007447"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022757"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023145"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023733"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023749"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023796"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023821"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PIRSR" id="PIRSR601461-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PIRSR" id="PIRSR601461-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="RuleBase" id="RU000454"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS51767"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029242.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="361" mass="39798" checksum="BE9320D00E0861DE" modified="2019-05-08" version="1">KTIREVMKEKGVLVEYLKHHKTDPASKYRFNEYNVAFEPMMYMDVILEPPPQNFLVLFDTGSSNLWVPSVYCQSQACSNHARFNPSQSSTFSSNGQTFSLQYSSGSFTGFFGYDIMTVSVISVTQQSFGLSEKEPGTNFIYAQFDDILGMAYSSLVVGGATTALQGMLQENLLSQPIFSFYLSSQSTSQYGGEVIFGGVDSSLYSRQIYWAPVTQELYWQIGIDDVYGEFAIGGQATGWCSQGCQAIVDKGTSLLTVPQQLMNSFLQDVGAQENKYGEYVVDCSSVQSLPTISFTINGVSFPLPPSAYILNNNGYCSVGVEPMYLPSENGQPLLILGDVFLRQYYSVCDMGNNRIGFVAVA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMR8</accession>
    <name>A0A452IMR8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">C-type lectin domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IMR8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033099.1">
        <property type="protein sequence ID" value="ENSGAGP00000029141.1"/>
        <property type="gene ID" value="ENSGAGG00000020766.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="CDD" id="cd00037">
        <property type="entry name" value="CLECT"/>
        <property type="match status" value="8"/>
    </dbReference>
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        <property type="entry name" value="Macrophage mannose receptor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000016">
        <property type="entry name" value="macrophage mannose receptor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="macrophage mannose receptor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000025">
        <property type="entry name" value="Mannose receptor C-type 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000030">
        <property type="entry name" value="Mannose receptor C-type 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000027">
        <property type="entry name" value="Mannose receptor, C type 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.100.10">
        <property type="entry name" value="Mannose-Binding Protein A, subunit A"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001304">
        <property type="entry name" value="C-type_lectin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016186">
        <property type="entry name" value="C-type_lectin-like/link_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050111">
        <property type="entry name" value="C-type_lectin/snaclec_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018378">
        <property type="entry name" value="C-type_lectin_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016187">
        <property type="entry name" value="CTDL_fold"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22803">
        <property type="entry name" value="MANNOSE, PHOSPHOLIPASE, LECTIN RECEPTOR RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00059">
        <property type="entry name" value="Lectin_C"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01504">
        <property type="entry name" value="PNCREATITSAP"/>
    </dbReference>
    <dbReference type="SMART" id="SM00034">
        <property type="entry name" value="CLECT"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56436">
        <property type="entry name" value="C-type lectin-like"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00615">
        <property type="entry name" value="C_TYPE_LECTIN_1"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50041">
        <property type="entry name" value="C_TYPE_LECTIN_2"/>
        <property type="match status" value="8"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="2" id="KW-0472">Membrane</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="2" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="1146"/>
            <end position="1169"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="3">
        <location>
            <begin position="33"/>
            <end position="129"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="3">
        <location>
            <begin position="155"/>
            <end position="272"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="3">
        <location>
            <begin position="297"/>
            <end position="412"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="3">
        <location>
            <begin position="441"/>
            <end position="559"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="3">
        <location>
            <begin position="588"/>
            <end position="704"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="3">
        <location>
            <begin position="732"/>
            <end position="856"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="3">
        <location>
            <begin position="878"/>
            <end position="988"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="3">
        <location>
            <begin position="1017"/>
            <end position="1134"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS50041"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029141.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1215" mass="138331" checksum="4DE5153CC8B15942" modified="2019-05-08" version="1">WGRVRLFIYLCPNLKQLHSVLVASGINSNSALTWHQARKSCQQQNAELLSITELHEQTYLTANSELWIGLNRLNSNGGWQWIGGSPFRYLNWAPGSPSLESEKICGLLQSGNGKWKNQQCDQKLGYICKKGNSSLDSFIIPSGRPIKCPSGWIPYSGHCYMLYRAPKIWKDALSSCRKEGGDLASIHNIEEYSFTVSQLGYKPTDELWIGLNDLKIHMYFDWSDGTPVTYTKWLHEEPSHANNRKEDCVSMKGEDGYWADDVCEKKLGYICKMKPLAEESEKVEVIIPGCQKGWIRHGFYCYSVGQTSVTFSEAKKICEGNKGYLVTVKDRYEQAFLTVLIGFSPVKYFWIALSNVEEQGTFKWADGEAVLFTHWNSGMPGREPGCVAMRTGTTAGLWDVVNCEEKSMFLCKQLVEGATPPPVPTTTPAPSCPDGWHSRAHSSSCFKNGRKHMKTWFDARDFCRAIGGDLASIHSEEDQNVINSLERAHYYQSYWMGMSAFDPDGGFTWSDGSPVNYENWADGEPNNYDGNEKCAMFNEYTDMQWNDMFCEKFGDWTCQIKKGATLKPEPSTTFEYTYKVSEDGWIIYEDKAYYFSHETLPMEKAREYCKKNSGDLVVIEGESKRKFLWKYSFYNDFRESLYIGLSVSLDKKFSWIDGTPLNYVAWSPNEPNFANNDENCVVMYTQTGMWNDLNCGAPNSFICERLNNITHSTVAPTSPAPLGGCPQNWFLFNNKCFKIFGSSENESLTWHAARTACINLEGNLASIPSEEVQAFLITHLKDASTDTWIGLNDVNSEYTFLWTDGSGVSYTSWAKGFPNYRSEVNVDCKRSLNYFDKEAGKWKDEDCQMNKSYICQRNADPKLYNFQTTVPVSGFTLYGNSSYSLISSRMNWEEAQKNCKAEYSELASILDPYSQSFLWLQVLKIGQPVWIGLNSNTSGFYKWTDNWRIKYSKWASEEPKQNLACVYLDVDGTWKTGSCDENYCSVCKRSDVLAPTELPQLPGKCPTSDIRRSWIPFHGHCYYIESSEMESWPQSSMECVRLGASLVSIENMAEMNFLLHHLDAFASDSRQFWIGMYKNVDGEWTWADNSAVEFVNWKEREPTDGYHEYCVDMDASDGGWRSYYCSADKRFICKIPKNEAVPAHHLSGMAVILVLLILVGAGLAAYIFFKKKRHNQLPIDVTFDNTLYCNRDAVPAASDSKYLVANIEQNEQAML</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMR9</accession>
    <name>A0A452IMR9_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="5">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the STRIP family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMR9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033122.1">
        <property type="protein sequence ID" value="ENSGAGP00000029160.1"/>
        <property type="gene ID" value="ENSGAGG00000020767.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007010">
        <property type="term" value="P:cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040185">
        <property type="entry name" value="Far11/STRP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021819">
        <property type="entry name" value="Far11/STRP_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012486">
        <property type="entry name" value="Far11/STRP_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13239">
        <property type="entry name" value="PROTEIN REQUIRED FOR HYPHAL ANASTOMOSIS HAM-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13239:SF6">
        <property type="entry name" value="STRIATIN-INTERACTING PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11882">
        <property type="entry name" value="DUF3402"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07923">
        <property type="entry name" value="N1221"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01293">
        <property type="entry name" value="DUF3402"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01292">
        <property type="entry name" value="N1221"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Far11/STRP N-terminal" evidence="3">
        <location>
            <begin position="1"/>
            <end position="240"/>
        </location>
    </feature>
    <feature type="domain" description="Far11/STRP C-terminal" evidence="4">
        <location>
            <begin position="337"/>
            <end position="693"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="245"/>
            <end position="264"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="2">
        <location>
            <begin position="253"/>
            <end position="264"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007062"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="SMART" id="SM01292"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="SMART" id="SM01293"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029160.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="715" mass="82549" checksum="75DE7AA56926F1A2" modified="2019-05-08" version="1">MRLLDGLEVVNRDKRLKAARAVLYLAQGVFGECDVEAEVLHWSRHNNFLLFQMGTFSAFLELLNMEIDNSQACSSALRKPAISLADSTELRVLLSVMYLMVENIRVEAEADPPEWKSSRETFRTELSFPVHSEEPFALLLFTMVAKFCSGHAPHFPMKKVLLLLWKVLLFTLGGFEALQAMKTKKREELGLPPLPEDSIQVVRRMRAASPPTYTIELLDQQSSKRGHRSRRPLVKQDSLDIYNERDPFKNDEAGVDEEENDDVDSGIEGELDLMERDAIIQAGPAQRPPIERVSFPKGLPWAPKVRQKDIEHFLEASRNKFIGFTLGQDTDTLVGLPRPIHESVKTLKQHKYVSISDVQIKNEEELEKCPMSLGEEEVQETPCEMLYRAMLYNLPQYMIALLKILLAAAPTSKAKTDSINILADVLPEEMPITVLQSMKLGVDVNRHKEIIVKSISASLLLLLKHFKLNHIYQFEYVSQHLVFANCIPLILKFFNQNIMSYITAKNSISVLDYPHCTVHELPELTAESLEAGDNNQFCWRNLFSCINLLRLLNKLTKWKHSRTMMLVVFKSAPILKRALKVKQAMMQLYVLKLLKIQTKYLGRQWRKSNMKTMSAIYQKVRHRMNDDWAYGNDIDARPWDFQAEECTLRANIEAFNSRRYDKSQDSEFAPVDNCLQSVLGQPLELPEDFHYSYELWLEREVFSQPIRWEELLHYQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMS0</accession>
    <name>A0A452IMS0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">SMB domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMS0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033358.1">
        <property type="protein sequence ID" value="ENSGAGP00000029377.1"/>
        <property type="gene ID" value="ENSGAGG00000020186.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009986">
        <property type="term" value="C:cell surface"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004551">
        <property type="term" value="F:dinucleotide phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003676">
        <property type="term" value="F:nucleic acid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004528">
        <property type="term" value="F:phosphodiesterase I activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046034">
        <property type="term" value="P:ATP metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030505">
        <property type="term" value="P:inorganic diphosphate transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045599">
        <property type="term" value="P:negative regulation of fat cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009143">
        <property type="term" value="P:nucleoside triphosphate catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030500">
        <property type="term" value="P:regulation of bone mineralization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd16018">
        <property type="entry name" value="Enpp"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00091">
        <property type="entry name" value="NUC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.410.20">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.720.10">
        <property type="entry name" value="Alkaline Phosphatase, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.570.10">
        <property type="entry name" value="Extracellular Endonuclease, subunit A"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017850">
        <property type="entry name" value="Alkaline_phosphatase_core_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044929">
        <property type="entry name" value="DNA/RNA_non-sp_Endonuclease_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001604">
        <property type="entry name" value="Endo_G_ENPP1-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020821">
        <property type="entry name" value="ENPP1-3/EXOG-like_nuc-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044925">
        <property type="entry name" value="His-Me_finger_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002591">
        <property type="entry name" value="Phosphodiest/P_Trfase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036024">
        <property type="entry name" value="Somatomedin_B-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001212">
        <property type="entry name" value="Somatomedin_B_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10151">
        <property type="entry name" value="ECTONUCLEOTIDE PYROPHOSPHATASE/PHOSPHODIESTERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10151:SF77">
        <property type="entry name" value="ECTONUCLEOTIDE PYROPHOSPHATASE_PHOSPHODIESTERASE FAMILY MEMBER 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01663">
        <property type="entry name" value="Phosphodiest"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01033">
        <property type="entry name" value="Somatomedin_B"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00892">
        <property type="entry name" value="Endonuclease_NS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00477">
        <property type="entry name" value="NUC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00201">
        <property type="entry name" value="SO"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53649">
        <property type="entry name" value="Alkaline phosphatase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54060">
        <property type="entry name" value="His-Me finger endonucleases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90188">
        <property type="entry name" value="Somatomedin B domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00524">
        <property type="entry name" value="SMB_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50958">
        <property type="entry name" value="SMB_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="4" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <feature type="domain" description="SMB" evidence="7">
        <location>
            <begin position="10"/>
            <end position="50"/>
        </location>
    </feature>
    <feature type="domain" description="SMB" evidence="7">
        <location>
            <begin position="53"/>
            <end position="97"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50958"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029377.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="789" mass="90942" checksum="3DA74DC7F72256BD" modified="2025-10-08" version="2">MKVMTLSNISVKTCKGRCFERTFGSCRCDRDCVELGNCCLDFQETCIQPGKTHIWTCNKFRCGEKRWPENRCSCSDDCVENKDCCVNYYIVCQGAKSWVEEKCEDINEPQCPPGFAKPPVLLFSLDGFRAEYLHTWSGLLPVISKLQKCGTYTPNMRPVYPTKTFPNHYTIVTGLYPESHGIIDNKIYDPHRNVSFTLKNAEKFNPQWYQGQPIWLTAMHQGLKAGTFFWPGSDVRINGTFPDLYRKYNGSIPFEERVIAILHWLQLPKDERPHFYTLYLEEPDSSGHNFGPVSSGVILALQRVDNIVGMLMDGLKQMNLHKCLNIILLSDHGKLMCTHCSKTAYLSTYMENVQDITVIPGPAARLRPQNVPDEYFSYPDQHFKAFMKELLPKRFHYVNNDRIERVHFYLDPQWQLARKPSEIKYCNGGFHGSDNRFMSMQAIFIGFGPGFKFKTQVDPFENIEIYNLICDLLGITPAYNNGTHGSLNHLLKNPVYIPRHPEEESHPSVYPLMKTTSSLDIGCSCDLAYIELVINSVSALVKKIEKYSLPYGRPRVLQKRSTYYLLYHHQYVSGYSQDLNMTLWSAYTVTKYDNRTASVENLSNCLHVDIRIPSSSSKTCSFYKNHHQLSYGFLFPPMLWNYFYDVLLPKYTLARNGVNVVSGPVFDYDSNGLYDTPEKVKRLTHDPEVLVPTHYFIVLTSCKNISQTPLQCEGALDALSFIVPHRPDNSESCARHKRDSLWVEERMRFHTARVRDVELLTGLSFYQDRKQPVSDILQLKTYLPTFPME</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMS3</accession>
    <name>A0A452IMS3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Endophilin-B2</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="8">SH3 domain-containing GRB2-like protein B2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="6">Homodimer, and heterodimer with SH3GLB1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the endophilin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMS3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033108.1">
        <property type="protein sequence ID" value="ENSGAGP00000029146.1"/>
        <property type="gene ID" value="ENSGAGG00000021083.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061024">
        <property type="term" value="P:membrane organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07617">
        <property type="entry name" value="BAR_Endophilin_B2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd11944">
        <property type="entry name" value="SH3_Endophilin_B2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1270.60:FF:000017">
        <property type="entry name" value="endophilin-B2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.40:FF:000028">
        <property type="entry name" value="endophilin-B2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.60">
        <property type="entry name" value="Arfaptin homology (AH) domain/BAR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027267">
        <property type="entry name" value="AH/BAR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004148">
        <property type="entry name" value="BAR_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035640">
        <property type="entry name" value="Endophilin_B2_SH3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050384">
        <property type="entry name" value="Endophilin_SH3RF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14167:SF68">
        <property type="entry name" value="DREBRIN-LIKE PROTEIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14167">
        <property type="entry name" value="SH3 DOMAIN-CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03114">
        <property type="entry name" value="BAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14604">
        <property type="entry name" value="SH3_9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00721">
        <property type="entry name" value="BAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00326">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103657">
        <property type="entry name" value="BAR/IMD domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51021">
        <property type="entry name" value="BAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 10" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 9" id="KW-0728">SH3 domain</keyword>
    <feature type="domain" description="BAR" evidence="12">
        <location>
            <begin position="24"/>
            <end position="258"/>
        </location>
    </feature>
    <feature type="domain" description="SH3" evidence="11">
        <location>
            <begin position="314"/>
            <end position="374"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="10">
        <location>
            <begin position="169"/>
            <end position="198"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006697"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00038771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00040331"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00042180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS51021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029146.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="374" mass="41465" checksum="5811C01D3A2BC6B7" modified="2019-05-08" version="1">MDFNMKKLASDAGVFFSRAVQFTEEKLGQAEKTELDAHFESLLAKADCTKNWTEKILHQTEVLLQPNPSVRVEEFLYEKLDRKVPSRVTNGELLAQYMVEAANDFGPGTPYGKTLLKVGETQKRLGGAERDFMHSASINFLNPLRNFLEGDWRTITKERRILENRRLDLDACKARLKKAKAAEAKASAEHELRVAQTEFDRQAELTRLLLEGISSTHVNHLRCLHQFVESQATYYAQGYQYMLDLQKQLGSSKGEIFPGTFVGNAESTSPPPATTSPTTVAATIPVVATVGRAPNPATAVEGALNLNDVKPPASGTRKARVLYDYEAADSSELALLADEMITVYSLPGMDPDWLIGERGNQKGKVPVTYLELLS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMS4</accession>
    <name>A0A452IMS4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">Ras-GAP domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IMS4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033127.1">
        <property type="protein sequence ID" value="ENSGAGP00000029165.1"/>
        <property type="gene ID" value="ENSGAGG00000019812.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005938">
        <property type="term" value="C:cell cortex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903479">
        <property type="term" value="P:mitotic actomyosin contractile ring assembly actin filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05131">
        <property type="entry name" value="RasGAP_IQGAP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00201">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.506.10:FF:000004">
        <property type="entry name" value="IQ motif containing GTPase activating protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.506.10">
        <property type="entry name" value="GTPase Activation - p120gap, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000048">
        <property type="entry name" value="IQ_motif_EF-hand-BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000593">
        <property type="entry name" value="RasGAP_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023152">
        <property type="entry name" value="RasGAP_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001936">
        <property type="entry name" value="RasGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001202">
        <property type="entry name" value="WW_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149">
        <property type="entry name" value="RAS GTPASE-ACTIVATING PROTEIN WITH IQ MOTIF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149:SF12">
        <property type="entry name" value="RAS GTPASE-ACTIVATING-LIKE PROTEIN IQGAP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00612">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00616">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03836">
        <property type="entry name" value="RasGAP_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00015">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00323">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00456">
        <property type="entry name" value="WW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF143885">
        <property type="entry name" value="RGC domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00509">
        <property type="entry name" value="RAS_GTPASE_ACTIV_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50018">
        <property type="entry name" value="RAS_GTPASE_ACTIV_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01159">
        <property type="entry name" value="WW_DOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Ras-GAP" evidence="1">
        <location>
            <begin position="613"/>
            <end position="841"/>
        </location>
    </feature>
    <evidence type="ECO:0000259" key="1">
        <source>
            <dbReference type="PROSITE" id="PS50018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029165.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1260" mass="143902" checksum="0D02EF431B96B4BF" modified="2019-05-08" version="1">ADGKNPINFLYFPPELVAVDHINTAIRNCDANKTLVALMKPEAQLPVVHPFAAAVYQTELFNLQQQNAVRYLAHDELSIAVEMLSAVVLLNQALGSKDILAIKNHLRNPSVGFNNVEDENFQRYADTLLSIKSEASSQGQDYLSWNDVQNCIDMVNMQVQEENDRIVAVSYINEAIDQGNPGKTLETLLLPTVKLHDVNPTNARHYQDVLHDAKVQKCKESHDESAVLWLDEIQKGINDANSNIEKAANLALGTSMINNCLEKHDSQPVLDILQSPKFGLRVVPECAETYYKNLLEAKNLKTKEDSSEGPWLKLIMKNRYDYYYNVETGESTFVPPEGFIPKTSWLTSEEIQTIVGQVTADYNREQLWFANENLILQLQALARGFLVRKSYEERKGFLQKQEPSVIKIQVLLCNIKAQSLLLEHDLLFEVSCSMLSKVFSLWSFFQNEEIVKIQAFLRANKARDDYRTLIGAENPPLNVLRKFAYLLDQSDLDFQEELEVTRLREEVVTKIRSNQQLEKDLNLMDIKIGLLVKNRISLQDVVLHSKKLNKKSKNQVEEMVTGDKQGIKGLSKERRKQLEAYQHLFYLLQTNPMYLAKLIFQMPQNKSTKFMDTVIFTLYNYASNQREEYLLLKLFKTALEEEINSKVDQIQDIVTGNPTVIKMVVSFNRGARGQNSLRQLLAPVVKEIMDDKSLLINTSPVEVYKAWVNQLETQTGEASKLPYDVTTEQALEHPEVVNRLESSIQSLRTVTDKVLVSIFSSYGMRYIAKVLKNSLHEKFPDATEDELLKIVGNLLYYRYMNPAIVAPDGFDIIDMTAGGQIHSDQRRNLGCVAKVLQHAASNKLFEGESAHLSSMNTYLSQTYQKFRNFFQIACDVPEPEEKFNIDEYSDMVTLSKPVIYISIEEIINTHSLLLEHQDAIAPEQNDLLNELLKGLGAVPDVESFLGEGAVDPNDPNKENTLSQLAKTEISLSLTSKYELQEGEDQDLKGTKRLVVDVIRTQPGDTLLEILEIPASAQQESEHLKVVEKRAILDSKTPEKMKHSQSMLEDGQLPIEQKKRKIQRNLRTLEQAGLVSSANKYQEIINEIAKDIRNQRRYRHHRKAELVKLQQTLSALQSKTAFYEDQVNYYNTYIKTCLDNLTRKSIKLDGKVEEKGSKKLKHTSLKYTAARLHEKGVILEIEDLQTNQFKNVMFDITPGEEMGDFEIKAKFLGVQMEKVELHFQDLLQMQYEGVAVMKMFDKAKVNVNLLIFLLNKKFYGK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMS6</accession>
    <name>A0A452IMS6_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="8">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the major facilitator superfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMS6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033146.1">
        <property type="protein sequence ID" value="ENSGAGP00000029184.1"/>
        <property type="gene ID" value="ENSGAGG00000021090.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="CDD" id="cd17454">
        <property type="entry name" value="MFS_NaGLT1_MFSD4B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1250.20:FF:000508">
        <property type="entry name" value="Sodium-dependent glucose transporter 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1250.20">
        <property type="entry name" value="MFS general substrate transporter like domains"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036259">
        <property type="entry name" value="MFS_trans_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23121">
        <property type="entry name" value="SODIUM-DEPENDENT GLUCOSE TRANSPORTER 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23121:SF9">
        <property type="entry name" value="SODIUM-DEPENDENT GLUCOSE TRANSPORTER 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103473">
        <property type="entry name" value="MFS general substrate transporter"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 7" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 7" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="60"/>
            <end position="81"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="101"/>
            <end position="123"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="130"/>
            <end position="146"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="152"/>
            <end position="179"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="191"/>
            <end position="211"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="252"/>
            <end position="274"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="295"/>
            <end position="322"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="367"/>
            <end position="385"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="391"/>
            <end position="413"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="425"/>
            <end position="445"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="457"/>
            <end position="477"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="1"/>
            <end position="53"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="6">
        <location>
            <begin position="1"/>
            <end position="11"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="13"/>
            <end position="24"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008335"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029184.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="523" mass="55852" checksum="5DA82B7C58219688" modified="2019-05-08" version="1">MAGAGRKKQVRFARPEAEDTRRGGEGPAAEQSLLPEPAPRPEVIVAGGGRSRAGEGATPLLRWCISGVLCSAFLGLGMSIAVLGPTFQDLAANVNKNVSDIYYIFMGRSLGYLGGSVIGGIIFDCMNPHLLLGLTMLGTAVGLYAIPWCKRALLLTALMSVIGTSVGVLDTGGNVLALNIWGAEAGPHMQALHFSFALGAVVAPILTKMALSSFISPNFQTEDEKTNHSVPSDIPNALFGSALKLHSNVSFMWSYIVIGTYILIVSLLFVVLYSRTGLARDKAKSSLQKCQIAKYHYALLILLFLFFFCYVGAEVTYGSYIFNYAEVYADMKESEAASLNSLFWGAFAACRGLAICFATCLYPGTMILLSVIGSTVSSLFLALLSKHPISLWIGTAVYGASMATVFPSGISWIEQYTTIQGKSASLFVVGAALGEMCIPAVVGFLEGKLHNVPVMMYAALGTSVMTAVLFPVMYKLATSPSESKLKDAKSPVAPYRLTDVLEHELSWVNTHLSDACSKTSVSL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMS7</accession>
    <name>A0A452IMS7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Protein FAM118B</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">May play a role in Cajal bodies formation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="7">Nucleus</location>
            <location evidence="7">Nuclear body</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the FAM118 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMS7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033225.1">
        <property type="protein sequence ID" value="ENSGAGP00000029252.1"/>
        <property type="gene ID" value="ENSGAGG00000021159.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016604">
        <property type="term" value="C:nuclear body"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038916">
        <property type="entry name" value="FAM118"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR28623">
        <property type="entry name" value="PROTEIN FAM118B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR28623:SF1">
        <property type="entry name" value="PROTEIN FAM118B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0007">Acetylation</keyword>
    <keyword evidence="6" id="KW-0539">Nucleus</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="279"/>
            <end position="312"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00003199"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006491"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00014026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00034306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029252.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="312" mass="35092" checksum="3765D47CE7389750" modified="2019-05-08" version="1">MASTVSVNLDKDDTFDGIPPVKKPRKLLPSLKTKKPQELVLVIGTGISAAVAPQVPALKSWKGLIQALLDAAIDFDLLEEEESRKFQKGLQEDKNLVHVAHDLIQKLSPRTSNVRSTFFKDCLYEVFDDLESKMEDSGKQLLQSVLRLMENGALVLTTNFDNLLELYAADQGKQLESLDLTDEKKGEIQKLYENKSFLFLGCGRTVDDTTFQALFLEAVKRKSDLEHFMLVQRGDVDEFKKLRENMLDKGIKVISYGNEYADLPDYFERLTSEISMRDQTGLPREGQLNGSSSAHTEIKEGAAPEHLPLQLN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMS9</accession>
    <name>A0A452IMS9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Neurabin-1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="14">Neurabin-I</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="13">Neural tissue-specific F-actin-binding protein I</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="15">Protein phosphatase 1 regulatory subunit 9A</fullName>
        </alternativeName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="11">Synapse</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000033372.1">
        <property type="protein sequence ID" value="ENSGAGP00000029388.1"/>
        <property type="gene ID" value="ENSGAGG00000020904.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015629">
        <property type="term" value="C:actin cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030425">
        <property type="term" value="C:dendrite"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0014069">
        <property type="term" value="C:postsynaptic density"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007015">
        <property type="term" value="P:actin filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019722">
        <property type="term" value="P:calcium-mediated signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031175">
        <property type="term" value="P:neuron projection development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd06790">
        <property type="entry name" value="PDZ_neurabin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd09512">
        <property type="entry name" value="SAM_Neurabin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000010">
        <property type="entry name" value="Neurabin-1 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.50:FF:000008">
        <property type="entry name" value="Neurabin-1 isoform 1-like protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.42.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.50">
        <property type="entry name" value="Transcription Factor, Ets-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040645">
        <property type="entry name" value="Neurabin-1/2_PDZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043446">
        <property type="entry name" value="Neurabin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001478">
        <property type="entry name" value="PDZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036034">
        <property type="entry name" value="PDZ_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001660">
        <property type="entry name" value="SAM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013761">
        <property type="entry name" value="SAM/pointed_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16154">
        <property type="entry name" value="NEURABIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16154:SF22">
        <property type="entry name" value="NEURABIN-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00595">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17817">
        <property type="entry name" value="PDZ_5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07647">
        <property type="entry name" value="SAM_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00228">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00454">
        <property type="entry name" value="SAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50156">
        <property type="entry name" value="PDZ domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47769">
        <property type="entry name" value="SAM/Pointed domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50106">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50105">
        <property type="entry name" value="SAM_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="9" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="8 16" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="10" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="5" id="KW-0221">Differentiation</keyword>
    <keyword evidence="6" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0770">Synapse</keyword>
    <feature type="domain" description="PDZ" evidence="19">
        <location>
            <begin position="504"/>
            <end position="592"/>
        </location>
    </feature>
    <feature type="domain" description="SAM" evidence="18">
        <location>
            <begin position="1173"/>
            <end position="1236"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1"/>
            <end position="104"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="176"/>
            <end position="196"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="275"/>
            <end position="302"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="897"/>
            <end position="938"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="980"/>
            <end position="1022"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1031"/>
            <end position="1050"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1239"/>
            <end position="1283"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="16">
        <location>
            <begin position="600"/>
            <end position="627"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="16">
        <location>
            <begin position="690"/>
            <end position="833"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="17">
        <location>
            <begin position="1"/>
            <end position="12"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="17">
        <location>
            <begin position="33"/>
            <end position="49"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="17">
        <location>
            <begin position="182"/>
            <end position="195"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="17">
        <location>
            <begin position="1000"/>
            <end position="1019"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="1037"/>
            <end position="1048"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="17">
        <location>
            <begin position="1239"/>
            <end position="1273"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00034103"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00067399"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00076637"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00077125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00082439"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50105"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50106"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029388.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1283" mass="143433" checksum="0517952F8DEEBFAC" modified="2019-05-08" version="1">MMKTESSGERSTLRSASPHRNAYRTEFQALKSTFDKPKSDGDQKTKDEGEVSQQSRGRKYGSNVNRIKNLFMQMGMEPSESTGVTPKTRGKSGPLSPQRIRPKEFVEKADGSVVKLESSVSERISRFDTVHGGPSYSKFTETRKMFERNVHEMGHSNRYSPKKERVVSNELQDEWCSSKSNRGSTDSLDSLSSRTETISPTVSQLSAVFENTDSHTVIVLEKSENNEEYSVTGHYPLNLSSTVENLSSTVANLDGFSPLKDASAWSSPAKQTTSSVSAETFQQNSTSSTIREASNSTLPVSKTNEEIRKNKEASFDPVDVCAATKQELASINMPXHSQNAESACVGTGAADIPRNLVAGGDFAADVASDATESPYDDASGKEVQKDLNNFQSSHVYMHSDYNVYRVRSRYNSDWGETGTEQDDQDDSDENNYYEPDMEYAEVIGLPAEDEIPANRRIKFSSAPIKVFNTYSNEDYDRRNDEVDPVAASAEYELEKRVEKLELFPVELGKDEDGLGISIIGMGVGADAGLEKLGIFVKTVTEGGAAERDGRIQVNDQIVEVDGISLVGVTQNFAATVLRNTKGNVRFVIGREKPGQVSEVAQLISQTLEQERRQRELLEQHYAQYDADDDEVIYLLFILSNTGEYATDEEDEDMGPIVPGADMAIEVFDLPENDDMFSPTELDTTKLAHKFKELQIKHAVTEAEIQKLKTKLQAAENEKVRWEREKTQLQQNIEENKERMMKLESYWIEAQTLCHTVNEHLKETQSQYQALEKKYNKAKKLIKDFQQKELDFIKRQEAERKKIEELEKAHLAEVQGLQARIRDLEAEVFKLLKQNGSQVNNNNNIFERQTSFGEVSRGDPVESLDVEQVSCLDDLSQDFNEAVPETERLDSKALKTRAQLSVRNKRQRPTRTRLYDSISSTDGEDSLERKPSDSYSSPMHISKSLLPICVRASSPASDSGASSLSPVASSAAFSFDNVAENQEGGQQHKGGVLSHYARGDTPLSSPSKSSHSSETSPLHHPTNRNVLQDKDILCGHTGPQSSRATSPTMGQAEKIKRKLVDLGAPLRRSSHKGKKCREREVIRVACAWKEKASLLQDVCIRSYVDFDPSFCLQNFTFTDDFSPSSTSSADLSGLGAEPKPPGFSHSLALSSDEILDDGQSPKHSQCQNRAVQEWSIQQVSHWLMSLNFEQYVSEFTAQNINGEHLLQLDGSKLKTLGMTSSQDRAIIKKKLKEMKVSIERARKAQEKMEKQREKLRKKEQERLQRKSKKLDKSSSDATEGANEQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMV4</accession>
    <name>A0A452IMV4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">non-specific serine/threonine protein kinase</fullName>
            <ecNumber evidence="3">2.7.11.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="12">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033399.1">
        <property type="protein sequence ID" value="ENSGAGP00000029413.1"/>
        <property type="gene ID" value="ENSGAGG00000021255.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.20.90:FF:000007">
        <property type="entry name" value="Serine/threonine-protein kinase WNK1 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000006">
        <property type="entry name" value="Serine/threonine-protein kinase WNK1 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.20.90:FF:000012">
        <property type="entry name" value="Serine/threonine-protein kinase WNK1 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000494">
        <property type="entry name" value="serine/threonine-protein kinase WNK2 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.20.90">
        <property type="entry name" value="Phosphatidylinositol 3-kinase Catalytic Subunit, Chain A, domain 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056865">
        <property type="entry name" value="CCTL2_WNK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024678">
        <property type="entry name" value="Kinase_OSR1/WNK_CCT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050588">
        <property type="entry name" value="WNK_Ser-Thr_kinase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13902">
        <property type="entry name" value="SERINE/THREONINE-PROTEIN KINASE WNK WITH NO LYSINE -RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24889">
        <property type="entry name" value="CCTL2_WNK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12202">
        <property type="entry name" value="OSR1_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="10" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="13" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9" id="KW-0418">Kinase</keyword>
    <keyword evidence="8" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="7" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Protein kinase" evidence="15">
        <location>
            <begin position="192"/>
            <end position="450"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1"/>
            <end position="184"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="606"/>
            <end position="625"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1094"/>
            <end position="1135"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1209"/>
            <end position="1248"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1273"/>
            <end position="1302"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1384"/>
            <end position="1404"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="1862"/>
            <end position="1926"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="2097"/>
            <end position="2116"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="2156"/>
            <end position="2185"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="2197"/>
            <end position="2229"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="13">
        <location>
            <begin position="1953"/>
            <end position="1980"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="1"/>
            <end position="12"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="110"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="125"/>
            <end position="144"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="147"/>
            <end position="173"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="606"/>
            <end position="615"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="616"/>
            <end position="625"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="14">
        <location>
            <begin position="1116"/>
            <end position="1126"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="1220"/>
            <end position="1229"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="14">
        <location>
            <begin position="1230"/>
            <end position="1239"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="1882"/>
            <end position="1891"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="1903"/>
            <end position="1926"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="2100"/>
            <end position="2111"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="2161"/>
            <end position="2170"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012513"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00047899"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00048679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029413.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2229" mass="241235" checksum="0B4A2F8853FC6E78" modified="2019-05-08" version="1">METGCAASSAPPRGGGGASPAAARPPPADGSAAVGGLGRRSGAGMETAAVPERGPAPGLPPRGPGAREPRVGHLRFLRRSVVDSDQEEPPLELSEAEPRRIVLLSKTRRIIAERAKERHGEADLEPPAVEPAAEPKGAVPGVPATAEEQKELGRDAERKEPSDKDQSKSKKQEPEEEADMKAVSTSPDGRFLKFDIELGRGSFKTVYKGLDTETWVEVAWCELQDRKLTKVERQRFKEEAEMLKGLQHPNIVRFYDFWESCVKGKKCIVLVTELMTSGTLKTYLKRFKVMKPKVLRSWCRQILKGLLFLHTRTPPIIHRDLKCDNIFITGPTGSVKIGDLGLATLKRASFAKSVIGTPEFMAPEMYEEHYDESVDVYAFGMCMLEMATSEYPYSECQNAAQIYRKVTCGVKPASFEKVTDPEIKEIIGECICKNKEERYEIKDLLSHAFFAEDTGVRVELAEEDHGRKSSIALRLWVEDPKKLKGKPKDNGAIEFTFDLEKETPDDVAQEMVDSGFFHESDVKIVAKSIRDRVALIQWRRERIWPMIQSEDHQVSECLEKLKVPQAQQVQVTSLSHAGQQTLSALEEPEADQHLFQHNLPTSVTSVASDSTFDSGQGSTVYSDSQSSQQSVIYSSLPDSVPPAIQRVYSPPLSESQALPQSLQQLGHYQQPSVPGLTVAQAPPAVLSQRPQHYQEPAMTFPVIQPVTVASSQMGQSQPVLASQQQPIPQQAPLQQVLASQPVCPLQPAAHLLPQYQTQTSQVAATSTQLTPLQISTVRQLPHVPPSQIPQLPVIPAVTPLAGLDNLPPSLSDLPAANVPSVPAPSQYFTPTVIIPSLPMNPALPLASNPPALSMQAVNLPHTTVASLVLPCQTIVPNMSAATIPLLAVAPPGVSALPTHHAISHLSQQQMYQPAFQQIIQSDAPSPHHTQNMQAVSQQQLPPLPQSLQPSIIHPSEQTMSTSGVGNQIIGHPQYGLEIGAQMPVIPMQPSIVMPPPLQLQPEILPPCLGPESLSQLHGNLAAASATVPIDHCLPLQPTGLLQLQPDLSQPLGQQAPCSAIQSVTESTQEEHAIQEKQQTLSLCCESYMSPDVASGKEMSDSFEGVVGSGKQEGKPSKKHKKSTRVRSRQEKSSRPKLTILNVCNTGDKMVECQLETHNHKMVTFKFDLDGDAPEEIATYMVEKEFILQTEKETFIEQMKDIIDKAEDMLSEDTEGERSSDQGTSPQQDASRLEMNEENRQSQVKTPMYQQNVLHTGKRWFIICPVVENPTEEVPEFSPAVPQSKQAEGPDPEQSDDQQMGSAVTDTPSVFFCHQLPSQTSVFDSPCGTPAPLAQVPAAASSAVLQTKAEILAATLPGTAPHVPEQFVTNVGQPEYPPLHLVMPSQPDASAHNAASLEGPSDAQLAPKSSLQTLQEMDEDSCIPDVEISCCTVRPPSSILTVPAQGAEFCPVPALSDVVVLELQPAPQIPQVSEASQPGVMRHSFVNQLLPGGVVSDPLSLTGSQLQSPPQLPVAVSQQHTMLTQSQPVTCTGLGVGPLQPPCAVESDGEGPPRVDFADDTIKSLDEKLRNLLYQECVPTSAASAGTPDSFVPLEQGESELASLPPLNEELIPTLVLDSREPGLNVPDTFQEANAAQDQCVSLVPSAMPPYPVLASQALTVPFIPINPAAMETSAVLNNATATSHSRTEFRIQEPMLEKFETTSIDVSSSKVRVYETGSAANISSTSAVTTQGLQVAPASSTIPEQMETSRRNKEAKIKISSALTENLIQTSTTDREINNLQRNDFVDPGSCGKQSEVHGNSTPAKTIGRFSVISTHDELTLTSPHCLRYSAPPDVYLDELPSSPDLKTSVRRVQTASFLDVLSDHGSSDSGDESLRRKSVAAQLSPQSSIAANDITKKPASLLQRSGKTSSPGPDSPNGQGTKIPTINITSFHSQSSYMSSDNDSEFEDADMKKELQNLREKHMKEIAELQTQQKNEIEALFVRLGKPLPPNVGFLHSVPPTGRRRRTSKNKLKAGKLLNPLVQQLKTAPSNTSNLSVCKESLPKETTKTHAGSTEATAVTSSTSATFGNSVQTQQPCSVKASLSSDICSGLATEGGNAHGSSGQGSSTNSLVTCPEPLPPPTLQVQVQANNSNNKKGTFTDDLHKLVDQWTSKTVGAAQSKPSLNQLKQNQKRQDMEPKANPMQAQNETCGILKMRMPLPPIDAGPAQLQYQAQGPMSAKEDEEEDVK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN10</accession>
    <name>A0A452IN10_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Neurabin-1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="14">Neurabin-I</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="13">Neural tissue-specific F-actin-binding protein I</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="15">Protein phosphatase 1 regulatory subunit 9A</fullName>
        </alternativeName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
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            <taxon>Archelosauria</taxon>
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            <taxon>Durocryptodira</taxon>
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        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
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        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
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        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
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        </citation>
        <scope>IDENTIFICATION</scope>
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    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
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</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN11</accession>
    <name>A0A452IN11_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Carbamoyl-phosphate synthase [ammonia], mitochondrial</fullName>
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        </recommendedName>
        <alternativeName>
            <fullName evidence="14">Carbamoyl-phosphate synthetase I</fullName>
        </alternativeName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
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                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="12">Involved in the urea cycle of ureotelic animals where the enzyme plays an important role in removing excess ammonia from the cell.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>hydrogencarbonate + NH4(+) + 2 ATP = carbamoyl phosphate + 2 ADP + phosphate + 2 H(+)</text>
            <dbReference type="Rhea" id="RHEA:18029"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:17544"/>
            <dbReference type="ChEBI" id="CHEBI:28938"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:58228"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="6.3.4.16"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Mitochondrion</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="6.3.4.16" evidence="10"/>
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        <property type="gene ID" value="ENSGAGG00000021129.1"/>
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        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004087">
        <property type="term" value="F:carbamoyl-phosphate synthase (ammonia) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004088">
        <property type="term" value="F:carbamoyl-phosphate synthase (glutamine-hydrolyzing) activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
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    <dbReference type="GO" id="GO:0006541">
        <property type="term" value="P:glutamine metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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    <dbReference type="GO" id="GO:0000050">
        <property type="term" value="P:urea cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
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        <property type="entry name" value="GATase1_CPSase"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Carbamoyl-phosphate synthase large chain"/>
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    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
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    </dbReference>
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        <property type="match status" value="2"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1490.20">
        <property type="entry name" value="ATP-grasp fold, A domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.470.20">
        <property type="entry name" value="ATP-grasp fold, B domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1030.10">
        <property type="entry name" value="Carbamoyl-phosphate synthetase, large subunit oligomerisation domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011761">
        <property type="entry name" value="ATP-grasp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013815">
        <property type="entry name" value="ATP_grasp_subdomain_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036897">
        <property type="entry name" value="CarbamoylP_synth_lsu_oligo_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002474">
        <property type="entry name" value="CarbamoylP_synth_ssu_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036480">
        <property type="entry name" value="CarbP_synth_ssu_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029062">
        <property type="entry name" value="Class_I_gatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005479">
        <property type="entry name" value="CPAse_ATP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005483">
        <property type="entry name" value="CPSase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035686">
        <property type="entry name" value="CPSase_GATase1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006275">
        <property type="entry name" value="CPSase_lsu"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005480">
        <property type="entry name" value="CPSase_lsu_oligo"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058047">
        <property type="entry name" value="CPSase_preATP-grasp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017926">
        <property type="entry name" value="GATASE"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016185">
        <property type="entry name" value="PreATP-grasp_dom_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01369">
        <property type="entry name" value="CPSaseII_lrg"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF003671">
        <property type="entry name" value="PRK05294.1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF009455">
        <property type="entry name" value="PRK12815.1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11405:SF53">
        <property type="entry name" value="CARBAMOYL-PHOSPHATE SYNTHASE [AMMONIA], MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11405">
        <property type="entry name" value="CARBAMOYLTRANSFERASE FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25596">
        <property type="entry name" value="CPSase_L_D1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02786">
        <property type="entry name" value="CPSase_L_D2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02787">
        <property type="entry name" value="CPSase_L_D3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00988">
        <property type="entry name" value="CPSase_sm_chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00117">
        <property type="entry name" value="GATase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00098">
        <property type="entry name" value="CPSASE"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00099">
        <property type="entry name" value="CPSGATASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM01096">
        <property type="entry name" value="CPSase_L_D3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01097">
        <property type="entry name" value="CPSase_sm_chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48108">
        <property type="entry name" value="Carbamoyl phosphate synthetase, large subunit connection domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52021">
        <property type="entry name" value="Carbamoyl phosphate synthetase, small subunit N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52317">
        <property type="entry name" value="Class I glutamine amidotransferase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56059">
        <property type="entry name" value="Glutathione synthetase ATP-binding domain-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52440">
        <property type="entry name" value="PreATP-grasp domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50975">
        <property type="entry name" value="ATP_GRASP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00866">
        <property type="entry name" value="CPSASE_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00867">
        <property type="entry name" value="CPSASE_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51273">
        <property type="entry name" value="GATASE_TYPE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0021">Allosteric enzyme</keyword>
    <keyword evidence="7 15" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="4" id="KW-0436">Ligase</keyword>
    <keyword evidence="9" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="6 15" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="8" id="KW-0809">Transit peptide</keyword>
    <keyword evidence="2" id="KW-0835">Urea cycle</keyword>
    <feature type="domain" description="ATP-grasp" evidence="16">
        <location>
            <begin position="518"/>
            <end position="710"/>
        </location>
    </feature>
    <feature type="domain" description="ATP-grasp" evidence="16">
        <location>
            <begin position="1060"/>
            <end position="1251"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004173"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022436"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022533"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022598"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00044063"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00047359"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00059545"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00074238"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00082675"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00409"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50975"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029341.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1376" mass="151207" checksum="956561D0BCE54FC1" modified="2019-05-08" version="1">MGDEGICALHHKWHGSRVPWNFLQTHHCLVFGSSLLAGLCWLFAPPAPCQQLSVCFSPRYTETLTDPSYRGQILTMVNPIVGNGGVPDTAALDEMGLSKFLESDGIKVSGLLVLDYSNDYSHWRAVKTLGQWLQEEKVRDLWATRPDTRHGISILRKLLSDSLSNGNQEREILEVKVYGKGNPIKVVAVDCGLKNNAIRLLVKRGAEVHLVPWNHDFTGMEYDGLFLSGGPGDPMQAQELIQNLQKVLKSDRKEPLFGVSMGNLITGIAAGATSYKMPMANRGQNQPVMNLSNGQAFITAQNHGYALQSNTLPAGWKPLFVNVNDQTNEGIMHETKPIFTAQFNPEASAGPTDTEFLFDSFISLIKKGKGTTVASVLPKARTVASRLEVSKVLILGSGGLSIGQAGEFDYSGSQAVKAMKEENVKTVLMNPNIASVQTNEVGLKQADTVYFLPITPQFVTEVIKAERPDGLILGMGGQTALNCGVELFRRGVLKEYGVKVLGTSVEAIMATEDRQLFSDKLTEINEKIAPSFAVESIEDALNAAEKIGYPVMIRSAYALGGLGSGICPDKERLLDLGTKAFAVTNQILVERSVVGWKEIEYEVVRDASDNCITVCNMENVDAMGVHTGDSIVVAPSQTLSNEEFQMLRESAIKVVRHLGIVGECNIQYALHPTSLEYCIIEVNARLSRSSALASKATGYPLAFIAAKIALGIPLPEIKNVVSGKTSACFEPSLDYMVAKIPRWDLDRFQGTSSQIGSSMKSVGEVMAIGRTFEECFQKALRMCHPSVDGFTSRLPMNKEWPANLDLQKELSDPSSIRVYAIAKALHDGVPVDDIHKLTAIDRWFIYKMRDIVNMEKSLKELSSESIPEETMRRAKQIGFSDKQIGKCLGVTEAQSRELRLKKNIKPSVKQIDTLAAEYPAITNYLYVTYNGQENDVKFDDHGMMVLGCGPYHIGSSVEFDWCAVSSIRTLRQLGHKTVVVNCNPETVSTDFDECDRLYFEELSLERILDIYQHEKCAGCIISVGGQIPNNLAVPLYKNGVKIMGTNPMQIDRAEDRSVFSAVLDELNVAQAPWKAVSTLKDAVEFANSVGYPCLLRPSYVLSGSAMNVVFTEEEMKKFLAEATRVSQEHPVVLTKFIEGAREVEMDAVAKAGKVVSHAISEHVEDAGVHSGDATLMLPTQNISQGALEKVKIITKKIARAFSISGPFNVQFLVRGNDVLVIECNLRASRSFPFVSKTIGVDFIDVATKVMIGEKIDESSLPTLERPIIPSEYVGIKAPMFSWPRLRDADPILRCEMASTGEVACFGESIHSAFLKAMLSTGFKFPQKGILVGIQVSTKSLQIRAKKGRRYVDRVTENNKSGEYSLLPVLRNSVFLR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN14</accession>
    <name>A0A452IN14_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Cysteine/serine-rich nuclear protein N-terminal domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the AXUD1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IN14"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033228.1">
        <property type="protein sequence ID" value="ENSGAGP00000029254.1"/>
        <property type="gene ID" value="ENSGAGG00000021162.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043565">
        <property type="term" value="F:sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006915">
        <property type="term" value="P:apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031972">
        <property type="entry name" value="CSRNP_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023260">
        <property type="entry name" value="Cys/Ser-rich_nuc_prot"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13580:SF10">
        <property type="entry name" value="CYSTEINE_SERINE-RICH NUCLEAR PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13580">
        <property type="entry name" value="TGF-BETA INDUCED APOPTOSIS PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16019">
        <property type="entry name" value="CSRNP_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR02031">
        <property type="entry name" value="CYSSERRICHNP"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0010">Activator</keyword>
    <keyword evidence="3" id="KW-0053">Apoptosis</keyword>
    <keyword evidence="9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="11" id="KW-0732">Signal</keyword>
    <keyword evidence="7" id="KW-0804">Transcription</keyword>
    <keyword evidence="4" id="KW-0805">Transcription regulation</keyword>
    <feature type="signal peptide" evidence="11">
        <location>
            <begin position="1"/>
            <end position="18"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019548841" description="Cysteine/serine-rich nuclear protein N-terminal domain-containing protein" evidence="11">
        <location>
            <begin position="19"/>
            <end position="532"/>
        </location>
    </feature>
    <feature type="domain" description="Cysteine/serine-rich nuclear protein N-terminal" evidence="12">
        <location>
            <begin position="22"/>
            <end position="240"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="293"/>
            <end position="330"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="72"/>
            <end position="99"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="293"/>
            <end position="310"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008548"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022703"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF16019"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029254.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="532" mass="59576" checksum="C0D36B81B962AB90" modified="2019-05-08" version="1">MCLFFISVFLATSILKKTKRLKKNNVEFDQVTVFYFPRCQGFTSVPSRGGCTLGMMSKHSSSKQFTLAEFSKEQENVRREKLKERLKEEKLEALKWKLTVNGTAESEEASQLTTEDISDDDVDVSNVELEDGFFLQPYPAKKRRALLKTMGVKKIDREEKRELHSIRLSREDCGCDCQEFCDPETCSCSLAGIKCQMDHTSFPCGCTKDGCGNTEGRIEFNQARVQTHFIHTIMKLELEKNQQSSERNVEPESSFRDRLHQFGCSAGKNVFEERAVPLTPAFQFNMDLETVGENSCSSDMTDSSISSIQSEDSEEQYESVPSDKSQSDIDDDGLARILHFNDSDIEDNSNNCQDNLSCFHPADFFSANIEDQCDTTAEKLSTGKYSGHLSNISECLDENANQDAGCFLDETADTHCDGLSCCVSSSAEQCSKSYTDLSLSSDSLDLFQSFSDYNLGPLYNSLKEYENLDNFSALQFQLPNFPGFPQAGDQSTCFLESLIGLSESVPETPVPFTDNQLLEDAIKSSLMETVKV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN22</accession>
    <name>A0A452IN22_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="3">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IN22"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033264.1">
        <property type="protein sequence ID" value="ENSGAGP00000029287.1"/>
        <property type="gene ID" value="ENSGAGG00000021186.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005814">
        <property type="term" value="C:centriole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026636">
        <property type="entry name" value="MPHOSPH9"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14926">
        <property type="entry name" value="M-PHASE PHOSPHOPROTEIN 9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14926:SF1">
        <property type="entry name" value="M-PHASE PHOSPHOPROTEIN 9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="405"/>
            <end position="432"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="830"/>
            <end position="904"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="555"/>
            <end position="722"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="405"/>
            <end position="417"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="418"/>
            <end position="432"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="854"/>
            <end position="865"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="866"/>
            <end position="875"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="880"/>
            <end position="893"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029287.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1115" mass="126212" checksum="D4DD73506B95075B" modified="2019-05-08" version="1">MSSPSSNPNGVSSFSGKTSRSVIPVEVLASFMHDIQKSGGTDSEILKNCETRWIQLFKLVEKQCQEQIVVQQEQFHHQIQCIQNEIRHLVKLQNSNASWDSFNSKSSPAKHANNFSSLESQMGFCSETFEETGCIVNQLKSNEAESQPKAPAEHQGCCVNSISMNSGYGILSASELNPRESKDIKKCIGYPEKISKGQGDAPVTKKDSIAPNVQTKKEFLRKNEKYCPSLKDSVLVFPAEFEHEVTEGHCGKTNCKYLTSWAQRLKQNQPKRTNTEEEEYMNSVQGNDQAKKLTFEKTSVTAAASPSYTFYLSQPNESPNSLASDASGLTYWKLDEKEMYHSLPENFRNEFSSIFVTKVSSDQVLTLDPTLHMKPGEQNPGHCFVSTLGEKTPFSPDSIVDPTFSSHSDTDSFSHTSNITSPQLGGSPKYTSSTKVLHLDPWENNAFQNETKIGCFFPMPNADTSNDLLINTEDEESLTLTPSLIIQSQLPCGDNNLPEYNISVTSFEDPVMLSKIRQNLRKKHARHIADLRAYYDSEIHSLKQQLEESHKTASSEELKKINQNLVDRCDELDAALNEASTRMRTLENKNNMLELQVTDWREHFYAVSNTSKVLQERIEEMRTSHKEKDNTISRLKSRLKDLEEAFEQAYKLSNNKDARLKEENKMLQDLLGEYESLGKEHERVKDTLNATENKLVDANTQISDLKRTISKLEAQIKQVEHEHMLKLRHITESHLRTSHANKLTASDVNRRKWLIPGAEYSIFTGQPLENQESIMDNRLEETYTPSRHHSPPEKDPSLEDSLTNVMAKKENEAPDTPIMKAFKELEEGKAFKSWGTQTEKEDTSTTTSHRRQTVGFNESTFSANRSPEKYRDQQRPKRYSSPSGQRSSSLPPSNRKANIPTGREIMLAPVSVTYSPKRSPKENLSPGFSHLVGKNENTMTRFNILLDDLETVPASTLQHSNPRKRLQFLSLDDTEGRQHSGKGIDSCIKHSSCTESLHTGMKKGTVASAWEERNIPHTNEQSIPPPVILYDADSKYTSRIKTLAETERLFDELTQEKQQIEAALSRIPSSGGRTTLQARLNQETLEDRLERINRDLGSIRMTLKRFHVLRTSANL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN23</accession>
    <name>A0A452IN23_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">ACB domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Peroxisome</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IN23"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033229.1">
        <property type="protein sequence ID" value="ENSGAGP00000029255.1"/>
        <property type="gene ID" value="ENSGAGG00000021163.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005777">
        <property type="term" value="C:peroxisome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004165">
        <property type="term" value="F:delta(3)-delta(2)-enoyl-CoA isomerase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000062">
        <property type="term" value="F:fatty-acyl-CoA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd06558">
        <property type="entry name" value="crotonase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.226.10:FF:000084">
        <property type="entry name" value="Enoyl-CoA delta isomerase 2, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.80.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.226.10">
        <property type="entry name" value="2-enoyl-CoA Hydratase, Chain A, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.12.10">
        <property type="entry name" value="Lyase 2-enoyl-coa Hydratase, Chain A, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Acyl-CoA-binding_prot_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000582">
        <property type="entry name" value="Acyl-CoA-binding_protein"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035984">
        <property type="entry name" value="Acyl-CoA-binding_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029045">
        <property type="entry name" value="ClpP/crotonase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051053">
        <property type="entry name" value="ECH/Chromodomain_protein"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001753">
        <property type="entry name" value="Enoyl-CoA_hydra/iso"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014748">
        <property type="entry name" value="Enoyl-CoA_hydra_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014352">
        <property type="entry name" value="FERM/acyl-CoA-bd_prot_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43684">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43684:SF1">
        <property type="entry name" value="ENOYL-COA DELTA ISOMERASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00887">
        <property type="entry name" value="ACBP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00378">
        <property type="entry name" value="ECH_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00689">
        <property type="entry name" value="ACOABINDINGP"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47027">
        <property type="entry name" value="Acyl-CoA binding protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52096">
        <property type="entry name" value="ClpP/crotonase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00880">
        <property type="entry name" value="ACB_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51228">
        <property type="entry name" value="ACB_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0413">Isomerase</keyword>
    <keyword evidence="2" id="KW-0576">Peroxisome</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="ACB" evidence="4">
        <location>
            <begin position="4"/>
            <end position="89"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004275"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023140"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023235"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS51228"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029255.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="358" mass="39935" checksum="B7C1C4E5B2647BD5" modified="2019-05-08" version="1">MRVSQEDFEKAKDQLKLLKEDPGNEEKLKLYALFKQATEGPCSSPKPGMLDFVKKAKWDAWSSLGSLSKDSARQKYVDLVSSLVSSESSSQVKDTTPDSKHGYETLQVTTTDHITKIMLNRPEKKNAITTQMYKEIIQALEEAAKDDSVITVITGNGDYYCSGNDLNNFTNIPSGGIEKMAKDGAVLLENFVKHFIDFPKPLIAVVNGPAVGISVTLLGLFDIVYATDRATFHTPFSELGQSPEGCSSYMFPKIMGLSKANEMLLFNKKLTAGEAYAQGLVTEVFPDRTFEKEVWIRLKAYANLPKNSLALSKQLIRGVEKEKLHAVNCQECERLRERWLSDECLNAVMSFFGKKSKL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN29</accession>
    <name>A0A452IN29_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">SMB domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IN29"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033342.1">
        <property type="protein sequence ID" value="ENSGAGP00000029361.1"/>
        <property type="gene ID" value="ENSGAGG00000020186.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009986">
        <property type="term" value="C:cell surface"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004551">
        <property type="term" value="F:dinucleotide phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003676">
        <property type="term" value="F:nucleic acid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004528">
        <property type="term" value="F:phosphodiesterase I activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046034">
        <property type="term" value="P:ATP metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030505">
        <property type="term" value="P:inorganic diphosphate transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045599">
        <property type="term" value="P:negative regulation of fat cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009143">
        <property type="term" value="P:nucleoside triphosphate catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030500">
        <property type="term" value="P:regulation of bone mineralization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd16018">
        <property type="entry name" value="Enpp"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00091">
        <property type="entry name" value="NUC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.410.20:FF:000003">
        <property type="entry name" value="Ectonucleotide pyrophosphatase/phosphodiesterase family member 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.410.20:FF:000001">
        <property type="entry name" value="Ectonucleotide pyrophosphatase/phosphodiesterase family member 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.720.10:FF:000145">
        <property type="entry name" value="Uncharacterized protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.410.20">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.720.10">
        <property type="entry name" value="Alkaline Phosphatase, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.570.10">
        <property type="entry name" value="Extracellular Endonuclease, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017850">
        <property type="entry name" value="Alkaline_phosphatase_core_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044929">
        <property type="entry name" value="DNA/RNA_non-sp_Endonuclease_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001604">
        <property type="entry name" value="Endo_G_ENPP1-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020821">
        <property type="entry name" value="ENPP1-3/EXOG-like_nuc-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044925">
        <property type="entry name" value="His-Me_finger_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002591">
        <property type="entry name" value="Phosphodiest/P_Trfase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036024">
        <property type="entry name" value="Somatomedin_B-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001212">
        <property type="entry name" value="Somatomedin_B_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10151">
        <property type="entry name" value="ECTONUCLEOTIDE PYROPHOSPHATASE/PHOSPHODIESTERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10151:SF77">
        <property type="entry name" value="ECTONUCLEOTIDE PYROPHOSPHATASE_PHOSPHODIESTERASE FAMILY MEMBER 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01223">
        <property type="entry name" value="Endonuclease_NS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01663">
        <property type="entry name" value="Phosphodiest"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01033">
        <property type="entry name" value="Somatomedin_B"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00892">
        <property type="entry name" value="Endonuclease_NS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00477">
        <property type="entry name" value="NUC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00201">
        <property type="entry name" value="SO"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53649">
        <property type="entry name" value="Alkaline phosphatase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54060">
        <property type="entry name" value="His-Me finger endonucleases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90188">
        <property type="entry name" value="Somatomedin B domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00524">
        <property type="entry name" value="SMB_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50958">
        <property type="entry name" value="SMB_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="4" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="7">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019372890" description="SMB domain-containing protein" evidence="7">
        <location>
            <begin position="24"/>
            <end position="831"/>
        </location>
    </feature>
    <feature type="domain" description="SMB" evidence="8">
        <location>
            <begin position="31"/>
            <end position="71"/>
        </location>
    </feature>
    <feature type="domain" description="SMB" evidence="8">
        <location>
            <begin position="72"/>
            <end position="116"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50958"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029361.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="831" mass="95563" checksum="0BDE0F498662833E" modified="2019-05-08" version="1">PDPFKLLVLSACVLTIILGCIFGLKPSCSRDVKTCKGRCFERTFGSCRCDRDCVELGNCCLDFQETCIQPAHIWTCNKFRCGEKRWPENRCSCSDDCVENKDCCVNYYIVCQGAKSWVEEKCEDINEPQCPPGFAKPPVLLFSLDGFRAEYLHTWSGLLPVISKLQKCGTYTPNMRPVYPTKTFPNHYTIVTGLYPESHGIIDNKIYDPHRNVSFTLKNAEKFNPQWYQGQPIWLTAMHQGLKAGTFFWPGSDVRINGTFPDLYRKYNGSIPFEERVIAILHWLQLPKDERPHFYTLYLEEPDSSGHNFGPVSSGVILALQRVDNIVGMLMDGLKQMNLHKCLNIILLSDHGMETASCSKTAYLSTYMENVQDITVIPGPAARLRPQNVPDEYFSYPDQHFKAFMKELLPKRFHYVNNDRIERVHFYLDPQWQLARKPSEIKYCNGGFHGSDNRFMSMQAIFIGFGPGFKFKTQVDPFENIEIYNLICDLLGITPAYNNGTHGSLNHLLKNPVYIPRHPEEESHPSVYPLMKTTSSLDIGCSCDLAGLPIRDFQQQLNHTASEVKKIEKYSLPYGRPRVLQKRSTYYLLYHHQYVSGYSQDLNMTLWSAYTDNRTASVENLSNCLHVDIRIPSSSSKTCSFYKNHHQLSYGFLFPPNTKKSSYEVLLNSNIVPMYHEFKVLWNYFYDVLLPKYTLARNGVNVVSGPVFDYDSNGLYDTPEKVKRLTHDPEVLVPTHYFIVLTSCKNISQTPLQCEGALDALSFIVPHRPDNSESCARHKRDSLWVEERMRFHTARVRDVELLTGLSFYQDRKQPVSDILQLKTYLPTFPME</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN30</accession>
    <name>A0A452IN30_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Rab3 GTPase-activating protein catalytic subunit</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Endoplasmic reticulum</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Golgi apparatus</location>
            <location evidence="1">cis-Golgi network</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the Rab3-GAP catalytic subunit family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IN30"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033243.1">
        <property type="protein sequence ID" value="ENSGAGP00000029269.1"/>
        <property type="gene ID" value="ENSGAGG00000021013.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005783">
        <property type="term" value="C:endoplasmic reticulum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045700">
        <property type="entry name" value="Rab3GAP1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045698">
        <property type="entry name" value="Rab3GAP1_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026147">
        <property type="entry name" value="Rab3GAP1_conserved"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21422">
        <property type="entry name" value="RAB3 GTPASE-ACTIVATING PROTEIN CATALYTIC SUBUNIT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21422:SF9">
        <property type="entry name" value="RAB3 GTPASE-ACTIVATING PROTEIN CATALYTIC SUBUNIT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19533">
        <property type="entry name" value="Rab3-GAP_cat_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13890">
        <property type="entry name" value="Rab3-GTPase_cat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="9" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="6" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Rab3GAP catalytic subunit conserved" evidence="11">
        <location>
            <begin position="389"/>
            <end position="543"/>
        </location>
    </feature>
    <feature type="domain" description="Rab3GAP catalytic subunit C-terminal" evidence="12">
        <location>
            <begin position="555"/>
            <end position="758"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="303"/>
            <end position="350"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="363"/>
            <end position="398"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="309"/>
            <end position="319"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="322"/>
            <end position="344"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="373"/>
            <end position="384"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004222"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00008856"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00015817"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF13890"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF19533"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029269.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="758" mass="85164" checksum="877D1ADFB47B3658" modified="2019-05-08" version="1">GCPLTPLPPVSIAIRLTYVLQDWQQYFWPQQPPDIDALIGGEVGGLEFGKLPFGACEDPVSELHLAATWPHLTEGIIVDNDVYSDLDPIQAPQWSVRVRKADNLQCLLGDFLTEFFKLCRRKESTDEILGRSTFEEDGKEVADITHALSKLTEPAPVPIHKLSVTNMVHSAKKKIRKHRGVDDSPLNNEVLNTLMLVSKLSMHPSYCVHVYINLYSQFKSAPSDSLTYKLALCLCMINFYHGGVKGVAHLWQEFVVEMRRRWEKNYLIPELANGSPDLRCCLLHQKLQMLNCCIERKKARDEGKKPYTSDRSPSGSSGDTGKAVDRSGGDYQKETEKERGEVGKSWESWSDSEEEFFECLSDTEELKGNGQENGKKGGTKEGNKESANLKPEGRLHPHGKLTLLHPGEPLYIPVTQEPAPMTEDLLEEQSEVLAKLGTSAEGAHLRARMQSACLLSDMESFKAANPGCCLEDFVRWYSPRDYIEEEVVDEEGNRVLKGELSARMKIPSNMWVEAWETAKPVPARRQRRLFDDTREAEKVLHYLAVQKPADLARHLLPCVIHAAVLKVKEEETLEDIASVKKIIKQIISHSSKVLRFPNPEDKKLEEIIAQIMSAEAIIARARSLKAKFGIEKCDQEEEKEDLERFVNCLLEQPEVSVIGAGRGPAGSIIHKLFLNAQRVATMTSLDEELKRSGSSDERRLNLGAVTDFPPPAGRELILRITVPRPAPYSKPLPQRMYSVITKEDFRLAGAFSADTTFF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN32</accession>
    <name>A0A452IN32_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Aminopeptidase NAALADL1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="17">N-acetylated-alpha-linked acidic dipeptidase-like protein</fullName>
        </alternativeName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">Aminopeptidase with broad substrate specificity. Has lower activity with substrates that have Asp or Glu in the P2' position, or Pro in the P3' position. Lacks activity with substrates that have both Pro in the P3' position and Asp or Glu in the P2' position. Lacks carboxypeptidase activity. Lacks dipeptidyl-peptidase IV type activity.</text>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Apical cell membrane</location>
            <topology evidence="2">Single-pass type II membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the peptidase M28 family. M28B subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IN32"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033485.1">
        <property type="protein sequence ID" value="ENSGAGP00000029489.1"/>
        <property type="gene ID" value="ENSGAGG00000020963.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016324">
        <property type="term" value="C:apical plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004177">
        <property type="term" value="F:aminopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004180">
        <property type="term" value="F:carboxypeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016805">
        <property type="term" value="F:dipeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008237">
        <property type="term" value="F:metallopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.630.10:FF:000101">
        <property type="entry name" value="N-acetylated alpha-linked acidic dipeptidase like 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.930.40:FF:000001">
        <property type="entry name" value="N-acetylated-alpha-linked acidic dipeptidase 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.50.30.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.930.40">
        <property type="entry name" value="Transferrin receptor-like, dimerisation domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.630.10">
        <property type="entry name" value="Zn peptidases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046450">
        <property type="entry name" value="PA_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007484">
        <property type="entry name" value="Peptidase_M28"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039373">
        <property type="entry name" value="Peptidase_M28B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007365">
        <property type="entry name" value="TFR-like_dimer_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036757">
        <property type="entry name" value="TFR-like_dimer_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10404:SF36">
        <property type="entry name" value="GLUTAMATE CARBOXYPEPTIDASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10404">
        <property type="entry name" value="N-ACETYLATED-ALPHA-LINKED ACIDIC DIPEPTIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04389">
        <property type="entry name" value="Peptidase_M28"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04253">
        <property type="entry name" value="TFR_dimer"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52025">
        <property type="entry name" value="PA domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47672">
        <property type="entry name" value="Transferrin receptor-like dimerisation domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53187">
        <property type="entry name" value="Zn-dependent exopeptidases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0031">Aminopeptidase</keyword>
    <keyword evidence="9" id="KW-0106">Calcium</keyword>
    <keyword evidence="10" id="KW-0224">Dipeptidase</keyword>
    <keyword evidence="12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="6" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-0482">Metalloprotease</keyword>
    <keyword evidence="14" id="KW-0511">Multifunctional enzyme</keyword>
    <keyword evidence="5" id="KW-0645">Protease</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="Peptidase M28" evidence="19">
        <location>
            <begin position="216"/>
            <end position="374"/>
        </location>
    </feature>
    <feature type="domain" description="Transferrin receptor-like dimerisation" evidence="18">
        <location>
            <begin position="442"/>
            <end position="562"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004655"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005634"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022438"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022997"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023049"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023268"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00059290"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00068168"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00081462"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF04253"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="Pfam" id="PF04389"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029489.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="565" mass="63966" checksum="5B8C990BFF3D1BE5" modified="2019-05-08" version="1">KDIKMVSVSSNFTQRPHLAGTAENLRLAQQIQAQWKEFGLDSVELAHYDVLLSYPNETNPNYISIIDEHGTEIFNTSLFEPPPPGYETVTDVVPPYSAFSAPGVPEVKRLELLNSRLPALSKINTSLIPPEYTYRYEEARAVGLPKLPVHPIGYNDAEKLLRDMGGPAPPDSSWKGSLNVSYNIGPGFMSRYSTRKVKMHIHSSNKVTRIYNVIGTIRGATEPGWRPRRTIIFASWDAEEFGLLGSTEWAEENTKLLQARGVAYINADSSVEGNYTLRIDCTPLMYSLVYSLTREIPSPDEGFKGKSLYESWHENNPSTEYKDVPRINKLGSGNDFEVFFQRLGIASGRARYSKNWRVEKYSSYPVYHSVYETYEIVERFYDPTFKNHLTVAQVRGGLVFELADSVVLPFDCRDYAEALRNYVQVIYNLSKNHQMGLTTYSVSFDSLFSAANNFTEAATDFHRRLQQVDINNPIAVRALNDQLMFLERAFIDPLGLPGRPFYRHIIFAPSSHNKYAGESFPGIYDAMFDIENKADQRKAWEEVKRQISIAAFTVQAAAGTLKDLA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN33</accession>
    <name>A0A452IN33_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Serine protease</fullName>
            <ecNumber evidence="4">3.4.21.-</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="4">Belongs to the peptidase S1B family.</text>
    </comment>
    <dbReference type="EC" id="3.4.21.-" evidence="4"/>
    <dbReference type="AlphaFoldDB" id="A0A452IN33"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033239.1">
        <property type="protein sequence ID" value="ENSGAGP00000029265.1"/>
        <property type="gene ID" value="ENSGAGG00000021168.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000785">
        <property type="term" value="C:chromatin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008236">
        <property type="term" value="F:serine-type peptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006260">
        <property type="term" value="P:DNA replication"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.10.120">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009003">
        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008256">
        <property type="entry name" value="Peptidase_S1B"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14389:SF3">
        <property type="entry name" value="PROTEIN FAM111A-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14389">
        <property type="entry name" value="SI:CH1073-475A24.1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13365">
        <property type="entry name" value="Trypsin_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00839">
        <property type="entry name" value="V8PROTEASE"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50494">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 4" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="1 4" id="KW-0645">Protease</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 4" id="KW-0720">Serine protease</keyword>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="1"/>
            <end position="25"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="RuleBase" id="RU004296"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029265.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="558" mass="63823" checksum="90BA44F07C145E35" modified="2025-10-08" version="2">MKRDITENSPLQNPPPKRKRSRQTRESLLITDIHARLDKEKGKEIHLLGKKGIEGYVNLGMPLQYVPENSHFEMKFYKVNRNPKGVVGYRQFESKGKKCVIFYVSSFGNRLENNARYRILWCRQLVKDLSKLCVFAPEGETIKDALCKDGRFHSCLEEKEWKLVENGKCRTSNHCVDNLANKSFEVVVKTKQPFKARDCSRNDQSCMASEEGQGKTYHYFKPILLDLYPDLKKQSTVIDELFAKAFAKALEEGKSKRAVFKLYRENFGKETKNSILIKVHKRLGVLSDSVGFIEWANNGNNGSATCFVLHDKYVLTCHHDWASIISHSARVTFSYEDKYPKENSWFSLAPWFEISDKDLDFAVLKLEGDRSSLPAGLVRFSHPLPFNGLIYIIGHPDGQAKSADGCTVVPVFERKQECHCRVQRGQKVCNSVRCGSEDRQCIHMFTQRSFQEVINNPNVVTYDTSFFFGSSGSPVFNADGQLVAMHTAGYKYEKNKQLHSIVEFGFSMVAILAAIKKKYKAWYEFELLSLGEDARSCPGEHENFTSAYTNDIEMENLE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN34</accession>
    <name>A0A452IN34_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="10">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IN34"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033347.1">
        <property type="protein sequence ID" value="ENSGAGP00000029366.1"/>
        <property type="gene ID" value="ENSGAGG00000021101.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004222">
        <property type="term" value="F:metalloendopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016485">
        <property type="term" value="P:protein processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd08662">
        <property type="entry name" value="M13"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.390.10">
        <property type="entry name" value="Collagenase (Catalytic Domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.1380.10">
        <property type="entry name" value="Neutral endopeptidase , domain2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024079">
        <property type="entry name" value="MetalloPept_cat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000718">
        <property type="entry name" value="Peptidase_M13"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018497">
        <property type="entry name" value="Peptidase_M13_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042089">
        <property type="entry name" value="Peptidase_M13_dom_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008753">
        <property type="entry name" value="Peptidase_M13_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11733:SF141">
        <property type="entry name" value="MEMBRANE METALLO-ENDOPEPTIDASE-LIKE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11733">
        <property type="entry name" value="ZINC METALLOPROTEASE FAMILY M13 NEPRILYSIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01431">
        <property type="entry name" value="Peptidase_M13"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05649">
        <property type="entry name" value="Peptidase_M13_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00786">
        <property type="entry name" value="NEPRILYSIN"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55486">
        <property type="entry name" value="Metalloproteases ('zincins'), catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51885">
        <property type="entry name" value="NEPRILYSIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="6" id="KW-0482">Metalloprotease</keyword>
    <keyword evidence="2" id="KW-0645">Protease</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="Peptidase M13 N-terminal" evidence="9">
        <location>
            <begin position="44"/>
            <end position="460"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase M13 C-terminal" evidence="8">
        <location>
            <begin position="521"/>
            <end position="727"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="464"/>
            <end position="507"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023049"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF01431"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF05649"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029366.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="728" mass="83808" checksum="4AF6D2D95897A203" modified="2019-05-08" version="1">MALCGFIPPTDMQYGTNSGSICTTPGCVTAAARILQNMDPTTEPCTDFYQYACGGWLNWHVIPETSSRYSIFDILRDELEIILKGVLESPEQGDREAFRKAKMLYTSCMNESFIEQRDSLPLLEVLAMVGDWPVASADWNITKGNKPNWSMEEKLSTLNSRFNKRVFIDMFVWNDDQDSSRHIIYIDQPSLGMPSRDYYFNGGNYQRVREAYLQFMITIAKMIREDKNMAKDDTFVQQEMAKVMKLETEIANATAPAEERHDVTLLYNKMTLRELQEKFALNVSLGIGVCLSLPSQEFNWTLFIQGIMSSVNVQIHLDEEVVVYGAPYLQELKAIISNYSASTIQNYLVWRLVTDRVSSLSRRFKDARAHYRKALYGTTLEEARWRECVSYVNNNMENAVGALYVRETFAGESKRMVRDLIDKIREVFIETLDELQWMDETSKEKAREKAMAIKEQIGYPDYILEDQNEKLDQEYANLNFSEHKYFENSLENLRAGAQKSLKKLRERVDQDIWIIGAAVVNAFYSPNRNQIVFPAGILQPPFFSKHQPQALNFGGIGMVIGHEITHGFDDNGRNFDKDGNMFDWWSNFSAMHFKDQSRCMVYQYGNYTWELAGGQNVSGISTLGENIADNGGVRQAYKAYLKWLEREGKEPKLPGLDLTHKQLFFLNFAQVWCGSYRPEYASQSIKTDVHSPLKYRVMGSLQNFEAFSEAFHCKKGTAMHPVEKCRVW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN37</accession>
    <name>A0A452IN37_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">glutamate carboxypeptidase II</fullName>
            <ecNumber evidence="18">3.4.17.21</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>Release of an unsubstituted, C-terminal glutamyl residue, typically from Ac-Asp-Glu or folylpoly-gamma-glutamates.</text>
            <dbReference type="EC" id="3.4.17.21"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Membrane</location>
            <topology evidence="2">Single-pass type II membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the peptidase M28 family. M28B subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.4.17.21" evidence="18"/>
    <dbReference type="AlphaFoldDB" id="A0A452IN37"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033490.1">
        <property type="protein sequence ID" value="ENSGAGP00000029494.1"/>
        <property type="gene ID" value="ENSGAGG00000020963.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016805">
        <property type="term" value="F:dipeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004181">
        <property type="term" value="F:metallocarboxypeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd08022">
        <property type="entry name" value="M28_PSMA_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.930.40:FF:000001">
        <property type="entry name" value="N-acetylated-alpha-linked acidic dipeptidase 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.630.10:FF:000009">
        <property type="entry name" value="N-acetylated-alpha-linked acidic dipeptidase 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.50.30.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.930.40">
        <property type="entry name" value="Transferrin receptor-like, dimerisation domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.630.10">
        <property type="entry name" value="Zn peptidases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046450">
        <property type="entry name" value="PA_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007484">
        <property type="entry name" value="Peptidase_M28"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039373">
        <property type="entry name" value="Peptidase_M28B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007365">
        <property type="entry name" value="TFR-like_dimer_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036757">
        <property type="entry name" value="TFR-like_dimer_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10404:SF36">
        <property type="entry name" value="GLUTAMATE CARBOXYPEPTIDASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10404">
        <property type="entry name" value="N-ACETYLATED-ALPHA-LINKED ACIDIC DIPEPTIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04389">
        <property type="entry name" value="Peptidase_M28"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04253">
        <property type="entry name" value="TFR_dimer"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52025">
        <property type="entry name" value="PA domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47672">
        <property type="entry name" value="Transferrin receptor-like dimerisation domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53187">
        <property type="entry name" value="Zn-dependent exopeptidases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0121">Carboxypeptidase</keyword>
    <keyword evidence="12" id="KW-0224">Dipeptidase</keyword>
    <keyword evidence="15" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="14" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13" id="KW-0482">Metalloprotease</keyword>
    <keyword evidence="16" id="KW-0511">Multifunctional enzyme</keyword>
    <keyword evidence="5" id="KW-0645">Protease</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0735">Signal-anchor</keyword>
    <keyword evidence="6" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="11" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="9" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="Peptidase M28" evidence="20">
        <location>
            <begin position="230"/>
            <end position="432"/>
        </location>
    </feature>
    <feature type="domain" description="Transferrin receptor-like dimerisation" evidence="19">
        <location>
            <begin position="500"/>
            <end position="620"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004606"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005634"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022645"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022997"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023049"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023268"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00052003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00066561"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="Pfam" id="PF04253"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="Pfam" id="PF04389"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029494.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="623" mass="70422" checksum="6A560AE6BCAD4A92" modified="2019-05-08" version="1">MTGKLTSNYKIQFLVLTANTCPFLFPSNFTQRPHLAGTAENLRLAQQIQAQWKEFGLDSVELAHYDVLLSYPNETNPNYISIIDEHGTEIFNTSLFEPPPPGYETVTDVVPPYSAFSAPGVPEVKRLELLNSRKTIISAKINTSLIPPEYTYRYEEARAVGLPKLPVHPIGYNDAEKLLRDMGGPAPPDSSWKGSLNVSYNIGPGFMSRYSTRSVKMHIHSSNKVTRIYNVIGTIRGATEPDRYVILGGHRDSWVFGGIDPQSGAAVVHEIVRSYGKLKKEGWRPRRTIIFASWDAEEFGLLGSTEWAEENTKLLQARGVAYINADSSVEGNYTLRIDCTPLMYSLVYSLTREIPSPDEGFKGKSLYESWHENNPSTEYKDVPRINKLGSGNDFEVFFQRLGIASGRARYSKNWRVEKYSSYPVYHSVYETYEIVERFYDPTFKNHLTVAQVRGGLVFELADSVVLPFDCRDYAEALRNYVQVIYNLSKNHQMGLTTYSVSFDSLFSAANNFTEAATDFHRRLQQVDINNPIAVRALNDQLMFLERAFIDPLGLPGRPFYRHIIFAPSSHNKYAGESFPGIYDAMFDIENKADQRKAWEEVKRQISIAAFTVQAAAGTLKDLA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN39</accession>
    <name>A0A452IN39_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="9">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000033257.1">
        <property type="protein sequence ID" value="ENSGAGP00000029280.1"/>
        <property type="gene ID" value="ENSGAGG00000020783.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005938">
        <property type="term" value="C:cell cortex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0120025">
        <property type="term" value="C:plasma membrane bounded cell projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903479">
        <property type="term" value="P:mitotic actomyosin contractile ring assembly actin filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd21276">
        <property type="entry name" value="CH_IQGAP3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.418.10:FF:000013">
        <property type="entry name" value="IQ motif containing GTPase activating protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.506.10:FF:000004">
        <property type="entry name" value="IQ motif containing GTPase activating protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.418.10">
        <property type="entry name" value="Calponin-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.506.10">
        <property type="entry name" value="GTPase Activation - p120gap, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001715">
        <property type="entry name" value="CH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036872">
        <property type="entry name" value="CH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000048">
        <property type="entry name" value="IQ_motif_EF-hand-BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000593">
        <property type="entry name" value="RasGAP_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023152">
        <property type="entry name" value="RasGAP_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001936">
        <property type="entry name" value="RasGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001202">
        <property type="entry name" value="WW_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149">
        <property type="entry name" value="RAS GTPASE-ACTIVATING PROTEIN WITH IQ MOTIF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14149:SF10">
        <property type="entry name" value="RAS GTPASE-ACTIVATING-LIKE PROTEIN IQGAP3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00307">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00612">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00616">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03836">
        <property type="entry name" value="RasGAP_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00033">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00015">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00323">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47576">
        <property type="entry name" value="Calponin-homology domain, CH-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF143885">
        <property type="entry name" value="RGC domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50021">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00509">
        <property type="entry name" value="RAS_GTPASE_ACTIV_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50018">
        <property type="entry name" value="RAS_GTPASE_ACTIV_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01159">
        <property type="entry name" value="WW_DOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50020">
        <property type="entry name" value="WW_DOMAIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="4" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="1" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="Calponin-homology (CH)" evidence="8">
        <location>
            <begin position="35"/>
            <end position="150"/>
        </location>
    </feature>
    <feature type="domain" description="WW" evidence="7">
        <location>
            <begin position="668"/>
            <end position="701"/>
        </location>
    </feature>
    <feature type="domain" description="Ras-GAP" evidence="6">
        <location>
            <begin position="964"/>
            <end position="1197"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="4">
        <location>
            <begin position="159"/>
            <end position="186"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="1"/>
            <end position="11"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="12"/>
            <end position="22"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50020"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029280.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1582" mass="180571" checksum="BD4043533C66DE71" modified="2019-05-08" version="1">PSSCRESQARVSSDERLTADEMDEQRRQNVAYQYLCRLEEAKRWMEACLNEELASPTELEESLRNGVILAKLGHCFAPDIIPLKKIYDREQMRYEATGLHFRHTDNINHWREAMAHLGLPTIFHPETTDIYDKKNMPRVVYCIHALSLYLFKLGLAPQIQDLYGKVNFTEEEINNMKRELEKYGLQMPAFSKIGGILASELSVDEAAVHAAVLAINEAVERGVVEQTMVALHNPSAMLLNLRENLAAVYQEMLFQVKMEKAGNARNRVRADEDIYDRCLTQAEIQGNINKVNVHGALEYVDDALERQDPLALCQALQDPVLALRGVQRENAAWYLDQLSTDREQKALELGYVDLLEQEEVQAGVYAANERGDQEHAMRRAVSRINASIRRGEAMETVRELMSPEAQLPKAFPFAAQLYQHELSLLQQHHPQGELGQEELYVAVEMLSAVALIDRALEAGDVSGFWSSLVSPALGLSDIEDANAQRYFDDLLKLKRRSGKAGVAFLSWNDIQACVNSANLAVQDENDKILAIRLINEALAQDDPEKTLTALLLPSARFSAVTLPGARRYHDVLTAALKQKAQVTGDDGAALWWEEIQQGVSKANQDTEAARRLALGIAAINQAIKEGKPAQTARVLRNPDVSLRGIVAECAGAYQAQLAALLAAKRQAGNANHNWVRHQLKDGSVYYFSLQNFEGSWERPHDCVLNSTHLSREEIQSTITGVTAAHDRERLWESNAGFVVRLQARMRGYLVRQEFVARRHLLQEQLPAVIKIQVRSAARLPSRSHADGHDDIYIIQTYVRMWRARKRYRERLRYFRKNVSVHARDPPLSIVRRFVHLLEQSQHDFWEESELLKLREEVVKRIRSNQQLESDLNLMDIKIGLLVKNRITLQEVVSHCKKLTRKNKEQLSNLMAIDKQKGLKSLSKEKRQKLEAYQHLFYLLQTHPVYLAKLIFQMPQNKSTKFMESVVFTLYNYASNPREAYLLLQLFKTALQEEIRSKVDQLRDILTGNPTVIRLVVSFYRNARGQNALRQILGSPVQEVLQDKTLSIRTNPVDIYKAWINQLESQSGQKSKFPYDVSPEQALSHPEVQRRLDISIRNLLAVTDKFLSAITSSVDKIPYGMRYVAKVLKTSLAEKFPAAGEEDICKIVGNLLYYRFMNPAVVAPDGFDIVDISAGVALHPEHRRNLGSIAKILQHAAANKMFEGENGHLRVVNQYLEETHSKFRKFICAACSVPEPEERFNVDEYSEMVAVAKPVIYITVAELINTHKLLLEHQDSISPDHKDPLHELLEDLDELPTVQSLIGTCRLLVSFKPAPPLVSSFNTKQMLVDVIQSQPGDSLLEILRTAASEDQEAFHHHQMHRRALREAQTPDRLKRNRSLAGNSQLSMEEKKRKVIRNLRRLENLGVVDSANQYQEIINEMAKDIRNQRRYRQHRKGELVKLRQTLCGLNSKTSFYEEQIDYYNQYIRTCLDNLAASSKYVGKKQQSLHYTAARLFEKGVLLEIEDLPVSQFRNVIFDIIPCGEAGKFQVKAKFMGIDMEKFQLHYQDLLQLQYEGVAVMKLFDKAKVNVNLLIFLLNKKFFRK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN45</accession>
    <name>A0A452IN45_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Cyclin-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="2">Belongs to the cyclin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IN45"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033262.1">
        <property type="protein sequence ID" value="ENSGAGP00000029285.1"/>
        <property type="gene ID" value="ENSGAGG00000021185.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016538">
        <property type="term" value="F:cyclin-dependent protein serine/threonine kinase regulator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.472.10:FF:000009">
        <property type="entry name" value="cyclin-T2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.472.10">
        <property type="entry name" value="Cyclin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013763">
        <property type="entry name" value="Cyclin-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036915">
        <property type="entry name" value="Cyclin-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043198">
        <property type="entry name" value="Cyclin/Ssn8"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006671">
        <property type="entry name" value="Cyclin_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10026">
        <property type="entry name" value="CYCLIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00134">
        <property type="entry name" value="Cyclin_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00385">
        <property type="entry name" value="CYCLIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47954">
        <property type="entry name" value="Cyclin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="1 2" id="KW-0195">Cyclin</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Cyclin-like" evidence="3">
        <location>
            <begin position="58"/>
            <end position="157"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00023127"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="RuleBase" id="RU000383"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="SMART" id="SM00385"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029285.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="182" mass="20259" checksum="A7ABBBADD9A46654" modified="2019-05-08" version="1">MAAAASGPGGAGSAARGGGASLSASRWFFSREQLENTPTRRCGVEADKELSYRQQAANLIQDMGQRLNVSQLTINTAIVYMHRFYMHHSFTKFNRNIISPTALFLAAKVEEQPRKLEHVIKVAHACLHPQELQLDTKSDAYLQQAQELVILETIMLQTLGFEITIEHPHTDVVKCTQLVRGQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN46</accession>
    <name>A0A452IN46_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Crooked neck-like protein 1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="11">Crooked neck homolog</fullName>
        </alternativeName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="9">Involved in pre-mRNA splicing process. As a component of the minor spliceosome, involved in the splicing of U12-type introns in pre-mRNAs.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus speckle</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the crooked-neck family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IN46"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033365.1">
        <property type="protein sequence ID" value="ENSGAGP00000029382.1"/>
        <property type="gene ID" value="ENSGAGG00000021233.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016607">
        <property type="term" value="C:nuclear speck"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071014">
        <property type="term" value="C:post-mRNA release spliceosomal complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071011">
        <property type="term" value="C:precatalytic spliceosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000974">
        <property type="term" value="C:Prp19 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071007">
        <property type="term" value="C:U2-type catalytic step 2 spliceosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000245">
        <property type="term" value="P:spliceosomal complex assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000075">
        <property type="entry name" value="Crooked neck pre-mRNA-splicing factor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000117">
        <property type="entry name" value="Crooked neck pre-mRNA-splicing factor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000269">
        <property type="entry name" value="Crooked neck pre-mRNA-splicing factor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.10">
        <property type="entry name" value="Tetratricopeptide repeat domain"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003107">
        <property type="entry name" value="HAT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055430">
        <property type="entry name" value="HAT_Syf1_CNRKL1_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045075">
        <property type="entry name" value="Syf1-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011990">
        <property type="entry name" value="TPR-like_helical_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11246:SF3">
        <property type="entry name" value="CROOKED NECK-LIKE PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11246">
        <property type="entry name" value="PRE-MRNA SPLICING FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02184">
        <property type="entry name" value="HAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23231">
        <property type="entry name" value="HAT_Syf1_CNRKL1_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00386">
        <property type="entry name" value="HAT"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48452">
        <property type="entry name" value="TPR-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0507">mRNA processing</keyword>
    <keyword evidence="7" id="KW-0508">mRNA splicing</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0747">Spliceosome</keyword>
    <feature type="domain" description="Pre-mRNA-splicing factor Syf1/CRNKL1-like C-terminal HAT-repeats" evidence="14">
        <location>
            <begin position="266"/>
            <end position="416"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="543"/>
            <end position="564"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="12">
        <location>
            <begin position="401"/>
            <end position="471"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="13">
        <location>
            <begin position="548"/>
            <end position="564"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004324"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008644"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022664"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022728"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023187"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00055662"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00067469"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00075217"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="Pfam" id="PF23231"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029382.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="564" mass="68900" checksum="D2C7A191297A271D" modified="2019-05-08" version="1">MEMKNRQVNHARNIWDRAITTLPRVNQFWYKYTYMEEMLGNVAGARQVFERWMEWQPEEQAWHSYINFELRYKEVERARTIYERFVIVHPDVKNWIKYARFEEKHSYFAHARKVYERAVEFFGEEHMNEHLFVAFAKFEENQKEFERVRVIYKYALDRIPKQEAQELFKNYTVFEKKFGDRRGIEDIIVSKRRFQYEEEVKANPHNYDAWFDYLRLVESDADPDTVREVYERAIANVPPIQEKRHWKRYIYLWINYALYEELEAKDPERTRQVYQACVELIPHKKFTFAKMWLLYAQFEIRQKNLPFARRALGTSIGKCPKNKLFKGYIELELQLREFDRCRKLYEKFLEFTPENCTSWIKFAELETILGDTDRARAIYELAIGQPRLDMPEVLWKSYIDFEIEQEEYEKTRNLYRRLLQRTQHVKVWISFAQFELSAGKEESLLRCRQVYEEANKTMRNCEEKEERLMLLESWRNFEEEFGTDCNKERVDKLMPDKVKKRRKLKAEDGSDAGWEEYYDYIFPEDAANQPNLKLLAMAKLWKKQQQENETEEMDPDKDIDESES</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN48</accession>
    <name>A0A452IN48_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Cyclin-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="2">Belongs to the cyclin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IN48"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033258.1">
        <property type="protein sequence ID" value="ENSGAGP00000029281.1"/>
        <property type="gene ID" value="ENSGAGG00000021185.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016538">
        <property type="term" value="F:cyclin-dependent protein serine/threonine kinase regulator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.472.10:FF:000009">
        <property type="entry name" value="cyclin-T2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.472.10">
        <property type="entry name" value="Cyclin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013763">
        <property type="entry name" value="Cyclin-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036915">
        <property type="entry name" value="Cyclin-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043198">
        <property type="entry name" value="Cyclin/Ssn8"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006671">
        <property type="entry name" value="Cyclin_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10026">
        <property type="entry name" value="CYCLIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00134">
        <property type="entry name" value="Cyclin_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00385">
        <property type="entry name" value="CYCLIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47954">
        <property type="entry name" value="Cyclin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="1 2" id="KW-0195">Cyclin</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Cyclin-like" evidence="3">
        <location>
            <begin position="58"/>
            <end position="166"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00023127"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="RuleBase" id="RU000383"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="SMART" id="SM00385"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029281.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="188" mass="20818" checksum="7C2B7E8FA701D28A" modified="2019-05-08" version="1">MAAAASGPGGAGSAARGGGASLSASRWFFSREQLENTPTRRCGVEADKELSYRQQAANLIQDMGQRLNVSQLTINTAIVYMHRFYMHHSFTKFNRNDEGSGLKVNIISPTALFLAAKVEEQPRKLEHVIKVAHACLHPQELQLDTKSDAYLQQAQELVILETIMLQTLGFEITIEHPHTDVVKCTQLV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN74</accession>
    <name>A0A452IN74_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">HECT-type E3 ubiquitin transferase</fullName>
            <ecNumber evidence="4">2.3.2.26</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>S-ubiquitinyl-[E2 ubiquitin-conjugating enzyme]-L-cysteine + [acceptor protein]-L-lysine = [E2 ubiquitin-conjugating enzyme]-L-cysteine + N(6)-ubiquitinyl-[acceptor protein]-L-lysine.</text>
            <dbReference type="EC" id="2.3.2.26"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="3">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="2.3.2.26" evidence="4"/>
    <dbReference type="AlphaFoldDB" id="A0A452IN74"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033529.1">
        <property type="protein sequence ID" value="ENSGAGP00000029529.1"/>
        <property type="gene ID" value="ENSGAGG00000021328.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00143"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019871">
        <property type="term" value="F:sodium channel inhibitor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061630">
        <property type="term" value="F:ubiquitin protein ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007528">
        <property type="term" value="P:neuromuscular junction development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016567">
        <property type="term" value="P:protein ubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032801">
        <property type="term" value="P:receptor catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031623">
        <property type="term" value="P:receptor internalization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048814">
        <property type="term" value="P:regulation of dendrite morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051049">
        <property type="term" value="P:regulation of transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006511">
        <property type="term" value="P:ubiquitin-dependent protein catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd04033">
        <property type="entry name" value="C2_NEDD4_NEDD4L"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00078">
        <property type="entry name" value="HECTc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00201">
        <property type="entry name" value="WW"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1750.10:FF:000026">
        <property type="entry name" value="E3 ubiquitin-protein ligase HACE1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.2160.10:FF:000001">
        <property type="entry name" value="E3 ubiquitin-protein ligase NEDD4-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.2410.10:FF:000001">
        <property type="entry name" value="E3 ubiquitin-protein ligase NEDD4-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1750.10:FF:000001">
        <property type="entry name" value="E3 ubiquitin-protein ligase NEDD4-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.70.10:FF:000006">
        <property type="entry name" value="E3 ubiquitin-protein ligase NEDD4-like protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.70.10:FF:000008">
        <property type="entry name" value="E3 ubiquitin-protein ligase NEDD4-like protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.70.10:FF:000025">
        <property type="entry name" value="E3 ubiquitin-protein ligase NEDD4-like protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000047">
        <property type="entry name" value="Putative E3 ubiquitin-protein ligase NEDD4-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.20.70.10">
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.2160.10">
        <property type="entry name" value="Hect, E3 ligase catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.2410.10">
        <property type="entry name" value="Hect, E3 ligase catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1750.10">
        <property type="entry name" value="Hect, E3 ligase catalytic domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000008">
        <property type="entry name" value="C2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024928">
        <property type="entry name" value="E3_ub_ligase_SMURF1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050409">
        <property type="entry name" value="E3_ubiq-protein_ligase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000569">
        <property type="entry name" value="HECT_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035983">
        <property type="entry name" value="Hect_E3_ubiquitin_ligase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001202">
        <property type="entry name" value="WW_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036020">
        <property type="entry name" value="WW_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11254:SF443">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE NEDD4-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11254">
        <property type="entry name" value="HECT DOMAIN UBIQUITIN-PROTEIN LIGASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00168">
        <property type="entry name" value="C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00632">
        <property type="entry name" value="HECT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00397">
        <property type="entry name" value="WW"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF001569">
        <property type="entry name" value="E3_ub_ligase_SMURF1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00360">
        <property type="entry name" value="C2DOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00239">
        <property type="entry name" value="C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00119">
        <property type="entry name" value="HECTc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00456">
        <property type="entry name" value="WW"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49562">
        <property type="entry name" value="C2 domain (Calcium/lipid-binding domain, CaLB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56204">
        <property type="entry name" value="Hect, E3 ligase catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51045">
        <property type="entry name" value="WW domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50004">
        <property type="entry name" value="C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50237">
        <property type="entry name" value="HECT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01159">
        <property type="entry name" value="WW_DOMAIN_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50020">
        <property type="entry name" value="WW_DOMAIN_2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="7" id="KW-0808">Transferase</keyword>
    <keyword evidence="10" id="KW-0832">Ubl conjugation</keyword>
    <keyword evidence="9 12" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="C2" evidence="14">
        <location>
            <begin position="4"/>
            <end position="126"/>
        </location>
    </feature>
    <feature type="domain" description="WW" evidence="15">
        <location>
            <begin position="193"/>
            <end position="226"/>
        </location>
    </feature>
    <feature type="domain" description="WW" evidence="15">
        <location>
            <begin position="365"/>
            <end position="398"/>
        </location>
    </feature>
    <feature type="domain" description="WW" evidence="15">
        <location>
            <begin position="477"/>
            <end position="510"/>
        </location>
    </feature>
    <feature type="domain" description="WW" evidence="15">
        <location>
            <begin position="528"/>
            <end position="552"/>
        </location>
    </feature>
    <feature type="domain" description="HECT" evidence="16">
        <location>
            <begin position="639"/>
            <end position="973"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="154"/>
            <end position="201"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="244"/>
            <end position="310"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="406"/>
            <end position="475"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="13">
        <location>
            <begin position="164"/>
            <end position="175"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="13">
        <location>
            <begin position="281"/>
            <end position="290"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="13">
        <location>
            <begin position="440"/>
            <end position="451"/>
        </location>
    </feature>
    <feature type="active site" description="Glycyl thioester intermediate" evidence="11 12">
        <location>
            <position position="941"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000885"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00012485"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PIRSR" id="PIRSR001569-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00104"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50020"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50237"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029529.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="974" mass="112325" checksum="161414F67B817A4E" modified="2019-05-08" version="1">MASGAVEPVYGLSEDEDESRLLRVKVVSGIDLAKKDIFGASDPYVKLSLYVADENRELALVQTKTIKKTLNPKWNEEFYFRVNPTNHRLLFEVFDENRLTRDDFLGQVDVPLNHLPTEDPTMERPYTFKDFLLRPRSHKSRVKGFLRLKMAYMPKNGGQEEETTEQREESERAWEVVDSNDSVSHHQEELPPPPLPPGWEEKVDNLGRTYYVNHNNRTTQWHRPSLIDVASDSDNNIRQINQEAAHRRFRSRRHISEDLEPEPSESGDASEPWETISEEASISGDSLSLSLPPPPASPVSRTSPQELSDELNRRLQVTPDSNGEQFGSLIQRDPSSRLRSCSVTDAVAEQSHLSLPSVAYVHTTPGLPSGWEERKDAKGRTYYVNHNNRTTTWTRPIMQLAEDGAVGSAANSNNHLSEPQIRRPRSLSSPTVTLSAPLEFAKDSPVRRAVKDTLSNPQSPQPSPYNSPKPQHKVTQSFLPPGWEMRIAPNGRPFFIDHNTKTTTWEDPRLKFPVHLRSKASLNPNDLGPLPPGWEERIHLDGRTFYIDHRSQVLICGDIDTRDLVPSYDSKITQWEDPRLQNPAITGPAVPYSREFKQKYDYFRKKLKKPADIPNRFEMKLHRNNIFEESYRRIMSVKRSDVLKARLWIEFESEKGLDYGGVAREWFFLLSKEMFNPYYGLFEYSATDNYTLQINPNSGLCNEDHLSYFTFIGRVAGLAVFHGKLLDGFFIRPFYKMMLGKQITLKDMESVDSEYYNSLKWILENDPTELDLMFCIDEENFGQTYQVDLKPNGSEIMVTNENKREYIDLVIQWRFVNRVQKQMNAFLEGFTELLPIDLIKIFDENELELLMCGLGDVDVNDWRQHTIYKNGYCPNHPVIQWYWKAVLLMDAEKRIRLLQFVTGTSRVPMNGFAELYGSNGPQLFTIEQWGSPEKLPRAHTCFNRLDLPPYESFEDLREKLLMAVENAQGFEGVD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGB1</accession>
    <name>A0A452GGB1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Unconventional myosin-XVIIIa</fullName>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">May link Golgi membranes to the cytoskeleton and participate in the tensile force required for vesicle budding from the Golgi. Thereby, may play a role in Golgi membrane trafficking and could indirectly give its flattened shape to the Golgi apparatus. Alternatively, in concert with LURAP1 and CDC42BPA/CDC42BPB, has been involved in modulating lamellar actomyosin retrograde flow that is crucial to cell protrusion and migration. May be involved in the maintenance of the stromal cell architectures required for cell to cell contact. Regulates trafficking, expression, and activation of innate immune receptors on macrophages. Plays a role to suppress inflammatory responsiveness of macrophages via a mechanism that modulates CD14 trafficking. Acts as a receptor of surfactant-associated protein A (SFTPA1/SP-A) and plays an important role in internalization and clearance of SFTPA1-opsonized S.aureus by alveolar macrophages. Strongly enhances natural killer cell cytotoxicity.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Microtubule organizing center</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Golgi apparatus</location>
            <location evidence="2">trans-Golgi network</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="13">Golgi outpost</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 16">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <comment type="caution">
        <text evidence="16">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000000688.1">
        <property type="protein sequence ID" value="ENSGAGP00000000612.1"/>
        <property type="gene ID" value="ENSGAGG00000000355.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005815">
        <property type="term" value="C:microtubule organizing center"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032982">
        <property type="term" value="C:myosin filament"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016460">
        <property type="term" value="C:myosin II complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008094">
        <property type="term" value="F:ATP-dependent activity, acting on DNA"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003774">
        <property type="term" value="F:cytoskeletal motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031032">
        <property type="term" value="P:actomyosin structure organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006259">
        <property type="term" value="P:DNA metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903028">
        <property type="term" value="P:positive regulation of opsonization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043030">
        <property type="term" value="P:regulation of macrophage activation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd01386">
        <property type="entry name" value="MYSc_Myo18"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.1160:FF:000006">
        <property type="entry name" value="Myosin-XVIIIa isoform a"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.820:FF:000004">
        <property type="entry name" value="unconventional myosin-XVIIIa isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000020">
        <property type="entry name" value="unconventional myosin-XVIIIa isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.270.10:FF:000002">
        <property type="entry name" value="unconventional myosin-XVIIIa isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.720:FF:000007">
        <property type="entry name" value="unconventional myosin-XVIIIa isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.530:FF:000011">
        <property type="entry name" value="unconventional myosin-XVIIIa isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.340">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.1590">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.270.10">
        <property type="entry name" value="Myosin, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.1160">
        <property type="entry name" value="Vasodilator-stimulated phosphoprotein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000048">
        <property type="entry name" value="IQ_motif_EF-hand-BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002928">
        <property type="entry name" value="Myosin_tail"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036064">
        <property type="entry name" value="MYSc_Myo18"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020587">
        <property type="entry name" value="RecA_monomer-monomer_interface"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057772">
        <property type="entry name" value="SH3_Myo18a"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615">
        <property type="entry name" value="MYOSIN HEAVY CHAIN, NON-MUSCLE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615:SF13">
        <property type="entry name" value="UNCONVENTIONAL MYOSIN-XVIIIA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01576">
        <property type="entry name" value="Myosin_tail_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24556">
        <property type="entry name" value="SH3_Myosin-XVIIIa"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00015">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90257">
        <property type="entry name" value="Myosin rod fragments"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50163">
        <property type="entry name" value="RECA_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="16" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="7 16" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="9 17" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="12" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="8" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="11 16" id="KW-0505">Motor protein</keyword>
    <keyword evidence="10 16" id="KW-0518">Myosin</keyword>
    <keyword evidence="6 16" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Myosin motor" evidence="20">
        <location>
            <begin position="492"/>
            <end position="1264"/>
        </location>
    </feature>
    <feature type="domain" description="RecA family profile 2" evidence="19">
        <location>
            <begin position="628"/>
            <end position="694"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="132"/>
            <end position="391"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="803"/>
            <end position="823"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="1931"/>
            <end position="1981"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="2041"/>
            <end position="2206"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="2219"/>
            <end position="2272"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="2326"/>
            <end position="2357"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="2383"/>
            <end position="2550"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="17">
        <location>
            <begin position="1328"/>
            <end position="1432"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="17">
        <location>
            <begin position="1468"/>
            <end position="1757"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="18">
        <location>
            <begin position="212"/>
            <end position="225"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="234"/>
            <end position="245"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="275"/>
            <end position="286"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="18">
        <location>
            <begin position="297"/>
            <end position="306"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="307"/>
            <end position="331"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="18">
        <location>
            <begin position="370"/>
            <end position="387"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="2066"/>
            <end position="2075"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="2094"/>
            <end position="2109"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="2133"/>
            <end position="2142"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="2164"/>
            <end position="2174"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="2188"/>
            <end position="2197"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="18">
        <location>
            <begin position="2219"/>
            <end position="2231"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="2233"/>
            <end position="2242"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="2244"/>
            <end position="2263"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="2347"/>
            <end position="2356"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="2404"/>
            <end position="2423"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="18">
        <location>
            <begin position="2436"/>
            <end position="2445"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="18">
        <location>
            <begin position="2457"/>
            <end position="2475"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="18">
        <location>
            <begin position="2477"/>
            <end position="2510"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="2513"/>
            <end position="2536"/>
        </location>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <begin position="585"/>
            <end position="592"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004267"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004601"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00008314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00024182"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00058021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00073449"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000612.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2550" mass="285306" checksum="35117562B04C16EF" modified="2019-05-08" version="1">MCAFKGQAKASGDWGWGLVTGSGAGLCSALHSSAIFRVAVSAQLSLCLRLQPWGTDTWLTYCITLALFARTPCPAACSALSSQALPGPQPQLTVRVAVPVPAPCDIAMAFSSRFAFWEQKVKDEDKVLSVKKTGKEDGASENPNAGKDDPPASRSPEPPECPERVKSPEPLLNGKNQLPSQAPARSMNGLQGSRAGSLAQPEPLGEESQGLSRKRVVRVVRKVVRKVLPGEDSGCTKEQGRDGKSPEPASPAKKEEKAPPRVMSSPPLPASSPKPEPKETGPKDEISVGLRSLMSRGKTKQHRPRLRLAEKQEEEKPPELKTPPVRKEPEKPGSPSPAQEEGPVATPGPRAKQEQTVQPSGAKVEPVQVAPQNAAALGQAKAAAGPGSQLTPVEEAHRRLERIFMAALEPVASPSAQSQAKTEEQITAEEAWYETDKVWLVHKDGFSLASQLRADETSLLPEGKVKVKLDHDGAVLEVDEDDVEKANPPSCDRVEDLANLVYLNESSVLHTVRQRYGGNLIHTYAGPTMLVVNPLSSPSMYSEKVMHMFKGCRREDTSPHIYAVAQAAYRDMLMSRQDQSIVLLGSSGSGKTTCCQHLIQYLATIAGSTGKVFSVEKWQALYTILEAFGNSSTSMNGNATRFSQIVSLDFDQAGQVASASIQTMLLEKLRVTKRPTNEATFNVFYYLLAGTDNALRTELHFNHLAETSVFGIVPLSKPEEKQKAAQQFSKLQAAMKVLGISSDEQKAFWLVLGSIYHLGAAGATKAGRKQFARHEWAQKAAYLLGCSLEELSSSVFKHQPKGSLQRSTSFRQGPEEAGLGDGSGSKLTALECLEGLASGLYSELFTLLISLLNRALKSSQHSLCSMMLVDTPGFQNPELANQSRGASFEELCHNYAQERLQMLFHERTFTQELERYKEESIELALDDIEPSTSGSVATVDQASHQALVRSLARTDEARGLLWLLEEEALQPAGNEDTLLERLFAYYGPQEGDKKGHSPLLRSDKPRHFFLGHSYGTNWVEYDATGWLQYVRQNPASQNAVALLQDSQKKIISSLFMGRAGTAMVLSGSIAGLEGGSQLALRRATSMRKTFTTGVAAVKKKSLCIQIKLQVDALVDTIKKSKIHFVHCLLPKAEGWSGDPKGLAARRLSSTELDLHGEHCEAGLMQLDVPLLRAQVRGSRLLDALRMYRQGYPDHMVFSEFRRRFDVLAPHLTKKHGRNYIVTDEKRAVEELLESLDLEKSSSHMGLSRVFFRAGTLAKLEEQRDKQTSKNIAHFQAACRGFLARQQFKKRKIQELAIRCVQKNIQKNRGVKDWPWWKLFTTVRPLIEVQLTEDQIRGKDEEIQQLKGRLEKVERERNELRLGSDRLESRVAELTAELTDERHTGESASQLLDAETAERLRAEKEMKDLQIKYEALKKQMDSMEMEVMEARLIRAAELNGDLDDDDSGGEWRLKYERAIREVDFTKKRMQQEFEDKLELEQQSKRQQERRLADLHADNEECQRALQQLKKKCQRLTAELQDTKLHLEGQQGRNHELEKKQRRFDSELAQAHEEAQREKLLREKLSREKDMLIAEVFSLKQQLEDKDSDIMSFTQKVEALEAELQDVSSQESKDEASLAKVKKQLRDLEAKAKDQEEELDEQAGTIQLLEQAKLRLEMEMERLRQTHSKEVESRDEEVEEIRQSCQKKLKQMEVQLEEEYEDKQKVLREKRELESKLAAASDQVSQRDLETEKRLRKDLKRTKALLADAQIMLDHLKNNAPSKREIAQLRNQLEESEFTCAAAVKARKSMEVEIEDLHLQIDDISKAKAALEEQLSRLQREKNEVQSRLEEDQEDMNELMKKHKASVAQASRDLAQINDLQSQLEDANKEKQELQEKLQGLQSQLEFLEQSMVDKSLVSRQEAKIRELETRLEFERTQVKRLENLASRLKENMEKLTEERDQRAAAENREKEQNKRLQRQLRDSKEEMGELAKKETEASRKKHELEMDLESLEAANQSLQSDLKLAFKRIGDLQAAIEDEMESDDNEDLINSLQDVVTKYQKRKNKLEGDSDMDSELEDRVDGVKSWLSRNKGSSKAVSDDGSWKSSRPALNSAPKEGKEGREPKEVEERPVSVTSSLSYRKRLNLKDSIGGQGDSETLFSTLSERPASPERSFRKAKRSSAASSQEETGSVSSWRKSLGSEDPDDRGSVISQAFSEASSRARKGLEKRWSVTTADFDQASALSAPASRAASSRHGLEADEADARSTLSFTLSSPGSLRRSTSRLSEPLAGVSGALSPQLSRRAELGLETSTLGSRVDSRLCRSRSRLEELDDVSSVALSDTRSLYSQHSLGRSFSVPPRPRTSASDKMQLEDSDIRPVSHRSYLDPDLEAAISEVLNYKPIKFKRSSLDPDPDLDSEEDKRSVRSARSATRPESSERSFSNLRRSASALDCSRPRSSRGPRSSSSSSEEDSSEEERKKSARKHAKKKAKKKSKKKQLPLDSDSSSDSSSSSYHSSSSVKKGPKKSPASEGEGAAEGRDAEKSSKRLKKEEKQRKKQVDSLMMKYLYRPESD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGB3</accession>
    <name>A0A452GGB3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Unconventional myosin-XVIIIa</fullName>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">May link Golgi membranes to the cytoskeleton and participate in the tensile force required for vesicle budding from the Golgi. Thereby, may play a role in Golgi membrane trafficking and could indirectly give its flattened shape to the Golgi apparatus. Alternatively, in concert with LURAP1 and CDC42BPA/CDC42BPB, has been involved in modulating lamellar actomyosin retrograde flow that is crucial to cell protrusion and migration. May be involved in the maintenance of the stromal cell architectures required for cell to cell contact. Regulates trafficking, expression, and activation of innate immune receptors on macrophages. Plays a role to suppress inflammatory responsiveness of macrophages via a mechanism that modulates CD14 trafficking. Acts as a receptor of surfactant-associated protein A (SFTPA1/SP-A) and plays an important role in internalization and clearance of SFTPA1-opsonized S.aureus by alveolar macrophages. Strongly enhances natural killer cell cytotoxicity.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Microtubule organizing center</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Golgi apparatus</location>
            <location evidence="2">trans-Golgi network</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="13">Golgi outpost</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 16">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <comment type="caution">
        <text evidence="16">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000000660.1">
        <property type="protein sequence ID" value="ENSGAGP00000000588.1"/>
        <property type="gene ID" value="ENSGAGG00000000355.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005815">
        <property type="term" value="C:microtubule organizing center"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032982">
        <property type="term" value="C:myosin filament"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016460">
        <property type="term" value="C:myosin II complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008094">
        <property type="term" value="F:ATP-dependent activity, acting on DNA"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003774">
        <property type="term" value="F:cytoskeletal motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031032">
        <property type="term" value="P:actomyosin structure organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006259">
        <property type="term" value="P:DNA metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903028">
        <property type="term" value="P:positive regulation of opsonization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043030">
        <property type="term" value="P:regulation of macrophage activation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd01386">
        <property type="entry name" value="MYSc_Myo18"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.1160:FF:000006">
        <property type="entry name" value="Myosin-XVIIIa isoform a"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.820:FF:000004">
        <property type="entry name" value="unconventional myosin-XVIIIa isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000020">
        <property type="entry name" value="unconventional myosin-XVIIIa isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.270.10:FF:000002">
        <property type="entry name" value="unconventional myosin-XVIIIa isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.720:FF:000007">
        <property type="entry name" value="unconventional myosin-XVIIIa isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.530:FF:000011">
        <property type="entry name" value="unconventional myosin-XVIIIa isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.340">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.1590">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.270.10">
        <property type="entry name" value="Myosin, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.1160">
        <property type="entry name" value="Vasodilator-stimulated phosphoprotein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000048">
        <property type="entry name" value="IQ_motif_EF-hand-BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002928">
        <property type="entry name" value="Myosin_tail"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036064">
        <property type="entry name" value="MYSc_Myo18"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020587">
        <property type="entry name" value="RecA_monomer-monomer_interface"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057772">
        <property type="entry name" value="SH3_Myo18a"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615">
        <property type="entry name" value="MYOSIN HEAVY CHAIN, NON-MUSCLE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615:SF13">
        <property type="entry name" value="UNCONVENTIONAL MYOSIN-XVIIIA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01576">
        <property type="entry name" value="Myosin_tail_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24556">
        <property type="entry name" value="SH3_Myosin-XVIIIa"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00015">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90257">
        <property type="entry name" value="Myosin rod fragments"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50163">
        <property type="entry name" value="RECA_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="16" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="7 16" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="9 17" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="12" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="8" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="11 16" id="KW-0505">Motor protein</keyword>
    <keyword evidence="10 16" id="KW-0518">Myosin</keyword>
    <keyword evidence="6 16" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Myosin motor" evidence="20">
        <location>
            <begin position="513"/>
            <end position="1285"/>
        </location>
    </feature>
    <feature type="domain" description="RecA family profile 2" evidence="19">
        <location>
            <begin position="649"/>
            <end position="715"/>
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    <sequence length="2571" mass="287231" checksum="B61456FB02896915" modified="2019-05-08" version="1">MCAFKGQAKASGDWGWGLVTGSGAGLCSALHSSAIFRVAVSAQLSLCLRLQPWGTDTWLTYCITLALFARTPCPAACSALSSQALPGPQPQLTVRVAVPVPAPCDIAMAFSSRFAFWEQKVKDEDKVLSVKKTGKEDGASENPNAGKDDPPASRSPEPPECPERVKSPEPLLNGKNQLPSQAPARSMNGLQGSRAGSLAQPEPLGEESQGLSRKRVVRVVRKVVRKVLPGEDSGCTKEQGRDGKSPEPASPAKKEEKAPPRVMSSPPLPASSPKPEPKETGPKDEISVGLRSLMSRGKTKQHRPRLRLAEKQEEEKPPELKTPPVRKEPEKPGSPSPAQEEGPVATPGPRAKQEQTVQPSGAKVEPVQVAPQNAAALGQAKAAAGPGSQLTPVEEAHRRLERIFMAALEPVASPSAQSQGQADDAPAATFGPASAASLPQAKTEEQITAEEAWYETDKVWLVHKDGFSLASQLRADETSLLPEGKVKVKLDHDGAVLEVDEDDVEKANPPSCDRVEDLANLVYLNESSVLHTVRQRYGGNLIHTYAGPTMLVVNPLSSPSMYSEKVMHMFKGCRREDTSPHIYAVAQAAYRDMLMSRQDQSIVLLGSSGSGKTTCCQHLIQYLATIAGSTGKVFSVEKWQALYTILEAFGNSSTSMNGNATRFSQIVSLDFDQAGQVASASIQTMLLEKLRVTKRPTNEATFNVFYYLLAGTDNALRTELHFNHLAETSVFGIVPLSKPEEKQKAAQQFSKLQAAMKVLGISSDEQKAFWLVLGSIYHLGAAGATKAGRKQFARHEWAQKAAYLLGCSLEELSSSVFKHQPKGSLQRSTSFRQGPEEAGLGDGSGSKLTALECLEGLASGLYSELFTLLISLLNRALKSSQHSLCSMMLVDTPGFQNPELANQSRGASFEELCHNYAQERLQMLFHERTFTQELERYKEESIELALDDIEPSTSGSVATVDQASHQALVRSLARTDEARGLLWLLEEEALQPAGNEDTLLERLFAYYGPQEGDKKGHSPLLRSDKPRHFFLGHSYGTNWVEYDATGWLQYVRQNPASQNAVALLQDSQKKIISSLFMGRAGTAMVLSGSIAGLEGGSQLALRRATSMRKTFTTGVAAVKKKSLCIQIKLQVDALVDTIKKSKIHFVHCLLPKAEGWSGDPKGLAARRLSSTELDLHGEHCEAGLMQLDVPLLRAQVRGSRLLDALRMYRQGYPDHMVFSEFRRRFDVLAPHLTKKHGRNYIVTDEKRAVEELLESLDLEKSSSHMGLSRVFFRAGTLAKLEEQRDKQTSKNIAHFQAACRGFLARQQFKKRKIQELAIRCVQKNIQKNRGVKDWPWWKLFTTVRPLIEVQLTEDQIRGKDEEIQQLKGRLEKVERERNELRLGSDRLESRVAELTAELTDERHTGESASQLLDAETAERLRAEKEMKDLQIKYEALKKQMDSMEMEVMEARLIRAAELNGDLDDDDSGGEWRLKYERAIREVDFTKKRMQQEFEDKLELEQQSKRQQERRLADLHADNEECQRALQQLKKKCQRLTAELQDTKLHLEGQQGRNHELEKKQRRFDSELAQAHEEAQREKLLREKLSREKDMLIAEVFSLKQQLEDKDSDIMSFTQKVEALEAELQDVSSQESKDEASLAKVKKQLRDLEAKAKDQEEELDEQAGTIQLLEQAKLRLEMEMERLRQTHSKEVESRDEEVEEIRQSCQKKLKQMEVQLEEEYEDKQKVLREKRELESKLAAASDQVSQRDLETEKRLRKDLKRTKALLADAQIMLDHLKNNAPSKREIAQLRNQLEESEFTCAAAVKARKSMEVEIEDLHLQIDDISKAKAALEEQLSRLQREKNEVQSRLEEDQEDMNELMKKHKASVAQASRDLAQINDLQSQLEDANKEKQELQEKLQGLQSQLEFLEQSMVDKSLVSRQEAKIRELETRLEFERTQVKRLENLASRLKENMEKLTEERDQRAAAENREKEQNKRLQRQLRDSKEEMGELAKKETEASRKKHELEMDLESLEAANQSLQSDLKLAFKRIGDLQAAIEDEMESDDNEDLINSLQDVVTKYQKRKNKLEGDSDMDSELEDRVDGVKSWLSRNKGSSKAVSDDGSWKSSRPALNSAPKEGKEGREPKEVEERPVSVTSSLSYRKRLNLKDSIGGQGDSETLFSTLSERPASPERSFRKAKRSSAASSQEETGSVSSWRKSLGSEDPDDRGSVISQAFSEASSRARKGLEKRWSVTTADFDQASALSAPASRAASSRHGLEADEADARSTLSFTLSSPGSLRRSTSRLSEPLAGVSGALSPQLSRRAELGLETSTLGSRVDSRLCRSRSRLEELDDVSSVALSDTRSLYSQHSLGRSFSVPPRPRTSASDKMQLEDSDIRPVSHRSYLDPDLEAAISEVLNYKPIKFKRSSLDPDPDLDSEEDKRSVRSARSATRPESSERSFSNLRRSASALDCSRPRSSRGPRSSSSSSEEDSSEEERKKSARKHAKKKAKKKSKKKQLPLDSDSSSDSSSSSYHSSSSVKKGPKKSPASEGEGAAEGRDAEKSSKRLKKEEKQRKKQVDSLMMKYLYRPESD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGB6</accession>
    <name>A0A452GGB6_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="4">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
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        <property type="gene ID" value="ENSGAGG00000000476.1"/>
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        </source>
    </evidence>
    <sequence length="1016" mass="113464" checksum="1C844935BCFCDB93" modified="2019-05-08" version="1">MGKDRNFSRHFRIFIPKKHRQRFDEVVSQSLLSKLCRSKSEHHTNRLRRSRSEDHQERLLVSTRASSVPRSHEEASKGLRKTTSLVTSSVGSGAARRTVRVYKGSKSFGFTLRGHAPVWIESILPGSPAEKAALKVGDRILFLNGLDMRNCSHDKVVSMLQGSGAMPTLVVEEGMVHFPFDSESADSPNPSSALTSLQWVAEILPSSIKIQGRTFNQQLERLLTPPERYIICKSLEGFFQHRNIDTLIVDVYPVLDTPVKQVIWQFIYQLLTYEEQEHCQQKIARFLGYKSVAAAAEPEAEEQHRTLVRGASVCRGSIRTQGPEDGGTGAGHSDTGEVSIRLMPGERQAGDGTSLPETPNPKMMSAVYAELENRLIGSFAGKMANAPTHRDSPPVPDQANAAGVRKSGMTISWQSDPLPSQQCFYHPQSSIPSPSSTESNPYVSLDSSPAPSPKHMDYTLSPLSRRKKLFTFSKPPRSRDTDRFLDALSEQLGHRVTIVDDFLTPENDYEEMSFHDDQGSYVTNDMSSASEYISSSDDGSSLTYSTLSDHIPPPPLSPPPPPPQFHDSRPVTLSSEVTRNICSPLSKMLGSHSHPVPPPPPPPPPPPVPCAPPLLQRGLVHRRSESSHMSVKRLRWEQVENSEGTIWGQLGEDSDYDKLSDMVKYLDLELHFGTQKPAISLPEPTLLSESFKKKDVVEILSHKKAYNTSILIAHLKLSHIELRQILMTMESDRLESSHIKQLLLYAPDAEEVQQYQNYQDNPSKLSEPDQFVLQMLSVPEYKTRLRSLYCKTTLQEKAEEIKASYECICKASLELKSSKKLAKILEFVLAMGNYLNNGQPKTNKTTGFKINFLTELNTTKTVDGKSTFLHILAKSLSQHFPELLGFAKDLPTVPLAAKVNQRTLTADLNDLQTTISEIETACHNMPAAEEDKFAIVMNSFLESAHPTIKSLDDLQHKAMEEFSKVLSFFGEDSKVTTSESFFGIFAEFMSKFERALSDVQACESQRSPRMTSPLAW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGB7</accession>
    <name>A0A452GGB7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">TP53-binding protein 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Chromosome</location>
            <location evidence="2">Centromere</location>
            <location evidence="2">Kinetochore</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000000702.1">
        <property type="protein sequence ID" value="ENSGAGP00000000625.1"/>
        <property type="gene ID" value="ENSGAGG00000000496.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000776">
        <property type="term" value="C:kinetochore"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016604">
        <property type="term" value="C:nuclear body"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035861">
        <property type="term" value="C:site of double-strand break"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042393">
        <property type="term" value="F:histone binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140005">
        <property type="term" value="F:histone H4K20me2 reader activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000077">
        <property type="term" value="P:DNA damage checkpoint signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006303">
        <property type="term" value="P:double-strand break repair via nonhomologous end joining"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:2000042">
        <property type="term" value="P:negative regulation of double-strand break repair via homologous recombination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045830">
        <property type="term" value="P:positive regulation of isotype switching"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045944">
        <property type="term" value="P:positive regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd17745">
        <property type="entry name" value="BRCT_p53bp1_rpt1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd17724">
        <property type="entry name" value="BRCT_p53bp1_rpt2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd20383">
        <property type="entry name" value="Tudor_53BP1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.140:FF:000021">
        <property type="entry name" value="Tumor suppressor p53-binding protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.30:FF:000019">
        <property type="entry name" value="Tumor suppressor p53-binding protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10190:FF:000003">
        <property type="entry name" value="Tumor suppressor p53-binding protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10190:FF:000005">
        <property type="entry name" value="Tumor suppressor p53-binding protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.10190">
        <property type="entry name" value="BRCT domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015125">
        <property type="entry name" value="53-BP1_Tudor"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001357">
        <property type="entry name" value="BRCT_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036420">
        <property type="entry name" value="BRCT_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047249">
        <property type="entry name" value="BRCT_p53bp1-like_rpt1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047250">
        <property type="entry name" value="BRCT_p53bp1-like_rpt2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014722">
        <property type="entry name" value="Rib_uL2_dom2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047252">
        <property type="entry name" value="TP53BP1-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15321:SF3">
        <property type="entry name" value="TP53-BINDING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15321">
        <property type="entry name" value="TUMOR SUPPRESSOR P53-BINDING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09038">
        <property type="entry name" value="53-BP1_Tudor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18428">
        <property type="entry name" value="BRCT_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00292">
        <property type="entry name" value="BRCT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52113">
        <property type="entry name" value="BRCT domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF63748">
        <property type="entry name" value="Tudor/PWWP/MBT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50172">
        <property type="entry name" value="BRCT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="13" id="KW-0010">Activator</keyword>
    <keyword evidence="17" id="KW-0137">Centromere</keyword>
    <keyword evidence="3" id="KW-0158">Chromosome</keyword>
    <keyword evidence="8" id="KW-0227">DNA damage</keyword>
    <keyword evidence="15" id="KW-0234">DNA repair</keyword>
    <keyword evidence="12" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="5" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="9" id="KW-0995">Kinetochore</keyword>
    <keyword evidence="4" id="KW-0488">Methylation</keyword>
    <keyword evidence="16" id="KW-0539">Nucleus</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="14" id="KW-0804">Transcription</keyword>
    <keyword evidence="11" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="10" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="BRCT" evidence="20">
        <location>
            <begin position="1736"/>
            <end position="1804"/>
        </location>
    </feature>
    <feature type="domain" description="BRCT" evidence="20">
        <location>
            <begin position="1820"/>
            <end position="1920"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="62"/>
            <end position="97"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="156"/>
            <end position="179"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="628"/>
            <end position="657"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="731"/>
            <end position="763"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="776"/>
            <end position="822"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1018"/>
            <end position="1127"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1155"/>
            <end position="1209"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1252"/>
            <end position="1322"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1336"/>
            <end position="1417"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1589"/>
            <end position="1676"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="72"/>
            <end position="82"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="86"/>
            <end position="97"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="156"/>
            <end position="167"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="636"/>
            <end position="651"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="747"/>
            <end position="757"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="790"/>
            <end position="800"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="1075"/>
            <end position="1117"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="19">
        <location>
            <begin position="1265"/>
            <end position="1310"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="19">
        <location>
            <begin position="1596"/>
            <end position="1608"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="1663"/>
            <end position="1676"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004629"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022454"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022838"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023328"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00073180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50172"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000625.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1928" mass="206561" checksum="BF68810D6D18D217" modified="2019-05-08" version="1">MDPSQSQLDLGFSQQDTPCLIVEDSQPESAVTEEDTAHGCLSLLALPLPNCQSPVLEIMTSPQGSKLAAGGGEDRGLEEKPLTEPMDSSTHLDTTGSLSQVIDRLPQPTKTNRVPGMVAAAAEEAGEDASHCATPLLCAEDSGTSQLGFGALELSQSQDVEGDSTLSEEGRGHQLQPADTVPTCCNPMKNDPNEDLHIVRSEAVSTGESYTDVQCQQEKSGRSEVMSSSPRILQQHKEISRIQMSVCNVLESELPAEKRPDPCLQESEILSTQEDMFSQSHTTASESDSTITRVEVGGSVACTPAGTLHLLHLSGQGSLGQESLSTYSSGLITPSSGAFGSHPLISGSPTEQEEAKDEQMNISAPAEGGDLLQEKQRGDEPVEVGNPPVPPGPHVLPQVSTPVSQNAPAFVPGSFPIPSQPEFSHDIFISTQSLEEKPSGEEKGGALLGSCLPADVSETTDALRKISTGDPELSCKLVLSASECSQPTRTEGNTSSLKTNQGSENIQVEMDSTLQASELKPCSTESYSMQLRLSEDSQALLSENCEKPKREAVWATGEPTICPVAEGVAAEGSVAQMACIDHSCDSGSQTAATLSVQSEISSHTLGQEALTEIKEKLSAEASEFEAVPETLCEEQGGERQAEEKQLDEKGSSDPLTLSQGLILLRETKEHEGQEVMEVESSEAFGKNSEKLSDLRHGLVQMDLETRKKETPKDSNVVAEVKSMKDLSLKAAPEDVAKLSEVPPQPSVGESLQQQDDSWPQVKVDVAPGLASCAKMCPGSQGNKSQAEVDLELKQVQKDRQPPPAPLERGDEVPATPQESQQPHLMTLEKEVDMPEKPFKTSNLEQEEERAAEQSPCPSTGTLFHFALPKEGDVIQPLTSVTPSLIGQLKMGPRRHSTPIVDGSCPDRTIVTSDVTAEGSIGTIDVTVESAMASADVLEECEKGDSGAAPEADGKLCLRMNLATPVSEESEGSLPFSLEKPAASERKNGSSAVAGAVASSQKTLSVFSLVCEVQREDEARGHDLPASPFRGNLFSFPSTQEEEEEQEQQHDNPEAWQQHQKHIPCGEIKVSQPAEQDSKHQPSHAKEGEAMEADIEKEGKSAQEKLSKALAAHERDKGQQLWAGTSNKEVQTTGGSVLAPAAVSAATQTMARVSSQVEVGTSMAGQRPGQQDAKVQTEESGEKLANAPGDDTESLHSQGEEEFNLPQPPRGRIQRRHMRTIREVRTVVTRIITDVYYVDGAEVERKVIEETEEPIVDCHESETDISPSRTTGGSSLTSGDLGDISSLSSKVSSLHRTSSGASSGLSATHGGSSSGRGAGSFKGKVSGMEAGEFALPSGRGVLGKLSPRRGAGQPGSPLRAGQTGATACEEDEDAALGSRQGGKAPLTPRGRGRRGRPPSRATGTRDAAGSGLLGVEDISTTVSPEEKPFVRIVVAGPSDSADLSSGSSFVGLRVVAKWSSNAYFYSGTITQDVGAGKYKLLFDDGYECDVLGKDILLCDPIPLETEVTALSEDEYFSAGVVKGHRKESGELYYCIEKEGQRKWYKRMAVILSLEQGNKLREQFGLSPYEPITPLTKAADISLDNLVEGKRKRRSNFGSPGTSSSSTTPTRKGLESPRVAPGLLSGKRKLIASEDERSPAKRGRKSAAMKPGAVRAGEFVSPCESGDHTGEPSTLEEHHGPLPHNKTLFLGYAFLLTMATSSDKLAIHQKPSDGPAGSSEEEEEFSIPYNKEYTALQLRTGAGYILEDFNETQCNAAYQCLLIADQHCRTRKYFLCLASGIPCVSHVWVRDSCHANQLQNYRNYLLPAGYSLQEQRLLEWHPRENPFGNLKVLLVSNQQQNFLELWSEILMTGGAASVKQHYSNAQNKDIALGVFDVVVSDSSCPAAVLKCAEALRLPVVSQEWVIQSLIAGERVGYNKHPKYKHDYGPC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGB9</accession>
    <name>A0A452GGB9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">tRNA dimethylallyltransferase</fullName>
            <ecNumber evidence="4">2.5.1.75</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="17">Isopentenyl-diphosphate:tRNA isopentenyltransferase</fullName>
        </alternativeName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="19">Catalyzes the transfer of a dimethylallyl group onto the adenine at position 37 of both cytosolic and mitochondrial tRNAs, leading to the formation of N6-(dimethylallyl)adenosine (i6A37). Mediates modification of a limited subset of tRNAs: tRNA(Ser)(AGA), tRNA(Ser)(CGA), tRNA(Ser)(UGA), as well as partial modification of the selenocysteine tRNA(Ser)(UCA). TRIT1 is therefore required for selenoprotein expression.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>adenosine(37) in tRNA + dimethylallyl diphosphate = N(6)-dimethylallyladenosine(37) in tRNA + diphosphate</text>
            <dbReference type="Rhea" id="RHEA:26482"/>
            <dbReference type="Rhea" id="RHEA-COMP:10162"/>
            <dbReference type="Rhea" id="RHEA-COMP:10375"/>
            <dbReference type="ChEBI" id="CHEBI:33019"/>
            <dbReference type="ChEBI" id="CHEBI:57623"/>
            <dbReference type="ChEBI" id="CHEBI:74411"/>
            <dbReference type="ChEBI" id="CHEBI:74415"/>
            <dbReference type="EC" id="2.5.1.75"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Mitochondrion</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 20">Belongs to the IPP transferase family.</text>
    </comment>
    <dbReference type="EC" id="2.5.1.75" evidence="4"/>
    <dbReference type="AlphaFoldDB" id="A0A452GGB9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000703.1">
        <property type="protein sequence ID" value="ENSGAGP00000000626.1"/>
        <property type="gene ID" value="ENSGAGG00000000522.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0052381">
        <property type="term" value="F:tRNA dimethylallyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006400">
        <property type="term" value="P:tRNA modification"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.20.140:FF:000002">
        <property type="entry name" value="tRNA dimethylallyltransferase, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.20.140">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039657">
        <property type="entry name" value="Dimethylallyltransferase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018022">
        <property type="entry name" value="IPT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022755">
        <property type="entry name" value="Znf_C2H2_jaz"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00174">
        <property type="entry name" value="miaA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11088">
        <property type="entry name" value="TRNA DIMETHYLALLYLTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11088:SF89">
        <property type="entry name" value="TRNA DIMETHYLALLYLTRANSFERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01715">
        <property type="entry name" value="IPPT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12171">
        <property type="entry name" value="zf-C2H2_jaz"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="14 20" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="10" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="16" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="11 20" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8 20" id="KW-0808">Transferase</keyword>
    <keyword evidence="15" id="KW-0809">Transit peptide</keyword>
    <keyword evidence="9" id="KW-0819">tRNA processing</keyword>
    <keyword evidence="13" id="KW-0862">Zinc</keyword>
    <keyword evidence="12" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="Zinc finger double-stranded RNA binding" evidence="22">
        <location>
            <begin position="315"/>
            <end position="339"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="21">
        <location>
            <begin position="360"/>
            <end position="380"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004173"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00012665"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00017477"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022694"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00030893"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00049563"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00057810"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="RuleBase" id="RU003785"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="Pfam" id="PF12171"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000626.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="380" mass="43548" checksum="2A311E2CBB47D313" modified="2019-05-08" version="1">MISFVDPLVSNYTVVDFRNKAAALIEDIFAQEKIPIVVGGTNYYIESLLWKVLVDTKEKTSPSPGRAADRRVELEQLDGCELHCRLSQVDPEMAAKLHPHDKRKVARSLQVFEETGIPHSEFLHRQREEEGGGPLGGPLKYPNSCILWLYADQAALDERLDKRVDDMVAAGLLEELRDFHRRYNQEKVAENCQDYQHGIFQSIGFKEFHEYLTTEGKCPEETSTLLLEKGIQALKLVTKRYARKQNKWVRNRFLRRPGPNVPPVYGLEVSDLSQWDENVLKPALQIVESFIQGRQPLADPVKMESEINEDKRSHHVCELCNRVIIGDREWAAHTKSKSHLHHLKKRWKLHSKLSAVESQVNVTETQGVSEGTENLGQDRN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGC5</accession>
    <name>A0A452GGC5_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GGC5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000000708.1">
        <property type="protein sequence ID" value="ENSGAGP00000000631.1"/>
        <property type="gene ID" value="ENSGAGG00000000484.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015840">
        <property type="term" value="P:urea transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006833">
        <property type="term" value="P:water transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009952">
        <property type="entry name" value="Uroplakin-2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024831">
        <property type="entry name" value="Uroplakin-3"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15446">
        <property type="entry name" value="UROPLAKIN III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15446:SF17">
        <property type="entry name" value="UROPLAKIN-3A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07353">
        <property type="entry name" value="Uroplakin_II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="1" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="1">
        <location>
            <begin position="204"/>
            <end position="228"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000631.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="256" mass="27725" checksum="231839A41ECF61DA" modified="2019-05-08" version="1">DLPLSSLLALTVPRAQNLRPQIANPKLATNNPTLTTIALEKPFCVFDDSLSPGSSYVVYLYAMADSASTVSSAVTDNSSKPLNTTFQDTNGGQLGPYRAALFNVPNCVSPPLLADVVNVKKVSDVLKQYLFRVGDDVTCLYDPNFPGACNPPLAEDTTYRFKYLLVDLVAGVVKDQTLWSDPMKTLKQTSTIDTWPGRRSGGMIVITSILSTLIFLLVVGFLASIFFFTSTETTHASQTTQQAVPRSQDSSEGVNQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGD1</accession>
    <name>A0A452GGD1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Unconventional myosin-XVIIIa</fullName>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">May link Golgi membranes to the cytoskeleton and participate in the tensile force required for vesicle budding from the Golgi. Thereby, may play a role in Golgi membrane trafficking and could indirectly give its flattened shape to the Golgi apparatus. Alternatively, in concert with LURAP1 and CDC42BPA/CDC42BPB, has been involved in modulating lamellar actomyosin retrograde flow that is crucial to cell protrusion and migration. May be involved in the maintenance of the stromal cell architectures required for cell to cell contact. Regulates trafficking, expression, and activation of innate immune receptors on macrophages. Plays a role to suppress inflammatory responsiveness of macrophages via a mechanism that modulates CD14 trafficking. Acts as a receptor of surfactant-associated protein A (SFTPA1/SP-A) and plays an important role in internalization and clearance of SFTPA1-opsonized S.aureus by alveolar macrophages. Strongly enhances natural killer cell cytotoxicity.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Microtubule organizing center</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Golgi apparatus</location>
            <location evidence="2">trans-Golgi network</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="13">Golgi outpost</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 16">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <comment type="caution">
        <text evidence="16">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000000723.1">
        <property type="protein sequence ID" value="ENSGAGP00000000644.1"/>
        <property type="gene ID" value="ENSGAGG00000000355.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005815">
        <property type="term" value="C:microtubule organizing center"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
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        <property type="term" value="C:myosin filament"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016460">
        <property type="term" value="C:myosin II complex"/>
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        <property type="project" value="TreeGrafter"/>
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        <property type="term" value="F:actin filament binding"/>
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        <property type="term" value="F:ATP binding"/>
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        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
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        <property type="term" value="F:ATP-dependent activity, acting on DNA"/>
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        <property type="project" value="InterPro"/>
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        <property type="term" value="F:cytoskeletal motor activity"/>
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        <property type="project" value="UniProtKB-UniRule"/>
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        <property type="term" value="P:actomyosin structure organization"/>
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        <property type="project" value="TreeGrafter"/>
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        <property type="term" value="P:DNA metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
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    <dbReference type="GO" id="GO:1903028">
        <property type="term" value="P:positive regulation of opsonization"/>
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        <property type="project" value="UniProtKB-ARBA"/>
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    <dbReference type="GO" id="GO:0043030">
        <property type="term" value="P:regulation of macrophage activation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd01386">
        <property type="entry name" value="MYSc_Myo18"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Myosin-XVIIIa isoform a"/>
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    </dbReference>
    <dbReference type="FunFam" id="1.10.10.820:FF:000004">
        <property type="entry name" value="unconventional myosin-XVIIIa isoform X1"/>
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    <dbReference type="FunFam" id="3.40.850.10:FF:000020">
        <property type="entry name" value="unconventional myosin-XVIIIa isoform X1"/>
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    <dbReference type="FunFam" id="4.10.270.10:FF:000002">
        <property type="entry name" value="unconventional myosin-XVIIIa isoform X1"/>
        <property type="match status" value="1"/>
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    <dbReference type="FunFam" id="1.20.120.720:FF:000007">
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    <dbReference type="FunFam" id="1.20.58.530:FF:000011">
        <property type="entry name" value="unconventional myosin-XVIIIa isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.340">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.1590">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
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    <dbReference type="Gene3D" id="4.10.270.10">
        <property type="entry name" value="Myosin, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.1160">
        <property type="entry name" value="Vasodilator-stimulated phosphoprotein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000048">
        <property type="entry name" value="IQ_motif_EF-hand-BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
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        <property type="entry name" value="Myosin_tail"/>
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    <dbReference type="InterPro" id="IPR036064">
        <property type="entry name" value="MYSc_Myo18"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020587">
        <property type="entry name" value="RecA_monomer-monomer_interface"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057772">
        <property type="entry name" value="SH3_Myo18a"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615">
        <property type="entry name" value="MYOSIN HEAVY CHAIN, NON-MUSCLE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615:SF13">
        <property type="entry name" value="UNCONVENTIONAL MYOSIN-XVIIIA"/>
        <property type="match status" value="1"/>
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    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01576">
        <property type="entry name" value="Myosin_tail_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24556">
        <property type="entry name" value="SH3_Myosin-XVIIIa"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00015">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90257">
        <property type="entry name" value="Myosin rod fragments"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
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    </dbReference>
    <dbReference type="PROSITE" id="PS50163">
        <property type="entry name" value="RECA_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="16" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="7 16" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="9 17" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="12" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="8" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="11 16" id="KW-0505">Motor protein</keyword>
    <keyword evidence="10 16" id="KW-0518">Myosin</keyword>
    <keyword evidence="6 16" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
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        <location>
            <begin position="487"/>
            <end position="1259"/>
        </location>
    </feature>
    <feature type="domain" description="RecA family profile 2" evidence="19">
        <location>
            <begin position="623"/>
            <end position="689"/>
        </location>
    </feature>
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            <begin position="132"/>
            <end position="418"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
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        </location>
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            <begin position="1926"/>
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        </location>
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            <begin position="2036"/>
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        </location>
    </feature>
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        <location>
            <begin position="2214"/>
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        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="2321"/>
            <end position="2352"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="2378"/>
            <end position="2545"/>
        </location>
    </feature>
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        <location>
            <begin position="1323"/>
            <end position="1427"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="17">
        <location>
            <begin position="1463"/>
            <end position="1752"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="18">
        <location>
            <begin position="212"/>
            <end position="225"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="234"/>
            <end position="245"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="275"/>
            <end position="286"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="18">
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            <begin position="297"/>
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        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
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            <begin position="307"/>
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        </location>
    </feature>
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        <location>
            <begin position="399"/>
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        </location>
    </feature>
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            <begin position="2061"/>
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    </feature>
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            <begin position="2239"/>
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        </location>
    </feature>
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        <location>
            <begin position="2342"/>
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        </location>
    </feature>
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            <begin position="2399"/>
            <end position="2418"/>
        </location>
    </feature>
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        <location>
            <begin position="2431"/>
            <end position="2440"/>
        </location>
    </feature>
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        <location>
            <begin position="2452"/>
            <end position="2470"/>
        </location>
    </feature>
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        <location>
            <begin position="2472"/>
            <end position="2505"/>
        </location>
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    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="18">
        <location>
            <begin position="2508"/>
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        </location>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <begin position="580"/>
            <end position="587"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
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            <dbReference type="ARBA" id="ARBA00004267"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00004601"/>
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    </evidence>
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    </evidence>
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    </evidence>
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            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
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        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
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        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00024182"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00058021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00073449"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000644.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2545" mass="284470" checksum="703ADCAD1C5180D1" modified="2019-05-08" version="1">MCAFKGQAKASGDWGWGLVTGSGAGLCSALHSSAIFRVAVSAQLSLCLRLQPWGTDTWLTYCITLALFARTPCPAACSALSSQALPGPQPQLTVRVAVPVPAPCDIAMAFSSRFAFWEQKVKDEDKVLSVKKTGKEDGASENPNAGKDDPPASRSPEPPECPERVKSPEPLLNGKNQLPSQAPARSMNGLQGSRAGSLAQPEPLGEESQGLSRKRVVRVVRKVVRKVLPGEDSGCTKEQGRDGKSPEPASPAKKEEKAPPRVMSSPPLPASSPKPEPKETGPKDEISVGLRSLMSRGKTKQHRPRLRLAEKQEEEKPPELKTPPVRKEPEKPGSPSPAQEEGPVATPGPRAKQEQTVQPSGAKVEPVQVAPQNAAALGQAKLEPVASPSAQSQGQADDAPAATFGPASAASLPQAKTEEQITAEEAWYETDKVWLVHKDGFSLASQLRADETSLLPEGKVKVKLDHDGAVLEVDEDDVEKANPPSCDRVEDLANLVYLNESSVLHTVRQRYGGNLIHTYAGPTMLVVNPLSSPSMYSEKVMHMFKGCRREDTSPHIYAVAQAAYRDMLMSRQDQSIVLLGSSGSGKTTCCQHLIQYLATIAGSTGKVFSVEKWQALYTILEAFGNSSTSMNGNATRFSQIVSLDFDQAGQVASASIQTMLLEKLRVTKRPTNEATFNVFYYLLAGTDNALRTELHFNHLAETSVFGIVPLSKPEEKQKAAQQFSKLQAAMKVLGISSDEQKAFWLVLGSIYHLGAAGATKAGRKQFARHEWAQKAAYLLGCSLEELSSSVFKHQPKGSLQRSTSFRQGPEEAGLGDGSGSKLTALECLEGLASGLYSELFTLLISLLNRALKSSQHSLCSMMLVDTPGFQNPELANQSRGASFEELCHNYAQERLQMLFHERTFTQELERYKEESIELALDDIEPSTSGSVATVDQASHQALVRSLARTDEARGLLWLLEEEALQPAGNEDTLLERLFAYYGPQEGDKKGHSPLLRSDKPRHFFLGHSYGTNWVEYDATGWLQYVRQNPASQNAVALLQDSQKKIISSLFMGRAGTAMVLSGSIAGLEGGSQLALRRATSMRKTFTTGVAAVKKKSLCIQIKLQVDALVDTIKKSKIHFVHCLLPKAEGWSGDPKGLAARRLSSTELDLHGEHCEAGLMQLDVPLLRAQVRGSRLLDALRMYRQGYPDHMVFSEFRRRFDVLAPHLTKKHGRNYIVTDEKRAVEELLESLDLEKSSSHMGLSRVFFRAGTLAKLEEQRDKQTSKNIAHFQAACRGFLARQQFKKRKIQELAIRCVQKNIQKNRGVKDWPWWKLFTTVRPLIEVQLTEDQIRGKDEEIQQLKGRLEKVERERNELRLGSDRLESRVAELTAELTDERHTGESASQLLDAETAERLRAEKEMKDLQIKYEALKKQMDSMEMEVMEARLIRAAELNGDLDDDDSGGEWRLKYERAIREVDFTKKRMQQEFEDKLELEQQSKRQQERRLADLHADNEECQRALQQLKKKCQRLTAELQDTKLHLEGQQGRNHELEKKQRRFDSELAQAHEEAQREKLLREKLSREKDMLIAEVFSLKQQLEDKDSDIMSFTQKVEALEAELQDVSSQESKDEASLAKVKKQLRDLEAKAKDQEEELDEQAGTIQLLEQAKLRLEMEMERLRQTHSKEVESRDEEVEEIRQSCQKKLKQMEVQLEEEYEDKQKVLREKRELESKLAAASDQVSQRDLETEKRLRKDLKRTKALLADAQIMLDHLKNNAPSKREIAQLRNQLEESEFTCAAAVKARKSMEVEIEDLHLQIDDISKAKAALEEQLSRLQREKNEVQSRLEEDQEDMNELMKKHKASVAQASRDLAQINDLQSQLEDANKEKQELQEKLQGLQSQLEFLEQSMVDKSLVSRQEAKIRELETRLEFERTQVKRLENLASRLKENMEKLTEERDQRAAAENREKEQNKRLQRQLRDSKEEMGELAKKETEASRKKHELEMDLESLEAANQSLQSDLKLAFKRIGDLQAAIEDEMESDDNEDLINSLQDVVTKYQKRKNKLEGDSDMDSELEDRVDGVKSWLSRNKGSSKAVSDDGSWKSSRPALNSAPKEGKEGREPKEVEERPVSVTSSLSYRKRLNLKDSIGGQGDSETLFSTLSERPASPERSFRKAKRSSAASSQEETGSVSSWRKSLGSEDPDDRGSVISQAFSEASSRARKGLEKRWSVTTADFDQASALSAPASRAASSRHGLEADEADARSTLSFTLSSPGSLRRSTSRLSEPLAGVSGALSPQLSRRAELGLETSTLGSRVDSRLCRSRSRLEELDDVSSVALSDTRSLYSQHSLGRSFSVPPRPRTSASDKMQLEDSDIRPVSHRSYLDPDLEAAISEVLNYKPIKFKRSSLDPDPDLDSEEDKRSVRSARSATRPESSERSFSNLRRSASALDCSRPRSSRGPRSSSSSSEEDSSEEERKKSARKHAKKKAKKKSKKKQLPLDSDSSSDSSSSSYHSSSSVKKGPKKSPASEGEGAAEGRDAEKSSKRLKKEEKQRKKQVDSLMMKYLYRPESD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GGH1</accession>
    <name>A0A452GGH1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Clathrin light chain</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
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        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
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        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
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                <consortium name="Ensembl"/>
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        <scope>IDENTIFICATION</scope>
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        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
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        <scope>IDENTIFICATION</scope>
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        <text evidence="6">Clathrin is the major protein of the polyhedral coat of coated pits and vesicles.</text>
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    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 6">Cytoplasmic vesicle membrane</location>
            <topology evidence="1 6">Peripheral membrane protein</topology>
            <orientation evidence="1 6">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="6">Membrane</location>
            <location evidence="6">Coated pit</location>
            <topology evidence="6">Peripheral membrane protein</topology>
            <orientation evidence="6">Cytoplasmic side</orientation>
        </subcellularLocation>
        <text evidence="6">Cytoplasmic face of coated pits and vesicles.</text>
    </comment>
    <comment type="similarity">
        <text evidence="2 6">Belongs to the clathrin light chain family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GGH1"/>
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        <property type="protein sequence ID" value="ENSGAGP00000000670.1"/>
        <property type="gene ID" value="ENSGAGG00000000556.1"/>
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        <property type="component" value="Unassembled WGS sequence"/>
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            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
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                <person name="Cornelius J.A."/>
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                <person name="Henen B.T."/>
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                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
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        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="3017"/>
            <end position="3026"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="1">
        <location>
            <begin position="3044"/>
            <end position="3055"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF23415"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="Pfam" id="PF25281"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000000673.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="3173" mass="352928" checksum="DD2F768E8A0ED9F3" modified="2019-05-08" version="1">MCDSSAHKLLLLCGQSSDQEGDLILQTGTFNYQKFAEIIEDPEVAQLLSNTDPDNKASLTVSCLGEGDWSNLGHPQFQQLINLKLNPDPVLPEMDGVSEFAEYVSETVDVPTPFDLLEPPTSGGFLKLSKPCCYIFPGGRGDSALFAVNGFNILVDGGSERKSCFWKLVRHLDRIDSILLTHIGADNLPGINGLLQRKIAEQDEEQSQGSTNYSDWMKNLISPELGVVFFNVPEKLKMPESSMKVKRSIEEACLTLQYLNKLGIKSEPLYRVLSNTIEPITLFHKMGVGKLDMYILNPVKDSKEMQFLMQKWAGNSKAKTGIILANGKEGEISVPYLTSITALVVWLPASPTEKIVRVLFPGNAPQNKILEGLEKLKHLDLLRYPVATQKDMSSGLPPPSLKQTKIKQRTDSKESLKSSPKPVATKQARKEEASDDGAKEIKPEIVKEARTEKKVKEHTEKIPEKPAKPEKIKTESSDALKAEKRKLTKDKIGKKHLKEKVSKLDEKKDKEKKEIKKERKDFKKDDGKKEEKKDSKKEAKPELKKMSKPDLKPFTPEVRKTLYKAKVPGRIKTEKTRVKAEKEAPVEQKMSPAQKRPVQQEPGMADMAEQSLVLSSPEDLTKDFEELNYEEKVELQVAQEMTDIEASDAVMRGPETGGKRVPEMTAQGHVDEALILEENIMMKVTPLDSPDEGIITFDRERELPCEEKPVQLEVRAGQAETFEDERAALEEPLEIGEFAEKAERKEEAKLLPEVHKDERGQAPTHTEAAAREAQTPELKEKEEEQKISLDRKQREGEMTCEKYIEGMDTGEEGEKEKREDVIEKAELEETEEQYTEEKDMKTKEFYQLHQDEEKGTKFTAKEKKREFSEEISEKSKLLEKEVTKEEYSLRSAPVPPGATAEHISYIQDETIPGYSETEQTISDEEIHDEQEERIPHLKYDVSAYDISVPDHPGSFEAVHGIQATSITDIAAKGYISQESEILVYPTSIVAAPLAEEEHVSSATSVTECDKLSSFATSVAEDQSVASITAPQTEETGKSSLLLDTVNSIPSSRTEATQGLDYVPSAGTISPTSSLEEDKCFKSPPSEDFQALAEGERKFKAQKKGLQEETEEEEEEEEEDETPNIEMPEKLRGHYGAQLFTQQGHLADVLIESRTEASMLLLSTEETQLSQVDSCSGEAEERCISPDDSTVKMASPTQSGPTSAGHTPFHQSPVEEKLETVEAELFENDRDASVKKGEGQTSGVEKGAESESEPLKKAEAPAANDIPEEELPSVIQEKPGLANKVLLPMESMFPRNEKQELPVGKEHISYHKQKTIEISEKDDSKLYIPSTDIISKPEKQAEFKETEMMQDVSLEFAKYTAKEPVSTKGISPKNAEVPQQSQLESDKSSQIFIKEISPDDSKSLSFGEESLCKEKSSDISIREASPEEEKPLYFTGHTLEKEKSTHIFIKDISPEDTKSISLTEESPGKETSPDIFMKGTSPEDTKSLTFTEKSPAKETFLATSTNEFSPEAVIPLHFMQQSPATSEKSQETAAKDIIKEKPKSPLCPEHSPIKDMLPTEISSAEISPEDTRVFSLTEMSLARGKALEMCSQETFPDMKFLHFTESRRIQDKESSDVCSEDVKSCDFMEETQTKEKSPEMEDIYMKNLNYEKKVWFPEEMEEIRSKESQQKYDKERESTFLDEVQEYEKKSSLTVTEEEKIIHSMAYEGYALETVEKKTSDSEYGYFQEEDAPRQAGYLPKKEDYLCYGEDKPVPQETEAGHFKVDSAEAQEFARWEDKGLSLETNKMEISPTEITSSEKSEVVRATPSSQHVCIEEKQVKPSSAAKGREESLFPSKQDYSYLDHKEKAVLDLYAKSLDRALTVSPLPASSSALEKEVDTSTQELEKHFMTDQSKPVLQAEVLQGATVEISDADALNQETSDSKYMLDIEQNAESKSTEAPDDPRGSKEQSAAAFGLLYSTSPAEYEYVAASQKDSDAASKNSMHSFSGPYYPKSSSDIIGHSEDMGSFSQVEYHSESCTRTSSSPSEKEVCASKGDDHAKQLEQEEREPTPYPDERTFQYADIYDQVVVSGVGLPSLLLKDVDSLGHPDKEKLKPTFVASRTEKVSVSQKQLLYHISAKEEESCLYTSAEKELSSPTSPKDKVDSSFAYTGIQERYSPLAESDKHGMPSDLRFKEMQEGQNGLSSSTAAPKEPYLTLPPSSPGSRECTPPTYTIDAFYQEERTPYIEAAHPDENIISISQEQRSLSPMEPQPKSFYYEKAGRGEESTSDSELEKGAKEKSEKESKLPSPPQAPEKVSEEEVYADISVMEMAQVSELLHQDKHQESHLSTSIASSPCDPVQKEELGVEKTETATVIETGTAGYSTLTDTCRDVWASEMSYSSHISLQQGKDAEKSPKEAHFLVTDYHHAEIASKGYDKDKVIYLKSTAKYVKEEHQEGLALQSKEMDLSSSGFEFSSLEKHEPSIVFRHETSSSGHLGDKSLILKFSDALLSDQGKKDEYLEVSKKDTSVDSSSLSFSPLSPYKEDKLFSGAVEREVASQQRAKEDCPRLSEDASPEATTPVIHTTAESFYSHMLSAYTGAEPDTRSLWDVSPLIPADSVKTYPAETEGFVSAENCGSAFTCDVDDTLPCRIECQSTLMSPLSTEKQKQSCAFSPEQHLSVTETLMAMTSLPDVLTSFPPHQQEAAATANGPTEVSTSPQAFRSVQSAAEVKDEKALPDSDWTVSPSLGGEDMARQSRPSSLRSPEHIFQPFFPEAQRGGRTEDHGDASHEGEPCLGASSDNRQKFSSCEYKQRKGELSPSFINPSPHDFSEDSDLSQDEGHPSVKGRPHHTTGSDVQATMIEETPPTSVSDSGTSQSDSDVPPETEECPSITADAAMDSDEDADFLPVDKAVGTSHHGSTRSSHDPQPVPMIDPHPHPPHPDVCMVDPEMLVNEQNVSRTDKISKKDLKEKSKGVRKPLSKPKSSSPARKLDVKGKRSPTPVKQPLDRSPKSTTAKKERDGGQKQSKPRTQLVQPSHIEEKDELSRSSHSNPGKTLVNGIKTNPVSSSPKSSLSVLPGPPVYVDLAYIPNHCSGKNSDLEFFKRVRASYYVVSGNDPVNGEPSRVVLDALLEGKSQWGENLQVTLIPTHDTEVTREWYQQTHEKQQELNIMVLASSSTVVMQDESFPACKIEF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXC4</accession>
    <name>A0A452GXC4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">ATP binding cassette subfamily C member 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>17beta-estradiol 17-O-(beta-D-glucuronate)(in) + ATP + H2O = 17beta-estradiol 17-O-(beta-D-glucuronate)(out) + ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:60128"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:82961"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="15">
            <dbReference type="Rhea" id="RHEA:60129"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>ATP + H2O + xenobioticSide 1 = ADP + phosphate + xenobioticSide 2.</text>
            <dbReference type="EC" id="7.6.2.2"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="14">
            <text>leukotriene C4(in) + ATP + H2O = leukotriene C4(out) + ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:38963"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:57973"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="14">
            <dbReference type="Rhea" id="RHEA:38964"/>
        </physiologicalReaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the ABC transporter superfamily. ABCC family. Conjugate transporter (TC 3.A.1.208) subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GXC4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007883.1">
        <property type="protein sequence ID" value="ENSGAGP00000006791.1"/>
        <property type="gene ID" value="ENSGAGG00000004641.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016324">
        <property type="term" value="C:apical plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008559">
        <property type="term" value="F:ABC-type xenobiotic transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006869">
        <property type="term" value="P:lipid transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd18595">
        <property type="entry name" value="ABC_6TM_MRP1_2_3_6_D1_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18603">
        <property type="entry name" value="ABC_6TM_MRP1_2_3_6_D2_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd03250">
        <property type="entry name" value="ABCC_MRP_domain1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd03244">
        <property type="entry name" value="ABCC_MRP_domain2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000293">
        <property type="entry name" value="ATP binding cassette subfamily C member 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1560.10:FF:000001">
        <property type="entry name" value="ATP-binding cassette subfamily C member 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1560.10:FF:000007">
        <property type="entry name" value="ATP-binding cassette subfamily C member 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000074">
        <property type="entry name" value="Multidrug resistance-associated protein 5 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1560.10">
        <property type="entry name" value="ABC transporter type 1, transmembrane domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003593">
        <property type="entry name" value="AAA+_ATPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011527">
        <property type="entry name" value="ABC1_TM_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036640">
        <property type="entry name" value="ABC1_TM_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003439">
        <property type="entry name" value="ABC_transporter-like_ATP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017871">
        <property type="entry name" value="ABC_transporter-like_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050173">
        <property type="entry name" value="ABC_transporter_C-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24223">
        <property type="entry name" value="ATP-BINDING CASSETTE SUB-FAMILY C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24223:SF176">
        <property type="entry name" value="ATP-BINDING CASSETTE SUB-FAMILY C MEMBER 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00664">
        <property type="entry name" value="ABC_membrane"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00005">
        <property type="entry name" value="ABC_tran"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00382">
        <property type="entry name" value="AAA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90123">
        <property type="entry name" value="ABC transporter transmembrane region"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50929">
        <property type="entry name" value="ABC_TM1F"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00211">
        <property type="entry name" value="ABC_TRANSPORTER_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50893">
        <property type="entry name" value="ABC_TRANSPORTER_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11" id="KW-0445">Lipid transport</keyword>
    <keyword evidence="12 16" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="9" id="KW-1278">Translocase</keyword>
    <keyword evidence="5 16" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="10 16" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="100"/>
            <end position="120"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="179"/>
            <end position="202"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="208"/>
            <end position="227"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="289"/>
            <end position="313"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="333"/>
            <end position="353"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="662"/>
            <end position="685"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="716"/>
            <end position="737"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="793"/>
            <end position="810"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="816"/>
            <end position="836"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="901"/>
            <end position="922"/>
        </location>
    </feature>
    <feature type="domain" description="ABC transmembrane type-1" evidence="18">
        <location>
            <begin position="88"/>
            <end position="350"/>
        </location>
    </feature>
    <feature type="domain" description="ABC transporter" evidence="17">
        <location>
            <begin position="353"/>
            <end position="588"/>
        </location>
    </feature>
    <feature type="domain" description="ABC transmembrane type-1" evidence="18">
        <location>
            <begin position="672"/>
            <end position="958"/>
        </location>
    </feature>
    <feature type="domain" description="ABC transporter" evidence="17">
        <location>
            <begin position="994"/>
            <end position="1228"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009726"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022967"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00034018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00047523"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00047576"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50893"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50929"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006791.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="21">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1239" mass="137290" checksum="FF5CAA558745746F" modified="2019-05-08" version="1">MVLKGYRKPLEMEDLWDLKENMRTQQILSRFDRNMRAEVRKARRELERRRCKRSLCEAAPDHRNGLGKAQSQDILVLVTPSLPPGLDLGLLISFVADPTAYAWQGYLYALLLFVTALVQSLCLQQYFQLCFVLGMSLRTALMAAVYKKALTVSNATRKEYTQGETVNLMSADTQRFMDLATFIHQLWSAPLQIALSIIFLWWELGPSVLAGIGVMVLLIPINAVLATKARRIQMKNMKNKDQRMKIMNEILSGIKILKLFAWEPSFERQVGEIRVRELKGLLHFTNLQAFSIFIFSCAPFLVSVASFAVYVMVDEKNVLDAQKAFTSISLFNILRFPMATLPLVISSLVQTAVSTERLERYLGSGNLDPSAIKCDPSLSRPSRVTLDIKPGHLVAVVGAVGSGKSSLVSAVLGEMENLKGHINIQGSVAYVPQQAWIQNATLKDNILFGSDLDEARYQQVVEACALLPDLKLLPGGDLTEIGEKGINLSGGQKQRVSLARAVYSDADIYLLDDPLSAVDAHVGKHLFDQVLGPDGLLQGKTRILVTHSVSFLPRTDEIVVLVAGAVSEQGSYSTLLANGGAFAQLLSTYGSQEGSTPEGEATGMWAGAEPRQEAGLGPVEMFLSGVACRRSCVRGACGGWTPWDAATECAALQVKFSLYLRYLHAVGWGFSAWVFITYVAQYAAFMGSNLWLSDWTDDSLRYWNQTYPTSQRDMRIGVFGALGTVQAAFLLAGALLATRGAIRASRVLHQQLLSNILRVPMSFFDTTPTGRIVNRFAKVRDIFTVDETIPMSFRSWLTCFLGIISTLLMICLATPYFALIIIPLSILYFFLLRYYVSTSRQLRRLDSVTRSPIYSHFGETVSGLSVIRAYGHQERFLQHNDRIVDMNQKSVYSWIVSNRWLAVRLEFVGNLVVFSAALLAVITRDNLGSGIVGLSISSALNVSPTLNWLVRMTSELETNIVAVERVREYTEVENEAQWVTAQRPPPGWPSKGEIQITDYQVRYRPGLELVLHGLTCDIGSTEKVGVVGRTGAGKSSLTNCLFRILEAAGGKIVIDGVDISTIGLHDLRQSLTIIPQDPVLFSGTLRMNLDPFDQHSDEEIWKALELAHLKTYVLDLPERLSHQVSEAGENLSVGQRQLLCLARALLRKSKILILDEATAAVDLETDHLIQTTIRSEFANCTVLTIAHRLHTIMDSNRVMVLQAGRIVEYDSPERLLQQQGLFSTMAKDAGIVSAQATVL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXC8</accession>
    <name>A0A452GXC8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">Lysyl oxidase homolog</fullName>
            <ecNumber evidence="21">1.4.3.13</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="25 26">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="26" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="25" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="25" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="21">Mediates the post-translational oxidative deamination of lysine residues on target proteins leading to the formation of deaminated lysine (allysine).</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="18 21">
            <text>L-lysyl-[protein] + O2 + H2O = (S)-2-amino-6-oxohexanoyl-[protein] + H2O2 + NH4(+)</text>
            <dbReference type="Rhea" id="RHEA:24544"/>
            <dbReference type="Rhea" id="RHEA-COMP:9752"/>
            <dbReference type="Rhea" id="RHEA-COMP:12448"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15379"/>
            <dbReference type="ChEBI" id="CHEBI:16240"/>
            <dbReference type="ChEBI" id="CHEBI:28938"/>
            <dbReference type="ChEBI" id="CHEBI:29969"/>
            <dbReference type="ChEBI" id="CHEBI:131803"/>
            <dbReference type="EC" id="1.4.3.13"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>N(6)-acetyl-L-lysyl-[protein] + O2 + H2O = acetamide + (S)-2-amino-6-oxohexanoyl-[protein] + H2O2</text>
            <dbReference type="Rhea" id="RHEA:51648"/>
            <dbReference type="Rhea" id="RHEA-COMP:10731"/>
            <dbReference type="Rhea" id="RHEA-COMP:12448"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15379"/>
            <dbReference type="ChEBI" id="CHEBI:16240"/>
            <dbReference type="ChEBI" id="CHEBI:27856"/>
            <dbReference type="ChEBI" id="CHEBI:61930"/>
            <dbReference type="ChEBI" id="CHEBI:131803"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 21">
            <name>Cu cation</name>
            <dbReference type="ChEBI" id="CHEBI:23378"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="21">
            <name>lysine tyrosylquinone residue</name>
            <dbReference type="ChEBI" id="CHEBI:20489"/>
        </cofactor>
        <text evidence="21">Contains 1 lysine tyrosylquinone.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="4">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3 21">Secreted</location>
            <location evidence="3 21">Extracellular space</location>
        </subcellularLocation>
    </comment>
    <comment type="PTM">
        <text evidence="21">The lysine tyrosylquinone cross-link (LTQ) is generated by condensation of the epsilon-amino group of a lysine with a topaquinone produced by oxidation of tyrosine.</text>
    </comment>
    <comment type="similarity">
        <text evidence="5 21">Belongs to the lysyl oxidase family.</text>
    </comment>
    <comment type="caution">
        <text evidence="20">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="EC" id="1.4.3.13" evidence="21"/>
    <dbReference type="AlphaFoldDB" id="A0A452GXC8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007889.1">
        <property type="protein sequence ID" value="ENSGAGP00000006796.1"/>
        <property type="gene ID" value="ENSGAGG00000005478.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005507">
        <property type="term" value="F:copper ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004720">
        <property type="term" value="F:protein-lysine 6-oxidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048513">
        <property type="term" value="P:animal organ development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030199">
        <property type="term" value="P:collagen fibril organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006954">
        <property type="term" value="P:inflammatory response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:2000329">
        <property type="term" value="P:negative regulation of T-helper 17 cell lineage commitment"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009888">
        <property type="term" value="P:tissue development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000001">
        <property type="entry name" value="Lysyl oxidase 4 isoform X1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000008">
        <property type="entry name" value="Lysyl oxidase homolog 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000022">
        <property type="entry name" value="lysyl oxidase homolog 3 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.250.10">
        <property type="entry name" value="SRCR-like domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050912">
        <property type="entry name" value="LOX-like_protein"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001695">
        <property type="entry name" value="Lysyl_oxidase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019828">
        <property type="entry name" value="Lysyl_oxidase_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001190">
        <property type="entry name" value="SRCR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036772">
        <property type="entry name" value="SRCR-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45817:SF2">
        <property type="entry name" value="LYSYL OXIDASE HOMOLOG 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45817">
        <property type="entry name" value="LYSYL OXIDASE-LIKE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01186">
        <property type="entry name" value="Lysyl_oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00530">
        <property type="entry name" value="SRCR"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00074">
        <property type="entry name" value="LYSYLOXIDASE"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00258">
        <property type="entry name" value="SPERACTRCPTR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00202">
        <property type="entry name" value="SR"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56487">
        <property type="entry name" value="SRCR-like"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00926">
        <property type="entry name" value="LYSYL_OXIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00420">
        <property type="entry name" value="SRCR_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50287">
        <property type="entry name" value="SRCR_2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="14 21" id="KW-0186">Copper</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="15 20" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="16" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6 21" id="KW-0886">LTQ</keyword>
    <keyword evidence="9 21" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="17" id="KW-0539">Nucleus</keyword>
    <keyword evidence="13 21" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="26" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="11" id="KW-0677">Repeat</keyword>
    <keyword evidence="8 21" id="KW-0964">Secreted</keyword>
    <keyword evidence="10 23" id="KW-0732">Signal</keyword>
    <keyword evidence="12 21" id="KW-0801">TPQ</keyword>
    <feature type="signal peptide" evidence="23">
        <location>
            <begin position="1"/>
            <end position="26"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019265829" description="Lysyl oxidase homolog" evidence="23">
        <location>
            <begin position="27"/>
            <end position="756"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="24">
        <location>
            <begin position="44"/>
            <end position="145"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="24">
        <location>
            <begin position="170"/>
            <end position="281"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="24">
        <location>
            <begin position="308"/>
            <end position="408"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="24">
        <location>
            <begin position="418"/>
            <end position="528"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="288"/>
            <end position="314"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="22">
        <location>
            <begin position="298"/>
            <end position="313"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="70"/>
            <end position="134"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="83"/>
            <end position="144"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="114"/>
            <end position="124"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="247"/>
            <end position="257"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="333"/>
            <end position="397"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="346"/>
            <end position="407"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="377"/>
            <end position="387"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="495"/>
            <end position="505"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001935"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00007492"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022477"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022772"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023008"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00047861"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00052308"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00196"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="RuleBase" id="RU367046"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS50287"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006796.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="756" mass="84516" checksum="5754AA3E46BD2774" modified="2019-05-08" version="1">MRSSSAWAWQELLALLCSVWLWVCAAQHTGPTPTAQSQGPKLKFRLAGYPRKHNEGRIEVFYNEEWGTICDDDFTLANAHVLCRHLGFVAATGWAHSAKYGKGIGRVWLDNVNCGGSEKSIADCKSRGWGNSDCSHEEDAGVICKDEHIPGYVDSNVIEAEQNQVEELRLRPVVSGARKRLPVTEGVVEVRYKEGWAQICDEGWNPKNNRVVCGMMGFPAEKKVNRNFYKLYLERQQLSYRLHSVACTGNEVHISMCPFEFYKGNATAACKGGMPAVVSCMPGPLFTTGSSPKKKQQQRQQQQGQQRIRLKGGAKAGEGRVEVLKNSEWGTVCDDRWNLLTASVVCRELGFGSAKEALTGARMGQGMGPIHMNEVQCTGLEMSLWSCPYKNITAEDCKHSEDAGVRCNIPYMGYETTIRLSGGRSRFEGRVEVALGTGSSGGPAWGLICGDGWGTLEAMVACRQLGLGFANHGLQETWYWDASNVTEMVLSGVRCAGHEMALSHCQHHGNSLNCKRTDTRFAAGVICSETASDLLLHAPLVQETAYIEDRPLHMLYCAAEENCLSSTARNANWPYGHRRLLRFSSQIHNNGRADFRPKAGRHSWVWHECHRHYHSMDIFTHYDLLTPNGTKVAEGHKASFCLEDTECQEDVAKRYECANFGEQGITVGCWDLYRHDIDCQWIDITDVRPGNYILQVVINPNYEVAESDFTNNAMKCNCKYDGHRIWVHNCHIGDAFSEEANKRFEQYPGQLNNQIV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXC9</accession>
    <name>A0A452GXC9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">SOCS box domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="pathway">
        <text evidence="1">Protein modification; protein ubiquitination.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GXC9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007761.1">
        <property type="protein sequence ID" value="ENSGAGP00000006683.1"/>
        <property type="gene ID" value="ENSGAGG00000005388.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00143"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070531">
        <property type="term" value="C:BRCA1-A complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031436">
        <property type="term" value="C:BRCA1-BARD1 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004842">
        <property type="term" value="F:ubiquitin-protein transferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035556">
        <property type="term" value="P:intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0085020">
        <property type="term" value="P:protein K6-linked ubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.750.20:FF:000001">
        <property type="entry name" value="Ankyrin repeat and SOCS box containing 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000246">
        <property type="entry name" value="Ankyrin repeat and SOCS box containing 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.750.20">
        <property type="entry name" value="SOCS box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001496">
        <property type="entry name" value="SOCS_box"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036036">
        <property type="entry name" value="SOCS_box-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003903">
        <property type="entry name" value="UIM_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24171">
        <property type="entry name" value="ANKYRIN REPEAT DOMAIN-CONTAINING PROTEIN 39-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24171:SF8">
        <property type="entry name" value="BRCA1-ASSOCIATED RING DOMAIN PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00023">
        <property type="entry name" value="Ank"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07525">
        <property type="entry name" value="SOCS_box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01415">
        <property type="entry name" value="ANKYRIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="SMART" id="SM00253">
        <property type="entry name" value="SOCS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00969">
        <property type="entry name" value="SOCS_box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF158235">
        <property type="entry name" value="SOCS box-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50225">
        <property type="entry name" value="SOCS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50330">
        <property type="entry name" value="UIM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3 4" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <feature type="repeat" description="ANK" evidence="4">
        <location>
            <begin position="197"/>
            <end position="229"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="4">
        <location>
            <begin position="230"/>
            <end position="262"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="4">
        <location>
            <begin position="263"/>
            <end position="295"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="4">
        <location>
            <begin position="296"/>
            <end position="328"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="4">
        <location>
            <begin position="328"/>
            <end position="360"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="4">
        <location>
            <begin position="400"/>
            <end position="432"/>
        </location>
    </feature>
    <feature type="domain" description="SOCS box" evidence="6">
        <location>
            <begin position="552"/>
            <end position="594"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="42"/>
            <end position="65"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="46"/>
            <end position="58"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50225"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006683.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="594" mass="66147" checksum="124CFC58A3F164E5" modified="2019-05-08" version="1">MATKITATTNTQRKTIGHEEYSLYSSMSEDELIQLAIEQSLKEDPSGQSAAQNHQKSVMTAPGEPPTLTRPCSHNPPHQINAWHRLAAQSGGQAPGSSWGVRRRYDGSLFITPQPVELDPVVEAIKEGDESALLNMMKSGKNLSEPNKDGWLPLHEAAYYGQAGCLKVLQRACERKNVEAVRILVQYNADANHRCNRGWTALHEAVSRNDIEIIDILIKGGAKVESKNTYGITSLFVAAEGGQLEALRYLAKCGADINTQASDSASALYEACKNGHEKVVEFLLSQGADANKTNKDGLLPLHIASKKGNYEIVKMLLPVTSRTRVKRSGISPLHLAAERNNDDILEELIEAGYDVNTTLSFERARLYEDRRSTVLYFAVINNNIYATELLLQAGANPNIDLINPLLISIRHGCLKTMKLLLDHGANIDAYISSHPTTFPATIMFSMKYLSLLKFLMDLGCDADSCFKCQYGNGPHPVLDYRRDRLDANPVNKQPNVVQFCDMVSAPEMNRWAGPIIDVLLDYVGNVQLCARLKEHIDSYEDWAVVKEKAELPRPLAHLCRIKVRNVIGRNRIKLIDTLPLPGRLIRYLQYDYSQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXD8</accession>
    <name>A0A452GXD8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">Lysyl oxidase homolog</fullName>
            <ecNumber evidence="21">1.4.3.13</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="25 26">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="26" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="25" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="25" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="21">Mediates the post-translational oxidative deamination of lysine residues on target proteins leading to the formation of deaminated lysine (allysine).</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="18 21">
            <text>L-lysyl-[protein] + O2 + H2O = (S)-2-amino-6-oxohexanoyl-[protein] + H2O2 + NH4(+)</text>
            <dbReference type="Rhea" id="RHEA:24544"/>
            <dbReference type="Rhea" id="RHEA-COMP:9752"/>
            <dbReference type="Rhea" id="RHEA-COMP:12448"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15379"/>
            <dbReference type="ChEBI" id="CHEBI:16240"/>
            <dbReference type="ChEBI" id="CHEBI:28938"/>
            <dbReference type="ChEBI" id="CHEBI:29969"/>
            <dbReference type="ChEBI" id="CHEBI:131803"/>
            <dbReference type="EC" id="1.4.3.13"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>N(6)-acetyl-L-lysyl-[protein] + O2 + H2O = acetamide + (S)-2-amino-6-oxohexanoyl-[protein] + H2O2</text>
            <dbReference type="Rhea" id="RHEA:51648"/>
            <dbReference type="Rhea" id="RHEA-COMP:10731"/>
            <dbReference type="Rhea" id="RHEA-COMP:12448"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15379"/>
            <dbReference type="ChEBI" id="CHEBI:16240"/>
            <dbReference type="ChEBI" id="CHEBI:27856"/>
            <dbReference type="ChEBI" id="CHEBI:61930"/>
            <dbReference type="ChEBI" id="CHEBI:131803"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 21">
            <name>Cu cation</name>
            <dbReference type="ChEBI" id="CHEBI:23378"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="21">
            <name>lysine tyrosylquinone residue</name>
            <dbReference type="ChEBI" id="CHEBI:20489"/>
        </cofactor>
        <text evidence="21">Contains 1 lysine tyrosylquinone.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="4">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3 21">Secreted</location>
            <location evidence="3 21">Extracellular space</location>
        </subcellularLocation>
    </comment>
    <comment type="PTM">
        <text evidence="21">The lysine tyrosylquinone cross-link (LTQ) is generated by condensation of the epsilon-amino group of a lysine with a topaquinone produced by oxidation of tyrosine.</text>
    </comment>
    <comment type="similarity">
        <text evidence="5 21">Belongs to the lysyl oxidase family.</text>
    </comment>
    <comment type="caution">
        <text evidence="20">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="EC" id="1.4.3.13" evidence="21"/>
    <dbReference type="AlphaFoldDB" id="A0A452GXD8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007893.1">
        <property type="protein sequence ID" value="ENSGAGP00000006799.1"/>
        <property type="gene ID" value="ENSGAGG00000005478.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005507">
        <property type="term" value="F:copper ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004720">
        <property type="term" value="F:protein-lysine 6-oxidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048513">
        <property type="term" value="P:animal organ development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030199">
        <property type="term" value="P:collagen fibril organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006954">
        <property type="term" value="P:inflammatory response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:2000329">
        <property type="term" value="P:negative regulation of T-helper 17 cell lineage commitment"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009888">
        <property type="term" value="P:tissue development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000001">
        <property type="entry name" value="Lysyl oxidase 4 isoform X1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000008">
        <property type="entry name" value="Lysyl oxidase homolog 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.250.10:FF:000022">
        <property type="entry name" value="lysyl oxidase homolog 3 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.250.10">
        <property type="entry name" value="SRCR-like domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050912">
        <property type="entry name" value="LOX-like_protein"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001695">
        <property type="entry name" value="Lysyl_oxidase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019828">
        <property type="entry name" value="Lysyl_oxidase_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001190">
        <property type="entry name" value="SRCR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036772">
        <property type="entry name" value="SRCR-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45817:SF2">
        <property type="entry name" value="LYSYL OXIDASE HOMOLOG 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45817">
        <property type="entry name" value="LYSYL OXIDASE-LIKE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01186">
        <property type="entry name" value="Lysyl_oxidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00530">
        <property type="entry name" value="SRCR"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00074">
        <property type="entry name" value="LYSYLOXIDASE"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00258">
        <property type="entry name" value="SPERACTRCPTR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00202">
        <property type="entry name" value="SR"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56487">
        <property type="entry name" value="SRCR-like"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00926">
        <property type="entry name" value="LYSYL_OXIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00420">
        <property type="entry name" value="SRCR_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50287">
        <property type="entry name" value="SRCR_2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="14 21" id="KW-0186">Copper</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="15 20" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="16" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6 21" id="KW-0886">LTQ</keyword>
    <keyword evidence="9 21" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="17" id="KW-0539">Nucleus</keyword>
    <keyword evidence="13 21" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="26" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="11" id="KW-0677">Repeat</keyword>
    <keyword evidence="8 21" id="KW-0964">Secreted</keyword>
    <keyword evidence="10 23" id="KW-0732">Signal</keyword>
    <keyword evidence="12 21" id="KW-0801">TPQ</keyword>
    <feature type="signal peptide" evidence="23">
        <location>
            <begin position="1"/>
            <end position="26"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019359250" description="Lysyl oxidase homolog" evidence="23">
        <location>
            <begin position="27"/>
            <end position="750"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="24">
        <location>
            <begin position="44"/>
            <end position="145"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="24">
        <location>
            <begin position="170"/>
            <end position="281"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="24">
        <location>
            <begin position="308"/>
            <end position="408"/>
        </location>
    </feature>
    <feature type="domain" description="SRCR" evidence="24">
        <location>
            <begin position="418"/>
            <end position="522"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="288"/>
            <end position="314"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="22">
        <location>
            <begin position="298"/>
            <end position="313"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="70"/>
            <end position="134"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="83"/>
            <end position="144"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="114"/>
            <end position="124"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="247"/>
            <end position="257"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="333"/>
            <end position="397"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="346"/>
            <end position="407"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="377"/>
            <end position="387"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="489"/>
            <end position="499"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001935"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00007492"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022477"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022772"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023008"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00047861"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00052308"/>
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    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00196"/>
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    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="RuleBase" id="RU367046"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS50287"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006799.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="750" mass="84191" checksum="A505759FF585C364" modified="2019-05-08" version="1">MRSSSAWAWQELLALLCSVWLWVCAAQHTGPTPTAQSQGPKLKFRLAGYPRKHNEGRIEVFYNEEWGTICDDDFTLANAHVLCRHLGFVAATGWAHSAKYGKGIGRVWLDNVNCGGSEKSIADCKSRGWGNSDCSHEEDAGVICKDEHIPGYVDSNVIEAEQNQVEELRLRPVVSGARKRLPVTEGVVEVRYKEGWAQICDEGWNPKNNRVVCGMMGFPAEKKVNRNFYKLYLERQQLSYRLHSVACTGNEVHISMCPFEFYKGNATAACKGGMPAVVSCMPGPLFTTGSSPKKKQQQRQQQQGQQRIRLKGGAKAGEGRVEVLKNSEWGTVCDDRWNLLTASVVCRELGFGSAKEALTGARMGQGMGPIHMNEVQCTGLEMSLWSCPYKNITAEDCKHSEDAGVRCNIPYMGYETTIRIVGGRTPTEGRVEVKRGSKWGTVCSEGWTTKEAMVACRQLGLGYSLHAATETWYWDASNVTEMVLSGVRCAGHEMALSHCQHHGNSLNCKRTDTRFAAGVICSETASDLLLHAPLVQETAYIEDRPLHMLYCAAEENCLSSTARNANWPYGHRRLLRFSSQIHNNGRADFRPKAGRHSWVWHECHRHYHSMDIFTHYDLLTPNGTKVAEGHKASFCLEDTECQEDVAKRYECANFGEQGITVGCWDLYRHDIDCQWIDITDVRPGNYILQVVINPNYEVAESDFTNNAMKCNCKYDGHRIWVHNCHIGDAFSEEANKRFEQYPGQLNNQIV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXD9</accession>
    <name>A0A452GXD9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">C2H2-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">May be involved in transcriptional regulation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GXD9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007901.1">
        <property type="protein sequence ID" value="ENSGAGP00000006806.1"/>
        <property type="gene ID" value="ENSGAGG00000005488.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000111">
        <property type="entry name" value="GLI family zinc finger 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000478">
        <property type="entry name" value="Zinc finger protein 133"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000631">
        <property type="entry name" value="zinc finger protein 189 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000295">
        <property type="entry name" value="zinc finger protein 19"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000358">
        <property type="entry name" value="zinc finger protein 24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000269">
        <property type="entry name" value="Zinc finger protein 287"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000016">
        <property type="entry name" value="zinc finger protein 37 homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001119">
        <property type="entry name" value="zinc finger protein 408"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000277">
        <property type="entry name" value="Zinc finger protein 48"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001270">
        <property type="entry name" value="zinc finger protein 583 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001437">
        <property type="entry name" value="Zinc finger protein 594"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000609">
        <property type="entry name" value="zinc finger protein 621"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000495">
        <property type="entry name" value="zinc finger protein 668"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000009">
        <property type="entry name" value="zinc finger protein 79 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000642">
        <property type="entry name" value="Zinc finger with KRAB and SCAN domains 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="17"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050752">
        <property type="entry name" value="C2H2-ZF_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24384">
        <property type="entry name" value="FINGER PUTATIVE TRANSCRIPTION FACTOR FAMILY-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24384:SF242">
        <property type="entry name" value="ZINC FINGER PROTEIN 628"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13465">
        <property type="entry name" value="zf-H2C2_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="17"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="17"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="17"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="11" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="4" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="5" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13" id="KW-0539">Nucleus</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="12" id="KW-0804">Transcription</keyword>
    <keyword evidence="10" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="9" id="KW-0832">Ubl conjugation</keyword>
    <keyword evidence="8" id="KW-0862">Zinc</keyword>
    <keyword evidence="7 14" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="16">
        <location>
            <begin position="239"/>
            <end position="266"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="16">
        <location>
            <begin position="267"/>
            <end position="294"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="16">
        <location>
            <begin position="295"/>
            <end position="322"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="16">
        <location>
            <begin position="323"/>
            <end position="350"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="16">
        <location>
            <begin position="351"/>
            <end position="378"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="16">
        <location>
            <begin position="379"/>
            <end position="406"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="16">
        <location>
            <begin position="407"/>
            <end position="434"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="16">
        <location>
            <begin position="435"/>
            <end position="462"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="16">
        <location>
            <begin position="463"/>
            <end position="491"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="16">
        <location>
            <begin position="492"/>
            <end position="519"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="16">
        <location>
            <begin position="519"/>
            <end position="546"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="16">
        <location>
            <begin position="547"/>
            <end position="574"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="16">
        <location>
            <begin position="575"/>
            <end position="602"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="16">
        <location>
            <begin position="603"/>
            <end position="630"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="16">
        <location>
            <begin position="631"/>
            <end position="658"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="16">
        <location>
            <begin position="659"/>
            <end position="686"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="16">
        <location>
            <begin position="687"/>
            <end position="714"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="1"/>
            <end position="91"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="175"/>
            <end position="225"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00003767"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006806.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="720" mass="79992" checksum="3934BC521F860E65" modified="2019-05-08" version="1">MRSLDAPAQCTQTRDSGAGRSRPSSIPDTWVPSRAPGPSVRRGRPCLQPRAGGIACERGAGDGEAAEAGNRCPEEGGETPPSPRRGSAGGSVQLGSACRSLFSWQVSLPDPTPSTSLLSPSLSLKAQLGCWGNPSPSSPLREGFLGGCFSHSCFLLSDKPLVILSHPDPWLDSLSLPAGARTEEGNSKQEGPAGAKLHGTSHCPKQGGDACGGQQQGAPLGQRPNDLTQLASIPAEKPYKCHECNKSYSLSSNLSRHRRSHTGERSHKCTQCGKSFHFNSQLVQHQISHTGERPYKCPDCGKSFGVSSNLLQHQRIHWVERPYSCANCEKSFGHSSLLAQHQRRHMDERPYKCGECGKAFGVSSALIRHQRIHTGERPYKCLVCGKSFNLSSNLFTHRRIHWDEKPYKCPECGRSFSQSSSLSRHQISHTGERPYKCPECGKCFNQSSLLIRHQRIHTGERPYRCSQCWKSFLRSSQLKRHQRMVHTGERPYKCPDCGKSFNQKSNLKQHQRIHTGEPYKCPQCGKCFRVSSHLIRHQVTHTGERPYKCSICGRSFSLSSHLIRHRIVHTEDKPYVCSECGKKFSLSSRLDRHLRTHTGERPYKCPNCGSGFVDSSALIKHQRIHTGERPYTCPECGKSFGRSSYLITHQRTHTGERPYKCPDCGKSFVGSSDLNKHQRTHTGERPYNCPECGKSFSRSSNLIAHQRIHARSFTPAQGCG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXE4</accession>
    <name>A0A452GXE4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Thrombospondin type-1 domain-containing protein 7A</fullName>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cell projection</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000007899.1">
        <property type="protein sequence ID" value="ENSGAGP00000006804.1"/>
        <property type="gene ID" value="ENSGAGG00000005480.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042995">
        <property type="term" value="C:cell projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030036">
        <property type="term" value="P:actin cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001525">
        <property type="term" value="P:angiogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000014">
        <property type="entry name" value="Thrombospondin type 1 domain containing 7A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000017">
        <property type="entry name" value="Thrombospondin type 1 domain containing 7A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000018">
        <property type="entry name" value="Thrombospondin type 1 domain containing 7A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000019">
        <property type="entry name" value="Thrombospondin type 1 domain containing 7A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000020">
        <property type="entry name" value="Thrombospondin type 1 domain containing 7A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000027">
        <property type="entry name" value="Thrombospondin type 1 domain containing 7A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000031">
        <property type="entry name" value="Thrombospondin type 1 domain containing 7A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000050">
        <property type="entry name" value="Thrombospondin type 1 domain containing 7B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000015">
        <property type="entry name" value="Thrombospondin, type I, domain containing 7A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.20.100.10">
        <property type="entry name" value="Thrombospondin type-1 (TSP1) repeat"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051418">
        <property type="entry name" value="Spondin/Thrombospondin_T1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000884">
        <property type="entry name" value="TSP1_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036383">
        <property type="entry name" value="TSP1_rpt_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044004">
        <property type="entry name" value="TSP1_spondin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056991">
        <property type="entry name" value="TSP1_TSH7A-B_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11311">
        <property type="entry name" value="SPONDIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11311:SF7">
        <property type="entry name" value="THROMBOSPONDIN TYPE-1 DOMAIN-CONTAINING PROTEIN 7B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19030">
        <property type="entry name" value="TSP1_ADAMTS"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19028">
        <property type="entry name" value="TSP1_spondin"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23308">
        <property type="entry name" value="TSP1_TSH7A-B_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00090">
        <property type="entry name" value="TSP_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00209">
        <property type="entry name" value="TSP1"/>
        <property type="match status" value="17"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF82895">
        <property type="entry name" value="TSP-1 type 1 repeat"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50092">
        <property type="entry name" value="TSP1"/>
        <property type="match status" value="15"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0037">Angiogenesis</keyword>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="13" id="KW-0966">Cell projection</keyword>
    <keyword evidence="8" id="KW-0221">Differentiation</keyword>
    <keyword evidence="11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10 16" id="KW-0472">Membrane</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="6 17" id="KW-0732">Signal</keyword>
    <keyword evidence="5 16" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9 16" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="17">
        <location>
            <begin position="1"/>
            <end position="24"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019451124" description="Thrombospondin type-1 domain-containing protein 7A" evidence="17">
        <location>
            <begin position="25"/>
            <end position="1581"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="1531"/>
            <end position="1555"/>
        </location>
    </feature>
    <feature type="domain" description="Spondin-like TSP1" evidence="18">
        <location>
            <begin position="150"/>
            <end position="198"/>
        </location>
    </feature>
    <feature type="domain" description="Spondin-like TSP1" evidence="18">
        <location>
            <begin position="307"/>
            <end position="360"/>
        </location>
    </feature>
    <feature type="domain" description="Spondin-like TSP1" evidence="18">
        <location>
            <begin position="573"/>
            <end position="631"/>
        </location>
    </feature>
    <feature type="domain" description="Spondin-like TSP1" evidence="18">
        <location>
            <begin position="709"/>
            <end position="765"/>
        </location>
    </feature>
    <feature type="domain" description="Spondin-like TSP1" evidence="18">
        <location>
            <begin position="973"/>
            <end position="1016"/>
        </location>
    </feature>
    <feature type="domain" description="Spondin-like TSP1" evidence="18">
        <location>
            <begin position="1223"/>
            <end position="1276"/>
        </location>
    </feature>
    <feature type="domain" description="Spondin-like TSP1" evidence="18">
        <location>
            <begin position="1346"/>
            <end position="1405"/>
        </location>
    </feature>
    <feature type="domain" description="Thrombospondin type-1" evidence="19">
        <location>
            <begin position="1450"/>
            <end position="1489"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="1559"/>
            <end position="1581"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="15">
        <location>
            <begin position="1561"/>
            <end position="1570"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004316"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022657"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00069078"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF19028"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="Pfam" id="PF23308"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006804.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1581" mass="177629" checksum="1B052C9F3B54B746" modified="2019-05-08" version="1">MFCCCCIFPCLGFGFIGHWGRCIGDCGSGGIRSRAVWCVHVEGWTAHHSNCGLRDKPENQRSCFQVCDWHLELFEWEVSEWQICVLVPFAHGEVKPRTTDCVTAQHGLQHRSVHCIQKLNRTIVASEICEYFIPQPPVEQACLIPCPRDCVVSEFSKWSECSKDCVRSLQHRTRFVISPPLYGGSNCPNLTESRICDVPISCFLGEEEYIYSLQVGPWSKCRLPHLKEVSLIGRTMLDFSSDSAERSTFKLHSYKSHHYSKPWDVEIGYQTRQVRCTRSDGKNAMLSLCMQETIPLTFQSCVMPKDCETSVWSMWSTCSKTCHSGDLSPGFRSRHRSIKYIAVGTGKECPELEEKEACNVGGKELLHPCPRFSWRTSEWKACQVTLLLDGQDPRRHKHTALCGGGIQVREVYCVQSVIESGTHKLKEVTRPVDKKLCVGPAPSASQLCNVPCSSDCLVSSWSAWGPCVHENCQDHQGRKGFRMRQRHVVVEPIGMSQGCPHLVESIPCEDPMCYQWVISEGVCVPDHGKCGHGNRILIAICKNNKGDEVSHTLCSEFLHPETMVCEVPCAVDCVVNDWSEWSSCSHSCSSKNAEGKQSRTKSILALPGEGGKPCPPLRALQEYRLCNDHICTQFYWETSPWGPCSEDTLVTALNATISWNGEATCGVGLQTRKVFCVKNNFDQVTNKRCPDSLRPETMRPCLLPCKKDCVVTPFSEWTHCPMTCQHDNATAIKQFRYRIIIQNAANGGQDCPDTLYEERECEDIPVCPVYRWKTHKWSHCILVPDSVRQGILGINEACGRGLQSRAVVCLSDNDLAADVTECMKWTEPMPALVQECLIPCKDDCTFTPWSKFTACSSDCESTKSRRRSLTGRSRKRDKCQNTVVYPQVETEACPCETFITQPHGNWSDCIIAEGKTELQLGMRFQGETKQCGEGVRLRAIACYDKTGKLVEPSYCSSSGYIEESCVIPCPFDCKLSDWSSWASCSASCGAGMKIRSKWLKEKPYNGGRPCPTLDLKNQAQVYEAVPCYTECNQYSWVVEPWSPCKINSEQNSLHCGDGIQTRKIRCVNTAEDGEGETVPITQCDQSEIPNEAQKCNLYCPNECVMSDWGQWSQCPQVCDPNIMQTRTRHVLRSPLNTKTCPKDSQMQPCVLNQNCFQYQYNLTDWSTCQLSENATCGQGIQRRLLSCMRSDGKSVNMNYCEQLNLEKPEKLSIHCHVECSVNCQLSKWSAWSECSNTCGIGGQMIRTRSIIMQSQGEGRLCPTQLTQYRNCPVKPCYTWILGAWSPCKVEGGQCGEGFQIRNLTCIVHDASMSVPSKQVKNAFCGELPSKDNALQLPCSVPCPGDCHLTGWSEWSSCELTCIDGRSFETTGHQSRSRTFIIQSFENRDSCPEQVLETRPCSGGKCYNYAWKTSLWRDNERTVWCQRSDGVNVTGGCSVHTQPAETRHCNPECKKPFTYCTQSGVCSCERGYTEIMRSNGFLDYCMKVPGLEDKKADVKTSAGKNKPVNSKMHDIFKGWSIQPFNPDGRLKIWVYGVSAGGFLIILFLIFTSYLVCKKPKQHQSTPPQQKPLTFAYDGDMDM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXF0</accession>
    <name>A0A452GXF0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Centromere protein J</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the TCP10 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000007877.1">
        <property type="protein sequence ID" value="ENSGAGP00000006785.1"/>
        <property type="gene ID" value="ENSGAGG00000005474.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005814">
        <property type="term" value="C:centriole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015631">
        <property type="term" value="F:tubulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061511">
        <property type="term" value="P:centriole elongation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060271">
        <property type="term" value="P:cilium assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.450.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009852">
        <property type="entry name" value="CENPJ_C_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047002">
        <property type="entry name" value="Tcp10_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026581">
        <property type="entry name" value="TCP10L/CENPJ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058029">
        <property type="entry name" value="Tubulin-bd_CENPJ"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10331">
        <property type="entry name" value="T COMPLEX PROTEIN 10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10331:SF25">
        <property type="entry name" value="T-COMPLEX PROTEIN 10A-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07202">
        <property type="entry name" value="Tcp10_C"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25779">
        <property type="entry name" value="Tubulin-bind_CPAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="CENPJ tubulin-binding region" evidence="5">
        <location>
            <begin position="294"/>
            <end position="356"/>
        </location>
    </feature>
    <feature type="domain" description="Centromere protein J C-terminal" evidence="4">
        <location>
            <begin position="1185"/>
            <end position="1218"/>
        </location>
    </feature>
    <feature type="domain" description="Centromere protein J C-terminal" evidence="4">
        <location>
            <begin position="1258"/>
            <end position="1289"/>
        </location>
    </feature>
    <feature type="domain" description="Centromere protein J C-terminal" evidence="4">
        <location>
            <begin position="1296"/>
            <end position="1328"/>
        </location>
    </feature>
    <feature type="domain" description="Centromere protein J C-terminal" evidence="4">
        <location>
            <begin position="1332"/>
            <end position="1362"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="233"/>
            <end position="258"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="426"/>
            <end position="455"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="487"/>
            <end position="515"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="621"/>
            <end position="678"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="696"/>
            <end position="780"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="997"/>
            <end position="1137"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="1149"/>
            <end position="1184"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="2">
        <location>
            <begin position="905"/>
            <end position="971"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="3">
        <location>
            <begin position="236"/>
            <end position="247"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="427"/>
            <end position="455"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="496"/>
            <end position="515"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="621"/>
            <end position="630"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="3">
        <location>
            <begin position="719"/>
            <end position="729"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="758"/>
            <end position="775"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="1005"/>
            <end position="1016"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="1021"/>
            <end position="1032"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="1038"/>
            <end position="1048"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="1049"/>
            <end position="1058"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="3">
        <location>
            <begin position="1065"/>
            <end position="1082"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="1112"/>
            <end position="1137"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="1149"/>
            <end position="1173"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="1174"/>
            <end position="1184"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00005627"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="Pfam" id="PF07202"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="Pfam" id="PF25779"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006785.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="8">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1374" mass="154690" checksum="7E114B01DCCCEB7A" modified="2019-05-08" version="1">MEADTLKDEDLMAEWTANDPARLVSSPSKVTEKAMCFLPPGQINSTNGSFNEQFTPFCYSADSSMMGSNGSSSDNRFHGQAFQDAKQLNYLANSVLFPHLLASSITQTGIVEANKSSNLETLSPVAAPPAPQINQEPTCYRNMEQSPGQQEQVMMEQMEQLQRLVTEQQKIITYYNPGFSVYPGTPSQLIAMVPTLPRFPATLFPVQLPLENSSPVQNHGHLQTSPSIMQTSLQRCSPASSPNNNCSETLDESPLKEETYPNTLPAIKEEEGEQTDEQIPLSPFGIRMDARTRNLEDRPIRPGIGVRQKTFEEFVEEQLKVDSQITENHQQNFCEAKATPRESFLKRGEGIARLEKNKENLAKEQGKHPRRVSFCQNIFSFPHQQDAGKLQLGKHLNRLHQRSSSPTVLASNEKNTNCTISMSEIKAQVDSKTRDPEQCPEERREGGGGEECAKEDPAVPLQMNWAELLQQCQEQITEMKLQIGEKNKAQSTDSLGHSDRADRSSMESTVQKDLDIVDQRVKGNLTQKAEKPLEILQEGSKLQNLEGSQTKQMDWSTGLKFTSGMKGISSCHNEDYTVSKSQESQSGTTTGFKMVNDRIMKVTHKSTQGMDKKKNITSVSHQQEWQSNRSAAYRWKAPPLSSGSGCTSTDSEDDPKSYCTRSPTMHVPQKVGHTDKNLDLSDADYATDELSGAEDLALKKHTKPPPKKLSAQEIKAKQDLSLSTSSSDSGNRVARLRGNKACSSLRKSPFRPPRSARGGREPEPRSERNVEDGKNVELQSSPSTCDLVASLFPAFKSKETLAEERAQRKSVLGTLEAVQRRLTDKLEELEKEVGVYHGETPLLAMMKEEQEKARHFLRTQMDQFETSKAQELNCLEEYKRDCIHTPKKEKVEFKKYATAARVTREDVKSEEIQMLKQQIAGLQEEFRRNESHWHAAHGKLKSQIEMLTKQNLELRDELRVSEHQRLEAEKKSGAVDFISRKSETPVSEAILRGTSSLANLEERSLQSSPKSRNTTPVGRKTPVEKHPPRDVNMKAMKSTVQRTESLKSVTRESREKTPSNHFHSRSTTPTGRRTPHPGRLTPFEPEKVMHQSSLNEQRTYGRKSPVIPVSHLTVSKETNSSSYIKGRFSSTSGSSEDTVLFRNQNEDVCSSASCSNNEETQTSPPKSTLSRRSSLYEESKKRDEEVREKIEYADGKVEEVLTDGRRIITFRNGTKKEISADKRMTVVTFFNGDVKKIMPDQRVIYYYADAETTHTTYPNGLEVLQFPNNQIEKHHPDGTKEIVFPDQTVKRFYDGGLEETVFPDGTVVKVEKNGDKMVMFSNGQKEIHTSQFKRREYPDGTIKTVYSSGQQETKYSSGRVRIKDEEGNIILNKK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXF3</accession>
    <name>A0A452GXF3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Follistatin-related protein 3</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="13">Follistatin-like protein 3</fullName>
        </alternativeName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Secreted</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GXF3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007911.1">
        <property type="protein sequence ID" value="ENSGAGP00000006816.1"/>
        <property type="gene ID" value="ENSGAGG00000005498.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048185">
        <property type="term" value="F:activin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032926">
        <property type="term" value="P:negative regulation of activin receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001503">
        <property type="term" value="P:ossification"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030510">
        <property type="term" value="P:regulation of BMP signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00104">
        <property type="entry name" value="KAZAL_FS"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.60.30:FF:000025">
        <property type="entry name" value="Follistatin-related protein 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.60.30:FF:000028">
        <property type="entry name" value="Follistatin-related protein 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.290.10:FF:000021">
        <property type="entry name" value="follistatin-related protein 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.60.30">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.290.10">
        <property type="entry name" value="TGF-beta binding (TB) domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003645">
        <property type="entry name" value="Fol_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015369">
        <property type="entry name" value="Follistatin/Osteonectin_EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002350">
        <property type="entry name" value="Kazal_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036058">
        <property type="entry name" value="Kazal_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050653">
        <property type="entry name" value="Prot_Inhib_GrowthFact_Antg"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017878">
        <property type="entry name" value="TB_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036773">
        <property type="entry name" value="TB_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10913">
        <property type="entry name" value="FOLLISTATIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10913:SF16">
        <property type="entry name" value="FOLLISTATIN-RELATED PROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09289">
        <property type="entry name" value="FOLN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21333">
        <property type="entry name" value="FST_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07648">
        <property type="entry name" value="Kazal_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00274">
        <property type="entry name" value="FOLN"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00280">
        <property type="entry name" value="KAZAL"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF100895">
        <property type="entry name" value="Kazal-type serine protease inhibitors"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57581">
        <property type="entry name" value="TB module/8-cys domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51465">
        <property type="entry name" value="KAZAL_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51364">
        <property type="entry name" value="TB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="6" id="KW-0892">Osteogenesis</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0964">Secreted</keyword>
    <keyword evidence="4 14" id="KW-0732">Signal</keyword>
    <keyword evidence="9" id="KW-0804">Transcription</keyword>
    <keyword evidence="7" id="KW-0805">Transcription regulation</keyword>
    <feature type="signal peptide" evidence="14">
        <location>
            <begin position="1"/>
            <end position="19"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019586988" description="Follistatin-related protein 3" evidence="14">
        <location>
            <begin position="20"/>
            <end position="244"/>
        </location>
    </feature>
    <feature type="domain" description="TB" evidence="15">
        <location>
            <begin position="19"/>
            <end position="78"/>
        </location>
    </feature>
    <feature type="domain" description="Kazal-like" evidence="16">
        <location>
            <begin position="88"/>
            <end position="152"/>
        </location>
    </feature>
    <feature type="domain" description="Kazal-like" evidence="16">
        <location>
            <begin position="176"/>
            <end position="228"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022855"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00069916"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00077850"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS51364"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS51465"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006816.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="244" mass="26448" checksum="0791DF1691CCFFA8" modified="2019-05-08" version="1">MLISRWLLSTSTPLTSVGGICWLQQGKEAKCTMILKADVTWEECCGNSNVDVAWSNYTQPGNKISLLGFLGLVPCHPCKETCEGVECGPGKVCKMKHGRPHCACAPDCSSLPRKLQVCGSDGYTYRDECELLTAKCRDHPDLEVMYQGKCKKSCSSVVCPGTHTCVVDQTGSAHCVMCRTAPCPDPSTLDHTLCGNNNVTYPSACHLRRATCFLGRSIGVRHYGSCLAVAKFPLDVGDAEENYV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXF6</accession>
    <name>A0A452GXF6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Thrombospondin type-1 domain-containing protein 7A</fullName>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cell projection</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000007907.1">
        <property type="protein sequence ID" value="ENSGAGP00000006812.1"/>
        <property type="gene ID" value="ENSGAGG00000005480.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000007917.1">
        <property type="protein sequence ID" value="ENSGAGP00000006822.1"/>
        <property type="gene ID" value="ENSGAGG00000005480.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042995">
        <property type="term" value="C:cell projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030036">
        <property type="term" value="P:actin cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001525">
        <property type="term" value="P:angiogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000014">
        <property type="entry name" value="Thrombospondin type 1 domain containing 7A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000017">
        <property type="entry name" value="Thrombospondin type 1 domain containing 7A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000018">
        <property type="entry name" value="Thrombospondin type 1 domain containing 7A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000019">
        <property type="entry name" value="Thrombospondin type 1 domain containing 7A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000020">
        <property type="entry name" value="Thrombospondin type 1 domain containing 7A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000027">
        <property type="entry name" value="Thrombospondin type 1 domain containing 7A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000031">
        <property type="entry name" value="Thrombospondin type 1 domain containing 7A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000050">
        <property type="entry name" value="Thrombospondin type 1 domain containing 7B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.100.10:FF:000015">
        <property type="entry name" value="Thrombospondin, type I, domain containing 7A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.20.100.10">
        <property type="entry name" value="Thrombospondin type-1 (TSP1) repeat"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051418">
        <property type="entry name" value="Spondin/Thrombospondin_T1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000884">
        <property type="entry name" value="TSP1_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036383">
        <property type="entry name" value="TSP1_rpt_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044004">
        <property type="entry name" value="TSP1_spondin_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11311">
        <property type="entry name" value="SPONDIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11311:SF7">
        <property type="entry name" value="THROMBOSPONDIN TYPE-1 DOMAIN-CONTAINING PROTEIN 7B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19030">
        <property type="entry name" value="TSP1_ADAMTS"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19028">
        <property type="entry name" value="TSP1_spondin"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00090">
        <property type="entry name" value="TSP_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00209">
        <property type="entry name" value="TSP1"/>
        <property type="match status" value="17"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF82895">
        <property type="entry name" value="TSP-1 type 1 repeat"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50092">
        <property type="entry name" value="TSP1"/>
        <property type="match status" value="15"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0037">Angiogenesis</keyword>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="13" id="KW-0966">Cell projection</keyword>
    <keyword evidence="8" id="KW-0221">Differentiation</keyword>
    <keyword evidence="11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10" id="KW-0472">Membrane</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="6 15" id="KW-0732">Signal</keyword>
    <keyword evidence="5" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="15">
        <location>
            <begin position="1"/>
            <end position="36"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5044604475" description="Thrombospondin type-1 domain-containing protein 7A" evidence="15">
        <location>
            <begin position="37"/>
            <end position="1512"/>
        </location>
    </feature>
    <feature type="domain" description="Spondin-like TSP1" evidence="16">
        <location>
            <begin position="180"/>
            <end position="228"/>
        </location>
    </feature>
    <feature type="domain" description="Spondin-like TSP1" evidence="16">
        <location>
            <begin position="337"/>
            <end position="390"/>
        </location>
    </feature>
    <feature type="domain" description="Spondin-like TSP1" evidence="16">
        <location>
            <begin position="603"/>
            <end position="661"/>
        </location>
    </feature>
    <feature type="domain" description="Spondin-like TSP1" evidence="16">
        <location>
            <begin position="739"/>
            <end position="795"/>
        </location>
    </feature>
    <feature type="domain" description="Spondin-like TSP1" evidence="16">
        <location>
            <begin position="1003"/>
            <end position="1046"/>
        </location>
    </feature>
    <feature type="domain" description="Spondin-like TSP1" evidence="16">
        <location>
            <begin position="1253"/>
            <end position="1306"/>
        </location>
    </feature>
    <feature type="domain" description="Spondin-like TSP1" evidence="16">
        <location>
            <begin position="1376"/>
            <end position="1435"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004316"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022657"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00069078"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF19028"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006822.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1512" mass="170453" checksum="5AD82CC63EB91F3F" modified="2019-05-08" version="1">MFLRDLWVSAHWAWQGMRNILLLLVLSMLQTTDLEARKDHQFVWKTGHWGRCIGDCGSGGIRSRAVWCVHVEGWTAHHSNCGLRDKPENQRSCFQVCDWHLELFEWEVSEWQICVLVPFAHGEVKPRTTDCVTAQHGLQHRSVHCIQKLNRTIVASEICEYFIPQPPVEQACLIPCPRDCVVSEFSKWSECSKDCVRSLQHRTRFVISPPLYGGSNCPNLTESRICDVPISCFLGEEEYIYSLQVGPWSKCRLPHLKEVSLIGRTMLDFSSDSAERSTFKLHSYKSHHYSKPWDVEIGYQTRQVRCTRSDGKNAMLSLCMQETIPLTFQSCVMPKDCETSVWSMWSTCSKTCHSGDLSPGFRSRHRSIKYIAVGTGKECPELEEKEACNVGGKELLHPCPRFSWRTSEWKACQVTLLLDGQDPRRHKHTALCGGGIQVREVYCVQSVIESGTHKLKEVTRPVDKKLCVGPAPSASQLCNVPCSSDCLVSSWSAWGPCVHENCQDHQGRKGFRMRQRHVVVEPIGMSQGCPHLVESIPCEDPMCYQWVISEGVCVPDHGKCGHGNRILIAICKNNKGDEVSHTLCSEFLHPETMVCEVPCAVDCVVNDWSEWSSCSHSCSSKNAEGKQSRTKSILALPGEGGKPCPPLRALQEYRLCNDHICTQFYWETSPWGPCSEDTLVTALNATISWNGEATCGVGLQTRKVFCVKNNFDQVTNKRCPDSLRPETMRPCLLPCKKDCVVTPFSEWTHCPMTCQHDNATAIKQFRYRIIIQNAANGGQDCPDTLYEERECEDIPVCPVYRWKTHKWSHCILVPDSVRQGILGINEACGRGLQSRAVVCLSDNDLAADVTECMKWTEPMPALVQECLIPCKDDCTFTPWSKFTACSSDCESTKSRRRSLTGRSRKRDKCQNTVVYPQVETEACPCETFITQPHGNWSDCIIAEGKTELQLGMRFQGETKQCGEGVRLRAIACYDKTGKLVEPSYCSSSGYIEESCVIPCPFDCKLSDWSSWASCSASCGAGMKIRSKWLKEKPYNGGRPCPTLDLKNQAQVYEAVPCYTECNQYSWVVEPWSPCKINSEQNSLHCGDGIQTRKIRCVNTAEDGEGETVPITQCDQSEIPNEAQKCNLYCPNECVMSDWGQWSQCPQVCDPNIMQTRTRHVLRSPLNTKTCPKDSQMQPCVLNQNCFQYQYNLTDWSTCQLSENATCGQGIQRRLLSCMRSDGKSVNMNYCEQLNLEKPEKLSIHCHVECSVNCQLSKWSAWSECSNTCGIGGQMIRTRSIIMQSQGEGRLCPTQLTQYRNCPVKPCYTWILGAWSPCKVEGGQCGEGFQIRNLTCIVHDASMSVPSKQVKNAFCGELPSKDNALQLPCSVPCPGDCHLTGWSEWSSCELTCIDGRSFETTGHQSRSRTFIIQSFENRDSCPEQVLETRPCSGGKCYNYAWKTSLWRDNERTVWCQRSDGVNVTGGCSVHTQPAETRHCNPECKKPFTYCTQVRGCCRRYFRNSLISNILEAF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXF9</accession>
    <name>A0A452GXF9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">E3 ubiquitin-protein ligase RNF126</fullName>
            <ecNumber evidence="5">2.3.2.27</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="18">RING finger protein 126</fullName>
        </alternativeName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="24">
            <text>S-ubiquitinyl-[E2 ubiquitin-conjugating enzyme]-L-cysteine + [acceptor protein]-L-lysine = [E2 ubiquitin-conjugating enzyme]-L-cysteine + N(6)-ubiquitinyl-[acceptor protein]-L-lysine.</text>
            <dbReference type="EC" id="2.3.2.27"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="4">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="subunit">
        <text evidence="16">Interacts with CCDC50, EGFR, FLT3 and SCAMP3. Interacts with BAG6 (via ubiquitin-like domain); required for BAG6-dependent ubiquitination of proteins mislocalized to the cytosol. Interacts with CDKN1A. Interacts with AICDA.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="2.3.2.27" evidence="5"/>
    <dbReference type="AlphaFoldDB" id="A0A452GXF9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007925.1">
        <property type="protein sequence ID" value="ENSGAGP00000006829.1"/>
        <property type="gene ID" value="ENSGAGG00000005505.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061630">
        <property type="term" value="F:ubiquitin protein ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000209">
        <property type="term" value="P:protein polyubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd16801">
        <property type="entry name" value="RING-H2_RNF126"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.40.10:FF:000253">
        <property type="entry name" value="E3 ubiquitin-protein ligase RNF126"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039571">
        <property type="entry name" value="RNF126_RING-H2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001841">
        <property type="entry name" value="Znf_RING"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15710">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE PRAJA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15710:SF21">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE RNF126"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13639">
        <property type="entry name" value="zf-RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00184">
        <property type="entry name" value="RING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57850">
        <property type="entry name" value="RING/U-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50089">
        <property type="entry name" value="ZF_RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="14" id="KW-0007">Acetylation</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="15" id="KW-0539">Nucleus</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0808">Transferase</keyword>
    <keyword evidence="13" id="KW-0832">Ubl conjugation</keyword>
    <keyword evidence="11" id="KW-0833">Ubl conjugation pathway</keyword>
    <keyword evidence="12" id="KW-0862">Zinc</keyword>
    <keyword evidence="10 19" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="RING-type" evidence="21">
        <location>
            <begin position="213"/>
            <end position="254"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="21"/>
            <end position="45"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="77"/>
            <end position="110"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="261"/>
            <end position="295"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="20">
        <location>
            <begin position="85"/>
            <end position="98"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="20">
        <location>
            <begin position="99"/>
            <end position="110"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="20">
        <location>
            <begin position="275"/>
            <end position="295"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00061906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00067468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00080437"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006829.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="295" mass="32406" checksum="10079C57F5463FAC" modified="2019-05-08" version="1">KPFENQHDYICPRCESGFIEELPEEPRNAENGSSSSASTSDQSRHPFENVDQHLFTLPQGYGQFAFGIFDDSFEFPAFGSSGQSEDNRDSENRREREHQSRHRYGARQPRARLTARRATGRHEGVPTLEGIIQQLVNGIIAPTTIPNLGLGPWGVLHSNPMDYAWGANGLDAIITQLLNQFENTGPPPADKEKIQALPTVQITEEHVGSGLECPVCKEDYTLGENVRQLPCNHLFHNGCIVPWLEQHDTCPVCRKSLSGQNTATNPPGLTGMNFSSSSSSSSSSSPSNENSSNNS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXG2</accession>
    <name>A0A452GXG2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">C2H2-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="13">Involved in histone 3'-end pre-mRNA processing by associating with U7 snRNP and interacting with SLBP/pre-mRNA complex. Increases histone 3'-end pre-mRNA processing but has no effect on U7 snRNP levels, when overexpressed. Required for cell cycle progression from G1 to S phases.</text>
    </comment>
    <comment type="subunit">
        <text evidence="14">Interacts with the SLBP/pre-mRNA complex but not with SLBP alone. Interacts with LSM11 in a U7 snRNP-dependent manner.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GXG2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007897.1">
        <property type="protein sequence ID" value="ENSGAGP00000006803.1"/>
        <property type="gene ID" value="ENSGAGG00000005488.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000111">
        <property type="entry name" value="GLI family zinc finger 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000478">
        <property type="entry name" value="Zinc finger protein 133"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000631">
        <property type="entry name" value="zinc finger protein 189 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000295">
        <property type="entry name" value="zinc finger protein 19"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000358">
        <property type="entry name" value="zinc finger protein 24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000269">
        <property type="entry name" value="Zinc finger protein 287"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002343">
        <property type="entry name" value="Zinc finger protein 33A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000016">
        <property type="entry name" value="zinc finger protein 37 homolog"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001119">
        <property type="entry name" value="zinc finger protein 408"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002090">
        <property type="entry name" value="Zinc finger protein 473"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000340">
        <property type="entry name" value="zinc finger protein 473 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:001270">
        <property type="entry name" value="zinc finger protein 583 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000609">
        <property type="entry name" value="zinc finger protein 621"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000495">
        <property type="entry name" value="zinc finger protein 668"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000642">
        <property type="entry name" value="Zinc finger with KRAB and SCAN domains 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="19"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050752">
        <property type="entry name" value="C2H2-ZF_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24384">
        <property type="entry name" value="FINGER PUTATIVE TRANSCRIPTION FACTOR FAMILY-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24384:SF218">
        <property type="entry name" value="ZINC FINGER PROTEIN 502"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="17"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="18"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="17"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="18"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="12" id="KW-0539">Nucleus</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="11" id="KW-0804">Transcription</keyword>
    <keyword evidence="9" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="8" id="KW-0832">Ubl conjugation</keyword>
    <keyword evidence="7" id="KW-0862">Zinc</keyword>
    <keyword evidence="6 15" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="17">
        <location>
            <begin position="19"/>
            <end position="46"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="17">
        <location>
            <begin position="47"/>
            <end position="74"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="17">
        <location>
            <begin position="75"/>
            <end position="102"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="17">
        <location>
            <begin position="103"/>
            <end position="130"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="17">
        <location>
            <begin position="131"/>
            <end position="158"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="17">
        <location>
            <begin position="159"/>
            <end position="186"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="17">
        <location>
            <begin position="187"/>
            <end position="214"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="17">
        <location>
            <begin position="236"/>
            <end position="263"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="17">
        <location>
            <begin position="264"/>
            <end position="291"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="17">
        <location>
            <begin position="316"/>
            <end position="343"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="17">
        <location>
            <begin position="344"/>
            <end position="371"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="17">
        <location>
            <begin position="372"/>
            <end position="399"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="17">
        <location>
            <begin position="400"/>
            <end position="427"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="17">
        <location>
            <begin position="428"/>
            <end position="455"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="17">
        <location>
            <begin position="456"/>
            <end position="483"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="17">
        <location>
            <begin position="484"/>
            <end position="511"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="17">
        <location>
            <begin position="517"/>
            <end position="544"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="17">
        <location>
            <begin position="545"/>
            <end position="572"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="277"/>
            <end position="308"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="16">
        <location>
            <begin position="286"/>
            <end position="299"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00054787"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00062613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006803.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="605" mass="69354" checksum="BD773E3CC96D3F28" modified="2019-05-08" version="1">MRLSNLSRHRRSHTGERSHKCTQCGKSFHFNSQLVQHQISHTGERPYKCPDCGKSFGVSSNLLQHQRIHWVERPYSCANCEKSFGHSSLLAQHQRRHMDERPYKCGECGKAFGVSSALIRHQRIHTGERPYKCLVCGKSFNLSSNLFTHRRIHWDEKPYKCPECGRSFSQSSSLSRHQISHTGERPYKCPECGKCFNQSSLLIRHQRIHTGERPYRCSQRSNNMQHQRIHTGERPYKCTDCGKSFSLSSRLLQHQMVHTGERPYKCPDCGKSFNQKSNLKQHQRIHTGERPYRRPDYGESHSLSPPQGIHKLERPYKCPQCGKCFRVSSHLIRHQVTHTGERPYKCSICGRSFSLSSHLIRHRIVHTEDKPYVCSECGKKFSLSSRLDRHLRTHTGERPYKCPNCGSGFVDSSALIKHQRIHTGERPYTCPECGKSFGRSSYLITHQRTHTGERPYKCPDCGKSFVGSSDLNKHQRTHTGERPYNCPECGKSFSRSSNLIAHQRIHARGRTKRERANTSPKCGKSFPWSSQLLQHQRSHTRERPYQCPDCGKGFGESLALIQHQRTHTRDRVEPLVCLHLGAGLAAGCFQGPAWFAMSGQAGSLP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXG8</accession>
    <name>A0A452GXG8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">RCK N-terminal domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>K(+)(in) = K(+)(out)</text>
            <dbReference type="Rhea" id="RHEA:29463"/>
            <dbReference type="ChEBI" id="CHEBI:29103"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GXG8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007809.1">
        <property type="protein sequence ID" value="ENSGAGP00000006723.1"/>
        <property type="gene ID" value="ENSGAGG00000005143.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005228">
        <property type="term" value="F:intracellular sodium-activated potassium channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015271">
        <property type="term" value="F:outward rectifier potassium channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.720:FF:000011">
        <property type="entry name" value="Potassium channel subfamily T member 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.720:FF:000034">
        <property type="entry name" value="Potassium channel subfamily T member 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.70:FF:000168">
        <property type="entry name" value="Potassium sodium-activated channel subfamily T member 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.70">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.720">
        <property type="entry name" value="NAD(P)-binding Rossmann-like Domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003929">
        <property type="entry name" value="K_chnl_BK_asu"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013099">
        <property type="entry name" value="K_chnl_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047871">
        <property type="entry name" value="K_chnl_Slo-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003148">
        <property type="entry name" value="RCK_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10027">
        <property type="entry name" value="CALCIUM-ACTIVATED POTASSIUM CHANNEL ALPHA CHAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10027:SF14">
        <property type="entry name" value="POTASSIUM CHANNEL SUBFAMILY T MEMBER 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03493">
        <property type="entry name" value="BK_channel_a"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07885">
        <property type="entry name" value="Ion_trans_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22614">
        <property type="entry name" value="Slo-like_RCK"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81324">
        <property type="entry name" value="Voltage-gated potassium channels"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51201">
        <property type="entry name" value="RCK_N"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="10" id="KW-0407">Ion channel</keyword>
    <keyword evidence="8" id="KW-0406">Ion transport</keyword>
    <keyword evidence="9 13" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-0630">Potassium</keyword>
    <keyword evidence="5" id="KW-0631">Potassium channel</keyword>
    <keyword evidence="3" id="KW-0633">Potassium transport</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 13" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 13" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="2" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="13">
        <location>
            <begin position="96"/>
            <end position="115"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="13">
        <location>
            <begin position="145"/>
            <end position="163"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="13">
        <location>
            <begin position="206"/>
            <end position="227"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="13">
        <location>
            <begin position="233"/>
            <end position="252"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="13">
        <location>
            <begin position="264"/>
            <end position="289"/>
        </location>
    </feature>
    <feature type="domain" description="RCK N-terminal" evidence="14">
        <location>
            <begin position="306"/>
            <end position="442"/>
        </location>
    </feature>
    <feature type="domain" description="RCK N-terminal" evidence="14">
        <location>
            <begin position="729"/>
            <end position="869"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1"/>
            <end position="28"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1145"/>
            <end position="1174"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="8"/>
            <end position="17"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="1159"/>
            <end position="1174"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022538"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022826"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022958"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00034430"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS51201"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006723.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1174" mass="134025" checksum="3D2E82680648A748" modified="2019-05-08" version="1">MARAKLKNSPSEINSHVKTVPPATTEDVHGVSPLLPARRMGSLGSDVGQRPHAEDFSMDSSLSQVQVEFYVNENTFKERLKLFFIKNQRSSLRIRLFNFSLKLLTCLLYIVRVLLDNPAPIMIVSKHLSVCLMFRGNIWEQIFRISFILEMINTVPFIITIFWSPLRNLFIPVFLNCWLAKYALENMINDLHRAIQRTQSAMFNQVLILICTLLCLVFTGTCGIQHLERAGDNLSLFKSFYFCIVTFSTVGYGDVTPKIWPSQLLVVIMICVALVVLPLQFEELVYLWMERQKSGGNYSRHRAQTEKHVVLCVSSLKIDLLMDFLNEFYAHPRLQDYYVVILCPTEMDIQVRRVLQIPLWSQRVIYLQGSALKDQDLMRAKMDNGEACFILSSRNEADRTAADHQTILRAWAVKDFAPNCPLYVQILKPENKFHVKFADHVVCEEECKYAMLALNCVCPATSTLITLLVHTSRGQEGQESPEQWQRMYGRCSGNEVYHIRMGDSKFFMEYEGKSFTYAAFHAHKKYGVCLIGIRREENKSILLNPGPRHIMTASDTCFYINITKEENSAFIFKQEEKQKKKGFAGRGIYDGPSRLPVHSIIASMGTVAMDLQNTECRPVNSSKLTLPSENGSGSRRPSIAPVLELADSSSLLPCDLLSDQSEDEMTQSDDEGLTVVEYVKGYPPNSPYIGSSPTLCHLLPVKAPFCCLRLDKGCKHNSYEDAKAYGFKNKLIIVSAETAGNGLYNFIVPLRAYYRSRKELNPIVLLLDNKPENHFLEAICCFPMVYYMEGTIDNLDSLLQCGIIYADNLVVVDKESTMSAEEDYMADAKTIVNVQTMFRLFPSLSIITELTHPSNMRFMQFRAKDSYSLALSKLEKKERENGSNLAFMFRLPFAAGRVFSISMLDTLLYQSFVKDYMITITRLLLGLDTTPGSGYLCAMKITEDDLWIRTYGRLFQKLCSSSAEIPIGIYRTESHMFSTSEVRNSQISINVEDCEDTKDVKEHWNIKTSHHRNSCSSDQSEHPLLRRKSMQWARRLSRKGNKHTSKTAEWISQQRLSLYRRSERQELSELVKNRMKHLGLPTTGYEDVANLTASDVMNRVNLGYLQDEMNDHQNTLSYVLINPPPDTRLELNDIVYLIRSDPLAHVSNDSHSRKSSCSKKLDPCNPETRDETQL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXJ3</accession>
    <name>A0A452GXJ3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Deoxyribonuclease</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 10">Belongs to the DNase I family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GXJ3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007932.1">
        <property type="protein sequence ID" value="ENSGAGP00000006836.1"/>
        <property type="gene ID" value="ENSGAGG00000005510.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005783">
        <property type="term" value="C:endoplasmic reticulum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004530">
        <property type="term" value="F:deoxyribonuclease I activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006308">
        <property type="term" value="P:DNA catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd10282">
        <property type="entry name" value="DNase1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.60.10.10:FF:000007">
        <property type="entry name" value="Deoxyribonuclease"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.60.10.10">
        <property type="entry name" value="Endonuclease/exonuclease/phosphatase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016202">
        <property type="entry name" value="DNase_I"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036691">
        <property type="entry name" value="Endo/exonu/phosph_ase_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005135">
        <property type="entry name" value="Endo/exonuclease/phosphatase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11371">
        <property type="entry name" value="DEOXYRIBONUCLEASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11371:SF28">
        <property type="entry name" value="DEOXYRIBONUCLEASE-1-LIKE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03372">
        <property type="entry name" value="Exo_endo_phos"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000988">
        <property type="entry name" value="DNase_I_euk"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00130">
        <property type="entry name" value="DNASEI"/>
    </dbReference>
    <dbReference type="SMART" id="SM00476">
        <property type="entry name" value="DNaseIc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56219">
        <property type="entry name" value="DNase I-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5 10" id="KW-0255">Endonuclease</keyword>
    <keyword evidence="7" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="9" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6 10" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="3 10" id="KW-0540">Nuclease</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 13" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="13">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019167695" description="Deoxyribonuclease" evidence="13">
        <location>
            <begin position="21"/>
            <end position="310"/>
        </location>
    </feature>
    <feature type="domain" description="Endonuclease/exonuclease/phosphatase" evidence="14">
        <location>
            <begin position="25"/>
            <end position="273"/>
        </location>
    </feature>
    <feature type="active site" evidence="11">
        <location>
            <position position="99"/>
        </location>
    </feature>
    <feature type="active site" evidence="11">
        <location>
            <position position="154"/>
        </location>
    </feature>
    <feature type="disulfide bond" description="Essential for enzymatic activity" evidence="12">
        <location>
            <begin position="193"/>
            <end position="230"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007359"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022759"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PIRNR" id="PIRNR000988"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PIRSR" id="PIRSR000988-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PIRSR" id="PIRSR000988-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="Pfam" id="PF03372"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006836.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="310" mass="33977" checksum="37289114C4228F1A" modified="2019-05-08" version="1">MGLPLRTLLLLLLLCPAGAPFRICAFNLQRFASPKAATAAVMDTLVKIVSRCDIAVLQEVMDAKGQAVPALISALHRSAGPDSYTALGSPLLGAGNYQERYVFVYRSGCTQVLDSYVYLDENPDRPDAFAREPFVVHFSLPSKVLPTLVLVPQHTVPKKAEAEIDALYDVFLDVQARWGTEDIMFLGDFNADCGYVAKKRWGQIQLRREPGFHWLIGDTADTTVRNSTHCAYDRIVVHGERCLGLVVPGSAQPFDFPAMFGLTETEALKVSDHYPVEVQLELNTAPRGHCPAGPALLPLAMLLARLGGWL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXJ4</accession>
    <name>A0A452GXJ4_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="11">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="domain">
        <text evidence="6">The CKK domain binds microtubules.</text>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the CAMSAP1 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000007838.1">
        <property type="protein sequence ID" value="ENSGAGP00000006750.1"/>
        <property type="gene ID" value="ENSGAGG00000005443.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036449">
        <property type="term" value="C:microtubule minus-end"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051011">
        <property type="term" value="F:microtubule minus-end binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030507">
        <property type="term" value="F:spectrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031122">
        <property type="term" value="P:cytoplasmic microtubule organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007026">
        <property type="term" value="P:negative regulation of microtubule depolymerization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031175">
        <property type="term" value="P:neuron projection development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd22265">
        <property type="entry name" value="UDM1_RNF168"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.20.360:FF:000001">
        <property type="entry name" value="Calmodulin-regulated spectrin-associated protein 3 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.418.10">
        <property type="entry name" value="Calponin-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.20.360">
        <property type="entry name" value="CKK domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032940">
        <property type="entry name" value="CAMSAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031372">
        <property type="entry name" value="CAMSAP_CC1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR058042">
        <property type="entry name" value="CAMSAP_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022613">
        <property type="entry name" value="CH_CAMSAP_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001715">
        <property type="entry name" value="CH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036872">
        <property type="entry name" value="CH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014797">
        <property type="entry name" value="CKK_CAMSAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038209">
        <property type="entry name" value="CKK_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011033">
        <property type="entry name" value="PRC_barrel-like_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21595:SF3">
        <property type="entry name" value="CALMODULIN-REGULATED SPECTRIN-ASSOCIATED PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21595">
        <property type="entry name" value="PATRONIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17095">
        <property type="entry name" value="CAMSAP_CC1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11971">
        <property type="entry name" value="CAMSAP_CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08683">
        <property type="entry name" value="CAMSAP_CKK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25532">
        <property type="entry name" value="CH_CAMSAP2_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01051">
        <property type="entry name" value="CAMSAP_CKK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47576">
        <property type="entry name" value="Calponin-homology domain, CH-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50346">
        <property type="entry name" value="PRC-barrel domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50021">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51508">
        <property type="entry name" value="CKK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4 7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="5" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="3 6" id="KW-0493">Microtubule</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Calponin-homology (CH)" evidence="9">
        <location>
            <begin position="217"/>
            <end position="332"/>
        </location>
    </feature>
    <feature type="domain" description="CKK" evidence="10">
        <location>
            <begin position="1458"/>
            <end position="1592"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="352"/>
            <end position="483"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="671"/>
            <end position="690"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="760"/>
            <end position="803"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="817"/>
            <end position="863"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1051"/>
            <end position="1163"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1235"/>
            <end position="1258"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1295"/>
            <end position="1447"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="868"/>
            <end position="898"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="377"/>
            <end position="389"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="671"/>
            <end position="689"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="760"/>
            <end position="782"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="790"/>
            <end position="803"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="823"/>
            <end position="834"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="850"/>
            <end position="861"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="1056"/>
            <end position="1065"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="1102"/>
            <end position="1114"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="1116"/>
            <end position="1127"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="1136"/>
            <end position="1150"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="1295"/>
            <end position="1341"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="8">
        <location>
            <begin position="1356"/>
            <end position="1367"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="1375"/>
            <end position="1387"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="1388"/>
            <end position="1403"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="1404"/>
            <end position="1420"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00841"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS51508"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006750.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1597" mass="178391" checksum="D1879590994510CF" modified="2019-05-08" version="1">MVDVDVCAVGDSIRRKMDALTDSAVEIVPLELYDSARAKIAANLQWICAKAYGIDNIPEELKDPFYIDQYEQEHIKPPVIKLLLSSELYCRVCSLILKGDQVAALQGHQSVIQALSRKGIYVMESDDTPVSESDLASAPIKMGSHMAMIDALMMAYTVEMISIEKVVASVKRFSTFSASKELPYDLEDAMVFWINKVNLKMREITEKEVKLKQQLLESPGHQKVRYRRDHLSSRQLPYFPLLEDLMKDGSDGAALLAVIHYYCPEQMKLDDICLKEVTSIADSLYNIQLLREFSNEYLNKCFYLTLEDMLYAPSVLKPNVMVFIAELFWWFENVKPDFVQPRDIQEIKDAKTVLQQKSSRPPVPISNATKRSFMASPASTNPADLQPSIQLPPEACNSGKGSPAFSPSHPLLPLRQKQQKSLQGEDSSGHRHRSNSLTRVDGQPRGSVLAWPEKKPRPLSQPTPFALHHATSSDVDPGSGDSISLARSISKDSLASNIVNLTPKHQSHPSSIKTNGKSLLNNVEIEDEDEELIAIIRSDEIPDRGDLELKSVSSRAPSLVATAWSSKTQSEAIESKPDSFYLEPLMPAVLKLAKEKQMINKEDECGEGKQRGFITKRLNEGHLPMVRKKANSSHGDHDINRTFTPISSSDFTAVADPCTADSMTLSEAGLEAPSPLASSSLDPSSQEQSTGGFFLHTAKVDDDVTSKVNVSYVKSTNSHVPSTTWTMVRQDSDSDLDAEVAEQDLVVVDDHPVVTKYLGEEESAKLQEDMKVKEHEDKDDASGRSSPCLSTISQVSSVSMTSGSVRMTNFAERKLQRLNSYETKSSTSSSQKTTPDGSESCPAPLTTWKQKREQSPTRQNKDNVNLLASELVQLHMQLEEKRRAIEAQKKKMEALSARQRLKLGKAAFLHVVKKGKAVDVLQPLKPEHFAKEYSRNSGEDLDEVSLSSKSEEFLVRGEEREEILYDSQEVTKVKMQENIAFVELHKPKESAAMHEIEKSKIISAALLEDNGSEVVDISECDLSIEKLNETISTLQQAILKISHQQELLMKSPTVPSPSTRNNSQDQKVKPSIHFVEPLSPTGMSNLRKPPRLGQGRSPRSGKPIELKVAKDRHQNSTRIKTPTQSVETLPHLRPFPSNNSFRTPTETGFENSPDHGSGSQDKFFFDSYRRHDESNQQACVLSTSKDANIISEMSKDVNSYLKEGLNSSDGSGKENIPVDEPLRSKVNLIEVDLSDLKAPDEEGEIESQDSSTGIINEGDQKSGLGFFFKDEQKAEDELAKKRAAFLLKQQRKAEEARIRKQQLEAEVEQKRDEARRKAEEDRIRKEEEKARRELIKQEYLRKKQQQILEEQGLGKPKSKPKKPRPKSVHREESYSDSGTKCSSTPDNLSSAQSGSSLSLASAATTEPESIHSGGTPSQRVESMESLPILSRNPSRNTERDWENASTASSIASVAEYTGPKLFKEPSSKSNKPIIHNAISHCCLAGKVNEPHKNSILEELEKCDANHYIILFRDAGCQFRALYCYYPDTEEIYKLTGTGPKSINKKMIDKLYKYSSDRKQFNVIPAKTMSVSVDALTIHNYLWQAKRPAVPKKTQTRK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXJ5</accession>
    <name>A0A452GXJ5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Basigin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="9">Basolateral cell membrane</location>
            <topology evidence="9">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GXJ5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007966.1">
        <property type="protein sequence ID" value="ENSGAGP00000006866.1"/>
        <property type="gene ID" value="ENSGAGG00000005534.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030424">
        <property type="term" value="C:axon"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016323">
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        <property type="term" value="P:homophilic cell adhesion via plasma membrane adhesion molecules"/>
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        <property type="entry name" value="Neuroplastin b"/>
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        <property type="entry name" value="Immunoglobulins"/>
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        <property type="entry name" value="Ig-like_fold"/>
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        <property type="entry name" value="Ig_sub"/>
    </dbReference>
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        <property type="entry name" value="BASIGIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10075">
        <property type="entry name" value="BASIGIN RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Ig_3"/>
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    </dbReference>
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        <property type="entry name" value="BASIGIN"/>
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    </dbReference>
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    </dbReference>
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    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="5" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="7" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="8" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="4 12" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-0675">Receptor</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="13" id="KW-0732">Signal</keyword>
    <keyword evidence="12" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="12" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="13">
        <location>
            <begin position="1"/>
            <end position="24"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5018992310" description="Basigin" evidence="13">
        <location>
            <begin position="25"/>
            <end position="272"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="213"/>
            <end position="234"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="14">
        <location>
            <begin position="27"/>
            <end position="108"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="14">
        <location>
            <begin position="111"/>
            <end position="208"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="242"/>
            <end position="272"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004115"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
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        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023768"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023876"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006866.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="272" mass="29498" checksum="BBAF7E13E8F6BDCB" modified="2019-05-08" version="1">MRAAGATGLVLLGALLCAGRGASGADPIVETSPNVTSGLDVVISCNISEPPSPITGHMWKKGDKILTSDKETSDYTTYKIGEVNGDSSGVYECIFETDPPVSRTVYVTVKPHVTAYKKSEHGNEGDTGVMICRCNSYPPVSQWTWYKIKSDVNEAIINGTEKFIIKSSGNKTELRIANLDIEKDPGEYICNATNELGTSGAVVGLRVRSRLAALWPFLGIVAEVLILVTIIFIYEKRRKPDEVLDDDDGGSAPLKSNPPNHKDKNVRQRNAN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXJ7</accession>
    <name>A0A452GXJ7_9SAUR</name>
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        <recommendedName>
            <fullName evidence="6">Fibroblast growth factor</fullName>
            <shortName evidence="6">FGF</shortName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1 6">Belongs to the heparin-binding growth factors family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GXJ7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007938.1">
        <property type="protein sequence ID" value="ENSGAGP00000006841.1"/>
        <property type="gene ID" value="ENSGAGG00000005513.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008083">
        <property type="term" value="F:growth factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
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        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
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        <property type="term" value="P:positive regulation of cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
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        <property type="entry name" value="beta-trefoil_FGF22"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Fibroblast growth factor"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Fibroblast_GF_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008996">
        <property type="entry name" value="IL1/FGF"/>
    </dbReference>
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        <property type="entry name" value="FIBROBLAST GROWTH FACTOR"/>
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    </dbReference>
    <dbReference type="Pfam" id="PF00167">
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    </dbReference>
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        <property type="entry name" value="HBGFFGF"/>
    </dbReference>
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        <property type="entry name" value="IL1HBGF"/>
    </dbReference>
    <dbReference type="SMART" id="SM00442">
        <property type="entry name" value="FGF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50353">
        <property type="entry name" value="Cytokine"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="3" id="KW-0221">Differentiation</keyword>
    <keyword evidence="4" id="KW-0339">Growth factor</keyword>
    <keyword evidence="5" id="KW-0497">Mitogen</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="6">
        <location>
            <begin position="1"/>
            <end position="25"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5018816054" description="Fibroblast growth factor" evidence="6">
        <location>
            <begin position="26"/>
            <end position="185"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="25"/>
            <end position="48"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="30"/>
            <end position="42"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007936"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023030"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="RuleBase" id="RU049442"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006841.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="185" mass="20565" checksum="8E1AA306CA617160" modified="2019-05-08" version="1">MGRQVAAGPVSLWLCLALSLLTGSAHRDPAAPSGSNPSGAPSRRPRSYNHLEGDVRWRRLYVSTHFFLCIDSSGKVQGTRNNNTGLGILEIRSVRVGVVAIKSVHTGFYLAMNKKGKVYGSKEYNPNCKFKERIEENGYNTYASLRWRPGNRPMFLALNSKGRPRQGGKARRQHLSTHFLPLLVS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXK0</accession>
    <name>A0A452GXK0_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="15">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GXK0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007974.1">
        <property type="protein sequence ID" value="ENSGAGP00000006874.1"/>
        <property type="gene ID" value="ENSGAGG00000002943.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd08849">
        <property type="entry name" value="ArfGap_ASAP2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd13251">
        <property type="entry name" value="PH_ASAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.220.150:FF:000002">
        <property type="entry name" value="arf-GAP with SH3 domain, ANK repeat and PH domain-containing protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.950:FF:000001">
        <property type="entry name" value="Arf-GAP with SH3 domain, ANK repeat and PH domain-containing protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000006">
        <property type="entry name" value="Arf-GAP with SH3 domain, ANK repeat and PH domain-containing protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000012">
        <property type="entry name" value="Arf-GAP with SH3 domain, ANK repeat and PH domain-containing protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.950">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.220.150">
        <property type="entry name" value="Arf GTPase activating protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.60">
        <property type="entry name" value="Arfaptin homology (AH) domain/BAR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027267">
        <property type="entry name" value="AH/BAR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037278">
        <property type="entry name" value="ARFGAP/RecO"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001164">
        <property type="entry name" value="ArfGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038508">
        <property type="entry name" value="ArfGAP_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043593">
        <property type="entry name" value="ASAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004148">
        <property type="entry name" value="BAR_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037844">
        <property type="entry name" value="PH_ASAP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45854:SF4">
        <property type="entry name" value="ARF-GAP WITH SH3 DOMAIN, ANK REPEAT AND PH DOMAIN-CONTAINING PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45854">
        <property type="entry name" value="ASAP FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01412">
        <property type="entry name" value="ArfGap"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16746">
        <property type="entry name" value="BAR_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00169">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00018">
        <property type="entry name" value="SH3_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00405">
        <property type="entry name" value="REVINTRACTNG"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00105">
        <property type="entry name" value="ArfGap"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00233">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00326">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57863">
        <property type="entry name" value="ArfGap/RecO-like zinc finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103657">
        <property type="entry name" value="BAR/IMD domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50115">
        <property type="entry name" value="ARFGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7 8" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="2 9" id="KW-0728">SH3 domain</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="10" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="PH" evidence="13">
        <location>
            <begin position="131"/>
            <end position="223"/>
        </location>
    </feature>
    <feature type="domain" description="Arf-GAP" evidence="14">
        <location>
            <begin position="247"/>
            <end position="369"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="8">
        <location>
            <begin position="410"/>
            <end position="445"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="8">
        <location>
            <begin position="446"/>
            <end position="478"/>
        </location>
    </feature>
    <feature type="domain" description="SH3" evidence="12">
        <location>
            <begin position="735"/>
            <end position="798"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="114"/>
            <end position="138"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="526"/>
            <end position="547"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="657"/>
            <end position="685"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="114"/>
            <end position="126"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="11">
        <location>
            <begin position="663"/>
            <end position="680"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50115"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006874.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="798" mass="90295" checksum="81C4CA1BA4BECF58" modified="2019-05-08" version="1">MYLSNKQNLSPVGHQFEDRPTQQRLKYLLKVNEIKIKKGVDLLQNLIKYFHAQCNFFQDGLKAVESLKPSIETLSTDLHTIKQAQDEERKQLIHLRDILKSALQVEQKEDSQIRQSTAYSLHQPQGNKEHGTERSGSLYKKSDGIRKVWQKRKCSVKNGFLTISHGTANRPPAKLNLLTCQVKTNPEEKKCFDLISHDRTYHFQAEDEQECQIWTSVLQNSKEEALNNAFKGDENTGENNIVQELTKEIISEVQRMTGNDVCCDCGAQDPSWLSTNLGILTCIECSGIHRELGVHYSRMQSLTLDVLGTSELLLAKNIGNAGFNEIMEFCLPADETIKPNPSSDMNARKDYITAKYFERKYARKKHVDNAAKLYSLCEAVKARDIFGLVQVYADGVDLTEKIPLANGHEQDETALHLAVRSVDRTSLHIVDFLVQNSGNLDKQTYKGSTALHYCCLTDNVECLKLLLRGKASIEIANEAGETPLDIAKRLKHTHCEELLTQALSGRFNSHVHVEYEWRLLHEDLDESDDDVDEKLQQPSPNRREDRPVSFYQLGSNQLQPNVASLARDAANIAKDKQRSFMPSILQNETYGAILSGSPPPIQPAVPVTTSAPPLPPRNIGKVQFHCTLYFAFFIDILFPAPPPPVAKTASVIEALNQQSKQPPAVPQSKPIPPPLPPQPPSRISQKKSWSNKIYYLTHFVSLISGLGPTRLQLHCLSFVMFHYLYIHIFVLQSKTKPKRVKAIYNCVADNPDELTFSEGDIIVVDGEEDQEWWVSIFTCAFISLVFHSKYAHTFRVHW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXK5</accession>
    <name>A0A452GXK5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">ATP binding cassette subfamily C member 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>17beta-estradiol 17-O-(beta-D-glucuronate)(in) + ATP + H2O = 17beta-estradiol 17-O-(beta-D-glucuronate)(out) + ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:60128"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:82961"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="15">
            <dbReference type="Rhea" id="RHEA:60129"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>ATP + H2O + xenobioticSide 1 = ADP + phosphate + xenobioticSide 2.</text>
            <dbReference type="EC" id="7.6.2.2"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="14">
            <text>leukotriene C4(in) + ATP + H2O = leukotriene C4(out) + ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:38963"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:57973"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="14">
            <dbReference type="Rhea" id="RHEA:38964"/>
        </physiologicalReaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the ABC transporter superfamily. ABCC family. Conjugate transporter (TC 3.A.1.208) subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GXK5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007976.1">
        <property type="protein sequence ID" value="ENSGAGP00000006876.1"/>
        <property type="gene ID" value="ENSGAGG00000004641.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016324">
        <property type="term" value="C:apical plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008559">
        <property type="term" value="F:ABC-type xenobiotic transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006869">
        <property type="term" value="P:lipid transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd18595">
        <property type="entry name" value="ABC_6TM_MRP1_2_3_6_D1_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18603">
        <property type="entry name" value="ABC_6TM_MRP1_2_3_6_D2_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd03250">
        <property type="entry name" value="ABCC_MRP_domain1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd03244">
        <property type="entry name" value="ABCC_MRP_domain2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000293">
        <property type="entry name" value="ATP binding cassette subfamily C member 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1560.10:FF:000001">
        <property type="entry name" value="ATP-binding cassette subfamily C member 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1560.10:FF:000007">
        <property type="entry name" value="ATP-binding cassette subfamily C member 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000074">
        <property type="entry name" value="Multidrug resistance-associated protein 5 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1560.10">
        <property type="entry name" value="ABC transporter type 1, transmembrane domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003593">
        <property type="entry name" value="AAA+_ATPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011527">
        <property type="entry name" value="ABC1_TM_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036640">
        <property type="entry name" value="ABC1_TM_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003439">
        <property type="entry name" value="ABC_transporter-like_ATP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017871">
        <property type="entry name" value="ABC_transporter-like_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050173">
        <property type="entry name" value="ABC_transporter_C-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24223">
        <property type="entry name" value="ATP-BINDING CASSETTE SUB-FAMILY C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24223:SF176">
        <property type="entry name" value="ATP-BINDING CASSETTE SUB-FAMILY C MEMBER 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00664">
        <property type="entry name" value="ABC_membrane"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00005">
        <property type="entry name" value="ABC_tran"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00382">
        <property type="entry name" value="AAA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90123">
        <property type="entry name" value="ABC transporter transmembrane region"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50929">
        <property type="entry name" value="ABC_TM1F"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00211">
        <property type="entry name" value="ABC_TRANSPORTER_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50893">
        <property type="entry name" value="ABC_TRANSPORTER_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11" id="KW-0445">Lipid transport</keyword>
    <keyword evidence="12 16" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="9" id="KW-1278">Translocase</keyword>
    <keyword evidence="5 16" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="10 16" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="54"/>
            <end position="74"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="133"/>
            <end position="156"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="162"/>
            <end position="180"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="228"/>
            <end position="252"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="272"/>
            <end position="292"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="629"/>
            <end position="652"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="683"/>
            <end position="705"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="754"/>
            <end position="771"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="777"/>
            <end position="797"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="862"/>
            <end position="883"/>
        </location>
    </feature>
    <feature type="domain" description="ABC transmembrane type-1" evidence="18">
        <location>
            <begin position="24"/>
            <end position="289"/>
        </location>
    </feature>
    <feature type="domain" description="ABC transporter" evidence="17">
        <location>
            <begin position="321"/>
            <end position="547"/>
        </location>
    </feature>
    <feature type="domain" description="ABC transmembrane type-1" evidence="18">
        <location>
            <begin position="683"/>
            <end position="919"/>
        </location>
    </feature>
    <feature type="domain" description="ABC transporter" evidence="17">
        <location>
            <begin position="955"/>
            <end position="1189"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009726"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022967"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00034018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00047523"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00047576"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50893"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50929"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006876.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="21">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1200" mass="133151" checksum="C302C826F6AD0ABE" modified="2019-05-08" version="1">IGDYPKAWLAKALIRTFVGNLLKSVAFKVVHDLLVFVSPQLLKLLISFVADPTAYAWQGYLYALLLFVTALVQSLCLQQYFQLCFVLGMSLRTALMAAVYKKALTVSNATRKEYTQGETVNLMSADTQRFMDLATFIHQLWSAPLQIALSIIFLWWELGPSVLAGIGVMVLLIPINAVLMKNMKNKDQRMKIMNEILSGIKILKLFAWEPSFERQVGEIRVRELKGLLHFTNLQAFSIFIFSCAPFAVYVMVDEKNVLDAQKAFTSISLFNILRFPMATLPLVISSLVQTAVSTERLERYLGSGNLDPSNFSWLHGYCFYTLFTSLVAAQRCSESQLSRPSRVTLDIKPGHLVAVVGAVGSGKSSLVSAVLGEMENLKGHINIQGSVAYVPQQAWIQNATLKDNILFGSDLDEARYQQVVEACALLPDLKLLPGGDLTEIGEKGINLSGGQKQRVSLARAVYSDADIYLLDDPLSAVDAHVGKHLFDQVLGPDGLLQGKTRILVTHSVSFLPRTDEIVVLVAGAVSEQGSYSTLLANGGAFAQLLSTYGSQEGSTPEGEGWGLHHWRWGFPMPCIFHSTLFGLMLLCGEGRGVACRRSCVRGACGGWTPWDAATECAALQVKFSLYLRYLHAVGWGFSAWVFITYVAQYAAFMGSNLWLSDWTDDSLRYWNQTYPTSQRDMRIGVFGALAAFLLAGALLATRGAIRASRVLHQQLLSNILRVPMSFFDTTPTGRIVNRFAKDIFTVDETIPMSFRSWLTCFLGIISTLLMICLATPYFALIIIPLSILYFFLLRYYVSTSRQLRRLDSVTRSPIYSHFGETVSGLSVIRAYGHQERFLQHNDRIVDMNQKSVYSWIVSNRWLAVRLEFVGNLVVFSAALLAVITRDNLGSGIVGLSISSALNITQTLNWLVRMTSELETNIVAVERVREYTEVENEAQWVTAQRPPPGWPSKGEIQITDYQVRYRPGLELVLHGLTCDIGSTEKVGVVGRTGAGKSSLTNCLFRILEAAGGKIVIDGVDISTIGLHDLRQSLTIIPQDPVLFSGTLRMNLDPFDQHSDEEIWKALELAHLKTYVLDLPERLSHQVSEAGENLSVGQRQLLCLARALLRKSKILILDEATAAVDLETDHLIQTTIRSEFANCTVLTIAHRLHTIMDSNRVMVLQAGRIVEYDSPERLLQQQGLFSTMAKDAGIVSAQATVL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXK8</accession>
    <name>A0A452GXK8_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="17">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="15">Cytoplasmic vesicle</location>
            <location evidence="15">Secretory vesicle</location>
            <location evidence="15">Synaptic vesicle membrane</location>
            <topology evidence="15">Peripheral membrane protein</topology>
            <orientation evidence="15">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Endoplasmic reticulum</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Endosome membrane</location>
            <topology evidence="2">Peripheral membrane protein</topology>
            <orientation evidence="2">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="14">Melanosome membrane</location>
            <topology evidence="14">Peripheral membrane protein</topology>
            <orientation evidence="14">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="13">Postsynaptic density</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the dysbindin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GXK8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007853.1">
        <property type="protein sequence ID" value="ENSGAGP00000006765.1"/>
        <property type="gene ID" value="ENSGAGG00000005453.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:1904115">
        <property type="term" value="C:axon cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="GOC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031083">
        <property type="term" value="C:BLOC-1 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005783">
        <property type="term" value="C:endoplasmic reticulum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010008">
        <property type="term" value="C:endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033162">
        <property type="term" value="C:melanosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0014069">
        <property type="term" value="C:postsynaptic density"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030672">
        <property type="term" value="C:synaptic vesicle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048490">
        <property type="term" value="P:anterograde synaptic vesicle transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031175">
        <property type="term" value="P:neuron projection development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060155">
        <property type="term" value="P:platelet dense granule organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:2000300">
        <property type="term" value="P:regulation of synaptic vesicle exocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007531">
        <property type="entry name" value="Dysbindin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16294:SF5">
        <property type="entry name" value="DYSBINDIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16294">
        <property type="entry name" value="DYSTROBREVIN BINDING PROTEIN 1 DYSBINDIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04440">
        <property type="entry name" value="Dysbindin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="12" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="7" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="6" id="KW-0967">Endosome</keyword>
    <keyword evidence="10" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0770">Synapse</keyword>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1"/>
            <end position="26"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="127"/>
            <end position="197"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1"/>
            <end position="16"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="16">
        <location>
            <begin position="130"/>
            <end position="150"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="16">
        <location>
            <begin position="187"/>
            <end position="197"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00008686"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00034105"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00037798"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00037838"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006765.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="197" mass="21846" checksum="0B57B6D9C3073349" modified="2019-05-08" version="1">MSSPGATENNQRSSSELDAEHAQKVLDMEHTQQMKLKERQKFFEEAFQQDMEQYLSTGYLQIAERREPIGSMSSMEVNVDMLEQMDLMDMSDQEALDVFLNSGEDDSILSPILGPESKTCQNEISLQVPSQSELRQKLSSSSSTWTDSASQGTSEVGESPVIQSDEEEVQVDTALAAVNTERKGTSDDSDESDSQTV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9C3</accession>
    <name>A0A452H9C3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">C3H1-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9C3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013059.1">
        <property type="protein sequence ID" value="ENSGAGP00000011408.1"/>
        <property type="gene ID" value="ENSGAGG00000008803.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035198">
        <property type="term" value="F:miRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035196">
        <property type="term" value="P:miRNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010608">
        <property type="term" value="P:post-transcriptional regulation of gene expression"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.10:FF:000070">
        <property type="entry name" value="zinc finger CCCH domain-containing protein 7B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.10">
        <property type="entry name" value="Tetratricopeptide repeat domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011990">
        <property type="entry name" value="TPR-like_helical_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019734">
        <property type="entry name" value="TPR_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039691">
        <property type="entry name" value="ZC3H7A/B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000571">
        <property type="entry name" value="Znf_CCCH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036855">
        <property type="entry name" value="Znf_CCCH_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14928">
        <property type="entry name" value="MICRO-RNA BINDING ZINC FINGER CCCH DOMAIN-CONTAINING PROTEIN 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14928:SF6">
        <property type="entry name" value="ZINC FINGER CCCH DOMAIN-CONTAINING PROTEIN 7B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00642">
        <property type="entry name" value="zf-CCCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00028">
        <property type="entry name" value="TPR"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00356">
        <property type="entry name" value="ZnF_C3H1"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90229">
        <property type="entry name" value="CCCH zinc finger"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48452">
        <property type="entry name" value="TPR-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50005">
        <property type="entry name" value="TPR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50103">
        <property type="entry name" value="ZF_C3H1"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3 9" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="8" id="KW-0802">TPR repeat</keyword>
    <keyword evidence="6 9" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 9" id="KW-0863">Zinc-finger</keyword>
    <feature type="repeat" description="TPR" evidence="8">
        <location>
            <begin position="82"/>
            <end position="115"/>
        </location>
    </feature>
    <feature type="domain" description="C3H1-type" evidence="11">
        <location>
            <begin position="536"/>
            <end position="560"/>
        </location>
    </feature>
    <feature type="domain" description="C3H1-type" evidence="11">
        <location>
            <begin position="668"/>
            <end position="690"/>
        </location>
    </feature>
    <feature type="domain" description="C3H1-type" evidence="11">
        <location>
            <begin position="806"/>
            <end position="834"/>
        </location>
    </feature>
    <feature type="domain" description="C3H1-type" evidence="11">
        <location>
            <begin position="938"/>
            <end position="966"/>
        </location>
    </feature>
    <feature type="zinc finger region" description="C3H1-type" evidence="9">
        <location>
            <begin position="536"/>
            <end position="560"/>
        </location>
    </feature>
    <feature type="zinc finger region" description="C3H1-type" evidence="9">
        <location>
            <begin position="668"/>
            <end position="690"/>
        </location>
    </feature>
    <feature type="zinc finger region" description="C3H1-type" evidence="9">
        <location>
            <begin position="806"/>
            <end position="834"/>
        </location>
    </feature>
    <feature type="zinc finger region" description="C3H1-type" evidence="9">
        <location>
            <begin position="938"/>
            <end position="966"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="417"/>
            <end position="436"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00339"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50103"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011408.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1009" mass="113758" checksum="494C216A1E6D8221" modified="2019-05-08" version="1">MERQKRKEEIEKGLQFIQSTLPLSQEDYEAFIQKLVQNLFAEGNDLFREKDFKLSLVQYVEGLNVADYAASDEVTIPIELLCKLHVNRAACYFAMGLYEKALEDSEKALMLDKENIRALFRKARSLNELGRHKEAYECNSRCLLSLPHDESVTQLGQELAQKLGLRVRKAYKRPQELETFSLLSNGTSASLANQMTSNGLGSIDDIETDFSMDLQCIPAPLATSIPVSEGAVLSASDMDAKGLPTSLPAATLLHSPDCVSLPVPESVEDFTDGEVIGEELDSLLDSLADGSQYPLSLVQGTIPTNLPTEMPQLIPVFPGGTPLLPPVVTANIPINQHPLPPASFGLVMDPTKQLSSSVLDAFEPTGGSGGSPLDSLDSLDLLPYTEPRLDALDAFGTSRGSLDALDSFAVEEAGPQELRHTAGSQKPSPAISLSGVSHPTIPKGAELLQSQPEAVIPNTALLVKNPLASTHVFKQACHLCYPKTGPKAGDYTYREGLEHKCKRDILLGRLRNSEDKTWKRIRPRPTKTNFVGSYYLCKDMINKQDCKYGDNCTFAYHQEEIDVWTEERKGTLNRDLLFDPLGGIKRGNLTIAKLLQEHQGIFTFLCEICFDSKPRIISKGTKDSPAVCSNLAAKHSFYDNKCLVHIVRSTSLKYSKIRQFQEHFQFDVCRHEVRYGCLREDSCHFAHSFIELKVWLLQQYSGMTHEDIVQESKKYWQLMEVHASKATSGTAAPRAPTSSSFDLQMKFVCGQCWRNGQVVEPDKDLKYCSAKARHCWTKERRVLLVMSKAKRKWVSVRPLPSIRNFPQQYDLCIHAQNGRKCQYVGNCSFAHSPEERDMWTFMKENKILDMQQTYDMWLKKHNPGKPGEGTPLTSREGEKQIQMPTDYADIMMGYHCWLCGKNSNSKKQWQQHIQSEKHKEKVFTSDNDSSCWNYRFPMGEFRLCDRFQKTKACPEGDKCCCAHGQDELTEWLDRREVLKQKLAKARKDMLLCPKDDDFGKYNFLLRDGV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9C5</accession>
    <name>A0A452H9C5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3 16">Guanylate cyclase</fullName>
            <ecNumber evidence="3 16">4.6.1.2</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1 16">
            <text>GTP = 3',5'-cyclic GMP + diphosphate</text>
            <dbReference type="Rhea" id="RHEA:13665"/>
            <dbReference type="ChEBI" id="CHEBI:33019"/>
            <dbReference type="ChEBI" id="CHEBI:37565"/>
            <dbReference type="ChEBI" id="CHEBI:57746"/>
            <dbReference type="EC" id="4.6.1.2"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="15">Belongs to the adenylyl cyclase class-4/guanylyl cyclase family.</text>
    </comment>
    <dbReference type="EC" id="4.6.1.2" evidence="3 16"/>
    <dbReference type="AlphaFoldDB" id="A0A452H9C5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013011.1">
        <property type="protein sequence ID" value="ENSGAGP00000011366.1"/>
        <property type="gene ID" value="ENSGAGG00000002083.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004016">
        <property type="term" value="F:adenylate cyclase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005525">
        <property type="term" value="F:GTP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004383">
        <property type="term" value="F:guanylate cyclase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016941">
        <property type="term" value="F:natriuretic peptide receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017046">
        <property type="term" value="F:peptide hormone binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004672">
        <property type="term" value="F:protein kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035556">
        <property type="term" value="P:intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007168">
        <property type="term" value="P:receptor guanylyl cyclase signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07302">
        <property type="entry name" value="CHD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd14042">
        <property type="entry name" value="PK_GC-A_B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000371">
        <property type="entry name" value="Guanylate cyclase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:001106">
        <property type="entry name" value="Guanylate cyclase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.1230:FF:000004">
        <property type="entry name" value="Guanylate cyclase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.2300:FF:000101">
        <property type="entry name" value="Guanylate cyclase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.2300">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.1230">
        <property type="entry name" value="Nucleotide cyclase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001054">
        <property type="entry name" value="A/G_cyclase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018297">
        <property type="entry name" value="A/G_cyclase_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001828">
        <property type="entry name" value="ANF_lig-bd_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001170">
        <property type="entry name" value="ANPR/GUC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050401">
        <property type="entry name" value="Cyclic_nucleotide_synthase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029787">
        <property type="entry name" value="Nucleotide_cyclase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028082">
        <property type="entry name" value="Peripla_BP_I"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001245">
        <property type="entry name" value="Ser-Thr/Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11920:SF494">
        <property type="entry name" value="ATRIAL NATRIURETIC PEPTIDE RECEPTOR 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11920">
        <property type="entry name" value="GUANYLYL CYCLASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01094">
        <property type="entry name" value="ANF_receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00211">
        <property type="entry name" value="Guanylate_cyc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07714">
        <property type="entry name" value="PK_Tyr_Ser-Thr"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00255">
        <property type="entry name" value="NATPEPTIDER"/>
    </dbReference>
    <dbReference type="SMART" id="SM00044">
        <property type="entry name" value="CYCc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55073">
        <property type="entry name" value="Nucleotide cyclase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53822">
        <property type="entry name" value="Periplasmic binding protein-like I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00458">
        <property type="entry name" value="ANF_RECEPTORS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00452">
        <property type="entry name" value="GUANYLATE_CYCLASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50125">
        <property type="entry name" value="GUANYLATE_CYCLASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="14 16" id="KW-0141">cGMP biosynthesis</keyword>
    <keyword evidence="17" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9" id="KW-0342">GTP-binding</keyword>
    <keyword evidence="13 15" id="KW-0456">Lyase</keyword>
    <keyword evidence="10" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="11" id="KW-0675">Receptor</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="5" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8" id="KW-1133">Transmembrane helix</keyword>
    <feature type="domain" description="Protein kinase" evidence="19">
        <location>
            <begin position="539"/>
            <end position="808"/>
        </location>
    </feature>
    <feature type="domain" description="Guanylate cyclase" evidence="20">
        <location>
            <begin position="883"/>
            <end position="1013"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="30"/>
            <end position="55"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="17">
        <location>
            <begin position="820"/>
            <end position="847"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="18">
        <location>
            <begin position="43"/>
            <end position="55"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001436"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012202"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023293"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="RuleBase" id="RU000405"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="RuleBase" id="RU003431"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011366.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1068" mass="119565" checksum="F22E6CF01FBAAC82" modified="2019-05-08" version="1">MSADASPEPRDQWTVCRNACGRSTGAVCRPPKSWRPSHLNGSAGPGGGSQGQGCEGVGVGRGSLSLAVVLPERNVSYPWAWPRVGPAIRLALEALERGDPPLLPGPLAVRLHFRSSELEGACSEYVAPLNAVDLKLYHDPDVLFGPGCVYPAASVGRFASHWRLPLITAGAVATGFSHKQEHYGTTVRTGPSAPKLGAFVSQLHAHFNWSARAALLYADHKTDDRPYYFTIEGVYQELQEALNLSVRYHIYAPEEVTDASPVFPLTVIYVCGPLEMLHQIMRLAQRESLTNGDYVFFYVDVFGESLRASATRDALKPWQNKQSQDVGLRQAFQTVLVITYHEPQNPEYQHFQTQLILRAKEEFKVEALMSLSRGQMNLVAGCFYDGVLLYAMALNETLAEGGSKKDGTRIVQKMRDRKFQGVTGLVSMDSNNDRDTDFSLWVMGDPENGEYEVGHYAGAEKKLNWLGRPIQWVKGAPPPDNPPCVFDVDDPSCDKSGCDGPAPLPWAACILLGLSPEKLMLEKELASMLWRIRWDELQFGNAERYYKGAGSRLTLSLVRTPIPVHPGNVVAVKHVNKKRIELTRQVLFELKHMRDIQFNHLTRFIGACIDPPNVCIVTEYCPRGSLQDILENESINLDWMFRYSLINDIVKGMAFLHNSIIGHHGSLKSSNCVVDSRFVLKITDYGLASFRSSCDAEDSHALYAKKLWTAPELLQQGRLPLPGMQKADVYSFSIILQEIALRNGPFYIEGMDLSPKEIVQKVRNSQKPFFRPSIDMGLHSEELAVLMERCWAQEPAERPDFSQIKIFIRRFNKEGSTSILDNLLSRMEQYANNLEKLVEERTQAYLEEKRKAENLLYQILPHSVAEQLKRGETVRAEAFDSVTIYFSDIVGFTALSAESTPMQVVTLLNDLYTCFDAIIDNFDVYKVETIGDAYMVVSGLPVRNGKLHANEIVRMALALLEAVKTFKIRHRPNDQLRLRIGIHTGPVCAGVVGLKMPRYCLFGDTVNTASRMESNGQALKIHVSSTTKEVLDEFGCFELELRGDVEMKGKGKMRTYWLLGERTDDSII</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9C8</accession>
    <name>A0A452H9C8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Elongation factor 1-alpha</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="8">This protein promotes the GTP-dependent binding of aminoacyl-tRNA to the A-site of ribosomes during protein biosynthesis.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1 8">Belongs to the TRAFAC class translation factor GTPase superfamily. Classic translation factor GTPase family. EF-Tu/EF-1A subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9C8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013068.1">
        <property type="protein sequence ID" value="ENSGAGP00000011413.1"/>
        <property type="gene ID" value="ENSGAGG00000008805.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005525">
        <property type="term" value="F:GTP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003924">
        <property type="term" value="F:GTPase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003746">
        <property type="term" value="F:translation elongation factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="CDD" id="cd01883">
        <property type="entry name" value="EF1_alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd03693">
        <property type="entry name" value="EF1_alpha_II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd03705">
        <property type="entry name" value="EF1_alpha_III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.30.10:FF:000005">
        <property type="entry name" value="Elongation factor 1-alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000090">
        <property type="entry name" value="Elongation factor 1-alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.30.10:FF:000168">
        <property type="entry name" value="Elongation factor 1-alpha 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.30.10">
        <property type="entry name" value="Translation factors"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="HAMAP" id="MF_00118_A">
        <property type="entry name" value="EF_Tu_A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004161">
        <property type="entry name" value="EFTu-like_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031157">
        <property type="entry name" value="G_TR_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR054696">
        <property type="entry name" value="GTP-eEF1A_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000795">
        <property type="entry name" value="T_Tr_GTP-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050100">
        <property type="entry name" value="TRAFAC_GTPase_members"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009000">
        <property type="entry name" value="Transl_B-barrel_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009001">
        <property type="entry name" value="Transl_elong_EF1A/Init_IF2_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004539">
        <property type="entry name" value="Transl_elong_EF1A_euk/arc"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00483">
        <property type="entry name" value="EF-1_alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF008969">
        <property type="entry name" value="PRK12317.1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23115">
        <property type="entry name" value="TRANSLATION FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22594">
        <property type="entry name" value="GTP-eEF1A_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00009">
        <property type="entry name" value="GTP_EFTU"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03144">
        <property type="entry name" value="GTP_EFTU_D2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00315">
        <property type="entry name" value="ELONGATNFCT"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50465">
        <property type="entry name" value="EF-Tu/eEF-1alpha/eIF2-gamma C-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50447">
        <property type="entry name" value="Translation proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00301">
        <property type="entry name" value="G_TR_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51722">
        <property type="entry name" value="G_TR_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 8" id="KW-0251">Elongation factor</keyword>
    <keyword evidence="7 8" id="KW-0342">GTP-binding</keyword>
    <keyword evidence="2" id="KW-0488">Methylation</keyword>
    <keyword evidence="4 8" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="6" id="KW-0648">Protein biosynthesis</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Tr-type G" evidence="10">
        <location>
            <begin position="5"/>
            <end position="242"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="444"/>
            <end position="463"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007249"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022768"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022917"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="RuleBase" id="RU000325"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS51722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011413.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="463" mass="50524" checksum="3BF62DFD9DD2E120" modified="2019-05-08" version="1">MGKEKTHINIVVIGHVDSGKSTTTGHLIYKCGGIDKRTIEKFEKEAAEMGKGSFKYAWVLDKLKAERERGITIDISLWKFETTKYYITIIDAPGHRDFIKNMITGTSQADCAVLIVAAGVGEFEAGISKNGQTREHALLAYTLGVKQLIVGINKMDSTEPPYSEKRYDEIVKEVSAYIKKIGYNPATVPFVPISGWHGDNMLEPSPNMPWFKGWKVERKEGNASGVSLLEALDTILPPTRPTDKPLRLPLQDVYKIGGIGTVPVGRVETGILRPGMVVTFAPVNITTEVKSVEMHHEALSEALPGDNVGFNVKNVSVKDIRRGNVCGDSKSDPPQEAAQFTSQVIILNHPGQISAGYSPVIDCHTAHIACKFAELKEKIDRRSGKKLEDNPKSLKSGDAAIVEMIPGKPMCVESFSQYPPLGRFAVRDMRQTVAVGVIKNVEKKSGGAGKVTKSAQKAQKAGK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9C9</accession>
    <name>A0A452H9C9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Cathepsin O</fullName>
            <ecNumber evidence="13">3.4.22.42</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="12">Proteolytic enzyme possibly involved in normal cellular protein degradation and turnover.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>The recombinant human enzyme hydrolyzes synthetic endopeptidase substrates including Z-Phe-Arg-NHMec and Z-Arg-Arg-NHMec.</text>
            <dbReference type="EC" id="3.4.22.42"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Lysosome</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the peptidase C1 family.</text>
    </comment>
    <dbReference type="EC" id="3.4.22.42" evidence="13"/>
    <dbReference type="AlphaFoldDB" id="A0A452H9C9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013017.1">
        <property type="protein sequence ID" value="ENSGAGP00000011371.1"/>
        <property type="gene ID" value="ENSGAGG00000008781.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005764">
        <property type="term" value="C:lysosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008234">
        <property type="term" value="F:cysteine-type peptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd02248">
        <property type="entry name" value="Peptidase_C1A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.70.10:FF:000079">
        <property type="entry name" value="Cathepsin O"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.70.10">
        <property type="entry name" value="Cysteine proteinases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038765">
        <property type="entry name" value="Papain-like_cys_pep_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000169">
        <property type="entry name" value="Pept_cys_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025660">
        <property type="entry name" value="Pept_his_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013128">
        <property type="entry name" value="Peptidase_C1A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000668">
        <property type="entry name" value="Peptidase_C1A_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039417">
        <property type="entry name" value="Peptidase_C1A_papain-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12411">
        <property type="entry name" value="CYSTEINE PROTEASE FAMILY C1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00112">
        <property type="entry name" value="Peptidase_C1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00705">
        <property type="entry name" value="PAPAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00645">
        <property type="entry name" value="Pept_C1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54001">
        <property type="entry name" value="Cysteine proteinases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00139">
        <property type="entry name" value="THIOL_PROTEASE_CYS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00639">
        <property type="entry name" value="THIOL_PROTEASE_HIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="9" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="5" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="10" id="KW-0458">Lysosome</keyword>
    <keyword evidence="3" id="KW-0645">Protease</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="6" id="KW-0788">Thiol protease</keyword>
    <keyword evidence="7" id="KW-0865">Zymogen</keyword>
    <feature type="domain" description="Peptidase C1A papain C-terminal" evidence="15">
        <location>
            <begin position="67"/>
            <end position="279"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004371"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008455"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022807"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023145"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023228"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00051025"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00053492"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00066464"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00072046"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="SMART" id="SM00645"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011371.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="280" mass="31461" checksum="BFED6036E4593BC9" modified="2019-05-08" version="1">MCKWEWESIKRIRSLNSLSSDDNTTAFYGINQFSHFFPEDFKAIYLKSRPYSVPRYTEVLNRKETPLPAKFDWRDKNVVTQVRNQQTCGGCWAFSVVGGIESAYAIKMKILEELSVQQVIDCSYNNYGCHGGSTVSALSWLNQTQVKLVKDSEYAFKAQTGLCHYFQSSDIGVSITDYTAYDFSGQEDKMMKKLVNCGPLAVTVDAVSWQDYLGGIIQYHCSSGKANHAVLITGFDRTGSIPYWIVRNSWGRTWGINGYAHIKIGSNVCGIADTVSSVFV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9D0</accession>
    <name>A0A452H9D0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="19">Guanylate cyclase</fullName>
            <ecNumber evidence="19">4.6.1.2</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="17">Receptor for the C-type natriuretic peptide NPPC/CNP hormone. Has guanylate cyclase activity upon binding of its ligand. May play a role in the regulation of skeletal growth.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="1 19">
            <text>GTP = 3',5'-cyclic GMP + diphosphate</text>
            <dbReference type="Rhea" id="RHEA:13665"/>
            <dbReference type="ChEBI" id="CHEBI:33019"/>
            <dbReference type="ChEBI" id="CHEBI:37565"/>
            <dbReference type="ChEBI" id="CHEBI:57746"/>
            <dbReference type="EC" id="4.6.1.2"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="18">Belongs to the adenylyl cyclase class-4/guanylyl cyclase family.</text>
    </comment>
    <dbReference type="EC" id="4.6.1.2" evidence="19"/>
    <dbReference type="AlphaFoldDB" id="A0A452H9D0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013013.1">
        <property type="protein sequence ID" value="ENSGAGP00000011367.1"/>
        <property type="gene ID" value="ENSGAGG00000002083.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004016">
        <property type="term" value="F:adenylate cyclase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005525">
        <property type="term" value="F:GTP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004383">
        <property type="term" value="F:guanylate cyclase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016941">
        <property type="term" value="F:natriuretic peptide receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017046">
        <property type="term" value="F:peptide hormone binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004672">
        <property type="term" value="F:protein kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035556">
        <property type="term" value="P:intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001503">
        <property type="term" value="P:ossification"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007168">
        <property type="term" value="P:receptor guanylyl cyclase signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07302">
        <property type="entry name" value="CHD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd14042">
        <property type="entry name" value="PK_GC-A_B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000270">
        <property type="entry name" value="Guanylate cyclase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:001106">
        <property type="entry name" value="Guanylate cyclase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.1230:FF:000004">
        <property type="entry name" value="Guanylate cyclase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.2300:FF:000101">
        <property type="entry name" value="Guanylate cyclase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.2300">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.1230">
        <property type="entry name" value="Nucleotide cyclase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001054">
        <property type="entry name" value="A/G_cyclase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018297">
        <property type="entry name" value="A/G_cyclase_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001828">
        <property type="entry name" value="ANF_lig-bd_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001170">
        <property type="entry name" value="ANPR/GUC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050401">
        <property type="entry name" value="Cyclic_nucleotide_synthase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029787">
        <property type="entry name" value="Nucleotide_cyclase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028082">
        <property type="entry name" value="Peripla_BP_I"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001245">
        <property type="entry name" value="Ser-Thr/Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11920:SF494">
        <property type="entry name" value="ATRIAL NATRIURETIC PEPTIDE RECEPTOR 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11920">
        <property type="entry name" value="GUANYLYL CYCLASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01094">
        <property type="entry name" value="ANF_receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00211">
        <property type="entry name" value="Guanylate_cyc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07714">
        <property type="entry name" value="PK_Tyr_Ser-Thr"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00255">
        <property type="entry name" value="NATPEPTIDER"/>
    </dbReference>
    <dbReference type="SMART" id="SM00044">
        <property type="entry name" value="CYCc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55073">
        <property type="entry name" value="Nucleotide cyclase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53822">
        <property type="entry name" value="Periplasmic binding protein-like I"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00458">
        <property type="entry name" value="ANF_RECEPTORS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00452">
        <property type="entry name" value="GUANYLATE_CYCLASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50125">
        <property type="entry name" value="GUANYLATE_CYCLASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="16 19" id="KW-0141">cGMP biosynthesis</keyword>
    <keyword evidence="20" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="14" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10" id="KW-0342">GTP-binding</keyword>
    <keyword evidence="15 18" id="KW-0456">Lyase</keyword>
    <keyword evidence="11" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="8" id="KW-0892">Osteogenesis</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="13" id="KW-0675">Receptor</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="5" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9" id="KW-1133">Transmembrane helix</keyword>
    <feature type="domain" description="Protein kinase" evidence="21">
        <location>
            <begin position="515"/>
            <end position="788"/>
        </location>
    </feature>
    <feature type="domain" description="Guanylate cyclase" evidence="22">
        <location>
            <begin position="863"/>
            <end position="993"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="20">
        <location>
            <begin position="800"/>
            <end position="827"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001436"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022855"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023293"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00043880"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="RuleBase" id="RU000405"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="RuleBase" id="RU003431"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS50125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011367.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1048" mass="117449" checksum="3274B69FB3D16B2A" modified="2019-05-08" version="1">LPQPSMPAESPLPLPYALAAVLCRGPGANRSLSLAVVLPERNVSYPWAWPRVGPAIRLALEALERGDPPLLPGPLAVRLHFRSSELEGACSEYVAPLNAVDLKLYHDPDVLFGPGCVYPAASVGRFASHWRLPLITAGAVATGFSHKQEHYGTTVRTGPSAPKLGAFVSQLHAHFNWSARAALLYADHKTDDRPYYFTIEGVYQELQEALNLSVRYHIYAPDKLTHPTRVLQGTYPQRGRDLHKAGTLLAQRESLTNGDYVFFYVDVFGESLRASATRDALKPWQNKQSQDVGLRQAFQTVLVITYHEPQNPEYQHFQTQLILRAKEEFKVELGYSLMNLVAGCFYDGVLLYAMALNETLAEGGSKKDGTRIVQKMRDRKFQGVTGLVSMDSNNDRDTDFSLWVMGDPENGEYEVGAALSQGGSSVLRTVPAAEPPPRAGQCWEQDSPLVSPAPLSTLAIVALGTGLTLVMFGISSFLIFRKLMLEKELASMLWRIRWDELQFGNAERYYKGAGSRLTLSLRGSSYGSLMTAQGKYQIFANTGHFKGNVVAVKHVNKKRIELTRQVLFELKHMRDIQFNHLTRFIGACIDPPNVCIVTEYCPRGSLQDILENESINLDWMFRYSLINDIVKGMAFLHNSIIGHHGSLKSSNCVVDSRFVLKITDYGLASFRSSCDAEDSHALYAKKLWTAPELLQQGRLPLPGMQKADVYSFSIILQEIALRNGPFYIEGMDLSPKEIVQKVRNSQKPFFRPSIDMGLHSEELAVLMERCWAQEPAERPDFSQIKIFIRRFNKEGSTSILDNLLSRMEQYANNLEKLVEERTQAYLEEKRKAENLLYQILPHSVAEQLKRGETVRAEAFDSVTIYFSDIVGFTALSAESTPMQVVTLLNDLYTCFDAIIDNFDVYKVETIGDAYMVVSGLPVRNGKLHANEIVRMALALLEAVKTFKIRHRPNDQLRLRIGIHTGPVCAGVVGLKMPRYCLFGDTVNTASRMESNGQALKIHVSSTTKEVLDEFGCFELELRGDVEMKGKGKMRTYWLLGERTDDSII</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9D3</accession>
    <name>A0A452H9D3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">RING-type E3 ubiquitin transferase</fullName>
            <ecNumber evidence="4">2.3.2.27</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="25 26">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="26" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="25" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="25" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="27">
            <text>S-ubiquitinyl-[E2 ubiquitin-conjugating enzyme]-L-cysteine + [acceptor protein]-L-lysine = [E2 ubiquitin-conjugating enzyme]-L-cysteine + N(6)-ubiquitinyl-[acceptor protein]-L-lysine.</text>
            <dbReference type="EC" id="2.3.2.27"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="3">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="2.3.2.27" evidence="4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013023.1">
        <property type="protein sequence ID" value="ENSGAGP00000011376.1"/>
        <property type="gene ID" value="ENSGAGG00000008266.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000785">
        <property type="term" value="C:chromatin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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    <dbReference type="GO" id="GO:0031981">
        <property type="term" value="C:nuclear lumen"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061630">
        <property type="term" value="F:ubiquitin protein ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd19845">
        <property type="entry name" value="Bbox1_TIF1a_C-VI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd19828">
        <property type="entry name" value="Bbox2_TIF1a_C-VI"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05502">
        <property type="entry name" value="Bromo_tif1_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd15622">
        <property type="entry name" value="PHD_TIF1alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.40.10:FF:000123">
        <property type="entry name" value="E3 ubiquitin-protein ligase TRIM33"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.920.10:FF:000024">
        <property type="entry name" value="Transcription intermediary factor 1-alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000074">
        <property type="entry name" value="Tripartite motif containing 66"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.920.10">
        <property type="entry name" value="Bromodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003649">
        <property type="entry name" value="Bbox_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001487">
        <property type="entry name" value="Bromodomain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036427">
        <property type="entry name" value="Bromodomain-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018359">
        <property type="entry name" value="Bromodomain_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019786">
        <property type="entry name" value="Zinc_finger_PHD-type_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000315">
        <property type="entry name" value="Znf_B-box"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011011">
        <property type="entry name" value="Znf_FYVE_PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001965">
        <property type="entry name" value="Znf_PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019787">
        <property type="entry name" value="Znf_PHD-finger"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001841">
        <property type="entry name" value="Znf_RING"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017907">
        <property type="entry name" value="Znf_RING_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45915">
        <property type="entry name" value="TRANSCRIPTION INTERMEDIARY FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45915:SF4">
        <property type="entry name" value="TRANSCRIPTION INTERMEDIARY FACTOR 1-ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00439">
        <property type="entry name" value="Bromodomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00628">
        <property type="entry name" value="PHD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00643">
        <property type="entry name" value="zf-B_box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00502">
        <property type="entry name" value="BBC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00336">
        <property type="entry name" value="BBOX"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00297">
        <property type="entry name" value="BROMO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00249">
        <property type="entry name" value="PHD"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00184">
        <property type="entry name" value="RING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57845">
        <property type="entry name" value="B-box zinc-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47370">
        <property type="entry name" value="Bromodomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57903">
        <property type="entry name" value="FYVE/PHD zinc finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57850">
        <property type="entry name" value="RING/U-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00633">
        <property type="entry name" value="BROMODOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="BROMODOMAIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="ZF_BBOX"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01359">
        <property type="entry name" value="ZF_PHD_1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="ZF_PHD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00518">
        <property type="entry name" value="ZF_RING_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50089">
        <property type="entry name" value="ZF_RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="14 19" id="KW-0103">Bromodomain</keyword>
    <keyword evidence="13" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="15" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="17" id="KW-0539">Nucleus</keyword>
    <keyword evidence="26" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0678">Repressor</keyword>
    <keyword evidence="16" id="KW-0804">Transcription</keyword>
    <keyword evidence="12" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="6" id="KW-0808">Transferase</keyword>
    <keyword evidence="10" id="KW-0833">Ubl conjugation pathway</keyword>
    <keyword evidence="11" id="KW-0862">Zinc</keyword>
    <keyword evidence="9 18" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="RING-type" evidence="23">
        <location>
            <begin position="63"/>
            <end position="117"/>
        </location>
    </feature>
    <feature type="domain" description="B box-type" evidence="24">
        <location>
            <begin position="144"/>
            <end position="197"/>
        </location>
    </feature>
    <feature type="domain" description="B box-type" evidence="24">
        <location>
            <begin position="204"/>
            <end position="245"/>
        </location>
    </feature>
    <feature type="domain" description="PHD-type" evidence="22">
        <location>
            <begin position="712"/>
            <end position="759"/>
        </location>
    </feature>
    <feature type="domain" description="Bromo" evidence="21">
        <location>
            <begin position="818"/>
            <end position="873"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="32"/>
            <end position="51"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="521"/>
            <end position="542"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="664"/>
            <end position="705"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="902"/>
            <end position="936"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="20">
        <location>
            <begin position="521"/>
            <end position="540"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="20">
        <location>
            <begin position="664"/>
            <end position="673"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="20">
        <location>
            <begin position="910"/>
            <end position="919"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00012483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022491"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
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        </source>
    </evidence>
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        </source>
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        </source>
    </evidence>
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    </evidence>
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        </source>
    </evidence>
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        <source>
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        </source>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        <source>
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    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="936" mass="106294" checksum="52A387544ADB8E4D" modified="2019-05-08" version="1">MLLEVFKKATLYAETTEPKPPKKNINLLLVASDSDDENEHAESRQGPAERGGEAAQLNLLDTCAVCGQHIQSRRPKLLPCLHSFCLRCLPQPDRYLMLPPAAPSPSPVGVIRCPVCSQECAERHIIDNFFVKDTTEVPSSTVEKSNQVCTSCEDNAEANGFCVECVEWLCKTCIRAHQRVKFTKDHTVRQKEEVSPEAVGVNSQRPVFCPYHKKEQLKLYCETCDKLTCRDCQLLEHKEHRYQFIEEAFQNQKVIIDTLITKLMEKTKYIKYTGKQIQNRILEANQNQKQVEQDIKVAIFTLMVEINKKGKALLHQLESLSKDHRIKLLQQQQEVASLSKQLEHVMNFSKWAVSSGSSTALLYSKRLITYRLRYLLRARCDASPVTNTTVQFHCDPSFWAQNIFNLGSLVIEDKETPPHMPKSPVMEPNLQPQSGLPSNQLSKFPTQISLAQLRLQHMQQQVMAQRQQAQRRVGPVGLPNSRILGPMQHTPTSHQVNYHCSREKTCLIVLLPRQLQXSAASSTASSTTSTPSSPTVTSAAGYDGKTFGPPMVDLSSPVGSSYTLPSLPDVRVTQAMVYSFYFNVNLLSGSSSRPPGADSTHRIPVVMLEPIRIKQESGTSHENYDFPIVIVKQETDEESRSRNTNFSRSILTSLLLDSHHNTTSDKTVLRTDAPDSTGDRPGILQENTSSGKTAWMGTPHIGDIRKEDDPNEDWCAVCQNGGELLCCEKCPKVFHLSCHVPSLMNFPRGEWICTFCRDLSKPEVEYDCDNPNQSSEKRKLEGYMGLAPIDRRKCERLLLYLYCHEMSLAFQDPVPPTVPEYYKIIKKPMDLSTIKKRLQVNHPLYMKPEDFVADFRLIFQNCAEFNEPDSEVADAGMKLEAYFEELLKTLYPEREFPIQPYCRSERENPEFSDDSDDDFVQPRKKRLKGEERQLLK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9D4</accession>
    <name>A0A452H9D4_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="3">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the PPDPF family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9D4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013075.1">
        <property type="protein sequence ID" value="ENSGAGP00000011420.1"/>
        <property type="gene ID" value="ENSGAGG00000008809.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026754">
        <property type="entry name" value="PPDPF"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14572">
        <property type="entry name" value="PANCREATIC PROGENITOR CELL DIFFERENTIATION AND PROLIFERATION FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15060">
        <property type="entry name" value="PPDFL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR02071">
        <property type="entry name" value="PPDPFACTOR"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="22"/>
            <end position="44"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="23"/>
            <end position="32"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006609"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011420.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="114" mass="12120" checksum="8D2094516944D52E" modified="2019-05-08" version="1">MAAIPSSGSLVATHDYYRRRLGSTSSNSSCGSAEYSGEVIPHHPGLPKSDPGHWWASFFFGKPTHPIMTTVSESPENSGTFQVANGMITCGLAQEVMRKRHASEPSKQSAGPSA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9D7</accession>
    <name>A0A452H9D7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Anoctamin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="8">Membrane</location>
            <topology evidence="8">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 8">Belongs to the anoctamin family.</text>
    </comment>
    <comment type="caution">
        <text evidence="8">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9D7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013029.1">
        <property type="protein sequence ID" value="ENSGAGP00000011381.1"/>
        <property type="gene ID" value="ENSGAGG00000007328.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005229">
        <property type="term" value="F:intracellularly calcium-gated chloride channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046983">
        <property type="term" value="F:protein dimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032394">
        <property type="entry name" value="Anoct_dimer"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007632">
        <property type="entry name" value="Anoctamin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049452">
        <property type="entry name" value="Anoctamin_TM"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12308">
        <property type="entry name" value="ANOCTAMIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12308:SF28">
        <property type="entry name" value="ANOCTAMIN-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16178">
        <property type="entry name" value="Anoct_dimer"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04547">
        <property type="entry name" value="Anoctamin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6 8" id="KW-0472">Membrane</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 8" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 8" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="315"/>
            <end position="341"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="391"/>
            <end position="410"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="467"/>
            <end position="491"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="511"/>
            <end position="530"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="551"/>
            <end position="572"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="671"/>
            <end position="693"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="721"/>
            <end position="745"/>
        </location>
    </feature>
    <feature type="domain" description="Anoctamin dimerisation" evidence="12">
        <location>
            <begin position="76"/>
            <end position="301"/>
        </location>
    </feature>
    <feature type="domain" description="Anoctamin transmembrane" evidence="11">
        <location>
            <begin position="304"/>
            <end position="874"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="36"/>
            <end position="65"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="873"/>
            <end position="900"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009671"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="RuleBase" id="RU280814"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF04547"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF16178"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011381.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="909" mass="106429" checksum="D4F061C9CEADFE24" modified="2019-05-08" version="1">VHSNNDPRFPTVPFIFHPVAKDVNIYFDELEGVNSPSKDDDSLLHHGNLTSTSDDASRLETVGEGVPDKNKSNGLYFRDGKCRIDYILVYRKSSPQTEKREIFERNIRAEGLQMEKESSLTNSDIIFVKLHAPWEVLGRYAEVMNVRMPFRRKIYYLHRRHKFMNRIERQISRFQGWLPRKPMRLDKETLPDLEENDCYTAPFSQQRIHHFIIHNKDTFFNNATRSRIVHHILQRVKYEEGKNKIGLNRLLTNGSYEAAFPLHEGSYRSKNSIRTHGAENHRHLLYECWASWGMWYKYQPLDLVRRYFGEKIGLYFAWLGWYTGMLFPAAFIGLFVFLYGVTTLNHCQVSKEVCQATDIIMCPICDKFCPFMRLSDSCVYAKVTHLFDNGATVFFAVFMAVWATVFLEFWKRRRAVIAYDWDLIDWEEEEEEIRPQFEAKYSKKERMNPISGKPEPYQAFADKCSRLIVSASGIFFMICVVIAAVFGIVIYRVVTVSTFAAFKWALIRNNSQVATTGTAVCINFCIIMLLNVVKQPRTESEWENSFTLKMFLFQFVNLNSSTFYIAFFLGRFTGHPGAYLRLINRWRLEECHPSGCLIDLCMQMGIIMVLKQTWNNFMELGYPLIQNWWTRRKLYQEHGSQRKTSFPQWEKDYNLQPMNAYGLFDEYLEMILQFGFTTIFVAAFPLAPLLALLNNIIEIRLDAYKFVTQWRRPLASRAKDIGIWYGILEGIGILSVITNAFVVAVTSDFIPRLVYAYKYGPCAGQGEAGQKCMVGYVNASLSVFLVSDFENRSEPTSNGSEFSGSPLKYCRYRDYRDPPHSPVPYGYTLQFWHVLAARLAFIIVFEHLVFCIKHLISYLIPDLPKDLRDRMRREKYLIQEMMYEAELERLQKERKERKKNGKSHHNEWP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9E2</accession>
    <name>A0A452H9E2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Tyrosine-protein kinase</fullName>
            <ecNumber evidence="13">2.7.10.2</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="9 13">
            <text>L-tyrosyl-[protein] + ATP = O-phospho-L-tyrosyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10596"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.10.2"/>
        </reaction>
    </comment>
    <comment type="similarity">
        <text evidence="13">Belongs to the protein kinase superfamily. Tyr protein kinase family.</text>
    </comment>
    <dbReference type="EC" id="2.7.10.2" evidence="13"/>
    <dbReference type="AlphaFoldDB" id="A0A452H9E2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013087.1">
        <property type="protein sequence ID" value="ENSGAGP00000011431.1"/>
        <property type="gene ID" value="ENSGAGG00000008815.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004715">
        <property type="term" value="F:non-membrane spanning protein tyrosine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000399">
        <property type="entry name" value="Tyrosine-protein kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000053">
        <property type="entry name" value="Tyrosine-protein kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.505.10:FF:000074">
        <property type="entry name" value="Tyrosine-protein kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.505.10">
        <property type="entry name" value="SH2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050198">
        <property type="entry name" value="Non-receptor_tyrosine_kinases"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001245">
        <property type="entry name" value="Ser-Thr/Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000980">
        <property type="entry name" value="SH2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036860">
        <property type="entry name" value="SH2_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008266">
        <property type="entry name" value="Tyr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020635">
        <property type="entry name" value="Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24418">
        <property type="entry name" value="TYROSINE-PROTEIN KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07714">
        <property type="entry name" value="PK_Tyr_Ser-Thr"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00017">
        <property type="entry name" value="SH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00018">
        <property type="entry name" value="SH3_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00401">
        <property type="entry name" value="SH2DOMAIN"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00109">
        <property type="entry name" value="TYRKINASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00252">
        <property type="entry name" value="SH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00326">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00219">
        <property type="entry name" value="TyrKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55550">
        <property type="entry name" value="SH2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00109">
        <property type="entry name" value="PROTEIN_KINASE_TYR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50001">
        <property type="entry name" value="SH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6 12 13" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="5 13" id="KW-0418">Kinase</keyword>
    <keyword evidence="4 12 13" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7 10" id="KW-0727">SH2 domain</keyword>
    <keyword evidence="1 11" id="KW-0728">SH3 domain</keyword>
    <keyword evidence="3 13" id="KW-0808">Transferase</keyword>
    <keyword evidence="8 13" id="KW-0829">Tyrosine-protein kinase</keyword>
    <feature type="domain" description="SH3" evidence="15">
        <location>
            <begin position="13"/>
            <end position="75"/>
        </location>
    </feature>
    <feature type="domain" description="SH2" evidence="14">
        <location>
            <begin position="78"/>
            <end position="170"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="16">
        <location>
            <begin position="187"/>
            <end position="442"/>
        </location>
    </feature>
    <feature type="binding site" evidence="12">
        <location>
            <position position="215"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022999"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023137"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00051245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00191"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="RuleBase" id="RU362096"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50001"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011431.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="448" mass="51725" checksum="DBDDBD9CDC467803" modified="2019-05-08" version="1">SAPLPGSMNSRLGTSPLYTSLWDFDARTQTEISFQAGDLFQVIRQEGEWWWARKVDGSNRILGDSISTHIYFLSSSSWFFGQISRSEALHRLLSEKNKTGAFLIRISEKKDTDYVLSVRDDSIVRHYKIWRNSQGNLYMNAALSFPDLLSLVEHYKAKNLTHGLRLTIPCWKPLPHWDDWERPREEFSFIKKLGAGYFGEVYEGYWKKKVKVAIKTIPKADLTYQATFKNEIEVMKNLRHKHILSLYAISSVGDPVYIITELMLKGNLLDFLRGKSEREHLEMTDLVDMASQVADGMCYLESQNFVHRDLAARNILVGENNICKVGDFGLARLIKDDVYLSYSRNIPYKWTAPEAISHGRYSTKSDVWSFGILLYEIITYGQIPYPGMSNSEVCQKVQTGFQMSCPPKCPPMLYEIMRKCWHLSPDQRPDFQHLKGKLHSFTHYENPG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9E3</accession>
    <name>A0A452H9E3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">NPC intracellular cholesterol transporter 2</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="7">Epididymal secretory protein E1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the NPC2 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9E3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000012832.1">
        <property type="protein sequence ID" value="ENSGAGP00000011205.1"/>
        <property type="gene ID" value="ENSGAGG00000008655.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015485">
        <property type="term" value="F:cholesterol binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033344">
        <property type="term" value="P:cholesterol efflux"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032367">
        <property type="term" value="P:intracellular cholesterol transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00916">
        <property type="entry name" value="Npc2_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.770:FF:000001">
        <property type="entry name" value="NPC intracellular cholesterol transporter 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.770">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014756">
        <property type="entry name" value="Ig_E-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003172">
        <property type="entry name" value="ML_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033916">
        <property type="entry name" value="ML_Npc2-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039670">
        <property type="entry name" value="NPC2-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11306">
        <property type="entry name" value="NIEMANN PICK TYPE C2 PROTEIN NPC2-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11306:SF68">
        <property type="entry name" value="NPC INTRACELLULAR CHOLESTEROL TRANSPORTER 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02221">
        <property type="entry name" value="E1_DerP2_DerF2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00737">
        <property type="entry name" value="ML"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81296">
        <property type="entry name" value="E set domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0964">Secreted</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <feature type="domain" description="MD-2-related lipid-recognition" evidence="8">
        <location>
            <begin position="24"/>
            <end position="145"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006370"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00021477"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00032516"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="SMART" id="SM00737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011205.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="149" mass="16382" checksum="25872CBAA2155716" modified="2019-05-08" version="1">SLQLTQYFLLPTALSSGPLLEFLFVLAGSKDGSITEVNVSPCPTQPCLLQKGTSYSVNVTFSSKIDSQGSKAKVYGEILHVDVPFPIPEPDGCKSGIQCPIEKGHSYSYLNKLPVKSEYPSIKLIVQWELIDDQGQMIFCWKIPVQITS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9F0</accession>
    <name>A0A452H9F0_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="8">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452H9F0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013041.1">
        <property type="protein sequence ID" value="ENSGAGP00000011391.1"/>
        <property type="gene ID" value="ENSGAGG00000008776.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004386">
        <property type="term" value="F:helicase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016787">
        <property type="term" value="F:hydrolase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd17979">
        <property type="entry name" value="DEXHc_DHX34"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18791">
        <property type="entry name" value="SF2_C_RHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.1080:FF:000013">
        <property type="entry name" value="ATP-dependent RNA helicase DHX34"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000540">
        <property type="entry name" value="probable ATP-dependent RNA helicase DHX34"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000725">
        <property type="entry name" value="probable ATP-dependent RNA helicase DHX34"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.1080">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011545">
        <property type="entry name" value="DEAD/DEAH_box_helicase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048333">
        <property type="entry name" value="HA2_WH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007502">
        <property type="entry name" value="Helicase-assoc_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014001">
        <property type="entry name" value="Helicase_ATP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001650">
        <property type="entry name" value="Helicase_C-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18934">
        <property type="entry name" value="ATP-DEPENDENT RNA HELICASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18934:SF221">
        <property type="entry name" value="ATP-DEPENDENT RNA HELICASE DHX34-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00270">
        <property type="entry name" value="DEAD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21010">
        <property type="entry name" value="HA2_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00271">
        <property type="entry name" value="Helicase_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04408">
        <property type="entry name" value="WHD_HA2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00487">
        <property type="entry name" value="DEXDc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00847">
        <property type="entry name" value="HA2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00490">
        <property type="entry name" value="HELICc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51192">
        <property type="entry name" value="HELICASE_ATP_BIND_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51194">
        <property type="entry name" value="HELICASE_CTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="3" id="KW-0347">Helicase</keyword>
    <keyword evidence="2" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="1" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Helicase ATP-binding" evidence="6">
        <location>
            <begin position="169"/>
            <end position="329"/>
        </location>
    </feature>
    <feature type="domain" description="Helicase C-terminal" evidence="7">
        <location>
            <begin position="367"/>
            <end position="535"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="720"/>
            <end position="829"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="720"/>
            <end position="732"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="5">
        <location>
            <begin position="796"/>
            <end position="811"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022806"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS51192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS51194"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011391.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="829" mass="91751" checksum="2B5569C5DCCA2D51" modified="2019-05-08" version="1">WEDPSPSRGHQGGRWDWSCPEMRRRLEELYFREQDYIRAGSEDSRQFWAFFERLQRFQTLKAEGELAPRAPPARADPSAALGLAGLPQEYDPRYRINLSVLSGDVEGLAQGRSGGPRRGPGVPAERLSQFRAALLHYLDFAQKQSFAKLAKLQRERAALPIAQYRDRLLRAVAQNQVVVVAGDTGCGKSTQVPQYLLAAGYSHVACTQPRRIACISLAKRVGFESLHQYGAEVGYQIRFESTRSLATKIVFLTEGLLLRQVQRELALPGYQVLIADEVHERHLHSDFLLGVLRRLLPSRPDLKLVLMSATINIRLFAGYFGGAPVVQVPGRLFPITVVYQPIPQEEAAASGKPERLDPRPYLRVLQAIDHKYPPEERGDLLVFLSGVAEIGAVLEAAQAYATHTQRWVVLPLHSTLSLAEQDKVFDMPPSGVRKCIVSTNIAETSVTIDGVRFVLDSGKVKELSYDPQAKLQRLQEFWISRASAEQRKGRAGRTGPGVCYRLYAESDYDAFAPYPVPEIQRVALDALVLQMKSMGLGDPRTFPFLEPPPQSSLEMAVRYLREQGALDEAESLTPIGSLLAQLPVDVVIGKMLVLGALFGLAGPVLTVAAALSVQSPFLRPVRANPDCATARRPLESPHGDPLTLLNTFNEWVQVKSEWAGGSRKWCRRRGLEEHRLYEAANLRRQFQELLRDHGLLEAVQGPRDSYARQSRHRVRQELHRLQRRHEETEGRQRKVLKLQGGEAGSSSDEGEGARSAGPPGVDIQVRGRGPQCGAEGPPRLTAHWGQSQHQEKEPGRGVGSSGLEWGSGGWEPGLLGSDPALGGEWGPAG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9F5</accession>
    <name>A0A452H9F5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">LRRCT domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452H9F5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013042.1">
        <property type="protein sequence ID" value="ENSGAGP00000011392.1"/>
        <property type="gene ID" value="ENSGAGG00000008796.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000770">
        <property type="entry name" value="Uncharacterized protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000483">
        <property type="entry name" value="Cys-rich_flank_reg_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001611">
        <property type="entry name" value="Leu-rich_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025875">
        <property type="entry name" value="Leu-rich_rpt_4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003591">
        <property type="entry name" value="Leu-rich_rpt_typical-subtyp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050541">
        <property type="entry name" value="LRR_TM_domain-containing"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000372">
        <property type="entry name" value="LRRNT"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24369">
        <property type="entry name" value="ANTIGEN BSP, PUTATIVE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24369:SF210">
        <property type="entry name" value="CHAOPTIN-RELATED"/>
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    </dbReference>
    <dbReference type="Pfam" id="PF12799">
        <property type="entry name" value="LRR_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13855">
        <property type="entry name" value="LRR_8"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00364">
        <property type="entry name" value="LRR_BAC"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SMART" id="SM00369">
        <property type="entry name" value="LRR_TYP"/>
        <property type="match status" value="10"/>
    </dbReference>
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        <property type="entry name" value="LRRCT"/>
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    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="L domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51450">
        <property type="entry name" value="LRR"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="1" id="KW-0433">Leucine-rich repeat</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="2 5" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="5">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5018973556" description="LRRCT domain-containing protein" evidence="5">
        <location>
            <begin position="21"/>
            <end position="536"/>
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        <location>
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    </feature>
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            <begin position="303"/>
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    </feature>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
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            <dbReference type="SAM" id="SignalP"/>
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    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="SMART" id="SM00013"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="SMART" id="SM00082"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011392.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="10">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="536" mass="59380" checksum="57FAE38361103A52" modified="2019-05-08" version="1">MSRLWIMVLVLLWIPVMAIAQLPGNAQCPVNCSCFFSQWFVRCSNASLSSLPPGIPNATVELDLQHNQFSTLSAGFFPELAEISIIYLGSSGIHQVEPGAFQGVKNLYHLHLDNNLLEQVPEGVFENLTNLIFLHLQHNQIAHLLPGGFSSLKQLSVLDLSNNLLLELSDQALHGLPRLRRLYLSANRITNVSSKALPGSLRALSLDENQLTSVPAAIRISPMLSTLQLSGNPIRKLTSLSFGRRLRSLRQLFLDGLALEQITNLAFTRLRWLELLSLRNNSLESLPPLSSLKSLSTLYLTGNKWRCDCNLIWLRTWQKKVLRKERSPVECSSPEVLQGQLLVDIELQKLTCPPFGMDSTTLSPSENTNTPMAIAPPRDIPLPGAATTATTTTSTTLITTKRLPSSTTHSATSLHHGLERWDPCLADHISSVHIRTKGNTSLVVSWAFSGDQDQFEGVHYLGQRRGCPKTSLSLSMNTFFQSRHLPKSVGGACRTLENTDRTLLCSPLCQGYLHPSFQTTGHWPGPHISFPLWNSI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9G3</accession>
    <name>A0A452H9G3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Microtubule-associated protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 7">Cytoplasm</location>
            <location evidence="1 7">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000012858.1">
        <property type="protein sequence ID" value="ENSGAGP00000011226.1"/>
        <property type="gene ID" value="ENSGAGG00000008671.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005874">
        <property type="term" value="C:microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043005">
        <property type="term" value="C:neuron projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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        <property type="term" value="F:microtubule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
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    <dbReference type="GO" id="GO:0000226">
        <property type="term" value="P:microtubule cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="project" value="TreeGrafter"/>
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        <property type="entry name" value="MAP2/MAP4/Tau"/>
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    <dbReference type="InterPro" id="IPR001084">
        <property type="entry name" value="MAP_tubulin-bd_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11501">
        <property type="entry name" value="MICROTUBULE-ASSOCIATED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11501:SF16">
        <property type="entry name" value="MICROTUBULE-ASSOCIATED PROTEIN 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00418">
        <property type="entry name" value="Tubulin-binding"/>
        <property type="match status" value="3"/>
    </dbReference>
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    </dbReference>
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    <proteinExistence type="predicted"/>
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    <keyword evidence="4 7" id="KW-0493">Microtubule</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
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    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU000686"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011226.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2403" mass="260449" checksum="4A6DC88621B8AB28" modified="2019-05-08" version="1">MADFDHNLSLADALTEPPAEIEEEVKRDFIATLEAEKFDDVVGETVGKTDYIPLLDDDDDAKAGSQEPKSKPHTDGGQVESTSAVGPAVLENGDHGTEDDSTVFPREITDEKMSYKEFLDRNETWAVDDRDLCFEPQSIFRPMEGTEPFRMHREDVLCDLLLRPSEMETRLPFTEHFEASEEVHAPHAAMMVPELPSLGSPYSPAEVIDPSAFITLDSTTESLLNIEAPARVTVPEENWLGAEHAVEELDESSFVEPPEPTRITDAAEKYLPGSPVAAAPAAVPPALTEPTGETKATDTPQVEPTESVPVMGDESVPIVEELMVFEPSVEQHKSAVNEPPTVLSAPTVVMEQKMTVTEASTLGADSTPVLENVEENKLSPSKHTEHLLKPNEELPELAVPVEAKEPLTLIEIKETLPEKTAPAADLTIVETLKEEAELSHAHHTEPPQKKSPLEPTGVPTAQVRQANKSSDRRFGRAKPAAVPGADVPEELLVGLPEQKSTDPKADPFSMAELGWVSGTFSRTRAPHKKAAGQLLSMPPEFVESQRDVPRESWDSEGSPVIVKKKKKKQKQKRNQQPRTMELGDENVEVSKAPKVPLFAAELQKPDVPFVLPAEGVKEHSAHSREGQRKGTSNVLRDAKAGTESHLVDDQNFISSPTAGQQSLKTEGPFKFLLDAKAGEVMKQEEMKADSLFQSKSKNKQVPMEKLECKAEHTKMGDPSSALTQTKPIEISFIDKNKKIECEHSDLEVSLSKGINLNKVETPLETKPVEINLIDENKEIKCTSHKHVAADETVPSEVQTPGGILMGEKPKKRSSDEKSRKAENGFSKQPAVLEIKTDTTNLPAAAKRVNKTKETDSFDKSQETGSITSEPPLETATDITKVPLIALVSDKPKEGIAGQNKKADFSFSEQPFVLETKTDTAKLHAPAETVDITNIGFTDKSKQIGFDHPAVTDPSMALVTDKPKKRGGDGKNKKVKNSSEQPVLMEVEIDPDKLPAVVKTAGKTKEIIFTDEDKEPGVAIAEHSLEDMTGPSTAKVAEKPKKRTSDGKNKKAEKGYFEQPFFLEAKEDTSSLPTVMEKDDKTKKVSFIDKGKETDFTTADHLLEGATATMEVQGPITTLVTEEPKNGITGKSEKAEFSISEQPDVAKLPASAETISKTKEDSLMDKSKETGFTTDSSRALMSDKPKEKGSDRKGKKVGKSSFEQPFLLETKIDTSKLPTVVEKAEKMKEMGFADIGKEPIFTTTEHQLENLTDTTKVQVPTMKLITEKPKIKESNEKSFFEQLVPSDYKMDTAQSETLTKTKETPFSDKGKETDFTTLEHLLEDIRDTAKAHIPMTELVADKPKKRCSDGKSKKIENTAEHLAVLEAKTDSIKLPAVLKEANKIKETHSTDKGQETQFTISEHLLENITDTAKIQTPITPLVTGKPEETNKGNSTKADVHLEKPFLLETKVDAATLPGGADEIVDKTEAISSTDESQVAGFTILEHQTMTDPSPALVTNKSKKRGNDGKNREAKNASEQPVLLETKTDKSKIQPPIVTDLEYQMEEMGLVDENQNIKNFPSKHQMLWDNKGHLFTPFTQSGAVGVDRVGKASIFPEYPVKDEGWKVPVPPAVVLEGTNKESSTSDKREENKHSHYEHSVTLEGEDAGVQAFTSVKEGDKIKVTSAGDKKKKGKWPSLDSPVKQDAKAGRDEVHTSILAEIDDKEISTECASLEHLVKETDAMKRTIPTDAKVTDKMEGSSSAGEKDVGCISPELTVQWENKAEAVVLQVLATAEVVNKAKEAELGEQSFSECPVSLEKKTDAAEVASVTLRDAQTEEISPTDKIKSHSEHSEGDAADKMKAGLKDFQALKSEEDKYADLPQKQIDGSGQKVVKETKKEERVKATEQIKGYMRPTKSRGLPAPPPRPAVQAREKPRQLKANGMSRQRQEKAKPEEIKPVELVTGSDITAPPNKELPPSPEKKSKPSVSTPSAKPAATKTKPLSTTSPKRPASATPGQNKKPTSPTAGPTSATTPKRSATSTTRPSTLTPKDTKPKVTDAKSPDKRTSLSKPLSATTPRAAVKVSPATPRTTAASPITTASGPRSTATSPPKRPTSIKTETKLADARKTTAKSPSADLSRPKSAPANSAKSSATTPTATTPGTLASPGVATSRPKPKPAAARPTTASSATPEAKKPSTVKAPLKTSTVPKPPRPTSSVSAPDLKNIRSKIGSTDNIKHQPGGGRAKVEKKPESAGAARKSEPNAVSKMATTKTTVTKEGAQKQPNGKVQIQSKKVDISKVSSKCGSKTNIKHKPGGGDVKIENQKLNFKEKAQAKVGSLDNVGHVPAGGTVKIESHKLTFREKAKARTDHGADIVVSKSPNFSSSTSPWCSTSVNESLGSITSSSPRQQPAPHANDLSAVLVQQGL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9G5</accession>
    <name>A0A452H9G5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="23">1-phosphatidylinositol 3-phosphate 5-kinase</fullName>
            <ecNumber evidence="3">2.7.1.150</ecNumber>
            <ecNumber evidence="4">2.7.11.1</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="26">FYVE finger-containing phosphoinositide kinase</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="27">PIKfyve</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="25">Phosphatidylinositol 3-phosphate 5-kinase type III</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="24">Serine-protein kinase PIKFYVE</fullName>
        </alternativeName>
    </protein>
    <organism evidence="34 35">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="35" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="34" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="34" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.1"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="19">
            <dbReference type="Rhea" id="RHEA:17990"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="20">
            <text>a 1,2-diacyl-sn-glycero-3-phospho-(1D-myo-inositol) + ATP = a 1,2-diacyl-sn-glycero-3-phospho-(1D-myo-inositol-5-phosphate) + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:44680"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:57795"/>
            <dbReference type="ChEBI" id="CHEBI:57880"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="20">
            <dbReference type="Rhea" id="RHEA:44681"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="21">
            <text>a 1,2-diacyl-sn-glycero-3-phospho-(1D-myo-inositol-3-phosphate) + ATP = a 1,2-diacyl-sn-glycero-3-phospho-(1D-myo-inositol-3,5-bisphosphate) + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:13609"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:57923"/>
            <dbReference type="ChEBI" id="CHEBI:58088"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.1.150"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="21">
            <dbReference type="Rhea" id="RHEA:13610"/>
        </physiologicalReaction>
    </comment>
    <comment type="subunit">
        <text evidence="22">Component of the PI(3,5)P2 regulatory complex/PAS complex, at least composed of PIKFYVE, FIG4 and VAC14. VAC14 nucleates the assembly of the complex and serves as a scaffold by pentamerizing into a star-shaped structure, which can bind a single copy each of PIKFYVE and FIG4 and coordinates their activities. Interacts (via chaperonin-like domain) with RABEPK; the interaction recruits RABEPK to the endosomal membrane. Interacts with SPAG9. Interacts with EGFR.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="18">Cytoplasmic vesicle</location>
            <location evidence="18">Phagosome membrane</location>
            <topology evidence="18">Peripheral membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Early endosome membrane</location>
            <topology evidence="1">Peripheral membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Late endosome membrane</location>
            <topology evidence="2">Peripheral membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="2.7.1.150" evidence="3"/>
    <dbReference type="EC" id="2.7.11.1" evidence="4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013053.1">
        <property type="protein sequence ID" value="ENSGAGP00000011402.1"/>
        <property type="gene ID" value="ENSGAGG00000008802.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031901">
        <property type="term" value="C:early endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000139">
        <property type="term" value="C:Golgi membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031902">
        <property type="term" value="C:late endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030670">
        <property type="term" value="C:phagocytic vesicle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000285">
        <property type="term" value="F:1-phosphatidylinositol-3-phosphate 5-kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016308">
        <property type="term" value="F:1-phosphatidylinositol-4-phosphate 5-kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0052810">
        <property type="term" value="F:1-phosphatidylinositol-5-kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008104">
        <property type="term" value="P:intracellular protein localization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035556">
        <property type="term" value="P:intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032438">
        <property type="term" value="P:melanosome organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030593">
        <property type="term" value="P:neutrophil chemotaxis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0090385">
        <property type="term" value="P:phagosome-lysosome fusion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046488">
        <property type="term" value="P:phosphatidylinositol metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903426">
        <property type="term" value="P:regulation of reactive oxygen species biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd04448">
        <property type="entry name" value="DEP_PIKfyve"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd03334">
        <property type="entry name" value="Fab1_TCP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd15725">
        <property type="entry name" value="FYVE_PIKfyve_Fab1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd17300">
        <property type="entry name" value="PIPKc_PIKfyve"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.810.10:FF:000001">
        <property type="entry name" value="1-phosphatidylinositol 3-phosphate 5-kinase FAB1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.10:FF:000206">
        <property type="entry name" value="1-phosphatidylinositol 3-phosphate 5-kinase isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.40.10:FF:000057">
        <property type="entry name" value="1-phosphatidylinositol 3-phosphate 5-kinase isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.800.10:FF:000004">
        <property type="entry name" value="1-phosphatidylinositol 3-phosphate 5-kinase isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.50.7.10:FF:000007">
        <property type="entry name" value="1-phosphatidylinositol 3-phosphate 5-kinase isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.810.10">
        <property type="entry name" value="2-Layer Sandwich"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.50.7.10">
        <property type="entry name" value="GroEL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.800.10">
        <property type="entry name" value="Phosphatidylinositol Phosphate Kinase II Beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.10">
        <property type="entry name" value="Winged helix-like DNA-binding domain superfamily/Winged helix DNA-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002423">
        <property type="entry name" value="Cpn60/GroEL/TCP-1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000591">
        <property type="entry name" value="DEP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027409">
        <property type="entry name" value="GroEL-like_apical_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043548">
        <property type="entry name" value="PIKfyve"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037378">
        <property type="entry name" value="PIKfyve_DEP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044769">
        <property type="entry name" value="PIKfyve_PIPKc"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027483">
        <property type="entry name" value="PInositol-4-P-4/5-kinase_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002498">
        <property type="entry name" value="PInositol-4-P-4/5-kinase_core"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027484">
        <property type="entry name" value="PInositol-4-P-5-kinase_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027410">
        <property type="entry name" value="TCP-1-like_intermed_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036388">
        <property type="entry name" value="WH-like_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036390">
        <property type="entry name" value="WH_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000306">
        <property type="entry name" value="Znf_FYVE"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017455">
        <property type="entry name" value="Znf_FYVE-rel"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011011">
        <property type="entry name" value="Znf_FYVE_PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46715">
        <property type="entry name" value="1-PHOSPHATIDYLINOSITOL 3-PHOSPHATE 5-KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46715:SF1">
        <property type="entry name" value="1-PHOSPHATIDYLINOSITOL 3-PHOSPHATE 5-KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00118">
        <property type="entry name" value="Cpn60_TCP1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00610">
        <property type="entry name" value="DEP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01363">
        <property type="entry name" value="FYVE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01504">
        <property type="entry name" value="PIP5K"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00049">
        <property type="entry name" value="DEP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00064">
        <property type="entry name" value="FYVE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00330">
        <property type="entry name" value="PIPKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57903">
        <property type="entry name" value="FYVE/PHD zinc finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52029">
        <property type="entry name" value="GroEL apical domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54849">
        <property type="entry name" value="GroEL-intermediate domain like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56104">
        <property type="entry name" value="SAICAR synthase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46785">
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    <keyword evidence="13 29" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="17" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="9" id="KW-0967">Endosome</keyword>
    <keyword evidence="11 29" id="KW-0418">Kinase</keyword>
    <keyword evidence="15" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="16" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0479">Metal-binding</keyword>
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    <keyword evidence="12" id="KW-0862">Zinc</keyword>
    <keyword evidence="10 28" id="KW-0863">Zinc-finger</keyword>
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            <begin position="165"/>
            <end position="225"/>
        </location>
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            <begin position="1778"/>
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            <begin position="1"/>
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    <feature type="region of interest" description="Disordered" evidence="30">
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            <begin position="61"/>
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            <begin position="491"/>
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            <begin position="1529"/>
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            <begin position="1714"/>
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            <begin position="1798"/>
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            <begin position="70"/>
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    <evidence type="ECO:0000256" key="28">
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        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00781"/>
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    </evidence>
    <evidence type="ECO:0000256" key="30">
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            <dbReference type="SAM" id="MobiDB-lite"/>
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            <dbReference type="PROSITE" id="PS50178"/>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011402.1"/>
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    </evidence>
    <evidence type="ECO:0000313" key="35">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2118" mass="239881" checksum="844745E56CCC61CC" modified="2019-05-08" version="1">MATDDRQSPTLDSVSDLPRSPSSPSHLTHFKPLTPDQDEPPFKSAYSSFVNLFRFGKDDGRLASPAESGQGEQQALSGTWSSPQHTSRAQSLRSPVPYKKQLTGELQRRASATLDTRRKVEPSLSGHDPRTAVQLRSLSTVLKRLKEIMEGKSQDSDLKQYWMPDSQCKECYDCSEKFTTFRRRHHCRLCGQIFCSRCCNQEIPGKFMGYTGDLRACTYCRKIALSYAHSTDSNSIGEDLNALSDSTCCVSVLEPGEPRTPVGSRKASRNIFLEEDLTWQSLIHPDSSTTTLSTRLVSVQEDPGKSPARNRSASITNLSLDRSASPMVPAYETSVSPQPNRTYLKAETNEDERKILLDSVQLKDLWKKICHHNSGMEFQDHRYWLRTHPNCIVGKELVNWLIRNGHITTRAQAIAIGQALVDGRWLDCVSHHDQIFRDEYALYRPLQSTEFSETPSPDSDSVNSVEGHSEPSWFKDIKFDDSDTEQIADEGEDNLANSASPSKRTSVSSFQSTMDSDSAASINLNVELDNVNFHIKKHSKYPHVPPHPADQKEYLNPDNGGQQMISISDAFIKESLFNRRVEEKSKELFFTPLGWHHSNLDLLREENGEKQAMERLLSANHNHMMALLQQLLNNESLSLSWRDIIVPVVCQVVQTVRPDVKNRDDDMDIRQFVHIKKIPGGKKFDSVVVNGFVCTKNVAHKKMNTYIKNPKILLLKCSIEYLYREETKFTCIDPIVLQEREFLKNYVQRIVDVRPTLVLVEKTVSRIAQDMLLEHGITLVINVKPQVLDRISRMTQGDLVMSMDQLLTKPHLGTCHKFYMQVFQLLNDQTKTLMFFEGCPPHLGCTIKLRGAADYELTRVKEILIFMVCVAYHSQLEISFLMDEFAMPPSLTKNFHTLIESQGYENGHQDLFNGEEFSTVVRDVELSPEKVPMVSESVSSDEASTLEQRTLFEEDQDDDDDVLREGSSLRHQELLRMEYSLPVSSNITTPVPESRLPFHNTDQHPGTLYSQHLETLKQPGDLQDSKSQMRVFRDPLQDDTGLYVMEEVTSSEDRLKAYSSAFKQELKDVILCISPVITFHEPFLLTEMGMRCPARDYFPEQVYLSPFLNKEYKELESRRKKQLLRDLSGLQGINGSIQAKAVQVLPTHELVNTRIAEHLGDSQSLARMLADYRARGGRIQQKNADPFAQCKDASGVPGGKAGGRNEEDERGLVRGESVWSHKVDCLSPVNHQRLCVLFSSSSAQSSNAPSACVSPWIVTMEFYGKNDLTLGVFLERYCFRPSYQCPSMFCETPMVHHVRRFVHGQGCVQIILKELDSPVPGYQHTILTYSWCRLCKQVTPVVPLSNDSWSMSFAKYLELRFYGHQYTRRANAEPCGHSIHHDYHQYFSYNQMVASFSYSPIRLLEVCVPHPKIYIKRQAPLKVTILQDLKDFSQKVSQVYLAVDDRLTSLKTDTFSKTREEKMEDLFAQKEMEEVEFRNWIEKIQARMLSPSLDTSQQLQSVFESLIAKKQSLCEMLQAWNNRLQDLFQQEKGRKRPSVPPSPGRLRQGEENKISAMDASPRNTSPALQNGEKDDRFLTTLSSQSSTSSTHLQLPTPPEIVSEQLTGGPSFANSLSEPDTTSSSEDIFDGHSLGSTDSQVKEKSTMKAILANLLPGNSYNPIPFPFDPDKHYLMYEHERVPIAVCEKEPSSIIAFALSCKEYRNGLDELSKASLKSSSEEGLPPNSMSDNKPKSSSPVRLPETNVGQANRTAEAEQPKKVSGVLSFFRGTGGKSPDLSAQKKETLRGADSAYYQVGQMGKEGPESQGAETQDEADGGDAQKKQLANPHVELQFSDANAKFYCRIYYAGEFHKMRDVILGSREEDFIRSLSHSLPWQARGGKSGAAFYVTEDDRFILKQMPRLEVQSFLDFAPHYFTYITNAVQQKKPTALAKILGVYRIGYKNSQNNTEKKLDLLVMENLFYGRKMAQVFDLKGSLRNRNVKTDTGKESCDVVLLDENLLKMVRDNPLYIRSHCKAVLRASIHSDSHFLSSHLIIDYSLLVGRDDTSNELVVGIIDYIRTFTWDKKLEMVVKSTGILGGQGKMPTVVSPELYRTRFCEAMDKYFLMVPDHWTGLGLNC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9G6</accession>
    <name>A0A452H9G6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Anoctamin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="8">Membrane</location>
            <topology evidence="8">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 8">Belongs to the anoctamin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9G6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013057.1">
        <property type="protein sequence ID" value="ENSGAGP00000011406.1"/>
        <property type="gene ID" value="ENSGAGG00000007328.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005229">
        <property type="term" value="F:intracellularly calcium-gated chloride channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046983">
        <property type="term" value="F:protein dimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032394">
        <property type="entry name" value="Anoct_dimer"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007632">
        <property type="entry name" value="Anoctamin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049452">
        <property type="entry name" value="Anoctamin_TM"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12308">
        <property type="entry name" value="ANOCTAMIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12308:SF28">
        <property type="entry name" value="ANOCTAMIN-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16178">
        <property type="entry name" value="Anoct_dimer"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04547">
        <property type="entry name" value="Anoctamin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6 8" id="KW-0472">Membrane</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 8" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 8" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="279"/>
            <end position="305"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="355"/>
            <end position="374"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="431"/>
            <end position="457"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="477"/>
            <end position="497"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="525"/>
            <end position="546"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="645"/>
            <end position="667"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="695"/>
            <end position="719"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="8">
        <location>
            <begin position="770"/>
            <end position="791"/>
        </location>
    </feature>
    <feature type="domain" description="Anoctamin dimerisation" evidence="11">
        <location>
            <begin position="55"/>
            <end position="265"/>
        </location>
    </feature>
    <feature type="domain" description="Anoctamin transmembrane" evidence="10">
        <location>
            <begin position="268"/>
            <end position="813"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="812"/>
            <end position="839"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009671"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="RuleBase" id="RU280814"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="Pfam" id="PF04547"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF16178"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011406.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="848" mass="99256" checksum="23ECFFF359BE062A" modified="2025-10-08" version="2">MFPPTPKEKKIKPFILNHTEICSSLGSNGDSQLCFGNCFCSSQGVPDKNKSNGLYFRDGKCRIDYILVYRKSSPQTEKREIFERNIRAEGLQMEKESSLTNSDIIFVKLHAPWEVLGRYAEVMNVRMPFRIERQISRFQGWLPRKPMRLDKETLPDLEENDCYTAPFSQQRIHHFIIHNKDTFFNNATRSRIVHHILQRVKYEEGKNKIGLNRLLTNGSYEAAFPLHEGSYRSKNSIRTHGAENHRHLLYECWASWGMWYKYQPLDLVRRYFGEKIGLYFAWLGWYTGMLFPAAFIGLFVFLYGVTTLNHCQVSKEVCQATDIIMCPICDKFCPFMRLSDSCVYAKVTHLFDNGATVFFAVFMAVWATVFLEFWKRRRAVIAYDWDLIDWEEEEEEIRPQFEAKYSKKERMNPISGKPEPYQAFADKCSRLIVSASGIFFMICVVIAAVFGIVIYRVVTVSTFAAFKWALIRNNSQVATTGTAVCINFCIIMLLNLYEKVALLLTNLEQPRTESEWENSFTLKMFLFQFVNLNSSTFYIAFFLGRFTGHPGAYLRLINRWRLEECHPSGCLIDLCMQMGIIMVLKQTWNNFMELGYPLIQNWWTRRKLYQEHGSQRKTSFPQWEKDYNLQPMNAYGLFDEYLEMILQFGFTTIFVAAFPLAPLLALLNNIIEIRLDAYKFVTQWRRPLASRAKDIGIWYGILEGIGILSVITNAFVVAVTSDFIPRLVYAYKYGPCAGQGEAGQNSSLSLYRDYRDPPHSPVPYGYTLQFWHVLAARLAFIIVFEHLVFCIKHLISYLIPDLPKDLRDRMRREKYLIQEMMYEAELERLQKERKERKKNGKSHHNEWP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9G8</accession>
    <name>A0A452H9G8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Microtubule-associated protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 7">Cytoplasm</location>
            <location evidence="1 7">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000012864.1">
        <property type="protein sequence ID" value="ENSGAGP00000011231.1"/>
        <property type="gene ID" value="ENSGAGG00000008671.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005874">
        <property type="term" value="C:microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043005">
        <property type="term" value="C:neuron projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008017">
        <property type="term" value="F:microtubule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000226">
        <property type="term" value="P:microtubule cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031175">
        <property type="term" value="P:neuron projection development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027324">
        <property type="entry name" value="MAP2/MAP4/Tau"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001084">
        <property type="entry name" value="MAP_tubulin-bd_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11501">
        <property type="entry name" value="MICROTUBULE-ASSOCIATED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11501:SF16">
        <property type="entry name" value="MICROTUBULE-ASSOCIATED PROTEIN 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00418">
        <property type="entry name" value="Tubulin-binding"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00229">
        <property type="entry name" value="TAU_MAP_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51491">
        <property type="entry name" value="TAU_MAP_2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2 7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6 7" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="4 7" id="KW-0493">Microtubule</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1"/>
            <end position="21"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="50"/>
            <end position="82"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="276"/>
            <end position="309"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="436"/>
            <end position="508"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="526"/>
            <end position="587"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="616"/>
            <end position="661"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="785"/>
            <end position="874"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="944"/>
            <end position="982"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1021"/>
            <end position="1052"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1119"/>
            <end position="1199"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1484"/>
            <end position="1518"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1612"/>
            <end position="1635"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1661"/>
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        </location>
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            <begin position="1813"/>
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        </location>
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            <begin position="1865"/>
            <end position="2254"/>
        </location>
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    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="2299"/>
            <end position="2360"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="276"/>
            <end position="286"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="436"/>
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        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="543"/>
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    <feature type="compositionally biased region" description="Basic residues" evidence="8">
        <location>
            <begin position="562"/>
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    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
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            <begin position="616"/>
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    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="636"/>
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    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="651"/>
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        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="812"/>
            <end position="822"/>
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    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="850"/>
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    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
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    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
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        </location>
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        <location>
            <begin position="1484"/>
            <end position="1498"/>
        </location>
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        <location>
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            <begin position="1680"/>
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        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="1819"/>
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        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="1870"/>
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        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="1924"/>
            <end position="1936"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="1963"/>
            <end position="1986"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="1999"/>
            <end position="2012"/>
        </location>
    </feature>
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        <location>
            <begin position="2013"/>
            <end position="2027"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="2028"/>
            <end position="2044"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="2065"/>
            <end position="2086"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="2096"/>
            <end position="2105"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="2118"/>
            <end position="2168"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="2311"/>
            <end position="2327"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="2328"/>
            <end position="2342"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU000686"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011231.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2360" mass="255983" checksum="B216927FAD3982F6" modified="2019-05-08" version="1">MADFDHNLSLADALTEPPAEIEEEVKRDFIATLEAEKFDDVVGETVGKTDYIPLLDDDDDAKAGSQEPKSKPHTDGGQVESTSAVGPAVLENGDHGTEDDSTVFPREITDEKMSYKEFLDRNETWAVDDRDLCFEPQSIFRPMEGTEPFRMHREDVLCDLLLRPSEMETRLPFTEHFEASEEVHAPHAAMMVPELPSLGSPYSPAEVIDPSAFITLDSTTESLLNIEAPARVTVPEENWLGAEHAVEELDESSFVEPPEPTRITDAAEKYLPGSPVAAAPAAVPPALTEPTGETKATDTPQVEPTESVPVMGDESVPIVEELMVFEPSVEQHKSAVNEPPTVLSAPTVVMEQKMTVTEASTLGADSTPVLENVEENKLSPSKHTEHLLKPNEELPELAVPVEAKEPLTLIEIKETLPEKTAPAADLTIVETLKEEAELSHAHHTEPPQKKSPLEPTGVPTAQVRQANKSSDRRFGRAKPAAVPGADVPEELLVGLPEQKSTDPKADPFSMAELGWVSGTFSRTRAPHKKAAGQLLSMPPEFVESQRDVPRESWDSEGSPVIVKKKKKKQKQKRNQQPRTMELGDENVEVSKAPKVPLFAAELQKPDVPFVLPAEGVKEHSAHSREGQRKGTSNVLRDAKAGTESHLVDDQNFISSPTAGQQSLKTEGPFKFLLDAKAGEVMKQEEMKADSLFQSKSKNKQVPMEKLECKAEHTKMGDPSSALTQTKPIEISFIDKNKKIECEHSDLEVSLSKGINLNKVETPLETKPVEINLIDENKEIKCTSHKHVAADETVPSEVQTPGGILMGEKPKKRSSDEKSRKAENGFSKQPAVLEIKTDTTNLPAAAKRVNKTKETDSFDKSQETGSITSEPPLETATDITKVPLIALVSDKPKEGIAGQNKKADFSFSEQPFVLETKTDTAKLHAPAETVDITNIGFTDKSKQIGFDHPAVTDPSMALVTDKPKKRGGDGKNKKVKNSSEQPVLMEVEIDPDKLPAVVKTAGKTKEIIFTDEDKEPGVAIAEHSLEDMTGPSTAKVAEKPKKRTSDGKNKKAEKGYFEQPFFLEAKEDTSSLPTVMEKDDKTKKVSFIDKGKETDFTTADHLLEGATATMEVQGPITTLVTEEPKNGITGKSEKAEFSISEQPDVAKLPASAETISKTKEDSLMDKSKETGFTTDSSRALMSDKPKEKGSDRKGKKVGKSSFEQPFLLETKIDTSKLPTVVEKAEKMKEMGFADIGKEPIFTTTEHQLENLTDTTKVQVPTMKLITEKPKIKESNEKSFFEQLVPSDYKMDTAQSETLTKTKETPFSDKGKETDFTTLEHLLEDIRDTAKAHIPMTELVADKPKKRCSDGKSKKIENTAEHLAVLEAKTDSIKLPAVLKEANKIKETHSTDKGQETQFTISEHLLENITDTAKIQTPITPLVTGKPEETNKGNSTKADVHLEKPFLLETKVDAATLPGGADEIVDKTEAISSTDESQVAGFTILEHQTMTDPSPALVTNKSKKRGNDGKNREAKNASEQPVLLETKTDKSKIQPPIVTDLEYQMEEMGLVDENQNIKNFPSKHQMLWDNKGHLFTPFTQSGAVGVDRVGKASIFPEYPVKDEGWKVPVPPAVVLEGTNKESSTSDKREENKHSHYEHSVTLEGEDAGVQAFTSVKEGDKIKVTSAGDKKKKGKWPSLDSPVKQDAKAGRDEVHTSILAEIDDKEISTECASLEHLVKETDAMKRTIPTDAKVTDKMEGSSSAGEKDVGCISPELTVQWENKAEAVVLQVLATAEVVNKAKEAELGEQSFSECPVSLEKKTDAAEVASVTLRDAQTEEISPTDKIKSHSEHSEGDAADKMKAGLKDFQALKSEEDKYADLPQKQIDGSGQKVVKETKKEERVKATEQIKGYMRPTKSRGLPAPPPRPAVQAREKPRQLKANGMSRQRQEKAKPEEIKPVELVTGSDITAPPNKELPPSPEKKSKPSVSTPSAKPAATKTKPLSTTSPKRPASATPGQNKKPTSPTAGPTSATTPKRSATSTTRPSTLTPKDTKPKVTDAKSPDKRTSLSKPLSATTPRAAVKVSPATPRTTAASPITTASGPRSTATSPPKRPTSIKTETKLADARKTTAKSPSADLSRPKSAPANSAKSSATTPTATTPGTLASPGVATSRPKPKPAAARPTTASSATPEAKKPSTVKAPLKTSTVPKPPRPTSSVSAPDLKNIRSKIGSTDNIKHQPGGGRVQIQSKKVDISKVSSKCGSKTNIKHKPGGGDVKIENQKLNFKEKAQAKVGSLDNVGHVPAGGTVKIESHKLTFREKAKARTDHGADIVVSKSPNFSSSTSPWCSTSVNESLGSITSSSPRQQPAPHANDLSAVLVQQGL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9J0</accession>
    <name>A0A452H9J0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">IPT/TIG domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the plexin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9J0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013146.1">
        <property type="protein sequence ID" value="ENSGAGP00000011483.1"/>
        <property type="gene ID" value="ENSGAGG00000008707.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002116">
        <property type="term" value="C:semaphorin receptor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017154">
        <property type="term" value="F:semaphorin receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007162">
        <property type="term" value="P:negative regulation of cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050772">
        <property type="term" value="P:positive regulation of axonogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030334">
        <property type="term" value="P:regulation of cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008360">
        <property type="term" value="P:regulation of cell shape"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd12792">
        <property type="entry name" value="RasGAP_plexin_B2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000203">
        <property type="entry name" value="Plexin B2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000798">
        <property type="entry name" value="Plexin B2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.506.10:FF:000035">
        <property type="entry name" value="Plexin b2a"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.506.10">
        <property type="entry name" value="GTPase Activation - p120gap, domain 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014756">
        <property type="entry name" value="Ig_E-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002909">
        <property type="entry name" value="IPT_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031148">
        <property type="entry name" value="Plexin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013548">
        <property type="entry name" value="Plexin_cytoplasmic_RasGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046800">
        <property type="entry name" value="Plexin_RBD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057533">
        <property type="entry name" value="PSI_Plexin-B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041019">
        <property type="entry name" value="TIG1_plexin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22625">
        <property type="entry name" value="PLEXIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22625:SF9">
        <property type="entry name" value="PLEXIN-B2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08337">
        <property type="entry name" value="Plexin_cytopl"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20170">
        <property type="entry name" value="Plexin_RBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24317">
        <property type="entry name" value="PSI_Plexin-B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01833">
        <property type="entry name" value="TIG"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17960">
        <property type="entry name" value="TIG_plexin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00429">
        <property type="entry name" value="IPT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81296">
        <property type="entry name" value="E set domains"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="12" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="9" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="8" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0675">Receptor</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <keyword evidence="4" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7" id="KW-1133">Transmembrane helix</keyword>
    <feature type="domain" description="IPT/TIG" evidence="13">
        <location>
            <begin position="208"/>
            <end position="301"/>
        </location>
    </feature>
    <feature type="domain" description="IPT/TIG" evidence="13">
        <location>
            <begin position="302"/>
            <end position="388"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="12">
        <location>
            <begin position="577"/>
            <end position="604"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010297"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="SMART" id="SM00429"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011483.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1187" mass="134690" checksum="1D5DFBE076F0A325" modified="2019-05-08" version="1">MPNHWLWSPDGVCMAITSAHPQNMSRKAPGKVELTVSPLPELTESDMLWCQFGETTNIAELRDGTIICDPPTTIPHTPKGQDHVSVSLILTFGEEKMFFASATYPFYDCEEEDCPQFLNPNPQIIPMEFPTIVHFEGKNLENYQVNELIPSSHLTYMTLYNCSYGRSDCSLCLAADPGYDCVWCEEEGKPNKCIYKNLCKTSTNTCPHPVITRIEPKTGPLSGGILLTIMGSNLGIKAEDVKSITVAGTECLFKEDFYSVSIRYLIVCQAGNVSVPSQGKIEVDINGKRGSSTSEVQFTYQQPHPTGISPTEGPRAGGTILTITGNNLETGSQKDVRVLLGDVPCNITDFGEQIVCITGQHESGGKVQVTVDHGDTENQLNDPWFTYSENPTVTKFFPKNSFNITKINESTIVFSSPPILEDPENYRITTLIKMDGCSLVLNAENNDPFKYVADPTFENFTDGIKKQVNKLIHAKGSNLNKAMTIEEAQAFVGAEPCNIKTLTEKDLYCEPPEVQPQPKKRQKRDTINNLPEFIVKFGLREWILGRVEYDTRVSDVPLDVILPSVIFPMVAIIIISIYCYRRKSQQAEREYEKIKSQLEGLEESVRDRCKKEFTDLMIEMEDQTNDVSEAGIPVLDYKMYTDRVFFLPSKDGEKDVMITGKLDIPEARKQTVEQALNQFSNLLNSKSFLINFIHTLEKQREFSAREKVYFASLLTVALHGKLEYYTDIMRTLFLELMEQYVVAKNPKLMLRRSETVVERMLSNWMAICLYQYLKDNAGEPLYKLFKAIKHQVEKGPVDAVLMKAKYTLNDTGLLGDDVEYTQLTVNVYVQDGGTDSVPVKVLNCDTISQVKEKIIDQVYRNLPCSQWPKADSVVLVVFKPASNSHVLWDDFSIFVFLKVRDGATLILSKIGISQQPEDNQQDVPGERHALLEDENKVWHLVRPADEVDEGKSKRGSMKEKERTKAITEIYLTRLLSVKGTLQQFVDNFFHSVLNSNHVVPPAVKYFFDFLDEQAEKHELKDEDTIHIWKTNSLPLRFWVNILKNPHFIFDVHVHEVVDASLSVIAQTFMDACTRTEHKLSRDSPSNKLLYAKEISNYKKMVEDYYKGIRQMVQVSDQDMNTHLAEISRAHTDSLNTLVALHQLYQYTNKYYDEIINALEEDPAAQKMQLAFRLQQIAAALENKVTDL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9J7</accession>
    <name>A0A452H9J7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="22">Integrin beta</fullName>
        </recommendedName>
    </protein>
    <organism evidence="27 28">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="28" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="27" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="27" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="20">Heterodimer of an alpha and a beta subunit. Interacts with FLNB. Interacts with HAX1. ITGAV:ITGB6 interacts with FBN1. ITGAV:ITGB6 interacts with TGFB1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell junction</location>
            <location evidence="1">Focal adhesion</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2 22">Cell membrane</location>
            <topology evidence="2 22">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 22">Belongs to the integrin beta chain family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9J7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013096.1">
        <property type="protein sequence ID" value="ENSGAGP00000011439.1"/>
        <property type="gene ID" value="ENSGAGG00000007913.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009986">
        <property type="term" value="C:cell surface"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005925">
        <property type="term" value="C:focal adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034685">
        <property type="term" value="C:integrin alphav-beta6 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033627">
        <property type="term" value="P:cell adhesion mediated by integrin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098609">
        <property type="term" value="P:cell-cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007160">
        <property type="term" value="P:cell-matrix adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007229">
        <property type="term" value="P:integrin-mediated signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.100:FF:000004">
        <property type="entry name" value="Integrin beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000043">
        <property type="entry name" value="Integrin beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000075">
        <property type="entry name" value="Integrin beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000328">
        <property type="entry name" value="Integrin beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.410:FF:000002">
        <property type="entry name" value="Integrin beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.1240.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.100">
        <property type="entry name" value="Cytochrome c1, transmembrane anchor, C-terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1510">
        <property type="entry name" value="ntegrin, alpha v. Chain A, domain 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.410">
        <property type="entry name" value="von Willebrand factor, type A domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040622">
        <property type="entry name" value="EGF_integrin_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057073">
        <property type="entry name" value="EGF_integrin_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015812">
        <property type="entry name" value="Integrin_bsu"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014836">
        <property type="entry name" value="Integrin_bsu_cyt_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012896">
        <property type="entry name" value="Integrin_bsu_tail"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036349">
        <property type="entry name" value="Integrin_bsu_tail_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002369">
        <property type="entry name" value="Integrin_bsu_VWA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032695">
        <property type="entry name" value="Integrin_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057243">
        <property type="entry name" value="Integrin_I-EGF_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036465">
        <property type="entry name" value="vWFA_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10082">
        <property type="entry name" value="INTEGRIN BETA SUBUNIT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10082:SF11">
        <property type="entry name" value="INTEGRIN BETA-6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23105">
        <property type="entry name" value="EGF_integrin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18372">
        <property type="entry name" value="I-EGF_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08725">
        <property type="entry name" value="Integrin_b_cyt"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07965">
        <property type="entry name" value="Integrin_B_tail"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00362">
        <property type="entry name" value="Integrin_beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF002512">
        <property type="entry name" value="Integrin_B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01186">
        <property type="entry name" value="INTEGRINB"/>
    </dbReference>
    <dbReference type="SMART" id="SM00187">
        <property type="entry name" value="INB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01241">
        <property type="entry name" value="Integrin_b_cyt"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01242">
        <property type="entry name" value="Integrin_B_tail"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69687">
        <property type="entry name" value="Integrin beta tail domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69179">
        <property type="entry name" value="Integrin domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53300">
        <property type="entry name" value="vWA-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00243">
        <property type="entry name" value="I_EGF_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS52047">
        <property type="entry name" value="I_EGF_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-0106">Calcium</keyword>
    <keyword evidence="12 22" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="13" id="KW-0965">Cell junction</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="17 21" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="19" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="15 22" id="KW-0401">Integrin</keyword>
    <keyword evidence="11" id="KW-0460">Magnesium</keyword>
    <keyword evidence="16 23" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="18" id="KW-0675">Receptor</keyword>
    <keyword evidence="28" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0677">Repeat</keyword>
    <keyword evidence="8" id="KW-0732">Signal</keyword>
    <keyword evidence="6 22 23" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="14 23" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="23">
        <location>
            <begin position="661"/>
            <end position="683"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin beta subunit VWA" evidence="24">
        <location>
            <begin position="2"/>
            <end position="407"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin beta subunit tail" evidence="26">
        <location>
            <begin position="577"/>
            <end position="660"/>
        </location>
    </feature>
    <feature type="domain" description="Integrin beta subunit cytoplasmic" evidence="25">
        <location>
            <begin position="684"/>
            <end position="730"/>
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    </feature>
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        <location>
            <begin position="150"/>
            <end position="157"/>
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    <feature type="disulfide bond" evidence="21">
        <location>
            <begin position="205"/>
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    <feature type="disulfide bond" evidence="21">
        <location>
            <begin position="379"/>
            <end position="623"/>
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    <feature type="disulfide bond" evidence="21">
        <location>
            <begin position="405"/>
            <end position="409"/>
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    <feature type="disulfide bond" evidence="21">
        <location>
            <begin position="420"/>
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    <feature type="disulfide bond" evidence="21">
        <location>
            <begin position="429"/>
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    <feature type="disulfide bond" evidence="21">
        <location>
            <begin position="434"/>
            <end position="443"/>
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    <feature type="disulfide bond" evidence="21">
        <location>
            <begin position="470"/>
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    <feature type="disulfide bond" evidence="21">
        <location>
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    <feature type="disulfide bond" evidence="21">
        <location>
            <begin position="477"/>
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    <feature type="disulfide bond" evidence="21">
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            <begin position="511"/>
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    <feature type="disulfide bond" evidence="21">
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            <begin position="513"/>
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    <feature type="disulfide bond" evidence="21">
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            <begin position="518"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00004246"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
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            <dbReference type="ARBA" id="ARBA00023037"/>
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            <dbReference type="ARBA" id="ARBA00023170"/>
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        <source>
            <dbReference type="PIRSR" id="PIRSR002512-1"/>
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    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="RuleBase" id="RU000633"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="SAM" id="Phobius"/>
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            <dbReference type="SMART" id="SM00187"/>
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        <source>
            <dbReference type="SMART" id="SM01241"/>
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    </evidence>
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        <source>
            <dbReference type="SMART" id="SM01242"/>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011439.1"/>
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    </evidence>
    <evidence type="ECO:0000313" key="28">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
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    </evidence>
    <sequence length="742" mass="82038" checksum="D90AD026B82FA51F" modified="2019-05-08" version="1">MNYTDSSGVHGRCDAPENLLSKGCQLKFIEFPISKVEIHKNKPLSIGIQNNSDVTQISPQKLTFRLRPGNEETIEIKVRQTEDYPVDLYYLMDLSASMDDDLNTIKELGSTLSKEMSKLTSNFRLGFGSFVEKPVSPFIKTTAEEINNPCRSVPRDCLPTFGYKHVLPLTNDAERFNEIVKEQKISANIDTPEGGFDAIMQAAVCKEKIGWRNDSLHLLVFVSDADSHFGMDSKLAGIVIPNDGHCHLDQNNEYSMSTVLEYPTIGQLTDKLVQNNVLLIFAVTKEQVYFYENYAKLIPGATVGQLQKDSGNILQLIIDAYKELRSEVELEVLGDTEGLNLSFTAICNNGSIFPHQRKCSHIKVGETVSFNMTVSLPTCEKRRRQIVIKPVGLSDALEIDIQLECSCNCQKEAEMNSSKCSKGKGSFECGVCACNPGYMGPYCECNENSLSTNSCKRSPEQSSCSGRGDCYCGQCICHFSVYGNIYGLYCECDNFSCVRFKGLQCGGNGDCDCGECVCHRGWTGEYCNCTTDTNSCVSEDGILCSGRGECICGTCVCTNPGASGSTCERCPTCGDPCNSKRSCIECHLASDVQSQAECNEKCRLVDATVNSAEDFSKDKSIPCSLQGENECVTTFLMATDEQGRTIIHSIKQKDCPQPPNIPMIMLGVSLAILLIGIVLLCIWKLLVSFQDRKEVAKFEAERSKAKWQTGTNPLYRGSTTTFKNVTYKHQEKQKVDITADFY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9K3</accession>
    <name>A0A452H9K3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">E3 ubiquitin-protein ligase CBL</fullName>
            <ecNumber evidence="12">2.3.2.27</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="12">E3 ubiquitin-protein ligase which accepts ubiquitin from specific E2 ubiquitin-conjugating enzymes, and transfers it to substrates, generally promoting their degradation by the proteasome.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>S-ubiquitinyl-[E2 ubiquitin-conjugating enzyme]-L-cysteine + [acceptor protein]-L-lysine = [E2 ubiquitin-conjugating enzyme]-L-cysteine + N(6)-ubiquitinyl-[acceptor protein]-L-lysine.</text>
            <dbReference type="EC" id="2.3.2.27"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="12">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="domain">
        <text evidence="12">The N-terminus is composed of the phosphotyrosine binding (PTB) domain, a short linker region and the RING-type zinc finger. The PTB domain, which is also called TKB (tyrosine kinase binding) domain, is composed of three different subdomains: a four-helix bundle (4H), a calcium-binding EF hand and a divergent SH2 domain.</text>
    </comment>
    <dbReference type="EC" id="2.3.2.27" evidence="12"/>
    <dbReference type="AlphaFoldDB" id="A0A452H9K3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000012900.1">
        <property type="protein sequence ID" value="ENSGAGP00000011261.1"/>
        <property type="gene ID" value="ENSGAGG00000008688.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00143"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045121">
        <property type="term" value="C:membrane raft"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001784">
        <property type="term" value="F:phosphotyrosine residue binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030971">
        <property type="term" value="F:receptor tyrosine kinase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017124">
        <property type="term" value="F:SH3 domain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061630">
        <property type="term" value="F:ubiquitin protein ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007166">
        <property type="term" value="P:cell surface receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016567">
        <property type="term" value="P:protein ubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="GO" id="GO:0023051">
        <property type="term" value="P:regulation of signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd09920">
        <property type="entry name" value="SH2_Cbl-b_TKB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.40.10:FF:000015">
        <property type="entry name" value="E3 ubiquitin-protein ligase CBL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.505.10:FF:000007">
        <property type="entry name" value="E3 ubiquitin-protein ligase CBL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.238.10:FF:000182">
        <property type="entry name" value="E3 ubiquitin-protein ligase CBL-C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.930.20">
        <property type="entry name" value="Adaptor protein Cbl, N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.238.10">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.505.10">
        <property type="entry name" value="SH2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024162">
        <property type="entry name" value="Adaptor_Cbl"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014741">
        <property type="entry name" value="Adaptor_Cbl_EF_hand-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036537">
        <property type="entry name" value="Adaptor_Cbl_N_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003153">
        <property type="entry name" value="Adaptor_Cbl_N_hlx"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014742">
        <property type="entry name" value="Adaptor_Cbl_SH2-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024159">
        <property type="entry name" value="Cbl_PTB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011992">
        <property type="entry name" value="EF-hand-dom_pair"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036860">
        <property type="entry name" value="SH2_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001841">
        <property type="entry name" value="Znf_RING"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017907">
        <property type="entry name" value="Znf_RING_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23007">
        <property type="entry name" value="CBL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23007:SF12">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE CBL-C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02262">
        <property type="entry name" value="Cbl_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02761">
        <property type="entry name" value="Cbl_N2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02762">
        <property type="entry name" value="Cbl_N3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13920">
        <property type="entry name" value="zf-C3HC4_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00184">
        <property type="entry name" value="RING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47473">
        <property type="entry name" value="EF-hand"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47668">
        <property type="entry name" value="N-terminal domain of cbl (N-cbl)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57850">
        <property type="entry name" value="RING/U-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55550">
        <property type="entry name" value="SH2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51506">
        <property type="entry name" value="CBL_PTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00518">
        <property type="entry name" value="ZF_RING_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50089">
        <property type="entry name" value="ZF_RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8 12" id="KW-0106">Calcium</keyword>
    <keyword evidence="4 12" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0729">SH3-binding</keyword>
    <keyword evidence="3 12" id="KW-0808">Transferase</keyword>
    <keyword evidence="9" id="KW-0832">Ubl conjugation</keyword>
    <keyword evidence="6 12" id="KW-0833">Ubl conjugation pathway</keyword>
    <keyword evidence="7 12" id="KW-0862">Zinc</keyword>
    <keyword evidence="5 11 12" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="Cbl-PTB" evidence="15">
        <location>
            <begin position="13"/>
            <end position="319"/>
        </location>
    </feature>
    <feature type="domain" description="RING-type" evidence="14">
        <location>
            <begin position="349"/>
            <end position="388"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="437"/>
            <end position="480"/>
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        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
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    <sequence length="577" mass="63244" checksum="245DDE41726D2738" modified="2019-05-08" version="1">MAVAGGPGARRGPFALSAAARHSLDKALQGLEKLQRLVEQPGLGLRNSPPCLLQLLPQMRQHLLLIRGQPGASLSCLWEAGYFPVFINNLQRKVKQATKLFKGDPAGIFQEGSASRRMLTKLSLIFSHMLGELRALIPNGQDQGHQYQPSQPPAKAFWRGTWGARSLVSWSEFQAGLQRVHPMAPGPMAAALRATMDLTCSDHVSIFEFDIFTRLFQPWPTLLKNWTHLAVMHPGYVAFLTYDEVRARLQTYSNKPGSYVFRLSCTRLGQWAIGYVSQDGSILQTIPQDRPLFQALIEGHKEGFYLYPDGKNVNPDLTELTESTAQSRIEVSPDQAQLYSQMGSTFQLCKICAENDKDVRLQPCGHLLCRDCLNTWQQVQAPTCPFCRREIQGHEEIRIDPVGVQGAGAAPDEDDDHLEDVESVLRELAALRKVGGWGYQGGERGSGRAPLPRHRGCPVPRSSRWRLAGTGDNPGPFVAAASGATHEVKGQITEPAQGNEQPLPPPSTPTKQLQPTPLLGGGVKPRSPDTIPLPEPGRESRSSGSQPPCSIPAAPTSLPGLGQESRIPGFRVTRQKM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9K6</accession>
    <name>A0A452H9K6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="19">G-protein coupled receptors family 1 profile domain-containing protein</fullName>
        </recommendedName>
    </protein>
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        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
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            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="16">Receptor for R-spondins that potentiates the canonical Wnt signaling pathway and acts as a stem cell marker of the intestinal epithelium and the hair follicle. Upon binding to R-spondins (RSPO1, RSPO2, RSPO3 or RSPO4), associates with phosphorylated LRP6 and frizzled receptors that are activated by extracellular Wnt receptors, triggering the canonical Wnt signaling pathway to increase expression of target genes. In contrast to classical G-protein coupled receptors, does not activate heterotrimeric G-proteins to transduce the signal. Involved in the development and/or maintenance of the adult intestinal stem cells during postembryonic development.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Golgi apparatus</location>
            <location evidence="1">trans-Golgi network membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9K6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000012914.1">
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        <property type="gene ID" value="ENSGAGG00000008716.1"/>
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        <property type="component" value="Unassembled WGS sequence"/>
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    <dbReference type="GO" id="GO:0005886">
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        <property type="project" value="UniProtKB-SubCell"/>
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        <property type="term" value="F:G protein-coupled peptide receptor activity"/>
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        <property type="project" value="TreeGrafter"/>
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    <dbReference type="GO" id="GO:0016500">
        <property type="term" value="F:protein-hormone receptor activity"/>
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        <property type="project" value="InterPro"/>
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    <dbReference type="GO" id="GO:0007189">
        <property type="term" value="P:adenylate cyclase-activating G protein-coupled receptor signaling pathway"/>
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    <dbReference type="GO" id="GO:0090263">
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        <property type="entry name" value="leucine-rich repeat-containing G-protein coupled receptor 4 isoform X1"/>
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        <property type="entry name" value="Uncharacterized protein"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Rhodopsin 7-helix transmembrane proteins"/>
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        <property type="entry name" value="Ribonuclease Inhibitor"/>
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    </dbReference>
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    </dbReference>
    <proteinExistence type="predicted"/>
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    <keyword evidence="12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="10" id="KW-0297">G-protein coupled receptor</keyword>
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    <keyword evidence="9" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="4" id="KW-0433">Leucine-rich repeat</keyword>
    <keyword evidence="11 17" id="KW-0472">Membrane</keyword>
    <keyword evidence="13" id="KW-0675">Receptor</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="6 18" id="KW-0732">Signal</keyword>
    <keyword evidence="15" id="KW-0807">Transducer</keyword>
    <keyword evidence="5 17" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 17" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="18">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019242370" description="G-protein coupled receptors family 1 profile domain-containing protein" evidence="18">
        <location>
            <begin position="24"/>
            <end position="862"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="512"/>
            <end position="537"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="549"/>
            <end position="573"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="593"/>
            <end position="614"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="635"/>
            <end position="656"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="676"/>
            <end position="701"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
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            <begin position="722"/>
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        </location>
    </feature>
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            <begin position="757"/>
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        </location>
    </feature>
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        <location>
            <begin position="528"/>
            <end position="774"/>
        </location>
    </feature>
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        <source>
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        </source>
    </evidence>
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    </evidence>
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        </source>
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        </source>
    </evidence>
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        </source>
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    </evidence>
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        </source>
    </evidence>
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        </source>
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        </source>
    </evidence>
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        </source>
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        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="862" mass="95108" checksum="241BA6C0F216DCDE" modified="2019-05-08" version="1">MDTSCPASALLLLCFLGLPGASSGGLASPAGRWQRGCPTLCQCEQDGMLLRADCSDRGLTAVPTNLSLFTSYLDLSMNNITKLPSNPLHNLRFLEELRLDANHINYVPPNCFSGLVSLRHLWLDDNSLTEIPVQAFRSLSALQAMTLALNKIHHIPDYAFGNLSSLVVLHLHNNRIYSLGKKCFDGLHNLETLDLNYNSLDEFPIAIRTLTNLKELGFHSNNIKSIPEHAFVGNPSLITIHFYDNPIQLVGQSAFQHLPELRTLTLNSASQITEFPDLTGTTSLESLTLTGAQVTSLPKVACDQLPNLQVLDLSYNLLQDLPCFTACKKLQKIDLHHNEIYEIKANTFQQLVALRSLDLAWNKIAVIHPSAFSSLPSLIKLDLSSNLLSSFPVTGLHGLTHLKLTGNHDLQSLISSENFPELKVIEMPYAYQCCAFGTCESHYKISSQWNKEENSSADDFHKKDSGMLHIQDEHDFEDFLLDFEEDLKAHHSVQCTPSPGPFKPCDHLFGSWLIRIGVWTIVGLAFICNALLTATVFRSPLYISSIKLLLGLIAIVNMLMGLSSGVLASMDAFTFGSFAQYGAWWESGVGCQVTGLLSIFASETSIFLLTLAALERGFSVKYATKCKTKSSFASVKIAIFFCFMLALIIAVVPLLSESKYGVSPLCLPLPFGEPSSMGFMVALVLLNSLCFLVMTVAYTKLYCSLEKGELDNIWDCSMIKHIALLLFTNCILYCPVAFLSFSSLLNLTFISPEVIKSILLVIVPLPACLNPLLYILFNPHFKEDLGSLRKQTLLWSRSKHTSLISVNSDDVEKQSCDSTQALVTFASTSISYDVPASNSLMPSSYQTTESCNLSPVAFVPCR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H9K9</accession>
    <name>A0A452H9K9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="19">G-protein coupled receptors family 1 profile domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
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        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
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            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="16">Receptor for R-spondins that potentiates the canonical Wnt signaling pathway and acts as a stem cell marker of the intestinal epithelium and the hair follicle. Upon binding to R-spondins (RSPO1, RSPO2, RSPO3 or RSPO4), associates with phosphorylated LRP6 and frizzled receptors that are activated by extracellular Wnt receptors, triggering the canonical Wnt signaling pathway to increase expression of target genes. In contrast to classical G-protein coupled receptors, does not activate heterotrimeric G-proteins to transduce the signal. Involved in the development and/or maintenance of the adult intestinal stem cells during postembryonic development.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell membrane</location>
            <topology evidence="2">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Golgi apparatus</location>
            <location evidence="1">trans-Golgi network membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H9K9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000012905.1">
        <property type="protein sequence ID" value="ENSGAGP00000011266.1"/>
        <property type="gene ID" value="ENSGAGG00000008716.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
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        <property type="term" value="C:plasma membrane"/>
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        <property type="project" value="UniProtKB-SubCell"/>
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        <property type="term" value="F:protein-hormone receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
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        <property type="term" value="P:adenylate cyclase-activating G protein-coupled receptor signaling pathway"/>
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        <property type="term" value="P:positive regulation of canonical Wnt signaling pathway"/>
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    </dbReference>
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        <property type="entry name" value="7tmA_LGR5"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="leucine-rich repeat-containing G-protein coupled receptor 4 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Uncharacterized protein"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Rhodopsin 7-helix transmembrane proteins"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="GPCR_Rhodpsn"/>
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        <property type="entry name" value="GPCR_Rhodpsn_7TM"/>
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        <property type="entry name" value="Gphrmn_rcpt_fam"/>
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        <property type="entry name" value="Leu-rich_rpt"/>
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        <property type="entry name" value="Leu-rich_rpt_typical-subtyp"/>
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        <property type="entry name" value="LRR_5"/>
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        <property type="entry name" value="LRR_dom_sf"/>
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    <dbReference type="InterPro" id="IPR000372">
        <property type="entry name" value="LRRNT"/>
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    <dbReference type="PANTHER" id="PTHR24372">
        <property type="entry name" value="GLYCOPROTEIN HORMONE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24372:SF71">
        <property type="entry name" value="LEUCINE-RICH REPEAT-CONTAINING G-PROTEIN COUPLED RECEPTOR 5"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="7tm_1"/>
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        <property type="entry name" value="LRR_5"/>
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        <property type="entry name" value="LRR_8"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00373">
        <property type="entry name" value="GLYCHORMONER"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00237">
        <property type="entry name" value="GPCRRHODOPSN"/>
    </dbReference>
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        <property type="entry name" value="LRR_BAC"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SMART" id="SM00365">
        <property type="entry name" value="LRR_SD22"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00369">
        <property type="entry name" value="LRR_TYP"/>
        <property type="match status" value="12"/>
    </dbReference>
    <dbReference type="SMART" id="SM00013">
        <property type="entry name" value="LRRNT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81321">
        <property type="entry name" value="Family A G protein-coupled receptor-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52058">
        <property type="entry name" value="L domain-like"/>
        <property type="match status" value="2"/>
    </dbReference>
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        <property type="entry name" value="G_PROTEIN_RECEP_F1_2"/>
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    </dbReference>
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        <property type="entry name" value="LRR"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="10" id="KW-0297">G-protein coupled receptor</keyword>
    <keyword evidence="14" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="4" id="KW-0433">Leucine-rich repeat</keyword>
    <keyword evidence="11 17" id="KW-0472">Membrane</keyword>
    <keyword evidence="13" id="KW-0675">Receptor</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="6 18" id="KW-0732">Signal</keyword>
    <keyword evidence="15" id="KW-0807">Transducer</keyword>
    <keyword evidence="5 17" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 17" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="18">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5018974065" description="G-protein coupled receptors family 1 profile domain-containing protein" evidence="18">
        <location>
            <begin position="24"/>
            <end position="886"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="536"/>
            <end position="561"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="573"/>
            <end position="597"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="617"/>
            <end position="638"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="659"/>
            <end position="680"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="700"/>
            <end position="725"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="746"/>
            <end position="769"/>
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    </feature>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="781"/>
            <end position="801"/>
        </location>
    </feature>
    <feature type="domain" description="G-protein coupled receptors family 1 profile" evidence="19">
        <location>
            <begin position="552"/>
            <end position="798"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004166"/>
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        <source>
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    </evidence>
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        </source>
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        <source>
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        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00093766"/>
        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50262"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011266.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="886" mass="97857" checksum="23BE6C4BE73DC139" modified="2019-05-08" version="1">MDTSCPASALLLLCFLGLPGASSGGLASPAGRWQRGCPTLCQCEQDGMLLRADCSDRGLTAVPTNLSLFTSYLDLSMNNITKLPSNPLHNLRFLEELMLQNNQLKQVPTEALQNLRSLQSLRLDANHINYVPPNCFSGLVSLRHLWLDDNSLTEIPVQAFRSLSALQAMTLALNKIHHIPDYAFGNLSSLVVLHLHNNRIYSLGKKCFDGLHNLETLDLNYNSLDEFPIAIRTLTNLKELGFHSNNIKSIPEHAFVGNPSLITIHFYDNPIQLVGQSAFQHLPELRTLTLNSASQITEFPDLTGTTSLESLTLTGAQVTSLPKVACDQLPNLQVLDLSYNLLQDLPCFTACKKLQKIDLHHNEIYEIKANTFQQLVALRSLDLAWNKIAVIHPSAFSSLPSLIKLDLSSNLLSSFPVTGLHGLTHLKLTGNHDLQSLISSENFPELKVIEMPYAYQCCAFGTCESHYKISSQWNKEENSSADDFHKKDSGMLHIQDEHDFEDFLLDFEEDLKAHHSVQCTPSPGPFKPCDHLFGSWLIRIGVWTIVGLAFICNALLTATVFRSPLYISSIKLLLGLIAIVNMLMGLSSGVLASMDAFTFGSFAQYGAWWESGVGCQVTGLLSIFASETSIFLLTLAALERGFSVKYATKCKTKSSFASVKIAIFFCFMLALIIAVVPLLSESKYGVSPLCLPLPFGEPSSMGFMVALVLLNSLCFLVMTVAYTKLYCSLEKGELDNIWDCSMIKHIALLLFTNCILYCPVAFLSFSSLLNLTFISPEVIKSILLVIVPLPACLNPLLYILFNPHFKEDLGSLRKQTLLWSRSKHTSLISVNSDDVEKQSCDSTQALVTFASTSISYDVPASNSLMPSSYQTTESCNLSPVAFVPCR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJE0</accession>
    <name>A0A452HJE0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Voltage-dependent P/Q-type calcium channel subunit alpha</fullName>
        </recommendedName>
    </protein>
    <organism evidence="25 26">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="26" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="25" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="25" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="20">Voltage-sensitive calcium channels (VSCC) mediate the entry of calcium ions into excitable cells and are also involved in a variety of calcium-dependent processes, including muscle contraction, hormone or neurotransmitter release, gene expression, cell motility, cell division and cell death. The isoform alpha-1A gives rise to P and/or Q-type calcium currents.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>Ca(2+)(in) = Ca(2+)(out)</text>
            <dbReference type="Rhea" id="RHEA:29671"/>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="20">Membrane</location>
            <topology evidence="20">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="18">Belongs to the calcium channel alpha-1 subunit (TC 1.A.1.11) family. CACNA1A subfamily.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000017131.1">
        <property type="protein sequence ID" value="ENSGAGP00000014997.1"/>
        <property type="gene ID" value="ENSGAGG00000008380.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043025">
        <property type="term" value="C:neuronal cell body"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045202">
        <property type="term" value="C:synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="GOC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005891">
        <property type="term" value="C:voltage-gated calcium channel complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008331">
        <property type="term" value="F:high voltage-gated calcium channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098703">
        <property type="term" value="P:calcium ion import across plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007268">
        <property type="term" value="P:chemical synaptic transmission"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.238.10:FF:000063">
        <property type="entry name" value="Voltage-dependent N-type calcium channel subunit alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.350:FF:000011">
        <property type="entry name" value="Voltage-dependent N-type calcium channel subunit alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.350:FF:000013">
        <property type="entry name" value="Voltage-dependent N-type calcium channel subunit alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.70:FF:000023">
        <property type="entry name" value="Voltage-dependent R-type calcium channel subunit alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.70">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.2180">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.350">
        <property type="entry name" value="Voltage-gated potassium channels. Chain C"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005448">
        <property type="entry name" value="CACNA1A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031649">
        <property type="entry name" value="GPHH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005821">
        <property type="entry name" value="Ion_trans_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014873">
        <property type="entry name" value="VDCC_a1su_IQ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050599">
        <property type="entry name" value="VDCC_alpha-1_subunit"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002077">
        <property type="entry name" value="VDCCAlpha1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027359">
        <property type="entry name" value="Volt_channel_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45628">
        <property type="entry name" value="VOLTAGE-DEPENDENT CALCIUM CHANNEL TYPE A SUBUNIT ALPHA-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45628:SF3">
        <property type="entry name" value="VOLTAGE-DEPENDENT P_Q-TYPE CALCIUM CHANNEL SUBUNIT ALPHA-1A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08763">
        <property type="entry name" value="Ca_chan_IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16905">
        <property type="entry name" value="GPHH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00520">
        <property type="entry name" value="Ion_trans"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00167">
        <property type="entry name" value="CACHANNEL"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01632">
        <property type="entry name" value="PQVDCCALPHA1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01062">
        <property type="entry name" value="Ca_chan_IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81324">
        <property type="entry name" value="Voltage-gated potassium channels"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10 19 20" id="KW-0106">Calcium</keyword>
    <keyword evidence="6 20" id="KW-0107">Calcium channel</keyword>
    <keyword evidence="5 20" id="KW-0109">Calcium transport</keyword>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="15" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="16" id="KW-0407">Ion channel</keyword>
    <keyword evidence="13" id="KW-0406">Ion transport</keyword>
    <keyword evidence="14 22" id="KW-0472">Membrane</keyword>
    <keyword evidence="8 19" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="26" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0677">Repeat</keyword>
    <keyword evidence="23" id="KW-0732">Signal</keyword>
    <keyword evidence="7 22" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="12 22" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="2" id="KW-0813">Transport</keyword>
    <keyword evidence="11 20" id="KW-0851">Voltage-gated channel</keyword>
    <feature type="signal peptide" evidence="23">
        <location>
            <begin position="1"/>
            <end position="20"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019280087" description="Voltage-dependent P/Q-type calcium channel subunit alpha" evidence="23">
        <location>
            <begin position="21"/>
            <end position="1807"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="22">
        <location>
            <begin position="610"/>
            <end position="628"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="22">
        <location>
            <begin position="648"/>
            <end position="668"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="22">
        <location>
            <begin position="680"/>
            <end position="698"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="22">
        <location>
            <begin position="742"/>
            <end position="764"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="22">
        <location>
            <begin position="855"/>
            <end position="880"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="22">
        <location>
            <begin position="915"/>
            <end position="933"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="22">
        <location>
            <begin position="945"/>
            <end position="968"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="22">
        <location>
            <begin position="974"/>
            <end position="992"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="22">
        <location>
            <begin position="1036"/>
            <end position="1054"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="22">
        <location>
            <begin position="1127"/>
            <end position="1151"/>
        </location>
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    <feature type="domain" description="Voltage-dependent calcium channel alpha-1 subunit IQ" evidence="24">
        <location>
            <begin position="1288"/>
            <end position="1322"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="21">
        <location>
            <begin position="48"/>
            <end position="68"/>
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    </feature>
    <feature type="region of interest" description="Disordered" evidence="21">
        <location>
            <begin position="192"/>
            <end position="224"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="21">
        <location>
            <begin position="237"/>
            <end position="582"/>
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    <feature type="region of interest" description="Disordered" evidence="21">
        <location>
            <begin position="1324"/>
            <end position="1364"/>
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    <feature type="region of interest" description="Disordered" evidence="21">
        <location>
            <begin position="1385"/>
            <end position="1405"/>
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            <begin position="1449"/>
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    <feature type="region of interest" description="Disordered" evidence="21">
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            <begin position="1489"/>
            <end position="1807"/>
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            <begin position="53"/>
            <end position="63"/>
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            <begin position="204"/>
            <end position="217"/>
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            <begin position="237"/>
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    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="21">
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            <begin position="291"/>
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    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="21">
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            <begin position="346"/>
            <end position="387"/>
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        <location>
            <begin position="453"/>
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    <feature type="compositionally biased region" description="Pro residues" evidence="21">
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            <begin position="496"/>
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            <begin position="508"/>
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    <feature type="compositionally biased region" description="Polar residues" evidence="21">
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            <begin position="542"/>
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            <begin position="567"/>
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            <begin position="1536"/>
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        <location>
            <position position="826"/>
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            <dbReference type="ARBA" id="ARBA00022692"/>
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            <dbReference type="RuleBase" id="RU003808"/>
        </source>
    </evidence>
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            <dbReference type="SAM" id="MobiDB-lite"/>
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            <dbReference type="Ensembl" id="ENSGAGP00000014997.1"/>
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    <evidence type="ECO:0000313" key="26">
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    </evidence>
    <sequence length="1807" mass="206614" checksum="67D3D8E894EECB71" modified="2019-05-08" version="1">MLRAGSFCLFHSLFLPLSPFLPFQRRSCPSPWAFWLIFHHLFPTSRPPGNHAPLSTPQPQGQSHRTRRGELHATGARNRGGVDGDPAPLMLAHVCTCQDEQEEEEVANQKLALQKAKEVAEVSPLSAASMSIAVKEQQKNQKFAKSVWEQRTSEMRKHNLLASREALYSEMDPEDPWKVAYARAMRPDIKTHLDRPLVVDPQENRNNNTNRTRPNEPSLEQRFSQQRAEDFLRKQARYHERSREFSSSRSYEQPDAEGLEVRRQRRGSKETNLNQDGTYRELDYPSQGYESSREQGYRDSASERIKAVDPHRRHQHRQGGSKESRSGSPRMAEGERDHRRHRAQRRMGEEGGGSRDEGSKVERRLRHREGSRPSRRDVEGEQPDGERRRRHRHAAQSTYDTDGKRDDRERRHRRRKENQGPSPVPGPNLSTTRPIQQDMGRHEPQVAEDIDNMKNNKLATTEPSTPLNASHLQSPAKVGHHPPPNSQNAANRQVPSNPPQPGPPKTPENSLIVTNPSTQNNPAKAAKKPEHTTVDIPATFTPPINNTVVQVNKNANPDPLPKKEEEKKEEEDGEQEENGPKPMVPYSSMFILSTTNPFRRLCHYIVNLRYFEMCILMVIAMSSIALAAEDPVQPNATRNNVLRYFDYVFTGVFTFEMVIKMVDLGLVLHQGAYFRDLWNILDFIVVSGALVAFAFTGSSKGKDINTIKSLRVLRVLRPLKTIKRLPKLKAVFDCVVNSLKNVLNILIVYMLFMFIFAVVAVQLFKGKFFYCTDESKEFEKDCRGEYLVYEKDNEVKAQKREWKKYEFHYDNVLWALLTLFTVSTGEGWPQVLKHSVDATYENQGPSPGYRMEMSIFYVVYFVVFPFFFVNIFVALIIITFQEQGDKMMEEYSLEKNEQSFQYRMWQFVVSPPFEYTIMAMIALNTIVLMMKFDKASTAYENVLKMFNHVFTSLFSLECLLKIMAFGVLNYFRDAWNIFDFVTVLGSITDILVTEFGNNFINLSFLRLFRAARLIKLLRQGYTIRILLWTFVQSFKALPYVCLLIAMLFFIYAIIGMQVFGNIGIEDEEEDSAITEHNNFRTFFQALMLLFRSATGEAWHEIMLSCLSGKPCDENSGIKEHECGNEFAYFYFVSFIFLCSFLMLNLFVAVIMDNFEYLTRDSSILGPHHLDEYVRVWAEYDPAACGRIHYKDMYSLLRVISPPLGLGKKCPHRVACKRLLRMDLPVADDNTVHFNSTLMALIRTALDIKIAKGGADKQQMDAELRKEMMAIWPNLSHKNLDLLVTPHKSTDLTVGKIYAAMMIMEYYRQSKAKKLQAMREEQNRTPLMFQRMEPPSPTQEGVPGQDALPSSQLDQGGGLSAPEGGIKESQSWVTERAQAMFQKTGTWSPERGHAEDMPNSRPNSQSVEMREIGKDGYSDSDHYLPMEGHGRAASMPRLPAENQVRGYFFEPSRDSTTITDASPMKRSASMLGHSRSRGIRLDDYSLERVVPEENQRHHQRRRERSHRASERSLSRYADVDTGLGTDLSMTTPLGDLPPKERDQERGRPKDRKHHHHHHHHRHHHPSSDKERHEHSRPRSRDRRWSRSPSEGREHVPHRQGSSSVSGSPVLSTSGTSTPRRGRRQLPQTPSTPRPHVSYSPVVRKVISTGPQQPPQSVHHPSRISTPVPRRYPLLPAEPQVVERPGRSPAMERHTVPLRNDSPRAYRHASSRWSAPHVSEPHPGPRRGGYYRSPDCAAPPSEDAFSYETTHISSSGRSPRTSRAGGGSSPSRPGRRLPNGYFHSHGAAKPRSRHGLHEPSSETDEDDCC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJE3</accession>
    <name>A0A452HJE3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Cytochrome P450 family 4 subfamily B member 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="9">
            <name>heme</name>
            <dbReference type="ChEBI" id="CHEBI:30413"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the cytochrome P450 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJE3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017168.1">
        <property type="protein sequence ID" value="ENSGAGP00000015029.1"/>
        <property type="gene ID" value="ENSGAGG00000011253.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0020037">
        <property type="term" value="F:heme binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005506">
        <property type="term" value="F:iron ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004497">
        <property type="term" value="F:monooxygenase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016705">
        <property type="term" value="F:oxidoreductase activity, acting on paired donors, with incorporation or reduction of molecular oxygen"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd20678">
        <property type="entry name" value="CYP4B-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.630.10:FF:000005">
        <property type="entry name" value="cytochrome P450 4F22 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.630.10">
        <property type="entry name" value="Cytochrome P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001128">
        <property type="entry name" value="Cyt_P450"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002401">
        <property type="entry name" value="Cyt_P450_E_grp-I"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036396">
        <property type="entry name" value="Cyt_P450_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050196">
        <property type="entry name" value="Cytochrome_P450_Monoox"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24291">
        <property type="entry name" value="CYTOCHROME P450 FAMILY 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24291:SF201">
        <property type="entry name" value="CYTOCHROME P450, FAMILY 4, SUBFAMILY B, POLYPEPTIDE 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00067">
        <property type="entry name" value="p450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00463">
        <property type="entry name" value="EP450I"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00385">
        <property type="entry name" value="P450"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48264">
        <property type="entry name" value="Cytochrome P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="3 9" id="KW-0349">Heme</keyword>
    <keyword evidence="6 9" id="KW-0408">Iron</keyword>
    <keyword evidence="8 10" id="KW-0472">Membrane</keyword>
    <keyword evidence="4 9" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7" id="KW-0503">Monooxygenase</keyword>
    <keyword evidence="7" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="10" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="6"/>
            <end position="25"/>
        </location>
    </feature>
    <feature type="binding site" description="axial binding residue" evidence="9">
        <location>
            <position position="427"/>
        </location>
        <ligand>
            <name>heme</name>
            <dbReference type="ChEBI" id="CHEBI:30413"/>
        </ligand>
        <ligandPart>
            <name>Fe</name>
            <dbReference type="ChEBI" id="CHEBI:18248"/>
        </ligandPart>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004586"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023033"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PIRSR" id="PIRSR602401-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015029.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="13">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="480" mass="55205" checksum="EFDA20AA8E53830B" modified="2019-05-08" version="1">MTTPSSLLLYPTAGFCLIYVLLKAIQLYRKRQQLLKAFESFPGPPSHWLYGHSSQISQGGELNQLLSWAKEYPYAYPRWIGGFMASLTVNHPDYAKTVFGHGGKGLLILHGPKWFQHRRLLTPGFHYDVLKPYVSLMAESTKVMLDKWEWLITVDKSVELFEHVSLMTLDSIMKCAFSCHSNCQTDRDSDYYIKAVYDLTYLVFHRIRFFPFHNDFIYRISSQGRRFQEACRLAHQHTDKVIKERKTSLQDQQELEKIQKKKYLDFLDILLCAKDENGAGLSDEDLRAEVDTFMFEGHDTTASGVSWLLYCMALHPEHQQRCREEIKEILGDRETIQWDDLGKMTYSTMCIKESLRLYPPVPGVSRLLSKPITFHDGRTVPEGLFSLFLLKGPILPSVFDPMRFSPENVSKRHSHAFLPFAAGARNCIGKQFAMNELKVALALTLLRFELAPDPAKPPKEIPQIVLKSSNGIHLLLRKLP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJE5</accession>
    <name>A0A452HJE5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Ig-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJE5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017136.1">
        <property type="protein sequence ID" value="ENSGAGP00000015002.1"/>
        <property type="gene ID" value="ENSGAGG00000011340.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009897">
        <property type="term" value="C:external side of plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042288">
        <property type="term" value="F:MHC class I protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0002323">
        <property type="term" value="P:natural killer cell activation involved in immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00096">
        <property type="entry name" value="Ig"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015631">
        <property type="entry name" value="CD2/SLAM_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024303">
        <property type="entry name" value="NK_rcpt_2B4_Ig_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12080:SF56">
        <property type="entry name" value="NATURAL KILLER CELL RECEPTOR 2B4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12080">
        <property type="entry name" value="SIGNALING LYMPHOCYTIC ACTIVATION MOLECULE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13895">
        <property type="entry name" value="Ig_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11465">
        <property type="entry name" value="Receptor_2B4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="3" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 5" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="5">
        <location>
            <begin position="1"/>
            <end position="18"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019507406" description="Ig-like domain-containing protein" evidence="5">
        <location>
            <begin position="19"/>
            <end position="247"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="6">
        <location>
            <begin position="125"/>
            <end position="213"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004370"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015002.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="247" mass="27357" checksum="93D217F62EB45CFF" modified="2019-05-08" version="1">VNDAQALCVLLWLPASRGVDVGGKWESVQLLPLKLPEPWAEITWEVTLDSAETYRIVTFNKGDPDHGSSPHFINRVTFYPENLSLQIDPVKKSDSGLYSLQIETGGGLIDITKFHVSVFNRVRQPNLTVLSTHPKLGQCNVTLSCSVPGADRVTYSWSQGTSRIPSDRDHQLPGNQSLLHVEINANSSDIFYRCNASNRASWATDAADGKSLCHSPLLKGLISFWGVLTGASYVRHERSLSRSFDHR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJE6</accession>
    <name>A0A452HJE6_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="13">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="9">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000017173.1">
        <property type="protein sequence ID" value="ENSGAGP00000015032.1"/>
        <property type="gene ID" value="ENSGAGG00000008780.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00054">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="12"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000005">
        <property type="entry name" value="Fibrillin 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000194">
        <property type="entry name" value="Latent transforming growth factor beta binding protein 2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000469">
        <property type="entry name" value="Latent transforming growth factor beta binding protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.290.10:FF:000004">
        <property type="entry name" value="latent-transforming growth factor beta-binding protein 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000019">
        <property type="entry name" value="latent-transforming growth factor beta-binding protein 1 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000014">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000077">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 3 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.290.10:FF:000001">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 3 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.290.10:FF:000002">
        <property type="entry name" value="Latent-transforming growth factor beta-binding protein 3 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000115">
        <property type="entry name" value="latent-transforming growth factor beta-binding protein 4 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000024">
        <property type="entry name" value="Putative latent-transforming growth factor beta-binding protein 2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000273">
        <property type="entry name" value="Putative latent-transforming growth factor beta-binding protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="16"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.290.10">
        <property type="entry name" value="TGF-beta binding (TB) domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026823">
        <property type="entry name" value="cEGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001881">
        <property type="entry name" value="EGF-like_Ca-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013032">
        <property type="entry name" value="EGF-like_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000152">
        <property type="entry name" value="EGF-type_Asp/Asn_hydroxyl_site"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018097">
        <property type="entry name" value="EGF_Ca-bd_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009030">
        <property type="entry name" value="Growth_fac_rcpt_cys_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049883">
        <property type="entry name" value="NOTCH1_EGF-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017878">
        <property type="entry name" value="TB_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036773">
        <property type="entry name" value="TB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052080">
        <property type="entry name" value="vWF_C/EGF_Fibrillin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47333:SF4">
        <property type="entry name" value="EGF-LIKE DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47333">
        <property type="entry name" value="VON WILLEBRAND FACTOR C AND EGF DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12662">
        <property type="entry name" value="cEGF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07645">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="12"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12661">
        <property type="entry name" value="hEGF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00683">
        <property type="entry name" value="TB"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="17"/>
    </dbReference>
    <dbReference type="SMART" id="SM00179">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="15"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57184">
        <property type="entry name" value="Growth factor receptor domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57581">
        <property type="entry name" value="TB module/8-cys domain"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00010">
        <property type="entry name" value="ASX_HYDROXYL"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00022">
        <property type="entry name" value="EGF_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01186">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01187">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51364">
        <property type="entry name" value="TB"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7 9" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="4 9" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="3" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="8" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <feature type="domain" description="EGF-like" evidence="11">
        <location>
            <begin position="60"/>
            <end position="92"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="11">
        <location>
            <begin position="258"/>
            <end position="290"/>
        </location>
    </feature>
    <feature type="domain" description="TB" evidence="12">
        <location>
            <begin position="405"/>
            <end position="448"/>
        </location>
    </feature>
    <feature type="domain" description="TB" evidence="12">
        <location>
            <begin position="525"/>
            <end position="577"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="11">
        <location>
            <begin position="685"/>
            <end position="726"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="11">
        <location>
            <begin position="727"/>
            <end position="763"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="11">
        <location>
            <begin position="770"/>
            <end position="809"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="11">
        <location>
            <begin position="850"/>
            <end position="890"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="11">
        <location>
            <begin position="975"/>
            <end position="1016"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="11">
        <location>
            <begin position="1059"/>
            <end position="1099"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="11">
        <location>
            <begin position="1144"/>
            <end position="1185"/>
        </location>
    </feature>
    <feature type="domain" description="TB" evidence="12">
        <location>
            <begin position="1247"/>
            <end position="1299"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="11">
        <location>
            <begin position="1321"/>
            <end position="1359"/>
        </location>
    </feature>
    <feature type="domain" description="TB" evidence="12">
        <location>
            <begin position="1421"/>
            <end position="1473"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1"/>
            <end position="31"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="105"/>
            <end position="132"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="375"/>
            <end position="400"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="598"/>
            <end position="621"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1527"/>
            <end position="1562"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="8"/>
            <end position="18"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="375"/>
            <end position="399"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="1550"/>
            <end position="1562"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="9">
        <location>
            <begin position="64"/>
            <end position="74"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="9">
        <location>
            <begin position="82"/>
            <end position="91"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="9">
        <location>
            <begin position="262"/>
            <end position="272"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="9">
        <location>
            <begin position="280"/>
            <end position="289"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004498"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS51364"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015032.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1608" mass="173523" checksum="30796562B4644424" modified="2019-05-08" version="1">KLSAPIHRSQSGASSQPCSPAGAGQARDPTPQRARLKCLEDQIQPLGWSHVSLFTPFHVPPAVCDPPCQNKGSCSRPQLCICRSGFQGAQCEEVVPERVYHPPGSVAALQPPAGSLPKRAGSAEREGSPRGAQTPILKQALAGTQGRIVFPVSRHTGSSRTVRRYPAGNTQLMSNALPNGNGHEQSSSGPHAAYQEHTLSLWGANLTEKIRKIKIVFTPTICKQTCRSGHCYNSCQKGDTTTLYSQGGQNHDPKSGFRIYFCQIPCLNGGRCVGRDECWCPSNSTGKFCHLPVPKLEKNQVGRVPKTLASGSIKQSMYTLPLSNQLASLNPSLVNVHIKHPPEATVQIHQVARVRGDLGQSEENSVEAVLLSQLPTGPQHSYSNGNSNSITTESGQQQRPPELMGRCFRETLHGQCANPLEGLAKFEDCCGSVGLFWGVNRCIPCPPRPAHPVIENRQVECPQGYKRLNRSHCQDINECLMLGLCKDTECVNTRGSFLCTCKPGTMLDPSRSRCVSDKAVSMEQGLCYRSAAEGVCALPLAQSITKQICCCSRVGKGWGRNCEKCPMLGSETFKEICPAGHGYTYSSSDIRLSMRKAEAEELPFSSEEQGESSDQTPTQAAKRLQLQEALSGGVTGTLARAGTSAAEGVLGCDATLVAGTADATQEPAPHPGQWPVTSPCLSLSEMDRCASVPDPCGPGTCVNLPDRYSCICGPGYQLHPGLPQCIDDDECMRDPCAGKGHCINSVGSYSCLCYPGYTLLASQGTQTCQDLNECEQPSMCQGGQCTNTPGSYHCECARGYIMGHTGQCEDIDECADPSTCPRGRCVNALGSYECLSCAAGYQPMNGRCVDDDECFTVGICAHGLCINLAGSFRCSCYHGYEVTADEKSCQDINECATHNACPMGLCINSEGSFSCAACGAGYTVSRDGSMCEDIDECSSATACPLGICTNTDGSFACRACDAGYMLSSNRFTCEDMNECLASEICSPNGECLNSHGSYFCICAPGFSNAAGGVSCQGEQLGQMAAAKCALDPGYLFIVQGPVYTHRTASVLIFILSLPDVDECADKSRCLRGQCFNTEGSYRCLCENGFRHSQETDDCVDVDECVDFGDTVCGTWRCQNTPGSYRCVMGCQPGFHRTPLGDCIDIDECANETLCGSHAFCDNTDGSFRCLCDSGYENLPPGQDCVDVNECELMVAVCGTALCENVEGSFLCLCLSDHEEYNTEAGECRPRAGMGTGMAPRPAGSERKECYYHISDVQLCDSVLAKNTTKEECCCTEGAAWGDNCETHPCPILVEYSEICPSGKGYIPVEGTLLFGQTSYTDADECEMFGPEICRNGHCVNTVPGYKCFCRSGYLYDSSRLECVDQDECENEMSCVNGECLNTAGSFHCFCGLPLVLDVTRNRCINVSSRAEDPEEHEIHLDICWQRVSNYICNQPLRGRRTTYTECCCRYGEAWSQDCALCPHRASEDFAHLCNVARGETEREPGLQERPGYEYGPALEDSHYGLYSPDLGPYYNYLSPEYGAPDASFPQRELSSEVGGSTFLRTPQLPPAQSQPRYLPSQTGEGLQTLCTFPNPCSHNQAASPDRLACGQELETGFRENGACSRGDS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJF0</accession>
    <name>A0A452HJF0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">3-beta hydroxysteroid dehydrogenase/isomerase domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="2">Belongs to the 3-beta-HSD family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJF0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017141.1">
        <property type="protein sequence ID" value="ENSGAGP00000015007.1"/>
        <property type="gene ID" value="ENSGAGG00000010832.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016616">
        <property type="term" value="F:oxidoreductase activity, acting on the CH-OH group of donors, NAD or NADP as acceptor"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006694">
        <property type="term" value="P:steroid biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.720:FF:000495">
        <property type="entry name" value="3 hydroxysteroid dehydrogenase, putative"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.720">
        <property type="entry name" value="NAD(P)-binding Rossmann-like Domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002225">
        <property type="entry name" value="3Beta_OHSteriod_DH/Estase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036291">
        <property type="entry name" value="NAD(P)-bd_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050425">
        <property type="entry name" value="NAD(P)_dehydrat-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10366:SF847">
        <property type="entry name" value="3 BETA-HYDROXYSTEROID DEHYDROGENASE TYPE 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10366">
        <property type="entry name" value="NAD DEPENDENT EPIMERASE/DEHYDRATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01073">
        <property type="entry name" value="3Beta_HSD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51735">
        <property type="entry name" value="NAD(P)-binding Rossmann-fold domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51257">
        <property type="entry name" value="PROKAR_LIPOPROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="1 2" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="3-beta hydroxysteroid dehydrogenase/isomerase" evidence="3">
        <location>
            <begin position="25"/>
            <end position="295"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="RuleBase" id="RU004475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="Pfam" id="PF01073"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015007.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="384" mass="42386" checksum="3741565444C207BD" modified="2019-05-08" version="1">MLFGGRRAREGGRAGTQESGTTVYLVTGGCGFLGSHLVRMLVERSPNVAEVRVFDLQLDPALGQLGTGEVKVTLILGDISSPTDVGAAVKGADVVIHAASLVDVWGRVPPEKITEVNVHGTKNVIDACIEHGTQYLIYTSSMEVVGPNIKGDPFYRGNEDSAYDAQHEQPYPLSKAQAERMVIEANGTPVRGGRWLVTCALRPTGIYGEKHPLMKEFYEKGLQTGRRLLRAIPVSTEHGRVYVGNVAWMHVLVARQMQEAPAAIGGQVYFCYDASPYKSYEDFNMEILASCGFRLLGSQPVVPHWLLRCLALANDLLRRLLRPFFSYAPLLNPYTLAVASTTFTVQTDKAQRHFGYQPLYSWEESRARTVAWLQEVDAQREAGK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJF1</accession>
    <name>A0A452HJF1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">U2 snRNP-associated SURP motif-containing protein</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="12">140 kDa Ser/Arg-rich domain protein</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="13">U2-associated protein SR140</fullName>
        </alternativeName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="10">Interacts with ERBB4.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the splicing factor SR family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJF1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017182.1">
        <property type="protein sequence ID" value="ENSGAGP00000015037.1"/>
        <property type="gene ID" value="ENSGAGG00000010747.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006396">
        <property type="term" value="P:RNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd21370">
        <property type="entry name" value="cwf21_SR140"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12223">
        <property type="entry name" value="RRM_SR140"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.790:FF:000006">
        <property type="entry name" value="U2 snRNP-associated SURP motif-containing protein isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.90:FF:000013">
        <property type="entry name" value="U2 snRNP-associated SURP motif-containing protein isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.330:FF:000177">
        <property type="entry name" value="U2 snRNP-associated SURP motif-containing protein-like isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.90">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.330">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.420">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.790">
        <property type="entry name" value="Surp module"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006569">
        <property type="entry name" value="CID_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008942">
        <property type="entry name" value="ENTH_VHS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013170">
        <property type="entry name" value="mRNA_splic_Cwf21_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012677">
        <property type="entry name" value="Nucleotide-bd_a/b_plait_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035979">
        <property type="entry name" value="RBD_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000504">
        <property type="entry name" value="RRM_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051485">
        <property type="entry name" value="SR-CTD_assoc_factor"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047488">
        <property type="entry name" value="SR140_cwf21"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035009">
        <property type="entry name" value="SR140_RRM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000061">
        <property type="entry name" value="Surp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035967">
        <property type="entry name" value="SWAP/Surp_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23140">
        <property type="entry name" value="RNA PROCESSING PROTEIN LD23810P"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23140:SF0">
        <property type="entry name" value="U2 SNRNP-ASSOCIATED SURP MOTIF-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04818">
        <property type="entry name" value="CID"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08312">
        <property type="entry name" value="cwf21"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00076">
        <property type="entry name" value="RRM_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01805">
        <property type="entry name" value="Surp"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01115">
        <property type="entry name" value="cwf21"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00582">
        <property type="entry name" value="RPR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00360">
        <property type="entry name" value="RRM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00648">
        <property type="entry name" value="SWAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48464">
        <property type="entry name" value="ENTH/VHS domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54928">
        <property type="entry name" value="RNA-binding domain, RBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109905">
        <property type="entry name" value="Surp module (SWAP domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51391">
        <property type="entry name" value="CID"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="RRM"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="SURP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0007">Acetylation</keyword>
    <keyword evidence="8" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="9" id="KW-0539">Nucleus</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6 14" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="5" id="KW-0832">Ubl conjugation</keyword>
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        <location>
            <begin position="186"/>
            <end position="267"/>
        </location>
    </feature>
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        <location>
            <begin position="342"/>
            <end position="385"/>
        </location>
    </feature>
    <feature type="domain" description="CID" evidence="18">
        <location>
            <begin position="446"/>
            <end position="591"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="52"/>
            <end position="185"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="695"/>
            <end position="754"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="768"/>
            <end position="942"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="15">
        <location>
            <begin position="9"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="15">
        <location>
            <begin position="56"/>
            <end position="67"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="15">
        <location>
            <begin position="79"/>
            <end position="91"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="15">
        <location>
            <begin position="99"/>
            <end position="135"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="15">
        <location>
            <begin position="152"/>
            <end position="162"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="15">
        <location>
            <begin position="698"/>
            <end position="718"/>
        </location>
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    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="15">
        <location>
            <begin position="722"/>
            <end position="739"/>
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    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="15">
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        <location>
            <begin position="863"/>
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        </location>
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    <feature type="compositionally biased region" description="Basic residues" evidence="15">
        <location>
            <begin position="904"/>
            <end position="942"/>
        </location>
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        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
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    </evidence>
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        <source>
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    </evidence>
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            <dbReference type="ARBA" id="ARBA00022884"/>
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    </evidence>
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    </evidence>
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    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00067522"/>
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        <source>
            <dbReference type="ARBA" id="ARBA00083599"/>
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    </evidence>
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        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00176"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
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    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50102"/>
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        <source>
            <dbReference type="PROSITE" id="PS50128"/>
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    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS51391"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015037.1"/>
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    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
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    </evidence>
    <sequence length="942" mass="108898" checksum="D7F3804FA110851A" modified="2019-05-08" version="1">MSAAKRTLSKKEQEELKKKEDEKAAAEIYEEFLAAFEGSDGNKVKTFVRGGIVNASKDEHETDEKRGKIYKPSARFADQKNQPNQPSSNERPPSLLMIETKKPPLKKGEKEKKKSNLELFKEELKQIQEERDERHKTKGRLSRFEPPQSDSDGQRRSMDAPSRRNRSSVLDDYAPGSHDVGDPSTTNLYLGNINPQMNEEMLCQEFGRFGPLASVKIMWPRTDEERARERNCGFVAFMNRRDAERALKNLNGKMIMSFEMKLGWGKAVPIPPHPIYIPPSMMEHTLPPPPSGLPFNAQPRERLKNPNAPMLPPPKNKEDFEKTLSQAIVKVVIPTERNLLALIHRMIEFVVREGPMFEAMIMNREINNPMFRFLFENQTPAHVYYRWKLYSILQGDAPTKWRTEDFRMFKNGSFWRPPPLNPYLHGMSEEQETEAFVEEPNKKGALKEEQRDKLEEILRGLTPRKNDIGDAMVFCLNNAEAAEEIVDCVTESLSILKTPLPKKIARLYLVSDVLYNSSAKVANASYYRKFFETKLCQIFSDLNATYRTIQGHLQSENFKQRVMTCFRAWEDWAIYPEPFLIKLQNIFLGLVNIIEEKEIEEVPDDLDGAPIEEELDGAPLEDVDGIPIDAAPIDDLDGIPIKSLDDDLDGIPLDTSEESKKNEPIFKVAPSKWEAVDEAELEAQAVTTSKWELFDQHEESEEEENQNQEEESEDEEDTQSSKSEEQHMYSNPIKEEMPESKSSIKYSEMSEEKRAKLREIELKVMKFQDELESGKRPKKPGQSFQEQVEHYRDKLLQREKEKELERERERDKKDKEKSESRSKDKKEKEECTPTRKERKRRHSTSPSPSRSSGSRRAKSPSPKSERSERSERSHKETSRSRSSHKDSPRDVSKKAKRSPSGSRTPKRSRRSRSRSPKKSGKKSRSQSRSPHRSHKKSKKSKH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJF6</accession>
    <name>A0A452HJF6_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="8">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000017190.1">
        <property type="protein sequence ID" value="ENSGAGP00000015044.1"/>
        <property type="gene ID" value="ENSGAGG00000011031.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046982">
        <property type="term" value="F:protein heterodimerization activity"/>
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        <property type="project" value="InterPro"/>
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        <property type="term" value="P:Ras protein signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="entry name" value="HFD_SOS1_rpt1"/>
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    </dbReference>
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        <property type="entry name" value="HFD_SOS1_rpt2"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="PH_SOS"/>
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    </dbReference>
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        <property type="entry name" value="REM"/>
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        <property type="entry name" value="Son of sevenless homolog 1 (Drosophila)"/>
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    </dbReference>
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        <property type="entry name" value="son of sevenless homolog 1 isoform X1"/>
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    </dbReference>
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        <property type="entry name" value="Dbl homology (DH) domain"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Histone, subunit A"/>
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        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
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        <property type="entry name" value="Ras guanine-nucleotide exchange factors catalytic domain"/>
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        <property type="entry name" value="Son of sevenless (SoS) protein Chain: S domain 1"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Histone-fold"/>
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        <property type="entry name" value="PH-like_dom_sf"/>
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        <property type="entry name" value="Ras-like_GEF"/>
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        <property type="entry name" value="Ras_G-nucl-exch_fac_CS"/>
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        <property type="entry name" value="Ras_GEF_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001895">
        <property type="entry name" value="RASGEF_cat_dom"/>
    </dbReference>
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        <property type="entry name" value="RASGEF_cat_dom_sf"/>
    </dbReference>
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        <property type="entry name" value="SOS1_NGEF_PH"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23113">
        <property type="entry name" value="GUANINE NUCLEOTIDE EXCHANGE FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23113:SF150">
        <property type="entry name" value="SON OF SEVENLESS HOMOLOG 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00617">
        <property type="entry name" value="RasGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="RasGEF_N"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="RhoGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22697">
        <property type="entry name" value="SOS1_NGEF_PH"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="RasGEF"/>
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    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="DBL homology domain (DH-domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47113">
        <property type="entry name" value="Histone-fold"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48366">
        <property type="entry name" value="Ras GEF"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="DH_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="RASGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50009">
        <property type="entry name" value="RASGEF_CAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50212">
        <property type="entry name" value="RASGEF_NTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1 2" id="KW-0344">Guanine-nucleotide releasing factor</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="DH" evidence="6">
        <location>
            <begin position="198"/>
            <end position="293"/>
        </location>
    </feature>
    <feature type="domain" description="PH" evidence="4">
        <location>
            <begin position="408"/>
            <end position="511"/>
        </location>
    </feature>
    <feature type="domain" description="N-terminal Ras-GEF" evidence="7">
        <location>
            <begin position="562"/>
            <end position="706"/>
        </location>
    </feature>
    <feature type="domain" description="Ras-GEF" evidence="5">
        <location>
            <begin position="744"/>
            <end position="983"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="984"/>
            <end position="1065"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="1107"/>
            <end position="1298"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="3">
        <location>
            <begin position="1044"/>
            <end position="1064"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="3">
        <location>
            <begin position="1139"/>
            <end position="1151"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="3">
        <location>
            <begin position="1168"/>
            <end position="1188"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="1208"/>
            <end position="1217"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="1276"/>
            <end position="1285"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022658"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00168"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="PROSITE" id="PS50009"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50010"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015044.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1298" mass="149736" checksum="00D705D10A600334" modified="2019-05-08" version="1">MQPPPQPYEFFSEENGPKWRGLFVPALRKVQQQVHPSFSAKEDSLYYIEELILQLLNKLCIAQPRTVQDVEERVQKTFPHPIDKWAIADAQSAIEKRKRRNPLLLPVDKIHPLLKEVLGYKVDYHVSLYIVAVLEYISADILKLAGNYVFNIRHFEISQQDIKVSMCADKVLMDMFDQDVIGLVSLCEDEPSSSGELNYYELVRTEIAEERQYLRELNLIIKVFREAFLSNRKLFTPYDIDTIFSNISDIHELTVKLLGLIEDTVEMTDESSPHPLAGSCFEDLAEEQAFDPYETLSQDILSPRFHEHFNNLMAKPAVALHFQQLQECSEDEEDRECLKQAITALLNLQCSMERIYNKYSPRRRPWETICRFYNRQVRSKHLAIKKMNEIQKNIDGWEGKDIGQCCNEFIMEGALTRIGAKHERHIFLFDGLLISCKTNHGQSRLPGYSNTEYRLKEKIIMRKMQIIDKDDTCEYKHAFELASKDENSLVFAAKSAEEKNNWMAALISLQYRSTLDRMLDSVLLQEENEQPLRLPSPNVYRFVVEDSEENIVFEDNLQSRNGIPIIKGGTVVKLIERLTYHMYADPNFVRTFLTTYRSFCKPQELLSLLIERFEIPEPEPTEADRLAIEKGEQPISADLKRFRKEYVQPVQLRILNVFRHWVEHHFYDFERDLELLDKLETFISSVRGKSMKKWVESIAKIIKRKKQAQANGISHNITFESQPPPIEWHICHLGQFEMDLMTLHPIEIARQLTLLESDLYRAVQPSELVGSVWTKEDKEINSPNLLKMIRHTTNLTLWFEKCIVETENFEERVAVLSRIIEILQVFQELNNFNGVLEIVSAMNSVSVYRLDHTFEALQERKKKVLDEAVELSQDHFKKYLAKLKSINPPCVPFFGIYLTNILKTEEGNPDFLKRQGKELINFSKRRKVAEITGEIQQYQNQPYCLRIEPEIRRFFENLNPMGNAPEKEFTDYLFNKSLEIEPRNCKQPPRFPRKTTYPLKSPGIRPNTGRHCSTSGTLRGHPTPLEREPCKISFSRITEAEQESAASAPTSPNTPSTPPVSASSERSVFLDIDLNSSYGNNSIFAPVHLPQSKSFFSSCGSIHKLSEEPLVPPPLPPRKKFDQEASNSKGSLKSDDDPPAIPPRQPPPPKIQPRVPAYNNPFDQLLHSPPPPPPRDPLPDTPPPVPLRPPEHFINCPFNLQPPPVGRFHRDPDRFREASTCPSSPNTPPSTPSPRVLRRCYMLSSSHNNVAHPPAPPVPPRQNSSPQLPKLPPKTYKRELSHPPLDRQFLLENAETPQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJF7</accession>
    <name>A0A452HJF7_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="11">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="8">Presynapse</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJF7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017191.1">
        <property type="protein sequence ID" value="ENSGAGP00000015045.1"/>
        <property type="gene ID" value="ENSGAGG00000009356.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030424">
        <property type="term" value="C:axon"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048788">
        <property type="term" value="C:cytoskeleton of presynaptic active zone"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098882">
        <property type="term" value="F:structural constituent of presynaptic active zone"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007274">
        <property type="term" value="P:neuromuscular synaptic transmission"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048167">
        <property type="term" value="P:regulation of synaptic plasticity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.1490">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019323">
        <property type="entry name" value="ELKS/CAST"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18861">
        <property type="entry name" value="ELKS/RAB6-INTERACTING/CAST PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18861:SF3">
        <property type="entry name" value="ERC PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10174">
        <property type="entry name" value="Cast"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57997">
        <property type="entry name" value="Tropomyosin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0966">Cell projection</keyword>
    <keyword evidence="5 9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0770">Synapse</keyword>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1"/>
            <end position="44"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="147"/>
            <end position="174"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="211"/>
            <end position="276"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="332"/>
            <end position="846"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="1"/>
            <end position="13"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="14"/>
            <end position="25"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
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        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
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        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00034106"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015045.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="853" mass="98547" checksum="A659C3202ABEC46D" modified="2019-05-08" version="1">MYGSARTISNLEGSPSRSPRLPRSPRLGHRRTSSGGGGGTGKTLSMENIQSLNAAYATSGPMYLSDHEGVASTTFPKGTMTLGRATNRAVYGGRVTAMGSSPNIASAGLSHTDVLSYTDQHGGLTTSSHHHHHQVPSMLRQVRDSTMLDLQAQLKELQRENDLLRKELDIKDSKLGSSMNSIKTFWSPELKKERVLRKEEAARMSVLKEQMRVSHEENQHLQLTIQALQDELRTQRDLNHLLQQESGNRGTEHFTIELTEENFRRLQAEHDRQAKELFLLRKTLEEMELRIETQKQTLNARDESIKKLLEMLQSKGLPSKGMEDDNERARRMAEAESQVNHLEVILDQKEKENIHLREELHRRTQLQPEPAKTKALQTVIEMKDTKIASLERNIRDLEDEIQMLKTNGVLNIEDREEEIKQIEVYKSHSKFMKNKIDQLKQELSKKESELLALQTKLETLSNQNSDCKQHIEVLKESLTAKEQRAAILQTEVDALRLRLEEKETFLNKKTKQLQDLTEEKGTLAGEIRDMKDMLEVKERKINVLQKKIENLQEQLRDKDKQLSNLKDRVKSLQTDSSNTDTALATLEEALSEKERIIERLKEQRERDDRERLEEIESFKKENKDLKEKVNALQAELTEKESNLIDLKEHASSLASTGLKRDSKLKSLEIAIEQKKEECSKLEAQLKKAHEAEEEARLNPEFADRMKQLDKEASYYREECGKAQAEVDRLLEILKEVENEKNDKDKKIAELERHVKDQNKKVANLKHNQQMEKKKNVQLLEEVRRREDNMTDNSQHLQLEELMNALEKTRQELDATKARLASTQQSLAEKEAHLANLRIERRKQLEEILEMNIP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJG2</accession>
    <name>A0A452HJG2_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="7">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HJG2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017193.1">
        <property type="protein sequence ID" value="ENSGAGP00000015047.1"/>
        <property type="gene ID" value="ENSGAGG00000010747.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006396">
        <property type="term" value="P:RNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd21370">
        <property type="entry name" value="cwf21_SR140"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12223">
        <property type="entry name" value="RRM_SR140"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.790:FF:000006">
        <property type="entry name" value="U2 snRNP-associated SURP motif-containing protein isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.330:FF:000177">
        <property type="entry name" value="U2 snRNP-associated SURP motif-containing protein-like isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.90">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.70.330">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.420">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.790">
        <property type="entry name" value="Surp module"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006569">
        <property type="entry name" value="CID_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008942">
        <property type="entry name" value="ENTH_VHS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013170">
        <property type="entry name" value="mRNA_splic_Cwf21_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012677">
        <property type="entry name" value="Nucleotide-bd_a/b_plait_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035979">
        <property type="entry name" value="RBD_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000504">
        <property type="entry name" value="RRM_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051485">
        <property type="entry name" value="SR-CTD_assoc_factor"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047488">
        <property type="entry name" value="SR140_cwf21"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035009">
        <property type="entry name" value="SR140_RRM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000061">
        <property type="entry name" value="Surp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035967">
        <property type="entry name" value="SWAP/Surp_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23140">
        <property type="entry name" value="RNA PROCESSING PROTEIN LD23810P"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23140:SF0">
        <property type="entry name" value="U2 SNRNP-ASSOCIATED SURP MOTIF-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04818">
        <property type="entry name" value="CID"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08312">
        <property type="entry name" value="cwf21"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00076">
        <property type="entry name" value="RRM_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01805">
        <property type="entry name" value="Surp"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01115">
        <property type="entry name" value="cwf21"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00582">
        <property type="entry name" value="RPR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00360">
        <property type="entry name" value="RRM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00648">
        <property type="entry name" value="SWAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54928">
        <property type="entry name" value="RNA-binding domain, RBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109905">
        <property type="entry name" value="Surp module (SWAP domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51391">
        <property type="entry name" value="CID"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50102">
        <property type="entry name" value="RRM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50128">
        <property type="entry name" value="SURP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1 2" id="KW-0694">RNA-binding</keyword>
    <feature type="domain" description="RRM" evidence="4">
        <location>
            <begin position="187"/>
            <end position="268"/>
        </location>
    </feature>
    <feature type="domain" description="SURP motif" evidence="5">
        <location>
            <begin position="343"/>
            <end position="386"/>
        </location>
    </feature>
    <feature type="domain" description="CID" evidence="6">
        <location>
            <begin position="447"/>
            <end position="584"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="52"/>
            <end position="186"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="688"/>
            <end position="747"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="761"/>
            <end position="935"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="9"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="56"/>
            <end position="67"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="79"/>
            <end position="91"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="99"/>
            <end position="135"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="152"/>
            <end position="162"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="3">
        <location>
            <begin position="691"/>
            <end position="711"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="715"/>
            <end position="732"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="780"/>
            <end position="828"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="856"/>
            <end position="886"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="3">
        <location>
            <begin position="897"/>
            <end position="935"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00176"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50102"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS51391"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015047.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="935" mass="108178" checksum="EBB60324812A9809" modified="2019-05-08" version="1">MSAAKRTLSKKEQEELKKKEDEKAAAEIYEEFLAAFEGSDGNKVKTFVRGGIVNASKDEHETDEKRGKIYKPSARFADQKNQPNQPSSNERPPSLLMIETKKPPLKKGEKEKKKSNLELFKEELKQIQEERDERHKTKGRLSRFEPPQSDSDGQRRSMDAPSRRNRSSGVLDDYAPGSHDVGDPSTTNLYLGNINPQMNEEMLCQEFGRFGPLASVKIMWPRTDEERARERNCGFVAFMNRRDAERALKNLNGKMIMSFEMKLGWGKAVPIPPHPIYIPPSMMEHTLPPPPSGLPFNAQPRERLKNPNAPMLPPPKNKEDFEKTLSQAIVKVVIPTERNLLALIHRMIEFVVREGPMFEAMIMNREINNPMFRFLFENQTPAHVYYRWKLYSILQGDAPTKWRTEDFRMFKNGSFWRPPPLNPYLHGMSEEQETEAFVEEPNKKGALKEEQRDKLEEILRGLTPRKNDIGDAMVFCLNNAEAAEEIVDCVTESLSILKTPLPKKIARLYLVSDVFFEFFIEINFETKLCQIFSDLNATYRTIQGHLQSENFKQRVMTCFRAWEDWAIYPEPFLIKLQNIFLGLVNIIEEKEIEEVPDDLDGAPIEEELDGAPLEDVDGIPIDAAPIDDLDGIPIKSLDDDLDGIPLDTSEESKKNEPIFKVAPSKWEAVDEAELEAQAVTTSKWELFDQHEESEEEENQNQEEESEDEEDTQSSKSEEQHMYSNPIKEEMPESKSSIKYSEMSEEKRAKLREIELKVMKFQDELESGKRPKKPGQSFQEQVEHYRDKLLQREKEKELERERERDKKDKEKSESRSKDKKEKEECTPTRKERKRRHSTSPSPSRSSGSRRAKSPSPKSERSERSERSHKETSRSRSSHKDSPRDVSKKAKRSPSGSRTPKRSRRSRSRSPKKSGKKSRSQSRSPHRSHKKSKKSKH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJG4</accession>
    <name>A0A452HJG4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">cAMP-dependent protein kinase</fullName>
            <ecNumber evidence="1">2.7.11.11</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="8">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.11"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="7">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.11"/>
        </reaction>
    </comment>
    <dbReference type="EC" id="2.7.11.11" evidence="1"/>
    <dbReference type="AlphaFoldDB" id="A0A452HJG4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017198.1">
        <property type="protein sequence ID" value="ENSGAGP00000015052.1"/>
        <property type="gene ID" value="ENSGAGG00000010963.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005952">
        <property type="term" value="C:cAMP-dependent protein kinase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004691">
        <property type="term" value="F:cAMP-dependent protein kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034237">
        <property type="term" value="F:protein kinase A regulatory subunit binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24353:SF153">
        <property type="entry name" value="CAMP-DEPENDENT PROTEIN KINASE CATALYTIC SUBUNIT 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24353">
        <property type="entry name" value="CYCLIC NUCLEOTIDE-DEPENDENT PROTEIN KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6 9" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="5" id="KW-0418">Kinase</keyword>
    <keyword evidence="4 9" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="3" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Protein kinase" evidence="11">
        <location>
            <begin position="46"/>
            <end position="313"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="248"/>
            <end position="313"/>
        </location>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="75"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00012444"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00047292"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00047454"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015052.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="313" mass="35209" checksum="9C50C85ABD773BD3" modified="2025-10-08" version="2">MLCRPSSEATNSPVSHGYKEFLAKGKEDFLKKWENPSQNTACLDQFERVKTLGMGSFGRVMLVKHKETGNYYAMKVLEKQKVRLVKLKQIEHTLNEKRILQAINFPLLIKLEYSFKDNSNLESHAWFYDLKPENLLLDQQGYIKVMDFGFAKRVKGRTWSLCGTPEYLAPEIILSKDYNKAVDWWALGVLIYEMAPIQIYEKIVSGKVFGLGEGTLVRFGGGFANQGSATCGTSSKHDRRADFEQHPATLSAPAHPSPLREQEAEAWFCGSQAPPPPRLPPTWRFPAPPPLLRQSVDGPSKGEGAERQPQAAP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJG8</accession>
    <name>A0A452HJG8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Bromo domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJG8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017200.1">
        <property type="protein sequence ID" value="ENSGAGP00000015054.1"/>
        <property type="gene ID" value="ENSGAGG00000010274.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05513">
        <property type="entry name" value="Bromo_brd7_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.920.10">
        <property type="entry name" value="Bromodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001487">
        <property type="entry name" value="Bromodomain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036427">
        <property type="entry name" value="Bromodomain-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051831">
        <property type="entry name" value="Bromodomain_contain_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021900">
        <property type="entry name" value="DUF3512"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22881">
        <property type="entry name" value="BROMODOMAIN CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22881:SF12">
        <property type="entry name" value="BROMODOMAIN-CONTAINING PROTEIN 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00439">
        <property type="entry name" value="Bromodomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12024">
        <property type="entry name" value="DUF3512"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00503">
        <property type="entry name" value="BROMODOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00297">
        <property type="entry name" value="BROMO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47370">
        <property type="entry name" value="Bromodomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50014">
        <property type="entry name" value="BROMODOMAIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3 6" id="KW-0103">Bromodomain</keyword>
    <keyword evidence="7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0539">Nucleus</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0804">Transcription</keyword>
    <keyword evidence="2" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="Bromo" evidence="9">
        <location>
            <begin position="144"/>
            <end position="214"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="29"/>
            <end position="126"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="247"/>
            <end position="272"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="287"/>
            <end position="314"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="31"/>
            <end position="41"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="43"/>
            <end position="53"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="8">
        <location>
            <begin position="54"/>
            <end position="65"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="66"/>
            <end position="102"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023117"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00035"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50014"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015054.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="651" mass="74143" checksum="DCF5181FDAF8501F" modified="2025-10-08" version="2">MYMEILKHKPDQKYVEKPLKLVLKVGGNEVTELSTGSSGHDSSLYEDKTEHEKHKDRKRKKRKKGEKQIPGEEKEKRKRKVKEDKKKRDREHAESEGERELKCQTPVRLELPPEKPLASSLAKQEEVEQTPLQEALNQLVRQLQRKDPNAFFSFPVTDFIAPGYSMIIKHPMDFSTMKEKIKNNGYQSIEELKDNFRLMCTNAMIYNKPETIYYKAAKKLLHSGMKILSQERIQSLKQSIDFMADLQQSRKQKDKMELQGNGEDDNLLGKDEDSMEADIKAFKTPNKEHKKKDKDLLEEKLRNNNLEREQEQIDRIIRESGGKLTRRLVNSQCEFERRKPDGTTTLGLLNPFDPNAGEPGYSPMKLGMTAGRLQSGVNTLQGFKEDKRNKVTPVVYLNYGPYSSYAPAYDSTFANISKDDSDLIYSTYGEDSNPECFSIHDFLAKSQDYPCVMADGLLDVLTKGGHSRSLRELETPMVDAEGQHGRSDTIKDVEIIETDAANKLDSNNDRLTALKAVTNFGIHVEEFDSEEAEIFQKKLDETTKLLRELQNAQNERLSTKPPPNMICLLGPSYKEMHLAERVTNNLKELAQQVTPGDIVSTYGIRKAMGISVPVPDTEASSVDLTDDFPKPLSSTTSLECNQVPLSHFHYP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJG9</accession>
    <name>A0A452HJG9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">SUI1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the SUI1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJG9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017206.1">
        <property type="protein sequence ID" value="ENSGAGP00000015060.1"/>
        <property type="gene ID" value="ENSGAGG00000011376.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003743">
        <property type="term" value="F:translation initiation factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010468">
        <property type="term" value="P:regulation of gene expression"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0080090">
        <property type="term" value="P:regulation of primary metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd11566">
        <property type="entry name" value="eIF1_SUI1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.780.10:FF:000003">
        <property type="entry name" value="Eukaryotic translation initiation factor 1b"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.780.10">
        <property type="entry name" value="SUI1-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001950">
        <property type="entry name" value="SUI1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036877">
        <property type="entry name" value="SUI1_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005874">
        <property type="entry name" value="SUI1_euk"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01160">
        <property type="entry name" value="SUI1_MOF2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10388">
        <property type="entry name" value="EUKARYOTIC TRANSLATION INITIATION FACTOR SUI1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01253">
        <property type="entry name" value="SUI1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF004499">
        <property type="entry name" value="SUI1_euk"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55159">
        <property type="entry name" value="eIF1-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50296">
        <property type="entry name" value="SUI1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0007">Acetylation</keyword>
    <keyword evidence="2" id="KW-0396">Initiation factor</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="4" id="KW-0648">Protein biosynthesis</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="SUI1" evidence="6">
        <location>
            <begin position="31"/>
            <end position="101"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00005422"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022540"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022917"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50296"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015060.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="103" mass="11735" checksum="0291A44634BB58B4" modified="2019-05-08" version="1">MSTIQNLQSFDPFADATKGDDLLPAGTEDYIHIRIQQRNGRKTLTTVQGIADDYDKKKLVKAFKKKFACNGTVIEHPEYGEVIQLQGDQRKNICQFLLELLIH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJH1</accession>
    <name>A0A452HJH1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Bromo domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJH1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017210.1">
        <property type="protein sequence ID" value="ENSGAGP00000015064.1"/>
        <property type="gene ID" value="ENSGAGG00000010274.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05513">
        <property type="entry name" value="Bromo_brd7_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.920.10:FF:000022">
        <property type="entry name" value="Putative bromodomain-containing protein 9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.920.10">
        <property type="entry name" value="Bromodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001487">
        <property type="entry name" value="Bromodomain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036427">
        <property type="entry name" value="Bromodomain-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051831">
        <property type="entry name" value="Bromodomain_contain_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021900">
        <property type="entry name" value="DUF3512"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22881">
        <property type="entry name" value="BROMODOMAIN CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22881:SF12">
        <property type="entry name" value="BROMODOMAIN-CONTAINING PROTEIN 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00439">
        <property type="entry name" value="Bromodomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12024">
        <property type="entry name" value="DUF3512"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00503">
        <property type="entry name" value="BROMODOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00297">
        <property type="entry name" value="BROMO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47370">
        <property type="entry name" value="Bromodomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50014">
        <property type="entry name" value="BROMODOMAIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3 6" id="KW-0103">Bromodomain</keyword>
    <keyword evidence="7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0539">Nucleus</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0804">Transcription</keyword>
    <keyword evidence="2" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="Bromo" evidence="9">
        <location>
            <begin position="140"/>
            <end position="210"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="25"/>
            <end position="119"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="243"/>
            <end position="268"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="283"/>
            <end position="310"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="27"/>
            <end position="37"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="39"/>
            <end position="49"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="8">
        <location>
            <begin position="50"/>
            <end position="61"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="62"/>
            <end position="98"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023117"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00035"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50014"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015064.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="641" mass="72672" checksum="07E6F2B723223E3C" modified="2019-05-08" version="1">QTLKNKKHKYVEKPLKLVLKVGGNEVTELSTGSSGHDSSLYEDKTEHEKHKDRKRKKRKKGEKQIPGEEKEKRKRKVKEDKKKRDREHAESEGERELKCQTPVRLELPPEKPLASSLAKQEEVEQTPLQEALNQLVRQLQRKDPNAFFSFPVTDFIAPGYSMIIKHPMDFSTMKEKIKNNGYQSIEELKDNFRLMCTNAMIYNKPETIYYKAAKKLLHSGMKILSQERIQSLKQSIDFMADLQQSRKQKDKMELQGNGEDDNLLGKDEDSMEADIKAFKTPNKEHKKKDKDLLEEKLRNNNLEREQEQIDRIIRESGGKLTRRLVNSQCEFERRKPDGTTTLGLLNPFDPNAGEPGYSPMKLGMTAGRLQSGVNTLQGFKEDKRNKVTPVVYLNYGPYSSYAPAYDSTFANISKDDSDLIYSTYGEDSNPECFSIHDFLAKSQDYPCVMADGLLDVLTKGGHSRSLRELETPMVDAEGQHGRSDTIKDIIETDAANKLDSNNDRLTALKAVTNFGIHVEEFDSEEAEIFQKKLDETTKLLRELQNAQNERLSTKPPPNMICLLGPSYKEMHLAERVTNNLKELAQQVTPGDIVSTYGIRKAMGISVPVPDTEASSVDLTDDCQESKKAASFSGNESGPIAI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJH3</accession>
    <name>A0A452HJH3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Retinol dehydrogenase 16</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1 2">Belongs to the short-chain dehydrogenases/reductases (SDR) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJH3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017207.1">
        <property type="protein sequence ID" value="ENSGAGP00000015061.1"/>
        <property type="gene ID" value="ENSGAGG00000011337.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016491">
        <property type="term" value="F:oxidoreductase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008202">
        <property type="term" value="P:steroid metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.720">
        <property type="entry name" value="NAD(P)-binding Rossmann-like Domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036291">
        <property type="entry name" value="NAD(P)-bd_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002347">
        <property type="entry name" value="SDR_fam"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43313:SF4">
        <property type="entry name" value="17-BETA-HYDROXYSTEROID DEHYDROGENASE TYPE 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43313">
        <property type="entry name" value="SHORT-CHAIN DEHYDROGENASE/REDUCTASE FAMILY 9C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00106">
        <property type="entry name" value="adh_short"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00081">
        <property type="entry name" value="GDHRDH"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00080">
        <property type="entry name" value="SDRFAMILY"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51735">
        <property type="entry name" value="NAD(P)-binding Rossmann-fold domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006484"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="RuleBase" id="RU000363"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015061.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="5">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="202" mass="22773" checksum="F5CEF49F993F4CA0" modified="2019-05-08" version="1">HHLKELPGLWGLVNNAGVALPLVPNEWLTKDDFVRVLDVNLIGLIEVTLSFLPLVKRARGRVVNMASILGRLAFYGGGYCPSKYGVEAFSDSLRRELRPFGMKIAIVEPGYFRTPISDMQWERLPSAGRMHALTREILTTKPSTNLYLVTDCMEHALTSCHPRTRYSGGWDAQFFYIPLSYLPTALADWVLTWGWPRPAQAV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJH8</accession>
    <name>A0A452HJH8_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="7">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the actin family. ARP1 subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJH8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017213.1">
        <property type="protein sequence ID" value="ENSGAGP00000015066.1"/>
        <property type="gene ID" value="ENSGAGG00000011333.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005815">
        <property type="term" value="C:microtubule organizing center"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd10216">
        <property type="entry name" value="ASKHA_NBD_Arp1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.420.40:FF:000188">
        <property type="entry name" value="Actin like 6B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.420.40:FF:000502">
        <property type="entry name" value="Actin-Related Proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.640.10:FF:000008">
        <property type="entry name" value="alpha-centractin isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.420.40">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.640.10">
        <property type="entry name" value="Actin, Chain A, domain 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004000">
        <property type="entry name" value="Actin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020902">
        <property type="entry name" value="Actin/actin-like_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004001">
        <property type="entry name" value="Actin_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043129">
        <property type="entry name" value="ATPase_NBD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11937">
        <property type="entry name" value="ACTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00022">
        <property type="entry name" value="Actin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00190">
        <property type="entry name" value="ACTIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00268">
        <property type="entry name" value="ACTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53067">
        <property type="entry name" value="Actin-like ATPase domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00432">
        <property type="entry name" value="ACTINS_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01132">
        <property type="entry name" value="ACTINS_ACT_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="5" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="3" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00038483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015066.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="329" mass="37154" checksum="0BC7F76B100F4DAA" modified="2019-05-08" version="1">MAGALEGDLFIGPKAEEHRGLLSIRYPMEHGIVRDWNDMERIWQYVYSKDQLQTFSEEHPVLLTEAPLNPCKNREKAAEVFFETFNVPALFISMQAVLSLYATGRTTGVVLDAGDGVTHAVPIYEGFAMPHSIMRVDVAGRDVSRYLRLLLRKEGYDFHTSAEFEVVKMIKERACYLSINPQKDEALETEKVLYNLPDGSTLEVGPARFRAPELLFQPDLVGDESEGIHEVLAFAIQKSDMDLRRTLFANIVLSGGSTLFKGFGDRLLSEVKKLAPKDVKIKISAPQERLYSTWIGGSILASLDTFKKMWVSKKEYEEDGARAIHRKTF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJI0</accession>
    <name>A0A452HJI0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">Auxilin</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="11">DnaJ homolog subfamily C member 6</fullName>
        </alternativeName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="9">Forms a complex composed of HSPA8, CLTC and DNAJC6. Interacts with HSPA8/HSC70 in an ATP-dependent manner; this interaction stimulates the HSPA8's ATPase activity. Interacts with CLTC; this interaction produces a local change in heavy-chain contacts, creating a detectable global distortion of the clathrin coat. Interacts with AP2A2. Interacts with DNM1(GTP-bound form); this interaction allows clathrin-coated vesicle (CCV) formation at the plasma membrane.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasmic vesicle</location>
            <location evidence="1">Clathrin-coated vesicle</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJI0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017218.1">
        <property type="protein sequence ID" value="ENSGAGP00000015070.1"/>
        <property type="gene ID" value="ENSGAGG00000011384.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030136">
        <property type="term" value="C:clathrin-coated vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0014069">
        <property type="term" value="C:postsynaptic density"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098793">
        <property type="term" value="C:presynapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="GOC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030276">
        <property type="term" value="F:clathrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004721">
        <property type="term" value="F:phosphoprotein phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017124">
        <property type="term" value="F:SH3 domain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072583">
        <property type="term" value="P:clathrin-dependent endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016191">
        <property type="term" value="P:synaptic vesicle uncoating"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd06257">
        <property type="entry name" value="DnaJ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd14563">
        <property type="entry name" value="PTP_auxilin_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1110:FF:000001">
        <property type="entry name" value="cyclin-G-associated kinase isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.190.10:FF:000255">
        <property type="entry name" value="putative tyrosine-protein phosphatase auxilin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.110:FF:000002">
        <property type="entry name" value="putative tyrosine-protein phosphatase auxilin isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1110">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.110">
        <property type="entry name" value="DnaJ domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.190.10">
        <property type="entry name" value="Protein tyrosine phosphatase superfamily"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001623">
        <property type="entry name" value="DnaJ_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036869">
        <property type="entry name" value="J_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029021">
        <property type="entry name" value="Prot-tyrosine_phosphatase-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014020">
        <property type="entry name" value="Tensin_C2-dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029023">
        <property type="entry name" value="Tensin_phosphatase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23172">
        <property type="entry name" value="AUXILIN/CYCLIN G-ASSOCIATED KINASE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23172:SF4">
        <property type="entry name" value="TYROSINE-PROTEIN PHOSPHATASE AUXILIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10409">
        <property type="entry name" value="PTEN_C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00271">
        <property type="entry name" value="DnaJ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01326">
        <property type="entry name" value="PTEN_C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52799">
        <property type="entry name" value="(Phosphotyrosine protein) phosphatases II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49562">
        <property type="entry name" value="C2 domain (Calcium/lipid-binding domain, CaLB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46565">
        <property type="entry name" value="Chaperone J-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51182">
        <property type="entry name" value="C2_TENSIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50076">
        <property type="entry name" value="DNAJ_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51181">
        <property type="entry name" value="PPASE_TENSIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0143">Chaperone</keyword>
    <keyword evidence="8" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="4" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="5" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0729">SH3-binding</keyword>
    <feature type="domain" description="Phosphatase tensin-type" evidence="14">
        <location>
            <begin position="55"/>
            <end position="222"/>
        </location>
    </feature>
    <feature type="domain" description="C2 tensin-type" evidence="15">
        <location>
            <begin position="228"/>
            <end position="366"/>
        </location>
    </feature>
    <feature type="domain" description="J" evidence="13">
        <location>
            <begin position="866"/>
            <end position="930"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="431"/>
            <end position="522"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="537"/>
            <end position="794"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="12">
        <location>
            <begin position="434"/>
            <end position="456"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="464"/>
            <end position="475"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="510"/>
            <end position="522"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="664"/>
            <end position="684"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="692"/>
            <end position="707"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="745"/>
            <end position="755"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="756"/>
            <end position="772"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004132"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023036"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023186"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00064305"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00069335"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00075670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS51181"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS51182"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015070.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="930" mass="101435" checksum="5F7B8498A7ACF7F0" modified="2019-05-08" version="1">MKESENKGASSPDMESTYGGGLLDMVKGGAGRLFSNLKDNLKDTLKDTSSKVMQSVASYTKGELDISYVTSRIIVMSFPAEGVELGFRNHIEDVRTFLDSRHPDHYTVFNLSQKSYRSAKFHNRVSECSWPMRQAPSLHNLYAVCKNMHNWLQQNPKNVCVIHCMDGRAASAVLVSAMFCFCHLFSSPVPAIQLLNSKRPGIGLWPSHRRYLGYICDLIAEKPIDPHCKPLAIKSVCLSPVPCFNKQRNGCRPFCDILIGETRIFTTSQEYERMKEFRVQEGKVLIPLGVTVHGDVVLSVYHMRSTIGGRLQAKMTNTQIFQIQFHTGFIALGTTMLKFTKPELDACDSPEKYPQLFHVTLEIEIESADRQTDLTPPWENFTTKDINASILFSSHQEHQDALALAGRGPVDAHQDNIKNVGQGTFLSSLSWQDQKSDKSSSHPTSEDRAALVHEESEQSDDELLSLSSQHSNASSDKTHGTPKHSKKQQEPPAPLPPEDVDLLGLDGSPMSKNFPTQSTALPSNSDLLSDLFGVSDSRAHSQGVQAAAEDVFHVGGPGSAHSTPRRSAASASPSPAPRVGEATAFDPFGTRPKQSGPDLLGSFLGSSNATGDPFLQATRSPSPTVHSDPFHMASSTPTVAVQPDVSGGWEWSNKSGGLGMGSKSAATSPTGSLHSTPTHQSKPQTLDPFADLGALGANLAGGSSFASKPTTPTGISGGFPPMGSPQKPSPQPMGGSGWQQGAGYSWQQSQSKSQQNIPHASPQNRPNYNVSFSAMPGGQNDRGKGPANADSKPKVSADFEDLLSGQGFNAHKDKKGPRTIAEMRKEEMAKDMDPEKLKILEWIEGKERNIRALLSTMHTVLWAGETKWKPIGMADLVTPEQVKKVYRRAVLVVHPDKATGQPYEQYAKMIFMELNDAWSEFENQGQKPLY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJI2</accession>
    <name>A0A452HJI2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Retinol dehydrogenase 16</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1 3">Belongs to the short-chain dehydrogenases/reductases (SDR) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJI2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017222.1">
        <property type="protein sequence ID" value="ENSGAGP00000015074.1"/>
        <property type="gene ID" value="ENSGAGG00000011337.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016491">
        <property type="term" value="F:oxidoreductase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008202">
        <property type="term" value="P:steroid metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.720:FF:000074">
        <property type="entry name" value="Retinol dehydrogenase type 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.720">
        <property type="entry name" value="NAD(P)-binding Rossmann-like Domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036291">
        <property type="entry name" value="NAD(P)-bd_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002347">
        <property type="entry name" value="SDR_fam"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43313:SF4">
        <property type="entry name" value="17-BETA-HYDROXYSTEROID DEHYDROGENASE TYPE 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43313">
        <property type="entry name" value="SHORT-CHAIN DEHYDROGENASE/REDUCTASE FAMILY 9C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00106">
        <property type="entry name" value="adh_short"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00081">
        <property type="entry name" value="GDHRDH"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00080">
        <property type="entry name" value="SDRFAMILY"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51735">
        <property type="entry name" value="NAD(P)-binding Rossmann-fold domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="4">
        <location>
            <begin position="1"/>
            <end position="17"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019525649" description="Retinol dehydrogenase 16" evidence="4">
        <location>
            <begin position="18"/>
            <end position="258"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006484"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="RuleBase" id="RU000363"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015074.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="7">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="258" mass="29000" checksum="717966BC171E1C01" modified="2019-05-08" version="1">MWLYLAALLGLYFLCRWYRERQIVENLREKYVLITGCDSGFGNLLAKQLDVRGLRVLAACLTQQGAEQLRKATSERLQTVILDVTRTDSIAAATAWVKERVGDKGLWGLVNNAGVALPLVPNEWLTKDDFVRVLDVNLIGLIEVTLSFLPLVKRARGRVVNMASILGRLAFYGGGYCPSKYGVEAFSDSLRRELRPFGMKIAIVEPGYFRTPISDMQRHLECLEQSWRNARPEVKESYGQHYFDSCDSSPQGVGFSVG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJI5</accession>
    <name>A0A452HJI5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Bromo domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJI5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017223.1">
        <property type="protein sequence ID" value="ENSGAGP00000015075.1"/>
        <property type="gene ID" value="ENSGAGG00000010274.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05513">
        <property type="entry name" value="Bromo_brd7_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.920.10">
        <property type="entry name" value="Bromodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001487">
        <property type="entry name" value="Bromodomain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036427">
        <property type="entry name" value="Bromodomain-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051831">
        <property type="entry name" value="Bromodomain_contain_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021900">
        <property type="entry name" value="DUF3512"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22881">
        <property type="entry name" value="BROMODOMAIN CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22881:SF12">
        <property type="entry name" value="BROMODOMAIN-CONTAINING PROTEIN 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00439">
        <property type="entry name" value="Bromodomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12024">
        <property type="entry name" value="DUF3512"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00503">
        <property type="entry name" value="BROMODOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00297">
        <property type="entry name" value="BROMO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47370">
        <property type="entry name" value="Bromodomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50014">
        <property type="entry name" value="BROMODOMAIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3 6" id="KW-0103">Bromodomain</keyword>
    <keyword evidence="7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0539">Nucleus</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0804">Transcription</keyword>
    <keyword evidence="2" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="Bromo" evidence="9">
        <location>
            <begin position="144"/>
            <end position="214"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="29"/>
            <end position="126"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="247"/>
            <end position="272"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="287"/>
            <end position="314"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="31"/>
            <end position="41"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="43"/>
            <end position="53"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="8">
        <location>
            <begin position="54"/>
            <end position="65"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="66"/>
            <end position="102"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023117"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00035"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50014"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015075.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="671" mass="76364" checksum="D3D06C8D1A7C33BD" modified="2025-10-08" version="2">MYMEILKHKPDQKYVEKPLKLVLKVGGNEVTELSTGSSGHDSSLYEDKTEHEKHKDRKRKKRKKGEKQIPGEEKEKRKRKVKEDKKKRDREHAESEGERELKCQTPVRLELPPEKPLASSLAKQEEVEQTPLQEALNQLVRQLQRKDPNAFFSFPVTDFIAPGYSMIIKHPMDFSTMKEKIKNNGYQSIEELKDNFRLMCTNAMIYNKPETIYYKAAKKLLHSGMKILSQERIQSLKQSIDFMADLQQSRKQKDKMELQGNGEDDNLLGKDEDSMEADIKAFKTPNKEHKKKDKDLLEEKLRNNNLEREQEQIDRIIRESGGKLTRRLVNSQCEFERRKPDGTTTLGLLNPFDPNAGEPGYSPMKLGMTAGRLQSGVNTLQGFKEDKRNKVTPVVYLNYGPYSSYAPAYDSTFANISKDDSDLIYSTYGEDSNPECFSIHDFLAKSQDYPCVMADGLLDVLTKGGHSRSLRELETPMVDAEGQHGRSDTIKDVEIIETDAANKLDSNNDRLTALKAVTNFGIHVEEFDSEEAEIFQKKLDETTKLLRELQNAQNERLSTKPPPNMICLLGPSYKEMHLAERVTNNLKELAQQVTPGDIVSTYGIRKAMGISVPVPDTEASSVDLTDGEYRVYLDMCCSRAKLHLQVIVVLLVATAEVKVESQLLVSLSFLK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJI7</accession>
    <name>A0A452HJI7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Bromo domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJI7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017227.1">
        <property type="protein sequence ID" value="ENSGAGP00000015079.1"/>
        <property type="gene ID" value="ENSGAGG00000010274.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05513">
        <property type="entry name" value="Bromo_brd7_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.920.10:FF:000022">
        <property type="entry name" value="Putative bromodomain-containing protein 9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.920.10">
        <property type="entry name" value="Bromodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001487">
        <property type="entry name" value="Bromodomain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036427">
        <property type="entry name" value="Bromodomain-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051831">
        <property type="entry name" value="Bromodomain_contain_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021900">
        <property type="entry name" value="DUF3512"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22881">
        <property type="entry name" value="BROMODOMAIN CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22881:SF12">
        <property type="entry name" value="BROMODOMAIN-CONTAINING PROTEIN 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00439">
        <property type="entry name" value="Bromodomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12024">
        <property type="entry name" value="DUF3512"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00503">
        <property type="entry name" value="BROMODOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00297">
        <property type="entry name" value="BROMO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47370">
        <property type="entry name" value="Bromodomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50014">
        <property type="entry name" value="BROMODOMAIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3 6" id="KW-0103">Bromodomain</keyword>
    <keyword evidence="7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0539">Nucleus</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0804">Transcription</keyword>
    <keyword evidence="2" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="Bromo" evidence="9">
        <location>
            <begin position="141"/>
            <end position="211"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="27"/>
            <end position="120"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="244"/>
            <end position="269"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="284"/>
            <end position="311"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="28"/>
            <end position="38"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="40"/>
            <end position="50"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="8">
        <location>
            <begin position="51"/>
            <end position="62"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="63"/>
            <end position="99"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023117"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00035"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50014"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015079.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="668" mass="75863" checksum="7639E25A3B021A75" modified="2019-05-08" version="1">DNDSTLKHCKYVEKPLKLVLKVGGNEVTELSTGSSGHDSSLYEDKTEHEKHKDRKRKKRKKGEKQIPGEEKEKRKRKVKEDKKKRDREHAESEGERELKCQTPVRLELPPEKPLASSLAKQEEVEQTPLQEALNQLVRQLQRKDPNAFFSFPVTDFIAPGYSMIIKHPMDFSTMKEKIKNNGYQSIEELKDNFRLMCTNAMIYNKPETIYYKAAKKLLHSGMKILSQERIQSLKQSIDFMADLQQSRKQKDKMELQGNGEDDNLLGKDEDSMEADIKAFKTPNKEHKKKDKDLLEEKLRNNNLEREQEQIDRIIRESGGKLTRRLVNSQCEFERRKPDGTTTLGLLNPFDPNAGEPGYSPMKLGMTAGRLQSGVNTLQGFKEDKRNKVTPVVYLNYGPYSSYAPAYDSTFANISKDDSDLIYSTYGEDSNPECFSIHDFLAKSQDYPCVMADGLLDVLTKGGHSRSLRELETPMVDAEGQHGRSDTIKDVEIIETDAANKLDSNNDRLTALKAVTNFGIHVEEFDSEEAEIFQKKLDETTKLLRELQNAQNERLSTKPPPNMICLLGPSYKEMHLAERVTNNLKELAQQVTPGDIVSTYGIRKAMGISVPVPDTEASSVDLTDGEYRVYLDMCCSRAKLHLQVIVVLLVATAEVKVESQLLVSLSFLK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJK0</accession>
    <name>A0A452HJK0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Polyadenylate-binding protein-interacting protein 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="6">Acts as a repressor in the regulation of translation initiation of poly(A)-containing mRNAs. Its inhibitory activity on translation is mediated via its action on PABPC1. Displaces the interaction of PABPC1 with poly(A) RNA and competes with PAIP1 for binding to PABPC1. Its association with PABPC1 results in disruption of the cytoplasmic poly(A) RNP structure organization.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the PAIP2 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJK0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017237.1">
        <property type="protein sequence ID" value="ENSGAGP00000015088.1"/>
        <property type="gene ID" value="ENSGAGG00000011392.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000900">
        <property type="term" value="F:mRNA regulatory element binding translation repressor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045947">
        <property type="term" value="P:negative regulation of translational initiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040396">
        <property type="entry name" value="PAIP2-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009818">
        <property type="entry name" value="PAM2_motif"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13154">
        <property type="entry name" value="POLYADENYLATE-BINDING PROTEIN-INTERACTING PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13154:SF2">
        <property type="entry name" value="POLYADENYLATE-BINDING PROTEIN-INTERACTING PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07145">
        <property type="entry name" value="PAM2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0810">Translation regulation</keyword>
    <keyword evidence="4" id="KW-0832">Ubl conjugation</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006858"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022845"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00037448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00040702"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015088.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="135" mass="16031" checksum="24064BC381ACBF20" modified="2019-05-08" version="1">MLWRLQVKGSTMKDPSRSSTSPSIINEDVIINGHSHEDDNPFAEYMWMENEEEFNRQIEEELWEEEFIERCFQEMLEEEEEHEWFIPARDLPQTMDQIQDQFNDLVISDSSSLEDLVVKSNLNPNAKEFVPGVKY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJK7</accession>
    <name>A0A452HJK7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Large ribosomal subunit protein uL14m</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="8">39S ribosomal protein L14, mitochondrial</fullName>
        </alternativeName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Mitochondrion</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the universal ribosomal protein uL14 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJK7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017246.1">
        <property type="protein sequence ID" value="ENSGAGP00000015096.1"/>
        <property type="gene ID" value="ENSGAGG00000011397.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005743">
        <property type="term" value="C:mitochondrial inner membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:1990904">
        <property type="term" value="C:ribonucleoprotein complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005840">
        <property type="term" value="C:ribosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003735">
        <property type="term" value="F:structural constituent of ribosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006412">
        <property type="term" value="P:translation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00337">
        <property type="entry name" value="Ribosomal_uL14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.150.20:FF:000004">
        <property type="entry name" value="39S ribosomal protein L14, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.150.20">
        <property type="entry name" value="Ribosomal protein L14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="HAMAP" id="MF_01367">
        <property type="entry name" value="Ribosomal_uL14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000218">
        <property type="entry name" value="Ribosomal_uL14"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036853">
        <property type="entry name" value="Ribosomal_uL14_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21037">
        <property type="entry name" value="39S RIBOSOMAL PROTEIN L14, MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21037:SF3">
        <property type="entry name" value="LARGE RIBOSOMAL SUBUNIT PROTEIN UL14M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00238">
        <property type="entry name" value="Ribosomal_L14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01374">
        <property type="entry name" value="Ribosomal_L14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50193">
        <property type="entry name" value="Ribosomal protein L14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0687">Ribonucleoprotein</keyword>
    <keyword evidence="4" id="KW-0689">Ribosomal protein</keyword>
    <keyword evidence="3" id="KW-0809">Transit peptide</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004173"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010745"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022980"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023274"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00040118"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00042938"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015096.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="169" mass="18490" checksum="F78747EE695F1FA1" modified="2019-05-08" version="1">MRAGDRAGGAGSLRPVRLVIRISMAFLSRQLGLLLTQVNRAVEQRHFSITGACSAIQKLTRVRVVDNSALGNTPYHRPPKCIHVYNKTGVGKVGDKILLAIKGEKKKALIVGHRMPGPCMTPRFDSNNVVLIEDNGNPTGTRIKTPIPLILRKREGEFSKVLAIAQHLV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJK9</accession>
    <name>A0A452HJK9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Bromo domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJK9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017225.1">
        <property type="protein sequence ID" value="ENSGAGP00000015077.1"/>
        <property type="gene ID" value="ENSGAGG00000010274.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05513">
        <property type="entry name" value="Bromo_brd7_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.920.10">
        <property type="entry name" value="Bromodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001487">
        <property type="entry name" value="Bromodomain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036427">
        <property type="entry name" value="Bromodomain-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051831">
        <property type="entry name" value="Bromodomain_contain_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021900">
        <property type="entry name" value="DUF3512"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22881">
        <property type="entry name" value="BROMODOMAIN CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22881:SF12">
        <property type="entry name" value="BROMODOMAIN-CONTAINING PROTEIN 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00439">
        <property type="entry name" value="Bromodomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12024">
        <property type="entry name" value="DUF3512"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00503">
        <property type="entry name" value="BROMODOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00297">
        <property type="entry name" value="BROMO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47370">
        <property type="entry name" value="Bromodomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50014">
        <property type="entry name" value="BROMODOMAIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3 6" id="KW-0103">Bromodomain</keyword>
    <keyword evidence="7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0539">Nucleus</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0804">Transcription</keyword>
    <keyword evidence="2" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="Bromo" evidence="9">
        <location>
            <begin position="144"/>
            <end position="214"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="29"/>
            <end position="126"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="247"/>
            <end position="272"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="287"/>
            <end position="314"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="31"/>
            <end position="41"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="43"/>
            <end position="53"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="8">
        <location>
            <begin position="54"/>
            <end position="65"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="66"/>
            <end position="102"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023117"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00035"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50014"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015077.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="666" mass="75932" checksum="08FC4FC34BB72BFA" modified="2025-10-08" version="2">MYMEILKHKPDQKYVEKPLKLVLKVGGNEVTELSTGSSGHDSSLYEDKTEHEKHKDRKRKKRKKGEKQIPGEEKEKRKRKVKEDKKKRDREHAESEGERELKCQTPVRLELPPEKPLASSLAKQEEVEQTPLQEALNQLVRQLQRKDPNAFFSFPVTDFIAPGYSMIIKHPMDFSTMKEKIKNNGYQSIEELKDNFRLMCTNAMIYNKPETIYYKAAKKLLHSGMKILSQERIQSLKQSIDFMADLQQSRKQKDKMELQGNGEDDNLLGKDEDSMEADIKAFKTPNKEHKKKDKDLLEEKLRNNNLEREQEQIDRIIRESGGKLTRRLVNSQCEFERRKPDGTTTLGLLNPFDPNAGEPGYSPMKLGMTAGRLQSGVNTLQGFKEDKRNKVTPVVYLNYGPYSSYAPAYDSTFANISKDDSDLIYSTYGEDSNPECFSIHDFLAKSQDYPCVMADGLLDVLTKGGHSRSLRELETSFSYLDKGEGHYYSIIETDAANKLDSNNDRLTALKAVTNFGIHVEEFDSEEAEIFQKKLDETTKLLRELQNAQNERLSTKPPPNMICLLGPSYKEMHLAERVTNNLKELAQQVTPGDIVSTYGIRKAMGISVPVPDTEASSVDLTDGEYRVYLDMCCSRAKLHLQVIVVLLVATAEVKVESQLLVSLSFLK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJL0</accession>
    <name>A0A452HJL0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Cytochrome P450 family 4 subfamily B member 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="9">
            <name>heme</name>
            <dbReference type="ChEBI" id="CHEBI:30413"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the cytochrome P450 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJL0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017249.1">
        <property type="protein sequence ID" value="ENSGAGP00000015099.1"/>
        <property type="gene ID" value="ENSGAGG00000011253.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0020037">
        <property type="term" value="F:heme binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005506">
        <property type="term" value="F:iron ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004497">
        <property type="term" value="F:monooxygenase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016705">
        <property type="term" value="F:oxidoreductase activity, acting on paired donors, with incorporation or reduction of molecular oxygen"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd20678">
        <property type="entry name" value="CYP4B-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.630.10:FF:000005">
        <property type="entry name" value="cytochrome P450 4F22 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.630.10">
        <property type="entry name" value="Cytochrome P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001128">
        <property type="entry name" value="Cyt_P450"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002401">
        <property type="entry name" value="Cyt_P450_E_grp-I"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036396">
        <property type="entry name" value="Cyt_P450_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050196">
        <property type="entry name" value="Cytochrome_P450_Monoox"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24291">
        <property type="entry name" value="CYTOCHROME P450 FAMILY 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24291:SF201">
        <property type="entry name" value="CYTOCHROME P450, FAMILY 4, SUBFAMILY B, POLYPEPTIDE 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00067">
        <property type="entry name" value="p450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00463">
        <property type="entry name" value="EP450I"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00385">
        <property type="entry name" value="P450"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48264">
        <property type="entry name" value="Cytochrome P450"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="3 9" id="KW-0349">Heme</keyword>
    <keyword evidence="6 9" id="KW-0408">Iron</keyword>
    <keyword evidence="8 10" id="KW-0472">Membrane</keyword>
    <keyword evidence="4 9" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7" id="KW-0503">Monooxygenase</keyword>
    <keyword evidence="7" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="10" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="20"/>
            <end position="40"/>
        </location>
    </feature>
    <feature type="binding site" description="axial binding residue" evidence="9">
        <location>
            <position position="439"/>
        </location>
        <ligand>
            <name>heme</name>
            <dbReference type="ChEBI" id="CHEBI:30413"/>
        </ligand>
        <ligandPart>
            <name>Fe</name>
            <dbReference type="ChEBI" id="CHEBI:18248"/>
        </ligandPart>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004586"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023033"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PIRSR" id="PIRSR602401-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015099.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="13">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="492" mass="56459" checksum="D7E4D621D027ACFA" modified="2019-05-08" version="1">TSELSPVPSTRLFATPSSLLLYPTAGFCLIYVLLKAIQLYRKRQQLLKAFESFPGPPSHWLYGHSSQISQGGELNQLLSWAKEYPYAYPRWIGGFMASLTVNHPDYAKTVFGHGGKGLLILHGPKWFQHRRLLTPGFHYDVLKPYVSLMAESTKVMLDKWEWLITVDKSVELFEHVSLMTLDSIMKCAFSCHSNCQTDRDSDYYIKAVYDLTYLVFHRIRFFPFHNDFIYRISSQGRRFQEACRLAHQHTDKVIKERKTSLQDQQELEKIQKKKYLDFLDILLCAKDENGAGLSDEDLRAEVDTFMFEGHDTTASGVSWLLYCMALHPEHQQRCREEIKEILGDRETIQWDDLGKMTYSTMCIKESLRLYPPVPGVSRLLSKPITFHDGRTVPEGLFSLFLLKGPILPSVFDPMRFSPENVSKRHSHAFLPFAAGARNCIGKQFAMNELKVALALTLLRFELAPDPAKPPKEIPQIVLKSSNGIHLLLRKLP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJL2</accession>
    <name>A0A452HJL2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Retinol dehydrogenase 16</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1 3">Belongs to the short-chain dehydrogenases/reductases (SDR) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJL2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017251.1">
        <property type="protein sequence ID" value="ENSGAGP00000015101.1"/>
        <property type="gene ID" value="ENSGAGG00000011337.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016491">
        <property type="term" value="F:oxidoreductase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008202">
        <property type="term" value="P:steroid metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.720:FF:000074">
        <property type="entry name" value="Retinol dehydrogenase type 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.720">
        <property type="entry name" value="NAD(P)-binding Rossmann-like Domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036291">
        <property type="entry name" value="NAD(P)-bd_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002347">
        <property type="entry name" value="SDR_fam"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43313:SF4">
        <property type="entry name" value="17-BETA-HYDROXYSTEROID DEHYDROGENASE TYPE 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43313">
        <property type="entry name" value="SHORT-CHAIN DEHYDROGENASE/REDUCTASE FAMILY 9C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00106">
        <property type="entry name" value="adh_short"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00081">
        <property type="entry name" value="GDHRDH"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00080">
        <property type="entry name" value="SDRFAMILY"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51735">
        <property type="entry name" value="NAD(P)-binding Rossmann-fold domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="4">
        <location>
            <begin position="1"/>
            <end position="17"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019105359" description="Retinol dehydrogenase 16" evidence="4">
        <location>
            <begin position="18"/>
            <end position="299"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006484"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="RuleBase" id="RU000363"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015101.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="7">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="299" mass="33734" checksum="BD83EBC781D599D8" modified="2019-05-08" version="1">MWLYLAALLGLYFLCRWYRERQIVENLREKYVLITGCDSGFGNLLAKQLDVRGLRVLAACLTQQGAEQLRKATSERLQTVILDVTRTDSIAAATAWVKERVGDKGLWGLVNNAGVALPLVPNEWLTKDDFVRVLDVNLIGLIEVTLSFLPLVKRARGRVVNMASILGRLAFYGGGYCPSKYGVEAFSDSLRRELRPFGMKIAIVEPGYFRTPISDMQWERLPSAGRMHALTREILTTKPSTNLYLVTDCMEHALTSCHPRTRYSGGWDAQFFYIPLSYLPTALADWVLTWGWPRPAQAV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJL4</accession>
    <name>A0A452HJL4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Propionyl-CoA carboxylase alpha chain, mitochondrial</fullName>
            <ecNumber evidence="4">6.4.1.3</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="16">Propanoyl-CoA:carbon dioxide ligase subunit alpha</fullName>
        </alternativeName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="17">
            <text>butanoyl-CoA + hydrogencarbonate + ATP = (2S)-ethylmalonyl-CoA + ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:59520"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:17544"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:57371"/>
            <dbReference type="ChEBI" id="CHEBI:60909"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="17">
            <dbReference type="Rhea" id="RHEA:59521"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>propanoyl-CoA + hydrogencarbonate + ATP = (S)-methylmalonyl-CoA + ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:23720"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:17544"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:57327"/>
            <dbReference type="ChEBI" id="CHEBI:57392"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="6.4.1.3"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="18">
            <dbReference type="Rhea" id="RHEA:23721"/>
        </physiologicalReaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>biotin</name>
            <dbReference type="ChEBI" id="CHEBI:57586"/>
        </cofactor>
    </comment>
    <comment type="pathway">
        <text evidence="3">Metabolic intermediate metabolism; propanoyl-CoA degradation; succinyl-CoA from propanoyl-CoA: step 1/3.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Mitochondrion matrix</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="6.4.1.3" evidence="4"/>
    <dbReference type="AlphaFoldDB" id="A0A452HJL4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017231.1">
        <property type="protein sequence ID" value="ENSGAGP00000015082.1"/>
        <property type="gene ID" value="ENSGAGG00000010472.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00945">
        <property type="reaction ID" value="UER00908"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005759">
        <property type="term" value="C:mitochondrial matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004658">
        <property type="term" value="F:propionyl-CoA carboxylase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016042">
        <property type="term" value="P:lipid catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd06850">
        <property type="entry name" value="biotinyl_domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1490.20:FF:000003">
        <property type="entry name" value="acetyl-CoA carboxylase isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.470.20:FF:000028">
        <property type="entry name" value="Methylcrotonoyl-CoA carboxylase subunit alpha, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.50.100">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.700.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1490.20">
        <property type="entry name" value="ATP-grasp fold, A domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.470.20">
        <property type="entry name" value="ATP-grasp fold, B domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011761">
        <property type="entry name" value="ATP-grasp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013815">
        <property type="entry name" value="ATP_grasp_subdomain_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005481">
        <property type="entry name" value="BC-like_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001882">
        <property type="entry name" value="Biotin_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050856">
        <property type="entry name" value="Biotin_carboxylase_complex"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011764">
        <property type="entry name" value="Biotin_carboxylation_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005482">
        <property type="entry name" value="Biotin_COase_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000089">
        <property type="entry name" value="Biotin_lipoyl"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005479">
        <property type="entry name" value="CPAse_ATP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041265">
        <property type="entry name" value="PCC_BT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016185">
        <property type="entry name" value="PreATP-grasp_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011054">
        <property type="entry name" value="Rudment_hybrid_motif"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011053">
        <property type="entry name" value="Single_hybrid_motif"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18866">
        <property type="entry name" value="CARBOXYLASE:PYRUVATE/ACETYL-COA/PROPIONYL-COA CARBOXYLASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18866:SF33">
        <property type="entry name" value="METHYLCROTONOYL-COA CARBOXYLASE SUBUNIT ALPHA, MITOCHONDRIAL-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02785">
        <property type="entry name" value="Biotin_carb_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00289">
        <property type="entry name" value="Biotin_carb_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02786">
        <property type="entry name" value="CPSase_L_D2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18140">
        <property type="entry name" value="PCC_BT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00878">
        <property type="entry name" value="Biotin_carb_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56059">
        <property type="entry name" value="Glutathione synthetase ATP-binding domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52440">
        <property type="entry name" value="PreATP-grasp domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51246">
        <property type="entry name" value="Rudiment single hybrid motif"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51230">
        <property type="entry name" value="Single hybrid motif"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50975">
        <property type="entry name" value="ATP_GRASP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50979">
        <property type="entry name" value="BC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00188">
        <property type="entry name" value="BIOTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50968">
        <property type="entry name" value="BIOTINYL_LIPOYL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00866">
        <property type="entry name" value="CPSASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00867">
        <property type="entry name" value="CPSASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="9 19" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="15" id="KW-0092">Biotin</keyword>
    <keyword evidence="6" id="KW-0436">Ligase</keyword>
    <keyword evidence="12" id="KW-0442">Lipid degradation</keyword>
    <keyword evidence="13" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="10" id="KW-0460">Magnesium</keyword>
    <keyword evidence="14" id="KW-0464">Manganese</keyword>
    <keyword evidence="7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8 19" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="11" id="KW-0809">Transit peptide</keyword>
    <feature type="domain" description="Biotin carboxylation" evidence="22">
        <location>
            <begin position="1"/>
            <end position="446"/>
        </location>
    </feature>
    <feature type="domain" description="ATP-grasp" evidence="21">
        <location>
            <begin position="118"/>
            <end position="315"/>
        </location>
    </feature>
    <feature type="domain" description="Lipoyl-binding" evidence="20">
        <location>
            <begin position="541"/>
            <end position="618"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001953"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004305"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005060"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00013050"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00018058"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022598"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022963"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023211"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023267"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00031557"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00048208"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00049495"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00409"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50975"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS50979"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015082.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="618" mass="68536" checksum="D27D3F41FAA6355A" modified="2019-05-08" version="1">MPNFKHVIRPTNYNRTNHMLKLKFRTNVPVISGSCASVHVKMADEAVCVGPAPTSKSYLNMDAIMEAIKKTRAQAVHPGYGFLSENKEFARRLASEGVTFVGPDTHAIQAMGDKIESKLLAKNAKVNTIPGFDGVVKDADEAVRIAREIGYPVMIKASAGGGGKGMRIAWNDEETREGFRFSSQEAASSFGDDRLLIEKFIDNPRHIEIQVLADKHDNALWLNERECSIQRRNQKVVEEAPSTFLDPETRRAMGEQAVALAKAVKYSSAGTVEFLVDTKKNFYFLEMNTRLQVEHPITECITGLDLVQEMIRVAKGYPLRHKQADIPINGWAVECRVYAEDPYKSFGLPSIGRLSRYQEPLHVSNVRVDSGIQEGSDISIYYDPMISKLITYGSDRAEALQRMEEALDNYVIRGVTHNIALLREVIIHPRFVQGDVSTKFLPDVYPDGFKGHKLTDLEKRELLAIATSLYMAEQLRSQQFLGSPRISLAKLDLNNWELSIQLGDGTYSVTVSNNGSTFSVEVDGMKLNVTSEWNLASPLLSVTIDGTQRTIQCLSRDAAGNISIQFLGTVVSEGQEICVIEAMKMQNSMIAAKTGKVKALHCKAGETVGEGDLLIELE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJL5</accession>
    <name>A0A452HJL5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">C2H2-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJL5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017254.1">
        <property type="protein sequence ID" value="ENSGAGP00000015104.1"/>
        <property type="gene ID" value="ENSGAGG00000011399.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031981">
        <property type="term" value="C:nuclear lumen"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0022603">
        <property type="term" value="P:regulation of anatomical structure morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000029">
        <property type="entry name" value="GLI family zinc finger 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000295">
        <property type="entry name" value="zinc finger protein 19"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002343">
        <property type="entry name" value="Zinc finger protein 33A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000690">
        <property type="entry name" value="Zinc finger protein 354C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:002090">
        <property type="entry name" value="Zinc finger protein 473"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000176">
        <property type="entry name" value="zinc finger protein 70"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000502">
        <property type="entry name" value="Zinc finger protein 710"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23226:SF170">
        <property type="entry name" value="MYONEURIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23226">
        <property type="entry name" value="ZINC FINGER AND SCAN DOMAIN-CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="7"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="2" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <keyword evidence="4 7" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="9">
        <location>
            <begin position="42"/>
            <end position="69"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="9">
        <location>
            <begin position="70"/>
            <end position="97"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="9">
        <location>
            <begin position="98"/>
            <end position="125"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="9">
        <location>
            <begin position="126"/>
            <end position="153"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="9">
        <location>
            <begin position="154"/>
            <end position="181"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="9">
        <location>
            <begin position="182"/>
            <end position="209"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="9">
        <location>
            <begin position="210"/>
            <end position="240"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1"/>
            <end position="25"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="14"/>
            <end position="25"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006991"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015104.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="249" mass="27930" checksum="B68E2C313B07A858" modified="2019-05-08" version="1">MNSQSFSLPTMVPSSSVINSSVTSNLPTNNPPSKAVLISKPFKCSECGRSFAQLTELELHEMSHAAERPYRCELCGKTFAQTSALVKHQRVHTGEKPYKCPMCGKTFALSSGLVLHKRIHTGERPHTCPLCGKKFISSSHLALHLRSHTGERKYKCSVCGKLFLQSSHLVRHKAIHTGEKPFKCEDCGKNFGRASHLKTHRRVHTGERPFKCTQCEKAFTQKLHMRTHSGERPYHLGDTENEWPVNVGI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJL6</accession>
    <name>A0A452HJL6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Bromo domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HJL6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017262.1">
        <property type="protein sequence ID" value="ENSGAGP00000015110.1"/>
        <property type="gene ID" value="ENSGAGG00000010274.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05513">
        <property type="entry name" value="Bromo_brd7_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.920.10:FF:000022">
        <property type="entry name" value="Putative bromodomain-containing protein 9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.920.10">
        <property type="entry name" value="Bromodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001487">
        <property type="entry name" value="Bromodomain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036427">
        <property type="entry name" value="Bromodomain-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051831">
        <property type="entry name" value="Bromodomain_contain_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021900">
        <property type="entry name" value="DUF3512"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22881">
        <property type="entry name" value="BROMODOMAIN CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22881:SF12">
        <property type="entry name" value="BROMODOMAIN-CONTAINING PROTEIN 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00439">
        <property type="entry name" value="Bromodomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12024">
        <property type="entry name" value="DUF3512"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00503">
        <property type="entry name" value="BROMODOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00297">
        <property type="entry name" value="BROMO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47370">
        <property type="entry name" value="Bromodomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50014">
        <property type="entry name" value="BROMODOMAIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3 6" id="KW-0103">Bromodomain</keyword>
    <keyword evidence="7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0539">Nucleus</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0804">Transcription</keyword>
    <keyword evidence="2" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="Bromo" evidence="9">
        <location>
            <begin position="148"/>
            <end position="218"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="34"/>
            <end position="125"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="251"/>
            <end position="276"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="291"/>
            <end position="318"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="35"/>
            <end position="45"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="47"/>
            <end position="57"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="8">
        <location>
            <begin position="58"/>
            <end position="69"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="70"/>
            <end position="106"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023117"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00035"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50014"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015110.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="536" mass="61298" checksum="62F9A84B28A69828" modified="2019-05-08" version="1">MGKKHKKHKSDKHPYEEYVEKPLKLVLKVGGNEVTELSTGSSGHDSSLYEDKTEHEKHKDRKRKKRKKGEKQIPGEEKEKRKRKVKEDKKKRDREHAESEGERELKCQTPVRLELPPEKPLASSLAKQEEVEQTPLQEALNQLVRQLQRKDPNAFFSFPVTDFIAPGYSMIIKHPMDFSTMKEKIKNNGYQSIEELKDNFRLMCTNAMIYNKPETIYYKAAKKLLHSGMKILSQERIQSLKQSIDFMADLQQSRKQKDKMELQGNGEDDNLLGKDEDSMEADIKAFKTPNKEHKKKDKDLLEEKLRNNNLEREQEQIDRIIRESGGKLTRRLVNSQCEFERRKPDGTTTLGLLNPFDPNAGEPGYSPMKLGMTAGRLQSGVNTLQGFKEDKRNKVTPVVYLNYGPYSSYAPAYDSTFANISKDDSDLIYSTYGEDSNPECFSIHDFLAKSQDYPCVMADGLLDVLTKGGHSRSLRELETPMVDAEGQHGRSDTIKDVEIIETDAANKLDSNNDRLTALKAVTNFGIHVEEFDSEGK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJL9</accession>
    <name>A0A452HJL9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Propionyl-CoA carboxylase alpha chain, mitochondrial</fullName>
            <ecNumber evidence="4">6.4.1.3</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="17">Propanoyl-CoA:carbon dioxide ligase subunit alpha</fullName>
        </alternativeName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>butanoyl-CoA + hydrogencarbonate + ATP = (2S)-ethylmalonyl-CoA + ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:59520"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:17544"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:57371"/>
            <dbReference type="ChEBI" id="CHEBI:60909"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="18">
            <dbReference type="Rhea" id="RHEA:59521"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>propanoyl-CoA + hydrogencarbonate + ATP = (S)-methylmalonyl-CoA + ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:23720"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:17544"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:57327"/>
            <dbReference type="ChEBI" id="CHEBI:57392"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="6.4.1.3"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="19">
            <dbReference type="Rhea" id="RHEA:23721"/>
        </physiologicalReaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>biotin</name>
            <dbReference type="ChEBI" id="CHEBI:57586"/>
        </cofactor>
    </comment>
    <comment type="pathway">
        <text evidence="3">Metabolic intermediate metabolism; propanoyl-CoA degradation; succinyl-CoA from propanoyl-CoA: step 1/3.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Mitochondrion matrix</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="6.4.1.3" evidence="4"/>
    <dbReference type="AlphaFoldDB" id="A0A452HJL9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017236.1">
        <property type="protein sequence ID" value="ENSGAGP00000015087.1"/>
        <property type="gene ID" value="ENSGAGG00000010472.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00945">
        <property type="reaction ID" value="UER00908"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005759">
        <property type="term" value="C:mitochondrial matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004658">
        <property type="term" value="F:propionyl-CoA carboxylase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016042">
        <property type="term" value="P:lipid catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd06850">
        <property type="entry name" value="biotinyl_domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1490.20:FF:000003">
        <property type="entry name" value="acetyl-CoA carboxylase isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.50.100:FF:000029">
        <property type="entry name" value="propionyl-CoA carboxylase alpha chain, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.50.100">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.700.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1490.20">
        <property type="entry name" value="ATP-grasp fold, A domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.470.20">
        <property type="entry name" value="ATP-grasp fold, B domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011761">
        <property type="entry name" value="ATP-grasp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013815">
        <property type="entry name" value="ATP_grasp_subdomain_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005481">
        <property type="entry name" value="BC-like_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001882">
        <property type="entry name" value="Biotin_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050856">
        <property type="entry name" value="Biotin_carboxylase_complex"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011764">
        <property type="entry name" value="Biotin_carboxylation_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005482">
        <property type="entry name" value="Biotin_COase_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000089">
        <property type="entry name" value="Biotin_lipoyl"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005479">
        <property type="entry name" value="CPAse_ATP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041265">
        <property type="entry name" value="PCC_BT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016185">
        <property type="entry name" value="PreATP-grasp_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011054">
        <property type="entry name" value="Rudment_hybrid_motif"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011053">
        <property type="entry name" value="Single_hybrid_motif"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18866">
        <property type="entry name" value="CARBOXYLASE:PYRUVATE/ACETYL-COA/PROPIONYL-COA CARBOXYLASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18866:SF33">
        <property type="entry name" value="METHYLCROTONOYL-COA CARBOXYLASE SUBUNIT ALPHA, MITOCHONDRIAL-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02785">
        <property type="entry name" value="Biotin_carb_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00289">
        <property type="entry name" value="Biotin_carb_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00364">
        <property type="entry name" value="Biotin_lipoyl"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02786">
        <property type="entry name" value="CPSase_L_D2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18140">
        <property type="entry name" value="PCC_BT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00878">
        <property type="entry name" value="Biotin_carb_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56059">
        <property type="entry name" value="Glutathione synthetase ATP-binding domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52440">
        <property type="entry name" value="PreATP-grasp domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51246">
        <property type="entry name" value="Rudiment single hybrid motif"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51230">
        <property type="entry name" value="Single hybrid motif"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50975">
        <property type="entry name" value="ATP_GRASP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50979">
        <property type="entry name" value="BC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00188">
        <property type="entry name" value="BIOTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50968">
        <property type="entry name" value="BIOTINYL_LIPOYL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00866">
        <property type="entry name" value="CPSASE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00867">
        <property type="entry name" value="CPSASE_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="9 20" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="16" id="KW-0092">Biotin</keyword>
    <keyword evidence="6" id="KW-0436">Ligase</keyword>
    <keyword evidence="12" id="KW-0442">Lipid degradation</keyword>
    <keyword evidence="13" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="10" id="KW-0460">Magnesium</keyword>
    <keyword evidence="15" id="KW-0464">Manganese</keyword>
    <keyword evidence="7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="14" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="8 20" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="11" id="KW-0809">Transit peptide</keyword>
    <feature type="domain" description="Biotin carboxylation" evidence="23">
        <location>
            <begin position="1"/>
            <end position="405"/>
        </location>
    </feature>
    <feature type="domain" description="ATP-grasp" evidence="22">
        <location>
            <begin position="105"/>
            <end position="302"/>
        </location>
    </feature>
    <feature type="domain" description="Lipoyl-binding" evidence="21">
        <location>
            <begin position="552"/>
            <end position="627"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001953"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004305"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005060"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00013050"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00018058"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022598"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022963"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023211"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023267"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00031557"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00048208"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00049495"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00409"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50968"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS50975"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS50979"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015087.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="627" mass="69524" checksum="C005EE4D0F939CBF" modified="2019-05-08" version="1">TGHECLLLSIKTLGYPGYIILPFQVHVKMADEAVCVGPAPTSKSYLNMDAIMEAIKKTRAQAVHPGYGFLSENKEFARRLASEGVTFVGPDTHAIQAMGDKIESKLLAKNAKVNTIPGFDGVVKDADEAVRIAREIGYPVMIKASAGGGGKGMRIAWNDEETREGFRFSSQEAASSFGDDRLLIEKFIDNPRHIEIQVLADKHDNALWLNERECSIQRRNQKVVEEAPSTFLDPETRRAMGEQAVALAKAVKYSSAGTVEFLVDTKKNFYFLEMNTRLQVEHPITECITGLDLVQEMIRVAKGYPLRHKQADIPINVVWGGRLYVRVDSGIQEGSDISIYYDPMISKLITYGSDRAEALQRMEEALDNYVIRGVTHNIALLREVIIHPRFVQGDVSTKFLPDVYPDGFKGHKLTDLEKRELLAIATSLYMAEQLRSQQFLGSPRISLAKLDLNNWELSIQLGDGTYSVTVSNNGSTFSVEVDGMKLNVTSEWNLASPLLSVTIDGTQRTIQKFTRDTGLNILKYFSFCLFLKYKLRILTKLAAELSKYMPEKAVEDTTSILRSPMPGAVVAVSVKPGDLVSEGQEICVIEAMKMQNSMIAAKTGKVKALHCKAGETVGEGDLLIELE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMH2</accession>
    <name>A0A452IMH2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Lipase</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1 7">Belongs to the AB hydrolase superfamily. Lipase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMH2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000032998.1">
        <property type="protein sequence ID" value="ENSGAGP00000029047.1"/>
        <property type="gene ID" value="ENSGAGG00000020969.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016788">
        <property type="term" value="F:hydrolase activity, acting on ester bonds"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016042">
        <property type="term" value="P:lipid catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1820:FF:000057">
        <property type="entry name" value="Lipase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1820">
        <property type="entry name" value="alpha/beta hydrolase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000073">
        <property type="entry name" value="AB_hydrolase_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029058">
        <property type="entry name" value="AB_hydrolase_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025483">
        <property type="entry name" value="Lipase_euk"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11005">
        <property type="entry name" value="LYSOSOMAL ACID LIPASE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00561">
        <property type="entry name" value="Abhydrolase_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000862">
        <property type="entry name" value="Steryl_ester_lip"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53474">
        <property type="entry name" value="alpha/beta-Hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="3 7" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="4 7" id="KW-0442">Lipid degradation</keyword>
    <keyword evidence="5" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0732">Signal</keyword>
    <feature type="domain" description="AB hydrolase-1" evidence="9">
        <location>
            <begin position="87"/>
            <end position="360"/>
        </location>
    </feature>
    <feature type="active site" description="Nucleophile" evidence="8">
        <location>
            <position position="181"/>
        </location>
    </feature>
    <feature type="active site" description="Charge relay system" evidence="8">
        <location>
            <position position="325"/>
        </location>
    </feature>
    <feature type="active site" description="Charge relay system" evidence="8">
        <location>
            <position position="354"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00010701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022963"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PIRNR" id="PIRNR000862"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PIRSR" id="PIRSR000862-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF00561"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029047.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="379" mass="43872" checksum="E026A2C3806B5D4A" modified="2019-05-08" version="1">MYVSRNVINRRIFFHRNKMWWFIAVVCLIQAITSSEELIKKRNMNPETFMNISQLICYEGYPSEEYEVLTEDGYYLKINRIPYGPKPAVLLQHGLFGQGSNWIENLANNSLGFMLADAGYDVWIGNSRGTTWSQRHQKFSVGQEEFWDFSFHEMGIYDLQAMINFILQETGQKQLYYVGYSQGGSMGFIAFSSMPQLAQKVKMFFALAPAVTVGHSKSPVVKMLLMPEEDLLGKREFRWWSKTMREFAAKICCYDLPNRLCANAFFFAGGFNEKNINMVCAVKSGEFKYFDYGSENPAKYNQSVPPFYKIEDMTVPTAVWSGGEDWMVDPKDIDMLLPHITNLIYHKNISDWNHWDFTWGLDAAKRVYSEIIALIGKYP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMH6</accession>
    <name>A0A452IMH6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">2'-phosphotransferase</fullName>
            <ecNumber evidence="3">2.7.1.160</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="2">Catalyzes the last step of tRNA splicing, the transfer of the splice junction 2'-phosphate from ligated tRNA to NAD to produce ADP-ribose 1''-2'' cyclic phosphate.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="7">
            <text>2'-phospho-[ligated tRNA] + NAD(+) = mature tRNA + ADP-alpha-D-ribose 1'',2''-cyclic phosphate + nicotinamide</text>
            <dbReference type="Rhea" id="RHEA:23324"/>
            <dbReference type="Rhea" id="RHEA-COMP:11106"/>
            <dbReference type="Rhea" id="RHEA-COMP:11107"/>
            <dbReference type="ChEBI" id="CHEBI:17154"/>
            <dbReference type="ChEBI" id="CHEBI:57540"/>
            <dbReference type="ChEBI" id="CHEBI:76596"/>
            <dbReference type="ChEBI" id="CHEBI:82883"/>
            <dbReference type="ChEBI" id="CHEBI:85027"/>
            <dbReference type="EC" id="2.7.1.160"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <dbReference type="EC" id="2.7.1.160" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452IMH6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033022.1">
        <property type="protein sequence ID" value="ENSGAGP00000029069.1"/>
        <property type="gene ID" value="ENSGAGG00000021032.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0052751">
        <property type="term" value="F:GDP-mannose hydrolase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000215">
        <property type="term" value="F:tRNA 2'-phosphotransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.79.10:FF:000087">
        <property type="entry name" value="Nudix (nucleoside diphosphate linked moiety X)-type motif 22"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.20.170.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.79.10">
        <property type="entry name" value="Nucleoside Triphosphate Pyrophosphohydrolase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.970">
        <property type="entry name" value="RNA 2'-phosphotransferase, Tpt1/KptA family, N-terminal domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015797">
        <property type="entry name" value="NUDIX_hydrolase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000086">
        <property type="entry name" value="NUDIX_hydrolase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055295">
        <property type="entry name" value="NUDT22/NUDT9-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002745">
        <property type="entry name" value="Ptrans_KptA/Tpt1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042081">
        <property type="entry name" value="RNA_2'-PTrans_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042080">
        <property type="entry name" value="RNA_2'-PTrans_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31835">
        <property type="entry name" value="URIDINE DIPHOSPHATE GLUCOSE PYROPHOSPHATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31835:SF1">
        <property type="entry name" value="URIDINE DIPHOSPHATE GLUCOSE PYROPHOSPHATASE NUDT22"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01885">
        <property type="entry name" value="PTS_2-RNA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56399">
        <property type="entry name" value="ADP-ribosylation"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55811">
        <property type="entry name" value="Nudix"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51462">
        <property type="entry name" value="NUDIX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="6" id="KW-0460">Magnesium</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Nudix hydrolase" evidence="8">
        <location>
            <begin position="114"/>
            <end position="275"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00003343"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012007"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00047949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51462"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029069.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="464" mass="51107" checksum="71D655FE349D620E" modified="2019-05-08" version="1">MDPEISIMLQCPSPKGLAETAVRAELSPAYNRRQLPGGQAWIDAVWEARCRHSPWLFNGSKFRLHSAQLDGGSLTFCLGLTCYKDFLGTNRAGMARHLQQQGRQDFGDSQAYLAEPLGVGAMVHTANDCFVFLRRSLRVGEAPGLVDIPGGHPEPQAVVGDVPEESIRLQDLPRQMVVKEIFTSILREIRDEVNLPLPTLSQPVLLGIARNQTSAGRASAEFYVRCSLTSEQVKQRYEIGGPEAQESTSIIFIKREDVLTLEQTGEMWRELCPSAKGANPVVHLSKTLSYVLRHGAAQLGLEMGADGFVDVAALLSLPRFGGVSVADVRHVVETNEKCRFALRSHPSDGRLQIRANQGHSLQVSELELIPLLEPTALPQTMVHGTYLRHWPAICRGGLSRMGRNHIHLAPGLPGDGHVLSDGIQFYRSANGVILTPGDAEGLLPPRYFQRVLQLRPDRRLLPLE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMH7</accession>
    <name>A0A452IMH7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Nck-associated protein 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="6">Cell projection</location>
            <location evidence="6">Lamellipodium membrane</location>
            <topology evidence="6">Single-pass membrane protein</topology>
            <orientation evidence="6">Cytoplasmic side</orientation>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="7">Belongs to the HEM-1/HEM-2 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMH7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000032977.1">
        <property type="protein sequence ID" value="ENSGAGP00000029028.1"/>
        <property type="gene ID" value="ENSGAGG00000019065.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031258">
        <property type="term" value="C:lamellipodium membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031209">
        <property type="term" value="C:SCAR complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030031">
        <property type="term" value="P:cell projection assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030866">
        <property type="term" value="P:cortical actin cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048812">
        <property type="term" value="P:neuron projection morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019137">
        <property type="entry name" value="Nck-associated_protein-1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12093">
        <property type="entry name" value="NCK-ASSOCIATED PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12093:SF11">
        <property type="entry name" value="NCK-ASSOCIATED PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09735">
        <property type="entry name" value="Nckap1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="1" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="5" id="KW-0966">Cell projection</keyword>
    <keyword evidence="4" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="3" id="KW-1133">Transmembrane helix</keyword>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="551"/>
            <end position="576"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="562"/>
            <end position="576"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00037839"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00037947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00039689"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029028.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="979" mass="112150" checksum="037B518B59609E5E" modified="2019-05-08" version="1">FGFCLTNQLELFEVVNQHENKGNPVSIVYLDFQKTVNFDLTKNYLDLIITYTTLMILLSRIEERKAIIGLYNYAHEMTHGASDREYPRLGQMIVDYENPLKKMMEEFVPHSKSLSDALISLQMVYPRRNLSADQWRNAQLLSLISAPSTMLNPAQSDTMPCEYLSLDAMEKWIIFGFILCHGILNSDATALNLWKLALQSSSCLSLFRDEVFHIHKAAEDLFVNIRGYNKRINDIRECKEAAVSHAGSMHRERRKFLRSALKELATVLSDQPGLLGPKALFVFMALSFARDEIIWLLRHADNMPKKSADDFIDKHIAELIFYMEELRAHVRKYGPVMQRYYVQYLSGFDAVVLNELVQNLSVCPEDESIIMSSFVNTMTSLSVKQVEDGEVFDFRGMRLDWFRLQAYTSVSKASLSLADHRELGKMMNTIIFHTKMVDSLVEMLVETSDLSIFCFYSRAFEKMFQQCLELPSQSRYSIAFPLLCTHFMSCTHELCPEERHHIGDRSLSLCNMFLDEMAKQARNLITDICTEQCTLSDQLLPKHCAKTISQAVNKKSKKQTGKKGEPEREKPGVESMRKNRLVVTNLDKLHTALSELCFSINYVPNMVVWEHTFTPREYLTSHLEIRFTKSIVGMTMYNQATQEIAKPSELLTSVRAYMTVLQSIENYVQIDITRVFNNVLLQQTQHLDSHGEPTITSLYTNWYLETLLRQVSNGHIAYFPAMKAFVNLPTENELTFNAEEYSDISEMRALSELLGPYGMKFLSESLMWHISSQVAELKKLVVENVEVLTQMRTSFDKPDQMAALFKRLSCGVLKRMTIIGVILSFRSLAQELLNLRHFFFFSENISPEEEYKIACLLMVFVAVSLPTLASNVMSQYSPAIEGHCNNIHCLAKAINQIAAALFTIHKGSIEDQLTATKIRLKNQTKTLNICCNSDLPFRCTVIVQESPFLTMDLLESCFPYVLLRNAYHAVYKQSVTSSA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMH8</accession>
    <name>A0A452IMH8_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="7">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the major facilitator superfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMH8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033110.1">
        <property type="protein sequence ID" value="ENSGAGP00000029148.1"/>
        <property type="gene ID" value="ENSGAGG00000021090.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1250.20:FF:000508">
        <property type="entry name" value="Sodium-dependent glucose transporter 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1250.20">
        <property type="entry name" value="MFS general substrate transporter like domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036259">
        <property type="entry name" value="MFS_trans_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23121">
        <property type="entry name" value="SODIUM-DEPENDENT GLUCOSE TRANSPORTER 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23121:SF9">
        <property type="entry name" value="SODIUM-DEPENDENT GLUCOSE TRANSPORTER 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103473">
        <property type="entry name" value="MFS general substrate transporter"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 6" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 6" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 6" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="6"/>
            <end position="28"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="35"/>
            <end position="52"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="135"/>
            <end position="157"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="178"/>
            <end position="205"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="250"/>
            <end position="268"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="274"/>
            <end position="295"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="307"/>
            <end position="325"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="337"/>
            <end position="357"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008335"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029148.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="372" mass="40523" checksum="B3F5DDBE1DF36BA8" modified="2019-05-08" version="1">LYHFLLLYGFWITFIGLTMLGTAVGLYAIPWCKRALLLTALMSVIGTSVGVLDTGGNVLALNIWGAEAGPHMQALHFSFALGAVVAPILTKMALSSFISPNFQTEDEKTNHSVPSDIPNALFGSALKLHSNVSFMWSYIVIGTYILIVSLLFVVLYSRTGLARDKAKSSLQKCQIAKYHYALLILLFLFFFCYVGAEVTYGSYIFNYAEVYADMKESEAASLNSLFWGAFAACRGLAICFATCLYPGTMILLSVIGSTVSSLFLALLSKHPISLWIGTAVYGASMATVFPSGISWIEQYTTIQGKSASLFVVGAALGEMCIPAVVGFLEGKLHNVPVMMYAALGTSVMTAVLFPVMYKLFVYLHLEHFCYVV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMI1</accession>
    <name>A0A452IMI1_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="12">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell projection</location>
            <location evidence="1">Lamellipodium</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMI1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000032982.1">
        <property type="protein sequence ID" value="ENSGAGP00000029033.1"/>
        <property type="gene ID" value="ENSGAGG00000021012.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030027">
        <property type="term" value="C:lamellipodium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035556">
        <property type="term" value="P:intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030032">
        <property type="term" value="P:lamellipodium assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd21265">
        <property type="entry name" value="CH_alphaPIX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd01225">
        <property type="entry name" value="PH_Cool_Pix"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12060">
        <property type="entry name" value="SH3_alphaPIX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.40:FF:000034">
        <property type="entry name" value="Rho guanine nucleotide exchange factor (GEF) 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000094">
        <property type="entry name" value="Rho guanine nucleotide exchange factor 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.390:FF:000001">
        <property type="entry name" value="rho guanine nucleotide exchange factor 7 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.418.10">
        <property type="entry name" value="Calponin-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.900.10">
        <property type="entry name" value="Dbl homology (DH) domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.390">
        <property type="entry name" value="L1 transposable element, trimerization domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035788">
        <property type="entry name" value="AlphaPIX_SH3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001715">
        <property type="entry name" value="CH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036872">
        <property type="entry name" value="CH_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035899">
        <property type="entry name" value="DBL_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000219">
        <property type="entry name" value="DH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001331">
        <property type="entry name" value="GDS_CDC24_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032409">
        <property type="entry name" value="GEF6/7_CC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046376">
        <property type="entry name" value="PH_Cool_Pix"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46026:SF2">
        <property type="entry name" value="RHO GUANINE NUCLEOTIDE EXCHANGE FACTOR 6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46026">
        <property type="entry name" value="RHO-TYPE GUANINE NUCLEOTIDE EXCHANGE FACTOR, ISOFORM F"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16523">
        <property type="entry name" value="betaPIX_CC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00307">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00169">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00621">
        <property type="entry name" value="RhoGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16615">
        <property type="entry name" value="RhoGEF67_u1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16614">
        <property type="entry name" value="RhoGEF67_u2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07653">
        <property type="entry name" value="SH3_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00452">
        <property type="entry name" value="SH3DOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00033">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00233">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00325">
        <property type="entry name" value="RhoGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00326">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47576">
        <property type="entry name" value="Calponin-homology domain, CH-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48065">
        <property type="entry name" value="DBL homology domain (DH-domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50021">
        <property type="entry name" value="CH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00741">
        <property type="entry name" value="DH_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50010">
        <property type="entry name" value="DH_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0966">Cell projection</keyword>
    <keyword evidence="6" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0344">Guanine-nucleotide releasing factor</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 5" id="KW-0728">SH3 domain</keyword>
    <feature type="domain" description="Calponin-homology (CH)" evidence="11">
        <location>
            <begin position="1"/>
            <end position="111"/>
        </location>
    </feature>
    <feature type="domain" description="SH3" evidence="8">
        <location>
            <begin position="161"/>
            <end position="220"/>
        </location>
    </feature>
    <feature type="domain" description="DH" evidence="10">
        <location>
            <begin position="242"/>
            <end position="324"/>
        </location>
    </feature>
    <feature type="domain" description="DH" evidence="10">
        <location>
            <begin position="329"/>
            <end position="381"/>
        </location>
    </feature>
    <feature type="domain" description="PH" evidence="9">
        <location>
            <begin position="403"/>
            <end position="508"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="111"/>
            <end position="148"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="6">
        <location>
            <begin position="674"/>
            <end position="712"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="7">
        <location>
            <begin position="123"/>
            <end position="132"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="7">
        <location>
            <begin position="133"/>
            <end position="144"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004510"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022658"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50010"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029033.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="729" mass="82144" checksum="EC51187B1FE2D466" modified="2019-05-08" version="1">MNPEERLVTWLISLGVLNSPKKMIGDPEQFLKVSLKDGVVLCKLVNRLLPGHVEKYCLEPKNEADCISNIQAFLKGCATLKVELFDARDLYAGENFSKVLSTLAAVNKATEDQPSERPCSRLSSHSAAASSSQTNLHGTTSITSRVLRRQSKPVEMVENGSYQLVVKARFNFKQTNEDELSVNKGDIIYVTRVEEGGWWEGTLNGKTGWFPSNYVKEIKSADKPLSPKALKGQESPQLTKNYYTVVLQSILETERDYAKELQSLLGTYLRPLQSYDKLSVVDIASLLGNVEEICSFQLTLNQALEECANDELEKFMENQGASNPGILILTTSLSKPFVRLDKYVTLLQELERHMEEAHPDHEDVLKAFTSFKSLMSQCQELRKRKQLELQILSESIPGWEGEDIKTMGNVIYMSQVMVQCGASEEKEERYFMLFSNVLLMLSASPRMTGFIYQGKLPLTGMTLTKLEGTEGTEHAFEITGNMIERITVSCNSSQDLHEWLDHLQRLTKGTISKTQSWSAHSTFSSTGQLRGPLEPPKILKPWTLSCLRPAPPLRPSAALGYKERMSYILKDSSKSPKTMKKFLPKRKTERKASDEEFVIRKSTAALEEDAQILKVIEAYCTGAGFQQTLSSGSRKDSLPQVLLPEEEKIIIEETRSNGQTVTEEKSLVDTVYALKDQVKELKQESKKMKQCLEEELKSRKDLEKLVRRLLKQTDECARDETGRKSSVLA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMI9</accession>
    <name>A0A452IMI9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">Mannose receptor C-type 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="14">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000033020.1">
        <property type="protein sequence ID" value="ENSGAGP00000029067.1"/>
        <property type="gene ID" value="ENSGAGG00000020766.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030246">
        <property type="term" value="F:carbohydrate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006897">
        <property type="term" value="P:endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd23407">
        <property type="entry name" value="beta-trefoil_Ricin_MRC1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00037">
        <property type="entry name" value="CLECT"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="CDD" id="cd00062">
        <property type="entry name" value="FN2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.80.10.50:FF:000032">
        <property type="entry name" value="macrophage mannose receptor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000014">
        <property type="entry name" value="Macrophage mannose receptor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000016">
        <property type="entry name" value="macrophage mannose receptor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000023">
        <property type="entry name" value="Macrophage mannose receptor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000031">
        <property type="entry name" value="macrophage mannose receptor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000025">
        <property type="entry name" value="Mannose receptor C-type 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000030">
        <property type="entry name" value="Mannose receptor C-type 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000027">
        <property type="entry name" value="Mannose receptor, C type 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.80.10.50">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.10.10">
        <property type="entry name" value="Fibronectin, type II, collagen-binding"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.100.10">
        <property type="entry name" value="Mannose-Binding Protein A, subunit A"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001304">
        <property type="entry name" value="C-type_lectin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016186">
        <property type="entry name" value="C-type_lectin-like/link_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050111">
        <property type="entry name" value="C-type_lectin/snaclec_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018378">
        <property type="entry name" value="C-type_lectin_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016187">
        <property type="entry name" value="CTDL_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000562">
        <property type="entry name" value="FN_type2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036943">
        <property type="entry name" value="FN_type2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035992">
        <property type="entry name" value="Ricin_B-like_lectins"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000772">
        <property type="entry name" value="Ricin_B_lectin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22803">
        <property type="entry name" value="MANNOSE, PHOSPHOLIPASE, LECTIN RECEPTOR RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24562">
        <property type="entry name" value="CysR_MRC2_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00040">
        <property type="entry name" value="fn2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00059">
        <property type="entry name" value="Lectin_C"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00013">
        <property type="entry name" value="FNTYPEII"/>
    </dbReference>
    <dbReference type="SMART" id="SM00034">
        <property type="entry name" value="CLECT"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SMART" id="SM00059">
        <property type="entry name" value="FN2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00458">
        <property type="entry name" value="RICIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56436">
        <property type="entry name" value="C-type lectin-like"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50370">
        <property type="entry name" value="Ricin B-like lectins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00615">
        <property type="entry name" value="C_TYPE_LECTIN_1"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50041">
        <property type="entry name" value="C_TYPE_LECTIN_2"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51092">
        <property type="entry name" value="FN2_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50231">
        <property type="entry name" value="RICIN_B_LECTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8" id="KW-0106">Calcium</keyword>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0254">Endocytosis</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6" id="KW-0430">Lectin</keyword>
    <keyword evidence="10 15" id="KW-0472">Membrane</keyword>
    <keyword evidence="12" id="KW-0675">Receptor</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="5 16" id="KW-0732">Signal</keyword>
    <keyword evidence="4 15" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9 15" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="16">
        <location>
            <begin position="1"/>
            <end position="18"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019280096" description="Mannose receptor C-type 1" evidence="16">
        <location>
            <begin position="19"/>
            <end position="1412"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="1343"/>
            <end position="1366"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-II" evidence="18">
        <location>
            <begin position="163"/>
            <end position="214"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="17">
        <location>
            <begin position="223"/>
            <end position="338"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="17">
        <location>
            <begin position="366"/>
            <end position="483"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="17">
        <location>
            <begin position="508"/>
            <end position="623"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="17">
        <location>
            <begin position="652"/>
            <end position="771"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="17">
        <location>
            <begin position="788"/>
            <end position="904"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="17">
        <location>
            <begin position="932"/>
            <end position="1055"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="17">
        <location>
            <begin position="1077"/>
            <end position="1188"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="17">
        <location>
            <begin position="1215"/>
            <end position="1333"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022583"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022734"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50041"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS51092"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029067.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="21">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1412" mass="160477" checksum="9483E749E38997D0" modified="2019-05-08" version="1">MAVYLLLIFLSLITTSIQLPGSGIFLIHNEDHKLCVKAQSSNSVIIVTCNQDDDSQKFRWVSDHQILSMALKLCLGVPSMKDQAAITLYPCNKTSELQQWECRNKTLLAIQGEDLFFNSAHGEERKVMLSKGSGSWNRWKIHGTTENLCSQGYEDMFTLLGNAFGAPCVFPFKFNSTWYAECTKDRKAGNKQGFPWCATSTDFDEDSLFGYCPLKDFWITNPFTGTHYQINSNSALTWHQARKSCQQQNAELLSITELHEQTYLTGLFGRANSELWIGLNRLNSNGGWQWIGGSPFRYLNWAPGSPSLESEKICGLLQSGNGKWKNQQCDQKLGYICKKGNSSLDSFIIPSGDFRPIKCPSGWIPYSGHCYMLYRAPKIWKDALSSCRKEGGDLASIHNIEEYSFTVSQLGYKPTDELWIGLNDLKIHMYFDWSDGTPVTYTKWLHEEPSHANNRKEDCVSMKGEDGYWADDVCEKKLGYICKMKPLAEESEKVEVIIPGCQKGWIRHGFYCYSVGQTSVTFSEAKKICEGNKGYLVTVKDRYEQAFLTVLIGFSPVKYFWIALSNVEEQGTFKWADGEAVLFTHWNSGMPGREPGCVAMRTGTTAGLWDVVNCEEKSMFLCKQLVEGATPPPVPTTTPAPSCPDGWHSRAHSSSCFKHFQNGRKHMKTWFDARDFCRAIGGDLASIHSEEDQNVINSLAHYYQSYWMGMSAFDPDGGFTWSDGSPVNYENWADGEPNNYDGNEKCAMFNEYTDMQWNDMFCEKFGDWTCQIKKEYTYKVSEDGWIIYEDKAYYFSHETLPMEKAREYCKKNSGDLVVIEGESKRKFLWKYSFYNDFRESLYIGLSVSLDKKFSWIDGTPLNYVAWSPNEPNFANNDENCVVMYTQTGMWNDLNCGAPNSFICERLNNITHSTVAPTSPAPLGGCPQNWFLFNNKCFKIFGSSENESLTWHAARTACINLEGNLASIPSEXAAASFLITHLKDASTDTWIGLNDVNSEYTFLWTDGSGVSYTSWAKGFPNYRSEADCVMMIKDPIEEAGKWKDEDCQMNKSYICQRNADPKLYNFQTTVPVSGFTLYGNSSYSLISSRMNWEEAQKNCKAEYSELASILDPYSQSFLWLQVLKIGQPVWIGLNSNVTSGFYKWTDNWRIKYSKWASEEPKQNLACVYLDVDGTWKTGSCDENYCSVCKRSDAPTELPQLPGKCPTSDIRRSWIPFHGHCYYIESSEMESWPQSSMECVRLGASLVSIENMAEMNFLLHHLDAFASDSRQFWIGMYKNVDGNYLTFNNNSAVEFVNWKEREPTDGYHEYCVDMDASDGGWRSYYCSADKRFICKIPKTVPAHHLSGMAVILVLLILVGAGLAAYIFFKKKRHNQLPIDVTFDNTLYCNRDAVPAASDSKYLVANIEQNEQAML</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMJ3</accession>
    <name>A0A452IMJ3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Macro domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="2">
            <text>alpha-NAD(+) + H2O = ADP-D-ribose + nicotinamide + H(+)</text>
            <dbReference type="Rhea" id="RHEA:68792"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:17154"/>
            <dbReference type="ChEBI" id="CHEBI:57967"/>
            <dbReference type="ChEBI" id="CHEBI:77017"/>
        </reaction>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the MacroD-type family. MacroD1/2-like subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMJ3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033043.1">
        <property type="protein sequence ID" value="ENSGAGP00000029088.1"/>
        <property type="gene ID" value="ENSGAGG00000021044.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005654">
        <property type="term" value="C:nucleoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140293">
        <property type="term" value="F:ADP-ribosylglutamate hydrolase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006974">
        <property type="term" value="P:DNA damage response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140291">
        <property type="term" value="P:peptidyl-glutamate ADP-deribosylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042278">
        <property type="term" value="P:purine nucleoside metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd02908">
        <property type="entry name" value="Macro_OAADPr_deacetylase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.220.10:FF:000003">
        <property type="entry name" value="O-acetyl-ADP-ribose deacetylase MACROD2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.220.10">
        <property type="entry name" value="Leucine Aminopeptidase, subunit E, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002589">
        <property type="entry name" value="Macro_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043472">
        <property type="entry name" value="Macro_dom-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11106:SF93">
        <property type="entry name" value="ADP-RIBOSE GLYCOHYDROLASE MACROD1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11106">
        <property type="entry name" value="GANGLIOSIDE INDUCED DIFFERENTIATION ASSOCIATED PROTEIN 2-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01661">
        <property type="entry name" value="Macro"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00506">
        <property type="entry name" value="A1pp"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52949">
        <property type="entry name" value="Macro domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51154">
        <property type="entry name" value="MACRO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="1" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="4">
        <location>
            <begin position="1"/>
            <end position="18"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019163918" description="Macro domain-containing protein" evidence="4">
        <location>
            <begin position="19"/>
            <end position="316"/>
        </location>
    </feature>
    <feature type="domain" description="Macro" evidence="5">
        <location>
            <begin position="133"/>
            <end position="304"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00049015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00093460"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS51154"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029088.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="316" mass="35348" checksum="1746BF9881DF8F41" modified="2019-05-08" version="1">MPGVLMTLLGRRLSGALGQLLGGWGKGRIPWVLGSVRRGTWSCREPSTSSLPCGTVQLVSRAFRSWAAASPLGMAAKVDLTTSTDWKEAKTYLKGLNQKQRREHYYTKDFVTLKDIDTWKKMAKSARLKQPEETKFPKDNHLNEKISLVRRDITKLEVDAIVNAVDGCIHRAAGELLKEECRSLNGCETGKAKLSCGYRLPAKYVIHAVGPIAQGEPSAAQEKELENCYKNSLKLAVENKLRTVAFPCLSTGVFGYPSDAAAEVVLRTLREWLEVNKEKVDRLVICVFQEKDEEIYKEKLPFYFPIDTTEQLAAQP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMJ5</accession>
    <name>A0A452IMJ5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">Mannose receptor C-type 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="14">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000033005.1">
        <property type="protein sequence ID" value="ENSGAGP00000029053.1"/>
        <property type="gene ID" value="ENSGAGG00000020766.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030246">
        <property type="term" value="F:carbohydrate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006897">
        <property type="term" value="P:endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd23407">
        <property type="entry name" value="beta-trefoil_Ricin_MRC1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00037">
        <property type="entry name" value="CLECT"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="CDD" id="cd00062">
        <property type="entry name" value="FN2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.80.10.50:FF:000032">
        <property type="entry name" value="macrophage mannose receptor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000014">
        <property type="entry name" value="Macrophage mannose receptor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000016">
        <property type="entry name" value="macrophage mannose receptor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000023">
        <property type="entry name" value="Macrophage mannose receptor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000031">
        <property type="entry name" value="macrophage mannose receptor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000025">
        <property type="entry name" value="Mannose receptor C-type 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000030">
        <property type="entry name" value="Mannose receptor C-type 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000027">
        <property type="entry name" value="Mannose receptor, C type 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.80.10.50">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.10.10">
        <property type="entry name" value="Fibronectin, type II, collagen-binding"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.100.10">
        <property type="entry name" value="Mannose-Binding Protein A, subunit A"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001304">
        <property type="entry name" value="C-type_lectin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016186">
        <property type="entry name" value="C-type_lectin-like/link_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050111">
        <property type="entry name" value="C-type_lectin/snaclec_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018378">
        <property type="entry name" value="C-type_lectin_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016187">
        <property type="entry name" value="CTDL_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000562">
        <property type="entry name" value="FN_type2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036943">
        <property type="entry name" value="FN_type2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035992">
        <property type="entry name" value="Ricin_B-like_lectins"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000772">
        <property type="entry name" value="Ricin_B_lectin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22803">
        <property type="entry name" value="MANNOSE, PHOSPHOLIPASE, LECTIN RECEPTOR RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24562">
        <property type="entry name" value="CysR_MRC2_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00040">
        <property type="entry name" value="fn2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00059">
        <property type="entry name" value="Lectin_C"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00013">
        <property type="entry name" value="FNTYPEII"/>
    </dbReference>
    <dbReference type="SMART" id="SM00034">
        <property type="entry name" value="CLECT"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SMART" id="SM00059">
        <property type="entry name" value="FN2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00458">
        <property type="entry name" value="RICIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56436">
        <property type="entry name" value="C-type lectin-like"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50370">
        <property type="entry name" value="Ricin B-like lectins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00615">
        <property type="entry name" value="C_TYPE_LECTIN_1"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50041">
        <property type="entry name" value="C_TYPE_LECTIN_2"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51092">
        <property type="entry name" value="FN2_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50231">
        <property type="entry name" value="RICIN_B_LECTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8" id="KW-0106">Calcium</keyword>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0254">Endocytosis</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6" id="KW-0430">Lectin</keyword>
    <keyword evidence="10 15" id="KW-0472">Membrane</keyword>
    <keyword evidence="12" id="KW-0675">Receptor</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="5 16" id="KW-0732">Signal</keyword>
    <keyword evidence="4 15" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9 15" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="16">
        <location>
            <begin position="1"/>
            <end position="18"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019314034" description="Mannose receptor C-type 1" evidence="16">
        <location>
            <begin position="19"/>
            <end position="1446"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="1377"/>
            <end position="1400"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-II" evidence="18">
        <location>
            <begin position="163"/>
            <end position="214"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="17">
        <location>
            <begin position="223"/>
            <end position="338"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="17">
        <location>
            <begin position="366"/>
            <end position="483"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="17">
        <location>
            <begin position="508"/>
            <end position="623"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="17">
        <location>
            <begin position="652"/>
            <end position="773"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="17">
        <location>
            <begin position="802"/>
            <end position="918"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="17">
        <location>
            <begin position="946"/>
            <end position="1069"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="17">
        <location>
            <begin position="1091"/>
            <end position="1202"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="17">
        <location>
            <begin position="1229"/>
            <end position="1347"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022583"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022734"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50041"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS51092"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029053.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="21">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1446" mass="164287" checksum="CC8669DD6AB3F874" modified="2019-05-08" version="1">MAVYLLLIFLSLITTSIQLPGSGIFLIHNEDHKLCVKAQSSNSVIIVTCNQDDDSQKFRWVSDHQILSMALKLCLGVPSMKDQAAITLYPCNKTSELQQWECRNKTLLAIQGEDLFFNSAHGEERKVMLSKGSGSWNRWKIHGTTENLCSQGYEDMFTLLGNAFGAPCVFPFKFNSTWYAECTKDRKAGNKQGFPWCATSTDFDEDSLFGYCPLKDFWITNPFTGTHYQINSNSALTWHQARKSCQQQNAELLSITELHEQTYLTGLFGRANSELWIGLNRLNSNGGWQWIGGSPFRYLNWAPGSPSLESEKICGLLQSGNGKWKNQQCDQKLGYICKKGNSSLDSFIIPSGDFRPIKCPSGWIPYSGHCYMLYRAPKIWKDALSSCRKEGGDLASIHNIEEYSFTVSQLGYKPTDELWIGLNDLKIHMYFDWSDGTPVTYTKWLHEEPSHANNRKEDCVSMKGEDGYWADDVCEKKLGYICKMKPLAEESEKVEVIIPGCQKGWIRHGFYCYSVGQTSVTFSEAKKICEGNKGYLVTVKDRYEQAFLTVLIGFSPVKYFWIALSNVEEQGTFKWADGEAVLFTHWNSGMPGREPGCVAMRTGTTAGLWDVVNCEEKSMFLCKQLVEGATPPPVPTTTPAPSCPDGWHSRAHSSSCFKHFQNGRKHMKTWFDARDFCRAIGGDLASIHSEEDQNVINSLERAHYYQSYWMGMSAFDPDGGFTWSDGSPVNYENWADGEPNNYDGNEKCAMFNEYTDMQWNDMFCEKFGDWTCQIKKGATLKPEPSTTFEYTYKVSEDGWIIYEDKAYYFSHETLPMEKAREYCKKNSGDLVVIEGESKRKFLWKYSFYNDFRESLYIGLSVSLDKKFSWIDGTPLNYVAWSPNEPNFANNDENCVVMYTQTGMWNDLNCGAPNSFICERLNNITHSTVAPTSPAPLGGCPQNWFLFNNKCFKIFGSSENESLTWHAARTACINLEGNLASIPSEXAAASFLITHLKDASTDTWIGLNDVNSEYTFLWTDGSGVSYTSWAKGFPNYRSEADCVMMIKDPIEEAGKWKDEDCQMNKSYICQRNADPKLYNFQTTVPVSGFTLYGNSSYSLISSRMNWEEAQKNCKAEYSELASILDPYSQSFLWLQVLKIGQPVWIGLNSNVTSGFYKWTDNWRIKYSKWASEEPKQNLACVYLDVDGTWKTGSCDENYCSVCKRSDAPTELPQLPGKCPTSDIRRSWIPFHGHCYYIESSEMESWPQSSMECVRLGESSLVSIENMAEMNFLLHHLDAFASDSRQFWIGMYKNVDGEWTWADNSAVEFVNWKEREPTDGYHEYCVDMDASDGGWRSYYCSADKRFICKIPKITEVEPSEKLSDNKGRTKDEAVPAHHLSGMAVILVLLILVGAGLAAYIFFKKKRHNQLPIDVTFDNTLYCNRDAVPAASDSKYLVANIEQNEQAML</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMJ8</accession>
    <name>A0A452IMJ8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="29">Protein kinase C and casein kinase substrate in neurons 2</fullName>
        </recommendedName>
    </protein>
    <organism evidence="27 28">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="28" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="27" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="27" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cell junction</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="7">Cell projection</location>
            <location evidence="7">Ruffle membrane</location>
            <topology evidence="7">Peripheral membrane protein</topology>
            <orientation evidence="7">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasmic vesicle membrane</location>
            <topology evidence="1">Peripheral membrane protein</topology>
            <orientation evidence="1">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="5">Early endosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Membrane</location>
            <location evidence="4">Caveola</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="6">Recycling endosome membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="8">Belongs to the PACSIN family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMJ8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033048.1">
        <property type="protein sequence ID" value="ENSGAGP00000029093.1"/>
        <property type="gene ID" value="ENSGAGG00000021048.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070161">
        <property type="term" value="C:anchoring junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005901">
        <property type="term" value="C:caveola"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005769">
        <property type="term" value="C:early endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055038">
        <property type="term" value="C:recycling endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032587">
        <property type="term" value="C:ruffle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005543">
        <property type="term" value="F:phospholipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030036">
        <property type="term" value="P:actin cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070836">
        <property type="term" value="P:caveola assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006897">
        <property type="term" value="P:endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097320">
        <property type="term" value="P:plasma membrane tubulation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030100">
        <property type="term" value="P:regulation of endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07679">
        <property type="entry name" value="F-BAR_PACSIN2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd11998">
        <property type="entry name" value="SH3_PACSIN1-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.40:FF:000014">
        <property type="entry name" value="Kinase C and casein kinase substrate in neurons protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1270.60:FF:000205">
        <property type="entry name" value="Protein kinase C and casein kinase substrate in neurons protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.60">
        <property type="entry name" value="Arfaptin homology (AH) domain/BAR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027267">
        <property type="entry name" value="AH/BAR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031160">
        <property type="entry name" value="F_BAR_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001060">
        <property type="entry name" value="FCH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035743">
        <property type="entry name" value="PACSIN1/PACSIN2_SH3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037453">
        <property type="entry name" value="PACSIN2_F-BAR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23065">
        <property type="entry name" value="PROLINE-SERINE-THREONINE PHOSPHATASE INTERACTING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23065:SF14">
        <property type="entry name" value="PROTEIN KINASE C AND CASEIN KINASE SUBSTRATE IN NEURONS PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00611">
        <property type="entry name" value="FCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14604">
        <property type="entry name" value="SH3_9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00452">
        <property type="entry name" value="SH3DOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00055">
        <property type="entry name" value="FCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00326">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103657">
        <property type="entry name" value="BAR/IMD domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51741">
        <property type="entry name" value="F_BAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="15" id="KW-0965">Cell junction</keyword>
    <keyword evidence="10" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="20" id="KW-0966">Cell projection</keyword>
    <keyword evidence="16 23" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="11" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="21" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="19" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="13" id="KW-0254">Endocytosis</keyword>
    <keyword evidence="14" id="KW-0967">Endosome</keyword>
    <keyword evidence="17" id="KW-0446">Lipid-binding</keyword>
    <keyword evidence="18" id="KW-0472">Membrane</keyword>
    <keyword evidence="12" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="28" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9 22" id="KW-0728">SH3 domain</keyword>
    <feature type="domain" description="F-BAR" evidence="26">
        <location>
            <begin position="71"/>
            <end position="340"/>
        </location>
    </feature>
    <feature type="domain" description="SH3" evidence="25">
        <location>
            <begin position="444"/>
            <end position="504"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="24">
        <location>
            <begin position="373"/>
            <end position="440"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="24">
        <location>
            <begin position="385"/>
            <end position="414"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="24">
        <location>
            <begin position="422"/>
            <end position="434"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004282"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004345"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004412"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00004565"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00004599"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00010461"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022583"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023121"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01077"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="26">
        <source>
            <dbReference type="PROSITE" id="PS51741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029093.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="28">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="29">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="504" mass="58127" checksum="6503C31DC56F618F" modified="2019-05-08" version="1">MILIDSRVSQWCSLLLYREAGAHVAFLVQSEDLYAAANGSPRTGASWVGGRWFGWSRAKQGPSRPTPTRRRGAGEQLEVGNYKRTVKRIDDGHRLCSDLMNCIHERARIEKTYAQQLTEWAKRWKQLVEKGPQYGTVEKAWHAVMSEAEKVSELHLDIKSALMNDDFEKIKNWQKEAFHKQMMGGFKETKEAEDGYRKAQKPWAKKLKEVEAAKKAYHAACKEEKLAISREMNSKADPALNPEQLKKLQDKVEKSKQDVLKTKEKYEKSLKDLDHVTPQYMENMEQVFEQCQQFEEKRLRFFREVLLEVQKHLDLSNDVSYKNIYRELEQNIKTADAIEDLRWFRANHGPGMSMNWPQFEEWSADFNRTLSRREKKKASDGVTLTGVNQTGDQALQHTKHSSVSSSEKNQSYPTEWSDEESSNPFSSTDANGDTNPFDEDASPAMEVRVRALYDYEGQEHDELSFKAGDELTKTEDEDEQGWCKGRLDNGQVGLYPANYVEAIQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMK2</accession>
    <name>A0A452IMK2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">Mannose receptor C-type 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="11">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000033135.1">
        <property type="protein sequence ID" value="ENSGAGP00000029173.1"/>
        <property type="gene ID" value="ENSGAGG00000020766.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006897">
        <property type="term" value="P:endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd23407">
        <property type="entry name" value="beta-trefoil_Ricin_MRC1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00037">
        <property type="entry name" value="CLECT"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="CDD" id="cd00062">
        <property type="entry name" value="FN2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.80.10.50:FF:000032">
        <property type="entry name" value="macrophage mannose receptor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000016">
        <property type="entry name" value="macrophage mannose receptor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000023">
        <property type="entry name" value="Macrophage mannose receptor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000031">
        <property type="entry name" value="macrophage mannose receptor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000025">
        <property type="entry name" value="Mannose receptor C-type 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000030">
        <property type="entry name" value="Mannose receptor C-type 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.100.10:FF:000027">
        <property type="entry name" value="Mannose receptor, C type 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.80.10.50">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.10.10">
        <property type="entry name" value="Fibronectin, type II, collagen-binding"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.100.10">
        <property type="entry name" value="Mannose-Binding Protein A, subunit A"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001304">
        <property type="entry name" value="C-type_lectin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016186">
        <property type="entry name" value="C-type_lectin-like/link_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050111">
        <property type="entry name" value="C-type_lectin/snaclec_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018378">
        <property type="entry name" value="C-type_lectin_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016187">
        <property type="entry name" value="CTDL_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000562">
        <property type="entry name" value="FN_type2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036943">
        <property type="entry name" value="FN_type2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013806">
        <property type="entry name" value="Kringle-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035992">
        <property type="entry name" value="Ricin_B-like_lectins"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000772">
        <property type="entry name" value="Ricin_B_lectin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22803">
        <property type="entry name" value="MANNOSE, PHOSPHOLIPASE, LECTIN RECEPTOR RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24562">
        <property type="entry name" value="CysR_MRC2_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00040">
        <property type="entry name" value="fn2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00059">
        <property type="entry name" value="Lectin_C"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00013">
        <property type="entry name" value="FNTYPEII"/>
    </dbReference>
    <dbReference type="SMART" id="SM00034">
        <property type="entry name" value="CLECT"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SMART" id="SM00059">
        <property type="entry name" value="FN2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00458">
        <property type="entry name" value="RICIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56436">
        <property type="entry name" value="C-type lectin-like"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57440">
        <property type="entry name" value="Kringle-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50370">
        <property type="entry name" value="Ricin B-like lectins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00615">
        <property type="entry name" value="C_TYPE_LECTIN_1"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50041">
        <property type="entry name" value="C_TYPE_LECTIN_2"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51092">
        <property type="entry name" value="FN2_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50231">
        <property type="entry name" value="RICIN_B_LECTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="2" id="KW-0254">Endocytosis</keyword>
    <keyword evidence="10" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7 12" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-0675">Receptor</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="3 12" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 12" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="1304"/>
            <end position="1327"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-II" evidence="14">
        <location>
            <begin position="131"/>
            <end position="182"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="13">
        <location>
            <begin position="196"/>
            <end position="311"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="13">
        <location>
            <begin position="337"/>
            <end position="454"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="13">
        <location>
            <begin position="479"/>
            <end position="594"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="13">
        <location>
            <begin position="623"/>
            <end position="719"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="13">
        <location>
            <begin position="740"/>
            <end position="856"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="13">
        <location>
            <begin position="884"/>
            <end position="1012"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="13">
        <location>
            <begin position="1031"/>
            <end position="1142"/>
        </location>
    </feature>
    <feature type="domain" description="C-type lectin" evidence="13">
        <location>
            <begin position="1171"/>
            <end position="1288"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004167"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022583"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50041"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS51092"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029173.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="17">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1373" mass="156081" checksum="7FF2617CA63D1774" modified="2019-05-08" version="1">KLCVKAQSSNSVIIVTCNQDDDSQKFRWVSDHQILSMALKLCLGVPSMKDQAAITLYPCNKTSELQQWECRNKTLLAIQGEDLFFNSAHGEERKVMLSKGSGSWNRWKIHGTTENLCSQGYEDMFTLLGNAFGAPCVFPFKFNSTWYAECTKDRKAGNKQGFPWCATSTDFDEDSLFGYCPLKDNIDKDFWITNPFTGTHYQINSNSALTWHQARKSCQQQNAELLSITELHEQTYLTGLFGRANSELWIGLNRLNSNGGWQWIGGSPFRYLNWAPGSPSLESEKICGLLQSGNGKWKNQQCDQKLGYICKKGNSSLDSFIIPSGRPIKCPSGWIPYSGHCYMLYRAPKIWKDALSSCRKEGGDLASIHNIEEYSFTVSQLGYKPTDELWIGLNDLKIHMYFDWSDGTPVTYTKWLHEEPSHANNRKEDCVSMKGEDGYWADDVCEKKLGYICKMKPLAEESEKVEVIIPGCQKGWIRHGFYCYSVGQTSVTFSEAKKICEGNKGYLVTVKDRYEQAFLTVLIGFSPVKYFWIALSNVEEQGTFKWADGEAVLFTHWNSGMPGREPGCVAMRTGTTAGLWDVVNCEEKSMFLCKQLVEGATPPPVPTTTPAPSCPDGWHSRAHSSSCFKHFQNGRKHMKTWFDARDFCRAIGGDLASIHSEEDQNVINIVFLPKVNYENWADGEPNNYDGNEKCAMFNEYTDMQWNDMFCEKFGDWTCQIKKGATLKPEPSTTFDGWIIYEDKAYYFSHETLPMEKAREYCKKNSGDLVVIEGESKRKFLWKYSFYNDFRESLYIGLSVSLDKKFSWIDGTPLNYVAWSPNEPNFANNDENCVVMYTQTGMWNDLNCGAPNSFICERLNNITHSTVAPTSPAPLGGCPQNWFLFNNKCFKIFGSSENESLTWHAARTACINLEGNLASIPSEEVQAFLITHLKDASTDTWIGLNDVNSEYTFLWTDGSGVSYTSWAKGFPNYRPYSANLTQADCVMMIKDPIEEAGKWKDEDCQMNKSYICQRNAGEFFLTKLPISGFTLYGNSSYSLISSRMNWEEAQKNCKAEYSELASILDPYSQSFLWLQVLKIGQPVWIGLNSNVTSGFYKWTDNWRIKYSKWASEEPKQNLACVYLDVDGTWKTGSCDENYCSVCKRSDVLAPTELPQLPGKCPTSDIRRSWIPFHGHCYYIESSEMESWPQSSMECVRLGASLVSIENMAEMNFLLHHLDAFASDSRQFWIGMYKNVDGEWTWADNSAVEFVNWKEREPTDGYHEYCVDMDASDGGWRSYYCSADKRFICKIPKILKKDEAVPAHHLSGMAVILVLLILVGAGLAAYIFFKKKRHNQLPIDVTFDNTLYCNRDAVPAASDSKYLVANIEQNEQAML</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMK3</accession>
    <name>A0A452IMK3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">ER lumen protein-retaining receptor</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasmic vesicle</location>
            <location evidence="1">COPI-coated vesicle membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2 14">Endoplasmic reticulum membrane</location>
            <topology evidence="2 14">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Golgi apparatus membrane</location>
            <topology evidence="3">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4 14">Belongs to the ERD2 family.</text>
    </comment>
    <comment type="caution">
        <text evidence="14">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMK3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033291.1">
        <property type="protein sequence ID" value="ENSGAGP00000029312.1"/>
        <property type="gene ID" value="ENSGAGG00000021206.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030663">
        <property type="term" value="C:COPI-coated vesicle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000139">
        <property type="term" value="C:Golgi membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046923">
        <property type="term" value="F:ER retention sequence binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006621">
        <property type="term" value="P:protein retention in ER lumen"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016192">
        <property type="term" value="P:vesicle-mediated transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000133">
        <property type="entry name" value="ER_ret_rcpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10585">
        <property type="entry name" value="ER LUMEN PROTEIN RETAINING RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00810">
        <property type="entry name" value="ER_lumen_recept"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00660">
        <property type="entry name" value="ERLUMENR"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00951">
        <property type="entry name" value="ER_LUMEN_RECEPTOR_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00952">
        <property type="entry name" value="ER_LUMEN_RECEPTOR_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="13" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="7 14" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="8" id="KW-0931">ER-Golgi transport</keyword>
    <keyword evidence="11 14" id="KW-0472">Membrane</keyword>
    <keyword evidence="9 14" id="KW-0653">Protein transport</keyword>
    <keyword evidence="12 14" id="KW-0675">Receptor</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6 14" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="10 14" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="5 14" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="62"/>
            <end position="80"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="120"/>
            <end position="138"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="150"/>
            <end position="168"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="14">
        <location>
            <begin position="180"/>
            <end position="199"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004129"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004477"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004653"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00010120"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022892"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="RuleBase" id="RU000634"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029312.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="203" mass="23881" checksum="6F09E335ECF23483" modified="2019-05-08" version="1">MNIFRILGDVSHLLAMIILLVKIWRSKSCAGISGKSQILFALVFTTRYLDLFTTFISLYNTVMKVIFLVCAYITVYMIYVKFQKTFDREHDSFRLEFLLVPVTGLSFLENHSFTALEILWTFSIYLESVAILPQLFMISKTGEAETITTHYLFFLGLYRALYLANWIWRYYTENFYDQIAVVSGVVQTIFYCDFFYLYVTKGR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMK4</accession>
    <name>A0A452IMK4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4 15">Very-long-chain (3R)-3-hydroxyacyl-CoA dehydratase</fullName>
            <ecNumber evidence="4 15">4.2.1.134</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">Catalyzes the third of the four reactions of the long-chain fatty acids elongation cycle. This endoplasmic reticulum-bound enzymatic process, allows the addition of two carbons to the chain of long- and very long-chain fatty acids/VLCFAs per cycle. This enzyme catalyzes the dehydration of the 3-hydroxyacyl-CoA intermediate into trans-2,3-enoyl-CoA, within each cycle of fatty acid elongation. Thereby, it participates to the production of VLCFAs of different chain lengths that are involved in multiple biological processes as precursors of membrane lipids and lipid mediators.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>(3R)-hydroxyhexadecanoyl-CoA = (2E)-hexadecenoyl-CoA + H2O</text>
            <dbReference type="Rhea" id="RHEA:39159"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:61526"/>
            <dbReference type="ChEBI" id="CHEBI:74278"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="13">
            <dbReference type="Rhea" id="RHEA:39160"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="14">
            <text>a very-long-chain (3R)-3-hydroxyacyl-CoA = a very-long-chain (2E)-enoyl-CoA + H2O</text>
            <dbReference type="Rhea" id="RHEA:45812"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:83728"/>
            <dbReference type="ChEBI" id="CHEBI:85440"/>
            <dbReference type="EC" id="4.2.1.134"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="14">
            <dbReference type="Rhea" id="RHEA:45813"/>
        </physiologicalReaction>
    </comment>
    <comment type="pathway">
        <text evidence="2 15">Lipid metabolism; fatty acid biosynthesis.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="15">Endoplasmic reticulum membrane</location>
            <topology evidence="15">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 15">Belongs to the very long-chain fatty acids dehydratase HACD family.</text>
    </comment>
    <comment type="caution">
        <text evidence="15">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="EC" id="4.2.1.134" evidence="4 15"/>
    <dbReference type="AlphaFoldDB" id="A0A452IMK4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033016.1">
        <property type="protein sequence ID" value="ENSGAGP00000029063.1"/>
        <property type="gene ID" value="ENSGAGG00000021030.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00094"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0102158">
        <property type="term" value="F:very-long-chain (3R)-3-hydroxyacyl-CoA dehydratase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030497">
        <property type="term" value="P:fatty acid elongation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030148">
        <property type="term" value="P:sphingolipid biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042761">
        <property type="term" value="P:very long-chain fatty acid biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007482">
        <property type="entry name" value="Tyr_Pase-like_PTPLA"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11035">
        <property type="entry name" value="VERY-LONG-CHAIN (3R)-3-HYDROXYACYL-COA DEHYDRATASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11035:SF16">
        <property type="entry name" value="VERY-LONG-CHAIN (3R)-3-HYDROXYACYL-COA DEHYDRATASE 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04387">
        <property type="entry name" value="PTPLA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="15" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="11 15" id="KW-0275">Fatty acid biosynthesis</keyword>
    <keyword evidence="7 15" id="KW-0276">Fatty acid metabolism</keyword>
    <keyword evidence="5 15" id="KW-0444">Lipid biosynthesis</keyword>
    <keyword evidence="9 15" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="12 15" id="KW-0456">Lyase</keyword>
    <keyword evidence="10 15" id="KW-0472">Membrane</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6 15" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 15" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="16"/>
            <end position="38"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="187"/>
            <end position="207"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005194"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007811"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00013122"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022516"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022832"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023160"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023688"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023727"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="RuleBase" id="RU363109"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029063.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="229" mass="26700" checksum="408820F3A3FB3752" modified="2019-05-08" version="1">GNPLPVPLCTMNTYLFIYYLIQFCGHSWIFTNMIIRFLSFGKDSLADTFYSIGLVMRLCQSLSILELLHILVGIEENRFLPRFLQITERIIILFVVITSQEEVQGKYVVCVLFFLWNLLDVVRYTYCMLSVSGTYYGALTWLNYTLWIPVYPLSILAKAFAIYESLPYFESFGTYSIKLPFPVALSIYFPYILKLYLAMLFVGMYFITTYLCSERKAHLGASYIKEKRS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMK8</accession>
    <name>A0A452IMK8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">THAP-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IMK8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033041.1">
        <property type="protein sequence ID" value="ENSGAGP00000029086.1"/>
        <property type="gene ID" value="ENSGAGG00000021046.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052958">
        <property type="entry name" value="IFN-induced_PKR_regulator"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006612">
        <property type="entry name" value="THAP_Znf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46289:SF16">
        <property type="entry name" value="52 KDA REPRESSOR OF THE INHIBITOR OF THE PROTEIN KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46289">
        <property type="entry name" value="52 KDA REPRESSOR OF THE INHIBITOR OF THE PROTEIN KINASE-LIKE PROTEIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05485">
        <property type="entry name" value="THAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00692">
        <property type="entry name" value="DM3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00980">
        <property type="entry name" value="THAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57716">
        <property type="entry name" value="Glucocorticoid receptor-like (DNA-binding domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50950">
        <property type="entry name" value="ZF_THAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4 5" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="1" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0862">Zinc</keyword>
    <keyword evidence="2 5" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="THAP-type" evidence="6">
        <location>
            <begin position="1"/>
            <end position="86"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00309"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50950"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029086.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="118" mass="13837" checksum="038A0CB81993D003" modified="2019-05-08" version="1">MPNFCAAPNCTRKSTQSDLAFFRFPRDPVRCQRWVENCRRADLEDKTPDQLNKHYRLCAKHFETSMICRSSPYRTVLRDNAVPTIFDLTSHLNNPHSRHRKRIKELALHLPTSGETLL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IML0</accession>
    <name>A0A452IML0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Unc-13 homolog A</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IML0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033147.1">
        <property type="protein sequence ID" value="ENSGAGP00000029185.1"/>
        <property type="gene ID" value="ENSGAGG00000020811.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031594">
        <property type="term" value="C:neuromuscular junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098831">
        <property type="term" value="C:presynaptic active zone cytoplasmic component"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042734">
        <property type="term" value="C:presynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030672">
        <property type="term" value="C:synaptic vesicle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="C:terminal bouton"/>
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        <property type="term" value="F:diacylglycerol binding"/>
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        <property type="term" value="F:syntaxin-1 binding"/>
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        <property type="term" value="F:zinc ion binding"/>
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        <property type="project" value="UniProtKB-KW"/>
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    <dbReference type="GO" id="GO:0061789">
        <property type="term" value="P:dense core granule priming"/>
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        <property type="project" value="TreeGrafter"/>
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        <property type="term" value="P:presynaptic dense core vesicle exocytosis"/>
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        <property type="project" value="TreeGrafter"/>
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    <dbReference type="GO" id="GO:0035249">
        <property type="term" value="P:synaptic transmission, glutamatergic"/>
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        <property type="project" value="TreeGrafter"/>
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    <dbReference type="GO" id="GO:0016081">
        <property type="term" value="P:synaptic vesicle docking"/>
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    <dbReference type="GO" id="GO:0016082">
        <property type="term" value="P:synaptic vesicle priming"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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    <dbReference type="CDD" id="cd20859">
        <property type="entry name" value="C1_Munc13-2-like"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="C2A_Munc13"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="C2B_Munc13"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Protein unc-13 homolog B"/>
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        <property type="entry name" value="Protein unc-13 homolog B"/>
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        <property type="entry name" value="Protein unc-13 homolog B"/>
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        <property type="entry name" value="Protein unc-13 homolog B"/>
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        <property type="entry name" value="C2_domain_sf"/>
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        <property type="entry name" value="MUN_dom"/>
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        <property type="entry name" value="Munc13_1"/>
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        <property type="entry name" value="PKC_DAG/PE"/>
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    <dbReference type="InterPro" id="IPR027080">
        <property type="entry name" value="Unc-13"/>
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        <property type="entry name" value="Unc-13_C2B"/>
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    <dbReference type="PANTHER" id="PTHR10480">
        <property type="entry name" value="PROTEIN UNC-13 HOMOLOG"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="PROTEIN UNC-13 HOMOLOG A"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="C1_1"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06292">
        <property type="entry name" value="MUN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00360">
        <property type="entry name" value="C2DOMAIN"/>
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        <property type="entry name" value="DUF1041"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49562">
        <property type="entry name" value="C2 domain (Calcium/lipid-binding domain, CaLB)"/>
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    </dbReference>
    <dbReference type="SUPFAM" id="SSF57889">
        <property type="entry name" value="Cysteine-rich domain"/>
        <property type="match status" value="1"/>
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        <property type="match status" value="2"/>
    </dbReference>
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    </dbReference>
    <dbReference type="PROSITE" id="PS00479">
        <property type="entry name" value="ZF_DAG_PE_1"/>
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    </dbReference>
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        <property type="entry name" value="ZF_DAG_PE_2"/>
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    </dbReference>
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    <keyword evidence="1" id="KW-0268">Exocytosis</keyword>
    <keyword evidence="2" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <keyword evidence="4" id="KW-0863">Zinc-finger</keyword>
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        <location>
            <begin position="1"/>
            <end position="97"/>
        </location>
    </feature>
    <feature type="domain" description="Phorbol-ester/DAG-type" evidence="8">
        <location>
            <begin position="470"/>
            <end position="520"/>
        </location>
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        <location>
            <begin position="576"/>
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        </location>
    </feature>
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        <location>
            <begin position="1005"/>
            <end position="1148"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
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            <begin position="176"/>
            <end position="318"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="346"/>
            <end position="378"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="195"/>
            <end position="204"/>
        </location>
    </feature>
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        <location>
            <begin position="205"/>
            <end position="224"/>
        </location>
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    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="231"/>
            <end position="246"/>
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        <location>
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            <end position="266"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
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            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
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            <dbReference type="PROSITE" id="PS50004"/>
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        <source>
            <dbReference type="PROSITE" id="PS51258"/>
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    </evidence>
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        <source>
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    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="12">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1277" mass="144624" checksum="2A49A2795F7E9978" modified="2019-05-08" version="1">TFYSLFAVKKAKFDGAQEKFNTYVTLKVQNVKSTTIAVRGNLPCWEQDFMFEINRLDLGLTVEVWNKGLIWDTMVGTVWIPLQTIRQSNEEGPGEWLTLDSQVIMTDNEISGTKDPTFHRILLDTHFELPLDIPEEEARYWAKKLEQLNAMKDQDDVNTGYLIPLRHVVASGPLPCETLTGDNRPFADDDPDSTMDDRDSDYRSETSNSIPPPYYTTSQPNASVHQYPMRHQQQSSRDSYMDSVQNYKMDYSDQRPISPTGSSRYASSGELSQGSSQLSDDFDPDNESLRGSELDERDGDSYHSCHSSVSYHDDSPRWEEDFYPEEEDEDYSENEQDFPKVEAEVLRDREADKAKPPDISAGMQRGHLPGAGQSTGPSVHGCLGRSPCPVPSDSCLSSRPGGGLIIIDSMPDIRKRKPIPLVSDLSLVQSRKAGITSALASSTLNNEELKNHVYKKTLQALIYPISSTTPHNFEVWTATTPTYCYECEGLLWGIARQGMRCTECGVKCHEKCQDLLNADCLQRAAEKSSKHGAEDRTQNIIMVLKDRMKIRERNKPEIFELIQEIFAVSKATHTQQMKAVKQSVLDGTSKWSAKISITVVCAQGLQAKDKTGSSDPYVTVQVGKTKKRTKTIYGNLNPVWEENFHFECHNSSDRIKVRVWDEDDDIKSRVKQRFKRESDDFLGQTIIEVRTLSGEMDVWYNLDKRTDKSAVSGAIRLHISVEIKGEEKVAPYHVQYTCLHENLFHFVTDIQNNGVVKIPEAKGDDAWKVYFDETAQEIVDEFAMRYGVESIYQAMTHFACLSSKYMCPGVPAVMSTLLANINAYYAHTTASTNVSASDRFAASNFGKERFVKLLDQLHNSLRIDLSMYRNNFPASSPERLQDLKSTVDLLTSITFFRMKVQELQSPPRASQVVKDCVKACLNSTYEYIFNNCHELYSREYQTDPVPPEEQGPSIKNLDFWSKLITLIVSIIEEDKNSYTPCLNQFPQELNVGKISAEVMWNLFAQDMKYAMEEHDKHRLCKSADYMNLHFKVKWLYNEYVTELPAFKSRVPEYPAWFEPFVIQWLDENEEVSRDFLHGALERDKKDGFQQTSEHALFSCSVVDVFSQLNQSFEIIKKLECPDPQIVGHYMRRFAKTISNVLLQYAEIISKDFALYCSKEKEKPCILMNNIQQLRVQLEKMFEAMGGKELDAEASDILKELQVKLNNVLDELSRVFATSFQPHIEECVKQMGDILSQVKGTGNVPASACSSVAQDADNQVSAGGLGGPVGRGTGGTAG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IML2</accession>
    <name>A0A452IML2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">CWF21 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IML2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033064.1">
        <property type="protein sequence ID" value="ENSGAGP00000029108.1"/>
        <property type="gene ID" value="ENSGAGG00000021058.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003729">
        <property type="term" value="F:mRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd21376">
        <property type="entry name" value="cwf21_SRRM3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.420">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013170">
        <property type="entry name" value="mRNA_splic_Cwf21_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047489">
        <property type="entry name" value="SRRM3_cwf21"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029360">
        <property type="entry name" value="SRRM_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052109">
        <property type="entry name" value="SRRM_Domain-Containing"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR34755">
        <property type="entry name" value="SERINE/ARGININE REPETITIVE MATRIX PROTEIN 3-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR34755:SF2">
        <property type="entry name" value="SERINE_ARGININE REPETITIVE MATRIX PROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08312">
        <property type="entry name" value="cwf21"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15230">
        <property type="entry name" value="SRRM_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01115">
        <property type="entry name" value="cwf21"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="CWF21" evidence="2">
        <location>
            <begin position="52"/>
            <end position="97"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1"/>
            <end position="36"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="151"/>
            <end position="584"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="1">
        <location>
            <begin position="164"/>
            <end position="182"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="1">
        <location>
            <begin position="197"/>
            <end position="209"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="1">
        <location>
            <begin position="217"/>
            <end position="241"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="1">
        <location>
            <begin position="255"/>
            <end position="273"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="1">
        <location>
            <begin position="289"/>
            <end position="306"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="373"/>
            <end position="384"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="1">
        <location>
            <begin position="385"/>
            <end position="399"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="401"/>
            <end position="425"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="1">
        <location>
            <begin position="442"/>
            <end position="456"/>
        </location>
    </feature>
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        <location>
            <begin position="479"/>
            <end position="492"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="1">
        <location>
            <begin position="493"/>
            <end position="504"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="1">
        <location>
            <begin position="505"/>
            <end position="514"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="516"/>
            <end position="535"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="1">
        <location>
            <begin position="536"/>
            <end position="551"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="563"/>
            <end position="573"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="SMART" id="SM01115"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029108.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="584" mass="66510" checksum="BD71800971757392" modified="2019-05-08" version="1">MALYNNGAKMPSPQEAANGFPQPGASSTWHKPEEEVRLVEPSLVKKAHREILDHERKRRVELKCMELQEMMEEQGYSEEEIRQKVGTFRQMLMEKEGVLTREDQHGRQIVIENHHMEQYAVEYPAYGEGCLLPCDCPADCYRDHCSHREHRLKRRMSSSASPPPKKKKKKKSGHRRSRKKRKPGSERSCDSSSPLRKEKKKKNGKKHRRDRSESGSRKKRRRRSRSPKNKRKEKTKERKRSPSNSPARRCHRHSSCSSHSASLSSDYSHSKSSGRLTPKHREDGHKPSSLRSSRSPSSSRTWHSRSATPHQNGHKGSAQNGRHSHGTPGEKQQDRLDSASPSPRGHVKTELPSPRSKGGRRGPRSLRSVSPEKGGRCSGGDKYRRCSRSASLHRHKGRSKGQASAQKESPHSLSHTDYSSDSEGSACSHPYPAARGVEKNHGAHLKKVKDRHHRGRPNSCSQSPVKHSRHNSERRKSLSRSSSWSSSSSPSKSRSRSREKRAARSRSQSPSQKKNSSRERDNEPRTRHSDTDPARARRRSRSYSPIRKRRRDSPSFMEPRRITSPSQGSSRRASPQAWLPWCPG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMN1</accession>
    <name>A0A452IMN1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Chloride channel CLIC-like protein 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the chloride channel MCLC family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMN1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033322.1">
        <property type="protein sequence ID" value="ENSGAGP00000029342.1"/>
        <property type="gene ID" value="ENSGAGG00000021220.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000033324.1">
        <property type="protein sequence ID" value="ENSGAGP00000029344.1"/>
        <property type="gene ID" value="ENSGAGG00000021220.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005783">
        <property type="term" value="C:endoplasmic reticulum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005254">
        <property type="term" value="F:chloride channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009231">
        <property type="entry name" value="Chloride_chnl_CLIC-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR34093">
        <property type="entry name" value="CHLORIDE CHANNEL CLIC-LIKE PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR34093:SF1">
        <property type="entry name" value="CHLORIDE CHANNEL CLIC-LIKE PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05934">
        <property type="entry name" value="MCLC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="6 9" id="KW-0472">Membrane</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0732">Signal</keyword>
    <keyword evidence="4 9" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5 9" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="10">
        <location>
            <begin position="1"/>
            <end position="16"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5044604582" description="Chloride channel CLIC-like protein 1" evidence="10">
        <location>
            <begin position="17"/>
            <end position="559"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="188"/>
            <end position="209"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="221"/>
            <end position="238"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="9">
        <location>
            <begin position="337"/>
            <end position="358"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="393"/>
            <end position="437"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="493"/>
            <end position="559"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="68"/>
            <end position="95"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="412"/>
            <end position="421"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="515"/>
            <end position="526"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="540"/>
            <end position="551"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005944"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00015571"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029342.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="559" mass="63597" checksum="473CEA2793B4E69E" modified="2019-05-08" version="1">MLVPLVLCATLMIGSGEIQDDEWIDPTDMLNYDAASGTMRRPAKVNYDDSENREKPDAVVSLPSTAAVSECHRKLESLIQKIEEYEKKEKTKLSESHSIYVFRRYLNKILIEAGRLGLPDENIGDVHYDAEIILTSQALLEINRFLNEEDWKSGALDDAVNNILINFKHHNFEVWKWRFEDTFGIDPYNLLMVLLCLLCITVTVATELWTHIGWYTQVKRILFISFLISCGWNWIYLYKIAFAQHQAEVARMGNYDTVCDKKIDWSESLFEWLRSSWTFQDDPCQKYYETILVNPIWLVPPTKVLAVTFTNFVTEPLKHIGKGIGEFIKALMKEIPMMLQIPVLIILAVAVLGFCYGAGRSVATLKHLTGPDRKPSPSLPSSDRQHWEQIHYSAGDSDSNYRRNVGAISGGPHDRGDDHMRLQNGNRSSEVLLPGGTPNAQREEHYKCGLHPVSQDQTIFRVKTNVDQNCDREIALEEQKLCTLGVEQESEKNCEPQASCSLKKEKTNKQNSVEKPGDLEKTEKGSLSRLMEGTEETKEPLNSTEQNQQPTIEDLPGTP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMN5</accession>
    <name>A0A452IMN5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Endophilin-B2</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="8">SH3 domain-containing GRB2-like protein B2</fullName>
        </alternativeName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="6">Homodimer, and heterodimer with SH3GLB1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the endophilin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMN5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033100.1">
        <property type="protein sequence ID" value="ENSGAGP00000029142.1"/>
        <property type="gene ID" value="ENSGAGG00000021083.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061024">
        <property type="term" value="P:membrane organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd07617">
        <property type="entry name" value="BAR_Endophilin_B2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd11944">
        <property type="entry name" value="SH3_Endophilin_B2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.40:FF:000028">
        <property type="entry name" value="endophilin-B2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.60">
        <property type="entry name" value="Arfaptin homology (AH) domain/BAR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027267">
        <property type="entry name" value="AH/BAR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004148">
        <property type="entry name" value="BAR_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035640">
        <property type="entry name" value="Endophilin_B2_SH3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050384">
        <property type="entry name" value="Endophilin_SH3RF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14167:SF68">
        <property type="entry name" value="DREBRIN-LIKE PROTEIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14167">
        <property type="entry name" value="SH3 DOMAIN-CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03114">
        <property type="entry name" value="BAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14604">
        <property type="entry name" value="SH3_9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00721">
        <property type="entry name" value="BAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00326">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103657">
        <property type="entry name" value="BAR/IMD domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51021">
        <property type="entry name" value="BAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 10" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 9" id="KW-0728">SH3 domain</keyword>
    <feature type="domain" description="BAR" evidence="12">
        <location>
            <begin position="24"/>
            <end position="291"/>
        </location>
    </feature>
    <feature type="domain" description="SH3" evidence="11">
        <location>
            <begin position="342"/>
            <end position="402"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="10">
        <location>
            <begin position="211"/>
            <end position="238"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006697"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00038771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00040331"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00042180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS51021"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029142.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="402" mass="44687" checksum="3E1F03A6A8919FE2" modified="2019-05-08" version="1">MDFNMKKLASDAGVFFSRAVQFTEEKLGQAEKTELDAHFESLLAKADCTKNWTEKILHQTEVLLQPNPSVRVEEFLYEKLDRKVPSRVTNGELLAQYMVEAANDFGPGTPYGKTLLKVGETQKRLGGAERDFMHSASINFLNPLRNFLEGDWRTITKERRILENRRLDLDACKARLKKAKAAEAKASCEGDAVPDFQETRPRNYILSASASALWSDEVEKAEHELRVAQTEFDRQAELTRLLLEGISSTHVNHLRCLHQFVESQATYYAQGYQYMLDLQKQLGRFPGTFVGNAESTSPPPATTSPTTVAATIPVVATVGRAPNPATAVEGALNLNDVKPPASGTRKARVLYDYEAADSSELALLADEMITVYSLPGMDPDWLIGERGNQKGKVPVTYLELLS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMN6</accession>
    <name>A0A452IMN6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">C2H2-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="11">Belongs to the Sp1 C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMN6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033058.1">
        <property type="protein sequence ID" value="ENSGAGP00000029102.1"/>
        <property type="gene ID" value="ENSGAGG00000021056.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd22537">
        <property type="entry name" value="SP3_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000014">
        <property type="entry name" value="Transcription factor Sp3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000026">
        <property type="entry name" value="Transcription factor Sp3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.160.60:FF:000061">
        <property type="entry name" value="Transcription factor Sp3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013087">
        <property type="entry name" value="Znf_C2H2_type"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23235">
        <property type="entry name" value="KRUEPPEL-LIKE TRANSCRIPTION FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23235:SF3">
        <property type="entry name" value="TRANSCRIPTION FACTOR SP3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00096">
        <property type="entry name" value="zf-C2H2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00355">
        <property type="entry name" value="ZnF_C2H2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00028">
        <property type="entry name" value="ZINC_FINGER_C2H2_1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50157">
        <property type="entry name" value="ZINC_FINGER_C2H2_2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0010">Activator</keyword>
    <keyword evidence="7" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="2" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="9" id="KW-0804">Transcription</keyword>
    <keyword evidence="6" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <keyword evidence="4 12" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C2H2-type" evidence="14">
        <location>
            <begin position="598"/>
            <end position="627"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="14">
        <location>
            <begin position="628"/>
            <end position="657"/>
        </location>
    </feature>
    <feature type="domain" description="C2H2-type" evidence="14">
        <location>
            <begin position="658"/>
            <end position="685"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="1"/>
            <end position="66"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="275"/>
            <end position="316"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="13">
        <location>
            <begin position="20"/>
            <end position="44"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="13">
        <location>
            <begin position="292"/>
            <end position="316"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00038409"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029102.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="758" mass="79631" checksum="028A9D601B4A20BD" modified="2019-05-08" version="1">MAALDVDSSSHSDYLQHGNGAASASAGAAAPQDAQPSPLALLAATCSKIGPPSPEEDEAAAAAAHSAGATGDLSSVQLAGTPNRWEVLSAAPTTIKDEAGNIVQIPGAATVTSSGQYVLPIQSLQNQQIFSVAPGSDSSNGTVSNVQYQVIPQIQTADGQQVQLGFATSSDNGGINQETGQIQIIPGSNQTIIASGTPSANIQNLLSQTGQVQVQGVAIGGSSFPGQAQVVANVPLGLPGNITFVPINSVDLDSLGLNSGSQTMTAGINADGHLINTGQAMDSSDTSERTGEQVSPEITETTTDNDLFVPTSSSSQLPVTIDSTSILEQNANSLTTNSGQVHSSDLQGNYIQTSVSDETQAQNIQVSTAQPIVQHIQLQESQQPTSQAQIVQGIAQQTIHGVQASQGISQQALQNLQLQLNPGTFLIQAQTVTPSGQITWQTFQVQGVQNLQNLQIQNAPGQQITLTPVQTLTLGQVAAGGALTSTPVSLSTAQLPNLQTVTVNSIDSAGIQLHPGENADSPADIRIKEEEPDPEEWQLSGDSTLNTNDLSHLRVQVVDEEGDQTHQEGKRLRRVACTCPNCKEGGGRGSNLGKKKQHICHIPGCGKVYGKTSHLRAHLRWHSGERPFVCTWMFCGKRFTRSDELQRHRRTHTGEKKFVCPECSKRFMRSDHLAKHIKTHQNKKGIHSSSTVLASVEATPDDTLITAGGTTLILANIQQGSVSGIGTVNTSGTSNQDILTNTEIPLQLVTVSGNETME</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN00</accession>
    <name>A0A452IN00_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Serologically defined colon cancer antigen 8</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IN00"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033446.1">
        <property type="protein sequence ID" value="ENSGAGP00000029454.1"/>
        <property type="gene ID" value="ENSGAGG00000021276.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007098">
        <property type="term" value="P:centrosome cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030010">
        <property type="term" value="P:establishment of cell polarity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="P:neuron migration"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="P:tube formation"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="entry name" value="SDCCAG8"/>
    </dbReference>
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        <property type="entry name" value="SEROLOGICALLY DEFINED COLON CANCER ANTIGEN 8"/>
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    </dbReference>
    <dbReference type="PANTHER" id="PTHR34343:SF1">
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    </dbReference>
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        <property type="entry name" value="CCCAP"/>
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    </dbReference>
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        <location>
            <begin position="34"/>
            <end position="60"/>
        </location>
    </feature>
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            <end position="106"/>
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    </feature>
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    <evidence type="ECO:0000256" key="1">
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            <dbReference type="SAM" id="MobiDB-lite"/>
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    </evidence>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029454.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="5">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="707" mass="82319" checksum="EB5E2408241EFB4E" modified="2019-05-08" version="1">MREAREESDLERYQRSLRDRASKSIHQLKYALVEGDPTAGESEVAFDQSSTGNDESGETAWQELQHSHAVNQLKALLREQEGLESEVSPSRRRKMSPSRTSEAADQSLPTLCDLVPIINDQSQYIHHLEAEVKFCKEELSGMKDRIQVVVLENEKLNEELKSLTVEQTLREQTLLDASANVRNSWTVGGDDSYLCQPAKSSPSHNKEQVLATRDTAETEKWQLELEKLKRLYQERTDTLEAQVRSLRKDLSESEKNCEDLKGRLKHQESLIATSNYNRVGGLCLKCAQHEAVLSQTHSNVHMHTIERLTKERDDLMGALVSVRRDLSEMQQRESSAYEQVKQAVQMTEEANLEKMKALVQCEQLKSEMERQKDRLERELIAQLDKRAGEKEALRGEMKKEREDLASTVMVLSQNVATLEARLERIIREKNSIANQLEEAQNQLTSQEMEVNKVCGEMRFQLNQTKMKKDEAEKEHREYRTKTIRELEIKDQEIEKLGLELNENKLRLEQAQQDAARAKDECLKLTELLGKSEHQLHLTRLKKESIQHSFSNEAKTRALQAEQREQELTQKMQQMEAQHDKTVNELDSLLTSQNTFIAKLKEECCMLARKLEHVTEKNRSEVSQLSQENEYFHDRLEKMQKRNDELEEQCIQHGRMHERMKLRLHQLDKHCQTTAQQLVELLNKQNQLFRERQMLTEEVQFLRTQIEA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN01</accession>
    <name>A0A452IN01_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">AKNA domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IN01"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033310.1">
        <property type="protein sequence ID" value="ENSGAGP00000029331.1"/>
        <property type="gene ID" value="ENSGAGG00000021212.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052655">
        <property type="entry name" value="AKNA_Centrosome-Trans_reg"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022150">
        <property type="entry name" value="AKNA_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21510">
        <property type="entry name" value="AKNA DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21510:SF16">
        <property type="entry name" value="PROTEIN AKNAD1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12443">
        <property type="entry name" value="AKNA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="AKNA" evidence="3">
        <location>
            <begin position="440"/>
            <end position="527"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="320"/>
            <end position="340"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="629"/>
            <end position="656"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="1">
        <location>
            <begin position="489"/>
            <end position="551"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="645"/>
            <end position="656"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="Pfam" id="PF12443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029331.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="947" mass="107868" checksum="B4A94A94F9FE9077" modified="2019-05-08" version="1">MEYNKDIQINTSKSRQPDHLKKQVNNILLDNFDYPEDATDEEQEDLPYDGDLERTYHHNNESNNLKACVSVENISNNFLNLTCSEINRSLKEEINYETQQLSQEKVFSHRMPATRTNGIMTEMAIEAPVSGMPLETELSVGAFSSLDRKEHFTNSNISDVLLRHFSKEELSNTGQLIDCETIPETSFTESVDETVLTKTRISECTKPTLRTEQWAKHFEDYHLNRQEEKSEDDYEDKHLIDENKFVIDDRATSYGDEKNCIQENSQLIAENEDTNNFKNIRNNHSHQKGLFERTGSSHELKYGQGQVNYCLPDFSKVPPKVKIPKRNNSDKSAPTMKRTKFSPNLVGKSAIVNDVLETINYFDSVEVKNQEEEMRIPELLQQLELLTKHAEAQNRIDHLRFDPKIYTHSASPNPNLSIHSRCMGAASEESVFHPLTVPIQPLLGLSQASFSGLQSGKIISSLPSASSAGEVHCLNPNLMQNTTKGEKMSQMLKEQAEQLKAKVEEFSKCVIQEAFPLQDRCLVLKELKGYLDTLERKYLATKEEHRGLQLQNYNHKSISVGEFDPDRKVEGEIFRLGMLLEDVKEKIDDSICSPCPSSFTTSPMSPSLTQSESVCSSCSCLRESPVVSSISDPPERSATEVNFHNNERNKGENRSHAIDVIPKKTSQLSLQNDNCDPFPEIQLGLQKRESCAYVKGMEPLEKAGLLANKHSSDVMQCFLPHEADNGPGGLESHRQLILSQKRNIDQENPKGSLNFWQMKSPPKTKPYNMCNLEHKIINLNVRQEQALQYRDWNINFCTSGQRNKCSRESDKNNIASSQKKRIVANSVCTSRQPDQFVYRISDKDNLETPQTCYSGMDDTIILSPQHLASRKDYGNKSIINIRNRHTSYTDTKILNSTLDHAIQTATSLQETTERMVQVVAEDLAKVKTHRRRRLKLNILSRNTIKLR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN03</accession>
    <name>A0A452IN03_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Rab3 GTPase-activating protein catalytic subunit</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Endoplasmic reticulum</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Golgi apparatus</location>
            <location evidence="1">cis-Golgi network</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the Rab3-GAP catalytic subunit family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IN03"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033240.1">
        <property type="protein sequence ID" value="ENSGAGP00000029266.1"/>
        <property type="gene ID" value="ENSGAGG00000021013.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005783">
        <property type="term" value="C:endoplasmic reticulum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045700">
        <property type="entry name" value="Rab3GAP1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045698">
        <property type="entry name" value="Rab3GAP1_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026147">
        <property type="entry name" value="Rab3GAP1_conserved"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21422">
        <property type="entry name" value="RAB3 GTPASE-ACTIVATING PROTEIN CATALYTIC SUBUNIT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21422:SF9">
        <property type="entry name" value="RAB3 GTPASE-ACTIVATING PROTEIN CATALYTIC SUBUNIT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19533">
        <property type="entry name" value="Rab3-GAP_cat_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13890">
        <property type="entry name" value="Rab3-GTPase_cat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="9" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="6" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Rab3GAP catalytic subunit conserved" evidence="11">
        <location>
            <begin position="621"/>
            <end position="775"/>
        </location>
    </feature>
    <feature type="domain" description="Rab3GAP catalytic subunit C-terminal" evidence="12">
        <location>
            <begin position="787"/>
            <end position="990"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="535"/>
            <end position="582"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="595"/>
            <end position="630"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="541"/>
            <end position="551"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="554"/>
            <end position="576"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="605"/>
            <end position="616"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004222"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00008856"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00015817"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF13890"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF19533"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029266.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="990" mass="111338" checksum="62D2F5ADFB93D2DF" modified="2019-05-08" version="1">MAADSEPESEVFEITDFTTASEWERFISKVEEVLNDWKLIGVSSGKPLEKGVYTTGIWEEKSDEISFADFRFSITHHYLVQEASGKDGKEELVEDVLPLSMQDLLCMNNDFPPRAHCLVRWYGLQEFVVIAPAANNDAVLSESKCNLLLSSVSIALGNTGCQVPLFVQIHHKWRRMYVGECQGPGVRTDFEMVHLRKVPNQYTHLSGLLDIFKSKIGCPLTPLPPVSIAIRLTYVLQDWQQYFWPQQPPDIDALIGGEVGGLEFGKLPFGACEDPVSELHLAATWPHLTEGIIVDNDVYSDLDPIQAPQWSVRVRKADNLQCLLGDFLTEFFKLCRRKESTDEILGRSTFEEDGKEVADITHALSKLTEPAPVPIHKLSVTNMVHSAKKKIRKHRGVDDSPLNNEVLNTLMLFLFPDDADKLTDGSEARLSTSRNNPPPESEEYNLYSQFKSAPSDSLTYKLALCLCMINFYHGGVKGVAHLWQEFVVEMRRRWEKNYLIPELANGSPDLRCCLLHQKLQMLNCCIERKKARDEGKKPYTSDRSPSGSSGDTGKAVDRSGGDYQKETEKERGEVGKSWESWSDSEEEFFECLSDTEELKGNGQENGKKGGTKEGNKESANLKPEGRLHPHGKLTLLHPGEPLYIPVTQEPAPMTEDLLEEQSEVLAKLGTSAEGAHLRARMQSACLLSDMESFKAANPGCCLEDFVRWYSPRDYIEEEVVDEEGNRVLKGELSARMKIPSNMWVEAWETAKPVPARRQRRLFDDTREAEKVLHYLAVQKPADLARHLLPCVIHAAVLKVKEEETLEDIASVKKIIKQIISHSSKVLRFPNPEDKKLEEIIAQIMSAEAIIARARSLKAKFGIEKCDQEEEKEDLERFVNCLLEQPEVSVIGAGRGPAGSIIHKLFLNAQRVATMTSLDEELKRSGSSDERRLNLGAVTDFPPPAGRELILRITVPRPAPYSKPLPQRMYSVITKEDFRLAGAFSADTTFF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN06</accession>
    <name>A0A452IN06_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">SMB domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IN06"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033315.1">
        <property type="protein sequence ID" value="ENSGAGP00000029336.1"/>
        <property type="gene ID" value="ENSGAGG00000020186.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009986">
        <property type="term" value="C:cell surface"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004551">
        <property type="term" value="F:dinucleotide phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003676">
        <property type="term" value="F:nucleic acid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004528">
        <property type="term" value="F:phosphodiesterase I activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046034">
        <property type="term" value="P:ATP metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030505">
        <property type="term" value="P:inorganic diphosphate transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045599">
        <property type="term" value="P:negative regulation of fat cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009143">
        <property type="term" value="P:nucleoside triphosphate catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030500">
        <property type="term" value="P:regulation of bone mineralization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd16018">
        <property type="entry name" value="Enpp"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00091">
        <property type="entry name" value="NUC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.410.20:FF:000003">
        <property type="entry name" value="Ectonucleotide pyrophosphatase/phosphodiesterase family member 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.410.20:FF:000001">
        <property type="entry name" value="Ectonucleotide pyrophosphatase/phosphodiesterase family member 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.720.10:FF:000145">
        <property type="entry name" value="Uncharacterized protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.410.20">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.720.10">
        <property type="entry name" value="Alkaline Phosphatase, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.570.10">
        <property type="entry name" value="Extracellular Endonuclease, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017850">
        <property type="entry name" value="Alkaline_phosphatase_core_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044929">
        <property type="entry name" value="DNA/RNA_non-sp_Endonuclease_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001604">
        <property type="entry name" value="Endo_G_ENPP1-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020821">
        <property type="entry name" value="ENPP1-3/EXOG-like_nuc-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044925">
        <property type="entry name" value="His-Me_finger_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002591">
        <property type="entry name" value="Phosphodiest/P_Trfase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036024">
        <property type="entry name" value="Somatomedin_B-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001212">
        <property type="entry name" value="Somatomedin_B_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10151">
        <property type="entry name" value="ECTONUCLEOTIDE PYROPHOSPHATASE/PHOSPHODIESTERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10151:SF77">
        <property type="entry name" value="ECTONUCLEOTIDE PYROPHOSPHATASE_PHOSPHODIESTERASE FAMILY MEMBER 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01223">
        <property type="entry name" value="Endonuclease_NS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01663">
        <property type="entry name" value="Phosphodiest"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01033">
        <property type="entry name" value="Somatomedin_B"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00892">
        <property type="entry name" value="Endonuclease_NS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00477">
        <property type="entry name" value="NUC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00201">
        <property type="entry name" value="SO"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53649">
        <property type="entry name" value="Alkaline phosphatase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54060">
        <property type="entry name" value="His-Me finger endonucleases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90188">
        <property type="entry name" value="Somatomedin B domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00524">
        <property type="entry name" value="SMB_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50958">
        <property type="entry name" value="SMB_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="4" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="7" id="KW-0472">Membrane</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="29"/>
            <end position="54"/>
        </location>
    </feature>
    <feature type="domain" description="SMB" evidence="8">
        <location>
            <begin position="61"/>
            <end position="101"/>
        </location>
    </feature>
    <feature type="domain" description="SMB" evidence="8">
        <location>
            <begin position="102"/>
            <end position="146"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50958"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029336.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="869" mass="99706" checksum="AF5A1204D4E59888" modified="2019-05-08" version="1">QDPGSVSATENQLACQGFQNDLIVTFNSVFILFPILQVLSACVLTIILGCIFGLKPSCSRDVKTCKGRCFERTFGSCRCDRDCVELGNCCLDFQETCIQPAHIWTCNKFRCGEKRWPENRCSCSDDCVENKDCCVNYYIVCQGAKSWVEEKCEDINEPQCPPGFAKPPVLLFSLDGFRAEYLHTWSGLLPVISKLQKCGTYTPNMRPVYPTKTFPNHYTIVTGLYPESHGIIDNKIYDPHRNVSFTLKNAEKFNPQWYQGQPIWLTAMHQGLKAGTFFWPGSDVRINGTFPDLYRKYNGSIPFEERVIAILHWLQLPKDERPHFYTLYLEEPDSSGHNFGPVSSGVILALQRVDNIVGMLMDGLKQMNLHKCLNIILLSDHGMETASCSKTAYLSTYMENVQDITVIPGPAARLRPQNVPDEYFSCLENLTCEEDPDQHFKAFMKELLPKRFHYVNNDRIERVHFYLDPQWQLARKPSEIKYCNGGFHGSDNRFMSMQAIFIGFGPGFKFKTQVDPFENIEIYNLICDLLGITPAYNNGTHGSLNHLLKNPVYIPRHPEEESHPSVYPLMKTTSSLDIGCSCDLAGLPIRDFQQQLNHTASEVKKIEKYSLPYGRPRVLQKRSTYYLLYHHQYVSGYSQDLNMTLWSAYTDNRTASVENLSNCLHVDIRIPSSSSKTCSFYKNHHQLSYGFLFPPNTKKSSYEVLLNSNIVPMYHEFKVLWNYFYDVLLPKYTLARNGVNVVSGPVFDYDSNGLYDTPEKVKRLTHDPEVLVPTHYFIVLTSCKNISQTPLQCEGALDALSFIVPHRPDNSESCRHKRDSLWVEERMRFHTARVRDVELLTGLSFYQDRKQPVSDILQLKTYLPTFPME</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IN08</accession>
    <name>A0A452IN08_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Rab3 GTPase-activating protein catalytic subunit</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Endoplasmic reticulum</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Golgi apparatus</location>
            <location evidence="1">cis-Golgi network</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the Rab3-GAP catalytic subunit family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IN08"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033245.1">
        <property type="protein sequence ID" value="ENSGAGP00000029271.1"/>
        <property type="gene ID" value="ENSGAGG00000021013.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005783">
        <property type="term" value="C:endoplasmic reticulum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045700">
        <property type="entry name" value="Rab3GAP1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045698">
        <property type="entry name" value="Rab3GAP1_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026147">
        <property type="entry name" value="Rab3GAP1_conserved"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21422">
        <property type="entry name" value="RAB3 GTPASE-ACTIVATING PROTEIN CATALYTIC SUBUNIT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21422:SF9">
        <property type="entry name" value="RAB3 GTPASE-ACTIVATING PROTEIN CATALYTIC SUBUNIT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19533">
        <property type="entry name" value="Rab3-GAP_cat_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13890">
        <property type="entry name" value="Rab3-GTPase_cat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="9" id="KW-0333">Golgi apparatus</keyword>
    <keyword evidence="6" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Rab3GAP catalytic subunit conserved" evidence="11">
        <location>
            <begin position="387"/>
            <end position="541"/>
        </location>
    </feature>
    <feature type="domain" description="Rab3GAP catalytic subunit C-terminal" evidence="12">
        <location>
            <begin position="553"/>
            <end position="756"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="301"/>
            <end position="348"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="361"/>
            <end position="396"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="307"/>
            <end position="317"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="320"/>
            <end position="342"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="371"/>
            <end position="382"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004222"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00008856"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00015817"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF13890"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF19533"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029271.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="756" mass="84833" checksum="16BE1DAD99C7E103" modified="2019-05-08" version="1">GCPLTPLPPVSIAIRLTYVLQDWQQYFWPQQPPDIDALIGGEVGGLEFGKLPFGACEDPVSELHLAATWPHLTEGIIVDNDVYSDLDPIQAPQWSVRVRKADNLQCLLGDFLTEFFKLCRRKESTDEILGRSTFEEDGKEVADITHALSKLTEPAPVPIHKLSVTNMVHSAKKKIRKHRGVDDSPLNNEVLNTLMLVSKLNQTVISLSSRNLYSQFKSAPSDSLTYKLALCLCMINFYHGGVKGVAHLWQEFVVEMRRRWEKNYLIPELANGSPDLRCCLLHQKLQMLNCCIERKKARDEGKKPYTSDRSPSGSSGDTGKAVDRSGGDYQKETEKERGEVGKSWESWSDSEEEFFECLSDTEELKGNGQENGKKGGTKEGNKESANLKPEGRLHPHGKLTLLHPGEPLYIPVTQEPAPMTEDLLEEQSEVLAKLGTSAEGAHLRARMQSACLLSDMESFKAANPGCCLEDFVRWYSPRDYIEEEVVDEEGNRVLKGELSARMKIPSNMWVEAWETAKPVPARRQRRLFDDTREAEKVLHYLAVQKPADLARHLLPCVIHAAVLKVKEEETLEDIASVKKIIKQIISHSSKVLRFPNPEDKKLEEIIAQIMSAEAIIARARSLKAKFGIEKCDQEEEKEDLERFVNCLLEQPEVSVIGAGRGPAGSIIHKLFLNAQRVATMTSLDEELKRSGSSDERRLNLGAVTDFPPPAGRELILRITVPRPAPYSKPLPQRMYSVITKEDFRLAGAFSADTTFF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GP00</accession>
    <name>A0A452GP00_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Winged helix Storkhead-box1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452GP00"/>
    <dbReference type="Ensembl" id="ENSGAGT00000003919.1">
        <property type="protein sequence ID" value="ENSGAGP00000003398.1"/>
        <property type="gene ID" value="ENSGAGG00000002749.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000977">
        <property type="term" value="F:RNA polymerase II transcription regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019391">
        <property type="entry name" value="Storkhead-box_WHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040126">
        <property type="entry name" value="STOX1/2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22437:SF2">
        <property type="entry name" value="STORKHEAD-BOX PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22437">
        <property type="entry name" value="WINGED HELIX DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10264">
        <property type="entry name" value="WHD_Storkhead"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Winged helix Storkhead-box1" evidence="2">
        <location>
            <begin position="63"/>
            <end position="141"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="1"/>
            <end position="32"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="338"/>
            <end position="402"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="452"/>
            <end position="524"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="635"/>
            <end position="670"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="758"/>
            <end position="797"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="826"/>
            <end position="852"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="18"/>
            <end position="32"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="1">
        <location>
            <begin position="353"/>
            <end position="378"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="379"/>
            <end position="398"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="1">
        <location>
            <begin position="463"/>
            <end position="472"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="473"/>
            <end position="495"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="496"/>
            <end position="506"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="635"/>
            <end position="658"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="758"/>
            <end position="769"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="785"/>
            <end position="797"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF10264"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000003398.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="870" mass="97844" checksum="1062882DC668BB9F" modified="2019-05-08" version="1">MKKTRSTTLRRAWPSSDFSDRASDRMRSRSEKDYRLHKHFPPAFIAQASRGYMTSGDVSPINMSPISQSQFIPLGEILCLAISAMNSARKQVTQEALMEHLTTCFPGVPTPSPEILRHTLNMLVRERKIYPTPDGYFIVTPQTYFITPSLIRTNSKWYHLDERIPDRSQCTSPQQGTITPSTSGCVRDRTLLKNHYDSCHCCREDMHSIHASTLQRKSAKDCKDSCCPPSLCQVPPTEKSKSTVNFSYKAETLSKPKDVEKQSKKFGLKLFRLSFKKDKAKQLANFSAQFPPEEWPLRDEDTPTSIPREVEMEIIRRINPDLTVENVMRHTALMKKLEEEKAQRSKAGSSAHHSGRSKKSRNHRKSHGKSRSHSKTRVSKGDPSDGSHLDIPGEREYGFYDPLTRSPREGCFIIEHKGDNFIMHSNPSVIESHFPMTPEWDVSGELAKRRTEMPFPEPSRGSSHSKVHRSHSHTQDRRSRNERSNKAKERSRSMDNSKGPLSSATLGTPEDVAEGCSPDDQTTSQAYIDDSTLRPSQSLSHQRALISSASYKDVCIPEKVSGNVETPNSCGLLEQIKPTENLPSYSELNSCTTKSAIDDYFQCNTSSETVLTAPSPLGKNKEDHDTVTLTDGIKKLSPSERQSHHIAREPGVHKEESPKGPSGGPVVAGQTPEVIANGRLVQQHNSDSSGLDKRKEIFSKDTLFKPLHNTLSVNSYHKSSAPLLKPHQKTPSDTLPVRCEKLEQAIVTSVTQVLPVSQRQQEPIGNQEATFDYYNVSDDDDSEEGTNKNAEEEKNRDDVGTMQWLLEREKERDLQRKFEKNLTLLAPKETENSSNQRATHSARLDSMDSSSITVDSGFNSPRKLLMCCWV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GP02</accession>
    <name>A0A452GP02_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">ubiquitinyl hydrolase 1</fullName>
            <ecNumber evidence="2">3.4.19.12</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1 14 15">
            <text>Thiol-dependent hydrolysis of ester, thioester, amide, peptide and isopeptide bonds formed by the C-terminal Gly of ubiquitin (a 76-residue protein attached to proteins as an intracellular targeting signal).</text>
            <dbReference type="EC" id="3.4.19.12"/>
        </reaction>
    </comment>
    <dbReference type="EC" id="3.4.19.12" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452GP02"/>
    <dbReference type="Ensembl" id="ENSGAGT00000004015.1">
        <property type="protein sequence ID" value="ENSGAGP00000003481.1"/>
        <property type="gene ID" value="ENSGAGG00000002686.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004843">
        <property type="term" value="F:cysteine-type deubiquitinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036503">
        <property type="term" value="P:ERAD pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016579">
        <property type="term" value="P:protein deubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd06466">
        <property type="entry name" value="p23_CS_SGT1_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd06463">
        <property type="entry name" value="p23_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.70.10:FF:000012">
        <property type="entry name" value="ubiquitin carboxyl-terminal hydrolase 19 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.790:FF:000004">
        <property type="entry name" value="ubiquitin carboxyl-terminal hydrolase 19 isoform X9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="6.10.140.2220:FF:000004">
        <property type="entry name" value="ubiquitin carboxyl-terminal hydrolase 19 isoform X9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.790:FF:000074">
        <property type="entry name" value="Ubiquitin-specific peptidase 19"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.790">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.2220">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.70.10">
        <property type="entry name" value="Cysteine proteinases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007052">
        <property type="entry name" value="CS_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008978">
        <property type="entry name" value="HSP20-like_chaperone"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038765">
        <property type="entry name" value="Papain-like_cys_pep_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001394">
        <property type="entry name" value="Peptidase_C19_UCH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050185">
        <property type="entry name" value="Ub_carboxyl-term_hydrolase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028889">
        <property type="entry name" value="USP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018200">
        <property type="entry name" value="USP_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002893">
        <property type="entry name" value="Znf_MYND"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21646">
        <property type="entry name" value="UBIQUITIN CARBOXYL-TERMINAL HYDROLASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21646:SF74">
        <property type="entry name" value="UBIQUITIN CARBOXYL-TERMINAL HYDROLASE 19"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04969">
        <property type="entry name" value="CS"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00443">
        <property type="entry name" value="UCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16602">
        <property type="entry name" value="USP19_linker"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01753">
        <property type="entry name" value="zf-MYND"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54001">
        <property type="entry name" value="Cysteine proteinases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF144232">
        <property type="entry name" value="HIT/MYND zinc finger-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49764">
        <property type="entry name" value="HSP20-like chaperones"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51203">
        <property type="entry name" value="CS"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00972">
        <property type="entry name" value="USP_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50235">
        <property type="entry name" value="USP_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01360">
        <property type="entry name" value="ZF_MYND_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50865">
        <property type="entry name" value="ZF_MYND_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <keyword evidence="4 7" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="CS" evidence="11">
        <location>
            <begin position="46"/>
            <end position="135"/>
        </location>
    </feature>
    <feature type="domain" description="CS" evidence="11">
        <location>
            <begin position="351"/>
            <end position="453"/>
        </location>
    </feature>
    <feature type="domain" description="USP" evidence="9">
        <location>
            <begin position="567"/>
            <end position="1151"/>
        </location>
    </feature>
    <feature type="domain" description="MYND-type" evidence="10">
        <location>
            <begin position="861"/>
            <end position="903"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="1"/>
            <end position="49"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="142"/>
            <end position="253"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="463"/>
            <end position="508"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="520"/>
            <end position="546"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="16"/>
            <end position="49"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="167"/>
            <end position="177"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="8">
        <location>
            <begin position="223"/>
            <end position="240"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="488"/>
            <end position="505"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000707"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00012759"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50235"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50865"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS51203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000003481.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1151" mass="126013" checksum="432BC89C06D9141B" modified="2019-05-08" version="1">MSSSPSAPGQRGAARGLEDATSKKKQKDRANQESKESKRPPGSDPKKDLLLDWKQNADEVIVKLNLGSGALKLEEVDAAFTDTDCVVRLPDGRQWSGQFYGEIESSCSKVQGKKGKFLQLVLQKKIPLHTWASLLKKRKDGSKELAKGAVSPNGKEQPLPAQATPEEPARSKREFPNSKRAPGRGEAPEEKSPASPGLQSGPSAKRAGYLKVALTEEEPNAGVPGGLEPGKGPSGRGGSLQNGRATCGDAATGLTPPLMKAEVLQKEPVLVGMQPLAAPSESFPQHMAPCLEKRALQPAASQCLAAPGEGPEAVLAPASPLLSRDAEKSDWSKEDVALEAAADEPEPMVSLSLVKNDSYEKGSDSVVVHVYVKEIHRETSKVLFREQDFTLVFQTSDANFLRVHPSCGPRTVFRWQVKLRNLIEPDQCTYNFTVSRINICLKKRHSQRWGGLEAPAARGAVGGAKVAMPTGPTPLDKSQPGGNQHPLASKEEARAGEKEKPRAEDGALEGVVARTATEHVTVKQEPHVPSPKPMCMVPPMTHSPVSSESVEEDEEEEEKKVCLPGFTGLVNLGNTCFMNSVIQSLSNTRELRDYFHDRSFEAEINYTNPLGTGGRLAIGFAVLLRALWKGTHHAFQPSKLKAIVASKASQFTGYAQHDAQEFMAFLLDGLHEDLNRIQNKPYTETVDSDGRPDEVVAEEAWQRHKMRNDSFIVDLFQGQYKSKLVCPVCAKVSITFDPFLYLPVPLPQKQKVLTVYYFAKEPHKKPVKFLVSISKENSSAAEVLDSVAHSMRVKPENLRLAEVIKSRFHRVFPPSQSLDTVSPTDLLLCFEVLSPELAKERVVVLQVQQRPQVPSVPISKCAACQKAQQSEDEKLKRCTRCYRVGYCNVACQKTHWPDHKALCRPENIGFPFLISVPESRLTYSRLAQLLEGYARYSVSVFQPPFQLGPQPLLPEKLDLPARSSCVAVPPAPEAGDANRAPHRQEPQLSTLELHPELGEASAVPRSSALGSDSGCSEHSLEVQGEGCWVKEPSYERGPKPEAAVPGYQHAPDALSAHATQFYINKIDGASREHKLEEKGDAPLELTDDCSLALVWKNNERLKEFVLVESKELECVEDPGSASEAARAGHFTLGQCLNLFTKPEVLAPEEAW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GP03</accession>
    <name>A0A452GP03_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">CNH domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endomembrane system</location>
            <topology evidence="1">Peripheral membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the VAM6/VPS39 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GP03"/>
    <dbReference type="Ensembl" id="ENSGAGT00000004175.1">
        <property type="protein sequence ID" value="ENSGAGP00000003619.1"/>
        <property type="gene ID" value="ENSGAGG00000002901.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0012505">
        <property type="term" value="C:endomembrane system"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006914">
        <property type="term" value="P:autophagy"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034058">
        <property type="term" value="P:endosomal vesicle fusion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001180">
        <property type="entry name" value="CNH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032914">
        <property type="entry name" value="Vam6/VPS39/TRAP1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019452">
        <property type="entry name" value="VPS39/TGF_beta_rcpt-assoc_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019453">
        <property type="entry name" value="VPS39/TGFA1_Znf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12894">
        <property type="entry name" value="CNH DOMAIN CONTAINING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12894:SF49">
        <property type="entry name" value="VAM6_VPS39-LIKE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00780">
        <property type="entry name" value="CNH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10366">
        <property type="entry name" value="Vps39_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10367">
        <property type="entry name" value="zf-Vps39_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00036">
        <property type="entry name" value="CNH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50219">
        <property type="entry name" value="CNH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="CNH" evidence="4">
        <location>
            <begin position="15"/>
            <end position="292"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004184"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00038201"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50219"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000003619.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="841" mass="96545" checksum="3AAA5E953EE87D96" modified="2019-05-08" version="1">MHDAFEPVPILEKLPLQIDCLAAWEEWLLVGTKQGHLLLYRIRKDIGEVTSSESGCCNRFEVTLEKSNKNFSKKIQQIHVVSQFKILVSLLENNIYVHDLLTFQQITTVSKAKGASLFTCDLQHSDTGEEVLRMCVAVRKKLQLYFWKDREFYELQGDFSVPDVPKSMAWCENSICVGFKRDYYLIRVDGKGSIKELFPTGKQLEPLVAPLADGKVAVGQDDLTVVLNEEGICTQKGALNWTDIPIAMEHQPPYIIAVLPRYVEIRTFEPRLLVQSIELQRPRFITSGGTNIVYVASNHFVWRLLPVSIATQIQQLLQDKQFELALQLAEMKDDSDSEKQQQIHHIKNLYAFNLFCQKRFDESMQVFAKLGTDPTHVMGLYPDLLPTDYRKQLQYPNPLPVLSGAELEKAHLALIDYLTQKRSQLVKKLNDSDHQSSTSPLMEGTPTIKSKKKLLQIIDTTLLKCYLHTNVALVAPLLRLENNHCHIEESEHVLKKAHKYSELIILYEKKGLHEKALQVLVDQSKKANSPLKGHERTVQYLQHLGTENLHLVFLYSTWVLRDFPDDGLKIFTEDLPEVESLPRDKVLNFLIESFKSLAIPYLEHIIHVWEETGSEFHNCLIQLYCEKVQGLMKEYLSSFPAGLLEERALLLGRMGKHEQALIIYVHILKDTKMAELYCHKHYDRSKDGNKDVYLALLRMYLSPPSVHCLGPIKMELLEPQANLQAALQVLELHHSKLDTTKAINLLPANTQISEIRIFLEKVLEENAQKKRFNQVLKNLLHAEFLRVQEERILHQQVKCIITEEKVCTVCKKKIGNSAFARYPNAVVVHYFCSKEVSTADT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GP09</accession>
    <name>A0A452GP09_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Caspase family p20 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the peptidase C14A family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GP09"/>
    <dbReference type="Ensembl" id="ENSGAGT00000004205.1">
        <property type="protein sequence ID" value="ENSGAGP00000003641.1"/>
        <property type="gene ID" value="ENSGAGG00000002937.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004197">
        <property type="term" value="F:cysteine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043525">
        <property type="term" value="P:positive regulation of neuron apoptotic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1460">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029030">
        <property type="entry name" value="Caspase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033139">
        <property type="entry name" value="Caspase_cys_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002398">
        <property type="entry name" value="Pept_C14"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011600">
        <property type="entry name" value="Pept_C14_caspase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001309">
        <property type="entry name" value="Pept_C14_p20"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015917">
        <property type="entry name" value="Pept_C14A"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10454">
        <property type="entry name" value="CASPASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10454:SF155">
        <property type="entry name" value="CASPASE-14-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00656">
        <property type="entry name" value="Peptidase_C14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00115">
        <property type="entry name" value="CASc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52129">
        <property type="entry name" value="Caspase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01122">
        <property type="entry name" value="CASPASE_CYS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50208">
        <property type="entry name" value="CASPASE_P20"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Caspase family p20" evidence="2">
        <location>
            <begin position="37"/>
            <end position="148"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00010134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="PROSITE" id="PS50208"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000003641.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="226" mass="26020" checksum="C255A6EFCD3CF601" modified="2019-05-08" version="1">MSSPPPIPYKSKCLGHSDMYDMSKTRVAFMMCVTTGRRGAEKDIEVMNKWFDKYQFETPSGQCIDPNGEFYRQKSLEEFRDQINQSEDEISCCLITLMSHGRSHGLIQGKDGDKVDLDDIFALFNNIQCPKLQEKPKIFIIQACRGEDHGVEEIEHEFKEADCTCNLIAIRRLPTASDYYVVYSTQKGKRDEAKLGPSYIHELWLIPVGVCVREYRIWPFINAISD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GXE7</accession>
    <name>A0A452GXE7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="23">Nitric oxide synthase 3</fullName>
            <ecNumber evidence="7">1.14.13.39</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="20">Constitutive NOS</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="26">EC-NOS</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="24">NOS type III</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="25">Nitric oxide synthase, endothelial</fullName>
        </alternativeName>
    </protein>
    <organism evidence="30 31">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="31" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="30" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="30" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="21">
            <text>2 L-arginine + 3 NADPH + 4 O2 + H(+) = 2 L-citrulline + 2 nitric oxide + 3 NADP(+) + 4 H2O</text>
            <dbReference type="Rhea" id="RHEA:19897"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:15379"/>
            <dbReference type="ChEBI" id="CHEBI:16480"/>
            <dbReference type="ChEBI" id="CHEBI:32682"/>
            <dbReference type="ChEBI" id="CHEBI:57743"/>
            <dbReference type="ChEBI" id="CHEBI:57783"/>
            <dbReference type="ChEBI" id="CHEBI:58349"/>
            <dbReference type="EC" id="1.14.13.39"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="21">
            <dbReference type="Rhea" id="RHEA:19898"/>
        </physiologicalReaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="2">
            <name>(6R)-L-erythro-5,6,7,8-tetrahydrobiopterin</name>
            <dbReference type="ChEBI" id="CHEBI:59560"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="4">
            <name>FAD</name>
            <dbReference type="ChEBI" id="CHEBI:57692"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>FMN</name>
            <dbReference type="ChEBI" id="CHEBI:58210"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="3">
            <name>heme b</name>
            <dbReference type="ChEBI" id="CHEBI:60344"/>
        </cofactor>
    </comment>
    <comment type="subunit">
        <text evidence="22">Homodimer. Interacts with NOSIP and NOSTRIN. Interacts with HSP90AB1. Forms a complex with ASL, ASS1 and SLC7A1; the complex regulates cell-autonomous L-arginine synthesis and citrulline recycling while channeling extracellular L-arginine to nitric oxide synthesis pathway.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="5">Cell membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the NOS family.</text>
    </comment>
    <dbReference type="EC" id="1.14.13.39" evidence="7"/>
    <dbReference type="AlphaFoldDB" id="A0A452GXE7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000007880.1">
        <property type="protein sequence ID" value="ENSGAGP00000006788.1"/>
        <property type="gene ID" value="ENSGAGG00000000423.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050660">
        <property type="term" value="F:flavin adenine dinucleotide binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010181">
        <property type="term" value="F:FMN binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0020037">
        <property type="term" value="F:heme binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050661">
        <property type="term" value="F:NADP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004517">
        <property type="term" value="F:nitric-oxide synthase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006809">
        <property type="term" value="P:nitric oxide biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00795">
        <property type="entry name" value="NOS_oxygenase_euk"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.440.10:FF:000001">
        <property type="entry name" value="Endothelial nitric oxide synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.360:FF:000003">
        <property type="entry name" value="Nitric oxide synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1230.10:FF:000001">
        <property type="entry name" value="Nitric oxide synthase, brain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.360">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.990.10">
        <property type="entry name" value="NADPH-cytochrome p450 Reductase, Chain A, domain 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.340.10">
        <property type="entry name" value="Nitric Oxide Synthase, Chain A, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1230.10">
        <property type="entry name" value="Nitric Oxide Synthase, Chain A, domain 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.440.10">
        <property type="entry name" value="Nitric Oxide Synthase,Heme Domain,Chain A domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.30.10">
        <property type="entry name" value="Translation factors"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003097">
        <property type="entry name" value="CysJ-like_FAD-binding"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001094">
        <property type="entry name" value="Flavdoxin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008254">
        <property type="entry name" value="Flavodoxin/NO_synth"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029039">
        <property type="entry name" value="Flavoprotein-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023173">
        <property type="entry name" value="NADPH_Cyt_P450_Rdtase_alpha"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050607">
        <property type="entry name" value="NOS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044943">
        <property type="entry name" value="NOS_dom_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044940">
        <property type="entry name" value="NOS_dom_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044944">
        <property type="entry name" value="NOS_dom_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012144">
        <property type="entry name" value="NOS_euk"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004030">
        <property type="entry name" value="NOS_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036119">
        <property type="entry name" value="NOS_N_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017938">
        <property type="entry name" value="Riboflavin_synthase-like_b-brl"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43410:SF1">
        <property type="entry name" value="NITRIC OXIDE SYNTHASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43410">
        <property type="entry name" value="NITRIC OXIDE SYNTHASE OXYGENASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00667">
        <property type="entry name" value="FAD_binding_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00258">
        <property type="entry name" value="Flavodoxin_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02898">
        <property type="entry name" value="NO_synthase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000333">
        <property type="entry name" value="NOS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00369">
        <property type="entry name" value="FLAVODOXIN"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52218">
        <property type="entry name" value="Flavoproteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56512">
        <property type="entry name" value="Nitric oxide (NO) synthase oxygenase domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF63380">
        <property type="entry name" value="Riboflavin synthase domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50902">
        <property type="entry name" value="FLAVODOXIN_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="16" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="8" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="14" id="KW-0274">FAD</keyword>
    <keyword evidence="11" id="KW-0285">Flavoprotein</keyword>
    <keyword evidence="12" id="KW-0288">FMN</keyword>
    <keyword evidence="10 27" id="KW-0349">Heme</keyword>
    <keyword evidence="18 27" id="KW-0408">Iron</keyword>
    <keyword evidence="19" id="KW-0472">Membrane</keyword>
    <keyword evidence="13 27" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="15" id="KW-0521">NADP</keyword>
    <keyword evidence="17" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="9" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="31" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Flavodoxin-like" evidence="29">
        <location>
            <begin position="472"/>
            <end position="655"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="28">
        <location>
            <begin position="1"/>
            <end position="36"/>
        </location>
    </feature>
    <feature type="binding site" description="axial binding residue" evidence="27">
        <location>
            <position position="136"/>
        </location>
        <ligand>
            <name>heme b</name>
            <dbReference type="ChEBI" id="CHEBI:60344"/>
        </ligand>
        <ligandPart>
            <name>Fe</name>
            <dbReference type="ChEBI" id="CHEBI:18248"/>
        </ligandPart>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001917"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00001950"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00001970"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00001974"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00006267"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00012989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022617"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022630"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022643"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022827"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022857"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00029794"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00047419"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00049679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00049764"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00049791"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00049814"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00049818"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="PIRSR" id="PIRSR000333-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="29">
        <source>
            <dbReference type="PROSITE" id="PS50902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="30">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000006788.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="31">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="839" mass="93219" checksum="0C2D93FD82CB166E" modified="2019-05-08" version="1">GWVHPGAHPSSLVSPFPGPGAGVHAGSDPNSPQPQLCPSVLQELPCTGKRCLGSLMFPKQMVCQADEGPPAKEELLPLARDFIHQYYSSIKRADSRVHMQRVQEVEQEIRATGTYQLLEPELIFGAKQAWRNAARCLGRIQWNKLQVFDARDCASVQEMFGFLCTHLQYATNRGNLRSAITIFPQRMPGRGDFRIWNSQLIRYAGYPQPDGSVLGDPANVDITELCIQHGWTPGHGRFDVLPLLLQAPDEAPELFLLPPELVLEVPIQHPTLAWFGELGLRWHALPAVANMLLEIGGLQFPAAPFNGWYVGSEIGMRNFCDSNRYNLLQEVALHMGLDTGKTSSLWKDKAAVEINVAVLHSFQVAKVTIVDHHVATESFVKHMDNELQTRGGCPADWVWIVPPISGSLTPVFHQEMVNYLLSPCFRYQPDAWKVYTSKGTSITRKKTFKEVANAVKFSAKLMVHVMARRVKATILYATETGKSRTYAWNLGELFRRAFDPKVLCMDEYDIVSLEHETLVLVVTSTFGNGDPPESGESFAKALMEMSCPYVSSTQPEQQKSYKVRFNSVSQSDGLVASWKKKRKESSNTDSAGALGTLRFSVFGLGSRAYPHFCAFAHAVDTRLEELGGERILAMGEGDELCAQEETFRTWAKRVFQAACDTFCVGDDAEEAAREIFTTRRSWKRQKYQLSVHANPPDMLTGLSQVHKRRVFPCRVLSVQNLQSKESSRSTLLVRLDTAGQPELQYLPGDHVGIFPANRLELVQELLERVEDPPPDEEPVAVEFLEKATSALGAGNLVLVPGGSQGGFWGGQCQCLGGPMRVWGDPGASAWRIPWGVLRG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H0N2</accession>
    <name>A0A452H0N2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">AMP deaminase</fullName>
            <ecNumber evidence="9">3.5.4.6</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="8">AMP deaminase plays a critical role in energy metabolism. Catalyzes the deamination of AMP to IMP and plays an important role in the purine nucleotide cycle.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="9">
            <text>AMP + H2O + H(+) = IMP + NH4(+)</text>
            <dbReference type="Rhea" id="RHEA:14777"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:28938"/>
            <dbReference type="ChEBI" id="CHEBI:58053"/>
            <dbReference type="ChEBI" id="CHEBI:456215"/>
            <dbReference type="EC" id="3.5.4.6"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 9">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
    </comment>
    <comment type="pathway">
        <text evidence="2">Purine metabolism; IMP biosynthesis via salvage pathway; IMP from AMP: step 1/1.</text>
    </comment>
    <comment type="similarity">
        <text evidence="3 9">Belongs to the metallo-dependent hydrolases superfamily. Adenosine and AMP deaminases family.</text>
    </comment>
    <dbReference type="EC" id="3.5.4.6" evidence="9"/>
    <dbReference type="AlphaFoldDB" id="A0A452H0N2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000009214.1">
        <property type="protein sequence ID" value="ENSGAGP00000007995.1"/>
        <property type="gene ID" value="ENSGAGG00000006352.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00591">
        <property type="reaction ID" value="UER00663"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003876">
        <property type="term" value="F:AMP deaminase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046033">
        <property type="term" value="P:AMP metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097009">
        <property type="term" value="P:energy homeostasis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032264">
        <property type="term" value="P:IMP salvage"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="CDD" id="cd01319">
        <property type="entry name" value="AMPD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.800.20:FF:000001">
        <property type="entry name" value="AMP deaminase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.20.20.140:FF:000035">
        <property type="entry name" value="Probable amp deaminase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.800.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.20.20.140">
        <property type="entry name" value="Metal-dependent hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006650">
        <property type="entry name" value="A/AMP_deam_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006329">
        <property type="entry name" value="AMPD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032466">
        <property type="entry name" value="Metal_Hydrolase"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01429">
        <property type="entry name" value="AMP_deaminase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11359">
        <property type="entry name" value="AMP DEAMINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11359:SF3">
        <property type="entry name" value="AMP DEAMINASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19326">
        <property type="entry name" value="AMP_deaminase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF001251">
        <property type="entry name" value="AMP_deaminase_met"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51556">
        <property type="entry name" value="Metallo-dependent hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00485">
        <property type="entry name" value="A_DEAMINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 9" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="4 9" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7" id="KW-0546">Nucleotide metabolism</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0862">Zinc</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001947"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004955"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00006676"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023080"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00054146"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PIRNR" id="PIRNR001251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007995.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="764" mass="88890" checksum="A7D163F0187CBFFD" modified="2019-05-08" version="1">MDGKYKEIAEELFSRSLVESDMRSAPYEFPEDSPIEQLEERRQRLERQISQDVKLEPDILLRAKQDFLKIDSAADLQLYKEQNEDLVDHLPKERDVLLEREFQRVTISGEETCGVPFTDLLDAAKSVVKALFIREKYMGLSLQSFCKTTARFLEELSGKPLETRFYEEIPETPVAADAPVHPPFTDQHPYETCLPEAMPADLGFGLRMVSGVIHVYTKRDVVDRSTELDLLYPDLQEFIADMNVLMALIINGPIKSFCYRRLQYLSSKFQMHVLLNEMKELAAQKKVPHRDFYNIRKVDTHIHASSCMNQKHLLRFIKRTMKKHLDEIVHVEKGKEQTLKEVFETMNLTAYDLSVDTLDVHADRNTFHRFDKFNTKYNPIGESILREIFIKTDNQVSGKYFAHIIKEVMSDLEESKYQNAELRLSIYGRSRDEWDKLAKWAVSHKVHSNNVRWLVQVPRLFDVYRTKKQLANFQEMLENIFLPLFEATIYPASHLELHLFLEHVEGFDSVDDESKPEHRIFNQDSPLPGTWVEEDNPPYSYYLYYMYANMTVLNHLRRKRGFHTFVLRPHCGEAGPIHHLVSAFMLSENISHGLLLRKAPVLQYLYYLAQIGIAMSPLSNNSLFLSYHRNPLPEYLSRGLMVSLSTDDPLQFHFTKEPLMEEYSIATQVWKLSSCDMCELARNSVLMSGFSHKVKSYWLGPNYLKEGSEGNDIRRTNVPDIRVSYRFETLCQELTLITQAVQTAELETIQEEDSLIVSSSLGMQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H0N6</accession>
    <name>A0A452H0N6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">C2H2-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">May be involved in transcriptional regulation.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the krueppel C2H2-type zinc-finger protein family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H0N6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000009223.1">
        <property type="protein sequence ID" value="ENSGAGP00000008004.1"/>
        <property type="gene ID" value="ENSGAGG00000006323.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000122">
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    <keyword evidence="10" id="KW-0804">Transcription</keyword>
    <keyword evidence="8" id="KW-0805">Transcription regulation</keyword>
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        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
    <sequence length="682" mass="77637" checksum="8CAF56EFA19DCAA7" modified="2019-05-08" version="1">GRDDTNISDCIPRASRPWICLTTASVGLERPEENFSSHSDLITCNRINLEETRYMCHECGKSFSRSSTLITHCRIHTGEKAYTCSECGKSFNRGLDLITHRTIHVGEKPYGCSECGKCFIHRSGLILHQRMHTGETPYTCAECGKSFRCSSHLFRHQRIHTGEKPYTCTECGKSFNRSSNLITHRRIHTGETPYMCSECGKSFSRRSHLFRHERIHTGEMPYTCSECGKSFNQSSALITHRRIHTGEMPYTCSECGKSVSHISALITHRRIHTGEKPYACTVCGKRFSRNSHLIRHQRIHTGEKPYTCTECGKSFNQRSSLITHCRIHTGETPYMCTECGKSFSHCSALTTHCRSHTGERPYTCSECGKSFNQSSHLITHRRIHGGEKPYGCSECGKCFIHRSDLILHQRIHTGEMPNTCPECGKNFTCSSQLFRHQRIHTEEKPYKCTECGKSFSRNSHLIRHQRIHTGEKPYTCTECGKSFNQSSNLITHRTSHTGKTQYTCSECGKNFSQSSRIHTGEKPYACTVCGKRFSRNSHLIRHQRIHTGEKPYTCTVCGKRFSRNSHLIRHQRIHTGEKPYTCTECGKSFNRSSSLITHYTVHTGETPYTCSECGKSFNQSSALITHHRIHTGEMPYTCSESGKGREGKITFLLLSSTQRAFFCSCIFRSNREGKCHKQISKS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H0N8</accession>
    <name>A0A452H0N8_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="12">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H0N8"/>
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        <property type="gene ID" value="ENSGAGG00000006312.1"/>
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        <property type="project" value="UniProtKB-KW"/>
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    </dbReference>
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        <property type="term" value="F:zinc ion binding"/>
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        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045892">
        <property type="term" value="P:negative regulation of DNA-templated transcription"/>
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        <property type="project" value="TreeGrafter"/>
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        <property type="entry name" value="Polyhomeotic-like protein 2 isoform 1"/>
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        <property type="entry name" value="SAM/pointed_sf"/>
    </dbReference>
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        <property type="entry name" value="POLYCOMB GROUP PROTEIN"/>
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    </dbReference>
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    </dbReference>
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        <property type="entry name" value="ZF_FCS"/>
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    </dbReference>
    <proteinExistence type="predicted"/>
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    <keyword evidence="6" id="KW-0238">DNA-binding</keyword>
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    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <keyword evidence="4 8" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="FCS-type" evidence="11">
        <location>
            <begin position="770"/>
            <end position="804"/>
        </location>
    </feature>
    <feature type="domain" description="SAM" evidence="10">
        <location>
            <begin position="919"/>
            <end position="983"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
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            <begin position="1"/>
            <end position="32"/>
        </location>
    </feature>
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        </location>
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            <begin position="422"/>
            <end position="489"/>
        </location>
    </feature>
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        <location>
            <begin position="823"/>
            <end position="851"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="7"/>
            <end position="26"/>
        </location>
    </feature>
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        <location>
            <begin position="101"/>
            <end position="112"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="113"/>
            <end position="130"/>
        </location>
    </feature>
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        <location>
            <begin position="140"/>
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        </location>
    </feature>
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            <end position="259"/>
        </location>
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            <begin position="267"/>
            <end position="280"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="9">
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            <begin position="371"/>
            <end position="383"/>
        </location>
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        <location>
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            <end position="396"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="425"/>
            <end position="451"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="470"/>
            <end position="489"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="823"/>
            <end position="833"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00367"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50105"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS51024"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007922.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="983" mass="106436" checksum="F5F41040374A5B3C" modified="2019-05-08" version="1">MENESNATTCSACTTTTTTTTTSSSSRTQPPQISVYSGSDRHAVQVIQQALHRPPSSAAQYLQQMYAAQQQHLMLQTAALQQQHLSSTQFQSLAAVPQASLSGGRQCSSPTGSVTQQSSISQTSINLSTSPTPPQLISRSQTSNTTSSSITQQTMLLGSTSPTLSASQAQMYLRAQMLIFTPATTVAAVPSDIPVVSSCSSSCQSAATQVQNLTLRHQKLGVLCSSQNGPPKSSSQAQSLALCTNKTVTGTKASQSDSPDSNRKGESPTSECRSTSVTRTSSIHQLIAPASYSAIQPHSLIKHQQIPLHSPPPKISHHQLILQQQQVQPITLQTPPDQEPPPSQHYVPLQSHGLPPVPNSVQSQHCSPVHIHPPPLTISPTPSPSAQQSVVVSPTPAHSPSQSPTIIIHPQALIQSQPHPLMPSAVQSEQNPQQPTTSQVRPVAQQLNLPSHLTVPPSPVVHIGPVDQPNLMSPGQQTMSPTPHQQYPTLQSVPIPVATPPQLSTSPTQIQQLPLQSVQSLQMQPEILSQGQVLVQNTLVSEEELPAAEALVQLPFQTLPPPQTVAVNLQVQPSVPVETPVIYQVENVCEEEMPEESECVRMVRTPTPPTLSPPIITLGNGEELASEAPLSEQVGLPAVASSVSASVIKSPSDPSHASIPPPPLLLPAAATRSNITSMPNSIPNLENKPPQAIVKPQILTHVIEGFVIQEGLEPFPVSRSSLLVEQPAKKRLLMEGQVMSVESELQNNTKHAENSSDTEIEDIITEEGLDEMENDLLKCEFCGKMGYANNFLRSKRFCSMSCAKRHNVSCTKKFGLFTSDKNSHWNRKSDSRSLGRRARRPSGSDGSSREHFLRQLPVTYPSAEDDLVSHDDGVPAAMTTRLRRQSERERERELRELKIRKMPENVDLLPSVQADPSVWTVDEVWAFIRSLPGCQDIADEFRAQEIDGQALLLLKEDHLMNAMNIKLGPALKICARINSLKES</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H0N9</accession>
    <name>A0A452H0N9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">Acetylcholine receptor subunit gamma</fullName>
        </recommendedName>
    </protein>
    <organism evidence="25 26">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="26" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="25" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="25" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">After binding acetylcholine, the AChR responds by an extensive change in conformation that affects all subunits and leads to opening of an ion-conducting channel across the plasma membrane.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>K(+)(in) = K(+)(out)</text>
            <dbReference type="Rhea" id="RHEA:29463"/>
            <dbReference type="ChEBI" id="CHEBI:29103"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>Na(+)(in) = Na(+)(out)</text>
            <dbReference type="Rhea" id="RHEA:34963"/>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
        </reaction>
    </comment>
    <comment type="subunit">
        <text evidence="2">Pentamer of two alpha chains, and one each of the beta, delta, and gamma chains.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="17">Postsynaptic cell membrane</location>
            <topology evidence="17">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="20">Belongs to the ligand-gated ion channel (TC 1.A.9) family. Acetylcholine receptor (TC 1.A.9.1) subfamily. Gamma/CHRNG sub-subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H0N9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000009371.1">
        <property type="protein sequence ID" value="ENSGAGP00000008137.1"/>
        <property type="gene ID" value="ENSGAGG00000006424.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0022848">
        <property type="term" value="F:acetylcholine-gated monoatomic cation-selective channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004888">
        <property type="term" value="F:transmembrane signaling receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd19029">
        <property type="entry name" value="LGIC_ECD_nAChR_G"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd19064">
        <property type="entry name" value="LGIC_TM_nAChR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.390:FF:000010">
        <property type="entry name" value="Nicotinic acetylcholine receptor subunit epsilon"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.390:FF:000036">
        <property type="entry name" value="Nicotinic acetylcholine receptor subunit gamma"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.70.170.10:FF:000012">
        <property type="entry name" value="Nicotinic acetylcholine receptor subunit gamma"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.70.170.10">
        <property type="entry name" value="Neurotransmitter-gated ion-channel ligand-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.390">
        <property type="entry name" value="Neurotransmitter-gated ion-channel transmembrane domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006202">
        <property type="entry name" value="Neur_chan_lig-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036734">
        <property type="entry name" value="Neur_chan_lig-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006201">
        <property type="entry name" value="Neur_channel"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036719">
        <property type="entry name" value="Neuro-gated_channel_TM_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038050">
        <property type="entry name" value="Neuro_actylchol_rec"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006029">
        <property type="entry name" value="Neurotrans-gated_channel_TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018000">
        <property type="entry name" value="Neurotransmitter_ion_chnl_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002394">
        <property type="entry name" value="Nicotinic_acetylcholine_rcpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18945">
        <property type="entry name" value="NEUROTRANSMITTER GATED ION CHANNEL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02931">
        <property type="entry name" value="Neur_chan_LBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02932">
        <property type="entry name" value="Neur_chan_memb"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00254">
        <property type="entry name" value="NICOTINICR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00252">
        <property type="entry name" value="NRIONCHANNEL"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90112">
        <property type="entry name" value="Neurotransmitter-gated ion-channel transmembrane pore"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF63712">
        <property type="entry name" value="Nicotinic receptor ligand binding domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00236">
        <property type="entry name" value="NEUROTR_ION_CHANNEL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="16 22" id="KW-0407">Ion channel</keyword>
    <keyword evidence="9 22" id="KW-0406">Ion transport</keyword>
    <keyword evidence="15" id="KW-1071">Ligand-gated ion channel</keyword>
    <keyword evidence="10 22" id="KW-0472">Membrane</keyword>
    <keyword evidence="14" id="KW-0628">Postsynaptic cell membrane</keyword>
    <keyword evidence="12" id="KW-0675">Receptor</keyword>
    <keyword evidence="26" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="8" id="KW-0770">Synapse</keyword>
    <keyword evidence="5 22" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 22" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="3 22" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="22">
        <location>
            <begin position="235"/>
            <end position="257"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="22">
        <location>
            <begin position="269"/>
            <end position="289"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="22">
        <location>
            <begin position="301"/>
            <end position="324"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="22">
        <location>
            <begin position="467"/>
            <end position="489"/>
        </location>
    </feature>
    <feature type="domain" description="Neurotransmitter-gated ion-channel ligand-binding" evidence="23">
        <location>
            <begin position="18"/>
            <end position="234"/>
        </location>
    </feature>
    <feature type="domain" description="Neurotransmitter-gated ion-channel transmembrane" evidence="24">
        <location>
            <begin position="241"/>
            <end position="484"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00003328"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00011293"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023257"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00034104"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00034430"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00036239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00038482"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00039195"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="RuleBase" id="RU000687"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="Pfam" id="PF02931"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="Pfam" id="PF02932"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000008137.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="506" mass="57743" checksum="EA864ED614EC93B8" modified="2019-05-08" version="1">MSQMCASVSLSVLCRNQEEKLLNDLMTNYNRNLRPAQGEGDIINVTLKLTLTNLISLNEREETLTTNIWIEMKWCDYRLQWDPDEYDNIQILRVPSTEVWLPDIVLENNIDGVFEITLYCNTLLDSNGCLLWMPPAIYRTSCAIFVTYFPFDWQNCSMVFQSQTYSANEINLLLMEEDGQTIEWIVIDPEAFTENGEWAIKHRPAKKIVNAERFTPDDLGYQQVVFYLIIQRKPLFYVINIIVPCVLISSVAVLVYFLPAKAGGQKCTVSINVLLAQTVFLFLIAQKVPETSDAMPLIGKYLTFLLVVTVVIVANAVIVLNVSLRTPNTHAMSQKVRQICLRQLPHFLGMHMRRSDMAPRLLLARRRSSLGLMVKADEYMLWKARTELVFEKQKERAGLMKAVLEKIGKGLENRCTQDLCHSLVQAGPEIWACVDACNHIAKTLREQNDFDNENEEWILVGRVIDRVCFLVMASLFVLGTVSIFLMAHFNQAPAAPFPGDPKCYLP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H102</accession>
    <name>A0A452H102_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="15">Anion exchange protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Electrogenic sodium/bicarbonate cotransporter with a Na(+):HCO3(-) stoichiometry varying from 1:2 to 1:3. May regulate bicarbonate influx/efflux at the basolateral membrane of cells and regulate intracellular pH.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="13">
            <text>2 hydrogencarbonate(out) + Na(+)(out) = 2 hydrogencarbonate(in) + Na(+)(in)</text>
            <dbReference type="Rhea" id="RHEA:72215"/>
            <dbReference type="ChEBI" id="CHEBI:17544"/>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="12">
            <text>3 hydrogencarbonate(out) + Na(+)(out) = 3 hydrogencarbonate(in) + Na(+)(in)</text>
            <dbReference type="Rhea" id="RHEA:72219"/>
            <dbReference type="ChEBI" id="CHEBI:17544"/>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Basolateral cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="15">Membrane</location>
            <topology evidence="15">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 15">Belongs to the anion exchanger (TC 2.A.31) family.</text>
    </comment>
    <comment type="caution">
        <text evidence="15">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H102"/>
    <dbReference type="Ensembl" id="ENSGAGT00000009265.1">
        <property type="protein sequence ID" value="ENSGAGP00000008042.1"/>
        <property type="gene ID" value="ENSGAGG00000006327.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016323">
        <property type="term" value="C:basolateral plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008509">
        <property type="term" value="F:monoatomic anion transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008510">
        <property type="term" value="F:sodium:bicarbonate symporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005452">
        <property type="term" value="F:solute:inorganic anion antiporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051453">
        <property type="term" value="P:regulation of intracellular pH"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.570:FF:000001">
        <property type="entry name" value="Anion exchange protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.930.10:FF:000002">
        <property type="entry name" value="Anion exchange protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.570">
        <property type="entry name" value="Helical hairpin bin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.930.10">
        <property type="entry name" value="Mannitol-specific EII, Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013769">
        <property type="entry name" value="Band3_cytoplasmic_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011531">
        <property type="entry name" value="HCO3_transpt-like_TM_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003020">
        <property type="entry name" value="HCO3_transpt_euk"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003024">
        <property type="entry name" value="Na/HCO3_transpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016152">
        <property type="entry name" value="PTrfase/Anion_transptr"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00834">
        <property type="entry name" value="ae"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11453">
        <property type="entry name" value="ANION EXCHANGE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11453:SF10">
        <property type="entry name" value="ELECTROGENIC SODIUM BICARBONATE COTRANSPORTER 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07565">
        <property type="entry name" value="Band_3_cyto"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00955">
        <property type="entry name" value="HCO3_cotransp"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01231">
        <property type="entry name" value="HCO3TRNSPORT"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01232">
        <property type="entry name" value="NAHCO3TRSPRT"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55804">
        <property type="entry name" value="Phoshotransferase/anion transport protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="9 15" id="KW-0406">Ion transport</keyword>
    <keyword evidence="10 15" id="KW-0472">Membrane</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0915">Sodium</keyword>
    <keyword evidence="11" id="KW-0739">Sodium transport</keyword>
    <keyword evidence="6" id="KW-0769">Symport</keyword>
    <keyword evidence="5 15" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 15" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="3 15" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="469"/>
            <end position="492"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="504"/>
            <end position="533"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="553"/>
            <end position="571"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="583"/>
            <end position="601"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="680"/>
            <end position="698"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="718"/>
            <end position="736"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="766"/>
            <end position="785"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="15">
        <location>
            <begin position="806"/>
            <end position="827"/>
        </location>
    </feature>
    <feature type="domain" description="Band 3 cytoplasmic" evidence="18">
        <location>
            <begin position="107"/>
            <end position="387"/>
        </location>
    </feature>
    <feature type="domain" description="Bicarbonate transporter-like transmembrane" evidence="17">
        <location>
            <begin position="441"/>
            <end position="827"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="40"/>
            <end position="62"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="238"/>
            <end position="265"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="411"/>
            <end position="434"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="16">
        <location>
            <begin position="40"/>
            <end position="49"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="16">
        <location>
            <begin position="50"/>
            <end position="62"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="251"/>
            <end position="260"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004554"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010993"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022847"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023053"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023201"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00035820"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00036309"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00037277"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="RuleBase" id="RU362035"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="Pfam" id="PF00955"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF07565"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000008042.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="837" mass="94305" checksum="EC2E410675A84744" modified="2019-05-08" version="1">MEDEAVLDRGASFLKHVCDEEEVEGHHTVYIGVHVPKSYRRRRRHRRRPGLKEKKEREKISEYYSDKSDVENADETSSGILKPLISPAAERIRFILGEEDDSPAPPQLFTELDELLAVDGQEMEWKETARWIKFEEKVEQGGERWSKPHVATLSLHSLFELRTCIEKGSILLDLEATSLPQVVEMVVDNQIETGLLKLDMKDKVTYTLLRKHRHQTKKSNLRSLADIGKTVSSASRMFTNPDNGSPAMTHRNLTSTSLNDISDKPDKEQLKNKFMKKLPRDAEASNVLVGEVDFLENPFIAFVRLQQAVMLGALTEVPVPTRFLFILLGPKGKAKSYHEIGRSIATLMSDEVFHDIAYKAKDRQDLIAGIDEFLDEVIVLPPGEWDPAIRIEPPKSLPSSDKRKNMYSGGENVQMNGDTPHDAGHGGGGHGDTEELQRTGRFCGGLIKDVKRKLPVFASDFYDALNIQALSAILFIYLATVTNAITFGGLLGDATDNMQGVLESFLGTAVTGAIFCLFAGQPLTILSSTGPVLVFERLLFNFSKDHGFNYLEFRLWIGLWSAFQCLILVATDASFLVKYFTRFTEEGFSSLISFIFIYDAFKKMIKLADYYPINSHFKVDYITFYSCVCMPPEPELAGAPFRRERHQYNNTIDWSSLTKKECWKYGGYLVGNNCNYVPDITLMSFILFFGTYTCSMALKKFKTSRYFPTTARKLISDFAIILSILIFCGIDALVGVDTPKLIVPSEFKPTSPHRGWFIPPFGGNPWWVYLAAAIPALLVTILIFMDQQITAVIVNRKEHKLKVGALYLARVQLMHGGFCSFLSLPYSQVTGSLKLEP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H105</accession>
    <name>A0A452H105_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">Acetylcholine receptor subunit gamma</fullName>
        </recommendedName>
    </protein>
    <organism evidence="25 26">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="26" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="25" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="25" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">After binding acetylcholine, the AChR responds by an extensive change in conformation that affects all subunits and leads to opening of an ion-conducting channel across the plasma membrane.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>K(+)(in) = K(+)(out)</text>
            <dbReference type="Rhea" id="RHEA:29463"/>
            <dbReference type="ChEBI" id="CHEBI:29103"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>Na(+)(in) = Na(+)(out)</text>
            <dbReference type="Rhea" id="RHEA:34963"/>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
        </reaction>
    </comment>
    <comment type="subunit">
        <text evidence="2">Pentamer of two alpha chains, and one each of the beta, delta, and gamma chains.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="17">Postsynaptic cell membrane</location>
            <topology evidence="17">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="20">Belongs to the ligand-gated ion channel (TC 1.A.9) family. Acetylcholine receptor (TC 1.A.9.1) subfamily. Gamma/CHRNG sub-subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H105"/>
    <dbReference type="Ensembl" id="ENSGAGT00000009370.1">
        <property type="protein sequence ID" value="ENSGAGP00000008136.1"/>
        <property type="gene ID" value="ENSGAGG00000006424.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0022848">
        <property type="term" value="F:acetylcholine-gated monoatomic cation-selective channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004888">
        <property type="term" value="F:transmembrane signaling receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd19029">
        <property type="entry name" value="LGIC_ECD_nAChR_G"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd19064">
        <property type="entry name" value="LGIC_TM_nAChR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.390:FF:000010">
        <property type="entry name" value="Nicotinic acetylcholine receptor subunit epsilon"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.390:FF:000036">
        <property type="entry name" value="Nicotinic acetylcholine receptor subunit gamma"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.70.170.10:FF:000012">
        <property type="entry name" value="Nicotinic acetylcholine receptor subunit gamma"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.70.170.10">
        <property type="entry name" value="Neurotransmitter-gated ion-channel ligand-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.390">
        <property type="entry name" value="Neurotransmitter-gated ion-channel transmembrane domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006202">
        <property type="entry name" value="Neur_chan_lig-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036734">
        <property type="entry name" value="Neur_chan_lig-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006201">
        <property type="entry name" value="Neur_channel"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036719">
        <property type="entry name" value="Neuro-gated_channel_TM_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038050">
        <property type="entry name" value="Neuro_actylchol_rec"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006029">
        <property type="entry name" value="Neurotrans-gated_channel_TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018000">
        <property type="entry name" value="Neurotransmitter_ion_chnl_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002394">
        <property type="entry name" value="Nicotinic_acetylcholine_rcpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18945">
        <property type="entry name" value="NEUROTRANSMITTER GATED ION CHANNEL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02931">
        <property type="entry name" value="Neur_chan_LBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02932">
        <property type="entry name" value="Neur_chan_memb"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00254">
        <property type="entry name" value="NICOTINICR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00252">
        <property type="entry name" value="NRIONCHANNEL"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90112">
        <property type="entry name" value="Neurotransmitter-gated ion-channel transmembrane pore"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF63712">
        <property type="entry name" value="Nicotinic receptor ligand binding domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00236">
        <property type="entry name" value="NEUROTR_ION_CHANNEL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="16 22" id="KW-0407">Ion channel</keyword>
    <keyword evidence="9 22" id="KW-0406">Ion transport</keyword>
    <keyword evidence="15" id="KW-1071">Ligand-gated ion channel</keyword>
    <keyword evidence="10 22" id="KW-0472">Membrane</keyword>
    <keyword evidence="14" id="KW-0628">Postsynaptic cell membrane</keyword>
    <keyword evidence="12" id="KW-0675">Receptor</keyword>
    <keyword evidence="26" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6 22" id="KW-0732">Signal</keyword>
    <keyword evidence="8" id="KW-0770">Synapse</keyword>
    <keyword evidence="5 22" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 22" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="3 22" id="KW-0813">Transport</keyword>
    <feature type="signal peptide" evidence="22">
        <location>
            <begin position="1"/>
            <end position="18"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5022249596" description="Acetylcholine receptor subunit gamma" evidence="22">
        <location>
            <begin position="19"/>
            <end position="516"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="22">
        <location>
            <begin position="242"/>
            <end position="264"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="22">
        <location>
            <begin position="276"/>
            <end position="296"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="22">
        <location>
            <begin position="308"/>
            <end position="331"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="22">
        <location>
            <begin position="474"/>
            <end position="496"/>
        </location>
    </feature>
    <feature type="domain" description="Neurotransmitter-gated ion-channel ligand-binding" evidence="23">
        <location>
            <begin position="25"/>
            <end position="241"/>
        </location>
    </feature>
    <feature type="domain" description="Neurotransmitter-gated ion-channel transmembrane" evidence="24">
        <location>
            <begin position="248"/>
            <end position="491"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00003328"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00011293"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023257"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00034104"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00034430"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00036239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00038482"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00039195"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="RuleBase" id="RU000687"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="Pfam" id="PF02931"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="Pfam" id="PF02932"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000008136.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="516" mass="58947" checksum="F0FA6C85B4F4571D" modified="2019-05-08" version="1">MLALMSWLPLSLLPPLVSVLCRNQEEKLLNDLMTNYNRNLRPAQGEGDIINVTLKLTLTNLISLNEREETLTTNIWIEMKWCDYRLQWDPDEYDNIQILRVPSTEVWLPDIVLENNIDGVFEITLYCNTLLDSNGCLLWMPPAIYRTSCAIFVTYFPFDWQNCSMVFQSQTYSANEINLLLMEEDGQTIEWIVIDPEAFTENGEWAIKHRPAKKIVNAERFTPDDLGYQQVVFYLIIQRKPLFYVINIIVPCVLISSVAVLVYFLPAKAGGQKCTVSINVLLAQTVFLFLIAQKVPETSDAMPLIGKYLTFLLVVTVVIVANAVIVLNVSLRTPNTHAMSQKVRQICLRQLPHFLGMHMRRSDMAPRLLLARRRSSLGLMVKADEYMLWKARTELVFEKQKERAGLMKAVLEKIGKGLENRCTQDLCHSLVQAGPEIWACVDACNHIAKTLREQNDFDNENEEWILVGRVIDRVCFLVMASLFVLGTVSIFLMAHFNQAPAAPFPGDPKCYLPYSD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H108</accession>
    <name>A0A452H108_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Glucosamine-6-phosphate isomerase</fullName>
            <ecNumber evidence="9">3.5.99.6</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="9">Glucosamine-6-phosphate isomerase</fullName>
        </alternativeName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="8">Catalyzes the reversible conversion of alpha-D-glucosamine 6-phosphate (GlcN-6P) into beta-D-fructose 6-phosphate (Fru-6P) and ammonium ion, a regulatory reaction step in de novo uridine diphosphate-N-acetyl-alpha-D-glucosamine (UDP-GlcNAc) biosynthesis via hexosamine pathway. Deamination is coupled to aldo-keto isomerization mediating the metabolic flux from UDP-GlcNAc toward Fru-6P. At high ammonium level can drive amination and isomerization of Fru-6P toward hexosamines and UDP-GlcNAc synthesis. Has a role in fine tuning the metabolic fluctuations of cytosolic UDP-GlcNAc and their effects on hyaluronan synthesis that occur during tissue remodeling. Seems to trigger calcium oscillations in mammalian eggs. These oscillations serve as the essential trigger for egg activation and early development of the embryo.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="9">
            <text>alpha-D-glucosamine 6-phosphate + H2O = beta-D-fructose 6-phosphate + NH4(+)</text>
            <dbReference type="Rhea" id="RHEA:12172"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:28938"/>
            <dbReference type="ChEBI" id="CHEBI:57634"/>
            <dbReference type="ChEBI" id="CHEBI:75989"/>
            <dbReference type="EC" id="3.5.99.6"/>
        </reaction>
    </comment>
    <comment type="subunit">
        <text evidence="3 9">Homohexamer.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 9">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 9">Belongs to the glucosamine/galactosamine-6-phosphate isomerase family.</text>
    </comment>
    <dbReference type="EC" id="3.5.99.6" evidence="9"/>
    <dbReference type="AlphaFoldDB" id="A0A452H108"/>
    <dbReference type="Ensembl" id="ENSGAGT00000009435.1">
        <property type="protein sequence ID" value="ENSGAGP00000008194.1"/>
        <property type="gene ID" value="ENSGAGG00000006492.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004342">
        <property type="term" value="F:glucosamine-6-phosphate deaminase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042802">
        <property type="term" value="F:identical protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005975">
        <property type="term" value="P:carbohydrate metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006043">
        <property type="term" value="P:glucosamine catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006046">
        <property type="term" value="P:N-acetylglucosamine catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019262">
        <property type="term" value="P:N-acetylneuraminate catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd01399">
        <property type="entry name" value="GlcN6P_deaminase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1360:FF:000004">
        <property type="entry name" value="Glucosamine-6-phosphate isomerase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1360">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="HAMAP" id="MF_01241">
        <property type="entry name" value="GlcN6P_deamin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006148">
        <property type="entry name" value="Glc/Gal-6P_isomerase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004547">
        <property type="entry name" value="Glucosamine6P_isomerase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018321">
        <property type="entry name" value="Glucosamine6P_isomerase_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037171">
        <property type="entry name" value="NagB/RpiA_transferase-like"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00502">
        <property type="entry name" value="nagB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11280">
        <property type="entry name" value="GLUCOSAMINE-6-PHOSPHATE ISOMERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11280:SF8">
        <property type="entry name" value="GLUCOSAMINE-6-PHOSPHATE ISOMERASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01182">
        <property type="entry name" value="Glucosamine_iso"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF100950">
        <property type="entry name" value="NagB/RpiA/CoA transferase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01161">
        <property type="entry name" value="GLC_GALNAC_ISOMERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7 9" id="KW-0119">Carbohydrate metabolism</keyword>
    <keyword evidence="4 9" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6 9" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Glucosamine/galactosamine-6-phosphate isomerase" evidence="10">
        <location>
            <begin position="16"/>
            <end position="231"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005526"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00011643"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023277"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00045295"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="RuleBase" id="RU361197"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="Pfam" id="PF01182"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000008194.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="271" mass="30280" checksum="4DA7A5933D3A8A8A" modified="2019-05-08" version="1">MKLIILENYAQASEWAAKYIRNRIIQFNPGPDKYFTLGLPTGSTPLGCYKKLIEYYKNGDLSFKFVKTFNMDEYVGLPRDHLESYHSFMWNNFFKHIDISPENTHILDGNAADLQAECDAFEGKIKAAGGIELFVGGIGPDGHIAFNEPGSSLVSRTRVKTLAMDTILANARFFDGDLSKVPTMALTVGVGTVMDAREVMILITGAHKAFALYKAIEEGVNHMWTVSAFQQHPSTVFVCDEDATLELKVKTVKYFKGPHWGCKTIWRPGPL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HAP2</accession>
    <name>A0A452HAP2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Dedicator of cytokinesis 8</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="3">Belongs to the DOCK family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000013502.1">
        <property type="protein sequence ID" value="ENSGAGP00000011790.1"/>
        <property type="gene ID" value="ENSGAGG00000007768.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031252">
        <property type="term" value="C:cell leading edge"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903905">
        <property type="term" value="P:positive regulation of establishment of T cell polarity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:2000406">
        <property type="term" value="P:positive regulation of T cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007264">
        <property type="term" value="P:small GTPase-mediated signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd08696">
        <property type="entry name" value="C2_Dock-C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.410:FF:000002">
        <property type="entry name" value="Dedicator of cytokinesis protein 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.410">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037808">
        <property type="entry name" value="C2_Dock-C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027007">
        <property type="entry name" value="C2_DOCK-type_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026791">
        <property type="entry name" value="DOCK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043161">
        <property type="entry name" value="DOCK_C_lobe_A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027357">
        <property type="entry name" value="DOCKER_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046769">
        <property type="entry name" value="DOCKER_Lobe_A"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23317">
        <property type="entry name" value="DEDICATOR OF CYTOKINESIS DOCK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23317:SF74">
        <property type="entry name" value="DEDICATOR OF CYTOKINESIS PROTEIN 8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06920">
        <property type="entry name" value="DHR-2_Lobe_A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14429">
        <property type="entry name" value="DOCK-C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51650">
        <property type="entry name" value="C2_DOCK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51651">
        <property type="entry name" value="DOCKER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0344">Guanine-nucleotide releasing factor</keyword>
    <keyword evidence="1" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="C2 DOCK-type" evidence="4">
        <location>
            <begin position="420"/>
            <end position="589"/>
        </location>
    </feature>
    <feature type="domain" description="DOCKER" evidence="5">
        <location>
            <begin position="1431"/>
            <end position="1651"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022658"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00983"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS51650"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS51651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011790.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="8">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1651" mass="188192" checksum="C944A2C4C2AABF57" modified="2019-05-08" version="1">SSFYAPFCLPSYLFICTSPRDPVRFQSCVSLQQEAPRPLNVLCDETGRTTLTSCDFNLRSLQPDPRLENLLQHVSVEDFERQNEEARRANRHAELFALYPAMDEEDAVEIRPVPDCPKEHLGNRILVKLQTLKFDIEIEPLFACIALYDVKERKKISENFHCDLNSEQFKGFLRSHTPCIDSSSQARSAVFSVTYPSSDIYLVVKIEKVLQQGEIGDCAEPYMVLKESEGGKSKEKVEKLKAQAESFCQRLGKYRMPFAWAPISLTNFFNFSTLEREMSEAESLNGNRKGTTGDRRTMMLQARRLSERSLNSEDNYAVSNFKATSLTVNTFIKQEGDRLSDEDLFKFLTDYKRSSSLQRRVKSIPGSLKLEISPASEVLSCCLTPELLQVKPFAENRSRPQKEILEFPVREVYVPHTVYRNLLYVYPQRLNFANRLASARNITIKIQFMCGEDATCAMPIIFGKSSGPEFLQEMYTAVTYHNKSPDFYEEVKIKLPAKLTEKHHLLFTFYHISCQPKQGACVETLLGYSWLPILLNDRLQTGHYCLPVALDKLPLNYSMHSPEKVPSQSPPIKWVEGHKGVFNVEVQAVSSIHTQDNHLEKFFTLCHSLESQVTFPIRVMEQKITEAALEHELKLSIICLNSSRLEPLVLFLHLVLEKLFQLAVQPMVIAGQTANFSQFAFESVVAIVNSLHNSKDLSKDQHGRNCLLASYVYYVFRLPEPQRDIARLESRYYTFGRTTAVSVGSRLLQSRVMSCSNPDIHGTQTATDEEVKNIMLSKVREERKLELALQMVVSTGMVRETVFKYAWFFFELLIKSMAQYVHNMDKQDTPRRSRFSDRFKDDITTIVNVVTSEIETEQAEKINISLAFFLYDLLSLMDRGFVFNLIRHYCNQLSNKLNTLPTLISMRLEFLRILCSHEHYLNLNLFFMTSVSAPVSPSPSLSSQVGITHFQDQKIASMFDLSADYRQQHFLAGLLFTELAAALDTEAEGKAISAVHSLLSSHDLDQRCLKPEVKVKIAALYLPLVGIILDALPQLYDFTEARSGKGRTGSPDEDQESSNPINQSVAWAIAGNQFNLLRTSGAPLSSLPYKQYNTLSPDTTRNLLICFLWIMKNAEQSLIRKWIADLPSMQLNRILDLLFICVSCFEYKVCGTFHRCFILLYDLISKSRSPLCTLILLTYVALLGNDRFPGLNENLRWRKEQTQWRQANERLDKTKAELDQEALISGNLATEANLIILDMQENIIQASSAVDCKDNLLGGVLKVLVNSLSCDQSTTYLTHCFATLRALIAKFGDLLFEEEMEQCADLCQRVLHHCSSSIDITRIQACATLYLLMRHSFSSISNFARVKMQVTMSLASLVGKSSDFNEEYLRRSLRTILAYAEEDMDMQPTMFPVQVKELLRNLNSILSDTVKMREFQEDPEMLMDLMYRIAKGYQTSPDLRLTWLQNMAEKHAKKKCYTEAAMCLVHAAALVAEYLSMLEDHKYLPVGSVSFQNISSNVLEESAVSDDILSPDEDGICSGRYFSESGLVGLLEQAAELFSTGGLYETVNEVYKIVIPILEAHRDFRKLTSTHSKLQKAFDSIINKGQKRMFGTYFRVGFYGSKFGDLDEQEFVYKEPAITKLPEISHRLEGLKLRGCKFCFESFWNFCLGQQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HAP9</accession>
    <name>A0A452HAP9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">IF rod domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="3 4">Belongs to the intermediate filament family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HAP9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013532.1">
        <property type="protein sequence ID" value="ENSGAGP00000011817.1"/>
        <property type="gene ID" value="ENSGAGG00000008332.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045095">
        <property type="term" value="C:keratin filament"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030280">
        <property type="term" value="F:structural constituent of skin epidermis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045109">
        <property type="term" value="P:intermediate filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031424">
        <property type="term" value="P:keratinization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.1160:FF:000001">
        <property type="entry name" value="Keratin type II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.170:FF:000004">
        <property type="entry name" value="Keratin, type II cytoskeletal 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.500:FF:000001">
        <property type="entry name" value="Type II keratin 23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.170">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.500">
        <property type="entry name" value="Single helix bin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.1160">
        <property type="entry name" value="Vasodilator-stimulated phosphoprotein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018039">
        <property type="entry name" value="IF_conserved"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039008">
        <property type="entry name" value="IF_rod_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032444">
        <property type="entry name" value="Keratin_2_head"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003054">
        <property type="entry name" value="Keratin_II"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45616">
        <property type="entry name" value="GATA-TYPE DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45616:SF19">
        <property type="entry name" value="KERATIN 90"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00038">
        <property type="entry name" value="Filament"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16208">
        <property type="entry name" value="Keratin_2_head"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01276">
        <property type="entry name" value="TYPE2KERATIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM01391">
        <property type="entry name" value="Filament"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF64593">
        <property type="entry name" value="Intermediate filament protein, coiled coil region"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00226">
        <property type="entry name" value="IF_ROD_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51842">
        <property type="entry name" value="IF_ROD_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 5" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="1 4" id="KW-0403">Intermediate filament</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="IF rod" evidence="6">
        <location>
            <begin position="99"/>
            <end position="410"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="5">
        <location>
            <begin position="96"/>
            <end position="123"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="5">
        <location>
            <begin position="152"/>
            <end position="235"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="5">
        <location>
            <begin position="308"/>
            <end position="388"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022754"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00061646"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="RuleBase" id="RU000685"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS51842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011817.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="432" mass="48489" checksum="FFEC5004911BDFD6" modified="2019-05-08" version="1">MSTKASVVPHGSCGSKPLMPPFSYMGPACCGGYRAVRSGYGYGLGYGGAGYRIGGAGYGFSGGFAPGAGGIHEVTVNQNLLAPLNLEIDPTIQKVRKEEKEQIKTLNNKFASFIDKVRFLEQQNKVLETKWCLLQEQKTAKSNIEPMFEAYINNLRRQLESLGSERLRLDGELKNMQDTVEDFRIRYEDEVNKRTAAENEFVVLKKDVDTAYMNKVELEAKVDALTDEMNFLRALYEAELSHFQSQISDTSVVLSMDNSRNLDLDGIIAEVKAQYEDIANRSRAEAESWYQTKYEELQLTAGRHGDDLRNTKIEISEMNRMIQRLHSEIDNVKKQCASLQTAIADAEQRGELALKDAQAKLAELEEALQKTKADMARQLREYQELMNVKLALDIEIATYRKLLEGEENRLAGEGVGAVNICKYSGSQMLDTQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HAQ0</accession>
    <name>A0A452HAQ0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">PIPK domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HAQ0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013533.1">
        <property type="protein sequence ID" value="ENSGAGP00000011818.1"/>
        <property type="gene ID" value="ENSGAGG00000009108.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016308">
        <property type="term" value="F:1-phosphatidylinositol-4-phosphate 5-kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046854">
        <property type="term" value="P:phosphatidylinositol phosphate biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.800.10:FF:000001">
        <property type="entry name" value="phosphatidylinositol 4-phosphate 5-kinase type-1 gamma"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.810.10">
        <property type="entry name" value="2-Layer Sandwich"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.800.10">
        <property type="entry name" value="Phosphatidylinositol Phosphate Kinase II Beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027483">
        <property type="entry name" value="PInositol-4-P-4/5-kinase_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002498">
        <property type="entry name" value="PInositol-4-P-4/5-kinase_core"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027484">
        <property type="entry name" value="PInositol-4-P-5-kinase_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023610">
        <property type="entry name" value="PInositol-4/5-P-5/4-kinase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23086:SF34">
        <property type="entry name" value="PHOSPHATIDYLINOSITOL 4-PHOSPHATE 5-KINASE TYPE-1 BETA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23086">
        <property type="entry name" value="PHOSPHATIDYLINOSITOL-4-PHOSPHATE 5-KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01504">
        <property type="entry name" value="PIP5K"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00330">
        <property type="entry name" value="PIPKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56104">
        <property type="entry name" value="SAICAR synthase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51455">
        <property type="entry name" value="PIPK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="3" id="KW-0418">Kinase</keyword>
    <keyword evidence="3" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="PIPK" evidence="5">
        <location>
            <begin position="25"/>
            <end position="395"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00781"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS51455"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011818.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="504" mass="56687" checksum="9A44FE01B0FD5274" modified="2019-05-08" version="1">MSSVTENGDAPAGKQNEEKTYKKTASSAIKGTIQLGIGYTVGNLTSKPDRDVLMQDFYVVESVFLPSEGSSLTPAHHYPDFRFKTYAPLAFRYFRELFGIKPDDYLYSICSEPLIELSNPGASGSLFFVTSDDEFIIKTVQHKEAEFLQKLLPGYYMNLNQNPRTLLPKFYGLYCVQSGGINIRIVVMNNVLPRAMKMHFTYDLKGSTYKRRASRKEREKSNPTFKDLDFLQEMHEGLYFDSETYSALMKTLQRDCRVLESFKIMDYSLLLGINVLDSITKEKDGESSQNASDAKRHGGQKVLYSTAMESIQGPGKSGESVITEATNTMGGIPAKSHKGEKLLLFMGIIDILQSYRLMKKLEHSWKALVYDGDTVSVHRPSFYADRFLKFMSSRVFKKVQALKSSPSKRRCSSIAALKATSQEIVSAVSQEWKDEKHDMLTEGQSFSSLDEEVLGSRHHPDLVPSTPSLFEAASLATTISSSSLYVDEQYQHDGTLLYSSSNRE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HAQ3</accession>
    <name>A0A452HAQ3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Scaffolding anchor of CK1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the FAM83 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000013530.1">
        <property type="protein sequence ID" value="ENSGAGP00000011815.1"/>
        <property type="gene ID" value="ENSGAGG00000009036.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019901">
        <property type="term" value="F:protein kinase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030509">
        <property type="term" value="P:BMP signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.870.10:FF:000004">
        <property type="entry name" value="protein FAM83H isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.870.10">
        <property type="entry name" value="Endonuclease Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050944">
        <property type="entry name" value="FAM83"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012461">
        <property type="entry name" value="SACK1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16181">
        <property type="entry name" value="PROTEIN FAM83A-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16181:SF29">
        <property type="entry name" value="PROTEIN FAM83A-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07894">
        <property type="entry name" value="SACK1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56024">
        <property type="entry name" value="Phospholipase D/nuclease"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Scaffolding anchor of CK1" evidence="5">
        <location>
            <begin position="15"/>
            <end position="297"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="76"/>
            <end position="101"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="350"/>
            <end position="387"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="454"/>
            <end position="773"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="353"/>
            <end position="373"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="460"/>
            <end position="470"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="489"/>
            <end position="498"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="515"/>
            <end position="525"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="637"/>
            <end position="649"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="673"/>
            <end position="682"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006937"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="Pfam" id="PF07894"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011815.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="773" mass="85733" checksum="D7429132467F3FB8" modified="2019-05-08" version="1">MALSQVQCLDDSHVNWRSSESKPEFFYSEEQRLALEALVSEGPEAFYEVLKKENIRDFLSELELKKIMESLETYDPGSEYIPPHRGGDSDGDCTSQGGEGELAPSLEYWPQRSDRSIPQLDLGWPETVAYRGVTRAMVYMQPPIDGQAHIKEVVRKMIVQAQKVVAVVMDMFTDVDIFKDLLDAGFKRKVGVYIIVDETNAKYFLQMCERAQMHAGHLKNLRVRTTGGTEFFTHSATKFKGALAQKFMFVDGDRAMCGSYSFTWSAARTDRNLISVLSGQVVEAFDKQFQELYLMSRGLSLKSIPMDEEPEPEPFTLPSVMPVTPANAVVKRLINPKYALVKAKSADQISKMSSERQADAGRNKIMDSKDKALPEGQAAERQNDMADLAPPIHPGLLNMEKANMFDYLPTWVEPDPDPSEILGYINIIDPTIKNVKLSQMNRIKVCDVSQANAQHRQMLKSKELEAKKSQEPSPASREQREAPPTPIKEAPKPVESPKLRQVAEPANADHGPGENPTQAQATASDHQLRRQEAVSQEAQGTAAMCPVNGLEGEEDEEDYLTLTTTTPTPPPPXSNASSTSDELFEVRGRFGPLRRTNSDHIPNGGNLHLQRKMSDPHISRGTYVSPLGSLQSPPNLRLEDAVQRRDDNIRMAQCHPWTSPKNPGPSKYKADGTSAKKPTTATSPPPYWQSKGFSSARNVQPRHLSQGISRATGKPLSSLPESQRTADEVRTPLGIPLTKLSQSKHLKARVGGGQQVSTDSKRRLPGAASRKDQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HAQ4</accession>
    <name>A0A452HAQ4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Nebulette</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HAQ4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013538.1">
        <property type="protein sequence ID" value="ENSGAGP00000011823.1"/>
        <property type="gene ID" value="ENSGAGG00000009109.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005925">
        <property type="term" value="C:focal adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd09447">
        <property type="entry name" value="LIM_LASP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.110.10:FF:000087">
        <property type="entry name" value="LIM zinc-binding domain-containing Nebulette"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.40:FF:000007">
        <property type="entry name" value="nebulin isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.110.10">
        <property type="entry name" value="Cysteine Rich Protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.40">
        <property type="entry name" value="SH3 Domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051759">
        <property type="entry name" value="LIM-SH3_domain_protein"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000900">
        <property type="entry name" value="Nebulin_repeat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036028">
        <property type="entry name" value="SH3-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001452">
        <property type="entry name" value="SH3_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001781">
        <property type="entry name" value="Znf_LIM"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46218">
        <property type="entry name" value="LASP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46218:SF3">
        <property type="entry name" value="NEBULETTE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00412">
        <property type="entry name" value="LIM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00880">
        <property type="entry name" value="Nebulin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14604">
        <property type="entry name" value="SH3_9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00452">
        <property type="entry name" value="SH3DOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00132">
        <property type="entry name" value="LIM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00227">
        <property type="entry name" value="NEBU"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00326">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57716">
        <property type="entry name" value="Glucocorticoid receptor-like (DNA-binding domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50044">
        <property type="entry name" value="SH3-domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00478">
        <property type="entry name" value="LIM_DOMAIN_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50023">
        <property type="entry name" value="LIM_DOMAIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51216">
        <property type="entry name" value="NEBULIN"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50002">
        <property type="entry name" value="SH3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5 6" id="KW-0440">LIM domain</keyword>
    <keyword evidence="2 6" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="1 7" id="KW-0728">SH3 domain</keyword>
    <keyword evidence="4 6" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="LIM zinc-binding" evidence="9">
        <location>
            <begin position="3"/>
            <end position="63"/>
        </location>
    </feature>
    <feature type="domain" description="SH3" evidence="8">
        <location>
            <begin position="213"/>
            <end position="273"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022443"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023038"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011823.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="12">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="273" mass="31443" checksum="A3275A1F54B23F75" modified="2019-05-08" version="1">MNPQCARCGKVVYPTEKVNCLDKYWHKGCFHCEVCKMALNMNNYKGYEKKPYCNAHYPKQSFTTVADTPENLRLKQQSELQSQVKYKRDFEESKGRGFSIVTDTPELQRLKRTQEQISNVKYHEDFEKTKGRGFTPVVDDPVTERVRKNTQVVSDAAYKGVHPHIVEMDRRPGIIVAPVLPGAYQQSQSQGYGYMHQTSMSSMRSMHSQPHSAGLRTYRAMYDYSAQDEDEVSFRDGDYIINVQPIDDGWMFGTVQRTGKTGMLPANYIEYVN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HAQ7</accession>
    <name>A0A452HAQ7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">PIPK domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HAQ7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013537.1">
        <property type="protein sequence ID" value="ENSGAGP00000011822.1"/>
        <property type="gene ID" value="ENSGAGG00000009108.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016308">
        <property type="term" value="F:1-phosphatidylinositol-4-phosphate 5-kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046854">
        <property type="term" value="P:phosphatidylinositol phosphate biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.800.10:FF:000001">
        <property type="entry name" value="phosphatidylinositol 4-phosphate 5-kinase type-1 gamma"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.810.10">
        <property type="entry name" value="2-Layer Sandwich"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.800.10">
        <property type="entry name" value="Phosphatidylinositol Phosphate Kinase II Beta"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027483">
        <property type="entry name" value="PInositol-4-P-4/5-kinase_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002498">
        <property type="entry name" value="PInositol-4-P-4/5-kinase_core"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027484">
        <property type="entry name" value="PInositol-4-P-5-kinase_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023610">
        <property type="entry name" value="PInositol-4/5-P-5/4-kinase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23086:SF34">
        <property type="entry name" value="PHOSPHATIDYLINOSITOL 4-PHOSPHATE 5-KINASE TYPE-1 BETA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23086">
        <property type="entry name" value="PHOSPHATIDYLINOSITOL-4-PHOSPHATE 5-KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01504">
        <property type="entry name" value="PIP5K"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00330">
        <property type="entry name" value="PIPKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56104">
        <property type="entry name" value="SAICAR synthase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51455">
        <property type="entry name" value="PIPK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="2" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="3" id="KW-0418">Kinase</keyword>
    <keyword evidence="3" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="PIPK" evidence="5">
        <location>
            <begin position="25"/>
            <end position="395"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00781"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS51455"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011822.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="540" mass="60540" checksum="01258EB041BCEEF4" modified="2019-05-08" version="1">MSSVTENGDAPAGKQNEEKTYKKTASSAIKGTIQLGIGYTVGNLTSKPDRDVLMQDFYVVESVFLPSEGSSLTPAHHYPDFRFKTYAPLAFRYFRELFGIKPDDYLYSICSEPLIELSNPGASGSLFFVTSDDEFIIKTVQHKEAEFLQKLLPGYYMNLNQNPRTLLPKFYGLYCVQSGGINIRIVVMNNVLPRAMKMHFTYDLKGSTYKRRASRKEREKSNPTFKDLDFLQEMHEGLYFDSETYSALMKTLQRDCRVLESFKIMDYSLLLGINVLDSITKEKDGESSQNASDAKRHGGQKVLYSTAMESIQGPGKSGESVITEATNTMGGIPAKSHKGEKLLLFMGIIDILQSYRLMKKLEHSWKALVYDGDTVSVHRPSFYADRFLKFMSSRVFKKVQALKSSPSKRRCSSIAALKATSQEIVSAVSQEWKDEKHDMLTEGQSFSSLDEEVLGSRHHPDLVPSTPSLFEAASLATTISSSSLYVDEQYQHDGTLLYSSSKGLPSSSTFTLEDSAIYLTSEQSTLEMENDNASVLDVYL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HB11</accession>
    <name>A0A452HB11_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Rho GTPase-activating protein 26</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="10">Rho-type GTPase-activating protein 26</fullName>
        </alternativeName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell junction</location>
            <location evidence="2">Focal adhesion</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Endosome membrane</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HB11"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013674.1">
        <property type="protein sequence ID" value="ENSGAGP00000011936.1"/>
        <property type="gene ID" value="ENSGAGG00000004156.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010008">
        <property type="term" value="C:endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005925">
        <property type="term" value="C:focal adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd01249">
        <property type="entry name" value="BAR-PH_GRAF_family"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd07636">
        <property type="entry name" value="BAR_GRAF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.555.10:FF:000006">
        <property type="entry name" value="Rho GTPase activating protein 26"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000116">
        <property type="entry name" value="Rho GTPase activating protein 26"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1270.60:FF:000001">
        <property type="entry name" value="Rho GTPase-activating protein 26"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.60">
        <property type="entry name" value="Arfaptin homology (AH) domain/BAR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.555.10">
        <property type="entry name" value="Rho GTPase activation protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027267">
        <property type="entry name" value="AH/BAR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004148">
        <property type="entry name" value="BAR_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035483">
        <property type="entry name" value="GRAF_BAR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047234">
        <property type="entry name" value="GRAF_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047225">
        <property type="entry name" value="PH_GRAF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000198">
        <property type="entry name" value="RhoGAP_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12552">
        <property type="entry name" value="OLIGOPHRENIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12552:SF4">
        <property type="entry name" value="RHO GTPASE-ACTIVATING PROTEIN 26"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16746">
        <property type="entry name" value="BAR_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00169">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00620">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00233">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00324">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103657">
        <property type="entry name" value="BAR/IMD domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50238">
        <property type="entry name" value="RHOGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0965">Cell junction</keyword>
    <keyword evidence="9" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="6" id="KW-0967">Endosome</keyword>
    <keyword evidence="5" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="8" id="KW-0472">Membrane</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="PH" evidence="11">
        <location>
            <begin position="265"/>
            <end position="369"/>
        </location>
    </feature>
    <feature type="domain" description="Rho-GAP" evidence="12">
        <location>
            <begin position="375"/>
            <end position="560"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004608"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00020525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00032211"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50238"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011936.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="717" mass="82200" checksum="4A990B2C7C08A486" modified="2019-05-08" version="1">MGLPALEFSDCCLDSPQFRERLKSHEAELDKTNKFIKELIKDGKSLVAALKNLSSAKRKFADSLNEFKFRCIGDAETDDEICIAKSLQEFATVLRNLEDERMRMIENAGEVLITPLEKFRKEQIGAAKDAKKKYDKETEKYCGVLEKHLNLSSKKKESQLQEADSQVDLVRQHFYEVSLEYVFKVQEVQERKMFEFVEPLLAFLQGLFTFYHHGYELAKDFSDFKTELTISIQNTRNRFEGTRSEVESLMRKMKENPEEHRNISPFTMEGYLYVQEKRHFGTSWVKHYCTYQRESKRITMVPFDQKSGGKGGEDEAVTLKSCIRRKTDSIEKRFCFDVEAVDRPGVITMQALSEEDRKLWMEAMDGREPVRRDTAQLDNIGFSIIRKCINAVETRGINEQGLYRIVGVNSRVQKLLSILMDPKTASETETDICAEWEIKTITSALKTYLRMLPGPLMMYQFQRSFIKAAKLENQESRISEIHNLVHRLPEKNRQMLQMLMNHLANVANNHKQNLMTVANLGVVFGPTLLRPQEETVAAIMDIKFQNIVVEILIENHEKIFNTVPETLPPNSQLLLLSHKKNTDSKPPSCSERPLTLFHTAQANENQETRNSIINSSLESVISSNVNSFLHSNSTLRPSLNSSDPDLEVIKPSRPNSLSADTPFSLHLPTSSTLSVTYTQVVEDHLEISWSGIHNAISHAATETICFKPSHKKPWLSE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HB15</accession>
    <name>A0A452HB15_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Rho GTPase-activating protein 26</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="10">Rho-type GTPase-activating protein 26</fullName>
        </alternativeName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell junction</location>
            <location evidence="2">Focal adhesion</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Endosome membrane</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HB15"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013666.1">
        <property type="protein sequence ID" value="ENSGAGP00000011929.1"/>
        <property type="gene ID" value="ENSGAGG00000004156.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010008">
        <property type="term" value="C:endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005925">
        <property type="term" value="C:focal adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd01249">
        <property type="entry name" value="BAR-PH_GRAF_family"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd07636">
        <property type="entry name" value="BAR_GRAF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.555.10:FF:000006">
        <property type="entry name" value="Rho GTPase activating protein 26"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000116">
        <property type="entry name" value="Rho GTPase activating protein 26"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1270.60:FF:000001">
        <property type="entry name" value="Rho GTPase-activating protein 26"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.60">
        <property type="entry name" value="Arfaptin homology (AH) domain/BAR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.555.10">
        <property type="entry name" value="Rho GTPase activation protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027267">
        <property type="entry name" value="AH/BAR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004148">
        <property type="entry name" value="BAR_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035483">
        <property type="entry name" value="GRAF_BAR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047234">
        <property type="entry name" value="GRAF_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047225">
        <property type="entry name" value="PH_GRAF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000198">
        <property type="entry name" value="RhoGAP_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12552">
        <property type="entry name" value="OLIGOPHRENIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12552:SF4">
        <property type="entry name" value="RHO GTPASE-ACTIVATING PROTEIN 26"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16746">
        <property type="entry name" value="BAR_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00169">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00620">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00233">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00324">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103657">
        <property type="entry name" value="BAR/IMD domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50238">
        <property type="entry name" value="RHOGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0965">Cell junction</keyword>
    <keyword evidence="9" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="6" id="KW-0967">Endosome</keyword>
    <keyword evidence="5" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="8" id="KW-0472">Membrane</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="PH" evidence="12">
        <location>
            <begin position="265"/>
            <end position="369"/>
        </location>
    </feature>
    <feature type="domain" description="Rho-GAP" evidence="13">
        <location>
            <begin position="375"/>
            <end position="560"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="634"/>
            <end position="669"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="634"/>
            <end position="643"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="11">
        <location>
            <begin position="660"/>
            <end position="669"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004608"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00020525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00032211"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50238"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011929.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="669" mass="76954" checksum="849E7FE15D0FE7E7" modified="2019-05-08" version="1">MGLPALEFSDCCLDSPQFRERLKSHEAELDKTNKFIKELIKDGKSLVAALKNLSSAKRKFADSLNEFKFRCIGDAETDDEICIAKSLQEFATVLRNLEDERMRMIENAGEVLITPLEKFRKEQIGAAKDAKKKYDKETEKYCGVLEKHLNLSSKKKESQLQEADSQVDLVRQHFYEVSLEYVFKVQEVQERKMFEFVEPLLAFLQGLFTFYHHGYELAKDFSDFKTELTISIQNTRNRFEGTRSEVESLMRKMKENPEEHRNISPFTMEGYLYVQEKRHFGTSWVKHYCTYQRESKRITMVPFDQKSGGKGGEDEAVTLKSCIRRKTDSIEKRFCFDVEAVDRPGVITMQALSEEDRKLWMEAMDGREPVRRDTAQLDNIGFSIIRKCINAVETRGINEQGLYRIVGVNSRVQKLLSILMDPKTASETETDICAEWEIKTITSALKTYLRMLPGPLMMYQFQRSFIKAAKLENQESRISEIHNLVHRLPEKNRQMLQMLMNHLANVANNHKQNLMTVANLGVVFGPTLLRPQEETVAAIMDIKFQNIVVEILIENHEKIFNTVPETLPPNSQLLLLSHKKNTDSKPPSCSERPLTLFHTAQANENQETRNSIINSSLESVISSNVNSFLHSNSTLRPSLNSSDPDLEVIKPSRPNSFTPTPWPTKPKMA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HB17</accession>
    <name>A0A452HB17_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Peptide-methionine (R)-S-oxide reductase</fullName>
            <ecNumber evidence="4">1.8.4.12</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="4">Methionine-sulfoxide reductase that specifically reduces methionine (R)-sulfoxide back to methionine. While in many cases methionine oxidation is the result of random oxidation following oxidative stress, methionine oxidation is also a post-translational modification that takes place on specific residues.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="4">
            <text>L-methionyl-[protein] + [thioredoxin]-disulfide + H2O = L-methionyl-(R)-S-oxide-[protein] + [thioredoxin]-dithiol</text>
            <dbReference type="Rhea" id="RHEA:24164"/>
            <dbReference type="Rhea" id="RHEA-COMP:10698"/>
            <dbReference type="Rhea" id="RHEA-COMP:10700"/>
            <dbReference type="Rhea" id="RHEA-COMP:12313"/>
            <dbReference type="Rhea" id="RHEA-COMP:12314"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:16044"/>
            <dbReference type="ChEBI" id="CHEBI:29950"/>
            <dbReference type="ChEBI" id="CHEBI:45764"/>
            <dbReference type="ChEBI" id="CHEBI:50058"/>
            <dbReference type="EC" id="1.8.4.12"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="3">
            <text>[thioredoxin]-disulfide + L-methionine + H2O = L-methionine (R)-S-oxide + [thioredoxin]-dithiol</text>
            <dbReference type="Rhea" id="RHEA:21260"/>
            <dbReference type="Rhea" id="RHEA-COMP:10698"/>
            <dbReference type="Rhea" id="RHEA-COMP:10700"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:29950"/>
            <dbReference type="ChEBI" id="CHEBI:50058"/>
            <dbReference type="ChEBI" id="CHEBI:57844"/>
            <dbReference type="ChEBI" id="CHEBI:58773"/>
            <dbReference type="EC" id="1.8.4.14"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="4">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
        <text evidence="4">Binds 1 zinc ion per subunit.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1 4">Belongs to the MsrB Met sulfoxide reductase family.</text>
    </comment>
    <dbReference type="EC" id="1.8.4.12" evidence="4"/>
    <dbReference type="AlphaFoldDB" id="A0A452HB17"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013679.1">
        <property type="protein sequence ID" value="ENSGAGP00000011941.1"/>
        <property type="gene ID" value="ENSGAGG00000009203.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033745">
        <property type="term" value="F:L-methionine-(R)-S-oxide reductase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033743">
        <property type="term" value="F:peptide-methionine (R)-S-oxide reductase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030091">
        <property type="term" value="P:protein repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006979">
        <property type="term" value="P:response to oxidative stress"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="2.170.150.20:FF:000017">
        <property type="entry name" value="Uncharacterized protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.170.150.20">
        <property type="entry name" value="Peptide methionine sulfoxide reductase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028427">
        <property type="entry name" value="Met_Sox_Rdtase_MsrB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002579">
        <property type="entry name" value="Met_Sox_Rdtase_MsrB_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011057">
        <property type="entry name" value="Mss4-like_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00357">
        <property type="entry name" value="peptide-methionine (R)-S-oxide reductase MsrB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10173">
        <property type="entry name" value="METHIONINE SULFOXIDE REDUCTASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10173:SF56">
        <property type="entry name" value="METHIONINE-R-SULFOXIDE REDUCTASE B3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01641">
        <property type="entry name" value="SelR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51316">
        <property type="entry name" value="Mss4-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51790">
        <property type="entry name" value="MSRB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="2 4" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="MsrB" evidence="5">
        <location>
            <begin position="40"/>
            <end position="130"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007174"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00049261"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="RuleBase" id="RU365044"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS51790"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011941.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="172" mass="19099" checksum="DB2E64266EB459F9" modified="2019-05-08" version="1">MSAFNLLHLVTKSQPVALRACGLPSGSCRDKKNCKVVFSQQDLRKRLTPLQYHVTQEKGTESAFEGEYTHHKAYGIYKCVVCGTPLFKSETKFDSNSGWPSFYDVISPDAITFTDDFSYGMHRIETGCSQVSVQHTETILIVKAANIVIASYPLTVIHKAFSSLNTNNLNVA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HB19</accession>
    <name>A0A452HB19_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Peptide-methionine (R)-S-oxide reductase</fullName>
            <ecNumber evidence="7">1.8.4.12</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="7">Methionine-sulfoxide reductase that specifically reduces methionine (R)-sulfoxide back to methionine. While in many cases methionine oxidation is the result of random oxidation following oxidative stress, methionine oxidation is also a post-translational modification that takes place on specific residues.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="7">
            <text>L-methionyl-[protein] + [thioredoxin]-disulfide + H2O = L-methionyl-(R)-S-oxide-[protein] + [thioredoxin]-dithiol</text>
            <dbReference type="Rhea" id="RHEA:24164"/>
            <dbReference type="Rhea" id="RHEA-COMP:10698"/>
            <dbReference type="Rhea" id="RHEA-COMP:10700"/>
            <dbReference type="Rhea" id="RHEA-COMP:12313"/>
            <dbReference type="Rhea" id="RHEA-COMP:12314"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:16044"/>
            <dbReference type="ChEBI" id="CHEBI:29950"/>
            <dbReference type="ChEBI" id="CHEBI:45764"/>
            <dbReference type="ChEBI" id="CHEBI:50058"/>
            <dbReference type="EC" id="1.8.4.12"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="6">
            <text>[thioredoxin]-disulfide + L-methionine + H2O = L-methionine (R)-S-oxide + [thioredoxin]-dithiol</text>
            <dbReference type="Rhea" id="RHEA:21260"/>
            <dbReference type="Rhea" id="RHEA-COMP:10698"/>
            <dbReference type="Rhea" id="RHEA-COMP:10700"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:29950"/>
            <dbReference type="ChEBI" id="CHEBI:50058"/>
            <dbReference type="ChEBI" id="CHEBI:57844"/>
            <dbReference type="ChEBI" id="CHEBI:58773"/>
            <dbReference type="EC" id="1.8.4.14"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="7">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
        <text evidence="7">Binds 1 zinc ion per subunit.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1 7">Belongs to the MsrB Met sulfoxide reductase family.</text>
    </comment>
    <dbReference type="EC" id="1.8.4.12" evidence="7"/>
    <dbReference type="AlphaFoldDB" id="A0A452HB19"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013678.1">
        <property type="protein sequence ID" value="ENSGAGP00000011940.1"/>
        <property type="gene ID" value="ENSGAGG00000009203.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033745">
        <property type="term" value="F:L-methionine-(R)-S-oxide reductase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033743">
        <property type="term" value="F:peptide-methionine (R)-S-oxide reductase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030091">
        <property type="term" value="P:protein repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006979">
        <property type="term" value="P:response to oxidative stress"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="2.170.150.20:FF:000004">
        <property type="entry name" value="Peptide-methionine (R)-S-oxide reductase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.170.150.20">
        <property type="entry name" value="Peptide methionine sulfoxide reductase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="HAMAP" id="MF_01400">
        <property type="entry name" value="MsrB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028427">
        <property type="entry name" value="Met_Sox_Rdtase_MsrB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002579">
        <property type="entry name" value="Met_Sox_Rdtase_MsrB_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011057">
        <property type="entry name" value="Mss4-like_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00357">
        <property type="entry name" value="peptide-methionine (R)-S-oxide reductase MsrB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10173">
        <property type="entry name" value="METHIONINE SULFOXIDE REDUCTASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10173:SF56">
        <property type="entry name" value="METHIONINE-R-SULFOXIDE REDUCTASE B3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01641">
        <property type="entry name" value="SelR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51316">
        <property type="entry name" value="Mss4-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51790">
        <property type="entry name" value="MSRB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="4 7" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 7" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="MsrB" evidence="9">
        <location>
            <begin position="52"/>
            <end position="174"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="173"/>
            <end position="196"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="184"/>
            <end position="196"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007174"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00048488"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00049261"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU365044"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51790"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011940.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="196" mass="21865" checksum="B14060DAE5705B4F" modified="2019-05-08" version="1">MAAIGRRTGCLVLQLMKQNRKTFYDQLREHNKTAALLGSCRDKKNCKVVFSQQDLRKRLTPLQYHVTQEKGTESAFEGEYTHHKAYGIYKCVVCGTPLFKSETKFDSNSGWPSFYDVISPDAITFTDDFSYGMHRIETGCSQCGAHLGHIFDDGPRPTGKRYCINSASLSFEPVDKNNAGEGSGSTSPAQTDKTEL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HB20</accession>
    <name>A0A452HB20_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Peptide-methionine (R)-S-oxide reductase</fullName>
            <ecNumber evidence="7">1.8.4.12</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="7">Methionine-sulfoxide reductase that specifically reduces methionine (R)-sulfoxide back to methionine. While in many cases methionine oxidation is the result of random oxidation following oxidative stress, methionine oxidation is also a post-translational modification that takes place on specific residues.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="7">
            <text>L-methionyl-[protein] + [thioredoxin]-disulfide + H2O = L-methionyl-(R)-S-oxide-[protein] + [thioredoxin]-dithiol</text>
            <dbReference type="Rhea" id="RHEA:24164"/>
            <dbReference type="Rhea" id="RHEA-COMP:10698"/>
            <dbReference type="Rhea" id="RHEA-COMP:10700"/>
            <dbReference type="Rhea" id="RHEA-COMP:12313"/>
            <dbReference type="Rhea" id="RHEA-COMP:12314"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:16044"/>
            <dbReference type="ChEBI" id="CHEBI:29950"/>
            <dbReference type="ChEBI" id="CHEBI:45764"/>
            <dbReference type="ChEBI" id="CHEBI:50058"/>
            <dbReference type="EC" id="1.8.4.12"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="6">
            <text>[thioredoxin]-disulfide + L-methionine + H2O = L-methionine (R)-S-oxide + [thioredoxin]-dithiol</text>
            <dbReference type="Rhea" id="RHEA:21260"/>
            <dbReference type="Rhea" id="RHEA-COMP:10698"/>
            <dbReference type="Rhea" id="RHEA-COMP:10700"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:29950"/>
            <dbReference type="ChEBI" id="CHEBI:50058"/>
            <dbReference type="ChEBI" id="CHEBI:57844"/>
            <dbReference type="ChEBI" id="CHEBI:58773"/>
            <dbReference type="EC" id="1.8.4.14"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="7">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
        <text evidence="7">Binds 1 zinc ion per subunit.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1 7">Belongs to the MsrB Met sulfoxide reductase family.</text>
    </comment>
    <dbReference type="EC" id="1.8.4.12" evidence="7"/>
    <dbReference type="AlphaFoldDB" id="A0A452HB20"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013675.1">
        <property type="protein sequence ID" value="ENSGAGP00000011937.1"/>
        <property type="gene ID" value="ENSGAGG00000009203.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033745">
        <property type="term" value="F:L-methionine-(R)-S-oxide reductase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033743">
        <property type="term" value="F:peptide-methionine (R)-S-oxide reductase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030091">
        <property type="term" value="P:protein repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006979">
        <property type="term" value="P:response to oxidative stress"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="2.170.150.20:FF:000004">
        <property type="entry name" value="Peptide-methionine (R)-S-oxide reductase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.170.150.20">
        <property type="entry name" value="Peptide methionine sulfoxide reductase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="HAMAP" id="MF_01400">
        <property type="entry name" value="MsrB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028427">
        <property type="entry name" value="Met_Sox_Rdtase_MsrB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002579">
        <property type="entry name" value="Met_Sox_Rdtase_MsrB_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011057">
        <property type="entry name" value="Mss4-like_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00357">
        <property type="entry name" value="peptide-methionine (R)-S-oxide reductase MsrB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10173">
        <property type="entry name" value="METHIONINE SULFOXIDE REDUCTASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10173:SF56">
        <property type="entry name" value="METHIONINE-R-SULFOXIDE REDUCTASE B3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01641">
        <property type="entry name" value="SelR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51316">
        <property type="entry name" value="Mss4-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51790">
        <property type="entry name" value="MSRB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 7" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="4 7" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
    <keyword evidence="3 7" id="KW-0862">Zinc</keyword>
    <feature type="signal peptide" evidence="7">
        <location>
            <begin position="1"/>
            <end position="21"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5018817576" description="Peptide-methionine (R)-S-oxide reductase" evidence="7">
        <location>
            <begin position="22"/>
            <end position="177"/>
        </location>
    </feature>
    <feature type="domain" description="MsrB" evidence="9">
        <location>
            <begin position="33"/>
            <end position="155"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="154"/>
            <end position="177"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="165"/>
            <end position="177"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007174"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00048488"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00049261"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU365044"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51790"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011937.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="177" mass="19356" checksum="6E22E90AB51A3EAC" modified="2019-05-08" version="1">MRALPLLSLCLAAAGIPAGSCRDKKNCKVVFSQQDLRKRLTPLQYHVTQEKGTESAFEGEYTHHKAYGIYKCVVCGTPLFKSETKFDSNSGWPSFYDVISPDAITFTDDFSYGMHRIETGCSQCGAHLGHIFDDGPRPTGKRYCINSASLSFEPVDKNNAGEGSGSTSPAQTDKTEL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HB22</accession>
    <name>A0A452HB22_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Peptide-methionine (R)-S-oxide reductase</fullName>
            <ecNumber evidence="4">1.8.4.12</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="4">Methionine-sulfoxide reductase that specifically reduces methionine (R)-sulfoxide back to methionine. While in many cases methionine oxidation is the result of random oxidation following oxidative stress, methionine oxidation is also a post-translational modification that takes place on specific residues.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="4">
            <text>L-methionyl-[protein] + [thioredoxin]-disulfide + H2O = L-methionyl-(R)-S-oxide-[protein] + [thioredoxin]-dithiol</text>
            <dbReference type="Rhea" id="RHEA:24164"/>
            <dbReference type="Rhea" id="RHEA-COMP:10698"/>
            <dbReference type="Rhea" id="RHEA-COMP:10700"/>
            <dbReference type="Rhea" id="RHEA-COMP:12313"/>
            <dbReference type="Rhea" id="RHEA-COMP:12314"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:16044"/>
            <dbReference type="ChEBI" id="CHEBI:29950"/>
            <dbReference type="ChEBI" id="CHEBI:45764"/>
            <dbReference type="ChEBI" id="CHEBI:50058"/>
            <dbReference type="EC" id="1.8.4.12"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="3">
            <text>[thioredoxin]-disulfide + L-methionine + H2O = L-methionine (R)-S-oxide + [thioredoxin]-dithiol</text>
            <dbReference type="Rhea" id="RHEA:21260"/>
            <dbReference type="Rhea" id="RHEA-COMP:10698"/>
            <dbReference type="Rhea" id="RHEA-COMP:10700"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:29950"/>
            <dbReference type="ChEBI" id="CHEBI:50058"/>
            <dbReference type="ChEBI" id="CHEBI:57844"/>
            <dbReference type="ChEBI" id="CHEBI:58773"/>
            <dbReference type="EC" id="1.8.4.14"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="4">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
        <text evidence="4">Binds 1 zinc ion per subunit.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1 4">Belongs to the MsrB Met sulfoxide reductase family.</text>
    </comment>
    <dbReference type="EC" id="1.8.4.12" evidence="4"/>
    <dbReference type="AlphaFoldDB" id="A0A452HB22"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013677.1">
        <property type="protein sequence ID" value="ENSGAGP00000011939.1"/>
        <property type="gene ID" value="ENSGAGG00000009203.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033745">
        <property type="term" value="F:L-methionine-(R)-S-oxide reductase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033743">
        <property type="term" value="F:peptide-methionine (R)-S-oxide reductase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030091">
        <property type="term" value="P:protein repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006979">
        <property type="term" value="P:response to oxidative stress"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="2.170.150.20:FF:000017">
        <property type="entry name" value="Uncharacterized protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.170.150.20">
        <property type="entry name" value="Peptide methionine sulfoxide reductase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028427">
        <property type="entry name" value="Met_Sox_Rdtase_MsrB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002579">
        <property type="entry name" value="Met_Sox_Rdtase_MsrB_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011057">
        <property type="entry name" value="Mss4-like_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR00357">
        <property type="entry name" value="peptide-methionine (R)-S-oxide reductase MsrB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10173">
        <property type="entry name" value="METHIONINE SULFOXIDE REDUCTASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10173:SF56">
        <property type="entry name" value="METHIONINE-R-SULFOXIDE REDUCTASE B3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01641">
        <property type="entry name" value="SelR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF51316">
        <property type="entry name" value="Mss4-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51790">
        <property type="entry name" value="MSRB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="2 4" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="5" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="4" id="KW-0862">Zinc</keyword>
    <feature type="transmembrane region" description="Helical" evidence="5">
        <location>
            <begin position="37"/>
            <end position="58"/>
        </location>
    </feature>
    <feature type="domain" description="MsrB" evidence="6">
        <location>
            <begin position="161"/>
            <end position="251"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007174"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00049261"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="RuleBase" id="RU365044"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS51790"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011939.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="293" mass="31511" checksum="47D005A109059034" modified="2019-05-08" version="1">MVGSPPNNTQGAVCSGERSITPGSPALRLGWCPAGRGWPLLALLCSMLIFGISCRCLGISRHASRGGRACHSSPGGGKGGAAGAAPCALCRSSRSAWRPLAYQQLLPLFFASDFVGEDITVMSAFNLLHLVTKSQPVALRACGLPSGSCRDKKNCKVVFSQQDLRKRLTPLQYHVTQEKGTESAFEGEYTHHKAYGIYKCVVCGTPLFKSETKFDSNSGWPSFYDVISPDAITFTDDFSYGMHRIETGCSQVSVQHTETILIVKAANIVIASYPLTVIHKAFSSLNTNNLNVA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HB23</accession>
    <name>A0A452HB23_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Centrosomal protein of 162 kDa</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Microtubule organizing center</location>
            <location evidence="1">Centrosome</location>
            <location evidence="1">Centriole</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the CEP162 family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000013687.1">
        <property type="protein sequence ID" value="ENSGAGP00000011949.1"/>
        <property type="gene ID" value="ENSGAGG00000009206.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005879">
        <property type="term" value="C:axonemal microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034451">
        <property type="term" value="C:centriolar satellite"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005814">
        <property type="term" value="C:centriole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005654">
        <property type="term" value="C:nucleoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060271">
        <property type="term" value="P:cilium assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038774">
        <property type="entry name" value="CEP162-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR34031">
        <property type="entry name" value="CENTROSOMAL PROTEIN OF 162 KDA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR34031:SF1">
        <property type="entry name" value="CENTROSOMAL PROTEIN OF 162 KDA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0970">Cilium biogenesis/degradation</keyword>
    <keyword evidence="7 9" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="5" id="KW-0493">Microtubule</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="21"/>
            <end position="126"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="187"/>
            <end position="227"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="495"/>
            <end position="563"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1101"/>
            <end position="1147"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="631"/>
            <end position="665"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="707"/>
            <end position="843"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="883"/>
            <end position="910"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="936"/>
            <end position="1049"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="1162"/>
            <end position="1189"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="9">
        <location>
            <begin position="1239"/>
            <end position="1297"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="78"/>
            <end position="94"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="97"/>
            <end position="111"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="216"/>
            <end position="227"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="10">
        <location>
            <begin position="495"/>
            <end position="508"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="535"/>
            <end position="545"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="553"/>
            <end position="562"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="1111"/>
            <end position="1129"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004114"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009485"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00021406"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022794"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011949.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1330" mass="152795" checksum="F85040DE075E9D63" modified="2019-05-08" version="1">MAHRFSKAELDEQFEQFLKESLSDDSFENSKTKSTILETLGQPRRKELKKKDPVPWWISKDDSDDGGMLGTNGSFLKSQKDSHPMVEVDEEAHTGKMQLQRSDGVSVSLSRDSLEPDDSVVASGPKQSILGAGLDTLEEQEEKEQFFAKLEKGGSYTIDYSRLNKELDSNDSVLLAAFACNEQNVEQVEDESKHEEKCGNYSEDFEDETESNYPSKNEDNEVEQTVRSDADLSIQGQDEKTGMLAKVVLLDSQDSTMETQKVIEQQDAAVTEHQTPEVTKDEINGTAGISYGQTNSDIEALQQAYCHIDQSLGDTDEQRINLSAMENAEFPVQDASQNNEDYAKNISTTESDLLTVEELMKPIGADSSHARGFDLEPVSPVKLTDSRAAEFMSHLPFKEHKNDAVLEKQSLDFLFEKHSKEEESVFLQATSNEGSHQKGTDKDQIDKEQLNLLRQKILQDSALLHQDSKINKACQSSSWRNTGLDSVINKQIMHKSTRSTAPLHKKKSPIGSHILVRSSGYGKPTSPLKQLSTTSEKRASKEALKKPGLKAKSPPDKARQKETLFTTRVKRSAINESASKKANNMVTSSQSVVNALGQQIKEMDSCVQSQNDSSLFPVKNCERELYLLKRVKEAEEKLKAAHDLIQQLKDLFSQKEKEMENKMVEMKMQHDKEVFRLSQENYVLQTKVNSMEELSKEKKWLHLGATEAVTEEKLKQIQKEIQDQENLLRGYQQENERLYKQMKELQIQNKKNEERMFKENQHLTSELASLREQMDKNNFLSRLEQDSKSTRNQSFTELISELRVAQKEEAKLLEEIKRLKQDKQALEVDLGQMRRERDLARAQIVSTSGEKSYEIKIIEESYKQEISLLKKRLQWYAENQELLDRDAVRLKDAKEEIEKLKLQIKEMETILKRRHPNSLPALIYAAAAASEGGGDISAKSNTIDFLERRIKKLETELEGKDDEAKKSLRAMEQQFQKIKLQYEQRLVELEELLAYKLMNEPQKLHDSKASFTAIEQELRSLKEDHQITVKNLQTEIKNLRNENSQLELKKCERSDEDVQTIEYQVEQAHAKARLVRLNQELITKSREIQDLTKTVERLQKERRTMLSNHNSSDKTDSKEKSTEILKKDILASSKRNASTTEPFPDTLDEKIYQPHNFADSHISEVLQENARLKDELERLSLEMSQQRVKSQAALAHSESNIRRTEEDTAEYIAALRASHQREVEKILSQHAVEHSASKVAELNSKISRQEILIKHLQEQVNKLHREQEALAVSQTREEILQKEMTKLLEELREAKERQSPEMKHFLCLERKIKNMEIRHVQREQELQQVK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HB28</accession>
    <name>A0A452HB28_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Rho GTPase-activating protein 26</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="10">Rho-type GTPase-activating protein 26</fullName>
        </alternativeName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell junction</location>
            <location evidence="2">Focal adhesion</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Endosome membrane</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HB28"/>
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        <property type="protein sequence ID" value="ENSGAGP00000011945.1"/>
        <property type="gene ID" value="ENSGAGG00000004156.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
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        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
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    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
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        <property type="entry name" value="BAR-PH_GRAF_family"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="BAR_GRAF"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Rho GTPase activating protein 26"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000116">
        <property type="entry name" value="Rho GTPase activating protein 26"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Rho GTPase-activating protein 26"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.60">
        <property type="entry name" value="Arfaptin homology (AH) domain/BAR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
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    <dbReference type="Gene3D" id="1.10.555.10">
        <property type="entry name" value="Rho GTPase activation protein"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="AH/BAR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004148">
        <property type="entry name" value="BAR_dom"/>
    </dbReference>
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        <property type="entry name" value="GRAF_BAR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047234">
        <property type="entry name" value="GRAF_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047225">
        <property type="entry name" value="PH_GRAF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
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        <property type="entry name" value="RhoGAP_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12552">
        <property type="entry name" value="OLIGOPHRENIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="RHO GTPASE-ACTIVATING PROTEIN 26"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="BAR_3"/>
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    </dbReference>
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    </dbReference>
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        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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    <keyword evidence="5" id="KW-0343">GTPase activation</keyword>
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        <location>
            <begin position="265"/>
            <end position="369"/>
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            <begin position="375"/>
            <end position="560"/>
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        </source>
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        <source>
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        </source>
    </evidence>
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    </evidence>
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        </source>
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        </source>
    </evidence>
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        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
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        </source>
    </evidence>
    <sequence length="686" mass="78636" checksum="FE516AA47C8EEC3C" modified="2019-05-08" version="1">MGLPALEFSDCCLDSPQFRERLKSHEAELDKTNKFIKELIKDGKSLVAALKNLSSAKRKFADSLNEFKFRCIGDAETDDEICIAKSLQEFATVLRNLEDERMRMIENAGEVLITPLEKFRKEQIGAAKDAKKKYDKETEKYCGVLEKHLNLSSKKKESQLQEADSQVDLVRQHFYEVSLEYVFKVQEVQERKMFEFVEPLLAFLQGLFTFYHHGYELAKDFSDFKTELTISIQNTRNRFEGTRSEVESLMRKMKENPEEHRNISPFTMEGYLYVQEKRHFGTSWVKHYCTYQRESKRITMVPFDQKSGGKGGEDEAVTLKSCIRRKTDSIEKRFCFDVEAVDRPGVITMQALSEEDRKLWMEAMDGREPVRRDTAQLDNIGFSIIRKCINAVETRGINEQGLYRIVGVNSRVQKLLSILMDPKTASETETDICAEWEIKTITSALKTYLRMLPGPLMMYQFQRSFIKAAKLENQESRISEIHNLVHRLPEKNRQMLQMLMNHLANVANNHKQNLMTVANLGVVFGPTLLRPQEETVAAIMDIKFQNIVVEILIENHEKIFNTVPETLPPNSQLLLLSHKKNTDSKPPSCSERPLTLFHTAQANENQETRNSIINSSLESVISSNVNSFLHSNSTLRPSLNSSDPDLEVIKPSRPNSLKGYILSPTSSKNPINSTTTHCYCSAISVP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJG1</accession>
    <name>A0A452HJG1_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="7">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HJG1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017199.1">
        <property type="protein sequence ID" value="ENSGAGP00000015053.1"/>
        <property type="gene ID" value="ENSGAGG00000010747.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006396">
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd12223">
        <property type="entry name" value="RRM_SR140"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="U2 snRNP-associated SURP motif-containing protein isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.70.330:FF:000177">
        <property type="entry name" value="U2 snRNP-associated SURP motif-containing protein-like isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.790">
        <property type="entry name" value="Surp module"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="CID_dom"/>
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        <property type="entry name" value="ENTH_VHS"/>
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        <property type="entry name" value="mRNA_splic_Cwf21_dom"/>
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        <property type="entry name" value="Nucleotide-bd_a/b_plait_sf"/>
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        <property type="entry name" value="RBD_domain_sf"/>
    </dbReference>
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        <property type="entry name" value="RRM_dom"/>
    </dbReference>
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        <property type="entry name" value="SR-CTD_assoc_factor"/>
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        <property type="entry name" value="SR140_cwf21"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035009">
        <property type="entry name" value="SR140_RRM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000061">
        <property type="entry name" value="Surp"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035967">
        <property type="entry name" value="SWAP/Surp_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23140">
        <property type="entry name" value="RNA PROCESSING PROTEIN LD23810P"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23140:SF0">
        <property type="entry name" value="U2 SNRNP-ASSOCIATED SURP MOTIF-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04818">
        <property type="entry name" value="CID"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00076">
        <property type="entry name" value="RRM_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01805">
        <property type="entry name" value="Surp"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="cwf21"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="RPR"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="RRM"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="SWAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54928">
        <property type="entry name" value="RNA-binding domain, RBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF109905">
        <property type="entry name" value="Surp module (SWAP domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="CID"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50128">
        <property type="entry name" value="SURP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1 2" id="KW-0694">RNA-binding</keyword>
    <feature type="domain" description="RRM" evidence="4">
        <location>
            <begin position="187"/>
            <end position="268"/>
        </location>
    </feature>
    <feature type="domain" description="SURP motif" evidence="5">
        <location>
            <begin position="343"/>
            <end position="386"/>
        </location>
    </feature>
    <feature type="domain" description="CID" evidence="6">
        <location>
            <begin position="447"/>
            <end position="584"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="52"/>
            <end position="186"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="749"/>
            <end position="918"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="9"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="56"/>
            <end position="67"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="3">
        <location>
            <begin position="79"/>
            <end position="91"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="99"/>
            <end position="135"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="152"/>
            <end position="162"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="763"/>
            <end position="811"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="839"/>
            <end position="869"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="3">
        <location>
            <begin position="880"/>
            <end position="918"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00176"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50102"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS51391"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015053.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="918" mass="106265" checksum="D81D4DF98C352114" modified="2019-05-08" version="1">MSAAKRTLSKKEQEELKKKEDEKAAAEIYEEFLAAFEGSDGNKVKTFVRGGIVNASKDEHETDEKRGKIYKPSARFADQKNQPNQPSSNERPPSLLMIETKKPPLKKGEKEKKKSNLELFKEELKQIQEERDERHKTKGRLSRFEPPQSDSDGQRRSMDAPSRRNRSSGVLDDYAPGSHDVGDPSTTNLYLGNINPQMNEEMLCQEFGRFGPLASVKIMWPRTDEERARERNCGFVAFMNRRDAERALKNLNGKMIMSFEMKLGWGKAVPIPPHPIYIPPSMMEHTLPPPPSGLPFNAQPRERLKNPNAPMLPPPKNKEDFEKTLSQAIVKVVIPTERNLLALIHRMIEFVVREGPMFEAMIMNREINNPMFRFLFENQTPAHVYYRWKLYSILQGDAPTKWRTEDFRMFKNGSFWRPPPLNPYLHGMSEEQETEAFVEEPNKKGALKEEQRDKLEEILRGLTPRKNDIGDAMVFCLNNAEAAEEIVDCVTESLSILKTPLPKKIARLYLVSDVFFEFFIEINFETKLCQIFSDLNATYRTIQGHLQSENFKQRVMTCFRAWEDWAIYPEPFLIKLQNIFLGLVNIIEEKEIEEVPDDLDGAPIEEELDGAPLEDVDGIPIDAAPIDDLDGIPIKSLDDDLDGIPLDTSEESKKNEPIFKVAPSKWEAVDEAELEAQAVTTSKWELFDQHEESEEEENQKTKYTNNSFQPSFIFSQLSKTCEYCLVKMYFKWLLILQLKVMKFQDELESGKRPKKPGQSFQEQVEHYRDKLLQREKEKELERERERDKKDKEKSESRSKDKKEKEECTPTRKERKRRHSTSPSPSRSSGSRRAKSPSPKSERSERSERSHKETSRSRSSHKDSPRDVSKKAKRSPSGSRTPKRSRRSRSRSPKKSGKKSRSQSRSPHRSHKKSKKSKH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HJG3</accession>
    <name>A0A452HJG3_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="8">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HJG3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000017196.1">
        <property type="protein sequence ID" value="ENSGAGP00000015050.1"/>
        <property type="gene ID" value="ENSGAGG00000011031.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046982">
        <property type="term" value="F:protein heterodimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007265">
        <property type="term" value="P:Ras protein signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd22914">
        <property type="entry name" value="HFD_SOS1_rpt1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd22915">
        <property type="entry name" value="HFD_SOS1_rpt2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd01261">
        <property type="entry name" value="PH_SOS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00155">
        <property type="entry name" value="RasGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd06224">
        <property type="entry name" value="REM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00160">
        <property type="entry name" value="RhoGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000068">
        <property type="entry name" value="Son of sevenless homolog 1 (Drosophila)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.20.10:FF:000029">
        <property type="entry name" value="son of sevenless homolog 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.900.10:FF:000015">
        <property type="entry name" value="son of sevenless homolog 1 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.3060">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.900.10">
        <property type="entry name" value="Dbl homology (DH) domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.20.10">
        <property type="entry name" value="Histone, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.840.10">
        <property type="entry name" value="Ras guanine-nucleotide exchange factors catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.870.10">
        <property type="entry name" value="Son of sevenless (SoS) protein Chain: S domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035899">
        <property type="entry name" value="DBL_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000219">
        <property type="entry name" value="DH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009072">
        <property type="entry name" value="Histone-fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008937">
        <property type="entry name" value="Ras-like_GEF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000651">
        <property type="entry name" value="Ras-like_Gua-exchang_fac_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019804">
        <property type="entry name" value="Ras_G-nucl-exch_fac_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023578">
        <property type="entry name" value="Ras_GEF_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001895">
        <property type="entry name" value="RASGEF_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036964">
        <property type="entry name" value="RASGEF_cat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055251">
        <property type="entry name" value="SOS1_NGEF_PH"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23113">
        <property type="entry name" value="GUANINE NUCLEOTIDE EXCHANGE FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23113:SF150">
        <property type="entry name" value="SON OF SEVENLESS HOMOLOG 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00617">
        <property type="entry name" value="RasGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00618">
        <property type="entry name" value="RasGEF_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00621">
        <property type="entry name" value="RhoGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22697">
        <property type="entry name" value="SOS1_NGEF_PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00233">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00147">
        <property type="entry name" value="RasGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00229">
        <property type="entry name" value="RasGEFN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00325">
        <property type="entry name" value="RhoGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48065">
        <property type="entry name" value="DBL homology domain (DH-domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47113">
        <property type="entry name" value="Histone-fold"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48366">
        <property type="entry name" value="Ras GEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50010">
        <property type="entry name" value="DH_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00720">
        <property type="entry name" value="RASGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50009">
        <property type="entry name" value="RASGEF_CAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50212">
        <property type="entry name" value="RASGEF_NTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1 2" id="KW-0344">Guanine-nucleotide releasing factor</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="DH" evidence="6">
        <location>
            <begin position="198"/>
            <end position="388"/>
        </location>
    </feature>
    <feature type="domain" description="PH" evidence="4">
        <location>
            <begin position="441"/>
            <end position="544"/>
        </location>
    </feature>
    <feature type="domain" description="N-terminal Ras-GEF" evidence="7">
        <location>
            <begin position="595"/>
            <end position="739"/>
        </location>
    </feature>
    <feature type="domain" description="Ras-GEF" evidence="5">
        <location>
            <begin position="777"/>
            <end position="1013"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="3">
        <location>
            <begin position="1014"/>
            <end position="1277"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="3">
        <location>
            <begin position="1074"/>
            <end position="1094"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="3">
        <location>
            <begin position="1118"/>
            <end position="1130"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="3">
        <location>
            <begin position="1147"/>
            <end position="1167"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="1187"/>
            <end position="1196"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="3">
        <location>
            <begin position="1255"/>
            <end position="1264"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022658"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00168"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50009"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50010"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000015050.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1277" mass="147795" checksum="C12362DBF141E0DA" modified="2019-05-08" version="1">MQPPPQPYEFFSEENGPKWRGLFVPALRKVQQQVHPSFSAKEDSLYYIEELILQLLNKLCIAQPRTVQDVEERVQKTFPHPIDKWAIADAQSAIEKRKRRNPLLLPVDKIHPLLKEVLGYKVDYHVSLYIVAVLEYISADILKLAGNYVFNIRHFEISQQDIKVSMCADKVLMDMFDQDVIGLVSLCEDEPSSSGELNYYELVRTEIAEERQYLRELNLIIKVFREAFLSNRKLFTPYDIDTIFSNISDIHELTVKLLGLIEDTVEMTDESSPHPLAGSCFEDLAEEQAFDPYETLSQDILSPRFHEHFNNLMAKPAVALHFQSIAEGFKEAVQYVLPRLMLIPVYHCLHYFELLQQLQECSEDEEDRECLKQAITALLNLQCSMERIYNKYSPRRRPWETICRFYNRQVRSKHLAIKKMNEIQKNIDGWEGKDIGQCCNEFIMEGALTRIGAKHERHIFLFDGLLISCKTNHGQSRLPGYSNTEYRLKEKIIMRKMQIIDKDDTCEYKHAFELASKDENSLVFAAKSAEEKNNWMAALISLQYRSTLDRMLDSVLLQEENEQPLRLPSPNVYRFVVEDSEENIVFEDNLQSRNGIPIIKGGTVVKLIERLTYHMYADPNFVRTFLTTYRSFCKPQELLSLLIERFEIPEPEPTEADRLAIEKGEQPISADLKRFRKEYVQPVQLRILNVFRHWVEHHFYDFERDLELLDKLETFISSVRGKSMKKWVESIAKIIKRKKQAQANGISHNITFESQPPPIEWHICHLGQFEMDLMTLHPIEIARQLTLLESDLYRAVQPSELVGSVWTKEDKEINSPNLLKMIRHTTNLTLWFEKCIVETENFEERVAVLSRIIEILQVFQELNNFNGVLEIVSAMNSVSVYRLDHTFEERKKKVLDEAVELSQDHFKKYLAKLKSINPPCVPFFGIYLTNILKTEEGNPDFLKRQGKELINFSKRRKVAEITGEIQQYQNQPYCLRIEPEIRRFFENLNPMGNAPEKEFTDYLFNKSLEIEPRNCKQPPRFPRKTTYPLKSPGIRPNTGRHCSTSGTLRGHPTPLEREPCKISFSRITEAEQESAASAPTSPNTPSTPPVSASSERSVFLDIDLNSSYGILKSDDDPPAIPPRQPPPPKIQPRVPAYNNPFDQLLHSPPPPPPRDPLPDTPPPVPLRPPEHFINCPFNLQPPPVGRFHRDPDRFREASTCPSSPNTPPSTPSPRVLRRCYMLSSSHNNVAHPPAPPVPPRQNSSPQLPKLPPKTYKRELSHPPLDRQFLLENAETPQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HRQ3</accession>
    <name>A0A452HRQ3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Regulatory factor X-associated protein RFXANK-binding domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HRQ3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000020257.1">
        <property type="protein sequence ID" value="ENSGAGP00000017758.1"/>
        <property type="gene ID" value="ENSGAGG00000013182.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006357">
        <property type="term" value="P:regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.290.30">
        <property type="entry name" value="Regulatory factor X-associated C-terminal binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038308">
        <property type="entry name" value="RFXAP_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029316">
        <property type="entry name" value="RFXAP_RFXANK-bd"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15110">
        <property type="entry name" value="REGULATORY FACTOR X-ASSOCIATED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15110:SF2">
        <property type="entry name" value="REGULATORY FACTOR X-ASSOCIATED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15289">
        <property type="entry name" value="RFXA_RFXANK_bdg"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Regulatory factor X-associated protein RFXANK-binding" evidence="2">
        <location>
            <begin position="72"/>
            <end position="197"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="24"/>
            <end position="154"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="1">
        <location>
            <begin position="27"/>
            <end position="36"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="72"/>
            <end position="85"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="1">
        <location>
            <begin position="86"/>
            <end position="109"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="1">
        <location>
            <begin position="119"/>
            <end position="129"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF15289"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000017758.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="202" mass="21275" checksum="30F769CF8048B01D" modified="2019-05-08" version="1">TEGPGGSSSRALRPGLVFLCEVCPGEGDGDAAEEGEPAGGESTASPEELEDEEAAAASGGETPAGGGGCKSCTYQGCSETTTQVAKQRKPWMCKRHRNKMYKDKYKRKKSDQALGGAGVPPAQGGAATPGAPPSLEISVTKQRTGSIGDRPARPTLLEQVLNQKRLSLLRSPEVVQFLQKQQQLLSQQALEQRQQQFPGAPV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HRQ4</accession>
    <name>A0A452HRQ4_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="8">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the SLC13A/DASS transporter (TC 2.A.47) family. NADC subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HRQ4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000020243.1">
        <property type="protein sequence ID" value="ENSGAGP00000017746.1"/>
        <property type="gene ID" value="ENSGAGG00000012896.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015139">
        <property type="term" value="F:alpha-ketoglutarate transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015138">
        <property type="term" value="F:fumarate transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017153">
        <property type="term" value="F:sodium:dicarboxylate symporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015141">
        <property type="term" value="F:succinate transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071285">
        <property type="term" value="P:cellular response to lithium ion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd01115">
        <property type="entry name" value="SLC13_permease"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001898">
        <property type="entry name" value="SLC13A/DASS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10283">
        <property type="entry name" value="SOLUTE CARRIER FAMILY 13 MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10283:SF82">
        <property type="entry name" value="SOLUTE CARRIER FAMILY 13 MEMBER 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00939">
        <property type="entry name" value="Na_sulph_symp"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0406">Ion transport</keyword>
    <keyword evidence="5 7" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0915">Sodium</keyword>
    <keyword evidence="6" id="KW-0739">Sodium transport</keyword>
    <keyword evidence="3 7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 7" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="6" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="6"/>
            <end position="25"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="32"/>
            <end position="59"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="79"/>
            <end position="97"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="109"/>
            <end position="131"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="258"/>
            <end position="280"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="324"/>
            <end position="343"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="363"/>
            <end position="389"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="410"/>
            <end position="427"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="447"/>
            <end position="463"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="475"/>
            <end position="496"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="502"/>
            <end position="522"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="534"/>
            <end position="553"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006772"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023201"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000017746.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="574" mass="63341" checksum="7E3E7D2496BFBB21" modified="2019-05-08" version="1">RRSFWIILWIAKSAVLCVLIVCVLFQEAKCGYVIILMALFWCMEVLPLGVTALFPVVLFPLMNIMDSTKVCMEYLKDSNMLFLGGLMVAIAVEHWNLHKRIALRVLLIVGVRPALLIMGFMSVTAFLSMWISNTATTAMMIPIAQAVLDQLHKSEQETSPVGHVAESINKAFELQEVPDKSDPSLCNSSPVPASEEEKLEDVKQLEEKHLTFCKGMSLSVCYAASIGGIATLTGSTPNLVMKGQMDALFPTNNNVVNFASWFMFAFPTMLLLLALSWIWLQIMFLGFDFRKNLGFGANASMKAKQKMAYRIIKDEHKKLGSMRFAEIAVTVLFLILVVLWFTRDPGFIPGWATNVFNKNGESYVTDATVVMFISLLMFLIPSEIPTYLYSTKQIPVLPALLDWKTVNQKMPWNIVILLGGGFALAKGSEESGLSEWLGSKLTPLQNMSSPIIVLLLCLLVATFTECTSNVATTTLFLPILAAMAQTICLNPLYVMLPCTLSASLAFMLPVATPPNAIAFSYGQLKVIDMAKAGSVLNILGVLTIMLAINTWGYPLFDLNHFPVWANSTSAASCK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HRQ8</accession>
    <name>A0A452HRQ8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Mothers against decapentaplegic homolog</fullName>
            <shortName evidence="8">MAD homolog</shortName>
            <shortName evidence="8">Mothers against DPP homolog</shortName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="8">SMAD family member</fullName>
        </alternativeName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="8">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="8">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="1 8">Belongs to the dwarfin/SMAD family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HRQ8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000020262.1">
        <property type="protein sequence ID" value="ENSGAGP00000017763.1"/>
        <property type="gene ID" value="ENSGAGG00000013188.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071144">
        <property type="term" value="C:heteromeric SMAD protein complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070411">
        <property type="term" value="F:I-SMAD binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009653">
        <property type="term" value="P:anatomical structure morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030509">
        <property type="term" value="P:BMP signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060395">
        <property type="term" value="P:SMAD protein signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007179">
        <property type="term" value="P:transforming growth factor beta receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd10490">
        <property type="entry name" value="MH1_SMAD_1_5_9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.200.10:FF:000001">
        <property type="entry name" value="Mothers against decapentaplegic homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.520.10:FF:000001">
        <property type="entry name" value="Mothers against decapentaplegic homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.200.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.520.10">
        <property type="entry name" value="SMAD MH1 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013790">
        <property type="entry name" value="Dwarfin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003619">
        <property type="entry name" value="MAD_homology1_Dwarfin-type"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013019">
        <property type="entry name" value="MAD_homology_MH1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017855">
        <property type="entry name" value="SMAD-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001132">
        <property type="entry name" value="SMAD_dom_Dwarfin-type"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008984">
        <property type="entry name" value="SMAD_FHA_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036578">
        <property type="entry name" value="SMAD_MH1_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13703:SF41">
        <property type="entry name" value="MOTHERS AGAINST DECAPENTAPLEGIC HOMOLOG 9"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13703">
        <property type="entry name" value="SMAD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03165">
        <property type="entry name" value="MH1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03166">
        <property type="entry name" value="MH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00523">
        <property type="entry name" value="DWA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00524">
        <property type="entry name" value="DWB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56366">
        <property type="entry name" value="SMAD MH1 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49879">
        <property type="entry name" value="SMAD/FHA domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51075">
        <property type="entry name" value="MH1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51076">
        <property type="entry name" value="MH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2 8" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7 8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6 8" id="KW-0804">Transcription</keyword>
    <keyword evidence="5 8" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="4" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="MH1" evidence="10">
        <location>
            <begin position="16"/>
            <end position="140"/>
        </location>
    </feature>
    <feature type="domain" description="MH2" evidence="11">
        <location>
            <begin position="236"/>
            <end position="430"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="182"/>
            <end position="218"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="192"/>
            <end position="218"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00005545"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="RuleBase" id="RU361195"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS51075"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS51076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000017763.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="430" mass="48567" checksum="8ECF357F537C0974" modified="2019-05-08" version="1">MHSSTPISSLFSFTSPAVKRLLGWKQGDEEEKWAEKAVDSLVKKLKKKKGAMEELERALSCPGQPSKCVTIPRSLDGRLQVSHRKGLPHVIYCRVWRWPDLQSHHELKPLECCEFPFGSKQKEVCINPYHYRRVETPVLPPVLVPRHSEFNPHLSLLAKFRSASLHNEPLMPHNATYPDSFQQTPCTPFPSSPSNMFSQSPSSISYPNSPGSSSGPGSPYQLTDFRPVCYEEPQHWCSVAYYELNNRVGETFQASSRSILIDGFTDPSNNKNRFCLGLLSNVNRNSTIENTRRHIGKGVHLYYVGGEVYAECVSDSSIFVQSRNCNYQHGFHPATVCKIPSGCSLKIFNNQLFAQLLAQSVNHGFEVVYELTKMCTIRMSFVKGWGAEYHRQDVTSTPCWIEIHLHGPLQWLDKVLTQMGSPHNPISSVS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HRR2</accession>
    <name>A0A452HRR2_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="8">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the SLC13A/DASS transporter (TC 2.A.47) family. NADC subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HRR2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000020193.1">
        <property type="protein sequence ID" value="ENSGAGP00000017698.1"/>
        <property type="gene ID" value="ENSGAGG00000012896.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015139">
        <property type="term" value="F:alpha-ketoglutarate transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015138">
        <property type="term" value="F:fumarate transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017153">
        <property type="term" value="F:sodium:dicarboxylate symporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015141">
        <property type="term" value="F:succinate transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071285">
        <property type="term" value="P:cellular response to lithium ion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd01115">
        <property type="entry name" value="SLC13_permease"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001898">
        <property type="entry name" value="SLC13A/DASS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10283">
        <property type="entry name" value="SOLUTE CARRIER FAMILY 13 MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10283:SF82">
        <property type="entry name" value="SOLUTE CARRIER FAMILY 13 MEMBER 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00939">
        <property type="entry name" value="Na_sulph_symp"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0406">Ion transport</keyword>
    <keyword evidence="5 7" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0915">Sodium</keyword>
    <keyword evidence="6" id="KW-0739">Sodium transport</keyword>
    <keyword evidence="3 7" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 7" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="6" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="15"/>
            <end position="38"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="45"/>
            <end position="72"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="92"/>
            <end position="110"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="122"/>
            <end position="144"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="250"/>
            <end position="272"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="316"/>
            <end position="335"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="355"/>
            <end position="381"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="402"/>
            <end position="419"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="439"/>
            <end position="455"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="467"/>
            <end position="488"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="494"/>
            <end position="514"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="7">
        <location>
            <begin position="526"/>
            <end position="545"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006772"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023201"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000017698.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="572" mass="62647" checksum="D870ED855E89597E" modified="2019-05-08" version="1">MQCCSSSVATKSAVIGLQACGLVSQSAVLCVLIVCVLFQEAKCGYVIILMALFWCMEVLPLGVTALFPVVLFPLMNIMDSTKVCMEYLKDSNMLFLGGLMVAIAVEHWNLHKRIALRVLLIVGVRPALLIMGFMSVTAFLSMWISNTATTAMMIPIAQAVLDQLHKSEQETSPVGHVAESINKAFELIGVIASNPTLEEKHLTFCKGMSLSVCYAASIGGIATLTGSTPNLVMKGQMDALFPTNNNVVNFASWFMFAFPTMLLLLALSWIWLQIMFLGFDFRKNLGFGANASMKAKQKMAYRIIKDEHKKLGSMRFAEIAVTVLFLILVVLWFTRDPGFIPGWATNVFNKNGESYVTDATVVMFISLLMFLIPSEIPTYLYSTKQIPVLPALLDWKTVNQKMPWNIVILLGGGFALAKGSEESGLSEWLGSKLTPLQNMSSPIIVLLLCLLVATFTECTSNVATTTLFLPILAAMAQTICLNPLYVMLPCTLSASLAFMLPVATPPNAIAFSYGQLKVIDMVSTGSVLNILGVLTIMLAINTWGYPLFDLNHFPVWANSTSTQHLETKSPRC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HRR3</accession>
    <name>A0A452HRR3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Zinc transporter ZIP4/12 EF-hand domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000020249.1">
        <property type="protein sequence ID" value="ENSGAGP00000017752.1"/>
        <property type="gene ID" value="ENSGAGG00000013178.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140410">
        <property type="term" value="F:monoatomic cation:bicarbonate symporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005385">
        <property type="term" value="F:zinc ion transmembrane transporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030003">
        <property type="term" value="P:intracellular monoatomic cation homeostasis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071578">
        <property type="term" value="P:zinc ion import across plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049406">
        <property type="entry name" value="ZIP4_12_EF-hand"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050799">
        <property type="entry name" value="ZIP_Transporter"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12191">
        <property type="entry name" value="SOLUTE CARRIER FAMILY 39"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12191:SF21">
        <property type="entry name" value="ZINC TRANSPORTER ZIP4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21116">
        <property type="entry name" value="EF-hand_Zip"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="1" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="1">
        <location>
            <begin position="257"/>
            <end position="280"/>
        </location>
    </feature>
    <feature type="domain" description="Zinc transporter ZIP4/12 EF-hand" evidence="2">
        <location>
            <begin position="198"/>
            <end position="238"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF21116"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000017752.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="332" mass="34542" checksum="68B2C5D55368A0EE" modified="2019-05-08" version="1">MGLSAVWCVPASALPLCPQCISVPDIFMLVGKAGTGDTNLTATELLPLGAGVLLYLTDPVATCAAVRGGSXYLVLSAISLFRPNSPKAPGPGVRGLRETCSANAQQLPALWSTAVSGQVAADMAGQVLVTIMYHVLLGDCFHALPPPHYFLGYIFRRYGNESQNLTLAGESGALGRGGLGSVAWVPPTPSVTPHLCLQMCLSPGDLLEIYSLDPDAGISGSDFTSLSPALIQQQLSRACSARHLSAPGRQLTMAERYIYGSLATLVICLCALLGIVLLLCTACASAYPYIIQGFVSLAVGSLTGDALLHLIPQVSSWACRGPTPGTTCTARK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HRR4</accession>
    <name>A0A452HRR4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="36">Tyrosine-protein kinase receptor</fullName>
            <ecNumber evidence="36">2.7.10.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="42 43">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="43" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="42" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="42" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="31 36">
            <text>L-tyrosyl-[protein] + ATP = O-phospho-L-tyrosyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10596"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.10.1"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cell projection</location>
            <location evidence="3">Axon</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cell projection</location>
            <location evidence="2">Dendrite</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="5">Cytoplasm</location>
            <location evidence="5">Perinuclear region</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Endosome membrane</location>
            <topology evidence="4">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="30">Postsynaptic density</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="36">Belongs to the protein kinase superfamily. Tyr protein kinase family. Insulin receptor subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.10.1" evidence="36"/>
    <dbReference type="AlphaFoldDB" id="A0A452HRR4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000020268.1">
        <property type="protein sequence ID" value="ENSGAGP00000017768.1"/>
        <property type="gene ID" value="ENSGAGG00000013192.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043679">
        <property type="term" value="C:axon terminus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043197">
        <property type="term" value="C:dendritic spine"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010008">
        <property type="term" value="C:endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048471">
        <property type="term" value="C:perinuclear region of cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0014069">
        <property type="term" value="C:postsynaptic density"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043235">
        <property type="term" value="C:receptor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048403">
        <property type="term" value="F:brain-derived neurotrophic factor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060175">
        <property type="term" value="F:brain-derived neurotrophic factor receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:1990416">
        <property type="term" value="P:cellular response to brain-derived neurotrophic factor stimulus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010976">
        <property type="term" value="P:positive regulation of neuron projection development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051897">
        <property type="term" value="P:positive regulation of phosphatidylinositol 3-kinase/protein kinase B signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0099551">
        <property type="term" value="P:trans-synaptic signaling by neuropeptide, modulating synaptic transmission"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd05855">
        <property type="entry name" value="IgI_TrkB_d5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05093">
        <property type="entry name" value="PTKc_TrkB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000239">
        <property type="entry name" value="BDNF/NT-3 growth factors receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000034">
        <property type="entry name" value="Tyrosine-protein kinase receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000486">
        <property type="entry name" value="Tyrosine-protein kinase receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000033">
        <property type="entry name" value="Tyrosine-protein kinase receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000319">
        <property type="entry name" value="Tyrosine-protein kinase receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000483">
        <property type="entry name" value="Cys-rich_flank_reg_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013098">
        <property type="entry name" value="Ig_I-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001611">
        <property type="entry name" value="Leu-rich_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000372">
        <property type="entry name" value="LRRNT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020777">
        <property type="entry name" value="NTRK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020455">
        <property type="entry name" value="NTRK2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031635">
        <property type="entry name" value="NTRK_LRRCT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050122">
        <property type="entry name" value="RTK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001245">
        <property type="entry name" value="Ser-Thr/Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008266">
        <property type="entry name" value="Tyr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020635">
        <property type="entry name" value="Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002011">
        <property type="entry name" value="Tyr_kinase_rcpt_2_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416:SF136">
        <property type="entry name" value="BDNF_NT-3 GROWTH FACTORS RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416">
        <property type="entry name" value="TYROSINE-PROTEIN KINASE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07679">
        <property type="entry name" value="I-set"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13855">
        <property type="entry name" value="LRR_8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16920">
        <property type="entry name" value="LRRCT_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07714">
        <property type="entry name" value="PK_Tyr_Ser-Thr"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01939">
        <property type="entry name" value="NTKRECEPTOR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01941">
        <property type="entry name" value="NTKRECEPTOR2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00109">
        <property type="entry name" value="TYRKINASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00082">
        <property type="entry name" value="LRRCT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00013">
        <property type="entry name" value="LRRNT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00219">
        <property type="entry name" value="TyrKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52058">
        <property type="entry name" value="L domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00109">
        <property type="entry name" value="PROTEIN_KINASE_TYR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00239">
        <property type="entry name" value="RECEPTOR_TYR_KIN_II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="19 33 35" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="7" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="28" id="KW-0966">Cell projection</keyword>
    <keyword evidence="8" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="18" id="KW-0221">Differentiation</keyword>
    <keyword evidence="25" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="16" id="KW-0967">Endosome</keyword>
    <keyword evidence="27" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="29" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="17" id="KW-0418">Kinase</keyword>
    <keyword evidence="10" id="KW-0433">Leucine-rich repeat</keyword>
    <keyword evidence="23 38" id="KW-0472">Membrane</keyword>
    <keyword evidence="20" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="15 33 35" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="9 36" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="26 36" id="KW-0675">Receptor</keyword>
    <keyword evidence="43" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="14" id="KW-0677">Repeat</keyword>
    <keyword evidence="13 39" id="KW-0732">Signal</keyword>
    <keyword evidence="22" id="KW-0770">Synapse</keyword>
    <keyword evidence="11" id="KW-0808">Transferase</keyword>
    <keyword evidence="12 36 38" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="21 38" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="24" id="KW-0829">Tyrosine-protein kinase</keyword>
    <feature type="signal peptide" evidence="39">
        <location>
            <begin position="1"/>
            <end position="32"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019017591" description="Tyrosine-protein kinase receptor" evidence="39">
        <location>
            <begin position="33"/>
            <end position="809"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="38">
        <location>
            <begin position="418"/>
            <end position="441"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="41">
        <location>
            <begin position="198"/>
            <end position="283"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="40">
        <location>
            <begin position="525"/>
            <end position="794"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="37">
        <location>
            <begin position="460"/>
            <end position="485"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="37">
        <location>
            <begin position="472"/>
            <end position="482"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="32">
        <location>
            <position position="663"/>
        </location>
    </feature>
    <feature type="binding site" evidence="33">
        <location>
            <begin position="531"/>
            <end position="539"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="33 35">
        <location>
            <position position="559"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="site" description="Interaction with SHC1" evidence="34">
        <location>
            <position position="503"/>
        </location>
    </feature>
    <feature type="site" description="Interaction with PLCG1" evidence="34">
        <location>
            <position position="804"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004279"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004489"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004556"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022614"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00023137"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="ARBA" id="ARBA00034105"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="31">
        <source>
            <dbReference type="ARBA" id="ARBA00051243"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="32">
        <source>
            <dbReference type="PIRSR" id="PIRSR620777-50"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="33">
        <source>
            <dbReference type="PIRSR" id="PIRSR620777-51"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="34">
        <source>
            <dbReference type="PIRSR" id="PIRSR620777-52"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="35">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="36">
        <source>
            <dbReference type="RuleBase" id="RU000312"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="37">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="38">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="39">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="40">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="41">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="42">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000017768.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="43">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="809" mass="90858" checksum="4448792AFBC34632" modified="2019-05-08" version="1">MVSWRRRWQGPGLARLWGFCCLVFGSWRALLACPAPCTCSASRIWCGEPAPGIVSFPVLERSAERENITDIYIANQRKLESINNNEVGLYVGLKNLTVVDSGLKFVSRQAFLKNTNLQYINFSRNKLSSLYKKTFRHLNLSDLILVDNPFKCSCDVMWIKIFQETKFSNPETQDFYCINDNNKRIPLSDMQIPNCGLPSANLSVYNLTIVEGRSTTLYCEATGVPSPNVSWVVANLVSKHESNTSKNPASLTIRNVSSKDSGIRISCIAENIVGEDQASADLVVFFAPNITFIEPPTPDHHWCIPFTVKGNPKPSLQWFYEGAILNESEYICTKIHVSNQSEYHGCLQLDNPTHLNNGAYTLLAKNEYGEDEKRVDAHFMSMPGDDYETTPNDIGDTTNNSNQITSTDVSNKENEDSITVYVVVGIAALVCTGLVIMLIILKFGRHSKFGMKGPSSVISNDDDSASPLHHISNGSNTPSSSEGGPDAVIIGMTKIPVIENPQYFGITNSQLKPDTFVQHIKRHNIVLKRELGEGAFGKVFLAECYNLSPEQDKILVAVKTLKDASDNARKDFHREAELLTNLQHEHIVKFYGVCVEGDPLIMVFEYMKHGDLNKFLRAHGPDAVLMAEGNLLAELTQSQMLHIAQQIASGMVYLASQHFVHRDLATRNCLVGENLLVKIGDFGMSRDVYSTDYYRVGGHTMLPIRWMPPESIMYRKFTTESDVWSLGVVLWEIFTYGKQPWYQLSNNEVIECITQGRVLQRPRTCPKEVYDLMLGCWQREPHMRLNIKEIHSLLQNLAKASPVYLDILG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HS20</accession>
    <name>A0A452HS20_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">tripeptidyl-peptidase II</fullName>
            <ecNumber evidence="3">3.4.14.10</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>Release of an N-terminal tripeptide from a polypeptide.</text>
            <dbReference type="EC" id="3.4.14.10"/>
        </reaction>
    </comment>
    <comment type="similarity">
        <text evidence="2 8">Belongs to the peptidase S8 family.</text>
    </comment>
    <dbReference type="EC" id="3.4.14.10" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452HS20"/>
    <dbReference type="Ensembl" id="ENSGAGT00000020401.1">
        <property type="protein sequence ID" value="ENSGAGP00000017884.1"/>
        <property type="gene ID" value="ENSGAGG00000011247.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004177">
        <property type="term" value="F:aminopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008240">
        <property type="term" value="F:tripeptidyl-peptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd04857">
        <property type="entry name" value="Peptidases_S8_Tripeptidyl_Aminopeptidase_II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.20.25.690">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.3170">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.3080">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.200">
        <property type="entry name" value="Peptidase S8/S53 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000209">
        <property type="entry name" value="Peptidase_S8/S53_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036852">
        <property type="entry name" value="Peptidase_S8/S53_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022398">
        <property type="entry name" value="Peptidase_S8_His-AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023828">
        <property type="entry name" value="Peptidase_S8_Ser-AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050131">
        <property type="entry name" value="Peptidase_S8_subtilisin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015500">
        <property type="entry name" value="Peptidase_S8_subtilisin-rel"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034051">
        <property type="entry name" value="TPP_II_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048384">
        <property type="entry name" value="TPPII_GBD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048383">
        <property type="entry name" value="TPPII_Ig-like-1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022229">
        <property type="entry name" value="TPPII_Ig-like-2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046940">
        <property type="entry name" value="TPPII_Ig-like_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43806">
        <property type="entry name" value="PEPTIDASE S8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43806:SF14">
        <property type="entry name" value="TRIPEPTIDYL-PEPTIDASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00082">
        <property type="entry name" value="Peptidase_S8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12580">
        <property type="entry name" value="TPPII"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21316">
        <property type="entry name" value="TPPII_GBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21223">
        <property type="entry name" value="TPPII_Ig-like-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00723">
        <property type="entry name" value="SUBTILISIN"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52743">
        <property type="entry name" value="Subtilisin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51892">
        <property type="entry name" value="SUBTILASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00137">
        <property type="entry name" value="SUBTILASE_HIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00138">
        <property type="entry name" value="SUBTILASE_SER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0031">Aminopeptidase</keyword>
    <keyword evidence="6 8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="5 8" id="KW-0645">Protease</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7 8" id="KW-0720">Serine protease</keyword>
    <feature type="domain" description="Peptidase S8/S53" evidence="9">
        <location>
            <begin position="34"/>
            <end position="499"/>
        </location>
    </feature>
    <feature type="domain" description="Tripeptidyl-peptidase II first Ig-like" evidence="11">
        <location>
            <begin position="520"/>
            <end position="633"/>
        </location>
    </feature>
    <feature type="domain" description="Tripeptidyl-peptidase II galactose-binding" evidence="12">
        <location>
            <begin position="653"/>
            <end position="740"/>
        </location>
    </feature>
    <feature type="domain" description="Tripeptidyl peptidase II second Ig-like" evidence="10">
        <location>
            <begin position="776"/>
            <end position="876"/>
        </location>
    </feature>
    <feature type="active site" description="Charge relay system" evidence="8">
        <location>
            <position position="43"/>
        </location>
    </feature>
    <feature type="active site" description="Charge relay system" evidence="8">
        <location>
            <position position="263"/>
        </location>
    </feature>
    <feature type="active site" description="Charge relay system" evidence="8">
        <location>
            <position position="448"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001910"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00011073"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012462"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022438"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF00082"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="Pfam" id="PF12580"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF21223"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF21316"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000017884.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="892" mass="98220" checksum="0204ABB5EC552CE7" modified="2025-10-08" version="2">MAGWTHEPFPFHGLLPKKETGAASFLGRYPEYDGRGVLLAVLDTGVDPGAPGMQITTDGKPKIIDIIDTTGSGDVNTSTIVEPKDGEIVGLSGRTLKIPTNWVNPSGKYHIGIKNAYDFYPKALKERIQKERKEKQWDPVHRLLLAEACRKLDEFDAVHNSPSQTTKLMKEEFQSQVELLNSLEKKYNDPGPVYDCLVWHDGETWRACIDSSECGDFTKCTVLGTYRERQEYGSFGISEMLNYSVNIYDEGNLLSIVTSGGAHGTHVASIAAGYFPEEPERNGVAPGAQILAIKIGDTRLSTMETGTGLIRAMIEAMKYKCDLVNYSYGEATHWPNSGRICEVINEAVWKHNVIYVSSAGNNGPCLSTVGCPGGTTSSVIGVGAYVSPDMMVAEYSLREKLPANQYTWSSRGPSTDGALGVSISAPGGAIASVPNWTLRGTQLMNGTSMSSPNACGGIALVLSGLKANDVHYTVHSVRRALENTAVKAENIEVFAQGHGIIQVDKAYDYLIQNSSFTSNVGFTVTVGNNRGIYLRDPVQIAAPSDRGVGIEPVFPENTENTKRISLQLHLALTSNAAWVQCPSHLELMNQCRHINVRVDPRGLREGLHYTEVCGYDIAAPNAGPLFRVPITLIIPARIDESSHYDLSYTDVHFKPGQIRRHFIEVPQGATWAEVTVCSCSTDVTAKFVLHAVQLVKQKAYRSHEFYKFSSLPEKGSVTEAFPVLGGKTIEFCIARWWASLNDVSIDYTISFHGIVCATPQLNIHASEGIVRFDVQSTLKYEDIAPCINLKSWVQTLRPVSTKIKPLGSRDVLPNNRQLYEMILTYSFHQPKSGEVTPSCPLLCELLYESEFDSQLWIIFDQNKRQMGSGDAYPHQKQASILHPDPKFMHLTA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HS23</accession>
    <name>A0A452HS23_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">MAPK/MAK/MRK overlapping kinase</fullName>
            <ecNumber evidence="6">2.7.11.22</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="22">MOK protein kinase</fullName>
        </alternativeName>
    </protein>
    <organism evidence="26 27">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="27" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="26" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="26" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="20">Able to phosphorylate several exogenous substrates and to undergo autophosphorylation. Negatively regulates cilium length in a cAMP and mTORC1 signaling-dependent manner.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.22"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.11.22"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cell projection</location>
            <location evidence="3">Cilium</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="5">Belongs to the protein kinase superfamily. CMGC Ser/Thr protein kinase family. CDC2/CDKX subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.22" evidence="6"/>
    <dbReference type="AlphaFoldDB" id="A0A452HS23"/>
    <dbReference type="Ensembl" id="ENSGAGT00000020362.1">
        <property type="protein sequence ID" value="ENSGAGP00000017846.1"/>
        <property type="gene ID" value="ENSGAGG00000013244.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005929">
        <property type="term" value="C:cilium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004693">
        <property type="term" value="F:cyclin-dependent protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd07831">
        <property type="entry name" value="STKc_MOK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000402">
        <property type="entry name" value="MAPK/MAK/MRK overlapping kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000271">
        <property type="entry name" value="MAPK/MAK/MRK overlapping kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050117">
        <property type="entry name" value="MAPK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24055">
        <property type="entry name" value="MITOGEN-ACTIVATED PROTEIN KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="14 23 24" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="17" id="KW-0966">Cell projection</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="13" id="KW-0418">Kinase</keyword>
    <keyword evidence="15" id="KW-0460">Magnesium</keyword>
    <keyword evidence="11" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="12 23 24" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="16" id="KW-0539">Nucleus</keyword>
    <keyword evidence="9" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="27" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8 24" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="10" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Protein kinase" evidence="25">
        <location>
            <begin position="4"/>
            <end position="285"/>
        </location>
    </feature>
    <feature type="binding site" evidence="23">
        <location>
            <position position="33"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004138"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00006485"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00012425"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00047811"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00048367"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00058464"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00069211"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00076383"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="RuleBase" id="RU000304"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000017846.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="420" mass="48458" checksum="F86329B1BBC481A4" modified="2019-05-08" version="1">MNKYKPIGKIGEGTFSDVLKTQSLRDGNYYACKQMKQHFESIEQVNSLREIQALRRLNPHPNILMLHEVVFDRKSGCLALICELMDMNIYELIRGRRKPLSEKKIMNYMYQLCKSLDHMHRNGIFHRDVKPENILIKQDTLKLGDFGSCRSVYSKQPYTEYISTRWYRAPECLLTDGYYSYKMDMWSSGCVFYEITSFHPLFPGSNELDQISKIHDILGTPAKKTLNKFKQSRAMSFDFPFKKGTGLSPHVPNISPKSLSLMYAMIEYDPDQRIGAHQALQHPYFQELRTAEKQAVTMHRKMRLAENIVGQESINLWCISKDGNRQHSLRKTQENPIRHHGPSCGIELPKLNVSGVTKLTSYPSPSFHSVFTEPATNGEVPVLQPIKYIGTNHKSEKQKELKSSMKQYHLPAIERRGGGY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HS24</accession>
    <name>A0A452HS24_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">BTB domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>K(+)(in) = K(+)(out)</text>
            <dbReference type="Rhea" id="RHEA:29463"/>
            <dbReference type="ChEBI" id="CHEBI:29103"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="17">Presynaptic cell membrane</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HS24"/>
    <dbReference type="Ensembl" id="ENSGAGT00000020380.1">
        <property type="protein sequence ID" value="ENSGAGP00000017863.1"/>
        <property type="gene ID" value="ENSGAGG00000013255.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000020382.1">
        <property type="protein sequence ID" value="ENSGAGP00000017865.1"/>
        <property type="gene ID" value="ENSGAGG00000013255.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043679">
        <property type="term" value="C:axon terminus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032590">
        <property type="term" value="C:dendrite membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032809">
        <property type="term" value="C:neuronal cell body membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042734">
        <property type="term" value="C:presynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008076">
        <property type="term" value="C:voltage-gated potassium channel complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005251">
        <property type="term" value="F:delayed rectifier potassium channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001508">
        <property type="term" value="P:action potential"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051260">
        <property type="term" value="P:protein homooligomerization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.70:FF:000011">
        <property type="entry name" value="Potassium channel, voltage-gated Shaw-related subfamily C, member 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.350:FF:000014">
        <property type="entry name" value="Potassium channel, voltage-gated Shaw-related subfamily C, member 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.710.10:FF:000002">
        <property type="entry name" value="Potassium voltage-gated channel subfamily C member 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.70">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.710.10">
        <property type="entry name" value="Potassium Channel Kv1.1, Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.350">
        <property type="entry name" value="Voltage-gated potassium channels. Chain C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000210">
        <property type="entry name" value="BTB/POZ_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005821">
        <property type="entry name" value="Ion_trans_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003968">
        <property type="entry name" value="K_chnl_volt-dep_Kv"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003974">
        <property type="entry name" value="K_chnl_volt-dep_Kv3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005405">
        <property type="entry name" value="K_chnl_volt-dep_Kv3.4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021645">
        <property type="entry name" value="Shal-type_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011333">
        <property type="entry name" value="SKP1/BTB/POZ_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003131">
        <property type="entry name" value="T1-type_BTB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028325">
        <property type="entry name" value="VG_K_chnl"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027359">
        <property type="entry name" value="Volt_channel_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11537:SF126">
        <property type="entry name" value="POTASSIUM VOLTAGE-GATED CHANNEL SUBFAMILY C MEMBER 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11537">
        <property type="entry name" value="VOLTAGE-GATED POTASSIUM CHANNEL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02214">
        <property type="entry name" value="BTB_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00520">
        <property type="entry name" value="Ion_trans"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11601">
        <property type="entry name" value="Shal-type"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00169">
        <property type="entry name" value="KCHANNEL"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01583">
        <property type="entry name" value="KV34CHANNEL"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01491">
        <property type="entry name" value="KVCHANNEL"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01498">
        <property type="entry name" value="SHAWCHANNEL"/>
    </dbReference>
    <dbReference type="SMART" id="SM00225">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54695">
        <property type="entry name" value="POZ domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81324">
        <property type="entry name" value="Voltage-gated potassium channels"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="15" id="KW-0966">Cell projection</keyword>
    <keyword evidence="14" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="16" id="KW-0407">Ion channel</keyword>
    <keyword evidence="12" id="KW-0406">Ion transport</keyword>
    <keyword evidence="13 20" id="KW-0472">Membrane</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="9" id="KW-0630">Potassium</keyword>
    <keyword evidence="7" id="KW-0631">Potassium channel</keyword>
    <keyword evidence="4" id="KW-0633">Potassium transport</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="11" id="KW-0770">Synapse</keyword>
    <keyword evidence="6 20" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="10 20" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="2" id="KW-0813">Transport</keyword>
    <keyword evidence="8" id="KW-0851">Voltage-gated channel</keyword>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="213"/>
            <end position="232"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="269"/>
            <end position="290"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="302"/>
            <end position="323"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="370"/>
            <end position="391"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="412"/>
            <end position="429"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="441"/>
            <end position="470"/>
        </location>
    </feature>
    <feature type="domain" description="BTB" evidence="21">
        <location>
            <begin position="36"/>
            <end position="140"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="479"/>
            <end position="566"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="521"/>
            <end position="531"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="19">
        <location>
            <begin position="555"/>
            <end position="566"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022538"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022826"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022882"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022958"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00034111"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00034430"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="SMART" id="SM00225"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000017863.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="621" mass="69598" checksum="C6B8B8039BE06B0C" modified="2019-05-08" version="1">MISSVCVSSYRGRKSGNKPPSKTCLKEEMAKGEESDKITINVGGTRHETYKSTLRTLPGTRLAWLADPDAQSNFDFDGKSNEFFFDRHPGIFSYVLNYYRTGKLHCPADICGPLFEEELTYWGIDETDVEPCCWMTYRQHRDAEEALDIFESPEPGAGGEEAEEEGGREMALQRLGMDDRSPGAAGAKGGGGCCRNWQPRMWALFEDPYSSRAARVVAFASLFFILVSITTFCLETHEAFNRNVNVTETMVVGNTTRVLLTNKVETEPVLTYIEGVCVLWFTLEFLVRIICCPDKLLFIKKLLNIIDFVAILPFYLEVGLSGLSSKAARDVLGFLRVVRFVRILRIFKLTRHFVGLRVLGHTLRASTNEFLLLIIFLALGVLIFATMIYYAERIGAKTSDQSGTDQTHFKNIPIGFWWAVVTMTTLGYGDMYPKTWSGMLVGALCALAGVLTIAMPVPVIVNNFGMYYSLAMAKQKLPKKKKKHIPRPAPLDSPTYCKSEDNSPRNSTQSDTCPLAAEEVVVERKRSDSKQNGDANTVLSDEEQPLSSLHEEKRPMRRSSTRDKTKKAATCFLLNASDFSCTADGGIRKAKNNWPFPTSSSYPNLHQPRIQPMKLLRDLTE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HS25</accession>
    <name>A0A452HS25_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">COMM domain-containing protein 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="2">Belongs to the COMM domain-containing protein 1 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HS25"/>
    <dbReference type="Ensembl" id="ENSGAGT00000020407.1">
        <property type="protein sequence ID" value="ENSGAGP00000017889.1"/>
        <property type="gene ID" value="ENSGAGG00000013226.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005768">
        <property type="term" value="C:endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055070">
        <property type="term" value="P:copper ion homeostasis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:2000009">
        <property type="term" value="P:negative regulation of protein localization to cell surface"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1902306">
        <property type="term" value="P:negative regulation of sodium ion transmembrane transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031398">
        <property type="term" value="P:positive regulation of protein ubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032434">
        <property type="term" value="P:regulation of proteasomal ubiquitin-dependent protein catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd04749">
        <property type="entry name" value="Commd1_MURR1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017920">
        <property type="entry name" value="COMM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033776">
        <property type="entry name" value="COMMD1_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037351">
        <property type="entry name" value="Murr1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21199">
        <property type="entry name" value="COMM DOMAIN-CONTAINING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21199:SF1">
        <property type="entry name" value="COMM DOMAIN-CONTAINING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07258">
        <property type="entry name" value="COMM_domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF17221">
        <property type="entry name" value="COMMD1_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51269">
        <property type="entry name" value="COMM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="COMM" evidence="4">
        <location>
            <begin position="114"/>
            <end position="182"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="3">
        <location>
            <begin position="157"/>
            <end position="184"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00016551"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00093455"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS51269"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000017889.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="185" mass="20417" checksum="3B56C8BD5C9D49B4" modified="2019-05-08" version="1">LAGGKGSLGGLLSGIAQATYYGHAGITEELLRGQLYPEAAPHEFRALLAKMSGVLKSVASADMDLNQLEAFLTAQTKKQGGITADQAAVIAKFWKNHRTKIRESLINQSRWDNTLKNMNWRVDLKSQSRHVDQINTPVAIVEMELGKNGQESEFLCLEFDEAKVNQMLKKLSEIEESITALTQTT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HS27</accession>
    <name>A0A452HS27_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">WD repeat-containing protein 20</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="9">Interacts with USP12; promotes translocation of USP12/WDR20 to the plasma membrane. Component of the USP12/WDR20/WDR48 deubiquitinating complex. Interacts with USP46; contributes to the cytoplasmic localization of the USP46/WDR20 complex. Component of the USP12/DMWD/WDR48 deubiquitinating complex.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HS27"/>
    <dbReference type="Ensembl" id="ENSGAGT00000020367.1">
        <property type="protein sequence ID" value="ENSGAGP00000017851.1"/>
        <property type="gene ID" value="ENSGAGG00000013248.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:000101">
        <property type="entry name" value="WD repeat-containing protein 20 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036322">
        <property type="entry name" value="WD40_repeat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001680">
        <property type="entry name" value="WD40_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051362">
        <property type="entry name" value="WD_repeat_creC_regulators"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14107">
        <property type="entry name" value="WD REPEAT PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14107:SF5">
        <property type="entry name" value="WD REPEAT-CONTAINING PROTEIN 20"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00400">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00320">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50978">
        <property type="entry name" value="WD40 repeat-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50082">
        <property type="entry name" value="WD_REPEATS_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50294">
        <property type="entry name" value="WD_REPEATS_REGION"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0007">Acetylation</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5 11" id="KW-0853">WD repeat</keyword>
    <feature type="repeat" description="WD" evidence="11">
        <location>
            <begin position="263"/>
            <end position="295"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="408"/>
            <end position="457"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="481"/>
            <end position="501"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="12">
        <location>
            <begin position="423"/>
            <end position="435"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="443"/>
            <end position="457"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="12">
        <location>
            <begin position="482"/>
            <end position="495"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022574"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00062479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00070598"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000017851.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="589" mass="64853" checksum="E10D5002BAACE9A0" modified="2019-05-08" version="1">MLLSKMAAEGGGKEMNEIKTQFTTREGLYKLLSHSEYSRPNRVPFNSQGSNPVRVSFVNVNDQSGNGDRLCFNVGRELYFYIYKGVRKAADLSKPIDKRIYKGTQPTCHDFNHLTATAESVSLLVGFSAGQVQLIDPIKKETSKLFNEERLIDKSRVTCVKWVPGSESLFLVAHSSGNMYLYNVEHTCGTTAPHYQLLKQGEGFAVHTCKSKSTRNPLLKWTVGEGALNEFAFSPDGKFLACVSQDGFLRVFNFDSVELHGTMKSYFGGLLCVCWSPDGKYIVIGGEDDLVTVWSFVDCRVIARGHGHKSWVSVVAFDPYTTSVEESDPMEFSGSDEDFQDLLHFGRDRANSTQSRLSKRNSTDSRPVSVTYRFGSVGQDTQLCLWDLTEDILFPHQPLSRARTHTNVMNATSPPAGSGGTNSGSNGNSITTPGNSVPPPLPRSNSLPHSAVSNAGSKSSVMDGAIAAGVSKFATLSLHDRKERHHEKDHKRNHSMGHISSKSSDKLNLVTKTKTDPAKTLGTPLCPRMEDVPLLEPLICKKIAHERLTVLIFLEDCIVTACQEGFICTWARPGKVIMAFSSLEHGTVK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HS28</accession>
    <name>A0A452HS28_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Tripeptidyl-peptidase 2</fullName>
            <ecNumber evidence="3">3.4.14.10</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="9">Tripeptidyl aminopeptidase</fullName>
        </alternativeName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>Release of an N-terminal tripeptide from a polypeptide.</text>
            <dbReference type="EC" id="3.4.14.10"/>
        </reaction>
    </comment>
    <comment type="similarity">
        <text evidence="2 10">Belongs to the peptidase S8 family.</text>
    </comment>
    <dbReference type="EC" id="3.4.14.10" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452HS28"/>
    <dbReference type="Ensembl" id="ENSGAGT00000020392.1">
        <property type="protein sequence ID" value="ENSGAGP00000017875.1"/>
        <property type="gene ID" value="ENSGAGG00000011247.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004177">
        <property type="term" value="F:aminopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008240">
        <property type="term" value="F:tripeptidyl-peptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd04857">
        <property type="entry name" value="Peptidases_S8_Tripeptidyl_Aminopeptidase_II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.710">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.3170">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.3080">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.200">
        <property type="entry name" value="Peptidase S8/S53 domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000209">
        <property type="entry name" value="Peptidase_S8/S53_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036852">
        <property type="entry name" value="Peptidase_S8/S53_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022398">
        <property type="entry name" value="Peptidase_S8_His-AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023828">
        <property type="entry name" value="Peptidase_S8_Ser-AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050131">
        <property type="entry name" value="Peptidase_S8_subtilisin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015500">
        <property type="entry name" value="Peptidase_S8_subtilisin-rel"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034051">
        <property type="entry name" value="TPP_II_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022232">
        <property type="entry name" value="TPPII_C_art"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046939">
        <property type="entry name" value="TPPII_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048384">
        <property type="entry name" value="TPPII_GBD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048383">
        <property type="entry name" value="TPPII_Ig-like-1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022229">
        <property type="entry name" value="TPPII_Ig-like-2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046940">
        <property type="entry name" value="TPPII_Ig-like_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43806">
        <property type="entry name" value="PEPTIDASE S8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43806:SF14">
        <property type="entry name" value="TRIPEPTIDYL-PEPTIDASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00082">
        <property type="entry name" value="Peptidase_S8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12580">
        <property type="entry name" value="TPPII"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12583">
        <property type="entry name" value="TPPII_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21316">
        <property type="entry name" value="TPPII_GBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21223">
        <property type="entry name" value="TPPII_Ig-like-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00723">
        <property type="entry name" value="SUBTILISIN"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52743">
        <property type="entry name" value="Subtilisin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51892">
        <property type="entry name" value="SUBTILASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00137">
        <property type="entry name" value="SUBTILASE_HIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00138">
        <property type="entry name" value="SUBTILASE_SER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0031">Aminopeptidase</keyword>
    <keyword evidence="7 10" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="6 10" id="KW-0645">Protease</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8 10" id="KW-0720">Serine protease</keyword>
    <feature type="domain" description="Peptidase S8/S53" evidence="12">
        <location>
            <begin position="34"/>
            <end position="467"/>
        </location>
    </feature>
    <feature type="domain" description="Tripeptidyl-peptidase II first Ig-like" evidence="15">
        <location>
            <begin position="488"/>
            <end position="601"/>
        </location>
    </feature>
    <feature type="domain" description="Tripeptidyl-peptidase II galactose-binding" evidence="16">
        <location>
            <begin position="621"/>
            <end position="708"/>
        </location>
    </feature>
    <feature type="domain" description="Tripeptidyl peptidase II second Ig-like" evidence="13">
        <location>
            <begin position="744"/>
            <end position="930"/>
        </location>
    </feature>
    <feature type="domain" description="Tripeptidyl peptidase II C-terminal" evidence="14">
        <location>
            <begin position="977"/>
            <end position="1036"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="964"/>
            <end position="984"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="970"/>
            <end position="984"/>
        </location>
    </feature>
    <feature type="active site" description="Charge relay system" evidence="10">
        <location>
            <position position="43"/>
        </location>
    </feature>
    <feature type="active site" description="Charge relay system" evidence="10">
        <location>
            <position position="231"/>
        </location>
    </feature>
    <feature type="active site" description="Charge relay system" evidence="10">
        <location>
            <position position="416"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001910"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00011073"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012462"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00020244"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022438"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00032232"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF00082"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="Pfam" id="PF12580"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="Pfam" id="PF12583"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="Pfam" id="PF21223"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF21316"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000017875.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1216" mass="134791" checksum="B865D98703852C56" modified="2025-10-08" version="2">MAGWTHEPFPFHGLLPKKETGAASFLGRYPEYDGRGVLLAVLDTGVDPGAPGMQITTDGKPKIIDIIDTTGSGDVNTSTIVEPKDGEIVGLSGRTLKKERKEKQWDPVHRLLLAEACRKLDEFDAVHNSPSQTTKLMKEEFQSQVELLNSLEKKYNDPGPVYDCLVWHDGETWRACIDSSECGDFTKCTVLGTYRERQEYGSFGISEMLNYSVNIYDEGNLLSIVTSGGAHGTHVASIAAGYFPEEPERNGVAPGAQILAIKIGDTRLSTMETGTGLIRAMIEAMKYKCDLVNYSYGEATHWPNSGRICEVINEAVWKHNVIYVSSAGNNGPCLSTVGCPGGTTSSVIGVGAYVSPDMMVAEYSLREKLPANQYTWSSRGPSTDGALGVSISAPGGAIASVPNWTLRGTQLMNGTSMSSPNACGGIALVLSGLKANDVHYTVHSVRRALENTAVKAENIEVFAQGHGIIQVDKAYDYLIQNSSFTSNVGFTVTVGNNRGIYLRDPVQIAAPSDRGVGIEPVFPENTENTKRISLQLHLALTSNAAWVQCPSHLELMNQCRHINVRVDPRGLREGLHYTEVCGYDIAAPNAGPLFRVPITLIIPARIDESSHYDLSYTDVHFKPGQIRRHFIEVPQGATWAEVTVCSCSTDVTAKFVLHAVQLVKQKAYRSHEFYKFSSLPEKGSVTEAFPVLGGKTIEFCIARWWASLNDVSIDYTISFHGIVCATPQLNIHASEGIVRFDVQSTLKYEDIAPCINLKSWVQTLRPVSTKIKPLGSRDVLPNNRQLYEMILTYSFHQPKSGEVTPSCPLLCELLYESEFDSQLWIIFDQNKRQMGSGDAYPHQYSLKLEKGDYTIRLQIRHEQNSELDRIKDLPFVVSHRLSNTLSLDIYENHSLALLGKKKSNSLTLPPKHSHPFFVTSLPDDKIPKGAGPGCYLVGALTLSKTELGKKADVITVHYHLIPSPTKTKNGSKEKEREPEKEKDVKEEFAEALRDLKIQWMTKLDTSDVFNELKEAFPNHLPLYVARLHQLDSEKERLKRLDEIIEAANTVISHIDQTALAVYFVMKTDPRPDAATIKNDMEKQKSTLVDALCRKGSALADQLLHLQAQDGAVSSDAEGKDDDQESCIGVLSETFCETTKWTDLFDTKVLTFAYKHALANKMYGRGLKFATKLAEEKPTKENLKNCIQLMKLLGWTHCASFSENWLPVMYPPDYCVF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HS29</accession>
    <name>A0A452HS29_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">BTB domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>K(+)(in) = K(+)(out)</text>
            <dbReference type="Rhea" id="RHEA:29463"/>
            <dbReference type="ChEBI" id="CHEBI:29103"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="17">Presynaptic cell membrane</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HS29"/>
    <dbReference type="Ensembl" id="ENSGAGT00000020385.1">
        <property type="protein sequence ID" value="ENSGAGP00000017868.1"/>
        <property type="gene ID" value="ENSGAGG00000013255.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043679">
        <property type="term" value="C:axon terminus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032590">
        <property type="term" value="C:dendrite membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032809">
        <property type="term" value="C:neuronal cell body membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042734">
        <property type="term" value="C:presynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008076">
        <property type="term" value="C:voltage-gated potassium channel complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005251">
        <property type="term" value="F:delayed rectifier potassium channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001508">
        <property type="term" value="P:action potential"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051260">
        <property type="term" value="P:protein homooligomerization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.70:FF:000011">
        <property type="entry name" value="Potassium channel, voltage-gated Shaw-related subfamily C, member 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.350:FF:000014">
        <property type="entry name" value="Potassium channel, voltage-gated Shaw-related subfamily C, member 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.710.10:FF:000002">
        <property type="entry name" value="Potassium voltage-gated channel subfamily C member 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.70">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.710.10">
        <property type="entry name" value="Potassium Channel Kv1.1, Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.350">
        <property type="entry name" value="Voltage-gated potassium channels. Chain C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000210">
        <property type="entry name" value="BTB/POZ_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005821">
        <property type="entry name" value="Ion_trans_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003968">
        <property type="entry name" value="K_chnl_volt-dep_Kv"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003974">
        <property type="entry name" value="K_chnl_volt-dep_Kv3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005405">
        <property type="entry name" value="K_chnl_volt-dep_Kv3.4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021645">
        <property type="entry name" value="Shal-type_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011333">
        <property type="entry name" value="SKP1/BTB/POZ_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003131">
        <property type="entry name" value="T1-type_BTB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028325">
        <property type="entry name" value="VG_K_chnl"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027359">
        <property type="entry name" value="Volt_channel_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11537:SF126">
        <property type="entry name" value="POTASSIUM VOLTAGE-GATED CHANNEL SUBFAMILY C MEMBER 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11537">
        <property type="entry name" value="VOLTAGE-GATED POTASSIUM CHANNEL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02214">
        <property type="entry name" value="BTB_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00520">
        <property type="entry name" value="Ion_trans"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11601">
        <property type="entry name" value="Shal-type"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00169">
        <property type="entry name" value="KCHANNEL"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01583">
        <property type="entry name" value="KV34CHANNEL"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01491">
        <property type="entry name" value="KVCHANNEL"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01498">
        <property type="entry name" value="SHAWCHANNEL"/>
    </dbReference>
    <dbReference type="SMART" id="SM00225">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54695">
        <property type="entry name" value="POZ domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81324">
        <property type="entry name" value="Voltage-gated potassium channels"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="15" id="KW-0966">Cell projection</keyword>
    <keyword evidence="14" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="16" id="KW-0407">Ion channel</keyword>
    <keyword evidence="12" id="KW-0406">Ion transport</keyword>
    <keyword evidence="13 20" id="KW-0472">Membrane</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="9" id="KW-0630">Potassium</keyword>
    <keyword evidence="7" id="KW-0631">Potassium channel</keyword>
    <keyword evidence="4" id="KW-0633">Potassium transport</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="11" id="KW-0770">Synapse</keyword>
    <keyword evidence="6 20" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="10 20" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="2" id="KW-0813">Transport</keyword>
    <keyword evidence="8" id="KW-0851">Voltage-gated channel</keyword>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="213"/>
            <end position="232"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="269"/>
            <end position="290"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="302"/>
            <end position="323"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="370"/>
            <end position="391"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="412"/>
            <end position="429"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="441"/>
            <end position="470"/>
        </location>
    </feature>
    <feature type="domain" description="BTB" evidence="21">
        <location>
            <begin position="36"/>
            <end position="140"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="479"/>
            <end position="566"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="521"/>
            <end position="531"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="19">
        <location>
            <begin position="555"/>
            <end position="566"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022538"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022826"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022882"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022958"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00034111"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00034430"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="SMART" id="SM00225"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000017868.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="618" mass="69051" checksum="5621681FF1F2DF0D" modified="2019-05-08" version="1">MISSVCVSSYRGRKSGNKPPSKTCLKEEMAKGEESDKITINVGGTRHETYKSTLRTLPGTRLAWLADPDAQSNFDFDGKSNEFFFDRHPGIFSYVLNYYRTGKLHCPADICGPLFEEELTYWGIDETDVEPCCWMTYRQHRDAEEALDIFESPEPGAGGEEAEEEGGREMALQRLGMDDRSPGAAGAKGGGGCCRNWQPRMWALFEDPYSSRAARVVAFASLFFILVSITTFCLETHEAFNRNVNVTETMVVGNTTRVLLTNKVETEPVLTYIEGVCVLWFTLEFLVRIICCPDKLLFIKKLLNIIDFVAILPFYLEVGLSGLSSKAARDVLGFLRVVRFVRILRIFKLTRHFVGLRVLGHTLRASTNEFLLLIIFLALGVLIFATMIYYAERIGAKTSDQSGTDQTHFKNIPIGFWWAVVTMTTLGYGDMYPKTWSGMLVGALCALAGVLTIAMPVPVIVNNFGMYYSLAMAKQKLPKKKKKHIPRPAPLDSPTYCKSEDNSPRNSTQSDTCPLAAEEVVVERKRSDSKQNGDANTVLSDEEQPLSSLHEEKRPMRRSSTRDKTKKAATCFLLNASDFSCTADGGIRKGTDETQRHLVDRVAARVGEMEDRGTSMDT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HS30</accession>
    <name>A0A452HS30_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="7">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the CD225/Dispanin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HS30"/>
    <dbReference type="Ensembl" id="ENSGAGT00000020415.1">
        <property type="protein sequence ID" value="ENSGAGP00000017894.1"/>
        <property type="gene ID" value="ENSGAGG00000013267.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051607">
        <property type="term" value="P:defense response to virus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046597">
        <property type="term" value="P:host-mediated suppression of symbiont invasion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045071">
        <property type="term" value="P:negative regulation of viral genome replication"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035455">
        <property type="term" value="P:response to interferon-alpha"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035456">
        <property type="term" value="P:response to interferon-beta"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034341">
        <property type="term" value="P:response to type II interferon"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060337">
        <property type="term" value="P:type I interferon-mediated signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007593">
        <property type="entry name" value="CD225/Dispanin_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051517">
        <property type="entry name" value="IFITM_antiviral_protein"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13999">
        <property type="entry name" value="INTERFERON INDUCIBLE TRANSMEMBRANE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13999:SF4">
        <property type="entry name" value="INTERFERON-INDUCED TRANSMEMBRANE PROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04505">
        <property type="entry name" value="CD225"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 6" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 6" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 6" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="35"/>
            <end position="58"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="86"/>
            <end position="105"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004370"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000017894.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="132" mass="14084" checksum="3B103402FC6DBC82" modified="2019-05-08" version="1">SCSTVSSQHGAGRPGSQVHLHRTREVLPLQPPHCFVLLSLFSTFFCNVCCLGFTALVFSFKARDRKVLGDTDGAGSNGKTAKCLNIAVLVLGCSVVLVFIILLSVKLHRSSVRGEDQGLLYLNHPSGFSPAG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HS33</accession>
    <name>A0A452HS33_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Sulfotransferase</fullName>
            <ecNumber evidence="3">2.8.2.-</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1 3">Belongs to the sulfotransferase 1 family.</text>
    </comment>
    <dbReference type="EC" id="2.8.2.-" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452HS33"/>
    <dbReference type="Ensembl" id="ENSGAGT00000020397.1">
        <property type="protein sequence ID" value="ENSGAGP00000017880.1"/>
        <property type="gene ID" value="ENSGAGG00000013102.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008146">
        <property type="term" value="F:sulfotransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000863">
        <property type="entry name" value="Sulfotransferase_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11783">
        <property type="entry name" value="SULFOTRANSFERASE SULT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00685">
        <property type="entry name" value="Sulfotransfer_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 3" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Sulfotransferase" evidence="4">
        <location>
            <begin position="38"/>
            <end position="260"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00005771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="RuleBase" id="RU361155"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="Pfam" id="PF00685"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000017880.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="266" mass="30963" checksum="BC9EF6839D0EAB38" modified="2019-05-08" version="1">EGYLFFLLSNLLFPSCEYKDRFVFLYLHVCPLTLSSLGTNWMLEILSLIRSDGDPGWVCSVLNWDRVPWVEYGPGLEAARKYPPPRLLCFHLPVQLFPKSLQRSKAKIIYTLRCPKDVVVSLYHFSRIVRLFKDPDSLDCFLEDFLSGNEPCGGRELRHRWTIKFGCLPQDLQGSVRRICHFLGKELSEEQVAAVVENVSFQSMKGNKISNFSQMSDKYMDHQKGEFLRKGICGDWRNHLSEAQSQRFDTVFREWMQGLGMTFPWD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HS36</accession>
    <name>A0A452HS36_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Sulfotransferase</fullName>
            <ecNumber evidence="3">2.8.2.-</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1 3">Belongs to the sulfotransferase 1 family.</text>
    </comment>
    <dbReference type="EC" id="2.8.2.-" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452HS36"/>
    <dbReference type="Ensembl" id="ENSGAGT00000020336.1">
        <property type="protein sequence ID" value="ENSGAGP00000017824.1"/>
        <property type="gene ID" value="ENSGAGG00000013102.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008146">
        <property type="term" value="F:sulfotransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000863">
        <property type="entry name" value="Sulfotransferase_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11783">
        <property type="entry name" value="SULFOTRANSFERASE SULT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00685">
        <property type="entry name" value="Sulfotransfer_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 3" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Sulfotransferase" evidence="4">
        <location>
            <begin position="54"/>
            <end position="284"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00005771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="RuleBase" id="RU361155"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="Pfam" id="PF00685"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000017824.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="290" mass="33193" checksum="3A1E0CA8BCA62090" modified="2019-05-08" version="1">MHWDLNPEFQWRGRLWELWDSYGIATHSAMLQKSTLASVHGQVVSGPLTLSSLGTNWMLEILSLIRSDGDPGWVCSVLNWDRVPWVEYGPGLEAARKYPPPRLLCFHLPVQLFPKSLQRSKAKIIYTLRCPKDVVVSLYHFSRIVRLFKDPDSLDCFLEDFLSGNVLYGSWFHHVTGWMGLKANKNIFSITYEDGCLPQDLQGSVRRICHFLGKELSEEQVAAVVENVSFQSMKGNKISNFSQMSDKGCGLVVRGICGDWRNHLSEAQSQRFDTVFREWMQGLGMTFPWD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HS38</accession>
    <name>A0A452HS38_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">tripeptidyl-peptidase II</fullName>
            <ecNumber evidence="3">3.4.14.10</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>Release of an N-terminal tripeptide from a polypeptide.</text>
            <dbReference type="EC" id="3.4.14.10"/>
        </reaction>
    </comment>
    <comment type="similarity">
        <text evidence="2 8">Belongs to the peptidase S8 family.</text>
    </comment>
    <dbReference type="EC" id="3.4.14.10" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452HS38"/>
    <dbReference type="Ensembl" id="ENSGAGT00000020395.1">
        <property type="protein sequence ID" value="ENSGAGP00000017878.1"/>
        <property type="gene ID" value="ENSGAGG00000011247.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004177">
        <property type="term" value="F:aminopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008240">
        <property type="term" value="F:tripeptidyl-peptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd04857">
        <property type="entry name" value="Peptidases_S8_Tripeptidyl_Aminopeptidase_II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.710">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.20.25.690">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.3170">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.3080">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.200">
        <property type="entry name" value="Peptidase S8/S53 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000209">
        <property type="entry name" value="Peptidase_S8/S53_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036852">
        <property type="entry name" value="Peptidase_S8/S53_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022398">
        <property type="entry name" value="Peptidase_S8_His-AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023828">
        <property type="entry name" value="Peptidase_S8_Ser-AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050131">
        <property type="entry name" value="Peptidase_S8_subtilisin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015500">
        <property type="entry name" value="Peptidase_S8_subtilisin-rel"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034051">
        <property type="entry name" value="TPP_II_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046939">
        <property type="entry name" value="TPPII_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048384">
        <property type="entry name" value="TPPII_GBD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048383">
        <property type="entry name" value="TPPII_Ig-like-1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022229">
        <property type="entry name" value="TPPII_Ig-like-2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046940">
        <property type="entry name" value="TPPII_Ig-like_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43806">
        <property type="entry name" value="PEPTIDASE S8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43806:SF14">
        <property type="entry name" value="TRIPEPTIDYL-PEPTIDASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00082">
        <property type="entry name" value="Peptidase_S8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12580">
        <property type="entry name" value="TPPII"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21316">
        <property type="entry name" value="TPPII_GBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21223">
        <property type="entry name" value="TPPII_Ig-like-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00723">
        <property type="entry name" value="SUBTILISIN"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52743">
        <property type="entry name" value="Subtilisin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51892">
        <property type="entry name" value="SUBTILASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00137">
        <property type="entry name" value="SUBTILASE_HIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00138">
        <property type="entry name" value="SUBTILASE_SER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0031">Aminopeptidase</keyword>
    <keyword evidence="6 8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="5 8" id="KW-0645">Protease</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7 8" id="KW-0720">Serine protease</keyword>
    <feature type="domain" description="Peptidase S8/S53" evidence="10">
        <location>
            <begin position="34"/>
            <end position="506"/>
        </location>
    </feature>
    <feature type="domain" description="Tripeptidyl-peptidase II first Ig-like" evidence="12">
        <location>
            <begin position="527"/>
            <end position="640"/>
        </location>
    </feature>
    <feature type="domain" description="Tripeptidyl-peptidase II galactose-binding" evidence="13">
        <location>
            <begin position="660"/>
            <end position="747"/>
        </location>
    </feature>
    <feature type="domain" description="Tripeptidyl peptidase II second Ig-like" evidence="11">
        <location>
            <begin position="783"/>
            <end position="969"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="1014"/>
            <end position="1037"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="1022"/>
            <end position="1037"/>
        </location>
    </feature>
    <feature type="active site" description="Charge relay system" evidence="8">
        <location>
            <position position="43"/>
        </location>
    </feature>
    <feature type="active site" description="Charge relay system" evidence="8">
        <location>
            <position position="270"/>
        </location>
    </feature>
    <feature type="active site" description="Charge relay system" evidence="8">
        <location>
            <position position="455"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001910"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00011073"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012462"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022438"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="Pfam" id="PF00082"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF12580"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF21223"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="Pfam" id="PF21316"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000017878.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1192" mass="132127" checksum="FD66AEB47FFFD091" modified="2025-10-08" version="2">MAGWTHEPFPFHGLLPKKETGAASFLGRYPEYDGRGVLLAVLDTGVDPGAPGMQITTDGKPKIIDIIDTTGSGDVNTSTIVEPKDGEIVGLSGRTLKIPTNWVNPSGKYHIGIKNAYDFYPKALKERIQKERKEKQWDPVHRLLLAEACRKLDEFDAVHNSPSQHLSCFKFTTKLMKEEFQSQVELLNSLEKKYNDPGPVYDCLVWHDGETWRACIDSSECGDFTKCTVLGTYRERQEYGSFGISEMLNYSVNIYDEGNLLSIVTSGGAHGTHVASIAAGYFPEEPERNGVAPGAQILAIKIGDTRLSTMETGTGLIRAMIEAMKYKCDLVNYSYGEATHWPNSGRICEVINEAVWKHNVIYVSSAGNNGPCLSTVGCPGGTTSSVIGVGAYVSPDMMVAEYSLREKLPANQYTWSSRGPSTDGALGVSISAPGGAIASVPNWTLRGTQLMNGTSMSSPNACGGIALVLSGLKANDVHYTVHSVRRALENTAVKAENIEVFAQGHGIIQVDKAYDYLIQNSSFTSNVGFTVTVGNNRGIYLRDPVQIAAPSDRGVGIEPVFPENTENTKRISLQLHLALTSNAAWVQCPSHLELMNQCRHINVRVDPRGLREGLHYTEVCGYDIAAPNAGPLFRVPITLIIPARIDESSHYDLSYTDVHFKPGQIRRHFIEVPQGATWAEVTVCSCSTDVTAKFVLHAVQLVKQKAYRSHEFYKFSSLPEKGSVTEAFPVLGGKTIEFCIARWWASLNDVSIDYTISFHGIVCATPQLNIHASEGIVRFDVQSTLKYEDIAPCINLKSWVQTLRPVSTKIKPLGSRDVLPNNRQLYEMILTYSFHQPKSGEVTPSCPLLCELLYESEFDSQLWIIFDQNKRQMGSGDAYPHQYSLKLEKGDYTIRLQIRHEQNSELDRIKDLPFVVSHRLSNTLSLDIYENHSLALLGKKKSNSLTLPPKHSHPFFVTSLPDDKIPKGAGPGCYLVGALTLSKTELGKKAGQSAAKRQGKFKKDVITVHYHLIPSPTKTKNGSKEKEREPEKEKDVKEEFAEALRDLKIQWMTNDMEKQKSTLVDALCRKGSALADQLLHLQAQDGAVSSDAEGKDDDQESCIGVLSETFCETTKWTDLFDTKVLTFAYKHALANKMYGRGLKFATKLAEEKPTKENLKNCIQLMKLLGWTHCASFSENWLPVMYPPDYCVF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452I3Q3</accession>
    <name>A0A452I3Q3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">trypsin</fullName>
            <ecNumber evidence="11">3.4.21.4</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="10">
            <text>Preferential cleavage: Arg-|-Xaa, Lys-|-Xaa.</text>
            <dbReference type="EC" id="3.4.21.4"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Secreted</location>
            <location evidence="2">Extracellular space</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="3.4.21.4" evidence="11"/>
    <dbReference type="Ensembl" id="ENSGAGT00000025244.1">
        <property type="protein sequence ID" value="ENSGAGP00000022146.1"/>
        <property type="gene ID" value="ENSGAGG00000015915.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007586">
        <property type="term" value="P:digestion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00190">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.10:FF:000008">
        <property type="entry name" value="Cationic trypsin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.10:FF:000005">
        <property type="entry name" value="Serine protease 37"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.10.10">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009003">
        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043504">
        <property type="entry name" value="Peptidase_S1_PA_chymotrypsin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001314">
        <property type="entry name" value="Peptidase_S1A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050127">
        <property type="entry name" value="Serine_Proteases_S1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001254">
        <property type="entry name" value="Trypsin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018114">
        <property type="entry name" value="TRYPSIN_HIS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033116">
        <property type="entry name" value="TRYPSIN_SER"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24264:SF68">
        <property type="entry name" value="TRYPSIN-3-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24264">
        <property type="entry name" value="TRYPSIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00089">
        <property type="entry name" value="Trypsin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00722">
        <property type="entry name" value="CHYMOTRYPSIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00020">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50494">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50240">
        <property type="entry name" value="TRYPSIN_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00134">
        <property type="entry name" value="TRYPSIN_HIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00135">
        <property type="entry name" value="TRYPSIN_SER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0222">Digestion</keyword>
    <keyword evidence="9" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6 12" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="4 12" id="KW-0645">Protease</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0964">Secreted</keyword>
    <keyword evidence="7 12" id="KW-0720">Serine protease</keyword>
    <keyword evidence="13" id="KW-0732">Signal</keyword>
    <keyword evidence="8" id="KW-0865">Zymogen</keyword>
    <feature type="signal peptide" evidence="13">
        <location>
            <begin position="1"/>
            <end position="15"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019149066" description="trypsin" evidence="13">
        <location>
            <begin position="16"/>
            <end position="269"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase S1" evidence="14">
        <location>
            <begin position="24"/>
            <end position="267"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001913"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022757"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023145"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00036320"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00038868"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="RuleBase" id="RU363034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000022146.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="269" mass="29528" checksum="581FAA8D3CBB751A" modified="2019-05-08" version="1">MEIILFTLLLSATAASQLEDDDKIIGGYECSPHSQPWQVYFTYGSDYRWCGGSLINEWWIISAAHCYKRPLWVGSKLSVGPRPSTLVAHLGEHDTSADEGTEQHIQVAKAIQFPQYNQYNMDNDIMLVKLAQPARFGTYVQPIPISRSCPVAGMEYLVSGWGNILTSGVQYPDVLQCLNVPILSDSACRAVYPGXAVYPGRITKNMFCAGYIEGGKDSCQGDSGGPVVYNGELMGVVSWGIGCAQKNYPGVYTTVCNYVSWIEEVIANN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452I3R2</accession>
    <name>A0A452I3R2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">Cadherin domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">Cadherins are calcium-dependent cell adhesion proteins.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 14">Cell membrane</location>
            <topology evidence="1 14">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452I3R2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000025255.1">
        <property type="protein sequence ID" value="ENSGAGP00000022156.1"/>
        <property type="gene ID" value="ENSGAGG00000016273.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005912">
        <property type="term" value="C:adherens junction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016342">
        <property type="term" value="C:catenin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008013">
        <property type="term" value="F:beta-catenin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045296">
        <property type="term" value="F:cadherin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034332">
        <property type="term" value="P:adherens junction organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016339">
        <property type="term" value="P:calcium-dependent cell-cell adhesion via plasma membrane cell adhesion molecules"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000902">
        <property type="term" value="P:cell morphogenesis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0044331">
        <property type="term" value="P:cell-cell adhesion mediated by cadherin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007043">
        <property type="term" value="P:cell-cell junction assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007156">
        <property type="term" value="P:homophilic cell adhesion via plasma membrane adhesion molecules"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd11304">
        <property type="entry name" value="Cadherin_repeat"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.900.10:FF:000007">
        <property type="entry name" value="Cadherin 22"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.60:FF:000009">
        <property type="entry name" value="Cadherin 24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.60:FF:000012">
        <property type="entry name" value="Cadherin 24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.60:FF:000017">
        <property type="entry name" value="Cadherin 24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.60:FF:000014">
        <property type="entry name" value="Cadherin 8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.60">
        <property type="entry name" value="Cadherins"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.900.10">
        <property type="entry name" value="TCF3-CBD (Catenin binding domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039808">
        <property type="entry name" value="Cadherin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002126">
        <property type="entry name" value="Cadherin-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015919">
        <property type="entry name" value="Cadherin-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020894">
        <property type="entry name" value="Cadherin_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000233">
        <property type="entry name" value="Cadherin_Y-type_LIR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027397">
        <property type="entry name" value="Catenin-bd_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24027:SF311">
        <property type="entry name" value="CADHERIN-22"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24027">
        <property type="entry name" value="CADHERIN-23"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01049">
        <property type="entry name" value="CADH_Y-type_LIR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00028">
        <property type="entry name" value="Cadherin"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00205">
        <property type="entry name" value="CADHERIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00112">
        <property type="entry name" value="CA"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49313">
        <property type="entry name" value="Cadherin-like"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00232">
        <property type="entry name" value="CADHERIN_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50268">
        <property type="entry name" value="CADHERIN_2"/>
        <property type="match status" value="5"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8 13 14" id="KW-0106">Calcium</keyword>
    <keyword evidence="9 14" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="3" id="KW-0165">Cleavage on pair of basic residues</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="11 16" id="KW-0472">Membrane</keyword>
    <keyword evidence="5" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="4 14 16" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="10 16" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="16">
        <location>
            <begin position="730"/>
            <end position="751"/>
        </location>
    </feature>
    <feature type="domain" description="Cadherin" evidence="17">
        <location>
            <begin position="234"/>
            <end position="314"/>
        </location>
    </feature>
    <feature type="domain" description="Cadherin" evidence="17">
        <location>
            <begin position="339"/>
            <end position="383"/>
        </location>
    </feature>
    <feature type="domain" description="Cadherin" evidence="17">
        <location>
            <begin position="384"/>
            <end position="498"/>
        </location>
    </feature>
    <feature type="domain" description="Cadherin" evidence="17">
        <location>
            <begin position="499"/>
            <end position="603"/>
        </location>
    </feature>
    <feature type="domain" description="Cadherin" evidence="17">
        <location>
            <begin position="603"/>
            <end position="720"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022685"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="RuleBase" id="RU003318"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="RuleBase" id="RU004357"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50268"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000022156.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="921" mass="99880" checksum="14C9EDCB1D401E34" modified="2019-05-08" version="1">MSASSLKATLHAASQCTVVMALPGFSWTVLLLLCTLEQLGALAMHTEAEGPGLEQPGALAMHTEAEGPGLEQPGALAMHTEAEGPGLEQPGALAMHTEAEGPGLEQPGALAMHTEAEGPGLEQPGALAMHTETEGPGLEQPGALAMHTEAEGPGLEQPGALVMHTEAGGPGLEQLGALAMHTMHMEAGGPGLEQSGDSMLLAGGRVKRGWVWNQFFVVEEYTGTEPLYVGKIHSDSDEGDGSIKYTISGEGAGTIFLIDEVTGDIHATERLDREQKTFYTLRAQARDRQTNQLLEPESEFIIKVQDINDSEPRFLEGPYIGSVAELSPIGVIRTAVANLDRETQDRYEVVIRATDMAGQLGGLSGSTTVTIVITDVNDNPPRFPQKMYQFSIVETALVGTAIGRVKAEDSDVGENTDMMYQIKDEEGVQMFKVTTDSNTQEAIITVQKPLDYEAKMVHTVVVEALNKFVDPRFVDLGTFRDQTIVRVSVLDVDEPPEFRPPSDLMEVQEDAYVGSVVGVVTALDPDAANNPVRYAIDHNTDLERIFDIDANTGTIVTVKVLDRETAGWHNITVLAMEADNRSQMSRASLHIRILDVNDNPPELAMPYEAAVCEDAKPGQLIQTISVVDRDEPQSGHRFYFTLASEATNNHHFSLLDIQDNTAAVHTQRVGFNRQEQDIYFLPILVVDSGPPALSSTGTLTIRVCGCDSTGAIQSCNSTAYVMSATLSPGALIALLVCILILIVLVLLILTLKRHHKSHLTSEEDEDMRDNVIKYNDEGGGEQDTEAYDMSALRSLYDFSELKGSDGGGGPGDGGMTSNLPSELHALPQWFQNPDLDFSMFRDYISRKVDQADEDLSVPPYDSFQTYAFEGSDSSLPSLSSINTWSTSSEQDFSYLSSWGPHFRQLAALYAGRKGEEESEES</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452I3R5</accession>
    <name>A0A452I3R5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Agouti-signaling protein</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="8">Agouti switch protein</fullName>
        </alternativeName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452I3R5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000025258.1">
        <property type="protein sequence ID" value="ENSGAGP00000022158.1"/>
        <property type="gene ID" value="ENSGAGG00000016275.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031779">
        <property type="term" value="F:melanocortin receptor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005184">
        <property type="term" value="F:neuropeptide hormone activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009755">
        <property type="term" value="P:hormone-mediated signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032438">
        <property type="term" value="P:melanosome organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.760.10">
        <property type="entry name" value="Agouti domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007733">
        <property type="entry name" value="Agouti"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027300">
        <property type="entry name" value="Agouti_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036836">
        <property type="entry name" value="Agouti_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16551">
        <property type="entry name" value="AGOUTI RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16551:SF1">
        <property type="entry name" value="AGOUTI-SIGNALING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF05039">
        <property type="entry name" value="Agouti"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00792">
        <property type="entry name" value="Agouti"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57055">
        <property type="entry name" value="Agouti-related protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS60024">
        <property type="entry name" value="AGOUTI_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51150">
        <property type="entry name" value="AGOUTI_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6 9" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="7" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="5" id="KW-0960">Knottin</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0964">Secreted</keyword>
    <keyword evidence="4 11" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="11">
        <location>
            <begin position="1"/>
            <end position="29"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019554111" description="Agouti-signaling protein" evidence="11">
        <location>
            <begin position="30"/>
            <end position="132"/>
        </location>
    </feature>
    <feature type="domain" description="Agouti" evidence="12">
        <location>
            <begin position="93"/>
            <end position="132"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="63"/>
            <end position="88"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="69"/>
            <end position="80"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="9">
        <location>
            <begin position="93"/>
            <end position="108"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="9">
        <location>
            <begin position="100"/>
            <end position="114"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="9">
        <location>
            <begin position="107"/>
            <end position="125"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="9">
        <location>
            <begin position="111"/>
            <end position="132"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="9">
        <location>
            <begin position="116"/>
            <end position="123"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00017885"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022854"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00033432"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00494"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS51150"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000022158.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="132" mass="15084" checksum="088E7D07CD66683F" modified="2019-05-08" version="1">MMRINMKIENKNLFLALFLCSILLSTAYSHMIFEENRDTNLATSNKMKLPDLPSISIVELTKTSRKVSRKEAEKKKESKVKSHRPRPTLPANCVATWKNCKPPSPPCCDVCTFCHCRLFQTVCYCRVGNPNC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452I3R6</accession>
    <name>A0A452I3R6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">trypsin</fullName>
            <ecNumber evidence="11">3.4.21.4</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="10">
            <text>Preferential cleavage: Arg-|-Xaa, Lys-|-Xaa.</text>
            <dbReference type="EC" id="3.4.21.4"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Ca(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29108"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Secreted</location>
            <location evidence="2">Extracellular space</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="3.4.21.4" evidence="11"/>
    <dbReference type="Ensembl" id="ENSGAGT00000025257.1">
        <property type="protein sequence ID" value="ENSGAGP00000022159.1"/>
        <property type="gene ID" value="ENSGAGG00000015915.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007586">
        <property type="term" value="P:digestion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00190">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.10:FF:000008">
        <property type="entry name" value="Cationic trypsin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.10:FF:000166">
        <property type="entry name" value="Trypsin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.10.10">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009003">
        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043504">
        <property type="entry name" value="Peptidase_S1_PA_chymotrypsin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001314">
        <property type="entry name" value="Peptidase_S1A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050127">
        <property type="entry name" value="Serine_Proteases_S1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001254">
        <property type="entry name" value="Trypsin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018114">
        <property type="entry name" value="TRYPSIN_HIS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033116">
        <property type="entry name" value="TRYPSIN_SER"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24264:SF68">
        <property type="entry name" value="TRYPSIN-3-LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24264">
        <property type="entry name" value="TRYPSIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00089">
        <property type="entry name" value="Trypsin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00722">
        <property type="entry name" value="CHYMOTRYPSIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00020">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50494">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50240">
        <property type="entry name" value="TRYPSIN_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00134">
        <property type="entry name" value="TRYPSIN_HIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00135">
        <property type="entry name" value="TRYPSIN_SER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0222">Digestion</keyword>
    <keyword evidence="9" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6 12" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="4 12" id="KW-0645">Protease</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0964">Secreted</keyword>
    <keyword evidence="7 12" id="KW-0720">Serine protease</keyword>
    <keyword evidence="13" id="KW-0732">Signal</keyword>
    <keyword evidence="8" id="KW-0865">Zymogen</keyword>
    <feature type="signal peptide" evidence="13">
        <location>
            <begin position="1"/>
            <end position="15"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019304225" description="trypsin" evidence="13">
        <location>
            <begin position="16"/>
            <end position="251"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase S1" evidence="14">
        <location>
            <begin position="24"/>
            <end position="249"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001913"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022757"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023145"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00036320"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00038868"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="RuleBase" id="RU363034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000022159.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="251" mass="27531" checksum="0435927F74323276" modified="2019-05-08" version="1">MEIILFTLLLSATAASQLEDDDKIIGGYECSPHSQPWQVYFTYGSDYRWCGGSLINEWWIISAAHCYKRPLWVGSKLSVGPRPSTLVAHLGEHDTSADEGTEQHIQVAKAIQFPQYNQYNMDNDIMLVKSCPVAGMEYLVSGWGNILTSGVQYPDVLQCLNVPILSDSACRAVYPGXAVYPGRITKNMFCAGYIEGGKDSCQGDSGGPVVYNGELMGVVSWGIGCAQKNYPGVYTTVCNYVSWIEEVIANN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452I3R7</accession>
    <name>A0A452I3R7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Centrosomal protein of 78 kDa</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
            <location evidence="2">Cilium basal body</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Microtubule organizing center</location>
            <location evidence="1">Centrosome</location>
            <location evidence="1">Centriole</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="8">Belongs to the CEP78 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452I3R7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000025260.1">
        <property type="protein sequence ID" value="ENSGAGP00000022161.1"/>
        <property type="gene ID" value="ENSGAGG00000016274.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005814">
        <property type="term" value="C:centriole"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036064">
        <property type="term" value="C:ciliary basal body"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0044782">
        <property type="term" value="P:cilium organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030317">
        <property type="term" value="P:flagellated sperm motility"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000057">
        <property type="entry name" value="Centrosomal protein of 78 kDa"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.10.10:FF:000070">
        <property type="entry name" value="Centrosomal protein of 78 kDa"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.10.10">
        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026212">
        <property type="entry name" value="Cep78"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001611">
        <property type="entry name" value="Leu-rich_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032675">
        <property type="entry name" value="LRR_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24110">
        <property type="entry name" value="CENTROSOMAL PROTEIN OF 78 KDA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24110:SF3">
        <property type="entry name" value="CENTROSOMAL PROTEIN OF 78 KDA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13516">
        <property type="entry name" value="LRR_6"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR02062">
        <property type="entry name" value="CENTROSOME78"/>
    </dbReference>
    <dbReference type="SMART" id="SM00368">
        <property type="entry name" value="LRR_RI"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52047">
        <property type="entry name" value="RNI-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0966">Cell projection</keyword>
    <keyword evidence="5" id="KW-0969">Cilium</keyword>
    <keyword evidence="4" id="KW-0970">Cilium biogenesis/degradation</keyword>
    <keyword evidence="10" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="366"/>
            <end position="386"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="636"/>
            <end position="760"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="10">
        <location>
            <begin position="464"/>
            <end position="508"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="658"/>
            <end position="667"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="671"/>
            <end position="708"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="729"/>
            <end position="743"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="751"/>
            <end position="760"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004114"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004120"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022794"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023069"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00061070"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00069623"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000022161.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="760" mass="84709" checksum="34AC2E05F6FDF276" modified="2019-05-08" version="1">MIDSVKVRRQSTLDFYSHYEFLCAVQGSVPLKTVKADVNQGILELNADRLKASDWAPLLNAIRHNKTLTSIVIRSFHQQGLGESGADRPKTYFRRRIPAIRSKDMTTQLCKAIRGCLNISSALKNVELEGLLLRERDLIFLTKGLAKTSSLESLSLAHCPIGDGGLETICQSVRNSANIKSINFTGCSLTWRGAEHLANVLKHQAMKRHGEAWAESLRYRRPDLDCMAGLRRITLNCNTLVGDRGARAFAECLGEDLWLKALDMQQCGISNDGAKSLLDAFQTNTTLVILDVRKNPLIDHTLMKKLIERVLMNGNSASSEYKWLTSPSSKEVLKPKSKKRTIVLGSGRKGKATIRIVIWRHRGLASKKPIHPGKSPSDKDTYSPKPLPPGAYGFLPWRTAERAKRCRGFPVDGTQELPLKFQTGIPVKVTVESASASETEETEGTLEDIMHDPDLTESPVKTNVRRYQQLRTELEECLLKLKEERRARVKADERIMELEVENARLRNLNFSLSEVLHAQSVTNMILEDEGVLGSIETSFQKFHAFLDLLKDAGLGQLATIAGIDQSDFGLLGHPQMNSTVSKPAIVQKEKSYEEERQEQTQTSKVLVNNTCAIPREIQEFQVTVQQQQGLLGETEAQMDNQGKEEETSKNSRPSSSEKSVKLSEQTKGHSLKQSATNQCSFSDSNVNLKSKGSKKFSATSSEDSQNGSSKKHIYRTNETGIVNVAVRGDQNKHRTADHSRNDTEGFDSEIQESIHSMKSI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452I423</accession>
    <name>A0A452I423_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Thioredoxin domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the protein disulfide isomerase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452I423"/>
    <dbReference type="Ensembl" id="ENSGAGT00000025400.1">
        <property type="protein sequence ID" value="ENSGAGP00000022286.1"/>
        <property type="gene ID" value="ENSGAGG00000016308.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005783">
        <property type="term" value="C:endoplasmic reticulum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003756">
        <property type="term" value="F:protein disulfide isomerase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006457">
        <property type="term" value="P:protein folding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd02997">
        <property type="entry name" value="PDI_a_PDIR"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.30.10:FF:000029">
        <property type="entry name" value="protein disulfide-isomerase A5 isoform X2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.30.10">
        <property type="entry name" value="Glutaredoxin"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051063">
        <property type="entry name" value="PDI"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046374">
        <property type="entry name" value="PDI_a_PDIR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036249">
        <property type="entry name" value="Thioredoxin-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017937">
        <property type="entry name" value="Thioredoxin_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013766">
        <property type="entry name" value="Thioredoxin_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45672:SF2">
        <property type="entry name" value="PROTEIN DISULFIDE-ISOMERASE A5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45672">
        <property type="entry name" value="PROTEIN DISULFIDE-ISOMERASE C17H9.14C-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00085">
        <property type="entry name" value="Thioredoxin"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00421">
        <property type="entry name" value="THIOREDOXIN"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52833">
        <property type="entry name" value="Thioredoxin-like"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00194">
        <property type="entry name" value="THIOREDOXIN_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51352">
        <property type="entry name" value="THIOREDOXIN_2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Thioredoxin" evidence="2">
        <location>
            <begin position="87"/>
            <end position="208"/>
        </location>
    </feature>
    <feature type="domain" description="Thioredoxin" evidence="2">
        <location>
            <begin position="209"/>
            <end position="337"/>
        </location>
    </feature>
    <feature type="domain" description="Thioredoxin" evidence="2">
        <location>
            <begin position="338"/>
            <end position="459"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006347"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="PROSITE" id="PS51352"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000022286.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="472" mass="54321" checksum="B13CD23CD65A0FB7" modified="2019-05-08" version="1">MLLLLLFSAAAAENPLRFLSDVAQEVKGRGTIAWIDCGDTESRKVCKKMKADPNSKEKGVELLHYKDGVFHTEYNRAVTLKSVVAFLKDPEGAPLWEEDPEAKDVVHVDSEKEFRRLLKKEDRPLLMMFYAPWCGVCKRMMPSYQQAATELKGTYVLAGMNVHSTEFEKIKEDYNVRGYPTICYFEKGKFLFHYENYGATAKDIAEWLQNPQPPQPQAPETPWADEENAVYHLTDEDFDKFIKEHSSVLVMFHAPWCGHCKKMKPEYEMAAEMLHVTSDSPGVLAAVDATVNKAVAERFHISGFPTLKYFQDGEEKYTLPQLRTKKKIVDWLQNPQAPPPPELAWEEKLTNVIHLAGEDFRESLKKKKHALVMFYAPWCPHCKNSIPHFTTAAELFKEDRKIAYAAVDCAKEQNHDLCKQEGVDGYPTFNYYNYGKFVEKYNGERGESGFATFMRTLRERDHERVGKKKDEL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452I430</accession>
    <name>A0A452I430_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Thioredoxin domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the protein disulfide isomerase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452I430"/>
    <dbReference type="Ensembl" id="ENSGAGT00000025409.1">
        <property type="protein sequence ID" value="ENSGAGP00000022294.1"/>
        <property type="gene ID" value="ENSGAGG00000016308.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005783">
        <property type="term" value="C:endoplasmic reticulum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003756">
        <property type="term" value="F:protein disulfide isomerase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006457">
        <property type="term" value="P:protein folding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd02997">
        <property type="entry name" value="PDI_a_PDIR"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.30.10">
        <property type="entry name" value="Glutaredoxin"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051063">
        <property type="entry name" value="PDI"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046374">
        <property type="entry name" value="PDI_a_PDIR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036249">
        <property type="entry name" value="Thioredoxin-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017937">
        <property type="entry name" value="Thioredoxin_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013766">
        <property type="entry name" value="Thioredoxin_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45672:SF2">
        <property type="entry name" value="PROTEIN DISULFIDE-ISOMERASE A5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45672">
        <property type="entry name" value="PROTEIN DISULFIDE-ISOMERASE C17H9.14C-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00085">
        <property type="entry name" value="Thioredoxin"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00421">
        <property type="entry name" value="THIOREDOXIN"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52833">
        <property type="entry name" value="Thioredoxin-like"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00194">
        <property type="entry name" value="THIOREDOXIN_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51352">
        <property type="entry name" value="THIOREDOXIN_2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0472">Membrane</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="2" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="12"/>
            <end position="38"/>
        </location>
    </feature>
    <feature type="domain" description="Thioredoxin" evidence="3">
        <location>
            <begin position="82"/>
            <end position="216"/>
        </location>
    </feature>
    <feature type="domain" description="Thioredoxin" evidence="3">
        <location>
            <begin position="217"/>
            <end position="345"/>
        </location>
    </feature>
    <feature type="domain" description="Thioredoxin" evidence="3">
        <location>
            <begin position="346"/>
            <end position="467"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006347"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS51352"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000022294.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="480" mass="54954" checksum="3A1EABBBCABA7CE5" modified="2025-10-08" version="2">MLLTPLDFLWLPLVLITITVDLQLLYLLIGLMLLLLLFSAAAAENPLRFLSDVAQEVKGRGTIAWIDCGDTESRKVCKKMKADPNSKEKGVELLHYKDGVFHTEYNRAVTLKSVVAFLKDPEGAPLWEEDPEAKDVVVCLWCGVCKRMMPSYQQAATELKGTYVLAGMNVHSTEFEKIKEDYNVRGYPTICYFEKGKFLFHYENYGATAKDIAEWLQNPQPPQPQAPETPWADEENAVYHLTDEDFDKFIKEHSSVLVMFHAPWCGHCKKMKPEYEMAAEMLHVTSDSPGVLAAVDATVNKAVAERFHISGFPTLKYFQDGEEKYTLPQLRTKKKIVDWLQNPQAPPPPELAWEEKLTNVIHLAGEDFRESLKKKKHALVMFYAPWCPHCKNSIPHFTTAAELFKEDRKIAYAAVDCAKEQNHDLCKQEGVDGYPTFNYYNYGKFVEKYNGERGESGFATFMRTLRERDHERVGKKKDEL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452I434</accession>
    <name>A0A452I434_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Thioredoxin domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the protein disulfide isomerase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452I434"/>
    <dbReference type="Ensembl" id="ENSGAGT00000025411.1">
        <property type="protein sequence ID" value="ENSGAGP00000022296.1"/>
        <property type="gene ID" value="ENSGAGG00000016308.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005783">
        <property type="term" value="C:endoplasmic reticulum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003756">
        <property type="term" value="F:protein disulfide isomerase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006457">
        <property type="term" value="P:protein folding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd02997">
        <property type="entry name" value="PDI_a_PDIR"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.30.10:FF:000029">
        <property type="entry name" value="protein disulfide-isomerase A5 isoform X2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.30.10">
        <property type="entry name" value="Glutaredoxin"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051063">
        <property type="entry name" value="PDI"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046374">
        <property type="entry name" value="PDI_a_PDIR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036249">
        <property type="entry name" value="Thioredoxin-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017937">
        <property type="entry name" value="Thioredoxin_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013766">
        <property type="entry name" value="Thioredoxin_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45672:SF2">
        <property type="entry name" value="PROTEIN DISULFIDE-ISOMERASE A5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45672">
        <property type="entry name" value="PROTEIN DISULFIDE-ISOMERASE C17H9.14C-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00085">
        <property type="entry name" value="Thioredoxin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00421">
        <property type="entry name" value="THIOREDOXIN"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52833">
        <property type="entry name" value="Thioredoxin-like"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00194">
        <property type="entry name" value="THIOREDOXIN_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51352">
        <property type="entry name" value="THIOREDOXIN_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0472">Membrane</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="2" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="6"/>
            <end position="31"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="2">
        <location>
            <begin position="38"/>
            <end position="57"/>
        </location>
    </feature>
    <feature type="domain" description="Thioredoxin" evidence="3">
        <location>
            <begin position="205"/>
            <end position="333"/>
        </location>
    </feature>
    <feature type="domain" description="Thioredoxin" evidence="3">
        <location>
            <begin position="334"/>
            <end position="455"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006347"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS51352"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000022296.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="468" mass="53643" checksum="8DA963864BF7EB2E" modified="2019-05-08" version="1">LTPLDFLWLPLVLITITVGLLCCSPVSLISYHVTYLQLLYLLIGLMLLLLLFSAAAAENPLRFLSDVAQEVKGRGTIAWIDCGDTESRKVCKKMKADPNSKEKGVELLHYKDGVFHTEYNRAVTLKSVVAFLKDPEGAPLWEEDPEAKDVVVCLWMNVHSTEFEKIKEDYNVRGYPTICYFEKGKFLFHYENYGATAKDIAEWLQNPQPPQPQAPETPWADEENAVYHLTDEDFDKFIKEHSSVLVMFHAPWCGHCKKMKPEYEMAAEMLHVTSDSPGVLAAVDATVNKAVAERFHISGFPTLKYFQDGEEKYTLPQLRTKKKIVDWLQNPQAPPPPELAWEEKLTNVIHLAGEDFRESLKKKKHALVMFYAPWCPHCKNSIPHFTTAAELFKEDRKIAYAAVDCAKEQNHDLCKQEGVDGYPTFNYYNYGKFVEKYNGERGESGFATFMRTLRERDHERVGKKKDEL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452I437</accession>
    <name>A0A452I437_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">PKD domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the VPS10-related sortilin family. SORCS subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452I437"/>
    <dbReference type="Ensembl" id="ENSGAGT00000025414.1">
        <property type="protein sequence ID" value="ENSGAGP00000022299.1"/>
        <property type="gene ID" value="ENSGAGG00000013961.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006892">
        <property type="term" value="P:post-Golgi vesicle-mediated transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd15482">
        <property type="entry name" value="Sialidase_non-viral"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.60.270:FF:000001">
        <property type="entry name" value="Sortilin related VPS10 domain containing receptor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000083">
        <property type="entry name" value="Sortilin-related VPS10 domain containing receptor 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.70.80:FF:000001">
        <property type="entry name" value="Sortilin-related VPS10 domain-containing receptor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.70.80">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.120.10.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.60.270">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000601">
        <property type="entry name" value="PKD_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035986">
        <property type="entry name" value="PKD_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031777">
        <property type="entry name" value="Sortilin_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031778">
        <property type="entry name" value="Sortilin_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006581">
        <property type="entry name" value="VPS10"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050310">
        <property type="entry name" value="VPS10-sortilin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12106">
        <property type="entry name" value="SORTILIN RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12106:SF8">
        <property type="entry name" value="VPS10 DOMAIN-CONTAINING RECEPTOR SORCS1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00801">
        <property type="entry name" value="PKD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15902">
        <property type="entry name" value="Sortilin-Vps10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15901">
        <property type="entry name" value="Sortilin_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00602">
        <property type="entry name" value="VPS10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF110296">
        <property type="entry name" value="Oligoxyloglucan reducing end-specific cellobiohydrolase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49299">
        <property type="entry name" value="PKD domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50093">
        <property type="entry name" value="PKD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7 10" id="KW-0472">Membrane</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="3 10" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 10" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="793"/>
            <end position="811"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="11">
        <location>
            <begin position="527"/>
            <end position="567"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="831"/>
            <end position="858"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010818"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50093"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000022299.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="858" mass="95962" checksum="850214FBD4248C09" modified="2019-05-08" version="1">MTACSPFLSDLIYVPIAKPATDWIVLEQWSARAHAAAERPFNAQYITCRMQNCTEATRSKPFPGYIDPDSLIVQDDYVFVQLTSGGRPHYYVSYRRSVFAQMKLPKYSLPKDMHVISTDENQVFAAVQEWNQNDTYNLYISDTRGVYFTLALENIKSSRGLEGNVMIDLYEVAGIKGMFLANKKIDSQVKTFITYNKGRDWRLLQAPDTDLRGNPVHCLLPYCSLHLHLKVSENPYTSGNIASRDTAPSIIVASGNIGTELSDNDISMFVSSDAGNTWRQIFEEEHSVLYLDQGGVLVAMKHTSLPIRHLWLSFDEGRSWSKYSFTSTPLFVDGVLGEPGEETLIMTVFGHFSHRSEWQLVKVDYKSIFDRRCAEEDYRPWQLHSQGEACIMGAKRIYRKRKSEKKCMQGKYAGAMASEPCVCTEADFDCDYGYERHTNGQCLPAFWYNPSSLSKDCSFGQSYLNSTGYRKVVSNNCTDGVREQYTAKPQQCPGKAPHGLRIITSDGKLTAEQGHNVTLMVQLEEVDFGDGIAVSYANLSSTEDGIKHVYQNVGIFRVTVQVENSLGLDSAVLYLHVTCPLEHVHLSLPFVTTKNKEVNATAVLWPSQVGTLTYIWWFGNNTEPLVTLEGSITFTFSVEGMNTITVQVSAGNTILQDTKTIAVYERFQSLRLSFSPNLDEYNPDIPEWRRDVSRVIKRALVEATSISSKHILVAVLPGLPTSAELFVLPCQEATGENKRPAEDLDQISEILIHKLNQNLVQFELKPGVRVLVHAAHLSAVPLVDLTPSHSGSAMLMLLSVVFVGLAVFVIYKFKRYDLPLPLPPLSTDSGLTSMTLEQPSHESPMFSSSRPPSPGPDV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452I438</accession>
    <name>A0A452I438_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">C3H1-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="9">Belongs to the muscleblind family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452I438"/>
    <dbReference type="Ensembl" id="ENSGAGT00000025418.1">
        <property type="protein sequence ID" value="ENSGAGP00000022303.1"/>
        <property type="gene ID" value="ENSGAGG00000016381.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005654">
        <property type="term" value="C:nucleoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043484">
        <property type="term" value="P:regulation of RNA splicing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1370.210:FF:000004">
        <property type="entry name" value="Muscleblind like splicing regulator 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1370.210:FF:000002">
        <property type="entry name" value="Muscleblind-like 1 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1370.210">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR054429">
        <property type="entry name" value="Znf-CCCH_Muscleblind-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000571">
        <property type="entry name" value="Znf_CCCH"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12675">
        <property type="entry name" value="MUSCLEBLIND-LIKE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12675:SF7">
        <property type="entry name" value="MUSCLEBLIND-LIKE PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00642">
        <property type="entry name" value="zf-CCCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22628">
        <property type="entry name" value="zf-CCCH_10"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14608">
        <property type="entry name" value="zf-CCCH_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00356">
        <property type="entry name" value="ZnF_C3H1"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50103">
        <property type="entry name" value="ZF_C3H1"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4 10" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="7 10" id="KW-0862">Zinc</keyword>
    <keyword evidence="6 10" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C3H1-type" evidence="11">
        <location>
            <begin position="13"/>
            <end position="41"/>
        </location>
    </feature>
    <feature type="domain" description="C3H1-type" evidence="11">
        <location>
            <begin position="47"/>
            <end position="73"/>
        </location>
    </feature>
    <feature type="domain" description="C3H1-type" evidence="11">
        <location>
            <begin position="209"/>
            <end position="237"/>
        </location>
    </feature>
    <feature type="domain" description="C3H1-type" evidence="11">
        <location>
            <begin position="245"/>
            <end position="271"/>
        </location>
    </feature>
    <feature type="zinc finger region" description="C3H1-type" evidence="10">
        <location>
            <begin position="13"/>
            <end position="41"/>
        </location>
    </feature>
    <feature type="zinc finger region" description="C3H1-type" evidence="10">
        <location>
            <begin position="47"/>
            <end position="73"/>
        </location>
    </feature>
    <feature type="zinc finger region" description="C3H1-type" evidence="10">
        <location>
            <begin position="209"/>
            <end position="237"/>
        </location>
    </feature>
    <feature type="zinc finger region" description="C3H1-type" evidence="10">
        <location>
            <begin position="245"/>
            <end position="271"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00038226"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50103"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000022303.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="410" mass="45156" checksum="ADA7CB36AD9F8256" modified="2019-05-08" version="1">MAVSVTPIRDTKWLTLEVCREFQRGTCSRPDTECKFAHPSKSCQVENGRVIACFDSLKGRCSRENCKYLHPPPHLKTQLEINGRNNLIQQKNMAMLAQQMQLANAMMPGAPLQPVNYMPQSFQKRGGEGSPYYIDIDTNYQYHTFKPMFSVAPSLATNASAAFNPYLGPVSPGLVPAEILPTAPMLVAGNPGVPVPAAAAAAAQKLMRTDRLEVCREYQRGNCNRGENDCRFAHPADSAMIDTNDNTVTVCMDYIKGRCSREKCKYFHPPAHLQAKIKAAQYQVNQAAAAQAAATAAAMGIPQAVLPPLPKRPALEKTNGATAVFNTGIFQYQQALANMQLQQHTAFLPPGKYLFSLCRAKYATKLVFSICNKLLKVSVLIVIDIYPVVVRCFLLRVRKGSLGITVFVIR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IDS2</accession>
    <name>A0A452IDS2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">GCF C-terminal domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the GCF family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IDS2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000029593.1">
        <property type="protein sequence ID" value="ENSGAGP00000026018.1"/>
        <property type="gene ID" value="ENSGAGG00000019007.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000398">
        <property type="term" value="P:mRNA splicing, via spliceosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012890">
        <property type="entry name" value="GCFC2-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022783">
        <property type="entry name" value="GCFC_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12214">
        <property type="entry name" value="GC-RICH SEQUENCE DNA-BINDING FACTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12214:SF4">
        <property type="entry name" value="INTRON LARGE COMPLEX COMPONENT GCFC2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07842">
        <property type="entry name" value="GCFC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0539">Nucleus</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="GCF C-terminal" evidence="6">
        <location>
            <begin position="298"/>
            <end position="451"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="16"/>
            <end position="46"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="246"/>
            <end position="267"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="4">
        <location>
            <begin position="129"/>
            <end position="170"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="5">
        <location>
            <begin position="246"/>
            <end position="258"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="Pfam" id="PF07842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000026018.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="576" mass="66675" checksum="64911C6C4ADE2A41" modified="2019-05-08" version="1">MQVACRKHQLARTQGDYIPLDASHRAPVSQKKACSDTESEDDSDHDTILHFAPKTKSLKQRMAEYVVPKSDGASEDDEIQNMWEEQQLKKAVRLPQGMNVDTSLHKSRPMKIKFDPSVSLPPVNLEIVKKRLNTRITSLQDLHRAHQREHEKYTQDIESSKSTIQELERSSNAALNYKFYRSMKTYVESLIDCLNEKLLHINELESAMQMLLQQQAITHLKRRQEDLKNESAYLITKYRTCRRQARESLGKSDHHEGMSSDDELSAAEKTDFQKNKDDILQESKKICEDVHADFCNIRNILLKFQQWRQKFPDTYYNAYISLCLPKLLSPLIRIQLIGWNPLEQNFIDLNEMPWFRAIEEFTDAKNMPQSKRNDDPDQTILPTIIEKTILPKITGFVEHVWDPLSTSQTVSLVQLCKNVFEEHSLSKNESSKAKEDLVNSIVSRMKKSIEEDLLSNILLWDGIILEDTLHELGLGKLLNRYLLLILLNAPPGPDNVEKCNKVVACLPERWFKELKSGSTLPQLANFSQHLLQSAHTLYKNNCRDLILLLVKVNALHVAEDFIEEYKLEYLKSIVRK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IDS3</accession>
    <name>A0A452IDS3_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="14">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the Su(H) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IDS3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000029504.1">
        <property type="protein sequence ID" value="ENSGAGP00000025937.1"/>
        <property type="gene ID" value="ENSGAGG00000018936.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000029508.1">
        <property type="protein sequence ID" value="ENSGAGP00000025941.1"/>
        <property type="gene ID" value="ENSGAGG00000018936.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000029518.1">
        <property type="protein sequence ID" value="ENSGAGP00000025951.1"/>
        <property type="gene ID" value="ENSGAGG00000018936.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005654">
        <property type="term" value="C:nucleoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005667">
        <property type="term" value="C:transcription regulator complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001228">
        <property type="term" value="F:DNA-binding transcription activator activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048513">
        <property type="term" value="P:animal organ development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000122">
        <property type="term" value="P:negative regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007221">
        <property type="term" value="P:positive regulation of transcription of Notch receptor target"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd01176">
        <property type="entry name" value="IPT_RBP-Jkappa"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.1450:FF:000001">
        <property type="entry name" value="Recombining binding protein suppressor of hairless"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.80.10.50:FF:000003">
        <property type="entry name" value="recombining binding protein suppressor of hairless"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000074">
        <property type="entry name" value="Recombining binding protein suppressor of hairless, putative"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.80.10.50">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.1450">
        <property type="entry name" value="LAG1, DNA binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015350">
        <property type="entry name" value="Beta-trefoil_DNA-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036358">
        <property type="entry name" value="BTD_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040159">
        <property type="entry name" value="CLS_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014756">
        <property type="entry name" value="Ig_E-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008967">
        <property type="entry name" value="p53-like_TF_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015351">
        <property type="entry name" value="RBP-J/Cbf11/Cbf12_DNA-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037095">
        <property type="entry name" value="RBP-J/Cbf11_DNA-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038007">
        <property type="entry name" value="RBP-Jkappa_IPT"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10665">
        <property type="entry name" value="RECOMBINING BINDING PROTEIN SUPPRESSOR OF HAIRLESS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09270">
        <property type="entry name" value="BTD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09271">
        <property type="entry name" value="LAG1-DNAbind"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF20144">
        <property type="entry name" value="TIG_SUH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01268">
        <property type="entry name" value="BTD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01267">
        <property type="entry name" value="LAG1_DNAbind"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF110217">
        <property type="entry name" value="DNA-binding protein LAG-1 (CSL)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81296">
        <property type="entry name" value="E set domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49417">
        <property type="entry name" value="p53-like transcription factors"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0010">Activator</keyword>
    <keyword evidence="7" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="5" id="KW-0914">Notch signaling pathway</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0678">Repressor</keyword>
    <keyword evidence="9" id="KW-0804">Transcription</keyword>
    <keyword evidence="6" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="RBP-J/Cbf11/Cbf12 DNA binding" evidence="12">
        <location>
            <begin position="12"/>
            <end position="143"/>
        </location>
    </feature>
    <feature type="domain" description="Beta-trefoil DNA-binding" evidence="13">
        <location>
            <begin position="144"/>
            <end position="293"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="433"/>
            <end position="465"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="433"/>
            <end position="446"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="447"/>
            <end position="465"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00009704"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022491"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022976"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="SMART" id="SM01267"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="SMART" id="SM01268"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000025937.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="465" mass="51964" checksum="786A7D43A201EF9F" modified="2019-05-08" version="1">MRNYLKERGDQTVLILHAKVAQKSYGNEKRFFCPPPCVYLMGSGWKKKKEQMERDGCSEQESQPCAFIGIGNSDQEMQQLNLEGKNYCTAKTLYISDSDKRKHFMLSVKMFYGNSDDIGVFLSKRIKVISKPSKKKQSLKNADLCIASGTKVALFNRLRSQTVSTRYLHVEGGNFHASSQQWGAFYIHLLDDDESEGEEFTVRDGYIHYGQTVKLVCSVTGMALPRLIIRKVDKQTALLDADDPVSQLHKCAFYLKDTERMYLCLSQERIIQFQATPCPKEPNKEMINDGASWTIISTDKAEYTFYEGMGPVHAPVTPVPVVESLQLNGGGDVAMLELTGQNFTPNLRVWFGDVEAETMYRCAESMLCVVPDISAFREGWRWVRQPVQVPVTLVRNDGIIYSTSLTFTYTPEPGPRPHCSAAGAILRANSSLMASSETNTNSEGSYTNVSTNPTNVTSSTATVVS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IDS7</accession>
    <name>A0A452IDS7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">non-specific serine/threonine protein kinase</fullName>
            <ecNumber evidence="2">2.7.11.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the protein kinase superfamily. STE Ser/Thr protein kinase family. STE20 subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000029602.1">
        <property type="protein sequence ID" value="ENSGAGP00000026026.1"/>
        <property type="gene ID" value="ENSGAGG00000019004.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000003">
        <property type="entry name" value="TRAF2 and NCK-interacting protein kinase isoform 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001180">
        <property type="entry name" value="CNH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051700">
        <property type="entry name" value="STE20_Ser-Thr_kinase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47096">
        <property type="entry name" value="MISSHAPEN LIKE KINASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47096:SF1">
        <property type="entry name" value="MISSHAPEN LIKE KINASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00780">
        <property type="entry name" value="CNH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00036">
        <property type="entry name" value="CNH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50219">
        <property type="entry name" value="CNH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0418">Kinase</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="4" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Protein kinase" evidence="7">
        <location>
            <begin position="1"/>
            <end position="234"/>
        </location>
    </feature>
    <feature type="domain" description="CNH" evidence="8">
        <location>
            <begin position="987"/>
            <end position="1274"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="221"/>
            <end position="292"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="343"/>
            <end position="385"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="447"/>
            <end position="815"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="233"/>
            <end position="252"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="6">
        <location>
            <begin position="262"/>
            <end position="280"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="456"/>
            <end position="471"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="480"/>
            <end position="490"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="548"/>
            <end position="559"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="574"/>
            <end position="592"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="600"/>
            <end position="617"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="674"/>
            <end position="703"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="723"/>
            <end position="737"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="749"/>
            <end position="767"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="6">
        <location>
            <begin position="774"/>
            <end position="787"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00008874"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00012513"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50219"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000026026.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1300" mass="148720" checksum="5A3CE946D253D7EB" modified="2019-05-08" version="1">MDVTGDEEEEIKQEINMLKKYSHHRNIATYYGAFIKKNPPGMDDQLWLVMEFCGAGSVTDLIKNTKGNTLKEEWIAYICREILRGLSHLHQHKVIHRDIKGQNVLLTENAEVKLVDFGVSAQLDRTVGRRNTFIGTPYWMAPEVIACDENPDATYDFKSDLWSLGITAIEMAEGAPPLCDMHPMRALFLIPRNPAPRLKSKKWSKKFQSFIESCLVKNHSQRPTTEQLMKHPFIRDQPNERQVRIQLKDHIDRTKKKRGEKDETEYEYSGSEEEEEENDSGEPSSILNLPGESTLRRDFLRLQLANKERSEALRRQQLEQQQRENEEHKRQLLAERQKRIEEQKEQRRRLEEQQRREKEMRKQQEREQRRHYEEQMRREEERRRAEHEQEYIRRQLEEEQRQLEILQQQLLQEQALLLEYKRKQLEEQRQAERLQRQLQQERDYLVSLQQQQQQRQDQRPAEKKPLYHYKEGMNASEKPAWAKEVEERSRLNRQSSPAMPHKAANRISDPNLPPRSESFSISGVQPARTPPMLRPVDPQIPHLVTVKSQGTSLSGSQSLHEQPSKGMSGFQEALTVTSHRTEMPRQNSDPTSENPPLPPRIEKFDRSSWLRQEEDVPPKVPQRTTSISPALARKNSPGNGTALGARLGSQPIRASNPDLRRTEPIIESPLQRTSSGSSSSSSTPSSQPSSQEGSQPGSQAGSSERNRVRGNTRPEGSPVLSHELSKMKQEDSRDVTRPSRPADLTALAKELRELRIEETNRPLKKVTDYSSSSEESESSEEEEEDGESETHDGTVAVSDIPRLIPTGVPGSNEPYNLGMVASHGLETSNADTFSSISREGTLMVRETTGEKKRSSHAASNGFAGHINLPDLVQQSHSPAGTPTEALGRVSTHAQDMDSVAEYGMGSSTKASFTPFVDTRVYQTSPTDEDEDDDEESSATALFTSELLRQEQAKLNEARKISVVNVNPTNIRPHSDTPEIRKYKKRFNSEILCAALWGVNLLVGTENGLMLLDRSGQGKVYNLINRRRFQQMDVLEGLNVLVTISGKKNKLRVYYLSWLRNRILHNDPEVEKKQGWITVGDLEGCIHYKVVKYERIKFLVIALKNAVEIYAWAPKPYHKFMAFKSFADLQHKPLLVDLTVEEGQRLKVIFGSHTGFHVIDVDSGNSYDIYIPSHIQGNITPHAIVILPKTDGMEMLVCYEDEGVYVNTYGRITKDVVLQWGEMPTSVAYIHSNQIMGWGEKAIEIRSVETGHLDGVFMHKRAQRLKFLCERNDKVFFASVRSGGSSQVFFMTLNRNSMMNW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IDT7</accession>
    <name>A0A452IDT7_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="15">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="10">
            <text>ATP + H2O = ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:13065"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000029562.1">
        <property type="protein sequence ID" value="ENSGAGP00000025991.1"/>
        <property type="gene ID" value="ENSGAGG00000013176.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000785">
        <property type="term" value="C:chromatin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140658">
        <property type="term" value="F:ATP-dependent chromatin remodeler activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003682">
        <property type="term" value="F:chromatin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042393">
        <property type="term" value="F:histone binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034728">
        <property type="term" value="P:nucleosome organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd18666">
        <property type="entry name" value="CD1_tandem_CHD1-2_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18661">
        <property type="entry name" value="CD2_tandem_CHD1-2_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18054">
        <property type="entry name" value="DEXHc_CHD2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18793">
        <property type="entry name" value="SF2_C_SNF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.60:FF:000106">
        <property type="entry name" value="Chromodomain-helicase-DNA-binding protein 2 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.50.40:FF:000008">
        <property type="entry name" value="Chromodomain-helicase-DNA-binding protein 2 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.50.40:FF:000014">
        <property type="entry name" value="Chromodomain-helicase-DNA-binding protein 2 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10810:FF:000007">
        <property type="entry name" value="Chromodomain-helicase-DNA-binding protein 2 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000130">
        <property type="entry name" value="Chromodomain-helicase-DNA-binding protein 2 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.50.40">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.60">
        <property type="entry name" value="Homeodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.10810">
        <property type="entry name" value="Tandem AAA-ATPase domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040793">
        <property type="entry name" value="CDH1_2_SANT_HL1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056302">
        <property type="entry name" value="CHD1-2/Hrp3_HTH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025260">
        <property type="entry name" value="CHD1-like_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016197">
        <property type="entry name" value="Chromo-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000953">
        <property type="entry name" value="Chromo/chromo_shadow_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023780">
        <property type="entry name" value="Chromo_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023779">
        <property type="entry name" value="Chromodomain_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014001">
        <property type="entry name" value="Helicase_ATP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001650">
        <property type="entry name" value="Helicase_C-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038718">
        <property type="entry name" value="SNF2-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049730">
        <property type="entry name" value="SNF2/RAD54-like_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000330">
        <property type="entry name" value="SNF2_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45623:SF19">
        <property type="entry name" value="CHROMODOMAIN-HELICASE-DNA-BINDING PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45623">
        <property type="entry name" value="CHROMODOMAIN-HELICASE-DNA-BINDING PROTEIN 3-RELATED-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18375">
        <property type="entry name" value="CDH1_2_SANT_HL1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13907">
        <property type="entry name" value="CHD1-like_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00385">
        <property type="entry name" value="Chromo"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00271">
        <property type="entry name" value="Helicase_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23588">
        <property type="entry name" value="HTH_CHD1_Hrp3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00176">
        <property type="entry name" value="SNF2-rel_dom"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00298">
        <property type="entry name" value="CHROMO"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00487">
        <property type="entry name" value="DEXDc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01176">
        <property type="entry name" value="DUF4208"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00490">
        <property type="entry name" value="HELICc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54160">
        <property type="entry name" value="Chromo domain-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00598">
        <property type="entry name" value="CHROMO_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50013">
        <property type="entry name" value="CHROMO_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51192">
        <property type="entry name" value="HELICASE_ATP_BIND_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51194">
        <property type="entry name" value="HELICASE_CTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="7" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="4" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="3" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="9" id="KW-0539">Nucleus</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <keyword evidence="8" id="KW-0804">Transcription</keyword>
    <keyword evidence="6" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="Chromo" evidence="12">
        <location>
            <begin position="28"/>
            <end position="120"/>
        </location>
    </feature>
    <feature type="domain" description="Chromo" evidence="12">
        <location>
            <begin position="145"/>
            <end position="207"/>
        </location>
    </feature>
    <feature type="domain" description="Helicase ATP-binding" evidence="13">
        <location>
            <begin position="247"/>
            <end position="417"/>
        </location>
    </feature>
    <feature type="domain" description="Helicase C-terminal" evidence="14">
        <location>
            <begin position="546"/>
            <end position="697"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1"/>
            <end position="29"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="828"/>
            <end position="866"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1248"/>
            <end position="1404"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="11">
        <location>
            <begin position="1"/>
            <end position="25"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="1248"/>
            <end position="1260"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="1261"/>
            <end position="1270"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="1274"/>
            <end position="1296"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="1315"/>
            <end position="1325"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="1336"/>
            <end position="1348"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="1358"/>
            <end position="1370"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="1371"/>
            <end position="1390"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00049360"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50013"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS51192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS51194"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000025991.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1404" mass="164068" checksum="73ADCA6A74CAB109" modified="2019-05-08" version="1">EDDDFETDSDDLIEMTGEGADEQQDNSETIEKVMDVRIGKKGATGASTTVYETEANGNPSADFDPEKDAGEVQYLIKWKGWSYIHSTWESEESLQQQKVKGLKKLENFKKKEDEIKQWLAKVTPEDVEYFSCQQELAAELNKQYQIVERVIAHSRKLSSNEPEYLCKWMGLPYAECSWEDEALISKKFQHCIDSFNHRNNSKTIPTRDCKVLKQRPRFVALKKQPSYIGGENLELRDYQLEGLNWLAHSWCKNNSVILADEMGLGKTIQTISFLSYLFYQHQLYGPFLIVVPLSTLTSWQREFEIWAPEINVVVYIGDMMSRNTIREYEWIHSQTKRLKFNALITTYEILLKDKTVLGSINWAFLGVDEAHRLKNDDSLLYKTLIDFKSNHRLLITGTPLQNSLKELWSLLHFIMPEKFEFWEDFEDDHGKGRENGYQSLHKVLEPFLLRRVKKDVEKSLPAKVEQILRVEMSAPQKQYYKWILTRNYKALSKGTRGSTSGFLNIVMELKKCCNHCYLIKPPEENERETGMEMLMSLIRSSGKLILLDKLLTRLRERGNRVLIFSQMVRMLDILAEYLAIKHYPFQRLDGSIKGEIRKQALDHFNAEGSEDFCFLLSTRAGGLGINLASADTVVIFDSDWNPQNDLQAQARAHRIGQKKQVNIYRLVTKGTVEEEIIERAKKKMVLDHLVIQRMDTTGRTVLDNNSGRSNSNPFNKEELTAILKFGAEDLFKEIEGEESEPQEMDIDEILRLAETRENEVSSSATDELLSQFKVANFATMEEEETELDERSQRDWDDIIPEEQRKKVEEEERQKELEEIYMLPRIRSSTKKVQIHRSSGSESETDDTDDDKRPKRRGRPRSVRKDTVEGFTDAEIRRSSDSFMILFFPLRLECIARDAELVDKSVADLKRLGELIHNSCVSAMQEYEEQLKENPGEGKGPGKRRGPTLKISGVQVNVKSIIQHEEEFEMLHKSIPTDPEEKRKYCLTCRVKAAHFDVDWGVEDDSRLLLGIYEHGYGNWELIKTDPELKLTDKILPVETDKKPQGKQLQTRVDYLLKLLKKELEKKETVKDGEEVSEGMAKCPWGGHWTLLRRKLPPAFLDHEILCLNGLMRPVKKALKQLDKPDKGLTVQEQLEHTRNCLLKIGDRISECLKAYSDQEHIKLWRRNLWIFVSKFTEFDARKLHKLYKMAHKKRSQEEETHAQACVFLPADRGDWQRDRKFNYGGNSNQVWGGERHHQYEQHWYKDHHYGDRRSRGDPHRSSGNYRPNNLSRKRPYEQYNSDRDHRGHRDYYDRHHHDSKRRRSDEFRPQNYHQQEFRRMSDHRPPMGYHGPGPSDHYRSFHTDKLGDYKQPLPPPHSQLSDPRSPPSQKSPHESKSPLDHRSPLDRSLEQKNNPDYNWNVRKT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IDT8</accession>
    <name>A0A452IDT8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Paraneoplastic antigen Ma-like C-terminal domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IDT8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000029612.1">
        <property type="protein sequence ID" value="ENSGAGP00000026036.1"/>
        <property type="gene ID" value="ENSGAGG00000019017.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR026523">
        <property type="entry name" value="PNMA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048270">
        <property type="entry name" value="PNMA_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23095">
        <property type="entry name" value="PARANEOPLASTIC ANTIGEN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23095:SF17">
        <property type="entry name" value="PARANEOPLASTIC ANTIGEN MA1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14893">
        <property type="entry name" value="PNMA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Paraneoplastic antigen Ma-like C-terminal" evidence="2">
        <location>
            <begin position="100"/>
            <end position="234"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="257"/>
            <end position="280"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="257"/>
            <end position="266"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF14893"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000026036.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="280" mass="31345" checksum="F36131F0506E1D35" modified="2019-05-08" version="1">MAVHLLEDWCKGMHVDPKRTVFWEEGAFAVLCELPVVPSETAVEDSAWKLIATGSQSPPAPASDAEFLKKMSAVLGKEGETLADIPGLLGFDPRAPRLGAEWVVPDAEKRRCLAECLRGPALDVIHTVKLSNPRVGVKDCLEALDHTFGRTENSEDIYCKFLNARQQKGEKVSAYIQRLEKLLQRIIMRGAVAVEQMDRTRLAQIVRGIQYQNPILLHLRLRERQENPPSYSRLIKEVREEEEMQAAGEVLEVQSHQATGTASMQTPKALMVNPQEELTQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IE42</accession>
    <name>A0A452IE42_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="3">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IE42"/>
    <dbReference type="Ensembl" id="ENSGAGT00000029536.1">
        <property type="protein sequence ID" value="ENSGAGP00000025968.1"/>
        <property type="gene ID" value="ENSGAGG00000018960.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033179">
        <property type="term" value="C:proton-transporting V-type ATPase, V0 domain"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046961">
        <property type="term" value="F:proton-transporting ATPase activity, rotational mechanism"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.132.50">
        <property type="entry name" value="ATP synthase (C/AC39) subunit, domain 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036079">
        <property type="entry name" value="ATPase_csu/dsu_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002843">
        <property type="entry name" value="ATPase_V0-cplx_csu/dsu"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016727">
        <property type="entry name" value="ATPase_V0-cplx_dsu"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044911">
        <property type="entry name" value="V-type_ATPase_csu/dsu_dom_3"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11028">
        <property type="entry name" value="VACUOLAR ATP SYNTHASE SUBUNIT AC39"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01992">
        <property type="entry name" value="vATP-synt_AC39"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103486">
        <property type="entry name" value="V-type ATP synthase subunit C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0406">Ion transport</keyword>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0813">Transport</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000025968.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="104" mass="11827" checksum="587000EB4299291E" modified="2019-05-08" version="1">MSSFPELYFNVDSGYLEGLVRGFKAGVLSQGDYVNLVQCESLEDLKLHLQSTDYGNFLANEASPLTVSVIDDKLKEKMVVEFRHMRNHAYEPLASFLDFITCRN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IE44</accession>
    <name>A0A452IE44_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">TBC1 domain family member 5</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="11">Interacts with MAP1LC3A, MAP1LC3B, MAP1LC3C, GABARAP, GABARAPL1, GABARAPL2. Interacts with VPS29 and VPS35; indicative for an association with retromer CSC subcomplex. MAP1LC3A and VPS29 compete for binding to TBC1D5. Interacts with AP2M1; indicative for an association with the AP2 complex. Interacts with ULK1 and ATG13 (phosphorylated); indicative for an association with the activated ULK1-ATG13-FIP200 complex. Interacts with ATG9A; the interactions seems to be restricted to the AP2-clathrin-associated fraction of ATG9A.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasmic vesicle</location>
            <location evidence="1">Autophagosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Endosome membrane</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IE44"/>
    <dbReference type="Ensembl" id="ENSGAGT00000029708.1">
        <property type="protein sequence ID" value="ENSGAGP00000026121.1"/>
        <property type="gene ID" value="ENSGAGG00000018625.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005776">
        <property type="term" value="C:autophagosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010008">
        <property type="term" value="C:endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006914">
        <property type="term" value="P:autophagy"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.472.80:FF:000010">
        <property type="entry name" value="Putative TBC1 domain family member 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.8.270:FF:000011">
        <property type="entry name" value="TBC1 domain family member 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.8.270">
        <property type="entry name" value="putative rabgap domain of human tbc1 domain family member 14 like domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.472.80">
        <property type="entry name" value="Ypt/Rab-GAP domain of gyp1p, domain 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000195">
        <property type="entry name" value="Rab-GAP-TBC_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035969">
        <property type="entry name" value="Rab-GAP_TBC_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22957:SF337">
        <property type="entry name" value="TBC1 DOMAIN FAMILY MEMBER 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22957">
        <property type="entry name" value="TBC1 DOMAIN FAMILY MEMBER GTPASE-ACTIVATING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00566">
        <property type="entry name" value="RabGAP-TBC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00164">
        <property type="entry name" value="TBC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47923">
        <property type="entry name" value="Ypt/Rab-GAP domain of gyp1p"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50086">
        <property type="entry name" value="TBC_RABGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8" id="KW-0072">Autophagy</keyword>
    <keyword evidence="10" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="6" id="KW-0967">Endosome</keyword>
    <keyword evidence="4" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="9" id="KW-0472">Membrane</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="7" id="KW-0653">Protein transport</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <feature type="domain" description="Rab-GAP TBC" evidence="14">
        <location>
            <begin position="78"/>
            <end position="355"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="500"/>
            <end position="522"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="773"/>
            <end position="804"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="13">
        <location>
            <begin position="500"/>
            <end position="510"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="13">
        <location>
            <begin position="773"/>
            <end position="783"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004419"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004608"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023006"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00062017"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00072014"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50086"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000026121.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="804" mass="90963" checksum="E5604D119A330D8A" modified="2019-05-08" version="1">MYHSISENSHPLQTEEQEIGIDPLQGYLNKPEEDCWREGKGARGFLACAVPSIKEWEELFVNNNYLATIRLKGINGQLRSSRFRSVCWKLFLNVLPQDRSQWINKTSKLRTWYNKIKEIHITNPRKAGQQDLIINNPLSQDEGSLWNKFFQDKELRAMIEQDVTRTFPEMQYFQQESVRKILTDVLFCYARENEQLLYKQGMHELLAPIIFILHCDHQAFLHASEAAQPSEEMKALLNPGYLEHDAYAMFTHLMKTAEHWFSTFEHDGQKEKDAMMTPIPFARPQDLGPSIAIVTKVNQIQDHLLKKHDIELYMHLNRLEIAPQIYGLRWVRLLFGREFPLQDLLVVWDALFADSITLDLVDYIFVAMLLYIRDALISSNYQTCLGLLMHYPPIGDVHSLILRALFLRDPKKNPRPVTYQFQQNLDYYKACGADLMNKTRAKVAPLNINKVSNSLLNFGRKLIAPAMASGSPAGPAISASSSSFPSAVMPIRTIVEPLQHQHLQAQQQQQRMMKSESMPVQLSKGDVVTGSEAQMSVPAQNLTALQGQSSKNVSSSQSIESLSAGREYTGSPPLSASKKDFFFSTISRSRSHSKTMGRKESEEELETQISFLQGQLNDLEAMCKYCAKMMNTHLGNIQEVILQEHLEKEDEILISLIKDILKGSLRFNQSQLEAEENEQITIEDDHYCSSSQNKEMDDALKGSVMAKQHFLSAGNSGSRSTDDYILVSKEEEGNKGAVSEHELSLQAHKKMATKNEAPSHSSLVFSDPLMGSISASSSNLSSSPDDDSNNSKDSDFTIVSPLDI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IE45</accession>
    <name>A0A452IE45_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Adhesion G-protein coupled receptor G1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="21">G-protein coupled receptor 56</fullName>
        </alternativeName>
    </protein>
    <organism evidence="27 28">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="28" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="27" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="27" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="22">Heterodimer of 2 chains generated by proteolytic processing; the large extracellular N-terminal fragment (ADGRG1 NT) and the membrane-bound C-terminal fragment (ADGRG1-CT) predominantly remain associated and non-covalently linked. ADGRG1 NT self-associates in a trans-trans manner; the homophilic interaction enhances receptor signaling. Interacts with TGM2. Interacts with heparin; leading to the reduction of ADGRG1 shedding. Interacts with COL3A1. Part of a GPCR-tetraspanin complex at least consisting of ADGRG1, CD81, eventually CD9, and GNA11 in which CD81 is enhancing the association of ADGRG1 with GNA11.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cell membrane</location>
            <topology evidence="3">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Membrane raft</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Secreted</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IE45"/>
    <dbReference type="Ensembl" id="ENSGAGT00000029305.1">
        <property type="protein sequence ID" value="ENSGAGP00000025763.1"/>
        <property type="gene ID" value="ENSGAGG00000018800.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045121">
        <property type="term" value="C:membrane raft"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004930">
        <property type="term" value="F:G protein-coupled receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008201">
        <property type="term" value="F:heparin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007189">
        <property type="term" value="P:adenylate cyclase-activating G protein-coupled receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007166">
        <property type="term" value="P:cell surface receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1070.10:FF:000117">
        <property type="entry name" value="adhesion G-protein coupled receptor G1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.220.50">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1070.10">
        <property type="entry name" value="Rhodopsin 7-helix transmembrane proteins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057244">
        <property type="entry name" value="GAIN_B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046338">
        <property type="entry name" value="GAIN_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017981">
        <property type="entry name" value="GPCR_2-like_7TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000832">
        <property type="entry name" value="GPCR_2_secretin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003910">
        <property type="entry name" value="GPR1/GPR3/GPR5"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000203">
        <property type="entry name" value="GPS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040679">
        <property type="entry name" value="PLL"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12011">
        <property type="entry name" value="ADHESION G-PROTEIN COUPLED RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12011:SF318">
        <property type="entry name" value="ADHESION G-PROTEIN COUPLED RECEPTOR G1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00002">
        <property type="entry name" value="7tm_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01825">
        <property type="entry name" value="GPS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18587">
        <property type="entry name" value="PLL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00249">
        <property type="entry name" value="GPCRSECRETIN"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01422">
        <property type="entry name" value="GPR56ORPHANR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00303">
        <property type="entry name" value="GPS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50261">
        <property type="entry name" value="G_PROTEIN_RECEP_F2_4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50221">
        <property type="entry name" value="GAIN_B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="12" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="5" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="10" id="KW-0221">Differentiation</keyword>
    <keyword evidence="17" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="15" id="KW-0297">G-protein coupled receptor</keyword>
    <keyword evidence="19" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7" id="KW-0358">Heparin-binding</keyword>
    <keyword evidence="16 23" id="KW-0472">Membrane</keyword>
    <keyword evidence="13" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="18" id="KW-0675">Receptor</keyword>
    <keyword evidence="28" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0964">Secreted</keyword>
    <keyword evidence="9 24" id="KW-0732">Signal</keyword>
    <keyword evidence="20" id="KW-0807">Transducer</keyword>
    <keyword evidence="8 23" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="14 23" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="11" id="KW-0832">Ubl conjugation</keyword>
    <feature type="signal peptide" evidence="24">
        <location>
            <begin position="1"/>
            <end position="18"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019067002" description="Adhesion G-protein coupled receptor G1" evidence="24">
        <location>
            <begin position="19"/>
            <end position="674"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="23">
        <location>
            <begin position="388"/>
            <end position="413"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="23">
        <location>
            <begin position="425"/>
            <end position="444"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="23">
        <location>
            <begin position="459"/>
            <end position="477"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="23">
        <location>
            <begin position="497"/>
            <end position="519"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="23">
        <location>
            <begin position="557"/>
            <end position="579"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="23">
        <location>
            <begin position="591"/>
            <end position="613"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="23">
        <location>
            <begin position="619"/>
            <end position="641"/>
        </location>
    </feature>
    <feature type="domain" description="GAIN-B" evidence="25">
        <location>
            <begin position="224"/>
            <end position="381"/>
        </location>
    </feature>
    <feature type="domain" description="G-protein coupled receptors family 2 profile 2" evidence="26">
        <location>
            <begin position="389"/>
            <end position="644"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004285"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00019701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022674"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023040"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00023224"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00033134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00093505"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS50221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="26">
        <source>
            <dbReference type="PROSITE" id="PS50261"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000025763.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="28">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="674" mass="76299" checksum="8BAE4082CC7E7249" modified="2019-05-08" version="1">MMDRLLLALLFLLQGVDGSSHHTEDFRFCGERNQTKKSYITYQMRPESNIITIENSAEMLKISAPFEPNLVNFHLPDILGNYRLCLYWYQSDRIFNLTYGKNNYTLSTKAHPSFNFSNPSAPQNRNGVPVLSKVSYAYGRGLRNTSLSSAAVYSFSIRDISKVHEIEEELRNLENYMKNPNKAIGRKTSTRAPYGELLRLESDLEQVMFEGECKTFGTSTVRATVLKIGPSKASQDLSLMSELEVDREVHGFAVNLPRILFAQARGRGRNAERRVLLMDINSQALFQDQNGSRVLGEKVIGITVGNTSVSGLQQDVVLKFFHDQLPRNVTPQCVFWDMDSSHGHGSWKSDGCATEMGNNQTVCRCNHLTYFAVLMMSSPEIDYIHKEYLTIITYIGCIISAVASLFTIFLFCCSRRKHRDGTVNIYIHLNLLGAIFLLDVSFLMTEHLALIGSEAACKAGGMFLHLSLLGCLTWMGIEGYNLYRLVVEVFNTYVEHFMLKLCLVGWGLPIFIVSVIFLANQANYGLFHIEVFESLEKSTNATICWIKAPLIHNIVNLGFFSLVFLFNTAMLGAMVREILRQKNRGHELKHALVLLGLSLVLGIPWALAFFSFSSGSFRLIIIYLFTIINTLQGFLIFLWYWTMVLQVRKLKSYPSLNNSDSTRLPISTSRMNAD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IE47</accession>
    <name>A0A452IE47_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">SH3 domain-binding glutamic acid-rich-like protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="8">Appears to function as an adapter protein that bridges proteins together or proteins with mRNAs. May function as a ubiquitin ligase-substrate adapter. Additionally, associates with translating cytoplasmic ribosomes and may promote the expression of specific mRNAs.</text>
    </comment>
    <comment type="subunit">
        <text evidence="9">Monomer. Interacts with PFN1/Profilin-1. Interacts with ERBB2. Interacts with ATG12. Interacts with BECN1. Interacts with translating ribosomes.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytosol</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 10">Belongs to the SH3BGR family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IE47"/>
    <dbReference type="Ensembl" id="ENSGAGT00000029541.1">
        <property type="protein sequence ID" value="ENSGAGP00000025973.1"/>
        <property type="gene ID" value="ENSGAGG00000018967.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017124">
        <property type="term" value="F:SH3 domain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd03030">
        <property type="entry name" value="GRX_SH3BGR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.30.10:FF:000065">
        <property type="entry name" value="SH3 domain-binding glutamic acid-rich-like protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.30.10">
        <property type="entry name" value="Glutaredoxin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006993">
        <property type="entry name" value="Glut_rich_SH3-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051033">
        <property type="entry name" value="SH3BGR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036249">
        <property type="entry name" value="Thioredoxin-like_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12232:SF5">
        <property type="entry name" value="ADAPTER SH3BGRL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12232">
        <property type="entry name" value="SH3 DOMAIN-BINDING GLUTAMIC ACID-RICH-LIKE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04908">
        <property type="entry name" value="SH3BGR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF008142">
        <property type="entry name" value="SH3-bind_E-rich_L"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52833">
        <property type="entry name" value="Thioredoxin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-0472">Membrane</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0729">SH3-binding</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004514"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007764"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023036"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00045879"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00047101"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PIRNR" id="PIRNR008142"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000025973.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="114" mass="12796" checksum="520C8B6AE0E362ED" modified="2019-05-08" version="1">MVIRVYIASSSGSTAIKKQQQDVLGFLEANKIEFEEKDIAANEENRKWMRENVPEESRPVSGNPLPPRIFNDSRYLGDYEAFFEARENNAVYAFLGLTAPPGSKEAEALAKQQA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IE49</accession>
    <name>A0A452IE49_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">TBC1 domain family member 5</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="11">Interacts with MAP1LC3A, MAP1LC3B, MAP1LC3C, GABARAP, GABARAPL1, GABARAPL2. Interacts with VPS29 and VPS35; indicative for an association with retromer CSC subcomplex. MAP1LC3A and VPS29 compete for binding to TBC1D5. Interacts with AP2M1; indicative for an association with the AP2 complex. Interacts with ULK1 and ATG13 (phosphorylated); indicative for an association with the activated ULK1-ATG13-FIP200 complex. Interacts with ATG9A; the interactions seems to be restricted to the AP2-clathrin-associated fraction of ATG9A.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasmic vesicle</location>
            <location evidence="1">Autophagosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Endosome membrane</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IE49"/>
    <dbReference type="Ensembl" id="ENSGAGT00000029713.1">
        <property type="protein sequence ID" value="ENSGAGP00000026126.1"/>
        <property type="gene ID" value="ENSGAGG00000018625.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005776">
        <property type="term" value="C:autophagosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010008">
        <property type="term" value="C:endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006914">
        <property type="term" value="P:autophagy"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.472.80:FF:000010">
        <property type="entry name" value="Putative TBC1 domain family member 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.8.270:FF:000011">
        <property type="entry name" value="TBC1 domain family member 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.8.270">
        <property type="entry name" value="putative rabgap domain of human tbc1 domain family member 14 like domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.472.80">
        <property type="entry name" value="Ypt/Rab-GAP domain of gyp1p, domain 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000195">
        <property type="entry name" value="Rab-GAP-TBC_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035969">
        <property type="entry name" value="Rab-GAP_TBC_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22957:SF337">
        <property type="entry name" value="TBC1 DOMAIN FAMILY MEMBER 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22957">
        <property type="entry name" value="TBC1 DOMAIN FAMILY MEMBER GTPASE-ACTIVATING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00566">
        <property type="entry name" value="RabGAP-TBC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00164">
        <property type="entry name" value="TBC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47923">
        <property type="entry name" value="Ypt/Rab-GAP domain of gyp1p"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50086">
        <property type="entry name" value="TBC_RABGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8" id="KW-0072">Autophagy</keyword>
    <keyword evidence="10" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="6" id="KW-0967">Endosome</keyword>
    <keyword evidence="4" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="9" id="KW-0472">Membrane</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="7" id="KW-0653">Protein transport</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <feature type="domain" description="Rab-GAP TBC" evidence="14">
        <location>
            <begin position="82"/>
            <end position="358"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="1"/>
            <end position="57"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="503"/>
            <end position="525"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="765"/>
            <end position="796"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="13">
        <location>
            <begin position="1"/>
            <end position="14"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="13">
        <location>
            <begin position="39"/>
            <end position="55"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="13">
        <location>
            <begin position="503"/>
            <end position="513"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="13">
        <location>
            <begin position="765"/>
            <end position="775"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004419"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004608"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023006"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00062017"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00072014"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50086"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000026126.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="796" mass="90360" checksum="4A1E1DFCFDD5768F" modified="2019-05-08" version="1">MYHSISENSHPLQTEEQEIGIDPLQGYLNKPEEGDSRHRGGRRASRSRSRQRHSGSRKEWEELFVNNNYLATIRLKGINGQLRSSRFRSVCWKLFLNVLPQDRSQWINKTSKLRTWYNKIKEIHITNPRKAGQQDLIINNPLSQDEGSLWNKFFQDKELRAMIEQDVTRTFPEMQYFQQESVRKILTDVLFCYARENEQLLYKQGMHELLAPIIFILHCDHQAFLHASEAAQPSEEMKALLNPGYLEHDAYAMFTHLMKTAEHWFSTFEHDGQKKDAMMTPIPFARPQDLGPSIAIVTKVNQIQDHLLKKHDIELYMHLNRLEIAPQIYGLRWVRLLFGREFPLQDLLVVWDALFADSITLDLVDYIFVAMLLYIRDALISSNYQTCLGLLMHYPPIGDVHSLILRALFLRDPKKNPRPVTYQFQQNLDYYKACGADLMNKTRAKVAPLNINKVSNSLLNFGRKLIAPAMASGSPAGPAISASSSSFPSAVMPIRTIVEPLQHQHLQAQQQQQRMMKSESMPVQLSKGDVVTGSEAQMSVPAQNLTALQGQSSKNVSSSQSIESLSAGREYTGSPPLSASKKDFFFSTISRSRSHSKTMGRKESEEELETQISFLQGQLNDLEAMCKYCAKMMNTHLGNIQEVILQEHLEKEDEILISLIKDILKGSLRFNQSQLEAEENEQITIEDDHYCSSSQNKEMDDALKGSVMANRSTDDYILVSKEEEGNKGAVSEHELSLQAHKKMATKNEAPSHSSLVFSDPLMGSISASSSNLSSSPDDDSNNSKDSDFTIVSPLDI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IE50</accession>
    <name>A0A452IE50_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Ankyrin repeat domain-containing protein 17</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="13">Gene trap ankyrin repeat protein</fullName>
        </alternativeName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000029766.1">
        <property type="protein sequence ID" value="ENSGAGP00000026171.1"/>
        <property type="gene ID" value="ENSGAGG00000018637.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045087">
        <property type="term" value="P:innate immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd22502">
        <property type="entry name" value="KH-I_ANKRD17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000046">
        <property type="entry name" value="Ankyrin repeat and KH domain-containing protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000114">
        <property type="entry name" value="ankyrin repeat and KH domain-containing protein 1 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000062">
        <property type="entry name" value="ankyrin repeat domain-containing protein 17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000012">
        <property type="entry name" value="ankyrin repeat domain-containing protein 17 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000131">
        <property type="entry name" value="ankyrin repeat domain-containing protein 17 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1370.10:FF:000031">
        <property type="entry name" value="ankyrin repeat domain-containing protein 17 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000014">
        <property type="entry name" value="ankyrin repeat domain-containing protein 17 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000161">
        <property type="entry name" value="ankyrin repeat domain-containing protein 17 isoform X3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1370.10">
        <property type="entry name" value="K Homology domain, type 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051631">
        <property type="entry name" value="Ankyrin-KH/SAM_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047375">
        <property type="entry name" value="KH-I_ANKRD17"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004087">
        <property type="entry name" value="KH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004088">
        <property type="entry name" value="KH_dom_type_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036612">
        <property type="entry name" value="KH_dom_type_1_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23206:SF1">
        <property type="entry name" value="ANKYRIN REPEAT DOMAIN-CONTAINING PROTEIN 17"/>
        <property type="match status" value="1"/>
    </dbReference>
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</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IE52</accession>
    <name>A0A452IE52_9SAUR</name>
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        <submittedName>
            <fullName evidence="9">Uncharacterized protein</fullName>
        </submittedName>
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    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="5">Plays a role in the regulation of cell morphology and cytoskeletal organization. Required in the cortical actin filament dynamics and cell shape. Part of the striatin-interacting phosphatase and kinase (STRIPAK) complexes. STRIPAK complexes have critical roles in protein (de)phosphorylation and are regulators of multiple signaling pathways including Hippo, MAPK, nuclear receptor and cytoskeleton remodeling. Different types of STRIPAK complexes are involved in a variety of biological processes such as cell growth, differentiation, apoptosis, metabolism and immune regulation.</text>
    </comment>
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        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the STRIP family.</text>
    </comment>
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        <property type="gene ID" value="ENSGAGG00000019067.1"/>
    </dbReference>
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        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007010">
        <property type="term" value="P:cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040185">
        <property type="entry name" value="Far11/STRP"/>
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    <dbReference type="InterPro" id="IPR021819">
        <property type="entry name" value="Far11/STRP_C"/>
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        <property type="entry name" value="Far11/STRP_N"/>
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        <property type="entry name" value="PROTEIN REQUIRED FOR HYPHAL ANASTOMOSIS HAM-2"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="STRIATIN-INTERACTING PROTEIN 1"/>
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        <property type="entry name" value="DUF3402"/>
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        <property type="match status" value="1"/>
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    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Far11/STRP N-terminal" evidence="7">
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            <begin position="48"/>
            <end position="346"/>
        </location>
    </feature>
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            <begin position="443"/>
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        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="319"/>
            <end position="405"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Gly residues" evidence="6">
        <location>
            <begin position="1"/>
            <end position="13"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="30"/>
            <end position="43"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="339"/>
            <end position="360"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="6">
        <location>
            <begin position="361"/>
            <end position="374"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007062"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00046123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="SMART" id="SM01292"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="SMART" id="SM01293"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000026094.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="820" mass="94206" checksum="BEDC58FB76CC8FE4" modified="2019-05-08" version="1">MEPGGVGGCGGPGPLIVNNKQRGGLLPTGKGREFNRNQRKDSEGYSESPDLEFEYADTDRWSAELSELYSYTEGPEFQLNRKCFEEDFRVHVLDKKWTELDNSQHRTHAMRLLDGLEVTAREKRLKVARAILYVAQGTFGECSSEAEVQAWMRYNIFLLLEVGTFNALVELLNMEIDNSAACSSAVRKPAISLADSTDLRVLLNIMYLIIETVRQEAEGDKVEWKTMRQTFRAELGAPLYNNEPFSVMLFGMVTKFCSGHAPHFPMKKVLLLLWKTVLCTLGGFEELQAMKAEKREILGLPPLPEDSIKVIRNMRAASPPASASDLIEQQQKRGRREHKALIKQDNLDAFNERDPYKADDSREEEEENDDDNSLEGETFPLERDEVMPPPIQHPPTDRLTCPKGLPWAPKVREKDIEMFLESSRSKFIGYTLGSDTNTVVGLPRPIHESIRTLKLHKYTSIAEVQVQMEEEYLRSPLSGREEEVEQVPTETLYQGLLPSLPQYMIALLKILLAAAPTSKAKTDSINILADVLPEEMPTTVLQSMKLGVDVNRHKEIIVKAISAVLLLLLKHFKLNHVYQFEYMAQHLVFANCIPLILKFFNQNIMSYITAKNSISVLDYPYCVVHELPELTAESLEAGDNNQFCWRNLFSCINLLRILNKLTKWKHSRTMMLVVFKSAPILKRALKVKQAMMQLYVLKLLKVQTKYLGRQWRKSNMKTMSAIYQKVRHRLNDDWAYGNDLDARPWDFQAEECALRASIERFNSRRYDRAHSNPDFLPVDNCLQSVLGQRVELPEDFQMNYDLWLEREVFSRPISWEELLQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IE53</accession>
    <name>A0A452IE53_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Microtubule-associated protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cell membrane</location>
            <topology evidence="3">Peripheral membrane protein</topology>
            <orientation evidence="3">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cell projection</location>
            <location evidence="4">Axon</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cell projection</location>
            <location evidence="2">Dendrite</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1 18">Cytoplasm</location>
            <location evidence="1 18">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="5">Cytoplasm</location>
            <location evidence="5">Cytosol</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000029548.1">
        <property type="protein sequence ID" value="ENSGAGP00000025980.1"/>
        <property type="gene ID" value="ENSGAGG00000018969.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030424">
        <property type="term" value="C:axon"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030425">
        <property type="term" value="C:dendrite"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005874">
        <property type="term" value="C:microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008017">
        <property type="term" value="F:microtubule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000226">
        <property type="term" value="P:microtubule cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031175">
        <property type="term" value="P:neuron projection development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027324">
        <property type="entry name" value="MAP2/MAP4/Tau"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001084">
        <property type="entry name" value="MAP_tubulin-bd_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002955">
        <property type="entry name" value="Tau"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11501">
        <property type="entry name" value="MICROTUBULE-ASSOCIATED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11501:SF14">
        <property type="entry name" value="MICROTUBULE-ASSOCIATED PROTEIN TAU"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00418">
        <property type="entry name" value="Tubulin-binding"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01261">
        <property type="entry name" value="TAUPROTEIN"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00229">
        <property type="entry name" value="TAU_MAP_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51491">
        <property type="entry name" value="TAU_MAP_2"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="14" id="KW-0007">Acetylation</keyword>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="17" id="KW-0966">Cell projection</keyword>
    <keyword evidence="8 18" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="16 18" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="9" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="15" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0488">Methylation</keyword>
    <keyword evidence="11 18" id="KW-0493">Microtubule</keyword>
    <keyword evidence="10" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="12" id="KW-0677">Repeat</keyword>
    <keyword evidence="13" id="KW-0832">Ubl conjugation</keyword>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1"/>
            <end position="123"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="284"/>
            <end position="374"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="445"/>
            <end position="689"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="800"/>
            <end position="820"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="9"/>
            <end position="31"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="57"/>
            <end position="67"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="76"/>
            <end position="93"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="293"/>
            <end position="306"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="314"/>
            <end position="326"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="334"/>
            <end position="374"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="474"/>
            <end position="491"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="504"/>
            <end position="523"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="19">
        <location>
            <begin position="524"/>
            <end position="537"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="538"/>
            <end position="552"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="573"/>
            <end position="583"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="19">
        <location>
            <begin position="584"/>
            <end position="602"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="616"/>
            <end position="627"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="19">
        <location>
            <begin position="628"/>
            <end position="648"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004279"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004413"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004489"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004514"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="RuleBase" id="RU000686"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000025980.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="844" mass="91298" checksum="F1956E5FEAC99DAB" modified="2019-05-08" version="1">MKWNKRNRGRDNFCNRMTEQRQDFNMMEDHSMSQAKQIPSGYPLQIPVDDGSDEPISETSDAKSTPTMEDATAPLVEERAHEDRIAARQHVEIPEGTTAEEAGVGATPNLEDQAAGDGAQEELSSPRLWESVEYGIQEHVASEIQPEKQAGEVLPSTVVPLCEMQTGEQAAAIPARAEATIPVPPGTYEDFRAFEESQEAVSRTELWDRGGKEDIGVGKALADEAGHDVCTEEFAASAVSEFSQSVPRAGMFYHEKRAAQFDKWHPGVDDVGYEPVGNSHIQDKASHLYSRGKYKEEAVGHEKDEDRDIDETIPQDSPSSLGQQDSPLLEIAEEAVKEHSFPAEGQPRDNLAEISRETPVTEREGHKAGQILDEKRQWFSGKGYDDVTKMEPSESMHQVEVEANIIREGEISGSPLDTTKFPDRLNKDVADRSVSVQQEMGGMVQAMEKQEAPKKKPATRVSEKQVSRVPHLKAHIDSKDKDGIDIEEKKPKTSTPSSAKPLTDRSSITPQRPSSVSTTPLKNPSSPAESSVPASTPKRVSSITSRPASTGTKETKPKGPEMKSGMKMAAPRSATQTQKSPANTTRIPAKTPTAPKTPPSAAGRKEQRKPPTTAAKSEKXEPAKSGDRSGYSSPGSPGTPGSRSRTPSLPTPPNREPKKVAVVRTPPKSPASAKSRLQTSTAPMPDLKNVRSKIGSTENLKHQPGGGKVQIVYKPVDLSHVTSKCGSLGNIHHKPGGGQVEVKSEKLDFKEKVQSKIGSLDNITHVPGGGNKKIESHKLTFRENAKAKTDHGAEIVYKSPTISGDASPRRLSNVSSTGSINMVDSPQLATLADEVSASLAKQGL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IE54</accession>
    <name>A0A452IE54_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="15">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="10">
            <text>ATP + H2O = ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:13065"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000029718.1">
        <property type="protein sequence ID" value="ENSGAGP00000026131.1"/>
        <property type="gene ID" value="ENSGAGG00000013176.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000785">
        <property type="term" value="C:chromatin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0140658">
        <property type="term" value="F:ATP-dependent chromatin remodeler activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003682">
        <property type="term" value="F:chromatin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042393">
        <property type="term" value="F:histone binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0034728">
        <property type="term" value="P:nucleosome organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd18666">
        <property type="entry name" value="CD1_tandem_CHD1-2_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18661">
        <property type="entry name" value="CD2_tandem_CHD1-2_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18054">
        <property type="entry name" value="DEXHc_CHD2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18793">
        <property type="entry name" value="SF2_C_SNF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.60:FF:000106">
        <property type="entry name" value="Chromodomain-helicase-DNA-binding protein 2 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.50.40:FF:000008">
        <property type="entry name" value="Chromodomain-helicase-DNA-binding protein 2 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.50.40:FF:000014">
        <property type="entry name" value="Chromodomain-helicase-DNA-binding protein 2 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10810:FF:000007">
        <property type="entry name" value="Chromodomain-helicase-DNA-binding protein 2 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000130">
        <property type="entry name" value="Chromodomain-helicase-DNA-binding protein 2 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.50.40">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.60">
        <property type="entry name" value="Homeodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.10810">
        <property type="entry name" value="Tandem AAA-ATPase domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR040793">
        <property type="entry name" value="CDH1_2_SANT_HL1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056302">
        <property type="entry name" value="CHD1-2/Hrp3_HTH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025260">
        <property type="entry name" value="CHD1-like_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016197">
        <property type="entry name" value="Chromo-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000953">
        <property type="entry name" value="Chromo/chromo_shadow_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023780">
        <property type="entry name" value="Chromo_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023779">
        <property type="entry name" value="Chromodomain_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014001">
        <property type="entry name" value="Helicase_ATP-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001650">
        <property type="entry name" value="Helicase_C-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038718">
        <property type="entry name" value="SNF2-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049730">
        <property type="entry name" value="SNF2/RAD54-like_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000330">
        <property type="entry name" value="SNF2_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45623:SF19">
        <property type="entry name" value="CHROMODOMAIN-HELICASE-DNA-BINDING PROTEIN 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45623">
        <property type="entry name" value="CHROMODOMAIN-HELICASE-DNA-BINDING PROTEIN 3-RELATED-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF18375">
        <property type="entry name" value="CDH1_2_SANT_HL1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13907">
        <property type="entry name" value="CHD1-like_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00385">
        <property type="entry name" value="Chromo"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00271">
        <property type="entry name" value="Helicase_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23588">
        <property type="entry name" value="HTH_CHD1_Hrp3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00176">
        <property type="entry name" value="SNF2-rel_dom"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00298">
        <property type="entry name" value="CHROMO"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00487">
        <property type="entry name" value="DEXDc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01176">
        <property type="entry name" value="DUF4208"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00490">
        <property type="entry name" value="HELICc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54160">
        <property type="entry name" value="Chromo domain-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00598">
        <property type="entry name" value="CHROMO_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50013">
        <property type="entry name" value="CHROMO_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51192">
        <property type="entry name" value="HELICASE_ATP_BIND_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51194">
        <property type="entry name" value="HELICASE_CTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="7" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="4" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="3" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="9" id="KW-0539">Nucleus</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <keyword evidence="8" id="KW-0804">Transcription</keyword>
    <keyword evidence="6" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="Chromo" evidence="12">
        <location>
            <begin position="28"/>
            <end position="120"/>
        </location>
    </feature>
    <feature type="domain" description="Chromo" evidence="12">
        <location>
            <begin position="145"/>
            <end position="207"/>
        </location>
    </feature>
    <feature type="domain" description="Helicase ATP-binding" evidence="13">
        <location>
            <begin position="247"/>
            <end position="417"/>
        </location>
    </feature>
    <feature type="domain" description="Helicase C-terminal" evidence="14">
        <location>
            <begin position="540"/>
            <end position="691"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1"/>
            <end position="29"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="821"/>
            <end position="869"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1227"/>
            <end position="1332"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="11">
        <location>
            <begin position="1"/>
            <end position="25"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="825"/>
            <end position="834"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="1243"/>
            <end position="1253"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="1264"/>
            <end position="1276"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="1286"/>
            <end position="1298"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="1299"/>
            <end position="1318"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00049360"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50013"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS51192"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS51194"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000026131.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1332" mass="154996" checksum="6D6E65A7B505E7C8" modified="2019-05-08" version="1">EDDDFETDSDDLIEMTGEGADEQQDNSETIEKVMDVRIGKKGATGASTTVYETEANGNPSADFDPEKDAGEVQYLIKWKGWSYIHSTWESEESLQQQKVKGLKKLENFKKKEDEIKQWLAKVTPEDVEYFSCQQELAAELNKQYQIVERVIAHSRKLSSNEPEYLCKWMGLPYAECSWEDEALISKKFQHCIDSFNHRNNSKTIPTRDCKVLKQRPRFVALKKQPSYIGGENLELRDYQLEGLNWLAHSWCKNNSVILADEMGLGKTIQTISFLSYLFYQHQLYGPFLIVVPLSTLTSWQREFEIWAPEINVVVYIGDMMSRNTIREYEWIHSQTKRLKFNALITTYEILLKDKTVLGSINWAFLGVDEAHRLKNDDSLLYKTLIDFKSNHRLLITGTPLQNSLKELWSLLHFIMPEKFEFWEDFEDDHGKGRENGYQSLHKVLEPFLLRRVKKDVEKSLPAKVEQILRVEMSAPQKQYYKWILTRNYKALSKGTRGSTSGFLNIVMELKKCCNHCYLIKPPEENERETGMEMSSGKLILLDKLLTRLRERGNRVLIFSQMVRMLDILAEYLAIKHYPFQRLDGSIKGEIRKQALDHFNAEGSEDFCFLLSTRAGGLGINLASADTVVIFDSDWNPQNDLQAQARAHRIGQKKQVNIYRLVTKGTVEEEIIERAKKKMVLDHLVIQRMDTTGRTVLDNNSGRSNSNPFNKEELTAILKFGAEDLFKEIEGEESEPQEMDIDEILRLAETRENEVSSSATDELLSQFKVANFATMEEEETELDERSQRDWDDIIPEEQRKKVEEEERQKELEEIYMLPRIRSSTKKVHDSDAESKRRIHRSSGSESETDDTDDDKRPKRRGRPRSVRKDTVEGFTDAEIRRSSDSFMILFFPLRLECIARDAELVDKSVADLKRLGELIHNSCVSAMQEYEEQLKENPGEGKGPGKRRGPTLKISGVQVNVKSIIQHEEEFEMLHKSIPTDPEEKRKYCLTCRVKAAHFDVDWGVEDDSRLLLGIYEHGYGNWELIKTDPELKLTDKILPVETDKKPQGKQLQTRVDYLLKLLKKELEKKETVKDGEEVSEGMAKCPWGGHWTLLRRKLPPAFLDHEILCLNGLMRPVKKALKQLDKPDKGLTVQEQLEHTRNCLLKIGDRISECLKAYSDQEHIKLWRRNLWIFVSKFTEFDARKLHKLYKMAHKKRSQEEEVRSETCSLADIVLCLFFTSRHHHDSKRRRSDEFRPQNYHQQEFRRMSDHRPPMGYHGPGPSDHYRSFHTDKLGDYKQPLPPPHSQLSDPRSPPSQKSPHESKSPLDHRSPLDRSLEQKNNPDYNWNVRKT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IE56</accession>
    <name>A0A452IE56_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Sarcolemmal membrane-associated protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Associates with the striatin-interacting phosphatase and kinase (STRIPAK) core complex, forming the extended (SIKE1:SLMAP)STRIPAK complex. The (SIKE1:SLMAP)STRIPAK complex dephosphorylates STK3 leading to the inhibition of Hippo signaling and the control of cell growth. May play a role during myoblast fusion.</text>
    </comment>
    <comment type="subunit">
        <text evidence="17">Homodimer. Interacts with myosin. Interacts with SIKE1 and both associate with the STRIPAK core complex composed of PP2A catalytic and scaffolding subunits, the striatins (PP2A regulatory subunits), the striatin-associated proteins MOB4, STRIP1 and STRIP2, PDCD10 and members of the STE20 kinases, such as STK24 and STK26. Interacts (via FHA domain) with STK3 (when phosphorylated); the interaction associates STK3 with the STRIPAK complex.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="15">Cell membrane</location>
            <location evidence="15">Sarcolemma</location>
            <topology evidence="15">Single-pass type IV membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Microtubule organizing center</location>
            <location evidence="1">Centrosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Endoplasmic reticulum membrane</location>
            <topology evidence="2">Single-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="13">Mitochondrion membrane</location>
            <topology evidence="13">Single-pass type IV membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="16">Belongs to the SLMAP family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IE56"/>
    <dbReference type="Ensembl" id="ENSGAGT00000029318.1">
        <property type="protein sequence ID" value="ENSGAGP00000025773.1"/>
        <property type="gene ID" value="ENSGAGG00000018788.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031966">
        <property type="term" value="C:mitochondrial membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042383">
        <property type="term" value="C:sarcolemma"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0072659">
        <property type="term" value="P:protein localization to plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1900825">
        <property type="term" value="P:regulation of membrane depolarization during cardiac muscle cell action potential"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd21911">
        <property type="entry name" value="CC1_SLMAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd22679">
        <property type="entry name" value="FHA_SLMAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.200.20:FF:000003">
        <property type="entry name" value="sarcolemmal membrane-associated protein isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.200.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051176">
        <property type="entry name" value="Cent_Immune-Sig_Mod"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000253">
        <property type="entry name" value="FHA_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008984">
        <property type="entry name" value="SMAD_FHA_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15715">
        <property type="entry name" value="CENTROSOMAL PROTEIN OF 170 KDA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15715:SF22">
        <property type="entry name" value="SARCOLEMMAL MEMBRANE-ASSOCIATED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00498">
        <property type="entry name" value="FHA"/>
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    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49879">
        <property type="entry name" value="SMAD/FHA domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50006">
        <property type="entry name" value="FHA_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="9 19" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="4" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="12" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="7" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="11" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8" id="KW-1133">Transmembrane helix</keyword>
    <feature type="domain" description="FHA" evidence="21">
        <location>
            <begin position="28"/>
            <end position="85"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="20">
        <location>
            <begin position="436"/>
            <end position="461"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="19">
        <location>
            <begin position="172"/>
            <end position="199"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="19">
        <location>
            <begin position="265"/>
            <end position="387"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="19">
        <location>
            <begin position="490"/>
            <end position="517"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="20">
        <location>
            <begin position="436"/>
            <end position="445"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004389"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
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            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00046294"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00057671"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00060409"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00061687"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00066015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00074026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS50006"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000025773.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="536" mass="61438" checksum="AF8CC10DEBCB287D" modified="2019-05-08" version="1">MPSALAIFTCRPNSHPFQERHVYLDEPVKIGRSVARCRPAQNNATFDCKVLSRNHALVWFDHKTGKFYLQDTKSSNGTFINSQRLSRGSEESPPCEILSGDIIQFGVDVTENTRKVTHGCIVSTIKLFLPDGMEARLRSDVIHAPLPSPVDKVAANTPSMYSQELFQLSQYLQEALHREQMLEQKLATLQRLLAVTQEASDTSWQALIDEDRLLSRLEVMGNQLQACSKNQTEDSIRKELVALQEDKHNYETTAKESLRRVLQEKIEVVRKLSEVERSLSNTEDECTHLKEMNERTQEELRELANKYNGAVNEIKDLSDKLKVAERRQEEIQQKGQAEKKEMQHKINEMEEREQELQAKIEALQADNDFTNERLTALQVRLEHLQEKTLKEHNSLEHFLSKSGGDCTFIHQFIECQNKFIDEGRLTKVEETKLLRENQARAKESDLSDTLSPSKEKSSDDTTGKFGCKFLQNGFGKNVLLSDELQGAQSETETKQEIQQLHKELIEAQELARTSKQKCFELQEKGIFNLTVSLAYK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IE58</accession>
    <name>A0A452IE58_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Microtubule-associated protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cell membrane</location>
            <topology evidence="3">Peripheral membrane protein</topology>
            <orientation evidence="3">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cell projection</location>
            <location evidence="4">Axon</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cell projection</location>
            <location evidence="2">Dendrite</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1 18">Cytoplasm</location>
            <location evidence="1 18">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="5">Cytoplasm</location>
            <location evidence="5">Cytosol</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000029554.1">
        <property type="protein sequence ID" value="ENSGAGP00000025985.1"/>
        <property type="gene ID" value="ENSGAGG00000018969.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030424">
        <property type="term" value="C:axon"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030425">
        <property type="term" value="C:dendrite"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005874">
        <property type="term" value="C:microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008017">
        <property type="term" value="F:microtubule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000226">
        <property type="term" value="P:microtubule cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031175">
        <property type="term" value="P:neuron projection development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027324">
        <property type="entry name" value="MAP2/MAP4/Tau"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001084">
        <property type="entry name" value="MAP_tubulin-bd_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002955">
        <property type="entry name" value="Tau"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11501">
        <property type="entry name" value="MICROTUBULE-ASSOCIATED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11501:SF14">
        <property type="entry name" value="MICROTUBULE-ASSOCIATED PROTEIN TAU"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00418">
        <property type="entry name" value="Tubulin-binding"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01261">
        <property type="entry name" value="TAUPROTEIN"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00229">
        <property type="entry name" value="TAU_MAP_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51491">
        <property type="entry name" value="TAU_MAP_2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="14" id="KW-0007">Acetylation</keyword>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="17" id="KW-0966">Cell projection</keyword>
    <keyword evidence="8 18" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="16 18" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="9" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="15" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0488">Methylation</keyword>
    <keyword evidence="11 18" id="KW-0493">Microtubule</keyword>
    <keyword evidence="10" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="12" id="KW-0677">Repeat</keyword>
    <keyword evidence="13" id="KW-0832">Ubl conjugation</keyword>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1"/>
            <end position="123"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="284"/>
            <end position="374"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="445"/>
            <end position="623"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="765"/>
            <end position="785"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="9"/>
            <end position="31"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="57"/>
            <end position="67"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="76"/>
            <end position="93"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="293"/>
            <end position="306"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="314"/>
            <end position="326"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="334"/>
            <end position="374"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="474"/>
            <end position="495"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="507"/>
            <end position="517"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="19">
        <location>
            <begin position="518"/>
            <end position="536"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="550"/>
            <end position="561"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="19">
        <location>
            <begin position="562"/>
            <end position="582"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004279"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004413"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004489"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004514"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="RuleBase" id="RU000686"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000025985.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="809" mass="87803" checksum="2781F4D07C9EAA89" modified="2019-05-08" version="1">MKWNKRNRGRDNFCNRMTEQRQDFNMMEDHSMSQAKQIPSGYPLQIPVDDGSDEPISETSDAKSTPTMEDATAPLVEERAHEDRIAARQHVEIPEGTTAEEAGVGATPNLEDQAAGDGAQEELSSPRLWESVEYGIQEHVASEIQPEKQAGEVLPSTVVPLCEMQTGEQAAAIPARAEATIPVPPGTYEDFRAFEESQEAVSRTELWDRGGKEDIGVGKALADEAGHDVCTEEFAASAVSEFSQSVPRAGMFYHEKRAAQFDKWHPGVDDVGYEPVGNSHIQDKASHLYSRGKYKEEAVGHEKDEDRDIDETIPQDSPSSLGQQDSPLLEIAEEAVKEHSFPAEGQPRDNLAEISRETPVTEREGHKAGQILDEKRQWFSGKGYDDVTKMEPSESMHQVEVEANIIREGEISGSPLDTTKFPDRLNKDVADRSVSVQQEMGGMVQAMEKQEAPKKKPATRVSEKQVSRVPHLKAHIDSKDKDGIDIEEKKPKGPEMKSGMKMAAPRSATQTQKSPANTTRIPAKTPTAPKTPPSAAGRKEQRKPPTTAAKSEKXEPAKSGDRSGYSSPGSPGTPGSRSRTPSLPTPPNREPKKVAVVRTPPKSPASAKSRLQTSTAPMPDLKNVRSKIGSTENLKHQPGGGKVQIMNKKLDLSNIQSKCGSKDNIKHSPGGGSVQIVYKPVDLSHVTSKCGSLGNIHHKPGGGQVEVKSEKLDFKEKVQSKIGSLDNITHVPGGGNKKIESHKLTFRENAKAKTDHGAEIVYKSPTISGDASPRRLSNVSSTGSINMVDSPQLATLADEVSASLAKQGL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IE59</accession>
    <name>A0A452IE59_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">TBC1 domain family member 5</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="11">Interacts with MAP1LC3A, MAP1LC3B, MAP1LC3C, GABARAP, GABARAPL1, GABARAPL2. Interacts with VPS29 and VPS35; indicative for an association with retromer CSC subcomplex. MAP1LC3A and VPS29 compete for binding to TBC1D5. Interacts with AP2M1; indicative for an association with the AP2 complex. Interacts with ULK1 and ATG13 (phosphorylated); indicative for an association with the activated ULK1-ATG13-FIP200 complex. Interacts with ATG9A; the interactions seems to be restricted to the AP2-clathrin-associated fraction of ATG9A.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasmic vesicle</location>
            <location evidence="1">Autophagosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Endosome membrane</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IE59"/>
    <dbReference type="Ensembl" id="ENSGAGT00000029723.1">
        <property type="protein sequence ID" value="ENSGAGP00000026136.1"/>
        <property type="gene ID" value="ENSGAGG00000018625.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005776">
        <property type="term" value="C:autophagosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010008">
        <property type="term" value="C:endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006914">
        <property type="term" value="P:autophagy"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015031">
        <property type="term" value="P:protein transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.472.80:FF:000010">
        <property type="entry name" value="Putative TBC1 domain family member 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.8.270:FF:000011">
        <property type="entry name" value="TBC1 domain family member 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.8.270">
        <property type="entry name" value="putative rabgap domain of human tbc1 domain family member 14 like domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.472.80">
        <property type="entry name" value="Ypt/Rab-GAP domain of gyp1p, domain 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000195">
        <property type="entry name" value="Rab-GAP-TBC_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035969">
        <property type="entry name" value="Rab-GAP_TBC_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22957:SF337">
        <property type="entry name" value="TBC1 DOMAIN FAMILY MEMBER 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22957">
        <property type="entry name" value="TBC1 DOMAIN FAMILY MEMBER GTPASE-ACTIVATING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00566">
        <property type="entry name" value="RabGAP-TBC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00164">
        <property type="entry name" value="TBC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47923">
        <property type="entry name" value="Ypt/Rab-GAP domain of gyp1p"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51257">
        <property type="entry name" value="PROKAR_LIPOPROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50086">
        <property type="entry name" value="TBC_RABGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8" id="KW-0072">Autophagy</keyword>
    <keyword evidence="10" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="6" id="KW-0967">Endosome</keyword>
    <keyword evidence="4" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="9" id="KW-0472">Membrane</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="7" id="KW-0653">Protein transport</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <feature type="domain" description="Rab-GAP TBC" evidence="14">
        <location>
            <begin position="41"/>
            <end position="318"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="13">
        <location>
            <begin position="463"/>
            <end position="485"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="13">
        <location>
            <begin position="463"/>
            <end position="473"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004419"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004608"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022927"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023006"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00062017"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00072014"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS50086"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000026136.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="599" mass="68460" checksum="25FFFC0F0F9DDBAF" modified="2019-05-08" version="1">MLRVFNNVHASFLSCSKEWEELFVNNNYLATIRLKGINGQLRSSRFRSVCWKLFLNVLPQDRSQWINKTSKLRTWYNKIKEIHITNPRKAGQQDLIINNPLSQDEGSLWNKFFQDKELRAMIEQDVTRTFPEMQYFQQESVRKILTDVLFCYARENEQLLYKQGMHELLAPIIFILHCDHQAFLHASEAAQPSEEMKALLNPGYLEHDAYAMFTHLMKTAEHWFSTFEHDGQKEKDAMMTPIPFARPQDLGPSIAIVTKVNQIQDHLLKKHDIELYMHLNRLEIAPQIYGLRWVRLLFGREFPLQDLLVVWDALFADSITLDLVDYIFVAMLLYIRDALISSNYQTCLGLLMHYPPIGDVHSLILRALFLRDPKKNPRPVTYQFQQNLDYYKACGADLMNKTRAKVAPLNINKVSNSLLNFGRKLIAPAMASGSPAGPAISASSSSFPSAVMPIRTIVEPLQHQHLQAQQQQQRMMKSESMPVQLSKGDVVTGSEAQMSVPAQNLTALQGQSSKNVSSSQSIESLSAGREYTGSPPLSASKKDFFFSTISRSRSHSKTMGRKESEEELETQISFLQGQLNDLEAMCKYCAKMMNTHLGK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMJ1</accession>
    <name>A0A452IMJ1_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="7">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the major facilitator superfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMJ1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033125.1">
        <property type="protein sequence ID" value="ENSGAGP00000029163.1"/>
        <property type="gene ID" value="ENSGAGG00000021090.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1250.20:FF:000508">
        <property type="entry name" value="Sodium-dependent glucose transporter 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1250.20">
        <property type="entry name" value="MFS general substrate transporter like domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036259">
        <property type="entry name" value="MFS_trans_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23121">
        <property type="entry name" value="SODIUM-DEPENDENT GLUCOSE TRANSPORTER 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23121:SF9">
        <property type="entry name" value="SODIUM-DEPENDENT GLUCOSE TRANSPORTER 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103473">
        <property type="entry name" value="MFS general substrate transporter"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 6" id="KW-0472">Membrane</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 6" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 6" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="31"/>
            <end position="53"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="60"/>
            <end position="76"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="160"/>
            <end position="182"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="203"/>
            <end position="230"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="275"/>
            <end position="293"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="299"/>
            <end position="321"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="333"/>
            <end position="353"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="6">
        <location>
            <begin position="365"/>
            <end position="385"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008335"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029163.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="431" mass="46464" checksum="D234A366B31A7DEE" modified="2019-05-08" version="1">YVCFKPGMSIAVLGPTFQDLAANVNKNVSDIYYIFMGRSLGYLGGSVIGGIIFDCMNPHLLLGLTMLGTAVGLYAIPWCKRALLLTALMSVIGTSVGVLDTGNFALGAVVAPILTKMALSSFISPNFQTEDEKTNHSVPSDIPNALFGSALKLHSNVSFMWSYIVIGTYILIVSLLFVVLYSRTGLARDKAKSSLQKCQIAKYHYALLILLFLFFFCYVGAEVTYGSYIFNYAEVYADMKESEAASLNSLFWGAFAACRGLAICFATCLYPGTMILLSVIGSTVSSLFLALLSKHPISLWIGTAVYGASMATVFPSGISWIEQYTTIQGKSASLFVVGAALGEMCIPAVVGFLEGKLHNVPVMMYAALGTSVMTAVLFPVMYKLATSPSESKLKDGGGSEDQKALFSERYQKVTYNCFCASWTGALCFLLC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IMJ4</accession>
    <name>A0A452IMJ4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Lipase</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1 6">Belongs to the AB hydrolase superfamily. Lipase family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IMJ4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000033026.1">
        <property type="protein sequence ID" value="ENSGAGP00000029073.1"/>
        <property type="gene ID" value="ENSGAGG00000020969.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016788">
        <property type="term" value="F:hydrolase activity, acting on ester bonds"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016042">
        <property type="term" value="P:lipid catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1820:FF:000012">
        <property type="entry name" value="Lipase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1820">
        <property type="entry name" value="alpha/beta hydrolase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000073">
        <property type="entry name" value="AB_hydrolase_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029058">
        <property type="entry name" value="AB_hydrolase_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025483">
        <property type="entry name" value="Lipase_euk"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11005">
        <property type="entry name" value="LYSOSOMAL ACID LIPASE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00561">
        <property type="entry name" value="Abhydrolase_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000862">
        <property type="entry name" value="Steryl_ester_lip"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53474">
        <property type="entry name" value="alpha/beta-Hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="3 6" id="KW-0442">Lipid degradation</keyword>
    <keyword evidence="4" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0732">Signal</keyword>
    <feature type="domain" description="AB hydrolase-1" evidence="8">
        <location>
            <begin position="93"/>
            <end position="380"/>
        </location>
    </feature>
    <feature type="active site" description="Nucleophile" evidence="7">
        <location>
            <position position="187"/>
        </location>
    </feature>
    <feature type="active site" description="Charge relay system" evidence="7">
        <location>
            <position position="345"/>
        </location>
    </feature>
    <feature type="active site" description="Charge relay system" evidence="7">
        <location>
            <position position="374"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00010701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022963"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PIRNR" id="PIRNR000862"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PIRSR" id="PIRSR000862-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF00561"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000029073.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="399" mass="45656" checksum="F8DCE88437A17614" modified="2019-05-08" version="1">MKNLIRAARGILPPQARQAGCLWGLACRRSLVPRDISIPEQLIATRRYGGYQLICYEGYPSEEYEVLTEDGYYLKINRIPYGREKPTDEGPKPAVLLQHGLFGQGSNWIENLANNSLGFMLADAGYDVWIGNSRGTTWSQRHQKFSVGQEEFWDFSFHEMGIYDLQAMINFILQETGQKQLYYVGYSQGGSMGFIAFSSMPQLAQKVKMFFALAPAVTVGHSKSPVVKMLLMPEEALQDLLGKREFRWWSKTMREFAAKICCYDLPNRLCANAFFFAGGFNEKNINMSRMDVYTGHFPDETSIKNILHWAQTASETCIFVFQSVPPFYKIEDMTVPTAVWSGGEDWMVDPKDIDMLLPHITNLIYHKNISDWNHWDFTWGLDAAKRVYSEIIALIGKYP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IUT3</accession>
    <name>A0A452IUT3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">PAX-interacting protein 1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="6">PAX transactivation activation domain-interacting protein</fullName>
        </alternativeName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IUT3"/>
    <dbReference type="Ensembl" id="ENSGAGT00000035878.1">
        <property type="protein sequence ID" value="ENSGAGP00000031635.1"/>
        <property type="gene ID" value="ENSGAGG00000022720.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031981">
        <property type="term" value="C:nuclear lumen"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006974">
        <property type="term" value="P:DNA damage response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060261">
        <property type="term" value="P:positive regulation of transcription initiation by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd17714">
        <property type="entry name" value="BRCT_PAXIP1_rpt1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd17710">
        <property type="entry name" value="BRCT_PAXIP1_rpt2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd17711">
        <property type="entry name" value="BRCT_PAXIP1_rpt3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd17730">
        <property type="entry name" value="BRCT_PAXIP1_rpt4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd17712">
        <property type="entry name" value="BRCT_PAXIP1_rpt5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd18440">
        <property type="entry name" value="BRCT_PAXIP1_rpt6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10190:FF:000042">
        <property type="entry name" value="PAX interacting protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10190:FF:000046">
        <property type="entry name" value="PAX interacting protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.10190">
        <property type="entry name" value="BRCT domain"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001357">
        <property type="entry name" value="BRCT_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036420">
        <property type="entry name" value="BRCT_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051579">
        <property type="entry name" value="DDR_Transcriptional_Reg"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23196">
        <property type="entry name" value="PAX TRANSCRIPTION ACTIVATION DOMAIN INTERACTING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23196:SF1">
        <property type="entry name" value="PAX-INTERACTING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00533">
        <property type="entry name" value="BRCT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16589">
        <property type="entry name" value="BRCT_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12738">
        <property type="entry name" value="PTCB-BRCT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16770">
        <property type="entry name" value="RTT107_BRCT_5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00292">
        <property type="entry name" value="BRCT"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52113">
        <property type="entry name" value="BRCT domain"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50172">
        <property type="entry name" value="BRCT"/>
        <property type="match status" value="5"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-0227">DNA damage</keyword>
    <keyword evidence="4" id="KW-0539">Nucleus</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="BRCT" evidence="8">
        <location>
            <begin position="1"/>
            <end position="67"/>
        </location>
    </feature>
    <feature type="domain" description="BRCT" evidence="8">
        <location>
            <begin position="68"/>
            <end position="157"/>
        </location>
    </feature>
    <feature type="domain" description="BRCT" evidence="8">
        <location>
            <begin position="578"/>
            <end position="671"/>
        </location>
    </feature>
    <feature type="domain" description="BRCT" evidence="8">
        <location>
            <begin position="688"/>
            <end position="749"/>
        </location>
    </feature>
    <feature type="domain" description="BRCT" evidence="8">
        <location>
            <begin position="842"/>
            <end position="923"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="159"/>
            <end position="258"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="399"/>
            <end position="430"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="522"/>
            <end position="565"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="7">
        <location>
            <begin position="812"/>
            <end position="833"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="7">
        <location>
            <begin position="163"/>
            <end position="181"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023858"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00030146"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50172"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000031635.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1060" mass="120829" checksum="67223400A2304A54" modified="2019-05-08" version="1">MVIQLLKAGKAKEVSYNALASHIISEDGDNPEVGESREVFDLPVVKPSWVILSVQCGALLPVNGFSPESCQIFFGVTACLSQISSEDRSVLWALITFYGGDCQLSLNNKCTHLIVPEPKGEKYECACKRESIKIVTPDWVLDSISDKAKKEEISYHPRLIIYEEGEEEEEEDDAENEEQDSQNEASTDEKLSSPASSREGSPLGDQAFSPKPISADKTKGELMFDDSSDSSPEKQERNLNWTPAEVPQTAAAKRRLPQGKDSGLINLCANVPPVPGNILPPEIRGNIMASVQGTQGSERPEMMTTWSPAVRTLRNITNSADIQQINRPSNVAHILQSLSAPTKSLEQQVNHSQQGHTNAVLFSQVKLTPETHLLQQQQHQTQQQQQQHHPLLHLQPQQLMQLQQQQQQQPQIPQQPFPQQQSHQFSQQQQQVHQQHQFAQQLQFPQQQLHPQQLHRPQQQLQFQQQHALQQQLHQLQQQQLQQQQLQQIQQQNLQQQQLQQQQQQIQQQQLQQQHLQRLHQQHQQQQMQTQAPQHLTQTSQALQHQVPAQQLQHHQQQQQPQLQQHQLFGHDPAIEIPEEYFLLGCVFAIADYPEQMSDKQLLATWKRIIQAHGGTVDPTLTSRCTHLLCESQVSNMYAQAVRERKRCITAHWLNSILKKKKMIPPHRALHFPVAFPPGGKPCSQHIISVTGFVDSDRDDLKLMAYLAGAKYTGYLCRSNTVLICKEPTGLKYEKAKEWRIPCVNAQWICDILLGNFEALRQIQHSRYTVFNLQDPLSPTQHLVLNLLDAWRVPLKVSSELLMGVRLPLKPKQNESSLQPSSKRARIEDIPPPTKKLSPELTPMVLFTGFEPTQVQQYIKKLYILGGEVAESAQKCTHLVASKVTRTVKFLTAISVVKHIVTPEWLEECFKCQKFVDEQNFMLRDAEAEVLFCFSLEESLKRAQVAPLFKGKYFYITPGICPSLSTMKAIVDCAGGKVLSKQPSFRKLMEHKQNKSLSAIILISCENDLHLCREYFARGIDVHNAEFVLTGVLTQTLDYESYPFYMKTKKCDCWENVFIY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IUT4</accession>
    <name>A0A452IUT4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Sperm acrosome membrane-associated protein 3</fullName>
            <ecNumber evidence="5">3.2.1.17</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Lysozymes have primarily a bacteriolytic function; those in tissues and body fluids are associated with the monocyte-macrophage system and enhance the activity of immunoagents. Has strong bacteriolytic activity against M.luteus and V.cholerae, weak bacteriolytic activity against P.aeruginosa and no activity against A.hydrophila.</text>
    </comment>
    <comment type="function">
        <text evidence="13">Sperm surface membrane protein that may be involved in sperm-egg plasma membrane adhesion and fusion during fertilization. It could be a potential receptor for the egg oligosaccharide residue N-acetylglucosamine, which is present in the extracellular matrix over the egg plasma membrane. The processed form has no detectable bacteriolytic activity in vitro.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>Hydrolysis of (1-&gt;4)-beta-linkages between N-acetylmuramic acid and N-acetyl-D-glucosamine residues in a peptidoglycan and between N-acetyl-D-glucosamine residues in chitodextrins.</text>
            <dbReference type="EC" id="3.2.1.17"/>
        </reaction>
    </comment>
    <comment type="subunit">
        <text evidence="4">Interacts with ASTL.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 15">Belongs to the glycosyl hydrolase 22 family.</text>
    </comment>
    <dbReference type="EC" id="3.2.1.17" evidence="5"/>
    <dbReference type="AlphaFoldDB" id="A0A452IUT4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000036026.1">
        <property type="protein sequence ID" value="ENSGAGP00000031767.1"/>
        <property type="gene ID" value="ENSGAGG00000022815.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001669">
        <property type="term" value="C:acrosomal vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036126">
        <property type="term" value="C:sperm flagellum"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003796">
        <property type="term" value="F:lysozyme activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042742">
        <property type="term" value="P:defense response to bacterium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007342">
        <property type="term" value="P:fusion of sperm to egg plasma membrane involved in single fertilization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031640">
        <property type="term" value="P:killing of cells of another organism"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd16897">
        <property type="entry name" value="LYZ_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.530.10:FF:000001">
        <property type="entry name" value="Lysozyme C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.530.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001916">
        <property type="entry name" value="Glyco_hydro_22"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000974">
        <property type="entry name" value="Glyco_hydro_22_lys"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023346">
        <property type="entry name" value="Lysozyme-like_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11407">
        <property type="entry name" value="LYSOZYME C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11407:SF25">
        <property type="entry name" value="SPERM ACROSOME MEMBRANE-ASSOCIATED PROTEIN 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00062">
        <property type="entry name" value="Lys"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00137">
        <property type="entry name" value="LYSOZYME"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00135">
        <property type="entry name" value="LYZLACT"/>
    </dbReference>
    <dbReference type="SMART" id="SM00263">
        <property type="entry name" value="LYZ1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53955">
        <property type="entry name" value="Lysozyme-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51348">
        <property type="entry name" value="GLYCOSYL_HYDROL_F22_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0929">Antimicrobial</keyword>
    <keyword evidence="9" id="KW-0081">Bacteriolytic enzyme</keyword>
    <keyword evidence="11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="12" id="KW-0326">Glycosidase</keyword>
    <keyword evidence="10" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0964">Secreted</keyword>
    <keyword evidence="16" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="16">
        <location>
            <begin position="1"/>
            <end position="19"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5018977896" description="Sperm acrosome membrane-associated protein 3" evidence="16">
        <location>
            <begin position="20"/>
            <end position="142"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000632"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00010859"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00011780"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012732"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00016370"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022529"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022638"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023295"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00024656"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00054602"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="RuleBase" id="RU004440"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000031767.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="142" mass="15806" checksum="194179C1ECC80BFC" modified="2019-05-08" version="1">MKALTLLSLLGCLITANKAKIFSRCELAYMLHEEGLDGYEGYSLANWICMAFFESGFNSAAEDDNADGSTDYGIFQINSRVWCNNYRSPTENLCHIPCTGTVPNPTPRSGKSRCRIPAHVIADWTMHCKGRDLSEWVDGCDL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IUT5</accession>
    <name>A0A452IUT5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">SH2 domain containing 3C</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IUT5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000036099.1">
        <property type="protein sequence ID" value="ENSGAGP00000031829.1"/>
        <property type="gene ID" value="ENSGAGG00000022854.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005085">
        <property type="term" value="F:guanyl-nucleotide exchange factor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001784">
        <property type="term" value="F:phosphotyrosine residue binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007264">
        <property type="term" value="P:small GTPase-mediated signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd10337">
        <property type="entry name" value="SH2_BCAR3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.840.10:FF:000007">
        <property type="entry name" value="SH2 domain containing 3C (Predicted)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.505.10:FF:000013">
        <property type="entry name" value="SH2 domain-containing protein 3C isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.840.10">
        <property type="entry name" value="Ras guanine-nucleotide exchange factors catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.505.10">
        <property type="entry name" value="SH2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023578">
        <property type="entry name" value="Ras_GEF_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001895">
        <property type="entry name" value="RASGEF_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036964">
        <property type="entry name" value="RASGEF_cat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000980">
        <property type="entry name" value="SH2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051853">
        <property type="entry name" value="SH2-Ras-GEF_adapter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036860">
        <property type="entry name" value="SH2_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044102">
        <property type="entry name" value="SH2_SHEP1/BCAR3/NSP1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14247">
        <property type="entry name" value="BREAST CANCER ANTI-ESTROGEN RESISTANCE PROTEIN 3 HOMOLOG-LIKE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14247:SF6">
        <property type="entry name" value="SH2 DOMAIN-CONTAINING PROTEIN 3C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00617">
        <property type="entry name" value="RasGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00017">
        <property type="entry name" value="SH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00147">
        <property type="entry name" value="RasGEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00252">
        <property type="entry name" value="SH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48366">
        <property type="entry name" value="Ras GEF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55550">
        <property type="entry name" value="SH2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51257">
        <property type="entry name" value="PROKAR_LIPOPROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50009">
        <property type="entry name" value="RASGEF_CAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50001">
        <property type="entry name" value="SH2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0344">Guanine-nucleotide releasing factor</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1 3" id="KW-0727">SH2 domain</keyword>
    <feature type="domain" description="SH2" evidence="5">
        <location>
            <begin position="62"/>
            <end position="161"/>
        </location>
    </feature>
    <feature type="domain" description="Ras-GEF" evidence="6">
        <location>
            <begin position="436"/>
            <end position="704"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="260"/>
            <end position="351"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="301"/>
            <end position="312"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="4">
        <location>
            <begin position="313"/>
            <end position="325"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022999"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00168"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00191"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50001"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="PROSITE" id="PS50009"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000031829.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="9">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="710" mass="78947" checksum="2708202B39786085" modified="2019-05-08" version="1">MTERCSLWSALSAAACCFYRGSFMQVQFSKEKYILDSSPEKLHKELEEELKLSSTDLRSHVWYHGRIPREVSESLVQRNGDFLIRDSLTSLGDYVLTCRWRNEPLHFKINKVMVKSSDGRTHIQYLFEQESFDNVPALVRFYVGNRKAVSEQSGAIIYCPINRTFPLRYLEASYGLANGKHSGPHSPASQKGGHIKRRSITMTDGLTADKITRGEGCPTSVSLPHHRDIIRNCAVSVDQIQDLHSPMSPISESPLSPAYSTVTRLKPQGGQAGAITPASPVIRRSSEPQLCPGNKPLTDLADSTHSTPCHGNSTASPSSSVNSYSDPDTGHYCQLHPTSPVGQERPTHDTKQLPTKSYAERLKVEGGQRVTAENGSSEAGQRMKGEQDILGFVLPTVEMGSSFNPVAFQSLLIPLENKPLEMAVLKKVKELLAEVDAKALARHITKVDCLVARTLGVTKEMQRLMGVSSGMELLTLPHGHQLRLDLLERFHTMSIMIAVDILGCTGSTEERAALLHKTIQLAAELKSTLGNMFSFAAVMNALEMTQISRLEHTWMALRQRHTEGAILYEKKLKPFLKSLNEGKEGPPLTNTTFPHIVPLITLLERDAVLLDSPEPWENTDNGVEVVMAHLEAARMVAHHGGLYHTNAEVKLQGFQGKAELLEIFSTEFQMRLLWGSRGAESSQRERYEKFDKVLTALSYKLEPSVRFSEL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IUT8</accession>
    <name>A0A452IUT8_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">ATP-dependent 6-phosphofructokinase</fullName>
            <shortName evidence="16">ATP-PFK</shortName>
            <shortName evidence="16">Phosphofructokinase</shortName>
            <ecNumber evidence="16">2.7.1.11</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="16">Phosphohexokinase</fullName>
        </alternativeName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="16">Catalyzes the phosphorylation of D-fructose 6-phosphate to fructose 1,6-bisphosphate by ATP, the first committing step of glycolysis.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="15">
            <text>beta-D-fructose 6-phosphate + ATP = beta-D-fructose 1,6-bisphosphate + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:16109"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:32966"/>
            <dbReference type="ChEBI" id="CHEBI:57634"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.1.11"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 16">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="activity regulation">
        <text evidence="16">Allosterically activated by ADP, AMP, or fructose 2,6-bisphosphate, and allosterically inhibited by ATP or citrate.</text>
    </comment>
    <comment type="pathway">
        <text evidence="2 16 17">Carbohydrate degradation; glycolysis; D-glyceraldehyde 3-phosphate and glycerone phosphate from D-glucose: step 3/4.</text>
    </comment>
    <comment type="subunit">
        <text evidence="16">Homo- and heterotetramers.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="16">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="17">Belongs to the phosphofructokinase type A (PFKA) family. ATP-dependent PFK group I subfamily. Eukaryotic two domain clade "E" sub-subfamily.</text>
    </comment>
    <comment type="similarity">
        <text evidence="16">Belongs to the phosphofructokinase type A (PFKA) family. ATP-dependent PFK group I subfamily. Eukaryotic two domain clade 'E' sub-subfamily.</text>
    </comment>
    <comment type="caution">
        <text evidence="16">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="EC" id="2.7.1.11" evidence="16"/>
    <dbReference type="AlphaFoldDB" id="A0A452IUT8"/>
    <dbReference type="Ensembl" id="ENSGAGT00000036069.1">
        <property type="protein sequence ID" value="ENSGAGP00000031804.1"/>
        <property type="gene ID" value="ENSGAGG00000022790.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00109">
        <property type="reaction ID" value="UER00182"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005945">
        <property type="term" value="C:6-phosphofructokinase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003872">
        <property type="term" value="F:6-phosphofructokinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016208">
        <property type="term" value="F:AMP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070095">
        <property type="term" value="F:fructose-6-phosphate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042802">
        <property type="term" value="F:identical protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048029">
        <property type="term" value="F:monosaccharide binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061621">
        <property type="term" value="P:canonical glycolysis"/>
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        <property type="term" value="P:fructose 1,6-bisphosphate metabolic process"/>
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        <property type="term" value="P:fructose 6-phosphate metabolic process"/>
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        <property type="project" value="InterPro"/>
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        <property type="entry name" value="Eukaryotic_PFK"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="PRK03202.1"/>
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    </dbReference>
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        <property type="entry name" value="ATP-DEPENDENT 6-PHOSPHOFRUCTOKINASE, PLATELET TYPE"/>
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    </dbReference>
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        <property type="entry name" value="PHFRCTKINASE"/>
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    </dbReference>
    <proteinExistence type="inferred from homology"/>
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    <keyword evidence="4 16" id="KW-0021">Allosteric enzyme</keyword>
    <keyword evidence="10 16 17" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="3 16" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="13 16 17" id="KW-0324">Glycolysis</keyword>
    <keyword evidence="14" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9 16 17" id="KW-0418">Kinase</keyword>
    <keyword evidence="11 16" id="KW-0460">Magnesium</keyword>
    <keyword evidence="7 16" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8 16 17" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6 16 17" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Phosphofructokinase" evidence="18">
        <location>
            <begin position="21"/>
            <end position="320"/>
        </location>
    </feature>
    <feature type="domain" description="Phosphofructokinase" evidence="18">
        <location>
            <begin position="400"/>
            <end position="685"/>
        </location>
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            <begin position="1"/>
            <end position="387"/>
        </location>
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            <begin position="400"/>
            <end position="781"/>
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    <feature type="active site" description="Proton acceptor" evidence="16">
        <location>
            <position position="163"/>
        </location>
    </feature>
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        <location>
            <position position="28"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
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        <location>
            <begin position="91"/>
            <end position="92"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
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        <location>
            <begin position="161"/>
            <end position="163"/>
        </location>
        <ligand>
            <name>substrate</name>
            <note>ligand shared between dimeric partners</note>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="198"/>
        </location>
        <ligand>
            <name>substrate</name>
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        </ligand>
    </feature>
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        <location>
            <begin position="205"/>
            <end position="207"/>
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        </ligand>
    </feature>
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        <location>
            <position position="261"/>
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        </ligand>
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        <location>
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        <location>
            <position position="469"/>
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        <ligand>
            <name>beta-D-fructose 2,6-bisphosphate</name>
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        </ligand>
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        <location>
            <begin position="526"/>
            <end position="530"/>
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            <position position="564"/>
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            <end position="573"/>
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        </ligand>
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            <position position="653"/>
        </location>
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            <begin position="659"/>
            <end position="662"/>
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            <position position="733"/>
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        </ligand>
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        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
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    </evidence>
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    </evidence>
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        <source>
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        <source>
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        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00048070"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="HAMAP-Rule" id="MF_03184"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="PIRNR" id="PIRNR000533"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF00365"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000031804.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="781" mass="85945" checksum="96F257C73E99A734" modified="2019-05-08" version="1">MAELPADKKFFENLSGAGKAIGVLTSGGDAQGMNAAVRAVVRMGIYVEAKVYFIYEGYQGMVDGGDNIVEVSWESVSSILQVGGTVIGSARCKEFRTREGRLKAAFNLIQRGITNLCVIGGANLFREEWSGLLEELAQSGKIEEEAVKKYVYLNIVGMVGSIDNDFCGTDMTIGTDSALHRIIEVVDAIMTTAQSHQRTFVLEVMGRHCGYLALVSALACGADWVFIPEYPPEEGWEDQMCVKLSENRARKKRLNIIIVAEGAIDSHNKPITSEKVKDLVVQQLGFDTRVTILGHVQRGGTPSAFDRILASRMGVEAVLALLEATPDTPACVVSLSGNQAVRLPLMECVQMTQEVQKAMDERKFCEAVRLRGRSFENNLNTYKLLSHRKPDTELPKTNFNVAVLNVGAPAAGMNAAVRSAVRVGITEGHNMFAVIDGFEGFANGQIKEISWGDVGGWTGQGGSLLGTKRTLPAKYLEKIADQMRTNNINALMVIGGFEAYESCLQLAEARARFEEFCIPICVLPATISNNVPGTDLSIGADTALNAIVETCDRIKQSASGTKRRVFIIETMGGYCGYLANMGGLAAGADAAYIFEEKFDIRELQANVEHLTEKMKTSIQRGLVLRNENSNENYTTDFIYQLYSEEGKGVFDCRKNVLGHMQQGGAPSPFDRNFGTKVSAKAVQWISKKLRETYRRGKVFANTEDSVCLLGMRRRNLIFQPVVELKDQADFVHRIPKEQWWLKLRPLMKILAKYKTSYDVSDSGQLEHVARHSPKEAETGAI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IUU0</accession>
    <name>A0A452IUU0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Striatin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="14">Calmodulin-binding scaffolding protein which is the center of the striatin-interacting phosphatase and kinase (STRIPAK) complexes. STRIPAK complexes have critical roles in protein (de)phosphorylation and are regulators of multiple signaling pathways including Hippo, MAPK, nuclear receptor and cytoskeleton remodeling. Different types of STRIPAK complexes are involved in a variety of biological processes such as cell growth, differentiation, apoptosis, metabolism and immune regulation.</text>
    </comment>
    <comment type="subunit">
        <text evidence="15">Part of the core of STRIPAK complexes composed of PP2A catalytic and scaffolding subunits, the striatins (PP2A regulatory subunits), the striatin-associated proteins MOB4, STRIP1 and STRIP2, PDCD10 and members of the STE20 kinases, such as STK24 and STK26. Interacts with CTTNBP2; this interaction may regulate dendritic spine distribution of STRN. Activation of glutamate receptors weakens the interaction with CTTNBP2.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cell projection</location>
            <location evidence="3">Dendritic spine</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Peripheral membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the WD repeat striatin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IUU0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000036104.1">
        <property type="protein sequence ID" value="ENSGAGP00000031834.1"/>
        <property type="gene ID" value="ENSGAGG00000022855.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043197">
        <property type="term" value="C:dendritic spine"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032991">
        <property type="term" value="C:protein-containing complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070016">
        <property type="term" value="F:armadillo repeat domain binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051721">
        <property type="term" value="F:protein phosphatase 2A binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0044877">
        <property type="term" value="F:protein-containing complex binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd00200">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Striatin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.300:FF:000001">
        <property type="entry name" value="striatin isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:000079">
        <property type="entry name" value="striatin isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:000211">
        <property type="entry name" value="striatin isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013258">
        <property type="entry name" value="Striatin_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020472">
        <property type="entry name" value="WD40_PAC1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019775">
        <property type="entry name" value="WD40_repeat_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036322">
        <property type="entry name" value="WD40_repeat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001680">
        <property type="entry name" value="WD40_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051488">
        <property type="entry name" value="WD_repeat_striatin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15653">
        <property type="entry name" value="STRIATIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR15653:SF2">
        <property type="entry name" value="STRIATIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08232">
        <property type="entry name" value="Striatin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00400">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00320">
        <property type="entry name" value="GPROTEINBRPT"/>
    </dbReference>
    <dbReference type="SMART" id="SM00320">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50978">
        <property type="entry name" value="WD40 repeat-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00678">
        <property type="entry name" value="WD_REPEATS_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50082">
        <property type="entry name" value="WD_REPEATS_2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50294">
        <property type="entry name" value="WD_REPEATS_REGION"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="13" id="KW-0966">Cell projection</keyword>
    <keyword evidence="11 18" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="12" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="10" id="KW-0770">Synapse</keyword>
    <keyword evidence="7 17" id="KW-0853">WD repeat</keyword>
    <feature type="domain" description="Striatin N-terminal" evidence="20">
        <location>
            <begin position="59"/>
            <end position="186"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="17">
        <location>
            <begin position="431"/>
            <end position="472"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="17">
        <location>
            <begin position="484"/>
            <end position="516"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="17">
        <location>
            <begin position="537"/>
            <end position="569"/>
        </location>
    </feature>
    <feature type="repeat" description="WD" evidence="17">
        <location>
            <begin position="674"/>
            <end position="715"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="134"/>
            <end position="158"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="278"/>
            <end position="324"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="337"/>
            <end position="357"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="18">
        <location>
            <begin position="70"/>
            <end position="125"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="285"/>
            <end position="298"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004552"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00009616"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022574"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00058972"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00063979"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00073140"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00221"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="Pfam" id="PF08232"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000031834.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="752" mass="82751" checksum="65118955A1A957D9" modified="2019-05-08" version="1">MDEQAGPGVFFSNNNNNNALLLPPGGAGPGLEPGEAAAGGSLAAAAPAPVSAARAQYSVPGILHFLQHEWGRFEAERAEWEAERAELQAQIAFLQGERKGQENLKKDLVRRIKMLEYALKQERAKYHKLKYGTELNQGDMKPPNYDSDEGNETEVQPQQNSQLIWKQGRQLLRQYLQEVGYTDTILDVKSKRVRALLGISSDASDRENGKNQESMVNGTEAEVKENAMIGKPELTDSASLLETFKFLENAAADFSDEDEDEDIEREKTMIDTSTMVRRRVVSDSSEDRDTEEALKEFDFLVGSDDGDGESRSSGDGTDWGPNRSKLQDMLANLRDVDDLPSLQPSVTSPSRPSSSRLIEHEINRTDEVEALTFPPTSGKSFIMGADEALESELGLGELAGLTVANEADSLTYDIANNKDALRKTWNPKFTLRSHFDGIRGLAFHPVEPVLITASEDHTLKMWNLQKTAPAKKSASLDVEPIYTFRAHSGPVLCVVMSSNGEQCYSGGTDGLIQAWNTTNPNIDPYDSYDPSVLRGAFVGHTDAVWGLVYSGAHQRLLSCSADGTIRLWKASEIAPALNIFNDNQEMGIPSSVDLVSSDPSHMVASFNSGHTCIFNIETRQRILTLESNRDTTANSSCQINRVVSHPTLPISITAHEDRHIKFYDNNTGKLIHAMVAHLDAVTSLAVDPNGLYLMSGSHDCSIRLWNLESKTCIQEFTAHRKKFDESIHDVAFHPSKCYIASAGADALAKVFV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IUU2</accession>
    <name>A0A452IUU2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">PEST proteolytic signal-containing nuclear protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">May be involved in cell cycle regulation.</text>
    </comment>
    <comment type="subunit">
        <text evidence="3">Interacts with UHRF2/NIRF.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IUU2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000036036.1">
        <property type="protein sequence ID" value="ENSGAGP00000031777.1"/>
        <property type="gene ID" value="ENSGAGG00000022824.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043161">
        <property type="term" value="P:proteasome-mediated ubiquitin-dependent protein catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016567">
        <property type="term" value="P:protein ubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029169">
        <property type="entry name" value="PCNP"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16523">
        <property type="entry name" value="PEST PROTEOLYTIC SIGNAL-CONTAINING NUCLEAR PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR16523:SF6">
        <property type="entry name" value="PEST PROTEOLYTIC SIGNAL-CONTAINING NUCLEAR PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15473">
        <property type="entry name" value="PCNP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0007">Acetylation</keyword>
    <keyword evidence="9" id="KW-0131">Cell cycle</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0832">Ubl conjugation</keyword>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1"/>
            <end position="93"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="158"/>
            <end position="202"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="1"/>
            <end position="12"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="10">
        <location>
            <begin position="13"/>
            <end position="24"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="25"/>
            <end position="37"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="59"/>
            <end position="69"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="163"/>
            <end position="173"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00002646"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00011097"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022059"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023306"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000031777.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="202" mass="22148" checksum="E750FF1C531DEF9F" modified="2019-05-08" version="1">MAGRERENERRLLRMLRTRGRRWGKMADSKAGEDKPKQPQADGGPEEEAEKPVKTKTVSSSNGGESSSRSAEKRLADEEAEDFTTKPAPAKMSKFGFAIGSQTTKKAPVISIKLGANKPKEPAPTLAPKTLSVAAVFNEDDDSEPEEMPPEAKMRMKNIGRDTPTSAGPNSFNKGKHGFSDNQKLWERNIKSHLGNVPEDDD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IUU3</accession>
    <name>A0A452IUU3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Myosin heavy chain 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="13">Muscle myosin is a hexameric protein that consists of 2 heavy chain subunits (MHC), 2 alkali light chain subunits (MLC) and 2 regulatory light chain subunits (MLC-2).</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Myofibril</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 14">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <comment type="caution">
        <text evidence="14">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000036074.1">
        <property type="protein sequence ID" value="ENSGAGP00000031809.1"/>
        <property type="gene ID" value="ENSGAGG00000009599.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030016">
        <property type="term" value="C:myofibril"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032982">
        <property type="term" value="C:myosin filament"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016460">
        <property type="term" value="C:myosin II complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000146">
        <property type="term" value="F:microfilament motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006936">
        <property type="term" value="P:muscle contraction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd01377">
        <property type="entry name" value="MYSc_class_II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000003">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000004">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000006">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000013">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000003">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000007">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000008">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.4820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.360:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000024">
        <property type="entry name" value="Myosin heavy chain, isoform J"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.720:FF:000001">
        <property type="entry name" value="Myosin heavy chain, muscle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.340">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.370">
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.4820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.2420">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.360">
        <property type="entry name" value="Myosin S1 fragment, N-terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008989">
        <property type="entry name" value="Myosin_S1_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002928">
        <property type="entry name" value="Myosin_tail"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004009">
        <property type="entry name" value="SH3_Myosin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014751">
        <property type="entry name" value="XRCC4-like_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615">
        <property type="entry name" value="MYOSIN HEAVY CHAIN, NON-MUSCLE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615:SF79">
        <property type="entry name" value="MYOSIN-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02736">
        <property type="entry name" value="Myosin_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01576">
        <property type="entry name" value="Myosin_tail_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90257">
        <property type="entry name" value="Myosin rod fragments"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51844">
        <property type="entry name" value="SH3_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12 14" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="7 14" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="8 15" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0488">Methylation</keyword>
    <keyword evidence="10 14" id="KW-0505">Motor protein</keyword>
    <keyword evidence="11" id="KW-0514">Muscle protein</keyword>
    <keyword evidence="9 14" id="KW-0518">Myosin</keyword>
    <keyword evidence="6 14" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0787">Thick filament</keyword>
    <feature type="domain" description="Myosin N-terminal SH3-like" evidence="17">
        <location>
            <begin position="34"/>
            <end position="83"/>
        </location>
    </feature>
    <feature type="domain" description="Myosin motor" evidence="16">
        <location>
            <begin position="87"/>
            <end position="545"/>
        </location>
    </feature>
    <feature type="domain" description="Myosin motor" evidence="16">
        <location>
            <begin position="553"/>
            <end position="682"/>
        </location>
    </feature>
    <feature type="region of interest" description="Actin-binding" evidence="14">
        <location>
            <begin position="559"/>
            <end position="581"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="15">
        <location>
            <begin position="746"/>
            <end position="1088"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="15">
        <location>
            <begin position="1140"/>
            <end position="1821"/>
        </location>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <begin position="180"/>
            <end position="187"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004657"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00008314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022433"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023179"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00038612"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS51844"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000031809.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <evidence type="ECO:0008006" key="20">
        <source>
            <dbReference type="Google" id="ProtNLM"/>
        </source>
    </evidence>
    <sequence length="1833" mass="211598" checksum="A40DCF361D9BB70D" modified="2019-05-08" version="1">AMSSDAEMAVFGVAAPFLRKSEKERIEDQNKPFDAKTSVFVAHPKESFVKSTIQSREGGKVTVKTEKGETLTVKEDQIFSMNPPKYDKIEDMAMMTHLHEPGVLYNLKERYAAWMIYTYSGLFCVTINPYKWLPVYNPEVVAAYRGKKRQEAPPHIFSISDSAYQFMLTDRENQSILITGESGAGKTVNTKRVIQYFATIAASGEKKKEEPGKMQGTLEDQIISANPLLEAFGNAKTVRNDNSSRFGKFIRIHFGTTGKLASADIETYLLEKSRVTFQLKAERSYHIFYQIMSNKKPELIEMLLITTNPYDFPFVSQGEISVASIDDQEELIATDSAIDILGFSSEEKTAIYKLTGAVLHYGNLKFKQKQREEQAEPDGTEVADKAAYLMNLNSADLLKAMCCPRVKVGNEYVTKGQTVQQVHNSVGALAKAVYEKMFLWMVVRINQQLDTKQPRQYFIGVLDIAGFEIFDYNSLEQLCINFTNEKLQQFFNHHMFVLEQEEYKKEGIEWTFIDFGMDLAACIELIEKVSIQFKCLIHLRILYLLGVVLILEGLFPENLNKLMSNLRSTHPHFVRCIIPNETKTPGAMEHELVLHQLRCNGVLEGIRICRKGFPSRIVYADFKQRYRILNASAVPEGHFMDSKKASEKLLGSIDIDQTQYKFGHTKVFFKAGLLGLLEEMRDDKLAEIMTRTQARCRGFLMRVEYQRMVERRESVFCIQYNVRSFMNVKHWPWMKLYFKIKPLLRSAEAEKEMATMKEEFEKTKEELAKSEAKRKELEEKMVTLMQEKNDLQLQVQSEAEGLADAEERCDQLIKTKIQLEAKIKEVTERAEDEEEMNAELTAKKRKLEDECSELKKDIDDLELTLAKVEKEKHATENKVKNLTEEMAVLDETIAKLTKEKKALQEAHQQTLDDLQAEEDKVNTLTKAKTKLEQQVDDLEGSLEQEKKIRMDLERAKRKLEGDLKLAQESTMDLENDKQQLEEKLKKKDFEISQFLSRIEDEQVLAIQLQKKIKELQARIEELEEEIEAERASRAKAEKQRGDLSRELEEISERLEEAGGATAVQIEMNKKREAEFQKMRRDLEEATLQHEATAAALRKKHADSTAELGEQIDNLNLETVSKAKPLMDLSSMNLSSYCLNTANLEKMCRTLEDQLSEIKTKEEEHQRMINDLNAQRARLQTESGEFARQVDEKDALISQLSRGKQAFTQQIEELKRHLEEEIKAKNALAHALQSARHDCDLLREQYEEEQEAKGELQRALSKANSEVAQWRTKYETDAIQRTEELEDAKKKLAQRLQDAEEHVEAVNSKCASLEKTKQRLQNEVEDLMIDVERSNAACAALDKKQKNFDKVLAEWKQKYEETQAELEASQKESRSLSTELFKMKNSYEETLDQLETLKRENKNLQQEISDLTEQIAEGGKTIHELEKVKKQIEQEKSEIQSALEEAEASLEHEEGKILRIQLELNQVKSEIDRKIAEKDEEIDQLKRNHIRVVESMQSTLDAEIRSRNEAIRLKKKMEGDLNEMEIQLSHANRLAAEAQKNLRNTQGVLKDTQIHLDDALRSQEDLKEQVAMVERRANLMLAEIEELRSALEQTERARKVAEQELLDASERVQLLHTQNTSLINTKKKLETDISQIQSEMEETIQEARNAEEKAKKAITDAAMMAEELKKEQDTSAHLERMKKNLDQTVKDLQLRLDEAEQLALKGGKKQIQKLEARVRELEGEVDNEQKRSADAIKGVRKYERRVKELTYQSEEDRKNILRLQDLVDKLQLKVKSYKRQAEESEELSNVNLTKFRKIQHELEEAEERADIAESQVNKLRVKSREIHRKIEGEE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IUU4</accession>
    <name>A0A452IUU4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Olfactomedin-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="3">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IUU4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000036111.1">
        <property type="protein sequence ID" value="ENSGAGP00000031839.1"/>
        <property type="gene ID" value="ENSGAGG00000022862.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005615">
        <property type="term" value="C:extracellular space"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003112">
        <property type="entry name" value="Olfac-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050605">
        <property type="entry name" value="Olfactomedin-like_domain"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23192:SF37">
        <property type="entry name" value="OLFACTOMEDIN-LIKE PROTEIN 2B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23192">
        <property type="entry name" value="OLFACTOMEDIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02191">
        <property type="entry name" value="OLF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00284">
        <property type="entry name" value="OLF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF101898">
        <property type="entry name" value="NHL repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51132">
        <property type="entry name" value="OLF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <feature type="domain" description="Olfactomedin-like" evidence="5">
        <location>
            <begin position="306"/>
            <end position="563"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="270"/>
            <end position="292"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="279"/>
            <end position="292"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00446"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS51132"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000031839.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="563" mass="63838" checksum="564308E850FFBBF9" modified="2019-05-08" version="1">SVSTLWNITQPGGTEVSRNLTRENMDATGNVDVIWSEPNTRDKENCSSSLPDIRSSLQRDAAAAYTHTEEKYKERFLQDEMISQQINSVESLQQLLPGPEAMKPKQPLQRKMHVRSRLPSKPTIIRGITYYKAKSTESENDLEEQDESFSGDHTVDLLIEDQLLRHNSHERSSTTRMPAPVGGPLTLRNIAPTIEYISQALPATLKTTTAPGTITMSGSVEEPATWTPATLTHTGTQAPTTAANLPTASPRKATIAMTTTTVHISTTTGSNVDAFRSEPPTTVSPTLPATPRQNLDEEDLRSIIGRCKDTLSTISGPTTQNTYGRNEGAWMKDPLAKDERIYVTNYYYGNTLVEFRNLENFKQGRWSNSYKLPYSWIGTGHAVYNGSLYYNRAFTRNIIKYDLKQRYVAAWAMLHDVAYEEATPWRWQGHSDVDFAVDENGLWIIYPAINYEGFNQEVIVLSKLNAADLSTQKETTWRTGLRRNFYRNSFVICGVLYAVDSYNHRNANISYAFDTHTNTQIIPRLLFENEYSYITQIDYNPKDRLLYAWDNGHQVTYHVIFAY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2025-10-08" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IV51</accession>
    <name>A0A452IV51_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">6-phosphofructokinase type C</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="13">Phosphofructo-1-kinase isozyme C</fullName>
        </alternativeName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="14">
            <text>beta-D-fructose 6-phosphate + ATP = beta-D-fructose 1,6-bisphosphate + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:16109"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:32966"/>
            <dbReference type="ChEBI" id="CHEBI:57634"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.1.11"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="pathway">
        <text evidence="2">Carbohydrate degradation; glycolysis; D-glyceraldehyde 3-phosphate and glycerone phosphate from D-glucose: step 3/4.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000036172.1">
        <property type="protein sequence ID" value="ENSGAGP00000031894.1"/>
        <property type="gene ID" value="ENSGAGG00000022790.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000036173.1">
        <property type="protein sequence ID" value="ENSGAGP00000031895.1"/>
        <property type="gene ID" value="ENSGAGG00000022790.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00109">
        <property type="reaction ID" value="UER00182"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005945">
        <property type="term" value="C:6-phosphofructokinase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003872">
        <property type="term" value="F:6-phosphofructokinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016208">
        <property type="term" value="F:AMP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070095">
        <property type="term" value="F:fructose-6-phosphate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042802">
        <property type="term" value="F:identical protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048029">
        <property type="term" value="F:monosaccharide binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061621">
        <property type="term" value="P:canonical glycolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030388">
        <property type="term" value="P:fructose 1,6-bisphosphate metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006002">
        <property type="term" value="P:fructose 6-phosphate metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.460:FF:000001">
        <property type="entry name" value="ATP-dependent 6-phosphofructokinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.460:FF:000003">
        <property type="entry name" value="ATP-dependent 6-phosphofructokinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.450:FF:000064">
        <property type="entry name" value="Phosphofructokinase, platelet b"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.450">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.460">
        <property type="entry name" value="Phosphofructokinase domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009161">
        <property type="entry name" value="6-Pfructokinase_euk"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022953">
        <property type="entry name" value="ATP_PFK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015912">
        <property type="entry name" value="Phosphofructokinase_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000023">
        <property type="entry name" value="Phosphofructokinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035966">
        <property type="entry name" value="PKF_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR02478">
        <property type="entry name" value="6PF1K_euk"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13697:SF5">
        <property type="entry name" value="ATP-DEPENDENT 6-PHOSPHOFRUCTOKINASE, PLATELET TYPE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13697">
        <property type="entry name" value="PHOSPHOFRUCTOKINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00365">
        <property type="entry name" value="PFK"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00476">
        <property type="entry name" value="PHFRCTKINASE"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53784">
        <property type="entry name" value="Phosphofructokinase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00433">
        <property type="entry name" value="PHOSPHOFRUCTOKINASE"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0021">Allosteric enzyme</keyword>
    <keyword evidence="9" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11" id="KW-0324">Glycolysis</keyword>
    <keyword evidence="8" id="KW-0418">Kinase</keyword>
    <keyword evidence="10" id="KW-0460">Magnesium</keyword>
    <keyword evidence="6" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Phosphofructokinase" evidence="15">
        <location>
            <begin position="2"/>
            <end position="210"/>
        </location>
    </feature>
    <feature type="domain" description="Phosphofructokinase" evidence="15">
        <location>
            <begin position="296"/>
            <end position="581"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022533"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023152"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00041248"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00041363"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00048070"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="Pfam" id="PF00365"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000031894.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="677" mass="74757" checksum="691DAC32A6FB9F22" modified="2019-05-08" version="1">MVVXGDGSLTGANLFREEWSGLLEELAQSGKIEEEAVKKYVYLNIVGMVGSIDNDFCGTDMTIGTDSALHRIIEVVDAIMTTAQSHQRTFVLEVMGRHCGYLALVSALACGADWVFIPEYPPEEGWEDQMCVKLSENRARKKRLNIIIVAEGAIDSHNKPITSEKVKDLVVQQLGFDTRVTILGHVQRGGTPSAFDRILASRMGVEAVLALLEATPDTPACVVSLSGNQAVRLPLMECVQMTQEVQKAMDERKFCEAVRLRGRSFENNLNTYKLLSHRKPDTELPKDDLSCSTNFNVAVLNVGAPAAGMNAAVRSAVRVGITEGHNMFAVIDGFEGFANGQIKEISWGDVGGWTGQGGSLLGTKRTLPAKYLEKIADQMRTNNINALMVIGGFEAYLGLLELSAAREKYDEFCVPMVMVPATVSNNVPGSDFSIGADTALNTITDTCDRIKQSASGTKRRVFIIETMGGYCGYLANMGGLAAGADAAYIFEEKFDIRELQANVEHLTEKMKTSIQRGLVLRNENSNENYTTDFIYQLYSEEGKGVFDCRKNVLGHMQQGGAPSPFDRNFGTKVSAKAVQWISKKLRETYRRGKVFANTEDSVCLLGMRRRNLIFQPVVELKDQADFVHRIPKEQWWLKLRPLMKILAKYKTSYDVSDSGQLEHVARHSPKEAETGAI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IV53</accession>
    <name>A0A452IV53_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">1-acylglycerol-3-phosphate O-acyltransferase ABHD5</fullName>
            <ecNumber evidence="5">2.3.1.51</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="19">Abhydrolase domain-containing protein 5</fullName>
        </alternativeName>
    </protein>
    <organism evidence="28 29">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="29" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="28" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="28" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="20">Coenzyme A-dependent lysophosphatidic acid acyltransferase that catalyzes the transfer of an acyl group on a lysophosphatidic acid. Functions preferentially with 1-oleoyl-lysophosphatidic acid followed by 1-palmitoyl-lysophosphatidic acid, 1-stearoyl-lysophosphatidic acid and 1-arachidonoyl-lysophosphatidic acid as lipid acceptor. Functions preferentially with arachidonoyl-CoA followed by oleoyl-CoA as acyl group donors. Functions in phosphatidic acid biosynthesis. May regulate the cellular storage of triacylglycerol through activation of the phospholipase PNPLA2. Involved in keratinocyte differentiation. Regulates lipid droplet fusion.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="24">
            <text>1-(5Z,8Z,11Z,14Z-eicosatetraenoyl)-sn-glycero-3-phosphate + (9Z)-octadecenoyl-CoA = 1-(5Z,8Z,11Z,14Z)-eicosatetraenoyl-2-(9Z)-octadecenoyl-sn-glycero-3-phosphate + CoA</text>
            <dbReference type="Rhea" id="RHEA:37455"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57387"/>
            <dbReference type="ChEBI" id="CHEBI:74938"/>
            <dbReference type="ChEBI" id="CHEBI:74941"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="24">
            <dbReference type="Rhea" id="RHEA:37456"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="25">
            <text>1-(9Z-octadecenoyl)-sn-glycero-3-phosphate + (5Z,8Z,11Z,14Z)-eicosatetraenoyl-CoA = 1-(9Z)-octadecenoyl-2-(5Z,8Z,11Z,14Z)-eicosatetraenoyl-sn-glycero-3-phosphate + CoA</text>
            <dbReference type="Rhea" id="RHEA:37443"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57368"/>
            <dbReference type="ChEBI" id="CHEBI:74544"/>
            <dbReference type="ChEBI" id="CHEBI:74928"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="25">
            <dbReference type="Rhea" id="RHEA:37444"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="26">
            <text>1-(9Z-octadecenoyl)-sn-glycero-3-phosphate + (9Z)-octadecenoyl-CoA = 1,2-di-(9Z-octadecenoyl)-sn-glycero-3-phosphate + CoA</text>
            <dbReference type="Rhea" id="RHEA:37131"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57387"/>
            <dbReference type="ChEBI" id="CHEBI:74544"/>
            <dbReference type="ChEBI" id="CHEBI:74546"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="26">
            <dbReference type="Rhea" id="RHEA:37132"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="2">
            <text>1-(9Z-octadecenoyl)-sn-glycero-3-phosphate + hexadecanoyl-CoA = 1-(9Z)-octadecenoyl-2-hexadecanoyl-sn-glycero-3-phosphate + CoA</text>
            <dbReference type="Rhea" id="RHEA:37143"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57379"/>
            <dbReference type="ChEBI" id="CHEBI:74544"/>
            <dbReference type="ChEBI" id="CHEBI:74551"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="2">
            <dbReference type="Rhea" id="RHEA:37144"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="16">
            <text>1-(9Z-octadecenoyl)-sn-glycero-3-phosphate + octadecanoyl-CoA = 1-(9Z-octadecenoyl)-2-octadecanoyl-sn-glycero-3-phosphate + CoA</text>
            <dbReference type="Rhea" id="RHEA:37147"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57394"/>
            <dbReference type="ChEBI" id="CHEBI:74544"/>
            <dbReference type="ChEBI" id="CHEBI:74552"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="16">
            <dbReference type="Rhea" id="RHEA:37148"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="21">
            <text>1-hexadecanoyl-sn-glycero-3-phosphate + (9Z)-octadecenoyl-CoA = 1-hexadecanoyl-2-(9Z-octadecenoyl)-sn-glycero-3-phosphate + CoA</text>
            <dbReference type="Rhea" id="RHEA:33187"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57387"/>
            <dbReference type="ChEBI" id="CHEBI:57518"/>
            <dbReference type="ChEBI" id="CHEBI:64839"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="21">
            <dbReference type="Rhea" id="RHEA:33188"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="22">
            <text>1-octadecanoyl-sn-glycero-3-phosphate + (9Z)-octadecenoyl-CoA = 1-octadecanoyl-2-(9Z-octadecenoyl)-sn-glycero-3-phosphate + CoA</text>
            <dbReference type="Rhea" id="RHEA:37163"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57387"/>
            <dbReference type="ChEBI" id="CHEBI:74560"/>
            <dbReference type="ChEBI" id="CHEBI:74565"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="22">
            <dbReference type="Rhea" id="RHEA:37164"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>a 1-acyl-sn-glycero-3-phosphate + an acyl-CoA = a 1,2-diacyl-sn-glycero-3-phosphate + CoA</text>
            <dbReference type="Rhea" id="RHEA:19709"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57970"/>
            <dbReference type="ChEBI" id="CHEBI:58342"/>
            <dbReference type="ChEBI" id="CHEBI:58608"/>
            <dbReference type="EC" id="2.3.1.51"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="1">
            <dbReference type="Rhea" id="RHEA:19710"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="23">
            <text>eicosanoyl-CoA + 1-(9Z-octadecenoyl)-sn-glycero-3-phosphate = 1-(9Z)-octadecenoyl-2-eicosanoyl-sn-glycero-3-phosphate + CoA</text>
            <dbReference type="Rhea" id="RHEA:37451"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57380"/>
            <dbReference type="ChEBI" id="CHEBI:74544"/>
            <dbReference type="ChEBI" id="CHEBI:74937"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="23">
            <dbReference type="Rhea" id="RHEA:37452"/>
        </physiologicalReaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Lipid droplet</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="17">Belongs to the peptidase S33 family. ABHD4/ABHD5 subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.3.1.51" evidence="5"/>
    <dbReference type="AlphaFoldDB" id="A0A452IV53"/>
    <dbReference type="Ensembl" id="ENSGAGT00000036014.1">
        <property type="protein sequence ID" value="ENSGAGP00000031756.1"/>
        <property type="gene ID" value="ENSGAGG00000022810.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005811">
        <property type="term" value="C:lipid droplet"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003841">
        <property type="term" value="F:1-acylglycerol-3-phosphate O-acyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0052689">
        <property type="term" value="F:carboxylic ester hydrolase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006631">
        <property type="term" value="P:fatty acid metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055088">
        <property type="term" value="P:lipid homeostasis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010891">
        <property type="term" value="P:negative regulation of triglyceride storage"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006654">
        <property type="term" value="P:phosphatidic acid biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010898">
        <property type="term" value="P:positive regulation of triglyceride catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.1820:FF:000019">
        <property type="entry name" value="1-acylglycerol-3-phosphate O-acyltransferase ABHD5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.1820">
        <property type="entry name" value="alpha/beta hydrolase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000073">
        <property type="entry name" value="AB_hydrolase_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029058">
        <property type="entry name" value="AB_hydrolase_fold"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR42886:SF34">
        <property type="entry name" value="1-ACYLGLYCEROL-3-PHOSPHATE O-ACYLTRANSFERASE ABHD5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR42886">
        <property type="entry name" value="RE40534P-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00561">
        <property type="entry name" value="Abhydrolase_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00111">
        <property type="entry name" value="ABHYDROLASE"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53474">
        <property type="entry name" value="alpha/beta-Hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="15" id="KW-0012">Acyltransferase</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="10" id="KW-0221">Differentiation</keyword>
    <keyword evidence="11" id="KW-0276">Fatty acid metabolism</keyword>
    <keyword evidence="7" id="KW-0444">Lipid biosynthesis</keyword>
    <keyword evidence="8" id="KW-0551">Lipid droplet</keyword>
    <keyword evidence="12" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="13" id="KW-0594">Phospholipid biosynthesis</keyword>
    <keyword evidence="14" id="KW-1208">Phospholipid metabolism</keyword>
    <keyword evidence="29" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="AB hydrolase-1" evidence="27">
        <location>
            <begin position="74"/>
            <end position="240"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00000816"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004502"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00013211"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022516"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022677"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022832"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023209"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023264"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023315"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00036296"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00038097"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00040731"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00042413"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00045357"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00047525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00047543"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00047849"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00048632"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00048770"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="ARBA" id="ARBA00049561"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="27">
        <source>
            <dbReference type="Pfam" id="PF00561"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="28">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000031756.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="29">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="318" mass="35799" checksum="F4D1213D20EF85CC" modified="2019-05-08" version="1">MAEDGHLEASSEGSGWLSSWLPAWCPTSLPHLKEAEEKILKCITSTYSKRYVFISNGNKIWTLTFSQDLSYKTPLVLLHGFGGGVGLWALNFEELCENRTVYAFDILGFGRSSRPQFDTDAEEAENQFVEAVEEWRQEVGLDKMILLGHNLGGFLAAAYSLKYPSRVKHLILVEPWGFPERPDNAEQERPIPIWIKALGAMLSPFNPLAGLRLAGPFGLSLVQRLRPDFKRKYSSMFDDNTVTEYIYHCNVQSPSGETAFKNMTVPYGWAKRPMLQRIAHMHQDIPITVIYGARSCIDGNSGSTIQSLRPDSYVKTIV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IV54</accession>
    <name>A0A452IV54_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">SMB domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IV54"/>
    <dbReference type="Ensembl" id="ENSGAGT00000036236.1">
        <property type="protein sequence ID" value="ENSGAGP00000031957.1"/>
        <property type="gene ID" value="ENSGAGG00000022901.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003676">
        <property type="term" value="F:nucleic acid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0047429">
        <property type="term" value="F:nucleoside triphosphate diphosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009143">
        <property type="term" value="P:nucleoside triphosphate catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd16018">
        <property type="entry name" value="Enpp"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00091">
        <property type="entry name" value="NUC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.410.20:FF:000001">
        <property type="entry name" value="Ectonucleotide pyrophosphatase/phosphodiesterase family member 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.570.10:FF:000005">
        <property type="entry name" value="ectonucleotide pyrophosphatase/phosphodiesterase family member 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.720.10:FF:000145">
        <property type="entry name" value="Uncharacterized protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.410.20">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.720.10">
        <property type="entry name" value="Alkaline Phosphatase, subunit A"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.570.10">
        <property type="entry name" value="Extracellular Endonuclease, subunit A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017850">
        <property type="entry name" value="Alkaline_phosphatase_core_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044929">
        <property type="entry name" value="DNA/RNA_non-sp_Endonuclease_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001604">
        <property type="entry name" value="Endo_G_ENPP1-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020821">
        <property type="entry name" value="ENPP1-3/EXOG-like_nuc-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044925">
        <property type="entry name" value="His-Me_finger_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002591">
        <property type="entry name" value="Phosphodiest/P_Trfase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036024">
        <property type="entry name" value="Somatomedin_B-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001212">
        <property type="entry name" value="Somatomedin_B_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10151">
        <property type="entry name" value="ECTONUCLEOTIDE PYROPHOSPHATASE/PHOSPHODIESTERASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10151:SF107">
        <property type="entry name" value="ECTONUCLEOTIDE PYROPHOSPHATASE_PHOSPHODIESTERASE FAMILY MEMBER 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01223">
        <property type="entry name" value="Endonuclease_NS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01663">
        <property type="entry name" value="Phosphodiest"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01033">
        <property type="entry name" value="Somatomedin_B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00892">
        <property type="entry name" value="Endonuclease_NS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00477">
        <property type="entry name" value="NUC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00201">
        <property type="entry name" value="SO"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53649">
        <property type="entry name" value="Alkaline phosphatase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54060">
        <property type="entry name" value="His-Me finger endonucleases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90188">
        <property type="entry name" value="Somatomedin B domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00524">
        <property type="entry name" value="SMB_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50958">
        <property type="entry name" value="SMB_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="6" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="4" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <feature type="domain" description="SMB" evidence="7">
        <location>
            <begin position="51"/>
            <end position="94"/>
        </location>
    </feature>
    <feature type="domain" description="SMB" evidence="7">
        <location>
            <begin position="95"/>
            <end position="139"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50958"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000031957.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="780" mass="89023" checksum="E98E5DFF1C84DE75" modified="2019-05-08" version="1">AFPVYIAQLQEPVFGNYSEHSDSRLLAMFHLIVVFLRTGITLVLKLSFVLFTVSCRNRCNEPFNKKGQGCQCDGGCKERQNCCWDYEGTCLEPTQIWTCTKLRCGETRITGSHCSCSEDCLEKKDCCVDYKSICKAEYLQTWSSLLPNIEKLKTCGTHSKYMRAAYPTKTFPNHYTIVTGLYPESHGIIDNNMYDVNLSQSFSLSGQAKFDPVWWKGQPIWLTAMYQNLKAVSKYSGHSYGPVSGGVIKALQLADEALGMLMEGLKQRNLHNCVNLIVLADHGMDKTYCNQLEYMTDYFNQINFYMYDGPAARIRAYNVPEDYFTFDSEGIVKNLTCRKPNQHFKPYLTPNLPKRFHYANNIRIDKVHLMVDRQWLAVRNKDYTFCGGGNHGYDNEFKSMEAIFMAHGPSFKEKTEVEPFENIDIYNLMCDLLHIKPAPNNGTHGSLNHLLKTAFYNPSHSKEESPPSSCPFTTLTPLDELGCTCAPLVSILEAINQRLNLTIEEIKKAETYNLPYGRPRVLQKQNTYCLLHHHQYVSGYSHNIWMPLWSAYTVNKTADSSSLPSTVPDCLRADVRIPATQSQNCSDYQLDLNITHGFLYPPNFNATGPEQYDALLTSNIVPMYEEFKKIWNYFHDVLLLKYAAERNGLNVISGPVFDYNYDGHFDALHEITQHINNTQIPVPTHYFVMLTSCKNQSNTPLKCVGSLDVSSFIIPHRPDNSESCAVSMSQWVEERVQAHAARVRDVELVTGLDFYQDRDQPLSEILQLKTFLPTFETIIN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IV57</accession>
    <name>A0A452IV57_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">C2 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasmic vesicle membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="11">Endomembrane system</location>
            <topology evidence="11">Single-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the synaptotagmin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IV57"/>
    <dbReference type="Ensembl" id="ENSGAGT00000036020.1">
        <property type="protein sequence ID" value="ENSGAGP00000031761.1"/>
        <property type="gene ID" value="ENSGAGG00000022814.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030424">
        <property type="term" value="C:axon"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030659">
        <property type="term" value="C:cytoplasmic vesicle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070382">
        <property type="term" value="C:exocytic vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098793">
        <property type="term" value="C:presynapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="GOC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005544">
        <property type="term" value="F:calcium-dependent phospholipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030276">
        <property type="term" value="F:clathrin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001786">
        <property type="term" value="F:phosphatidylserine binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000149">
        <property type="term" value="F:SNARE binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048791">
        <property type="term" value="P:calcium ion-regulated exocytosis of neurotransmitter"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:1903531">
        <property type="term" value="P:negative regulation of secretion by cell"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006906">
        <property type="term" value="P:vesicle fusion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd08388">
        <property type="entry name" value="C2A_Synaptotagmin-4-11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd08404">
        <property type="entry name" value="C2B_Synaptotagmin-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000039">
        <property type="entry name" value="Synaptotagmin 11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000051">
        <property type="entry name" value="Synaptotagmin 11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000008">
        <property type="entry name" value="C2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001565">
        <property type="entry name" value="Synaptotagmin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10024">
        <property type="entry name" value="SYNAPTOTAGMIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10024:SF115">
        <property type="entry name" value="SYNAPTOTAGMIN-11"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00168">
        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00399">
        <property type="entry name" value="SYNAPTOTAGMN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00239">
        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49562">
        <property type="entry name" value="C2 domain (Calcium/lipid-binding domain, CaLB)"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50004">
        <property type="entry name" value="C2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0106">Calcium</keyword>
    <keyword evidence="10" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="9 12" id="KW-0472">Membrane</keyword>
    <keyword evidence="5" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="4 12" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="8 12" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="12">
        <location>
            <begin position="16"/>
            <end position="37"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="13">
        <location>
            <begin position="156"/>
            <end position="278"/>
        </location>
    </feature>
    <feature type="domain" description="C2" evidence="13">
        <location>
            <begin position="290"/>
            <end position="425"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004156"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006996"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00037847"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="PROSITE" id="PS50004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000031761.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="430" mass="47923" checksum="899307FC3D8D02BF" modified="2019-05-08" version="1">MAEIASVRPGLDVPPIVAGLIGATVLVVSVSVTVFVWTCCHQQAEKKHNPPPYKFIHMLKGISIYPETLSNKKKIIRIRRDKNGTVKETGRGNLLVDAAEAGLVGSEKAPNGMNAALCIDQLPIKVDYGDELSPDQSITPGGSKISSPSSPEEDVLLGTLTLSVDYNFPKKALVVTIQEAHGLPVMDEQTQGSDPYIKMTILPDKRHRVKTRVLRKTLDPVFDETFTFYGIPYSQLQDLVLHFLVLSFDRFSRDDVIGEVMVPLAGVDPSTGKVQLTREIIKRNIQKCISRGELQVSLSYQPVAQRMTVVVLKARHLPKMDITGLSGNPYVKVNVYYGRKRIAKKKTHVKKCTLNPIFNESFIYDIPVDLLPDISTEFLVIDFDRTTKNEVVGRLILGAHSITTGGVEHWREVCENPRKQIAKWHSLSEY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IV58</accession>
    <name>A0A452IV58_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">mitogen-activated protein kinase kinase</fullName>
            <ecNumber evidence="9">2.7.12.2</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="10">
            <text>L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:17989"/>
            <dbReference type="Rhea" id="RHEA-COMP:9863"/>
            <dbReference type="Rhea" id="RHEA-COMP:11604"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29999"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:83421"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.12.2"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="11">
            <text>L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:46608"/>
            <dbReference type="Rhea" id="RHEA-COMP:11060"/>
            <dbReference type="Rhea" id="RHEA-COMP:11605"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30013"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:61977"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.12.2"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="12">
            <text>L-tyrosyl-[protein] + ATP = O-phospho-L-tyrosyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10596"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.12.2"/>
        </reaction>
    </comment>
    <comment type="similarity">
        <text evidence="8">Belongs to the protein kinase superfamily. STE Ser/Thr protein kinase family. MAP kinase kinase subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.12.2" evidence="9"/>
    <dbReference type="AlphaFoldDB" id="A0A452IV58"/>
    <dbReference type="Ensembl" id="ENSGAGT00000036201.1">
        <property type="protein sequence ID" value="ENSGAGP00000031922.1"/>
        <property type="gene ID" value="ENSGAGG00000022911.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008545">
        <property type="term" value="F:JUN kinase kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004713">
        <property type="term" value="F:protein tyrosine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0033554">
        <property type="term" value="P:cellular response to stress"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd06616">
        <property type="entry name" value="PKc_MKK4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000090">
        <property type="entry name" value="Dual specificity mitogen-activated protein kinase kinase 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000126">
        <property type="entry name" value="Dual specificity mitogen-activated protein kinase kinase 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48013:SF15">
        <property type="entry name" value="DUAL SPECIFICITY MITOGEN-ACTIVATED PROTEIN KINASE KINASE 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48013">
        <property type="entry name" value="DUAL SPECIFICITY MITOGEN-ACTIVATED PROTEIN KINASE KINASE 5-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6 13 14" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="5" id="KW-0418">Kinase</keyword>
    <keyword evidence="4 13 14" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1 14" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="3" id="KW-0808">Transferase</keyword>
    <keyword evidence="7" id="KW-0829">Tyrosine-protein kinase</keyword>
    <feature type="domain" description="Protein kinase" evidence="16">
        <location>
            <begin position="142"/>
            <end position="407"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="1"/>
            <end position="33"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="15">
        <location>
            <begin position="1"/>
            <end position="25"/>
        </location>
    </feature>
    <feature type="binding site" evidence="13">
        <location>
            <position position="171"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023137"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00038035"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00038999"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00049014"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00049299"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00051693"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="RuleBase" id="RU000304"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000031922.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="439" mass="49251" checksum="E0FCD3B6861FD936" modified="2019-05-08" version="1">MATPSPSSGSSSGSGSSSGSSTPGSQAHTAVSSMQDVHLDTFSAESSKRKALKLNFANPPFKSTARFTLNTSGVPFQNPHIDTQATLGSNHIVPFTFKRREADLIQKLSSRLQERLRTHSIESSGKLKISPEQHWDFTAEDLKDLGEIGRGAYGSVNKMVHKPSGQIMAVKRIRSTVDEKEQKQLLMDLDVVMRSSDCPYIVQFYGALFREGDCWICMELMSTSFDKFYKYVYSVLDDVIPEEILGKITLATVKALNHLKENLKIIHRDIKPSNILLDRNGNIKLCDFGISGQLVDSIAKTRDAGCRPYMAPERIDPSASRQGYDVRSDVWSLGITLYELATGRFPYPKWNSVFDQLTQVVKGDPPQLSNSEEREFSPSFINFVNLCLTKDESKRPKYKELLKHPFILMYEERTVDVACYVCKILDQMPTTPSSPMYVD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IV62</accession>
    <name>A0A452IV62_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Phosphorylase b kinase regulatory subunit</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="7">Phosphorylase b kinase catalyzes the phosphorylation of serine in certain substrates, including troponin I.</text>
    </comment>
    <comment type="pathway">
        <text evidence="2 7">Glycan biosynthesis; glycogen metabolism.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 7">Cell membrane</location>
            <topology evidence="1 7">Lipid-anchor</topology>
            <orientation evidence="1 7">Cytoplasmic side</orientation>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 7">Belongs to the phosphorylase b kinase regulatory chain family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IV62"/>
    <dbReference type="Ensembl" id="ENSGAGT00000036182.1">
        <property type="protein sequence ID" value="ENSGAGP00000031904.1"/>
        <property type="gene ID" value="ENSGAGG00000022485.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00163"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005964">
        <property type="term" value="C:phosphorylase kinase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005977">
        <property type="term" value="P:glycogen metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008928">
        <property type="entry name" value="6-hairpin_glycosidase_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011613">
        <property type="entry name" value="GH15-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045583">
        <property type="entry name" value="KPBA/B_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008734">
        <property type="entry name" value="PHK_A/B_su"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10749">
        <property type="entry name" value="PHOSPHORYLASE B KINASE REGULATORY SUBUNIT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10749:SF8">
        <property type="entry name" value="PHOSPHORYLASE B KINASE REGULATORY SUBUNIT BETA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00723">
        <property type="entry name" value="Glyco_hydro_15"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19292">
        <property type="entry name" value="KPBB_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48208">
        <property type="entry name" value="Six-hairpin glycosidases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 7" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="6 7" id="KW-0119">Carbohydrate metabolism</keyword>
    <keyword evidence="7" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="4 7" id="KW-0321">Glycogen metabolism</keyword>
    <keyword evidence="7" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="7" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0636">Prenylation</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="GH15-like" evidence="8">
        <location>
            <begin position="36"/>
            <end position="758"/>
        </location>
    </feature>
    <feature type="domain" description="Phosphorylase b kinase regulatory subunit alpha/beta C-terminal" evidence="9">
        <location>
            <begin position="819"/>
            <end position="977"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004342"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005131"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022600"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023277"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU364123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF00723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF19292"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000031904.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="993" mass="113737" checksum="8B0FD8E21C3F213E" modified="2019-05-08" version="1">SGYSPTKAHASKSSVYEPLKSINLPRPEGESLWDKLDHYYRIVKSTVLLYQSPTTGLFPTKTFGDNQKAKVQDSLYCAACAWALALAYRRIDDDKGRTHELEHSAIKCMRGILYCCMRQADKVQQFKQDPRPSKCLHSVFSAHSGDEVFSHDEYGHLQINAVSLFLLYLVEMISSGLQIIYNTDEVSFIQNLVFCVERAYRVPDFGIWERGSKYNNGSTELHSSSVGLAKAALEAINGFNLFGNQGCSWSIIFVDLDAHNRNGQTLCSLLPRESRSHNTDAALLPCISYPAFAVDDDALYSQTLDKIVRKLKGKYGFKRFLRDGYRTSLEDRNRRYYKPAEIKVMEYVNGTHNIGNPQQVKEYQELLDHLLQHSSEGYPVVPKYYYVPADFVELEKKNPGSQKRFPSNSGRDGKFFLWGQALYIIAKLLVDGLVNPKDIDPVGRYIPPQDQRNVSMRYSNQGPLENDLVVHVALIAESQRLQVFLNTYGIQTQTPQQVEPIQIWAQQELVKAYFHLGINDKLGLSGRPDRPIGCLGTSKIYRILGKTVVCYPIIFDLSDFYMSQDVMLLIDDIKNALQFIKQYWKMHGRPLFLVLIREDNIRGSRFNPILDMLAAFKKGVVGGVKVHADRVQTLISGAVVEQLDFLRISETEEPPDFKSFEELELPKHSKVKRQTSTPNVSELEQQPEVNINDWKIRSTYEILQKLHDCNCLASQAMLLSILLKREGPNFITKEGTISDHIERIYRRAGSKKLWFVYNIFVSPHITTLLVHGKQVKCTEVLNSAYELKIRAGDMPPRDLYQLSPSEVKQLLLDILQPQQQGRCWLNRRQIDGSLNRTPHGFYDRVWQILERTPNGVIVAEKYLPQQPTLSDMTMYEMNFSLLVEDMLQNIDQPEYRQIIVELLMVVSVILERNPELEFQDKVDLDKLVKEAFHYFQKDEIRLKGVEKQDDMTSFYNTPPMGKRGTCSYLTKAVMNLLLEGEVKPCSDDPCTIS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IV63</accession>
    <name>A0A452IV63_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="13">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1 10">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IV63"/>
    <dbReference type="Ensembl" id="ENSGAGT00000036292.1">
        <property type="protein sequence ID" value="ENSGAGP00000032011.1"/>
        <property type="gene ID" value="ENSGAGG00000022196.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005903">
        <property type="term" value="C:brush border"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005902">
        <property type="term" value="C:microvillus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016459">
        <property type="term" value="C:myosin complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000146">
        <property type="term" value="F:microfilament motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007015">
        <property type="term" value="P:actin filament organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030048">
        <property type="term" value="P:actin filament-based movement"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006897">
        <property type="term" value="P:endocytosis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd23767">
        <property type="entry name" value="IQCD"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.4820:FF:000013">
        <property type="entry name" value="LOW QUALITY PROTEIN: unconventional myosin-Ib"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.190:FF:000007">
        <property type="entry name" value="Myosin-ib isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.530:FF:000004">
        <property type="entry name" value="Unconventional myosin ID"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.720:FF:000004">
        <property type="entry name" value="unconventional myosin-Ib isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.190">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.20.240.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000048">
        <property type="entry name" value="IQ_motif_EF-hand-BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR010926">
        <property type="entry name" value="Myosin_TH1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13140">
        <property type="entry name" value="MYOSIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13140:SF277">
        <property type="entry name" value="UNCONVENTIONAL MYOSIN-IB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00612">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06017">
        <property type="entry name" value="Myosin_TH1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00015">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="4"/>
    </dbReference>
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        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="TH1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9 10" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="5 10" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="6" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="8 10" id="KW-0505">Motor protein</keyword>
    <keyword evidence="7 10" id="KW-0518">Myosin</keyword>
    <keyword evidence="4 10" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="Myosin motor" evidence="11">
        <location>
            <begin position="1"/>
            <end position="712"/>
        </location>
    </feature>
    <feature type="domain" description="TH1" evidence="12">
        <location>
            <begin position="934"/>
            <end position="1118"/>
        </location>
    </feature>
    <feature type="region of interest" description="Actin-binding" evidence="10">
        <location>
            <begin position="589"/>
            <end position="611"/>
        </location>
    </feature>
    <feature type="binding site" evidence="10">
        <location>
            <begin position="131"/>
            <end position="138"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00008314"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023175"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023203"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS51456"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS51757"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000032011.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1118" mass="129438" checksum="510AA7332C3CD21B" modified="2019-05-08" version="1">MGRWSTCNAGRALSQYWRHNNIRTSKRCHGSAPAAALWSFECSQALSKGVKVVSAFYIHGSSLNVVINFQKFMFFSFISLIGKSSVNRYSSIHGYKESKTKYQGQNKLFKINEAYRSLRDQDKDQCILITGESGAGKTEASKLVMSYVAAVCGKGAEVNQVKEQLLQSNPVLEGKNLIMHLENTFLPVLHHSSRSGKYMDIEFDFKGDPLGGVISNFLHISGPKGERAINKDKLKLERDFSRYNYLSLDSARVNGVDDAVNFRTVRNAMQIVGFMDHETQSILEVVAAVLKLGNIEFKPESRVNGLDESKIKDKNELKEICELTGIDQSVLERAFSFRTVEAKQEKVSTTLNVAQAYYARDALAKNLYSRLFSWLVTRINESIKAQTKVRKKVMGVLDIYGFEIFEDNSFEQFIINYCNEKLQQIFIELTLKEEQEEYIREDIEWTHIEYFNNAIICDLIENNQTGILAMLDEECLRPGTVTDDTFLEKLNQVCATHQHFESRMSKCSRFLNDTSLPHSCFRIQHYAGKVMYQVEGFVDKNNDLLYRDLSQAMWKANHSLIKALFPEGNPAKINLKRPPTAGSQFKASVATLMKNLQTKNPNYIRCIKPNDKKAAHIFNEALVCHQVRYLGLLENVRVRRAGYAFRQPYELCLERYKMLCKQTWPHWRGPARAGVEVLFNELEIPEEEYSFGRSKIFIRNPRTLFKLEDLRKQRLEDLATLIQKIYRGWKCRTHFLLMKRSQIVIASWYRRYAQQKKYQRIKSSAIVVQSYIRGWKARKLLRELKHQKRCKEAATTIAAYWHGTQARKELRRLKEEARNKHAIAVIWAYWLGYKVRREYRKFFRANAGRKIYEFTLQKIMQKYFLEMKNKLPSLSPIDKNWPARPYLFLDSTHKELKRIFHLWRCKKYRDHFTDQQKLIYEEKLEASELFKDKKALYPASVGQPFHGAYLEISKNPKYKKLKDAVEEKIIIAEVVNKINRANGKCASRIFLLTKNNVLLADQKSGHIKSEVPLGDVTKVSMSSQNDGFFAVHLKEGSGAAGKGDFLFSSDHLIEMATKLYRTTLSQTKQKLNIEISDEFLVQFKQDKVCVKFIQGNQKNGNIPTCKRKNNRLLEVAVP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IV65</accession>
    <name>A0A452IV65_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Myocardin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="16 17">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="17" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="16" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="16" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="10">Smooth muscle cells (SM) and cardiac muscle cells-specific transcriptional factor which uses the canonical single or multiple CArG boxes DNA sequence. Acts as a cofactor of serum response factor (SRF) with the potential to modulate SRF-target genes. Plays a crucial role in cardiogenesis, urinary bladder development, and differentiation of the smooth muscle cell lineage (myogenesis). Positively regulates the transcription of genes involved in vascular smooth muscle contraction.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IV65"/>
    <dbReference type="Ensembl" id="ENSGAGT00000036206.1">
        <property type="protein sequence ID" value="ENSGAGP00000031927.1"/>
        <property type="gene ID" value="ENSGAGG00000022912.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003713">
        <property type="term" value="F:transcription coactivator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0055007">
        <property type="term" value="P:cardiac muscle cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045944">
        <property type="term" value="P:positive regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051147">
        <property type="term" value="P:regulation of muscle cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051145">
        <property type="term" value="P:smooth muscle cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.720.30:FF:000008">
        <property type="entry name" value="Myocardin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.140.2040">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.150.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.720.30">
        <property type="entry name" value="SAP domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043451">
        <property type="entry name" value="Myocardin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004018">
        <property type="entry name" value="RPEL_repeat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003034">
        <property type="entry name" value="SAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036361">
        <property type="entry name" value="SAP_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22793:SF11">
        <property type="entry name" value="MYOCARDIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR22793">
        <property type="entry name" value="MYOCARDIN-RELATED TRANSCRIPTION FACTOR-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02755">
        <property type="entry name" value="RPEL"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02037">
        <property type="entry name" value="SAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00707">
        <property type="entry name" value="RPEL"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00513">
        <property type="entry name" value="SAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF68906">
        <property type="entry name" value="SAP domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51073">
        <property type="entry name" value="RPEL"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50800">
        <property type="entry name" value="SAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0010">Activator</keyword>
    <keyword evidence="6 13" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="9" id="KW-0539">Nucleus</keyword>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="17" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="8" id="KW-0804">Transcription</keyword>
    <keyword evidence="5" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="4" id="KW-0832">Ubl conjugation</keyword>
    <feature type="repeat" description="RPEL" evidence="12">
        <location>
            <begin position="18"/>
            <end position="43"/>
        </location>
    </feature>
    <feature type="repeat" description="RPEL" evidence="12">
        <location>
            <begin position="62"/>
            <end position="87"/>
        </location>
    </feature>
    <feature type="repeat" description="RPEL" evidence="12">
        <location>
            <begin position="106"/>
            <end position="131"/>
        </location>
    </feature>
    <feature type="domain" description="SAP" evidence="15">
        <location>
            <begin position="368"/>
            <end position="402"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="155"/>
            <end position="282"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="319"/>
            <end position="346"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="451"/>
            <end position="482"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="14">
        <location>
            <begin position="629"/>
            <end position="694"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="13">
        <location>
            <begin position="522"/>
            <end position="556"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="173"/>
            <end position="229"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="14">
        <location>
            <begin position="248"/>
            <end position="265"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="14">
        <location>
            <begin position="451"/>
            <end position="464"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="465"/>
            <end position="482"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="629"/>
            <end position="652"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="14">
        <location>
            <begin position="660"/>
            <end position="671"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00057404"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00069373"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00401"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="PROSITE" id="PS50800"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000031927.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="895" mass="97804" checksum="E1D70AA7ADF66072" modified="2019-05-08" version="1">MTLLGSEHSLLIRSKFRSVLQLRLQQRRTREQLADQGIMPPLKSPSAFHEQRKNLERAKTEDYLKHKIRSRPEQSDLVNMHILQDSAAEGSIQATQMKLKRARLADDLNEKIALRPGPLELVEKNIIPVDSAVKEAIKGAQGSFSKSADAFAFEEDSSNDGLSPEQTRREDLQGSTESPAATKNSEAISSATSGTTQDHSRTSENNVTDLTSVQNSQCASPKQVNGQQSPPLPMPAVVKSKSSSDSKNRHKKPKDTKPKVKKLKYHQYIPPDQKAEKSPPPMDSAYARLLQQQQLFLQLQILSQQQQQHFSYPGIHQAQQKQPSEQMVKSSNSSSTPVTNMPLSPVKTTFSGQTCVSSIKPGPLPSNLDDLKVSELRQQLRIRGLPVSGTKTALMERLRPFQDCSGSVVPNFNEITTVTFPVTPTNTLSSYQSQPSASMLSSSFYQFGSTSSSPPISPASSDLSVSGSLPDTFNDGPMTSPQFGLQPSPVHGSAEESLMGSMNGGSIQLELEGIDTEKDKMLVEKQKVINELTWKLQQEQRQVEELRMQLQKRKRNNGLEEKQPPTQHFFGVPIKQENTVSSCPFSSKQTALKAQANSLDKLSNCGTPQLPHIVNSLCMESAGQSTITSSTFLSPQCSPQHSPLGTAKSPQHISLPPSPNSHYLLSVSPGSQGEGRSVSPQASNRLRPAPQMTRSQQMDELLDVLIESGEMPAIAKEDRSCLQKVPQITVSSGNASASLPKASAPFEHVSSTQLSFDHCTGSNDTHLEVLLNSQSPLGKVSEIPLLKMGGEESHFEGLIDGFSSKVADELLTSQEIPLSPMETQLSPSSAEGSGLQMSFTESPWETMEWLDLTPPSSATAFGSFTTTGPNLFNIDFLDVTDLSLNTTMDLHLQQW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IV69</accession>
    <name>A0A452IV69_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Neurofibromin</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="14">Neurofibromatosis-related protein NF-1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="11">Stimulates the GTPase activity of Ras. NF1 shows greater affinity for Ras GAP, but lower specific activity. May be a regulator of Ras activity.</text>
    </comment>
    <comment type="subunit">
        <text evidence="12">Interacts with HTR6. Interacts with SPRED2.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
            <location evidence="2">Nucleolus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000036252.1">
        <property type="protein sequence ID" value="ENSGAGP00000031973.1"/>
        <property type="gene ID" value="ENSGAGG00000022899.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005730">
        <property type="term" value="C:nucleolus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008289">
        <property type="term" value="F:lipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050793">
        <property type="term" value="P:regulation of developmental process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="CDD" id="cd13313">
        <property type="entry name" value="PH_NF1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd05130">
        <property type="entry name" value="RasGAP_Neurofibromin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00170">
        <property type="entry name" value="SEC14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000070">
        <property type="entry name" value="Neurofibromin 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.525.10:FF:000004">
        <property type="entry name" value="Neurofibromin 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.506.10:FF:000015">
        <property type="entry name" value="Neurofibromin isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.506.10:FF:000023">
        <property type="entry name" value="Neurofibromin isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.525.10">
        <property type="entry name" value="CRAL-TRIO lipid binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.506.10">
        <property type="entry name" value="GTPase Activation - p120gap, domain 1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001251">
        <property type="entry name" value="CRAL-TRIO_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036865">
        <property type="entry name" value="CRAL-TRIO_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR054071">
        <property type="entry name" value="PH_NF1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039360">
        <property type="entry name" value="Ras_GTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023152">
        <property type="entry name" value="RasGAP_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001936">
        <property type="entry name" value="RasGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10194:SF142">
        <property type="entry name" value="NEUROFIBROMIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10194">
        <property type="entry name" value="RAS GTPASE-ACTIVATING PROTEINS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13716">
        <property type="entry name" value="CRAL_TRIO_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21877">
        <property type="entry name" value="PH_NF1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00616">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00323">
        <property type="entry name" value="RasGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00516">
        <property type="entry name" value="SEC14"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52087">
        <property type="entry name" value="CRAL/TRIO domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50191">
        <property type="entry name" value="CRAL_TRIO"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00509">
        <property type="entry name" value="RAS_GTPASE_ACTIV_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50018">
        <property type="entry name" value="RAS_GTPASE_ACTIV_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0007">Acetylation</keyword>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="3" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="8" id="KW-0446">Lipid-binding</keyword>
    <keyword evidence="9" id="KW-0472">Membrane</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="Ras-GAP" evidence="16">
        <location>
            <begin position="1223"/>
            <end position="1418"/>
        </location>
    </feature>
    <feature type="domain" description="CRAL-TRIO" evidence="17">
        <location>
            <begin position="1547"/>
            <end position="1705"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="2750"/>
            <end position="2805"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="15">
        <location>
            <begin position="2767"/>
            <end position="2793"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="15">
        <location>
            <begin position="2796"/>
            <end position="2805"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004604"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023121"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00059799"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00062501"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00074788"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00081423"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="PROSITE" id="PS50018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS50191"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000031973.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2805" mass="316298" checksum="C7BCA3C9C8E9A63F" modified="2019-05-08" version="1">HVSRCLFNFILPIKAGQQNTHTKVSTEHNKECLINISKYKFSLVISGLTNILKNVNNMRIFGEAAEKNLYLSQLIILDTLEKCLAGQPKDTMRLDETMLVKQLLPEICHFIHTYREGNQHAAELRNSASGVLFSLSCNNFNAVFSRISTRLQELTVCSEDNADVHDIELLQYISVDCIKLKRILQETVFKFKALKKVAQLAVINSLEKAFWNWVENYPDEFTKLYQTPQTDMADCAEKLFDLVDGFAESTKRKAAVWPLQIILLVLCPEIIQDIAKDVVEETKINKKLFLDNLRKALASHSGSRQLTESAAIACVKLCKASTYINWEDNSVIFLLVQSMVDDLKTLLFNPSKPFSRGNQNADVDLMIDCLVSCFRINPQNNPYLKICLAQNSPSTFHYVLVNSLQRIITNSEVDWWPKIDAVYCHSVELRSMFSETLQKAVQSCGAHPAIRMTPSLTFKEKMTSLKFKEKPTDLETRSYKFLLLSMVKLLHADPKLLLYNPRKQGPETQSSTAELITGLVQLVPQSNMPEIAQEAMEALLVLHQLDSIDLWNPDAPIETFWEISSQMLFYICKKLTGHQMLSSTEILKWLREILICRNKFLLKNKQSDRSSCHFLFLYGVGCDVSGGGASQLSLDHEEMRCAPGATLRKGKGNSSMESTAGCSGTPICRQAQTKLEVALYMFLWSPDIEAVLVAMSCFRHLCEEADIRCGVDEVSVHNFLPNYNTFMEFASVSNMMSTGRAALQKRVMALLRRIEHPTAGNTEAWEDTHAKWEQATKLILNYPKTKMEDGQVTDSLHKTIVRRRMSHVSGGGSIDLSDTDSLQEWINMTGFLCALGGVCLQHRSNAGLATYSPPMGPINERKGSMISMISTEGNAETPVSKFIDRLLSLMVCNHEKVGLQIRTNIKDLVGLELSPALYPMLFNKLKNTISKFFDSQGQVLLTETNTQFVEQTIAIMKNLLDNHTDGSSEHLGQASIETMMLNLVRYVRVLGNLVHAIQIKTKLCQLVEVMMERRDDLSFCQEMKFRNKMVEYLTDWVMGTSNQATDEDVKCLTRDLDQASMEAVVSLLAGLPLQPEEGDGVELMEAKSQLFLKYFTLFMNLLNDCSEVEDESAQTGGRKRGMSRRLASLRHCTVLAMSNLLNANVDSGLMHSIGLGYHKDLQTRATFMEVLTKILQQGTEFDTLAETVLADRFERLVELVTMMGDQGELPIAMALANVVPCSQWDELARVLVTLFDSRHLLYQLLWNMFSKEVELADSMQTLFRGNSLASKIMTFCFKVYGATYLQKLLEPLLRTVIMSSEWQHVSFEVDPTRLEHTESLEANQRSLLQMTEKFFHAIINSSSEFPPQLRSVCHCLYQVVSQRFPQNSIGAVGSAMFLRFINPAIVSPYEAGILDKKPPPRIERGLKLMSKILQSIANHVLFTKEEHMRPFNDFVKSNFDAARRFFLDIASDCPASDTVNHSLSFISDGNVLALHRLLWNNQEKIGQYLSSNRDHKAVGRRPFDKMATLLAYLGPPEHKPVADTHWSSLNLTSSKFEEFMTRHQVHEKEEFKALKTLNIFYQAGTSKAGNPVFYYIARRFKTGQINGDLLIYHVLLTLKPYYAKPYEIVVDLTHAGPSNRFKTDFLSKWFVVFPGFAYENVSAVYIYNCNSWVREYTKYHERLLTGLKGSKRLIFIDSPGKLAEHIEHDQQKLPAATLALEEDLKVFHNALKLAHKDTKVSIKVGSTAVQVTSADRTRVLGQTVFLNDIYYASEIEEICLVDENQFTLTIANQGTPLTFMHQECEAIVQSIIHIRTRWELSQPDSIPQHTKIRPKDVPGTLLNIALLNLGSSDPSLRSAAYNLLCALTCTFNLKIEGQLLETSGLCIPANNTLFIVSISKTLAANEPHLTLEFLEECISGFSKSSIELKHLCLEYMTPWLSNLVRFCKHNDDAKRQRVTAILDKLITMTINEKQMYPSIQAKIWGSLGQITDLLDVVLDSFIKTSATGGLGSIKAEVMADTAVALASGNVKLVSSKVIGRMCKIIDKTCLSPTPTLEQHLMWDDIAILARYMLMLSFNNSLDVAAHLPYLFHVVTLLVATGPLSLRASTHGLVINIIHSLCTCSQLHFSEETKQVLRLSLTEFSLPKFYLLFGISKVKSAAVIAFRSSYRDRSFSPGSYERETFALTSLETVTEALLEIMEACMRDIPTCKWLDQWTDLAQKFAFQYNPSLQPRALVVFGCISKRVSHGQIKQIIRILSKGLESCLKGPDNYNSQVLIEATVIALTKLQPLLNKDSPMHKSLFWVAMAVLQLDEVNLYSAGTALLEQNLHTLDSLRVFNDKSPEEVFMEIRRPLEWHCKQMDHFVGLNFNSNFNFALVGHLLKGYRHPSPTIVARTVRILHTLLALVNKHRNCDKFEVNTQSVAYLAALLTVSEEVRSRCSLKHRKSLLLTEVSMENVPMDTYPIRHSDTSCRTLKENQPWSSPKGSDRHLAANYPTVGQISPRTRKSMSLDMGQPSQANTKKLLGTRKSFDHLISDTKAPKRQEMESGITTPPKMRRVAENDYDMETQRISPQQHPHLRKVSVSESNVLLDEEVLTDPKIQALLLTVLATLVKYTTDEFDQRILYEYLAEASVVFPKVFPVVHNLLDSKINTLLSLCQDPNLLNPIHGIVQSVVYHEESPPQYQTSYLQSFGFNGLWRFAGPFSKQTQIPDYAELIVKFLDALIDTYLPGIDEETSEESLLTPTSPYPPAVQSQLSITANLNLSNSMTSLATSQHSPGIDKENVELSPTTGHSNSGRTRHGSASQVQKQRSAGSFKRHSSKKIV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J6T3</accession>
    <name>A0A452J6T3_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="3">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the N-CoR nuclear receptor corepressors family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000040981.1">
        <property type="protein sequence ID" value="ENSGAGP00000036201.1"/>
        <property type="gene ID" value="ENSGAGG00000025594.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000785">
        <property type="term" value="C:chromatin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046966">
        <property type="term" value="F:nuclear thyroid hormone receptor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003714">
        <property type="term" value="F:transcription corepressor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000122">
        <property type="term" value="P:negative regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051571">
        <property type="entry name" value="N-CoR_corepressor"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13992">
        <property type="entry name" value="NUCLEAR RECEPTOR CO-REPRESSOR RELATED NCOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13992:SF5">
        <property type="entry name" value="NUCLEAR RECEPTOR COREPRESSOR 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="28"/>
            <end position="52"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="66"/>
            <end position="88"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="139"/>
            <end position="160"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="464"/>
            <end position="515"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="645"/>
            <end position="725"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="806"/>
            <end position="889"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="911"/>
            <end position="1004"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1039"/>
            <end position="1118"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1130"/>
            <end position="1149"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1250"/>
            <end position="1321"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="1357"/>
            <end position="1378"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="66"/>
            <end position="81"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="139"/>
            <end position="153"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="670"/>
            <end position="695"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="806"/>
            <end position="820"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="2">
        <location>
            <begin position="824"/>
            <end position="835"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="2">
        <location>
            <begin position="855"/>
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            <dbReference type="ARBA" id="ARBA00010097"/>
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    <sequence length="1401" mass="152272" checksum="E1DCBF2B5ABB6ABC" modified="2019-05-08" version="1">MNAARYSVPPVLQPAPHQVIANIPEGVRLPTTRPTRPPPPLIPSSKTTMPSEKPSFIMGGSISQGTPGTYLTSHSQASYSQEPAKPSVGSISLGLPRQQESAKPASIPYIKQEEFSPRSQNSQPEGLLVRAQHEGVVRGTTTTIQEGSITRGTPANKVPVETIPSLRGSITQGTPALSQSGIAADALLKGTITRLATEDSSPEKCREEAAAKGHVIYEGKSGHILSYDRLICRTLPRGSPHAELKERTVLSGSIMQGTPRATTESFEEGLKYPKQIKRESPPIRTFEGAITKGKPYDGVTTIKEMGRSIHEIPRQDLLSQESRKTPEIVQSARPIMEGSISQGTPIKYESNSGQSAIKHNVKSLITGPSKLPRGMSQLEMVPENVKAVERAKYEDVKTVEAVRSRHTSVVSSGTSVLRSTLHEAPKSQLSPGIYDDTNARRTPVNYQSPMSRSSPMMNRVTEAGISSGKPANHERKTTLTPTQRESVPAKSPVSGVDPVVTHSPFDSHHRGSTPEVYRSHLPPHLDSAMQFHRALDPAAAAYLFQRQLSPTPGYPSQYQLYAMENTRQTILNDYITSQQMQVNLRPDVTRGLSPREQTLALPYAGARGIIDLNNMAPTILVPHPGGTSTPPMERITYIPGTQLTFPTRPYNPASMSPGHPSHLASAAANAEREREKERERERERERERERERERIAAVPAEHYLRPGTEQPGRPGSHGYVRSPSPSVRTQENMLQQRPSIFQGTNGTSVITPLDPAAQLRIMQLTPGAPSITPGLPASRYNTAADALAALVDAAASAPQMEVVKAKEHKHEGTRIEENMGRRSTVVSEQQQQMEQKTPETEKRSVQCPYTSAAFSSGKSQGPSSSVVYSEAGKDKGPPPKSRYEEELRTRGKTTITAANFIDVIITRQIASDKDARDRGSQSSDSSSSLSSHRYEAPGDAIEVISPANSPVPPQEKLQSYQLEAPKPSQAESKQYEGPVHRYRTQQESPSPQQPPLPSSQAEGIGHVPRTHRLITLADHICQIITQDFARNQVASQVSLQPPTSTFQNSAPASATTPTRAKASNRYSPESQSVHHQRPSARVSPENLSDKSRARPGKSPERSHVSSEPYEPISPPQVPVVHEKQDNIILLSQRNEPAEQRTDSRSPGSISYLPSFFTKLENTSPMVKSKKQEIFRKLNSSGGGDSDMATAQPGTEIFNLPAVTTSGAVSSRGHSFADPASNLGLEDIIRKALMGSFDDKGEDHGVVMSQTMGIAPGSSNPAVSANSEARREEANPSPNSGGVGKQKLIGKSNSRKSKSPIPGQGYLGTERPSSVSSVHSEGDYHRQTPVWAWEDRPSSTGSTQFPYNPLTMRMLSSTPPTSIACAPPSMSQAATHQQNRIWEREPAPLLSAQYETLSDSDD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J6T5</accession>
    <name>A0A452J6T5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Nuclear receptor corepressor 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="17">Mediates transcriptional repression by certain nuclear receptors. Part of a complex which promotes histone deacetylation and the formation of repressive chromatin structures which may impede the access of basal transcription factors. Participates in the transcriptional repressor activity produced by BCL6. Recruited by ZBTB7A to the androgen response elements/ARE on target genes, negatively regulates androgen receptor signaling and androgen-induced cell proliferation. Mediates the NR1D1-dependent repression and circadian regulation of TSHB expression. The NCOR1-HDAC3 complex regulates the circadian expression of the core clock gene ARTNL/BMAL1 and the genes involved in lipid metabolism in the liver.</text>
    </comment>
    <comment type="subunit">
        <text evidence="18">Forms a large corepressor complex that contains SIN3A/B and histone deacetylases HDAC1 and HDAC2. This complex associates with the thyroid receptor (TR) and the retinoid acid receptor (RAR) in the absence of ligand. Interacts directly with RARA; the interaction is facilitated with RARA trimethylation. Component of the N-Cor repressor complex, at least composed of CBFA2T3, HEXIM1, NCOR1, NCOR2, HDAC3, TBL1X, TBL1XR1, CORO2A and GPS2. Interacts with ZBTB33; the interaction serves to recruit the N-CoR complex to promoter regions containing methylated CpG dinucleotides. Interacts with TRIM28 and KDM3A. Interacts (via the RD1 domain) with BAZ1A (via its N-terminal); the interaction corepresses a number of NCOR1-regulated genes. Interacts with BCL6, C1D, DACH1, HEXIM1, HDAC7, RORA, RORC, SAP30, SIAH2, SIN3A and SIN3B. May interact with DEAF1. Interacts with RXRA. Interacts with SETD5. Interacts with VDR. Interacts with ZBTB7A. Interacts with AR. Interacts with HDAC3.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the N-CoR nuclear receptor corepressors family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000040974.1">
        <property type="protein sequence ID" value="ENSGAGP00000036194.1"/>
        <property type="gene ID" value="ENSGAGG00000025594.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000785">
        <property type="term" value="C:chromatin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005654">
        <property type="term" value="C:nucleoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017053">
        <property type="term" value="C:transcription repressor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046966">
        <property type="term" value="F:nuclear thyroid hormone receptor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003714">
        <property type="term" value="F:transcription corepressor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
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        <property type="term" value="P:chromatin organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:1902532">
        <property type="term" value="P:negative regulation of intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
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        <property type="term" value="P:negative regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
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        <property type="term" value="P:rhythmic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
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        <property type="match status" value="2"/>
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        <property type="entry name" value="nuclear receptor corepressor 1 isoform X4"/>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="N-CoR_GPS2_interact"/>
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    </dbReference>
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    </dbReference>
    <dbReference type="PANTHER" id="PTHR13992">
        <property type="entry name" value="NUCLEAR RECEPTOR CO-REPRESSOR RELATED NCOR"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="NUCLEAR RECEPTOR COREPRESSOR 1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
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        <property type="match status" value="2"/>
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        <property type="match status" value="2"/>
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    <keyword evidence="9" id="KW-0007">Acetylation</keyword>
    <keyword evidence="12" id="KW-0090">Biological rhythms</keyword>
    <keyword evidence="8" id="KW-0156">Chromatin regulator</keyword>
    <keyword evidence="11" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="13" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="4" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="15" id="KW-0539">Nucleus</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0678">Repressor</keyword>
    <keyword evidence="14" id="KW-0804">Transcription</keyword>
    <keyword evidence="10" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="7" id="KW-0832">Ubl conjugation</keyword>
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    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="1921"/>
            <end position="1939"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="19">
        <location>
            <begin position="1970"/>
            <end position="1981"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="2089"/>
            <end position="2098"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="19">
        <location>
            <begin position="2099"/>
            <end position="2113"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="2114"/>
            <end position="2123"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="2137"/>
            <end position="2154"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="2300"/>
            <end position="2316"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="2419"/>
            <end position="2429"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010097"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022491"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023108"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00044171"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00054129"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00065205"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50090"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="PROSITE" id="PS51293"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="PROSITE" id="PS51294"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000036194.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2452" mass="272603" checksum="A1EB10A030F0CE29" modified="2019-05-08" version="1">MSSSGYPPNQGAFSTEQNRYPTHSVQYTFSNPRHQQEFAVPDYRSSHLEVSQATQLLQQQQQQQQQLRRRPSLLSEFHPGSDRPQERRTGYEQQFHPGPSQTDHDSLESKRPRLEQVSDSHFQRVSAAAVLPLVHPLQEGLRSADIKKEPAFGGKHEGPSSPVSGQPCGEDQNASPSKLSKEELIQSMDRVDREIAKVEQQILKLKKKQQQLEEEAAKPPEPERPVSPPPVEQKHRSIVQIIYDENRPLYNQPSDTKVYHENIKTNQVMRKKLILFFKRRNHARKQREQKICQRYDQLMEAWEKKVDRIENNPRRKAKESKTREYYEKQFPEIRKQREQQERFQRVGQRGAGLSATIARSEHEISEIIDGLSEQENNEKQMRQLSVIPPMMFDAEQRRVKFINMNGLMEDPMKVYKDRQFMNVWTDHEKEIFKEKFVQHPKNFGLIASYLERKNVPDCVLYYYLTKKNENYKALVRRNYGKRRGRNQQQIARPSQEEKVEEKVEEEKAEKTEKKEEEKKDEEEKDEKEESKENTKEKDKTEIAAEETEQATPRGRKTANSQGRRKGRITRSMTNEAAAANAAAAATEEPPPPLPPPPEPSSAEPVETSRWTEEEMEIAKKGLVEHGRNWAAIAKMVGTKSEAQCKNFYFNYKRRHNLDNLLQQHKQKSSRRPREERDVSQCESVASTVSAQEDEDIEASNEEENPEDSEGAENSSDTESAPSPSPVEASKPSEDAMSESSASRITTETTAEQESAIKTVPSTSPSSAAQSTQTAENQNIKLQVKEEISNEAEEPMEMDSRSHPAEAKPVLSLPVHTKAEPVDAEMRLPENIQVKIESDTKERDLEKPKEMSEPEEMDFSLSQQSNTARPEPNSDNDSSATCSADEDVDGEPERQRIFTMDSKPSLLNPTGSILVSSTIKQGQMDLQQLHHRAAVIPPMVSCSPCNVSVGTPISGYALYQRHIKAMHESALLEEQRQRQEHLDMEYRSSASPCGTSKSPNVEWEGKSVAYMPYVEVKRAIEQEAQVQNTAGRSASPYRLSPREISKASPQPDMNAARYSVPPVLQPAPHQVIANIPEGVRLPTTRPTRPPPPLIPSSKTTMPSEKPSFIMGGSISQQEEFSPRSQNSQPEGLLVRAQHEGVVRGTTTTIQEGSITRGTPANKVPVETIPSLRGSITQGTPALSQSGIAADALLKGTITRLATEDSSPEKCREEAAAKGHVIYEGKSGHILSYDTIKNVREGTRSPRAAHEISLKRSYDTMEGNIKQGMSMRESPVSAPLEGLICRTLPRGSPHAELKERTVLSGSIMQGTPRATTESFEEGLKYPKQIKRESPPIRTFEGAITKGKPYDGVTTIKEMGRSIHEIPRQDLLSQESRKTPEIVQSARPIMEGSISQGTPIKYESNSGQSAIKHNVKSLITGPSKLPRGMSQLEMVPENVKAVERAKYEDVKTVEAVRSRHTSVVSSGTSVLRSTLHEAPKSQLSPGIYDDTNARRTPVNYQSPMSRSSPMMNRVTEGMLSFLPANHERKTTLTPTQRESVPAKSPVSGVDPVVTHSPFDSHHRGSTPEVYRSHLPPHLDSAMQFHRALDPAAAAYLFQRQLSPTPGYPSQYQLYAMENTRQTILNDYITSQQMQVNLRPDVTRGLSPREQTLALPYAGARGIIDLNNMAPTILVPHPGGTSTPPMERITYIPGTQLTFPTRPYNPASMSPGHPSHLASAAANAEREREKERERERERERERERERERIAAVPAEHYLRPGTEQPGRPGSHGYVRSPSPSVRTQENMLQQRPSIFQGTNGTSVITPLDPAAQLRIMQLTPGAPSITPGLPASRYNTAADALAALVDAAASAPQMEVVKAKEHKHEGTRIEENMGRRSTVVSEQQQQMEQKTPETEKRSVQCPYTSAAFSSGKSQGPSSSVVYSEAGKDKGPPPKSRYEEELRTRGKTTITAANFIDVIITRQIASDKDARDRGSQSSDSSSSLSSHRYEAPGDAIEVISPANSPVPPQEKLQSYQLEAPKPSQAESKSYEGPVHRYRTQQESPSPQQPPLPSSQAEGIGHVPRTHRLITLADHICQIITQDFARNQVASQVSLQPPTSTFQNSAPASATTPTRAKASNRYSPESQSVHHQRPSARVSPENLSDKSRARPGKSPERSHVSSEPYEPISPPQVPVVHEKQDNIILLSQRNEPAEQRTDSRSPGSISYLPSFFTKLENTSPMVKSKKQEIFRKLNSSGGGDSDMATAQPGTEIFNLPAVTTSGAVSSRGHSFADPASNLGLEDIIRKALMGSFDDKGEDHGVVMSQTMGIAPGSSNPAVSANSEARREEANPSPNSGGGVGKQKLIGKSNSRKSKSPIPGQGYLGTERPSSVSSVHSEGDYHRQTPVWAWEDRPSSTGSTQFPYNPLTMRMLSSTPPTSIACAPPSMSQAATHQQNRIWEREPAPLLSAQYETLSDSDD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J6U2</accession>
    <name>A0A452J6U2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Nuclear receptor corepressor 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="23 24">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="24" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="23" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="23" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="17">Mediates transcriptional repression by certain nuclear receptors. Part of a complex which promotes histone deacetylation and the formation of repressive chromatin structures which may impede the access of basal transcription factors. Participates in the transcriptional repressor activity produced by BCL6. Recruited by ZBTB7A to the androgen response elements/ARE on target genes, negatively regulates androgen receptor signaling and androgen-induced cell proliferation. Mediates the NR1D1-dependent repression and circadian regulation of TSHB expression. The NCOR1-HDAC3 complex regulates the circadian expression of the core clock gene ARTNL/BMAL1 and the genes involved in lipid metabolism in the liver.</text>
    </comment>
    <comment type="subunit">
        <text evidence="18">Forms a large corepressor complex that contains SIN3A/B and histone deacetylases HDAC1 and HDAC2. This complex associates with the thyroid receptor (TR) and the retinoid acid receptor (RAR) in the absence of ligand. Interacts directly with RARA; the interaction is facilitated with RARA trimethylation. Component of the N-Cor repressor complex, at least composed of CBFA2T3, HEXIM1, NCOR1, NCOR2, HDAC3, TBL1X, TBL1XR1, CORO2A and GPS2. Interacts with ZBTB33; the interaction serves to recruit the N-CoR complex to promoter regions containing methylated CpG dinucleotides. Interacts with TRIM28 and KDM3A. Interacts (via the RD1 domain) with BAZ1A (via its N-terminal); the interaction corepresses a number of NCOR1-regulated genes. Interacts with BCL6, C1D, DACH1, HEXIM1, HDAC7, RORA, RORC, SAP30, SIAH2, SIN3A and SIN3B. May interact with DEAF1. Interacts with RXRA. Interacts with SETD5. Interacts with VDR. Interacts with ZBTB7A. Interacts with AR. Interacts with HDAC3.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the N-CoR nuclear receptor corepressors family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000040980.1">
        <property type="protein sequence ID" value="ENSGAGP00000036200.1"/>
        <property type="gene ID" value="ENSGAGG00000025594.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000785">
        <property type="term" value="C:chromatin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005654">
        <property type="term" value="C:nucleoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017053">
        <property type="term" value="C:transcription repressor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046966">
        <property type="term" value="F:nuclear thyroid hormone receptor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003714">
        <property type="term" value="F:transcription corepressor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006325">
        <property type="term" value="P:chromatin organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:1902532">
        <property type="term" value="P:negative regulation of intracellular signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000122">
        <property type="term" value="P:negative regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048511">
        <property type="term" value="P:rhythmic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00167">
        <property type="entry name" value="SANT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.1880:FF:000004">
        <property type="entry name" value="nuclear receptor corepressor 1 isoform X4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.60:FF:000026">
        <property type="entry name" value="Nuclear receptor corepressor 2 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.430:FF:000001">
        <property type="entry name" value="Nuclear receptor corepressor 2 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.430">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.1880">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.60">
        <property type="entry name" value="Homeodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009057">
        <property type="entry name" value="Homeodomain-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017930">
        <property type="entry name" value="Myb_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051571">
        <property type="entry name" value="N-CoR_corepressor"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031557">
        <property type="entry name" value="N-CoR_GPS2_interact"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001005">
        <property type="entry name" value="SANT/Myb"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017884">
        <property type="entry name" value="SANT_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13992">
        <property type="entry name" value="NUCLEAR RECEPTOR CO-REPRESSOR RELATED NCOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13992:SF5">
        <property type="entry name" value="NUCLEAR RECEPTOR COREPRESSOR 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15784">
        <property type="entry name" value="GPS2_interact"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00249">
        <property type="entry name" value="Myb_DNA-binding"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00717">
        <property type="entry name" value="SANT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF46689">
        <property type="entry name" value="Homeodomain-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51294">
        <property type="entry name" value="HTH_MYB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50090">
        <property type="entry name" value="MYB_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51293">
        <property type="entry name" value="SANT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0007">Acetylation</keyword>
    <keyword evidence="12" id="KW-0090">Biological rhythms</keyword>
    <keyword evidence="8" id="KW-0156">Chromatin regulator</keyword>
    <keyword evidence="11" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="13" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="4" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="15" id="KW-0539">Nucleus</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="24" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0678">Repressor</keyword>
    <keyword evidence="14" id="KW-0804">Transcription</keyword>
    <keyword evidence="10" id="KW-0805">Transcription regulation</keyword>
    <keyword evidence="7" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="SANT" evidence="21">
        <location>
            <begin position="307"/>
            <end position="358"/>
        </location>
    </feature>
    <feature type="domain" description="Myb-like" evidence="20">
        <location>
            <begin position="489"/>
            <end position="539"/>
        </location>
    </feature>
    <feature type="domain" description="SANT" evidence="21">
        <location>
            <begin position="492"/>
            <end position="543"/>
        </location>
    </feature>
    <feature type="domain" description="HTH myb-type" evidence="22">
        <location>
            <begin position="496"/>
            <end position="543"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1"/>
            <end position="74"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="97"/>
            <end position="121"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="369"/>
            <end position="512"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="546"/>
            <end position="779"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="908"/>
            <end position="932"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1019"/>
            <end position="1039"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1391"/>
            <end position="1442"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1572"/>
            <end position="1652"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1733"/>
            <end position="1816"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1838"/>
            <end position="1934"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1969"/>
            <end position="2048"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="2060"/>
            <end position="2079"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="2180"/>
            <end position="2251"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
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            <begin position="2287"/>
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        <location>
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            <dbReference type="ARBA" id="ARBA00065205"/>
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            <dbReference type="SAM" id="MobiDB-lite"/>
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    <evidence type="ECO:0000259" key="20">
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            <dbReference type="PROSITE" id="PS50090"/>
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    <sequence length="2331" mass="258402" checksum="EEC39EF839D6030A" modified="2019-05-08" version="1">MSSSGYPPNQGAFSTEQNRYPTHSVQYTFSNPRHQQEPAFGGKHEGPSSPVSGQPCGEDQNASPSKLSKEELIQSMDRVDREIAKVEQQILKLKKKQQQLEEEAAKPPEPERPVSPPPVEQKHRSIVQIIYDENRPLYNQPSDTKVYHENIKTNQVMRKKLILFFKRRNHARKQREQKICQRYDQLMEAWEKKVDRIENNPRRKAKESKTREYYEKQFPEIRKQREQQERFQRVGQRGAGLSATIARSEHEISEIIDGLSEQENNEKQMRQLSVIPPMMFDAEQRRVKFINMNGLMEDPMKVYKDRQFMNVWTDHEKEIFKEKFVQHPKNFGLIASYLERKNVPDCVLYYYLTKKNENYKALVRRNYGKRRGRNQQIARPSQEEKVEEKVEEEKAEKTEKKEEEKKDEEEKDEKEESKENTKEKDKTEIAAEETEQATPRGRKTANSQGRRKGRITRSMTNEAAAANAAAAATEEPPPPLPPPPEPSSAEPVETSRWTEEEMEIAKKGLVEHGRNWAAIAKMVGTKSEAQCKNFYFNYKRRHNLDNLLQQHKQKSSRRPREERDVSQCESVASTVSAQEDEDIEASNEEENPEDSEGAENSSDTESAPSPSPVEASKPSEDAMSESSASRITTETTAEQESAIKTVPSTSPSSAAQSTQTAENQNIKLQVKEEISNEAEEPMEMDSRSHPAEAKPVLSLPVHTKAEPVDAEMRLPENIQVKIESDTKERDLEKPKEMSEPEEMDFSLSQQSNTARPEPNSDNDSSATCSADEDVDGEPERQRIFTMDSKPSLLNPTGSILVSSTIKQGQMDLQQLHHRAAVIPPMVSCSPCNVSVGTPISGYALYQRHIKAMHESALLEEQRQRQEHLDMEYRSSASPCGTSKSPNVEWEVLQPAPHQVIANIPEGVRLPTTRPTRPPPPLIPSSKTTMPSEKPSFIMGGSISQGTPGTYLTSHSQASYSQEPAKPSVGSISLGLPRQQESAKPASIPYIKQEEFSPRSQNSQPEGLLVRAQHEGVVRGTTTTIQEGSITRGTPANKVPVETIPSLRGSITQGTPALSQSGIAADALLKGTITRLATEDSSPEKCREEAAAKGHVIYEGKSGHILSYDTIKNVREGTRSPRAAHEISLKRSYDTMEGNIKQGMSMRESPVSAPLEGLICRTLPRGSPHAELKERTVLSGSIMQGTPRATTESFEEGLKYPKQIKRESPPIRTFEGAITKGKPYDGVTTIKEMGRSIHEIPRQDLLSQESRKTPEIVQSARPIMEGSISQGTPIKYESNSGQSAIKHNVKSLITGPSKLPRGMSQLEMVPENVKAVERAKYEDVKTVEAVRSRHTSVVSSGTSVLRSTLHEAPKSQLSPGIYDDTNARRTPVNYQSPMSRSSPMMNRVTEAGISSGKPANHERKTTLTPTQRESVPAKSPVSGVDPVVTHSPFDSHHRGSTPEVYRSHLPPHLDSAMQFHRALDPAAAAYLFQRQLSPTPGYPSQYQLYAMENTRQTILNDYITSQQMQVNLRPDVTRGLSPREQTLALPYAGARGIIDLNNMAPTILVPHPGGTSTPPMERITYIPGTQLTFPTRPYNPASMSPGHPSHLASAAANAEREREKERERERERERERERERERIAAVPAEHYLRPGTEQPGRPGSHGYVRSPSPSVRTQENMLQQRPSIFQGTNGTSVITPLDPAAQLRIMQLTPGAPSITPGLPASRYNTAADALAALVDAAASAPQMEVVKAKEHKHEGTRIEENMGRRSTVVSEQQQQMEQKTPETEKRSVQCPYTSAAFSSGKSQGPSSSVVYSEAGKDKGPPPKSRYEEELRTRGKTTITAANFIDVIITRQIASDKDARDRGSQSSDSSSSLSSHRYEAPGDAIEVISPANSPVPPQEKLQSYQLEAPKPSQAESDPNRQYEGPVHRYRTQQESPSPQQPPLPSSQAEGIGHVPRTHRLITLADHICQIITQDFARNQVASQVSLQPPTSTFQNSAPASATTPTRAKASNRYSPESQSVHHQRPSARVSPENLSDKSRARPGKSPERSHVSSEPYEPISPPQVPVVHEKQDNIILLSQRNEPAEQRTDSRSPGSISYLPSFFTKLENTSPMVKSKKQEIFRKLNSSGGGDSDMATAQPGTEIFNLPAVTTSGAVSSRGHSFADPASNLGLEDIIRKALMGSFDDKGEDHGVVMSQTMGIAPGSSNPAVSANSEARREEANPSPNSGGVGKQKLIGKSNSRKSKSPIPGQGYLGTERPSSVSSVHSEGDYHRQTPVWAWEDRPSSTGSTQFPYNPLTMRMLSSTPPTSIACAPPSMSQAATHQQNRIWEREPAPLLSAQYETLSDSDD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J6U8</accession>
    <name>A0A452J6U8_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="13">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the N-CoR nuclear receptor corepressors family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000040985.1">
        <property type="protein sequence ID" value="ENSGAGP00000036205.1"/>
        <property type="gene ID" value="ENSGAGG00000025594.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="term" value="C:chromatin"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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        <property type="term" value="C:nucleoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032991">
        <property type="term" value="C:protein-containing complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046966">
        <property type="term" value="F:nuclear thyroid hormone receptor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003714">
        <property type="term" value="F:transcription corepressor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000122">
        <property type="term" value="P:negative regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00167">
        <property type="entry name" value="SANT"/>
        <property type="match status" value="2"/>
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        <property type="entry name" value="Nuclear receptor corepressor 2 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Nuclear receptor corepressor 2 isoform 1"/>
        <property type="match status" value="1"/>
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        <property type="match status" value="1"/>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Homeodomain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Homeodomain-like_sf"/>
    </dbReference>
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        <property type="entry name" value="Myb_dom"/>
    </dbReference>
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        <property type="entry name" value="N-CoR_corepressor"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031557">
        <property type="entry name" value="N-CoR_GPS2_interact"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001005">
        <property type="entry name" value="SANT/Myb"/>
    </dbReference>
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        <property type="entry name" value="SANT_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13992">
        <property type="entry name" value="NUCLEAR RECEPTOR CO-REPRESSOR RELATED NCOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13992:SF5">
        <property type="entry name" value="NUCLEAR RECEPTOR COREPRESSOR 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15784">
        <property type="entry name" value="GPS2_interact"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00249">
        <property type="entry name" value="Myb_DNA-binding"/>
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    </dbReference>
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        <property type="match status" value="2"/>
    </dbReference>
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        <property type="match status" value="2"/>
    </dbReference>
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        <property type="entry name" value="HTH_MYB"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="SANT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="6" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="9" id="KW-0539">Nucleus</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0678">Repressor</keyword>
    <keyword evidence="8" id="KW-0804">Transcription</keyword>
    <keyword evidence="5" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="SANT" evidence="11">
        <location>
            <begin position="419"/>
            <end position="470"/>
        </location>
    </feature>
    <feature type="domain" description="HTH myb-type" evidence="12">
        <location>
            <begin position="602"/>
            <end position="640"/>
        </location>
    </feature>
    <feature type="domain" description="SANT" evidence="11">
        <location>
            <begin position="602"/>
            <end position="640"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1"/>
            <end position="120"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="149"/>
            <end position="180"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
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            <begin position="209"/>
            <end position="233"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="480"/>
            <end position="598"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="643"/>
            <end position="876"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1005"/>
            <end position="1029"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1116"/>
            <end position="1136"/>
        </location>
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    <feature type="region of interest" description="Disordered" evidence="10">
        <location>
            <begin position="1488"/>
            <end position="1539"/>
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        <location>
            <begin position="1669"/>
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        <location>
            <begin position="1830"/>
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            <begin position="1935"/>
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        <location>
            <begin position="2066"/>
            <end position="2145"/>
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    </feature>
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        <location>
            <begin position="1"/>
            <end position="33"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
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</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J753</accession>
    <name>A0A452J753_9SAUR</name>
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        <recommendedName>
            <fullName evidence="11">SANT domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
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        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
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    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
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    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
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        <subcellularLocation>
            <location evidence="1">Nucleus</location>
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        <text evidence="2">Belongs to the N-CoR nuclear receptor corepressors family.</text>
    </comment>
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    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0678">Repressor</keyword>
    <keyword evidence="8" id="KW-0804">Transcription</keyword>
    <keyword evidence="5" id="KW-0805">Transcription regulation</keyword>
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        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="792"/>
            <end position="808"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="816"/>
            <end position="838"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="1089"/>
            <end position="1103"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="1653"/>
            <end position="1678"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="1789"/>
            <end position="1803"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="1807"/>
            <end position="1818"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="1838"/>
            <end position="1848"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="1854"/>
            <end position="1872"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="1903"/>
            <end position="1914"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="2022"/>
            <end position="2031"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="10">
        <location>
            <begin position="2032"/>
            <end position="2046"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="2047"/>
            <end position="2056"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="10">
        <location>
            <begin position="2070"/>
            <end position="2087"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="2233"/>
            <end position="2249"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="10">
        <location>
            <begin position="2351"/>
            <end position="2361"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010097"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022491"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS51293"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000036199.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2384" mass="264417" checksum="FEFAA658A26E86B2" modified="2019-05-08" version="1">MSSSGYPPNQGAFSTEQNRYPTHSVQYTFSNPRHQQEFAVPDYRSSHLEVSQATQLLQQQQQQQQQLRRRPSLLSEFHPGSDRPQERRTGYEQQFHPGPSQTDHDSLESKRPRLEQVSDSHFQRVSAAAVLPLVHPLQEGLRSADIKKEPAFGGKHEGPSSPVSGQPCGEDQNASPSKLSKEELIQSMDRVDREIAKVEQQILKLKKKQQQLEEEAAKPPEPERPVSPPPVEQKHRSIVQIIYDENRPLYNQPSDTKVYHENIKTNQVMRKKLILFFKRRNHARKQREQKICQRYDQLMEAWEKKVDRIENNPRRKAKESKTREYYEKQFPEIRKQREQQERFQRVGQRGAGLSATIARSEHEISEIIDGLSEQENNEKQMRQLSVIPPMMFDAEQRRVKFINMNGLMEDPMKVYKDRQFMNVWTDHEKEIFKEKFVQHPKNFGLIASYLERKNVPDCVLYYYLTKKNENYKALVRRNYGKRRGRNQQIARPSQEEKVEEKVEEEKAEKTEKKEEEKKDEEEKDEKEESKENTKEKDKTEIAAEETEQATPRGRKTANSQGRRKGRITRSMTNEAAAANAAAAATEEPPPPLPPPPEPSSAEPVETSRWTEEEMEIAKKGNPSKSSRRPREERDVSQCESVASTVSAQEDEDIEASNEEENPEDSEGAENSSDTESAPSPSPVEASKPSEDAMSESSASRITTETTAEQESAIKTVPSTSPSSAAQSTQTAENQNIKLQVKEEISNEAEEPMEMDSRSHPAEAKPVLSLPVHTKAEPVDAEMRLPENIQVKIESDTKERDLEKPKEMSEPEEMDFSLSQQSNTARPEPNSDNDSSATCSADEDVDGEPERQRIFTMDSKPSLLNPTGSILVSSTIKQGQMDLQQLHHRAAVIPPMVSCSPCNVSVGTPISGYALYQRHIKAMHESALLEEQRQRQEHLDMEYRSSASPCGTSKSPNVEWEVLQPAPHQVIANIPEGVRLPTTRPTRPPPPLIPSSKTTMPSEKPSFIMGGSISQGTPGTYLTSHSQASYSQEPAKPSVGSISLGLPRQQESAKPASIPYIKQEEFSPRSQNSQPEGLLVRAQHEGVVRGTTTTIQEGSITRGTPANKVPVETIPSLRGSITQGTPALSQSGIAADALLKGTITRLATEDSSPEKCREEAAAKGHVIYEGKSGHILSYDTIKNVREGTRSPRAAHEISLKRSYDTMEGNIKQGMSMRESPVSAPLEELKERTVLSGSIMQGTPRATTESFEEGLKYPKQIKRESPPIRTFEGAITKGKPYDGVTTIKEMGRSIHEIPRQDLLSQESRKTPEIVQSARPIMEGSISQGTPIKYESNSGQSAIKHNVKSLITGPSKLPRGMSQLEMVPENVKAVERAKYEDVKTVEAVRSRHTSVVSSGTSVLRSTLHEAPKSQLSPGIYDDTNARRTPVNYQSPMSRSSPMMNRVTEAGISSGKPANHERKTTLTPTQRESVPAKSPVSGVDPVVTHSPFDSHHRGSTPEVYRSHLPPHLDSAMQFHRALDPAAAAYLFQRQLSPTPGYPSQYQLYAMENTRQTILNDYITSQQMQVNLRPDVTRGLSPREQTLALPYAGARGIIDLNNMAPTILVPHPGGTSTPPMERITYIPGTQLTFPTRPYNPASMSPGHPSHLASAAANAEREREKERERERERERERERERERIAAVPAEHYLRPGTEQPGRPGSHGYVRSPSPSVRTQENMLQQRPSIFQGTNGTSVITPLDPAAQLRIMQLTPGAPSITPGLPASRYNTAADALAALVDAAASAPQMEVVKAKEHKHEGTRIEENMGRRSTVVSEQQQQMEQKTPETEKRSVQCPYTSAAFSSGKSQGPSSSVVYSEAGKDKGPPPKSRYEEELRTRGKTTITAANFIDVIITRQIASDKDARDRGSQSSDSSSSLSSHRYEAPGDAIEVISPANSPVPPQEKLQSYQLEAPKPSQAESKQYEGPVHRYRTQQESPSPQQPPLPSSQAEGIGHVPRTHRLITLADHICQIITQDFARNQVASQVSLQPPTSTFQNSAPASATTPTRAKASNRYSPESQSVHHQRPSARVSPENLSDKSRARPGKSPERSHVSSEPYEPISPPQVPVVHEKQDNIILLSQRNEPAEQRTDSRSPGSISYLPSFFTKLENTSPMVKSKKQEIFRKLNSSGGGDSDMATAQPGTEIFNLPAVTTSGAVSSRGHSFADPASNLGLEDIIRKALMGSFDDKGEDHGVVMSQTMGIAPGSSNPAVSANSEARREEANPSPNSGGVGKQKLIGKSNSRKSKSPIPGQGYLGTERPSSVSSVHSEGDYHRQTPVWAWEDRPSSTGSTQFPYNPLTMRMLSSTPPTSIACAPPSMSQAATHQQNRIWEREPAPLLSAQYETLSDSDD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J762</accession>
    <name>A0A452J762_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">40S ribosomal protein S4</fullName>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1 6">Belongs to the eukaryotic ribosomal protein eS4 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452J762"/>
    <dbReference type="Ensembl" id="ENSGAGT00000040990.1">
        <property type="protein sequence ID" value="ENSGAGP00000036209.1"/>
        <property type="gene ID" value="ENSGAGG00000025649.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0022627">
        <property type="term" value="C:cytosolic small ribosomal subunit"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019843">
        <property type="term" value="F:rRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003735">
        <property type="term" value="F:structural constituent of ribosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006412">
        <property type="term" value="P:translation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd06087">
        <property type="entry name" value="KOW_RPS4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00165">
        <property type="entry name" value="S4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.30:FF:000005">
        <property type="entry name" value="40S ribosomal protein S4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.50.740:FF:000001">
        <property type="entry name" value="40S ribosomal protein S4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.290.10:FF:000051">
        <property type="entry name" value="40S ribosomal protein S4, X isoform"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.50.740">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.290.10">
        <property type="entry name" value="RNA-binding S4 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="HAMAP" id="MF_00485">
        <property type="entry name" value="Ribosomal_eS4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005824">
        <property type="entry name" value="KOW"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014722">
        <property type="entry name" value="Rib_uL2_dom2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000876">
        <property type="entry name" value="Ribosomal_eS4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032277">
        <property type="entry name" value="Ribosomal_eS4_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013845">
        <property type="entry name" value="Ribosomal_eS4_central_region"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038237">
        <property type="entry name" value="Ribosomal_eS4_central_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041982">
        <property type="entry name" value="Ribosomal_eS4_KOW"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013843">
        <property type="entry name" value="Ribosomal_eS4_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002942">
        <property type="entry name" value="S4_RNA-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036986">
        <property type="entry name" value="S4_RNA-bd_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11581">
        <property type="entry name" value="30S/40S RIBOSOMAL PROTEIN S4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11581:SF0">
        <property type="entry name" value="SMALL RIBOSOMAL SUBUNIT PROTEIN ES4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16121">
        <property type="entry name" value="40S_S4_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00467">
        <property type="entry name" value="KOW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00900">
        <property type="entry name" value="Ribosomal_S4e"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08071">
        <property type="entry name" value="RS4NT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF002116">
        <property type="entry name" value="Ribosomal_S4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00363">
        <property type="entry name" value="S4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50889">
        <property type="entry name" value="S4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 6" id="KW-0687">Ribonucleoprotein</keyword>
    <keyword evidence="4 6" id="KW-0689">Ribosomal protein</keyword>
    <keyword evidence="3 6 7" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="2 6" id="KW-0699">rRNA-binding</keyword>
    <feature type="domain" description="RNA-binding S4" evidence="8">
        <location>
            <begin position="24"/>
            <end position="88"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007500"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022730"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022980"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023274"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PIRNR" id="PIRNR002116"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00182"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="SMART" id="SM00363"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000036209.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="245" mass="27531" checksum="3757AF084BEEBC08" modified="2019-05-08" version="1">MLDKLTGVFAPRPSTGPHKLRECLPLIIFLRNRLKYALTGDEVKKICMQRFIKIDGKVRTDITYPAGFMDVISIEKTGEHFRLVYDTKGRFAVHRITAEEAKYKLCKVRKIFVGTKGIPHLVTHDARTIRYPDPLIKVNDTIQIDLETGKITDFIKFDTGNLCMVTGGANLGRIGVITNRERHPGSFDVVHVKDANGNSFATRLSNIFVIGKGNKPWISLPRGKGIRLTIAEERDKRLAAKQSSS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J765</accession>
    <name>A0A452J765_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="23">Tyrosine-protein kinase receptor</fullName>
            <ecNumber evidence="23">2.7.10.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="28 29">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="29" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="28" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="28" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="18 23">
            <text>L-tyrosyl-[protein] + ATP = O-phospho-L-tyrosyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10596"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.10.1"/>
        </reaction>
    </comment>
    <comment type="subunit">
        <text evidence="19">Tetramer of 2 alpha and 2 beta chains linked by disulfide bonds. The alpha chains contribute to the formation of the ligand-binding domain, while the beta chain carries the kinase domain. Interacts with PIK3R1 and with the PTB/PID domains of IRS1 and SHC1 in vitro when autophosphorylated on tyrosine residues. Forms a hybrid receptor with INSR, the hybrid is a tetramer consisting of 1 alpha chain and 1 beta chain of INSR and 1 alpha chain and 1 beta chain of IGF1R. Interacts with ARRB1 and ARRB2. Interacts with GRB10. Interacts with RACK1. Interacts with SOCS1, SOCS2 and SOCS3. Interacts with 14-3-3 proteins. Interacts with NMD2. Interacts with MAP3K5. Interacts with STAT3. Found in a ternary complex with IGF1 and ITGAV:ITGB3 or ITGA6:ITGB4. Interacts (nascent precursor form) with ZFAND2B.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="23">Belongs to the protein kinase superfamily. Tyr protein kinase family. Insulin receptor subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.10.1" evidence="23"/>
    <dbReference type="AlphaFoldDB" id="A0A452J765"/>
    <dbReference type="Ensembl" id="ENSGAGT00000041160.1">
        <property type="protein sequence ID" value="ENSGAGP00000036350.1"/>
        <property type="gene ID" value="ENSGAGG00000025684.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030424">
        <property type="term" value="C:axon"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005899">
        <property type="term" value="C:insulin receptor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005009">
        <property type="term" value="F:insulin receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043560">
        <property type="term" value="F:insulin receptor substrate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005010">
        <property type="term" value="F:insulin-like growth factor receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043548">
        <property type="term" value="F:phosphatidylinositol 3-kinase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071333">
        <property type="term" value="P:cellular response to glucose stimulus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043410">
        <property type="term" value="P:positive regulation of MAPK cascade"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051897">
        <property type="term" value="P:positive regulation of phosphatidylinositol 3-kinase/protein kinase B signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046328">
        <property type="term" value="P:regulation of JNK cascade"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="CDD" id="cd00064">
        <property type="entry name" value="FU"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000050">
        <property type="entry name" value="Tyrosine-protein kinase receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.220.10:FF:000007">
        <property type="entry name" value="Tyrosine-protein kinase receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000087">
        <property type="entry name" value="Tyrosine-protein kinase receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000108">
        <property type="entry name" value="Tyrosine-protein kinase receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:002452">
        <property type="entry name" value="Tyrosine-protein kinase receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.200.20:FF:000026">
        <property type="entry name" value="Tyrosine-protein kinase receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.20.20:FF:000001">
        <property type="entry name" value="Tyrosine-protein kinase receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.80.20.20:FF:000020">
        <property type="entry name" value="Tyrosine-protein kinase receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.220.10">
        <property type="entry name" value="Hormone Receptor, Insulin-like Growth Factor Receptor 1, Chain A, domain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.80.20.20">
        <property type="entry name" value="Receptor L-domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006211">
        <property type="entry name" value="Furin-like_Cys-rich_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006212">
        <property type="entry name" value="Furin_repeat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009030">
        <property type="entry name" value="Growth_fac_rcpt_cys_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017441">
        <property type="entry name" value="Protein_kinase_ATP_BS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000494">
        <property type="entry name" value="Rcpt_L-dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036941">
        <property type="entry name" value="Rcpt_L-dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050122">
        <property type="entry name" value="RTK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001245">
        <property type="entry name" value="Ser-Thr/Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008266">
        <property type="entry name" value="Tyr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020635">
        <property type="entry name" value="Tyr_kinase_cat_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016246">
        <property type="entry name" value="Tyr_kinase_insulin-like_rcpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002011">
        <property type="entry name" value="Tyr_kinase_rcpt_2_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416:SF106">
        <property type="entry name" value="INSULIN-LIKE GROWTH FACTOR 1 RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24416">
        <property type="entry name" value="TYROSINE-PROTEIN KINASE RECEPTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00757">
        <property type="entry name" value="Furin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07714">
        <property type="entry name" value="PK_Tyr_Ser-Thr"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01030">
        <property type="entry name" value="Recep_L_domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000620">
        <property type="entry name" value="Insulin_receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00109">
        <property type="entry name" value="TYRKINASE"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00261">
        <property type="entry name" value="FU"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00219">
        <property type="entry name" value="TyrKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57184">
        <property type="entry name" value="Growth factor receptor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52058">
        <property type="entry name" value="L domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00109">
        <property type="entry name" value="PROTEIN_KINASE_TYR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00239">
        <property type="entry name" value="RECEPTOR_TYR_KIN_II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="11 21 22" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="5" id="KW-0165">Cleavage on pair of basic residues</keyword>
    <keyword evidence="15" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="17" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10" id="KW-0418">Kinase</keyword>
    <keyword evidence="13 25" id="KW-0472">Membrane</keyword>
    <keyword evidence="9 21 22" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="3 23" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="16 23" id="KW-0675">Receptor</keyword>
    <keyword evidence="29" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
    <keyword evidence="4" id="KW-0808">Transferase</keyword>
    <keyword evidence="6 23 25" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="12 25" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="14" id="KW-0829">Tyrosine-protein kinase</keyword>
    <feature type="transmembrane region" description="Helical" evidence="25">
        <location>
            <begin position="29"/>
            <end position="45"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="25">
        <location>
            <begin position="777"/>
            <end position="802"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="27">
        <location>
            <begin position="333"/>
            <end position="453"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="27">
        <location>
            <begin position="454"/>
            <end position="552"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="27">
        <location>
            <begin position="679"/>
            <end position="772"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="26">
        <location>
            <begin position="843"/>
            <end position="1145"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="24">
        <location>
            <begin position="520"/>
            <end position="543"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="24">
        <location>
            <begin position="1174"/>
            <end position="1233"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="24">
        <location>
            <begin position="529"/>
            <end position="540"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="24">
        <location>
            <begin position="1185"/>
            <end position="1194"/>
        </location>
    </feature>
    <feature type="active site" description="Proton donor/acceptor" evidence="20">
        <location>
            <position position="1006"/>
        </location>
    </feature>
    <feature type="binding site" evidence="21">
        <location>
            <position position="853"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="21 22">
        <location>
            <position position="877"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="21">
        <location>
            <begin position="924"/>
            <end position="930"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="21">
        <location>
            <begin position="1010"/>
            <end position="1011"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="21">
        <location>
            <position position="1024"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022685"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023137"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00051243"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00065231"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="PIRSR" id="PIRSR000620-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="PIRSR" id="PIRSR000620-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="RuleBase" id="RU000312"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="26">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="27">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="28">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000036350.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="29">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1233" mass="140026" checksum="49DEF64A2F0AA5B3" modified="2019-05-08" version="1">FPPIIVIFCFSFLPSVKVLKSCSRPKGICAYRVFTLLFIVLLYFLNASCSIFPCSVFAVCPSACSKRACTDQNECCHPECLGSCTVPDNNTACVACRNYYYEGICVPTCPLNTYKFEGWRCVTKDFCSKVPATEMSDYEKFVIHNDECMAECPSGFIRNGSQSMFCSPCEGPCPKICEEEKTKTIDSVTSAQMLQGCTVLKGNLLINIRRGNNIASELENFMGLIETVTGYVKIRHSHALVSLSFLKNLRYILGEEQLDGNYSFYVLDNQNLQQLWDWNHHNLTIKEGKMYFAFNPKLCVSEIYRMEDVSGTKGRQSKGDINPRNNGERASCESHILNFVSNTTLKNRIKLTWERYRPPDYRDLISFTVYYKEAPFKNVTEYDGQDACGSNSWNMVDVDLPPNKENNPGILLQGLKPWTQYAIYVKAVTLTMMENHHIHGAKSEIVYIRTNASVPSIPLDVISASNTSSQLIVKWNPPSLPNGNLSYYIVRWQQQPQDSYLYRHNYCSKDKVPIRRYADETIDTEEATEPTKPEGSEGEKGPCCACPKTEAEKQAEKEEAEYRKVFEDFLHNSIFVPRPDRKRRDLFRIANTTLSSRNKNTTVPDHFANVSDVEESEMEYPFYESKVEGRERTLISHLQPFTLYRIDIHSCNHEAEKLGCSASNFVFARTMPAEGADNIPGTVTWEGKEDNAIYLKWSEPVNPNGLILMYEIKYGQHGEEKRECVSRQEYRKFGGAKLTHLNPGNYSARVQATSLAGNGSWTEPVSFYVQPKHYGNFLHLIIVLPIAVLLVIGGLLIMLYVFNKKRNSDRLGNGVLYASVNPEYFSASDVYVPDEWEVPREKITMCRELGQGSFGMVYEGIAKGVVKDEPETRVAIKTVNESASMRERIEFLNEASVMKEFNCHHVVRLLGVVSQGQPTLVIMELMTRGDLKSYLRSLRPDTEVRVWVGICDWKQFFPSTSHFYLLILAQNNPGQSPPTLKKMIQMAGEIADGMAYLNANKFVHRDLAARNCMVAEDFTVKIGDFGMTRDIYETDYYRKGGKGLLPVRWMSPESLKDGVFTTHSDVWSFGVVLWEIATLAEQPYQGMSNEQVLRFVMEGGLLEKPDNCPDMLFELMRMCWQYNPKMRPSFLEIIGSIKDELDPAFKEVSFFYSEENKPPDTEELDLETENMESIPLDPSSTLQPSDKHSGHKAENGPGVVVLRASFEERQPYAHMNGGRKNERALPLPQSSAC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J768</accession>
    <name>A0A452J768_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">Protein turtle homolog B</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="22">Immunoglobulin superfamily member 9B</fullName>
        </alternativeName>
    </protein>
    <organism evidence="28 29">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="29" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="28" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="28" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="18">Transmembrane protein which is abundantly expressed in interneurons, where it may regulate inhibitory synapse development. May mediate homophilic cell adhesion.</text>
    </comment>
    <comment type="subunit">
        <text evidence="20">Found in a complex with MAGI2 and NLGN2, where it interacts with MAGI2 (via PDZ 5 and PDZ 6 domains).</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="17">Postsynaptic cell membrane</location>
            <topology evidence="17">Single-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="16">Postsynaptic density</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="19">Belongs to the immunoglobulin superfamily. Turtle family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000041140.1">
        <property type="protein sequence ID" value="ENSGAGP00000036335.1"/>
        <property type="gene ID" value="ENSGAGG00000025752.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098982">
        <property type="term" value="C:GABA-ergic synapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043005">
        <property type="term" value="C:neuron projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0014069">
        <property type="term" value="C:postsynaptic density"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007155">
        <property type="term" value="P:cell adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000272">
        <property type="entry name" value="Immunoglobulin superfamily member 9B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000323">
        <property type="entry name" value="Immunoglobulin superfamily member 9B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000389">
        <property type="entry name" value="Immunoglobulin superfamily member 9B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000531">
        <property type="entry name" value="Immunoglobulin superfamily member 9B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000226">
        <property type="entry name" value="protein turtle homolog B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000245">
        <property type="entry name" value="protein turtle homolog B isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000321">
        <property type="entry name" value="protein turtle homolog B isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013106">
        <property type="entry name" value="Ig_V-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051170">
        <property type="entry name" value="Neural/epithelial_adhesion"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12231">
        <property type="entry name" value="CTX-RELATED TYPE I TRANSMEMBRANE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12231:SF240">
        <property type="entry name" value="PROTEIN TURTLE HOMOLOG B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13927">
        <property type="entry name" value="Ig_3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07686">
        <property type="entry name" value="V-set"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="5"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="1" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="15" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="11 24" id="KW-0472">Membrane</keyword>
    <keyword evidence="2" id="KW-0488">Methylation</keyword>
    <keyword evidence="8" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="14" id="KW-0628">Postsynaptic cell membrane</keyword>
    <keyword evidence="29" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="5 25" id="KW-0732">Signal</keyword>
    <keyword evidence="10" id="KW-0770">Synapse</keyword>
    <keyword evidence="4 24" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="9 24" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="25">
        <location>
            <begin position="1"/>
            <end position="24"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019215229" description="Protein turtle homolog B" evidence="25">
        <location>
            <begin position="25"/>
            <end position="1341"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="24">
        <location>
            <begin position="728"/>
            <end position="753"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="26">
        <location>
            <begin position="43"/>
            <end position="120"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="26">
        <location>
            <begin position="144"/>
            <end position="230"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="26">
        <location>
            <begin position="233"/>
            <end position="325"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="26">
        <location>
            <begin position="329"/>
            <end position="420"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="26">
        <location>
            <begin position="425"/>
            <end position="509"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="27">
        <location>
            <begin position="517"/>
            <end position="609"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="27">
        <location>
            <begin position="619"/>
            <end position="717"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="23">
        <location>
            <begin position="763"/>
            <end position="820"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="23">
        <location>
            <begin position="992"/>
            <end position="1040"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="23">
        <location>
            <begin position="1100"/>
            <end position="1327"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="23">
        <location>
            <begin position="992"/>
            <end position="1001"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="23">
        <location>
            <begin position="1020"/>
            <end position="1035"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="23">
        <location>
            <begin position="1113"/>
            <end position="1122"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="23">
        <location>
            <begin position="1131"/>
            <end position="1143"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="23">
        <location>
            <begin position="1156"/>
            <end position="1167"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="23">
        <location>
            <begin position="1201"/>
            <end position="1219"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="23">
        <location>
            <begin position="1250"/>
            <end position="1267"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="23">
        <location>
            <begin position="1279"/>
            <end position="1289"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="23">
        <location>
            <begin position="1318"/>
            <end position="1327"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023257"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00034105"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00037805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00058005"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00061549"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00062279"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00069171"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00075032"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="26">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="27">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="28">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000036335.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="29">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1341" mass="147452" checksum="A29470B5EDC3F261" modified="2019-05-08" version="1">MVGSTMIWYVATLIASVITTRGLAAQGAHGSREEPEFVTARAGESVILGCDVIHPLTGQPPPYVVEWFKFGVPIPIFIKFGFYPPHVDPEYAGRASLHDKASLRIEQIRSEDQGWYECKVLMLDQQYDTFHNGSWVHLTVNAPPTFTETPPQYVEAKEGSSVTLTCMAFGNPKPIVTWLREGNLLGASEKYQVSDGSLTVMSVSREDRGAYTCRAYSIQGEAMHTTRLLVQGPPFIVSPPENITVNISQDALFTCQAEAYPGNMTYTWYWEEENVYFKNDLKLRVRILIDGTLIIFRVKSDDTGKYTCIPSNSLGRSPSASAYLTVQYPARVVNMPPILYVPIGIHGYIRCPVEAEPPVTFVKWNKDGRPLRFEKYSGWNLLEDGSIRIEEVTEEALGTYTCVPYNALGTMGQSPPARLVLKDPPYFTVLPGWEYRQEAGRELLIPCAAAGDPFPVIAWRKVGKPSRSKHNTLPGGSLQFRSLSKDDHGEWECIATNVVTSITASTHLTVIGTSPHAPGSVHVMVSMTSANVSWEPGYDGGYEQTFSVWMKRAQFGPHDWLSLPVPTGPSWLLVDTLEPETAYQFSVLAQNKLGTSAFSEVVTVNTLAFPVTTPEPLVLVTPPRCLTANRTQQGVLLSWLPPANHSFPIDHYIMEFRVAERWEILDDAILGSDNDFFAKDLSQDTWYEFRVLAVMQDLISEPSNIAGVSSTDIFPPPDLTEDGLARPVLAGIVATICFLAAAILFSTLTACFVNKQRKRKLKRKKDPPLSITHCRKSLESPLSSGKVSPESIRTLRPPSESSDDQGQQGKRMLSPSKEKELSLYKKTKRAISSKKYSVSKAEAEAEATTPIELISRGPDGRFVMDPSEMEPSLKTRRIEGFPFVEETDMYPEFRQSDEENDDPIVPSSVAALKSQLTPLSSSQESYLQPPAYSPRFHRGLEGTGALESHLQATGQARPPAPRPFHHSQFYGYLSSSSPGEMDPPPFYMPEVSPLSSVMSSPPLNPEGPFGHPTIPEENGENASNSTLPLTQTPTGGRSPEPWGRAEFPFSSLEASPMIFPQQLHQCEVAECVQLTACLPRGLPPSSLQVPASYPGILPLEAPKSWTGKPPGRSQPSVPTTTKWQDKPMQPMASQGQLRHTSQGMGIPVLPYHEPSESVSHSGTSTFGLDTRWYEPQPRPRPSPRQVRRAEPSLHQVVLQPSRLSPLTQSPLSSRNSSPELTARARPRPGLIQQAEISEITLQPPAAVSFSRKSTPSTGSPAQSSRGESPSYRPAMAFTSLATGYPSSECPSLPGDSMDVFGQIPSPRRAGEEILRPEPTQTTLSTSGYLGSVVETSFSSAQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452J769</accession>
    <name>A0A452J769_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="9">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
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            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
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        <scope>IDENTIFICATION</scope>
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            <begin position="607"/>
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    <sequence length="1220" mass="134052" checksum="86CD981F7C0510FA" modified="2019-05-08" version="1">MLDQQYDTFHNGSWVHLTVNAPPTFTETPPQYVEAKEGSSVTLTCMAFGNPKPIVTWLREGNLLGASEKYQVSDGSLTVMSVSREDRGAYTCRAYSIQGEAMHTTRLLVQGPPFIVSPPENITVNISQDALFTCQAEAYPGNMTYTWYWEEENVYFKNDLKLRVRILIDGTLIIFRVKSDDTGKYTCIPSNSLGRSPSASAYLTVQYPARVVNMPPILYVPIGIHGYIRCPVEAEPPVTFVKWNKDGRPLRFEKYSGWNLLEDGSIRIEEVTEEALGTYTCVPYNALGTMGQSPPARLVLKDPPYFTVLPGWEYRQEAGRELLIPCAAAGDPFPVIAWRKVGKPSRSKHNTLPGGSLQFRSLSKDDHGEWECIATNVVTSITASTHLTVIGTSPHAPGSVHVMVSMTSANVSWEPGYDGGYEQTFSVWMKRAQFGPHDWLSLPVPTGPSWLLVDTLEPETAYQFSVLAQNKLGTSAFSEVVTVNTLAFPVTTPEPLVLVTPPRCLTANRTQQGVLLSWLPPANHSFPIDHYIMEFRVAERWEILDDAILGSDNDFFAKDLSQDTWYEFRVLAVMQDLISEPSNIAGVSSTDIFPPPDLTEDGLARPVLAGIVATICFLAAAILFSTLTACFVNKQRKRKLKRKKDPPLSITHCRKSLESPLSSGKVSPESIRTLRPPSESSDDQGQQGKRMLSPSKEKELSLYKKTKRAISSKKYSVSKAEAEAEATTPIELISRGPDGRFVMDPSEMEPSLKTRRIEGFPFVEETDMYPEFRQSDEENDDPIVPSSVAALKSQLTPLSSSQESYLQPPAYSPRFHRGLEGTGALESHLQATGQARPPAPRPFHHSQFYGYLSSSSPGEMDPPPFYMPEVSPLSSVMSSPPLNPEGPFGHPTIPEENGENASNSTLPLTQTPTGGRSPEPWGRAEFPFSSLEASPMIFPQQLHQCEVAECVQLTACLPRGLPPSSLQVPASYPGILPLEAPKSWTGKPPGRSQPSVPTTTKWQDKPMQPMASQGQLRHTSQGMGIPVLPYHEPSESVSHSGTSTFGLDTRWYEPQPRPRPSPRQVRRAEPSLHQVVLQPSRLSPLTQSPLSSRNSSPELTARARPRPGLIQQAEISEITLQPPAAVSFSRKSTPSTGSPAQSSRGESPSYRPAMAFTSLATGYPSSECPSLPGDSMDVFGQIPSPRRAGEEILRPEPTQTTLSTSGYLGSVVETSFSSAQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452GP10</accession>
    <name>A0A452GP10_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">Arf-GAP domain and FG repeat-containing protein 1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="17">Nucleoporin-like protein RIP</fullName>
        </alternativeName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasmic vesicle</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452GP10"/>
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        <property type="gene ID" value="ENSGAGG00000002814.1"/>
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        <property type="component" value="Unassembled WGS sequence"/>
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    <dbReference type="GO" id="GO:0031410">
        <property type="term" value="C:cytoplasmic vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
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        <property type="project" value="TreeGrafter"/>
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        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
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        <property type="term" value="F:GTPase activator activity"/>
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        <property type="project" value="InterPro"/>
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        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
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        <property type="term" value="P:acrosome assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
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        <property type="term" value="P:intermediate filament organization"/>
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        <property type="project" value="TreeGrafter"/>
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        <property type="term" value="P:spermatid nucleus differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="entry name" value="ArfGap_AGFG1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Arf-GAP domain and FG repeat-containing protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Arf GTPase activating protein"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Longin domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052248">
        <property type="entry name" value="Arf-GAP_FG-repeat_protein"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037278">
        <property type="entry name" value="ARFGAP/RecO"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001164">
        <property type="entry name" value="ArfGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038508">
        <property type="entry name" value="ArfGAP_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46134:SF3">
        <property type="entry name" value="ARFGAP WITH FG REPEATS 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46134">
        <property type="entry name" value="DRONGO, ISOFORM F"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01412">
        <property type="entry name" value="ArfGap"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00405">
        <property type="entry name" value="REVINTRACTNG"/>
    </dbReference>
    <dbReference type="SMART" id="SM00105">
        <property type="entry name" value="ArfGap"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57863">
        <property type="entry name" value="ArfGap/RecO-like zinc finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50115">
        <property type="entry name" value="ARFGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="15" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="4" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="9" id="KW-0221">Differentiation</keyword>
    <keyword evidence="12" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="6" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="14" id="KW-0539">Nucleus</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="11" id="KW-0744">Spermatogenesis</keyword>
    <keyword evidence="3" id="KW-0813">Transport</keyword>
    <keyword evidence="10" id="KW-0862">Zinc</keyword>
    <keyword evidence="8 18" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="Arf-GAP" evidence="20">
        <location>
            <begin position="11"/>
            <end position="135"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="170"/>
            <end position="195"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004541"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022871"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00069633"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00078983"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50115"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000003493.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="573" mass="59695" checksum="FD75E801657EBB68" modified="2019-05-08" version="1">MAASAKRKQEEKHLKLLREMSSLPPNRKCFDCDQRGPTYTDMTVGSFVCTSCSGILRGLNPPHRVKSISMTTFTQQEIEFLQKHGNEVCKQIWLGLFDDRSSAIPDFRDPQKVKEFLQEKYEKKRWYVPPEQAKVVASVHASISGSSASSTSSTPEVKPLKSLLGESAPALHLNKGTPSQSPVIVRSQGQQQQEKKQFDLLSDLGSDIFAAPAPQSTATANFANFAHFNSHAAQNSANAGFANFDAFGQSSGSSNFGGFPTASQSPFQPQNTAFRTLSSSCSFGEFTSAFPLQAVHSGSAGSMNANFAHFDNFPKSSSADFGAFSASQSNTTATAASKVAVNKPNLQSADKYAALANLDNIFSTGQGGSEQGSGFGTVVTAPSGPAVSAPSQPSASSDNSAFGTVPVAATAQTQPASSSVPASFGATPSTNPFVAAPVAPVAPSTNPFQTNSRGVTGLSGAMHSHIFPQAHFAATFGTASMSMPAGFGTPASYSLPTSFSGNFQQPAFPTQGAFPQQTAFAQQPNGFAAFGQAKPVVTPFGQVAAVGVSSNPFMGAAPTGQFPTGSSSTNPFL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H0P1</accession>
    <name>A0A452H0P1_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="9">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452H0P1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000009228.1">
        <property type="protein sequence ID" value="ENSGAGP00000008009.1"/>
        <property type="gene ID" value="ENSGAGG00000006363.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005834">
        <property type="term" value="C:heterotrimeric G-protein complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001917">
        <property type="term" value="C:photoreceptor inner segment"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001750">
        <property type="term" value="C:photoreceptor outer segment"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001664">
        <property type="term" value="F:G protein-coupled receptor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031683">
        <property type="term" value="F:G-protein beta/gamma-subunit complex binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005525">
        <property type="term" value="F:GTP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003924">
        <property type="term" value="F:GTPase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007188">
        <property type="term" value="P:adenylate cyclase-modulating G protein-coupled receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001580">
        <property type="term" value="P:detection of chemical stimulus involved in sensory perception of bitter taste"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00066">
        <property type="entry name" value="G-alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.400.10:FF:000001">
        <property type="entry name" value="Guanine nucleotide-binding protein G(I) subunit alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000256">
        <property type="entry name" value="Guanine nucleotide-binding protein G(t) subunit alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.400.10">
        <property type="entry name" value="GI Alpha 1, domain 2-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001408">
        <property type="entry name" value="Gprotein_alpha_I"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001019">
        <property type="entry name" value="Gprotein_alpha_su"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011025">
        <property type="entry name" value="GproteinA_insert"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10218">
        <property type="entry name" value="GTP-BINDING PROTEIN ALPHA SUBUNIT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10218:SF68">
        <property type="entry name" value="GUANINE NUCLEOTIDE-BINDING PROTEIN G(T) SUBUNIT ALPHA-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00503">
        <property type="entry name" value="G-alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00318">
        <property type="entry name" value="GPROTEINA"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00441">
        <property type="entry name" value="GPROTEINAI"/>
    </dbReference>
    <dbReference type="SMART" id="SM00275">
        <property type="entry name" value="G_alpha"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47895">
        <property type="entry name" value="Transducin (alpha subunit), insertion domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51882">
        <property type="entry name" value="G_ALPHA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4 7" id="KW-0342">GTP-binding</keyword>
    <keyword evidence="6" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="3 8" id="KW-0460">Magnesium</keyword>
    <keyword evidence="1 8" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="2 7" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0807">Transducer</keyword>
    <feature type="binding site" evidence="7">
        <location>
            <begin position="145"/>
            <end position="146"/>
        </location>
        <ligand>
            <name>GTP</name>
            <dbReference type="ChEBI" id="CHEBI:37565"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="7">
        <location>
            <begin position="170"/>
            <end position="176"/>
        </location>
        <ligand>
            <name>GTP</name>
            <dbReference type="ChEBI" id="CHEBI:37565"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="8">
        <location>
            <position position="176"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="7">
        <location>
            <begin position="195"/>
            <end position="199"/>
        </location>
        <ligand>
            <name>GTP</name>
            <dbReference type="ChEBI" id="CHEBI:37565"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="7">
        <location>
            <begin position="264"/>
            <end position="267"/>
        </location>
        <ligand>
            <name>GTP</name>
            <dbReference type="ChEBI" id="CHEBI:37565"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="7">
        <location>
            <position position="321"/>
        </location>
        <ligand>
            <name>GTP</name>
            <dbReference type="ChEBI" id="CHEBI:37565"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023224"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PIRSR" id="PIRSR601019-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PIRSR" id="PIRSR601019-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000008009.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="349" mass="39672" checksum="2059FEABBE08F033" modified="2019-05-08" version="1">MGSGASAEDKKLAKRSKELEKKLQEDADKEAKTVKLLLLGKHGSWATLQIIHQDGYTQEECLEFKSIIYGNVLQSILAIIRAMSTLGIDYAESGRGDDGRQLCNLADSIEEGTMPLELVDCIKKLWKDAGVQACFDRAAEYQLNDSASYYLNQLDRITAPDYLPNEQDVLRSRVKTTGIIETKFSVKDLNFRMFDVGGQRSERKKWIHCFEGVTCIIFCGALSAYDMVLVEDDEVNRMHESLHLFNSICNHKFFAATSIILFLNKKDLFEEKIKKVHLSICFPDYDGPNTFEDAGNYIKNQFLDLNMRKDVKEIYSHMTCATDTQNVKFVFDAVTDVIIKENLKDCGLF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H0P5</accession>
    <name>A0A452H0P5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">Acetylcholine receptor subunit gamma</fullName>
        </recommendedName>
    </protein>
    <organism evidence="25 26">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="26" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="25" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="25" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">After binding acetylcholine, the AChR responds by an extensive change in conformation that affects all subunits and leads to opening of an ion-conducting channel across the plasma membrane.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>K(+)(in) = K(+)(out)</text>
            <dbReference type="Rhea" id="RHEA:29463"/>
            <dbReference type="ChEBI" id="CHEBI:29103"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>Na(+)(in) = Na(+)(out)</text>
            <dbReference type="Rhea" id="RHEA:34963"/>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
        </reaction>
    </comment>
    <comment type="subunit">
        <text evidence="2">Pentamer of two alpha chains, and one each of the beta, delta, and gamma chains.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="17">Postsynaptic cell membrane</location>
            <topology evidence="17">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="20">Belongs to the ligand-gated ion channel (TC 1.A.9) family. Acetylcholine receptor (TC 1.A.9.1) subfamily. Gamma/CHRNG sub-subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H0P5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000009376.1">
        <property type="protein sequence ID" value="ENSGAGP00000008142.1"/>
        <property type="gene ID" value="ENSGAGG00000006424.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0022848">
        <property type="term" value="F:acetylcholine-gated monoatomic cation-selective channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004888">
        <property type="term" value="F:transmembrane signaling receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd19029">
        <property type="entry name" value="LGIC_ECD_nAChR_G"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd19064">
        <property type="entry name" value="LGIC_TM_nAChR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.390:FF:000010">
        <property type="entry name" value="Nicotinic acetylcholine receptor subunit epsilon"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.390:FF:000036">
        <property type="entry name" value="Nicotinic acetylcholine receptor subunit gamma"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.70.170.10:FF:000012">
        <property type="entry name" value="Nicotinic acetylcholine receptor subunit gamma"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.70.170.10">
        <property type="entry name" value="Neurotransmitter-gated ion-channel ligand-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.390">
        <property type="entry name" value="Neurotransmitter-gated ion-channel transmembrane domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006202">
        <property type="entry name" value="Neur_chan_lig-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036734">
        <property type="entry name" value="Neur_chan_lig-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006201">
        <property type="entry name" value="Neur_channel"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036719">
        <property type="entry name" value="Neuro-gated_channel_TM_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038050">
        <property type="entry name" value="Neuro_actylchol_rec"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006029">
        <property type="entry name" value="Neurotrans-gated_channel_TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018000">
        <property type="entry name" value="Neurotransmitter_ion_chnl_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002394">
        <property type="entry name" value="Nicotinic_acetylcholine_rcpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18945">
        <property type="entry name" value="NEUROTRANSMITTER GATED ION CHANNEL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02931">
        <property type="entry name" value="Neur_chan_LBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02932">
        <property type="entry name" value="Neur_chan_memb"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00254">
        <property type="entry name" value="NICOTINICR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00252">
        <property type="entry name" value="NRIONCHANNEL"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Nicotinic receptor ligand binding domain-like"/>
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    </dbReference>
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        <property type="entry name" value="NEUROTR_ION_CHANNEL"/>
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    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="16 22" id="KW-0407">Ion channel</keyword>
    <keyword evidence="9 22" id="KW-0406">Ion transport</keyword>
    <keyword evidence="15" id="KW-1071">Ligand-gated ion channel</keyword>
    <keyword evidence="10 22" id="KW-0472">Membrane</keyword>
    <keyword evidence="14" id="KW-0628">Postsynaptic cell membrane</keyword>
    <keyword evidence="12" id="KW-0675">Receptor</keyword>
    <keyword evidence="26" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="8" id="KW-0770">Synapse</keyword>
    <keyword evidence="5 22" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 22" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="3 22" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="22">
        <location>
            <begin position="258"/>
            <end position="280"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="22">
        <location>
            <begin position="292"/>
            <end position="312"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="22">
        <location>
            <begin position="324"/>
            <end position="347"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="22">
        <location>
            <begin position="490"/>
            <end position="512"/>
        </location>
    </feature>
    <feature type="domain" description="Neurotransmitter-gated ion-channel ligand-binding" evidence="23">
        <location>
            <begin position="41"/>
            <end position="257"/>
        </location>
    </feature>
    <feature type="domain" description="Neurotransmitter-gated ion-channel transmembrane" evidence="24">
        <location>
            <begin position="264"/>
            <end position="507"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00003328"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00011293"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
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        </source>
    </evidence>
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        </source>
    </evidence>
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        <source>
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        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00034104"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00034430"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00036239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00038482"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00039195"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="RuleBase" id="RU000687"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="Pfam" id="PF02931"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="Pfam" id="PF02932"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000008142.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="529" mass="60505" checksum="B5A06746529C6A5A" modified="2019-05-08" version="1">MQRQHPVPTPSPEWLWGILHRKTPTRTLLAVLGSVLCRNQEEKLLNDLMTNYNRNLRPAQGEGDIINVTLKLTLTNLISLNEREETLTTNIWIEMKWCDYRLQWDPDEYDNIQILRVPSTEVWLPDIVLENNIDGVFEITLYCNTLLDSNGCLLWMPPAIYRTSCAIFVTYFPFDWQNCSMVFQSQTYSANEINLLLMEEDGQTIEWIVIDPEAFTENGEWAIKHRPAKKIVNAERFTPDDLGYQQVVFYLIIQRKPLFYVINIIVPCVLISSVAVLVYFLPAKAGGQKCTVSINVLLAQTVFLFLIAQKVPETSDAMPLIGKYLTFLLVVTVVIVANAVIVLNVSLRTPNTHAMSQKVRQICLRQLPHFLGMHMRRSDMAPRLLLARRRSSLGLMVKADEYMLWKARTELVFEKQKERAGLMKAVLEKIGKGLENRCTQDLCHSLVQAGPEIWACVDACNHIAKTLREQNDFDNENEEWILVGRVIDRVCFLVMASLFVLGTVSIFLMAHFNQAPAAPFPGDPKCYLP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H0Q2</accession>
    <name>A0A452H0Q2_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="4">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000009381.1">
        <property type="protein sequence ID" value="ENSGAGP00000008147.1"/>
        <property type="gene ID" value="ENSGAGG00000006245.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
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        <property type="term" value="C:nucleus"/>
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        <property type="project" value="InterPro"/>
    </dbReference>
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        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039779">
        <property type="entry name" value="RFX-like"/>
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    <dbReference type="InterPro" id="IPR057321">
        <property type="entry name" value="RFX1-4/6/8-like_BCD"/>
    </dbReference>
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        <property type="entry name" value="RFX1_trans_act"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12619:SF17">
        <property type="entry name" value="DNA-BINDING PROTEIN RFX2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12619">
        <property type="entry name" value="RFX TRANSCRIPTION FACTOR FAMILY"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="1"/>
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    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
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        <location>
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            <end position="160"/>
        </location>
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            <begin position="262"/>
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    <feature type="compositionally biased region" description="Polar residues" evidence="1">
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            <begin position="1"/>
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            <begin position="31"/>
            <end position="49"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="281"/>
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            <dbReference type="Pfam" id="PF25340"/>
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    </evidence>
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        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="663" mass="73753" checksum="66571CA0A1E8F516" modified="2019-05-08" version="1">MQNSEGGSDSATSVALRPSASAQAPVVQPVPASQQRVLVQTTSSSPKGAQMQQISIPRVQQVSQQVQTVQHVYPSQVQYVEGGEAVYTNGTIRAAYSYNTEAQIYAPSSAASYFEPQGGGAQVTTAASSPTAIPSHSMVGITMDIGGSQILSSSGAYLIHGGMENTRHSLSLTSRSSPATVCSPEGIVITGPGILEMAIENLQKNEGITSHKSSLLNSHVSYYWTLQPFSCFYHYYGIRLKPESPLNHLQEDTQYMAMRQQPIHQKQRYRPAQKMDGIRESGSNSSLHTTPEQSVAAQSQHHQQFIDVTHVFPDFPAPDLGNVLLQEGVTMNDVKTLQLLYRRHCEATLDVVMNLQFHYIEKLWQSFWSSKTPSSDGSTTMPSSEEEHEGVIPKDKLITLCKYDPILKWMRSCDHILYQALVEILIPDVLRPVPGTLTQAIRNFAKSLEGWLMNAMSDFPPQVVQTKVGVVSAFAQTLRRYTSLNHLAQAARAVLQNTSQINQMLSDLNRVDFANVQEQASWVCQCEEGMVQKLEQDFKLTLQQQSSLDQWASWLDNVVTQVLKHHEDSPSFPKAARQFLLKWSFYSSMVIRDLTLRSAASFGSFHLIRLLYDEYMFYLVEHRVAQATGETPIAVMGEVSTIYDSLIFGXQPLFLKDSLVFGT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H0Q4</accession>
    <name>A0A452H0Q4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">Spartin</fullName>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">Lipophagy receptor that plays an important role in lipid droplet (LD) turnover in motor neurons. Localizes to LDs and interacts with components of the autophagy machinery, such as MAP1LC3A/C proteins to deliver LDs to autophagosomes for degradation via lipophagy. Lipid transfer protein required for lipid droplet degradation, including by lipophagy. Can bind and transfer all lipid species found in lipid droplets, from phospholipids to triglycerides and sterol esters but the direction of lipid transfer by spartin and its cargos are unknown. May be implicated in endosomal trafficking, or microtubule dynamics, or both. Participates in cytokinesis.</text>
    </comment>
    <comment type="subunit">
        <text evidence="16">Interacts with ITCH and WWP1. Interacts (via MIT domain) with IST1; leading to the recruitment of SPART to midbodies. Interacts with MAP1LC3A and MAP1LC3C.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Lipid droplet</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Midbody</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H0Q4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000009148.1">
        <property type="protein sequence ID" value="ENSGAGP00000007937.1"/>
        <property type="gene ID" value="ENSGAGG00000006322.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005811">
        <property type="term" value="C:lipid droplet"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030496">
        <property type="term" value="C:midbody"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008289">
        <property type="term" value="F:lipid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051301">
        <property type="term" value="P:cell division"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016042">
        <property type="term" value="P:lipid catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006869">
        <property type="term" value="P:lipid transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061724">
        <property type="term" value="P:lipophagy"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030514">
        <property type="term" value="P:negative regulation of BMP signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd02679">
        <property type="entry name" value="MIT_spastin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.80:FF:000009">
        <property type="entry name" value="spartin isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.80">
        <property type="entry name" value="Phosphotransferase system, lactose/cellobiose-type IIA subunit"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007330">
        <property type="entry name" value="MIT_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036181">
        <property type="entry name" value="MIT_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009686">
        <property type="entry name" value="Senescence/spartin_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045036">
        <property type="entry name" value="Spartin-like"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21068">
        <property type="entry name" value="SPARTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21068:SF43">
        <property type="entry name" value="SPARTIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF06911">
        <property type="entry name" value="Senescence"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00745">
        <property type="entry name" value="MIT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF116846">
        <property type="entry name" value="MIT domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="11" id="KW-0007">Acetylation</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="10" id="KW-0442">Lipid degradation</keyword>
    <keyword evidence="8" id="KW-0551">Lipid droplet</keyword>
    <keyword evidence="13" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="12" id="KW-0445">Lipid transport</keyword>
    <keyword evidence="14" id="KW-0446">Lipid-binding</keyword>
    <keyword evidence="7" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0813">Transport</keyword>
    <keyword evidence="9" id="KW-0832">Ubl conjugation</keyword>
    <feature type="domain" description="MIT" evidence="19">
        <location>
            <begin position="18"/>
            <end position="96"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="18">
        <location>
            <begin position="142"/>
            <end position="174"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="18">
        <location>
            <begin position="151"/>
            <end position="169"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004214"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004502"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022677"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022963"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023055"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023121"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00054810"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00064034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00067916"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="SMART" id="SM00745"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007937.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="596" mass="64814" checksum="B48F56EAE7A15320" modified="2019-05-08" version="1">MEQEQDPAIPEDANIKAIKEEFKKAFLFVNKGLNTDELGQKEEARNYYKQGLDHLLRGVSIPSQGSECTGSQWDSARQMQQKMQETLQNVRTRMNILEQNTPAMQTNAITMNVPLEGPKLYPVIPTNEKPERPPAPYSLIPLSAPPAENENVATTSKGQLPAMSPSSPKAHSLPHEAPPVYTPQATDGHYTVSYGTDSGEFSSVGEDYYTKCTQPPALESLGVDADELILIPQGVQIFFVSPDGQVSAPSYPGYLRIVKFLDTDGAMAQNHPPAFLQVCDWLYPLMCNQSPVLCCNTGVYMFPDLMSQVPGSHVGVVLSSELPAADRELFEDLLKQMSDLRIQALEASSDVINLAQTVRIQPEPEGGERERELPEWSEKVAHGILSGASWLSWGLVKGAEYTGKAIHKGASKLREHIQPEEKPLEVNPTVAKGLHVAKQATGGAVKVSRFLVEGVCSIASSIGKELAPHVKKHGSKLVPESLKKDKDGKSTFDGAMVVAASGIQGFSTVWQGLERAAKCIANSVSTETVHTVQHKYGDDAGHATDNAMNSAINVGVTAFNIDNIGIKAIVKKTAKETSYAVLDEYKVLEKEKKDGR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H0Q7</accession>
    <name>A0A452H0Q7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Acyl-CoA synthetase short-chain family member 3, mitochondrial</fullName>
            <ecNumber evidence="2">6.2.1.1</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="5">Acetate--CoA ligase 3</fullName>
        </alternativeName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="6">
            <text>butanoate + ATP + CoA = butanoyl-CoA + AMP + diphosphate</text>
            <dbReference type="Rhea" id="RHEA:46172"/>
            <dbReference type="ChEBI" id="CHEBI:17968"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:33019"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57371"/>
            <dbReference type="ChEBI" id="CHEBI:456215"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="6">
            <dbReference type="Rhea" id="RHEA:46173"/>
        </physiologicalReaction>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the ATP-dependent AMP-binding enzyme family.</text>
    </comment>
    <dbReference type="EC" id="6.2.1.1" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452H0Q7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000009386.1">
        <property type="protein sequence ID" value="ENSGAGP00000008152.1"/>
        <property type="gene ID" value="ENSGAGG00000006436.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005759">
        <property type="term" value="C:mitochondrial matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003987">
        <property type="term" value="F:acetate-CoA ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050218">
        <property type="term" value="F:propionate-CoA ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006629">
        <property type="term" value="P:lipid metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.300.30:FF:000017">
        <property type="entry name" value="Acyl-CoA synthetase short-chain family member 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.300.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.12780">
        <property type="entry name" value="N-terminal domain of ligase-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025110">
        <property type="entry name" value="AMP-bd_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045851">
        <property type="entry name" value="AMP-bd_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020845">
        <property type="entry name" value="AMP-binding_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000873">
        <property type="entry name" value="AMP-dep_synth/lig_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042099">
        <property type="entry name" value="ANL_N_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43347">
        <property type="entry name" value="ACYL-COA SYNTHETASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43347:SF3">
        <property type="entry name" value="ACYL-COA SYNTHETASE SHORT-CHAIN FAMILY MEMBER 3, MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00501">
        <property type="entry name" value="AMP-binding"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13193">
        <property type="entry name" value="AMP-binding_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56801">
        <property type="entry name" value="Acetyl-CoA synthetase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00455">
        <property type="entry name" value="AMP_BINDING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="7">
        <location>
            <begin position="1"/>
            <end position="24"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019510151" description="Acyl-CoA synthetase short-chain family member 3, mitochondrial" evidence="7">
        <location>
            <begin position="25"/>
            <end position="553"/>
        </location>
    </feature>
    <feature type="domain" description="AMP-dependent synthetase/ligase" evidence="8">
        <location>
            <begin position="48"/>
            <end position="233"/>
        </location>
    </feature>
    <feature type="domain" description="AMP-dependent synthetase/ligase" evidence="8">
        <location>
            <begin position="239"/>
            <end position="373"/>
        </location>
    </feature>
    <feature type="domain" description="AMP-binding enzyme C-terminal" evidence="9">
        <location>
            <begin position="437"/>
            <end position="515"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00006432"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00013275"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00040004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00042755"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00047935"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF00501"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF13193"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000008152.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="553" mass="60794" checksum="46EE58AEAFB5C14B" modified="2019-05-08" version="1">AARGICRVTLWCSIHAFVTHYALVQESVADGSFELNICYNAIDRHVESGHADKIAIIYDSPVTNTKQAITYKEVLEQVSKLADVMVKYGVKKGDCVVIYMPMIPQTIYTMLACARIGAIHSLIFGGFASKELSSRIDHAKPKLVVTATFGIEPGRRVEYMPLLERALEMIQHQPEKVLIYSRPHLGTVPLVSGRDLDWDEEIAKAKPHDCVAVRSDHPLYILYTSGTTGVPKGKPVGTPDAGAYFRVLAEHGVAAFFTAPTAIRAIRQQDPEADLGKQYCLTRFRTLFLAGERCDVETLEWSKKVFKVPVLDHWWQTETGSAITASCIGLGNSTTPPPGQAGKSVPGYNVMVLDDNKQPVKAKTLGNIVVKLPLPPGAFSGLWKNQETFKELYFQKFPGYYDTMDAGCMDEDGYLYVMSRIDDVINVAGHRISAGAIEESVLSHGAVADCAVVGQEDALKGLVPLALCVLRNGIKIEEEKVLEEIVKRVRESIGPVAAFRKAVFVKQIPKTRSGKIPRSALSALVSGKPYQITPTIEDPGVFKHVEEVLKKQK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H0Q9</accession>
    <name>A0A452H0Q9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2 7">Protein arginine N-methyltransferase 5</fullName>
            <ecNumber evidence="1 7">2.1.1.320</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="7">Arginine methyltransferase that can both catalyze the formation of omega-N monomethylarginine (MMA) and symmetrical dimethylarginine (sDMA).</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="6 7">
            <text>L-arginyl-[protein] + 2 S-adenosyl-L-methionine = N(omega),N(omega)'-dimethyl-L-arginyl-[protein] + 2 S-adenosyl-L-homocysteine + 2 H(+)</text>
            <dbReference type="Rhea" id="RHEA:48108"/>
            <dbReference type="Rhea" id="RHEA-COMP:10532"/>
            <dbReference type="Rhea" id="RHEA-COMP:11992"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:29965"/>
            <dbReference type="ChEBI" id="CHEBI:57856"/>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
            <dbReference type="ChEBI" id="CHEBI:88221"/>
            <dbReference type="EC" id="2.1.1.320"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="7">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="7">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="7">Belongs to the class I-like SAM-binding methyltransferase superfamily.</text>
    </comment>
    <dbReference type="EC" id="2.1.1.320" evidence="1 7"/>
    <dbReference type="AlphaFoldDB" id="A0A452H0Q9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000009153.1">
        <property type="protein sequence ID" value="ENSGAGP00000007942.1"/>
        <property type="gene ID" value="ENSGAGG00000003291.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035243">
        <property type="term" value="F:protein-arginine omega-N symmetric methyltransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032259">
        <property type="term" value="P:methylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006355">
        <property type="term" value="P:regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd02440">
        <property type="entry name" value="AdoMet_MTases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.20.20.150:FF:000008">
        <property type="entry name" value="Protein arginine N-methyltransferase 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.150:FF:000029">
        <property type="entry name" value="Protein arginine N-methyltransferase 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.20.20.150">
        <property type="entry name" value="Divalent-metal-dependent TIM barrel enzymes"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.70.160.11">
        <property type="entry name" value="Hnrnp arginine n-methyltransferase1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.150">
        <property type="entry name" value="Vaccinia Virus protein VP39"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Arg_MeTrfase"/>
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        <property type="entry name" value="Arg_MeTrfase_PRMT5"/>
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        <property type="entry name" value="PRMT5"/>
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        <property type="entry name" value="PRMT5_TIM"/>
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        <property type="entry name" value="SAM-dependent_MTases_sf"/>
    </dbReference>
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        <property type="entry name" value="PROTEIN ARGININE N-METHYLTRANSFERASE 5"/>
        <property type="match status" value="1"/>
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    <dbReference type="PANTHER" id="PTHR10738:SF0">
        <property type="entry name" value="PROTEIN ARGININE N-METHYLTRANSFERASE 5"/>
        <property type="match status" value="1"/>
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    <dbReference type="Pfam" id="PF05185">
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        <property type="entry name" value="Skb1_MeTrfase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53335">
        <property type="entry name" value="S-adenosyl-L-methionine-dependent methyltransferases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51678">
        <property type="entry name" value="SAM_MT_PRMT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="3 7 11" id="KW-0489">Methyltransferase</keyword>
    <keyword evidence="7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5 7 9" id="KW-0949">S-adenosyl-L-methionine</keyword>
    <keyword evidence="4 7 11" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="PRMT5 TIM barrel" evidence="13">
        <location>
            <begin position="18"/>
            <end position="271"/>
        </location>
    </feature>
    <feature type="domain" description="PRMT5 arginine-N-methyltransferase" evidence="12">
        <location>
            <begin position="278"/>
            <end position="445"/>
        </location>
    </feature>
    <feature type="domain" description="PRMT5 oligomerisation" evidence="14">
        <location>
            <begin position="449"/>
            <end position="602"/>
        </location>
    </feature>
    <feature type="active site" description="Proton donor/acceptor" evidence="8">
        <location>
            <position position="416"/>
        </location>
    </feature>
    <feature type="active site" description="Proton donor/acceptor" evidence="8">
        <location>
            <position position="425"/>
        </location>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="305"/>
        </location>
        <ligand>
            <name>S-adenosyl-L-methionine</name>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <begin position="314"/>
            <end position="315"/>
        </location>
        <ligand>
            <name>S-adenosyl-L-methionine</name>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <position position="373"/>
        </location>
        <ligand>
            <name>S-adenosyl-L-methionine</name>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="9">
        <location>
            <begin position="400"/>
            <end position="401"/>
        </location>
        <ligand>
            <name>S-adenosyl-L-methionine</name>
            <dbReference type="ChEBI" id="CHEBI:59789"/>
        </ligand>
    </feature>
    <feature type="site" description="Critical for specifying symmetric addition of methyl groups" evidence="10">
        <location>
            <position position="308"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00011935"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00018777"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022603"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022691"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00048612"/>
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    </evidence>
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        <source>
            <dbReference type="PIRNR" id="PIRNR015894"/>
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    </evidence>
    <evidence type="ECO:0000256" key="8">
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    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PIRSR" id="PIRSR015894-3"/>
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    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01015"/>
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    </evidence>
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        <source>
            <dbReference type="Pfam" id="PF05185"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="Pfam" id="PF17285"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="Pfam" id="PF17286"/>
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    </evidence>
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        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000007942.1"/>
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    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="604" mass="68569" checksum="6FB6E2D2FE79BDA7" modified="2019-05-08" version="1">ATGPTWGTARSRAYTPAGFDFLCMPVFHPRYRREFSHQPARGRPGPHTRSDLLLSGRDWNTLIVGKLSPWIRPDSKVEAVRRNSEAAMLQELNFGAYLGLPAFLLPLTQGENPNLARLLSTHIHTGHHSTMFWMRVPLLAPEDLRDDLIENEPVPVPEDGSGDEKTWIWWHNFRTLCDYNKRIAVALEVGPDLPSNHVIDRWLGEPIKAAILPTSIFLTNKKGFPVLSKMHQRLVFRLLKLEVQFIVWGAHHHPEKEFCSYLQYLEYLSQNRPPPSAYELFAKGYEDYLQSPLQPLMDNLESQTYEVFEKDPIKYSQYQQAIYRCLLDRVPEEEKDTNVQVVMVLGAGRGPLVNATLRASRQASQRVKIYAVEKNPNAVVTLESWQYEEWGSQVTVVSSDMREWSAPEQADLLVSELLGSFADNELSPECLDGAQHCLRAGGVSIPSTYTSFLAPISSSKLYNEVRACREKDRHPEAQFEMPYVVRLHNFHQLAPPQPCFTFHHPHPDPARDKSQYRQLEFAVEVNTVLHGFAGYFETTLYGDIMLSEGGADPLQPLAVRAGERVRVAFWRCAGPRKVWYEWAVTAPGCSALHNPSGRSSTIGL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H110</accession>
    <name>A0A452H110_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">Acetylcholine receptor subunit gamma</fullName>
        </recommendedName>
    </protein>
    <organism evidence="26 27">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="27" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="26" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="26" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="1">After binding acetylcholine, the AChR responds by an extensive change in conformation that affects all subunits and leads to opening of an ion-conducting channel across the plasma membrane.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>K(+)(in) = K(+)(out)</text>
            <dbReference type="Rhea" id="RHEA:29463"/>
            <dbReference type="ChEBI" id="CHEBI:29103"/>
        </reaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>Na(+)(in) = Na(+)(out)</text>
            <dbReference type="Rhea" id="RHEA:34963"/>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
        </reaction>
    </comment>
    <comment type="subunit">
        <text evidence="2">Pentamer of two alpha chains, and one each of the beta, delta, and gamma chains.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="17">Postsynaptic cell membrane</location>
            <topology evidence="17">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="20">Belongs to the ligand-gated ion channel (TC 1.A.9) family. Acetylcholine receptor (TC 1.A.9.1) subfamily. Gamma/CHRNG sub-subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H110"/>
    <dbReference type="Ensembl" id="ENSGAGT00000009375.1">
        <property type="protein sequence ID" value="ENSGAGP00000008141.1"/>
        <property type="gene ID" value="ENSGAGG00000006424.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0022848">
        <property type="term" value="F:acetylcholine-gated monoatomic cation-selective channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004888">
        <property type="term" value="F:transmembrane signaling receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd19064">
        <property type="entry name" value="LGIC_TM_nAChR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.390:FF:000010">
        <property type="entry name" value="Nicotinic acetylcholine receptor subunit epsilon"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.390:FF:000036">
        <property type="entry name" value="Nicotinic acetylcholine receptor subunit gamma"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.70.170.10:FF:000012">
        <property type="entry name" value="Nicotinic acetylcholine receptor subunit gamma"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.70.170.10">
        <property type="entry name" value="Neurotransmitter-gated ion-channel ligand-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.390">
        <property type="entry name" value="Neurotransmitter-gated ion-channel transmembrane domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006202">
        <property type="entry name" value="Neur_chan_lig-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036734">
        <property type="entry name" value="Neur_chan_lig-bd_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006201">
        <property type="entry name" value="Neur_channel"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036719">
        <property type="entry name" value="Neuro-gated_channel_TM_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038050">
        <property type="entry name" value="Neuro_actylchol_rec"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006029">
        <property type="entry name" value="Neurotrans-gated_channel_TM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018000">
        <property type="entry name" value="Neurotransmitter_ion_chnl_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002394">
        <property type="entry name" value="Nicotinic_acetylcholine_rcpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR18945">
        <property type="entry name" value="NEUROTRANSMITTER GATED ION CHANNEL"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Neur_chan_LBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02932">
        <property type="entry name" value="Neur_chan_memb"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00254">
        <property type="entry name" value="NICOTINICR"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00252">
        <property type="entry name" value="NRIONCHANNEL"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90112">
        <property type="entry name" value="Neurotransmitter-gated ion-channel transmembrane pore"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF63712">
        <property type="entry name" value="Nicotinic receptor ligand binding domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00236">
        <property type="entry name" value="NEUROTR_ION_CHANNEL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="11" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="13" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="16 22" id="KW-0407">Ion channel</keyword>
    <keyword evidence="9 22" id="KW-0406">Ion transport</keyword>
    <keyword evidence="15" id="KW-1071">Ligand-gated ion channel</keyword>
    <keyword evidence="10 22" id="KW-0472">Membrane</keyword>
    <keyword evidence="14" id="KW-0628">Postsynaptic cell membrane</keyword>
    <keyword evidence="12" id="KW-0675">Receptor</keyword>
    <keyword evidence="27" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="8" id="KW-0770">Synapse</keyword>
    <keyword evidence="5 22" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 22" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="3 22" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="22">
        <location>
            <begin position="280"/>
            <end position="302"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="22">
        <location>
            <begin position="314"/>
            <end position="334"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="22">
        <location>
            <begin position="346"/>
            <end position="369"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="22">
        <location>
            <begin position="512"/>
            <end position="534"/>
        </location>
    </feature>
    <feature type="domain" description="Neurotransmitter-gated ion-channel ligand-binding" evidence="24">
        <location>
            <begin position="63"/>
            <end position="279"/>
        </location>
    </feature>
    <feature type="domain" description="Neurotransmitter-gated ion-channel transmembrane" evidence="25">
        <location>
            <begin position="286"/>
            <end position="529"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="23">
        <location>
            <begin position="1"/>
            <end position="31"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="23">
        <location>
            <begin position="1"/>
            <end position="15"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00003328"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00011293"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023257"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023286"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00034104"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00034430"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00036239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00038482"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00039195"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="RuleBase" id="RU000687"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="Pfam" id="PF02931"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="Pfam" id="PF02932"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000008141.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="551" mass="61960" checksum="0C3696979AC11A4B" modified="2019-05-08" version="1">MDSSVLGSAAGNQGSSRDHPHPSPSQGWGGEQGVASPLAVFTLYLFLQGLGSSSSLLVLLADEEKLLNDLMTNYNRNLRPAQGEGDIINVTLKLTLTNLISLVSRTMLYSAPSPLFQKWCDYRLQWDPDEYDNIQILRVPSTEVWLPDIVLENNIDGVFEITLYCNTLLDSNGCLLWMPPAIYRTSCAIFVTYFPFDWQNCSMVFQSQTYSANEINLLLMEEDGQTIEWIVIDPEAFTENGEWAIKHRPAKKIVNAERFTPDDLGYQQVVFYLIIQRKPLFYVINIIVPCVLISSVAVLVYFLPAKAGGQKCTVSINVLLAQTVFLFLIAQKVPETSDAMPLIGKYLTFLLVVTVVIVANAVIVLNVSLRTPNTHAMSQKVRQICLRQLPHFLGMHMRRSDMAPRLLLARRRSSLGLMVKADEYMLWKARTELVFEKQKERAGLMKAVLEKIGKGLENRCTQDLCHSLVQAGPEIWACVDACNHIAKTLREQNDFDNENEEWILVGRVIDRVCFLVMASLFVLGTVSIFLMAHFNQAPAAPFPGDPKCYLP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H111</accession>
    <name>A0A452H111_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="8">Transporter</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="8">Belongs to the sodium:neurotransmitter symporter (SNF) (TC 2.A.22) family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H111"/>
    <dbReference type="Ensembl" id="ENSGAGT00000009277.1">
        <property type="protein sequence ID" value="ENSGAGP00000008053.1"/>
        <property type="gene ID" value="ENSGAGG00000006398.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043005">
        <property type="term" value="C:neuron projection"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098793">
        <property type="term" value="C:presynapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="GOC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051378">
        <property type="term" value="F:serotonin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005335">
        <property type="term" value="F:serotonin:sodium:chloride symporter activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006865">
        <property type="term" value="P:amino acid transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000175">
        <property type="entry name" value="Na/ntran_symport"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037272">
        <property type="entry name" value="SNS_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF037979">
        <property type="entry name" value="Na_transp"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11616">
        <property type="entry name" value="SODIUM/CHLORIDE DEPENDENT TRANSPORTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11616:SF129">
        <property type="entry name" value="TRANSPORTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00209">
        <property type="entry name" value="SNF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00176">
        <property type="entry name" value="NANEUSMPORT"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF161070">
        <property type="entry name" value="SNF-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00610">
        <property type="entry name" value="NA_NEUROTRAN_SYMP_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00754">
        <property type="entry name" value="NA_NEUROTRAN_SYMP_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50267">
        <property type="entry name" value="NA_NEUROTRAN_SYMP_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5 10" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0915">Sodium</keyword>
    <keyword evidence="8" id="KW-0769">Symport</keyword>
    <keyword evidence="3 8 10" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="4 10" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="2 8" id="KW-0813">Transport</keyword>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="47"/>
            <end position="65"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="77"/>
            <end position="98"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="119"/>
            <end position="147"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="210"/>
            <end position="227"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="239"/>
            <end position="256"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="287"/>
            <end position="308"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="320"/>
            <end position="345"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="371"/>
            <end position="394"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="422"/>
            <end position="447"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="453"/>
            <end position="476"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="1"/>
            <end position="29"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="17"/>
            <end position="29"/>
        </location>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="53"/>
        </location>
        <ligand>
            <name>Na(+)</name>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="55"/>
        </location>
        <ligand>
            <name>Na(+)</name>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="56"/>
        </location>
        <ligand>
            <name>Na(+)</name>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="60"/>
        </location>
        <ligand>
            <name>Na(+)</name>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="294"/>
        </location>
        <ligand>
            <name>Na(+)</name>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="326"/>
        </location>
        <ligand>
            <name>Na(+)</name>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="392"/>
        </location>
        <ligand>
            <name>Na(+)</name>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="395"/>
        </location>
        <ligand>
            <name>Na(+)</name>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="binding site" evidence="6">
        <location>
            <position position="396"/>
        </location>
        <ligand>
            <name>Na(+)</name>
            <dbReference type="ChEBI" id="CHEBI:29101"/>
            <label>1</label>
        </ligand>
    </feature>
    <feature type="disulfide bond" evidence="7">
        <location>
            <begin position="159"/>
            <end position="168"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004141"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PIRSR" id="PIRSR600175-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PIRSR" id="PIRSR600175-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="RuleBase" id="RU003732"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000008053.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="531" mass="59620" checksum="F62FB3AB07F03D9C" modified="2019-05-08" version="1">MSEQNSPVQVDSLPRQAIQTNPSHTANPVLLTTQTDSRDKWSKKVDFLLSVIGFAVDLGNVWRFPYICYQNGGGAFLIPYTLMAIFGGVPLFYMELALGQFHRTGAISIWKRICPIFKGIGFAICIIALYVSFYYNTIIAWALYYFYSSFTGTLPWTSCDNPWNTPNCTNYFGKSNVTWTTFSKSPAEEFYMRKVLEIQKSSGLYDVGGIRWQLLLCLFLLFTIVYFSLWKGVKTSGKVVWVTATLPYIVLLILLVRGATLPGAWRGIVFYLRPDWEKLMSTVWVDAAAQIFFSLGPGFGVLLALASYNHFNNNCYRDAIVTSVVNCLTSFLSGFVIFTVLGYMAEMRDVEVEDVAKDKGPSLLFITYPEAIANMVGSTFFAIIFFLMMITLGLDSTFGGLEAVITAVMDEYPHILAKQRELFVLGLITICFLACLCLSFQGGAYVVKLLEEFGAGCSILAVVLLEAIAVSWFYGIQRFCDDVKAMLGFTPGTFWKVCWVAISPAFLAVRLPHPRHNYIRGRPDRREACQH</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H114</accession>
    <name>A0A452H114_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Protein inturned</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="10">Inturned planar cell polarity effector homolog</fullName>
        </alternativeName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell surface</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Cilium basal body</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the inturned family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H114"/>
    <dbReference type="Ensembl" id="ENSGAGT00000009373.1">
        <property type="protein sequence ID" value="ENSGAGP00000008139.1"/>
        <property type="gene ID" value="ENSGAGG00000006468.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009986">
        <property type="term" value="C:cell surface"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005929">
        <property type="term" value="C:cilium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060271">
        <property type="term" value="P:cilium assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001736">
        <property type="term" value="P:establishment of planar polarity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016192">
        <property type="term" value="P:vesicle-mediated transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000327">
        <property type="entry name" value="Protein inturned"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.42.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043987">
        <property type="entry name" value="CCZ1/INTU/HSP4_longin_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043989">
        <property type="entry name" value="CCZ1/INTU/HSP4_longin_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043988">
        <property type="entry name" value="CCZ1/INTU_longin_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039151">
        <property type="entry name" value="INTU"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001478">
        <property type="entry name" value="PDZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036034">
        <property type="entry name" value="PDZ_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21082">
        <property type="entry name" value="PROTEIN INTURNED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21082:SF4">
        <property type="entry name" value="PROTEIN INTURNED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19031">
        <property type="entry name" value="Intu_longin_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19032">
        <property type="entry name" value="Intu_longin_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19033">
        <property type="entry name" value="Intu_longin_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50156">
        <property type="entry name" value="PDZ domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50106">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0966">Cell projection</keyword>
    <keyword evidence="7" id="KW-0970">Cilium biogenesis/degradation</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="5" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="PDZ" evidence="12">
        <location>
            <begin position="210"/>
            <end position="271"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="116"/>
            <end position="146"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="688"/>
            <end position="737"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="136"/>
            <end position="146"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="688"/>
            <end position="710"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="720"/>
            <end position="729"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004120"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004241"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00010034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00015639"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022794"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00032633"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50106"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000008139.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="930" mass="104383" checksum="0EA6E98A83A05B4B" modified="2019-05-08" version="1">MKWPLLCSMFSVGRYSIPRPVTRNLEDLDSVQSILLHSDLEPEWLDSVQKNGELFYLELSEGEEETLLQDSSSEVPAVNHVRFRENEAEVIQEGSRKERKYEPRLKKITKILKKKSLLPKHSSKKGTGGSDRHPSGPTSILKHQSTQKVGVIVQQQYKDVCVYVNPKRLPNDGAGEKLKLLEALIGIVHQSSWSNRRAEKQGEQNRSNKGINEEKLVVHGLVPGSSAIKTGQILIGDALVAVNDVDVNAENIERVLSCIPGPMQVKLTFETSVFEHEGTSQQRYKQAQSAMNNLVKLLWGEDNMDLQLAIQGMPHIVMYLTLKLDSETSKDEQEILYQYPISEASQKLKSVRGIFLTLCDMLENVTGAHIISSSLFLCGKLVQVVYWKESDKLLVIGLPAESVPLSQLRNMITDVVRTLKFMYGSLDSAFCQVENISRLDHFFNLFFQRAIQPARLNSNASPSAQHYDSCSALFLDNLPGIRWLTLAQEIKVEIDTALSDLEAADFAELSEDYYDMRRLYMILGSCLFYKGCLIGNHLPKEDLIDIALYSRHYCLLPLAAEQRIGQLVIWREVFPHHHLQPSEASGVTGYREPEARYFLLIVGLRHFMLCVLLETGGCTSKAIGNPGPDCIYVDQVKATILQLEGIDSSIEERLESPAIPCLSCADWFLPASRDKLESFTTPPILSKLQNTSKTATSPTSKRTLFTDSSLRTRKPSPTRSSGGSDNGSEGPEEGGGLNLLATVDQVTRKKHTLPNPFHLGNLKKNLSEKDTELYNVVKLTSGPENTLFHYISLETMQGIFITPTHKEVQQLSGSIHPQLIKNFHRCCLSIRSIFQQAVMKEKKKNADSGSNSDSRKASSDLNLVREHGVLFECYPDNWTDQKKPPPTMTYWVVGRLFLHPKPQEFYVCFHDSVTEVAVELAFKLSFGLAT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H115</accession>
    <name>A0A452H115_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Protein inturned</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="10">Inturned planar cell polarity effector homolog</fullName>
        </alternativeName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell surface</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Cilium basal body</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the inturned family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H115"/>
    <dbReference type="Ensembl" id="ENSGAGT00000009380.1">
        <property type="protein sequence ID" value="ENSGAGP00000008146.1"/>
        <property type="gene ID" value="ENSGAGG00000006468.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009986">
        <property type="term" value="C:cell surface"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005929">
        <property type="term" value="C:cilium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060271">
        <property type="term" value="P:cilium assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001736">
        <property type="term" value="P:establishment of planar polarity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016192">
        <property type="term" value="P:vesicle-mediated transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000327">
        <property type="entry name" value="Protein inturned"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.42.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043987">
        <property type="entry name" value="CCZ1/INTU/HSP4_longin_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043989">
        <property type="entry name" value="CCZ1/INTU/HSP4_longin_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043988">
        <property type="entry name" value="CCZ1/INTU_longin_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039151">
        <property type="entry name" value="INTU"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001478">
        <property type="entry name" value="PDZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036034">
        <property type="entry name" value="PDZ_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21082">
        <property type="entry name" value="PROTEIN INTURNED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21082:SF4">
        <property type="entry name" value="PROTEIN INTURNED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19031">
        <property type="entry name" value="Intu_longin_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19032">
        <property type="entry name" value="Intu_longin_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19033">
        <property type="entry name" value="Intu_longin_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50156">
        <property type="entry name" value="PDZ domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50106">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0966">Cell projection</keyword>
    <keyword evidence="7" id="KW-0970">Cilium biogenesis/degradation</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="5" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="PDZ" evidence="12">
        <location>
            <begin position="214"/>
            <end position="275"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1"/>
            <end position="49"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="120"/>
            <end position="150"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="670"/>
            <end position="720"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="9"/>
            <end position="19"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="140"/>
            <end position="150"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="670"/>
            <end position="693"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="703"/>
            <end position="712"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004120"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004241"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00010034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00015639"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022794"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00032633"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50106"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000008146.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="913" mass="101690" checksum="36EA56E342CE4D84" modified="2019-05-08" version="1">MEEPLSLEMPREPAPRQEEEAAGGDESSETGSCSASFSGGSDDLEPEWLDSVQKNGELFYLELSEGEEETLLQDSSSEVPAVNHVRFRENEAEVIQEGSRKERKYEPRLKKITKILKKKSLLPKHSSKKGTGGSDRHPSGPTSILKHQSTQKVGVIVQQQYKDVCVYVNPKRLPNDGAGEKLKLLEALIGIVHQSSWSNRRAEKQGEQNRSNKGINEEKLVVHGLVPGSSAIKTGQILIGDALVAVNDVDVNAENIERVLSCIPGPMQVKLTFETSVFEHEGTSQQRYKQAQSAMNNLVKLLWGEDNMDLQLAIQGMPHIVMYLTLKLDSETSKDEQEILYQYPISEASQKLKSVRGIFLTLCDMLENVTGAHIISSSLFLCGKLVQVVYWKESDKLLVIGLPAESVPLSQLRNMITDVVRTLKFMYGSLDSAFCQVENISRLDHFFNLFFQRAIQPARLNSNASPSAQHYDSCSALFLDNLPGIRWLTLAQEIKVEIDTALSDLEAADFAELGCLIGNHLPKEDLIDIALYSRHYCLLPLAAEQRIGQLVIWREVFPHHHLQPSEASGVTGYREPEARYFLLIVGLRHFMLCVLLETGGCTSKAIGNPGPDCIYVDQVKATILQLEGIDSSIEERLESPAIPCLSCADWFLPASRDKLESFTTPPILSKLQNTSKTATSPTSKRTLFTDSSLRTRKPSPTRSSGGSDNGSEGPEEGGGLNLLATVDQVTRKKHTLPNPFHLGNLKKNLSEKDTELYNVVKLTSGPENTLFHYISLETMQGIFITPTHKEVQQLSGSIHPQLIKNFHRCCLSIRSIFQQAVMKEKKKNADSGSNSDSRKASSDLNLVREHGVLFECYPDNWTDQKKPPPTMTYWVVGRLFLHPKPQEFYVCFHDSVTEVAVELAFKLSFGLAT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H116</accession>
    <name>A0A452H116_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">Ankyrin repeat domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="6">Ubiquitin-binding protein that specifically recognizes and binds 'Lys-63'-linked ubiquitin. Does not bind 'Lys-48'-linked ubiquitin. Positively regulates the internalization of ligand-activated EGFR by binding to the Ub moiety of ubiquitinated EGFR at the cell membrane.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Late endosome</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H116"/>
    <dbReference type="Ensembl" id="ENSGAGT00000009285.1">
        <property type="protein sequence ID" value="ENSGAGP00000008059.1"/>
        <property type="gene ID" value="ENSGAGG00000006410.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005770">
        <property type="term" value="C:late endosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021832">
        <property type="entry name" value="ANKRD13"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR055285">
        <property type="entry name" value="ANKRD13_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003903">
        <property type="entry name" value="UIM_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12447">
        <property type="entry name" value="ANKYRIN REPEAT DOMAIN-CONTAINING PROTEIN 13"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12447:SF4">
        <property type="entry name" value="ANKYRIN REPEAT DOMAIN-CONTAINING PROTEIN 13A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11904">
        <property type="entry name" value="ANKRD13_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00726">
        <property type="entry name" value="UIM"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50330">
        <property type="entry name" value="UIM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="5" id="KW-0472">Membrane</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <feature type="repeat" description="ANK" evidence="7">
        <location>
            <begin position="25"/>
            <end position="57"/>
        </location>
    </feature>
    <feature type="domain" description="Ankyrin repeat" evidence="9">
        <location>
            <begin position="141"/>
            <end position="456"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="521"/>
            <end position="577"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="521"/>
            <end position="534"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="8">
        <location>
            <begin position="547"/>
            <end position="560"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="8">
        <location>
            <begin position="567"/>
            <end position="577"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004236"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004603"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00024956"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF11904"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000008059.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="577" mass="66342" checksum="E3257DDC2F55CCF4" modified="2019-05-08" version="1">MLFCFKSLKLVVKNICKDVDQRDPRGRTLLHLAVSLGYIESAEVLLRHKADVTKENAQGWTVLHEAVSTGDPEMAYLILQHRDYHQTSATLGGVPELLQKINETSDFYVEMKWEFTSWVPLVSRVCPNDTCRIWKSGAKLRVDITLLGFENMSWERGKRSLIFKGEDTGGWAELIEINHDDKVVTTERFEISQQMKRLTLGSVMPERKEVERRLTSPIINTCLDTKNIAFERNTSGFWVWRTEKAEGVNGYEAKVYTANNVNVVTKIRTEHLTEEEKKRYKADRNPLESFLGTVEHEYGVQDLKTEYATMNNPTAITLEEYFDPKFDLKDRDIGRPKEVTIRTQKFKATLWMCEEFPLSLMEQITPIIDLMARASAHFARLKDFITLEFPPGFPVKIEIPLFHVLNARITFENVNGCRTAEETSSQTAEGAECELASTASNFEVNQSVFEIPKSYHTQHDGRNVRVQDEDNEIMQFAIQQSLLESSGNKEALGMHSNGAVAYAQDFNIQYQRALQESYIMSSSTPHVSTPNESSSFDKDLQLAMELSVREQQEQEKRRREEEEEEFQQALQRSLTEK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H117</accession>
    <name>A0A452H117_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">Mediator of RNA polymerase II transcription subunit 24</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="8">Mediator complex subunit 24</fullName>
        </alternativeName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the Mediator complex subunit 24 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H117"/>
    <dbReference type="Ensembl" id="ENSGAGT00000009538.1">
        <property type="protein sequence ID" value="ENSGAGP00000008282.1"/>
        <property type="gene ID" value="ENSGAGG00000006579.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016592">
        <property type="term" value="C:mediator complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003712">
        <property type="term" value="F:transcription coregulator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060261">
        <property type="term" value="P:positive regulation of transcription initiation by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021429">
        <property type="entry name" value="Mediator_Med24"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12898">
        <property type="entry name" value="MEDIATOR OF RNA POLYMERASE II TRANSCRIPTION SUBUNIT 24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12898:SF1">
        <property type="entry name" value="MEDIATOR OF RNA POLYMERASE II TRANSCRIPTION SUBUNIT 24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11277">
        <property type="entry name" value="Med24_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0010">Activator</keyword>
    <keyword evidence="7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0804">Transcription</keyword>
    <keyword evidence="4" id="KW-0805">Transcription regulation</keyword>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007864"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00019693"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00031960"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000008282.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="982" mass="110111" checksum="27139038EEF22AC4" modified="2019-05-08" version="1">MKVVNLKQAILQAWKERWSDYQWAINMKKFFPKGATWDILNLAEALLEQAMIGPSPNPLILSYLKYAISSQMVSYSTVLMAISKFDDFSRDLCVQSLLEIMDMFCDRLSCHGKAEECIGLCRALMSALNWMLRCAAFYTEKFKDVLDQVAAENQLKMCLERLEKILGSTKNRALIHIGKLEETSSWSAIEQSLIKLGENLSVLSNPQLRSQADECISLIKSIPTMLSAHSEQLNKTGFPTVHAVVLLEGTMNLTGETQPLVEQLMMVKRMQRIPSPLFVLEIWKACFVGLIESPEGTEELKWTAFTFLKIPQVLVKLKKYPQGEKDFTEDVNCAFEFLLKLTPLLDKADQRCNCDCTSLLLQECSKQGLLSEAHMTNLVAKRTADREHAPRLKSAENANIQPNPGLILRAEPTVTNILKTMDADHSKSPEGLLGVLGHMLSGKSLDLLLAAAAATGKLKSFARKFIKLNEFTKHISGEGSKAASVRALLFDISFLMLCHVAQTYGSEVILSESSPTGEVPFFETWMQTCMPEEGKILNPDHPCFRPDSTKVESLVALLNSSSEMKLVQMKWHEACLSISAAILEILNAWENGVLTFESIQKITDNIKGKVCSMAVCTMAWLVAHVRMLGLDERDKSLQMIRQLATSLQSENTLQFYNERVVIMSSILERMCADVLQQTATQIRFPSTGTDPIPYWNLLPPKKPIKEVLMGVFTKVLEKGWVDSRSIHIFDTLLHMGGVYWFCNNLVKELLKETRKEHTLRAVELLYAIFCLDMQQLTLSLLGHILPSLLTDSSKWHTLMDPPGKALAKLSVWCALSSYSSHNKGQASARQKKRHREDIEDYISLFPLDDTQPSKLMRLLSSNEEDPNVLSSPTDRSMSSSLSASQLHTVSMRDPLNRVLANLFLLISSILGAKTAGTHTQFVQWFMEECVDCLEQGSRGSILQFMPFTMVLLQELLEILIPGELRQGSCVQHGASQTQHGAD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H119</accession>
    <name>A0A452H119_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Protein inturned</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="10">Inturned planar cell polarity effector homolog</fullName>
        </alternativeName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell surface</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
            <location evidence="1">Cilium basal body</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3">Belongs to the inturned family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H119"/>
    <dbReference type="Ensembl" id="ENSGAGT00000009378.1">
        <property type="protein sequence ID" value="ENSGAGP00000008144.1"/>
        <property type="gene ID" value="ENSGAGG00000006468.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009986">
        <property type="term" value="C:cell surface"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005929">
        <property type="term" value="C:cilium"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060271">
        <property type="term" value="P:cilium assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001736">
        <property type="term" value="P:establishment of planar polarity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016192">
        <property type="term" value="P:vesicle-mediated transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.42.10:FF:000327">
        <property type="entry name" value="Protein inturned"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.42.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043987">
        <property type="entry name" value="CCZ1/INTU/HSP4_longin_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043989">
        <property type="entry name" value="CCZ1/INTU/HSP4_longin_3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043988">
        <property type="entry name" value="CCZ1/INTU_longin_2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039151">
        <property type="entry name" value="INTU"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001478">
        <property type="entry name" value="PDZ"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036034">
        <property type="entry name" value="PDZ_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21082">
        <property type="entry name" value="PROTEIN INTURNED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR21082:SF4">
        <property type="entry name" value="PROTEIN INTURNED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19031">
        <property type="entry name" value="Intu_longin_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19032">
        <property type="entry name" value="Intu_longin_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19033">
        <property type="entry name" value="Intu_longin_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50156">
        <property type="entry name" value="PDZ domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50106">
        <property type="entry name" value="PDZ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="9" id="KW-0966">Cell projection</keyword>
    <keyword evidence="7" id="KW-0970">Cilium biogenesis/degradation</keyword>
    <keyword evidence="6" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="5" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="PDZ" evidence="12">
        <location>
            <begin position="210"/>
            <end position="271"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="116"/>
            <end position="146"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="662"/>
            <end position="711"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="136"/>
            <end position="146"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="662"/>
            <end position="684"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="11">
        <location>
            <begin position="694"/>
            <end position="703"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004120"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004241"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00010034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00015639"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022794"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00032633"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50106"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000008144.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="904" mass="101417" checksum="69E426AA2979D4A0" modified="2019-05-08" version="1">MKWPLLCSMFSVGRYSIPRPVTRNLEDLDSVQSILLHSDLEPEWLDSVQKNGELFYLELSEGEEETLLQDSSSEVPAVNHVRFRENEAEVIQEGSRKERKYEPRLKKITKILKKKSLLPKHSSKKGTGGSDRHPSGPTSILKHQSTQKVGVIVQQQYKDVCVYVNPKRLPNDGAGEKLKLLEALIGIVHQSSWSNRRAEKQGEQNRSNKGINEEKLVVHGLVPGSSAIKTGQILIGDALVAVNDVDVNAENIERVLSCIPGPMQVKLTFETSVFEHEGTSQQRYKQAQSAMNNLVKLLWGEDNMDLQLAIQGMPHIVMYLTLKLDSETSKDEQEILYQYPISEASQKLKSVRGIFLTLCDMLENVTGAHIISSSLFLCGKLVQVVYWKESDKLLVIGLPAESAFCQVENISRLDHFFNLFFQRAIQPARLNSNASPSAQHYDSCSALFLDNLPGIRWLTLAQEIKVEIDTALSDLEAADFAELSEDYYDMRRLYMILGSCLFYKGCLIGNHLPKEDLIDIALYSRHYCLLPLAAEQRIGQLVIWREVFPHHHLQPSEASGVTGYREPEARYFLLIVGLRHFMLCVLLETGGCTSKAIGNPGPDCIYVDQVKATILQLEGIDSSIEERLESPAIPCLSCADWFLPASRDKLESFTTPPILSKLQNTSKTATSPTSKRTLFTDSSLRTRKPSPTRSSGGSDNGSEGPEEGGGLNLLATVDQVTRKKHTLPNPFHLGNLKKNLSEKDTELYNVVKLTSGPENTLFHYISLETMQGIFITPTHKEVQQLSGSIHPQLIKNFHRCCLSIRSIFQQAVMKEKKKNADSGSNSDSRKASSDLNLVREHGVLFECYPDNWTDQKKPPPTMTYWVVGRLFLHPKPQEFYVCFHDSVTEVAVELAFKLSFGLAT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H121</accession>
    <name>A0A452H121_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="13">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Lipid-anchor</topology>
            <topology evidence="1">GPI-anchor</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452H121"/>
    <dbReference type="Ensembl" id="ENSGAGT00000009290.1">
        <property type="protein sequence ID" value="ENSGAGP00000008064.1"/>
        <property type="gene ID" value="ENSGAGG00000005614.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030424">
        <property type="term" value="C:axon"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043025">
        <property type="term" value="C:neuronal cell body"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098552">
        <property type="term" value="C:side of membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008046">
        <property type="term" value="F:axon guidance receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007156">
        <property type="term" value="P:homophilic cell adhesion via plasma membrane adhesion molecules"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050808">
        <property type="term" value="P:synapse organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd00096">
        <property type="entry name" value="Ig"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="CDD" id="cd06263">
        <property type="entry name" value="MAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000240">
        <property type="entry name" value="MAM domain containing glycosylphosphatidylinositol anchor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000262">
        <property type="entry name" value="MAM domain containing glycosylphosphatidylinositol anchor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000303">
        <property type="entry name" value="MAM domain containing glycosylphosphatidylinositol anchor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.200:FF:000019">
        <property type="entry name" value="MAM domain containing glycosylphosphatidylinositol anchor 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000165">
        <property type="entry name" value="MAM domain containing glycosylphosphatidylinositol anchor 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000243">
        <property type="entry name" value="MAM domain-containing glycosylphosphatidylinositol anchor protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.200">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050958">
        <property type="entry name" value="Cell_Adh-Cytoskel_Orgn"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013098">
        <property type="entry name" value="Ig_I-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000998">
        <property type="entry name" value="MAM_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45080">
        <property type="entry name" value="CONTACTIN 5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45080:SF35">
        <property type="entry name" value="MAM DOMAIN-CONTAINING GLYCOSYLPHOSPHATIDYLINOSITOL ANCHOR 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07679">
        <property type="entry name" value="I-set"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13927">
        <property type="entry name" value="Ig_3"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00629">
        <property type="entry name" value="MAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SMART" id="SM00137">
        <property type="entry name" value="MAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50060">
        <property type="entry name" value="MAM_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="7" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="8" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="3" id="KW-0336">GPI-anchor</keyword>
    <keyword evidence="10" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="9" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="6" id="KW-0472">Membrane</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <feature type="domain" description="Ig-like" evidence="12">
        <location>
            <begin position="19"/>
            <end position="119"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="12">
        <location>
            <begin position="126"/>
            <end position="224"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="12">
        <location>
            <begin position="234"/>
            <end position="320"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="12">
        <location>
            <begin position="332"/>
            <end position="428"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="12">
        <location>
            <begin position="434"/>
            <end position="525"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="12">
        <location>
            <begin position="532"/>
            <end position="619"/>
        </location>
    </feature>
    <feature type="domain" description="MAM" evidence="11">
        <location>
            <begin position="730"/>
            <end position="905"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004609"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022622"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50060"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000008064.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="913" mass="102693" checksum="7F134B3D89E36C3F" modified="2019-05-08" version="1">YKGSLVLFLSGKRTFVPPPTVRIVHSGLACNIEEERYSERVYTIREGETLELTCLVTGHPRPQIRWTKTAGSASDRFQDSSVFNETLRITNIQRHQGGRYYCKAENGLGSPAIKSIRVDVYYLDDPIVTVHQSIGEAKEQFYYERTVFLRCVANSNPPVRYSWRRGQEVLLQGSDKGVEIYEPFFTQGETKILKLKNLRPQDYANYSCIASVRSVCNIPDKMVSFRLSNKTASPSIKLLVDDPIVVNPGEAITLVCVTTGGEPAPTLTWVRSIGALPEKTVLNGGTLTIPAITLDDTGTYSCIANNNVGNPAKKSTNIIVRALKKGRFWITPDPYHKDDNIQIGREVKISCQVEAVPSEELTFSWFKNGRPLRSSERMVITQTDPDVSPGTTNLDIIDLKFTDFGTYTCVASLKGGGISDISIDVNISSSTVPPNLTVPQEKSPLVTREGDIIELQCQVTGKPKPIILWSRADKEIAMPDGSMQMESYDGILRIVNVSREMTGTYRCQTSQYNGFNVKPREALVQLIVQYPPAVEPTFLEIRQGQGRSITMSCRVLRAYPTRVLTYEWRLGNKLLRTGQFDAHDSTEYIVSSLSRDSYGIYNCNIINEAGAGRCSFLVTGKAYAPEFYYDTYSPLWQNRPRVYSYSLQWTQMNPDAVDRILAYRLGIRQAGQQRWWEQEITVNGNIQKGELITYNLTELIKPEAYEVRLTPITRFGEGDSTIRVIKYREFHCGFEDGNICLFTQDDTDNFDWTKQSTATRDTKYTPNTGPNADRSGSKEGFYMYIETSRPRLEGEKARLLSPVFSVAPKNPYGVTNTAYCFSFYYHMYGQHIGSLNVYLRLKGQTTIENPVWSSSGNKGQHWNQARVNIHPTTSFQLIFEGIRGPGIEGDIAIDDVSIVEGECMKADQPANSK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H125</accession>
    <name>A0A452H125_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Acyl-CoA synthetase short-chain family member 3, mitochondrial</fullName>
            <ecNumber evidence="4">6.2.1.1</ecNumber>
            <ecNumber evidence="3">6.2.1.17</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="8">Acetate--CoA ligase 3</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="6">Propionate--CoA ligase</fullName>
        </alternativeName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>acetate + ATP + CoA = acetyl-CoA + AMP + diphosphate</text>
            <dbReference type="Rhea" id="RHEA:23176"/>
            <dbReference type="ChEBI" id="CHEBI:30089"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:33019"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57288"/>
            <dbReference type="ChEBI" id="CHEBI:456215"/>
            <dbReference type="EC" id="6.2.1.1"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="1">
            <dbReference type="Rhea" id="RHEA:23177"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="9">
            <text>butanoate + ATP + CoA = butanoyl-CoA + AMP + diphosphate</text>
            <dbReference type="Rhea" id="RHEA:46172"/>
            <dbReference type="ChEBI" id="CHEBI:17968"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:33019"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57371"/>
            <dbReference type="ChEBI" id="CHEBI:456215"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="9">
            <dbReference type="Rhea" id="RHEA:46173"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="10">
            <text>propanoate + ATP + CoA = propanoyl-CoA + AMP + diphosphate</text>
            <dbReference type="Rhea" id="RHEA:20373"/>
            <dbReference type="ChEBI" id="CHEBI:17272"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:33019"/>
            <dbReference type="ChEBI" id="CHEBI:57287"/>
            <dbReference type="ChEBI" id="CHEBI:57392"/>
            <dbReference type="ChEBI" id="CHEBI:456215"/>
            <dbReference type="EC" id="6.2.1.17"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="10">
            <dbReference type="Rhea" id="RHEA:20374"/>
        </physiologicalReaction>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the ATP-dependent AMP-binding enzyme family.</text>
    </comment>
    <dbReference type="EC" id="6.2.1.1" evidence="4"/>
    <dbReference type="EC" id="6.2.1.17" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452H125"/>
    <dbReference type="Ensembl" id="ENSGAGT00000009383.1">
        <property type="protein sequence ID" value="ENSGAGP00000008149.1"/>
        <property type="gene ID" value="ENSGAGG00000006436.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005759">
        <property type="term" value="C:mitochondrial matrix"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003987">
        <property type="term" value="F:acetate-CoA ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0050218">
        <property type="term" value="F:propionate-CoA ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006629">
        <property type="term" value="P:lipid metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd05967">
        <property type="entry name" value="PrpE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.12780:FF:000011">
        <property type="entry name" value="Acetyl-coenzyme A synthetase 2-like, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.300.30:FF:000017">
        <property type="entry name" value="Acyl-CoA synthetase short-chain family member 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.300.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.12780">
        <property type="entry name" value="N-terminal domain of ligase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR025110">
        <property type="entry name" value="AMP-bd_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045851">
        <property type="entry name" value="AMP-bd_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020845">
        <property type="entry name" value="AMP-binding_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000873">
        <property type="entry name" value="AMP-dep_synth/lig_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042099">
        <property type="entry name" value="ANL_N_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43347">
        <property type="entry name" value="ACYL-COA SYNTHETASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43347:SF3">
        <property type="entry name" value="ACYL-COA SYNTHETASE SHORT-CHAIN FAMILY MEMBER 3, MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00501">
        <property type="entry name" value="AMP-binding"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF13193">
        <property type="entry name" value="AMP-binding_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56801">
        <property type="entry name" value="Acetyl-CoA synthetase-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00455">
        <property type="entry name" value="AMP_BINDING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0443">Lipid metabolism</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="11" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="11">
        <location>
            <begin position="1"/>
            <end position="24"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019047817" description="Acyl-CoA synthetase short-chain family member 3, mitochondrial" evidence="11">
        <location>
            <begin position="25"/>
            <end position="612"/>
        </location>
    </feature>
    <feature type="domain" description="AMP-dependent synthetase/ligase" evidence="12">
        <location>
            <begin position="48"/>
            <end position="432"/>
        </location>
    </feature>
    <feature type="domain" description="AMP-binding enzyme C-terminal" evidence="13">
        <location>
            <begin position="496"/>
            <end position="574"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006432"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012985"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00013275"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023098"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00029726"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00040004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00042755"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00047935"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00049004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF00501"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="Pfam" id="PF13193"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000008149.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="612" mass="67249" checksum="D95B43C474524B4E" modified="2019-05-08" version="1">AARGICRVTLWCSIHAFVTHYALVQESVADGSFELNICYNAIDRHVESGHADKIAIIYDSPVTNTKQAITYKEVLEQVSKLADVMVKYGVKKGDCVVIYMPMIPQTIYTMLACARIGAIHSLIFGGFASKELSSRIDHAKPKLVVTATFGIEPGRRVEYMPLLERALEMIQHQPEKVLIYSRPHLGTVPLVSGRDLDWDEEIAKAKPHDCVAVRSDHPLYILYTSGTTGVPKGVVRPTGGYAVMLNWTMSGIYGLRPGEVWWAASDLGWVVGHSYVCYAPLLHGNTTVLYEGKPVGTPDAGAYFRVLAEHGVAAFFTAPTAIRAIRQQDPEADLGKQYCLTRFRTLFLAGERCDVETLEWSKKVFKVPVLDHWWQTETGSAITASCIGLGNSTTPPPGQAGKSVPGYNVMVLDDNKQPVKAKTLGNIVVKLPLPPGAFSGLWKNQETFKELYFQKFPGYYDTMDAGCMDEDGYLYVMSRIDDVINVAGHRISAGAIEESVLSHGAVADCAVVGQEDALKGLVPLALCVLRNGIKIEEEKVLEEIVKRVRESIGPVAAFRKAVFVKQIPKTRSGKIPRSALSALVSGKPYQITPTIEDPGVFKHVEEVLKKQK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452H129</accession>
    <name>A0A452H129_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Arf-GAP domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452H129"/>
    <dbReference type="Ensembl" id="ENSGAGT00000009300.1">
        <property type="protein sequence ID" value="ENSGAGP00000008074.1"/>
        <property type="gene ID" value="ENSGAGG00000006421.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0098793">
        <property type="term" value="C:presynapse"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="GOC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031267">
        <property type="term" value="F:small GTPase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007420">
        <property type="term" value="P:brain development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032012">
        <property type="term" value="P:regulation of ARF protein signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008277">
        <property type="term" value="P:regulation of G protein-coupled receptor signaling pathway"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0036465">
        <property type="term" value="P:synaptic vesicle recycling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd08847">
        <property type="entry name" value="ArfGap_GIT2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000013">
        <property type="entry name" value="ARF GTPase-activating protein GIT1 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.220.150:FF:000003">
        <property type="entry name" value="ARF GTPase-activating protein GIT2 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.170:FF:000015">
        <property type="entry name" value="ARF GTPase-activating protein GIT2 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.170">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.220.150">
        <property type="entry name" value="Arf GTPase activating protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.330">
        <property type="entry name" value="Nucleotidyltransferases domain 2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR037278">
        <property type="entry name" value="ARFGAP/RecO"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001164">
        <property type="entry name" value="ArfGAP_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038508">
        <property type="entry name" value="ArfGAP_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047161">
        <property type="entry name" value="GIT-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032352">
        <property type="entry name" value="GIT1/2_CC"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022018">
        <property type="entry name" value="GIT1_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013724">
        <property type="entry name" value="GIT_SHD"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46097:SF4">
        <property type="entry name" value="ARF GTPASE-ACTIVATING PROTEIN GIT2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46097">
        <property type="entry name" value="G PROTEIN-COUPLED RECEPTOR KINASE INTERACTING ARFGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01412">
        <property type="entry name" value="ArfGap"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12205">
        <property type="entry name" value="GIT1_C"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16559">
        <property type="entry name" value="GIT_CC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF08518">
        <property type="entry name" value="GIT_SHD"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00405">
        <property type="entry name" value="REVINTRACTNG"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00105">
        <property type="entry name" value="ArfGap"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00555">
        <property type="entry name" value="GIT"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57863">
        <property type="entry name" value="ArfGap/RecO-like zinc finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50115">
        <property type="entry name" value="ARFGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6 8" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="7 10" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="1" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="2" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <keyword evidence="4 9" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="Arf-GAP" evidence="12">
        <location>
            <begin position="1"/>
            <end position="124"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="8">
        <location>
            <begin position="166"/>
            <end position="198"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="379"/>
            <end position="398"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="486"/>
            <end position="529"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="541"/>
            <end position="614"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="10">
        <location>
            <begin position="444"/>
            <end position="478"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="11">
        <location>
            <begin position="385"/>
            <end position="398"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="486"/>
            <end position="499"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="554"/>
            <end position="567"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="11">
        <location>
            <begin position="594"/>
            <end position="609"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00288"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50115"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000008074.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="771" mass="86384" checksum="1BF1E62C31307A7B" modified="2019-05-08" version="1">MSKRLRSSEVCADCSAQDPCWASVNRGILICDECCSVHRSLGRHISQVRHLKHTPWPPTLLQMVETLYNNGANSIWEHSLLDPASVMSGRRKANPQDKVHPNKAEFIRAKYQMLAFVHRLPCREDDSVTAKDLSKQLHSSVRTGNLETCLRLLSLGAQANFFHPEKGNTPLHVAAKAGQILQAELLAVYGADPGTQDSNGKTPVDYARQGGHHELAERLIEIQYELTDRLAFYLCGRKPDHKNGQHFIIPQMADSSLDLSELAKAAKKKLQSLSNHLFEELAMDVYDEVDRRETDAVWLATQNHSTLVTETTLVPFLPVNPEYSSTRNQGRQKLARFNAHEFATLVIDILSDAKRRQQGNPVTSCKENVELILKAINSQHSSESQDNDQPDYDSVASDEDTDLDANAAKANRQKSLDSDVSDGPVTAQEYMQVKNALVASEVKIQQLMKVNINLSDELRTMQKKLQTLQSENTNLRRQATTNIYQVQTGSEYTDPNNHSSLKRRPSARGSRPMSMYETGSGQKPYLPMGEVTYPEESITRLQPFPPHASKLEKQNSMPESDYDNPSPSLELDETGPGRKGRQRSIIWQGEGSIPENSDTDATPSSTLPSTEDVIRKTEQITKNIQELLRAAQENKHDRPLERASMLKLRHSLSCFSTLVPWAERAPLQPLTHQQPDPTSCYIPCSERIHVAVTEMAALFPKKPKSELVRTSLRLLTSSAYRLQSECKKTLPIETCPTTDIQLVTQQVIQCAYDIAKAAKQLVTITTKENNN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HAR4</accession>
    <name>A0A452HAR4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Death domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HAR4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013529.1">
        <property type="protein sequence ID" value="ENSGAGP00000011814.1"/>
        <property type="gene ID" value="ENSGAGG00000009100.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd01670">
        <property type="entry name" value="Death"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.533.10">
        <property type="entry name" value="Death Domain, Fas"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052457">
        <property type="entry name" value="Ankyrin-DD_containing_protein"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011029">
        <property type="entry name" value="DEATH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000488">
        <property type="entry name" value="Death_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24125">
        <property type="entry name" value="ANKYRIN REPEAT AND DEATH DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24125:SF0">
        <property type="entry name" value="ANKYRIN REPEAT AND DEATH DOMAIN-CONTAINING PROTEIN 1A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00023">
        <property type="entry name" value="Ank"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="3"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01415">
        <property type="entry name" value="ANKYRIN"/>
    </dbReference>
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        <property type="match status" value="10"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
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        <property type="match status" value="6"/>
    </dbReference>
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    </dbReference>
    <dbReference type="PROSITE" id="PS50017">
        <property type="entry name" value="DEATH_DOMAIN"/>
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    </dbReference>
    <proteinExistence type="predicted"/>
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        <location>
            <begin position="47"/>
            <end position="79"/>
        </location>
    </feature>
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            <begin position="80"/>
            <end position="112"/>
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        <location>
            <begin position="113"/>
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        </location>
    </feature>
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            <begin position="148"/>
            <end position="180"/>
        </location>
    </feature>
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            <begin position="181"/>
            <end position="213"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="1">
        <location>
            <begin position="214"/>
            <end position="246"/>
        </location>
    </feature>
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            <begin position="247"/>
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        </location>
    </feature>
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            <begin position="280"/>
            <end position="312"/>
        </location>
    </feature>
    <feature type="domain" description="Death" evidence="2">
        <location>
            <begin position="388"/>
            <end position="462"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="PROSITE" id="PS50017"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011814.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="475" mass="53598" checksum="E2F20150587F16D1" modified="2019-05-08" version="1">MEDGLVSEEEILLHSEKEFHEAAKRNDIARMQELIRTRVNVKAKNNTDRTALHWAAGAGNEQAVRLLLDHEVPVDDKDSFGMNALLLSAWFGHLRVLQILVNAGAKINCVDKNGRNLLHCAAQRGHIKVMEFIMEDLEDVHVDKTDKEKNTALHLAAKNGHPTVLQKIVEIGQDLEEKNMEGLTALHLAVEGGHYDCVKVLLEAGCDVNAQTQKAMTCLHYAALHGYEDMARIFIDAGIYIDAVNHQNASAVHIAVLQNFPAIVKVLIDTECNLDIPDNRQQTPLHIAAEHGRQDIAEMILIAGVNLKLIDKQGKTSLDVAARGNHINLADMIIKADRFYKWEKDNLNSDSDSWVAKHLTFKQDHKLETQHIRSVMWRLATKYLKPSEWKKLAHYWKFTDAHIRAIEQQWAGTKSYREHGHRMLLIWLHGVIMAGENPIKGLYEGLVGICRRDLAESIRKKANAESTSPRKCTAM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HAR7</accession>
    <name>A0A452HAR7_9SAUR</name>
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        <submittedName>
            <fullName evidence="4">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
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            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
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    <dbReference type="Ensembl" id="ENSGAGT00000013548.1">
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        <property type="gene ID" value="ENSGAGG00000009113.1"/>
    </dbReference>
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        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="term" value="C:Z disc"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071691">
        <property type="term" value="P:cardiac muscle thin filament assembly"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="entry name" value="NEBU/NEBL"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000900">
        <property type="entry name" value="Nebulin_repeat"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11039:SF48">
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    </dbReference>
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    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="4"/>
    </dbReference>
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    </dbReference>
    <proteinExistence type="predicted"/>
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    <keyword evidence="3" id="KW-0472">Membrane</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="1" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="3" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="3">
        <location>
            <begin position="113"/>
            <end position="133"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011833.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="153" mass="17356" checksum="9110886B5B67C1DD" modified="2019-05-08" version="1">MLCNYCTIPDTPEIERIKNTQKNISSVSSKEVKHATAISDPPEIRRVKENQKNISNVQYKEQPCKATPVTVTPEMERVKKNQEIVHYKGQPAKATSVSVTPEMERVKKNQDNISSASCLNLFALFIFITGYAFQFRFDVINASHCIIHLKVNS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
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    <name>A0A452HAR9_9SAUR</name>
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        <recommendedName>
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        </recommendedName>
    </protein>
    <organism evidence="3 4">
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            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
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            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HAR9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013536.1">
        <property type="protein sequence ID" value="ENSGAGP00000011821.1"/>
        <property type="gene ID" value="ENSGAGG00000009100.1"/>
    </dbReference>
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    </dbReference>
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
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        <property type="match status" value="1"/>
    </dbReference>
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    </dbReference>
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    </dbReference>
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        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
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        <property type="entry name" value="DEATH-like_dom_sf"/>
    </dbReference>
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        <property type="entry name" value="Death_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24125">
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    </dbReference>
    <dbReference type="PANTHER" id="PTHR24125:SF0">
        <property type="entry name" value="ANKYRIN REPEAT AND DEATH DOMAIN-CONTAINING PROTEIN 1A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00023">
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        <property type="match status" value="1"/>
    </dbReference>
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        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00531">
        <property type="entry name" value="Death"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01415">
        <property type="entry name" value="ANKYRIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
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        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47986">
        <property type="entry name" value="DEATH domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="7"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50017">
        <property type="entry name" value="DEATH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="repeat" description="ANK" evidence="1">
        <location>
            <begin position="17"/>
            <end position="49"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="1">
        <location>
            <begin position="50"/>
            <end position="82"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="1">
        <location>
            <begin position="83"/>
            <end position="115"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="1">
        <location>
            <begin position="118"/>
            <end position="150"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="1">
        <location>
            <begin position="151"/>
            <end position="183"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="1">
        <location>
            <begin position="184"/>
            <end position="216"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="1">
        <location>
            <begin position="217"/>
            <end position="249"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="1">
        <location>
            <begin position="250"/>
            <end position="282"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="1">
        <location>
            <begin position="283"/>
            <end position="315"/>
        </location>
    </feature>
    <feature type="domain" description="Death" evidence="2">
        <location>
            <begin position="391"/>
            <end position="465"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="PROSITE" id="PS50017"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011821.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="478" mass="53715" checksum="3D0A46E92F86CDBF" modified="2019-05-08" version="1">MQELIRTRVNVKAKNNTDRTALHWAAGAGNEQAVRLLLDHEVPVDDKDSFGMNALLLSAWFGHLRVLQILVNAGAKINCVDKNGRNLLHCAAQRGHIKVMEFIMEDLEDVHVDKTDKMDKTAFHLAAEHGRLEVVEFLIGSGCSHSVKDKEKNTALHLAAKNGHPTVLQKIVEIGQDLEEKNMEGLTALHLAVEGGHYDCVKVLLEAGCDVNAQTQKAMTCLHYAALHGYEDMARIFIDAGIYIDAVNHQNASAVHIAVLQNFPAIVKVLIDTECNLDIPDNRQQTPLHIAAEHGRQDIAEMILIAGVNLKLIDKQGKTSLDVAARGNHINLADMIIKADRFYKWEKDNLNSDSDSWVAKHLTFKQDHKLETQHIRSVMWRLATKYLKPSEWKKLAHYWKFTDAHIRAIEQQWAGTKSYREHGHRMLLIWLHGVIMAGENPIKGLYEGLVGICRRDLAESIRKKANAESTSPRKCTAM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HAS2</accession>
    <name>A0A452HAS2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">BOS complex subunit NCLN</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Endoplasmic reticulum membrane</location>
            <topology evidence="1">Single-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the nicastrin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HAS2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013542.1">
        <property type="protein sequence ID" value="ENSGAGP00000011827.1"/>
        <property type="gene ID" value="ENSGAGG00000009044.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005789">
        <property type="term" value="C:endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0009966">
        <property type="term" value="P:regulation of signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd03882">
        <property type="entry name" value="M28_nicalin_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.630.10:FF:000021">
        <property type="entry name" value="Nicalin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.630.10">
        <property type="entry name" value="Zn peptidases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016574">
        <property type="entry name" value="Nicalin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007484">
        <property type="entry name" value="Peptidase_M28"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR31826">
        <property type="entry name" value="NICALIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04389">
        <property type="entry name" value="Peptidase_M28"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53187">
        <property type="entry name" value="Zn-dependent exopeptidases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="8" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7" id="KW-0472">Membrane</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 10" id="KW-0732">Signal</keyword>
    <keyword evidence="3" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6" id="KW-1133">Transmembrane helix</keyword>
    <feature type="signal peptide" evidence="10">
        <location>
            <begin position="1"/>
            <end position="41"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019516748" description="BOS complex subunit NCLN" evidence="10">
        <location>
            <begin position="42"/>
            <end position="416"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase M28" evidence="11">
        <location>
            <begin position="216"/>
            <end position="370"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004389"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007717"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00034873"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="Pfam" id="PF04389"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011827.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="416" mass="46030" checksum="14591AE0C2B9D81C" modified="2019-05-08" version="1">MLEEAGEVLESVLKASCLPLSFLLFVPAVLLLLGPPPAAEAAHEFTVYRMQQYELGGQPYGTRNAVLNTEARTLEADVLSRRCVMMRLVDFSYEQYQKALRQSAGAVVIILPKAMSALPQDVVRQFMEIEPEMLAMETIVPVYFAVEDEELLSIYEQTRVASTSQGSASAAEVLLHTATANGFQMVTSGAQSKAVSDWLITSLEGRLTGLGGEDLPTIVIVAHYDSFGVAPWLSHGADSNGSGISGLLELARLFSRLYTYKRTHAGYNLLFFASGGGKFNYQGTKRWLEDNLDHTDSSLLQDNVAFVLCLDTLGRGNSLHLHVSKPPKEGTLQHAFLRELEMVVASQFPEVKFSMVHKKINLAEDMLAWEHERFAIRRLPAFTISHLESHRDGLRNSIMDLRSVALRGGHQQKISC</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HAS4</accession>
    <name>A0A452HAS4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">ATP-dependent clpX-like chaperone, mitochondrial</fullName>
            <ecNumber evidence="2">3.6.4.10</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="18">ATP-dependent Clp protease ATP-binding subunit clpX-like, mitochondrial</fullName>
        </alternativeName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="21" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="15">ATP-dependent chaperone that functions as an unfoldase. As part of the ClpXP protease complex, it recognizes specific protein substrates, unfolds them using energy derived from ATP hydrolysis, and then translocates them to the proteolytic subunit (CLPP) of the ClpXP complex for degradation. Thanks to its chaperone activity, it also functions in the incorporation of the pyridoxal phosphate cofactor into 5-aminolevulinate synthase, thereby activating 5-aminolevulinate (ALA) synthesis, the first step in heme biosynthesis. This chaperone is also involved in the control of mtDNA nucleoid distribution, by regulating mitochondrial transcription factor A (TFAM) activity.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="14">
            <text>ATP + H2O = ADP + phosphate + H(+)</text>
            <dbReference type="Rhea" id="RHEA:13065"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="3.6.4.10"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="14">
            <dbReference type="Rhea" id="RHEA:13066"/>
        </physiologicalReaction>
    </comment>
    <comment type="subunit">
        <text evidence="16">Homohexamer that forms a ring structure; this hexamerization requires ATP binding. Component of the ClpXP complex formed by the assembly of two CLPP heptameric rings with two CLPX hexameric rings, giving rise to a symmetrical structure with two central CLPP rings flanked by a CLPX ring at either end of the complex. Interacts with TFAM.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Mitochondrion matrix</location>
            <location evidence="1">Mitochondrion nucleoid</location>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="3.6.4.10" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452HAS4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013566.1">
        <property type="protein sequence ID" value="ENSGAGP00000011847.1"/>
        <property type="gene ID" value="ENSGAGG00000009121.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042645">
        <property type="term" value="C:mitochondrial nucleoid"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016887">
        <property type="term" value="F:ATP hydrolysis activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046983">
        <property type="term" value="F:protein dimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051603">
        <property type="term" value="P:proteolysis involved in protein catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd19497">
        <property type="entry name" value="RecA-like_ClpX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.8.60:FF:000002">
        <property type="entry name" value="ATP-dependent Clp protease ATP-binding subunit ClpX"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000378">
        <property type="entry name" value="ATP-dependent Clp protease ATP-binding subunit clpX-like, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:003247">
        <property type="entry name" value="ATP-dependent Clp protease ATP-binding subunit clpX-like, mitochondrial"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.8.60">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003593">
        <property type="entry name" value="AAA+_ATPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050052">
        <property type="entry name" value="ATP-dep_Clp_protease_ClpX"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003959">
        <property type="entry name" value="ATPase_AAA_core"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019489">
        <property type="entry name" value="Clp_ATPase_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR059188">
        <property type="entry name" value="Znf_CLPX-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR059067">
        <property type="entry name" value="Znf_ribbon_CLPX-like"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF003745">
        <property type="entry name" value="PRK05342.1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48102:SF7">
        <property type="entry name" value="ATP-DEPENDENT CLP PROTEASE ATP-BINDING SUBUNIT CLPX-LIKE, MITOCHONDRIAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR48102">
        <property type="entry name" value="ATP-DEPENDENT CLP PROTEASE ATP-BINDING SUBUNIT CLPX-LIKE, MITOCHONDRIAL-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07724">
        <property type="entry name" value="AAA_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10431">
        <property type="entry name" value="ClpB_D2-small"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF26040">
        <property type="entry name" value="Zn_ribbon_CLPX_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00382">
        <property type="entry name" value="AAA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM01086">
        <property type="entry name" value="ClpB_D2-small"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51902">
        <property type="entry name" value="CLPX_ZB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="10" id="KW-0007">Acetylation</keyword>
    <keyword evidence="8" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="12" id="KW-0143">Chaperone</keyword>
    <keyword evidence="6" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="11" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="13" id="KW-1135">Mitochondrion nucleoid</keyword>
    <keyword evidence="5" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="3" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0809">Transit peptide</keyword>
    <keyword evidence="7" id="KW-0862">Zinc</keyword>
    <feature type="domain" description="ClpX-type ZB" evidence="20">
        <location>
            <begin position="86"/>
            <end position="139"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="61"/>
            <end position="94"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="592"/>
            <end position="626"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="62"/>
            <end position="76"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="19">
        <location>
            <begin position="77"/>
            <end position="88"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="592"/>
            <end position="603"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="19">
        <location>
            <begin position="604"/>
            <end position="615"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004436"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00012554"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023186"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023271"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00050527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00058714"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00065499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00073042"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00077425"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS51902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011847.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="626" mass="69165" checksum="0D93164957004802" modified="2019-05-08" version="1">MKRSSPYTIYWHFRCLLINGITCGRTCLAVLGRIGTFETQSLRRIPFRTFSETPAYFASKDGVSKDGSGDGNKKSVSEGGSKKSSSGNSGKGGNQLRCPKCGDLCTHVETFVSSTRFVKCEKCHHFFVVLSEADSKKSIIKEPESAAEAVKLAFQQKPPPPPKKIYNYLDKYVVGQSFAKKVLSVAVYNHYKRIYNNIPANLRQQAEVEKQTSLTPRELEIRRREDEYRFTKLLQIAGISPHGNALGASMQQQVNQQMPQEKRGGEVLDSTHDDINLEKSNILLLGPTGSGKTLLAQTLAKCLDVPFAICDCTTLTQAGYVGEDIESVIAKLLQDANYNVEKAQQGIVFLDEVDKIGSVPGIHQLRDVGGEGVQQGLLKLLEGTIVNVPEKNSRKLRGETVQVDTTNVLFVASGAFNGLDRIISRRKNEKYLGFGTPSNLGKGRRAAAAADLANISGESNTQQDIEEKDRLLRHVEARDLIEFGMIPEFVGRLPVVVPLHSLDEKTLVRILTEPRNAVVPQYQALFSMDKCELTVTEDALKAIARLALDRKTGARGLRSIMEKLLLEPMFEVPNSDIVCVEVDKEVVEGKKEPGYIRAPTKDTSEEEYDSGVEEEGWPRQADAANN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HB08</accession>
    <name>A0A452HB08_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="4">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="4 5">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="5" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="4" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="4" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HB08"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013653.1">
        <property type="protein sequence ID" value="ENSGAGP00000011920.1"/>
        <property type="gene ID" value="ENSGAGG00000004156.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd01249">
        <property type="entry name" value="BAR-PH_GRAF_family"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.555.10:FF:000006">
        <property type="entry name" value="Rho GTPase activating protein 26"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000116">
        <property type="entry name" value="Rho GTPase activating protein 26"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1270.60:FF:000086">
        <property type="entry name" value="Rho GTPase-activating protein 10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.60">
        <property type="entry name" value="Arfaptin homology (AH) domain/BAR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.555.10">
        <property type="entry name" value="Rho GTPase activation protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027267">
        <property type="entry name" value="AH/BAR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004148">
        <property type="entry name" value="BAR_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047234">
        <property type="entry name" value="GRAF_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047225">
        <property type="entry name" value="PH_GRAF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000198">
        <property type="entry name" value="RhoGAP_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12552">
        <property type="entry name" value="OLIGOPHRENIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12552:SF4">
        <property type="entry name" value="RHO GTPASE-ACTIVATING PROTEIN 26"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16746">
        <property type="entry name" value="BAR_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00169">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00620">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00233">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00324">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103657">
        <property type="entry name" value="BAR/IMD domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50238">
        <property type="entry name" value="RHOGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="5" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="PH" evidence="2">
        <location>
            <begin position="235"/>
            <end position="339"/>
        </location>
    </feature>
    <feature type="domain" description="Rho-GAP" evidence="3">
        <location>
            <begin position="345"/>
            <end position="530"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="3">
        <source>
            <dbReference type="PROSITE" id="PS50238"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011920.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="677" mass="77769" checksum="AF0000A9B80A91E0" modified="2019-05-08" version="1">MGLPALEFSDCCLDSPQFRERLKSHEAELDKTNKFIKELIKDGKSLVAALKNLSSAKRKFADSLNEFKFRCIGDIENAGEVLITPLEKFRKEQIGAAKDAKKKYDKETEKYCGVLEKHLNLSSKKKESQLQEADSQVDLVRQHFYEVSLEYVFKVQEVQERKMFEFVEPLLAFLQGLFTFYHHGYELAKDFSDFKTELTISIQNTRNRFEGTRSEVESLMRKMKENPEEHRNISPFTMEGYLYVQEKRHFGTSWVKHYCTYQRESKRITMVPFDQKSGGKGGEDEAVTLKSCIRRKTDSIEKRFCFDVEAVDRPGVITMQALSEEDRKLWMEAMDGREPVRRDTAQLDNIGFSIIRKCINAVETRGINEQGLYRIVGVNSRVQKLLSILMDPKTASETETDICAEWEIKTITSALKTYLRMLPGPLMMYQFQRSFIKAAKLENQESRISEIHNLVHRLPEKNRQMLQMLMNHLANVANNHKQNLMTVANLGVVFGPTLLRPQEETVAAIMDIKFQNIVVEILIENHEKIFNTVPETLPPNSQLLLLSHKKNTDSKPPSCSERPLTLFHTAQANENQETRNSIINSSLESVISSNVNSFLHSNSTLRPSLNSSDPDLEVIKPSRPNSCSATSGAQSSHQYQCMHQFMIFDVFLHIWCEAVMHSCLSSIPSSYRLGIRM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HB33</accession>
    <name>A0A452HB33_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Rho GTPase-activating protein 26</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="10">Rho-type GTPase-activating protein 26</fullName>
        </alternativeName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell junction</location>
            <location evidence="2">Focal adhesion</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Endosome membrane</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HB33"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013689.1">
        <property type="protein sequence ID" value="ENSGAGP00000011951.1"/>
        <property type="gene ID" value="ENSGAGG00000004156.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010008">
        <property type="term" value="C:endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005925">
        <property type="term" value="C:focal adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd01249">
        <property type="entry name" value="BAR-PH_GRAF_family"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd07636">
        <property type="entry name" value="BAR_GRAF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.555.10:FF:000006">
        <property type="entry name" value="Rho GTPase activating protein 26"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000116">
        <property type="entry name" value="Rho GTPase activating protein 26"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1270.60:FF:000001">
        <property type="entry name" value="Rho GTPase-activating protein 26"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.60">
        <property type="entry name" value="Arfaptin homology (AH) domain/BAR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.555.10">
        <property type="entry name" value="Rho GTPase activation protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027267">
        <property type="entry name" value="AH/BAR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004148">
        <property type="entry name" value="BAR_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035483">
        <property type="entry name" value="GRAF_BAR"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047234">
        <property type="entry name" value="GRAF_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047225">
        <property type="entry name" value="PH_GRAF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000198">
        <property type="entry name" value="RhoGAP_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12552">
        <property type="entry name" value="OLIGOPHRENIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12552:SF4">
        <property type="entry name" value="RHO GTPASE-ACTIVATING PROTEIN 26"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16746">
        <property type="entry name" value="BAR_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00169">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00620">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00233">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00324">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103657">
        <property type="entry name" value="BAR/IMD domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50238">
        <property type="entry name" value="RHOGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0965">Cell junction</keyword>
    <keyword evidence="9" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="6" id="KW-0967">Endosome</keyword>
    <keyword evidence="5" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="8" id="KW-0472">Membrane</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="PH" evidence="11">
        <location>
            <begin position="265"/>
            <end position="369"/>
        </location>
    </feature>
    <feature type="domain" description="Rho-GAP" evidence="12">
        <location>
            <begin position="375"/>
            <end position="560"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004608"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00020525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00032211"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50238"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011951.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="688" mass="79260" checksum="BF5823844C1C0FDE" modified="2019-05-08" version="1">MGLPALEFSDCCLDSPQFRERLKSHEAELDKTNKFIKELIKDGKSLVAALKNLSSAKRKFADSLNEFKFRCIGDAETDDEICIAKSLQEFATVLRNLEDERMRMIENAGEVLITPLEKFRKEQIGAAKDAKKKYDKETEKYCGVLEKHLNLSSKKKESQLQEADSQVDLVRQHFYEVSLEYVFKVQEVQERKMFEFVEPLLAFLQGLFTFYHHGYELAKDFSDFKTELTISIQNTRNRFEGTRSEVESLMRKMKENPEEHRNISPFTMEGYLYVQEKRHFGTSWVKHYCTYQRESKRITMVPFDQKSGGKGGEDEAVTLKSCIRRKTDSIEKRFCFDVEAVDRPGVITMQALSEEDRKLWMEAMDGREPVRRDTAQLDNIGFSIIRKCINAVETRGINEQGLYRIVGVNSRVQKLLSILMDPKTASETETDICAEWEIKTITSALKTYLRMLPGPLMMYQFQRSFIKAAKLENQESRISEIHNLVHRLPEKNRQMLQMLMNHLANVANNHKQNLMTVANLGVVFGPTLLRPQEETVAAIMDIKFQNIVVEILIENHEKIFNTVPETLPPNSQLLLLSHKKNTDSKPPSCSERPLTLFHTAQANENQETRNSIINSSLESVISSNVNSFLHSNSTLRPSLNSSDPDLEVIKPSRPNSLPQRYIPCHHMANINTEQNWVQDSSCQRTHSL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HB38</accession>
    <name>A0A452HB38_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">Rho GTPase-activating protein 26</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="10">Rho-type GTPase-activating protein 26</fullName>
        </alternativeName>
    </protein>
    <organism evidence="13 14">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="14" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="13" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="13" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cell junction</location>
            <location evidence="2">Focal adhesion</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Endosome membrane</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HB38"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013694.1">
        <property type="protein sequence ID" value="ENSGAGP00000011956.1"/>
        <property type="gene ID" value="ENSGAGG00000004156.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005856">
        <property type="term" value="C:cytoskeleton"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010008">
        <property type="term" value="C:endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005925">
        <property type="term" value="C:focal adhesion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005096">
        <property type="term" value="F:GTPase activator activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007165">
        <property type="term" value="P:signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd01249">
        <property type="entry name" value="BAR-PH_GRAF_family"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.555.10:FF:000006">
        <property type="entry name" value="Rho GTPase activating protein 26"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.29.30:FF:000116">
        <property type="entry name" value="Rho GTPase activating protein 26"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.1270.60:FF:000001">
        <property type="entry name" value="Rho GTPase-activating protein 26"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.1270.60">
        <property type="entry name" value="Arfaptin homology (AH) domain/BAR domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.29.30">
        <property type="entry name" value="Pleckstrin-homology domain (PH domain)/Phosphotyrosine-binding domain (PTB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.555.10">
        <property type="entry name" value="Rho GTPase activation protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027267">
        <property type="entry name" value="AH/BAR_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004148">
        <property type="entry name" value="BAR_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047234">
        <property type="entry name" value="GRAF_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011993">
        <property type="entry name" value="PH-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001849">
        <property type="entry name" value="PH_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047225">
        <property type="entry name" value="PH_GRAF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008936">
        <property type="entry name" value="Rho_GTPase_activation_prot"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000198">
        <property type="entry name" value="RhoGAP_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12552">
        <property type="entry name" value="OLIGOPHRENIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12552:SF4">
        <property type="entry name" value="RHO GTPASE-ACTIVATING PROTEIN 26"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16746">
        <property type="entry name" value="BAR_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00169">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00620">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00233">
        <property type="entry name" value="PH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00324">
        <property type="entry name" value="RhoGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF103657">
        <property type="entry name" value="BAR/IMD domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48350">
        <property type="entry name" value="GTPase activation domain, GAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50729">
        <property type="entry name" value="PH domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50003">
        <property type="entry name" value="PH_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50238">
        <property type="entry name" value="RHOGAP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0965">Cell junction</keyword>
    <keyword evidence="9" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="6" id="KW-0967">Endosome</keyword>
    <keyword evidence="5" id="KW-0343">GTPase activation</keyword>
    <keyword evidence="8" id="KW-0472">Membrane</keyword>
    <keyword evidence="14" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="PH" evidence="11">
        <location>
            <begin position="234"/>
            <end position="338"/>
        </location>
    </feature>
    <feature type="domain" description="Rho-GAP" evidence="12">
        <location>
            <begin position="344"/>
            <end position="529"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004246"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004608"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00020525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022468"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022949"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00032211"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50003"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="PROSITE" id="PS50238"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011956.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="676" mass="77529" checksum="95AFD3FAD8293943" modified="2019-05-08" version="1">MQTTVPAPPKASCSDRGIPGDLSSAKRKFADSLNEFKFRCIGDAETDDEICIAKSLQEFATVLRNLEDERMRMIENAGEVLITPLEKFRKEQIGAAKDAKKKYDKETEKYCGVLEKHLNLSSKKKESQLQEADSQVDLVRQHFYEVSLEYVFKVQEVQERKMFEFVEPLLAFLQGLFTFYHHGYELAKDFSDFKTELTISIQNTRNRFEGTRSEVESLMRKMKENPEEHRNISPFTMEGYLYVQEKRHFGTSWVKHYCTYQRESKRITMVPFDQKSGGKGGEDEAVTLKSCIRRKTDSIEKRFCFDVEAVDRPGVITMQALSEEDRKLWMEAMDGREPVRRDTAQLDNIGFSIIRKCINAVETRGINEQGLYRIVGVNSRVQKLLSILMDPKTASETETDICAEWEIKTITSALKTYLRMLPGPLMMYQFQRSFIKAAKLENQESRISEIHNLVHRLPEKNRQMLQMLMNHLANVANNHKQNLMTVANLGVVFGPTLLRPQEETVAAIMDIKFQNIVVEILIENHEKIFNTVPETLPPNSQLLLLSHKKNTDSKPPSCSERPLTLFHTAQANENQETRNSIINSSLESVISSNVNSFLHSNSTLRPSLNSSDPDLEVIKPSRPNSCSATSGAQSSHQYQCMHQFMIFDVFLHIWCEAVMHSCLSSIPSSYRLGIRM</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HB40</accession>
    <name>A0A452HB40_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">Ig-like domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HB40"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013701.1">
        <property type="protein sequence ID" value="ENSGAGP00000011963.1"/>
        <property type="gene ID" value="ENSGAGG00000009213.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004672">
        <property type="term" value="F:protein kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd20972">
        <property type="entry name" value="IgI_2_Titin_Z1z2-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000256">
        <property type="entry name" value="myopalladin isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000399">
        <property type="entry name" value="myopalladin isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013098">
        <property type="entry name" value="Ig_I-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003598">
        <property type="entry name" value="Ig_sub2"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47633">
        <property type="entry name" value="IMMUNOGLOBULIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47633:SF10">
        <property type="entry name" value="PALLADIN, CYTOSKELETAL ASSOCIATED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07679">
        <property type="entry name" value="I-set"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00408">
        <property type="entry name" value="IGc2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="Ig-like" evidence="2">
        <location>
            <begin position="287"/>
            <end position="376"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="2">
        <location>
            <begin position="457"/>
            <end position="553"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="53"/>
            <end position="83"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="98"/>
            <end position="174"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="198"/>
            <end position="239"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="627"/>
            <end position="662"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="53"/>
            <end position="77"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="1">
        <location>
            <begin position="101"/>
            <end position="123"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="132"/>
            <end position="144"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="156"/>
            <end position="170"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="1">
        <location>
            <begin position="203"/>
            <end position="214"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="224"/>
            <end position="235"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="1">
        <location>
            <begin position="642"/>
            <end position="654"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011963.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="685" mass="74390" checksum="F637FEFEAA635EA0" modified="2019-05-08" version="1">MSEASSFHDFFYDSLSGIQDGEVSNELSAFLSKEEINKSLDLAREAIANSVKENQNVEQDFTHNFNTSPNSSEYTSSRKTKVHEQEVLGRLQVSSPNSLFSTTQTLPKQQKQQPPTSPQAASSICTVPGRQGSISPSLTTSPSFIRSLRNAERCSPNMQKTSPKSKSMSQGDAPFRNKLCDKAAMFIEELSSIFREAAKARGRSPNGDSSSPDSGYLSPKKKQSATINASVNQSHGKACPEIKPEAKLPETHHNGELFPDRKNVTQENETAFHHELTNPGSYQSSAPRFTQKLRSQEVAEGSKVLLECRVAGNPTPHVRWFCEGKELQNSPDIQIHCESGGLHSLVIAEAFEDDTGRYTCLASNSVGSDSTSAEVFIEGVSSTDSESESLVFKSKSGAMPQAQKKTTSVSLTIGSSTPKAGVTTAVIQPISVPIQQVQSPTSYLYRLDGSKPVCSTPIFTKELQNSTASEGQVVVLECRIRGAPPLHVEWFRQGVEIQDSPDFRILQKKPRSAAELEEICTLVIAETFPEDAGTFTCTATNEYGSVASSAQLTVCSANSESFSHELSTRESNSDDFQHFPPPPPILEISSLELPPKKHTESQQVNNTEFRPNIAAVQLQFNSAEKKTSGIHPSHEANGVINSKASSDKSAPAPAVLLSPTKEPPPVLVYTCFVSPGKPKAAEVYW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HB47</accession>
    <name>A0A452HB47_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">CUB domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasmic vesicle</location>
            <location evidence="1">Secretory vesicle</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="7">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HB47"/>
    <dbReference type="Ensembl" id="ENSGAGT00000013708.1">
        <property type="protein sequence ID" value="ENSGAGP00000011970.1"/>
        <property type="gene ID" value="ENSGAGG00000009219.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030133">
        <property type="term" value="C:transport vesicle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="CDD" id="cd00041">
        <property type="entry name" value="CUB"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="CDD" id="cd00112">
        <property type="entry name" value="LDLa"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.400.10:FF:000034">
        <property type="entry name" value="Low-density lipoprotein receptor-related protein 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.290:FF:000013">
        <property type="entry name" value="Membrane frizzled-related protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.290:FF:000005">
        <property type="entry name" value="Procollagen C-endopeptidase enhancer 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="4.10.400.10:FF:000065">
        <property type="entry name" value="Transmembrane protease serine 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.400.10">
        <property type="entry name" value="Low-density Lipoprotein Receptor"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.290">
        <property type="entry name" value="Spermadhesin, CUB domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000859">
        <property type="entry name" value="CUB_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036055">
        <property type="entry name" value="LDL_receptor-like_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023415">
        <property type="entry name" value="LDLR_class-A_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002172">
        <property type="entry name" value="LDrepeatLR_classA_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035914">
        <property type="entry name" value="Sperma_CUB_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24251:SF30">
        <property type="entry name" value="MEMBRANE FRIZZLED-RELATED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24251">
        <property type="entry name" value="OVOCHYMASE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00431">
        <property type="entry name" value="CUB"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00057">
        <property type="entry name" value="Ldl_recept_a"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00261">
        <property type="entry name" value="LDLRECEPTOR"/>
    </dbReference>
    <dbReference type="SMART" id="SM00042">
        <property type="entry name" value="CUB"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00192">
        <property type="entry name" value="LDLa"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57424">
        <property type="entry name" value="LDL receptor-like module"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49854">
        <property type="entry name" value="Spermadhesin, CUB domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01180">
        <property type="entry name" value="CUB"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01209">
        <property type="entry name" value="LDLRA_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50068">
        <property type="entry name" value="LDLRA_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="6" id="KW-0968">Cytoplasmic vesicle</keyword>
    <keyword evidence="4 8" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="5" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="2 10" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="10">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019343290" description="CUB domain-containing protein" evidence="10">
        <location>
            <begin position="23"/>
            <end position="394"/>
        </location>
    </feature>
    <feature type="domain" description="CUB" evidence="11">
        <location>
            <begin position="66"/>
            <end position="182"/>
        </location>
    </feature>
    <feature type="domain" description="CUB" evidence="11">
        <location>
            <begin position="230"/>
            <end position="343"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="26"/>
            <end position="81"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="8">
        <location>
            <begin position="189"/>
            <end position="201"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="8">
        <location>
            <begin position="196"/>
            <end position="214"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="8">
        <location>
            <begin position="208"/>
            <end position="223"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="8">
        <location>
            <begin position="350"/>
            <end position="362"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="8">
        <location>
            <begin position="357"/>
            <end position="375"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="8">
        <location>
            <begin position="369"/>
            <end position="384"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004398"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023329"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00059"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00124"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS01180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000011970.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="394" mass="42514" checksum="1A35CB89FC72CB1B" modified="2019-05-08" version="1">MGSLLLLLLALLLGIIISQLQSSRPAGPSSVTASTQHHDATITAPASTAEPMATVPPPATTPEPSCGGSLQGPEGSLSSPRYPAPYPPNTLCIWRIQVEDGLAIQLKIESLSLEGTRWCLFDRVELYEELGAASGSPWGGTSRFCGDVAPPTINTNSNRLRVIFVSDNSVGALGFTAHYRAILPSDKSCAWDEFFCDQGLCIHLGLVCDGFHDCTDKSDEANCSSRHKECGGSLTNLEGQFSTPSHPQPYPHQQLCLWQIAVPVGHVIDLQFHNFSLEWQEDCTFDFVEVHDSAGTDASSLLGRFCGSQLPPILTSSRHVMTVLFMADEGVAEDGFFATYRARNATEKTCSPSEFSCSNGECQARELVCDGWHDCLDGSDEFNCTNVSPPSFGE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HRS0</accession>
    <name>A0A452HRS0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="36">Tyrosine-protein kinase receptor</fullName>
            <ecNumber evidence="36">2.7.10.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="42 43">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="43" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="42" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="42" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="31 36">
            <text>L-tyrosyl-[protein] + ATP = O-phospho-L-tyrosyl-[protein] + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:10596"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.10.1"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="3">Cell projection</location>
            <location evidence="3">Axon</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cell projection</location>
            <location evidence="2">Dendrite</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="5">Cytoplasm</location>
            <location evidence="5">Perinuclear region</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Endosome membrane</location>
            <topology evidence="4">Single-pass type I membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="30">Postsynaptic density</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="36">Belongs to the protein kinase superfamily. Tyr protein kinase family. Insulin receptor subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.10.1" evidence="36"/>
    <dbReference type="AlphaFoldDB" id="A0A452HRS0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000020273.1">
        <property type="protein sequence ID" value="ENSGAGP00000017773.1"/>
        <property type="gene ID" value="ENSGAGG00000013192.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043679">
        <property type="term" value="C:axon terminus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043197">
        <property type="term" value="C:dendritic spine"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010008">
        <property type="term" value="C:endosome membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048471">
        <property type="term" value="C:perinuclear region of cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0014069">
        <property type="term" value="C:postsynaptic density"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043235">
        <property type="term" value="C:receptor complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048403">
        <property type="term" value="F:brain-derived neurotrophic factor binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0060175">
        <property type="term" value="F:brain-derived neurotrophic factor receptor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:1990416">
        <property type="term" value="P:cellular response to brain-derived neurotrophic factor stimulus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007399">
        <property type="term" value="P:nervous system development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
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        <property type="term" value="P:trans-synaptic signaling by neuropeptide, modulating synaptic transmission"/>
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        <property type="project" value="TreeGrafter"/>
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        <property type="entry name" value="IgI_TrkB_d5"/>
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        <property type="entry name" value="PTKc_TrkB"/>
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        <property type="match status" value="2"/>
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        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Ribonuclease Inhibitor"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
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        <property type="entry name" value="Cys-rich_flank_reg_C"/>
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        <property type="entry name" value="Ig-like_fold"/>
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        <property type="entry name" value="BDNF_NT-3 GROWTH FACTORS RECEPTOR"/>
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        <property type="entry name" value="PK_Tyr_Ser-Thr"/>
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        <property type="entry name" value="TYRKINASE"/>
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        <property type="entry name" value="TyrKc"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52058">
        <property type="entry name" value="L domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="IG_LIKE"/>
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    </dbReference>
    <dbReference type="PROSITE" id="PS00107">
        <property type="entry name" value="PROTEIN_KINASE_ATP"/>
        <property type="match status" value="1"/>
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        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
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    <dbReference type="PROSITE" id="PS00109">
        <property type="entry name" value="PROTEIN_KINASE_TYR"/>
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    <dbReference type="PROSITE" id="PS00239">
        <property type="entry name" value="RECEPTOR_TYR_KIN_II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="19 33 35" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="7" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="28" id="KW-0966">Cell projection</keyword>
    <keyword evidence="8" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="18" id="KW-0221">Differentiation</keyword>
    <keyword evidence="25" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="16" id="KW-0967">Endosome</keyword>
    <keyword evidence="27" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="29" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="17" id="KW-0418">Kinase</keyword>
    <keyword evidence="10" id="KW-0433">Leucine-rich repeat</keyword>
    <keyword evidence="23 38" id="KW-0472">Membrane</keyword>
    <keyword evidence="20" id="KW-0524">Neurogenesis</keyword>
    <keyword evidence="15 33 35" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="9 36" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="26 36" id="KW-0675">Receptor</keyword>
    <keyword evidence="43" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="14" id="KW-0677">Repeat</keyword>
    <keyword evidence="13 39" id="KW-0732">Signal</keyword>
    <keyword evidence="22" id="KW-0770">Synapse</keyword>
    <keyword evidence="11" id="KW-0808">Transferase</keyword>
    <keyword evidence="12 36 38" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="21 38" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="24" id="KW-0829">Tyrosine-protein kinase</keyword>
    <feature type="signal peptide" evidence="39">
        <location>
            <begin position="1"/>
            <end position="32"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5018968650" description="Tyrosine-protein kinase receptor" evidence="39">
        <location>
            <begin position="33"/>
            <end position="796"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="38">
        <location>
            <begin position="405"/>
            <end position="428"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="41">
        <location>
            <begin position="174"/>
            <end position="259"/>
        </location>
    </feature>
    <feature type="domain" description="Protein kinase" evidence="40">
        <location>
            <begin position="512"/>
            <end position="781"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="37">
        <location>
            <begin position="447"/>
            <end position="472"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="37">
        <location>
            <begin position="459"/>
            <end position="469"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="32">
        <location>
            <position position="650"/>
        </location>
    </feature>
    <feature type="binding site" evidence="33">
        <location>
            <begin position="518"/>
            <end position="526"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="33 35">
        <location>
            <position position="546"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="site" description="Interaction with SHC1" evidence="34">
        <location>
            <position position="490"/>
        </location>
    </feature>
    <feature type="site" description="Interaction with PLCG1" evidence="34">
        <location>
            <position position="791"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004251"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004279"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004489"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004530"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00004556"/>
        </source>
    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022614"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00022753"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00022902"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="ARBA" id="ARBA00023137"/>
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    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
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    </evidence>
    <evidence type="ECO:0000256" key="26">
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            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="27">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="28">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="29">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="30">
        <source>
            <dbReference type="ARBA" id="ARBA00034105"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="31">
        <source>
            <dbReference type="ARBA" id="ARBA00051243"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="32">
        <source>
            <dbReference type="PIRSR" id="PIRSR620777-50"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="33">
        <source>
            <dbReference type="PIRSR" id="PIRSR620777-51"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="34">
        <source>
            <dbReference type="PIRSR" id="PIRSR620777-52"/>
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    </evidence>
    <evidence type="ECO:0000256" key="35">
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            <dbReference type="PROSITE-ProRule" id="PRU10141"/>
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    </evidence>
    <evidence type="ECO:0000256" key="36">
        <source>
            <dbReference type="RuleBase" id="RU000312"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="37">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="38">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="39">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="40">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="41">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="42">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000017773.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="43">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="796" mass="89276" checksum="EEA51807A5F6344A" modified="2019-05-08" version="1">MVSWRRRWQGPGLARLWGFCCLVFGSWRALLACPAPCTCSASRIWCGEPAPGIVSFPVLERSAERENITDIYIANQRKLESINNNEVGLYVGLKNLNFSRNKLSSLYKKTFRHLNLSDLILVDNPFKCSCDVMWIKIFQETKFSNPETQDFYCINDNNKRIPLSDMQIPNCGLPSANLSVYNLTIVEGRSTTLYCEATGVPSPNVSWVVANLVSKHESNTSKNPASLTIRNVSSKDSGIRISCIAENIVGEDQASADLVVFFAPNITFIEPPTPDHHWCIPFTVKGNPKPSLQWFYEGAILNESEYICTKIHVSNQSEYHGCLQLDNPTHLNNGAYTLLAKNEYGEDEKRVDAHFMSMPGDGSGPILDPELYDYETTPNDIGDTTNNSNQITSTDVSNKENEDSITVYVVVGIAALVCTGLVIMLIILKFGRHSKFGMKGPSSVISNDDDSASPLHHISNGSNTPSSSEGGPDAVIIGMTKIPVIENPQYFGITNSQLKPDTFVQHIKRHNIVLKRELGEGAFGKVFLAECYNLSPEQDKILVAVKTLKDASDNARKDFHREAELLTNLQHEHIVKFYGVCVEGDPLIMVFEYMKHGDLNKFLRAHGPDAVLMAEGNLLAELTQSQMLHIAQQIASGMVYLASQHFVHRDLATRNCLVGENLLVKIGDFGMSRDVYSTDYYRVGGHTMLPIRWMPPESIMYRKFTTESDVWSLGVVLWEIFTYGKQPWYQLSNNEVIECITQGRVLQRPRTCPKEVYDLMLGCWQREPHMRLNIKEIHSLLQNLAKASPVYLDILG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HRS1</accession>
    <name>A0A452HRS1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="3">protein-tyrosine-phosphatase</fullName>
            <ecNumber evidence="3">3.1.3.48</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="21 22">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="22" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="21" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="16 23">
            <text>O-phospho-L-tyrosyl-[protein] + H2O = L-tyrosyl-[protein] + phosphate</text>
            <dbReference type="Rhea" id="RHEA:10684"/>
            <dbReference type="Rhea" id="RHEA-COMP:10136"/>
            <dbReference type="Rhea" id="RHEA-COMP:20101"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:43474"/>
            <dbReference type="ChEBI" id="CHEBI:46858"/>
            <dbReference type="ChEBI" id="CHEBI:61978"/>
            <dbReference type="EC" id="3.1.3.48"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the protein-tyrosine phosphatase family. Receptor class 2B subfamily.</text>
    </comment>
    <dbReference type="EC" id="3.1.3.48" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452HRS1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000020247.1">
        <property type="protein sequence ID" value="ENSGAGP00000017750.1"/>
        <property type="gene ID" value="ENSGAGG00000012313.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000020251.1">
        <property type="protein sequence ID" value="ENSGAGP00000017754.1"/>
        <property type="gene ID" value="ENSGAGG00000012313.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004725">
        <property type="term" value="F:protein tyrosine phosphatase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="CDD" id="cd00063">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="CDD" id="cd06263">
        <property type="entry name" value="MAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000019">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase kappa isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.200:FF:000006">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase T isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000152">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase T isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000009">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase U isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000025">
        <property type="entry name" value="receptor-type tyrosine-protein phosphatase U isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.200">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013320">
        <property type="entry name" value="ConA-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003961">
        <property type="entry name" value="FN3_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057598">
        <property type="entry name" value="Fn3_PTPRU"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036116">
        <property type="entry name" value="FN3_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR007110">
        <property type="entry name" value="Ig-like_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036179">
        <property type="entry name" value="Ig-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003599">
        <property type="entry name" value="Ig_sub"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013151">
        <property type="entry name" value="Immunoglobulin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000998">
        <property type="entry name" value="MAM_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051622">
        <property type="entry name" value="R-tyr_protein_phosphatases"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24051:SF11">
        <property type="entry name" value="PROTEIN TYROSINE PHOSPHATASE, RECEPTOR TYPE, M"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24051">
        <property type="entry name" value="SUSHI DOMAIN-CONTAINING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00041">
        <property type="entry name" value="fn3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23144">
        <property type="entry name" value="Fn3_PTPRU"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00047">
        <property type="entry name" value="ig"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00629">
        <property type="entry name" value="MAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00020">
        <property type="entry name" value="MAMDOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00060">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00409">
        <property type="entry name" value="IG"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00137">
        <property type="entry name" value="MAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49899">
        <property type="entry name" value="Concanavalin A-like lectins/glucanases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49265">
        <property type="entry name" value="Fibronectin type III"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48726">
        <property type="entry name" value="Immunoglobulin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50853">
        <property type="entry name" value="FN3"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50835">
        <property type="entry name" value="IG_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00740">
        <property type="entry name" value="MAM_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50060">
        <property type="entry name" value="MAM_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="14" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="8" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="15" id="KW-0393">Immunoglobulin domain</keyword>
    <keyword evidence="11 17" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="9" id="KW-0904">Protein phosphatase</keyword>
    <keyword evidence="13" id="KW-0675">Receptor</keyword>
    <keyword evidence="22" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7" id="KW-0677">Repeat</keyword>
    <keyword evidence="6" id="KW-0732">Signal</keyword>
    <keyword evidence="5 17" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="10 17" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="17">
        <location>
            <begin position="681"/>
            <end position="702"/>
        </location>
    </feature>
    <feature type="domain" description="MAM" evidence="18">
        <location>
            <begin position="1"/>
            <end position="122"/>
        </location>
    </feature>
    <feature type="domain" description="Ig-like" evidence="19">
        <location>
            <begin position="124"/>
            <end position="215"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="20">
        <location>
            <begin position="222"/>
            <end position="317"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="20">
        <location>
            <begin position="320"/>
            <end position="418"/>
        </location>
    </feature>
    <feature type="domain" description="Fibronectin type-III" evidence="20">
        <location>
            <begin position="419"/>
            <end position="525"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00006396"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00013064"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022912"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023170"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023319"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00051722"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="PROSITE" id="PS50060"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="19">
        <source>
            <dbReference type="PROSITE" id="PS50835"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="20">
        <source>
            <dbReference type="PROSITE" id="PS50853"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000017750.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="873" mass="97563" checksum="FD484A8769A5B548" modified="2019-05-08" version="1">MPSGSFMLVNTSGRFAEQRAHLLLPHLKENDTHCIDFHYYVSSKSGSNPGSLNVYVKVNNGPLGNPIWNMSGAATGTWNRAELAISTFWPNFYQVIFEVVTSGHQGYLAIDEVKVLGHPCTKTPHFLRLQSVEVNAGQFATFQCTANGKTNSGDRLWLQGIYVRDAPLKDIKVSNDRRFVALFNVVNATKRDAGNYRCMIRTEGGAGVSSYAELIVKEPPVPIAPPQLSSVGATYLWIQLNANSINGDGPIIEREVEYHTSSGSWYDRQPVDSTSYKIGHLDPDTEYEISVLLTRPGEGGTGSPGPALRTRTKCADPMRGPRKLEVLEIKSRQITIRWEPFGYNVTRCHRYNLTVHYRYQAGGQEQVREEVSWDTESSHPQHTITNLSPYTNVSVKLVLMNPEGRKESEELVVQTDEDVPSAVPLESIQGSTFEEKIFLQWREPVQTYGVITLYEITYKAVSSFDPEIDLSNQSGRVLKLGNETHYLFFGLYPGTTYSFTIRASTAKGFGPPITNQFTTKISAPSMPPYELETPLNQTDSTVTILLKPAQSRGAPVSVYQIVVEEERPRRTKKTTEILKCYPVPIHFQNASLLNSHYYFAAEFPANSLQAAQSFTIGDNRTYSGFWNTPLLPHKSYSIYFQAASRANGETKIDCVRVATKGAATRKPDPEPEKETDHTVKIAGVIAGILLFVIIFLGVVLVMKKRRSYHSYTYYLKLAKKRKETLSSTRQEMTVMVNSMDKSYTEQGTNCDEAFSFMDTHNLNGRSVSSPSSFTMKTNTLSTTVPNSYYPDETHTMASDTSSLVQSHTYKKRDPVDVPYQTGQLHPAIRVADLLQHITQMKCAEGYGFKEEYEVSARSIGLIVVPSVVLALNL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HRS3</accession>
    <name>A0A452HRS3_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Sulfotransferase</fullName>
            <ecNumber evidence="5">2.8.2.-</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 5">Belongs to the sulfotransferase 1 family.</text>
    </comment>
    <dbReference type="EC" id="2.8.2.-" evidence="5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000020261.1">
        <property type="protein sequence ID" value="ENSGAGP00000017762.1"/>
        <property type="gene ID" value="ENSGAGG00000013102.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008146">
        <property type="term" value="F:sulfotransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000433">
        <property type="entry name" value="Estrogen sulfotransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000863">
        <property type="entry name" value="Sulfotransferase_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11783">
        <property type="entry name" value="SULFOTRANSFERASE SULT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00685">
        <property type="entry name" value="Sulfotransfer_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 5" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Sulfotransferase" evidence="7">
        <location>
            <begin position="63"/>
            <end position="295"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="295"/>
            <end position="329"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="6">
        <location>
            <begin position="300"/>
            <end position="313"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="RuleBase" id="RU361155"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF00685"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000017762.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="329" mass="37822" checksum="27738CDF746E529D" modified="2019-05-08" version="1">MHWDLNPEFQWRGRLWELWDSYGIATHSAMLQKSTLASVHGRTSPNFVLCVVVSGPLTLSSLGTNWMLEILSLIRSDGDPGWVCSVLNWDRVPWVEYGPGLEAARKYPPPRLLCFHLPVQLFPKSLQRSKAKIIYTLRCPKDVVVSLYHFSRIVRLFKDPDSLDCFLEDFLSGNVLYGSWFHHVTGWMGLKANKNIFSITYEELQQDLQGSVRRICHFLGKELSEEQVAAVVENVSFQSMKGNKISNFSQMSDKYMDHQKGEFLRKGICGDWRNHLSEAQSQRFDTVFREWMQGLASPTPQTPYPQPEPFPEPPPHHEPLIXPQSSFSL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HRS4</accession>
    <name>A0A452HRS4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">C3H1-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HRS4"/>
    <dbReference type="Ensembl" id="ENSGAGT00000020266.1">
        <property type="protein sequence ID" value="ENSGAGP00000017766.1"/>
        <property type="gene ID" value="ENSGAGG00000013068.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035198">
        <property type="term" value="F:miRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035196">
        <property type="term" value="P:miRNA processing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.10">
        <property type="entry name" value="Tetratricopeptide repeat domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.1000.10">
        <property type="entry name" value="Zinc finger, CCCH-type"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011990">
        <property type="entry name" value="TPR-like_helical_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019734">
        <property type="entry name" value="TPR_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR039691">
        <property type="entry name" value="ZC3H7A/B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036236">
        <property type="entry name" value="Znf_C2H2_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000571">
        <property type="entry name" value="Znf_CCCH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036855">
        <property type="entry name" value="Znf_CCCH_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14928">
        <property type="entry name" value="MICRO-RNA BINDING ZINC FINGER CCCH DOMAIN-CONTAINING PROTEIN 7"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR14928:SF13">
        <property type="entry name" value="ZINC FINGER CCCH DOMAIN-CONTAINING PROTEIN 7A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00642">
        <property type="entry name" value="zf-CCCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00028">
        <property type="entry name" value="TPR"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00356">
        <property type="entry name" value="ZnF_C3H1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57667">
        <property type="entry name" value="beta-beta-alpha zinc fingers"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90229">
        <property type="entry name" value="CCCH zinc finger"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48452">
        <property type="entry name" value="TPR-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50103">
        <property type="entry name" value="ZF_C3H1"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1 4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 4" id="KW-0862">Zinc</keyword>
    <keyword evidence="2 4" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C3H1-type" evidence="5">
        <location>
            <begin position="593"/>
            <end position="615"/>
        </location>
    </feature>
    <feature type="domain" description="C3H1-type" evidence="5">
        <location>
            <begin position="729"/>
            <end position="757"/>
        </location>
    </feature>
    <feature type="domain" description="C3H1-type" evidence="5">
        <location>
            <begin position="865"/>
            <end position="887"/>
        </location>
    </feature>
    <feature type="zinc finger region" description="C3H1-type" evidence="4">
        <location>
            <begin position="593"/>
            <end position="615"/>
        </location>
    </feature>
    <feature type="zinc finger region" description="C3H1-type" evidence="4">
        <location>
            <begin position="729"/>
            <end position="757"/>
        </location>
    </feature>
    <feature type="zinc finger region" description="C3H1-type" evidence="4">
        <location>
            <begin position="865"/>
            <end position="887"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="PROSITE" id="PS50103"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000017766.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="930" mass="105856" checksum="55A56662FFE77CB8" modified="2019-05-08" version="1">MSSVPEVRSIRQTDVLMGLVRNLFNEGNDAYREGAWGRSLSHYTEALNISDYANSEEIHVSNEILEKLHVNRIACYSNLRLHDKVLEDCEIVLRLNENNFRALYRKSKALNELGRYKKAYDVVAKCSLAVPQDESVIKLTQELAQKLGLKIRKAYVRAKVRKIFILFNLILKEKQETLSTVSSANFVPEMSSDLASIPVQSISSIMPLQVEKSSLPSVVLANGGNVSFSMPEACLDCGDGDDDDISIGEELDELLDSVPDSNENQTTVVRGPIPSASVAPNIAFSTTLLGTLPVGAGFVPAASFSEMYSQPLASSLENFCSSLNTFPISDSKRDLSRDGTPTLTSNNSSLLLINGPNTFFGTEDYMGIAGQTRNDCSHIFSSGTANIAASLVGRNPLEGTHELRQACQLCFVKTGPKLLDFTYHPNLEHKCKKDILIGKIKNSEDKSWKKIRPRPTKTQYVGPYYICKDVAAEEECRYPGHCTFAYCQEEIDVWTLERKGAFNREAVFGGNGNINFTVSQLLQEHHGLFMFLCEKCFDHKPRIISKRNKDNSSCSHPAMHDFEDNKCLVHILRETTVKYSKIRSFHDQCQLDLCRHEVRYGCLREDECFYAHSLVELKVWIMQNETGISHDTIAQESKKYWQNMEANAHGAQILGNQMKHGSLNLKMKFVCAQCWRHGQVSEPDKNKKYCSAKKAGHPWTKDRRVVLVMSNERKKWMTIRPLPTKKQVPSQFDLCNHIASGKKCQYVGNCSFAHSPEEREMWTYMKENSIQDMEQLYEMWLKSQKPEKEDTATQANKENGKQIHMPTDYAEVTVDFHCWMCGKNCNSEKQWQGHISSEKHKEKVFHTEDDQNCWQHRFPTGYFNICDRYMAGTCIEGNSCKFAHGNAELREWEERRDVLRMKLSKARKDHLIAPNDNDFGKYSFLFKDLN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HRT1</accession>
    <name>A0A452HRT1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Sulfotransferase</fullName>
            <ecNumber evidence="5">2.8.2.-</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 5">Belongs to the sulfotransferase 1 family.</text>
    </comment>
    <dbReference type="EC" id="2.8.2.-" evidence="5"/>
    <dbReference type="AlphaFoldDB" id="A0A452HRT1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000020284.1">
        <property type="protein sequence ID" value="ENSGAGP00000017783.1"/>
        <property type="gene ID" value="ENSGAGG00000013102.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008146">
        <property type="term" value="F:sulfotransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000433">
        <property type="entry name" value="Estrogen sulfotransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000863">
        <property type="entry name" value="Sulfotransferase_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11783">
        <property type="entry name" value="SULFOTRANSFERASE SULT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00685">
        <property type="entry name" value="Sulfotransfer_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 5" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Sulfotransferase" evidence="7">
        <location>
            <begin position="71"/>
            <end position="304"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="310"/>
            <end position="347"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="310"/>
            <end position="320"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="RuleBase" id="RU361155"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="Pfam" id="PF00685"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000017783.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="378" mass="42193" checksum="E9B9E4E48DA78821" modified="2019-05-08" version="1">MFISLCCETQGIWVLGSPGKVWGDQSAPKHSNFWGSGSFTRSKLVTKLGGFHADPHILDVVSGPLTLSSLGTNWMLEILSLIRSDGDPGWVCSVLNWDRVPWVEYGPGLEAARKYPPPRLLCFHLPVQLFPKSLQRSKAKIIYTLRCPKDVVVSLYHFSRIVRLFKDPDSLDCFLEDFLSGNVLYGSWFHHVTGWMGLKANKNIFSITYEELQQDLQGSVRRICHFLGKELSEEQVAAVVENVSFQSMKGNKISNFSQMSDKYMDHQKGEFLRKGICGDWRNHLSEAQSQRFDTVFREWMQGLGSRWQFSDQSPHGSSLPGSPCSMFPAASPNRGSPSPAPQASGRPSLLPAQAGFLLFAHTVRLEGDALAPAWPMAG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="25" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HRT5</accession>
    <name>A0A452HRT5_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">MADS-box domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HRT5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000020289.1">
        <property type="protein sequence ID" value="ENSGAGP00000017788.1"/>
        <property type="gene ID" value="ENSGAGG00000013199.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042826">
        <property type="term" value="F:histone deacetylase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046983">
        <property type="term" value="F:protein dimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007507">
        <property type="term" value="P:heart development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045944">
        <property type="term" value="P:positive regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00265">
        <property type="entry name" value="MADS_MEF2_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.1810.10:FF:000001">
        <property type="entry name" value="Myocyte-specific enhancer factor 2A homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.1810.10">
        <property type="entry name" value="Transcription factor, MADS-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022102">
        <property type="entry name" value="HJURP_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033896">
        <property type="entry name" value="MEF2-like_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002100">
        <property type="entry name" value="TF_MADSbox"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036879">
        <property type="entry name" value="TF_MADSbox_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11945">
        <property type="entry name" value="MADS BOX PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11945:SF837">
        <property type="entry name" value="MYOCYTE ENHANCER FACTOR 2D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12347">
        <property type="entry name" value="HJURP_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00319">
        <property type="entry name" value="SRF-TF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00404">
        <property type="entry name" value="MADSDOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00432">
        <property type="entry name" value="MADS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55455">
        <property type="entry name" value="SRF-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00350">
        <property type="entry name" value="MADS_BOX_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50066">
        <property type="entry name" value="MADS_BOX_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0010">Activator</keyword>
    <keyword evidence="7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="6" id="KW-0539">Nucleus</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0804">Transcription</keyword>
    <keyword evidence="2" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="MADS-box" evidence="9">
        <location>
            <begin position="1"/>
            <end position="61"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="176"/>
            <end position="207"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="401"/>
            <end position="501"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="354"/>
            <end position="382"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="180"/>
            <end position="192"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="401"/>
            <end position="414"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="427"/>
            <end position="436"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="449"/>
            <end position="459"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50066"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000017788.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="501" mass="54460" checksum="83196433EC359198" modified="2019-05-08" version="1">MGRKKIQIQRITDERNRQVTFTKRKFGLMKKAYELSVLCDCEIALIIFNHSNKLFQYASTDMDKVLLKYTEYNEPHESRTNADIIETLRKKGFNGCDSPEPDGDDSIDQSPLMEDKYRKASEDLDILFKRYGSAVPAPNFAMPVTVPVTNQNTLQFSNPGGSLVTQSLVTSSLTDPRLLSPQQPALQRNTVSPGLPQRPASAGAMLGGDLTNTNGACPSPVGNGYVSARASPGLLPVSNGNTMGKIIPAKSPPPPSHSTQLAANSRKPDLRVITSQGAKGLMHHLQNAQRLGVSQATHSLTTPVVSVATPSLLTQGLPFSAMPTAYNTDYQLTSAELSSLPAFSSPSGLSLGNISAWQQQHQQQQQQQQQQQQQQQQQQQQQHLVPVSLSNLIQGGHLSHTTTLTVNTNPNISIKSEPVSPNRERNTATPLSSFPHQSRHEPTGRSPVDSLSSNASSYEGNERDDPARADFGSSLGLLRPTTEAEGESPSVKRVRLDAWVT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HRT6</accession>
    <name>A0A452HRT6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Kynurenine--oxoglutarate transaminase 3</fullName>
            <ecNumber evidence="6">2.6.1.63</ecNumber>
            <ecNumber evidence="5">2.6.1.7</ecNumber>
            <ecNumber evidence="4">4.4.1.13</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="17">Cysteine-S-conjugate beta-lyase 2</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="13">Kynurenine aminotransferase 3</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="14">Kynurenine aminotransferase III</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="15">Kynurenine--glyoxylate transaminase</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="16">Kynurenine--oxoglutarate transaminase III</fullName>
        </alternativeName>
    </protein>
    <organism evidence="24 25">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="25" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="24" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="24" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="22">Catalyzes the irreversible transamination of the L-tryptophan metabolite L-kynurenine to form kynurenic acid (KA), an intermediate in the tryptophan catabolic pathway which is also a broad spectrum antagonist of the three ionotropic excitatory amino acid receptors among others. May catalyze the beta-elimination of S-conjugates and Se-conjugates of L-(seleno)cysteine, resulting in the cleavage of the C-S or C-Se bond. Has transaminase activity towards L-kynurenine, tryptophan, phenylalanine, serine, cysteine, methionine, histidine, glutamine and asparagine with glyoxylate as an amino group acceptor (in vitro). Has lower activity with 2-oxoglutarate as amino group acceptor (in vitro).</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="20">
            <text>3-hydroxy-L-kynurenine + glyoxylate = xanthurenate + glycine + H2O</text>
            <dbReference type="Rhea" id="RHEA:65900"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:36655"/>
            <dbReference type="ChEBI" id="CHEBI:57305"/>
            <dbReference type="ChEBI" id="CHEBI:58125"/>
            <dbReference type="ChEBI" id="CHEBI:71201"/>
            <dbReference type="EC" id="2.6.1.63"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="20">
            <dbReference type="Rhea" id="RHEA:65901"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>L-kynurenine + 2-oxoglutarate = kynurenate + L-glutamate + H2O</text>
            <dbReference type="Rhea" id="RHEA:65560"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:16810"/>
            <dbReference type="ChEBI" id="CHEBI:29985"/>
            <dbReference type="ChEBI" id="CHEBI:57959"/>
            <dbReference type="ChEBI" id="CHEBI:58454"/>
            <dbReference type="EC" id="2.6.1.7"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="18">
            <dbReference type="Rhea" id="RHEA:65561"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="19">
            <text>L-kynurenine + glyoxylate = kynurenate + glycine + H2O</text>
            <dbReference type="Rhea" id="RHEA:65896"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:36655"/>
            <dbReference type="ChEBI" id="CHEBI:57305"/>
            <dbReference type="ChEBI" id="CHEBI:57959"/>
            <dbReference type="ChEBI" id="CHEBI:58454"/>
            <dbReference type="EC" id="2.6.1.63"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="19">
            <dbReference type="Rhea" id="RHEA:65897"/>
        </physiologicalReaction>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="21">
            <text>an S-substituted L-cysteine + H2O = a thiol + pyruvate + NH4(+)</text>
            <dbReference type="Rhea" id="RHEA:18121"/>
            <dbReference type="ChEBI" id="CHEBI:15361"/>
            <dbReference type="ChEBI" id="CHEBI:15377"/>
            <dbReference type="ChEBI" id="CHEBI:28938"/>
            <dbReference type="ChEBI" id="CHEBI:29256"/>
            <dbReference type="ChEBI" id="CHEBI:58717"/>
            <dbReference type="EC" id="4.4.1.13"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="21">
            <dbReference type="Rhea" id="RHEA:18122"/>
        </physiologicalReaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>pyridoxal 5'-phosphate</name>
            <dbReference type="ChEBI" id="CHEBI:597326"/>
        </cofactor>
    </comment>
    <comment type="pathway">
        <text evidence="12">Amino-acid degradation; L-kynurenine degradation; kynurenate from L-kynurenine: step 1/2.</text>
    </comment>
    <comment type="subunit">
        <text evidence="3">Homodimer.</text>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the class-I pyridoxal-phosphate-dependent aminotransferase family.</text>
    </comment>
    <dbReference type="EC" id="2.6.1.63" evidence="6"/>
    <dbReference type="EC" id="2.6.1.7" evidence="5"/>
    <dbReference type="EC" id="4.4.1.13" evidence="4"/>
    <dbReference type="AlphaFoldDB" id="A0A452HRT6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000020276.1">
        <property type="protein sequence ID" value="ENSGAGP00000017776.1"/>
        <property type="gene ID" value="ENSGAGG00000013198.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00334">
        <property type="reaction ID" value="UER00726"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005739">
        <property type="term" value="C:mitochondrion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0047804">
        <property type="term" value="F:cysteine-S-conjugate beta-lyase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0047315">
        <property type="term" value="F:kynurenine-glyoxylate transaminase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016212">
        <property type="term" value="F:kynurenine-oxoglutarate transaminase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030170">
        <property type="term" value="F:pyridoxal phosphate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0097053">
        <property type="term" value="P:L-kynurenine catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="CDD" id="cd00609">
        <property type="entry name" value="AAT_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.640.10:FF:000024">
        <property type="entry name" value="Kynurenine--oxoglutarate transaminase 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.90.1150.10:FF:000021">
        <property type="entry name" value="Kynurenine--oxoglutarate transaminase 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.90.1150.10">
        <property type="entry name" value="Aspartate Aminotransferase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.640.10">
        <property type="entry name" value="Type I PLP-dependent aspartate aminotransferase-like (Major domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004839">
        <property type="entry name" value="Aminotransferase_I/II_large"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051326">
        <property type="entry name" value="Kynurenine-oxoglutarate_AT"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015424">
        <property type="entry name" value="PyrdxlP-dep_Trfase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015421">
        <property type="entry name" value="PyrdxlP-dep_Trfase_major"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015422">
        <property type="entry name" value="PyrdxlP-dep_Trfase_small"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43807">
        <property type="entry name" value="FI04487P"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43807:SF6">
        <property type="entry name" value="KYNURENINE--OXOGLUTARATE TRANSAMINASE 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00155">
        <property type="entry name" value="Aminotran_1_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53383">
        <property type="entry name" value="PLP-dependent transferases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0032">Aminotransferase</keyword>
    <keyword evidence="11" id="KW-0456">Lyase</keyword>
    <keyword evidence="10" id="KW-0663">Pyridoxal phosphate</keyword>
    <keyword evidence="25" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="9" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Aminotransferase class I/classII large" evidence="23">
        <location>
            <begin position="31"/>
            <end position="400"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001933"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00007441"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00011738"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00012224"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012751"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00013010"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00019100"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022576"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022898"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023239"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00024016"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00029778"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00030993"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00031198"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00031371"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00031600"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00047478"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00047677"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00047888"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00049325"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="ARBA" id="ARBA00054518"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="Pfam" id="PF00155"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="24">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000017776.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="25">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="420" mass="47330" checksum="E0D12FCC77EB8430" modified="2019-05-08" version="1">MAFKHKNAKRIEGLDVNVWVEFTKVAADSSIVNLGQGLPDISPPSYVKEELAKAAKVDRLNQYTRGFGHPLLVKALSQVYEKVCGRKIDPYTDILVTVGGYGSLFSAIQGLIEEGDEVIIIEPFFDCYEPMVKMAGGKPVFIPLRYKTVVGNSASSADWVLDPTELASKFNSRTKAIILNTPHNPIGKVFTKEELQVIADLCIKHDTLCLSDEVYEWLVYKGSKHIKIATLPGMWERTITIGSAGKTYSVTGWKLGWSIGPQHLIKHLQVVQQNTLYTCPTPLQEALAQALWIDFKRMDDPDCYFYLLSRELEGKRNRMAQLLQEVGLKPVIPDGGYFMIVDVSALNVDLSDMEANQHYDYKFVKWMIKSKKLSAIPLTAFCGPETKKQFEKYIRFCFIKQDSTLDAAEVILKNWNKPTS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HRT7</accession>
    <name>A0A452HRT7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="9">MADS-box domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HRT7"/>
    <dbReference type="Ensembl" id="ENSGAGT00000020277.1">
        <property type="protein sequence ID" value="ENSGAGP00000017777.1"/>
        <property type="gene ID" value="ENSGAGG00000013199.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
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        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
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    <dbReference type="GO" id="GO:0000981">
        <property type="term" value="F:DNA-binding transcription factor activity, RNA polymerase II-specific"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042826">
        <property type="term" value="F:histone deacetylase binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046983">
        <property type="term" value="F:protein dimerization activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000978">
        <property type="term" value="F:RNA polymerase II cis-regulatory region sequence-specific DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007507">
        <property type="term" value="P:heart development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045944">
        <property type="term" value="P:positive regulation of transcription by RNA polymerase II"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
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        <property type="entry name" value="MADS_MEF2_like"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Myocyte-specific enhancer factor 2A homolog"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Transcription factor, MADS-box"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="HJURP_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033896">
        <property type="entry name" value="MEF2-like_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002100">
        <property type="entry name" value="TF_MADSbox"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036879">
        <property type="entry name" value="TF_MADSbox_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11945">
        <property type="entry name" value="MADS BOX PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11945:SF837">
        <property type="entry name" value="MYOCYTE ENHANCER FACTOR 2D"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12347">
        <property type="entry name" value="HJURP_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00319">
        <property type="entry name" value="SRF-TF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00404">
        <property type="entry name" value="MADSDOMAIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00432">
        <property type="entry name" value="MADS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55455">
        <property type="entry name" value="SRF-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00350">
        <property type="entry name" value="MADS_BOX_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50066">
        <property type="entry name" value="MADS_BOX_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-0010">Activator</keyword>
    <keyword evidence="7" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="3" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="6" id="KW-0539">Nucleus</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0804">Transcription</keyword>
    <keyword evidence="2" id="KW-0805">Transcription regulation</keyword>
    <feature type="domain" description="MADS-box" evidence="9">
        <location>
            <begin position="1"/>
            <end position="61"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="176"/>
            <end position="207"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="8">
        <location>
            <begin position="400"/>
            <end position="505"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="7">
        <location>
            <begin position="353"/>
            <end position="381"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="180"/>
            <end position="192"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="400"/>
            <end position="413"/>
        </location>
    </feature>
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        <location>
            <begin position="426"/>
            <end position="435"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="8">
        <location>
            <begin position="448"/>
            <end position="458"/>
        </location>
    </feature>
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        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023015"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00023159"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023163"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS50066"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000017777.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="505" mass="54992" checksum="95165F0E81C579FF" modified="2019-05-08" version="1">MGRKKIQIQRITDERNRQVTFTKRKFGLMKKAYELSVLCDCEIALIIFNHSNKLFQYASTDMDKVLLKYTEYNEPHESRTNADIIETLRKKGFNGCDSPEPDGDDSIDQSPLMEDKYRKASEDLDILFKRYGSAVPAPNFAMPVTVPVTNQNTLQFSNPGGSLVTQSLVTSSLTDPRLLSPQQPALQRNTVSPGLPQRPASAGAMLGGDLTNTNGACPSPVGNGYVSARASPGLLPVSNGNTMGKIIPAKSPPPPSHSTQLAANSRKPDLRVITSQGAKGLMHHLNAQRLGVSQATHSLTTPVVSVATPSLLTQGLPFSAMPTAYNTDYQLTSAELSSLPAFSSPSGLSLGNISAWQQQHQQQQQQQQQQQQQQQQQQQQQHLVPVSLSNLIQGGHLSHTTTLTVNTNPNISIKSEPVSPNRERNTATPLSSFPHQSRHEPTGRSPVDSLSSNASSYEGNERDDPARADFGSSLGLLRPTTEAEGESPSVKRVRLDAWHPQAQWL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HRT9</accession>
    <name>A0A452HRT9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="13">Thymidine kinase</fullName>
            <ecNumber evidence="13">2.7.1.21</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="10">
            <text>thymidine + ATP = dTMP + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:19129"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:17748"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:63528"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.1.21"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="10">
            <dbReference type="Rhea" id="RHEA:19130"/>
        </physiologicalReaction>
    </comment>
    <comment type="subunit">
        <text evidence="9">Homotetramer. Tetramerization from dimerization is induced by ATP and increases catalytic efficiency due to a high affinity for thymidine. Tetramerization is inhibited by phosphorylation at Ser-13. Interacts (via the KEN box) with FZR1.</text>
    </comment>
    <comment type="similarity">
        <text evidence="1 14">Belongs to the thymidine kinase family.</text>
    </comment>
    <dbReference type="EC" id="2.7.1.21" evidence="13"/>
    <dbReference type="AlphaFoldDB" id="A0A452HRT9"/>
    <dbReference type="Ensembl" id="ENSGAGT00000020226.1">
        <property type="protein sequence ID" value="ENSGAGP00000017729.1"/>
        <property type="gene ID" value="ENSGAGG00000013169.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004797">
        <property type="term" value="F:thymidine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0071897">
        <property type="term" value="P:DNA biosynthetic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046104">
        <property type="term" value="P:thymidine metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.60.20:FF:000028">
        <property type="entry name" value="Thymidine kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000761">
        <property type="entry name" value="Thymidine kinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.60.20">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001267">
        <property type="entry name" value="Thymidine_kinase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020633">
        <property type="entry name" value="Thymidine_kinase_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11441">
        <property type="entry name" value="THYMIDINE KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11441:SF0">
        <property type="entry name" value="THYMIDINE KINASE, CYTOSOLIC"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00265">
        <property type="entry name" value="TK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF035805">
        <property type="entry name" value="TK_cell"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57716">
        <property type="entry name" value="Glucocorticoid receptor-like (DNA-binding domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00603">
        <property type="entry name" value="TK_CELLULAR_TYPE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 13" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="2 13" id="KW-0237">DNA synthesis</keyword>
    <keyword evidence="6 13" id="KW-0418">Kinase</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="5 13" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3 13" id="KW-0808">Transferase</keyword>
    <keyword evidence="7" id="KW-0862">Zinc</keyword>
    <feature type="active site" description="Proton acceptor" evidence="11">
        <location>
            <position position="63"/>
        </location>
    </feature>
    <feature type="binding site" evidence="12">
        <location>
            <position position="146"/>
        </location>
        <ligand>
            <name>substrate</name>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00007587"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022634"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00046642"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00048113"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="PIRSR" id="PIRSR035805-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="PIRSR" id="PIRSR035805-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="RuleBase" id="RU000544"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="RuleBase" id="RU004165"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000017729.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="189" mass="21062" checksum="B92519DD94C5AE0A" modified="2019-05-08" version="1">MRRVRRFQIAQYKCLVIKYAKDSRYCTNGVSTHDRNTMEAFSACYLKDVQQEALGSAVIGIDEGQFFPDIVEFCEAMANAGKTVIVAALDGTFQRKAFGSILNLVPLAESVVKLNAVCMECYREASYTKRLGAEREVEVIGGSDKYHSVCRVCYFRKRPQQAGPENKENLPVGAKQVQTLGCSGSRRPQ</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="28" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HS21</accession>
    <name>A0A452HS21_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="1">[histone H3]-lysine(4) N-trimethyltransferase</fullName>
            <ecNumber evidence="1">2.1.1.354</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
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        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
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                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="7">
            <text>L-lysyl(4)-[histone H3] + 3 S-adenosyl-L-methionine = N(6),N(6),N(6)-trimethyl-L-lysyl(4)-[histone H3] + 3 S-adenosyl-L-homocysteine + 3 H(+)</text>
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            <dbReference type="ChEBI" id="CHEBI:61961"/>
            <dbReference type="EC" id="2.1.1.354"/>
        </reaction>
    </comment>
    <dbReference type="EC" id="2.1.1.354" evidence="1"/>
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    </evidence>
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        <source>
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    </evidence>
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        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="138" mass="16014" checksum="A5A14D48522FABCC" modified="2019-05-08" version="1">LIQDISYRNERLLRCSQCKIARYCDTRCQKEAWQEHKRECKCLKSIEPNFPPDSVRLVGRIVFKLLGQSAACPSEKLYSFSDLQSNIEELSEEMKEGLKHLAQTLQLYLRVEIQDAFQLLPAIDIFQIFAKVSVKTDS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HS41</accession>
    <name>A0A452HS41_9SAUR</name>
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        <recommendedName>
            <fullName evidence="4">Tripeptidyl-peptidase 2</fullName>
            <ecNumber evidence="3">3.4.14.10</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="9">Tripeptidyl aminopeptidase</fullName>
        </alternativeName>
    </protein>
    <organism evidence="17 18">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="18" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="17" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="17" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>Release of an N-terminal tripeptide from a polypeptide.</text>
            <dbReference type="EC" id="3.4.14.10"/>
        </reaction>
    </comment>
    <comment type="similarity">
        <text evidence="2 10">Belongs to the peptidase S8 family.</text>
    </comment>
    <dbReference type="EC" id="3.4.14.10" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452HS41"/>
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        <property type="protein sequence ID" value="ENSGAGP00000017883.1"/>
        <property type="gene ID" value="ENSGAGG00000011247.1"/>
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        <property type="project" value="TreeGrafter"/>
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    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
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        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008240">
        <property type="term" value="F:tripeptidyl-peptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
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        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
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    </dbReference>
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        <property type="entry name" value="Peptidase S8/S53 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Peptidase_S8/S53_dom"/>
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        <property type="entry name" value="Peptidase_S8/S53_dom_sf"/>
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        <property type="entry name" value="Peptidase_S8_His-AS"/>
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        <property type="entry name" value="Peptidase_S8_subtilisin-rel"/>
    </dbReference>
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        <property type="entry name" value="TPP_II_domain"/>
    </dbReference>
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        <property type="entry name" value="TPPII_C_art"/>
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        <property type="entry name" value="TPPII_GBD"/>
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        <property type="entry name" value="TPPII_Ig-like-1"/>
    </dbReference>
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    </dbReference>
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        <property type="entry name" value="TPPII_Ig-like_sf"/>
    </dbReference>
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        <property type="entry name" value="PEPTIDASE S8"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="TRIPEPTIDYL-PEPTIDASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Peptidase_S8"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="TPPII"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="TPPII_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21316">
        <property type="entry name" value="TPPII_GBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21223">
        <property type="entry name" value="TPPII_Ig-like-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00723">
        <property type="entry name" value="SUBTILISIN"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52743">
        <property type="entry name" value="Subtilisin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51892">
        <property type="entry name" value="SUBTILASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00137">
        <property type="entry name" value="SUBTILASE_HIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00138">
        <property type="entry name" value="SUBTILASE_SER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0031">Aminopeptidase</keyword>
    <keyword evidence="7 10" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="6 10" id="KW-0645">Protease</keyword>
    <keyword evidence="18" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8 10" id="KW-0720">Serine protease</keyword>
    <feature type="domain" description="Peptidase S8/S53" evidence="12">
        <location>
            <begin position="34"/>
            <end position="506"/>
        </location>
    </feature>
    <feature type="domain" description="Tripeptidyl-peptidase II first Ig-like" evidence="15">
        <location>
            <begin position="527"/>
            <end position="640"/>
        </location>
    </feature>
    <feature type="domain" description="Tripeptidyl-peptidase II galactose-binding" evidence="16">
        <location>
            <begin position="660"/>
            <end position="747"/>
        </location>
    </feature>
    <feature type="domain" description="Tripeptidyl peptidase II second Ig-like" evidence="13">
        <location>
            <begin position="783"/>
            <end position="969"/>
        </location>
    </feature>
    <feature type="domain" description="Tripeptidyl peptidase II C-terminal" evidence="14">
        <location>
            <begin position="1016"/>
            <end position="1075"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="11">
        <location>
            <begin position="1003"/>
            <end position="1023"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="11">
        <location>
            <begin position="1009"/>
            <end position="1023"/>
        </location>
    </feature>
    <feature type="active site" description="Charge relay system" evidence="10">
        <location>
            <position position="43"/>
        </location>
    </feature>
    <feature type="active site" description="Charge relay system" evidence="10">
        <location>
            <position position="270"/>
        </location>
    </feature>
    <feature type="active site" description="Charge relay system" evidence="10">
        <location>
            <position position="455"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
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        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00011073"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012462"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00020244"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022438"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00032232"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="12">
        <source>
            <dbReference type="Pfam" id="PF00082"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="Pfam" id="PF12580"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="Pfam" id="PF12583"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="15">
        <source>
            <dbReference type="Pfam" id="PF21223"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="16">
        <source>
            <dbReference type="Pfam" id="PF21316"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="17">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000017883.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1139" mass="126619" checksum="1D5B56032E50A674" modified="2025-10-08" version="2">MAGWTHEPFPFHGLLPKKETGAASFLGRYPEYDGRGVLLAVLDTGVDPGAPGMQITTDGKPKIIDIIDTTGSGDVNTSTIVEPKDGEIVGLSGRTLKIPTNWVNPSGKYHIGIKNAYDFYPKALKERIQKERKEKQWDPVHRLLLAEACRKLDEFDAVHNSPSQHLSCFKFTTKLMKEEFQSQVELLNSLEKKYNDPGPVYDCLVWHDGETWRACIDSSECGDFTKCTVLGTYRERQEYGSFGISEMLNYSVNIYDEGNLLSIVTSGGAHGTHVASIAAGYFPEEPERNGVAPGAQILAIKIGDTRLSTMETGTGLIRAMIEAMKYKCDLVNYSYGEATHWPNSGRICEVINEAVWKHNVIYVSSAGNNGPCLSTVGCPGGTTSSVIGVGAYVSPDMMVAEYSLREKLPANQYTWSSRGPSTDGALGVSISAPGGAIASVPNWTLRGTQLMNGTSMSSPNACGGIALVLSGLKANDVHYTVHSVRRALENTAVKAENIEVFAQGHGIIQVDKAYDYLIQNSSFTSNVGFTVTVGNNRGIYLRDPVQIAAPSDRGVGIEPVFPENTENTKRISLQLHLALTSNAAWVQCPSHLELMNQCRHINVRVDPRGLREGLHYTEVCGYDIAAPNAGPLFRVPITLIIPARIDESSHYDLSYTDVHFKPGQIRRHFIEVPQGATWAEVTVCSCSTDVTAKFVLHAVQLVKQKAYRSHEFYKFSSLPEKGSVTEAFPVLGGKTIEFCIARWWASLNDVSIDYTISFHGIVCATPQLNIHASEGIVRFDVQSTLKYEDIAPCINLKSWVQTLRPVSTKIKPLGSRDVLPNNRQLYEMILTYSFHQPKSGEVTPSCPLLCELLYESEFDSQLWIIFDQNKRQMGSGDAYPHQYSLKLEKGDYTIRLQIRHEQNSELDRIKDLPFVVSHRLSNTLSLDIYENHSLALLGKKKSNSLTLPPKHSHPFFVTSLPDDKIPKGAGPGCYLVGALTLSKTELGKKADVITVHYHLIPSPTKTKNGSKEKEREPEKEKDVKEEFAEALRDLKIQWMTKLDTSDVFNELKEAFPNHLPLYVARLHQLDSEKERLKRLDEIIEAANTVISHIDQTALAVYFVMKTDPRPDAATIKKYLTMIWKNRMMLKVKMTIKRAA</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HS44</accession>
    <name>A0A452HS44_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">BTB domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="22 23">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="23" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="22" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="22" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="18">
            <text>K(+)(in) = K(+)(out)</text>
            <dbReference type="Rhea" id="RHEA:29463"/>
            <dbReference type="ChEBI" id="CHEBI:29103"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cell membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="17">Presynaptic cell membrane</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452HS44"/>
    <dbReference type="Ensembl" id="ENSGAGT00000020384.1">
        <property type="protein sequence ID" value="ENSGAGP00000017867.1"/>
        <property type="gene ID" value="ENSGAGG00000013255.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043679">
        <property type="term" value="C:axon terminus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032590">
        <property type="term" value="C:dendrite membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032809">
        <property type="term" value="C:neuronal cell body membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045211">
        <property type="term" value="C:postsynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042734">
        <property type="term" value="C:presynaptic membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008076">
        <property type="term" value="C:voltage-gated potassium channel complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005251">
        <property type="term" value="F:delayed rectifier potassium channel activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0001508">
        <property type="term" value="P:action potential"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051260">
        <property type="term" value="P:protein homooligomerization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.287.70:FF:000011">
        <property type="entry name" value="Potassium channel, voltage-gated Shaw-related subfamily C, member 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.350:FF:000014">
        <property type="entry name" value="Potassium channel, voltage-gated Shaw-related subfamily C, member 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.710.10:FF:000002">
        <property type="entry name" value="Potassium voltage-gated channel subfamily C member 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.287.70">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.710.10">
        <property type="entry name" value="Potassium Channel Kv1.1, Chain A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.350">
        <property type="entry name" value="Voltage-gated potassium channels. Chain C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000210">
        <property type="entry name" value="BTB/POZ_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005821">
        <property type="entry name" value="Ion_trans_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003968">
        <property type="entry name" value="K_chnl_volt-dep_Kv"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003974">
        <property type="entry name" value="K_chnl_volt-dep_Kv3"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005405">
        <property type="entry name" value="K_chnl_volt-dep_Kv3.4"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR021645">
        <property type="entry name" value="Shal-type_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011333">
        <property type="entry name" value="SKP1/BTB/POZ_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003131">
        <property type="entry name" value="T1-type_BTB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR028325">
        <property type="entry name" value="VG_K_chnl"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027359">
        <property type="entry name" value="Volt_channel_dom_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11537:SF126">
        <property type="entry name" value="POTASSIUM VOLTAGE-GATED CHANNEL SUBFAMILY C MEMBER 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11537">
        <property type="entry name" value="VOLTAGE-GATED POTASSIUM CHANNEL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02214">
        <property type="entry name" value="BTB_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00520">
        <property type="entry name" value="Ion_trans"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF11601">
        <property type="entry name" value="Shal-type"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00169">
        <property type="entry name" value="KCHANNEL"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01583">
        <property type="entry name" value="KV34CHANNEL"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01491">
        <property type="entry name" value="KVCHANNEL"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01498">
        <property type="entry name" value="SHAWCHANNEL"/>
    </dbReference>
    <dbReference type="SMART" id="SM00225">
        <property type="entry name" value="BTB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54695">
        <property type="entry name" value="POZ domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81324">
        <property type="entry name" value="Voltage-gated potassium channels"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="15" id="KW-0966">Cell projection</keyword>
    <keyword evidence="14" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="16" id="KW-0407">Ion channel</keyword>
    <keyword evidence="12" id="KW-0406">Ion transport</keyword>
    <keyword evidence="13 20" id="KW-0472">Membrane</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="9" id="KW-0630">Potassium</keyword>
    <keyword evidence="7" id="KW-0631">Potassium channel</keyword>
    <keyword evidence="4" id="KW-0633">Potassium transport</keyword>
    <keyword evidence="23" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="11" id="KW-0770">Synapse</keyword>
    <keyword evidence="6 20" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="10 20" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="2" id="KW-0813">Transport</keyword>
    <keyword evidence="8" id="KW-0851">Voltage-gated channel</keyword>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="213"/>
            <end position="232"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="269"/>
            <end position="290"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="302"/>
            <end position="323"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="370"/>
            <end position="391"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="412"/>
            <end position="429"/>
        </location>
    </feature>
    <feature type="transmembrane region" description="Helical" evidence="20">
        <location>
            <begin position="441"/>
            <end position="470"/>
        </location>
    </feature>
    <feature type="domain" description="BTB" evidence="21">
        <location>
            <begin position="36"/>
            <end position="140"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1"/>
            <end position="23"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="479"/>
            <end position="566"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="521"/>
            <end position="531"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="19">
        <location>
            <begin position="555"/>
            <end position="566"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004651"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022448"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022538"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022826"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022882"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022958"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023065"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023303"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00034111"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00034430"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="SMART" id="SM00225"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="22">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000017867.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="23">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="595" mass="66417" checksum="2DEDF1C198B9A8E2" modified="2019-05-08" version="1">MISSVCVSSYRGRKSGNKPPSKTCLKEEMAKGEESDKITINVGGTRHETYKSTLRTLPGTRLAWLADPDAQSNFDFDGKSNEFFFDRHPGIFSYVLNYYRTGKLHCPADICGPLFEEELTYWGIDETDVEPCCWMTYRQHRDAEEALDIFESPEPGAGGEEAEEEGGREMALQRLGMDDRSPGAAGAKGGGGCCRNWQPRMWALFEDPYSSRAARVVAFASLFFILVSITTFCLETHEAFNRNVNVTETMVVGNTTRVLLTNKVETEPVLTYIEGVCVLWFTLEFLVRIICCPDKLLFIKKLLNIIDFVAILPFYLEVGLSGLSSKAARDVLGFLRVVRFVRILRIFKLTRHFVGLRVLGHTLRASTNEFLLLIIFLALGVLIFATMIYYAERIGAKTSDQSGTDQTHFKNIPIGFWWAVVTMTTLGYGDMYPKTWSGMLVGALCALAGVLTIAMPVPVIVNNFGMYYSLAMAKQKLPKKKKKHIPRPAPLDSPTYCKSEDNSPRNSTQSDTCPLAAEEVVVERKRSDSKQNGDANTVLSDEEQPLSSLHEEKRPMRRSSTRDKTKKAATCFLLNASDFSCTADGGIRKGLSNPL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HS48</accession>
    <name>A0A452HS48_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="17">E3 ubiquitin-protein ligase TRIM71</fullName>
            <ecNumber evidence="5">2.3.2.27</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="20">Protein lin-41 homolog</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="18">RING-type E3 ubiquitin transferase TRIM71</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="19">Tripartite motif-containing protein 71</fullName>
        </alternativeName>
    </protein>
    <organism evidence="29 30">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="30" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="29" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="29" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="21">E3 ubiquitin-protein ligase that cooperates with the microRNAs (miRNAs) machinery and promotes embryonic stem cells proliferation and maintenance. Binds to miRNAs and participates in post-transcriptional repression of transcripts. Required to maintain proliferation and prevent premature differentiation of neural progenitor cells during early neural development.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>S-ubiquitinyl-[E2 ubiquitin-conjugating enzyme]-L-cysteine + [acceptor protein]-L-lysine = [E2 ubiquitin-conjugating enzyme]-L-cysteine + N(6)-ubiquitinyl-[acceptor protein]-L-lysine.</text>
            <dbReference type="EC" id="2.3.2.27"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="3">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">P-body</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4">Belongs to the TRIM/RBCC family.</text>
    </comment>
    <dbReference type="EC" id="2.3.2.27" evidence="5"/>
    <dbReference type="AlphaFoldDB" id="A0A452HS48"/>
    <dbReference type="Ensembl" id="ENSGAGT00000020416.1">
        <property type="protein sequence ID" value="ENSGAGP00000017895.1"/>
        <property type="gene ID" value="ENSGAGG00000013271.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000932">
        <property type="term" value="C:P-body"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035198">
        <property type="term" value="F:miRNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061630">
        <property type="term" value="F:ubiquitin protein ligase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0017148">
        <property type="term" value="P:negative regulation of translation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043161">
        <property type="term" value="P:proteasome-mediated ubiquitin-dependent protein catabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000209">
        <property type="term" value="P:protein polyubiquitination"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031047">
        <property type="term" value="P:regulatory ncRNA-mediated gene silencing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd19812">
        <property type="entry name" value="Bbox1_TRIM71_C-VII"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd19796">
        <property type="entry name" value="Bbox2_TRIM71_C-VII"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd14954">
        <property type="entry name" value="NHL_TRIM71_like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd16589">
        <property type="entry name" value="RING-HC_TRIM71_C-VII"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.120.10.30:FF:000013">
        <property type="entry name" value="E3 ubiquitin-protein ligase TRIM71"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.120.10.30:FF:000025">
        <property type="entry name" value="E3 ubiquitin-protein ligase TRIM71"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.120.10.30:FF:000080">
        <property type="entry name" value="E3 ubiquitin-protein ligase TRIM71"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000527">
        <property type="entry name" value="E3 ubiquitin-protein ligase TRIM71"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.40.10:FF:000435">
        <property type="entry name" value="Tripartite motif containing 71, E3 ubiquitin protein ligase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="4.10.830.40">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.160.60">
        <property type="entry name" value="Classic Zinc Finger"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.120.10.30">
        <property type="entry name" value="TolB, C-terminal domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.40.10">
        <property type="entry name" value="Zinc/RING finger domain, C3HC4 (zinc finger)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011042">
        <property type="entry name" value="6-blade_b-propeller_TolB-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017868">
        <property type="entry name" value="Filamin/ABP280_repeat-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001298">
        <property type="entry name" value="Filamin/ABP280_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014756">
        <property type="entry name" value="Ig_E-set"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001258">
        <property type="entry name" value="NHL_repeat"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050952">
        <property type="entry name" value="TRIM-NHL_E3_ligases"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000315">
        <property type="entry name" value="Znf_B-box"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001841">
        <property type="entry name" value="Znf_RING"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013083">
        <property type="entry name" value="Znf_RING/FYVE/PHD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR017907">
        <property type="entry name" value="Znf_RING_CS"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24104">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE NHLRC1-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24104:SF55">
        <property type="entry name" value="E3 UBIQUITIN-PROTEIN LIGASE TRIM71"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00630">
        <property type="entry name" value="Filamin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01436">
        <property type="entry name" value="NHL"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00643">
        <property type="entry name" value="zf-B_box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00336">
        <property type="entry name" value="BBOX"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00557">
        <property type="entry name" value="IG_FLMN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00184">
        <property type="entry name" value="RING"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57845">
        <property type="entry name" value="B-box zinc-binding domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF81296">
        <property type="entry name" value="E set domains"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF101898">
        <property type="entry name" value="NHL repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57850">
        <property type="entry name" value="RING/U-box"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50194">
        <property type="entry name" value="FILAMIN_REPEAT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51125">
        <property type="entry name" value="NHL"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50119">
        <property type="entry name" value="ZF_BBOX"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00518">
        <property type="entry name" value="ZF_RING_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50089">
        <property type="entry name" value="ZF_RING_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="15 25" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="6" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="9" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="30" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="10" id="KW-0677">Repeat</keyword>
    <keyword evidence="14" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="16" id="KW-0943">RNA-mediated gene silencing</keyword>
    <keyword evidence="8" id="KW-0808">Transferase</keyword>
    <keyword evidence="12" id="KW-0833">Ubl conjugation pathway</keyword>
    <keyword evidence="13" id="KW-0862">Zinc</keyword>
    <keyword evidence="11 22" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="RING-type" evidence="27">
        <location>
            <begin position="12"/>
            <end position="90"/>
        </location>
    </feature>
    <feature type="domain" description="B box-type" evidence="28">
        <location>
            <begin position="181"/>
            <end position="228"/>
        </location>
    </feature>
    <feature type="domain" description="B box-type" evidence="28">
        <location>
            <begin position="232"/>
            <end position="260"/>
        </location>
    </feature>
    <feature type="repeat" description="Filamin" evidence="23">
        <location>
            <begin position="425"/>
            <end position="526"/>
        </location>
    </feature>
    <feature type="repeat" description="NHL" evidence="24">
        <location>
            <begin position="539"/>
            <end position="582"/>
        </location>
    </feature>
    <feature type="repeat" description="NHL" evidence="24">
        <location>
            <begin position="586"/>
            <end position="629"/>
        </location>
    </feature>
    <feature type="repeat" description="NHL" evidence="24">
        <location>
            <begin position="635"/>
            <end position="676"/>
        </location>
    </feature>
    <feature type="repeat" description="NHL" evidence="24">
        <location>
            <begin position="680"/>
            <end position="723"/>
        </location>
    </feature>
    <feature type="repeat" description="NHL" evidence="24">
        <location>
            <begin position="727"/>
            <end position="770"/>
        </location>
    </feature>
    <feature type="repeat" description="NHL" evidence="24">
        <location>
            <begin position="774"/>
            <end position="814"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="26">
        <location>
            <begin position="26"/>
            <end position="49"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="26">
        <location>
            <begin position="129"/>
            <end position="187"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="25">
        <location>
            <begin position="282"/>
            <end position="316"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="25">
        <location>
            <begin position="340"/>
            <end position="367"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="26">
        <location>
            <begin position="149"/>
            <end position="163"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000900"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004201"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00008518"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012483"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022884"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023158"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00040205"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00041679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00042007"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00043228"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00059249"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00024"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00087"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="24">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00504"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="25">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="26">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="27">
        <source>
            <dbReference type="PROSITE" id="PS50089"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="28">
        <source>
            <dbReference type="PROSITE" id="PS50119"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="29">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000017895.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="30">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="814" mass="88730" checksum="FA8669A39CE668D9" modified="2019-05-08" version="1">MASFPEADFQLCPLCKEMCGSPAPISSNSSTSSSSSQTSNSSSAGGSSCAPSRRLHVLPCLHAFCRQCLEAQRQHPGAGDALKLRCPICDQKVVISEPSGMDALPSSNFLLSNLLDVVVVAAAAAEEQKNNGRAGGGGGSSNNRHHHGSSPAAASAAAPSPSSSGGGGSASLLLRRPHSRQGEPRCSSCDEGNAASSRCLDCQEHLCDNCVRAHQRVRLTKDHFIERFGGVLHFYCDTCSVPICRECTMGRHVGHSFIYLQDALQDSRTLTIQLLADAQQGRQAIQLSIEQAQAVAEQVEMKAKVMQSEVKAMTSRHKKALEERECELLWKVEKIRQVKAKSLYLQVEKLRQNLNKLDNTISAVQQVLEEGRSMDILLARDRMLAQVQELKNVRGLLQPQEDDRVMFTPPDQALYMAIKSMGFVSSGAFAPLTKATGEGLKRALQGKVASFTVIGYDHDGEPRLSGGDMISAVVMGPDGNLFGADVNDQQNGTYLVSYRPQLEGEHLVSVMMCNQHIENSPFKVMVKSGRSYIGIGLPGLSFGSEGDNDGKLCRPWGVCVDKEGYIMVADRSNNRIQVFKPCGTFHHKFGTLGSRPGQFDRPAGVACDISRRIVVADKDNHRIQIFTFDGQFILKFGEKGTKNGQFNYPWDVAVNAEGKILVSDTRNHRIQLFGPDGVFLNKYGFEGALWKHFDSPRGVTFNHEGHLVVTDFNNHRLLVIHPDCQSARFLGSEGTSNGQFLRPQGVAVDQEGRIIVADSRNHRVQIFESNGSFLCKFGTQGSGFGQMDRPSGIAVTPDGMIVVVDFGNNRILIF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HS49</accession>
    <name>A0A452HS49_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Tripeptidyl-peptidase 2</fullName>
            <ecNumber evidence="5">3.4.14.10</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="15">Tripeptidyl aminopeptidase</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="17">Tripeptidyl-peptidase II</fullName>
        </alternativeName>
    </protein>
    <organism evidence="26 27">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="27" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="26" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="26" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="16">Cytosolic tripeptidyl-peptidase that releases N-terminal tripeptides from polypeptides and is a component of the proteolytic cascade acting downstream of the 26S proteasome in the ubiquitin-proteasome pathway. It plays an important role in intracellular amino acid homeostasis. Stimulates adipogenesis.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>Release of an N-terminal tripeptide from a polypeptide.</text>
            <dbReference type="EC" id="3.4.14.10"/>
        </reaction>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="4 18">Belongs to the peptidase S8 family.</text>
    </comment>
    <dbReference type="EC" id="3.4.14.10" evidence="5"/>
    <dbReference type="Ensembl" id="ENSGAGT00000020389.1">
        <property type="protein sequence ID" value="ENSGAGP00000017872.1"/>
        <property type="gene ID" value="ENSGAGG00000011247.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004177">
        <property type="term" value="F:aminopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008240">
        <property type="term" value="F:tripeptidyl-peptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd04857">
        <property type="entry name" value="Peptidases_S8_Tripeptidyl_Aminopeptidase_II"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.710:FF:000001">
        <property type="entry name" value="Tripeptidyl peptidase 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.20.25.690:FF:000002">
        <property type="entry name" value="Tripeptidyl peptidase 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.3170:FF:000001">
        <property type="entry name" value="Tripeptidyl peptidase 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.200:FF:000003">
        <property type="entry name" value="Tripeptidyl peptidase 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.200:FF:000009">
        <property type="entry name" value="tripeptidyl-peptidase 2 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.710">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.20.25.690">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.3170">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.3080">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.200">
        <property type="entry name" value="Peptidase S8/S53 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000209">
        <property type="entry name" value="Peptidase_S8/S53_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036852">
        <property type="entry name" value="Peptidase_S8/S53_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022398">
        <property type="entry name" value="Peptidase_S8_His-AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR023828">
        <property type="entry name" value="Peptidase_S8_Ser-AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050131">
        <property type="entry name" value="Peptidase_S8_subtilisin-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015500">
        <property type="entry name" value="Peptidase_S8_subtilisin-rel"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034051">
        <property type="entry name" value="TPP_II_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022232">
        <property type="entry name" value="TPPII_C_art"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046939">
        <property type="entry name" value="TPPII_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048384">
        <property type="entry name" value="TPPII_GBD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR048383">
        <property type="entry name" value="TPPII_Ig-like-1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022229">
        <property type="entry name" value="TPPII_Ig-like-2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR046940">
        <property type="entry name" value="TPPII_Ig-like_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43806">
        <property type="entry name" value="PEPTIDASE S8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR43806:SF14">
        <property type="entry name" value="TRIPEPTIDYL-PEPTIDASE 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00082">
        <property type="entry name" value="Peptidase_S8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12580">
        <property type="entry name" value="TPPII"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12583">
        <property type="entry name" value="TPPII_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21316">
        <property type="entry name" value="TPPII_GBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21223">
        <property type="entry name" value="TPPII_Ig-like-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00723">
        <property type="entry name" value="SUBTILISIN"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52743">
        <property type="entry name" value="Subtilisin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51892">
        <property type="entry name" value="SUBTILASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00137">
        <property type="entry name" value="SUBTILASE_HIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00138">
        <property type="entry name" value="SUBTILASE_SER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="13" id="KW-0007">Acetylation</keyword>
    <keyword evidence="7" id="KW-0031">Aminopeptidase</keyword>
    <keyword evidence="8" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="11 18" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="14" id="KW-0539">Nucleus</keyword>
    <keyword evidence="9" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="10 18" id="KW-0645">Protease</keyword>
    <keyword evidence="27" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="12 18" id="KW-0720">Serine protease</keyword>
    <keyword evidence="20" id="KW-0732">Signal</keyword>
    <feature type="signal peptide" evidence="20">
        <location>
            <begin position="1"/>
            <end position="28"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5019479467" description="Tripeptidyl-peptidase 2" evidence="20">
        <location>
            <begin position="29"/>
            <end position="1284"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase S8/S53" evidence="21">
        <location>
            <begin position="57"/>
            <end position="522"/>
        </location>
    </feature>
    <feature type="domain" description="Tripeptidyl-peptidase II first Ig-like" evidence="24">
        <location>
            <begin position="543"/>
            <end position="656"/>
        </location>
    </feature>
    <feature type="domain" description="Tripeptidyl-peptidase II galactose-binding" evidence="25">
        <location>
            <begin position="676"/>
            <end position="763"/>
        </location>
    </feature>
    <feature type="domain" description="Tripeptidyl peptidase II second Ig-like" evidence="22">
        <location>
            <begin position="799"/>
            <end position="985"/>
        </location>
    </feature>
    <feature type="domain" description="Tripeptidyl peptidase II C-terminal" evidence="23">
        <location>
            <begin position="1045"/>
            <end position="1104"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="1030"/>
            <end position="1052"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="1038"/>
            <end position="1052"/>
        </location>
    </feature>
    <feature type="active site" description="Charge relay system" evidence="18">
        <location>
            <position position="66"/>
        </location>
    </feature>
    <feature type="active site" description="Charge relay system" evidence="18">
        <location>
            <position position="286"/>
        </location>
    </feature>
    <feature type="active site" description="Charge relay system" evidence="18">
        <location>
            <position position="471"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001910"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00011073"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00012462"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00020244"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022438"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00032232"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00054482"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00075739"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="SAM" id="SignalP"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="21">
        <source>
            <dbReference type="Pfam" id="PF00082"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="22">
        <source>
            <dbReference type="Pfam" id="PF12580"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="Pfam" id="PF12583"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="Pfam" id="PF21223"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="Pfam" id="PF21316"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000017872.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1284" mass="142347" checksum="93384CAD978CA493" modified="2019-05-08" version="1">WDRDGNRRRCRGLVGPVALSLGVMAAAAEEPFPFHGLLPKKETGAASFLGRYPEYDGRGVLLAVLDTGVDPGAPGMQITTDGKPKIIDIIDTTGSGDVNTSTIVEPKDGEIVGLSGRTLKIPTNWVNPSGKYHIGIKNAYDFYPKALKERIQKERKEKQWDPVHRLLLAEACRKLDEFDAVHNSPSQTTKLMKEEFQSQVELLNSLEKKYNDPGPVYDCLVWHDGETWRACIDSSECGDFTKCTVLGTYRERQEYGSFGISEMLNYSVNIYDEGNLLSIVTSGGAHGTHVASIAAGYFPEEPERNGVAPGAQILAIKIGDTRLSTMETGTGLIRAMIEAMKYKCDLVNYSYGEATHWPNSGRICEVINEAVWKHNVIYVSSAGNNGPCLSTVGCPGGTTSSVIGVGAYVSPDMMVAEYSLREKLPANQYTWSSRGPSTDGALGVSISAPGGAIASVPNWTLRGTQLMNGTSMSSPNACGGIALVLSGLKANDVHYTVHSVRRALENTAVKAENIEVFAQGHGIIQVDKAYDYLIQNSSFTSNVGFTVTVGNNRGIYLRDPVQIAAPSDRGVGIEPVFPENTENTKRISLQLHLALTSNAAWVQCPSHLELMNQCRHINVRVDPRGLREGLHYTEVCGYDIAAPNAGPLFRVPITLIIPARIDESSHYDLSYTDVHFKPGQIRRHFIEVPQGATWAEVTVCSCSTDVTAKFVLHAVQLVKQKAYRSHEFYKFSSLPEKGSVTEAFPVLGGKTIEFCIARWWASLNDVSIDYTISFHGIVCATPQLNIHASEGIVRFDVQSTLKYEDIAPCINLKSWVQTLRPVSTKIKPLGSRDVLPNNRQLYEMILTYSFHQPKSGEVTPSCPLLCELLYESEFDSQLWIIFDQNKRQMGSGDAYPHQYSLKLEKGDYTIRLQIRHEQNSELDRIKDLPFVVSHRLSNTLSLDIYENHSLALLGKKKSNSLTLPPKHSHPFFVTSLPDDKIPKGAGPGCYLVGALTLSKTELGKKAGQSAAKRQGKFKKDVITVHYHLIPSPTKTKNGSKEKEREPEKEKDVKEEFAEALRDLKIQWMTKLDTSDVFNELKEAFPNHLPLYVARLHQLDSEKERLKRLDEIIEAANTVISHIDQTALAVYFVMKTDPRPDAATIKNDMEKQKSTLVDALCRKGSALADQLLHLQAQDGAVSSDAEGKDDDQESCIGVLSETFCETTKWTDLFDTKVLTFAYKHALANKMYGRGLKFATKLAEEKPTKENLKNCIQLMKLLGWTHCASFSENWLPVMYPPDYCVF</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HS51</accession>
    <name>A0A452HS51_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="2">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="2 3">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="3" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="2" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="2" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000020356.1">
        <property type="protein sequence ID" value="ENSGAGP00000017840.1"/>
        <property type="gene ID" value="ENSGAGG00000013240.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032184">
        <property type="term" value="F:SUMO polymer binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052119">
        <property type="entry name" value="ElonginBC-PRC2_ViralRestrict"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23187">
        <property type="entry name" value="FLJ44216 PROTEIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23187:SF3">
        <property type="entry name" value="SUMO-INTERACTING MOTIF-CONTAINING PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="3" id="KW-1185">Reference proteome</keyword>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="232"/>
            <end position="262"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="248"/>
            <end position="261"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000017840.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="473" mass="53955" checksum="4385A758367BF9C4" modified="2019-05-08" version="1">MRXEQPLLETSNSVLTAVKQKPPEPAKLGPVNKAWLHKLRYCRKTPVHHLFLQGVVQDEESHQNAHRRAQPISSRKLSMVSTTMEENFPKGTLQFLMDFVSCHHYPPKEIVSHVIRNILLSSETGKMLKDAYMLLMKIQILHPATATMVGWDWELLRYVMEEQEETLPGRFLFLQYVVQTLEDDFQRNLRLRFLQKSIAKEVLSCDRCFSNVKEVIEWLVAAITGLGFSQPRKQQQKAPCPSSGADWSISSPGPDSAQNDSETGKAEAVLPAFLSQKVVLLLQRMLSIAVEVDRSPNCSTNKIADVIFPYLVNIPLRSQREAFLNSMESQLLRCKVLELLFQHSCDMPASLPLSLSKILYFLGRHSLLLQYQDHEVTWLRWDEMLQHLILLLTSYRTVVLEHLRSSLSERMDLIISNAKPQLQANDSITSVDIELHIKNFSSRLLQILGKSLPPQIMDKVCALEALLLTIATT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HS58</accession>
    <name>A0A452HS58_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="5">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452HS58"/>
    <dbReference type="Ensembl" id="ENSGAGT00000020361.1">
        <property type="protein sequence ID" value="ENSGAGP00000017845.1"/>
        <property type="gene ID" value="ENSGAGG00000013245.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005525">
        <property type="term" value="F:GTP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003924">
        <property type="term" value="F:GTPase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR053254">
        <property type="entry name" value="Arf-like_GTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006689">
        <property type="entry name" value="Small_GTPase_ARF/SAR"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46724:SF1">
        <property type="entry name" value="ADP RIBOSYLATION FACTOR LIKE GTPASE 10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46724">
        <property type="entry name" value="ADP-RIBOSYLATION FACTOR-LIKE PROTEIN 9-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00025">
        <property type="entry name" value="Arf"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00328">
        <property type="entry name" value="SAR1GTPBP"/>
    </dbReference>
    <dbReference type="SMART" id="SM00177">
        <property type="entry name" value="ARF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00178">
        <property type="entry name" value="SAR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51417">
        <property type="entry name" value="ARF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2 3" id="KW-0342">GTP-binding</keyword>
    <keyword evidence="4" id="KW-0460">Magnesium</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="1 3" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <feature type="binding site" evidence="3">
        <location>
            <begin position="19"/>
            <end position="26"/>
        </location>
        <ligand>
            <name>GTP</name>
            <dbReference type="ChEBI" id="CHEBI:37565"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="4">
        <location>
            <position position="26"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="4">
        <location>
            <position position="44"/>
        </location>
        <ligand>
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="3">
        <location>
            <position position="66"/>
        </location>
        <ligand>
            <name>GTP</name>
            <dbReference type="ChEBI" id="CHEBI:37565"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="3">
        <location>
            <begin position="120"/>
            <end position="123"/>
        </location>
        <ligand>
            <name>GTP</name>
            <dbReference type="ChEBI" id="CHEBI:37565"/>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00023134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="PIRSR" id="PIRSR606689-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="PIRSR" id="PIRSR606689-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000017845.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="180" mass="20004" checksum="A2759368350C291F" modified="2019-05-08" version="1">APLLSWQPDFSMRQVLVLGLKGAGKSSVLHYISSEAAKDRTAPTQGFNSVQVYADGFQIDLLELGGSQNLRFYWNQYLSQAHVLVFVVDSADRPHLPTARQELHWLLAEDPRLPLVVLANKQDKSDALSVAELHRELALHTLNGQREFFLLPTSATHTALATATSIRHVKSLLVELLSQL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452HS59</accession>
    <name>A0A452HS59_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">Sulfotransferase</fullName>
            <ecNumber evidence="5">2.8.2.-</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 5">Belongs to the sulfotransferase 1 family.</text>
    </comment>
    <dbReference type="EC" id="2.8.2.-" evidence="5"/>
    <dbReference type="AlphaFoldDB" id="A0A452HS59"/>
    <dbReference type="Ensembl" id="ENSGAGT00000020399.1">
        <property type="protein sequence ID" value="ENSGAGP00000017882.1"/>
        <property type="gene ID" value="ENSGAGG00000013102.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008146">
        <property type="term" value="F:sulfotransferase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000433">
        <property type="entry name" value="Estrogen sulfotransferase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000863">
        <property type="entry name" value="Sulfotransferase_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11783">
        <property type="entry name" value="SULFOTRANSFERASE SULT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00685">
        <property type="entry name" value="Sulfotransfer_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 5" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Sulfotransferase" evidence="6">
        <location>
            <begin position="22"/>
            <end position="257"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="RuleBase" id="RU361155"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="Pfam" id="PF00685"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000017882.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="263" mass="30160" checksum="85F4CBAEB1B2B6BB" modified="2019-05-08" version="1">SLVGNSRELKVVSGPLTLSSLGTNWMLEILSLIRSDGDPGWVCSVLNWDRVPWVEYGPGLEAARKYPPPRLLCFHLPVQLFPKSLQRSKAKIIYTLRCPKDVVVSLYHFSRIVRLFKDPDSLDCFLEDFLSGNVLYGSWFHHVTGWMGLKANKNIFSITYEDGCLPQDLQGSVRRICHFLGKELSEEQVAAVVENVSFQSMKGNKISNFSQMSDKYMDHQKGEFLRKGICGDWRNHLSEAQSQRFDTVFREWMQGLGMTFPWD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452I3T0</accession>
    <name>A0A452I3T0_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">E1 ubiquitin-activating enzyme</fullName>
            <ecNumber evidence="4">6.2.1.45</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="9">Ubiquitin-activating enzyme E1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="14 15">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="15" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="14" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="14" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="1">
            <text>ATP + ubiquitin + [E1 ubiquitin-activating enzyme]-L-cysteine = AMP + diphosphate + S-ubiquitinyl-[E1 ubiquitin-activating enzyme]-L-cysteine.</text>
            <dbReference type="EC" id="6.2.1.45"/>
        </reaction>
    </comment>
    <comment type="pathway">
        <text evidence="2">Protein modification; protein ubiquitination.</text>
    </comment>
    <comment type="similarity">
        <text evidence="3 11">Belongs to the ubiquitin-activating E1 family.</text>
    </comment>
    <dbReference type="EC" id="6.2.1.45" evidence="4"/>
    <dbReference type="AlphaFoldDB" id="A0A452I3T0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000025280.1">
        <property type="protein sequence ID" value="ENSGAGP00000022176.1"/>
        <property type="gene ID" value="ENSGAGG00000016282.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00143"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031510">
        <property type="term" value="C:SUMO activating enzyme complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0019948">
        <property type="term" value="F:SUMO activating enzyme activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004839">
        <property type="term" value="F:ubiquitin activating enzyme activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016925">
        <property type="term" value="P:protein sumoylation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd01491">
        <property type="entry name" value="Ube1_repeat1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd01490">
        <property type="entry name" value="Ube1_repeat2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.2660:FF:000001">
        <property type="entry name" value="Ubiquitin-activating enzyme E1 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.12550:FF:000001">
        <property type="entry name" value="Ubiquitin-activating enzyme E1 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.720:FF:000015">
        <property type="entry name" value="Ubiquitin-activating enzyme E1 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.30.180:FF:000001">
        <property type="entry name" value="ubiquitin-like modifier-activating enzyme 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.290.60:FF:000002">
        <property type="entry name" value="Ubiquitin-like modifier-activating enzyme 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.50.50.80:FF:000001">
        <property type="entry name" value="ubiquitin-like modifier-activating enzyme 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.720">
        <property type="entry name" value="NAD(P)-binding Rossmann-like Domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.30.180">
        <property type="entry name" value="Ubiquitin-activating enzyme E1, FCCH domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.50.50.80">
        <property type="entry name" value="Ubiquitin-activating enzyme E1, inactive adenylation domain, subdomain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.12550">
        <property type="entry name" value="Ubiquitin-activating enzyme E1, inactive adenylation domain, subdomain 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.2660">
        <property type="entry name" value="Ubiquitin-activating enzyme E1, SCCH domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.290.60">
        <property type="entry name" value="Ubiquitin-activating enzyme E1, UFD domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032420">
        <property type="entry name" value="E1_4HB"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR032418">
        <property type="entry name" value="E1_FCCH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042302">
        <property type="entry name" value="E1_FCCH_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045886">
        <property type="entry name" value="ThiF/MoeB/HesA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000594">
        <property type="entry name" value="ThiF_NAD_FAD-bd"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018965">
        <property type="entry name" value="Ub-activating_enz_E1_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042449">
        <property type="entry name" value="Ub-E1_IAD_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038252">
        <property type="entry name" value="UBA_E1_C_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR019572">
        <property type="entry name" value="UBA_E1_SCCH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042063">
        <property type="entry name" value="Ubi_acti_E1_SCCH"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035985">
        <property type="entry name" value="Ubiquitin-activating_enz"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018075">
        <property type="entry name" value="UBQ-activ_enz_E1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018074">
        <property type="entry name" value="UBQ-activ_enz_E1_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033127">
        <property type="entry name" value="UBQ-activ_enz_E1_Cys_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000011">
        <property type="entry name" value="UBQ/SUMO-activ_enz_E1-like"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01408">
        <property type="entry name" value="Ube1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10953">
        <property type="entry name" value="UBIQUITIN-ACTIVATING ENZYME E1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10953:SF4">
        <property type="entry name" value="UBIQUITIN-ACTIVATING ENZYME E1 C-TERMINAL DOMAIN-CONTAINING PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16191">
        <property type="entry name" value="E1_4HB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16190">
        <property type="entry name" value="E1_FCCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF09358">
        <property type="entry name" value="E1_UFD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00899">
        <property type="entry name" value="ThiF"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF10585">
        <property type="entry name" value="UBA_E1_SCCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01849">
        <property type="entry name" value="UBIQUITINACT"/>
    </dbReference>
    <dbReference type="SMART" id="SM00985">
        <property type="entry name" value="UBA_e1_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF69572">
        <property type="entry name" value="Activating enzymes of the ubiquitin-like proteins"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00536">
        <property type="entry name" value="UBIQUITIN_ACTIVAT_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00865">
        <property type="entry name" value="UBIQUITIN_ACTIVAT_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8 11" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="5 11" id="KW-0436">Ligase</keyword>
    <keyword evidence="6 11" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="15" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="7 11" id="KW-0833">Ubl conjugation pathway</keyword>
    <feature type="domain" description="Ubiquitin-activating enzyme E1 C-terminal" evidence="13">
        <location>
            <begin position="935"/>
            <end position="1066"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="12">
        <location>
            <begin position="1"/>
            <end position="59"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="12">
        <location>
            <begin position="28"/>
            <end position="43"/>
        </location>
    </feature>
    <feature type="active site" description="Glycyl thioester intermediate" evidence="10">
        <location>
            <position position="645"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00000488"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004906"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00005673"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00012990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022598"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022786"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00030371"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU10132"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="RuleBase" id="RU000519"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="13">
        <source>
            <dbReference type="SMART" id="SM00985"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="14">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000022176.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1071" mass="119695" checksum="D9017B62E3FD5B85" modified="2019-05-08" version="1">MRRSRHWCEGTRMSSSPLSKKRRVSGSEPKTGSNCSSGNSVQSDPRAAPANGMAKNGSDVDIDESLYSRQLYVLGHEAMKRMQNASVLVSGLRGLGVEIAKNIILGGVKSVTLHDTEPAQWADLASQFYLREEDLGKNRAEVSQPRLAELNSYVPVSVHNGALTEEVLSPFQVVVLTNAPLEEQLRVGGVCHSKGIKMVVADTRGLFGQLFCDFGDEMVVTDTNGEQPLSAMISMITKGCPGEVTCLDEARHGFESGDFVSFTEVDGMEELNRCPPMEIKVLGPYTFSIGDTSSFSEYVRGGIVSQVKMPKKMSFKPLSVALAEPDFVVTDFGKFDRPAHLHLAFQALHDFQRQHQRLPRPRSEVDAAEVLALTQAVNQRAPPALKQEKLNEGLIKELAYQAAGDLAPVNAFIGGLAAQEVMKACSGKFMPIMQWLYFDALECLPEDSKEALTEEQCRPRNSRYDGQIAVFGADLQAKLGQQKYFLVGAGAIGCELLKNFALLGLACGDGGEVTVTDMDTIEKSNLNRQFLFRPWDVTKMKSDTAAAAVRQMNPNIHITSHQDRVGPDTERVYDDDFFEGLDGVANALDNVDARMYMDRRCVYYRKPLLESGTLGTKGNVQVVIPFLTESYSSSQDPPEKSIPICTLKNFPNAIEHTLQWARDEFEGLFKQPAESVNQYISDPKFMERTLKLPGTQPLEVLEAVNKSLVSERPRSWADCVGWACRHWHCQYSNNIRQLLHNFPPHQKTNSGTLFWSGPKRCPHPLTFDESNPLHMDYIVAAANLFAQTYGITGTRDVGAVAELLHQVQVPEFTPKSGVRIHISDQELQNANASVDDSRLEELKSSLPNPEDLQSFRMYPIDFEKDDDTNFHMDFIVAASNLRAENYDIPPADRHKSKLIAGKIIPAIATTTAAVVGLVCLELYKVVQGHKRLESYKNGFLNLALPFFGFSEPIACPRNKYYDIEWTLWDRFEVQGVQPSGEEMTLRQFLAYFKSEHRLEITMLSQGVSMLYSFFMPPAKLRERHDQPMTEIVTKVSKKKIGRHVKALVFELCCNDESECDIEVPYVRYTIR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452I441</accession>
    <name>A0A452I441_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">C3H1-type domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="9">Belongs to the muscleblind family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452I441"/>
    <dbReference type="Ensembl" id="ENSGAGT00000025419.1">
        <property type="protein sequence ID" value="ENSGAGP00000022304.1"/>
        <property type="gene ID" value="ENSGAGG00000016381.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005654">
        <property type="term" value="C:nucleoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043484">
        <property type="term" value="P:regulation of RNA splicing"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="entry name" value="Muscleblind like splicing regulator 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1370.210:FF:000002">
        <property type="entry name" value="Muscleblind-like 1 isoform 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1370.210">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR054429">
        <property type="entry name" value="Znf-CCCH_Muscleblind-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000571">
        <property type="entry name" value="Znf_CCCH"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12675">
        <property type="entry name" value="MUSCLEBLIND-LIKE PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12675:SF7">
        <property type="entry name" value="MUSCLEBLIND-LIKE PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00642">
        <property type="entry name" value="zf-CCCH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF22628">
        <property type="entry name" value="zf-CCCH_10"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14608">
        <property type="entry name" value="zf-CCCH_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00356">
        <property type="entry name" value="ZnF_C3H1"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50103">
        <property type="entry name" value="ZF_C3H1"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4 10" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="7 10" id="KW-0862">Zinc</keyword>
    <keyword evidence="6 10" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="C3H1-type" evidence="11">
        <location>
            <begin position="13"/>
            <end position="41"/>
        </location>
    </feature>
    <feature type="domain" description="C3H1-type" evidence="11">
        <location>
            <begin position="47"/>
            <end position="73"/>
        </location>
    </feature>
    <feature type="domain" description="C3H1-type" evidence="11">
        <location>
            <begin position="178"/>
            <end position="206"/>
        </location>
    </feature>
    <feature type="domain" description="C3H1-type" evidence="11">
        <location>
            <begin position="214"/>
            <end position="240"/>
        </location>
    </feature>
    <feature type="zinc finger region" description="C3H1-type" evidence="10">
        <location>
            <begin position="13"/>
            <end position="41"/>
        </location>
    </feature>
    <feature type="zinc finger region" description="C3H1-type" evidence="10">
        <location>
            <begin position="47"/>
            <end position="73"/>
        </location>
    </feature>
    <feature type="zinc finger region" description="C3H1-type" evidence="10">
        <location>
            <begin position="178"/>
            <end position="206"/>
        </location>
    </feature>
    <feature type="zinc finger region" description="C3H1-type" evidence="10">
        <location>
            <begin position="214"/>
            <end position="240"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004496"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00038226"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50103"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000022304.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="379" mass="41424" checksum="6B122F40184B0FC7" modified="2019-05-08" version="1">MAVSVTPIRDTKWLTLEVCREFQRGTCSRPDTECKFAHPSKSCQVENGRVIACFDSLKGRCSRENCKYLHPPPHLKTQLEINGRNNLIQQKNMAMLAQQMQLANAMMPGAPLQPVPMFSVAPSLATNASAAFNPYLGPVSPGLVPAEILPTAPMLVAGNPGVPVPAAAAAAAQKLMRTDRLEVCREYQRGNCNRGENDCRFAHPADSAMIDTNDNTVTVCMDYIKGRCSREKCKYFHPPAHLQAKIKAAQYQVNQAAAAQAAATAAAMGIPQAVLPPLPKRPALEKTNGATAVFNTGIFQYQQALANMQLQQHTAFLPPGKYLFSLCRAKYATKLVFSICNKLLKVSVLIVIDIYPVVVRCFLLRVRKGSLGITVFVIR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452I445</accession>
    <name>A0A452I445_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">BRCT domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
            <location evidence="2">Microtubule organizing center</location>
            <location evidence="2">Centrosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452I445"/>
    <dbReference type="Ensembl" id="ENSGAGT00000025421.1">
        <property type="protein sequence ID" value="ENSGAGP00000022306.1"/>
        <property type="gene ID" value="ENSGAGG00000016303.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035861">
        <property type="term" value="C:site of double-strand break"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006281">
        <property type="term" value="P:DNA repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:2000781">
        <property type="term" value="P:positive regulation of double-strand break repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:1990166">
        <property type="term" value="P:protein localization to site of double-strand break"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10190:FF:000018">
        <property type="entry name" value="DNA topoisomerase 2-binding protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.10190">
        <property type="entry name" value="BRCT domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001357">
        <property type="entry name" value="BRCT_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036420">
        <property type="entry name" value="BRCT_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042479">
        <property type="entry name" value="Slf1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057595">
        <property type="entry name" value="TopB1_SLF1_BRCT"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46677">
        <property type="entry name" value="SMC5-SMC6 COMPLEX LOCALIZATION FACTOR PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46677:SF1">
        <property type="entry name" value="SMC5-SMC6 COMPLEX LOCALIZATION FACTOR PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23294">
        <property type="entry name" value="BRCT_TopB1_SLF1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16770">
        <property type="entry name" value="RTT107_BRCT_5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00292">
        <property type="entry name" value="BRCT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52113">
        <property type="entry name" value="BRCT domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="9" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="5" id="KW-0227">DNA damage</keyword>
    <keyword evidence="6" id="KW-0234">DNA repair</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="BRCT" evidence="10">
        <location>
            <begin position="3"/>
            <end position="80"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="9">
        <location>
            <begin position="672"/>
            <end position="705"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="9">
        <location>
            <begin position="706"/>
            <end position="738"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="9">
        <location>
            <begin position="740"/>
            <end position="772"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="SMART" id="SM00292"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000022306.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="929" mass="107058" checksum="115FEFF1D7C31ABD" modified="2019-05-08" version="1">MACAVQKRRIQLTGFKKQEKEAIAELLLTLDCVFIDTEQYRNCTHLIAKQLSKSEKFLAACAAGKWILTKDYIINSAESGRWLDETTYEWGYKIEKDSHYSPQMQSAPKRWREELTCSGALGAFHRWKVILLVKEGDKRRDSITRVLEAGKAIIYPPQKEVRAVTHVFTDNKSVTIEREKCLFEAPYYPVQYIGDYLFENEIQNITEDNEMNHLLAEQEARDTSDMQLVAMKNALIKHMYFTQVRKKQGYTFYVVFHKYLPFQVKNIDLCRGILCILEGLVEGHFFTDVITELAATQKYFIPPVQLLHSLLEHMLQGNTDTVFSAKFFHILYILLQLDPPWKSPSMLRYYLEFLQCPICMKGTWSLLEMLVRSCLYCKSFCHAVSVSEKKNEQRVVHKTLLKFFFNLVKAEVQALTRRLCEGANSQLLQVMPQTVLLKTFWLGSETSLLFTKYINILVDWVIHSYREKFKTNDAFKDEVAILLIGILGTVVEYWILSGFMMDRNVLHQVADDLANYIAISCDDFSTQELKMFICSIPSLWLQMFVAEAVFKNLCLRRNITISAEPLSLHKIVYSCLPALGEVGMHETGKVQKVKKKKIGQWPCPESQRALLMLNGDKQNQTKRLPDLPELTFNKFPPLSKKLRTKAEVEMSNTKENRHPLDEEMHFYKTNIKGETALHIACIRNKVERLIQLLPFPGTDINVKDYAGWTPLHEACNHGNTVCVREILQRCPEVDLLSQVDGVTPLHDALSNGHVEIGKLLLQRGGPLLLRQRDFYGKLPLDYVESLPVKQELLDIVHPKETVKDFCERVEKDFHNQQRELWLILFNKMLLNFCSVYNLSSPSPLSFKAHLNSLPLVAINGCKLKSRSLTDWLVDLYVGEVQTFQKLPEFIQEISGNLKKSFGEEIYPLLATLETIAMATQVATFHLSNN</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452I446</accession>
    <name>A0A452I446_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Procollagen-lysine,2-oxoglutarate 5-dioxygenase 1</fullName>
            <ecNumber evidence="3">1.14.11.4</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="15">Lysyl hydroxylase 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="16">
            <text>L-lysyl-[collagen] + 2-oxoglutarate + O2 = (5R)-5-hydroxy-L-lysyl-[collagen] + succinate + CO2</text>
            <dbReference type="Rhea" id="RHEA:16569"/>
            <dbReference type="Rhea" id="RHEA-COMP:12751"/>
            <dbReference type="Rhea" id="RHEA-COMP:12752"/>
            <dbReference type="ChEBI" id="CHEBI:15379"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:16810"/>
            <dbReference type="ChEBI" id="CHEBI:29969"/>
            <dbReference type="ChEBI" id="CHEBI:30031"/>
            <dbReference type="ChEBI" id="CHEBI:133442"/>
            <dbReference type="EC" id="1.14.11.4"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Fe(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29033"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="2">
            <name>L-ascorbate</name>
            <dbReference type="ChEBI" id="CHEBI:38290"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="13">Rough endoplasmic reticulum membrane</location>
            <topology evidence="13">Peripheral membrane protein</topology>
            <orientation evidence="13">Lumenal side</orientation>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="1.14.11.4" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452I446"/>
    <dbReference type="Ensembl" id="ENSGAGT00000025424.1">
        <property type="protein sequence ID" value="ENSGAGP00000022309.1"/>
        <property type="gene ID" value="ENSGAGG00000016382.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030867">
        <property type="term" value="C:rough endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005506">
        <property type="term" value="F:iron ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031418">
        <property type="term" value="F:L-ascorbic acid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008475">
        <property type="term" value="F:procollagen-lysine 5-dioxygenase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="CDD" id="cd23004">
        <property type="entry name" value="GT_LH1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.620">
        <property type="entry name" value="q2cbj1_9rhob like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050757">
        <property type="entry name" value="Collagen_mod_GT25"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057589">
        <property type="entry name" value="GT_PLOD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044861">
        <property type="entry name" value="IPNS-like_FE2OG_OXY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029044">
        <property type="entry name" value="Nucleotide-diphossugar_trans"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005123">
        <property type="entry name" value="Oxoglu/Fe-dep_dioxygenase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006620">
        <property type="entry name" value="Pro_4_hyd_alph"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001006">
        <property type="entry name" value="Procol_lys_dOase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10730:SF5">
        <property type="entry name" value="PROCOLLAGEN-LYSINE,2-OXOGLUTARATE 5-DIOXYGENASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10730">
        <property type="entry name" value="PROCOLLAGEN-LYSINE,2-OXOGLUTARATE 5-DIOXYGENASE/GLYCOSYLTRANSFERASE 25 FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03171">
        <property type="entry name" value="2OG-FeII_Oxy"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25342">
        <property type="entry name" value="GT_PLOD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00702">
        <property type="entry name" value="P4Hc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53448">
        <property type="entry name" value="Nucleotide-diphospho-sugar transferases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51471">
        <property type="entry name" value="FE2OG_OXY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01325">
        <property type="entry name" value="LYS_HYDROXYLASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8" id="KW-0223">Dioxygenase</keyword>
    <keyword evidence="6" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10" id="KW-0408">Iron</keyword>
    <keyword evidence="11" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <keyword evidence="7" id="KW-0847">Vitamin C</keyword>
    <feature type="domain" description="Fe2OG dioxygenase" evidence="17">
        <location>
            <begin position="656"/>
            <end position="747"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001954"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00001961"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012264"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022896"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022964"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00037819"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00040791"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00042560"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00047930"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS51471"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000022309.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="747" mass="85232" checksum="D6CE553753A39CAC" modified="2019-05-08" version="1">MSQDCREEPPGGACLVRGSENLAAPGVGRSGFPLCPAPRCTGLGPVCRQHRGWNRTRGDQDPPQVASMLPQLLLLPGLVLTLLIPEGSCGSKQEENLLVLTVATQETEGFKRFKRSAQFFNYKVQVLGLDEEWRGEDKKAGGGGQKVRLLKSALKEYADKEDLIILVTESYDVLFASGPAELLKKFKQAKSKVVFSAETFIYPDRRLEAKYPPVRDGKRFLGSGGFIGYAPNLNKLVEDWKGLDNDSDQLFYTNIFLDPEKRENINITLDQRCRIFQNLNGALDEVALKFENARVRARNVEYDTLPVLIHGNGPTKLQLNYLGNYIPRAWTFETGCTVCDEGLRSLTGFKDDVLPLVLIGIFIKQPTPFLSQFFLRLQNFQYPKKRTQLFIHNHEQHHQSRVDTFVKEHGKEYHAVKVIGPEDHMENADARNMGMDLCRQDPDCEYYFSLDAEVVLKNSKVLRILIEQNKQVIAPLVSRLGKLWSNFWGALSPDGYYARSEDYVDIVQGRRVGIWNVPYISSIYVIKGSTLRSQLNQGDLFHSGKLDADMAFCHNVRDQRVFMFVTNRHQFGHILSLEDYQTTHLHNDLWQIFSNPEDWKEKYIHENYTLALKGKLVETPCPDVYWFPIFTDVACDELVEEMEHFGQWSTGDNTTQFELAFVVRYKPDEQPSLMPHHDASTFTINIALNRVGIDYEGGGCRFLRYNCSIRAPRKGWTLLHPGRLTHYHEGLPTTKGTRYIAVSFLDP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452I447</accession>
    <name>A0A452I447_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Laminin subunit beta-1</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="17">Laminin B1 chain</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="15">Laminin-1 subunit beta</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="19">Laminin-10 subunit beta</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="16">Laminin-12 subunit beta</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="20">Laminin-2 subunit beta</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="18">Laminin-6 subunit beta</fullName>
        </alternativeName>
        <alternativeName>
            <fullName evidence="21">Laminin-8 subunit beta</fullName>
        </alternativeName>
    </protein>
    <organism evidence="27 28">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="28" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="27" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="27" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="13">Laminin is a complex glycoprotein, consisting of three different polypeptide chains (alpha, beta, gamma), which are bound to each other by disulfide bonds into a cross-shaped molecule comprising one long and three short arms with globules at each end. Beta-1 is a subunit of laminin-1 (laminin-111 or EHS laminin), laminin-2 (laminin-211 or merosin), laminin-6 (laminin-311 or K-laminin), laminin-8 (laminin-411), laminin-10 (laminin-511) and laminin-12 (laminin-213). Interacts with ITGB1.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
            <location evidence="1">Extracellular space</location>
            <location evidence="1">Extracellular matrix</location>
            <location evidence="1">Basement membrane</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="22">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
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        <property type="protein sequence ID" value="ENSGAGP00000022295.1"/>
        <property type="gene ID" value="ENSGAGG00000014979.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005606">
        <property type="term" value="C:laminin-1 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043259">
        <property type="term" value="C:laminin-10 complex"/>
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        <property type="project" value="UniProtKB-ARBA"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005178">
        <property type="term" value="F:integrin binding"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
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        <property type="term" value="F:structural constituent of synapse-associated extracellular matrix"/>
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    </dbReference>
    <dbReference type="GO" id="GO:0009887">
        <property type="term" value="P:animal organ morphogenesis"/>
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    </dbReference>
    <dbReference type="GO" id="GO:0007411">
        <property type="term" value="P:axon guidance"/>
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    </dbReference>
    <dbReference type="GO" id="GO:0070831">
        <property type="term" value="P:basement membrane assembly"/>
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    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
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    </dbReference>
    <dbReference type="GO" id="GO:0034446">
        <property type="term" value="P:substrate adhesion-dependent cell spreading"/>
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    </dbReference>
    <dbReference type="GO" id="GO:0009888">
        <property type="term" value="P:tissue development"/>
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        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd22300">
        <property type="entry name" value="cc_LAMB1_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd00055">
        <property type="entry name" value="EGF_Lam"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000011">
        <property type="entry name" value="Cadherin EGF LAG seven-pass G-type receptor"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000083">
        <property type="entry name" value="Laminin subunit alpha"/>
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    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000084">
        <property type="entry name" value="Laminin subunit alpha 3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000065">
        <property type="entry name" value="Laminin subunit beta 1"/>
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    </dbReference>
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        <property type="entry name" value="Laminin subunit beta 1"/>
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    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000130">
        <property type="entry name" value="Laminin subunit beta 1"/>
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    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000138">
        <property type="entry name" value="Laminin subunit beta 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000145">
        <property type="entry name" value="Laminin subunit beta 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.170.300.10:FF:000004">
        <property type="entry name" value="Laminin subunit beta 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000135">
        <property type="entry name" value="Laminin subunit beta 4"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000280">
        <property type="entry name" value="Laminin subunit beta 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.170.300.10:FF:000001">
        <property type="entry name" value="Laminin subunit beta-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.260:FF:000225">
        <property type="entry name" value="Laminin subunit beta-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.260">
        <property type="entry name" value="Galactose-binding domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.170.300.10">
        <property type="entry name" value="Tie2 ligand-binding domain superfamily"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056558">
        <property type="entry name" value="LAMB1-4_helical"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050440">
        <property type="entry name" value="Laminin/Netrin_ECM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013015">
        <property type="entry name" value="Laminin_IV_B"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008211">
        <property type="entry name" value="Laminin_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002049">
        <property type="entry name" value="LE_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR056863">
        <property type="entry name" value="LMN_ATRN_NET-like_EGF"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10574:SF233">
        <property type="entry name" value="LAMININ SUBUNIT BETA-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10574">
        <property type="entry name" value="NETRIN/LAMININ-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00053">
        <property type="entry name" value="EGF_laminin"/>
        <property type="match status" value="11"/>
    </dbReference>
    <dbReference type="Pfam" id="PF24973">
        <property type="entry name" value="EGF_LMN_ATRN"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23219">
        <property type="entry name" value="LAMB1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21199">
        <property type="entry name" value="LAMININ_IV_B"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00055">
        <property type="entry name" value="Laminin_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00011">
        <property type="entry name" value="EGFLAMININ"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="6"/>
    </dbReference>
    <dbReference type="SMART" id="SM00180">
        <property type="entry name" value="EGF_Lam"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="SMART" id="SM00136">
        <property type="entry name" value="LamNT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="13"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01248">
        <property type="entry name" value="EGF_LAM_1"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50027">
        <property type="entry name" value="EGF_LAM_2"/>
        <property type="match status" value="10"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51116">
        <property type="entry name" value="LAMININ_IVB"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51117">
        <property type="entry name" value="LAMININ_NTER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="7" id="KW-0084">Basement membrane</keyword>
    <keyword evidence="8" id="KW-0130">Cell adhesion</keyword>
    <keyword evidence="9 23" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="10 22" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0272">Extracellular matrix</keyword>
    <keyword evidence="11" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="12 22" id="KW-0424">Laminin EGF-like domain</keyword>
    <keyword evidence="4" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="28" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0964">Secreted</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <feature type="domain" description="Laminin N-terminal" evidence="26">
        <location>
            <begin position="1"/>
            <end position="220"/>
        </location>
    </feature>
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            <begin position="221"/>
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        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="24">
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        </location>
    </feature>
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            <begin position="348"/>
            <end position="407"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="24">
        <location>
            <begin position="408"/>
            <end position="459"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin IV type B" evidence="25">
        <location>
            <begin position="499"/>
            <end position="717"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="24">
        <location>
            <begin position="723"/>
            <end position="770"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="24">
        <location>
            <begin position="771"/>
            <end position="816"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="24">
        <location>
            <begin position="817"/>
            <end position="866"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="24">
        <location>
            <begin position="978"/>
            <end position="1034"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="24">
        <location>
            <begin position="1035"/>
            <end position="1082"/>
        </location>
    </feature>
    <feature type="domain" description="Laminin EGF-like" evidence="24">
        <location>
            <begin position="1083"/>
            <end position="1129"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="23">
        <location>
            <begin position="1247"/>
            <end position="1274"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="23">
        <location>
            <begin position="1345"/>
            <end position="1407"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="23">
        <location>
            <begin position="1455"/>
            <end position="1609"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="22">
        <location>
            <begin position="250"/>
            <end position="259"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="22">
        <location>
            <begin position="315"/>
            <end position="324"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="22">
        <location>
            <begin position="378"/>
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    </feature>
    <feature type="disulfide bond" evidence="22">
        <location>
            <begin position="431"/>
            <end position="440"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="22">
        <location>
            <begin position="443"/>
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        </location>
    </feature>
    <feature type="disulfide bond" evidence="22">
        <location>
            <begin position="723"/>
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        </location>
    </feature>
    <feature type="disulfide bond" evidence="22">
        <location>
            <begin position="725"/>
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        </location>
    </feature>
    <feature type="disulfide bond" evidence="22">
        <location>
            <begin position="744"/>
            <end position="753"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="22">
        <location>
            <begin position="771"/>
            <end position="783"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="22">
        <location>
            <begin position="773"/>
            <end position="790"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="22">
        <location>
            <begin position="792"/>
            <end position="801"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="22">
        <location>
            <begin position="836"/>
            <end position="845"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="22">
        <location>
            <begin position="978"/>
            <end position="990"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="22">
        <location>
            <begin position="1007"/>
            <end position="1016"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="22">
        <location>
            <begin position="1035"/>
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        </location>
    </feature>
    <feature type="disulfide bond" evidence="22">
        <location>
            <begin position="1037"/>
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        </location>
    </feature>
    <feature type="disulfide bond" evidence="22">
        <location>
            <begin position="1056"/>
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    </feature>
    <feature type="disulfide bond" evidence="22">
        <location>
            <begin position="1083"/>
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    </feature>
    <feature type="disulfide bond" evidence="22">
        <location>
            <begin position="1085"/>
            <end position="1102"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="22">
        <location>
            <begin position="1104"/>
            <end position="1113"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004302"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022530"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
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    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00022869"/>
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    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022889"/>
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    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
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    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
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    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
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    </evidence>
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        <source>
            <dbReference type="ARBA" id="ARBA00023292"/>
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    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00065312"/>
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    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00071083"/>
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    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00075282"/>
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    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00075415"/>
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    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00076920"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="ARBA" id="ARBA00076958"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="ARBA" id="ARBA00082919"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="ARBA" id="ARBA00083431"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="ARBA" id="ARBA00083813"/>
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    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00460"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="23">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS50027"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS51116"/>
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    </evidence>
    <evidence type="ECO:0000259" key="26">
        <source>
            <dbReference type="PROSITE" id="PS51117"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000022295.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="28">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1621" mass="179446" checksum="2EE8B6034F42FE1E" modified="2019-05-08" version="1">MRGLECRRGLWVWGMGLGGGAQADLRQLLVSSTAGLRQAARLSWLHAACRKQVWLLGGEAPRLKAPPPPAESRCSQHYINICCFLFLFFLKTFRPAAMLIERSSDFGKTWKVYRYFAYDCENSFPGISTGPMKKVDDIICDSRYSDIEPSTEGEVIYRALDPAFRIEDPYSPRIQNLLKITNLRISFTKLHTLGDNLLDSRMEIKEKYYYAIYDMVVRGNCFCYGHASECAPVDGFNEEVEGMVHGHCMCRHNTKGLNCELCLDFYHDLPWRPAEGRHSNACKKCNCNGHSIQCHFDMAVYMATGNTSGGVCDDCQHNTMGHNCEQCKPFYFQHPEREIRDPNICEPCNCDPLGSQNGGICDRYTDFSAGLIAGQCRCKLFVEGERCDLCKEGFYGLSADNPLGCQSCACNPLGTLPGGNPCDSETGSCYCKRLATGRHCEQCLPEHWGLSNDMDGCRPCDCDQGGAINNNCSSETGQCQCRPHMIGRQCNEVESGYYFMSLDHYIYEAEEANFGPGVSIVERQYSQDRTPSWTGIGFVRVPEGAYLEFYVDNIPYSMEYDVVIRYEPQLPDHWEKVVITVSRPGKIPTSSRCGNTVPDDDNQTVSLSPGSRYVVLPRPVCFEKGLNYTVRLELSQYSSFDTEVESPYTLIDSLVLMPYCKSLDIFTVGGSGEDVVTNSAWETFQRYRCLENSRSVVKTPMTDVCKNIIFSISALLHETALSCQCNPQGSLSSVCDSNGGRCQCRPNVVGRRCDRCAPGTFGFGPNGCKSCECHLQGSVSSFCHPETGQCYCFHGVYGRQCDRCLPGHWGFPSCQPCQCNGHTDDCDPYTGECLNCRDSTTGHNCERCLGGYYGDPVLGSGDHCRPCPCPDGPESGRQFASSCYQDPITLQIVCVCNVGYIGSRCDECASGFFGDPEERGGVCQPCQCNNNIDMTDPGACDEQSGKCLNCLYQTEGENCQLCKYGYYGHALQQDCRKCVCNYLGTVREHCNSSEDCQCEKTTGQCQCLPNVTGQNCDRCAPNTWRLASGTGCELCECDVARSFRPSCNEFTGQCQCMPGFGGRTCRECQEFFWGDPDVACQACDCDPRGIQTPQCNRSTGLCACKEGVEGPHCDKCTRGYSGIFPDCVPCHQCFAVWDVIISELTNRTQRFLERAKILKISGVSGPYQQTINTMEEKLNEIKSILAQNPAAEPLKNIGNLFEELDLISAGALDSITKYFQMSLEAEERVNASTVHPDSLVELSAQTRQEVEDLISEQEAQFKAKQEEQSRLLDELAGKLQSLDLSEVAEKTCGTPSGASCADSECGGLNCRTDEGEKKCGGPGCEGLVTVAHSAWQKAMDFDRDILSALAEVEELSKMVSEAKQRADKAKQNAQEVLLKTNATKEQVDRSNEDLRNLIKQIREFLMQDSADLDSIEAVANEVLKMEMPSSPQQLQTLTEDIRERVESLSDVEIILQQSAGDIARAEMLLEEANKASKSAKDVKVTAAIVKEALEEAEKAQNAAGKAIKQAEEDIRGTQDLLTSVLDSELHAKYKTVENLIAKKTEESADARKKAGMLQNEAKSLLAQANSKLQLLKDLESTYENNQKVLEDKAKQLVELEETVRSLLQAISQKVAVYSTCL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452I450</accession>
    <name>A0A452I450_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Procollagen-lysine,2-oxoglutarate 5-dioxygenase 1</fullName>
            <ecNumber evidence="3">1.14.11.4</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="15">Lysyl hydroxylase 1</fullName>
        </alternativeName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-05" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="16">
            <text>L-lysyl-[collagen] + 2-oxoglutarate + O2 = (5R)-5-hydroxy-L-lysyl-[collagen] + succinate + CO2</text>
            <dbReference type="Rhea" id="RHEA:16569"/>
            <dbReference type="Rhea" id="RHEA-COMP:12751"/>
            <dbReference type="Rhea" id="RHEA-COMP:12752"/>
            <dbReference type="ChEBI" id="CHEBI:15379"/>
            <dbReference type="ChEBI" id="CHEBI:16526"/>
            <dbReference type="ChEBI" id="CHEBI:16810"/>
            <dbReference type="ChEBI" id="CHEBI:29969"/>
            <dbReference type="ChEBI" id="CHEBI:30031"/>
            <dbReference type="ChEBI" id="CHEBI:133442"/>
            <dbReference type="EC" id="1.14.11.4"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1">
            <name>Fe(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29033"/>
        </cofactor>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="2">
            <name>L-ascorbate</name>
            <dbReference type="ChEBI" id="CHEBI:38290"/>
        </cofactor>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="13">Rough endoplasmic reticulum membrane</location>
            <topology evidence="13">Peripheral membrane protein</topology>
            <orientation evidence="13">Lumenal side</orientation>
        </subcellularLocation>
    </comment>
    <dbReference type="EC" id="1.14.11.4" evidence="3"/>
    <dbReference type="AlphaFoldDB" id="A0A452I450"/>
    <dbReference type="Ensembl" id="ENSGAGT00000025426.1">
        <property type="protein sequence ID" value="ENSGAGP00000022311.1"/>
        <property type="gene ID" value="ENSGAGG00000016382.1"/>
    </dbReference>
    <dbReference type="Ensembl" id="ENSGAGT00000025429.1">
        <property type="protein sequence ID" value="ENSGAGP00000022314.1"/>
        <property type="gene ID" value="ENSGAGG00000016382.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030867">
        <property type="term" value="C:rough endoplasmic reticulum membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005506">
        <property type="term" value="F:iron ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031418">
        <property type="term" value="F:L-ascorbic acid binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008475">
        <property type="term" value="F:procollagen-lysine 5-dioxygenase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="CDD" id="cd23004">
        <property type="entry name" value="GT_LH1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.620">
        <property type="entry name" value="q2cbj1_9rhob like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050757">
        <property type="entry name" value="Collagen_mod_GT25"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057589">
        <property type="entry name" value="GT_PLOD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR044861">
        <property type="entry name" value="IPNS-like_FE2OG_OXY"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029044">
        <property type="entry name" value="Nucleotide-diphossugar_trans"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005123">
        <property type="entry name" value="Oxoglu/Fe-dep_dioxygenase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006620">
        <property type="entry name" value="Pro_4_hyd_alph"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001006">
        <property type="entry name" value="Procol_lys_dOase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10730:SF5">
        <property type="entry name" value="PROCOLLAGEN-LYSINE,2-OXOGLUTARATE 5-DIOXYGENASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10730">
        <property type="entry name" value="PROCOLLAGEN-LYSINE,2-OXOGLUTARATE 5-DIOXYGENASE/GLYCOSYLTRANSFERASE 25 FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF03171">
        <property type="entry name" value="2OG-FeII_Oxy"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF25342">
        <property type="entry name" value="GT_PLOD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00702">
        <property type="entry name" value="P4Hc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53448">
        <property type="entry name" value="Nucleotide-diphospho-sugar transferases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51471">
        <property type="entry name" value="FE2OG_OXY"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01325">
        <property type="entry name" value="LYS_HYDROXYLASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8" id="KW-0223">Dioxygenase</keyword>
    <keyword evidence="6" id="KW-0256">Endoplasmic reticulum</keyword>
    <keyword evidence="12" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10" id="KW-0408">Iron</keyword>
    <keyword evidence="11" id="KW-0472">Membrane</keyword>
    <keyword evidence="4" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="9" id="KW-0560">Oxidoreductase</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0732">Signal</keyword>
    <keyword evidence="7" id="KW-0847">Vitamin C</keyword>
    <feature type="domain" description="Fe2OG dioxygenase" evidence="17">
        <location>
            <begin position="581"/>
            <end position="672"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001954"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00001961"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00012264"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022824"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022896"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022964"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023002"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023004"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00037819"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00040791"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00042560"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00047930"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="PROSITE" id="PS51471"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000022311.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="672" mass="77376" checksum="093BEF5E69AB32C0" modified="2019-05-08" version="1">CETDEGGTSLFTLCFLLPTENLLVLTVATQETEGFKRFKRSAQFFNYKVQVLGLDEEWRGEDKKAGGGGQKVRLLKSALKEYADKEDLIILVTESYDVLFASGPAELLKKFKQAKSKVVFSAETFIYPDRRLEAKYPPVRDGKRFLGSGGFIGYAPNLNKLVEDWKGLDNDSDQLFYTNIFLDPEKRENINITLDQRCRIFQNLNGALDEVALKFENARVRARNVEYDTLPVLIHGNGPTKLQLNYLGNYIPRAWTFETGCTVCDEGLRSLTGFKDDVLPLVLIGIFIKQPTPFLSQFFLRLQNFQYPKKRTQLFIHNHEQHHQSRVDTFVKEHGKEYHAVKVIGPEDHMENADARNMGMDLCRQDPDCEYYFSLDAEVVLKNSKVLRILIEQNKQVIAPLVSRLGKLWSNFWGALSPDGYYARSEDYVDIVQGRRVGIWNVPYISSIYVIKGSTLRSQLNQGDLFHSGKLDADMAFCHNVRDQRVFMFVTNRHQFGHILSLEDYQTTHLHNDLWQIFSNPEDWKEKYIHENYTLALKGKLVETPCPDVYWFPIFTDVACDELVEEMEHFGQWSTGDNTTQFELAFVVRYKPDEQPSLMPHHDASTFTINIALNRVGIDYEGGGCRFLRYNCSIRAPRKGWTLLHPGRLTHYHEGLPTTKGTRYIAVSFLDP</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452I453</accession>
    <name>A0A452I453_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">PKD domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Membrane</location>
            <topology evidence="1">Single-pass type I membrane protein</topology>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2">Belongs to the VPS10-related sortilin family. SORCS subfamily.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452I453"/>
    <dbReference type="Ensembl" id="ENSGAGT00000025433.1">
        <property type="protein sequence ID" value="ENSGAGP00000022318.1"/>
        <property type="gene ID" value="ENSGAGG00000013961.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005794">
        <property type="term" value="C:Golgi apparatus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006892">
        <property type="term" value="P:post-Golgi vesicle-mediated transport"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd15482">
        <property type="entry name" value="Sialidase_non-viral"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.60.270:FF:000001">
        <property type="entry name" value="Sortilin related VPS10 domain containing receptor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.10:FF:000083">
        <property type="entry name" value="Sortilin-related VPS10 domain containing receptor 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.70.80:FF:000001">
        <property type="entry name" value="Sortilin-related VPS10 domain-containing receptor 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.70.80">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.120.10.10">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.60.270">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.10">
        <property type="entry name" value="Immunoglobulins"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013783">
        <property type="entry name" value="Ig-like_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000601">
        <property type="entry name" value="PKD_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035986">
        <property type="entry name" value="PKD_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031777">
        <property type="entry name" value="Sortilin_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031778">
        <property type="entry name" value="Sortilin_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006581">
        <property type="entry name" value="VPS10"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050310">
        <property type="entry name" value="VPS10-sortilin"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12106">
        <property type="entry name" value="SORTILIN RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12106:SF8">
        <property type="entry name" value="VPS10 DOMAIN-CONTAINING RECEPTOR SORCS1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00801">
        <property type="entry name" value="PKD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15902">
        <property type="entry name" value="Sortilin-Vps10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15901">
        <property type="entry name" value="Sortilin_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00602">
        <property type="entry name" value="VPS10"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF110296">
        <property type="entry name" value="Oligoxyloglucan reducing end-specific cellobiohydrolase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49299">
        <property type="entry name" value="PKD domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50093">
        <property type="entry name" value="PKD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="8" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="7 10" id="KW-0472">Membrane</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="4" id="KW-0732">Signal</keyword>
    <keyword evidence="3 10" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="6 10" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="10">
        <location>
            <begin position="783"/>
            <end position="801"/>
        </location>
    </feature>
    <feature type="domain" description="PKD" evidence="11">
        <location>
            <begin position="517"/>
            <end position="557"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="821"/>
            <end position="848"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004479"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00010818"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS50093"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000022318.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="848" mass="94726" checksum="96A5540EE94D7282" modified="2019-05-08" version="1">MTACSPFLSDLIYVPIAKPATDWIVLEQWSARAHAAAERPFNAQYITCRMQNCTEATRSKPFPGYIDPDSLIVQDDYVFQCLFPSHSKSFFPMQNPTPKSLDMHVISTDENQVFAAVQEWNQNDTYNLYISDTRGVYFTLALENIKSSRGLEGNVMIDLYEVAGIKGMFLANKKIDSQVKTFITYNKGRDWRLLQAPDTDLRGNPVHCLLPYCSLHLHLKVSENPYTSGNIASRDTAPSIIVASGNIGTELSDNDISMFVSSDAGNTWRQIFEEEHSVLYLDQGGVLVAMKHTSLPIRHLWLSFDEGRSWSKYSFTSTPLFVDGVLGEPGEETLIMTVFGHFSHRSEWQLVKVDYKSIFDRRCAEEDYRPWQLHSQGEACIMGAKRIYRKRKSEKKCMQGKYAGAMASEPCVCTEADFDCDYGYERHTNGQCLPAFWYNPSSLSKDCSFGQSYLNSTGYRKVVSNNCTDGVREQYTAKPQQCPGKAPHGLRIITSDGKLTAEQGHNVTLMVQLEEVDFGDGIAVSYANLSSTEDGIKHVYQNVGIFRVTVQVENSLGLDSAVLYLHVTCPLEHVHLSLPFVTTKNKEVNATAVLWPSQVGTLTYIWWFGNNTEPLVTLEGSITFTFSVEGMNTITVQVSAGNTILQDTKTIAVYERFQSLRLSFSPNLDEYNPDIPEWRRDVSRVIKRALVEATSISSKHILVAVLPGLPTSAELFVLPCQEATGENKRPAEDLDQISEILIHKLNQNLVQFELKPGVRVLVHAAHLSAVPLVDLTPSHSGSAMLMLLSVVFVGLAVFVIYKFKRYDLPLPLPPLSTDSGLTSMTLEQPSHESPMFSSSRPPSPGPDV</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452I457</accession>
    <name>A0A452I457_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="10">BRCT domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="2">Cytoplasm</location>
            <location evidence="2">Cytoskeleton</location>
            <location evidence="2">Microtubule organizing center</location>
            <location evidence="2">Centrosome</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452I457"/>
    <dbReference type="Ensembl" id="ENSGAGT00000025420.1">
        <property type="protein sequence ID" value="ENSGAGP00000022305.1"/>
        <property type="gene ID" value="ENSGAGG00000016303.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005813">
        <property type="term" value="C:centrosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035861">
        <property type="term" value="C:site of double-strand break"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006281">
        <property type="term" value="P:DNA repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:2000781">
        <property type="term" value="P:positive regulation of double-strand break repair"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:1990166">
        <property type="term" value="P:protein localization to site of double-strand break"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.10190:FF:000018">
        <property type="entry name" value="DNA topoisomerase 2-binding protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.10190">
        <property type="entry name" value="BRCT domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001357">
        <property type="entry name" value="BRCT_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036420">
        <property type="entry name" value="BRCT_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042479">
        <property type="entry name" value="Slf1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR057595">
        <property type="entry name" value="TopB1_SLF1_BRCT"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46677">
        <property type="entry name" value="SMC5-SMC6 COMPLEX LOCALIZATION FACTOR PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR46677:SF1">
        <property type="entry name" value="SMC5-SMC6 COMPLEX LOCALIZATION FACTOR PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF23294">
        <property type="entry name" value="BRCT_TopB1_SLF1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16770">
        <property type="entry name" value="RTT107_BRCT_5"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SMART" id="SM00292">
        <property type="entry name" value="BRCT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52113">
        <property type="entry name" value="BRCT domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="3"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="9" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="3" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="7" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="5" id="KW-0227">DNA damage</keyword>
    <keyword evidence="6" id="KW-0234">DNA repair</keyword>
    <keyword evidence="8" id="KW-0539">Nucleus</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4" id="KW-0677">Repeat</keyword>
    <feature type="domain" description="BRCT" evidence="10">
        <location>
            <begin position="3"/>
            <end position="80"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="9">
        <location>
            <begin position="665"/>
            <end position="698"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="9">
        <location>
            <begin position="699"/>
            <end position="731"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="9">
        <location>
            <begin position="733"/>
            <end position="765"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004300"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022763"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023204"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="SMART" id="SM00292"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000022305.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="917" mass="105896" checksum="C70A7853A90C2549" modified="2019-05-08" version="1">MACAVQKRRIQLTGFKKQEKEAIAELLLTLDCVFIDTEQYRNCTHLIAKQLSKSEKFLAACAAGKWILTKDYIINSAESGRWLDETTYEWGYKIEKDSHYSPQMQSAPKRWREELTCSGALGAFHRWKVILLVKEGDKRRDSITRVLEAGKAIIYPPQKEVRAVTHVFTDNKSVTIEREKCLFEAPYYPVQYIGDYLFEVNNEMNHLLAEQEARDTSDMQLVAMKNALIKHMYFTQVRKKQGYTFYVVFHKYLPFQVKNIDLCRGILCILEGLVEGHFFTDVITELAATQKYFIPPVQLLHSLLEHMLQGNTDTVFSAKFFHILYILLQLDPPWKSPSMLRYYLEFLQCPICMKGTWSLLEMLVRSCLYCKSFCHAVSVSEKKNEQRVVHKTLLKFFFNLVKAEVQALTRRLCEGANSQLLQVMPQTVLLKTFWLGSETSLLFTKYINILVDWVIHSYREKFKTNDAFKDEVAILLIGILGTVVEYWILSGFMMDRNVLHQVADDLANYIAISCDDFSTQELKMFICSIPSLWLQMFVAEAVFKNLCLRRNITISAEPLSLHKIVYSCLPALGEVGMHETGKVQKVKKKKIGQWPCPESQRALLMLNGDKQNQTKRLPDLPELTFNKFPPLSKKLRTKAEVEMSNTKENRHPLDEEMHFYKTNIKGETALHIACIRNKVERLIQLLPFPGTDINVKDYAGWTPLHEACNHGNTVCVREILQRCPEVDLLSQVDGVTPLHDALSNGHVEIGKLLLQRGGPLLLRQRDFYGKLPLDYVESLPVKQELLDIVHPKETVKDFCERVEKDFHNQQRELWLILFNKMLLNFCSVYNLSSPSPLSFKAHLNSLPLVAINGCKLKSRSLTDWLVDLYVGEVQTFQKLPEFIQEISGNLKKSFGEEIYPLLATLETIWLLITEYSY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="29" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452I458</accession>
    <name>A0A452I458_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="14">Dynamin-type G domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="15 16">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="16" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="15" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="15" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Mitochondrion outer membrane</location>
            <topology evidence="1">Multi-pass membrane protein</topology>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452I458"/>
    <dbReference type="Ensembl" id="ENSGAGT00000025436.1">
        <property type="protein sequence ID" value="ENSGAGP00000022321.1"/>
        <property type="gene ID" value="ENSGAGG00000016388.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005741">
        <property type="term" value="C:mitochondrial outer membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005525">
        <property type="term" value="F:GTP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003924">
        <property type="term" value="F:GTPase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008053">
        <property type="term" value="P:mitochondrial fusion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051646">
        <property type="term" value="P:mitochondrion localization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd09912">
        <property type="entry name" value="DLP_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.110:FF:000012">
        <property type="entry name" value="Mitofusin 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.300:FF:000214">
        <property type="entry name" value="Mitofusin 2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.110">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.300">
        <property type="entry name" value="P-loop containing nucleotide triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045063">
        <property type="entry name" value="Dynamin_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006884">
        <property type="entry name" value="Fzo/mitofusin_HR2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR030381">
        <property type="entry name" value="G_DYNAMIN_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027094">
        <property type="entry name" value="Mitofusin_fam"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10465:SF1">
        <property type="entry name" value="MITOFUSIN-2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10465">
        <property type="entry name" value="TRANSMEMBRANE GTPASE FZO1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00350">
        <property type="entry name" value="Dynamin_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF04799">
        <property type="entry name" value="Fzo_mitofusin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF111479">
        <property type="entry name" value="Fzo-like conserved region"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51718">
        <property type="entry name" value="G_DYNAMIN_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8 12" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="10" id="KW-0342">GTP-binding</keyword>
    <keyword evidence="5" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="11 13" id="KW-0472">Membrane</keyword>
    <keyword evidence="9" id="KW-0496">Mitochondrion</keyword>
    <keyword evidence="4" id="KW-1000">Mitochondrion outer membrane</keyword>
    <keyword evidence="3" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="16" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="2 13" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="7 13" id="KW-1133">Transmembrane helix</keyword>
    <keyword evidence="6" id="KW-0832">Ubl conjugation</keyword>
    <feature type="transmembrane region" description="Helical" evidence="13">
        <location>
            <begin position="577"/>
            <end position="595"/>
        </location>
    </feature>
    <feature type="domain" description="Dynamin-type G" evidence="14">
        <location>
            <begin position="93"/>
            <end position="342"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="12">
        <location>
            <begin position="648"/>
            <end position="685"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004374"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022692"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022787"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022989"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023134"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="14">
        <source>
            <dbReference type="PROSITE" id="PS51718"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="15">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000022321.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="16">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="705" mass="81087" checksum="31EF1350B914112C" modified="2019-05-08" version="1">MSLLFTRSKSIVAVKKDKRHMAEVNASPLKHFVTAKKKINGIFEQLAAYIQESSTFLEETYRNVELDPVTTEEQVLEVKGYLSKVSGISEVLARRHMKVAFFGRTSNGKSTVINAMLWDKVLPSGIGHTTNCFLRVEGTDGHDAFLLTEGSEEKRSVKTVNQLAHALHQNEFLHAGSLVSVMWPNSKCPLLKDDLVLMDSPGIDVTTELDSWIDKFCLDADVFVLVANSESTLMQTEKQFFHKVNERLSRPNIFILNNRWDASASEPEYMEEVRRQHMERCTSFLVDELGVVDRAQAGDRIFFVSAKEVLNARIQKAQGMPEGGGALAEGFQVRMFEFQNFERRFEECISQSAVKTKFEQHTVRAKQIAEDVRLIMDSLHIAAQEQRVYCLEMREERQERLGFIDKQLELLTQDYKMKIKQITEEVERQVSNAMAEEIRRLSVLVDDYQMDFHPSPVVLKVYKNELHRHIEEGLGRNMSDRCSNTITASLQTMQQEMIDGLKPLLPVSLRGQIDMLIPRQCFALSYDLNCDKLCADFQEDVEFHFSLGWTMLVNRFLGPKNSRRALMGYNDQVWKAVGWRLIALSFGLYGLLYVYERLTWTMKAKERAFKRQFVEYASEKLQLIVSYTGSNCSHQVQQELAGTFAHLCQQVDVTRENLEQEIAAMNKKIEILDSLQSKAKLLRNKAGWLDSELNMFTHQYLQQSR</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="23" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452I459</accession>
    <name>A0A452I459_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="4">TNFR-Cys domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="5 6">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="6" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="5" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="5" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="caution">
        <text evidence="1">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452I459"/>
    <dbReference type="Ensembl" id="ENSGAGT00000025443.1">
        <property type="protein sequence ID" value="ENSGAGP00000022327.1"/>
        <property type="gene ID" value="ENSGAGG00000016393.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030336">
        <property type="term" value="P:negative regulation of cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0010972">
        <property type="term" value="P:negative regulation of G2/M transition of mitotic cell cycle"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd13409">
        <property type="entry name" value="TNFRSF8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.50.10">
        <property type="entry name" value="Tumor Necrosis Factor Receptor, subunit A, domain 2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR031466">
        <property type="entry name" value="MIIP"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001368">
        <property type="entry name" value="TNFR/NGFR_Cys_rich_reg"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052862">
        <property type="entry name" value="TNFR_superfamily_member_8"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034002">
        <property type="entry name" value="TNFRSF8_N"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47497">
        <property type="entry name" value="TUMOR NECROSIS FACTOR RECEPTOR SUPERFAMILY MEMBER 8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47497:SF1">
        <property type="entry name" value="TUMOR NECROSIS FACTOR RECEPTOR SUPERFAMILY MEMBER 8"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF15734">
        <property type="entry name" value="MIIP"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00020">
        <property type="entry name" value="TNFR_c6"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00208">
        <property type="entry name" value="TNFR"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57586">
        <property type="entry name" value="TNF receptor-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50050">
        <property type="entry name" value="TNFR_NGFR_2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3" id="KW-0472">Membrane</keyword>
    <keyword evidence="6" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0812">Transmembrane</keyword>
    <keyword evidence="3" id="KW-1133">Transmembrane helix</keyword>
    <feature type="transmembrane region" description="Helical" evidence="3">
        <location>
            <begin position="668"/>
            <end position="689"/>
        </location>
    </feature>
    <feature type="repeat" description="TNFR-Cys" evidence="1">
        <location>
            <begin position="436"/>
            <end position="478"/>
        </location>
    </feature>
    <feature type="domain" description="TNFR-Cys" evidence="4">
        <location>
            <begin position="436"/>
            <end position="478"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="23"/>
            <end position="75"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="2">
        <location>
            <begin position="120"/>
            <end position="176"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="2">
        <location>
            <begin position="23"/>
            <end position="32"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="1">
        <location>
            <begin position="437"/>
            <end position="452"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00206"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="SAM" id="Phobius"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="4">
        <source>
            <dbReference type="PROSITE" id="PS50050"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="5">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000022327.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="883" mass="97662" checksum="39363AA8945B4B69" modified="2019-05-08" version="1">MQKMESEQLERLRHANQDLLERLKAKQEEIKKRLPRKPLSASSLSRRTTPAERAVPVPKRGKENQVCGMKGTTDPVTLVSVEPGASAARAALRSPLKPAVCGREGRGVLHCRAGAHPDSLRAERSFTPAASITRETSRVARDGSALRSPEEESSPVEHRESRKQSTMLHGAQERGRPWAEAEMALGRTPAEGSGRQQTAIRDSRTPKSILLTPGGKEAKKEAGRVTFLSDPEEYTIPANSWSARPFLGYDWIAGLLETDSSLSEKSEQYFTELREFRQVNKEACVHEEALGPEAVDFLAPEQEADLDSSSHQCVYCYRINRRLFTIPVDSQSACPVCKTLRARRPPETLEEPAYIRVSIPRSTLLPAYKYKIHRRKSFEPADDLALPSTLITSPQSCNVLENWLYDETSQRCCYQCPSGYHKRKECPTDPDRDCGQCEPEHFLNREQGKPHCKLCVSCKPEYNLVEKVPCSFNSRRVCECRAGLFCKLAIQSSCVQCISHTVCKPGFGVLARGTSEVDVSCQECPSGTFSDRHSSTERCKPHTDCAKLNKVATDSGNSTHDQVCMDPPPTHLTPASLSAKFSNRPKDSDRAEVPLLTEQPMTVRGQHFHTATKPVTSASILMSDAASDDANWTTAGTVDGSDTKHAAGTSPTSDIATEEIAKTGDGSFVPWGAVFLCSMVLLAGMLAVWKRRVCKRRIFTPNGKTLDLYPVTTCRPKSSSQGSDLMKNCAKRFSMLKADKESEERELLNRTPIVETNNNLVSISEKNPGPDSSLTEVIQSTGNATDYPADSRVRDHTNNRIDKIYIMKADTVIVGSISEVPVSKNCAVRGCENDAKAEENMEENELAMHYPQQETESFPGSDVMIPVEEEGKEFHHPTTATEK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="34" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IDU6</accession>
    <name>A0A452IDU6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">phosphatidylinositol-4,5-bisphosphate 3-kinase</fullName>
            <ecNumber evidence="2">2.7.1.153</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="11 12">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="12" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="11" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="11" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="catalytic activity">
        <reaction evidence="5">
            <text>a 1,2-diacyl-sn-glycero-3-phospho-(1D-myo-inositol-4,5-bisphosphate) + ATP = a 1,2-diacyl-sn-glycero-3-phospho-(1D-myo-inositol-3,4,5-trisphosphate) + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:21292"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:57836"/>
            <dbReference type="ChEBI" id="CHEBI:58456"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.1.153"/>
        </reaction>
        <physiologicalReaction direction="left-to-right" evidence="5">
            <dbReference type="Rhea" id="RHEA:21293"/>
        </physiologicalReaction>
    </comment>
    <comment type="pathway">
        <text evidence="1">Phospholipid metabolism; phosphatidylinositol phosphate biosynthesis.</text>
    </comment>
    <comment type="similarity">
        <text evidence="6">Belongs to the PI3/PI4-kinase family.</text>
    </comment>
    <dbReference type="EC" id="2.7.1.153" evidence="2"/>
    <dbReference type="AlphaFoldDB" id="A0A452IDU6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000029624.1">
        <property type="protein sequence ID" value="ENSGAGP00000026048.1"/>
        <property type="gene ID" value="ENSGAGG00000019027.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00220"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005943">
        <property type="term" value="C:phosphatidylinositol 3-kinase complex, class IA"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005944">
        <property type="term" value="C:phosphatidylinositol 3-kinase complex, class IB"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016303">
        <property type="term" value="F:1-phosphatidylinositol-3-kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046934">
        <property type="term" value="F:1-phosphatidylinositol-4,5-bisphosphate 3-kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-EC"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035005">
        <property type="term" value="F:1-phosphatidylinositol-4-phosphate 3-kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016477">
        <property type="term" value="P:cell migration"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043491">
        <property type="term" value="P:phosphatidylinositol 3-kinase/protein kinase B signal transduction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048015">
        <property type="term" value="P:phosphatidylinositol-mediated signaling"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd08399">
        <property type="entry name" value="C2_PI3K_class_I_gamma"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.70:FF:000006">
        <property type="entry name" value="Phosphatidylinositol 4,5-bisphosphate 3-kinase catalytic subunit gamma isoform"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.40.150:FF:000087">
        <property type="entry name" value="Phosphatidylinositol 4,5-bisphosphate 3-kinase catalytic subunit gamma isoform"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.10.20.770:FF:000001">
        <property type="entry name" value="Phosphatidylinositol 4,5-bisphosphate 3-kinase catalytic subunit gamma isoform"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.10.20.770">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.40.150">
        <property type="entry name" value="C2 domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.70">
        <property type="entry name" value="Phosphatidylinositol 3-kinase, accessory domain (PIK)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR016024">
        <property type="entry name" value="ARM-type_fold"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035892">
        <property type="entry name" value="C2_domain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015433">
        <property type="entry name" value="PI3/4_kinase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002420">
        <property type="entry name" value="PI3K-type_C2_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR003113">
        <property type="entry name" value="PI3K_ABD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001263">
        <property type="entry name" value="PI3K_accessory_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR042236">
        <property type="entry name" value="PI3K_accessory_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000341">
        <property type="entry name" value="PI3K_Ras-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045580">
        <property type="entry name" value="PIK3CG_ABD"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR029071">
        <property type="entry name" value="Ubiquitin-like_domsf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10048:SF34">
        <property type="entry name" value="PHOSPHATIDYLINOSITOL 4,5-BISPHOSPHATE 3-KINASE CATALYTIC SUBUNIT GAMMA ISOFORM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10048">
        <property type="entry name" value="PHOSPHATIDYLINOSITOL KINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00792">
        <property type="entry name" value="PI3K_C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00794">
        <property type="entry name" value="PI3K_rbd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00613">
        <property type="entry name" value="PI3Ka"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19710">
        <property type="entry name" value="PIK3CG_ABD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00142">
        <property type="entry name" value="PI3K_C2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00144">
        <property type="entry name" value="PI3K_rbd"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00145">
        <property type="entry name" value="PI3Ka"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48371">
        <property type="entry name" value="ARM repeat"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49562">
        <property type="entry name" value="C2 domain (Calcium/lipid-binding domain, CaLB)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54236">
        <property type="entry name" value="Ubiquitin-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51547">
        <property type="entry name" value="C2_PI3K"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51544">
        <property type="entry name" value="PI3K_ABD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51546">
        <property type="entry name" value="PI3K_RBD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51545">
        <property type="entry name" value="PIK_HELICAL"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="4" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="3" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="12" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="PI3K-ABD" evidence="7">
        <location>
            <begin position="34"/>
            <end position="139"/>
        </location>
    </feature>
    <feature type="domain" description="PI3K-RBD" evidence="9">
        <location>
            <begin position="215"/>
            <end position="307"/>
        </location>
    </feature>
    <feature type="domain" description="C2 PI3K-type" evidence="10">
        <location>
            <begin position="355"/>
            <end position="522"/>
        </location>
    </feature>
    <feature type="domain" description="PIK helical" evidence="8">
        <location>
            <begin position="542"/>
            <end position="715"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004805"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00012010"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023981"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00880"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS51544"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS51545"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="PROSITE" id="PS51546"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS51547"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000026048.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="715" mass="82878" checksum="A68A1150016D81D8" modified="2019-05-08" version="1">MELGNYEQPVVMREEAKRRRRRMKPHSPASNLSSVELISIEFILPTSNKQVQETMLLEIAGNCTVEQMKAQIWMRAIEMSQTTDFYHKFPPDQFILQYQKKGQWYEIYDKHQVIQTLDCILYWKVLEKKVGKISVIQKQKPAQAVQEFQTQLNDLIGYDVTDVSNVHDDELEFTRRRLVTPRMIEVACRDPKLYSMHPWITSKPLPEYVLKKITNNNIFIIIHRGTTSQKIKVSIDDTPDMILQSFFTKMAKKKSLMDIPEDHSELDFVLRVCGRDEYITGETPIKDFHWIRQCLKNGEEIHLVLDDPPDPKEDEVQKEEWPLVDDCTGVTGYHEQLTIDGKDHERVFTISLWDCNRKFRVKIVGIDIPVLPRNTELTVFVEANIQHGQQLLSQRRTISKPFTEEVLWNTWVEFDIKIKDLPKGALLNLQIYCGKAQALSTKVNLQSHDSPNSDSKCKTQLLYYVNILLIDHRSLLRSGEYVLHMWKISGKGEEQGSINADKLTSATNPDKENSMAISIVLDKYCHPIALPKHRITSDPQGDRTRAEMPNQLRKQLEAIIDTDPLNPLTPEDKELLWHFRYESIKHPKAYPKLLSSVKWGQQEIVAKTYQLLAKREAWDHSSLDVGLTMQLLDCNFSDENVRAMAVQKLESLEDDDVLHYLLQLVQAVKFEPYHDSALARFLLKRGLRVSTSKLNNIKVTGIVKIDCFSGENHVY</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IDU9</accession>
    <name>A0A452IDU9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">RAVE complex protein Rav1 C-terminal domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000029432.1">
        <property type="protein sequence ID" value="ENSGAGP00000025874.1"/>
        <property type="gene ID" value="ENSGAGG00000018632.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043291">
        <property type="term" value="C:RAVE complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007035">
        <property type="term" value="P:vacuolar acidification"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="2.130.10.10:FF:000150">
        <property type="entry name" value="Dmx-like 2, isoform CRA_c"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.130.10.10">
        <property type="entry name" value="YVTN repeat-like/Quinoprotein amine dehydrogenase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052208">
        <property type="entry name" value="DmX-like/RAVE_component"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022033">
        <property type="entry name" value="Rav1p_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015943">
        <property type="entry name" value="WD40/YVTN_repeat-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036322">
        <property type="entry name" value="WD40_repeat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001680">
        <property type="entry name" value="WD40_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13950:SF12">
        <property type="entry name" value="DMX-LIKE PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13950">
        <property type="entry name" value="RABCONNECTIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12234">
        <property type="entry name" value="Rav1p_C"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00400">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00320">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50978">
        <property type="entry name" value="WD40 repeat-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="1" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0677">Repeat</keyword>
    <keyword evidence="2" id="KW-0853">WD repeat</keyword>
    <feature type="domain" description="RAVE complex protein Rav1 C-terminal" evidence="5">
        <location>
            <begin position="1120"/>
            <end position="1399"/>
        </location>
    </feature>
    <feature type="domain" description="RAVE complex protein Rav1 C-terminal" evidence="5">
        <location>
            <begin position="1449"/>
            <end position="1664"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="419"/>
            <end position="462"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="662"/>
            <end position="696"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="4">
        <location>
            <begin position="445"/>
            <end position="458"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="4">
        <location>
            <begin position="669"/>
            <end position="696"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022574"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="Pfam" id="PF12234"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000025874.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1728" mass="192821" checksum="4E55B171212C0DB3" modified="2019-05-08" version="1">MNLHQVLTGAVNPGDNCFSVGSLHDQPLTAYASGCDIVILGTDFERLQIIPGAKHGNIQVGCVDCSMQQGKIAASYGNMICIFEPVTVLKQKSNHPGMYYQWQKSGQILLDSIAHNITWDPTGTRLLTGSNCLRLWSNVNLENQSEDGNPERTDVSFGDWRCMWQCKTASQVYLMKFSPDGEFFATAGKDDCLVKVWYNTDSWRSAVTPQSSSPQKQAQEVDFSFVYLAHPRSVNGFSWRKTSKYMPRGAVCNVLLTCCKDNVCRLWAETLLPNDSLSCAGGYNHWTEPVNLTNNFKRNASSKERMQNALELNLRPFRRGRRRSLALIAHTGYSPHQQDPHEVHRNTPLHANALCHFHIAASINPATDIPLLPSITSLSLSENEEKSGPFVVHWLNNKELHFTLSVEVFLQHLKKSFEQHSSETSTEESNQLDGKSDEEAADSGEEQKGNQEKKELSSEKSIPNSGFVPLSSAVIDHEIEILLSEWSKNADMLFSIHPMDGSLLVWHVDWLDEYQPGMFRQVQVSFVSRIPVAFPTGDANSLCRSIVMYACTKNVDLAIQQGKQKPLGLTRSSSMLISSAHNKSTNSLKLSIFTPNVMMISKHADGSLNQWLVSFAEESAFSTVLSISHKSRYCGHRFHLNDLACHSVLPLLLTTSHHNALRSPDVDSTKQTNDTSNSGESPLRSQNRDTTSVTSQDPNAVYSELILWRVDPVGPLSFSGGVSELARINSLHVSAFSNVAWLPTLIPSYCLGAYCNSPSACFVASDGQYLRLYQAVIDAKKLLCELSNPEISKYVGEVFNIISQQSTARPGCIVELDAITELHGSKTQLLHVFQEDFILNNNEEKIPEKNNCLSDSGNLQNGFSEKFYLIVIEYTQNRSLLHMWNLHLKSIPVSVDEKIDSNIDEAVSHHEERYSSSLDPEKIQSPFTQKYRACRANLQSTSRLTLSSEMVYSQELNLPEGVEIISVKPSAGHLSSSSVYPVCYAPYLLATSCSDGKVRFWRCRVNPEAVISSMPEYTINSNVMYIWEEWPLLVEDGLLSSSTVNVPGRPIEVSCAHTNRLAVAYKQMTSKNSRCSQDFMMHVSIFECESTGGSCWILEQTLHLDEISIMLDSAISIDSNLMAYNKQEVYLSNNERILPSTKHLVHLDWMSKEDGSHILTVGIGSKLYMFGQLVGNMHDQSSKEMSALPLWENTKVTSFCRFVLLRCVDLVSSVEGSPPLPVSLSWVRDGILVVGMDCEMHVYSQWQSPSKQESVMSDSYSGSTPSITSLMKQSQSAGSGLHPPKRTLTRSMTSLAQKLSGKRTAFDLSVEMEDSGLFEAAHALCPTLPQYHPFQLLELMDLGKVRRAKAILSHLVKCIAGEVVAINESESNHDRRLRSLTISASGSTARDPKTFNKTENTDYTEIDSVPPLPLYALLAADDDCSHFCLEKSNNQNNLNKQNGIPDENYEDLFQNSSLNTDDLTVFEADEDNTKPKIIDLSQYNPTYFGPEHAQVLSRHLLHSSLPGLTRMEQMSLMALADTIATTSTDIGESRDRNQGGETLDECGLKFLLAVQLHTFLTTSLPPVYRAQLLHQGLSTRHFAWAFHSVAEEELLNMLPAVQKGDPTWSELRAMGVGWWVRNIHSLRRCIEKVAKAAFQRNNDPLDAAIFYLAMKKKAVIWGLYSPFDKNNLSPLLFKSSKKKVPNIPFTDACTLVIIMCSEVYSFLFFTGSCHTSTLAKNVLIFFGL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IDV0</accession>
    <name>A0A452IDV0_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="12">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="12 13">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="13" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="12" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="12" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IDV0"/>
    <dbReference type="Ensembl" id="ENSGAGT00000029629.1">
        <property type="protein sequence ID" value="ENSGAGP00000026053.1"/>
        <property type="gene ID" value="ENSGAGG00000019029.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0035102">
        <property type="term" value="C:PRC1 complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003682">
        <property type="term" value="F:chromatin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003677">
        <property type="term" value="F:DNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042393">
        <property type="term" value="F:histone binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045892">
        <property type="term" value="P:negative regulation of DNA-templated transcription"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="CDD" id="cd09577">
        <property type="entry name" value="SAM_Ph1_2_3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.150.50:FF:000011">
        <property type="entry name" value="Polyhomeotic-like protein 2 isoform 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.60.160">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.150.50">
        <property type="entry name" value="Transcription Factor, Ets-1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR050548">
        <property type="entry name" value="PcG_chromatin_remod_factors"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001660">
        <property type="entry name" value="SAM"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013761">
        <property type="entry name" value="SAM/pointed_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR012313">
        <property type="entry name" value="Znf_FCS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR038603">
        <property type="entry name" value="Znf_FCS_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12247">
        <property type="entry name" value="POLYCOMB GROUP PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR12247:SF20">
        <property type="entry name" value="POLYHOMEOTIC-LIKE PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16616">
        <property type="entry name" value="PHC2_SAM_assoc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00536">
        <property type="entry name" value="SAM_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF21319">
        <property type="entry name" value="zf-FCS_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00454">
        <property type="entry name" value="SAM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF47769">
        <property type="entry name" value="SAM/Pointed domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50105">
        <property type="entry name" value="SAM_DOMAIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51024">
        <property type="entry name" value="ZF_FCS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="6" id="KW-0238">DNA-binding</keyword>
    <keyword evidence="3" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="7" id="KW-0539">Nucleus</keyword>
    <keyword evidence="13" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0862">Zinc</keyword>
    <keyword evidence="4 8" id="KW-0863">Zinc-finger</keyword>
    <feature type="domain" description="FCS-type" evidence="11">
        <location>
            <begin position="731"/>
            <end position="765"/>
        </location>
    </feature>
    <feature type="domain" description="SAM" evidence="10">
        <location>
            <begin position="880"/>
            <end position="944"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="1"/>
            <end position="28"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="237"/>
            <end position="256"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="372"/>
            <end position="454"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="509"/>
            <end position="533"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="577"/>
            <end position="650"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="705"/>
            <end position="729"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="9">
        <location>
            <begin position="788"/>
            <end position="870"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="1"/>
            <end position="15"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="372"/>
            <end position="388"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="9">
        <location>
            <begin position="389"/>
            <end position="399"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="400"/>
            <end position="428"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="509"/>
            <end position="531"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="9">
        <location>
            <begin position="612"/>
            <end position="625"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="9">
        <location>
            <begin position="634"/>
            <end position="650"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="9">
        <location>
            <begin position="711"/>
            <end position="723"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022771"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023125"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00367"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="10">
        <source>
            <dbReference type="PROSITE" id="PS50105"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="11">
        <source>
            <dbReference type="PROSITE" id="PS51024"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="12">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000026053.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="13">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="944" mass="99671" checksum="7417181C38A457E1" modified="2019-05-08" version="1">METESEQNSNSTNGSSGSGGSTRPQISQMSLYERQAVQALQALQRQPNAAQYFHQFMLQQQLNSAQLHSLAAVQQATIAASRQASSPNTSNPQQTTTTQASINLATTSAAQLISRSQSVSSPSATTLTQSVLLGNTTSPPLNQSQAQMYLRVQNLAVRSQQTSVANAQLQSSAQKAALPGNSQTSGLPQATSASQTLAVAQASSGNTAQSLNLSQGATGSNGVSGGVVAGAGSQATTGVGQAASSGPGGSCQRKGTGVVQPLPVAAAQAVTVSQGSQTETENAAATKKAEADGGQQTVGMNLTRTATPAPSQTLISSATYTQIQPHSLIQQQQQIHLQKQVVIQQQIAIHHQQQFQHRQSQLLHTATHLQLAQQQQQQASPLTQQQQQAPPPQQQPSPPQNQQQAQTLVVQPMLQSQPQPIQLQPDSPCQTATKSPVPIQSKPPSAPIKPPQLGAAKVSAAQQPPPHIPVQVVGSRQQGSAQAQALGLAQITPTASAPRGMPAVVQPISQARATSPSSQAPATASPQEAPPLTTGVNLAQVQGTAHVVKSTASSPVMAQVPAAFYMQPVQLPGKPQTLAVKRKAESEEEKEESPSATTLLPGKSSPVAESPKTMEEKSGFGEKSEPVPSTTPNAPTSETASVTTTSSPAPTLAMVSRQTGDSKPPQAIVKPQILTHIIEGFVIQEGAEPFPVGCSQLLKESEKLLQGGAPSGQSENQSSNSPGGDSASLELDKKTNLLKCEYCGKYAPANQFRGSKRFCSMTCAKRYNVSCSHQFRLQRKKMKEFQEANYARVRRRGPRRSSSEIARAKIQGKRHRGQEDSSRGSDNSSYDEALSPTSPGPLSMRASHGERDLTNSNMAPPTPDLHGINPDFLSSNPSRWSVEEVYEFIASLQGCQEIAEEFRSQEIDGQALLLLKEEHLMSAMNIKLGPALKICAKINILKET</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IDV4</accession>
    <name>A0A452IDV4_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">RAVE complex protein Rav1 C-terminal domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="3 4">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="4" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="3" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="3" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000029437.1">
        <property type="protein sequence ID" value="ENSGAGP00000025879.1"/>
        <property type="gene ID" value="ENSGAGG00000018632.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0043291">
        <property type="term" value="C:RAVE complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007035">
        <property type="term" value="P:vacuolar acidification"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR052208">
        <property type="entry name" value="DmX-like/RAVE_component"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022033">
        <property type="entry name" value="Rav1p_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036322">
        <property type="entry name" value="WD40_repeat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001680">
        <property type="entry name" value="WD40_rpt"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13950:SF12">
        <property type="entry name" value="DMX-LIKE PROTEIN 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13950">
        <property type="entry name" value="RABCONNECTIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12234">
        <property type="entry name" value="Rav1p_C"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00320">
        <property type="entry name" value="WD40"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50978">
        <property type="entry name" value="WD40 repeat-like"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="4" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="RAVE complex protein Rav1 C-terminal" evidence="2">
        <location>
            <begin position="935"/>
            <end position="1214"/>
        </location>
    </feature>
    <feature type="domain" description="RAVE complex protein Rav1 C-terminal" evidence="2">
        <location>
            <begin position="1264"/>
            <end position="1467"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="234"/>
            <end position="277"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="1">
        <location>
            <begin position="477"/>
            <end position="511"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="1">
        <location>
            <begin position="260"/>
            <end position="273"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="1">
        <location>
            <begin position="484"/>
            <end position="511"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="2">
        <source>
            <dbReference type="Pfam" id="PF12234"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="3">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000025879.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="4">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1532" mass="171293" checksum="95427771DD546925" modified="2019-05-08" version="1">RCFPPDFNFCVWYNTDSWRSAVTPQSSSPQKQAQEVDFSFVYLAHPRSVNGFSWRKTSKYMPRGAVCNVLLTCCKDNVCRLWAETLLPNDSLSCAGGYNHWTEPVNLTNNFKRNASSKERMQNALELNLRPFRRGRRRSLALIAHTGYSPHQQDPHEVHRNTPLHANALCHFHIAASINPATDIPLLPSITSLSLSENEEKSGPFVVHWLNNKELHFTLSVEVFLQHLKKSFEQHSSETSTEESNQLDGKSDEEAADSGEEQKGNQEKKELSSEKSIPNSGFVPLSSAVIDHEIEILLSEWSKNADMLFSIHPMDGSLLVWHVDWLDEYQPGMFRQVQVSFVSRIPVAFPTGDANSLCRSIVMYACTKNVDLAIQQGKQKPLGLTRSSSMLISSAHNKSTNSLKLSIFTPNVMMISKHADGSLNQWLVSFAEESAFSTVLSISHKSRYCGHRFHLNDLACHSVLPLLLTTSHHNALRSPDVDSTKQTNDTSNSGESPLRSQNRDTTSVTSQDPNAVYSELILWRVDPVGPLSFSGGVSELARINSLHVSAFSNVAWLPTLIPSYCLGAYCNSPSACFVASDGQYLRLYQAVIDAKKLLCELSNPEISKYVGEVFNIISQQSTARPGCIVELDAITELHGSKTQLLHVFQEDFILNNNEEKIPEKNNCLSDSGNLQNGFSEKFYLIVIEYTQNRSLLHMWNLHLKSIPVSVDEKIDSNIDEAVSHHEERYSSSLDPEKIQSPFTQKYRACRANLQSTSRLTLSSEMVYSQELNLPEGVEIISVKPSAGHLSSSSVYPVCYAPYLLATSCSDGKVRFWRCRVNPEAVISSMPEYTINSNVMYIWEEWPLLVEDGLLSSSTVNVPGRPIEVSCAHTNRLAVAYKQMTSKNSRCSQDFMMHVSIFECESTGGSCWILEQTLHLDEISIMLDSAISIDSNLMAYNKQEVYLSNNERILPSTKHLVHLDWMSKEDGSHILTVGIGSKLYMFGQLVGNMHDQSSKEMSALPLWENTKVTSFCRFVLLRCVDLVSSVEGSPPLPVSLSWVRDGILVVGMDCEMHVYSQWQSPSKQESVMSDSYSGSTPSITSLMKQSQSAGSGLHPPKRTLTRSMTSLAQKLSGKRTAFDLSVEMEDSGLFEAAHALCPTLPQYHPFQLLELMDLGKVRRAKAILSHLVKCIAGEVVAINESESNHDRRLRSLTISASGSTARDPKTFNKTENTDYTEIDSVPPLPLYALLAADDDCSHFCLEKSNNQNNLNKQNGIPDENYEDLFQNSSLNTDDLTVFEADEDNTKPKIIDLSQYNPTYFGPEHAQVLSRHLLHSSLPGLTRMEQMSLMALADTIATTSTDIGESRDRNQGGETLDECGLKFLLAVQLHTFLTTSLPPVYRAQLLHQGLSTRHFAWAFHSVAEEELLNMLPAVQKGDPTWSELRAMGVGWWVRNIHSLRRCIEKVAKAAFQRNNDPLDAAIFYLETPFDKNNLSPLLFKSSKKKVPNIPFTDACTLVIIMCSEVYSFLFFTGSCHTSTLAKNVLIFFGL</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IDV9</accession>
    <name>A0A452IDV9_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="2">non-specific serine/threonine protein kinase</fullName>
            <ecNumber evidence="2">2.7.11.1</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="9 10">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="10" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="9" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="9" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the protein kinase superfamily. STE Ser/Thr protein kinase family. STE20 subfamily.</text>
    </comment>
    <dbReference type="EC" id="2.7.11.1" evidence="2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000029591.1">
        <property type="protein sequence ID" value="ENSGAGP00000026016.1"/>
        <property type="gene ID" value="ENSGAGG00000019004.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004674">
        <property type="term" value="F:protein serine/threonine kinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.510.10:FF:000003">
        <property type="entry name" value="TRAF2 and NCK-interacting protein kinase isoform 4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.200.20">
        <property type="entry name" value="Phosphorylase Kinase, domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.510.10">
        <property type="entry name" value="Transferase(Phosphotransferase) domain 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001180">
        <property type="entry name" value="CNH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011009">
        <property type="entry name" value="Kinase-like_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000719">
        <property type="entry name" value="Prot_kinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008271">
        <property type="entry name" value="Ser/Thr_kinase_AS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051700">
        <property type="entry name" value="STE20_Ser-Thr_kinase"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47096">
        <property type="entry name" value="MISSHAPEN LIKE KINASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR47096:SF1">
        <property type="entry name" value="MISSHAPEN LIKE KINASE 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00780">
        <property type="entry name" value="CNH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00069">
        <property type="entry name" value="Pkinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00036">
        <property type="entry name" value="CNH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00220">
        <property type="entry name" value="S_TKc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56112">
        <property type="entry name" value="Protein kinase-like (PK-like)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50219">
        <property type="entry name" value="CNH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50011">
        <property type="entry name" value="PROTEIN_KINASE_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00108">
        <property type="entry name" value="PROTEIN_KINASE_ST"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5" id="KW-0418">Kinase</keyword>
    <keyword evidence="10" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0723">Serine/threonine-protein kinase</keyword>
    <keyword evidence="4" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Protein kinase" evidence="7">
        <location>
            <begin position="1"/>
            <end position="234"/>
        </location>
    </feature>
    <feature type="domain" description="CNH" evidence="8">
        <location>
            <begin position="1336"/>
            <end position="1623"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="221"/>
            <end position="292"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="342"/>
            <end position="399"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="488"/>
            <end position="511"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="542"/>
            <end position="618"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="829"/>
            <end position="880"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="6">
        <location>
            <begin position="896"/>
            <end position="1164"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="233"/>
            <end position="252"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="6">
        <location>
            <begin position="262"/>
            <end position="280"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="492"/>
            <end position="507"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="544"/>
            <end position="558"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="567"/>
            <end position="576"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="598"/>
            <end position="615"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="897"/>
            <end position="908"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="6">
        <location>
            <begin position="923"/>
            <end position="941"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="949"/>
            <end position="966"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="6">
        <location>
            <begin position="1023"/>
            <end position="1052"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="1072"/>
            <end position="1086"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="6">
        <location>
            <begin position="1098"/>
            <end position="1116"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Acidic residues" evidence="6">
        <location>
            <begin position="1123"/>
            <end position="1136"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00008874"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00012513"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022527"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50011"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="PROSITE" id="PS50219"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000026016.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1649" mass="186330" checksum="200E88067C09FB11" modified="2019-05-08" version="1">MDVTGDEEEEIKQEINMLKKYSHHRNIATYYGAFIKKNPPGMDDQLWLVMEFCGAGSVTDLIKNTKGNTLKEEWIAYICREILRGLSHLHQHKVIHRDIKGQNVLLTENAEVKLVDFGVSAQLDRTVGRRNTFIGTPYWMAPEVIACDENPDATYDFKSDLWSLGITAIEMAEGAPPLCDMHPMRALFLIPRNPAPRLKSKKWSKKFQSFIESCLVKNHSQRPTTEQLMKHPFIRDQPNERQVRIQLKDHIDRTKKKRGEKDETEYEYSGSEEEEEENDSGEPSSILNLPGESTLRRDFLRLQLANKERSEALRRQQLEQQQRENEEHKRQLLAERQKRIEEQKEQRRRLEEQQRREKEMRKQQEREQRRHYEEQMRREEERRRAEHEQEYKRKQLEEQRQAERLQRQLQQERDYLVSLQQQQQQRQDQRPAEKKPLYHYKEGMNASEKPAWAKEVQQQYLNNSPVLPAKQNHSSVARHLAPFITGKDVAPQSNIKPSSNISKSQPSAKPNLQDLRALDPWQVNPLVSAKPRQVPPIIKISKPRSQEQQHTQKGEEAIWTHSQIRLGHQERSKKNDGVISLKGSRSQGCSPVRKSGLEVTSSSPHVSKGINSHLPSNGVKMDFKEHNGDRGRKFAHSLMPTSPLFPSRSTPSLNNIDLMCDHLDLSASSPEHILRKMKTQALHKELHASFESIPGLLPSTGIHRTQMQQKIGRSMDQLPLPSLGLSPQGSHWNEAILAPLSSCSRSSSVDLLSFPLNSNLIQASQVCRGNAPNSMSSRVAPGSFSNSSYCSPKLSPQVCNGKSFVNLPVIKTFLSSSTPDLRDVRQLNSEQPLVEERSRLNRQSSPAMPHKAANRISDPNLPPRSESFSISGVQPARTPPMLRPVDPQIPHLVTVKSQGTSLSGSQSLHEQPSKGMSGFQEALTVTSHRTEMPRQNSDPTSENPPLPPRIEKFDRSSWLRQEEDVPPKVPQRTTSISPALARKNSPGNGTALGARLGSQPIRASNPDLRRTEPIIESPLQRTSSGSSSSSSTPSSQPSSQEGSQPGSQAGSSERNRVRGNTRPEGSPVLSHELSKMKQEDSRDVTRPSRPADLTALAKELRELRIEETNRPLKKVTDYSSSSEESESSEEEEEDGESETHDGTVAVSDIPRLIPTGVPGSNEPYNLGMVASHGLETSNADTFSSISREGTLMVRETTGEKKRSSHAASNGFAGHINLPDLVQQSHSPAGTPTEALGRVSTHAQDMDSVAEYGMGSSTKASFTPFVDTRVYQTSPTDEDEDDDEESSATALFTSELLRQEQAKLNEARKISVVNVNPTNIRPHSDTPEIRKYKKRFNSEILCAALWGVNLLVGTENGLMLLDRSGQGKVYNLINRRRFQQMDVLEGLNVLVTISGKKNKLRVYYLSWLRNRILHNDPEVEKKQGWITVGDLEGCIHYKVVKYERIKFLVIALKNAVEIYAWAPKPYHKFMAFKSFADLQHKPLLVDLTVEEGQRLKVIFGSHTGFHVIDVDSGNSYDIYIPSHIQGNITPHAIVILPKTDGMEMLVCYEDEGVYVNTYGRITKDVVLQWGEMPTSVAYIHSNQIMGWGEKAIEIRSVETGHLDGVFMHKRAQRLKFLCERNDKVFFASVRSGGSSQVFFMTLNRNSMMNW</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IE60</accession>
    <name>A0A452IE60_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="12">Ankyrin repeat domain-containing protein 17</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="13">Gene trap ankyrin repeat protein</fullName>
        </alternativeName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000029776.1">
        <property type="protein sequence ID" value="ENSGAGP00000026181.1"/>
        <property type="gene ID" value="ENSGAGG00000018637.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045087">
        <property type="term" value="P:innate immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd22502">
        <property type="entry name" value="KH-I_ANKRD17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd22249">
        <property type="entry name" value="UDM1_RNF168_RNF169-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000046">
        <property type="entry name" value="Ankyrin repeat and KH domain-containing protein 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000114">
        <property type="entry name" value="ankyrin repeat and KH domain-containing protein 1 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000062">
        <property type="entry name" value="ankyrin repeat domain-containing protein 17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000012">
        <property type="entry name" value="ankyrin repeat domain-containing protein 17 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000131">
        <property type="entry name" value="ankyrin repeat domain-containing protein 17 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1370.10:FF:000031">
        <property type="entry name" value="ankyrin repeat domain-containing protein 17 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000014">
        <property type="entry name" value="ankyrin repeat domain-containing protein 17 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000161">
        <property type="entry name" value="ankyrin repeat domain-containing protein 17 isoform X3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1370.10">
        <property type="entry name" value="K Homology domain, type 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051631">
        <property type="entry name" value="Ankyrin-KH/SAM_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047375">
        <property type="entry name" value="KH-I_ANKRD17"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004087">
        <property type="entry name" value="KH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004088">
        <property type="entry name" value="KH_dom_type_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036612">
        <property type="entry name" value="KH_dom_type_1_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23206:SF1">
        <property type="entry name" value="ANKYRIN REPEAT DOMAIN-CONTAINING PROTEIN 17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23206">
        <property type="entry name" value="MASK PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00023">
        <property type="entry name" value="Ank"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="9"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00013">
        <property type="entry name" value="KH_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01415">
        <property type="entry name" value="ANKYRIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="24"/>
    </dbReference>
    <dbReference type="SMART" id="SM00322">
        <property type="entry name" value="KH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54791">
        <property type="entry name" value="Eukaryotic type KH-domain (KH-domain type I)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="19"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="19"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50084">
        <property type="entry name" value="KH_TYPE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8" id="KW-0007">Acetylation</keyword>
    <keyword evidence="9 14" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="10 16" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-0391">Immunity</keyword>
    <keyword evidence="4" id="KW-0399">Innate immunity</keyword>
    <keyword evidence="3" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="2" id="KW-0488">Methylation</keyword>
    <keyword evidence="11" id="KW-0539">Nucleus</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="15" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="6" id="KW-0832">Ubl conjugation</keyword>
    <feature type="repeat" description="ANK" evidence="14">
        <location>
            <begin position="155"/>
            <end position="187"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="14">
        <location>
            <begin position="188"/>
            <end position="220"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="14">
        <location>
            <begin position="221"/>
            <end position="253"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="14">
        <location>
            <begin position="255"/>
            <end position="287"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="14">
        <location>
            <begin position="288"/>
            <end position="320"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="14">
        <location>
            <begin position="321"/>
            <end position="353"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="14">
        <location>
            <begin position="354"/>
            <end position="386"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="14">
        <location>
            <begin position="418"/>
            <end position="450"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="14">
        <location>
            <begin position="451"/>
            <end position="483"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="14">
        <location>
            <begin position="484"/>
            <end position="516"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="14">
        <location>
            <begin position="518"/>
            <end position="550"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="14">
        <location>
            <begin position="939"/>
            <end position="971"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="14">
        <location>
            <begin position="972"/>
            <end position="1004"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="14">
        <location>
            <begin position="1006"/>
            <end position="1038"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="14">
        <location>
            <begin position="1039"/>
            <end position="1071"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="14">
        <location>
            <begin position="1100"/>
            <end position="1132"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="14">
        <location>
            <begin position="1133"/>
            <end position="1165"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="14">
        <location>
            <begin position="1168"/>
            <end position="1200"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="14">
        <location>
            <begin position="1201"/>
            <end position="1233"/>
        </location>
    </feature>
    <feature type="domain" description="K Homology" evidence="18">
        <location>
            <begin position="1540"/>
            <end position="1610"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1328"/>
            <end position="1387"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1400"/>
            <end position="1471"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1502"/>
            <end position="1532"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1691"/>
            <end position="1780"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="1798"/>
            <end position="1921"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="2067"/>
            <end position="2110"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="17">
        <location>
            <begin position="2144"/>
            <end position="2165"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="16">
        <location>
            <begin position="663"/>
            <end position="759"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="17">
        <location>
            <begin position="1330"/>
            <end position="1340"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="17">
        <location>
            <begin position="1341"/>
            <end position="1370"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="17">
        <location>
            <begin position="1451"/>
            <end position="1460"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="1502"/>
            <end position="1519"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="17">
        <location>
            <begin position="1735"/>
            <end position="1779"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="17">
        <location>
            <begin position="1798"/>
            <end position="1813"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="1842"/>
            <end position="1863"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="17">
        <location>
            <begin position="1873"/>
            <end position="1890"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="1895"/>
            <end position="1904"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="17">
        <location>
            <begin position="1910"/>
            <end position="1919"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="2067"/>
            <end position="2083"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="17">
        <location>
            <begin position="2099"/>
            <end position="2109"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="17">
        <location>
            <begin position="2146"/>
            <end position="2160"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022588"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022859"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00067263"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00081278"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00117"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="SMART" id="SM00322"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000026181.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2338" mass="248447" checksum="59FDBD2F7FE23C9E" modified="2019-05-08" version="1">MLETASKLLLSSTADGADLRTVDPETQARLEALLEAAGIGKLSTADGKAFADPEVLRRLTSSVSCALDEAAAALTRMRAESTANAGQTDNRSLAEACSEGDVNAVRKLLIEGRSVNEHTEEGESLLCLACSAGYYELAQVLLAMHANVEDRGIKGDITPLMAAANGGHVKIVKLLLAHGADVNAQSSTGNTALTYACAGGYVDVVKVLLESGASIEDHNENGHTPLMEAGSAGHVEVARVLLENGAGINTHSNEFKESALTLACYKGHLEMVRFLLEAGADQEHKTDEMHTALMEACMDGHVEVARLLLDSGAQVNMPADSFESPLTLAACGGHVELAALLIERGANLEEVNDEGYTPLMEAAREGHEEMVALLLGQGANINAQTEETQETALTLACCGGFLEVADFLIKAGADIELGCSTPLMEAAQEGHLELVKYLLAAGANVHATTATGDTALTYACENGHTDVADVLLQAGADLEHESEGGRTPLMKAARAGHVCTVQFLISKGANVNRTTANNDHTVLSLACAGGHLAVVELLLAHGADPTHRLKDGSTMLIEAAKGGHTSVVCYLLDYPNNLLSAPAPDATQLTPPSHDLNRAPRVPVQALPMVVPPQEPDKPPANVATTLPIRNKAASKQKSSTHLPANNQDVQGYITNQSPESIVEEAQGKLTELEQRIKEAIEKNAQLQSLELAHADQLTKEKIEELNKTREEQIQKKQKILEELQKVERELQLKTQQQLKKQYLEVKAQRIQLQQQQQQQQSCQHLGLLTPVGVGEQLSEGDYARLQQVDPILLKDDPQQPAMQMGFAPIQPLAMPQALPLATGSLPPGSIANLTELQGVIVGQPVLGQAQLAGLGQGILTETQQGLMVASPAQTLNDTLDDIMAAVSGRASAMSNTPTHSIATSVSQPQTPAPSPIISPSAMLPIYPAIDIDAQTESNHDTALTLACAGGHEELVQTLLERGANIEHRDKKGFTPLILAATAGHVGVVEILLDNGADIEAQSERTKDTPLSLACSGGRQEVVELLLARGANKEHRNVSDYTPLSLAASGGYVNIIKILLNAGAEINSRTGSKLAMNGHTAAVKLLLDMGSDINAQIETNRNTALTLACFQGRTEVVSLLLDRKANVEHRAKTGLTPLMEAASGGYAEVGRVLLDKGADVNAPPVPSSRDTALTIAADKGHYKFCELLISRGGHIDVRNKKGNTPLWLAANGGHLDVVQLLVQAGADVDAADNRKITPLMAAFRKGHVKVVRYLVKEVNQFPSDSECMRYIATITDKEMLKKCHLCMESIVQAKDRQAAEANKNASILLEELDLEKLREESRRLALAAKREKRKEKRRKKKEEQRRKLEEIEAKNKENFELQAAQEKEELKTEDEPEVLMEPPSATTTTTIGISATWTTLAGSHGKRNNTITTTSSKRKSRKNKINPDNVQILFDDQLPISYGQPEKANGESKSSSTSESGDSDNMRISSCSDESSNSSLLLCIFLRCSEPVSSFHEVTSNSLSTCTKSGPSPLSSPNGKLTIASPKRGQKREEGWKEVVRRSKKVSVPSTVISRVIGRGGCNINAIREFTGAHIDIDKQKDKTGDRIITIRGGTESTRQATQLINALIKDPDKEIDELIPKNRLKSSSTSQTATALTVPAISSAPTHKTVKNPVNNVRPGFPVSLPLAYPPPQFAHALLAAQTFQQIRPPRLPMTHFGGTFPPAQSTWGPFPVRPLSPARATNSPKPHMVPRHNSQNSSGSQVNSTSSLTTSLTTATSSTASAVPVSSTNGSPSSPSVRRQLFVTVVKTSNATTTTITTTASNNSTAPTNATYPIPTAKEHYPVSSPSSPSPPAQPVAVSRNSPSDCVTTSPNKGASSSDQEAASPPVVETSNLASSGQLSASSLGSSLVHPAHQQSPGSLSQEARLPPQQPQVPLPDPRMVVPPNLAAVTSSSAPVVGSTNAPMTYPLPQTSMGSTQPPAKMETPAIRPPAHGTNTGHKTPAPVQNCSVAILNVNHIKRPHSVPSSVQLPSTLNTQTASQNSAHPANKSVGTNFSATLPFGPFSTLFENSSTSAHAFWGGSVVSSQTTPESIRSGKSSFLPNSEPLHRSDTSKAPGFRPPLQRPAPSPSGKCVCVCVVSSAECLHFGTLNIFLSSGVRAPSPTPSAVSLGSEKPSNASQDRKVPVPIGTERSARIRQTGTSTPSVIGSNLATPVGHSGIWSFEGIGANQGMKYRLTDWCHSGMGTHMIHRPMSDPGVFSHQVMERDSTGIVTPSGTFHQHVPGMPFSVYGNAMIPPVAPIADGTGGPIFNGPHAADPSWNSLIKMVSNSTENNGPQTVWTGTWTPHMNSVHMNQLG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IE61</accession>
    <name>A0A452IE61_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="5">KOW domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="6 7">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="7" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="6" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="6" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="similarity">
        <text evidence="1">Belongs to the universal ribosomal protein uL24 family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IE61"/>
    <dbReference type="Ensembl" id="ENSGAGT00000029323.1">
        <property type="protein sequence ID" value="ENSGAGP00000025778.1"/>
        <property type="gene ID" value="ENSGAGG00000018815.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0015934">
        <property type="term" value="C:large ribosomal subunit"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003735">
        <property type="term" value="F:structural constituent of ribosome"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006412">
        <property type="term" value="P:translation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd06089">
        <property type="entry name" value="KOW_RPL26"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.30:FF:000009">
        <property type="entry name" value="60S ribosomal protein L26"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.30">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="HAMAP" id="MF_01326_A">
        <property type="entry name" value="Ribosomal_uL24_A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005824">
        <property type="entry name" value="KOW"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014722">
        <property type="entry name" value="Rib_uL2_dom2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005825">
        <property type="entry name" value="Ribosomal_uL24_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR005756">
        <property type="entry name" value="Ribosomal_uL24_euk/arc"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041988">
        <property type="entry name" value="Ribosomal_uL24_KOW"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008991">
        <property type="entry name" value="Translation_prot_SH3-like_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR01080">
        <property type="entry name" value="rplX_A_E"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11143">
        <property type="entry name" value="60S RIBOSOMAL PROTEIN L26 FAMILY MEMBER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00467">
        <property type="entry name" value="KOW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF16906">
        <property type="entry name" value="Ribosomal_L26"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SMART" id="SM00739">
        <property type="entry name" value="KOW"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50104">
        <property type="entry name" value="Translation proteins SH3-like domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01108">
        <property type="entry name" value="RIBOSOMAL_L24"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="7" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0687">Ribonucleoprotein</keyword>
    <keyword evidence="2" id="KW-0689">Ribosomal protein</keyword>
    <feature type="domain" description="KOW" evidence="5">
        <location>
            <begin position="48"/>
            <end position="76"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="1"/>
            <end position="21"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="4">
        <location>
            <begin position="122"/>
            <end position="145"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00010618"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022980"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023274"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="5">
        <source>
            <dbReference type="SMART" id="SM00739"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="6">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000025778.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="145" mass="17246" checksum="048CB25B5FDEB6B6" modified="2019-05-08" version="1">MKFNPFVTSDRSKNRKRHFNAPSHIRRKIMSSPLSKELRQKYNVRSMPIRKDDEVQVVRGHYKGQQIGKVVQVYRKKYVTYIERVQREKANGTTVHVGIHPSKVVITRLKLDKDRKKILERKAKSRQVGKEKGKYKEETIEKMQE</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IE62</accession>
    <name>A0A452IE62_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Peptidase S1 domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="8 9">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="9" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="8" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="8" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="AlphaFoldDB" id="A0A452IE62"/>
    <dbReference type="Ensembl" id="ENSGAGT00000029694.1">
        <property type="protein sequence ID" value="ENSGAGP00000026107.1"/>
        <property type="gene ID" value="ENSGAGG00000019075.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004252">
        <property type="term" value="F:serine-type endopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00190">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.40.10.10:FF:000120">
        <property type="entry name" value="Putative serine protease"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.40.10.10">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009003">
        <property type="entry name" value="Peptidase_S1_PA"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR043504">
        <property type="entry name" value="Peptidase_S1_PA_chymotrypsin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001314">
        <property type="entry name" value="Peptidase_S1A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001254">
        <property type="entry name" value="Trypsin_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018114">
        <property type="entry name" value="TRYPSIN_HIS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR033116">
        <property type="entry name" value="TRYPSIN_SER"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271:SF69">
        <property type="entry name" value="GRANZYME A"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24271">
        <property type="entry name" value="KALLIKREIN-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00089">
        <property type="entry name" value="Trypsin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00722">
        <property type="entry name" value="CHYMOTRYPSIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00020">
        <property type="entry name" value="Tryp_SPc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF50494">
        <property type="entry name" value="Trypsin-like serine proteases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50240">
        <property type="entry name" value="TRYPSIN_DOM"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00134">
        <property type="entry name" value="TRYPSIN_HIS"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00135">
        <property type="entry name" value="TRYPSIN_SER"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="5" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="3 6" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="1 6" id="KW-0645">Protease</keyword>
    <keyword evidence="9" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="4 6" id="KW-0720">Serine protease</keyword>
    <keyword evidence="2" id="KW-0732">Signal</keyword>
    <feature type="domain" description="Peptidase S1" evidence="7">
        <location>
            <begin position="34"/>
            <end position="259"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022825"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="RuleBase" id="RU363034"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="7">
        <source>
            <dbReference type="PROSITE" id="PS50240"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000026107.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="9">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="263" mass="29197" checksum="915B91DD806D2E08" modified="2019-05-08" version="1">MISQGPFPAYNSMILWQAWHWAAPGLFLDSCVDIIGGYAVTPHSRPYMALIKGKEYCGGALIKENWVLTAAHCKIKVVLGAHSLSKKEKEQQTFKIARPISFPCFNNQTKENDIMLLQLQQRAKINRAVKTIKIPNTDTDPKPGTKCQVAGWGITKNQERTPSDILREVNITIIDRKICNDQNHYNYNPMITMNMVCAGDENGGKDSCDGDSGGPLICNGELKGITSFGRAKKCGTPQYPGVYARLTKKHLLWIKQTIGGDID</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="33" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IE63</accession>
    <name>A0A452IE63_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="18">Microtubule-associated protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="20 21">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="21" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="20" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="20" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="3">Cell membrane</location>
            <topology evidence="3">Peripheral membrane protein</topology>
            <orientation evidence="3">Cytoplasmic side</orientation>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="4">Cell projection</location>
            <location evidence="4">Axon</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="2">Cell projection</location>
            <location evidence="2">Dendrite</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="1 18">Cytoplasm</location>
            <location evidence="1 18">Cytoskeleton</location>
        </subcellularLocation>
        <subcellularLocation>
            <location evidence="5">Cytoplasm</location>
            <location evidence="5">Cytosol</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000029561.1">
        <property type="protein sequence ID" value="ENSGAGP00000025990.1"/>
        <property type="gene ID" value="ENSGAGG00000018969.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030424">
        <property type="term" value="C:axon"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005829">
        <property type="term" value="C:cytosol"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030425">
        <property type="term" value="C:dendrite"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005874">
        <property type="term" value="C:microtubule"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008017">
        <property type="term" value="F:microtubule binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000226">
        <property type="term" value="P:microtubule cytoskeleton organization"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0031175">
        <property type="term" value="P:neuron projection development"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027324">
        <property type="entry name" value="MAP2/MAP4/Tau"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001084">
        <property type="entry name" value="MAP_tubulin-bd_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002955">
        <property type="entry name" value="Tau"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11501">
        <property type="entry name" value="MICROTUBULE-ASSOCIATED PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR11501:SF14">
        <property type="entry name" value="MICROTUBULE-ASSOCIATED PROTEIN TAU"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00418">
        <property type="entry name" value="Tubulin-binding"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01261">
        <property type="entry name" value="TAUPROTEIN"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00229">
        <property type="entry name" value="TAU_MAP_1"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51491">
        <property type="entry name" value="TAU_MAP_2"/>
        <property type="match status" value="4"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="14" id="KW-0007">Acetylation</keyword>
    <keyword evidence="6" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="17" id="KW-0966">Cell projection</keyword>
    <keyword evidence="8 18" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="16 18" id="KW-0206">Cytoskeleton</keyword>
    <keyword evidence="9" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="15" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0488">Methylation</keyword>
    <keyword evidence="11 18" id="KW-0493">Microtubule</keyword>
    <keyword evidence="10" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="21" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="12" id="KW-0677">Repeat</keyword>
    <keyword evidence="13" id="KW-0832">Ubl conjugation</keyword>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="29"/>
            <end position="69"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="262"/>
            <end position="302"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="317"/>
            <end position="352"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="423"/>
            <end position="667"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="19">
        <location>
            <begin position="765"/>
            <end position="843"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="57"/>
            <end position="67"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="271"/>
            <end position="284"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="292"/>
            <end position="302"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="452"/>
            <end position="469"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="482"/>
            <end position="501"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="19">
        <location>
            <begin position="502"/>
            <end position="515"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="516"/>
            <end position="530"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="19">
        <location>
            <begin position="551"/>
            <end position="561"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="19">
        <location>
            <begin position="562"/>
            <end position="580"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="19">
        <location>
            <begin position="594"/>
            <end position="605"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="19">
        <location>
            <begin position="606"/>
            <end position="626"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004245"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004279"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00004413"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00004489"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00004514"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022475"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022701"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023136"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023212"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="ARBA" id="ARBA00023273"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="RuleBase" id="RU000686"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000025990.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="21">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="843" mass="91526" checksum="2D6BA43CB1C69A6D" modified="2019-05-08" version="1">MKWNKRNRGRDNFCNRMTEQRQDFNMMEDHSMSQAKQIPSGYPLQIPVDDGSDEPISETSDAKSTPTMEDATAPLVEERAHEDRIAARQHVEIPEGTTEELSSPRLWESVEYGIQEHVASEIQPEKQAGEVLPSTVVPLCEMQTGEQAAAIPARAEATIPVPPGTYEDFRAFEESQEAVSRTELWDRGGKEDIGVGKALADEAGHDVCTEEFAASAVSEFSQSVPRAGMFYHEKRAAQFDKWHPGVDDVGYEPVGNSHIQDKASHLYSRGKYKEEAVGHEKDEDRDIDETIPQDSPSSLGQQDSPLLEIAEEAVKEHSFPAEGQPRDNLAEISRETPVTEREGHKAGQILDEKRQWFSGKGYDDVTKMEPSESMHQVEVEANIIREGEISGSPLDTTKFPDRLNKDVADRSVSVQQEMGGMVQAMEKQEAPKKKPATRVSEKQVSRVPHLKAHIDSKDKDGIDIEEKKPKTSTPSSAKPLTDRSSITPQRPSSVSTTPLKNPSSPAESSVPASTPKRVSSITSRPASTGTKETKPKGPEMKSGMKMAAPRSATQTQKSPANTTRIPAKTPTAPKTPPSAAGRKEQRKPPTTAAKSEKXEPAKSGDRSGYSSPGSPGTPGSRSRTPSLPTPPNREPKKVAVVRTPPKSPASAKSRLQTSTAPMPDLKNVRSKIGSTENLKHQPGGGKVQIMNKKLDLSNIQSKCGSKDNIKHSPGGGSVQIVYKPVDLSHVTSKCGSLGNIHHKPGGGQVEVKSEKLDFKEKVQSKIGSLDNITHVPGGGNKKREKGKDDERGAQNGVSQSAGPQALLTEPAIPEESPLPDLQPDGDANRVSQADLPRECQSQD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IE74</accession>
    <name>A0A452IE74_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="11">Ankyrin repeat domain-containing protein 17</fullName>
        </recommendedName>
        <alternativeName>
            <fullName evidence="12">Gene trap ankyrin repeat protein</fullName>
        </alternativeName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Nucleus</location>
        </subcellularLocation>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000029791.1">
        <property type="protein sequence ID" value="ENSGAGP00000026196.1"/>
        <property type="gene ID" value="ENSGAGG00000018637.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005737">
        <property type="term" value="C:cytoplasm"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005634">
        <property type="term" value="C:nucleus"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003723">
        <property type="term" value="F:RNA binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0045087">
        <property type="term" value="P:innate immune response"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd22502">
        <property type="entry name" value="KH-I_ANKRD17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd22249">
        <property type="entry name" value="UDM1_RNF168_RNF169-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000114">
        <property type="entry name" value="ankyrin repeat and KH domain-containing protein 1 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000062">
        <property type="entry name" value="ankyrin repeat domain-containing protein 17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000012">
        <property type="entry name" value="ankyrin repeat domain-containing protein 17 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000131">
        <property type="entry name" value="ankyrin repeat domain-containing protein 17 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.1370.10:FF:000031">
        <property type="entry name" value="ankyrin repeat domain-containing protein 17 isoform X1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000014">
        <property type="entry name" value="ankyrin repeat domain-containing protein 17 isoform X2"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000161">
        <property type="entry name" value="ankyrin repeat domain-containing protein 17 isoform X3"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.25.40.20:FF:000584">
        <property type="entry name" value="Uncharacterized protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.25.40.20">
        <property type="entry name" value="Ankyrin repeat-containing domain"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1370.10">
        <property type="entry name" value="K Homology domain, type 1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR051631">
        <property type="entry name" value="Ankyrin-KH/SAM_domain"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002110">
        <property type="entry name" value="Ankyrin_rpt"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036770">
        <property type="entry name" value="Ankyrin_rpt-contain_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR047375">
        <property type="entry name" value="KH-I_ANKRD17"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004087">
        <property type="entry name" value="KH_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004088">
        <property type="entry name" value="KH_dom_type_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036612">
        <property type="entry name" value="KH_dom_type_1_sf"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23206:SF1">
        <property type="entry name" value="ANKYRIN REPEAT DOMAIN-CONTAINING PROTEIN 17"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR23206">
        <property type="entry name" value="MASK PROTEIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00023">
        <property type="entry name" value="Ank"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="Pfam" id="PF12796">
        <property type="entry name" value="Ank_2"/>
        <property type="match status" value="8"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00013">
        <property type="entry name" value="KH_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01415">
        <property type="entry name" value="ANKYRIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00248">
        <property type="entry name" value="ANK"/>
        <property type="match status" value="22"/>
    </dbReference>
    <dbReference type="SMART" id="SM00322">
        <property type="entry name" value="KH"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48403">
        <property type="entry name" value="Ankyrin repeat"/>
        <property type="match status" value="3"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF54791">
        <property type="entry name" value="Eukaryotic type KH-domain (KH-domain type I)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50297">
        <property type="entry name" value="ANK_REP_REGION"/>
        <property type="match status" value="16"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50088">
        <property type="entry name" value="ANK_REPEAT"/>
        <property type="match status" value="16"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50084">
        <property type="entry name" value="KH_TYPE_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="8 13" id="KW-0040">ANK repeat</keyword>
    <keyword evidence="9 15" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="7" id="KW-0391">Immunity</keyword>
    <keyword evidence="4" id="KW-0399">Innate immunity</keyword>
    <keyword evidence="3" id="KW-1017">Isopeptide bond</keyword>
    <keyword evidence="2" id="KW-0488">Methylation</keyword>
    <keyword evidence="10" id="KW-0539">Nucleus</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="5" id="KW-0677">Repeat</keyword>
    <keyword evidence="14" id="KW-0694">RNA-binding</keyword>
    <keyword evidence="6" id="KW-0832">Ubl conjugation</keyword>
    <feature type="repeat" description="ANK" evidence="13">
        <location>
            <begin position="155"/>
            <end position="187"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="13">
        <location>
            <begin position="188"/>
            <end position="220"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="13">
        <location>
            <begin position="221"/>
            <end position="253"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="13">
        <location>
            <begin position="286"/>
            <end position="310"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="13">
        <location>
            <begin position="354"/>
            <end position="386"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="13">
        <location>
            <begin position="387"/>
            <end position="419"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="13">
        <location>
            <begin position="420"/>
            <end position="452"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="13">
        <location>
            <begin position="454"/>
            <end position="486"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="13">
        <location>
            <begin position="875"/>
            <end position="907"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="13">
        <location>
            <begin position="908"/>
            <end position="940"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="13">
        <location>
            <begin position="942"/>
            <end position="974"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="13">
        <location>
            <begin position="975"/>
            <end position="1007"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="13">
        <location>
            <begin position="1036"/>
            <end position="1068"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="13">
        <location>
            <begin position="1069"/>
            <end position="1101"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="13">
        <location>
            <begin position="1104"/>
            <end position="1136"/>
        </location>
    </feature>
    <feature type="repeat" description="ANK" evidence="13">
        <location>
            <begin position="1137"/>
            <end position="1169"/>
        </location>
    </feature>
    <feature type="domain" description="K Homology" evidence="17">
        <location>
            <begin position="1463"/>
            <end position="1533"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1264"/>
            <end position="1459"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1629"/>
            <end position="1718"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="1736"/>
            <end position="1857"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="2005"/>
            <end position="2048"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="16">
        <location>
            <begin position="2082"/>
            <end position="2103"/>
        </location>
    </feature>
    <feature type="coiled-coil region" evidence="15">
        <location>
            <begin position="599"/>
            <end position="695"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic residues" evidence="16">
        <location>
            <begin position="1266"/>
            <end position="1276"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="16">
        <location>
            <begin position="1277"/>
            <end position="1306"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="16">
        <location>
            <begin position="1320"/>
            <end position="1335"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="16">
        <location>
            <begin position="1387"/>
            <end position="1396"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="16">
        <location>
            <begin position="1405"/>
            <end position="1414"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1415"/>
            <end position="1442"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="16">
        <location>
            <begin position="1673"/>
            <end position="1717"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="16">
        <location>
            <begin position="1736"/>
            <end position="1751"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1780"/>
            <end position="1801"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="16">
        <location>
            <begin position="1811"/>
            <end position="1828"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="1833"/>
            <end position="1842"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="16">
        <location>
            <begin position="1848"/>
            <end position="1857"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="2005"/>
            <end position="2021"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="16">
        <location>
            <begin position="2037"/>
            <end position="2047"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="16">
        <location>
            <begin position="2084"/>
            <end position="2098"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022481"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022499"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022588"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022843"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022859"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00023043"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00023054"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00023242"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00067263"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00081278"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00023"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00117"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="SAM" id="Coils"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="17">
        <source>
            <dbReference type="SMART" id="SM00322"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="18">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000026196.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="2284" mass="242608" checksum="725592E49179CDF0" modified="2019-05-08" version="1">MLETASKLLLSSTADGADLRTVDPETQARLEALLEAAGIGKLSTADGKAFADPEVLRRLTSSVSCALDEAAAALTRMRAESTANAGQTDNRSLAEACSEGDVNAVRKLLIEGRSVNEHTEEGESLLCLACSAGYYELAQVLLAMHANVEDRGIKGDITPLMAAANGGHVKIVKLLLAHGADVNAQSSTGNTALTYACAGGYVDVVKVLLESGASIEDHNENGHTPLMEAGSAGHVEVARVLLENGAGINTHSNEFKESALTLACYKGALAALLIERGANLEEVNDEGYTPLMEAAREGHEEMVALLLGQGNYNRANINAQTEETQETALTLACCGGFLEVADFLIKAGADIELGCSTPLMEAAQEGHLELVKYLLAAGANVHATTATGDTALTYACENGHTDVADVLLQAGADLEHESEGGRTPLMKAARAGHVCTVQFLISKGANVNRTTANNDHTVLSLACAGGHLAVVELLLAHGADPTHRLKDGSTMLIEAAKGGHTSVVCYLLDYPNNLLSAPAPDATQLTPPSHDLNRAPRVPVQALPMVVPPQEPDKPPANVATTLPIRNKAASKQKSSTHLPANNQDVQGYITNQSPESIVEEAQGKLTELEQRIKEAIEKNAQLQSLELAHADQLTKEKIEELNKTREEQIQKKQKILEELQKVERELQLKTQQQLKKQYLEVKAQRIQLQQQQQQQQSCQHLGLLTPVGVGEQLSEGDYARLQQVDPILLKDDPQQPAMQMGFAPIQPLAMPQALPLATGSLPPGSIANLTELQGVIVGQPVLGQAQLAGLGQGILTETQQGLMVASPAQTLNDTLDDIMAAVSGRASAMSNTPTHSIATSVSQPQTPAPSPIISPSAMLPIYPAIDIDAQTESNHDTALTLACAGGHEELVQTLLERGANIEHRDKKGFTPLILAATAGHVGVVEILLDNGADIEAQSERTKDTPLSLACSGGRQEVVELLLARGANKEHRNVSDYTPLSLAASGGYVNIIKILLNAGAEINSRTGSKLAMNGHTAAVKLLLDMGSDINAQIETNRNTALTLACFQGRTEVVSLLLDRKANVEHRAKTGLTPLMEAASGGYAEVGRVLLDKGADVNAPPVPSSRDTALTIAADKGHYKFCELLISRGGHIDVRNKKGNTPLWLAANGGHLDVVQLLVQAGADVDAADNRKITPLMAAFRKGHVKVVRYLVKEVNQFPSDSECMRYIATITDKEMLKKCHLCMESIVQAKDRQAAEANKNASILLEELDLEKLREESRRLALAAKREKRKEKRRKKKEEQRRKLEEIEAKNKENFELQAAQEKEELKTEDEPEVLMEPPSATTTTTIGISATWTTLAGSHGKRNNTITTTSSKRKSRKNKINPDNVQILFDDQLPISYGQPEKANGESKSSSTSESGDSDNMRISSCSDESSNSNKVISEVTSNSLSTCTKSGPSPLSSPNGKLTIASPKRGQKREEGWKEVVRRSKKVSVPSTVISRVIGRGGCNINAIREFTGAHIDIDKQKDKTGDRIITIRGGTESTRQATQLINALIKDPDKEIDELIPKNRLKTVNNSLMGIKVTTVAVQSTSQTATALTVPAISSAPTHKTVKNPVNNVRPGFPVSLPLAYPPPQFAHALLAAQTFQQIRPPRLPMTHFGGTFPPAQSTWGPFPVRPLSPARATNSPKPHMVPRHNSQNSSGSQVNSTSSLTTSLTTATSSTASAVPVSSTNGSPSSPSVRRQLFVTVVKTSNATTTTITTTASNNSTAPTNATYPIPTAKEHYPVSSPSSPSPPAQPVAVSRNSPSDCVTTSPNKGASSSDQEAASPPVVETSNLASSGQLSASSLGSSLVHPAHQQSPGSLSQEARLPPQQPQVPLPDPRMVVPPNLAAVTSSSAPVVGSTNAPMTYPLPQTSMGSTQPPAKMETPAIRPPAHGTNTGHKTPAPVQNCSVAILNVNHIKRPHSVPSSVQLPSTLNTQTASQNSAHPANKSVGTNFSATLPFGPFSTLFENSSTSAHAFWGGSVVSSQTTPESIRSGKSSFLPNSEPLHRSDTSKAPGFRPPLQRPAPSPSGKCVCVCVVSSAECLHFGTLNIFLSSGVRAPSPTPSAVSLGSEKPSNASQDRKVPVPIGTERSARIRQTGTSTPSVIGSNLATPVGHSGIWSFEGIGANQGMKYRLTDWCHSGMGTHMIHRPMSDPGVFSHQVMERDSTGIVTPSGTFHQHVPAGYMDFPKGMPFSVYGNAMIPPVAPIADGTGGPIFNGPHAADPSWNSLIKMVSNSTENNGPQTVWTGTWTPHMNSVHMNQLG</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="24" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IE76</accession>
    <name>A0A452IE76_9SAUR</name>
    <protein>
        <submittedName>
            <fullName evidence="1">Uncharacterized protein</fullName>
        </submittedName>
    </protein>
    <organism evidence="1 2">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="2" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="1" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="1" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <dbReference type="Ensembl" id="ENSGAGT00000029333.1">
        <property type="protein sequence ID" value="ENSGAGP00000025788.1"/>
        <property type="gene ID" value="ENSGAGG00000018821.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="2" id="KW-1185">Reference proteome</keyword>
    <evidence type="ECO:0000313" key="1">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000025788.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="2">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="155" mass="17475" checksum="E6C841EA70A6A91D" modified="2019-05-08" version="1">MQAPAVREHTPVSEAHHHATEISLWTVVAAIXPVSEADHHTTEISLWTVVAAIQAVERKVNSHATRLLNLEGRTGTTEKKLTDCERTAMEFGNQLESKWTILGTLIQENNFLQRRLENMENLLRNRNFWILRLPPGTKGEVPKVTISQLVLGVQT</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="27" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IE78</accession>
    <name>A0A452IE78_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="6">Synapsin ATP-binding domain-containing protein</fullName>
        </recommendedName>
    </protein>
    <organism evidence="7 8">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="8" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="7" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="7" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="4">Synapse</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="1">Belongs to the synapsin family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IE78"/>
    <dbReference type="Ensembl" id="ENSGAGT00000029798.1">
        <property type="protein sequence ID" value="ENSGAGP00000026202.1"/>
        <property type="gene ID" value="ENSGAGG00000019121.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030672">
        <property type="term" value="C:synaptic vesicle membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0007269">
        <property type="term" value="P:neurotransmitter secretion"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="FunFam" id="3.30.470.20:FF:000042">
        <property type="entry name" value="Synapsin III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.1490.20">
        <property type="entry name" value="ATP-grasp fold, A domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.30.470.20">
        <property type="entry name" value="ATP-grasp fold, B domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR013815">
        <property type="entry name" value="ATP_grasp_subdomain_1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001359">
        <property type="entry name" value="Synapsin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR020898">
        <property type="entry name" value="Synapsin_ATP-bd_dom"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10841">
        <property type="entry name" value="SYNAPSIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10841:SF6">
        <property type="entry name" value="SYNAPSIN III"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02750">
        <property type="entry name" value="Synapsin_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR01368">
        <property type="entry name" value="SYNAPSIN"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF56059">
        <property type="entry name" value="Glutathione synthetase ATP-binding domain-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="2" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="8" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0770">Synapse</keyword>
    <feature type="domain" description="Synapsin ATP-binding" evidence="6">
        <location>
            <begin position="1"/>
            <end position="140"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="5">
        <location>
            <begin position="145"/>
            <end position="315"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="5">
        <location>
            <begin position="156"/>
            <end position="180"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Low complexity" evidence="5">
        <location>
            <begin position="189"/>
            <end position="234"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="235"/>
            <end position="246"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Pro residues" evidence="5">
        <location>
            <begin position="252"/>
            <end position="261"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Polar residues" evidence="5">
        <location>
            <begin position="276"/>
            <end position="306"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00008243"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00023018"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00034103"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="6">
        <source>
            <dbReference type="Pfam" id="PF02750"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="7">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000026202.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="8">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="330" mass="35961" checksum="00ACC6155EA07B78" modified="2019-05-08" version="1">MGKIKVENQYDFQDMASMVAMAKTYATTEAFIDSKYDIRIQKIGSNYKAYMRTSISGNWKANTGSAMLEQVAMTERYRLWADSCAEMFGGLDICAVKAVHSKDGKDYIIEVMDSSMPLIGEHVEEDTQLVADLVVSKMTQLLVTAGSIPSPPRPWPQQIQPQSMKSQMQPQPGPQLGQPSQPRPPPQGGPRQPQGSQPPRTGAPPQQRLSPQGQQPLSPQSSSPQQQRSPGSPQLTRSSTGTSPVQASKPGIFPPQQPRPPAKGRAPAQQGLGETPKQQASPHPHLNKSQSLTNTFSISESSQRGNANEDEAKAETIRNLRKSFASLFSD</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="31" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IUT2</accession>
    <name>A0A452IUT2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="16">ATP-dependent 6-phosphofructokinase</fullName>
            <shortName evidence="16">ATP-PFK</shortName>
            <shortName evidence="16">Phosphofructokinase</shortName>
            <ecNumber evidence="16">2.7.1.11</ecNumber>
        </recommendedName>
        <alternativeName>
            <fullName evidence="16">Phosphohexokinase</fullName>
        </alternativeName>
    </protein>
    <organism evidence="19 20">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="20" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="19" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="19" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="16">Catalyzes the phosphorylation of D-fructose 6-phosphate to fructose 1,6-bisphosphate by ATP, the first committing step of glycolysis.</text>
    </comment>
    <comment type="catalytic activity">
        <reaction evidence="15 16 17">
            <text>beta-D-fructose 6-phosphate + ATP = beta-D-fructose 1,6-bisphosphate + ADP + H(+)</text>
            <dbReference type="Rhea" id="RHEA:16109"/>
            <dbReference type="ChEBI" id="CHEBI:15378"/>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
            <dbReference type="ChEBI" id="CHEBI:32966"/>
            <dbReference type="ChEBI" id="CHEBI:57634"/>
            <dbReference type="ChEBI" id="CHEBI:456216"/>
            <dbReference type="EC" id="2.7.1.11"/>
        </reaction>
    </comment>
    <comment type="cofactor">
        <cofactor evidence="1 16">
            <name>Mg(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:18420"/>
        </cofactor>
    </comment>
    <comment type="activity regulation">
        <text evidence="16">Allosterically activated by ADP, AMP, or fructose 2,6-bisphosphate, and allosterically inhibited by ATP or citrate.</text>
    </comment>
    <comment type="pathway">
        <text evidence="2 16 17">Carbohydrate degradation; glycolysis; D-glyceraldehyde 3-phosphate and glycerone phosphate from D-glucose: step 3/4.</text>
    </comment>
    <comment type="subunit">
        <text evidence="16">Homo- and heterotetramers.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="16">Cytoplasm</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="17">Belongs to the phosphofructokinase type A (PFKA) family. ATP-dependent PFK group I subfamily. Eukaryotic two domain clade "E" sub-subfamily.</text>
    </comment>
    <comment type="similarity">
        <text evidence="16">Belongs to the phosphofructokinase type A (PFKA) family. ATP-dependent PFK group I subfamily. Eukaryotic two domain clade 'E' sub-subfamily.</text>
    </comment>
    <comment type="caution">
        <text evidence="16">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="EC" id="2.7.1.11" evidence="16"/>
    <dbReference type="AlphaFoldDB" id="A0A452IUT2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000036064.1">
        <property type="protein sequence ID" value="ENSGAGP00000031799.1"/>
        <property type="gene ID" value="ENSGAGG00000022790.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00109">
        <property type="reaction ID" value="UER00182"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005945">
        <property type="term" value="C:6-phosphofructokinase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016020">
        <property type="term" value="C:membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0003872">
        <property type="term" value="F:6-phosphofructokinase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016208">
        <property type="term" value="F:AMP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0070095">
        <property type="term" value="F:fructose-6-phosphate binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0042802">
        <property type="term" value="F:identical protein binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0046872">
        <property type="term" value="F:metal ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0048029">
        <property type="term" value="F:monosaccharide binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0061621">
        <property type="term" value="P:canonical glycolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030388">
        <property type="term" value="P:fructose 1,6-bisphosphate metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006002">
        <property type="term" value="P:fructose 6-phosphate metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="CDD" id="cd00764">
        <property type="entry name" value="Eukaryotic_PFK"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.460:FF:000001">
        <property type="entry name" value="ATP-dependent 6-phosphofructokinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.460:FF:000003">
        <property type="entry name" value="ATP-dependent 6-phosphofructokinase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.50.450:FF:000043">
        <property type="entry name" value="ATP-dependent 6-phosphofructokinase, platelet type"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.450">
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.50.460">
        <property type="entry name" value="Phosphofructokinase domain"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="HAMAP" id="MF_03184">
        <property type="entry name" value="Phosphofructokinase_I_E"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR009161">
        <property type="entry name" value="6-Pfructokinase_euk"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR022953">
        <property type="entry name" value="ATP_PFK"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR041914">
        <property type="entry name" value="PFK_vert-type"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015912">
        <property type="entry name" value="Phosphofructokinase_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000023">
        <property type="entry name" value="Phosphofructokinase_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035966">
        <property type="entry name" value="PKF_sf"/>
    </dbReference>
    <dbReference type="NCBIfam" id="TIGR02478">
        <property type="entry name" value="6PF1K_euk"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="NCBIfam" id="NF002872">
        <property type="entry name" value="PRK03202.1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13697:SF5">
        <property type="entry name" value="ATP-DEPENDENT 6-PHOSPHOFRUCTOKINASE, PLATELET TYPE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR13697">
        <property type="entry name" value="PHOSPHOFRUCTOKINASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00365">
        <property type="entry name" value="PFK"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF000533">
        <property type="entry name" value="ATP_PFK_euk"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00476">
        <property type="entry name" value="PHFRCTKINASE"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF53784">
        <property type="entry name" value="Phosphofructokinase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00433">
        <property type="entry name" value="PHOSPHOFRUCTOKINASE"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12" id="KW-0007">Acetylation</keyword>
    <keyword evidence="4 16" id="KW-0021">Allosteric enzyme</keyword>
    <keyword evidence="10 16 17" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="3 16" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="13 16 17" id="KW-0324">Glycolysis</keyword>
    <keyword evidence="14" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="9 16 17" id="KW-0418">Kinase</keyword>
    <keyword evidence="11 16" id="KW-0460">Magnesium</keyword>
    <keyword evidence="7 16" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="8 16 17" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="5" id="KW-0597">Phosphoprotein</keyword>
    <keyword evidence="20" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="6 16 17" id="KW-0808">Transferase</keyword>
    <feature type="domain" description="Phosphofructokinase" evidence="18">
        <location>
            <begin position="28"/>
            <end position="327"/>
        </location>
    </feature>
    <feature type="domain" description="Phosphofructokinase" evidence="18">
        <location>
            <begin position="403"/>
            <end position="688"/>
        </location>
    </feature>
    <feature type="region of interest" description="N-terminal catalytic PFK domain 1" evidence="16">
        <location>
            <begin position="1"/>
            <end position="390"/>
        </location>
    </feature>
    <feature type="region of interest" description="C-terminal regulatory PFK domain 2" evidence="16">
        <location>
            <begin position="403"/>
            <end position="784"/>
        </location>
    </feature>
    <feature type="active site" description="Proton acceptor" evidence="16">
        <location>
            <position position="170"/>
        </location>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="35"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <begin position="98"/>
            <end position="99"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
        </ligand>
    </feature>
    <feature type="binding site" description="in other chain" evidence="16">
        <location>
            <begin position="168"/>
            <end position="170"/>
        </location>
        <ligand>
            <name>substrate</name>
            <note>ligand shared between dimeric partners</note>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="205"/>
        </location>
        <ligand>
            <name>substrate</name>
            <note>ligand shared between dimeric partners</note>
        </ligand>
    </feature>
    <feature type="binding site" description="in other chain" evidence="16">
        <location>
            <begin position="212"/>
            <end position="214"/>
        </location>
        <ligand>
            <name>substrate</name>
            <note>ligand shared between dimeric partners</note>
        </ligand>
    </feature>
    <feature type="binding site" description="in other chain" evidence="16">
        <location>
            <position position="268"/>
        </location>
        <ligand>
            <name>substrate</name>
            <note>ligand shared between dimeric partners</note>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="296"/>
        </location>
        <ligand>
            <name>substrate</name>
            <note>ligand shared between dimeric partners</note>
        </ligand>
    </feature>
    <feature type="binding site" description="in other chain" evidence="16">
        <location>
            <begin position="302"/>
            <end position="305"/>
        </location>
        <ligand>
            <name>substrate</name>
            <note>ligand shared between dimeric partners</note>
        </ligand>
    </feature>
    <feature type="binding site" description="in other chain" evidence="16">
        <location>
            <position position="472"/>
        </location>
        <ligand>
            <name>beta-D-fructose 2,6-bisphosphate</name>
            <dbReference type="ChEBI" id="CHEBI:58579"/>
            <note>allosteric activator; ligand shared between dimeric partners</note>
        </ligand>
    </feature>
    <feature type="binding site" description="in other chain" evidence="16">
        <location>
            <begin position="529"/>
            <end position="533"/>
        </location>
        <ligand>
            <name>beta-D-fructose 2,6-bisphosphate</name>
            <dbReference type="ChEBI" id="CHEBI:58579"/>
            <note>allosteric activator; ligand shared between dimeric partners</note>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="567"/>
        </location>
        <ligand>
            <name>beta-D-fructose 2,6-bisphosphate</name>
            <dbReference type="ChEBI" id="CHEBI:58579"/>
            <note>allosteric activator; ligand shared between dimeric partners</note>
        </ligand>
    </feature>
    <feature type="binding site" description="in other chain" evidence="16">
        <location>
            <begin position="574"/>
            <end position="576"/>
        </location>
        <ligand>
            <name>beta-D-fructose 2,6-bisphosphate</name>
            <dbReference type="ChEBI" id="CHEBI:58579"/>
            <note>allosteric activator; ligand shared between dimeric partners</note>
        </ligand>
    </feature>
    <feature type="binding site" description="in other chain" evidence="16">
        <location>
            <position position="630"/>
        </location>
        <ligand>
            <name>beta-D-fructose 2,6-bisphosphate</name>
            <dbReference type="ChEBI" id="CHEBI:58579"/>
            <note>allosteric activator; ligand shared between dimeric partners</note>
        </ligand>
    </feature>
    <feature type="binding site" evidence="16">
        <location>
            <position position="656"/>
        </location>
        <ligand>
            <name>beta-D-fructose 2,6-bisphosphate</name>
            <dbReference type="ChEBI" id="CHEBI:58579"/>
            <note>allosteric activator; ligand shared between dimeric partners</note>
        </ligand>
    </feature>
    <feature type="binding site" description="in other chain" evidence="16">
        <location>
            <begin position="662"/>
            <end position="665"/>
        </location>
        <ligand>
            <name>beta-D-fructose 2,6-bisphosphate</name>
            <dbReference type="ChEBI" id="CHEBI:58579"/>
            <note>allosteric activator; ligand shared between dimeric partners</note>
        </ligand>
    </feature>
    <feature type="binding site" description="in other chain" evidence="16">
        <location>
            <position position="736"/>
        </location>
        <ligand>
            <name>beta-D-fructose 2,6-bisphosphate</name>
            <dbReference type="ChEBI" id="CHEBI:58579"/>
            <note>allosteric activator; ligand shared between dimeric partners</note>
        </ligand>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00001946"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00004679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022490"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022533"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022553"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022679"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022741"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022777"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022840"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022842"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022990"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023152"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00048070"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="HAMAP-Rule" id="MF_03184"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="PIRNR" id="PIRNR000533"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="18">
        <source>
            <dbReference type="Pfam" id="PF00365"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="19">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000031799.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="20">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="784" mass="86185" checksum="E75C75CB89D9EB76" modified="2019-05-08" version="1">MQGDDREQLPQSSFLQHLQNLSGAGKAIGVLTSGGDAQGMNAAVRAVVRMGIYVEAKVYFIYEGYQGMVDGGDNIVEVSWESVSSILQVGGTVIGSARCKEFRTREGRLKAAFNLIQRGITNLCVIGGANLFREEWSGLLEELAQSGKIEEEAVKKYVYLNIVGMVGSIDNDFCGTDMTIGTDSALHRIIEVVDAIMTTAQSHQRTFVLEVMGRHCGYLALVSALACGADWVFIPEYPPEEGWEDQMCVKLSENRARKKRLNIIIVAEGAIDSHNKPITSEKVKDLVVQQLGFDTRVTILGHVQRGGTPSAFDRILASRMGVEAVLALLEATPDTPACVVSLSGNQAVRLPLMECVQMVSSGRRASLHLFINYNKVSFENNLNTYKLLSHRKPDTELPKTNFNVAVLNVGAPAAGMNAAVRSAVRVGITEGHNMFAVIDGFEGFANGQIKEISWGDVGGWTGQGGSLLGTKRTLPAKYLEKIADQMRTNNINALMVIGGFEAYLGLLELSAAREKYDEFCVPMVMVPATVSNNVPGSDFSIGADTALNTITDTCDRIKQSASGTKRRVFIIETMGGYCGYLANMGGLAAGADAAYIFEEKFDIRELQANVEHLTEKMKTSIQRGLVLRNENSNENYTTDFIYQLYSEEGKGVFDCRKNVLGHMQQGGAPSPFDRNFGTKVSAKAVQWISKKLRETYRRGKVFANTEDSVCLLGMRRRNLIFQPVVELKDQADFVHRIPKEQWWLKLRPLMKILAKYKTSYDVSDSGQLEHVARHSPKEAETGAI</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IUV1</accession>
    <name>A0A452IUV1_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Phosphorylase b kinase regulatory subunit</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="7">Phosphorylase b kinase catalyzes the phosphorylation of serine in certain substrates, including troponin I.</text>
    </comment>
    <comment type="pathway">
        <text evidence="2 7">Glycan biosynthesis; glycogen metabolism.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 7">Cell membrane</location>
            <topology evidence="1 7">Lipid-anchor</topology>
            <orientation evidence="1 7">Cytoplasmic side</orientation>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 7">Belongs to the phosphorylase b kinase regulatory chain family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IUV1"/>
    <dbReference type="Ensembl" id="ENSGAGT00000036180.1">
        <property type="protein sequence ID" value="ENSGAGP00000031902.1"/>
        <property type="gene ID" value="ENSGAGG00000022485.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00163"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005964">
        <property type="term" value="C:phosphorylase kinase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005977">
        <property type="term" value="P:glycogen metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008928">
        <property type="entry name" value="6-hairpin_glycosidase_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011613">
        <property type="entry name" value="GH15-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045583">
        <property type="entry name" value="KPBA/B_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008734">
        <property type="entry name" value="PHK_A/B_su"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10749">
        <property type="entry name" value="PHOSPHORYLASE B KINASE REGULATORY SUBUNIT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10749:SF8">
        <property type="entry name" value="PHOSPHORYLASE B KINASE REGULATORY SUBUNIT BETA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00723">
        <property type="entry name" value="Glyco_hydro_15"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19292">
        <property type="entry name" value="KPBB_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48208">
        <property type="entry name" value="Six-hairpin glycosidases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 7" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="6 7" id="KW-0119">Carbohydrate metabolism</keyword>
    <keyword evidence="7" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="4 7" id="KW-0321">Glycogen metabolism</keyword>
    <keyword evidence="7" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="7" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0636">Prenylation</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="GH15-like" evidence="8">
        <location>
            <begin position="24"/>
            <end position="746"/>
        </location>
    </feature>
    <feature type="domain" description="Phosphorylase b kinase regulatory subunit alpha/beta C-terminal" evidence="9">
        <location>
            <begin position="812"/>
            <end position="970"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004342"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005131"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022600"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023277"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU364123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF00723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF19292"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000031902.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="986" mass="113109" checksum="D2625C4CEA8A7A47" modified="2019-05-08" version="1">VASWQDQMIINLPRPEGESLWDKLDHYYRIVKSTVLLYQSPTTGLFPTKTFGDNQKAKVQDSLYCAACAWALALAYRRIDDDKGRTHELEHSAIKCMRGILYCCMRQADKVQQFKQDPRPSKCLHSVFSAHSGDEVFSHDEYGHLQINAVSLFLLYLVEMISSGLQIIYNTDEVSFIQNLVFCVERAYRVPDFGIWERGSKYNNGSTELHSSSVGLAKAALEAINGFNLFGNQGCSWSIIFVDLDAHNRNGQTLCSLLPRESRSHNTDAALLPCISYPAFAVDDDALYSQTLDKIVRKLKGKYGFKRFLRDGYRTSLEDRNRRYYKPAEIKVMEYVNGTHNIGNPQQVKEYQELLDHLLQHSSEGYPVVPKYYYVPADFVELEKKNPGSQKRFPSNSGRDGKFFLWGQALYIIAKLLVDGLVNPKDIDPVGRYIPPQDQRNVSMRYSNQGPLENDLVVHVALIAESQRLQVFLNTYGIQTQTPQQVEPIQIWAQQELVKAYFHLGINDKLGLSGRPDRPIGCLGTSKIYRILGKTVVCYPIIFDLSDFYMSQDVMLLIDDIKNALQFIKQYWKMHGRPLFLVLIREDNIRGSRFNPILDMLAAFKKGVVGGVKVHADRVQTLISGAVVEQLDFLRISETEEPPDFKSFEELELPKHSKVKRQTSTPNVSELEQQPEVNINDWKIRSTYEILQKLHDCNCLASQAMLLSILLKREGPNFITKEGTISDHIERIYRRAGSKKLWFVYNIFKFASSIAPHITTLLVHGKQVKCTEVLNSAYELKIRAGDMPPRDLYQLSPSEVKQLLLDILQPQQQGRCWLNRRQIDGSLNRTPHGFYDRVWQILERTPNGVIVAEKYLPQQPTLSDMTMYEMNFSLLVEDMLQNIDQPEYRQIIVELLMVVSVILERNPELEFQDKVDLDKLVKEAFHYFQKDEIRLKGVEKQDDMTSFYNTPPMGKRGTCSYLTKAVMNLLLEGEVKPCSDDPCTIS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="32" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IUV2</accession>
    <name>A0A452IUV2_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="21">Metalloendopeptidase</fullName>
            <ecNumber evidence="21">3.4.24.-</ecNumber>
        </recommendedName>
    </protein>
    <organism evidence="26 27">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="27" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="26" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="26" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="cofactor">
        <cofactor evidence="20 21">
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
        </cofactor>
        <text evidence="20 21">Binds 1 zinc ion per subunit.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Secreted</location>
        </subcellularLocation>
    </comment>
    <comment type="caution">
        <text evidence="19">Lacks conserved residue(s) required for the propagation of feature annotation.</text>
    </comment>
    <dbReference type="EC" id="3.4.24.-" evidence="21"/>
    <dbReference type="AlphaFoldDB" id="A0A452IUV2"/>
    <dbReference type="Ensembl" id="ENSGAGT00000036049.1">
        <property type="protein sequence ID" value="ENSGAGP00000031787.1"/>
        <property type="gene ID" value="ENSGAGG00000022834.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005576">
        <property type="term" value="C:extracellular region"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005509">
        <property type="term" value="F:calcium ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0004222">
        <property type="term" value="F:metalloendopeptidase activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0008270">
        <property type="term" value="F:zinc ion binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030154">
        <property type="term" value="P:cell differentiation"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006508">
        <property type="term" value="P:proteolysis"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="CDD" id="cd00041">
        <property type="entry name" value="CUB"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="CDD" id="cd00054">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="CDD" id="cd04281">
        <property type="entry name" value="ZnMc_BMP1_TLD"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.290:FF:000013">
        <property type="entry name" value="Membrane frizzled-related protein"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.10.25.10:FF:000022">
        <property type="entry name" value="Metalloendopeptidase"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.290:FF:000004">
        <property type="entry name" value="Metalloendopeptidase"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Metalloendopeptidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.60.120.290:FF:000011">
        <property type="entry name" value="Metalloendopeptidase"/>
        <property type="match status" value="1"/>
    </dbReference>
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        <property type="entry name" value="Metalloendopeptidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.390.10:FF:000004">
        <property type="entry name" value="Metalloendopeptidase"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.390.10">
        <property type="entry name" value="Collagenase (Catalytic Domain)"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.10.25.10">
        <property type="entry name" value="Laminin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.60.120.290">
        <property type="entry name" value="Spermadhesin, CUB domain"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR015446">
        <property type="entry name" value="BMP_1/tolloid-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000859">
        <property type="entry name" value="CUB_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000742">
        <property type="entry name" value="EGF"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001881">
        <property type="entry name" value="EGF-like_Ca-bd_dom"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR000152">
        <property type="entry name" value="EGF-type_Asp/Asn_hydroxyl_site"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR018097">
        <property type="entry name" value="EGF_Ca-bd_CS"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR024079">
        <property type="entry name" value="MetalloPept_cat_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR049883">
        <property type="entry name" value="NOTCH1_EGF-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001506">
        <property type="entry name" value="Peptidase_M12A"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR006026">
        <property type="entry name" value="Peptidase_Metallo"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR035914">
        <property type="entry name" value="Sperma_CUB_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR034036">
        <property type="entry name" value="ZnMP_TLD/BMP1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24251:SF45">
        <property type="entry name" value="METALLOENDOPEPTIDASE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR24251">
        <property type="entry name" value="OVOCHYMASE-RELATED"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01400">
        <property type="entry name" value="Astacin"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00431">
        <property type="entry name" value="CUB"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Pfam" id="PF07645">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF14670">
        <property type="entry name" value="FXa_inhibition"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PIRSF" id="PIRSF001199">
        <property type="entry name" value="BMP_1/tolloid-like"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00480">
        <property type="entry name" value="ASTACIN"/>
    </dbReference>
    <dbReference type="SMART" id="SM00042">
        <property type="entry name" value="CUB"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="SMART" id="SM00181">
        <property type="entry name" value="EGF"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00179">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SMART" id="SM00235">
        <property type="entry name" value="ZnMc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF57196">
        <property type="entry name" value="EGF/Laminin"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF55486">
        <property type="entry name" value="Metalloproteases ('zincins'), catalytic domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF49854">
        <property type="entry name" value="Spermadhesin, CUB domain"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51864">
        <property type="entry name" value="ASTACIN"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS00010">
        <property type="entry name" value="ASX_HYDROXYL"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01180">
        <property type="entry name" value="CUB"/>
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01186">
        <property type="entry name" value="EGF_2"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50026">
        <property type="entry name" value="EGF_3"/>
        <property type="match status" value="2"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS01187">
        <property type="entry name" value="EGF_CA"/>
        <property type="match status" value="2"/>
    </dbReference>
    <proteinExistence type="predicted"/>
    <keyword evidence="12" id="KW-0106">Calcium</keyword>
    <keyword evidence="2" id="KW-0217">Developmental protein</keyword>
    <keyword evidence="9" id="KW-0221">Differentiation</keyword>
    <keyword evidence="15 20" id="KW-1015">Disulfide bond</keyword>
    <keyword evidence="4 19" id="KW-0245">EGF-like domain</keyword>
    <keyword evidence="16" id="KW-0325">Glycoprotein</keyword>
    <keyword evidence="10 20 21" id="KW-0378">Hydrolase</keyword>
    <keyword evidence="6 18 20" id="KW-0479">Metal-binding</keyword>
    <keyword evidence="13 20 21" id="KW-0482">Metalloprotease</keyword>
    <keyword evidence="5 20 21" id="KW-0645">Protease</keyword>
    <keyword evidence="27" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="8" id="KW-0677">Repeat</keyword>
    <keyword evidence="3" id="KW-0964">Secreted</keyword>
    <keyword evidence="7 21" id="KW-0732">Signal</keyword>
    <keyword evidence="11 18 20" id="KW-0862">Zinc</keyword>
    <keyword evidence="14" id="KW-0865">Zymogen</keyword>
    <feature type="signal peptide" evidence="21">
        <location>
            <begin position="1"/>
            <end position="22"/>
        </location>
    </feature>
    <feature type="chain" id="PRO_5018817665" description="Metalloendopeptidase" evidence="21">
        <location>
            <begin position="23"/>
            <end position="980"/>
        </location>
    </feature>
    <feature type="domain" description="Peptidase M12A" evidence="25">
        <location>
            <begin position="114"/>
            <end position="314"/>
        </location>
    </feature>
    <feature type="domain" description="CUB" evidence="23">
        <location>
            <begin position="316"/>
            <end position="428"/>
        </location>
    </feature>
    <feature type="domain" description="CUB" evidence="23">
        <location>
            <begin position="429"/>
            <end position="541"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="24">
        <location>
            <begin position="541"/>
            <end position="582"/>
        </location>
    </feature>
    <feature type="domain" description="CUB" evidence="23">
        <location>
            <begin position="585"/>
            <end position="697"/>
        </location>
    </feature>
    <feature type="domain" description="EGF-like" evidence="24">
        <location>
            <begin position="697"/>
            <end position="737"/>
        </location>
    </feature>
    <feature type="domain" description="CUB" evidence="23">
        <location>
            <begin position="741"/>
            <end position="853"/>
        </location>
    </feature>
    <feature type="domain" description="CUB" evidence="23">
        <location>
            <begin position="854"/>
            <end position="970"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="22">
        <location>
            <begin position="77"/>
            <end position="99"/>
        </location>
    </feature>
    <feature type="active site" evidence="17 20">
        <location>
            <position position="208"/>
        </location>
    </feature>
    <feature type="binding site" evidence="18 20">
        <location>
            <position position="207"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <note>catalytic</note>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18 20">
        <location>
            <position position="211"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <note>catalytic</note>
        </ligand>
    </feature>
    <feature type="binding site" evidence="18 20">
        <location>
            <position position="217"/>
        </location>
        <ligand>
            <name>Zn(2+)</name>
            <dbReference type="ChEBI" id="CHEBI:29105"/>
            <note>catalytic</note>
        </ligand>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="177"/>
            <end position="199"/>
        </location>
    </feature>
    <feature type="disulfide bond" evidence="20">
        <location>
            <begin position="179"/>
            <end position="180"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004613"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00022473"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00022525"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022536"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022670"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00022723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="ARBA" id="ARBA00022729"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="8">
        <source>
            <dbReference type="ARBA" id="ARBA00022737"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="9">
        <source>
            <dbReference type="ARBA" id="ARBA00022782"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="10">
        <source>
            <dbReference type="ARBA" id="ARBA00022801"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="11">
        <source>
            <dbReference type="ARBA" id="ARBA00022833"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="12">
        <source>
            <dbReference type="ARBA" id="ARBA00022837"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="13">
        <source>
            <dbReference type="ARBA" id="ARBA00023049"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="14">
        <source>
            <dbReference type="ARBA" id="ARBA00023145"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="15">
        <source>
            <dbReference type="ARBA" id="ARBA00023157"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="16">
        <source>
            <dbReference type="ARBA" id="ARBA00023180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="17">
        <source>
            <dbReference type="PIRSR" id="PIRSR001199-1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="18">
        <source>
            <dbReference type="PIRSR" id="PIRSR001199-2"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="19">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU00076"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="20">
        <source>
            <dbReference type="PROSITE-ProRule" id="PRU01211"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="21">
        <source>
            <dbReference type="RuleBase" id="RU361183"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="22">
        <source>
            <dbReference type="SAM" id="MobiDB-lite"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="23">
        <source>
            <dbReference type="PROSITE" id="PS01180"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="24">
        <source>
            <dbReference type="PROSITE" id="PS50026"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="25">
        <source>
            <dbReference type="PROSITE" id="PS51864"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="26">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000031787.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="27">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="980" mass="110793" checksum="95EB7D7FED817B88" modified="2019-05-08" version="1">MGWAAPLGTLLLLLLRVPAARGSWGLAEEESLEQQPGYRDPCKAAAFWGDIALDEEDLKLFQIDRTIDLTKHLDENTRHTTDGRRNATLSRRPTEEEDKDLLLSATGSSPRIRRATTSRAERIWPGGVIPYVIGGNFTGSQRAIFKQAMRHWEKHTCVTFVERTDEESFIVFTYRTCGCCSYVGRRGGGPQAISIGKNCDKFGIVAHELGHVVGFWHEHTRPDRDQHVTIIRENIQQGQEYNFLKMEAGEVNSLGETYDFDSIMHYARNTFSRGVFLDTILPRRDDNGVRPTIGQRIRLSQGDIAQARKLYKCPACGETLQDTTGNFSAPGFPNGYPSYSHCVWRISVTPGEKIILNFTSMDLFKSRLCWYDYVEVRDGYWRKAPLLGRFCGDKIPEPLISTDSRLWIEFRSSSNILGKGFFAVYEAICGGEIHKDVGQIQSPNYPDDYRPSKECVWRITVSEGFHVGLTFQAFEIERHDSCAYDYLEIRDGLTEDSPLMGHFCGYEKPEDIKSSSNKVWMKFASDGSINKAGFAANFFKEVDECSRPDNGGCEQRCMNTLGSYKCACDPGYELTADKKSCEAACGGFITKLNGTITSPGWPKEYPTNKNCVWQVVAPAQYRISLQFEVFELEGNDVCKYDYVEVRSGLASDSKLHGKFCGSEKPEVITSYSNNLRLEFKSDNAVSKKGFKVHFFSDKDECSKDNSGCQHECVNTFGSYVCQCRNGFMLHENGHDCKEAGCEHKVNSAEGTMASPNWPDKYPSRKECTWNISTTPGHRVKITFNEFEIEQHQECAYDHLEMYDGPNSKSPILGRFCGSKKPDPVVASTNKMFLRFYSDASVQRKGFQAKHSTECGGRLKADVRTKDLYSHAQFGDNNYPGQSNCDWVIVAEDGYGVELIFQTFELEEEADCGYDYMEIYDGYDSTAPRLGRFCGSGPLEEIYSAGDSIMIRFHTDDTINKRGFHARYTSTKFQDVLHMRK</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="26" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IUV6</accession>
    <name>A0A452IUV6_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="7">Phosphorylase b kinase regulatory subunit</fullName>
        </recommendedName>
    </protein>
    <organism evidence="10 11">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="11" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="10" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="10" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="function">
        <text evidence="7">Phosphorylase b kinase catalyzes the phosphorylation of serine in certain substrates, including troponin I.</text>
    </comment>
    <comment type="pathway">
        <text evidence="2 7">Glycan biosynthesis; glycogen metabolism.</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1 7">Cell membrane</location>
            <topology evidence="1 7">Lipid-anchor</topology>
            <orientation evidence="1 7">Cytoplasmic side</orientation>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="3 7">Belongs to the phosphorylase b kinase regulatory chain family.</text>
    </comment>
    <dbReference type="AlphaFoldDB" id="A0A452IUV6"/>
    <dbReference type="Ensembl" id="ENSGAGT00000036185.1">
        <property type="protein sequence ID" value="ENSGAGP00000031907.1"/>
        <property type="gene ID" value="ENSGAGG00000022485.1"/>
    </dbReference>
    <dbReference type="UniPathway" id="UPA00163"/>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005964">
        <property type="term" value="C:phosphorylase kinase complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005886">
        <property type="term" value="C:plasma membrane"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005516">
        <property type="term" value="F:calmodulin binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005977">
        <property type="term" value="P:glycogen metabolic process"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniPathway"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008928">
        <property type="entry name" value="6-hairpin_glycosidase_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR011613">
        <property type="entry name" value="GH15-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR045583">
        <property type="entry name" value="KPBA/B_C"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008734">
        <property type="entry name" value="PHK_A/B_su"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10749">
        <property type="entry name" value="PHOSPHORYLASE B KINASE REGULATORY SUBUNIT"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR10749:SF8">
        <property type="entry name" value="PHOSPHORYLASE B KINASE REGULATORY SUBUNIT BETA"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00723">
        <property type="entry name" value="Glyco_hydro_15"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF19292">
        <property type="entry name" value="KPBB_C"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF48208">
        <property type="entry name" value="Six-hairpin glycosidases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="5 7" id="KW-0112">Calmodulin-binding</keyword>
    <keyword evidence="6 7" id="KW-0119">Carbohydrate metabolism</keyword>
    <keyword evidence="7" id="KW-1003">Cell membrane</keyword>
    <keyword evidence="4 7" id="KW-0321">Glycogen metabolism</keyword>
    <keyword evidence="7" id="KW-0449">Lipoprotein</keyword>
    <keyword evidence="7" id="KW-0472">Membrane</keyword>
    <keyword evidence="7" id="KW-0636">Prenylation</keyword>
    <keyword evidence="11" id="KW-1185">Reference proteome</keyword>
    <feature type="domain" description="GH15-like" evidence="8">
        <location>
            <begin position="52"/>
            <end position="808"/>
        </location>
    </feature>
    <feature type="domain" description="Phosphorylase b kinase regulatory subunit alpha/beta C-terminal" evidence="9">
        <location>
            <begin position="858"/>
            <end position="1016"/>
        </location>
    </feature>
    <evidence type="ECO:0000256" key="1">
        <source>
            <dbReference type="ARBA" id="ARBA00004342"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="2">
        <source>
            <dbReference type="ARBA" id="ARBA00005131"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="3">
        <source>
            <dbReference type="ARBA" id="ARBA00007128"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="4">
        <source>
            <dbReference type="ARBA" id="ARBA00022600"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="5">
        <source>
            <dbReference type="ARBA" id="ARBA00022860"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="6">
        <source>
            <dbReference type="ARBA" id="ARBA00023277"/>
        </source>
    </evidence>
    <evidence type="ECO:0000256" key="7">
        <source>
            <dbReference type="RuleBase" id="RU364123"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="8">
        <source>
            <dbReference type="Pfam" id="PF00723"/>
        </source>
    </evidence>
    <evidence type="ECO:0000259" key="9">
        <source>
            <dbReference type="Pfam" id="PF19292"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="10">
        <source>
            <dbReference type="Ensembl" id="ENSGAGP00000031907.1"/>
        </source>
    </evidence>
    <evidence type="ECO:0000313" key="11">
        <source>
            <dbReference type="Proteomes" id="UP000291020"/>
        </source>
    </evidence>
    <sequence length="1032" mass="118389" checksum="3A1ACDDC29DF9039" modified="2019-05-08" version="1">MVALTLEAFRKWLLLAKNWCESLTVVTACSVYEPLKSINLPRPEGESLWDKLDHYYRIVKSTVLLYQSPTTGLFPTKTFGDNQKAKVQDSLYCAACAWALALAYRRIDDDKGRTHELEHSAIKCMRGILYCCMRQADKVQQFKQDPRPSKCLHSVFSAHSGDEVFSHDEYGHLQINAVSLFLLYLVEMISSGLQIIYNTDEVSFIQNLVFCVERAYRVPDFGIWERGSKYNNGSTELHSSSVGLAKAALEAINGFNLFGNQGCSWSIIFVDLDAHNRNGQTLCSLLPRESRSHNTDAALLPCISYPAFAVDDDALYSQTLDKIVRKLKGKYGFKRFLRDGYRTSLEDRNRRYYKPAEIKLFDGIECEFPLFFIFMMIDGIFRGNPQQVKEYQELLDHLLQHSSEGYPVVPKYYYVPADFVELEKKNPGSQKRFPSNSGRDGKFFLWGQALYIIAKLLVDGLVNPKDIDPVGRYIPPQDQRNVSMRYSNQGPLENDLVVHVALIAESQRLQVFLNTYGIQTQTPQQVEPIQIWAQQELVKAYFHLGINDKLGLSGRPDRPIGCLGTSKIYRILGKTVVCYPIIFDLSDFYMSQDVMLLIDDIKNALQFIKQYWKMHGRPLFLVLIREDNIRGSRFNPILDMLAAFKKGVVGGVKVHADRVQTLISGAVVEQLDFLRISETEEPPDFKSFEELELPKHSKVKRQTSTPNVSELEQQPEVNINDWKIRSTYEILQKLHDCNCLASQAMLLSILLKREGPNFITKEGTISDHIERIYRRAGSKKLWFVYNIFVAAFTQKFASSIAPHITTLLVHGKQVKCTEVLNSAYELKIRAGDMPPRDLYQLSPSEVKQLLLDILQPQQQGRCWLNRRQIDGSLNRTPHGFYDRVWQILERTPNGVIVAEKYLPQQPTLSDMTMYEMNFSLLVEDMLQNIDQPEYRQIIVELLMVVSVILERNPELEFQDKVDLDKLVKEAFHYFQKDEIRLKGVEKQDDMTSFYNTPPMGKRGTCSYLTKAVMNLLLEGEVKPCSDDPCTIS</sequence>
</entry>
<entry dataset="TrEMBL" created="2019-05-08" modified="2026-01-28" version="30" xmlns="https://uniprot.org/uniprot">
    <accession>A0A452IUV7</accession>
    <name>A0A452IUV7_9SAUR</name>
    <protein>
        <recommendedName>
            <fullName evidence="20">Myosin heavy chain 1</fullName>
        </recommendedName>
    </protein>
    <organism evidence="18 19">
        <name type="scientific">Gopherus agassizii</name>
        <name type="common">Agassiz's desert tortoise</name>
        <dbReference type="NCBI Taxonomy" id="38772"/>
        <lineage>
            <taxon>Eukaryota</taxon>
            <taxon>Metazoa</taxon>
            <taxon>Chordata</taxon>
            <taxon>Craniata</taxon>
            <taxon>Vertebrata</taxon>
            <taxon>Euteleostomi</taxon>
            <taxon>Archelosauria</taxon>
            <taxon>Testudinata</taxon>
            <taxon>Testudines</taxon>
            <taxon>Cryptodira</taxon>
            <taxon>Durocryptodira</taxon>
            <taxon>Testudinoidea</taxon>
            <taxon>Testudinidae</taxon>
            <taxon>Gopherus</taxon>
        </lineage>
    </organism>
    <reference evidence="19" key="1">
        <citation type="journal article" date="2017" name="PLoS ONE" volume="12" first="e0177708" last="e0177708">
            <title>The Agassiz's desert tortoise genome provides a resource for the conservation of a threatened species.</title>
            <authorList>
                <person name="Tollis M."/>
                <person name="DeNardo D.F."/>
                <person name="Cornelius J.A."/>
                <person name="Dolby G.A."/>
                <person name="Edwards T."/>
                <person name="Henen B.T."/>
                <person name="Karl A.E."/>
                <person name="Murphy R.W."/>
                <person name="Kusumi K."/>
            </authorList>
            <dbReference type="PubMed" id="28562605"/>
        </citation>
        <scope>NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]</scope>
    </reference>
    <reference evidence="18" key="2">
        <citation type="submission" date="2025-08" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <reference evidence="18" key="3">
        <citation type="submission" date="2025-09" db="UniProtKB">
            <authorList>
                <consortium name="Ensembl"/>
            </authorList>
        </citation>
        <scope>IDENTIFICATION</scope>
    </reference>
    <comment type="subunit">
        <text evidence="13">Muscle myosin is a hexameric protein that consists of 2 heavy chain subunits (MHC), 2 alkali light chain subunits (MLC) and 2 regulatory light chain subunits (MLC-2).</text>
    </comment>
    <comment type="subcellular location">
        <subcellularLocation>
            <location evidence="1">Cytoplasm</location>
            <location evidence="1">Myofibril</location>
        </subcellularLocation>
    </comment>
    <comment type="similarity">
        <text evidence="2 14">Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Myosin family.</text>
    </comment>
    <dbReference type="Ensembl" id="ENSGAGT00000036057.1">
        <property type="protein sequence ID" value="ENSGAGP00000031792.1"/>
        <property type="gene ID" value="ENSGAGG00000009599.1"/>
    </dbReference>
    <dbReference type="Proteomes" id="UP000291020">
        <property type="component" value="Unassembled WGS sequence"/>
    </dbReference>
    <dbReference type="GO" id="GO:0030016">
        <property type="term" value="C:myofibril"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-SubCell"/>
    </dbReference>
    <dbReference type="GO" id="GO:0032982">
        <property type="term" value="C:myosin filament"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-KW"/>
    </dbReference>
    <dbReference type="GO" id="GO:0016460">
        <property type="term" value="C:myosin II complex"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0051015">
        <property type="term" value="F:actin filament binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="InterPro"/>
    </dbReference>
    <dbReference type="GO" id="GO:0005524">
        <property type="term" value="F:ATP binding"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="UniProtKB-UniRule"/>
    </dbReference>
    <dbReference type="GO" id="GO:0000146">
        <property type="term" value="F:microfilament motor activity"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="GO" id="GO:0006936">
        <property type="term" value="P:muscle contraction"/>
        <property type="evidence" value="ECO:0007669"/>
        <property type="project" value="TreeGrafter"/>
    </dbReference>
    <dbReference type="FunFam" id="1.10.10.820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000003">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000004">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000006">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.340:FF:000013">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000002">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000003">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000007">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.370:FF:000008">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.5.4820:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.58.530:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="2.30.30.360:FF:000001">
        <property type="entry name" value="Myosin heavy chain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="3.40.850.10:FF:000024">
        <property type="entry name" value="Myosin heavy chain, isoform J"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="FunFam" id="1.20.120.720:FF:000001">
        <property type="entry name" value="Myosin heavy chain, muscle"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.10.10.820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.340">
        <property type="match status" value="5"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.370">
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.5.4820">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.58.530">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="6.10.250.2420">
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="3.40.850.10">
        <property type="entry name" value="Kinesin motor domain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="2.30.30.360">
        <property type="entry name" value="Myosin S1 fragment, N-terminal"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Gene3D" id="1.20.120.720">
        <property type="entry name" value="Myosin VI head, motor domain, U50 subdomain"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR036961">
        <property type="entry name" value="Kinesin_motor_dom_sf"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR001609">
        <property type="entry name" value="Myosin_head_motor_dom-like"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR008989">
        <property type="entry name" value="Myosin_S1_N"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR002928">
        <property type="entry name" value="Myosin_tail"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR027417">
        <property type="entry name" value="P-loop_NTPase"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR004009">
        <property type="entry name" value="SH3_Myosin"/>
    </dbReference>
    <dbReference type="InterPro" id="IPR014751">
        <property type="entry name" value="XRCC4-like_C"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615">
        <property type="entry name" value="MYOSIN HEAVY CHAIN, NON-MUSCLE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PANTHER" id="PTHR45615:SF79">
        <property type="entry name" value="MYOSIN-4"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF00063">
        <property type="entry name" value="Myosin_head"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF02736">
        <property type="entry name" value="Myosin_N"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="Pfam" id="PF01576">
        <property type="entry name" value="Myosin_tail_1"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PRINTS" id="PR00193">
        <property type="entry name" value="MYOSINHEAVY"/>
    </dbReference>
    <dbReference type="SMART" id="SM00242">
        <property type="entry name" value="MYSc"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF90257">
        <property type="entry name" value="Myosin rod fragments"/>
        <property type="match status" value="4"/>
    </dbReference>
    <dbReference type="SUPFAM" id="SSF52540">
        <property type="entry name" value="P-loop containing nucleoside triphosphate hydrolases"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS50096">
        <property type="entry name" value="IQ"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51456">
        <property type="entry name" value="MYOSIN_MOTOR"/>
        <property type="match status" value="1"/>
    </dbReference>
    <dbReference type="PROSITE" id="PS51844">
        <property type="entry name" value="SH3_LIKE"/>
        <property type="match status" value="1"/>
    </dbReference>
    <proteinExistence type="inferred from homology"/>
    <keyword evidence="12 14" id="KW-0009">Actin-binding</keyword>
    <keyword evidence="7 14" id="KW-0067">ATP-binding</keyword>
    <keyword evidence="8" id="KW-0175">Coiled coil</keyword>
    <keyword evidence="5" id="KW-0963">Cytoplasm</keyword>
    <keyword evidence="4" id="KW-0488">Methylation</keyword>
    <keyword evidence="10 14" id="KW-0505">Motor protein</keyword>
    <keyword evidence="11" id="KW-0514">Muscle protein</keyword>
    <keyword evidence="9 14" id="KW-0518">Myosin</keyword>
    <keyword evidence="6 14" id="KW-0547">Nucleotide-binding</keyword>
    <keyword evidence="19" id="KW-1185">Reference proteome</keyword>
    <keyword evidence="3" id="KW-0787">Thick filament</keyword>
    <feature type="domain" description="Myosin N-terminal SH3-like" evidence="17">
        <location>
            <begin position="46"/>
            <end position="95"/>
        </location>
    </feature>
    <feature type="domain" description="Myosin motor" evidence="16">
        <location>
            <begin position="99"/>
            <end position="793"/>
        </location>
    </feature>
    <feature type="region of interest" description="Actin-binding" evidence="14">
        <location>
            <begin position="670"/>
            <end position="692"/>
        </location>
    </feature>
    <feature type="region of interest" description="Disordered" evidence="15">
        <location>
            <begin position="1137"/>
            <end position="1157"/>
        </location>
    </feature>
    <feature type="compositionally biased region" description="Basic and acidic residues" evidence="15">
        <location>
            <begin position="1139"/>
            <end position="1157"/>
        </location>
    </feature>
    <feature type="binding site" evidence="14">
        <location>
            <begin position="192"/>
            <end position="199"/>
        </location>
        <ligand>
            <name>ATP</name>
            <dbReference type="ChEBI" id="CHEBI:30616"/>
 
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